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INTRODUCTION TO SECTION I THE NEUROBIOLOGY
OF THINKING
Anyone who can read these words, including you,
experiences conscious awareness. What explains
this? Is the conscious experience of other people “the
same” as my own? Is our conscious experience
qualitatively different from what may or may not be
experienced by nonhuman animals? These and similar
questions have fascinated some of humankind’s most
celebrated thinkers for as long as humans have been
leaving records of their cogitation. They are among the
most profound that a human being can ask. Not
surprising, then, is the fact that questions that relate
either directly or indirectly to the phenomenon
consciousness and the related construct of cognition
(i.e., thinking) have been taken up by many different
scholarly disciplines, including philosophy,
evolutionary biology, anthropology, economics,
linguistics, computer science, and psychology. What
distinguishes cognitive neuroscience, the focus of this
book, from these and other disciplines, is its
grounding in the methods and traditions of
neuroscience, and the primacy that it places on
understanding the biological bases of mental
phenomena. Now it bears noting that cognitive
neuroscience doesn’t concern itself only with
consciousness and conscious awareness. Indeed, I
feel confident asserting that the vast majority of
articles that have been published in, say, the Journal
of Cognitive Neuroscience (one of many scientific
journals publishing peer-reviewed reports of research
in the field) don’t even explicitly address the idea of
consciousness, much less use the word.
Nonetheless, it is also true that it can be difficult to
entertain detailed thoughts about human behavior
and the principles that govern it without straying into
ideas such as those that opened this paragraph. This
clear relationship to profound philosophical questions
is one of the qualities that differentiates cognitive
neuroscience from other physical and biological
sciences, including many other domains of
neuroscience. Like other physical and biological
scientists, cognitive neuroscientists design and carry
out rigorously controlled experiments that yield
objective, measurable data, and seek to relate these
data to mechanistic models of how a natural system
works. And, as suggested above, the system studied
by cognitive neuroscientists is one that is also of
direct interest to scholars who study it from a very
different perspective (e.g., from philosophy, from
anthropology, and from cognitive psychology). This
overlap turns out to be both a blessing and a curse.
It’s a blessing in that the cognitive neuroscientist can
draw on ideas and observations from a vast array of
rich intellectual traditions. And because they are
studying questions that are fundamental to the human
condition, almost any thinking person that a cognitive
neuroscientist encounters is likely to be interested in
what they do for a living. (Much easier, in the humble
opinion of the author, to be a cognitive neuroscientist
attending a social function full of strangers than, say,
an economist, or a cell biologist, or a particle
physicist.) The curse is found on the flip side of this
coin. Because scholars in different fields are often
interested in the same object of study (whether it be
visual perception, or aesthetics, or antisocial
behavior), cognitive neuroscience research can often
be characterized by people working in other fields as
asking the wrong question, or asking it in the wrong
way, or of generating findings that are fundamentally
irrelevant for understanding the question at hand.
Being aware of this sociological context surrounding
cognitive neuroscience is INTRODUCTION TO
SECTION I 3 just one piece of knowledge that can be
helpful in evaluating the implications and importance
of any particular set of facts and ideas that will be
described in this book. COGNITIVE NEUROSCIENCE?
OR “HUMAN NEUROSCIENCE”? OR
“NEUROSCIENCE-WITHDIRECT-IMPLICATIONSFOR-
UNDERSTANDINGHUMAN-BEHAVIOR”? How were
the topics to be covered in this book selected? More
importantly, when teaching cognitive neuroscience,
where does one draw the line that defines the
boundaries of the discipline? This is a difficult
question that doesn’t have a definitive answer. It’s
really just an accident of history that the confluence of
neuroscientific methods and studies of human
behavior happened first, or, at least, influentially, with
domains of behavior and function that are studied by
cognitive psychology (e.g., visual perception, language
function, memory), as opposed to, for example, social
behavior, personality, emotion, or psychopathology.
As a result, one sees the label “cognitive
neuroscience” appearing earlier in the literature than,
for example, affective neuroscience or, certainly,
social cognitive neuroscience. As a consequence, the
term “cognitive neuroscience” has come to be used in
many contexts, and not always with precisely the
same meaning. In one context, “cognitive
neuroscience” can be used to refer to the tools and
methods used to study the neural bases of human
behavior (e.g., brain scans, recording electrical
potentials at the scalp, brain stimulation). This might
seem like a misnomer, however, if the behavior being
studied doesn’t directly relate to cognition (e.g., the
study of sleep or of the maturation of various brain
systems during adolescence). The label “cognitive
neuroscience” can also seem like a bad fit for
research that spans multiple traditional categories of
brain and/or behavior. To pick one real-world
example, let’s consider the study of how neural
systems measured with brain scans relate to
performance on an economic decision-making task by
high trait-anxiety vs. low traitanxiety prisoners (all
classified as “psychopaths”), and the comparison of
these data with data from neurological patients with
frank damage to the ventromedial prefrontal cortex.
Should this be classified as cognitive neuroscience
research? On one level, I hope that the answer is
“yes,” because we will be considering this research in
Chapter 17! However, to do so is to not explicitly
acknowledge the equally important contributions to
this research from clinical psychology, from affective
neuroscience, from neuroeconomics, and from
neuropsychology. In view of the above, to capture the
interdisciplinary breadth of much of what we’ll be
considering in this book, might it perhaps have been
better to entitle it “Essentials of Human
Neuroscience”? Here the answer is an unequivocal
“no,” because to do so would be to exclude the fact
that understanding the neural bases of almost all
domains of human behavior requires a thorough
knowledge of analogous neural functioning in
nonhuman animals. This is because, as we shall see
in almost every chapter of this book, technical and
ethical limitations of what we can measure in humans
require us to draw heavily on the results of research
performed with nonhuman animals. Hence, the idea,
implied in the title of this section, that the most
precise title for this book might be “Essentials of
Neuroscience with Direct Implications for
Understanding Human Behavior.” Now I’m no expert
in academic publishing, but I’m fairly confident that,
were I to have actually proposed this title, my editor
would have rejected it. If nothing else, there probably
aren’t many universities where one can find a course
that’s called “Introduction to Neuroscience with
Direct Implications for Understanding Human
Behavior.” Nonetheless, this is probably as good a
summary, in 10 words or less, of what this book hopes
to cover. And so, with these considerations in mind,
we’ll stick with the label “cognitive neuroscience.” It’s
not perfect, but if one is comfortable with a
reasonably broad definition of cognition as thinking,
behaving, and the factors on which these depend,
then this label will serve us reasonably well. CHAPTER
1 KEY THEMES ● Although the phenomenon of
consciousness and the related construct of cognition
(i.e., thinking) are the focus of many different scholarly
disciplines, what distinguishes cognitive
neuroscience is its grounding in the methods and
traditions of neuroscience, and the primacy that it
places on understanding the neurobiological bases of
mental phenomena. ● There are two levels at which
the term “cognitive neuroscience” is used: broadly, it
has come to refer to the neuroscientific study of most
domains of human behavior; narrowly, it refers to the
study of neural bases of thinking – what influences it,
what it consists of, and how it is controlled. ● The
roots of cognitive neuroscience can be traced back to
a nineteenth-century debate over two ways of thinking
about brain function that both remain relevant today:
localization of function vs. mass action. ● Mid-to-late
nineteenth-century research, and the vigorous debate
that accompanied it, led to models of localization of
three functions: motor control (localized to posterior
frontal lobes); vision (localized to occipital lobes); and
speech production (localized to the left posterior
inferior frontal gyrus). ● Motor control research
introduced the principle of topographic
representation, that is, adjacent parts of the body can
be represented on adjacent parts of the cerebral
cortex. ● Studying an aspect of cognition requires
careful thought about the validity of the function to be
studied; and not all aspects of human behavior can be
studied with the same sets of assumptions, or even
with the same methods. ● The discipline of cognitive
neuroscience could not exist without discoveries
yielded by research with nonhuman animals. ● At the
dawn of the twentieth century, scientists were
studying the brain and behavior from three related, but
distinct, perspectives that would eventually give rise
to cognitive neuroscience as we know it today:
systems neuroscience, behavioral
neurology/neuropsychology, and experimental
psychology. INTRODUCTION AND HISTORY
INTRODUCTION AND HISTORY 5 CONTENTS KEY
THEMES A BRIEF (AND SELECTIVE) HISTORY
Construct validity in models of cognition Localization
of function vs. mass action The first scientifically
rigorous demonstrations of localization of function
The localization of motor functions The localization of
visual perception The localization of speech WHAT IS
A BRAIN AND WHAT DOES IT DO? LOOKING AHEAD
TO THE DEVELOPMENT OF COGNITIVE
NEUROSCIENCE END-OF-CHAPTER QUESTIONS
REFERENCES OTHER SOURCES USED FURTHER
READING 6 SECTION I: THE NEUROBIOLOGY OF
THINKING A BRIEF (AND SELECTIVE) HISTORY
Although the term “cognitive neuroscience” as a
moniker for a scientific discipline has only been with
us for a few decades, the field has roots that extend
back thousands of years. Ancient Egyptians, Greeks,
and Romans all had ideas about the corporeal bases
of human thoughts and emotions, although many of
these did not specify a role for the brain. In preparing
the bodies of deceased nobles for the afterlife, for
example, ancient Egyptians removed and discarded
the brain as an early step in the mummification
process. The internal organs that were deemed to be
important were preserved in urns that were entombed
along with the body. In most ancient civilizations for
which there are records, up through and including
Roman civilization, the heart was believed to be the
organ of thought. By the time we get to Enlightenment–
era Europe, however, the central importance of
“neuro” for cognition was widely accepted. One highly
influential (and, more recently, ridiculed) example was
that of German anatomists Franz Josef Gall (1758–
1828) and Johann Gaspar Spurzheim (1776–1832),
who developed a highly detailed scheme, known as
phrenology, for how they thought that the shape of
different parts of the skull related to one’s personality
and mental capacities. The underlying premise was
that the relative bigness or smallness of various parts
of the brain would produce convexities or concavities
in the overlying skull. A skilled phrenologist, then,
could learn something about an individual by
palpating that person’s skull. A bulge in the eye socket
(inferred from “bulgy eyes”) would mean a
predilection toward language, whereas an indentation
near the left ear would correspond to a relative
absence of the trait of “destructiveness” (Figure 1.1).
One can see how such a scheme, if it had any validity,
would have obvious utility for diagnosing maladies of
the brain, as well as for assessing personality and
aptitude. (Indeed, for a period during the 1800s it was
[mis] used in this way quite extensively, particularly in
England and in the United States.) Construct validity in
models of cognition For at least the past 100 years,
the psychology and neuroscience communities have
viewed virtually all tenets of the phrenological
enterprise as being scientifically invalid. For starters,
the very “functions” that phrenologists assigned to
different parts of the scalp were derived from Gall’s
intuition rather than from a principled theoretical
framework. Let’s consider, for example,
conscientiousness. Now it is true, of course, that an
organism without a brain cannot exhibit
conscientiousness, and, therefore, the phenomenon
could not exist without a brain. However, might it not
be the case that “conscientiousness” is just a label
that we, as denizens of highly organized societies,
have given to a certain collection of attributes
characteristic of some peoples’ behavior and
personality? This can be illustrated from two
perspectives. The first is a caveat about inferring the
existence of a discrete neural correspondence to
every describable aspect of behavior. For example,
when a student sends me a thank-you note for having
written a letter of recommendation for her, I will
consider her to have displayed conscientiousness.
But could it not be that this reflects the fact that she
was conditioned during her upbringing to seek positive
reinforcement from her parents (“You’re such a good
girl for writing those thank-you notes!”), and that her
note to me is “merely” the product of an association
that she has formed between writing a thank-you note
and this reinforcement? Were this the case, it
wouldn’t make sense to think of conscientiousness as
a discrete mental faculty. (And if there’s no such
faculty, then there’s no “thing” to localize to a part of
the brain.) The second perspective is that, as it turns
out, there is a branch of psychology in which
conscientiousness is a valid construct, but it is as one
of the so-called “big-5” personality traits. But because
each of these five traits (openness-toexperiences,
conscientiousness, extraversion, agreeableness, and
neuroticism) is a statistical composite derived from
multiple measurements (such as responses to many-
morethan-five questions on personality assessment
questionnaire), none of them can be construed as a
unitary entity that might be localized to one brain
system. And so, from this perspective, because
conscientiousness isn’t any one thing, it doesn’t seem
reasonable to expect it to localize to one part of the
brain.

The exercise of thinking through the phrenological

treatment of conscientiousness highlights two
concepts that are highly relevant to contemporary
cognitive neuroscience. The first is fundamentally
“cognitive”: a model of the neural instantiation of a
cognitive function depends critically on the validity of
the function that it seeks to explain. The question of
construct validity will be important for every domain of
behavior that we consider in this book. For many, the
formal models of the construct under study will come
from one of the “different scholarly disciplines”
invoked in the Introduction to Section I.
INTRODUCTION AND HISTORY 7 A second concept
invoked by phrenology is fundamentally
“neuroscience,” and its legacy from the Age of Reason
is more complicated. On the one hand, we now know
that subtle, idiosyncratic variations in gross shape
from one brain to another have little, if anything, to do
with the “kind of person” that one is. We also
recognize that the assignments of function that Gall
gave to various parts of the brain were not based on
rigorous science and turned out to be altogether
wrong. A third point is that the very selection and
definition of functions that phrenologists mapped
onto the brain lacked systematicity and rigor. There
was, however, at the core of the phrenological
enterprise, a powerful idea that has continued to
animate many debates about brain function up to the
present time – the idea of localization of function.
Localization of function vs. mass action The principle
of localization of function refers to the idea that
different aspects of brain function, such as visual
perception vs. the control of our emotions vs. our
talents as musicians, are governed by, and therefore
localizable to, different “centers” in the brain. An
analogy might be that different functions of the body –
extracting oxygen from 2 2 2 3 3 3 4 4 4 6 6 10 8 9 9 9 19
18 20 7 1 17 10 10 11 11 11 12 12 12 15 15 14 14 13 13
18 18 21 21 21 20 19 19 34 35 35 35 30 31 31 31 24 24
25 24 25 25 26 26 26 23 23 28 28 28 29 29 29 27 27 27
22 32 32 32 33 33 20 15 16 16 16 6 5 5 5 * FIGURE 1.1 A
phrenological map of the anatomical organization of
mental faculties. They are organized into two primary
categories (bold), with subcategories (underlined)
under each. The first primary category is Affective
Faculties: Propensities:* – Alimentiveness; 1 –
Destructiveness; 2 – Amativeness; 3 –
Philoprogenitiveness; 4 – Adhesiveness; 5 –
Inhabitiveness; 6 – Combativeness; 7 – Secretiveness;
8 – Acquisitiveness; 9 – Constructiveness;
Sentiments: 10 – Cautiousness; 11 – Approbativeness;
12 – Self-esteem; 13 – Benevolence; 14 – Reverence;
15 – Firmness; 16 – Conscientiousness; 17 – Hope; 18
– Marvelousness; 19 – Ideality; 20 – Mirthfulness; 21 –
Imitation. The second primary category is Intellectual
Faculties: Perceptive: 22 – Individuality; 23 –
Configuration; 24 – Size; 25 – Weight and resistance;
26 – Coloring; 27 – Locality; 28 – Order; 29 –
Calculation; 30 – Eventuality; 31 – Time; 32 – Tune; 33
– Language; Reflective: 34 – Comparison; 35 –
Causality. Source: Spurzheim, Johann C. 1834.
Phrenology or the Doctrine of the Mental
Phenomenon, 3rd ed. Boston: Marsh, Capen, and
Lyon. Public Domain. 8 SECTION I: THE
NEUROBIOLOGY OF THINKING blood vs. pumping
blood vs. filtering blood – are each accomplished by
different organs (i.e., the lungs, the heart, and the
kidneys) that are located in different parts of the body.
This notion can be contrasted with an alternative idea,
mass action, according to which a particular function
can’t necessarily be localized to a specific area of the
brain, and, conversely, any given area of the brain
can’t be thought of as a “center” that is specialized for
any one function. To stick with our analogy to familiar
parts of the body below the neck, we can illustrate the
principle of mass action by zeroing in on the kidney.
The overall function of the kidney – filtering blood – is
carried out in the same way by the top portion and the
middle portion and the bottom portion. To understand
how the kidney does its job, one could study in detail
the inner workings of only the top, or only the middle,
or only the bottom of this organ, and one would learn
the same thing from each. In effect, then, different
zones of the kidney are “interchangeable” with
respect to understanding their functions. Now project
yourself back in time a few centuries to a time when
what I’ve just written hasn’t yet been discovered. It is
an era when biomedical research techniques are
limited, and the best tool that you have for studying
the function of an organ is to damage a portion of it
and then observe the consequent impact of this
damage on its function. Your kidney research would
indicate that damaging comparable-sized regions of
the upper vs. middle vs. lower kidney has the same
effect in all cases: an overall decline in the efficacy of
blood filtration. Thus, you would have discovered that
a principle of mass action applies to the kidney: a
larger lesion results in a larger diminishment of the
rate of blood filtration; smaller lesions result in a
smaller diminishment of the rate of blood filtration;
and, critically, the effects of damage to different parts
of the kidney are the same. Now, let’s return to the
functions of the brain. In the decades following the
introduction of phrenology, and in some instances in
reaction to it, scientists and physicians began
pursuing the idea of localization of function in the
brain with methods reflecting the maturation of the
scientific method that was occurring in many
branches of science, from biology to chemistry to
physics. At this general level, this entailed the a priori
articulation of falsifiable hypotheses (i.e., devising
experiments that, if successful, can rule out
previously plausible ideas and stating how various
possible outcomes would be interpreted prior to
performing the experiment) and the design of
controlled laboratory experiments that could be
replicated in other laboratories. An important advance
for studies of the brain, in particular, was the careful
analysis of the behavioral consequences resulting
from damage to a particular brain structure. This
method has come to be known as neuropsychology.
Armed with this approach, nineteenth-century
scientists began to disprove many specific
localizationist claims from phrenology. Perhaps most
influential were the studies of French scientist Pierre
Flourens (1794–1867). A tireless critic of phrenology,
Flourens did much of his experimental work with
pigeons and dogs. This research was influential on two
levels. First, particular experiments of Flourens
disproved specific phrenological claims, such as his
demonstration that damage to the cerebellum
disrupted locomotor coordination, but had no effect
on amativeness (i.e., predilection for sexual arousal),
as Gall’s model would have predicted. Secondly, and
at a broader level, Flourens’ studies of the brain
largely failed to find evidence for localization of many
functions. Thus, although damage to the brain
invariably produced marked disruption of behaviors
associated with judging, remembering, and
perceiving, these impairments seemed to occur
regardless of what part of the brain had been
damaged. By inference, such results seemed to
indicate that all regions of the brain contributed
equally to these behaviors. (Note that the same was
not true of Flourens’ studies of the brainstem, to
which, for example, the cerebellum belongs.) (Figure
1.2). Another important concept to come out of the
work of Flourens was derived from the fact that, over
time, animals with experimental damage to a part of
cortex often recovered to presurgical levels of
functioning. Because this occurred without evident
repair of the damaged tissue itself, it was assumed
that intact areas of the brain had taken over this
function. This gave rise to the concept of
equipotentiality, the idea that any given piece of
cortical tissue had the potential to support any brain
function. Roughly 50 years prior to the writing of this
textbook, and 130 years after the heyday of
phrenology, neuroscientists Charles Gross and
Lawrence Weiskrantz wrote that “The ‘heroic age of
our field’ was opened by Gall (1835) . . . [who]
stimulated the search for centers and gave the mass
action-localization pendulum its first major swing”
(Gross and Weiskrantz, 1964). Implied in this quote
was that understanding the localization–mass action
dichotomy would provide insight into understanding
key contemporary problems in neuroscience. Indeed,
this theme will prove to be useful for understanding
many of the concepts and controversies that are
prominent in contemporary cognitive neuroscience.
INTRODUCTION AND HISTORY 9 A Frontal lobe
Central sulcus Parietal lobe Occipital lobe
Cerebellum Sylvian fissure Temporal lobe Corpus
callosum Frontal lobe Limbic lobe B Caudate nucleus
Fornix Temporal lobe Brainstem Cerebellum Occipital
lobe Thalamus Limbic lobe Parietal lobe Central
sulcus FIGURE 1.2 The human brain (see Chapter 2 for
labels and terminology). A. Lateral view. B. Medial
view. Source: Dr. Colin Chumbley/Science Source.
The first scientifically rigorous demonstrations of
localization of function Although the concepts
advocated by Gall, on the one hand, and Flourens, on
the other hand, still resonate today, the same cannot
be said for most of the “facts” that arose from their
work. Rather, it was in the mid-to-late 1800s, during
what can be seen as the first return of the pendulum
back toward localization, that we see the emergence
of principles of brain function that, at least to a first
order of approximation, have held up through to the
present day. These involved the functions of motor
control, vision, and language. The localization of
motor functions Beginning in the 1860s, British
neurologist John Hughlings Jackson (1835–1911)
described the systematic trajectory of certain focal
seizures that appear to start in the fingers and spread
along the arm toward the trunk, sometimes ending
with a loss of consciousness. From this distinctive
pattern, which has since come to be known as the 10
SECTION I: THE NEUROBIOLOGY OF THINKING
Jacksonian march (it’s as though the seizure is
“marching” along the body), Jackson proposed that
the abnormal brain activity presumed to be the cause
of this progression of abnormal muscle contractions
begins in a part of the brain that controls the fingers,
then moves continuously along the surface of the
brain, progressively affecting areas that control the
palm of the hand, the wrist, the forearm, and so forth.
There were two important implications of Jackson’s
theory. The first was quite simply the proposal that the
capacity for movement of the body (i.e., motor
control) is a function that is localized within the brain.
The second was what has come to be understood as a
fundamental principle underlying the organization of
function within many portions of the brain, which is
that their organization can mirror the organization of
the body (or, as we shall see, of a particular part of the
body). Specifically, in this case, the proposal was that
the area of the brain that controls the muscles of the
fingers is adjacent to the area of the brain that
controls the muscles of the palm, which is adjacent to
the area of the brain that controls the muscles of the
wrist, and so forth. Thus, the functions of what came
to be known as the motor cortex are physically laid out
on the surface of the brain in a kind of map of the body
(that is, in a somatotopy). In this way, the idea of a
lawful, topographic organization of function was
introduced in the void left by the (by-now-largely-
discredited) arbitrary, willy-nilly scheme of the
phrenologists. (The principle and characteristics of
somatotopy will be considered in detail in Chapter 5
and Chapter 8.) Although the ideas that Jackson
proposed were based on careful observation of
patients, an idea such as the somatotopic
organization of motor cortex couldn’t be definitively
evaluated without either direct observation or,
preferably, manipulation of the brain itself. The ability
to undertake such definitive empirical investigation
became possible because Age-of-Enlightenment
advances in thinking about how science should be
conducted, such as the importance of the scientific
method, were being paralleled by technical advances
that afforded improved experimental methods. Of
particular importance was the development of
methods for performing aseptic (i.e., sterile) surgery.
These enabled experimenters to keep animals alive for
weeks or longer after performing the craniotomy that
was necessary to create a lesion or to manipulate the
functioning of the brain. Prior to this, infection would
often limit postsurgical survival to a matter of hours or
days. This technical advance had several important
consequences, one of which was that it opened the
way for direct stimulation of, and subsequently
recording from, the brain of an intact animal
(techniques that fall under the category of
neurophysiology). Thus, it was that German
physicians Gustav Fritsch (1838–1927) and Eduard
Hitzig (1838–1907) reported that electrical stimulation
of anterior portions of the cerebral cortex of a dog, in
the frontal lobe, produced movements on the
opposite side of the body (Fritsch and Hitzig, 1870).
Noteworthy were two facts. First, comparable
stimulation of a more posterior brain region, the
parietal lobe, did not produce body movements
(Figure 1.3). Based on this demonstration of the
anatomical specificity of their effect, Fritsch and
Hitzig explicitly challenged the idea of equipotentiality
that had been advocated by Flourens. Second, the
part of the body affected by electrical stimulation
(e.g., neck, forelimb, hind limb) varied systematically
with the positioning of the stimulating electrode. This
observation very directly supported Jackson’s idea of
a somatotopic organization of motor functions in the
brain.
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