Professional Documents
Culture Documents
Overcoming Blindness
7.1 Neural Development Is Guided by the Interaction of Genetic and Environmental
Factors
7.2 Development of the Nervous System Can Be Divided into Six Distinct Stages
BOX 7.1 Transgenic and Knockout Mice
BOX 7.2 Degeneration and Regeneration of Nervous Tissue
7.3 Lifelong Synapse Rearrangement Is Guided by Experience
7.4 Experience Can Alter Gene Expression to Affect Brain Development
The Cutting Edge ■ Harnessing Glia to Reverse an Inherited Brain Disorder
7.5 The Brain Continues to Change as We Grow Older
Visual Summary
10 Vision
From Eye to Brain
When Seeing Isn’t Seeing
10.1 The Retina Transduces Light into Neuronal Activity
10.2 Properties of the Retina Shape Many Aspects of Our Vision
10.3 Neural Signals Travel from the Retina to Several Brain Regions
BOX 10.1 Eyes with Lenses Have Evolved in Several Phyla
10.4 Neurons at Different Levels of the Visual System Have Very Different Receptive
Fields
10.5 Color Vision Depends on Special Channels from the Retinal Cones through
Cortical Area V4
BOX 10.2 Most Mammalian Species Have Some Color Vision
10.6 The Many Cortical Visual Areas Are Organized into Two Major Streams
10.7 Visual Neuroscience Can Be Applied to Alleviate Some Visual Deficiencies
The Cutting Edge ■ Seeing the Light
Visual Summary
12 Sex
Evolutionary, Hormonal, and Neural Bases
Genitals and Gender: What Makes Us Male and Female?
SEXUAL BEHAVIOR
12.1 Reproductive Behavior Can Be Divided into Four Stages
12.2 The Neural Circuitry of the Brain Regulates Reproductive Behavior
The Cutting Edge ■ Sexual Experience Solidifies Neural Circuits for Mating
12.3 The Hallmark of Human Sexual Behavior Is Diversity
12.4 For Many Vertebrates, Parental Care Determines Offspring Survival
SEXUAL DIFFERENTIATION
12.5 Sex Determination and Sexual Differentiation Occur Early in Development
12.6 Gonadal Hormones Direct Sexual Differentiation of the Brain and Behavior
BOX 12.1 The Paradoxical Sexual Differentiation of the Spotted Hyena
12.7 Do Fetal Hormones Masculinize Human Behaviors in Adulthood?
Visual Summary
13 Homeostasis
Active Regulation of the Internal Environment
Harsh Reality TV
13.1 Homeostasis Maintains a Consistent Internal Environment: The Example of
Thermoregulation
BOX 13.1 Physiological and Behavioral Thermoregulation Are Integrated
FLUID REGULATION
13.2 Water Shuttles between Two Body Compartments
13.3 Two Internal Cues Trigger Thirst
FOOD AND ENERGY REGULATION
13.4 Nutrient Regulation Helps Prepare for Future Needs
13.5 A Hypothalamic Appetite Controller Integrates Multiple Hunger Signals
The Cutting Edge ■ Friends with Benefits
13.6 Obesity and Eating Disorders Are Difficult to Treat
BOX 13.2 Body Fat Stores Are Tightly Regulated, Even after Surgical Removal of Fat
Visual Summary
16 Psychopathology
Biological Basis of Behavioral Disorders
“The Voice”
16.1 Schizophrenia Is the Major Neurobiological Challenge in Psychiatry
BOX 16.1 Long-Term Effects of Antipsychotic Drugs
16.2 Depression Is the Most Prevalent Mood Disorder
The Cutting Edge ■ Can Our Genes Tell Us Which Drugs to Use?
BOX 16.2 The Season to Be Depressed?
16.3 Extreme Mood Cycles Define Bipolar Disorder
16.4 There Are Several Types of Anxiety Disorders
BOX 16.3 Tics, Twitches, and Snorts: The Unusual Character of Tourette's Syndrome
Visual Summary
Appendix
Glossary
References
Author Index
Subject Index
Preface
Twenty-four years ago, a new kind of textbook was published for University courses that were often called “Brain
and Behavior.” As the field evolved, the book’s title metamorphosed from Biological Psychology to Behavioral
Neuroscience, but the same drive to provide a definitive and comprehensive survey of the neurosciences lies at the
heart of all our efforts. We strive to keep the book up-to-date while keeping a conversational tone to make this
wealth of information not just accessible, but fascinating. The biggest change in this new edition is the development of
Learning Objectives for each segment of the book, with the idea that telegraphing what’s to come will focus
readers’ attention and facilitate learning. As you finish each section of text, it would be a good idea to go back and
read the associated Learning Objectives to see whether in fact you incorporated the material. If not, a quick review
of that text may be in order.
As always, there have been plenty of new findings to add to this edition. In fact, the problem we face is which of
the many, many new findings to leave out—those that are not quite essential for a survey of the field. We are pretty
picky about what we add, and still it seems like a deluge of new information and ideas. Hundreds of new papers are
cited in this edition. If that sounds like a lot, let us give you a perspective on how many new papers were omitted.
On our newsfeed site (www.biopsychology.com/news/), 1,299 new links were added in 2018 alone. Those are just
the findings that were important enough to get the attention of mass media reporters. As we note in Chapter 1, over
40,000 new articles indexed under “neuroscience” appeared that year in PubMed. It would take a thick tome just to
list the titles of the papers from 2018!
While being very, very selective in sampling this flood of findings, we have made substantial changes in every
chapter. For example, in Chapter 3 we have a new figure comparing “kiss and run” synapses with more traditional
models of synaptic transmission. Chapter 5 has new material about a hormone secreted from bone that acts on the
hypothalamus to reduce appetite. We totally reorganized Chapter 7 for a more streamlined approach and discuss the
growing doubts about whether amyloid deposits cause Alzheimer’s. Chapter 9 needed a new figure comparing
transduction in the five taste receptors. Chapter 13 talks about yet another factor affecting appetite, glucagon-like
peptide 1. Chapter 16 now discusses the logic of pharmacogenomics to treat depression, while Chapter 18 was
thoroughly reorganized and includes more about executive function. Honestly, we could go on like this for every
chapter. Clearly this is an exciting era in the neurosciences. As Lewis Carrol put it, “We must run as fast as we can
just to stay in place!”
We’ve also kept several very popular features from previous editions: The Cutting Edge appears in each
chapter, where we explore some of the most exciting examples of recent research, and each chapter ends with a
Visual Summary, where you can see graphic reminders as you review the principle findings that we just presented.
These Visual Summaries really shine online, where with just a click you can review figures, animations, and quizzes to
help integrate the material. We also continue to open each chapter with a gripping vignette, relating someone’s real-
life experiences that will be better understood as the content of the chapter unfolds, and we again replaced several of
these vignettes as more recent events bring to the surface many of the important issues in behavioral neuroscience.
Likewise we’ve retained the marginal glossary that makes it easy to find the definitions that unlock the material, as
well as two features to let you burrow in on a particular subject: the online supplements called A Step Further cited
throughout the text, and the Recommended Reading at the close of each chapter.
You might think that approaching the quarter-century mark we’d be jaded about improving and revising our
presentations, but we still love it, perhaps because the dynamic and exciting pace of neuroscience research shows no
sign of abating soon. As always, we welcome all feedback, praise or criticism, cuts or additions, from our readers.
You can email us directly at behavneuro@gmail.com.
Acknowledgments
We continue to feel so lucky to work with the inestimable team at Sinauer Associates, now a part of Oxford
University Press, whose deep skills and generous guidance transform our hundreds of files, thousands of email
attachments and sometimes scrambled emails into yet another beautiful book. Again, we feel so grateful to benefit
from the experience and exquisite taste of others. In particular, the book could not exist without the contributions of
Senior Acquisition Editor Syd Carroll, Production Editor Alison Hornbeck, Production Manager Joan Gemme,
Book Designer and Production Specialist Annette Rapier, and Media and Supplements Editor Zan Carter and her
crew. We also fondly bid adieu to the recently retired Chris Small, Production Manager for all our previous editions.
We hope you’re enjoying yourself, Chris, but how could you abandon us!? A cadre of commandos delved deep in
the archives to deal with copyrights and permissions, so we salute you Michele Beckta, Mark Siddall, and Tracy
Marton. We’d also like to thank our copy editor Lou Doucette, and our longtime art studio, Dragonfly Media, who
bring amazing skill and commitment to make us look good.
We must also thank the founder of Sinauer Associates, Andy Sinauer, for his unwavering support over the years,
with a touch of sadness upon his retirement. We are so proud to be a part of Andy’s tremendous legacy, begun all
those years ago with From Neuron to Brain, creating gorgeous books that make even the most complex topics
accessible and enjoyable.
By this point in the evolution of the book, we have benefited from the wisdom and advice of hundreds of
colleagues who have generously served as reviewers of past editions. Although we don’t have the space to list them
all, we want to acknowledge that in many ways the book you are holding is the product of a whole community of
neuroscientists. In this, the Ninth Edition, the following colleagues have provided invaluable critique and commentary:
This invaluable online resource helps students make connections between the science of behavioral neuroscience and
their daily lives and keeps them apprised of the latest developments in the field. The site includes links to thousands
of news stories, all organized both by keyword and by textbook chapter. The site is updated 3–4 times per week, so
it includes up-to-the-minute information. NewsLink updates are also available on Facebook
(facebook.com/behavioralneuroscience).
Value Options
eBook
(ISBN 978-1-60535-937-3)
Ideal for self-study, the Behavioral Neuroscience, Ninth Edition, enhanced eBook delivers the full suite of digital
resources in a format that is independent from any courseware or learning management system platform. The
enhanced eBook is available through leading higher education eBook vendors.
Looseleaf Textbook
(ISBN 978-1-60535-936-6)
Behavioral Neuroscience, Ninth Edition is also available in a three-hole-punched, looseleaf format. Students can
take just the sections they need to class and can easily integrate instructor material with the text.
Introduction
Scope and Outlook
Machine or Human?
In the near future depicted in the HBO series Westworld, people visit a theme park set in the Old West, with steam
locomotives, saloons, and brothels, populated with androids, called “hosts,” to entertain humans. The mechanical hosts
provide their guests with anything, from casual banter to gunfights, harmless flirting to kinky sex, the only restriction
being that the robots are never to harm the humans. The android hosts are so lifelike in appearance and behavior that
visitors may have a hard time distinguishing whether someone is a fellow guest or a robot. To make the androids’
simulation of humans complete, they are given backstories, false memories of a life before their appearance for each
new batch of guests. Importantly, none of the androids know that they are mechanical beings rather than humans. It’s
probably not much of a spoiler to say that several plot lines in the series hinge on androids slowly discovering their true
nature, moving from shock and shame that they are mere machines, to openly rebelling from the notion that they are to
be used, and abused, as mere playthings for the humans.
We aren’t told too much about how the android “brains” in Westworld work, because, of course, such technology
remains far outside our grasp, so the writers, reduced to mere speculation, remain rather vague. But apparently the
knowledge and personality for any particular android lies in a “control unit,” a golf-ball-size device that can be extracted
from the head of one host and implanted into the head of another, interchangeable body. Presumably, if we had enough
knowledge and surgical skill, we could remove your brain from your head and connect it up to the head of some other
body. Would you still be you? Even if we put your brain into a body of the opposite sex? Come to think of it, are you
entirely sure there is a brain in your head, and not one of those control units?
Our aim in this book is to help you learn what is known so far about how brains work, and about how much more
we have yet to learn. We will explore the many ways in which the structures and actions of the brain produce mind
and behavior. But that is only half of our task. We are also interested in the ways in which behavior and experience
modify the structures and actions of the brain. One of the most important lessons we want to convey is that
interactions between brain and behavior are reciprocal. The brain controls behavior and, in turn, behavior and
experience alter the brain.
We hope to give an interesting account of the main ideas and research in behavioral neuroscience, which is of
great popular as well as scientific interest. Most important, we try to communicate our own interest and excitement
about the mysteries of mind and body.
Brain Explorer
1.1 The Brain Is Full of Surprises
Le arning Obje ctive s
After reading this section, you should be able to:
1.1.1 Name the main type of cells found in the brain, and name the connections between them.
1.1.2 List the names of some of the many fields of study related to behavioral neuroscience.
1.1.3 Describe five different perspectives taken in understanding the biology of behavior.
I used to think that the brain was the most wonderful organ in my body. Then I realized who was telling me this.
—Emo Philips
(American comedian)
Of course we should always consider the source when evaluating an idea, but even so, the brain indeed seems like a
pretty wonderful organ. For one thing, brains produced the entire extent of human knowledge, everything we
understand about the universe, however limited that may be. Brains also produced every written description of that
hard-won knowledge (including this book you hold in your hands), as well as every work of visual art, from doodles
to the sweeping frescos on the ceiling of the Sistine Chapel.
Most of us have a hard time grasping the idea of a billion of anything, but your head contains an estimated 86
billion nerve cells, or neurons (from the Greek word for “nerve” or “cord”) (Herculano-Houzel, 2012). Each neuron
contacts many other cells at points called synapses, so there are trillions of those between your ears. A specialized
extension of neurons, called an axon, is microscopically slender, yet it may be several feet long. We’ll learn that
axons produce electrical impulses that travel hundreds of miles per hour. Figure 1.1 offers a list of just a few of the
things we will learn about the human brain in the course of this book. All this hardware isn’t just for show—it allows
you to take in all the information in that figure in less than a minute.
1.1 Your Brain by the Numbers The cerebral cortex is the outermost portion of the brain.
1. Describing behavior
2. Observing the development of behavior and its biological characteristics over the life-span
3. Studying the biological mechanisms of behavior
4. Studying applications of behavioral neuroscience—for example, its application to dysfunctions of human behavior
5. Studying the evolution of behavior
These perspectives are discussed in the sections that follow, and Table 1.1 illustrates how each perspective can be
applied to three kinds of behavior.
We compare species to learn how the brain and behavior have evolved
Nature is conservative. Once particular features of the body or behavior evolve, they may be maintained for millions
of years and may be seen in animals that otherwise appear very different. For example, the electrical messages used
by nerve cells (see Chapter 3) are essentially the same in a jellyfish, a cockroach, and a human being. Some of the
chemical compounds that transmit messages through the bloodstream (hormones) are also the same in diverse
animals (see Chapter 5). Species share these conserved characteristics because the features first arose in a shared
ancestor (Box 1.1 on the next page). But mere similarity of a feature between species does not guarantee that the
feature came from a common ancestral species. Similar solutions to a problem may have evolved independently in
different classes of animals.
Charles Darwin’s theory of evolution through natural selection is central to all modern biology. From this
perspective emerge two rather different emphases: (1) the continuity of behavior and biological processes among
species that reflects shared ancestry and (2) the species-specific differences in behavior and biology that have
evolved as adaptations to different environments.
Study Questions
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1.2 Three Approaches Relate Brain and Behavior
Le arning Obje ctive s
After reading this section, you should be able to:
1.2.1 Differentiate between the independent and dependent variables in scientific experiments.
1.2.2 Name the type of research in which a part of the brain is manipulated to observe effects on behavior, and
offer examples.
1.2.3 Name the type of research in which behavior or experience is manipulated to observe effects on the brain,
and offer examples.
1.2.4 Describe correlational research about the brain and behavior, and offer examples.
1.2.5 Explain why the brain must be capable of changing its structure, and name the term to describe that
changeability.
Behavioral neuroscientists use three approaches to understand the relationship between brain and behavior: somatic
intervention, behavioral intervention, and correlation. In the most common approach, somatic intervention (Figure
1.3A), we alter a structure or function of the brain or body to see how this alteration changes behavior. Here,
somatic intervention is the independent variable, and the behavioral effect is the dependent variable; that is, the
resulting behavior depends on how the brain has been altered. For example, in response to mild electrical stimulation
of one part of her brain, not only did one patient laugh, but she found whatever she happened to be looking at
amusing (Fried et al., 1998).
1.3 Three Main Approaches to Studying the Neuroscience of Behavior (A) In somatic intervention, investigators
change the body structure or chemistry of an animal in some way and observe and measure any resulting behavioral
effects. (B) Conversely, in behavioral intervention, researchers change an animal’s behavior or its environment and try to
ascertain whether the change results in physiological or anatomical changes. (C) Measurements of both kinds of variables
allow researchers to arrive at correlations between somatic changes and behavioral changes. (D) Each approach enriches
and informs the others.
In later chapters we describe many kinds of somatic intervention with both humans and other animals, as in the
following examples:
A hormone is administered to some animals but not to others; various behaviors of the two groups are later
compared.
A part of the brain is stimulated electrically, or by use of light to stimulate only a particular class of neurons, and
behavioral effects are observed.
A connection between two parts of the nervous system is cut, and changes in behavior are measured.
The approach opposite to somatic intervention is psychological or behavioral intervention (Figure 1.3B). In this
approach, the scientist intervenes in the behavior or experience of an organism and looks for resulting changes in
body structure or function. Here, behavior is the independent variable, and change in the body is the dependent
variable. Among the examples that we will consider in later chapters are the following:
Putting two adults of opposite sex together may lead to increased secretion of certain hormones.
Exposing a person or animal to a visual stimulus provokes changes in electrical activity and blood flow in parts of
the brain.
Training of animals in a maze is accompanied by electrical, biochemical, and anatomical changes in parts of their
brains.
The third approach to brain-behavior relations, correlation (Figure 1.3C), consists of finding the extent to which
a given body measure varies with a given behavioral measure. Later we will examine the following questions, among
others:
Are people with large brains more intelligent than people with smaller brains (a topic we’ll take up later in this
chapter)?
Are individual differences in sexual behavior correlated with levels of certain hormones in the individuals?
Is the severity of schizophrenia correlated with the magnitude of changes in brain structure?
Such correlations should not be taken as proof of causal relationship. For one thing, even if a causal relation
exists, the correlation does not reveal its direction—that is, which variable is independent and which is dependent.
For another, two factors might be correlated only because a third, unknown factor affects the two factors measured.
If you and your study partner get similar scores on an exam, that’s not because your performance caused her to get
the score she did, or vice versa. What a correlation does suggest is that the two variables are linked in some way—
directly or indirectly. Such a correlation often stimulates investigators to formulate hypotheses and to test them by
somatic or behavioral intervention. Only by moving on to such intervention approaches can we establish whether one
variable is causing changes in the other.
Combining these three approaches yields the circle diagram of Figure 1.3D, incorporating the basic approaches to
studying relationships between bodily processes and behavior. It also emphasizes the theme that the relations
between brain and behavior are reciprocal: each affects the other in an ongoing cycle of bodily and behavioral
interactions. We will see examples of this reciprocal relationship throughout the book.
1.4 The Role of Play in Brain Development A brain region involved in processing odors (the posterodorsal portion of
the medial amygdala) was smaller in male rats housed individually than in males housed together and allowed to play. Other
nearby regions were identical in the two groups. (After B. M. Cooke et al., 2000. Behav Brain Res 117: 107–113.)
Here’s an example of how social influences can affect human brain function. When people were asked to put a
hand into moderately hot water (47°C), part of the brain became active, presumably because of the discomfort
involved (Rainville et al., 1997). But people who were led to believe the water would be very hot had a more
activated brain than did those led to believe the discomfort would be minimal (Figure 1.5), even though the water
was the same temperature for everyone. The socially induced psychological expectation affected the magnitude of
the brain response, even though the physical stimulus was exactly the same. (By the way, the people with the more
activated brains also reported feeling more pain.)
1.5 Pictures of Pain People told to expect only mild discomfort from putting a hand into 47°C water (left) showed less
activation in a particular brain region (the anterior cingulate cortex) than did people expecting more discomfort (right) from
water of the very same temperature. Areas of high activation are indicated by orange, red, and white.
In most cases, biological and social factors continually interact and affect each other in an ongoing series of events
as behavior unfolds. For example, the level of the hormone testosterone in circulation can affect dominance behavior
and aggression (see Chapter 15). The dominance may be exhibited in a great variety of social settings, ranging from
playing chess to physical aggression. In humans and other primates, the level of testosterone correlates positively
with the degree of dominance and with the amount of aggression exhibited. Winning a contest, whether a game of
chess or a boxing match, raises the level of testosterone; losing a contest lowers the level. Thus, at any moment the
level of testosterone is determined, in part, by recent dominant-submissive social experience, and the level of
testosterone determines, in part, the degree of dominance and aggression in the future. Of course, social and cultural
factors also help determine the frequency of aggression; cross-cultural differences in rates of aggression exist that
cannot be correlated with hormone levels, and ways of expressing aggression and dominance are influenced by
sociocultural factors.
Perhaps nothing distinguishes neuroscience from the other sciences more clearly than this fascination with
neuroplasticity and the role of experience. Neuroscientists have a pervasive interest in how experience physically
alters the brain and therefore affects future behavior. We will touch on this theme in every chapter of this book.
Study Questions
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1.3 Behavioral Neuroscientists Use Several Levels of Analysis
Le arning Obje ctive s
After reading this section, you should be able to:
1.3.1 Name and describe the scientific approach of explaining mechanisms at simpler and simpler levels.
1.3.2 Give a survey of important ongoing questions about the relationship between the brain and behavior.
1.3.3 Offer estimates of the extent of neurological and psychiatric disorders.
1.3.4 Explain the importance of research with animals for neuroscience, and discuss the ethics of such research.
Scientific explanations of systems or structures or functions usually involve breaking them down into smaller parts, as
a way of understanding them. This approach is known as reductionism. In principle, it is possible to reduce each
explanatory series down to the molecular or atomic level, though for practical reasons this extent of reductionism is
rare. For example, most chemists deal with large, complex molecules and the laws that govern them; seldom do they
seek explanations in terms of subatomic quarks and bosons.
Understanding behavior often requires several levels of biological analysis. The units of each level of analysis are
simpler in structure and organization than those of the level above. The levels of analysis range from social
interactions to the brain, continuing to successively less complex units until we arrive at single nerve cells and their
even simpler, molecular constituents.
Naturally, in all fields different problems are carried to different levels of analysis, and fruitful work is often being
done simultaneously by different workers at several levels (Figure 1.6). Thus, in their research on visual perception,
cognitive neuroscientists advance analytical descriptions of behavior. They try to determine how the eyes move while
looking at a visual pattern, or how the contrast among parts of the pattern determines its visibility. Meanwhile, other
behavioral neuroscientists study the differences in visual abilities among species and try to determine the adaptive
significance of these differences. For example, how is the presence (or absence) of color vision related to the life of a
species? At the same time, other investigators trace out brain structures and networks involved in different kinds of
visual discrimination. Still other scientists try to ascertain the electrical and chemical events that occur in the brain
during vision.
1.6 Levels of Analysis in Behavioral Neuroscience The scope of behavioral neuroscience ranges from the level of the
individual interacting with others, to the level of the molecule. Depending on the question at hand, investigators use
different techniques to focus on these many levels, but always with an eye toward how their findings apply to behavior.
1.7 “Tell Me Where Is Fancy Bred, Or in the Heart Or in the Head?” (A) The parts of the brain highlighted here
become especially active when a person thinks about his or her romantic partner. (B) Different brain regions are activated
when people perform four different language tasks. The techniques used to generate such images are described in Chapter
2.
The relationship between the brain and behavior is, on the one hand, very mysterious because it is difficult to
understand how a physical device, the brain, could be responsible for our subjective experiences of fear, love, and
awe. Yet despite this mystery, we all use our brains every day. Perhaps it is the “everyday miracle” aspect of the
topic that has generated so much folk wisdom about the brain. Think of it as “neuromythology.”
Sometimes these popular ideas about the brain are in line with our current knowledge, but in many cases we know
they are false. For example, the notion that we normally use only 10% of our brain is commonplace—a survey of
teachers found that nearly half of them agreed with this notion (Howard-Jones, 2014)—but it is patent nonsense.
Brain scans make it clear that the entire brain is activated by even fairly mundane tasks. Indeed, although the areas of
activation shown in Figure 1.7 appear rather small and discrete, we will show in Box 2.3 that experimenters must
work very hard to create images that separate activation related to a particular task from the background of
widespread, ongoing brain activity.
We offer a list of other commonly held beliefs about the brain and behavior on the website in A STEP FURTHER
1.1 : NEUROMYTHOLOGY: FACTS OR FABLES? Throughout the book we offer such opportunities for you to
explore a given topic in more detail on the website, bn9e.com.
1.8 The Toll of Brain Disorders Estimated numbers of people in the United States with neurological disorders (A) and
number of people worldwide with psychiatric disorders (B). (Part A after C. L. Gooch et al., 2017. Ann Neurol 81: 479–
484; B after H. Ritchie and M. Roser, 2019. "Mental Health". Published online at OurWorldInData.org. Retrieved from:
https://ourworldindata.org/mental-health. Underlying data available from http://ghdx.healthdata.org/gbd-results-tool.)
The toll of these disorders is enormous, in terms of both individual suffering and social costs (Demyttenaere et al.,
2004). The National Advisory Mental Health Council has estimated that direct and indirect costs of behavioral and
brain disorders amount to $400 billion a year in the United States alone. For example, the cost for treatment of
dementia (severely disordered thinking) exceeds the costs of treating cancer and heart disease combined. The World
Health Organization (2004) estimates that over 15% of all disease burden, in terms of lost productivity, is due to
mental disorders. The high cost in suffering and expense has compelled researchers to try to understand the
mechanisms involved in these disorders and to try to alleviate or even prevent them.
In this quest, the distinction between clinical and laboratory approaches begins to fade away. For example, when
clinicians encounter a pair of twins, one of whom has schizophrenia while the other seems healthy, the discovery of
structural differences in their brains (Figure 1.9) immediately raises questions for laboratory scientists: Did the
structural differences arise before the symptoms of schizophrenia, or the other way around? Were the brain
differences present at birth, or did they arise during puberty? Does medication that reduces symptoms affect brain
structure? When genes associated with schizophrenia in people are introduced into mice, will that change the mouse
brains (see Figure 16.7)? This last question is just one instance of when working with animals is essential, an issue we
address next.
1.9 Identical Twins but Nonidentical Brains and Behavior In these images of the brains of identical twins, the fluid-
filled cerebral ventricles are prominent as dark “butterfly” shapes. The brain of the twin with schizophrenia (A) has the
enlarged cerebral ventricles that some researchers believe are characteristic of this disorder. The other twin does not have
schizophrenia; his brain (B) clearly has smaller ventricles.
1.10 Car Firebombed by Animal Rights Activists The extremists targeted the cars and homes of two scientists who
work with animals at the University of California in Santa Cruz in 2008. The next year, the car of a researcher at UCLA
was torched.
Perhaps in a future where robots can be made that look and act like humans, methods will be available to clearly
see all the processes at work in a living, working human brain. In the meantime, there’s no substitute for research
with animal subjects. Every chapter in this book is teeming with information that was gathered from humane
experiments with animals.
Study Questions
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1.4 The History of Research on the Brain and Behavior Begins in Antiquity
Le arning Obje ctive s
After reading this section, you should be able to:
1.4.1 Trace the historical point at which the brain was recognized as the control unit for behavior.
1.4.2 Discuss the importance of the Renaissance in better understanding human anatomy.
1.4.3 Explain Descartes’s contribution to early neuroscience and his now-discredited ideas of dualism.
1.4.4 Trace the history of phrenology and the relationship to modern thinking about brain and behavior.
1.4.5 Discuss the difficult question of consciousness and the explosion of neuroscience research.
Only recently have scientists recognized the central role of the brain in controlling behavior. When Egyptian pharaoh
Tutankhamen was mummified (about 1300 BCE ), five important organs were preserved in his tomb: liver, lungs,
stomach, intestines, and heart. All these organs were considered necessary to ensure the pharaoh’s continued
existence in the afterlife. The brain, however, was picked out through the nostrils (Figure 1.11) and thrown away.
Although the Egyptian version of the afterlife entailed considerable struggle, the brain was not considered an asset.
1.11 Brain Removal Kit Ancient Egyptians had little regard for the brain. During mummification, they would use tools
like these to reach through the nostrils to pick out brain pieces and throw them away.
Neither the Hebrew Bible (written from the twelfth to the second century BCE ) nor the New Testament ever
mentions the brain. However, the Bible mentions the heart hundreds of times and makes several references each to
the liver, the stomach, and the bowels as the seats of passion, courage, and pity, respectively. “Teach us … that we
may gain a heart of wisdom” (Psalms 90:12).
The heart is also where Aristotle (about 350 BCE ), the most prominent scientist of ancient Greece, located mental
capacities. We still reflect this ancient notion when we call people kindhearted, openhearted, fainthearted,
hardhearted, or heartless and when we speak of learning by heart. Aristotle considered the brain to be only a
cooling unit to lower the temperature of the hot blood from the heart.
Also about 350 BCE , the Greek physician Herophilus (the “Father of Anatomy”) advanced our knowledge of the
nervous system by dissecting bodies of both people and animals. He traced nerves from muscles and skin into the
spinal cord and noted that each region of the body is connected to separate nerves.
A second-century Greco-Roman physician, Galen, treated the injuries of gladiators. His reports of behavioral
changes caused by injuries to the heads of gladiators drew attention to the brain as the controller of behavior. Galen
advanced the idea that animal spirits—a mysterious fluid—passed along nerves to all regions of the body. But
Galen’s ideas about the anatomy of the human brain were very inaccurate because dissecting humans was illegal in
Rome at that time.
Renaissance scientists began to understand brain anatomy
The eminent Renaissance painter and scientist Leonardo da Vinci (1452–1519) studied the workings of the human
body and laid the foundations of anatomical drawing. He especially pioneered in providing views from different
angles and cross-sectional representations. His artistic renditions of the body included portraits of the nerves in the
arm and the fluid-filled ventricles in the brain (Figure 1.12).
1.12 Leonardo da Vinci’s Changing View of the Brain (A) In an early representation, Leonardo simply copied old
schematic drawings that represented the fluid-filled cerebral ventricles as a linear series of chambers. (B) Later he made a
drawing based on direct observation: after making a cast of the ventricles of an ox brain by pouring melted wax into the
brain and letting it set, he cut away the tissue to reveal the true shape of the ventricles.
Renaissance anatomists emphasized the shape and appearance of the external surfaces of the brain because these
were the parts that were easiest to see when the skull was removed. It was immediately apparent to anyone who
looked that the brain has an extraordinarily complex shape. To Renaissance artists like Michelangelo (1475–1564),
this marvelous structure was God’s greatest gift to humankind. So, in Michelangelo’s painting on the ceiling of the
Sistine Chapel, God seems to ride the form of the human brain when bestowing life to Adam (Meshberger, 1990),
while in another scene God’s throat resembles the base of the brain (Suk and Tamargo, 2010).
In 1633, René Descartes (1596–1650) wrote an influential book (De Homine [On Man]) in which he tried to
explain how the behavior of animals, and to some extent the behavior of humans, could be like the workings of a
machine. Descartes proposed the concept of spinal reflexes and a neural pathway for them (Figure 1.13).
Attempting to relate the mind to the body, Descartes suggested that the two come into contact in the pineal gland,
located within the brain. He suggested the pineal gland for this role because (1) whereas most brain structures are
double, located symmetrically in the two hemispheres, the pineal gland is single, like consciousness, and (2) he
believed, erroneously, that the pineal gland exists only in humans and not in animals.
1.13 An Early Account of Reflexes In this depiction of an explanation by Descartes, when a person’s toe touches fire,
the heat causes nervous activity to flow up the nerve to the brain (blue arrows). From there the nervous activity is
“reflected” back down to the leg muscles (red arrows), which contract, pulling the foot away from the fire; the idea of
activity being reflected back is what gave rise to the word reflex. In Descartes’s time, the difference between sensory and
motor nerves had not yet been discovered, nor was it known that nerve fibers normally conduct in only one direction.
Nevertheless, Descartes promoted thinking about bodily processes in scientific terms, and this focus led to steadily more
accurate knowledge and concepts.
As Descartes was preparing to publish his book, he learned that the Catholic Church had forced Galileo to
renounce his teaching that Earth revolves around the sun, threatening to execute him if he did not recant. Fearful that
his own speculations about mind and body could also incur the wrath of the church, Descartes withheld his book
from publication. It did not appear in print until 1662, after his death. Descartes believed that if people were nothing
more than intricate machines, they could have about as much free will as a pocket watch, with no opportunity to
make the moral choices that were so important to the church. He asserted that humans, at least, had a nonmaterial
soul as well as a material body. This notion of dualism spread widely and left other philosophers with the task of
determining how a nonmaterial soul could exert influence over a material body and brain. Mainstream neuroscientists
reject dualism and insist that all the workings of the mind can also, in theory, be understood as purely physical
processes in the material world, specifically in the brain.
1.14 Old and New Phrenology (A) In the early nineteenth century, certain “faculties,” such as skill at mathematics or a
tendency toward aggression, were believed to be directly associated with particular brain regions. Phrenologists used
diagrams like this one to measure bumps on the skull, which they took as an indication of how fully developed each brain
region was in an individual, and hence how fully that person should display particular qualities. (B) Today, technology
enables us to roughly gauge how active different parts of the brain are when a person is performing various tasks (see
Chapter 2). But virtually the entire brain is active during any task, so the localization of function that such studies provide is
really a measure of where peak activity occurs, rather than a suggestion of a single region involved in a particular task. (B
after M. J. Nichols and W. T. Newsome, 1999. Nature 402: C35–C38.)
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DANCE ON STILTS AT THE GIRLS’ UNYAGO, NIUCHI
I see increasing reason to believe that the view formed some time
back as to the origin of the Makonde bush is the correct one. I have
no doubt that it is not a natural product, but the result of human
occupation. Those parts of the high country where man—as a very
slight amount of practice enables the eye to perceive at once—has not
yet penetrated with axe and hoe, are still occupied by a splendid
timber forest quite able to sustain a comparison with our mixed
forests in Germany. But wherever man has once built his hut or tilled
his field, this horrible bush springs up. Every phase of this process
may be seen in the course of a couple of hours’ walk along the main
road. From the bush to right or left, one hears the sound of the axe—
not from one spot only, but from several directions at once. A few
steps further on, we can see what is taking place. The brush has been
cut down and piled up in heaps to the height of a yard or more,
between which the trunks of the large trees stand up like the last
pillars of a magnificent ruined building. These, too, present a
melancholy spectacle: the destructive Makonde have ringed them—
cut a broad strip of bark all round to ensure their dying off—and also
piled up pyramids of brush round them. Father and son, mother and
son-in-law, are chopping away perseveringly in the background—too
busy, almost, to look round at the white stranger, who usually excites
so much interest. If you pass by the same place a week later, the piles
of brushwood have disappeared and a thick layer of ashes has taken
the place of the green forest. The large trees stretch their
smouldering trunks and branches in dumb accusation to heaven—if
they have not already fallen and been more or less reduced to ashes,
perhaps only showing as a white stripe on the dark ground.
This work of destruction is carried out by the Makonde alike on the
virgin forest and on the bush which has sprung up on sites already
cultivated and deserted. In the second case they are saved the trouble
of burning the large trees, these being entirely absent in the
secondary bush.
After burning this piece of forest ground and loosening it with the
hoe, the native sows his corn and plants his vegetables. All over the
country, he goes in for bed-culture, which requires, and, in fact,
receives, the most careful attention. Weeds are nowhere tolerated in
the south of German East Africa. The crops may fail on the plains,
where droughts are frequent, but never on the plateau with its
abundant rains and heavy dews. Its fortunate inhabitants even have
the satisfaction of seeing the proud Wayao and Wamakua working
for them as labourers, driven by hunger to serve where they were
accustomed to rule.
But the light, sandy soil is soon exhausted, and would yield no
harvest the second year if cultivated twice running. This fact has
been familiar to the native for ages; consequently he provides in
time, and, while his crop is growing, prepares the next plot with axe
and firebrand. Next year he plants this with his various crops and
lets the first piece lie fallow. For a short time it remains waste and
desolate; then nature steps in to repair the destruction wrought by
man; a thousand new growths spring out of the exhausted soil, and
even the old stumps put forth fresh shoots. Next year the new growth
is up to one’s knees, and in a few years more it is that terrible,
impenetrable bush, which maintains its position till the black
occupier of the land has made the round of all the available sites and
come back to his starting point.
The Makonde are, body and soul, so to speak, one with this bush.
According to my Yao informants, indeed, their name means nothing
else but “bush people.” Their own tradition says that they have been
settled up here for a very long time, but to my surprise they laid great
stress on an original immigration. Their old homes were in the
south-east, near Mikindani and the mouth of the Rovuma, whence
their peaceful forefathers were driven by the continual raids of the
Sakalavas from Madagascar and the warlike Shirazis[47] of the coast,
to take refuge on the almost inaccessible plateau. I have studied
African ethnology for twenty years, but the fact that changes of
population in this apparently quiet and peaceable corner of the earth
could have been occasioned by outside enterprises taking place on
the high seas, was completely new to me. It is, no doubt, however,
correct.
The charming tribal legend of the Makonde—besides informing us
of other interesting matters—explains why they have to live in the
thickest of the bush and a long way from the edge of the plateau,
instead of making their permanent homes beside the purling brooks
and springs of the low country.
“The place where the tribe originated is Mahuta, on the southern
side of the plateau towards the Rovuma, where of old time there was
nothing but thick bush. Out of this bush came a man who never
washed himself or shaved his head, and who ate and drank but little.
He went out and made a human figure from the wood of a tree
growing in the open country, which he took home to his abode in the
bush and there set it upright. In the night this image came to life and
was a woman. The man and woman went down together to the
Rovuma to wash themselves. Here the woman gave birth to a still-
born child. They left that place and passed over the high land into the
valley of the Mbemkuru, where the woman had another child, which
was also born dead. Then they returned to the high bush country of
Mahuta, where the third child was born, which lived and grew up. In
course of time, the couple had many more children, and called
themselves Wamatanda. These were the ancestral stock of the
Makonde, also called Wamakonde,[48] i.e., aborigines. Their
forefather, the man from the bush, gave his children the command to
bury their dead upright, in memory of the mother of their race who
was cut out of wood and awoke to life when standing upright. He also
warned them against settling in the valleys and near large streams,
for sickness and death dwelt there. They were to make it a rule to
have their huts at least an hour’s walk from the nearest watering-
place; then their children would thrive and escape illness.”
The explanation of the name Makonde given by my informants is
somewhat different from that contained in the above legend, which I
extract from a little book (small, but packed with information), by
Pater Adams, entitled Lindi und sein Hinterland. Otherwise, my
results agree exactly with the statements of the legend. Washing?
Hapana—there is no such thing. Why should they do so? As it is, the
supply of water scarcely suffices for cooking and drinking; other
people do not wash, so why should the Makonde distinguish himself
by such needless eccentricity? As for shaving the head, the short,
woolly crop scarcely needs it,[49] so the second ancestral precept is
likewise easy enough to follow. Beyond this, however, there is
nothing ridiculous in the ancestor’s advice. I have obtained from
various local artists a fairly large number of figures carved in wood,
ranging from fifteen to twenty-three inches in height, and
representing women belonging to the great group of the Mavia,
Makonde, and Matambwe tribes. The carving is remarkably well
done and renders the female type with great accuracy, especially the
keloid ornamentation, to be described later on. As to the object and
meaning of their works the sculptors either could or (more probably)
would tell me nothing, and I was forced to content myself with the
scanty information vouchsafed by one man, who said that the figures
were merely intended to represent the nembo—the artificial
deformations of pelele, ear-discs, and keloids. The legend recorded
by Pater Adams places these figures in a new light. They must surely
be more than mere dolls; and we may even venture to assume that
they are—though the majority of present-day Makonde are probably
unaware of the fact—representations of the tribal ancestress.
The references in the legend to the descent from Mahuta to the
Rovuma, and to a journey across the highlands into the Mbekuru
valley, undoubtedly indicate the previous history of the tribe, the
travels of the ancestral pair typifying the migrations of their
descendants. The descent to the neighbouring Rovuma valley, with
its extraordinary fertility and great abundance of game, is intelligible
at a glance—but the crossing of the Lukuledi depression, the ascent
to the Rondo Plateau and the descent to the Mbemkuru, also lie
within the bounds of probability, for all these districts have exactly
the same character as the extreme south. Now, however, comes a
point of especial interest for our bacteriological age. The primitive
Makonde did not enjoy their lives in the marshy river-valleys.
Disease raged among them, and many died. It was only after they
had returned to their original home near Mahuta, that the health
conditions of these people improved. We are very apt to think of the
African as a stupid person whose ignorance of nature is only equalled
by his fear of it, and who looks on all mishaps as caused by evil
spirits and malignant natural powers. It is much more correct to
assume in this case that the people very early learnt to distinguish
districts infested with malaria from those where it is absent.
This knowledge is crystallized in the
ancestral warning against settling in the
valleys and near the great waters, the
dwelling-places of disease and death. At the
same time, for security against the hostile
Mavia south of the Rovuma, it was enacted
that every settlement must be not less than a
certain distance from the southern edge of the
plateau. Such in fact is their mode of life at the
present day. It is not such a bad one, and
certainly they are both safer and more
comfortable than the Makua, the recent
intruders from the south, who have made USUAL METHOD OF
good their footing on the western edge of the CLOSING HUT-DOOR
plateau, extending over a fairly wide belt of
country. Neither Makua nor Makonde show in their dwellings
anything of the size and comeliness of the Yao houses in the plain,
especially at Masasi, Chingulungulu and Zuza’s. Jumbe Chauro, a
Makonde hamlet not far from Newala, on the road to Mahuta, is the
most important settlement of the tribe I have yet seen, and has fairly
spacious huts. But how slovenly is their construction compared with
the palatial residences of the elephant-hunters living in the plain.
The roofs are still more untidy than in the general run of huts during
the dry season, the walls show here and there the scanty beginnings
or the lamentable remains of the mud plastering, and the interior is a
veritable dog-kennel; dirt, dust and disorder everywhere. A few huts
only show any attempt at division into rooms, and this consists
merely of very roughly-made bamboo partitions. In one point alone
have I noticed any indication of progress—in the method of fastening
the door. Houses all over the south are secured in a simple but
ingenious manner. The door consists of a set of stout pieces of wood
or bamboo, tied with bark-string to two cross-pieces, and moving in
two grooves round one of the door-posts, so as to open inwards. If
the owner wishes to leave home, he takes two logs as thick as a man’s
upper arm and about a yard long. One of these is placed obliquely
against the middle of the door from the inside, so as to form an angle
of from 60° to 75° with the ground. He then places the second piece
horizontally across the first, pressing it downward with all his might.
It is kept in place by two strong posts planted in the ground a few
inches inside the door. This fastening is absolutely safe, but of course
cannot be applied to both doors at once, otherwise how could the
owner leave or enter his house? I have not yet succeeded in finding
out how the back door is fastened.