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 Behavioral
Neuroscience

GEORGE SPILICH
Washington College

Spilich1e_FM.indd 1 12/19/22 7:46 PM

 VICE PRESIDENT Amanda Miller
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Library of Congress Cataloging-in-Publication Data
Names: Spilich, George, author.
Title: Behavioral neuroscience / George Spilich, Washington College.
Description: First edition. | [Hoboken, NJ] : Wiley, [2023] | Includes index.
Identifiers: LCCN 2022053829 (print) | LCCN 2022053830 (ebook) | ISBN 9781118547380 (print) |
ISBN 9781119889625 (adobe pdf) | ISBN 9781119505662 (epub)
Subjects: LCSH: Psychobiology—Textbooks. | Neuropsychology—Textbooks.
Classification: LCC QP360 .S696 2023 (print) | LCC QP360 (ebook) | DDC
612.8—dc23/eng/20221215
LC record available at [Link]
LC ebook record available at [Link]
Printed in the United States of America.
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Spilich1e_FM.indd 2 12/19/22 7:46 PM

 About the Author

GE ORG E S P ILICH is a behavioral neuroscientist who earned degrees from the University
of Wisconsin, the University of Texas at El Paso, and the University of Pittsburgh, where he
was at the Learning Research and Development Center. He is currently an emeritus faculty at
Washington College, the first college founded after the American Revolution, where he was
the department chair for 20 years, was the first John Toll Professor, and won the Lindback
Foundation Teaching Award. In academic year 1988–1989, he replaced two-time Nobel lau-
reate Linus Pauling as the Fulbright Researcher for Yugoslavia and spent the year as visiting
professor in the departments of neurology and nuclear medicine at university hospitals. He
was the inaugural curator of the Cromwell Center for Teaching and Learning at Washing-
ton College. His research interests explore the processes that underlie encoding, storage, and
retrieval of information in an effort to understand how a biological system creates and stores
expertise and then how disease and damage disrupt that mechanism.

The human brain has 100 billion neurons, each neuron is connected to 10 thousand
other neurons. Sitting on your shoulders is the most complicated object in the
known Universe.

—MICHIO KAKU

It is essential to understand our brains in some detail if we are to assess correctly

our place in this vast and complicated universe we see all around us.

—FRANCIS CRICK, NOBEL LAUREATE

I’m just looking for clues at the scene of the crime.

—JOSEPH FIDLER WALSH

iii

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 Preface

Understanding human behavior from a biological perspective is the goal of behavioral neu-
roscience. The field incorporates biology, genetics, developmental biology, gerontology, psy-
chopharmacology, cognitive psychology, and even sociology to create a multidisciplinary
approach to understanding behavior.
Behavioral neuroscientists consider questions such as, How does our evolutionary past
influence our behavior today? To what extent is our behavior controlled by our genes? How
do our senses convert raw input into what we call reality? How do we regulate our internal
processes to stay alive? Why are some substances so addictive? What does it mean to have a
sex or gender identity? Why do I sleep (or not!), and why do I dream? Why can’t I remember
what I learned last year? Why do some people become depressed, anxious, or even have a total
break from reality? Is beauty only a construction of my mind? These and thousands of other
questions are the purview of behavioral neuroscience, one of the most exciting of human
endeavors.

Features Unique to Behavioral

Neuroscience
Meets Students Where They Are: Some students will take this course in a traditional
face-to-face lecture/discussion format; some will enroll in an online class, and others will
take a hybrid course. Students choosing to enroll in behavioral neuroscience often come
from two groups: those preparing for a career in medicine or science, and those students
who are fulfilling a general education science requirement and choose behavioral neurosci-
ence to learn more about themselves. Behavioral Neuroscience is written to reach all those
audiences.
Making Science Approachable and Inviting: Students following a course of study
leading to a career as a physician, medical/physician’s assistant, nurse, or substance abuse/
addictions counselor; students preparing for graduate work in behavioral neuroscience; and
those preparing to take the MCAT will benefit from the focus on science. However, all stu-
dents, including those following a different career path, will benefit from an accessible writing
style and frequent examples of how behavioral neuroscience speaks to the issues of everyday
life.
Encourages Active Learning through Effective Use of Media: Behavioral Neu-
roscience engages students with a wealth of multimedia created or curated by the author and
puts that media to use often at the service of conveying traditionally difficult-to-master con-
cepts. Every chapter contains multiple activities that engage students’ imagination while con-
veying content.
Videos, Dynamic Visualizations, and Online Demonstrations: The Check It Out,
Try It Out, and Think About It activities include video interviews, dynamic visualizations,
interactive images, and links to online demonstrations that can be watched as many times as
the student feels is necessary.

iv

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whimsical portraits. This portrait was of Rudolf II
the Holy Roman Emperor, depicted as Vertumnus,
Measuring Visual Acuity and Contrast
the Roman god of the seasons.

Sensitivity MCAT
that only differ in their skin tone or the characteristics of hair. In Check It Out: What Is It Like
Children
to Be Facemay experience
blind? difficulties
you can meet Kristenreading
and learnthe board what
firsthand in class, elderly
it is like drivers
to have may have
prosopagnosia. PREFAC E v
trouble reading road signs, and some of us struggle to read text or see
You might be surprised to find that Apple co-founder Steve Wozniak, primatologist Jane objects without the
help of corrective
Goodall,Itactor lenses. For a variety of reasons, it is useful to accurately measure acuity,
Check OutBrad Pitt, and
activities Colorado
direct governor
students Johna Hickenlooper
to watch short video that areconveys
all believed to experi-
concepts and
or sharpness,
ence some of vision.
aspects of One standard measure of acuity is the Snellen chart, which can be
prosopagnosia.
then the student answers questions which reinforce their understanding of the material.
found on the wall of almost every physician’s office. The Snellen chart presents letters of
diminishing size to the viewer, who tries to identify the letters from a standard distance. Per-
fectCheck
vision is It
characterized
Out What as 20/20 (or 6/6
Is It Like toin Bemetric
Facecountries),
blind? which means the individual
sees at 20 feet (6 meters) what a person with normal vision sees at the same distance. By the
same measure, an individual with 20/40 vision sees at 20 feet It can(6 meters)
be hardwhat a personthe
to imagine with
normal vision sees at 40 feet (12 meters). everyday life of an individual with
The Snellen chart is used as a clinical screening tool prosopagnosiato diagnose common given that vision prob-
recog-
lems, but more sensitive measures also exist. For research purposes, nizing the scientists oftenseems
faces of others prefer tothe
Landolt C task (sometimes called the Landolt ring) as a measure be at the of heart of our
visual social
acuity expe- it
because
provides a finer level of analysis. In this test, a computer delivers rience. How might
a series of face
visualblindness
stimuli in

wavebreakmedia/Adobe Stock
the form of rings with a segment missing. On each trial, the change your behavior?
observer is asked to Watch
specify thethe
video What is it like to be face
direction of the missing segment using the numeric keypad on a computer. With each correct
blind? and then answer these
response, the stimulus reduces in size until the observer can no longer detect the missing seg-
questions.
ment (see Try It Out: The Landolt C Task).
Acuity is one useful measure of visual capabilities, but [Link] is WasnotKristen
the only a child or a young
measure or even
the only important measure. Another important measure is contrast adult when she learnedContrast
sensitivity. she had is
the difference between the intensity of a stimulus and its background prosopagnosia?
(Figure What does
5.5). that
Contrast
tell you about the adaptability
allows us to distinguish an object from its surroundings, known as the figure–ground orga-
of humans?
nization. If you think distinguishing an object from its surroundings is easy, it is only because
2. Kristen spoke about how her face blindness was an
our ancestors have been working on it for eons. You may remember impediment and how the itstory
mightofbethe
an princess
asset.
How might you have accommodated being face blind in a similar situation?
and the pea from your childhood in which the sensitive princess could detect even the small-
3. In
est pea behavioral
under neuroscience
her bedding. we often
In like see that
fashion, an behaviors
individual or with
functions
highthat are essential
contrast for ourcan
sensitivity
detect asurvival
stimulusare that
hardwired
differsinto their
only own dedicated
slightly structure.
in intensity fromWhat does the existence of the FFA
its background.
suggest about the importance of the ability to recognize who is in our clan and who is not in
Contrast sensitivity comes into play in many real-world tasks. Older drivers who test very
the evolution of humans?
successfully on the static Snellen test may perform poorly on measures of contrast sensitivity.
Their ability to perceive the road at night or in foggy conditions is considerably impaired when
compared to that of a typical younger driver.
Try It Out activities have the student perform some activity related to the chapter content
that deepens their understanding of the content and makes it memorable.

Try It Out The Landolt C Task

You can test your own vision using a Landolt C task by downloading the Freiburg
vision and contrast test (FrACT), which is available from your instructor. Watch
the brief video Freiburg Visual Acuity Task, collect your own measure of visual
acuity, interpret it in light of the worksheet for this activity, and then answer
these questions.

1. What advantages might there be to the Landolt C task over typical paper tests such as the
Snellen chart? What are the disadvantages?
2. Visual acuity is very important to your success. Search the internet using the phrase “How to
protect my vision” and identify two risks to your vision and what you can do to protect your
vision from them.

Spilich1e_FM.indd 5 12/19/22 7:49 PM

 JHVEPhoto / Adob
vi PREFACE a. b.
FIGURE 13.1 Predator and prey a. Predators such as the lion may sleep longer because they are
only at risk from larger predators. b. Prey such as the deer sleep less because they must be constantly on
Think About It activities provide the student with a topical issue in behavioral neurosci-
guard against a wide range of threats.
ence and then guides them to consider the implications of what they are learning in their life.

Think About It Has Civilization Changed How We Sleep?

Humans are a remarkably diverse spe-
cies, but if there is one trait that binds us

Gorodenkoff / Adobe Stock

all together, it is sleep. Everyone sleeps,
regardless of sex, culture, or personal
beliefs. Given the universality of sleep,
you might think that how we sleep has not
changed over time, but there is consider-
able evidence to suggest that our current
sleep habits are a recent development in
our history as a species.
In modern industrialized nations, people often wake after dawn, go to work or school, come
home and stay awake several hours after sunset, and then sleep through the night in one con-
tinuous stretch. This sleep pattern is known as monophasic sleep because it consists of a single
continuous event. But what if monophasic sleep is not our natural sleep pattern? Before the inven-
tion of electric lights, candles and lanterns were an expensive and not very effective source of
illumination, and before they were invented, our only source of illumination after dark was a fire.
Historians have recognized in reading diaries and other personal accounts of life in the pre-
industrial age that many people engaged in biphasic sleep. They went to bed as night fell, slept
2–4 hours, awoke around midnight and looked after their children or perhaps tended the fire, and
then went back to sleep for 4–5 more hours before awakening for their day. The Industrial Age
and its appetite for workers who could be present for a long and continuous shift plus the easy
availability of electric lights may have dramatically changed a basic human behavior.

Questions
1. Search the internet using the terms “biphasic sleep or second sleep in history.” Do
you think the evidence for second sleep in the preindustrial age is convincing? Why
or why not?
2. As a general rule, predators sleep more than prey. Humans today sleep like preda-
tors, yet some argue that our natural sleep pattern is biphasic sleep. Would bipha-
sic sleep be more adaptive for our distant ancestors whose existence in the African
savanna was more tenuous?
3. It is assumed that the Industrial Age created an environment in which it was advan-
tageous to replace biphasic sleep with monophasic sleep. In today’s world, people
may be part of a team that is distributed across several time zones. Would biphasic
sleep be an advantage for a person whose career requires them to interact part of the
day with people on the other side of the globe?

Spilich1e_FM.indd 6 12/19/22 7:50 PM

 A fewSimilarly, a plant
examples will product
help drive called
the point curare
home. For inhibits theknow
instance, we receptor
that a for acetylcholine (Ach) at
particular
variant
theofjunction
the adrenalin
where receptor,
a neuronthe αactivates
1 receptor,astimulates narrowingparalysis.
muscle, causing of the bloodForvessels
this reason, curare or
(vasoconstriction) and thereby increases blood pressure. Individuals suffering from low blood
similar drugs are often administered by anesthesiologists during surgery to keep the patient
pressure (hypotension) may benefit from an α1 agonist. On the other hand, an α1 receptor
immobile.
antagonist Without
decreases respiratoryinassistance,
vasoconstriction curare
the body, thereby can leadblood
decreasing to death by asphyxiation.
pressure. These
drugs are often recommended for treating high blood pressure (hypertension).
Similarly, a plant product called curare inhibits the receptor for acetylcholine (Ach) at PREFAC E vii
the junction where a neuron activates a muscle, causing paralysis. For this reason, curare or
similar drugs are often administered by anesthesiologists during surgery to keep the patient
Vibrant Illustrations and Interactivities: Before drug Neuroscience sets Agonist drug Antagonist drug
immobile. WithoutBehavioral a new
respiratory assistance, curare can lead standard
to death by asphyxiation.
for quality of images used in the text to depict concepts and neuroanatomy.

Natural
Before drug substance Agonist drug Antagonist drug

160 C H A P T E R 6 Audition, Chemosenses, and Cutaneous Senses

160 C H A P T E R 6 Audition, Chemosenses, and Cutaneous Senses Natural
substance
Frontal lobe Our sense of smell begins in the nasal cavity. Inside this space are
ofFrontal lobe
cerebrum
a seriesOur of sense
ridgesofknownsmell as
begins in the nasal
the turbinates thatcavity.
tumble Inside
the this space
air for better are
of cerebrum Antagonist
Olfactory tract a series ofbyridges
reception known as
the olfactory the turbinates
system. that tumble
The olfactory bulb the sitsair for better
at the base
Olfactory tract reception by the olfactory system. The olfactory bulb Agonist
sits at the base drug
of the skull just below eyebrow level and underneath the frontal region Cell
Olfactory bulb of the skull just below eyebrow level and underneath the drug
frontal region
Olfactory bulb of the brain. Olfactory receptors extend through the tiny perforations membrane
inofthe
thecribriform
brain. Olfactory platereceptors extend through
cavity. the tinyreceptors
perforations
Antagonist
Olfactory receptors and into the nasal These are
Olfactory receptors in the cribriform
embedded in the nasalplate and into
mucosa, andthe nasal
they
Agonist
are cavity.
constantly receptorsdrug
Thesesampling are Cell
the
drug
embedded in the nasal mucosa, and they are constantly
air for molecules called odorants, which are defined as any substance sampling the membrane
air for
that molecules
stimulates thecalled odorants,
olfactory [Link],areolfactory
defined as any substance
receptors trans-
thatthe
mit stimulates
resultingthe olfactory
signal to thesystem.
olfactory Finally,
tract,olfactory
which sends receptors trans-
the signal
mit the resulting signal to the olfactory tract,
into the brain. These key structures of the olfactory system are pre-which sends the signal
Nasal into the brain. These
Nasal
cavity sented in Figure 6.15. key structures of the olfactory system are pre-
Receptor
sented in Figure
cavity
site A look at the 6.15.
path that olfactory data travels provides important
clues A
Receptor to look at the pathof
the importance that
ourolfactory
sense of datasmelltravels
(Figure provides important
6.16). Odorants
clues to the
site traveling importance of our sense of smell (Figure
into the nasal cavity reach olfactory receptors embedded 6.16). Odorants in
traveling
the Normalinto
olfactory the nasal(step
epithelium cavity1).reach
Signalsolfactory
from the receptors
olfactoryembedded
bulb travelin Blocked
FIGURE 3.13 Agonists upthetocellular
Normal olfactory
a corticalepithelium
region known (stepas1).
theSignals
piriform from the olfactory
cortex bulb travel
that is dedicated
Blocked to cellular
and FIGURE 3.13 Agonists
antagonists at the up to
cellular a cortical
activity
olfaction (step 2). region
Fromknown
there asthethe piriform
signal cortex
diverges that
into is cellular
dedicated
multiple paths:to activity
toolfaction (step 2). From therewiththe signal diverges into multiple
and antagonists at the activity Enhanced activity paths:
receptor An agonist docks to a the amygdala (involved emotions),
Enhanced the entorhinal cortex and
receptor An agonist docks to a to the amygdala
hippocampus (involved
(associated with
with emotions),
memory), and the entorhinal
cellular
the thalamus cortex the
(where and
receptor and activates it, whereas
receptor and activates it, whereas
cellular
hippocampus
different senses (associated
are with (step
integrated) memory), 3).
activity The activity
andsignals
the thalamus
finally (where
terminate the
ananantagonist
antagonist blocks thereceptor
blocks the receptor
different senses are integrated) (step 3). The signals
in the frontal lobe (specifically the orbitofrontal cortex) and the insula, finally terminate
FIGURE 6.15 from from
The activating.
activating.
human olfactory system Your sense in the frontal lobe (specifically the orbitofrontal cortex) and the insula,
ofFIGURE 6.15 with
smell begins Thereceptors
human located
olfactory system
at the Your
top of the sense where sophisticated attentional and emotional decisions are created
nasal where
(step 4).sophisticated attentional and emotional decisions are created
of smell
cavity andbegins with receptors
just beneath located
the frontal [Link] the top
These of the nasal
receptors (stepThe 4).take-home message is that although olfaction is less import-
cavity from
extend and just
the beneath
olfactorythe frontal
bulb [Link]
through These receptorsin
perforations
extend from the
Process
olfactory
images
bulbofthrough
help
tinyThe
students
perforations
grasp
ant than mechanisms
the
The take-home
either orthat
visionmessage underlie
is
audition complex
that(asalthough
evidenced byprocesses
olfaction is less import-
it receiving less
the cribriform plate at the base the skull. olfactory in
the cribriformsuch asthe
plate at olfaction.
base of the skull. The olfactory
receptors sample the air for odorants and transmit the
ant than either vision or audition (as evidenced by it receiving less
receptorssignal
resulting sample the olfactory
to the air for odorants and transmit
tract, which thethe
transmits
resulting
signal intosignal to the olfactory tract, which transmits the
the brain.
signal into the brain.
2 The resulting signal travels 4 The signal terminates in
2 from
The the
resulting signal
olfactory travels
bulb to 4 the
The signallobe
frontal terminates
as the in
from
the the olfactory
piriform cortex. bulb to the frontaloflobe
perception as the
smell.
the piriform cortex. perception of smell.
Olfactory Frontal lobe
Olfactory
bulb Frontal lobe
1 Odorants enter the nose bulb
1 and
Odorants enter
stimulate the nose
olfactory Olfactory
and stimulate olfactory Olfactory
receptors Limbic system
receptors embedded in the Limbic system
receptors embedded in the receptors
olfactory epithelium.
olfactory epithelium. 2 Piriform Thalamus
2 Piriform
cortex Thalamus
cortex Hypothalamus
1
Odorant 1 Hypothalamus
Odorant
3
3

3 At the piriform cortex, the signal

3 divides,
At the piriform
engagingcortex, the signal
both emotional
divides,
and memoryengaging both
centers emotional
in the brain.
and memory centers in the brain.

FIGURE 6.16 The olfactory circuit The olfactory circuit begins in the nasal cavity and ends in the brain.
FIGURE 6.16 The olfactory circuit The olfactory circuit begins in the nasal cavity and ends in the brain.

Spilich1e_FM.indd 7 12/19/22 7:51 PM

 tialtive
information
processes suchis as
integrated
conscious and then
thought, associated
planning, with theWhen
and judgment. proper youresponse
are payingof your muscles.
atten-
When you
tion to an reach
in-classout to catchplanning
discussion, a ball, your
you are pathusing
to youryour parietal
degree, lobe,whether
or deciding and when you are finding
attending
youra party
waythe night before
around your aneighborhood
big exam is reallyor theputting
wisest choice, you are
together usingof
a piece your frontal lobe.
furniture that came unas-
sembled, Theyour
parietal lobe is
parietal the place
lobe where information from your many senses as well as spa-
is engaged.
tial information is integrated and then associated with the proper response of your muscles.
Your temporal lobe has much of the responsibility for language comprehension and pro-
When you reach out to catch a ball, you are using your parietal lobe, and when you are finding
duction.
your wayWhen
aroundyou areneighborhood
your chatting with a friendtogether
or putting about aclasses
piece ofand life, your
furniture temporal
that came unas-lobe is busy
viii PREFACE comprehending your friend’s speech and creating your verbal responses. The occipital lobe
sembled, your parietal lobe is engaged.
createsYouryour visual world.
temporal lobe hasFinally,
much of there is a fifth lobe
the responsibility that is comprehension
for language somewhat hidden and pro-in the lateral
duction. When
In addition, you are chatting
the enhanced e-text with a friend about
includes classes and life,distributed
35 interactivities your temporalthroughout
lobe is busy the
chapters comprehending your friend’s
that bring illustrations tospeech and creatingthe
life, reinforcing youractive
verballearning
[Link]
The occipital lobe this text
offers. creates your visual world. Finally, there is a fifth lobe that is somewhat hidden in the lateral

Central sulcus
Postcentral gyrus
248 CHAPTER 9 Reward, Addiction, and Psychopharmacology
Central sulcus
Parietal lobe
Postcentral gyrus
Snyder and Pert solved the puzzle by turning it on its head; opioids workFrontal as they do
lobe
Parietal lobe
because they mimic substances that our brain naturally creates. When you earn an A on an
assignment, master a difficult skill, thrill to a lover’s kiss, or see an old Frontal lobeInsula
friend after a long
(projected to surface)
absence, that pleasurable sensation is generated by your brain secreting Insula natural opioids. When
opioids are created internally, they are known as endogenous opioids. (projected to surface)
Opioids generated
Occipital lobe
outside the body such as morphine, heroin, and OxyContin, are exogenous opioids. Given
Occipital lobe
that exogenous opioids mimic the natural effects of endogenous opioids that are manufac-
Temporal lobe
tured by your own brain, you can now readily see why they are so addictive. TemporalPert’s
lobe work on
endogenous opioids was the start of an illustrious career, as you can see in this Advances in
Cerebellum Behavioral Neuroscience: Candace Pert and Opioid Receptors.
Cerebellum
Subsequent work with nonhuman species suggests that Anterior the opioid receptor has a long
Anterior
evolutionary history and one of its important functions is to enhance the development and
maintenance oflateral
Right
Right strongview
lateral social bonds, which support pair bonding and family ties. Given the
view
importance of social bonds among primates in general and humans in particular, it should not
FIGURE 4.4 Major lobes of the brain and associated anatomical landmarks The frontal,
be surprising
Major thatofexogenous
lobes the opioids are very reinforcing andlandmarks
highly addictive.
FIGURE 4.4
parietal, temporal, and occipital lobesbrain and associated
of the brain anatomical
are readily visible in this lateral view, as are theThe
sulcifrontal,
parietal,
and gyri that define them. The insula is hidden underneath the temporal and parietallateral
temporal, and occipital lobes of the brain are readily visible in this lobes. view, as are the sulci
and gyri that define [Link]
Synthetic The insula isMorphine
hidden underneath
and herointhe aretemporal
compounds andcreated
parietalfrom
[Link] and as
such are considered natural opioids. However, chemists have also devised synthetic opioids in
an effort to create an analgesic that relieves pain but is not addictive. So far that goal has not
Advancesbeen reached, but the
in Behavioral effort has led to synthetic
Neuroscience: opioids that
These profiles are more
in each powerful
chapter than any
highlight an natu-
indi-
ral opioid. Some examples of synthetic opioids are oxycodone, methadone
vidual whose work has significantly enhanced our understanding of the field. Students (often used as a sub-
are
directed tostitute for heroin in treatment), fentanyl, and carfentanil. These last two are extremely potent;
the researcher’s lab web page and answer brief questions about their work that are
fentanyl is thought to be 100 times more potent than morphine, and carfentanil is considered
designed to encourage the student to see the connection between science in the lab and their
to be 10,000 more potent than morphine. Using the potency of morphine as a reference point,
own lives. Table 9.6 provides a glimpse into the relative potency of common synthetic opioids.

Advances in Behavioral Neuroscience Candace Pert and Opioid Receptors

there must be an endogenous or innate opioid receptor in the

human brain. Although she was told that the search for this
receptor was a dead end and she should go back to her other
work, she came into the lab on a weekend and conducted the
experiment that made her world famous. Other discoveries
would soon follow.
Everett Collection Inc.

In a career that spanned four decades, Pert published over

250 research articles and changed the way we think about how
the brain affects our overall health. In her later work, she was a
pioneer of mind–body medicine and a proponent of the concept
that emotions have a powerful effect on physical health. She was
affectionately known as the Mother of Psychoneuroimmunology
and the Goddess of Neuroscience and encouraged her students
Sometimes in life, you have to listen to your own inner voice and and collaborators to think outside the box of accepted knowledge.
do what you think is right, even when others disagree. Candace Although Dr. Pert died in 2013, her discoveries and insights con-
Pert was an internationally renowned neuroscientist whose curi- tinue to influence the field today.
osity and intuition about nature led her to rewrite our understand-
Questions
ing of the link between our brain and our health.
Dr. Pert was born in 1946 in New York City; she graduated 1. One way to end opioid addiction would be to perma-
from Bryn Mawr College in 1970 and then entered the Ph.D. pro- nently blockade opioid receptors. Assuming for a
gram at Johns Hopkins, where she worked in the laboratory of moment that this is even possible, what unpleasant con-
Solomon Snyder, one of the world’s premier pharmacologists. Pert sequences might this cure create?
earned her doctorate in pharmacology from the Johns Hopkins 2. Pert was one of the founders of the field of psychoneu-
School of Medicine in 1974. roendocrinology. Search the internet for the term, and
In 1972, while still a graduate student in Snyder’s lab, Pert (a) summarize in one sentence what the field is about,
began to think seriously about a vexing question: what is the and (b) note one disorder that the field may provide a
neural mechanism that allows us to feel pleasure? Pert believed treatment for.

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 PREFAC E ix

Access to Actual Data: Behavioral Neuroscience provides instructors and their students
with access to data from actual experiments. Examples of data sets include:

• The effects of mild concussions upon attention

• Differences in visual acuity that distinguish varsity athletes from nonathletes
• The effects of low levels of alcohol upon visual search

Depending upon the course prerequisites regarding statistics, the student could be asked
to interpret a graph; interpret the results of a statistical test; or determine the independent and
dependent variables in a data set, select the appropriate statistical test, perform the analysis
themselves, and interpret the results. Instructors are provided with a completed and anno-
tated solution to all three levels of analysis so that students at every level of statistical prepa-
ration can think critically about the analysis and interpretation of data.
Ethics: Where appropriate, students are encouraged to consider the ethical implications
of scientific and clinical issues.
Emphasis on Major Themes: Major themes that are threaded throughout each chap-
ter include inhibitory and excitatory mechanisms that operate at multiple levels to regulate
behavior; the influence of our evolutionary past upon human behavior today; and clinical and
real-world implications.
Behavioral Neuroscience in the Real World: Your brain is fascinating in and of
itself but seeing how behavioral neuroscience speaks to everyday issues is an important part
of its appeal. Why are sugars and fats so appealing when consuming too much of them is
unhealthy? Why are some people unable to carry a tune? How is it that some people can per-
ceive the seams of a baseball being thrown at them at 90 miles an hour from 60 feet away in
less than two-tenths of a second? Why do some people have to engage in elaborate behavioral
rituals to leave their house? Why are some people suspicious of people that they have never
met? Behavioral Neuroscience considers these and many other meaningful questions.
MCAT Behavioral neuroscience is a popular major as well as a popular course for stu-
dents preparing for a career in medicine, and for that reason, topics that are tested on the
MCAT are clearly identified throughout the text. The Instructor’s Companion Website con-
tains a complete list of topics covered on the MCAT at the time of writing, and those topics
which Behavioral Neuroscience covers are clearly indicated.

To the Student
For most of my adult life, I have been fascinated by the human brain and how it creates behav-
ior, particularly higher-order cognitive behaviors such as memory and skilled visual percep-
tion, and how changes in one’s underlying neurology lead to cognitive disorders. I have spent
literally thousands of hours thinking about how I can best convey the subject of behavioral
neuroscience to students, and I have also spent more than 15,000 hours in front of students
trying my best to explain complex concepts in an interesting and accessible way. My goal in
writing this book is to make behavioral neuroscience as interesting and exciting to you as it is
to me, and to do so in a way that you will carry that understanding forward in your life. Con-
versations with my students have taught me much and so I hope to engage in conversation
with you also. Feel free to contact me directly at BNbyGS@[Link].

To the Instructor
In addition to the unique features listed previously, you will find that Behavioral Neuroscience
has the pedagogical learning aids that you have come to rely on.
Chapters are organized into manageable sections with learning objectives clearly spelled
out and addressed in each section. Every section ends with a section summary and compre-
hension checks in the form of multiple-choice questions that verify the student has acquired
the main themes. The art sets a new standard for quality, and interactive images that students
can use to check their comprehension are included in each chapter. There are dozens of videos
associated with the text that illustrate concepts; these can be folded into class presentations

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 x PREFACE

or made available via your school’s content management system. The Instructor’s Companion
Website includes many other videos that, while not directly referenced in the textbook, will be
of use to instructors. There is a computerized test bank for the easy construction of multiple-­
choice exams, PowerPoint presentations for every chapter, and an Instructor’s Manual with
a variety of tools and tips to benefit both seasoned faculty and those who may be teaching
behavioral neuroscience for the first time. Conversations with departmental colleagues about
new discoveries that we want to share with our students were always an important part of my
professional life and I hope to engage with you in such dialogues going forward. Feel free to
contact me directly at BNbyGS@[Link].

Acknowledgements
Writing a textbook is an enormous undertaking that involves many Andreas Keil, University of Florida
people. I’d like to thank my professors, whose wisdom and knowledge Mark Kristal, SUNY Buffalo
greatly influenced me: Tony Caggiula, Jim Devine, Judith Goggin, Charles Kutscher, Syracuse University
Harry Harlow, Dick Keesey, Alan Lesgold, Dominic Massaro, Don Donald Leitner, Saint Joseph’s University
Moss, Fred Mote, Chuck Perfetti, Ed Striker, Jim Voss, and Randy Daniel McConnell, University of Central Florida
Whitworth. During the writing process, I benefited from illuminat- Maura Mitrushina, California State University, Northridge
ing email conversations with hundreds of colleagues, ranging from Michael Owren, Emory University
Nobel laureates to first-year faculty at colleges and universities across Patricia Sharp, Bowling Green State
the globe. My colleagues in my own department and across academic
I was fortunate to fall into the hands of a talented team at Wiley.
divisions at Washington College provided me with a model of how to
My development editor, Rebecca Heider, was invaluable in creating
be a scholar and a professional educator. Every professor knows that
this text and taught me the nuts and bolts of academic publishing.
students teach us as much as we teach them, and I am grateful for all
Lauren Olesky, Dorothy Sinclair, Valerie Vargas, and Carolyn Wells
that my students have taught me. They continue to do so.
have provided expert guidance and advice at every point in the pro-
I want to thank the colleagues who reviewed the original pro-
cess. Rajeev Kumar and Sukhwinder Singh oversaw the creation of
posal and early samples of a book envisioned several years ago and
media that is essential to the text. This project started with a phone
whose feedback helped shape the development of what is now Behav-
call from Glenn Wilson, and his wisdom and energy have propelled
ioral Neuroscience; among them:
this project across the finish line. Thank you, Glenn. You are the very
J. Timothy Cannon, University of Scranton best.
Jeffrey Carlton, California State University, Sacramento Finally, I am indebted to Becky, my wife and partner in all things,
Rolf Greenwald, Central Washington University who tolerated my physical and mental absences throughout the writ-
Jeffrey Grimm, Western Washington University ing. I could not have done this without you.
Michael Hoane, Augusta University

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 Brief Contents

1 Evolution and Genetics 1

2 The Neuron and Action Potential 27

3 Synapses and Neurotransmitters 51

4 The Nervous System 78

5 Vision 110

6 Audition, Chemosenses, and Cutaneous Senses 144

7 Homeostatic Regulation of the Internal Environment 174

8 The Endocrine System 200

9 Reward, Addiction, and Psycho­pharmacology 227

10 Sexual Reproduction and Behavior 261

11 Neurodevelopment Across the Life Span 289

12 Learning and Conditioning 314

13 Sleep, Dreaming, and Circadian Rhythms 338

14 Language, Music, and the Brain 365

15 Cognitive Neuroscience 391

16 Psychiatric and Neurological Disorders 423

17 Social Neuroscience 456

INDEX 485

xi

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 Contents

1 Evolution and Genetics 1

Check It Out: Guillain-Barré Syndrome 37
2.3 Neural Transmission of a Signal 38
Stories of the Brain 2 Energy Produced through Imbalances 38
Ion Exchange 39
1.1 Human Origins 2 The Sodium-Potassium Pump 40
The History of the Earth 2 Think About It: Disrupting the Sodium-Potassium Pump 42
The Evolution of Mammals 3 The Action Potential 42
The Evolution of Primates 4 The All-or-None Law 44
The Evolution of Humans 4 2.4 White Matter, Gray Matter, and Glia 45
Check It Out: Are You Part Neanderthal? 5 White and Gray Matter 46
1.2 The Genetic Basis of Evolution 7 Glia 46
Mendel and the Theory of Heritability 7
Modern Genetics 9 Think About It: White or Gray Matter? 47
Think About It: Genetic Engineering 11 Conclusion 48
Genetic Variation 12 Critical Thinking Questions 49 / Dig Deeper into
Advances in Behavioral Neuroscience: Doudna, Behavioral Neuroscience 49 / Chapter 2 Before You Go On
Charpentier, and CRISPR-Cas9 14 Answers 49 / Chapter 2 Glossary 50
1.3 Evolution and the Environment 15
Natural Selection 15 3 Synapses and Neurotransmitters 51
The Evolution of Color Vision 16
Adaptations to Extreme Environments 17 Stories of the Brain 52
Try It Out: Are You Ready to Climb Everest? 18 3.1 The Synapse 52
Evolution and the Human Diet 19 The Presynaptic Mechanism 53
1.4 The Evolution of Social Behavior 21 The Postsynaptic Mechanism 53
Evidence for the Heritability of Behavior 21 A Closer Look at Interneuron Communication 55
The Survival Benefits of Social Behavior 22 Chemical versus Electrical Synapses 56
The Biological Basis for Modern Behavior 23 Check It Out: Electrical Synapses and Atrial Fibrillation 58
Check It Out: My Genetic History 24 3.2 Receptors and Postsynaptic Effects 59
Conclusion 25 Ionotropic and Metabotropic Receptors 59
Critical Thinking Questions 25 / Dig Deeper into Downregulation and Upregulation 61
Behavioral Neuroscience 25 / Chapter 1 Before You Go On Think About It: How Upregulation Explains Nicotine
Answers 26 / Chapter 1 Glossary 26 Cravings 62
Autoreceptors and Heteroreceptors 62
2 The Neuron and Action 3.3 An Overview of Major Neurotransmitters 64
Four Key Neurotransmitter Systems 64
Potential 27
Check It Out: The Sleepy Sickness Epidemic 66
Stories of the Brain 28 Other Neurotransmitters of Note 68
3.4 Disorders and Treatments Related
2.1 An Introduction to the Neuron 28
to Neurotransmission 71
An Overview of the Nervous System 28
Synaptopathies 71
The Discovery of the Neuron 29
Advances in Behavioral Neuroscience: Dimitri Kullmann
Check It Out: Staining a Neuron 30
and the Silent Synapse 72
2.2 The Structure of a Neuron 31
Alcohol, Drugs, and Addiction 73
Eukaryotic Cell Structure 31
Drug Treatments Targeting Neurotransmission 73
Neuron Components 32
Conclusion 75
Advances in Behavioral Neuroscience: Mark Harnett and Critical Thinking Questions 75 / Dig Deeper into
the Role of Dendrites 33 Behavioral Neuroscience 76 / Chapter 3 Before You Go On
Types of Neurons 34 Answers 76 / Chapter 3 Glossary 77

xii

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4 The Nervous System 78

The Geniculostriate Pathway 122
The Tectopulvinar Pathway 124
Stories of the Brain 79 Check It Out: The Eye of the Batter 124
4.1 Damage to the Visual Stream 125
An Overview of the Nervous System 79 5.4 The Visual Cortex 126
Divisions of the Nervous System 80 The Primary Visual Cortex 127
Gross Anatomy of the Brain 82 Processing in the Visual Cortex 128
Check It Out: Gross Anatomy of the Brain 83 Advances in Behavioral Neuroscience: Margaret Wong-Riley
The Brainstem 84 and the PVC 130
The Spinal Cord 85 The Extrastriate Cortex 131
Advances in Behavioral Neuroscience: Nicholas Check It Out: Motion Aftereffects 133
Hatsopoulos and Spinal Cord Injury Repair 87 The Ventral and Dorsal Streams 133
4.2 Major Structures and Support Systems 5.5 Color Vision 134
of the CNS 88 The Trichromatic Theory 135
The Cerebral Cortex 88 Color Vision Abnormalities 136
The Cerebellum 91 Color Sensitivity Across the Visual Spectrum 138
Essential Support Systems of the CNS 92 Think About It: Safety and Color Vision 139
Check It Out: Comparing Sheep and Human Brains 93 The Opponent Process Theory 139
4.3 Major Subcortical Processing Circuits 95 Try It Out: The Checkerboard Illusion 140
Check It Out: The Ventricular and Glymphatic Systems 95 Conclusion 141
The Corpus Callosum: Information Transfer 96 Critical Thinking Questions 141 / Dig Deeper into
The Thalamus and Hypothalamus: Integration Behavioral Neuroscience 142 / Chapter 5 Before You Go
and Regulation 96 On Answers 142 / Chapter 5 Glossary 143
The Limbic System: Emotion and Memory 96
The Basal Ganglia: Initiation and Control of Movement 98
Check It Out: Delgado’s Bull 99 6 Audition, Chemosenses, and
4.4 The Peripheral Nervous System 100 Cutaneous Senses 144
Divisions of the Peripheral Nervous System 101
The Neuromuscular Junction 102 Stories of the Brain 145
The Cranial Nerves 103 6.1 Audition: Hearing and the Ear 145
Think About It: Myasthenia Gravis and the Neuromuscular The Physics of Sound 145
Junction 104 The Anatomy of the Ear 146
Conclusion 106 Try It Out: Test the Limits of Your Hearing 146
Critical Thinking Questions 107 / Dig Deeper into Transmission of Sound Waves Through the Ear 149
Behavioral Neuroscience 107 / Chapter 4 Before You Go Addressing Hearing Deficits 151
On Answers 108 / Chapter 4 Glossary 108
Check It Out: Experience a Hearing Deficit and a Cochlear
Implant 152
5 Vision 110 6.2 Auditory Processing and the Vestibular
System 153
Stories of the Brain 111 Auditory Processing in the Brain 153
5.1 An Introduction to Vision 112 Amplitude and Frequency Coding 154
The Physics of Light 112 Auditory Localization 156
The Architecture of the Human Eye 113 Think About It: Auditory Sensitivity and Age 156
How the Focal Plane Affects Our Vision 114 The Vestibular System 157
Measuring Visual Acuity and Contrast Try It Out: Experience Auditory Localization 157
Sensitivity 115 6.3 The Chemosenses: Olfaction 159
Try It Out: The Landolt C Task 115 The Anatomy and Physiology of Olfaction 159
The Visual Processing Stream 116 Olfactory Receptors 161
5.2 The Retina 117 Individual Differences in Olfaction 161
Photoreceptors 117 Check It Out: Conversation with an Anosmic 162
Retinal Processing Beyond Rod and Cones 119 6.4 The Chemosenses: Gustation 163
Check It Out: Peeking into the Retina Through OCT 119 The Anatomy and Physiology of Taste 163
5.3 The Subcortical Centers 122 Taste Receptors 165

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Individual Differences in Taste 165

Try It Out: Are You a Picky Eater? 165
8 The Endocrine System 200

Advances in Behavioral Neuroscience: Linda Bartoshuk Stories of the Brain 201

and Supertasters 166
8.1 An Introduction to the Endocrine System 201
6.5 The Cutaneous Senses 167
The Discovery of Hormones as Messengers 201
Pressure and Vibration 168
Comparing the Endocrine and Nervous Systems 202
Pain 169
Try It Out: Fast- and Slow-Response Systems 203
Check It Out: Global Kinesthetic Deficit 169
Glands and Hormones 203
Conclusion 171
Negative Feedback and Endocrine Function 205
Critical Thinking Questions 171 / Dig Deeper into
8.2 Endocrine Glands in the CNS 206
Behavioral Neuroscience 171 / Chapter 6 Before You Go
The Hypothalamus, Pituitary Gland, and Pineal
On Answers 172 / Chapter 6 Glossary 173
Gland 206

7 Homeostatic Regulation of the Pituitary Hormones 208

Endocrine Regulation of Growth 209
Internal Environment 174 Think About It: Synthetic Growth Hormone Use
and Abuse 210
Stories of the Brain 175 8.3 Endocrine Glands in the Periphery 212
7.1 An Overview of Homeostasis 175 The Thymus and Thyroid 212
Balance and Change in the Body 175 The Pancreas and Glucoregulation 213
Feedback Systems 177 Check It Out: Glucoregulation 214
Try It Out: Negative Feedback Systems in Nature and Think About It: Preventing Type 2 Diabetes 215
Technology 177 The Adrenal Glands and Stress 216
7.2 Homeostatic Regulation of Temperature 178 The Gonads and Sexual Development 217
Ectothermic versus Endothermic 8.4 The Stress Response 218
Thermoregulation 178 The HPA Axis 218
The Neural Basis of Thermoregulation 178 The General Adaptational Syndrome 221
Hyperthermia and Hypothermia 180 Post-Traumatic Stress Disorder 222
Think About It: Avoiding Exercise-Induced Advances in Behavioral Neuroscience: Carrie Hill Kennedy
Hyperthermia 180 and PTSD 223
7.3 Homeostatic Regulation of Fluids Through Conclusion 224
Thirst 183 Critical Thinking Questions 224 / Dig Deeper into
Volumetric Thirst and the Renin–Angiotensin Behavioral Neuroscience 225 / Chapter 8 Before You Go
System 183 On Answers 225 / Chapter 8 Glossary 226
Osmotic Thirst and Cellular Sodium
Concentration 184
Disorders Related to Fluid Imbalance 185
9 Reward, Addiction, and
Try It Out: Recognizing the Dangers of Fluid Imbalance 185 Psycho­pharmacology 227
7.4 Homeostatic Regulation of Hunger and Satiety 186
The Gastrointestinal Tract 186 Stories of the Brain 228
The Neural Basis of Hunger and Satiety 188 9.1 The Biological Basis for Reward 228
Advances in Behavioral Neuroscience: Philip Teitelbaum Discovering the Biological Basis for Reward 229
and the Motivation to Eat 188 Reward Centers in the Brain 229
Hormonal Regulation of Hunger 189 Think About It: The Ethics of Deep Brain Stimulation 230
Hormonal Regulation of Satiety 190 The Mesolimbic and Mesocortical Pathways 231
7.5 Disorders of Hunger and Satiety 191 The Reward Circuit and Human Behavior 232
Obesity 192 9.2 Addiction as Dysfunction of the Reward Circuit 234
Prader-Willi Syndrome 194 The Genetic Basis for Addiction 234
Anorexia Nervosa 195 Substance Abuse Disorders 235
The Challenges of Treating Disordered Eating 196 Risk Factors for Addiction 236
Conclusion 197 Try It Out: Diagnosing Addictive Behavior 236
Critical Thinking Questions 197 / Dig Deeper into 9.3 The Essentials of Psychopharmacology 238
Behavioral Neuroscience 198 / Chapter 7 Before You Go How Drugs Enter the Body 238
On Answers 199 / Chapter 7 Glossary 199 Pharmacodynamics 240

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Think About It: Behavioral Tolerance and Drug Critical Thinking Questions 286 / Dig Deeper into
Overdoses 242 Behavioral Neuroscience 286 / Chapter 10 Before You Go
Pharmacokinetics 242 On Answers 287 / Chapter 10 Glossary 288
9.4 Intoxicants 244
Alcohol 244
Check It Out: The Effects of Alcohol on Visual Search 246
11 Neurodevelopment Across the
Opioids 247 Life Span 289
Advances in Behavioral Neuroscience: Candace Pert and
Opioid Receptors 248 Stories of the Brain 290
Marijuana 250 11.1 Prenatal Development of Brain
9.5 Stimulants and Other Commonly Abused Structures 290
Drugs 252 Embryonic Neurodevelopment 291
Nicotine 252 Think About It: Developing Brains at Risk 292
Caffeine 253 Fetal Neurodevelopment 293
Think About It: Do We Abuse Caffeine? 254 11.2 Prenatal Neurodevelopment at the Cellular
Other Commonly Abused Drugs 254 Level 295
Conclusion 257 Neurodevelopment as an Organizational
Critical Thinking Questions 258 / Dig Deeper into Process 295
Behavioral Neuroscience 258 / Chapter 9 Before You Go Check It Out: Visualizing Neuromigration 297
On Answers 259 / Chapter 9 Glossary 259 Guidance for Neuromigration 300
11.3 Postnatal Neurodevelopment 301
10 Sexual Reproduction and Environmental Influences on Postnatal
Neurodevelopment 301
Behavior 261
Advances in Behavioral Neuroscience: Marian Diamond and
Stories of the Brain 262 Neuroplasticity 302
The Adolescent Brain 304
10.1 An Overview of Sex and Reproduction 262 Try It Out: Are You a Risk Taker? 305
Asexual and Sexual Reproduction 263 The Aging Brain 305
Male and Female Reproductive Anatomy 264 11.4 Neurodevelopmental Disorders 307
Sex versus Gender 265 Defects in Neurulation 307
Try It Out: Sex Roles and Gender Roles 267 Check It Out: Spina Bifida 308
10.2 Prenatal Sexual Organization 268 Fetal Alcohol Syndrome 308
Genetic Influences on Biological Sex 268 Attention-Deficit/Hyperactivity Disorder 309
Sexual Differentiation in Utero 269 Williams Syndrome 310
Hormonal Influences on Prenatal Sexual Schizophrenia 310
Development 270 Conclusion 311
Try It Out: The D2:D4 Ratio 271 Critical Thinking Questions 311 / Dig Deeper into
10.3 The HPG Axis and Life-Span Sexual Behavioral Neuroscience 312 / Chapter 11 Before You Go
Development 273 On Answers 312 / Chapter 11 Glossary 313
Puberty 273
Think About It: Environmental Effects on Puberty
Onset 274 12 Learning and Conditioning 314
Hormonal Control of the Menstrual Cycle 276
Hormonal Activation of Parental Behavior 277 Stories of the Brain 315
Sexuality and Aging 278 12.1 Classical Conditioning 315
10.4 Sexual and Reproductive Behaviors 279 Innate Versus Learned Behavior 315
The Human Sexual Response 279 Pavlov and the Discovery of Classical
Sexual Orientation 282 Conditioning 316
Advances in Behavioral Neuroscience: Simon LeVay Check It Out: Experience Conditioning 317
and the Neural Basis of Sexual Try It Out: Classical Conditioning in Everyday Life 319
Orientation 283 Watson and the Little Albert Experiment 319
The Biological Basis of Attraction 284 Think About It: Ethics in Psychology Research 319
Check It Out: Are Average Faces More Attractive? 285 Conditioned Taste Aversion 320
Conclusion 286 12.2 Operant Conditioning 322

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 xvi CO N TENTS

The Discovery of Operant Conditioning 322

Reinforcement and Punishment 322
14 Language, Music, and the
Conditioning Phenomena 323 Brain 365
Try It Out: Turning Vices into Virtues 324
Stories of the Brain 366
Reinforcement Schedules 325
Try It Out: Operant Conditioning in Everyday Life 326 14.1 The Evolution of Human Language 366
12.3 Learning and the Brain 327 Communication in the Animal Kingdom 366
Where Memories Are Stored 327 Check It Out: The McGurk Effect 367
The Neuronal Basis of Learning 329 The FOXP2 Gene 368
Long-Term Potentiation 330 Evidence for Speech in Our Ancestry 368
Check It Out: Long-Term Potentiation 331 14.2 The Neural Basis of Language 370
NMDA and AMPA Receptors 332 The Discovery of Broca’s and Wernicke’s Areas 370
Advances in Behavioral Neuroscience: Karl Deisseroth and The Geschwind–Wernicke Model of Language 372
Optogenetics 334 The Two-Circuit Model of Language Comprehension
Conclusion 335 and Production 373
Critical Thinking Questions 335 / Dig Deeper into Contributions of the Right and Left Hemispheres 374
Behavioral Neuroscience 335 / Chapter 12 Before You Go Try it Out: The Dichotic Listening Task 375
On Answers 336 / Chapter 12 Glossary 336 14.3 The Neural Basis of Reading and Writing 377
The Invention of Reading and Writing 377
Reading Processes 378
13 Sleep, Dreaming, and Circadian Eye Movements and Reading 379
Rhythms 338 Reading and Writing Disorders 380
Check It Out: Reading Through the Eyes of a Person with
Stories of the Brain 339 Dyslexia 381
13.1 The Mystery of Sleep 339 14.4 The Neural Basis of Music 383
Evolutionary Patterns of Sleep 339 The Evolutionary Basis of Music 383
Think About It: Has Civilization Changed How We Sleep? 341 Characteristics of Music 384
The Purpose of Sleep 342 Think About It: Williams Syndrome 384
13.2 Sleep Stages and Dreaming 344 Music Processing in the Brain 385
The Study of Sleep 344 Advances in Behavioral Neuroscience: Isabelle Peretz and
The Stages of Sleep 345 Neural Processing of Music 385
Try It Out: What’s Your Sleep IQ? 347 Try It Out: Test Your Own Tone Discrimination 386
Dreaming 347 Conclusion 387
Try It Out: Are You a Lucid Dreamer? 348 Critical Thinking Questions 388 / Dig Deeper into
13.3 The Biological Basis of Sleep 349 Behavioral Neuroscience 388 / Chapter 14 Before You Go
Sensory Stimulation and Sleep 349 On Answers 389 / Chapter 14 Glossary 389
The Reticular Activating System and Sleep 350
The Brainstem, Hypothalamus, and Sleep 351 15 Cognitive Neuroscience 391
Neurotransmitters and Sleep 351
Advances in Behavioral Neuroscience: Gina Poe and the Stories of the Brain 392
Role of Sleep in Memory 352 15.1 Memory 392
13.4 Sleep Disruptions 353 The Three-Store Model of Memory 392
Drugs and Sleep 353 Sensory Memory 393
Sleep Disorders 355 Check It Out: Experience Echoic Memory 394
Think About It: Sleep Insufficiency 356 Working Memory 395
13.5 Circadian Rhythms 357 Try It Out: Measure Your Working Memory 396
Melatonin and the Sleep–Wake Cycle 358 Long-Term Memory 396
The Suprachiasmatic Nucleus 359 Amnesia 397
When Diurnal and Circadian Rhythms Are Think About It: A Life Lived Moment by Moment 398
Mismatched 359 15.2 Attention 399
Conclusion 361 Aspects of Attention 399
Critical Thinking Questions 362 / Dig Deeper into The Neural Basis of Attention 400
Behavioral Neuroscience 362 / Chapter 13 Before You Go Check It Out: Change Blindness 401
On Answers 363 / Chapter 13 Glossary 363 Hemineglect 401

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Think About It: Why Do Actors Enter from Stage Left? 402 Check It Out: The Facts About Strokes 445
15.3 Object Recognition 404 Traumatic Brain Injury 445
Visual Object Recognition 404 Try It Out: Concussions and Cognition 447
Challenges to Object Recognition 405 Dementia 447
Visual Agnosia 406 Movement Disorders 449
Prosopagnosia 407 Check It Out: Understanding Tourette’s Syndrome 451
Check It Out: What Is It Like to Be Face blind? 408 Conclusion 452
15.4 Executive Function 409 Critical Thinking Questions 452 / Dig Deeper into
Aspects of Executive Function 409 Behavioral Neuroscience 453 / Chapter 16 Before You Go
The Neural Basis of Executive Function 410 On Answers 454 / Chapter 16 Glossary 454
Advances in Behavioral Neuroscience: Marie Banich and
Executive Function 412 17 Social Neuroscience 456
Evaluating Executive Function 412
Try It Out: The Stroop Task 413 Stories of the Brain 457
Psychosurgery as a Treatment for Executive 17.1 Emotions 457
Dysfunction 413 Defining Emotion 457
Think About It: Moniz, Psychosurgery, and the The Evolutionary Basis of Emotions 458
Lobotomy 414 Advances in Behavioral Neuroscience: Leda Cosmides and
15.5 Hemispheric Specialization 415 Evolutionary Psychology 459
Anatomical Differences Between the Theories of Emotions 460
Hemispheres 415 The Neural Basis of Emotions 461
Sperry and the Split-Brain Procedure 416 17.2 Communicating Internal States 463
Check It Out: Do We Have Two Brains? 418 Facial Expressions 463
The Mystery of Left-Handedness 418 Social and Duchenne Smiles 464
Try It Out: Are You Right- or Left-Handed? 419 Try It Out: Cultural Differences in Emotional
Conclusion 420 Expressions 464
Critical Thinking Questions 420 / Dig Deeper into Deception as an Adaptive Strategy 465
Behavioral Neuroscience 420 / Chapter 15 Before You Go Check It Out: What’s in a Smile? 465
On Answers 421 / Chapter 15 Glossary 421 Theory of Mind 466
Check It Out: Can You Tell the Truth from a Lie? 466
16 Psychiatric and Neurological 17.3 Beauty and Attraction 469
External Tells for Health and Reproductive
Disorders 423 Fitness 469
Ovulation and Attractiveness 472
Stories of the Brain 424 Hormonal Influences on Pair Bonding 472
16.1 Mood Disorders 424 17.4 Human Social Networks 473
An Overview of Depression 425 Evolutionary Advantages of Group
The Neural Basis of Depression 426 Membership 473
Advances in Behavioral Neuroscience: Avshalom Caspi and Signaling Group Membership 474
the Genetic Basis of Depression 427 Ingroups and Outgroups 475
Treatment of Depression 428 Prejudice and Stereotypes 475
Bipolar Disorder 430 Think About It: The Implicit Association Test 477
16.2 Anxiety Disorders 432 17.5 Autism Spectrum Disorder and Emotional
General Anxiety Disorder 433 Blindness 478
Panic Disorder 434 Autism Spectrum Disorder 479
Obsessive–Compulsive Disorder 435 Difficulties in Processing Emotional Cues 480
Think About It: OCD, Compulsions, and Rituals 436 The Neurology of Asperger’s Syndrome 480
16.3 Consciousness Disorders 437 Check It Out: Lindsay’s Social World 481
Schizophrenia 438 Conclusion 482
Think About It: The Cost of Schizophrenia 441 Critical Thinking Questions 482 / Dig Deeper into
Delusions of Identity 442 Behavioral Neuroscience 482 / Chapter 17 Before You Go
16.4 Neurological Disorders 443 On Answers 483 / Chapter 17 Glossary 484
Bacterial and Viral Infections 443
Cerebrovascular Accidents 443 INDEX 485

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