Table Of ContentFunctional Neuroscience
Springer Science+Business Media, LLC
Oswald Steward, Ph.D.
Reeve-Irvine Research Center
University of California, Irvine
Functional N euroscience
with 306 Illustrations, 89 in color
Springer
Oswald Steward, Ph.D.
Reeve-Irvine Research Center
University of California, Irvine
1105 Gillespie Neuroscience Research Facility
Irvine, California 92697 -4292
USA
Library of Congress Cataloging-in -Publication Data
Steward, Oswald.
Functional neuroscience / Oswald Steward.
p. cm.
Includes bibliographical references.
ISBN 978-0-387-98543-5 ISBN 978-1-4612-1198-3 (eBook)
DOI 10.1007/978-1-4612-1198-3
1. Neurosciences. 1. Title.
[DNLM: 1. Nervous System Physiology. 2. Nervous System
physiopathology. 3. Nervous System-anatomy & histology. WL 102
S849f 1998]
RC341.S85 2000
612.8-dc21
DNLMIDLC
for Library of Congress 98-21387
CIP
Printed on acid-free paper.
© 2000 Springer Science+Business Media New York
Originally published by Springer-Verlag New York, Jnc. in 2000
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987 6 5 4 321
ISBN 978-0-387-98543-5 SPIN 10680218
To
Kathy, Jessie, and Ossie
Preface
This book is designed to be used as the core cusing first on the basic cell biology of neu
neuroscience text for medical students. It rons and glia, and then on signaling pro
presents the key information that is required cesses. In this, I have made certain choices
for a basic understanding of the operation of that reflect the way the medical curriculum
the nervous system. My goal in writing this is organized in most medical schools. For ex
text was to integrate and blend two success ample, there is no chapter that covers resting
ful styles. One is the style that has dominated membrane potential and action potential, or
the neuroanatomical teaching known as the elemental aspects of synaptic transmis
"functional neuroanatomy", in which the sion. These topics are usually taught in the
focus is on regional anatomy, and how le physiology courses in most medical curric
sions at particular sites disrupt different ula. The present book assumes a background
pathways. The second is the style favored by in these subjects, and then moves on to
neurophysiologists known as "systems neu those aspects of synaptic transmission that
roscience" in which the focus is on the phys are especially important for neurons in the
iological operation of the different functional central nervous system (that is, integration
systems. I use the term "functional neuro of information from multiple inputs, interac
science" to refer to the resulting blend. tions between different kinds of neurotrans
The overall organization of the presenta mitter systems, and integration of excitation
tion, as reflected by the chapter headings, is and inhibition).
traditional. We begin by considering the The third section of the book is the heart
ground plan of the nervous system (that is, of what I call functional neuroscience. In
regional anatomy and the organization at this section, each of the key functional sys
different levels of the neuraxis). Then, we tems are considered one by one. We begin
consider cellular aspects of neuroscience, fo- with the somatosensory pathways, and
vII
viii PREFACE
then consider the motor pathways and sen How
TO ApPROACH THE SUBJECT
sory motor integration. The reason for con
OF NEUROSCIENCE
sidering somatosensory and then somato
motor is that the two are mapped in parallel The information that is encompassed by
throughout the neuraxis, and there impor Functional Neuroscience is of critical impor
tant function interactions between sensory tance to physicians of all types. The impor
and motor components at each level. We tance of the information to a specialist is ob
then consider the systems that control the vious, however, family physicians must also
output of the somatic motor pathways, the use this information on virtually a daily
cerebellum and the basal ganglia, and fol basis.
low with a consideration of the special Whenever a patient walks into your office
senses: vision, audition, vestibular system, for whatever reason, you will be carrying out
and the chemical senses (taste and smell). a basic neurological exam. This means ob
The final section considers the systems that serving the patient for any signs and symp
mediate associative and integrative func toms suggestive of disease of the nervous
tions, and the "higher" functions mediated system. When you first see the patient, you
by the cerebral cortex. check for general demeanor. When you
Some instructors who are used to the shake their hand, you check their hand
"functional neuroanatomy" style may be strength. You watch their eyes to detect any
surprised that there is no separate chapter on abnormalities in eye movement. You observe
the cranial nerves. Instead cranial nerves be how they hold themselves in order to detect
longing to a particular functional system are any postural asymmetries. You listen to what
described in the chapter pertaining to that they say to detect any difficulties with speech
system. The cranial nerves that are involved or difficulties with cognition. Part of your
in somatic sensation are described in the so routine neurological exam will involve
matosensory section; the cranial nerves that watching your patient move across the
control the muscles of the face are described room. This will give you an initial indication
in the motor system, and the cranial nerves of their motor status. You will likely do a
that control autonomic functions are de simple check of reflexes, and you may carry
scribed in the chapter on the autonomic ner out simple tests of eye movements to track
vous system, and finally there is a separate visual stimuli. You may ask to the patient to
chapter that considers the control of eye stand with their eyes closed to check their
movement. balance, to touch their nose or ear with their
This book is designed for medical stu eyes closed, to rapidly pat the back and front
dents. I have attempted to keep the presen of their hands to check for cerebellar func
tation as concise as is consistent with accu tion. And then of course, you will listen to
racy. There is minimal consideration of what the patients say about any problems
original data, and the only references are the they are experiencing.
ones indicated as sources for the figures. The Everything that you learn in this way is
cost of maintaining brevity is that I do not then integrated into an opinion about the
cite the thousands of individuals whose re patient's functional situation. If you detect
search forms the basis of the material pre no abnormalities, and in the absence of any
sented herein, or provide a view of how the complaints from the patient, you will con
knowledge was actually derived. Also miss clude that their nervous system is function
ing is a consideration of the limitations of our ing normally. Any complaints, however
present knowledge. For this reason, instruc minor, will cause you to formulate hypothe
tors will need to provide supplementary ma ses about what may be wrong. The impor
terial for graduate students or medical stu tant question that you must ask yourselves
dents interested in neuroscience research. is, "is there a lesion involving the nervous
ix
PREFACE
system, and if so where is it localized?". The each stage, students must make use of infor
answer to these questions will tell you mation learned at earlier stages. This be
whether you handle treatment yourself or comes apparent when trying to understand
refer the patient to a specialist. the consequences of lesions at particular
Learning neuroscience is a challenge. One sites. To understand the consequences of a
reason is that it is difficult to begin by pre lesion involving the brain stem, one must
senting the "big picture" and then move to know the distribution of the somatosensory
the details. Instead, in most cases one can't pathways, the motor pathways, the organi
understand the big picture without knowing zation of the cranial nerve nuclei, the de
the details. Usually it is not until the end of scending autonomic pathways, and the func
the course that all the pieces start to "come tional role that the cerebellum plays in
together". This causes anxiety. The most im modulating motor output. The big picture
portant advice I can give is, be patient. The comes from the details. This type of learning
big picture will eventually become clear. is challenging and sometimes frustrating, but
The other difficulty is in the nature of the IS what you need to know.
information that must be learned. Physicians An important skill that physicians must
must understand the pathways that mediate develop is the ability to identify structures
particular functions. One must be able to from histological sections, CT scans, or mag
predict the site of a lesion on the basis of a netic resonance images (MRI). This skill re
certain symptom complex, and one must be quires an understanding of regional neu
able to predict the symptoms from a known roanatomy and three-dimensional structure.
lesion. This requires integration of a variety In this regard, memorization of a particular
of types of information. Students must also photograph or drawing is not sufficient,
learn about the way signaling occurs in the since any section or image may be taken at a
nervous system. This is a different kind of in slightly different angle or magnification. One
formation, more conceptual in nature than must be able to generalize in order to recog
the memorization required to know where a nize structures in configurations that have
particular pathway lies. Some students excel not previously been seen. Although sections
at learning one type of information, others and diagrams are included in this book, the
excel at learning the other. It may help to full development of the ability to identify
recognize individual differences in order to structures will require practice with a variety
focus one's efforts for maximal efficiency. images. Students are encouraged to explore
One important aspect of neuroscience is image-based self-study programs which are
that one can't learn something and then put available by subscription, or to use any of a
it aside to move on to something else. At number of human brain atlases.
Acknowledgments
I would like to thank two colleagues have The original drawings and diagrams in this
played an especially important role in shap book are the work of a single artist, Ms. Ann
ing my philosophy regarding the teaching of Dunn. I would like to acknowledge her creativ
neuroscience. The first is Edwin R. Rubel, ity and her tolerance for the uneven schedules,
formerly of the University of Virginia, and and what must sometimes have seemed like
now at the University of Washington. Dr. artistic whims of the author. Thanks also to
Rubel's lectures on sensory systems were my Paula Falk for help in assembling the figures
first exposure to what I now call functional and to Elaine Lowe, who helped with the
neuroscience. The second key colleague is proofreading of the manuscript.
Dr. Gary Banker who directed the neuro Finally, I would like to thank William Day,
science course at the University of Virginia, Senior Medical Editor at Springer-Verlag
and was a master in the "functional neu from the time I began this project through
roanatomy" style. Both of these individuals the time that I submitted the manuscript. His
as well as other colleagues as the University confidence in the concept of the book helped
of Virginia will certainly detect elements of to motivate me during those times when
their own teaching style in this book. progress was slow.
xi
Contents
Part I
The Ground plan of the Nervous System .......................... .......... 1
1. Ground Plan of the Nervous System I
Principles of Central Nervous System Development ........ ..... .. ................. 3
2. Ground Plan of the Nervous System II
Levels of the Neuraxis ............ ...... .. .. ......... .. ...... .. .. ... ..... 25
3. Central Nervous System Vasculature, Blood-Brain Barrier,
and Cerebrospinal Fluid .... ............. .......... .. .................. 51
Part 2
Cellular Basis of Neuronal Signaling ....................................... 67
4. Cell Biology of Neurons ...... ......................................... 69
5. Cell Biology of Glia ........ ... ........... .. ....... .. .......... ........ 91
6. Presynaptic Mechanisms Underlying Neurotransmitter Synthesis,
Storage, Release and Inactivation ................... .... ........ .. ...... 107
7. Postsynaptic Mechanisms: Neurotransmitter Receptors
and Signal Transduction Cascades ........ .......... ... .............. ... . 127
8. Neuronal Integration ..... ... .................... ................... .. 145
xiii
