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《探索大脑奥秘》第3版:神经科学经典教材
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"《神经科学:探索大脑》第三版是由马克·F·贝尔(Mark F. Bear)、巴里·康纳斯(Barry W. Connors)和迈克尔·帕拉迪索(Michael A. Paradiso)三位神经科学领域的权威共同编著的一本备受赞誉的本科教科书。该书以其亲和力强的风格、出色的插图和卓越的教学方法,深入浅出地讲解了大脑生物学以及行为背后的神经系统。在第三版中,作者们对味觉和嗅觉、昼夜节律、大脑发育以及发育障碍等主题进行了扩展讨论,提供了更为全面的内容。 新增的章节包括分子机制和功能性脑成像技术,这使得读者能够紧跟最新的科学研究进展。"Path of Discovery"(发现之旅)系列由顶尖研究者撰写,聚焦于当前的重大神经科学研究成果,为学习者提供了深入了解领域前沿的机会。此外,书中还配有一本附有穿孔式自我测试手册的《人类神经解剖图解》,帮助学生巩固神经解剖学知识。 本书的辅助教材非常丰富,包含一个内置的学生CD-ROM,供学习者进行互动式学习;一个教师资源CD-ROM,提供教学材料和答疑支持;一个配套网站Connection Website,提供在线课程资料和互动讨论区;以及LiveAdvise: Neuroscience在线学生辅导服务,进一步增强了学习体验和支持。 版权方面,所有权利受保护,未经版权所有者书面许可,任何形式的复制或存储及检索系统使用均需遵守严格的版权规定。《神经科学:探索大脑》第三版不仅是一本教育工具,也反映了神经科学研究的最新动态,对于理解大脑功能及其在日常生活和医学中的应用具有重要意义。"
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To our ears, the singing of birds may be simply a pleasant
harbinger of spring, but for birds, it is part of the serious
business of sex and reproduction. Singing is strictly a male
function for many species, performed for the purpose of
attracting and keeping a mate and for warning off poten-
tial rivals. Studies of two bird species with different habits
of reproduction and singing have revealed some fascinating
clues about the control and diversity of sexual dimor-
phisms in the brain.
Zebra finches are popular pets, but their wild habitat is
the harsh Australian desert. To breed successfully, birds
require dependable sources of food, but in the desert,
food comes only with sporadic and unpredictable rains.
Zebra finches must therefore be ready and willing to
breed whenever food and a mate are available, in any sea-
son. Wild canaries, on the other hand, live in the more
predictable environment of the Azores and (where else?)
the Canary Islands. They breed seasonally during spring
and summer, and do not reproduce during fall and winter.
The males of both species are passionate singers, but they
differ greatly in the size of their repertoires. Zebra finches
belt out one simple ditty all their lives, and cannot learn
new ones. Canaries learn many elaborate songs, and they
add new ones each spring.The different behaviors of zebra
finches and canaries require different mechanisms of neural
control.
The birds’ sexually dimorphic behavior— singing—is
generated by dramatically dimorphic neural structures.
Birds sing by forcing air past a special muscularized organ
called the syrinx, which encircles the air passage. The
muscles of the syrinx are activated by motor neurons of
the nucleus of cranial nerve XII, which are in turn con-
trolled by a set of higher nuclei collectively called the vocal
control regions, or VCRs (Figure A). In zebra finches and
canaries, VCR size is five or more times larger in males
than in females.
The development of VCRs and singing behavior is under
the control of steroid hormones. However, the very differ-
ent seasonal requirements of zebra finches and canaries
are paralleled by distinctly different modes of steroidal
control. Zebra finches apparently require early doses of
steroids to organize their VCRs, and later androgens to
activate them. If a hatchling female zebra finch is exposed
to testosterone or estradiol, its VCRs will be larger than
those of normal females when it reaches adulthood. If the
masculinized female is given more testosterone as an
adult, its VCRs will grow larger still, and she will then sing
like a male. Females that are not exposed to steroids
when young are unresponsive to testosterone as adults.
By contrast, the song system in canaries seems to be
independent of early steroid exposure, yet it bursts into
full service each spring. If female canaries are given an-
drogens for the first time as adults, they will begin
singing within a few weeks.The androgens of males surge
naturally each spring; their VCRs double in size as neurons
grow larger dendrites and more synapses, and singing
commences. Remarkably, neurogenesis, the birth of neu-
rons, continues throughout adulthood in songbird
brains, further contributing to the VCR circuitry during
the mating season. By fall, male androgen levels drop, and
the canary song system shrinks in size as his singing abates.
In a sense, the male canary rebuilds much of his song
control system anew each year as courtship begins. This
may enable him to learn new songs more easily and, with
his enlarged repertoire, gain some advantage in attracting
a mate.
OF SPECIAL INTEREST
Box 17.1
Bird Songs and Bird Brains
Male
Syrinx
Cranial
nerve XII
Female
Cranial
nerve XII
Syrinx
FIGURE A
Blue circles represent the vocal control regions in male and female zebra finches.
It should never be too late to follow a new idea. That is
what I told myself when, at nearly thirty years old, I aban-
doned my career as a medical doctor, realizing I would be
happier as a scientist. In Chris Miller’s laboratory at Bran-
deis University, I was introduced to potassium channels.
That was the beginning of an exciting adventure for me—
a mixture of “chance and design,” to use Alan Hodgkin’s
words. I think in my case it was mostly chance.
The year was 1986, when biophysicists imagined ion
channels to be membrane pores with selectivity filters
and gates.This essentially correct view had been deduced
by Clay Armstrong, Bertil Hille, and others through
thoughtful analysis of electrophysiological recordings. But
ion channels were not quite “molecular” in the same way
biochemists viewed enzymes. No one had ever visualized
a potassium channel protein. In fact, potassium channel
genes had not yet been identified, so even their amino
acid sequences were a mystery. I began to study what are
known as high-conductance Ca
2⫹
-activated potassium
channels, which we isolated from mammalian skeletal mus-
cle and reconstituted into lipid membranes. My question
was a humble one: How does a scorpion toxin inhibit
these potassium channels? Admittedly, this was not a very
hot topic, in fact you might say it was cold, but that
made no difference to me. I was having fun learning chan-
nel biophysics, and I found the mechanism of toxin inhi-
bition interesting, even if it seemed unimportant. It be-
came clear to me that the toxin functions as a plug on
the pore, and it interacts with ions inside the pore. I spent
long hours trying to imagine what the channel might look
like, and how it could selectively conduct ions at such a
high rate.
About a year into my toxin studies, the potassium chan-
nel field got a huge boost when the laboratories of Lily
and Yuh Nung Jan, Mark Tanouye, and Olaf Pongs reported
the cloning of the Shaker channel from Drosophila. As luck
would have it, I found during a late night experiment at a
Cold Spring Harbor course that the Shaker channel was
sensitive to scorpion toxins. I knew immediately that I
could use scorpion toxins together with site-directed
mutagenesis to identify which amino acids form the ion
conduction pore. That would be valuable information be-
cause the amino acid sequence had no assigned function.
The toxin led me directly to the pore and to other in-
teresting aspects of potassium channels, such as how many
subunits they have. After a few years at Harvard Medical
School, where I had taken a faculty position, my labora-
tory defined which amino acids form the selectivity filter
of the Shaker channel. Conservation of these amino acids
in different potassium channels seemed to underscore the
fact that nature had arrived at a single solution for selec-
tive K
⫹
conduction across the cell membrane. I began to
realize then that I would not understand nature’s solution
without actually seeing the atomic structure (Figure A).
I needed to become a membrane protein biochemist and
X-ray crystallographer. I abandoned my nicely advancing
career as an electrophysiologist at Harvard and moved to
Rockefeller University to concentrate on learning the
new techniques. I was told that I was committing career
suicide because of the difficulty with membrane proteins
and my complete lack of experience. But it made little dif-
ference to me. My reasoning was simple: I would rather
crash and burn trying to solve the problem than not try
at all.Though the lab was initially small, we were very de-
termined. It was a thrilling time because we knew we
were working on a good problem, and we were passion-
ate about it.Through hard work, perseverance, and more
than a little luck, a very beautiful piece of nature slowly
revealed itself to us. It was in fact more beautiful than I
ever could have imagined.
PATH OF DISCOVERY
Box 3.4
The Atomic Structure of a
Potassium Channel
by Roderick MacKinnon
FIGURE A
The protein structure of the potassium channel selectivity filter
(from two of four subunits) is yellow; oxygen atoms are red
spheres. Electron density (blue mesh) shows K
⫹
ions (green
spheres) lined up along the pore. Inside the filter, each K
⫹
ion
binding site is surrounded by eight oxygen atoms, which appear
to mimic the water molecules surrounding the hydrated K
⫹
ion
below the filter. (Courtesy of Dr. Roderick MacKinnon.)
Of Special Interest Boxes
This content complements the
text by enhancing the connection
between neuroscience and real
life, with discussion of brain
disorders, human case studies,
drugs, new technology, and more.
Path of Discovery Boxes
24 new boxes, written by leading
neuroscience researchers, high-
light current discoveries and
achievements of individuals in
the field of neuroscience. Path of
Discovery boxes from the previ-
ous two editions are available on
the Instructor’s Resource CD and
online at http://
connection.lww.com/go/bear.
xiv USER’S GUIDE
BearcFM.qrk(xiv-xiv).ps 1/6/06 9:32 AM Page xiv
spinal cord runs anterior to posterior. The top side of the spinal cord is the
dorsal side, and the bottom side is the ventral side.
If we look down on the nervous system, we see that it may be divided
into two equal halves (Figure 7.2b). The right side of the brain and spinal
cord is the mirror image of the left side. This characteristic is known as
bilateral symmetry. With just a few exceptions, most structures within the
nervous system come in pairs, one on the right side and the other on the
left. The invisible line running down the middle of the nervous system is
called the midline, and this gives us another way to describe anatomical
references. Structures closer to the midline are medial; structures farther
away from the midline are lateral. In other words, the nose is medial to
the eyes, the eyes are medial to the ears, and so on. In addition, two struc-
tures that are on the same side are said to be ipsilateral to each other; for
example, the right ear is ipsilateral to the right eye. If the structures are on
opposite sides of the midline, they are said to be contralateral to each
other; the right ear is contralateral to the left ear.
To view the internal structure of the brain, it is usually necessary to slice
it up. In the language of anatomists, a slice is called a section; to slice is to
section. Although one could imagine an infinite number of ways we might
cut into the brain, the standard approach is to make cuts parallel to one of
the three anatomical planes of section. The plane of the section resulting from
splitting the brain into equal right and left halves is called the midsagittal
plane (Figure 7.3a). Sections parallel to the midsagittal plane are in the
sagittal plane.
The two other anatomical planes are perpendicular to the sagittal plane and
to one another. The horizontal plane is parallel to the ground (Figure
7.3b). A single section in this plane could pass through both the eyes and
the ears. Thus, horizontal sections split the brain into dorsal and ventral
parts. The coronal plane is perpendicular to the ground and to the sagit-
tal plane (Figure 7.3c). A single section in this plane could pass through
both eyes or both ears, but not through all four at the same time. Thus, the
coronal plane splits the brain into anterior and posterior parts.
Key Terms
Appearing in bold throughout the
text, key terms are also listed at
the end of each chapter and
defined in the glossary.
The Golgi Stain
The Nissl stain, however, does not tell the whole story. A Nissl-stained neu-
ron looks like little more than a lump of protoplasm containing a nucleus.
Neurons are much more than that, but how much more was not recog-
nized until the publication of the work of Italian histologist Camillo Golgi
(Figure 2.2). In 1873, Golgi discovered that by soaking brain tissue in a sil-
ver chromate solution, now called the Golgi stain, a small percentage of
neurons became darkly colored in their entirety (Figure 2.3). This revealed
that the neuronal cell body, the region of the neuron around the nucleus
that is shown with the Nissl stain, is actually only a small fraction of the
total structure of the neuron. Notice in Figures 2.1 and 2.3 how different
histological stains can provide strikingly different views of the same tissue.
Today, neurohistology remains an active field in neuroscience, along with
its credo: “The gain in brain is mainly in the stain.”
The Golgi stain shows that neurons have at least two distinguishable
parts: a central region that contains the cell nucleus, and numerous thin
tubes that radiate away from the central region. The swollen region con-
taining the cell nucleus has several names that are used interchangeably:
cell body, soma (plural: somata), and perikaryon (plural: perikarya). The
thin tubes that radiate away from the soma are called neurites and are of
two types: axons and dendrites (Figure 2.4).
The cell body usually gives rise to a single axon. The axon is of uniform
diameter throughout its length, and if it branches, the branches generally
extend at right angles. Because axons can travel over great distances in the
body (a meter or more), it was immediately recognized by the histologists
of the day that axons must act like “wires” that carry the output of the
neurons. Dendrites, on the other hand, rarely extend more than 2 mm in
USER’S GUIDE xv
Further Reading
Recent review articles are identi-
fied at the end of each chapter to
guide further study.
Review Questions
Chapter review questions provoke
thought and help students test
their comprehension of each
chapter’s major concepts.
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An Illustrated Guide to Human
Neuroanatomy
This exceptional appendix to Chapter
7 includes an extensive self-quiz with
labeling exercises that enable
students to assess their knowledge of
neuroanatomy.
Self-Quiz
These brief vocabulary reviews found
in Chapter 7 enhance students’
understanding of nervous system
anatomy.
▼ SELF-QUIZ
This review workbook is designed to help you learn the neuroanatomy that
has been presented. Here we have reproduced the images from the Guide;
instead of labels, however, numbered leader lines (arranged clockwise)
point to the structures of interest. Test your knowledge by filling in the ap-
propriate names in the spaces provided. To review what you have learned,
quiz yourself by putting your hand over the names. This technique greatly
facilitates the learning and retention of anatomical terms. Mastery of the
vocabulary of neuroanatomy will serve you well as you learn about the
functional organization of the brain in the remainder of the book.
236
CHAPTER 7 • APPENDIX: AN ILLUSTRATED GUIDE TO HUMAN NEUROANATOMY
THE LATERAL SURFACE OF THE BRAIN
(a) Gross Features
3
4
1
2
6
5
7
8
9
1. _______________________________
2. _______________________________
3. _______________________________
4. _______________________________
(b) Selected Gyri, Sulci, and Fissures
5. _______________________________
6. _______________________________
7. _______________________________
8. _______________________________
9. _______________________________
xvi USER’S GUIDE
(b) Selected Gyri, Sulci, and Fissures. The cerebrum is
noteworthy for its convoluted surface. The bumps are
called gyri, and the grooves are called sulci or, if they are
especially deep, fissures. The precise pattern of gyri and
sulci can vary considerably from individual to individual,
but many features are common to all human brains. Some
of the important landmarks are labeled here. The post-
central gyrus lies immediately posterior to the central sul-
cus, and the precentral gyrus lies immediately anterior to
the central sulcus. The neurons of the postcentral gyrus
are involved in somatic sensation (touch; Chapter 12),
and those of the precentral gyrus control voluntary move-
ment (Chapter 14). Neurons in the superior temporal
gyrus are involved in audition (hearing; Chapter 11).
▼ SURFACE ANATOMY OF THE BRAIN 209
(c) Cerebral Lobes and the Insula. By convention,
the cerebrum is subdivided into lobes named after the
bones of the skull that lie over them. The central sulcus
divides the frontal lobe from the parietal lobe. The tem-
poral lobe lies immediately ventral to the deep lateral
(Sylvian) fissure. The occipital lobe lies at the very back
of the cerebrum, bordering both parietal and temporal
lobes. A buried piece of the cerebral cortex, called the
insula (Latin for “island”), is revealed if the margins of
the lateral fissure are gently pulled apart (inset). The
insula borders and separates the temporal and frontal
lobes.
(0.5X)
Superior temporal
gyrus
Lateral (Sylvian)
fissure
Postcentral gyrus
Precentral gyrus
Central sulcus
(0.6X)
Occipital lobe
Parietal lobe
Frontal lobe
Temporal lobe
Insula
▼ SELF-QUIZ
Take a few moments right now and be sure you understand the meaning
of these terms:
anterior ventral contralateral
rostral midline midsagittal plane
posterior medial sagittal plane
caudal lateral horizontal plane
dorsal ipsilateral coronal plane
BearcFM.qrk(xvi-xvi).ps 1/6/06 9:33 AM Page xvi
LiveAdvise Neuroscience
This online student tutoring service offers
access to live help from experienced
neuroscience tutors. You can chat with
expert educators and get help while
studying for tests or working on
assignments. See the LiveAdvise code
packaged with this text for more
information on this free service.
ADDITIONAL LEARNING RESOURCES
This powerful learning suite also includes:
Student Resource CD-ROM
Features Answers to Review Questions, Labeling
Exercises, Glossary of Key Terms, and Video Clips
from Acland’s Video Atlas of Human Anatomy.
Materials are also available on the companion
website: http://connection.lww.com/go/bear.
USER’S GUIDE xvii
FIGURE 2.7
The internal structure of a typical neuron.
Neuronal
membrane
Nucleus
Golgi apparatus
Polyribosomes
Smooth ER
Axon
Axon
hillock
Rough ER
Mitochondrion
Ribosomes
Microtubules
Comprehensive Art Program
Exceptional artwork engages
readers and illuminates content.
BearcFM.qrk(i-xxxviii).ps 12/1/05 10:08 AM Page xvii
xviii
Acknowledgments
Back in 1993, when we began in earnest to write the first
edition, we had the good fortune to work closely with a
remarkably dedicated and talented group of individu-
als—Betsy Dilernia, Caitlin Duckwall, and Suzanne
Meagher—who helped us bring the book to fruition.
Remarkably, the team is still intact 12 years later, and,
we modestly suggest, practice makes perfect. We are
proud of the third edition, and very grateful to the con-
tinuing invaluable contributions of Betsy, Caitlin, and
Suzanne.
Betsy is our developmental editor. As always, she kept
us in line with her purple pencil. We are especially grate-
ful for the standard of excellence that Betsy established
and held us to. The clarity and consistency of the writing
are due to her remarkable efforts. In addition, she helped
us improve the layout of the book to make it more read-
er-friendly. Caitlin’s studio produced the new art, and
the results speak for themselves. Caitlin took our some-
times fuzzy concepts and made them a beautiful reality.
Finally, we are forever indebted to Suzanne, who assist-
ed us at every step. It is no exaggeration to say that with-
out her incredible assistance, loyalty, and dedication to
this project, the book would never have been completed.
Suzanne, you are the best!
For the current edition, we have the pleasure of
acknowledging a new “team member,” Elizabeth
Connolly. Elizabeth is an associate development editor at
Lippincott Williams & Wilkins. She worked very closely
with us from start to finish, helping us to meet a
demanding schedule. Her efficiency, flexibility, and good
humor were greatly appreciated.
In the publishing industry, editors seem to come and
go with alarming frequency. Yet two senior editors at
Lippincott Williams & Wilkins have stayed the course
and been unwavering advocates for our project: Nancy
Evans and Susan Katz. Thanks to you and the entire staff
under your direction. It has been a pleasure working
with you.
We again acknowledge the architects and current
trustees of the undergraduate neuroscience curriculum
at Brown University. We thank Mitchell Glickstein, Ford
Ebner, James McIlwain, Leon Cooper, James Anderson,
Leslie Smith, John Donoghue, and John Stein for all they
did to make undergraduate neuroscience great at Brown.
We gratefully acknowledge the research support provid-
ed to us over the years by the National Institutes of
Health, the Whitehall Foundation, the Alfred P. Sloan
Foundation, the Klingenstein Foundation, the Charles A.
Dana Foundation, the National Science Foundation, the
Keck Foundation, the Human Frontiers Science
Program, the Office of Naval Research, and the Howard
Hughes Medical Institute. We thank our colleagues in
the Brown University Department of Neuroscience and
in the Department of Brain and Cognitive Science at the
Massachusetts Institute of Technology for their support
of this project and for helpful advice. A key feature of the
book is the Path of Discovery boxes in which neurosci-
entists describe their research. We thank our new
Discovery authors for these fascinating contributions. We
also thank the anonymous, but very helpful, colleagues
at other institutions who gave us comments on the ear-
lier editions. We are grateful to the scientists who pro-
vided us with figures illustrating their research results. In
addition, many students and colleagues helped us to
improve the new edition by informing us about recent
research, pointing out errors in the first edition, and sug-
gesting better ways to describe or illustrate concepts. We
thank them all, including Gül Dölen, Nancy Kanwisher,
Chris Moore, Steve Mouldin, Luiz Pessoa, Wolfram
Shultz, and Dick Wurtman.
We thank our loved ones for standing by us despite the
countless weekends and evenings lost to preparing this
book.
Last, but not least, we wish to thank the thousands of
students we have had the privilege to teach neuroscience
to over the past 25 years.
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