Brodmann’s Areas 477
Brodmann’s Areas
E G Jones, University of California, Davis, CA, USA
ã 2004 Published by Elsevier Ltd.
This article is reproduced from the previous edition ã 2004,
Elsevier B.V.
The concept of the cortical area or cortical field has
its origins in comparative cytoarchitectonic studies of
the cerebral cortex made in the first decade of the
twentieth century by a number of anatomists, partic-
ularly Bolton, Lewis, Campbell, Brodmann, C Vogt,
and O Vogt. Among these names, that of Brodmann is
perhaps the best remembered since the numbers he
assigned to the cortical fields he identified have
entered into the common vocabulary of cortical anat-
omy and physiology. The numbers simply indicate the
order in which Brodmann studied them (Figure 1).
Cytoarchitectonics, and myeloarchitectonics, the
complementary technique that followed it, have as
their basis the recognition that different regions of
the cerebral cortex have reproducible differences in
cell and fiber layering and packing density. Architec-
tonics is the art of determining where the boundaries
fall between different areas. The early impetus to this
type of study came first from phrenology, then from
demonstrations of the electrical excitability of the
motor cortex, and from the recognition of the depen-
dence of vision on posterior regions of the cerebral
cortex. The identification of cytoarchitectonic areas
and their borders thus became a search for areas of
the cerebral cortex with distinct functions. The visual,
motor, and somatic sensory areas were the first iden-
tified and later auditory, associational, and many
further subdivisions of the original areas have been
described. In the human brain Brodmann recognized
nearly 50 areas, Campbell less than 20, C Vogt and
O Vogt more than 200. Progressively fewer were
found in the cortices of lower mammals, though Brod-
mann argued that the sensory and motor areas could
be identified in all. At the moment, most neuroscien-
tists would probably agree that Brodmann’s divisions
come somewhere near what seems reasonable, though
after a period of the converse, the trend now appears
to be going in the direction of a finer and finer parcel-
lation, akin to that of the Vogts, as functional sub-
divisions are recognized within areas by new techniques
and as imaging studies reveal new functional localiza-
tions. Brodmann’s numbered fields have been given
names in other nomenclatural schemes, and individ-
ual areas in modern experimental studies have some-
times been given specific names, but in general there is
a remarkable correspondence between schemes.
Where new fields have been described, they com-
monly represent further parcellations of larger regions
of cortex identified as single fields by Brodmann.
The persistence of the architectonic field concept
and its resurgence after the attacks on it of the 1950s
and 1960s, stem from the facts that structural varia-
tion can be seen across the cortex and that many of
these structural variations can be correlated with dis-
tinct patterns of input–output connections, with
distinct physiological properties, and often with
definable aspects of sensation, perception, cognition,
and motor control.
A modern definition of a cortical field would prob-
ably encompass the following points: (1) It is an area
of singular cellular and fiber architecture often with
boundaries that are particularly sharp and recogniz-
able even by the inexperienced eye. The border
between areas 17 and 18 in the visual cortex is
probably the most clear-cut. Borders between other
areas are not always so sharp and one area may
appear to give place to another over a gradient of
architectonic change. This may represent a truly
blurred border or be imposed by stretching of the
cortex in the floor of a sulcus or on the crown of a
gyrus and then present a trap for the unwary who
may see a distinct field where one does not exist. (2)
It is an area that receives afferent fibers from the
major population of cells in a defined relay nucleus
of the thalamus. This input stops abruptly at an
architectonic border and often serves to demonstrate
the position of a sharp border when architectonics
alone suggests a gradient of change. There appears to
be no overlap between inputs from different specific
thalamic nuclei to adjacent fields. Fibers arising from
so-called nonspecific thalamic nuclei and from sub-
sidiary populations of cells in a relay nucleus, how-
ever, often disregard these borders. (3) It receives a
set of corticocortical and commissural axons from a
limited, defined, and constant set of other cortical
areas. These tend to respect the same boundaries
delimited by architectonics and by the distribution
of thalamic afferents. (4) It gives rise to a constant
set of output connections to a limited and constant
set of cortical and subcortical targets, including
returning fibers to those thalamic nuclei (specific
and nonspecific) from which it receives inputs.
(5) A topographic map of a receptive periphery
such as the visual field, auditory space, the body
surface, or of the body’s musculature can be detected
in certain cortical fields by evoked potential or
microelectrode mapping or by electrical stimulation
478 Brodmann’s Areas

4
8 31 2
5
7a
8
7b

19

46
39 18
10

45 43 41
52 42
17
47 22
11
21 18
19
38 37

20

4 31 2
6 5
8 7

9
31
24
23
32 19
33

10 18
26
29
30
25 27
12 17
34 35
11
32

13 18
36 19
35

27

Figure 1 Brodmann’s map of the areas in the human cerebral cortex. (Top) lateral view; (bottom) medial view. From Brodmann K (1910)
Feinere Anatomie des Grosshirns. In: Lewandowsky M (ed.) Handbuch der Neurologie, Vol. 1: Allgemeinere Neurologie, pp. 206–307.
Berlin: Springer.

of the field. In association with these maps, the neu-
rons in the field may display physiological properties
that distinguish them from those in other fields.
Aggregation of subpopulations of cells with different
functional properties can occur in a field and some-
times, as in some of the visually related areas, these
aggregations may have definable anatomical char-
acteristics. These characteristics may then serve as
a basis for further subdividing the cortical field.
(6) Deactivation of a field by ablation, cooling, or
injection of drugs may lead to loss of a particu-
lar sensory or motor function or of other behaviors.
(7) The field may show a characteristic pattern of
histochemical or immunocytochemical staining dif-
ferent from that of its neighbors. The so-called
chemoarchitectonic borders defined in this manner
commonly coincide with those defined by cell and
fiber staining.
Obviously, not every cortical field can be demon-
strated to meet all these criteria, primarily because of

the difficulty in eliciting physiological responses in
many of them or in defining the nature of a deficit
resulting from their deactivation. Similarly, argu-
ments invariably ensue about the sharpness of certain
architectonic borders and about whether particular
architectonic changes indicate the existence of new
fields or simply variation within a field, a variation
perhaps resulting from a local concentration of cells
projecting to the same site or with a particular physio-
logical property. The connectional basis of the cortical
field, especially that defined by connections with a
particular principal thalamic nucleus, probably remains
the most reliable anatomical guide for deciding when a
region of cortex should be called a cortical area or field
independent of its neighbors and for determining its
boundaries. It is to be hoped that eventually molecu-
lar-genetic markers will be discovered that determine
this reproducible parcellation of the cerebral cortex on
the basis of structural and functional properties.
See also: Cerebral Cortex; Memory Consolidation:
Cerebral Cortex; Topographic Maps: Molecular
Mechanisms.
Brodmann’s Areas 479
Further Reading
Bailey P and von Bonin G (1951) The Isocortex of Man. Urbana,
IL: University of Illinois Press.
Brodmann K (1909) Vergleichende Lokalisationslehre der
Grosshirnrinde in ihren Prinzipien dargestellt auf Grund des
Zellenbaues. Leipzig: Barth.
Brodmann K (1910) Feinere Anatomie des Grosshirns. In:
Lewandowsky M (ed.) Handbuch der Neurologie, Vol. 1: All-
gemeinere Neurologie, pp. 206–307. Berlin: Springer.
Campbell AW (1905) Histological Studies on the Localization of
Cerebral Function. London: Cambridge University Press.
Flechsig P (1920) Anatomie des menschlichen Gehirns und Rücken-
marks auf myelogenetischer Grundlage. Leipzig: Thieme.
Jones EG (1985) The Thalamus. New York: Plenum.
Jones EG and Burton H (1976) Areal differences in the laminar
distribution of thalamic afferents in cortical fields of the insular,
parietal and temporal regions of primates. Journal of Compara-
tive Neurology 168: 197–248.
Lewis JW and Van Essen DC (2000) Mapping of architectonic
subdivisions in the macaque monkey, with emphasis on
parieto-occipital cortex. Journal of Comparative Neurology
428: 79–111.
Peters A and Jones EG (eds.) (1984–1994) Cerebral Cortex,
vols. 1–10. New York: Plenum.
von Economo C and Koskinas CN (1925) Die Cytoarchitektonik
der Hirnrinde des erwachsenen Menschen. Berlin: Springer.