Ain Anatomy

Human Brain Anatomy
I N C O M P U T E R I Z E D I M A G E S
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Human Brain Anatomy
I N COMPUTERI ZED I MAGES S E C O N D E D I T I O N
HANNA DAMASI O, M.D.
Professor of Neurology and Director
of the Human Neuroanatomy and Neuroimaging Laboratory
at the Department of Neurology
University of Iowa College of Medicine
OXPORD
UNIVERSITY PRESS
2005
OXFORD
UNIVERSITY PRESS
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Library of Congress Cataloging-in-Publication Data
Damasio, Hanna.
Human brain anatomy in computerized images / Hanna Damasio.2nd ed.
p. ; cm.
Includes bibliographical references and index.
ISBN-13 978-0-19-516561-6
ISBN 0-19-516561-6
1. BrainTomographyAtlases. 2. BrainMagnetic resonance imagingAtlases. I.
Title.
[DNLM: 1. Brainanatomy & histologyAtlases. 2. Image Interpretation,
Computer-AssistedAtlases. 3. Magnetic Resonance ImagingAtlases. WL 17 D155h 2005]
QM455.D23 2005
611'.81dc22 2004053157
9 8 7 6 5 4 3 2 1
Printed in China
on acid-free paper
To Antonio
Preface
T
his is the second and entirely revised edition of Human Brain Anatomy in
Computerized Images. The principal aim of the book remains unchanged:
to assist clinicians and researchers in the analysis of human neuroanatomic im-
ages obtained with computerized tomographic techniques and to serve as a tool
for teaching neuroanatomy.
The organization of the second edition is similar to that of the first, but all
the images are new and come from new brain specimens. The book begins, in
Chapters 2-4, with a description of the external anatomy of the brains used in
Chapters 7-9 to depict the different sets of axial and coronal slices. The three
main specimens are a dolichocephalic brain, the brachicephalic brain of a per-
son of Caucasian descent, and the brain of a person of Asian descent, which al-
lowed me to include an extreme brachicephalic example. These brains are shown
in the same views as in the first edition, and their sulci and gyri are identified.
A collection of 26 new normal brains follows the first three, shown from sev-
eral viewslateral, mesial, top, and bottomwith the major sulci identified.
Half are from men and half are from women; gender, age, and handedness are
noted for all.
The chapters depicting axial and coronal slices obtained from the first three
brains include a larger number of slices than in the first edition because the
slice interspace was reduced to 5 mm, to provide greater detail. Also, each mag-
netic resonance slice was stripped of all non-brain structures to facilitate the la-
beling of anatomic details. The original magnetic resonance images for each
slice are posted in the upper and lower corners of the page.
At the beginning of each section, I added an image of the brain with col-
ored gyri and a grid depicting the position of slices. A small black-and-white
image showing the appropriate slice position also has been added to each page
with axial or coronal slices. Sequences of axial slices parallel to the canto-meatal
line, at a rostral tilt of 30 and a caudal tilt of +10, are shown for all three
brains, as are the coronal slices for the first two axial sequences. A parasagittal
[vii]
sequence is shown for the first brain, and Brodmann's cytoarchitectonic fields
Preface are shown in an axial sequence.
A new chapter was added to address issues of brain structure quantification.
A final chapter, providing examples of lesioned brains seen in varied inci-
dences of cut, fulfills the same role as in the first edition but is based on new
cases.
Iowa City, Iowa H.D.
[viii]
Acknowledgments
I
would like to thank all the colleagues who commented on the first edition of
this book and encouraged me to prepare a better second edition. I would also
like to thank Jeff House and Fiona Stevens at Oxford University Press, respec-
tively, the editors of the first and second editions, for their steady support and
guidance.
I am grateful to Joel Bruss, who assisted me in the preparation of many im-
ages, collaborated with John Allen and me in Chapter 6, and helped me with
the index and the rechecking of the final images. He also prepared the images
used in the dust jacket. I will not forget his good will and good humor. I am
equally grateful to Jon Spradling, who, as always, assisted me in virtually all as-
pects of image processing with his usual patience and expertise.
I also acknowledge Sonya Mehta, who was always ready to answer questions
regarding the use of graphics programs; Jocelyn Cole, who traced all the brain
specimens used in the book; Keary Saul and Julie Sexton, who assisted me in
the preparation of the final manuscript; and my assistant Neal Purdum, who
facilitated communication with the editors and solved all sorts of last-minute
problems. I thank all of them deeply for their professionalism and good will.
Finally, I cannot thank Antonio Damasio enough for putting up with all the
hours I spent glued to the computer.
[ix]
Contents
List of Structures Identified in the Figures and Their Abbreviations xiii
1 Introduction 1
2 Exterior Description of a Normal Dolichocephalic Brain 11
Sulci Gyri Brodmann's fields
3 Exterior Description of a Brachicephalic Brain 31
Sulci Gyri
4 Exterior Description of Another Brachicephalic Brain 39
Sulci Gyri
5 An Alphabet of Normal Brains 47
6 Quantifying Neuroanatomic Differences 77
In collaboration with John S. Allen and Joel Bruss
7 Sections through Dolicho 97
Canto-meatal incidence: axial slices 100
Canto-meatal incidence: coronal slices 126
Hyperextension incidence: axial slices 160
Hyperextension incidence: coronal slices 188
Posterior fossa incidence: axial slices 222
Parasagittal incidence 248
Brodmann's fields 286
8 Sections through Brachi-1 293
[xi]
Canto-meatal incidence: coronal slices 320
Contents Hyperextension incidence: axial slices 352
Hyperextension incidence: coronal slices 378
9 Sections through Brachi-2 435
Hyperextension incidence: axial slices 462
10 Application to Lesion Studies 515
A left parietal lesion 517
A left calcarine lesion 521
A right temporal lesion 525
A left frontal lesion (subcortical) 529
References 533
Index of Anatomical Structures Seen in the Figures 537
[xii]
List of Structures
Identified in the
Images and Their
Abbreviations
Sulci
anterior insular sulci
anterior occipital sulcus
ascending branch of sylvian fissure
ascending ramus of cingulate sulcus
calcarine fissure
central sulcus
cingulate sulcus
circular sulcus
collateral sulcus
frontomarginal sulcus
H-shaped sulci
hippocampal fissure
horizontal branch of sylvian fissure
inferior frontal sulcus
inferior temporal sulcus
intraparietal sulcus
lateral occipital sulcus
lateral orbital sulcus
lateral temporo-occipital sulcus
lingual sulcus
middle frontal sulcus
midfrontal sulcus
midorbital sulcus
occipitoparietal sulcus
olfactory sulcus
paracentral sulcus
postcentral sulcus
ants
AOS
abSF
arCingS
CalcF
CS
CingS
CircS,*
ColS
FmS
HsS
HipF
hbSF
IPS
ITS
IPS
LOS
LatOrbS
LTOS
LingS
MFS
mFS
mOrbS
OPS
OlfS
paraCS
postCS
[xiii]
List of Structures Identified in the
Images and Their Abbreviations
posterior insular sulci
precentral sulcus
rhinal sulcus
suborbital sulcus
subparietal sulcus
superior frontal sulcus
superior temporal sulcus
inferior ramus
superior ramus
sylvian fissure
temporo-occipital sulcus
transverse occipital sulcus
transverse temporal sulcus
Gyri, central structures, cerebellum, and
amygdala
anterior commissure
caudate nucleus
body of caudate nucleus
head of caudate nucleus
tail of caudate nucleus
cerebellum
cerebellar hemispheres
cerebellar tonsils
cerebellar vermis
cingulate gyrus
claustrum
colliculi
inferior
superior
corpus callosum
beak
splenium
cuneus
fornix
frontal lobe
frontal pole
fusiform gyrus
gyrus rectus
Heschl's gyrus
hippocampus
hypothalamus
inferior frontal gyrus
pars opercularis
pars orbitalis
posts
preCS
RhS, rhS
sOrbS
sPS
SFS
STS
STSir
STSsr
SF
TOS
trOS, trsOS
trTS
brain stem
a
ac
c
be
he
tc
cereb
cerebH
cerebT
cerebV
CingG
3
ic
sc
CC
bCC, bcc
sCC, sec
Gun
for
FL
FP
FusiG
Grec
HG
hip
ht
IFG
IFG, po
IFG, porb
[xiv]
pars triangularis
inferior parietal lobule
angular gyrus
supramarginal gyrus
inferior temporal gyrus
insula
anterior
posterior
internal capsule
anterior
posterior
lateral occipital gyri
lingual gyrus
mamillary body
medulla
mesencephalon
middle frontal gyrus
middle temporal gyrus
occipital pole
orbital frontal gyri
anterior orbital gyrus
lateral orbital gyrus
middle orbital gyrus
posterior orbital gyrus
pallidum
paracentral gyrus
parahippocampal gyrus
parietal lobe
planum temporale
pons
postcentral gyrus
posterior commissure
precentral gyrus
precuneus
putamen
quadrigeminal plate
retrosplenial area
spinal cord
superior frontal gyrus
superior parietal lobule
superior temporal gyrus
temporal lobe
temporal pole
temporo-occipital gyrus
thalamus
IFG, pt
IPL
AG
SMG
ITG
Ins
1
2
aic
pic
LOG
LingG
mb
med
mes
MFC
MTG
OP
OrbG
aOrbG
lOrbG
mOrbG
pOrbG
pa
paraCG
paHG
PL
PT
pons
postCG
pc
preCG, preCg
preCun
pu
qpl
rSp
sc
SFG
SPL
STG
TL
TP
TOG
th
List of Structures Identified in the
Images and Their Abbreviations
[xv]
Human Brain Anatomy
I N C O M P U T E R I Z E D I M A G E S
Chapter 1
Introduction
T
his is a completely revised edition of an atlas designed with two goals in
mind: the first is to assist clinicians and researchers in analyzing human
neuroanatomic images obtained with computerized tomographic techniques;
the second is to serve as a tool for teaching neuroanatomy to the students who
are becoming acquainted with anatomic data either at the computer screen or
by means of radiologic transparencies rather than through the study of post-
mortem specimens. The images in the atlas depict the structures of the telen-
cephalon (cerebral cortex, basal ganglia, and thalamus) and cerebellum, but the
focus is on the cerebral cortex.
The refinement of imaging technologies such as X-ray computerized tomog-
raphy and magnetic resonance (CT and MR) has given neuroscientists and clin-
icians concerned with brain function and brain diseases access to the macroscopic
neuroanatomy and neuropathology of living humans. The amount of structural
information these technologies make available is remarkable, and both research
and clinical practice are inconceivable today without their support. Yet proper
interpretation of the information contained in tomographic scans is a difficult
task, and if the purpose of the interpretation is to produce a fine neuroanatomic
description, the reliability of the result is often problematic. There are several rea-
sons behind these difficulties. One has to do with the fact that, more often than
not, the imaging sections generated by a scanner are different from the idealized
brain sections we have committed to memory or can find in standard brain at-
lases. The difficulty is compounded by the fact that superficially the imaging sec-
tions resemble the standard brain sections and the observer can be more easily
misled. In brief, the axial incidence of section according to which most modern
scans are obtained is not the same as in standard atlases and can vary from scan
to scan. If this reservation applies to axial incidences, it applies even more strongly
to coronal incidences because they are, by convention, obtained at 90 angles from
the axial incidence chosen for a given person. Thus, coronal incidences vary from
study to study as much as axial incidences do.
[1]
One may well ask why incidences vary so much. The answer is that varia-
Human Brain Anatomy tion is unavoidable for diverse reasons: the design of the head (an incidence
I N C OM P U TE R I Z E D I M AG E S parallel to the canto-meatal line is quite different in a dolichocephalic head and
in a brachicephalic head, as shown in the first sequence of C hapter 7 and of
C hapters 8 and 9), the mobility of the spine and neck (older persons with stiffer
necks often lie on the scanning table with hyperextended heads and end up be-
ing scanned in these incidences; examples of such an incidence are shown in
the second axial sequence of C hapters 7-9), and the need to select a special
plane to target the region where a lesion is presumably located (as in a poste-
rior fossa incidence required to better visualize the cerebellum and brain stem,
shown in the third axial sequence in any of C hapters 7-9). Furthermore, inci-
dences are chosen in relation to external landmarks, most often the canto-
meatal line. U nfortunately, the relation of the canto-meatal line to brain struc-
tures is not consistent from subject to subject, as can be seen in Figure 1-1.
For practical purposes then, tomographic brain sections come in a wide
range of incidences, few of which match the textbook image of brain sections
available in traditional brain atlases. This reduces the "fit" between the section
to be interpreted and the section kept in our mind or in the library atlas. The
chance of a poor fit is further increased by the sheer ambiguity of the anatomic
features depicted in cross-sectionone sulcus looks like any other sulcus when
you see its section in a two-dimensional (2-D ) thin sliceand by the very real
difficulty of mentally transforming 2-D surfaces into 3-D volumes. Be that as
it may, this is how anatomic localization must be carried out: by taking a 2-D
section and mentally picturing its position as a component of the 3-D volume
where it belongs or by acquiring M R images with the purpose of creating 3-D
reconstructions of the brain and proceeding with direct observation and iden-
tification of sulci and gyri in that unique brain. I n brief, images that correspond
closely to the sections obtained in routine imaging procedures are not easy to
find in standard atlases.
The problems posed by the variation of scanning incidence are made worse
by an often neglected biologic reality: individual anatomic variation. When one
considers the anatomy of the human brain across a large number of individu-
als, one is struck both by the similarity and by the uniqueness of its macro-
scopic features. The similarity is easy to appreciate: we do find the same major
components in each and every specimen. All normal brains have a brain stem,
a cerebellum, and two hemispheres; and each hemisphere can be subdivided
into four lobes (frontal, parietal, occipital, and temporal), which maintain their
relative positions. M oreover, as we look at the lateral surface of every brain, we
can find two prominent sulci, the central sulcus and the sylvian fissure; and
every mesial brain surface shows us a similarly placed corpus callosum. The
similarity ends here, however. U sing plain, naked-eye analysis, we can appreci-
ate that from one individual to the next even the central sulcus and sylvian fis-
sure travel in different courses and that, even within the same individual, there
are clear differences between the two hemispheres (see Fig. 1-2). When we an-
alyze the secondary sulci within each lobe, the differences between individuals
and between the two hemispheres of the same individual become even more
pronounced. C hapters 2-5 amply demonstrate this point.
[2]
In neuroanatomy, then, there is a permanent tension between similarity
and singularity, between anatomic constants and individual variation. This ten- Introduction
sion is not a mere curiosity. It has many practical implications. Neuroscien-
tists studying normal populations or using the lesion method must take into
account similarity and singularity. Likewise, neurologists and neurosurgeons
must determine not only the pathologic nature of lesions but also where they
are with precision, especially when they need to plan an optimal surgical ap-
proach or predict future deficits and improvements. Finally, it is important to
make medical students and graduate students aware of anatomic uniqueness
because they must learn to deal with its consequences in both clinical practice
and research.
The issue of anatomic uniqueness has gained importance because of the
spectacular developments in neuroimaging technologies. As recently as the
1970s, researchers and clinicians used to make guesses and attempted to assess
the merit of those guesses by descending surgically on the living brain, scalpel
in hand, or by studying the real thing at the autopsy table. Images from CT and
MR have changed this situation radically. In particular, MR provides such faith-
ful anatomic details at such fine resolution that it has permitted the emergence
of a new field: human neuroanatomy in vivo.
The only way of eliminating the problems posed by individual anatomic
variation is to obtain a 3-D brain reconstruction for every subject and to use it
for a customized analysis of the individual's anatomy. In the many circum-
stances in which a 3-D reconstruction is either not possible or practical, how-
ever, errors of interpretation can be minimized by becoming familiar with com-
mon profiles of variation in key neuroanatomic structures. That is something
I hope to help readers achieve.
This book provides a collection of normal human brains in 3-D recon-
struction (labeled A-Z), which demonstrates a wide range of anatomic varia-
tion. It also provides a comprehensive collection of 174 sections obtained in a
single, dolichocephalic brain, using six different incidences of cut, to guaran-
tee that the same structure identified in the section of one incidence is also
present in the section from another incidence that intersects it. This reveals
unequivocally the degree to which different types of incidence alter the ap-
pearance of the same anatomic structure. Moreover, axial and coronal sections
are also included at the same incidences in two other brains, of brachicephalic
configuration, chosen for having about as different an overall shape as the
"standard" brain used in the first collection. The illustrations of feature vari-
ation as well as this latter set of sections are meant to ease problems of inter-
pretation by providing examples of brain sections obtained axially and coro-
nally at the most frequently used incidences of cut. As a whole, they also will
help the reader develop a sense of the "correction" that must be applied to the
images of some individuals in whom the typical structures do not fit the stan-
dard image.
The atlas is based on a technique developed in my laboratory, known as
Brainvox (Damasio and Frank, 1992; Frank et al., 1997). It allows for the re-
construction of the brain in three dimensions, at a level of detail compatible
with identifying all major sulci and gyri with the same degree of precision
[3]
achievable at the autopsy table. Moreover, once identified, the sulci can be traced
H u m a n B r a i n Anatomy on the 3-D image and the outline of the tracings can be transferred automati-
I N C O MP U T E R I Z E D I MA G E S cally to the original 2-D images. Because the 3-D reconstruction obtained with
Brainvox is not a surface rendering but a voxel rendering, the image of a 3-D
volume can be cut in any spatial plane; for example, axial, coronal, parasagit-
tal, oblique, and the color-coded sulci tracings from the 3-D image can be car-
ried onto the new cut, thus maintaining sulci identification. I n fact, we can go
further than we would at autopsy because we are not limited to one plane of
cut; we are free to cut and recut as necessary. A ny sulcus or gyms seen in a
2-D slice can be interrogated and studied in the 3-D image, automatically by-
passing any "guessing" process.
Brainvox has been used extensively in the study of patients with focal brain
lesions (a few examples of such lesions are given in C hapter 10). T he technique
has also been used in the preparation of neurosurgical interventions by pro-
viding high-quality anatomic images of a particular brain rendered in 3-D, with
the sulci identified and marked. I t is a helpful tool in the stereotactically guided
implantation of intracerebral electrodes, and it allows for checking of the loca-
tion of such electrodes after implantation.
Brainvox also has been used to study the detailed anatomy of the brains of
apes. E rroneous beliefs regarding the comparative anatomy of the frontal lobe
in humans and apes have been rectified on the basis of Brainvox data (S e-
mendeferi and Damasio, 2000; S emendeferi et al., 1997). Figure 1-3 shows an
example of each extant ape's brain and permits comparison with a human brain
and with the brain of a macaque. (T he mean volume and proportional values
given in the figure come from S emendeferi and Damasio, 2000, and, for the
macaque, from S emendeferi et al., 1997.)
C hapter 2 shows a variety of views of a normal dolichocephalic brain from a
fully right-handed C aucasian man, 47 years of age, a brain different from that
shown in the first edition. T he sulci are identified in one set of images and the
gyri in another. T he complete names of the different anatomic structures, fol-
lowed by the respective abbreviations used in the plates, are provided in a single
table. T he chapter also includes a set of images with the markings of Brodmann's
cytoarchitectonic regions (Brodmann 1909; 1999) superimposed on the gyri.
C hapter 3 depicts another normal brain, with a different overall configura-
tion. For this chapter, I selected the brachicephalic brain of a C aucasian woman,
also fully right-handed, 43 years of age. T his brain too is different from the
brachicephalic example of the first edition. I t helps illustrate, in C hapter 8, how
the same slice incidence can appear so different in a brain of different overall
shape.
C hapter 4 shows another brachicephalic brain, that of a 34-year-old, fully
right-handed man of A sian descent. I added this example to show how the slice
incidences shown in C hapters 7 and 8 have a very different appearance in a
shorter and wider brain. (T he slices through this brain are shown in C hapter 9.)
C hapter 5 presents a series of 26 normal brains, labeled A -Z . T hese brains
are presented in 3-D reconstruction and are seen from the left and right lateral
aspects, from the two mesial surfaces, as well as from above and below. T his
[4]
series, also different from the one presented in the first edition, allows the reader
to appreciate better the variability of sulci and gyri and to see how it accounts Introduction
both for right-left differences and for differences among individuals.
The brains in Chapter 5 are organized in five groups. The chapter begins
with the brains of eight young, fully right-handed subjects, followed by a group
of four strongly left-handed subjects and another of six subjects with mixed
handedness. Next, there is a pair of right-handed identical twins, followed by
six older fully right-handed subjects. The brains come from 12 women and 12
men; the two identical twins are female. The woman's brain is always on the
right; the matching man's brain is depicted on the left. Age, gender, and hand-
edness scores are provided for each case.
Chapter 6 is new and deals with quantitative analyses. It was prepared in
collaboration with John Allen and Joel Bruss. The chapter provides numerical
results obtained in the segmentation of large numbers of normal brains into
different lobes. It also depicts a collection of 46 brains, half male and half fe-
male, in which the different lobes were segmented and color-coded, together
with tables for the sizes of the segmented volumes.
Chapter 7 shows three different axial sequences and the corresponding coro-
nal sequences for two of them as well as one set of parasagittal slices. All come
from the dolichocephalic brain described in Chapter 2. The sulci are indicated
in the images on the left and the gyri, in those on the right. For this second edi-
tion, I chose smaller spacing of the slices, only 5 mm apart, and marked the
sulci and gyri on slices in which all nonbrain structures were removed, to in-
crease the clarity of the image. However, the "radiologic image," with the non-
brain structures in place, is visible in the upper and lower corners of each page,
along with a marking of the position of the cut as seen on the right and left
hemispheres, in both lateral and mesial views.
Chapter 8 shows the first brachicephalic brain (described in Chapter 3) in
the three axial and the two coronal incidences of Chapter 7.
Chapter 9 shows the second brachicephalic brain (described in Chapter 4)
in the same three axial incidences.
Chapter 10 provides examples of lesions seen in 3-D reconstruction and in
varied incidences of cuts.
All images in this second edition are new and were obtained in a General
Electric Signa scanner operating at 1.5 Tesla, using the following protocol:
SPGR/50, TR 24, TE 7, 1 NEX matrix 256 X 192, FOV 24 cm.
For each brain, three separate MR sequences with 1.5-1.6 mm contiguous
coronal slices were obtained and averaged post hoc using the Automated Im-
age Registration (AIR 3.03, UCLA; Woods et al., 1992), to improve resolution.
The final 3-D volume was obtained with Brainvox, which was also used to
reslice the first three brains and obtain the different sets of images. Adobe Pho-
toshop and Adobe Illustrator programs were used subsequently to label the
images and smoothen the peripheries, but I did not attempt to eliminate all
pixellation. My priority was to produce images that would reflect what is seen
in high-quality MR scans rather than to artificially smooth out images for es-
thetic purposes. All brain sections are shown according to radiologic conven-
[5]
tion, the right hemisphere appearing on the left and the left hemisphere, on
Human B r ai n Anatomy the right.
I N C O M P U T E R I Z E D I M A G E S A ll brains come from the patient and normal subject registries of the D ivi-
sion of C ognitive N euroscience at the U niversity of I owa. A ll scans were ob-
tained with informed consent, in accordance with institutional and federal
guidelines.
[6]
Figure 1-1
Transparent heads showing the left hemisphere of the brains
described in Chapters 2-4. The canto-meatal line is shown in red.
A few slices parallel to the canto-meatal line are shown to
exemplify how a plane set on the basis of external landmarks
intersects different brain regions in heads of different configura-
tion. Top: The dolichocephalic brain shown in Chapter 2.
Bottom: The two brachicephalic brains shown in Chapters 3 and
4, respectively.
[7]
Figure 1-2
Top: Sulci of the three brains discussed in Chapters 2-4
superimposed on the image of an averaged brain (based on 50
normal brains) after each of the three brains was morphed to the
averaged brain (using Woods et al., 1999 Average Image Regis-
tration). Note how the sulci remain individualized and at times
quite far from their counterpart in the other brains. Bottom: Re-
sults of the same process applied to the two identical twin brains
shown in Chapter 5 (brains S and T). Note that the superposi-
tion of sulci is better for the twins than for the three brains in
the top image. Still, the superposition is not perfect.
.human
cerebrum
FL
TL
_ErOL
insula
volume in cc % of cerebrum
1233.8_
454.8
223.2
448.8_
17.4
36,8
18.1
36,4 _
1.4
bonobo
cerebrum
FL
TL
p-OL
insula
volume in cc_% of cerebrum
_ 290.5 _ _
101.5
48.2
109.1
3,7
34.9
16.6
37.6
1.3
orangutan volume in cc % of cerebrum
cerebrum
_EL
_P-OL _
insula
391.2
142.3
65.7_
138.8
4,8
36.3
16.8_
35.5
1.2
chimpanzee volumei ncc Hofcerebrum
cerebrum 279,7
FL
_TL.
P-OL
insula
97.2
48.2_
103.6
3,1
34.8
17.2
37.0
1.1
gibbon vol umei ncc %ofcerebrum
cerebrum
FL
TL
P-OL
insula
71.6
203
13.5
28.1
0.7
28.4
18.9
39.2
1.0
gorilla
cerebrum
FL _
TL
P-OL
insula
vol umei nct %ofcerebrum
355.9
124.8 35.1
49.8 14.0
142.6 ' 39.9
5.3 1.4
Figure 1-3
I lunian and e\uni ape brai ns \viih the niiiior lobes L okii' - L oded.
L ' t iiler i-Mi.li brain are
1
t be mean valut;.^ lor iib.solnie brain .i/e and
lur iliL
1
perL enia^e oeijupied bv eaeli lobe (as report ed in
Seniendek-fi and I J.iiiiasiu, 2000; Semendefcri el al., 19971. Note
rhesus volume in cc
cerebrum 62.7
llic marked di l l e' renee in absoluie si/e and the remarkable simi-
larity ol lira in lobe proporlions among the human ,ind ape tir.it us.
Mrains L ire shown in their approximat e size reladon.
[9]
T
he brain chosen for this chapter is a typical dolichocephalic brain of a fully
right-handed, 47-year-old man. It conforms to the most commonly de-
picted global brain image, what one might call a "standard" brain. I have called
it Dolicho.
This brain is shown first from the lateral view (Figs. 2-1, 2-2), then from
the mesial view (Figs. 2-3, 2-4). Both hemispheres are seen together, the left
on the upper tier and the right on the lower tier, to allow an easy right-left
comparison of the different structures. For each hemisphere, a straight lateral
or mesial view is shown together with a downward tilt for the lateral view and
an upward tilt for the mesial view. In the latter image, the cerebellum and brain
stem have been removed to allow complete inspection of the mesio-inferior
temporal lobe.
It is important to realize how asymmetric the sulci are. The asymmetry be-
tween the right and left hemispheres is certainly not limited to the sylvian fissure,
the one structure traditionally considered asymmetric. By comparing the image
on the left, showing the sulci (Figs. 2-1, 2-3, 2-5), with the image on the right,
showing the gyri (Figs. 2-2, 2-4, 2-6), on each set of two facing pages, the reader
can easily see which sulci provide the limits for each of the gyri. These images are
followed in the upper tier by a view from the top and a view of the inferior sur-
face of the brain (the cerebellum was removed in the latter view to permit obser-
vation of the inferior surface of the temporal and occipital lobes) and in the lower
tier by images showing the frontal and occipital perspectives (Figs. 2-5,2-6). These
images are followed by the same views of the brain seen in Figures 2-2, 2-4, and
2-6 but with the approximate limits of Brodmann's cytoarchitectonic fields su-
perimposed on the colored gyri (Figs. 2-7, 2-8, 2-9). The chapter closes with two
images showing hidden structures: Figure 2-10 shows the superior aspect of the
superior temporal gyrus after removal of the frontal and parietal lobes, and Fig-
ures 2-11 and 2-12 show the insula after removal of the overlaying fronto-
parietal operculum and upper portion of the superior temporal gyrus.
Chapter 2
Exterior
Description of
a Normal
Dolichocephalic
Brain
[11]
When one looks at lateral views of the brain, two major sulci should be iden-
Human Brain Anatomy tified: (1) the central sulcus (CS), which separates the frontal from the parietal
I N CO M P U T E R I Z E D I M AG E S lobe, and (2) the sylvian fissure (SF), which separates those two lobes from the
temporal (Figs. 2-1, 2-2).
T he central sulcus is often a continuous valley with an antero-inferior course
descending from the interhemispheric fissure toward the sylvian fissure. I t usu-
ally does not reach the sylvian fissure (as in D olicho and Brachi-1), but it may
do so, as in Brain-A, the left hemisphere of Brachi-2, the right hemisphere of
Brain-K, and the left hemisphere of Brain-V. T he lower end of the sulcus con-
tinues in its main direction or makes a backward kink (compare Brain-A and
-B for an example).
T he sylvian fissure adopts an anteroposterior and mostly horizontal course,
especially in the left hemisphere, where it is usually longer. T he anterior end of
the sylvian fissure branches into two rami, which course in the frontal lobe.
T hese are known as the ascending (abSF) and the horizontal (hbSF) branches of
the sylvian fissure. T he rami may constitute two independent sulci with a com-
mon origin, as in the left hemisphere of the present brain, or have a common
trunk and then separate into individual branches, as in the right hemisphere.
T he sylvian fissure has a horizontal and long course in the left hemisphere
and terminates in this fashion (as in this brain and in Brain-F, -N , and -O ), or
it may terminate by turning down at the end (as in Brain-D , -L, and -Z ). I t may
also split in two small branches as seen in Brachi-2 and Brain-M . I n the right
hemisphere, however, it tends to have a shorter horizontal course, after which
it turns upward at the end (as in the present brain, Brachi-2, and most of the
brains shown in Chapter 5).
T he description above conforms to the classical concept we have of the right
and left SF in right-handed subjects. However, as we will see in Chapter 5, there
are exceptions (see, e.g., Brain-G , that of a fully right-handed subject). I n Brain-
G , there is a complete reversal of the course of the SF. I t is the right SF that
turns sharply down at its end and the left that shows an upward course. See
also Chapter 6 for both types of presentation.
T here are two sulci parallel to the central sulcus. Anteriorly, the precentral
sulcus (preCS) most often shows two separate segments, as in Brachi-1 (also
Brain-E , -J, -S, and -T ). T ogether with the central sulcus, the precentral sulcus
defines the precentral gyrus (preCG ). I n its posterior and superior sector, the
precentral gyrus contains Brodmann's field 4, which continues in the depth of
the central sulcus in its inferior segment. I t corresponds to the primary motor
cortex. T he anterior and inferior two-thirds of the precentral gyrus contain
Brodmann's field 6, which corresponds to the premotor cortex. (For cytoar-
chitectonic markings, see Figs. 2-7, 2-8, 2-9.)
T he postcentral sulcus (postCS) is posterior to the central sulcus and roughly
parallel to it. T his sulcus can be continuous (as in the right hemisphere of
D olicho; Brachi-1 and -2; Brain-F, -L, and -W; and the left hemisphere of Brain-
H, -I , and -J) or it may have two or three separate segments (as in the left hemi-
sphere of D olicho and Brachi-1 and the right hemisphere of Brain-J and -S).
T ogether with the CS, it defines the postcentral gyrus (postCG ), which contains
Brodmann's fields 3, 1, and 2 (the primary somatosensory cortices).
[12]
Two additional sulci should be identified: on the lateral surface of the frontal
lobe, the superior frontal sukus (SFS) and the inferior frontal sulcus (IFS). Both Normal Dolichocephalic Brain
course in a postero-anterior direction, beginning at about the precentral sulcus
but not necessarily connected to it, and are essentially parallel to each other.
The superior frontal sulcus is as often continuous as discontinuous. The best
views to examine the SFS are superior and anterior (Figs. 2-5, 2-7). In Dolicho
and Brachi-1, the SFS is discontinuous in both hemispheres, but in Brachi-2, it
is continuous in the right hemisphere and discontinuous in the left. The SFS
constitutes the lateral border of the superior frontal gyrus (SFG).
The inferior frontal sulcus is more often than not continuous (as in the left
hemispheres of the three main brains in Chapters 2-4), although it can be in-
terrupted (as in the right hemispheres of the same brains). It constitutes the
upper limit of the inferior frontal gyrus (IFG) and, together with the superior
frontal sulcus, limits the middle frontal gyrus (MFC).
The middle and superior frontal gyri share several prefrontal association
cortices in Brodmann's fields 9 (anteriorly), 8 (middle), and 6 (posteriorly).
This posterior segment is the upper expansion of premotor area 6, which we
first noted in the postcentral gyrus. The anterior segment of the middle frontal
gyrus has a separate cytoarchitectonic area, Brodmann's field 46, which is also
part of the prefrontal sector.
The inferior frontal gyrus is almost exclusively occupied by the frontal op-
erculum. The operculum is subdivided by the previously mentioned ascending
and horizontal rami of the sylvian fissure. The segment between the precentral
sulcus and the ascending ramus constitutes the pars opercularis (IFG, po) and
contains Brodmann's field 44, usually regarded as a premotor region, while the
sector between the abSF and hbSF constitutes the pars triangularis (IFG, pt).
The latter contains Brodmann's field 45, which is regarded as part of the pre-
frontal sector. This cytoarchitectonic area extends inferiorly into the upper sec-
tor of the third and most inferior component of the frontal operculum, the pars
orbitalis (IFG, porb). The inferior segment of the pars orbitalis contains the lat-
eral segment of Brodmann's field 47, also part of the prefrontal sector. With
respect to the left hemisphere, the combination of fields 44 and 45 is also known
as "Broca's area."
The very anterior tip of the frontal lobe is the frontal pole (FP). It has no
macroscopic limits and is occupied by Brodmann's field 10.
We saw that the most anterior gyrus in the parietal lobe was the postcen-
tral gyrus limited by the postcentral sulcus and containing the primary so-
matosensory cortices. In the superolateral surface of the parietal lobe, another
important sulcus must be identified: the intraparietal sulcus (IPS). The IPS has
an anteroposterior course beginning at the upper half of the postcentral sulcus.
It can be continuous or interrupted and is best seen in superior and posterior
views (Figs. 2-5, 2-7). It separates the parietal lobe into two lobules. The first
is the superior parietal lobule (SPL), which is occupied by the somatosensory as-
sociation cortex of Brodmann's field 7. Squeezed in between this field and the
superior sector of the primary somatosensory cortices is another somatosen-
sory association cortex, Brodmann's field 5. The second is the inferior parietal
lobule (IPL), itself subdivided into two independent gyri, the supramarginal
[13]
( S M G ) anteriorly and the angular ( AG ) posteriorly. The angular gyrus may be
H u m a n B r a i n Anatomy seen as the continuation of the middle temporal gyrus into the parietal lobe
I N C O M P U TE R I Z E D I M AG E S and is constituted of multimodal association cortex ( Brodmann's field 39) ,
while the supramarginal gyrus sits above the superior temporal gyrus. The lat-
ter is also multimodal association cortex and contains Brodmann's field 40.
The angular gyrus and the superior parietal lobule are contiguous with the
lateral occipital gyri, which in humans are occupied by visual association cor-
tices ( Brodmann's fields 19 and 18) . There is no sulcal separation between the
parietal and the occipital lobe on the lateral surface of the brain.
Let us return now to the sylvian fissure and the temporal lobe. P arallel and
inferior to the sylvian fissure are two sulci, seen on the lateral surface: the su-
perior temporal ( S TS ) and the inferior temporal ( I TS ) . The superior temporal
sulcus can be continuous, as in the three main brains and in most of the brains
in C hapter 5, or divided into two or three segments, as in Brain-C and -R . I t is
usually readily identifiable and forms the lower limit of the superior temporal
gyrus ( S TG ) , also known as the first temporal gyrus. The lateral aspect of the
gyrus is occupied by Brodmann's field 22, an auditory association cortex. The
superior surface of the gyrus, hidden by the frontal and parietal opercula and
accessible to inspection only when the frontal and parietal lobes are lifted ( Figs.
2-10, 3-7, 4-7) , is occupied by the transverse temporal gyrus or Heschl's gyrus
( H G ) , which contains the primary auditory cortex ( Brodmann's fields 41 and
42) and is limited by the transverse temporal sulci ( trTS ) . H eschl's gyrus is most
often a single gyrus, but it can be double or partially divided in two. A double
H G is found more often in the right hemisphere. The segment of the superior
temporal gyrus posterior to H eschl's gyrus is the planum temporale ( P T) , and
in the left hemisphere it is part of Wernicke's area. The sector anterior to the
H eschl's gyrus is the planum polare. Both plana are constituted by auditory as-
sociation cortex ( Brodmann's field 22) .
The inferior temporal sulcus is more difficult to identify because of its many
segments. H owever, both the left hemisphere of D olicho and the right hemi-
sphere of Brain-E display a long and continuous inferior temporal sulcus. To-
gether with the superior temporal sulcus the I TS limits the middle temporal
gyrus ( M TG ) or second temporal gyrus, which is occupied by higher-order as-
sociation cortex in Brodmann's field 21 ( anteriorly) and by Brodmann's field
37 ( posteriorly) . The anterior tip of the temporal lobe constitutes the temporal
pole ( TP ) or field 38; as with the frontal pole, there are no macroscopic ana-
tomic landmarks to demarcate the posterior limit of the polar region. The pos-
terior end of the temporal lobe is found at the temporo-occipital incisura, at
which a vertical sulcus, the anterior occipital sulcus ( AO S ) separates the tempo-
ral from the occipital lobe.
C ontinuing into the inferior and mesial surfaces of the temporal lobe ( see
Figs. 2-2, 2-3, 2-5) , we find yet another anteroposterior sulcus: the temporo-
occipital ( TO S ) , which can be continuous, as in Brain-F, the right hemisphere
of Brain -I and -Q, and the left hemisphere of Brain-A, or most often separated
into two, three, or even four segments. I t marks the mesial limit to the inferior
temporal gyrus ( I TG ) or third temporal gyrus. The lateral limit of this gyrus is
the inferior temporal sulcus. This gyrus is occupied by higher-order visual as-
[ 1 4 ]
sociation cortex: Brodmann's field 20 anteriorly and the continuation of field
37 posteriorly. Normal Dolichocephalic Brain
Mesial to the TOS, the most notable sulcus is the collateral (ColS). This sul-
cus marks the mesial limit of the fourth temporal gyrus, also known as the
temporo-occipital gyrus (TOG) or fusiform gyrus of the temporal lobe. As was the
case with the previous gyrus, it holds Brodmann's fields 20 and 37. The collat-
eral sulcus also constitutes the lateral limit of the parahippocampal gyrus
(paHG), or fifth temporal gyrus, largely occupied by Brodmann's field 28 (the
entorhinal cortex). Buried inside the collateral sulcus are some important cy-
toarchitectonic fields, namely, fields 35 laterally and 36 mesially, which can
come to the surface of the posterior segment of the parahippocampal gyrus and
all along the mesial edge of the fourth temporal gyrus, respectively. The ante-
rior and mesial sector of the parahippocampal gyrus is occupied by field 34 and
the posterior and mesial sector, by field 27. The anteriormost tip of the tem-
poral lobe in both the inferior and mesial views forms the temporal pole (Brod-
mann's field 38, as mentioned earlier).
The collateral sulcus and the temporo-occipital sulcus can continue poste-
riorly into the occipital lobe and respectively limit the lingual (LingG) and
fusiform (FusiG) gyri, which contain early visual association cortex (Brodmann's
fields 18 and 19).
The calcarine sulcus or calcarine fissure (CalcF) is the most important sulcus
in the mesial view of the occipital lobe (Figs. 2-3, 2-4). This sulcus is extremely
variable in its course and configuration (see mesial views of brains in Chapter
5). It can be extraordinarily different in the right and left hemispheres of the
same individual (as in Dolicho) or fairly similar (as in Brachi-1 and -2). Ante-
riorly, the calcarine joins another major sulcus that separates the occipital from
the parietal lobe, the occipitoparietal sulcus (OPS) and then continues its ante-
rior course. Posteriorly, the CalcF can terminate inside the mesial surface of the
hemisphere (see Brain-D) or extend to the pole and even to the outer surface
(as in the present case, Fig. 2-5). It can terminate in a single straight line or,
more typically, in a T or Y shape (see Brain-F and -T, or the left hemisphere
of -H, -K, and -L). The primary visual cortex is mostly buried in the depth of
the calcarine sulcus, and it overflows to a variable degree onto the inferior and
superior banks of the calcarine region. It corresponds to Brodmann's field 17.
Between the calcarine sulcus and the occipitalparieto sulcus rests the cuneus
(Cun), occupied by early visual association cortex (part of Brodmann's fields
18 and 19).
Apart from the sulci described above, the most notable sulcus in the brain's
mesial surface is the cingulate (CingS). It is parallel to the corpus callosum (CC)
and to its limiting sulcus, the callosomarginal sulcus. It can be continuous, as in
the left hemisphere of Dolicho and the right hemisphere of Brachi-1 and -2, or
interrupted, as in one or the other hemisphere of several brains in Chapter 5.
It is not unusual to see a double cingulate sulcus, as in the right hemisphere of
Dolicho or the left hemisphere of Brachi-1 and -2. The posterior end of the cin-
gulate sulcus bends upward in an anterior concave shape, creating the ascend-
ing ramus of the cingulate sulcus (arCingS). Another short sulcus can be found
posteriorly to the ascending branch and still parallel to the corpus callosum,
[15]
the subparietal sulcus (sPS), which is most often separate but can also be con-
Human Brain Anatomy tinuous with the cingulate sulcus (see Brain-B). The cingulate sulcus and the
I N C O M P U TE R I Z E D I M AG E S subparietal sulcus form the superior border of the cingulate gyrus. This is lim-
bic cortex, Brodmann's field 24 in the anterior half and field 23 in the poste-
rior. Further back, in the superior section of the posterior half, there is also
Brodmann's field 31 and even further back, in the retrosplenial region (Sp), there
are Brodmann's fields 26, 29, and 30. Above the subparietal sulcus and between
the parieto-occipital sulcus (posteriorly) and the ascending branch of the cin-
gulate sulcus (anteriorly) lies the mesial parietal region or precuneus (preC un).
This is somatosensory association cortex (the continuation of Brodmann's field
7). The most anterior sector contains Brodmann's field 5. Buried anteriorly in
the callosomarginal sulcus, we find Brodmann's field 33.
The ascending branch of the cingulate sulcus usually terminates just behind
the superior and mesial extension of the central sulcus and is the landmark of
choice to find the mesial extension of the central sulcus. Just anterior to the
central sulcus we usually find a more or less vertical sulcus, the paracentral
(paraC S), which arises either as a side branch of the cingulate sulcus, as in
Brachi-1 and the left hemisphere of Brachi-2, or as a sulcus descending from
the outer edge of the interhemispheric space, as in D olicho and the right hemi-
sphere of Brachi-2. The region, limited by the paracentral sulcus, cingulate sul-
cus, and ascending branch of the cingulate sulcus is called the paracentral lob-
ule or paracentral gyrus (paraC G ), which is mostly occupied by primary motor
cortex or field 4 of Brodmann. The most posterior sector, behind the edge of
the central sulcus, is occupied by the extension of the primary somatosensory
cortices, Brodmann's fields 3, 1, and 2.
Anterior to the paracentral sulcus and limited inferiorly by the cingulate sul-
cus lies the mesial surface of the superior frontal gyrus. The major cytoarchi-
tectonic component is the large expansion of field 6, premotor cortex, which
on the medial surface constitutes the supplementary motor area. Fields 8 and 9
sit just in front. Again, the tip of the frontal lobe corresponds to the frontal
pole. Just in front of field 24, in a narrow area parallel to it, is Brodmann's
field 32.
U nder the lower portion of the inferior and anterior end of the cingulate
sulcus lies the mesial portion of the orbital region of the frontal lobe. I n the
mesial view, we can recognize a small anteroposterior sulcus, the suborbital sul-
cus (SO rbS). I n the inferior view, the most mesial, anteroposterior sulcus is the
medial orbital sulcus (mO rbS) or olfactory sulcus, which marks the lateral bor-
der of the gyrus rectus (G rec). Lateral to this sulcus we find a host of orbital
sulci forming a pattern reminiscent of the letter H and usually known as H-
shaped sulci (HsS). These sulci help define the anterior, posterior, medial, and
lateral orbital gyri (aO rbG , pO rbG , mO rbG , and lO rbG , respectively). Brod-
mann's field 11, 12, 13, 14, and 15 occupy this region. Brodmann's field 25 is
located in the most posterior and mesial region of the orbital surface of the
frontal lobe.
The insula is completely buried under the frontoparietal opercula and the
anterior and superior sector of the superior temporal gyrus. When these are ex-
cised, a large oval shaped region of cortex is made visible (see Figs. 2-11, 2-12).
[16]
The insula is limited by the circular sulcus. The insula is divided into anterior
and posterior sectors. Located anteriorly we find the anterior or short insular Normal Dolichocephalic Brain
sulci, which serve as limits to the anterior or short insular gyri; posteriorly we
find the posterior or long insular suki, which limit the posterior or long insular
gyri.
In Figures 2-7,2-8, and 2-9, the dotted lines delineate the approximate lim-
its of Brodmann's cytoarchitectonic regions. It is important to realize that no
visible landmarks correspond to these borders. All we can currently do is ven-
ture on approximations, which continues to rely on the informative maps of
Brodmann and his followers (e.g., Sarkissow et al., 1955; Braak, 1978). Fortu-
nately, this task is made easier by the fact that some Brodmann's areas do fol-
low sulcal patterns. As a result, talking about areas 40 and 39 is equivalent to
talking about the supra-marginal and the angular gyrus, and mentioning area
37 is certainly more efficient than referring to the posterior sector of the mid-
dle, inferior and 4th temporal gyri.
As a complement to the reading of this chapter, I recommend Duvernoy
(1999), an anatomic description of postmortem brains, and Ono et al. (1990),
a detailed study of sulcal variability.
[17]
Figure 2-1
Lateral views of Dolicho with marked sulci. See text for details.
[18]
Figure 2-2
Lateral views of Dolicho with marked gyri. See text for details.
[19]
Figure 2-3
Mesial views of Dolicho with marked sulci. See text for details.
[20]
Figure 2-4
Mesial views of Dolicho with marked gyri. See text for details.
[21]
Figure 2-5
Bottom and top views (top) and front and back views (bottom)
of Dolicho with marked sulci. See text for details.
[22]
Figure 2-6
of Dolicho with marked gyri. See text for details.
[23]
Figure 2-7
Lateral views of Dolicho with Brodmann's cytoarchitectonic
areas marked on the same views seen in Figure 2-2. See text for
details.
[24]
Figure 2-8
Mesial views of Dolicho with Brodmann's cytoarchitectonic ar-
eas marked on the same views seen in Figure 2-4. See text for
details.
[25]
Figure 2-9
of Dolicho with Brodmann's cytoarchitectonic areas marked on
the same views seen in Figure 2-6. See text for details.
[26]
Figure 2-10
Unroofing of the superior temporal gyrus. The frontal and pari-
etal lobes were removed above the sylvian fissure and above a
straight line drawn from the end of the sylvian fissure to the edge
of the brain (top), according to the method described by
Geschwind and Levitzky (1968). Observation from above (bot-
tom) allows inspection of the superior surface of the STG. A
transverse running sulcus and gyrus are prominently seen. The
transverse temporal sulcus is on the bottom l e f t. The transverse
gyrus of Heshl is colored red on the bottom right. It corresponds
roughly to Brodmann's fields 41/42, the primary auditory cor-
tices. Behind HG is a triangular region, colored green. This is
known as the planum temporale, and corresponds to auditory
association cortex, Brodmann's field 22, as does the rest of the
superior temporal gyrus.
[27]
Figure 2-11
Image of the left and right insula after portions of the frontal,
parietal, and temporal lobes which cover it have been removed.
Sulci are marked. See text for details.
[28]
Figure 2-12
Image of the left and right insula after portions of the frontal,
parietal, and temporal lobes covering it have been removed. Gyri
are marked. See text for details.
[29]
Chapter 3
Exterior
Description of a
Brachicephalic Brain
A nother normal brain is depicted in this section, that of a 43-year-old, right-
jL~Vhanded Caucasian woman. The overall configuration of this brain is quite
different from the one in the previous chapter. It is a brachicephalic brain, and
I refer to it as Brachi-1. A s with Dolicho, I begin at the outer surface and iden-
tify the major sulci (Figs. 3-1, 3-3, 3-5). Figures 3-2, 3-4, and 3-6 show the
gyri. Figure 3-7 shows both the sulci and the gyri in the superior surface of the
superior temporal gyrus. The description of the sulci and gyri in Chapter 2 en-
tirely applies here. The images in this chapter should be compared with the cor-
responding images in the previous chapter. No images of cytoarchitectonic fields
are shown for this brain.
[31]
Figure 3-1
Lateral views of Brachi-1 with marked sulci. See Chapter 2 for
details.
[32]
Figure 3-2
Lateral views of Brachi-1 with marked gyri. See Chapter 2 for
details.
[33]
Figure 3-3
Mesial views of Brachi-1 with marked sulci. See Chapter 2 for
details.
[34]
Figure 3-4
Mesial views of Brachi-1 with marked gyri. See Chapter 2 for
details.
[35]
Figure 3-5
of Brachi-1 with marked sulci. See Chapter 2 for details.
[36]
Figure 3-6
Bottom and top views (top] and front and back views (bottom)
of Brachi-1 with marked gyri. See Chapter 2 for details.
[37]
Figure 3-7
Unroofing of the superior temporal gyrus. See Chapter 2 and leg-
end of Figure 2-10 for details.
[38]
A nother brachicephalic brain, Brachi-2, is shown in this chapter. This is the
JLJLbrain of a 34-year-old, fully right-handed man of Asian descent. The rea-
son for depicting a second brachicephalic brain is that brachicephalic brains
vary quite considerably among themselves. The difference between Dolicho and
Brachi-2 is even more pronounced than that between Dolicho and Brachi-1.
The overall configuration shows a shorter anterior-posterior diameter and a
larger left-right diameter, best appreciated by comparing Figures 4-5 and 4-6
with Figures 2-5 and 2-6. Compare also to Figures 3-5 and 3-6.
The external morphologic description remains the same, but note how the
different overall configuration leads to different structures appearing in the
slices obtained through this brain (see Chapter 9). In spite of these differences,
however, the sulci and gyri are the same as described in Chapter 2 and the pro-
portions of the brain, lobes, and gyri are the same as those in the Caucasian
brains depicted in Chapter 6.
This brain shows a good example of a partially doubled Heschl's gyrus (see
Fig. 4-7). Also, note that in the right hemisphere there is a clear middle frontal
sulcus> (MFS), which divides the middle frontal gyrus in two.
Chapter 4
Exterior
Description
of Another
Brachicephalic
Brain
[39]
Figure 4-1
Lateral views of Brachi-2 with marked sulci. See text of Chapter
2 for details.
[40]
Figure 4-2
Lateral views of Brachi-2 with marked gyri. See text of Chapter
2 for details.
[41]
Figure 4-3
Mesial views of Brachi-2 with marked sulci. See text of Chapter
2 for details.
[42]
Figure 4-4
Mesial views of Brachi-2 with marked gyri. See text of Chapter
2 for details.
[43]
Figure 4-5
of Brachi-2 with marked sulci. See text of Chapter 2 for details.
[44]
Figure 4-6
of Brachi-2 with marked gyri. See text of Chapter 2 for details.
[45]
Figure 4-7
Unroofing of the superior temporal gyrus. See text of Chapter 2
and legend of Figure 2-10 for details.
[46]
C
hapter 2 provided a description of the different sulci on the brain's lateral
and mesial surfaces based on the most common appearance of those struc-
tures. The purpose of this chapter is different. Rather than concentrating on
"average" neuroanatomic design, I emphasize the large range of variation that
entirely normal structures may exhibit. Neuroanatomic interpretations are un-
likely to be accurate unless such diversity is taken into account.
This chapter is made up of a collection of normal adult brains, labeled A-Z.
Each brain is depicted in its lateral and mesial surfaces, seen from the right and
left, from above and below. The reader should inspect these brains and appre-
ciate the large variance among individual specimens and even between the two
hemispheres of the same brain. Not all sulci were identified so that the surfaces
could be left uncluttered but the main landmark sulci highlighted. Unmarked
sulci should be easy to recognize using the description provided in Chapter 2.
Most of the brains depicted in this chapter belong to persons below age 50
(range 18-50, mean 29 years). The last six belong to persons 61-79 years of age
(mean 70). As was to be expected, the sulci in the latter group are wider than
those in the younger group. However, there are some remarkable exceptions.
Brain W (of a 75-year-old woman) has notably tight sulci, not much different
from those of brain D (of a 22-year-old woman). On the other hand, brain K
(of a 50-year-old man) shows remarkably wide sulci, not that different from
those in the majority of older subjects.
Half the brains are from women (always depicted on the right) and half
from men (depicted on the left). Brains S and T are from women that happen
to be identical twins. The pairs were matched, as well as possible, for handed-
ness and age, providing a realistic comparison of the morphology of male and
female brains. Note, for instance, how the gender difference that has been re-
ported for the splenium, the posterior sector of the corpus callosum, can eas-
ily be challenged. Compare the splenium in brains C and D (of a man and a
woman, respectively): neither shows the "typical" enlargement attributed to the
Chapter 5
An Alphabet of
Normal Brains
[47]
female brain, although both brains M and N (also of a man and a woman, re-
Human B r ai n Anatomy spectively) show such an enlargement. Comparison of brains O and P or U and
I N COMPUT E R I Z E D I MA G E S V shows the bulging of the splenium in men but not women. T he reported dif-
ference can be seen in brains A and B as well as in H and G . Out of the 12 fe-
male brains, six show bulging of the splenium and six do not; out of the 12
male brains, four show splenial bulging and eight do not. T he twin brains, T
and S (both women), show bulging of the splenium. A mixture of these pre-
sentations can also be seen in Chapter 6. For example, the pair 14F and 14M
presents with a typical bulging pattern, while the pair 7F and 7M shows more
bulging in the man.
No systematic difference between the genders can be found on inspection
of the sulcal pattern. What is remarkable in the sulcal pattern is the variability
among subjects, even between the right and left hemispheres of the same per-
son. Marked differences can be appreciated even in the major sulci and all the
more so in secondary or tertiary sulci. T he fact that the sulci in the two twin
brains are much closer to each other than any other two brains in the collec-
tion is interesting. However, equally interesting is the fact that even the sylvian
fissure and the central sulcus, two sulci that develop early, are quite different
in these two persons whose genetic makeup is the same. (S ee Figure 1-2 to ap-
preciate the lack of superposition of these sulci when the brains are morphed
to the same space.)
One other purpose of this chapter is to compare brains according to hand-
edness. T he chapter begins with brains from extreme right-handers, brains A -H
(handedness scores of+ 85 to +1000 in the Oldfield-G eschwind questionnaire).
I t continues with four brains from extreme left-handers, brains I-L (scores of
80 to 100) and six brains of subjects with mixed handedness, brains M-R
(scores of 3 0 to +65). Both twins are fully right-handed, as are the six older
subjects.
T he feature of external morphology most connected to right handedness
(and, by extension, with left hemisphere dominance for language) is the course
of the sylvian fissure. We expect it to be longer and to terminate with a down-
ward turn in the left hemisphere and to be shorter and to have a terminal up-
swing in the right hemisphere. S uch a typical sylvian fissure can be seen in brains
D , Y, and Z , all strong right-handed subjects; but it is also found in brain K,
that of a left-handed man (80), and in brain N, that of a woman with mixed
handedness ( 3 0). T he reverse pattern might be expected in strong left-
handers, but that is not the case in the exemplars at hand. T he remaining three
strong left-handers (brains I , J, and L) have rather symmetrically coursing syl-
vian fissures, as happens with most mixed-handedness cases. T here is a very
good example of a reversal of the sylvian fissure pattern (downward turn on
the right and upward turn on the left) in brain G , that of a fully right-handed
man ( + 100), as well as in several of the brains seen in Chapter 6, where all sub-
jects are strongly right-handed; see for instance the pair 17 F and 17M). A lso,
one of the most typical right-left asymmetry patterns in the whole of this chap-
ter is that in brain O, a man of mixed handedness.
[48]
Chapter 5 Figures
[49]
Figure 5-1
Brain A. Right-handed (+90) man, 27 years of age.
[50]
Figure 5-2
Brain B. Right-handed (+90) woman, 27 years of age.
[51]
Figure 5-3
Brain C. Right-handed ( + 100) man, 24 years of age.
[52]
Figure 5-4
Brain D. Right-handed (+85) woman, 22 years of age.
[53]
Figure 5-5
Brain E. Right-handed (+90) man, 20 years of age.
[54J
Figure 5-6
Brain F. Right-handed ( + 100) woman, 18 years of age.
[55]
Figure 5-7
Brain G. Right-handed ( + 100) man, 32 years of age.
[56]
Figure 5-8
Brain H. Right-handed (+90) woman, 28 years of age.
[57]
Figure 5-9
Brain I. Left-handed (-100) man, 42 years of age.
[58]
Figure 5-10
Brain J. Left-handed (-100) woman, 39 years of age.
[59]
Figure 5-11
Brain K. Left-handed (-80) man, 50 years of age.
[60]
Figure 5-12
Brain L. Left-handed (-95) woman, 50 years of age.
[61]
Figure 5-13
Brain M. Mixed-handed ( 30) man, 28 years of age.
[62]
t
Brain N. Mked-handed ( 30) woman, 26 years of age.
[63]
Figure 5-15
Brain O. Mixed-handed ( 2 5 ) man, 19 years of age.
[64]
Figure 5-16
Brain P. Mixed-handed (+35) woman, 31 years of age.
[65]
Figure 5-17
Brain Q. Mixed-handed (+65) man, 29 years of age.
[66]
Figure 5-18
Brain R. Mixed-handed (+40) woman, 22 years of age.
[67]
Figure 5-19
Brain S. Right-handed ( + 100) woman, 23 years of age. Identical
twin of the woman in Figure 5-20.
[68]
Figure 5-20
Brain T. Right-handed ( + 100) woman, 23 years of age. Identi-
cal twin of the woman in Figure 5-19.
[69]
Figure 5-21
Brain U. Right-handed ( + 100) man, 73 years of age.
[70]
Figure 5-22
Brain V. Right-handed ( + 100) woman, 68 years of age.
[71]
Figure 5-23
Brain W. Right-handed ( + 100) man, 75 years of age.
[72]
Figure 5-24
Brain X. Right-handed (+100) woman, 79 years of age.
[73]
Figure 5-25
Brain Y. Right-handed (+95) man, 61 years of age.
[74]
Figure 5-26
Brain Z. Right-handed ( + 100) woman, 63 years of age.
[75]
T
his chapter demonstrates that, in addition to the qualitative analysis shown
in the previous sections, modern imaging techniques can be used for
the quantitative investigation of brain structure. Three-dimensional (3-D) mag-
netic resonance imaging (MRI) allows us to identify and measure surface brain
structures, deep structures can be delineated using a combination of two-
dimensional (2-D) slice orientations, and a number of new techniques permit
the quantitative study of structural MRIs: for example, the automated segmen-
tation of MRIs into gray matter (GM), white matter (WM), and cerebral spinal
fluid (CSF) provides quantitative information about the global and regional
composition of the brain (Fig. 6-1) (Grabowski et aL, 2000).
Useful quantitative data on the structure of the brain can be obtained with
the relatively small number of metrics relevant for analyzing MR images. These
metrics include linear and angular measurements of surface structures, such as
sulci, and volumetric measurements of any specific three-dimensional structure
or region of interest (ROI). Automated tissue segmentation allows us to deter-
mine GM and WM volumes of structures and to calculate the ratio of the two
in an ROI or the gray matter density of a region (GM/[GM + WM]).
Quantitative analysis of brain structures with MRI has become a valuable re-
search tool over the past decade. However, when reading this literature, it is im-
portant to keep in mind that such analysis of brain MR images is not standardized.
Comparisons across studies conducted by different research groups need to take
into account factors such as voxel size and slice thickness, which determine the spa-
tial resolution of the MR images. Quantitative studies also employ different sam-
pling and parcellation methods, which may contribute to variability in the results
obtained by different groups. Given the rapid pace of technological development
in the field and the vagaries of anatomic parcellation, methodologic issues are of
particular importance in meta-analyses of MRI studies (e.g., Brierly et al., 2002).
In this chapter, we present results based on extensive quantitative studies of
brain size conducted in 87 healthy, right-handed adults between the ages of 22 and
Chapter 6
Quantifying
Neuroanatomic
Differences
In collaboration with
John S. Allen and Joel Bruss
[77]
88 years (44 men and 43 women) and in 25 congenitally deaf adults (Allen et al.,
2002,2003a-c; Emmorey et al., 2003). These studies demonstrate the value of MRI
for understanding how brain size and the proportional composition of the brain's
major lobes and sectors vary according to gender, age, disease, or congenital ab-
normality. Data from these studies allow us to address basic questions about the
development, evolution, and function of the human brain (Allen et al., 2004a).
A Comment on Automated Methods for Quantitative Magnetic
Resonance Imaging Research
Given that tracing ROIs on contiguous MRI slices is such a laborious undertak-
ing (see below), it is not surprising that various researchers have tried to develop
automated methods to parcellate the brain (for examples of automated methods,
see Nopoulos et al., 2000; Good et al., 2000; Sowell et al., 2003). These automated
methods ease the task of data collecting, although there is an inevitable trade-off
in loss of precision. Brain structures are not identified by anatomic landmarks
but, rather, according to statistical brain maps. Another attraction of automated
methods for determining regional brain volumes (or to compare relative volumes
across subject groups) is that they allow investigators with little neuroanatomic
expertise to conduct neuroanatomic research. While there is nothing inherently
wrong with such facilitation, there are often unforeseen consequences when a re-
searcher depends on the output from a "black box" at a critical step in data anal-
ysis. Neuroimaging researchers need to keep in mind that the MRI "brains" they
analyze are highly processed digital signals rendered into virtual structures, and
anatomical knowledge is of the essence to perform anatomically based research.
One of the main drawbacks of automated methods for volumetric brain re-
search is that all require resizing brains into a common anatomic space before
analysis can be done (e.g., Tailarach and Tournoux, 1988). Although resizing
serves to "control" for brain size in some comparative contexts, the resizing it-
self may attenuate differences between groups of interest. Furthermore, resiz-
ing methods may treat different regions of the brain differently, which may dis-
tort regional volume relationships both within and between individual brains;
there is the possibility for significant errors when the volumetric analysis is car-
ried out in resized rather than "original" brains (Allen et al. 2004b). Norma-
tive data on brain volume are becoming more widely available in publications
and via sources such as the Internet Brain Volume Database (www.cma.mgh.
harvard.edu/ibvd/). This database, derived from more than 80 publications,
provides a continuously updated source of volumetric MRI data organized by
anatomic region, age, sex, and pathology. Since brains vary by size and tissue
composition (i.e., amount of GM versus WM) between the sexes and across the
life span, it is essential to develop an accurate picture of normal variation be-
fore investigating changes in brain size or composition that might be associated
with various diseases or conditions. As mentioned above, however, investiga-
tors need to be wary about the methodologic sources of variability (e.g., have
the brains been resized before quantification?) when examining patterns of bi-
ologic variability obtained by combining data from different research sources.
[78]
Human Brain Anatomy
IN COMPUTERI ZED I MAGES
Method and Rules for Pa reel lat ion of the Major Lobes
The process of dividing brain structures into various regions is called parcella-
tion. The ultimate goal of parcellation in MRI research is to identify and delin-
eate neuroanatomic regions that are of functional, developmental, pathologic,
or evolutionary significance. Different researchers have used different methods
to obtain brain parcellations. Here, we describe the principles used in our lab-
oratory. For ROIs that are fully or partially bounded by structures seen on the
surface of the hemisphere, parcellation proceeds first by marking the limiting
structures on the surface of the 3-D rendered brain; these markings are trans-
ferred automatically to 2-D slices of the brain (Frank et al, 1997). Tracing of
ROIs is done by hand on contiguous slices. The area traced in each ROI is mul-
tiplied by the slice thickness to determine a slice volume. Then, the volumes of
each slice are summed to create a volume for the 3-D ROI structure. To main-
tain a consistent orientation for tracing ROIs, all brains are realigned into a com-
mon position. Consistent position is necessary to ensure that those ROI limits
that have to be set using "arbitrary" lines and planes will be set in the same fash-
ion across different brains. Consistent position also helps improve intertracer
reliability since brain structures appear quite differently in different slice orien-
tations. The brains are reoriented so that the coronal planes are perpendicular
to a plane containing the anterior and posterior commissures (the ac-pc plane).
The criteria followed in our laboratory for the parcellation of the frontal, tem-
poral, parietal, and occipital lobes, along with the insula and cingulate gyrus, are
presented below (see Fig. 6-2). The quantitative volumetric results of this parcel-
lation applied to 46 healthy, right-handed adults have been presented fully in Allen
et al. (2002, 2003b) and are also discussed in subsequent sections of this chapter.
Frontal Lobe
The major sulcal boundaries of the frontal lobe are the sylvian fissure, the cen-
tral sulcus, and the cingulate sulcus. The superoposterior boundary on the
mesial surface is the ascending branch of the cingulate sulcus and the mesial
extension of the central sulcus. Since these sulci do not join, an arbitrary line
linking the two is drawn along the surface of the brain in the parasagittal cut,
in which the end of the central sulcus is seen. The cingulate gyrus and insula
are excluded from the frontal lobe, and, because the ascending branch of the
cingulate sulcus is posterior to the mesial termination of the central sulcus, a
small sliver of the mesial portion of the postcentral gyrus (parietal lobe) is nec-
essarily included in the frontal lobe ROI.
Temporal Lobe
The temporal lobe is bounded by the sylvian fissure superiorly and the parahip-
pocampal fissure mesially. On the mesial surface, the posterior boundaries are
formed by the occipital cut (see Occipital Lobe, below) and an arbitrary line
drawn from the junction of the occipitoparietal and calcarine sulci to the most
inferior point of the splenium of the corpus callosum. On the lateral surface,
[79]
Quantifying Neuroanatomic
Differences
the boundary between the temporal and occipital lobes is formed by the oc-
cipital cut. An additional limiting line is drawn from the end of the sylvian
fissure to a point along the lateral portion of the occipital cut. This point is
one-quarter of the distance from the preoccipital notch (point 3) to the occip-
itoparietal sulcus (point 1), as measured along the surface of the brain in the
coronal slice containing the occipital cut. This line forms the boundary between
the temporal and parietal lobes.
Occipital Lobe
The occipital lobe has no "natural" sulcal boundaries. We consider it limited
anteriorly by the occipital cut (see below), with the exception of the super-
omesial boundary, where instead the occipitoparietal sulcus is used to separate
the occipital from the parietal lobe.
The occipital cut forms the anterior boundary of the occipital lobe and the pos-
terior boundaries of the parietal and temporal lobes. It is a plane defined by the fol-
lowing three points: the superior end of the occipitoparietal sulcus (point 1) and
the junction of the occipitoparietal and calcarine sulci (point 2) on the mesial sur-
face and the preoccipital notch on the lateral surface (point 3). This oblique plane
is automatically rendered as a slice by the Brainvox image analysis program (Frank
et al., 1997). On this oblique slice, the hemisphere being parcellated is traced, gen-
erating the limiting line on the surface of the brain that corresponds to the "cut."
In coronal slices, this line appears as two points on the surface of the brain.
Parietal Lobe
The boundaries are the central sulcus, sylvian fissure, ascending branch of the
cingulate sulcus, occipital cut, and the arbitrary line separating it from the tem-
poral lobe as described above. The parietal lobe is separated from the posterior
cingulate by a line drawn from the origin of the ascending branch of the cin-
gulate sulcus to the junction of the occipitoparietal and calcarine sulci.
Cingulate Gyrus
We treat the cingulate gyrus as a separate structure instead of including it under
the frontal and parietal lobes. The cingulate gyrus is bounded by the cingulate sul-
cus and the callosal sulcus. The inferior boundary of the anterior cingulate gyrus
is formed by a line drawn from the antero-inferior end of the sulcus to the poste-
rior point of the rostrum of the corpus callosum (this corresponds to the antero-
inferior end of the callosal sulcus). The posterior cingulate is separated from the
parietal and temporal lobes by the two arbitrary lines described above. In cases of
double cingulate sulci, the outer sulcus is chosen as the boundary for the gyrus.
Insula
We also treat the insula as a separate structure. The insula is defined by the cir-
cular sulcus, which can be clearly seen in coronal cuts. Anteriorly, the insula
may appear as a small amount of gray matter embedded in the surrounding tis-
sue of the frontal lobe. The insula is separated from the rest of the hemisphere
Human Brain Anatomy
IN COMP UT E RI Z E D I MAGES
[80]
by a line linking the deepest extent of both ends of the circular sulcus. When
the claustrum becomes visible, the line is edited to exclude it.
Proportions of the Major Lobes
The parcellation of a group of 46 healthy, right-handed individuals between the
ages of 22 and 49 years (Figs. 6-3 to 6-10) indicates that there are no sex dif-
ferences in the proportional representations of the major lobes (lobe volume
divided by hemisphere volume) and that volumetric variation is consistently
greater than variation in proportions (Allen et al., 2002). In both men and
women, the frontal lobes comprise about 38% of the hemispheres (range
36%-43%), the temporal lobe about 22% (range 19%-24%), the parietal lobe
about 25% (range 21%-28%), and the occipital lobe about 9% (range
7%-12%); the cingulate gyrus comprises about 4.3% of the left hemisphere and
3.8% of the right, reflecting a sulcal asymmetry that will be discussed below.
Although there is not major proportional variability for any of the princi-
pal lobes, individual variation in lobe proportions does occur. For example, af-
ter controlling for brain size, there is a significant negative correlation between
frontal and parietal lobe volumes: people with large parietal lobes tend to have
small frontal lobes and vice versa. Subjects 11F (Fig. 6-6) and 6M (Fig. 6-5)
have proportionally large frontal and small parietal lobes. A similar but weaker
inverse relationship can also be seen between the temporal and occipital lobes
(e.g., subject 1M in Fig. 6-3). The inverse relationship between the frontal and
parietal lobes suggests that the position of the central sulcus, which is one of
the earliest sulci to appear in development, is a critical determinant of the rel-
ative sizes and proportions of these lobes. Theories that suggest relative frontal
lobe expansion over the course of human evolution (e.g., Deacon, 1997) are
not supported by this finding, since the selection for individuals with relatively
larger frontal lobes would also select for people with relatively smaller parietal
lobes. There is no evidence for parietal lobe contraction during human evolu-
tion, nor would that be expected given that tool use, a hallmark of hominid
cognitive evolution, depends on parietofrontal circuits (Johnson-Frey, 2003).
Further evidence against frontal lobe expansion in hominid evolution comes
from comparative anatomy. Volumetric MRI studies suggest that the proportional
volumetric representation of the principal lobes of the human brain follows the
pattern seen in our closest relatives, the great apes (see Fig. 1-3) (Semendeferi and
Damasio, 2000; Semendeferi et al., 2002). In general, across mammalian species,
the proportional representation of different parts of the brain tends to remain fairly
constant, even while volume varies tremendously (Jerison, 1973; Finlay and Dar-
lington, 1995; Finlay et al., 2001). It comes as no surprise then that within our own
species volumetric variability exceeds proportional variability.
Brain Volume and Tissue Composition: Sex Differences
Both postmortem and MRI volumetric studies of the brain have found that on
average, men's brains are larger than women's brains (e.g., Pakkenberg and
Voight, 1964; Ho et al., 1980; Skullerud, 1985; Peters et al., 1998; Courchesne
Differences
[81]
et al., 2000) but that women have a higher proportion of gray matter (versus
white matter) than men do (Filipek et al, 1994; Passe et al., 1997; Peters et al.,
1998; Gur et al., 1999). The overall size difference remains even after control-
ling for body size (Holloway, 1980; Peters et al., 1998; Nopoulos et al., 2000).
Similarly, the gray-white difference is also present after controlling for brain
size: if one compares a male and a female brain with the same overall volume,
the woman's brain will contain, on average, more gray matter and less white
matter than the man's brain (Peters et al., 1998; Gur et al., 1999).
We have studied volumetric sex differences in brain structures (Allen et al.,
2002, 2003b) in the group of 46 healthy, right-handed individuals between the
ages of 22 and 49 years mentioned above. There were 23 men (mean age 32.1
years) and 23 women (mean age 32.6 years) in the group. On average, male brains
are about 12% larger than female brains (left hemisphere and left hemicerebel-
lum: mean 618.9 cc, standard deviation [s.d.] 54.7 for men and 547.0 cc, s.d. 47.8
for women; right hemisphere and right hemicerebellum: mean 621.7 cc, s.d. 58.3
for men and 552.7 cc, s.d. 49.5 for women). Male brains were no more variable
than female brains. Although there was not a significant difference in size be-
tween the two hemispheres, both men and women were more likely to have larger
right rather than left hemispheres. In general, the hemisphere patterns hold for
each of the major lobes and substructures of the brain. However, the occipital
lobe appears to be somewhat less sexually dimorphic than other regions.
Segmentation of the brain into gray and white matter provides additional in-
sights into the volumetric differences between the sexes. As discussed above,
women (left hemisphere gray/white ratio: mean 1.35, s.d. 0.09; right hemisphere:
mean 1.35, s.d. 0.09) tend to have relatively more gray matter than men (left
hemisphere gray/white ratio: mean 1.27, s.d. 0.10; right hemisphere: mean 1.25,
s.d. 0.10). Statistical analyses indicate that the higher gray/white ratio in women
is not simply a matter of brain size (if the brain were simply a sphere, the sur-
face/volume ratio would increase as the volume decreased). For example, the
cerebra of 7F (Fig. 6-5) and of 16M (Fig. 6-8) are about the same size, 1,020 cc;
but the gray/white ratio of the woman is 1.37, while that of the man is 1.10.
The sex difference in cerebral volume is due more to differences in white
matter volume than gray matter: analyses of covariance indicate that the
gray/white ratio difference between the sexes disappears when controlling for
WM volume, indicating that it is variability in white matter volume that most
profoundly influences this relationship (Allen et al., 2003b). There is substantial
variation among regions in the relative composition of gray and white matter.
This is not surprising given the relative amounts of cortex and white matter core
that are found in different parcellated regions (e.g., occipital versus frontal lobes).
On the other hand, some gray-white composition patterns may reflect other fac-
tors. For example, effect size statistics indicate that the right parietal lobe is the
most sexually dimorphic for the gray/white ratio compared to all other regions
of the brain (female gray/white ratio: mean 0.95, s.d. 0.08; male: mean 0.85, s.d.
0.08). This is due to the fact that women have nearly as much gray matter in
this region as men (59.8 cc versus 62.5 cc), while at the same time they have sub-
stantially less white matter (63.0 cc versus 74.2 cc). While the functional signif-
icance of this pattern is not yet clear, it could be a result of the distribution of
sex steroid receptors in the brain (Reiss et al., 1995; Goldstein et al., 2001).
Human Brain Anatomy
IN COMPUTERIZED IMAGES
[82]
In terms of sex differences, a region of the brain that has drawn much at-
tention over the years is the corpus callosum (CC) (de Lacoste-Utamsing and
Holloway, 1982; Bishop and Wahlsten, 1997). Although some studies have shown
that it might be larger in women than men, most in vivo neuroimaging studies
have shown that it is larger in men, reflecting the overall volume difference (e.g.,
Sullivan et al., 2001). Earlier studies based on postmortem material may not have
controlled for the effects of age when comparing men and women, which could
certainly have led to spurious results concerning sexual dimorphism in the CC.
The CC is a highly variable structure, as can be noted by inspection of the mesial
surfaces of the 46 brains presented in Figures 6-3 to 6-10. Subjects 6F and 7M
(both Fig. 6-5) have proportionally small CCs, while 14F (Fig. 6-7) and 19M
(Fig. 6-9) have relatively large CCs for their respective total brain volumes.
Although quantitative studies of the corpus callosum have traditionally con-
sidered the surface area of the structure (as it appears on the mesial surface
when the brain is bisected through the hemispheric fissure), in our studies we
have defined it as a three-dimensional structure encompassing the band of mid-
line white matter inferior to the cingulate gyrus and superior to the lateral ven-
tricles (see Allen et al., 2002, for a detailed description). Our results confirm
that it is significantly larger in men than women (10.6 cc, s.d. 1.62 vs. 9.68 cc,
s.d. 1.26). However, the corpus callosum makes up a significantly larger pro-
portion of the total white matter in women (mean 0.0236, s.d. 0.0026) than in
men (mean 0.0219, s.d. 0.0026). Since the corpus callosum is composed pri-
marily of myelinated axonal fiber tracts, this result may indicate that a sub-
stantial portion of the overall white matter volume sex difference may be due
to glial cells or blood vessels that contribute to the "white matter" seen in MR
images. In other words, the "excess" of white matter relative to gray matter in
men may not be due to a net increase in neuronal connectivity.
Interhemispheric Asymmetry
In terms of overall volume, the brain is a fairly symmetric structure. However,
there is a tendency for right-handed individuals to have a slightly larger right
hemisphere (Allen et al., 2002). Some regions of the brain show pronounced
asymmetries, while others are fairly symmetric (Toga and Thompson, 2003).
For example, the left cingulate gyrus is substantially larger in volume than the
right, especially in males: we found that the mean left cingulate gyrus volume
was 23.4 cc compared to 19.7 cc in the right (Allen et al., 2002). This may be
due to the fact that the left cingulate sulcus is more likely to be "doubled" on
the left side rather than the right (Paus et al., 1996), although the opposite pat-
tern also occurs with some frequency. See brains C, E, and G, in Chapter 5 sub-
jects and 8M (Fig. 6-5), 3M (Fig. 6-4), and IF (Fig. 6-3) in this chapter for ex-
amples of cingulate sulcus asymmetry. Brain A shows a symmetric pattern in
the cingulate sulcus, while brains O and K show reversed asymmetry.
The frontal and occipital petalias comprise another well-known cerebral
asymmetry (Chui and Damasio, 1980; Zilles et al., 1996). Seen from above or
in some axial sections, the human brain often protrudes slightly anteriorly in
the right hemisphere (frontal petalia) and posteriorly in the left hemisphere
[83]
Differences
(occipital petalia). In practical terms, the right frontal lobe goes farther into the
frontal bone than the left, and the left occipital lobe goes farther back into the
occipital bone than the right, creating bone depressions, the petalias. We have
found that the petalia may correspond to asymmetries in the distribution of
white matter (but not gray matter), since white matter volumes tend to be larger
in the right frontal and left occipital lobes compared to their opposites (Allen
et al., 2003b). Brains B and C in Chapter 5 provide particularly good examples
of the "typical" petalia pattern.
One of the best-known asymmetries in the brain involves the planum tem-
porale (PT) (Geschwind and Levitsky, 1968; Westbury et al., 1999). The PT in-
cludes all of the cortex on the superior surface of the temporal lobe that is pos-
terior to Heschl's gyrus, terminating at the posterior end of the sylvian fissure
(see Figs. 2-10, 3-7, and 4-7). In right-handers, this structure is clearly larger
in the left hemisphere. In a recent study of 25 right-handed adults (14 women
and 11 men), we found that the left PT is 23.6% larger in the left hemisphere
compared to the right (Emmorey et al., 2003). Although it was once thought
that this asymmetry might be an anatomic pattern that corresponded to the
evolution of spoken language (Galaburda et al., 1978), recent research on great
apes has shown that a larger left PT is something that we share with our clos-
est relatives, who do not possess language (Gannon et al., 1998; Hopkins et al.,
1998). The relationship among PT asymmetry, language dominance, and hand-
edness is not clear (Moffat et al., 1998). Congenitally deaf individuals also have
a larger left PT, indicating that the pattern does not arise as a result of the pro-
cessing of spoken language (Emmorey et al., 2003; see discussion below).
The course of the sylvian fissure provides another common instance of brain
asymmetry, one that in all likelihood is related to PT asymmetry. In right-
handers, the sylvian fissure tends to turn upward in the right hemisphere and
to extend more posteriorly in the left (Rubens et al., 1976). Since the PT is de-
fined by the extension of the sylvian fissure, this asymmetry is usually consis-
tent with the left PT being larger than the right.
The angle of the posterior sylvian fissure can be quantified in MR images
by taking the ac-pc plane as a reference. In groups of left- and right-handers
studied in our laboratory, we have found that, while right-handers have the ex-
pected pronounced upward course in the right rather than left posterior syl-
vian fissure (average of 36.7 upward angle in the right hemisphere compared
to 22.6 in the left), there is no such asymmetry in left-handers (31.5 angle in
the left hemisphere and 33.4 in the right). These patterns can be visually con-
firmed by examining the sylvian fissures of the brains of the 46 right-handed
individuals illustrated in this chapter and comparing them to the left-handed
individuals represented in brains I-O in Chapter 5.
Brain Volume and Tissue Composition: Aging Effects
Age is a fundamental component of human variation, and changes in brain size
and composition over the life span reflect this (Raz, 1999). Over the past decade,
volumetric MRI analyses have added much to our understanding of many as-
[84]
Human Brain Anatomy
pects of brain development and aging (e.g., Cowell et al., 1994; Coffey et al.,
1998; Tang et al., 2001; Sowell et al., 2003). The human newborn has a brain
that is about 30% of adult size, and brain growth is rapid until the age of 2
years. By the age of 2, the average child's brain volume is over 1,000 cc, and
growth contipues, at a much slower rate, between the ages of 2 and 14 years
(Courchesne et al., 2000). It has long been known that the aging brain is char-
acterized by volume reduction, with an increase in the size and depth of sulci.
Recent MRI research indicates, however, that age-associated brain atrophy does
not occur in a uniform manner, although different studies have produced con-
flicting results. For example, age-associated volume reductions have been more
pronounced in the frontal lobe compared to other brain regions (Cowell et al.,
1994; Jernigan et al., 2001), although others have found that the frontal and
temporal lobes age at similar rates (Bartzokis et al., 2001). Within the tempo-
ral lobe, the hippocampus may be more sensitive to age than the amygdala
(Pruessner et al. 2001).
Several aging studies have found that gray and white matter volumes do not
change over the life span at the same rate (Jernigan et al., 2001; Bartzokis et al.,
2001; Guttman et al., 1998). Gray matter volume declines throughout adult-
hood and into old age at a more or less linear rate. By contrast, some studies
have shown that white matter volumes may actually increase slowly through
adulthood, peaking in the 40-50 year range (Bartzokis et al., 2001). After 60
years of age, there is a precipitous decline in white matter volume (Guttman et
al., 1998). Brain aging patterns in men and women are largely similar, although
some studies show that the gray matter decline may be slower in women than
in men (Cowell et al., 1994).
We have analyzed aging patterns in the gray and white matter in a group of
87 healthy, right-handed adults (44 women and 43 men), ranging in age from
22 to 88 years (Allen et al., 2003a). Regression analyses indicate that for both
men and women gray matter declines linearly across the life span, while white
matter volumes remain steady or actually increase slightly throughout much of
adulthood, peaking around age 45-55 years. This is followed by the precipitous
decline of white matter volume noted above. These patterns hold approximately
for the hemispheres and for the principal lobes. Considering the cerebrum as
a whole, we can predict, based on the regression equations, that from age 30 to
70 years white matter volume will decrease 6.4% in women and 5.5% in men
and that gray matter will decline 9.9% in women and 9.1% in men. Over this
period, the gray/white ratio will drop from 1.29 to 1.26 in women and from
1.22 to 1.20 in men. When we compare volume predictions at 30 and 80 years
of age, the gray matter volume shows a decrease of 12.4% in women and 11.4%
in men and the white matter decrease is 25.0% in women and 21.6% in men.
The predicted gray/white ratio increases to 1.50 in women and 1.43 in men, re-
flecting the fact that white matter loss occurs much more quickly past the age
of 70 than gray matter loss.
Brains U through Z in Chapter 5 are from healthy subjects older than 60
years. They demonstrate that normal aging patterns and individual variations
in aging become important components of variability in the brain morphology
of normal, healthy subjects greater than 60 years of age. Brain U, which comes
[85]
Differences
Human Brain Anatomy
IN COMPUTERI ZED IMAGES
from a 73-year-old male, exhibits the pronounced sulcal widening that results
from loss of gray and, especially, white matter. By contrast, brain Y, which be-
longs to a somewhat younger individual, shows only a mild degree of sulcal
widening. The visual difference between brains U and Y is confirmed by quan-
titative measures: the gray/white ratio for brain U is 1.38, while that for Y is
1.19. This indicates that white matter loss in brain U is markedly more advanced
than it is in brain Y, thus the higher gray/white ratio. Individual variation in
aging can be seen by comparing brain X, which belongs to a 79-year-old fe-
male, to brain U. Despite the fact that the brain X subject is 6 years older than
the brain U subject, her brain is markedly less atrophied.
Heschl's Gyrus in Congenitally Deaf Individuals
Earlier, we mentioned that congenitally deaf subjects appear to have the same
asymmetry in the planum temporale as hearing subjects (Emmorey et al., 2003).
Heschl's gyrus (HG), the small transverse gyrus located on the surface of the
temporal lobe, is the approximate site of the primary auditory cortex
(Rademacher et al., 1993, 2001). By comparing HG in congenitally deaf and
hearing individuals, it may be possible to gain insight into the impact the sen-
sory environment has on the development of a specific neural structure.
We investigated a group of 25 congenitally deaf individuals and 25 age- and
sex-matched hearing comparison subjects with respect to gray and white mat-
ter composition of several structures in the temporal lobe, including the supe-
rior temporal gyrus as a whole (its posterior sector, the planum temporale), and
Heschl's gyrus (Emmorey et al., 2003). Heschl's gyrus is defined anteriorly by
the first transverse temporal sulcus and posteriorly by Heschl's sulcus (the sec-
ond transverse sulcus). When more than one sulcus is present, the most ante-
rior one is used. Heschl's gyrus is visible on the superior surface of the tempo-
ral lobe in a "planum cut", a preparation in which the frontoparietal operculum
has been removed. Accurate parcellation of HG is greatly facilitated using sur-
face landmarks and multiple 2-D slice orientations.
Although the planum temporale is similar in deaf and hearing subjects, we
found that these two groups had differences in other structures. Most impor-
tantly, the gray/white ratio in Heschl's gyrus was significantly higher in deaf
compared to hearing subjects. This appears to be due primarily to a reduction
in white matter volume: after correcting for overall temporal lobe volume, there
was no difference in Heschl's gyrus gray volume between deaf and hearing sub-
jects but HG white matter volume was significantly smaller in deaf subjects.
This result suggests that auditory deprivation from birth results in less myeli-
nation, fewer fibers projecting to and from the auditory cortex, and/or greater
axonal pruning in the congenitally deaf.
[86]
Figure 6-1
Automated segmentation of a magnetic resonance brain volume.
Gray matter is represented in dark blue, white matter in cyan,
and cerebral spinal fluid in red. Each figure is composed of
10 contiguous coronal slices, offset slightly to convey the three-
dimensional structure.
Figure 6-2
Parcellation of the major sectors of the cerebrum. Landmarks on
the surface of the brain, such as sulci, are used to guide tracing
of regions of interest (ROI) on contiguous coronal slices. In the
top row, lateral and mesial views of the left hemisphere are pre-
sented; cyan line corresponds to the position of the coronal slice
shown between the two hemisphere views. Parcellation of ROIs
on this coronal slice is indicated by green lines. In the middle
row, lateral and mesial views of the hemisphere are presented
with the major sectors colored in following parcellation on con-
tiguous coronal slices (1.5 mm thick) through the whole brain.
Four representative coronal slices demonstrating parcellation of
the brain are presented in the bottom row; their positions are
indicated by lines 1-4 on the lobes in the middle row. FL, red;
PL, green; TL, blue; OL, yellow; Cing, magenta; Ins, teal. Itc, lat-
eral temporal cut; oc, occipital cut; mpc, mesial parietal cut; mtc,
mesial temporal cut.
[87]
Figures 6-3 to 6-10
Lateral and mesial views of 46 parcellated brains with gray and white matter volumes (in cubic millimeters) for left and right hemi-
spheres: 23 male (M) and 23 female (F) (females on the left and males on the right; brains are presented in descending order of to-
tal volume for each sex. All brain images are rendered to the same scale. The number following the subject number is the subject's
age in years at the time of the magnetic resonance scan. Frontal lobe, red; parietal lobe, green; temporal lobe, blue; occipital lobe,
yellow; cingulate gyrus, magenta.
Figure 6-3
Parcellation of the cerebrum and volume data for male and fe-
male subjects 1 and 2.
[88]
IF 23
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
114884
83411
78785
37271
16523
White
110782
47503
79199
18675
4675
Left
Gray
113646
81995
70367
37688
17825
White
111924
43274
71845
24531
7421
1M 23
Frontal
Temporal
Parietal
Occipital
Cingulate
^^m^. _
Right
Gray
122890
99398
65851
29709
20852
White
122750
55995
79088
16714
6738
^p-
Left
Gray
118233
96384
71494
26771
19993
White
116544
54785
81579
18148
8393
2F 26
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
107633 105901
66115
67862
34936
21773
33063
66734
19498
6519
Left
Gray
107064
76400
62501
29837
19682
White
95771
39474
65148
18854
7923
2M 46
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
111558 134797
88266
63373
32490
15826
62083
81923
19343
5523
Left
Gray
112701
86714
63269
32941
15982
White
126925
58483
83031
23221
8931
Figure 6-4
male subjects 3-5.
[89]
3F 41
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
92576
71077
59917
36471
15909
White
98659
42683
74287
21839
5143
Left
Gray
92584
66126
63812
34294
15878
White
99756
40536
75528
21412
7278
3M 25
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
115500
91433
69649
36805
17836
White
111458
52965
82503
23223
5682
Left
Gray
109075
86375
72932
36197
21882
White
104765
48757
84460
24668
9390
4F 24
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
105623
75370
65532
31135
14671
White
94858
36539
75778
16326
4570
Left
Gray
105188
74198
62568
30098
14630
White
96042
36297
67674
18544
6395
4M 32
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
115368
80601
76742
33961
18316
White
116119
45330
86637
20707
5389
Left
Gray
111965
82249
71548
30715
24175
White
106326
46698
86126
21749
10415
5F 24
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
96119 100259
63599
63877
29970
14703
36764
84901
20185
5150
Left
Gray White
98084 99843
62479
63473
32286
11937
35724
79679
24749
6111
5M 23
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
113992 114729
89685
68391
30887
14012
51446
88064
19924
5006
Left
Gray White
111395 113429
77652
75041
35416
17798
42071
87945
25602
7590
Figure 6-5
Parcellation of the cerebrum and volume data for male and
female subjects 6-8.
[90]
6F 26
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
103524 98169
72454
58773
36907
13464
42692
61790
21719
3567
Left
Gray White
102647 91948
71393
54473
38339
12115
40424
63247
24538
5251
6M 22
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
118797 128975
88306
63261
26963
13485
54770
77117
14539
4783
Left
Gray White
127333 139931
82070
50432
30538
14989
46342
57626
21747
7392
-^^
7F 23
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
110389
66418
69192
27048
15019
White
96916
31062
70197
13355
4380
Left
Gray
111351
75639
58868
26864
14501
White
93280
37665
61730
15889
5981
7M 24
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
109792
83882
69220
32597
16269
White
106403
42326
79588
21333
5559
Left
Gray
116020
80514
69138
34798
13267
White
108541
39024
79336
24448
5533
8F 41
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
96793 97578
72653 40173
57404 61460
27864 16251
14362 4091
Left
Gray White
99521 95006
68233 36478
57733 63477
29577 20614
11451 5128
8M 32
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
101196 111313
64265 39680
72996 99407
34262 24153
15796 5843
Left
Gray White
102990 110455
76279 49820
66577 88844
28449 21911
17586 8638
Figure 6-6
[91]
9F 39
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
99585
74735
59061
29866
13481
White
87623
36586
60107
16289
3408
Left
Gray
94113
78152
57595
23762
16692
White
84643
39263
59780
15261
5706
9M 45
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
107026
73948
62025
30849
13410
White
120092
44668
82277
20784
5529
Left
Gray
103570
79512
62418
24370
14398
White
117836
50970
78276
19370
7959
10F 28
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
96613 93635
68715 34701
62196 62305
24149 12047
13174 4490
Left
Gray White
96648 89800
71443 36953
54839 55328
24410 14132
15957 7128
10M 28
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
106103 103380
83187 45688
64390 75212
28988 16304
15617 4945
Left
Gray White
105243 96829
82616 46027
67967 78625
26652 18078
14386 6333
11F 42
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
94961 102228
66620
48422
23995
12881
42973
55421
14665
4452
Left
Gray
91653
62781
50920
23878
14039
White
96670
40809
58919
16133
6546
11M 43
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
94407 113973
69432
58752
31699
13356
43975
82730
27674
5422
Left
Gray
96405
74810
57420
25892
15432
White
112035
46981
80297
23341
8759
Figure 6-7
male subjects 12-14.
[92]
12F 36
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
94687
60292
65320
25774
16517
White
84335
30507
68174
14731
4253
Left
Gray
95024
69904
58344
24570
11142
White
81583
37697
62278
15417
4632
12M 38
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray
106107
79990
54183
26612
16471
White
117441
47058
71690
19841
6710
Left
Gray
102185
73779
62092
25824
17111
White
109943
40860
73247
22020
9019
13F 47
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
90756 91944
67537 36901
55294 63071
25260 15048
12273 4348
Left
Gray White
89656 83409
59687 31595
59520 69549
29165 20514
12402 5427
13M 42
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
103786 104071
82889 48384
62273 76965
24181 13864
12352 4393
Left
Gray White
101134 104282
76424 43264
58997 73407
31582 21592
15402 7206
14F 43
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
89799 89803
64719 35843
55167 67358
28855 16040
11756 3322
Left
Gray White
86249 87027
66030 36473
52794 62672
27082 17973
15004 6389
14M 24
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
104907 99505
79372 42289
60361 67768
28287 16063
12353 4179
Left
Gray White
106041 96821
81008 43296
56478 65929
24600 16452
16099 6531
Figure 6-8
[93]
15F 26
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
85513 93660
60158 32548
48870 60441
31416 21025
16979 5628
Left
Gray
85618
65374
49975
24266
12142
White
88750
36805
62953
19051
6044
15M 24
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
104639 92950
79177 39211
66755 73786
26768 14362
12981 3967
Left
Gray White
98837 89565
79913 40138
66823 70555
24180 14526
15055 6821
16F 32
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
93574 84679
60731 30269
63480 61391
24506 14049
13089 4232
Left
Gray
90742
63941
60143
21932
17326
White
79975
33549
61285
15012
7252
16M 38
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
92788 102536
73354
54402
28425
10738
44630
73196
20078
3774
Left
Gray
94837
66472
59243
30884
11970
White
100663
35513
77122
23549
6253
17F 28
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
92418 80806
66764 32334
57575 61125
22372 11862
16356 4445
Left
Gray White !
93907 79977
71282 36154
53923 55465
20984 11857
12168 5064
17M 42
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
100599 94659
69858 36351
52786 60893
34741 24089
12104 4462
Left
Gray White
96896 93662
77868 42257
58216 64832
23177 17330
16322 6980
w hite White
White
Figure 6-9
[94]
18F 32
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
85183 78352
64148 34854
58768 64573
22906 10777
14313 4015
Left
Gray White
85737 76785
65656 35529
51573 61241
24128 14310
11651 5409
18M 26
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
99255 88063
71693 36205
61648 66470
28017 14517
15807 4808
Left
Gray White
100376 86538
69159 32038
64925 64385
28766 16367
15559 6001
19F 27
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
84695 80640
62535 33812
53137 57759
20876 9528
14066 4068
Left
Gray White
86749 80855
59766 31558
49834 53772
24332 14685
11173 6597
19M 28
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
94495 90140
71856 39676
54354 68812
23631 14106
13872 4568
Left
Gray White
90793 89489
74006 42616
60145 67255
20980 14344
13097 5048
20F 35
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
88651 78612
65132 33563
52041 53863
24379 12226
12082 3421
Left
Gray White
89964 79984
62840 33507
53050 498%
20882 11290
9375 4013
20M 29
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
91676 88583
64712 36208
56012 74212
25761 16615
13899 5174
Left
Gray White
93344 90461
63665 35250
5291 1 65903
31263 20454
13372 5529
21F 28 Right Left
Frontal
Temporal
Parietal
Occipital
Cingulate
Gray
82421
61638
43663
26564
13229
White
86260
36797
46564
14223
4162
Gray
83263
59141
41526
28515
13334
White
80744
35156
47957
19237
5914
22F 41 Right Left
Frontal
Temporal
Parietal
Occipital
Cingulate
Gray
84105
59818
50628
25030
10783
White
79429
32721
54151
13133
3043
Gray
81525
60058
52812
21164
10929
White
73833
33078
57115
13162
4412
Figure 6-10
male subjects 21-23.
21M 49
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
83259 89690
58720 34309
50715 65215
34112 20140
12737 4472
Left
Gray White
86368 87486
61148 34625
52504 67738
30637 19013
12520 5423
22M 33
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
84087 84820
77437 41193
47486 57257
22365 14798
13345 4734
Left
Gray White
86240 80119
66731 33771
61096 66723
21058 14912
12915 6462
23M 23
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
85490 83443
65606 37808
49722 62786
25806 16438
9338 2606
Left
Gray White
81152 81195
62561 33449
52171 62562
24843 18423
14586 6805
[95]
23F 37
Frontal
Temporal
Parietal
Occipital
Cingulate
Right
Gray White
82133 78424
61405 33222
46446 53627
23503 12154
11124 3614
Left
Gray White
81006 78859
56847 30732
47395 49817
23781 15134
13717 6003
D
olicho, the brain described in Chapter 2, is here sliced into several axial, coro-
nal, and parasagittal sequences. The goal is to demonstrate the role played
by different incidences of sections in the apparent relations of anatomic structures.
To obtain these sections at the desired angle to a set external landmark and at
levels that include most structures in posterior and anterior fossae, I relied on the
capabilities of Brainvox (Damasio and Frank, 1992; Frank et al., 1997). First, the
canto-meatal line, the reference most often used in the alignment of routine mag-
netic resonance (MR) and computed tomographic (CT) scans, was drawn on both
the left and the right; second, a plane that included these two lines was constructed;
then, using another Brainvox program, MP-FIT, parallel planes at a distance of 5
mm were constructed so as to encompass the whole brain and posterior fossa con-
tents. The result is the 24 axial sections seen in Figures 7-1 through 7-25.
There is a reason to choose a set of cuts parallel to the canto-meatal line for
the first sequence, usually designated as "axial" slices or cuts; this is the orien-
tation seen in most axial sequences obtained in standard MR studies, even if it
is not exactly the incidence most often used in brain cutting sessions and in
most anatomy atlases. The incidence most commonly used for the latter is closer
to the inferior orbito-meatal line, which has a rostral angulation of approxi-
mately 10 in relation to the canto-meatal line. Of note, that was the incidence
chosen in the first edition of this atlas.
The next section depicts a series of coronal slices obtained at 90 to the canto-
meatal axial slices (Figs. 7-26 to 7-58). Again, this is the incidence most often
seen in coronal slices of standard MR sequences, given that the coronal sequence
is taken in a plane perpendicular to that of the axial sequence. The individual
brain slices are obtained at 5 mm intervals, as everywhere else in this atlas.
Often, MR studies are conducted with a rostral angulation, a so-called neg-
ative tilt. This is often necessary in older subjects, who may have changes in the
cervical spine that do not permit head flexion, and in intubated patients, whose
heads must be hyperextended. In those same circumstances, CTs also are ob-
tained with a negative tilt. The incidence of 30 chosen for Figures 7-59 to
7-85 is meant to give an idea of the configuration of the brain sections in such
Chapter 7
Sections through
Dolicho
[97]
Human Brain Anatomy
IN COMPUTERI ZED IMAGES
a situation. (This sequence corresponds roughly to the 2 0 sequence seen in
the first edition of this atlas.) Obviously, there is no "standard" negative tilt.
The reader should take note of the marked changes in appearance of certain struc-
tures when moving from an axial sequence parallel to the canto-meatal line to a se-
quence obtained with a rostral negative tilt. For instance, observe what happens to
the calcarine fissure and to the two main components of the mesial occipital lobe: the
infracalcarine region, with the lingual and fusiform gyri, and the supracalcarine sec-
tor, with the cuneus. Compare Figures 7-66 through 7-71 with Figures 7-12 through
7-15. The cuts in Figures 7-66 and 7-67 barely slice through the temporal lobe, but
they show the most posterior sector of the calcarine fissure in the polar region of the
occipital lobe. This same sector of the visual cortex is seen in Figures 7-12 and 7-13.
However, in the latter images from the canto-meatal axial sequence, the visual cor-
tex appears in the same cut in which most of the temporal lobes and a middle sector
of the frontal lobe can be seen. If we look for the corresponding frontal sector in the
hyperextension axial sequence, we find it in Figures 7-76 through 7-79, in which no
occipital structures are seen and only frontal and parietal structures are visible.
As happened with the canto-meatal axial sequence, the negative tilt series is
followed by the corresponding coronal series. This sequence was obtained at 90
from the corresponding axial sequence, which translates into 120 to the canto-
meatal line (see Figs. 7-86 to 7-118). Note that here, at the point at which the cuts
slice through the beak of the corpus callosum, they also slice through the poste-
rior sector of the temporal pole. In the coronal sequence (corresponding to the
axial sequence parallel to the canto-meatal line), this sector of the temporal pole
appears only when the slices pass through the body of the callosum, well behind
the beak (Figs. 7-37, 7-38). In relation to the frontal operculum, slices 11 and 12
(Figs. 7-37,7-38) cut through its posterior regions, the pars opercularis, and even
the lower sector of the precentral gyrus, while in Figures 7-95 and 7-96 cuts 9
and 10 at the level of the beak of the callosum still slice through the pars trian-
gularis, barely touching the anterior sector of the pars opercularis. With respect
to the characteristic juncture of the calcarine fissure and the occipitoparietal sul-
cus (in a ><-like shape) seen in the right hemisphere of cut 26 (Fig. 7-111),
note that it coincides with a cut through the posterior region of the paracentral
gyrus in the coronal hyperextension sequence. However, in the coronal sections
perpendicular to the canto-meatal line (slice 27 in Fig. 7-53), the superior sec-
tor of the cut shows the most posterior regions of the cuneus. To see the para-
central gyrus, we need to move back to Figures 7-46 and 7^7 (cuts 20 and 21).
The last axial sequence (Figs. 7-119 to 7-144) is a posterior fossa incidence,
often used when the posterior fossa is the target region. This sequence was ob-
tained using a caudal rotation of 10 to the canto-meatal plane. Once again, the
relationships vary. For example, while the left pars triangularis first appears in Fig-
ure 7-128 (when the slice reveals anterior temporal lobe, no occipital lobe or cor-
pus callosum, and mid-sectors of the cerebellum), in the hyperextended axial se-
quence this same frontal region is visible together with the beak and body of the
callosum and the parietal lobe (Fig. 7-77). On the other hand, in the canto-meatal
axial sequence, this frontal sector is seen in slices 9 and 10 (Fig. 7-11) along with
a large sector of the temporal lobe and the lower sector of the occipital lobe.
The parasagittal sequences were also obtained at 90 to the axial sequence.
They usually proceed from the left to the right hemisphere, parallel to the in-
[98]
terhemispheric fissure. The incidence of the axial sections exerts its influence
in the position the section has on the film (more or less tilted posteriorly) but
does not interfere with the relative position of the different structures, given
that the sequence is always meant to be parallel to the interhemispheric fissure.
One parasagittal sequence (Figs. 7-145 to 7-183) is offered in this chapter, start-
ing in the left temporal lobe and proceeding to the right. The sections are also
set at 5 mm intervals with the exception of slices 18 and 19, which are sepa-
rated by 10 mm, 5 mm on each side of the interhemispheric plane.
The last sequence in the chapter is again a canto-meatal axial sequence (the
same as in Figs. 7-2 through 7-25). Here, the images show the approximate
limits of the cytoarchitectonic fields of Brodmann. For the same reasons to
which I alluded earlier, MR images, even those with the best resolution, do not
allow identification of cytoarchitectonic fields. All we can do is approximate us-
ing gyral and sulcal information related to the available data from microscopic
studies. As noted, it is reasonable to refer to the pars opercularis of the inferior
frontal gyrus as Brodmann's field 44 or to the supramarginal gyrus as area 40;
but we must keep in mind that we are simply changing the name we attribute
to a macroscopically identifiable sector of the brain, the pars opercularis or the
supramarginal gyrus, because it has recognizable macroscopic limits and not
commenting on the microscopic characteristics of the region. Brodmann's ter-
minology is used when we designate regions that occupy more than one macro-
scopically identifiable sector (e.g., area 37 in the temporal lobe). For these rea-
sons and because several readers commented on the lack of cytoarchitectonic
field identification in the previous edition of this atlas, I decided to include it
here. It will be important, however, to examine this last sequence together with
the images that define the gyri because only correct identification of the gyri
allows for translation into Brodmann's terminology.
In each of the sequences, the image on the left page has the sulci identified
and the image on the right page has the gyri and subcortical structures identi-
fied. The reader should note that even in images carefully placed at the same
level in the right and left hemispheres, the structures appearing on either side
are not mirror images of each other.
Only the clearly recognizable subcortical structures were labeled. No attempt
was made to label likely sites of structures not seen, nor were the several seg-
ments of the ventricular system labeled, the main purpose of the atlas being the
recognition of cortical structures.
The actual MR image of the cut used for anatomic description is shown on
the upper and lower outside corners of each page. Note that the MR images and
the stripped brain slices in the center are precisely the same. Next to the cen-
terfold, the four views of the brain, right and left, lateral and mesial views, show
the setting of the level of the cuts used on that page. All slices are shown in the
radiologic convention: right hemisphere on the left, left hemisphere on the right.
The first image of each sequence shows the right and left hemispheres, in dor-
solateral and mesial views, as seen in Chapter 2, with the gyri colored (Figs. 2-2,
2 4, and 2-6) with a superimposed grid corresponding to the slices to follow.
Above is the image of the transparent head with the left hemisphere visible. The
canto-meatal line is drawn in red, and a sample of slices is marked to show the
angulation of the slices of the section in relation to the canto-meatal line.
Sections through Dolicho
[99]
Canto-meatal
Incidence:
Axial Slices
[100]
Figure 7-1
Left hemisphere of Dolicho seen through the transparent head
on top. The canto-meatal line is marked in red. Black lines rep-
resent a sample of the slices obtained parallel to the canto-meatal
line. Underneath are the two hemispheres in lateral and mesial
views, as seen in Figures 2-2 and 2-4, with gyri individually col-
ored. The lines correspond to the slices parallel to the canto-
meatal line depicted in this section.
[101]
Figure 7-2
Slices 1 and 2 parallel to the canto-meatal line in Dolicho.
Identification of sulci.
[102]
Figure 7-3
Identification of gyri.
[103]
Figure 7-4
[104]
Figure 7-5
Slices 3 and 4 parallel to the canto-meatal line in Dolicho. Iden-
tification of gyri.
[105]
Figure 7-6
[106]
Figure 7-7
[107]
Figure 7-8
[108]
Figure 7-9
[109]
Figure 7-10
[110]
Figure 7-11
[111]
Figure 7-12
[112]
Figure 7-13
[113]
Figure 7-14
[114]
Figure 7-15
[115]
Figure 7-16
[116]
Figure 7-17
[117]
Figure 7-18
[118]
Figure 7-19
[119]
Figure 7-20
[120]
Figure 7-21
tification of gyri.
[121]
Figure 7-22
[122]
Figure 7-23
[123]
Figure 7-24
[124]
Figure 7-25
tification of gyri.
[125]
Canto-meatal
Incidence:
Coronal Slices
[126]
Figure 7-26
Left hemisphere of Dolicho seen through the transparent head parallel to the canto-meatal line. Underneath are the two hemi-
on top. The canto-meatal line is marked in red. Black lines rep- spheres in lateral and mesial views, as seen in Figures 2-2 and
resent a sample of slices obtained at 90 to the canto-meatal line, 2-4, with gyri individually colored. Lines correspond to slices ob-
which is the coronal incidence when the axial slices are obtained tained at 90 to the canto-meatal line as depicted in this section.
[127]
Figure 7-27
Slices 1 and 2 at 90 to the canto-meatal line in Dolicho. Identi-
fication of sulci.
[128]
Figure 7-28
fication of gyri.
[129]
Figure 7-29
fication of sulci.
[130]
Figure 7-30
fication of gyri.
[131]
Figure 7-31
fication of sulci.
[132]
Figure 7-32
fication of gyri.
[133]
Figure 7-33
fication of sulci.
[134]
Figure 7-34
fication of gyri.
[135]
Figure 7-35
Slices 9 and 10 at 90 to the canto-meatal line in Dolicho. Iden-
tification of sulci.
[136]
Figure 7-36
tification of gyri.
[137]
Figure 7-37
[138]
Figure 7-38
tification of gyri.
[139]
Figure 7-39
[140]
Figure 7-40
tification of gyri.
[141]
Figure 7-41
[142]
Figure 7-42
tification of gyri.
[143]
Figure 7-43
[144]
Figure 7-44
tification of gyri.
[145]
Figure 7-45
[146]
Figure 7-46
tification of gyri.
[147]
Figure 7-47
[148]
Figure 7-48
tification of gyri.
[149]
Figure 7-49
[150]
Figure 7-50
tification of gyri.
[151]
Figure 7-51
[152]
Figure 7-52
tification of gyri.
[153]
Figure 7-53
[154]
Figure 7-54
tification of gyri.
[155]
Figure 7-55
[156]
Figure 7-56
tification of gyri.
[157]
Figure 7-57
Slices 31, 32, and 33 at 90 to the canto-meatal line in Dolicho.
[158]
Figure 7-58
Slices 31, 32, and 33 at 90 to the canto-meatal line in Dolicho.
[159]
Hyperextension
Incidence:
Axial Slices
[160]
Figure 7-59
resent a sample of slices obtained at -30 to the canto-meatal
line, which is the incidence found with a hyperextended head.
Underneath are the two hemispheres in lateral and mesial views,
as seen in Figures 2-2 and 2-4, with gyri individually colored.
Lines correspond to slices obtained at 30 to the canto-meatal
line as depicted in this section.
[161]
Figure 7-60
[162]
Figure 7-61
tification of gyri.
[163]
Figure 7-62
[164]
Figure 7-63
tification of gyri.
[165]
Figure 7-64
[166]
Figure 7-65
tification of gyri.
[167]
Figure 7-66
Slices 7 and 8 at 3 0 to the canto-meatal line in Dolicho. Iden-
[168]
Figure 7-67
tification of gyri.
[169]
Figure 7-68
Slices 9 and 10 at 3 0 to the canto-meatal line in Dolicho. Iden-
[170]
Figure 7-69
tification of gyri.
[171]
Figure 7-70
Slices 11 and 12 at 30 to the canto-meatal line in Dolicho.
[172]
Figure 7-71
[173]
Figure 7-72
[174]
Figure 7-73
[175]
Figure 7-74
[176]
Figure 7-75
[177]
Figure 7-76
[178]
Figure 7-77
Slices 17 and 18 at -30 to the canto-meatal line in Dolicho.
[179]
Figure 7-78
Slices 19 and 20 at -30 to the canto-meatal line in Dolicho.
[180]
Figure 7-79
Slices 19 and 20 at 3 0 to the canto-meatal line in Dolicho.
[181]
Figure 7-80
[182]
Figure 7-81
[183]
Figure 7-82
[184]
Figure 7-83
[185]
Figure 7-84
Slices 25, 26, and 27 at -30 to the canto-meatal line in Dolicho.
[186]
Figure 7-85
Slices 25,26, and 27 at 30 to the canto-meatal line in Dolicho.
[187]
Hyperextension
Incidence:
Coronal Slices
[188]
Figure 7-86
Left hemisphere of Dolicho seen through the transparent head on
top. The canto-meatal line is marked in red. Black lines repre-
sent a sample of slices obtained at +60 to the canto-meatal line,
which is the coronal incidence when the axial slices are obtained
with a hyperextended head. Underneath are the two hemispheres
in lateral and mesial views as seen in Figures 2-2 and 2-4, with
gyri individually colored. Lines correspond to slices obtained at
+60 to the canto-meatal line as depicted in this section.
[189]
Figure 7-87
Slices 1 and 2 at +60 to the canto-meatal line in Dolicho. Iden-
[190]
Figure 7-88
tification of gyri.
[191]
Figure 7-89
[192]
Figure 7-90
tification of gyri.
[193]
Figure 7-91
[194]
Figure 7-92
tification of gyri.
[195]
Figure 7-93
[196]
Figure 7-94
tification of gyri.
[197]
Figure 7-95
[198]
Figure 7-96
tification of gyri.
[199]
Figure 7-97
Slices 11 and 12 at +60 to the canto-meatal line in Dolicho.
[200]
Figure 7-98
[201]
Figure 7-99
[202]
Figure 7-100
[203]
Figure 7-101
[204]
Figure 7-102
[205]
Figure 7-103
[206]
Figure 7-104
[207]
Figure 7-105
[208]
Figure 7-106
[209]
Figure 7-107
[210]
Figure 7-108
[211]
Figure 7-109
[212]
Figure 7-110
[213]
Figure 7-111
[214]
Figure 7-112
[215]
Figure 7-113
[216]
Figure 7-114
[217]
Figure 7-115
[218]
Figure 7-116
[219]
Figure 7-117
[220]
Figure 7-118
[221]
Posterior Fossa
Incidence:
Axial Slices
[222]
Figure 7-119
resent a sample of slices obtained at +10 to the canto-meatal
line, which is the incidence of axial slices when posterior fossa
structures are the target. Underneath are the two hemispheres in
lateral and mesial views, as seen in Figures 2-2 and 2-4, with gyri
individually colored. Lines correspond to slices obtained at +10
to the canto-meatal line as depicted in this section.
[223]
Figure 7-120
[224]
Figure 7-121
tification of gyri.
[225]
Figure 7-122
[226]
Figure 7-123
tification of gyri.
[227]
Figure 7-124
[228]
Figure 7-125
tification of gyri.
[229]
Figure 7-126
[230]
Figure 7-127
tification of gyri.
[231]
Figure 7-128
[232]
Figure 7-129
tification of gyri.
[233]
Figure 7-130
[234]
Figure 7-131
[235]
Figure 7-132
[236]
Figure 7-133
[237]
Figure 7-134
[238]
Figure 7-135
[239]
Figure 7-136
[240]
Figure 7-137
[241]
Figure 7-138
[242]
Figure 7-139
[243]
Figure 7-140
[244]
Figure 7-141
[245]
Figure 7-142
Slices 23, 24, and 25 at +10 to the canto-meatal line in Dolicho.
[246]
Figure 7-143
Slices 23, 24, and 25 at +10 to the canto-meatal line in Dolicho.
[247]
Parasagittal Incidence
[248]
Figure 7-144
Dolicho seen from the back and front (top) and from the top and obtained parallel to the interhemispheric fissure depicted in this
bottom (below), as in Figure 2-6. Black lines represent slices section. They begin at the outer surface of the left hemisphere.
[249]
Figure 7-145
Left hemisphere parasagittal slices 1 and 2 in Dolicho. Identifi-
cation of sulci.
[250]
Figure 7-146
cation of gyri.
[251]
Figure 7-147
cation of sulci.
[252]
Figure 7-148
cation of gyri.
[253]
Figure 7-149
cation of sulci.
[254]
Figure 7-150
cation of gyri.
[255]
Figure 7-151
cation of sulci.
[256]
Figure 7-152
cation of gyri.
[257]
Figure 7-153
cation of sulci.
[258]
Figure 7-154
cation of gyri.
[259]
Figure 7-155
Left hemisphere parasagittal slices 11 and 12 in Dolicho. Identi-
fication of sulci.
[260]
Figure 7-156
fication of gyri.
[261]
Figure 7-157
fication of sulci.
[262]
Figure 7-158
fication of gyri.
[263]
Figure 7-159
fication of sulci.
[264]
Figure 7-160
fication of gyri.
[265]
Figure 7-161
fication of sulci.
[266]
Figure 7-162
fication of gyri.
[267]
Figure 7-163
Right hemisphere parasagittal slices 19 and 20 in Dolicho. Iden-
[268]
tification of gyri.
[269]
Figu re 7-16 4
Figure 7-165
[270]
Figure 7-166
tification of gyri.
[271]
Figure 7-167
[272]
Figure 7-168
tification of gyri.
[273]
Figure 7-169
[274]
Figure 7-170
tification of gyri.
[275]
Figure 7-171
[276]
Figure 7-172
tification of gyri.
[277]
Figure 7-173
[278]
Figure 7-174
tification of gyri.
[279]
Figure 7-175
[280]
Figure 7-176
tification of gyri.
[281]
Figure 7-177
[282]
Figure 7-178
tification of gyri.
[283]
Figure 7-179
[284]
Figure 7-180
tification of gyri.
[285]
Brodmann's Fields
[286]
Figure 7-181
Right and left hemispheres of Dolicho in mesial and lateral views
with the gyri colored and Brodmann's cytoarchitectonic fields
marked, as in Figures 2-7 and 2-8. Black lines correspond to
axial slices obtained parallel to the canto-meatal line, as in Fig-
ure 7-1, corresponding to the slices depicted in this section.
[287]
Figure 7-182
Slices 1-6 parallel to the canto-meatal line with Brodmann's
fields in Dolicho.
[288]
Figure 7-183
fields marked in Dolicho.
[289]
Figure 7-184
[290]
Figure 7-185
[291]
T
he main goal of this chapter is to show that, in different individuals, axial se-
quences obtained with the same incidences can produce d i f f er en t images. The
procedures are the same used for Dolicho. The two most typical sequences used in
magnetic resonance (MR) and computed tomographic studies are shown: parallel
to the canto-meatal line (Figs. 8-1 to 8-25) and the 30-degree rostral angulation
sequence obtained with the hyperextended head (Figs. 8-57 to 8-81). As in the pre-
vious chapter, a set of coronal sections to each of the axial sequences is also shown
(Figs. 8-26 to 8-56 and 8-82 to 8-110). The posterior fossa incidence with a cau-
dal 10 tilt is shown in Figures 8-111 to 8-135. Again, sulci are marked on the left-
hand side and gyri, on the right. The MR cuts from which the images were ob-
tained and the levels of the cuts are depicted as in Chapter 7. The first image of
each sequence shows, as in Chapter 7, the two hemispheres seen from the dorso-
lateral and mesial views, with a superimposed grid corresponding to the slices to
follow. The brain images used here are the same as those in Figures 3-2, 3-4, and
3-6. Above is the image of the transparent head and the left hemisphere of Brachi-
1, with the canto-meatal line drawn in red and a sample of the slices in the section.
The comments made in Chapter 7 apply here too: the relative appearance
among the different structures depends on the incidence of the cut. The reader
should compare the slices in one sequence to those in the others. It is also im-
portant to compare the images seen in this chapter to those obtained with the
same incidence in Chapter 7. See, for instance, the first axial incidence in these
two chapters, parallel to the canto-meatal line. Compare the slices that first show
the calcarine fissure, slice 10 in Figures 8-10 and 8-11 and slice 11 in Figures
7-12 and 7-13. There is a different overall configuration of the slice (Brachi-1 is
rounder than Dolicho) but also considerable similarity in the visible structures,
particularly in the frontal and temporal lobes. The structures in the occipital lobe,
however, are quite different. Slice 10 in this chapter barely touches the occipital
lobe, while the corresponding cut in Chapter 7 (slice 11 in Figs. 7-12, 7-13) al-
most reaches the cuneus (which is visible in slice 12). In Brachi-1, we must ad-
vance to slice 13 (Fig. 8-15) to catch the very first sliver of the cuneus.
Chapter 8
Sections through
Brachi-1
[293]
Canto-meatal
Incidence: Axial Slices
[294]
Figure 8-1
Left hemisphere of Brachi-1 seen through the transparent head
resent a sample of slices obtained parallel to the canto-meatal
ored. Lines correspond to slices parallel to the canto-meatal line
as depicted in this section.
[295]
Figure 8-2
Slices 1 and 2 parallel to the canto-meatal line in Brachi-1. Iden-
[296]
Figure 8-3
tification of gyri.
[297]
Figure 8-4
[298]
Figure 8-5
tification of gyri.
[299]
Figure 8-6
[300]
Figure 8-7
tification of gyri.
[301]
Figure 8-8
[302]
Figure 8-9
tification of gyri.
[303]
Figure 8-10
[304]
Figure 8-11
tification of gyri.
[305]
Figure 8-12
Slices 11 and 12 parallel to the canto-meatal line in Brachi-1.
[306]
Figure 8-13
[307]
Figure 8-14
[308]
Figure 8-15
[309]
Figure 8-16
[310]
Figure 8-17
[311]
Figure 8-18
[312]
Figure 8-19
[3131
Figure 8-20
[314]
Figure 8-21
[315]
Figure 8-22
[316]
Figure 8-23
[317]
Figure 8-24
[318]
Figure 8-25
[319]
Canto-meatal
Incidence: Coronal
Slices
[320]
Figure 8-26
resent a sample of slices obtained at 90 to the canto-meatal line,
parallel to the canto-meatal line. Underneath are the two hemi-
spheres in lateral and mesial views, as seen in Figures 3-2 and
3-4, with gyri individually colored. Lines correspond to slices ob-
tained at 90 to the canto-meatal line as depicted in this section.
[321]
Figure 8-27
Slices 1 and 2 at 90 to the canto-meatal line in Brachi-1. Iden-
[322]
Figure 8-28
tification of gyri.
[3231
Figure 8-29
[324]
Figure 8-30
tification of gyri.
[325]
Figure 8-31
[326]
Figure 8-32
tification of gyri.
[327]
Figure 8-33
[328]
Figure 8-34
tification of gyri.
[329]
Figure 8-35
[330]
Figure 8-36
tification of gyri.
[331]
Figure 8-37
[332]
Figure 8-38
tification of gyri.
[333]
Figure 8-39
[334]
Figure 8-40
tification of gyri.
[335]
Figure 8-41
[336]
Figure 8-42
tification of gyri.
[337]
Figure 8-43
[338]
Figure 8-44
tification of gyri.
[339]
Figure 8-45
[340]
Figure 8-46
tification of gyri.
[341]
Figure 8-47
[342]
Figure 8-48
tification of gyri.
[343]
Figure 8-49
[344]
Figure 8-50
tification of gyri.
[345]
Figure 8-51
[346]
Figure 8-52
tification of gyri.
[347]
Figure 8-53
[348]
Figure 8-54
tification of gyri.
[349]
Figure 8-55
[350]
Figure 8-56
tification of gyri.
[351]
Hyperextension
[352]
Figure 8-57
resent a sample of slices obtained at 30 to the canto-meatal
as seen in Figures 32 and 3-4, with gyri individually colored.
Lines correspond to slices obtained at 30 to the canto-meatal
line as depicted hi this section.
[353]
Figure 8-58
Slices 1, 2, 3, and 4 at 30 to the canto-meatal line in Brachi-
1. Identification of sulci.
[354]
Figure 8-59
1. Identification of gyri.
[355]
Figure 8-60
[356]
Figure 8-61
tification of gyri.
[357]
Figure 8-62
[358]
Figure 8-63
tification of gyri.
[359]
Figure 8-64
[360]
Figure 8-65
tification of gyri.
[361]
Figure 8-66
Slices 11 and 12 at 30 to the canto-meatal line in Brachi-1.
[362]
Figure 8-67
[363]
Figure 8-68
[364]
Figure 8-69
Slices 13 and 14 at -30 to the canto-meatal line in Brachi-1.
[365]
Figure 8-70
Slices 15 and 16 at 3 0 to the canto-meatal line in Brachi-1.
[366]
Figure 8-71
[3671
Figure 8-72
[368]
Figure 8-73
[369]
Figure 8-74
[370]
Figure 8-75
[371]
Figure 8-76
[372]
Figure 8-77
[373]
Figure 8-78
[374]
Figure 8-79
[375]
Figure 8-80
Slices 25, 26 and 27 at 30 to the canto-meatal line in Brachi-
[376]
Figure 8-81
Slices 25, 26 and 27 at -30 to the canto-meatal line in Brachi-
[3771
Hyperextension
Incidence:
Coronal Slices
[378]
Figure 8-82
Left hemisphere of Brachi-1 seen through the transparent head on
sent a sample of slices obtained at +60 to the canto-meatal line,
with a hyperextended head. Underneath are the two hemispheres
in lateral and mesial views, as seen in Figures 3-2 and 3-4, with
gyri individually colored. Lines correspond to slices obtained at
+60 to the canto-meatal line as depicted in this section.
[379]
Figure 8-83
Slices 1 and 2 at +60 to the canto-meatal line in Brachi-1. Iden-
[380]
Figure 8-84
tification of gyri.
[381]
Figure 8-85
[382]
Figure 8-86
tification of gyri.
[383]
Figure 8-87
[384]
Figure 8-88
tification of gyri.
[385]
Figure 8-89
[386]
Figure 8-90
tification of gyri.
[387]
Figure 8-91
[388]
Figure 8-92
tification of gyri.
F3891
Figure 8-93
Slices 11 and 12 at +60 to the canto-meatal line in Brachi-1.
[390]
Figure 8-94
[391]
Figure 8-95
[392]
Figure 8-96
[393]
Figure 8-97
[394]
Figure 8-98
[395]
Figure 8-99
[396]
Figure 8-100
[397]
Figure 8-101
[398]
Figure 8-102
[399]
Figure 8-103
[400]
Figure 8-104
[401]
Figure 8-105
[402]
Figure 8-106
f4031
Figure 8-107
[404]
Figure 8-108
[405]
Figure 8-109
Slices 27, 28, and 29 at +60 to the canto-meatal line in Brachi-
[406]
Figure 8-110
Slices 27, 28, and 29 at +60 to the canto-meatal line in Brachi-
[407]
Posterior Fossa
[408]
Figure 8-111
Left hemisphere of Brachi-1 seen through the transparent head on
sent a sample of slices obtained at +10 to the cantomeatal line,
which is the incidence of axial slices obtained when posterior
fossa structures are the target. Underneath are the two hemi-
spheres in lateral and mesial views, as seen in Figures 3-2 and 3-4,
with gyri individually colored. Lines correspond to slices obtained
at +10 to the canto-meatal line as depicted in this section.
[409]
Figure 8-112
[410]
Figure 8-113
tification of gyri.
[411]
Figure 8-114
[412]
Figure 8-115
tification of gyri.
[413]
Figure 8-116
[414]
Figure 8-117
tification of gyri.
[415]
Figure 8-118
[416]
Figure 8-119
tification of gyri.
[417]
Figure 8-120
[418]
Figure 8-121
tification of gyri.
[419]
Figure 8-122
[420]
Figure 8-123
[421]
Figure 8-124
[422]
Figure 8-125
[423]
Figure 8-126
[424]
Figure 8-127
[425]
Figure 8-128
[426]
Figure 8-129
[427]
Figure 8-130
[428]
Figure 8-131
[429]
Figure 8-132
[430]
Figure 8-133
[431]
Figure 8-134
[432]
Figure 8-135
[433]
T
he three axial sequences used in the previous two chapters were obtained
in Brachi-2. This chapter illustrates how incidences chosen in relation to
external landmarks reveal remarkably different structures in subjects with dif-
ferent cranial configurations. Let us continue with the example chosen for the
previous chapter, namely, the comparison among slices parallel to the canto-
meatal line. Once again, compare the slices in which the calcarine sulcus first
appears (Figs. 9-12, 9-13 and Figs. 7-12, 7-13 and 8-10, 8-11). In Figs. 9-12
and 9-13, slice 11, the most anterior segment of the calcarine sulcus, shows up
for the first time. This slice shows the corpus callosum anteriorly, sliced at the
very top of the beak of the callosum, together with the superior regions of the
frontal operculum. On the other hand, in Dolicho (Figs. 7-12, 7-13), the cal-
losum is cut in the middle of the beak and the frontal operculum is sliced
through the middle portion rather than through its superior sector. We must
advance to Figure 7-17 (slice 16) to see the superior regions of the pars oper-
cularis. In fact, in Brachi-2, slices 11 and 12 of the axial sequence parallel to the
canto-meatal line are much closer to slices 11 and 12 in Figure 7-132, the pos-
terior fossa incidence for Dolicho.
As in the two preceding chapters, sulci are marked on the left page and gyri,
on the right. The magnetic resonance images from which the anatomic images
were obtained and the levels of the cuts are as in Chapter 7. The gyri images
used to illustrate the setting of the slices in each sequence are those used in
Chapter 4 (Figs. 4-2, 4-4, and 4-6).
Chapter 9
Sections through
Brachi-2
[435]
Canto-meatal
[436]
Figure 9-1
resent a sample of slices obtained parallel to the canto-meatal
ored. Lines correspond to slices parallel to the canto-meatal line
as depicted in this section.
[437]
Figure 9-2
[438]
Figure 9-3
tification of gyri.
[439]
Figure 9-4
[440]
Figure 9-5
tification of gyri.
[441]
Figure 9-6
[442]
Figure 9-7
tification of gyri.
[443]
Figure 9-8
[444]
Figure 9-9
tification of gyri.
[445]
Figure 9-10
[446]
Figure 9-11
tification of gyri.
[447]
Figure 9-12
[448]
Figure 9-13
[449]
Figure 9-14
[450]
Figure 9-15
[451]
Figure 9-16
[452]
Figure 9-17
[453]
Figure 9-18
[454]
Figure 9-19
[455]
Figure 9-20
[456]
Figure 9-21
[457]
Figure 9-22
[458]
Figure 9-23
[459]
Figure 9-24
[460]
Figure 9-25
[461]
Hyperextension
[462]
Figure 9-26
resent a sample of slices obtained at 30 to the canto-meatal
as seen in Figures 4-2 and 4-4, with gyri individually colored.
Lines correspond to slices obtained at -30 to the canto-meatal
line as depicted in this section.
[463]
Figure 9-27
[464]
Figure 9-28
Slices 1, 2, 3, and 4 at -30 at +60 to the canto-meatal line in
Brachi-2. Identification of gyri.
[4651
Figure 9-29
[466]
Figure 9-30
tification of gyri.
F4671
Figure 9-31
[468]
Figure 9-32
tification of gyri.
[469]
Figure 9-33
[470]
Figure 9-34
tification of gyri.
[471]
Figure 9-35
[472]
Figure 9-36
[473]
Figure 9-37
[474]
Figure 9-38
[475]
Figure 9-39
[476]
Figure 9-40
[477]
Figure 9-41
[478]
Figure 9-42
[479]
Figure 9-43
[480]
Figure 9-44
[481]
Figure 9-45
[482]
Figure 9-46
[483]
Figure 9-47
[484]
Figure 9-48
[485]
Figure 9-49
Slices 25, 26, and 27 at 3 0 to the canto-meatal line in Brachi-
[486]
Figure 9-50
Slices 25, 26, and 27 at 30 to the canto-meatal line in Brachi-
[487]
Posterior Fossa
[488]
Figure 9-51
resent a sample of slices obtained at + 10 to the canto-meatal
line, which is the incidence of axial slices obtained when poste-
rior fossa structures are the target. Underneath are the two hemi-
spheres in lateral and mesial views, as seen in Figures 4-2 and
4-4, with gyri individually colored. Lines correspond to slices ob-
tained at + 10 to the canto-meatal line as depicted in this section.
[489]
Figure 9-52
[490]
Figure 9-53
tification of gyri.
[491]
Figure 9-54
[492]
Figure 9-55
tification of gyri.
[493]
Figure 9-56
[494]
Figure 9-57
tification of gyri.
[495]
Figure 9-58
[496]
Figure 9-59
tification of gyri.
[497]
Figure 9-60
[498]
Figure 9-61
tification of gyri.
[499]
Figure 9-62
[500]
Figure 9-63
[501]
Figure 9-64
[502]
Figure 9-65
[503]
Figure 9-66
[504]
Figure 9-67
[505]
Figure 9-68
[506]
Figure 9-69
[507]
Figure 9-70
Slices 19 and 20 at +10 to the canto-meata! line in Brachi-2.
[508]
Figure 9-71
[509]
Figure 9-72
[510]
Figure 9-73
[511]
Figure 9-74
[512]
Figure 9-75
[513]
Chapter 10
Application to
Lesion Studies
T
his chapter aims at demonstrating how variability due to different inci-
dences of cut can play a role in the determination of lesion location. I use
four examples of relatively circumscribed lesions in distinct locations: left pari-
etal, left calcarine, right temporal, and left frontal. All lesions resulted from
stroke.
Two or more incidences of sections are given for each case to make the point
that quick inspection is unlikely to yield a correct reading. Also shown are a
transparent head and the appropriate hemisphere with lines marking the slices.
The sulci most relevant for a correct interpretation of the lesion location are
identified.
[515]
Figures 10-1 and 10-2 show a left parietal infarct seen from the left, through a
transparent head, in slices parallel to the canto-meatal line (Fig. 10-1 A) and the
corresponding coronal slices (Fig. 10-2A). Figures 10-15 and 10-25 show the
sort of axial and coronal slices that would be revealed if the head had been hy-
perextended.
The images obtained parallel to the canto-meatal line and perpendicular to
it would probably be interpreted as showing damage in the left parietal lobe (in
the inferior parietal lobule, mostly in the supramarginal gyrus). By contrast, the
images obtained with a negative rostral tiltthe "hyperextension images" (Fig.
1015)could easily be misinterpreted as showing damage in the posterior
temporo-occipital region, possibly extending into the angular gyrus. An erro-
neous reading would arise from the relation of the lesion to the posterior re-
gion of the lateral ventricle (at a point at which the frontal lobe barely shows
up anteriorly). Observation of the coronal slices (Fig. 10-25) might dispel the
erroneous reading given that the lesion is located very high in each cut. How-
ever, the fact that it is seen in a coronal slice behind the splenium of the cor-
pus callosum and at the posterior end of the cerebellum and beyond might still
suggest that it was located in the occipital lobe. Two clues should help the reader
reach the correct interpretation. First, cuts 1 and 2 show the very anterior end
of the calcarine fissure, prior to it being joined by the occipitoparietal sulcus
(the juncture occurs only in cut 3; from that point on, the sulcus that is seen
mesially is the occipitoparietal sulcus, with the calcarine fissure disappearing
inferiorly). Second, the intraparietal sulcus appears in cut 5. This reveals that
the lesion is lateral to this sulcus and therefore not in the occipital lobe but
rather in the parietal.
[517]
A Left Parietal Lesion
(Figs. 10-1, 10-2)
Figure 10-1
Two axial incidences in the brain of a 53-year-old, fully right-
handed woman with a left parietal infarct. The lesion is seen
through the transparent head on the surface of the left hemi-
sphere in the three-dimensionally reconstructed brain. The
canto-meatal line is drawn as a dotted line. The position of the
slices (seen on the right) is marked with solid lines. The most im-
portant sulci needed to localize the lesion are identified and
marked. (A) Slices obtained parallel to the canto-meatal line. (B)
Slices obtained in hyperextension.
Figure 10-2
Coronal slices corresponding to the axial slices seen in Figure
10-1.
[519]
As in the previous example, I chose two axial orientationsparallel to the canto-
meatal line (Fig. 10-3A) and with a negative rostral tilt (Fig. 10-35)along
with their corresponding coronal cuts (Figs. 10-4A, 10-45) and the mesial view
of the left hemisphere (seen through a transparent head from the right). When
we examine the axial cuts, regardless of the incidence, there is no doubt that
the lesion is in the calcarine region. However, if we begin by inspecting Figure
10-35, it would be possible to read these images as if the lesion were placed
more anteriorly in the occipital lobe. It would even be possible to think about
a mesial temporo-occipital location rather then a purely occipital one.
Inspection of the coronal slices in either panel A or B of Figure 10-4 should
leave no doubt that the lesion does involve the calcarine fissure in its anterior
sector. At some point, regardless of the angulation to the canto-meatal line,
coronal slices will show the joining of the calcarine fissure and the occipi-
toparietal sulcus in a characteristic shape (><), in which the upper arms of
the V shapes (><) corresponds to the occipitoparietal sulcus and the lower to
the calcarine fissure (slices 1 and 2 in A, slices 3 and 4 in 5). Inspection of these
images shows that the damage involves the lower arm of the left side, the cal-
carine fissure, and that it does so at the juncture of the two and farther back.
[521]
A Left Calcarine Lesion
(Figs. 10-3, 10-4)
Figure 10-3
Two axial incidences in the brain of a 42-year-old, fully right-
handed man with a left calcarine infarct. The lesion is seen
through the transparent head on the mesial surface of the left
hemisphere in the three-dimensionally reconstructed brain. The
canto-meatal line is drawn as a dotted line. The position of the
slices (seen on the right) is marked with solid lines. The most im-
portant sulci needed to localize the lesion are identified and
Slices obtained in hyperextension.
Figure 10-4
Coronal slices corresponding to the axial slices seen in Figure
10-3.
[523]
Figures 10-5 and 10-6 show a right posterior temporal lobe lesion seen through
a transparent head from the right. Figure 105 shows the axial slices parallel to
the canto-meatal line (A) and the corresponding coronal slices (B). The inter-
pretation of these images is straightforward because the incidence is close to
that of images in traditional anatomic atlases. The position of the lesion in A
cannot possibly be interpreted as anything but the posterior temporal region.
The lesion is at the edge of the temporal lobe, behind the superior temporal
sulcus, next to the cerebellum, and behind a very anterior image of the hip-
pocampus; the amygdalae are still visible, and the anterior cingulate begins to
be revealed. The subsequent two slices simply confirm the first impression.
The coronal slices in Figure 10-55, showing the lesion at the level of the
splenium and tail end of the hippocampus (slices 1 and 2) just below the trigone
(slices 2 and 3), do not leave much doubt about the location of the lesion. On
the other hand Figure 10-6 may raise questions. In Figure 10-6A, I show three
slices of a posterior fossa incidence. In spite of the fact that the lesion is seen
in slices that cut the frontal lobe relatively high up, above the beak of the cor-
pus callosum, the interpretation is not difficult. The lack of ambiguity is due,
once again, to the recognizable sylvian fissure, superior temporal sulcus, and
hippocampus. Visualization of the cerebellum also helps: the amount of cere-
bellum seen along with the obviously high cut through the frontal lobe indi-
cate that the slice was obtained with a posterior fossa incidence. The interpre-
tation would be less clear if the lesion were placed in the parietal lobe (see the
higher cuts in the posterior fossa sequences in Chapters 7-9).
In Figure 10-65, however, it would be more difficulty to decide where in
the temporal lobe the lesion is located. The negative rostral tilt (hyperextension
incidence) may suggest that all we see in these slices is a long longitudinal cut
through the temporal lobe, perhaps with some occipital structures appearing at
the tail end of the slice. This erroneous reading would make one assume that
the lesion is in the anterior temporal lobe, just behind the temporal pole. How-
ever, as we have seen, this is not the case. The non-canonical presentation of
the temporal and occipital structures would lead to a misinterpretation. A cor-
rect reading depends on recognizing the following: (1) that slice 3 shows the
amygdalae and the hippocampus in a long longitudinal sequence, (2) that in
this same slice only the very top of the cerebellar vermis is seen, and (3) that
not even a hint of frontal lobe is seen. These features tell us that these slices
were obtained with a negative rostral tilt, thus revealing the occipital lobe even
in slice 1.
[525]
A Right Temporal Lesion
(Figs. 10-5, 10-6)
Figure 10-5
Axial and coronal slices in the brain of a 31-year-old, fully right-
handed woman with a right posterior temporal infarct. The
lesion is seen through the transparent head on the surface of the
right hemisphere in the three-dimensionally reconstructed brain.
The canto-meatal line is drawn as a dotted line. The position of
the slices (seen on the right] is marked with solid lines. The most
important sulci needed to localize the lesion are identified and
Corresponding coronal slices.
Figure 10-6
Two sets of axial slices in the same brain seen in Figure 10-5.
(A) Slices obtained with a posterior fossa incidence. (B) Slices
obtained in hyperextension.
[527]
Figures 10-7 and 10-8 show the surface projection of a left subcortical frontal
lesion seen through a transparent head on the left hemisphere as a faint gray
shadow. The lesion was traced in Brainvox and would not have been visible
from mere inspection of the lateral surface of the hemisphere.
Figure 10 7 shows the coronal slices (A) in which the scan was obtained and
which correspond to axial cuts (B) obtained parallel to the inferior orbito-
meatal line rather than the canto-meatal line, a rostral 10 tilt from the lat-
ter. In these slices, after the sulci in the left hemisphere have been identified, it
becomes apparent that the lesion is in the white matter beneath the precentral
gyrus, at the level of the inferior frontal sulcus.
In Figure 10-8A, we see two slices obtained in hyperextension and, in B,
slices obtained with a posterior fossa incidence. Without the help of a three-
dimensionally reconstructed brain, it is difficult to identify the sulci. One might
be tempted to think that in A the lesion is in the parietal lobe and in B it might
be in the anterior frontal region. Comparison of the images in A with those in
Figures 8-75 and 8-76 or 9-43 and of the images in B with Figures 8-122 and
9-62 of the previous chapters would help to reach the correct interpretation
better than a comparison with the corresponding images in Chapter 7. Even so,
unequivocal identification of sulci would be best achieved with the help of a
three-dimensional image and transfer of such identification onto the two-
dimensional slices, as demonstrated here.
A Left Frontal Lesion
(Subcortical)
(Figs. 10-7, 10-8)
[529]
Figure 10-7
Coronal and axial slices in the brain of a 41-year-old, fully right-
handed woman with a left subcortical frontal infarct. The lesion
is projected onto the surface of the left hemisphere in the three-
dimensionally reconstructed brain seen through the transparent
head. The canto-meatal line is drawn as a dotted line. The posi-
tion of the slices (seen on the right) is marked with solid lines.
The most important sulci needed to localize the lesion are iden-
tified and marked. (A) Coronal slices obtained during the native
magnetic resonance acquisition, which is perpendicular to the
axial slices (), obtained parallel to the inferior orbito-meatal
line, a rostral 10 to the canto-meatal line.
[530]
Figure 10-8
Two sets of axial slices in the same brain seen in Figure 10-7. (A)
Axial slices obtained in hyperextension. (B) Axial slices obtained
in a posterior fossa incidence.
[531]
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[536]
Human Brain Anatomy
All pages in which a given structure appears in Chapters 1-4, 6, and 10 are noted. The structures that
appear in Chapter 5 are noted as 50-75. For structures that appear in Chapters 7-9 only the first page in
which they appear, in each sequence, is noted. The exception is the parasagital sequence in Chapter 7 in
which the first appearance of structure is noted both for the left and the right hemispheres.
Amygdala (a), 109, 141, 173, 201, 229, 261,
273, 301, 335, 363, 391, 415, 443, 473,
495, 526, 527
Anterior commisure (ac), 21, 35, 43, 111,
141, 177, 203, 305, 335, 391
Anterior occipital sulcus (AOS), 18, 22, 32,
36, 40, 44, 50-75, 108, 152, 166, 212,
234, 252, 278, 302, 344, 358, 400, 418,
448, 468, 500, 526, 527
Calcarine fissure (CalcF), 8, 20, 22, 34, 36,
42, 44, 50-75, 112, 148, 166, 210, 234,
262, 268, 304, 342, 360, 398, 420, 446,
468, 500, 519, 522, 523
Caudate nucleus
body of caudate nucleus (be), 117, 141,
203, 237, 263, 271, 307, 335, 369, 393,
421, 449, 479, 501
head of caudate nucleus (he), 111, 137,
177, 199, 231, 263, 271, 303, 331, 365,
389, 417, 447, 475, 497
tail of caudate nucleus (tc), 113, 174, 209,
239, 263, 273, 305, 339, 399, 419, 479
Central sulcus (CS), 8, 18, 20, 22, 32, 34,
36, 40, 42, 44, 50-75, 87, 114, 138, 178,
202, 234, 250, 268, 306, 334, 368, 392,
420, 448, 476, 498, 518, 519, 530, 531
Cerebellum
cerebellar hemispheres (cerebH), 19, 21,
23, 33, 35, 37, 41, 43, 45, 103, 149,
163, 207, 225, 255, 269, 297, 341, 355,
395, 413, 439, 465, 493
cerebellar tonsils (cerebT), 21, 35, 43,
103, 151, 163, 207, 227, 265, 269, 297,
345, 355, 397, 413, 439, 465, 493
cerebellar vermis (cerebV), 21, 35, 43,
111, 151, 165, 211, 229, 267, 269,
299, 343, 355, 399, 415, 443, 465,
497
Cingulate gyrus (CingG), 21, 35, 87, 109,
133, 177, 195, 231, 265, 269, 303, 327,
367, 385, 415, 477, 495
Cingulate sulcus (CingS), 8, 20, 34, a42,
50-75, 87, 108, 132, 176, 194, 228, 264,
268, 302, 324, 366, 384, 414, 444, 476,
492, 522, 523
ascending ramus of cingulated sulcus
(arCingS), 8, 20, 22, 34, 36, 42, 44,
50-75, 120, 148, 182, 212, 244, 264,
314, 340, 372, 400, 428, 456, 482, 508,
522, 523
Circular sulcus (circS,*), 108, 138, 174, 200,
230, 302, 334, 362, 388, 416, 444, 472,
494
Claustrum (3), 111, 139, 177, 199, 233, 305,
333, 365, 389, 417, 445, 497
Collateral sulcus (ColS), 20, 22, 34, 36, 42,
44, 50-75, 104, 140, 166, 200, 226, 258,
270, 298, 336, 358, 390, 412, 438, 468,
470, 522, 523, 526, 527
Colliculi
inferior colliculi (ic), 111, 149, 173, 209,
233, 267, 269, 305, 343, 397, 419, 447,
473, 499
superior colliculi (sc), 111, 147, 173, 209,
235, 267, 269, 305, 341, 397, 421, 447,
473,501
Corpus callosum (CC), 21, 35, 43, 115, 139,
177, 201, 235, 267, 269, 307, 333, 369,
391, 421, 449, 479, 499
Index of Anatomical
Structures Seen in
the Figures
[537]
Human Brain Anatomy
Corpus callosum (continued)
beak of callosum (bcc), 21, 35, 43, 111,
137, 179, 199, 233, 265, 269, 305, 331,
367, 389, 447, 477, 497
splenium of callosum (sec), 21, 35, 43,
113, 147, 175, 209, 237, 265, 269, 307,
341, 365, 399, 423, 451, 475, 503, 526
Cuneus (Gun), 21, 35, 43, 113, 155, 167,
215, 237, 263, 269, 309, 345, 361, 403,
425, 451, 471, 505
Fornix (for), 21, 35, 43, 141, 177, 203, 235,
267, 269, 307, 335, 367, 393, 419, 447,
499
Frontal lobe (FL), 9, 87
Frontal pole (FP), 19, 21, 23, 33, 35, 37, 41,
43, 45, 107, 129, 179, 191, 227, 257,
269, 301, 323, 367, 381, 413, 443, 477,
491
Frontomarginal sulcus (FmS), 22, 36, 108,
128, 178, 190, 228, 258, 270, 302, 322,
370, 380, 412
Fusiform gyrus (FusiG), 21, 23, 35, 37, 43,
45, 111, 151, 169, 211, 235, 259, 273,
305, 345, 359, 401, 421, 447, 469, 499
Gyrus rectus (Grec), 21, 23, 35, 37, 43, 45,
107, 133, 175, 193, 227, 267, 269, 299,
325, 363, 383, 411, 441, 473, 491
Heschl's gyrus (HG), 27, 38, 46, 113, 143,
177, 205, 235, 255, 277, 305, 337, 367,
395, 419, 447, 475, 497
H shaped sulci (HsS), 20, 22, 34, 36, 42, 44,
130, 174, 192, 228, 258, 270, 300, 324,
364, 380, 412, 442, 472, 492
Hippocampal fissure (HipF), 108
Hippocampus (hip), 109, 143, 173, 203,
231, 259, 273, 301, 337, 361, 393, 415,
443, 471, 495, 526, 527
Hypothalamus (ht), 111, 143, 175, 203, 233,
265, 269, 335, 393, 417, 447, 475, 497
Inferior frontal gyrus (IFG), 19, 23, 33, 37,
41,45
pars opercularis (IFG,po), 19, 33, 41, 113,
139, 177, 199, 233, 253, 281, 305, 329,
367, 389, 417, 447, 477, 497
pars orbitalis (IFG.porb), 19, 33, 41, 109,
129, 177, 193, 229, 255, 281, 303, 327,
365, 383, 413, 445, 477, 495
pars triangularis (IFG,pt), 19, 33, 41, 111,
133, 177, 195, 231, 255, 279, 303, 325,
367, 383, 415, 445, 477, 495
Inferior frontal sulcus (IPS), 18, 22, 32, 36,
40, 44, 50-75, 108, 128, 180, 190, 228,
254, 276, 300, 326, 366, 382 ,414, 446,
494, 527, 530, 531
Inferior parietal lobule (IPL), 19, 33, 37, 41,
45
angular gyrus (AG), 19, 23, 33, 37, 41,
45, 115, 149, 171, 211, 237, 253, 275,
311, 343, 365, 403, 427, 451, 475, 505
supra-marginal gyrus (SMG), 19, 23, 33,
37, 41, 45, 115, 143, 177, 209, 239, 251,
279, 309, 337, 369, 397, 423, 451, 477,
503
Inferior temporal gyrus (ITG), 19, 21, 23,
33, 35, 37, 41, 43, 45, 107, 141, 169,
199, 227, 253, 279, 297, 335, 359, 389,
411,439,469,491
Inferior temporal sulcus (ITS), 18, 22, 32,
36, 40, 44, 50-75, 104, 140, 168, 198,
226, 252, 280, 296, 332, 360, 386, 410,
438, 468, 490, 526, 527
Insula (Ins), 9, 87
anterior gyri (1), 29, 109, 139, 177, 199,
231, 259, 277, 303, 331, 365, 389, 417,
445, 475, 497
anterior sulci (ants), 28
posterior gyri (2), 29, 109, 141, 175, 203,
231, 257, 277, 303, 335, 365, 391, 417,
445, 475, 497
posterior sulci (posts), 28
Internal capsule
anterior (aic), 111, 141, 177, 203, 233,
265, 271, 305, 335, 367, 391, 419, 447,
477, 497
posterior (pic), 113, 141, 177, 205, 235,
265, 271, 305, 337, 367, 395, 419, 447,
477, 497
Intra-parietal sulcus (IPS), 18, 22, 32, 36,
40, 44, 50-75, 116, 146, 172, 214, 256,
272, 310, 340, 364, 402, 424, 452, 476,
506, 518
Lateral occipital gyri (LOG), 19, 23, 33, 37,
41, 45, 109, 153, 165, 213, 235, 253,
273, 305, 347, 359, 403, 421, 447, 469,
501
Lateral occipital sulcus (LOS), 18, 22, 32,
36, 40, 44, 114, 150, 170, 212, 236, 252,
278, 308, 346 ,404, 426, 446, 472, 502,
526
Lateral temporo-occipital sulcus (LTOS),
20, 22, 34, 36, 42, 44, 104, 140, 168,
196, 224, 258, 274, 296, 332, 358, 388,
410, 440, 468, 492, 526, 527
Lingual gyrus (LingG), 21, 23, 35, 37, 43,
45, 111, 149, 167, 211, 235, 263, 269,
305, 343, 361, 399, 419, 447, 469 ,499
Lingual sulcus (LingS), 20, 22, 34, 36, 42,
110, 152, 166, 212, 234, 264, 268, 306,
344, 362, 400, 420, 448, 468, 500
Mamillary body (mb), 21, 35, 109, 445, 473
Medulla (med), 19, 21, 23, 33, 35, 37, 41,
43, 45, 103, 149, 163, 205, 225, 2a67,
269, 297, 341, 355, 395, 411, 439, 465,
491
Mesencephalon (mes), 21, 35, 43, 111, 145,
173, 205, 233, 265, 269, 303, 339, 363,
395, 417, 473
Middle frontal gyrus (MFC), 19, 23, 33, 37,
41, 45, 109, 129, 181, 193, 229, 255,
275, 305, 325, 383, 415, 445, 479, 495
[538]
Middle frontal sulcus (MFS), 40, 44, 110,
446, 480, 494
Middle temporal gyrus (MTG), 19, 23, 33,
37, 41, 45, 107, 141, 171, 199, 227, 251,
281, 299, 333, 361, 365, 389, 413, 441,
471, 491
Midfrontal sulcus (mFS), 18, 22, 32, 36, 40,
114, 128, 184, 194, 234, 262, 270, 312,
324, 386, 424, 502
Occipital lobe (OL), 9, 87
Occipital pole (OP), 23, 37, 45
Occipitoparietal sulcus (OPS), 8, 20, 22, 34,
42, 50-75, 114, 152, 174, 214, 238, 264,
268, 308, 344, 364, 402, 424, 450, 474,
504, 519, 522, 523
Olfactory sulcus (OlfS), 50-75, 106, 134
Orbital frontal gyri
anterior orbital gyrus (aOrbG), 21, 23,
35, 37, 43, 45, 109, 133, 177, 193, 227,
259, 273, 301, 327, 367, 383, 413, 443,
477, 493
lateral orbital gyrus (lOrbG), 21, 23, 35,
37, 43, 45, 109, 133, 177, 193, 229, 255,
277, 301, 325, 365, 383, 413, 443, 477,
493
middle orbital gyrus (mOrbG), 19, 23,
35, 37, 43, 45, 107, 133, 175, 193, 227,
263, 269, 299, 325, 365, 383, 413, 441,
473, 491
posterior orbital gyrus (pOrbG), 21, 23,
35, 37, 43, 45, 109, 135, 175, 195, 229,
259, 273, 301, 331, 363, 385, 413, 443,
473, 493
Orbital frontal sulci
lateral orbital sulcus (LOrbS), 18, 22, 32,
36, 40, 44, 110, 130, 176, 192, 228, 254,
278, 300, 322, 366, 382, 412, 444, 478,
494
mid orbital sulcus (mOrbS), 20, 22, 34,
36, 42, 44, 106, 130, 174, 190, 226, 266,
268, 298, 324, 362, 380, 412, 440, 472,
490
suborbital sulcus (sOrbS), 20, 34, 42, 108,
128, 176, 190, 228, 322, 366, 380, 412,
442, 476, 494
Pallidum (pa), 111, 141, 177 ,203, 233, 263,
273, 305, 335, 365, 391, 419, 447, 477,
497
Paracentral gyrus (paraCG), 21, 35, 43, 121,
143, 183, 209, 243, 267, 269, 315, 333,
373, 395, 429, 457, 483, 507
Paracentral sulcus (paraCS), 20, 34, 42, 120,
140, 184, 208, 242, 266, 268, 314, 334,
374, 426, 456, 484, 506
Parahippocampal gyrus (paHG), 21, 23, 35,
37, 43, 45, 107, 143, 171, 201, 229, 259,
273, 299, 335, 359, 363, 391, 415, 441,
471, 493
Parietal lobe (PL), 9, 87
Planum temporale (PT), 27, 38, 46
Pons (pons), 21, 23, 33, 35, 37, 41, 43, 45,
105, 145, 167, 203, 227, 265, 269, 297,
339, 357, 393, 413, 441, 467, 493
Postcentral gyrus (postCG), 19, 23, 33, 37,
41, 45, 115, 141, 177, 203, 235, 251,
271, 307, 335, 367, 393, 421, 449, 477,
499
Postcentral sulcus (postCS), 8, 18, 22, 32,
36, 40, 44, 50-75, 114, 142, 178, 204,
240, 250, 272, 308, 336, 368, 394, 422,
450, 476, 502, 518, 519, 530, 531
Posterior commisure (pc), 21, 35, 43,
501
Precentral gyrus (preCG, preCg), 19, 23, 33,
37, 41, 45, 111, 139, 179, 201, 233, 251,
271, 305, 333, 367, 391, 419, 449, 477,
499
Precentral sulcus (preCS), 8, 18, 22, 32, 36,
40, 44, 50-75, 112, 136, 178, 200, 232,
252, 270, 304, 332, 368, 390, 418, 448,
476, 498, 518, 519, 527, 530, 531
Precuneus (preCun), 21, 35, 43, 115, 149,
173, 215, 239, 267, 269, 273, 311, 343,
365, 405, 425, 453, 477, 507
Putamen (pu), 111, 137, 177, 199, 231, 261,
271, 303, 333, 365, 389, 417, 445, 475,
497
Quadrigeminal plate (qpl), 21, 35, 43
Retrosplenial area (rSp), 21, 35, 43, 113,
149, 175, 211, 237, 263, 269, 307, 343,
365, 399, 423, 449, 475, 501
Pxhinal sulcus (RhS, rhS), 20, 34, 42, 106,
172, 300, 360, 412, 440, 470, 492
Spinal cord (sc), 19, 21, 23, 33, 35, 37, 41,
43, 45, 151, 205, 225, 269, 345, 355,
395, 491
Subparietal sulcus (sPS), 20, 34, 42, 116,
148, 174, 214, 238, 264, 268, 310, 340,
366, 400, 454, 476, 504, 519, 522,
523
Superior frontal gyrus (SFG), 19, 21, 23, 33,
35, 37, 41, 43, 45, 111, 129, 183, 193,
231, 259, 269, 305, 323, 373, 381, 415,
445, 481, 495
Superior frontal sulcus (SFS), 18, 22, 32, 36,
40, 44, 50-75, 112, 128, 182, 192, 230,
256, 274, 302, 324, 370, 380, 414, 444,
478, 492, 518, 527, 530, 531
Superior parietal lobule (SPL), 19, 23, 33,
37, 41, 45, 119, 147, 173, 215, 241, 259,
271, 313, 343, 365, 403, 429, 453, 477,
509
Superior temporal gyrus (STG), 19, 23, 33,
37, 41, 45, 109, 139, 175, 199, 229, 251,
279, 301, 333, 363, 389, 415, 443, 475,
493
Superior temporal sulcus (STS), 18, 22, 32,
36, 40, 44, 50-75, 106, 138, 172, 200,
228, 250, 278, 298, 332, 362, 386, 412,
438, 470, 490, 526, 527, 530, 531
Index
[539]
Human Brain Anatomy
IN COMPUTERI ZED I MAGES
Superior temporal sulcus (continued)
inferior ramus (STSir), 114, 150, 176,
214, 238, 250, 282, 310, 364, 404, 426,
474, 502
superior ramus (STSsr), 8, 116, 150, 176,
214, 238, 250, 280, 310, 346, 366, 402,
426, 452, 474, 502
Sylvian fissure (SF), 8, 18, 22, 32, 36, 40, 44,
50-75, 87, 108, 138, 176, 198, 230, 250,
278, 302, 330, 364, 386, 416, 444, 476,
494, 518, 519, 526, 527, 530, 531
ascending branch of sylvian fissure (abSF),
8, 18, 22, 32, 40, 50-75, 110, 136, 176,
198, 232, 252, 278, 304, 330, 366, 388,
416, 446, 476, 496, 527, 530, 531
horizontal branch of sylvian fissure
(hbSF), 8, 18, 32, 40, 50-75, 110, 134,
176, 196, 254, 280, 302, 326, 366, 414,
444, 476, 494, 530, 531
Temporal lobe (TL), 9, 87
Temporal pole (TP), 19, 21, 23, 33, 35, 37,
41, 43, 45, 103, 137, 169, 197, 225, 257,
273, 297, 331, 361, 385, 411, 439, 469,
491
Temporo-occipital gyrus (TOG), 21, 23, 35,
37, 43, 45, 105, 141, 167, 201, 227, 257,
277, 297, 335, 359, 389, 411, 439, 469,
491
Thalamus (th), 21, 35, 43, 113, 143, 175,
203, 235, 263, 269, 305, 337, 365, 393,
419, 447, 475, 499
Transverse occipital sulcus (trOS, trsOS),
18, 22, 32, 36, 40, 44, 114, 156, 170,
218, 238, 256, 272, 308, 350, 362, 420,
452, 474, 504, 526, 527
Transverse temporal sulcus (trTS), 27, 38,
46, 112, 142, 176, 208, 234, 254, 278,
306, 336, 364, 392, 418, 446, 474, 502,
527
Uncus (uncus), 21, 35, 43
[540]

Ain Anatomy

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Human Brain Anatomy

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