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The Oxford Handbook of

EEG
FREQUENCY
Edited by
PHILIP GABLE, MATTHEW MILLER,
EDWARD BERNAT
 Great Clarendon Street, Oxford, OX2 6DP,
United Kingdom
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 Contents

List of Contributors ix
Foreword xiii
Preface xv

PA RT I
1. Introduction: Methods for Collecting EEG Data for Frequency
Analyses in Humans  3
Philip A. Gable and Matthew W. Miller
2. Logic behind EEG Frequency Analysis: Basic Electricity and
Assumptions  15
Kyle J. Curham and John J. B. Allen

3. From Neural Oscillations to Cognitive Processes  40

Andreas Keil and Nina Thigpen
4. Time-​Frequency Decomposition Methods for Event-​Related
Potential Analysis  65
Selin Aviyente
5. Time Frequency Analyses in Event-​Related Potential
Methodologies  88
Anna Weinberg, Paige Ethridge,
Belel Ait Oumeziane, and Dan Foti
6. The Relationship Between Evoked and Induced EEG/​MEG
Changes: Going Beyond Labels  115
Ali Mazaheri
7. Frequency Analysis of the Monkey Neocortical Local
Field Potential  131
Steven L. Bressler
vi   Contents

PA RT I I
8. Gamma Activity in Sensory and Cognitive Processing  145
Daniel Strüber and Christoph S. Herrmann
9. Frontal Midline Theta as a Model Specimen of Cortical Theta  178
James F. Cavanagh and Michael X Cohen
10. The Role of Alpha and Beta Oscillations in the Human EEG during
Perception and Memory Processes  202
Sebastian Michelmann, Benjamin Griffiths,
and Simon Hanslmayr
11. Theory and Research on Asymmetric Frontal Cortical Activity as
Assessed by EEG Frequency Analyses  220
Eddie Harmon-​Jones, Taylor Popp, and Philip A. Gable
12. Oscillatory Activity in Sensorimotor Function  259
Bernadette C. M. van Wijk

PA RT I I I
13. EEG Frequency Development across Infancy and Childhood  293
Kimberly Cuevas and Martha Ann Bell
14. Developmental Research on Time-​Frequency Activity in
Adolescence and Early Adulthood  324
Stephen M. Malone, Jeremy Harper, and William G. Iacono
15. Theta-Beta Power Ratio: An Electrophysiological Signature of
Motivation, Attention and Cognitive Control  352
Dennis J. L. G. Schutter and J. Leon Kenemans
16. Cortical Source Localization in EEG Frequency Analysis  377
Wanze Xie and John E. Richards
17. Frequency Characteristics of Sleep  401
Alpár S. Lázár, Zsolt I. Lázár, and Róbert Bódizs
18. A Review of Oscillatory Brain Dynamics in Schizophrenia  434
Kevin M. Spencer
19. EEG Frequency Techniques for Imaging Control Functions
in Anxiety  464
Jason S. Moser, Courtney Louis, Lilianne Gloe,
Stefanie Russman Block, and Spencer Fix
 Contents   vii

PA RT I V
20. Bivariate Functional Connectivity Measures for Within-​and
Cross-​Frequency Coupling of Neuronal Oscillations  495
J. Matias Palva and Satu Palva
21. Multivariate Methods for Functional Connectivity Analysis  514
Selin Aviyente
22. Brain Stimulation Approaches to Investigate EEG Oscillations  532
Florian H. Kasten and Christoph S. Herrmann
23. Parameterizing Neural Field Potential Data  563
Bradley Voytek

Index 579
 List of Contributors

John J. B. Allen, Distinguished Professor of Psychology, Department of Psychology,

University of Arizona, Tucson, AZ
Selin Aviyente, Professor, Department of Electrical and Computer Engineering,
Michigan State University, East Lansing, MI
Martha Ann Bell, University Distinguished Professor, Department of Psychology, Virginia
Tech, Blacksburg, VA
Róbert Bódizs, Senior Research Fellow, Semmelweis University, Institute of Behavioural
Sciences, Budapest, Hungary
Steven L. Bressler, Professor, Center for Complex Systems and Brain Sciences, Department
of Psychology, Florida Atlantic University, Boca Raton, FL
James F. Cavanagh, Associate Professor, Department of Psychology, University of New
Mexico, Albuquerque, NM
Michael X Cohen, Assistant Professor, Donders Centre for Neuroscience & Radboud
University Medical Center, Nijmegen, Netherlands
Kimberly Cuevas, Associate Professor, Department of Psychological Sciences,
University of Connecticut, Storrs, CT
Kyle J. Curham, Department of Psychology, University of Arizona, Tucson, AZ
Paige Ethridge, Department of Psychology, McGill University, Montreal, QC, Canada
Spencer Fix, Department of Psychology, University of Maryland College Park, College
Park, MD
Dan Foti, Associate Professor, Department of Psychological Sciences, Purdue University,
West Lafayette, IN
Philip A. Gable, Associate Professor of Psychology, Department of Psychological and
Brain Sciences, University of Delaware, Newark, DE
Lilianne Gloe, Department of Psychology, College of Social Science, Michigan State
University, East Lansing, MI
Benjamin Griffiths, School of Psychology, University of Birmingham, Edgbaston,
Birmingham, UK
x   LIST OF CONTRIBUTORS

Simon Hanslmayr, Professor, Centre for Cognitive Neuroimaging, School for Neuroscience
and Psychology, University of Glasgow, UK
Eddie Harmon-​Jones, Professor, School of Psychological Science, University of New
South Wales, Sydney, Australia
Jeremy Harper, Department of Psychiatry and Behavioral Sciences, University of
Minnesota, Minneapolis, MN
Christoph S. Herrmann, Professor, Experimental Psychology Lab, Carl von Ossietzky
University, Oldenburg, Germany
William G. Iacono, Professor, Department of Psychology, University of Minnesota,
Minneapolis, MN
Florian H. Kasten, Experimental Psychology Lab, Carl von Ossietzky University,
Oldenburg, Germany
Andreas Keil, Professor of Psychology, Department of Psychology and Center for the
Study of Emotion & Attention, University of Florida, Gainesville, FL.
J. Leon Kenemans, Professor, Biopsychology and Psychopharmacology, Utrecht
University, Utrecht, Netherlands
Alpár S. Lázár, Associate Professor, School of Health Sciences, University of East
Anglia, Norwich, UK
Zsolt I. Lázár, Assistant Professor, Faculty of Physics, Babeş-​Bolyai University, Cluj-​
Napoca, Romania
Courtney Louis, Department of Psychology, College of Social Science, Michigan State
University, East Lansing, MI
Stephen M. Malone, Research Assistant Professor and Co-​investigator, Minnesota
Center for Twin & Family Research, University of Minnesota, Minneapolis, MN
Ali Mazaheri, Associate Professor, School of Psychology, University of Birmingham,
Edgbaston, Birmingham, UK
Sebastian Michelmann, Princeton Neuroscience Institute, Princeton, NJ
Matthew W. Miller, Associate Professor, School of Kinesiology, Auburn University,
Auburn, AL
Jason Moser, Professor, Department of Psychology College of Social Science, Psychology,
Michigan State University, East Lansing, MI
Belel Ait Oumeziane, Department of Psychological Sciences, Purdue University, West
Lafayette, IN
 LIST OF CONTRIBUTORS    xi

J. Matias Palva, Research Director, Department of Neuroscience and Biomedical

Engineering, Aalto University, Helsinki
Satu Palva, Professor, Centre for Cognitive Neuroimaging, School of Psychology and
Neuroscience, University of Glasgow, Glasgow
Taylor Popp, Carnegie Mellon University, Pittsburgh, PA
John E. Richards, Professor, Department of Psychology, University of South Carolina,
Columbia, SC
Stefanie Russman Block, Department of Psychology, College of Social Science, Michigan
State University, East Lansing, MI
Dennis J. L. G. Schutter, Professor, Experimental Biopsychology of Motivation and
Emotion, Utrecht University, Utrecht, Netherlands
Kevin M. Spencer, Research Health Scientist, Veterans Affairs Boston Healthcare
System; and Associate Professor of Psychiatry, Harvard Medical School, Boston, MA
Daniel Strüber, Adjunct Professor, Experimental Psychology Lab, Carl von Ossietzky
University, Oldenburg, Germany
Nina Thigpen, Research Scientist at Google X, The Moonshot Factory, Mountain
View, CA.
Bernadette C. M. Van Wijk, Marie Curie Research Fellow at the University of Amsterdam,
Amsterdam, Netherlands
Bradley Voytek, Associate Professor, Department of Cognitive Science, UC San Diego,
San Diego, CA
Anna Weinberg, Assistant Professor, Department of Psychology, McGill University,
Montreal, QC, Canada
Wanze Xie, Assistant Professor, School of Psychological and Cognitive Sciences, Peking
University, Beijing, China
 Foreword

Steven J. Luck

When Hans Berger conducted his pioneering human EEG recordings in the 1920s, his
first major discovery was the alpha rhythm, a 10-​Hz oscillation that grew larger when the
subject’s eyes were closed. A few years later, Lord Adrian (my intellectual great-​great-​
grandfather) showed that the alpha rhythm also varied according to whether the subject
was focusing intensely or daydreaming. Thus the study of EEG oscillations was born.
But this area of research underwent a protracted childhood, because the scientists
of the mid-​twentieth century could not easily see smaller oscillations amidst the cha-
otic twists and turns of the scalp EEG. To pull out specific neural processes from the
complex and noisy EEG, they began to use signal averaging techniques that can isolate
the brain potentials that are triggered by specific events such as the onset of a light (the
event-​related potentials or ERPs). However, these techniques assume that the phase of
the signal is constant across trials, and the application of signal averaging to EEG data
eliminates or hopelessly distorts oscillating activity. Indeed, for the first 30 years of my
own research career, I viewed the alpha rhythm as a nuisance that should be suppressed
lest it contaminate my precious ERP waveforms.
All of this began to change in the 1980s and 1990s, partly driven by high-​profile studies
of local field potential oscillations in animals and partly driven by the application of
time-​frequency analysis methods to human EEG recordings. The brain oscillations that
were obscured by signal averaging could now be visualized and quantified. A new gen-
eration of researchers began studying human EEG oscillations and linking them with
microelectrode data from animals and computational models of brain dynamics.
In science, the introduction of a new approach often leads to a burst of progress
followed by the realization that things are not as simple as they seem. The new wave of
oscillation research followed this common path, with the discovery of important new
phenomena being accompanied by conceptual and methodological challenges. One
such challenge—​related to the famous Heisenberg Uncertainty Principle—​is that pre-
cision in the time-​domain is inversely related to precision in the frequency domain. The
more precisely you determine the frequency of an oscillation, the less you can say about
when the oscillation was present.
A second important challenge is the difficulty of distinguishing between bona fide
oscillations and other kinds of neural events. Part of the genius of the Fourier transform
xiv   Foreword

is that any waveform—​whether or not it contains oscillations—​can be reconstructed by

summing together a set of sinusoids. As a result, when you apply a method such as the
Fourier transform to a time-​domain signal such as the EEG, you will always see activity
at some frequencies, whether or not the signal is actually oscillating. And what if the
brain is oscillating, but not in a sinusoidal manner? This can lead to completely artifac-
tual results, such as the apparent coupling of the amplitude of one frequency with the
phase of another frequency.
A third key challenge is the interpretation of brain oscillations. Virtually any system
containing multiple interconnected parts will oscillate when energy is introduced.
Those oscillations can be essential to the function of the system, as when the sound of a
single violin fills a concert hall or when a clock keeps perfect time. But oscillations can
also be a sign of trouble, as when a bridge shakes violently following an earthquake or
when a seizure spreads throughout the brain. The fact that the brain oscillates, and that
the oscillations vary across states or tasks, does not mean that the oscillations them-
selves are playing a functional role in the brain’s computations. They may be epiphe-
nomenal. Or they may be fundamental. Distinguishing between these possibilities may
require invasive recordings in animals or experimental manipulations of oscillations via
brain stimulation.
The study of EEG oscillations has reached a key point in its development. The appli-
cation of time-​frequency methods to EEG data is now commonplace, aided by open
source data analysis packages such as FieldTrip and EEGLAB. Labs that eschewed
these methods for many years—​including my own—​are now examining oscillations
alongside traditional event-​related potentials. However, this success means that more
researchers are using time-​frequency analyses without understanding the challenges
involved in properly quantifying and interpreting oscillatory activity. As a result, this
edited volume has appeared at the perfect time.
The chapters in this volume will give scientists of all career stages the knowledge
they need to understand how oscillations arise in the brain, how they can be accurately
quantified, and how they can be appropriately interpreted. I encourage readers to think
deeply about the many important issues that are raised in these chapters, especially with
regard to the three challenges I have outlined. But if you think deeply and follow the
“best practices” described in this volume, you will be able to see brain activity that would
otherwise be invisible. And you may make important new discoveries about the human
mind and brain.
 Preface

Motivation

The time is ripe for a comprehensive book on the array of historical and cutting-​edge
frequency and time-​frequency approaches to studying EEG/​ERP because of the wide-​
spread interest in frequency research. There is a great need for a book organizing the di-
verse and important methods of EEG frequency analyses and interpreting the resultant
measures.
One stream of research comes from traditional (band-​based) frequency analyses.
Likewise, understanding the cutting-​edge frequency analyses which may not be familiar
to many EEG researchers is increasingly important for investigators applying frequency
analyses. However, there is a major need for a comprehensive handbook on analyses
within this domain. Although research has been rapidly accumulating over the last
decade, there has not been sufficient organization of research on the topic. We believe
this comprehensive handbook is increasingly necessary to help delineate the boundaries
of the area, the major scientific questions that need to be addressed, and the core theor-
etical frameworks that can guide future research and development.
Thus, a specific goal of this book is to bring together these various scientific
perspectives and research approaches within a single reference volume that provides an
integrated, cutting-​edge overview of the current state of the field. This volume comprises
contributions from leading researchers within various allied disciplines.
The use of electroencephalography (EEG) to study the human mind has seen tre-
mendous growth across a vast array of disciplines due to increased ease of use and
affordability of the technology. EEG is a non-​invasive measure of electrical brain ac-
tivity. Typically, researchers investigate the EEG signal using either time-​domain (e.g.,
ERP) analyses or frequency analyses. Several books have examined practicalities of
conducting ERP analyses and interpreting various ERP measures. However, a compre-
hensive book has yet to be developed organizing the numerous ways to process EEG
frequency and interpreting frequency measures linked to cognitive, affective, and motor
processes.
We (editors Philip, Matt, and Ed) felt a great need for a book organizing the diverse
and fascinating methods of EEG frequency analyses and interpreting the resultant
measures. Frequency analyses provide unique assessments of neural functioning,
neural connectivity, and “resting” neural activity studied by EEG researchers. Further,
xvi   Preface

frequency-​domain measures are reliably associated with cognitive, affective, and motor
processes of great interest to neuroscientists and psychological scientists. For example,
asymmetrical activation of the frontal cortex as measured by the inverse of alpha-​band
activity is closely linked with motivation and emotion. In addition, analyses examining
the synchrony of EEG frequencies recorded from different scalp locations allow
researchers to examine brain connectivity without having to incur the costs of magnetic
resonance imaging.

ORIGIN

As EEG frequency researchers, we wanted a resource that introduced the myriad ways
in which EEG frequency analyses are being investigated. This volume began while the
lead editor, Philip, was on sabbatical and seeking to begin a new chapter in his career.
During that time, he visited with Matt Miller and the project quickly developed into a
collaborative project. After developing the project more, Matt and Philip brought Ed
Bernat on board.
As individual editors, we each had areas of expertise in EEG frequency research, but
in developing this volume, we quickly discovered that each of our individual areas of
expertise were far different from each other. Three editors were necessary to even try
to cover the breadth of EEG frequency research being conducted. In addition, we did
not want to create a handbook that focused on EEG frequency research specifically in
emotion, cognition, or clinical applications. Instead, we wanted to provide a survey of
the breadth of work being conducted with EEG frequency research. Try as we might, we
also acknowledge this handbook will inevitably fail to cover everyone’s interest across all
topics. To that end, we hope to receive feedback from readers so that future editions of
this handbook can be expanded to encompass the ever-​growing field of EEG frequency
research.
Together, the project has been a long labor, but well worth the time and effort to de-
velop the resource. We have been especially excited to work with leading experts in the
field as they develop chapters for the volume. We are excited for you as the reader to see
what we have gotten to see throughout the editing process: the excitement and develop-
ment of EEG frequency research across a wide range of fields and programs of research.

ORGANIZATION

To aid in reading the handbook, much thought and structuring has been given to the
organization of the chapters. As a whole, the book provides a systematic summary of
EEG frequency analyses and applications. Individual chapters give depth to each type
 Preface   xvii

of frequency analysis and interpretation of resultant measures. Chapters are organized

into three sections.
The first section of the book is focused on basics of EEG frequency research and
linking frequency analyses to other components of EEG research, such as event-​related
potential (ERP) components and the fundamentals of inference from EEG recording.
For the second section, contributors focus on specific EEG frequency components that
are commonly studied using traditional frequency bands of activity to study specific
psychological processes related to cognition, motivation, and perception. The third
section focuses on EEG frequency analyses in special populations and altered states. The
fourth section of the handbook concludes with chapters focused on advancing method-
ology used in EEG frequency research.
The initial chapters in the first section describe methods for collecting EEG data for
frequency analyses in humans as well as the basics of electrical activity and assumptions
regarding the EEG signal. Following these chapters, contributors consider how the
oscillations in the EEG signal may give rise to psychological phenomena, and how
ERPs can be decomposed into time-​frequency components. The chapters go on to con-
sider the relationships between evoked (ERP-​related) and induced EEG activity before
shedding light upon frequency analyses of LFPs in non-​human primates, which may
inform frequency analyses of human EEG. The second section begins with the role of
gamma oscillations in cognitive and sensory processing. Then, frontal midline theta,
which is often linked to cognitive control, is addressed, followed by the role of alpha and
beta oscillations in perception and memory. Next, research on asymmetries in frontal
alpha oscillations are linked with motivation, followed by a description of the role of
oscillatory activity in sensorimotor function. Moving into the third section of the book,
the first chapter in this section focuses on changes in EEG frequency throughout in-
fancy, childhood, adolescence, and early adulthood. Next, follows an examination of
the ratio of theta to beta power in motivation and attentional control as they relate to
normal and abnormal behavior. Then, the characteristics of the oscillations in persons
with schizophrenia are described, followed by an examination of how frequency
analyses clarify control processes in people with anxiety. The concluding section of the
handbook begins with specialized frequency analyses for source localization and brain
connectivity before concluding with chapters describing how to manipulate oscillatory
activity with brain stimulation and how to parameterize neural field potential data to,
for example, tease apart true oscillations from aperiodic signals.

LIMITATIONS OF THE
CURRENT VOLUME

As with any book, the current volume is not a complete work on the topic of EEG fre-
quency analyses. In addition, this is not the first book on EEG frequency research. Many
xviii   Preface

excellent books have been published this topic. Here we mention what the current book
does not include and refer the reader to additional resources available in other books.
One of the most inspiring resources for us as editors is Steven J. Luck and Emily
S. Kappenman’s Oxford Handbook on Event-​Related Potential Components. This book
has been used in our courses and labs, as well as by countless other EEG researchers. It
focuses on the excellent work that has investigated the spectrum of ERP components
derived from EEG research. We were inspired to build a similar handbook that would
cover EEG frequency analyses. The current book does not address ERP research in
much detail. The closest chapters addressing ERP research are Chapters 4 and 5. For
those desiring a more comprehensive volume on ERP analyses, please see Luck and
Kappenman.
Another extraordinary resource is Mike X. Cohen’s Analyzing Neural Time Series
Data: Theory and Practice. This book has served as the primary resource for many EEG
researchers, including us, to learn how to conduct time-​frequency analyses, and to teach
our students how to perform the analyses. The book is particularly helpful in guiding
the reader from the mathematical bases of frequency analyses to the implementation
of these analyses in MATLAB. These analyses are referenced throughout our book and
include fast Fourier transforms, complex Morlet wavelet convolution, intertrial phase
clustering, surface Laplacian filtering, phase-​and power-​based connectivity measures,
and cross-​frequency coupling. Chapters in our book that address these “how-​to” topics
include Chapters 4, 5, 19, 20, 21, and 23.
Finally, Chapter 1 of this volume describes how to collect EEG data. For readers
who which to have greater detail about implementing and collect EEG from human
participants, we recommend Dickter and Kieffaber’s EEG Methods for the Psychological
Sciences. This book is an excellent resource for researchers beginning to implement EEG.
Pa rt I
 Chapter 1

INTRODU C T I ON
Methods for Collecting EEG Data for Frequency
Analyses in Humans

PHILIP A. GABLE AND MATTHEW W. MILLER

1.1 Chapter Aims

This chapter aims to provide a structure for readers to understand the methodology
behind collecting EEG (electroencephalography) presented in the subsequent chapters.
It is important to first understand the research methods involved in recording EEG fre-
quency before delving into more advanced frequency analyses and the interpretations.
As researchers, we focus this first chapter on a brief introduction to the topic of EEG
methodology and scientific practices. To begin, however, we feel it is important to lay
down definitions for terms used throughout the book.

1.2 Definitions of EEG

Frequency Research

EEG refers to the recording of electrical brain activity from the human scalp. It is one
of the most common methods for measuring brain functioning in areas of mind, brain,
and behavior science. EEG data contain rhythmic activity or waves that may reflect
neural oscillations, or fluctuations in the excitability of populations of neurons (more
on this later). These rhythmic fluctuations are typically described using two main
descriptors. The first is frequency, which is the speed of the wave, and it is measured
in hertz (Hz), which is the number of wave cycles per second. The second is power,
which is the squared amplitude of the wave. The greater the power of an oscillation, the
greater the energy of that oscillation. All of the chapters in this work discuss frequency
4    PHILIP A. GABLE and MATTHEW W. MILLER

with most referencing power. Sometimes researchers investigate the phase of the wave,
which is the position of the wave measured in radians or degrees. Many of the chapters
included here discuss phase.
The brain produces rhythms in multiple frequencies, which can be isolated from
the raw EEG signal using multiple techniques described by Curhamn & Allen in
Chapter 2, and Voytek in Chapter 23). Different psychological processes are linked to
different frequencies, which are often grouped into bands. The most commonly studied
bands include delta (1–​4 Hz), theta (4–​8 Hz), alpha (8–​12 Hz), beta (13–​30 Hz), and
gamma (lower gamma 30–​80 Hz; upper gamma 80–​150 Hz). While these are not the
only frequency bands, these are the bands most typically associated with processes
of mind and behavior measured by EEG. Importantly, these bands are not defined
without reason, but instead reflect biological changes at the cellular level (see Cohen,
2014 and Buzsaki, 2006 for reviews). However, these bands are not rigid and may vary
depending on individual differences, such as brain development, structure, and chem-
istry. Chapters in the second section of this work note how different frequency bands are
associated with cognitive, motivational, and sensorimotor processes.
These definitions are by no means complete. Individual chapters provide more pre-
cise definitions of terms used. With this initial framework, readers should be able to
venture into subsequent chapters focusing in more detail on these definitions. Because
EEG is a rather complex measure, we focus in more detail on the physiological basis and
scientific methods used to record and process EEG.

1.3 Physiological Basis of EEG

EEG is measured because all nerve cells communicate using electrical signals, sending
information throughout the brain and to the rest of the body. Within a neuron, an action
potential is an electrical wave that runs from the axon hillock at the cell body to the axon
terminals. At the axon terminals, the action potential causes neurotransmitter to be
released. This neurotransmitter crosses the synaptic gap and binds to receptors on the
membrane of the postsynaptic cell. Binding to the receptor causes voltage changes by
activating ion channels or second messengers that either excite or inhibit the postsynaptic
neuron. The summation of this voltage change in the membrane of the dendrites and cell
body of the postsynaptic neuron is called a postsynaptic potential. Postsynaptic potentials
tend to occur locally, rather than moving down the axon. This allows postsynaptic
potentials to summate rather than to cancel, resulting in voltage changes that have larger
amplitudes and can be recorded on the cortical surface or at the scalp.
When tens of thousands to millions of neurons are excited or inhibited at the same
time, the voltage change outside the cell (extracellular potential) can be recorded at the
scalp using EEG, which measures the sum of electrical activity from excitatory and in-
hibitory postsynaptic potentials over this collection of neurons. The activity can only be
recorded on the scalp surface because tissue (cerebrospinal fluid, meninges, skull, and
 INTRODUCTION   5

skin) between the neurons and scalp conducts the electrical signal. In addition, for elec-
trical activity to be projected to the scalp, cellular alignment must be precisely arranged
in parallel so that their effects cumulate to project the electrical activity to the scalp (see
Curhamn & Allen, Chapter 2; Keil & Thigpen, Chapter 3). Neurons must be arranged
so that the cluster of neurons all have dendrites at one pole and axons departing at the
opposite pole. This arrangement is called an open field and occurs when neurons are
organized in layers. The cortex, cerebellum, and parts of the thalamus tend to have this
open field arrangement of neurons resulting from pyramidal cells.

1.4 A Dual Nature

Due to the electrical basis of the EEG signal, EEG has excellent ability to tell us when
something is happening in the brain. This is called temporal resolution and is one of
the greatest strengths of the EEG signal. The EEG signal measures neural activity at the
accuracy of milliseconds, which allows for the ongoing measurement of psychological
processes as they unfold (Luck, 2014).
However, the EEG signal is limited in its ability to measure where something is
occurring. This is called spatial resolution and is one of the greatest weaknesses of the
EEG signal. Depending on where the source of the EEG signal is generated, the orien-
tation of the open field neurons might not be parallel to the scalp, thus generating EEG
signals in multiple directions. In addition, resistors (e.g., the skull) in the tissue between
the neurons and scalp can cause the EEG signal to spread out. Because of the volume
conduction through the head, as well as the orientation of the pyramidal cells emitting
the signal, the spatial location of the signal is difficult to ascertain. As Keil and Thigpen
(Chapter 3) note, a difference in frequency power between two experimental conditions
could be the result of a different number of neurons activated, the temporal order in
which they were activated, or neurons with different orientations being activated. To
address EEG’s limited spatial resolution, cortical source localization techniques have
been developed and are reviewed by Xie and Richards (Chapter 19). In sum, using an
analogy from Steve Luck’s ERP Boot Camp, the EEG signal is like being able to see every
frame of a movie as it unfolds. However, because of the low spatial resolution, the movie
appears a bit blurry.

1.5 EEG Equipment and Recording

The earliest method of EEG measurement was implemented by Hans Berger in the
late 1920s. In his early experiments, he used two sponges soaked in saline connected
to an amplifier (Berger, 1929). While the equipment and processing of EEG signal has
advanced considerably since that time, the basic components remain similar. EEG
6    PHILIP A. GABLE and MATTHEW W. MILLER

electrodes are placed on or near the head, the signal from the electrode is transmitted to
an amplifier, and the signal is digitized and recorded.
In psychological research labs, EEG is usually recorded from 32, 64, 128, or more
electrodes. In other research (e.g., sleep, nonhuman) fewer electrodes (2–​8 electrodes)
is more typical. When larger numbers of electrodes are used, electrodes are mounted in
an electrode cap or net. When fewer electrodes are used, electrodes may be positioned
individually on the head using a bonding agent. Electrodes can either be wet electrodes
or dry electrodes. Wet electrodes are made of silver, silver-​chloride, or tin, and a con-
ductive gel or liquid is placed inside or around the electrode. Dry electrode systems
use electrodes coated in gold, silver, or nickel, and place electrodes directly on the scalp
without a conductive medium. Wet electrodes generally have higher signal quality, but
dry electrodes may be preferred when high impedance levels are tolerable, or when re-
cording for long periods.
Electrode systems will have active or passive electrodes. Active electrodes con-
tain a small pre-​amplification unit directly attached to the conductive metal in
the electrode. This allows the EEG signal picked up at the sensor to be immedi-
ately amplified before additional environmental noise can be introduced. Passive
electrodes do not have amplification at the electrode, and instead carry the signal to
an amplifier about a meter away. Compared to passive electrodes, active electrodes
minimize noise introduced during signal transmission, tolerate high impedance re-
cording, and reduce participant preparation time. Passive electrodes have a lower
profile to benefit transcranial magnetic stimulation over the cap and can be used
inside an MRI bore.
Electrode placement is predominantly based on the 10–​20 system (Jasper, 1958).
Electrodes are named using the first letter to refer to the brain region under the electrode
from anterior to posterior (e.g., Fp—​frontal pole, F—​frontal, C—​central, P—​parietal,
T—​temporal, O—​occipital). Numbers following the letter are used to indicate the lateral
position of the electrodes. Ascending odd numbers indicate sites more lateral over the
left hemisphere of the brain, whereas ascending even numbers indicate sites more lateral
over the right hemisphere of the brain. The letter Z is used to designate medial sites. In
addition to the recording electrodes, EEG also requires a ground electrode, which assists
in reducing electrical noise, as well as a reference electrode placed on the head or face.
The raw EEG signal is usually filtered during recording. Signals below 0.1 Hz or above
200 Hz are removed because the frequency bands of interest fall within this range.
A filter at 60 Hz (in North and South America) or 50 Hz (in Europe, Asia, and Africa)
may also be used to further reduce electrical noise from alternating current.

1.6 Artifacts

The quality of the EEG is crucial to EEG frequency analysis. To best record EEG signal
reflecting brain activity, researchers must remove signal that occurs because of anything
other than neural activity. Signal that is not naturally present is called artifact. These
 INTRODUCTION   7

can be biological (e.g., muscle movement) or nonbiological (e.g., electrical noise). Much
artifact can by eliminated by taking preventative measures. Artifact that cannot be
prevented should be removed.
Muscle artifact, or electromyography (EMG), is one of the most common types of
artifacts and is usually high in frequency (100–​500 Hz). Usually, this falls outside of the
frequency range typically investigated by researchers. However, some muscle artifact
may seep into lower frequencies. Researchers can reduce muscle artifact by instructing
participants to limit their muscle movements. Muscle artifact that does occur can be
removed through visual inspection, filtering, and automatic artifact detection algo-
rithm. It should be noted that some muscle artifact may be related to the experiment
(e.g., sensorimotor studies). In such cases, it may be beneficial to measure EMG at the
site movement is expected (e.g., the hand), then control for it in analyses.
Eye movements are another common type of artifact. The eyeball is polarized which
causes large artifact in the form of voltage changes resulting from moving the eyes. Like
dealing with muscle artifact, eye movement artifact can be removed from signal using
recordings near the eyes called electro-​oculograms (EOG). One pair of EOG electrodes
are placed above and below the eye, while another pair is placed just lateral to either eye
on the temple. Eye blinks also create eye movement artifact. It is preferable to correct
blink artifact using an artifact reduction algorithm based on regression, principal com-
ponent analyses, or independent component analyses.
Artifacts occurring in the environment are the result of nonbiological factors. The
most common sources of these artifacts are the result of external electrical noise coming
from compact fluorescent lightbulbs, data hubs, or electrical junctions. Grounding will
aid in reducing these sources of noise, as will electromagnetically shielded rooms.

1.7 Frequency Processing

Once an EEG signal is recorded, the raw data must go through several processing
steps before it is in a format useable in analyses. The raw signal is collected in the
time-​domain but must be converted into a frequency-​domain representation. One
way this can be accomplished is in the form of a power spectrum, which collapses fre-
quency data across time to map the frequencies present. A frequency analysis can be
conducted over windows that are minutes, seconds, or milliseconds in length; these are
called epochs. Epochs that are seconds or minutes can be analyzed for power spectra
using a Fourier transform, which decomposes a signal into a series of sine and cosine
functions of various frequencies. The function of each frequency begins with its own
phase. A Fourier transformation assumes that the epoch repeats infinitely forward
and infinitely backward in time. A process called windowing is used to prevent artifact
created from the Fourier transform. However, windowing can also introduce artifact
and data loss into the frequency analysis. Overlapping epochs is a way to prevent dis-
continuity, data loss near the ends of the epoch, and to help meet the assumptions of the
Fourier transform in windowing.
8    PHILIP A. GABLE and MATTHEW W. MILLER

One of the most common forms of signal frequency processing is to use a fast
Fourier transform (FFT). An FFT provides the spectrum of frequency power for a
period, which is often averaged across a range of frequencies comprising a band (e.g.,
theta). It also provides a spectrum of phase. The power spectrum reflects the energy
of each frequency determined by the squared amplitude of the wave. The phase spec-
trum reflects the phase in radians or degrees of the sine or cosine wave at each interval
(e.g., 1/​T). Most frequency analyses focus exclusively on frequency power. However,
there is increasing interest in examining the phase of frequencies (e.g., see Michelmann,
Griffiths, & Hanslmayr, Chapter 10; Palva & Palva, Chapter 20). Crucially, the FFT has
two limitations: it poorly depicts changes in the frequency spectrum over time, and it
assumes the EEG data are stationary during the period to which the FFT is applied. To
overcome these limitations, time-​resolved frequency decomposition techniques are
growing in popularity, particularly wavelet analyses (e.g., complex Morlet wavelets),
which reveal changes in power at various frequencies with excellent temporal precision
(see Aviyente, Chapter 4; Weinberg, Ethridge, Oumeziane, & Foti, Chapter 5).

1.8 Experimental Design

Regardless of the EEG recording and processing, researchers need to be considerate of

experimental design. In any experiment, researchers should manipulate a single vari-
able at a time to ensure internal reliability, but this procedure is challenging in EEG
research. This follows because manipulating one variable between conditions may inad-
vertently change a second variable between conditions that affects EEG. For example, if
an experimental condition attempts to manipulate participants’ motivation while they
are physically responding to stimuli, then EEG linked to motivation may change (see
Harmon-​Jones, Popp, & Gable, Chapter 11) but so may the vigor of their responses and
EEG linked to sensorimotor function (see van Wijk, Chapter 12). Thus, it is crucial that
researchers attempt to experimentally control for such factors in their experimental de-
sign or statistically account for them by collecting covariates, such as EMG to index the
vigor of motor responses.

1.8.1 Reproducibility in Electrophysiology: Challenges

and Recommendations
A scientific discipline benefits from reproducible results because they strengthen
confidence that they reflect the way a system operates under certain conditions.1

1 There have been special issues in the International Journal of Psychophysiology and Psychophysiology

devoted to reproducibility and, relatedly, open science in cognitive electrophysiology, and readers are
encouraged to read these special issues (Kappenman & Keil, 2017; Larson & Moser, 2017).
 INTRODUCTION   9

We use Goodman, Fanelli, and Ioannidis’s (2016) definition of reproducible results

as “obtaining the same results from the conduct of an independent study whose
procedures are as closely matched to the original experiment as possible” (pp. 2–​3).
Although there are different definitions of what it means to obtain “the same results”
(Open Science Collaboration, 2015), we hope that closely matched studies yield
effects with confidence intervals that overlap substantially (for details on using confi-
dence intervals in EEG studies, see Groppe [2017]), thus allowing us to make precise
inferences about the true size of the effect being studied. Since reproducible results are
crucial for a discipline, it is important to consider what can be done to obtain them.
We discuss several research practices that increase the likelihood that an original re-
sult will be reproducible and that subsequent studies reproduce the original result,
as well as challenges faced by cognitive electrophysiology researchers attempting
to do so.

1.9 Pre-​R egister Specific Hypotheses

Each study should test specific hypotheses because results that confirm hypotheses
are more likely to be true than results based on exploratory analyses (Ioannidis, 2005).
Thus, it is crucial that researchers do not rewrite their hypotheses to fit with their results,
a practice known as HARKing (hypothesizing after results are known), as it exaggerates
the confidence the reader has that the results are true. One method to avoid HARKing
is pre-​registering hypotheses using the Open Science Framework (osf.io), aspredicted.
org, or other repositories. However, formulating specific hypotheses for cognitive elec-
trophysiology studies can be difficult, especially if researchers want to frame them stat-
istically. For example, a researcher may be confident in predicting that an experimental
condition will affect EEG activity, but they may struggle to define “EEG activity” as
a dependent variable. In ERP studies, this is less of a concern because the dependent
variables (ERP components) are well-​characterized (Kappenman & Luck, 2012); how-
ever, there are fewer well-​characterized time-​frequency variables. Although researchers
can be vague about defining their time-​frequency variables, they should then use stat-
istical analyses with strict corrections for multiple comparisons (see Cohen, 2014). This
reduces the likelihood of making Type I errors, but consequently increases the likeli-
hood of making Type II errors. (Researchers may also consider data-​driven region-​of-​
interest approaches; see Brooks et al., 2017). Therefore, time-​frequency analyses will
benefit from having well-​defined dependent variables (Indeed, this was an initial im-
petus for this book!).
For example, if a researcher believes an experimental manipulation is likely to influence
EEG activity related to cognitive control, they can have a clearly defined dependent vari-
able of oscillatory activity in the theta frequency bandwidth measured from frontal mid-
line electrodes over a certain time (Cavanagh & Cohen, Chapter 9). In a pre-​registration,
researchers should use a more precise definition of their dependent variable than “frontal
10    PHILIP A. GABLE and MATTHEW W. MILLER

midline theta”. For example, they could specify that they will determine the wavelet of 4–​
8 Hz that exhibits the greatest peak power between 200 and 600 ms after stimulus onset
at electrode Fz for each participant, and then compute the average power of this wavelet
during the 200–​600 ms time epoch at Fz. They could also introduce some flexibility into
the specification of the dependent variable by noting that they will choose a different band-
width, epoch, and/​or electrode if the grand average time-​frequency plot (averaged across
all conditions) reveals an unexpected time-​frequency and/​or scalp distribution. In this
example, the average across all conditions avoids biasing the analysis in favor of choosing
a time-​frequency window exhibiting differences between conditions. Besides reducing
HARKing and facilitating the specification of dependent variables, pre-​registration is cru-
cial for holding researchers accountable to their research design, statistical analyses, and
sample size; however, none of these positive features of pre-​registration work if researchers
deviate from their pre-​ registration without properly noting the deviation (Claesen
et al., 2019).

1.10 Increase Power and Conduct

A Priori Power Calculations

Another way that researchers can increase the likelihood that study results are reprodu-
cible is by increasing the power of their studies (Ioannidis, 2005). Alarmingly, Button
and colleagues’ (2013) analysis of neuroscience studies found their average power was
very low (8–​31%). There are two general ways that researchers can increase the power of
their studies. First, researchers should attempt to maximize the effect they are studying
and minimize its variance (i.e., increase the standardized effect size). This can be done
by using strong experimental manipulations and reliable dependent variables, such
as those discussed in this work. Additionally, researchers should increase the signal
to noise ratio in their studies by optimizing the number of trials and collecting good
EEG data (Cohen, 2017; Luck, 2014).2 Further, when possible, researchers should use
within-​subjects designs, which is already the case in many cognitive electrophysi-
ology studies. Second, researchers should collect larger samples, which is particu-
larly important when testing between-​subjects effects or within-​between subject
interaction effects. Cognitive electrophysiology studies can require a lot of time to
collect and process data, so collecting more participants may seem burdensome, es-
pecially for researchers investigating small–​medium effects. For example, a two-​tailed

2
A good way for researchers to benefit the field (and their citation count) may be for them to establish
the number of trials required for different time-​frequency variables in different paradigms, which has
been done for ERP variables (e.g., Rietdijk et al., 2014). It is worth noting that simply adding more trials
may not increase the signal to noise ratio, since participants may fidget more toward the end of long data
collections, consequently increasing noise.
 INTRODUCTION   11

dependent t-​test for an effect size of dz =​0.35, an alpha =​.05, and power =​.90 requires
88 participants, according to G*Power 3.1.9.4 (Faul et al., 2009). Indeed, it is likely that
researchers will often find themselves studying small-​medium effects. Specifically,
when researchers conduct a priori power calculations to determine their sample sizes,
they should assume that the effect sizes in the extant literature are inflated, due to pub-
lication bias by researchers and journals (i.e., only publishing significant results) (for
a more detailed discussion on sample size calculations in EEG studies, see Larson &
Carbine, 2017). Although it is difficult to collect and process large samples, it is crucial
to the reproducibility of cognitive electrophysiology studies. To reduce the demands
large sample sizes impose, researchers who mentor doctoral students, review for and
sit on the editorial boards of journals, are involved in hiring decisions about faculty and
post-​doctoral researchers, and are involved in promotion and tenure decisions should
reconsider expectations about the speed of science and the number of publications (for
further discussion on these issues, see Bradley (2017) and Yeung [2019]). Also, if a re-
searcher is concerned about allocating a lot of time to a study that may not yield signifi-
cant results, they can conduct a sequential analysis where they pause data collection
after a pre-​specified sample has been collected and then determine whether to continue
data collection based on if the incremental results are significant (given an adjusted
alpha level) and if the incremental results suggest an effect size that is too small to be of
interest (Lakens, 2014).

1.11 Make Methods, Materials,

and Data Open

In addition to pre-​registered hypotheses and adequately powered studies, researchers

should also make their methods, materials, and data accessible3. In so doing, they
will allow other researchers to “methodologically reproduce” the original study
(Goodman et al., 2016), which should increase the likelihood of observing the same
results. Researchers should make their stimuli and stimulus presentation scripts
available and include specific details about instructions given to participants and the
equipment used for the study. Further, researchers should also make signal and stat-
istical processing scripts available and provide their data when possible so that other
researchers can attempt to reproduce analyses or explore new ones. There are various
ways to provide this information, including on the Open Science Framework and
GitHub (github.com).

3
There is a special issue in the International Journal of Psychophysiology devoted to open science in
human electrophysiology, and readers are encouraged to read it (Clayson, Keil, & Larson, 2022).
12    PHILIP A. GABLE and MATTHEW W. MILLER

1.12 Replicate and Expand

The recommendations were made to increase the likelihood that researchers’ ori-
ginal results will be reproducible, but the recommendations also apply to researchers
attempting to reproduce original results. Crucially, researchers attempting to repro-
duce original results should also attempt to replicate and expand an original finding
(Cohen, 2017), preferably increasing the sample size by two and half times the ori-
ginal (Simonsohn, 2015). Specifically, cognitive electrophysiology will benefit if most
studies include an attempt to reproduce an original result and then add a new result
(e.g., by adding a new experimental condition). With results from replication attempts,
more precise estimates about the direction and size of effects can be made. Of course,
it is nearly impossible to reproduce a study methodologically. For example, different
researchers may have different criteria for manually rejecting trials, and different in-
dependent component analyses may yield different components. However, researchers
can still come quite close to a methodological reproduction, especially if methods and
materials are available for them to use. A challenge for cognitive electrophysiologists
is that they often employ different signal processing methods, such as subtracting or
not subtracting the ERP from an epoch of EEG data prior to convolving the data with
a wavelet. To this end, researchers should use the same methods as the original study,
unless a different method is clearly superior. Of course, whether a different method is
clearly superior is debatable; thus it is incumbent upon the researcher conducting the
methodological reproduction to state their case in a compelling way.

1.13 Explore

Some of the most exciting and reproducible effects in cognitive electrophysiology have
been discovered by accident (e.g., Kutas & Federmeier, 2011), meaning that they would
not have occurred if researchers had only conducted confirmatory research testing a
priori hypotheses. Thus, it is imperative that researchers conduct exploratory analyses
in addition to confirmatory analyses. However, results from exploratory analyses should
be clearly labeled as such to avoid misleading readers to having excessive confidence in
the result. Ideally, then, a study will test pre-​registered confirmatory hypotheses that
replicate and expand an original result and conduct exploratory analyses. Compelling
results from the exploratory analyses can then serve as a priori hypotheses in future
confirmatory research. Finally, some of the most exciting exploratory research may
come from analyzing old data in new ways. For example, Voytek (Chapter 23) proposes
exciting new analytical methods that researchers can apply; his signal processing scripts
are freely available (https://​voytek​lab.com/​code), and researchers can use these scripts
to analyze their old data or other openly available data.
 INTRODUCTION   13

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 CHAPTER 2

L O GIC BEHI ND E E G
F REQU ENCY A NA LYSI S
Basic Electricity and Assumptions

KYLE J. CURHAM AND JOHN J. B. ALLEN

2.1 Introduction

The brain is an electrochemical machine. Although nerve cells differ greatly in their
size and morphology, all pass messages using electrical signals, sending information
throughout the brain and to the rest of the body via the spinal cord. Within any neuron,
an action potential is a wave of electrical activity that travels along the nerve membrane.
Action potentials are triggered by the summation of input from other neurons using
chemical neurotransmitters, which create voltage potential changes in the post-​synaptic
neuron. Surface-​recorded EEG is blind to the activity of single neurons but can non-​
invasively measure the electrical activity resulting from summated excitatory and in-
hibitory post-​synaptic potentials over millions of these signals in humans. EEG has
excellent temporal resolution, but poor spatial resolution; it can tell us when something
is happening in the brain, but not precisely where it is happening.
EEG gives an incomplete picture of the electrical activity occurring in the brain. The
cortex is the outermost layer of the brain, and the primary generator of the electrical
activity we measure with EEG. Our ability to detect cortical activity largely depends
on the parallel arrangement of cortical pyramidal neurons. When millions of parallel
neurons fire simultaneously, their electrical activity adds together to generate a signal
large enough to detect at the scalp. However, when neural populations fire incoherently,
or when they are not arranged in a parallel formation, the electrical activity does not
add constructively, so there is no observed signal at the scalp. This chapter introduces
basic concepts in electricity in signal processing, with some practical considerations
for data collection and analysis, to help readers understand EEG measurement and
interpretation.
16    KYLE J. CURHAM and JOHN J. B. ALLEN

2.2 Electricity—​Voltage, Current,

and Resistance

Every physical thing is made up of atoms, which in turn are made up of fundamental
particles including protons, neutrons, and electrons. Protons have a positive charge,
neutrons have a neutral charge, and electrons have a negative charge. Charges of the
same sign repel, and opposite charges attract. A few simple experiments demonstrate the
existence of electrical charge. For example, rubbing a balloon on a wool sweater makes
the balloon negatively charged as electrons move from the wool to the balloon (Figure
2.1). The degree of attraction or repulsion between two point charges is proportional
to the product of the charges divided by the inverse squared distance between them.
Therefore, when the electron-​rich balloon is brought into proximity of the electron-​
poor wool, or any neutral surface such as a piece of paper or the wall, the balloon will
be attracted. If you subsequently rub a second balloon in the same way, the two balloons
will repel each other since they are both negatively charged.
Most of the time, atoms have equal numbers of protons and electrons. However,
atoms can become ionized when electrons are removed or added, resulting in a net
charge. Electricity is the phenomenon that describes the behavior and movement of
charge. In general, it doesn’t matter whether it is the electrons or ions that are moving.
The flow of charge can be accomplished either by the transfer of electrons from atom to
atom, or, in the case of electrophysiology, by the diffusion of charged ions across cellular
membranes.
The degree to which electrons are free to move from atom to atom varies by material
type. For example, in metals, the outermost electrons are so loosely bound that they
freely move in the space between atoms at room temperature. Because these unbound
electrons are free to travel from atom to atom, they are called free electrons. The relative
mobility of electrons within a material is known as electrical conductivity. Conductivity
is determined by the types of atoms in a material, and how the atoms are linked to-
gether with one another. Materials with few or no free electrons are called insulators,
and materials with many free electrons are called conductors. The directed motion of
electrons is called electrical current. Just like water flowing through a pipe, electrons
move within the empty space between atoms. Under normal conditions, the motion of
free electrons in a conductor is random, with no particular direction or speed. However,
electrons can be influenced to move in a coordinated fashion through a conductive ma-
terial by supplying a voltage. Voltage is the “pressure” that pushes on free electrons to
cause them to flow. The ability of a current to flow from one location to another depends
on the resistance. In insulating materials, such as glass or rubber, electrons have little
freedom to move from atom to atom. The less freedom electrons have to move from
atom to atom, the greater the resistance to the flow of charge. A conductor’s resistance
generally increases as its length increases or its diameter decreases. It is again useful to
refer to the water analogy: water can flow more easily through a short, wide pipe than a
long, narrow pipe. The international system of units (SI) of current is called the ampere.
 LOGIC BEHIND EEG FREQUENCY ANALYSIS    17

(a)

Positive Charges and Electrons are

Present in Equal quantities in the
Sweater and the Ballon

(b)

Electrons are Transferred

from the Sweater to the Balloon

Figure 2.1 Before rubbing the balloon against the sweater (A), no net accumulation of electrons
exists on the balloon. After rubbing the balloon on the sweater (B), the balloon has accumulated an
excess of electrons, and the resultant negative charge of the balloon and positive charge of the sweater
creates a force of attraction sufficient to keep the balloon from being pulled to the ground by gravity.
Figure credit: K. Ehrmann.
18    KYLE J. CURHAM and JOHN J. B. ALLEN

Table 2.1 Guide to electrical symbols and terminology

Symbol Term Definition Unit

E Voltage Electromotive force Volts (V)

I Current Rate of flow Amperes (A)
R Resistance Opposition to current Ohm (Ω)
C Capacitance Ratio of the change in charge Farad (F)
to the change in voltage
P Power Rate of work Watt (w)
W Energy Ability to do work Watt-​second (Joule)

One ampere is defined as one coulomb of charge (or 6 × 1018 electrons) flowing past a
given point in a conductor in one second. The volt is the unit of pressure, that is, the
amount of electromotive force (EMF) required to push a current of one ampere through a
conductor with a resistance of one ohm, or 1 volt/​ampere.
When resistance is high, electrons tend to gather on one side of the insulating material
and the positive ions tend to gather on the other, effectively storing potential energy in
an electric field. At some point, the voltage across the material will exceed a threshold
known as the dielectric constant, at which point current begins to flow. This tendency
for high resistance to result in charge separation is known as capacitance. The amount of
charge stored in the capacitor is directly proportional to the surface area of the dielectric
(the electrical insulator polarized by the electric field) (Table 2.1).

2.3 Circuits

An electrical circuit consists of closed conductive paths between circuit elements.

Elements may consist of resistors, capacitors, voltage sources, or current sources.
Electrical components may be wired in series or parallel with each other (Figure 2.2).
In many cases, circuits consist of some complex combination of series and parallel
components. However, these circuits may often be represented by simpler equivalent
circuits with identical electrical properties.
To gather some intuition for the flow of current in simple circuits, we refer back to
the water analogy. The resistance increases as the pipe gets longer. This is equivalent to
stringing together multiple resistors in series. The rule to combine resistors in series is
additive:

Rseries = R1 + R2 + …+ Rn
 LOGIC BEHIND EEG FREQUENCY ANALYSIS    19

(a) R1

+
V1 R2

R3
(b)

+
V1 R1 R2 R3

Figure 2.2 Simple series (left) and parallel (right) circuits. Circuit components labeled with R
indicate resistors. Circuit components labeled with V indicate voltage sources.

Conversely, adding multiple resistors in parallel will decrease the overall resistance:

1 1 1 1
= + + …+
R parallel R1 R2 Rn

As more “pipes” are added, the water has more paths to escape, decreasing the overall
resistance to flow. Capacitors wired in series or parallel follow the same rules, but
reversed:

1 1 1 1
= + + …+
Cseries C1 C2 Cn

C parallel = C1 + C2 + …+ Cn

Most circuits are some complex combination of series and parallel. We can approach
these circuits one piece at a time, deriving a new equivalent circuit at each step
(Figure 2.3).
The next few sections explore examples of equivalent circuit representations, and
we use equivalent circuit representations to learn the voltages and currents at every
point in the original complex circuit. We later show how models of neurons can
be represented as a simple equivalent circuit of capacitors, resistors, and voltage
sources.
20    KYLE J. CURHAM and JOHN J. B. ALLEN

(a) R1

+
V1 R1 R2 R3

(b) R1

+
V1 Req

(c) R1

+
V1 Req

Figure 2.3 Reducing a complex circuit (left) to a simple equivalent circuit (right). In an inter-
mediate step, we combine parallel resistors 2 and 3 (middle). Next, we combine resistor 1 with the
equivalent resistor from the intermediate step.

2.4 Direct current

Circuits come in two basic flavors: direct current (DC) and alternating current (AC).
The difference depends on whether the voltage and current change directions over
time. DC circuits maintain currents flowing in a constant direction within a closed
loop, whereas AC circuits have current that repeatedly reverses direction. A DC electric
source feeds from one terminal to a set of circuit elements and then back to the other
terminal, in a complete circuit. Figures 2.2 and 2.3 are both DC circuits due to their con-
stant voltage power source. Note some resistors are connected in parallel, while others
are connected in series. At each step, we can combine resistors according to the rules in
Section 2.3 to derive a simpler equivalent circuit.
Some circuits may contain both resistors and capacitors. These are known as RC
circuits. In a simple circuit with one resistor and one capacitor in series, the capacitor
must discharge through the resistor. This discharge occurs at an exponential rate
determined by the RC time constant. The time constant indicates the number of seconds
for the capacitor to become 63.2% charged, or, equivalently, the time for current flow
 LOGIC BEHIND EEG FREQUENCY ANALYSIS    21

to have slowed by 63.2% from its starting value. This choice of time constant has an in-
tuitive explanation: at any moment in time, the rate of change in voltage is equal to the
voltage divided the time constant. For example, a 1 mF cap and a 1 kΩ resistor yields a
time constant of one second. If the capacitor is charged to 5 volts, the voltage will fall at
a rate of 5 V/​s. If the capacitor is charged to 2 volts, the voltage will fall at a rate of 2 V/​s.

2.5 Alternating Current

In contrast to DC signals, some sources of electricity produce AC, where voltages and
currents periodically reverse direction and switch back and forth between positive and
negative polarity. The electricity that comes from an American wall outlet is an example
of AC. The current in North America is 120 VAC and changes direction 60 times per
second. AC circuits can exhibit more interesting behaviors than DC circuits. For ex-
ample, at low frequencies, a capacitor acts like an open circuit, so no current flows in the
dielectric. However, when driven by an AC source, a capacitor will only accumulate a
limited amount of charge before the potential difference changes polarity and the charge
is returned to the source. The higher the frequency, the less charge will accumulate and
the smaller the opposition to the current.
Both resistors and capacitors resist the flow of current when a voltage is applied.
However, unlike in DC circuits, resistance may be frequency dependent. Frequency-​
dependent resistance is known as impedance, a complex-​valued quantity that can be
broken into two parts: magnitude (the ratio of the voltage amplitude to the current
amplitude) and phase (quantifies how much the current lags the voltage). Alternatively,
we can break impedance down into its real and imaginary parts. Like in DC circuits,
the real part of the impedance acts like resistance, resisting the flow of electric current.
The imaginary part is called reactance, and it quantifies the opposition to a change in
the current of a capacitive circuit element. Ideal capacitors are purely reactive, that is,
they have zero resistance, and the impedance of a resistor is purely real, or resistive.
Note this implies the current in a capacitor always lags the voltage by 90°. Impedance
devices add like resistors in a DC circuit. For a set of components in series, the total im-
pedance is the sum of the component impedances. To obtain the impedance of parallel
circuit components, the inverse total impedance is given by the sum of the inverses of
the component impedances.
Using different combinations of resistors and capacitors, RC circuits can be used to
attenuate some frequencies, while allowing others to propagate through the circuit un-
affected. For example, wiring a resistor in series with a load, and a capacitor in parallel
with the same load, significantly attenuates high-​frequency signals. Conversely, wiring
a resistor in parallel with the load, and the capacitor in series, attenuates low-​frequency
signals. In Figure 2.4, at low frequencies, the reactance of the capacitor will be very large
compared to the resistance of the resistor. This means that the voltage across the cap-
acitor will be much larger than the voltage across the resistor. At high frequencies the
22    KYLE J. CURHAM and JOHN J. B. ALLEN

reverse is true: the voltage across the resistor is larger than across the capacitor. In other
words, low frequencies pass to the output, and high frequencies are attenuated. This is
known as a low-​pass RC filter. Similarly, we can construct a high-​pass filter by swapping
the resistor and capacitor. The frequency cutoff for these filters is determined by the
time-​constant of the circuit, which is derived from the resistance and capacitance. The
cutoff frequency for an RC circuit is:

1
fc =
2πRC

In the low-​pass configuration, frequencies just above the cutoff are attenuated to half
their original amplitude. Conversely, in the high-​pass configuration, frequencies just
below the cutoff are attenuated to half amplitude. The amount of attenuation increases
as you move farther beyond the cutoff frequency. Figure 2.4 shows the amplitude roll-​off
as a function of frequency for a low-​pass filter.

(a)

R1

Vin C1 Vout

(b) Bode Diagram

0
Magnitude (dB)

—10

—20

—30
0
Phase (deg)

—45

—90

Frequency (Hz)

Figure 2.4 Low-​pass RC filter (left). The parallel arm with the capacitor provides a low im-
pedance path for high frequency signals. However, the capacitor saturates for low-​frequency
signals, providing a high-​impedance path. Low-​frequencies signals are thus preferentially
observed at Vout. Frequency response of the low-​pass RC filter (right). The signal is not appre-
ciably attenuated below the cutoff frequency of 50 Hz (shown in red). At frequencies just above
50 Hz, the signal magnitude is cut in half. As frequency increases, the amount of attenuation
increases.
 LOGIC BEHIND EEG FREQUENCY ANALYSIS    23

2.6 Hodgkin–​Huxley Model

We can use a simple circuit model to describe the electrical properties of neurons,
including the initiation and propagation of action potentials. Action potentials are the
result of the diffusion of sodium and potassium ions across neural membranes. The
Hodgkin–​Huxley model treats each component of a neuron as an electrical element in
the circuit (Figure 2.5), where current is propagated by the movement of ions across cell
membranes.
The cell membrane is represented by a capacitance (Cm). Cellular membranes are
highly resistive, and act as a dielectric material due to their relatively impermeability.
In the absence of special proteins called ion channels, ions like sodium and potassium
are unable to diffuse across the membrane, effectively turning the membrane into the
dielectric of a capacitor. As ionic currents add or subtract from the charge accumulating
inside the neuron, ions line up along the cell membrane. The differing concentration
of ions on either side of the membrane results in a net voltage potential, represented by
voltage sources (En). To maintain these concentration gradients, neurons have active
sodium-​potassium pumps that exchange two sodium ions into the extracellular space
for three potassium ions in the intracellular space.
Sodium and potassium ion channels are represented by electrical conductances (GNA,
GK) that depend on both voltage and time. As the voltage potential increases, the per-
meability of the membrane is selectively modulated, that is, conductance is increased,
for specific ion species. The flux of sodium or potassium ions across the membrane is
represented by ionic currents (Ip). Since the membrane is not perfectly impermeable,
leak channels are also included, represented by another conductance (GL).
The Hodgkin–​Huxley circuit model exhibits similar dynamics to real neurons. The
amount of injected current controls the emergence of a stable limit cycle. For a suffi-
ciently large input current, the circuit will exhibit repeating “action potentials” at a min-
imum firing rate. This means that either the neuron is not firing at all (corresponding
to zero frequency) or is firing at the minimum firing rate. Increasing the injected
current beyond the minimum threshold increases the firing rate of the neuron. When

Vm
inside
INa IK IL

GNa GK GL
CM

ENa EK EL

outside

Figure 2.5 Equivalent circuit diagram for the Hodgkin–​Huxley model.

©2003 James M. Bower and David Beeman.
24    KYLE J. CURHAM and JOHN J. B. ALLEN

the neuron fires, a series of channel activations occur to produce the action potential.
As the membrane potential approaches threshold, sodium ion channels begin to rap-
idly open, depolarizing the membrane (i.e., discharging the capacitor). The influx of
sodium changes the voltage gradient between the intracellular and extracellular space,
increasing the membrane potential. Once the polarity of the potential changes direction
(when enough sodium ions have crossed the membrane), sodium ion channels begin
to deactivate (decreasing sodium conductance). As the sodium channels close, potas-
sium channels begin to open (increasing potassium conductance), resulting in an efflux
of potassium ions to the extracellular space, restoring the membrane potential to the
resting state following a brief hyperpolarization.

2.7 Filtering

As discussed, simple RC circuits can selectively attenuate certain components or

features of a signal, and not others. This is known as filtering. Filters come in a variety of
forms (RC circuits, mechanical and optical filters, digital signal processing, etc.). Here
we examine two of the most common digital filters used in electrophysiology. The re-
sponse of a system to a brief input signal, or impulse, is called the impulse response. The
impulse response of a finite impulse response (FIR) filter settles to zero in finite time.
Given a finite sample of nonzero input values, an FIR filter will always yield a finite
sample of nonzero output values. This contrasts with an infinite impulse response (IIR)
filter, which does not settle in finite time. The RC filters seen in Section 2.6 are examples
of IIR filters since the capacitors (or inductors) in the RC filter never completely relax
following an impulse.

2.8 Analog-​to-​Digital
Signal Conversion

Most FIR filters are implemented using digital signal processing. Our discussion of elec-
tricity thus far has dealt with analog signals, which are continuous in both time and in
voltage. An economy of representation can be achieved by sampling discrete points in
both the time and voltage domains, a process of creating a digital signal, which has a tem-
poral resolution determined by the sampling rate and a voltage resolution determined
by the resolution of the analog-​to-​digital converter (Figure 2.6). For example, a 16-​bit
converter will allow 1016 of 65,536 discrete voltage values, and a sampling rate of 1,000
Hz will allow one value every millisecond.
With sufficiently large sampling rates, a digital signal can closely approximate the
analog signal it is attempting to represent (Figure 2.7). However, several considerations
 LOGIC BEHIND EEG FREQUENCY ANALYSIS    25

2 Discrete-time sampled 2 Digitally sampled (3 bits)

1.5 1.5
1 1
0.5 0.5
0
µV

0
–0.5 –0.5
–1 –1
–1.5 –1.5
–2 –2
0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8 0 0.2 0.4 0.6 0.8 1 1.2 1.4 1.6 1.8 2
Seconds Seconds

Figure 2.6 A signal sampled at 20 Hz. Discrete-​time sampling (left panel) allows for con-
tinuous y-​axis (μV) values, whereas digitally-​sampled signals (right panel) must use a limited
number of y-​axis values. The three bit converter illustrated here (right panel) allows for 23 =​8 dis-
tinct values, providing only a coarse approximation of the signal voltage. The right panel depicts
the discrete sample value (red circle) and the 3-​bit digital equivalent (red line), and the discrep-
ancy (dashed vertical black lines).

are essential to ensure signal fidelity when digitizing an analog waveform. In order to
recover all components of a periodic waveform, it is necessary to use a sampling rate at
least twice the highest waveform frequency. This is known as the Nyquist sampling rate,
and it determines whether or not aliasing will occur. Similarly, for a given sampling rate,

20 Hz Sampling 40 Hz Sampling 100 Hz Sampling

2 2 2
1 1 1
4 bits
µV

0 0 0
—1 —1 —1
—2 —2 —2
0 0.5 1 1.5 2 0 0.5 1 1.5 2 0 0.5 1 1.5 2

2 2 2
1 1 1
5 bits

0 0 0
µV

—1 —1 —1
—2 —2 —2
0 0.5 1 1.5 2 0 0.5 1 1.5 2 0 0.5 1 1.5 2

2 2 2
1 1 1
8 bits

0 0 0
µV

—1 —1 —1
—2 —2 —2
0 0.5 1 1.5 2 0 0.5 1 1.5 2 0 0.5 1 1.5 2
Seconds Seconds Seconds

Figure 2.7 A comparison of a signal (black line) sampled (red line) at three sampling rates (20,
40, 100 Hz) and using three different converter resolutions (4-​bit, 5-​bit, and 8-​bit) that allow for
16, 32, and 128 distinct μV values. Low bit-​resolution was used here for illustrative purposes; com-
mercial converters are typically 12-​bit (4,096 values) or 16-​bit (65,536 values).
26    KYLE J. CURHAM and JOHN J. B. ALLEN

Signal Aliasing

18Hz @ 20 samples/s

0 0.1 0.2 0.3 0.4 0.5 0.6 0.7 0.8 0.9 1

Time (s)

Figure 2.8 Signal aliasing due to insufficient sampling rate. Samples from an 18-​Hz signal were
sampled at a rate of 20 Hz. Identical samples are obtained for a 2-​Hz sine wave sampled at 20 Hz.
These digitized signals are indistinguishable from each other.

the highest frequency signal that can be represented is one-​half the sampling rate, and
this is known as the Nyquist frequency. Aliasing occurs when the set of samples obtained
from the analog signal are indistinguishable from samples from a lower-​frequency
signal. For example, in Figure 2.8, an 18-​Hz signal is sampled at 20 Hz, well below the
Nyquist sampling rate. For this 20-​Hz sampling rate, the Nyquist frequency is 10 Hz.
Although the signal is in fact an 18-​Hz signal, the sampled signal appears as a 2-​Hz sine
wave, so the signal is not well characterized. In general, signals that are x Hz above the
Nyquist frequency will appear as a signal x Hz below the Nyquist frequency. Sampling
above the Nyquist frequency will prevent aliasing but may still not characterize the
signal well in the time domain. As a general guideline, it is recommended to sample at
least 5× the highest frequency of interest to get a good signal.

2.9 Digital Filtering

FIR and IIR digital filters can be used to process digital signals. A simple moving average
is an example of an FIR filter. If we take the average across the last five data samples
and shift the five-​sample window forward by one sample at each timestep, the result is
a moving average window. We can denote the value of the filtered signal (x) at the nth
timepoint using summation notation:

2
1
y (n) = ∑ 5 x (n − k )
k = −2
Another random document with
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his illegal barter with the Indians to the great consternation of the
Pilgrims.
Beaver, otter and valuable deer skins found their way in great
quantities to the new truck house, while Merrymount became “a sort
of a drunkard’s resort and gambling hall” and worse. The jolly host of
Merrymount, revelling in “riotous prodigality,” had a great Maypole
erected for the entertainment of visiting factors who wanted to dance
and frisk with pleasurable Indian maids. Fishing and trading vessels
along the coast much preferred to do business with the open-handed
Morton rather than the close-fisted Pilgrims. Therefore, trading
goods were as easily come by at Merrymount as were beaver skins.
Everyone was quite happy about the situation except the fathers at
Plymouth.
Morton had even discovered their profitable new trading grounds
in Maine. The first year the Pilgrims extended their fur trading
operations to Maine, in 1625, they gathered in 700 pounds of beaver
besides some other furs on the Kennebec, mostly in exchange for
the corn they had by then learned to grow at Plymouth. But now
Morton was outbidding them with his more attractive trading goods
and getting nearly everything of value in that vicinity too. The very
existence of the Plymouth Colony seemed to be at stake.
Complaints, cajolings and threats were of no avail. Morton, the
lawyer, was too clever. Something had to be done and force was the
final resort. Captain Standish was sent with some soldiers to arrest
the obnoxious neighbor on the charge that he was violating the royal
proclamation prohibiting trade in guns and powder.
“Captain Shrimp,” as Morton contemptuously called Standish,
succeeded in his mission only after a fight. He captured Morton and
took him to Plymouth. There he tried his best to have the erstwhile
“host of Merrymount” hanged. But in the end Morton’s only
punishment was to be shipped back to England.
As it came about, the Wessagusset area was no sooner
eliminated as a competitor for the beaver trade than the Pilgrims
were faced with new and much more formidable rivals in the same
neighborhood. These were the Puritans, the advance guard of whom
arrived at Salem under Captain John Endicott in 1628. As recruits of
the great Massachusetts Bay Company they were soon coming by
the thousands. Under the leadership of Governor John Winthrop
they overran the Boston Bay region. And there wasn’t much the
fathers at Plymouth could do about these new neighbors except to
offer religious advice on the relative merits of Brownism and
Separation.

The local competition for beaver now being almost overwhelming,

the Pilgrims found it necessary to go more and more afield to meet
their required quota of pelts. Excursions north to the Kennebec in
Maine were the most fruitful, not only because there was much fine
fur to be had from the natives there but because trading goods on
occasion could be obtained from fishing and trading ships off that
coast. Once, too, in exchange for corn the Pilgrims picked up four
hundred pounds worth of trucking stuff from some Englishmen who
were abandoning their plantation on Monhegan Island. And they
acquired an additional stock of truck from a French ship which had
been wrecked at Sagadahoc.
To the south along the coast they never met with much success.
When they had first gone out to Narragansett Bay on a trading
voyage in 1623, for instance, the Indians there disdained their
meagre offerings. The Narragansett were much too happy with the
goods being furnished by the Dutch traders from Manhattan. Both
the Narragansett and their neighbors, the Pequot, held on to their
furs for the Hollanders, and they were powerful enough to get away
with it. They had Dutch guns and powder, as well as the Hollanders
themselves if need be, to back them up. In fact, by 1625 the Dutch
had a fortified trading post on an island in Narragansett Bay and two
similar forts near-by on the mainland.
Governor Bradford, in exasperation, gave out a warning that all the
region along the coast to the southeast of Plymouth was English
territory. He as much as demanded that the New Netherlanders stop
trading there. But it was useless. The Dutchmen simply ignored the
warnings. However they did offer to enter into direct trading relations
themselves with Plymouth and sent a mission there in 1627, with
sugar, linen and other goods, to talk it over.
This mission was headed up by Isaack de Rasieres, the chief
trader as well as the Secretary of New Netherland. Rasieres appears
to have had an ulterior motive in making the visit. He brought along a
stock of wampum, the strings of highly polished shell-beads that the
Dutchmen had been accustomed to getting from the Narragansett
and the Pequot. With wampum, Rasieres pointed out to the Pilgrims,
he had been doing a great business among Indians who didn’t have
the means of manufacturing it, especially along the Hudson River
where Henry Hudson himself had first discovered strings of shell-
beads circulating as a kind of money.
The Hollander cannily suggested that wampum might be used to
just as great advantage on the Kennebec by the Pilgrims. No doubt,
by this means, he hoped to direct their attention more to Maine and
away from the Dutch trading preserves in Long Island Sound and the
Narragansett country. Offering to sell wampum to the Englishmen at
a fair price, probably he hoped also to keep them from dealing direct
for it with the Narragansett and the Pequot.
Although the Pilgrim fathers were a bit suspicious of Rasieres’
motives, nevertheless they did find the shell-money to be most
“vendable” on the Kennebec. So much so, in fact, that in a very short
time with the aid of this medium of exchange they were enabled to
cut off the fur trade of that region from the fishermen and other
independent traders who had been accustomed to barter there. In
the meantime they also developed their own sources of wampum
among some of the Massachusetts coastal tribes who, as it turned
out, also possessed the means of manufacturing it.
The Pilgrims secured their rights up the Kennebec River by
obtaining a patent to a strip of land fifteen miles wide on both sides
of that river. They built a trucking house at Cushenoc (Augusta)
which they kept stocked with coats, shirts, rugs, blankets, corn,
biscuit, peas, prunes, and other supplies. With the help of wampum
for exchange they drove a brisk trade among the Abnaki Indians in
those parts. In 1630 they extended their operations even farther
north, setting up a trading post on the Penobscot River at Pentagoet,
now Castine, Maine.
The Penobscot trading post originated as a private venture for
which the Pilgrim, Isaac Allerton, along with some partners in
England promoted a patent. Allerton, once a London tailor, had risen
in prominence at Plymouth in New England to stand second only to
the governor. It would appear that he and his overseas partners
shared the rights to trade in the territory north of the Kennebec with
other leading Pilgrims in return for the loan of wampum, shallops,
supplies and servants.
But the partnership accounts became jumbled, because of
Allerton’s financial deceits according to the Pilgrims. And Edward
Ashley, the “profane young man” sent out from England as factor,
didn’t help things by his personal conduct.
Actually, young Ashley did well enough in the bartering
department, acquiring over a thousand pounds of beaver and otter
the first season. However he seems to have gone native in the most
offensive sense to the Pilgrim fathers, living “naked” among the
savages and committing “uncleannes with Indean women.” Also, it
was discovered, he was trading shot and powder with the savages
and not even accounting for the profits from this unholy trade.
Such behavior, of course, could not be countenanced. Ashley was
arrested and shipped back to England by the Plymouth partners,
while they themselves took over complete operating control of the
enterprise.
All of which was not to Isaac Allerton’s liking. Deserting the
partnership, he retaliated by setting himself up as a competitor in
Maine. In 1633 he settled some “base fellows” in a new trading post
at the mouth of the Machias River, close by the present Canadian
border. There he did his best to cut off Plymouth’s commerce with
the more northerly tribes who were then taking their furs to the
Penobscot and Kennebec trucking houses.
 The Pilgrims had other competitors in Maine also. In 1630 a
trading post was established at Pemaquid Point under a patent
obtained by two English merchants for twelve thousand acres
between the Damariscotta and Muscongus Rivers. About the same
time John Oldham and Richard Vines obtained a grant at the mouth
of the Saco River, which remained a most profitable fur trading
center for some years.
Then there was the Laconia Company, organized by Sir
Ferdinando Gorges and Captain John Mason to put new life in their
Piscataqua colony. Laconia, described as a vast hinterland area of
rivers and lakes, was an extension of their original patent to the
“Province of Maine.” Gorges and Mason now planned to send
cargoes of Indian trucking goods up the Piscataqua River into
Laconia, to Lake Champlain, to be bartered for peltries. Thus they
hoped to compete with the Dutch and the French for the hinterland
trade.
Captain Walter Neale, as governor of the Laconia Company, and
Ambrose Gibbons, as factor, did very well from 1630 to 1633,
establishing several trading posts on the Piscataqua River. But they
never reached Lake Champlain. Maybe they had it confused with
Lake Winnepesaukee. In any case, as they discovered, the rivers of
Maine flowed from the north, not from the west, and they couldn’t
penetrate deep enough into the interior to tap the hinterland trade of
the Dutch and the French.
Probably the most dangerous of the Pilgrims’ rivals in Maine about
this time was the trading post on Richmond’s Island off Cape
Elizabeth. Thomas Morton, the jolly host of Merrymount, had traded
here as early as 1627. After his banishment from America one of his
most roguish associates, Walter Bagnall, took over the island and is
said to have gained 1,000 pounds sterling from his trade in a period
of three years. Then this “wicked fellow” was murdered by the
natives.
Bagnall’s successor, John Winter, also did a flourishing business
on Richmond’s Island as factor for some English merchants,
employing some sixty men at one time in both fishing and fur trading
activities. The records indicate that he was about as unscrupulous in
his dealings as was his predecessor, cheating and otherwise
mistreating the Indians at every turn. He charged them at the rate of
thirty-three pounds for a hogshead of brandy which cost him seven.
For powder which cost him twenty pence per pound he raised the
rate to three shillings in trade. But the natives preferred his goods.
In spite of such competition, however, Maine became the Pilgrims’
chief source of furs. In one period of two years the trading post at
Cushenoc alone is known to have gathered in more than 7,000
pounds of beaver. The route of the beaver-laden shallops from the
Maine coast was indeed Plymouth’s life-line. Wampum played no
small role in the success of this extended operation which was so
vital to the colony’s existence, although the tight control exercised
over the fur trade by the Pilgrim fathers was the main factor.
Always the peltry traffic had been invested in certain leaders at
Plymouth who managed the whole trade in the interests of the
communalty in order that the Pilgrims’ debt to the merchants in
England might be paid. In 1627 William Bradford, Isaac Allerton,
Edward Winslow, Miles Standish and a few others in partnership
undertook to assume this entire indebtedness. In return, these
“undertakers” were to enjoy any and all profits from the traffic in
peltries for six years. They were empowered to do what they pleased
with all furs and trucking goods in the common store, and they alone
were to use the colony’s trading posts, truck houses and shallops.
From that time forward the Plymouth Plantation enjoyed real
economic success. In 1628 furs to the value of 659 English pounds
were exported in one cargo. Although no total figures are available
for 1628 to 1630, a very large amount of peltry went to England
according to Governor Bradford. From 1631 to 1636 over 12,000
pounds of beaver and 1,000 pounds of otter were shipped, most of
which brought 20 shillings a pound and none less than 14 shillings.
Bradford said that the beaver alone during these years brought
10,000 pounds sterling and that the otter was more than sufficient to
pay all costs of transport and auction. Considering the size of the
plantation this was indeed big business.
 In spite of the financial bungling of the partnership’s agents in
England, and even outright irregularities there, the debt was paid off.
Additional profits from the sale of beaver skins bought many other
things the Pilgrims needed, and in the end it was the earnings from
the fur trade that provided the foundation for Plymouth’s next
economic development, cattle farming.
In the meantime the Pilgrims’ profitable trade in furs on the Maine
coast and on rivers leading to the interior was jealously regarded by
the French who, after all, considered that territory within their own
limits. And neighboring French traders were doing something about
it.
 X
A Border Fixed on the Coast of Maine
THE planting of trading posts farther and farther up the coast of
Maine by the New Englanders did not go unchallenged by the
French. Although in English eyes it had been well established ever
since Captain Argall’s raids in the Bay of Fundy that English claims
to the coast now extended well above the 45th parallel, even to the
St. Lawrence valley, no such admission had ever been made by the
French. There were always French traders in Acadia (Nova Scotia)
who envisaged their preserves as extending well down the coast of
Maine and who intended to resist further encroachments in those
parts.
In fact, as it came about, these fur traders protected and
maintained the French claim to much of this coastal region during an
incident of the Thirty Years’ War which put most of New France
under the English flag for three years, from 1629 to 1632.
Prior to this episode the affairs of New France had reached a
turning point. In 1627 Samuel de Champlain’s difficulties became
acute. That year the Iroquois, his irreconcilable enemies, renewed
their bloody savagery in the interior, while along the coast the
English were closing off the very entrance to the valley of the St.
Lawrence with their plantations and trading posts and their increased
shipping.
Although Champlain’s company had a good season in furs,
collecting 22,000 that summer which could be sold in France at ten
francs each, the expense of operation had made the outlook none
too bright relatively for the shareholders who expected huge returns.
Besides the salaries of the Viceroy in France and of Champlain as
his lieutenant governing the colony, interpreters now cost as much
as a thousand francs a season and sailors six hundred, while factors
and other servants came correspondingly high. And there were other
increased expenses that bid fair to cut deep into profits.
As a result the directors of the company made things difficult for
their governor on the St. Lawrence. They particularly balked at his
urgent request for extra funds to strengthen the fortifications of
Quebec, even though only a wooden palisade and a few small
cannon protected the French trading post.
Champlain, with characteristic fortitude, made the best of his
difficulties. But he was most uneasy about the English at his
doorway. And well he might be!
As early as 1620 Sir George Calvert, English Baron of Baltimore,
had adventured a plantation in Newfoundland. Three years later King
James granted him quasi-regal proprietorship of the southeastern
peninsula of that island between Trinity and Placentia Bays. In 1627
he came out with his family, built a fine house at Ferryland, and
showed every intention of establishing a great fur trading and fishing
colony in the form of a British dominion at this strategic gateway to
New France.
Still more alarming were the activities of a Scotsman, Sir William
Alexander, who had been stirred by accounts of the founding of New
Spain, New France, New Netherland and New England to attempt
the founding of a New Scotland. In 1621, with complete disregard for
French claims, the English king had granted to him all the vast
peninsula between the Atlantic Ocean and the Gulf of St. Lawrence!
Today that would be Nova Scotia, New Brunswick and part of
Quebec Province. This princely domain was to be divided into
feudal-type Scotch baronies complete with a hierarchy of hereditary
titles.
Alexander’s attempts to establish permanent trading posts in New
Scotland, or Nova Scotia, were failures until 1628. That year
seventy-two of his colonists managed to set up for business at
Poutrincourt’s old quarters at Port Royal in the Bay of Fundy. There
they found themselves to be the near neighbors of a young French
fur trader at Cape Sable named Charles de la Tour who was under
the impression that the country belonged to him.
 Monsieur La Tour had inherited his rights from Saint Just after that
pioneer finally returned to France in 1623. Since then he had been
living the independent life of a wilderness lord with French retainers
and aboriginal subjects, and he had no intention of yielding his
estate or fur-trading privileges to anyone, much less foreigners.
La Tour did not immediately attempt to oust Alexander’s Scots
from Port Royal as he was on much the weaker side at the time, but
he did stand firm and unsubmissive at Cape Sable and he resisted
all efforts of the English to bring him over to their side. This was not
easy, for nearly all of New France was soon in the hands of the
enemy.
In 1628 a French colonizing fleet sent out by the Company of New
France under the aegis of Cardinal de Richelieu was captured by
privateers boldly operating in the Gulf of St. Lawrence with English
letters of marque. In another year Champlain’s worst fears were
realized when this same fleet of privateers, under the leadership of
the Kirke brothers, appeared before Quebec in force enough to
insure its capitulation.
Champlain himself was taken prisoner at Quebec. But under the
terms of the surrender he marched out of the fort with his arms and
all of his own furs, as did other head men. The factors, servants and
workmen were allowed to take but one beaver skin each before the
English took over the factory. Then, after running up the English flag,
the Kirkes were said to have driven an immediate trade with Indians
and others remaining about the fort for some 2,000 skins in addition
to those they found in the storehouse. They even ended up with the
pelts they had allowed the starving garrison under the terms of
surrender, taking them in trade for food.
Sir William Alexander, uniting his interests with the Kirkes to form
the Scottish and English Company for the peltry trade of New
France, sent out a second fleet to Port Royal in 1629 under the
command of his son. When the fleet arrived nearly half the people
who had been left at Port Royal the year before were found to have
died from one cause or another. But young Alexander relieved the
survivors, strengthened the fort, and put new life into the colony. His
vessels managed to acquire a satisfactory number of pelts in the Bay
of Fundy during that summer. He also captured a French ship. And
as further proof of the substance of New Scotland he took back on
his return voyage an Indian chief of the region who wanted to
conclude an alliance with the English king.
In the meantime it developed that shortly before Champlain’s
capitulation a treaty of peace had been signed between England and
France. But, although King Charles agreed very soon to restore
Quebec to the French under the terms of the treaty, the negotiations
dragged on for three years. During that time the Scottish Company
continued to drive a great trade for pelts on the St. Lawrence and
elsewhere, while the French pressed not only their claims to furs
appropriated from stores at Quebec after the actual signing of the
peace treaty but insisted also upon the return of all of New France.
That included Acadia of course and the evacuation of the fort at Port
Royal.
At first, Charles was not disposed to disturb Alexander in his
occupation of “Nova Scotia.” In fact he encouraged the Scot to
further efforts in that direction. However, he needed gold at the time
much more than he needed another Scotland, so he was not
reluctant to do a little bargaining. When his wife’s brother, the French
king, agreed to pay a long overdue and substantial dowry, Charles
agreed to return Port Royal to the French. Sir William’s dream was
thereupon finished. The Scots demolished their fort, and young
Alexander surrendered Port Royal to the Chevalier Isaac de Razilly
who had been sent out by the Company of New France as governor
of all Acadia.
This was a tremendous victory for Charles de la Tour and his
French fur traders, who had not only stubbornly maintained
themselves during the incident of the Scottish occupation of Acadia,
but had managed harassments intended to hinder further
encroachments up the coast of Maine by the Englishmen in those
more southerly parts.
In 1631 the Frenchmen had paid a visit to the Pilgrims’ most
northerly outpost at Pentagoet on the Penobscot. Taking advantage
of the absence of the factor and most of his company, they were able
to surprise a few “simple” servants by a ruse. First, pretending they
had “newly come from the sea” and that their vessel was in need of
repairs, a “false Scot” among them fell to admiring the Englishmen’s
muskets. Then, talking the servants into letting them examine the
guns, they gained possession of the Englishmen’s weapons and
promptly made away with some four or five hundred pounds worth of
Pilgrim goods, including three hundredweight of beaver.
In time, however, after the ousting of the Scots from the Bay of
Fundy, the Frenchmen were able to do more than simply harass the
Englishmen on the coast of Maine. The truck house at Machias in
Maine, built by Isaac Allerton in 1633, was even more of a challenge
than the Penobscot post had been. Allerton had no sooner settled
his traders there than “La Tour, governor of the French in those
parts, making claim to the place, came to displant them, and finding
resistance, killed two of the men and carried away the other three
and the goods.”
Allerton himself later went to Port Royal to protest. But he was told
by La Tour that “he had authority from the king of France, who
challenged all from Cape Sable to Cape Cod, wishing them to take
notice and to certify the rest of the English, that, if they traded to the
east of Pemaquid, he would make prize of them.” When Allerton
asked to see Monsieur La Tour’s commission the Frenchman replied
that “his sword was commission sufficient, where he had strength to
overcome; where that wanted, he would show his commission.”
And the French made good on their challenge, for in 1635
Monsieur Charles d’Aunay, one of Governor Razilly’s lieutenants,
came in a man-of-war to the Pilgrims’ trading post on the Penobscot.
By a show of this force he took possession of the trucking house in
the name of the King of France. Making an inventory of the goods he
found there at prices he set himself, d’Aunay “made no payment for
them, but told them in convenient time he would do it if they came for
it. For the house and fortification etc. he would not allow nor account
anything, saying that they which build on another man’s ground do
forfeit the same.” Then he put the English traders in a shallop and
sent them back to Plymouth.
 The Pilgrim fathers were enough upset about the loss of their
goods and trading post to do something violent about it. With the
approval of the Massachusetts Bay Colony, but without much in the
way of material assistance from their Puritan brethren, they hired “a
fair ship of above 300 tun well fitted with ordnance” to retake
Pentagoet. This ship was under the command of a Captain Girling. If
the captain was successful he was to have 700 pounds of beaver; if
not, nothing. Along with him went Captain Miles Standish and twenty
Plymouth men in their own bark to resettle their trading post after
Girling had driven out the French.
It would appear, however, that the French must have had notice of
the impending attack. So firmly were they entrenched behind
earthworks that Captain Girling’s gunfire could not dislodge them
and all the big ship’s powder was exhausted without effect. No
landing was attempted. Standish, frustrated, returned to Plymouth in
his bark, and Captain Girling went his own way.
The French now remained in permanent possession of what was
soon known as the “Mission of Pentagoet.” And so the line was
finally drawn as a practical matter between the English and the
French on the Maine coast, although England continued to maintain
officially that it was more northerly, at the 45th parallel.
Mutual distrust, born of the religious differences between English
Puritan and French Catholic, often erupted in charges and counter-
charges that sometimes threatened security on either side of the
line. In fact, fear of possible French aggression was in part
responsible for the eventual formation of the New England
Confederation in 1643. On the other side of the picture however,
there was much guarded trade between Frenchman and Englishman
as it suited their pocketbooks. As a matter of fact the Puritans
actually traded with the French conquerors of Pentagoet shortly after
d’Aunay captured the Pilgrim trading post.
But, even so, this coastal border created by the rival fur traders of
these two nations was maintained with only minor variations for
years, even after competition for the beaver trade was no longer the
reason for its existence.
 Acadian fur traders and the rivers of Maine and Massachusetts
that ran in the wrong direction had effectively contained the English
on the northern coasts. The St. Lawrence River route to the interior,
to the great lake country that teemed with fur-bearers, remained safe
to the French traders, who resumed their profitable westward
penetration of the hinterland.
It remained to be seen, however, if the French could prevent the
Dutch of New Netherland and the English of Virginia from draining
off this hinterland trade. The Connecticut, Hudson, Delaware, and
Susquehanna Rivers all ran in the right direction to tap it.
 XI
The Bay of Virginia
WHEN Sir George Calvert, Baron of Baltimore, founded his
province of “Avalon” in Newfoundland he was partly motivated by
religious urgings. Being a Catholic convert he earnestly wanted to
provide a haven in the new world for the persecuted Catholics of
England. But this court favorite, who as the son of a humble
Yorkshire farmer had figuratively pulled himself up by his own
bootstraps to ennoblement, was also a profit-minded promoter. He
expected a rich return on his newly gained proprietorship.
However, after investing many thousands of pounds at Ferryland,
Lord Baltimore had little to show in the way of profits. The future, too,
looked as bleak as the cold Newfoundland winters, as barren as the
sterile soil. True, there were fish to be had for the taking—weather
permitting. But the trade in pelts was only fair, for the more profitable
fur frontier had long since passed into the interior of Canada. And
agricultural pursuits, the traditional support of English feudal manors,
were almost impossible due to the inclemencies of the climate.
The Lord Proprietor of Avalon was soon looking southward, his
eyes resting appraisingly on Virginia. There, the warmer weather and
richer soil seemed to offer more appropriate conditions for the kind of
province he had in mind. There, in the lush tidewater of the
Chesapeake Bay country, he could hope for a better trade in pelts
too.
As for possible profits from Newfoundland cod, he would gladly
consign them to those hearty fishermen who by nature were better
able to cope with the rigors of the northern climate!
So it came about that in 1629 his lordship wrote to King Charles
asking that a precinct of land in Virginia be granted to him with the
same quasi-regal privileges he enjoyed under his Newfoundland
patent. Then, without waiting for a reply, he embarked for
Jamestown where he let it be known that he was looking for a new
plantation site, and forthwith began cruising about Chesapeake Bay
with the air of a man who was confident that the king would approve
his plans.
This was anything but politic. No Virginians, Anglican or Puritan,
wanted a Catholic in their midst. They didn’t want one even as a
neighbor in their great bay, and especially such an influential convert
as Baltimore. Who knew what Romish plot he might be promoting?
But even more alarming was the threat that this ambitious baron
posed to their fur trade in the Chesapeake.

The tidewater had come to represent a treasure house of pelts to

the merchants and factors on the James River, at Jamestown and at
Kecoughtan, and to scores of traders plying their shallops and
pinnaces in the bay. On the York, the Rappahannock and the
Potomac Rivers these traders regularly visited the villages of the
Indians, their shallops returning to the James River plantations laden
with skins. And across the bay on the Eastern Shore there was still
another lively fur trading center at Accomac where beaver and
muskrat abounded in the near-by streams and marshes.
The most profitable branch of this trade was that in the Potomac
River, where the Spaniards of the sixteenth century had been the
first of record to barter with the natives for pelts and hides. Captain
Samuel Argall, who succeeded Captain John Smith in 1609 as the
colony’s main support for trade with the Indians, made several early
visits to the villages of the Patowomecks there. For hatchets and
hoes, no doubt, he took good quantities of beaver, otter, raccoon and
deer skins as well as the corn that was so badly needed at
Jamestown. In 1612 he really whetted the Patowomecks’ appetite for
English goods when he produced a copper kettle for their chief.
Many English traders, with and without permits, followed Captain
Argall into the Potomac River, but especially members of the Virginia
Council who more or less controlled the Chesapeake trade as a
perquisite of office.
The Eastern Shore was opened up for trade in 1619 by Ensign
Thomas Savage who was one of the earliest professional factors in
the Chesapeake tidewater. He had spent several years as a youthful
hostage with Chief Wahunsonacock of the Powhatan Confederacy
after having been left with this old Powhatan in 1608 by Captain
Christopher Newport. During that time young Savage became a
favorite of the natives and learned much about the dialects and
customs of the tribes in the tidewater country. Later, when his
services as an interpreter came to be in demand by Virginia officials,
he was commissioned an “Aunchient,” or ensign, on the staff of the
Master of Ordnance at Jamestown. It was in this capacity on
expeditions of discovery in the Potomac and Patuxent Rivers and on
the Eastern Shore that he began bartering on his own account.
Like Captain John Smith before him, Ensign Savage learned of the
great trade in furs being had by the Frenchmen on the “backside of
Virginia.” In 1621 his account of their deeper penetration of the
hinterland reached the Virginia Company Council in London. At the
same time the company received reports of Dutch traders operating
along the Virginia coasts. A detailed account of this competition was
rendered by Thomas Dermer, the New England fur trader, who in
addition to having paid the visit to Jamestown went into the
Delaware and Hudson Rivers. He reported that in both those rivers
he had “found divers ships of Amsterdam and Horne who yearly had
there a great and rich trade in Furrs.”
The Dutch encroachments prompted a special letter, too, from the
governor and his council at Jamestown. They urged the company to
undertake “so certaine and gainefull” voyages for pelts as might be
made “to their infinite gaine” in the Delaware and Hudson, both of
which they pointed out were “within our lymits.” The Dutch
interlopers on the coasts were almost as challenging to the
Virginians, now that they themselves were well rooted, as was the
possibility of French competition creeping down the rivers that flowed
into Chesapeake Bay from the northern hinterland.
 Almost—but not quite as challenging. For one thing, the settlers on
the James at this time had neither the ships to trade coastwise nor
the trade goods necessary to compete with the Dutch. More
importantly, however, right at their back door was the vast tidewater
of the bay and, beyond that, great stretches of lake country in the
north, all fertile with fur bearers to be had for the taking—provided
they got there before the French. This could be accomplished with
home-made shallops and less fancy trade goods than were needed
to compete with the Dutch.
So it came about that the Virginians left any exploitation of the
Delaware and Hudson Rivers to the merchants in England and,
individually with their more limited resources, extended their own
trading operations in their “Bay of Virginia.”
Scores of immigrants participated in this tidewater fur trade,
including a great many who had the advantage of a knowledge of the
Algonquin tongue because of having cohabited with the savages.
Some of the latter were no more than villainous runaways, cautiously
returning from sanctuary with the natives to share the profits of the
boom in pelts. Others, such as Robert Poole and Henry Spelman,
had been left with the Indians as hostages when they were youths,
very much in the same manner as Thomas Savage. Like him, they
too had learned the dialects and habits of various tribes, acted as
spies for the English colonists and then, later, were useful as
interpreters.
Robert Poole traded professionally for the joint-stock, making
voyages in the company’s pinnace and rendering his accountings to
the colony’s treasurer at Jamestown. But he and Thomas Savage
also obtained commissions through members of the council and,
renting shallops from more prosperous colonists for voyages to the
villages of the Indians, did very well for themselves in this way.
Probably neither of them traded for pelts as extensively however as
did Henry Spelman who eventually managed to get financial backing
in England and ships of his own.
Young Spelman was the incorrigible scion of a distinguished
English family. According to his own testimony he had been sent out
to Virginia in 1609 at fourteen years of age because of “beinge in
displeasure of my fryndes and desirous to see other countries.”
Captain John Smith left him with Powhatan’s people, no doubt as the
best way to get rid of him. Henry Spelman was not as popular with
the natives as Thomas Savage, except perhaps with the Indian
maidens. In fact he appears to have escaped with his life only
because of the intervention of Pocahontas who, continuing her
interest in white men after the affair with Captain John Smith,
succeeded eventually to marrying an Englishman named John Rolfe.
Spelman’s entire career in the colony was one of manifold and
dangerous deceit. As a youth living with the Indians, he was accused
of treacherously laying a death trap for some of his fellow
Englishmen at Jamestown. Later, when he was employed as an
official interpreter for the colony, his tenure in office was cut short
after he was tried by the House of Burgesses and degraded from his
rank for inciting Opechancanough, the new Powhatan, against the
English governor.
In 1622, when Opechancanough finally loosed his stored-up
hatred against the English, killing some three hundred fifty men,
women and children along the James River in a surprise massacre,
Henry Spelman and a trading partner, Captain Raleigh Croshaw,
were bartering in the Potomac. Opechancanough conspired with
Chief Japazaws of the Patowomecks to have the traders murdered.
But the Patowomeck king double-crossed the Powhatan. Instead of
killing the Englishmen, he entered into an alliance with them,
permitting them to build a fortified trading post on his lands.
All the evidence would indicate that Japazaws was a very wily
politician. He liked English kettles and knives as much as some of
his red neighbors disliked him, and he never did get around to
knuckling under to Opechancanough. No doubt the fort was a smart
move on his part. In any event, because of Japazaws’ need for white
friends with guns Captains Spelman and Croshaw escaped death.
Henry Spelman was not so fortunate the following year however.
On a trading voyage farther up the Potomac River, at the later site of
Washington, D. C., he was killed by the Anacostan Indians after
being captured in an ambush along with Henry Fleet and some other
traders and servants. Just a few days before his death Spelman had
betrayed a warrior who hazarded warning him of his own intended
murder. It appears he even witnessed the agony of the red man’s
tortured execution. Then, by some sort of final justice, he died at the
hands of the same savages to whom he had given over the kindly
warrior.
Aqua vitae may have had something to do with this murderous
episode, for, like the French and the Dutch before them and like the
English who were to follow in New England, some traders in Virginia
taught the savages to drink strong waters as a means of extracting a
richer profit during barter. And what a frightful chance they took!
Emotionally volatile and capricious by nature, the red man was
even more unpredictable of course when he was under the influence
of liquor. He might lose his trading sense to the advantage of the
white men who purposely plied him with aqua vitae during the
feasting that always preceded barter, but when he realized he had
been cheated he was apt to reach for another jug and his tomahawk
at the same time. No doubt many a trader suffered a slow and
bestial death while looking into the bloodshot eyes of savages galled
by stomach fires that he himself had kindled.
Henry Fleet, who had been captured with Spelman, managed the
predicament in which he found himself with more skill than his friend,
indeed with some foresight. While no one in the colony heard from
him in over four years, and all thought he was dead, Fleet not only
contrived to escape execution by the savages, but spent his captivity
to real advantage. By the time he was finally released he had
learned enough about the Algonquin language and had become
sufficiently fraternal with his captors to engage in a profitable trade in
skins with them. Then, returning to London, where he had excellent
family connections, he acquired financial backing from William
Cloberry and other well-known merchants there. Afterward, as
Captain Henry Fleet, he traded in the Chesapeake for some years in
his own ships and later played a significant role in the genesis of the
war between the Virginians and Lord Baltimore over the fur trade.