Robert Morris Stern, William J. Ray, Karen S. Quigley - Psychophysiological Recording-Oxford University Press (2001)

PSYCHOPHYSIOLOGICAL RECORDING
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PSYCHOPHYSIOLOGICAL
RECORDING
Second Edition
Robert M. Stern
William J. Ray
Karen S. Quigley
OXFORD
UNIVERSITY PRESS
2001
OXFORD
UNIVERSITY PRESS
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Copyright © 2001 by Oxford University Press, Inc.

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Library of Congress Cataloging-in-Publication Data

Stern, Robert Morris, 1937-
Psychophysiological recording / Robert M. Stern, William J. Ray, Karen S. Quigley.—2nd ed.
p. cm.
ISBN 0-19-511358-6; ISBN 0-19-511359-4 (pbk)
1. Psychophysiology—Research—Methodology I.Ray, William J., 1945- II. Quigley, Karen S.
QP360 .S79 2000
612.8'028'7—dc21 99-049560
9 8 7 6 5 4 3 2 1
Printed in the United States of America
on acid-free paper
Dedicated to
R. C. Davis
(1902-1961)
Teacher, Scholar, and Father
Preface
This second edition of our book was written 20 years after the first. There
have been great technological advances made during this period, ad-
vances that now make possible the recording of additional physiological
measures from brain and periphery under a variety of conditions not
thought possible when we wrote the first edition. Consider the fact that
functional magnetic resonance imaging (fMRI) is now being used in sev-
eral laboratories to monitor changes in blood flow in the brain, and on
a recent NASA mission the electrogastrogram (EGG) was used to measure
gastric myoelectric activity from a freely moving astronaut in zero grav-
ity.
We have seen a considerable increase in interest in psychophysiology
during the past 20 years, partly due to the availability of new recording
techniques, and, we believe, partly due to the birth and enormous growth
in the area of cognitive neuroscience. Just as psychophysiology (even
before it was called that) contributed both recording techniques and new
hypotheses to the study of emotions, it is now making equally valuable
contributions to the study of cognition.
This second edition of our book, like the first, was written for those
who wish to begin studying psychophysiology. We assumed no previous
knowledge of psychophysiology or of related areas such as physiology or
instrumentation. Therefore, chapters dealing with these areas are pre-
sented first. We aim to provide the reader with a good foundation to
begin working in, as well as understanding, psychophysiology.
Every chapter has been updated, and a chapter dealing with signal
processing has been added. The book is organized into three parts. Part
I deals with background material that we feel is essential to an under-
standing of psychophysiology. It concludes with a chapter on safety and
ethics. Part II presents separate chapters on the psychophysiology of the
brain, muscles, eyes, respiratory system, gastrointestinal system, cardio-
vascular system, skin, and signal processing. Part III deals with appli-
cations of psychophysiological recording to research and clinical use.
In addition to serving as the primary text for a course in psychophys-
iology, this book can function as a supplementary text for courses in
biological psychology, neuropsychology, behavioral neuroscience, cog-
nitive neuroscience, human physiology, medical and nursing courses that
deal with the recording of physiological measures, and beginning courses
in biomedical engineering.
The authors thank our students and others who have commented on
the first edition of the book. We also thank the following individuals who
have helped with the preparation of this second edition. Kari Logel was
a super assistant to the authors. Peter J. Gianaros contributed material
to chapter 5, and he read and offered constructive comments on the
entire manuscript. Kenneth R. Jones, at the University of North Carolina,
also read the entire manuscript and made many valuable suggestions.
We are also grateful to Richard G. Lyons of Sunny Vale, California, who
contributed his signal processing expertise to our new chapter 14. It is
also our pleasure to acknowledge the help, encouragement, and patience
of our editor at Oxford University Press, Joan Bossert, and her assistant
Constanza Morales-Mair.
VIM PREFACE
Contents
PART I. G E N E R A L E L E M E N T S OF P S Y C H O P H Y S I O L O G Y
1. Psychophysiology 3
Short History and Long Past 5
2. Neurons and Muscles: The Sources of

Psychophysiological Recordings 12
Organization of the Nervous System 14
Function of Nerve and Muscle Cells 21
Bioelectric Potentials 33
3. Equipment Used in Psychophysiological Recording 36

Electrodes and Transducers 36
Polygraphs 40
Computers 44
4. Psychophysiological Recordings 47
Spontaneous Responses 48
Tonic Activity 49
Phasic Activity 50
5. Some Basic Principles of Psychophysiology 52

Arousal and Habituation 52
Orienting, Defensive, and Startle Responses 56
Homeostasis and Autonomic Balance 59
Law of Initial Values 62
Stimulus-Response Specificity and Individual
Response Stereotypy 65
6. Safety and Ethics in a Psychophysiology Laboratory 70
Safetyy 70
Additional Safety Principles 71
Ethical Considerations 72
PART II. P S Y C H O P H Y S I O L O G Y OF S P E C I F I C O R G A N S
AND SYSTEMS
7. Brain: Electroencephalography and Imaging 79

Spontaneous EEGs 79
Event-Related Potentials 91
Brain Imaging Techniques 100
8. Muscles: Electromyography 106

Physiological Basis 108
Recording Procedure 109
Typical Recordings 114
Common Problems 115
Analysis and Quantification 121
9. Eyes: Pupillography and Electrooculography 125

Pupillography 125
Eye Movements 130
Electrooculography 135
10. Respiratory System 142

Recording Procedures 147
Typical Recording 152
11. Gastrointestinal Motility: Electrogastrography 157

Recording Procedure 164
Typical Recordings 166
Common Problems 170
12. Cardiovascular System: Heart Rate; Cardiac Output;

and Blood Pressure, Volume, and Flow 178
Heart Rate or Heart Period 186
Cardiac Output 192
X CONTENTS
Blood Pressure 194
Blood Volume and Flow 199
13. Skin: Electrodermal Activity 206

Terminology 207
Skin Conductance 211
Skin Potential 217
14. Signal Processing 221

Assessing Basal Activity 226
Assessing Change 227
Assessing Global Aspects of Physiological Signals 230
PART III. APPLICATIONS
15. Applications of Psychophysiological Recording 245

Five Categories of Psychophysiological Studies 246
Conclusions 255
Glossary 263
Index 275
CONTENTS XI
PART I
General Elements of
Psychophysiology
1
Psychophysiology
Psychophysiology is relatively new as a separate discipline; in the mid-

1950s a group of physiological psychologists began referring to them-
selves as psychophysiologists. However, the subject matter of psycho-
physiology—the interaction of mind and body—has been studied for
centuries by people trained as philosophers, physicists, physicians, phys-
iologists, and, most recently, psychologists.
John Stern (1964) defined the work of psychophysiology as "any re-
search in which the dependent variable (the subject's response) is a phys-
iological measure and the independent variable (the factor manipulated
by the experimenter) a behavioral one." If subjects are shown slides, some
of landscapes and some of car accident scenes, and the subjects' heart
rates are recorded, we have an example of a psychophysiological exper-
iment according to Stern's definition. The dependent variable is heart rate
and the independent variable is type of slide (landscape or car accident
scene). This study would exemplify the typical psychophysiological ex-
periment in which something was done to the subject and the subject's
physiological responses were recorded. Rather than viewing slides, the
subject might have been solving problems, experiencing an embarrassing
situation, waiting to receive an electric shock, or watching a radar screen
for signs of enemy planes. Rather than heart rate, the physiological re-
sponse recorded might have been sweating; a change in blood pressure,
muscle potentials, or cortisol levels in saliva; change in the size of the
pupil of the eye; alterations in brain waves, respiration, stomach motility,
or penis size; or any of several other bodily changes.
Stern's definition of psychophysiology is not incorrect, but with the
passage of time it has become too limiting. The type of research he was
defining, as just described, examined the physiological changes that ac-
companied certain psychological or behavioral manipulations. More re-
cent experiments conducted by psychophysiologists show that it is
equally tenable to manipulate physiological variables and examine be-
havioral changes. In a study typical of newer research, the heart rate of
3
subjects was modified by biofeedbackk and their ability to withstand pain
measured. (A glossary is provided at the back of the book that includes
definitions of technical terms. These terms are italicized the first time they
appear in the text.) The dependent variable in this case is a behavioral
one: the indication of how much pain the subject can tolerate. The in-
dependent variable is a physiological one: the subject's heart rate.
Psychophysiologists are not the only group of behavioral scientists
who study the relationship of physiological and psychological variables.
Psychophysiologists are a subset of a larger group of behavioral scientists
who were referred to as physiological psychologists until recently, and
are now referred to as biological psychologists, psychobiologists, or be-
havioral neuroscientists. What are some of the differences in the ap-
proaches used by psychophysiologists and these other types of biological
psychologists? Other biological psychologists usually study the effects of
their manipulation of the brain or other parts of the nervous system on
some aspect of behavior. The independent variable might be destruction
of a part of the brain, while the dependent variable might be eating
behavior. Such research must be conducted on nonhuman animals and
only rarely on human beings. Most psychophysiologists, on the other
hand, study the responses of humans rather than nonhuman animals;
therefore, such researchers must limit their techniques of data collection
to the surface recording of bioelectric signals. Harmless electrodes are
attached to the skin over the organ of interest. The techniques of psy-
chophysiology have the advantage of not greatly interfering with normal
behavior, particularly when some of the newer methods (such as am-
bulatory assessment) are used so that the subjects can move freely (Fah-
renberg & Myrtek, 1996). Conversely, because surface recording is used,
for example, from the scalp (rather than from deep in the brain), psy-
chophysiologists must sacrifice some degree of immediate biological ex-
actitude, that is, they cannot gain access to the precise source of the
bioelectric signal. A biological psychologist might drill a hole through the
bony skull of the subject, perhaps a cat, and place a very small electrode
on a single cell in a precise part of the brain. This researcher could then
record the electrical signal from this cell while studying, for example,
pleasure. The psychophysiologist placing electrodes on the surface of the
scalp must record the activity of perhaps millions of cells and cannot say
much about the nature of the cells, their number, their location, and so
on. But the psychophysiologist usually records from human subjects. So
when studying pleasure, such a researcher can simply ask subjects to
describe how they feel, rather than needing to make assumptions about
their feeling state based on observable behavior, as biological psycholo-
gists must do. And if the biological psychologist discovers some relation-
ship between the subjects' brain activity and their behavior, this type of
researcher still must question whether that same relationship would be
found with human subjects. Neither approach, that of psychophysiolo-
4 G E N E R A L E L E M E N T S OF P S Y C H O P H Y S I O L O G Y
gists or that of other biological psychologists, is better than the other.
They have both contributed and will continue to contribute to our overall
understanding of the relationship of physiological and psychological var-
iables.
Short History and Long Past
The history of psychophysiology as a separate discipline is quite brief. The

formal development of psychophysiology began in the 1950s, when a
group composed mainly of psychologists met informally under the lead-
ership of R. C. Davis. In 1960 this group organized the Society for Psy-
chophysiological Research, with Chester Darrow as the first president.
Research communications among this group were initiated in 1955,
when Albert Ax began a newsletter dealing with research and instru-
mentation in psychophysiology. In 1964 this newsletter developed into
the journal Psychophysiology, with Ax as the editor; this journal became
the society's official publication. Two articles in the first issue of Psycho-
physiology are of historical interest: "Psychophysiology, Yesterday, Today,
and Tomorrow" by Darrow (1964), and "Goals and Methods of Psycho-
physiology" by Ax (1964). The table of contents of this first issue of
Psychophysiology reveals that in five of the eight original research articles
the galvanic skin response (or electrodermal activity, as we would say
today—see chapter 13) was the measure of interest, and there were no
articles dealing with brain activity. By contrast, in a recent issue of Psy-
chophysiology (November 1999), out of a total of thirteen articles, nine
dealt with brain activity, and there were no articles in which electro-
dermal activity was measured. This change in the psychophysiological
measures of interest to most (but not all) researchers is a reflection of a
great increase in interest in cognitive functioning, plus improvements in
the equipment used to record brain activity and the availability of com-
puters and appropriate software to analyze the signals (see chapter 7).
In the remainder of this chapter, we will examine the past of psycho-
physiology, including the understanding of the electrical properties of the
skin, and conclude with a brief discussion of the development of instru-
mentation.
The Past of Psychophysiology

The early Greeks were interested in the location within the body of in-
tellectual, emotional, and instinctual functioning. The philosopher Plato
suggested that humans possessed a tripartite organization. He believed
that rational faculties were located in the head. Passions were said to be
located in the spinal marrow, which related them to the heart. The in-
stincts—or lower appetites, as they were sometimes called—were said to
PSYCHOPHYSIOLOGY 5
be located in the spinal cord below the diaphragm, where they could
influence the liver. There were parallels to Plato's system outside the
Mediterranean area. In other parts of the world, the body was also
thought of as being organized into different functional entities (e.g., chak-
ras in the metaphysics of India) that performed differential psychological
and physiological functions with respect to the development of the indi-
vidual and the utilization of energy. In all of this philosophical investi-
gation, there was little of what we today refer to as experimentation.
Because of his belief that our senses deceive us, Plato rejected the idea of
experimentation and placed pure thought above empirical observation as
the means to achieve knowledge. In a vein similar to Plato's, Chinese
science and medicine rejected dissection as a method that would lead to
meaningful answers, relying instead on more holistic concepts of human
physiology and functioning.
Yet there is evidence of a sort of psychophysiology before the Renais-
sance. Mesulam and Perry (1972) have shown, through a reexamination
of texts from the third century B.C. to the eleventh century A.D., that
there was considerable empirical sophistication in the writings of Eras-
istratos, Galen, and Ibn Sina.
Erasistratos, a physician during the time of Alexander, is credited with
the following example of clinical psychophysiological observation. A gen-
eral of the time married. The general's son by a previous marriage fell in
love with the same woman (his stepmother) but, realizing that his love
could not be brought out into the open, he resolved not to show his
feelings. The boy then became ill and almost died. After a number of
other doctors had failed to help the boy, Erasistratos worked with him
and decided that the physical problems must be related to a problem of
the mind. This conclusion, though accurate, was not surprising because
contemporary medicine held that the mind and body affected each other.
What is interesting from our standpoint is the method Erasistratos used
to determine the source of the boy's problem. The technique illustrates
an early study in lie detection. Erasistratos observed the reactions of the
boy as various people came to his room. In a later account of this episode,
Plutarch reported that certain signs—"stammering speech, sudden
sweats, irregular palpitations of the heart"—were all present in the boy
whenever the stepmother came to see him. Thus, Erasistratos realized
and correctly diagnosed the problem as being due to the relationship
between the boy and his stepmother. Mesulam and Perry remind us that
Erasistratos was actually an early psychophysiologist developing a theory
of stimulus-response specificity, a concept which will be discussed in
chapter 5.
Galen, a second-century physician who is often thought of as a father
of modern physiology, reported a similar case of lovesickness he diag-
nosed based on the subject's irregular pulse when she heard the name
of her lover. A tenth-century example of these same psychophysiological
6 G E N E R A L ELEMENTS OF P S Y C H O P H Y S I O L O G Y
principles is found in the work of Ibn Sina (Avicenna), who is sometimes
referred to as the Persian Galen. Again, Ibn Sina utilized the method of
elevated pulse rate to determine the person with whom one was in love.
Understanding the Electrical Properties

of the Skin
It was not until the end of the eighteenth century and the experiments
of Luigi Galvani in Italy that the stage was set for the further development
of psychophysiology (Hoff, 1936). Galvani's contribution was the dem-
onstration that animals produce electricity that originates within the or-
ganism itself. Before this time, it was known that muscles of a frog, for
example, would contract when connected to an electrical source, but it
was not known that the muscles were capable of producing an electrical
impulse of their own.
Galvani's research, together with the demonstration of the effect of
applying electricity to paralyzed muscles, led to much speculation con-
cerning how electricity could improve people's health. One theory sug-
gested that diseases could be diagnosed by measuring changes in the
distribution of electrical current in the body. (It is interesting to note that
a similar theory is presently offered as the basis of acupuncture.) A second
theory stated that there was a connection between electricity, animal
magnetism, suggestibility, and hysteria. In particular, it was thought that
through the utilization of a magnet, a hysterical symptom such as func-
tional paralysis of an arm could be transferred to the opposite side of the
body and the originally affected side thus restored to normal functioning.
This was referred to as "transfert" by Charcot. It was through experi-
mentation with this phenomenon that Vigouroux first observed skin re-
sistance level changes. (For an up-to-date discussion of skin resistance
and other aspects of electrodermal activity, see chapter 13.) Vigouroux
measured skin resistance while the hysterical symptom was transferred
from side to side. The hysterical symptom observed was a loss of sensi-
tivity in part of the body, classically referred to as a conversion reaction
or hysterical anesthesia. When the anesthesia was transferred from one
side of the body to the other, Vigouroux noted that skin resistance levels
taken from the insensitive side of the body were higher than those of the
normally functioning side. After a number of these alternations, the skin
resistance of each side of the body remained similar. Thus Vigouroux
may have provided us with the first documentation of the habituation of
skin resistance, that is, the diminution of a response to repeated stimu-
lation (see chapter 5).
Fere, another early worker in this field, was also interested in Char-
cot's theory of hysteria. Before turning to the electrical properties of the
skin, Fere performed research that utilized a hand dynamometer. In these
studies he noted the pressure on the dynamometer as hysterics were
PSYCHOPHYSIOLOGY 7
presented with either sensory stimulation or material of an emotionally
arousing nature. His apparent goal in these studies was to obtain some
measure of the excitation of the nervous system. To this end, he later
undertook another set of studies in which a current was applied to the
anterior surface of the forearm of subjects as they were presented with
emotional and sensory stimulation. In these studies, Fere measured the
change in current flow as a function of the stimuli. Thus Fere reported
in 1888 the first study of skin resistance responses. In addition, work
was also being performed on skin conductance by Hermann, Tarchanoff,
Sticker, Sommer, and others. The interested reader may consult Neu-
mann and Blanton (1970) for a brief history and an excellent bibliog-
raphy. At this time, we will turn to the experiments of Mueller, Veraguth,
and Jung, which helped bring international attention to the study of skin
resistance. As is often the case in science, similar or identical discoveries
are made throughout the world by scientists working independently of
one another. This was the case with Mueller, a Swiss engineer, who
observed that changes in skin resistance appeared to correlate with
changes in psychological state. Mueller consulted with Veraguth, a neu-
rologist, and each independently wrote a series of papers on the subject,
apparently without any knowledge of the earlier work in the area. Ver-
aguth believed that he had found a new reflex, sensitive to emotional
factors, that would be important in dealing with psychiatric problems.
It was in this connection that Veraguth influenced Jung, who com-
bined the measure of skin resistance with a word-association procedure.
Jung developed a procedure in which 100 words were said to an indi-
vidual with the instruction, "Answer as quickly as possible with the first
word that occurs to you." Jung timed the responses and then repeated
the list (Jung, 1910). From this procedure, Jung sought to identify areas
of the person's life which were emotionally important. In one series of
experiments (Jung, 1907; Peterson and Jung, 1907; Ricksher and Jung,
1908), Jung and his colleagues studied the skin resistance responses of
normal and abnormal populations. He also examined changes in respi-
ration that were concomitant with the skin resistance response. In the
last study just cited, Jung concluded that skin resistance responses were
related to attention to the stimulus and the ability to associate it with
previous occurrences, either conscious or unconscious. He also stated
that physical stimuli elicit a greater response than psychological ones
and that the reaction is greater in normal populations than in patholog-
ical ones.
Instrumentation
The first instrument capable of reproducing a continuous record of a
rapidly changing bioelectrical event was the capillary electrometer de-
veloped in the 1870s by Marey (Geddes and Baker, 1968). This instru-
ment consisted of a tube filled with sulfuric acid and mercury. The elec-
trical activity would change the shape of the mercury meniscus, and
through the use of high-intensity light the variations in the contour of
the meniscus would form the basis for recording. This was the first in-
strument to record and display the electrical activity of a frog's heart in
the 1880s; the recording was made by Sir John Burdon-Sanderson. In
1887, Waller first recorded the electrical activity from the human heart
using electrodes on the skin.
Although the capillary electrometer stimulated research in the re-
cording of bioelectrical events, there were problems with the device.
These problems prompted Einthoven—who is known as the father of
electrocardiography—to develop a better device for recording electrical
activity, the string galvanometer. This instrument proved to be reliable,
and even though it was developed in the 1900s it was not fully replaced
until the 1940s. A string galvanometer can be seen in the National Mu-
seum of American History at the Smithsonian Institution in Washington,
D.C., along with some later devices for measuring the electrical activity
of the heart.
In the 1920s additional physiological responses from human beings
were recorded, and research and clinical interest soon followed. In 1929
Berger reported the first human electroencephalogram (EEC), which mea-
sured electrical activity from the brain. Berger not only named the EEC,
he was also the first to report the alpha rhythm and beta rhythm. (See
chapter 7 for more about brain wave activity.) In his early recording, he
pushed platinum wires into the scalp. Later he placed plate electrodes on
the front and back of the head and utilized the Einthoven string galva-
nometer. For a time, he also tried placing a silver spoon in the subject's
mouth as the reference electrode, but later he abandoned the idea. In his
work, Berger demonstrated EEC changes related to eye opening, large-
scale stimuli, and mental activity and attention. He also observed the
EEC in brain-damaged individuals. Within ten to fifteen years, the EEC
had become a clinical tool utilizing multichannel ink writing instruments.
With the introduction of the vacuum tube and then the transistor, psy-
chophysiological recording was soon to become readily available as both
a clinical and research tool.
The field of psychophysiological instrumentation has grown at a tre-
mendous rate, especially since the introduction of the integrated circuit
and the personal computer (PC). In many laboratories, the ink-writing
polygraph has been replaced by a PC equipped with an analog to digital
(A/D) board and data acquisition and analysis software. (See chapter 3
for more about instrumentation.)
This revolution in instrumentation has brought with it a change in
the skills needed by the psychophysiologist. At one time, every psycho-
physiologist had to construct electronic circuits for particular recording
needs, and even the electrodes had to be constructed by hand for each
PSYCHOPHYSIOLOGY 9
application. Today commercially available equipment is excellent and ex-
tremely reliable. With less emphasis on required electronic skills, the psy-
chophysiologist has more time to devote to theoretical and empirical
work related to the interaction of psychological and physiological factors.
References to the specific material covered in the following chapters are

provided at the end of each chapter, but some more general sources of
information about psychophysiology are also available. Andreassi (1995),
in the third edition of his text, "Psychophysiology: Human Behavior and
the Physiological Response," provides many excellent examples of appli-
cations of the psychophysiological recording techniques covered in this
book. Hugdahl (1995) reviews numerous studies in the area of cognitive
psychophysiology in his book, Psychophysiology: The Mind-Body Perspec-
tive. A more advanced treatment of the concepts and techniques used in
psychophysiological recording can be found in Cacioppo and Tassinary
(1990), Principles of Psychophysiology: Physical, Social, and Inferential Ele-
ments. A new Handbook of Psychophysiology (Cacioppo, Tassinary, &
Berntson, 2000) is available. The URL of the web-page for the Society
for Psychophysiological Research is http://www.sprweb.org; the site has
a variety of information about psychophysiology and about the society,
including information about the annual meeting and the society's jour-
nal, Psychophysiology. Other journals devoted to psychophysiological re-
search include the International Journal of Psychophysiologyy and the Journal
of Psychophysiology.
References
Andreassi, J. L. (1995). Psychophysiology: Human behavior and physiological re-
sponse (3rd ed.). Hillsdale, NJ: Erlbaum.
Ax, A. F. (1964). Goals and methods of psychophysiology. Psychophysiology,
1, 8-25.
Cacioppo, J. T., & Tassinary, L. G. (1990). Principles of psychophysiology: Phys-
ical, social, and inferential elements. Cambridge: Cambridge University
Press.
Cacioppo, J. T., Tassinary, L. G., & Berntson, G. G. (2000). Handbook of psy-
chophysiology. Cambridge: Cambridge University Press.
Darrow, C. W. (1964). Psychophysiology, yesterday, today, and tomorrow.
Psychophysiology, 1, 4-7.
Fahrenberg, J., & Myrtek, M. (Eds.) (1996). Ambulatory assessment. Computer-
assisted psychological and psychophysiological methods in monitoring and field
studies. Seattle: Hogrefe & Huber.
Hugdahl, K. (1995). Psychophysiology: The mind-body perspective. Cambridge,
MA: Harvard University Press.
Geddes, L. A., & Baker, L. E. (1968). Principles of applied biomedical instrumen-
tation. New York: Wiley.
Hoff, H. (1936). Galvani and the pregalvanian electrophysiologists. Annals of
Science, 1, 147-172.
Jung, C. G. (1907). On the psychological relations of the association experi-
ment. Journal of Abnormal Psychology, 7, 247-255.
Jung, C. G. (1910). The association method. American Journal of Psychology,
21, 219-269.
Mesulam, M., & Perry, J. (1972). The diagnosis oflove-sickness: Experimental
psychophysiology without the polygraph. Psychophysiology, 9, 546-551.
Neumann, E., & Blanton, R. (1970). The early history of electrodermal re-
search. Psychophysiology, 6, 453-475.
Peterson, F., & Jung, C. G. (1907). Psychophysical investigations with the
galvanometer and insane individuals. Brain, 30, 143-182.
Ricksher, C., & Jung, C. G. (1908). Further investigations on the galvanic
phenomenon and respiration in normal and insane individuals. Journal
of Abnormal Psychology, 2, 189-217.
Stern, J. A. (1964). Towards a definition of psychophysiology. Psychophysi-
ology, 1, 90-91.
PSYCHOPHYSIOLOGY 11
2
Neurons and Muscles
The Sources of Psychophysiological Recordings
The bodily responses that are the subject of psychophysiological study

originate as electrochemical changes in neurons (nerve cells), muscles,
and gland cells. These signals spread from their sources through the body
to the skin surface, appearing to a recording electrode on the body surface
in a somewhat altered form. Understanding the genesis of bioelectric po-
tentials will help in interpreting the surface potentials and serve as a
reminder that whatever their relation to behavior, psychophysiological
responses reflect the functioning of neurons, muscles, and glands.
The nervous system controls the bodily functions measured by psy-
chophysiologists, including muscle action, organ function, and glandular
activity. For example, the coordinated contraction of thousands of skeletal
muscle cells moves us through our environment; enables us to react to
its changes; and is the mechanism of singing, smiling, sitting, and eating.
Cardiac muscle cells pump blood to the lungs and through the body;
smooth muscle cells move food through the digestive system from one
end to the other and generate the uterine forces which move the fetus
from gestation to birth. All these activities require coordinated action of
muscle cells. They must be coordinated with each other, both within and
between muscles, as well as with other organ systems and environmental
events. The random twitchings of thousands of cells could produce nei-
ther effective action nor sustained life. A rapid communication system is
provided by the nervous system, a group of cells specialized for the trans-
mission of information (see figure 2.1). Neurons with their branching
processes pervade the body, interconnecting sense organs with muscle
and gland systems. Events occurring at one place in the body are quickly
and reliably reported to structures elsewhere. This conveying of infor-
mation throughout the body is elaborate in complex organisms, where
billions of nerve cells ensure that the simplest muscular reflex is coor-
12
Figure 2.1. The nervous system. Redrawn with permission from A. W. Ham,
1974, Histology (7th ed.). Philadelphia: Lippincott.
dinated with posture, other reflexes, organ states, events in the environ-
ment, and the experience of the organism.
Organization of the Nervous System
There are at least 100 billion nerve cells in the human nervous system
(Guyton, 1987); some knowledge of their organization is useful in un-
derstanding their function. Neurons are usually found in bundles that
reflect their origin and their destination. These bundles, called nerves in
the peripheral nervous system (PNS) and tracts within the central nervous
system (CNS), may include only a few neurons or millions of them. The
PNS includes all neurons outside the bony enclosures of the spinal col-
umn and skull, and the CNS is composed of all those cells inside it. Note
that this is a distinction of convenience because the PNS and CNS are
not really separate systems. Indeed, some neurons are contained partly
in the CNS and partly in the PNS.
Neurons are somewhat oddly shaped cells of the body. A schematized
neuron, is depicted in figure 2.2. The receiving end of a neuron is
equipped with dendrites that accept chemical messages from other neu-
rons or, in the case of sensory neurons that generate a signal within the
neuron, when a sensory stimulus is received (e.g., a dendrite transmits
information about touch when pressure deforms it). Dendrites extend
from the neuron's cell body, which contains the genetic material and the
energy-generating machinery of the cell. Extending away from the cell
body is a long process called the axon; the axon ends in axon terminals.
Signals internal to a neuron typically pass from the dendrites to the cell
body to the axon and finally to the axon terminals. At the axon terminals,
chemical substances called neurotransmitters are released into the extra-
cellular fluid. Neurotransmitters diffuse across the extracellular space be-
Figure 2.2. Major components of a neuron. The double slash marks through
the axon indicate that the full length of the axon has not been drawn.
tween two neurons and are received at receptors on the dendrites of a
nearby neuron. If the neurotransmitter finds a matching receptor, then
an electrical change is initiated in the neuron receiving the chemical
message. We will discuss the function of these neuronal components in
more detail later.
Neurons can be functionally distinguished by the direction in which
they conduct impulses. Neurons conducting impulses toward a particular
structure are afferent neurons (often the point of reference is the CNS);
those conducting impulses away from a particular structure are efferent
neurons. Functionally independent afferent and efferent neurons, such
as those conducting sensory information to the CNS, and motor infor-
mation away from the CNS can be, and often are, found in the same
nerve bundle.
Sensory Systems
Sensory neurons, which translate sensory signals such as light or pres-
sure into neural signals send information from sensory receptors to the
CNS. Sensory neurons usually do not synapse before entering the CNS;
this means that a single neuron transmits information from a sensory
receptor in a muscle, the skin, or an internal organ directly to the spinal
cord or brain; a notable exception occurs in the retina of the eye. Once
information reaches the spinal cord or the brain, other neurons convey
that information to sites both within the CNS and, in some cases, to
motor neurons leading back out of the CNS. The nervous system does
not connect muscles directly to other muscles or glands, nor does it con-
duct sensory information from the environment directly to the muscles.
Rather, the sensory information goes first to the CNS, from which it is
redistributed both inside and outside the CNS. This arrangement provides
the capacity for integrating incoming sensory signals with conditions
elsewhere in the body where the CNS acts as the "executive system" in
charge of coordinating action and function throughout the body.
Motor Systems
Anatomically the motor neurons can be divided into two subsystems, the
somatic motor system and the autonomic nervous system. Although his-
torically the autonomic subdivision of the PNS was considered a motor
system, a considerable number of sensory fibers are present within au-
tonomic nerve bundles (Loewy, 1990). Thus, the autonomic nervous sys-
tem is more correctly considered both a motor and sensory system for
control of, and feedback from, the internal organs and glands. However,
because much more is known about the efferent function of the ANS, we
will only consider here its role as an efferent system for control of organs
and glands.
N E U R O N S AND M U S C L E S 15
Figure 2.3. The segmental arrangement of the spinal cord. Redrawn by per-
mission from E. Gardner, (1975), Fundamentals of neurology (6th ed.), Phila-
delphia: Saunders.
Somatic System. The somatic nervous system is composed of efferent neu-

rons that project out of the CNS and innervate the skeletal musculature.
The dendrites of motor neurons that receive incoming messages are
found within the central horn (i.e., gray matter) of the spinal cord at
each spinal segment (there is one segment for each vertebra; see figure
2.3). The motor neurons in each spinal nerve extend without synapse to
16 G E N E R A L E L E M E N T S OF PSYCHOPHYSIOLOGY
a group of striate (skeletal) muscles at approximately the level of the
vertebral segment from which the neuronal processes emerged. The ax-
ons of each motor neuron run close to one another in the spinal nerve
and then separate from one another near the target muscle, so that a
single motor neuron typically terminates at the motor end plates of sev-
eral muscle cells. Each motor neuron and the group of muscle cells in-
nervated by that motor neuron is called a motor unit—all the muscle cells
will necessarily contract together in response to action potentials origi-
nating in the motor neuron. In the somatic motor system, this route from
the CNS to the motor unit is called the final common path, reflecting the
invariant nature of the system once an action potential has been gen-
erated in the spinal motor neuron.
Considering this arrangement of motor units, one might predict that
where there is fine control of muscular action, the motor units would be
small when compared to those muscles over which we have relatively
crude control. Sure enough, motor units are much smaller in the muscles
controlling eye movements than in the large muscles of the back. This
difference in control is referred to as the size principle. Not only do smaller
motor units control fewer muscle fibers and thus confer finer muscle
control than do large motor units, but smaller motor units also are re-
cruited faster than larger motor units. This faster recruitment of smaller
motor units allows for movements to be initiated gradually and smoothly,
which is particularly important when large muscle masses are activated
(Guyton & Hall, 1996).
Autonomic System. The autonomic nervous system (ANS) consists of two

major branches or divisions, the sympathetic nervous system and the para-
sympathetic nervous system. The two divisions of the ANS are anatomically
quite different than the somatic system. Efferent neuronal fibers from the
somatic system exit the CNS and innervate a muscle without synapse.
Efferent fibers from the ANS, on the other hand, emerge from the CNS
and synapse once outside the CNS before reaching the target organ or
gland. The anatomical structure formed by the synapses between the
neurons exiting the spinal cord and the dendrites and cell bodies of re-
ceiving neurons is called a ganglion (plural is ganglia). Ganglia for the
sympathetic nervous system lie in a chain near the spinal cord and ver-
tebral column. Ganglia for the parasympathetic nervous system typically
lie in the wall of the target organ. Thus, even the anatomy of the two
major autonomic branches is quite dissimilar. We also now recognize a
third division of the ANS: the enteric nervous system, which regulates
much of the function of the gastrointestinal system.
Sympathetic division. The sympathetic division is composed of those
neurons that originate within the thoracic and lumbar segments of the
spinal cord and project to ganglia lying along the vertebral column just
outside the cord (see figure 2.4). Because of its anatomical arrangement,
Figure 2.4. Sympathetic division of the autonomic nervous system. This figure
illustrates the preganglionic axons exiting the spinal cord, the ganglia making
up the sympathetic chain, and the postganglionic fibers innervating target
organs and glands. Figure used with permission from B. Pansky, D. J. Allen,
& G. C. Budd, 1988, Review of neuroscience (2nd ed.), New York: Macmillan.
the sympathetic division has also been called the thoracolumbar system.
There is a ganglion through which efferent fibers of the sympathetic sys-
tem pass for each spinal segment from the first thoracic to the third
lumbar segment. Many of these ganglia are interconnected to form the
chain of ganglia on each side of the vertebral column. Efferent neurons
from the spinal cord (i.e., preganglionic neurons) that enter the sympa-
thetic chain may synapse at the same segmental level where they exit
from the cord, or they may extend up or down the chain to enter other
ganglia, or they may even pass directly through the chain without syn-
apse. The efferent fibers leaving the sympathetic chain (also known as
postganglionic neurons) innervate smooth muscles and glands in the
skin, eyes, mucous membranes, and viscera. Most postganglionic sym-
pathetic fibers extend directly from cell bodies in the chain ganglia to a
target organ. However, postganglionic fibers innervating the smooth
muscles and glands of the abdomen and pelvic area originate in ganglia
lying near the sympathetic chain (e.g., the celiac and inferior mesenteric
ganglia; see figure 2.4). The rule is that there is a preganglionic and a
postganglionic fiber providing autonomic input to a target organ; one
interesting exception is the case of the adrenal medulla. A single pregan-
glionic neuron extends without synapse directly to the adrenal medulla,
where norepinephrine and epinephrine are released into the blood stream
as hormones (Edwards, 1990). This is a somewhat unusual mode of de-
livery for the ANS, where substances are typically released as neurotrans-
mitters onto a postsynaptic cell. However, the secretory cells that release
norepinephrine and epinephrine are developmentally derived from ner-
vous system cells, and function essentially as a specialized type of post-
ganglionic neuron (Guyton & Hall, 1996).
The neurotransmitter released at the ganglia onto the dendrites of the
postganglionic neuron is acetylcholine. Here, acetylcholine activates cho-
linergic receptors of the nicotinic subtype (which, as the name implies,
respond to nicotine). The neurotransmitter released by the terminals of
the sympathetic postganglionic neurons onto target organs and glands is
norepinephrine, with the exception of the eccrine sweat glands, where
acetylcholine is the transmitter (see chapter 13 for a description). Thus,
the norepinephrine released into the bloodstream as a hormone by the
adrenal medulla activates most targets innervated by the sympathetic
division.
Parasympathetic division. The second major division of the ANS is the
craniosacral, or parasympathetic division. The parasympathetic division
is composed of preganglionic neurons whose cell bodies lie in the brain
stem and in the sacral segments of the spinal cord, and synapse onto
postganglionic neurons in or near the target organ (see figure 2.5). Post-
ganglionic neurons in the cranial division innervate the eyes, the mucosa
of the nose and mouth, the salivary glands, the heart, the lungs and
bronchi, and the abdominal organs. Postganglionic neurons in the sacral
division innervate the genitalia and organs of the pelvic cavity, such as
the bladder and bowel. As with the sympathetic system, the neurotrans-
mitter released by the preganglionic fibers onto the postganglionic den-
drites is acetylcholine, which activates cholinergic receptors of the nico-
tinic subtype. As in the somatic system, the neurotransmitter at the
terminals of the postganglionic neurons in the parasympathetic division
is acetylcholine, and here it activates cholinergic receptors of the mus-
carinic subtype.
The parasympathetic division innervates many of the same target or-
gans and glands as the sympathetic system, although the overlap is not
complete. When both divisions of the autonomic nervous system inner-
vate the same target, we say that the target is dually innervated. This is
a common pattern in autonomic anatomy. Often, activation of the two
Figure 2.5. Parasympathetic division of the autonomic nervous system. This
figure illustrates the preganglionic axons exiting the brain and spinal cord
and entering ganglia. Postganglionic axons leaving ganglia and entering a
target organ or gland are also shown. Where ganglia are located very near
or within the target organ, the postganglionic cell bodies and axons are not
shown. Figure used with permission from B. Pansky, D. J. Allen & G. C. Budd,
1988, Review of neuroscience (2nd ed.), New York: Macmillan.
branches exerts differential effects on the target organ. For example, the
heart receives input from both the sympathetic and parasympathetic
branches of the ANS. Increasing sympathetic activation of the heart leads
to an increased rate of beating of the heart. Conversely, reducing activity
in the sympathetic branch innervating the heart causes the heart beat
to slow. At the heart, the parasympathetic system acts in a manner op-
posite to the sympathetic. Thus, increasing parasympathetic activation of
the heart decreases the heart rate and decreasing parasympathetic acti-
vation increases the heart rate. It should be kept in mind that both
branches of the ANS can be tonically active (meaning that there is some
ongoing activity in the nerves) even when the body is at rest. This permits
both increases and decreases in activity in either branch to make changes
in the functional state of a given target organ. It used to be assumed that
when activity in one branch increased, that activity in the other auto-
nomic branch necessarily decreased (referred to as a reciprocal mode of
ANS control). This is not always true, however. Sometimes activity in
one autonomic branch increases or decreases, with no change in the
activity of the other branch (uncoupled mode of ANS control), or activity
in both branches can simultaneously increase or decrease (coactivational
mode of ANS control). These three modes of ANS control—reciprocal,
uncoupled, and coactivational—can occur for any target organ receiving
innervation from both autonomic branches. These multiple modes permit
subtle changes in the function of the target organ that are not possible
with only reciprocal activation (see chapter 12 and Berntson, Cacioppo,
& Quigley, 1991, for more discussion of this issue). Table 2.1 depicts the
usual effects of the activation of the sympathetic and parasympathetic
divisions on various organs.
Function of Nerve and Muscle Cells
Nerve cells and muscle cells are alike in two important ways; they are
elongated and they are excitable. Elongation refers to the long, narrow
shape of nerve and muscle cells. Cells are called excitable when a stimulus
that occurs at one point on the cell causes an electrical disturbance that
spreads over much or all of the cell. These two properties combine to
create the distribution of local events over large areas and great distances.
Indeed, neural excitation may be propagated for a meter or more in some
nerve cells.
Excitability: Resting Potential and

Action Potential
When nerve and muscle cells are inactive, an electrical potential can be
observed across the cell membrane. The inside of the cell is approximately
-90 mV compared to the outside (see figure 2.6). This means that unlike
charges have been separated, with positive charges lying along the out-
side of the cell membrane and negative charges along the inside of the
cell membrane. The fact that charges are separated also means that there
is a capacity to perform work. The resting nerve or muscle cell is like a
tiny flashlight battery in which charges are held apart, to be used when
a path between the positive and negative charges is created. The sepa-
ration of charges is termed polarization, and the resulting steady electrical
potential between the inside and outside of excitable cells is called the
resting potential.
The resting potential is due to the resting cell's permeability to potas-
sium (K + ), the cell's relative impermeability to ions such as sodium
Table 2.1. Effects of the Autonomic Nervous System on Selected Target Organs and Glands
Increased Increased
Structure Function Parasympathetic Activity Sympathetic Activity
Eyes: Iris Control of light to the

eye
-radial muscle —a Contraction (pupil dilates)
-sphincter muscle Contraction (pupil con- —a
stricts)
Eyes: Ciliary muscle (at- Accommodation Constriction (near vision) Relaxation (far vision)
tached to the lens)
Nasal glands Secretion Increased secretion —a
Lacrimal glands Tear production Increased secretion —a
Salivary glands Saliva production Increased potassium and Some increase in potassium, water
water secretion (profuse, and amylase secretion (viscous
thin secretion) secretion)
Gastrointestinal Muscle tone and motility Increased motility and tone Some decrease in motility and tone
Pancreas (islets) Insulin secretion Increased release Decreased (cc2-adrenergic) and in-
creased (p2-adrenergic) release
Heart: S-A node Rate control Slows rate Increases rate

Heart: atria and ventri- Contractility control Some decrease in contrac- Increase in contractility
cles tility
Bronchi of lungs Bronchial muscle tone Contraction and narrowing Relaxation and dilation of bronchi
of bronchi
Adrenal medulla Secretion of catechola- —a Secretion of catecholamines
mines (epinephrine &
norepinephrine
Arterioles: skeletal muscle Arterial tone Slight dilation Both constriction and dilation
Arterioles: skin Arterial tone —a Constriction
Sweat glands Sweat production —a Increased sweating
Pilomotor muscles Erection of hair —a Initiation of "goose flesh" or
"goose bumps"
Bladder: muscle wall Muscle tone Contraction Relaxation

Bladder: sphincter Muscle tone Relaxation Contraction
Male genitalia Sexual behavior Erection Emission & ejaculation
Note: The table is organized with organs and glands innervated by the cranial portion of the parasympathetic division at the top, the effectors innervated by the
thoracolumbar (sympathetic) division in the middle, and the effectors innervated by the sacral portion of the parasympathetic division at the bottom of the table.
Thus, the table roughly reflects the anatomical organization of the ANS. The horizontal spaces in the table separate the cranial, thoracolumbar and sacral
divisions. Information in the table was compiled from Berne & Levy (1998), Guyton & Hall (1996), and Loewy & Spyer (1990).
"Indicates that there is no known functional effect of the branch indicated.
Figure 2.6. The resting and action potentials. The figure illustrates the resting
potential of a neuron (approx. -90 mV), and the rapid depolarization and
repolarization that occurs when an action potential is fired. Above the illus-
tration of the potentials is a depiction of the way in which these potentials
are measured, using an electrode that impales the axon and records the elec-
trical potential of the inside of the cell relative to the outside of the cell. Figure
used with permission from A. C. Guyton & J. E. Hall, 1996, Textbook of medical
physiology (9th ed.), Philadelphia: Saunders.
(Na + ), and an energy-requiring Na + -K + pump in the cell's membrane

which helps to maintain the relative negativity of the inner surface of
the cell membrane. The cell membrane is semipermeable, meaning that
some substances can pass through the membrane whereas others cannot.
The major contributor to the resting potential is the Na+-K+pump which
requires metabolic energy and pumps 3 Na + ions out of the cell while
simultaneously bringing 2 K + ions into the cell. As the pump works, there
is a greater accumulation of positive ions on the outside of the cell and
a relative negativity on the inside of the cell (i.e., fewer positive ions).
Together, these factors result in a cell with the potential to do the work
of the nervous system or create muscular activity. For a more compre-
hensive description of the basis of the resting potential, see Guyton and
Hall (1996) or Berne and Levy (1998).
When the resting potential of the cell is reduced slightly at a point on
the membrane (i.e., the inside of the cell becomes more positive relative
to the outside), several local events occur. First, the membrane momen-
tarily becomes more permeable to sodium, so that sodium ions begin to
enter the cell from outside. Local currents are then created as ions move
across the membrane. However, if the current flow is small (only a few
millivolts), the disturbance will remain localized and likely will not spread
to excite the entire cell. However, if a stronger stimulus (i.e., a larger
decrease in membrane potential or depolarization) is applied to the
membrane, this creates a very different effect. If the depolarization
reaches a certain level, termed threshold (in the neighborhood of a 20-
40 mV reduction in transmembrane potential), then the membrane be-
comes very permeable to sodium, which enters the cell quickly. This
results in a rapid change across the membrane such that the inside of
the neuron briefly becomes positively charged. The increase in sodium
permeability, large sodium influx, and quick reduction of negative poten-
tial in the cell occurs explosively and ends only when all of the energy
stored at that point on the membrane has been exhausted. When the cell
membrane becomes briefly positive, then the approximately 90 mV rest-
ing potential has been discharged.
Just after the influx of sodium, sodium inactivation gates close and
the membrane again becomes highly impermeable to sodium. Subse-
quently, the Na + —K + pump begins to force sodium out of the cell in
exchange for potassium ions that enter the cell. This mechanism restores
the resting potential. The explosive chain of events during which the
potential across the membrane is discharged is called the action potential
of excitable cells and is the mechanism of excitation in nerve and muscle
cells. The relatively large local currents produced by initiation of an ac-
tion potential are sufficient to propagate this potential to nearby portions
of the cell by reducing the membrane potential there to threshold value,
and thereby triggering an action potential. This is analogous to the burn-
ing of a fuse on a firecracker, where the burning fuse successively ignites
all along the length of the fuse. Interestingly, the action potential is un-
diminished as it moves over the cell, because energy is stored all along
the length of the cell. As a result, the excitation of a given nerve cell is
uniform each time an action potential is fired; this excitation is indepen-
dent of the size or physical nature of the stimulus, as long as depolari-
zation exceeds the threshold. The fact that the action potential is either
(a) initiated when depolarization reaches threshold or (b) not initiated
when changes in polarity are below threshold is referred to as the all-or-
none principle. This principle implies that if the stimulus is great enough
to cause an action potential, the cell will be completely depolarized, re-
gardless of further increases in the amplitude of the stimulus. On the
other hand, a subthreshold stimulus will produce only local changes that
are not propagated along the length of the cell.
The consequences of action potentials are quite different in muscle
and nerve cells. We will first examine muscle cells and the activity of the
three types of muscles found in the body—striate, smooth, and cardiac—
and then return to our discussion of nerve cells.
Muscles
Groups of Muscle Cells: Muscles. Acting alone, single muscle cells would
be incapable of producing the great tensions required to move the long
bones and to support the skeleton against the force of gravity. In fact,
muscle cells are ordinarily arranged in groups, with like cells having the
same function. These groups, of course, are muscles. Muscles are of two
very different kinds—striate (skeletal) and smooth—with cardiac (or
heart) muscle exhibiting some of the properties of each. Cardiac muscle
is striated like skeletal muscle but operates as a syncytium, or intercon-
nected unit, like most smooth muscle does.
Striate muscle. Striate, or striped muscle is sometimes called skeletal
muscle because it attaches to the bones and is responsible for support and
movement of skeletal body parts. Striate muscle is relatively fast-acting,
although there is a wide variation in contraction speed from the slow
muscles of the back to the fast muscles of the eyelid. Not all striate mus-
cles are attached to bones at both ends. For example, the muscles of facial
expression arise from and attach to connective tissue; thus smiling and
frowning do not involve the skeleton. However, most striate muscles are
arranged around bones, arising from tendons and ligaments on one side
of a joint and inserting into bone on the other side of the joint, so that
muscle shortening produces tension across the joint. If tension is suffi-
cient, there will be movement of bone around the joint. Due to their
anatomical placement, other muscles may oppose this action, so their
action can prevent movement or move the bone in the opposite direction.
At complex joints such as the ankle, hip, wrist, and neck, many muscles
surround the joint, providing support and movement in many directions.
The cells of striate muscle are electrically insulated from each other.
As a result, an action potential spreading over a cell does not affect neigh-
boring cells. This means that both the initiation of contraction and co-
ordinated contractions of groups of muscle cells must come from outside
the muscle by way of the nervous system. Striate muscle deprived of its
nerve supply is completely relaxed unless action potentials are artificially
generated by electrical or chemical stimulation. Paralysis following severe
damage to the spinal cord is an example of intact, otherwise healthy
muscles that are incapable of generating tension because they are re-
ceiving no external input.
The cellular basis of striate or skeletal muscle contraction is relatively
well understood and so we will explore it in some detail. The skeletal
muscle cell, or fiber, is composed of slender myofibrils. Each fibril is made
up of tiny filaments, which are the contracting units of the cell. These
filaments, which lie longitudinally within the fiber, are of two kinds: actin
26 G E N E R A L ELEMENTS OF PSYCHOPHYSIOLOGY
and myosin. In a noncontracted state, adjacent actin and myosin fila-
ments overlap slightly but are not bound to one another. When con-
traction is initiated, bonds between the two types of filaments are quickly
made, the fibers are pulled alongside one another such that the filaments
overlap to a greater extent, and then the bond is broken. This process is
often described as a kind of "ratcheting" movement with repeated se-
quences of bond formation, pulling together of the fibers, and breaking
of the bond as the muscle fibers become overlapped to a greater and
greater extent with each sequence. As the two kinds of filaments slide
alongside one another, the myofibril shortens along its longitudinal axis.
When this occurs in many adjacent myofibrils it causes contraction or
shortening of the entire muscle fiber.
Skeletal muscle contraction is initiated when a neuron sends a mes-
sage to the muscle at the particular neuromuscular junction that is the
point of contact between a nerve cell axon terminal and a muscle cell.
Release of the neurotransmitter acetylcholine onto the muscle at the neu-
romuscular junction causes an action potential to be propagated in the
muscle fibers. The muscle action potential then causes calcium to be
released. The influx of calcium to the filaments begins a chemical reac-
tion, part of which is used to initiate the ratcheting and sliding motion
of the actin and myosin filaments. A thorough account of the contractile
process may be found in Guy ton and Hall (1996) or Berne and Levy
(1998).
The time required for contraction varies greatly in different muscles,
but it is always much longer than the action potential that precedes it.
Even in relatively fast skeletal muscle, the contraction-relaxation se-
quence to a single stimulus may not be complete for 30-200 ms even
though the action potential lasts only 1-5 ms. In addition, the action
potential can be complete before contraction even begins.
Under normal circumstances the action potential is always of the same
magnitude in a given cell, and it might be expected that the contractile
response of the muscle cell would always be the same (i.e., that it would
result in the same shortening or increase in tension). However, this is
not always the case. If the relaxation phase following contraction is com-
pleted, then the tension produced by a succeeding action potential will
indeed be similar. If, however, a second action potential is propagated
before the previous contraction is complete, a further increase in tension
results. The amount of increased tension depends on the interval between
the two stimuli, with shorter intervals producing greater increases in
tension. This effect, called frequency summation, continues as the fre-
quency of stimulation increases, with each succeeding increase in fre-
quency adding a diminishing amount of tension until further increases
add no more tension (figure 2.7). At this point, the muscle cell is in a
state of continuing maximal tension called tetanus. Thus, a single striate
(skeletal) muscle cell is capable of finely graded contractions that depend
Figure 2.7. The process of tetanization. The figure demonstrates how tetanus
develops in striate muscle as the frequency of stimulation of a muscle in-
creases. Note that contraction is smooth and sustained once the rate of stim-
ulation becomes sufficiently high. Figure used with permission from A. C.
Guyton & J. E. Hall, 1996, Textbook of medical physiology (9th ed.), Philadel-
phia. Saunders.
on the rate at which action potentials are generated. Whereas striate

muscle is capable of tetanic contraction, smooth muscle is not. Smooth
muscle, such as that found in the stomach, can also sustain maximal
tension, however, it uses a different mechanism than tetanization.
Smooth muscle. Smooth muscle is found in layers around the hollow
organs of the gastrointestinal tract, around the uterus and bladder, and
surrounding the arteries. Smooth muscle also controls constriction and
dilation of the pupils and the limited action of some body hair (as when
hair "stands on end" or you get "goose bumps"). Smooth muscle con-
tracts slowly. An action potential may produce a complete contraction-
relaxation cycle only after a second or two. There are two types of smooth
muscle: multi-unit and unitary. Multi-unit smooth muscle operates much
like striate muscle cells because each cell is physically separate from its
neighboring cells and activation of single cells generally occurs when
there is input from a neuron. In contrast, action potentials generated in
unitary smooth muscle cells spread to other smooth muscle cells via gap
junctions (low-resistence connections) that permit the spread of electrical
current between cells. This ability to spread activation over a large num-
ber of cells means that a contraction in one part of a smooth muscle
may, in effect, spread over the muscle with minimal or even no neural
input. For example, hormones or mechanical changes (such as stretch)
can produce smooth muscle contraction that is initiated without any
neural input to the muscle.
In addition, the resting potential of some smooth muscle cells (partic-
ularly those of the gastrointestinal tract) fluctuates rhythmically. These
spontaneous changes in the resting potential, called slow waves, can
generate action potentials and hence contractions. Contraction of smooth
muscle can be both initiated and maintained under local control as a
result of spontaneous rhythmic variations and direct electrical conduc-
tion between cells.
Cardiac muscle. Cardiac muscle acts faster than smooth muscle, but
the interconnection of cells is similar such that electrical activity can
spread quickly over many cells. The heart has two regions of functionally
interconnected muscle cells, the atrial syncytium and the ventricular syn-
cytium. Syncytia are composed of cells connected by gap junctions. Thus,
the muscle cells of the atrial syncytium conduct action potentials virtu-
ally simultaneously, as do the cells of the ventricular syncytium. There
is, however, a slight delay between contraction of the atria and contrac-
tion of the ventricles because electrical current passes between these two
syncytia via a specialized bundle of conducting fibers called the atrioven-
tricular (A/V) bundle. This delay is important because it allows time for
blood to move from the atria to the ventricles so that there is sufficient
blood in the ventricles before contraction to move blood into the arterial
system (see chapter 12 for more details).
Neurons
Conduction in a Neuron. In individual nerve cells, an action potential is
generated when a sufficient increase in positive potential occurs at the
dendrites or cell body. At the point where the axon exits from the cell
body is a portion of the axon known as the axon hillock, where changes
in potential coming from all around the cell body and dendrites are
summed and if threshold is reached, an action potential is generated. The
rate at which the action potential is conducted down the axon depends
on two factors: (a) the diameter of the axon, and (b) whether or not the
axon is myelinated. Conduction speed is faster in large diameter axons
than in small diameter ones, the speed being proportional to the square
root of the diameter.
The other condition that greatly increases the speed of the impulse
along the nerve fiber is the presence of a myelin sheath. Many nerve cells
that conduct over long distances are individually encased in a sheath of
fatty material called myelin. This sheath gives nerve tracts their char-
acteristic white appearance. Indeed, areas of myelinated and unmyelin-
ated fibers in the brain are termed white and gray matter, respectively.
Myelin is highly resistive to the passage of current, so that the resting
potential does not develop in myelinated areas. Indeed, if the entire axon
were covered in myelin, no impulse would be able to be conducted be-
cause the sheath would prevent any current spread. However, the myelin
sheath contains tiny gaps where the axon membrane is exposed (these
gaps are only about 2-3 mm long). These gaps, called nodes of Ranvier,
occur about every 1 mm along the axon and permit local current flows
through one of these exposed sections of membrane, which then depo-
larizes the membrane to threshold at the next node thereby generating
an action potential at the subsequent node. Thus, the impulse "jumps"
across adjacent nodes of Ranvier along the length of the neuron. This
action, termed saltatory conduction, greatly speeds conduction, since no
time is spent depolarizing the membrane between the nodes. Indeed, the
impulse moves about 40-80 times faster in a myelinated fiber than in
an unmyelinated fiber of the same diameter. For example, Morell and
Norton (1980) point out that a human spinal cord that contained only
unmyelinated axons would need to be several yards in diameter to con-
duct impulses as quickly as it can in its normal, myelinated state. Con-
duction velocities as fast as 120 m/s (which is greater than the length
of a football field in 1 s) can be observed in large diameter myelinated
axons, whereas small, unmyelinated axons may propagate signals as
slowly as 0.25 m/s (Guyton & Hall, 1996).
When the action potential reaches the end of the neuron, it causes a
momentary release of neurotransmitters stored within the axon termin-
als. This process describes the action of single nerve cells; electrical events
occurring at the receiving end of the cell cause the release of chemicals
from the transmitting end of the cell. The interconnections of numerous
neurons, with each other and with sense organs, muscles, and glands
are the basis for the function of the nervous system.
Communication between Neurons. Until now, we have mostly described

events along the length of the axons. But since we noted earlier that,
once started, the action potential is unchanged as it passes down the
axon, then it must be the processes happening at the synapses between
the neurons that are responsible for the variety of our experience and
the subtlety of our actions.
Receptor or generator potentials. In the sensory neurons, dendrites gen-
erate messages when receptors are activated by sensory stimuli. These
messages can be in the form of stimuli that are mechanical (e.g., pressure
or vibration), thermal (e.g., heat or cold), nociceptive (e.g., painful), elec-
tromagnetic (e.g., light), or chemical (e.g., taste or smell). These stimuli
act on specialized receptors and generate a depolarizing potential or re-
ceptor potential across the receptor membrane. When a receptor potential
is generated, there is a change in electrical potential of the neuron that
is proportional to the amount of stimulation received (such a proportional
response is called graded). An action potential may be initiated when
graded receptor potentials from multiple sensory receptors are combined
with other changes in potential coming in at points all over the cell.
Therefore, sensory signals differ as a result of the degree of stimulation
received at the sensory receptors and due to the number and frequency
of action potentials generated, not because of differences between action
potentials across various sensory systems. Thus, our knowledge of the
world around us and within us is a function of our sensory receptors and
their sensitivity.
The synapse: EPSPs and IPSPs. Most nerve cells terminate in the im-
mediate vicinity of other nerve cells. The region where two neurons meet
and where neurotransmission can take place between them is called a
synapse. Adjacent neurons do not touch one another; instead there is a
small space between them called a synaptic deft. When an impulse arrives
at the axon terminals, it does not "hop" the gap between the presynaptic
and postsynaptic cells. Instead, an action potential in the presynaptic cell
causes the release of a neurotransmitter from the axon terminals of that
cell. The neurotransmitter then diffuses across the synaptic cleft and ar-
rives at the membrane of the postsynaptic cell. The neurotransmitter next
causes depolarization or hyperpolarization of the dendritic membrane of
the postsynaptic cell. The effect of the neurotransmitter is determined by
the chemical composition of the transmitter substance. Moreover, a neu-
rotransmitter can be effective only if it finds a receptor whose structure
matches the structure of the transmitter. If the neurotransmitter partially
depolarizes the postsynaptic cell membrane, then an excitatory postsynap-
tic potential (EPSP) has occurred. It is excitatory because it moves the
transmembrane voltage closer to threshold level, making the cell easier
to excite. It is worth noting that the transmitter released by a single
action potential from a single terminal is not sufficient to produce an
action potential in a postsynaptic cell. The effect of multiple action po-
tentials in many presynaptic cells is required to excite a postsynaptic cell.
Some neurotransmitters make a postsynaptic cell harder to excite. This
is accomplished by hyperpolarizing the membrane, making the inside of
the postsynaptic cell more negative with respect to the outside, so that a
greater subsequent depolarization is required to bring the membrane to
threshold. This hyperpolarizing potential is termed an inhibitory postsy-
naptic potential (IPSP). Both EPSPs and IPSPs are graded potentials.
The postsynaptic cell, then, is bombarded by a continuing flow of neu-
rotransmitters, some of which make the cell more likely to fire (that is,
generate an action potential) and some of which make the cell less likely
to fire. The result of this bombardment is a continually fluctuating trans-
membrane potential. However, when the postsynaptic cell's membrane
potential reaches threshold, an action potential is produced in the cell
and is conducted over the length of the cell in the manner previously
described.
From these synaptic events several important observations can be
made. First, impulses are normally conducted in the nervous system in
only one direction. This occurs because action potentials are propagated
from the axon hillock toward the terminals and because neurotransmit-
ters are released only at axon terminals, not at the dendrites. Second,
although the conducted impulse in a given neuron is "all or none," the
N E U R O N S AND MUSCLES 31
potentials at the synapse are finely graded. Finally, it is at synapses that
the integration of nervous control of behavior takes place. Excitatory and
inhibitory inputs from multiple neurons converge to produce or inhibit
activity in a given postsynaptic cell.
Motor or end plate potentials. Besides stimulating or inhibiting other
neurons, nerve cells may release neurotransmitters onto muscle or gland
cells, and thereby influence their action. In the case of striate muscle
cells, an axon terminal supplies each muscle cell, arriving at a specialized
structure called the motor end plate. There the neurotransmitter acetyl-
choline produces a reduction in the resting potential of the muscle cell
(the end plate potential) that nearly always initiates a muscle action
potential. In some smooth muscle, the transmitter (typically acetylcholine
or norepinephrine) is released at several places along the terminal ends
of the axon and diffuses over the muscle cells (Guyton & Hall, 1996). By
analogous mechanisms, neuronal action potentials can produce glan-
dular secretion.
A simple reflex: The function of receptor, synaptic, and motor potentials.
The three actions of neurons—receptive, synaptic, and motor—are easily
seen in the simple monosynaptic reflex arc shown in figure 2.8. In most
striate muscles, there are receptors that respond to stretching by initi-
Figure 2.8. The monosynaptic reflex. The figure depicts the monosynaptic
patellar tendon reflex. This reflex occurs when a tap on the patellar tendon
stretches muscle fibers of the quadriceps muscle. The dendrites of a sensory
neuron sense the stretch and pass this information to the cell body in the
dorsal root ganglion (just outside the spinal cord) and then into the spinal
cord. In the cord, there is a synapse between the axon terminals of the sen-
sory neuron and the dendrites of the motor neuron. The signal travels across
the synapse and initiates an action potential in the motor neuron, which
results in contraction of the quadriceps muscle and an extension of the lower
leg. Figure used with permission from G. G. Matthews, 1998, Neurobiology:
Molecules, cells, and systems, Oxford, England: Blackwell Science.
ating action potentials in sensory neurons that terminate inside the spinal
cord. At that synapse, a neurotransmitter is released which produces an
EPSP in a motor neuron which in turn projects back to muscle fibers in
the originating muscle. If the EPSP is of sufficient magnitude or if other
excitatory impulses are arriving, action potentials will be initiated in the
postsynaptic neuron. Once the impulse reaches the axon terminals, neu-
rotransmitter will be released at the motor end plate and a muscle action
potential will spread over the muscle, thus initiating movement.
Bioelectric Potentials
The bioelectric events—both action potentials and the graded, synaptic

potentials just described for neurons and muscles—provide an electro-
chemical record of neuromuscular functioning. By recording the appear-
ance and passage of action potentials or graded potentials, the researcher
can provide a representation of the action of nerves and muscles. By
placing an appropriate electrode in or near a nerve or muscle cell and
amplifying the observed potentials, the psychophysiologist records the
activity of neural, muscle, and gland cells underneath the electrode. In
general, the psychophysiologist's goal is to relate some aspect of behavior
to the function of large systems of cells in the body which may be spread
over a considerable area. For instance, we often record the electrical
processes that initiate the beating of the heart (the electrocardiogram or
EKG), the synchronous graded potential changes in millions of cortical
brain cells (the electroencephalogram, EEC), or changes in the activity of
many motor units in a muscle or muscle group (the electromyogram, or
EMG). Moreover, as noted in chapter 1, the psychophysiologist, who usu-
ally works with human subjects, is typically restricted to recording these
multicellular events from a relatively great distance, usually from the skin
surface.
What are the relationships between cellular bioelectric events and the
surface recorded potentials which are the primary interest of the psycho-
physiologist? In short, they are often difficult to determine. When action
potentials or graded potentials occur simultaneously in a great number
of cells, their summed potentials may be large enough to be recorded
from the skin surface. However, the summed potential is difficult to relate
to specific cellular functions; there are several reasons for this. First, cells
that are nearer the recording electrode will contribute more to the re-
corded potential than those that are farther away. If, for example, a re-
cording electrode is over a muscle, the motor units firing synchronously
nearer the skin surface will contribute more to the surface EMG than
motor units deep within the muscle tissue. Similarly, most of the activity
recorded from the scalp as an EEC is the result of graded potentials arising
from cortical brain cells rather than subcortical brain structures. Second,
N E U R O N S AND MUSCLES 33
the orientation of cells firing action potentials can influence the appear-
ance of a surface-recorded signal. Retreating action potentials (moving
away from the electrode) will be seen as increasingly negative, whereas
advancing potentials will be seen as increasingly positive. Repolarization
also can produce measurable voltages at the skin surface. For instance,
the T wave of the EKG is produced by the rapid repolarization of hundreds
of muscle cells in the ventricles of the heart. Thus, a surface-recorded
signal is a combination of potentials from many cells, some nearer the
electrode, some further away, some which are moving toward the elec-
trode and some away, some from cells which are depolarizing, others
from cells that are repolarizing. The difficulties in interpreting such po-
tential combinations is considerable.
Nevertheless, the potentials recorded from surface electrodes do result
from underlying nerve and muscle action. Surface-recorded potentials
result from either action potentials or graded potentials spreading over
nerve or muscle cells and then through the body via the interstitial (ex-
tracellular) fluid that surrounds all cells of the body. Thus, very large
signals such as those arising from depolarizing and repolarizing heart
muscle which manifest as the surface EKG can be recorded between any
two points on the body surface. However, the amplitude and waveform
of the potentials spreading through the interstitial fluid are greatly
changed as they spread. Like the ripple that forms in a pond into which
a stone has been thrown, the potential becomes smaller as it spreads. In
fact, the amplitude is so diminished that the waveform of the action
potential from a single cell cannot be recorded from the surface of the
skin. The amplitude of a signal is further reduced in passing through the
high impedance of the skin. This is the reason that psychophysiologists
recording a bioelectrical signal often must take great care to abrade the
skin over which an electrode is to be placed. Abrading reduces the im-
pedance of the skin to the passage of current between the tissue and the
electrode. Recording principles such as the importance of abrading the
skin will be discussed in greater detail where appropriate in chapters 7-
13.
Despite the potential complications in interpretation, one can, by care-
ful analysis and a knowledge of the underlying anatomy, relate surface-
recorded signals to certain aspects of neuronal and muscular functioning.
Thus, although there is not a simple mapping between the activity of a
group of neurons and a signal on the skin's surface, knowledge about
the underlying source of the signal recorded from the skin increases the
likelihood that one can make appropriate inferences about the "meaning"
of the signal. For example, during isometric contractions (where muscle
tension is increased but muscle length does not change), EMG amplitude
from electrodes placed over skeletal muscle fibers correlates reasonably
well with the force generated, at least for moderate to high levels of
contraction (Woods & Bigland-Ritchie, 1983). Likewise, the waves of the
EKG can be used to determine the timing of the electrical signal initiating
cardiac contraction. The potentials with which the psychophysiologist
works are, therefore, neither mysterious indices of cognitive or emotional
function nor simple translations of physiological processes carried to the
body surface. Rather, they are complex ramifications of the bioelectric
spread of action potentials in a conductive medium, the human body.
References
Berne, R. M., & Levy, M. N. (1998). Physiology (4th ed.). St. Louis, MO:
Mosby.
Berntson, G. G., Cacioppo, J. T., & Quigley, K. S. (1991). Autonomic deter-
minism: The modes of autonomic control, the doctrine of autonomic
space and the laws of autonomic constraint. Psychological Review 98,
459-487.
Edwards, A. V. (1990). Autonomic control of endocrine pancreatic and ad-
renal function. In A. D. Loewy & K. M. Spyer (Eds.), Central regulation of
autonomic functions (pp. 286-309). New York: Oxford University Press.
Gardner, E. (1975). Fundamentals of neurology (6th ed.). Philadelphia: Saun-
ders.
Guyton, A. C. (1987). Basic neuroscience: Anatomy and physiology. Philadel-
phia: Saunders.
Guyton, A. C., & Hall, J. E. (1996). Textbook of medical physiology (9th ed.).
Philadelphia: Saunders.
Ham, A. W. (1974). Histology (7th ed.). Philadelphia: Lippincott.
Loewy, A. D. (1990). Anatomy of the autonomic nervous system: An over-
view. In A. D. Loewy & K. M. Spyer (Eds.), Central regulation of autonomic
functions (pp. 1-16). New York: Oxford University Press.
Matthews, G. G. (1998). Neurobiology: Molecules, cells, and systems. Oxford:
Blackwell Science.
Morell, P., & Norton, W. T. (1980). Myelin. Scientific American, 242, 88-117.
Pansky, B., Allen, D. J., & Budd, G. C. (1988). Review of neuroscience (2nd ed.).
New York: Macmillan.
Woods, J. J., & Bigland-Ritchie, B. (1983). Linear and non-linear surface
EMG/force relationships in human muscles. American Journal of Physical
Medicine, 62, 287-299.
3
Equipment Used in
Psychophysiological Recording
In chapter 2 we discussed the sources of the bioelectric potentials that

can be recorded from the surface of the skin. The purpose of this chapter
is to trace the bioelectric signals from the skin, across the junction with
an electrode or transducer, into a polygraph or computer where it is
filtered and amplified, and finally to the point where it is displayed and
analyzed.
Electrodes and Transducers
We will describe only those electrodes designed to be placed directly on

the surface of the skin—cutaneous electrodes—although other types of
electrodes exist (such as needle electrodes) for both human and animal
work. In the second part of this section, we will discuss transducers for
measuring temperature, respiration, and blood volume.
Electrodes
Electrodes are usually small metal discs attached to the subject's skin for
the purpose of recording the underlying electrical activity. Two electrodes
must always be used and their location depends upon the particular phys-
iological signal of interest. We discuss electrode location in the chapters
dealing with specific measures. The one overriding concern in selecting
electrodes and electrode paste and in preparing the skin is to provide a
low-impedance, electrochemically stable path for the bioelectric potential
from the skin to the input of the polygraph. The low impedance is nec-
36
essary to keep the small bioelectric potentials from being seriously atten-
uated in crossing the skin, that is, before reaching the electrode. Chemical
stability prevents the development of unstable potentials at the electrode,
which would affect the biopotential. Electrodes are more than simply
terminals or contact points from which voltages can be obtained on the
surface of the body. Electrodes aid in converting ionic potentials gener-
ated by nerve, muscle, or gland cells within the body into electrical po-
tentials that can be measured. Complications arise, however, when met-
als such as those commonly used for electrodes are placed in contact
with an electrolytic substance, such as electrode paste or the skin. At the
electrode-electrolyte interface, an electrochemical reaction is produced
that creates a difference in voltage between the metal and the electrolytic
solution. In this manner, voltages may be produced at the electrode site
independent of the bioelectric event in the body. Thus, it is possible to
record unwanted voltages produced at the electrodes in addition to the
desired psychophysiological signal.
The potential or voltage produced by the electrodes themselves (the
bias potential or offset potential, as it is referred to by some investigators)
is a function of many factors, among them the particular metal, the type
of electrolytic solution, and the temperature. In a historical study, Lykken
(1959) constructed electrodes from different types of metals, placed them
in pairs in a saline solution, and then measured the potential difference
between the electrodes over time. After an hour, Lykken found that some
metals showed greater potential difference than others. For example, he
found platinum to produce a potential of 320 mV; silver, 94 mV; zinc,
100 mV; and chlorided silver (silver-silver chloride), only 2.5 mV. Thus,
it is not difficult to understand why silver-silver chloride electrodes are
preferred in most laboratories today. The advantage of silver-silver chlo-
ride electrodes is that (1) they introduce a relatively small initial mea-
surement error (bias potential); (2) they show a relatively small drift of
potential with use; and (3) they minimally develop polarization potentials.
With prolonged use, every pair of electrodes shows some polarization,
the buildup of a counter-electromotive force, which has the effect of an
apparent increase in subject resistance. Polarization can be thought of as
the unequal distribution of ions on the two electrodes as a function of
the passage of current through the electrolyte. That is, one electrode
becomes positive in relation to the other, and the two electrodes therefore
produce a potential or voltage of their own. Polarization generally can
be reduced by leaving the electrodes connected together in a saline so-
lution for several hours.
The electrodes in frequent use today are the floating or cup-shaped
type that is seen in most research labs and a disposable version which is
used by both labs and clinical settings. With this type of electrode, the
metal part does not come in direct contact with the skin; it contacts the
skin through a "cushion" of electrode paste. The floating or cup-shaped
E Q U I P M E N T USED IN P S Y C H O P H Y S I O L O G I C A L R E C O R D I N G 37
Figure 3.1. Commonly used floating or cup-shaped electrodes.
electrode (figure 3.1) is attached to the subject by an adhesive collar. The

collar has an adhesive material on both sides that sticks to both the skin
and the electrode, holding the electrode securely to the skin. Because of
the jelly like consistency of the electrode paste, the floating electrode is
less disturbed by small movements than electrodes rigidly affixed to the
skin. Using these electrodes, it is even possible to record muscle potentials
from moving athletes without significant artifact. Electrode paste is also
used for two other reasons. First, it lowers the impedance between the
electrode and the skin which is its most important function. Second, some
types of electrode paste also help to stick the electrode to the skin.
An early study conducted by Lewes (1965) demonstrated that for
some types of recording, many different electrode pastes will work satis-
factorily. This study recorded the electrical activity of the heart using
several types of electrode paste as well as no paste. In each case, the
heart signal was as good as that recorded with standard electrode paste.
What makes this interesting is that a standard laboratory electrode paste
was compared with mayonnaise, French mustard, tomato paste, hand
cream, and toothpaste! It should be noted that although almost any-
thing works for heart rate recording, most of these substances probably
will not suffice for signals of lower voltages, such as those recorded from
muscles and the brain. Also, the excessive salt contained in most stan-
dard laboratory electrode pastes makes them unsuitable for recording
some aspects of skin conductance and skin potential. In later chapters in
this book dealing with individual physiological measures, we will em-
phasize when the characteristics of the electrode paste used are impor-
tant. Although silver-silver chloride electrodes and improved electrode
pastes have reduced many of the problems of recording, one must still
be concerned about proper skin preparation in order to insure the best
possible signal. Because the outer layer of the skin is mainly dead cells,
dirt, and grease, these must be removed to lower impedance before the
electrodes are applied. There are numerous methods for reducing skin
impedance. One method is to abrade the area with fine sandpaper,
an abrasive cloth, or an electrode paste which contains an abrasive
material. The skin is then cleaned and dead cells wiped away with either
alcohol or water. (This procedure must not be used when recording elec-
trodermal activity; see chapter 13.) One should abrade the skin until
it is somewhat reddish but not bloody. The report of an ad hoc com-
mittee of the Society for Psychophysiological Research (Putnam, John-
son, & Roth, 1992) offers guidelines for reducing the risk of disease
transmission when applying electrodes to abraded skin. Once the elec-
trodes are in place, resistance should be checked with an impedance
meter. For a good recording, the impedance should be well below
10,000 ft.
The type of electrode and electrode paste used and the care that must
be taken in skin preparation are a function of the signal to be recorded.
Usually large amplitude signals can be satisfactorily recorded with min-
imal effort. However, weak signals or signals of low frequency require
more stable electrodes and lower skin impedance. The greatest care must
be taken when recording DC potentials, such as in skin potentials and
certain brain potentials.
Transducers
When energy changes form, it is said to be transduced. Transducers are
used to convert physiological events to electrical potentials suitable for
electronic amplification. All physiological responses must be transduced
to voltage for the polygraph or computer even when the measure of
interest is resistance or conductance. It is often convenient to record some
result of nerve or muscle action rather than to record the action poten-
tials themselves. For example, the pupil of the eye constricts and dilates
through the contraction of muscle cells within the eye. While changes
in pupil diameter are an interesting response, the muscle action potentials
that produce them are difficult or nearly impossible to record directly.
Alternatively, the result of the action potentials, the pupil diameter, may
be more accessible and can be used to reflect underlying activity. If we
transduced changes in pupil diameter to changes in some electrical pa-
rameter, those changes could be introduced to the physiological amplifier.
In fact, we can use the amount of light reflected from the pupil onto a
photosensitive resistor for the purpose. The possibilities for transducers
are endless. Researchers can transduce any result of combined nerve,
muscle, and glandular action to some form of electrical change. In prac-
tice, however, transducers typically respond to cardiovascular, respira-
tory, and somatic muscle changes with changes in their resistance to the
passage of current. We will describe several of the more commonly used
transducers here.
Most strain gauge transducers works on the principle that tension or
strain in a metallic conductor changes the resistance and therefore the
flow of electricity. A constant electrical voltage is applied to the metal,
and changes in its resistance (due to changes in tension) are noted. One
common application of the strain gauge is in the measurement of respi-
ration. The specific type of gauge commonly utilized is a small, flexible
tube filled with mercury which is placed in an extended position across
the subject's chest or around the body. As the person inhales, the chest
enlarges and increases the tension in the tube, thereby increasing the
resistance in the circuit. In this manner, changes in resistance reflect
respiratory movements (see chapter 10).
Another common transducer is the photoconduction cell (or photo cell),
which varies its resistance with the amount of light that hits it. Research-
ers often use such cells to indicate changes in blood volume in some body
area. Light in the red to infrared range passes through living tissue, such
as the skin, but such light is poorly transmitted and readily reflected by
blood. This principle is used to determine the amount of blood passing
through a finger, toe, ear lobe, penis, or clitoris. A small light is focused
on, for example, the finger and mounted next to the photo cell. As the
blood enters the finger, the amount of light reflected back to the cell
changes; thus one can observe a change in the resistance of the cell. This
resistance change corresponds to changes in the amount of blood (see
chapter 12).
A third type of transducer is a thermoresistive transducer or thermistor,
a device that changes its resistance in relation to its temperature. One
popular use of this transducer is the measurement of skin temperature,
such as from the hand. As the temperature of the hand increases, the
resistance of the thermistor changes.
Polygraphs
Polygraphs record a physiological signal in a continuous analog fashion

rather than digitally sampling it in discrete units as is the case with
computers. Many psychophysiological laboratories employ a multichan-
nel polygraph. Such devices can display one or many physiological events
recorded on a paper chart. A polygraph generally has three separate
components through which the signal passes before being displayed. The
first is a coupler or signal conditioner, which is designed to make the
GENERAL ELEMENTS OF PSYCHOPHYSIOLOGY

electrical characteristics of all signals compatible, regardless of the type
of transducer from which the signals originated. The second component
is a preamplifier; the third is a main amplifier, which has the function of
producing an output of sufficient voltage to drive the transcribing pens
or standardize the voltage for computer storage and analysis.
Couplers
The coupler conditions the psychophysiological signal coming from the
subject. In some cases, the coupler does nothing more than supply the
signal to the preamplifier; in other cases, the coupler changes the form of
the signal to meet the requirements of the amplifier. Sometimes the cou-
pler is used to provide external voltage to a transducer, electronic balanc-
ing, or calibration. Other couplers provide selective filtering or perform in-
tegration or rate computation. There are specialized couplers designed for
heart rate, EMG, respiration, EEC, and other standard measures.
Filtering
Filtering of the signal can occur in the coupler and/or in either the pre-
amplifier or power amplifier. Filters remove or reduce (attenuate) certain
parts of the input signal and allow other parts of the signal to pass. Low-
pass /liters allow only frequencies below a certain frequency to pass. For
example, if a low-pass filter set at 12 Hz is introduced into a circuit, then
all frequencies above 12 Hz would be attenuated or reduced. A high-pass
filter is similar to a low-pass filter, except that it allows frequencies only
above a certain frequency to pass unattenuated. A notch filter will atten-
uate only a small range of frequencies while allowing all others to pass
undiminished. A notch filter may be described as a band-reject filter.
Notch filters are most often set at 60 Hz (the frequency of AC current in
the United States) or at 50 Hz (the frequency of current in Europe and
other areas) to remove unwanted interference. Figure 3.2 shows a re-
Figure 3.2. Recording of an EKG with and without a 60-Hz notch filter.
E Q U I P M E N T USED I N P S Y C H O P H Y S I O L O G I C A L R E C O R D I N G 41
Figure 3.3. Recording of an EKG with different time constants.
cording of an EKG with and without the 60-Hz notch filter. Another
type of filter is the band-pass filter. This filter is really a combination
of a high-pass and a low-pass filter adjusted such that only a certain
range of frequencies can pass. For example, a researcher interested in the
alpha rhythm (8-12 Hz) of the EEC, can set a band-pass filter to pass
only those frequencies between 8 and 12 Hz and to attenuate all other
frequencies.
Depending on how they are set, some couplers and amplifiers can
handle either DC or AC signals. DC circuitry is designed for the recording
of DC signals and signals of very low frequency. AC circuitry is designed
for situations in which one wants to record higher frequencies and elim-
inate low frequencies. For example, EKG and EEC signals are generally
recorded utilizing AC circuitry to avoid recording slow changes in poten-
tial, which would cause the signal to "drift" and thus make the changes
of interest more difficult to interpret.
The reduction of the low frequencies in an AC circuit is determined
by the time constant utilized. Stated simply, the time constant is defined
as the amount of time required for a rectangular signal (step function)
to return to 63% of its voltage. To illustrate this, figure 3.3 shows the
same EKG recorded with different time constants. Note the difference in
the baseline of the AC and DC recordings.
Amplification
From the coupler the signal goes to the preamplifier, which increases
the amplitude of the signal to a level that can be accepted by the power
amplifier. The function of the amplifier is to increase the size of the
physiological signal recorded at the surface of the skin to approxi-
mately 1 V. This typically will require an amplification factor or gain of
at least 1,000. That is, a millivolt (1/1000 of a volt) would have to be
amplified 1,000 times to equal 1 V. Since it is possible for nonphysio-
logical signals (e.g., 60 Hz) to be picked up and amplified with the desired
signal, there exists a term to describe the amount of signal (the biopo-
tential) in relation to other electrical activity (generally referred to as
noise). This is the signal-to-noise ratio. This ratio gives some indication of
how much noise, or nondesired electrical activity, any electronic instru-
ment will introduce. A term related to signal-to-noise ratio is common
mode rejection. This refers to the amount of noise or interference pres-
ent at the output of a differential amplifier. The higher the number in
both cases, the better the physiological signal will be in relation to the
noise of the total electronic environment in which the recording is taking
place.
Today, the basic unit of construction for biomedical equipment
such as amplifiers is the integrated circuit (1C). The integrated circuit is a
multicomponent device (transistors, resistors, capacitors, diodes, and so
forth) in which the components have been miniaturized and designed
for specific purposes. For example, it is possible to purchase an inex-
pensive high-gain amplifier only a centimeter or so in size. An 1C of
this size may contain tens of thousands of components. Because of the
1C, the psychophysiologist today is rarely required to construct com-
plete circuits, or even to repair the existing ones, as was the case previ-
ously. However, it is important to have some knowledge of the types,
properties, and functions of amplifiers, filters, and related coupling
devices.
Display
After the signal has been filtered and amplified, the power amplifier pro-
vides the voltage necessary to drive the recording device, usually a robust
galvanometer. The galvanometer is an electromechanical device which
uses a moving coil to drive the polygraph pen. Polygraphs can be pur-
chased with ink-writing, heat-writing, or light-writing systems. Ink writ-
ing is the most common because the initial cost is less and the recording
paper is considerably less expensive.
Computers
The computer can be utilized to record and quantify data as long as the
physiological signal can be converted into a numerical form. The device
used to perform this function is the analog-to-digital (A-D) converter. The
function of this device is to take a continuous signal and convert it into
discrete steps. One way of thinking about an A-D converter is to imagine
it as a voltmeter that samples at a very fast rate and then stores each of
the samples. The speed at which the converter samples is referred to as
the sampling rate. A good rule of thumb is that the sampling rate per
second should be at least two to five times faster than the fastest fre-
quency component of interest in the signal. This rule, the Nyquist rela-
tion, is discussed more fully in chapter 14.
Computers can also perform the functions described previously in
terms of a polygraph, although they do this in a digital rather than an-
alog manner. In terms of filtering, for example, the digital computer
would use a mathematical formula rather than electronic circuitry to
emphasize particular frequency components in a physiological signal. We
will describe some of these mathematical transformations in the chapter
on signal processing (chapter 14).
The computer also can be used to reduce the data. If the incoming
signal is that of the biopotential from the heart and the desired measure
is heart rate, then one does not need every value of the signal that is
sampled. One would really only need to know when the biopotential was
above a certain level, as in the case of the larger spike in the EKG, because
this occurs precisely once per beat. A researcher could program the com-
puter to compare each voltage coming from the A-D converter with a
given voltage that was known to be present only when there was an
EKG spike. A clock within the computer could be started and the time
between each heart beat recorded and stored for future analysis.
A computer can also be utilized to recognize established psychophys-
iological patterns. For example, a researcher can program a computer to
check a given EEC against a certain standard to determine if specific
parameters are met. One example of such a use might be the determi-
nation of epileptic activity in the EEC. Or the computer might be used to
determine whether certain frequencies, alpha for example, are present in
the EEC signal. Using this type of analysis, the researcher could program
a computer to give feedback or a type of stimulus only when certain
preestablished conditions were met. For example, the presence of alpha
activity without the presence of any eye movement could be one such
combination. Although computers provide a means of making quick de-
cisions concerning data, it must always be remembered that the com-
puter makes its decisions in accordance with the instructions given to it.
That is, a computer cannot discriminate any better than the person who
programs it.
The ultimate value of the computer is that, if it is programmed prop-
erly, it can be designed to run entire experiments and even to analyze
the data at the conclusion. This is particularly powerful in so-called real-
time or on-line environments. For example, one could conduct a biofeed-
back experiment in real time; that is, the computer would provide feed-
back to the person in relation to actual physiological changes that were
being made at that moment.
If data from an experiment were previously recorded on the computer
disk or other storage device and the computer analyzed the data, the
computer usage would be considered off-line and not in real time. That
is, the experimenter could analyze the data in any manner or sequence
desired without regard to how the events happened in real time. Let us
end on one cautionary note. Although computers can quickly record and
analyze physiological data, it is still critical that the researcher have a
means for visually inspecting the data to determine that what is being
recorded is an accurate physiological signal undistorted by, for example,
movement, coughing, or electrical interference from other equipment.
References
Brown, C. C. (1967). Methods in psychophysiology. Baltimore, MD: Williams

and Wilkins.
Cornsweet, T. M. (1963). The design of electric circuits in the behavioral sciences.
New York: Wiley.
Cromwell, L., Arditti, M., Weibell, F. J., Pfeiffer, E. A., Steele, B., & Labok, J.
(1976). Medical instrumentation for health care. Englewood Cliffs, NJ:
Prentice-Hall.
Cromwell, L., Weibell, F.J., Pfeiffer, E. A., & Usselmann, L. B. (1973). Bio-
medical instrumentation and measurements. Englewood Cliffs, N. J.:
Prentice-Hall.
Dewhurst, D. J. (1976). An introduction to biomedical instrumentation. Oxford:
Pergamon.
Ferris, C. D. Introduction to bioelectrodes. (1974). New York: Plenum.
Geddes, L. A. (1972). Electrodes and the measurement of bioelectric events. New
York: Wiley-Interscience.
Lewes, D. (1965). Electrode jelly in electrocardiography. British Heart Journal
27, 105-115.
Lykken, D. T. (1959). Properties of electrodes used in electrodermal measure-
ment. Journal of Comparative and Physiological Psychology, 52, 629-634.
Putnam, L. E., Johnson, R., & Roth, W. T. (1992). Guidelines for reducing the
risks of disease transmission in the psychophysiology laboratory. Psy-
chophysiology, 29, 127-141.
E Q U I P M E N T USED I N P S Y C H O P H Y S I O L O G I C A L R E C O R D I N G 45
Venables, P. H., & Martin, L. (1967). A manual of psychophysiological methods.
Amsterdam: North-Holland.
Welkowitz, W., & Deutsch, S. (1976). Biomedical instruments. Theory and de-
sign. New York: Academic Press.
Zucker, M. H. (1969). Electronic circuits for the behavioral and biomedical sci-
ences. San Francisco: Freeman.
4
Psychophysiological Recordings
... somatic responses abound. One has but to observe them on

a set of recording instruments to believe that they are by far the
most numerous responses of the organism. It is clear that any
overt response, vocal utterance, or bodily movement is surrounded
by a wide penumbra of them, and it may not be too bold a guess
to say that whenever there is any evidence of a stimulus affecting
the individual, something in his periphery or viscera is set into
motion. Not infrequently these may be seen when there is no
other means at hand for detecting that the person has been
stimulated.
(Davis, Buchwald, and Frankmann, 1955)
Somatic responses such as heart rate and muscle potentials do indeed

abound. Electrical activity can be recorded from the surface of the skin
at any moment. Background electrical activity is always present and
spontaneous changes in activity occasionally appear, even when it seems
that an individual is relaxing or asleep. This makes the task of recording
and correctly interpreting a subject's evoked response to a specific stim-
ulus or situation a challenge.
Most psychophysiological recordings can be analyzed in terms of three
types of activity: spontaneous, tonic (background), and phasic (evoked re-
sponses).
Figure 4.1 shows a section of a typical recording. We will refer back
to this figure in this chapter as we define and discuss spontaneous, tonic,
and phasic activity.
47
Figure 4.1 Recording of respiration EKG, and skin Conductance showing
spontaneous, tonic, and phasic activity.
Spontaneous Responses
What does a spontaneous physiological response to an unknown stimulus

look like? It is the same as a physiological response to a known stimulus
such as a tone, a shock, or a snake. It may take the form of a change in
heart rate or skin conductance in the palm of the hand, muscle potential,
and so on. Spontaneous activity is a name given in ignorance to what
may be several different types of responses. What is referred to as spon-
taneous activity is in reality a change in physiological activity that occurs
in the absence of any known stimuli.
Point B in Figure 4.1 shows an increase in heart rate and an increase
in skin conductance to an unknown stimulus. In most if not all cases,
such a spontaneous response would be the result of some CNS activity.
For example, the subject might have just had an anxious thought about
what procedure the experimenter was going to perform. However, unless
the subject is asked about these thoughts immediately following the re-
sponse, the experimenter would not know the cause of the so-called spon-
taneous activity. Changes in heart rate, skin conductance and/or other
physiological responses that follow a deep breath, movement, or a
cough—that is, following identifiable stimuli—are not considered to be
spontaneous responses. Point C, in figure 4.1 shows such a series of
events. Spontaneous ANS responses that involve no CNS activity have
not been studied by psychophysiologists and probably occur rarely if at
all. An example would be an increase in skin conductance that results
from localized biochemical changes immediately under the recording
electrode.
Why are spontaneous physiological responses of interest? When con-
ducting research, it is important to be aware of the existence of such
responses in order to avoid misinterpreting the data resulting from a
study of the effects of some known stimulus on specific physiological
activity. An obvious example would be the presentation of a stimulus to
the subject while the subject is making a spontaneous response. The net
effect might be a response greater than normal; then again, it might be
smaller if the spontaneous response is in the opposite direction. In either
case, quantification of the stimulus-contingent response will be difficult.
Tonic Activity
Tonic activity is often referred to as the background level or resting level

of activity of a particular physiological measure. However, resting level
is not a good description of tonic activity because what we are really
discussing is the level of activity when a subject is (1) not making a
spontaneous response and (2) not making a discrete response to a known
stimulus. The subject might be highly aroused and certainly not resting.
For example, the tonic level of one individual's heart rate while waiting
in a dentist's office prior to a root canal operation might be 130 beats
per minute. Another subject in the same situation, perhaps someone not
so bothered by dental work, might have a tonic level of heart rate of 80
beats per minute. Thus, tonic level is simply the level of activity of some
ANS or CNS measure at a particular point in time prior to stimulation.
In figure 4.1, point A or point D would be considered examples of tonic
activity. Point A is just prior to a spontaneous response; point D precedes
a response to a tone.
There are two separate reasons for recording and caring about tonic
level. First, tonic level is of interest in its own right as a measure of the
activity of the ANS or of muscles or brain when the subject is not re-
sponding to a specific stimulus. In such cases it is a complex result of
many factors, including CNS reactions to environmental and internal
stimuli, plus the delicate interplay between the sympathetic and para-
sympathetic branches of the ANS. People with essential hypertension and
tension headaches are examples of individuals with extreme tonic levels
of responding. Individuals with essential hypertension have high blood
pressure due to elevated levels of sympathetic nervous system activity
which keep their blood vessels in a highly contracted state. Muscle po-
tential recordings from individuals who suffer from frequent tension
headaches reveal that even when they are not experiencing headaches,
the muscles of their head and/or neck show higher levels of activity than
those of people who do not have tension headaches.
PSYCHOPHYSIOLOGICAL RECORDINGS 49
The second reason tonic level is of interest is that in some cases the
size of a response to a specific stimulus depends upon the tonic level as
measured immediately prior to the stimulus. For example, if one's heart
rate is already close to the highest level that that person's heart can beat,
it is unlikely that any stimulus will make it go much higher. This rela-
tionship is referred to as the law of initial values. This is a general principle
of psychophysiology and will be discussed more fully in chapter 5.
The tonic level measured immediately prior to stimulation is referred
to as the baseline, the level of activity against which we compare the
phasic response to a stimulus. One important issue in psychophysiological
recording is how long the baseline period should be. The answer is that
it should be long enough to provide a stable prestimulus level and long
enough to provide sufficient data for an appropriate analysis. On the
other hand, it should be short enough so that the subject is neither bored
nor anxious about the delay in instructions. For example, in recording
the electrical activity that accompanies the contractions of the stomach
(the electrogastrogram or EGG; see chapter 11), we find that the signal
stabilizes in about 6 min. However, even if the EGG stabilized in 2-3 min,
we would still need to record for a minimum of 4 min because that is
the minimum length of time needed for the type of analysis used with
EGG recording. Gerin, Pieper, and Pickering (1994) presented data show-
ing that a 5-min baseline period is sufficient for cardiovascular recording.
They also provide evidence that the baseline is not effected by the antic-
ipation of a stressful task. In addition to the length of the baseline period,
a second related issue is what the subject should be instructed to do
during the baseline period. Most experimenters just tell the participants
to sit quietly and relax. But not all subjects will or can follow those
instructions. Jennings, Kamarck, Stewart, Eddy, and Johnson (1992)
have suggested giving subjects a simple cognitive task that will create
what they call a "vanilla baseline" and standardize the mental activity
of all subjects during the baseline period.
Most psychophysiologists are less interested in tonic level than in re-
sponses to specific stimuli. The present authors, about 50 years after
Schlosberg (1954), support his warning that this emphasis on stimulus-
contingent responses (that is, phasic responses) and relative neglect of
tonic levels is analogous to not seeing the forest for the trees.
Phasic Activity
Phasic activity is a discrete response to a specific stimulus—an evoked

response. Point E in figure 4.1 shows a phasic response to a tone. Phasic
activity can be an increase or a decrease in either the frequency or am-
plitude of a response, or a more complex change in wave form or latency.
The most important factor to consider when quantifying phasic activ-
ity is that the subject's response to, for example, a slide of an American
flag is not being made against a background of zero activity. The subject
is constantly responding to internal as well as external stimuli—somatic
responses abound. Two difficulties sometimes occur: (1) determining the
magnitude of a subject's phasic response to a specific stimulus and sep-
arating it from other phasic responses and from spontaneous activity,
and (2) attempting to introduce a correction factor for the magnitude of
the phasic activity as a function of the preceding tonic activity, that is,
the law of initial values. We address these issues in chapter 5 and where
relevant in Part II, for example in chapter 13.
References
Davis, R. C., Buchwald, A. M., & Frankmann, R. W. (1955). Autonomic and
muscular responses and their relation to simple stimuli. Psychological
Monographs, 69 (Whole No. 405).
Gerin, W., Pieper, C., & Pickering, T. G. (1994). Anticipatory and residual
effects of an active coping task on pre- and post-stress baselines. Journal
of Psychosomatic Research, 38, 138-149.
Jennings, J. R., Kamarck, T., Stewart, C., Eddy, M., & Johnson, P. (1992).
Alternate cardiovascular baseline assessment techniques: Vanilla or rest-
ing baseline. Psychophysiology, 29, 742-750.
Schlosberg, H. (1954). Three dimensions of emotion. Psychological Review, 61,
81-88.
PSYCHOPHYSIOLOGICAL RECORDINGS 51
5
Some Basic Principles of
Psychophysiology
The basic principles of psychophysiological recording are generalizations

workers in this field have arrived at based on thousands of psychophys-
iological experiments. A familiarity with these principles will not only
provide the reader with information about certain relationships between
psychological and physiological variables but will also alert the reader to
factors—other than the independent variable—that might influence the
data collected in a psychophysiological experiment. Therefore, in this
chapter we will discuss some basic relationships that are of interest in
themselves but that sometimes make it difficult to see the effects of other
variables being studied or to interpret the results properly. With an un-
derstanding of these generalizations, the reader should be better equipped
to design new psychophysiological studies and to understand earlier
publications.
Arousal and Habituation
Arousal
The concept of arousal is so basic to psychophysiology that we will discuss
it first. New students of psychophysiology, when asked what specific prob-
lem they would like to study, often respond in one of the following two
ways: (1) "I would like to record a physiological measure of arousal and
determine the relationship of arousal to some behavior, such as problem
solving," or (2) "I would like to record a physiological measure so that
I can see how aroused participants will get when I show them slides of
something gruesome, such as a car crash." These students are usually
making two assumptions. First, they assume that arousal extends along
an unidimensional continuum from deep sleep to high agitation. Second,
52
they assume that one's position on the arousal continuum can be deter-
mined by measuring any one of several physiological variables, such as
heart rate or skin conductance. Before we discuss why both assumptions
are too simplistic, let us briefly relate them to the theories of some well-
known investigators.
The concept of arousal or activation has its roots in Cannon's (1915)
notion of the unified body preparing for fight or flight. Duffy (1957) ex-
tended this concept to include the intensity aspect of all behavior. As
shown in figure 5.1, Duffy hypothesized an inverted U-shaped curve re-
lating level of activation to performance. If our measure of performance
is how fast our participants can run the 100-m dash and our measure
of level of activation is their heart rate, we might well obtain an inverted
U-shaped curve. Those who are too minimally aroused may start slowly,
and those who are too highly aroused may commit false starts or suffer
from a feeling of "spongy knees" which often accompanies stage fright.
For a more detailed account of Duffy's theory, we refer the interested
reader to her book Activation and Behavior (Duffy, 1962). Other early
investigators who were interested in the relationship of arousal to be-
havior were Lindsley (1952) and Malmo (1959).
Let us examine some of Lacey's (1967) criticisms of activation theory.
Lacey suggested that there are at least three different forms of arousal:
cortical, autonomic, and behavioral. He pointed out that each is very
complex, not a simple continuum. He presented evidence showing that
Figure 5.1. Duffy's hypothesized inverted U-shaped function relating level of

activation and performance.
S O M E B A S I C P R I N C I P L E S OF P S Y C H O P H Y S I O L O G Y 53
one form of arousal cannot always be used as a valid measure of another
form of arousal. As an example of the complexities that exist within one
form of arousal, consider autonomic indices of sexual arousal when het-
erosexual participants view slides of opposite-sex nudes. If we measure
vasomotor activity in the finger, we would probably not detect any mean-
ingful changes and might conclude that our participants are not aroused;
however, if we record heart rate, we might come to a different conclusion.
The explanation is that sexual arousal prior to climax is primarily a par-
asympathetic response, while vasomotor activity in the finger is governed
by the sympathetic nervous system. Heart rate is controlled by both the
parasympathetic and sympathetic systems; thus, we should expect to see
some heart rate changes associated with the parasympathetic changes
that accompany sexual excitement.
Another major criticism of activation theory can be found in the prin-
ciple of stimulus-response specificity. Basically, this principle states that
specific stimulus contexts—for example, noticing that your wallet or
purse is missing—bring about certain patterns of responding, not just an
increase or decrease in an unidimensional activation continuum. The
pattern might include an increase in muscle tension, skin conductance,
and respiratory amplitude, but a decrease in respiration rate and heart
rate.
As a final point in his criticism of activation theory, Lacey referred to
a special case of stimulus-response specificity which he termed directional
fractionation. Picture the following scene and predict what is happening
in terms of Lacey's three forms of arousal—cortical, autonomic, and be-
havioral. Four soldiers are in enemy territory, and it is late at night. One
soldier is on guard duty while the others sleep. Suddenly there is an
approaching noise in the darkness. What happens to the soldier's level
of arousal? If we measure EEC activity, we would probably observe an
increase in cortical arousal. If we measure skin conductance and heart
rate as autonomic indices, however, we would likely find an increase in
skin conductance, but a decrease in heart rate. This is what Lacey meant
by directional fractionation—response directions are not uniform. Fur-
thermore, if we observe the physical behavior, we would see that the
soldier is probably standing very still, looking attentively toward the
source of the noise and trying to determine if it is being made by friend
or foe. A concept of activation based on a unidimensional continuum
cannot deal adequately with such complexities.
Habituation
The concept of habituation is just as basic to psychophysiology as the
concept of arousal. In a sense, the two concepts are complementary.
Whereas arousal suggests heightened responding to a stimulus, habitu-
ation is the reduction of responding that occurs to the repeated presen-
tation of the same stimulus. Habituation is a fascinating process to study
in itself because of its ubiquity across a broad range of species and re-
sponse systems; it is also important to be aware of habituation as an
experimenter because it is commonly observed in psychophysiological
experiments. For example, responses observed at the end of an experi-
ment may be significantly smaller than those at the beginning because
of habituation to the laboratory setting or to the repeated presentation
of stimuli with similar features.
There are two general forms of habituation, short-term and long-term.
Short-term habituation occurs within a single testing session. Figure 5.2
shows the short-term habituation of heart rate responses to a repeatedly
presented 60 dB tone. Long-term habituation occurs when responses de-
crease in size over the course of multiple testing sessions separated by
days or even weeks.
Numerous investigations have shown that responses generally habit-
uate slowly to stimuli that are unique, complex, or intense. Responses
habituate rapidly to stimuli that are presented frequently and slowly to
stimuli that are infrequent. Habituation may be somewhat inhibited
if the participant is required to make a behavioral response (e.g., to rate
the subjective intensity of the stimuli) or if other stimuli—different from
the repeated stimulus—are presented within an interstimulus interval.
Despite the wide range of species and responses that display habitu-
ation, there is no universally accepted explanation of why habituation
Figure 5.2. Habituation of the heart rate response to a moderate tone during
trials 5,10, and 15. Each data point represents the heart rate of one fictitious
participant.
SOME BASIC P R I N C I P L E S OF PSYCHOPHYSIOLOGY 55

occurs. Among the most well-known theories of habituation are Soko-
lov's (1963) comparator theory of habituation and Groves and Thomp-
son's (1970) dual-process theory of habituation.
Sokolov's comparator theory of habituation compares sensory infor-
mation to a "neuronal model" of an anticipated stimulus. It was thought
that the subject formed a model of a stimulus after the stimulus was
presented many times. An assumption of the comparator theory was that
when sensory input (i.e., a perceived stimulus) matches a model, a re-
sponse to the input is inhibited. A mismatch between the input and the
model was hypothesized to result in a response (e.g., orienting). The am-
plitude or strength of a given response was said to be a function of how
well sensory input matches the model. Early in stimulus repetition, re-
searchers observe strong responses because the mismatch between sen-
sory input and the evolving model is large. After repetition, the model
comes to represent the stimulus better; consequently, the mismatch be-
tween a stimulus and its model declines and response strength weakens.
Groves and Thompson proposed that two opposing processes—habit-
uation and sensitization—simultaneously occur during the course of re-
peated stimulus presentation. According to Groves and Thompson, the
habituation process mediates response decrement while sensitization acts
to inflate response size. Habituation was thought to occur by decreased
neuronal transmission along the stimulus-response pathway. Sensitiza-
tion was thought to occur by increased activation of an organism's "state
system," which governs general arousal. Groves and Thompson argued
that a response incrementing process, such as sensitization, was a nec-
essary postulate because increased—along with decreased—responding
is commonly observed during the course of stimulus repetition, especially
when the repeated stimulus is intense. For more information about how
these two theories relate to the habituation of psychophysiological re-
sponses, see Graham (1973) and Stephenson and Siddle (1983).
Orienting, Defensive, and

Startle Responses
For more than seven decades, a considerable amount of research has

been directed toward the study of orienting, defensive, and startle responses.
The measurement of these responses has been utilized extensively in the
study of development, emotion, learning, and attention in both humans
and animals (see Campbell, Hayne, & Richardson, 1992, and Lang, Si-
mons, & Balaban, 1997, for an overview). Because of the substantive role
these responses play in the study of behavioral and cognitive processes,
it is important to distinguish orienting, defensive, and startle response
profiles. As an experimenter, it is also important to be aware of the types
of stimuli that elicit these basic responses.
56 G E N E R A L E L E M E N T S OF PSYCHOPHYSIOLOGY
Orienting Response
The orienting response directs our attention to novel stimuli and en-
hances sensory processing: it is the "what-is-it?" response. Beginning
with Pavlov, Russian physiologists and psychologists studied this re-
sponse extensively. Pavlov first became aware of the phenomenon when
dogs, which had been conditioned by his students, failed to perform prop-
erly when he entered the laboratory. The problem was that the dogs paid
attention to Pavlov—that, is they made an orienting response to him—
instead of to the stimulus being presented by the student. Pavlov noted:
It is this reflex which brings about the immediate response in men and
animals to the slightest changes in the world around them, so that
they immediately orientate their appropriate receptor organ in accor-
dance with the perceptible quality in the agent bringing about the
change, making a full investigation of it. The biological significance of
this reflex is obvious. If the animal were not provided with such a reflex
its life would hang at any moment by a thread. (1927, p. 12)
Some of the major components of the orienting response include (1)

decreased irrelevant motor activity; (2) high-frequency, low-voltage EEC
activity; (3) peripheral vasoconstriction and cephalic vasodilation; (4) in-
creased skin conductance; (5) delayed respiration followed by an increase
in amplitude and a decrease in frequency; and (6) heart rate deceleration
(Graham, 1979; Graham & Clifton,1966; Lynn, 1966; Sokolov, 1963;
Turpin, 1983, 1986). Note that peripheral vasoconstriction and cephalic
vasodilation during orienting have been difficult to replicate and some
authors have even reported that vasoconstriction in both the periphery
and head occur to novel stimuli.
It appears that the function of this response is to prepare us to deal
with novel stimuli—for example, the unexpected noise of an acorn falling
on dry leaves while we are resting under an oak tree deep in the woods.
Once we determine that the stimulus possesses no significance, there is
no reason why we should prepare to deal with, for example, the noise
made by other falling acorns. Because the orienting response is highly
sensitive to stimulus novelty, it habituates rapidly after stimulus repeti-
tion (that is, after the stimulus becomes a common event).
Defensive Response
Defensive responses are thought to protect us from the possible dangers
of intense, painful, or threatening stimuli. Additionally, defensive re-
sponses trigger physiological adjustments that prepare an organism for
action (e.g., "fight or flight"). These adjustments typically involve (1)
increased skeletal muscle blood flow, (2) decreased blood flow to the gut,
S O M E BASIC P R I N C I P L E S OF P S Y C H O P H Y S I O L O G Y 57
(3) increased blood pressure, (4) peripheral and cephalic vasoconstriction,
and (5) heart rate acceleration (Cannon, 1915; Sokolov, 1963; Turpin,
1979, 1983, 1986; Turpin & Siddle, 1978; Viken, Johnson, & Knutson,
1991). In general, defensive responses habituate slowly, but depending
upon the intensity of a stimulus and the extent to which a repeated
stimulus actually signals danger or induces pain, this response may ha-
bituate quite rapidly. In addition, Turpin (1986) suggested that homeo-
static mechanisms may inhibit the repeated expression of a defensive
response in a short time period.
Startle Response
The startle response is elicited by an intense stimulus with a sudden or
abrupt onset such as the crack of a lightening bolt. Graham (1992) con-
tends that the function of the startle response is to interrupt or disengage
an organism from ongoing activity. The human startle response involves
both somatic and cardiovascular components. Using high-speed photog-
raphy and the sudden shot of a pistol behind their participants, Landis
and Hunt (1939) were among the first to note that the startle response
involves a reflexive eyeblink and whole-body jerk. The startle response is
also characterized by an immediate heart rate acceleration and rapid
habituation of elicited physiological responses. Contemporary psycho-
physiologists employ stimuli of much less intensity than gunfire (e.g.,
short bursts of white noise) and typically focus on the eyeblink and heart
rate components. It is important to note that components of the startle
response differ from those of the defensive response primarily in speed.
For example, the heart rate acceleration during startle typically peaks at
approximately 4 s, whereas heart rate acceleration during a defensive
response peaks at approximately 30 s (Reyes del Paso, Godoy, & Vila,
1993; Turpin, 1986). These differences are consistent with the functional
importance attributed to each response; startle responses are thought of
as fast acting and interruptive, but defensive responses involve more wide
spread neurohumoral adjustments that facilitate behavioral action.
A practical reason for becoming aware of the circumstances under
which orienting, defensive, and startle responses normally occur, and
what they look like, is that the physiological response of participants to
an independent variable in an experiment might be confounded by (that
is, intermixed with) these responses. For example, whenever a series of
slides is presented, the subject's responses to the first few will be partly
caused by the content of those slides and partly by an orienting response.
One way to deal with this problem is to disregard the data from the first
few trials. Defensive or startle responses may be elicited during an ex-
periment if participants hear a nearby door slam.
Homeostasis and
Autonomic Balance
Homeostasis
The word homeostasis has been used to describe both a state of the or-
ganism and a process which takes place within the organism. When we
refer to the homeostatic state, we are identifying equilibrium, stability,
constancy, and the like. What we are really describing is a steady-state
internal environment providing the right temperature, nourishment, oxy-
gen, and fluids for optimal functioning of all cells. In our opinion, it is
no more meaningful to specify the homeostatic state of the whole organ-
ism than it is to specify the stability of an entire city. In both cases, some
parts (physiological systems or neighborhoods) may be very stable, while
others are not. A typical statement appearing in the literature is: "Some
animals are more successful than others in maintaining homeostasis."
We feel that such a statement should always be qualified. In actuality,
some animals are more successful in maintaining, for example, constant
temperature, whereas others are more successful in maintaining constant
blood pressure.
The mechanism that underlies the homeostatic process is negative
feedback. However, the presence of the homeostatic state—that is, sta-
bility—is not invariably a sign of negative feedback. An example of a
negative feedback system outside of our bodies is the thermostat that
controls the temperature of a room. The thermostat senses the temper-
ature; the warmer the room, the less heat it allows in.
Let us now examine Davis's (1958) analysis of the homeostatic control
of body temperature in human beings. When we become too hot, regu-
lation is accomplished by the evaporation of water secreted through the
skin. Temperature is regulated, but what is happening to the sweating
mechanism? Davis' answer was that every bit of reduced variation in
temperature is brought about by increased variation in sweating. This
he called heterostasis.
The general concept of homeostasis as first stated by Claude Bernard—
"Le fixite du milieu interieur est la condition de la vie libre" ("The sta-
bility of the internal environment is the condition of a healthy life")—
and later supported by Cannon (1939) in his book The Wisdom of the
Body captured the imagination of physiologists and psychologists alike.
The maintenance of equilibrium was accepted by some as the unifying
principle of motivation and, indeed, as a model for many other aspects
of behavior.
We believe that the following quotation from Davis (1958) puts the
concept of homeostasis in better perspective: "Homeostasis exists, of
SOME BASIC PRINCIPLES OF PSYCHOPHYSIOLOGY 59

course, with respect to certain variables, and one should try to find out
what those are. But it is to be understood as a special case of the more
general conception of the response of systems to inputs. There is no com-
pulsion to think that the organism is an elaborate machine for the pur-
pose of getting itself back to the status quo ante, or, indeed, for any other
purpose" (p. 13).
Here is a practical example why the psychophysiologist should be
aware of the possible complicating effects of homeostatic inputs. Stern
(1976) studied heart-rate responses between the GET SET and GO of a
race. In the initial design, subjects were instructed to get down on their
hands in a typical sprinter's starting position, just prior to the GET SET.
But the homeostatic response of the cardiovascular system to this pos-
tural change was a dramatic slowing of the heart. This effect was so
great and long-lasting that it was not possible to determine the effect on
heart rate of receiving the GET SET and waiting for the GO—the variable
of interest in the study. To resolve this problem, in subsequent experi-
ments subjects began from a standing position.
Autonomic Balance
As mentioned in chapter 2, most internal organs, such as the heart, are
innervated by both branches of the autonomic nervous system: the sym-
pathetic nervous system (SNS) and the parasympathetic nervous system
(PNS). The rate at which the heart beats is determined by the relative
excitation from the SNS and PNS, or the autonomic balance. Autonomic
balance may, therefore, be considered one specific part of homeostasis.
Eppinger and Hess (1915) were the first to classify people as vagotonics
or sympathicotonics. Vagotonics (from vagus nerve, the primary parasym-
pathetic nerve) are individuals who show unusually large responses to
drugs that stimulate the PNS. Sympathicotonics are persons who show
unusually large responses to drugs that stimulate the SNS. The interested
reader is referred to early work on autonomic balance by Gellhorn, Cor-
tell, and Feldman (1941) and Darrow (1943).
Wenger (1972) developed a technique for comparing an individual's
resting scores on a group of ANS measures with the scores of other
individuals and, in so doing, came up with an estimate of autonomic
balance for each subject. Each score, called A, falls somewhere along a
continuum, with low numbers indicating SNS dominance and high val-
ues indicating PNS dominance. For a given group of individuals, Wenger
found that the scores are normally distributed; that is, few people—va-
gotonics and sympathicotonics—obtain extreme scores, and most fall in
the middle. Wenger studied autonomic balance in children, college stu-
dents, military personnel, and hospitalized groups. Interestingly, Wenger
and his associates reported a higher incidence of psychosomatic, psy-
chotic, neurotic, and physical disorders in people with low A scores.
60 GENERAL ELEMENTS OF PSYCHOPHYSIOLOGY

Modes of Autonomic Control
A more recent theoretical model of autonomic function (Berntson, Ca-
cioppo, & Quigley, 1991, 1993) suggests that the sympathetic and par-
asympathetic nervous systems do not always act in an antagonistic or
reciprocal fashion—that is, autonomic activity does not always fall some-
where along a continuum that extends from sympathetic to parasym-
pathetic dominance. The major thesis of this model is that the two divi-
sions can act independently, reciprocally, or even coactively (i.e., increase
or decrease together). Furthermore, measures such as heart rate, which
are influenced by both ANS divisions, provide little information about the
specific mode of autonomic control underlying a given response. For in-
stance, an increase in heart rate may be due to one of several possible
patterns of autonomic activation: decreased parasympathetic activity, in-
creased sympathetic activity, or even increased or decreased activity in
both branches. In order to summarize all of the possible modes, Berntson
et al. (1993, p. 297) advanced the bivariate representation of autonomic
control shown in figure 5.3. The figure shows relative units of parasym-
pathetic and sympathetic activity along the lower left and right axes of
the model, respectively. The bidirectional arrows represent the possible
modes of autonomic control over organs like the heart which receive
both sympathetic and parasympathetic input. Arrows on the upper left
and right axes represent the independent (uncorrelated) activities of the
two autonomic divisions. The vertical arrow in the middle of the diagram
represents a mode of coactivation, where activation of the two divisions
may increase or decrease simultaneously. The intersecting horizontal ar-
row represents a reciprocal mode of control, where increased activity in
one branch is accompanied by decreased activity in the other branch.
The mode of autonomic control that underlies a particular response
is often determined by the selective pharmacological blockade of sym-
pathetic or parasympathetic activity. In humans, this is particularly dif-
ficult, so many of the blockade studies that lend support to the model
have been performed in nonhuman animals. For instance, Quigley and
Berntson (1990) showed that heart rate deceleration to an orienting
stimulus is no longer observed after parasympathetic blockade in the rat.
This finding suggests that heart rate deceleration during an orienting
response is mediated by increased parasympathetic activity; however,
Quigley and Berntson also observed a notable acceleration to novel stim-
uli after parasympathetic blockade. This unexpected finding suggested
that an orienting response may be mediated by an increase in the activity
level of both autonomic divisions (i.e., coactivation), but the increase in
parasympathetic activity—which is abolished after blockade—has a more
powerful effect on the heart than does increased sympathetic activity. In
this case, the increase in parasympathetic activity is said to mask the
effects of the sympathetic division.
SOME BASIC P R I N C I P L E S OF P S Y C H O P H Y S I O L O G Y 61
Autonomic Space
Figure 5.3. Possible modes of autonomic nervous system control. Reproduced

with permission from G. G. Berntson, J. T. Cacioppo, & K. S. Quigley, 1993;
"Cardiac psychophysiology and autonomic space in humans: Empirical per-
spectives and conceptual implications." Psychological Bulletin, 114, 296-322.
Noninvasive estimates of sympathetic (e.g., cardiac pre-ejection period)

and parasympathetic (e.g., respiratory sinus arrhythmia) influences on
cardiac activity have been well validated and are becoming increasingly
employed in human research. We discuss these measures further in chap-
ter 12.
Law of Initial Values
In chapter 4, which dealt with tonic versus phasic measures of ANS

activity, the law of initial values was briefly mentioned. In this section, we
cover the relationship of the size of a response to the prestimulus level
in greater detail.
Wilder (1967) was the first to call attention to this relationship and
to name it the law of initial values (LIV). He stated that any increasing
function would be smaller if the prestimulus level were higher and larger
if the prestimulus level were lower. We would feel more comfortable with
this formulation if Wilder had called it something like the principle of
initial values. It is a principle that is often supported, but it does not hold
at all prestimulus levels, for all subjects, or for all psychophysiological
measures.
Before discussing when and where this principle does seem to hold,
we must clarify what it conveys. Figure 5.4 shows fictitious data for the
heart-rate (HR) response (increase in HR) to some stimulus as a function
of prestimulus HR. Each data point represents the response of a different
subject. This figure indicates that for these data, the magnitude of the
increase in HR was strongly related to the prestimulus level. As would
be predicted by the LIV, the greater the prestimulus level, the smaller the
response to stimulation.
What physiological explanation is there for the LIV? Most psycho-
physiologists believe that homeostasis (negative feedback) is responsible
for the LIV. If we think of the various feedback mechanisms that set the
functional limits for HR and other variables, then a subject whose pre-
Figure 5.4. Heart rate response to a stimulus as a function of prestimulus

heart rate. Each dot represents the heart rate for one fictitious participant.
SOME BASIC P R I N C I P L E S OF P S Y C H O P H Y S I O L O G Y 63
stimulus level was 100 (subject B) was closer to the limit than a subject
whose prestimulus level was 50 (subject A). Looking again at the func-
tion depicted in figure 5.4, which subject would be expected to show a
greater increase in HR, A or B? We are now at the crux of the problem.
Of course, subject A would be expected to show a greater absolute in-
crease. But what about subject B? Shouldn't we inflate that subjects score,
because stronger homeostatic mechanisms would have to be overcome
due to a higher prestimulus level near the HR limit? Probably! Our an-
swer would be a definite "yes" if there were sound evidence that it was
more difficult—for example, took more metabolic energy—for subject B
to go from an HR of 100 to 105 than for subject A to go from 50 to 65.
Most psychophysiologists assume that this is the case, and the methods
for dealing with the LIV discussed below are based on this assumption.
If we examine the evidence for the LIV from empirical studies, forget-
ting for the moment the previously mentioned assumption concerning
the physiological basis, we find the following. For HR, most investigators
have found that their results support the LIV. Most psychophysiologists
have also found that skin resistance follows the LIV, but skin conductance
usually does not. A few additional studies have found support for the LIV
with blood pressure and respiration.
How would one know if a data set supported the LIV? An investigator
could either calculate the correlation between the prestimulus levels and
the magnitude of the response to stimulation, or could construct a scat-
tergram such as figure 5.4. If a significant correlation does exist, how
might we remove the effect of this relationship in further analyses?
Two methods are available for statistically neutralizing the LIV. La-
cey's (1956) Autonomic Lability Score (ALS) is obtained from the following
formula for each subject separately for each measure:
In this formula, Xz represents the individual's prestimulus level and Yz

represents a poststimulus level, expressed in units of standard deviations
from the total sample; rxy is the correlation for the sample between pres-
timulus and poststimulus levels. The constants 10 and 50 transform the
resulting scores to a distribution with a mean of 50 and a standard de-
viation of 10. In other words, the ALS is a measure of the relative mag-
nitude of a given subject's response compared to that which would be
predicted from the linear regression between variables X and Y for the
entire group. A score of 50 would indicate an average size response, 60
would indicate 1 standard deviation above the mean, and so on.
The second statistical method used to minimize the LIV is analysis of
covariance (Benjamin, 1967). We will not describe in this book how to
perform an analysis of covariance, but we will only make some general
comments. Lacey's ALS is really a special case of the analysis of covari-
ance. In the former, each individual's score is adjusted based on the
correlation between group prestimulus and poststimulus scores. In the
latter case, the group data are adjusted in much the same way so that
the poststimulus scores of two groups that differed at prestimulus levels
can be compared.
Comparing the size of responses of two subjects or two groups is very
difficult. Lykken (1968) has suggested that we get a range of possible
scores on each response measure of interest for each subject, then present
the stimulus, and assign a score to each subject based on her/his mini-
mum and maximum scores. Another possibility is to preselect subjects
for two (or more) groups so that pairs of subjects have similar prestimulus
levels.
This matter is so complex that when trying to write about it, we feel
the way Cannon must have felt when Wilder wrote to him about the
LIV. Cannon wrote back to Wilder that they were both too old to un-
dertake the task. Cannon then was 68 years old, and Wilder was 44
(Wilder, 1967). Berntson et al. (1993), however, have recently attempted
to relate the LIV to the model of autonomic control just described.
Stimulus-Response Specificity and

Individual Response Stereotypy
Stimulus-Response Specificity
At the beginning of this chapter, we mentioned the concept of stimulus-
response specificity, which makes a unidimensional activation continuum
indefensible. The soldier standing guard duty at night who suddenly
heard a noise displayed a specific pattern of responding which has been
called directional fractionation. The soldier showed cortical arousal and
an increase in skin conductance but a decrease in heart rate. Directional
fractionation is a special case of stimulus-response specificity. What is
germane to this discussion is that most other soldiers would have shown
a similar pattern of responding in that stimulus situation. By definition,
stimulus-response specificity exists if a stimulus brings about a similar
pattern of physiological responding among most subjects.
Some investigators sought evidence of stimulus-response specificity to
support William James's theory of emotion. James said that the percep-
tion of bodily changes is what constitutes emotion. Ax (1953) created a
laboratory situation in which one group of subjects was made angry and
another fearful. He recorded a large number of bodily responses and
found that about half of them differentiated between the fear and anger
situations. Davis (1957) sought response patterns to some simple stimuli
with no notion of uncovering the physiological correlates of fear, repul-
sion, and the like. The various bodily responses recorded showed signif-
S O M E BASIC P R I N C I P L E S OF P S Y C H O P H Y S I O L O G Y 65
leant differentiation among the four stimulus situations: paced key press-
ing, listening to noises, looking at pictures, and receiving cutaneous
stimulation. More recently, Stern and Koch (1996) showed that those
subjects who report symptoms of motion sickness during illusory self-
motion show a characteristic pattern of decreased respiratory sinus ar-
rhythmia amplitude coupled with increased heart rate and gastric tach-
y arrhythmia.
Individual Response Stereotypy

Individual response stereotypy refers to idiosyncratic responding. Will psy-
chiatric patients who frequently complain of head and neck pain show a
different pattern of bodily responses in a stress situation than patients
who frequently complain of heart palpitation? That was the basic ques-
tion asked by Malmo and Shagass (1949) in the first of a series of studies
on what they came to call symptom specificity. They recorded heart rate
changes, muscle potentials from the neck, and other physiological mea-
sures from both groups of patients. The exciting finding was that even
when the stress consisted of only moderate thermal stimulation, and the
subjects were not reporting that they were in pain, the head and neck
complainers showed a significant increase in muscle potential from the
neck, while the group that normally complained of palpitations showed
a significant change in their heart rate.
Lacey and his co-workers conducted several studies during the 1950s
(e.g., Lacey and Lacey, 1958) to see if the principle of symptom specificity
would hold for nonpsychiatric patients. Using various groups of subjects
and several different stressors, they found that individuals tend to respond
by showing the greatest degree of activity in the same physiological sys-
tem, no matter what the stress. For many subjects, their pattern of phys-
iological responding was repeated from stressor to stressor. This is what
we mean by individual response stereotypy.
Roessler and Engel (1977) made the point that stimulus-response
specificity and individual response stereotypy are not mutually exclusive.
For example, in the Davis (1957) study previously mentioned, the male
subjects were shown a slide of a nude woman. Most, but not all, of the
subjects displayed what Davis called the P pattern: increased skin con-
ductance, peripheral vasoconstriction, and heart rate slowing. This would
be an example of stimulus-response specificity. The fact that a few sub-
jects responded differently, perhaps with a heart rate increase, may well
be due to their idiosyncratic cardiac responding—another example of
individual response stereotypy.
Stimulus-response specificity and individual response stereotypy prob-
ably exist to some degree in all psychophysiological studies. The practical
question is, how serious is this problem, and what can and should be
done about it? We cannot make a blanket statement concerning the pro-
portion of the total variance in all future studies which will be attribut-
able to stimulus-response specificity and individual response stereotypy,
as it will no doubt vary considerably. However, the problem does not
appear serious if careful consideration is given to the selection of physi-
ological responses to be measured and the number of subjects. Stimulus-
response specificity tells us that we should record not just one but several
physiological measures and examine the pattern of responses to our var-
ious stimulus situations. Individual response stereotypy, on the other
hand, alerts us to the problem of a few subjects making idiosyncratic
responses in a situation where quite a different pattern of responses might
be expected.
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6
Safety and Ethics in a
Psychophysiology Laboratory
Safety
While the laboratory electrical environment is probably at least as safe

as the modern home and ordinarily poses little threat of shock, there are
important implications for the treatment of the subject, particularly con-
cerning grounding.
It is standard procedure in some labs to attach a ground lead to the
subject when recording psychophysiological measures. The reason for the
ground connection in these cases is not as protection for the subject, but
rather to minimize unwanted electrical signals. If left "floating," or un-
grounded, the subject acts as an antenna, picking up unwanted voltages
from the air, much as a radio antenna picks up radio signals. These
voltage variations, particularly 60 Hz, are then amplified and appear on
the record, obscuring the desired biological signal. However, a ground
lead may expose the subject to electrical hazard. If the subject is con-
nected to ground through a low-impedance lead, then any device not at
ground potential which touches the subject or which the subject touches,
such as a lamp with an old two-prong plug and a short circuit, will cause
a current to flow through the subject to ground. Furthermore, subjects
seldom need to be grounded today to obtain artifact-free recordings. Mod-
ern differential amplifiers provide sufficient common mode rejection to
reject, at the amplifier input, those transient signals the subject ground
was meant to eliminate. Some older polygraphs, however, specify that a
ground lead be attached for certain types of psychophysiological record-
ing. We suggest that the experimenter carefully read the user manual
for the instrument before deciding to use a subject ground.
If a piece of equipment is moved for repair and its ground is not re-
connected properly, the case of that unit becomes a potential hazard.
Therefore, ground connections should be routinely checked. If the labo-
70
ratory is moved, it is important to check in the new facility that the third
prong in wall sockets is, indeed, attached to the building ground. It
quickly becomes obvious if your recorder is not receiving power, but it
will not be obvious if it loses its ground connection.
Additional Safety Principles
Plans for Medical Emergency

There is always the possibility that a medical emergency will occur while
a subject is under your care. The emergency may be related to the re-
cording you are doing or independent of it. In any event, there should
be a prepared course of action. The plan will vary depending on the
location of laboratory. In a hospital, qualified persons and proper emer-
gency equipment, as well as established emergency procedures, already
exist. However, in an academic setting, emergency procedures are usually
not so clear, but every laboratory must have a plan of action in case of
an emergency. It is imperative that there be a first aid kit and a phone
in the laboratory next to which there is a list of numbers to call in case
of an emergency. We also recommend that laboratory personnel receive
training in basic first aid and, if possible, cardiopulmonary resuscitation
(CPR).
Unusual Recordings
While recording, you may observe an EKG, EEC, or other psychophysi-
ological response that appears abnormal. The EKG signal may be irreg-
ular, or missing a component. EEC records may appear to demonstrate
"spike and dome" waves indicative of medical problems. The student
psychophysiologist should not mention such apparent abnormalities to
the subject. In most cases, the unusual recording is not indicative of a
medical problem. Any suggestion to the subject that there may be some-
thing wrong with the subject's heart, brain, or other organ may seriously
worry the subject for no reason. There are many reasons for unusual
recordings; interpretation should be made by an expert. Such recordings
should be shown to an instructor or other person responsible for the
laboratory. This individual will make the interpretation, after consulting
with a medical doctor if deemed necessary, and advise the subject when
appropriate.
The Use of Electric Shock on

Human Subjects
It was formerly a common procedure to use electric shock as a noxious
or arousing stimulus while recording psychophysiological data. More re-
SAFETY & ETHICS IN P S Y C H O P H Y S I O L O G Y L A B O R A T O R Y 71

cently, electric shock has been less used, both because it is ethically ques-
tionable and because of the danger of serious injury. While a complete
account of the potential circuits created through the subject by the in-
troduction of shock is beyond the scope of this chapter, we feel that the
dangers inherent in the use of electric shock are great enough to advise
against its use where bioelectric potentials are being recorded simulta-
neously. Indeed, many of the "shockers" currently in the storerooms of
psychology departments could not pass any reasonable test for electrical
safety. Certainly any device that involves a 117 V AC supply—that is,
one that plugs into a wall outlet—should not be used. Battery-powered
shock generators are less dangerous, but they should be used only under
direct supervision and with the greatest care. Electrodes, for example,
should be placed close to one another and far from vital organs. Currents
should never be passed through the body.
Ethical Considerations
Before recording begins, the psychophysiologist must consider ethical

questions related to his procedure. Does the procedure involve physical
stress or even the threat of it? Could the recording result in anxiety,
shame, guilt, or embarrassment? Is the subject's physical safety assured?
Will all promises made to the subject be fulfilled? Is the subject to be fully
informed about the procedure? Will information about the subject be
confidential? Is coercion involved? The list of questions can go on and
on, but the psychophysiologist must raise these questions and answer
them.
While there may be a desire to absolve oneself of ethical responsibility
by turning it over to others—the instructor, the "use of human subjects
committee" (often referred to as institutional review board or IRB), or
even the subject—responsibility must remain with the experimenter.
What is more, psychophysiological recording, by its very nature raises
ethical questions. For example, in EGG recording, should male experi-
menters apply electrodes to the skin over the abdomen of female subjects?
We recommend gender matching when electrodes are applied so as to
reduce any embarrassment or anxiety on the part of the subject. Some
use of human subjects committees now require this. Ethical issues are
also raised when, for example, recording blood pressure, the electroen-
cephalogram, or electrodermal activity provides the experimenter or tech-
nician with information about the subject not known to others and prob-
ably not even to the subject. Also, as detailed earlier in the chapter, some
discomfort and an element of danger are necessarily involved in record-
ing.
What if the subject becomes frightened of the electrodes? Who is re-
sponsible? How did the situation come about? Is the equipment com-
pletely safe? How sure should one be of its safety before attaching it to
another human being?
Ethical questions balance the rights of the subject with the possible
benefits of the procedure. The rights of the subject include the right to
privacy, the right to know what is to occur, freedom from physical dan-
ger, and freedom from fear, anger, embarrassment, or other behavioral
disturbance. Benefits may include benefits to the experimenter (who may
be learning a technique), benefits to the subject (in the case of an EMG
relaxation procedure, for example), and benefits to society through an
advance in scientific knowledge resulting from the experiment.
How can one evaluate the dangers inherent in doing psychophysio-
logical research, and how can they be balanced with the possible benefits
of recording? In the final analysis, each psychophysiologist must make
his/her own ethical judgment. Scientists whose research involves human
subjects have studied the ethics of working with human beings and have
devised principles that will guide the psychophysiologist. The American
Psychological Association (1992) has published an article entitled "Eth-
ical Principles of Psychologists and Code of Conduct." The article deals
with ethical standards for all aspects of the varied work of psychologists,
but only the standards that relate directly to research are listed here.
Where interpretation of the principles is not clear, the psychophysiologist
should refer to the complete article, where the standards are presented
in greater detail.
The Ethical Standards

6.06 Planning Research.
(a) Psychologists design, conduct, and report research in accordance

with recognized standards of scientific competence and ethical re-
search.
(b) Psychologists plan their research so as to minimize the possibility
that results will be misleading.
(c) In planning research, psychologists consider its ethical acceptability
under the Ethics Code. If an ethical issue is unclear, psychologists
seek to resolve the issue through consultation with institutional
review boards, animal care and use committees, peer consultations,
or other proper mechanisms.
(d) Psychologists take reasonable steps to implement appropriate pro-
tections for the rights and welfare of human participants, other
persons affected by the research, and the welfare of animal subjects.
6.07 Responsibility.
(a) Psychologists conduct research competently and with due concern

for the dignity and welfare of the participants.
S A F E T Y & E T H I C S IN P S Y C H O P H Y S I O L O G Y L A B O R A T O R Y 73
(b) Psychologists are responsible for the ethical conduct of research
conducted by them or by others under their supervision or control.
(c) Researchers and assistants are permitted to perform only those
tasks for which they are appropriately trained and prepared.
(d) As part of the process of development and implementation of re-
search projects, psychologists consult those with expertise concern-
ing any special population under investigation or most likely to be
affected.
6.08 Compliance With Law and Standards.
Psychologists plan and conduct research in a manner consistent with

federal and state law and regulations, as well as professional standards
governing the conduct of research, and particularly those standards
governing research with human participants and animal subjects.
6.09 Institutional Approval.
Psychologists obtain from host institutions or organizations appropriate

approval prior to conducting research, and they provide accurate in-
formation about their research proposals. They conduct the research
in accordance with the approved research protocol.
6.10 Research Responsibilities.
Prior to conducting research (except research involving only anony-

mous surveys, naturalistic observations, or similar research), psychol-
ogists enter into an agreement with participants that clarifies the na-
ture of the research and the responsibilities of each party.
6.11 Informed Consent to Research.
(a) Psychologists use language that is reasonably understandable to

research participants in obtaining their appropriate informed con-
sent (except as provided in Standard 6.12, Dispensing with In-
formed Consent). Such informed consent is appropriately docu-
mented.
(b) Using language that is reasonably understandable to participants,
psychologists inform participants of the nature of the research; they
inform participants that they are free to participate or to decline to
participate or to withdraw from the research; they explain the fore-
seeable consequences of declining or withdrawing; they inform par-
ticipants of significant factors that may be expected to influence
their willingness to participate (such as risks, discomfort, adverse
effects, or limitations on confidentiality, except as provided in Stan-
dard 6.15, Deception in Research); and they explain other aspects
about which the prospective participants inquire.
(c) When psychologists conduct research with individuals such as stu-
dents or subordinates, psychologists take special care to protect the
prospective participants from adverse consequences of declining or
withdrawing from participation.
(d) When research participation is a course requirement or opportu-
nity for extra credit, the prospective participant is given the choice
of equitable alternative activities.
(e) For persons who are legally incapable of giving informed consent,
psychologists nevertheless (1) provide an appropriate explanation,
(2) obtain the participant's assent, and (3) obtain appropriate per-
mission from a legally authorized person, if such substitute consent
is permitted by law.
6.12 Dispensing with Informed Consent.
Before determining that planned research (such as research involving

only anonymous questionnaires, naturalistic observations, or certain
kinds of archival research) does not require the informed consent of
research participants, psychologists consider applicable regulations and
institutional review board requirements, and they consult with col-
leagues as appropriate.
6.13 Informed Consent in Research Filming or Recording.
Psychologists obtain informed consent from research participants prior

to filming or recording them in any form, unless the research involves
simply naturalistic observations in public places and it is not antici-
pated that the recording will be used in a manner that could cause
personal identification or harm.
6.14 Offering Inducements for Research Participants.
(a) In offering professional services as an inducement to obtain re-

search participants, psychologists make clear the nature of the
services, as well as the risks, obligations, and limitations.
(b) Psychologists do not offer excessive or inappropriate financial or
other inducements to obtain research participants, particularly
when it might tend to coerce participation.
6.15 Deception in Research.
(a) Psychologists do not conduct a study involving deception unless

they have determined that the use of deceptive techniques is jus-
tified by the study's prospective scientific, educational, or applied
value and that equally effective alternative procedures that do not
use deception are not feasible.
S A F E T Y & E T H I C S IN P S Y C H O P H Y S I O L O G Y L A B O R A T O R Y 75
(b) Psychologists never deceive research participants about significant
aspects that would affect their willingness to participate, such as
physical risks, discomfort, or unpleasant emotional experiences.
(c) Any other deception that is an integral feature of the design and
conduct of an experiment must be explained to participants as early
as is feasible, preferably at the conclusion of their participation, but
no later than at the conclusion of the research. (See also Standard
6.18, Providing Participants with Information about the Study.)
6.16 Sharing and Utilizing Data.
Psychologists inform research participants of their anticipated sharing

or further use of personally identifiable research data and of the pos-
sibility of unanticipated future uses.
6.17 Minimizing Invasiveness.
In conducting research, psychologists interfere with the participants or

milieu from which data are collected only in a manner that is war-
ranted by an appropriate research design and that is consistent with
psychologists' roles as scientific investigators.
6.18 Providing Participants with Information about the Study.
(a) Psychologists provide a prompt opportunity for participants to ob-

tain appropriate information about the nature, results, and conclu-
sions of the research, and psychologists attempt to correct any mis-
conceptions that participants may have.
(b) If scientific or humane values justify delaying or withholding this
information, psychologists take reasonable measures to reduce the
risk of harm.
6.19 Honoring Commitments.
Psychologists take reasonable measures to honor all commitments they

have made to research participants.
References
American Psychological Association (1992). Ethical principles of psycholo-
gists and code of conduct. American Psychologist, 47,1597-1628. (Also
available at www.APA.org). Copyright © 1992 by the American Psy-
chological Association. Reprinted with permission.
PART II
Psychophysiology of Specific
Organs and Systems
7
Brain
Electroencephalogmphy and Imaging
The presence of recognizable electrical rhythms of the brain has excited

the curiosity and imagination of both professionals and laypeople alike.
Psychophysiologists, neurologists, and science fiction writers have been
intrigued by the presence of brain activity and the possibility of having
an objective noninvasive marker that reflects underlying psychological
processes. Some have taken these ideas to the extreme by suggesting that
these measures can tell us what someone is thinking or feeling or even
if they are telling the truth. Although it is not that simple, some re-
searchers have begun to use brain activity to help physically disabled
individuals to communicate, to move paralyzed limbs, or even to reduce
seizure disorders. As discussed in this chapter, the understanding of cor-
tical processes through various forms of brain imaging is a complex task.
Scientists have come to appreciate the complicated relationship that exists
between electrocortical measures and cognitive, emotional, and behav-
ioral processes. There is a variety of techniques available to help us un-
derstand the functioning of the brain. We begin with the measurement
of electrical activity recorded from the scalp, electroencephalography
(EEC).
Spontaneous EEGs
Electroencephalography is a technique for recording electrical activity

from the scalp related to cortical activity. The EEC in humans was first
described by Hans Berger in 1929. In his first set of papers, Berger sought
to determine what activities were related to the EEC. As a good scientist,
he first determined that EEC was actually related to brain activity and
not other physiological activity. In order to do this he showed that EEC
was not related to cerebral pulsations, cerebral blood flow, blood flow
79
through scalp vessels, heart rate activity, muscle activity, eye movements
or electrical properties of the skin. Berger took his studies beyond the
physiological level and was one of the first to suggest that periodic fluc-
tuations of the EEC might be related in humans to cognitive processes
such as arousal, memory, and consciousness. Today, we carry on this
work by using EEC measures to help us understand such processes as
sleep, attention, emotion, and preparation for movement.
To record an EEC, electrical signals of only a few microvolts must be
detected on the scalp. This can be accomplished by amplifying the differ-
ential between two electrodes, at least one of which is placed on the scalp.
Since the signal must be amplified almost one million times, care must
be taken that the resulting signal is indeed actual EEC and not artifact.
Later in the chapter, we will discuss some possible artifacts, but for now
let us examine the EEC. The rhythmic variations of the EEC are contin-
ually present at the surface of the scalp from well before birth to death.
In fact, the absence of the EEC for 24 hr has been used as an indicator
of "brain death." The various frequencies and distributions of specific
patterns of the EEC wax and wane, providing the brain researcher and
clinician with a constant record of the changing patterns of electrical
activity of the brain. Some aspects of the EEC may appear almost random
while other fluctuations appear periodic. We have a variety of signal-
processing techniques to help us describe the EEC, but in general we use
two basic parameters: amplitude (how large the signal is) and frequency
(how fast the signal cycles). Some EEC patterns are extremely reliable
and can be visually observed, as would have been required in the days
before computer analysis. These patterns have been identified by Greek
letters such as a (alpha), (3 (beta), and 0 (theta).
Alpha activity can be seen in about three-fourths of all individuals
when they are awake and relaxed. Asking these individuals to relax and
close their eyes will result in recurring periods of several seconds in which
the EEC consists of relatively large, rhythmic waves of about 8-12 Hz.
This is the alpha rhythm, the presence of which has been related to re-
laxation and the lack of active cognitive processes. If someone who dis-
plays alpha activity is asked to perform cognitive activity such as solving
an arithmetic problem in their head, alpha activity will no longer be
present in the EEC. This is referred to as alpha blocking. Typically, with
cognitive activity the alpha rhythm is replaced by high-frequency, low-
amplitude EEC activity referred to as beta activity.
Beta activity occurs when one is alert. Traditionally, lower-voltage var-
iations ranging from about 18 to 30 Hz have been referred to as beta
and higher frequency lower-voltage variations ranging from about 30 to
70 Hz or higher as gamma. Recent work suggests that gamma activity*
is related to the brain's ability to integrate a variety of stimuli into a
coherent whole. For example, Catherine Tallon-Baudry and her col-
leagues (1997) showed individuals pictures of a hidden Dalmatian dog
80 P S Y C H O P H Y S I O L O G Y OF S P E C I F I C O R G A N S AND S Y S T E M S
which was difficult to see because of the black and white background.
After training individuals to see the dog, there were differences in the
gamma band response suggesting differential responses to meaningful
versus nonmeaningful stimuli.
Additional patterns of spontaneous EEC activity include delta activity
(0.5-4 Hz), theta activity (5-7 Hz), and lambda and K-complex waves
and sleep spindles, which are not defined solely in terms of frequency.
Theta activity refers to EEC activity in the 4-8 Hz range. Walter (1953),
who introduced the term theta rhythm, suggested that theta was seen
at the cessation of a pleasurable activity. More recent research has theta
associated with such processes as hypnagogic imagery, REM (rapid eye
movement) sleep, problem solving, attention, and hypnosis. In a review
of theta activity, Schacter (1977) suggested that there are actually two
different types of theta activity. First there is theta activity associated
with low levels of alertness as would be seen as one falls asleep. Second,
there is theta activity associated with attention and active and efficient
processing of cognitive and perceptual tasks. This is consistent with the
suggestion of Vogel, Broverman, and Klaiber (1968) that there are two
types of behavioral inhibition, one associated with a gross inactivation
of an entire excitatory process resulting in less active behavioral states
and one associated with selective inactivity as seen in overlearned pro-
cesses.
Delta activity is low frequency (0.5-4 Hz) associated with sleep in
healthy humans as well as with pathological conditions such as brain
tumors. This is also the predominant frequency of human infants during
the first two years of life. Figure 7.1 displays some commonly observed
EEC waveforms.
Figure 7.1. An EEC from an adult with electrodes at various sites on the
scalp.
BRAIN 81
Recording Procedure
Although procedures for recording EEC activity have improved greatly
over the past 20 years with the incorporation of computer-controlled and
digital amplifiers, one must still carefully consider how EEC is recorded
and the possible artifacts that can compromise the data. Because EEC
voltages are minuscule (several millionths of a volt) and thus must be
amplified by a factor of a million, the possibility of recording electrical
interference looms at every stage of instrumentation from the electrodes
through to the recorder. These potentials can easily be confused with the
legitimate EEC. Specific problems will be described in the appropriate sec-
tions of this chapter.
Electrodes and Electrode Placement Where the electrodes are placed and
how many are used depend on the purpose of the recording. Today,
almost all EEC procedures use a variety of EEC helmets with up to 2 5 6
electrodes built into the helmet, although it is also still possible to record
EEC from only two electrodes. Those recording helmets that use 128 to
256 electrodes are generally referred to as dense array EEC recordings.
If the spatial distribution of some aspect of the EEC is the research ques-
tion, then multiple electrodes distributed over the scalp are required. Of
course, one can record from many fewer electrodes depending upon the
empirical questions that are being asked. For example, if one is only
interested in EEC responses associated with movement, then one may
chose to record from regions of the scalp lying above the motor areas of
the brain.
Historically, the system of locating electrodes in EEC is referred to as
the International 10-20 system shown in figure 7.2 (Jasper, 1958). The
name 10-20 refers to the fact that electrodes in this system are placed
at sites 10% and 20% from four anatomical landmarks. In the front the
nasion (the bridge of the nose) is used. In the rear of the head, the inion
(the bump at the back of the head just above the neck) is used. The left
and right landmarks are the preauricular points (depressions in front of
the ears above the cheekbone). In this system, the letters refer to areas
of the brain; 0 = occipital, P = parietal, C = central, F = frontal, and
T = temporal. Numerical subscripts indicate laterality (odd numbers left,
even right) and degree of displacement from the midline (subscripted z).
Thus, C3 describes an electrode over the central region of the brain on
the left side, whereas Cz would refer to an electrode placed at the top of
the scalp above the central area. These lead locations are simply conven-
tional and you may see in the literature nonstandard electrode location
that are used in order to examine a particular research question.
Two specific types of EEC recording are called monopolar and bipolar
recordings. In order to understand this point we must remember that
Figure 7.2. The international 10-20 electrode system.
EEC recordings reflect the difference in voltage between a signal at two
electrodes. This means that if the exact cortical signal were present at
two separate sites on which our electrodes were placed then we would
record a straight line reflecting no difference in activity between the two
sites. Of course, this never happens because there are always differences
in activity between recording sites. In monopolar recordings the idea is
to find a site that is not reflective of EEC activity per se to use as a
reference site. Common sites used for this purpose are the ear (or ears),
the mastoid, or even the nose. Both ears can be used as a reference site
by connecting clip electrodes to each ear and then connecting these to-
gether as if they were one electrode. This technique is referred to as
"linked ears." Thus, EEC activity from the various sites on the scalp could
be referenced to the ear (or ears) to give the difference between each site
and the reference electrode. Other researchers have suggested that a use-
ful reference to use is that of the average reference. This procedure ba-
sically takes a network of electrodes spaced across the scalp and math-
ematically averages these together. This mathematical average value is
then used as the reference.
In bipolar recording, each electrode is located to record from an active
site on the scalp. Thus, one could compare the difference in EEC activity
between the right frontal area with that of the left frontal area. One might
use such a procedure to infer whether, for example, the left or right
hemisphere was more involved in a particular task. This type of proce-
dure has traditionally been used in clinical settings to identify unusual
pathological waveforms, such as epileptic discharges.
Recording Equipment Several types of signal-conditioning and amplifica-

tion equipment may be used. Large hospitals and research laboratories
employ multiple-channel electroencephalographs designed expressly for
EEC recording. Many research labs also utilize a paperless EEC system
which is controlled, displayed, and stored by the computer such that
paper recording is not required. Psychophysiological laboratories may
have one or more multipurpose polygraphs with signal-conditioning
equipment that permits EEC recording. The amplifier used must have
high gain capabilities and must be able to reproduce accurately the EEC
signal at all desired frequencies. The amplifier must be able to amplify
waveforms from almost DC (0 Hz) to more than 100 Hz. Typical ampli-
fication is a million or more times with little distortion. The amplifier
must have an input impedance of several million ohms to prevent atten-
uation of the EEC signal. Also, the amplifier must have high common-
mode rejection to reject ground-referred interference signals, such as 60-
Hz interference from lights. Some systems may include a 60-Hz notch
filter, which increases the ability to exclude interference of that frequency
from the record, although distortion of waveforms at frequencies near 60
Hz may be introduced. It is helpful to have signal-conditioning filters that
84 P S Y C H O P H Y S I O L O G Y OF S P E C I F I C ORGANS AND SYSTEMS

are used to emphasize a chosen range of EEC frequencies and to minimize
others. Most commercial EEC recording systems include both high-pass
and low-pass filters so that a "frequency window" can be constructed by
setting the filters selectively to eliminate frequencies above and below
those of interest. An important question we discuss in other chapters (see
chapters 3 and 14) is the nature of a filter and what is referred to as roll
off or sharpness. This simply asks the question of how does a 60-Hz filter,
for example, affect nearby frequencies at 59 Hz and below and 61 Hz
and above. Some systems describe the filters in terms of a time constant,
which is basically the time it takes for an AC signal to fall two-thirds of
its initial amplitude. According to the formula relating time constants
and frequency, a time constant of 1 s, for example, attenuates all fre-
quencies below 0.16 Hz, whereas a time constant of 0.03 would reduce
signals below 5.3 Hz. Alternatively, a filter may be described in terms of
the frequencies that are attenuated.
Procedure As with most psychophysiological recording procedures, the

basic requirement is to ensure that the areas under the EEC electrodes
are free of hair, oils, and dead skin. If one were attaching a few electrodes,
the areas to which electrodes will be attached are rubbed with alcohol
or acetone to remove oils and dead skin. This should be done with care
and patience, for the resistance between any electrode pair must be under
5,000 fl to obtain a good recording. One may also slightly abrade the
skin using a variety of procedures. An electrode paste (usually containing
sodium chloride as the electrolyte) is rubbed into the skin at the electrode
site. A similar procedure is used with electrode helmets in which as
blunted needle is used to move the hair away from the electrode and a
syringe like device used to inject electrode paste between the electrode
and the scalp. It is imperative that the electrodes and the instruments
used to prepare the scalp be disinfected between users to avoid spreading
any pathogens. As mentioned in chapter 3, a committee of the Society
for Psychophysiological Research developed a set of guidelines for the
proper care and sterilization of EEC equipment (Putnam, Johnson, &
Roth, 1992). Because of these concerns and with the development of
dense array systems, a new technology is being developed that requires
no skin abrasion prior to the application of EEC electrodes. Dense array
systems use 128 or 256 electrodes; these new systems use a slightly
different procedure because their amplifiers do not require lowering the
impedance to 5,000 O as do the traditional systems. These systems are
simply soaked in a solution and then applied directly to the head.
Whether using these systems or the traditional methods, one must mea-
sure the quality of the connection, the impedance, between the scalp and
the electrode. This measurement enables the researcher to ensure that
all electrodes are functioning properly.
BRAIN 85
Figure 7.3. Resting EEC—alpha predominance.
Typical Recordings
EEGs recorded under a variety of conditions are shown in figures 7.3,
7.4, and 7.5. Figure 7.3 shows four recordings obtained simultaneously
from a resting subject. The four locations on the scalp from which the
tracings were made are indicated in the figure: two temporal, one central,
and one frontal. The high-amplitude waves seen in all the recordings
are typical alpha rhythms. Figure 7.4 shows four recordings from the
same scalp areas as in figure 7.3, but from an alert subject. Here we
Figure 7.4. Alert EEC—beta rhythm.
Figure 7.5. Delta waves recorded during deep sleep.
see typical low-amplitude fast waves—beta rhythms—along with some

larger spikes. Sleep research is an area that makes much use of EEGs to
determine if a subject is asleep and, if so, the stage of sleep. Figure 7.5
shows very the high-amplitude, very slow waves characteristic of delta
rhythms. A record such as this would be obtained from a deeply sleeping
subject.
Common Problems
Given that the EEC signal is amplified more than a million times, artifacts
can be a real problem in the EEC and misinterpreted as an actual signal.
BRAIN 87
For example, a student once interpreted an EEC to indicate a person was
having an epileptic seizure when in fact the subject was chewing gum.
Any type of movement can be seen in the EEC including muscle move-
ment from facial expressions and talking. Another common artifact is
related to eye movements which can interfere with the EEC signal. Today
it is standard procedure when measuring the EEC to place electrodes
around the eye to detect eye movement in any direction. These proce-
dures are described in chapter 9. A variety of algorithms is available for
removing eye movement signals from the EEC. The beating of the heart,
which occurs at about once per second, also sometimes appears in an in
a EEC record. External factors such as elevator motors or electric lights
also can produce artifacts. Most of these problems can be dealt with by
filters that reduce the unwanted frequencies. One of the best ways for
students to learn about artifacts is to learn how to produce them on
command. The presence of 60-Hz noise in the record is the most common
difficulty in EEC recording. Figure 7.6 shows a continuous EEC in which
60 Hz is present in the first section but has been removed from the second
section. Usually 60 Hz is recognized by its invariant frequency and am-
plitude. However, when it is of low amplitude, the presence of 60 Hz
interference is not always as clear as in the example. This interference is
often difficult to eliminate, in spite of the high common-mode rejection
of modern differential amplifiers and the use of 60-Hz notch filters, be-
cause the source of the interference may be hundreds of times as powerful
as the EEC signal generators themselves. If 60-Hz interference persists, it
must be dealt with, since it will either obliterate the EEC or at least
confound its interpretation. One common source of 60-Hz interference is
fluorescent light fixtures. They may simply be turned off. Also, the inter-
ference may emanate from stimulus devices such as slide projectors, tape
recorders, computer monitors, electric motors, and nearby electrical
transmission lines.
The equipment and the subject may be isolated from all electrical
interference by being enclosed in a grounded metal shield, but this is
usually not necessary with newer recording equipment. For EEC record-
Figure 7.6. 60-Hz interference in an EEC record.
88 PSYCHOPHYSIOLOGY OF SPECIFIC ORGANS AND SYSTEMS

Figure 7.7. Physiological interference in EEC records.
ing, it is necessary that the subject be muscularly relaxed; head move-

ments, jaw clenching, and frowning introduce large artifacts into the
EEC. Subjects are usually instructed to fixate visually on some target or
to close their eyes. Figure 7.7 demonstrates the effect of some of these
physiological sources of interference on the spontaneous EEC. It is useful
for the technician to be able to see the subject in order to identify better
the source of artifacts.
BRAIN 89
Analysis and Quantification
Historically, EEC technicians in clinical settings underwent extensive
training so that they could recognize the visual patterns of EEC related
to sleep stages and neurological disorders. Some frequencies, such as the
alpha rhythm, are easy to recognize while detecting the presence of other
EEC frequencies is more difficult. Since visual pattern recognition is sub-
jective, EEC researchers have sought to develop quantitative procedures
for describing EEC activity. In order to do a quantitative analysis, it is
first necessary to convert the continuous analog EEC signal into a digital
form, which is accomplished by an analog-to-digital converter. Once the
signal is represented as individual numbers in a time series, these num-
bers can be manipulated mathematically. One of the first questions that
must be determined is the sampling rate of the digital converter so that
an accurate EEC record can be obtained. Based on a variety of engineer-
ing studies, the smallest sampling rate recommended is that of twice the
highest frequency that one wishes to detect. Thus, if one wanted to study
an EEC signal between 4 and 30 Hz, one would have to record the EEC
at a sampling rate of at least 60 Hz. However, most researchers sample
at four to eight times the highest frequency under consideration to ensure
accurate detection of the EEC.
Frequency Analysis One of the most common frequency analysis tech-

niques is that of Fourier analysis. The technique is named after the French
mathematician Fourier, who suggested that any given time series can be
described as a corresponding sum of sine and cosine functions. Using this
information he described how to determine in the frequency domain the
amplitude and phase information of a known temporal signal. One simple
way of understanding this procedure is to imagine that one had a variety
of templates which represent each frequency band under consideration.
Thus, one could have an 8-Hz template, a 9-Hz template, a 10-Hz tem-
plate, and so forth. By simply placing each template on top of the signal,
you could determine how closely the signal fit that template. This is
basically the procedure that Fourier analysis uses. It takes an EEC signal
over time and describes it in term of how much of each frequency is
represented in the signal. Thus, Fourier converts a time-based signal to
a frequency-based signal. In the 1960s a mathematical algorithm, re-
ferred to as the Fast Fourier Transform (FFT), was developed to speed
computations of this procedure; FTT is used by most computer programs
today. A more recent technique for frequency analysis, especially for
short segments, is that of wavelet analysis (this is introduced in chapter
14).
An analysis technique related to Fourier analysis is that of coherence
analysis. Whereas Fourier analysis gives the frequency for a given elec-
trode, coherence gives the covariance of this measure for a pair of elec-

trodes. Thus, coherence tells you how the EEC signal at each of two
electrodes is related to each electrode. In simple terms, coherence reflects
the manner in which two signals covary at a particular frequency. That
is to say if the EEC at the right frontal electrode and the left frontal
electrode both demonstrated a frequency of 8 Hz, then we would see
greater coherence between the two electrodes than if they did not show
the same frequency. In doing the coherence analysis, one can also obtain
a measure of phase or synchrony. In other words, we can determine if
two signals of the same frequency have peaks and valleys at the same
time. Using coherence, Thatcher and his colleagues (1987, 1992) have
studied how the brains of children develop patterns of EEC activity in
different areas as they mature.
Event-Related Potentials
Evoked Responses
If a flash of light is viewed by a subject who has one electrode on the
rear of his scalp and another on his earlobe, a predictable sequence of
voltage variations will be recorded. A very small positive deflection (less
than a microvolt) will follow the flash by about 40 ms. This response will
be followed by a large negative deflection lasting 10 to 30 ms and peaking
around 60 ms after the flash. Immediately following this wave there
appears a fairly large, positive wave with a maximum amplitude occur-
ring about 80 ms after the flash. This pattern is quite predictable; it
follows each successive light flash, although, it should be stressed, with
some variability from flash to flash. This succession of waveforms is
termed the visual evoked response. When the distribution of the responses
is examined, it is found to be of maximum amplitude over the occipital
area of the brain, and to be less widely distributed than most spontaneous
rhythms.
Other sensory-evoked responses also can be demonstrated. A sharp
sound reliably produces an auditory evoked response. The response is max-
imal over the vertex of the brain and usually entirely absent from the
occipital area. It has been shown that the brain's response to discrete
sounds can be traced from the brain stem to the cortex in recordings
from an electrode on the scalp. In such records (termed brain stem evoked
responses, or BSERs), a distinct wave of positive voltage reflects each level
of neural activity as the effects of the stimulus move through the brain.
In the same manner, local stimulation of the skin surface in most body
locations results in a somatic evoked potential, the waveform and distri-
bution of which are dependent upon the area stimulated.
In general, evoked responses regardless of the nature of the stimulus
are referred to as event-related responses (ERPs). Unlike the spontaneous
BRAIN 91
EEC, which is recorded in a continuous fashion over a period of time,
ERPs are time locked to specific stimuli or responses. In the literature a
distinction is sometimes made between endogenous and exogenous ERPs.
Exogenous ERPs are seen to be controlled largely by the physical nature
of the stimulus itself. Endogenous ERPs, on the other hand, are those
that are influenced by the individual's perception or interpretation of the
event. Most of the ERP research of interest to psychophysiologists would
fall within the endogenous ERPs designation. Overall, the ERP is smaller
in voltage than the EEC and requires averaging procedures over many
trials for patterns to be clearly seen. The most common ERP procedure
is to time lock the EEC signal to a particular tone or visual stimulus. By
repeating the stimulus a number of times and averaging the electrocort-
ical signal to each of these stimuli, it is possible to see a signal as displayed
in figure 7.8. In this figure the stimulus presented at time 0 and the
resulting ERP is seen to reflect the brain's processing of the information
component of the stimulus. Traditionally ERPs are referred to in terms of
whether the deflection is negative or positive and when the deflection
occurs. Thus, a P300 component is a positive component occurring
about 300 ms after the stimulus. It should be noted that the timing of
Figure 7.8. Averaged event-related response to acoustic stimuli. Waves I to

VI comprise acoustic brain stem potentials. Components from 100 ms latency
on are considered endogenous components. Reprinted with permission from
B. Rockstroh, T. Elbert, A. Canavan, W. Lutzenberger, & N. Birbaumer, 1982,
Slow cortical potentials and behaviour, Baltimore: Urban & Schwarzenberg.
92 P S Y C H O P H Y S I O L O G Y OF S P E C I F I C O R G A N S AND SYSTEMS
the components are not precise but relative. While it is true that a P300
will follow an N200, the P300 may occur later than 300 ms. In viewing
graphs of the ERP, a general procedure is to show the negative compo-
nents as going upwards and the positive ones as downward. Unfortu-
nately, this protocol is not always followed and one should carefully
check the axes of a particular graph to determine how the ERP is plotted.
To add to the confusion, ERP components may be abbreviated so that a
N100 negative component may be referred to as Nl, or a negative de-
flection that occurs approximately 200 ms after the stimulus referred to
asN2.
In terms of time, the initial components of the ERP reflect automatic
processing with the later components being more controlled and related
to the cognitive processing of the stimulus. For example, if a pain stimulus
was delivered to your right finger, an initial response would be seen on
the left side of the cortex. At about 250 ms, an evoked response is seen
that some researchers believe to be associated with the subjective re-
sponse of pain. One of the most well known of the ERP components is
that of P 3 00, which in actuality can appear anywhere from 300 to 800
ms after the response. P300 is seen as reflecting cognitive processing and
has been used in a variety of paradigms. For example, this component is
larger if individuals are told to respond to a stimulus than if they are
instructed to ignore it. One common P300 paradigm is that of the odd-
ball. In this procedure, a series of tones with a similar frequency is played
in which a tone of a different frequency is played randomly. The novel
stimulus or "oddball" results in an increase in the amplitude of the P300.
A related component involved with linguistic processing is that of the
N400. This component seems to be especially related to linguistic expec-
tation. For example, if you were to hear "Mary had a little ..." you
would probably expect the word "lamb" to come next. However, if you
heard "Mary had a little pizza" then you would see an increase in the
N400 component of an ERP.
Slow Potentials
If you were told that after you heard a tone a picture would follow a few
seconds later, you would notice a slow negative potential being generated
once the tone sounded. This slow negative potential generally measured
at the vertex is the contingent negative variation (CNV). The CNV is gen-
erated in the laboratory by presenting a first or warning stimulus which
signals that a second stimulus will follow in a specific time period. In
most studies the second stimulus signals cognitive or task processing.
Walter (1967) described the CNV as an expectancy measure, because the
first stimulus suggests that the second will follow.
Another form of event-related potential are very slow potentials that
precede and accompany movement or other activities. If we ask a par-
BRAIN 93
ticipant to press a button whenever desired, we discover that as early as
a second before movement begins, a recognizable EEC waveform starts
to develop. A recording made with an electrode placed over the central
areas of the cortex displays increasingly negativity until, in the few mil-
liseconds before a movement occurs, there is often a slight positive dip
in the wave followed by a steep negative slope, which is terminated si-
multaneously with the beginning of the movement. The beginning of the
movement is accompanied by a large positive deflection and a recovery
to the original baseline. This complex of waveforms is not uniformly dis-
tributed. Technically, this slow increase in surface negativity is referred
to as the Bereitschaftspotential (BP) or the readiness potential (RP).
The readiness potential is maximal at the vertex and initially equal in
amplitude over both hemispheres of the brain. One research paradigm is
to signal to the person which hand to use to make the movement. Prior
to the movement, this potential begins to lateralize and becomes maximal
over the motor cortex contralateral to the body part moved. Some re-
searchers (e.g., Kutas and Donchin, 1980) have suggested that this be-
ginning of lateralization reflects the point in time at which the response
side is determined (i.e., to move the left or right hand). Since the infor-
mation contained within the RP includes nonmotor processes as well as
motor processes, researchers have suggested that by subtracting the re-
sponse from one hemisphere from that of the opposite hemisphere, it
would be possible to obtain a purer measure of motoric preparation for
a response. This measure, referred to as the lateralized readiness potential
(LRP), has become an important tool in the study of the neural basis of
human cognitive-motor processing. Figure 7.9 illustrates the steps re-
quired to calculate the LRP.
To summarize, the development of this measure was based upon the
assumption that the asymmetry of the RP could be used as an index for
Figure 7.9. Derivation of the lateralized readiness potential. The top panel
shows idealized brain potentials from left C3 and right C4 scalp sites in a
warned reaction time task, when subjects know in advance of the imperative
stimulus the hand to be used to make a correct response. In the middle panel,
potentials associated with left-hand movements are shown on the left; those
associated with right-hand movements are shown on the right (WS=warning
stimulus; IS=imperative stimulus). As subjects prepare to make a movement,
a negativity develops that is maximum at scalp sites contralateral to the
responding hand. The asymmetry in these potentials is illustrated by sub-
tracting the potential recorded at the site ipsilateral to the movement from
that recorded contralateral to the movement. Then the difference potentials
for left-hand and right-hand movements are averaged to yield the lateralized
readiness potential, bottom panel. Redrawn with permission from Coles,
M. G. H., 1989, "Modern mind-brain reading: Psychophysiology, physiology,
and cognition," Psychophysiology, 216, 251-269.
the preparation of specific motor acts. To eliminate any RP asymmetries
that may contain activity lateralized with respect to nonmotoric pro-
cesses, the LRP was calculated as the difference between recording sites
contralateral and ipsilateral to the responding hand, averaged over left-
and right-hand responses (see de Jong, Wierda, Mulder, & Mulder, 1988;
Gratton, Coles, Sirevaag, Eriksen, & Donchin, 1988, for alternative ways
to calculate the LRP). The LRP's special significance in cognitive and
sensorimotor research stems from the fact that this component offers a
continuous analog measure of the differential engagement of the left
hand versus right hand associated with cued or uncued voluntary re-
actions (see Hackley & Miller, 1995, for a review of this work).
The growing popularity of the LRP is due to the fact that its neuroan-
atomical and functional correlates are better understood than those of
most other endogenous event-related potentials. Surface and depth re-
cording indicate that the LRP is mainly generated by primary motor
cortex. Moreover, the foreperiod LRP was found to be twice as large
preceding complex movements (subjects were requested to press a se-
quence of three keys, using the index, ring, and middle fingers) than
preceding simple ones (only index finger keystroke was required). Also,
it has been reported that lateralization tends to be larger preceding a
short sequence (one press with the index finger) than preceding a longer
sequence (three presses with the same finger). These and other studies
support the hypothesis that lateralized preparatory activity in motor cor-
tex varies with specific properties of the planned movement.
The event related potentials, including evoked responses, the readiness
potentials, and CNVs, are generally much smaller in amplitude than
spontaneous EEGs and are therefore often not discernible in the raw or
untreated record. In order to examine ERPs, special recording and data
treatment procedures are necessary.
Physiological Basis
What is the source of these recurring rhythmic potentials and event-
related potentials? Until now, their definition and description have been
entirely in terms of electrical comparisons between various points on the
scalp surface and an earlobe, for example. But, what is under there? The
brain, of course, with its more than 10 billion nerve cells, most of them
synapsing with thousands of others. There are six layers of cells in the
cortex which are referred to as layer I through layer VI. There are also
other tissues in the brain. There are glial cells, whose functions are not
yet entirely understood but which do generate a resting potential. There
are also the fluids of the circulatory cerebrospinal system. The scalp itself,
the skull, and the meninges covering the brain intervene between elec-
trode and brain. However, the genesis of the EEC is clearly in brain tissue
proper. Elul (1972) presented the arguments for this conclusion in careful

and fascinating detail for those who would pursue the question. What
structures actually produce EEC has been debated. One theory suggests
that the EEC is generated by pyramidal cells in layers IV and V of the
cortex.
Where a large group of cortical neurons is driven by identifiable vol-
leys of afferent neural activity, as in the case of evoked potentials, the
genesis of surface recorded waves is easily understood in terms of the
mechanisms just described. But in the case of spontaneous rhythms, it is
not at all obvious why the dendrites and synaptic connections of large
groups of cortical neurons would vary in synchrony in a manner capable
of producing the alpha rhythm, for example. While no firm answer can
yet be given, it is probably that subcortical brain structures, particularly
the thalamus, provide synchronizing signals to broad cortical areas. Brain
stem mechanisms have also been shown to control some aspects of the
EEC. Ascending discharges from the reticular formation cause a shift from
alpha and slower rhythms to faster, less synchronized waves in cortical
EEGs.
Recording Procedure
Electrodes Because some event-related potentials approach DC, the se-
lection and preparation of electrodes must be undertaken with the
greatest care. Otherwise, sizable slowly varying offset potentials may de-
velop between electrodes, which can either obscure ERPs or confound
their interpretation. Even with the best electrodes and most careful pro-
cedures, some drift often develops during ERP recording. Because silver-
silver chloride electrodes are relatively nonpolarizing, are stable, and have
a relatively low noise level, they are generally the electrodes of choice
although gold, silver, or platinum discs have also been used for recording
evoked potentials. Of all electrode types, the recessed pellets (composed
of a compressed mixture of silver and silver chloride) are the most stable.
Recording Equipment The amplifier must be capable of following voltages

from DC to more than 100 Hz without distortion. It must have high
input impedance and common-mode rejection capabilities equal to those
required for recording the spontaneous EEC. Filtering and 60-Hz notch
filters are useful but less vital to ERP recording than recording sponta-
neous EEGs.
Averaging Some ERPs can be observed in a single trial before or after

stimulation or responding, and some new techniques are being developed
for single trial analysis. However, because ERPs are generally hidden in
the spontaneous EEC, most techniques use signal-averaging procedures
in which single trials are added together and averaged. ERPs are, by their
nature, time-locked to known events, while spontaneous variations are
BRAIN 97
not. If we compute an average of many repetitions of the event in which
the event always occurs at the same point, then that portion of the EEC
that regularly precedes or follows the event will remain in the average.
The spontaneous variations will tend to cancel out over successive oc-
currences because a given wave is as likely to be positive as negative at
any point in time. The result is a relative enhancement of the time-locked
EEC, the ERP. A very rough representation of ERPs can be made by
carefully measuring the amplitude of the EEC trace at several points
which are time-locked to an event on 20 or more repetitive trials. The
averages of these points can reveal the presence of an evoked or readiness
potential. Today, this process is accomplished by a variety of computer
programs.
Procedure The procedure for ERP recording is the same as that for re-
cording spontaneous EEGs, but extra care should be given to electrode
attachment.
Typical Recordings
The effects of averaging are demonstrated in figure 7.10. In figure
7.10(a), EEGs for five single trials are compared to their average. The
effect of increasing the number of trials on the averaged EEC is shown
in figure 7.10(b). Figure 7.11 shows a comparison of somatic, auditory,
and visual evoked potentials. Two types of slow potentials sometimes seen
in the EEC are shown in figure 7.12. Figure 7.12(a) depicts the readiness
potential (RP) and figure 7.12(b) the contingent negative variation,
(CNV).
Common Problems
Because ERPs are so small (1-10 |0,V), it is possible for an artifact occur-
ring on a single trial to influence the appearance of the average. Addi-
tionally, time-linked artifacts may occur and be emphasized by averaging.
In visually evoked responses, eye blinks may follow the visual stimulus
on several trials with approximately the same latency. To preclude eye-
blink effects in ERPs, electrodes are often placed near the orbit to monitor
blinks and eye movements. Then, either the EEC record or the single trial
record is examined for artifact, and only trials free from artifacts are
included in the average.

It should be recognized that considerable signal manipulation occurs in
the averaging process. The averaged waveform is not necessarily a
clearer version of that which occurs in each individual trial. In fact,
Figure 7.10. The effects of averaging evoked potentials: (a) five single trials
compared to their average; (b) the effect of increasing the number of trials
on the average.
differences between single trial ERPs might well be of interest but are lost
in the averaging. Furthermore, ERPs do not occur with exactly the same
latency from trial to trial, and this latency jitter will significantly influ-
ence the shape of the averaged ERP. Nevertheless, ERPs can be mean-
ingfully subjected to numerical analysis.
To the degree that the waveform of an ERP is reliable, it can be com-
pared to similar ERPs obtained under different circumstances. For in-
BRAIN 99
Figure 7.11. Comparison of somatic, auditory, and visual evoked potentials.
stance, the latency, or time from the event to specified points on the
waveform, can be measured and compared in the usual statistical fash-
ion. Amplitudes may also be measured at specified points relative to the
event. The slope of an identifiable aspect of the wave and areas enclosed
in waveforms can also be calculated. Figure 7.13(a) shows various pos-
sibilities for quantifying an idealized average evoked potential, while fig-
ure 7.13(b) demonstrates common approaches to quantifying slow po-
tentials, the CNV and RP.
Brain Imaging Techniques
EEC and ERPs have a real value in determining the time course of a
response because they reflect millisecond changes within the electrical
activity of the cortex. However, knowing where EEC activity takes place
on the scalp does not in turn give you certainty concerning where the
activity originated in the brain. This is referred to as the inverse problem.
100 P S Y C H O P H Y S I O L O G Y OF SPECIFIC ORGANS AND SYSTEMS

Figure 7.12. Examples of slow wave potentials: (a) the readiness potential;
(b) the contingent negative variation (CNV).
The problem reflects the fact that given a distribution of EEC activity on
the scalp, there is a variety of possible distributions in the cortex that
could lead to the same pattern of scalp activity. Other processes—such
as the fact that electrical activity does not move uniformly through the
brain and that there exists variation in the thickness of different individ-
uals' skulls which influences how the brain's activity is distributed on
the scalp—also add to the problem.
If you have ever placed a magnet under a piece of paper covered with
iron filings, you know that by changing the position of the magnet you
can change the pattern of filings on the paper. You can also do a similar
procedure with electrical activity generated within the brain. Such a pro-
cedure is called dipole modeling. Using computers, one determines what
type of pattern on the scalp would be produced by different generators
in the brain. The pattern generated by the computer could then be com-
pared to actual recorded EEC data. The computer can continue to move
the dipole within the imagined brain until the theoretical pattern of EEC
matches the actual pattern of EEC activity. Although dipole modeling
offers one way of determining localization of activity, there are better
BRAIN 101
Figure 7.13. Quantification of ERPs: (a) measurement of evoked potentials;
(b) measurement of contingent negative variation and a readiness potential.
methods for determining more exact localization of processes in the cor-

tex. These are magnetoencephalography (MEG), positron emission tomogra-
phy (PET), and magnetic resonance imaging (MRI).
Magnetoencephalography uses a SQUID (superconducting quantum
interference device) to detect the small magnetic field gradients exiting
and entering the surface of the head that are produced when neurons
are active. MEG signals are similar to EEC ones but have one important
advantage: magnetic fields are not distorted when they pass through the
cortex and the skull, which makes localization of sources more accurate
than EEC. It should be noted that MEG is only sensitive to tangential
activity, which limits it to activity located in the sulci or cortical folds.
102 PSYCHOPHYSIOLOGY OF SPECIFIC O R G A N S AND SYSTEMS

In order to make a measurement, an individual simply places his or her
head within the sensing device typically containing a large array of sen-
sors that do not require physical contact with the head. Since measuring
magnetic fields using MEG is a complex process requiring liquid helium
(which must be supercooled 24 hr hours a day), the price of this system
is expensive both to acquire and to maintain.
Positron emission tomography systems measure variations in cerebral
blood flow that are correlated with brain activity. It is through blood flow
that the brain obtains oxygen and glucose from which it gets its energy.
By measuring changes in blood flow in different brain areas, it is possible
to infer which areas of the brain are more or less active during particular
tasks. Blood flow using PET is measured by injecting a tracer (a radio-
active isotope) into the blood stream which is recorded by the PET scan-
ner (a gamma ray detector). The general procedure is to make a mea-
surement during a control task which is subtracted from the reading
taken during an experimental task. Although it takes some time to make
a PET reading (which reduces its value in terms of temporal resolution),
the procedure is able to illustrate specific areas of the brain that are active
during different types of processing. Since PET can measure almost any
molecule that can be radioactively labeled, it can be used to answer spe-
cific questions about perfusion, metabolism, and neurotransmitter turn-
over. Some of PET's main disadvantages include expense, the need for a
cyclotron to create radioactive agents, the injection of radioactive tracers
(which limits the number of experimental sessions that can be run for a
given individual), and limited temporal resolution.
Like PET, functional magnetic resonance imaging (fMRI) is based on
the fact that blood flow increases in active areas of the cortex. However,
it uses a different technology from PET; in fMRI local magnetic fields are
measured in relation to an external magnet. Specifically, hemoglobin,
which carries oxygen in the bloodstream, has different magnetic prop-
erties before and after oxygen is absorbed. Thus by measuring the ratio
of hemoglobin with and without oxygen, the fMRI is able to map changes
in cortical blood and infer neuronal activity. Although fMRI has the same
temporal disadvantage as PET, it has a number of advantages including
better spatial resolution and the ability to do repeated images on one
individual.
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BRAIN 105
8
Muscles
Electromyogmphy
There are many reasons for recording muscle activity. Patterns of muscle
action summate to produce movement and maintain posture; these pat-
terns form the domain of kinesiologists and sports psychologists. Physical
therapists record muscle action to document disabilities, to measure ther-
apeutic progress, and to evaluate orthotic and prosthetic devices. The
psychologist studying learning measures muscular activity in order to
record the development of motor skills. The psychophysiologist, too, rec-
ords muscle action, often when no movement occurs, as in the case of
tension headaches or, more generally, to study the patterns of bodily
reaction to stimulation, as in the startle response. One of the most sig-
nificant uses of our muscles, as Darwin (1872) pointed out, is to com-
municate emotional expressions. Another use of muscles is to help us
move quickly as when we are startled. Today, many psychophysiologists
use the startle reflex as a way to measure emotionality. Let us examine
this reflex in some detail.
The startle reflex, present in humans and other animals, is elicited by
an intense stimulus of sudden onset which reaches maximum intensity
in a short time. It has a short latency and occurs within 20 to 40 ms
after the presentation of a 92 dB white noise burst in humans and other
animals (Davis, 1984). The magnitude of the startle response can be
decreased by presenting a less intense stimulus prior to the startle stim-
ulus. This is called pre-pulse inhibition. Although less well studied, some
stimuli (e.g., a light preceding the startle tone) have been shown to in-
crease the startle response. This is called pre-pulse facilitation. An appli-
cation of the prepulse work has been as a correlate of thought disorders
in schizophrenics. In humans the startle reflex is often measured as the
strength of the eye blink created by the orbicularis oculi muscle after an
acoustic stimulus. We can measure many facets of the acoustic eye blink
106
startle reflex: these include blink magnitude, latency to blink, and habit-
uation rate of blink response.
The startle response has been demonstrated in both humans and an-
imals to reflect or result from reflexive modulation in relation to affective
content (cf. Lang, 1995). The startle reflex also has been shown to be
increased in magnitude during high (relative to low) arousal conditions
(Vrana, Spence, & Lang, 1988) and during negative affective states such
as fear, anger, and sadness relative to positive affective states such as joy
and relaxation (Lang, 1995). Although the overall magnitude of the star-
tle reflex decreases after repeated presentations of the startle stimulus,
(i.e., the reflex shows habituation), the reflex continues to be stronger
during negative relative to positive affect even after repeated presenta-
tions of the acoustic startle probe (Bradley, Lang, & Cuthbert, 1993). See
figure 8.1 for an example of the startle reflex.
The question now arises about how to measure muscular activity. The
means of recording muscle action range from filming whole body move-
ments to the recording of action potentials from motor units within a
single muscle. The choice of method should be made with careful regard
for the aims of the procedure; each method has its particular virtues and
shortcomings. Electromyography (EMG) is best suited for examining the
way in which tension develops within a muscle, for determining the firing
rates of particular motor units in relation to the recruitment of others,
and for revealing activity too small to produce visible movement. But
electromyography requires the attachment of electrodes, usually with ac-
companying wires that restrict movement. Furthermore, as will be de-
tailed later in the chapter, EMG signals are subject to interference and
distortion when movement occurs.
The EMG is a record over time of electrical potentials originating in
muscles. EMGs may be obtained either by inserting electrodes into the
muscle or by placing electrodes on the skin over the muscle or muscle
group of interest. This chapter will describe only the techniques for the
latter procedure, called surface electromyography. For a more in-depth
Figure 8.1. An example of the startle reflex. This figure shows an electro-
myogram (EMG) recorded from below the eye with the occurrence of a startle
probe. In order to quantify the response, this signal would be rectified (all the
negative values would be made positive) and integrated (the area under the
curve would be computed).
MUSCLES 107
discussion of the skeletomotor system, see Cacioppo, Tassinary, & Frid-
lund (1990).
Physiological Basis
The cellular basis of EMGs is muscle action potentials spreading over

skeletal muscle cells following neural stimulation. The intracellular result
of muscle action potentials is contraction, as described in detail in chapter
2. A wave of depolarization can be recorded at a distance as a momen-
tary difference in potential between electrodes spaced over the muscle.
Since skeletal muscles are functionally divided into motor units which
are activated in unison, action potentials are produced simultaneously in
all the cells of a unit. The cells of a motor unit are usually distributed
through a muscle rather than being concentrated at some point, and the
firing of a motor unit produces a chord, so to speak, of action potential
from the muscle. Seen from electrodes on the surface of the muscle, this
chord will appear as a wave, the amplitude of which will be a function
of the number of cells in the unit and their proximity to the electrode.
Distant cells will contribute less to the wave than nearby cells will. If the
interstitial fluid is considered to be homogeneous, then the contribution
of a single action potential to the surface potential seen by an electrode
will decrease as the square root of the distance from cell to electrode.
Since the cells of the motor unit are usually dispersed longitudinally as
well as radially, the sum of the individual cellular action potentials will
differ in waveform from that of the single cells. The result is that the
surface-recorded EMG resulting from the firing of a single motor unit will
probably be unique, unlike that from other motor units within the mus-
cle. When a skeletal muscle is nearly relaxed, the periodic firing of each
of several motor units can be distinguished in terms of its amplitude,
frequency, and waveform. In fact, in an early study Basmajian (1963),
recording from needle electrodes inserted into the muscle, showed that
voluntary control could be exercised over individual motor units. He re-
ported that individuals were able to increase or decrease the rate of firing
of single motor units.
However, as muscle activity increases, the time between action poten-
tial spikes becomes shorter and shorter as motor units increase their rate
of firing. Other motor units are recruited until the firings of single motor
units fuse with each other to produce a complex waveform that is no
longer interpretable in terms of a motor unit.
More important for most applications, EMG can be related to the ten-
sion created in the muscle. In isometric contractions (when there is no
movement), there is a close correspondence between the surface EMG
summed over time and tension developed in the muscle over a wide range
of tensions. However, the relationship does not hold for isotonic mea-
surement, where movement occurs. This is partly because the tension
developed in a muscle in response to a standard excitatory stimulus is a
function of its length; in general, a muscle produces a smaller increment
in tension at shorter initial lengths. Since a muscle becomes progressively
shorter in an isotonic contraction, the length-tension relationship is con-
tinually changing. Also, as the muscle shortens, its fibers move under
the electrodes, altering the relationship of the electrodes to the signal-
generating tissue. Nevertheless, it has been determined that during con-
stant velocity shortening there is a direct relationship between the inte-
grated EMG and the tension exerted by the muscle.
The observed EMG varies from muscle to muscle, depending on the
size of the muscle, the distribution of motor units within it, the size of
the motor units, and the anatomy. Examples of typical recordings from
various muscles are illustrated later in this chapter.
Recording Procedure
Since the surface recorded EMG may vary in amplitude from a few mi-
crovolts to more than a millivolt, depending on the muscle and its state
of contraction, the requirements for recording systems and procedures
will vary with the intended use. If, for example, the aim of recording is
simply to know when (or if) an arm or leg moved, the recording system
need not be able to follow the tiny variations in tension which continually
occur. A relatively simple system and somewhat crude procedures will
suffice. However, to record the bursts attributable to single motor units,
or to follow the buildup of tension before movement and when movement
does occur, a system capable of high differential amplification must be
used and accompanied by patient and careful recording procedures. The
equipment and procedures described here are those for recording micro-
volt signals. The same equipment and procedures may be used for less
sensitive recording, but with the amplifier sensitivity appropriately re-
duced.
Electrodes
EMG electrodes should have the following characteristics: low impedance,
low electrode potential (nonpolarizing), stability not subject to movement
artifact, small size, and lightweight. If the electrode is either a combina-
tion of silver and silver chloride or carefully chlorided silver, it will be
very stable, relatively nonpolarizing, and develop the smallest electrode
potential of any available material. If, in addition, the electrode housing
is constructed so that the electrode is of the floating, or liquid junction
type—that is, the conductor is not directly in contact with the skin—
then movement artifacts are greatly reduced. Floating silver-silver chlo-
MUSCLES 109
ride electrodes are commercially available in several configurations (see
chapter 3).
Electrode Placement
The electrodes of a pair will record the difference in electrical potential
between them originating in nearby, and to a lesser degree, distant mus-
cle tissue. The placement of the pair and the distance between them,
therefore, determines which muscles will contribute to the recorded EMG.
The two principal considerations are: (1) both electrodes should be over
the same muscle or muscle group, and (2) the pair should, where pos-
sible, be on a line parallel with the muscle fibers. A third factor, the
distance between the electrodes of a pair, is dependent upon the length
of the muscle and the desired discreteness of the EMG. Closely spaced
electrodes (1 or 2 cm between them) will generally be superior for ob-
serving the activity of single motor units than a widely spaced pair, but
not as good for obtaining an index of overall muscle tension.
In the case of the muscles moving the long bones of the skeleton, the
application of these principles is relatively simple. For example, in order
to record EMGs from biceps, the two leads, separated by 2 or 3 cm, are
centered over the belly of the muscle (which may be located by palpation)
in a line parallel with the underlying bone, that is, a line drawn from
elbow to shoulder. If several recordings are to be made from the same
subject, or if recordings from several subjects are to be compared, then
care should be taken to identify the placements in terms of anatomical
landmarks (see figure 8.2).
Since an electrode pair will record muscle potentials from nearby mus-
cles, as well as the muscle under study, electrode placement and the
interpretation of EMGs is difficult where several different muscles are near
either electrode. Similarly, deep muscles may influence EMGs from elec-
trodes over a superficial muscle.
In the forearm, the muscles moving the fingers, those bending the
wrist, and those rotating the wrist are considerably interwoven. An EMG
recorded from leads anywhere on the forearm will include, to some de-
gree, contributions from all three muscles. A similar situation exists in
the back, neck, head, and (to a lesser degree) the lower and upper leg.
Electrode placement in these situations is arbitrary, and serviceable EMGs
can be obtained by applying the principles just enumerated. There have,
however, been attempts to standardize EMG lead placements. Some of
Figure 8.2. Atlas of EMG electrode placements. From J. T. Cacioppo and L. G.

Tassinary (Eds.), 1990, Principles of Psychophysiology, New York: Cambridge
University Press. Reprinted with the permission of Cambridge University
Press.
these are described by Cacioppo, Tassinary, & Fridlund (1990). Histori-
cally, Lippold (1967) includes descriptions of ten lead arrangements
which were originally devised by J. F. Davis (1952). These placements,
while they may be useful as guides, are arbitrary, and at least one of
them (the forehead lead) violates the principles of electrode placement
previously described (Davis, Brickett, Stern, & Kimball, 1978). That is,
some individuals place frontalis electrodes horizontally across the fore-
head when in fact the muscles run in a vertical manner. We must reem-
phasize that regardless of the placement of electrodes, the EMG recorded
from most locations will include contributions from other muscles. More
will be said on this subject in the section on interpretation of EMGs.
Recording Equipment
The requirements for amplification of EMGs are as follows: high gain,
high input impedance, frequency response from 1 to 1,000 Hz. Since
there is little or no DC in the EMG, capacitively coupled (AC) recording
systems are preferred and selectively amplify fast voltage changes, thus
ignoring steady states and very slow changes. Typical amplification sys-
tems include the clinical electromyograph, laboratory multipurpose poly-
graphs, and laboratory polygraphs with associated signal conditioning
apparatus. Any of these systems is adequate if properly used. Since the
frequencies in the EMG signal range up to 1,000 Hz, special consideration
should be given to the frequency characteristics of the recording device.
Most ink writing recorders are unable to follow reliably frequencies above
60 or 80 Hz. Thus, one should use an oscilloscope computer display for
accurate portrayal of the signal. In using a computer system, it must be
determined that the sampling frequency of the analog-to-digital conver-
sion is sufficiently high to record the highest frequency components in
the EMG.
Some EMG units, particularly those intended for biofeedback training,
include no provision for displaying the raw, untreated EMG. Rather, their
output is an auditory signal or visual (meter) display of the sum of the
EMG. Such systems cannot be used for studying EMGs, although they
may be useful for biofeedback training. Many EMG recorders include a
provision for rectifying and, often, for integrating the signal as an addi-
tion or an alternative to direct recording. The function of integrators is
described later in this chapter.
Procedure
Electrode attachment must be accomplished with great care. After the
electrode sites are chosen, carefully measured, and marked according to
predetermined criteria, any hair under the site should be moved. The
skin at the site is then cleaned with alcohol. Then the skin is abraded to
remove the high-impedance dead surface layer of skin. The exposed area
may be rubbed with electrode jelly, although the jelly will reduce the
effectiveness of a electrode adhesive if it is allowed to spread beyond the
central, active portion of the electrode. Also, any connection between
the electrode paste or jelly at the sites of the two electrodes will reduce
the recorded EMG. The electrodes must be firmly attached to the skin.
Any movement of the electrodes will produce large artifacts.
The electrodes are prepared by coating or filling (in the case of recessed
electrodes) them with electrode jelly. Some EMG electrodes are held in
place by suction. Commercial electrodes that are affixed with collars with
adhesive on both sides, are particularly convenient. Some laboratories
employ a third, ground lead, for EMG recording, which is similarly at-
tached. The impedance between the active leads of a channel should be
tested with an impedance meter (preferably not an ohmmeter, because
it will pass a DC current through the electrodes). The impedance of the
pair should be 5,000 (1 or less. Higher electrode impedance will reduce
the signal introduced to the amplifier and provide a possible source of
interference. If the impedance is greater than 5,000 fi, the electrodes
should be removed and reapplied after the underlying skin has been fur-
ther abraded and new electrode jelly applied. The time spent in carefully
applying electodes will be more than repaid by clear, interpretable EMGs
later in the recording session.
The amplifier gain should be adjusted to the appropriate level for the
type of recording to be generated. If single motor unit reponses or other
low-level signals are of interest, then the gain will be at or near the
maximum the amplifier allows. The range of input voltages in this case
will be from less than 1 jiV to 50 uV or more. If EMGs are to be recorded
during movement or isometric tension, then the gain will need to be
reduced considerably. EMGs in this application will range upward from
50 jiV to more than a millivolt. Unfortunately, the entire range of activ-
ities cannot be usefully recorded at a single gain setting. Small prepara-
tory EMGs are often missed because the attenuation is set so that the
largest EMGs occurring during contraction can be recorded without ex-
ceeding the limits of the recording system.
As previously stated, the frequency of EMG ranges from about 1 Hz
to more than 1,000 Hz, although most of the signal power is between
10 and 150 Hz. If there is a signal-conditioning filter, it should be set for
a band pass in this range.
Once the amplifiers are adjusted to the desired gain, a standard cali-
bration signal should be introduced at the input. Some EMG units and
polygraphs are equipped with an internal calibration signal. The ampli-
fiers should be adjusted so that calibration signals produce identical out-
put for each amplifier. Calibration signals recorded this way will serve as
the basis for quantifying the EMGs.
MUSCLES 113
Typical Recordings
Figure 8.3 shows the electrode location for recording from the biceps.
Figure 8.4 shows the firing of single motor units recorded at high gain
from: (a) biceps, (b) lumbricalis, and (c) forehead. In figure 8.5 one can
see successive contraction and relaxation recorded from a muscle. Figure
8.6 shows EMG during an arm movement under the following conditions:
(a) slow speed, medium-gain, and (b) high speed, high-gain recording of
the beginning of movement in (a). Figure 8.6 reveals the differences be-
tween high- and medium-gain EMG recording. In Figure 8.6 (a) the EMG
can be followed throughout the movement and can be used to document
the beginning and end of the movement and something of the effort
expended. In figure 8.6 (b) the maximum tension is forfeited in order to
be able to examine the developing tension before the movement begins.
Figure 8.7 displays frequency histograms of the EMG recorded from the
forearm in the following states: (a) relaxation, (b) supporting 100 g, (c)
supporting 435 g, (d) during a 30-kg squeeze.
Figure 8.3. Electrode locations for recording from the biceps.
114 PSYCHOPHYSIOLOGY OF S P E C I F I C ORGANS AND SYSTEMS

Figure 8.4. Single motor units recorded fro the following locations: (a) biceps;
(b) lumbricalis; (c) forehead.
Common Problems
As is the case with EEC, the appearance of 60-Hz interference in the

recorded EMG is by far the most common difficulty. The problem may
arise from any of several sources, most of which are discussed in chapter
7. Figure 8.8 displays an EMG from the biceps, the initial part of which
is contaminated by 60-Hz noise. In the case of EMGs, the 60-Hz notch
filters currently available on many polygraphs should not be used. As
may be noted in figure 8.8, 60 Hz is near the center of the most powerful
portion of the EMG frequency range, so that eliminating it and attenu-
ating nearby frequencies will distort EMG considerably. Figure 8.9 dem-
MUSCLES 115
Figure 8.5. Contraction and relaxation of a muscle.
onstrates artifacts generated by movement and by a loose electrode.

When interference begins to appear in a previously clear record, the most
likely source is a loose electrode or connection.
Other physiologically generated signals may contaminate the EMG.
The electrical potential generated by the heart (the EKG) commonly ap-
pears, particularly in neck and back leads. This may be overcome by
moving the leads closer together or by repositioning the subject. A
ground lead attached nearby (not between the two active leads) will often
reduce the EKG in the EMG leads. The EKG can often be eliminated by
orienting the axis of the EMG electrodes at right angles to the axis of the
heart. EEC is clearly present in figure 8.10; the frontal EEC quite reliably
contributes to forehead EMG. The EEC is recognizable as a band of low-
amplitude, somewhat faster activity. The EEC contribution can be re-
duced but not eliminated by reducing the space between electrodes to 2
cm or less.
While no obvious distortion appears from incorrect electrode place-
ment, the resulting EMG may not emanate from the intended muscle.

The most common error is to place the two leads of a pair of electrodes
at right angles to the fibers of the muscle. This greatly attenuates and
distorts the signal, as may be seen in figure 8.11 (a), taken from pairs of
leads correctly and incorrectly placed over biceps. Alternatively, the two
electrodes are sometimes incorrectly placed on different muscles. EMG
will then represent the difference between the activity of the two muscle
groups rather than that of either one of them. The forehead placement
Figure 8.6. EMG during an arm movement: (a) slow speed, medium gain; (b)
high speed, high gain.
MUSCLES 117
Figure 8.7. Frequency histograms of power in the EMG during four tensions:
(a) relaxed arm; (b) 100-g weight; (c) 435-g weight; (d) 30-kg squeeze.
118
Figure 8.8. EMG recorded from thebicep with 60-Hz interference
Figure 8.9. Artifacts generated by movement and a loose electrode.
119
Figure 8.10. EMG recorded from the forehead contaminated with frontal EEC.
Figure 8.11. EMG recorded with electrodes correctly and incorrectly placed
(a) from the bicep;
Figure 8.11. (continued) EMG recorded with electrodes correctly and incor-
rectly placed (b) from the frontalis.
suggested by a number of biofeedback practitioners is an example of such

a placement and was studied by Davis, Brickett, Stern, and Kimball,
1978. Figure 8.11(b) compares records from these leads to a simulta-
neously recorded EMG from a single frontalis muscle.
In what follows, methods are suggested for the analysis of surface re-
corded EMGs. Considerable caution must be demonstrated, however, in
assigning their muscle origin. It is common to use such terms as "fron-
talis EMG," or "corrugator" or "buccinator" EMG, for example. But, as
indicated earlier, such specific designation is unwarranted because the
surface record reflects distant as well as local activity. Particularly in
anatomical areas where there are several muscles and where, as is often
the case, they are arranged in superficial and deep layers under the same
electrode pair, one simply cannot discern which muscle or which com-
bination of them is responsible for the momentary EMG. It is preferable
to indicate the position of the electrode in terms of anatomical landmarks.
MUSCLES 121
Nevertheless, EMGs are amenable to quantitative analysis. EMGs,
where there is no movement, will usually appear similar to those of figure
8.4. Where discharges of single motor units are recognizable by their
amplitude, they may simply be counted, so that an analysis of counts per
unit time results. The ultimate limit of this counting procedure occurs
when the individual spikes fuse to form the complex waves seen in figure
8.5. Here the analysis of the untreated record becomes very difficult. If
Figure 8.12. EMG preparatory to an arm movement: (a) raw EMG; (b) rectified
signal; (c) rectified and filtered using two filters.

the record is divided into equal time segments, various apects of the
complex waveform in each may be measured. The largest deflections in
each segment, or the average of some number of the largest in each
segment, may be used as an approximation of the amplitude of the EMG.
These are tedious procedures, and the relation of any single measurement
to the tension developed by the muscle is questionable. For these reasons,
EMGs are often subjected to considerable alteration within the recorder's
output stages.
The most common treatment first specifies the EMG and then inte-
grates the rectified signal. Rectification combines negative and positive
deviations around a specified value; it is as though the EMG record was
folded lengthwise through the middle of the trace and the underlying
waves traced onto the top. Integration of the rectified record with respect
to time effectively sums continuously the area above and below the trace,
which at any moment represents the absolute EMG amplitude. The in-
tegral may be continually displayed or discretely "dumped" onto the
polygraph for writeout whenever the integrated voltage reaches some
predetermined level. Both integrals permit evaluation of EMGs in terms
of amplitude-time relations. As was indicated earlier, the integrated EMG
does correspond linearly to muscle tension over a wide range of isometric
contractions. Rectification, integration, and filtering can be accomplished
by computer as well as in the output stages of the EMG recorder. Figure
8.12 displays: (a) the raw EMG preparatory to arm movement, (b) rec-
tification of the same record, and (c) rectified, filtered EMG using two
filters.
Where the EMG is time-locked to an identifiable event, such as an
external stimulus or a movement, a researcher can construct averages
of many trials of the rectified EMG in a manner identical to that followed
for averaged evoked potentials from the brain (see chapter 7).
References
Basmajian, V. (1963). Control and training of individual motor units. Science,

141, 440-441.
Bradley, M. M., Lang, P. ]., & Cuthbert, B. N. (1993). Emotion, novelty, and
the startle reflex: Habituation in humans. Behavioral Neuroscience, 107,
970-980.
Cacioppo, J., Tassinary, L., & Fridlund, A. (1990). The skeletomotor system.
In J. Cacopppo & L. Tassinary (Eds.), Principles of psychophysiology: Phys-
ical, social and inferential elements. Cambridge: Cambridge University
Press.
Darwin, C. (1965). The expression of emotions in man and animals. Chicago:
University of Chicago Press. (Reprint of edition by D. Appleton & Co,
London, 1872).
Davis, C. M., Brickett, P., Stern, R. M., & Kimball, W. H. (1978). Tension in
the two frontales: Electrode placement and artifact in the recording of
forehead EMG. Psychophysiology, 15, 591-593.
MUSCLES 123
Davis, J. F. (1952). A manual of surface electromyography. Montreal: Laboratory
for Psychological Studies, Allen Memorial Institute of Psychiatry.
Davis, M. (1984). The Mammalian Startle Response. In R. C. Eaton (Ed.),
Neural Mechanisms of Startle Behavior. New York: Plenum.
Lang, P. J. (1995). The emotion probe. Studies of motivation and attention.
American Psychologist, 50, 372-385.
Lippold, 0. C. J. (1967). Electromyography. In P. H. Venables and I. Martin
(Eds.), A manual of psychophysiological methods. Amsterdam: North-
Holland.
Vrana, S. R., Spence, E. L., & Lang, P. J. (1988). The startle probe response:
A new measure of emotion?. Journal of Abnormal Psychology, 97, 487-
491.
124 PSYCHOPHYSIOLOGY OF S P E C I F I C O R G A N S AND SYSTEMS

9
Eyes
Pupillography and Electrooculogmphy
Investigators have claimed that various responses of our eyes can be used
to determine a great variety of things, from interest in sexual feelings, to
the relative degree of processing occurring in the brain. Some of the more
common areas of research have included relating the size of the pupil to
arousal and examining eye movements in relation to processes such as
reading and dreaming. Of the various measures possible, psychophysiol-
ogists have been most interested in pupillography, or the measurement of
the size of the pupils and eye movement. In this chapter, we will discuss
pupillography and electrooculography, one technique for studying the po-
sition of the eyes.
Pupillography
In both Eastern and Western cultures, a tradition dating back hundreds

of years views the pupils as "windows of the soul." There are stories
about merchants who were able to sell their wares by watching the
changes in pupil size as the buyer first looked at an item; when the sellers
noticed a pupillary response, they knew which item the buyer was really
interested in and could set their price accordingly.
Not only has the pupillary response made its way into our folklore,
but as Janisse (1977) and Hess (1975) have noted in their brief histories
of the area, scientists over the past 200 years have also been intrigued
by the response. For example, in the 1870s Darwin related pupillary
dilation to emotional responses such as fear and surprise. In the twentieth
century there was interest in the pupillary response in different popula-
tions, such as children, people with schizophrenia, and groups given var-
ious drugs. Although there has been scattered interest in the pupillary
response for many years, it was not until the work of Hess in the early
125
1960s that this response gained the attention of numerous psychologists
and other researchers.
Hess (1975) described his first "experiment" in which he showed a
series of pictures, including landscapes and a nude female, to his research
assistant, James Polt. After a number of pictures, Hess noticed that Polt's
pupils became larger in response to the picture of the nude. Following
this, Hess and Polt designed other experiments to understand better how
psychological stimuli affect pupillary size. The first summary of this re-
search was presented by Hess (1965) in a Scientific American article in
which he suggested that attraction to a stimulus resulted in pupillary
dilation and that pupillary constriction was the outcome of viewing stim-
uli that were "distasteful or unappealing." Not only did Hess argue that
females responded with greater dilation than males to a pleasant stimu-
lus, such as a picture of a mother and baby, but also that people whose
pupils were dilated were judged more appealing. He examined this ques-
tion by showing two pictures of a female to males. The two pictures were
identical except that in one picture the pupils were shown to be larger
than in the other. Although the males reported that they could not dis-
tinguish one picture from the other, they consistently preferred the one
with the larger pupils. In the middle ages, the drug belladonna (atropine)
was used to produce large-diameter pupils, which were symbolic of
beauty in women, hence the name belladonna, "beautiful woman." Jan-
isse (1977) reviewed the work of Hess and others in this area and pointed
out that there is still much controversy over the relationship of psycho-
logical factors and pupillary responses, especially Hess' aversion-
constriction hypothesis.
Although we know that light intensity is a major determinant of the
pupillary response, it is the psychological factors including cognitive,
emotional, and motor processes that have been of interest to psycho-
physiologists (see Beatty, 1982; Sirevaag & Stern, 2000, for reviews). For
example, when students were asked to commit a mock crime, those with
knowledge of the crime scene showed larger pupillary responses to pho-
tographs of the scene than individuals without any knowledge of the
crime (Lubow & Fein, 1996) In fact, these authors reported that 50% of
those who took part in the mock crime and 100% of those who did not
could be correctly identified from pupillary responses. Other work sug-
gests that mental effort or mental load also plays a role in pupillary
dilation. For example, complex sentences have been shown to produce
larger pupillary changes than simple ones (Just & Carpenter, 1993).
Likewise, it has been reported that pupil dilation increases as an individ-
ual is asked to remember an increasing number of digits (Granholm,
Asarnow, Sarkin, & Dykes, 1996). Other intriguing areas of research
involving pupillary responses are those involving psychopathology (e.g.,
Rubin, 1974; Steinhauer & Hakerem, 1992), those examining the inter-
relationship of pupillary responses with other psychophysiological vari-

ables such as heart rate (e.g., Libby, Lacey, and Lacey, 1973; van der
Molen, Boomsma, Jennings, & Nieuwboer, 1989), and those that use
pupillary responses as a measure of short-term activation during infor-
mation processing or motor preparation (e.g., Beatty and Wagoner,
1978; Richer & Beatty, 1987). This last area of research has implications
for human factors research; that is, pupil diameter can be used to infer
work load and fatigue in groups such as airline pilots (cf. Sirevaag &
Stern, 1999). For further information, the interested reader should con-
sult the special issue of the Journal of Psychophysiology, 1991 (3), on
pupillary response.
Physiological Basis
Physiologically, dilation and constriction of the pupil require the involve-
ment of the autonomic nervous system in the following manner. The
sympathetic nervous system dilates the pupil through a contraction of
the medial fibers of the iris. Parasympathetic activation contracts the
circular muscle or the iris and brings about the constriction of the pupil.
There are a number of reflexes, as well as other factors, which can bring
these responses into play. Tryon (1975) has noted 23 such factors, which
are presented in table 9.1. This table points out the complexity of per-
forming carefully controlled studies with this response measure.
Recording Procedure
Photographing the Eye One of the techniques used by Hess (1972) was
photography of the eye with a 16 mm camera. The procedure was as
follows. A subject would come into the laboratory, sit down, and look
into a boxlike apparatus approximately 2 ft (0.60m) X 2 ft X 2 ft in size.
The far side of the apparatus consisted of a screen on which slides could
be projected. Mounted on one side of the apparatus was a 16 mm camera
aimed at a mirror placed outside of the subject's line of vision which
reflected the eye. Hess reported an almost perfect concordance between
the pupillary responses of the two eyes and thus suggested that only one
eye need be recorded. The pupillary changes were recorded on infrared
film, which permitted filming under various lighting conditions and al-
lowed for sharper definition between the pupil and the iris than found
with standard negative films. The largest drawback to the filming tech-
nique was the film processing time and scoring, usually by hand, which
was an expensive and time-consuming procedure. Even a small study
required a large number of pupillary measurements. According to Janisse
(1977), 20,000 separate measurements would not be unusual; one study
used 100,000 measurements. Today, computers allow for easier mea-
surements to be made.
EYES 127
Table 9.1. Sources, and Descriptions, Regarding Variation in Pupil Size
Sources Descriptions
1. Light reflex Pupil constricts with increased intensity of
illumination and dilates with decreased
intensity of illumination
2. Darkness reflex Momentary dilation due to interrupting a
constant adapting light; different from the
light reflex
3. Consensual reflex Stimulation of one eye affects both eyes
equally; failure called dynamic anisocoria.
4. Near reflex Constriction due to decreasing the point of
focus
5. Lid-closure reflex Momentary contraction followed by redila-
tion
6. Pupillary unrest (hippus) Continuous changes in pupil diameter
7. Psychosensory reflex Restoration of diminished reflexes due to ex-
ternal stimulation
8. Age Decreased diameter and increased variability
with age
9. Habituation Pupil diameter decreases, speed of contrac-
tion increases, magnitude of reflex de-
creases
10. Fatigue Diameter decreases, amplitude and fre-
quency of hippus increase; age amplifies
these effects
1 1 . Alertness and relaxation Alertness suggestions decrease and relaxa-
tion suggestions increase pupil size
12. Binocular summation Constriction is greater when both eyes are
stimulated
13. Wavelength (pupillomo- Larger dilation to chromatic than achro-
tor Purkinje phenom- matic stimuli; as intensity of illumination
ena) is increased, proportionately more con-
striction is elicited by shorter wavelengths
14. Alcohol Dilates the pupil in proportion to the per-
centage of alcohol in the blood
15. Sexual preference Dilation to sexually stimulating material
16. Psychiatric diagnosis Abnormal pupillary responses in schizo-
phrenics and neurotics
1 7. Pupil size Stimuli involving larger pupils elicit more
dilation
18. Political attitude Dilation for preferred political figures
19. Semantic stimuli Small pupil diameters associated with high
recognition thresholds
20. Taste Pleasant taste elicits dilation
2 1 . Information processing Increasing dilation to increasingly difficult
load problems
22. Task-relevant response Having to make a motor response aug-
ments pupillary response
23. Incentive Increases diameter on easy problems only
Source: From W. W. Tryon 1975, "Pupillometry: A survey of sources of variation," Psychophysi-
ology, 12, 90-93. Reprinted by permission. Research documenting each factor can be found in
Tryon (1975).
Figure 9.1. Eye monitoring system.
Electronic Scanning Electronic scanning is the most highly developed

method for pupillary measurements. One such system is shown in figure
9.1. As can be seen in the figure, the electronic scanning device uses a
television camera that records the eye. Most units have a filter that allows
infrared light, which cannot be seen by the subject, to be projected on
the eye. Within the control unit a special circuit detects the amount of
light reflected from the pupil and cornea. The greater the dilation, the
less light that is reflected. The television monitor also allows the experi-
menter to check the picture quality from which the pupil measurement
is made for possible artifacts. The television scanning type of system also
gives the experimenter immediate feedback concerning the pupillary re-
sponse.
Common Problems
Since the measurement of pupillary responses is machine specific, the
most common problems, other than gross movements, are related to the
manner in which the measurement is taken. Video systems are limited
EYES 129
by the scanning rate of the system, which is typically 60 samples per
second. There are other problems the researcher should consider, such
as whether the system will allow the subject to wear eyeglasses or contact
lenses; the maximum amount of head movement the subject can make;
how comfortable the subject is; whether the subject can make verbal
responses while still giving accurate pupillary data; and so forth. There
is one additional problem with television systems which relates to the
manner in which the scanner determines the size of the pupil; some
systems require extensive calibration for each subject and for a given
subject under different conditions.

The analysis procedure of Hess (1972) compared the changes in pupillary
responses of subjects while they observed a control slide and a stimulus
slide, both of which were shown for 10s. Since both slides were equated
for brightness and contrast, Hess interpreted any change in pupillary
response to be of an informational or emotional nature. During the con-
trol and stimulus situations, about 20 frames of the film were taken for
each slide and then analyzed. In his early studies, Hess simply compared
pupil area during the showing of the control and stimulus slides. How-
ever, in his later work, the measure of percentage change in diameter
was used. Television scanning devices use similar measurement units.
Eye Movements
Eye movement has been used by psychophysiologists to infer cognitive

processing. For example, it is possible to record the manner in which
good and poor readers move their eyes while reading and to infer from
these movements differences in their cognitive approach to the material.
Oster and Stern (1980) found that good readers, when changing from
general reading to detailed information reading, did not change the
length of time between the fixation points but increased the amount of
time they paused during each fixation. Poor readers did not change the
amount of time they paused; instead, they decreased the amount of ma-
terial between the fixation points. From the eye-movement records, these
researchers were able to infer cognitive processing.
Eye movements have also been used as an indirect measure to infer
the presence of dreams, fantasy, and differential processing by the two
hemispheres of the human brain. According to the classification system
developed by Dement and Kleitman (1957), the initial stages of sleep are
characterized by a slowing in EEC frequency and a lack of eye movement.
In the so-called fifth stage of sleep, however, a paradoxical phenomenon
appears. The person's physiological responses begin to vary more, in-
eluding an increase in eye movements and a reduction in chin EMG. If
someone is awakened during rapid eye movement or REM sleep there is
a high probability that dreaming will be reported. In one of the early
studies, Dement and Kleitman (1957) reported that 80% of their subjects
were dreaming when awakened from REM sleep, whereas only 7% re-
ported dreaming when awakened from non-REM sleep. Thus, for re-
searchers interested in studying subjective factors such as dreaming, fan-
tasy, and flow of consciousness, the measurement of eye movements
offers an objective indication of their occurrence.
Eye movement has also been thought to be an indirect measure of
hemispheric activation. Based on early work by Day (1964) and Duke
(1968), Bakan (1969) suggested that the direction in which one looks
after being asked a question is related to the hemisphere of the brain
which has been activated in response to the question. The present theory
is that right hemispheric activation produces an initial left eye movement,
whereas left hemispheric activation produces a right eye movement. This
reasoning suggests that by observing eye movement, a researcher can
determine which hemisphere is being used in the processing of various
types of material. For example, it has been suggested that the left hemi-
sphere is more active in the analysis and production of verbal material,
whereas the right hemisphere is more active in the comprehension of
spatial and musical material. Research has suggested that there is some
validity to this claim, although the total picture is not simple (see Ehr-
lichman and Weinberger, 1978, for a review of hemispheric activity in
relation to eye movements).
Various studies have reported that individuals with schizophrenia dis-
play dysfunctions of a particular type of slow eye movement referred to
as smooth pursuit (see Clementz & Sweeney, 1990, for a review). The
smooth pursuit system allows our eyes to follow relatively slowly moving
objects such a pendulum on a clock. Researchers can measure the dif-
ference between a target's movement and that of a person's eye move-
ment. In addition to schizophrenics showing pursuit eye movement dys-
functions, studies have reported that first-degree relatives also show
dysfunctions. This has suggested to some that eye movement dysfunc-
tions may be a biological marker for schizophrenia.
Physiological Basis and Classification

Control of Eye Movement Movement of the eye is controlled by six mus-
cles that are innervated by the third, fourth, and sixth cranial nerves.
These muscles, working in antagonistic pairs, coordinate movement of
both eyes in horizontal, vertical, and circular directions. For example,
when both eyes look to the right, the left medial rectus muscles and the
right lateral rectus are activated, while the right medial and the left lat-
eral muscles are inhibited. In a similar fashion, vertical movement is
EYES 131
controlled by the superior and inferior rectus muscles. The final two mus-
cles, the superior oblique and the inferior oblique, are involved in rota-
tional movements of the eye. Although we have described the three
groups of muscles separately, in reality all are involved in each movement
of the eye.
Types of Eye Movement As one reads a page of printed material, the eyes
move from one fixation point to another. The voluntary jump from one
fixation point to another is referred to as saccadic eye movement. These
movements are fast and characterized by a high initial acceleration and
final deceleration. Some scientists have even speculated that the saccade
represents the fastest somatic movement that any muscular system in
the body can produce.
In contrast to saccades, which are quick, the eyes display slow move-
ments known as smooth pursuit movements. These movements appear
not to be under voluntary control and can be elicited by having a person
view a moving visual field. Independent of saccadic movements, the
smooth movements help to stabilize on the retina the image of the mov-
ing object being viewed. In the same manner that smooth movements
represent the eye's response to external movement, compensatory move-
ments of the eyes represent movement on the part of the person—either
of the head or the body or both.
Another type of eye movement is referred to as nystagmus movements,
which consist of oscillatory motion of the eye. Specific examples include
optokinetic nystagmus, which may be elicited by a moving pattern con-
taining repeated patters; vestibular nystagmus, which may be elicited by
head movement that stimulates the semicircular canals; and spontaneous
or gaze nystagmus, which is an anomaly of the eye related to certain
neurological disorders.
Two additional types of eye movements are torsional eye movements
and vergence eye movements. Torsional eye movements are rotational
movements about the line of gaze and are smooth and compensatory.
Vergence movements are the mechanism by which binocular fixation is
maintained. The eyes thus move in opposite directions, so that an object
moving toward or away from the eyes always appears as one object.
These vergence eye movements are relatively slower than most other
types of eye movements. Table 9.2 outlines the various types of eye move-
ments, along with the size, speed, and latency of each.
Recording Procedure
Eye position, the direction of gaze, and eye movement, or the change in
gaze, can be measured by various means. One simple method is to watch
the person's eyes or even to record their movement with a movie camera.
The problems with just looking include being obstrusive and not obtain-
Table 9.2. Types of Eye Movements
Size
(degrees Latency Speed Possible
Type Description of arc) (ms) (degrees/s) Recording Methods
Saccadic movements Conjugate fast eye movements 5-50 100-500 100-500 Photography
that carry the eyes from one Corneal reflection
fixation point to another Photoelectric
Electrooculography (EOG)
Smooth pursuit move- Conjugate involuntary slow eye 1-60 200 1-30 EOG
ments movements to follow slowly Photoelectric
moving targets Photography
Compensatory Smooth conjugate involuntary 1-30 10-100 1-30 Photography
movements movements used to compensate Photoelectric
for passive or active move- EOG
ments of the head or body
Vergence movements Nonconjugate movements of the 1-15 — 6-15 Corneal Reflection
eye to maintain binocular vi- Photography
sion. Vergence movements are Photoelectric
smoother and slower than con- EOG
jugate pursuit and compensa-
tory movements
Torsional or rolling eye Involuntary movements around — — — EOG
movements the line of gaze that compen- Photography
sate for the displacement of the Photoelectric
visual vertical
Minature eye move- Tiny involuntary movements that less than — — Contact lens
ments occur during periods of fixafixation 1 Corneal reflection
These methods have been classi-
fied into these categories
(continued)
Table 9.2. Continued
Size
(degrees Latency Speed Possible
Type Description of arc) (ms) (degrees/s) Recording Methods
Flicks Sharp saccadic movements

Drifts Slow movements between flicks
Tremor Rapid, oscillating movements
Nystagmoid Eye movements of an oscillating EOG
or unstable nature classified Photoelectric
into three categories
Ocular or Movement of the eyes in trying to
optokinetic follow a nonhomogeneous field
that is continuously moving
past the observer
Vestibular Compensating movements to
overcome problems due to im-
pairment of vestibular nerve
Spontaneous or cen- Occurs when the gaze is directed
tral nystagmus peripherally and is usually a
sign of impairment of the cen-
tral visual and vestibular path-
ways
Intraocular move- Pupillary reflex contraction to Photography
ments change in illumination
Source: From B. T. Tursky, 1974, "Recording of human eye movement." In R. F. Thompson and M. M. Paterson (Eds.), Bioelectric recording techniques, New York: Academic
Press. Reprinted by permission.
Note: — indicates not applicable.
ing a permanent record of the movements and position for further anal-
ysis. A movie camera provides a record of the eyes, but as with pupillary
responses, the amount of data to be scored is very large and almost
prohibitive. Thus, a number of alternate methods have been developed.
We will briefly describe these techniques here; the interested reader
should consult Young and Sheena (1975) or Stern and Dunham (1990)
for more detailed descriptions.
Corneal Reflection
One traditional method is the corneal reflection method. The front surface
of the cornea acts as a convex mirror to reflect light. Eye position deter-
mines the position of the corneal reflection. The reflected light is imaged
through a lens onto film, video equipment, or other light-sensitive de-
vices, such as a photo cell. One disadvantage of this device is that it
requires the subject's head to remain in a stable position or that head
position be calculated for each measurement to ensure an accurate mea-
surement.
TV Scanning and Other Techniques

Other techniques require scanning of the eye with a television camera.
Through the use of photodetectors, the boundary between the iris and
the sclera (the limbus) can be detected and the position inferred. Most
limbus-boundary techniques, as these are referred to, use infrared light
for better illumination of the eye. Another method includes fitting the
subject with a contact lens that fits tightly and moves with the eye. Small
mirrors are ground into the lens and reflect light onto a recording device.
Although contact lenses are one of the most accurate means of eye move-
ment recording, they may impose some discomfort and danger to the
subject. Other techniques are also being developed, such as those that
use fiber optics. Recently, with the incorporation of online computing
systems, video methods of eye movement tracking are becoming more
realistic. However, both online video equipment and other techniques
with one exception, are beyond the means of most psychophysiological
laboratories. The one exception, and the only technique that does not
interfere with normal vision, is electrooculography.
Electrooculography
Electrooculography (EOG) is a method of recording eye movements and

position that utilizes equipment commonly found in psychophysiological
laboratories. As the name implies, EOG is an electrical technique that
records potential differences from electrodes placed around the eyes, the
EYES 135
Figure 9.2. The eye as a dipole. Note the movement of the eye and the cor-
responding tracing on the recording system.
corned-retinal potential. This potential is seen as the result of the difference

in potential between the front and the back of the eyeball; that is, the
cornea remains 0.40 to 1.0 mV positive with respect to the retina. Thus,
the eye may be considered a battery which, as it rotates, carries with it
a potential field or dipole which can be measured by placing electrodes
on adjacent tissue. This relationship is shown in figure 9.2. This figure
shows that, as the eye moves, the potential at the electrode becomes more
positive or negative depending upon the direction of movement. EOG can
be used to record eye movements up to ± 70°. Typical accuracy is ±1.5-
2.0°.
Electrodes and Electrode Placement

The most commonly used electrodes for recording eye movements are
miniature type (11 mm) silver-silver chloride electrodes; several com-
mercial companies manufacture such electrodes. The electrodes are
placed as shown in figure 9.3. Horizontal eye movements are generally
recorded with an electrode placed at the outer edge of each eye (outer
canthi). It is important to place the electrodes as near to the eye as
possible, since the DC potential decreases as the electrodes are placed
farther from the eye. Vertical eye movements are recorded with the elec-
trodes above and below the eye. In placing the electrodes, careful align-
ment is required to eliminate any vertical component in the horizontal
measurement, and vice versa. The possibility of cross talk must be further
determined by having the subject move the eyes in a horizontal manner
and observing the vertical record, and by having the subject move the
eyes in a vertical manner and observing the horizontal record. Figure
9.4 shows a record of a subject instructed to move right and left and
then up and down. Notice the small amount of cross talk in the two
records.
Procedure
After the subject has been informed about the experimental procedure,
the electrodes are placed around the eyes. To insure proper conductance
and reduce drift, the skin should be lightly abraded and all excess facial
oils removed. Although alcohol is often used when electrodes are placed
on other parts of the body, it is suggested alcohol not be used for EOG
recordings because of the possible discomfort or damage to the eyes. A
cloth or cotton ball wet with water will serve to remove excess oils. Once
the electrode site is cleaned, silver-silver chloride electrodes are placed
on the skin following standard procedures. That is, when adhesive collars
are used, one side is attached to the electrode mount and the electrode
cup is filled with electrode paste. Once sufficient paste is applied to insure
complete filling, the electrode and collar are attached to the skin, and the
leads are connected to the preamplifier and amplifier.
Recording Equipment
The signal is amplified using a DC amplifier capable of reproducing volt-
ages in the range of 15 to 200 uV. As with any DC amplifier, the use of
Figure 9.3. Electrode placement for EOG recording of both horizontal and
vertical eye movements.
EYES 137
Figure 9.4. Horizontal and vertical eye movement records from a subject who
looked first right and then left, and then up and down.
short shielded cable and a high common-mode rejection preamplifier re-

duces noise. The frequency range of the amplifier depends on the partic-
ular eye movement response under consideration. For example, if only
eye position is of interest, then it is only necessary to have an amplifier
capable of reproducing accurately in the range of 0-15 Hz. If one wishes
more exact reproduction of the amplitude of a saccade, then a frequency
response of more than 100 Hz is necessary. If one is interested only in
the number of saccades, then an AC recording is appropriate, and this
eliminates any problem of drift.
Typical Recordings
Figure 9.5 shows the difference between DC and AC recording of eye
movement. Notice that from the DC chart one may derive eye position,
but that only the occurrence of a saccade, and not absolute position, can
be determined from the AC recording. Figure 9.6 shows horizontal and
vertical eye movement recording from a subject who moved the eyes in
a circle.
Figure 9.5. DC and AC records of vertical eye movements.
138 PSYCHOPHYSIOLOGY OF SPECIFIC ORGANS AND S Y S T E M S

Figure 9.6. Eye movement records obtained while an individual's eyes traced
a circle.
Common Problems
There are relatively few problems with the technique just described. The
major problem is slow DC drift of the baseline over time. New amplifiers
and electrodes make this less of a problem today than previously. Another
potential problem, depending upon the nature of the experiment, would
be for the subject to change the gaze by turning the head. That is, the
subject's eye position would not have changed in relation to the head
and thus would not be recorded, but the position would have changed
in relation to the environment. This can be prevented by the use of a
device such as a chin rest or bite board for keeping the head stable. The
chief source of bioelectrical artifacts is muscle potentials. It should also
be noted that the potential difference between the retina and the cornea
may vary with such factors as light adaptation, diurnal variations, the
degree of alertness, and the sex of the subject. These problems are in-
cluded here to alert the researcher to the complexity of the area and the
difficulty of conducting well-controlled studies. For sophisticated appli-
cations of eye-movement techniques, the researcher should consult a
more detailed reference (Oster & Stern, 1980; Stern & Dunham, 1990;
Young & Sheena, 1975).

The analysis and quantification of eye movement data vary with the aim
of the study. If eye movement is used as a control—as might be the case
in an EEC alpha biofeedback study—then one simply looks at the record
to determine if the eyes were moving at a particular time. One can like-
wise look at the record alone to determine saccadic movement and du-
ration during a reading task. If a more precise determination of the ori-
entation of the eyes is required, it is necessary to have performed initial
calibrations. That is, the subject should be instructed to look straight
ahead, up, down, left, and right at stimuli a known distance away, with
each of these points being marked on the EOG record. It is also possible
to combine the outputs of the horizontal and vertical traces on a plotter
EYES 139
or screen to obtain a representation of where the eyes are looking at any
one time. It should be pointed out that the combination of the vertical
and horizontal traces may introduce errors that make pinpoint accuracy
impossible. These errors may be somewhat reduced by a method of vector
EOG recording (Uenoyama, Uenoyama, and linuma, 1964). As with any
electrophysiological variable, a computer may be used for pattern rec-
ognition; the computer then determine the presence, duration, and di-
rection of saccades, or a measure of eye position. Oster and Stern (1980)
and Stern and Dunham (1990) describe in some detail the program and
hardware used in their studies of eye movement during reading; the in-
terested researcher should consult these reports.
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Beatty, J., & Wagoner, B. L. (1978). Pupillometric signs of the brain activation
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Dement, W., & Kleitman, N. (1957). Relation of eye movement during sleep
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Experimental Psychology, 53, 339-346.
Duke, J. (1968). Lateral eye movement behavior. Journal of General Psychol-
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Ehrlichman, H., & Weinberger, A. (1978). Lateral eye movements and hemi-
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EYES 141
10
Respiratory System
Respiration refers to the process by which oxygen is supplied to cells and

carbon dioxide is removed. The aspects of respiration that psychophysi-
ologists usually measure are breathing rate and amplitude, the latter
being a measure of the depth of breathing. The normal rate of respiration
in humans is about 12-16 breaths per minute, and the usual depth (tidal
volume, or total volume of each breath) is about 400-500 ml for healthy
adults. Breathing amplitude can be measured either directly (the true
volume of the lungs) or indirectly (using a measure such as the circum-
ference of the chest). Some methods of direct volume measurement also
allow one to assess the nature and amounts of gases like C02 that are
being expired from the lungs. In addition, one can measure aspects of
the respiratory cycle such as the inspiratory duty cycle (also called the
inspiration fraction) which is the ratio of inspiratory duration to the total
respiratory cycle duration. Another measure that provides important in-
formation about the influence of the central nervous system on respira-
tion is the mean inspiratory flow, which appears to reflect central inspir-
atory drive and which can be quantified using the ratio of tidal volume
to inspiratory duration (Wientjes, 1992).
Relatively few studies in the psychophysiological literature focus on
respiration as the response of primary interest, although respiratory ma-
nipulations and measures have played a prominent role in studies of anx-
iety disorders and relaxation (Clark & Hirschman, 1990; Fried, 1993;
Papp, Klein, & Gorman, 1993), asthma (e.g., Grossman & Wientjes, 1989;
Isenberg, Lehrer, & Hochron, 1992), emotion and stress (Boiten, Frijda,
& Wientjes, 1994; Grossman, 1983), and speech (Winkworth, Davis, Ellis,
& Adams, 1994). In fact, Wientjes (1992) has suggested that technological
advances make respiration measures more useful for psychophysiologists
because newer measures provide more information about respiratory func-
tion while also being less invasive than older techniques.
142
When psychophysiologists do record respiration, it is often used as a
check for possible artifacts in other response measures caused either by
breathing irregularities or by changes in breathing due to an experimental
manipulation that might confound (i.e., systematically alter) the measure
of interest, such as heart rate. A deep breath, intentional or not, can often
bring about a greater change in autonomic nervous system function than
will manipulation of the independent variable. For example, Stern and
Anschel (1968) had subjects take four different types of breaths and re-
corded the effects on finger pulse volume, heart rate, and skin resistance.
The types of breaths taken are shown in figure 10.1, and the effects on
various response measures are shown in figure 10.2. The extent of the
ANS disturbance varied directly with the depth of the inspiration, with
deeper breaths leading to a decrease in skin resistance, an increase fol-
lowed by a decrease in heart rate, and vasoconstriction in the finger.
The effect of respiration on heart rate has long been of interest to
psychophysiologists. Respiratory sinus arrhythmia (RSA) was first de-
scribed by Ludwig in 1847 (see Daly, 1985), and refers to the rhythmic
increases and decreases in heart rate produced by normal respiration in
many subjects. As a person inhales, the heart rate increases; as the per-
son exhales, the heart rate decreases (see figure 12.5). Respiratory sinus
arrhythmia appears to arise from both afferent connections from the
Figure 10.1. Mean relative amplitude of four types of breaths. The graphs are
schematic depictions drawn using only the onset, peak, and offset values
derived from the original records (which is why they are triangular in shape).
For a typical respiratory record, see figure 10.4. Redrawn with permission
from R. M. Stern and C. Anschel, 1968, "Deep inspirations as stimuli for
responses of the autonomic nervous system," Psychophysiology, 5, 132-141.
R E S P I R A T O R Y SYSTEM 143
Figure 10.2. Mean group decrease in skin resistance (facing page, top panel),
heart rate response (facing page, bottom panel), and finger pulse volume
(above) to four respiratory stimuli. Redrawn with permission from R. M. Stern
and C. Anschel, 1968, "Deep inspirations as stimuli for responses of the au-
tonomic nervous system," Psychophysiology, 5, 132-141.
lungs to the central nervous system (CNS), and from CNS respiratory
rhythm generators (Berntson, Cacioppo, & Quigley, 1993; deBurghDaly,
1985). Respiratory sinus arrhythmia under many circumstances provides
a reasonable estimate of the effects of the parasympathetic nervous sys-
tem on the heart, and is particularly useful to the psychophysiologist
because RSA is measured noninvasively. For additional information, see
chapter 12.
Physiological Basis
Respiration is modified by both the central nervous system and the au-
tonomic nervous system, particularly the parasympathetic branch. Res-
piratory centers in the medulla and pons contain respiratory generator
neurons which spontaneously fire bursts of action potentials that initiate
inspiration. These brainstem areas have connections to the cortex, the
hypothalamus, and other parts of the brainstem. In addition, respiration
is also modified by several autonomic reflexes arising from the lungs,
upper airways, heart, and blood vessels. For example, if the lungs become
overly inflated during inspiration, stretch receptors transmit impulses
through the vagus nerve to the brainstem respiratory centers, which then
inhibit inspiration and prevent further overinflation of the lungs. This
reflex, called the Hering-Breuer reflex, does not operate during normal
respiration, but appears to protect against overinflation.
Respiratory activity is highly responsive to changes in the concentra-
tions of carbon dioxide and oxygen in the blood. Chemosensitive areas
in the brainstem that are part of the respiratory center are directly stim-
ulated by an increase of hydrogen ions in the cerebrospinal fluid, the
concentration of which is largely determined by the amount of carbon
dioxide in the blood. Although the concentration of oxygen in the blood
has no direct effect on the respiratory centers, it is sensed by peripheral
chemosensitive areas near the large vessels of the heart. Increases in
carbon dioxide sensed by the central chemoreceptors and/or decreases in
oxygen sensed by the peripheral chemoreceptors lead to the transmission
of impulses to the respiratory center and initiation of inspiration. Inter-
estingly, the level of carbon dioxide is much more important for the reg-
ulation of breathing than the level of oxygen. For additional information
on the physiological basis of respiration, see Guyton and Hall (1996).
The mechanical changes in the thorax that accompany inspiration
and expiration are particularly important for the psychophysiologist be-
cause they are noninvasively measurable aspects of breathing. Figure
10.3 illustrates some of the differences in size and shape of the thorax
that occur during inspiration and expiration. Just before inspiration be-
Figure 10.3. Depiction of the ribcage and diaphragm during expiration (left)
and inspiration (right). Notice that during inspiration the ribcage is much
larger both because the ribs have been lifted up and out and because the
diaphragm is more flattened. Redrawn with permission from A. C. Guyton
and J. E. Hall, 1996, Textbook of medical physiology (9th ed.), Philadelphia:
W. B. Saunders.

gins, the diaphragm is relaxed, and in this state, it forms a dome-shaped
structure that juts up into the area below the rib cage. Also during this
state, some of the muscles (the external intercostals) that attach to ad-
jacent ribs are relaxed, and the ribs will therefore slant downward and
forward from their attachments at the spine. When inspiration is initi-
ated, the diaphragm contracts and flattens, which creates a negative pres-
sure in the thorax which forces the lungs open. In addition, the external
intercostal muscles contract; this raises the ribs so that they are more
nearly perpendicular with the spine, and also pulls the sternum forward.
This action causes not only more negative pressure inside the thorax,
but also enlarges the circumference of the thorax during inspiration. It
is this larger inspiratory circumference that can be measured and used
by the psychophysiologist to indicate the phases of respiration. Expiration
generally occurs mostly by passive recoil of the chest against the negative
pressure created during inspiration. Thus, during expiration, the dia-
phragm and external intercostal muscles relax, which allows the ribcage
to fall down and back toward the spine again. Contraction of the abdom-
inal muscles and activation of the internal intercostal muscles can be
used for more active expiration, but these more active processes are not
typical for normal, quiet breathing and generally are recruited only dur-
ing heavy breathing or disease states such as emphysema.
Recording Procedures
Five methods of recording respiratory variables will be briefly described:

spirometry, the air-pressure pneumograph, impedance pneumography, air
temperature, and respiratory inductive plethysmography. We will also de-
scribe one other method, the strain gauge, in greater detail because it
has been (and is likely to continue to be) used often in psychophysiolog-
ical laboratories.
Spirometry
A spirometer is a device that measures the volume of air expired with
each breath. The subject either breathes through a wide tube, or into a
face mask attached to tubing leading to the spirometer. With a nose clip
in place to prevent air escaping from the nose, the subject breathes into
the spirometer and the volume of air displaced by a breath is measured.
Thus, respiratory or tidal volume can be measured directly, and respi-
ratory rate can be derived from the changes in the amount of carbon
dioxide that occur over the breathing cycle. Because the nose clip, and
face mask or tubing make normal breathing difficult, and also provide
resistance to breathing, measurements made with a spirometer are typ-
ically considered too intrusive to be used in most psychophysiological
RESPIRATORY SYSTEM 147

studies except for calibration of less invasive measures at one or two time
points during an experiment. An example of a study using spirometric
measurements is that of Miller and Wood (1994) on air-way reactivity
in asthmatic children in response to an emotionally evocative video.
Impedance Pneumogmphy
Impedance is the opposition to current in an AC circuit and it varies
with the volume of a conductor, among other things. The thorax of a
breathing subject can be viewed as a conductor when AC current is
applied across the chest. The current used to record the impedance pneu-
mograph is very high frequency (typically 20-30 kHz) and cannot harm
the subject. The impedance waveforms are produced by the variations in
voltage produced by transthoracic impedance changes which occur with
each breath. When the lungs are filled with air, impedance of the chest
is higher than when the lungs are not full.
As with other indirect respiratory methods, the measure of respiratory
amplitude obtained using impedance pneumography is relative and can-
not be compared across subjects although indirect methods such as this
do yield absolute respiration rate. Using an indirect method, respiratory
amplitude is a function of the changes in circumference of the thorax at
the specific location where the measuring device is placed. This means
that it is not possible to compare amplitude data across subjects or across
sessions due to differences in the placement location of the measuring
device and body composition of different subjects.
Because the cost of this method is considerably higher than that of
the strain gauge method (discussed later) and because the measures of
volume obtained are no more accurate than less expensive girth methods,
impedance pneumography has not seen wide use in psychophysiology.
Recently, suggestions have been made that impedance-derived respira-
tion may be recorded using the impedance cardiograph, which would
remove the need for separate electrodes and equipment for measuring
respiration if one is already measuring impedance cardiographic wave-
forms (Ernst, Litvack, Lozano, Cacioppo, and Berntson, 1999; see chapter
12 for more information on impedance cardiography).
Air Temperature
The air we inhale is cooler than the air we exhale. This difference in
temperature is the basis of one method of recording respiration. Either a
thermistor or a thermocouple is used to sense changes in air temperature,
typically in the vicinity of the subject's nose. Thermistors are semicon-
ductors that change resistance with changes in temperature. Thermocou-
ples change their voltage output with changes in temperature. Very small
thermistors and thermocouples—about the size of the head of a pin—are

available, thus making their placement relatively simple. The sensor is
typically clipped to the nose or taped in place with the sensor in front of
the nostril. The subject breathes in and inhales the relatively cool room
air across the sensor; exhaling, the subject warms the air.
Investigators usually report that the subject forgets about the device
on or near the nose after a few minutes. This ease of adjustment is im-
portant when measuring respiration, since anything that calls the sub-
ject's attention to her respiration makes it difficult for her to breathe
normally. There is the problem, however, that many subjects breathe
partially or even totally through their mouths, making such measures
problematic. In addition, any moisture on the temperature-sensitive de-
vice (as might happen after a sneeze) will alter the device's sensitivity to
temperature. A study, using an air temperature-based device, of the ef-
fects of paced respiration on feelings of anxiety was conducted by Clark
and Hirschman (1990).
Respiratory Inductive Plethysmography

A relatively newer measurement of respiratory function developed in the
late 1970s has begun to make its way into psychophysiological labora-
tories. This measure, called respiratory inductive plethysmography, uses
AC current to produce changes in self-inductance in coils of wire that
encircle the upper chest and abdomen. Self-inductance is the property of
a coil of wire that opposes any change in the current that is passing
through the coil (it is somewhat analogous to resistance). This property
manifests when the voltage applied to a coil changes rapidly, which in
turn, produces a change in the current in the coil. Alternations in the
direction of current flow (which occur at a frequency of 60 Hz in North
America) induce changing magnetic fields around the wires carrying this
alternating current. In turn, these changing magnetic fields act to oppose
the flow of current in the wire. Self-inductance is partly a function of the
cross-sectional area of the coil in which AC current is flowing. For mea-
suring respiration changes, AC current is passed through coils wrapped
around the abdomen and thorax, and self-inductance in the coils fluc-
tuates as the cross-sectional area of the thorax changes with each in-
halation and exhalation. These respiratory-related self-inductance
changes result in variations in voltage output which, when calibrated,
are proportional to lung tidal volume.
The measure relies on an assumption that the changes in thoracic
cross-sectional area are essentially a function of two components: the
volume of the ribcage, and the volume of the abdomen. A ribcage band
containing wires in a zig-zag configuration (the coil) is placed over the
sternum with the top of the band just below the axillae (the "arm pits").
An abdominal band, also wired in a zig-zag configuration, is placed be-
tween the lower ribs and the iliac crests (the anterior protrusions of the

hip bones). The voltage output of these two bands together reflects the
change in tidal volume, but only after these measures are first calibrated
to a spirometer-based volume.
Calibration, generally speaking, is a process whereby a measure with-
out an absolute reference is compared to an absolute reference such that
the relative measure can be reported in units of absolute magnitude. In
respiratory inductive plethysmography, calibration is performed by si-
multaneously recording changes in voltage in the abdominal and ribcage
coils along with changes in lung volume recorded by a spirometer. Then,
as long as the coils do not move during the experiment, the investigator
knows how a given change in the voltage output of the abdominal and
ribcage bands is related to lung volume. This calibration procedure makes
what would otherwise be a relative measure of respiratory amplitude a
reasonably good estimate of absolute volume.
Because the equipment is somewhat expensive and requires not only
the respiratory inductive plethysmograph but also a spirometer, this tech-
nology will likely continue to be used in only a few psychophysiological
laboratories. For more information about this technique, the interested
reader can refer to Cohn et al. (1982) and Chadha et al. (1982). An
example of a psychophysiological study using respiratory inductive pleth-
ysmography is found in Winkworth et al. (1994).
Strain Gauges
One of the most common, and cost-effective, approaches to measuring
respiration has been the use of a girth method to measure thoracic cir-
cumference (or girth) using a strain gauge. A strain gauge wraps around
the thorax and measures the degree of strain placed on the measuring
device as the circumference of the thorax increases. As a person inhales,
the thorax becomes larger, increasing the stretch on the strain gauge; as
the person exhales, the thorax becomes smaller, decreasing the stretch
on the gauge. As previously noted for impedance pneumography, girth
methods yield only relative measures of respiratory amplitude, and these
cannot be compared across subjects unless the amount of stretch is cal-
ibrated to a known volume of air displaced by the lungs. Even within a
session for a given subject, care must be taken that the strain gauge does
not move; if there is any such movement, measures of amplitude recorded
early in the session may differ from those recorded later in the same
session.
Transducer. One commonly used transducer is a length of silastic (syn-

thetic rubber) tubing filled with mercury. The tubing is placed around
the chest and a small current is passed through it. As the subject inhales,
the cross-sectional area of the mercury in the tube is made smaller,
thereby increasing the resistance in the tube. As the subject exhales, the

resistance decreases. In a mercury-in-silastic strain gauge, air bubbles
can develop in the gauge after several months even if they are not used.
Therefore, the continuity, or the presence of an uninterrupted circuit in
the gauge should always be checked with an ohmmeter before the gauge
is used. Gauges lacking continuity are worthless and must be discarded—
and they must be discarded properly because they contain mercury. For-
tunately, mercury strain gauges are relatively inexpensive.
Another popular strain gauge device relies on the properties of a pi-
ezoelectric crystal material that changes electrical potential when it is
deformed. Such crystals are typically placed in a protective plastic device
and the crystal is attached to an elastic strap placed around the subject.
Deformation of the crystal which occurs when the subject inhales is mea-
sured as a change in voltage.
Transducer Placement. The respiratory transducer is usually placed

around the chest somewhere between the nipple line and the base of the
sternum. There is, however, considerable variability as to the point of
maximum displacement of the chest or abdomen. It is suggested that the
subject take a few deep breaths and the gauge be placed accordingly.
Indeed, one problem with recording of respiration only at the abdomen
or at the chest is that subjects may change the pattern of breathing
within a recording session. For example, someone feeling relaxed may
breathe using predominantly abdominal movements, however, if the per-
son became anxious, then she may shift to breathing using primarily
chest movements. For these reasons, some investigators prefer to record
from two devices simultaneously, one over the chest and one over the
abdomen.
Recording Equipment. The mercury strain gauge is normally wired into

a Wheatstone bridge circuit (see Marshall-Goodell, Tassinary, & Cacioppo,
1990, for details) to obtain a DC voltage output proportional to the re-
sistance change produced by the change in size of the subject's chest
with each breath. The specialized respiration couplers that plug into poly-
graphs are bridge circuits. The signal obtained is relatively large, so a
great deal of amplification is not necessary. Piezoelectric devices do not
require a bridge circuit, because they produce voltages that can be seen
by the polygraph or a computer (although some amplification may be
necessary).
Procedure. After the subject is seated, the gauge is attached by being

stretched slightly and placed around the subject, with the silastic or pi-
ezoelectric part in front. For silastic devices, the wires will wrap around
the subject's back and they may be held together with an alligator clip
behind the subject's back. The gauge can be placed over shirts, blouses,
sweaters, and other garments that are not too bulky. The investigator
should check to make sure that the belt is tight enough to show deflec-
tions with each breath, but not so snug as to be uncomfortable for the
subject.
Typical Recording
A typical recording of respiration is shown in figure 10.4. Inspiration in

this illustration drove the pen upward, which is the convention for de-
picting respiratory traces.
Potential Problems
A problem that can occur in recording respiration is shown in figure
10.4. When the subject took a very deep breath, the pen went off the
recording paper. Obviously, quantification of this portion of the recording
would not be possible. These lost data would be no problem if the inves-
tigator was interested only in knowing when the subject took a deep
breath. However, if one wanted to measure the amplitude of such breaths
in the future, one could (a) re-zero the pen so that the point of maximum
exhalation on the recording which is almost always the same point, is
at the bottom edge of the paper, or (b) reduce the amount of amplification
of the signal. In general, respiratory signals collected directly by a com-
puter acquisition program will not have the limitations on range that
can be problematic with a polygraph's chart recorder; one must take care
however, to make sure that there is sufficient amplification to faithfully
reproduce the signal (i.e., so that even shallow breaths can be detected).
Movement of a strain gauge up or down the subject's chest or abdo-
men will cause what may look like changes in the amplitude of the
respiration record and/or a change in the base level (see figure 10.5). In
order to minimize such movement, silastic gauges may be taped to the
subject's shirt or skin, with one piece of tape placed vertically on each
side at the point where the wires pass into the gauge. Care should be
taken not to put tape on the gauge itself. For additional information on
the use of strain gauges for recording respiratory function, see Lorig &
Schwartz (1990).
Figure 10.4. Typical recording of respiration with a single deep breath. The
upper channel is a timing channel making ticks once per second.
Figure 10.5. Respiratory tracing with timing channel. Point A shows what
happens when the subject pulls on or touches the strain gauge. Point B shows
a section of the record when the gauge has slipped and loosened.
Respiration Rate
In determining respiration rate, one may score a paper record by hand
or use a computer to mark events in the respiratory record and then
compute respiration rate. When scoring by hand, the most accurate
method is to use a ruler or calipers to measure the distance between
several successive cycles (e.g., 30 s or 1 min of data on the polygraph
paper) and then divide the number of full cycles within the measured
area by the total distance measured. Finally, the researcher converts the
measure of the number of cycles/cm into units of breaths/minute using
the known chart speed of the polygraph record (e.g., 50 cm/min). This
permits the researcher to count how many cycles would occur on aver-
age within a 1-min period, even though one did not measure exactly 1
min. With this method, there is no problem of deciding whether or not
to include partial cycles at the beginning or end of a measurement epoch.
When scoring respiration using a computer program, it is important
to remember that the computer scoring is only as good as the program
and the quality of the data input to the program. Typically, respiration
rate can be calculated by marking events such as the onset of a breath
and the onset of the next breath and converting the duration of a full
respiratory cycle into a value of breaths/minute. Alternatively, other
places may be marked on the respiratory record, e.g., peaks, which may
be better for records where the peaks are more definitive and can be more
accurately marked than onsets. Also, it is a good idea to inspect any
marks placed by a computer program to see that onsets or peaks are
marked accurately. It is not a good idea to assume that the computer
is "perfect"; often computer-acquired respiration records are both
computer-scored as well as inspected by eye for accurate detection of
events in the record.

Respiration Amplitude
Only the rather invasive technique of spirometry (or calibration of an-
other technique like respiratory inductive plethysmography by spirom-
etry) yields absolute respiratory amplitude values, and these measures
are rather uncommon in most psychophysiological studies. For assessing
breathing amplitude using relative measures, one can simply measure
the amplitude of the inspiratory waveform (using the signal level of the
breath at its onset as the baseline) in millimeters and report all experi-
mental comparisons in these arbitrary units. Recall, however, that am-
plitudes should only be compared within session and within subjects un-
less calibration to an absolute volume has been performed. As with
respiratory rate, amplitude measures can be made by hand or using a
computer program. When making amplitude assessments by hand, it is
typical to measure the amplitude of a small number of cycles and com-
pute a mean for these chosen cycles. For example, one could take a mean
of five pre-stimulus respiratory cycles, and a mean of five poststimulus
respiratory cycles and compute a pre-stimulus to poststimulus change in
relative respiratory amplitude. When computing respiratory amplitude
using a computer, one can typically use the entire record (e.g., compare
the entire pre-stimulus period with the entire poststimulus period) be-
cause the computer does most of the difficult measurement work.
In addition to measuring amplitude, one can use respiratory amplitude
measures to detect very deep breaths or other respiratory irregularities
(e.g., a yawn or sneeze) so that subsequent disturbances in ANS response
measures can be discounted. One must decide how much deeper than
normal the respiration amplitude can be before causing disruptive effects
in other measures. As a rule of thumb, it is suggested that if the ampli-
tude of a respiratory cycle is twice as large as or greater than the previous
cycle, other psychophysiological responses that occur for at least the next
20 s should be considered "suspect" data.
Respiratory Events
Respiratory events such as inspiratory time or inspiratory duty cycle (the
ratio of inspiratory duration to the total breathing cycle) have been used
in some studies of respiratory activity. These measures may provide a
more complete picture of the workings of the respiratory system. For
example, Wientjes (1992) notes that both tidal volume and respiration
rate are multiply determined by other features of respiratory function.
Thus, respiration rate may increase because of an increase in the inspir-
atory rate, an increase in expiratory rate, or both. For processes that
selectively influence either inspiration or expiration, reporting only the
overall respiration rate may lead the investigator to miss a true change
in respiratory function. As an example of a process that leads to changes
Figure 10.6. Measures of potential interest in a typical respiratory cycle.
in both inspiratory and expiratory times, consider speaking. Speaking can

lead to a shortening of inspiratory time because we tend to make a brief,
large inhalation just before beginning to speak. In addition, we typically
lengthen expiratory time during speech as we exhale air in a slower,
more controlled fashion than during normal breathing at rest (Wink-
worth et al., 1994).
Respiratory events would be scored just as described for respiratory
amplitude, with particular points in the waveform marked either by hand
or using a computer program. As noted previously, when using a com-
puter to place event markers in the data, it is important to look at the
raw data to be sure that there are no abnormalities, and to be sure that
marks are being placed at reasonable locations by the computer. Respi-
ratory events that may be of interest to the psychophysiologist are de-
picted in figure 10.6.
References
Berntson, G. G., Cacioppo, J. T., & Quigley, K. S. (1993). Respiratory sinus
arrhythmia: Autonomic origins, physiological mechanisms, and psycho-
physiological implications. Psychophysiology, 30, 183-196.
Boiten, F. A., Frijda, N. H., & Wientjes, C. J. E. (1994). Emotions and respi-
ratory patterns: Review and critical analysis. International Journal ofPsy-
Carry, P. Y., Baconnier, P., Eberhard, A., Cotte, P., & Benchetrit, G. (1997).
Evaluation of respiratory inductive plethysmography: Accuracy for anal-
ysis of respiratory waveforms. Chest, I I I , 910-915.
Clark, D. M., & Hirschman, R. (1990). Effects of paced respiration on anxiety
reduction in a clinical population. Biofeedback and Self-Regulation, 15,
273-284.
Chadha, T. S., Watson, H., Birch, S., Jenouri, G. A., Schneider, A. W., Cohn,
M. A., & Sackner, M. A. (1982). Validation of respiratory inductive pleth-

ysmography using different calibration procedures. American Review of
Respiratory Diseases, 125, 644-649.
Cohn, M. A., Rao, A. S. V., Broudy, M., Birch, S., Watson, H., Atkins, N.,
Davis, B., Stott, F. D., & Sackner, M. A. (1982). The respiratory inductive
plethysmograph: A new non-invasive monitor of respiration. Bulletin Eu-
ropeen de Physiopathologie Respiratoire, 18, 643-658.
de Burgh Daly, M. (1985). Interactions between respiration and circulation.
In N. S. Cherniack and J. G. Widdicombe (Eds.), Handbook of Physiology:
The Respiratory System II (pp. 529-594). Bethesda, MD: American Phys-
iological Society.
Ernst, J. M., Litvack, D. A., Lozano, D., Cacioppo, J. T., & Berntson, G. G.
(1999). Impedance pneumography: Noise as signal in impedance car-
diography. Psychophysiology, 36, 333-338.
Fried, R. (1993). The psychology and physiology of breathing. New York: Ple-
num.
Grossman, P. (1983). Respiration, stress, and cardiovascular function. Psy-
Grossman, P., & Wientjes, C. T. E. (1989). Respiratory disorders: Asthma and
hyperventilation syndrome. In G. Turpin. (Ed.), Handbook of clinical psy-
chophysiology (pp. 519-554). Chichester: Wiley.
Guyton, A. C., & Hall, J. E. (1996). Textbook of medical physiology (9th ed.).
Philadelphia: W. B. Saunders.
Isenberg, S. A., Lehrer, P. M., & Hochron, S. (1992). The effects of suggestion
and emotional arousal on pulmonary function in asthma: A review and
a hypothesis regarding vagal mediation. Psychosomatic Medicine, 54,
192-216.
Lorig, T. S., & Schwartz, G. E. (1990). The pulmonary system. In J. T. Ca-
cioppo and L. G. Tassinary (Eds.), Principles ofpsychophysiology: Physical,
social, and inferential elements (pp. 580-598). New York: Cambridge Uni-
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(pp. 113-148). New York: Cambridge University Press.
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children: A cholinergically mediated confluence of pathways. Journal of
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of the autonomic nervous system. Psychophysiology, 5, 132-141.
Wientjes, C. J. E. (1992). Respiration in psychophysiology: Methods and ap-
plications. Biological Psychology, 34, 179-203.
Winkworth, A. L., Davis, P. J., Ellis, E., & Adams, R. D. (1994). Variability
and consistency in speech breathing during reading: Lung volumes,
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search, 37, 535-556.

11
Gastrointestinal Motility
Electrogastrography
The psychophysiology of the GI system is a relatively unexplored area.

This is surprising when one considers the landmark work of Wolf and
Wolff (1943) in which they described the secretory and motility changes
of their fistulated subject to various stress situations, and the recent work
of Muth, Koch, Stern, and Thayer (1999) who have demonstrated that
behavioral stressors, which influence autonomic and cardiovascular reac-
tivity, also influence gastric activity; there are however, few studies re-
ported in between. But the paucity of psychophysiology research on the
GI system is not so surprising when we consider the instrumentation and
measurement problems of obtaining data from far inside this constantly
changing many-meter-long system.
As a consequence of these problems, no psychophysiological studies
of absorption are known to the authors, and few studies of gastric acid
secretion have been conducted by psychophysiologists. However, several
studies of motor activity have been conducted, particularly in the more
easily accessible two ends of the GI tract, the esophagus and rectum, and
also in the stomach. In this chapter the major emphasis will be on the
motor activity of the stomach as measured with the noninvasive method
of electrogastrography.
Electrogastrography refers to the recording of electrogastrograms
(EGGs). Electrogastrograms reflect gastric myoelectrical activity as it is
recorded from the abdominal surface with cutaneous electrodes. EGGs
are sinusoidal waves recurring at a rate of 3 cycles per minute (cpm) in
healthy humans. This predominant frequency is usually discern able by
visual inspection of the signal, but computer analysis is essential for
quantitative study of EGG recordings. The stomach is also the source of
abnormally fast or slow usually dysrhythmic myoelectrical signals, the
tachygastrias and bradygastrias. Acute or chronic shifts from normal 3
cpm EGG signals to the gastric dysrhythmias are associated with a variety
157
of clinical syndromes and symptoms, particularly nausea. In contrast to
the abnormalities in frequency such as the gastric dysrhythmias, the
amplitude, duration, waveform, and wave propagation characteristics of
the EGG have been infrequently studied.
Psychophysiological and pathophysiological investigations of EGG
characteristics in health and disease have increased greatly since the first
edition of this book was published. The International EGG Society (IEGGS)
was established in 1995, and commercial companies are now selling EGG
hardware and software including ambulatory models.
Physiological Basis
The GI system extends from the mouth to the rectum and includes the
mouth, esophagus, stomach, small intestine, large intestine, and rectum.
The three functions of the GI system are movement of food through the
alimentary tract, secretion of substances that aid in digestion or protect
the alimentary tract, and absorption of the digestive end products.
The GI tract may be considered to be a series of muscular tubes that
have been modified to perform region-specific digestive functions, that is,
transit of food from esophagus to stomach, mixing and emptying of in-
gested foods from the stomach into the duodenum, and absorption of
micronutrients from the small intestine. Other specialized tubes (i.e., the
cecum; ascending, transverse, and descending colon; and rectum) con-
serve water, electrolytes, and nutrients and evacuate wastes. These func-
tions require exquisite control and integration of relevant neural, mus-
cular, and hormonal systems within the GI tract.
The purpose of this section is to describe the relationship of gastric
motor activity to gastric myoelectric activity and the EGG. We include
more information about the physiological basis of the EGG than we did
for the other psychophysiological measures included in this book because
we thought that less was known about EGG by most psychophysiologists.
Readers desiring additional information about the physiology of the GI
system are referred to Johnson, Christensen, Jacobsen, and Schultz
(1987), and for more information about EGG the reader is referred to
Stern, Koch, and Muth (2000).
Relationship between Gastric Myoelectric

Activity and Gastric Motor Activity
The gastric contractions that occur at 3 cpm during the mixing and
emptying of meals are the result of coordinated electromechanical cou-
pling of circular layer smooth muscle cells. What are the electrical and
mechanical events within the smooth muscle that underlie the mechan-
ical work performed by the stomach?
Figure 11.1. The stomach and its principal regions. Inset shows major layers
of gastric wall. Origin of gastric pacesetter potentials (pacemaker signals) in-
dicated by stippled region on gastric body. Reprinted with permission from
K. L. Koch, 1993, "Stomach." In M. M. Schuster (Ed.), Atlas of gastrointestinal
motility in health and disease, Baltimore: Williams & Wilkins.
Gastric Slow Waves Gastric slow waves are the electrical events that
control gastric contractions. The slow waves result from spontaneous
depolarizations of the longitudinal muscle in the region of the juncture
of the fundus and body on the greater curvature (see figure 11.1). The
outer layer of the circular muscle may participate in the genesis of slow
waves. From this region, named the pacemaker area, the depolarization
wavefront moves circumferentially and distally toward the distal antrum.
The normal slow-wave frequency in humans is 3 cpm. The slow wave
does not move into the fundic area, which is electrically silent. The slow
wave is a spontaneous event, sodium-mediated and omnipresent, and is
associated with very low amplitude contractile activity (You & Chey,
1984).
The slow-wave coordinates the frequency and propagation velocity of
gastric contractions. That is, the slow wave brings the circular muscle
GASTROINTESTINAL MOTILITY 159

layer near the point of depolarization, and if physical, neural, and/or
hormonal signals are appropriate for contraction, the depolarization
threshold is reached and circular muscle contraction occurs. Because
circular muscle contractions are linked with the slow wave, the circular
muscle contractions occur at the slow-wave frequency (3 cpm in hu-
mans) and the contractions propagate at the slow-wave velocity (0.8-4
cm/s). For these reasons the slow waves have also been called pacesetter
potentials and electrical control activity. Slow waves are considered myo-
genic phenomena, but extrinsic neural input may modulate the rhythm-
icity of depolarization.
Gastric Spike Potentials The electrical events underlying circular smooth

muscle contractions are plateau and spike potentials. Depolarization of
the circular muscle, in contrast to the longitudinal muscle, is very fast
(i.e., spikes). The spikes may or may not occur on plateau potentials,
which are associated with the slow wave. The spikes reflect fluxes of
calcium passing through the circular muscle membrane. Contractions of
the circular muscle may increase tone and/or intraluminal pressure, par-
ticularly if they form concentric ring contractions. Such strong contrac-
tions may be recorded with strain gauges, intraluminal pressure trans-
ducers, or perfused catheters.
In summary, gastric slow waves are present at all times and control
the frequency and propagation velocity of spike potentials (i.e., circular
muscle contractions) when the latter are elicited by the appropriate stim-
uli. Gastric slow waves and spike potentials are the myoelectric compo-
nents of gastric contractions. It is these contractions that perform the
work of mixing and emptying foodstuffs. Slow waves and spike potentials
from the stomach may be recorded from electrodes sewn to the serosa
(the outer surface of the stomach) or from electrodes applied to the gastric
mucosa (the inner surface of the stomach). Because slow waves occur
within a conducting medium (i.e., the body), they are also recorded with
fidelity from electrodes positioned on the skin (i.e., the EGG). Figures 11.2
and 11.3 shows gastric myoelectric activity recorded from serosal and
cutaneous electrodes during motor quiescence (figure 11.2) and during
gastric peristalsis (figure 11.3).
Relationship of the EGG to

Gastric Myoelectric Activity and
Gastric Motor Activity
EGG and Gastric Myoelectric Activity Nelsen and Kohatsu (1968) simul-
taneously recorded the electrical activity from electrodes implanted on
the serosal surface of the stomach and EGGs from 13 patients. They found
an excellent correspondence between the frequency of the signals ob-
Figure 11.2. Gastric myoelectrical activity during motor quiescence. Paceset-
ter potentials begin in the pacemaker area located in the proximal gastric
body along the greater curve as shown by the darkened area. Slow waves
spread circumferentially and distally from the pacemaker region and migrate
through the antrum (as shown by serosal electrodes, B, C, and D). The slow-
wave migration ends at the pylorus. As the slow wave dissolves in the ter-
minal antrum, another slow wave begins to migrate distally from the pace-
maker region. Thus, as shown in the figure, three slow waves will propagate
from proximal to distal stomach every 60 s, i.e., 3 cpm slow waves. As shown
at point A, the cutaneous electrogastrogram (EGG) reflects the dipole created
by the migrating slow wave.
tained from the EGG and the internal electrodes. Another comparison of
EGG and serosal recordings from dogs by Smout, van der Schee, and
Grashuis (1980b) indicated a perfect correspondence between the fre-
quency of the signals.
In an effort to study the relationship of the EGG to internal electrical
activity of the stomach without involving surgery, several investigators
have compared the EGG to simultaneously recorded mucosal signals. The
mucosal signals are obtained from swallowed electrodes (i.e., electrodes
inside the stomach). Hamilton, Bellahsene, Reichelderfer, Webster, and
Bass (1986) compared EGG and mucosal signals from 20 human subjects
during fasting, after ingesting milk, and in one case, during a period of
spontaneous dysrhythmia. They summarized their findings as follows:
We did find that the surface recordings were of similar visual form as
those obtained directly from the mucosa simultaneously. In addition,
G A S T R O I N T E S T I N A L MOTILITY 161
Figure 11.3. Gastric myoelectrical activity during gastric peristalsis. Action
potentials occur during gastric circular muscle contraction. The action po-
tentials are linked to the gastric slow waves or pacesetter potentials as shown
in the extracellular recordings from the serosal electrodes (B, C, and D). As
the slow wave linked with action potentials migrates distally along the gastric
body and antrum, one gastric peristaltic wave occurs, and one EGG wave is
recorded from the surface electrodes. During gastric peristaltic contractions,
the EGG amplitude is generally increased. Compare this figure with figure
11.2. Reprinted with permission from K. L. Koch, 1993, "Stomach." In M. M.
Schuster (Ed.), Atlas of gastrointestinal motility in health and disease, Baltimore:
Williams & Wilkins.
frequency analysis determined that the two simultaneously obtained

signals were of the same frequency. Finally when the rare arrhythmic
events occurred, they were detected in both the mucosal and cutaneous
signals. Therefore, the signal obtained from the skin does seem to ac-
curately reflect the BER [basic electrical rhythm] as measured directly
from the stomach mucosa (p. 37).
Mintchev, Ott, and Bowes (1997) made simultaneous serosal and EGG
recordings from dogs in whom they had created dysrhythmias by surgical
means. They reported that the EGG could be used to detect severely ab-
normal gastric myoelectric activity 93% of the time, and mild abnor-
malities 74% of the time.
EGG and Gastric Motor Activity Smallwood and his colleagues published
a number of studies (e.g., Smallwood, 1978; Smallwood & Brown, 1983)
in which they examined the frequency of the EGG and made many ad-
vances in techniques for analysis of the EGG signal. In some studies (e.g.,
Brown, Smallwood, Duthie, & Stoddard, 1975) they compared the EGG
signal with intragastric pressure recordings. Their findings were the same
as those from other laboratories—when contractions occurred, they oc-
curred at the same frequency as the EGG signals; and whereas the EGG
showed 3 cpm almost continuously for most subjects, contractions as
recorded with intragastric pressure instruments did not.
It should be noted that the simultaneous presence of 3-cpm EGG and
the absence of changes in intragastric pressure do not necessarily indicate
that the EGG is unrelated to contractions. The possibility exists that the
EGG is a more sensitive measure of gastric contractile activity than the
pressure-sensitive probes. That is, the EGG may reflect increases in elec-
trical activity (i.e., spike activity) during low-level contractile events that
do not alter gastric intraluminal pressure. In fact, Vantrappen, Hostein,
Janssens, Vanderweerd, and De Wever (1983) indicated that low-
amplitude 5-cpm motor activity is always present in the dog. In addition,
You and Chey (1984) have shown that in dogs the 5-cpm pacesetter
potentials correlated well with low-amplitude contractions recorded by
strain gauges sewn to serosa but correlated poorly with intraluminal
pressure changes.
From 1980 to the present, published reports have appeared that not
only suggest that the EGG provides information about frequency of con-
tractions but also, indeed, that the amplitude of the EGG is related to the
degree of contractile activity (Smout, 1980; Smout, van der Schee, &
Grashuis 1980a, 1980b). One of the major contribution of Smout and
his colleagues was to point out that the amplitude of the EGG increases
when a contraction occurs. They concluded that the pacesetter potential
and the second potential, which is related to contractions, are reflected
in the EGG. Abell, Tucker, and Malagelada (1985) conducted a study in
which they compared the EGG signal from healthy human subjects with
the electrical signal recorded from the mucosal surface of the stomach,
and intraluminal pressure. They summarized their findings as follows:
"Antral phasic pressure activity, when present, was accompanied by an
increase in amplitude and/or a change in shape of both the internal and
external EGG" (p. 86).
Koch and Stern (1985) reported a close correspondence between the
amplitude of EGG waves and the amplitude of peristaltic antral contrac-
tions observed during simultaneous EGG-fluoroscopy recordings in four
healthy subjects. Hamilton et al. (1986) reported that fluoroscopy re-
vealed contractions in the antrum that correlated with three- and four-
fold increases in amplitude of the EGG.
The relationship of the amplitude of EGG waves to contractions is
complex and not totally understood at this time. However, in addition to
the studies mentioned, there is considerable indirect evidence linking am-
plitude changes in the EGG with strength of contractile activity. For ex-
ample, in situations where increased contractile activity would be ex-
G A S T R O I N T E S T I N A L MOTILITY 163
pected (e.g., eating, after swallowing barium), EGG amplitude increases
(Hamilton et al., 1986; Jones & Jones, 1985; Koch, Stewart, & Stern,
1987). And in patients with diabetic gastroparesis, where one would
expect weak contractility activity, Hamilton et al. (1986) found no in-
crease in the amplitude of EGG after eating.
Can EGG amplitude alone be used to infer reliably the presence or
absence of GI contractions? Not at this time. It is possible that with im-
proved methods of measuring contractile activity we will find that all
myoelectric activity is accompanied by some contractile activity (see Mor-
gan, Schmalz, & Szurszewski, 1978; Vantrappen et al., 1983; You &
Chey, 1984) and that the amplitude of the EGG is related to the intensity
or strength of contractile activity. A significant question then becomes:
Can the amplitude of the EGG be used to determine whether the accom-
panying gastric contractile activity is of sufficient strength to do the mo-
tor work of the stomach, i.e., mixing and propelling?
Several investigators (e.g., Bruley, des Varannes, Mizrahi, Curran,
Kandasamy & Dubois, 1991; Dubois and Mizrahi, 1994) have been ex-
amining the possibility of using the EGG as an indirect measure of gastric
emptying. Chiloiro, Riezzo, Guerra, Reddy, and Giorgio (1994) simulta-
neously recorded gastric emptying using ultrasound and the power in
the normal 3-cpm EGG from healthy subjects. The correlations ranged
from 0.68 to 0.96. Other investigators (e.g., Chen, Richards & McCallum,
1993) have demonstrated a negative relationship between the presence
of dysrhythmias in the EGG and gastric emptying. And Bortolotti, Sarti,
Barara, and Brunelli (1990) have demonstrated the presence of tachy-
gastria in patients suffering from idiopathic gastroparesis, that is, patients
with severely delayed gastric emptying with no known cause.
In summary, the frequency of the EGG is identical to the frequency of
gastric pacesetter potentials recorded from the mucosal or serosal surface
of the stomach. There is no general agreement, however, on the inter-
pretation of the amplitude of the EGG. Indirect evidence from several
studies has demonstrated that amplitude increases during an increase in
contractile activity; however, the amplitude of the EGG alone cannot be
used to determine the presence or absence of contractions.
Recording Procedure
Electrodes
Silver-silver chloride electrodes should be used. The size is not important,
but the electrical stability is. Since a relatively small, very slowly varying
potential will be recorded, the electrodes should show little bias or, as
some refer to it, offset potential.
Electrode Placement
EGG recording is done with bipolar electrodes placed on the skin surface
usually over the antrum of the stomach, plus a reference electrode on
the right side of the subject's abdomen. The optimal recording sites will
depend on the nature of the signal desired: e.g., largest possible amplitude;
lowest artifact from EKG, respiration and subject movement; and the
position of the subject's internal organs, particularly the antrum of the
stomach and the diaphragm (Mirizzi & Scafoglieri, 1983). The exact
placement of the electrodes is not important if the frequency of the EGG
is what is of interest, and it usually is. In fact, the EGG can be recorded
from the subject's two wrists, but the amplitude will be low compared
to recordings from the abdomen because the wrist electrodes are far
from the source of the signal, namely, the gastric pacemaker in the
antrum. For good EGG recordings from most subjects we recommend
placing one active electrode on the subject's left side approximately 6
cm from the midline and just below the lowest rib. The second active
electrode should be placed on the midline just above the umbilicus. The
reference electrode may be placed anywhere on the right side of the
subject's abdomen.
Recording Equipment
The EGG is a relatively weak and slow biological signal. Therefore, a high-
quality recording system is needed that can amplify and process a 100-
800 (iV signal in the frequency range of 0.5-15.0 cpm. Polygraphs with
the appropriate amplifiers can record the EGG. In addition, several com-
panies make appropriate amplifiers and analog-to-digital boards that pre-
pare the EGG signal for analysis, with the proper software, on a PC.
Ambulatory EGG equipment is also available that stores the data on a
memory card for later analysis. Whatever equipment is used, it is im-
portant to have a visual display of the raw signal so that you can be
confident that a good signal is being processed.
Procedure
Subjects should be instructed what and when to eat prior to an EGG
recording session because the contents of the stomach will effect the EGG.
For most studies, we instruct subjects to fast for at least four hours prior
to the experimental session. In other studies subjects are asked to come
to the lab after an overnight fast and are given a standard small breakfast
such as two pieces of toast and juice. The best EGG recordings will be
obtained if the subject lies supine (that is, on the back) or reclines in a
comfortable chair. If the subject must move, ambulatory recording equip-

ment should be used with Fetrodes, which are miniature preamplifiers
that are attached to the electrodes and reduce movement artifacts. If
more than one recording system is available, EGGs may be taken simul-
taneously from multiple abdominal locations using a single reference
electrode. Prior to the attachment of the electrodes, the subject's skin
must be prepared by shaving away hair (if excessive), abrading, and
cleaning with alcohol.
Typical Recordings
Eating and Sham Feeding

A number of investigators have reported that eating a nutritive meal
increases the amplitude or power of normal 3-cpm activity and produces
a brief frequency decrease. Figure 11.4 shows a typical EGG recording
from a healthy subject prior to and after eating. Note the increase in the
amplitude and regularity of the normal 3-cpm signal after eating.
Stern, Crawford, Stewart, Vasey, and Koch (1989) have used a sham
feeding procedure to examine cephalic-vagal influences on the EGG. Fol-
lowing a 15-min baseline period, subjects were required to chew and
expectorate a hotdog and roll. After another 10-min baseline period,
subjects were given a second hotdog to eat normally. The effect on the
EGG of eating the hotdog was as expected: a large increase in the ampli-
tude of the 3-cpm EGG wave that lasted several minutes. The effect
on the EGG of sham feeding was an equally large but short-lasting
Figure 11.4. Effects of eating on the EGG. Note the increase in amplitude and
regularity of the 3 cpm activity after eating.

Figure 11.5. Running spectral analysis of the EGG of a subject who reported
that the experience of sham feeding was not disgusting. Note the low level
of activity at approximately 2.5 cpm before sham feeding and the increase in
power during sham feeding and during eating.
increase in the amplitude of the EGG. The EGG of a typical subject

from this study is depicted in figure 11.5 as a running spectral analysis.
The use of such analyses will be discussed in the section on Analysis
and Quantification: for now, however, note that frequency is plotted on
the X axis, time on the Y axis, and power is the third dimension that
seems to rise from the page. This type of plot is referred to as a pseudo-
three-dimensional (3-D) plot. It was of interest to note that two subjects
who reported after the session that the experience of chewing and
expectorating the hotdog was disgusting showed a decrease rather than
an increase in the amplitude of their EGG during sham feeding (see
figure 11.6).

Figure 11.6. Running spectral analysis of the EGG from a subject who re-
ported that the experience of sham feeding was disgusting. This subject
showed power at approximately 2.8 cpm before sham feeding and a decrease
during sham feed. The subject showed the typical increase in power during
eating. Reprinted with permission from R. M. Stern, H. E. Crawford, W. R.
Stewart, M. W. Vasey, & K. L. Koch, 1989, "Sham feeding. Cephalic-vagal
influences on gastric myoelectric activity," Digestive Diseases and Sciences, 34,
521-527.
Cold Pressor Jest

Stern, Vasey, Hu, and Koch (1991) examined the effects of the cold pres-
sor test on EGG activity. A subject who had recently eaten was asked to
put one hand into a container of ice water (4°C) for 1 min, take it out
for 15s, put it back for 1 min, and so on, for a total of 20 min. Figure
11.7 shows the running spectral analysis of the EGG from a typical sub-
ject from this study. As can be seen, there was a significant attenuation
of EGG 3-cpm activity starting at the point in time when the subject put
one hand in ice water. Tachygastria was not seen in response to the cold
pressor test, a procedure that induces pain in addition to stress.

Motion Sickness
In the first experiment that attempted to relate changes in gastric myo-
electric activity to the development of symptoms of motion sickness, Stern
et al. (1985) obtained EGGs from 21 healthy human subjects who were
seated within an optokinetic drum, the rotation of which produced vec-
tion (illusory self-motion). Fourteen subjects developed symptoms of mo-
tion sickness during rotation, and in each subject the EGG frequency
shifted from the normal 3 cpm to 4-9 cpm, indicating tachygastria. Fig-
ure 11.8 shows an example of the EGG recording of one of these subjects.
Note that the frequency of the subject's EGG was 3 cpm prior to drum
rotation, but it changed to 6 cpm (tachygastria) after about 4 min of
rotation. The subject reported nausea after 6 min and requested that the
drum be stopped after 11 min. In six of seven asymptomatic subjects, the
3-cpm EGG pattern was unchanged during rotation. Figure 11.9 shows
a portion of one of these EGG recordings. It was concluded from this and
several subsequent studies that the sensory mismatch created by the il-
lusory self-motion produced tachygastria and symptoms of motion sick-
ness in susceptible subjects.
Figure 11.7. Running spectral analysis of a subject who had eaten just prior
to putting one hand in ice water. Note the inhibition of 3 cpm power when
the subject put the hand in the cold water and the gradual recovery. Re-
printed with permission from R. M. Stern, M. W. Vasey, S. Hu, & K. L. Koch,
1991, "Effects of cold stress on gastric myoelectric activity," Journal of Gas-
trointestinal Activity, 3, 225-228.

Figure 11.8. (a) EGG activity recorded from upper, middle, and lower elec-
trodes (E1-E3) prior to drum rotation. The EGG frequency is 3 cpm. (b) EGG
from same subject after start of drum rotation. Note presence of tachygastria
(6 cpm). Tachygastria began at 4 min, and the subject reported nausea at 6
min. At 11 min, he requested that drum rotation be stopped. Reprinted with
permission from R. M. Stern, K. L. Koch, H. W. Leibowitz, I. Lindblad, C. Shu-
pert, & W. R. Stewart, 1985, "Tachygastria and motion sickness," Aviation,
Space and Environmental Medicine, 56, 1074-1077.
Common Problems
A quick deflection usually indicates that the subject moved, disturbing

the subject-electrode interface. A slow drift in one direction for a minute
or more usually indicates electrode or amplifier malfunction.
The most obvious thing to check if an EGG recording is flat or almost
flat is to see if there is sufficient amplification and a proper band-pass
filter. Keep in mind that you are recording microvolts not millivolts, and
if the subject has considerable fat between the stomach and the electrode,
the normally weak signal will be further attenuated. Also be aware that
if the subject has not eaten for several hours, the readings might show
very low amplitude EGG because the stomach is quiescent.
In some cases a strong respiratory signal will dominate the EGG. It is
not possible to filter out the respiratory signal because its frequency is
too close to that of the EGG. The extent to which respiration is seen in
an EGG record depends on how close the diaphragm of the subject is to
the stomach. This will vary from subject to subject and will also depend

on the subject's posture. The basic problem is that in some cases the
subject's stomach moves with each breath, producing the unwanted res-
piratory artifact in the EGG. The only sure way to know if an EGG signal
with a frequency at 10-15 cpm is being generated by the stomach or
intestines or by respiration is to record respiration separately.
Spectral Analysis
Spectral analysis is currently the most commonly used method of quan-
tifying the EGG signal. Spectral analysis typically uses the Fast Fourier
Transform (FFT) to convert a signal in the time domain into the fre-
quency domain using a series of coefficients describing the amplitudes
and phase relationships of its independent sinusoidal waveforms. This
transformation is analogous to a prism transforming white light into the
visible spectrum of colors. (See chapter 14 for more about FFT and signal
analysis.)
The output of a spectral analysis is the squared magnitude of the
Fourier transform and is typically graphed as a curve showing the
Figure 11.9. (a) EGG activity prior to drum rotation. The EGG frequency is 3
cpm. (b) The EGG from the same subject 7 min after start of drum rotation.
Subject reported no symptoms of motion sickness and the EGG frequency
remained 3 cpm during 15 min of drum rotation. Reprinted with permission
from R. M. Stern, K. L. Koch, H. W. Leibowitz, I. Lindblad, C. Shupert, & W. R.
Stewart, 1985, "Tachygastria and motion sickness," Aviation, Space and En-
vironmental Medicine, 56, 1074-1077.

strength, or power, of the frequencies into which the original signal can
be decomposed. Although power has a very specific meaning in mathe-
matics and physics, we may think of it as an index of the amplitude of
the sine waves of a particular frequency that would be required in order
to recreate the EGG record. In the analysis of EGG recordings we are
usually interested in the power within the following three frequency
bands: 0-2.25, 2.5-3.5, and 3.75-9.75. The exact cutoffs for these bands
vary from lab to lab. The first frequency band represents the often found
but poorly understood ultra slow rhythm referred to as bradygastria. The
second encompasses the normal electrical rhythm of the healthy human
gastric antrum (3 cpm). The third includes frequencies commonly asso-
ciated with nausea and is referred to as tachygastria when regular, and
gastric tachyarrhythmia when dysrhythmic.
Van der Schee, Smout, and Grashuis (1982) have described an exten-
sion of this method that uses running spectral analysis to depict EGG
data. Running spectral analysis, with overlapping power spectra dis-
played as a function of time, yields both frequency and time information.
The more conventional spectral analysis provides power only as a func-
tion of frequency, not time. With running spectral analysis, frequency,
power, and time can be depicted two-dimensionally either with a pseudo
3-D display or with a grayscale plot. Figures 11.5 and 11.6 show two
examples of running spectral analyses, and Figure 11.7 shows an addi-
tional example.
A brief description of the procedure used to convert raw EGG data to
a pseudo-3-D display follows. The first step in any quantification proce-
dure is to insure that quality data are being analyzed. Hence, time must
be taken to insure the quality of EGG recordings before complex analysis
procedures are undertaken. For single-channel recording for most sub-
jects see our recommendation in the section on Electrode Placement. The
amplifying and recording system should filter out signals below 0.5 cpm
and above 15 cpm. With these filter settings one can record ultra slow
rhythms (0.5-2.0 cpm) but still eliminate shifts in baseline due to DC
potentials. Frequencies higher than 30 cpm are filtered out to avoid dom-
ination of the gastric signal by EKG. Respiration can also obscure the
EGG but its frequency range falls near that of tachygastria and, rather
than remove it with analog filters at the time of the recording, it is pref-
erable to remove it later with more precise and flexible digital filters or
by using a separate respiration recording to select visually and exclude
data that contain respiration artifact.
From the amplifier the EGG signal goes to an analog-to-digital (A/D)
converter where it is digitized into a series of numerical values repre-
senting discrete voltage levels of the input signal. Thus the analog EGG
signal is converted to a digital time series that can then be subjected to
a wide range of analyses. A/D conversion units typically allow sampling
at a wide range of speeds. Given an EGG signal which has had frequencies

faster than 15 cpm or .25 Hz attenuated at the time of recording, a
sampling rate of at least 1 Hz is required. We recommend a sampling
rate of 4.267 Hz, which yields 256 data points per minute.
Once the EGG signal has been digitized, it must be preprocessed in
order to meet the assumptions of spectral analysis. Because the dominant
frequency in the EGG is relatively slow, we recommend the use of at least
4-min data windows. Thus, at 4.267 samples per second, a 4-min seg-
ment would be composed of 1,024 data points. It is generally desirable
to center data for spectral analysis around a mean of zero. This is easily
accomplished by subtracting the mean of the segment from each data
point. Additionally, the EGG is likely to contain some very slow compo-
nents that reflect a shifting baseline or other undetermined factors. Such
extremely low frequency shifts and simple linear trends should be re-
moved to provide a clean spectrum. A high-pass digital filter which at-
tenuates frequencies below .01 Hz will accomplish such trend removal.
One can also remove simple linear trends by fitting a least-squares re-
gression line and subtracting it from the data segment.
After preprocessing, the data segment is Fourier transformed and the
spectral density estimates calculated. In order to produce a running spec-
tral analysis, one should overlap consecutive data segments by, for ex-
ample, 75%. In other words, segment 1 includes minutes 1-4, segment
2 includes minutes 2-5, etc. Thus, 1 min of new information is provided
in each consecutive power spectrum. These overlapping power spectra
can be plotted in a pseudo-3-D fashion to allow easy viewing of changes
in power at various frequencies as a function of time (see figures 11.5,
11.6, and 11.7).
While such running spectral analyses do provide a useful way to view
frequency and power changes over time, it is important to note that
transient changes in the EGG may go unnoticed if they are small. If such
transient changes are large enough they will appear but only as a grad-
ual change with the peak in spectral density appearing in the pseudo-3-D
display several minutes after it occurred in real time. Thus, running spec-
tral analysis may not be appropriate for experiments in which very short
duration stimulus induced changes are expected. For such cases adaptive
spectral analysis methods are recommended; see, for example, Lin and
Chen (1994). Similarly, spectral analysis is useful only when the EGG
signal contains a significant amount of cyclical activity. This is usually
the case for 3-cpm activity, but some gastric phenomena occur intermit-
tently and may not appear in a spectral plot. For more appropriate meth-
ods of analysis for quantification of very brief duration, intermittent phe-
nomena, see, for example, Holzl, Loffler, and Muller's (1985) discussion
of so-called zoom FFTs, and Lin and Chen's (1994) discussion of the
exponential distribution.
Once spectral power estimates have been calculated, data reduction is
usually performed. Our lab typically calculates percentage of total power

estimates for the bradygastria, normal 3 cpm, and the tachygastria bands
previously mentioned. This is accomplished by summing the power es-
timates for a given band, dividing this sum by the total spectral power,
and multiplying by 100%. However, when looking at changes in per-
centage of power as a function of, for example, exposure to a stressor,
the calculations of percentage of 3-cpm activity and tachygastria can be
grossly distorted if there is a large change in the amount of bradygastria.
In such cases we recommend calculating the ratio of power in the normal
3-cpm and tachygastria bands before and after presentation of the stim-
ulus. Various other methods of data reduction can be found in the lit-
erature. Chen and McCallum (1991) have proposed focusing on the fre-
quency peak which contains the dominant power of the EGG signal. It is
possible to analyze both changes in the power of this peak and changes
in the frequency of the dominant peak. Smout, Jebbink, and Samson
(1994) have suggested an instability factor, which is essentially a mea-
sure of the frequency variability of this dominant peak. The best method
for reducing the EGG frequency and power data from an EFT is still un-
settled. In most cases it will depend on the question to be answered and
all of the previously mentioned methods have merit.
Future Directions
EGG, because of its noninvasive nature, will continue to aid basic re-
searchers in their quest for additional information about gastric myo-
electric activity, gastric motility, and their relationship in normal and
pathophysiological conditions. Applied research using the EGG by gastro-
enterologists is increasing rapidly largely due to the ease and reliability
of its use in detecting gastric dysrhythmias and the recently established
relationship between gastric dysrhythmias and upper GI disorders in-
cluding delayed gastric emptying and nausea. Much of this research has
been supported by pharmaceutical companies, and we anticipate that this
will continue. A related exciting new area that requires EGG recording
in order to assess results is electrical pacing of the stomach. Research is
currently being carried out on dogs and on a small number of humans
in cases of gastric paresis where the stomach has ceased contracting and
no drugs are helpful. A less dramatic but related area of research that
we have planned is biofeedback of EGG for individuals with gastric dys-
rhythmias in an effort to restore normal 3-cpm activity and thereby re-
lieve nausea. The recent increase in the use of the EGG by gastroenter-
ologists has brought with it refinements in both hardware and software,
including ambulatory units that have flown on space shuttle flights. We
predict that with the availability of this new equipment additional psy-
chophysiologists will soon be using the EGG to study the effects of stress
and emotions on gastric activity.

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12
Cardiovascular System
Heart Rate; Cardiac Output; and Blood Pressure,
Volume, and Flow
The heart is a muscle, referred to as the myocardium, which begins func-

tioning within the fourth week of embryonic development and continues
to beat 3 billion to 4 billion times throughout life. About the size of a
fist, the heart weighs less than a pound and contracts about 60-75 times
a minute. As it beats, the heart moves blood to various organs, including
the lungs (pulmonary circulation), the heart (coronary circulation), and
the rest of the body (systemic circulation). Psychophysiologists have long
been interested in the functioning of the heart and circulation and have
focused their attention on such measures as heart rate, blood pressure,
blood volume, and blood flow. This chapter will discuss the recording and
analysis of these and related measures, after providing a brief overview
of cardiovascular physiology.
Physiological Basis
The heart consists of a left and right pump, each with two chambers,
the atrium and the ventricle. The right pump supplies blood to the pul-
monary system, and the left pump supplies oxygenated blood to the rest
of the body. Venous blood, low in oxygen, returns to the heart and enters
the right atrium. Contraction of the right atrium forces blood into the
right ventricle. The right ventricle then contracts, forcing blood into the
lungs through the pulmonary artery. Blood oxygenated in the lungs enters
the left side of the heart. The blood is pumped from the left atrium into
the left ventricle. From there, the contraction, or systolic action of the
heart muscle, forces the blood into the aorta, the major artery of the
heart, for distribution throughout the body. Distribution of blood through-
out the body is accomplished by a system of arteries, smaller vessels
178
referred to as arterioles, and capillaries. Blood returns to the heart through
the veins. At rest, the majority of the blood is in the veins, which act
as reservoirs.
The amount of blood pumped by the heart per unit time is referred to
as cardiac output. At rest, the cardiac output is approximately 5 liters per
minute, but this amount may increase 400% to 600% during extreme
exercise. There also is a change in the distribution of blood during ex-
ercise. For example, at rest, the muscles receive only 15% of the cardiac
output, with the brain receiving 14%, and the kidney and liver receiving
22% and 27%, respectively. However, with extreme exercise, muscles
may receive 75% of the total blood flow. This shift of blood flow during
exercise is controlled in part by the sympathetic nervous system (SNS).
Thus, sympathetic innervation causes vasoconstriction in some parts of
the body, which provides more blood for use at the muscles. In addition,
local metabolic effects of lowered oxygen at the muscles leads to vaso-
dilation in the vessels of the exercising muscles. These vasodilatory effects
appear to occur because of the release of humoral (fluid-borne) sub-
stances that are presumably released from the tissues that need additional
oxygen. In summary, humoral factors are released in response to the
local nutritional needs of the tissues of the body, whereas SNS innerva-
tion directs the overall distribution of blood.
The SNS shifts blood flow not only during exercise but also in response
to temperature changes of the body, postural changes, and strong emo-
tional responses such as fear. For example, if body temperature becomes
elevated, inhibition of the SNS produces a vasodilation of the arterioles
of the skin. This in turn increases the flow of warm blood to the skin,
with a resulting heat loss. When a person changes posture, blood is also
shunted to particular areas of the body to maintain blood pressure. When
a person stands from a seated position, a drop in blood pressure is reg-
istered at pressure sensors called baroreceptors in the chest and neck.
These sensors send a message to the brainstem, which activates the SNS
throughout the body and results in vasoconstriction in the peripheral
blood vessels which returns the blood pressure to prestanding levels. Fear
and other strong emotions can also result in increased sympathetically
mediated vasoconstriction, and this activation is specific to particular vas-
cular beds. For example, Anderson, Wallin, and Mark (1987) showed
that during a mental arithmetic task, activity increased in muscle sym-
pathetic nerves innervating the leg, but did not increase muscle sympa-
thetic outflow to the arm. Thus, sympathetic activity makes an important
contribution to the redistribution of blood flow when the body's needs
change, and this redistribution is directed very precisely to reduce blood
flow to certain vascular regions and provide additional blood flow to areas
with the greatest need. It should be pointed out that the parasympathetic
nervous system (PNS) is generally considered to have a relatively minor
effect on the blood vessels.
CARDIOVASCULAR SYSTEM 179

Vasoconstriction and vasodilation change the resistance to blood flow
through the vessels. This change in resistance directly affects both blood
flow and blood pressure. This relationship, the physiological analog to
Ohm's law, states that blood flow through a vein or artery equals the
difference in pressure between the two ends of the vessel divided by the
resistance to the flow of blood. Thus, the resistance of a constricted vessel
would be greater than that of a dilated one and a vessel with high resis-
tance will have slower blood flow and higher pressure than a vessel with
low resistance. This relationship holds not only for individual vessels but
also for the entire body, and may be written as follows:
Cardiac Output (amount of blood pumped per unit of time) =

Arterial Blood Pressure / Total Peripheral Resistance
(resistance of all vessels in the systemic circulation).
Electrical Activity of the Heart

The electrical activity of the heart as recorded at the surface of the skin
is measured by a technique known as electrocardiography. The printed
record is called the electrocardiogram or EKG. (The abbreviation EKG is
derived from the German spelling, but the abbreviation ECG is sometimes
also used.) A typical EKG is illustrated in figure 12.1. A careful reading
of an EKG can discern how electrical activity spreads across the muscle
of the heart. The sinoatrial node (S-A node) is a small strip of muscle
located in the upper part of the right atrium. It is at this point that the
initial impulse begins that triggers the contraction of the heart. The cells
of the S-A node are also referred to as pacemaker cells because of their
ability to originate an electrical impulse. The electrical impulse passes
from the S-A node through the atria to the atrioventricular node (A-V
node). With this passage, the atrial muscle depolarizes; this is represented
in the EKG by the P wave. Mechanical contraction of the atria follows
depolarization after a brief (ms) interval. At this point, the blood is being
passed into the ventricles. The impulse is delayed briefly (about 0.09 s)
in the A-V node, allowing time for the atrial contraction to produce com-
plete ventricular filling. It then passes through the common bundle of
His and into the ramifications of the Purkinje network. These fibers trans-
mit the impulse almost immediately through the ventricular system and
result in blood being pushed into the lungs and body. It is the depolari-
zation of the ventricles that produces the characteristic QRS complex so
predominant in the ECG. The ventricles then repolarize; this is repre-
sented by the T wave of the ECG. The precise relationships between each
component of the ECG are a valuable aid in determining disorders of the
heart, such as premature ventricular contractions, sinus bradycardia, fib-
rillation, and tachycardia. Some abnormal rhythms, or arrhythmias, are
Sequential electrical events Electrocardiographic
of the cardiac cycle representation
1 . I mpulse from the sinus node Not visible
2. Depolarization of the atria P wave
3. Depolarization of the A— V node Isoelectric
4. Repolarization of the atria Usually obscured by
the QRS complex
5. Depolarization of the ventricles QRS complex
a. intraventricular septum a. initial portion
b. right and left ventricles b. central and terminal portions
6. Activated state of the ventricles ST segment: isoelectric
immediately after depolarization
7. Repolarization of the ventricles T wave
8. After-potentials following repolar- U wave
ization of the ventricles
Figure 12.1. Prototypical EKG and electrophysiological events producing

characteristic features of the EKG. Redrawn with permission from R. Philips
and M. Feeney, 1973, Cardiac rhythms, Philadelphia: Saunders.
shown in figure 12.2. High heart rate (tachycardia) or low heart rate
(bradycardia) may be a function of numerous conditions, which include
pathology as well as factors related to diet, posture, exercise, emotional
excitement, and mental activity. The last two factors have been of par-
ticular interest to psychophysiologists.
Cardiac Responding
Early studies on the effects of stressors on the cardiovascular system sug-
gested that there was a strong and concerted action of the sympathetic
nervous system in response to potent stressors such as fear stimuli. This
systemwide increase in sympathetic activational effects on the cardiovas-
cular system produced concurrent increases in heart rate and blood pres-
sure as well as other "arousal-related" responses such as increased ac-
C A R D I O V A S C U L A R SYSTEM 181
Figure 12.2. Some arrhythmias apparent in the EKG.
tivity of the sweat glands (i.e., increased skin conductance) and increased
breathing rate. However, research with more mild to moderate stressors
has demonstrated much greater variety in the patterns of cardiovascular
responding, suggesting that the concept of a unitary "arousal" response
pattern is misleading. Beatrice and John Lacey and their colleagues
showed that so-called arousal responses in visceral organs controlled by
the autonomic nervous system did not always occur in concert with one
another, and were determined in large part by the situation or context
in which these responses occurred (Lacey, 1967; Lacey, Kagan, Lacey, &
Moss, 1963). The Laceys noted that on some occasions changes in the
heart rate and skin conductance were inversely related, so that heart
rate decreases were accompanied by skin conductance increases. Such a
pattern of differential responsiveness in two or more physiological systems
being controlled by the autonomic nervous system was referred to as
directional fractionation and has been observed extensively across physio-
logical systems. For a further discussion of these issues, see chapter 5.
Also as an offshoot of their work on visceral responses to sensory stimuli
and environmental stressors, the Laceys derived an intake-rejection hy-

pothesis of cardiac responding which suggested that cardiac deceleration
permits sensory information to be processed more effectively, whereas
cardiac acceleration promotes the rejection of or inattention to intense,
unpleasant, or painful sensory stimuli. This hypothesis is somewhat
counterintuitive because it proposes that a peripheral physiological sys-
tem can affect the central nervous system, rather than the other way
around. Later work by Obrist and colleagues questioned the role of heart
rate in producing an attentive state because pharmacological blockade of
the cardiac deceleration preceding a reaction time response did not pre-
vent the participant from producing a quick response (Obrist, Webb, Sut-
terer, & Howard, 1970). These data, however, do not preclude the pos-
sibility that changes in blood pressure, rather than heart rate, may
provide feedback to the central nervous system and either facilitate or
inhibit sensory processing. Indeed, neurophysiological evidence supports
the idea that blood pressure increases can blunt the perception of pain,
as well as inhibit sensorimotor activity and reflex action. The mere pres-
ence of feedback signals from the periphery to the central nervous system
does not necessarily mean that the feedback to the central nervous sys-
tem is sufficiently potent under normal physiological conditions to pro-
vide any useful information to the organism. Thus, we do not yet know
whether heart rate deceleration or blood pressure decreases are simply
outputs of an "attentive" nervous system or whether they may be caus-
ally related to altered attentional abilities.
Changes in the cardiovascular system are also linked to motor activity,
and bodily movement is typically accompanied by an increase in heart
rate. This phenomenon is known as cardiac-somatic coupling (Obrist,
1981, 1982). Movement demands blood flow to fulfill the metabolic needs
of skeletal muscle activity. This does not imply, however, that cardiac
changes occur only in response to changes in metabolic need, and indeed,
cardiac changes in excess of metabolic requirements occur frequently, a
phenomenon called additional heart rate (Turner, 1994). Preparatory
cardiac responses, especially to novel situational demands, appear to be
common and may represent an attempt by the body to be ready for
whatever challenge is forthcoming. After repeated presentations of a sit-
uational demand, the cardiac response is typically titrated more closely
to the actual metabolic needs of the demand; that is additional heart rates
are smaller with succeeding presentations of a demand.
One can see from figure 12.3 that the heart is innervated by both the
sympathetic and parasympathetic branches of the ANS. Activation of the
parasympathetic nervous system results in a decrease in heart rate,
whereas a decrease in parasympathetic activity results in an increase in
heart rate. Conversely, an increase in sympathetic activation of the heart
results in an increase in heart rate and the force of contraction of the
heart (contractility), whereas a decrease in sympathetic activation to the
heart decreases heart rate and contractility. Although the two branches
Figure 12.3. Central and peripheral nervous system input to the heart. This
figure illustrates both the parasympathetic (left side) and sympathetic (right
side) autonomic inputs to the heart. (Reprinted by permission from J. F.
Green, 1987, Fundamental cardiovascular and pulmonary physiology, 2nd ed.,
Philadelphia: Lea & Fibiger,
often act in a reciprocal fashion to control the heart such that activity
in one autonomic branch increases while activity in the other autonomic
branch decreases, this reciprocal relationship between the two branches
does not invariably characterize cardiac control (Berntson, Cacioppo, &
Quigley, 1991). Under some circumstances, activity in both autonomic
branches may increase (or decrease) concurrently (a coactivational
mode); in other cases, activity in one branch may change while activity
in the other branch does not (an uncoupled mode). See chapter 5 for
further discussion of this issue. Because basal heart rate as well as
changes in heart rate may arise from different combinations of autonomic
nervous system activity, heart rate alone can not reveal the underlying
activity of the ANS. For this reason, cardiovascular researchers interested
in understanding the contributions of the ANS to heart rate activity must
combine measures of autonomic effects on the heart with those of heart
rate.
Measures of Autonomic Effects

on the Heart
Respiratory Sinus Arrhythmia Respiratory sinus arrhythmia (RSA) is an
oscillation in heart period due to the respiratory cycle (Forges, 1986;
Berntson et al., 1997). Changes in RSA appear to reflect the activity of
the vagus nerve, and an increase in RSA is strongly positively correlated
with increases in parasympathetic influence on the heart. RSA is mea-
sured by assessing how much heart period changes from beat to beat.
One simple way it can be measured is by using the maximal difference
(in ms) between the heart period associated with inspiration and the
heart period associated with expiration (peak-valley method). During in-
spiration, heart period is shorter than during expiration and this differ-
ence in heart period across the respiratory cycle can index parasympa-
thetic changes. Fast Fourier Transform (FFT) or other methods (e.g., the
Forges and Bohrer method; Forges & Bohrer, 1990) can also assess the
degree of oscillation in heart period from beat to beat that falls within
the respiratory frequency. These techniques can provide similar results
to those using the peak-valley method (Grossman, van Beek, & Wientjes,
1990). For additional information on the physiological basis and mea-
surement of RSA, see Berntson et al. (1997) and Berntson, Cacioppo, &
Quigley (1993). For additional information on techniques such as FFT,
see chapter 14.
Pre-ejection Period One of the most promising estimates of sympathetic

effects on the heart currently seems to be the systolic time interval known
as the pre-ejection period (PEP). This is the time between the electrical
signal to the ventricles initiating contraction and the ejection of blood

from the left ventricle into the aorta. PEP can serve as an estimate of
sympathetic influences on the heart because it usually reflects changes
in cardiac contractility (or the force of contraction). PEP is negatively
correlated with sympathetic activity. One must use caution, however, in
making inferences about PEP because it also may change with large
hemodynamic alterations such as when a person changes posture (in
this case, the change in PEP is not related only to sympathetic influences
on the heart). PEP is measured using two devices simultaneously, the
EKG and impedance cardiography. The impedance cardiograph permits
a researcher to record when changes in the velocity of blood flow occur
in the thorax—in particular, those associated with the ejection of blood
flow into the aorta. For further information on the measurement and
sympathetic basis of PEP see Berntson et al. (1994), Newlin and Levenson
(1979), and Sherwood, et al. (1990).
Heart Rate or Heart Period
Recording Procedure
Although the electrical activity of the heart had been noted previously,
it was not until the start of the twentieth century that a device capable
of faithfully reproducing this activity was developed. This instrument was
developed in Holland by Willem Einthoven, who is the founder of modern
electrocardiography. Einthoven's device, the string galvanometer, contin-
ued in use until the advent of vacuum tube amplifiers in the 1930s and
1940s. Einthoven also experimented with other aspects of EKG work,
including various types of electrodes. He was later awarded the Nobel
Prize for this work.
Electrodes and Their Placement The first electrodes used for EKG recording
were buckets of saline solution in which the arms and legs were im-
mersed. Silver-silver chloride electrodes are commonly used today. Al-
though there exists a standardized system for electrode placement on the
limbs and chest utilized for medical diagnosis, all that is required for an
EKG of sufficient quality to measure heart rate is that two electrodes be
placed on the skin fairly far apart. Psychophysiologists use standard limb
leads designated as follows: I, one electrode on each arm; II, right arm
and left leg; III, left arm and left leg (see figure 12.4). An electrode place-
ment which is less sensitive to body movement than standard limb leads
entails placing electrodes on the torso with one electrode on the distal
end of the right collarbone and the other on the lower left rib cage. This
has been referred to as a modified Lead II placement because the resulting
EKG is very similar to that obtained by a Lead II recording made from
the limbs (figure 12.4). This placement minimizes artifacts due to move-
Figure 12.4. EKGs derived from standard limb leads I, II, and III and from a
modified Lead II. Leads I and III are less often used by psychophysiologists
because they result in a smaller R spike (see first and second channels). Lead
II and modified lead II (right collarbone and lower left rib) are more often
used by psychophysiologists because these leads produce a pronounced R
spike. These four leads were recorded simultaneously from the same person.
ment because the leads are placed on areas that are relatively free of fatty
tissue and muscle that may shift during movement. A Lead II or modified
Lead II placement is typical in the psychophysiology lab because it gen-
erally provides an ECG with a large ORS complex relative to the other
components (figure 12.4). One and only one large spike per cycle makes
quantification of heart rate much easier.
Recording Equipment The QRS complex picked up from the surface of the
skin is approximately 1 mV. Often, the signal is amplified, either by a
hardware amplifier or by software. In conjunction with the amplification
process, the heart signal can be further conditioned by the utilization of
filters, as discussed in chapters 3 and 14. When possible, it is better to
remove electrical noise from the recording environment, rather than filter
the signal to remove the effects of that noise. However, because the high-
est frequency of interest in the QRS complex is approximately 12 Hz, one
can set a high-pass filter at 30-35 Hz, which will both reduce the prob-
lem of 60-Hz electrical noise and remove many muscle artifacts. If one
wants simply to determine the distance between successive heart beats,
one can use an instrument called a cardiotachometer (or cardiotach);
this instrument is triggered by the R spike in the QRS complex of each
heart beat and gives an output in the form of heart rate that is calculated
from the preceding interbeat interval. Heart rate is displayed on the Y
axis; with each beat, a new calculation is performed and the new heart
rate displayed as a horizontal line on the cardiotach record (see bottom
panel of figure 12.5 for an example). A cardiotach display also reveals
the oscillations in heart rate referred to previously as RSA. Thus, the
figure shows that heart rate increases and decreases in a cyclical fashion
that is roughly coincident with respiratory inhalation and exhalation (see
figure 12.5). The EKG is generally recorded using an AC amplifier. In
this way, one obtains a more stable baseline than would be seen with
DC amplification, and this improves visualization of the signal for either
computer or cardiotachometer analyses.
Procedure What follows is the standard procedure for most psychophys-

iological experiments in which heart rate is measured. The subject is
seated in a comfortable chair and is given information about the exper-
iment. It is a good idea to describe briefly how heart rate is measured
and to emphasize that no electricity will be penetrating the subject
through the electrodes, but rather that the equipment measures the nat-
ural electrical activity of the body without harm. The skin may be cleaned
with alcohol on a gauze pad to remove dead skin and skin oils. When
standard silver-silver chloride electrodes are used, high conductivity elec-
trode gel is placed in the electrode cup and the electrode is attached to
the skin with an adhesive collar or tape. Disposable electrodes, which are
commonly used in psychophysiology laboratories, are gelled by the man-
ufacture and are ready to apply.
Typical Recordings and

Common Problems
Figure 12.4 depicts EKGs recorded from the three standard limb leads,
along with a modified Lead II placement. Since the voltage of the EKG
188 P S Y C H O P H Y S I O L O G Y OF SPECIFIC O R G A N S AND SYSTEMS

Figure 12.5. EKG, respiration, and cardiotach records. Note that the oscilla-
tory changes in heart rate which can be most easily seen in the bottom
channel coincide (albeit with a brief delay) with the respiratory changes in
the second channel. These shifts in interbeat interval are much less apparent
with the EKG depiction (top channel).
on the surface of the skin is relatively large, there are few problems when
recording. The most common problems are 60 Hz interference and move-
ment artifacts. However, these problems can be corrected without great
expense by careful choice and preparation of the electrode site and the
electrodes. An additional factor that becomes important if the researcher
employs a computer or cardiotachometer is that the QRS complexes must
be easily distinguishable electronically from one another and significantly
larger than the P and T waves. As previously noted and as seen in figure
12.4, limb lead II or the modified Lead II usually results in an EKG with
these desired characteristics. For additional information about recording
heart rate or period, consult Papillo & Shapiro (1990).

In the early days of psychophysiological research, the distance between
R waves on a paper record was measured with a ruler, and this measure
was either converted to heart rate or utilized directly as the measure for
analysis. Today most researchers use either a cardiotachometer or a com-
puter for direct analysis of the timing between beats. There are two com-
monly used measures of cardiac activity: heart rate and heart period (or
interbeat interval). Heart rate is defined as the number of beats per unit
time (usually in minutes). For example, a change in beats per minute
may be used to determine the effects of biofeedback training, with the
experimenter measuring the number of beats occurring in a 10-min
training period relative to the number of beats occurring in a 10-min
rest period. Interbeat interval or heart period is determined by measuring
the time between R waves (usually in msec). Note that heart rate is
simply the reciprocal of heart period. Thus, the timing of the heart beats
during 1 min can be reported as either 85 beats/min or 706 ms depend-
ing upon the unit of measure used. Because the time units are different
for heart period (ms) and heart rate (min), one can convert from one to
the other using the following:
Heart period (in ms) = 60000/heart rate (in bpm)
Heart rate (in bpm) = 60000/heart period (in ms)
Often, it has been assumed that the choice of cardiac metric or mea-
sure is not important. However, heart rate and heart period are not lin-
early related to each other. To illustrate this, figure 12.6 shows the re-
lationship between change in activity of the parasympathetic branch and
heart rate, and the relationship between parasympathetic change and
heart period based on data recorded from dogs. This figure shows that
while a given change in activity of an autonomic nerve will result in
approximately the same change in heart period for very different baseline
heart period values, the same is not true of heart rate. For example, an
increase in stimulation frequency of the parasympathetic nerve of 2 Hz
results in a change of 70-72 ms in heart period whether the resting
(baseline) heart period is 875 ms or 350 ms. However, when the dog's
heart is beating at a heart period of 875 ms (or 68.6 bpm) the change
in heart rate with a 2 Hz increment in parasympathetic activation is 5.1
bpm, whereas at a baseline heart period of 350 ms, the same 2 Hz change
in autonomic input to the heart results in a heart rate change of 29.2
bpm. The same relative linearity also tends to be true for sympathetic
Figure 12.6. Heart period and heart rate as a function of parasympathetic
stimulation frequency. Data used for illustration are from a study of vagus
nerve (parasympathetic) stimulation in dogs by Parker, Celler, Potter, & Mc-
Closkey, 1984. To illustrate the nonlinearity in change scores that would
appear as a result of using heart rate rather than heart period, a 2-Hz change
in stimulation from 2-4 Hz and from 16-18 Hz are shown for both heart
rate and heart period. Lower panel: Note that a 2-Hz increase in stimulation
changes heart period a similar amount regardless of whether the stimulation
began at 2 Hz or 16 Hz. Upper panel: In contrast, a 2-Hz increase in stim-
ulation of the vagus at 2 Hz changes heart rate by 29.2 bpm, whereas a 2-
Hz increase from 16 to 18 Hz only changes heart rate by 5.1 bpm. These
data are derived from the same subjects, but the transformation of heart
period (which is relatively linear with respect to the parasympathetic input)
to heart rate introduces a statistical artifact which alters the apparent changes
if one were to use heart rate as the metric of cardiac function.
impact on the heart period, and seems to be true for a variety of mam-
malian species, including humans (Berntson, Cacioppo, & Quigley, 1995;
Parker, Celler, Potter, & McCloskey, 1984). Therefore, the amount of
cardiac change that the psychophysiologist reports as a result of some
environmental or psychological manipulation will differ depending upon
the metric chosen to represent the change in cardiac function. Also note
that these effects will be most pronounced when baseline autonomic
changes are large. Thus, Berntson et al. (1995) have recommended that
heart period be used as the metric of choice when (a) the change in
cardiac function is likely to be a result of autonomic effects (e.g., for many
of the short-term cardiac responses seen in the psychophysiology labo-
ratory), and (b) when the changes in cardiac function are large, where
errors due to the nonlinear relationship between autonomic inputs and
heart rate will be significant.
In addition to the choice of cardiac measure, one must decide whether
to represent cardiac function by computing a mean value for cardiac
function over a number of beats (unit of analysis is cardiac time) or over
a period of time (unit of analysis is real time). Heart period permits anal-
ysis of cardiac function in either cardiac time or real time, whereas heart
rate can be represented appropriately only in real time (Berntson et al.,
1995; Graham, 1978). As depicted in figure 12.7, heart period can be
measured beat by beat or be computed for a particular time period. To
compute heart period in real time, it is important to use a weighted
averaging procedure so that the average reflects the proportion of time
that each beat contributes to the overall average. See figure 12.7 for an
example of computing an average weighted heart period or heart rate
for a given epoch.
Cardiac Output
Cardiac output (generally in liters per minute) represents one of the most
important cardiodynamic variables because it determines in large part
the amount of oxygen and nutrients available to the brain, the heart,
and to skeletal muscles that is necessary to sustain life and respond to
the internal and external stimuli with which we are constantly con-
fronted. In the past, most measurements of cardiac output were made
invasively, which limited their use to clinical settings. However, psycho-
physiologists now use indirect noninvasive measurements of cardiac out-
put. The two noninvasive procedures most commonly used in the labo-
ratory are impedance cardiography (ZCG) and combined-Doppler
ultrasound. The combined-Doppler ultrasound procedure, however is cur-
rently too expensive to be found in many psychophysiological labs; this
has made impedance cardiography the current method of choice. Impe-
dance cardiography is a technique that provides a measure of changes

Figure 12.7. Calculation of weighted average heart period and heart rate.
The upper part of the figure shows a series of four intervals representing four
interbeat intervals of 750, 730, 745, and 775 ms, respectively. The next part
of the figure depicts a clock running along with the interbeat intervals. In
this case, the clock began timing as the first interbeat interval began. Sam-
pling of a value by the computer is shown in the next line, with a sample of
the data taken at each 100 ms interval beginning with the start of the clock
and time zero. Calculations are shown for deriving both weighted heart period
(examples of a 1-s and 3-s weighted average) and for weighted heart rate
(example of a 1-s weighted average).
in blood flow in the thorax. From those changes one can derive measures
of stroke volume. Recall that cardiac output is the product of heart rate
and stroke volume. Thus, combining measures from the EKG and the
impedance cardiogram, one can derive a noninvasive estimate of cardiac
output. It appears that cardiac output derived from impedance cardi-
ography is likely to be a biased estimate of the true absolute cardiac
output, but the relative measures of changes in cardiac output are valid.
For a review of various methods for measuring cardiac output, including
noninvasive measures, see Ehlers, Mylrea, Waterson, and Calkins (1986).
For more information on impedance cardiography see Sherwood et al.
(1990), and Miller and Horvath (1978).
Blood Pressure
Since the seventeenth century, scientists have had some understanding

of the function of the circulatory system, but it was not until 1733 that
blood pressure was first measured. Before that time, it was believed that
blood circulation was maintained at each heart beat by a force of
100,000 pounds. Steven Hales, a scientist and Anglican priest, was the
first person to measure blood pressure in animals and dispel what he
considered to be "unsatisfactory conjectures." Hales inserted a flexible
tube, or cannula, into an artery of a horse which was, in turn, connected
to a long glass tube open at one end. With each beat of the horse's heart,
the blood would rise or fall within the tube, thus yielding a measure of
the animal's blood pressure (Cohen, 1976). An instrument is still in use
for animal research based on the same principle as Hales's device. A
refinement of this direct method still remains the most accurate measure
of human blood pressure, although there is only a limited number of
situations (e.g., surgery) where it can be applied.
Before considering methods of measuring blood pressure, let us review
the pumping of the heart as it relates to pressure. It was previously stated
that blood low in oxygen returns to the right side of the heart from the
body, enters the right atrium, then flows into the right ventricle and from
there is pumped into the lungs. Following this, the oxygenated blood
returns to the heart through the left atrium and then into the left ven-
tricle for distribution throughout the body. It is the contraction of the
left ventricle that produces the necessary pressure to move the blood
throughout the body. Sufficient pressure must be generated to push blood
through the arteries, capillaries, and veins. The larger arteries and veins
require smaller differences in pressure across the length of the vessel for
blood flow than do the small vessels where resistance to flow will be
relatively high.
The maximal, or systolic, blood pressure occurs when the ventricle of
the heart contracts. Following the period of cardiac contraction, there is
relaxation (diastole) of the ventricle, during which blood pressure is at a
minimum; a measurement at this time yields diastolic pressure. In gen-
eral, diastolic pressure varies mostly with peripheral resistance, whereas
systolic pressure is related to both peripheral resistance and stroke vol-
ume. Blood pressure is reported as systolic pressure over diastolic pres-
sure. The standard unit of measurement is millimeters of mercury, ab-
breviated as "mmHg." The typical blood pressure of a healthy college
student is 120/80 mmHg, although various factors, such as age, diet,
posture, and weight, are important.
Pulse pressure is the pressure difference between systolic and diastolic
pressures. In the case of the typical college student, the pulse pressure
would be about 40 mmHg (120 — 80 = 40). Some researchers calculate

the mean arterial pressure, which is generally defined as diastolic pressure
plus 1/3 pulse pressure.
Recording Procedure
In clinical practice, both direct and indirect measures of blood pressure
are used. The direct measure uses a catheter that is inserted into a vessel
or heart cavity. The direct method is infrequently used in psychophysi-
ological research and the interested reader should consult a medically
oriented text (e.g., Guyton & Hall, 1996), for more details. For additional
information about non-invasive recording of blood pressure consult Pap-
illo and Shapiro (1990).
Indirect Measurement The most common indirect measure of blood pres-

sure employs a sphygmomanometer. This device, commonly found in phy-
sicians' offices, consists of a pressure cuff connected to a tube containing
mercury. The present method dates back to the beginning of this century
and was introduced by Korotkoff. The following is a step-by-step proce-
dure for determining both systolic and diastolic blood pressure.
The blood pressure cuff is placed on the upper arm, with a stethoscope
over the brachial artery and just below the cuff (figure 12.8). When the
cuff is inflated to a pressure sufficient to cut off all arterial blood flow, no
sound is heard through the stethoscope. As the pressure in the cuff is
slowly reduced, the so-called Korotkoff sounds (tapping sounds) appear.
The cuff pressure at which the first sound is heard is read as systolic
pressure and is referred to as the start of phase I. As the pressure in the
cuff continues to decrease, the sounds take on a murmuring quality
(phase II) and then become clearer and louder (phase III). Following this,
Korotkoff sounds become muffled (phase IV). This lasts for the next 5 to
6 mmHg fall in pressure, until the sounds disappear completely (phase
V). In general medical practice and typically in the psychophysiology
laboratory, the pressure at phase V is considered the diastolic blood pres-
sure. Certain methodological and experimental problems exist in record-
ing blood pressure. First, blood pressure is not static, even though our
use of absolute numbers might lead us to believe otherwise. Rather, as
pointed out more than 200 years ago by Hales, there is a natural vari-
ation in the system. A normal individual's blood pressure can vary as
much as 30 mmHg during a 1-min recording using a direct blood pres-
sure technique (Tursky, 1974).
When using a sphygmomanometer to obtain indirect blood pressure,
a number of factors must be taken into account. First, there are factors
related to the technique itself. Repeated measurements within a short
period will produce (through the inflation and deflation of the pressure
cuff) temporary tissue changes that result in different blood pressure read-
ings. Second, there are factors related to the instrument being used. Most
Figure 12.8. Auscultatory method for measuring blood pressure. The aus-
cultatory method relies on the changes in sounds in a microphone located
over the artery, which has been briefly occluded. First, the occluding cuff is
pumped full of air to a relatively high pressure (above systolic pressure). Then,
air is gradually bled from the cuff. The blood pressure at which the first
arterial sounds appear is read as the systolic blood pressure. Diastolic pressure
is typically read as the pressure in the cuff when the last arterial sound
disappears. When using this method, mean arterial pressure is calculated
from the systolic and diastolic pressures (see text for the formula). Figure is
from L. A. Geddes, 1991, Handbook of blood pressure measurement, Clifton, NJ:
Humana Press.
notably, the size of the cuff can change the blood pressure recorded. To
determine the appropriate cuff size, the circumference of the upper arm
is first determined. The cuff width should be at least 40% of arm circum-
ference, and the cuff length should be at least 80% of the circumference
(Bailey & Bauer, 1993). In addition, the closed pressure system of the
instrument must be free of all leaks (e.g., of air or mercury) in order for
the measurements to be valid. Finally, blood pressure can vary due to
factors related to the person's situation. For example, several investigators
have found higher blood pressure readings when they are taken in a
clinical setting than when they are taken during a normal day (a phe-
nomenon known as "white-coat hypertension"). Similarly, a subject
coming for a psychophysiological experiment may initially experience
anxiety or fear concerning the situation or the equipment, and thus dem-
onstrate blood pressures that differ from normal readings before the ex-
periment has even begun. Another set of factors that also should be
considered is related to individual differences in the skills of the experi-

menter. That is, since the presence and absence of Korotkoff sounds re-
quire a judgment on the part of the experimenter, the precision with
which the recording is made and the skill of the experimenter must also
be considered.
Automated Indirect Measurement Automated devices for measuring blood

pressure are available and can be utilized with a polygraph or computer,
or the measurements can simply be read from a visual display on the
front panel of the instrument. These devices employ either a microphone
in the cuff (auscultatory method) or a pressure transducer in the cuff
(oscillometric method). The auscultatory method utilizes Korotkoff sounds
transduced by the microphone in the cuff in the same way as would be
done by a human observer using a manual recording system. Systolic
pressure is taken to be that point at which the first Korotkoff sound
appears on the polygraph record and diastolic pressure the level at which
the last sound appears. With this method, it is easier to accurately de-
termine systolic than diastolic pressure. The oscillometric method utilizes
oscillations in pressure in the cuff to determine systolic, diastolic, and
mean arterial pressure (Borow & Newberger, 1982). With this method,
following inflation of the cuff to a pressure above the systolic pressure,
the cuff is deflated in steps. The systolic pressure is taken as the pressure
when the oscillations in the cuff first begin to get larger, the mean arterial
pressure is taken as the point when the cuff oscillations are maximal in
size, and the diastolic pressure is taken as the point when the cuff pres-
sure oscillations no longer become smaller in amplitude (see figure 12.9).
A major problem with both the manual and automated procedures
for obtaining indirect measures of blood pressure is that a cuff must be
inflated and deflated. This is sometimes a stronger stimulus for the subject
than the independent variable and it also limits the number of readings
that can be taken in a given session. The development of continuous
blood pressure recording devices removes the need for repeated cuff in-
flations. These devices are in use in many psychophysiology labs, but the
most affordable devices were removed from the market. Currently, several
relatively expensive beat-to-beat recording devices are available, and it is
likely that less expensive devices will reappear on the market. The two
most commonly used devices for the psychophysiological lab use either
arterial tonometry or vascular unloading (also known as the method of
Penaz). Blood pressure is measured using the arterial tonometry method
by placing a sensor over an artery that overlies bone. Slight pressure is
applied to the artery, and the pressure in the partially flattened artery is
recorded as the arterial pressure. The most common site for tonometric
measurements is the radial artery at the wrist. Care must be taken that
the sensor is placed directly over the artery and that no movement of
the artery or sensor takes place during the recording session (Kemmotsu,
Ueda, Otsuka, Yamamura, Winter, & Eckerle, 1991). The vascular un-

Figure 12.9. Oscillometric method for measuring blood pressure. The oscil-
lometric method relies on the transduction of oscillations in the cuff pressure
to derive blood pressure. The occluding cuff is pumped full of air to a relatively
high pressure (above systolic pressure). Then, air is gradually bled from the
cuff. Oscillations in the pressure inside the cuff are used to mark systolic,
mean arterial, and diastolic pressures. Systolic pressure is determined when
pressure oscillations first appear as the pressure in the occluding cuff is bled
off. Mean arterial pressure is taken as the point where the pressure oscilla-
tions are maximal in size, and diastolic pressure is taken as the point when
the last pressure oscillations in the cuff disappear. Figure is from L. A. Geddes,
1991, Handbook of blood pressure measurement, Clifton, NJ: Humana Press.
loading method typically uses a cuff on the finger to clamp the vascular
volume of the finger at a specific level which is maintained from beat to
beat. The volume is measured using a plethysmographic device, which
reflects changes in blood volume beneath the sensor (see section on blood
volume); then, using a servo-control system, changes are made in the
pressure within the finger cuff, and the artery is returned to its previous
volume. The amount of pressure change needed in the finger cuff to
reestablish the volume in the artery is a function of the arterial pressure
underlying the cuff (Shapiro et al., 1996).

The measurement and quantification of blood pressure data are related
and often limited to the particular measuring device used. For example,
even when the cuff method is automated, it is not recommended that the
experimenter take more than one reading per minute (Shapiro et al.,
1996). Beat-to-beat recordings permit blood pressure determinations for
each beat, which will result in a greater reliability of measurements.
Because of the variability in blood pressure that was noted previously, it
is recommended that multiple blood pressure readings taken from a re-
cording epoch be averaged to provide a more stable estimate of the blood
pressure for that epoch.
Blood Volume and Flow
Many individuals have experienced vasomotor changes, such as cold feet

and hands, while waiting for someone special or preparing to take an
important examination; others have experienced a flushing or blushing
during emotional arousal. As was mentioned earlier in this chapter,
changes in blood volume may be produced by either a decrease or an
increase in sympathetic nervous system activity.
The two most commonly used vasomotor measures in psychophysio-
logical research are blood volume and pulse volume. Blood volume re-
cordings reflect relatively slow changes (i.e., changes in tonic level) in
the amount of blood in an arm, leg, finger, or toe. Pulse volume is a
phasic measure of the pulsatile change in blood flow related both to the
pumping action of the heart and to the vasodilation and constriction of
vessels in the periphery. Thus, pulse volume is a measure of the ampli-
tude of individual pulses.
Recording Procedure
The most common measure of blood volume used in psychophysiological
laboratories today is that of plethysmography. The term "plethys-
mography" comes from a Greek word that can be translated as "enlarge-
ment" or "fullness." Technically, a plethysmograph is a device for mea-
suring the change in volume of a given structure such as an arm, leg,
finger, or even the entire body. Thus, a true plethysmograph would con-
sist of a sealed chamber in which the arm, leg, or other structure is
placed. Since the chamber is sealed, any change in the volume within
the chamber would cause a recordable pressure change.
Although there are a variety of techniques for measuring blood vol-
ume and pulse volume, the three most commonly used are (1) photo-
electric changes, (2) impedance changes, and (3) volume changes re-
corded with a strain gauge.
Photoelectric Plethysmography By far the most popular plethysmographic

technique employs a photocell placed over an area of tissue perfused with
blood. There are two basic variations of this method: a light source can
pass through the tissue segment (transillumined) or bounce off the tissue
(backscattered). The amount of light that passes through or is reflected
back is received by a photoelectric transducer and converted into elec-
trical energy for recording. The light source produces light in the infrared
range (7000-9000 A). Since electromagnetic radiation in this frequency
is scattered by blood, the output of a photodetector may be considered
related to the amount of blood within the region (Jennings, Tahmoush,
& Redmond, 1980; Tursky & Jamner, 1982). Currently, most plethys-
mographic devices utilize either a light-emitting diode (LED) or a photo-
transistor that has minimal effects on the underlying skin and blood ves-
sels so that the device does not alter the tissue being recorded from.
If the light is transmitted through the tissue to a photodetector on the
other side, only a limited number of sites are convenient (e.g., the ear-
lobe). However, as demonstrated by Stern (1974), blood flow changes in
the earlobe are not particularly responsive to typical laboratory tasks and
stressors such as viewing slides of auto accidents or nudes, or performing
mental arithmetic. With the backscattered photoelectric plethysmo-
graphic technique, the light source and the photodetector are both lo-
cated on the same side of the tissue and thus can be placed almost any-
where on the body. The backscattered photoplethysmograph is more
sensitive to the vascular fluctuations occurring close to the surface of the
skin, whereas the transillumined photoplethysmograph is sensitive to
both skin and deeper tissue vascular changes. For an example of a recent
study employing photoplethysmography, see Spence, Shapiro, and Zaidel
(1996).
Impedance Plethysmography In chapter 10, impedance was mentioned as

a method of recording respiration. It can also be used to record blood
volume changes. One of the major advantages of the impedance method
is that it is not prone to artifacts caused by EKG, EMG, and other electrical
activity observed at the surface of the skin. The procedure employs two
electrodes on the skin through which a high-frequency alternating cur-
rent is passed. Since human tissue is a good conductor of alternating
current, and since changes in the amount of blood volume in a given
tissue segment produce changes in electrical impedance, it is possible to
infer indirectly the change of blood volume taking place. This technique
is the same as impedance cardiography in which the segment of interest
is the torso. For a recent study using impedance plethysmography in body
segments other than the torso, see White and Montgomery (1996).
Strain Gauge Plethysmography Previously we discussed the application of

strain gauges for measuring changes in respiration. Using the same prin-
ciple, strain gauges may also be used to measure changes in blood vol-
ume and pulse volume. The strain gauge is placed around the finger, toe,
or other body segment; changes in resistance or voltage of the strain
gauge can be considered an indirect measurement of blood volume

changes. The strain gauge is well suited for application on the penis
where other devices work less well. Indeed, use of the penile strain gauge
has become especially important in studies of male responses to erotic
stimuli and studies of treatments for sex offenders (see Farrall, 1992, for
a discussion of the use of the penile plethysmograph for assessing sexual
arousal).
Venous Occlusion Plethysmography Venous occlusion plethysmography is

a recording technique for measuring changes in blood flow to segments
of the limbs. This technique generally requires two cuffs, one placed distal
to the limb segment of interest, and one placed proximal to the limb
segment. The distal cuff is inflated to a pressure above the systolic arterial
pressure in order to prevent blood flow into and out of the distal limb
segment. The proximal cuff is inflated to a pressure level that is sufficient
to eliminate venous flow from the limb segment, but which does not
prevent arterial flow into the segment. A strain gauge is placed around
the limb segment being measured, and the change in limb circumference
per unit of time is used to infer the rate of blood flow into the segment.
The advantage of this method is that arterial blood flow into the isolated
limb segment can be measured independent of other possible sources of
blood flow change (e.g., venous flow). One obvious limitation of the ve-
nous occlusion technique is that measurements cannot be taken contin-
uously; even at very short intervals, numbness or pain in the limb can
result. In addition, measurements can be altered by movement of the
limb. For a recent example of the use of the venous occlusion pie thy s-
mographic technique, see Lindqvist, Melcher, and Hjemdahl (1997).
Typical Recordings and

Common Problems
Figure 12.10 shows two typical recordings of vasomotor activity: blood
volume and pulse volume. As noted previously, the blood volume mea-
sure represents the relatively slow changes of blood in a particular struc-
ture, such as an arm or a leg. Measurements of blood volume are made
using a DC amplifier connected to an appropriate preamplifier and to
whatever type of transducer is used. In figure 12.10, a photoelectric cell
sensed the blood volume changes. Likewise, pulse volume changes were
recorded with a photoelectric cell on the finger, except that the amplifier
used was AC coupled. Recall that AC coupling removes some of the
slower frequency shifts in the DC coupled signal. Pulse volume recordings
reflect the changes in blood flow to an area with each beat of the heart.
Jennings, Tahmoush, and Redmond (1980) and Tursky and Jamner
(1982) reviewed many of the factors that influence vasomotor changes
and discussed problems of interpretation from the indirect measures that
have been described here. For example, changes in room temperature
Figure 12.10. Blood volume and pulse volume recorded simultaneously. Note
that the blood volume was DC coupled whereas the pulse volume was AC
coupled. AC coupling removes the slow drift.
during a study from one day to the next could result in differential results
unrelated to the experimental situation, especially using the measure of
pulse volume. Another problem is related to the wide variation in skin
characteristics (e.g., location of vessels), which makes absolute compar-
isons between subjects impossible. Even within the same subject, difficulty
in precise placement of the transducer makes comparisons only relative.
It should also be pointed out that since different areas of the body contain
differential distributions of muscles and blood vessels, it is difficult to com-
pare results between studies when varying transducer sites are used. Fi-
nally, one must remember that blood flow is a complex function of pres-
sure in the vasculature, the radius of the blood vessels, and the viscosity
of the blood. Moreover, flow changes may occur in a given segment by
means of arterial or venous flow changes either into or out of the segment
of interest. Thus, one must be cautious in making interpretations about
the mechanisms underlying indirectly measured blood flow changes.

Because of the relative nature of the vasomotor measurements performed
by most psychophysiologists, experimenters typically examine changes
for each participant from a baseline period and compare this to the ex-
perimental or treatment period. The change between baseline and treat-
ment is generally expressed as a percentage. The magnitude of an indi-
vidual pulse is determined by simply measuring the difference between
the lowest point and the peak and a computer can be programmed to
detect the minimum and maximum points. Some researchers have used
an integrating coupler to perform a similar function. Because integration
of a curve represents the area under the curve, this may also represent
relative pulse volume changes.

As always in psychophysiology, new methodological advances will
likely enhance our ability to measure a greater number and variety of
hemodynamic variables, which will permit a more comprehensive picture
of the complexities of the cardiovascular system.
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13
Skin
Electrodermal Activity
Electrodermal recording continues stoutly to provide useful data in

spite of being abused by measurement techniques which range from
the arbitrary to the positively weird.
(Lykken and Venables, 1971, p. 656)
Electrodermal activity (EDA) has been recorded in thousands of psycho-

physiological studies. Why is this such a popular measure? Many who
record EDA today share the basic belief expressed by Carl Jung in 1907
and also by present-day lie detector operators that verbal responses do
not tell all, but that EDA does reveal the secrets of "mental life." Neu-
mann and Blanton (1970), in a fascinating history of early EDA research
which was referred to in chapter 1, noted the "mind-reading" view of
EDA that was made popular by Jung's word-association experiments. Pe-
terson (1907), a student of Jung, described the experiments as follows:
"It is like fishing in the sea of the unconscious, and the fish that likes
the bait best jumps to the hook. . . . Every stimulus accompanied by an
emotion produced a deviation of the galvanometer to a degree of direct
proportion to the liveliness and actuality of the emotion aroused"
(p. 805).
Today we think of EDA as a measure of the state of the organism's
interaction with its environment. EDA reflects not only emotional re-
sponding; it is also elicited by cognitive activity (e.g., Siddle, 1991). Edel-
berg (1972) reminded us that the skin has a special significance because
it both receives outside information and responds to signals from within.
He further states, "We can listen in on such signals by taking advantage
of the fact that their arrival at the skin is heralded by measurable elec-
trified changes that we call electrodermal activity" (p. 368). If we think
206
of our skin as a giant receptor separating us from the rest of the world,
is it any wonder that responses obtained from it would be of interest to
psychologists?
From an evolutionary standpoint, the survival value of EDA is a per-
plexing issue and one not easily studied. Why do the palms of our hands
and the bottoms of our feet sweat when we are anxious? Darrow sug-
gested (1933) that this response has persisted through evolution because
it aids grasping—for example, grasping of vines in flight, grasping of clubs
when fighting, grasping of tennis rackets, and so on. Edelberg (1973) has
written about the relationship of changes in the electrical activity of the
skin to survival functions such as locomotion, manipulation, and defense.
Terminology
The terms used to describe EDA have changed over the years. The term
used at the turn of the nineteenth century was "psychogalvanic reflex."
Later the term "galvanic skin response" was used. Today most psycho-
physiologists favor the term electrodermal activity.
The electrical activity of the skin can be measured in two ways. First,
a small current can be passed through the skin from an external source
and the resistance to the passage of current then measured. This tech-
nique was first used by Fere in 1888 and is referred to as the exosomatic
method. The second method, the endosomatic technique, was first used
by Tarchanoff in 1889; it measures the electrical activity at the surface
of the skin, with no externally imposed current. The exosomatic method
has been modified today into the measurement of skin conductance (SC),
the reciprocal of skin resistance. The endosomatic method is still used to
measure skin potential (SP), but skin conductance recording is used today
by most researchers.
When describing electrodermal activity, one can discuss basal activity
(tonic) versus the response to a stimulus (phasic) (see chapter 4). When
referring to tonic electrodermal activity, the convention is to use the word
level (L); when discussing phasic activity, the convention is to use the
word response (R). Therefore, the four common descriptions of electro-
dermal activity are as follows:
skin conductance level (SCL);

skin conductance response (SCR);
skin potential level (SPL);
skin potential response (SPR).
As stated in chapter 4, psychophysiological recordings show a third

type of activity in addition to tonic and phasic activity: spontaneous or
SKIN 207
nonspecific activity. Electrodermal activity is no exception. Spontaneous
electrodermal activity appears in records obtained using both the SC and
SP techniques.
Table 13.1 summarizes the various skin conductance measures used
by researchers and provides some expected values; this information is
from Dawson, Schell, and Filion (1990). The reader is referred to their
chapter for a more detailed discussion of electrodermal activity.
Figure 13.1 shows a simultaneous recording of SC and SP. The SPR
is usually biphasic, with the negative component followed by the positive
component. In some cases SPR is uniphasic negative. The latency of SCR
and the negative phase of SPR are usually between 1.0 and 3.0 s. Note
that SCL and SPL changed during the few seconds of this recording. These
relatively slow or gradual changes in SCL and SPL are not considered to
be examples of SCR or SPR. The latter terms are reserved for the more
Table 13.1 Electrodermal Measures, Definitions, and Typical Values

Typical
Measure Definition Values
Skin conductance Tonic level of electrical con- 2-20 microsiemens
level (SCL) ductivity of skin (uS)
Change in SCL Gradual changes in SCL mea- 1-3 |0,S
sured at two or more
points in time
Frequency of NS- Number of SCRs in absence of 1-3 per min
SCRs identifiable eliciting stimu-
lus
ER-SCR ampli- Phasic increase in conduc- 0.2-1.0 |iS
tude tance shortly following
stimulus onset
ER-SCR latency Temporal interval between 1-3 s
stimulus onset and SCR ini-
tiation
ER-SCR rise time Temporal interval between 1-3 s
SCR initiation and SCR
peak
ER-SCR half re- Temporal interval between 2-10 s
covery time SCR peak and point of 50%
recovery of SCR amplitude
ER-SCR habitua- Number of stimulus presenta- 2-8 stimulus
tion (trials to tions before two or three presentations
habituation) trials with no response
ER-SCR habitua- Rate of change of ER-SCR 0.01-0.5 (iS per
tion (slope) amplitude trial
Source: From M. Dawson, A. Schell, and D. Filion, 1990, "The electrodermal system." In J. T.
Cacioppo and L. G. Tassinary (Eds.), Principles ofpsychophysiology, Cambridge: Cambridge University
Press. Reprinted by permission.

Figure 13.1. A simultaneous recording of skin potential and skin conduc-
tance.
rapid responses to stimuli, which show the latency and shape of the
responses in figure 13.1.
Physiological Basis
We turn now to a brief summary of what is known about the underly-

ing physiological mechanisms of electrodermal activity. For additional
information, consult Dawson, Schell, and Filion (1990) or Boucsein
(1992).
We still do not know a great deal about the complex relationship of
the central nervous system to EDA. Boucsein (1992) has proposed a two-
component model of this relationship. According to Boucsein, there are
two separate portions of the central nervous system that are involved in
the control of EDA activity: an ipsilateral system—the hypothalamus,
anterior thalamus, and cingulate gyrus; and a centralateral system—the
lateral frontal cortex, particularly the premotor cortex, and parts of the
basal ganglia. Boucsein goes on to propose that the ipsilateral component
controls EDA when the stimulus is of an emotional or affective nature,
and the contralateral component controls EDA during orienting, cogni-
tion, and locomotion. For more details of this model the reader is referred
to Boucsein's comprehensive book.
Peripherally, we know that eccrine sweat glands (a special type of
sweat gland) are intimately involved in EDA. Eccrine sweat glands are
concentrated in the palms of the hands and soles of the feet. What makes
them of particular interest to psychologists is that they respond primarily
to "psychic" stimulation, whereas other sweat glands respond more to
increases in temperature. The eccrine sweat glands are innervated by the
SKIN 209
sympathetic branch of the ANS, but the chemical transmitter at the post-
ganglionic synapse is acetylcholine, not noradrenaline, as would be ex-
pected in the sympathetic nervous system (Shields, MacDowell, Fairchild,
& Campbell, 1987). This is worth noting, because some investigators
make the mistake of generalizing from EDA recording to all other psy-
chophsyiological activity. Generalizing from one channel of data is always
risky in psychophsyiology (see chapter 5), but particularly so when the
single measure used is an exception to the rule.
The eccrine sweat glands, which can be thought of as tiny tubes with
their openings at the surface of the skin, act as variable resistors wired
in parallel. Figure 13.2 shows a simplified model. Depending upon the
degree of sympathetic activation, sweat rises toward the surface of the
skin in varying amounts and in varying numbers of sweat glands. The
higher the sweat rises in a given gland, the lower the resistance in that
variable resistor. In some cases, but certainly not all, sweat overflows
onto the surface of the skin. This hydration of the skin with salty sweat
increases SCL and SPL. Remember, conductance is the reciprocal of re-
sistance. Years ago, it was thought that EDA was determined solely by
the amount of sweat on the surface of the skin. We now know this is
not so. Even in those cases where stimulation does not result in sweat
at the surface of the skin, changes in EDA are often found because even
a slight rise of the sweat in the glands will change the values of the
variable resistors shown in figure 13.2. If we wished to quantify the EDA
at a given moment, we would sum the values of all the active resistors
that are wired in parallel. The sum of resistors in parallel equals the sum
of their reciprocals, or conductance. This is one reason for using skin
conductance rather than skin resistance when describing exosomatic
EDA. A second reason, which will be discussed in the section on Analysis
and Quantification, is that skin conductance, unlike skin resistance, does
not have to be corrected for base level (see the discussion of the law of
initial values in chapter 5). In addition to the hydration of the skin and
Figure 13.2. A simplified model of sweat gland activity with the individual
glands depicted as variable resistors wired in parallel.

the number of active sweat glands, other factors that may be involved
in EDA include a membrane in the sweat duct wall that effects the reab-
sorption of sweat (Edelberg, 1972) and changes in pressure in the duct
that effect the opening of the pores in the skin (Edelberg, 1993). And for
skin potential, Christie and Venables (1971) demonstrated that SPL of
extremely relaxed subjects is determined largely by the concentration of
potassium at the surface of the skin.
Skin Conductance
Recording Procedure
Electrodes Silver-silver chloride electrodes should be used to minimize
polarization, which can affect the subject's conductance. The size of the
electrodes will affect conductance; the size used should be stated. Stan-
dard commercial electrodes are a good size and attach conveniently with
double-sided adhesive collars. The collars also help control the size of the
skin area that comes in contact with the electrode jelly. The contact area,
not the size of the electrode, effects the conductance values. Extremely
small electrodes should be avoided, because the smaller the contact area
the greater the current density (the amount of current flowing per unit
of electrode area). Too high a current density will effect the recordings
and could even stimulate underlying tissue.
The electrode jelly is the conductive medium between the electrodes
and the skin. Commercial EKG or EEC electrode jellies can be used for SC
recording, but if they contain near saturation concentrations of saline,
such jellies will effect recordings over time. Instructions for making elec-
trode jelly that will not effect recordings are given in Venables and Chris-
tie (1973, p. 80).
Electrode Placement The electrodes are attached where the concentration

of eccrine sweat glands is the highest—the palmar surface of the hands
or fingers or the soles of the feet. Either bipolar or monopolar placements
may be used. Figure 13.3(a) shows two typical bipolar placements. Figure
13.3(b) depicts a monopolar placement.
Recording Equipment Older equipment for recording EDA impressed a

direct current across two electrodes and indicated changes in skin resis-
tance. Newer circuits, instead of imposing a constant current, impose a
constant voltage, resulting in a direct readout of skin conductance. Either
system can be used; in fact, they are both used in the same way. In the
section on Common Problems, the conversion from resistance values
(ohms) to conductance (microsiemens or the older term micromhos) is
discussed. Considerable amplification is required to record what can be a
SKIN 211
Figure 13.3. Typical bipolar and monopolar electrode placements, (a) Bipolar;
(b) monopolar.
very weak signal. A reliable calibration signal (normally found in com-

mercial skin conductance couplers) is also needed in order to quantify
responses.
Procedure If bipolar electrodes are to be used, the chosen sites are simply
washed with soap and water and the electrodes are attached. With mono-
polar placements, after both sites have been cleaned, the skin over the
inactive site must be abraded. At the start of recording, the appropriate
degree of amplification must be selected. One way to tell if there is suf-
ficient gain is to have the subject take a deep breath. If no response is
seen, the amplification should be increased. If, on the other hand, the
record frequently goes off scale while the subject is relaxing, the gain is
too high. After the desired amplification is set, the calibration signal
should be recorded.
Factors to Control When Recording EDA The following subject variables

have been found to effect EDA: age, sex, race, and stage of menstrual
cycle. Environmental factors that effect EDA include temperature, hu-
midity, time of day, day of week, and season. For a discussion of the
specific effects of these variables, see Venables and Christie (1973) or
Boucsein (1992).

Figure 13.4. A simultaneous recording of SCL (a) and SCR (b) to a noise.
Typical Recordings
Figure 13.4(a) shows the skin conductance level recorded on a low-
amplification channel to a loud noise; figure 13.4(b) shows the same
signal recorded on a second higher-amplification channel to obtain the
skin conductance response. Figure 13.5 is a record of the spontaneous
skin conductance activity of a subject considered to be a labile responder,
that is, one who showed numerous responses for which there were no
known stimuli.
Common Problems
Simultaneous Recording of Level and Response If one is interested in si-
multaneously recording SC level and response, or SP level and response,
then there is a problem. Basically, the problem is how to set one amplifier
in order to amplify two signals, or two aspects of the same signal, when
the signals or aspects vary greatly in magnitude. The amount of ampli-
fication needed in order to see the relatively small phasic responses will
be much too great to observe the relatively large tonic level; the record
will be driven off the scale by shifts in the tonic level. Before discussing
methods for solving this problem, we should mention why one might
Figure 13.5. Spontaneous skin conductance activity.
SKIN 213
want to record level and response simultaneously. One possibility is that
the investigator might be interested in the two separate measures as a
function of some stimulus situation. The more usual reason for simul-
taneous recording is an interest in phasic activity (the size of the discrete
response), together with the realization that the size of the response is
somewhat dependent on the tonic level prior to stimulation, particularly
if skin resistance rather than conductance is recorded (see the discussion
of the law of initial values in chapter 5).
One method of solving this recording problem is to split the signal
coming from the subject. One signal is then fed into a high-amplification
channel for SCR, and the other into a relatively low-amplification DC
channel for SCL (see figure 13.4). But this method is expensive, because
it requires two channels of equipment. An alternative method provides
phasic and tonic information within one channel and is available on
some commercial equipment. A bucking voltage or offsetting circuit is
used. With the subject in the circuit, the experimenter turns a knob that
is calibrated in units appropriate for measurement of tonic level, until the
signal is zeroed. At this moment, the experimenter can read the size of
the internal signal needed to oppose the subject's tonic level, the con-
ductance level, or potential level. With amplification turned up, the ex-
perimenter can then observe the relatively small phasic responses.
Converting from Resistance Readings to Conductance As previously stated,

some older equipment provides output in terms of resistance (ohms) rather
than conductance (micromhos). Note that the older unit of measurement
for conductance "mhos" is "ohms" spelled backwards. The current unit of
measurement for conductance is microsiemen or |iS (see table 13.1). The
conversion itself is a simple matter, since resistance is the reciprocal of con-
ductance. However, a warning is in order. Beware when converting the
amplitude of a response from resistance to conductance. Do not subtract
the poststimulus resistance from the prestimulus resistance and then con-
vert the difference to conductance. Rather, convert the prestimulus resis-
tance to conductance and convert the poststimulus resistance to conduc-
tance and then subtract the two conductances.
Figure 13.6 shows a typical response recorded on equipment that pro-
vides the output in resistance (ohms). The amplitude of the response in
conductance units (|J.S) was obtained as follows:
Poststimulus level = 90,000 ohms = 11 |0,S
Prestimulus level = 100,000 ohms = 10 uS
Amplitude of response = 1 jiS

Amplitude Figure 13.7 shows typical skin conductance waveforms. Mea-
surement of the amplitude of figure 13.7(a) is simple: conductance at the
Figure 13.6. A typical response recorded on equipment that provides the
output in resistance (ohms). The size of the response when converted to con-
ductance is shown to be 1 ^iS.
peak minus conductance prior to the response. Difficulties arise, however,

when a second response occurs before the first recovers, as in figure
13.7(b) or 13.7(d). One way of measuring the amplitude of the second
wave in figure 13.7 (b) is shown in figure 13.7(c). Most psychophysiol-
ogists do not use this approach, however, preferring instead to measure
the amplitude of each successive wave from its point of origin, as shown
in figure 13.7(d).
When conductance change is used, it usually is not necessary to cor-
rect for base level conductance prior to the response. Table 13.2 shows
why this is so. The skin resistance and skin conductance response of two
hypothetical subjects are compared. Subject A's prestimulus level of re-
sistance is very high compared to that of subject B. Therefore, it might
be said that A's electrodermal system has much further to decrease than
B's or that the size of B's responses should be inflated when compared
with A's because of the differences in prestimulus level. Simplistically, it
is easier for A to respond—show a decrease in resistance—than it is for
B. In the example given, both subjects decreased 10,000 H. This was
actually a much greater response for B, considering B's low prestimulus
level; so, the size of the response must be "corrected." The lower half of
table 13.2 shows the same data converted to conductance. Note that
here the size of B's response is much greater than A's, as it should be.
SKIN 215
Figure 13.7. Criteria for measurement of skin conductance amplitude (H).
Method d is preferred over method c for measurement of compound responses.
Redrawn with permission from R. Edelberg, 1967, "Electrical properties of
the skin." In C. C. Brown (Ed.), Methods in psychophysiology, Baltimore: Wil-
liams and Wilkins.
There is no need to correct the conductance data. Conductance data are

usually relatively normally distributed, eliminating the need for a trans-
formation.
One possible transformation that should be considered, however, has
been described in detail by Lykken, Rose, Luther, and Maley (1966) and
can be applied to all psychophysiological data. This technique enables us
to express the amplitude of a response relative to a particular subject's
minimum and maximum conditions of activation.
Latency The latency of EDA refers to the time from stimulus to onset of
the response. The latency of EDA is usually 1.0 to 3.0 s. It is difficult to
measure accurately because SCR does not begin abruptly but instead
starts gradually. This makes onset difficult to determine.
Recovery Time The onset phase of an SCR—the time from the start of
the response until it peaks—depends largely on the amplitude of the re-
sponse and is, therefore, of little interest to most investigators. Edelberg
(1972) and others suggest that the rate of recovery of an SCR is some-
what independent of amplitude, mediated by the central nervous system
and of interest to psychologists. The specific measure of recovery time
216 P S Y C H O P H Y S I O L O G Y OF SPECIFIC O R G A N S AND SYSTEMS

Table 13.2. A Comparision of Skin Resistance and Skin
Conductance Responses in Relationship to Prestimulus
Levels
Resistance (ohms)
Subject A Subject B
Prestimulus 100,000 20,000
Poststimulus 90,000 10,000
Amplitude of response 10,000 10,000
Conductance (|iS)
Prestimulus 10 50
Poststimulus 11 100
Amplitude of response 1 50
used, rec t/2, is the time from the peak to the point at which the response
has recovered to 50% amplitude. Numerous authors (e.g., Christie, 1976)
relate the SCR recovery rate to the nature of the testing situation (e.g.,
threatening versus task-orienting) and to the nature of the subject (e.g,
schizophrenics versus normals).
Frequency Counting nonspecific SCRs is not as easy as it seems. First,

one must set the criteria for a response; 0.01 fiS is commonly used. Then
one must rule out SCRs that occur within so many seconds of a known
stimulus, such as a deep breath. We suggest discounting all ANS re-
sponses for approximately 20 s after such a disturbance (see Stern and
Anschel, 1968).
Skin Potential
Recording Procedure
Endosomatic recording of EDA is performed with no excitation current.
What is recorded is the skin potential over a site rich in eccrine sweat
glands, with reference to an inactive site. The recording equipment must
contain a stable, high-gain amplifier with a high-input impedance. Stan-
dard silver-silver chloride electrodes are used, as with SC recording. In
SP recording, it is crucial to minimize electrode bias. The degree of bias,
or difference in potential, between a pair of electrodes can be measured
by immersing them in a saline solution and measuring the potential with
the same high-input impedance recorder that will be used to measure
SKIN 217
Figure 13.8. Three different waveforms commonly found in skin potential
recording. Negativity is recorded upward by convention and amplitude is
measured from the same prestimulus baseline.
skin potential. The bias potential should be less than 1 mV. The size of
the electrodes is not critical. Monopolar placement must be used (figure
13.3), and the inactive site must be abraded. The basic recording pro-
cedure is similar to that used for the recording of SC.
Typical Recordings Figure 13.8 shows three different waveforms com-

monly seen in SP recording. Note that negativity is recorded upward by
convention; the top figure depicts a simple negative wave. The middle
tracing shows a typical biphasic SP response. The lower figure shows a
more unusual triphasic waveform.
Common Problems
Electrode drift is the most common problem in recording skin potential.
Since SP recording must be carried out with relatively high amplification,
any slight potential between the electrodes will be recorded as a slow
wave, indistinguishable from a slow change in skin potential level. As
previously mentioned, electrode bias should be measured prior to record-
ing. In addition, electrode bias should be checked after a recording session
in order to be confident that the source of the record was the skin po-
tential of the subject rather than electrode potential.
Amplitude It is difficult to quantify the amplitude of SPRs. The conven-
tional method of quantifying a biphasic response is to measure the neg-
ative response from the preresponse level to its peak and then measure
the positive response from the same preresponse level to its peak (see
figure 13.7). The difficulty is that the amplitude of the negative wave is
attenuated, by some unknown amount, by the onset of the positive wave.
Latency Latency of the first wave of the SPR can be measured much
like latency in SCR (as previously discussed). The values obtained should
be similar.
References
Boucsein, W. (1992). Electrodermal activity. New York: Plenum.
Christie, M. J. (1976). Electrodermal activity. In 0. W. Hill (Ed.), Modern trends
in psychosomatic medicine. Vol. 3. London: Butterworth.
Christie, M. J., & Venables, P. H. (1971). Characteristics of palmar skin po-
tential and conductance in relaxed human subjects. Psychophysiology, 8,
523-532.
Darrow, C. W. (1933). The functional significance of the galvanic skin reflex
and perspiration on the backs and palms of the hands. Psychological Bul-
letin, 30, 712.
Dawson, M. E., Schell, A. M. & Filion, L. (1990). The electrodermal system.
In J. T. Cacioppo and L. G. Tassinary (Eds.), Principles of psychophysiology
(pp. 295-324). Cambridge: Cambridge University Press.
Edelberg, R. (1967). Electrical properties of the skin. In C. C. Brown (Ed.),
Methods in psychophysiology (pp. 1-53). Baltimore: Williams & Wilkins.
Edelberg, R. (1972). The electrodermal system. In N. S. Greenfield & R. A.
Sternbach (Eds.), Handbook of psychophysiology (pp. 367-418). New York:
Holt, Rinehart & Winston.
Edelberg, R. (1973). Mechanisms of electrodermal adaptations for locomotion,
manipulation, or defense. In E. Stellar and J. M. Sprague (Eds.), Progress
in physiological psychology Vol. 5 (pp. 155-209). New York: Academic
Press.
Edelberg, R. (1993). Electrodermal mechanisms: A critique of the two-effector
hypothesis and a proposed replacement. In J. C. Roy, W. Boucsein, D. C.
Fowles, & J. H. Gruzelier (Eds.), Progress in electrodermal research (pp. 7-
30). New York: Plenum.
Jung, C. G. (1907). On psychophysical relations of the associative experiment.
Journal of Abnormal Psychology, 7, 247-255.
Lykken, D. T., Rose, R., Luther, B., & Maley, M. (1966). Correcting pycho-
physiological measures for individual differences in range. Psychological
Bulletin, 66, 481-484.
Lykken, D. T., & Venables, P. H. (1971). Direct measurement of skin conduc-
tance: A proposal for standardization. Psychophysiology, 8, 656-672.
SKIN 219
Neumann, E., & Blanton, R. (1970). The early history of electrodermal re-
search. Psychophysiology, 6, 453-475.
Peterson, F. (1907). The galvanometer as a measure of emotions. British
Medical Journal, 2, 804-806.
Shields, S. A., MacDowell, K. A., Fairchild, S. B., & Campbell, M. L. (1987). Is
mediation of sweating cholernergic, adrenergic, or both? A comment on
the literature. Psychophysiology, 24, 312-319.
Siddle, D. A. T. (1991). Orienting, habituation, and resource allocation: An
associative analysis. Psychophysiology, 28, 245-259.
Stern, R. M., & Anschel, C. (1968). Deep inspirations as stimuli for responses
of the autonomic nervous system. Psychophysiology, 5, 132-141.
Venables, P. H., & Christie, M. H. (1973). Mechanisms, instrumentation, re-
cording techniques, and quantification of responses. In W. F. Prokasy
and D. C. Raskin (Eds.), Electrodermal activity in psychological research
(pp. 2-124). New York: Academic Press.

14
Signal Processing
You have designed a cognitive intervention intended to increase atten-

tional abilities in young children. You would like to use a psychophysi-
ological method to determine if there are any effects of your intervention
on attentional control. Before you begin a study, you will need to choose
the psychophysiological variable(s) that will permit you to answer your
question. You base this decision on the conceptualization of your re-
search question and on what is known about the impact of the relevant
psychological variable (e.g., attention) on the measured physiological var-
iable^). In this case, because your question concerns possible changes in
attentional ability, you will want to choose a dependent variable that
reflects changes in attentional processes.
The next important issue is to decide, for each measured physiological
variable, whether you are interested in recording basal (tonic) activity or
event-related (phasic) activity (see chapter 4). What is considered to be
the basal or tonic level of activity in a physiological system is typically
determined by your question and by the physiological system of interest.
In general, you can derive tonic activity from the ongoing level of activity
in the system when the organism is at rest. The length of time needed
to assess tonic level appropriately will differ across physiological systems,
but in many cases it will be on the order of several minutes. Time periods
shorter than this can be unstable measures of tonic activity because of
the occurrence of phasic events that can drastically alter the mean value
of physiological function during the period. On the other hand, time pe-
riods longer than this are often difficult to obtain because subjects do not
like to sit still for more than a few minutes, and very long time periods
(i.e., hours) may be altered by circadian effects. Phasic responses typically
are measured over time periods that capture the full response from basal
level until the physiological parameter returns to it's basal level. The
speed with which a physiological system responds and activity returns
221
Figure 14.1. Sample analog skin conductance signal. Both basal skin con-
ductance levels and two skin conductance responses are depicted (units are
microSiemens).
to baseline differs from system to system. As we mentioned in chapter

13, eccrine sweat gland activity measured as skin conductance is an
example of a system from which one can record both tonic and phasic
activity. Basal skin conductance, also referred to as skin conductance
level, changes relatively slowly (i.e., several seconds to minutes; see figure
14.1). In addition, skin conductance can also show short-term changes
known as skin conductance responses, which are superimposed on this
background skin conductance level. These responses typically last several
seconds and are distinct from the background level of activity (figure
14.1). Skin conductance responses may be nonspecific, (meaning that we
do not know of any event leading to their occurrence) or responses can
be event-related (meaning that the skin conductance response appears to
have occurred as a result of an experimenter-controlled external event
such as a stimulus display, or an unplanned or subject-controlled event
such as a sneeze).
Your decision about whether to record physiological level or response
(or both) will influence your recording parameters. First, you'll need to
determine the length of the time period that you wish to record. Note
that if you want to measure tonic activity then the recording period is
likely to be long relative to what is required to record a response. For
example, skin conductance or heart rate level is often determined over
one to several minutes, whereas a phasic skin conductance or heart rate
response is determined over one to several seconds. The time period
length should be based on what is known about the tonic levels and
phasic responsiveness of the physiological system of interest. Second, you
will need to decide on a sampling rate at which you want to record the
signal. Sampling rate, measured as samples per second, refers to the rate
at which your original (analog) physiological signal is converted to a
digital form, or set of numbers, by a computer (figure 14.2). The analog
signal is a waveform that is continuous across time. When the computer

Figure 14.2. Sample analog skin conductance signal and some digital values
illustrating sampling. A sample (a conversion from the analog signal to a
digital value) is taken at each vertical arrow over time. Every fourth digital
value is shown below the analog signal. Digital values are shown in
microSiemen units.
samples the signal it assigns a numerical (digital) value based on the

current value of the analog signal. In order to reproduce the original
signal accurately in digital form, it is important to sample frequently
enough so that the signal is reproduced faithfully. In figure 14.3, the
lower panel depicts a digital reconstruction of the analog signal depicted
in the upper panel. Notice that if you sample too slowly, the recreated,
digitized signal will not look like the original, analog signal. Also note
that although there are lines drawn between the dots, that the only
Figure 14.3. Sample analog skin conductance signal (upper panel) and the
recreated digital signal (lower panel). Each digital sample is represented by a
solid circle in the lower panel and the samples are connected by a line. Note
that when the signal level changes rapidly, the sampling rate shown is less
able to recreate faithfully the original analog signal than when the signal
changes slowly.
SIGNAL PROCESSING 223

information one has in a digitized signal are the data points represented
by the dots (which are numbers). Indeed, a digitized signal is also known
as a discrete-time signal because we only know the values of the signal
at discrete time points, and not at the points in between. Thus, the illus-
tration of a signal using "connect-the-dots" is a depiction only; we cannot
know for sure what the values of the analog signal are between the
sampled points. In order to reproduce all of the elements of interest from
the analog signal in the digitized waveform, one needs to determine the
highest frequency component of interest in the signal. As we mentioned
in chapter 3, it is necessary to sample at least twice the highest frequency
of interest in order to reproduce the signal; in general, it is a good idea
to sample even faster than that (e.g., four to eight times the highest
frequency of interest; Cacioppo, Tassinary & Fridlund, 1990). This rule
requiring a sampling rate of at least twice the highest frequency com-
ponent in the signal is referred to as the Nyquist relation. When sampling
is too slow, a digital form of the physiological signal will be produced
that does not accurately represent the original signal which is a problem
known as aliasing (figure 14.4). It may seem surprising, but recording
period length and sampling rate are related issues because the memory
capacity limitations of a computer used for collecting physiological data
results in a trade-off between these two experimental parameters. The
longer one's recording time, the more memory is required in order to
store the physiological data. Likewise, the faster the sampling rate, the
more numbers must be stored per unit of time and the greater the mem-
ory requirements for the computer storing the data.
One final decision that should be made prior to data collection is
whether you will use a filter when collecting the data. Filters remove,
or filter out, unwanted frequencies of activity from your physiological
signals. One of the most important sources of unwanted variation that
can be removed using a filter is 60 Hz noise. Such noise arises from
electrical devices, computer screens, and a variety of other sources.
Fluorescent lighting, a 120 Hz noise source, is particularly problematic
in the psychophysiology lab. In general, it is better to shield the source
of the noise from the subject or to shield the subject from the noise.
However, in some circumstances, the environment may not be easily
alterable. In this case, unwanted noise can be removed by filtering it
out of the recorded signal. In addition to removing unwanted noise,
filters can remove other unnecessary frequencies from the data. For
example, when recording an EEC, one may want to examine a particular
frequency of activity such as alpha activity in the EEC, but alpha activity
may not be apparent to the unaided eye. However, a filter could be
applied to remove frequencies above and below the alpha band, permitting
a relatively clear view of predominantly alpha activity. Analog filters
can be applied either online (during data collection), or digital filters
can be applied using a computer either online or offline (after the data

Figure 14.4. An illustration of aliasing: (a) depicts an exemplar analog phys-
iological signal; (b) depicts a digital recreation of the original signal where a
sample was recorded at each vertical arrow. Note that the sampling rate was
too slow to recreate faithfully the original signal, a problem known as alias-
ing.
collection). Filters can be achieved either using electronic circuitry (i.e.,

a hardware filter) or using mathematical procedures such as FFT (dis-
cussed later in this chapter) which are based in software. A hardware
filter is typically applied when the data are collected, however, a software
filter can be applied to the data as they are collected or afterward. If
one is unsure of the effect of a chosen filter on the data, it is most
conservative to record the data unfiltered, and then to apply the filter
after data collection and check what effect the filter has on the data.
Investigators should also be cautious that filters do not remove desired
elements of a physiological signal; they should also be aware that filters
are not perfect: a 60 Hz filter removes some frequencies of activity around
60 Hz along with the 60 Hz activity.

In the days before computers were a ubiquitous part of the psycho-
physiology laboratory, physiological signals were typically recorded with
a polygraph and scored from a paper record by hand. Today, computers
and relatively inexpensive recording equipment, and computer programs
for collecting and reducing physiological signals make it possible for most
labs to collect data directly into a computer and reduce the data offline
after the experiment is over. After collecting your physiological data, the
next step is to reduce it. Reducing physiological data refers to the process
by which the large sets of numbers recorded by the computer are reduced
to a smaller set of numbers that represent the events of interest during
the experiment. For instance, although you may have sampled a plethys-
mographic signal at 250 samples/s, you may ultimately decide that the
numbers you want to use in a statistical analysis are the mean values of
the plethysmographic signal for each 1-min period in the data set. Thus,
you will reduce the large data set of numbers (i.e., the digitized signal
representing the plethysmographic signal at 250 samples/s) to a set of
numbers representing the mean value of the signal once per minute.
Indeed, signal averaging is a very common method for reducing noise in
data (for an example, see chapter 7 on event-related potentials). These
averaged values then are submitted to statistical analyses and/or are
plotted in graphical form in order to see changes as a result of your
experimental manipulation or to see differences across different individ-
uals. Next we will consider some of the ways in which you can assess
your reduced physiological data.
Assessing Basal Activity
Assessing basal or tonic activity is typically accomplished by computing

mean values over a predefined period within the experiment. Mean values
typically provide a reasonably representative view of the overall level of
activity of the signal of interest. For some physiological signals, or under
some circumstances, one may choose another measure of central ten-
dency such as the median. The median is less sensitive than the mean
to outliers in a data set. Thus, for data sets containing a sufficient number
of outliers that the mean and median differ considerably, one may get a
better estimate of the general level of activity using the median rather
than the mean. In addition, one should be sensitive to whether an outlier
(or group of outliers) could be a "real" physiological phenomenon (e.g.,
a severe bradycardia in an infant) or a result of movement-related or
other artifact-related perturbation of the digitizing process. One may wish
even to remove a "real" event, for instance a precipitous bradycardia,
for purposes of creating a meaningful representation of the average level
of activity without the influence of a single punctate event. One final
issue concerns the choice of measure both for representing the status of
the physiological system and for performing statistical analyses. Each
physiological system will require a different solution for choosing a unit
of measurement and deciding whether there is a need to transform the
measurements for analysis purposes. When possible, it is suggested that
units of measurement be chosen on the basis of the characteristics of the
physiological system of interest (Levey, 1980). For an example, see the
rationale given in chapter 12 for choosing heart period instead of heart
rate as the cardiac metric. For more specific recommendations on the
choice of units of measurement for physiological variables, the reader is
urged to consult the relevant chapters in this volume. The reader should
also consult the literature for relevant information on the following gen-
eral measurement issues (Levey, 1980) and discussions on measurement
of skin conductance (Boucsein, 1993; Fowles et al., 1981; Lykken &
Venables, 1971), cardiac function (Berntson, Cacioppo & Quigley, 1995),
electromyography (Cacioppo, Tassinary & Fridlund, 1990), ocular re-
sponses (Stern & Dunham, 1990), respiratory responses (Lorig &
Schwartz, 1990), and sexual responses (Geer & Head, 1990).
Assessing Change
Decisions about how to assess changes in physiology are somewhat more

difficult than those for assessing tonic levels of activity. As we have men-
tioned previously, this is partly due to the fact that, in some cases, the
amplitude of a response may be in part dependent on the baseline level
of activity. We will discuss several of the methods used for assessing
change and indicate some of the advantages and disadvantages associ-
ated with each measure.
Change Scores
Change scores are computed by taking a measure during a phasic re-
sponse (either the mean or the peak) and subtracting a measure of base-
line activity. Thus, change scores reflect the change from the baseline to
the response. The use of change scores has been debated in the psycho-
logical literature, with no firm consensus emerging for different physio-
logical measures. This is because several issues influence whether or not
a change score is a reliable measure of change in a physiological system
over time. One issue concerns whether the baseline and task levels of
activity are independent of one another. This issue is related to the law
of initial values (LIV) (see chapter 5). The LIV states that the initial state
of a system will limit the degree to which the system can change it's
state. More specifically, the LIV states that the higher the basal level of
function in a given physiological system, the more the system can de-
crease its function or the less the system can increase its function. The

reverse is true when the basal level is low. There are situations in which
physiological limits do impose a constraint on the amount of change (see
Berntson, Uchino & Cacioppo, 1994). For example, there are physiolog-
ical and metabolic limits on the rate with which the heart can beat and
under normal physiological conditions there is a maximal rate. However,
it is also clear that the LIV does not always occur. That is why we prefer
to call it a "principle"; see also Myrtek and Foerster (1986). It is impor-
tant to keep in mind that when physiological functions are near a "ceil-
ing" or "floor" (i.e., near their physical limits), that responses can be
constrained by the system. At least with regard to cardiovascular mea-
sures, within many typical laboratory reactivity paradigms the correla-
tion between the baseline and the change score appears to be small,
suggesting that the LIV is not consistently problematic (Llabre et al.,
1991; Seraganian et al., 1985). However, the most conservative ap-
proach is probably to assess whether the baseline and task levels are
correlated, and if so, decide if it is important to remove the effect of the
baseline using one of the techniques discussed here.
The second issue concerns the issue of test-retest reliability of change
scores. Test-retest reliability refers to the idea that a change score mea-
sured in a person in response to a task on one occasion should be similar
to the change score measured in that same person in response to the
same task on another occasion. If a measure is unreliable, it means that
we cannot be sure that one would see a similar change score within a
person if the study was completed again. In cardiovascular reactivity
studies, where one typically measures changes in cardiovascular function
as a result of a task, challenge or stressor, the reliability of change scores
has been assessed. Cardiovascular measures such as heart rate and sys-
tolic blood pressure have been shown to have relatively good reliability,
whereas diastolic blood pressure has poorer reliability (Llabre et al., 1991;
Seraganian et al., 1985). For a discussion of reliability for several psy-
chophysiological measures, see Tomarken (1995).
Residualized Change Scores and

Covariance Analyses
Residualized change scores are scores derived by computing a regression
equation that reveals the relationship between the phasic or task level and
the baseline level. Task level is regressed onto baseline level; then the dif-
ference between the expected value depicted by the regression line and the
observed value becomes the new residualized change score (see figure
14.5). These change scores are sometimes used when there is a relation-
ship between the task and baseline levels and the investigator wishes to
remove the effect of the baseline. Note that unlike change scores (i.e., task
- baseline), the residualized change score is dependent on the sample from
which it is drawn. All data in the sample will be used to construct the re-
Figure 14.5. Derivation of a residual change score. As an example, task-level
heart rates were regressed onto baseline-level heart rates. The residual is the
difference (in bpm) between the observed value (solid circles) and the expected
value for the baseline level (dark square). The expected value falls on the
regression line that best fits the task and baseline level data. The residual
represents the heart rate change that occurs independent of the baseline.
gression equation from which the residuals are derived; therefore, differ-
ences across samples will result in somewhat different regression equa-
tions, and thus residual scores will vary from sample to sample. In this
regard, change scores are easier to interpret than residualized change
scores, because they are comparable across studies. A technique that is
conceptually similar to the use of residualized changes scores is analysis of
covariance (ANCOVA). Using ANCOVA, one can use the baseline level as
a covariate of the task level and thus statistically "remove" the effects of
the baseline from the task level. Comparisons of change scores and resi-
dualized change scores in cardiovascular reactivity research suggest that
they are usually similar in reliability; thus, the simple change score is
sometimes recommended because of its ease of interpretation and calcu-
lation (Kamarck et al., 1992; Llabre et al., 1991; Manuck et al., 1989).
Again, however, the best strategy is to assess the relationship between the
baseline and task levels so that one can make an educated decision about
whether or not to remove the effects of the baseline.
Percent Change Scores

Another type of change score that is used is the percent change score. It
is calculated by dividing the difference between the task and the baseline

by the baseline (i.e., [task level - baseline level]/baseline level). Thus,
the change in physiological activity is now represented as a function of
the original baseline level. There are some caveats that should be con-
sidered when using percent change scores. First, percent change scores
typically are not normally distributed, making them difficult to use in
parametric statistical tests unless they are first transformed. Furthermore,
the necessity of a transformation for statistical purposes after already
making the percentage transformation means that the data have now
been transformed twice and thus have become difficult to understand in
the sense of "real world" interpretability. Finally, little work has been
done assessing the reliability of percent change scores. Because of the
difficulty of interpreting transformed percent change scores and a lack of
information on their reliability, percent change scores are not usually
recommended unless the investigator can provide evidence of reliability.
Assessing Global Aspects of

Physiological Signals
Recent advances in signal processing capabilities with computers allow

the psychophysiologist to assess easily more global aspects of a physio-
logical signal. These techniques can provide an overall look at the vari-
ation in the signal over time, the frequency of that variation, and whether
the variation forms an overall pattern that is not visible to the naked
eye. Analytical tools that permit examination of a signal over time are
called time domain techniques. Tools that permit the investigator to ex-
amine the frequency characteristics of a signal are called frequency domain
techniques. Time domain techniques tend to be useful for showing what
a signal looks like over time. However, low-frequency patterns in a signal,
particularly in a signal recorded over a very long time period, can be
difficult to envision using time domain techniques. Instead, frequency
domain techniques often provide a more useful way of characterizing a
physiological signal because they serve to unearth patterns in the data
that are not visible to the eye. The interested reader is urged to consult
sources that can give a more detailed explanation of the various tech-
niques detailed here; useful sources of additional information are sug-
gested in each section. For an accessible overview of digital signal proc-
essing, the reader is urged to consult Lyons (1997).
Fourier Analysis or Fast

Fourier Transforms
Biological signals are typically composed of complex-looking waveforms.
Sometimes, we may want to decompose those waveforms into their com-

ponent frequencies because we believe that there is biological and/or
psychological "meaning" associated with a particular frequency of activ-
ity. For instance, there is an oscillation in heart period that is coincident
with the respiratory frequency (approximately 0.12-0.4 Hz in adult hu-
mans). This oscillation in heart period (or heart rate) is due to central
nervous system respiratory oscillators and feedback from the lungs, and
under some circumstances this oscillation will have a visible impact on
the heart period matching the frequency of respiration (see figure 12.5).
The oscillation appears to be associated with central parasympathetic
outflow to the heart; thus it is a frequency of particular interest for some
psychophysiologists (see chapter 12 for more information).
Fast Fourier Transform (FFT) is a frequency domain procedure by
which physiological signals comprised of waveforms with different fre-
quencies and different amplitudes are decomposed so that one can ob-
serve the underlying frequencies that make up the signal. The FFT as-
sumes that the underlying waveforms are sine waves; although this is
not necessarily a correct assumption for biological signals, it can still
serve as a reasonable approximation to reality. The FFT process is anal-
ogous to taking sine waves of many different frequencies and seeing how
each of the sine waves fits the signal of interest. Mathematically, the FFT
steps through a series of sine waves of successively greater frequencies,
checking a template of each frequency against the signal and quantifying
how much of the signal matches each of the frequency templates. The
FFT results in a graphical depiction called the power spectrum or spectral
density plot that illustrates how a given signal sampled over time is de-
composed into it's component sine waves which differ in frequency and
amplitude. The spectral density plot is useful because it summarizes how
much activity in the original signal arose from each frequency of sine
wave that was fitted to the signal. Figures 14.6 and 14.7 depict the
"template-matching" that is performed by an FFT. Notice that each of
the templates matches some part of the original signal, but that some
frequencies contribute more to the original signal than others. The spec-
tral density plot that results shows which frequency templates were most
prominent in the signal (i.e., greater power), and those which were a
small component of the signal (i.e., less power). Figure 14.8 shows time
domain and frequency domain depictions of two single frequency sine
waves and a signal made up of the two individual sine waves. Figure
14.8 (a) and 14.8 (b) each show a sine wave of one frequency and
amplitude. The left panels of the figure show the sine waves in the time
domain and the right panels show the decomposed sine wave in the
frequency domain. Figure 14.8 (c) depicts the combination of the sine
waves shown in figure 14.8 (a) and 14.8 (b) and illustrates how the
decomposition of the signal in the frequency domain reveals both fre-
quency components (one at f() and the other at 2 f () ). Figure 14.9 illus-
trates two different frequency domain depictions, one showing amplitude

and the other showing power (or spectral density) of the data shown in
the right side of figure 14.8 (c). Figure 14.9 shows that the signal is
made up of two different frequencies and that the amplitudes and spectral
power of the two frequency components differ. Indeed, power is simply
the (amplitude)2.
Biological signals are not made up simply of sine waves. FFT can still
be used, however, as long as the investigator is aware that the interpre-
tation of the power spectra for biological signals is less straightforward
than the interpretation of power spectra produced for pure sine waves.
Some real physiological signals (heart period and respiration) are depicted
in figure 14.10 and demonstrate that the resulting power spectra will be
more complex (e.g., has more than one peak) for more complex physio-
logical signals. Thus, respiration, which is relatively sinusoidal for a phys-
iological signal, gives a relatively clean, single-peaked power spectrum.
However, heart period, which is composed of multiple frequency com-
ponents, gives a more complex, multipeaked power spectrum. There are
other computational issues that must be addressed when using FFT, such
as the need to window the signals to remove the abrupt frequency shifts
that can occur at the beginning and end of a recording period, which
the investigator should understand when performing FFT analyses. For
a discussion of the issues related to using time domain and frequency
domain analyses such as FFT with psychophysiological signals, see Por-
Figure 14.6. Four templates of different frequencies that were used to con-
struct the signal shown in the top panel of Fig. 14.7.

Figure 14.7. The top panel illustrates a curve constructed from the templates
shown in figure 14.6. The bottom panel illustrates a spectral analysis (FFT)
of the signal in the upper panel with four characteristic frequencies emerging
as peaks.
ges & Bohrer (1990) for a very thorough treatment of the methodological
issues and caveats for interpretation. For illustrations of the use of FFTs
to analyze EGG signals, see chapter 11 and Stern, Koch, and Vasey
(1990). The reader desiring additional information about FFTs should
consult Lyons (1997, chapter 4).
Coherence Analysis
At times researchers are interested in the question of how two sep-
arate physiological signals might be related. The two panels of figure
14.10 depict data from respiration and the heart. Although the graphs
suggest that the signals are similar, there are computational methods
which can help you to determine how similar they are. One method is
that of coherence (see Forges & Bohrer, 1990, for a description of the
method). An easy way to understand coherence is to see it as a corre-
lation procedure for relating two signals. In addition, the correlation is
squared and thus coherence values range from 0 to 1.00. The upper
panel of figure 14.11 shows the same data depicted in figure 14.10.
However, the lower panel of figure 14.11 now shows a coherence plot

of the two signals in the upper panel. To provide a better understanding
of coherence, let us take another example: brain activity, namely EEC.
In this case we want to know if two signals coming from two parts of
the brain are similar. You might first ask if the two signals are of the
same frequency. The measure of coherence gives you the answer to this
question by determining the frequency components that make up each
signal (like an FFT analysis) and then examining the correlation between
the two signals at each frequency. Thus, two signals can be alike at one
frequency but not necessarily at another. You can also have a situation
in which the signal recorded over one part of the brain is similar to that
over another part except that the signals do not occur synchronously—
instead one signal is delayed behind the other one. A coherence analysis
Figure 14.8. Time and frequency domain depictions of sine waves. The left
panels illustrate sine waves in the time domain and the frequency domain
characterizations of those waves are shown in the right panels, (a) A sine
wave with an amplitude of 1 and a frequency of f(). (b) A sine wave with an
amplitude of 0.4 and a frequency of 2f0. (c) The sum of the two sine waves
shown in (a) and (b) and how both frequencies and amplitudes are revealed
by the frequency domain depiction on the right. Reprinted with permission
from R. G. Lyons, 1997, Understanding digital signal processing, Reading, MA:
Addison-Wesley.

Figure 14.9. Two frequency domain depictions of sine waves. These frequency
domain illustrations are based on the sine waves shown in Figure 14.8. The
left panel is identical to the depiction in the right side of Figure 14.8 (c) and
illustrates the amplitude and frequency components of the sum of two sine
waves. The right panel shows a different frequency domain depiction where
amplitude has been squared to produce power. This depiction shows the result
of a FFT analysis. Reprinted with permission from R. G. Lyons, 1997, Under-
standing digital signal processing, Reading, MA: Addison-Wesley.
can detect this delay or lag between two signals (called the phase angle).
Using this analysis we could determine that EEC over one brain region
is delayed by some defined period from that over another region. An
example of the use of coherence analysis of the EEC during a movement
task can be found in Ford, Goethe, and Dekker (1986).
Wavelet Analysis
Wavelet analysis can be viewed as being similar to Fourier analysis in
that they are both means of transforming data unfolding over time into
a series of frequency components. However, wavelet analysis can be used
to assess the frequency components of data that are not amenable to
Fourier analysis, which is problematic for viewing short-term changes in
a physiological signal. Indeed, FFT requires a signal to be relatively stable
or stationary over the time period from which frequency components are
derived.
Figure 14.12 shows the output of a wavelet analysis (bottom plot) for
a signal (top plot) that changes in frequency during the time period.
Notice how the changes in the signal are reflected immediately in the
wavelet analysis. The basic idea is that rather than examining a signal
in terms of sine waves of varying frequencies (as in FFT), various time
segments of the signal are instead viewed as shifted and scaled versions
of a particular mathematical function, namely, the wavelet. Such a pro-
cedure is useful for examining in detail an aspect of the physiological
signal that is different from that preceding and following it. For example,
if you were to look at the optical illusion that consists either of two faces
or a vase depending on your perception, wavelet analysis would allow

you to examine the EEC both before and after the optical illusion switched
from one perception to the other. Figure 14.13 shows such an analysis
of the change in EEC with the "flipping" of a Necker cube illusion from
one perspective to the other. For a recent discussion of the use of wavelet
analysis see Samar, Bopardikar, Rao, and Swartz (1999).
Figure 14.10. Power spectra of two physiological signals, (a) Two physiolog-
ical signals, respiration and heart period, over a 2-min period, (b) The power
density spectra for the two physiological signals shown in (a). Note the close
concordance between the prominent peak in the respiratory spectrum and
the higher frequency peak in the heart period spectrum. These peaks co-occur
with the predominant respiratory frequency. Reprinted with permission from
S. W. Forges and R. E. Bohrer, 1990, "The analysis of periodic processes in
psychophysiological research." In J. T. Cacioppo and L. G. Tassinary (Eds.);
Principles of psychophysiology: Physical, social and inferential elements, New
York: Cambridge University Press.
Figure 14.11. Coherence analysis of two physiological signals. The top panel
shows the respiration and heart period data from Figure 14.10. The bottom
panel depicts the coherence analysis between the respiration and heart pe-
riod. Note that coherence is high (i.e., the correlation is strong) over some
portions of the epoch, but weaker over other portions. Reprinted with per-
mission from S. W. Forges and R. E. Bohrer, 1990, "The analysis of periodic
processes in psychophysiological research." In J. T. Cacioppo and L. G. Tas-
sinary (Eds.), Principles of psychophysiology: Physical, social, and inferential ele-
ments, New York: Cambridge University Press.
Nonlinear Dynamical Systems (Chaos)
Analysis
In contrast to the traditional signal processing procedures that decompose
a signal like the EEC into its component frequencies and thus reflect a
limited amount of information (one dimensional), the dynamical systems
view suggests that a time series reflects the effects of all other variables
participating in the dynamics of the system. The theoretical basis for this
view derives from a variety of mathematical theorems (e.g., Whitney
embedding theorem). Given that complex dynamic systems (such as the
human nervous system) have an enormous number of interrelated de-
pendent variables which are impossible to measure directly, the theorems
suggest that if we can measure any single variable with sufficient accu-
racy, sufficiently often, and for sufficiently long periods of time, then it is
possible to make quantitatively meaningful inferences about the dynamic
structure of the entire system from the behavior of that single variable.
From this perspective we have a theoretical foundation to explain why
the nonlinear dynamic or chaotic approach may offer a characterization
of behavior that is richer than that obtained by classical measures. One
important nonlinear, dynamical measure of interest is that of dimension-
ality, referred to as d. A periodic oscillation (e.g., a sine wave) would
have a dimension of 1. A quasi-periodic oscillation (e.g., two dispropor-
tionate frequencies) would result in a dimension of 2. Truly random noise
would have a dimension approaching infinity. Many studies have used
dimensionality as a measure of neural processing. Lutzenberger, Preissl,
and Pulvermuller (1995) suggest that the dimensionality measure rep-
resents a relative measure of the number of neural ensembles recruited
Figure 14.12. Output of a wavelet analysis (bottom plot) for a signal (top
plot) that changes in frequency quickly during the time period. Notice how
the changes in the signal are reflected immediately in the wavelet analysis.

Figure 14.13. Wavelet analysis of EEC as the Necker cube illusion "switched."
The top plot represents the output of the wavelet analysis in terms of time
and frequency. The middle plot is EEC data recorded at the left occipital site
of the scalp. The participant moved her finger slightly when the Necker cube
"switched"; this was measured with an accelerometer, as shown in the bot-
tom panel.
during a specific cognitive task. Applying this thinking on a global brain

level, one might, for example, expect to see creativity to be associated
with greater dimensionality. In terms of cardiovascular processes, given
that the healthy heart shows greater variability than the abnormal one,
one important question is whether regularity or the lack of nonlinear
processing might be associated with the lack of health. An interesting
theoretical application of the nonlinear approach is that of psychopa-
thology (Globus & Arpaia, 1994). For an introduction to nonlinear dy-
namics especially in relation to physiology, see Elbert et al. (1994); Prit-
chard & Duke (1995); and Theiler (1990).
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PART III
Applications
15
Applications of Psychophysiological
Recording
Once an individual is able to record psychophysiological measures suc-

cessfully, the question arises as to how these measures might be used in
understanding the interdependence between psychological and physio-
logical processes. What fields of study might significantly benefit from
psychophysiological approaches?
Before beginning a discussion of applications of psychophysiological
recording, we wish to refer back to some of the principles of psycho-
physiology introduced in chapter 5. Researchers have at times ignored
these principles—these complexities—and have attempted to place such
concepts as emotionality or arousal along a continuum. These same re-
searchers would then try in vain to find a single physiological measure
that would show a high correlation with a given psychophysiological
concept. These simplistic studies were performed even in the midst of
more appropriate conceptualizations of psychophysiological functioning.
For example, in his review of psychophysiological approaches to the eval-
uation of psychotherapy outcome, Lacey (1959) emphasized that no one
measure of bodily arousal was adequate in relation to either psycholog-
ical process variables or other physiological measures. In this review and
elsewhere, Lacey (as well as other pioneers Chester Darrow, and R. C.
Davis) suggested that there are patterns of psychophysiological respond-
ing. This patterning is related to both the type of external stimuli (either
sensory or "ideational," in Darrow's terminology, or "stimulus response
specificity" in Lacey's) and the individual's physiological response pat-
tern, which Lacey referred to as individual response stereotypy. Although
Darrow (1929) presented his conceptualizations more than seventy years
ago, today one still sees researchers attempting to use a single psycho-
physiological measure as an indicant of emotionality and arousal without
regard to either situational or individual variables. With this cautionary
245
note, let us now examine some examples of recent psychophysiological
research.
Five Categories of
Psychophysiological Studies
If one were to review the journal Psychophysiology as well as books in

psychophysiology one could divide psychophysiological studies into those
that emphasize any one of five separate aspects. These categories are not
seen as mutually exclusive or all inclusive but are presented here to offer
an organizing principle for discussing recent psychophysiological re-
search. These approaches, or emphases, are: (1) response variables, (2)
stimulus/situational variables, (3) subject variables, (4) correlational var-
iables, and (5) the applications of psychophysiological research.
Response Variables
These studies are primarily concerned with properties of the psycho-
physiological response. Examples include examining the relation between
electroencephalogram (EEC) alpha activity and metabolic activity in the
thalamus, because EEC alpha activity has been thought to be modulated
by the thalamus (Larson et al., 1998); neuroanatomical correlates of
electrodermal activity (Fredrikson et al., 1998), the study of pulse wave
transit time and its relationship to blood pressure (Steptoe, Smulyan, &
Gribbin, 1976); ways of measuring the psychophysiological response, as
in the case of frontalis EMG placements (Davis, Brickett, Stern, & Kimball,
1978); problems of instrumentation, as in the effect of time constants on
the evoked potential P300 component (Duncan-Johnson & Donchin,
1979); problems of interpretation and quantification, as in the differences
in heart rate and heart period measurements (Graham, 1978; Quigley &
Berntson, 1996); and methods of statistical analysis using more than one
occurrence or more than one psychophysiological measure (Vasey &
Thayer, 1987). From its beginning in 1964, Psychophysiology has con-
sidered these questions to be at the heart of psychophysiological research
and the journal includes a separate section for advances in instrumen-
tation as well as methodology.
Stimulus/Situational Variables
Studies that emphasize stimulus/situational variables are generally de-
signed to examine differential psychophysiological responding as a func-
tion of different types of stimuli, either internal or external to the indi-
vidual. These studies predate the formal study of psychophysiology and
246 APPLICATIONS
are among the first psychological studies of the last century. For example,
in 1888 Fere published a report on the effects of stimuli such as colored
glasses, the sounds of tuning forks, and stimuli involving taste and smell
on electrodermal responses (Fere, 1888/1976). C. G. Jung, at the turn of
the twentieth century, noted the electrodermal response to various emo-
tionally arousing and neutral words. Other studies noted the effects of
music on heart rate and respiration. In the twentieth century, Darrow
(1929) reviewed the research concerning psychophysiological reactions
to ideational and sensory stimuli. This tradition has been continued by
a variety of researchers (e.g., Ray & Cole, 1985; Turpin, 1986).
Subject Variables
Studies that examine these variables emphasize particular characteristics
of the subjects themselves. These characteristics may be subdivided into
five general classes: (1) state factors, (2) trait factors, (3) psychopatho-
logical factors, (4) organizational factors, and (5) evolving factors. State
variables emphasize certain naturally occurring states within the indi-
vidual. One commonly researched state factor is sleep. For example, Bur-
gess, Kleiman, and Trinder (1999) examined cardiac activity during sleep
onset; McDonald, Shallenberger, Kortesko, and Kinzy (1976) recorded
spontaneous electrodermal responses during sleep. Other studies have
looked at such states as hypnosis (e.g., Graffin, Ray, & Lundy, 1995;
Dumas, 1977; Hilgard et al., 1974), and menstrual cycle (e.g., Little &
Zahn, 1974; see Bell, Christie, & Venables, 1975, for a review of this
area).
Trait or personality factors emphasize those characteristics of an in-
dividual which are seen as persisting in a variety of situations. One ap-
proach has been to examine psychophysiological variables in twins. An
example of this work would be an examination of EEG characteristics in
monozygotic and dizygotic twins (Christian et al., 1996). Other studies
have examined individual difference characteristics such as hostility (Sul
& Wan, 1993), locus of control (Ray, 1974), stimulation seeking (Zuck-
erman, Murtaugh, and Siegel, 1974), and anxiety (Heller, Nitschke,
Etienne, & Miller, 1997; Neary and Zuckerman, 1976).
Underlying many of the specific studies of personality or trait variables
are the numerous theories of personality types, such as that of Jung
(1971) who in 1920 divided people into thinking, feeling, sensing, and
intuitive types; Eysenck (1953), who classified people in terms of neu-
roticism and introversion/extroversion; and Pavlov (1928), who dis-
cussed temperaments in terms of Hippocrates' four types (i.e., choleric,
melancholic, sanguine, and phlegmatic). All of these theories had impli-
cations for psychophysiological research. In the last part of the twentieth
century, measures of personality involving the "Big Five" (extraversion,
neuroticism, openness, agreeableness, and conscientiousness) have be-
A P P L I C A T I O N S OF P S Y C H O P H Y S I O L O G I C A L RECORDING 247
come the standard measure of individual differences and are sparking a
variety of psychophysiological studies.
The third class of subject variables is the psychopathological or phy-
siopathological factors. In studies emphasizing these factors, subjects are
divided according to some preestablished diagnostic system, such as the
Diagnostic and statistical manual of mental disorders (4th ed.) (DSM) (APA,
1994). These systems include both psychological disorders such as
schizophrenia (Clementz, 1996; R6(3ner, Rockstroh, Cohen, Wagner, &
Elbert, 1999; Yee, Nuechterlein, & Dawson, 1998) and psychopathy
(Blackburn, 1979; Raskin and Hare, 1978; Williamson, Harpur, & Hare,
1991), as well as physiological disorders such as ulcers, hypertension,
and headaches (Shapiro, Jammer, & Goldstein, 1997). The task of the
psychophysiological researcher is to determine the important parameters
of the psychological-physiological interface. One approach is to examine
individuals who are unable to identify or communicate affective experi-
ences (Roedema & Simons, 1999).
A fourth category is that of organizational differences, including such
variables as race and gender. Although most psychophysiological studies
have been performed with Caucasians who are male, there is evidence
to suggest that both race and gender play a role in differential psycho-
physiological responding (Vrana & Rollock, 1998). Electrodermal activity
in a variety of studies has been shown to be sensitive to racial differences
(cf. Fisher & Kotses, 1973; James, Worland, and Stern, 1976; Lieblich,
Kugelmass, and Ben-Shakhar, 1973), as has the electrogastrogram (EGG)
(e.g., Stern, Breen, Watanabe, & Perry, 1996). Likewise, gender differ-
ences have been shown to be a significant variable in a number of psy-
chophysiological studies (e.g., Beaumont & Mayes, 1977; Davidson &
Schwartz, 1976; Heiman, 1977; Ketterer & Smith, 1977; Ray, Morell,
Frediani, & Tucker, 1976). Both race and gender variables are extremely
relevant in attempting to understand how hormonal, nervous system,
and experimental influences interact to produce the observed psycho-
physiological response.
We refer to the fifth class of subject variables to as evolving factors.
This category includes studies that examine a psychophysiological vari-
able as it evolves over time. The clearest example of this type of research
is psychophysiological studies within the area of lifespan development.
So far, research in this area has focused on the differences between the
electrophysiological responding of older and younger individuals. For ex-
ample, Morgan Geisler, Covington, Polich, & Murphy (1999) examined
event-related potential differences in young (age 20s) and older (age 60s)
adults. The examination of age differences in EEC dates back to the 1930s
and the work of Hans Berger. Other psychophysiological research has
attempted to determine psychophysiological responses in the developing
human organism. For example, Schaefer (1975) used EEC to examine
248 APPLICATIONS
neonatal responses to external stimulation; Graham and Jackson (1970)
have used heart rate measures.
Correlational Variables
For many researchers, these factors are both the most interesting to discuss
and the most difficult to study. They are interesting to discuss because they
include such traditional philosophical issues as the mind-brain question
and the relation of bodily change to emotion and behavior. But, they are
difficult to research because, by definition, these studies attempt to un-
derstand activity on one level of human activity from its correlation with
activity on another level. The most commonly used classification of ac-
tivity levels is a modern version of the Platonic organization presented in
chapter 1. The modern version divides psychological activity into (1) cog-
nitive activity, which may include so-called higher activity such as cre-
ativity, awareness, and consciousness; (2) emotional activity; and (3) in-
stinctual activity. Whereas the psychological researcher correlates activity
from one of these three levels with the other, the psychophysiological re-
searcher correlates each with physiological responding. Psychophysiology
offers researchers a window into cognitive and emotional processing. One
fruitful area has been the study of cognition and attention (see Osman,
1998, for an overview). Another area has been that of language proc-
essing, particularly the use of electrocortical measures to study language
structure and expectation (see Kutas, 1997, for an overview).
Another example of this type of research is found in the area of brain
lateralization and activation. Doyle, Ornstein, and Galin (1974) suggested
that EEC power relationships between the two cerebral hemispheres could
be used to differentiate different types of cognitive processing, such as
linguistic versus spatial processing. In a similar vein, a variety of studies
have suggested that EEC measures might also be related to the processing
of negative or positive emotional material (Davidson, 1995). Other stud-
ies outside of the hemispheric paradigm have sought to discover corre-
lations between the electrical activity of the brain and such concepts as
creativity, altered states of consciousness, and meditative experiences.
Psychophysiological correlations with instinctual activity such as sexu-
ality have also been observed and recorded. Although there are many
studies on the correlations of cognitive, emotional, and instinctual be-
havior with psychophysiological variables, both the basic physiological
mechanisms and the philosophical questions concerning the similarities
among these levels of analysis remain unresolved.
Application Studies
The final category we will discuss includes those studies that attempt to
understand psychophysiological principles as they are applied in real-
A P P L I C A T I O N S OF PSYCHOPHYSIOLOGICAL RECORDING 249

world settings. Three examples these types of research are biofeedback,
lie detection, and brain plasticity and pain.
Biofeedback. Until the 1960s, it was assumed that autonomic nervous

system (ANS) responses had to be elicited rather than emitted, thus in-
volving the procedures of classical conditioning. Central nervous system
(CNS) responses, on the other hand, were considered to be voluntary and
thus reinforced by the techniques of operant conditioning. These specu-
lations were largely untested until Miller (1963) questioned the tradi-
tional differences between classical and operant conditioning. Miller sug-
gested that there is really only one type of learning; classical and operant
conditioning are two manifestations of the same phenomenon under dif-
ferent conditions. It follows that if there is only one type of learning,
autonomic responses should be conditionable through operant methods.
Thus, the 1960s saw a number of research studies directed at this ques-
tion.
The initial studies were performed with animals. In order to demon-
strate autonomic conditioning, it was first thought necessary to show
that there was no somatic intervention. For example, if one wanted to
condition the heart, it would be important to show that the heart rate
changes were not due to respiratory changes. The control of somatic
mediation was achieved by paralyzing the animal's CNS.
However, when working with human beings, the question became
more difficult, as the reviews by Kimmel (1967) and Katkin and Murray
(1968) point out. Not only was there a possibility of somatic mediation,
but there was also the question of cognitive mediation. The question
became how to conceptualize the situation in which arousing or angry
thoughts produced a change in heart rate. Could this be considered an
example of human autonomic conditioning? Although there was much
debate, the question was never really answered.
The conditioning question was reformulated into a question of control.
That is, rather than asking if autonomic conditioning were possible, re-
searchers looked to find under what conditions an individual could dem-
onstrate physiological self-control. Research in "operant conditioning"
was replaced by an interest in "biofeedback." Biofeedback is a deceptively
simple technique in which information from physiological responses such
as EEC, muscle tension, and so forth is fed back to the individual, with
the final goal being the self-regulation of the physiological function.
The transition from operant conditioning to biofeedback involved a
movement from theory to practicality. Initial research questions focused
on the type of feedback that was optimal and the importance of aware-
ness in learning self-control. Clinical researchers suggested that the man-
ner in which patients self-regulated their hypertension, for example, was
less important than the possibility of returning their blood pressure to
within normal limits. Biofeedback became the new panacea for a number
250 APPLICATIONS
of disorders including headaches, epilepsy, cardiac arrhythmias, and den-
tal and neuromuscular disorders. However, 10 years after the first bio-
feedback studies, the therapeutic efficacy was still in question for a large
number of disorders (Ray, Raczynski, Rogers, & Kimball, 1979) and re-
mains so today. However, there are a number of success stories which
make this work intriguing. For example, a variety of studies from the
Tubingen lab of Niels Birbaumer reported that teaching epileptics to mod-
ify slow-wave EEC behavior resulted in a reduction of seizures. These
studies were also performed with children with attentional disorders as
well as healthy volunteers (see Rockstroh, Elber, Canavan, Lutzenberger,
& Birbaumer, 1989 for a review). In these studies, slow-wave activity
was recorded along the midline of the cortex. Changes in the EEC moved
a symbolic rocket ship from left to right on a computer screen during a
6-s period. The task was to direct the rocket ship toward one of two
locations on the screen through the generation or suppression of cortical
negativity. The results from these studies suggest that individuals can
learn to control slow cortical potentials in both a negative and positive
direction within about 80 to 160 feedback trials. Another important use
of EEC biofeedback technology has been to teach paralyzed individuals to
communicate. Birbaumer and his colleagues describe the case of an in-
dividual completed paralyzed who was able to use his brain waves to
slowly spell out words and communicate with the outside world (Birbau-
mer et al, 1999).
Lie Detection. Lie detection refers to a variety of procedures whereby

ANS measures such as blood pressure, pulse rate, and electrodermal ac-
tivity are recorded from an individual while the person is asked two types
of questions: relevant and comparison. The relevant questions deal with
specifics of a crime under investigation or some aspect of the individual's
personal life. The latter variety are sometimes used in preemployment
screening by government agencies that deal with security matters. The
use of lie detection in preemployment screening by private companies has
been outlawed in the United States.
Comparison questions are always included in the interrogation process
in order to determine the magnitude of the individual's ANS responses
in answering nonrelevant questions alone. As we discussed in chapter 5,
individual responding using various psychophysiological measures varies
greatly from person to person. Without this information, an individual
might be falsely accused of being guilty or suppressing information just
because of large ANS responses made to all questions.
The formulation of the questions to be asked a crime suspect or job
applicant for a position in a governmental security agency is a highly
technical and somewhat controversial matter. Raskin and his colleagues
(e.g., Podlesny and Raskin, 1977) recommend the use of the control-
question technique. The control questions (used instead of neutral ques-

tions), formulated during a pretest interview, are selected so that the
individual will probably be deceptive or at least very concerned about
them. An example of a control question used in a theft case would be,
"During the first eighteen years of your life, did you ever steal something
from someone who trusted you?" The rationale for this technique is ex-
plained by Podlesny and Raskin (1977): "The control question approach
attempts to set up a situation wherein a subject who is truthful concern-
ing relevant questions will be more concerned about the control questions
and will produce greater responses to them than to the relevant ques-
tions. Similarly, deception on relevant questions should make the subject
more responsive to relevant than to control questions" (p. 786). Another
discussion of the control-question technique can be found in Honts,
Kircher, and Raskin (1995). The interested reader is also referred to an
article by Furedy (1996) in which he raises ethical problems that he
claims are inherent in the use of the control-question method.
Lykken (1974, 1979, 1991) has questioned many of the claims and
assumptions of proponents of traditional lie detection procedures such as
the control-question test. According to Lykken, psychophysiological re-
cordings cannot and should not be used to determine whether or not
someone is lying. He suggests an alternative and related use, however.
Lykken's procedure, referred to as the guilty-knowledge test, is based
upon the presence or absence of differential responsivity to items of in-
formation that only the guilty suspect would recognize as being relevant.
If the person being questioned did not murder the girl in the red dress,
a question about red dresses should bring about ANS responses no larger
than responses to a question about blue dresses.
We believe that there are two assumptions that must be examined in
order to test the validity of any of these procedures. The first assumption
is that lying (or possessing guilty knowledge) will cause heightened ANS
responding. This is true for most but not all individuals. The second as-
sumption is that the ANS responses used in the lie detection procedure
are not under the suspect's voluntary control. This assumption is more
questionable, particularly in light of work in the area of biofeedback (see
chapter 20 in Stern & Ray, 1977). It should be noted that individuals
can learn, through the use of biofeedback over many trials, to control
their ANS responses, but when participants are provided with feedback
of their physiological responses during an interrogation, they are easier
to detect than subjects in a no-feedback control group (Stern, Breen,
Watanabe, & Perry, 1981).
The psychophysiological response measures used in lie detection sit-
uations are more relevant to this book than are the interrogation pro-
cedures. The following response variables have been used in laboratory
and field tests: respiration, blood pressure, electrodermal activity, pulse
rate, pulse volume, muscular tension, eye movement, and eye blinks.
Curiously, in laboratory tests, electrodermal activity is usually the best
252 APPLICATIONS
single indicator of deception, but in actual field use, the cardiovascular
measures discriminate best. This apparent discrepancy may be a function
of basic differences in laboratory and field situations. These differences
include level of emotional involvement of the subjects and differences
between the background and training of the people doing the interro-
gating. Laboratory lie detection studies are usually conducted by psy-
chologists interested in psychophysiology. Field testing in lie detection is
usually done by individuals with a background in police work. These two
groups of investigators not only have different backgrounds and goals
but also use different equipment and recording techniques. They have a
lot to offer each other but, unfortunately, few individuals belong to both
the American Polygraph Association and to scientific psychological re-
search societies.
Lie detection raises a number of important questions for psychophys-
iologists to consider. Some of these questions are as follows:
1. Does a person's physiology actually mirror that person's state of

emotional arousal?
2. Can we differentiate between increases in emotional arousal caused
by lying versus other states such as worry or embarrassment?
3. Are there some individuals, such as psychopaths, for whom we can-
not use the lie detector?
4. Can a person, through biofeedback or other techniques, learn to
modify his physiological responding and thus "fool" a lie detection
operator?
5. To what extent can augmented physiological feedback aid in the
detection of deception?
6. What does traditional theory in psychophysiology concerning ha-
bituation, sensitization, motivation, stimulus presentation, arousal,
and so forth have to offer the field of lie detection, and vice versa?
Lie detection has received only minor interest by psychophysiologists,

the result being little long-term systematic research in the area. Yet as
an area with both applied and theoretical implications, it appears to offer
the psychophysiological researcher numerous opportunities and chal-
lenges. For a bibliography of lie detection, see Ansley and Horvath
(1977). Other reviews that may be consulted include Davis (1961) and
Orne, Thackray, and Paskewitz (1972).
Cognitive Neuroscience: Cognitive neuroscience is a multileveled ap-

proach to the understanding of the mind, including cognitive, emotional,
and motor processes. Psychophysiology has emerged as an important tool
that aids researchers and clinicians in describing the underlying mech-
anisms in both normal and pathological functions. In this section we will
sample a variety of cognitive areas that have made use of psychophysi-
ological techniques.
Cortical plasticity and reorganization after injury is one area that has
used psychophysiological approaches, especially brain mapping tech-
niques, to obtain a better understanding of brain processes. If a person
is in an accident that results in the removal of an arm or leg, the area
of the brain that was associated with bodily sensation and movement of
that limb changes and reorganizes itself in a new way. Using magne-
toencephalogram (MEG) techniques, Ramachandran and his colleagues
(see Ramachandran & Hirstein, 1998, for a review) described four indi-
viduals who had their arms amputated. In all four individuals, these
researchers found that the area of the cortex associated with the hand
changed to become associated with the face and upper arm. In fact, if a
cotton swab was lightly rubbed on one individual's lower face, he re-
ported feeling sensations in the missing arm. Likewise, warm water trick-
ling down the face would be experienced both on the face and down the
length of the missing arm. Given that a traumatic experience such as
losing an arm could cause cortical reorganization, one could also ask if
normal experience itself could have an influence. The answer to this
question turns out to be yes. Using MEG techniques, Elbert and his col-
leagues compared string players who had played instruments throughout
their lives with a group of controls (Elbert, Panter, Wienbruch, Rock-
stroh, & Taub, 1995). Not only was the cortical representation of the
fingers of the left hand of string players larger than that in controls, but
the amount of cortical reorganization in the representation of the finger-
ing digits was correlated with the age at which the person had begun to
play.
Another area that has benefited from psychophysiological techniques
is that of attention. An interesting question in attention is how we pick
out one aspect of the multitude of sounds, images, sensations, and so on
that constantly confront us (see Luck, 1998 for an overview of this ques-
tion). A great variety of ERP studies have been performed to determine
when particular types of information are processed in the cortex. For
example, Hillyard and colleagues (1973) studied the influence of attend-
ing or not attending to a particular stimulus. They found that when one
attends to a stimulus, the first negative component (Nl) of the ERP is
larger than when one did not. Since this Nl component peaks at ap-
proximately 100 ms after a stimulus presentation, these researchers con-
cluded that selective attention acts at a very early stage of processing.
Another type of attention research has examined if deaf or blind individ-
uals have enhanced capabilities of sensory processing. For example,
Roder and colleagues (1999) report a differential pattern of Nl activity
across the scalp in individuals who are blind versus sighted individuals
with a blindfold. They also reported localization abilities of blind individ-
uals to be superior to sighted ones, but only when the stimuli were pre-
sented in the periphery.
254 APPLICATIONS
As we look at an object, we see it as a whole even through various
aspects of it (color, smell, taste, texture, complexity, etc.) are encoded and
processed in different parts of the brain. This question has come to be
called the "binding" question in that it asks how the brain puts or binds
together various aspects of an experience to produce a coherent whole.
One answer to this question has come from EEC research. A variety of
studies have shown that oscillatory synchronization of the EEC in the
gamma band (30-80 Hz) is associated with feature binding in a variety
of modalities including vision (see Singer & Gray, 1995; Tallon-Baudry
& Bertrand, 1999, for a review of this work). Using EEC measures and
wavelet analysis, Tallon-Baudry and her colleagues showed enhanced
gamma band activity when an individual recognized a hidden figure (a
Dalmatian dog) but not when the figure appeared as meaningless black
blobs on a grey background. Likewise, Keil Miiller, Ray, Gruber, & Elbert
(1999) showed enhanced gamma band activity when an ambiguous fig-
ure (either a happy or sad face depending on orientation) switched to a
new perspective. Using classical conditioning to establish a relationship
between a visual and tactile stimulus, Miltner Braun, Witte, & Taub
(1999) showed greater EEC gamma band coherence between areas in
the brain that are involved in visual and tactile experiences. Once the
conditioning relationship between the visual and tactile stimuli was ex-
tinguished, the EEC coherence decreased.
In each of the areas discussed—cortical reorganization, attention, and
binding—psychophysiological techniques helped to answer questions
concerning temporal and spatial aspects of the process. In these examples,
the evoked potential was particularly important for understanding when
certain types of processing took place, whereas the brain mapping tech-
niques were useful for describing which areas were involved in a partic-
ular activity. Other areas of research, such as motor processing and emo-
tional experience and expression, have also benefited from psycho-
physiological studies; the interested student should consult such journals
as Psychophysiology for reports of recent work.
Conclusions
With the availability of small laboratory computers and newer instru-

ments for the recording of physiological measures, it is clear that we will
see an increasing number of psychophysiological studies being performed
in traditional areas of psychology. We also predict an increase in real-
world applications of psychophysiology as greater use is made of telem-
etry, ambulatory recordings, and nonintrusive recording techniques
(Fahrenberg & Myrtek, 1996). As mentioned previously, psychophysio-
logical studies in the area of lifespan development, especially aging and

neonatal development, are beginning to increase in both number and
sophistication. Other areas, such as biofeedback and behavioral medicine,
sleep, and pain, are continuing to draw the interest of researchers and
clinicians who can make significant contributions based on their under-
standing of psychophysiological principles.
Just as psychophysiologists are able to contribute to new and devel-
oping areas, they must also be ready to integrate new aspects of these
areas into their research. For example, the standard statistical procedure
of determining significance through the use of techniques such as anal-
ysis of variance may be combined with newer nonlinear models of human
functioning. For example, researchers in the area of behavioral medicine
ask how a number of treatment techniques can combine to form a sum
greater than any of the parts alone. Likewise, developmental studies dem-
onstrate the importance of certain changes of physiological state (e.g.,
puberty) that result in irreversible processes impossible to discuss from a
simple acquisition and extinction paradigm.
We end this book as we began. In the beginning, we indicated that
psychophysiology had a short history and a long past. We described the
subject matter of psychophysiology as not new but as represented by
broad questions studied for centuries by individuals trained as philoso-
phers, physicists, physicians, physiologists, and psychologists. We would
like to point out that as the name implies, psychophysiology is at the
heart of one of the most fascinating and enduring of these issues: the
relationship between mind and body. It is our hope that with the aid of
psychophysiological recording you will gain insights into the basic nature
of this and other fundamental questions.
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A P P L I C A T I O N S OF P S Y C H O P H Y S I O L O G I C A L R E C O R D I N G 261
Glossary
Acetylcholine: neurotransmitter at all parasympathetic postganglionic and all

preganglionic peripheral synapses and at the neuromuscular junction.
Actin: contractile protein in muscle fibrils; functions in conjunction with my-
osin.
Action Potential: sequence of changes in potential associated with impulse
conduction in nerves and muscles.
Adrenaline: see Epinephrine.
Adrenal Medulla: portion of the adrenal gland which secretes epinephrine.
Aliasing: faulty representation of a digitized analog signal due to an insuffi-
cient sampling rate; an aliased signal will contain spurious frequency com-
ponents that are not present in the original analog signal and will not contain
some higher frequency components present in the original signal.
Alpha Blocking: the change in the EEC from alpha to other rhythms, usually
beta.
Alpha Rhythm: relatively large, rhythmic brain waves of approximately 8-12
Hz, as recorded in an EEC; thought to be related to relaxation.
Analog signal: a continuous physiological signal.
Analog-to-Digital (A-D) Converter: device which converts a continuous physi-
ological signal into discrete steps, enabling a computer to quantify the data.
Antrum: lower part of the stomach, from which EGG is recorded.
Arousal: often used synonymously with activation.
Artifact: stray and unwanted electrical signal in a physiological recording.
Atrioventricular Node (A-V Node): a small area of electrically active muscle in
the right atria in which excitatory impulses arriving from the sinoatrial node
are passed along to the conducting system of the heart.
263
Auditory Evoked Response: A predictable series of brain wave forms and dis-
tribution following each of a series of sounds; of maximum amplitude over
the vertex of the brain.
Autocorrelation: the correlation between a digitized physiological record and
itself lagged by some time period.
Autonomic Lability Score (ALS): a method of standardizing phasic ANS scores.
Autonomic Nervous System (ANS): ganglia, nerves, and plexuses which reg-
ulate activities of the viscera, heart, blood vessels, smooth muscle, and glands.
Autoregulation: one factor which increases the rate of blood flow during ex-
ercise; it responds to the local nutritional needs of the body.
Axon: neuron fiber processes that generally conduct impulses away from the
cell body; long neuron fibers.
Band-pass Filter: a combination of a highpass and a lowpass filter set so that
only a certain range of frequencies can pass.
Baroreceptors: sensory receptors found predominantly in the large arteries
which detect changes in pressure when deformed.
Beta Rhythm: rhythmic brain waves of approximately 13-30 Hz, as recorded
in an EEC, most often observed when the subject is alert.
Biofeedback: information concerning the functioning of internal organs, usu-
ally obtained through the use of electronic recording equipment.
Bioelectric Potential: a difference in electrical potential as recorded from dif-
ferent parts of a living organism.
Bipolar Recording: a recording method involving no reference electrode but
rather a comparison of two active sites.
Blood Volume: a measure of the relatively slow changes in the amount of
blood in an arm, finger, or other body structure.
Bradygastria: an abnormally slow rate of contractions of the stomach.
Brain Stem Evoked Response (BSER): an evoked response which originates in
the brain stem area.
Bucking Voltage: an off-setting voltage used to neutralize an undesirable stand-
ing potential.
Capacitance: the property that permits a body or circuit to store an electrical
charge; is equal to the stored charge divided by the voltage, and is expressed
in farads.
Cardiac Output (CO): the amount of blood circulated through the body per
unit of time (usually in liters/minute).
Cardiac-somatic Coupling: influence of motor activity on cardiovascular activ-
ity.
Cardiotachometer: a device for measuring heart rate that electronically deter-
mines the time from one QRS complex to the next and displays the infor-
mation graphically in terms of beats per minute.
264 GLOSSARY
Central Nervous System (CNS): neural material contained in the spinal cord
and brain.
Chaos: see nonlinear dynamical analysis
Chemoreceptors: sensory receptors sensitive to changes in the chemical milieu
surrounding the receptor.
Common Mode Rejection Ratio: a measure of the ability of a differential am-
plifier to reject noise or interference.
Compensatory Movement: movement of the head or body in response to ex-
ternal movement.
Contingent Negative Variation (CNV): a slow negative potential, recorded in an
EEC, typically occurring between a warning signal and a signal which in-
structs the subject to respond; similar to the readiness potential.
Corneal-retinal Potential: the potential between the cornea and retina of the
eye; the basis of the EOG.
Cortical: describes a function or process associated with the cerebral cortex.
Coupler: used in the first stage of some polygraphs.
Defensive Response: pattern of responding which serves to limit action of a
stimulus on an organism in order to protect the organism from possible dan-
gers of intense stimulation.
Delta Rhythm: low EEC frequencies (0.5-4 Hz) associated with sleep.
Dendrite: neuron fiber processes that generally conduct impulses toward the
cell body; short neuron fibers.
Depolarization: reduction or neutralization of polarity; typically occurs when
the inside of an electrically excitable cell becomes less negative relative to the
outside and therefore is closer to reaching threshold for initiating an action
potential.
Diastolic Blood Pressure (DBF): the minimal pressure in the vascular system
at the point where the measurement is taken; diastolic blood pressure occurs
at the lowest point of the pressure pulse; units of millimeters of mercury
(mmHg).
Digital Filtering: eliminates certain frequencies while others are allowed to
pass.
Digital signal: a signal recreated from sequential, discrete samples of an analog
physiological signal.
Directional Fractionation: a pattern of ANS responses characterized by some
physiological responses suggestive of decreased arousal (e.g., a heart rate
decrease) coupled with others suggestive of increased arousal (e.g., a skin
conductance response).
Eccrine Sweat Glands: sweat glands, located primarily in the palms of hands
and soles of feet, innervated by the sympathetic branch of the ANS; differ
from other sweat glands because they respond primarily to "psychic" stim-
ulation as opposed to increases in temperature.
GLOSSARY 265
Electrical Control Activity (EGA): basic electrical rhythm which controls the
rhythmic contractions of the lower part of the stomach (antrum) by making
the membrane temporarily susceptible to the generation of another potential
i.e., the electrical response activity (ERA).
Electrical Response Activity (ERA): electrical potential consisting of one or
more spikes superimposed on the decline phase of the electrical control ac-
tivity (EGA) and preceding contractions of the lower portion of the stomach
(antrum).
Electrocardiography (EKG or ECG): a technique for recording the electrical
activity of the heart.
Electrode: conductor through which electricity enters or leaves a medium.
Electrodermal Activity (EDA): a measure of various electrical properties of the
skin.
Electroencephalography (EEC): a technique for recording over time variations
in electrical potentials observed from electrodes on the scalp.
Electrogastrography (EGG): a method for recording gastric myoelectric activity
from the surface of the skin over the lower part of the stomach.
Electrolyte: a chemical compound, such as sodium chloride, that dissociates
into ions when dissolved, forming a conductor.
Electromyography (EMG): a technique for recording the time, voltage graph
of electrical potentials originating in muscles either from the surface of the
skin (surface EMG) or from electrodes inserted into the muscle.
Electrooculography (EOG): a technique which records changes in the cor-
nearetinal potential as a function of the movement of the eyes.
Epinephrine (also called Adrenaline): hormone secreted by the adrenal medulla
which stimulates glucose production; acts on sympathetically innervated or-
gans in a way similar to norepinephrine.
Evoked Response: a neural response to an abrupt stimulus such as a flash of
light.
Excitatory postsynaptic potential (EPSP): a depolarizing graded potential that
moves the cell membrane nearer to the threshold for initiating an action
potential.
Filter: a device which removes or reduces certain parts of the input signal
while allowing other parts of the signal to remain.
Fistulated: an animal with an artificial opening from an internal organ such
as the stomach to the outside.
Fluoroscope: a device used to view objects exposed to X-rays.
Fourier Analysis: a method of breaking down a time series into the sine waves
that make it up; a specific version of Fourier analysis often used by psycho-
physiologists is the Fast Fourier Transform or FFT.
Frequency domain techniques: techniques for analyzing data represented by
different frequencies (or in the frequency domain).
266 GLOSSARY
Gain: amount of amplification.
Galvanic Skin Response (GSR): term replaced by electrodermal activity (EDA).
Galvanometer: a device used to determine the presence, strength, and direction
of an electric current.
Gamma band activity: EEC activity in the 30 Hz to 70 Hz range assumed to
be related to the brain's ability to integrate a variety of stimuli into a coherent
whole.
Ganglion: a collection of nerve cells outside of the central nervous system.
Gastroparesis: an abnormal condition where the stomach fails to empty.
Gaze Nystagmus: see spontaneous nystagmus.
Generator Potential: the condition of a receptor after it has been excited and
partially depolarized. If depolarization exceeds the axonal threshold, the gen-
erator potential produces a neural impulse.
Graded Potential: a change in potential across the cell membrane that is pro-
portional to the amount of stimulation received. (See also excitatory postsy-
naptic potential and inhibitory postsynaptic potential.)
Ground: the connection (usually through a low-resistance conductor and the
round prong on a three-pronged plug) of an electrical circuit or device with
the earth.
Habituation: cessation of responding due to repeated presentation of the same
stimulus.
Heart period (HP) or Interbeat Interval (IBI): the time between two consecutive
heart beats; units usually in milliseconds.
Heart rate (HR): the rate of beating of the heart; units usually in beats/
minute.
Hering-Breuer Reflex: inspiratory reflex evoked by stretch receptors in lung
tissue that prevents overdistension of the lungs.
High-pass Filter: allows only frequencies above a certain frequency to pass.
Homeostatic State: steady-state internal environment providing the right tem-
perature, nourishment, oxygen, and fluids for optimum functioning of all cells
in a given region of the body.
Hormone: a chemical messenger which acts on receptors at relatively distant
sites from its source and which reaches the receptor by way of the blood
stream.
Hyperpolarization: enhancement of polarity; typically occurs when the inside
of a neuron becomes more negatively charged relative to the outside and
therefore less likely to generate an action potential.
Impedance: total resistance to current of an AC circuit; varies with the volume
of a conductor and other factors.
Impedance cardiography: a non-invasive technique whereby a high frequency,
low amplitude alternating current is passed through the chest to determine
GLOSSARY 267
stroke volume, cardiac output, and systolic time intervals (such as pre-
ejection period).
Individual Response Stereotypy: refers to the fact that individuals differ from
one another in their pattern of bodily response to different situations; a given
individual tends to show the greatest degree of activity in the same physio-
logical system no matter what the situation.
Inhibitory postsynaptic potential (IPSP): a hyperpolarizing graded potential that
moves the cell membrane further from the threshold for initiating an action
potential.
Inspiratory duty cycle (or inspiration fraction): the inspiratory duration divided
by the total duration of a respiratory cycle.
Integrated Circuit (1C): a multicomponent device (transistors, resistors, capac-
itors, etc.) in which the components have been miniaturized and designed for
specific purposes.
International 10-20 System: a system which makes possible standardized
placement of EEC electrodes on the scalp.
Interstitial Fluid: fluid found between the cells of the body; serves as a nutrient
medium for the cells.
Ion: atom which has lost or gained an electron and is thereby capable of
conducting electricity; a charged particle.
Joint Interval Histogram: method of presenting the results of frequency analysis
of an EEC.
Korotkoff Sounds: used to determine blood pressure; the sounds are caused by
turbulence in the blood as it spurts through the tiny arterial opening under
the blood pressure cuff during each systole.
Latency: duration from stimulus presentation to response onset.
Lateralized readiness potential (LRP): A cortical measure of preparation for a
motor response computed by subtracting the readiness potential of one hemi-
sphere from that of the other.
Law of Initial Values (LIV): states that the initial state of a physiological system
will limit the degree to which the system can change it's state; higher initial
levels will limit further increases in function and lower initial levels will limit
further decreases in function; this "law" is not always observed.
Low-pass Filter: allows only frequencies below a certain frequency to pass.
Magnetic resonance imaging (MRI): A system for measuring brain activity
based on blood flow increases in active areas of the cortex. Hemoglobin which
carries oxygen in the bloodstream has different magnetic properties before
and after oxygen is absorbed. Magnetic fields are measured in MRI in relation
to an external magnet.
Magnetoencephalogram (MEG): uses a SQUID (superconducting quantum in-
terference device) to detect the small magnetic field gradients exiting and
entering the surface of the head that are produced when neurons are active.
268 GLOSSARY
Mastoid Process: a nipple-shaped protrusion of the temporal bone located be-
hind the ear.
Mean Arterial Pressure (MAP): the average blood pressure in the vascular
system at the point where the measurement is taken; also diastolic blood
pressure plus one-third the pulse pressure; units of millimeters of mercury
(mmHg).
Mean Inspiratory Flow: the tidal volume divided by the duration of inspiration;
units typically milliliters/second.
Microsiemen: the unit of conductance that has replaced micromho.
Monopolar Recording: a recording method involving placement of one or more
electrodes at a site(s) where a relatively large amount of electrical activity is
generated, and the other electrode (known as the reference electrode) at a
site where electrical activity is minimal.
Motor (or Endplate) Potential: sudden depolarization across the motor endplate
of the neuromuscular junction that typically initiates an action potential in
the muscle.
Motor Unit: motor neuron and the muscle fibers it innervates.
Myelin Sheath: white, fatty substance that forms a sheath around the axons
of some nerves.
Myosin: contractile protein in muscle fibers; functions in conjunction with
actin.
Nodes of Ranvier: local constriction in the myelin sheath.
Nonlinear dynamical analysis: a broad theoretical approach that seeks to de-
scribe patterns in random appearing signal such as the EEC.
Norepinephrine (also called Noradrenaline): neurotransmitter at all sympathetic
postganglionic synapses (except sweat glands); also acts as a hormone when
released from the adrenal medulla.
Notch Filter: reduces only a small range of frequencies while allowing all
others to pass undiminished (e.g., 60 Hz notch filter).
Nystagmus Movement: oscillatory movement of the eye.
Off-line Computer Usage: use of a computer to analyze stored data; allows for
analysis without regard to how events happened in real time, as opposed to
on-line or real time usage.
Ohm's Law: the relationship between current, voltage and resistance in any
electrical circuit and described by the equation, I (current) = V (voltage) -=-
R (resistance).
On-Line: the functioning of a computer which is working in the same time
frame as the experiment it is controlling.
Optokinetic Nystagmus: oscillatory eye movements elicited by a moving pattern
containing repeated patterns.
Orienting Response: a specific pattern of reactions which occurs in response to
novel stimuli.
GLOSSARY 269
Outlier: a data point or observation that is extreme relative to the data from
the rest of the sample.
Pacemaker Cells (Pacesetter Potentials): cells which originate the electrical im-
pulse which produces contraction.
Parasympathetic Nervous System (PNS): craniosacral division of the autonomic
nervous system; usually activation of this system induces effects opposite to
those of the sympathetic nervous system.
Peripheral Nervous System: all neural material outside the brain and spinal
cord.
Positron emission tomography (PET): radioactive system to measure variations
in cerebral blood flow that are correlated with brain activity.
Phasic (or Event-related) Activity: a discrete response to a specific stimulus.
Photoconductive Cell: consists of a light source and a photoelectric cell which
varies in electrical activity in proportion to the amount of light that hits the
cell.
Pilomotor Muscles: those muscles that control elevation of body hair.
Plethysmography: the measurement of the size of a part of the body as a
function of the volume it contains (typically volume of blood or air).
Pneumograph: a distensible air-filled tube, used for measuring respiration, that
is placed around the subject's chest and connected to a pressure-sensitive
device, an amplifier, and a computer or recorder.
Polarization: the difference in charge between the outside and inside of a
membrane.
Polygraph: a device for recording two or more signals which are amplified
and then written out on paper.
Power Spectrum (or Spectral Density Plot): a depiction of a physiological signal
in the frequency domain that shows the component frequencies making up
that signal.
Preamplifier: a device which conditions and amplifies input signals before they
reach the power amplifier.
Pre-ejection Period (PEP): the time between the initiation of the electrical sig-
nal to the heart (usually the Q wave of the ECG) and the beginning of ejection
of blood from the left ventricle; units are milliseconds.
Pre-pulse Inhibition: The act of presenting a less intense stimulus prior to the
startle stimulus which decreases the startle response.
Psychogalvanic Reflex (PGR): an outdated term for skin conductance response.
Pulse Pressure (PP): the difference between the systolic blood pressure and
the diastolic pressure; i.e., systolic blood pressure minus diastolic pressure;
units of millimeters of mercury (mmHg).
Pulse Volume: a measure of the amplitude of individual pulses of blood in the
vascular system.
270 GLOSSARY
Pulse Wave Transit Time (PWTT): time it takes for a pulse wave to travel from
the heart to a distant location; related to blood pressure.
Pupillography: the measurement of the size of the pupil.
Readiness Potential: a slow negative potential, as recorded in an EEC, which
precedes (by as much as 1.5 sec) and accompanies movement or other re-
sponses.
Real Time: (See On-Line).
Receptor (or Generator) Potential: a graded change in potential across the
membrane after a receptor has been bound; if depolarization occurs and ex-
ceeds threshold, the receptor potential produces a neural impulse.
Resistance: a measure of the opposition that a conductor offers to the passage
of current; it is the reciprocal of conductance.
Respiratory Sinus Arrhythmia (RSA): the rhythmic increases and decreases in
heart rate produced by normal respiration; heart rate is increased during
inspiration and decreased during expiration.
Resting Potential: the state of a neuron when it is conducive to transmitting
a neural impulse specifically, when there is a difference in charge across its
membrane. Also called the polarized state.
Reticular Activating System (RAS): reticular formation, thalamus, hypothala-
mus, and related structures which function to maintain appropriate states of
arousal.
Saccadic Eye Movement: rapid jumping of the eyes from one fixation point to
another, as during reading.
Sampling Rate: frequency with which sequential discrete samples of an analog
physiological signal are recorded; units typically in Hz (samples/second).
S-A Node: see Sinoatrial Node.
Schmitt Trigger: a device which electronically checks for a certain voltage level
and then signals a computer when it occurs.
Sham feeding: feeding that does not reach the stomach. In humans modified
sham feeding is accomplished by having participants chew and then spit out
the food; in lower animals surgery is used to remove food that has been
swallowed from the gastrointestinal tract before it reaches the stomach.
Signal Conditioner: See Coupler.
Signal-to-noise Ratio: amount of signal (biopotential) in relation to other elec-
trical activity (noise).
Sinoatrial Node (S-A Node): a small strip of electrically active muscle located
in the upper part of the right atrium of the heart where the impulse that
produces contraction of the heart begins.
Skeletal Muscles: those that move the trunk and limbs.
Skin Conductance Level (SCL): the reciprocal of the measure of how much
resistance the skin of an organism in a state of rest or basal activity offers to
GLOSSARY 271
passage of an electrical current (i.e., the reciprocal of skin resistance level);
measured in mhos.
Skin Conductance Response (SCR): the reciprocal of the measure of how much
resistance the skin of an organism that is responding to a particular stimulus
offers to passage of an electrical current (i.e., the reciprocal of skin resistance
response); measured in mhos.
Skin Potential Level (SPL): measure of electrical activity at the surface of the
skin when the organism is in a state of rest or basal activity.
Skin Potential Response (SPR): measure of electrical activity at the surface of
the skin when the organism is responding to a specific stimulus.
Smooth Muscle: unstriped muscle usually located in visceral walls and blood
vessels.
Smooth Pursuit Movement: slow, apparently involuntary movements of the
eyes which occur when a person is viewing a moving visual field.
Somatic: refers to the body.
Somatic nervous system: the portion of the nervous system controlling the
skeletal muscles.
Spectral Analysis: a technique for determining the power of the frequencies
present in a physiological record.
Sphygmomanometer: a device used to measure blood pressure, consisting of a
pressure cuff connected to a vertical column containing mercury.
Spirometry: technique for directly measuring the volume of air inspired or
expired during breathing.
Spontaneous ANS Response: a change in ANS activity that occurs in the ab-
sence of any known stimulus.
Spontaneous Electroencephalogram: continually occurring patterns of brain
wave activity, as distinguished from event-related potentials.
Spontaneous Nystagmus (Gaze Nystagmus): oscillatory movements of the eye
related to certain neurological disorders.
Startle Response: occurs to an intense stimulus with a sudden onset; charac-
terized by a reflexive eyeblink, heart rate acceleration, and a rapid habitua-
tion.
Stepwise Cross Correlation: used, among other things, to determine time dif-
ferences in similar EEC activity recorded from two sites.
Stimulus-Response Specificity: principle that specific stimulus situations bring
about certain patterns of responding in most subjects, not just an increase or
a decrease in a unidimensional activation continuum.
Strain-Gauge Transducer: sensing device that changes in electrical resistance
as a function of the degree to which it is stretched; used to measure respi-
ration, for example.
Stretch receptors: sensory receptors sensitive to the stretch or deformation of
tissues.
272 GLOSSARY
Striated Muscle: see Skeletal Muscle.
String Galvanometer: early device used for recording the EKG.
Sympathetic Nervous System (SNS): thoracolumbar division of the autonomic
nervous system; usually activation of this system induces effects opposite
those of the parasympathetic nervous system.
Sympathicotonic: an individual who shows an unusually large response to
drugs which stimulate the sympathetic nervous system.
Symptom Specificity: See Individual Response Stereotypy.
Synapse: the gap between two contiguous neurons along with the axon ter-
minals of the presynaptic neuron and the receptive membrane of the post-
synaptic neuron.
Synaptic cleft: the space or gap between contiguous neurons.
Synaptic Functional Unit: a group of cortical neurons sharing the same pre-
synaptic input.
Systolic Blood Pressure (SBP): the maximal pressure in the vascular system
at the point where the measurement is taken; systolic blood pressure occurs
at the peak of the pressure pulse; units of millimeters of mercury (mmHg).
Tachygastria: an abnormally fast rate of incomplete gastric contractions often
associated with delayed gastric emptying and reports of nausea.
Telemetry: a method for collecting psychophysiological data involving a trans-
mitter (affixed to the subject) and receiver; eliminates need for direct connec-
tions between the subject and the recording equipment.
Thermistor: a device which changes electrical resistance in relation to changes
in its temperature.
Thermocouple: a device which changes voltage in relation to changes in its
temperature.
Thermoresistive Transducer (Thermistor: a device which changes in electrical
resistance in relation to its temperature.
Theta Activity: refers to EEC activity in the 4-8 Hz range.
Tidal volume: The amount of air entering the lungs in a single breath; units
typically in milliliters.
Time Constant: the amount of time required for a signal to return to 63 per-
cent of its voltage.
Time domain techniques: techniques for analyzing data represented across time
(or in the time domain).
Tonic Level: background or basal level of ANS or muscle activity.
Torsional Eye Movements: rotation movements around the line of gaze; smooth
and compensatory.
Transducer: a device that changes one form of energy or activity into another.
Vagotonic: an individual who shows an unusually large response to drugs
which stimulate the parasympathetic nervous system.
GLOSSARY 273
Vasomotor Activity: producing contraction or dilation in the walls of blood
vessels.
Vergence Eye Movements: movement of the eyes in opposite directions so that
objects moving toward or away from the eyes always appear as one object.
Vestibular Nystagmus: oscillatory eye movements elicited by head movement
in which the semicircular canals are stimulated.
Viscera: internal organs of the body.
Visual Evoked Response: a predictable succession of brain wave forms and
distribution observed after the subject is exposed to each of a series of flashes
of light; of maximum amplitude over the occipital areas of the brain.
Wavelet analysis: a technique for determining frequency components of a sig-
nal. One advantage of this technique is its ability to work with short term
signals.
274 GLOSSARY
Index
abdomen, 149 analysis of covariance (ANCOVA),

AC circuitry, 42, 138, 148, 149, 64, 229
201 ANCOVA. See analysis of covariance
acetylcholine, 19, 27, 32, 210 arousal, 52-54, 245
acoustic eye blink, 106-7 arousal-related response, 181-82
actin, 26-27 arrhythmia, 182
action potential arterial tonometry, 197
definition of, 25 arteriole, 22-23
at end of neuron, 30 artifact, 87-89, 116, 119
in individual nerve cells, 29 atrial syncytium, 29
measurement of, 24 attention, 254
in muscle fibers, 27 auditory evoked response, 91
activation theory. See arousal auscultatory measurement, 196
additional heart rate, 183 autonomic balance, 60
adrenal medulla, 19, 22 Autonomic Lability Score, 64
afferent neuron, 15 autonomic nervous system, 17-21
age, 248 and conditioning, 250
air temperature, 148-49 and deep breathing, 143
aliasing, 224, 225 effects on organs and glands, 22-
all-or-none principle, 25, 31 23
alpha activity, 80, 224, 246 and lie detection, 251, 252
alpha rhythm, 80 modes of control, 61-62
amplifiers See also parasympathetic nervous
electroencephalogram, 84 system; sympathetic nervous
electromyography, 113 system
electrooculography, 137-38 Avicenna. See Ibn Sina
event-related potential recording, Ax, Albert, 5, 65
97 axon, 14, 17, 18, 20, 29-30
polygraph, 43-44 axon hillock, 29
amputation, 254 axon terminal, 14
analog signal, 222-24
analog-to-digital converter, 44, band-pass filter, 42
45 baroreceptor, 179
275
basal activity. See tonic activity central nervous system, 14-17, 184,
baseline, 50 209, 250
behavioral neuroscience. See cerebral blood flow, 103
biological psychology change, physiological, 227-30
belladonna, 126 change scores, 227-28
Bereitschaftspotential. See readiness percent, 229-30
potentials residualized, 228-29
Berger, Hans, 9, 79-80 test-retest reliability, 228
Bernard, Claude, 59 chaos analysis. See nonlinear
beta activity, 80 dynamical systems (chaos)
bioelectric potentials, 33-35 analysis
biofeedback, 250-51, 252 Charcot, Jean, 7
biological psychology, 4 classical conditioning, 250
bipolar recording, 82, 84 cognitive neuroscience, 253-55
bladder, 23 cognitive processing, 130, 249
blood flow, 103, 179-80, 199-203 coherence analysis, 90-91, 233-35,
blood pressure, 179, 183, 194-99 237
auscultatory measurement, 196 cold pressor test, 168
automated measurement, 197-98 combined-Doppler ultrasound, 192
indirect measurement, 195-97 comparator theory of habituation, 56
oscillometric measurement, 197, computers, 44-45
198 and brain imaging, 101
blood volume, 199-203 and respiration rate, 153-54
common problems, 201-2 in signal processing, 222-23, 226
recording procedure, 199-201 conditioning, 250
typical recordings, 201 contact lens, 135
bradycardia, 181 contingent negative variation, 93,
brain, 79-103 101
and cognitive neuroscience, 254, control-question technique, 251-52
255 conversion reaction, 7
and eye movement, 131 corneal reflection, 135
lateralization and activation, 249 correlational variables, 249
preganglionic axons exiting, 20 cortex, 100, 101, 102, 103, 145,
See also electroencephalogram 254
brainstem, 145, 179 cortical neuron, 97
breathing amplitude, 142, 154 coupler, 41
breathing rate, 142, 153-55 craniosacral division. See
Burdon-Sanderson, Sir John, 9 parasympathetic nervous system
calcium, 27 Darrow, Chester, 5

calibration, 150 Davis, R. C., 5, 59, 65, 66
capillary electrometer, 8-9 DC circuitry, 42, 138, 201
carbon dioxide, 146 defensive response, 57-58
cardiac muscle, 29 delta activity, 81
cardiac output, 179-80, 192-93 dendrite, 14-15, 16, 30, 97
cardiac responding, 181-85 diaphragm, 146, 147
cardiac-somatic coupling, 183 diastolic blood pressure, 194-95
cardiotachometer, 188, 189, 190 digital signal, 223-24
cardiovascular system, 178-203 dipole modeling, 101
See also electrocardiography; heart directional fractionation, 54, 65, 182
cell body, 14 discrete-time signal, 224
276 INDEX
dual process theory of habituation, impedance and chemical stability,
56 36-37, 113
Duffy, E., 53 paste, 39, 85
polarization, 37
eccrine sweat glands, 209-10, 217, potential or voltage produced by,
222 37
ECG. See electrocardiography and skin conductance, 211, 212
EDA. See electrodermal activity skin preparation for, 39, 112-13
EEC. See electroencephalogram electroencephalogram (EEC), 79-91,
efferent fiber, 17-19 224
efferent neuron, 15, 18 alpha activity, 80, 224, 246
EGG. See electrogastrography analysis and quantification, 90-
Einthoven, Willem, 9, 186 91
EKG. See electrocardiography Berger's early work, 9, 79-80
electricity and brain, 100-101, 249, 255
of heart, 180-81 common problems, 87-89
safety in laboratory, 70-71 electrodes, 82-84
of skin, 7-8 and event-related potentials, 96-
electric shock, 71-72 98
electrocardiography (EKG/ECG), 180- recording procedure, 82-85
81 typical recordings, 86-87
arrhythmias, 182 wavelet analysis, 236
common problems, 189-90 electrogastrography (EGG), 157-74
electrodes, 186-87 amplitude, 163-64, 166, 167
recording procedure, 186-88 cold pressor test, 168
typical recordings, 188 common problems, 170-71
electrodermal activity (EDA), 206- eating and sham feeding, 166-67
19 electrodes, 164-65
amplitude, 214-16, 219 future directions, 174
common problems, 213-14, 218- and gastric motor activity, 162-
19 64
electrodes, 211, 212, 218 and gastric myoelectric activity,
latency, 216, 219 160-62
and lie detection, 252-53 and motion sickness, 169
physiological basis, 209-11 physiological basis, 158-64
recovery time, 216-17 recording procedure, 164-66
skin conductance, 207-17 spectral analysis, 171-74
skin potential, 217-19 typical recordings, 166-70
terminology, 207-9 electromyography (EMG), 106-23
typical recordings, 213, 218 analysis and quantification, 121-
electrode(s), 36-39 23
definition of, 36 common problems, 115-21
electrocardiography, 186-87 electrodes, 109-12, 116-17, 119-
electrodermal activity, 211, 212, 21
218 physiological basis, 108-9
electroencephalogram, 82-84 recording procedure, 109-13
electrogastrography, 164-65 typical recordings, 114
electromyography, 109-12, 116- electronic scanning, 129
17, 119-21 electrooculography (EOG), 135-40
electrooculography, 135-37 analysis and quantification, 139-
and event-related potentials, 9 7 40
INDEX 277
electrooculography (EOG) (Cont.) filters
common problems, 139 electroencephalogram, 84-85
electrodes, 135-37 polygraph, 41-42
procedure, 137 signal processing, 224-25
recording equipment, 137-38 final common path, 17
typical recordings, 138 fluorescent lighting, 224
EMG. See electromyography fMRI. See functional magnetic
emotion, 65, 245, 249 resonance imaging
EOG. See electrooculography Fourier analysis, 90, 230-33
epinephrine, 19 frequency analysis, 90-91
Eppinger, H., 60 frequency domain techniques, 230,
EPSP. See excitatory postsynaptic 231, 234
potential frequency summation, 27
Erasistratos, 6 "frequency window," 85
"Ethical Principles of Psychologists functional magnetic resonance
and Code of Conduct," 73 imaging (fMRI), 103
ethics, 72-76
event-related potentials, 91-100 Galen, 6
analysis and quantification, 98- Galvani, Luigi, 7
100 gamma activity, 80, 255
averaging, 97-98, 99 ganglia, 17-19, 20
common problems, 98 gap junction, 28
evoked responses, 91-93 gastric motor activity, 158-64
physiological basis, 96-97 gastric myoelectric activity, 158-
recording procedure, 97-98 64
slow potentials, 93-96 gastric peristalsis, 162
typical recordings, 98 gastric slow wave, 159-60
evoked responses, 91-93 gastric spike potential, 160
evolving factors, 248 gastrointestinal motility, 15 7-74
excitability, 21, 24-26 See also electrogastrography
excitatory postsynaptic potential gaze nystagmus, 132
(EPSP), 31, 33 gender, 248
experimentation, 6 generator potentials, 30-31
eyes glands, 12
acoustic blink, 106-7 "Goals and Methods of
corneal reflection, 135 Psychophysiology," 5
effects of autonomic nervous ground lead, 70
system, 22 Groves, P. M., 56
electroencephalogram, 88 guilty-knowledge test, 252
electrooculography, 135-40
movement of, 88, 130-35, 138- habituation, 54-56
39 Hales, Steven, 194
photographing of, 127 heart, 22, 29
pupillography, 125-30 anatomy of, 178-79
TV scanning, 135 electrical activity of, 180-81
measures of autonomic effects on,
Fast Fourier Transform (FFT), 90, 185-86
171, 185, 225, 230-33, pumping of, 194
235 See also electrocardiography
Fere, C., 7-8, 247 heart period, 190-93, 231, 232,
FFT. See Fast Fourier Transform 236-37
278 INDEX
heart rate, 186-93 LIV. See law of initial values
and autonomic nervous system, 20 lovesickness, 6-7
and cardiac responding, 181, 183 low-pass filter, 42, 85
and conditioning, 250 lung, 147
recording procedure, 186-88
and respiration, 143-45 magnetic field, 102-3
weighted average, 193 magnetoencephalogram (MEG), 102-
hemoglobin, 103 3, 254
Hering-Breuer reflex, 146 mean arterial pressure, 195
Hess, E. H., 125-26, 127, 130 mean inspiratory flow, 142
Hess, L, 60 mean value, 226
heterostasis, 59 measurement unit, 227
high-pass filter, 42, 85 median, 226
homeostasis, 59-60 medical emergency, 71
hysteria, 7-8 medulla, 145
hysterical anesthesia. See conversion MEG. See magnetoencephalogram
reaction monopolar recording, 82, 84
monosynaptic reflex, 32-33
Ibn Sina, 7 motion sickness, 169
idiosyncratic responding, 66 motor end plate, 32
impedance cardiography, 192-93 motor systems, 15-21
impedance plethysmography, 200 motor unit, 17
impedance pneumography, 148 Mueller (Swiss engineer), 8
individual response stereotypy, 66- multi-unit smooth muscle, 28
67, 245 muscle(s)
informed consent, 74-75 cells, 12, 21, 24-25
inhibitory postsynaptic potential controlling eye movement, 131-32
(IPSP), 31 electromyography, 106-23
inspiratory duty cycle, 142 groups of, 26
instinct, 5-6, 249 smooth, 28-29
intake-rejection hypothesis of cardiac striate, 26-28
responding, 183 myelin sheath, 29-30
inverse problem, 100-101 myosin, 27
IPSP. See inhibitory postsynaptic
potential Necker cube illusion, 236, 239
negative feedback, 59
James, William, 65 nerve, 14
Jung, Carl, 8, 206, 247 cell, 21, 24-25
See also neuron(s)
Korotkoff sounds, 195 nervous system, 12-14
organization of, 14-21
Lacey, Beatrice, 182 neuron(s), 12, 14-15
Lacey, John, 53-54, 64, 66, 182, communication between, 30-33
245 conduction in, 29-30
lateralized readiness potential, 94-96 major components of, 14
law, 74 monosynaptic reflex, 32-33
law of initial values (LIV), 50, 62- motor, 15-21
65, 227-28 sensory, 15
learning, 250 neurotransmitters, 14-15, 19, 31
lie detection, 6, 251-53 nodes of Ranvier, 29-30
limbus-boundary techniques, 135 noise, 224
INDEX 279
nonlinear dynamical systems (chaos) prepulse facilitation, 106
analysis, 238-39 psychobiology. See biological
norepinephrine, 19 psychology
notch filter, 42 psychopathological factors, 248
Nyquist relation, 44, 224 psychophysiological recording(s)
nystagmus movement, 132, 134 applications, 245-56
early instrumentation, 8-9
operant conditioning, 250 equipment used in, 36-45
optokinetic nystagmus, 132, 134 neurons and muscles as sources of,
organizational factors, 248 12-35
orienting response, 56-57 safety and ethics in laboratory, 70-
oscillometric measurement, 197, 198 76
oxygen, 146 spontaneous, tonic, and phasic
activity, 47-51
parasympathetic nervous system, 17, unusual, 71
19-21, 22-23, 60 See also specific recording procedures
and autonomic control, 61-62 Psychophysiology, 5, 246
and cardiac responding, 183-85 psychophysiology
passions, 5 basic principles of, 52-67
Pavlov, I. P., 57 and biological psychology, 4
peripheral nervous system, 14, 184 of brain, 79-103
personality factors, 247 of cardiovascular system, 178-203
phasic activity, 50-51, 221-22 definition of, 3-4
photoconduction cell, 40 future of, 255-56
photoelectric plethysmography, 199- of gastrointestinal system, 15 7-74
200 history of, 5-7
physiopathological factors, 248 of muscles, 106-23
piezoelectric devices, 150 of respiratory system, 142-55
Plato, 5-6 of skin, 206-19
plethysmography, 199-201, 226 studies, 246-55
Plutarch, 6 "Psychophysiology, Yesterday,
polarization, 21, 37 Today, and Tomorrow," 5
Polt, James, 126 pule pressure, 194-95
polygraph, 41-44 pulse volume, 199, 201
amplification, 43-44 pupillography, 125-30
coupler, 41 analysis and quantification, 130
display, 44 common problems, 129-30
filtering, 41-42 physiological basis, 127
pons, 145 recording procedure, 127, 129
positron emission tomography, 103
postganglionic axon, 20 QRS complex, 187-89
postganglionic fiber, 18-19
postganglionic neuron, 18, 19 race, 248
potassium, 21 readiness potentials, 94-96, 101
power spectrum, 231-32 reading, 130, 132
preamplifier, 43, 138 receptor potentials, 30-31
pre-ejection period, 185-86 reduction, of physiological data, 226
preganglionic axon, 20 REM (rapid eye movement) sleep,
preganglionic fiber, 19 131
preganglionic neuron, 18, 19 research ethical standards, 73-76
280 INDEX
respiratory inductive smooth pursuit movements, 131,
plethysmography, 149-50 132, 133
respiratory sinus arrhythmia, 143, Society for Psychophysiological
145, 185 Research, 5
respiratory system, 142-55 sodium, 25
physiological basis, 145-47 Sokolov, E. N., 56
potential problems in recordings, somatic responses, 47
152-53 somatic system, 16-17
recording procedures, 147-52 spectral analysis, 171-74
respiration amplitude, 142, 154 spectral density plot, 231
respiration power spectrum, 232, sphygmomanometer, 195-96
236-37 spinal cord, 16, 18, 20
respiration rate, 142, 153 spirometry, 147-48, 154
respiratory events, 154-55 spontaneous electroencephalogram.
response variables, 246 See electroencephalogram
responsibility, 73-74 spontaneous responses, 47-49
resting potential, 21, 24-25, 28-29 SQUID (superconducting quantum
ribcage, 146, 147, 150 interference device), 102
stability, 59
saccadic eye movement, 132, 133 standards, 74
safety, 70-72 startle response, 58, 106-7
saltatory conduction, 30 state variables, 247
sampling rate, 44, 222-25 Stern, John, 3
schizophrenia, 131 stimulus-response specificity, 6, 54,
self-control, 250 65-66
self-inductance, 149 stimulus/situational variables, 246-
sensitization, 56 47
sensory systems, 15 strain gauge, 150-52
signal-averaging procedures, 97-98 strain gauge plethysmography, 200-
signal processing, 221-39 201
signal-to-noise ratio, 43 strain gauge transducer, 40, 150
sine waves, 231, 234 striate muscle, 26-28
size principle, 17 string galvanometer, 9, 186
skeletal muscle. See striate muscle subject variables, 247-49
skin superconducting quantum
electrical properties of, 7-8 interference device. See SQUID
electrodermal activity, 206-19 sweat glands, 209-11
preparation for electrodes, 39, 112- sympathetic nervous system, 17-19,
13 22-23, 60
resistance, 7-8 and autonomic control, 61-62
skin conductance, 207-17 and blood flow, 179
conversion from resistance and cardiac responding, 183-85
readings, 214 sympathicotonics, 60
recording procedure, 211-12 synapse, 30-32, 97
responses, 222 synaptic cleft, 31
See also electrodermal activity syncytia, 29
skin potential, 217-19 systolic blood pressure, 194-95
sleep, 87, 130-31, 247
slow potentials, 93-96, 101 tachycardia, 181
smooth muscle, 28-29 Tallon-Baudry, Catherine, 80
INDEX 281
tetanus, 27-28 unitary smooth muscle, 28
thalamus, 246
thermistor, 41, 148 vagotonics, 60
thermocouple, 148 "vanilla baseline," 50
theta activity, 81 vascular unloading method, 197-
Thompson, R. F., 56 98
thoracolumbar system. See vasoconstriction, 179, 180
sympathetic nervous system vasodilation, 179, 180
thorax, 146, 148, 150 venous occlusion plethysmography,
threshold, depolarization, 25 201
time domain techniques, 230, 231, ventricular syncytium, 29
234 Veraguth (neurologist), 8
tonic activity, 49-50, 221-22, 226- vergence eye movement, 132, 133
27 vestibular nystagmus, 132, 134
tonometric measurement, 197 Vigouroux (scientist), 7
torsional eye movement, 132, 133 visual evoked response, 91
tract, neuron, 14
trait factors, 247 wavelet analysis, 235-36, 238-39
transducer, 40-41, 150 Wenger, M. A., 60
"transfert," 7 Wheatstone bridge circuit, 151
twins, 247 "white-coat hypertension," 196
282 INDEX

Robert Morris Stern, William J. Ray, Karen S. Quigley - Psychophysiological Recording-Oxford University Press (2001)

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PSYCHOPHYSIOLOGICAL RECORDING

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Copyright © 2001 by Oxford University Press, Inc.

Library of Congress Cataloging-in-Publication Data

2. Neurons and Muscles: The Sources of

3. Equipment Used in Psychophysiological Recording 36

5. Some Basic Principles of Psychophysiology 52

7. Brain: Electroencephalography and Imaging 79

8. Muscles: Electromyography 106

9. Eyes: Pupillography and Electrooculography 125

10. Respiratory System 142

11. Gastrointestinal Motility: Electrogastrography 157

12. Cardiovascular System: Heart Rate; Cardiac Output;

13. Skin: Electrodermal Activity 206

14. Signal Processing 221

PART III. APPLICATIONS

15. Applications of Psychophysiological Recording 245

Psychophysiology is relatively new as a separate discipline; in the mid-

Short History and Long Past

The history of psychophysiology as a separate discipline is quite brief. The

The Past of Psychophysiology

Understanding the Electrical Properties

References to the specific material covered in the following chapters are

The bodily responses that are the subject of psychophysiological study

Organization of the Nervous System

Somatic System. The somatic nervous system is composed of efferent neu-

Autonomic System. The autonomic nervous system (ANS) consists of two

Function of Nerve and Muscle Cells

Excitability: Resting Potential and

Eyes: Iris Control of light to the

Heart: S-A node Rate control Slows rate Increases rate

Bladder: muscle wall Muscle tone Contraction Relaxation

(Na + ), and an energy-requiring Na + -K + pump in the cell's membrane

on the rate at which action potentials are generated. Whereas striate

Communication between Neurons. Until now, we have mostly described

The bioelectric events—both action potentials and the graded, synaptic

In chapter 2 we discussed the sources of the bioelectric potentials that

Electrodes and Transducers

We will describe only those electrodes designed to be placed directly on

electrode (figure 3.1) is attached to the subject by an adhesive collar. The

Polygraphs record a physiological signal in a continuous analog fashion

GENERAL ELEMENTS OF PSYCHOPHYSIOLOGY

Brown, C. C. (1967). Methods in psychophysiology. Baltimore, MD: Williams

... somatic responses abound. One has but to observe them on

Somatic responses such as heart rate and muscle potentials do indeed

What does a spontaneous physiological response to an unknown stimulus

Tonic activity is often referred to as the background level or resting level

Phasic activity is a discrete response to a specific stimulus—an evoked

The basic principles of psychophysiological recording are generalizations

Arousal and Habituation

Figure 5.1. Duffy's hypothesized inverted U-shaped function relating level of

SOME BASIC P R I N C I P L E S OF PSYCHOPHYSIOLOGY 55

Orienting, Defensive, and

For more than seven decades, a considerable amount of research has

Some of the major components of the orienting response include (1)

SOME BASIC PRINCIPLES OF PSYCHOPHYSIOLOGY 59

60 GENERAL ELEMENTS OF PSYCHOPHYSIOLOGY

Figure 5.3. Possible modes of autonomic nervous system control. Reproduced

Noninvasive estimates of sympathetic (e.g., cardiac pre-ejection period)

Law of Initial Values