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No principle is more central to today's psychology, or to this book, than this:

Everything psychological is simultaneously biological. Your every idea, every mood.


every urge is a biological happening. You think, feel, and act with your body. You
relate to the world through your body. (Try laughing, crying, or loving without it.)
Without your body-your genes, your brain, your body chemistry, your appear
ance-you are, indeed, nobody. Although we find it convenient to talk separately of
biological and psychological influences on behavior, we need to remember: To
think, feel, or act without a body would be like running without legs.
Today's science is riveted on the most amazing parts of our body-our brain,
its component neural systems, and their genetic blueprints. The brain's ultimate
challenge? To understand itself. How does our brain organize and communicate
with itself? How does our heredity prewire the brain and our 'experience modify it?
How does the brain process the information we need to shoot a basketball? To de
light in a guitarist's notes? To remember our first kiss?
Our understanding of how the brain enables the mind has come a long way.
The ancient philosopher Plato correctly located the mind in the spherical head
his idea of the perfect form. His student, Aristotle, believed the mind was in the
heart, which pumps warmth and vitality to the body. The heart remains our sym
bol for love, but science has long overtaken philosophy on this issue. It's your
brain, not your heart, that falls in love.
On the timescale of human existence, the last 150 years are but a few ticks of
the clock. Yet that's how recently a scientific understanding of the brain-mind
connection began to emerge. We have come far since the early 1800s, when the
German physician Franz Gall invented phrenology, an ill-fated theory that claimed
bumps on the skull could reveal our mental abilities and our character traits. At
one point, Britain had 29 phrenological societies, and phrenologists traveled
North America giving skull readings (Hunt, 1993).
Phrenology did, however, correctly focus attention on the idea that various
brain regions have particular functions. Within little more than the last century.
we have also realized that the body is composed of cells; that among these are
nerve cells that conduct electricity and "talk" to one another by sending chemical
messages across'"a tiny gap that separates them; that specific brain systems serve
specific functions (though not the functions Gall supposed); and that from the
A wroncileaded tllMly
Despite Initial acceptaace of GaWs
speoolations, bumps Oft the skull tell us
nothing about the bt'aitisundertylng functions.
Nevertheless, some of his assumptions have
held true: DIfferent parts of the brain do
control different aspects of behavior.
J
57
58 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
!! biological psychology a branch
of psychology concerned with the
links between biology and behavior.
(Some biological psychologists
call themselves behavioral
neuroscientists, neuropsychologists,
behavior geneticists, physiological
psychologists. or biopsychologists.)
II neuron a nerve cell; the basic building
block of the nervous system.
11 dendrite the bushy. branching
extensions of a neuron that receive
messages and conduct impulses toward
the cell body.
'" axon the extension of a neuron,
ending in branching terminal fibers,
through which messages pass to other
neurons or to muscles or glands.
myelin [MY-uh-lin) sheath a layer of
fatty tissue segmentally encasing the
fibers of many neurons; enables vastly
greater transmission speed of neural
impulses as the impulse hops from one
node to the next.
action potential a neural impulse; a
brief electrical charge that travels down
an axon. The action potential is
generated by the movement of
positively charged atoms in and out of
channels in the axon's membrane.
information processed in these different brain systems, we construct our experience
of sights and sounds, meanings and memories., pain and passion. You and I are privi
leged to live in a time when discoveries about the interplay of our biology and behav
ior are occurring at an exhilarating pace. .
Throughout this book you will find examples of how our biology underlies our
behavior and mental processes. By studying the links between biological activity and
psychological events, biological psychologists are gaining a better understanding of
sleep and dreams, depression and schizophrenia, hunger and sex, stress and disease.
We therefore begin our study of psychology with a look at its biological roots.
Neural Communication
Preview: The body's information system is built from billions of
interconnected cells called neurons. To fathom our thoughts and actions,
memories and moods, we must first understand how neurons work and
communicate.
W
e are each a system composed of subsystems that are in turn composed of even
smaller subsystems. Tiny cells organize to form such body organs as our stom
ach, heart, and brain. These organs in turn form larger systems for digestion, circula
tion, and information processing. And those systems are part of an even larger
system-you, who in turn are a part of your family, community, and culture. We are
bio-psycho-social systems. To more deeply understand our behavior, we need to study
how these biological, psychological, and social systems work and interact.
In this book we start small and build from the bottom up-from neurons up to
the brain in this chapter, and to the environmental and cultural influences that in
teract with our biology in later chapters. We will also work from the top down, as we
consider how our thinking and emotions influence our brains and our health. At all
levels, psychologists examine how we process information-how we take in informa
tion; how we organize, interpret, and store it; and how we use it.
For scientists, it is a fortunate fact of nature that the information systems of
humans and other animals operate similarly-so similarly, in fact, that you could not
distinguish between small samples of brain tissue from a human and a monkey. This
similarity allows researchers to study relatively simple animals, such as squids and sea
slugs, to discover how our neural systems operate, and to study other mammals'
brains to understand the organization of our own. Though human brains are more
complex, they follow the same principles that govern all of the animal world.
Neurons
Our body's neural information system is complexity built from simplicity. Its building
blocks are neurons, or nerve cells. There are many different types of neurons, but all are
variations on the same theme (AGURE 2.1). Each consists of a cell body and its branching
fibers. The bushy dendrite fibers receive information, and the axon fibers pass it along
to other neurons or to muscles or glands. Unlike the short dendrites, axons are some
times very long, projecting several feet through the body. "Motor" neurons, which con
trol muscles, are the neural system's giant redwoods. A neuron canying orders to a leg
muscle has a cell body and axon roughly on the scale of a basketball attached to a rope
4 miles long. A layer of fatty tissue, called the myelin sheath, insulates the axons of
some neurons and helps speed their impulses. The myelin sheath's importance is evident
in multiple sclerosis, a disease in which the myelin sheath degenerates. The result is a
slowing of all communication to muscles and the eventual loss of muscle control.
CHAPTER 2 NEUROSCIENCE AND BEHAVIOR 59
Terminal branches of axon FIGURE 2.1
(form junctions with other cells) A motor neuron
Dendrites
- ~ - - ; 7 T (receive messages
/' / from other cells)
/.<
Axon
(passes messages away
from the cell body to
other neurons,
muscles, or glands)
Myelin sheath
(covers the axon of some
neurons and helps speed
Neural Impulse neural impulses)
(electrical signal traveling
: '\ . down the axon)
Depending on the type of fiber, the neural impulse travels at speeds ranging
from a sluggish 2 miles per hour to, in some myelinated fibers, a breakneck 200 or
more miles per hour. But even this top speed is 3 million times slower than that of
electricity through a wire. We measure brain activity in milliseconds (thousandths of
a second) and computer activity in nanoseconds (billionths of a second). That helps
to explain why, unlike the nearly instantaneous reactions of a high-speed computer,
your reaction to a sudden event, such as a child darting in front of your car, may take
a quarter-second or more. Your brain is vastly more complex than a computer, but
not faster at executing simple responses.
A neuron fires an impulse when it receives signals from sense receptors stimu
lated by pressure, heat, or light, or when it is stimulated by chemical messages from
neighboring neurons. The impulse, called the action potential, is a brief electrical
charge that travels down the axon.
Neurons, like batteries, generate electricity from chemical events. The
chemistry-to-electricity process involves the exchange of electrically charged atoms,
called ions. The fluid interior of a resting axon has an excess of negatively charged
ions, while the fluid outside the axon membrane has more positively charged ions.
This positive-outside/negative-inside state is called the resting potential. Uke a tightly
guarded facility, the axon's surface is very selective about what it allows in. We say the
axon's surface is selectively pmneo.ble. $0, for example. a resting axon has gates that
block positive sodium ions.
When a neuron fires, however, the security parameters change: The fust bit of
the axon opens its gates, rather like manhole covers flipping open, and the positively
charged sodium ions flood through the membrane channel (FIGURE 2.2, page 60),
This depolarizes that part of the axon, causing the axon's next channel to open. and
then the next, like dominoes falling, each one tripping the next During .. RSting
pause (the refractory period. rather like a camera flash pausing to recharge), abe neu
ron pumps the positively charged sodium atoms back outside. Then it can ~ ~
(In myelinated neurons, the action potential speeds up by hopping from one IDjeUn
"sausage" to the next, as shown in Figure 2.1.) The mind boggles when imagininc this
electrochemical process repeating up to 100 or even 1000 times a second. But this is
just the first of many astonishments.
The neuron is a miniature decision-making device that performs some complex
calculations. From hundreds, even thousands of other neurons, it receives signals on
its dendrites and cell body. Some of these signals are exdtatory, somewhat like push
"I SiRg the body electrJc."
Walt Whitman, "Children of Adam," 1855
2. This depolarization produces another action
potential a little farther along the axon. Gates in
this neighboring area now open, and more
positively charged atoms rush in, while the positively
charged atoms in the previous section of axon exit.
and an action potential.
FIGURE 2.2
Action potential
3. As the action potential continues
speedily down the axon, the first
section has now completely recharged.
+++
threshold the level of stimulation
required to trigger a neural impulse.
;- synapse [SIN-aps] the junction
between the axon tip of the sending
neuron and the dendrite or cell body of
the receiving neuron. The tiny gap at
this junction is called the synaptic gap
or cleft.
,i neurotransmitters chemical
messengers that traverse the
synaptic gaps between neurons.
When released by the sending neuron,
neurotransmitters travel across the
synapse and bind to receptor sites on
the receiving neuron, thereby
influencing whether that neuron will
generate a neural impulse.
"All information processing in the brain
involves neurons 'talking to' each other
at synapses."
Neuroscientist Solomon H. Snyder (1984)
ing a neuron's accelerator. Other signals are inhibitory, more like pushing its brake. If
excitatory signals minus inhibitory signals exceed a minimum intensity, called the
threshold, the combined signals trigger an action potential. (Think of it this way: If
the excitatory party animals outvote the inhibitory party poopers, the party's on.) The
action potential transmits down the axon, which branches into junctions with hun
dreds or thousands of other neurons and with the body's muscles and glands.
Increasing the stimulus above the threshold, however, will not increase the ac
tion potential's intensity. The neuron's reaction is an all-Dr-none response; like guns,
neurons either fire or they don't. The strength of the stimulus does not affect the ac
tion potential's speed.
How then do we detect the intensity of a stimulus? How do we distinguish a
gentle touch from a big hult.' A strong stimulus cannot trigger a stronger or faster im
pulse in a neuron-squeezing a trigger harder won't make a bullet go faster. But a
strong stimulus can trigger more neurons to fire, and to fire more often.
How Neurons Communicate
Neurons interweave so intricately that even with a microscope it is hard to see where
one neuron ends and another begins. Scientists once believed that the branching
axon of one cell fused with the dendrites of another in an uninterrupted fabric. Then
a Spanish anatomist, Santiago Ramon y Cajal (1852-1934), described gaps between
individual nerve cells and concluded that the individual neurons must function as
independent agents within the nervous system. At the same time, the British physiol
ogist Sir Charles Sherrington (1857-1952) noticed that neural impulses were taking
an unexpectedly long time to travel a neural pathway. Sherrington inferred there
must be a brief interruption in the transmission.
We now know that the axon terminal of one neuron is in fact separated from
the receiving neuron by a gap less than a millionth of an inch wide. Sherrington
called this junction the synapse, and the gap is called the synaptic gap or cleft. To
Cajal, these near-unions of neurons-"protoplasmic kisses," he called them-were
another of nature's marvels. How do the neurons execute the protoplasmic kiss? How
Cell body end of axon
1. Neuron stimulation causes a brief
change in electrical charge. If strong
enough, this produces depolarization
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 61
FIGURE 2.3
HoW neurons communicate
Sending neuron
1. Electrical impulses (action potentials) travel
from a neuron's dendrites, down the axon,
to the axon terminal.
i
\
\
.
.
/Sendin
g
neuron
2. When an action potential
3 The sending neuron normally reabsorbs
reaches an axon terminal,
excess neurotransmitter molecules,
it stimulates the release of
a process called reuptake.
neurotransmitter molecules
from sacs called vesicles.
These molecules cross the
synaptic gap and bind to
receptor sites on the
receiving neuron. This
allows electrically charged
.atoms (not pictured here)
to enter the receiving
neuron and excite or inhibit
Receptor sites on
a new action potential
Neurotransmitter
receiving neuron
does it cross the tiny synaptic gap? The answer, it turns out, is one of the important
scientific discoveries of our age.
When tbe action potential reaches the knoblike terminals at an axon's end, it
triggers the release of chefnical messengers, called neurotransmitters (FIGVRE 2.3).
Within 1!10,OOOth of a second. the neurotransmitter molecules cross the synaptic
gap and bind to receptor sites on the receiving neuron-as precisely as a key fits a
lock. For an instant, the neurotransmitter unlocks tiny channels at the receiving site.
This allows ions to enter the receiving neuron, thereby either exciting or inhibiting its
readiness to fire. Excess neurotransmitters are reabsorbed by the sending neuron in a
process called reuptake. Many drugs increase the availability of selected neurotrans
mitters by blocking their reuptake.
Most neurons have a resting rate of random firing, which either increases or de
. creases with input from other neurons and from chemicals that affect their sensitiv
ity. As we noted earlier, the neuron is democratic If it receives many more crdtatoxy
. than inhibitory messages, the c:dJ fires often. More electrical impulses flash down its
axon, releasing more packets of neurotransmitte.ts. which diffuseacross their synap
tic gaps to other neurons.
How Neurotransmitters Influence Us
Researchers have discovered dozens of different neurotransmitters. That raises ques
tions: Are certain neurotransmitters found only in specific places? What are their ef
fects? Can we boost or diminish these effects through drugs or diet? Could such
changes affect our moods, memories, or mental abilities?
"When it comes to the brain, if lJOU
want to see the action, follow the neu
fOtransmitters. "
Neuroscientist floyd Bloom (1993)
62 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
FIGURE 2.4
Neurotransmitter pathways
Each of the brain's differing chemical
messengers has designated pathways where
it operates, as shown here for dopamine and
serotonin (Carter, 1998).
"You are lJour synapses."
Joseph Le Doux. The Synaptic Self, 2002
Molecules and muscles
When your body moves, a flood of
acetylcholine molecules triggers the muscle
action.
Dopamine pathways Serotonin pathways
Later chapters explain the role of neurotransmitters in depression and eupho
ria, hunger and thinking, addictions and therapy. For now, let's glimpse how neuro
transmitters influence our motions and our emotions. We now know that a
particular neural pathway in the brain may use only one or two neurotransmitters
(FIGURE 2.4), and that particular neurotransmitters may have particular effects on be
havior and emotions. Table 2.1 offers examples of how these chemicals floating in
our brains influence our thoughts and emotions.
Acetylcholine (ACh) is one of the best-understood neurotransmitters. In ad
dition to its vital role in learning and memory, ACh is the messenger at every junc
tion between a motor neuron and skeletal muscle. Using powerful electron
microscopes, neurobiologists can magnify thinly sliced specimens of tissue enough to
see the sacs that store and release ACh molecules. When ACh is released to our mus
cle cells, the muscle contracts.
If the transmission of ACh is blocked, our muscles cannot contract. Curare, a
poison that certain South American Indians have put on the tips of their hunting
darts, occupies and blocks ACh receptor sites, leaving the neurotransmitter unable to
TABLE 2.1
SOME NEUROTRANSMlnERS AND THEIR FUNCTIONS
Neurotransmitter Function Examples of Malfunctions
Acetylcholine (ACh)
Dopamine
Serotonin
Norepinephrine
GABA (gamma
aminobutyric acid)
Glutamate
Enables muscle action,
learning. and memory
Influences movement, learn
ing, attention, and emotion
Affects mood, hunger. sleep,
and arousal
Helps control alertness and
arousal
A major inhibitory neuro
transmitter
A major excitatory neuro
transmitter; involved in
memory
Undersupply. as ACh-producing neurons
deteriorate, marks Alzheimer's disease
Excess dopamine receptor activity linked to
schizophrenia; starved of dopamine, the
brain produces the tremors and decreased
mobility of Parkinson's disease
Undersupply linked to depreSSion; Prozac
and some other antidepressant drugs raise
serotonin levels
Undersupply can depress mood
Undersupply linked to seizures, tremors,
and insomnia
Oversupply can overstimulate brain, pro
ducing migraines or seizures (which is why
some people avoid MSG, monosodium glu
tamate. in food)
-
CHAPTER 2
affect the muscles. Struck by one of these darts, an animal becomes paralyzed. Botu
lin, a poison that can form in improperly canned food, causes paralysis by blocking
ACh release from the sending neuron. (Injections of botulin-Botox-smooth wrin
kles by paralyzing the underlying facial muscles.) By contrast, the venom of the black
widow spider causes a synaptic flood of ACh. The result? Violent muscle contractions,
convulsions, and possible death.
The Endorphins
An exciting discovery about neurotransmitters occurred when Candace Pert and
Solomon Snyder (1973) attached a radioactive tracer to morphine, allowing them to
see exactly where in an animal's brain it was taken up. Pert and Snyder discovered
that the morphine, an opiate drug that elevates mood and eases pain, was taken up
by receptors in areas linked with mood and pain sensations.
It was hard to imagine why the brain would contain these "opiate receptors"
unless it had its own naturally occurring opiates. Why would the brain have a chemi
cal lock, unless it also had a corresponding key? Researchers soon confirmed that the
brain does indeed contain several types of neurotransmitter molecules similar to
morphine. Named endorphins (short for endogenous [produced within] morphine),
these natural opiates are released in response to pain and vigorous exercise. They may
therefore help explain all sorts of good feelings, such as the "runner's high." the
painkilling effects of acupuncture, and the indifference to pain in some severely in
jured people, such as David livingstone reported in his 1857 Missionary Travels:
I heard a shout. Starting, and looking half round, I saw the lion just in the act of
springing upon me. I was upon a little height, he caught my shoulder as he
sprang, and we both came to the ground below together. Growling horribly dose
to my ear, he shook me as a terrier does a rat. The shock produced a stupor simi
lar to that which seems to be felt by a mouse after the first shake of the cat. It
caused a sort of dreaminess in which there was no sense of pain nor feeling of
terror, though [I was] quite conscious of all that was happening. ... This peculiar
state is probably produced in all animals killed by the carnivora; and if so, is a
merciful provision by our benevolent Creator for lessening the pain of death.
How Drugs and Other Chemicals Alter Neurotransmission
If indeed the endorphins lessen pain and boost mood, why not flood the brain with ar
tificial opiates, thereby intensifying the brain's own "feel-good" chemistry? Oneprob
lem is that when flooded with opiate drugs such as heroin and morphine, the brain
may stop producing its own natural opiates. When the drug is withdrawn, the brain
may then be deprived of any form of opiate. for a drug addict, the result is agony that
persists until the brain resumes production of its natural opiates or receives more artifi
cial opiates. As we will see in later chapters, mood-altering drugs, from alcoholm nico
tine to heroin, share a common effect They trigger unpleasant, lingering aftereffcd:s.
For suppressing the body's own neurotransmitter prodUction, nature charges a pJic.e.
Various drugs affect communication at the ~ p s e . often by either excitin,cor in
hibiting neurons' firing. Agonists excite. An agonist can be a drug moh::ule that issimilar
enough to the neurotransmitter to mimic its efmcts or that blocks a neurofJ'aJ\SlU'O'S
reuptake (FIGURE 2.5, page 64). Some opiate drugs. for example. produce a t:empor.uy
"high" by amplifying normal sensations of arousal or pleasure. Antqgonists inbtit.. An
antagonist can be a drug molecule that inhibits a neurotransmit:teis release. Or itmay
be enough like the natural neurotransmitter to occupy its receptor sire and block its ef
fect but not similar enough to stimulate the receptor (rather like Comgn coins thJt fit
into, but won't operate, a soda or candy machine). As noted earlier, curare poison causes
paralysis by blocking acetylcholine receptors that produce muscle movement
NEUROSCIENCE AND BEHAVIOR 63
Physician Lewis Thomas. on the
endorphins:
"There it is, a biologically universal
act of mercy. I cannot explain it,
except to say that I would have put it
in had I been around at the very
beginning, sitting as a member of a
planning committee."
The Youngest Science. 1983
II acetylchoiine[ah-seat-el-KO-leen]
(ACh) a neurotransmitter that, among
its functions, triggers muscle
contraction
endorphJns[en-OOR-fins] "morphine
within"- natural, opiatelike
neurotransmitters linked to pain
control and to pleasure.
64 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
FIGURE 2.5
Agonists and antagonists
-Sending
\ neuron
} ,
Vesicles
containing
\
\ Action
neurotrans
\ potential
mitters
Synaptic .7.f' \
gap \. ." . .' .: i Neurotransmitter

"'"'"
Neurotransmitters carry a message from a
sending neuron across a synapse to receptor
sites on a receiving neuron.
Neurotransmitter molecule
Receiving cell
membrane
I
This neurotransmitter molecule
has a molecular structure that
precisely fits the receptor site
on the receiving neuron, much
as a key fits a lock. Receptor site on
receiving neuron
This agonist molecule excites. It
is similar enough in structure to
the neurotransmitter molecule
that it mimics its effects on the
receiving neuron. Morphine, for
instance, mimics the action of
endorphins by stimulating
receptors in brain areas involved
in mood and pain sensations.
Antagonist
blocks
This antagonist molecule
neurotransmitter
inhibits. It has a structure
similar enough to the
neurotransmitter to occupy its
receptor site and block its
action, but not similar enough to
stimulate the receptor. Curare
poisoning paralyzes its victims
by blocking ACh receptors
involved in muscle movement.
(a)
(b)
(c)
Neurotransmitter research is leading to new therapeutic drugs such as those
used to alleviate depression and schizophrenia. But designing a drug can be harder
than it sounds. A blood-brain barrier enables the brain to fence out unwanted chemi
cals circulating in the blood, and some .l:hemicals don't have the right shape to slither
through this barrier. Scientists know, for example, that the tremors of Parkinson's
disease result from the death of nerve cells that produce dopamine. Giving the pa
tient dopamine doesn't help, because dopamine cannot cross the blood-brain barrier.
But L-dopa, a raw material the brain can convert to dopamine, can sneak through,
enabling many patients to regain better muscular control.
________
Neural Communication
The body's circuitry, the nervous system, consists of billions of individual cells called neu
rons. A neuron receives signals from other neurons through its branching dendrites and cell
body, combines these signals in the cell body, and transmits an electrical impulse (the
action potential) down its axon. When these signals reach end of the axon, they stimu
late the release of chemical messengers called neurotransmitters. These molecules pass
on their excitatory or inhibitory messages as they traverse the synaptic gap between neu
rons and combine with receptor sites on neighboring neurons. Researchers are studying
neurotransmitters to discern their role in behavior and emotion. Some drugs (agonists) ex
-
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 65
cite by mimicking particular neurotransmitters or blocking their reuptake; others (antago
nervous system the bOdy's speedy,
nistS) inhibit by blocking neurotransmitters. electrochemical communication system,
consisting of all the nerve cells ofthe
CHECK YOURSELF: How do neurons communicate with one another?
peripheral and central nervous systems.
ASK YOURSELF: Can you recall a time when the endorphin response may have protected
II! central nervous system (eNS) the
you from feeling extreme pain? brain and spinal cord.
Answers to the Check Yourself questions can be found in tile yeUow appendix at tile end of tile book.
peripheral nervoussystem (PHS) the
sensory and motor neurons that
connect the central nervous system
(CNS) to the rest of the body .
nerves neural "cables" containing
many axons. These bundled axons,
The Nervous System which are part of the peripheral
nervous system, connect the central
Preview: To live is to take in information from the world and the body's
nervous system with muscles, glands,
tissues, to make decisions, and to send back information and orders to the and sense organs.
body's tissues. Neurons are the elementary components of our nervous
<!; sensory neurons neurons that carry
system. our body's speedy electrochemical information system.
incoming information from the sense
receptors to the central nervous system.
N
eurons communicating with other neurons form our body's primary information
III interneurons central nervous system
system, the nervous system (FIGURE 2.6). The brain and spinal cord form the
neurons that internally communicate
central nervous system (eNS). The peripheral nervous system (PNS) links the
and intervene between the sensory
central nervous system with the body's sense receptors, muscles, and glands. The sen inputs and motor outputs.
sory and motor axons carrying this PNS information are bundled into the electrical
If motor neurons neurons that carry
cables that we know as nerves. The optic nerve, for example, bundles a million axon
outgoing information from the central
fibers into a single cable carrying the information that each eye sends to the brain nervous system to the muscles and
glands.
(Mason & Kandel, 1991).
Information travels in the nervous system through three types of neurons. The
sensory neurons send information from the body's tissues and sensory organs in
ward to the brain and spinal cord, which process the information. This processing in
volves a second class of neurons, the central nervous system's own interneurons,
which enable its internal communication. The central nervous system then sends in
structions out to the body's tissues via the motor neurons. Our complexity, though,
resides mostly in ouI. interneuron systems. Our nervous system has a few million sen
sory neurons, a few million motor neurons, but billions and billions of interneurons. ..
FIGURE 2.6
The functional divisions of the human
nervous system

AutOlllllllk (controls
self-repbted actlon Gf
intemalGfP11Sand glands)
I
Somatic......
voluntaryaGlWlllents of
skeletal1lll5des)
66 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
somatic nervous system the division
of the periphe ral nervous system that
controls the body's skeletal muscles.
Also called the skeletal nervous system.
, autonomic [aw-tuhNAHMikJ
nervous system the part of the
peripheral nervous system that
controls the glands and the muscles of
the internal organs (such as the heart).
Its sympathetic division arouses; its
parasympathetic division calms.
. sympathetic nervous system the
division of the autonomic nervous
system that arouses the body, mobilizing
its energy in stressful situations.
parasympathetic nervous system the
division of the autonomic nervous
system that calms the body, conserving
its energy.
reflex a simple, automatic, inborn
response to a sensory stimulus, such as
the knee-jerk response.
FIGURE 2.7
The dual functions of the a.lonomlc
nervous system
The autonomic nervous system controls the
more autonomous (or self-regulating) internal
functions. Its sympathetic division arouses
and expends energy. Its parasympathetic
division calms and conserves energy. allowing
routine maintenance activity. For example.
sympathetic stimulation accelerates
heartbeat, whereas parasympathetic
stimulation slows it.
The Peripheral Nervous System
Our peripheral nervous system has two components-somatic and autonomic. The
somatic nervous system controls the movements of our skeletal muscles. As you
reach the bottom of this page, the somatic nervous system will report to your brain
the current state of your skeletal muscles and carry instructions back, triggering your
hand to turn the page.
Our autonomic nervous system controls the glands and the muscles
of our internal organs. Like an automatic pilot, it may sometimes be consciously
overridden. But usually it operates on its own (autonomously) to influence
our internal functioning, including our heartbeat, digestion, and glandular activity .
The autonomic nervous system is a dual system (FIGURE 2.7). The sympa
thetic nervous system arouses us for defensive action. If something alarms or en
rages you, the sympathetic system will accelerate your heartbeat, slow your
digestion, raise your blood sugar, dilate your arteries, and cool you with perspira
tion, making you alert and ready for action. (Lie detection machines measure such
stress responses, which mayor may not accompany lies.) When the stress subsides,
the parasympathetic nervous system produces opposite effects. It conserves en
ergy as it calms you by decreasing your heartbeat, lowering your blood sugar, and
so forth. In everyday situations, the sympathetic and parasympathetic nervous sys
tems work together to keep us in a steady internal state.
SYMPATHETIC PARASYMPATHETIC
NERVOUS SYSTEM NERVOUS SYSTEM
(arousing) Brain (calming)
Liver
Contracts
bladder
Allows blood
flow to
sex organs
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 67
The Central Nervous S y s t ~ m
From the simplicity of neurons "talking" to other neurons arises the complexity
of the central nervous system that enables our humanity-our thinking, feeling, and
acting. Tens of billions of neurons, each communicating with thousands of other
neurons, yield an ever-changing wiring diagram that dwarfs a powerful computer.
One of the great remaining scientific mysteries is how this neural machinery orga
niles itself into complex circuits capable of learning, feeling, and thinking.
Spinal Cord and Brain
The central nervous system's spinal cord is an information highway connecting the
peripheral nervous system to the brain. Ascending neural tracts send up sensory in
formation, and descending tracts send back motor-control information.
The neural pathways governing our reflexes, our automatic responses to stim
uli, illustrate the spinal cord's work. A simple spinal reflex pathway is composed of a
single sensory neuron and a single motor neuron. These often communicate through
an interneuron. The knee-jerk response, for example, involves one such simple path
way; a headless warm body could do it.
Another such pathway enables the pain reflex (FIGURE 2.8). When your fingers
touch a flame, neural activity excited by the heat travels via sensory neurons to in
terneurons in your spinal cord. These interneurons respond by activating motor neu
rons to the muscles in your arm. That's why it feels as if your hand jerks away not by
your choice, but on its own.
Because the simple pain reflex pathway runs through the spinal cord and out,
your hand jerks from the candle's flame before your brain receives and responds to the
information that causes you to feel pain. Information travels to and from the brain
by way of the spinal cord. Were the top of your spinal cord severed, you would not
feel such pain. Nor would you feel pleasure. Your brain would literally be out of touch
with your body. You would lose all sensation and voluntary movement in body re
gions whose sensory and motor neurons connect with the spinal cord below its point
of injury. You would exhibit the knee-jerk without feeling the tap. With the severing
of the brain center that keeps the brakes on erections, men paralyzed below the waist
are easily capable of an erection (a simple reflex) if their genitals are stimulated
Brain
Sensory neuron
(incoming information)
1. In this simple hand-withdrawal reflex, information

is carried from skin receptors along a sensory neuron


to the spinal cord (shown by the red arrows).
From here it is passed via interneurons to motor
neurons that lead to muscles in the hand and ana
(blue arrows).
"If the nervous system be cut off
between the brain and other parts, the
experiences of those other parts are
nonexistent for the mind. The eye is
blind, the ear deaf, the hand insensible
and motionless."
William lames, Principles of Psychology, 1890
2. Because thisRftex involves only the spinal cord.
the hand jerks away from the candle flame even
before information about the event has reached the FIGURE 2.8
brain, causing lhr experience of pain.
A simple reflex
S8 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
"The body is made up of millions and millions
of crumbs."
FIGURE 2.9
A simplified neural network
Neurons network with nearby neurons.
Encoded in these networks of inter-relating
neurons is your own enduring identity, your
sense of self that extends across the years.
(Goldstein, 2000). Females similarly paralyzed respond with vaginal lubrication. But,
depending on where and how completely the spinal cord is severed, they may be gen
itally unresponsive to erotic images and have no genital feeling (Kennedy & Over,
1990; Sipski & others, 1999). To produce bodily pain or pleasure, the sensory infor
mation must reach the brain. '
Neural Networks
The other part of your central nervous system, your brain, receives information, in
terprets it, and decides responses. In doing so, the brain functions rather like a com
puting machine. It receives slightly differing images of an object from the two eyes,
computes their difference, and instantly infers how far away the object must be to
project such a difference. As basketball star LeBron James releases his falling-away
jump shot, his agile brain performs an incredible number of instant computations,
adjusting for body position and movement, distance, and angle.
How does James's brain perform such computations? First, each neuron con
nects with thousands of others. To get a feel for the complexity of these interconnec
tions, consider that you could join two 8-studded Lego bricks 24 ways, and six bricks
nearly 103 million ways. With some 30 billion neurons, each having roughly 10,000
contacts with other neurons, we end up with something like 300 trillion cortical
synaptic connections. A grain-of-sand-sized speck of your brain contains 100,000
neurons and one billion "talking" synapses (Ramachandran & Blakeslee, 1998).
Being human takes a lot of nerve.
Neurons cluster into work groups called neural networks. To understand why
neurons tend to connect with nearby neurons, Stephen Kosslyn and Olivier Koenig
(1992, p. 12) invite us to "think about why cities exist; why don't people distribute
themselves more evenly across the countryside?" Like people networking with people,
neurons network with nearby neurons with which they can have short, fast connec
tions. As in FIGURE 2.9, the cells in each layer of a neural network connect with vari
ous cells in the next layer. Learning occurs as feedback strengthens connections that
produce certain results. Piano practice builds neural connections. New computer
models simulate neural networks, complete with excitatory and inhibitory connec
tions that gain strength with experience-and mimic the brain's capacity for learning.
Of course, the system inside each of us is more complicated than the networks
depicted in Figure 2.9. In our brains, one neural network is interconnected with other
networks that do different things. There are no arrows to tell us where one network
ends and the next begins; what distinguishes them is their specific functions. Each is
a subnetwork, contributing its little bit of information to the whole information
processing system that we call the brain.
Neurons in the brain connect
with one another to form networks
Inputs --........ Outputs
The brain learns by modifying
certain connections in response to feedback
-
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 69
(REVIEW AND
... neural networks interconnected
neural cells. With networks
can learn, as feedback strengthens or
The Nervous System
inhibits connections that produce
The central nervous system's neurons in the brain and spinal cord communicate with the certain results. Computer simulations
of neural networks show analogous
peripheral nervous system's sensory and motor neurons. The peripheral nervous system
learning.
has two main divisions. The somatic nervous system directs voluntary movements and re
flexes. The autonomic nervous system, through its sympathetic and parasympathetic divi
sions, controls our involuntary muscles and glands. like people clustering into
neighborhoods, neurons cluster into working networks.
CHECK YOURSELF: How does information flow through your nervous system as you pick
up a fork? Can you summarize this process?
ASK YOURSELF: Does our nervous system's design-with its synaptic gaps that chemical
messenger molecules cross in an imperceptibly brief instant-surprise you? Would you
have designed yourself differently?
Answers to the Check Yourself questions can be found in the yellow appendix at the end of the book.
The Brain
Preview: The known universe's most amazing organ is being probed and
mapped by a new generation of neural cartographers. lower-level brain
networks sustain basic life functions and enable memory, emotions, and
basic drives. Higher up, neural networks within the cerebral hemispheres
form specialized work teams that enable our perceiving, thinking, and
speaking.
I
na jar on a display shelf in Cornell University's psychology department resides the
well-preserved brain of Edward Bradford Titchener, a great turn-of-the-century ex
perimental psychologist and proponent of the study of consciousness. Imagine your
self gazing at that wrinkled mass of grayish tissue, wondering if in any sense
Titchener is still in there. t
You might answer that, without the living whir of electrochemical activity, there
could be nothing of Titchener in his preserved brai.fi..Consider then an experiment
about which the inquisitive Titchener himself might have daydreamed. Imagine that
just moments before his death, someone removed TItchener's brain from his body and
kept it alive by pumping enriched blood through it as it floated in a tank of cerebral
flUid. Would Titchener now still be in there? Further imagine, to carry our fantasy to
.. its limit, that someone transplanted the still-living brain into the body of a person
with severe brain damage. To whose home should the recovered patient return?
That we can imagine such questions illustrates how convinced we are that live
in our heads. And for good reason: The brain enables the mind: seeing. bearing. remem
bering, thinking, feeling, speaking. dmuning. It is thebrain that setf-Idlectivelyanalyzes
the brain. When we're thinking about our brain. we"1f! thinking with our brain-by ru
ing millions of synapses and releasing billions of neurotransmitter molecules. Indeed.
say neuroscientists, the mind is wfIIIt ,.brain does. But precisely wItt2 and how em the
mind's functions tied to the brainll.tt's first see how scientists such questions.
'Carl Sagan's Broca's Brain (1979) InspIred this questiIIn.
"'You're,certainlya lot less fun since the
operation."
"J am a brain, Watson. The rest of me
is a mere appendix...
Sherlock Holmes, in Arthur Conan Doyle's "The
of the Mazarin Stone"
70 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
- lesion IlEE-zhuhn] tissue destruction.
A brain lesion is a naturally or
experimentally caused destruction of
brain tissue.
'iY electroencephalogram (EEG) an
amplified recording of the waves of
electrical activity that sweep across
the brain's surface. These waves are
measured by electrodes placed on the
scalp.
, CT (computed tomography) scan a
series of x-ray photographs taken
from different angles and combined
by computer into a composite
representation of a slice through the
body. Also called CATscan.
PET (positron emission tomography)
scan a visual display of brain activity
that detects where a radioactive form of
glucose goes while the brain performs
a given task.
MRI (magnetic resonance imaging)
a technique that uses magnetic fields
and radio waves to produce computer
generated images that distinguish
among different types of soft tissue;
allows us to see structures within the
brain.
,
FIGURE 2.10
An electroencephalograph providing
amplified tracings of waves of
electrical activity in the brain
Here it is detecting brain response to sound,
making possible an early evaluation of what
may be a hearing impairment.
The Tools of Discovery
It is exciting to consider how fast and how far the neurosciences have progressed
within a lifetime. For centuries, the human brain lay largely beyond the reach of sci
ence. The neuron was too small to study with the naked eye, its impulses too faint to
record with ordinary electrodes. We were able to feel bumps on the skull, dissect and
analyze lifeless brains, and observe the effects of specific brain diseases and injuries.
But there were no tools high-powered yet gentle enough to explore the living brain.
Now, that has changed. Whether in the interests of science or medicine, we can se
lectively lesion tiny clusters of normal or defective brain cells, leaving their sur
roundings unharmed. We can probe the brain with tiny electrical pulses. We can
snoop on the messages of individual neurons and on the mass action of billions of
neurons. We can see color representations of the brain's energy-consuming activity.
These new tools and techniques have enabled a neuroscientific revolution.
Clinical Observations
The oldest method of studying brain-mind connections is to observe the effects of
brain diseases and injuries. Such observations were first recorded some 5000 years
ago. But it was not until the last two centuries that physicians began systematically
recording the results of damage to specific brain areas. Some noted that damage to
one side of the brain often caused numbness or paralysis on the body's opposite
side, suggesting that the right side of the body is wired to the brain's left side, and
vice versa. Others noticed that damage to the back of the brain disrupted vision,
and that damage to the left-front part of the brain produced speech difficulties.
Gradually, the brain was being mapped. Today, records of more than 1500 brain
injured patients have been assembled by University of Iowa researchers into the
largest-ever brain-damage registry. Their stories provide clues to our own brains'
inner workings.
Manipulating the Brain
Today's scientists, however, do not need to await brain injuries. They can electri
cally, chemically, or magnetically stimulate various parts of the brain and note the
effects. They can also surgically lesion (destroy) tissue in specific brain areas in an
imals. For example, a lesion in one area of the hypothalamus in a rat's brain re
duces eating, causing the rat to starve unless force-fed. A lesion in another area
produces overeating.
Recording the Brain's Electrical Activity
Right now, your mental activity is giving off telltale electrical, metabolic, and magnetic I
signals that would enable neuroscientists to eavesdrop on your brain. The tips of mod- ,I'
ern microelectrodes are so small they can detect the electrical pulse in a
single neuron, making possible some astonishingly precise findings. For
example, we can now detect exactly where the information goes in a
cat's brain, when someone strokes its whisker.
Electrical activity in the brain's billions of neurons sweeps in reg
ular waves across its surface. The electroencephalogram (EEG) is an
amplified tracing of such waves by an instrument called an electroen
cephalograph. Studying an EEG of the gross activity of the whole brain
is like studying the activity of a car engine by listening to the hum of
its motor. However, by presenting a stimulus repeatedly and having a
computer filter out brain activity unrelated to the stimulus, one can ,..
identify the electrical wave evoked by the stimulus (FIGURE 2.10).
j
11
CHAPTER Z
Neuroimaging Techniques
Other new windows into the-brain give us a Supermanlike ability to
see inside the brain without lesioning it. For example, the CT (com
puted tomography) scan examines the brain by taking x-ray pho
tographs that can reveal brain damage. Even more dramatic is the
PET (positron emission tomography) scan (FIGURE 2.11), which
depicts brain activity by showing each brain area's consumption of
its chemical fuel, the sugar glucose (see Figure 2.24, page 83). Active
neurons are glucose hogs. A person is given a temporarily radioactive
form of glucose, and the PET scan locates and measures the radioac
tivity, thereby detecting where this "food for thought" goes. By not
ing these "hot spots," researchers can see which brain areas are most
active as the person performs mathematical calculations, listens to
music, or daydreams.
Another new way of looking into the living brain exploits the fact that the cen
ters of atoms, including those in our brains, spin like tops. In MRI (magnetic reso
nance imaging) scans, the head is put in a strong magnetic field, which aligns the
spinning atoms. Then a brief pulse of radio waves disorients the atoms momentarily.
When the atoms return to their normal spin, they release detectable signals, which
become images of their concentrations. The result is a detailed
picture of the brain's soft tissues. For example, MRI scans reveal enlarged fluid-filled
brain areas in some patients who have schizophrenia, a disabling psychological disor
der (FIGURE 2.12).
Where the brain is especially active, blood goes. By taking pictures less than a
second apart, MRI scans can therefore show the brain lighting up (with increased
oxygen-laden bloodflow) as a research participant performs different mental func
tions. As the person looks at a face, a functional MRI machine detects blood rushing
to the back of the brain, which processes visual information (see Figure 2.20, page
80). Ask the participant to solve a verbal analogy problem, and part of the brain's left
side near the front will light up. Such snapshots of the brain's activity provide new in
sights into how the brain divides its labor. They also reveal when things happen. how
brain areas change with experience, and what brain areas work together. For example,
imaging research reveals that similar brain areas are involved in reading and remem
bering words (Posner & Raichle. 1998).
These new brain-imaging instruments are doing for psychological sciena! what
the microscope did for biology and the telescope for astronotDJ. To be learning about
the neurosciences now is like studying world geography while Magellan was exploring
the seas. The number of explorers grows: Membership in the interdisciplinary Society
for Neuroscience has quadrupled to 30,000 since 1971. Every year the explorers an
nounce new discoveries, which also generate new interpretations of old disaMries.
Here is a small sampling of new revelations:
PET scans show that the b13in areas that light up when people silently
say the name of an animal differ from those that light upwhen they
say the name of a tool (Martin & others, 1996).
MRI scans reveal a larger-tban-average neur.d area in tbcJcftbrain of
musicians who display pedectpitch (Schlaug & others. 1995).
MRI scans of bilingual peopIe's brains reveal that second languages are
represented in the same am;a as the first if learned early. and in differ
ing areas if learned late (lim &: others, 1997).
Functional MRI scans revl'3I that, during a myming task. men's brains
have a distinctively active left brain area, whereas womens brains are
active on both sides (Shayitz It others, 1995).
NEUROSCIENCE AND BEHAVIOR 71
FIGURE 2.11
The PET scan
To obtain a PET scan, researchers inject
volunteers with a low and harmless dose of a
short-lived radioactive sugar. Detectors
around the subject's head pick up the release
of gamma rays from the sugar. which has
concentrated in active brain areas. A
computer then processes and translates
these Signals into a map of the brain at work.
FIGURE 2.12
MRI scan of a heam., individual (left)
and a person with sdllzophrenia
(right)
Note the enlarged ftuid-filled brain region in
the image on the right.
72 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
brainstem the oldest part and central
core of the brain, beginning where the
spinal cord swells as it enters the skull;
the brainstem is responsible for
automatic survival functions.
" medulla [muh-DUL-uh] the base of
the brainstem; controls heartbeat and
breathing.
reticular formation a nerve network in
the brainstem that plays an important
role in controlling arousal.
thalamus [THAL-uh-muss] the brain's
sensory switchboard, located on top of
the brainstemj it directs messages to
the sensory receiving areas in the
cortex and transmits replies to the
cerebellum and medulla.
cerebellum [sehr-uh-BELL-um] the
"little brain" attached to the rear of the
brainstemj it helps coordinate
voluntary movement and balance.
Data from different brain-imaging techniques are appearing faster than anyone
can read and remember them, and researchers are assembling this information in
computer databases. This brain cartography will give all researchers instant access
through electronic networks to PET or MRI studies that reveal activity in a particular
brain area while a person, for example, solves math problems. Clearly, this is the
golden age of brain science.
lower-level Brain Structures
If you could open the skull and look inside, the first thing you might note is the
brain's si2e. In dinosaurs, the brain represents 1/100,000th of the body's weight, in
whales 1/10,000th, in elephants 1I600th, in humans 1/45th. It looks as though a
principle is emerging. But keep on. In mice the brain is 1I40th the body's weight, and
in marmosets 1I25th. So there are exceptions to the rule that the ratio of brain to
body weight provides a clue to a species' intelligence.
More useful clues to an animal's capacities come from the brain's structures. In
primitive vertebrate (backboned) animals, such as sharks, the brain primarily regu
lates basic survival functions: breathing, resting, and feeding. In lower mammals,
such as rodents, a more complex brain enables emotion and greater memory. In ad
vanced mammals, such as humans, the brain processes more information, enabling
us to act with foresight.
Biological history has not greatly altered the basic mechanisms for survival.
Rather, as the English neurologist John Hughlings Jackson recognized a century ago,
species have elaborated new brain systems on top of the old, much as the Earth's land
scape covers the old with the new. Digging down, one discovers the fossil remnants of
the past-brainstem components still performing much as they did for our distant an
cestors. Starting with the brainstem and working up, let's now explore the brain.
The Brainstem
The brain's basement-its oldest and innermost region-is the brainstem. It begins
where the spinal cord enters the skull and swells slightly, forming the medulla. Here
lie the controls for your heartbeat and breathing. If the top of a cat's brainstem is sev
ered from the rest of the brain above it, the animal will still breathe and live-and
even run, climb, and groom (Klemm, 1990). Bufcut off from the brain's higher re
gion, it won't purposefully run or climb to get food.
The brain stem is also the crossover point, where most nerves to and from each
side of the brain connect with the body's opposite side. This peculiar cross-wiring is
but one of many surprises the brain has to offer.
Inside the brainstem, between your ears, lies the reticular ("netlike") forma
tion, a finger-shaped network of neurons that extends from the spinal cord right up
to the thalamus (FIGURE 2.13, page 73). As the spinal cord's sensory input travels up to
the thalamus, some of it travels through the reticular formation, which filters incom
ing stimuli and relays important information to other areas of the brain. Among its
other functions, the reticular formation helps control arousal.
In 1949, Giuseppe Moruzzi and Horace Magoun discovered that electrically
stimulating the reticular formation of a sleeping cat almost instantly produced an
awake, alert animal. Magoun also severed a cat's reticular formation from higher
brain regions without damaging the nearby sensory pathways. The effect? The cat
lapsed into a coma from which it never awakened. Magoun could clap his hands by
the cat's ear, even pinch it; still, no response. The conclusion? The reticular forma
tion is involved in arousal. Later researchers discovered that elsewhere in the brain
stem are neurons whose activity is needed for sleep. (As you will see in Chapter 7,.our
brains are not idle while we sleep.)
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 73
Thalamus
Reticular
formation
FIGURE 2.13
The brainsJem and thalamus
The brainstem. including the
medulla. is an extension of the
spinal cord. The thalamus is
attached to its top. The reticular
formation passes through both
structures.
The Thalamus
Atop the brainstem sits the brain's sensory switchboard, a joined pair of egg-shaped
structures called the thalamus (see Figure 2.13). It receives information from all the
senses except smell and routes it to the higher brain regions that deal with seeing,
hearing, tasting, and touching. Think of the thalamus as being to sensory input what
London is to England's trains: a hub through which traffic passes en route to various
destinations. The thalamus also receives some of the higher brain's replies, which it
then directs to the cerebellum and medulla. The thalamus further appears to coordi
nate t h ~ brain's electrical oscillations, which slow during sleep and speed up to pro
duce waking consciousness (Uinas & Ribary, 2001).
The Cerebellum
Extending from the rear of the
brainstem is the cerebellum,
meaning "little brain," which is
what its two wrinkled hemispheres
rather look like (FIGURE 2.14). As
you will see in Chapter: 9, the
cerebellum enables one type of
nonverbal learning and memory.
However, its most obvious func
tion is coordinating 1'Oluntary
movement. When basdJall great
. Sammy Sosa smacks a home run
with a perfectly timed swing of the bat, ~ his ccn:bel
lum some credit. If you injured your cerebeJ1um. you
would likely have difficulty walking, keeping your bal
ance, or shaking hands. Your movements would be jerky
and exaggerated.
FIGURE 2.14 "
The brain's organ of agility
Hanging at the back of the brain, the
cerebellum coordinates our movements, as
when Mia Hamm directs the ball precisely.
74 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
Note: These lower brain functions all occur without any conscious ef
fort. This illustrates another of our recurring themes: Our brai!! processes
most information outside of our awareness. We are aware of the results of our
brain's labor (say, our current visual experience) but not of how we con
struct the visual image. Likewise, whether we are asleep or awake, our
brainstem manages its life-sustaining functions, freeing our higher brain
regions to dream or to think, talk, or savor a memory.
The limbic System
At the border ("limbus") of the brain's older parts and the cerebral hemi
spheres is a doughnut-shaped neural system, the limbic system (FIGURE
2.15). We will see in Chapter 9 how one limbic system component, the hip
pocampus, processes memory. (If animals or humans lose their hippocam
pus to surgery or injury, they become unable to lay down new memories of
facts and episodes.) For now, let's look at the limbic system's links to emo
tions such as fear and anger, and to basic motives such as those for food
Amygdala Hippocampus
and sex. As we will later see, the limbic system's influence on emotions
FIGURE 2.15
and motives occurs partly through its control of the body's hormones.
The limbic system
THE AMYGDALA In the limbic system, two almond-shaped neural clusters, called the
Limbic structures form a doughnut-shaped
amygdala, influence aggression and fear. In 1939, psychologist Heinrich Kluver and
neural system between the brain's older
neurosurgeon Paul Bucy surgically lesioned the part of a rhesus monkey's brain that
parts and its cerebral hemispheres.
Although part of the hormonal (endocrine) included the amygdala. The result? The normally ill-tempered monkey turned into
system, not the brain, the pituitary gland is the most mellow of creatures. Poke it, pinch it, do virtually anything that normally
controlled by the limbic system's
would trigger a ferocious response, and still the animal remained placid. In later
hypothalamus, just above it.
studies with other wild animals, including the lynx, wolverine, and wild rat, re
searchers noted the same effect. What then might happen if we electrically stimu
lated the amygdala in a normally placid domestic animal such as a cat? Do so in one
spot and the cat prepares to attack, hissing with its back arched, its pupils dilated, its
i! limbic system a doughnut-shaped hair on end. Move the electrode only slightly within the amygdala, cage the cat with a
system of neural structures at the
small mouse, and now it cowers in terror.
border of the brainstem and cerebral
These experiments confirm the amygdala's role in rage and fear, not to mention
hemispheres; associated with emotions
the perception of such emotions and the processing of emotional memories (Ander
such as fear and aggression and drives
son & Phelps, 2000; Poremba & Gabriel, 2001). Still, we must be careful not to think
such as those for food and sex.
of the amygdala as the control center for aggression and fear. The brain is not neatly
Includes the hippocampus, amygdala,
organized into structures that correspond to our categories of behavior. Actually, both
and hypothalamus.
aggressive and fearful behavior involve neural activity in all levels of the brain. Even
. amygdala [uh-MIG-duh-la] two
within the limbic system, stimulating neural structures other than the amygdala can
almond-shaped neural clusters that are
components of the limbic system and
evoke such behavior. If you put a charge to your car's dead battery, you can activate
are linked to emotion. the engine. Yet the battery is merely one link in an integrated system.
Given that amygdala lesions can trans
form violent monkeys into mellow ones, might
such lesions do the same in violent humans?
You might think so. But such psychosurgery has
produced varied results (Mark & Ervin, 1970;
Valenstein, 1986). In a few cases involving pa
tients who suffered brain abnormalities, it re
duced fits of rage, though sometimes with
devastating side effects on the patient's e v e r y ~
day functioning. for ethical reasons, and be,;
cause of the uncertainties involved, drastit
psychosurgery is rarely used. Perhaps, though,
as we learn more about how the brain controls
behavior, we will learn to alleviate brain disor
ders without creating new ones.
Hypothalamus
The amygdala
...
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 75
THE HYPOTHALAMUS Another of the limbic system's fascinating structures lies just
below (hypo) the thalamus, and so is called the hypothalamus. Neuroscientists, by
lesioning or stimulating different areas in the hypothalamus, have isolated within it
neural networks that perform specific maintenance duties for the body. Some
neural clusters influence hunger; still others regulate thirst, body temperature, and
sexual behavior.
The hypothalamus both monitors blood chemistry and takes orders from other
parts of the brain. For example, thinking about sex (in your brain's cerebral cortex) can
stimulate your hypothalamus to secrete hormones. Through these hormones, the hypo
thalamus controls the adjacent "master gland," the pituitary (see Figure 2.15, page 74),
which in tum influences hormone release by other glands, which the hypothalamus
monitors. (Note the interplay between the nervous and hormone systems: The brain in
fluences the hormone system, which in tum influences the brain.) The powerful little
hypothalamus also exerts control by triggering autonomic nervous system activity.
The story of a remarkable discovery about the hypothalamus illustrates
how progress in scientific research often occurs-when curious, open-minded in
vestigators make an unexpected observation. Two young McGill University neo
ropsychologists, James Olds and Peter Milner (1954), were trying to implant
electrodes in the reticular formations of white rats when they made a magnificent
mistake. In one rat, ttiey incorrectly placed an electrode in what was later discov
ered to be a region of the hypothalamus (Olds, 1975). Curiously, the rat kept re
turning to the place on its tabletop enclosure where it had been stimulated by this
misplaced electrode, as if seeking more stimulation. Upon discovering their mis
take, they alertly recognized that they had stumbled upon a brain center that pro
vides a pleasurable reward.
In a meticulous series of experiments, Olds (1958) went on to locate other
"pleasure centers," as he called them. (What the rats actually experience only they
know. And they aren't telling. TodaYs scientists do not want to attribute human reel
ings to rats, so they refer to reward centers, not "pleasure centers.") When OllIs al
lowed rats to trigger their own stimulation in tbcse areas by pressing a poW. he
noticed that they would sometimes do so at a fevaish pace-up to 7000 t:imrs pel'
hour-until they dropped from exhaustion. Moreover. they would do anythinc1D d.
this stimulation, even cross an dectrifiaf floor that a starving rat would not aoss
even to reach food (FIGURE 2.16, page 76).
Similar reward centers in or near the hypotbaIamus were later discovtRli in
many other species, including goldfish. dolphins. and monkeys. In fact, aninu1re
search has revealed both a general reward system that triggers the release ofthe neu
rotransmitter dopamine and specific centers associated with the pleasures of c3ing,
drinking, and sex. Animals, it seems, come equipped with built-in systems tb3t re
ward activities essential to survival.
Aggression as a brain state
Back arched and fur fluffed, this fierce cat is
ready to attack. Electrical stimulation o{a
cat's amygdala provokes reactions such as
the one shown here, suggesting its role in
emotions like rage. Which division of the
autonomic nervous system is activated by
such stimulation? (See page 76.)
The hypothalamus
This small but important structure, colored
red in this MRI brain scan photograph, helps
keep the body's internal environment in a
steady state by regulating thirst, hunger, and
body temperature. Its activity also influences
experiences of pleasureful reward.
"If you were designing a robot vehicle
to walk into me future and survive, ...
you'd wire it'up so that behavior that
ensured the survival of the :.elf Of the
species-like se)( and eating-would be
naturally reinforcing."
Candace Pert (1986)
hypothalalllOs(hi-po-THAL-uh-mussj
a neural sbudure lying below (hypo) the
thalamus; it directs several maintenance
activities (eating, drinking, body
temperature), helps govern the
endocrine system via the pituitary
gland, and is linked to emotion.
76 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
FIGURE 2.16
Rat with an implanted electrode
With an electrode implanted in a reward
center of its hypothalamus, the rat readily
crosses an electrified grid, accepting the
painful shocks, to press a lever that sends
electrical impulses to its "pleasure centers."
More recent experiments have found new ways of using limbic stimulation in
animals to control their actions. By using brain stimulation to reward rats for turning
left or right (when electrodes stimulate a brain region that makes them feel as if their
left or right whiskers have been touched), Sanjiv Talwar and his colleagues (2002)
were able to train rats that had never been outdoors to navigate natural environ
ments. By pressing buttons on a laptop, the researchers can direct a rat-which carries
a receiver, power source, and
video camera on a backpack-to
Ratbot on a pleasure cruise
turn on cue, climb trees, scurry
When stimulated by remote control,
along branches, and turn around
this rat could be guided to navigate
and come back down, suggesting
across a field and even up a tree.
future applications in search
and-rescue operations.
These are dramatic find
ings. They led people to wonder
whether humans, too, might
have limbic centers for pleasure.
Indeed we do. One neurosur
geon has used electrodes to calm
violent patients. Stimulated pa
tients reported mild pleasure;
The cat on page 75 is aroused via its
sympathetic nervous system.
The people who first dissected and
labeled the brain used the language of
scholars-latin and Greek. Their words
are actually attempts at graphic
description: For example, cortex means
"bark," cerebellum is "little brain," and
thalamus is "inner chamber."
however, unlike Olds' rats, they
were not driven to a frenzy by it (Deutsch, 1972; Hooper & Teresi, 1986). Some re
searchers believe that addictive disorders, such as alcoholism, drug abuse, and
binge eating, may stem from a reward deficiency syndrome-a genetically disposed
deficiency in the natural brain systems for pleasure and well-being that leads peo
ple to crave whatever provides that missing pleasure or relieves negative feelings
(Blum & others, 1996).
The Cerebral Cortex
Your cerebral cortex is an intricate covering of interconnected neural cells that, like
bark on a tree, forms a thin surface layer on your cerebral hemispheres. It is your
body's ultimate control and information-processing center.
With the elaboration of the cerebral cortex, tight g e ~ e t i c controls relax and the
organism's adaptability increases. Frogs and other amphibians have a small cortex
and operate extensively on preprogrammed genetic instructions; the larger cortex of
mammals offers increased capacities for learning and thinking, enabling them to be
more adaptable. What makes us distinctively human mostly arises from the complex
functions of our brain's crown, its cerebral cortex.
d
CHAPTER 2
Structure of the Cortex
If you opened a human skull, exposing the
brain, you would see a wrinkled organ, shaped
somewhat like the meat of an oversized wal
nut. Eighty percent of the brain's weight lies in
the ballooning left and right cerebral hemi
spheres, which are mostly filled with axon
connections between the brain's surface and
its other regions. The thin surface layer of the
cerebral hemispheres is the cerebral cortex, a
sheet of cells that is one-eighth of an inch
thick and contains some 20 to 23 billion nerve cells-an estimate projected by
sampling square-millimeter columns of cortical tissue, 180 of which average
299,052 neurons (de Courten-Myers, 2002). Supporting these billions of nerve
cells are nine times as many glial ceIls-"glue cells" that gUide neural connec
tions, provide nutrients and insulating myelin, and mop up ions and neurotrans
mitters. Some researchers believe that glial cells are more than neural nannies. By
"chatting" with neurons they may also participate in information transmission
and memory (Travis, 1994).
Looking at the brain, the first thing you would notice about the cerebral cortex
is its wrinkled surface, only about one-third of which would be visible. These folds
greatly increase the brain's surface area. If flattened, the brain's surface would be
roughly the size of a large pizza. (To fit a thin pizza crust inside a skull, we would need
to crumple it up.) In rats and other lower mammals, the cortex surface is smoother,
with less neural fabric (FIGURE 2.17).
Each brain hemisphere is divided into four regions, or lobes. Starting at the front
of your brain and going around over the top, there are the frontal lobes (behind
your forehead), the parietal lobes (at the top and to the rear), the occipital lobes
(at the back of your head), and the temporal lobes (just above your ears). These
ChimpalU'el!
Monter

l ,:/
The cetUAlarlex
If flattened, a human ClIIIIie.Iwould OMJr.....four
pages ofthis book. Achimpanzee's woutd QIIIfef one
Rat
page, a monkey's a postc;Md. and a rafs apostage
,JJ

4.
stamp. (From Scientific October1994, P.102.)
NEUROSCIENCE RND BEHRVIOR 77
Einstein's brain
For you and me, as well as for Albert
Einstein (whose brain is shown here), this
small wrinkled organ is far more complex
than the most sophisticated computer. What
you see here is only a portion of the brain's
outer layer. Most of its surface lies hidden
within its convoluted folds.
" cerebral [seh-REE-bruhIJ cortex the
intricate fabric of interconnected neural
cells that covers the cerebral
hemispheres; the body's ultimate control
and information-processing center.
!to glial cells cells in the nervous system
that support. nouris", and protect
neurons.
the portion of the
cerebral cortex lying just behind the
forehead; involved in speaking and
muscle movements and in making
plans and judgments.
oc, parietal [puh-RYE-uh-tuhl) lobes the
portion of the cerebral cortex lying at
the top ofthe head and toward the rear;
includes the seIlSOJ}' cortex.
" oaipilal (ahk-Sl ......... luhl] lobes the
portion-ofthe cerebral cortex lying at
the back of the head; includes the visual
areas, which receiYevisual information
from the opposite visual field.
temporal lobes the portion of the
cerebral cortex lying roughly above the
ears: Includes the auditory areas, each
of which receives auditory information
primarily from the opposite ear.
78 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
FIGURE 2.18
The basic subdivisions of the cortex
Demonstration: Try moving your right
hand in a circular motion, as if polishing a
table. Now start your right foot doing the
same motion synchronized with the hand.
Now reverse the foot motion (but not the
hand). Tough, huh? But easier if you try
moving the left foot opposite to the right
hand. The left and right limbs are
controlled by opposite sides of the brain.
So their opposed activities interfere less
with one another.
lobes are convenient geographic
subdivisions separated by promi
nent folds ("fissures") (FIGUR{
2.18). Each lobe carries out many
Parietal functions, and many functions
lobe
require the interplay of several
lobes.
Occipital
Functions of the Cortex
lobe
More than a century ago, autop
sies of people partially paralyzed
Temporal lobe
or speechless revealed damaged
cortex areas. But this rather crude
evidence did not convince re
searchers that specific parts of the
cortex perform specific functions.
After all, if control of speech and movement were diffused across the cortex, damage
to almost any area might produce the same effect. A television would go dead with its
power cord cut, but we would be deluding ourselves if we thought we had "localized"
the picture in the cord.
This analogy reminds us how easy it is to err when trying to localize brain func
tions. We draw lines around brain areas as in a meat market's beef chart, but complex
activities such as speaking, drawing, and shooting pool involve many brain areas. For
example, our experience of vocal music integrates brain activity in areas involved in
speech and music processing. Mereille Besson and colleagues (1998) discovered this
when recording electrical activity in the brains of French musicians listening to oper
atic solos sung a cappella (unaccompanied). The musicians' brains processed the
lyrics and tunes in separate brain areas en route to their experiencing "the exquisite
unity of vocal music."
MOTOR FUNCTIONS Scientists have, however, localized simpler brain functions. For ex
ample, in 1870, when German physicians Gustav Fritsch and Eduard Hitzig applied
mild electrical stimulation to the cortexes of dogs, they made an important discovery:
They could make different body parts move. The effects were selective: Stimulation
caused movement only when applied to an arch-shaped region at the back of the
frontal lobe, running roughly from ear to ear across the top of the brain. This arch we
now call the motor cortex (FIGURE 2.19). Moreover, when the researchers stimulated
specific parts of this region in the left or right hemisphere, specific body parts moved
on the opposite side of the body.
A half-century ago, neurosurgeons Otfrid Foerster in Germany and Wilder
Penfield in Montreal mapped the motor cortex in hundreds of wide-awake patients.
Before putting the knife to the brain, the surgeons needed to know the possible side
effects of removing different parts of the cortex. They painlessly (the brain has no
sensory receptors) stimulated different cortical areas and noted the body responses.
Like Fritsch and Hitzig, they found that when they stimulated different areas of the
motor cortex at the back of the frontal lobe, different body parts moved. (Kids,
don't try this without parental supervision.) They were now able to map the motor
cortex according to the body parts it controlled (Figure 2.19). Interestingly, those
areas of the body requiring precise control, such as the fingers and mouth, occupied
the greatest amount of cortical space.
Neuroscientist Jose Delgado demonstrated the mechanics of motor behavior.
In one monkey, he evoked a smiling response over 400,000 times. In a human pa- ..
tient, he stimulated a spot on the left motor cortex that triggered the right hand to
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 79
output: Motor cortex
(left hemisphere section
controls the body's right side)
Swallowing
make a fist. Asked to keep the fingers open during the next stimulation, the patient,
whose fingers closed despite his best efforts, remarked, "I guess, Doctor, that your
electricity is stronger than my will" (Delgado, 1969, p. 114). More recently, scien
tists have been able to predict a monkey's arm motion a tenth of a second before it
moves-by repeatedly measuring motor cortex activity preceding specific arm move
ments (Gibbs, 1996).
By Similarly eavesdropping on the brain, could we enable someone-perhaps a
paralyzed person-to move a rebotic limb or command a cursor to wrltee-mail or suIt
the Web? To find out, Brown University brain researchers implanted 100 tiny record
ing electrodes in the motor cortexes of three monkeys (Nicolelis & Chapin, 2002;
Serruya & others, 2002). As the monkeys used a joystick to move a cursor to follow a
moving red target (to gain rewards), the researchers recorded the neural firing pat
tern. Then they let their computer operate the joystick. When a monkey merely
thought about the move, the computer responded by moving the amor with nearly
the same proficiency as the monkey. The dr.unatic result raises hopes for those trying
to enable the human mind to control machines, hands-free.
SENSORY FUNCTIONS If the motor cortex sends messages out to the body, where are
incoming messages received in the corta7 Penfield identified a aJJtical area that
specializes in receiving information from the skin senses and from the movement
of body parts. This area, parallel to the motor cortex and just bebiD4it at the front
of the parietal lobes, we now call the sensory cortes: (Figure 2.19). Stimulate a
point on the top of this band of tissue, and a person may report bdnc touched on
the shoulder; stimulate some point on the side, and the person 11I3Jfcd something
on the face.
The more sensitive a body region. the greater the area of the SCI'ISOIY cortex de
voted to it; your supersensitive lips project to a larger brain area thm do your toes
Input: Sensory cortex
(left hemisphere section receives
input from the body's right side)
Trunk Hip
Knee
leg
Foot
Toes
FIGURE 2.19
Left hemisphere tissue devoted to
each body part in the motor cortex and
the sensory cortex
As you can see from this classic though
inexact representation, the amount of cortex
devoted to a body part is not proportional to
that part's size.. Rather, the brain devotes
more tissue to sensitive areas and to areas
requiring precise control. Thus, the fingers
have a greater representation in the cortex
than does the upper arm. However, the neural
wiring Is complex. Individual muscles are
linked with multiple neural dusters, which in
turn may also Iiak with other muscles.
an area at the rear ofthe
frontal labes that controls voluntary
movemetds,
sensoryc:octer the area at the front of
the parietal lobes that registers and
processes body sensations.
80 CHAPTER 2 NEUROSCIENCE RND BEHRVIOR
FIGURE 2.20
New technology shows the brain in
action
This functional MRI scan shows the visual
cortex-the occipital lobes-activated (color
representation of increased blood flow) as
the subject looks at faces. When the person
stops looking at faces, the region instantly
calms down.
FIGURE 2.21
The visual cortex and auditory cortex
The occipital lobes at the rear of the brain
receive input from the eyes. An auditory area
of the temporal lobes receives information
from the ears.
III association areas a reas of the
cerebral cortex that are not involved in
primary motor or sensory functionsj
rather, they are involved in higher
mental functions such as learning,
remembering, thinking, and speaking.
(Figure 2.19). (That's one reason we
kiss with our lips rather than touch
toes.) Similarly, rats have a large area
of the brain devoted to their whisker
sensations, owls to their hearing sensa
tions, and so forth. If a monkey or a
human loses a finger, the region of the
sensory cortex devoted to receiving
input from that finger branches to re
ceive sensory input from the adjacent
fingers. They then become more sensi
tive (Fox, 1984). MRI scans show that
well-practiced pianists likewise have a
larger-than-usual auditory cortex area
that encodes piano sounds, and that deaf people have an enhanced visual cortex
with greater peripheral vision (Bavelier & others, 2000; Pantev & others, 1998). The
brain is sculpted not only by our genes but also by our experience.
Scientists explored further and identified areas where the cortex receives
input from the other senses. At this moment you are receiving visual information
in the occipital lobes at the very back of your brain (FIGURE 2.20). A bad enough
bash there and you would go blind. Stimulated there, you might see flashes of light
or dashes of color. In a sense, we do have eyes in the back of our head! From your
occipital lobes, the visual information you are now processing goes to other areas
that specialize in tasks such as identifying words, detecting emotions, and recog
nizing faces.
,
Any sound you are now hearing
you processed with the auditory areas
in your temporal lobes (FIGURE 2.21).
Most of this auditory information trav
els a circuitous route from one ear to
the auditory receiving area above your
opposite ear. If you were stimulated
there, you might hear a sound. The
f
sound needn't be reaL Jv'lRl scans of
people with schizophrenia reveal that
,(
auditory areas of the temporal lobe are
Visual
active during auditory hallucinations
. cortex
(Lennox & others, 1999). Even the
phantom ringing sound experienced by
people with hearing loss is-if heard in
one ear-associated with activity in the
temporal lobe on the brain's opposite
side (Muhlnickel & others, 1998).
ASSOCIATION AREAS So far, we have pointed out small areas of the cortex that ei
ther receive sensory information or direct muscular responses. In humans, that
leaves a full three-fourths of the thin wrinkled layer, the cerebral cortex, uncommit
ted to sensory or muscular activity. What then goes on in this vast region of the
brain? Neurons in these association areas (the tan areas in FIGURE 2.22) integrate
information. They associate various sensory inputs with stored memories-a very
important part of thinking.
Electrically probing the association areas doesn't trigger any observable response.
So, unlike the sensory and motor areas, we can't so neatly specify the functions of the
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 81
Motor areas
,/'
Sensory areas
Association areas
Rat
Cat
Chimpanzee
association areas. Their silence seems to be what someone had in mind when formu
lating one of pop psychology's most widespread falsehoods: that we ordinarily use only
10 percent of our brains. This fabrkation-"one of the hardiest weeds in the garden of
psychology," writes Donald McBurney (1996, p. 44)-implies that if we could activate
our whole brain, we would be far smarter than those who drudge along on 10 percent
brain power. But from observing surgically lesioned animals and brain-damaged hu
mans, we know that the association areas are in fact not dormant. (The brain has no
appendix-no apparently purposeless tissue.) Rather, the association areas interpret,
integrate, and act on information processed by the sensory areas.
Our association areas in the frontal lobes enable us to judge, plan, and
process new memories. People with damaged frontal lobes may have intact memo
ries, score high on intelligence tests, and be able to bake a cake-yet be unable to
plan ahead to begin baking the cake for the birthday party. After the sister of famed
neurosurgeon Wilder Penfield suffered frontal lobe damage, she was unable to pre
pare the simplest of meals. She retained her knowledge of recipes, measurements,
and techniques, but she could not decipher the order of steps in preparing the meal
(Kimberg & others, 1998).
Frontal lobe damage also can alter personality, removing a person's inhibitions.
Consider the classic case of railroad worker Phineas Gage. One afternoon in 1848,
Gage, then 25 years old, was packing gunpE)wder into a rock with a tamping iron. A
spark ignited the gunpowder, shooting the rod up through his left cheek and out the
top of his skull, leaving his shish-kebab bed frontal lobes masSively damaged. To every
one's amazement, Gage was immediately able to sit up and speak, and after the wound
healed he returned to work. Although his mental abilities and memories were intact.
his personality was not The affable, soft..spoken Phineas Gage was now irritable. pro
fane, and dishonest. He eventually lost his job and
ended up earning his living as a fairground exhibit
This person, said his friends, was -no longer
Gage." With his frontal lobes ruptured. Gage's
moral compass became disconnected from his be
havior. The same loss of moral compiISS was re
cently discovered to be true of two people who as
young children had experienced fu:mtal lobe dam
age similar to Gage's. Both ofthese individuals re
covered, but they also matured as morally
I
.,.
f
deficient-stealing, lying, and abusin&and neglect
ing their out-of-wedlock chitdrm without remorse
(Dolan, 1999). Although raised in good homes,
they seemingly didn't know right from wrong.
j
j
.,
Human
FIGURE 2.22
Areas of the cortex in four mammals
More intelligent animals have increased
"uncommitted" or association areas of the
cortex. These vast areas of the brain are
responsible for integrating and acting on
information received and processed by
sensory areas.
Phineas Gage reconsidered
Using measurements of his skutt (which was
kept as amedical record) and modern
neuroimagingtechniques, researcher Hanna
Oamasio and her colleagues (1994) have
reconstructed the probable path of the rod
through Gage's brain.
82 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
,aphasia impairment oflanguage,
usually caused by left hemisphere
damage either to Broca's area
(impairing speaking) or to Wernicke's
area (impairing understanding).
Broca's area controls language
expression -an area of the frontal lobe,
usually in the left hemisphere. that
directs the muscle movements involved
in speech.
Ii! Wernicke's area controls language
reception-a brain area involved in
language comprehension and
expression; usually in the left temporal
lobe.
The association areas of the other lobes also perform mental functions. For ex
ample, the parietal lobes, parts of which were large and unusually shaped in Ein
stein's normal-weight brain, are involved in mathematical and spatial reasoning
(Witelson & others, 1999). An area on the underside of the right temporal lobe en
ables us to recognize faces. If a stroke or head injury destroyed this area of your
brain, you would still be able to describe facial features and to recognize someone's
gender and approximate age, yet be strangely unable to identify the person as, say,
Nelson Mandela or your next-door neighbor or even your spouse. But by and large,
complex mental functions such as learning and memory don't reside in anyone
place. There is no one spot in a rat's small association cortex that, when damaged,
will obliterate its ability to learn or remember a maze. Such functions are spread
throughout much of the cortex.
LANGUAGE Complex human abilities, such as language, result from the intricate co
ordination of many brain areas. For example, consider this curious finding: Damage
to anyone of several cortical areas can cause aphasia, an impaired use of language. It
is even more curious that some people with aphasia can speak fluently but cannot
read (despite good vision), while others can comprehend what they read but cannot
speak. Still others can write but not read, read but not write, read numbers but not
letters, or sing but not speak. This all is very puzzling. After all, we think of speaking
and reading, or writing and reading, or singing and speaking as merely different ex
amples of the same general ability. So, how did researchers solve this mystery? Con
sider these clues.
Que 1 In 1865, French physician Paul Broca reported that after damage to a
specific area of the left frontal lobe, later called Broca's area, a person would
struggle to form words while still being able to sing familiar songs and
comprehend speech.
Que 2 In 1874, German investigator Carl Wernicke discovered that after
damage to a specific area of the left temporal lobe (Wernicke's area) people
could speak only meaningless words. Asked to describe a picture that showed two
boys stealing cookies behind a woman's back, one patient responded: "Mother is
away her working her work to get her better, but when she's looking the two boys
looking the other part. She's working another time" (Geschwind, 1979).
Que 3 It was later discovered that reading aloud involves a third brain area.
The angular gyrus receives the visual information from the visual area and recodes
it into the auditory form, which Wernicke's area uses to derive its meaning.
Que 4 Nerve fibers interconnect these brain areas.
Norman Geschwind assembled these clues into an explanation of how we
use language. When you read aloud, the words (1) register in the visual area, (2)
are relayed to the angular gyrus that transforms the words into an auditory code
that is (3) received and understood in the nearby Wernicke's area and (4) sent to
Broca's area, which (5) controls the motor cortex, creating the pronounced word
(fIGURES 2.23 and 2.24). Depending on which link in this chain is damaged, a dif
ferent form of aphasia occurs. Damage to the angular gyrus leaves the person able
to speak and understand but unable to read. Damage to Wernicke's area disrupts
understanding. Damage to Broca's area disrupts speaking. The general prinCiple
bears repeating: Complex abilities result from the intricate coordination of many
brain areas.
Said another way, the brain operates by dividing its mental functions-speaking,
perceiving, thinking, remembering-into subfunctions. Our conscious experience
seems indivisible. Right now you are experiencing a whole visual scene as if your eyes
were video cameras projecting the scene into your brain. Actually, as you will see in
Chapter 5, your brain breaks vision into specialized subtasks, such as discerning
..
CHAPTER 2
5. Motor cortex
(word is pronounced)
2. Angular gyrus
(transforms visual representations
4. Broca's area ---f--I
i into an auditory code)
(controls speech
muscles via
the motor cortex)
1. Visual cortex
(receives written words
as visual stimulation)
\
1
3. Wernicke's area
(interprets auditory code)
color, depth, movement, and form. (After a localized stroke that destroys one of these
neural work teams, people may lose just one aspect of vision, such as the ability to
perceive movement.) These specialized neural networks, each having simultaneously
done its own thing, then feed their information to "higher-level" networks that com
bine the atoms of experience and relay them to progreSSively higher-level association
areas, enabling us to recognize a face as "Grandmother."
The same is true of reading a word: The brain computes the word's form,
sound, and meaning using different neural networks (Posner & Carr, 1992).
Thus fMRI scans show that jokes playing on meaning ("Why don't sharks bite
lawyers? ... Professional courtesy") are processed in a different brain area than
jokes playing on language sounds ("What kind oflights did Noah use on the ark? ...
Flood lights") (Goel & Dolan, 2001). Think about it: What you experience as a
continuous, indivisible stream of perception is actually but the visible tip of the
information-processing iceberg, most of which lies beneath the surface of your con
scious awareness.
To sum up, the mind's subsystems are localized in particular brain regions, yet
the brain acts as a unified whole. Moving your hand; recognizing faces; even perceiv
ing color, motion, and depth-all depend on specific neural networks. Yet complex
functions such as language, learning, and loving involve the coordination of many
brain areas. Both principles-specialization and integration-appear in research on
the two brain hemispheres.
(a) (b)
(c)
Hearing words Seeln, WiInb Speaking words
NEUROSCIENCE AND BEHAVIOR 83
FIGURE 2.23
Specialization and integration in
language
FIGURE 2.24
Brain activity When hearing, seeing,
..nd speaking words
PET scans such as these detect the activity of
different areas of the brain by measuring their
rebtive consumption of a temporarily
radioadive.torm oflhe brain's normal fuel,
&tucose.. This'series ofPET scans shows levels
efincn!ased brain activity in specific areas: (a)
when hearing a word-auditory cortex and
Wemic::te-S are!!; (b) when seeing a word
....cortex and angular gyruS; and (c) when
speaking a word - Broca's area and the motor
cortex. The red blotches show where the brain
Is rapidly consuming glucose.
4 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
plasticity the brain's capacity for
modification, as evident in brain
reorganization following damage
(especially in children) and in
experiments on the effects of
experience on brain development.
Brain Reorganization
Nurture's sculpting of the ever-changing brain is evident in studies of the brain's
plasticity. Most severed neurons will not regenerate (if your spinal cord were sev
ered, you likely would be permanently paralyzed). But neural tissue can reorganize in
response to damage.
In one experiment, neuroscientists severed the neural pathways for incoming
information from a monkey's arm. The area of the sensory cortex that formerly re
ceived this input gradually shifted its function and began to respond when re
searchers touched the animal's face (Pons & others, 1991). Similarly, if a laser beam
damages a spot in a cat's eye, the brain area that received input from that spot will
soon begin responding to stimulation from nearby areas in the cat's eye. Another ex
periment reconfigured the brains of newborn ferrets to feed information from their
eyes to brain regions normally assigned to hearing. Tests later showed the animals
could see lights with their auditory cortex (von Melchner & others, 2000). If a blind
person uses one finger to read Braille, the brain area dedicated to that finger expands
(Barinaga, 1992a). The sense of touch invades the part of the brain that normally
helps people see. PET scans also reveal activation of the visual cortex when blind peo
ple read Braille (Sadato & others, 1996). Among deaf people who communicate with
sign language, it is the temporal lobe area normally dedicated to auditory information
that waits in vain for stimulation. Finally, it looks for other signals to process, such as
those from the visual system.
If a body part is amputated, sensory fibers that terminate on adjacent areas of
the sensory cortex may invade the brain tissue that's no longer receiving sensory
input. As Figure 2.19 on page 79 shows, the hand is between the face and the arm re
gions on the sensory cortex. This explains a mysterious phenomenon: When stroking
the face of someone whose hand had been amputated, V. S. Ramachandran found the
person felt the sensations not only on his face but also on his nonexistent ("phan
tom") fingers. Ditto when stroking the arm, whose sensory fibers had also invaded
the brain area vacated by the hand. Note, too, that the toes region is adjacent to the
genitals. So what do you suppose was the experience of another person, whose lower
leg had been amputated, while having sexual intercourse? "I actually experience my
orgasm in my foot. And there it's much bigger than it used to be because it's no
longer just confined to my genitals" (Ramachandran & Blakeslee, 1998, p. 36).
Thus, the brain may not be as "hard-wired" as once thought. Unlike fixed
computer circuits, brain hardware changes with time. In response to changing
stimulation, the brain can either rewire itself with new synapses or (according to
another theory) select new uses for its prewired circuits (Gazzaniga, 1992; Kolb &
Whishaw, 1998). When one brain area is damaged, other areas may in time reorga
nize and take over some of its functions. If neurons are destroyed, nearby neurons
may partly compensate for the damage by making new connections that replace the
lost ones. These new connections are one way the brain struggles to recover from a
minor stroke.
New evidence suggests that, contrary to long-held belief, adult mice and hu
mans can also generate new brain cells (Kempermann & Gage, 1999; Van Praag &
others, 2002). Moreover, monkey brains have recently been discovered to form thou
sands of new neurons each day. These baby neurons originate deep in the brain and
then migrate to the thinking frontal lobe and form connections with neighboring
neurons (Gould & others, 1999). Such are the brain's ways of partially compensating
for the gradual loss of neurons with age. Master "stem celts" that can develop into
any type of brain cell have also been discovered in the fetal brain. If mass-produced in
a lab and injected into a damaged brain, might neural stem cells turn themselves into
replacements for lost brain cells? Might we someday be able to rebuild damaged
brains much as we reseed damaged lawns? Stay tuned.
CHAPTER 2
Our brains are most plastic
when we are young children
(Kolb, 1989). Children are born
with a surplus of neurons. If an
injury destroys one part of a
child's brain, the brain will com
pensate by putting other surplus
areas to work. Thus, if the speech
areas of an infant's left hemi
sphere are damaged, the right
hemisphere will take over much of
its language function.
As an extreme example of
plasticity, consider a 5-year-old
boy whose severe seizures, caused by a deteriorating left hemisphere, require removing
the entire hemisphere. What hope for the future would such a child have? Is there any
chance he might attend school and lead a normal life, or would he suffer permanent
retardation?
Astonishingly, one such individual was at last report an executive. Half his
skull was filled with nothing but cerebrospinal fluid-functionally it might as well
be sawdust-yet he had scored well above average on intelligence tests, had com
pleted college, and had attended graduate school (Smith & Sugar. 1975; A. Smith,
1987). Although paralyzed on the right side, this man (along with other such cases
of "hemispherectomy") testifies to the brain's extraordinary powers of reorganiza
tion when damaged before it is fully developed. Indeed. one Johns Hopkins med
ical team. reflecting on the 58 child hemispherectomies they have performed.
reports being "awed" by how well children retain their memory, personality, and
humor after removal of either brain hemisphere (Vining & others, 1997).
Our Divided Brains
For more than a century, clinical evidence has shown that the brain's two sides serve
differing functions. Accidents, strokes, and tumorS in the left hemisphere generally
impair reading, writing, speaking, arithmetic reasoning, and understanding. Similar
lesions in the right hemisphere seldom have such dramatic effects.
By 1960 the left hemisphere was well accepted as the "dominant" or "major"
hemisphere, and its silent companion to the right as the "subordinate" or "minor"
hemisphere. The left, verbal hemisphere is rather like the moon's facing side-the one
easiest to observe and study. The other side is there. of course, but backstage. (For
about 1 in 10 people, including one-fourth of all left-handers, speech is processed in
the right hemisphere-see page 89.) But then researchers found that the "minor"
right hemisphere was not so limited after all. The stoIy of this discovery is a fascinat
ing chapter in psychology's histoIy.
Splitting the Brain
In 1961, two Los Angeles neurosurgeons, Philip Vocd and Joseph Bop, speculated
that major epileptk seizures were caused by an amplification of abnormal brain3div
ity that reverberated between the two hemispheres. They therefore wondered wbdher
they could reduce seizures in their patients with uncontrollable epilepsy by cutting
communication between the hemispheres. To do this, Vogel and Bogen knew they
would have to sever the corpus callosum, the wide band of axon fibers connecting
the two hemispheres (FIGURE 2.25, page 86).
NEUROSCIENCE AND BEHAVIOR 85
Brain plasticity
Believe it or not, this is functioning
with only half a brain. Her right hemisphere
was surgically removed to eliminate seizures.
Now neurons in her left hemisphere have
made countless new connections to take over
the tasks once performed by her right
hemisphere.
"You wouldn't want to have a date
with the right hemisphere.
Michael Gazzaniga (2000)
corpus c:.aUosum [KOR-pus kah-lOW
sum] the large band of neural fibers
connecting the two brain hemispheres
and carrying messages between them.
36 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR <
Corpus callosum
FIGURE 2.25
The corpus callosum
This large band of neural fibers connects the
two brain hemispheres. To photograph the
half brain shown at left. the hemispheres
were separated by cutting through the
corpus callosum and lower brain regions. In
the view on the right. a surgeon has cut back
brain tissue to expose the corpus callosum
and bundles of fibers coming out from it.
FIGURE 2.26
The information highway from eye to
brain
Information from the left half of your field of
vision goes to your right hemisphere, and
information from the right half of your visual
field goes to your left hemisphere. which
usually controls speech. (Note. however, that
each eye receives sensory information from
both the right and left visual fields.) The data
received by either hemisphere are quickly
transmitted to the other across the corpus
callosum. In a person with a severed corpus
callosum, this information sharing does not
take place.
The surgeons had reason to believe such
an operation would not be incapacitating.
Psychologists Roger Sperry, Ronald Myers,
and Michael Gazzaniga had divided the
brains of cats and monkeys in this manner
with no serious ill effects. So Vogel and Bogen
operated. The result? The seizures were all but
eliminated and the patients with these split
brains were surprisingly normal. their per
sonalities and intellect hardly affected. Wak
ing from the surgery, one patient even
managed to quip that he had a "splitting
headache" (Gazzaniga, 1967).
Only a decade earlier, neuropsycholo
gist Karl Lashley had jested that maybe the corpus callosum served only "to keep the
hemispheres from sagging." The ingenious experiments of Sperry and Gazzaniga re
vealed that this broad band of more than 200 million nerve fibers, capable of trans
ferring more than a billion bits of information per second between the hemispheres,
has a more significant purpose. Their studies of split-brain people-"the most fasci
nating people on earth"-provided a key to understanding the two hemispheres' com
plementary functions.
As FIGURE 2.26 explains, the peculiar na
Left Right
ture of our visual wiring enabled the researchers
visual field visual field
to send information to the patient's left or right
brain-by having the patient stare at a spot and
then flashing a stimulus to its right or left.
(They could do this with you, too, but in your
intact brain the telltale hemisphere that re
ceived the information would instantly call the
news to its partner across the valley. The split
brain surgery severs the phone cables-the cor
pus callosum-across the valley.) Finally, the
researchers quizzed each hemisphere separately.
In an early experiment, Gazzaniga (1967)
asked split-brain people to stare at a dot as he
flashed HE-ART on a screen (FIGURE 2.27). Thus,
HE appeared in their left visual field (which
transmits to the right brain) and ART in their
right field (which transmits to the left brain).
When he then asked them what they had seen,
the patients said they saw ART and so were star
tled when their left hand (controlled by the right
brain) pointed to HE. Given an opportunity to
express itself, each hemisphere reported only
what it had seen. The left hand intuitively knew
what it could not verbally report.
Similarly, when a picture of a spoon was
flashed to their right hemisphere, the patients
could not say what they. saw. But when asked to
identifywhat they had seen by feeling an assort
ment of hidden objects with their left hand,
they readily selected the spoon. If the experi
Visual area Corpus Visual area
menter said, "Right!" the patient might reply,
ofleft callosum of right
hemisphere hemisphere "What? Right? How could I possibly pick out the
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 87
FIGURE 2.27
Testing the divided brain
When an experimenter
flashes the word HEART
"What word
across the visual field, the
did you see?"
HE
splitbrain person reports
seeing the portion of the
word transmitted to her left
hemisphere. However, if
asked to inalCate with her
or
left hand what she saw, she
points to the portion of the
word transmitted to her right
hemisphere. (From
"Point with Gazzaniga, 1983.)
your left hand
to the word
"look at the dot." Two words separated by a dot
yousaw.
w
are momentarily projected.
right object when I don't know what I saw?" It is, of course, the left hemisphere doing
the talking here, bewildered by what the nonverbal right hemisphere knows.
A few people who have had split-brain surgery have been for a time bothered
by the unruly independence of their left hand, which might unbutton a shirt
while the right hand buttoned it or put grocery store items back on the shelf after
the right hand put them in the cart. It was as if each hemisphere was thinking
"I've half a mind to wear my green (blue) shirt today." One woman, after a mas
sive stroke in her corpus callosum, complained that occasionally her left hand
would fly up to her throat and try to strangle her, requiring her right hand to
push it away (Ramachandran & Blakeslee, 1998). Indeed, said Sperry (1964),
split-brain surgery leaves people "with two separate minds." (Reading these re
ports, I fantasize a split-brain person enjoying a solitary game of "rocks, paper,
scissors"-left versus right hand.)
There are other cases of "alien hand syndrome," a neurological disorder in
which people experience one hand as operating with a mind of its own (rather
like Peter Sellers' Dr. Strange/ave, whose autonomous hand would steer his wheel
chair astray or make a Nazi salute). One alien hand patient's hand would some
times grasp her throat during sleep. Her solution: to sleep with the arm tied.
Another patient said it is as if "someone from the moon" controlled her band.
Some cases involve damage to the motor cortex on the brain side oppo.sik the
hand; most also involve some damage to the corpus callosum (Feinberg. 20(11;
Wilson, 2002). All illustrate one of this book's themes; the unconscious brain
can control our behavior without our conscious effort or will.
When the "two minds" are at odds, the left hemisphere seems to act.as the
brain's press agent, doing mental gymnastics to rationalize reactions it doa Dot
understand. If a patient follows an order sent to the right hemisphere C......).
the interpretive left hemisphere will offer a ready explanation ("I'm gome into
the house to get a Coke"). Thus, Michael Gazzaniga (1988) concludes dut the
conscious left hemisphere is an "interpreter- that instantly constructs thames to
explain our behavior. The unconscious out-of-sight mind that helps run our lives
seems to many psychologists rather like the autopilot that operates a jumhD jet
the conscious mind acts like the pilot who occasionally takes control of the jet. At
other times it acts as the mind's press secretary-the part that reports and inter
prets decisions made behind closed doors (Wegner, 2002; Wilson, 2002).
"Do not let your left hand know what
lJour right hand is doing."
Matthew 6:3
Question: If we flashed a red light to
the right hemisphere of a splitbrain
patient and flashed a green light to the
left hemisphere, would each observe its
own color? Would the person be aware
that the colors differ? What would the
person verbally report seeing?
on page 89.)
split brain a condition in which the
two hemispheres of the brain are
isolated by cutting the connecting
fibers (mainly those of the corpus
callosum) between them.
88 CHAPTER 2 NEUROSCIENCE FIND BEHAVIOR
Which one is happier?
Look at the center of one face, then the other.
Does one appear happier? Most people say
the right face does. Some researchers think
this is because the right hemisphere, which
is skilled in emotion processing, receives
information from the left half of each face
(when looking at its center).
Try this
Joe, a split-brain patient, can simultaneously
draw two different shapes.
These experiments demonstrate that
the right hemisphere understands simple
requests and easily perceives objects. With
a split brain, both hemispheres can com
prehend and follow an instruction to
copy-simultaneously-different figures with
the left and right hand (Franz & others,
2000). But the right hemisphere surpasses
the left at copying drawings, and also at
recognizing faces, perceiving differences, perceiving emotion, and expressing emo
tion through the more expressive left side of the face. Most of the body's paired or
gans-kidneys, lungs, breasts-perform identical functions, providing a backup
system should one side fail. Not so the brain's two halves. They are a biological odd
couple, serving differing functions, each seemingly with a mind of its own.
Studying Hemispheric Differences in the Intact Brain
What about the 99.99+ percent of us with undivided brains? Have scientists
found our hemispheres to be similarly specialized? Yes they have, in several differ
ent types of studies. For example, when a person performs a perceptual task, brain
waves, bloodflow, and glucose consumption reveal increased activity in the right
hemisphere; when a person speaks or calculates, activity increases in the left
hemisphere.
On occasion, hemispheric specialization has been even more dramatically
shown by briefly sedating an entire hemisphere. To check for the locus of language
before surgery, a physician may inject a sedative into the neck artery that feeds blood
to the hemisphere on its side of the body. Before the drug is injected, the patient is
lying down, arms in the air, conversing easily. You can likely predict what happens
when the drug flows into the artery going to the left hemisphere: Within seconds, the
person's right arm falls limp and, if the left hemisphere controls language, the subject
becomes speechless until the drug wears off. When the drug goes into the artery to
the right hemisphere, the left arm falls limp, but the person can still speak. With the
right rather than left hemisphere asleep, the person also has difficulty recognizing an
altered self-image (Keenan, 2001).
Other tests also confirm hemispheric specialization. For example, most people"'"
recognize a picture faster and more accurately when it is flashed to the right hemi
sphere. But they recognize a word faster and more accurately when it is flashed to the
left hemisphere. If a word is flashed to your right hemisphere, perception takes a frac
tion of a second longer-the length of time it takes to send the information through
the corpus callosum to the more verbal left hemisphere.
Which hemisphere would you suppose enables sign language among deaf
people? The right, because of its visual-spatial superiority? Or the left, because of
its preparedness to process language? Studies reveal that, just as hearing people
use the left hemisphere to process speech, deaf people use the left hemisphere to
read signs (Corina & others, 1992; Hickok & others, 2001). A stroke in the left
hemisphere will disrupt a deaf person's signing much as it would disrupt a hear
ing person's speaking. Broca's area is similarly involved in both spoken and signed
speech production (Corina, 1998). To the brain, language is language, whether
spoken or signed.
Although the left hemisphere is adept at making quick, literal interpreta
tions of language, the right hemisphere excels in making subtle inferences (Bee
man & Chiarello, 1998; Beeman & others, 1994; Bowden & Beeman, 1998). If
"primed" with the flashed word foot, the left hemisphere will be especially quick
to then recognize the closely associated word heel. But if primed with foot, cry, and
glass, the right hemisphere will more quickly recognize another word that is dis

CHAPTER 2
tantly related to all three (cut). And if given an insightlike problem
what word goes with high, district. and house?-the right hemisph_ere
has better access to the solution. (The right hemisphere more quickly
than the left recognizes that the solution is school.) As one patient
explained after suffering right-hemisphere stroke damage. "I under
stand words. but I'm missing the subtleties." Thus, the right hemi
sphere helps us modulate our speech to make meaning clear-as
when we ask "What's that in the road ahead?" instead of "What's
that in the road. a head?" (Heller, 1990).
Even babies exhibit asymmetry, by favoring the left side of their
mouth when beginning a smile, and the right side when using baby
language-babbling (FIGURE 2.28) From simply looking at the two
hemispheres, which appear alike to the naked eye, who would suppose that they
contribute so uniquely to the harmony of the whole? Yet a variety of observa
tions-of people with split brains and people with normal brains-converge beauti
fully, leaving little doubt that we have unified brains with specialized parts.
Brain Organization and Handedness
About 90 percent of the human population is right-handed. The remaining 10 per
cent (somewhat more among males, somewhat less among females) is left-handed.
Tests reveal that about 95 percent of right-handers process speech primarily in their
left hemispheres, which tend to be slightly larger (Springer & Deutsch, 1985). Left
handers are more diverse. More than half process speech in the left hemisphere, as
right-handers do. About a quarter process language in the right hemisphere; the other
quarter use both hemispheres more or less equally.
Is HANDEDNESS INHERITED? Judging from the cave drawings, the tools, and the
hand and arm bones of prehistOric humans, this veer to the right occurred long ago
in the development of our species (Corballis, 1989; Steele, 2000). Right-handedness
prevails in all human cultures. Moreover, it appears prior to culture's impact Ultra
sound observations of fetal thumb-sucking reveal that more than 9 in 10 fetuses suck
the right hand's thumb (Hepper & others. 1990). This bias for the right hand is
uniquely human; other individual primates are more eveJ}ly divided between Idties
and righties (though among those most like us-chimps and gorillas-only 35 pm::ent
are lefties). Among both chimps and humans, genetic factors appear to play a role
(Geschwind & others, 2002; Hopkins & others, 2001).
The search is underway for a gene that perhaps codes right-handedness (al
lowing those without it to veer toward either left- or right-handedness). Handed
ness is, however, one of but a few traits that genetically identical twins aren't
especially likely to share (Halpern & Coren, 1990). Nevertheless, observing 150
babies during the first 2 days after their birth, George Michel (1981) found that
two-thirds consistently preferred to lie with their heads turned to the right When
he again studied a sample of these babies at age 5 months, almost aU of the '"head
right" babies reached for things with their right hands, and almost all of the
left" babies reached with their left hands. Such findings, along with the UDitasal
prevalence of right-handers, indicate that either genes or some prenatal fadaI- in
fluence handedness.
So, Is IT ALL RIGHTTO BE LEfT-HANDED? Judpngbyoureverydayconversatbl.ldt
handedness is not all right. To be "coming out of left field" or to offer a "Ieft-kmded
compliment" is hardly better than to be "gauche" or "sinister" (words deria from
the French and Latin for "left"). On the other hand, right-handedness is "rigllton,"
which any "righteous" man" "in his right mind" usually is.
Left-handers are more numerous than usual among those with reading disabilities,
allergies, and migraine headaches (Geschwind & Behan, 1984). But left-handedness is
NEUROSCIENCE AND BEHAVIOR 89
FIGURE 2.28
Babies exhibit cerebral specialization
Consecutive video frames from a study by
Slobhan Holowka and Ann Petitto (2002)
show the left side of a baby's mouth opening
as a smile begins, the right side while
babbling. How about you? Do you more
easily smile with the left or with the right
side of your mouth?
Answers to questions on page 87: Yes.
No. Green.
90 CHAPTER:Z NEUROSCIENCE AND BEHAVIOR
Long-lived left-handers:
Benjamin Franklin (84)
Charlie Chaplin (88)
Pablo Picasso (92)
The Queen Mother (101)
Short-lived left-handers:
Babe Ruth (53)
Marilyn MOliroe (36)
Alexander the Great (33)
Kurt Cobain (27)
also more common among mUSICIans, mathematicians, professional baseball and
cricket players, architects, and artists, including such luminaries as Michelangelo,
Leonardo da Vinci, and Picasso.
2
Although left-handers must tolerate elbow jostling at
dinner parties, right-handed desks, and awkward scissors, the pros and cons of being a
lefty seem roughly equal. But researchers have turned up one con that deserves more at
tention: Left-handers seem to disappear with age!
THE CASE OF THE DISAPPEARING SOUTHPAWS While studying handedness, psycholo
gist Stanley Coren (1993) stumbled upon a rather stunning fact: With age, the per
centage of left-handers declines dramatically. In his initial sample of 5147 people, he
found that 15 percent of lO-year-olds, 5 percent of 50-year-olds, and less than 1 per
cent of those over age 80 were left-handers (FIGURE 2.29). Other researchers around the
world have confirmed Coren's finding. Intrigued, Coren and fellow sleuth Diane
Halpern (1991; Halpern & Coren, 1988, 1991, 1993) set out in search of an answer. (If
you were they, what explanations might have come to mind?)
Perhaps, Coren and Halpern first thought, childhood coercion causes natural left
ies to become right-handed as they age. (Many older people can recall having their left
hand slapped, tied down, or even balled into a fist with surgical tape when they tried to
use it.) If today's parents and teachers are more accepting of left-handedness than were
those of earlier times, the result might be more young left-handers. But North American
and European studies suggest that during the twentieth century, left-handers increased
only from about 6 percent of the population to 10 percent (Porac & others, 1980), too
little to explain the much greater percentage of left-handers among the very young.
Coren and his colleagues found further support for the stability of left-handedness in art
works dated from 15,400 to 3000 B.C., where 10 percent of the people were depicted as
left-handed. In modem art works, 11 percent are left-handers.
Might simple learning explain the slow and gradual disappearance of left-handers?
Might the world's being designed for right-handers make it easier for lefties to learn grad
ually to use their right hands? But again, no-preschoolers who switch handedness com
plete the process before adolescence. Handedness rarely switches after age 8 or 9, and
even before then it occurs only for specific coerced actions (such as eating or writing).
What else remains? Coren and Halpern dared to think the unthinkable: that
left-handers die younger. "That can't be true," skeptical colleagues replied when
Coren first voiced the idea. "If true, surely someone would have noticed by now. Be
sides, my paternal grandmother was left-handed and lived to 91." But vivid anecdotes
("I know a person who ...") are no substitute for conclusions based on large and
representative samples. Neither examples nor counterexamples prove a generality.
So Coren and Halpern decided to explore their morbid idea. They first reflected
on the lefties' known health risks. Left-handers are more likely to have experienced
birth stress, such as prematurity or the need for assisted respiration. They also en
dure more headaches, have more accidents (partly because of right-handed equip
ment), have more knee and joint problems, use more tobacco and alcohol, and
suffer more immune system problems (including allergies such as asthma, eczema,
and hay fever).
These handedness differences aren't huge (individual differences are much
greater). But might they, like gender differences in health risks, add up to differing
life spans? When Coren and Halpern studied a random sample of recently
deceased people of all ages they found that, on average, right-handers did live
eight or nine years longer. This difference dwindled a :bit when Coren, Halpern,
and other researchers excluded children. By comparing left-handed and right
handed former baseball and cricket players, they found the life-span difference re
'Strategic factors explain the higher-than-normal percentage of lefties in sports. for example, it helps a soccer team to
have left-footed players play the left side of the field (Wood & Aggleton, 1989).
"
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 91
Percentage of 14%
left-handedness
12
10
8
4
2
o
10 20 30 40 50 60 70 80 90
Age in years
duced-three years, Coren estimated-but still apparent (Aggleton & others, 1993;
Rogerson, 1994).
This stunning finding triggered an avalanche of publicity, much of it distorted,
which in turn triggered nasty letters ("You have some gall to publicly ridicule people
you have just condemned to an early death"), newspaper columns ("Dear professor ...
we strongly suggest that you beware of sinister lefties bearing grudges-and chain
saws"), and phone messages ("You right-handers think that you'llUve longer than
left-handers but you won't if we kill you first" [Halpern & others, 1996]). The find
ing also stimulated some follow-up research that produced no life-expectancy advan
tage for right-handers (Harris, 1993). One National Institute on Aging research team
followed 3800 East Boston adults for 6 years and found that, at any age, left-handers
were not more likely to die (SaUve & others, 1993). Coren (1993) responded that 6
years is not long enough to catch a statistically Significant handedness effect in a
sample this small.
The unfinished case of the disappearing southpaws illustrates the very heart of sd
ence-how it dares to ask researchable questions, even ftlose witlYunsettJing implica
tions. In the court of sdentific judgment., researchers are welcome to state new ideas but
they must expose themselves to counterexamination from opposing views and to heated
:lebate. Over time, science has a way of unmasking error, taking us closer In truth. and
often to an improved world. As it explains the absence of elder left-handers. it may also
point the way to safer, more comfortable environments for left -handed people.
FIGURE 2.29
The disappearing southpaws
The percentage of left-handers decreases
sharply in samples of older people.
(Adapted from Coren. 1993b.)
"Truth is arrived at blJ the painstaking
process of eliminating the untrue.
When you have eliminated the impossi
ble, whatever remains, however
improbable, must be the truth."
Sherlock Holmes in Arthur Conan Doyle's "The
Sign of the Four," 1890
The southpaw's hazardous life
In the book The Left-Hander Syndrome,
Stanley Corea illustrates risks posed by
a world made for right-handers. When
left-handers use a drill press, their left
arm may obscure their view.
92 CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
LEFT BRAIN/RIGHT BRAIN
hemispheres. Even when reading a
"Error flies from mouth to mouth, from pen to pen, and to destroy it
story, we use both hemispheres-the
'J
takes ages."
You've heard or read it many times:
Some people are "left-brained," oth
ers "right-brained." Leaping from the
new research on split and intact
brains, educators, management ad
visers, and self-help writers urge us
to harness the undeveloped half of
our brain. Do you want to develop
your creativity, music appreciation,
or emotional empathy? Well, get
your brain in balance. Awaken your
dormant right hemisphere. Advocate
"whole brain" education in your
schools. Unleash intuitive, right
brained management from the
chains of cold logic and statistics. Try
Drawing on the Right Side ofthe Brain
(the title of a million-copy bestseller
translated into 10 languages).
What should we make of all
this? By calling my friend Elsie
Voltaire, 1694-1778
"right-brained" have I explained why
she's such a zany free spirit? And
why her "left-brained" husband Bill
is so coolly analytical? Research does
show that each hemisphere serves
special functions. The left is more
logical, verbal, and able to deal with
things in sequence. The right is more
emotionally intuitive and expressive,
skilled at spatial relations, and able
to deal with things all at once. But
neuroscientists raise a caution flag:
Beware the fad of locating complex
human abilities such as science or
art in either hemisphere. "The left
right dichotomy in cognitive mode is
an idea with which it is very easy to
run wild," warned Roger Sperry
(1982). Actually, such complex ac
tivities as doing science or creating
art require the integration of both
left processing the words and find
ing meaning, the right appreciating
humor, imagery, and emotional con
tent (Hellige, 1993; Levy, 1985).
Why, then, do the populariza
tions of brain research so greatly
exaggerate the findings? In The Left
Handa Syndrome, University of British
Columbia psychologist Stanley Coren
(1993) illustrates how journalism
often oversimplifies and embellishes
science. He recalls hearing a conven
tion talk by Doreen Kimura, a psy
chologist at the University of Western
Ontario in London, Ontario. Kimura
reported that melodies fed to the left
ear were more easily recognized than
melodies fed to the right ear. Know
ing that the left ear sends most of
its information to the right hemi
sphere, she concluded that, among
her right-handed student volunteers,
the right brain was better at recog
nizing melodies.
A few days later, the New York
Times reported that "Doreen Kimura,
a psychologist from London, On
tario, has found that musical ability
is controlled by the right side of the
brain" (italics highlight the embell
ishment). Apparently drawing from
the Times story, a syndicated news
paper story then reported that "Lon
don psychologist, Dr. Doreen Kimura,
claims that musicians are right
brained!" (But Kimura studied uni
versity students, not musicians.) Later,
a follow-up newspaper article further
distorted the study: "An English psy
chologist has finally explained why
there are so many great left-handed
musicians."
Knowing that Kimura is not
English, did not study musicians,
and did not study left-handers,
Coren recalled the words of an
American editor: "Everything you
read in the newspaper is absolutely
true, except for the rare story of
Left br_/rlght brain
Is the adist a "right-brained" person? Such notions distort
sclentifidindings regarding the cerebral hemispheres. White
some areas of each hemisphere specialize in different tasks, the
brain's two sides cooperate to enable the person in whatever he
or she does.

....
'IY.WhiCh you happen to have first
'. and knowledge."
.;' What can happen is this: As
. nformation flows from scientist to
,reader, it gets simplified as well as
F.'embellished, much as gossip does in
from one person to the
:'next. A TV network picks up an in
: teresting finding, then reduces it to
,;a 30-second report with an ll-second
sound bite from the researcher. This
alerts a major newspaper to a story
angle, which in turn gets picked up
by popular science magazines and,
eventually, by supermarket maga
zines and tabloids.
At each step, notes Coren,
"Ideas become more speculative and
more distant from the actual re
search.... After a while. the neu
ropsychologist is no longer even
visible in the communication chain."
The rumors grow, accumulate, and
evolve into scientific misinforma
tion that becomes "'accepted truths,'
which show up in conversation and
writing in sentences that begin with,
'As everybody knows ... " or 'Scien
tists have shown that ...'" In tbeend,
sighs Coren, the public myth drowns
out the voices of dissenting sdeotists.
The moral is not to discount
everything you read. Surely Oscar
Wilde was too cynical when c:apress
ing gratitude "for modern journal
ism. By giving us the opinions of the
uneducated, it keeps us iD. touch
with the ignorance of the amunu
nity." (Some journalists cbed the
accuracy of their draft articks with
those they've interviewed.) The
moral is to beware. Reportas want
their stories to be ne'WdllOlthy.
When at their best, their (and
mine, in writing this book) isID ex
tract the essence-to simplitjowith
out becoming simplistic. WIIm at
their worst, they distort a pazel
shaped finding into a bm&ti.ck
shaped story: Some peoJ!Ie are
left-brained, others right-brai.nej, ..
CHAPTER 2 NEUROSCIENCE AND BEHAVIOR 93
__ __ ________
The Brain
The Tools of Discovery
Clinical observations have long revealed the general effects of damage to various areas of
the brain. But CT and MRI scans now reveal brain structures, and EEG, PET, and functional
MRI recordings reveal brain activity. By surgically lesioning or electrically stimulating spe
cific brain areas, by recording the brain's surface electrical activity. and by displaying
neural activity with computeraided brain scans, neuroscientists explore the connections
among brain, mind, and behavior.
lower-level Brain Structures
The brainstem begins where the spinal cord swells to form the medulla, which controls
heartbeat and breathing. Within the brainstem, the reticular formation controls arousal.
Atop the brainstem is the thalamus, the brain's sensory switchboard. The cerebellum, at
tached to the rear ofthe brainstem, coordinates muscle movement.
Between the brainstem and cerebral cortex is the limbic system, which is linked to
memory, emotions, and drives. One of its neural centers, the amygdala, is involved in re
sponses of aggression and fear. Another, the hypothalamus, is involved in various bodily
maintenance functions, pleasurable rewards. and the control of the hormonal system.
The Cerebral Cortex
Each hemisphere of the cerebral cortex-the neural fabric that covers the hemispheres
has four geographic areas: the frontal, parietal, occipital, and temporal lobes. Small, well
defined regions within these lobes control muscle movement and receive information from
the body senses. However, most of the cortex-its association areas-is uncommitted to
such functions and is therefore free to process other information.
Some brain regions serve specific functions (FIGURE 2.30, page 94). The brain di
vides its labor into specialized subtasks and then integrates the various outputs from its
neural networks. Thus, our emotions, thoughts, and behaviors result from the intricate co
ordination of manybrainareas. Language, for example, depends on a chain of events in
several brain regiOns. Ifone hemisphere is damaged early in life, the other will pick up
many of its functions,. thus demonstrating the brain"s plasticity. The brain becomes less
plastic later in tife. frequently, however, nearby neurons can partially compensate for dam
aged ones, as when aperson recovers from a stroke or brain injury.
Our Divided Brains
Clinical observations long ago revealed that the left cerebral hemisphere is crucial for lan
guage. Experiments on people with a severed cCKpUS callosum have refined our knowledge
of each hemisphefe"s special functions. Separatelytesting the two hemispheres. reo
searchers have coaIirmed that in most people theleft hemisphere is the more verbal, and
that the right heaIisfIIere ex:els in visual percep&. andlhe recognition of emotion. Stud
ies of healthy peopIewidt intact brains confirm tlliillteadl hemiSPhere makes unique contri
butions toIhe inteJ:faled functioning of the brain.
(HECK YOURSElf: Wdbin what brain region MMIIddamage be tikelyto disrupt your
ability to skip rope?Your abUity to sense tastes orsounds? In what brain region would
damage perhaps teMe you in a coma? Without thewry breath and heartbeat of life?
ASK YOURSELF: How might you feel with two separate brain hemispheres, both of which
controlled your thought and action but one of which dominated your consciousness and
speech? How might that affect your sense of self, as one indivisible person?
Answers to the (heck Yourself questlons can be found In the yellow appendix at the end of the book.
94 CHAPTER 2 NEUROSCIENCE FIND BEHFlVIOR
FIGURE 2.30
Cerebral cortex:
Corpus callosum: ---,
Brain structures and their functions
ultimate control and
axon fibers connecting
information-processing
two cerebral hemispheres
Right hemisphere
center
left . /
. / /;.
) !
Amygdala:
neural centers
In the limbic
system linked
to emotion
Hippocampus:
a structure in
the limbic
system linked
to memory
Thalamus: -------,
relays messages between
lower brain centers
and cerebral cortex
Hypothalamus: ----___.,.
controls maintenance
functions such as eating;
helps govern endocrine
system; linked to emotion
and reward
Pituitary: ------
master endocrine gland
Reticular formation: ----'"'"
helps control arousal
Medulla:
controls heartbeat and
breathing
Spinal cord:
pathway for neural fibers
traveling to and from brain;
controls simple reflexes
---------Cerebellum:
coordinates voluntary
movement and balance
Cerebral cortex Limbic system Brainstem
The Endocrine System
Preview: This chapter has focused on the body's speedy electrochemical
information There is, however, another communication system.
Hormones released by the endocrine glands form the body's slower
information system.
I
nterconnected with the nervous system is the second of the body's communication.
systems, the endocrine system (FIGURE 2.31). The endocrine system's glands se
crete another form of chemical messengers, hormones. Hormones originate in one
tissue, travel through the bloodstream, and affect other tissues, including the brain.
When they act on the brain, they influence our interest in sex, food, and aggression ...
Some hormones are chemically identical to neurotransmitters (those chemical
messengers that diffuse across a synapse and excite or inhibit an adjacent neuron).
The endocrine system and nervous system are therefore kindred systems: They both
secrete molecules that activate receptors elsewhere. But unlike the speedy nervous
system, zipping messages from eyes to brain to hand in a fraction of a second, en
endocrine [EN-duh-krinJ system the
docrine messages trudge along. If the nervous system's communication delivers mes
body's "slow" chemical communication
sages rather like e-mail, the endocrine system is the body's snail mail. Several seconds
system; a set of glands that secrete
or more may elapse before the bloodstream carries a hdrmone from an endocrine
hormones into the bloodstream.
gland to its target tissue. But these endocrine messages are often worth the wait be
hormones chemical messengers,
cause their effects usually outlast the effects of a neural message.
mostly those manufactured by the
endocrine glands, that are produced in
The endocrine system's hormones influence many aspects of our lives-growth,
one tissue and affect another. reproduction, metabolism, mood-working to keep everything in balance while we re
CHAPTER 2
'//'
Hypothalamus , ;':')) Pituitary gland
(brain region ); V.(( (secretes many different
controlling the' . i .., hormones. some ofwhkh
pituitary gland) f ; affect other glands)
Parathyroids
Thyroid gland

(help regulate the level
(affects metabolism.
./ :-! of calcium in the blood)
among other things) ..
I
Adrenal stands
(inner part. called
the medulla. helps
trigger the "fight or
! flight" response)
Pancreas
t
\ \
i
(regulates the
level of sugar
, in the blood)
"
! ( .'

\
" (secretes female
. '\; r
\
sex hormones)
TestiS
(secretes male sex
hormones) ,
spond to stress, exertion, and our own thoughts. In a moment of danger, for example,
the autonomic nervous system orders the adrenal glands on top of the kidneys to re
lease epinephrine and norepinephrine (also called adrenaline and noradrmaline). These
hormones increase heart rate, blood pressure, and blood sugar, providing us with a
surge of energy. When the emergency passes, the hormones-and the feelings of ex
citement-linger a while.
The most influential endocrine gland is the pituitarY ghmd, a pea-sized struc
ture located in the base of the brain, where it is controlled by an adjacent brain area
called the hypothalamus. The pituituy releases hormones that influence growth, and
its secretions also influence the release of hormones by other endocrine glands. The
pituitary then is really a sort of master gland (whose own master is the hypothala
mus). For example, under the brain's influence, the pituitary triggers your sex glands
to release sex hormones. These in tum influence your brain and behavior.
This feedback system (brain -+ pituitary --+ other glands -+ hormones -+ brain)
reveals the intimate connection of the nervous and endocrine systems: the nervous
system directing endocrine secretions, which then affect the nervous system. In fact,
the two systems are so closely connected that the distinction I>ct:Jm:n than some
times blurs. Researchers have disawered that neurotransmit:tc:ls an drift in the
brain's fluid to nerve receptors at distant sites, thereby affecting 0Ia3J1 akrtness or
mood (Agnati & others, 1992; Pert" 1986). In such cases, the distindion betwcm cer
tain neurotransmitters and their chemical twins-which are called hormones when
released by glands-is no longer so dear. Conducting and coordimting this whole
electrochemical orchestra is that maestro we caJI the brain.
We have glimpsed the truth ofour overriding principle: Evetything psychologi
cal is Simultaneously biological. This chapter has focused on how our thoughts. feel
ings, and actions arise from our specialized yet integrated brain. In chapters to come,
NEUROSCIENCE RND BEHAVIOR 95
FIGURE 2.31
- The body's major endocrine glands
Ii'! adrenal lah-DREEN-el] glands a pair
of endocrine glands just above the
kidneys.. The adrenals secrete the
hormones epinephrine (adrenaline) and
norepinephrine (noradrenaline), which
help to arouse the body in times of
stress.
pHultaryctand the endocrine
system's most influential gland. Under
the influence of the hypothalamus, the
pituitary regulates growth and controls
other endocrine glands.
CHAPTER 2 NEUROSCIENCE AND BEHAVIOR
ind and brain as holistic system
Roger Sperry's view, the brain creates and
ntrols the emergent mind, which in turn
'luences the brain. (Think vividly about
:ing into a lemon and you may salivate.)
,f the human brain were so simple
lat we (ould understand it, we would
oso simple that we couldn't."
Emerson M. Pugh, quoted by George E. Pugh,
The Biological Origin of Human Values, 1977
"
we will further explore the significance of the biological revolution in psychology. We
will see, for example, how
genes and experience jointly influence our personality, emotions, and
intelligence.
brain development underlies a child's mental development.
our sense organs and our brain enable us to see and hear.
the brain records memories.
aberrant brain anatomy and chemistry influence depression and schizophrenia;
and we'll see how biological treatments can alleviate these conditions.
our brain and body work to create our experiences of hunger and sexuality, anger
and fear, sleep and dreams.
mind and body together influence our vulnerability to disease and our capacity
for healing.
our species' evolutionary history may predispose us to hurt, help, or love certain
others.
From nineteenth-century phrenology to today's neuroscience we have come a
long way. Yet what is unknown still dwarfs what is known. We can describe the brain.
We can learn the functions of its parts. We can study how the parts communicate.
But how do we get mind out of meat? How does the electrochemical whir in a hunk
of tissue the size of a head of lettuce give rise to elation, a creative idea, or that mem
ory of Grandmother?
Much as gas and air, above certain concen
trations, give rise to something different-fire
so, believed Sperry, does the complex human
Mind brain give rise to something different: conscious
ness. The mind, he argued, emerges from the
brain's dance of ions, yet is not reducible to it.
Cells cannot be fully explained by the actions of
Brain
atoms, nor minds by the activity of cells. Psy
chology is rooted in biology which is rooted in
chemistry which is rooted in physics. Yet psy
chology is more than applied physics. As Jerome
Kagan (1998) reminds us, the meaning of the Gettysburg Address is not reducible to
neural activity. Sexual love is more than blood flooding to the genitals. Morality and
responsibility become possible when we understand the mind as a "holistic system
property," said Sperry (1992). We are not mere jabbering robots.
The mind boggles both at what is known and what is not. Interviews with lead
ing brain scientists reveal their own awe and wonder. Others ponder philosophical
mysteries: How does the material brain give rise to consciousness? To what extent can
a thing understand itself? The mind seeking to understand the brain-that is indeed
among the ultimate scientific challenges.
CHAPTER 2
NEUROSCIENCE AND BEHAVIOR 97
TERMS AND
psychology, p. 58
P5
8
.dlename, p. 58
potential, p. 59
CONCEPTS TO REMEMBER
brainstem. p. 72
sensory neurons. p. 65 medulla [muh-DUl-uhl. p. 72
interneurons. p. 65 reticular formation, p. 72
motor neurons. p. 65 thalamus [THAl-uh-mussl, p. 73
somatic nervous system. p. 66 cerebellum (sehr-uh-BEll-uml,
autonomic (aw-tuh-NAHM-ikl P73
nervous system. p. 66 limbic system, p. 74
sympathetic nervous system, p. 66 amygdala (uh-MIG-duh-Ia). p. 74
p. 60
parasympathetic nervous system. hypothalamus (hj-po-THAl-uh
Il's;vnllipse (SIN-aps), p. 60
p.66 75
rleurotransmitters, p. 61
reftex. p. 67 cerebral (seh-REE-bruhl) cortex,
.;... Icetylcholine (ah-seat-el-KOleen!
neural networks, p. 68 P7
6
L (ACh), p. 62
glial cells, p. n
lesion (lEE-zhuhn), p. 70
-;' endorphins (en-DOR-fins), p. 63
frontal lobes, p. n
electroencephalogram (EEG), p. 70
nervous system, p. 65
parietal [puh-RYE-uh-tuhl! lobes,
CT (computed tomography) scan,
central nervous system (CNS).
pn
P7
1
p.65
occipital (ahk-SIP-uh-tuhl) lobes.
PET (positron emission
peripheral nervous system (PNS).
tomography) scan, p. 71
pn
p.65
temporal lobes, p. 77
MRI (magnetic resonance imaging).
, nerveS, p. 65
P7
1
C!:J. . To continue your study and review of Neuroscience and Behavior, visit this book's ..
:. '< __' Web site at www.worthpublishers.com/myers. You will find Practice Tests, Review
:' I Activities, and many interesting articles and Web links for more information on top
, ics related to Neuroscience and Behavior.
motor cortex, p. 78
, sensory cortex, p. 79
association areas, p. 80
aphasia, p.82
Broca's area, p. 82
Wernicke's area, p. 82
plasticity, p. 84
corpus callosum (KORpus kah
p.85
split brain, p. 86
endocrine (ENduh-krin) system,
P94
hormones, p. 94
adrenal (ah-DREEN-ell glands,
P95
pituitary gland, P.95
i. ~