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In Search of the Definitive Brodmann's Map of Cortical Areas in Human

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 C O M ME N T A R Y
In Search of the Definitive Brodmann’s Map of
Cortical Areas in Human

Goran Simić, 1
* and Patrick R. Hof2
1
Department of Neuroscience, Croatian Institute for Brain Research, University of Zagreb Medical School, Zagreb 10000, Croatia
2
Fishberg Department of Neuroscience and Friedman Brain Institute, Icahn School of Medicine at Mount Sinai, New York, New
York 10029
The birth of the cytoarchitectonics (in Korbinian Brod-
mann’s own words cytoarchitectonics should be defined
as “the localization of the individual histological ele-
ments, their layering, and their parcellation in the adult
brain”) of the human cerebral cortex is credited to Mey-
nert, who in 186721868, by using one of the first syn-
thetic colors, Prussian (Berlin) blue (in painting, Parisian
blue, Fe4[Fe(CN)6]3, derived from hydrogen cyanide or
“blue acid”), noticed regional variations in the nerve
cells stratification in different parts of the gray matter
of the cerebral hemispheres (Meynert, 1868). During
the 20th century, many investigators, including Camp-
bell (1903), Brodmann (1909, 1910; for review see
Judas et al., 2012), Elliot Smith (1907), von Economo
and Koskinas (1925), Vogt and Vogt (1926), Bailey and
von Bonin (1951), Sarkisov et al. (1955), and others,
have produced cytoarchitectural maps of the human
cerebral cortex by dividing it into a number of areas or
fields. For this purpose, they were generally used the
Nissl staining method, which is based on “defatting”
brain tissue sections by passing through graded dilu-
tions of ethanol, rehydrating through decreasing ethanol
concentrations, and staining in a water solution of a
cationic dye (such as methylene blue, cresyl violet, thio-
nin, toluidine blue). The ethanol solutions act to differ-
entiate the stain, causing myelin and other components
to lose color and be “stripped off the cells”, whereas
negatively charged molecules within cell bodies retain
the coloration (Nissl, 1894). Although many maps have
been produced in attempts to make a definitive parti-
tion of the human cerebral cortex, Brodmann’s map
and especially his numerical system attached to
cytoarchitecture is by far the most widely recognized in
current use. The last (final) Brodmann’s map of the
human cerebral cortex is shown in Figure 1A2C, and
the list of cortical areas defined by Brodmann is pro-
vided in Table 1.
There is a common agreement that the popularity of
Brodmann’s numbering system is due to its simplicity
to recall, its relatively sharp delineations of the borders
between adjacent areas, its detail not so exhaustive to
C 2014 Wiley Periodicals, Inc.
V (wileyonlinelibrary.com)
The Journal of Comparative Neurology | Research in Systems Neuroscience 523:5–14 (2015)
be impossible to refer to, and the fact that Brodmann’s
observations of cytoarchitecture have stood the test of
time. In addition, Brodmann’s map is also the closest to
the modern definition of the cortical area (field), by
which an area, among several other features, is receiv-
ing afferent fibers from a particular (principal) nucleus
of the thalamus (Jones, 1985, 2009). Thalamic inputs,
except for the so-called nonspecific thalamic nuclei,
stop abruptly at an areal border, and there appears to
be no overlap between inputs from different specific
thalamic nuclei to adjacent fields (Jones, 1985, 2009).
Moreover, the chemoarchitectural borders, as defined
by a characteristic pattern of histochemical or immuno-
cytochemical staining, also commonly coincide with the
cytoarchitecture defined by Brodmann. It is therefore
not surprising that modified versions of Brodmann’s
map, often based on its extrapolation onto the Talairach
and Tournoux brain atlas (Talairach and Tournoux,
1988), are so frequently used in neuroimaging studies
to describe the localization of activation (Zilles and
Amunts, 2010). In practice, however, this led to the
occurrence of consistent mistakes and misinterpreta-
tions in relation to usage and quotation of Brodmann’s
maps and numbers.
Brodmann published a series of comparative neuroana-
tomical works on more than 60 mammalian species in
the Journal f €ur Psychologie und Neurologie between 1903
and 1908. These findings were summarized in his famous
book (Brodmann, 1909) and extended in two chapters,
which he contributed to contemporary handbooks
(Brodmann, 1910, 1914). The first (“original”) Brodmann’s
cytoarchitectonic map was first published in one of his
monographs (Brodmann, 1908a) and then reproduced
in his 1909 book, when he also added the map of the
Grant sponsor: Croatian Science Foundation; Grant number: 09/16
 Grant sponsor: Autism Speaks (to P.R.H.).
(to G.S.).
*CORRESPONDENCE TO: Goran Simić, MD, PhD, Department of Neuro-
science, Croatian Institute for Brain Research, University of Zagreb Medi-

cal School, Salata 12, Zagreb 10000, Croatia; E-mail: gsimic@hiim.hr
Received May 5, 2014; Revised May 28, 2014;
Accepted May 28, 2014.
DOI 10.1002/cne.23636
Published online May 30, 2014 in Wiley Online Library
5
  Ć1* and P.R. Hof
G. Simi

Figure 1. A,B: Brodmann’s regional map of the adult human cerebral cortex based on cytoarchitectural analysis, as reproduced by Lewan-
dowsky’s Handbook of Neurology (Brodmann, 1910). Brodmann intentionally projected borders between neighboring fields, which are found
in the depths of sulci such as in pericentral and retrosplenial regions (e.g., areas 2, 3, 5, 6, 18, 19, etc.), on the free surface of the hemi-
sphere. C: Brodmann’s map of the insula and the adjacent superior aspect of the superior temporal gyrus as it appears in his 1909 mono-
graph (Brodmann, 1909). I. ant., Anterior insular zone; I. post., posterior insular zone; sp, posterior ramus of the Sylvian fissure; sv,
vertical ramus of the Sylvian fissure; sh, horizontal ramus of the Sylvian fissure; t1, superior temporal sulcus. D: Cytoarchitectonic parcella-
tion of the orbitofrontal cortex, extrapolated from Brodmann (1914), as it appeared in Nieuwenhuys et al., 2008 (with permission); LOS,
lateral orbital sulcus; MOS, medial orbital sulcus; OLF, olfactory sulcus; TOS, transverse orbital sulcus. Images have been graphically
enhanced for clarity.

insular and adjacent cortices of the superior temporal
gyrus (shown in Fig. 1C) and grouped individual areas
into 11 larger (major) regions (“Hauptzonen”, Table 1,
second column; Brodmann, 1909). The second (and
final) Brodmann’s cytoarchitectonic map of cortical
regions was published in Lewandowsky’s Handbook of
Neurology in 1910 (Brodmann, 1910; Fig. 1A2C), in
which he added areas 12 (between areas 10 and 11 in
medial view) and 52 (between areas 43 and 41 in lat-
eral view) to the first (1908) map, and subdivided area
7 into two subfields (7a and 7b), apparently without
acknowledging these changes in the text (Markowitsch,
1993). To understand the changes made, one should
consult the text of his 1909 book, in which he had
already proposed these and many other changes to his
initial 1908 map (Judas et al., 2012). A misquotation
made in many current reproductions of Brodmann’s
map is that the 1910 map is reproduced, but incor-
rectly referred to, as representing the one published in
the 1909 book (e.g., Greenstein and Greenstein, 2000;
Zilles et al., 2002; Vogt et al., 2004; ten Donkelaar
et al., 2006; Jacobson and Marcus, 2008; Klein and
Behrens, 2009; Filley, 2011; Krebs et al., 2012; Amaral
and Strick, 2013; Kandel and Hudspeth, 2013; among
others). Consequently, the numbering system also does
not match the map.
The second most commonly encountered mistake is
that, even today, most textbooks cannot agree on the
total number of cortical areas described by Brodmann
(see Table 2). The reason for this is a misperception
that the “gaps” in numbers 13216 and 48251, which
are not shown on either of Brodmann’s maps of the
human cerebral cortex, occurred because Brodmann
described those areas in the primate brains but “could
not find homologous areas in the human brain” (e.g.,
Zilles and Amunts, 2010). Many experts thought that,
for example, “numbers 13216 are totally missing” and
that Brodmann had not described those areas in the

6 The Journal of Comparative Neurology | Research in Systems Neuroscience

 Brodmann’s map of cortical areas in human

TABLE 1.
Brodmann’s List of Human Cortical Areas Sorted According to Topographic Regions (First Column) According to Brodmann,
1908 (and Brodmann, 1905 and 1914, for Insula), or 11 Major Cytoarchitectonic Regions (Second Column) According to
Brodmann, 19091
Brodmann (1908), except for areas 13216 Brodmann (1909), except for areas 12
(Brodmann, 1905; Brodmann, 1914) and 52 (Brodmann, 1910)
Area No. Area name Area no. Area name Brodmann (1909) core feature
Regio rolandica Regio postcentralis
1 Area postcentralis intermedia 1 Area postcentralis intermedia Homogenetic: homotypical cortex
(six-layered pattern throughout life)
2 Area postcentralis caudalis 2 Area postcentralis caudalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
3 Area postcentralis oralis 3 Area postcentralis oralis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
43 Area subcentralis 43 Area subcentralis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
— Regio praecentralis
4 Area praecentralis 4 Area gigantopyramidalis Homogenetic: heterotypical cortex
gigantopyramidalis (secondary transformation to six-
layered pattern with reduction in
layers)
Lobus frontalis
6 Area frontalis agranularis 6 Area frontalis agranularis Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with reduction in
layers)
— Regio frontalis
8 Area frontalis intermedia 8 Area frontalis intermedia Homogenetic: homotypical cortex
(six-layered pattern throughout life)
9 Area frontalis granularis 9 Area frontalis granularis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
10 Area frontopolaris 10 Area frontopolaris Homogenetic: homotypical cortex
(six-layered pattern throughout life)
11 Area praefrontalis 11 Area praefrontalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
12 Area frontopolaris (5area Homogenetic: homotypical cortex
praefrontalis (six-layered pattern throughout life)
superomedialis)
44 Area opercularis 44 Area opercularis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
45 Area triangularis 45 Area triangularis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
46 Area frontalis media 46 Area frontalis media Homogenetic: homotypical cortex
(six-layered pattern throughout life)
47 Area orbitalis 47 Area orbitalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
Lobus parietalis Regio parietalis
5 Area praeparietalis 5 Area praeparietalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
7 Area parietalis superior 7 Area parietalis superior Homogenetic: homotypical cortex
(six-layered pattern throughout life)
39 Area parietalis inferior posterior 39 Area angularis Homogenetic: homotypical cortex
(5area angularis) (six-layered pattern throughout life)
40 Area parietalis inferior anterior 40 Area supramarginalis Homogenetic: homotypical cortex
(5area supramarginalis) (six-layered pattern throughout life)
Lobus occipitalis Regio occipitalis
17 Area striata 17 Area striata Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with increase in
layers)
18 Area occipitalis 18 Area occipitalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
19 Area praeoccipitalis 19 Area praeoccipitalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)

The Journal of Comparative Neurology | Research in Systems Neuroscience 7

  Ć1* and P.R. Hof
G. Simi

TABLE 1. Continued
Brodmann (1908), except for areas 13216 Brodmann (1909), except for areas 12
(Brodmann, 1905; Brodmann, 1914) and 52 (Brodmann, 1910)
Area No. Area name Area no. Area name Brodmann (1909) core feature
Lobus temporalis Regio temporalis
20 Area temporalis inferior 20 Area temporalis inferior Homogenetic: homotypical cortex
(six-layered pattern throughout life)
21 Area temporalis media 21 Area temporalis media Homogenetic: homotypical cortex
(six-layered pattern throughout life)
22 Area temporalis superior 22 Area temporalis superior Homogenetic: homotypical cortex
(six-layered pattern throughout life)
36 Area ectorhinalis 36 Area ectorhinalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
37 Area occipitotemporalis 37 Area occipitotemporalis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
38 Area temporopolaris 38 Area temporopolaris Homogenetic: homotypical cortex
(six-layered pattern throughout life)
41 Area (temporalis) transversa 41 Area temporalis transversa Homogenetic: homotypical cortex
interna s. anterior interna (anterior) (six-layered pattern throughout life)
42 Area (temporalis) transversa 42 Area temporalis transversa Homogenetic: homotypical cortex
externa s. posterior externa (posterior) (six-layered pattern throughout life)
— 52 Area parainsularis Homogenetic: homotypical cortex
(six-layered pattern throughout life)
50* Unnamed Homogenetic: homotypical cortex
(six-layered pattern throughout life)
Lobus limbicus Regio cingularis
— Subregio postcingularis:
23 Area limbica posterior ventralis 23 Area cingularis posterior Homogenetic: homotypical cortex
ventralis (six-layered pattern throughout life)
31 Area limbica posterior dorsalis 31 Area cingularis posterior Homogenetic: homotypical cortex
dorsalis (six-layered pattern throughout life)
— Subregio praecingularis:
24 Area limbica anterior ventralis 24 Area cingularis anterior Homogenetic: heterotypical cortex
ventralis (secondary transformation to six-
layered pattern with reduction in
layers)
32 Area limbica anterior dorsalis 32 Area cingularis anterior Homogenetic: heterotypical cortex
dorsalis (secondary transformation to six-
layered pattern with reduction in
layers)
33 Area praegenualis 33 Area praegenualis Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with reduction in
layers)
25 Area subgenualis (5area 25 Area subgenualis (5area Heterogenetic: rudimentary cortex
preterminalis) preterminalis) (isolated rudimentary layers)
— Regio retrosplenialis
26 Area ectosplenialis 26 Area ectosplenialis Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with reduction in
layers)
29 Area retrolimbica granularis 29 Area retrolimbica granularis Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with reduction in
layers)
30 Area retrolimbica agranularis 30 Area retrolimbica agranularis Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with reduction in
layers)
— Regio hippocampica
27 Area praesubicularis 27 Area praesubicularis Heterogenetic: striate cortex (several
secondary, further differentiated
layers)
28 Area entorhinalis ventralis 28 Area entorhinalis Heterogenetic: striate cortex (several
secondary, further differentiated
layers)

8 The Journal of Comparative Neurology | Research in Systems Neuroscience

 Brodmann’s map of cortical areas in human

TABLE 1. Continued
Brodmann (1908), except for areas 13216 Brodmann (1909), except for areas 12
(Brodmann, 1905; Brodmann, 1914) and 52 (Brodmann, 1910)
Area No. Area name Area no. Area name Brodmann (1909) core feature
34 Area enthronialis dorsalis 34 Area entorhinalis dorsalis Heterogenetic: striate cortex (several
secondary, further differentiated
layers)
35 Area perirhinalis 35 Area perirhinalis Heterogenetic: striate cortex (several
secondary, further differentiated
layers)
48* Area retrosubicularis Heterogenetic: striate cortex (several
(postsubicularis) secondary, further differentiated layers)
49* Area parasubicularis Heterogenetic: striate cortex (several
secondary, further differentiated
layers)
Regio insularis
— 52 Area parainsularis Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with increase in
layers)
13* Insula posterior — Insula posterior Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with increase in
layers)
14* Insula anterior — Insula anterior Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with increase in
layers)
15* Insula ventralis — Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with increase in
layers)
16* Insula olfactoria — Homogenetic: heterotypical cortex
(secondary transformation to six-
layered pattern with increase in
layers)
Regio olfactoria
Cortex primitivus
— Bulbus olfactorius Heterogenetic: primitive cortex
(without layers)
— Pedunculus olfactorius Heterogenetic: primitive cortex
(without layers)
— Tuberculum olfactorium Heterogenetic: primitive cortex
(without layers)
— Substantia perforata anterior Heterogenetic: primitive cortex
(without layers)
— Amygdala Heterogenetic: primitive cortex
(without layers)
Cortex rudimentarius
— Hippocampus Heterogenetic: rudimentary cortex
(isolated rudimentary layers)
— Gyrus dentatus Heterogenetic: rudimentary cortex
(isolated rudimentary layers)
— Subiculum Heterogenetic: rudimentary cortex
(isolated rudimentary layers)
— Indusium griseum Heterogenetic: rudimentary cortex
(isolated rudimentary layers)
— Septum pellucidum Heterogenetic: rudimentary cortex
(isolated rudimentary layers)
51* Area prepiriformis Heterogenetic: rudimentary cortex
(isolated laminar pattern)
1
The third column describes the core cytoarchitectural features of each area (homogenetic cortex means all types of cortex derived from the basic
six-layered pattern; heterogenetic cortex means all types of cortex that lack the six-layered pattern in ontogenesis and phylogenesis). A dash indi-
cates that Brodmann described but did not assign a number to a given structure. An asterisk indicates that Brodmann described but did not illus-
trate a given area in either of his maps.

The Journal of Comparative Neurology | Research in Systems Neuroscience 9

  Ć1* and P.R. Hof
G. Simi

Examples of Variable Numbers of Brodmann’s Areas Reported by Various Authors Either in Articles or in Textbooks, With
Reference
Gorman and Un€utzer, 1993
Toga and Mazziota, 2002
Purves et al., 2004
Van Essen, 2005
Nieuwenhuys et al., 2008
Strotzer, 2009
Olry, 2010
Zilles and Amunts, 2010
Olry and Haines, 2010
Geyer et al., 2011
Loukas et al., 2011
Seung, 2012
Kandel and Hudspeth, 2013
Amaral and Strick, 2013
TABLE 2.
the Authors’ Explanation
No. of areas Authors’ explanation
47 “Most neurologists have some familiarity with the system of Korbinian Brodmann,
which divides the cortex into 47 parts on the basis of cytoarchitecture.”
52 “Brodmann’s 1909 numerical map of the human brain, based on a
cytoarchitectonic study, recognized 52 discrete areas.”
About 50 “Korbinian Brodmann, who early in the twentieth century devoted his career to an
analysis of brain regions distinguished in this way, described about 50 distinct
cortical regions, or cytoarchitectonic areas.”
46 “Brodmann identified 28 neocortical areas in the Old World monkey and 46 in the
human.”
44 “The highest numbered area in Brodmann’s map is area 52; however, because
areas numbered 13–16 and 48–51 are lacking the total number of areas in his
map amounts to 44.”
44 “Areas 13216 were not defined for the human brain by Brodmann” and “Areas
48251 were not included in Brodmann’s maps of the human brain.”
44 “However, it should be pointed out that Brodmann’s areas are not 52 in number
as it may appear, but only 44 because areas 13216 and 48251 do not exist.”
43 “The insular cortex is segregated into areas 14216 in Old World monkeys (for
example, Cercopithecus) and into areas 13216 in prosimians (for example,
Lemuridae). Brodmann could not find homologous areas in the human brain.”
44 “However, Brodmann’s areas actually are not 52 in number, but only 44 because
areas 13216 and 48251 do not exist!”
43 “However, there are two gaps in the sequence: areas 12, 13, 14–16, and 48–51
were defined in lower mammals and non-human primates, but they are missing
in humans. This results in 43 areas in the human cortex.”
47 “In humans, he identified 47 histologically distinct regions using novel staining
techniques introduced by Nissl, and in primates, he described 52 different
regions.”
43 “Brodmann skipped 12216 and 48251. He reserved these numbers for cortical
areas in animals that appeared to have no analogues in the human cortex.”
52 “Brodmann distinguished 52 anatomically and functionally distinct areas in the
human cerebral cortex”; and map caption: “Brodmann’s division of the human
cerebral cortex into 52 functional areas.”
47 “Based on such cytoarchitectonic differences, Brodmann in 1909 divided the
cerebral cortex into 47 regions.” (reproduced from Martin, 2003)

human brain (e.g., Gorman and Un€utzer, 1993; Strotzer,
2009; among others).
The facts are quite the opposite. Fields 13216 would
correspond to the insular cortex, and Brodmann studied
this region in detail. Brodmann first published a subdivi-
sion of the human insula into three regions in 1903
(Brodmann, 1903); in his 1905 paper (Brodmann, 1905),
with regard to the four insular regions that he defined in
a monkey (Cercopithecus) brain, he specifically stated (p.
219) that “I do not hold it unlikely that, upon review of a
larger material and with consideration of comparative-
anatomical features, I will succeed to establish four
areas in humans too (Ich halte es nicht f€ur unwahrschein-
lich, dab es bei Durchsicht eines gr€oberen Materials und
mit Ber€ucksichtigung vergleichend-anatomischer Merkmale
auch beim Menschen gelingen wird, 4 Typen
aufzustellen).” Given that Brodmann was well aware of
the high variability in insular areas among different mam-
malian species, even when describing those areas in
great detail, he did not put numbers 13216 on the map
of the lemur’s brain (Brodmann, 1908b), because “he did
not want to destroy the clearness of the illustrations”
(Fix, 1993). As he did not pursue the cytoarchitectural
analysis of the human insula later (Fix, 1993), in his
1908 paper on the human brain (Brodmann, 1908a) he
simply stated (p. 244) that “insula, for which I discerned
4 areas, cannot be seen on the illustration” and restated
the same in a similar manner in his 1909 book (Brod-
mann, 1909, p. 146). Finally, in his legend to the 1914
chapter (Brodmann, 1914), he explicitly stated that
“human insular region contains areas 13216” (p. 101).
Unfortunately, his work on the insular cortex remained
unfinished, with his premature death in 1918. The insular
cortex is one of the most heterogeneous regions of the
cerebral cortex, both structurally and functionally, as
exemplified in pioneering comparative embryological and
cytoarchitectonic studies by Maximilian Rose (1928a,b)
as well as in several recent reports (Craig 2002, 2009;

10 The Journal of Comparative Neurology | Research in Systems Neuroscience


Butti et al., 2010; Gallay et al., 2012; Nieuwenhuys,
2012; Bauernfeind et al., 2013; Morel et al., 2013; Gu
et al., 2013; Barks et al., 2014; Evrard et al., 2014). In
fact, Rose (1928b) described as many as 31 cytoarchi-
tectural areas in the human insula. Moreover, it was
Rose who first suggested that the insula of great apes
and humans could be distinguished from that of most
monkeys and of small mammals by the abundant pres-
ence of a specific type of neurons (the so-called von
Economo neurons) in the ventral anterior insula (Rose,
1928b; see Seeley at al., 2012). Thus, Brodmann was far
ahead of his time when he envisioned that “the further
differentiation of the insular region into individual
cytoarchitectonic areas will be the object of future stud-
ies (Die genaue Lokalisation dieser Einzelfelder mub
sp€ateren Forschungen vorbehalten bleiben)” (Brodmann,
1909, p. 146; Garey, 2006, p. 176).
Some authors, perhaps being unaware that Brod-
mann’s numbers 13 and 14 were in fact already
accounted for by parts of the insular cortex, have
unfortunately transferred these presumed “unused”
numbers to remap the orbitofrontal cortex, first in mac-
aque monkeys and then in humans. For example,
Walker et al. (1940) designated the orbital and medial
parts of the orbitofrontal cortex as Brodmann’s areas
13 and 14, respectively. On the basis of such publica-
tions, some authors have recently subdivided orbital
and medial prefrontal cortex of the macaque monkeys
into areas 13m, 13l, 13a, 13b, and 14r and 14c (e.g.,
Carmichael and Price, 1994) and described their
“counterparts” in humans (e.g., Ong€€ ur et al., 2003),
bringing further confusion to the map for comparative
studies.
As with areas 13216, Brodmann also worked on
areas 48251 in the human brain but did not illustrate
them on either of his maps, presumably because of
their small size and to increase clarity. Although the
textual description of these areas is scarce, careful
reading of Brodmann’s 1909 book indicate that he ana-
lyzed these areas and assigned them numbers 48251
in the human cerebral cortex as well (Markowitsch,
1993).
Brodmann named area 48 the area retrosubicularis
(postsubicularis): “At the caudal end of the perirhinal
area (35) and lateral to the presubicular area (27) one
can also differentiate another special structural type in
man, which I first recognised as such after it had struck
me in lower animals, in which I described it as the ret-
rosubicular area (48) (Am kaudalen Ende der Area peri-
rhinalis (35) und lateral von der Area praesubicularis
(27) k€onnte man auch beim Menschen noch eine beson-
dere Strukturformation abtrennen, welche ist erst als
solche erkannt habe, nachdem sie mir bei niederen Tie-
The Journal of Comparative Neurology | Research in Systems Neuroscience
Brodmann’s map of cortical areas in human
ren aufgefallen war. Ich habe sie dort als Area retrosubi-
cularis (48) beschrieben)” (Brodmann, 1909, p. 150;
Garey, 2006, p. 126).
Area 49 was described by Brodmann as the area par-
asubicularis between the presubicular and entorhinal
areas (27 and 28, respectively), also as a part of the
hippocampal region at several places in the 1909 book.
For example, “the hippocampal region is, with the pre-
central and insular regions, the most absolutely con-
stant of the major, homogeneous structural regions in
mammals. It belongs to the heterogenetic formations
and is immediately recognizable in all mammalian
brains by its atypical lamination. It includes the cortex
medial to the posterior rhinal sulcus as far as the hip-
pocampal sulcus, that is to say, the head of the para-
hippocampal gyrus or part of the piriform lobule or its
homologues, and also the deep cortex of the hippocam-
pal sulcus itself; it consists of areas 27, 28, 35, 48 and
49 in our maps, as well as their subdivisions (Die hippo-
kampale Hauptregion ist neben der Regio praecentralis
und insularis das absolut konstanteste einheitliche
gr€obere Strukturgebiet in der S€augerreiche. Es geh€ort zu
den heterogenetischen Formationen und ist zufolge
seines atypischen Schichtungscharakters ohne weiteres
an allen Mammaliergehirnen identifizierbar. Es umfabt
die Rinde nach innen vom Sulcus rhinalis posterior bis
zum Sulcus hippocampi, also das Caput Gyri hippocampi
resp. Einen Teil des Lobulus pyriformis oder deren
Homologa, ferner die Tiefenrinde des Sulcus hippocampi
selbst; sie setzt sich in unseren Diagrammen aus den
Feldern 27, 28, 35, 48 und 49, sowie deren Unterabtei-
lungen zusammen)” (Brodmann, 1909, p. 150; Garey,
2006, p. 177).
Area 50 is probably the least described area by Brod-
mann (Markowitsch, 1993), presumably because “in the
temporal region one encounters the greatest difficulties
with structural homologies. In the accompanying map
(Figure 104) I include areas 20, 21, 22, 50 and 36 in
this region (Bei der Regio temporalis begegnet man mit
der strukturellen Homologie den gr€obten Schwierig-
keiten. Ich rechne auf der beigegebenen Hirnkarte (Figur
104) zu dieser Hauptzone die Felder 20, 21, 22, 50, und
36).” Then, after another sentence, “it is not at all pos-
sible to decide which of the three areas 21, 22 and 50
correspond to the middle and superior temporal gyri,
and thus the middle and superior temporal areas, of pri-
mates (vollends gar nicht zu entscheiden ist es, welches
von den drei Feldern 21, 22 und 50 der mittleren und
oberen Schl€afenwindung, also der Area temporalis supe-
rior und media bei den Primaten entspricht).”
(Brodmann, 1909, p. 1862187; Garey, 2006, p. 161).
Another reason for the scarce description of area 50,
which remained unnamed in the human brain, could be
11
  Ć1* and P.R. Hof
G. Simi
the fact that Brodmann was undecided of its actual
affinities: “whether area 50, lying at the upper border
of area 13, should also be counted in the insular region
I cannot decide for the present (Ob das am oberen
Rande von 13 gelegene Feld 50 auch zur insul€aren
Hauptregion zu rechnen ist, vermag ich einstweilen nicht
zu entscheiden)” (Brodmann, 1909, p. 150; Garey,
2006, p. 1642165).
Finally, Brodmann defined area 51 as the area prepiri-
formis and descibed it as a part of the olfactory region:
“in most mammals the region consists, apart from the
amygdala and the olfactory tubercle, only of a single
area (51) that is not further differentiated (into subfields
a, b, c, and d) (bei den meisten Mammaliern besteht die
Regio auber dem Mandelkern und dem Tuberculum olfac-
torium €uberhaupt nur aus einem einzigen nicht weiter dif-
ferenzierten Felde (Area 51a2Area 51d))” (Brodmann,
1909, p. 228; Garey, 2006, p. 1962197).
The early motivation for cytoarchitectural studies
came first from phrenology, then from demonstrations
of the electrical excitability of the motor cortex and
from the recognition of the dependence of vision on
posterior regions of the cerebral cortex. Later, areas
with somatic sensory, auditory, association, and other
distinct functions were, in turn, described. The identifi-
cation of cortical fields and their borders thus became,
and remains to this day, a search for areas of the cere-
bral cortex with distinct functions (Jones, 2009). For
Brodmann, the cortical areas were the ‘‘organs of the
brain”, each with a set of specific functions. Unlike
Flechsig (1896, p. 97), who distinguished four “psychic
centres” or “thought organs” (one frontal, one insular,
one parietal, and one temporal, as based on the asyn-
chronous myelinization of the different parts of the cor-
tex), Brodmann believed that “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 sur-
face that one can never justify any separate specially
differentiated ‘psychic’ centres within this whole (In
Wahrheit gibt es eben nur ein psychisches Zentrum: das
ist das Gehirn als Ganzes mit allen seinen Organen, die
bei jedem verwickelteren psychischen Vorgang entweder
s€amtliche oder in der Mehrzahl zugleich und in so weiter
Ausdehnung €uber die verschiedensten Teile der Rin-
denfl€ache in Aktion treten, dab von irgendeinem inner-
halb dieses Ganzen abzugrenzenden ‘psychischen’
Sonderzentrum niemals die Rede sein kann)” (Brodmann,
1909, p. 303; Garey, 2006, p. 251).
The trend now appears to be going in the direction
of a finer and finer parcellation, akin to that of the
Vogts, who described over 200 areas (Vogt and Vogt,
12 The Journal of Comparative Neurology | Research in Systems Neuroscience
1926), as functional subdivisions are recognized within
areas by new techniques such as evoked potentials and
microelectrode mapping, receptor autoradiographic
mapping (e.g., Palomero-Gallagher et al., 2013), and
functional magnetic resonance imaging and diffusion
tensor imaging (e.g., http://www.humanconnectome-
project.org/; Seung, 2012; Van Essen et al., 2013). As
an example, Palomero-Gallagher and collaborators
(2013) demonstrated a rich diversity of receptor binding
patterns (such as for kainate, benzodiazepine, sero-
tonin, N-methyl-D-aspartate, and a2-adrenergic recep-
tors) in Brodmann’s area 32, which plays a crucial role
in emotional processing and memory consolidation, that
provided a pharmacologically specific context to map-
ping efforts based strictly on cortical architecture. It
can be concluded that such modern imaging studies
are uncovering new functional localizations (areal acti-
vation “centroids” in terms of peak voxel coordinates)
with hopes that, eventually, molecular-genetic markers
will be discovered ultimately to determine a reproduci-
ble parcellation of the cerebral cortex on the basis of
both structural and functional properties. It will then be
safe to say that the definitive Brodmann’s map of the
human cerebral cortex is established.
CONFLICT OF INTERESTS STATEMENT
The authors have no conflict of interest to disclose.
ROLE OF AUTHORS
Both authors equally contributed to this work.
LITERATURE CITED
Amaral GA, Strick PL. 2013. The organization of the central
nervous system. In: Kandel ER, Schwartz JH, Jessell TM,
Siegelbaum SA, Hudspeth AJ, editors. Principles of neural
science, 5th ed. New York: McGraw-Hill. p 3372355.
Bailey P, von Bonin G. 1951. The isocortex of man. Urbana,
IL: University of Illinois Press.
Barks SK, Bauernfeind AL, Bonar CJ, Cranfield MR, de Sousa
AA, Erwin JM, Hopkins WD, Lewandowski AH, Mudakikwa
A, Phillips KA, Raghanti MA, Stimpson CD, Hof PR, Zilles
K, Sherwood CC. 2014. Variable temporoinsular cortex
neuroanatomy in primates suggests a bottleneck effect
in eastern gorillas. J Comp Neurol 522:8442860.
Bauernfeind AL, de Sousa AA, Avasthi T, Dobson SD, Raghanti
MA, Lewandowski AH, Zilles K, Semendeferi K, Allman
JM, Craig AD, Hof PR, Sherwood CC. 2013. A volumetric
comparison of the insular cortex and its subregions in
primates. J Hum Evol 64:2632279.
Brodmann K. 1903. Zur histologischen Lokalisation der Hirn-
rinde (die Inseltypen). Neurol Zentralbl 32:113321134.
Brodmann K. 1905. Beitr€age zur histologischen Lokalisation
der Grobhirnrinde. Dritte Mitteilung: Die Rindenfelder der
niederen Affen. J Psychol Neurol 4:1772226.
Brodmann K. 1908a. Beitr€age zur histologischen Lokalisation
der Grobhirnrinde. VI Mitteilung: Die cytoarchitektonische
Cortexgliederung des Menschen. J Psychol Neurol 10:
2312246.

Brodmann K. 1908b. Beitr€age zur histologischen Lokalisation
der Grobhirnrinde. VII Mitteilung: Die Cytoarchitektoni-
sche Cortexgliederung der Halbaffen (Lemuriden). J Psy-
chol Neurol 10:2872334.
Brodmann K. 1909. Vergleichende Lokalisationslehre der
Grobhirnrinde in ihren Prinzipien dargestellt auf Grund
des Zellenbaues. Leipzig: J.A. Barth.
Brodmann K. 1910. Feinere Anatomie des Grobhirns. In: Lew-
andowsky M, editor. Handbuch der Neurologie, 1. Band:
Allgemeine Neurologie. Berlin: Springer. p 2062307.
Brodmann K. 1914. Physiologie des Gehirns. In: Von Bruns P,
editor. Neue Deutsche Chirurgie, vol. 11. Stuttgart: Enke.
p 852426.
Butti C, Hof PR. 2010. The insular cortex: a comparative per-
spective. Brain Struct Funct 214:4772493.
Campbell AW. 1903. Histological studies on cerebral localiza-
tion. Proc R Soc Lond B Biol Sci 72:4882492.
Carmichael ST, Price JL. 1994. Architectonic subdivisions of
the orbital and medial prefrontal cortex in the macaque
monkey. J Comp Neurol 346:3662402.
Craig AD. 2002. How do you feel? Interoception: the sense of
the physiological condition of the body. Nat Rev Neuro-
sci 3:6552666.
Craig AD. 2009. How do you feel—now? The anterior insula
and human awareness. Nat Rev Neurosci 10:59270.
Elliot Smith G. 1907. A new topographical survey of the
human cerebral cortex, being an account of the distribu-
tion of the anatomically distinct cortical areas and their
relationship to the cerebral sulci. J Anat Physiol 41:
2372254.
Evrard HC, Logothetis NK, Craig AD. 2014. Modular architec-
tonic organization of the insula in the macaque monkey.
J Comp Neurol 522:64297.
Filley CM. 2011. Neurobehavioral anatomy. Boulder, CO: Uni-
versity Press.
Fix M. 1993. Brodmann’s numbers. Neurology 44:198421985.
Flechsig PE. 1896. Gehirn und Seele. Leipzig: Veit.
Garey LJ. 2006. Brodmann’s localization in the cerebral cor-
tex. Translation of Brodmann’s 1909 monograph
“Vergleichende Lokalisationslehre der Grobhirnrinde in
ihren Prinzipien dargestellt auf Grund des Zellenbaues”
(The principles of comparative localisation in the cerebral
cortex based on cytoarchitectonics). Berlin: Springer.
Gallay DS, Gallay MN, Jeanmonod D, Rouiller EM, Morel A.
2012. The insula of Reil revisited: multiarchitectonic
organization in macaque monkeys. Cereb Cortex 22:
1752190.
Geyer S, Weiss M, Reimann K, Lohmann G, Turner R. 2011.
Microstructural parcellation of the human cerebral cor-
tex—from Brodmann’s post-mortem map to in vivo map-
ping with high-field magnetic resonance imaging. Front
Hum Neurosci 5:19.
Gorman DG, Un€utzer J. 1993. Brodmann’s “missing” numbers.
Neurology 43:2262227.
Greenstein B, Greenstein A. 2000. Color atlas of neuro-
science: neuroanatomy and neurophysiology. Stuttgart:
Thieme.
Gu X, Hof PR, Friston KJ, Fan J. 2013. Anterior insular cortex and
emotional awareness. J Comp Neurol 521:337123388.
Jacobson S, Marcus EM. 2008. Neuroanatomy for the neuro-
scientist. New York: Springer.
Jones EG. 1985. The thalamus. New York: Plenum.
Jones EG. 2009. Brodmann’s areas. In: Squire LR, editor.
Encyclopedia of neuroscience. Oxford: Academic Press.
p 4772479.
Judas M, Cepanec M, Sedmak G. 2012. Brodmann’s map of
the human cerebral cortex—or Brodmann’s maps? Transl
Neurosci 3:67274.
The Journal of Comparative Neurology | Research in Systems Neuroscience
Brodmann’s map of cortical areas in human
Kandel ER, Hudspeth AJ. 2013. The brain and behavior. In:
Kandel ER, Schwartz JH, Jessell TM, Siegelbaum SA, Hud-
speth AJ, editors. Principles of neural science, 5th ed.
New York: McGraw-Hill. p 5220.
Klein JC, Behrens TEJ. 2009. Connectivity fingerprinting of
grey matter. In: Johansen-Berg H, Behrens TEJ, editors.
Diffusion MRI: from quantitative measurement to in-vivo
neuroanatomy. San Diego: Academic Press. p 3772402.
Krebs C, Weinberg J, Akesson E. 2012. Lippincott’s illustrated
review of neuroscience. Philadelphia: Lippincott, Williams
& Wilkins.
Loukas M, Pennell C, Groat C, Tubbs RS, Cohen-Gadol AA. 2011.
Korbinian Brodmann (186821918) and his contributions to
mapping the cerebral cortex. Neurosurgery 68:6211.
Markowitsch HJ. 1993. Brodmann’s numbers. Neurology 43:
186321864.
Martin JH. 2003. Neuroanatomy: text and atlas, 3rd ed. New
York: McGraw-Hill.
Meynert T. 1868. Der Bau der Grobhirnrinde und seine
€ortlichen Verschiedenheiter, nebst einem pathologischa-
natomischen Corollarium. Leipzig: Engelmann.
Morel A, Gallay MN, Baechler A, Wyss M, Gallay DS. 2013.
The human insula: architectonic organization and post-
mortem MRI registration. Neuroscience 236:1172135.
Nieuwenhuys R. 2012. The insular cortex: a review. Prog Brain
Res 195:1232163.
Nieuwenhuys R, Voogd J, van Huijzen C. 2008. The human
central nervous system. Berlin: Springer.
Nissl F. 1894. Uber € eine neue Untersuchungsmethode des
Centralorgans zur Feststellung der Localisation der Ner-
venzellen. Neurol Centralbl 13:5072508.
Olry R. 2010. Korbinian Brodmann (186821918). J Neurol
257:211222113.
Olry R, Haines DE. 2010. Korbinian Brodmann: the Victor Hugo of
cytoarchitectonic brain maps. J Hist Neurosci 19:1952198.
€ ur D, Ferry AT, Price JL. 2003. Architectonic subdivision of
Ong€
the human orbital and medial prefrontal cortex. J Comp
Neurol 460:4252449.
Palomero-Gallagher N, Zilles K, Schleicher A, Vogt BA. 2013.
Cyto- and receptor architecture of area 32 in human and
macaque brains. J Comp Neurol 521:327223286.
Purves D, Augustine GJ, Fitzpatrick D, Hall WC, LaMantia AS,
McNamara JO, Williams SM. 2004. Neuroscience, 3rd
ed. Sunderland, MA: Sinauer Associates.
Rose M. 1928a. Die Ontogenie der Inselrinde. Zugleich ein
Beitrag zur histogenetischen Rindeneinteilung. J Psychol
Neurol 36:1822209.
Rose M. 1928b. Die Inselrinde des Menschen und der Tieren.
J Psychol Neurol 37:4672624.
Sarkisov SA, Filimonoff IN, Kononowa EP, Preobraschenskaja
IS, Kukuew LA. 1955. Atlas of the cytoarchitectonics of
the human cerebral cortex. Moscow: Medgiz.
Seeley WW, Merkle FT, Gaus SE, Craig AD, Allman JM, Hof
PR. 2012. Distinctive neurons of the anterior cingulate
and frontoinsular cortex: a historical perspective. Cereb
Cortex 22:2452250.
Seung S. 2012. Connectome: how the brain’s wiring make us
who we are. New York: Houghton Mifflin Harcourt.
Strotzer M. 2009. One century of brain mapping using Brod-
mann areas. Clin Neuroradiol 19:1792186.
Talairach J, Tournoux P. 1988. Co-planar stereotaxic atlas of
the human brain. Stuttgart: Thieme.
ten Donkelaar HJ, Lammens M, Renier W, Hamel B, Hori A,
Verbist B. 2006. Development and developmental disorders
of the cerebral cortex. In: ten Donkelaar HJ, Lammens M,
Hori A, editors. Clinical neuroembryology: development and
developmental disorders of the human central nervous sys-
tem. Berlin: Springer. p 4292518.
13
  Ć1* and P.R. Hof
G. Simi
Toga AW, Mazziotta JC. 2002. Introduction to cartography of the
brain. In: Toga AW, Mazziotta JC, editors. Brain mapping:
the methods, 2nd ed. San Diego: Academic Press. p 3231.
Van Essen DC. 2005. Surface-based comparisons of macaque
and human cortical organization. In: Dehaene S, Duhamel
J-R, Hauser MD, Rizzolatti G, editors. From monkey brain
to human brain. Cambridge, MA: MIT Press. p 3219.
Van Essen DC, Smith SM, Barch DM, Behrens TE, Yacoub E,
Ugurbil K, WU-Minn Human Connectome Project
Consortium. 2013. The WU-Minn Human Connectome
Project: an overview. Neuroimage 80:62279.
Vogt C, Vogt O. 1926. Die vergleichend-architektonische und die
vergleich-reizphysiologische Felderung der Grobhirnrinde
unter besonderer Ber€ucksichtigung der menschlichen.
Naturwissenschaften 14:119021194.
14 The Journal of Comparative Neurology | Research in Systems Neuroscience
View publication stats
Vogt BA, Hof PR, Vogt LJ. 2004. Cingulate gyrus. In: Paxinos
G, Mai JK, editors. The human nervous system, 2nd ed.
San Diego: Academic Press. p 9152949.
Walker AE. 1940. A cytoarchitectural study of the prefrontal
area of the macaque monkey. J Comp Neurol 73:59286.
von Economo C, Koskinas GN. 1925. Die Cytoarchitektonik
der Hirnrinde des erwachsenen Menschen. Wien:
Springer.
Zilles K, Amunts K. 2010. Centenary of Brodmann’s map—con-
ception and fate. Nat Rev Neurosci 11:1392145.
Zilles K, Schleicher A, Palomero-Gallagher N, Amunts K. 2002.
Quantitative analysis of cyto- and receptor architecture
of the human brain. In: Toga AW, Mazziotta JC, editors.
Brain mapping: the methods, 2nd ed. San Diego: Aca-
demic Press. p 5732602.