Clinical

Neuroradiology Review Article

One Century of Brain Mapping Using Brodmann

Areas*
Michael Strotzer1

Abstract
100 years after their publication, Brodmann’s maps of the cerebral cortex are universally used to locate neuropsychological
functions. On the occasion of this jubilee the life and work of Korbinian Brodmann are reported. The core functions of each
single Brodmann area are described and Brodmann’s views on neuropsychological processes are depicted.

Key Words: Cerebral cortex · Neuropsychology

Clin Neuroradiol 2009;19:179–86

DOI 10.1007/s00062-009-9002-3

100 Jahre Hirnkartierung nach Brodmann

Zusammenfassung
100 Jahre nach ihrer Veröffentlichung wird Brodmanns Kartierung des zerebralen Kortex universell zur Lokalisation neu-
ropsychologischer Funktionen eingesetzt. Anlässlich dieses Jubiläums werden Leben und Werk von Korbinian Brodmann
dargestellt. Die wesentlichen Funktionen der einzelnen Brodmann-Areale werden beschrieben und Brodmanns Ansichten
über neuropsychologische Prozesse wiedergegeben.

Schlüsselwörter: Zerebraler Kortex · Neuropsychologie

Introduction er. He studied medicine in Munich, Würzburg, Berlin,

Exactly 100 years ago, Korbinian Brodmann’s famous
book was published [1]: “Vergleichende Lokalisations-
lehre der Großhirnrinde in ihren Prinzipien dargestellt
auf Grund des Zellenbaues” (The principles of compar-
ative localisation in the cerebral cortex based on cytoar-
chitectonics). Until today, Brodmann’s maps are uni-
versally used to locate neuropsychological functions in
the cerebral cortex, although Brodmann’s classificatory
system has been debated, refined and renamed for a
whole century.
Biography
Korbinian Brodmann (Figure 1) was born on Novem-
ber 17, 1868 in Liggersdorf, Germany, as son of a farm-
Freiburg, and Lausanne and intended to establish him-
self as a practitioner in the Black Forest. However, after
suffering from diphtheria in 1896 (1 year after receiving
his medical diploma), he began to work for Oskar Vogt,
director of the Neurologic Clinic Alexandersbad, and
decided to concentrate on neurology, psychiatry, and
brain research. In 1898, he received the degree of a doc-
tor of medicine from the University of Leipzig for his
thesis on chronic ependymal sclerosis. From 1901 to
1910, he performed his famous studies on comparative
cytoarchitectonics of the mammalian and human cortex
in the Neurobiological Laboratory, which was founded
by Oskar Vogt in Berlin in 1898. The basis of Brod-
mann’s localization is the subdivision of the cortex into

* Dedicated to my teacher Dr. Till Bretschneider (staff neuroradiologist;

Institute of Radiodiagnostics, University Hospital Regensburg, Germany).

1
Klinik Hohe Warte, Bayreuth, Germany.

Received: January 28, 2009; revision accepted: May 5, 2009

Published Online: July 20, 2009

Clin Neuroradiol 2009 · No. 3 © Urban & Vogel 179

Strotzer M. Brodmann Areas
Figure 1. This photo-
graph of Korbinian Brod-
mann served as frontis-
piece in the original
1909 edition of his book
[1] (from [2]).
areas with similar cellular and laminar structure using
the new staining method of Nissl. He studied rodents,
marsupials, primates, and human brains. Together with
Cécile and Oskar Vogt he described the different cyto-
architectonic structures of the pre- and postcentral gyri
before publishing his book, whose English translation
by Laurence J. Garey [2] is subject of this article. Sur-
prisingly, Brodmann’s professoral thesis on cortical ar-
chitecture was rejected by the Medical Faculty of Ber-
lin. However, he was welcomed to the Faculty of
Medicine in Tübingen, where he was appointed profes-
sor in 1913.
During his remarkable, but short career he worked
with Alois Alzheimer, Ludwig Binswanger, Hubert von
Grashey, Emil Kraepelin, and Franz Nissl. Brodmann
was a contemporary of many other famous neuroscien-
tists, e.g., Vladimir Betz, Pierre-Paul Broca, Santiago
Ramón Cajal, Alfred Campbell, Jean-Martin Charcot,
Harvey Cushing, Joseph Jules Déjerine, Guillaume
Duchenne, Siegmund Exner, Sigmund Freud, Norman
Geschwind, Camillo Golgi, Victor Horsley, John Hugh-
lings Jackson, Karl Kleist, Pierre Marie, and Carl Wer-
nicke.
In 1916, Brodmann took over the prosectorship at
the Nietleben Mental Asylum in Halle. He married
Margarete Francke in 1917 and their daughter Ilse was
born in 1918. After short illness he died in Munich on
180
August 22, 1918 at the age of 49 years, just after he had
begun to work as director of the group of histology at
the Psychiatric Research Center at the University of
Munich. Margarete Brodmann died on December 8,
1918, less than 4 months after her husband.
Brodmann’s biography and a lot of information ma-
terial concerning his life and work as well as the Kor-
binian Brodmann Museum in Hohenfels, Germany, is
available on the Korbinian Brodmann homepage (http://
www.korbinian-brodmann.de).
Brodmann’s Research in the Timeline
of Neuroanatomy
1858: R. Berlin analyzes the cytoarchitectonic structure
of the cerebral cortex.
1868: Theodor Meynert’s work about cytoarchitectonic
differences within the cerebral cortex is published.
1874: Vladimir Betz detects the giant pyramidal neu-
rons of the primary motor cortex (great cells of Betz).
1889: Victor Horsley develops a somatotopic map of the
monkey’s primary motor cortex using a stereotaxic
frame and needle electrodes.
1905: Alfred Campbell’s “Histological studies on the lo-
calisation of cerebral function” is published. He defines
17 cortical areas labeled by function [3].
1909: Korbinian Brodmann defines 52 discrete areas of
the cerebral cortex based on cytoarchitectonic features
[1].
1919: Cécile and Oskar Vogt continue Brodmann’s
work by describing more than 200 different cortical re-
gions.
1925: Constantin Economo and Georg Koskinas pre-
sent their cytoarchitectonic parcellation scheme of the
adult human brain with 107 cortical areas [4].
1925: cytoarchitectonic mapping of the cerebral isocor-
tex of the macaque monkey by Gerhardt von Bonin and
Percival Bailey.
1946: cytoarchitectonic studies of the cerebral cortex by
Karl S. Lashley and G. Clark.
1955: brain mapping and subdivision of Brodmann ar-
eas by S.A. Sarkissov.
Brodmann’s Localization in the Cerebral Cortex
The laminar cellular pattern of the cerebral cortex was
well recognized at that time and various layering
schemes with different nomenclatures and numberings
(varying from five to nine layers) were applied. Thus,
completely different layers carried identical names
while anatomically similar and homologous layers were
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Strotzer M. Brodmann Areas
Figure 2. The cortical areas of the lateral and medial surfaces of the human cerebral hemispheres (printed in the original edition of Brodmann’s
book [1]; from [8]).
given different names by different authors [5]. Accord-
ing to Brodmann the original pattern of the cortex is the
“primitive six-layered type” and all variations in cortical
structure (with the exception of some rudimentary cor-
tical zones) are derived from this cytoarchitecture [5].
Thus, Brodmann distinguished between two basic corti-
cal cytoarchitectonic patterns [5]:
(1) homogenetic formations (derived from the basic
six-layered type, covering the major part of the
hemispheres; today addressed as “isocortex”);
(2) heterogenetic formations (six-layered structure can-
not be demonstrated, e.g., in the rhinencephalon,
piriform lobe, cingulate gyrus adjacent to the corpus
callosum; now called “allocortex”).
Brodmann determined a cortical thickness in human
brains ranging from 2.3 mm to 4.5 mm, the giant pyrami-
dal cortex being always much thicker than the calcarine
cortex [6], and assumed a total cortical surface area of
1,900–2,200 cm2 [7]. He used the following scheme of
the basic six-layered cortex incorporating the nomen-
clature of various authors [5]:
I. molecular layer (lamina zonalis),
II. outer granular layer (lamina granularis externa),
III. pyramidal layer (lamina pyramidalis),
IV. inner granular layer (lamina granularis interna),
V. ganglion cell layer (lamina ganglionaris),
VI. spindle cell layer (lamina multiformis).
Brodmann analyzed regional variations in cell structure
of the cerebral cortex in detail. Studying different mam-
malian and human brains, he found a certain grade of
constancy regarding homologous cortical areas. Con-
cerning different cortical regions, however, a high grade
Clin Neuroradiol 2009 · No. 3 © Urban & Vogel
of variability was present. Based on exclusively anatom-
ic features, e.g., cell density, cell size, cortical thickness,
and composition of different layers, Brodmann per-
formed a parcellation of the cerebral cortex resulting in
a histological topographic map (Figure 2). He began his
studies in the rolandic region starting with the postcen-
tral gyrus, which could be separated into three different
areas by reproducible but certainly not absolutely sharp
borders (BA 1–3). Its main cytoarchitectonic features
are “the presence of a distinct granular layer and a lack
of giant pyramids” [8]. The precentral region (BA 4) “is
one of the most strikingly differentiated and cytoarchi-
tectonically delimitable structural regions of the whole
human cerebral cortex” [8]. It is “characterised by the
appearance of giant pyramids and the lack of an inner
granular layer” [8].
In this way, Brodmann defined 52 different areas
of the cerebral cortex. Their borders sometimes were
sharp (e.g., the boundaries of BA 44 and the anterior
border of BA 43) [8]. In spite of markedly different
cytoarchitectural features, Brodmann often found
ill-defined boundaries between the areas, e.g., the bor-
ders of BA 17–20, 23, 31, 37–39, 40, 46, and the poste-
rior border of BA 43 [8]. Not all areas show a homoge-
neous pattern of cytoarchitecture: BA 6 (agranular
frontal area) and BA 24 (ventral anterior cingulate
area) gradually change their structure in a dorsoven-
tral direction [8]. It is important to recognize, that the
borders between the areas often do not correspond ex-
actly to the sulci. The insular region of the human brain
was not parcellated by Brodmann. He found an ante-
rior agranular and a posterior granular area, separated
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Strotzer M. Brodmann Areas
a b
Figures 3a and 3b. Color-coded illustration of Brodmann’s cortical maps. Lateral (a) and medial surface (b).
by a line that is a prolongation of the central sulcus [8].
Nevertheless, he found it difficult to divide the insula
into separate areas and concluded as follows: “The
precise localisation of these individual fields must
await further investigation” [8].
Brodmann studied variations in cortical architec-
tonics among several species extensively. He also ana-
lyzed the ontogenetic development of the cortical lami-
nar structure in mammals and man. However, these
aspects are beyond the scope of this paper. Interesting-
ly, it is not clear whether Brodmann based his famous
maps (Figure 2) on the data of one single, adult, human
brain. This fact is one important source of criticism.
Brodmann Areas: Core Functions
A huge amount of functional imaging data is available
on the website of Trinity College, Hartford, CT, USA. It
reports on a meta-analysis of 478 published functional
magnetic resonance imaging (fMRI) studies to address
the issue of functional localization [9]. Figure 3 shows a
more recent illustration of Brodmann’s cortical maps
based on his original publication.
Areas 1, 2, and 3 (Intermediate, Caudal, and Rostral
Postcentral Area)
These areas represent the primary somatosensory pro-
jection cortex, which is responsible for somatosensory
perception. Its neurons are organized in a somatotopic
manner in the postcentral gyrus.
Area 4 (Giant Pyramidal Area)
The primary motor cortex is localized in the precentral
gyrus and is the main source of motor activation.
182
Area 5 (Preparietal Area)
This part of the superior parietal lobule and precuneus
is addressed as secondary somatosensory cortex and in-
volved in visuospatial processing.
Area 6 (Agranular Frontal Area)
The medial part corresponds to the supplementary mo-
tor area (SMA), the lateral one to the premotor area. Its
main functions are motor learning and planning as well
as motor activation of the hand (Exner’s area). It is part
of the dorsolateral frontal cortex (DLFC), which is in-
volved in working memory [10, 11].
Area 7 (Superior Parietal Area)
Anatomically, it includes the superior parietal lobule
laterally and the precuneus medially. It is involved in
counting and mathematics (dominant hemisphere), vi-
suospatial processing (especially in the nondominant
hemisphere), and episodic memory retrieval [12].
Area 8 (Intermediate Frontal Area)
Medially, it represents the pre-SMA (motor learning),
laterally, it includes the premotor cortex (motor action
planning) and the frontal eye field (voluntary eye move-
ment). It is part of the DLFC.
Area 9 (Granular Frontal Area)
As part of the DLFC it plays a role in working mem-
ory and higher cognitive processes (e.g., problem
solving). Together with BA 10, BA 11 and BA 46 it
builds the prefrontal cortex which is associated with
personality and is involved in memory and cognitive
tasks.
Clin Neuroradiol 2009 · No. 3 © Urban & Vogel
Strotzer M. Brodmann Areas
Area 10 (Frontopolar Area)
Together with BA 9 and BA 11 (prefrontal cortex) it is
responsible for the central executive.
Area 11 (Prefrontal Area)
It is part of the prefrontal cortex (see area 10) and in-
cludes the gyrus rectus and orbital gyri. Olfaction and
social behavior are important functional aspects.
Area 12 (Prefrontal Area)
This field was not included in Brodmann’s original maps
of the human brain. Later, it was defined as the superior
and medial part of BA 11.
Areas 13–16
Theses areas were not defined for the human brain by
Brodmann.
Area 17 (Striate Area)
The striate cortex (primary visual cortex) includes the
calcarine fissure region and the occipital pole. It has a
retinotopic organization and is responsible for visual
perception.
Area 18 (Occipital Area)
Functionally, it belongs to the secondary visual cortex
(color and motion recognition).
Area 19 (Preoccipital Area)
It is an element of the secondary visual cortex and like
BA 18 it includes a part of the fusiform (medial occipi-
totemporal) gyrus. Functions are object and face recog-
nition (fusiform face area [FFA]).
Area 20 (Inferior Temporal Area)
This field lies in the inferior temporal gyrus and is main-
ly involved in visual association processes (e.g., emo-
tional meaning of colors and facial expression).
Area 21 (Middle Temporal Area)
The middle temporal gyrus, a part of the auditory asso-
ciation cortex, is involved in higher-order audition pro-
cesses and speech reception (dominant hemisphere).
The posterior division belongs to the visual association
cortex and includes the FFA.
Area 22 (Superior Temporal Area)
The superior temporal gyrus houses the auditory asso-
ciation cortex and – in the dominant hemisphere togeth-
Clin Neuroradiol 2009 · No. 3 © Urban & Vogel
er with BA 21 and BA 42 – Wernicke’s area (speech
reception). Semantic memory is a function of the poste-
rior division of the superior temporal area.
Area 23 (Ventral Posterior Cingulate Area)
This part of the limbic cortex is associated with face fa-
miliarity and emotional processes.
Area 24 (Ventral Anterior Cingulate Area)
As another section of the limbic lobe it is involved in the
emotional assessment of somatosensory perception
(e.g., pain).
Area 25 (Subgenual Area)
Olfaction and working memory processing are two of
various possible functions of this area, which includes
the lamina terminalis and anterior perforate substance.
Area 26 (Ectosplenial Area)
The function of this small area near the posterior com-
missure is not well known.
Area 27 (Presubicular Area)
Memory encoding is the most important function of the
piriform gyrus.
Area 28 (Entorhinal Area)
Like BA 27 it is not based on the six-layered isocortex
but on the so-called allocortex (less than six – typically
three – layers of neurons). The entorhinal cortex forms
the main input to the hippocampus (important for mem-
ory encoding and consolidation) and is one of the first
areas to be affected in Alzheimer’s disease.
Area 29 (Granular Retrolimbic Area)
Only little is known about its function.
Area 30 (Agranular Retrolimbic Area)
Visual attention appears to be associated with BA 30.
Area 31 (Dorsal Posterior Cingulate Area)
This cortical field includes parts of the precuneus and
the posterior cingulate gyrus. It is thought to play a role
in visuospatial imagery, episodic memory retrieval, and
recognition of familiar faces.
Area 32 (Dorsal Anterior Cingulate Area)
Together with BA 24 it is responsible for the emotional
judgment of somatosensory awareness, specifically pain
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Strotzer M. Brodmann Areas
perception. It is also involved in motor action planning
and memory processes. Its dorsal and caudal partition is
equivalent to the rostral cingulate motor cortex (rCM).
Area 33 (Pregenual Area)
Little is known about its function. BA 33 is activated
during sexual arousal and plays a role in interoception.
Area 34 (Dorsal Entorhinal Area)
It belongs to the entorhinal cortex (see also BA 28) and
is important for memory, emotional assessment, and ol-
faction.
Area 35 (Perirhinal Area)
This area overlaps with the parahippocampal gyrus and
belongs to the limbic cortex. Involvement in memory
and emotional functions was observed (happiness, sad-
ness, disgust).
Area 36 (Ectorhinal Area)
The functions of this area are not quite clear. Involve-
ment in emotion perception has been suggested.
Area 37 (Occipitotemporal Area)
Visual and language functions are located in this area
(word and object recognition and naming, face recogni-
tion). It overlaps with the FFA.
Area 38 (Temporopolar Area)
Various putative roles are assigned to the temporal pole:
social and emotional processing, self-other distinction,
memory retrieval, and even humor comprehension
[13].
Area 39 (Angular Area)
Major functions are visual processing (face recognition,
reading), spatial imagery, visuomotor function (eye
movement), orthographic memory, and mathematics
(dominant hemisphere).
Area 40 (Supramarginal Area)
Spatial discrimination and – in the dominant hemisphere
– language (graphem-to-letter conversion) and numeri-
cal functions are represented in the supramarginal gyrus.
Area 41 (Anterior Transverse Temporal Area)
The primary auditory cortex is responsible for sound
perception. Its neurons are organized in a tonotopic and
periodotopic manner.
184
Area 42 (Posterior Transverse Temporal Area)
BA 42 plays a role as secondary auditory cortex and is
– in the dominant hemisphere – an element of Wer-
nicke’s area.
Area 43 (Subcentral Area)
The primary gustatory cortex is located in the frontal
and parietal opercular regions. BA 43 is also involved in
language perception.
Area 44 (Opercular Area)
In the dominant hemisphere, the opercular region of
the inferior frontal gyrus is part of Broca’s area (speech
expression). It is also involved in motor action of fingers
and toes. BA 44, 45 and 47 were termed as “subfrontal
area” by Brodmann.
Area 45 (Triangular Area)
The triangular region of the inferior frontal gyrus is also
part of Broca’s area. It is involved in tongue move-
ment.
Area 46 (Middle Frontal Area)
This division of the prefrontal cortex (see also BA 9) is
involved in motivation, attention, and several cognitive
tasks.
Area 47 (Orbital Area)
Memory, emotion, and olfaction are putative functions
of this element of the subfrontal cortex.
Areas 48–51
These areas were not included in Brodmann’s maps of
the human brain.
Area 52 (Parainsular Area)
It represents a narrow, band-like transitional area be-
tween the temporal cortex and the insula. There are no
data concerning the function of BA 52 in humans.
Brodmann’s Influence on Neuropathology and
Neuropsychology
The results of Brodmann’s treatise influenced the neu-
ropathologic view on secondary cellular changes in the
precentral region of subjects with amyotrophic lateral
sclerosis. Furthermore, a fundamental error concerning
the postcentral region in patients with tabes dorsalis
could be corrected: structural peculiarities, interpreted
as atrophy and erroneously attributed to tabes, revealed
Clin Neuroradiol 2009 · No. 3 © Urban & Vogel
Strotzer M. Brodmann Areas
a b
Figures 4a and 4b. Depiction of cortical activation (red) according to the Brodmann areas during neuropsychological tasks based on meta-analysis
data published by Lloyd [9]: motor action activation (a) and auditory perception (b).
to represent structural variations, which could be found
in all healthy human brains [14].
Brodmann was well aware of the meaning of field
topography for neuropathology and neuropsychology:
“Our comparative anatomical studies have furnished
such fundamentally new data concerning the histologi-
cal structure of the elements of the cerebral cortex, and
especially about homologies of individual elements, that
I believe we may even succeed in providing new direc-
tions for many physiological concepts” [15]. “There is
an undisputed axiom: physiologically dissimilar ele-
ments have dissimilar structures” [15].
Brodmann distinguished “strict localisers” and “half
localisers” as main groups of researchers. “The most ex-
treme localisational hypothesis states that the cerebral
cortex is divided into a large number of sharply sepa-
rated zones, rather like a topographical map, that cor-
respond exactly to the different sensory organs and mo-
tor apparatuses of the body” [15]. “Our findings (…)
prove with unassailable certainty the existence of a
strictly circumscribed regional localisation of specific
functions” [15]. “One must further accept that specific
complex processes occur mainly in one locality and oth-
ers mainly in another” [15]. Brodmann’s intuition was
that a particular anatomic structure corresponds to a
particular function. However, Brodmann recognized
recovery from physiological deficits due to cortical plas-
ticity [15] and accepted the principle of functional re-
placement. Thus, he cannot be classified as a “strict lo-
caliser” to use his own idiom.
One century after Brodmann’s publication there is
still a lively debate between “localizationists” and “an-
ti-localizationists”. The two major anti-localizationist
views are “holism” and “equipotentialism”. Both terms
Clin Neuroradiol 2009 · No. 3 © Urban & Vogel
are often used synonymously, although holism can be
regarded as a kind of compromise position between lo-
calizationism and equipotentialism. The essential point
for the equipotentialist is that no one area is function-
ally distinct from any other. Brodmann wrote: “The ac-
ceptance that all parts of the cerebral cortex participate
uniformly in all its functions must forthwith be consid-
ered as obsolete (…)” [15]. This is a clear statement
against equipotentialism. Holists, however, are of the
view that to understand how the brain subserves a given
function it is necessary to understand the whole brain or
at least large portions of it acting as integrated, func-
tional units. They deny that any particular function can
be understood as the isolated contribution of just one or
even a few, selected components [16]. Brodmann did
not completely reject the idea of holism: “In reality
there is only one psychic centre: the brain as a whole
with all its organs activated for every complex psychic
event, either all together or most at the same time and
so widespread over the different parts of the cortical
surface that one can never justify any separate specially
differentiated psychic centres within this whole. Natu-
rally this does not mean that all the individual organs
make equal physiological contributions to higher psy-
chic processes” [15]. Numerous functional imaging stud-
ies seem to support this holistic attitude. No single neu-
ropsychological task is fulfilled by the activity of only
one anatomically distinct area. The opposite is true: de-
pending on the number of experiments performed to
investigate a specific neuropsychological function, an
increasing number of active areas is found to be involved
[9]. The examples of action execution (Figure 4a) and
auditory perception (Figure 4b) are representative of
most neuropsychological tasks.
185
Strotzer M. Brodmann Areas
In the discussion about the meaning of cytoarchitec-
tonic cartography for cognitive neuroscience we must
not forget the importance of white matter connections.
In the late 19th century, localizationism was comple-
mented by the associationist school. Associationists (or
connectionists) stress the importance of the patterns of
white matter connection and explain neuropsychologi-
cal deficits using the disconnection theory of Karl Wer-
nicke [17]. Campbell, a contemporary of Brodmann,
based his cortical field map not only on cytoarchitec-
tonic features, but also on fiber anatomy and function
[18]. The science of connectional anatomy (hodology) is
presently reexplored using magnetic resonance tractog-
raphy based on diffusion tensor imaging (DTI).
Brodmann finished his reflections as follows: “A
wide field of fruitful activity is opened up to physiology
here through newly acquired anatomical localisational
information. Anyone wishing to undertake physiologi-
cal localisational studies will thus have to base his re-
search on the results of histological localisation” [15].
Indeed, most functional neuroimaging studies refer
their findings to Brodmann’s maps, which provide a
steady basis for comparison across experiments [9], a
kind of “lingua franca” of cortical location [18]. In view
of the huge number of functional imaging publications
Brodmann is still present, even 100 years after his great
publication.
Conflict of Interest Statement
The author declares that there is no actual or potential conflict of in-
terest in relation to this article.
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Address for Correspondence
Prof. Dr. Michael Strotzer
Klinik Hohe Warte
95445 Bayreuth
Germany
Phone (+49/921) 400-4402, Fax -4409
e-mail: Michael.strotzer@klinikum-bayreuth.de
Clin Neuroradiol 2009 · No. 3 © Urban & Vogel
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