Review Article • DOI: 10.2478/s13380-012-0009-x • Translational Neuroscience • 3(1) • 2012 • 67-74 Translational Neuroscience

BRODMANN’S MAP OF THE HUMAN CEREBRAL CORTEX Miloš Judaš1,*, Maja Cepanec1,2, – OR BRODMANN’S MAPS? Goran Sedmak1 Abstract 1Croatian Institute for Brain Research, The cytoarchitectonic map of the adult human cerebral cortex prepared by Korbinian Brodmann is probably the University of Zagreb School of Medicine, most widely used and reproduced cortical map. However, few people today realize that in the period between 10000 Zagreb, Croatia 1908 and 1914 (the first and last date of maps publication by Brodmann himself) that map was gradually developed, extended and subjected to some significant modifications. The aim of this article is to reproduce and 2Department of Speech and Language Pathology, briefly describe all versions of Brodmann’s map, trace its changes and highlight the importance of these changes University of Zagreb Faculty of Education – especially with respect to the speech and language related cortical regions of the left cerebral hemisphere. and Rehabilitation Sciences, 10000 Zagreb, Croatia Keywords • Cytoarchitectonics • Human brain mapping • History of neuroscience • Korbinian Brodmann

Received 24 February 2012 © Versita Sp. z o.o. Accepted 28 February 2012

Introduction high-quality histological microphotographs really consulted. This is probably due to the and camera lucida drawings of several areas fact that Brodmann’s original studies were During the first decade of the 20th century, of the human cerebral cortex to convince a published in German and in journals which are Korbinian Brodmann (1868-1918; for his reader that his map rests on a solid histological not easily accessible to many present students, biography, see [1]; for his contribution to foundation. That much was at least clear to and that English translation is available only neuroscience, see [2]) published a series Brodmann’s contemporaries who not only for his book published in 1909. Even the book of seminal contributions which represent read his publications but also attended remained untranslated for almost a century: the foundation of the modern comparative his public lectures and demonstrations of the first partial translation (a single chapter) cytoarchitectonics of the mammalian cerebral histological sections [15-18]. was published by Gerhard von Bonin in cortex. His principal work was published As pointed out in a recent review [19], one 1960 [22], and the complete translation was between 1903 and 1908 in the Journal für of Brodmann’s greatest merits was to integrate published by Lawrence Garey in 1994 [23]. Psychologie und Neurologie as a series of evolutionary ideas and the histological analysis Thus, most present authors seem to neglect seven monographs [3-9]. These findings were of the cortex with functional localization. a simple fact: there is no single „Brodmann’s summarized in his famous book [10] as well as Brodmann’s cortical map is probably the most map“ but instead several versions of it, which in two extensive chapters which he contributed widely known and used architectonic map of were gradually modified and completed to leading contemporary handbooks [11,12]. the human cerebral cortex and it has been between 1908 and 1914 (i.e., the dates of the Furthermore, Brodmann published two reproduced in a number of textbooks and first and the last publication of the mapby important supplementary studies in which handbooks. In fact, with the introduction of Brodmann himself). As already noted [21], he described his new architectonic findings novel neuroimaging techniques, Brodmann’s a common mistake is to reproduce a version in brains of apes (gibbons and chimpanzees) map has received ever increasing popularity of the map which was in fact published in and additional human specimens [13,14]. – probably the most popular example being 1910 (or 1914), but to quote the source as Although he planned to publish a complete the stereotaxic atlas of Talairach and Tournoux „Brodmann 1909“. cytoarchitectonic atlas of the human [19-21]. Although Brodmann’s map is so Therefore, the aim of this paper is to provide brain (and thus provide a comprehensive popular, it remains a sad fact that Brodmann’s reproductions and brief descriptions of all histological documentation for his cortical original papers today remain largely unknown versions of the „Brodmann’s map“, to trace its map), this plan was never realized due to the or even misquoted. Indeed, the very pattern of changes and highlight the importance of these outbreak of World War I and his premature reproducing and quoting „Brodmann’s map“ changes – especially with respect to the speech death in 1918. Nevertheless, Brodmann’s in a number of recent publications seems to and language related cortical regions of the left publications contain a sufficient number of suggest that the „original Brodmann“ was not cerebral hemisphere.

* E-mail: [email protected]

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Brodmann’s map: The original version

While Brodmann already published a partial map of motor/premotor, somatosensory and visual cortical areas in his initial studies on the human brain [3,4], the first complete version of the human cortical map (Figure 1) was published twice in 1908 [8,9], as figures 5 and 6 on p. 236 of his paper on areal parcellation of the human cortex [8] and reproduced as figures 34 and 35 on p. 316 of his paper on the cortex of lemurs [9]. The same version of the map was reproduced as figures 85 and 86 on p. 131 of his famous book [10]. It should be noted that this first version displays only lateral view of the left hemisphere and medial view of the right hemisphere, while the map of the insular and adjacent supratemporal cortex (Figure 2) was first published as a separate figure (figure 89 on p. 144) in the 1909 book [10], and this figure shows just two areas of the insular cortex (named „insula anterior + insula posterior“) and four supratemporal areas (areas 22, 41, 42 and 52). Whereas the map itself remained the same in all three publications [8-10] it should be noted that the accompanying text already displays significant modifications of Brodmann’s concepts in 1909 compared to 1908. The first significant change is related to his grouping of individual cortical fields (area) into larger regions (regio). While in the original study [8] there was just one (Rolandic) region encompassing both primary motor (area 4) and primary somatosensory cortex (areas 1, 2, 3 and 43) and all other areas were simply listed as topographically belonging to corresponding lobes, in the 1909 monograph [10] all areas were re-grouped in appropriate cytoarchitectonic regions (Table 1). Note also that in the original study the premotor cortex (area 6) was separated from the primary motor cortex (area 4) and listed under the „frontal Figure 1. The original version of the Brodmann’s map, as published in [8] (figs 5 and 6, p. 236) and reproduced lobe“. Accordingly, Brodmann published his in [9] (figs 34 and 35, p. 316) and in [10] (figs 85 and 86, p. 131). The legend in [8] and [9] simply stated first black-and-white diagram of major cortical „Lateral and medial view of the human cerebral hemisphere with cytoarchitectonic cortical areas“ and in [10] „Areal parcellation of the lateral and medial hemispheric surface in the human brain“. Names and regions (Figure 3) as figures 83 and 84 on p. 128 brief description of cortical areas were provided in the main text. Note that figure claims to represent a of his book [10]. The major changes represented single hemisphere, but in fact displays lateral view of the left and medial view of the right hemisphere. in this diagram are focused on the division of the frontal lobe cortex: previous Regio Rolandica consists of two regions (Regio praecentralis important changes that Brodmann mentioned is now divided in Regio postcentralis and Regio and Regio frontalis). However, even this new in the text (pp. 127-150) of his 1909 monograph praecentralis, while the entire frontal lobe now diagram of regions does not represent all (see below).

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Table 1. Brodmann’s list of cortical areas sorted according to topographical regions, i.e. cerebral lobes [8] or major cytoarchitectonic regions [10].

Brodmann (1908) [8] Brodmann (1909) [10]

Area no.: Area name: Area no.: Area name:

REGIO ROLANDICA REGIO POSTCENTRALIS 1 Area postcentralis intermedia 1 Area postcentralis intermedia 2 Area postcentralis caudalis 2 Area postcentralis caudalis 3 Area postcentralis oralis 3 Area postcentralis oralis 43 Area subcentralis 43 Area subcentralis - REGIO PRAECENTRALIS 4 Area praecentralis gigantopyramidalis 4 Area gigantopyramidalis LOBUS FRONTALIS 6 Area frontalis agranularis 6 Area frontalis agranularis - REGIO FRONTALIS 8 Area frontalis intermedia 8 Area frontalis intermedia 9 Area frontalis granularis 9 Area frontalis granularis 10 Area frontopolaris 10 Area frontopolaris 11 Area praefrontalis 11 Area praefrontalis 44 Area opercularis 44 Area opercularis 45 Area triangularis 45 Area triangularis 46 Area frontalis media 46 Area frontalis media 47 Area orbitalis 47 Area orbitalis LOBUS PARIETALIS REGIO PARIETALIS 5 Area praeparietalis 5 Area praeparietalis 7 Area parietalis superior 7 Area parietalis superior 39 Area parietalis inferior posterior (= Area angularis) 39 Area angularis 40 Area parietalis inferior anterior (= Area supramarginalis) 40 Area supramarginalis LOBUS OCCIPITALIS REGIO OCCIPITALIS 17 Area striata 17 Area striata 18 Area occipitalis 18 Area occipitalis 19 Area praeoccipitalis 19 Area praeoccipitalis LOBUS TEMPORALIS REGIO TEMPORALIS 20 Area temporalis inferior 20 Area temporalis inferior 21 Area temporalis media 21 Area temporalis media 22 Area temporalis superior 22 Area temporalis superior 36 Area ectorhinalis 36 Area ectorhinalis 37 Area occipitotemporalis 37 Area occipito-temporalis 38 Area temporopolaris 38 Area temporopolaris 41 Area (temporalis) transversa interna s. anterior 41 Area temporalis transversa interna (anterior) 42 Area (temporalis) transversa externa s. posterior 42 Area temporalis transversa externa (posterior) - 52 Area parainsularis LOBUS LIMBICUS REGIO CINGULARIS - Subregio postcingularis: 23 Area limbica posterior ventralis 23 Area cingularis posterior ventralis 31 Area limbica posterior dorsalis 31 Area cingularis posterior dorsalis - Subregio praecingularis: 24 Area limbica anterior ventralis 24 Area cingularis anterior ventralis 32 Area limbica anterior dorsalis 32 Area cingularis anterior dorsalis 33 Area praegenualis 33 Area praegenualis 25 Area subgenualis 25 Area subgenualis - REGIO RETROSPLENIALIS 26 Area ectosplenialis 26 Area ectosplenialis 29 Area retrolimbica granularis 29 Area retrolimbica granularis 30 Area retrolimbica agranularis 30 Area retrolimbica agranularis - REGIO HIPPOCAMPICA 27 Area praesubicularis 27 Area praesubicularis 28 Area entorhinalis ventralis 28 Area entorhinalis 34 Area enthronialis dorsalis 34 Area entorhinalis dorsalis 35 Area perirhinalis 35 Area perirhinalis - REGIO INSULARIS - - Insula anterior + Insula posterior - REGIO OLFACTORIA

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Brodmann’s map: The modified (final) version

The modified version of both the areal (Figure 4) and the regional (Figure 5) map appeared in 1910, as figure 49 on p. 212 (areal map) and color figures 83 and 84 on p. 228 (regional map) of his chapter in Lewandowsky’s handbook of neurology [11]. This final version of the areal map was reproduced as figures 76 and 77 on p. 100 of Brodmann’s chapter in 11. volume of the Neue Deutsche Chirurgie [12] which probably represents the source of most reproductions in the current literature (frequently wrongly Figure 2. The first and only version of the Brodmann’s map of the insular and adjacent supratemporal cortex quoted as stemmingfrom „Brodmann 1909“ i.e. [10] (fig. 89, p. 144). Legend: Regio insularis and Facies superior of the freely exposed Gyrus temporalis [10]). superior. J. ant. = agranular anterior insular region; J. post. = granular posterior insular region; sp = Ramus posterior; sv = Ramus verticalis; sh = Ramus horizontalis fissurae Sylvii; t1 = sulcus temporalis As already mentioned, these new maps superior. There are three cortical areas on the superior surface of the superior temporal gyrus: 52 = Area codified Brodmann’s evolving concept of major parainsularis; 41 = Area temporalis transversa interna s. anterior; 42 = Area temporalis transversa externa s. posterior. cytoarchitectonic regions. In 1908, there was just one region (Regio Rolandica) plus 5 lobes (excluding insula). In 1909 [10], p. 129 there were already 11 architectonic regions (see Table 1, right column): Rolandic region was split into precentral and postcentral region, limbic lobe was split into three regions (Regio cingularis, Regio retrosplenialis and Regio hippocampica), and two new regions (Regio insularis, Regio olfactoria) were introduced. Finaly, in 1910 [11] (pp. 225-230) one encounters confusing discrepancies between the regional color map of 1910 (Figure 5), its legend and the main text. Namely, the text states that there are eight „clinically most important“ regions (precentral, postcentral, frontal, insular, occipital, temporal, and cingular). Moreover, there is within the frontal region a separate „Subregio frontalis inferior“ and within the temporal region a separate „Regio (sic) supratemporalis“. On the other hand, the color map of regions and its legend also include hippocampal and retrosplenial region – but olfactory region of 1909 is now missing! In addition, Regio cingularis in the color map is subdivided into „anterior and posterior“, while in the text these Figure 3. The initial black-and-white diagram of major cytoarchitectonic regions ([10] figs 83 and 84, p. 128). two subregions are described as agranular Legend states briefly: „Major cytoarchitectonic regions in man. The olfactory region is not marked Subregio praecingularis and granular Subregio in the map“. However, on p. 129 in the text the following 11 regions are listed: 1. Regio postcentralis, postcingularis. Obviously, both Brodmann’s 2. Regio praecentralis, 3. Regio frontalis, 4. Regio insularis, 5. Regio parietalis, 6. Regio temporalis, 7. Regio occipitalis, 8. Regio cingularis, 9. Regio retrosplenialis, 10. Regio hippocampica and 11. Regio olfactoria. classification and terminology were in the state of continuous fluctuation. For example, area 7 is now subdivided in two and area 52 (area parainsularis) is now inserted A closer look at the revised areal map subfields (7a and 7b), area 12 is now introduced in the main map of the lateral hemispheric (Figure 4) reveals other important novelties. as dorso-medial half of the previous area 11, surface. However, none of these important

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changes were not even mentioned in the text of the 1910 handbook chapter! Instead, to understand these changes, one has to consult the text of his 1909 book [10] (pp. 136-150) where Brodmann proposed the following changes of his initial map: (a) Area 11 (Area praefrontalis) may be subdivided in three subfields or even separate areas (Area rostralis, Area recta and Area orbitalis interna); (b) Area 7 (Area parietalis superior) may be subdivided in two subfields, one in front of the superior parietal sulcus (Area parietalis superior anterior – 7a) and one behind it (Area parietalis superior posterior – 7b); (c) The posterior part of the area 36 (Area ectorhinalis) may also be described as a separate Area retrosubicularis (already described in animals as area 48); (d) Instead of only two areas on the superior surface of the superior temporal gyrus (as described in [8]), there are in fact three such areas – area 52 (Area parainsularis) being newly described one, and all three included in the „auditory cortex“; (e) In the insular cortex (p. 146) Brodmann distinguished four areas already in 1904, but now presents only anterior agranular and posterior granular area in the fig. 89 (see our Figure 2). However, he also previously described as Pars olfactoria insulae another area situated in the limen insulae, as well as another area at the transition from insula to orbitofrontal gyri; (f) In the Regio cingularis Brodmann describes two clearly distinct (anterior agranular vs. posterior granular) subregions – Subregio postcingularis and Subregio praecingularis – although they are not mentioned in either areal or regional map. Moreover, there is between these two subregions a transitional Area cingularis intermedia which is also not noted in the map. Thus, the modern reader has to cope with somewhat paradoxical situation. In the 1909 book there is original (unrevised) areal map, but careful reading of the text reveals a number of important changes and additions. On the other Figure 4. The modified (final) version of the Brodmann’s areal map ([11] figs 81 and 82, p. 226; reproduced as hand, in the [11] (and [12]) chapter there is figs 76 and 77, p. 101, in[12]). The legend in [11] is laconic („Cytoarchitectonic cortical areas after Brodmann“) and there is no description of changes in the accompanying [11] text – the reasons for new, revised map – but the accompanying text changes were listed on pp. 127-150 of the 1909 book [10] which featured the old map (sic). The legend in provides no description or explanation of these [12] states: „Cortical areal map according to cytoarchitecture (Brodmann). Compare with physiologically changes. To really understand the „Brodmann’s important major regions in figs 78 and 79 (= Figure 7 in this paper). 1. Regio praecentralis: areas 4 and 6; 2. Regio postcentralis: areas 1, 2, 3 and 43; 3. Regio frontalis: areas 8-12; 4. Regio insularis: areas 13-16; 5. map“, one has to entertain oneself with a Regio parietalis: areas 5, 7 (a + b), 39 and 40; 6. Regio temporalis: areas 20, 21, 22, 36, 37, 38, 41, 42 and 52; collated reading of Brodmann’s original papers 7. Regio occipitalis: areas 17-19; 8. Regio cingularis: areas 23-25 and 31-33; 9. Regio retrosplenialis: areas 26, 29 and 30; 10. Regio hippocampica: areas 27, 28, 41 and 35; 11. Regio olfactoria.“ Note that this is the published between 1908 and 1914. first time that insular cortex is described as consisting of areas 13-16. Note also that, quite conspiciously, areas 44, 45 and 47 are not mentioned in the legend.

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Significant and continuous1910, pp. 227-230), Brodmann introduced of the regional map ([13] fig. 49, p. 212; changes in the fronto-opercular designations 44 and 44a in his revised areal reproduced as [12] fig. 108, p. 166) as well as region (Broca’s area) map (Figure 4) and described areas 44, 45 and in the modified color version (Figure 7) of the 47 as a separate Subregio frontalis inferior in regional map ([12] fig. 78, p. 102). Moreover, the The most important (and probably the his new regional map (Figure 5). The influence terminology changed once more: to Subregio most confusing) changes are related to of Oscar Vogt is now explicitely recognized, infrafrontalis [13] and to Regio infrafrontalis Brodmann’s evolving conception of the as Brodmann pointed out that this new including the „Broca’s speech zone“ [12]. Thus, language-related fronto-opercular cortex. To region almost perfectly overlaps with Vogt’s Brodmann’s concept of Broca’s speech region understand these changes, one has to deal myeloarchitectonically defined Subregio steadily expanded in both topographical (from with two issues: (a) Brodmann’s elaboration unitostriata infrafrontalis. Thus, the entire left single area to the entire gyrus) and hierarchical of the initial architectonic parcellation, which inferior frontal gyrus was transformed into a status (from single area to the independent was obviously and heavily influenced by new, human-specific and language-related architectonic region). Namely, it started as concurrent myeloarchitectonic parcellation region, well-delineated by both cyto- and Broca’s area (44) and Broca’s region (44+45) in of the human frontal cortex as elaborated by myeloarchitectonic methods. This concept was 1908, but soon expanded to the entire inferior Oscar Vogt; and (b) Brodmann’s continuously further developed in subsequent publications frontal gyrus encompasing all three areas (44, changing as well as inconsistent and confusing [12-14] and in two further modifications of the 45, 47) and was progressively designated as changes of terminology. The original areal regional map. The entire inferior frontal gyrus (Sub)regio subfrontalis (1909), Subregio frontalis map [8-10] displays a single area 44 (Area is now depicted as separate subregion in the inferior (1910), Subregio infrafrontalis (1912) opercularis) as well as area 45 (Area triangularis) modified black-and-white version (Figure 6) and Regio infrafrontalis (1914). and area 47 (Area orbitalis) in the region of the inferior frontal gyrus. All three areas were initially described as parts of a single topographic region (Lobus frontalis in 1908) or a single architectonic region (Regio frontalis in 1909). In the initial description ([8] pp. 241- 242) Brodmann mentioned certain differences between rostral and caudal part of the area 44 (separated by sulcus diagonalis), but concluded that these differences are „so unimpressive“ (so unbeträchtlich) that they cannot lead to distinction of two subfields within the area 44. Moreover (Brodmann 1908a, pp. 244-245), he described area 44 as „Area Broca“ (Brocasche Stelle) and areas 44 and 45 together as „Regio Broca“ – leaving aside area 47 as not important for language. However, next year ([10] pp. 138- 139) he concluded that it may be justified to divide area 44 in two subareas (Area opercularis anterior – 44a + Area opercularis posterior – 44), but still regarded this difference as not very important because of the pronounced interindividual variability and inconstancy of the diagonal sulcus and thus retained a single area 44 in his areal map. On the other hand, he further noted in the text that area 47 displays „certain structural similarity“ with areas 44 and Figure 5. The modified color version of the Brodmann’s regional map ([11] figs 83 and 84, p. 228). Legend: Major 45, so that all three areas could be described cytoarchitectonic regions (Hauptzonen) in the man. Light brown = Regio frontalis with dark-brown subregion (Binnenfeld). Dark-brown = Subregio frontalis inferior. Dark-blue = Regio praecentralis. as a separate subregion – which on p. 137 he Orange = Regio postcentralis. Green = Regio parietalis (the small dark-green field isArea praeparietalis). designated as Regio subfrontalis, but only few Light-red = Regio occipitalis with „Binnenfeld“ 17. Dark-red = Area striata (area 17 in areal map on pages later (p. 139) renamed into Subregio figs. 81 and 82). Light-violet = Regio temporalis with subregion (Binnenfeld). Dark-violet = Regio supratemporalis. Black and white stripes = Regio insularis (anterior and posterior). Yellow = Regio subfrontalis. In his 1910 chapter (Brodmann cingularis (anterior and posterior). Yellow dots = Regio hippocampica. Light-blue = Regio retrosplenialis.

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Figure 6. The modified black-and-white version of the Brodmann’s regional Figure 7. The modified color version of the Brodmann’s regional map of major map of the frontal lobe ([13] figs 49 and 50, p. 212; reproduced as cytoarchitectonic regions ([12] figs 78 and 79, p. 102). Legend: „The Regio fig. 108, p. 166 in [14]). Legend in [13]: Lateral and medial view of praecentralis (areas 4 and 6) roughly corresponds to the electromotor zone, the human cerebral hemisphere (schema) showing the relationship and Regio postcentralis (areas 1-3) to the sensory zone. Area striata (17) is between Regio praecentralis (4+6) and Regio frontalis. Within the visual cortex (Sehsphäre), and Regio supratemporalis (T1) is auditory cortex frontal region there is Subregio infrafrontalis which, according to my (Hörsphäre?) whereas Regio hippocampica represents the olfactory cortex earlier studies, should be distinguished and covers an entire gyrus (Riechsphäre). The Regio infrafrontalis (F3) includes Broca’s speech zone (F3, i.e. inferior frontal gyrus). Area striata (17) is almost completely (Brocasche Sprachzone). The physiological role of Regio retrosplenalis (Rspl.) situated at the medial hemispheric surface. is completely unknown“.

Acknowledgements

This work was supported by Croatian Ministry 1081870-1878 to M.J. Authors declare no of Science, Education & Sport grant No. 108- conflict of interest.

References

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