Brodmann's Map of the Human Cerebral Cortex
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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 map by 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] 67 Translational Neuroscience 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). 68 Translational Neuroscience 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