Nolte’s The Human Brain: An Introduction to its Functional Anatomy

720 Pages·2015·0.3 MB·English

by  Todd Vanderah

7
th Edition
NOLTE’S

HUM AN
the

BR AIN
An Introduction to its Functional Anatomy
Todd W. Vanderah, PhD
Professor and Head
Department of Pharmacology
University of Arizona
Tucson, Arizona
Douglas J. Gould, PhD
Professor and Vice Chair
Department of Biomedical Sciences
Oakland University William Beaumont School
of Medicine
Rochester, Michigan
1600 John F. Kennedy Blvd.
Ste 1800
Philadelphia, PA 19103-2899
NOLTE’S THE HUMAN BRAIN: AN INTRODUCTION TO ITS
FUNCTIONAL ANATOMY, SEVENTH EDITION ISBN: 978-1-4557-2859-6
Copyright © 2016 by Elsevier, Inc. All rights reserved.
Previous editions copyrighted: 2009, 2002, 1999, 1993, 1988, 1981.
No part of this publication may be reproduced or transmitted in any form or by any means,
electronic or mechanical, including photocopying, recording, or any information storage and
retrieval system, without permission in writing from the publisher. Details on how to seek
permission, further information about the Publisher’s permissions policies and our arrangements
with organizations such as the Copyright Clearance Center and the Copyright Licensing Agency,
can be found at our website: www.elsevier.com/permissions.
This book and the individual contributions contained in it are protected under copyright by the
Publisher (other than as may be noted herein).
Notices
Knowledge and best practice in this field are constantly changing. As new research and
experience broaden our understanding, changes in research methods, professional practices,
or medical treatment may become necessary.
Practitioners and researchers must always rely on their own experience and knowledge in
evaluating and using any information, methods, compounds, or experiments described herein.
In using such information or methods they should be mindful of their own safety and the
safety of others, including parties for whom they have a professional responsibility.
With respect to any drug or pharmaceutical products identified, readers are advised to check
the most current information provided (i) on procedures featured or (ii) by the manufacturer
of each product to be administered, to verify the recommended dose or formula, the method
and duration of administration, and contraindications. It is the responsibility of practitioners,
relying on their own experience and knowledge of their patients, to make diagnoses, to
determine dosages and the best treatment for each individual patient, and to take all
appropriate safety precautions.
To the fullest extent of the law, neither the Publisher nor the authors, contributors, or
editors, assume any liability for any injury and/or damage to persons or property as a matter
of products liability, negligence or otherwise, or from any use or operation of any methods,
products, instructions, or ideas contained in the material herein.
Library of Congress Cataloging-in-Publication Data
Vanderah, Todd W., author.
Nolte’s The human brain : an introduction to its functional anatomy / Todd W. Vanderah,
Douglas J. Gould.—Seventh edition.
p. ; cm.
Human brain : an introduction to its functional anatomy
Preceded by: The human brain / John Nolte. 6th ed. c2009.
Includes bibliographical references and index.
ISBN 978-1-4557-2859-6 (pbk. : alk. paper)
I. Gould, Douglas J., author. II. Nolte, John. Human brain. Preceded by
(work): III. Title. IV. Title: Human brain : an introduction to its functional anatomy.
[DNLM: 1. Central Nervous System—anatomy & histology. 2. Brain—anatomy &
histology. 3. Nervous System Physiological Phenomena. WL 300]
QM451
611.8—dc23
2015008889
Content Strategist: Meghan Ziegler
Content Development Specialist: Marybeth Thiel
Publishing Services Manager: Catherine Jackson
Project Manager: Carol O’Connell
Design Direction: Amy Buxton
Printed in the United States of America
Last digit is the print number: 9 8 7 6 5 4 3 2 1
Preface
Over the past 5 to 6 years since the publication of the This edition continues to carry the tradition of cross-
sixth edition, our understanding of the human brain referencing from chapter to chapter in order that the
has continued to rapidly grow with the human brain reader understands how the nervous system all works
being the “unknown frontier” in understanding human together.
behavior. Many new techniques are being developed to In addition to the many traditions held in this new
further measure and understand the functions of the edition, we have added multiple-choice questions to the
human brain and its connections to the periphery. This end of each chapter on Student Consult that will allow
new edition upholds the academic rigor and new tech- for continued learning opportunities. In addition to
nologies established by the previous editions written these questions, other resources, including an electronic
by Dr. Jack Nolte. The basic science of the nervous collection of images and videos, are also available. This
system continues to help in our clinical understanding includes the images from the book and a series of ani-
of normal and abnormal behaviors, while emphasizing mations of three-dimensional brain reconstructions
methods and treatments that are being used in practice produced by Drs. John Sundsten and Kathleen Mulligan
today. Dr. Doug Gould and I have continued to stay true of the University of Washington. Our hope is that stu-
to the ideas on the inception of the book, keeping things dents, enthusiasts, and other faculty find these materials
simple to the facts and not trying to simply obfuscate. useful in their own learning and teaching.
We have updated terminology that is slowly changing, Our challenge in writing an up-to-date text in a rapidly
such as the removal of names from when things were growing field is to have a textbook that remains updated
first discovered but no longer are correct. The new while not including overwhelming information that may
edition has made some changes in figures, tables, and make the function of the human brain difficult to under-
illustrations to better understand and categorize impor- stand. There is hope that the updated version along
tant facts. The current edition continues to use the with added questions to each chapter will continue to
latest medical imaging, including diffuse tensor images aid in students’ ability to learn. We welcome comments
to delineate neuronal pathways. Suggested readings and suggestions for future editions and for our own
have been emphasized and references listed for more knowledge.
detailed information in each area of the nervous system. Todd W. Vanderah
iii
In Memoriam
goal was not only to educate students but to educate
future teachers as well. He continually scoured the
primary literature and behaved with childhood excite-
ment when discovering new explanations for human
brain function. His own love and excitement for the
nervous system would naturally bleed over to his stu-
dents and colleagues, encouraging them to explore and
question the human nervous system.
In working with Jack for over 15 years, my career and
take on life changed from teaching as a “job” to teaching
as an enjoyable hobby with benefits. His ability to make
teaching fun, telling jokes and giving examples, led to an
enriched student environment that resulted in students
wanting more. Our meetings most often included discus-
sions of how we could better educate students. Jack was
on the cutting edge of designing “case-based” instruc-
tion, showing videos of patients for teaching purposes,
having patients present in the classroom, and pushing
ideas of working and taking exams as groups, using
innovative technology, including the virtual brain and
interconnected vocabulary terms across multiple fields
of science. Jack’s desire to create a textbook that was
cutting edge yet to the point for learning purposes was
his continual love. He shared with me chapters, images,
and novel ideas while writing the next edition to continu-
ally produce a product that students would enjoy.
Our time together was not all work. Although most
know him as a professor of the nervous system, Jack
also enjoyed playing handball, woodworking, traveling,
cooking, wonderful deep red wines, a martini with blue
cheese olives, and the joy of eating oysters (things that
When it comes to learning the anatomy and basic func- often appeared in his textbook as examples of nervous
tions of the human nervous system, John “Jack” Nolte system function). In closing, I dedicate this new edition
has played a role as an author and/or a professor for to my teacher, colleague, and friend. I dearly miss Jack’s
hundreds of thousands of students, residents, and physi- enthusiasm for teaching, his humor, his friendship, and
cians. Jack viewed the human brain as an endlessly fas- of course his infamous Birkenstocks.
cinating playground that is forever changing. His lifetime Todd W. Vanderah
v
Video Contents
Video 1-1 An animated version of Figure 1-2, Video 19-2 Orientation of the striatum, seen
showing the major subdivisions of the central inside a translucent cerebral hemisphere.
nervous system.
Video 19-3 Peeling away a cerebral
Video 3-1 Lobes of the cerebral hemisphere. hemisphere to reveal the striatum and globus
pallidus.
Video 5-1 The ventricular system.
Video 20-1 Building a brain. An animation
Video 5-2 The ventricular system, seen inside a
beginning with the brainstem, cerebellum, and
translucent brain.
diencephalon and successively adding the
Video 5-3 Choroid plexus of the lateral, third, hippocampus, lateral ventricles, amygdala, basal
and fourth ventricles. ganglia and internal capsule, subcortical white
Video 6-1 The arterial system at the base of matter, and cerebral cortex.
the brain. Video 20-2 An animated version of Figure
Video 10-1 The spinal cord, showing its gross 20-1, showing the gross anatomy of the
cerebellum.
anatomy and general cross-sectional features.
Video 11-1 The brainstem, showing its gross Video 20-3 Projections of all three pairs of
deep cerebellar nuclei.
anatomy and general cross-sectional features, and
the paths of the medial lemniscus, spinothalamic Video 20-4 An animated version of Figure
tract, and corticospinal tract. 20-15, showing the longitudinal zones of
Video 15-1 Projections from the fastigial cerebellar cortex and the deep nuclei to which
these zones project.
nucleus to the brainstem.
Video 16-1 An animated version of Figure Video 20-5 An animated version of Figure
20-21A, showing the formation and course of the
16-22, showing the internal capsule, corona
superior cerebellar peduncle.
radiata, and neighboring structures.
Video 17-1 Thalamus, lateral ventricle, and Video 24-1 The hippocampus, fornix, and
amygdala, seen inside a translucent cerebral
optic nerve, chiasm, and tract.
hemisphere.
Video 17-2 Animated version of Figure 17-33,
Video 24-2 The hippocampus, fornix,
showing the visual pathway.
amygdala, and limbic lobe.
Video 18-1 The course of corticospinal axons
Video 24-3 Peeling away a cerebral
from motor cortex through the internal capsule
hemisphere to reveal major components of the
and into the cerebral peduncle.
limbic system.
Video 19-1 The basal nuclei, including the
subthalamic nucleus and substantia nigra.
xi
Introduction to the Nervous System
1  Chapter Outline
The Nervous System Has Central and Peripheral Parts
The Principal Cellular Elements of the Nervous System Are Neurons and Glial Cells
Neurons Come in a Variety of Sizes and Shapes, yet All Are Variations on the Same Theme
Neuronal Cell Bodies and Axons Are Largely Segregated Within the Nervous System
Neuronal Organelles Are Distributed in a Pattern That Supports Neuronal Function
Neuronal Cell Bodies Synthesize Macromolecules
Dendrites Receive Synaptic Inputs
Axons Convey Electrical Signals Over Long Distances
Organelles and Macromolecules Are Transported in Both Directions Along Axons
Synapses Mediate Information Transfer Between Neurons
Schwann Cells Are Glial Cells of the PNS
PNS Axons Can Be Myelinated or Unmyelinated
CNS Glial Cells Include Oligodendrocytes, Astrocytes, Ependymal Cells, and Microglial Cells
Some CNS Axons Are Myelinated by Oligodendrocytes, but Others Are Unmyelinated
Astrocytes Provide Structural and Metabolic Support to Neurons
Ependymal Cells Line the Ventricles
Microglial Cells Respond to CNS Injury
The aims of this book are to present and explain some  infiltrate virtually all parts of the body, conveying mes-
basic  anatomical  facts  about  how  the  brain  is  put  sages to and from the CNS. The CNS, ensconced in the 
together and to discuss some aspects of how it works.  skull and vertebral column, is composed of the brain 
This introductory chapter describes in a very general  and the spinal cord (Fig. 1-2). The brain itself has mul-
way the subdivisions of the nervous system, then focuses  tiple subdivisions and is composed of the forebrain, the 
on the cellular elements found within it and the ana- cerebellum, and the brainstem. The forebrain, in turn, 
tomical specializations that adapt these cellular ele- is composed of the two massive cerebral hemispheres 
ments to their respective functions. (separated  from  one  another  by  the  longitudinal
fissure—a long cleft between the two hemispheres) and 
the diencephalona; in an intact human brain most of the 
The Nervous System Has Central and
Peripheral Parts
The  nervous  system  is  broadly  subdivided  into  the  aMuch seemingly arcane neuroanatomical terminology has a Latin 
or Greek derivation that actually makes sense. In this case, enceph-
peripheral nervous system  (PNS)  and  the  central
alon is Greek for “in the head” (i.e., “brain”). Diencephalon means 
nervous system (CNS) (Fig. 1-1). The PNS is the collec-
“in between brain,” signifying that this part of the CNS is inter-
tion  of  spinal  and  cranial  nerves  whose  branches   posed between the cerebral hemispheres and the brainstem.
1
2 The Human Brain
numbers. There are around 100 billionb neurons in the 
human nervous system and a similar number of glial 
cells.
CNS
PNS
Neurons Come in a Variety of Sizes and
Shapes, yet All Are Variations on the
Same Theme
Neurons are in the business of conveying information. 
They do so by a combination of electrical and chemi-
cal signaling mechanisms: electrical signals are used 
to  convey  information  rapidly  from  one  part  of  a 
neuron to another, whereas chemical messengers are 
typically used to carry information between neurons. 
Hence  there  are  anatomically  specialized  zones  for 
collecting,  integrating,  conducting,  and  transmitting 
information (Fig. 1-3, Table 1-1). All neurons have a 
cell body (or somac) that supports the metabolic and 
synthetic  needs  of  the  rest  of  the  neuron.  Most 
neurons have a series of branching, tapering processes 
called dendrites (Greek for “like a tree”) that receive 
information  from  other  neurons  at  junctions  called 
synapsesd and one long, cylindrical process called an 
axon  that  conducts  information  away  from  the  cell 
body.  The  axon  gives  rise  to  a  series  of  terminal 
branches that form synapses on other neurons. Hence 
neurons are anatomically and functionally polarized, 
with electrical signals traveling most often in only one 
direction under ordinary physiological circumstances. 
(The molecular underpinnings of this anatomical and 
functional  polarization  are  discussed  in  Chapters  7  
Figure 1-1 Central  and  peripheral  nervous  systems.  The  central  to 9.)
nervous system (CNS) is encased in the skull and vertebral canal. The  Despite the basic similarity of neurons to one another, 
peripheral nervous system (PNS) is attached to the CNS, but its nerve  there is wide variability in the details of their shapes and 
fibers are distributed throughout the body. (Redrawn from Krstić RV:
sizes (Fig. 1-4). Certain aspects of somatic, dendritic, 
General histology of the mammal, Berlin, 1985, Springer-Verlag.)
and axonal morphology give rise to a descriptive termi-
nology  for  neurons. The  vast  majority  of  vertebrate 
neurons are multipolar, meaning that there are multiple 
dendritic projections from the cell body and almost 
always an axon as well (see Fig. 1-4A to E); in many cases 
diencephalon is hidden from view by the massive cere- the pattern of the dendritic processes is characteristic of 
bral hemispheres. The brainstem is that part of the CNS,  that type of neuron. Some neurons are bipolar (see Fig. 
exclusive of the cerebellum, that lies between the fore-
brain and the spinal cord.
The Principal Cellular Elements of the
Nervous System Are Neurons and
bIt is hard to get a sense of how big numbers like this really are, so 
Glial Cells
analogies sometimes help. If you could count one neuron per 
second, and took no breaks for anything else, it would take you 
Despite the large size and widespread distribution of  more than 3000 years to count 100 billion neurons!
the nervous system, it contains only two principal cat- cSometimes also referred to as the perikaryon. Karyon is Greek 
egories  of  cells—nerve cells,  or  neurons,  which  are  for “nucleus,” and, strictly speaking, the perikaryon is the cyto-
plasm surrounding the nucleus of a neuron. However, the term is 
the  information-processing  and  signaling  elements, 
commonly used to refer to the entire cell body.
and glial cells, which play a variety of supporting roles.  d“Synapse” is derived from two Greek words meaning “to fasten 
Both neurons and glial cells are present in enormous  together.”
CHAPTER 1 Introduction to the Nervous System 3
Cerebral
Diencephalon
hemisphere
Brainstem
Cerebellum
Spinal cord
Diencephalon
Brainstem
Cerebellum
A B C
Figure 1-2 Three-dimensional reconstruction of the entire CNS, seen from the left side (A), from directly in front (C), and from halfway between (B). 
The eyes are included with the reconstruction because, as described in Chapter 2, the retina develops as an outgrowth from the same neural tube 
that becomes the CNS. The arrow in C indicates the longitudinal fissure that separates the two cerebral hemispheres. (See Video 1-1.) (Courtesy Dr.
John W. Sundsten, Department of Biological Structure, University of Washington School of Medicine.)
Dendrite
Synapse
Axon
Figure 1-3 Schematic view of a typical neuron, indicating synaptic inputs to its dendrites (although other sites are possible) and the flow of electrical 
information down its axon, reaching synaptic endings on other neurons. Electrical information flow is unidirectional due to molecular specializations 
of various parts of neurons, as described in Chapters 7 and 8. The pink segments covering the axon represent the myelin sheath that coats many axons 
(see Figs. 1-27 and 1-33), and the gap in the axon represents a missing extent that might be as long as a meter in the longest axons.
4 The Human Brain
Table 1-1 Parts of a Typical Neuron
Part Description Major Organelles Primary Functions
Dendrites Tapered extensions of cell  Cytoskeleton, mitochondria Collect information from other 
body neurons
Soma (cell body) May have one, two, or many  Nucleus, Golgi apparatus, Nissl  Synthesize macromolecules, 
processes; typically one  substance, cytoskeleton,  integrate electrical signals*
axon, many dendrites mitochondria
Axon Single, cylindrical; may be  Cytoskeleton, mitochondria,  Conduct information to other 
many centimeters long; may  transport vesicles neurons
be myelinated or 
unmyelinated
Axon terminals (synaptic  Vesicle-filled apposition to  Synaptic vesicles, mitochondria Transmit information to other 
endings) part of another neuron; most  neurons
are axodendritic or 
axosomatic, but other 
configurations occur
*As discussed in Chapter 7, the final integration of electrical signals (i.e., conversion of synaptic potentials to trains of action potentials) 
typically occurs at the beginning of the axon.
1-4F) or unipolare (see Fig. 1-4G), having two processes  percentage of neurons with heavy metals, causing them 
or only one, respectively. There is a broad spectrum not  to stand out from an unstained or counterstained back-
only of neuronal shapes but also of neuronal sizes. Cell  ground (see Figs. 1-4 and 1-16A). More recently, however, 
bodies range from about 5 to 100 µm in diameter. Many  a variety of methods relying on microinjection, immu-
axons are short, only a millimeter or so in length; but  nocytochemical,  or  other  techniques  have  become 
some, like those that extend from the cerebral cortex to  available (Box 1-1). These now make it possible to cor-
the sacral spinal cord, measure a meter or more.f For  relate the structure of an individual neuron with aspects 
many years, the major technique available for studying  of its function.
the shapes and sizes of neurons was Golgi staining, a  Neurons can also be classified according to their con-
method  that  infiltrates  all  the  processes  of  a  small  nections. Sensory neurons either are directly sensitive 
to various stimuli (e.g., touch or temperature changes) 
or they receive direct connections from nonneuronal 
receptor cells. Motor neurons end directly on muscles, 
eAlthough true unipolar neurons are common in invertebrate 
glands, or other neurons in PNS ganglia. Most sensory 
nervous systems, vertebrate neurons with a unipolar appearance 
are actually pseudounipolar. They start out as bipolar neurons, but  and motor neurons live partly in the PNS and partly in 
during development the cell body expands asymmetrically (below),  the CNS (see Fig. 1-10), whereas almost all other neurons 
leaving behind a stalk from which both processes emerge. reside  entirely  in  the  CNS  and  interconnect  other 
neurons. Some are local interneurons and have all their 
processes confined to a single small area of the CNS. 
Others are projection neurons, with long axons con-
necting different areas, as in a neuron in the cerebral 
cortex whose axon reaches the spinal cord. In a strict 
sense, the human nervous system is composed almost 
entirely of interneurons and projection neurons: there 
are at most 20 million sensory fibers in all of the spinal 
and cranial nerves combined and no more than a few 
million motor neurons. Even taking into account the 
fDiagrams and drawings such as those in Figures 1-3 and 1-4 do 
autonomic neurons that innervate muscles and glands 
not convey a sense of the relative sizes of neurons and their parts. 
If you envision the cell body of the spinal motor neuron shown in  (see Chapter 10), more than 99% of our neurons are 
Figure 1-4D as being the size of a tennis ball, then its dendrites  interneurons or projection neurons. However, the words 
would spread out through a room-size volume and its axon would  “sensory” and “motor” are often used in a much broader 
correspond to a garden hose half a mile or so in length. Using the 
sense to refer to cells and axons that carry information 
same scale, the small interneuron shown in Figure 1-4B would be 
related  to  sensory  stimuli  and  to  the  generation  of 
a few pinheads in diameter; its axon would be a hair-thin process 
only 1 or 2 feet long. responses, respectively.