Cellular Structure of The Human Cerebral Cortex (2009)

Cellular Structure of the Human Cerebral Cortex
Constantin von Economo

Professor of Neurology and Psychiatry at the University of Vienna, Austria
Cellular Structure
of the Human
Cerebral Cortex
Translated and edited by
Lazaros C. Triarhou
Professor of Neuroscience and Chairman of Educational Policy,
University of Macedonia, Thessaloniki, Greece
88 gures including 48 original microphotographs, 4 in color, 12 tables, and fold-out, 2009
Basel Freiburg Paris London New York Bangalore

Bangkok Shanghai Singapore Tokyo Sydney
New English edition

Originally published in German and in French under the titles:
Zellaufbau der Grosshirnrinde des Menschen.
Zehn Vorlesungen mit 61 Abbildungen.
Berlin, Verlag Julius Springer, 1927.
Larchitecture cellulaire normale de lcorce crbrale.
Paris, Masson et Cie, Editeurs, 1927.
Frontispiece portrait information:

Previously unpublished photograph of Professor Constantin von Economo (18761931) from the
album dedicated by pupils and colleagues to Hofrat Professor Julius Wagner von Jauregg (1857
1940), winner of the 1927 Nobel Prize in Physiology or Medicine for his discovery of the therapeutic value of malaria inoculation in the treatment of dementia paralytica. Photo by Max
Schneider, Vienna. Courtesy: Bildarchiv, Institut fr Geschichte der Medizin, Medizinische Universitt Wien, Austria. Signature from a 1911 patient report, hand-written by Economo in Wagner
von Jaureggs neurology clinic (editors archive).
Library of Congress Cataloging-in-Publication Data

Economo, Constantin, Freiherr von, 18761931.
[Zellaufbau der Grosshirnrinde des Menschen. English]
Cellular structure of the human cerebral cortex / Constantin von Economo ;
translated and edited by Lazaros C. Triarhou.
p. ; cm.
Originally published also in French under the title: Larchitecture cellulaire normale de lcorce crbrale. 1927.
Includes bibliographical references and index.
ISBN 9783805590617 (hard cover : alk. paper)
1. Cerebral cortex--Cytology. I. Triarhou, Lazaros Constantinos, 1957- II. Title.
[DNLM: 1. Cerebral Cortex--cytology. WL 307 E19z 2009a]
QM455.E213 2009
612.825--dc22
2009011546
Copyright 2009 by S. Karger AG, P.O. Box, CH4009 Basel (Switzerland)

www.karger.com
Printed in Switzerland on acid-free paper by Reinhardt Druck, Basel
ISBN 9783805590617
e-ISBN 9783805590624
Dedicated
as an indication of admiration to
Hofrat Professor
Julius Wagner von Jauregg
in whose Psychiatric Clinic in Vienna
these lectures were given
Lazaros C. Triarhou
Professor of Neuroscience
University of Macedonia
156 Egnatia Ave.
GR54006 Thessaloniki (Greece)
This book was sponsored by the Academy of Athens to honor Constantin von Economo as the
first laureate of its Aristeion of Science Award (1928), with additional grant support from the Bodossakis Foundation and University of Macedonia, Greece; the Hellenic Neurological Society is
also acknowledged.
Disclaimer. The statements, opinions and data contained in this publication are solely those of the individual
authors and contributors and not of the publisher and the editor(s). The appearance of advertisements in the book is
not a warranty, endorsement, or approval of the products or services advertised or of their effectiveness, quality or
safety. The publisher and the editor(s) disclaim responsibility for any injury to persons or property resulting from any
ideas, methods, instructions or products referred to in the content.
All rights reserved. No part of this publication may be translated into other languages, reproduced or utilized
in any form or by any means electronic or mechanical, including photocopying, recording, microcopying, or by any
information storage and retrieval system, without permission in writing from the publisher.
Contents
VIII
Preface Acknowledgements
L.C. Triarhou
XVII
Vorwort Prface Prefazione Preface

C. von Economo
001
Introduction
032
Frontal Lobe
073
Parietal Lobe
095
Insular Lobe
102
Occipital Lobe
114
Temporal Lobe
133
Superior Limbic Lobe

Cingulate Gyrus and Retrosplenial Region
149
Hippocampal (Inferior Limbic) Lobe

Hippocampal Gyrus, Dentate Gyrus and Uncus
173
Conclusion
194
Appendix
An Outline of Cytoarchitectonics of the Adult Human Cerebral Cortex
G.N. Koskinas
227
References
235
List of Persons
237
Subject Index
This volume aims at awakening one of the classics of the neuroscience literature, Constantin
von Economos Zellaufbau der Grosshirnrinde
des Menschen (Cellular Structure of the Human
Cerebral Cortex) [1], which was simultaneously
published in French as The Normal Cellular Architecture of the Cerebral Cortex [2], followed by
an Italian version under the title The Cytoarchitectonics of the Human Cerebral Cortex [3], and
an initial English translation in due course [4].
More than a mere new translation, the present
book can be viewed as a slightly revised second
edition, complementing the Atlas of Cytoarchitectonics of the Adult Human Cerebral Cortex by
Economo and Koskinas [5]. The changes are discreet, maintaining the core structure of the book,
with modifications for the sake of functionality
and thoroughness.
While the voluminous German Textband (xl +
810 pp.) [6] that accompanied the original Atlas
[7] remains an invaluable encyclopedic resource,
the compact Zellaufbau [14] has the advantage
of a handy and topical overview of a more didactic nature, based on a series of teaching lectures
given by Professor von Economo, epitomizing his
updated views on cerebral cortical cytoarchitectonics [8, 9].
VIII
The German edition of the book (146 pp.) [1]

was structured into ten lectures: two introductory, two on the frontal lobe, two on the parietal
lobe (the second covering the insula as well), one
each on the occipital and temporal lobes, and two
on the limbic lobe (subdivided into the cingulate
gyrus or superior limbic lobe and the hippocampal gyrus or inferior limbic lobe). Each lecture
began with the exclamation M.H.! (Meine Herren! or Gentlemen!). The book is distinguished by
the typical intricate syntax with long clauses and
paragraphs.
The French edition (183 pp.) [2], edited by the
Belgian neuropathologist and Economos friend
Ludo van Bogaert (18971989) [10, 11], was restructured into eight chapters: the introduction
was combined into one, the frontal lobe still occupied two chapters, the parietal, insular, occipital and temporal lobes formed individual chapters, and the cingulate and hippocampal gyri
were combined into a single chapter on the grand
lobe limbique de Broca.
The Italian edition (192 pp.) [3] is pleasantly
comprehensible and precise in terminology. It
was overseen by Professor Giovanni Mingazzini
(18591929) of the University of Rome, a veteran
representative of the Italian school of neurology
[12, 13]. Like the German original, it contained

ten chapters. In his introductory commentary,
Mingazzini extols Economos oration as coupling Demosthenian eloquence with Horatian
simplicity.
The English edition (186 pp.) [4] was rendered
by Sam Parker (19011990), who also translated
Economos article Encephalitis lethargica and
encphalomylite subaigu diffuse of Cruchet
for the Journal of the American Medical Association [14]. A native of Kiev (then Russian Empire),
Parker obtained his B.S. from Columbia University, M.D. from Vienna University, and S.J.D. in
Psychiatric Criminology from New York University [15]. When he translated the book, he was a
young trainee in psychoanalysis under Sigmund
Freud (18561939) and an assistant to Julius Wagner von Jauregg (18571940) at the Vienna Psychiatric Clinic; he later became affiliated with the
New York Psychiatric Institute and Brooklyn
Kings County Hospital, and practised privately
in California [16, 17]. The book was expanded to
twelve chapters, splitting the insula from the second parietal lobe lecture, and adding a concluding chapter with Economos later views on cytoarchitectonic neuropathology and evolutionary
neuroscience: in the human cerebral cortex,
Economo contemplated an organ in transition
between its past and future evolution, with its
theoretical potentialities not fully realized [24].
For practical reasons, the present volume has
been restructured into nine chapters: one for each
of the seven defined cerebral lobes, plus the introductory and concluding chapters.
The new translation was based on the original
German and French texts; the Italian and the
previous English edition were consulted. Being
multilingual [25], Economo would use idiomatic
language variations in each edition. A comparison of all four editions [14], paragraph by paragraph, provided the means for a quality check in
the case of the occasional error, and a thesaurus
of synonyms or alternative terms toward a naturally flowing text. Mild variations or sentences
specifically appearing in one or the other of the

four previous editions have been included and
are indicated by citing the corresponding book.
In deference to Economos cosmopolitan character that incorporated the exquisite Triestine
and Viennese cultures into his Hellenic, Romanian and Hungarian heritage, and linguistic fluency that comprised Ancient and Modern Greek,
Latin, Italian, German, French and English [25],
all four original prefaces are reproduced by permission from Springer Verlag, ditions Elsevier
Masson, Licinio Capelli Editore, and Oxford
University Press, respectively.
Worthy heirs of Alcmaeon of Croton, the discoverer in 500 BC of the brain as the locus of
mind ( or leading principle) [26, 27],
Economo and Koskinas [6] meticulously collected a huge amount of information on the structure
of the human cerebral cortex over the years. This
new book provides the majesty of their quantitative data [9]. The work is considered to be unsurpassed in both its scientific eloquence and accurate photographic documentation [28], and represents the most comprehensive set of high
resolution images of cortical histology ever assembled [9].
The Economo-Koskinas system of areas was
adopted in classic neurologic works as opposed to
Brodmann areas by some of the foremost academic scientists in the field, including Otto Marburg (18741948) in Vienna [29], Alfons Maria
Jakob (18841931) in Hamburg [30], Kurt Goldstein (18781965) in Frankfurt am Main [31],
Maximilian Rose (18831937) in Vilnius [32],
Christofredo Jakob (18661956) in Buenos Aires
[26], and Gerhardt von Bonin (18901979) in
Chicago [33].
There are three major advantages in using
Economo-Koskinas areas over Brodmann areas:
(1) Timing of publication: Brodmann published his monograph in 1909. Economo began
work on cytoarchitectonics in 1912, with Koskinas joining in 1919; their Textband and Atlas [6,
IX
7] were published in 1925, almost two decades after Brodmann, and included 150 new discoveries
[34]. Economo continued with further studies
through 1931.
(2) Cytoarchitectonic fields defined: Brodmann defined 44 areas in the human, and 52 areas in the primate brain. Economo and Koskinas
defined 54 ground, 76 variant, and 107 modification areas in the human brain (table 1), plus more
than 60 additional transition areas (table 2), thus
availing of a greater resolution against Brodmann areas for the human cerebral hemispheres
by a factor of four. Brodmann correlations can be
found in the Atlas [5], a related review [35], and
the included poster. It is worth noting that, after
the publication of Zellaufbau [1], Economo and
Horn [36] further defined more than 20 modifications in the temporal lobe alone. The Economo-Koskinas cortical modifications are grouped
into five main structural types, histologically
distinct and probably functionally distinct as
well [37].
(3) Extrapolated vs. real surface designations:
Brodmann maps are commonly used to either
designate cytoarchitectonic areas as such, or as a
shorthand system to designate some region on
the cerebral surface [38]. Macroscopic extrapolation of Brodmann projection maps are effected
on the atlas of Talairach and Tournoux [39], rather than being based on real microscopic cytoarchitectonics. On the other hand, the unique sectioning method of Economo and Koskinas,
whereby each gyrus is dissected into blocks always perpendicular to the gyral surface, be it
dome, wall or sulcus floor, essentially offers a
mechanical solution to the generalized mapmakers problem of flattening nonconvex polyhedral surfaces [40], one of the commonest problems at the epicenter of cortical research.
Moreover, merely looking at figures 6, 74, 86,
87 and 88, one may ponder over the fact that, in
the interpretation of pseudo-colored patches of
slightly-enhanced cortical activity by modern
phrenologists equipped with the powerful tools
of functional MRI [9], parameters such as regional variations in cortical thickness and cell
density, which conceivably bear upon the integrated metabolic output measured locally, are
seldom considered. In other words, it makes
sense, in drawing conclusions about function, to
keep in mind that the lighting up in a neuroimaging experiment of a cortical area with fewer
cells per unit volume takes on a different meaning compared to a signal of a similar intensity
but in an area that is denser in cells or deeper in
thickness; the extensive quantitative data presented can be useful as potential correcting denominators.
Economo briefly relates information on the
function of various cytoarchitectonic areas. Since
the primary goal of the present edition is to preserve the treasure of his accrued cytoarchitectonic data, rather than presenting an updated review
on the function of each cerebral lobe or cortical
area, it would be beyond the scope of the book to
introduce further updates, which can be found in
modern sources.
An example is the insular lobe. We understand today that the insula has multiple roles,
such as in the conscious sense of disgust and nausea and interoception, i.e. the visceral feelings of
ones inner organs or gut feelings [41]; the anterior insula of the right hemisphere may provide a
basis for emotional awareness, i.e. the subjective
image of the material self as a sentient entity [41,
42]. Besides mediating the experience of disgust
through its connections from olfactory and gustatory centers (chemical exteroception) as well as
interoceptive cortical afferents, the anterior insula appears to mediate the perception of disgust
in others via a mirroring mechanism for emotions as well [43, 44].
It looks like greater progress has been made in
recent years with regard to function than structure, with histology faring much better in the test
of time over behavior. The following admission
supports such a thesis: Von Economo published
a book in 1929, in which appear concise detailed
descriptions of the brain including the insula.

Unfortunately, this publication, with comprehensive presentations illustrating the intricate
gyral and sulcal patterns, is a relatively unknown
source of reference. In the intervening decades,
little has transpired to expand our knowledge
surrounding the gross anatomic features of the
insular region or to illustrate the intricacies of its
function [45].
Economo repeatedly mentions the rhinencephalon (olfactory brain) a term popular at
the time and its association with the functions
of the limbic lobe. We now understand that the
mammalian hippocampal complex is critical to
remembering conscious experiences and also
needed for experiential or episodic memory, by
transferring information to longer-term memory, retrieving episodic memories, and participating in spatial navigation [41]. In the primate hippocampus, there appear to be both separate and
combined representations of objects and their
spatial location; the rodent hippocampus is critical to the capacity of integrating what, where
and when information in memory for single experiences, in other words, using combinations of
olfactory and spatial cues to judge the order of
presented events [46]. The entire hippocampal vicinity seems to be implicated in such functions,
further encoding olfaction; that is why parts of
the medial temporal lobe are called rhinal, entorhinal, and perirhinal cortices [47].
Clinically relevant to the rhinencephalon is
the notion that in Alzheimer disease, an association is found between psychophysical odor identification tasks and underlying neuropathologic
changes in areas critical to processing olfactory
information, defined by volumetric MRI measures of mesial temporal areas; thus, correlations
between the volume of the hippocampus and the
parahippocampal gyrus and the performance on
odor tasks even points to the potential of odor
identification tests as diagnostic batteries for an
early diagnosis [48].
It is worth reviewing some of the reviews that the

book had received when it was first published.
The neuropathologist Karl Neubrger (1890
1972) of Mnchen-Schwabing had commented
about the German edition: The work eliminates
the conceded lack of a short modern description
of the basic facts of cortical architectonics; because the well-known magnificent and highly
praised standard work that von Economo published two years ago with Koskinas is accessible
with difficulty to most and, due to its comprehensive detail, it is rather intended for the specialized researcher. In the new overview, one will
find an excellent and rapid orientation to all the
substantial issues of cellular structure a possibility most gratefully welcomed by anyone interested in brain anatomy. After a general introduction, virtually all important areas are discussed with a consideration of functional
questions as well and illustrated through excellent figures. For a future edition one might have
perhaps wished that figure 1 (the cytoarchitectonic area brain map) were accompanied by a
direct explanation of the abbreviated letter symbols ... By studying the work, one becomes again
and again more aware of the numerous interesting problems that a science, apparently as dry as
cytoarchitectonics, can offer [49].
The anatomist Erich Kallius (18671935) of
Heidelberg remarked: After Economo published
the large comprehensive work on the Cytoarchitectonics of the Adult Human Cerebral Cortex and
the Atlas, which unfortunately is not very affordable due to its high price, he wisely and thankfully decided to author this lucid manual, which
of course cannot reach the glory of the larger
work. The illustrations, which are mostly new,1
do not come close to the marvelous large plates of
the Atlas; however, they have the advantage of be-
As a matter of fact, all of the microphotographs depicting

cortical areas in the book are new, with none duplicated from
the Atlas (Editors note).
XI
ing much handier and very sharp, naturally forming a substantial part of the work ... People beyond the [neurology] circles will be very grateful
to the author for putting together this intelligible
overview of the cellular structure of the cerebral
cortex. Further important facts will be discovered, above all by comparative anatomy, opening
up yet another far-reaching sphere of research.
The fact that one can diagnose the sensory and
motor cortices based on structure is a fundamental discovery [50].
The British Journal of Surgery wrote of the
French edition: Although von Economos small
book on the cerebral cortex is more concise than
his large Atlas, the average reader will find in it
all that he wants. The clearness of the description
and the numerous large and excellent illustrations make it very readable, and valuable also as
a work of reference. It deals with the structure of
the cortex not only from an anatomical but also,
as it were, from a physiological point of view ...
One can see that the author has the outlook of a
neurologist who is accustomed to study the results of cerebral disease, rather than that of a pure
anatomist. The book has been brought fully up to
date and can be recommended as the best monograph on its subject [51].
The editors of Brain welcomed the German
and French editions: A very useful book, as it
presents the detailed cell structure of the cerebral
cortex in an easily assimilable form ... By his system of naming each of the cortical areas by a letter-group, the author preserves the gross anatomical nomenclature, and at the same time gives
an indication of the type of cortex present in each
area. This system seems to us a definite improvement on the numerical system of Brodmann ...
The book is beautifully illustrated, and the 46
large photomicrographs of the most distinctive
cortical areas make it a book which will be as useful for reference in the laboratory as for home
study. The French translation by Dr. Ludo van
Bogaert has been well done, and the addition in it
of headlines to the paragraphs in leaded type fa-
XII
cilitates reference. Both volumes are most welcome additions to the neurologists library [37].
On the other hand, the 1929 English translation by a training psychoanalyst rather than a
neuroanatomist, although welcomed, laid itself
open to criticism: We could wish that the translator had not followed the original so exactly in
some respects, as too literal a translation of the
German has made the text either unintelligible
or misleading in many places it is often necessary to refer to the German or the French edition
to arrive at the authors meaning. We could wish
too that, instead of the long chapter headings of
the original, the translator had adopted the useful paragraph headings of the French edition,
which greatly excels the English edition [52].
Moreover, in reviewing Economos subsequent
book Encephalitis Lethargica, the editors of Brain
had remarked: the translators knowledge of
neuropathology has been valuable, as without
such knowledge translations of technicalities are
apt to be incomprehensible, an error which was
not altogether avoided in the English translation
of the same authors book on the cerebral cortex
[53].
The Journal of Anatomy had been kinder: The
present work is a condensation an abstract of
the essentials of Professor von Economos great
book and Atlas. The author has taken the opportunity to introduce modifications and additions
which are new. For English anatomists and medical men generally, the present work will be a
boon, for it presents them with an excellent summary of the present state of knowledge concerning the microscopic structure of the cortical areas of the brain ... The final chapter, the most interesting of all, is devoted to the manifestations of
the living cortex, consciousness being regarded
as the specific sensory energy of the cerebrum. Finally, the natural law of progressive cerebration is
discussed [54].
Morton [55] wrongly mentions in the third
edition of Garrison and Morton an English
translation 1929 of Economo and Koskinas; he
most likely refers to the English edition [4] of

Zellaufbau [1].
Some additional details have been inserted into
the text based on the larger Textband [6], e.g. brief
comments on areas such as FC I, FDm(C), FDm(E),
FGi and FHL in the frontal lobe, TAm in the temporal lobe, and HB3 and HCg in the inferior limbic
(hippocampal) lobe, which are not mentioned by
Economo in any of the versions of Cellular Structure. The newly compiled table 2 presents all the
additional transition and modification areas described by Economo and Koskinas, beyond their
standard 107 cytoarchitectonic modifications
listed in table 1 (also newly compiled for this edition). Likewise, figure 1 has been supplemented
with the cortical maps of the superior and inferior hemispheric surfaces, beyond the lateral and
midsagittal (median) views appearing in the
original. The quantitative information on each
cortical layer for each cytoarchitectonic area (i.e.
layer thickness, cell density and cell size) has been
extracted from the text and segregated into practical tables (6 through 12); certain missing values
were supplemented from the data in the larger
work [6]. Thus, the reader can gain an integral
view of the data by lobe.
Two other departures from the original edition are the introduction of subheadings to facilitate reference following the example set by
van Bogaert in the French edition [2, 37], and the
elimination of the smaller font size originally
used in the detailed layer descriptions [14]; instead, those sections are now presented in fullsize text, with the corresponding layer at the beginning of each paragraph highlighted in bold
characters (a similar concept was applied to the
layer descriptions in the larger textbook).
The Appendix presents the English world
premire of a recently discovered manuscript of
Koskinas [34], privately published in Athens in
1931, in which Economos collaborator presents an
insightful analysis of the General Part of the larger Textband [6], thus forming an inextricable sup-
plement to Cellular Structure. The Outline has

been enhanced by the inclusion of the original illustrations [6] that Koskinas refers to (fig. 7388).
While the previous German [1] and English
[4] editions of Cellular Structure contained the
same references, the French [2] and Italian [3]
versions incorporated further articles from the
respective regional literature. The comprehensive bibliography section of 307 references at the
end of this volume combines all the 80 references
found in the four European editions [14], all the
references cited in the larger Textband [6], plus
additional entries related to points made in the
text as well as Economos sequel studies, with full
citation information thoroughly compiled over
the past four years. Moreover, for all citations in
the text to the larger work [6], inclusive pagination has been filled in. Cross-references to the
corresponding plates of the new Atlas [5] are given for each cytoarchitectonic area, as well as EK
(Economo-Koskinas) modification area coding
numbers (1 through 107) [35].
Many of the concepts that Economo and
Koskinas communed about remain at the core of
neuroscientific inquiry. Fundamental gender differences seem to exist in the cytoarchitecture of
the human neocortex regarding neuronal density,
neuropil volume, and laterality [56]. Microscopically defined borders usually differ from gross
anatomic landmarks, cytoarchitectonics reflecting the inner organization of cortical areas and
their morphofunctional correlates [57, 58]. Despite the integration of multifactorial descriptors
such as chemoarchitecture, angioarchitecture,
neurotransmitter, receptor and gene expression
patterns, as well as white matter tracts, it is clear
that the knowledge of the classical and tedious
to learn anatomy remains fundamental [59].
The structure of cortical layers incorporates, and
reflects, the form of their constitutive cells and
their functional connections; the underpinnings
of neuronal connectivity at the microscopic level
are paramount to interpreting any clues afforded
by neuroimaging pertinent to cognition.
XIII
Nothing may possibly conclude a discussion

on the human cerebral cortex better than the
crystallized thought of neuroanatomist Gerhardt
von Bonin: The cortex is both chaos and order,
and therein lies its strength ... Even if understanding is held to be subservient to doing, it is
still crucial to understand understanding ... Yet
the study of individuals, and an insight into the
structure of the organ of their minds [... one of
the most difficult and most fascinating of scientific problems ...] will lead to a better understanding of their abilities and their limitations, of the
ways in which they can build a society and thus
will indirectly help us to define that type of order
which is the good life [33].
The courtesy of numerous individuals is gratefully acknowledged, namely, staff members at the
Province Library in Foggia, Central Medical Library in Trieste, Marani Medical Center in Verona, Tiraboschi Library in Bergamo, Central
National and Anthropology Libraries in Florence, Mondino Central Library in Pavia, Cencelli Science Library in Rome, Human Physiology
Library in Bologna, Librairie Harteveld in Fribourg, Public and Medical Libraries of the University of Basel, Berlin State Library, New York
State Library, Library of Congress, National Library of Medicine, Lilly Medical Library of Indiana University, Neurology Department of Aiginiteion Hospital, Veterinary Anatomical Library
of Aristotelian University, National Library of
Greece, and the numerous German and Austrian
Libraries associated with the Subito Service. I
thank the Medical History Department of the
University of Vienna for the authors portrait in
the front matter, and Nikolaus Reiner of Carl
Reiner Chirurgische Instrumente in Vienna for
the photographs of figure 62.
Constantin von Economo was the first ever
recipient in 1928 of the Science Prize awarded
by the Academy of Athens for his audacious
studies on encephalitis lethargica, the dreadful
disease that he first defined anatomically and ex-
XIV
perimentally, and for his work on the cytoarchitectonics of the cerebral cortex, which truly opens
up new paths for studying localization in the
brain and gives an impetus to the elucidation of
its area variations according to structural types
[60]. Appropriately, it is a true joy and an honor
that the Academy has embraced the present endeavour under its auspices. Additional research
grants awarded by the Bodossakis Foundation
and the University of Macedonia, Greece, have
made this publication possible, and are dutifully
acknowledged, as is the support of the Hellenic
Neurological Society.
I thank Springer Rights and Permissions in
Heidelberg and in Vienna, Dpartement Livres
Elsevier-Masson in Issy-les-Moulineaux, Academic Permissions of Oxford University Press,
and Cappelli Editore in Bologna, for permission
to use previously published material.
I express my appreciation to Dr. h.c. Thomas
Karger for preserving the tradition and fine craft
of biomedical publishing and for a magnificent
collaboration; to S. Karger AG for the uncompromising quality standards in producing the Cellular Structure of the Human Cerebral Cortex in
conjunction with the Atlas of Cytoarchitectonics
of the Adult Human Cerebral Cortex, especially,
the staff members that I had the pleasure to cooperate with, as well as the additional staff who
once again contributed their service behind the
scenes and thus enabled me to bring this opus
into effect.
Lazaros C. Triarhou
Thessaloniki, September 2008
References
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2 Economo C von: Larchitecture cellulaire normale de lcorce crbrale
(translated and edited by L. van Bogaert). Paris, Masson et Cie, 1927.
3 Economo C: La citoarchitettonica della
corteccia cerebrale umana (translated
by C. Enderle, edited by G. Mingazzini).
Bologna, L. Cappelli, 1928.
4 Economo C von: The Cytoarchitectonics of the Human Cerebral Cortex
(translated by S. Parker). London,
Humphrey Milford-Oxford University
Press, 1929.
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Karger, 2008.
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Vorwort
Authors Foreword to the 1927 German Edition
Der Wunsch nach einem handlichen und bersichtlichen Leitfaden zum Studium der Cytoarchitektonik der Grosshirnrinde des Menschen
wurde mir von Psychiatern und insbesondere
von Hrern meiner Semestralvorlesungen ber
dieses Thema in letzter Zeit vielfach ausgesprochen.
Obschon ich nun vor kaum zwei Jahren ein
umfangreiches und ziemlich alles bis dahin Bekannte umfassendes Werk samt Atlas darber
herausgegeben habe, komme ich diesem berechtigten Verlangen nach einem kurzen Lehrbuche
gerne nach. Ich tue dies in der einfachsten Form,
der Verffentlichung des Textes dieser Vorlesungen, die ich seit dem Jahre 1923 in zweistndigem College ankndige. Obschon also diese
Vorlesungen keineswegs als ein blosser Auszug
des grossen Werkes zu werten sind, sondern mir
eher die Grundlage bei der Abfassung des Textes
desselben abgegeben haben und jetzt durch die
neuesten Forschungsergebnisse wieder vervollstndigt sind, so knnen sie das Studium des
Hauptwerkes und seines reichen Atlas durchaus
nicht ersetzen, denn eine erschpfende Darstellung der so komplizierten Architektur der Hirnrinde lsst sich in zehn Vorlesungen nicht geben.
Wohl aber knnen dieselben auch dem grndlichen Forscher die Handhabung des grossen
Werkes bedeutend erleichtern, indem sie ihm
vorerst einen raschen allgemeinen berblick ermglichen, worauf er sich im grossen Werke in
jenen Kapiteln, die sein spezielles Interesse beanspruchen, unmittelbar zurecht finden wird.
Der Hauptzweck dieses kleinen Buches ist es
jedoch fr Studierende und auch fr selbstarbeitende Untersucher, die notwendigsten Grundtatsachen des Zellaufbaues des Cortex in handlicher Form zusammenzufassen. Dementsprechend habe ich es vorgezogen, die 46 durchaus
neuen Photographien der wichtigsten Rindenfelder diesmal nicht als eigenen Atlas, sondern
gemeinsam mit den brigen Abbildungen und
Schemen direkt im Text erscheinen zu lassen. Bei
Darstellung der Photographien hat diesmal Dr.
Horn in dankenswertester Weise mitgearbeitet;
besten Dank sage ich auch nun wieder der Laborantin unserer psychiatrischen Klinik Frl. Strasky fr ihre Mhewaltung bei der Prparierung
des Materials. Auch hier hat akademischer Maler
Bruno Keilitz die von mir entworfenen Zeichnungen sorgfltigst und tadellos ausgefhrt.
Wenn die Lektre dieses kleinen Lehrbuches
ausser der Aufklrung, die es dem Studierenden
geben soll, auch noch den einen oder den andern
zu selbstndiger weiterer Erforschung des Neulandes der Architektonik der Rinde anregen
sollte, so wre alles erreicht, was man von einem
so kurzen Leitfaden erhoffen darf.
Wien-Gerasdorf, im Mrz 1927
XVII
Prface
Authors Preface to the 1927 French Edition
Ces derniers temps, divers psychiatres et auditeurs des mes confrences semestrielles sur
lArchitectonie cellulaire de lEcorce humaine
mavaient, maintes reprises, exprim le dsir de
possder un manuel pratique et gnral de cette
partie de lAnatomie crbrale.
Malgr que jaie publi, il y a deux ans peine,
un vaste Atlas runissant tout ce qui est actuellement connu dans ce domaine, cest avec plaisir
que je rponds leur dsir trs justifi, en publiant ce petit livre, dans sa forme la plus simple,
celle mme du texte de mes leons depuis 1923.
Ces confrences ne sont pas un simple extrait
du grand travail, elles en constituent plutt la
base, mais taye de nouvelles acquisitions. Leur
lecture ne peut donc en aucune manire remplacer ltude du travail principal, et lexpos dtaill de larchitecture crbrale si complexe dpasse
videment le cadre de ces quelques chapitres. A
ceux qui voudraient tudier ce sujet dune manire
XVIII
plus approfondie, ils fournissent cependant un

aperu densemble leur permettant de retrouver
rapidement, dans le grand Atlas, les chapitres qui
les intressent spcialement, leur facilitant ainsi
lusage du travail complet.
Le but de ce petit livre est donc avant tout de
fournir aux tudiants et aux chercheurs, sous une
forme aisment accessible, les notions indispensables la connaissance de larchitectonie cellulaire corticale. Aussi ai-je prfr insrer dans le
texte mme, avec les schmas et les autres dessins,
les 46 photographies nouvelles des principaux
champs corticaux, plutt que de les runir sparment sous forme dAtlas.
Si la lecture de ce livre, aprs avoir instruit les
tudiants, pouvait inciter lun ou lautre dentre
eux faire des recherches personnelles dans cette
terre nouvelle quest lArchitectonie corticale,
jaurais atteint largement ce quon est en droit
dattendre dun guide aussi abrg que celui-ci.
Prefazione
Authors Preface to the 1928 Italian Edition
Numerosi medici e numerosissimi studenti, e pi

specialmente quelli che hanno seguito i miei corsi semestrali, mi hanno espresso il desiderio di
avere una guida succinta e chiara allo studio della citoarchitettonica della corteccia cerebrale
delluomo. Quantunque io abbia gi due anni fa
pubblicato un voluminoso trattato che contiene
pressoch tutto quanto si conosce oggi sullargomento, pure, seguendo i molti inviti ricevuti, mi
piaciuto porre mano a questo breve manuale. A
tale scopo ho raccolto in volume il testo delle
conferenze che tengo fin dallanno 1923 e che
non rappresentano, quindi, un riassunto della
mia opera maggiore del 1925, ma piuttosto mi
hanno servito di base per il suo testo e sono state
aggiornate con i risultati delle ricerche pi recenti. Il presente volume non pu sostituire n il
testo n latlante dellopera maggiore per chi
voglia approfondire i problemi della citoarchitettonica poich non possibile ridurre in soli 10
capitoli una completa descrizione della tanto
complicata struttura della corteccia cerebrale
per pu tornare utile al ricercatore che voglia

studiare il maggior volume, dando un quadro
dinsieme dellargomento e permettendo quindi
una migliore e pi pronta comprensione degli
speciali capitoli, argomento del suo studio pi
profondo.
Pi che ad ogni altro, per, questo volume
destinato allo studente ed al medico che ami il
lavoro indipendente a questi porge raccolte in
breve volgere di pagine le nozioni fondamentali
relative alla citoarchitettonica della corteccia cerebrale.
Io ho preferito non raccogliere in un atlante le
46 nuove figure fotografiche che illustrano questo volume, ma intercalarle nel testo insieme con
le altre figure schematiche.
Considerer raggiunto lo scopo di questo libro se, oltre allavere servito come mezzo di cultura agli studiosi, varr a spingere qualcheduno
di essi a tentare, per proprio conto, lesplorazione
dellarchitettonica della corteccia cerebrale, tanto
ricca di problemi, tanto ricca di soddisfazioni.
Costantino Economo
Trieste, 1926
XIX
Preface
Authors Preface to the 1929 English Edition
The desire for a handy and concise introduction

to the study of the cytoarchitectonics of the human cerebrum has been expressed to me with
great frequency of late, not only by psychiatrists
but also by anatomists and physiologists who
have attended my University lectures on this subject. Although only 3 years have gone by since I
published a voluminous treatise on the subject,
which contained not only the fruit of my own
work, but also all that had been done before, I feel
that there is a place for such a short textbook as
this. The arrangement of the material follows
rather closely that which I have adopted in the
course of my lectures. The content of these lectures, as they appear in this little volume, is not
merely an abstract of the large work published in
1925; for the lectures were, on the contrary, the
basis for this book. But, although the present volume contains a few additions based on more recent research, it can in no way replace a careful
use of the larger work. It is manifestly impossible
to exhaust the complex architecture of the cortex
in so short a space as a dozen chapters. The present book is, however, intended to serve even the
specializing scientist as an aid to the more effi-
XX
cient use of the larger work. The general view and

the quick survey afforded by the present book
should lead to a rapid orientation in regard to
those chapters of the large work which attract
special attention. Its chief purpose, however, is to
bring the basic facts of the cell structure of the
human cortex within the reach of the student and
independent worker in as easily intelligible a
form as possible. It is for this reason that I have
chosen to use the 46 new photographs of the chief
cortical areas, not as a separate atlas but as illustrations in the text.
If the study of this little textbook should provoke one or another of its readers to an independent journey into the new world of cytoarchitectonics, in addition to the enlightenment which it
offers to students, then all the hopes I cherish for
it will have been realized.
It is my particular hope that the Englishspeaking public will find this book of advantage,
especially as a number of devoted English workers I mention only the names of Elliot Smith,
Campbell, Bolton were pioneers in this province, and very early called the attention of the scientific world to its importance.
Vienna
Introduction
Historical Remarks
The cerebral hemispheres are clad in a gray cortex, which abounds in nerve cells. With the description by Vicq dAzyr [1786], in the region of
the calcarine sulcus within the unusually thin
occipital cortex, of the characteristic white stria
of myelinated fibers that bears his name, it became apparent that the cerebral cortex is not
uniform in structure; rather, it shows regional
variations in both its structural form and thickness.
From the studies of Baillarger [1840, 1853] onwards, we distinguish six horizontally superimposed layers in the cerebral cortex. The regional
variations of the cerebral cortex in cellular composition and layer thickness often produce striking structural differences.
The Viennese psychiatrist Meynert [1867/1868,
1868, 1872a, b] placed emphasis on such differences in the 1860s. Thence, he is considered as the
true founder of the modern field of cytoarchitectonics of the cerebral cortex. Based on the multiplicity of structure in various parts of the cerebral
cortical surface, Meynert concluded, long before
Fritsch and Hitzig [1870], that the cortex contains, so to speak, many different organs, which,
depending on their particular structure, can have
specific functions. Meynert realized that the exact study of cortical areas would give birth to a
new cerebral organology founded on a firm anatomic basis, in contradistinction to the organology of Gall [18221825] that was fashionable at
the time.
Meynert became particularly interested in the
cortex of the calcarine sulcus; he was able to recognize in it eight layers, in contrast to the rest of
the cerebral cortex. The augmented granular layers and their high cellularity, which reminds one
of the structure of the retina, led Meynert to attribute an analogous sensory receptive function to
this part of the cortex, a concept no longer disputed.
In 1874, Betz [1874] described the giant pyramidal cells of the precentral gyrus that are named
after him; in agreement with Fritsch and Hitzig
[1870], he showed that these cells, both in humans and in animals, fall exclusively within a
primary zone that is excitable by direct electromotor stimuli. Betz [1881] carried out further
studies on the cellular structure of the entire cerebral cortex and confirmed the precepts of
Meynert regarding the division of the cortex into
multiple fields of varied structure.
Following Betz [1874, 1881], Lewis [1879, 1880,
1882], Lewis and Clarke [1878] and particularly
Hammarberg [1895], these studies were continued by Campbell [1903, 1905], Elliot Smith [1904b,
1904c, 1907], Brodmann [1903a, 1903b, 1905a,
1905b, 1906, 1908ac, 1909, 1914], Vogt [1902,
Fig. 1. a, b Cytoarchitectonic area maps of the human cerebral hemispheres showing a the lateral (convex) and b the
median (midsagittal) hemispheric facies. The lateral (Sylvian) fissure is laid open, with the frontal and parietal opercula lifted in the lateral view, and the hippocampal and callosal sulci laid open in the median view, to make deeper
lying areas, such as the insular region, visible. Drawings based on the corresponding maps of Economo and Koskinas
[1925], incorporating some later research results.
1903, 1906, 1910, 1911, 1913], Vogt and Vogt

[1902, 1910, 1919], Marinesco [1902, 1909, 1910a
c, 1911], Marinesco and Goldstein [1910, 1911,
1927], Marinesco and Mironesco [1910], and
Economo and Koskinas [1925]. Campbell [1903,
1905] and Elliot Smith [1904c, 1907] were the
first authors to produce maps of cortical area divisions.
Partition Scheme of the Cerebral Cortical

Surface into Areas
The first two discoveries mentioned namely,

that of Meynert on the specific structure of the
sensory visual cortex in the calcarine sulcus, and
that of Betz on the specific structure of the motor
zone with its giant cells in the precentral gyrus
have remained the foundations from which our
entire subsequent knowledge on cortical cytoarchitectonic areas has emerged.

The cortical maps of figure 1 represent our
newest division of the cortex into areas, based on
the larger work on the cytoarchitectonics of the
cerebral cortex [Economo and Koskinas, 1925],
and incorporating slight changes based on more
recent data. We have up to now been able to identify 109 different structural areas (tables 1, 2), the
details of which I shall return to later.1
The Six Cortical Layers
As already mentioned, the cerebral cortex consists of six superimposed layers, each of which is
usually named after its predominant cell type (cf.
fig. 2, which shows the transition from the dome
of a gyrus on the right side to its wall on the left;
for a comparison of the various terms adopted for
each layer at different periods, see table 5 in the
Atlas [Economo and Koskinas, 2008]).
The 107 modification areas defined by Economo and Koskinas [1925, 2008], plus areas HE 4 and HE 5 in the hippocampal
gyrus (cf. corresponding chapter).
Introduction
Fig. 1. c, d Cytoarchitectonic area maps of the human cerebral hemispheres showing c the dorsal (superior) and d
the ventral (inferior or basal) hemispheric facies. In the view of the ventral surface, the temporal pole is left intact on
the hemisphere shown on the left side of the drawing, while it has been removed on the right side, to expose the
orbital and insular regions. R, central sulcus of Rolando. Schemes added from Economo and Koskinas [1925, 2008].
Introduction
Table 1. The basic parcellation scheme of Economo and Koskinas [1925, pp. 218220] for the human cerebral cortex
consists of 54 ground areas, organized into 76 variants and 107 modifications (cf. tables 4, 6 and 7 in the Atlas [Economo and Koskinas, 2008] for classification details, Latin area names, and Brodmann area correlations). The EconomoKoskinas (EK) area numbers are also mentioned in the discussion of each individual cytoarchitectonic area in the text.
The sequence of the lobes in this table (FLIPOTH) has been kept in accordance with the order they originally
appear in the larger textbook [Economo and Koskinas, 1925] and the Atlas [Economo and Koskinas, 2008], whereas
in the series of lectures that form the matter of the present volume, the superior limbic lobe is discussed just before
the hippocampus, and the insula immediately following the parietal lobe (FPIOTLH).
EK area number
Area symbol
Area name
Frontal lobe (F)

Prerolandic region
1
2
3
4
5
6
7
8
9
10
FA
FA
FAop
FB
FBop
FC
FC L
FCBm
FC I
FCop
precentral area
giant pyramidal precentral area
opercular precentral area
agranular frontal area
opercular agranular frontal area
intermediate frontal area
limbic intermediate frontal area
magnocellular agranular intermediate frontal (Brocas) area
intermediate frontal area at beginning of insula
opercular intermediate frontal area
Anterior frontal (prefrontal) region

11
FD
12
FDm
13
FDp
14
FDop
15
FDL
16
FD
17
FD
18
FE
19
FE L
granular frontal area

magnocellular granular frontal area
parvicellular granular frontal area
opercular granular frontal area
limbic granular frontal area
middle granular frontal area
triangular granular frontal area
frontopolar area
limbic frontopolar area
Orbital (orbitomedial) region

20
FF
21
FF
22
FF
23
FG
24
FGi
25
FH
26
FHL
27
FH L
28
FJ
29
FK
30
FL1
31
FL2
32
FL3
33
FM
34
FMt
35
FN
granular orbital area

agranular orbital area
pretriangular orbital area
area of straight gyrus (area recta)
internal area of straight gyrus
prefrontal area
parolfactory prefrontal area
limbic prefrontal area
frontoinsular area
frontal piriform area
primary parolfactory area
secondary parolfactory area
tertiary parolfactory area
geniculate area
geniculate area of olfactory triangle
precommissural area
Table 1 (continued)
EK area number
Area symbol
Area name
Superior limbic lobe (L)

Anterior superior limbic region
36
LA1
37
LA2
38
LA3
39
LB1
40
LB2
precingulate agranular anterior limbic area

anterior cingulate agranular anterior limbic area
cingulate agranular anterior limbic area limitans
anterior ultracingulate area
area of indusium griseum
Posterior superior limbic region

41
LC1
42
LC2
43
LC3
dorsal posterior cingulate area

ventral posterior cingulate area
posterior cingulate area limitans
Retrosplenial subregion
44
45
46
47
48
LD
LE1
LE2
LF1
LF2
agranular retrosplenial area

superior retrosplenial area granulosa
inferior retrosplenial area granulosa
posterior ultracingulate area
ultracingulate area obtecta
Insular lobe (I)

49
50
51
52
53
54
IA1
IA2
IB
IBT
IC
ID
dorsal precentral insular area

ventral precentral insular area
postcentral insular area
postcentral insular area at temporal entrance
orbito-insular area
piriform insular area
Parietal lobe (P)

Postcentral (anterior parietal) region
55
PA1
56
PA2
57
PB2
58
PB1
59
PC
60
PD
giant pyramidal postcentral area

giant pyramidal postparacentral area
oral postcentral area simplex
oral postcentral area granulosa
intermediate postcentral area
caudal postcentral area
Superior parietal region

61
62
63
64
PE(D)
PEm
PEp
PE
superior parietal area (transition postcentral)

magnocellular superior parietal area
parvicellular superior parietal area
giant pyramidal posterior superior parietal area
Inferior parietal region

65
66
67
68
69
PF
PFt
PFop
PFcm
PG
supramarginal area
tenuicortical supramarginal area
opercular supramarginal area
magnocellular (posterior) supramarginal area
angular area
Introduction
Table 1 (continued)
EK area number
Area symbol
Area name
Basal parietal region

70
71
72
PH P
PH T
PH O
basal (temporooccipital) parietal area at parietal entrance

basal (temporooccipital) parietal area at temporal entrance
basal (temporooccipital) parietal area at occipital entrance
Occipital lobe (O)

73
74
75
76
77
78
79
OA2
OA1
OAm
OB
OB
OB
OC
anterior peristriate area

posterior peristriate area
magnocellular peristriate area
parastriate area
giant pyramidal parastriate boundary of parastriate area
maculae granulosae of parastriate area
striate area (granulosa)
Temporal lobe (T)

Supratemporal region
80
81
82
83
84
TA1
TA2
TB
TC
TD
posterior superior temporal area

anterior superior temporal area
magnocellular supratemporal area simplex
supratemporal area granulosa
intercalated supratemporal area
Temporal region proper

85
86
TE1
TE2
middle temporal area proper

inferior temporal area proper
Fusiform region
87
88
89
TF
TH
TH
fusiform area
hippocampotemporal area
agranular hippocampotemporal area
Temporopolar region
90
91
92
93
TG
TG
TJ
TK
temporopolar area
agranular temporopolar area
temporal piriform area
area of substantia perforata
Hippocampal (inferior limbic) lobe (H)

94
HA1
95
HA2
96
HA3
97
HB1
98
HB2
99
HC
100
HD1
101
HD2
102
HD3
103
HE1
104
HE1
105
HE2
106
HE3
107
HF
primary uncinate area

secondary uncinate area
tertiary uncinate area
primary parauncinate area
secondary parauncinate area
rhinal area limitans
presubicular area granulosa limitans
middle presubicular area granulosa
glomerular presubicular area granulosa
subicular glomerular pyramidal area
subicular pyramidal area simplex
pyramidal area of Ammons horn
pyramidal area of digitate gyrus of uncus
dentate area
Layer I is called the molecular layer. It is mostly composed of cortical gray matter, in which the
arbors of numerous dendrites and the terminal
axons form an intricate plexus hence termed
plexiform layer by Campbell [1903, 1905]. Cells of
this layer are few in number, they acquire a piriform or fusiform shape, and they are disposed
tangentially; they are called Cajal cells (also CajalRetzius cells), with a size of 46 m. The other
cell nuclei visible in layer I in figure 2 belong to
glia and vascular endothelial cells.
Layer II is the external granular layer. It is
composed of numerous small granule cells,
densely packed, with a round, polygonal or triangular shape.
Layer III immediately beneath layer II is called
pyramidal cell layer. It is thick and contains pyramidal cells which are fewer in number, but robust
and larger in size.
Layer IV or internal granular layer follows; it
is also composed of densely packed, extremely
small, mostly round or polymorphous granule
cells.
Layer V is called internal pyramidal layer or
ganglionic layer. It has a lighter appearance than
layer IV, and it is largely composed of fine, pyramidal neurons, although these hardly reach the
size of the pyramidal cells of layer III.
Layer VI is called spindle (fusiform) cell layer
and lies beneath layer V. It is composed of somewhat densely collected spindle-shaped (fusiform)
cells, with their long axis arranged perpendicularly to the cortical surface. It borders directly on
the white matter of gyri. Thick fiber bundles radiate from that white matter into layer VI; in
preparations stained for myelin, these are easily
traced as far up as layer III.
Subdivision of Layers into Sublayers

On a closer examination, the six individual cortical layers described do not exhibit a uniform
structure throughout their extent.
Introduction
Layer III for example, contains pyramidal

cells that gradually become larger, from the cortical surface towards the white matter; the largest
examples of such cells are found close to the border with layer IV (fig. 2). Thus, layer III is subdivided into an upper sublayer IIIa with small pyramidal cells, a middle-positioned sublayer IIIb
with medium-sized pyramidal cells, and a lower
sublayer IIIc with the largest pyramidal cells.
Layer V is also subdivided into two sublayers,
an upper Va and a lower Vb. However, such a division is based less on differences in cell size and
more on their packing density.
Layer VI is regularly subdivided into a denser,
upper sublayer VIa composed of larger cells, and
a lower, less compact sublayer VIb with fewer and
smaller cells, also being traversed by incoming
fiber bundles from the white matter (fig. 2).
Such divisions result in ten cortical layers (decalaminar structure) and show that the distinction of six layers can be both arbitrary and conventional. Nevertheless, on practical grounds, we
retain the six-layer division, especially since it has
received more support than any other scheme,
and also, because these layers can be clearly demarcated in histologic sections based on the cell
types that they contain.
Relation of Cytoarchitectonics to
Myeloarchitectonics
The six layers can be distinguished even in preparations stained with the Weigert myelin method,
based on the course of the horizontal, intracortical
myelinated fibers. The study of such fiber structures of the cerebral cortex, called myeloarchitectonics of the cerebral cortex, is not the subject of this
book. I nonetheless reproduce a scheme (fig. 3) that
shows the relation of cytoarchitectonic to myeloarchitectonic layers. Results from myeloarchitectonic
studies led to a mapping of the cortex into numerous fields, which to a great degree overlap with our
cytoarchitectonic fields depicted in figure 1.
Table 2. Additional cytoarchitectonic variations and designations mentioned by Economo and Koskinas [1925],
Economo [1927c, 1927d, 1928e, 1929d], and Economo and Horn [1930] in the human cerebral cortex beyond the original 107 modifications contained in table 1, bringing the total to over 170 cytoarchitectonically defined areas. Two
corsiva capitals after a lobe initial, e.g. PFD, denote transitional types of cortex between two adjacent cytoarchitectonic ground areas; a corsiva capital in parentheses, e.g. FC(B), denotes an admixture of structural elements from that
specific neighboring area. Area TA m was described by Economo and Koskinas [1925]; the four TJH transition modifications by Economo [1929d]; all the other temporal areas below by Economo and Horn [1930]. Areas HD a and HD p
adopted by Economo [1927d, 1928e] from Rose [1926b]; areas HE 4 and HE 5 described by Economo [1927c, d].
Area symbol
Area name
FA(B)
FBA
FBC
FB(C)
FBI
FB(C)op
FBCop
FC(B)
FC(D)
FC(D)op
FCDop
FDC
FDCop
FDm(C)
FDE
FDm(E)
FEF
FEm
FEDm
FFE
FH
LA(C)
IAB
PA
PC
PDE
PDF
PFD
PFt(op)
PFc
PFm
OA2(m)
TA2
TA2
TAm
TAG
TA2G
TBA1
TBA2
TBmp
TBlp
TBma
TBla
precentral area at beginning of agranular frontal area

agranular frontal area, precentral transition
agranular frontal area, intermediate frontal transition
agranular frontal area at beginning of intermediate frontal area
agranular frontal area at beginning of insula
agranular frontal area, (intermediate) in operculum
opercular agranular intermediate frontal area
intermediate frontal area (agranular transition)
intermediate frontal area at beginning of granular frontal area
opercular intermediate frontal area at beginning of granular frontal area
opercular intermediate granular frontal area
granular frontal area, intermediate frontal transition
opercular granular intermediate frontal area
magnocellular granular frontal area at beginning of intermediate frontal area
granular frontal area, frontopolar transition
magnocellular granular frontal area at beginning of frontopolar area
frontopolar area, orbital transition
magnocellular frontopolar area
magnocellular frontopolar area, granular frontal transition
orbital area, frontopolar transition
agranular prefrontal area
agranular anterior limbic area (posterior cingulate transition)
precentral insular area (postcentral transition)
giant pyramidal postcentral area
giant pyramidal intermediate postcentral area
caudal postcentral area, superior parietal transition
caudal postcentral area, supramarginal transition
supramarginal area, caudal postcentral transition
tenuicortical opercular supramarginal area
supramarginal area columnata
magnocellular supramarginal area
magnocellular anterior peristriate area
marginal superior temporal area
superior temporal area in supratemporal gyrus
magnocellular superior temporal area
superior temporal area, temporopolar transition
polar marginal superior temporal area
magnocellular supratemporal area simplex, posterosuperior transition
magnocellular supratemporal area simplex, anterosuperior transition
posteromedial magnocellular supratemporal area simplex
posterolateral magnocellular supratemporal area simplex
anteromedial magnocellular supratemporal area simplex
anterolateral magnocellular supratemporal area simplex
10
Table 2 (continued)
Area symbol
Area name
TBC
TC1
TC2
TD1
TD2
TGA
TG1A
TG1
TG2
TG2
TG2
TJ H1
TJ H2
TJ H3
TJ H4
HB3
HC1
HCg
HC(D)
HDa
HDp
HE4
HE5
magnocellular supratemporal area simplex, granulous transition

transverse supratemporal area granulosa
transverse supratemporal area simplex
intercalated supratemporal area granulosa
intercalated supratemporal area simplex
temporopolar area, supratemporal transition
parainsular temporopolar area, supratemporal transition
parainsular temporopolar area
temporopolar marginal area
temporopolar marginal area, lateral field
temporopolar marginal area, medial field
primary hippocampotemporal (periamygdalar) piriform transition area
secondary hippocampotemporal (periamygdalar) piriform transition area
tertiary hippocampotemporal (periamygdalar) piriform transition area
quaternary hippocampotemporal (periamygdalar) piriform transition area
tertiary parauncinate area
primary rhinal area limitans
granular rhinal area limitans
rhinal area limitans (presubicular granulous transition)
presubicular area granulosa pars anterioris
presubicular area granulosa pars posterioris
quaternary pyramidal area
quinary pyramidal area
The Three Main Cell Types
We saw, from the exposition above, that there are

three main groups of cells that make up the cerebral cortex: pyramidal, granule, and spindle (fusiform) cells (fig. 2, 3).
Pyramidal cells are prominent in layers III and
V; they have a triangular shape, being elongated
in the vertical direction. They emit a long dendrite from their upper pole, seen clearly in silverimpregnated specimens, which reaches as far up
as the molecular layer. On the other hand, dendrites issued from the base of the soma are oriented laterally and downwards, and terminate
within the immediate cortical vicinity at a short
distance from the soma. The axon is also issued
from the base of the soma, and usually invades
Introduction
the white matter of the gyrus, after giving off a

few horizontal collaterals. Pyramidal cells usually contain well-developed Nissl bodies, and a
relatively large and vesicular nucleus with a pronounced nucleolus. Their size may vary substantially; that is why we classify them according to
their height-to-width (H/W) ratio (Schlankheit or
cell slenderness) as small pyramidal cells (size
12/10 m), medium-sized pyramidal cells (about
25/15 m), and large pyramidal cells (about 30
45/1520 m). We classify the even larger pyramidal cells, which reach 50100/2560 m
(H/W), as giant cells.
Granule cells mainly populate layers II and IV.
They are round or polygonal, often triangular,
with a very scanty cytoplasm, as the nucleus occupies most of the cell. Conventional staining
11
Fig. 2. Section of the dome of a gyrus from the frontal lobe, showing the normal six-layered (hexalaminar) cortex.
The white matter (Mark in German), which is devoid of nerve cells, is seen on the lower-right hand corner. IVI, the
six superimposed cortical cell layers. !50.
12
Fig. 3. Comparison of the cytoarchitectonic

layers of cells (left) with the myeloarchitectonic arrangement of fibers (right) in the
human cerebral cortex. The left field, after
Brodmann [1909], shows the cellular structure in a Nissl-stained preparation. Cell layers: I, molecular or plexiform layer; II, external granular layer; III, pyramidal cell layer; IV,
internal granular layer; V, internal pyramidal
or ganglionic layer; VI, spindle or fusiform
cell layer. The right field, after Vogt [1906],
shows myelinated fiber pathways in a preparation stained with the Weigert method.
Myelin layers: 1, tangential layer or plexus of
Retzius; 2, dysfibrous layer; 3, suprastriate or
outer main layer (stripe of Kaes-Bekhterew);
4, external (outer) band of Baillarger (stria of
Gennari or Vicq dAzyr in the visual cortex);
5, interstriate layer; 5b, internal (inner) band
of Baillarger; 6, infrastriate layer. The outer
(external) and inner (internal) zones on the
left refer to the main zones of Kaes [1907]
and Mott [1907] or the fundamental layers
of Jakob and Onelli [1911].
methods do not generally reveal a distinct nucleolus. Their size varies from 4/4 to 8/10 m
(H/W). As a rule, one does not see any dendrites;
only with Cajals silver impregnation can one obtain evidence for short, often bushy (fasciculated)
processes, and also an axon that splits within the
cortex, not far from the soma.
Fusiform cells of layer VI are long and spindleshaped; their ovoid, vesicular and centrally placed
nucleus contains a readily identifiable round nucleolus. Nissl bodies are generally accumulated
towards the two poles of the cell. Both poles of the
spindle transform into long dendrites; according
to Ramn y Cajal [19001906, 1921, 1923], the
upper dendrite reaches as far up as the molecular
layer, whereas the lower dendrite passes downwards and appears to split while still within layer
Introduction
VI. The axon stems from the middle of the soma

or from its lower end, and goes into the underlying white matter of the gyrus. The size of spindle
cells varies between 30/15 and 15/10 m (H/W).
At times, they can also assume a more triangular
shape.
Special Cells
Besides these three fundamental cell types, which

constitute the main cellular mass of the cerebral
cortex, one also finds certain other cell types.
In the molecular layer, we consistently find
small horizontal and piriform cells of Cajal typical of layer I; their long axons stretch over great
distances parallel to the surface of the layer, where
13
Fig. 4. Two giant pyramidal cells of Betz in the precentral

gyrus. !250.
In layer IV of the visual cortex, and particularly in the so-called stria of Gennari [1782] or
Vicq dAzyr [1786], one also finds very large, flat,
horizontally-disposed stellate cells, called giant
stellate cells.
Finally, in the rhinencephalon (fig. 5) one also
finds characteristic cell types with a very peculiar
morphology, such as the tufted pampiniform cells
of Cajal, and our corkscrew or rod cells in the limbic gyrus and the transverse insular gyrus
[Economo and Koskinas, 1925; Economo, 1926d].
These cells were described as spindle cells of layer V and also reproduced in part by Hammarberg
[1895], Flechsig [1897, 1920], Ramn y Cajal
[19001906, 1921, 1923], Nikitin [1909] and Marinesco [1910a, 1910c].
These different cellular types, characteristic
of specific cortical regions, are called special cells
of the cerebral cortex. However, I repeat that for
the major part, the cortex is composed of the
three main types mentioned above: pyramidal,
granule, and spindle (fusiform) cells.
Cellular Density
they also split into terminal branches. Such axons

form a large part of the so-called tangential fibers.
I already described the giant cells of Betz in the
motor cortex. Most often they are pyramidal in
shape, thicker and coarser, reaching dimensions
of 60120/3080 m (H/W); they have an exceptionally large nucleus, a large nucleolus, a lipid
inclusion and pronounced tigroid (Nissl) bodies.
Their basal dendrites are strangely knotty and resemble radicels; the axon issued from the base enters the white matter of the gyrus. Figure 4 shows
two such giant cells in the human precentral gyrus, in striking contrast to the few large pyramidal cells seen on the right side of the field.
The giant or solitary cells of Meynert found in
layer V of the visual cortex have a similar structure but are smaller in size.
14
The number of cells in a cortical section varies

depending on cell type, cortical layer, and cortical area. We adopted a unit of measurement after
Hammarberg [1895], i.e. a cube measuring 0.1
mm on each side, and thus having a volume of
(0.1 mm)3 or 0.001 mm3. To determine the number of cells contained in such a cube, we take a
section of a known thickness (25 m) and count
the cells in a square measuring 0.1 ! 0.1 mm (exactly like in the haemacytometers used for blood
cell counts). We then multiply that count by the
number of sections contained in 0.1 mm (0.1 mm/
25 m = 4). For example, in figure 2, which was
photographed at a !50 magnification, 5 mm
correspond to 0.1 mm. We therefore count the
cells in a 5 ! 5 mm square, and then multiply
that count by 4 (since the section has a thickness
of 25 m, i.e. 1/4 of 0.1 mm) to obtain the total
number of cells per 0.001 mm3. Average cell count

estimates with this method are shown for each
isocortical layer in table 3.
Table 3. Average thickness and cell number for each

cortical layer of the isocortex. For the present edition,
and throughout the following chapters, the original cell
numbers given per (0.1 mm)3 or 0.001 mm3 were multiplied by 1,000 to yield cell counts per mm3.
Division of the Cerebral Cortex into Isocortex

and Allocortex
Layer
Thickness
mm
Cell content
/mm3
I
II
III
IV
V
VIa
VIb
0.20
0.15
0.75
0.20
0.40
0.50
0.30
5,000
65,000
20,000
80,000
17,000
20,000
10,000
The major part of the human cerebral cortex

shows the six typical layers described. Brodmann
[1908a, c, 1909, 1914] was able to demonstrate this
six-layered (hexalaminar) structure even in the
fetal brain, between the 6th and 8th months of
gestation; at that developmental stage, the layering can be seen even in regions of the cortex in
which, in the adult, the division into six layers is
partially or totally effaced.
We call the entire part of the cortex which features the six distinct layers in the adult, or which
at least has such six layers during embryonic development, isocortex according to Vogt [1903,
1911] and Vogt and Vogt [1902, 1919]. However,
the small part of the brain called rhinencephalon, in the strict sense, shows, both in the embryo
and in the adult, a fundamentally different structure with either incomplete or no lamination at
all. The ensemble of these territories constitutes
the allocortex and comprises the subcallosal gyrus, the intralimbic gyrus with the indusium, the
fasciolar and dentate gyri including Ammons
horn, the subiculum and presubiculum of the
hippocampal gyrus, the uncus, the medial and
lateral olfactory gyri, and perhaps the anterior
substantia perforata.
However, all these allocortical regions only
make up a small part of the human cerebral cortex, not more than 8% of the entire cortical surface. On the other hand, in animals, and among
macrosmatic species in particular, the allocortex
represents the major part of the cortical surface.
Figure 5 illustrates the relation of the isocortex
(white areas) to the allocortex (hatched areas) in
the human (fig. 5c, d) and the hedgehog (fig. 5a,
b) brain, drawn to scale. We see how large the ex-
Introduction
pansion of the allocortex is in this animal, whereas in humans, the corresponding part is reduced
to a very thin band encircling the corpus callosum and the fornix. The chapters on the superior
limbic and the hippocampal (inferior limbic) lobe
deal more extensively with the allocortex. For the
moment, we may leave the subject by reiterating
that the allocortex is also divided into numerous
areas of a very different structure.
Regional Structural Variations of the

Isocortex and Its Layers
Back to the isocortex: the six layers are almost always seen; however, the cortical thickness and
the individual constitution of the layers show
characteristic differences among various cortical
areas. Cortical thickness becomes diminished by
about 50% from its maximum value at the dome
of a gyrus to its minimal value at the valley floor.
Additionally, cortical thickness varies among
different areas of the cerebral hemispheres. Thus,
one can generally say that, on average, the thickness of the cortex of the gyri on the lateral facies
(convexity) of the cerebral hemispheres is about
3.5 mm, whereas in the gyri of the ventral (infe-
15
Fig. 5. a, b Lateral and median hemispheric facies in the hedgehog cerebrum [Brodmann, 1909]. c, d Lateral and
median cerebral hemispheric facies in the human cerebrum, brought to the same size [Economo, 1929a]. The allocortex is marked with horizontal hatching in both species; isocortex is left blank. In the hedgehog, the allocortex
comprises about 75% of the cerebral surface, in the human not more than 8% of the total surface. Olfaction, with
which the allocortex is associated, is more important for many animals; in comparison, vision and hearing acquired
a much more crucial role in humans.
rior) aspect it is only about 3.0 mm, becoming

even smaller (about 2.7 mm) on the median facies, where cortical thickness is the least conspicuous.
Figure 6 shows a diagram with the regional
differences in thickness of the cortex, depicted by
varying degrees of hatching. Thickness measurements span from 4.5 to 1.3 mm. The thickest
point of the cortex is found in the precentral gyrus, in the vicinity of the longitudinal cerebral
(interhemispheric) fissure and the anterior temporal lobe (fig. 6). Cortical thickness progressive-
16
ly diminishes towards the frontal and the occipital pole. The thinnest parts of cortex are situated
in the calcarine sulcus, especially in its floor, as
well as in the central sulcus of Rolando, at the anterior wall of the postcentral gyrus. Taking into
account the 50% reduction in cortical thickness
at the valley floor already mentioned, the mean
for the whole brain is about 2.5 mm.
Wherever the overall cortical thickness varies, it is evident that the thickness of individual
layers will not remain proportional or parallel either: in the frontal lobe, for example, layer III be-
Fig. 5. e, f Lateral and median hemispheric facies in the rabbit cerebrum, showing the relative proportions of homotypic isocortex (blank), simple heterotypic isocortex (vertical hatching), granulous heterotypic isocortex (crossed hatching) and allocortex (horizontal hatching). In this instance, the homotypic isocortex hardly comprises 40% of the total
cerebral surface. g Schematic representation of the concept of the rhinencephalon in the human brain, after an illustration of Villiger [1922]. The intralimbic gyrus which includes the subcallosal gyrus, indusium griseum, fascia
cinerea, fasciolar gyrus, dentate gyrus and the band of Giacomini is densely shaded; the anterior substantia perforata is cross-hatched; the rest of the rhinencephalon is lightly shaded. Compared to the red parts of figure 22a, b in
the Atlas [Economo and Koskinas, 2008] and figure 85b in the Appendix, this drawing illustrates that the concepts of
the rhinencephalon and the allocortex only overlap partially. Abbreviations: R, central sulcus of Rolando; cm, callosomarginal sulcus; po, parieto-occipital sulcus; C, calcarine sulcus; Cc, corpus callosum; fo, fornix. h Schematic drawing
of the ventral (or inferior) surface of the embryonic human cerebrum (anterior half), depicting the anatomic relationships of the allocortex, after an illustration of Villiger [1922]. Schemes supplemented from the original figures 108,
109, 61 and 62 in Economo and Koskinas [1925].
comes progressively thinner as one approaches

the frontal pole, whereas at the same time layer
IV increases. In other words, the thickness of
each layer varies according to a strict plan, depending on cortical area, orientation, and other
factors.
Introduction
Granular and Agranular Cortex
If we drew a diagram of the regional thicknesses

of every cortical layer, just as we have done for the
total cortical depth in figure 6, we would get totally different images for each layer. Thus, granu-
17
Cortical thickness
blank
under 2.0 mm
2.02.5 mm
2.53.0 mm
3.03.5 mm
3.54.0 mm
Fig. 6. Lateral (a) and median (b) facies of the human cerebral hemispheres with various hatching patterns denoting
the varying thickness of the cortex. (For the sake of simplicity, the drawing of the brain contours is based on the
schemes in Economo and Koskinas [1925].)
18
lar layers II and IV are poorest in cells exactly at

those points where the overall cortical thickness
is highest (cf. fig. 7 with fig. 6); these layers may
even disappear entirely in such areas, although
they increase in thickness and in cell density at
the thinnest cortical regions (e.g. the two poles).
The usual six-layered cortex (isocortex) is
called granular cortex when it possesses both
granular layers; the portions of the isocortex,
from which the two granular layers completely
disappear, are referred to as agranular cortex.
Figure 7 depicts the regional abundance of granule cells by means of hatching.
In addition to variations of the granular layers
in thickness and in cell density, the remaining
layers, i.e. layers III, V and VI, undergo other regional and local modifications, such as differences in cell type, cell size, and cytoarchitectonic organization.
In general, one may say that the entire cerebral
cortex anteriorly to the central sulcus of Rolando
contains fewer but more voluminous cells, while
the cortex posteriorly to the central sulcus contains smaller but more numerous cells. Pyramidal cells of layer III and spindle (fusiform) cells of
layer VI in front of the central sulcus are larger
and more conspicuous than those behind it. The
most conspicuous and largest pyramidal cells are
found immediately in front of the central sulcus,
but even within the frontal lobe, they gradually
diminish in size and become less robust as one
approaches the frontal pole. The same holds true
for pyramidal cells in layer V, which progressively thins as well, and becomes poorer in cells from
the central sulcus toward the occipital pole; on
the contrary, such a trend is reversed in the direction of the temporal lobe.
The rhinencephalon, on the other hand, and
all the other cortical regions bordering it, including the insula of Reil, present the particularity of
a dense secondary sublayer within layer V, with
pyramidal cells of a larger size than average, and
a distinct appearance with its high cellularity and
the cell disposition in rows. In the vicinity of the
Introduction
rhinencephalon, layer II cells often lose their

granular appearance and juxtaposition in a distinct layer; they become larger, acquiring a stellate or navicular form, and tend to gather in
small groups or aggregates.
Radial Cortical Striation
Apart from the arrangement of the cerebral cortex into horizontal layers, one frequently encounters a tendency for a radial striation, whereby
cells become disposed in perpendicular columns
to the external surface of a gyrus; such radial cellular arrangements may take the form of either
delicate strips or more solid columns.
A very delicate, perpendicular radiation is
characteristic of the parietal lobe, especially its
inferior regions, and extends as far as the superior temporal gyrus (T1). The remaining gyri of
the temporal lobe are marked by wider strips,
with cells arranged in distinct columns that traverse all six cortical layers.
In the occipital lobe, cells are densely packed,
forming thick but short radial columns. Such a
radial striation is almost absent from the frontal
lobe, the closest trace of similar arrangements being found in the pars triangularis of the inferior
frontal gyrus (F3), and, to a lesser extent, at the
foot of the three frontal gyri, and sometimes in
the frontal pole.
The Ideal Cerebral Cortical Map
With the exception of certain cases that I will discuss later, all these changes take place gradually,
in a stepwise mode. Nevertheless, there are in
certain places, as we shall see soon, very sudden
and abrupt changes in the cytoarchitectonics of
the cortex; in certain areas, this contributes to the
demarcation of precise and sharp boundaries.
If, after these modifications, one attempted to
construct an ideal map of the cerebral cortical
19
blank
Agranular cortex
Slightly granular cortex
Increasing
granule
cell
presence
Granulous cortex
(koniocortex)
Fig. 7. Lateral (a) and median (b) facies of the human cerebral hemispheres, in which the various patterns of hatching denote the varying abundance of granule cells in layers II and IV. (The basic scheme of the brain contours is from
Economo and Koskinas [1925].)
20
Fig. 8. The five fundamental structural types of isocortex: 1, agranular; 2, frontal; 3, parietal;
4, polar; 5, granulous or koniocortex [Economo, 1925a, 1929a; Economo and Koskinas, 1925].
surface, one would have to come up with something different from the scheme of figure 1. That
scheme is really artificial, because otherwise one
would need to enter, separately for each regional
and local variation of each individual layer and
its cells, and on different cortical maps, separate
diagrams, that would then have to be superimposed to form an ideal map of the cortex. Such a
map would consist of as many sheets as there are
cortical layers, with all the changes shown at every single point.
Despite such theoretically justified considerations, I retain, for the sake of practicality, the
area map of the cerebral surface that we had orig-
Introduction
inally constructed: first, because it gives a more

general idea of the entire cerebrum compared to
a so-called ideal map; and second, because there
are in actuality certain areas that can be rigorously delimited.
The Five Structural Types of Isocortex
So far I have described the constant variations

that one observes in each of the layers in different
regions. Alongside the parcellation into cortical
areas, such individualities also involve different
structural types of the six-layered cortex, which
21
characterize much more extensive sectors of the

isocortex than a single area. We distinguish five
such fundamental structural types (fig. 8).
Homotypic Isocortex
The most typical are the cortical types called
frontal (structural type 2), parietal (structural
type 3) and polar (structural type 4), named after
the region where they are most prominent.
The frontal type (cortical structural type 2) is
characterized by six distinct layers. It is thick,
with voluminous and robust pyramidal cells,
regularly arranged in layers III and V; so are the
regularly-placed spindle cells in layer VI. Both
granular layers are distinct, although not very
dense, and their granule cells have mostly a triangular shape.
The parietal type (cortical structural type 3) is
even more dinstinctly six-layered, owing to the
increased thickness and density of both granular
layers (II and IV), the granule cells of which appear round. In contrast, pyramidal cells of layers
III and V are smaller and slenderer, more numerous, and less orderly arranged. Especially in layer
V they are also less robust, being hardly larger
than the already small spindle-shaped (fusiform)
cells of layer VI.
The polar type (cortical structural type 4) is
found at the frontal and occipital poles; it is unusual, because of its thinness and its high cellularity. The granular layers are particularly well represented, rendering this structural type even more
clearly layered. The frontal polar type is different
from the occipital polar type, having a more pronounced layer V with numerous, large, and robust
pyramidal cells, whereas the cells of layer V in the
occipital pole are generally small and inconspicuous, and larger pyramidal cells are rarely seen.
There may be differences in the structure of
these three types of isocortex, but they all clearly
conserve the basic six-layered arrangement characteristic of the granular isocortex.
22
Heterotypic Isocortex
On the other hand, at some points of the isocortex, there can be abrupt and fundamental changes, such that not all six layers are recognizable.
Certain layers are so attenuated that they appear
virtually missing; cellular rearrangements may
also render individual layers indiscernible. These
regions, which do not distinctly reveal the original six layers, are designated heterotypic isocortex, as opposed to the homotypic isocortex with
the usual six-layered granular types (structural
types 2, 3 and 4).
Practically speaking, there are only two isocortical heterotypies in the human brain, called
agranular (structural type 1) and granulous
(structural type 5).
The agranular type (cortical structural type 1)
differs from the homotypic six-layered granular
cortex of the structural types 2, 3 and 4 in that it
totally lacks granule cells (fig. 8), hence its name.
In place of the absent granular layers (II and IV),
one finds cells with a changed aspect, having acquired the shape and size of medium-sized and
small pyramidal cells. Thus, type 1 is also called
agranular pyramidal type. With the disappearance of the granular layers, cells undergo changes
that render them indistinct from the neighboring
layers III and V; such a substitution is a typical
pyramidal transformation or pyramidization of
the cortical elements. The best example of an
agranular heterotypy is the cortex of the precentral gyrus (fig. 11).
The granulous type (cortical structural type
5) is the result of an opposite process (fig. 8). In
contrast to the usual type of granular cortex, not
only do layers II and IV contain large numbers of
granule-sized cells, but the other layers contain
similar small cells as well, in densities and numbers that bolster them into also being called granular layers. Such a change usually involves layer
III in its entirety or in part, and occasionally layers V or VI. Those layers can be hardly distinguished from the regular granular layers (II and
Fig. 9. Distribution of the five structural types of isocortex over the surface of a the lateral and b the median facies
of the human cerebral hemispheres. Structural type numbers as in figure 8.
Introduction
23
IV). Such a granulous transformation or granulization is evident in structural type 5. We have

termed this type of granulous or dusty cortex
koniocortex (from the Greek word = dust).
A classic example of this type is the calcarine cortex, in which the internal granular layer appears
to be duplicated (cf. also fig. 41).
Physiologic Significance and Topographic

Extent of Each Cortical Structural Type
The five structural types of cortex just described

extend over diverse regions of the cerebral cortical surface (fig. 9). The two heterotypic extremes
represent the most important particularities of
the isocortex (structural types 1 and 5, fig. 8). As
we shall see later in the course of our discussion,
the granulous cortex corresponds to the sensory
cortical domain with regard to localization, such
that we may consider it to be the sensory cortex
par excellence. The agranular cortex, on the other hand, by its localization in the precentral gyrus, seems to be primarily efferent and directly
motor in part, especially in regions containing
giant cells. Despite their importance, these two
heterotypies take up a minor part of the isocortex. In figure 9, the agranular heterotypic isocortex (structural type 1) is left blank, and the granulous heterotypic isocortex (koniocortex, structural type 5) is dotted; the remainder represents
homotypic isocortex.
The agranular efferent cortex (structural
type 1) covers the caudal third of the frontal
lobe, anteriorly to the central sulcus on the lateral and median hemispheric facies, i.e. the entire electromotor region, and thence stretches
over the entire anterior limbic gyrus and the parolfactory field of Broca (fig. 9); there is also an
agranular territory on the anterior insula, but
this is not directly continuous with the same
type of cortex on the lateral hemispheric aspect;
a third agranular field, narrow and shaped like
a comma, is found in the retrosplenial segment
24
of the limbic gyrus; finally, a fourth long stretch

is seen at the dome of the hippocampal gyrus
and the uncus, this last really being agranular
allocortex.
The granulous koniocortex (structural type 5)
is found in the posterior wall of the central sulcus, in the two walls and lips of the calcarine sulcus, on the deep transverse temporal gyrus of
Heschl in the lateral (Sylvian) fissure, in the caudal retrosplenial parts of the wall of the callosal
sulcus, and, finally, in the dorsal (upper) wall of
the hippocampal sulcus, this last really being allocortical koniocortex.
The remaining part of the cortical surface (in
so far as it is not clad in allocortex, cf. fig. 5), is
covered by the three structural types of homotypic isocortex, i.e. types 2, 3 and 4 (fig. 9).
The frontal type (structural type 2) is spread
over the rostral two-thirds of the frontal lobe, the
superior parietal lobule, including the dome of
the postcentral gyrus, and the anterior insular
gyri.
The parietal type (structural type 3) mainly
extends over the inferior parietal lobule, including the superior (T1) and fusiform (T4) temporal
gyri, and the rostralmost part of the occipital
lobe. It is worth pointing out that there is a rather well-defined territory in the rostral third of
the frontal lobe with an evident analogy to the
parietal type (structural type 3). The frontal
type, on the other hand, is also found in a block
on the middle (T2) and inferior (T3) temporal
gyri.
The six-layered isocortex (structural types 2
and 3), which occupies the largest part of the
cerebral cortical surface in the human brain,
must be principally (although not exclusively!)
effecting memory and associative functions (the
frontal regions process associative mechanisms,
and the parietal regions sensory associative
mechanisms); the same likely applies to structural type 4.
Parasensory Zones
Dome
Brink (edge)
In all likelihood, future research will reveal additional structural types, beyond the five described. Even now, we regularly see a special type
in a small band at the boundary between any koniocortex and the ordinary surrounding isocortex. Such a zone contains remarkably large pyramidal cells (asterisk in area OB in fig. 40). We
call this region a parasensory zone, and we think
that it serves a reflex efferent function, perhaps
that of attentive presentation to a stimulus.
Wall
wm
Valley
(sulcus oor)
Layer Variations and Layer Proportions
There is yet another typical cytoarchitectonic

variation in cortical structure that I ought to
mention, which is owed to the fact that the cortex
is folded in gyri. The immediate effect of such a
cortical arrangement is that the dome (Kuppel) of
a gyrus has a cortical lamination about twice as
thick as that of the valley floor (Tal). The attenuation of the cortex from the dome to the valley
occurs gradually at the wall (Wand), with the
brink (Kante) of the gyrus being at least as thick,
or even thicker, than the dome (fig. 10; cf. also
Economo and Koskinas [2008, p. 25]).
This gradual thinning of the cortex at the wall
and valley is not equally distributed to all the layers
(fig. 10). One can see that the thickness of layer I in
particular, and of layer II to a lesser degree, increases at the valley. The thickness of layer III becomes
somewhat reduced from the dome to the valley
proportionally to the overall thinning of the cortex, because layer III is always about 33% of the total cortex at the dome, wall and valley floor. Layer
IV also becomes thinner from the dome to the valley. Layers V and VI become substantially thinner,
the extent of such thinning evidenced by the fact
that, while both layers constitute together about
50% of the thickness of the cortex at the dome,
their combined thickness at the valley floor is not
more than 20% of an already very thin cortex.
Introduction
Fig. 10. Changing thickness of layers IVI at the dome,

brink (edge), wall, and valley (sulcus floor) of a gyrus. The
two granular layers (external and internal) are hatched;
wm, subcortical white matter.
Such relations are best exemplified in table 4,

which gives measurements of the cortical thickness at the dome, wall and valley floor in a small
precentral gyrus, along with the values for each
individual layer. The numbers further permit a
calculation of the proportion taken by each layer
at the dome, wall and valley for the sake of comparison (table 4). This clearly demonstrates that
toward the valley, layers I and II increase appreciably, layers III and IV remain about the same,
while layers V and VI substantially diminish in
thickness.
The Gyrus as an Organ
Layers V and VI are the ones from which cortical

efferent projection fibers are mainly issued to
reach more distal areas. Layers III and IV, on the
other hand, function essentially in the reception
of stimuli and the association of intercortical relations. In other words, the layer arrangement
25
within the various sections of a gyrus suggests a

division of labor.
We see a corresponding differentiation of
every part of a gyrus and its components, with
the dome (where layers V and VI are most pronounced) rather functioning as an effector organ,
and the wall and the valley being a receptive and
associative organ. Since the overdevelopment of
efferent layers V and VI necessarily occurs in the
anlage during embryonic life, there must be an
important genetic influence on the formation of
the gyri [Economo, 1926a, b].
The orientation of the cells, whose long axis
usually lies perpendicular to the surface at the
dome of a gyrus, also changes at the wall, and especially at the valley floor of the two lower layers;
the long axis of layer VI cells changes its orientation to lie obliquely to the surface at the wall, and
almost horizontally at the valley floor. In all the
layers, all cells at the dome are at a greater distance from each other, compared to the wall.
Cells of the dome are, on the other hand, larger
and more robust. At the brink (or edge) of a gyrus, i.e. at the transition point from dome to wall,
cells are usually so superimposed that they give
the impression of a radial striation.
The Allocortex and Its Different Structural

Areas
Following this brief description of the structure

of the isocortex, we must now consider the allocortex. Figure 5 gives an idea of its extent; in humans it mainly comprises the cortical region that
borders on the corpus callosum (including the
intralimbic gyrus with the indusium and the
fasciola cinerea or fasciolar gyrus), the entire upper wall of the hippocampal gyrus with the dentate gyrus, the entire distended anterior part of
the hippocampal gyrus, the uncus, and, lastly,
the adjacent structures of the lateral olfactory gyrus at the base of the brain, the substantia perforata and the olfactory tubercle (which no longer
26
belongs to the cortex sensu stricto), the olfactory

triangle, and the medial olfactory gyrus with the
subcallosal gyrus.
In these various regions the allocortex presents with structural differences, sufficiently discernible to allow a delimitation into various cytoarchitectonic areas here as well. A division into
areas is more difficult at the level of the cortical
margin, as the structure keeps changing there,
with fields gradually attenuating and finally disappearing (fig. 19, 22). But in the other regions of
the allocortex, the cytoarchitectonic areas have
much preciser boundaries than in the isocortex,
such boundaries occasionally being razor-sharp
or linear (fig. 5255).
Even in its primordial anlage, the allocortex
appears distinct from the six-layered isocortex.
Such a differentiation is already visible, as I have
shown [Economo, 1930c], as early as the 3rd
month of fetal life, at the level of the anlage of the
cortical plate of His [1904]. Throughout the extent of the hemispheric vesicle, one observes at
that stage, under the thin, reticular marginal zone
that forms the outermost layer of the vesicle wall,
a dense and thick cellular layer, rich in granule
cells and clearly delimited on both its external
and internal sides, called the cortical plate.
This cortical plate is formed by the migration
of neuroblasts from the embryonic matrix that
internally lines the cerebral vesicles. In their migratory course, neuroblasts traverse the intermediate layer, which is poor in cells and later forms
the myelinated white matter. The cortical plate of
His generates the cell layers of the cortex, and the
marginal zone produces cortical layer I, which is
poor in cells.
In the largest part of the cerebral vesicle wall,
which later develops into isocortex, one observes
at the 3rd month of fetal life that the dense and
thick cortical plate is separated from the matrix
by an intermediate layer. The same does not apply
to the region of the vesicle wall that gives rise to
the allocortex. There are three alternative possibilities [Economo, 1927d, 1928e]:
(1) A true cortical plate is missing, and only a superficial mass of neuroblasts, somewhat dense,
is seen, forming cellular bridges just beneath
the marginal zone, in association with the matrix and ganglia derived from it.
(2) A cortical plate is little developed, but at the
same time, neuroblasts in the intermediate
layer stay longer in the matrix and become incompletely fused with the cortical plate.
(3) The vesicle wall remains membranous throughout development, with neuroblasts only forming a narrow layer beneath the matrix.
In these three cases, the cortical plate is either
missing or incompletely developed. From such an
outline will derive, after the 3rd month, the allocortex of the rhinencephalon. Here, apart from a
marginal zone from which layer I (the molecular
layer) will be issued later, one does not see cellular
layers, but incompletely developed layers or layers with a different composition.
With the exception of layer I of the allocortex,
it is not often that one discerns cell layers in the
deeper cellular complement (e.g. substantia perforata). At other places one often gets the impression that individual (e.g. uncus) or multiple layers are missing. For example, in the outermost
cortical margin, one gets the impression (fig. 22)
that only layers V and VI, and more often VI
alone, lie directly beneath layer I (e.g. intralimbic
or olfactory gyrus or subiculum and Ammons
horn).
The attempt to draw homologies between the
layers of isocortex and allocortex is an absolutely
vain enterprise, because these two categories of
cortex are differentiated by their embryonic
structural origins. In spite of such considerations,
we describe the layers of the allocortex with roman numerals, analogously to the isocortex (cf.
subsequent figures), solely on practical grounds,
in order to preserve some uniformity in the sequence of layers, to render evident their relative
overlap, and also to denote the ensuing layer
transitions at the boundaries from the rhinencephalon to neighboring isocortical areas.
Introduction
Great differences in structure are also found

among various allocortical areas; there are zones
where cells may assume in part or in total a granular appearance (e.g. koniocortex of the dorsal
wall of the hippocampal gyrus) or, reversely,
evolve towards the purely pyramidal type (e.g.
Ammons horn). In other words, a granulization
or a pyramidization is seen, just like in the isocortex.
Cortical Area Boundaries
The boundaries between individual isocortical

areas are for the most part imprecise. This happens in many regions, where the transition from
one area to another is so gradual, that it can take
up half or even a full gyrus. One talks of a sharp
and precise transition when it is effected within a
few mm, as e.g. from the foot of the inferior frontal gyrus (F3) to the pars triangularis, and usually also in the transition from heterotypic to homotypic isocortex. A transition that is razorsharp and completed within a determined point
is called linear, as, e.g., in the transition from the
six-layered (hexalaminar) isocortex to the eightlayered (octalaminar) structure of the calcarine
sulcus, or between isocortex and allocortex.
The boundaries between isocortex and allocortex are in general linear, although the isocortex is generally heterotypic at such transition
points, a circumstance that renders the delimitation somewhat difficult.
Cortical Measurements
Before embarking on a detailed description of the

structure of the individual cortical areas, we must
acquaint ourselves with certain numbers regarding the size and relations of the cortex, necessary
for a general orientation.
27
Cortical and Layer Thickness

I already mentioned that the maximum cortical
thickness is 4.5 mm, and the minimal 1.21.4
mm; the average cortical thickness that we adopt
for calculations is 2.5 mm. Using such a mean,
the overall thickness for each layer is shown in
table 3.
Cortical Surface
The total surface expansion of the human cerebral cortex is in the vicinity of 220,000 mm2
[Wagner, 1864; Henneberg, 1910/1911], twothirds of which, or some 145,000 mm2, cover
walls and valleys, and the remaining one-third,
or about 75,000 mm2, corresponds to the free
surface of the cerebral tissue.
Specific Weight of the Cortex

According to Danilewsky [1880], the specific
weight of the cortical gray matter is 1.038 g/cm3
and that of the white matter 1.043 g/cm3.
Table 4. Absolute thickness measurements of the six isocortical layers, and proportional occupancy of the total
cortical thickness by each single isocortical layer (in parentheses), at the dome, wall and valley floor of a small
precentral gyrus.
Layer
Dome
mm (%)
Wall
mm (%)
Valley
mm (%)
I
II
III
IV
V
VI
0.24 (9)
0.17 (7)
0.85 (33)
0.25 (9)
0.55 (20)
0.57 (22)
0.26 (12)
0.17 (7)
0.81 (37)
0.18 (8)
0.40 (18)
0.39 (18)
0.42 (25)
0.25 (15)
0.51 (30)
0.20 (12)
0.16 (10)
0.14 (8)
Total
2.6 (100)
2.2 (100)
1.7 (100)
ures for the average thickness of each layer, we

can estimate the number of cells in a section of
cortex, with a surface of 1 mm2 and an average
depth of 2.5 mm, at about 63,000 cells. Knowing
that the surface of the brain is 220,000 times
1 mm2, we estimate the total cell number in the
cortex at about 14 ! 109. Of these, 8 ! 109 are
large and medium-sized cells of layers III, V and
VI, and the remaining 6 ! 109 are small cells in
all the layers.
Cortical Volume and Weight

With the help of these figures, and of those of the
average thickness (2.5 mm) and surface expansion of the cortex, it is possible to calculate the
volume and the weight of the cortical gray matter
for both hemispheres: they stand at about 560
cm3 and 581 g, while the white matter has a volume of 445 cm3 and a weight of 464 g.
Total Cell Number in the Cortex

According to our estimates, the average counts of
cells per mm3 in the isocortical layers are shown
in table 3. From these numbers, and with the fig-
28
Cellular Weight
The total volume occupied by cortical cells themselves is hardly more than 20.4 cm3, meaning that
the total cellular weight is not more than 21 g.
This can be roughly calculated from the volume
of a medium-sized pyramidal cell of a conical
shape, with a height of 25 m and a base of 20
m, multiplied by the specific weight (1.038 g/
cm3) and the average number of these cells (8 !
109); the same calculation can be made for small
cells. Details on all of these calculations can be
found in our larger work [Economo and Koskinas, 1925].
Gray Matter to Cell (G/C) Coefficient

Thus, we can gather that the ratio of the mass of
the cortical gray matter to the total cellular mass,
i.e. the gray matter to cell coefficient (G/C), is
(560 20.4):20.4 in humans, or about 27:1. The
lower the animal species, the greater becomes the
mass of cells as compared to the gray matter
[Nissl, 1898; Kaes, 1907]. In the G/C coefficient
we then have a factor which may in practice be
used as an index of the stage to which a brain has
evolved [Economo, 1926c].
[1897], Franceschi [1902], Rossi and Roussy [1906,

1907], van Bogaert [1925], and Bertrand and van
Bogaert [1925]); and familial Huntington chorea,
where one observes a rather peculiar proliferation of cells, mostly neuroglial cells, in layer IV
(cf. the works of Klpin [1909] and Lewy [1921]).
However, other diseases, such as mental retardation, are associated with neuronal fallout in all
layers [Hammarberg, 1895]. Cytoarchitectonic
cortical neuropathology is still in its infancy (cf.
concluding chapter).
The Seven Lobes of the Cerebral Hemispheres

Ontogenetic Alterations of the Cellular
Structure of the Cerebral Cortex
In its developmental course from infancy through

old age, the cerebral cortex undergoes considerable structural changes [His, 1904; Jakob and
Onelli, 1911] that are still incompletely understood. Therefore, I shall confine my scope to the
brains of adult men in their fourth decade of life.
However, we do know that the granular layers,
which are absent from the precentral gyrus and
the foot of the superior (F1) and middle (F2) frontal gyri in the adult brain, reach as far as the precentral gyrus in both childhood and fetal life.
We also know that layer II becomes appreciably
thinner with advancing age. It is quite possible
that additional age, race, and especially individual variations exist, as well as side differences between the left and right hemisphere. Further research should clarify these questions.
Neuropathologic Alterations of the Cellular

Structure of the Cerebral Cortex
We equally understand that certain diseases have

a predilection for specific layers or cell groups, as
for instance amyotrophic lateral sclerosis, which
affects the large pyramidal cells of layer V and
layer III (cf. the works of Dotto and Pusateri
Introduction
We can now consider the individual areas of the

cortex. To facilitate orientation, we divide the cerebral hemispheres into seven lobes (cf. tables 1
and 2). Their cytoarchitectonic boundaries do
not necessarily coincide with their gross anatomical (macroscopic) boundaries.
(1) The frontal lobe reaches from the floor of the
central sulcus of Rolando to the frontal pole,
over and around it to the substantia perforata
on the ventral (inferior) hemispheric aspect,
and to the limbic gyrus on the median aspect.
We designate all areas of this lobe with the initial capital letter F.
(2) The parietal lobe stretches behind the central
sulcus on the median hemispheric facies to the
limbic gyrus below, backwards until the interoccipital sulcus, and posteriorly to the parieto-occipital sulcus on the superolateral hemispheric facies (convexity); to the parietal lobe
belongs from a cytoarchitectonic viewpoint
the temporo-occipital transition zone on the
lateral and ventral hemispheric aspects, all the
way to the collateral sulcus. I discuss those regions in the parietal lobe chapter. We designate all areas of this lobe with the initial capital letter P.
(3) The insular lobe is inserted in the form of a
fan between the temporal, parietal, and frontal lobes, being separated from them by the
29
Table 5. Range of cortical thickness at the dome, and average cortical thickness at the wall of gyri in the most important cytoarchitectonic cortical areas. Wall thickness data supplemented from table III in Economo and Koskinas [1925,
p. 796]; blanks signify structures that only appear as dome formations.
Area
Gyrus thickness, mm
dome (range)
wall (mean)
Frontal lobe
FA
FB
FC
FCBm
FD
FD
FD
FDL
FDp
FE
FF
FG
FH
FJ
FK
FL
FM
FN
3.54.5
3.24.0
2.63.4
3.03.2
2.43.0
2.42.6
2.72.9
2.62.8
2.42.5
2.32.6
2.73.2
2.02.4
2.52.8
3.13.5
0.52.5
1.82.5
0.41.8
0.31.0
3.3
2.8
2.9
2.8
2.5
2.6
2.5
2.1
2.7
1.8
2.0
3.0
Parietal lobe
PA1
PA2
PB1
PB2
PC
PD
PE
PF
PG
PH
2.02.2
2.52.8
1.92.2
1.92.1
3.03.3
2.22.4
2.83.0
3.13.6
3.03.4
2.53.0
2.0
1.9
2.0
2.0
2.3
2.6
2.4
2.2
posterior, superior, and anterior insular sulci.

We designate all areas of this lobe with the initial capital letter I.
(4) The occipital lobe is a direct caudal prolongation of the parietal lobe, from which it is separated cytoarchitectonically by the parieto-occipital and interoccipital sulci. We designate
all areas of this lobe with the initial capital letter O.
30
Area
Gyrus thickness, mm
dome (range)
wall (mean)
Insular lobe
IA
IB
2.93.1
2.83.0
2.7
2.6
Occipital lobe
OA
OB
OC
2.32.6
1.82.2
1.82.3
1.8
1.6
2.0
Temporal lobe
TA
TB
TC
TD
TE
TF
TG
2.83.0
2.93.0
2.72.9
2.72.8
3.23.6
2.93.1
3.23.8
2.1
2.7
2.7
2.5
3.1
2.5
3.6
Superior limbic lobe

LA
2.62.9
LC
2.73.0
LD
2.22.4
LE
0.42.3
2.4
2.4
2.2
2.2
Hippocampal (inferior limbic) lobe

HA
2.83.0
HB
2.62.9
HC
2.42.7
HD
2.22.4
HE
1.23.0
HF
0.60.7
2.9
2.5
2.0
2.3
2.2
0.6
(5) The temporal lobe spreads from the floor of

the lateral (Sylvian) fissure to the collateral
sulcus; caudally, and on cytoarchitectonic
grounds, this lobe does not extend as far as the
preoccipital incisure (notch), but ends rostrally to it, half-way between the notch and the
caudal end of the lateral fissure. We designate
all areas of this lobe with the initial capital letter T.
(6) The superior limbic lobe represents the superior semicircular part of the fornicate gyrus
( grand lobe limbique of Broca), which on the
median aspect encircles the corpus callosum;
it includes the cingulate gyrus and the retrosplenial region as far as the isthmus. We designate all areas of this lobe with the initial capital letter L.
(7) The hippocampal (inferior limbic) lobe constitutes the inferior part of the fornicate gyrus
that is accompanied by the fimbria; it includes
the hippocampal and dentate gyri and the uncus. We designate all areas of this lobe with
the initial capital letter H.
These lobes form the subject matter of the
subsequent seven chapters, with the corresponding cytoarchitectonic areas treated.
Division of the Cerebral Cortex into

Cytoarchitectonic Areas
Individual lobes are composed of numerous, cytoarchitectonically diverse areas (fig. 1; tables 1,
2), which we identify, apart from their full Latin
names, with a series of graphic symbols composed of two capital characters.
The first character (roman type) denotes the
initial of the lobe in which an area is situated; for
example, the letter F precedes all the areas that
belong to the frontal lobe, the letter T all the areas
Introduction
of the temporal lobe, etc. A second character (corsiva type) denotes the sequence of a ground area
in its topographic anatomic succession within a
lobe (FA, FB, FC, etc.). Large-cell (magnocellular)
or small-cell (parvicellular) varieties, plus other
modifications, are indicated by a Latin, Greek or
number subscript (e.g. FDm stands for granular
frontal area, pars magnocellularis).
Transition zones from one area to another are
denoted by the use of the two corsiva capitals, following the lobe initial (e.g. FCD). A code with a
second corsiva capital in parentheses, such as the
designation FC(B), implies a part of area FC with
an admixture of the type of the neighboring area
FB.
For each area, I provide a figure with a microphotographic registration from the corresponding tissue section. The majority of figures are
shown at a !45 magnification; figures of entire
gyri are shown at magnifications of !8 to !15,
as marked in the captions. All the photographed
sections had a uniform thickness of 25 m.
Cortical Thickness of the Various

Cytoarchitectonic Areas at the Gyral Dome
Table 5 gives the thickness of gyri (at the dome
and wall) for the most important cortical areas.
Compare these numbers with figure 6.
31
Frontal Lobe
Cellular Structure
The frontal lobe is covered with a thick, well-developed cortex, not particularly rich in cells. The
robustness and orderly disposition of pyramidal
cells in layers III and V is quite remarkable and is
hardly seen anywhere else in the cerebral hemispheres. Layer VI is equally well formed and
shows relatively large and orderly arranged spindle (fusiform) cells, oriented in the direction of
the incoming radial myelinated fibers. In general, layers II and IV are less well developed, their
cells being mostly small and triangular; at some
places they may be absent altogether.
I shall begin the description of the areas of the
frontal lobe from the central sulcus of Rolando
and move stepwise to the frontal pole; in the
meantime, I remind you once again of the six-layered (hexalaminar) homotypic structure of the
isocortex (fig. 2), and furthermore the cortical
maps of figure 1, to which I shall refer repeatedly
during the discussion of the topography of individual areas.
Precentral Area FA
To begin with an exception, the first cortical field

that one encounters on the entire precentral gyrus
(fig. 1a, c) is the precentral area FA (area EK 1), a
heterotypic agranular cortex (fig. 7a, b, 9a, b). In

figure 11, it can readily be seen that the distinct
layers II and IV, which were characterized by the
presence of numerous granule cells in figure 2, are
absent from area FA. One may indeed note the cortical zones where the external and internal granular layers are normally present (layers III(II), III(IV)
and V(IV) in fig. 11). Yet, these layers have become
so deprived of cells and cell size is so large that one
can no longer speak of granular layers; they are no
longer granule cells, but rather pyramidal forms of
a small or a medium caliber. The result is that
from layer I all the way to the spindle (fusiform)
cells of layer VI, we have a unique, unusually thick
pyramidal layer, which occupies the entire space,
and which would normally correspond to the four
layers IIV. In this 2.53.0 mm thick layer, the pyramidal cells are not spread very regularly.
The cortex of area FA reveals an unusual overall thickness, measuring about 4.5 mm at the superomedial hemispheric edge, and staying between 3.5 and 3.2 mm at the opercular segment
of the precentral gyrus. All magnocellular layers
participate in the cortical thickening of area FA,
but in various degrees (cf. fig. 11 and 12, where
layers are marked on the sides of the microphotographs).
Layer I is rather poor in cells (table 6). The upper sublayer is more granular than the lower (better seen as Ia and Ib in fig. 12).
Fig. 11. Giant pyramidal precentral area FA (area EK 2). Caudal (posterior) brink of precentral gyrus, dorsal part.
!45.
Frontal Lobe
33
Fig. 12. Giant pyramidal precentral area FA (area EK 2). Dome of paracentral lobule. !45.
34
Layer III essentially comprises five secondary

sublayers: III(II), IIIa, IIIb, IIIc and III(IV). It measures about 1.5 mm in thickness, and its cells are
not arranged in a very orderly fashion. Layer II is
missing (fig. 11), having turned into the upper
part of layer III and accordingly denoted as layer
III(II). The cells are pyramidal instead of granule.
A real sublayer IIIa follows, consisting of small
pyramidal cells; then comes sublayer IIIb, with
medium-sized pyramidal cells, and sublayer IIIc,
with multiple, superimposed large pyramidal
cells; here, one frequently finds single, extremely
large pyramidal cells, 5060 m in height, with
distinct tigroid (Nissl) bodies. Underneath sublayer IIIc, instead of a granular layer IV, lies another layer of pyramidal cells, called layer III(IV)
and containing medium-sized pyramidal cells;
these cells replace the granule cells that normally
populate layer IV, having been subjected to pyramidization, such that area FA lacks a layer IV as
well.
Layer V lies directly beneath and is also thick
(table 6); its upper part is composed of smaller
pyramidal cells, which are still within the boundaries of layer IV; we accordingly denote this motif
as layer V(IV) (fig. 11). Immediately underneath it,
layer V consists of large and medium-sized pyramidal cells, arranged somewhat irregularly.
Giant Cells of Betz. Beneath the cells just described, especially in caudal sections of area FA,
in the deeper zone of layer V one finds solitary
examples or small groups of 34 elements, the colossal, well-known giant cells of Betz (Betzsche
Riesenzellen), readily discernible in the microphotographs of figures 11 and 12 as the largest
cells of the entire cerebral cortex. They are most
numerous and largest at the anterior segment of
the paracentral lobule and the dorsal tier of the
precentral gyrus closest to the superomedial
hemispheric edge. Ventrally, their number decreases rapidly, and they have already disappeared especially from the dome of the precentral
gyrus at the level of the inferior frontal sulcus
( f2). At the level of the inferior frontal gyrus (F3),
Frontal Lobe
they are only found in the posterior wall of the

precentral gyrus which dips into the central sulcus of Rolando, although area FA, with its typical
structure but now devoid of giant cells still
vests here part of the dome of the precentral gyrus. Therefore, the giant cells are not present in
the entire precentral area FA, but mainly found in
its caudal and dorsal segment, which is accordingly termed pars gigantopyramidalis or giant pyramidal precentral area FA (area EK 2) (cf. also
plates P1P4 in the Atlas [Economo and Koskinas, 2008]). It only reaches as far as the anterior
boundary of area FA at the superomedial hemispheric edge and the paracentral lobule, where
the anterior boundaries of areas FA and FA coincide (fig. 1a, c); ventrally, however, the anterior
margin of area FA becomes increasingly free of
giant cells. Giant cells are mostly found at the
deepest parts of layer V, forming (fig. 4) colossal,
plump, pyramidal, and bottle-shaped structures,
with numerous radicel-shaped processes. At
about the center of the soma lies the large, pale,
vesicular nucleus, with its large, darkly staining,
intense nucleolus; in the cytoplasm there are
large, distinct Nissl bodies; at the base of the
soma, one usually finds lipid- and pigment-laden
inclusions (left side of fig. 4). The cells of Betz are
accompanied by about half a dozen small satellite
cells (Trabantzellen) each. Layer V in area FA does
not lend itself to any further subdivision.
Layer VI has an upper sublayer VIa and a lower sublayer VIb. In general, spindle (fusiform)
cells are somewhat more orderly arranged than
the pyramidal cells of layers III and V. The transition to the white matter is gradual, and the border
imprecise.
Giant Pyramidal Precentral Area FA

Area FA (including modification FA ) coats the
largest part of the precentral gyrus, commencing
at its posterior boundary at the floor of the central sulcus of Rolando (fig. 1a, b); to be precise, it
35
reaches dorsally close to the superomedial hemispheric edge (fig. 1c) and also over the anterior
wall of the precentral gyrus in the precentral sulcus to the foot of the superior frontal gyrus (pF1).
Ventrally, its anterior boundary rapidly recedes,
in that it is already behind the rostral brink of the
precentral gyrus at the level of the superior frontal sulcus ( f1); at the level of f2, it again lies behind
the midpoint of the dome of the precentral gyrus
to rapidly sink into the ventral extreme of the
floor of the central sulcus of Rolando on top of
the lateral (Sylvian) fissure.
On the median hemispheric facies (fig. 1b),
this area occupies the anterior part of the paracentral lobule, continuing ventrally to the callosomarginal sulcus, anteriorly to the precentral
sulcus, and posteriorly to the paracentral fossa.
On the median facies, and in the region of F1 on
the superolateral hemispheric convexity, the
boundaries of the giant pyramidal area FA (area
EK 2) totally coincide with those of the precentral
area FA (area EK 1); however, from this point on
and further below, the anterior boundary of area
FA recedes much more rapidly than that of area
FA, such that at the level of the middle frontal gyrus (F2) the former has reached its posterior brink
and has totally sunk, at the vicinity of F3, in the
depths of its posterior wall; in other words, at the
level of F3, giant cells are only found in the deeper segments of the posterior wall of the precentral
gyrus sunk into the central sulcus of Rolando.
Paracentral Lobule
The thickness of area FA is not the same through
its entire extent; however, at both walls of the precentral gyrus, it becomes less attenuated than one
normally sees in the cortex. At the level of the
paracentral lobule (fig. 12; cf. also plates P3 and
P4 in the Atlas [Economo and Koskinas, 2008]),
and at the wall of the central sulcus, there are indications in area FA of an internal and partially
an external granular layer (II and IV). Betz cells
36
appear more numerous and compacted especially in the paracentral lobule; they almost form a
cellular lining and lie deeper, i.e. closer to the
white matter, than in the precentral gyrus.
Function
The precentral area corresponds to the highly excitable electromotor zone, and we must view it as
the prototype of a motor cortex. In dogs, it approximately encircles the cruciate sulcus. Area
FA is the source of origin of the largest part of the
pyramidal tract, but of other descending tracts as
well. Betz cells are considered to be the origin of
the pyramidal tract, as they degenerate in amyotrophic lateral sclerosis and as they undergo
marked regressive changes in certain forms of
hemiplegias. Nonetheless, we think that axons of
other large pyramidal cells of area FA also join
the pyramidal tract.
Agranular Frontal Area FB
Directly rostral, the area FA just described is succeeded by the agranular frontal area FB (area EK
4) in its entire extent (fig. 1a, c). Area FB shows an
extremely similar structure to area FA, except for
the complete absence of giant cells of Betz, a slight
general reduction in cortical thickness, and a finer, orderly radial disposition of its cells. Here
again, neither a layer II nor a layer IV is discernible in the form of a granular layer; thus, area FB
exhibits an agranular heterotypic structure (cortical structural type 1) as well. With the exception of layer VI, it also consists of robust, large
pyramidal cells. As just mentioned, its cells are
arranged radially in a more obvious manner than
area FA, rendering a more pleasing and orderly
cytoarchitectonic picture (fig. 13; cf. also plates
P6P8 in the Atlas [Economo and Koskinas,
2008]).
Table 6. Summative table of quantitative data in sixteen fundamental areas of the frontal lobe. Overall layer thickness based on the present work. Separate dome and wall data and some additional values supplemented from Tables I, III, V and VI in Economo and Koskinas [1925, pp. 794801].
Area
Area name
symbol
Cortical
layer
FA
I
III(II)
IIIa
IIIb
IIIc
III(IV)
V
FB
FC
FCBm
FD
precentral area
agranular frontal
area
intermediate
frontal area
magnocellular
agranular
intermediate
frontal (Brocas)
area
granular frontal
area
Frontal Lobe
Layer thickness at dome

mm
Layer thickness at wall

mm
Layer thickness overall

mm
0.18
0.20
0.18
1.47
1.40
1.43
0.80
0.70
0.85
VIa
VIb
1.00
0.70
0.70
0.40
I
III(II)
IIIa
IIIb
IIIc
III(IV)
V
VIa
VIb
0.22
0.27
0.16
I
II
IIIa
IIIb
IIIc
IV
Va
Vb
VIa
VIb
I
II
IIIa
IIIb
IIIc
IV
Va
Vb
VIa
VIb
I
II
IIIa
IIIb
1.40
1.20
1.25
0.22
1.50
0.50
0.90
0.60
0.03
0.47
0.33
0.37
0.26
0.12
0.38
0.17
0.25
0.15
1.00
0.90
1.00
0.20
0.25
0.15
0.46
0.43
0.50
0.70
0.45
0.50
0.30
0.21
0.18
0.27
0.18
0.24
0.18
1.00
1.00
1.00
0.16
0.18
0.17
0.46
0.40
0.43
0.70
0.40
0.55
0.25
0.21
0.18
0.23
0.20
0.20
0.19
0.78
0.89
0.80
0.50
1.37
0.25
0.95
Cell content
cells/mm3
7,000
55,000
30,000
20,000
15,000
13,500
16,000
25,000
15,000
Cell size, m*
H(minmax)/
W(minmax)
5/6
8/7
12/10
20/15
3060/20
20/10
2030/20
6120/3060
30/15
15/10
7,000
65,000
30,000
25,000
18,000
30,000
25,000
24,000
15,000
4/8
7/8
17/12
30/15
3580/2030
5,000
55,000
24,000
17,000
28,000
45,000
30,000
15,000
20,000
12,000
4/8
7/5
1520/812
2530/1012
3050/1020
1015/9
2025/1020
3040/20
30/15
15/10
5,000
55,000
28,000
26,000
20,000
60,000
32,000
16,000
20,000
12,500
4/8
1015/68
1520/812
2530/1030
3060/2025
615/410
2025/1020
2545/1520
2030/1015
20/10
9,000
75,000
32,000
16,000
46/810
515/412
1520/710
1540/720
3040/2025
30/15
15/10
37
Table 6 (continued)
Area
Area name
symbol
Cortical
layer
IV
Va1
Va2
Vb
VIa
VIb
FD
middle granular
frontal area
FE
triangular
granular frontal
area
frontopolar area
I
II
IIIa
IIIb/c
IV
Va
Vb
VIa
VIb
38
orbital area
(agranular
orbital area FF)
I
II
IIIa
IIIb
IV
Va1
Va2
Vb
VIa
VIb
FF
I
II
IIIa
IIIb
IV
V
VIa
VIb
FD

mm
I
II
IIIa
IIIb
IIIc
IV
Va
Vb
VIa
VIb

mm

mm
0.21
0.26
0.24
0.45
0.35
0.45
0.52
0.35
0.33
0.20
0.25
0.18
Cell content
cells/mm3
Cell size, m*
H(minmax)/
W(minmax)
0.90
85,000
35,000
20,000
12,000
35,000
15,000
610/510
1520/1520
2030/1525
2040/20
1530/1015
1520/10
0.27
0.22
0.26
0.20
9,000
65,000
0.82
0.80
0.81
32,500
0.24
0.40
0.45
0.36
0.27
0.30
0.35
0.30
0.25
0.35
75,000
30,000
37,000
20,000
46/810
67/46
1015/68
1520/710
2030/1015
68/58
20/1015
20/10
20/10
0.18
0.12
0.25
0.16
0.21
0.14
0.78
1.05
0.91
0.21
0.24
0.22
0.38
0.40
0.39
0.50
0.34
0.36
0.20
8,000
65,000
25,000
25,000
70,000
30,000
12,000
37,500
18,000
46/810
67/46
1015/10
2060/1540
68/58
1530/1525
20/20
20/810
15/810
0.24
0.15
0.25
0.22
0.22
0.15
5,000
90,000
0.66
0.63
0.60
0.29
0.22
0.30
0.43
0.33
0.40
0.45
0.50
0.23
0.20
40,000
30,000
80,000
50,000
40,000
28,000
40,000
20,000
45/610
57/56
1015/710
815/610
1530/1020
515/510
1015/810
2530/1014
1025/810
1520/10
1015/57
0.30
0.10
0.30
(0.10)
0.30
0.10
1.00
1.00
1.00
(0.06)
(0.04)
0.05
0.64
0.60
0.62
0.60
0.50
0.40
0.25
12,000
60,000
30,000
18,000
24,000
12,500
37,500
30,000
26,000
12,000
79/34
715/38
1520/710
2025/10
2530/1015
57/56
2030/1015
1015/710
1520/710
1015/57
0.78
0.70
0.70
0.87
Table 6 (continued)
Cell size, m*
H(minmax)/
W(minmax)
5,000
100,000
25,000
75,000
60,000
30,000
20,000
45/610
515/510
1030/815
6/610
1020/1015
15/10
1015/7
7,000
75,000
30,000
25,000
70,000
40,000
25,000
26,000
12,000
68/45
710/47
20/10
2530/1015
610/56
2030/15
1025/715
1015/57
10/7
16,000
60,000
(50,000)
40,000
20,000
23,000
15,000
820/10
520/510
1520/5
3040/710
(710/57)
2025/1020
2580/710
3040/1015
30/710
0.55
(0.14)
0.52
0.30
0.90
2,500
30,000
15,000
17,500
30,000
30,000
5/10
1020/10
3035/1520
2040/1520
30/10
30/10
0.36
0.90
0.34
17,500
25,000
45,000
40,000
40,000
17,000
56/56
1825/1013
2030/1015
2040/710
20/10
15/7
0.30
0.70
0.40
15,000
30,000
30,000
12,000
56/56
2060/710
20/10
15/7
0.30
0.80
16,000
3050/2030
Cortical
layer
FG
I
II
III
IV
V
VIa
VIb
0.19
0.18
0.44
0.19
0.35
0.39
0.45
0.23
0.16
0.56
0.16
0.25
0.22
0.20
I
II
IIIa/b
IIIc
IV
Va
Vb
VIa
VIb
0.22
0.12
0.23
0.10
0.22
0.11
0.72
0.64
0.68
(0.16)
0.15
0.15
0.46
0.37
0.42
0.52
0.55
0.26
0.29
0.28
0.08
0.27
0.07
0.90
1.26
1.08
25,000
0.00
0.10
(0.05)
0.70
0.55
0.62
0.80
0.60
0.37
0.35
FH
FJ
FK
FL
straight area
prefrontal area
frontoinsular
area
I
II
IIIa
IIIb
IV
Va
Vb
VIa
VIb

mm
Cell content
cells/mm3
Area
Area name
symbol
frontal piriform
area
I
II
III
V
VIa
VIb
0.55
(0.14)
0.52
0.30
0.60
0.30
parolfactory area
I
III
Va
Vb
VIa
VIb
0.36
0.90
0.34
0.30
0.20
FM
geniculate area
I
Vb
VIa
VIb
0.30
0.70
0.20
0.20
FN
precommissural
area
I
VIb
0.30
0.80
Frontal Lobe

mm
dome
structure
only
dome
structure
only
dome
structure
only
dome
structure
only

mm
0.20
0.16
0.32
0.18
0.35
0.80
0.80
0.28
0.08
1.06
0.50
39
Table 6 (continued)
* Editors note: As Koskinas explains in the Appendix, in the larger textbook the authors adopted a special notation
to represent cell size, consisting of a horizontal line (which looked like the fraction symbol, but was not intended to
be a true fraction in the mathematical sense), with ranges of cell height and cell width indicated, respectively, above
and beneath such a line. If one chose to consider such a notation as a fraction, one could then obtain the height-towidth (H/W) ratio of a cell, in other words, an index of its shape and slenderness (Schlankheit). Based on that criterion, Economo and Koskinas [1925] subdivided pyramidal cells into five categories, as Koskinas describes, from overly slender (H/W = 2.5) to flattened (H/W = 0.5). In mathematical terms, the two dimensions of a cell that express its
size (height and width) in a microphotograph represent an ordered pair, i.e. a pair of numbers written in a particular
order. The correct notation would be H, W, using angle brackets (not to be confused with the less-than < or greater-than > inequality operators) rather than parentheses, to extinguish a possible ambiguity with the notation of an
open interval on the real number line. Whenever a range is included instead of the average height or width, that
should be moreover expressed as an ordered pair of bounded intervals, written as [Hmin, Hmax], [Wmin, Wmax]. Such
a notation, although mathematically precise, might overwhelm the reader. On the other hand, a modern way, commonly used, of denoting the two dimensions of an object is H ! W. This convention, although practical, is arbitrary
with regard to connoting the cells surface area, i.e. the quantity expressing its size in an Euclidean plane, because it
might be construed as representing a mathematical product. Cell size, which can be calculcated from the cell dimensions H, W, will depend on the geometrical shape of any given cell: for example, if a cell appeared as a parallelogram,
the area would be bh, where b = W and h = H are the lengths of the base and the perpendicular height; if a cell were a
triangle, then the area would be 1/2 bh, where b = W and h = H are the base and altitude (measured perpendicular to
the base); if the cell appeared as an ellipse, then the area would be ab, where a = H/2 and b = W/2 are the semi-major and semi-minor axes; and if it were a circle, then the area would be 1/4 d2, with d = H = W representing the diameter. Taking into account all these considerations, the present edition abides by the original idea of the authors to
express cell height and width in a form resembling a fraction (rather than as an ordered pair or a mathematical product), with the only exception of substituting the fraction-like horizontal line () with a forward slash punctuation (/)
to maintain tidiness in the appearance of the Table rows. The closed range between minimum and maximum measured dimensions is denoted by means of an n-dash (), whereby e.g. 3060 m should be read as thirty to sixty
micrometers.
In this region, the cortex is about 4.0 mm thick

near the superomedial hemispheric edge, and
gradually diminishes to 3.2 mm in the dorsoventral direction.
Layer I only contains scanty small neurons
(table 6).
Layer III lies immediately beneath, containing in its upper layer III(II) the rather denser cells
of the former layer II that have undergone a pyramidal transformation. Then follows sublayer IIIa
of the small pyramidal cells, and sublayer IIIb of
the medium-sized pyramidal cells; sublayer IIIc
contains multiple rows of very large, slender, and
robust pyramidal cells. Sublayer IIIc is much
more distinctive here than in area FA . Underneath it lies the layer of medium-sized and large
pyramidal cells, transformed from the granule
40
cells of the former granular layer IV; rather than

forming an independent layer III(IV), these cells
instead mark the direct passage from layer III to
layer V.
Layer V lies directly beneath, with its border
on the deepest part of layer III being absolutely
indistinct and its cells about similar in size and
number to those of layer III, but arranged less orderly. As already stated, it does not contain any
giant cells of Betz. Here again, layer V does not
lend itself to any further subdivision into sublayers.
Layer VI comes next, containing larger spindle (fusiform) cells in its upper sublayer VIa, and
smaller spindle cells more loosely arranged in its
lower sublayer VIb. The demarcation vis--vis
the white matter is also blurred here.
Like area FA, area FB stretches from the callosomarginal sulcus on the median hemispheric
facies, over the superomedial hemispheric edge
and the entire frontal superolateral convexity, to
the opercular region; its rostrocaudal dimension
considerably diminishes as one traces the region
ventrally, such that, while it covers several wide
strips of the massive F1 on the superomedial edge
of the longitudinal cerebral fissure, it is reduced
to the precentral gyrus alone in the opercular region (fig. 1a, c). Thus, its expansion takes the
form of a triangle, the base of which is found on
the median hemispheric facies, and the tip at the
operculum of Rolando.
Its posterior boundary is formed by the anterior confines of area FA discussed earlier. This
boundary is rather sharp and can be recognized
in microscopic preparations by the criterion that
layer III, which is somewhat thicker in area FB ,
thickens abruptly at the boundary line from area
FA and area FB , and its cells become arranged in
much finer radial strands (cf. plate P5 in the Atlas
[Economo and Koskinas, 2008]).
The anterior boundary of area FB , which on the
superomedial hemispheric margin lies about 6 cm
rostral to its posterior boundary and passes
straight across F1 on the superolateral hemispheric convexity, steps back a good deal with the passage to F2 (fig. 1a), and stretches over F2 obliquely
in the ventral and caudal direction, such that at the
level of f2 it sinks into the inferior ramus of the precentral sulcus, and area FB here (at the level of F3)
occupies no more than the anterior segment of the
opercular part of the precentral gyrus. Overall,
area FB can be viewed as an example of how cytoarchitectonic cortical fields may not be delimited
by gyri and sulci, but instead course across them.
I mentioned earlier that this area shows certain
regular variations in thickness, and moreover, regional variations in the size of its cells, with pyramidal cells becoming progressively smaller toward the frontal pole; finally, cells in F1, caudally
in particular, near the superomedial hemispheric
edge, appear larger than anywhere else.
Frontal Lobe
Function
Area FB still belongs to the electromotor zone of
the cerebral cortex, although stronger currents
are necessary to provoke movements from this
part than from area FA . It seems likely that more
complex motor combinations are localized in area
FB ; for example, this area contains, among others, the writing center (on F2 [Economo, 1927d])
and the center for upright posture and walking (on
F1 [Economo, 1927d]). Thus, area FB seems to belong probably to motor cortex of a higher order.
Intermediate Frontal Area FC
Just like area FB extends as a wide dorsoventral

band over the entire superolateral convexity of
the frontal lobe, so does the intermediate frontal
area FC (area EK 6), forming a narrower band and
located immediately rostral to area FB (fig. 1ac).
In this region, the cortex is still rather thick.
However, its thickness diminishes as one moves
anteriorly, such that the caudal maximum of 3.5
mm at the superomedial hemispheric edge drops
to 3.1 mm at the same level rostrally, while ventrally, it sinks to 3.0 or even 2.9 mm. Pyramidal
cells are still quite large, but evidently somewhat
smaller than in area FB (fig. 14; cf. also plates P12
and P13 in the Atlas [Economo and Koskinas,
2008]).
The most striking difference is the appearance of thin and frequently interrupted, yet definite external and internal granular layers (II and
IV), both of which can better be seen at the gyral
walls than at the dome; both of these layers become continually more distinct toward the frontal pole. Their cells are actually small and triangular, rather than round granule cells; one can
see these characteristics in figure 14 by using a
magnifying loupe.
Layer I is thicker than in the two previous areas (table 6). As usual, it is poor in cells.
41
Fig. 13. Agranular frontal area FB (area EK 4). Midpoint of the posterior third of superior frontal gyrus (F1), dome at
superolateral hemispheric convexity. !45.
42
Fig. 14. Intermediate frontal area FC (area EK 6). Anterior part of the posterior third of superior frontal gyrus (F1), dome
at superolateral hemispheric convexity. !45.
Frontal Lobe
43
Layer II is well demarcated from layer I, and

also distinctly, although not too sharply, from
sublayer IIIa directly beneath. It is easily recognizable by its greater density of dwarf pyramidal
cells (Zwergpyramidenzellen).
Layer III is distinctly subdivided into sublayers IIIa (small pyramidal cells), IIIb (mediumsized pyramidal cells), and IIIc (with 50% large
pyramidal cells and the rest medium and small).
Sublayer IIIb is clearly the lightest stained of all.
Taking into account the quantitative measurements in area FB (table 6), it becomes obvious
that cell size decreases in an anterior direction.
Layer IV is less well defined from layer III
above and layer V below, and frequently interrupted, especially at the gyral brinks. This layer
is rather poor in cells, mostly small pyramidal.
Layer V is subdivided into an upper sublayer
Va fairly rich in cells, and a lower sublayer Vb,
containing half the number. Cells are mostly medium-sized pyramidal. Sublayer Va may be readily discerned in figure 14 as a dense cellular layer
interposed between IV and V; it contrasts highly
with the smaller and more sparsely arranged cells
of sublayer Vb, corresponding to just beneath the
mark V.
Layer VI comprises sublayers VIa and VIb
(table 6). The demarcation from the white matter
is no longer as indistinct as in the two previous
(caudal) areas.
Thus, area FC is already a granular homotypic,
and definitely six-layered (hexalaminar) structure. In the sagittal direction, this area covers 2
3 cm on F1 immediately in front of area FB . On
F2, however, it becomes much narrower (fig. 1a)
and its anterior boundary recedes sharply. In all,
one may say that the anterior boundary of this
area shares the tendency of frontal gyri to change
from the wide type of the posterior frontal lobe to
the narrower type of the anterior frontal lobe.
The modification at the foot of F3 (pF3), known
as Brocas area FCBm (area EK 8), will be discussed
in detail later.
44
Intermediate FC I and Agranular FBI Insular

Frontal Transition Areas
At the boundary between the upturned inner
wall of the operculum and the superior ramus of
the circular insular sulcus, the distinctive layer V
and particularly sublayer Va that characterize insular structures, with their robust pyramidal
cells (cf. table 8), are extended toward the opercular parts of the adjacent area FC (and FB as well)
at the depths of the lateral (Sylvian) fissure, such
that one may designate a transition zone termed
intermediate insular frontal area FC I (area EK 9,
table 1; also, a respective agranular insular frontal area FB I, cf. table 2; cf. also plate P16 in the
Atlas [Economo and Koskinas, 2008]). However,
one should keep in mind that very large differences are noted among individual brains [Economo and Koskinas, 1925, pp. 323330, 491, 492].
Function
Area FC also belongs to the electromotor zone, but
here again currents must be stronger than in area
FB to elicit functional responses. The center for
spying or exploratory eye movements (intentional
conjugated ocular movements) is located in the region of F2 vested by area FC. Brocas motor speech
area is located on pF3, as we shall see later, associated with a slight FC cytoarchitectonic modification. Thus, we must conclude that area FC subserves higher motor praxic functions.
Granular Frontal Area FD
Rostral to area FC, the cerebral cortex becomes

distinctly granular, and both layers II and IV can
be seen as conspicuous, thick layers, thus lending
the cerebral cortex a typical horizontally stratified appearance in the cytoarchitectonic picture
(fig. 15; cf. also plates P24 and P25 in the Atlas
[Economo and Koskinas, 2008]). Recognizable
Fig. 15. Granular frontal area FD (area EK 11). Middle third of superior frontal gyrus (F1) and transition zone to middle
frontal gyrus (F2). Dome at superolateral hemispheric convexity. !45.
Frontal Lobe
45
by this distinctive lamination, the granular frontal area FD (area EK 11) coats in its broad expansion the entire superolateral hemispheric convexity of the rostral one-third of the frontal lobe,
from the opercular pars triangularis to the callosomarginal sulcus on the median hemispheric
facies, beginning just in front of area FC and only
leaving free the frontal pole (fig. 1ac).
The entire cortex progressively diminishes in
thickness in all its parts in a posteroanterior direction. As I mentioned earlier, its cells also become smaller in the same direction. This is particularly valid for both pyramidal layers III and
V. However, at the superomedial hemispheric
edge and in F3, the decrease in pyramidal cell size
is somewhat slower, such that in both of these territories one still finds relatively large pyramidal
cells in area FD.
Cortical thickness in area FD averages 2.8 mm
on the domes, with a fluctuation between 3.0 mm
(in its most posterior zones and in the vicinity of
the longitudinal cerebral fissure) and 2.4 mm (in
its most anterior and ventral regions).
Layer I is rather thin and contains slightly
more numerous cells than the previous areas (table 6).
Layer II is overall distinct and contains small
granule cells and extremely small pyramidal
cells. The lower border with layer III is somewhat
indistinct.
Layer III is clearly thinner than in areas FA
FC. Its cells are also smaller, and above all, a sublayer IIIc is missing, as individual large pyramidal cells only appear sporadically in its deeper
zone, and in a number insufficient for forming a
continuous secondary sublayer. Sublayer IIIb
contains half as many cells as IIIa. The largest
pyramidal cells only appear isolated in the profundity of sublayer IIIb. Pyramidal cells here are
generally less slender than in the most posterior
segments of the frontal lobe (areas FC, FB and
FA).
Layer IV is rather striking, dense, and uninterrupted, containing small cells. Overall, these
46
are not round granule, but rather small triangular cells. Both the upper and the lower border of
layer IV are quite sharp.
Layer V can be recognized quite easily by the
fact that it is somewhat lighter, especially in its
lower zone, whereas its upper parts contain some
2.5 times as many cells; the cells of the upper sublayer Va are mostly small and somewhat slender
pyramidal; those of the deeper sublayer Vb are
somewhat larger.
Layer VI discloses a somewhat denser arrangement of spindle (fusiform) cells in its upper
sublayer VIa than in the areas described earlier;
consequently, it can be well differentiated from
the somewhat lighter layer V. The deeper sublayer
VIb only contains half as many cells; these are
also smaller in size. The upper border of layer VI,
as well as its lower border vis--vis the white matter, is much more distinct than in the previous
areas, as can be seen in figure 15.
Area FD occupies the entire anterior one-third
of the frontal lobe, from the callosomarginal sulcus on the median hemispheric facies to the pars
triangularis (caput) of F3 on the superolateral
hemispheric convexity (fig. 1b, c), such that it
only leaves the polar, orbital, and subrostral segments of the frontal lobe free. In this broad expansion, it often displays structural variations; I
already emphasized that its more caudal segments, as well as those on the superomedial hemispheric edge and on F3, have larger cells than its
rostral segments. Further, area FD modifies its
character at places in such a typical way that such
variations can be justly considered as cytoarchitectonically true independent areas, as I shall explain later.
Function
According to the latest pathophysiologic findings, one must locate in the anterior parts of the
frontal lobe vested by area FD certain intellectual
functions or single components thereof, namely,
functions of attention, will (psychomotor) and

emotion (a more detailed discussion can be found
in our repeatedly cited larger work [Economo
and Koskinas, 1925, pp. 360364], as well as in
the concluding chapter of this book).
Frontopolar Area FE
The frontopolar area FE (area EK 18), which covers the frontal pole like a cap (fig. 1a, b), lies directly in front of FD and differs from it in the
following four characteristics, which all appear
more pronounced (fig. 16; cf. also plate P31 in the
Atlas [Economo and Koskinas, 2008]): a typical
posteroanterior attenuation of cortical thickness, a reduction in cell size, a sharp demarcation from the white matter, and progressively
more conspicuous granular layers. Nonetheless,
the transition between areas FD and FE is only
gradual.
There seems to be a relation between the size
of gyri and the extension of this field, as area FE
only occupies the slimmer gyri. It is above all layer III that contributes to the overall reduction in
cortical thickness; accordingly, layers II and IV
become more noticeable, with layer II being relatively thicker. Particularly around the frontomarginal sulcus, the cortex exhibits a slight radial striation (not discernible in fig. 16).
Cortical thickness averages about 2.4 mm. I
already mentioned in the introductory chapter
that this thin cortex, which covers the narrow
gyri of the frontal pole (cortical structural type 4,
fig. 8, 9), presents many structural similarities to
the thin cortex of the equally narrow gyri of the
occipital pole. The most important difference between the two poles is that layer V in the frontal
pole is generally characterized by larger, more robust, and more numerous pyramidal cells, in
contrast to the occipital pole, where layer V mostly contains extremely small cells, while larger
cells are only seen sporadically. On the other
hand, the radial striation of the cortex and the
Frontal Lobe
density of both granular layers are far more distinct in the occipital lobe as opposed to the frontal. Those differences become evident if one compares figure 16 with figure 38.
Layer I in area FE is relatively thick and poor
in cells (table 6).
Layer II is more distinct and denser than elsewhere in the frontal lobe, and contains granule
and extremely small pyramidal cells. This layer
stands out well from the other layers.
Layer III in this area is relatively thin, without
a deep sublayer IIIc of large cells; it forms a single
pyramidal cell layer, with cells in sublayer IIIa being almost as large as those in sublayer IIIb.
Layer IV is distinctly separated from layer III
above and layer V below; it becomes more conspicuous than the layers already mentioned, owing to its cell density. Most of the cells are triangular, but quite small, resembling granule cells
(one can see that difference by comparing layer
IV in fig. 14 and 16).
Layer V has a denser upper and a looser lower
sublayer. Cells are fine pyramidal.
Layer VI is rather thick; its upper sublayer VIa
is 50% richer in cells than its deeper sublayer VIb.
The demarcation from the white matter is rather
sharp, but somewhat less so than in area FD.
On the median hemispheric facies, area FE
also reaches as far as the callosomarginal sulcus,
and occupies the frontal pole like a cap at F1 and
F2, however without covering the anteriormost
part of F3 or reaching as far as the lateral (Sylvian)
fissure, from which it is separated by areas FD
and FF (fig. 1a).
Area FE is not structured identically throughout its extent; the part that covers F1 contains
larger cells than the part covering F2. I already
spoke of the succinct radial striation at the pole.
The cytoarchitectonic variations of this area on
the median hemispheric facies will be discussed
in the latter part of this chapter.
47
Fig. 16. Frontopolar area FE (area EK 18). Anterior third of superior frontal gyrus (F1). Dome at frontal pole. !45.
48
Function
On grounds of the ataxic disturbances resulting
from lesions of the frontal pole and the anterior
orbital surface of the frontal lobe, one may infer
that area FE plays a role in certain static and postural (equilibrium) functions; certain frontopontocerebellar pathways conceivably originate in
this area.
Area of Straight Gyrus (Area Recta) FG
Structured very similarly to area FE and lying directly adjacent to it on the medial side of the orbital surface of the frontal pole, the area of the
straight gyrus or area recta FG (area EK 23) is laurel-leaf shaped and extends to the olfactory sulcus posteriorly, which is covered by the olfactory
root (fig. 1a, b, d, 21). Area FG occupies both walls
of this deep sulcus as well as both of its lips, particularly its medial lip; in that stretch, it becomes
thinner and stretches further back to the straight
gyrus (gyrus rectus) and the median hemispheric facies.
Its structural similarity to area FE rests on the
small size of its cells and the progressive diminution of its thickness to 2.42.0 mm, thus constituting one of the thinnest localities in the cerebral
cortex. However, one should keep in mind that
the largest part of this area occupies the walls of
the olfactory sulcus, where the cortex is normally
very thin. Such a thinning particularly involves
layer III, especially at the dome (fig. 17; cf. also
plates P35, P36 and P41b in the Atlas [Economo
and Koskinas, 2008]). Layer V, on the other hand,
differs from area FE in that its cells are extremely
compacted, assuming a band-like appearance at
the wall, which hints at the proximity of the rhinencephalon (cf. Introduction); layer VI is very
sharply delimited against the white matter.
Layer I is thinner than in the frontopolar field
(table 6); its sparse cells do not reveal any specific
particularity.
Frontal Lobe
Layer II is very distinct and dense, possessing

extremely small pyramidal cellular forms.
Layer III is, as already said, very thin and
rather poor in cells. It does not show a clear subdivision into IIIa and IIIb sublayers based on cell
size; the cells are short and triangular, rather
than slender and pyramidal.
Layer IV has a much lower density of cells
compared to the same layer in the frontopolar
area.
Layer V is relatively thick, and certainly the
most succinct layer of area FG. There are pyramidal cells in its denser and sometimes band-like
outer sublayer Va. The deeper sublayer Vb contains fewer and smaller cells, thus staining somewhat lighter.
Layer VI is thicker at the dome and clearly
thinner towards the wall. It is poorer in cells than
the corresponding layer in areas FAFE , and its
fusiform cells are smaller as well. The demarcation from the white matter is very sharp at the
walls, but substantially less so at the dome.
Internal Area Recta Modification FGi

A particular modification of the general cytoarchitecture of area FG is observed on the median
hemispheric facies, with layer V becoming relatively thicker at the dome than the walls; such a
behavior is the exact opposite of layer III, which
thickens at the walls and thins out at the dome
(table 6). On that basis, one can identify a separate area FG termed internal area of the straight
gyrus FGi (area EK 24) [Economo and Koskinas,
1925, pp. 388389].
Function
The location of area FG in the immediate vicinity
of known olfactory parts of the cerebral hemispheres and the presence of a dense layer V to
the point of frequently forming a true band-like
49
Fig. 17. Area recta or area of straight gyrus FG (area EK 23). Dome of straight gyrus (gyrus rectus) at orbital surface of
frontal pole. !45.
50
stria properties common among many olfactory constituents permit the supposition that
the cortex of area FG bears some distant relation
to the rhinencephalon, sensu lato.
Orbital Area FF and Prefrontal Area FH
At the basal junction of the lateral hemispheric

convexity of the frontal lobe with its orbital surface, the granular orbital area FF (area EK 20, also
written FFg to denote its granularity, cf. plate P32
in the Atlas [Economo and Koskinas, 2008]) lies
immediately in front of area FD , while the frontal
pole, as we already saw, is occupied by area FE
(fig. 1a, b). Area FF then covers the largest part on
the orbital surface of the frontal lobe, while area
FE only vests the polar parts of the orbital surface; the boundary between areas FF and FE usually lies somewhat anteriorly to the transverse ramus of the orbital sulcus. Posteriorly, area FF extends to the anterior ramus of the circular insular
sulcus (fig. 21). Medially, it stretches all the way
to the confines of area FG, i.e. almost to the olfactory sulcus. Thus, area FF covers the entire orbital part of F3, including its pars pretriangularis,
and a small orbital part of F2.
On the median hemispheric facies, inferoposteriorly to the polar area FE , lies the prefrontal
area FH (area EK 25; plate P37 in the Atlas [Economo and Koskinas, 2008]), covering the largest
part of the subrostral surface ventrally to the callosomarginal sulcus and reaching almost to the
olfactory sulcus, insofar as this territory is not already occupied by area FG. Caudally, area FH
stretches almost to the parolfactory field of Broca
(fig. 1b, d, 21).
Areas FF and FH are described together because of their structural similarities; as a matter
of fact, they would have constituted a single cytoarchitectonic entity had area FG not been inserted anteroposteriorly between the two like a
wedge. Accordingly, I only present a single microphotograph for the two of them (fig. 18). A
Frontal Lobe
characteristic of both areas is that, in contrast to

the structure of polar area FE and the anterior
parts of area FD , they show a relative and moderate increase in cell size, as well as a certain cortical thickening. At the same time, the cortex becomes impoverished in cells and less distinctly
stratified. An especially typical change in areas
FF and FH is the thickening of layers V and VI,
whereas layers II and IV not only share the overall cellular impoverishment, but also become
thinner.
Layer I contains somewhat more nerve cells
than usual (table 6).
Layer II varies in thickness, becoming minimal in the agranular caudal part; a similar trend
is found regarding cell content. Cells are of the
smallest pyramidal form.
Layer III increases in thickness compared to
areas FE and FG. However, cell density is quite
low thereby. Cells are rather slender pyramidal.
This layer often comprises acellular patches,
which, although normal, might give the impression of a pathologic alteration [Economo, 1929d].
Sublayers IIIa and IIIb are hardly discerned.
Layer IV varies in thickness rostrocaudally,
becoming close to zero in the agranular parts;
granule cells are very small.
Layer V is relatively well represented; in its anterior parts, this layer is not particularly cellular,
but it becomes denser caudally, especially in the
upper zones, such that in caudal segments one
may differentiate a distinctly denser, band-like
upper sublayer Va from an oligocellular sublayer
Vb.
Layer VI contains fewer, very slender spindle
(fusiform) cells. The border with the white matter
is again imprecise.
Agranular Modifications FF and FH

All these characteristics become even more
marked as one proceeds posteriorly towards the
cortical margin (Rindensaum) against the sub-
51
Fig. 18. Prefrontal area FH (area EK 25). Anterior frontal lobe, median hemispheric (midsagittal or interhemispheric)
facies. Dome dorsally to superior rostral sulcus. !45.
52
stantia perforata; thus, the caudalmost parts in

both of these areas finally become heterotypic
agranular in the modifications respectively
termed agranular orbital area FFa (area EK 21;
shown in plate P33 in the Atlas [Economo and
Koskinas, 2008]) and agranular prefrontal area
FHa (fig. 1a, b, d).
The gradual structural alterations of these
fields do not permit the establishment of a common cytoarchitectonic description applicable to
all of their segments. Besides their caudal agranularity, both areas FF and FH undergo further regional modifications that will be explained later.
On the orbital hemispheric surface, they approach each other within a few mm, as area FH
reaches the olfactory sulcus in caudal sections
(fig. 21). Posteriorly to these two areas, the cortex
begins to pass into the ventromedial cortical
margin against the substantia perforata (fig. 1b).
Function
The anteriormost parts of areas FF and FH may
still belong functionally to centers maintaining
equilibrium. The posterior parts, which have a
markedly developed layer V, probably bear some
relation to the rhinencephalon, sensu latiori.
The Frontal Cortical Margin and Its Ground

Areas
The areas FF, FG, and FH just described extend

almost to the cortical margin of the frontal lobe,
i.e. a large cortical band which, on the orbital
hemispheric facies, thinning progressively and
forming a curved line, passes into the anterior
substantia perforata; in the subrostral region on
the median hemispheric facies, this margin continues into the parolfactory field of Broca, which
in turn, thinning progressively towards the back,
ends in the lamina terminalis (a region that links
the two hemispheres) or, better said, the superfi-
Frontal Lobe
cial gray matter lining of the lamina commissuralis. Ventrally, the cortical margin rolls into the
substantia perforata, and dorsally into the indusium griseum on the external surface (dorsum) of
the corpus callosum (Balkenrcken). The cortical
margin is vested in allocortex [Economo,
1928e].
Thus, on the orbital hemispheric surface, area
FF is bordered posteriorly, in the direction towards the transverse insular gyrus, by the isocortical frontoinsular area FJ (area EK 28) (fig. 1a, d
and 21; cf. also plates P42 and P43 in the Atlas
[Economo and Koskinas, 2008]). The next bordering field caudally is the narrow allocortical
frontal piriform area FK (area EK 29), which covers the lateral olfactory gyrus; it forms the orbital
margin toward the substantia perforata on the
ventral facies of the frontal lobe, lateral to the root
of the olfactory tract (trigonum olfactorium).
On the median subrostral surface of the frontal lobe, area FH is bordered posteriorly by the
isocortical parolfactory area FL (fig. 1b, d), which
vests the parolfactory field (carrefour olfactif) of
Broca. Immediately behind area FL , the cortical
margin is constituted by the allocortical geniculate area FM (area EK 33), which covers the medial olfactory gyrus and extends medially from
the root of the olfactory tract, delimiting the cortical margin from the gray matter of the lamina
commissuralis on the median surface of the subrostral region (fig. 21). In other words, the lamina
commissuralis represents the continuation of the
substantia perforata on the median hemispheric
surface (fig. 1b, d).
Parolfactory Area FL , Geniculate Area FM and

Precommissural Area FN
In the subsequent expos, one should keep at

hand the schematic drawings of the median
(fig. 1b) and orbital (fig. 21) hemispheric facies, as
well as the microphotograph (fig. 19) of the horizontal section through Brocas parolfactory field.
53
Fig. 19. Parolfactory area FL (areas EK 3032). Geniculate area FM (area EK 33). Precommissural area FN (area EK 35).
Horizontal section through the subrostral region at the median hemispheric aspect of the frontal lobe, extending
from the parolfactory field (carrefour olfactif) of Broca to the anterior commissure. Left to right: Co.a., anterior commissure; l.t., lamina terminalis or rostralis, separated from the lamina terminalis of the opposite hemisphere at the
plus sign (+); Ggl.basal., nucleus basalis (ganglion basale of Meynert); Nucl.caudat., internal zone of the head of the
caudate nucleus; asterisk (*), parolfactory granular islands of caudate nucleus. FN, FM , FL3, 2, 1, FHL , FH, cortical areas
of this region. The three vertical arrows beneath the dashed line at the top of the figure respectively denote the area
of the parolfactory sulcus postremus (left arrow), the posterior parolfactory sulcus (middle arrow), and the middle
parolfactory sulcus (right arrow). The subcallosal gyrus extends between the first and second arrows; the geniculate
(or internal olfactory) gyrus, between the second and third arrows. The three vertical arrows above the dashed line
indicate, respectively, the border between the commissural white matter and its gray matter lining (extreme left arrow), the border between the commissural gray matter and the cerebral cortex proper (middle arrow), and the boundary between the parolfactory field and the six-layered isocortex (extreme right arrow), which corresponds to the anterior parolfactory sulcus. !8.
The captions explain the anatomical relations in

detail. Figure 19 is an overview microphotograph
shown at a magnification of !8; thus, it does not
depict cellular details, but rather, the topographic relations of the cortical layers.
54
On the median hemispheric facies (cf. fig. 1b,

19), area FH is succeeded caudally in the subrostral region, separated from it by the shallow vertical depression of the anterior parolfactory sulcus,
by the structure called Brocas parolfactory field
or parolfactory area FL , which extends to the

small vertical, very shallow middle parolfactory
sulcus, followed by the cortical margin of the
frontal lobe.
This margin is formed at this level by a small,
vertically running protrusion, the geniculate gyrus, which represents the continuation of the medial olfactory gyrus proceeding from the orbital
to the median hemispheric surface, i.e. the thin
gray cortical lamella termed geniculate area FM
(area EK 33; plates P40 and P41a in the Atlas
[Economo and Koskinas, 2008]) that follows the
medial olfactory root on the median surface of
the cerebral hemisphere.
This is followed caudally, eventually separated
by some shallow depressions of the posterior
parolfactory sulcus, by a thin lamella of cerebral
matter that links the two hemispheres (lamina
commissuralis); the gray matter of this lamella
often becomes thickened to form a small vertical
short bulge especially directly under the rostrum and on the lower (inferomedial) hemispheric edge that we call subcallosal gyrus, and whose
caudal slope may sometimes be delimited by a
shallow incisure (notch) that one might call parolfactory sulcus postremus (= hindmost; the
superlative of the Latin adjective posterus, the
comparative being posterior). Essentially, this
formation of the lamina commissuralis, or precommissural area FN (area EK 35; plates P40 and
P41a in the Atlas [Economo and Koskinas, 2008]),
no longer belongs to the cerebral cortex sensu
stricto, but rather corresponds to the fusion point
(Verwachsungsstelle) or junction (Verbindungslamelle) of the two anterior cerebral vesicles (Vorderhirnblschen), where the commissures, and
especially the corpus callosum, are formed. As
already mentioned, its gray matter continues further dorsally into the indusium griseum on the
dorsum of the corpus callosum (Balkenrcken);
on the inner side of the corpus callosum, it continues into the gray matter of the septum pellucidum; ventrally, this area rolls into the substantia perforata.
Frontal Lobe
In this entire territory, from the anterior parolfactory sulcus and moving anteroposteriorly,
the cortex continually diminishes in thickness;
however, the involvement of the individual layers
in such a diminution varies greatly. This phenomenon can best be seen in figure 19: on the
right side of the figure is Brocas parolfactory
field, on the left the white matter of the anterior
commissure, and in between the geniculate and
subcallosal gyri, and the lamina commissuralis
(rostralis or terminalis) sectioned at the midline.
To better convey the continuous, gradual
change of cytoarchitectonic cortical structure in
this region, I discuss these three areas, which
form a rostrocaudal continuum (in the order FL ,
FM, FN, cf. fig. 19) jointly. On account of the stepwise change of cellular composition in area FL ,
we further subdivide it into primary, secondary
and tertiary parolfactory area modifications FL1,
FL2 and FL3 (areas EK 30, 31 and 32, respectively;
cf. plates P39P41 in the Atlas [Economo and
Koskinas, 2008]). Moreover, the transition modification from area FH to area FL is termed parolfactory prefrontal area FHL (area EK 26; cf.
Economo and Koskinas [1925, pp. 396405] and
plate P39 in the Atlas [Economo and Koskinas,
2008]).
Layer I, in contrast to the other layers, actually becomes thicker and more cellular through
the entire extent of Brocas parolfactory field,
particularly in the caudal direction (from right to
left in fig. 19) and also distinctly towards the cortical margin; the tangential (myelinated) fibers in
its uppermost zone are increased to such a degree
that the cortical surface of the caudal parts assumes a whitish appearance (this is discerned
from the numerous neuroglial nuclei present in
layer I in fig. 19). As the other layers progressively thin, layer I is finally the only one remaining,
rolling alone into the lamina terminalis, rostralis
or commissuralis, i.e. the ventral fusion point of
the two cerebral hemispheres. In the middle
parolfactory sulcus, at the posterior boundary of
55
Brocas parolfactory field (area FL1, 2, 3) and the

anterior boundary of the geniculate gyrus (area
FM), layer I sends a wedge to the depths, at which
layers II, III, and the offshoot of layer IV seem to
end (visible in fig. 19 as a smaller spur issued
from layer I and directed from above left to the
depths).
Layer II, as mentioned earlier, diminishes in
thickness and cellularity in the posterior direction, already at the level of area FH; at the level of
area FL1 this layer has thinned considerably and
become impoverished in cells, peculiarly frayed,
and indistinctly delimited from layer III beneath.
However, the cells are somewhat larger than usual and have an irregular stellate-shaped form.
Layer II can hardly be recognized at the caudalmost parts of area FL2, and its actual locality is
only indicated by single round cellular conglomerates, which finally disappear, in modification
FL3, at the wedge of layer I of the middle parolfactory sulcus mentioned above.
Layer III, distinguished in area FH by its
thickness, first becomes slightly thicker in Brocas parolfactory field FL1, but then rapidly diminishes caudally within the modification FL2,
such that its lower border seems to approach
more and more the cortical surface. Layer III
then disappears abruptly in the modification FL3
and directly in front of the geniculate gyrus at the
aforementioned wedge of layer I. In the passage
from area FH caudally towards the middle parolfactory sulcus, layer III does not only become
progressively thinner, but also less cellular, even
completely acellular at places (these details can be
seen in fig. 19); its pyramidal cells, which were
originally long and slender, gradually become
smaller and less conspicuous in areas FL2 and
FL3.
Layer IV becomes, as already mentioned, continually thinner and less cellular in area FH caudally, and completely disappears within area FL .
Layer V, which in area FH was subdivided into
an upper sublayer Va with smaller but more compactly disposed cells, and a lower sublayer Vb
56
with larger but less densely packed cells, displays

more distinctly such a subdivision into two sublayers, as one passes caudally; thus, sublayer Va
appears strikingly as a compact, dark band in the
posterior part of area FL2, whereas the deeper
sublayer Vb in this region characteristically contains less densely packed, but particularly slender, lancet-shaped cells (Lanzettfrmige Zellen)
with long, whip-like upper dendrites. In the transition to the geniculate gyrus, i.e. in area FM,
which is separated from area FL3 by the wedge of
layer I of the middle parolfactory sulcus mentioned earlier, in which layers II, III (and IV) end,
layer V and almost exclusively its deeper sublayer Vb forms a loop about the bottom of that
wedge, and then passes over alone into the now
very thin cortex of area FM. It is found directly
subjacent to the single layer I, consisting of rather
disorderly cell conglomerates.
Layer VI cells, besides layer I and sublayer Vb,
also penetrate area FM under this wedge; that peculiar cellular compilation without a regular
lamination presents as allocortex, while area FL is
still heterotypic isocortex. In passing from area
FH over Brocas parolfactory field to the cortical
margin, layer VI also becomes considerably diminished in thickness, and yet it is still comparatively compact at the posterior parts in modification FL2 and rather sharply demarcated against
the white matter. In area FM the cells of layers V
and VI are so intermixed that one cannot discern
any lamination; neither is it advisable to attempt
to draw any exact homologies with the cells or the
layers of the isocortex. At the level of the posterior parolfactory sulcus, cells disappear altogether.
The ventral confines of areas FL and FM,
which become markedly thinner on the median
hemispheric surface (constituting there the caudal extreme of area FH, cf. fig. 1b, d), even reach
all the way to the olfactory sulcus on the orbital
surface (fig. 21); at the posterior extreme of that
sulcus, at the root of the olfactory tract, one finds
a small gray eminence, the trigonum olfactori-
um, which is vested by the cortical formation

called geniculate area of olfactory triangle FMt
(area EK 34), which essentially represents a very
slight modification of area FM (cf. plate P41b in
the Atlas [Economo and Koskinas, 2008]).
The precommissural area FN, caudal to area
FM, only consists (fig. 19) of a very thick layer I,
beneath which one finds some irregularly formed,
strikingly large, and also smaller polygonal stellate cells, forming small ganglionic nodules.
These cells are hardly related to the cells of area
FM, and not considered to be true cortical elements. Rather, they represent cells of the so-called
nucleus basalis ( ganglion basale of Meynert
[1872a]), which is located dorsally and anteriorly
to the optic tract at the base of the entire (smaller)
medioposterior part of the anterior substantia
perforata, which is covered by the diagonal band
of Broca (fig. 21); the nucleus basalis passes from
the ventral to the median surface with the stylum
septi, reaching there as far as the subrostral region, and then continuing with the septum pellucidum and the indusium griseum. In the depths
of the cerebral tissue appears the head of the corpus striatum (marked by the asterisk in fig. 19).
Frontoinsular Area FJ
On the median hemispheric surface, the cortex of

area FH continues, through a gradual anteroposterior thinning towards the still isocortical area
FL (fig. 1b, d), into the cortical margin of the
frontal lobe (area FM); in a like manner, on the
ventral (or orbital) surface of the cerebral hemisphere, the similarly structured area FF continues, through a thinning towards the still isocortical area FJ (fig. 21), into the cortical border of the
frontal lobe (area FK ), which separates it from the
substantia perforata. (Fig. 21 shows the topographic relations in a section from the anterior
part of the orbital cerebral surface, around the
substantia perforata; gross anatomic relations are
marked on the left side of the figure, which is a
Frontal Lobe
semi-schematic drawing of the right cerebral

hemisphere, and cytoarchitectonic area boundaries are marked on the right side of the figure,
which depicts the left cerebral hemisphere.)
This transition usually occurs on a transverse,
small bridging gyrus, which reaches from the
medial extreme of the orbital part of F3 (laterally
at the insertion of the root of the olfactory tract)
to the pole of the insula: the so-called transverse
insular gyrus. Its anterior wall concomitantly
forms the anterior, i.e. frontal wall of the insular
pyramid. It is vested in a peculiar, heterotypic isocortex, which constitutes the frontoinsular area
FJ (area EK 28).
Rostrally, area FJ is bordered by area FF on F3.
It continues caudally into the narrow, allocortical,
gray cortical margin which accompanies the lateral root of the olfactory tract, i.e. the so-called lateral olfactory gyrus or frontal piriform area FK
(area EK 29). This small gyrus represents the termination of the cerebral cortex and forms the transition to the substantia perforata (which, in the human brain, does no longer belong to true cortex).
Posteriorly to area FF, the cortex does not thin
immediately, but rather becomes initially enlarged in area FJ all the way to the dome of the
transverse insular gyrus; only then does it diminish rather rapidly in thickness caudally, to stop
entirely at the posterior boundary of the lateral
olfactory gyrus, where it becomes reduced to
solely a layer I, apparently continuing alone into
the anterior substantia perforata.
In addition to its general thickening to 3.5
mm, the cortex of area FJ undergoes some further appreciable changes (cf. fig. 20 and upper left
corner of fig. 22).
Layer I progressively increases in thickness
and cellularity as it courses caudally (table 6); in
its posteriormost parts, the myelinated fibers are
substantially increased beneath the cortical surface, imparting a white shimmering appearance
to sublayer Ia.
Layer II, appearing already thin and impoverished in cells in the posterior parts of area FF (i.e.
57
Fig. 20. Frontoinsular area FJ (area EK 28). Sagittal section through the dome of the transverse insular gyrus at the
caudal (posterior) boundary of the orbital surface of the frontal lobe. Right side of the figure is rostral, left is caudal.
!45.
58
Fig. 21. Schematic drawing of the region of the anterior substantia perforata. The left side of the figure depicts the
substantia perforata and its immediate vicinity; the right side of the figure shows cytoarchitectonic area boundaries
on the ventral aspect of the frontal pole. The temporal poles have been dissected out bilaterally, in order to expose
the entire orbital surface. Abbreviations on the left: Ch., optic chiasm; em.pro., parolfactory eminence or olfactory tubercle, derived from the head of the caudate nucleus; F1 and F3, superior and inferior frontal gyri; g.olf.l., lateral olfactory gyrus; g.olf.m., medial olfactory gyrus; g.r., gyrus rectus (straight gyrus); g.tr.is., transverse insular gyrus; l.t., lamina terminalis; pl.spt., septal plane, issued from the nucleus basalis; s.d., diagonal sulcus of substantia perforata; s.p.,
substantia perforata; s.pro.ps., parolfactory sulcus postremus; str.olf.l. and m., lateral and medial olfactory striae; str.
pro.hip., hippocampal parolfactory stria; styl.spt., septal stylum (diagonal band); T., dissected temporal pole; Tr.olf.,
olfactory tract; Trig.olf., olfactory triangle (tuber olfactorium). Abbreviations on the right: FF FN, frontal lobe areas; IC
and ID , insular lobe areas; TJ and TK , temporal lobe areas.
in modification FFa), becomes even thinner in

area FJ with a marked reduction in cell number;
however, its cells are larger and here, in the caudal parts of FJ, one can even observe quite large,
irregularly formed, stellate-shaped cells (clearly
visible on the upper left side of fig. 20). Here also,
layer II has a fringed appearance and reveals a
tendency in caudal parts for glomerulous cellular
accumulations (a characteristic of rhinencephalic formations [Economo, 1928e]).
Layer III thickens appreciably in area FJ, but
its cell content is reduced, such that oligocellular
patches appear occasionally. The cells themselves
are no longer fine pyramidal, but peculiarly elongated spindle (fusiform) cells, such that one might
Frontal Lobe
speak here of an actual cellular fusiform transformation (Verspindelung).

Layer IV, which also displayed acellular patches in the posterior parts of area FF (i.e. in modification FFa), is lost completely in area FJ, especially at the dome of the transverse insular gyrus;
as a result, layer III here borders immediately on
layer V. On the right (anterior) side of figure 20,
layer IV is still identifiable, but on the left (posterior) side of the figure it is missing.
Layer V in this region is also subdivided into
an almost band-like upper sublayer Va, with
small, compact cells, and a lower sublayer Vb
with larger cells, but only half as many (table 6).
The tendency of cells in upper sublayer Va to be
59
packed more densely, forming a cellular band,

becomes readily discernible at more caudal parts
of area FJ (i.e. on the left side of fig. 20); such a
tendency is typical for regions in the vicinity of
the rhinencephalon.
Rod and Corkscrew Cells. The cells of the
deeper sublayer Vb are absolutely spindle-shaped
with regard to their vertical direction, and mostly form very long rod-like and also often spirally
twined elements, extremely characteristic of this
region, that we term rod (Stbchen) and corkscrew
cells (Korkzieherzellen) [Economo and Koskinas,
1925; Economo, 1926d]. The largest of them may
reach 70100 m in height (cf. Introduction).
Such cells are specific for this area, as they are not
found anywhere else in the entire cerebral cortex,
except, as we shall see later, in the anterior segment of the superior limbic (cingulate) gyrus.
One can see these cells on the left side of figure
20, directly to the upper right of the indication
Vb, very clearly and in large numbers; from here
they can be followed in this layer through the entire microphotographic field. Thus, there is in
layer V a veritable fusiform transmutation of cells
that is even more pronounced than in layer III
[Economo, 1927d].
Layer VI, finally, which is also quite enlarged,
but somewhat poorer in cells than usual, displays
the overall cellular elongation of this region as
well, although to a moderate measure. The delimitation against the white matter is exceptionally indistinct, and the cells of this layer become
at places dispersed into the profundity, where
they meet with the cellular streaks of the claustrum and the substantia perforata.
Frontal Piriform Area FK and Area of

Substantia Perforata TK
As already mentioned (fig. 21), the cerebral cortex on the orbital surface rolls anteroposteriorly,
across the transverse insular and lateral olfactory
gyri (areas FJ and FK , respectively), into the sub-
60
stantia perforata (cf. also plate P41c, d in the Atlas

Figure 22 which represents a somewhat posterior midsagittal section through the substantia
perforata laterally to the olfactory tract illustrates in a clear way the cytoarchitectonic characteristics of this region. On the far left side of the
figure, one still sees the dome of the transverse
insular gyrus. Immediately to the right, i.e. moving caudally, one finds the allocortex of the lateral
olfactory gyrus (area FK ), rapidly thinning on the
gradually descending wall of the gyrus and becoming pointed. At the extreme brink on the far
right side of the figure, i.e. further caudally, we
note a cellular disposition, whereby the large, stellate-shaped cells of the nucleus basalis, organized
in groups as we observed in figure 19, are covered
by a thick layer I, the so-called precommissural
area FN (on the surface, this area corresponds to
the planum septale of the substantia perforata
marked pl.spt. in fig. 21). Still further caudally
(which would fall outside the figure to the right),
we should next encounter in cross-section the
myelinated fiber bundle of the optic tract.
Between the nucleus basalis (FN ) and the lateral olfactory gyrus (FK ), one sees in the same
figure a large formation (TK ) advancing towards
the cortical surface; it consists of groups of small
granule cells of a round or streaky form, reaching
very close to the surface, and it is only vested by
a continuation of layer I. This in turn appears
crisscrossed with lacunas, which give way to larger vessels toward the depth. This segment of the
substantia perforata tallies with the head of the
caudate nucleus, which, at this point, almost
reaches the surface, and protrudes, in the human
brain, as the olfactory tubercle (or parolfactory
eminence), filling the entire anterolateral territory of the substantia perforata between the frontal, insular and temporal lobes (labeled em.pro. in
fig. 21). On the cortical surface, reckoning this
formation as belonging to the areas of the temporal lobe, we termed it area of substantia perforata
TK (area EK 93).
Fig. 22. Frontoinsular area FJ (area EK 28). Frontal piriform area FK (area EK 29). Area of substantia perforata TK (area
EK 93). Precommissural area FN (area EK 35). Sagittal section through the anterior substantia perforata. Left to right:
transverse insular gyrus and its area FJ ; continuing backwards (i.e. to the right), lateral olfactory gyrus with area FK ,
where the cortex rapidly diminishes in thickness; Cl., last cellular traces of claustrum, reaching as far as the deeper
layers of the cortex of the lateral olfactory gyrus; Nucl.caudat., head of caudate nucleus, reaching the surface behind
the lateral olfactory gyrus (area FK ), which is denuded of cortex in this expansion, the head of the caudate forms
the anterolateral limit of the substantia perforata (area TK ); behind this area (to the right), the uninterrupted continuation of the nucleus basalis, which, along with its gray matter lining, forms the posteromedial limit of the substantia perforata (area FN ); finally, directly behind it, where the figure ends, the optic tract is found, in the angle of
which lie the cells of the supraoptic ganglion (not visible in this figure). !10.
In all, the substantia perforata comprises two

distinct areas (right side of fig. 21): the anterolateral area TK , in close association with the head of
the caudate nucleus, and the posteromedial area
FN, which shows a close association with the nucleus basalis, bordered by the optic tract, and extending with the stylum septi (marked styl.spt. in
fig. 21) of the diagonal band from the basal (or-
Frontal Lobe
bital) surface to the median hemispheric surface

and the subcallosal gyrus.
If we now return to the microphotograph of
the section through the parolfactory field on the
median hemispheric facies (fig. 19), we also see
here that, in the deeper parts of the tissue between the medial olfactory (FM) and the subcallosal (FN ) gyri, there are groups and streaks of
61
small cells (denoted by the asterisk in fig. 19) still

belonging to the head of the caudate nucleus,
which, however, at this point, i.e. the median surface, does not reach the surface yet, and thus does
not yet separate areas FM and FN from each other in the form of a wedge, like it does on the orbital cerebral surface (areas FN and FK ).
Let us now take another look at the individual
layers of the substantia perforata (fig. 22). At the
posterior part of the transverse insular gyrus
(right side of the figure), the cortex of area FJ begins to thin out.
Layer I thickens in the region of the lateral olfactory gyrus (frontal piriform area FK), and discloses at the surface numerous glial nuclei disposed in horizontal rows, suggesting the presence of numerous myelinated fiber bundles,
which impart in unstained specimens a whitish
appearance to this cortical margin. Those myelinated bundles, which consist of olfactory nerve
fibers, penetrate layer I and the rest of the cortex
at different places towards deeper regions of the
tissue (denoted by the plus sign in fig. 22).
Layer II becomes impoverished in cells; nonetheless, their size increases and they are grouped
into looser clusters, such that a distinctly continuous layer can no longer be discerned.
Layer III is equally poor in cells, revealing numerous irregular acellular patches; it is immediately lost at the level of the entrance of the olfactory fiber bundles.
Layers V and VI, whose elements are indistinguishable from each other, because of their reduced size and loss of typical form, also end
somewhat more posteriorly, at the level of entry
of the olfactory fibers. At many places, the cells of
layers V and VI are displaced by the incoming
myelinated fiber pathways into the depths of the
tissue, where they meet and become mixed with
the cells of the claustrum.
The cortex of the lateral olfactory gyrus just
described (FK ) does no longer display the typical
lamination; it belongs to allocortex, and a drawing of any homologies of its cell layers to those of
62
the isocortex is virtually impossible. We adopt the

layer designations II, III, etc. for cellular ensembles, only insofar as the cells themselves present
an apparent cohesion with those of the isocortex.
At the posterior boundary of the lateral olfactory gyrus, the cellular gray cortex ends completely; the shallow groove that separates it from
the substantia perforata is the continuation of
the posterior parolfactory sulcus of the median
hemispheric facies. This groove is only vested in
a layer I (in the human brain); beneath it, one encounters the cellular compartments and clusters
of the caudate nucleus, and further posteriorly,
the large stellate-shaped cells of the nucleus basalis. The shallow incisure between these last
two structures is called diagonal sulcus (marked
s.d. in fig. 21); medially, it passes into the posterior parolfactory sulcus. Further posteriorly, the
sulcus that delimits the septal plane (nucleus basalis, FN ) from the optic tract is the continuation
of the parolfactory sulcus postremus on the basal (orbital) cerebral facies (marked s.pro.ps. in
fig. 21).
Variant and Modification Areas of the Frontal

Lobe
In the preceding part of this chapter, I covered

the typical cellular structure of all the main areas
of the frontal lobe, beginning at the central sulcus, over the frontal pole, including the base of
the frontal lobe and the cortical margin at the
level of the substantia perforata and lamina commissuralis.
I described in detail the progressive decrease
in overall cortical thickness and size of pyramidal and spindle (fusiform) cells, as well as the
concomitant progressive reinforcement of both
granular layers, proceeding from the agranular
regions proper in the precentral region towards
the frontal pole; a certain reversal of many of
those parameters is observed from here towards
the more caudal regions at the base (orbital surface) of the cerebral hemispheres, all the way to
the cortical margin.
We sequentially divided this entire territory of
the frontal lobe into the corresponding areas,
from FA to FN, following the typical changes in
the overall cortical picture. However, it would be
erroneous to imagine that the manifold cortical
characteristics can be exhausted in such a few areas, particularly with regard to the major fields,
which occupy entire segments of the superolateral
hemispheric convexity overriding sulci and gyri.
I occasionally mentioned that these areas
display various regional modifications in their
structure. For example, regions in the vicinity of
the superomedial hemispheric border contain
larger cells than more ventral regions; also, caudal segments of each area usually contain larger
cells than anterior segments; further, all the areas
on F3 are characterized by large pyramidal cells,
and much more.
In addition, most areas (not all!), at the transition to a neighboring area, display a shorter or
longer stretch of a mixed cytoarchitectonic type,
with intermediate character marks between the
two fields. For example, we can distinguish a
faintly granular transition type between areas FB
and FC, which we call agranular-intermediate
transition frontal area FBC (cf. plate P9 in the Atlas [Economo and Koskinas, 2008]); the same applies to other transition points.
As already mentioned, we denote magnocellular and parvicellular area modifications with
the suffices m and p, respectively. Thus, in ground
area FD , we define a magnocellular granular frontal area FDm (area EK 12) and a parvicellular
granular frontal area FDp (area EK 13) (fig. 1a, c,
d; cf. also plates P19P23 in the Atlas [Economo
and Koskinas, 2008]). Transition modifications
of the area FDm that include an admixture of
structural elements characteristic of the adjacent
intermediate frontal area FC and frontopolar area
FE are denoted as the magnocellular granular
frontal area at the beginning of the intermediate
Frontal Lobe
frontal area FDm(C) and the magnocellular granular frontal area at the beginning of the frontopolar
area FDm(E), respectively (table 2; cf. also plates
P19 and P21 in the Atlas [Economo and Koskinas, 2008]).
Moreover, within an area, certain structural
variations recur consistently, depending on the
precise location within each gyrus; e.g. a certain
reinforcement of the granular layers is regularly
noted at the walls of gyri, even at loci where the
cortex is otherwise agranular. This is the case in
the posterior wall of the precentral gyrus in the
central sulcus of Rolando, which, although belonging to the agranular area FA, nnetheless
suggests the presence of the two granular layers;
the same is observed in the paracentral lobule
(fig. 12). Area FB is another example of a similar
situation.
Besides such general modifications in each
ground area, there are additional circumscribed,
typical structural alterations, which, owing to
their constant occurrence at the same site, might
conceivably justify their consideration as independent areas; to avoid complicating our exposition even further, we shall view them merely as
variants or modifications of the areas that we already know. Thus, I already described a distinct
modification in the more caudal and dorsal part
of area FA, characterized by the presence of giant
cells of Betz, which we call giant pyramidal precentral area FA (area EK 2).
Opercular Precentral Area FAop
The structure of area FA also becomes significantly modified at the inferior extreme of the
precentral gyrus, i.e. at the level of the operculum. Giant pyramidal cells of Betz are no longer
present at this point. The cortex diminishes considerably in thickness; its cells, particularly in
layer III (and in other layers as well) become
sparser, smaller, less finely formed, and lose, for
the most part, their otherwise pretty, vertical ra-
63
dial orientation toward the surface, such that the

cytoarchitectonic picture assumes a peculiarly
untidy aspect. Nor is the cortex of area FA any
longer truly agranular in the opercular region.
We denote this modification as opercular precentral area FAop (area EK 3). The degree to which
these characteristics develop and the magnitude
of their territorial span are subject to fluctuations
among individuals.
Opercular Agranular Frontal Area FBop
Area FB , which only extends as such at the level

of F3 on the precentral gyrus (fig. 1a), undergoes
an analogous structural regression near the inferior edge of the operculum, as typified in figure
23; this opercular agranular frontal area FBop
(area EK 5) can thus be viewed as a sample of this
kind of opercular cytoarchitectonic modification. On the contrary, the internal wall of the
operculum is usually clad in the classical FB formation. A comparison of area FB in figure 13
with area FBop in figure 23 shows very clearly the
attenuation of the entire cortex in the latter, the
decrease in the size of cells, their disorderly arrangement, the accentuation of layers II and IV,
and the loss of the orderly cellular disposition in
layer III, in sublayer IIIc, plus other factors.
The transition points between the opercular
segments of the agranular frontal area FB and the
intermediate frontal area FC are denoted as (intermediate) agranular frontal area in operculo FB(C)op
or opercular agranular intermediate frontal area
FBCop (table 2; depicted in plates P10 and P11 in
Limbic Intermediate Frontal Area FC L
Within ground area FC, we can distinguish two

further structural variations which can be differentiated as true individual areas, namely: (1) the
variant area FC L (discussed under this subhead-
64
ing), and (2) the modification area FCBm (detailed

under the next subheading).
The first one is found on the median hemispheric facies, in the region where area FC touches
the callosomarginal sulcus and also vests the posterior part of the frontolimbic transition gyrus.
The cellular structure of area FC becomes modified not only by becoming somewhat thinner
overall, but also by assuming the typical characteristics of the cortex of the superior limbic gyrus
that I detail in a subsequent chapter: such characteristics include an extremely marked increase of
cell density in layer V, especially its upper parts,
whereby such a sublayer Va almost becomes the
most striking cellular zone in the entire section (a
feature that witnesses the contiguity of the rhinencephalon [Economo, 1927d, 1928e]). This cytoarchitectonic change, which, as we shall see later,
recurs in all frontal areas that verge on the median hemispheric surface (fig. 1b), justifies by way
of its constancy the identification of this part of
area FC as a separate variant, the limbic intermediate (or intermedio-limbic) frontal area FC L (area
EK 7). I merely offer an example of the analogous
variant of area FD , i.e. the limbic granular frontal
area FD L (area EK 15), in the last microphotograph of this chapter (fig. 27; also depicted in plate
P17 of the Atlas [Economo and Koskinas, 2008]).
Magnocellular Agranular Intermediate

Frontal (Brocas) Area FCBm
Another, much more conspicuous variation of

area FC is Brocas area, found at the foot of F3
(pF3), i.e. the pars orbitalis (fig. 1a, 24). It is also
vested in a somewhat granular, i.e. intermediate,
magnocellular formation; but at this level the
cortex of area FC thickens to a certain extent and
assumes a fan-like, striated appearance; it possesses larger and more abundant cells (compare
fig. 14 with fig. 24). Moreover, the cells are disposed in vertical columns to the cortical surface,
more distinctly than elsewhere in area FC.
Fig. 23. Opercular agranular frontal area FBop (area EK 5). Operculum of Rolando, precentral gyrus, dome. !45.
Frontal Lobe
65
Fig. 24. Magnocellular agranular intermediate frontal (Brocas) area FCBm (area EK 8). Brink of the foot of inferior frontal gyrus (pF3). !45.
66
Layer III especially displays such a perpendicular columnar organization. It can be subdivided
into sublayers IIIa, IIIb and IIIc (table 6). The
cells of the deepest sublayer IIIc form a distinctively singular row of extremely large, slender pyramidal cells, which at certain places almost represent giant cells, attaining a height of 4570 m
and a width of 25 m.
Layers II and IV, on the other hand, are thinner, but much denser and more visible than in all
the other fields of area FC.
Layer V also contains large pyramidal cells
that are quite large (much larger than the cells of
layer VI, cf. table 6). One may also separate a
lighter, deeper sublayer Vb.
Layer VI is clearly pronounced and thick, but
without any characteristic particularities.
The altogether unusual size of the pyramidal
cells in this area, which sometimes even exceed
the pyramidal cells of area FB at the superomedial hemispheric edge, as well as the radial striation and the thickness of the cortex, all recall the
directly adjacent area FB ; however, the explicit
presence of both granular layers demonstrates
that this cortex appertains to the ground area FC.
We thus view this area as a transition structure
designated FCB , to which we add the suffix m owing to its magnocellularity, and accordingly call
this typical modification magnocellular agranular intermediate frontal area FCBm (area EK 8) or
Brocas area2 (cf. plates P14 and P15 in the Atlas
Function
Brocas area constitutes the specific structure
of the motor speech zone. It is known that not
only Brocas field, but the entire F3 with its three
parts opercular, triangular and orbital is a re-
2
It appears occasionally written FBCm in the larger work
[Economo and Koskinas, 1925, pp. 308332].
Frontal Lobe
cent evolutionary acquisition of the human brain.

This gyrus does not appear in the animal series;
at best, there are indications that it is found in the
young brain of the Orangutan.
Triangular Granular Frontal Area FD
Further rostrally, the cortex shows a tendency to

produce particularly large pyramidal cells in the
pars triangularis or so-called head (caput) of
F3 situated directly in front of its foot which,
as we saw earlier, belongs to the domain of the
homotypic, distinctly granular area FD. And yet,
within that ground area, it shows such a typical
and constant structural modification, that we
must designate it as a separate variant, which we
call triangular granular frontal area FD (area EK
17) (fig. 1a, c, d). The primary difference from the
structure of the rest of area FD is a thinning of its
cortex to 2.5 mm or even less (fig. 25; also plates
2008]); another, the striking fan-like radial striation of the cortex. Moreover:
Layers II and IV both appear robust, imparting a succinct horizontal lamination to the cortical cytoarchitectonic picture.
Layer III presents a distinct deep sublayer
IIIc, a phenomenon unusual for area FD , which
consistently contains larger, triangular pyramidal cells, laden with Nissl bodies. These pyramidal cells reach nearly giant cell size, i.e. 3060 m
in height and 2540 m in width (fig. 15, 25).
Layer V is lighter, especially in its deeper sublayer Vb, which also contains fewer cells, another
typical feature of this area.
Layer VI is well demarcated from the white
matter.
The transition from the thick cortical structure of area FCBm at pF3 to the thinner cortical
structure of area FD at the pars triangularis (or
caput) is very abrupt at the dorsal slope of the
pars ascendens of pF3, thus constituting one of
few truly sharp area boundaries.
67
Fig. 25. Triangular granular frontal area FD (area EK 17). Brink of pars triangularis of inferior frontal gyrus (F3). !45.
68
Function
Herv [1888a, b] had already noticed in 1888 the
giant size cells at the foot (pes) and head (caput)
of F3. The caput also has a likely function in the
motor speech mechanism of the cortex. Henschen [1918] places his song center (Gesangzentrum) in the pars triangularis.
Middle Granular Frontal Area FD
Dorsally to the caput of F3, but still within the

field of area FD on F2, another typical variant is
found (fig. 1a, c), which shows exactly the opposite structural particularities to those of the variant FD just described. We call it middle granular frontal area FD (area EK 16) (fig. 26; also
2008]). In this circumscribed area, the frontal
granular cortex becomes thicker; its cells are
more numerous, more tightly packed, and smaller in size.
Layers II and IV thicken considerably and appear denser in cells than anywhere else in the
frontal lobe.
Layer III contains at best medium-sized pyramidal cells, but they are more numerous and
densely placed than elsewhere; there is no sublayer IIIc.
Layer V is equally thick and abundant in cells;
it does not comprise the two secondary sublayers
that are seen in the rest of area FD.
Layer VI has an uncertain border against layer V.
On the basis of all these structural particularities, one remembers the parietal structural
type of cortex; thus, area FD represents cortical
structural type 3, whereas all the remaining anterior frontal lobe belongs to either structural
type 1 or type 2, as I mention in the introduction
(fig. 9a). This area FD merely covers 2 or 3 relatively wide secondary gyri of F2 immediately dorsally and anteriorly to the caput (fig. 1a, c). It is
Frontal Lobe
entirely surrounded otherwise by the typical cortex of area FD , and as a matter of fact by its magnocellular modification FDm dorsally and posteriorly, and by its parvicellular modification FDp
anteriorly and ventrally.
Function
The physiologic significance of this area remains
unknown, although it is interesting to recall that
this is where Flechsig [1895] places his anterior or
frontal association center, in a region surrounded
on all sides by the cortical structural type 2, and
constituting an isle of parietal structural type 3
that becomes lost anteriorly toward the frontal
pole; a similar cortical structural type 3 is precisely the one corresponding to the inferior parietal territories where Flechsig [1895, 1896a, 1896b]
places his large, posterior association center. There
is little doubt that such an analogy is incomplete,
but nonetheless sufficiently conspicuous to be
underscored.
Limbic Granular Frontal Area FD L
Area FD further presents on the median hemispheric facies the typical increase in cell density
in the upper zone of layer V that witnesses the
proximity of the limbic gyrus. This characteristic
becomes accentuated at the point where area FD
vests the anterior frontolimbic transition gyrus
between the limbic sulcus and the callosomarginal sulcus (fig. 1b). We call this variant limbic
granular frontal area FD L (area EK 15). Figure 27
(and plate P26 in the Atlas [Economo and Koskinas, 2008]) illustrates this cytoarchitectonic type,
which is equally typical of the earlier mentioned
area FC L.
69
Fig. 26. Middle granular frontal area FD (area EK 16). Dome of the middle third of middle frontal gyrus (F2). !45.
70
Fig. 27. Limbic granular frontal area FD L (area EK 15). Fold (pli) of frontolimbic passage, dome at median hemispheric facies. !45.
Frontal Lobe
71
Limbic Frontopolar Area FE L and Limbic

Prefrontal Area FH L
Similar structural changes pertain to the more

anteriorly situated variants of areas FE and FH,
i.e. the limbic frontopolar area FE L (area EK 19;
2008]) and the limbic prefrontal area FH L (area
EK 27; plates P38 and P39 in the Atlas [Economo
and Koskinas, 2008]) on the frontolimbic gyrus,
which are all characterized by the presence of a
band-like cellular disposition in layer V. As I explained in the introductory chapter, this particular overgrowth of layer V is rather typical of the
vicinity of the rhinencephalon sensu lato and of
certain parts of the rhinencephalon proper.
Pretriangular Orbital Area FF
Regarding area FF, I already spoke of its transformation from a granular to an almost agranular
type on the orbital hemispheric surface of the
frontal lobe. Here, I wish to merely mention in
brief the cytoarchitectonic structure of the pars
pretriangularis of F3, which we denote as its pretriangular orbital area FF variant (area EK 22)
(fig. 1a, d; cf. plate P34 in the Atlas [Economo and
Koskinas, 2008]), despite the fact that its change
is so great that it can be hardly ascribed to the
formation FF.
In particular, this small territory comprises
12 small gyri in front of the caput, which effect
the transition from the lateral to the basal hemispheric surface of the frontal lobe. An extreme
thinning of the cortex is observed, primarily affecting the upper cortical layers, and above all
layer III.
Layer III equally becomes especially impoverished in cells, even revealing acellular patches;
its cells are very small.
72
Layer IV is also thin and oligocellular.

Layers V and VI are the only layers that appear somewhat thicker and with larger cells, thus
justifying the incorporation of this territory more
or less into ground area FF.
The superficial cortical layers mostly recall
the structure of area FG, which they resemble
markedly; however, we do not wish to incorporate this field into ground area FG, because it does
not stand in any direct continuity with it.
Opercular Modifications FCop and FDop of

Intermediate and Granular Frontal Areas
I have only to add that at the points where areas

FC and FD coat the ventral surface and partially
the internal wall of the operculum, they usually
assume the earlier mentioned typical opercular
modifications of the cerebral cortex as well;
namely, a pronounced thickness of the granular
layers and an untidy cluttering of the sparser cells
of layer III, with an overall thinning of the cortex.
Thus, we could also speak of two modifications in this area, partially covering the antidiagonal and antitriangular gyri: the opercular intermediate frontal area FCop (area EK 10) and the
opercular granular frontal area FDop (area EK 14).
Finally, a transition form between these two
modifications is also found, which we call opercular intermediate granular frontal area FCDop
(fig. 1a, d and table 2; also appearing as FC(D)op
[Economo and Koskinas, 1925, p. 329] and FDCop
[Economo and Koskinas, 1925, p. 356]; cf. plates
P16, P18 and P20 in the Atlas [Economo and
Koskinas, 2008]).
Parietal Lobe
Macroscopic and Cytoarchitectonic

Boundaries
We now come to the discussion of the very extensive parietal lobe, which is the most important
lobe of the cerebral hemispheres after the frontal
lobe; as figure 1 shows, its cytoarchitectonic
boundaries outrun those usually admitted by
gross (macroscopic) anatomy.
It begins at the depths of the central sulcus of
Rolando and from there stretches on the median
(midsagittal) hemispheric facies backwards to
the parieto-occipital sulcus (S.po.); in the vicinity
of the limbic gyrus, it is bordered at first by the
horizontal ramus of the callosomarginal sulcus,
and further caudally by the subparietal sulcus,
such that it comprises the posterior part of the
paracentral lobule and the entire precuneus.
On the superolateral hemispheric convexity,
the parietal lobe extends from the valley floor of
the central sulcus of Rolando to the notch of the
parieto-occipital sulcus at the superomedial border of the longitudinal cerebral fissure, and further ventrally as far as the interoccipital sulcus,
also including the entire temporo-occipital intermediate region up to a somewhat imprecise posterior border, which approximately corresponds
to an imaginary extrapolation line projected
from the interoccipital sulcus perpendicularly
downwards and passing over to the inferomedial

hemispheric margin ventrally.
The lower boundary on the (lateral) hemispheric convexity is primarily the lateral (Sylvian) fissure and the continuation from its posterior extreme, at nearly a right angle, perpendicularly downwards, to form the anterior boundary
line of the temporo-occipital intermediate region. Thus, the preoccipital notch (or incisure)
and the anterior occipital sulcus, which in macroscopic anatomy are usually regarded as the
posterior boundary between this intermediate
region and the occipital lobe, fall within the temporo-occipital region, which, from the cytoarchitectonic viewpoint, now belongs to the inferior
parietal lobe, constituting, in our nomenclature,
its basal region (table 1). Accordingly, the most
caudal segments of the middle (T2) and inferior
(T3) temporal gyri largely fall within this basal
region of the parietal lobe as well.
On the ventral cerebral facies, this part of the
parietal lobe then stretches across the inferior
hemispheric margin into the common trunk of
convergence of the calcarine and parieto-occipital
sulci, becoming gradually narrower in the rostral
direction (fig. 1b, d). Therefore, with these new
cytoarchitectonic confines, the parietal lobe forms
an almost closed ring around the cerebral mantle;
its opening on the median hemispheric facies is
occupied by the retrosplenial region of the limbic

gyrus, as well as [Economo, 1929d] by the anterior
pointed extreme of the occipital lobe.
Cellular Structure
In its extensive spread, the parietal lobe presents

with very diverse structural areas; those in the
postcentral gyrus and the superior parietal lobule
reveal a certain analogy with the structure of the
frontal lobe (cortical structural type 2). However,
its cellular structure, especially in the lower and
basal parts, is fundamentally different from that
of the frontal cortex and typically presents as
cortical structural type 3 (cf. Introduction and
fig. 9); we accordingly call it the parietal type.
The cortex of this structural type is rather
thick, but never quite attains such high measurements as the thickest parts of the frontal cortex;
both granular layers (II and IV) are usually composed of true granule cells; they are extremely
distinct, thick, and highly cellular, such that the
horizontal lamination of the cortex is much more
conspicuous here than it is generally in the frontal lobe.
A further important characteristic is a fine radial striation that traverses all cortical layers, but
it is especially distinct in layer III. The parietal
structural type contains cells higher in number
but smaller in size than the frontal lobe, and layer
III has usually only medium-sized pyramidal
cells; cells the size of large pyramidal cells, which
we often saw forming a clearly delimited sublayer
IIIc in the frontal cortex, only occur here rarely.
I must also emphasize the lesser development
of layer VI, and particularly layer V, which progressively becomes thinner toward caudal hemispheric regions. Therefore, while layers V and VI
are still fairly well constituted in the anterior and
upper regions of the parietal lobe, i.e. in the postcentral gyrus and the superior parietal lobule, we
see them distinctively diminishing in the inferior
parietal lobule, and even more at the boundary
74
with the occipital lobe, where nerve cells become

markedly smaller in size; such a progressive thinning continues within the occipital lobe, as far as
the posterior hemispheric pole. I later discuss the
fact that the superior (T1) and fusiform (T4) temporal gyri display a certain resemblance to the
parietal cortical structural type, in contrast to T2
and T3, which are distinctively temporal in their
cytoarchitecture.
As already mentioned, the parietal cortical
structural type still reveals certain elements of
the frontal structural type on the postcentral gyrus and the superior parietal lobule, and that is
why I discuss those first.
Anterior (Postcentral) and Superior Parietal

Regions: Areas PA PE
The boundary between the frontal and the parietal lobe runs along the floor of the central sulcus
of Rolando, usually projecting up the wall of the
precentral gyrus. Such a boundary is generally
well defined by a sharp (totally colossal) reduction in cortical thickness, and by a sudden affluence of layers II and IV in granule cells.
On the postcentral gyrus alone, we may distinguish four ground areas, which stretch like
linear extensions from the operculum to the superomedial hemispheric border at the longitudinal cerebral fissure and then onto the paracentral
lobule. The first two of these areas, the giant pyramidal postcentral area PA and the oral postcentral area PB , lie superimposed in the anterior wall
of the postcentral gyrus; the intermediate postcentral area PC occupies the dome of the postcentral gyrus from the operculum to within the
paracentral lobule, and the caudal postcentral
area PD occupies the posterior wall of the postcentral gyrus and the facing anterior wall of the
adjacent gyrus, i.e. both walls of the postcentral
sulcus (fig. 1a, c).
The entire superior parietal lobule is vested by
the superior parietal area PE (fig. 1ac; table 1).
Fig. 28. Giant pyramidal postcentral area PA1 (area EK 55). Anterior wall of postcentral gyrus at the floor of the central
sulcus of Rolando. !40.
Giant Pyramidal Postcentral Area PA
At the floor of the central sulcus of Rolando, the

cerebral cortex which at the posterior wall of
the precentral gyrus was over 3.0 mm thick as the
agranular cortex of area FA becomes briskly
reduced to 2.0 mm or even less, rolling over into
the lower segment of the anterior wall of the postcentral gyrus (fig. 28). Concomitantly, layers II
and IV, which were barely discernible in the lower part of the posterior wall of the precentral gyrus, become altogether distinct, and in so doing,
the agranular, heterotypic cortex of area FA
changes into the six-layered (hexalaminar) agranular homotypic isocortex of area PA . The two
granular layers are compactly filled with small
Parietal Lobe
granule cells, appearing in the cytoarchitectonic

picture as dark blue horizontal strips. Layer V,
which has also thinned considerably, becomes at
the same time somewhat paler; one can observe
in it isolated giant cells of Betz spreading over
from area FA, although these appear more flattened and smaller than in the precentral gyrus
(cf. plate P66 in the Atlas [Economo and Koskinas, 2008]).
Characterized specifically by its reduced
thickness, the presence of giant cells of Betz, and
two distinct granular layers, this giant pyramidal
area PA occupies the valley and the lower onethird of the posterior wall of the postcentral gyrus in the central sulcus of Rolando in its entire
expansion. Specifically, area PA stretches from
75
the opercular end of the central sulcus, on the

dome surface of which it ascends only slightly
from the depths of the valley, to the crease of the
paracentral lobule, on the surface of which it
spreads widely, over the dorsal end of the central
sulcus (fig. 1a); it then expands over this entire
region as far as the horizontal and the vertical
rami of the callosomarginal sulcus, even widening towards the outside (fig. 1b), to follow the vertical ramus to the level where the latter cuts into
the superomedial hemispheric edge, and again
on to the superolateral cerebral convexity. In that
way, area PA encircles the upper part of the postcentral gyrus with its somewhat hook-shaped
modifications giant pyramidal postcentral area
PA1 (area EK 55) and giant pyramidal postparacentral area PA2 (area EK 56) (fig. 1ac).
At the wall of the postcentral gyrus, the cortex
of area PA has an average thickness of about 2.0
mm, which, however, reaches 2.8 mm at the level
of the paracentral lobule [Economo, 1928e]. Figure 28 shows this parietal area at its actual transition point from the floor of the central sulcus of
Rolando to its posterior wall.
Layer I has an average thickness, like at the
walls of gyri elsewhere. The layer is also poor in
cells, as usual (table 7).
Layer is very rich in cells. The more superficially located cells are real granule cells, whereas the more deeply located ones are rather triangular, such that the border of this layer with layer
III becomes indistinct.
Layer III is relatively thick for a wall formation. Cells are orderly arranged into three secondary sublayers: IIIa with small pyramidal
forms; IIIb much lighter and poorer in cells (medium-sized pyramidal), and IIIc containing
about 25% larger pyramidal cells. One may encounter some isolated examples of extremely
large pyramidal cells. In many places, especially
at the valley floor, layer III appears to be widely
and radially striated.
Layer IV forms a thick, compact and conspicuous layer.
76
Layer V stands in sharp contrast with the

compactness of layer IV by its paucity of cells,
which makes it appear lighter. But, as I already
mentioned, one always finds some examples of
giant pyramidal cells of Betz in this layer, recognizable from their relatively large size, Nissl (tigroid) bodies, lipid and pigment inclusions, varicose (nodose) dendrites, large vesicular nucleus
with a distinct nucleolus, and other cytologic features. And yet, these giant cells are not at all as
well developed as those of the precentral gyrus,
being frequently flattened and elongated, especially at the valley floor.
Layer VI is subdivided into an upper sublayer
VIa and a lower sublayer VIb. Only VIb, which is
poorer in cells than VIa, deserves the name of a
spindle (fusiform) cell layer. The more compact
and band-like sublayer VIa largely consists of
triangular and pyramidal cells. Demarcation
against the white matter is sharp.
Postcentral PA1 and Postparacentral PA2 Giant

Pyramidal Area Modifications
At its passage to the surface of the paracentral
lobule in the dorsal end of the central sulcus of
Rolando, this area changes its structure insofar as
any area usually changes between the wall of a
gyrus and its dome. The cortex becomes thicker,
up to 2.8 mm, and just as the cells themselves also
usually increase in size on the domes of gyri, we
see here that they are larger at the surface of the
paracentral lobule than at the wall of the postcentral gyrus. It even looks as if the giant cells of Betz
also conform to this general condition, but even
so, they always remain smaller, lesser developed
and fewer in number than in the anterior part of
the paracentral lobule occupied by the giant pyramidal precentral area FA (area EK 2).
On the basis of those cytoarchitectonic differences between the sulcus and the dome, we may
term the part of area PA at the surface of the paracentral lobule giant pyramidal postparacentral
Fig. 29. Oral postcentral area granulosa PB1 (area EK 58). Middle part of anterior wall of postcentral gyrus in the central sulcus of Rolando. !40.
area PA2 (area EK 56), and that in the sulcus giant

pyramidal postcentral area PA1 (area EK 55) (cf.
plates P59a, P62, P66 and P67 in the Atlas [Economo and Koskinas, 2008]).
As mentioned earlier, the giant pyramidal
area PA2 follows the vertical ramus of the callosomarginal sulcus from the median surface over
the superomedial hemispheric border at the longitudinal cerebral fissure, and continues on to
the lateral hemispheric convexity, reaching the
dorsocaudal part of the postcentral gyrus and the
superior parietal lobule, such that the remaining
areas of the postcentral gyrus (oral postcentral
Parietal Lobe
area PB and intermediate postcentral area PC) are

encircled by area PA2 at the edge of the interhemispheric (longitudinal) fissure like by a collar (cf.
fig. 1ac). To the degree in which such a loop occurs, one also finds some disseminated giant cells
in layer V of the other fields of the dorsalmost
parts of the postcentral gyrus (areas PB and PC),
which do not otherwise contain such cells; thus,
the giant cell cortex on the edge of the interhemispheric (longitudinal cerebral) fissure reaches
from the foot of the superior frontal gyrus (F1) as
far back as the anterior segment of the anterior
arcuate gyrus of the superior parietal lobule.
77
Table 7. Summative table of quantitative data in ten ground and variant areas of the parietal lobe. Overall layer thickness based on the present work. Separate dome and wall data and some additional values supplemented from tables
I, III, V and VI in Economo and Koskinas [1925, pp. 794801].
Area
symbol
Area name
Cortical
layer

mm

mm

mm
Cell content
cells/mm3
Cell size, m*

H(minmax)/
W(minmax)
PA
1postcentral
I
II
IIIa
IIIb
IIIc
IV
V
0.272
0.252
0.231
0.201
0.23
0.20
0.782
0.641
0.65
0.312
0.352
0.261
0.221
0.25
0.15
9,000
100,000
65,000
40,000
25,000
150,000
35,000
VIa
VIb
0.452
0.302
0.271
0.181
6/4
5/5
10/8
20/15
3550/20
5/5
15/10
50/25
1520/10
15/10
I
II
III
IV
V
VIa
VIb
0.26
0.24
0.44
0.34
0.28
0.26
0.34
0.26
0.29
0.47
0.29
0.21
0.25
0.14
I
II
IIIa
IIIb
IIIc
IV
V
VIa
VIb
0.30
0.26
0.24
0.20
0.27
0.23
0.72
0.56
0.64
0.32
0.20
0.18
0.12
0.28
0.24
0.26
0.18
0.30
0.22
PB1
PB2
PC
PD
78
giant
pyramidal area
PA1 and
2
postparacentral
giant pyramidal
area PA2
oral postcentral
area granulosa
oral postcentral
area simplex
intermediate
postcentral area
caudal postcentral
area
I
II
IIIa
IIIb
IIIc
VIa
VIb
0.60
0.50
wall
structure
only
0.50
0.37
0.22
0.23
0.86
0.36
0.38
0.60
0.25
0.30
0.45
0.30
0.20
0.23
0.23
IV
V
I
II
IIIa
IIIb
IIIc
IV
V
VIa
VIb
dome
structure
only
8,000
210,000
105,000
230,000
60,000
60,000
17,500
6/7
5/5
46/46
5/5
1520/1015
1520/915
15/10
8,000
100,000
80,000
60,000
60,000
120,000
45,000
60,000
17,500
510/57
510/57
10/7
15/10
1030/1020
610/610
1025/1020
15/10
15/10
8,000
130,000
35,000
28,000
25,000
0.35
0.40
130,000
30,000
1.10
45,000
20,000
6/7
67/56
1015/10
1520/15
30/15
3570/2535
68/68
1015/10
4060/2025
1820/10
1516/78
9,500
130,000
56,000
44,000
16,000
135,000
30,000
40,000
20,000
79/6
510/57
1015/710
1520/1015
3060/1525
5/5
1520/1015
15/10
1012/10
0.90
0.21
0.20
0.21
0.20
0.64
0.65
0.21
0.34
0.30
0.10
0.20
0.30
50,000
15,000
0.40
Table 7 (continued)
Area
symbol
Area name
Cortical
layer
PE
superior parietal
area
Ia
Ib
II
IIIa
IIIb
IIIc
IVa
IVb
Va
Vb
VIa1
VIa2
VIb
PF
PG
PH
supramarginal area
angular area
basal (temporooccipital) parietal

area
Ia
Ib
II
IIIa
IIIb
IIIc
IVa
IVb
Va
Vb
VIa
VIb
I
II
IIIa
IIIb
IIIc
IVa
IVb
IVc
V
VIa
VIb
I
II
IIIa
IIIb
IIIc
IV
V
VIa
VIb

mm

mm

mm
0.18
0.24
0.18
0.19
0.28
0.20
0.77
0.68
0.70
0.30
0.30
0.40
0.40
0.30
0.40
Cell content
cells/mm3
Cell size, m*

H(minmax)/
W(minmax)
9,000
2,500
110,000
50,000
25,000
20,000
65,000
80,000
35,000
17,000
17,000
25,000
10,000
5/8
5/8
510/510
1520/1015
2030/1520
3050/2025
56/56
9/810
2535/1520
12/12
25/15
15/12
1215/10
1015/10
67/67
46/46
1518/1015
1525/1015
1540/1020
56/56
710/710
1015/10
2025/1520
1530/815
15/8
0.62
0.30
0.40
0.18
0.26
0.30
0.25
0.30
0.30
0.32
13,000
4,000
120,000
1.00
0.76
1.00
30,000
0.40
0.29
0.40
0.45
0.90
90,000
115,000
35,000
40,000
30,000
15,000
0.47
0.34
0.70
0.45
0.46
0.24
0.22
0.20
0.20
0.18
0.22
0.20
8,000
95,000
0.72
0.80
0.80
25,000
0.35
0.30
0.35
0.58
0.60
0.60
0.34
0.36
0.30
0.55
0.22
0.26
0.22
0.22
0.20
0.22
10,000
110,000
0.70
0.60
0.90
40,000
0.24
0.60
0.50
0.20
0.20
0.50
0.40
0.10
0.22
0.65
100,000
40,000
40,000
12,000
1.00
1.20
0.65
100,000
140,000
60,000
18,000
25,000
18,000
56/6
512/510
1215/10
1220/1015
1235/1020
58/58
1012/1012
1520/10
1525/1015
1525/15
1015/710
7/6
412/410
18/815
1825/1018
1850/830
68/65
1220/15
15/12
15/10
* See footnote in table 6.
Parietal Lobe
79
Oral Postcentral Area PB
While area PA1 occupies the valley floor and the

lower one-third of the anterior wall of the postcentral gyrus, the upper two-thirds of the anterior wall are occupied by the altogether heterotypic cortex of area PB , which also extends as a
band parallel to area PA at the wall of the postcentral gyrus, from the operculum to the paracentral
lobule (fig. 1ac). Area PB can be distinguished
from area PA by the fact that most of the cells in
this region (fig. 29) appear much smaller in the
cortical sections, almost granular; the granular
layers themselves become thicker, such that the
cortical cross-section assumes a particular dustlike appearance, and the whole, from the lower
border of layer I all the way to layer V, appears
as a single, thickened granular layer (marked
II+III+IV in fig. 29). Thus, this area constitutes
an example of heterotypic cortex that we call granulous (or koniocortex). That heterotypy ensues
from a granulization of the cells in layer III, as
well as by a substantial increase in their number.
The thickness of the cortex in this region is reduced to a minimum (1.8 mm), and layer V becomes less marked and poorer in cells, as it is
filled with a thick fiber network, as one can observe in sections prepared with the Weigert myelin stain.
Oral Postcentral Area Modifications Granulosa

PB1 and Simplex PB2
The granulization is not uniform in intensity
through all the segments of area PB ; territories
showing the maximum granulization form strips
and islets throughout area PB . One may term its
more granulous part oral postcentral area granulosa PB1 (area EK 58) (shown in plates P59b, P60a,
P61 and P62 in the Atlas [Economo and Koskinas, 2008]), and the part with small cells, which
still possess a distinct pyramidal form, oral postcentral area simplex PB2 (area EK 57) (shown in
80
plates P60b and P61 in the Atlas [Economo and

Koskinas, 2008]).
Layer I is poor in cells, as usual (table 7).
Layer II is very compact.
Layer III contains an unusually large number
of cells for this layer; at a low power, layer III appears hardly distinguishable from layers II or IV
(denoted as II+III+IV in fig. 29). One nonetheless
observes that even in the granulous parts of layer
III, there are a few particularly long and slender
pyramidal cells that appear clearly as isolated examples among the large mass of smaller cells.
Layer IV beneath contains cells in a dense
manner. Its upper and lower borders are very indistinct; as already mentioned, one gets the impression that layers II, III and IV form a single,
thick granulous layer.
Layer V, on the other hand, forms a strikingly
light thin strip, very poor, with small and medium-sized pyramidal cells. Not infrequently,
especially at the depths of the central sulcus
throughout its entire expansion, as well as near
the superomedial hemispheric edge of the longitudinal cerebral fissure, one can see individual,
flat, and relatively small giant cells of Betz, which
have wandered thus far out of their domain (not
visible in fig. 29).
Layer VI forms an even more prominent band
in this region than in area PA, its cells being also
smaller than usual, usually not of a spindle (fusiform) shape, but rather triangular. In the upper
sublayer VIa they are almost quadruple in number compared to the lower sublayer VIb (table 7).
The density and band-like form of sublayer VIa
cause layer VI to stand out clearly from V, which
stains lightly and is oligocellular. The border with
the white matter is sharp.
With the increasing shallowness of the dorsal
segment of the central sulcus of Rolando toward
the superomedial hemispheric edge, the koniocortex of area PB also passes gradually to the surface of the paracentral lobule, where it occupies
an oval field (fig. 1ac), found inwards of the territory surrounded by areas PA1 and PA2 on the
posterior paracentral lobule. This koniocortex

preserves its own typical structure even at this
surface, being nonetheless subjected to the usual
changes at the passage from wall to dome; in other words, one observes a certain cortical thickening, an increase in cell size, and a slight reduction
in cell number. At the inferior extreme of the central sulcus, area PB does not pass to the surface of
the operculum, but ends rather bluntly in the
widened and shallow end of the sulcus.
Function
As we already know, the koniocortex represents a
sensory cortex, i.e. a cortical zone, into which sensory impulses relayed to the diencephalon, following their confluence from the sense organs
into the spinal cord, are admitted for further processing and assimilation. In area PB , in particular, one finds the tactile domain for the various
cutaneous sensations (tactile, thermesthesia,
etc.), and probably also the center for other tactile
sensations, including muscle sensibility from the
entire body. The opercular segments serve as cortical stations for sensations from the mouth, perhaps also for certain taste modalities. The entire
surface of the koniocortex of the tactile domain
in both hemispheres most likely occupies 3540
cm2. The fact that the pathways from the sensory
diencephalic nuclei radiate to the postcentral gyrus has been verified by anatomical fiber tract
studies as well. The tactile domain is more exhaustively treated in our larger work [Economo
and Koskinas, 1925, pp. 538544].
Intermediate Postcentral Area PC
This homotypic isocortical structure comprises

in its range the entire dome and usually both
brinks of the postcentral gyrus over its entire extent, from the operculum to the paracentral lobule. Thus, the intermediate postcentral area PC
Parietal Lobe
(area EK 59) forms a wide, although superficial,

band that is immediately adjacent throughout its
length to the afore-mentioned area PB (fig. 1a, c).
In its cellular structure it has a particular similarity to that of the frontal lobe (fig. 30; cf. also
plates P61P63 in the Atlas [Economo and Koskinas, 2008]), especially of areas FB and FC, but it
mainly differs from them in its distinctive granular layers and cell opulence, whereas these regions have in common a marked growth of layers
III and V. The exceptionally large and robust pyramidal cells of sublayer IIIc in area PC often exceed the largest pyramidal cells of the frontal lobe
in size and structural fineness, and are also perpendicularly arranged just as in the frontal lobe,
a situation that gives the cortex a radial, regularly striated appearance. The thickness of the cortex here varies between 3.03.3 mm.
Layer I is parvicellular and indigent in cells, as
usual (table 7).
Layer II contains granule and very small pyramidal cells.
Layer III almost reaches 1.0 mm in thickness,
like the posterior areas of the frontal cortex. It is
distinctly subdivided into a rather cellular, upper
sublayer IIIa containing small pyramidal cells; a
somewhat lighter intermediate sublayer IIIb with
medium-sized pyramidal cells, and a deep sublayer IIIc with large pyramidal cells. In sublayer
IIIc one further finds some very well-formed
large pyramidal cells nearly as large as giant cells;
these are particularly found in the anterior brink
of the gyrus, i.e. at the boundary with area PB ,
lending the area a distinctive appearance, and
characterizing it as a particular field among all
the other areas of the six-layered isocortex.
Layer IV is fairly thick and contains plentiful
granule cells.
Layer V is very well developed. Its main mass
is composed of medium-sized cells, but there are
also regular examples of some very large pyramidal cells. Compared to the compact layer IV, layer
V seems somewhat dissolved. In the region of the
superomedial hemispheric edge of the longitudi-
81
Fig. 30. Intermediate postcentral area PC (area EK 59). Dome and caudal brink of postcentral gyrus. !45.
82
nal cerebral fissure toward the median hemispheric facies, area PC also displays a few giant
cells of Betz of a smaller caliber; the same is true
for the part of area PC that passes over to the paracentral lobule.
Layer VI again contains robust spindle (fusiform) cells, in contrast to the two areas PA and PB
discussed earlier. These medium-sized cells are
about twice as numerous in sublayer VIa as in the
lower sublayer VIb. The demarcation from the
white matter is more or less precise.
Giant Pyramidal Intermediate Postcentral Area

Modification PC, a Parasensory Zone
At the operculum, area PC reaches on the postcentral gyrus almost as far as area FB does on the
precentral gyrus; the boundary between these
two areas varies widely among individuals with
regard to its exact position from the postcentral
to the precentral gyrus (fig. 1a, c). Near the superomedial hemispheric edge of the longitudinal
(cerebral) fissure, as I mentioned earlier for area
PA, giant cells of Betz appear in layer V of area PC.
It becomes rich in giant cells not only in sublayer
IIIc, but also in layer V, and for this reason, we
call this part of area PC giant pyramidal intermediate postcentral area PC (cf. plate P64 in the Atlas [Economo and Koskinas, 2008]). It extends
dorsally, over the superomedial hemispheric edge
of the longitudinal cerebral fissure, from the
paracentral lobule to the median hemispheric
surface, and takes up a position behind the upper
extreme of area PB in the concave space formed
by the hook-shaped loops of modifications PA1
and PA2 (fig. 1b, c).
Function
The physiologic role of area PC is only understood
to the extent that electric stimulation studies are
concerned; such a stimulation provokes two
Parietal Lobe
kinds of movements: first, global movements

passing to lower centers by means of direct descending pathways and representing complex
postural motions, and, second, special single
movements that first pass over the precentral gyrus and are only transmitted from there to lower
centers in the medulla.
At the boundary with koniocortex PB the cells
of sublayer IIIc in area PC sometimes have a particularly large size. Such marginal zones with
magnocellular characteristics are found rather
regularly at the boundaries of any koniocortex,
i.e. any sensory zone; we hence call them parasensory zones (cf. Introduction). Area PC distinctly reveals, apart from its marked granular
layers, the cortical structural type 2, and thus essentially recalls the frontal type of cortex.
Caudal Postcentral Area PD
At the posterior wall of the postcentral gyrus, the

gray cortex, which was thick at the dome of the
gyrus, thins again markedly out of proportion;
such an attenuation exceeds that usually seen
elsewhere between dome and wall, such that, by
taking also into consideration certain other
structural changes in the posterior wall of the
postcentral gyrus, we may view it as a separate
striated area, which we term caudal postcentral
area PD (area EK 60) (fig. 31; cf. also plate P65a in
the Atlas [Economo and Koskinas, 2008]). The
thickness of the cortex reaches 1.82.0 mm. In
contrast to the anterior wall of the postcentral
gyrus, its posterior wall is characterized by its
large cells, and especially the band-like, thick and
multilaminar disposition of pyramidal cells in
sublayer IIIc. This cellular superimposition also
differentiates area PD from all the other regions
of the superior parietal lobe, and especially the
anterior wall of the postcentral gyrus, that are in
other respects similarly structured at the walls.
Layer I is rather thin and contains perhaps
slightly more nerve cells than usual (table 7).
83
Fig. 31. Caudal postcentral area PD (area EK 60). Posterior wall of postcentral gyrus. !40.
Layer II is considerably thinner than in the

anterior wall, although thicker than usual. It is
very densely populated.
Layer III is very thick for a wall formation and
very cellular. It is separated into distinct sublayers IIIa, IIIb and IIIc. Cells are superimposed in
several dense rows, rendering a distinct appearance to the entire area.
Layer IV contains granule cells, densely
packed, and a few triangular cells.
Layer V, directly subjacent, is sharply delimited from layer IV, and stands out by its light
staining. There are sporadic, large pyramidal
cells.
84
Layer VI is relatively thin. Only one-half of

the cells are spindle-shaped (fusiform), the rest
being pyramidal. This unusual feature differentiates area PD from other areas. The upper sublayer VIa has twice as many cells than the lower
sublayer VIb. The demarcation against the white
matter is less sharp than at the anterior wall of the
postcentral gyrus.
Transition Area Modifications PDE and PFD

Area PD does not only occupy the posterior wall
of the postcentral gyrus in its entirety, i.e. all of
the postcentral sulcus, from the superomedial
hemispheric edge to the operculum, but also

passes over the floor of the postcentral sulcus to
its opposite wall that belongs to the superior and
inferior parietal lobules, and even stretches,
shaped like a tongue, backwards along the walls
of the interparietal sulcus.
In that caudal prolongation, the area does not
retain its pure structural type, but shows an admixture of the type of its adjacent superior parietal area PE , which we indicate by marking that
prolongation as transition area PDE (appearing
as PE(D) in fig. 1a; cf. also plate P65b in the Atlas
[Economo and Koskinas, 2008]). Dorsally, area
PD hardly reaches the superomedial hemispheric
edge, but persists only as far as the postcentral
sulcus. Ventrally, however, this area moves from
the end of the postcentral sulcus onto the surface
of the operculum, and covers the posterior and
inferior parts of that region with the somewhat
modified cortical structure of the superior-postcentral parietal transition area PDE (also written
as PE(D) or area EK 61); when, in turn, this shows
some characteristics of the supramarginal area
PF (area EK 65), it then becomes the supramarginal-postcentral transition area PFD (fig. 1a, d),
where it stretches directly up to the agranular
frontal area FB (area EK 4).
Superior Parietal Area PE
The entire superior parietal lobule is occupied by

a single area (fig. 1ac), beginning at the subparietal sulcus of the precuneus on the median hemispheric facies and reaching to the interparietal
sulcus on the superolateral convexity, and from
the postcentral sulcus or the vertical ramus of the
callosomarginal sulcus anteriorly to the parietooccipital sulcus on the median hemispheric surface and the upturned sulcus of Brissaud [1893]
on the superolateral convexity. Thus, area PE
covers all three arcuate gyri that compose the superior parietal lobule.
Parietal Lobe
In this expanded area, the cortex is thinner at

the most rostral and caudal parts, and thicker in
the middle (intermediate arcuate gyrus). Thickness varies between 2.6 and 3.0 mm. Moreover,
the anterior parts of area PE have in general larger cells, and the caudal parts smaller cells; these
modifications are respectively termed magnocellular PEm (area EK 62) and parvicellular PEp (area
EK 63) superior parietal areas (fig. 1ac; cf. also
plates P68P70 in the Atlas [Economo and Koskinas, 2008]). Area PE is further characterized by
the thickness of its granular layers and a markedly light stria in layer V visible to the naked eye.
There is also a distinct radial striation (fig. 32).
Layer I is poor in cells (table 7).
Layer II is distinct because of the dense arrangement of its authentic granule cells.
Layer III is subdivided into sublayers IIIa, IIIb
and IIIc. Sublayer IIIc contains cells that are not
collected in such a dense layer as in area PD. A
certain arrangement of the cells in radial stretches can be readily seen in figure 32 (the deeper part
of layer III).
Layer IV is very thick. It is subdivided into an
upper sublayer IVa with less compacted, roundish granule cells, and a deeper sublayer IVb with
more densely packed, usually triangular cells.
Sublayer IVb can be further subdivided into an
upper, denser subzone IVb1, and a looser, deeper
subzone IVb2. Due to the loose arrangement, and
the lighter staining of the upper sublayer IVa,
which separates the dense sublayer IIIc from the
dense subzone IVb1, the sublayer IIIc seems to
hang freely above the subzone IVb1, an appearance rather characteristic of area PE , becoming
particularly conspicuous in paler stained specimens. A certain radial cellular disposition is also
discernible in layer IV.
Layer V is subdivided into a rich upper sublayer Va, containing relatively large cells, and a
deeper and lighter sublayer Vb, with smaller and
fewer cells. Sublayer Vb typically appears as a
fine stria in the cytoarchitectonic picture under
very low magnifications. Sublayer Va also con-
85
Fig. 32. Superior parietal area PE (area EK 63). Dome of middle part of superior parietal lobule. !45.
86
tains some scattered larger cells; sublayer Vb only

possesses half as many cells. If we discount the
areas of the postcentral gyrus, then the superior
parietal lobule contains in its layer V the largest
cells, relatively speaking, of all the rest of the cortex behind the central sulcus. The best differentiation of area PE from any of the other areas of
the parietal lobule can be based on the relative
size of these cells, as well as the lighter staining of
layer V. Such a differentiation from other areas of
the inferior parietal lobule is clear, as that lobule
has only small-sized cells in layer V. Layers V and
VI also show a coarse, radial striation.
Layer VI is usually very thick at the domes of
the gyri. It is distinct and reveals a thick, dense,
strip-like upper sublayer VIa with large spindle
(fusiform) cells; sublayer VIb contains only relatively small spindle cells. One may separate two
tertiary subzones, a less dense upper subzone
VIa1 and a lower subzone VIa2 that contains larger triangular cells. The denser subzone VIa2 thus
appears as a darker band between the lighter subzone VIa1 and sublayer VIb. The typical spindle
cell form in layer VI of area PE even at the gyral
walls is also a differentiating mark from area PD
described earlier. The border against the white
matter is rather sharp.
Although the structure of the cortex in the superior parietal lobule is unquestionably similar to
the inferior parietal region (detailed next), and
although the distinctiveness of both granular layers characterizes this region as belonging to cortex behind the central sulcus, it nevertheless has
many resemblances, regarding the cellular structure and size in layers III and V, to the formations
of the intermediate region of the frontal lobe, i.e.
cortical structural type 2 (cf. Introduction). We
would therefore prefer to assign this region to the
frontal cortical type 2 (fig. 9), as it resembles it
closer than it does the parvicellular structures of
the inferior parietal lobule, the temporal or the
occipital lobe.
Function
The physiologic function of area PE is unknown;
nonetheless, electric stimulation of this region
provokes contralateral locomotor complexes in
both extremities. These impulses travel to the periphery through intrinsic paths in the corona radiata. A center for ocular movements is possibly
located in area PE.
Properties of Cortical Structural Type 3
Giant Pyramidal Posterior Superior Parietal

Area Modification PE
In its expansion over the superior parietal lobule,
area PE shows the regional structural changes
with respect to cortical thickness and cell size
that I mentioned at the beginning of this section.
The caudalmost part of this area on the posterior
arcuate gyrus always shows characteristic slender
giant cells at the vicinity of the occipital cortex;
after their occurrence in layer V, we term this
caudal part of the upper parietal area giant pyramidal posterior superior parietal area PE (area
EK 64) (fig. 1ac; cf. also plate P71 in the Atlas
Parietal Lobe
The inferior parietal lobule, as I mentioned earlier in this chapter, does not only include the supramarginal and angular gyri with both their
posterior transition gyri to the occipital lobe (parieto-occipital gyri), but also, in its basal region,
the entire temporo-occipital intermediate zone
all the way to the base of the cerebral hemispheres;
it is all vested in a typically structured, distinctly
stratified thick cortex, which is characterized by
extremely striking granular layers II and IV, and
also by a robust growth of layer III; on the other
hand, layers V and VI lose much of their importance in the cytoarchitectonic picture.
87
Almost all the cells of layer III have a uniformly medium calibre; therefore, the subdivision into
sublayers that is so prominent in the frontal and
the superior parietal lobes is not seen in this region. The cells of layers V and VI are remarkably
small, with layer V cells diminishing in size more
rapidly toward the occipital pole than those of
layer VI, such that in the caudal region of the
temporo-occipital intermediate zone of the parietal lobule, the relative size of cells of layer V is
even smaller than that of VI; this is the opposite
phenomenon of what occurs everywhere else in
the cerebral hemispheres, and becomes even
more accentuated at the occipital lobe.
Another typical characteristic of this entire
cortex is the fine radial (i.e. vertical) striation,
resulting from the disposition of cells of all layers particularly those of the upper layers and
with the exception of layer II in thin perpendicular columns. The intensity of such a vertical
striation varies depending on the location.
The cortex of this entire region belongs to the
structural type 3 (parietal type), already dealt
with in the Introduction; I take this opportunity
to mention that the superior (T1) and fusiform
(T4) temporal gyri also show suggestions of this
parietal type of cortical structure.
Inferior and Basal (Temporo-Occipital)

Parietal Regions: Areas PF PH
The division of this expanded region into cytoarchitectonic areas is far more difficult than in the
frontal lobe, because the cortex here shows a
greater number of small variations, more frequent
intermediate and transition types, and territories
that are less clearly demarcated and circumscribed
than in the frontal lobe. Our delimitation of areas
is thus somewhat arbitrary, and largely rests on
the macroscopic anatomic boundaries.
In the inferior parietal region, we recognize
two ground areas in the respective gyri, namely,
the supramarginal area PF on the supramarginal
88
gyrus, and the angular area PG on the angular

gyrus. In the basal parietal region, we recognize
the basal (temporo-occipital) parietal area PH and
its modifications, occupying the temporo-occipital intermediate zone (fig. 1ad; table 1).
Supramarginal Area PF
The supramarginal area PF (area EK 65) is the

field that most clearly reveals the characteristics
of the parietal cortex just mentioned (fig. 33; cf.
also plates P72P74 in the Atlas [Economo and
Koskinas, 2008]). The total cortical thickness
varies between 3.03.5 mm.
Layer I is relatively rich in cells (table 7) and
has a denser superficial sublayer Ia and a deeper
sublayer Ib with fewer cells.
Layer II seems at a first glance to be extremely thick because it does not stand out well from
sublayer IIIa; it contains extremely small, true
granule cells.
Layer III contains numerous small cells,
mostly medium-sized pyramidal cells; there are
no sublayers, but rather, an equal distribution of
cells over the entire cross-section. There are some
examples of large pyramidal cells sporadically
disseminated in the deeper zones of this layer
that might enable us to differentiate, though
rather indistinctly, sublayers IIIa and IIIb. Cells
are typically arranged in very narrow, parallel,
perpendicular streaks, which appear more clearly
in layer III than in any other layer.
Layer IV is very striking with its thickness
and cellular plenitude. Cells are denser in the
middle of the layer, where they form a subzone
IVb1, which appears as a somewhat darker band
than either sublayer IVa or subzone IVb2. Just
like in layer IV of the superior parietal lobule, although not so clearly, we can discern more typical granule cells in the superficial sublayer IVa
than in the lower rows of subzones IVb1 and IVb2,
which are filled with triangular cells. Cells of layer IV partially participate in the radial striation.
Fig. 33. Supramarginal area PF (area EK 65). Dome of supramarginal gyrus in inferior parietal lobule. !45.
Parietal Lobe
89
Layer V is relatively thin; it is neither subdivided into distinct sublayers nor lighter stained
like at the superior parietal lobule and the postcentral gyrus. It is difficult to demarcate it precisely from layer VI, because its pyramidal cells
are scarcely larger than the spindle cells of the latter, and are almost as compactly arranged. A border between layers V and VI can be discerned
only under higher magnifications based on the
change in cellular form. The presence of a vertical
radial striation is suggested in both layers V and
VI.
Layer VI is clearly subdivided into an upper
sublayer VIa, with medium-sized and small spindle (fusiform) cells, and a lower sublayer VIb,
with only half the number of cells of about the
same proportions. There is difficulty in distinguishing layer VI from layer V; the border between layer VI and the white matter, on the other
hand, is very sharp and clear.
Regional PF Area Modifications

By taking into account the cortical thickness, the
distinctive narrow radial striation, and cell size,
various regional differences in the structure of
this area can be distinguished (fig. 1).
The cortex is thinnest at the opercular supramarginal area PFop (area EK 67), as well as at the
small secondary gyri, i.e. the tenuicortical supramarginal area PFt (area EK 66) (cf. plates P75 and
P77 in the Atlas [Economo and Koskinas, 2008]).
The size of cells is largest at the posterior dorsal
parts of the supramarginal gyrus in the magnocellular supramarginal area PFm (part of area EK
68). The columnar striation is most marked at the
transition to the superior temporal gyrus (T1), in
the supramarginal area columnata PFc (part of
area EK 68) (fig. 1a, c, d; cf. also plate P75 in the
At the gyral walls of the secondary rami of the
interparietal sulcus, which, as mentioned above,
are occupied by the caudal postcentral area PD ,
90
one encounters numerous structural features in

area PF that remind of area PD , and thus, we may
speak of a supramarginal-postcentral transition
area PFD in these precincts (fig. 1a, d). The transition part of the supramarginal gyrus in the
operculum of Rolando is also occupied by such a
transition formation, which, as mentioned earlier,
stretches as far as the agranular frontal area FB .
Function
The anterior part of the supramarginal gyrus is
considered to be the center for muscular sense
and stereognosis. Lesions of the posteroventral
part cause apraxic and aphasic disturbances.
Angular Area PG
In the region of the angular gyrus, the cortex is

still of a fair thickness, although it is somewhat
thinner than in the supramarginal gyrus; cortical thickness in the angular area PG (area EK 69)
varies from 2.9 to 3.3 mm. The cellular structure
is very similar to area PF, only that the deeper
sublayers of the overall thinner layer III become
again magnocellular and there is also a tendency
for lighter staining at the region of layer V. The
radial striation is rather wider than in area PF,
rendering the horizontal lamination somewhat
better visible (fig. 34; cf. also plate P76 in the Atlas [Economo and Koskinas, 2008]).
Layer I is subdivided into two sublayers, with
the upper sublayer appearing denser (table 7).
Layer II is somewhat thinner and poorer in
cells than in area PF, largely consisting of extremely small pyramidal cells.
Layer III is thinner, clearly striated with vertical cellular columns, and can again be subdivided into sublayers IIIa, IIIb and IIIc, containing
medium-sized cells. The largest cells are found at
the deepest zones of layer III. Cells are arranged
in somewhat wider vertical radial columns.
Fig. 34. Angular area PG (area EK 69). Dome of angular gyrus in inferior parietal lobule. !45.
Parietal Lobe
91
Layer IV shows a denser intermediate subzone IVb1, like in area PF. The upper sublayer IVa
is structured of small granule cells, yet there are
also numerous triangular cells among them,
which in the deeper zones form the layers main
cellular mass; subzone IVb2 presents a more dissolute cellular disposition. The radial striation is
also evident in layer IV, but less so than in the
other layers.
Layer V stands out by virtue of a somewhat
lighter staining, which, although not very distinct, stands out better here than in area PF. Layer V is poorer in cells than in area PF. These cells
are not quite as large as the spindle (fusiform)
cells that lie just beneath. In this area, layers V
and VI again participate in the radial striation in
certain parts. The differentiation of layer V from
VI is especially difficult at gyral walls, since the
size and density of cells there is about the same
(cf. VI+V mark on the lower right side of fig. 34).
Layer VI is thicker than in area PF, but less
sharply demarcated from the white matter. It
contains spindle (fusiform) cells in its upper part,
and fewer and smaller cells in its deeper sublayer.
Towards the transition from the parietal to the
occipital lobe, there is a regular appearance of
only individual examples of larger cells in layer V.
At the transition of the angular gyrus to the middle (T2) and inferior (T3) temporal gyri, the radial striation becomes coarser and the horizontal
lamination less obvious.
Function
Lesions in the field of the angular gyrus provoke
alexia, acalculia, ideomotor apraxia, as well as
agraphia. Deeper lesions, particularly in the
posterior part, also cause visuomotor disturbances. Flechsig [1895, 1920] localizes his large
posterior association center in the inferior parietal lobule.
92
Basal (Temporo-Occipital) Parietal Area PH
The occipitotemporal intermediate zone (already

mentioned at the beginning of this chapter) is
also vested throughout its extent by cortical
structural type 3, all the way to the trunk of the
calcarine sulcus and the base of the cerebral
hemispheres; this is termed basal (temporo-occipital) parietal area PH (fig. 1ad).
The field shows much the same structure as
area PF, i.e. in contrast to area PG, a more distinct
lamination and a narrower radial striation
(fig. 35). But layers II and IV are somewhat thinner than in area PF and contain more triangular
cells. Layers V and VI are even more difficult to
distinguish than in area PF and their borders
are irregular. The total cortical thickness is less
than in areas PF and PG, varying between 2.7 and
2.9 mm at the domes of gyri.
Layer I contains two sublayers, Ia and Ib, of
which the upper sublayer Ia is denser (table 7).
Layer II contains mostly small granule cells
in the upper zone, and mostly small pyramidal
forms in the deeper zone.
Layer III is less distinctly divisible into three
sublayers than in area PG, although more so than
in area PF. Cells are mostly medium-sized. The
radial striation is just as striking and fine as in
area PF (cf. fig. 33 with fig. 35).
Layer IV contains cells mostly triangular and
pyramidal; sublayers can no longer be seen. The
vertical striation is indistinct.
Layer V is relatively thick but hardly distinguishable from subjacent layer VI, with cells of
the two layers often intermingled at their border;
such a border thus appears as a wavy, or undulate,
line (cf. V/VI in the middle of fig. 35). The cells of
layer V are actually smaller in general than in layer VI, especially at more caudal parts of this intermediate zone; only seldom they assume a fine
pyramidal form and are rather disorderly arranged. The vertical striation is not very distinct
in layers V and VI, nonetheless it is still recognizable.
Fig. 35. Basal (temporo-occipital) parietal area PH (area EK 70). Dome of superior temporo-occipital intermediate
zone. !45.
Parietal Lobe
93
Layer VI has in its upper sublayer VIa cells

that are not very clearly spindle-shaped (fusiform). Sublayer VIb is somewhat poor in cells; the
border vis--vis the white matter is sharp.
Occipital and Temporal Transition Forms

Thus, the main characteristic of the basal parietal
area PH is the fusion of layers V and VI into a
single layer.
Toward the angular area PG, it assumes the
modification form called basal parietal area at
the parietal entrance PH P (area EK 70) [Economo
and Koskinas, 1925, pp. 588593], as shown in
figure 1a and in plate P78 in the Atlas [Economo
and Koskinas, 2008].
Toward the temporal lobe the radial striation
becomes coarser and cells of layers III and V are
again larger in size; in the rostral direction they
gradually become more distinctly slender pyramidal. This transition modification is denoted as
the basal parietal area at the temporal entrance
PH T (area EK 71), shown in figure 1a and in plate
P80 in the Atlas [Economo and Koskinas, 2008].
As already said, this basal area reaches on the
basal surface of the cerebral hemispheres beyond
the occipitotemporal sulcus, as far as the trunk of
the calcarine sulcus, where it can no longer be
94
distinguished from the fusiform area TF (area EK

87) of the temporal lobe, into which it merges
(fig. 1d).
Toward the occipital lobe, the cells of layer V
become even smaller and the layer even lighter;
sporadic isolated large cells make their appearance, rendering layer V again more distinct as
one passes in that direction. Layer III somewhat
diminishes in thickness toward the occipital lobe
and also contains only isolated large cells. This
transition modification is denoted as basal parietal area at the occipital entrance PH O (area EK
72), shown in figure 1a and in plate P79 in the
Atlas [Economo and Koskinas, 2008].
Function
In neuropathology, lesions of the temporo-occipital intermediate zone lead to amnesic aphasia,
certain disturbances of color perception such
as dyschromatopsia, and spatial disorientation.
There is one important consideration: the entire
inferior parietal lobe, and hence the temporo-occipital intermediate zone as well, are largely rudimentary in the animal including the simian
brain; thus, this region can be considered as a recent phylogenetic acquisition in humans.
Insular Lobe
Anatomic Subdivisions
The small insular lobe is hidden between the

frontal and parietal lobes, under the operculum,
with its slender gyri disposed radially in the form
of a fan (fig. 1a). The central insular sulcus, which
is the deepest, divides the insula into a larger rostral part, the anterior insula, and a smaller caudal
part, the posterior insula, which consists of 23
gyri. The central insular sulcus is found at the
imaginary extrapolation line of the central sulcus
of Rolando.
The anterior insula, with the large size of the
cells in its pyramidal layers and the feeble development of its granular layers, presents a characteristic cytoarchitecture that recalls the frontal
cortical structural type 1, whereas the posterior
insula, with its characteristically thick and parvicellular granular layers, recalls the parietal cortical structural type 3 (fig. 8). Both insular segments, but particularly the anterior part, are distinguished by a typically dense and robust layer
V, which is often so marked that in stained sections it can be seen as a blue band even by the naked eye.
The insula is further characterized by the
presence of the claustrum, a thin buckle of gray
matter lying deeply in the white matter at about
1 cm from the cortical surface, and running al-
most parallel to it. The fine flattened cells of the

claustrum are disposed horizontally and adjoined indirectly, by means of indistinct cellular
rows, with the deepest cells of layer VI of the insular cortex, without though being in direct continuity. The claustrum is fan-shaped as well, and
lies between the insular cortex and the corpus
striatum, touching rostrally the cellular mass of
the substantia perforata, and caudally the cells of
the nucleus of the amygdala.
The surface of the insula is oriented in a quasisagittal plane, and is covered by the parietal and
frontal opercula. The apex of the radial fan of the
insular gyri constitutes the insular pole. At its anterior extreme, along the dome of the first (rostralmost) short gyrus, the insula makes a medial
loop at almost a right angle, such that its surface
forms a coronal plane. That plane is small, triangular, and borders the anterior ramus of the circular insular sulcus, which separates the transverse insular gyrus (the medial continuation of
the falciform gyrus and the insular pole) from the
inferior frontal gyrus (F3) at the orbital surface.
At the same time, this coronal plane forms the
rostral wall of the transverse gyrus; it is vested by
frontoinsular area FJ (area EK 28) already discussed in the frontal lobe chapter which occupies the entire transverse gyrus.
The remaining areas of the insular cortex, on

its wider, sagittal surface, are primarily the precentral insular area IA, lying anterior to the central insular sulcus, and the postcentral insular
area IB , lying behind it. These areas cover with
their homotypic cortex the entire lateral surface
of the insula. Only the insular pole (fig. 1a, d) is
covered by a different, heterotypic cortex, the orbito-insular area IC (area EK 53), which resembles
in structure the area FJ of the transverse gyrus
and is also directly annexed to it, effecting the
transition to the base of the substantia perforata.
Area IC is only separated from the substantia perforata by a thin margin of gray matter that accompanies the lateral olfactory bulb as the lateral
olfactory gyrus, the so-called piriform insular
area ID (area EK 54), which shows the same cytoarchitectonic structure as the anterior part of the
lateral olfactory gyrus the frontal piriform area
FK (area EK 29) of which it forms the cortical
continuation (fig. 1a, d, 21).
Precentral Insular Area IA
This anterior insular zone is of the cortical structural type 2 (frontal type), with a thickness between 3.0 and 3.5 mm (fig. 36). At the dome, the
cortex displays a delicate radial striation, which
becomes more evident and coarser at the gyral
walls. Cells of layer III are voluminous and orderly arranged, like in the frontal lobe. Cells of
layer V are also conspicuous and larger than
those of layer VI, which is typical of the frontal
structural type.
Layer I is very thick (table 8).
Layer II mostly contains small pyramidal
cells.
Layer III shows, as in the frontal lobe, pyramidal cells orderly arranged in three secondary
sublayers, IIIa, IIIb and IIIc; in sublayer IIIc, besides the medium-sized cells, there are some rare
examples of large but very slender pyramidal
cells.
96
Layer IV occasionally becomes interrupted.

For being a granular layer, it appears rather poor
in cells (oligocellular), most of them triangular.
Such a cytoarchitectonic particularity of the
granular layers shows that this area is of the
frontal cortical structural type. In rostral parts
of area IA , both granular layers are less distinct;
they progressively become denser and more
conspicuous caudally to the central insular
sulcus.
Layer V is the most typical layer of area IA . It
is subdivided into two sublayers: the upper sublayer Va represents a dense, true insular girdle
(Inselgrtel) with voluminous cells arranged in
straight lines, like rows of soldiers; the looser sublayer Vb underneath hardly contains half the
number of cells, thus appearing lighter than sublayer Va.
Layer VI presents some difficulty in accurately determining its thickness, because it passes indistinctly into the white matter, and because the
cells of sublayer VIb gradually pass over into the
structure of the claustrum further deeply without
an actual interruption. Spindle (fusiform) cells of
the upper sublayer are larger than those of the
deeper sublayer. In figure 36, the claustrum is no
longer visible, but one can see that the entire white
matter is pervaded by spindle (fusiform) cells.
Modification and Transition Areas

Area IA covers the anterior central gyrus as well
as, immediately in front of it, both short insular
gyri, all the way to the accessory gyrus, which
forms the medial passage to the transverse gyrus;
at that boundary, area IA is delimited from the
agranular frontoinsular area FJ (area EK 28).
The dorsal precentral insular area IA1 (area EK
49), contains more voluminous cells but fewer
granule cells than the ventral precentral insular
area IA2 (area EK 50) (cf. plates P53 and P54 in the
Fig. 36. Precentral insular area IA (areas EK 49 and 50). Anterior insular lobe. Dome of precentral insular gyrus. !45.
Insular Lobe
97
Table 8. Summative table of quantitative data in two ground areas of the insular lobe. Separate dome and wall data
and some additional values supplemented from tables I, III, V and VI of Economo and Koskinas [1925, pp. 794801].
Area
symbol
Area name
Cortical
layer

mm

mm

mm
IA
precentral insular
area
I
II
IIIa
IIIb
IIIc
IV
Va
Vb
VIa
VIb
0.23
0.08
0.22
0.14
0.23
0.10
0.74
0.84
0.70
0.15
0.24
0.39
0.77
0.41
0.22
0.15
0.20
0.45
IB
postcentral insular
area
I
II
III
IV
Va
Vb
VIa
VIb
0.32
0.18
0.78
0.31
0.21
0.32
0.43
0.30
0.60
0.45
0.22
1.20
0.34
0.25
0.78
0.20
0.30
0.20
0.80
0.30
0.50
0.50
0.30
0.30
0.70
Cell content
cells/mm3
Cell size, m*
H(minmax)/
W(minmax)
8,000
90,000
35,000
25,000
30,000
70,000
50,000
25,000
25,000
12,000
510/57
510/57
1520/1215
2025/1520
1540/1015
315/57
2540/20
1520/710
2025/1215
1215/810
5,000
100,000
32,000
130,000
40,000
30,000
35,000
20,000
47/510
414/514
1240/718
46/46
20/1215
1020/715
20/8
15/8
Caudally, the precentral insular area IA gradually passes over the central insular sulcus into
the postcentral insular area IB as the transition
area IAB (fig. 1a; table 2; cf. also plate P55 in the
Atlas [Economo and Koskinas, 2008]); the latter
is characterized by a condensation of the granular layers and a reduction in the size of its pyramidal cells.
Postcentral Insular Area IB
In the postcentral insular area IB (area EK 51), the

cortex is diminished to a thickness of only 2.8
mm (fig. 37; cf. also plate P56 in the Atlas [Economo and Koskinas, 2008]). This area displays the
cortical structural type 3 and has a parieto-temporal character. The radial striation is slight; an
98
evident horizontal lamination ensues from its

dense granular layers. It generally contains more
numerous and smaller cells than the area IA .
Layer I is thicker than in area IA and poor in
cells or oligocellular (table 8).
Layer II is clearly thicker than in area IA and
richer, partly composed of granule and partly of
small polygonal and stellate cells.
Layer III is also thickened; it is subdivided,
albeit indistinctly, into two secondary sublayers;
in the deeper sublayer IIIb, only isolated larger
cells are found. Layer III shows a radial striation,
similar to the parietal lobe.
Layer IV is densely populated; it mostly contains true granule cells. The granular layer also
displays in part a certain radial striation. The upper cells of layer V virtually touch layer IV and
sometimes mingle with it.
Fig. 37. Postcentral insular area IB (area EK 51). Posterior insular lobe. Dome of postcentral insular gyrus. !45.
Insular Lobe
99
Layer V presents an upper sublayer of large

pyramidal cells; this cellular row constitutes a
continuation of the layer V girdle in the anterior
insula, which here becomes less typical. Figure 37,
on the left side, shows the increased cellular density of the upper sublayer Va; however, cells are
smaller than those of sublayer Va in figure 36. In
general, layer V cells are smaller in area IB than
in area IA .
Layer VI is relatively thin; it is rather rich in
cells, with more numerous and slender spindle
(fusiform) cells, compared to the anterior insular
gyrus. The deep sublayer VIb is better delimited
from the white matter than in the anterior insula;
but here cells show a less clear, gradual transition
to the claustrum (which is still not visible in
fig. 37). The white matter is also filled with strings
of fusiform cells stretching toward the claustrum.
Caudally to the posterior ramus of the circular
insular sulcus (margo superior insulae), one observes the transition form of the postcentral insular area at the temporal entrance IB T (area EK 52)
with a radial columnar striation; area IB T leads
into the temporal lobe (cf. plate P57 in the Atlas
Claustrum
In cross-sections, the claustrum appears as a gray

cellular band, 0.500.75 mm in thickness, composed of many rod-like cells, 9/25 m in size (H/
W), disposed horizontally; they are rather numerous and are accompanied by plentiful satellite cells. The gray mass of the claustrum appears
quite smooth at the surface facing the corpus striatum. On the other hand, the external surface
sends festooned processes outbound, corresponding to the gyral formation of the insula;
thus, this claustral surface assumes an undulate
form.
100
Insular Pole, Orbitoinsular Area IC and Insular

Piriform Area ID
All insular gyri converge at the insular pole, with

the two ground areas IA and IB gradually passing
into the orbitoinsular area IC (area EK 53) of the
insular pole or polar insular gyrus (fig. 1a, d; cf.
also plate P58 in the Atlas [Economo and Koskinas, 2008]); in this region the cortex is still 2.5
mm thick.
On the basal cerebral facies, the lateral olfactory gyrus is medially annexed to the piriform insular area ID (area EK 54) as a cortical margin, i.e.
as a boundary with the substantia perforata (cf.
2008]). In figure 21, area ID can be seen ending
against the posterior area of substantia perforata
TK (area EK 93). Toward that cortical boundary,
the cortex of areas IC and ID rapidly thins out in
all the layers except the molecular layer (layer I),
which actually appears thicker and seems to continue into the substantia perforata.
Layer I borders beneath with a thin layer II
that appears as a preserved band containing
small stellate and navicular cells, 10/10 m in
size (H/W), as in areas FJ and FK (fig. 22). At the
transition to the substantia perforata, these cells
form glomerular accumulations.
Layer III still contains pyramidal cells, 20/10
m in size (H/W) in area IC , but their axis is
no longer oriented perpendicularly to the surface of the cortex; further, they are not as regularly spaced. The number of cells and the
thickness of layer III gradually diminish in the
basal and medial direction; in the region of the
olfactory gyrus, i.e. in area ID , many acellular
patches are found, until layer III progressively
disappears.
Layer IV is the first to rapidly disappear in
area IC, even at the boundary between the two
typical insular areas (IA and IB) and area IC, such
that area IC can be already considered in reality a
heterotypic agranular cortex.
Layers V and VI also become gradually attenuated in these areas; the weaving of their cells
continually loosens, and layers finally stop at
the point where incoming olfactory myelinated
fibers penetrate the substantia perforata. With
these white matter bundles, some rows of cells
drag deeply into the tissue and meet cellular rows
deriving from the claustrum and the substantia
perforata.
As already mentioned, areas IC and ID delimit the insula from the substantia perforata in the
frontal cortical boundary: area IC from area FJ,
and area ID from area FK (fig. 21).
Insular Lobe
Function
The physiologic significance of the insula is incompletely understood. Its basal and orbital
parts, which bear a direct anatomic relation to
the lateral olfactory gyri, probably pertain somehow to olfaction. The other regions, according
to our views, can be ascribed to the rhinencephalon sensu lato, just like the limbic lobe. The
insula is in effect found in the same relation vis-vis the lateral olfactory root, just as the limbic
lobe is found vis--vis the medial olfactory root.
Certain facts favor the view that both the insula and the limbic lobe are cortical representations of the autonomic and sympathetic nervous
system.
101
Occipital Lobe
Cytoarchitectonic Boundaries
The occipital lobe extends immediately caudally

to the parietal lobe, separated from it by the parieto-occipital sulcus on the median (mid-sagittal)
hemispheric facies. Both on the superolateral
(convex) and the ventral (inferior) hemispheric
facies, the cytoarchitectonic boundaries of the
occipital lobe lie somewhat more caudally than
gross (macroscopic) anatomic boundaries. These
cytoarchitectonic boundaries coincide with the
posterior boundaries of the parietal lobe (discussed in detail in the respective chapter). The
occipital lobe essentially abuts the parietal lobe
on all its parts (fig. 1a, c), with the exception of a
small territory at the anteriormost end of the calcarine sulcus, where it touches the limbic lobe
(fig. 1b, d).
On the median hemispheric facies, the occipital lobe is divided by the calcarine sulcus into two
parts, the dorsally lying cuneus and the ventrobasally lying lingual gyrus. The calcarine sulcus is
T-branched into two perpendicular ramifications
at the posterior hemispheric pole, and thus reaches to a certain degree the lateral hemispheric convexity; the anterior end of the calcarine sulcus
forms, together with the parieto-occipital sulcus,
the calcarine trunk, which stretches to the limbic
gyrus. The floor, walls and lips of the calcarine
sulcus are covered by heterotypic granulous iso-
cortex, i.e. the koniocortex of the striate area OC

(area EK 79), which represents the sensory cortex
for vision (fig. 1a, b). In cross-sections, this can be
easily identified macroscopically by the white myelinated horizontal stria of Gennari [1782] or Vicq
dAzyr [1786] that traverses the gray matter.
Concentric Areal Disposition around the

Calcarine Sulcus
The remaining cortex of the occipital lobe was divided by Elliot Smith [1907] into two areas, concentrically disposed around the striate area of the
calcarine sulcus. The outer, wider area, which is
called peristriate area OA, resembles in many respects the parietal cortex and is located all along
the posterior boundary of the parietal lobe. The
inner area, called parastriate area OB , is concentrically placed within the peristriate area and directly enshrouds the striate area (fig. 1a, b, d).
General Characteristics of the Occipital Cortex
What characterizes the entire occipital cortex is

an altogether remarkable horizontal lamination,
and a low thickness, between 2.5 and 2.0 mm or
even less at the sulcus floors; the thinnest parts of
the occipital cortex (down to 1.2 mm) are reck-
Table 9. Summative table of quantitative data in three ground areas of the occipital lobe. Separate dome and wall
data and some additional values supplemented from tables I, III, V and VI of Economo and Koskinas [1925, pp. 794
801].
Area
symbol
Area name
Cortical
layer

mm

mm

mm
Cell content
cells/mm3
Cell size, m*
H(minmax)/
W(minmax)
OA
peristriate
area
I
II
IIIa
IIIb
IIIc
IV
V
VIa
VIb
0.20
0.24
0.18
0.25
0.18
0.23
0.66
0.53
0.60
0.22
0.39
0.43
0.34
0.20
0.20
0.26
0.18
0.20
0.35
0.35
0.26
6,000
80,000
60,000
45,000
50,000
150,000
40,000
55,000
12,000
7/5
10/6
10/8
20/12
1225/820
68/68
812/810
2030/810
1520/8
0.16
0.18
0.20
0.18
0.15
0.18
0.46
0.48
0.45
3,500
150,000
85,000
70,000
60,000
IV
V
0.18
0.26
0.18
0.26
0.18
0.22
240,000
70,000
VIa
VIb
0.32
0.26
0.18
0.10
0.25
0.18
85,000
25,000
910/45
410/410
612/610
615/612
20/15
75/20
610/68
412/412
45/25
1820/1012
15/5
I
II
III
IVa
IVb
0.19
0.13
0.27
0.17
0.28
0.22
0.15
0.32
0.18
0.26
0.19
0.12
0.27
0.18
0.28
9,000
150,000
100,000
150,000
60,000
IVc
V
0.37
0.27
0.30
0.19
0.34
0.27
220,000
45,000
VIa
VIb
0.26
0.42
0.20
0.15
0.25
0.20
140,000
20,000
OB
OC
parastriate
area
striate area
I
II
IIIa
IIIb
IIIc
5/5
6/6
710/6
5/5
5/5
25/50
8/8
718/715
60/30
615/10
20/10
oned among the thinnest territories of isocortex.

It is also typical for the occipital lobe, in contrast
to the neighboring parietal lobe, to have a lightlystaining layer V and a reduced cell size in that
layer. The size of cells in layer V falls below that
of cells in layer VI, which are small as well. None-
Occipital Lobe
theless, it is characteristic of the occipital lobe to

sporadically show large cells in layer V that appear individually and are distinctly filled with tigroid (Nissl) bodies. Individual large cells also
occur in the deeper zone of layer III in the occipital lobe; the cells of all the layers, and particu-
103
larly of layer III, are often arranged in thick short

cellular columns. Both granular layers are remarkably thick and cell-dense, considering the
overall thinness of the occipital cortex.
Peristriate Area OA
The peristriate area OA is spread over the lateral,

median and ventral hemispheric facies, immediately caudally to the parietal areas. On the lateral
hemispheric convexity, its cortex reaches a thickness of only 2.6 mm; in the median hemispheric
facies, 2.3 mm. It is rich in small cells, with sporadic larger cells, and exhibits a radial striation
with large intervals and a marked horizontal
lamination (clearly seen in fig. 38). The cortical
thinness becomes evident to the eye if one compares figure 38 with figure 33 or, even more so,
with figure 11. The columnar arrangement becomes very distinct.
Layer I is very thin (table 9).
Layer II is very dense, filled mostly with small
pyramidal cells.
Layer III is difficult to separate from layer II in
its upper border, as the compactness of both layers
at that border is very alike. At the border with layer IV one occasionally observes isolated, very
large pyramidal cells; however, such cells are so
few in number that they do not form an actual
sublayer, such that a sublayer IIIc is virtually lacking. The cells of layer III are arranged in wide vertical dispositions, which are somewhat more massive than in the parietal lobe, but nonetheless not
representing robust cellular columns yet; columns
only appear in sublayer IIIb and not in sublayer
IIIa, and extend to the deeper layers IV and V.
Layer IV is the most typical layer of this area,
due to its unusual richness in round and ovoid
granule cells. They form a band which appears
deep blue to the naked eye in the stained specimens. This layer also displays a radial striation.
Layer V is very thin, lightly staining and with
small cells, hardly larger than those of layer IV;
104
the small cells are mostly polygonal, triangular or

fusiform. Finer and larger pyramidal cells appear
rather regularly, but only isolated. Often, and particularly at the gyral walls, the cells of layer V in
area OA are grouped together in wider and shorter columnar packages, projecting into the lower
part of layer III through layer IV, but also into layer VI beneath (double arrow in fig. 38). I emphasize the progressive decrease in cell size in layer V
in caudal parts of the hemispheres, in the vicinity
of the occipital pole; as we saw, the bulk of the
cells in layer V are already smaller than those in
layer VI at the posteriormost sections of the basal
parietal lobe. Such a situation is even more pronounced in the occipital lobe, allowing its easy
identification in the histologic tissue sections.
The lightly staining layer V is a characteristic of
the occipital lobe, as opposed to the structure of
the inferior parietal fields, and helps delimit the
cytoarchitectonic boundaries of the two lobes.
Layer VI differs from layer V in being rather
cell-dense, especially in the upper sublayer VIa,
which contains spindle (fusiform) cells. Sublayer
VIb is somewhat looser and thinner, yet its demarcation from the white matter is clearer compared to the parietal lobe. Here as well, and particularly in the gyral walls (lower left field of
fig. 38), the cells are distinctly grouped into thick
short columns, often in such a way that the columns alternate in certain localities with those of
layer V like a chess-board, or even forming peculiar S-shaped sockets.
Cytoarchitectonic Modifications of Peristriate

Area OA
This cytoarchitectonic picture of area OA is not
identical throughout its entire extent. On the lateral hemispheric facies, and in the vicinity of the
parietal lobe, as well as in the parieto-occipital
sulcus, we find greater numbers of larger cells in
the deeper part of layer III in area OA, such that
one might occasionally speak of a sublayer IIIc.
Fig. 38. Peristriate area OA (area EK 74). Dome and wall of the superior sagittal gyrus of the cuneus. !45.
Occipital Lobe
105
We have identified three modifications: the

posterior peristriate area OA1 (area EK 74), which
mostly occupies the inner part of area OA in the
vicinity of the parastriate area OB (area EK 76);
the anterior peristriate area OA2 (area EK 73),
which courses upwards the rear wall of the parieto-occipital sulcus, the anterior upper part of
the cuneus and a small stripe on the lateral hemispheric convexity caudally from the primary occipital sulcus and the transverse (parietal) occipital sulcus, all the way to the superomedial edge
of the hemisphere; and the magnocellular peristriate area OAm (area EK 75), covering a very
small field above the superior sagittal sulcus of
the cuneus on the median hemispheric facies and
between the secondary and tertiary occipital sulci (fig. 1ac; cf. also plates P81P83 in the Atlas
The histologic picture of area OA, as far as size
of cells and distinct columnar arrangement are
concerned, varies among gyri, with irregular
changes and substantial individual differences.
Parastriate Area OB
The parastriate area OB (plates P84P87 in the

Atlas [Economo and Koskinas, 2008]) lies concentrically within the area OA, being surrounded
by the latter on all sides in the form of a shell
(fig. 1a, b, d). On the lateral hemispheric facies,
the area OB forms only a narrow band within the
so-called semilunar sulcus, which embraces from
the outside the descending gyrus of Ecker [1869,
1873]; on the median hemispheric facies, the area
OB is somewhat thicker and reaches inbound up
to both lips of the calcarine sulcus, at loci that are
left free by the striate area OC, as I detail later. To
the outside, the area OB reaches dorsally almost
to the inferior sagittal sulcus of the cuneus, and
ventrally almost to the lingual sulcus with a
certain margin of individual variations. The cortex in this area has a thickness of only 1.82.0
mm (fig. 39), further dropping to 1.5 mm at the
106
gyral walls; it is therefore very thin, ranking

among the thinnest cerebral cortical areas. Nonetheless, the cortex is highly cellular and shows an
even more lightly-staining layer V than area OA .
The boundary of area OB vis--vis area OA is not
sharp at all and difficult to trace (plate P83 in the
Atlas [Economo and Koskinas, 2008]); on the
contrary, its delimitation from the calcarine field
OC is razor-sharp across the entire course of this
transition zone to the occipital pole and to the
median hemispheric facies (cf. plates P85P87 in
Layer I is very thin; it contains relatively large
and slender cells (table 9).
Layer II is extremely dense and intimately
fused with the upper parts of layer III, such that
one has difficulty distinguishing the two. One gets
the impression that they form a single, 0.4 mm
thick granular layer, were it not possible to draw a
sharper border between them under a higher magnification. This thick layer of small cells in layer II
and in sublayer IIIa, which forms a cornice over
the columnar mass of the remaining layer III, is
the most characteristic feature of area OB. Layer II
contains granule and triangular cells of a dimension that elsewhere is only found in layer III.
Layer III is rather thin, especially in comparison with its thickness in the parietal lobe. It comprises sublayers IIIa, IIIb and IIIc. Sublayer IIIa
can hardly be delimited from layer II, as mentioned above, because it contains cells to a certain
extent of a similar size as layer II, although sublayer IIIa contains more and larger pyramidal
cells. Sublayer IIIb appears somewhat lighter. Sublayer IIIc contains medium-sized and pyramidal
cells that may exceptionally reach a very large size
(middle of the field in fig. 39). Cells of layer III are
often arranged in radiating short columns, whose
base is formed by the large pyramidal cells of sublayer IIIc; the body of the columns is formed by
medium-sized cells in sublayer IIIb; the beams of
the cornice are formed by small cells of sublayer
IIIa and layer II together, an arrangement that
constitutes the typical picture of this area.
Fig. 39. Parastriate area OB (area EK 76). Wall of the superior sagittal gyrus of the cuneus. !40.
Layer IV is very dense, perhaps the densest

cellular layer of the entire cerebral cortex. It contains mostly round or ovoid, relatively large granule cells that stain very deeply, revealing a dark
horizontal stripe in the tissue sections.
Layer V is thin, a light band compared to layer
IV; most of its cells are even smaller than the
granule cells of layer IV, and definitely smaller
than the cells of layer VI. On rare occasions, one
may encounter large pyramidal cells filled with
tigroid (Nissl) bodies; they almost look like giant
cells amidst the pigmy elements in their vicinity.
Layer VI shows a dense, thin band in its upper
sublayer; the cells are mostly triangular, and a
few are spindle-shaped (fusiform). The lower,
lighter sublayer VIb has a looser structure and
contains more numerous spindle cells; it is very
Occipital Lobe
thin and clearly demarcated from the white matter.

In many fields of area OB , the cells of layers V
and VI are also arranged in groups similar to the
arrangement described above for area OA; this
can be seen at the lower part of figure 39, and at
the lower left corner of figure 40.
Giant Pyramidal Parastriate Boundary OB,

a Parasensory Zone
As mentioned earlier, the outer boundaries between areas OB and OA are difficult to define at
loci where the latter also contains large cells
(modification OAm). On the other hand, the inner
boundary with area OC at the lips of the calcarine
107
sulcus is the most exemplary type of an absolutely razor-sharp boundary between two areas (asterisk in fig. 40). Immediately in front of that
boundary line, and all along the transition of area
OB into the cortex of the calcarine sulcus in the
median hemispheric facies and the occipital pole,
one encounters a confined collection of unusually large giant pyramidal cells in sublayer IIIc of
area OB within a transition line at the most 1.0
2.0 mm thick that may reach 70/25 m (H/W)
in size (center field in fig. 40). That boundary
with striate area OC is called the giant pyramidal
parastriate boundary OB (area EK 77).
In earlier discussing the sensory tactile center
of postcentral area PA, I already remarked that, in
the immediate vicinity, i.e. the boundary of any
koniocortex (or sensory isocortex), one finds a
zone with such giant cells, which I call a parasensory zone.3 The functional significance of such
zones remains incompletely understood at present, but they probably have a stake in attention
warning, an adaptation reflex to sensory excitation. This boundary OB (plates P85P87 in the
Atlas [Economo and Koskinas, 2008]) is therefore the parasensory zone of the sensory field of
vision, i.e. of the OC koniocortex.
Striate Area OC
On the inner side of area OB, the cortical structure changes abruptly into the striate area OC
(area EK 79) as can be distinctly seen in the
middle of figure 40 to a granulization of most
cortical elements. On the other hand, in the left
field of figure 40, one sees the wide cellular columns and large, robust cells of area OB ; in the
central field, the giant cells of area OB; and in
3
This concept was formally presented by Economo [1926g]
at the Viennese Society for Psychiatry and Neurology on March
9, 1926, and further elaborated upon in an article [Economo,
1928a] published the year after the German edition of this
book.
108
the right field of figure 40, the picture only contains numerous small cells, hence being koniocortex. Moreover, the horizontal lamination is accentuated, and the number of layers increases,
such that suddenly four dark granular layers appear on the right, separated by four lightly stained
layers. Meynert [1872a, b] considered this region
a cortex with eight layers (octalaminar), four of
which are granular, and thus understood its sensory role.
Layer II is clearly demarcated, but not reinforced. Layer III is thin and only contains small
cells. By contrast, a most striking point is that layer IV is split into two, as a result of the appearance
of a white, horizontal band inside it, the so-called
stria of Gennari (asterisk in fig. 40), stretching
from the center of the figure, where giant pyramidal cells are still seen in layer IIIc, towards the
right as a light, acellular band (sublayer IVb) ascending somewhat obliquely. The lightly staining
stria of Gennari divides layer IV into an upper
granular sublayer IVa, a middle sublayer IVb (the
actual light stria of Gennari), and a lower sublayer
IVc with darkly staining granule cells. Through
such a division, sublayer IVa ends up lying higher
and closer to the cortical surface, whereas sublayer IVc lies somewhat deeper than usual.
Layer V lies directly beneath sublayer IVc and
also stains lightly. On the other hand, the even
deeper sublayer VIa contains such small and
dense cell elements, that Meynert [1872a, b] considered it as the deepest, i.e. the fourth granular
layer; sublayer VIb, just beneath, again appears
more lightly stained. Layer VI also displays a
quite abrupt passage from area OB to area OC,
becoming denser at that boundary, and better
marked off from layer V and from the white matter. In this location, the overall cortical thickness
is further reduced.
The transition from area OB to area OC that I
just mentioned occurs rather abruptly and suddenly at the parastriate boundary OB, directly
behind its giant pyramidal cells; the granulous
cytoarchitectonic type of koniocortex in area OC
Fig. 40. Parastriate area OB (area EK 76), giant pyramidal parastriate boundary OB (area EK 77) and striate area OC
(area EK 79). Upper lip of calcarine sulcus. The asterisk (*) denotes the transition from the parastriate to the striate
area. !40.
then covers both walls, the floor and large parts

of the lips of the calcarine sulcus through its entire extent, and to a lesser degree, the dome of the
occipital pole (fig. 1a, b, d). The cortical thickness
in the striate area varies from 2.5 to 2.0 mm and
falls to 1.5 mm at the walls. Together with area
OB , area OC ranks among the thinnest regions of
the cerebral cortex (fig. 6a, b). Figure 41, taken
from the lower wall of the calcarine sulcus, depicts in detail the full cytoarchitectonic picture of
striate area OC (cf. also plates P85P88 in the Atlas [Economo and Koskinas, 2008]).
Layer I is rather thin, but generally thicker
than in area OB . It contains triangular cells for
the most part (table 9).
Occipital Lobe
Layer II is also very thin, when strictly defined vis--vis layer III; it is rather rich in smaller
pyramidal cells, arranged in not more than 23
rows.
Layer III underneath is very thin as well. It
has a very high density of pyramidal cells, such
that layer III does not look like a compact granule
cell layer, but as a lighter stripe, contrasting with
the darker granular layers II and IV (fig. 40).
Layer IV reaches the maximum thickness for
the entire cerebral cortex. Its upper sublayer, the
superficial internal granular layer IVa, is poorly
delimited vis--vis layer III; it is only recognizable from the form of its cells, which are very
small round granule cells. The sublayer IVb, di-
109
Fig. 41. Striate area OC (area EK 79). Wall of calcarine sulcus. !45.
rectly beneath, is called the intermediate layer or

stria of Gennari; it is much lighter in color and
clearly defined from top and bottom. It contains
cells of the same caliber as those in sublayer IVa,
also staining very lightly. Additionally, in the
stria of Gennari, one also finds single examples
(12 every 100 m in the horizontal direction) of
the so-called giant stellate or solitary cells of
Meynert, disposed horizontally. Ramn y Cajal
[19001906, 1921, 1923] considers these specifi-
110
cally visual cells, and thinks that the terminal arborizations of the optic radiation, whose fibers
first constitute the stria of Gennari, terminate
upon them. In the cortical sections, these cells
usually do not appear as stellate, because they are
arranged horizontally and they are cut through
their thinnest dimension. They merit the term giant cells only in comparison with the size of the
surrounding granule cells and other small stellate cells of sublayer IVb. In figure 40 and 41 they
appear as somewhat larger black points, barely

discernible from the other granule cells. Beneath
the lightly staining sublayer IVb, one finds sublayer IVc, the deep internal granular layer, which
is the richest and densest granular layer of the
striate area, and the second most compact layer
in the entire cortex after layer IV of area OB . For
further details regarding the growth of sublayer
IVa through a granulization of the lower parts of
layer III, see Economo and Koskinas [1925, pp.
627628].
Layer V is very thin in comparison with layer
IV; it can be seen by the naked eye as a lighter
band in the stained preparations. While in fresh
cortical specimens the stria of Gennari, or sublayer IVb, strikes the observer as the lightest layer due to its myelinated fibers coursing amidst
the gray matter, in stained preparations it is layer
V that appears as the lightest of all the horizontal
layers (fig. 41); it is in effect very poor in cells,
compared to the dense layers in its vicinity, and
even poorer, and therefore lighter in color, than
sublayer IVb. Its upper and lower parts, which respectively border on the cell-rich sublayers IVc
and VIa, contain more numerous cells. A further
characteristic of layer V is the regular appearance
of sporadic, large pyramidal cells, filled with
chromophilic Nissl (tigroid) bodies. One finds
these so-called giant cells of Meynert throughout
the entire striate area in the middle parts of layer
V, either solitary or in groups of two or three at
intervals about 12 mm apart. These cells are
seen in layer V of figure 41 as distinct small black
triangles, and can only be discerned as giant cells
relative to the smaller roundish cells that surround them.
Layer VI has an upper sublayer VIa, a dense
cellular band consisting of compactly-positioned
small pyramidal cells. The small size and density
of these cells can easily give the impression of a
fourth granular layer, as Meynert [1872a, b] had
noted. Very few spindle (fusiform) cells appear
isolated among those pyramidal cells. The cellpoor (oligocellular), thick sublayer VIb contains
Occipital Lobe
more spindle cells than sublayer VIa, of small and

medium sizes, mostly arranged horizontally and
revealing a perpendicular or radial disposition
only at the dome adjoining the calcarine lips. The
boundaries with the white matter are very distinct.
Extension of Striate Area OC

With its particular koniocortical structure, the
striate area covers the floor, both walls and lips of
the calcarine sulcus. Rostrally, the territory of the
striate area does begin at the hemispheric surface, but at a promontory directed far forward at
the floor of the calcarine trunk and the parietooccipital sulcus, just about where the converging
trunk impinges on the limbic gyrus to form the
isthmus, covering slightly more the lower wall.
Usually, the striate area does not quite reach as
far as the anteriormost part of the trunk of the
calcarine sulcus; it stops at a point that corresponds to an imaginary frontal plane falling into
the splenium of the corpus callosum.
Caudally, the striate area OC becomes gradually enlarged, first on the ventral, lingual wall,
and then on the dorsal, cuneal wall of the trunk,
continually occupying greater districts of both.
At the junction of the calcarine sulcus with the
parieto-occipital sulcus, area OC outflows to the
ventral lip of the calcarine sulcus, i.e. the brink
of the lingual gyrus, where it appears over a
short expansion on the median hemispheric facies; at the dorsal or cuneal lips of the calcarine
sulcus, area OC only extends to the free surface
somewhat caudally (fig. 1b). Subsequently, the
striate area progressively clothes the domes of
the lingual and cuneal gyri, first as a small
boundary, and then, more caudally, in increasing thickness. That way, the lingual gyrus is
consistently covered more than the cuneus. Finally, area OC surrounds the T-shaped ramification of the calcarine sulcus at the occipital pole
like a wreath.
111
Thus, area OC is also apparent on the lateral

facies (convexity) of the human cerebral hemisphere, at least at the occipital pole; usually, it
only reaches the middle of the dome of the descending gyrus of Ecker, where it again abuts area
OB. In primates, on the other hand, the striate
area extends far on to the lateral hemispheric
convexity, i.e. as far as the so-called simian fissure
(Affenspalte); consequently, one can say that the
semilunar sulcus (sulcus lunatus), which frames
the gyrus of Ecker on the lateral hemispheric
convexity, is probably the human homolog4 of the
simian fissure of Elliot Smith [1904a].
Multilaminar Tendency of Area OC

In studying closer the layer disposition of the individual cellular forms in the striate area, one
readily distinguishes an even more detailed laminar division of the entire occipital cortex, were
one to enter into greater detail. Thus, one may e.g.
further break sublayers IVb, IVc and VIa down to
three secondary subzones each, based on cell
type, size and density. Layer V can be subdivided
into a further two sublayers. Therefore, in this
case, one can speak of area OC as being stratified
into 16 cortical layers (i.e. being hexakaidecalaminar).
Function
The Striate Area Is the Koniocortex of the

Primary Visual Domain
The heterotypic granulous cortex of striate area
OC is the visual koniocortex, i.e. the sensory cortex where the first retinal impressions deriving
from lower centers reach the cerebral hemi4
This issue was detailed in a subsequent study of Economo

[1930b], entitled On the question of the presence of the simian
fissure in humans in the light of cytoarchitectonics.
112
spheres. Thus, such a cortex constitutes the visual cortical domain par excellence, and this is an
indisputable fact. It is equally well known that
territories of area OC occupying the wall and the
dome of the cuneus correspond to the lower sector of the contralateral field of vision, whereas the
wall and dome of the lingual gyrus correspond to
the upper sector of the contralateral field of vision from both eyes. It is further acknowledged
that the polar part of area OC responds to macular vision; parts lying more anteriorly correspond
to the remaining (peripheral) visual field. However, another view holds that the entire floor of
the calcarine sulcus participates in macular vision.
The subdivision of the internal granular layer
into two secondary sublayers by the stria of Gennari has provided the basis for a theory of binocular vision by Brny [1925] and Kleist [1926].
Such a disposition occurs in humans and all animals with a binocular vision, whereas animals
with laterally placed eyes, i.e. with monocular vision, do not present with such a split of layer IV;
Volkmann [1926] has reported that a partial division of this layer can be found in animals with
partial binocular vision. One tends to conclude
that the two granular sublayers IVa and IVc issued from such a split are the cortical stations
that respond to the homonymous retinal sectors
of the opposite field from both eyes, and that the
intracortical superimposition of those two layers
translates the fusion of the corresponding retinal
images from the two eyes into one impression.
The thicker and denser sublayer IVc corresponds
to the contralateral, and the sublayer IVa to the
ipsilateral eye.
According to Lenz [1921], it is also possible
that primary color perception takes place in the
striate area OC. It is still questioned whether the
area OC is solely responsible for visual reception:
one finds fields in area OB whose structure recalls the koniocortex, by a patchy increase in cell
density and decrease in cell size (evidently, without a splitting of layer IV by a stria of Gennari).
We have designated those territories as maculae

granulosae of parastriate area OB (area EK 78),
without, however, prejudging their physiologic
role (plate P84 in the Atlas [Economo and Koskinas, 2008]).
cortex. Also, the striate area is more than four

times richer in cells than any other cortical region [Economo, 1926c].
Physiologic Significance of Areas OA and OB

Cellular Density of Area OC
The total surface of the visual koniocortex (area
OC) in both hemispheres together is most likely
about 50 cm2. The total number of cells in the
striate area bilaterally is around 1.4 !109, i.e. 10%
of the total number of cells in the entire cerebral
Occipital Lobe
It is possible that area OA responds to visual

memory, while area OB and its gigantocellular
zone OB have perhaps an associative function
on the one hand, and a reflective and motor function on the other. More details on this theme are
given in our larger work [Economo and Koskinas, 1925, pp. 653657].
113
Temporal Lobe
Cytoarchitectonic Boundaries
With regard to the temporal lobe, again, it is only

the study of the cellular structure of its cortex
that provides the necessary elements for an exact
anatomic demarcation from its surroundings.
The cytoarchitectonic boundaries of the temporal lobe are in fact somewhat more restricted
than the macroscopic (gross anatomic) boundaries: its posterior segment extends beyond the caudal extreme of the lateral (Sylvian) fissure, and belongs, with a cytoarchitectonic criterion, to the
parietal type of cortex (intermediate temporo-occipital area PH ) already described in the parietal
lobe chapter. In all other respects, the actual temporal lobe extends from the temporoinsular sulcus (margo insulae posterior), and from the floor
of the lateral (Sylvian) fissure over the ventral aspect of the lateral convexity and the base of the
cerebral hemispheres up to the hippocampal gyrus or the occipitotemporal sulcus (fig. 1a, b, d). In
this entire territory, the cortex proves to be quite
thick, marked by a progressive increase in thickness as it approaches the temporal pole (fig. 6).
The temporal lobe is composed of numerous
different cytoarchitectonic areas. I already emphasized in the parietal lobe chapter that the cellular composition of the superior (T1) and fusiform (T4) temporal gyri (fig. 43) is cytoarchitec-
tonically very similar to the cortical structural

type 3 of the inferior parietal region (fig. 33). Only
the middle (T2) and inferior (T3) temporal gyri
exhibit a pure temporal structure (fig. 45). Nevertheless, T1, T4, and all the remaining gyri of the
temporal lobe present sufficient characteristic cytoarchitectonic features to allow a clear identification of their typical temporal character, despite the masked regional particularities of the
diverse area modifications and their striking similarities to the parietal cortical structural type.
General Cytoarchitectonic Characteristics
In brief, these distinguishing characteristic features are the marked overall thickness of the cortex and the marked thickness of layer I; the progressive attenuation of the thickness of both
granular layers toward the temporal pole (fig. 7);
the peculiar appearance of layer II as frayed or irregularly interrupted, with clusters or bouquets
of cells tending to protrude into layer I; an aspect
of layer IV, which is characteristically divided by
bundles of incoming myelinated fibers into vertical columns of granule cells, separated by intercolumnar, acellular intervals up to 25 m wide
(granular columns are not contiguous to each
other, in contrast to the parietal lobe, where layer
IV is also radially segmented, but with the granular columns maintaining contiguity).
Layer III has generally larger, but fewer cells,
and is thinner than in the parietal lobe; this layer
also has the peculiarity of becoming not only relatively, but absolutely thinner at the dome of the
gyri compared to the walls.
Layers V and VI, on the other hand, are remarkably massive, especially compared to the
same layers in the parietal and occipital lobes,
where they lose much of their importance. Thus,
in thickness, cell density and cell robustness, layers V and VI surpass the upper layers, and stand
out most prominently in the cytoarchitectonic
picture, a feature that immediately differentiates
the temporal cortex from the granular structural
types of the frontal lobe.
Finally, the entire temporal cortex uniformly
shows a clearly demarcated and somewhat coarser radial striation, which may extend from layer
VI all the way up to layer II.
Through one or more of these criteria, the cortical areas of the temporal lobe can be recognized
correctly and consistently.
Areas and Regions of the Temporal Lobe
As many as five cytoarchitectonic areas can be

distinguished in T1 alone, namely, areas TA, TB ,
TC, TD and parts of TG. The modifications of area
TE are found in T2 and T3. Areas TF and TH belong to T4. Areas TJ, TK and parts of TG are found
in the temporopolar region (fig. 1a, b, d; table 1).
Superior Temporal Area TA
The superior temporal area TA comprises the major part of T1 (fig. 1a, d); it invests the entire long
dome on its lateral hemispheric aspect and its
lower wall, with the exception of its polar (anterior) segment, which is occupied by area TG, and
the dorsal surface of the lateral (Sylvian) fissure,
Temporal Lobe
which is occupied by areas TB , TC and TD. The

cortex of area TA has an average thickness of
about 3.0 mm. It is very similar to the structural
cortical type found in the inferior parietal region,
but the radial striation of area TA is somewhat
coarser and more marked than that of, for example, area PF ; such a striation stretches, as already
mentioned, from layer VI into layer II, segmenting layer IV into vertical columns (organ pipe formation) (fig. 42), with the cortex further showing
certain additional temporal characteristics.
Layer I is rather thick (table 10), with cells
generally assuming a radial orientation.
Layer II is rather thin, also not dense in cells,
and it appears peculiarly frayed, i.e. with alternating partial gaps and bush-like forms, protruding into layer I; these denser areas usually contain
cells of a triangular shape; the gaps are again very
poor in cells (denoted by a plus sign in fig. 42).
Layer III is rather thick, with such thickness
(parietal type) distinguishing the cortex of T1
from the true temporal formations of T2 and T3
(which have a thinner layer III); it also resembles
the cortex of the parietal lobe in possessing
throughout, and rather uniformly, medium-sized
pyramidal cells, such that a sublayer IIIc is hardly visible; one may only discern a clear sublayer
IIIc on the right half of figure 42, at the wall of the
superior temporal sulcus. Cells are arranged in
vertical columns somewhat wider than in the parietal lobe; they often appear to continue downwards in cellular columns passing through layer
IV, and even through layers V and VI, giving a
radial appearance. Some cellular columns apparently traverse all layers (denoted by the arrow in
fig. 42).
Layer IV is relatively thin and mostly contains small pyramidal cells grouped in small vertical columns about 50 m wide and 150 m
high, separated by narrower, acellular intervals.
Occasionally, these cellular columns extend into
the lower zone of layer III and the upper zone of
layer V.
115
Fig. 42. Superior temporal area TA . Ventral brink of the dome at the caudal part of the superior temporal gyrus (T1),
showing its posterior modification TA1 (area EK 80). !45.
116
Table 10. Summative table of quantitative data in seven ground areas of the temporal lobe. Overall layer thickness
based on the present work. Separate dome and wall data and some additional values supplemented from tables I,
III, V and VI in Economo and Koskinas [1925, pp. 794801].
Cell size, m*
H(minmax)/
W(minmax)
Area
symbol
Area name
Cortical
layer

mm

mm

mm
TA
superior temporal
area
I
II
III
0.23
0.18
0.86
0.25
0.16
0.69
0.22
0.10
0.95
7,000
85,000
40,000
IV
V
VIa
VIb
0.19
0.49
0.45
0.43
0.19
0.33
0.31
0.16
0.18
0.50
100,000
40,000
30,000
15,000
I
II
IIIa
IIIb
IIIc
0.24
0.20
0.25
0.15
0.23
0.20
0.88
0.80
0.90
IV
V
0.37
0.40
0.45
0.40
0.35
0.40
125,000
50,000
VIa
VIb
0.53
0.49
0.40
0.20
0.80
40,000
20,000
I
II
III
0.26
0.28
0.74
0.23
0.25
0.70
0.26
0.30
0.70
15,000
120,000
70,000
IV
V
VIa
VIb
0.45
0.53
0.39
0.25
0.40
0.55
0.35
0.20
0.55
0.50
150,000
40,000
40,000
15,000
7/7
58/58
810/810
40/25
47/47
9/8
20/9
12/8
I
II
III
IV
V
VIa
VIb
0.22
0.22
0.80
0.30
0.50
0.40
0.30
0.21
0.23
0.75
0.30
0.45
0.35
0.22
4,000
120,000
35,000
140,000
60,000
40,000
20,000
8/7
45/5
912/710
57/67
7/7
20/15
20/10
I
II
IIIa
IIIb
IIIc
0.24
0.17
0.24
0.20
0.25
0.15
0.86
0.95
0.75
10,000
85,000
35,000
25,000
30,000
IV
V
0.24
0.69
0.22
0.50
0.24
0.80
90,000
50,000
VIa
VIb
0.76
0.59
0.53
0.43
1.40
40,000
20,000
610/68
610/712
15/12
18/15
30/18
50/30
6/6
1020/815
2040/1520
3040/1520
2030/810
TB
TC
TD
TE
supratemporal
area simplex
supratemporal
area granulosa
intercalated supratemporal area
temporal area
proper
Temporal Lobe
0.90
0.60
0.20
0.23
0.80
0.35
0.50
0.75
Cell content
cells/mm3
5,000
80,000
55,000
30,000
33,000
8/7
6/5
1525/815
50/30
8/7
2030/1020
25/10
17/9
7/6
715/710
12/10
22/17
1025/818
3560/2530
9/8
15/10
50/25
25/15
1518/10
117
Table 10 (continued)
Area
symbol
Area name
Cortical
layer

mm

mm

mm
Cell content
cells/mm3
Cell size, m*
H(minmax)/
W(minmax)
TF
fusiform area
I
II
0.26
0.20
0.24
0.20
0.25
0.20
8,000
100,000
III
IV
Va
Vb
VIa
VIb
1.04
0.20
0.64
0.20
1.00
0.20
0.50
0.56
0.50
0.48
0.26
0.40
0.24
45,000
100,000
50,000
35,000
28,000
15,000
9/9
6/6
12/10
15/12
6/6
1012/1012
2540/1820
2030/1820
1220/1012
0.29
0.09
1.09
0.17
0.70
0.77
0.68
0.34
0.12
0.86
0.08
0.60
0.80
0.80
TG
temporopolar area
I
II
III
IV
V
VIa
VIb
0.75
0.30
0.08
1.20
0.15
0.80
1.50
8,000
60,000 (?)
25,000
50,000
35,000
40,000
12,000
10/7
520/510
1520/1220
10/8
2530/1218
30/20
20/10
Layer V shows a constant increase in thickness

and cell size from its caudal parietal boundary toward the temporal pole. It mostly contains medium-sized pyramidal cells, well formed, in contrast to the inferior parietal lobe; these cells are
also disposed into wide, vertical columns, separated by interposed bundles of myelinated fibers,
although such an arrangement is less striking
than in layer III. In the posterior part (area TA1),
cells are smaller and less robust, whereas towards
the pole (area TA2), true pyramidal forms predominate, being larger than those of layer VI.
Layer VI is subdivided into sublayers VIa and
VIb, with a 2: 1 ratio in spindle (fusiform) cell
number; the cellular groups of layer VI are distinctly separated by bundles of myelinated fibers.
In the posterior modification (TA1), layer VI still
contains numerous triangular cells; in the anterior modification (TA2), more typically spindleshaped cells. The anterior modification TA2
118
shows a less conspicuous radial striation than the

posterior modification TA1. The border between
these two modifications is at about the acoustic
sulcus, which corresponds superficially to the beginning of the gyrus of Heschl. The demarcation
of the cortex from the white matter is not very
sharp.
Posteriorly, area TA passes gradually, via the
transition gyrus that caudally secludes the lateral
(Sylvian) fissure, into the supramarginal gyrus,
i.e. area PF, which it resembles cytoarchitectonically in any event (fig. 1a). Anteriorly (fig. 1a), it
extends to a point about halfway between the rostral extremity of the temporal pole and the acoustic sulcus, and rises somewhat, both frontally and
caudally, on to the dorsal wall of T1. Moreover, in
the superior temporal sulcus, it occupies almost
the entire ventral wall of T1; in this region, the columnar organization of area TA is more evident,
and the cells of layer III are of a larger caliber.
A modification with magnocellular characteristics (area TAm) and striking cytoarchitectonic similarities to area TB is described on the ventral (inferior) surface of T1, opposite from the
transverse gyrus of Heschl [Economo and Koskinas, 1925, p. 686].
Function
Pathophysiologic studies localize the center for
the understanding of word phonetics (verbal audition) in the cortex of the posterior superior temporal area TA1 (area EK 80) of the left side (plates
P88b, P89 and P92 in the Atlas [Economo and
Koskinas, 2008]); the center for the understanding of word meaning (verbal cognition) in the caudal transition area from TA1 to PF ; and the center
for the understanding of music (musical intelligence) in the anterior superior temporal area TA2
(area EK 81, further subdivided into fields TA2
and TA2 by Economo and Horn [1930], cf. table 2
and fig. 68), and perhaps even, with some question, in the temporal pole.
Magnocellular Supratemporal Area

Simplex TB
The dorsal surface of the superior temporal gyrus

(T1) that lies within the lateral (Sylvian) fissure,
i.e. the so-called planum temporale, usually comprises one to two, and often three transverse gyri,
the most anterior of which generally goes by the
name of first gyrus of Heschl (HI). The second is
often designated as the second gyrus of Heschl
(HII). Further back, one observes additional small
transverse gyri or a small plane surface.
The cortex of this entire dorsal surface of T1
can be cytoarchitectonically divided into three
areas, namely, TB , TC and TD (fig. 1a). The magnocellular supratemporal area simplex TB (area
EK 82) occupies the major portion of that surface,
while area TC, still partially inside area TB , occu-
Temporal Lobe
pies the surface area of a coin5 22 mm in diameter

(about 4 cm2), mostly on HI and partly on HII. In
contrast, area TD is entirely hidden in the profundity of the lateral (Sylvian) fissure.
Anteriorly and posteriorly, area TA also reaches as a thin promontory onto the dorsal surface of
T1. Area TB , which ventrally and laterally progressively becomes area TA, shows a structural
type analogous to TA, with the exception that the
cellular columns, disposed as an organ pipe formation (plate P93 in the Atlas [Economo and
Koskinas, 2008]), span even more distinctively
over all the cortical layers (denoted by the asterisk in fig. 43). Additionally, area TB is characterized by the presence of large cells in sublayer IIIc,
which have the appearance of giant cells amidst
the medium-sized cells in their vicinity (fig. 43).
The total thickness of the cortex here is about
3.0 mm, which is rather great, considering that
the planum temporale inside the lateral (Sylvian)
fissure is actually the dorsal wall of T1, and wall
formations are usually rather thin. But it is worth
noting that the planum temporale does not distinctly display all the characteristic features of
wall structures.
Layer I is very thick (table 10).
Layer II has a typical temporal appearance,
being irregular and mottled. The radial striation
that spans over the entire cortex encompasses
layer II; cells appear to advance into layer I at
many places; this is better seen in sections thinner than that depicted in figure 43. Layer II is not
sharply demarcated from layer III.
Layer III consists of vertical cellular columns,
traceable from layer IV (even layer VI) all the way
to layer II. With a cell size criterion, this layer can
be subdivided into sublayers IIIa, IIIb and IIIc. In
sublayer IIIc, i.e. at the lowest points of the organ
pipes, there are also very large pyramidal cells
(sometimes mistakenly called giant cells) distrib-
10 Groschen of the First Austrian Republic.
119
Fig. 43. Magnocellular temporal area simplex TB (area EK 82). Dome of the third (transverse supratemporal) gyrus of
Heschl (HIII). !45.
120
uted among the large multi-layered, superimposed pyramidal cells of the same sublayer.
Layer IV is both unusually thick, for belonging to the temporal lobe, and relatively rich in
cells, comprising large granule cells and numerous triangular cells; not infrequently, larger pyramidal cells proceed from sublayer IIIc into layer IV. Over the entire expansion of layer IV, these
cells are arranged in dense vertical columns (separated by acellular intervals) that form the continuation of the layer III columns.
Layer V is relatively thin, but of a rather compatible character with the dorsal wall of T1. The
cells are small, and only here and there isolated
larger pyramidal cells appear. In layer V as well,
cell-poor radial stripes are interposed among the
columnar cell groupings; being in continuation
with those mentioned above, they impart the appearance of an organ pipe formation.
Layer VI is subdivided into a superficial sublayer VIa and a deep sublayer VIb. Sublayer VIa
has almost double the number of spindle (fusiform) cells than VIb; cells of sublayer VIb are
smaller in size. The spindle cells are arranged in
columns. Demarcation from the white matter is
sharp.
Within area TB , Economo and Horn [1930]
define posteromedial, posterolateral, anteromedial and anterolateral fields; moreover, they observe transitional structures with the neighboring areas TA and TC, denoted as TBA1, TBA2 and
TBC (table 2; cf. also fig. 64b and 68 in the concluding chapter).
Function
The physiologic significance of area TB remains
unknown. In its inner magnocellular territories
it undoubtedly contains the parasensory zone of
the primary auditory domain TC and as such, it is
probable that it may functionally subserve, at
least in part, the adaptative reflex movements in
response to a stimulus or reflex attention.
Temporal Lobe
Supratemporal Area Granulosa TC
The supratemporal area granulosa TC (area EK

83) borders directly on area TB . A comparison of
figure 43 with figure 44 reveals one of the clearest
structural differences between these two neighboring areas. Area TC is heterotypic granulous
isocortex, i.e. a sensory cortex, which, as already
said, occupies a surface area of 4 cm2 on the middle of HI, and partially HII (fig. 1a); it is surrounded on all sides by area TB , and only towards the
floor of the lateral (Sylvian) fissure is it continued
by an analogous structure, albeit a less typical
granulous form, the intercalated supratemporal
area TD (area EK 84).
Area TC is characterized by a reduced size of
the majority of its cells, i.e. a general granulization; by a very striking cellular density and thickness of the true granular layers; and by the radial
disposition of the granule cells in absolutely fine
and slender strands, which we termed rain shower formation (fig. 44; cf. also plate P94 in the Atlas
The cortex is about 3.0 mm thick a figure
rather high for koniocortex. At the same time, its
structure has other, sufficiently characteristic
particularities, e.g. a light (cell-poor) layer V.
Layer I is exceptionally thick and relatively
rich in cells (table 10).
Layer II is exceptionally striking with many
small granule cells and with the smallest pyramidal cells. Cells are often compiled in thick groups
that extend to layer III like cloudy stretches. The
border with layer III is not sharp at all.
Layer III is thinner than in area TB , and is unusually rich in cells for a layer III; they are mainly small pyramidal and granule cells. The borders
with layer II, and with layer IV as well, are totally
obliterated, as all of these layers contain cells of
quite the same size and shape, and often cells of
one layer pass into the neighboring layer; nonetheless, layer III is somewhat less cell-dense than
the rest. On occasion, examples of larger pyramidal cells are found sporadically, but regularly, in
121
Fig. 44. Supratemporal area granulosa TC (area EK 83). Dome of the middle segment of the first gyrus of Heschl (HI).
!45.
122
deeper loci of layer III; these give the impression

of being very large relative to the dwarf cells in
their vicinity.
Layer IV can reach a similar, although not as
great, thickness as in striate area OC. It is considerably dense with granule and very small pyramidal cells.
Layer V has likewise the appearance of a granular layer; but it is considerably poorer in cells,
and it characteristically appears as a thick lighter
band in the stained tissue sections (fig. 44). Since,
however, this layer is made of such small cells,
one gets the impression that it has only a single
thick granular layer, with the cells vertically arranged, as if they were tied from bottom to top in
pearl beads, stretching all the way to layer I.
Layer VI consists of a sublayer VIa with spindle (fusiform) and small pyramidal cells, giving
the appearance of a somewhat darker band, and
a thinner sublayer VIb, containing cells loosely.
The border with the white matter is sharp.
Economo and Horn [1930] further detail a
pars granulosa (area TC1) and a pars simplex (TC2)
within the transverse supratemporal area TC (table 2; cf. also fig. 68).
Function
Area TC thus belongs to the granulous heterotypic
isocortex or koniocortex, i.e. a sensory cortex,
which represents precisely the primary auditory
domain into which the acoustic fibers that emanate from the medial geniculate body terminate
(the acoustic radiation of Pfeifer [1920]). It is
worth noting that area TC does not form a completely closed territory of its own, as e.g. does
the striate area; on the contrary, frequent bands
and islets from the surrounding area TB reach
over the boundary, and one often encounters
here all the intermediate transition forms between the granular homotypic isocortex of area
TB and the granulous heterotypic isocortex of area
TC (fig. 1a).
Temporal Lobe
Furthermore, it is remarkable that this sensory

area TC occupies such a small space on the gyrus
of Heschl it covers a total surface of 8 cm2 in
both hemispheres. And yet, it is possible that area
TD, which I cover next, also belongs to the primary auditory domain, despite its lack of the granulization so typical of area TC. Such an annexation, however, would still not increase substantially the surface area of this sensory domain.
In this regard, it is worth noting that the method of Flechsig [1898, 1903, 1920] for tracing the
development of myelinated fibers has led to the
isolation of a primary auditory sensory domain
on HI, an equally small field practically overlapping with our cytoarchitectonic area TC. The perception (Auffassung ) of higher-frequency tones is
mostly effected at portions of this field located in
the profundity of the lateral (Sylvian) fissure, the
perception of lower-frequency tones more towards
its external aspect. A detailed discussion of this
question can be found in Economo and Koskinas
[1925, pp. 701710].
Intercalated Supratemporal Area TD
The intercalated supratemporal area TD (area EK

84) is found completely within the depths of the
lateral (Sylvian) fissure, i.e. medially to the supratemporal area granulosa TC (area EK 83) and the
magnocellular temporal area simplex TB (area EK
82) (fig. 1a, d).
The cortex has an average thickness of 2.8 mm
and shows no radial striation, such as seen in areas TC and TB ; its cells are disposed in a remarkably disorderly manner as I already described
for other opercular formations to the point that
a horizontal lamination is no longer apparent in
this mass. On the whole, this field is poor, with
medium-sized cells only making a sporadic appearance in it. One might consider this either as
a less well differentiated granulous cortex and
very poor in cells, or as a cortical territory with
an arrested development. I do not show a figure
123
of this rather small field (it can be found in plate

P95 of the Atlas [Economo and Koskinas, 2008]),
but only note the following characteristics:
Layer I is of an average thickness (table 10).
Layer II is rich in cells, mostly small granule
and very small pyramidal cells.
Layer III is rather poor in cells, these being
small; here larger cells are only seen isolated, with
a certain granulization, but without the typical
concomitant increase in number. The orientation
of the cells is greatly variable, as is the density of
their distribution. Cellular columns are not observed, and in all, the cell picture is rather disorderly.
Layer IV is quite thick. Although its cells are
rather numerous, they are irregularly arranged,
with acellular patches at places. For the most
part, they are granule cells.
Layer V is richer in cells than in area TC, but
these hardly exceed the granule cell size.
Layer VI shows a sublayer VIa with pyramidal
and spindle (fusiform) cells and a sublayer VIb
with only half as many cells, of about the same
size. Layer VI is clearly demarcated from the
white matter.
Area TD frequently extends forward from the
floor of the lateral (Sylvian) fissure to the robust
koniocortical area TC, and frontally, from area TC
to the bordering postcentral insular area at the
temporal entrance IB T (area EK 52). Similarly to
area TC, Economo and Horn [1930] further detail
a pars granulosa (area TD1) and a pars simplex
(TD2) within the intercalated supratemporal area
TD (table 2; cf. also fig. 68).
Function
Thus, we have, as mentioned, certainly in area
TC, and probably in area TD as well, the koniocortex of the primary auditory domain, into which
terminate the acoustic radiations issued from the
medial geniculate body. At the same time, we recognize that area TB , with its large pyramidal
124
cells, corresponds to the magnocellular field that

we regularly find in the immediate vicinity of any
koniocortex (cf. the parasensory zones of areas
PC and OB). Sound perception is localized in the
deepest part of the lateral (Sylvian) fissure; it is
possible that this region corresponds to our area
TD.
Temporal Area Proper TE
The cortical formation that covers T2 and T3 displays the structural characteristics of the temporal lobe in all aspects (fig. 1ac). At an extreme
thickness of 3.5 mm, the cortex of this area ranks
among the thickest regions of the entire cerebral
cortex (fig. 6).
Figure 45 gives a good idea of the particular
cortical cytoarchitecture of the temporal lobe.
Layer II appears frayed. Layer III is thin and contains large cells, although it is relatively poor in
cell numbers. Layer IV is segmented into the typical vertical cellular columns. Layers V and VI
are enormously (colossally) thick and disclose a
development that recalls the structure of the
frontal lobe.
In spite of their purely temporal character,
the cytoarchitectonic features of T2 and T3 (i.e.
size of cells, thickness of the cortex and deep layers, and overall cortical growth) recall the frontal
lobe, to a point that area TE must be viewed as
belonging to the cortical structural type 2 (fig. 9).
On the contrary, the cortical build of T1 and T4,
which resembles the parietal structures, with medium-sized cells and a modest growth of layers V
and VI, must be reckoned as belonging to the cortical structural type 3.
In the temporal area proper TE , and throughout its cortical thickness, cells are disposed in
vertical columns as well; such a characteristic,
along with the robustness of the deep layers V
and VI regarding both thickness and cell density,
differentiates area TE from pure frontal formations (fig. 45).
Fig. 45. Temporal area proper TE . Dome of the middle temporal gyrus (T2), showing its middle modification TE1 (area
EK 85). !45.
Temporal Lobe
125
Layer I is divided into an upper sublayer rich

in cells, and a lower sublayer poor in cells (table 10), which are mostly triangular and small
pyramidal.
Layer II is relatively thin and uneven with regard to both distribution and density of cells. It
has a frayed appearance; often, dense clusters of
cells swarm towards layer I (beneath the arrows
in fig. 45). The border vis--vis layer III is diffuse.
Layer III occupies less than one-third of the
total cortical height, which is relatively low for
this layer. It shows a rather disorderly repartition
of cells, which appear poorly formed and are
grouped into irregular columns; certain loci seem
to be quite poor in cells. One can distinguish
three separate sublayers in layer III, namely, IIIa,
IIIb and IIIc. Sublayer IIIc contains isolated large
cells. It is typical for area TE to often have a layer
III that is both relatively and absolutely thicker at
the gyral wall than at the dome; this is the inverse
relationship from what is usually seen in other
gyri.
Layer IV is segmented, as is typical for the
temporal cortex, in vertical columns, with oligocellular or acellular intervals among them (denoted by the plus sign in fig. 45). Above and below, the cells of layer IV, which are partly granule
and partly pyramidal, enter into the domains of
layers III and V, blending with adjacent rows of
pyramidal cells.
Layer V is rich in large cells, and contains two
cellular populations based on size. They are all
disposed in fine, radial stretches or columns. Pyramidal cells in layer V of area TE in T2 are finely formed, remarkably so if one compares them
with the irregular and small pyramidal cells in
layer V of the adjacent basal parietal area PH.
Such a characteristic constitutes an excellent criterion for studying the transition between the occipitotemporal intermediate zone (with a parietal
structure) to a true temporal region.
Layer VI is typically thick and very rich in
large cells distinctively disposed in radial col-
126
umns, i.e. in full accord with area TE . The upper

sublayer VIa contains robust spindle (fusiform)
cells, while the deeper sublayer VIb contains half
their number. The border of layer VI vis--vis the
white matter is uncertain; the transition is gradual, a particularity we already saw in cortical
structural type 2 in the frontal lobe, e.g. in the
intermediate frontal area FC.
Middle (TE1) and Inferior (TE2) Modifications

of Temporal Area Proper
The prevalence of layers V and VI in the cytoarchitectonic picture as we already know from
area TA (fig. 42) over the relatively thin and
poorer layer III, and the weak growth of layers II
and IV are striking characteristics of proper temporal formations. At the same time, T2 contains
larger cells in layer III than T3, and thus one may
divide this area into an upper half, the middle
temporal area proper TE1 (area EK 85) located on
T2 (plates P89 and P90 in the Atlas [Economo and
Koskinas, 2008]) and a lower half, the inferior
temporal area proper TE2 (area EK 86) covering
T3 (plate P91 in the Atlas [Economo and Koskinas, 2008]).
This formation in the middle segments of T2
and T3, which has some characteristics of the cortical structural type 2, continues caudally along
T2 and T3 as far as into the angular gyrus and the
basal parietal aspect, such that, right in the district of cortical structural type 3, we find a blending of these two cortical structural types in the
lower part of the angular gyrus and its basal vicinity.
The boundary between the temporal structural type 2 and the occipitotemporal intermediate
zone of the parietal lobe (structural type 3) roughly corresponds to an oblique line descending
from the posterior extreme of the lateral (Sylvian)
fissure backwards (fig. 1a). The boundary of area
TE with area TA lies at the depths of the superior
temporal sulcus, and usually on its dorsal wall.
Rostrally, area TE is gradually transformed

through a reduction of its granular layers to the
temporopolar area TG. At the depths of the internal temporal sulcus, area TE borders on the successive area TF.
Function
Area TE is the site of origin of the temporopontine fiber tracts, possibly in rapport with the
marked development of efferent layers V and VI.
Neuropathology teaches that lesions of this area
occasionally produce static ataxia and certain
disturbances of ocular movements; with lesions of
more caudal parts, in the vicinity of the occipitotemporal intermediate zone, amnesic aphasia has
been observed.
Fusiform Area TF
As mentioned earlier, the fusiform gyrus (T4) is

clad in a cortical formation having some of the
structural characteristics of the temporal lobe on
the one hand, but resembling the cortex of T1 on
the other, as it recalls the parietal structural type
3 (fig. 46), which it also borders caudally (fig. 9).
This is actually the basal parietal area PH which,
as described above, carries the parietal type of
cortex all the way to the ventral surface of the cerebral hemispheres (fig. 1b, d).
The fusiform area TF (area EK 87) still has a
thick cortex, albeit somewhat thinner than the
rest of the temporal gyri; it measures 2.53.0 mm
on average (fig. 6, 46). Both granular layers are
somewhat more distinct than in area TE of T2
and T3, a fact that accentuates their horizontal
lamination. Vertical striation is more delicate, resembling that of T1 and the posteriorly adjacent
parietal areas. However, what distinguishes area
TF from area PH is mainly the presence of a relatively well developed layer V with fine pyramidal
cells, in contrast to the rudimentary cells of PH.
Temporal Lobe
The clear demarcation of layer VI from the white

matter is another characteristic feature of area TF
(cf. also plates P96, P108 and P112 in the Atlas
Layer I is slightly thicker than average, with
fairly large cells (table 10).
Layer II is not very dense in cells, but still
more compact than layer II in T2 and T3. It mostly contains granule cells, and also some pyramidal cells. Like most parts of the temporal lobe,
this layer in area TF also shows local irregularities in density, often appearing fringy at the layer
I border.
Layer III is relatively thick and richer in cells,
which are predominantly medium-sized pyramidal. There are no marked sublayer divisions (parietal structural type). More voluminous cells are
only seldom observed. The medium-sized cells
that prevail are arranged in narrow perpendicular columns, which, though, are not very conspicuous.
Layer IV is quite compacted with large granule cells, arranged in columns, with alternating
multicellular and oligocellular zones (temporal
structural type). The upper and lower borders of
layer IV are imprecise.
Layer V is subdivided into a thin superficial
sublayer Va, immediately subjacent to layer IV
and composed of densely packed, small, triangular cells, and a deeper sublayer Vb with large cells,
less densely arranged than in Va. Sublayer Vb
thus stains more lightly compared to Va and helps
differentiate area TF from its surrounding temporal areas. Moreover, the robust size of cells in
layer V helps distinguish area TF from the parietal structure of the intermediate zone PH situated at its immediate caudal boundary in T4.
Sometimes, cells of sublayer Va are also less compactly arranged, rendering it paler, and making
layer V present with a darker intermediate band
between its upper and lower lighter rows, which
reminds of a similar structure in the uncus, as we
shall see later.
127
Fig. 46. Fusiform area TF (area EK 87). Dome of fusiform gyrus (T4). !45.
128
Layer VI is relatively thin and comprises an

upper, more compact sublayer VIa with cells of a
good size, partly triangular and partly spindle
(fusiform), and a deeper sublayer VIb, which is
thinner than elsewhere, with true spindle cells.
Demarcation from the white matter is considerably sharper than in other temporal areas.
Function
Area TF extends over the entire fusiform gyrus
and its inner wall through the collateral sulcus.
With lesions of the fusiform gyrus, disturbances
of visual orientation are observed.
Hippocampotemporal Area TH
The inferior wall of the fusiform gyrus in the collateral sulcus and the opposite wall of this sulcus,
which forms the inferior wall of the hippocampal
gyrus, are occupied by the hippocampotemporal
(or temporohippocampal) area TH (area EK 88),
which bears structural similarities to area TF.
Area TH is a thin rostrocaudal band that covers
the middle third of the collateral sulcus (fig. 1b,
d) and bounds the six-layered (hexalaminar) homotypic isocortex of the temporal lobe from the
allocortex of the hippocampal gyrus, the latter
forming part of the rhinencephalon.
In the same way, area TH occasionally reaches
somewhat over the inferior brink of the hippocampal gyrus, finding its way on to the dome
within a few mm. In this part, area TH shows an
analogy to both area TF and to the polar area TG,
which lies directly in front of TH, as explained
further below. I do not include a specific figure
for this small area, but its position and structure
can be discerned with precision in figure 52 and
53.
The cortex is 2.7 mm thick at the wall and displays a most distinct horizontal lamination,
which differentiates it from the temporal struc-
Temporal Lobe
tures discussed so far; at the same time, the vertical striation becomes indistinct or lost.
Layers II and III are poorly developed, containing few cells, while layers V and VI are particularly rich in cells.
Layer V further discloses a very distinct horizontal cell band, consisting of pyramidal cells
compactly arranged in multiple rows, a structure
that we typically find in all regions in the vicinity
of the rhinencephalon, i.e. in the insula and the
frontal areas neighboring the limbic gyrus, such
as FC L, FD L, FE L, FHL , FG, FJ, IA and IB . Layer
V often also contains a small lightly staining horizontal band, situated under the above-mentioned dark band.
Layer VI is dense in cells, and even more
sharply demarcated from the white matter than
in area TF or the other temporal fields; it mostly
contains triangular spindle cells.
Agranular Hippocampotemporal Area TH

Near the boundary of area TH with area TF, layers II and IV still display temporal characteristics, i.e. the fringed border towards layer I and the
columnar disposition of layer IV. But towards the
hippocampal gyrus, they become less compact,
and almost agranular at the boundary with the
hippocampal rhinencephalic areas. Such an inner border can be designated as the agranular
hippocampotemporal area TH (area EK 89)
(fig. 1b, d and 52; cf. also plates P108, P111 and
P112 in the Atlas [Economo and Koskinas,
2008]).
Temporopolar Area TG
All the areas of the temporal lobe described so far

course rostrocaudally along the longitudinal axis
of the various temporal gyri, and past them, to
within reach of the temporal pole. However, the
temporal pole is wreathed in a particular struc-
129
ture, the temporopolar area TG (area EK 90); it

further includes the anterior extreme of the dorsal (Sylvian) surface of T1 with its rostral secondary (transverse) gyri of Schwalbe [1881] (fig. 1a, b,
d), and the most rostral segments of T1T4 (plates
P97, P107 and P110 in the Atlas [Economo and
Koskinas, 2008]).
The transition of area TG to the other areas of
the various temporal gyri is gradual (cf. table 2
and fig. 68 in the concluding chapter for the specific transition modifications with area TA according to Economo and Horn [1930]), and consequently indistinct. In the rhinal sulcus, i.e. in
the rostral segment of the collateral sulcus, there
is a sharp distinction of the boundary of area TG
with the allocortex of the uncus. Economo and
Horn [1930] further subdivide the temporopolar
area TG into a parainsular (TG1) and a marginal
(TG2) part, the latter comprising a lateral (TG2)
and a medial (TG2) field (cf. table 2 and fig. 68 in
the concluding chapter).
The temporopolar cortex ranks among the
thickest of all cerebral cortical territories (fig. 6);
it varies from 3.0 to 4.0 mm. Its structure is mostly magnocellular (fig. 47), and pyramidal cells assume a somewhat special, guttiform shape, owing to the small number of dendrites. The cortex
is exiguously granular, at places actually becoming agranular. Vertical striation is not seen in this
area.
Layer I is thick and contains large cells of a
triangular or spindle shape (table 10).
Layer II, in contrast, is thin and is formed
only of one to two rows of pyramidal cells. It often shows discontinuities and acellular patches,
irregularly alternating with hoards of small cells;
still, due to its intense staining, layer II is readily
discerned as a dark, frequently interrupted, thin
band under layer I.
Layer III goes to the other extreme, in being
exceptionally thick. It is not very rich in cells; the
cells are hardly more than medium-sized, and
have a pointed ovoid or guttiform shape. Acellular
patches are frequent in layer III.
130
Layer IV is also exceptionally thin, and often

interrupted or fragmented as well. It contains
mostly small, pointed ovoid pyramidal cells.
There are no cellular columns observed in either
layer III or layer IV. At certain loci, especially in
the dorsal zone of area TG, layer IV presents as a
lightly-staining band that brings to mind the
acellular lamina dissecans of the neighboring allocortex of the hippocampal gyrus and the uncus; this pale band can be seen in figure 47 (cf.
also fig. 56, as well as the section on the uncinate
area HA in the hippocampal lobe chapter). (These
comments also apply to the cytoarchitectonic
picture of the hippocampotemporal area TH and
its agranular modification TH.)
Layer V has a variable thickness. It consists of
pyramidal cells, usually robust. The deeper part
of layer V is somewhat richer in cells and more
intensely stained, as cells are more voluminous
than in the upper part. The deeper part of layer V
is divided from the subjacent sublayer VIa by a
clear short band. Thus, layer V appears to lie between two light bands, a situation that strongly
recalls an analogous picture of certain parts of
the parauncinate area HB in the uncus.
Layer VI comprises an intensely stained upper sublayer VIa, being the densest in the entire
area TG. In tissue sections from this area, the increasingly deeper staining from the cortical surface towards the depth is striking and characteristic. Directly beneath, the deeper sublayer VIb
appears much lighter. In both sublayers VIa and
VIb, triangular cells predominate over fusiform.
Agranular Temporopolar Area TG

The temporopolar area becomes devoid of its
granular layers at its confines with the uncus, as
well as its proximity with the substantia perforata; one may hence designate in its dorsal part a
separate modification, the agranular temporopolar area TG (area EK 91) (fig. 1a, b, d; cf. also
plates P58c, d, P98, P99 and P103c, d in the Atlas
Fig. 47. Temporopolar area TG (area EK 90). Dome of temporal pole. !45.
Temporal Lobe
131
[Economo and Koskinas, 2008]). Such a cortex

becomes progressively thinner toward the substantia perforata, and is separated from it by a
thin cortical margin formed by the posterior crus
of the lateral olfactory gyrus (area TJ, described
next). At the confines between the uncus and the
lateral olfactory gyrus, the cells of layer II increasingly tend to become stellate and to form
small groups.
Layer III diminishes rapidly, becomes poorer

in cells, and shows numerous acellular patches.
Layers V and VI are thus the only prolongations found directly beneath layer I, and these
also terminate abruptly. In the deep parts of the
tissue, the cells of layers V and VI of area TJ are
in loose continuation with the deep cell groups of
the substantia perforata, the semilunar gyrus of
the uncus (area TJ H) and the nucleus of the
amygdala (cf. hippocampal lobe chapter and
fig. 59a).
Temporal Piriform Area TJ
This thin cortical margin, which forms the posterior boundary of the substantia perforata (fig. 1d
and 21), is the continuation of a thin layer of allocortical gray matter that accompanies the lateral olfactory root in its entire trajectory from the
olfactory tract and the olfactory triangle, into
area FK along the frontal transverse insular gyrus, and then as area ID along the polar insular
gyrus, whence its path bends in an acute angle
along the small falciform insular gyrus posteriorly, passing medially as area TJ to the edge of the
temporal pole, to finally terminate at the semilunar gyrus of the uncus as area TJ H.
The temporal piriform area TJ (area EK 92),
which occupies this part of the cortical margin
(cf. plates P58c, d and P98 in the Atlas [Economo
and Koskinas, 2008]), is composed of an incomplete cortex, similarly to the frontal piriform area
FK (area EK 29) and the piriform insular area ID
(area EK 54), forming a continuation of them
(fig. 22).
Layer I is exceptionally thick and contains in
its upper part numerous myelinated fibers.
Layer II cells show a tendency to increase in
size and to form small clusters.
132
Area of Substantia Perforata TK
Beyond this cortical margin, only layer I passes

over to the substantia perforata. We have named
this lateral (external) portion of the substantia
perforata, which lies between the temporal pole
and the base of the frontal lobe, the area of substantia perforata TK (area EK 93), although it
does not truly belong to cerebral cortex proper
(fig. 1d, 21, 22; cf. also plate P58c, d in the Atlas
[Economo and Koskinas, 2008]). The medial (internal) portion of the substantia perforata, already described in the frontal lobe chapter, is represented in the precommissural area FN (area EK
35).
In the region of area TK one finds, beneath the
thick layer I, irregular glomerulous clumps of
small stellate and granule cells, separated by passing myelinated fiber bundles; still deeper, one
may even see larger glomerulous accumulations
of cells; as I already clarified in the frontal lobe
chapter (fig. 22), they are related to the head of
the caudate nucleus.

Cingulate Gyrus and Retrosplenial Region
Anatomy of the Cingulate Gyrus
The present chapter covers the cytoarchitectonics of the upper (superior) half of the so-called
limbic lobe, including the retrosplenial region as
far as the isthmus (fig. 1b). This anatomical structure has been alternatively termed by different
authors [Economo, 1927d], such as gyrus cinguli
(gyrus of the cingulum) by Burdach [18191826],
circonvolution annulaire (annular convolution)
by Gerdy [1836], circonvolution de lourlet (convolution of the cingulum) by Foville [1844], gyrus
fornicatus (fornicate gyrus) by Ecker [1869, 1873],
circonvolution limbique (limbic convolution) and
grand lobe limbique (great limbic lobe) by Broca
[1877, 1878], lobus falciformis (falciform lobe) by
Schwalbe [249], and convolution of the corpus callosum by Turner [1890]. Whatever its name may
be, the limbic or cingulate gyrus is generally associated with the rhinencephalon sensu latiori.
Across the median (mid-sagittal) plane of the
cerebral hemisphere (consult fig. 57 in the next
chapter), the limbic gyrus (g.l.) follows the curvature of the corpus callosum, assuming an arched
form, and also the posterior pillars (crura) of the
fornix from the splenium (Spl.C.c.), downward
and forward, all the way to the fusion of the fimbria with the uncus (U.). Therefore, in its extension from the rostrum of the corpus callosum
(R.C.c.) to the uncus, the limbic gyrus forms an

almost closed ring, with an opening only underneath and to the front, where the parolfactory
field of Broca (C.Br.), the olfactory gyri and the
anterior substantia perforata (structures already
discussed) are interposed.
Ventrally to the posterior extreme of the corpus callosum, i.e. the splenium (fig. 57), the common trunk (Tr.) of convergence of the calcarine
(C.) and parieto-occipital sulci (s.po.) reaches forward as far as the cingulate gyrus and constricts
it at this point, as it were, to form a virtual isthmus (fig. 57). In that manner, the limbic lobe can
be subdivided into two segments (fig. 57): the superior limbic lobe or limbic gyrus (g.l.) dorsally,
which encircles the corpus callosum; and the inferior limbic lobe or hippocampal gyrus (g.h.) ventrally, which accompanies the fimbria (fi.).
Limbic and Intralimbic Gyrus
In this arch, the cortex of the limbic lobe is successively bounded at its periphery by the cortical
areas of the frontal, superior parietal and occipital lobes already mentioned (fig. 1b), then by the
basal part of the parietal lobe, and finally by the
temporal lobe; it belongs partly to isocortex, and
partly to allocortex. As isocortex it coats the ex-
ternal (outer) wall and the dome of the cingulate

gyrus. At the internal (inner) wall, inward of the
callosal and hippocampal sulci (s.c.c. and s.h. in
fig. 57), it undergoes a swift thinning and a metamorphosis to allocortex, shifting, at the depths of
these sulci, to hardly more than an absolutely
thin lamella folded over the external aspect of the
corpus callosum (Balkenrcken in German, literally meaning saddle or dorsum of the corpus callosum) and the fimbria.
At the point where this small stretch becomes
folded on the two above-mentioned formations
(i.e. the dorsum of the corpus callosum and the
fimbria) and enlarged to form a small bulge, it assumes the name of intralimbic gyrus. Just like in
the cingulate gyrus, one may also distinguish in
this inner concentric gyrus a simple, largely
membranous dorsal part that overlies the dorsum of the corpus callosum, the indusium griseum (Latin for gray veil derived from the Greek
verb = to dress or to coat), and a ventral
part, the dentate gyrus, that accompanies the
fimbria.
We designate all the areas of the limbic and
intralimbic gyrus, from the rostrum of the corpus callosum to the isthmus, with the initial capital letter L; those of the hippocampal and dentate
(intralimbic) gyrus from the isthmus to the end
of the uncus with the initial capital letter H
(fig. 1b, d).
Figure 4855 reproduce microphotographs of
consecutive cross-sections through the cingulate
gyrus at !15, !14, !12 or !10 magnifications.
In particular, figures 4850 correspond to the
cingulate gyrus, figure 51 to the isthmus, figures
52 and 53 to the hippocampal gyrus, and figures
54 and 55 to the uncus, so that the expansion and
shift of areas might be followed directly.
For a better understanding of these figures, I
include an additional series of drawings. Figure
57 in the following chapter presents a schematic
drawing of the entire limbic lobe; the radially disposed dashed lines (marked with letters at their
distal end) leading to the arch of the cingulate
134
gyrus indicate consecutive sections, fully drawn

in figures 5961. Thus, based on figure 1b and
the cross-sectional drawings, one can easily reconstruct the form of expansion of the cortex
over the various successive areas in this intricately structured region. Such cross-sections through
the limbic gyrus are denoted by the Greek letters
, , and in figure 57, and schematically depicted by the series of drawings in figure 61.
I would like to call attention [Economo, 1929d]
to the fact that the entire rhinencephalon reveals
a complex structure and numerous areas; a full
description and photographic documentation
would exceed the limits of this book, the essential
aim of which is to offer an overall survey of the
cerebral cortex. Therefore, I herein discuss and
illustrate only the fundamental points. Anyone
particularly interested in the structure of the
rhinencephalon and the allocortex will find exhaustive material in the respective sections of our
larger treatise [Economo and Koskinas, 1925,
chap. 8 and 13].
Agranular Anterior Limbic Area LA
The cortex of the entire anterior tier of the cingulate gyrus, from the rostrum of the corpus callosum to the extrapolated continuation line of the
central sulcus on the median hemispheric facies,
is agranular; the cortex of the posterior tier, all
the way to the isthmus, is granular (fig. 7b).
The agranular area LA exhibits certain structural modifications along its vast extent. One distinguishes three longitudinal concentric bands
(LA1, 2, 3): the precingulate agranular anterior limbic area LA1 (area EK 36), covering the dorsal and
external wall of the limbic gyrus; the cingulate
agranular anterior limbic area LA2 (area EK 37),
covering the dome; and the agranular anterior
limbic area limitans LA3 (area EK 38), covering
the internal wall of the cingulate gyrus within the
callosal sulcus (fig. 1b, 61), towards the depths
of which it rapidly thins out, to the point of the
Fig. 48. A coronal section (inclined anterosuperiorly) through the anterior tier of the cingulate gyrus (corresponding
to plane in fig. 57 and schematically drawn in fig. 61). The median hemispheric plane (and the dome of the cingulate gyrus) is on the top side of the figure. The internal wall of the cingulate gyrus (upper bank of the callosal sulcus)
is on the left side of the figure, opposite to the dorsal aspect of the corpus callosum (extreme left). The external wall
of the cingulate gyrus (lower bank of the cingulate sulcus) is on the right side of the figure. Precingulate agranular
anterior limbic area LA1 (area EK 36). Anterior cingulate agranular anterior limbic area LA2 (area EK 37). Cingulate
agranular anterior limbic area limitans LA3 (area EK 38). Anterior ultracingulate area LB1 (area EK 39). Area of indusium
griseum LB2 (area EK 40). The limbic granular frontal area FD L (area EK 15), a transition zone, is seen on the lower right
corner, in the cingulate sulcus. C.c., corpus callosum; s.cc., sulcus of corpus callosum; Z, ridge of layer I delimiting the
intralimbic gyrus. !12.
135
Fig. 49. A coronal section (inclined posterosuperiorly) through the posterior tier of the cingulate gyrus (corresponding to plane in fig. 57 and schematically drawn in fig. 61). The median hemispheric plane (and the dome of the
cingulate gyrus) is on the top side of the figure. The internal wall of the cingulate gyrus (upper bank of the callosal
sulcus) is on the left side of the figure, opposite to the dorsal aspect of the corpus callosum (extreme left). The external wall of the cingulate gyrus (lower bank of the cingulate sulcus) is on the right side of the figure. Ventral posterior
cingulate area LC2 (area EK 42). Posterior cingulate area limitans LC3 (area EK 43). Anterior ultracingulate area LB1 (area
EK 39). Area of indusium griseum LB2 (area EK 40). Posterior ultracingulate area LF1 (area EK 47). Ultracingulate area
obtecta LF2 (area EK 48). C.c., corpus callosum. !12.
136
cortex finally becoming absolutely thin and consisting only of layer I and a few subjacent cells.
This cortex continues by covering the external
surface of the corpus callosum as area LB (vide
infra). Figure 48 reproduces a microphotograph
which corresponds to the schematic drawing of
figure 61. It gives a good overview of all the topographic relations; the corpus callosum is on the
left side of the figure, and the cingulate gyrus on
the right.
The cortex of area LA1 has an average thickness of 2.5 mm (table 11).
Layer I is rather thick; sublayer Ib contains
three times as many cells as sublayer Ia. At the
dome of the cingulate gyrus in area LA2, where
the cortex reaches a total thickness of 2.9 mm,
layer I becomes even thicker and richer in cells.
This layer then becomes thinner as it passes on to
the internal wall of the cingulate gyrus, and as far
as the floor of the callosal sulcus, where it sends
a ridge (denoted by the letter Z in fig. 48) into the
deeper parts of the cortex, similar to the arrangement mentioned in the frontal lobe chapter when
speaking of the transition from the parolfactory
area FL to the geniculate area FM.
Layer II does not truly exist in the cortex of
the three areas in the anterior tier of the cingulate
gyrus; in its place one finds triangular and small
pyramidal cells of the upper zone of layer III. In
figure 48, one certainly sees (to the right) a layer
II with small granule cells in the external wall of
the cingulate gyrus near the floor of the cingulate
sulcus, but such a layer actually reaches the wall
upward only as far as the small vessel that penetrates the cortex from the outside; further toward
the dome, one does not really find traces of a definite layer II. The part of the wall below the blood
vessel belongs to the cortical transition zone of
the adjacent area FD L, as one recognizes from the
existence of an internal granular layer IV. Area
LA1 only commences towards the dome.
Layer III is subdivided into a more superficial,
parvicellular sublayer IIIa, and a deeper sublayer
IIIb. The pyramidal cells are only of small and
medium calibre, and their juxtaposition is quite

irregular; in all, the entire layer III is little developed. At the internal wall of the cingulate gyrus
(area LA3), layer III becomes extremely thin and
poor in cells, often showing acellular patches. It
appears to end abruptly at the aforesaid ridge (Z
in fig. 48) of layer I.
Layer IV is also missing from the entire extent
of areas LA1, 2, 3 as already mentioned; it appears
again only in the cingulate sulcus at anterior segments of the frontal lobe, where the granular areas FC, FD , FE and FH border on this region directly, with their variants FC L, FD L, FE L and
FH L.
In contrast to the inadequate growth of the superficial layers, the robust presence of layers V
and VI is typical for the anterior tier of the cingulate gyrus (table 11).
Layer V is thick in the external wall of the cingulate gyrus (areas LA1 and LA2); it only diminishes in thickness in the internal wall (area LA3).
It is subdivided into a thinner and more compact,
parvicellular sublayer Va, where cells are disposed in bands or stripes, most distinct at the external wall (area LA1), and still very evident at the
dome (area LA2). They begin to disappear at the
internal wall of the cingulate gyrus (area LA3).
This darker stripe is also visible in figure 48. The
deeper sublayer Vb is much thicker and lightly
staining; its cells are larger but less densely
packed.
Rod and Corkscrew Cells. At the dome in area
LA2, and especially at the internal angle of the
dome and the transition to area LA3, a portion of
the cells of sublayer Vb are conspicuously elongated, many being corkscrew-like and stretched
like spindles. These here are the peculiar rod and
corkscrew cells we have already recognized in the
frontoinsular area FJ (area EK 28, cf. Frontal
Lobe chapter); area FJ and this region of layer V
of the anterior cingulate gyrus are the only two
regions in which these cells are found. They were
already known as spindle cells in layer V of the
cingulate gyrus to Hammarberg [1895], Flechsig
137
138
[1897, 1920], Ramn y Cajal [19001906, 1921,

1923], Nikitin [1909], Marinesco [1910a, c] and
other authors.
They are best developed at the internal brink
of the cingulate gyrus, i.e. in the transition from
area LA2 to LA3; despite the low magnification,
they can even be seen in figure 48. These special
cells of the anterior cingulate gyrus (and the
transverse insular gyrus) can reach a height of
80 m, yet their width is only 710 m (table 11).
I showed that these cells derive from neuronal
precursors [Economo, 1926d].
In addition to these rod cells in sublayer Vb of
area LA3, one also finds very slender, lancetshaped pyramidal cells, especially toward the
floor of the callosal sulcus.
Layer VI is also relatively thick. One can distinguish an external sublayer VIa packed with
spindle cells, which in the internal wall of area
LA3 becomes substantially thinner and only contains small triangular and spindle (fusiform)
cells, and a sublayer VIb poorer in cells, yet rather thick, which also becomes substantially thinner in area LA3.
At certain places of the dome, layer VI and layer V together make up about two-thirds of the
total cortical thickness. This predominance of
the lower over the upper layers is exceptionally
Fig. 50. A horizontal section (inclined posterosuperiorly) through the retrosplenial region (corresponding to
plane in fig. 57 and schematically drawn in fig. 61).
The median hemispheric plane (and the dome of the cingulate gyrus) is on the top side of the figure. The internal
wall of the cingulate gyrus (upper bank of the callosal
sulcus) is on the center-left side of the figure, opposite to
the dorsal aspect of the corpus callosum (left). Posterior
cingulate area LC (areas EK 4143). Agranular retrosplenial area LD (area EK 44). Superior LE1 and inferior LE2
retrosplenial area granulosa (areas EK 45 and 46). Posterior ultracingulate area LF1 (area EK 47). Ultracingulate
area obtecta LF2 (area EK 48). Pyramidal (hippocampal)
area HE (areas EK 103106 and Economo 108 and 109).
Fascia dentata HF (area EK 107). Area of indusium griseum LB2 (area EK 40). !15.
typical of the cingulate gyrus. I shall talk immediately about the further relations of layers V and
VI at the cortical margin. The border with the
white matter is rather sharp.
From this description of the structural form of
areas LA1, 2, 3 one sees that all three modifications
are heterotypic agranular isocortex (cf. also plates
P44P47 and P52c, d in the Atlas [Economo and
Koskinas, 2008]).
Anterior Ultracingulate Area LB1 and Area of

Indusium Griseum LB2
At the depths of the callosal sulcus, the cortex

passes on to the dorsum of the corpus callosum
(fig. 61); in the vicinity of this passage point, the
molecular layer shows a deep protrusion, in the
form of a more or less well developed conical
ridge (Z in fig. 48), as described earlier, delimiting the cortex from the cortical margin (Rindensaum). On the other hand, the molecular layer,
however thin, passes over to the dorsum of the
corpus callosum, investing its entire width, all
the way to the opposite hemisphere; a few cells of
the upper sublayers and many cells of sublayer
Vb and layer VI push their way under the ridge
of layer I to the other side of it and form small,
loosely arranged cellular groups in the floor of
the callosal sulcus at the transition point to the
corpus callosum, even passing in part on to the
dorsum of the corpus callosum. Thus, embedded
in the angle between the corpus callosum and
the cingulate gyrus, and extended rostrocaudally, these cellular groups form a distinctive bandlike area at the depths of the callosal sulcus,
which we term anterior ultracingulate area LB1
(area EK 39).
The rest of the corpus callosum is clad only in
a thin molecular layer (immediately to the right
of the mark LB2 in fig. 48), practically without
nerve cells, with the exception of a band of small,
ovoid neurons that accompany the medial longitudinal stria of Lancisi [1712]. This part of the
139
Table 11. Summative table of quantitative data in eight modification areas of the superior limbic lobe. Separate
dome and wall thickness data and some additional values supplemented from tables I, III, V and VI in Economo and
Koskinas [1925, pp. 794801].
Area
symbol
Area name
Cortical
layer
LA1
precingulate agranular
anterior limbic area
Ia
Ib
IIIa
IIIb
Va
Vb

mm
wall
structure
only

mm
0.27
0.60
0.65
VIa
VIb
LA2
LC1
LC2
LC3
140
anterior cingulate
agranular anterior
limbic area
dorsal posterior
cingulate area
ventral posterior
cingulate area
posterior cingulate
area limitans
I
IIIa
IIIb
Va
Vb
0.45
0.35
0.27
0.82
0.80
dome
structure
only
VIa
VIb
0.57
0.37
I
II
IIIa
IIIb
IV
Va
Vb
VIa
VIb
0.24
0.26
0.24
0.23
0.66
0.65
0.33
0.24
0.48
0.41
0.40
0.35
0.37
0.15
0.25
0.00
0.25
0.18
I
II
IIIa
IIIb
IV
Va
Vb
VIa
VIb
I
IIIa
IIIb
Va
Vb
VIa
VIb
1.00
0.52
0.22
0.24
0.60
0.44
0.60
0.30
0.45
0.35
0.27
wall
structure
only
0.85
0.50
0.40
0.20
Cell content
cells/mm3
5,000
15,000
35,000
25,000
30,000
20,000
35,000
20,000
25,000
33,000
22,000
20,000
16,000
Cell size, m*
H(minmax)/
W(minmax)
55/510
15/710
20/1015
2530/1520
2530/1520
40/10
1520/710
1015/57
25,000
15,000
510/57
1520/1015
2025/15
30/15
40/1015
6080/710
2040/1015
15/10
6,000
130,000
30,000
25,000
125,000
65,000
30,000
35,000
15,000
7/45
515/310
1520/1015
2030/1520
510/510
2025/1520
2530/1215
1520/710
15/7
5,000
70,000
27,000
25,000
120,000
40,000
30,000
40,000
18,000
3/7
1015/510
18/18
2530/22
46/46
2030/1520
3040/1520
2030/1215
15/7
14,000
30,000
21,000
21,000
12,000
22,000
14,000
Table 11 (continued)
Area
symbol
Area name
Cortical
layer
LD
agranular retrosplenial
area
I
IIIa
IIIb
Va
Vb
VIa
VIb
LE1
LE2
superior retrosplenial
area granulosa
inferior retrosplenial
area granulosa
I
III
IV
Va
Vb

mm

mm
0.30
mostly wall
structure
0.80
0.60
0.30
0.20
0.31
0.41
0.35
mostly wall
structure
0.66
Cell content
cells/mm3
3,500
60,000
36,000
50,000
16,000
35,000
15,000
6/8
15/710
2030/1525
1520/1015
2040/1520
20/15
12/8
6,000
65,000
160,000
55,000
25,000
56/610
15/710
6/6
15/15
10/7
2040/1015
15/10
12/8
VIa
VIb
0.39
0.20
33,000
12,000
I
IV
V
0.36
0.34
0.70
6,000
160,000
25,000
0.40
0.20
27,000
10,000
mostly wall
structure
VIa
VIb
Cell size, m*
H(minmax)/
W(minmax)
56/610
6/6
15/15
3070/1015
15/10
12/8
indusium griseum, reposed entirely on the corpus callosum, is termed area of indusium griseum
LB2 (area EK 40).
The two modifications LB1 and LB2 constitute
areas of the so-called intralimbic gyrus; they belong to allocortex, as their cells do not disclose the
typical layering (cf. also plates P47, P50, P51 and
P52c, d in the Atlas [Economo and Koskinas,
2008]).
Transition Areas FC L, FDL, FE L and FH L

In the cellular structure discussed here, area LA
invests the anterior tier of the cingulate gyrus,
and gradually continues (fig. 1b) under the ros-
trum of the corpus callosum into the areas of the

parolfactory field of Broca (parolfactory prefrontal area FHL and parolfactory area FL). Dorsally in the cingulate sulcus, area LA successively passes over to the adjacent transition areas
FH L, FE L, FD L and FC L of the frontal lobe
(fig. 1b; cf. also plates P17, P26, P38, P44 and
P52d in the Atlas [Economo and Koskinas,
2008]), from which it can be distinguished first,
by the better development of layer III in the
frontal structures, and second, by the fact that
these adjacent areas possess granular layers (II
and IV), which are both lacking from the agranular area LA . Moreover, area LA extends into
these frontal transition areas the band-like
structure of sublayer Va, which is so character-
141
istic of all formations in the vicinity of the rhinencephalon and of area LA , as already discussed
in the frontal lobe chapter.
More caudally to area FC L, at the point where
the posterior frontolimbic transition gyrus leaps
dorsally over the callosomarginal sulcus, the
agranular area LA borders directly on the similarly agranular frontal area FB , such that the
agranular cortices of the frontal lobe and the cingulate gyrus are in immediate continuation
(fig. 1b, 7b). Further caudally, granular layers reappear in the cingulate gyrus, like they do anywhere in the cortex behind the central sulcus of
Rolando.
Areas LB1 and LB2, which lie in the angle, and
in part also on the dorsum of the corpus callosum, anteriorly (under the rostrum, fig. 1b) adjoin, respectively, the geniculate area FM, and the
precommissural area FN and the splenium; they
pass caudally along the dorsum of the corpus callosum, and both reach further caudally than area
LA, i.e. into the region of the splenium. Area LB1
terminates somewhat before the posterior curve
of the trunk of the corpus callosum towards the
splenium, whereas area LB2 continues as far as
the end of the corpus callosum, where it rolls into
the thin gray matter that covers the posteroventral surface of the corpus callosum.
Granular Posterior Cingulate Area LC
Caudally to the extrapolated projection line from

the central sulcus to the corpus callosum on the
median hemispheric facies, the cortex of the cingulate gyrus becomes granular again, and retains that feature all the way to the isthmus; it
thus resembles in structure the parietal cortex,
on which it actually borders at the precuneus
(fig. 1b, 7b).
The granular layer IV commences rostrally in
the internal wall of the cingulate gyrus, and as a
matter of fact in the depths of the callosal sulcus,
gradually spreading from here over the dome
142
and the external wall as well. The markedly overgrown layers V and VI which characterize the
anterior tier of the cingulate gyrus attenuate,
and their thickness returns to normal in its posterior tier. This entire granular field is termed
(granular) posterior cingulate area LC ; however,
from this field we must exclude a small part in the
internal wall of the cingulate gyrus, namely the
area over the splenium of the corpus callosum,
which forms an independent, retrosplenial region.
Within the posterior superior limbic region
LC one can further distinguish three subdivisions: the dorsal posterior cingulate area LC1 (area
EK 41) dorsally, which borders on the superior
parietal area PE ; the ventral posterior cingulate
area LC2 (area EK 42) ventrally; and lastly the posterior cingulate area limitans LC3 (area EK 43), a
part of LC that occupies a confined stretch on the
internal wall of the cingulate gyrus, between area
LA3 and the retrosplenial fields LD , LE and LF
discussed later (fig. 1b, 61).
The reciprocal topographic relations of these
areas and their disposition relative to the various
fields lying rostrally and caudally to them can be
seen by means of the markings , and (which
denote the consecutive cross-section planes in
fig. 57) or the schematic drawings of the same
sections shown in figure 61, and .
Figure 49 is a microphotograph corresponding to the section drawn in figure 61; it represents an intermediate location between the sections of figure 61 and . In that field, one can
sufficiently review all the particular relations
that I describe next, especially areas LC2 and LC3.
On the other hand, area LC1 (which can be seen
on the right side of fig. 61) actually falls outside
the microscopic field of figure 49, being located
on the next gyrus dorsally.
Areas LC1 and LC2 present sufficient analogies
to justify a description in common (cf. also plates
P48, P49 and P52a, b in the Atlas [Economo and
Koskinas, 2008]).
Dorsal LC1 and Ventral LC2 Posterior Cingulate

Areas
At the dome of areas LC1 and LC2 the cortex has
an average thickness of 2.53.0 mm. It is rather
packed with cells and clearly shows a horizontal
lamination with both its granular layers being
evident. Cells are more robust and more voluminous than in the anterior tier of the cingulate gyrus. Layer III again gains importance, while the
previous predominance of layers V and VI attenuates. One can distinguish area LC1 from the resembling region of the superior parietal lobule,
which it borders at the level of the subparietal sulcus, by the somewhat weaker development of layer III, as well as by the absence of the fine radial
striation that characterizes the parietal cortex.
Layer I is poorer in cells than the molecular
layer in the anterior tier of the cingulate gyrus
(table 11).
Layer II is well developed in area LC1, with
granule cells in its upper part, and triangular
cells in the deeper parts. Ventrally (areas LC2 and
LC3), layer II becomes poorer in cells, most of
them assuming a small, pyramidal form.
Layer III is thicker than layer III in the anterior tier of the cingulate gyrus, but still rather
thin and not particularly rich in cells. The cells of
sublayer IIIa are smaller than those of IIIb. A distinct sublayer IIIc cannot be seen. As a matter of
fact, the border with layer IV stains somewhat
more lightly (superior parietal structural type
cf. Parietal Lobe chapter). More ventrally (area
LC2), pyramidal cells are clearly larger.
Layer IV is very well developed and shows the
subdivision characteristic of the parietal lobe
especially the superior parietal lobule into an
upper, looser sublayer IVa, consisting of round
granule cells, and a lower sublayer IVb, consisting of packed pyramidal cells. Such a bipartition
is most distinct in area LC1, which directly borders on the parietal lobe; it becomes less evident
in area LC2.
Layer V is subdivided, although not very distinctly, into sublayers Va and Vb; the division becomes clearer as one moves ventrally, i.e. in area
LC2 (fig. 49) relative to area LC1. In the dorsal area
(LC1), the larger pyramidal cells are found compactly packed directly beneath layer IV, i.e. in
sublayer Va, whereas ventrally (area LC2), the larger pyramidal cells are found in sublayer Vb. In
area LC1, sublayer Va contains more than double
the number of cells of Vb (table 11). In area LC2,
sublayer Va contains 33% more cells than Vb, the
latter also staining relatively lightly. In these caudal territories of the cingulate gyrus one does not
encounter rod or corkscrew cells anymore.
Layer VI is relatively thin at the dorsal region
of the cingulate gyrus (area LC1). It is subdivided just like in the case of the superior parietal
lobule into a distinctive, clearly delimited sublayer VIa, packed with small spindle (fusiform)
cells, and a sublayer VIb, poorer in cells. In the
more ventral region of the cingulate gyrus (area
LC2), layer VI is substantially thicker, and sublayer VIa is not very clearly separated from sublayer VIb.
Areas LC1 and LC2 encircle the posterior segment of the corpus callosum in an arched form,
reaching backwards as far as the parieto-occipital
sulcus (fig. 1b, d).
Posterior Cingulate Area Limitans LC3

Area LC3 is situated inbound from area LC2, on the
ventral wall of the cingulate gyrus; it stretches all
the way to the floor of the callosal sulcus (fig. 49;
cf. also plate P50 in the Atlas [Economo and
Koskinas, 2008]). In that wall, the cortex thins
out very rapidly; individual layers undergo a considerable thinning as well, along with certain
changes in their cytoarchitectonic composition.
Layer I is thicker at the wall of the callosal
sulcus; it contains numerous myelinated fibers in
its superficial part, conferring a whitish appearance.
143
Layer II becomes extremely thin and poor in

cells (only very small pyramidal), losing its granular character at the LC3 territory.
Layer III becomes thinner with smaller and
fewer cells, and with acellular patches.
Layer IV likewise follows that general thinning compared to area LC2; it usually reaches
deep as far as the floor of the callosal sulcus (layer III(IV) in fig. 49).
Layer V shows further thinning and a cellular
rarefaction, but there are still some isolated, very
slender, lancet-shaped pyramidal cells in sublayer Vb, which are remarkably large, about 42/14
m (H/W).
Layer VI thins out very rapidly at the internal
wall and does not reach the sulcus floor, such that
it appears as if layer V borders directly on the
white matter at the lowest portion of area LC3.
Area LC3 extends only a few centimeters caudally from the highest point or upper extremity
(culmen) of the superior aspect of the trunk of
the corpus callosum (fig. 1b); it soon becomes
substituted by the retrosplenial fields, which
appear embedded into the floor of the callosal
sulcus. At that sulcus floor, layer I passes over
to the dorsum of the corpus callosum, as already described; at the angle of this passage,
cellular groups already noted in the anterior
cingulate gyrus accumulate under layer I,
such that here again in this part, in the angle
between the corpus callosum and the cingulate
gyrus and likewise onto the dorsum of the corpus callosum, area LB2 may be considered as a
continuation of area LB1.
In figure 49 one further sees, on the dorsum of
the corpus callosum at the floor of the callosal
sulcus and attached to the indusium griseum
(area LB2), an almond-shaped cellular accumulation, the ultracingulate area obtecta LF2 (area EK
48), which corresponds to the taenia tecta, considered later in the context of retrosplenial structures.
144
Agranular Retrosplenial Area LD and

Retrosplenial Area Granulosa LE
Areas LC1 and LC2 begin rostrally at the imaginary prolongation line of the central sulcus on the
median hemispheric facies; going backwards,
they cover, in a long arch, the dome of the posterior tier of the cingulate gyrus and its dorsal secondary gyri, as far as the subparietal sulcus of the
precuneus, to finally scuttle caudally into the
common trunk of the parieto-occipital and calcarine sulci (fig. 1b, d). On the other hand, area LC3
does not stretch that far caudally and it ends very
soon, because the internal wall of the cingulate
gyrus, where it coats the splenium of the corpus
callosum further augmented by the lateral disposition of the fornix [Economo, 1928e] is lined
with a particular cortical structure of heterotypic
cytoarchitecture, the so-called retrosplenial cortex (areas LD , LE and LF in fig. 1b). This structure
passes further back towards the internal brink of
the cingulate gyrus, reaching the dome at the level of the isthmus, and even extending to the external wall of the very thin isthmus itself.
The afore-mentioned series of cross-sections
in figure 57 (in the sequence , , g, f, e) offer an
exact idea of how the areas in this part of the cingulate gyrus successively change, substitute each
other or disappear. The schematic drawings of
figure 61 and , and of fig. 60g, f and e, correspond to these sections of figure 57.
The retrosplenial cortex comprises two bandlike areas, LD and LE (fig. 1b, d).
The outward-located agranular retrosplenial
area LD (area EK 44) is a thin band of granuloprival (agranular) cortex; it first abuts directly the
posterior extreme of area LC3 and then extends
caudally assuming an arched form to the internal brink of the cingulate gyrus. Figure 50 reproduces a microphotograph of a section through
the retrosplenial region, roughly corresponding
to the section in figure 61; area LD can be seen
as a thin zone just beneath the internal angle of
the dome of the cingulate gyrus. Layers II and IV
are missing from this region, such that the slim

medium-sized pyramidal cells of layers III and V
overlap. This thin heterotypic agranular zone LD
reaches a maximum thickness of only 2.03.0
mm in the internal brink of the cingulate gyrus
at the retrosplenial region, but it is over 6.07.0
cm long in the rostrocaudal direction (cf. also
plates P51, P52a, b and P109 in the Atlas [Economo and Koskinas, 2008]).
The other field, inwards from area LD , is a koniocortex that invests the deeper part of the internal wall; it is termed retrosplenial area granulosa
LE (its cytoarchitectonic modifications LE1 and
LE2 are detailed below; cf. also plates P51, P52a,
b, P104 and P109 in the Atlas [Economo and
Koskinas, 2008]). It virtually occupies all the remaining internal wall of the limbic gyrus, and it
is surrounded dorsally and outwards by the
agranular structure LD. The granular layers of
the heterotypic granulous (sensory) koniocortex
of area LE in no way adjoin the granular layers of
the ordinary granular homotypic isocortex of the
posterior cingulate area LC, being separated from
them, throughout, by the heterotypic agranular
strip of area LD (fig. 1b). Since the internal granular layers of the granular area LC and of the
granulous area LE are not equivalent in either
structure or ontogeny, we designate the granular
layer of area LE as layer III(IV).
Area LE becomes substantially thinner towards the floor of the callosal sulcus, its thickness being reduced from 2.5 to 1.7 mm. In figure
50, area LD occupies the upper one-third, while
area LE occupies the lower two-thirds of the internal wall of the cingulate gyrus facing the corpus callosum. One can readily see how this inner
thick granular layer III(IV) begins suddenly at
the boundary with area LD (which itself lacks a
layer IV) and, extending from here (in area LE1)
toward the sulcus floor, drifts more and more
toward the cortical surface, to finally end up
directly beneath layer I. In such a course, as granular layer III(IV) ascends to the surface, it progressively displaces layer III, which rapidly thins out
towards the sulcus floor to finally disappear altogether in area LE2.

Layer I is very thick (table 11), and rich in myelinated fibers in its superficial zones. This layer
increases in thickness as it approaches the sulcus
floor.
A true layer II is missing here, as noted for areas LC3 and LD (it is only still visible at the dorsal
boundary of area LD).
Layer III is composed of small, orderly arranged
pyramidal cells at the boundary between areas LE
and LD. It gradually thins out from the brink of the
gyrus toward the sulcus floor, practically disappearing completely, such that the subjacent granular layer III(IV) ends up lying directly beneath layer
I. On this account, one can distinguish an upper
part of the gyral wall, insofar as it still shows a distinct layer III, termed superior retrosplenial area
granulosa LE1 (area EK 45), and a lower part,
termed inferior retrosplenial area granulosa LE2
(area EK 46), in which a layer III cannot be discerned. The modification LE2 still shows a vestigial
layer III as a sketchy row of medium-sized, mostly
stellate cells, inserted between layers III(IV) and I.
Layer III(IV) denotes the granular layer that
makes its appearance suddenly at the gyral brink
of the inner boundary of area LE with the agranular area LD. It results from a secondary granulization of pyramidal cells at the deepest parts of
layer III in the koniocortex of area LE (fig. 50). As
stated above, it is not an imaginary continuation
whatsoever of the internal granular layer IV of
the homotypic area LC (which lies outside area
LD). By its proximity to the cortical surface, one
may conclude that layer III(IV) does not originate
from a real internal granular layer IV, with which
it bears no relation, but rather, from a granulization of layer III, which then continues to increase
toward the floor of the callosal sulcus to such an
extent that, as already said, layer III finally disappears completely, with granule cells coming to lie
immediately beneath layer I and producing a peculiar inclined rise line (fig. 50). These granule
cells are strikingly large and ovoid.
145
Layer V directly beneath is extremely thick at

the vicinity of the gyral brink and rapidly diminishes in thickness as well as number of cells as it
passes toward the valley. On the other hand, the
size of cells rather increases, such that in area LE2
one finds very slender, lancet-shaped pyramidal
cells, forming groups, with acellular patches
among them. Layer V is delimited from layers
III(IV) and VI by a lightly staining strip.
Layer VI also diminishes in thickness; it still
represents a compact layer beneath layer V in the
dorsal part of the wall (area LE1) and is quite conspicuous toward the depths of the sulcus. The
gradual general thinning is further accompanied
by reduced cell numbers per mm3 at various places. Such a reduction can reach 60% at the deepest
parts of area LE2 (table 11). The cells do not preserve their true spindle (fusiform) shape, but
rather become triangular and small.
Due to the cellular depletion of the two deeper
layers, as well as the disappearance of pyramidal
cells from layer III, a granulization prevails in the
overall cytoarchitectonic picture and characterizes this cortex as granulous, i.e. as koniocortex.
Since, however, this cortex develops from a sixlayered (hexalaminar) pattern, it can be considered as constituting another isocortical heterotypy.
Function
One might very well suppose that this sensory
cortex, located in the middle of the so-called
rhinencephalon (fig. 5g), represents the primary
olfactory cortical center, an interpretation further
supported by the argument that the retrosplenial
area granulosa LE is exceptionally well developed
in macrosmatic animals, to the point of extending in such species dorsally, over the superomedial edge or border of the cortical mantle, to the
superolateral hemispheric convexity.
146
Ultracingulate (Posterior Intralimbic) Area LF
At the floor of the internal wall of the cingulate

gyrus, layers III, III(IV) and IV stop completely;
only cells of layers V and VI roll over, alongside
layer I, to the dorsum of the corpus callosum,
where they form a thin bed (fig. 50).
At the angle of the valley floor of the callosolimbic retrosplenial region they form a fold somewhat thicker and denser in cells than the anterior
tier of the cingulate gyrus. One may consider this
point as an individual, band-like field of the angle of the corpus callosum, termed posterior ultracingulate area LF1 (area EK 47); it is characterized by fine, large, lancet-shaped pyramidal cells,
orderly arranged in rows (fig. 50, 61). These features clearly distinguish area LF1 from the cellular conglomerate of the anterior ultracingulate
area LB1, of which LF1 is a direct continuation (cf.
fig. 1b).
At the point of contact with the dorsum of
the corpus callosum, layer I thickens and contains increasingly more numerous cells (as already explained for the indusium griseum),
such that in cross-section it forms an oval field,
rich in cells, and, in the rostrocaudal direction,
a string-like collection of gray matter, termed
ultracingulate area obtecta LF2 (area EK 48).
That cellular string accompanies the taenia tecta towards the front with considerable individual variations. (We already saw area LF2 in figure 49, where a distinctly oval structure is observed; caudally, this cellular string continues
into the fasciola cinerea.)
Medial to this structure, only layer I is seen
again and a few ovoid cells, passing onto the dorsum of the corpus callosum as the area of indusium griseum LB2 (area EK 40).
Both areas LF1 and LF2 simply consist of a layer I with a cellular row beneath; they belong
along with areas LB1 and LB2 to allocortex (cf.
also plates P51 and P52a, b in the Atlas [Economo
and Koskinas, 2008]). Areas HE and HF (fig. 50)
are detailed in the next chapter.
Function
One could logically suppose, from its topographic relation with the koniocortex of area LE , which
is without question the primary sensory olfactory
center, that the entire retrosplenial region with all
its areas is in some way related to the function of
smell. It is often mentioned that even the entire
limbic gyrus belongs to the so-called rhinencephalon (fig. 5g).
However, when one thinks of the small role
that the sense of olfaction plays in humans, as
well as the comparatively meager development of
the koniocortex of area LE compared to animals,
it becomes more and more apparent that the anterior part of the limbic gyrus, which is still well
developed in humans, may have acquired a new
function in the course of evolution. Based on its
agranular motor cortical structural type 1, this
residual part of the limbic gyrus might possibly
mediate sympathetic efferent functions (cf. Insular Lobe chapter).
Transition of the Limbic Lobe to the Isthmus
In figure 50, on the dorsum of the corpus callosum, we see certain structures appearing between
the area obtecta LF2 and the area of indusium
griseum LB2, which already belong to the region
of the hippocampus and are hence designated
with the symbols HE and HF.
Since it is quite difficult to visualize the successive areas of the cingulate gyrus in their correlative shifts in its dome and walls, I provide a
set of schematic drawings of consecutive sections
through this entire region, to which I already referred:
Figure 57 schematically depicts the entire cingulate gyrus, from its outset at the paraterminal
gyrus (carrefour olfactif of Broca), around the
corpus callosum and the fimbria, all the way to
the uncus. The dashed lines indicate the planes to
which the schematic cross-sections of figures 59
61 correspond. The series of sections commences

at the uncus and hippocampus (fig. 59AE); a series of sections then follows through the isthmus
and retrosplenial region (fig. 60ag); then four
sections through the superior part of the limbic
gyrus (fig. 61), of which, specifically, figure
61 through the superior extreme of the retrosplenial region, fig. 61 through the posterior
one-third, figure 61 through the mid-point, and
figure 61 through the anteriormost part of the
rostral cingulate gyrus.
Figure 61 is a schematic representation corresponding to the microphotograph of figure 48,
namely a cross-section of the cingulate gyrus
with its cortex of agranular area LA, and, specifically, the external wall (area LA1), dome (area
LA2) and internal wall (area LA3, inside the callosal sulcus). From the depths of the sulcus, the
cortex courses again along the dorsum of the corpus callosum (C.c.) with areas LB1 and LB2.
Figure 61 schematically depicts the investment of the more caudal parts of the dorsum of
the corpus callosum by areas LB1 and LB2. On the
other hand, agranular areas LA become substituted by granular area LC3 at the sulcus depth. At
this level of the cingulate gyrus, areas LA and LC
briefly intermingle at the dome, over the central
one-third of the corpus callosum. At the depths
of the sulcus [Economo, 1929d], the cortex rolls
into the corpus callosum (C.c.) with areas LB1 and
LB2.
Figure 61 schematically shows (cf. microphotograph of fig. 49) how granular areas LC1,
LC2 and LC3 occupy the entire posterior tier of the
cingulate and its adjacent gyri.
Figure 61 schematically shows (cf. microphotograph of fig. 50) how granular areas LC become gradually substituted at the lateral internal
wall of the cingulate gyrus and dorsally displaced
by the agranular and granulous retrosplenial areas LD and LE . At the point where the cortex recurves over the corpus callosum, the new cellular
groups LF1 and LF2 make their appearance.
147
Such a substitution is complete in the next section (fig. 60g). The retrosplenial areas LE1, LE2
and LD cover the entire (thin) dome and wall of
the limbic gyrus ( g.l. in fig. 60). Area LB is substituted at the dorsum of the corpus callosum, in the
vicinity of the isthmus, by the lancet-shaped pyramidal cells of area LF1 and area obtecta LF2
(fig. 60f). The remaining sections (fig. 60ae, g)
illustrate the transition from retrosplenial to hippocampal structures via the isthmus. At the region of the isthmus, they reach driven back by
the hippocampal structures that make their appearance at the depths of the wall the lip of the
internal wall [Economo, 1927d].
To better understand such a transition, let us
imagine that the retrosplenial region between the
solid lines II and IIII in figure 57 is excised and
stretched, like in figure 58a, so that the gyri and
the sulci would appear distended and flattened
out. The caption of figure 58 sufficiently clarifies
the gross anatomic (macroscopic) relations. In
148
figure 58b, one sees all the areas of this region

with their boundaries readily discernible by
means of the various patterns of hatching, so that
the intricacy and complexity of the individual cytoarchitectonic fields, as well as the small individual details of cytoarchitectonic structure,
might be better studied in this scheme, as opposed to figure 1b, e.g. in the fasciolar gyrus (g.f.),
the gyri of Anders Retzius (g.ar.) [Retzius, 1898]
or their adjacent structures x and y. This scheme
moreover shows the intricate relations between
retrosplenial and hippocampal fields. A further
explanation is unnecessary. A closer study of figure 58 and the corresponding series of cross-sections in figure 60 will clarify much better than
words the way in which cytoarchitectonic areas
extend in this transition zone between the retrosplenial region, the isthmus, and the hippocampus. I discuss the transition to the hippocampal
gyrus in the following chapter.
Hippocampal (Inferior Limbic)

Lobe
Hippocampal Gyrus, Dentate Gyrus and Uncus
Preliminary Anatomic Remarks
We now arrive at the discussion of the hippocampal gyrus (the second limbic convolution of the
French literature [Economo, 1927d]).6 The agranular retrosplenial area LD (area EK 44), and the
superior LE1 and inferior retrosplenial area granulosa LE2 (areas EK 44 and 46) cover the entire
depth of the inner wall of the limbic gyrus at the
caudal segment of the corpus callosum (fig. 1b); at
the region of the isthmus, these areas gradually
rise, over the internal brink of the gyrus, toward
the dome (cf. fig. 58b and 60eg) avoiding hippocampal areas at the depths of the wall vesting
them entirely; they then sink into the external wall
of the hippocampal gyrus somewhat further down,
at the interface with the trunk of the calcarine sulcus. (The microphotograph of the section through
the isthmus in figure 51 approximately corresponds to the schematic drawing of figure 60e.)
The microphotographs shown in figures 5155 of this chapter represent a series of sections through the hippocampal gyrus. They do not follow the same order as the description of the
corresponding cytoarchitectonic areas in the text; rather, they
represent topographic views directly after the series of sections through the superior limbic gyrus (fig. 4850 of the previous chapter), beginning at the rostrum of the corpus callosum,
continuing around the splenium, and ending at the uncus.
On the other hand, the gray matter of the posterior ultracingulate area LF1 (area EK 47) and the
ultracingulate area obtecta LF2 (area EK 48) adheres closely to the indusium griseum LB2 (area EK
40, labeled i in fig. 60eg) at the corpus callosum;
it abandons the corpus callosum at the point where
the indusium ends, passing over to the fimbria ( fi.
in fig. 60e), which makes its appearance directly
underneath the end of the corpus callosum.
Hippocampal areas already appear entering
the space where areas LE2 and LF1 begin to split
(fig. 58b, 60ge), gradually replacing them in the
internal wall of the limbic gyrus (g.l.), which now
becomes the hippocampal gyrus (g.h.). Thus, the
hippocampal gyrus is in a way the direct downward and forward continuation of the limbic gyrus (fig. 1b, 57); at that passage, it comes to lie
immediately on the fusiform gyrus (T4) of the
temporal lobe.
More anteriorly, toward the temporal pole, the
hippocampal gyrus bends laterally, i.e. towards
the median hemispheric facies, in the shape of a
hook, simultaneously thickening to form the uncus (U). The fimbria (fi.) is finally attached to the
lateral surface of the hook-shaped uncinate bend
that faces the hemisphere, at the margin of the
velum terminale (inferior choroidal point) of
Aeby [1883].
Fig. 51. A perpendicular section traversing the isthmus. Inferior retrosplenial area granulosa LE2 (area EK 46). Rhinal
area limitans HC (area EK 99). Presubicular area granulosa HD(1, 2, 3): limitans (area EK 100), middle (area EK 101), glomerular (area EK 102). Subicular pyramidal area HE1 (areas EK 103104). Pyramidal area of Ammons horn HE2 (areas
EK 105106). Dentate area HF (area EK 107). Transition of the retrosplenial part of the limbic gyrus (on the right side
of the figure) to the part of the hippocampal gyrus (left side). The fimbria (fi.) has replaced the corpus callosum; l.af.,
lamina affixa, which roofs the inferior ventricular horn (V); s.f.d., fimbriodentate sulcus; f.d., fascia dentata; s.h., hippocampal sulcus. !14.
In this entire path, from the splenium of the

corpus callosum to the uncus, the fimbria maintains a similar relation to the hippocampal gyrus,
as the corpus callosum does to the limbic gyrus.
Just as the cortex of the limbic gyrus encircles the
dorsum of the corpus callosum (Balkenrcken),
being occupied at the retrosplenial region of this
bent passage by area LF1 and at the outer edge of
the dorsum of the corpus callosum by the vermiform thickening of area LF2, whilst the indusium
150
LB2 spreads as a fine lamella over the remainder

of the corpus callosum from one hemisphere to
the other, so does the gray cortex of the hippocampal gyrus henceforth merge ventrally of the
splenium of the corpus callosum into the fimbria,
i.e. the cessation of the callosal radiation on the
cornu of the fornix (fig. 60da).
The indusium (i in fig. 58a) does not continue
on to the fimbria, since the temporal and occipital lobes of the two hemispheres are not united;
rather, it becomes folded, ventrally and around

the splenium of the corpus callosum, and continues as a fine lamella of gray matter under the corpus callosum and into the septum pellucidum. In
this way, the internal wall of the limbic gyrus becomes the internal wall of the hippocampal gyrus. Just as the limbic gyrus is separated from the
corpus callosum by the callosal sulcus (S.cc. in
fig. 58a), so is the hippocampal gyrus now separated from the fimbria by the hippocampal sulcus. At the base of this hippocampal sulcus, the
cortical margin rolls into the fimbria ( fi. in
fig. 58), just like it earlier made the transition to
the corpus callosum. This transition point forms
a bulge on the inner side of the lateral ventricle,
Ammons horn (A in fig. 60).
However, the cortical margin does not lie as
flatly on the fimbria as it did earlier, in its passage
to the dorsum of the corpus callosum; rather, it
rolls once again spirally and is further covered by
a thin bilayer of gray matter, the fascia dentata of
Tarin [1750] ( f.d. in fig. 60ea). In cross-section,
this capuche-like covering assumes a horseshoe
shape. (Figure 52, a microphotograph of a section
approximately through the middle of the hippocampal gyrus as indicated in the schematic drawing of figure 60a, shows these relations rather
clearly.) This rolled-up cortical margin, which
accompanies the hippocampal gyrus from the
isthmus, or rather even from the splenium of the
corpus callosum, all the way to the uncus shaped
like a vermiform pad, constitutes the dentate gyrus (labeled g.d. in fig. 58a). It can be seen at the
base of the hippocampal sulcus (s.h.); it is separated from the fimbria through a shallow groove,
the fimbriodentate sulcus (s.f.d. in fig. 52).
The part of the dorsal (internal) wall of the
hippocampal gyrus, against which the dentate
gyrus rolls up, is called subiculum; the free part
of the wall above it, together with the brink of the
dome, constitutes the presubiculum.
Caudally, at the level of the splenium of the
corpus callosum, the dentate gyrus gradually
merges into the fasciolar gyrus ( g.f. in fig. 57, 58a)
and further distantly into the taenia obtecta (T.t.

in fig. 58a). Rostrally, the dentate gyrus directly
merges into the band (nastro or bandelette) of
Giacomini [1883] (cf. B.G. in fig. 57 and area HF
in fig. 1b), which transversely crosses, from the
outside toward the inside, the posterior extreme
of the uncus. The dentate gyrus terminates on the
lateral external surface of the uncus, facing the
interior of the hemispheres, at the velum terminale of Aeby [1883].
Immediately rostral to this point, on the external as well as the dorsal surface of the uncus,
there is the terminal expanse of the temporal
piriform area TJ (area EK 92), forming the posterior end of the small olfactory gyrus and finally
ending at the semilunar gyrus of the uncus. At
this end (vesting the cortex of the semilunar gyrus), area TJ appears somewhat modified in its
fundamental structure and becomes the transition hippocampotemporal piriform area TJ H (table 2). Its deepest cellular layers frequently reveal
a very close continuity with the cells of the nucleus of the amygdala (NA); for that reason, this terminal segment of the temporal piriform area is
also called periamygdalar piriform area [Economo, 1929d].
The schematic drawings of figure 59AG best
clarify the relations of this region and its areas at
the polar transition of the hippocampal gyrus to
the uncus.
Hippocampal Areas of the Entorhinal Region
The hippocampal gyrus comprises a granulous

cortical field, i.e. a koniocortex, specifically in its
superointernal wall surface in the presubiculum
(fig. 58b, 60af, 59BG at HD, and fig. 5154 at
HD1, 2, 3). Such a koniocortex is found in the immediate continuation of the granulous limbic
area LE1, 2 (retrosplenial area granulosa), replacing the latter in the deeper parts of the internal
wall, at the point where area LE begins to roll
into the gyral dome at the level of the isthmus
151
(fig. 1b, 58b, 60df); it is termed presubicular

area granulosa HD and sets out at the isthmus,
occupying the brink and upper dorsal wall of the
hippocampal gyrus, i.e. the presubiculum, and
covering in a rostrocaudal direction the entire
distance from the isthmus to the uncus curve
(fig. 1b, 5154).
The lower part of the wall (i.e. the subiculum),
and the transition territory to the fimbria (i.e.
Ammons horn), as well as the core of the upturned dentate gyrus, are all vested in their entire
rostrocaudal length by the pyramidal area HE ,
which is the immediate continuation of the posterior ultracingulate area LF1 (fig. 1b, 58b, 60ag);
both of these are solely composed of a molecular
layer and a rather thick subjacent layer of conspicuously large pyramidal cells. In the series of
microphotographs of figures 5154, this field of
the subiculum and Ammons horn (areas HE1
and HE2) can be seen very well.
The upturned margin of the dentate gyrus,
together with its horseshoe-shaped covering of
the fascia dentata (f.d.), forms the area dentata
HF, which is thus the continuation of the ultracingulate area obtecta LF2 (fig. 58b). To the
inside, one only finds the white matter string of
the fimbria (fi.); its superficial gray layer vests
the inferior ventricular horn with an extremely
thin leaf, the lamina affixa (marked l.af. in
fig. 51 and 52), which then continues to the opposite hemispheric side in the fields of the hindbrain.
All the areas mentioned attenuate in the rostrocaudal direction, stretching as fine strips of
cortex on the internal wall and dome of the hippocampal gyrus, which they cover (fig. 1b).
To the exterior of the presubicular area granulosa HD , which occupies the upper brink (presubiculum) of the hippocampal gyrus, the dome
is covered in its anteroposterior axis by area HC ,
which is also like a strip and constitutes the
posterior continuation of the cortical structures
of the uncus (fig. 1b, 58b, 59CG and 60ae).
The uncus is clad in a rather particularly struc-
152
tured cortex, which justifies its denomination

of entorhinal region (I also include the frontal,
pad-like extreme of the hippocampal gyrus
with the uncus). One can divide it into three areas, HA , HB and HC (fig. 1b, d). Those three
entorhinal areas, as well as areas HD , HE and
HF, do not belong to the ordinary six-layered
(hexalaminar) isocortex; having instead a quite
different structure, they form parts of the allocortex (fig. 5d).
The boundary between this allocortex of the
uncus and the isocortex of the temporal lobe
(i.e. the agranular temporopolar area TG and
the hippocampotemporal area TH ) is found anteriorly in the rhinal (olfactory) sulcus, posteriorly in the temporo-occipital (or collateral) sulcus, on the lower (ventral) lip of the hippocampal gyrus or its lower wall (fig. 5254, 59CG,
60a, b).
The microphotograph of figure 54 shows a
coronal section at the rostral extreme of the hippocampal gyrus, already falling into the region of
the uncus, and approximately corresponding to
the schematic section drawing of figure 59C; figure 55 shows a section through the polar anterior
part of the uncus. In both microphotographs one
can discern the areas of the uncus rather well (cf.
also in that regard the work of Levi [1903]). I now
move on to a discussion of the cytoarchitectonics
of these individual areas.
Fig. 52. Coronal section through the middle of the hippocampal gyrus with the presubiculum (HD ), subiculum
(HE1), alveus (A), Ammons horn (HE2) and dentate gyrus
(HF ). Agranular hippocampotemporal area TH (area EK
89). Rhinal area limitans HC (area EK 99). Presubicular
area granulosa HD(1, 2, 3): limitans (area EK 100), middle
(area EK 101), glomerular (area EK 102). Subicular glomerular pyramidal area HE1 (area EK 103). Subicular pyramidal area simplex HE1 (area EK 104). Pyramidal area of
Ammons horn HE2 (area EK 105). Dentate area HF (area
EK 107). Remaining abbreviations as in figure 50. !14.
153
Fig. 53. Coronal section passing through the anterior segment of the hippocampal gyrus and uncus, at the point
where the fascia dentata begins to cross the uncus as the band of Giacomini. Fusiform area TF (area EK 87). Agranular
temporopolar area TG (area EK 91). Parauncinate area HB (areas EK 9798). Granulous presubicular transition of rhinal
area limitans HC(D) . Presubicular area granulosa HD(1, 2, 3) (areas EK 100102). Subicular pyramidal area HE1(, ) (areas
EK 103104). Pyramidal area of Ammons horn HE2 (area EK 105). Pyramidal area of digitate gyrus of uncus HE3 (area
EK 106). Dentate area HF (area EK 107). Abbreviations as in figure 51. !10.
154
Fig. 54. Coronal section through the transition zone of the rostralmost segment of the hippocampal gyrus in the
uncus. Agranular temporopolar area TG (area EK 91). Uncinate area HA (areas EK 9496). Parauncinate area HB (areas
EK 9798). Granulous presubicular transition of rhinal area limitans HC(D) . Presubicular area granulosa HD (areas EK
100102). Pyramidal area HE(1, 2, 3): subicular (areas EK 103104) of Ammons horn (area EK 105) and of digitate gyrus
of uncus (area EK 106). !12.
Uncinate Area HA
This cortical area in the polar protrusion of the

uncus is characteristically thick and contains
large cells. Figure 56, at a higher magnification
(!45), shows the middle part of the cortex shown
in figure 55 at !10. Cortical thickness, particularly toward the polar segment, reaches 3.0 mm;
caudally, toward the (inner) concavity of the uncus, it diminishes to 2.0 mm.
Upon the macroscopic inspection of fresh
brain tissue, the surface of this cortex appears
whitish instead of grayish and has a peculiar honeycomb pattern, owing to the penetration of the
superficial zone of layer I by numerous myelinated fibers deriving from the olfactory tract (ra-
155
Fig. 55. Coronal section through the anterior pole of the uncus. Temporopolar area TG (area EK 90). Uncinate area HA
(areas EK 9495). V, inferior ventricular horn; NA , nucleus of the amygdala. !10.
dix lateralis). That reticular pattern ensues from

gray matter islets being interspersed amidst this
myelinated fiber network (the substantia reticulata of Arnold [1851]).
Although the entorhinal cortex of the uncus
and the hippocampal gyrus belong to the allocortex, they nevertheless display a clear and repetitive multilamination. Thus, one can clearly distinguish six definite layers, e.g. in areas HA and
HB , like in isocortex. However, the embryogenet-
156
ic evidence does not allow us to draw any homologies between this allocortex and the six layers of
isocortex, even if one were able to trace contiguities of cellular topography between individual
layers at the allocortical-isocortical boundary. It
should then be implicit that when we retain the
same layer numbering system, from superficiality to profundity, for the two cortical types, this
is done solely on practical grounds in order to establish the respective heights of layers in the two
Table 12. Summative table of quantitative data in six ground areas of the hippocampal (inferior limbic) lobe. Overall
layer thickness similar between dome and wall. Some additional values supplemented from tables I, III, V and VI in
Economo and Koskinas [1925, pp. 794801].
Area
symbol
Area name
Cortical
layer
Average layer
thickness, mm
HA
uncinate area
I
II
IIIa
IIIb
V
VIa
VIb
0.36
0.34
Cell content
cells/mm3
Cell size, m*
H(minmax)/W(minmax)
0.33
0.53
0.14
5,000
20,000
20,000
22,000
24,000
27,500
6,000
1025/810
3050/2030
1525/1015
2530/1518
3035/2025
2025/1015
1530/15
0.99
HB
parauncinate area
I
II
III
V
VIa
VIb
0.40
0.40
0.78
0.21
0.50
0.18
5,000
15,000
20,000
20,000
20,000
5,000
10/8
30/20
2530/1520
2030/1518
20/15
25/815
HC
rhinal area limitans
I
II
III
V
VIa
VIb
0.35
0.20
0.60
0.30
0.32
0.24
6,000
40,000
15,000
35,000
30,000
10,000
1018/78
10/20
25/19
35/25
20/1015
15/810
HD
presubicular area granulosa
I
II+III
V+VI
0.40
0.61
0.70
5,500
120,000
37,500
1012/810
815/8
1520/1015
HE
pyramidal area
I
V+VI
1.05
1.25
2,500
23,500
20/30
2080/1020
HF
dentate area
I
VI
0.50
0.15
1,500
250,000
10/8
1012/810
situations and also to indicate in part allocortical

layer proportions relative to the matched isocortical layers (cf. fig. 55).
Layer I contains plentiful glial nuclei in its superficial myelinated zone; its deeper zone contains nerve cells of a size that is exceptionally
large for a molecular layer (table 12).
Layer II does not appear as a continuous
granular layer; instead, it comprises individual
large roundish cellular groups of large stellate
cells (called glomeruli). In a 25-m-thick section, one can count about 30 cells in each glomerulus; the cells are polygonal, have a large nucleus and nucleolus, and stain very intensely. Occasionally, certain glomeruli are composed of
smaller cells (e.g. under the asterisk in fig. 56).
Between individual glomeruli, layer I reaches
deeply into the tissue all the way to layer III. It is
these globular glomeruli that shimmer through
the molecular layer as gray masses and give the
157
surface its mesh-like appearance. Glomeruli may

reach 0.5 mm in diameter.
Layer III beneath consists of a deeper sublayer
IIIb with regularly formed and arranged slender
pyramidal cells. The upper sublayer IIIa consists
of somewhat smaller, triangular and polygonal
cells, arranged very irregularly and grouped in
rather dense hoards at places; among them, one
finds oligocellular or acellular patches, which
impart a ragged appearance to the upper border
of layer III. An extremely cell-poor, lighter horizontal strip separates layer III from layer II.
Layer V(IV) is found just beneath layer III in
lieu of a layer IV also in this case; it is an almost
totally acellular, sharply demarcated white band
(lamina dissecans of Rose [1926b]), composed of
a bundle of myelinated fibers (fig. 56).
Layer V is thick and contains large pyramidal
cells; they stain deeply, are packed extremely
densely, and are arranged orderly like rows of soldiers.
Layer VI comprises a thicker sublayer VIa
with numerous spindle (fusiform) and polygonal
cells, giving a rather dense appearance; its upper
cellular rows bordering on layer V are sometimes
much looser in texture than its deeper rows bordering on sublayer VIb (fig. 56). On the contrary,
the altogether oligocellular sublayer VIb is extremely thin, parvicellular and lightly stained.
The demarcation vis--vis the subjacent white
matter is extremely sharp.
The entorhinal uncinate area HA vests the entire anterior polar segment of the uncus, as can be
seen in the coronal overview microphotograph of
figure 55 (at a magnification of !10), and in the
schematic drawings of figure 59AD. Dorsally, it
extends on the external crus of the uncus, without, however, reaching exactly as far as the band
of Giacomini; on the internal crus, which continues into the hippocampal gyrus, area HA imperceptibly passes caudally into the area HB (discussed next) as early as the (inner) concavity of
the uncus (fig. 1b, d).
158
Entorhinal Uncinate Area Modifications HA1, 2, 3

The term primary uncinate area HA1 (area EK
94; cf. plates P100 and P105 in the Atlas [Economo and Koskinas, 2008]) is reserved for the anterior segments of ground area HA that only have
a single white band above layer V [Economo,
1927d].
Quite often, that white band (IV in fig. 55 and
V(IV) in fig. 56) is doubled, such that layer V may
come to lie between two lightly stained striae.
Such a doubling particularly occurs on the internal crus (concavity) of the uncus; this modification is called secondary uncinate area HA2 (area
EK 95; cf. plates P105 and P106 in the Atlas
At the passage to the hippocampal gyrus, on
the other hand, the superior white stria is missing, and the remnant acellular stria thus solely
appears beneath layer V; we term this modification tertiary uncinate area HA3 (area EK 96; cf.
2008]).
Toward the rhinal sulcus, the glomeruli of layer II lose their fine roundish form and become
scarcer [Economo, 1928e].
This lamina dissecans is more or less clearly
observed in all multilaminar allocortical areas of
the hippocampal gyrus, e.g. in areas HA, HB , HC
and HD ; it first begins to disappear in the monostratified areas HE and HF ; on the outside of the
allocortex, it ends at the boundary with the isocortex (fig. 5). One may follow the structure
through in figures 5551.
Hippocampotemporal Piriform Transition Areas

TJH1, 2, 3, 4 in the Semilunar Gyrus
It is interesting to note that the small annular ambient gyrus of the uncus (piega circondante of
Sterzi [19141915] in the Italian literature [Economo, 1928e]) is also clad in a cortex of the entorhinal structural type HA .
Fig. 56. Uncinate area HA (areas EK 9495). Detail from figure 54. Beneath the asterisk (*), glomeruli of Arnolds substantia reticulata. !45.
159
Fig. 57. Schematic drawing of the cingulum to depict the position of sections in figure 58ab, 59AG, 60ag and
61. C. = Calcarine sulcus; C.Br. = Brocas parolfactory field; fi. = fimbria; g.ar. = gyri of Anders Retzius; g.d. = dentate
gyrus; g.f. = fasciolar gyrus; g.h. = hippocampal gyrus; g.l. = limbic (cingulate) gyrus; g.rl. = retrolimbic gyrus;
G.B. = band of Giacomini; R.cc. = rostrum of corpus callosum; s.cc. = sulcus of corpus callosum; s.h. = hippocampal
sulcus; Spl.cc. = splenium of corpus callosum; s.po. = parieto-occipital sulcus; Tr. = trunk of calcarine sulcus; U =
uncus.
On the other hand, the semilunar gyrus (piega

semilunare [Economo, 1928e]), within the ambient gyrus, is the direct continuation of the lateral
olfactory gyrus, as already stated; accordingly, it
is clad in a cortex of the cortical margin structural type TJ, which is quite different from HA
and sharply defined from it. This hippocampotemporal or periamygdalar piriform area TJ H is
allocortex and adjoins, with its deeper cell layers,
the nucleus of the amygdala (NA).
This transition area can be discerned in the
schematic drawings of figure 59AC, as well as
160
in plate P105 of the Atlas [Economo and Koskinas, 2008] (marked as 1 on the right side of the
microphotograph). Depending on the intimacy
of the cell continuity with the amygdala, as well
as on the cytoarchitectonic attributes of its super ficial cell layers, which beneath the molecular layer consist of stellate and pyramidal-like
cells, area TJ H may be subdivided into a further
three to four secondary modifications, such as
areas TJ H1 through TJ H4 (table 2) [Economo,
1929d].
Fig. 58. The retrosplenium in-between the levels II and IIII of figure 57. a Schematically stretched and with opened
sulci. b The same, with the areas of the limbic gyrus (LA LF ) and the hippocampal gyrus (HC HF ) shaded. f.d. = Fascia dentata; fi. = fimbria; g.ar. = gyri of Anders Retzius; g.d. = dentate gyrus; g.f. = fasciolar gyrus; g.h. = hippocampal
gyrus; g.l. = limbic gyrus; g.rl. = retrolimbic gyrus; i. = indusium griseum; s.c.c. = sulcus of corpus callosum; Spl.C.c. =
splenium of corpus callosum; T.t. = taenia tecta; Tr. = trunk of calcarine sulcus; y = the small warp of dwarf gyri issued
from the upturned segment of the fascia dentata; x = small secondary gyri of the subiculum.
Parauncinate Area HB
At the transition zone of the uncus to the hippocampal gyrus, area HA becomes the parauncinate
area HB (fig. 1b, d), vesting with its concavity the
dome of the anterior segment of the hippocampal
gyrus from the knee of the uncus up to the vicinity of the rhinal (collateral) sulcus, where it terminates caudally (fig. 53, 54; cf. also plate P105 in
the Atlas [Economo and Koskinas, 2008]). Area
HB only differs from area HA in specific points,
especially the form of layer II, which no longer
consists of glomeruli, but forms instead a band of
large cells. Overall, the cortex of area HB is somewhat thinner than area HA, measuring 2.5 mm in
thickness.
Layer I is unusually thick, just like in area HA

(table 12); it contains numerous myelinated fibers, too.
Layer II consists of voluminous, polygonal
and dark-staining stellate cells with a large nucleus and nucleolus, like in area HA . However, the
cells are no longer segregated in neat glomeruli,
forming instead an untidy, lacunar layer. At the
same time, this layer contains the largest cells in
this area. Below those large cells, i.e. between layers II and III, one finds a paucicellular white
band, more distinct than in area HA .
Layer III beneath consists of pyramidal cells,
more uniformly distributed than in area HA; but
even here, the tissue shows alternating denser
and clearer cellular patches. Here again, the up-
161
per border of the layer is often irregular and undulate.

Layer IV is totally absent from area HB as
well; in its place one does not find either granule
cells or the acellular white stria, like in area HA;
the pale lamina dissecans is hardly noticed touching layer V.
Layer V is composed of densely-packed pyramidal cells, deeply-staining, and arranged in a
single layer (table 12). A lightly-staining lamina
dissecans may also appear below layer V in area
HB in its rostrodorsal parts; on this basis, one can
subdivide area HB into two modifications, the
primary HB1 (area EK 97; cf. plates P101 and
P102a, b in the Atlas [Economo and Koskinas,
2008]) and the secondary parauncinate area HB2
(area EK 98; cf. plates P102b and P106 in the Atlas
[Economo and Koskinas, 2008]). A tertiary parauncinate area HB3 (table 2) is found on the dorsal
brink of the uncus (cf. plate P102b in the Atlas
[Economo and Koskinas, 2008]); in this modification, one further observes the gradual disappearance of layer III or, more precisely, its
merging into layer V, while layer VI can still be
clearly discerned [Economo and Koskinas, 1925,
p. 754].
Layer VI is more or less distinguished from
layer V by the pale secondary lamina dissecans,
when present; alternatively, the two layers may be
confounded, when the stria is missing. Layer VI
contains mostly real spindle (fusiform) cells, with
some pyramidal cells among them. As sublayer
VIb is thin and rather acellular, the demarcation
from the white matter is very sharp.
Area HB does not cover the hippocampal gyrus all the way to the depths of the rhinal sulcus;
it stops slightly underneath the inferior edge of its
wall, where it borders on the temporopolar area
TG, and as a matter of fact, its agranular margin
TG (area EK 91). Caudally, area HB only extends
for a very few cm; the course of areas HB and HA
can be traced in the schematic drawings of figure
59AF.
162
Rhinal Area Limitans HC
The rhinal area limitans HC stretches inwards

and posterior to area HB on the (inner) concavity
of the uncus, from a pointed anterior (fig. 1b),
over the entire remaining segment of the dome of
the hippocampal gyrus, to the isthmus caudally,
where it becomes pointed again (fig. 5451; cf.
also plates P102b, P103c, P104d, P107 and P110 in
the Atlas [Economo and Koskinas, 2008]). Its
maximum thickness is found approximately at
the anterior one-third of the hippocampal gyrus;
thus, it assumes the shape of a spindle (fig. 1b).
Ventrally, area HC accompanies and surrounds the koniocortex of the hippocampal presubicular area granulosa HD (detailed next) along
its entire rostrocaudal trajectory on the dorsal lip
of the hippocampal gyrus all the way to the isthmus, just like the koniocortex of area LE in the
retrosplenial region is bounded by the agranular
area LD.
The cortex of area HC is not as uniformly
structured as that of the other areas, being laden
with regional as well as individual variations of
different grades that would allow one to cytoarchitectonically subdivide it in various ways. Generally though, one may admit that its cortex is
thinner (reduced to 2.0 mm on the average), with
the cells in layers III, V, and VI more loosely organized, compared to area HB . One capital difference from area HB [Economo, 1929d] is that
layer II in area HC consists of a rather compact
and wide row of very small stellate cells.
Layer I remains rather thick (table 12) and
displays a cellular disposition similar to the other
areas of the uncus.
Layer II in this region does not yet contain
true granule cells; rather, it is composed of stellate, navicular and polygonal cells, which are
similar to the cells in layer II of the parauncinate
area HB , albeit considerably smaller. The layer is
rather dense and sharply delimited over and under, beginning nevertheless to acquire more the
appearance of a granular layer than in areas HA
and HB . Its cells gradually diminish in size toward the hippocampal sulcus (fig. 54), where layer II rolls into the thick granulous layer of the
koniocortex of the presubicular area granulosa
HD ; on the other hand, these polygonal pseudogranule cells nonetheless confirm again their relationship with the corresponding polygonal cells
of layer II in area HB by the fact that, in some of
the more rostral parts of area HC, sporadic glomeruli of larger cells occasionally stray into it
from area HB . In the rhinal (collateral) sulcus,
layer II merges into the similarly magnocellular
layer II of the temporopolar area TG (area EK 90).
In caudal parts of area HC, cells of layer II become
even smaller and assume a more granule-like appearance (fig. 51, 52).
Layer III consists of pyramidal cells; however,
their number varies greatly. At some places one
sees large acellular patches, and at others a rather
compact layer III. In the first situation, the overlying layer II, being quite rich in stellate cells,
markedly contrasts with the poorer layer III
(fig. 53, 54) and, as in such cases layers V and VI
also comprise acellular patches and are equally
poor in cells, area HC takes on a totally particular
appearance, almost as if it consisted solely of layers I and II; thus, it gets to look very much like the
koniocortex of area HD on which it borders. We
consider that modification a presubicular granulous transition HCD of the rhinal area limitans
HC (table 2); it seems to appear most often in the
caudal segment of the hippocampal gyrus near
the isthmus (fig. 51), and in its anteriormost segment in the vicinity of the (inner) concavity of the
uncus. On the other hand, at the dome of the
middle segment of the hippocampal gyrus, layer
III is usually well formed and rather cellular, and
layers V and VI are distinct, thus retaining the
distinctive cytoarchitectonic form of area HC
proper. Still, I must emphasize that, besides these
regional variations, area HC often shows substantial variations among individuals as well.
Layer IV is absent. Area HC, which constitutes
part of the allocortex, also borders on agranular
isocortical formations of the temporal lobe

(fig. 52), such as the agranular modifications of
the temporopolar area TG (area EK 91) and the
hippocampotemporal area TH (area EK 89) at the
temporo-occipital sulcus. Only in those places
behind the rhinal sulcus, where an actual hippocampotemporal bridge gyrus is found, can one
see granule cells issued from layer IV of fusiform
area TF (area EK 87) passing into the ventralmost
parts of area HC, and one could hence define a
granular modification HCg of the rhinal area limitans [Economo and Koskinas, 1925, p. 758].
Layer V thus borders directly on layer III.
There is a compact cellular middle zone of pyramidal cells, occasionally delimited from layer III
above by a paucicellular superior strip, and from
VI below by a similar inferior strip (cf. the lamina
dissecans described earlier in area HA). These
strips are not markedly clear in a consistent fashion.
Layer VI is very thin and contains cells obviously smaller than layer V. They are mostly triangular and irregular spindle (fusiform) cells, with
some polygonal forms among them. The sublayer
VIb is very poor in cells and thin, such that the
demarcation from the white matter is quite distinct here as well.
Cytoarchitectonic Modification HCD

As already mentioned, at loci where layers III, V
and VI are opulent in cells, the cortex of the rhinal area limitans HC appears robust and sufficiently thick. However, at loci where these three
layers become destitute, either through a uniform rarefaction or by the occurrence of acellular
patches, as is the case in its dorsal boundary (presubicular-rhinal granulous transition HCD), the
cortex becomes extremely frail; one may then
distinguish in an obvious way only the dense layer II, while the deeper paucicellular layers become indiscernible. As I pointed out earlier, considerable differences exist among individuals re-
163
Fig. 59. Seven sections (AG) through the uncus and hippocampal gyrus (cf. also fig. 57). B.G. = Band of Giacomini;
f.d. = fascia dentata; fi. = fimbria; G.h. = hippocampal gyrus; g.sl. = semilunar gyrus; NA = nucleus of the amygdala;
s.h. = hippocampal sulcus; s.pf. = substantia perforata; U = uncus; V = inferior ventricular horn; HA HF = areas of hippocampal gyrus; TJ, TG and TH = neighboring areas of temporal lobe; TJ H = hippocampotemporal piriform transition
area (on semilunar gyrus).
164
garding the cytoarchitectonics of this cortical

area [Economo, 1927c, d, 1928e].
Area HC occupies a large segment of the dome
of the hippocampal gyrus (fig. 1b) and only
reaches at its rostral extreme the superior brink
on its dorsal wall; caudally, this area does not
usually reach quite as far as the dorsal brink, but
it terminates just beneath it, as the brink itself is
occupied by the presubiculum and the koniocortex of the presubicular area granulosa HD described next (cf. schematic section drawings in
fig. 59CG and 60af). In some cases, the presubiculum and its area HD may even extend, in the
middle of the hippocampal gyrus, to the lower
half of the dome, and repel area HC ventrally.
At the isthmus, area HC (or rather its transition form HCD) seems to glide between the koniocortices of the retrosplenial area granulosa LE
and the presubicular area granulosa HD in such a
way that the rest of the koniocortex on the caudal
extreme of area HD occupies the dome, area HC
the ventral lip, and area LE the ventral wall of the
isthmus (fig. 51). However, as area HC here assumes the character of area HCD , i.e. the transition cytoarchitectonic structure already defined,
its demarcation from area HD is not easy and
somewhat subjective (cf. also plates P102b, P103d
and P104a, d in the Atlas [Economo and Koskinas, 2008]).
Presubicular Area Granulosa HD
The dorsal brink of the hippocampal gyrus is occupied in its entirety, from the (inner) concavity
of the uncus all the way to the isthmus and the
retrosplenial region, by a band of koniocortex, the
presubicular area granulosa HD , which from here
may stretch on the wall and sometimes the dome
of the hippocampal gyrus (fig. 1b, 59, 60; cf. also
plates P105, P106 and P109 in the Atlas [Economo
and Koskinas, 2008]). This area also becomes
pointed at its anterior and posterior extremes; its
maximum thickness corresponds with the mid-
point of the hippocampal gyrus. In its extent it

covers the entire superior part of the presubiculum (cf. areas HD1, HD2 and HD3 in fig. 5154).
Layer I is exceedingly thick (table 12); its superficial tier contains plentiful myelinated fibers.
Layer II+III reflects the combined layer II and
its direct continuation, layer III, featuring a granulization of its cells. The layer II component is
about 0.5 mm thick and consists of large ovoid
and round granule cells, densely packed. This is
the most distinctive and weighty layer of area HD
(best seen in fig. 52), assuming the aspect of a koniocortex; it can be traced all the way to the
boundary with the rhinal area limitans HC.
Layer V+VI is interposed between layer II+III
and the white matter of the gyrus; it contains
small pyramidal, triangular and spindle-shaped
(fusiform) cells in a characteristic paucity and indistinct demarcation vis--vis the white matter.
This layer is a combination of layers V and VI,
their individual attributes in terms of cellular
density and shape being virtually indiscernible.
A paucicellular band may be observed between layers II+III and V+VI, which is a continuation of the lamina dissecans of areas HC (fig. 52)
and HB . Towards the depths of the wall of the
hippocampal sulcus, layer II+III becomes gradually thinner, and the lamina dissecans appears to
rise toward the surface until it comes to lie immediately beneath layer I, at the point where layer
II+III terminates altogether (at the boundary
with area HE).
Cytoarchitectonic Modifications HD1, 2, 3

In an overview of the presubicular area granulosa
HD , the cytoarchitectonic feature that strikes
most is a predominance of the intensely staining
superficial granulous layer (II+III), ensuing from
its increased cellularity; the cellular layer V+VI
beneath is poorer in cells and thus less intense.
And yet, area HD is not uniformly structured
165
166
through its entire dorsoventral extent in the presubiculum; a tripartite division into three modifications can be traced all the way to its anterior
and posterior pointed extremes (fig. 51, 54; cf.
also plates P102bd, P104a, c, d, P107, P108 and
2008]):
The presubicular area granulosa limitans HD1
(area EK 100) is spread over the dome (fig. 52), at
the boundary with the rhinal area limitans HC.
The granulous layer II+III is actually thicker than
deeper down in the hippocampal sulcus, and also
less cellular; cells toward the immediate vicinity
of area HC are not yet completely transformed to
ovoid granule cells, the deeper zone (III) mostly
comprising triangular cells; further, layer V+VI is
still denser and somewhat thicker at that boundary.
The middle presubicular area granulosa HD2
(area EK 101) is an intermediate territory, lying
more dorsally; here, the granulous layer II+III diminishes in thickness, becoming at the same
time denser; in this modification all cellular elements have now become ovoid granule cells. On
the other hand, the deeper layer V+VI gradually
becomes excessively clear, the result of a cellular
rarefaction to such a degree that the layer may be
totally absent at places.
Fig. 60ag. Seven sections through the hippocampal

gyrus, isthmus, and retrosplenial region (cf. also fig. 57
and 58). A = Alveus; C.c. = corpus callosum; f.d. = fascia
dentata; fi. = fimbria; g.ar. = gyri of Anders Retzius; g.f. =
fasciolar gyrus; g.l. = limbic gyrus; g.rl. = retrolimbic gyrus; i. = indusium griseum; s.c. = sulcus of corpus callosum; Tr. = trunk of calcarine sulcus; V = inferior ventricular horn; x = small pads of Retzius gyrus; xx = beginning
of corpus callosum; HC HF = hippocampal gyrus areas;
TH and PH = neighboring temporal and parietal lobe
areas; OA and OB = occipital lobe areas; LB LF = superior
limbic gyrus areas.
The glomerular presubicular area granulosa

HD3 (area EK 102) is the last modification, already noted in the dorsal wall of the hippocampal
gyrus at the boundary with the subiculum, toward which layer II+III becomes even thinner,
but without losing its cellular density; on the contrary, its cells are grouped together into two to
three successive, spherical clusters or glomeruli
having a diameter of 0.30.5 mm. At the same
time, as one moves toward the subiculum, single,
extremely large, lancet-shaped pyramidal cells,
35/15 m in size (H/W), gradually make their appearance in the paucicellular layer V+VI with a
particularly long, slightly bulbous, and readily
visible vertical dendrite, issued from the cephalic pole of the cell and oriented toward the cortical surface. These cells, which seem to herald the
typical, large pyramidal cells of Ammons horn
(detailed next), come closer together and increase
in number as one approaches the subiculum.
As already explained, this area ends caudally
at the isthmus, in whose internal wall it can still
be recognized towards the back as an absolutely
thin field, where it directly borders on the koniocortex of the retrosplenial area granulosa LE
(fig. 1b, 5154, 59BG, 60af).
In figures 51 and 52 one can discern that the
granulous layer II+III is thicker and denser in
area HD compared to the more rostral sections of
figures 53 and 54. Thus, one might distinguish as
well, in accordance with Rose [1926b], a pars
anterioris HDa and a pars posterioris HDp. However, each of these divisions would still comprise
the secondary modifications HD1, 2, 3 [Economo,
1927d, 1928e, 1929d].
Pyramidal Area HE
Under the presubiculum, in the dorsal wall of the

hippocampal gyrus, the entire subiculum appears to be occupied by a single thick layer of
cells, lying immediately beneath a particularly
thick molecular layer (fig. 5153). We call this
167
Fig. 61. Four sections () through the limbic (cingulate) gyrus. LC LF = Areas in the posterior part; LA LB = areas
in the anterior part.
formation pyramidal area HE after the large pyramidal cells that compose it.
With only minimal structural variations, it
occupies the inferior part of the presubiculum
and the entire subiculum as variant area HE1, the
Ammons horn as modification area HE2, and finally the upturned cortical margin of the dentate
gyrus, on which the fascia dentata (f.d.) is
clapped. Through this entire extent, the pyramidal area only comprises a molecular and a pyramidal layer.
168
Layer I is exceptionally thick, reaching 0.6

0.8 mm at its thickest points; it comprises, in its
upper tier, a thick matrix of myelinated fibers; its
intermediate zone is also traversed by numerous
myelinated fibers, a feature distinguishable in
Nissl-stained specimens from the multitude of
satellite glia nuclei.
Layer V+VI beneath contains large pyramidal
cells, 3580/20 m in size (H/W), superimposed
in multiple tiers within a single pyramidal layer,
and disposed perpendicularly to the cortical sur-
face; their number is estimated at 12,00020,000

cells/mm3. The thickness of this pyramidal layer
differs from place to place; directly after the passage from the presubiculum to subiculum (modification area HE1), the thickness is about 1.5
mm or more; in the subiculum proper (modification area HE1), it becomes reduced to 0.81.0
mm; finally, in Ammons horn (variant area HE2),
this pyramidal layer forms an even denser and
darker, but relatively thinner band, 0.4 mm in
thickness. The upper border of this pyramidal
layer vis--vis the molecular layer is vague and
largely festooned (fig. 52). The lower border
against the white matter, however, is rather sharp
(linear). In the immediate vicinity of the white
matter, one notes a sublayer of stellate or spindleshaped (fusiform) cells, small, flat, and horizontally disposed; they form an absolutely thin margin and they are not visible under the low magnification power of figures 5154.
Subicular Pyramidal Area HE1

The pyramidal layer, with the secondary sublayer
just mentioned, lies in an extrapolated extension
projected from layer V+VI of the presubicular
area granulosa HD. However, a true identification of these two layers might be erroneous, like
it would also be with the deeper layers of isocortex (e.g. of areas TG and TH [Economo, 1928e]),
since the layers of area HE1 are quite different in
structure from those of area HD ; as already pointed out in the introductory chapter, the allocortex
develops from a layered anlage fundamentally
different from that of the isocortex. The pyramidal layer of the subiculum bears no true relation
at all to layers V and VI of the isocortex, since
even in the presubiculum (area HD), the deeper
cellular layers which we number V and VI only
for the sake of practicality are very different
from the similarly numbered isocortical layers.
Moreover, these two cellular layers are so scanty
in the modification areas HD2 and HD3 that they
seem to virtually disappear; consequently, the

continuity between the pyramidal cells of layer
V+VI of the modification area HD1 and those of
the ground area HE is intermittent. For further
details, see our larger work [Economo and Koskinas, 1925, pp. 767771].
We term this variant of the pyramidal area,
which vests the subiculum, subicular pyramidal
area HE1 (cf. plates P102d, P104c and P106P112
in the Atlas [Economo and Koskinas, 2008]). It
comprises two further modifications:
The glomerular subicular pyramidal area HE1
(area EK 103) occupies the upper segment of area
HE1, which still constitutes in actuality the most
inferior part of the presubiculum. That dorsalmost segment, which borders on the glomerular
presubicular area granulosa HD3 (area EK 102),
also shows a tendency of pyramidal cells, at the
border between the molecular and pyramidal
layers, to cluster into glomeruli, among which a
few area HD3 glomeruli may be scattered, containing small granule cells. The border between
the molecular and pyramidal layers displays
some isolated deep indentations, which reach
deeply into the pyramidal layer.
The pyramidal subicular area simplex HE1
(area EK 104) is found within the subiculum proper, where such a tendency dwindles, although the
border between the molecular and pyramidal layers remains wiggly, with occasional deep indentations (fig. 53). The pyramidal layer in this modification is considerably thinner at 0.8 mm.
Pyramidal Area of Ammons Horn HE2

The adjoining cytoarchitectonic structure is the
pyramidal area of Ammons horn HE2 (area EK
105; cf. plates P103a, P104b, P106P109 and P112
in the Atlas [Economo and Koskinas, 2008]),
which swings over to the dentate gyrus in an arch
and exhibits the same structure as already noted,
i.e. a thick molecular and a thinner pyramidal
layer; beneath the latter, one only finds the white
169
matter of the alveus (A in fig. 52), protruding into

the ventricle.
Layer I in Ammons horn is known to be extremely thick (fig. 52); it comprises a superficial
and an intermediate myelinated strip (fig. 53),
which are not readily visible in Nissl-stained
preparations, but can be deduced from the presence of an increased number of glial cells. Subjacent to that intermediate strip, and just above the
pyramidal layer, one finds the stratum radiatum,
a particularly dense and somewhat homogeneous,
vertically-striated structure (visible as a pale band
just above the pyramidal layer in fig. 52).
Layer V+VI follows with the pyramidal layer
(stratum cellulare) and a stratum oriens, thinner
than the stratum radiatum above, but also compact and apparently homogeneous, interspersed
just above the dense myelinated fibers of the alveus (fig. 52). These three structures (stratum radiatum, stratum cellulare, and stratum oriens)
course further into the subicular region described
above, but not very distinctly. The pyramidal layer of Ammons horn HE2, with about 35,000 cells/
mm3, mostly 40/20 m in size (H/W), is even
denser in cells than the subicular pyramidal
modifications HE1, such that in somewhat thicker sections one hardly discerns any intracellular
space. Moreover, the cells are all disposed perpendicularly towards the surface (i.e. radially),
having the appearance of the spokes of a wheel; as
at this point the Ammons horn makes the curve
to the dentate gyrus, the tips of the pyramidal
cells all converge toward the extrapolated center
of the dentate gyrus. The cell nucleus is strikingly large, vesicular, and contains a conspicuous,
deeply staining and shimmering nucleolus.
gyrus; all this along the entire caudorostral extent of the hippocampal gyrus, all the way to its
anterior bend that leads to the uncus (fig. 1b).
We have seen how entorhinal areas HA and
HB vest the entire anterior frontal part of the uncus bulge, whereas areas HC and HD terminate at
a point that corresponds to the hollow surface of
the uncinate knee. The hook-like part of the uncus, folded toward the interior of the hemispheres
and backwards, is vested in the entirety of its inner concave side by the subicular pyramidal area
HE1 (fig. 1b).
Layer I (the molecular layer) in the cortex of
the posterior crus of the uncus is extremely thick,
reaching 1.5 mm (fig. 53, 54).
Layer V+VI (the pyramidal layer) beneath it,
is equally thick; all of its exceedingly large pyramidal cells are disposed exactly perpendicularly,
with their tip pointing toward the cortical surface, just like one observes it in the cortex of Ammons horn. However, since this area follows a
convex plane, unlike the concave surface of Ammons horn, pyramidal cells here do not give the
impression of the radial spokes of a wheel, but
rather assume a fan-like form. On the other hand,
since the superficial faade of the uncus is frequently furled into small protrusions (digitate
gyri), one observes images analogous to those
shown in figure 53 and 54 at the mark HE3.
We have termed this variant of ground area
HE the pyramidal area of the uncus HE3 (area EK
106; cf. plates P103b, P105 and P106 in the Atlas
[Economo and Koskinas, 2008]). The rather complicated relations, shifts, reciprocal confines, and
backfolding of the various areas at the level of the
uncus pole can best be made out in the schematic
section series of figure 59AG.
Pyramidal Area of the Uncus HE3

Band of Giacomini and Pyramidal Areas HE4, 5
The pyramidal area HE , with its structural modifications outlined above, vests the lower part of
the presubiculum, the entire subiculum, Ammons horn, and also forms the core of the dentate
170
Based on gross anatomy, one generally assumes

that the fascia dentata continues at its crossing
over the uncus as the band of Giacomini [1883].
Yet, our cytoarchitectonic studies demonstrate

that such a concept is incomplete [Economo,
1925b].
When the dentate gyrus and the Ammons
horn reach the folded region of the uncus at the
anterior extreme of the hippocampal gyrus, the
entire dentate gyrus, as it uprolls its spiral fold,
together with the fascia dentata, tuck up over the
fimbrial part of the uncus (already vested by the
pyramidal area) to form a cap-shaped shield. The
brim of this hat-like formation constitutes the
fascia dentata; with its particular cytoarchitectonic structure, which consists of dual molecular
and granular layers and an interposed pyramidal
layer, the fascia dentata directly traverses the uncus as the band of Giacomini, from inside towards the outside (fig. 53). In a schematic drawing of a cross-section from this region (fig. 59D)
one may discern the pyramidal area of the uncus
HE3 (area EK 106) and, over it, the arch-shaped
granular and molecular layers of the fascia dentata, i.e. the band of Giacomini (B.G.), with the
pyramidal layer compressed between its two
granular sheets (cf. area HF in fig. 53 and B.G. in
fig. 59D).
In somewhat more caudal sections, one sees
the pyramidal area of the digitate gyrus of the uncus HE3 (area EK 106) and a further cortical layer,
which proceeds from the pyramidal area of Ammons horn HE2 (area EK 105) and forms a shield
over the posterior extreme of the uncus (fig. 59E ,
F). We have characterized that upturned shield
by sequentially terming its two territories quaternary (HE4) and quinary (HE5) pyramidal areas
(fig. 1b; table 2). Such is the composition of the
anterior segment of the pyramidal area.
In its caudal extreme toward the isthmus, the
presubiculum (area HD), the subiculum (area
HE1), Ammons horn (area HE2) and the dentate
gyrus (area HF ) narrow more and more, until, at
the isthmus itself, they occupy a territory covering at the most 2.5 mm in the depths of the internal wall of the limbic gyrus (cf. fig. 51 and 50 and
schematic drawings of fig. 60ag).
At this point, the presubicular area granulosa

HD has completely disappeared; however, the pyramidal area HE continues further caudally, as
already mentioned in the preceding chapter (cf.
the schematic cross-section drawings of fig. 60a
g and 58b), where it rolls over into the posterior
ultracingulate area LF1 (area EK 47), i.e. into the
magnocellular transition of the limbic gyrus that
folds on the dorsum of the corpus callosum. The
dentate gyrus, or better said, the dentate area HF,
gradually passes into the fasciola cinerea or fasciolar gyrus and finally into the ultracingulate
area obtecta LF2 (area EK 48).
In the schematic drawings of figure 60cf, one
can clearly discern that gradual passage of the
structures of the dentate gyrus and the fascia
dentata (f.d.) into the fasciolar gyrus (g.f.). This
figure is hatched similarly to figure 58, in order
to better demonstrate the areas covered by structures such as the intralimbic gyri of Anders
Retzius [Retzius, 1898] and the protrusions of the
subicular wall and dentate gyrus.
Dentate Area HF
At the point where the pyramidal area spirally

rolls up against the subiculum, i.e. the terminal
margin of the cortex, its termination thickens
like a knob; this comes about as the cortical margin rolls up to the uncus from outside inwards; it
then makes a hook-like turn from the inside outwards (HF in fig. 52 and schematic drawings of
fig. 59G and 60a). The thickened and upturned
knob-shaped structure thus formed remains always cloaked in the fold constituting the fascia
dentata (f.d.); it thus becomes evident that in
cross-section, the fascia dentata assumes the aspect of a horseshoe, which harbors the aforementioned knob of the cortical margin within its
concavity (fig. 51, 52).
The fascia dentata actually consists of two molecular layers, between which there is a dense layer of coarse granule cells (table 12). Of the two
171
molecular layers, the external is rather thick (0.5

mm) and adheres to the subiculum, fusing with
the molecular layer of the latter. The internal molecular layer, which is intimately and inseparably
fused with the molecular layer of the upturned
extreme of the pyramidal area, is extremely thin
(hardly reaching 0.2 mm in thickness); at certain
levels, it is pervaded by cells derived from the pyramidal layer to such a degree, that it becomes
difficult to recognize its molecular layer nature.
We termed this cortical margin, i.e. the cortex
of the dentate gyrus, whose internal segment is
formed by the pyramidal layer and whose external
aspect is covered by the fascia dentata, the dentate
area HF (area EK 107; cf. plates P103a, P104b,
P106P109 and P112 in the Atlas [Economo and
Koskinas, 2008]). It is bounded on the outside by
the white matter of the fimbria (fi.), which is covered superficially by a very fine layer of cell-poor
gray matter, and which rolls into the lamina affixa
to shut the ventricle off (marked V in fig. 52).
The dentate area HF contains coarse granule
cells and further lacks all the other attributes that
would allow us to consider it a sensory cortical
zone [Economo, 1928e].
Function
With regard to the physiologic function of the

hippocampal gyrus, it is more or less reasonable
to consider it as generally relating to the sense of
172
taste. Thus, the most obvious thing would probably be to regard the koniocortex of the presubicular area granulosa HD as the primary sensory gustatory domain; the subiculum, Ammons
horn and dentate gyrus, where the fimbria originates, could then perhaps function as efferent
centers at the disposal of the sense of taste, specifically having a secretory or even a vasomotor
role. Nevertheless, definite proof is missing in
order to really attribute such a primary gustatory
function to area HD. Although the fascia dentata
is largely composed of granule cells (vide supra),
it cannot be considered to be koniocortex, because firstly, its granule cells are too coarse, and
secondly, it lacks some other important attributes of the sensory cortex (e.g. a nerve terminal
plexus).
The anterior pole of the uncus, with its entorhinal region and the uncinate and parauncinate
areas HA and HB , is directly related to the external ramus of the olfactory root, whose myelinated fibers plunge into its molecular layer; thus,
that segment of the uncus does not in all likelihood belong to the organs of taste, but to those
subserving olfaction [Economo and Koskinas,
1925, pp. 788790]. But it is curious to note to this
end that, according to Ramn y Cajal [19001906,
1923], entorhinal areas are equally well and perhaps better developed in microsmatic animals
(humans and monkeys) than in macrosmatic animals [Economo, 1928e].
Conclusion
The Future of Cytoarchitectonics7
In the preceding chapters I covered the main areas of the human cerebrum and briefly surveyed
the cellular structure of its lobes, as an orientation to the field of cytoarchitectonics. Within the
scope of such a restricted review, it is impossible
to extensively discuss all the details. Anyone interested in the special questions raised in this
book will find them more exhaustively treated in
the larger work repeatedly cited [Economo and
Koskinas, 1925], as well as in the Atlas with its
numerous microphotographs [Economo and
Koskinas, 2008]. That material can serve as the
basis for individual and original study.
This chapter only appeared in the 1929 English edition

[Economo, 1929d] without any iconography. It is reproduced
here by permission from Oxford University Press. It treats cytoarchitectonic neuropathology, as well as the evolutionary concept of progressive cerebration that Economo [1928b] introduced after the publication of Zellaufbau, developing his bold
propositions on brain phylogeny and the neurobiology of the
gifted and talented (lite brains). He gave a related presentation on July 2, 1928, to the International Neurological Reunion
at La Salptrire in Paris in French [Economo, 1928c], and published an extended review the following year in German
[Economo, 1929a]. He made further presentations on December 4, 1929, on the occasion of the dedication of the Psychiatric
Institute and Hospital of New York in English [Economo, 1930e],
and gave the festive address to the Viennese Medical Association on March 20, 1931, in German [Economo, 1931e], with an
article concurrently published in Spanish [Economo, 1931f].
The present chapter is supplemented with illustrations (fig. 63
72) from Economos original articles.
Nevertheless, even a brief survey suffices to underscore the importance of studying the cerebral
cortex. The view of Meynert that the cortex consists of many different areas and must therefore be
regarded as constituting multiple nervous organs,
the varying structure of which forms the basis for
recognizing and differentiating their diverse functions, has been corroborated by our identification
of the sensory cortex, which can be recognized by
its cytoarchitectonic structure alone, as can the efferent cortex as well. It is certain that soon will follow the recognition of the importance of other
structural types, perhaps, e.g. those mediating
memory and higher mental functions.
Much is also expected from comparative anatomic studies in the field. The poor or robust development of some of those organs in the brains
of different animal species is of substantial importance in comparative physiology and psychology.
In the province of macroscopic anatomy, cytoarchitectonic studies form the basis for a new
and more accurate mapping of lobe and region
boundaries. I already had an opportunity, in the
preceding chapters, to define boundaries for the
occipital, temporal, parietal, and partly also for
the olfactory lobes, which differ from the boundaries hitherto laid down by gross anatomy
(fig. 62); contrary to the boundaries assigned for
these cerebral regions by gross anatomy, the de-
Fig. 62. A patented plaster model of the human brain, with the various cytoarchitectonic areas marked according to
the original Economo and Koskinas [1925, 2008] lettering system, and indicated by Economo with various colors on
the convexity and median facies of the cerebral hemispheres, was manufactured in the 1920s by Fabrikation Chirurgischer Instrumente Carl Reiner, Mariannengasse 17, Wien IX (still operating today in the same locality after four generations). Economo made a demonstration using such a model in his December 3, 1929 lecture at the Section on
Neurology of the New York Academy of Medicine [Economo, 1930f, g]. One additional model is on display at the Institut fr Geschichte der Medizin (Institute for the History of Medicine) of the University of Vienna, next to Economos
death mask. The numbers in the upper left frame (mid-sagittal hemispheric view) denote the five structural types of
isocortex (cf. fig. 8 and 9 in the introductory chapter). Photos courtesy of Nikolaus Reiner, Manufaktur Chirurgischer
Instrumente Carl Reiner GmbH, Vienna, Austria.
limitations upon which the cytoarchitectonic division of these parts of the brain rests are constant and characteristic structural details, which
can be corroborated at any time through the use
of the microscope.
One of the most interesting fields of research,
however, will be the study of individual normal
174
and pathologic differences in human cytoarchitectonics. Even now there are many problems attracting attention: the relations of areas to diverse
types of gyri; differences between the two cerebral hemispheres (fig. 63, 64), among various races and ages; the accurate study of the brains of the
mentally gifted, of exceptional individuals, of
Fig. 63. Comparison of the lateral (Sylvian) fissure between right and left hemisphere in six human brains (Gh.). The
small Arabic numerals at the small adjacent domes along the lateral fissure are for orientation and correspond to the
same numbers shown in figure 64 of the superior temporal fields in the respective brains. Originally figure 3 in the
study of Economo and Horn [1930] on the temporal lobe.
Conclusion
175
176
Fig. 64. Right-left hemispheric comparisons of the superior temporal fields in the six brains (Gh.) shown in figure 63.
The small Arabic numerals correspond to the small adjacent domes, along the lateral (Sylvian) fissure, depicted in
figure 63. Originally figures 49 in Economo and Horn [1930]. Individual cytoarchitectonic areas are listed in table 2
(Introduction).
Conclusion
177
mental defects (the brain in hereditary familial

amaurotic idiocy, congenital deafness and mutedness, mental retardation); the study of pathologic brains and those of psychotic patients.
Cellular, Laminar, Structural and Areal

Neuropathologic Changes
In applying our present knowledge of cytoarchitectonics to neuropathology, we must remember

the following principles. The three main types of
cells of the cortex (pyramidal, granule, and fusiform) are arranged in horizontal layers. The cortex consists of six such layers, each of which has
probably a certain function in the physiologic activity of the gray matter. On the basis of the relative development of each of these layers and their
different importance, we have been able to differentiate five types of cortical structure (cf. Introduction and fig. 8). These five structural types
are further subdivided into the different modification areas, which number more than 100 (tables 1, 2). The question is whether, in studying
pathologic brains, one may find certain diseases
that correspond to definite alterations of some of
these structural elements in short, whether one
might expect to find cellular, laminar, structural
(typical) or areal changes.
At the outset, however, one would have to exclude types of diseases that depend upon vascular
changes, because in those cases, the neuropathologic changes of the cerebral parenchyma depend
on the course of the blood vessels.
It has long been known that amyotrophic lateral sclerosis especially affects the giant cells of
Betz at an early stage, i.e. that this disease primarily strikes at the giant pyramidal precentral
area FA (area EK 2). Aside from the cells of Betz,
however, amyotrophic lateral sclerosis also affects the large pyramidal cells of layers III and V,
and possibly also cells of the other layers, in its
progressive course. Furthermore, the disease
passes beyond the boundaries of the precentral
178
area, especially in a rostral direction, where it

particularly affects the posterior parts of the
three frontal gyri, insofar as the latter are agranular (agranular frontal area FB or area EK 4). In
this case, then, we can say that a large part of cortical structural type 1 has been affected. On the
other hand, the studies of van Bogaert [1925],
Bertrand and van Bogaert [1925] and others have
shown that this disease does not necessarily stop
at this boundary: it may even reach over into
parts of the cortex behind the central sulcus of
Rolando, especially affecting layers III and V in
such regions. One can thus say that amyotrophic
lateral sclerosis is in reality a disease that primarily shows an areal spread. Further, I must state
that in this disease there is a certain susceptibility of layers III and V in many other brain regions. Such a vulnerability to a certain disease is
called pathoclisis by Vogt [1925].
Thus far we are not acquainted with any other
disease of the nervous system that shows an areal
circumscription similar to amyotrophic lateral
sclerosis in the agranular precentral frontal lobe
(extended motor region). The majority of organic
mental diseases, as Alfons Jakob [1920, 1921,
1923, 1927a, b] has shown, rather manifest with a
widely spread involvement of the cortex, although, it must be stated, with a certain predilection for certain regions. Thus, for example, in
general paralysis, Jakob finds that changes are
primarily localized in the anterior frontal lobe, in
some districts of the parietal, in the middle (T2)
and inferior (T3) temporal gyri on the temporal
pole, and also in the insula, the anterior half of
the cingulate gyrus, and especially in Ammons
horn. The agranular precentral region, the posterior segments of the inferior frontal gyrus (F3),
the postcentral gyrus, the entire occipital region,
and the superior temporal gyrus (T1) along with
the transverse gyrus of Heschl, usually show no
changes.
A typical case of senile dementia reveals a localization to, and an avoidance of, regions identical with those affected in general paralysis. One
might include in this type of distribution such

diseases which avoid the cortical structural types
1 and 5, especially affect cortical type 2, and only
partly involve changes of cortical types 3 and 4,
and might further presume a selective pathoclisis
of certain cortical structural types in these diseases. But here I must stop to point out that, although the cortical structural type 1 has been
mentioned among those spared by these diseases,
the anterior part of the cingulate gyrus, which is
affected, actually belongs just to this type 1. Progressive paralysis and senile dementia thus find
no true areal, and no absolute structural, dissemination in the cortex, their selectivity being only
a relative predilection of the disease for the cortical structural type 2. Jakob [1920, 1921, 1923,
1927a, b] has especially pointed out that spared
regions are precisely those which are the richest
in myelinated fibers; thus, one might assume that
such regions acquire their special resistance to
the diseases mentioned as a consequence to the
wealth of myelinated fibers. Even in cases of atypical (focal) general paralysis of Lissauer and atypical senile dementia with focal syndromes (Alzheimer disease), there is no apparent localization
of the process to any definite area, but merely a
displacement of the pathologic process posteriorly, towards the parietal and temporal lobes. At
the same time, the regions mentioned above as
being free from involvement are also spared in
these diseases.
Pick lobar atrophy also shows no restriction to
particular areas, but is distributed over the middle, inferior and fusiform temporal gyri (T2T4),
sparing the superior temporal gyrus (T1). In the
frontal lobe only the anterior one-half of the three
frontal gyri, the insula and the cingulate gyrus
are affected. In contrast to general paralysis and
the senile dementias, the presubiculum and the
subiculum are affected in this disease, whereas
Ammons horn is spared. Since Pick disease leaves
the inferior frontal gyrus (F3), the superior temporal gyrus (T1) and the lower parietal lobe free,
it is inaccurate for Gans [1922] to call these con-
Conclusion
ditions a disease of specifically human (i.e. recently acquired) cerebral structures, since F3 and
T1 are by themselves specifically human.
Cytoarchitectonic Neuropathology of the

Psychoses
The three organic psychoses mentioned here involve all six cortical layers, but they especially affect layer III; this is also the layer particularly affected in schizophrenia (dementia praecox) and
chronic alcoholism. Thus, the involvement of layer III probably occurs in all organic and functional psychoses characterized by dementia and a
disintegration of the personality. (It is as yet impossible to state to what extent individual symptoms, such as for instance Korsakoff syndrome,
hallucinations, etc., might or might not be due to
anatomic changes in layer III in specific cytoarchitectonic areas.)
In contrast with these rather purely mental
diseases involving primary changes in layer III,
Jakob [1920, 1921, 1923, 1927a, b] studied progressive degenerative brain disorders with motor
symptoms. He collectively grouped those under
the name of spastic pseudosclerosis and found
that, apart from the corpus striatum, the agranular precentral frontal region (type 1) was the gray
matter especially attacked; secondarily, the rest
of the frontal and also the temporal lobe were affected, layers V and VI, i.e. the efferent layers, being markedly involved in those diseases with a
characteristically motor symptomatology. He
further found that, in cases of senile dementia
that show a striking extrapyramidal muscular rigidity in addition to their mental signs, there was
also a marked involvement of layers V and VI. In
the same way, he found that in cases of chronic
lethargic encephalitis in which, aside from mental signs, the capacity for expression rather than
the essence of the personality had been destroyed,
there was an especial destruction of layers V and
VI. The same is true of Huntington chorea.
179
All these circumstances point to a systematic

relation between the agranular precentral regions
and layers V and VI with the extrapyramidal centers.
In summary, I can say, in light of our present
knowledge of the cytoarchitectonics of brain diseases, that, apart from amyotrophic lateral sclerosis, there are no general area constraints for neuropathologic processes, but that there is rather a
regional selectivity for certain structural types of
cortex. Furthermore, there is a certain layer affinity to disease, as a result of which we find layer III
markedly affected in psychotic personality disorders; on the other hand, diseases primarily affecting the capacity for expression and movement
show a particular affinity for the efferent layers V
and VI. More cannot be said at the moment.
The cytoarchitectonic changes in the cortex in
cases of severe general paralysis, which are caused
partly by the massive loss of nerve cells and partly by the hyperproliferation of glial cells and the
influx of infiltrating cells, are usually of such a
nature that they entirely blur and disturb the layer and area characteristics. Changes brought
about by other organic brain diseases hardly ever
reach such a degree of damage, but they, also, often disturb the picture considerably, as e.g. in
schizophrenia.
Mental Retardation
In the case of oligophrenia there has been no research carried out on a definitive cytoarchitectonic basis. The first studies in this province were
carried out by the Swedish researcher Karl Hammarberg [1895], who is also one of the founders of
cytoarchitectonics. His work remains of fundamental importance to this day. He considered oligophrenia less of a disease and pathologic change
of the cortex, and more of an arrest of development, which may manifest itself anatomically in
varying degrees of intensity in the cortex, i.e. either locally or diffusely, through a simple paucity
180
of cells, their small dimensions, incomplete cell

layers or even a definitely fetal level of development. In extreme cases, the brain may reveal only
a membrane forming the wall of the hemisphere,
e.g. in porencephaly. It also seems that a developmental inhibition of a purely localized nature has
a certain retarding effect on other parts of the
cortex that otherwise possess an inherent capacity for further development. The grades of mental
deficiency idiocy, imbecility and oligophrenia
differ anatomically only in the degree of such a
developmental inhibition.
Further cytoarchitectonic and myeloarchitectonic studies are thus likely to be attended with
rich fruits in the field of the psychoses as well as
in the defective conditions. Comparative anatomic studies of the cellular structure of the brain
of different animal species would be of the highest interest. Such investigations should give us a
basis for understanding the function of the cortex as a whole. At present, our knowledge on this
subject only warrants the following statements.
The Cerebrum as a Sensory Organ
Like higher sensory organs, the cerebrum is an

example of a paired development, due to the
evagination of vesicles from the tube-like anlage
of the encephalon; these protrusions occur at the
forebrain, just as the olfactory bulb protrudes
from the basal parts of that same forebrain, the
acoustic vesicles from the hindbrain, and the optic vesicles from the diencephalon. On the basis
of this type of development, we may consider the
cerebrum as an analog of the sensory organs, albeit a sensory organ that does not receive its sensations directly from the external world, but
rather from the lower parts of the central nervous
system, and is thus only indirectly connected
via the lower central nervous system with the
outside world. Its field of sensation is turned towards the inner world of the central nervous system itself.
The specific sensory energy of the cerebrum is

consciousness, an individual sensory mode, just
as the capacity for sensing light is the specific modality of the retina. That is the reason why it is
just as fruitless to try to explain consciousness on
the basis of any psychologic constellation of images, or in any other psychologic way, as it is
impossible to define or to explain the nature of
the sensation of light or any other of the higher
senses.
The function of the cerebrum has also a further similarity to that of sensory organs, especially the eye; we see that both voluntarily and
involuntarily, some cerebral process may often
be explained or described upon the analogy
with such an organ, e.g. the analogy between
mental attention and the fixation to a point
with the eyes. In addition to the simple reception of the processes that have taken place in the
central nervous system, it is a further function
of the cerebrum to effect an engraphic accumulation and connection (association) of such processes, as well as an efferent discharge of motor
energy towards the external world, very similar
to the insertion of some further sensory processes (for instance the conscious perception of
movements and colors), and even of a motor apparatus (the lens and the iris) into the sensory
organ of our eye.
Consciousness as the Specific Sensory Energy

of the Cerebrum
The reception of sensory stimuli that have

touched upon centers in the lower nervous system takes place in the brain, i.e. in consciousness,
on the sensory surfaces. As I was able to show in
the preceding chapters, these surfaces are characterized by a more or less intensive granulous
transformation or granulization of cortical cells
(koniocortex). We have become acquainted with
five such koniocortical areas, corresponding to
the five primary senses. However, it is very prob-
Conclusion
able that the total extent of the so-called sensory

cortex is not exhausted in those five koniocortical areas (vision, hearing, taste, olfaction and
touch).
It is necessary to assume that other, also very
typical and specific sensations, which, however,
are not included in the above-mentioned group of
our five senses, must also be represented in the
cerebral cortex, e.g. the sensation of vertigo, orgasm, and perhaps other organic sensations such
as hunger, thirst, and even complex sensations
such as anxiety, pleasure, etc., which first become
combined in the deeper parts of the central nervous system belonging to the domain of affects,
instincts and impulses, and then enter consciousness as the entity of an emotion, and so forth.
These, very likely sensory, surfaces of the brain
are not known yet. The reason may lie in the fact
that they possibly overlap in their extent with
sensory areas already known; alternatively, they
may be localized elsewhere, and their specific
differentiation might be of such a kind that our
present state of knowledge makes it impossible to
recognize them as such.
The Central Sulcus Divides the Cerebral

Hemispheres into a Motor and a Sensory Part
Whatever the ultimate truth of such matters may

be, it is nevertheless a striking fact that each one
of the five sensory fields that I delimited falls behind the central sulcus of Rolando, onto one of
the caudal portions of the cerebrum. This recalls
a rule laid down years ago by the Russian anatomist Betz [1874, 1881]: the central sulcus of Rolando divides the cerebral hemispheres into an
anterior motor and a posterior sensory part, just
like the lateral sulci of the spinal cord divide its
gray matter into a ventral motor and a dorsal sensory half. Although such a statement is a little too
general, it is nevertheless true insofar as the motor parts of the cortex are actually in front of the
central sulcus of Rolando, and the sensory be-
181
Fig. 65. A section of the cerebral hemisphere through the two walls of the central sulcus of Rolando. The wall of the
precentral gyrus (C.a., Convolutio anterioris) is seen on the left side of the figure, and the wall of the postcentral gyrus
(C.p., Convolutio posterioris) on the right side. Even at such a low power, one readily discerns the greater cortical thickness and large cell size in the precentral gyrus, against the thin cortex, small cell size, and dense cellular packing in
the postcentral gyrus. Thus, we here have two isocortical regions in direct vicinity, but so differently structured, that
it is difficult to imagine that they form part of the same organ. Magnification !12.5. Originally figure 7 in Economo
[1929a].
hind it (fig. 65). It is furthermore striking that

such a functional difference is accompanied by
an anatomic difference in cellular structure between the anterior and posterior parts of the cerebrum. The entire cortex anterior to the central
sulcus of Rolando discloses a striking growth of
large pyramidal cells and the pyramidal cell layers, with a simultaneous poor development, and
frequently a complete absence, of both granular
layers. The caudal parts of the cortex, on the other hand, reveal a marked growth of the granular
layers, and a lesser development of pyramidal
cells (fig. 66, 67).
182
Between the three sensory domains that are

most important to humans touch, vision and
hearing there is a wide cortical field, which, as
we already learned, is largely occupied by cortical
structural type 3. It is very probable that each of
the boundaries of cortical type 3 regions that
touches upon the above sensory domains in the
parietal (postcentral gyrus), occipital and temporal lobes, subserves the conscious perception of
these higher senses, as well as its memory impressions.
Next to these areas comes a series of centers,
as we learned from neuropathology, that are su-
perimposed on a scale of complexity, and these

are very important for mental perception. Thus,
for example, in addition to the granulous sensory
surface of the gyrus of Heschl, which may be considered as the center of pure tone perception,
there first comes a field that subserves the comprehension of spoken words, and then one that
subserves the understanding of word meaning
(fig. 68). In the same way, there are centers located next to the other two sensory fields, cortical
areas whose role is to gradually form higher complexes out of corresponding specific cortical excitations.
The combinations of these complexes of
higher order of one sensory surface with the complexes of other sensory surfaces take place in
more eccentric areas. Functional neuropathology
teaches us that a lesion of the segment of the lower parietal lobe that lies halfway between the sensory domains induces disturbances such as stereoagnosia, sensory alexia, the higher forms of
sensory aphasia, acalculia, etc., i.e. the so-called
cognitive disturbances. Cognitive disturbances
are those in which there is no agreement or identification between the primary conscious perception of an object by one of our senses, and the past
experience of that object, i.e. its memory impression stored away as a result of its earlier concep-
Fig. 66ac. Schematic drawing of anatomic area demarcation at the operculum of Rolando, a region with many
individual variations. The sensory tactile domain PB ,
with its granulous koniocortex which occupies the largest part of the rear wall of the central sulcus of Rolando
(R) in its entire dorsoventral extent terminates somewhat anteriorly, i.e. dorsally from the shallow ventral extreme of the central sulcus, but still within it. On the other hand, the agranular motor area FA ends either concomitantly with the sulcus, or reaches across with only a
small segment, ventrally from the sulcus end, yet still
within the operculum. Abbreviations (other than cytoarchitectonic area codes): s.po.i, inferior postcentral sulcus;
s.pr.i., inferior precentral sulcus; s.sc.a., anterior subcentral sulcus; s.sc.p., posterior subcentral sulcus. Originally
figure 1 in Economo [1930d].
Conclusion
183
Fig. 67. a Semi-schematic illustration of gross anatomic relations, with the ventral surface of the operculum shown
as well. b The extent of individual areas and their boundaries, marked by dotted lines, as determined from serial sections. The central sulcus of Rolando (R) includes its continuation on the ventral surface of the operculum and an
imaginary prolongation in the same direction further medially, representing a sharp boundary between the parietal
and the frontal structures. Abbreviations (other than cytoarchitectonic area codes): C.a., precentral gyrus; C.p., postcentral gyrus; Sylv.F.Ob., surface area of the lateral (Sylvian) fissure; od. (= oder), or. Originally figure 2 in Economo
[1930d].
tion complex by multiple senses. Lesions of the

caudal portions of this territory comprising the
angular gyrus and the basal segment of the inferior parietal lobule provoke disturbances of an
even higher order that give the impression of
mental and intellectual defects. Head [1926] has
called such defects asemantic, meaning that what
suffers in such cases is the understanding of the
importance of things of the external world; and
thus, the contact with the suitable, purposeful actions of conduct also suffers as a result. Therefore, in such cases there is a disturbance of the
receptive-conscious part of the mental personality. Thus, the notion exists of an intellectual defect
of a sensory-receptive nature.
184
Two Foci of Mental Activity, the Anterior in

the Middle Frontal Gyrus (F2) and the
Posterior in the Middle Parietal Gyrus (P2)
In the anterior parts of the cerebrum we also find

a superimposition of centers. In those parts of the
cortex directly in front of the central sulcus of
Rolando, i.e. on the precentral gyrus, we find a
zone primarily excitable by electricity, which
leads to individual limb movements (fig. 69): the
precentral area FA (area EK 1) or pure motor cortex. Then, in front of this area, there is a series of
motor centers for increasingly higher orders of
activity: first the center in the agranular frontal
area FB (area EK 4) for locomotor complexes,
Fig. 68. Schematic drawing of the superior temporal plane. a Designation of sulci and gyri. b Cytoarchitectonic areas.
c The neighboring influence of the gyrus of Heschl (first transverse gyrus or gyrus temporalis transversus magnus)
from the insular, temporal and parietal structures in the vicinity, and the organization of the gyrus of Heschl into
various regions in the mediolateral () and anteroposterior (aabb) direction. d Schematic organization of areas TC and TD of the gyrus of Heschl into a pars granulosa (areas TC1 and TD1) and a pars simplex (areas TC2 and TD2)
each. Originally figure 1 in Economo and Horn [1930]. Individual cytoarchitectonic areas are listed in table 2 (Introduction).
such as for walking upright, in the superior frontal gyrus (F1); then the centers for writing movements ventrally to that. Further ventrally, there
are the centers for the acts of mastication and
swallowing, as well as for phonation (area FB).
Rostrally, in the region of the intermediate frontal
area FC (area EK 6), the inferior frontal gyrus (F3)
carries the centers for motor speech (Brocas area
FCBm or area EK 8), and dorsal to this field there
are the centers for the combined trunk, head, and
ocular presentations and postures, especially the
so-called spying movements of the eyes, i.e. presentations of attention, in area FC. As with speech,
such movements involve many mental components, and it is remarkable that such a fact is expressed in the anatomic picture of corresponding
areas through a gradual increase in the number
of cortical granule cells.
Conclusion
More rostrally to this region, one finds the

segments of the granular frontal area FD (area EK
11) that are rich in granule cells. The experiences
of the war have taught us that the result of lesions
in this region (called prefrontal region) is a disturbance of attention, of psychomotor, of will, of the
emotional apparatus, the appearance of ethical
defects and morally regressive changes in character [Feuchtwanger, 1923]. Such observations lead
us to the localization of certain of the higher
mental functions in these regions of the brain.
But functions such as attention, psychomotor
state, will, ethical conduct etc. are various forms
of the personal activity, which is a motility of the
highest order, as it were. This is the component of
mental personality that I call the effector part.
As a matter of fact, those war-injured patients,
who had prefrontal lesions, showed intellectual
defects characteristic of certain forms of demen-
185
Fig. 69. The progressive change of the characteristics of layers I through VI in the human frontal cortex, from the
precentral region FA (left) to the polar region FG (right). Originally figure 18 in Economo [1929a].
186
tia, which we also encounter in psychoses that are

based on inflammatory changes (such as general
paralysis), and in purely functional psychoses,
such as schizophrenia, especially its catatonic
and hebephrenic forms. These are disturbances
of attention, will, conduct, and also purely ethical
defects, such as those encountered in moral insanity.
Since the inferior frontal gyrus (F3) primarily
subserves the function of speech, and the anterior segment of the superior frontal gyrus (F1) most
probably subserves static and equilibrating functions, then the anterior parts of the middle frontal gyrus (F2) and the frontal pole, subserve the
normal course of this higher, mental, individual
activity. This segment of F2 is very rich in granule
cells (middle granular frontal area FD or area
EK 16 and frontopolar area FE or area EK 18);
they often remind the parietal cortical structural
type 3. This center of the active mental personality in the prefrontal lobe is the counterpart of the
above-mentioned center of the sensory-receptive
mental personality in the posteroinferior parietal
lobe.
Sensory and Effector Personality,

Understanding and Reason
The neuropathology of the aphasias has already

taught us that speech functions are clearly divided into a motor and a sensory component. This
fact corresponds to the fundamental anatomic
cytoarchitectonic division of the cerebral hemispheres into an anterior, more effector part, and
a posterior, more receptive part. Such a division
into effector and receptive components is, as we
now realize, also observed in higher intellectual
functions, and even in the personality overall.
Centers for the effector part of the personality,
localized in anterior parts of the frontal lobe, primarily correspond to the creative activities of our
reason, while sensory-receptive parts correspond
to conscious awareness. However, this does not
Conclusion
mean that I am here defining two distinctly localizable or localized centers of intelligence, one
at the anterior and the other at the posterior half
of the cerebral hemispheres, apart from which
there are no other mental or intellectual capacities of a higher degree in the cerebrum. On the
contrary, I must emphasize the fact that the functioning of the entire cerebrum is necessary for
any kind of mental expression. I can nonetheless
say that there are cerebral parts, the injury of
which in no way disturbs so exclusively the mental condition of the individual as the injury of the
two centers mentioned above.
Let us return for a moment to the analogy of
the cerebrum with a sensory apparatus, such as
the eye. Just as the retina is sensitive to light in all
its parts and yet needs its totality to produce an
accurate image, so is the intact condition of the
entire cerebrum necessary for the execution of
perfect mental conscious functions. The retina,
however, possesses the macula lutea, a region
where sight is most potent and concentrated in a
focus. The same comparison holds for the two cerebral centers described above: these are very
specially differentiated cortical fields for the execution of intensified mental activity: two focal
points of intelligence around which the remaining functions are grouped, just like peripheral vision is grouped around the central vision of the
macula lutea.
Phylogenetic Differentiation of Animals
The cytoarchitectonic study of the brains of vertebrates has further shown that in addition to the
absence of the inferior frontal gyrus (F3), which
corresponds to the absence of developed speech,
there is also an underdevelopment of the two
fields for the execution of higher mental and intellectual functions described above. The supramarginal (PF ), angular (PG) and temporo-occipital (PH ) ground areas of the inferior parietal lobe
are all absent from lower vertebrates, even from
187
188
lower primates, and are poorly developed in the

great apes. The granular prefrontal areas of the
cerebral cortex are also poorly developed in animals, and the anterior segments of the granular
frontal area FD (especially its middle modification FD) seem to be missing.
In addition, the examination of animal brains
has shown that they often reveal large areas with
a single type of structure, which, in the human
brain, are divided into a number of well-defined,
individual areas. For instance, the first five
ground areas of the human superior parietal lobule, namely areas PA, PB , PC, PD and PE , only
form a single area in the mouse brain [Economo,
1930e]. Such an area in the animal brain reveals
an indistinctly mixed cytoarchitectonic structural type. Thus, we find that there is not only an
increase in the mass of the brain as we move from
simpler animals to humans, but there is also an
increase of areas, i.e. a special differentiation of
the various parts of the cortical surface, in addition to which humans possess very recent cerebral acquisitions. It is very plausible to connect
these structures with the specific human functions of intelligence.
Fig. 70. Rear view of both cerebral hemispheres in a human brain showing the presence of a large occipital
operculum. In the left occipital pole, one can see a sort
of Affenspalte, which runs in the dorsoventral direction
far laterally on the hemispheric convexity, while in the
right occipital pole the same abnormality is expressed
somewhat less strongly, but nonetheless still appearing
clearly. The two drawing variants originally appeared as
figure 24 (upper) in Economo [1929a] and as figure 45
(lower) in Economo [1930b]. ac.i = Inferior accessory
occipital sulcus; ac.s = superior accessory occipital sulcus; B = perpendicular occipital sulcus of Bischoff; C or
Calc. = calcarine sulcus; C(e) or Calc. ext. = external calcarine sulcus; Calc. i. = internal calcarine sulcus; ip = interparietal sulcus; o.i = inferior occipital sulcus; o.l.m = lateral occipital sulcus; o.tr1 = medial (dorsal) ramus of
transverse occipital sulcus; o.tr2 = lateral (ventral) ramus
of transverse occipital sulcus, also io, interoccipital sulcus; po = parieto-occipital sulcus (medial); s.l. or s.l.o =
opercular sulcus limitans.
Conclusion
Differences in the Human Ancestral Line
Studies on the skull casts from evolutionary stages of various prehistoric types, such as those that
Osborn [1910], Elliot Smith [1927], Tilney [1928]
and others have conducted, further taught us
[Economo, 1929a] that in the scale which begins
with Pithecanthropus erectus (the Java man,
skull capacity 940 cm3), and leads via the Eoanthropus dawsoni (Piltdown man, skull capacity
1,170 cm3), Homo rhodesiensis (Rhodesian man,
skull capacity 1,300 cm3), Homo sapiens neanderthalensis (Neanderthal man, skull capacity 1,400
cm3), and Homo sapiens sapiens (Cr-Magnon
and Pedmost man, skull capacity 1,500 cm3),
there is a gradual increase of precisely those cortical, basal, and parietal areas to which we have
ascribed through comparative neuroanatomic
research the higher human functions, even in
cases in which, exceptionally, the total capacity of
the brain might exceed that of Homo sapiens,
with its current skull capacity of 1,550 cm3 (e.g. a
continual increase of the speech center and of
both focal points of intelligence, the prefrontal
and the parietal).
We are even able to reconstruct some superficial cytoarchitectonic areas configurative of antediluvian8 man by the knowledge of some of
their gross anatomic particularities in certain
brain regions. For instance, we know from skull
casts that those ancestors of ours had what we call
an operculum occipitale very similar to that of
monkeys initially termed Affenspalte in German and later designated as sulcus lunatus by
Elliot Smith [1904a, 1927] (cf. also Economo
[1930b]). We now know, from the exceptional occurrence of such an occipital operculum in modern humans, that it corresponds to an extension
in the primary sensory visual areas over the lat-
Term as used by Economo [1929d]; modern explanations

include prehistoric, of or belonging to the time before the biblical flood, perhaps associated with Homo neanderthalensis or
Homo erectus.
189
eral convexity of the cerebral hemisphere, whereas in the normal modern brain this area is smaller and runs only on the median (interhemispheric) surface (fig. 70). Such an extension of the
sensory area over the lateral hemispheric aspect
reduces the space left in the lower parietal lobe
from higher mental functions of association, and
it is the increase in associative parts of the brain
in modern humans that has caused the recession
of the sensory cortex and the medial surfaces
[Economo, 1930e].
Thus, the species of vertebrates called Homo
sapiens not only reveals a corresponding increase
in brain mass, but primarily an increase in the
structural fineness and a differentiation of new
cerebral organs. This process of an increasing
brain mass in the upward evolutionary trend of a
species, as well as its progressive differentiation
of cytoarchitectonically specific structures and
the acquisition of new cerebral organs, is what I
wish to call progressive cerebration. This process
must be differentiated from the hitherto known
meaning of the simple phylogenetic course. With
regard to humans, this progressive cerebration is
already a fact.
The further study of the phylogeny of individual animal families discloses the curious fact
that, when one traces the evolution through the
entire ancestral line of an animal species, as for
example in the horse from the four-toed paleotherium, the three-toed architherium, and the hipparion to the single-toed horse, there is a gradual
increase in the growth of the brain (i.e. skull capacity) proportionally greater than the increase
in the size of the body. The same is true of the
dog, the swine, the bear, the rhinoceros, the hippopotamus, and even of birds that have persisted
past the Diluvian age (fig. 71). The modern bear,
e.g., has an absolutely and relatively larger brain
than the bear of the Ice Age, which was much
larger in body size.
190
The Natural Law of Progressive Cerebration
Progressive cerebration thus seems to be a general law of nature, which states that there is a continuous and general increase in the higher mental
functions of living creatures of our planet, in other words, a continuous increase in intelligence.
Whether this natural law of progressive cerebration is an evolutionary trend, a priori, inherent in
the genetic material of cells or whether it is a result of natural selection following upon the struggle for existence, is a question I cannot enter upon
here. It is very probable that the latter case is in
fact true.
Progressive cerebration does not only mean a
biologically quantitative increase in the capacities already present, but the possibility of the development of new cerebral organs in the cortex
that would also enhance the emergence of new
mental capabilities, a circumstance that opens up
entirely new perspectives for human evolution.
In this sense, it is quite possible that individual
variations that exceed the human average (insofar as they are based on anatomic changes of the
cortex), e.g. genius, great talent, criminality, and
psychosis, may be conceived as diversities or idiovariations through which nature achieves her results in the course of generations of selection. At
this point it may become possible for mankind to
control its future evolution through arbitrary eugenics. This also serves to demonstrate how important cytoarchitectonics may sometimes be in
enabling us to anatomically understand these
plus and minus variations of the noblest of our
organs.
Method
In conclusion, I must add a few words on the research method of cytoarchitectonics. The staining method of choice is the Nissl cytoplasmic
stain and any of its modifications (toluidine blue
or cresyl violet).
Fig. 71. Schematic depiction of the increase in brain mass exhibited by the modern forms of four animal species (right
column), compared with their ancestral forms (left column). For instance, in the modern domestic swine (Sus scrofa), the
neopallium reaches a size that constitutes approximately two-thirds of the lateral hemispheric surface, whereas in the
oligocene forms of this animal class, the neopallium only covered one-half of the lateral hemispheric surface. Olf. = Olfactory bulbs; Cer. = cerebrum; Cbl. = cerebellum. Reproduced from Osborn [1910] as figure 25 in Economo [1929a].
Conclusion
191
192
To obtain correct images of the cortical layers,

one must make certain that sections are perpendicular both to the cortical surface and to the
course of the gyrus in question (cf. fig. 13 in
Economo and Koskinas [2008]). This is absolutely necessary in order to have the cortex, and all its
layers, represented at their actual minimal thickness in each section. Only when such a requirement is fulfilled, can the different parts of the
cortex be compared to each other; this automatically entails that it is not possible at all to conduct
cytoarchitectonic studies with the usual large serial sections that pass in cross-section through
the entire brain (fig. 72). Rather, it is indispensable to block, as we have done (slice sectioning
method), each separate gyrus into a series of individual, small, perpendicular slices, by dividing
the entire hemisphere into analogous small blocks
(cf. also Economo and Koskinas [1925, pp. 249
258] and Economo and Koskinas [2008, pp. 13
19]).
After fixing the brain in 5% formalin for 2
days, every gyrus is cut into multiple small blocks,
about 4 mm thick, which must always remain
perpendicular to its surface. A further fixation
follows in 10% formalin for 24 h, washing of the
slices in running H2O for a few hours, and then
hardening in ascending concentrations of ethanol (50100%) for 8 days. From absolute ethanol,
the slices are placed into xylene for a few hours,
and then embedded in paraffin. This treatment

of the material ensures as little shrinkage as possible; thus, the mass of the cortex suffers no great
displacement in the preparations in comparison
with the normal state.
It is advisable to cut the sections at 25 m, because that thickness allows the recognition of
sufficient cell details and particularities of the
cellular layers that cannot be recognized in thinner sections. Sections are stained with 0.1%
Grblers toluidine blue, and the differentiation
of the stain is effected with aniline oil-alcohol. To
obtain an as thorough and regular stain of the
entire specimen as possible, we advise not to
mount the paraffin sections on slides before
staining, but rather to drop them directly into xylene from the microtome, in order to dissolve the
paraffin. Successive treatment with alcohol in
descending concentrations removes the xylene,
after which preparations are immersed in distilled H2O for 2 h, and then into the staining solution. This method ensures thoroughly and
equally-stained preparations, in which deeplystained cells always stand out against the uncolored ground substance. The preparations are
thus altogether suitable for photographic purposes. The method makes it possible to obtain wellstained preparations from all parts of the cerebral hemispheres.
Fig. 72. A series of six parasagittal sections moving

from medial to lateral planes from section 1 to section
6 schematically depicting occipital and parietal area
relationships in whole brain sections, based on cytoarchitectonic data from the miscoscopic study of perpendicular sections. Area OC is hatched in red, areas OA and
OB are left white, and parietal areas are hatched in blue;
the boundary between area OA and area OB is marked
with a black line. The neighboring gyri 1, 2, and
, and the caudal part of the transition gyrus I are covered by the occipital area OA (and OB ). Abbreviations
(other than cytoarchitectonic area codes) as in figure 70.
Originally figure 48 in Economo [1930b].
Conclusion
193
Appendix
An Outline of Cytoarchitectonics of the Adult Human
Cerebral Cortex
Georg N. Koskinas
Preface
No one can deny the importance and the complexity of the functions of the cerebral cortex and
mental phenomena. Given that each physiologic
function, as a general principle, presupposes a
English translation by Lazaros C. Triarhou.

The Outline of Cytoarchitectonics was published by Georg
N. Koskinas (18851975), Economos collaborator on the Atlas
and larger Textbook of cytoarchitectonics, in 1931 (the final
year of Economos life) in Greek, in a monograph explicating his
complete scientific works, as part of the petition for his ill-fated
candidacy to the Chair of Neurology and Psychiatry at the University of Athens following his repatriation to Greece from Vienna. I discovered a copy the only one known to be extant of
that hitherto unknown monograph in February 2005 at the National Library of Greece in Athens, in all likelihood personally
furnished by Koskinas; its octavo folia had never been cut. The
staff of the National Library kindly gave me permission to snip
and photograph all 114 pages of the monograph. The English
version of the unabridged Outline [Koskinas, 1931] is being published for the first time; in essence, it epitomizes the fundamental points of the first six chapters or General Part (pages 1258)
of the 1925 textbook [Economo and Koskinas, 1925], in a way
that only an insider, i.e. one of the original authors, could. Thus,
it complements in a meaningful way Economos Zellaufbau, the
latter being mainly an epitome of the last eight chapters or
Special Part (pp. 259792) of the larger textbook. Koskinass
overview is more exhaustive and highlights more details than
Economos introductory chapter; figures 7388, to which Koskinas refers, have been supplemented from the original Textband
[Economo and Koskinas, 1925] by permission from SpringerVerlag, Vienna.
corresponding anatomic basis, one comprehends

how elaborate the structure of an organ performing such complex and important functions must
be, and how meaningful the study of its structure
is in the quest for understanding and interpreting
such functions.
The fact that its study has not been exhausted
thus far is due to the nature of the problem, which
indeed presents the utmost difficulties stemming
from the complexity of cerebral cortical structure. Being fully aware of those difficulties, but of
the importance of the problem as well, we opted
to contribute to its solution to the extent of our
abilities, and proceeded with the present work.
Preliminary Observations
Introduction
Over millennia, the structure of the brain instigated the greatest interest in European civilization; ancient Greek science already accepted that
in the cerebral matter end the impressions from
the senses and from it stem the movements of the
body, that it is the seat of the intellect, consciousness and the psyche (Galen of Pergamon, c.130
200 AD). One might think that one is listening to

a contemporary scientist, if one hears that in the
third century BC, Erasistratus of Chios (c.304
250 BC) considered humans as the most intelligent beings, exactly because their brain is the
richest in gyri.
That knowledge as well as all the once-expressed great truths were never completely lost
for humanity. It is certainly true that in Western
Europe, under the ruins of civilization and under
the influence of superstition, they fell into oblivion for a long time; but in the Eastern Roman Nation, the uninterrupted experience of the devout
pupils and heirs of the great masters of antiquity,
up to their last celebrated representative Johannes
Actuarius (c.12751328 AD), owing to the preserving power of the elevated Byzantine civilization during the Middle Ages and its endurance,
preserved this treasure of knowledge and other
elements of culture and transmitted them, still
vivid, to their bairns, in the middle of the tempest
of the general intellectual decay and the ferociousness of its neighbors.
Directly relying on that knowledge, the new
generations of the peoples of Western Europe,
with the conquest of barbarity, betook themselves anew, from the Renaissance and hither, to
the solution of those problems. Thenceforth, the
old medieval spirit and the false interpretation of
religious dogma only seldom interfered with the
further evolution of such a knowledge, and as a
curiosity of the kind, we may mention, for example, that in 1805, Gall was forced to leave Vienna, because he placed the seat of mental powers in the brain. The view, according to which the
brain is the seat of our higher mental functions,
has been since recognized as generally correct.
Therefore, the study of its structure elicited more
and more interest. But the modality of its activity still remained enigmatic for some time. However, the knowledge on electricity (current, conduction, induction, capacitance, etc.) and the
certain analogy between these physical phenomena and the animal functions of the nervous sys-
Appendix
tem rendered the latter relatively comprehensible

after a while.
Thus, assisted by all natural sciences and specifically by comparative anatomy and brain physiology, particularly in the last decades, we understood, more and more, that a higher special evolution of the central nervous system corresponds
to the evolution of nervous phenomena as its material basis, from the simple reflex movement to
the rather complex purposive acts, and that the
utmost development of mental powers, all the way
to the ingenious creative achievements of the human spirit, has as its organic term and material
basis the paramount evolution of the fine structure
of the brain and especially its nerve cells. The creative ability of the individual, through which one
creates anew not only by means of general physical reproduction, but also through ones personal
knowledge, although already presenting in a rudimentary form in animals of higher orders,
nonetheless belongs only to humans to such a
great degree, at least on the Earth.
In the phylogenetic line of the living organisms, nature works creatively, sometimes very
slowly, other times in leaps, but anyhow constantly, towards the production of new more complex
forms and forces of life. Indeed, that creative force
of nature, which in the course of geologic eons
formed the wings of the eagle, indirectly rendered
humans able to construct wings, through the
growth of their knowledge, in order to conquer
gravity. That part of the general creative, even divine one might say, principle, which was transferred to the rather evolved organisms, has its organic expression, its seat or temple, in the brain;
therefore, we are justified in saying that the discovery of the structure and the nature of our noblest
organ is a value of the greatest efforts of science.
Similarly to sensory organs, e.g. the optic vesicles, which emanate from the diencephalon, the
cerebral hemispheric vesicles are partitioned in a
pair from the azygous holospheric prosencephalon; one might then consider the cerebral cortex
thus formed as a sensory organ, whose visual
195
field, however, is not pursuing the external world,

but the internal events of the central nervous system. The signals that arrive at such a sensory organ do not derive directly from the periphery, but
represent internal stimuli, which from the entire
nervous system end up in the brain, are all conceived concurrently and are subjected to further
processing to make up an integral. The cerebral
cortex also has the ability to accumulate those impressions of the impulses, such that superfluous
components of the energy of the stimulus, which
are not conducted again outbound via the simple
reflex arch as a direct result, become stored in it.
Becoming thus able to convert the past and present
energy into future energy, the organism gets rid of
the brute initial law of reflex action and is provided with individual freedom and personality.
Therefore, we may expect much enlightenment from the precise knowledge of the structure
of the cerebral cortex upon issues of the utmost
importance, such as the anatomic bases of the
natural course of mental phenomena and the relation of certain physical attributes to brain structure; on such issues rests already the will to appoach the problem of problems, the problem of the
psyche. Moreover, general issues of anatomy, comparative anatomy, physiologic localization, and
finally psychopathology and pathologic anatomy
are involved. Until now, one expected the solution
of these encompassing problems to derive, in total
or in part, from the gross anatomy of the brain.
Macroscopic anatomy, pathology and experimental physiology already offered us very deep, in
part final, views on certain of these problems, and
we can hope that the knowledge of the finer structure of the cerebral cortex will provide us with
new and more essential revelations in this field.
To what point can we generally hope that we
may advance in this field on the basis of anatomic
knowledge? When we, as anatomists, speak about
the problem of the psyche, by the term psyche we do
not imply some metaphysical being falling a priori
outside any anatomic and physiologic study, but
the psyche that appears to us from other humans as
196
the moral, intellectual, active, as the historical personality, acting upon ourselves. The concept of the
psyche, so narrowly considered, is only partially
an expression of the structure of the brain, in other words it can only be partially understood in anatomic terms. It bears the same relation to the
structure of the brain as the melody does to the
composition on the keyboard on which it is produced. In any event, its relation to the brain is even
closer, regarding the historical component of the
personality; although, perhaps, we cannot conceive it through microscopic anatomy, it is, nonetheless, certainly at least localized in the brain,
and materially connected somehow as an engram.
However, although the problem cannot perhaps
be fully solved from the structure, we can nevertheless deduce from it important conclusions, as
we may deduce, to carry on the above example,
from the quality or lack of strings and keys of the
piano, at least certain attributes and defects of the
melody. Thus, severe mental deficits, such as advanced forms of mental retardation, can always be
recognized from the structural aberrations of the
brain in general and of the cerebral cortex in particular, either from the absence or the aberrant
position and form of its cells.
Accordingly, may we not hypothesize that
lesser degrees of mental retardation, even those
permitting a social life, correspond to such defects in structure, albeit relatively milder, and
that the limit of resolution, to which we may anatomically monitor mental defects is merely a matter of the sophistication of our methods and the
acuity of our sight? And vice versa? If we contrast
the inherent absolute lack of music perception or
the ability to discriminate pitch and tempo, noticed in some people, with the profuse and largely inherent talent of others for music comprehension, is not it possible that, when we decipher the
sensory fields of the cerebral cortex, we might
discover the differences in structure and extent
that correspond to such extreme variations in
abilities, and thus, in a wider sense, consider certain of the bases of various abilities and genius as
purely anatomic? The study of the structure of

the cerebral cortex will thus lead us closest to the
problem of individual mental attributes and their
anatomic correlates. Even from that standpoint,
gross anatomy and pathology effected a fundamental and successful preparatory work through
the theory of the agnosias.
For the psychoses in particular, many await
the solution of plentiful issues from the fine anatomy and the pathologic anatomy of the cerebral
cortex, totally understandably to a certain extent,
but not exclusively. For those who think that all
brain functions can always be resolved, ultimately and definitely, in a large series of reflex phenomena, anatomy, i.e. the knowledge of these reflex archs, is everything. But often, one does not
sufficiently take into account that such an overestimation of anatomy is incorrect, although it is
generally known that, e.g., exogenous or endogenous chemical reactions (intoxication) exert a
great influence on mental manifestations, especially through the formation or the disruption of
the reflex archs.
A most common example of such a chemical,
i.e. hormonal, formation of an arch is the embracing reflex in the frog that is localized in the
upper thoracic segments, which only appears
during the period of the estrus. Many emotional,
therefore mental, functions, owe their occurrence to such hormonal causes, such as orgasm
and its highest ethical correlate, love, and perhaps anxiety (animals deprived of epinephrine
do not manifest any reflex anxiety anymore),
sentimentality (Basedow disease), and more.
It appears that in such manifestations and
their abnormal increase, the cerebral cortex only
plays a secondary role. Besides this, however,
even the knowledge of encephalitis lethargica
and its sequelae taught us that numerous mental
functions, normal and pathologic, which we hitherto considered as functionally subserved by the
cerebral cortex (will, initiative, psychomotorium,
elicitation of emotions, vividness of thoughts
etc.), in principle have their initial domain out-
Appendix
side the cortex, mostly in the gray masses of the

central ganglia. Thus, an entire group of functions can be distinguished, so-called mental,
whose initial location is either not at all, or at best
partially, found in the cortex.
Although today we consider the cerebral cortex and its complex structure as the material basis
of all the things that we call higher intellect,
memory, thought, consciousness and awareness,
perhaps, with the progress of our science, we shall
be nonetheless forced to detach from such a totality this or that component. And if we ponder that
each of those functions (e.g. the intellect) is not
elemental, but rather an entire complex of functions, it becomes very likely that some such complexes may also be localized outside the cortex, as
we have seen above, in this case the psychomotor,
among others.
These last observations of ours may appear to
perhaps limit the importance of the study of the
cerebral cortex, which we emphasized at the beginning of this introduction. But no one could
seriously think that we, having exactly undertaken a description of the finer structure of the cerebral cortex, undervalue the importance of its
anatomy; we merely wish to emphasize that we
must not expect everything from it alone, particularly in view of the fact that, although the cortex constitutes the most important organ of our
mental functions, the totality of the brain nonetheless participates in them, and significant components of mental events are partially localized
outside the cerebral cortex, and even might not
be localizable at all, in the common sense of the
word, in the nervous system.
We kept such limitations in mind during the
execution of our work, such that, certainly, nobody can accuse us of single-sidedly approaching
the theme.
It is obvious that early researchers, who laid
the fundamentals of cortical cytoarchitectonics
and distinguished various regions, already addressed important issues, such as to what extent
do defined histologic regions coincide with func-
197
tional and excitatory centers localized in the various lobes and gyri and to what extent may we
draw conclusions on the function of the cortex
from its structure.
We shall later see that the motor cortex has a
particular structure and that from such analyses
it ensues that the sensory cortex is immediately
recognized as such from its structure as well. And
the remaining issues, those related to our mental
functions and abilities, for which only general
short principles had been laid before, become
clarified in a new way under the light of such a
subdivision of the cortex into areas.
As is natural, and as said above, in such studies we must always keep in mind that our mental
abilities form, for the most part, complexes,
whose psychologic components do not necessarily coincide with their anatomophysiologic components; thus, e.g., the psychologic analysis of the
ability of speech almost in no way could lead to
the motor and sensory components of speech and
its remaining elements, which we know from pathology and anatomy.
But even for the understanding of the psychoses from an anatomic viewpoint, the knowledge
of the layer structure and areal subdivision of the
cortex opens up new paths. The issue is immediately raised whether systematic lesions of the cortex in layers are possible, especially in inherited
diseases. Here too, in recent years, pathology research, in its impatience to find a new field of
work on the grounds of cytoarchitectonics, led
into numerous immature works, whose potentially correct or erroneous results will only be
possible to discern later on the basis of normal
observations, such as those in our work.
It will be possible to subject all problems and
issues raised a long time ago to a successful investigation only when we gain, firstly, a precise
knowledge of the normal finer structure of the
cerebral cortex, and secondly, the knowledge of
the apparent variations from the typical structure, within normal limits. Anyone involved in
issues of brain physiology and psychopathology
198
realizes that, to reach such a goal, it is absolutely

and above all necessary to finally have a precise
description and an illustration through figures of
the total structure of the cortex, and must have
felt more and more the lack of such an integrative
presentation of the matter. To exactly replenish
that deficiency which we have also so often felt,
we expose in our work the description of the normal cytoarchitectonics of the cortex of the forebrain. And we hope that it may be possible for our
work to be used as the basis for future research, and
that is why we attributed importance mainly to the
presentation through images, i.e. in the Atlas.
History
Here we mention in detail the names and the
works of all researchers before us, who occupied
themselves with the investigation of the fine
structure and the cytoarchitectonics of the cerebral cortex. The fundamentals for the detailed investigation of cerebral cytoarchitecture, upon
which later researchers based their works, were
laid over decades. However, the phenomenon has
repeatedly occurred, especially in recent years,
even when works of great value were concerned,
that old fundamentals, or even knowledge gained
long ago, were presented under new alien names
or even dressed under a plenitude of neologisms
as allegedly new revelations.
To the extent that such transformations of already known information, due to the progress of
our knowledge, appear necessary, they must indeed be implemented (especially respecting the
priority of previous researchers, as far as essential
points are concerned). But whenever they are not
necessary, which is often the case, they only constitute a great obstacle for a correct understanding of things, because the reader, in order to find
the correspondence of the allegedly new items to
those already known for a long time, is subjected
to a retrospective mental work, of which we wanted to rid him. That is why in our analyses we
maintained, as far as it was possible, the accepted

subdivisions and names established by previous
authors, without introducing any new term, unless it corresponded to a new fact. In the next section we mention in their historical sequence the
most important fundamental works, on which
our knowledge of the structure of the cerebral
cortex, and particularly of cytoarchitectonics,
was based until now.
Experimental Design
As exposed in the history part, the first studies on
the fine structure of the cortex took place some
150 years ago. However, the study of the cytoarchitectonics of the cerebral cortex in particular
evolved over the past 70 years. Thus, Meynert
[1867/1868, 1868, 1872a, b] in Vienna, based on
his own studies on the cytoarchitectonics of the
brain, and even before the work of Fritsch and
Hitzig [1870], first conceived the idea of cortical
organology and realized as a matter of fact certain
of its principles, which apply even today; therefore, he is the one who deserves the honor of not
only having founded this discipline of science, but
also defined, once and for all, its successful directions. Subsequently, Betz [1874, 1881] developed
those thoughts and described the distinction of
cortical areas from a cytoarchitectonic aspect
that are partly independent of gyral boundaries.
Ramn y Cajal [19001906] described in detail
the forms of cells. Hammarberg [1895] published
for the first time brilliant normal images of the
rather different parts of cortex consisting of six
layers, as well as the rest, and correlated them
to corresponding pathologic images. Campbell
[1903, 1905] and Elliot Smith [1904b, c, 1907] were
the first to publish detailed and good maps of the
brain, providing the distribution of the cortical
surface into areas; Campbell [1903, 1905] in particular produced a valuable atlas with schematic
diagrams of the structure of the most important
of these areas, also taking myeloarchitectonics
Appendix
into account. Brodmann [1903a, b, 1905a, b, 1906,

1908ac, 1909, 1914], through most extensive
studies of the comparative anatomy of the cortex,
provided a wide scientific basis for future systematic research, and finally Vogt executed the preparatory works on the knowledge of myeloarchitectonics which are of utmost value until today
[Vogt, 1902, 1903, 1906, 1910, 1911, 1913; Vogt
and Vogt, 1902, 1910, 1919].
These are the landmarks of the study of the cytoarchitectonics of the cerebral cortex; in examining the various areas of the cortex, we repeatedly return to the works of these authors in particular, and by further building upon them, we
expose our new observations and develop our
views based on our photographic Atlas [Economo
and Koskinas, 1925, 2008], which will form an objective criterion for this new scientific branch and
will render anyone working on this topic independent of our present views and traditions,
therefore to a wide measure at least independent
even of our own subdivision of the cortex into areas and of our text. Any systematization in science must be considered only as a simple technical aid for the better understanding of nature and
for the intercommunication of scientists. As such,
it always involves a personal element. It forms, so
to say, the coordinates that we ourselves etch, but
which are conceivably foreign to the essence of the
phenomena. On the contrary, photographic images
are impersonal and speak by themselves.
Our work thus consists of the Atlas and of the
explanatory text included in the special volume.
The Atlas (the description of the text will follow the description of the Atlas) contains 134
photographs, 40 ! 40 cm in size, organized into
112 plates of the most different areas of the cortex;
102 of these figures, representing the most important parts of the cerebral cortex, were made under
a magnification of !100, such that 1 mm of the
scale in each figure corresponds to 10 m and
thus one can immediately discern the actual sizes
through direct measurements on the photographs. Scales on the sides of the figures allow not
199
only measurements of size, but also a direct orientation and definition of any point on the plate;
thus, e.g., if we say that a point in the figure lies at
a height of 20 cm and a width of 15 cm on the
scale, we define it precisely through these coordinates. Only 32 pictures, referring to the so-called
rhinencephalon, in which one is more interested
in the general topographic orientation rather than
the histologic details, were made under a magnification of !50, and some of those even under a
magnification of !25, which is marked especially
in each figure; therefore, in those figures, 1 mm
corresponds to 20 and 40 m, respectively. All the
figures are direct photographic prints of microphotographs without emendations (not photocopies or printed drawings), such that each detail
is genuine, without any addition.
The photographed sections of the cortex had a
thickness of 25 m, such that the various plates
might be compared to each other regarding number, size, density and position of cells, and the
data can be immediately and correctly calculated
from the plates. All of the photographs were made
with Zeiss Planar objective lenses, of a focal length
of 2 cm, such that the entire section thickness appears in the figure with the same clarity and with
all its elements. The exact location from which
each photograph was taken can be seen in figure
1 in the Atlas [Economo and Koskinas, 2008].
We reckon that the magnification of !100,
which we selected after multiple trials, is by far
the best, because it allows us to discern directly,
without using any other lens, every detail and
renders each measurement very simple.
The text, consisting of 810 pages in great octavo, incorporates, for almost all the complex
problems it covers, numerous original printed
figures and drawings made by us, through which
even the understanding of the most abstruse issues is incredibly facilitated. In order to be able to
include in our account this entire broad field of
cytoarchitectonics, we divided the text into a
General and a Special Part, comprising in all 14
chapters.
200
For an exact localization of each of the areas

we defined and described upon the various surfaces of the cerebral hemispheres, a certain compound and fundamental schematic representation of those surfaces was necessary, depicting all
of their details, i.e. sulci and gyri, so that the conclusions of our research would be recorded on the
fundamental compound scheme, thus defining
their location with precision.
Such a scheme was not available until now;
that is why we had to proceed with such a drawing ourselves (cf. fig. 19 in Economo and Koskinas [2008]). To this end, we put together pre-existing partial schemes by previous investigators,
with the related descriptions. And because, as
known, numerous albeit small variations are
observed among various brains (as far as direction, form and other parameters of sulci and gyri
are concerned, even the presence or absence of
certain of them of secondary importance), we examined several brains, and took for each group of
data the average; that is how we compiled our
fundamental schematic figure. We registered everything we found in our studies on that scheme;
moreover, this fundamental scheme can be used
towards a common realization in all future works
on the subject.
From that fundamental scheme we prepared
additional general drawings, in which we present, through the use of numbers or through different hatching or coloring, the thickness of the
cortex, the number and the presence or absence
of various types of nerve cells in regions and areas. Thus, one most readily gets an overall picture
of the distribution of these elements over the entire hemispheric surface.
General Remarks on the Cortex and Its Nerve

Cells
Any researcher who attempts to estimate the cortical thickness must set rules, according to which
one may define the boundaries of the cortex and
strictly adhere in all measurements. We obtained

our measurements in paraffin sections stained
with toluidine blue. We made the sections in the
only way appropriate for measurements, i.e. perpendicularly to the surface of each gyrus, and more
specifically its dome, walls and valley floors.
All the blocks, 250350 in number, into which
each brain was dissected, as well as the sections
that derived from them, were subjected to the action of the same chemical substances, for the
same period of time, and under the same temperature.
We examined each part of the cortex in the
brains of several subjects of about the same age.
We observed that cortical thickness varies in different brain areas (fig. 73) and in different segments of the same gyrus (dome, wall, valley
floor); the differences of these various parts are
influenced by the shape of the gyrus.
Differences are also observed among individuals; that is why for each part of the cerebral cortex we took the mean of the numbers gathered
from the various brains.
To provide the reader with an overall view of
those differences in thickness, we compiled comprehensive schematic drawings (fig. 74), in which
the differences in the thickness of the dome of the
various gyri are depicted through different degrees of shading. To this end, we further compiled schematic drawings, whereby the dome
thickness in various gyri is denoted by numbers
(cf. fig. 20 in the Atlas [Economo and Koskinas,
2008]). Finally, in additional schematic illustrations (fig. 75), we depicted the differences in
thickness of the various segments of each gyrus.
We calculated the volume of the cortex by
means of pure calculus methods, while Jger
[1914] had used toward that goal, before us, the
so-called surface-measuring method. The result
we obtained (V = 555 cm3) agrees, almost completely, with that of Jger [1914] (560 cm3). As far
as the surface of the cerebral cortex is concerned,
our estimates are in agreement with those by previous investigators as well. The surface area of the
Appendix
entire cerebral cortex ranges in Europeans from

S = 200,000 to S = 218,000 mm2, about 33% constituting free surface and the remaining 67% the
surface hidden inside sulci.
In the same chapter we describe the neurons
of the cortex, which differ a great deal from each
other, in regard to both their external and internal structure, i.e. shape, size, nucleus, nucleolus,
Nissl bodies, filaments, and finally contents. The
cells are classified into the following five categories: (1) pyramidal cells; (2) spindle cells; (3) granule cells; (4) Cajal cells; and (5) special cells.
(1) As far as pyramidal cells are concerned, we
describe their shape and size. We studied their size
in Nissl-stained or silver-impregnated specimens.
Concerning size, we found as more practical the
measurement of two dimensions, i.e. height (H),
from the base up to the point where the soma ends
and the dendrite begins, and width (W), i.e. the
maximum cross dimension, which in most cases
is found at the base of the cell. To represent the
size, we used a special symbol, consisting of a horizontal line, above which we indicate the height, or
actually its margins, and beneath it the width.
That notation, although looking like a fraction, is
not a fraction in the mathematical sense of the
word. After all, by dividing the height by the width,
i.e. by considering the notation as a fraction, we
obtain a number, the height-to-width (H/W) ratio,
which gives an idea of the shape of the cell; we
called that ratio cell slenderness (Schlankheit).
With regard to size, we distinguish the following six classes of pyramidal cells: (a) pyramidallike granule cells (H = 67 m, W = 5 m); (b)
small pyramidal cells (H = 1015 m, W = 710
m); (c) medium-sized pyramidal cells (H = 20
30 m, W = 1020 m); (d) large pyramidal cells
(H = 3050 m, W = 1520 m); (e) giant pyramidal cells, with a height slightly greater or the
largest from the previous category, but distinguished by being more compact, richer in cytoplasm, comprising at their base and soma more
compact processes, and other features, and (f) giant pyramidal cells of Betz or colossal cells.
201
Fig. 73. A horizontal section through

the left human cerebral hemisphere,
depicting the sizeable regional differences in cortical thickness. Weigert myelin stain. Abbreviations
(moving counterclockwise from
top): F1 = Superior frontal gyrus; F2 =
middle frontal gyrus; Ca = precentral
gyrus; R = central sulcus of Rolando;
Cp = postcentral gyrus, P = parietal
lobe; O = occipital lobe; L = limbic
gyrus. Originally figure 25 in Economo and Koskinas [1925].
202
As far as slenderness is concerned, we categorize the pyramidal cells into: () overly slender
(H/W = 2.5); () slender (H/W = 2.0); () medium
slender (H/W = 1.5); () short triangular (H/W =
1.0); and () flattened (H/W = 0.5).
Besides the above differences regarding size
and slenderness, pyramidal cells also display additional differences in form, depending on the
cortical area. Thus, for example, pyramidal cells
of the superior (T1) and middle (T2) temporal gyrus near the temporal pole are rounder in shape,
and that is why we call them guttiform (Tropfenfrmig).
In certain cortical areas, all the cells of the
other categories assume a more or less pyramidal
form; we termed such a transformation pyramidization (Pyramidisierung) of the cortex. Pyramidal cells present substantial individual differences from brain to brain, being larger and
slenderer in some, and smaller and less slender in
others. We discuss their physiology in chapter 4
[Economo and Koskinas, 1925] and depict all
those size and form differences in lucid schematic illustrations (fig. 76, 77).
(2) Regarding spindle (fusiform) cells, we describe the size, which we represent in the same
way as for the pyramidal cells, the shape and its
regional modifications, and their position.
We occasionally observed that pyramidal cells
of layer V and sometimes of layer III assume a
certain spindle-like form, a change that we called
spindle transformation (Verspindelung) of the
cells. We expose their physiology in chapter 4
[Economo and Koskinas, 1925] and juxtapose all
these form variations in figure 78.
(3) In the section on granule cells, which are
characterized by their extremely small size, we
describe the form, which is most variable (spherical, ovoid, triangular, rhomboid, stellate and pyramidal or spindle) and represent their size with
a symbol similar to the one used for the previous
categories. We expose their most variable disposition in groups under the description of each
area.
Appendix
We observed, with very few exceptions, a certain competition between the development of
these cells and of pyramidal cells.
In some areas, not only is the wealth of granule cells very great, but pyramidal and spindle
(fusiform) cells become smaller and assume a
rather granular-like form, such that the entire
section of the cortex appears as consisting of
granule cells only. We called such a transformation granulization (Verkrnelung), and that type
of cortex we named koniocortex.
We also describe the differences that we found
regarding the shape of granule cells among the
various areas and the segments of each gyrus. We
depict these granule cell varieties in figure 79. We
cover their physiology in chapter 4 [Economo
and Koskinas, 1925].
(4) Concerning Cajal cells (or Cajal-Retzius
cells), we describe the size, which we measured
everywhere in the cortex, as well as their shape
and arrangement, referring their detailed description, on the one hand, to the description of
each area, and the physiology, on the other hand,
to chapter 4 [Economo and Koskinas, 1925].
(5) Under the name special cells we combined
most diverse cellular shapes, each of which is
only found in certain areas of the cortex. In chapter 2 [Economo and Koskinas, 1925] we describe
the following main cell types, according to shape,
size and position: the giant cells of Betz, the giant
stellate or solitary cells of Meynert, the acoustic
cells of Cajal, the tufted cells (Quastenzellen) of
Klliker and Calleja, and lastly the cells that we
observed and described for the first time, which
we named from their shape rod and corkscrew
cells (Stbchen and Korkzieherzellen); we describe exactly the form, composition, size and position where they are found in utmost detail and
with precision. The varieties of the shapes of
these two cells are shown in figure 80.
In the section on cell size, because size varies
from area to area, we distinguished in that respect five grades of cortex, which we describe
with the terms () very magnocellular (sehr zell-
203
Fig. 74. Schematic drawing of the total cortical thickness (layers IVI combined) in a the superolateral convexity, and
b the median hemispheric facies. The substantial variation in cortical thickness is depicted by means of the gray shading intensity. The thinnest (02.0 mm) cortical localities are left white; each additional 0.25 mm in thickeness is incrementally marked by applying an extra coat of Indian ink, such that cortical localities with the greatest thickness of
4.55.0 mm (at the superomedial hemispheric edge of the precentral gyrus) appear black. Originally figures 26 and 27
in Economo and Koskinas [1925]. The posterior segments of the frontal lobe are the thickest in the entire cerebral hemisphere; more rostral segments progressively diminish in thickness toward the frontal pole, and somewhat less so in the
ventral direction; that progressive thinning becomes interrupted rostrally, at a segment lying between the two pointing
204
Fig. 75. Schematic illustration of the

variations in cortical diameter at the
dome, brink, wall and valley floor in
cross-sections of gyri of various
shapes. Types of gyri depending on
location: A = Frontal; B = central; C =
superior parietal; D = inferior parietal; E = orbital/basal; and F = calcarine. One further finds these individual types in additional areas beyond
those mentioned, which actually
represent their most frequent occurrence. Originally figure 30 in
Economo and Koskinas [1925].
gross), () magnocellular (zellgross), () mediummagnocellular (mittel zellgross), () parvicellular

(zellklein), and () very parvicellular (sehr zellklein) cortex. We present the results of the measurements of cell size in table VI of chapter 14
[Economo and Koskinas, 1925].
In the section on cell numbers we expose the
method of their estimation. To estimate cell number in a given area we always define the number
of cells contained in 0.001 mm3. To that end, it
suffices to count the cells in a sampling surface of
0.01 mm2 and to multiply that number by 4, given
that the section thickness of 25 m equals 1/4 !
arrows, where the cortex anteriorly to the midpoint of the

middle frontal gyrus (F2) exhibits a thickened islet in the
vicinity of two secondary transverse gyri. An imprecisely
delimited thickening (asterisk) is observed at the posterior
one-third of the superior frontal gyrus (F1). (Cortical thickness is broken down for layers I and VI in fig. 86, and for
layers III and V in fig. 87 below; layers II and IV are shown
in fig. 11 in the Atlas [Economo and Koskinas, 2008].)
Appendix
0.1 mm, therefore the volume corresponding to a

surface of 0.01 mm2 is only 1/4 ! 0.001 mm3.
One may effect such measurements on the microphotographs. But because these have a magnification of !100, one must take as a base the surface of a square with a side of 0.1 mm multiplied
100 times, that is 1 cm on the plate. We expose the
results of these measurements for each area in table V of chapter 14 [Economo and Koskinas,
1925].
In that respect we distinguished five types of
cortex as well, which we named (a) very cellular
(sehr zellreich), (b) cellular (zellreich), (c) medium-cellular (mittel zellreich), (d) oligocellular
(zellarm), and (e) very oligocellular (sehr zellarm)
cortex. We further observed that anteriorly the
cerebral cortex is rather magnocellular, but poorer in cells, whereas on the contrary, posteriorly
the cortex is rather parvicellular, but reversely,
richer in cells.
We attribute a lot of importance to the determination of the number of cells, because, as
205
Fig. 76. Semi-schematic drawing of various forms and sizes of pyramidal cells at a magnification of about !200:
1, dwarf; 2, small; 3, medium-sized; 4, large; 5, giant; 6, Betz giant or colossal. Originally figure 34 in Economo and
Koskinas [1925].
Fig. 77. Schematic drawing of various slender forms of pyramidal cells, according to cell height (H) over width (W)
ratio. Examples: a, overly slender (H/ W = 2.5); b, slender (H/ W = 2.0); c, medium-slender (H/ W = 1.5); d, flat-triangular
(H/ W = 1.0), and e, flat (H/ W = 0.5) pyramidal cells. Originally figure 35 in Economo and Koskinas [1925].
206
Fig. 78. Semi-schematic drawing of 6 common spindle (fusiform) cell forms: 1, triangular; 2 and 3, spindle proper;
4, triangular spindle; 5, dual spindle; 6, crescent. Originally figure 36 in Economo and Koskinas [1925].
Fig. 79. Semi-schematic drawing of granule cells with substantial variations in form: A, group from layer II (common triangular and spindle form, more isolated); B, group from layer IV (various forms, cells abutting each other);
B, columnar group from layer IV of temporal cortex; t, satellite cells (Trabantzellen). Originally figure 38 in Economo and Koskinas [1925].
Appendix
207
Fig. 80. Our small rod (Stbchen) and

corkscrew cells (Korkzieherzellen) of
the anterior limbic gyrus and the
transverse insular gyrus. Originally
figure 44 in Economo and Koskinas
[1925].
208
known, there are pathologic conditions accompanied by cell loss. In order thus to make a prediction about such a pathologic loss in a given
condition, we have to be aware of the normal
number of cells of that particular area. That is
why such measurements must be always conducted in the exactly corresponding areas of the
various brains, as normal differences exist among
them. Lacking such knowedge, reductions in cell
number were many times characterized as abnormal, although they were totally normal and
only due to differences among areas.
Through a comparison of the various areas of
the cerebral cortex regarding cortical thickness,
as well as cell number, size, form and distribution, it is possible to gradually crystallize a multitude of parameters for each area, which allow its
precise characterization.
From all our measurements of the thickness of
the cortex, and the number and size of cells in the
various areas, it was easy, by means of simple calculus, to estimate much more precisely than investigators before us the total number of cells in
the cerebral cortex, which we found to reach some
14 ! 109, of which about 6 ! 109 are the smallest
granule cells and 8 ! 109 all the remaining larger cells together.
Through calculus we also found that the total
volume of the cells of the cortex is 20.4 cm3, that
these make up only 5% of the entire cortical volume, that their total mass is about 21.5 g and
that the mass of a medium-sized pyramidal cell
is about 40 pg and that of a granule cell about
14 pg.
In the section on cellular density we define it
as the ratio of the stained cytoplasmic substance
in an area over the unstained ground substance.
Such a density depends on the number and size
of cells, increasing with more numerous and larger size cells, because accordingly the stained substance increases and the ground substance decreases. As is evident, the intensity of the stain
depends on the density; it might even be possible
for the intensity of the staining to be defined by
Appendix
means of photographic methods. Such a method,

difficult in its own, has not been defined yet; that
is why we introduce the concept of a density factor, readily calculated from the cell size and number. Toward that goal we suppose that we observe
a cube with a side of 0.1 mm, i.e. with a volume of
0.001 mm3, whereby all the cells in it appear as
projected on one face of a cube with a surface area
of 0.01 mm2.
Considering such projections as approximate
geometrical shapes, e.g. triangles for pyramidal
cells, we calculate from their dimensions in m
the surface of each projection in m2, and by
multiplying the number of cells contained in
0.001 mm3, we obtain the surface of all the projections in m2. We divide that surface by the
total surface of one facet, which is 0.01 mm2 (or
10,000 m2). That ratio constitutes the density
coefficient (Dichtigkeitskoeffizient) and is usually
greater than 1, more rarely equal to 1, which
means that the projections of the cells cover the
entire surface of the facet; most seldom it may rise
to 1.2, i.e. when the density is so great that cells
would partially overlap with each other [Economo and Koskinas, 1925, pp. 7476].
In the section on the disposition of cells we examine the orientation of their axis and relative position to each other. With regard to this last issue,
we observed that besides their common arrangement in fine layers, parallel to the cortical surface,
one also runs into another disposition, where the
cells form streaks vertical to the cortical surface,
in the direction of the so-called myelin radii. We
call that disposition radial (radire).
Because such a disposition differs among various areas, we provide figure 81, which schematically depicts those differences through the entire
cortex. Besides the laminar arrangement, one
also observes an additional disposition in groups,
which, depending on the number of the cells that
form them, are called clusters or huddles. Owing
to that arrangement, there are perforce lacunas
formed between such groups. We draw attention
to that fact for histopathologic comparative stud-
209
Striation (S.) hint

S. distinct
S. very narrow
Columnar disposition (c.d.)
S. ne and c.d.
Granulous S.
Fig. 81. Schematic drawing of the form and distribution of radial striation over the cortex of a the superolateral convex and b median hemispheric surfaces in the cytoarchitectonic picture. The density of the hatching gives a rough
measure of the distinctness and density (compactness) of the radial striation in the individual regions. Originally figures 45 and 46 in Economo and Koskinas [1925].
210
Fig. 82. Semi-schematic drawing of the embryonic developmental phases of the cerebral cortex from the neuroepithelium (ependymre Anlage) up to the full development of the pyramidal layer in the 5th prenatal month. le, li =
External and internal limiting membrane; M = germinal layer; M = matrix originating from M; n = neuroblasts; P =
pia mater; Py = pyramidal or cortical layer, from which the actual cellular cortex develops; RS = marginal zone (subsequently molecular layer); x = germinal layer in marginal zone, from which glial and Cajal (or Cajal-Retzius) cells of
the molecular layer will develop later; Z = intermediate zone, from which the white matter will develop; Z = cortical
subplate or border of intermediate zone Z (subsequently layer VIb). Originally figure 52 in Economo and Koskinas
[1925].
Appendix
211
ies, insofar as such normal rarification of cells

might erroneously be interpreted as a pathologic
change.
Subdivision of the Cortex into Layers and

Their Development
In the third chapter we detail the laminar subdivision of the cerebral cortex into layers. Any such
subdivision is somewhat subjective, hence different authors accepted a different number of layers;
that is why we provide a table comparing the correspondence of layers according to different investigators. We opted for the more practical subdivision into six layers of Bevan Lewis [1878, 1879,
1880]. This appears more or less straightforward
for most areas of the cerebral cortex, while in some
other areas one may observe an altogether different
subdivision. The former type we call, like previous
authors, isogenetic cortex or isocortex for short,
and the latter allogenetic cortex or allocortex.
At some places of the isogenetic cortex, layers
are completely distinguishable from each other;
this type was called homotypic isocortex. At other
places small variations appear, insofar as some of
the layers become rudimentary, disappear altogether, merge with other layers or even splinter
into multiple sublayers; that type was called heterotypic isocortex. The allocortex is not uniform,
but rather comprises diverse forms.
We next examine the embryonic development
of the cortex, showing how the isocortex and the
allocortex can be distinguished from each other
very early on. Regarding the allocortex, we also
proceed with its study in comparison with the
brains of animals.
In the semischematic figure 29 in the Atlas
[Economo and Koskinas, 2008] and in figure 82
we clearly depict the gradual development of the
cerebral cortex from the neuroepithelium (ependymre Anlage) until the perfect formation of the
so-called pyramidal layer during the fifth month
of fetal life.
212
In figure 83, we show the appearance of the

brain during the third month of fetal life, and in
figure 84 ten parallel sections through the brain
shown in figure 83. We depict the areas, in which
the above-mentioned types of cortex appear, in
color figure 22a, b in the Atlas [Economo and
Koskinas, 2008] and in figure 85.
Composition and Laminar Organization of

the Cortex
In the fourth chapter, we study the details and

significance of the laminar composition of the
cortex, and, as a matter of fact, separately of the
isocortex (homotypic and heterotypic) and the
allocortex, particularly the composition and the
details of each layer, as well as the physiologic significance.
We particularly emphasize that, like the total
thickness of the cortex, so the thickness of individual layers varies in different segments of each
gyrus; such a change further varies for each layer.
Thus, e.g. layers I and II are thinnest at the dome
and gradually thicken toward the valley floor,
where they reach their maximum thickness,
whereas layers V and VI are thickest at the dome
and gradually become thinner toward the valley
floor; that thinning is much greater than the respective thickening of layers I and II, such that
the reduction of the overall cortical thickness at
the valley is mostly due to the reduction of the
thickness of layers V and VI. Layers III and IV
retain about the same thickness.
We give the average thickness that we found
for each of the six layers in every part of the gyrus
(fig. 86, 87) and their relative proportions, i.e. the
absolute thickness of each layer and its percentage over the total cortical thickness in the same
part, and all that specifically for each area of the
cerebral cortex. Those numbers are contained in
tables I, II, III and IV in chapter 14 of our Textbook [Economo and Koskinas, 1925]; figure 27 in
the Atlas [Economo and Koskinas, 2008] and fig-
Fig. 83. Median facies of the right cerebral hemisphere (with the diencephalon and mesencephalon dissected out)
of a 3-month-old fetus with a crown-rump length around 6 cm, semi-schematically reconstructed from a series of
sections at an approximate linear magnification of !5. Roman numerals IX and the respective lines denote the
planes of the sections reproduced in figure 84. The dashed line on the median facies hems on the outside around
that part of the hemispheric wall, which develops into the allocortex. The dash-dot-dash line hems around that part
of the median hemispheric wall, at which the two hemispheres, without physically merging, stumble against each
other in the region of the trapezoidal field (Tr.). a = Angle where the dorsal (m.th.) and rostral (m.r.) inner boundaries
of the two hemispheres converge; Bo = olfactory bulb; C.c. = callosal anlage at commissural plate (K); C.i. = section
through the internal capsule and the diencephalon rolling into the cerebrum (telencephalodiencephalic border);
Coa = anterior commissure in commissural plate (K); F = crus fornicis (Fornixschenkel) in commissural plate (K); F.ch. =
choroid fissure as invagination of the medial hemispheric wall in the rostrocaudal direction; F.M. = interventricular
foramen of Monro; K = commissural plate of Hochstetter [1919], the broader, anteriorly-positioned median fusion of
the two cerebral hemispheres; L.i. = infrachoroid layer (lamina infrachorioidea) of His [1904]; L.s. = suprachoroid margin (limbus suprachorioideus) of His [1904]; L.t. = lamina terminalis; m.hy.th. = hypothalamic border (margo hypothalamicus); m.p. = posterior border (margo posterior); m.r. = adhesion border (margo reuniens); m.th. = thalamic border (margo thalamicus) [margo-ines (Latin) = border zone between the two cerebral hemispheres or between telencephalic and diencephalic structures]; opt. = optic chiasma; S = primary temporal pole; S.M. = hypothalamic sulcus
of Monro; Str. = corpus striatum; Tr. = trapezoidal field (Trapezfeld), whereby the medial wall of the hemispheric pouch
is thickened; V3 = third ventricle; Z = diencephalon. Originally figure 65 in Economo and Koskinas [1925].
Appendix
213
Fig. 84. A semi-schematic series of 10 coronal sections (IX) through the 3-month-old embryonic brain depicted in
figure 83. Section I courses through the occipital region, section X through the frontal region. The diencephalon and
mesencephalon are only outlined. a and b = Dorsal and ventral boundary marks of allocortical anlage (corresponding to the dashed line in fig. 83); B.olf. = olfactory bulb; C.c. = callosal anlage; C.i. = internal capsule; Co.a. = anterior
commissure; C.str. = corpus striatum; F = crus fornicis ascendans (aufsteigender Fornixschenkel); F.ch. = choroid fissure;
K = commissural plate of Hochstetter [1919]; L.s. = suprachoroid margin (Limbus suprachorioideus) of His [1904]; L.i. =
infrachoroid layer (lamina infrachorioidea) of His [1904]; L.m. = medullary margin (limbus medullaris); L.c. = cortical
margin (limbus corticalis); L.t. = lamina terminalis; M = matrix; m.hy.th. = hypothalamic border (margo hypothalamicus);
N.c. = caudate nucleus; N.l. = lenticular nucleus; Olf. = lateral olfactory gyrus; Opt. = optic chiasma; Pl.ch. = choroid
plexus; Py. = pyramidal layer or cortical plate (Pyramidenschicht or Rindenschicht); R = optic recess of third ventricle;
RS = marginal veil (Randschleier); S = primary temporal pole (uncus); T = taenia; Tr. = trapezoidal field (Trapezfeld); U =
inferior horn of lateral ventricle; V1 = lateral ventricle; V3 = third ventricle; Z = intermediate zone (Zwischenschicht).
Originally figure 66 in Economo and Koskinas [1925].
214
ure 10 in the present book give a schematic representation of these relations.

We think that such differences, like in principle any anatomic differences, correspond to
physiologic differences, which we discuss extensively in a subsequent section, exposing our theory of the gyrus as an organ.
In examining, as mentioned above, the particular composition of each of the six layers, we
emphasize our following specific observations:
Layer II granule cells in areas near the olfactory brain are larger, stain more intensely and are
arranged in clusters. On the contrary, in the parts
of the cortex that we call koniocortex these cells
are extremely small.
Layer III cells (fig. 88a, b) display most clearly
a radial disposition in the occipital lobe, which is
less marked in the frontal lobe.
Layer IV exhibits its highest density in the
striate area. In the frontal lobe the cells of this
layer are larger and thinner and have a shape that
approaches the pyramidal shape, whereas in the
calcarine area they are smaller and rounder. Layer IV in the temporal lobe displays a radial arrangement, as its cells are disposed in columns of
a high cellular density, clearly delimited from
each other through acellular spokes. We call such
a disposition cleavage (Zerklftung).
Layer V presents such a great cellular density
in the insula, because of cell size and number,
that in stained specimens it appears as a continuous band running through this area, which we
call the insular zone; elsewhere, on the contrary,
e.g. in the calcarine sulcus, the density of this layer is very small, such that it appears as a white
band. The cells of this layer, like those of layer IV,
present from area to area, like e.g. between the
frontal and the occipital cortex, a most striking
difference, which is not found in any of the other
layers (fig. 88c, d). It thus becomes more or less
probable that, although they constitute one and
the same layer everywhere, they nevertheless
have different functions.
Appendix
Layer VI cells are arranged perpendicularly to

the cortical surface at the dome, where the layer
assumes its maximum thickness; the cells are very
small at the walls; lastly, at the valleys, the cells are
most small, scanty, and arranged parallel to the
cortical surface. This particularity of layer VI has
special importance regarding the interpretation
of the overall generation of the gyri; it further confirms our theory of the gyrus as an organ, with its
different parts subserving different functions.
This layer is likely the most developed in humans, forming about 40% of the overall thickness
of the cortex (fig. 86c, d). From a comparison of
the six layers we conclude that, as far as the stability of their presence in the various cortical areas
is concerned, they must be ranked in the order
I VI V III IV II. On the other hand, as
far as the uniformity of their composition in the
various areas is concerned, they should be ranked
in the order I VI III V II IV, although
layers I and VI are those more constantly and
uniformly seen.
In the section on the physiology of the various
cortical layers, having reviewed the conclusions of
cytoarchitectonics and the previously expressed
opinions, we arrive at our own conclusions as follows: large pyramidal cells, mainly of layer V, but
in part also of sublayer IIIc, are the origin of motor pathways; layer VI, which according to the experiments of Nissl [1898] also carries the efferent
impulses, perhaps constitutes the origin of communication pathways between the cortex on the
one hand, and the optic thalamus, red nucleus, or
other deep centers (e.g. pons) on the other. The
view of Meynert [1872a, b] that this layer gives origin to communication pathways between different fields of the cortex must be rejected.
Layer IV is used for the reception of afferent
(sensory) stimuli and the establishment of short
communication pathways in the cortex. Layer III,
on the one hand, subserves higher receptive (sensory) functions, and, on the other, possibly transmits such impulses again to layers V and VI; regarding its long axons, which enter the white
215
216
matter, it subserves higher communication functions among various cortical areas. Layer I principally subserves the connection and propagation within the cortex itself, but to a lesser extent
than layer IV, either within the same gyrus or between neighboring gyri.
From our presentations, it can be concluded
that our attempt to attribute an individual physiologic function to each layer is totally justified.
The integrative function of each cortical area
thus appears to be a sum of the functions of its
various layers. For example, one and the same
motor center may have, besides its motor function, other secondary functions as well, such as
sensory or association functions.
In the same chapter, examining the issue of
the construction of a cerebral cortical map, we
observe the following: between various areas, it
is not only the diverse cortical layers, but also the

different attributes of each single layer that are
modified separately, such that one might not lay
clear-cut boundaries between the various areas;
therefore, their definition comprises a certain
degree of subjectivity and arbitrariness. That is
why an ideal map of the cerebral hemispheres
would be one that depicted all the properties of
all the layers. Such a map might be constructed,
e.g., if we first produced a series of maps, each of
which only depicted one and a single property of
a specific layer, i.e. at least four such properties
per layer, corresponding to the size, number, disposition and density of the cells. We should next
superimpose all these maps upon each other, so
that for each location we would see transparently all the attributes of all the layers. But because
this is not practically feasible, the delimitation of
Fig. 85. a Schematic drawing of the expansion of the agranular isocortex by means of blue shading and hatching.
The left cerebral hemisphere is viewed from its lateral convexity; it has been rendered transparent, with the sulci and
fields of the median facies projected on the convexity in red contours. The agranular cortex of the median facies is
shaded in lighter blue than that of the convexity. (Owing to its almost agranular cortex, the temporal pole is plainly
hatched in blue.) The allocortex, although to a large part likewise agranular, is no longer considered here, and hence
not marked in blue. The stretch of the agranular precentral cortex comes out very graphically in this figure, from the
operculum of Rolando (Op) and from within the central sulcus (RR) over the convexity and the widened configuration dorsally. It continues, over the superomedial hemispheric edge, towards the median facies; here the agranular
cortex occupies in a broad expansion the middle segment of the limbic lobe and its entire anterior segment (L), then
the parolfactory field of Broca (AB). It then courses around the inferomedial hemispheric margin, becoming thinner,
onto the orbital (ventral) surface and next onto the transverse insular gyrus and anterior insula (J), reappearing consequently on the convexity, though without reaching the operculum (Op), i.e. without apparently completing the
ring, but continuing instead from the anterior insular surface along the insular pole (JP) to the posterior of the temporal pole (TP) and from here again to the median surface as a comma-shaped strip (a) in the occipitotemporal sulcus
(ot), forming the allocortical-isocortical boundary and reaching all the way to the isthmus (Js). Furthermore, a thinner,
longer, crescent-shaped strip of agranular isocortex (b) exists in the retrosplenium of the limbic gyrus, completely
detached from the remaining agranular cortex. Originally figure 76 in Economo and Koskinas [1925]. This figure is in
essence a negative of figure 11 in Economo and Koskinas [2008] that depicts the two granular layers. b Ventral facies of the human cerebral hemispheres. Allogenetic cortex (allocortex) is drawn in red. All of the remaining cortex
is isogenetic (isocortex). In particular, parts left white depict ordinary six-layered (hexalaminar) granular cortex, i.e.
homotypic isocortex. Parts painted blue depict the differently structured, non-six-layered heterotypic isocortex. In this
latter category, deep blue areas represent the general granularity that gives rise to the granulous heterotypy of the
koniocortex; blue-colored parts represent the absence of granular layers associated with the agranular isocortical
heterotypy. The temporopolar region (sky blue, area TG ), although six-layered, can be considered as heterotypic isocortex, as layers II and IV become thinner and poorly populated toward the temporal pole. A clear predominance of
isocortex over allocortex is evident in the human brain; the allocortex which is virtually shrouded in heterotypic isocortex hardly occupies 9% of the entire cortical surface. Originally figure 58 in Economo and Koskinas [1925].
Appendix
217
Layer I
thickness
b
Layer VI
thickness
Fig. 86. Schematic depiction of the regional changes of cortical thickness in layer I (a, b) and layer VI (c, d) at the lateral convexity (a, c) and median hemispheric facies (b, d) through gray shading intensity. Thicker territories appear
darker. Originally figures 68, 69, 83 and 84 in Economo and Koskinas [1925].
the various areas from each other is only effected

on the basis of these main elements.
From the cytoarchitectonic study of the various areas it can be concluded that two areas, even
when related, are never exactly the same. On the
other hand, distant areas often present certain
similarities. If we wish to classify together all the
areas that present such similarities, we observe
that they can all be grouped into five general categories, i.e. there are five different general structural types of cerebral cortex. These are:
218
(1) The agranular pyramidal type, comprising

large cells, and as a matter of fact robust pyramidal and spindle cells, and lacking granule cells.
(2) The granular pyramidal (frontal) type,
consisting of well developed pyramidal cells and
comprising two granular layers.
(3) The parietal type, in which the granular
layers are much more distinctly developed, and
the other cells are much smaller.
(4) The polar type, composed of numerous
granule cells, and other features.
Layer III
thickness
b
Layer V
thickness
Fig. 87. Schematic depiction of the regional changes of cortical thickness in layer III (a, b) and layer V (c, d) at the lateral convexity (a, c) and median hemispheric facies (b, d) through gray shading intensity. Thicker territories appear
darker. Originally figures 72, 73, 77 and 78 in Economo and Koskinas [1925].
(5) The granulous type (koniocortex) characterized by the following: (a) the cells of all its layers are small and numerous, giving the impression of dust; (b) immediately beneath the granule cells, i.e. in layer V, it displays an exogenous
dense matrix of fibers, which we should consider as the cortical termination of sensory fibers;
(c) the spindle cells of layer VI often become triangular and pyramidal; and (d) the overall
thickness is smaller than the adjacent areas, because koniocortex is mainly found on the walls
Appendix
of gyri, where, as known, the thickness diminishes. With regard to this type, we emphasize
especially that immediately around the areas
where is appears, one observes spaces of the
agranular type or of the magnocellular intermediate type 1(2) or at least extra large cells, almost
giant cells.
Comparing the subdivision of the isogenetic
cortex into five general types and its above mentioned distinction into homotypic and heterotypic, we note that layers II, III and IV make up
219
Layer III
cell size
b
Layer V
cell size
Fig. 88. Regional variations in a, b average pyramidal cell size in layer III and c, d average nerve cell size in layer V on
the lateral convexity (a, c) and median hemispheric facies (b, d), schematically depicted by means of red coloring
intensity. Magnocellular territories appear darker, and parvicellular territories appear lighter. Originally figures 74,
75, 79 and 80 in Economo and Koskinas [1925]. a, b The sporadic presence of giant pyramidal cells in layer III is further
denoted by solid black triangles (their compactness offering a rough idea of the quantities of giant cells). One readily discerns that such giant cells are not present exclusively in magnocellular, but in parvicellular regions as well. In
general, smaller cells are found in thinner cortical segments of layer III, although fairly large pyramidal cells are individually found in deeper zones. Cortical regions, where layer III is as thin as in the occipital lobe, include areas FF
(plates P34 and P35 in the Atlas [Economo and Koskinas, 2008]) and FG especially at the walls and brinks of the olfactory sulcus (where layer III only occupies 26% of the entire cortical thickness). In the occipital lobe, large pyramidal
cells are present in sublayer IIIc of area OB (plate P85 in the Atlas [Economo and Koskinas, 2008]). c, d The sporadic
presence of giant cells in layer V is further denoted by hollow black triangles (their compactness offering a rough
idea of the quantities of giant cells). One readily discerns that giant cells are also present in certain parvicellular regions. Moreover, the presence of a cellular band emerging from dense arrangements of pyramidal cells in layer V is
indicated by radial lines, whereby the compactness of the lines gives a rough measure of the distinctness of this cellular band (or girdle), most commonly encountered in areas adjacent to the rhinencephalon.
220
the homotypic, while layers I and IV the heterotypic isocortex.

As far as the physiologic significance of these
five structural types is concerned, we uphold the
following views:
(1) The agranular pyramidal type of the frontal cortex has a motor function, but it is doubtful
whether the same happens with the same structural type in other areas.
(2) Concerning the granular pyramidal (frontal) type, which is mainly developed in the middle frontal brain, one could hence accept that it
subserves higher functions related to voluntary
movements, but in such a case it would be difficult to explain the fact that this type is equally
developed in the upper parietal lobe as well.
(3) The parietal type is very well developed in
the lower parietal, anterior occipital, and temporal lobe, i.e. in exactly those areas, whose lesions
disturb sensory perception. It thus appears that
this type subserves the combination of new sensory stimuli with old mnemonic impressions. We
are still ignorant of its function in the frontal
pole.
(4) As far as the polar type is concerned, it is
possible that from the physiologic viewpoint it is
a simple variation of the previous type.
(5) Our knowledge is clearer regarding the
granulous type (koniocortex). This is mainly developed in those areas in which, for physiologic
reasons, we were driven to localize the sensory
centers; it is therefore most likely that it primarily
subserves the reception of stimuli derived from
the sensory organs and carried by subcortical nuclei.
We find it imperative to observe here that, except for structural types 1 and 5, everything said
about the remaining types, owing to the lack of
great precision, only has a so-called heuristic
value.
By juxtaposing our demarcation of cortical
structural types to Flechsigs demarcation of centers, we observe that the primordial centers of
Flechsig [1894, 1920] coincide with those areas
Appendix
which from a cytoarchitectonic viewpoint appear

as heterotypic isocortex or as allocortex.
By examining the allocortex in particular, we
observe that it is not uniform, but rather consists
of three types that are very different from each
other, namely: striate, formed of multiple strata,
like the isocortex, but much more differentiated
than the isocortical layers; rudimentary, which
only comprises two layers, presenting a certain
analogy to isocortical layers I and VI, and occasionally also a third layer, analogous to layer V of
isocortex; and primordial, comprising a single
layer, analogous to layer I of isocortex, and disarrayed huddles of neurons, related in multiple
ways to the subcortical ganglia. The allocortex is
also subdivided into multiple areas, detailed in
the Special Part [Economo and Koskinas, 1925].
Subdivision of the Cortex into Areas
In chapter 5 [Economo and Koskinas, 1925] we

expose the general fundamentals of the subdivision of the cerebral cortex into areas and we also
examine individual variations and their physiologic significance.
We subdivided the cerebral cortex, depending
on the constituent cells of each of its various
parts, into areas that differ from each other from
a cytoarchitectonic aspect, according to the concept of distribution proposed by Meynert
[1867/1868, 1868, 1872a, b] and Betz [1874, 1881],
and effected by Campbell [1903, 1905].
Because anatomic structure presents differences from one brain to another, it follows that an
area can be considered as distinctly defined only
provided that, despite such individual variations,
it is found in every brain.
We based such a distribution on cytoarchitectonic differences alone, but for the naming and
characterization of the areas we took into account, for practical reasons, the macroscopic dispersion of the cortex, so that one would readily
and directly recognize the location of each area.
221
For the classification of the various cytoarchitectonic areas we subdivided the cortex into seven lobes: (1) frontal lobe (lobus frontalis); (2) superior limbic lobe (lobus limbicus superioris); (3)
insular lobe (lobus insulae); (4) parietal lobe (lobus parietalis); (5) occipital lobe (lobus occipitalis); (6) temporal lobe (lobus temporalis), and (7)
hippocampal or inferior limbic lobe (lobus hippocampi or limbicus inferioris).
We further subdivide most lobes into regions,
as follows: the frontal lobe into prerolandic, anterior frontal (prefrontal), and orbital (orbitomedial) regions; the superior limbic lobe into anterior,
posterior, and retrosplenial regions; the parietal
lobe into postcentral (anterior), superior, inferior,
and basal regions; and the temporal lobe into
supratemporal, temporal proper, fusiform, and
temporopolar regions (table 1; cf. also fig. 22c, d
in the Atlas [Economo and Koskinas, 2008]).
In the subdivision of the cerebrum into lobes
we followed the accepted conventions; we only
brought about modifications with regard to certain points, e.g. the parietal and occipital lobes, on
the basis of cytoarchitectonics, and we think that
anatomists in the future will vindicate us, insofar
as it is mostly the microscopic study that is bound
to solve many disputed, or at the very least vague,
issues of the macroscopic anatomy of the brain.
We depicted the distribution of cytoarchitectonic areas in the cerebral cortex in figure 1ad,
(cf. also fig. 2, 25 and 26 in the Atlas [Economo
and Koskinas, 2008]) which correspond to the
lateral, median, dorsal and ventral facies of the
cerebral hemispheres, respectively. One first discerns in them the larger ground cytoarchitectonic areas (Grundareae), and next the most important consistently observed differences, i.e. the
variants (Varianten), as well as the also consistently observed, albeit of lesser importance,
modifications (Modifikationen). We furnish
greater details under the specific description of
each cytoarchitectonic area.
Thus, we discerned about 54 ground cytoarchitectonic areas; by adding the variants, the to-
222
tal number increases to 76, and with the modifications we arrive at 107 cytoarchitectonic areas.
For the symbol notation of each area we did
not use the hitherto granted system of numbers,
which are arbitrary and say nothing, but we denoted each area by the initial capital letter of the
Latin name of the lobe in which it lies, to which
we add another capital in corsiva, which, depending on its sequence in the alphabet, denotes the
sequence of an area within a lobe. For example,
FB means the second ground area in the frontal
lobe. We further denoted the variations of areas
with the above symbols, where each one belongs,
adding a Latin or Greek subscript, which indicates a characteristic feature (e.g. m = magnocellular, p = parvicellular, = giant cellular).
Of the 107 cytoarchitectonic areas mentioned,
22 belong to the allocortex, another 22 to the heterotypic isogenetic cortex, and the remaining 63
to the homotypic isogenetic cortex. As far as the
boundaries between any two cytoarchitectonic
areas are concerned, we notice, on the one hand,
that they are clear between the isocortex and the
allocortex, whereas they become less distinct between homotypic and heterotypic isogenetic cortex; finally, among the various isocortical areas,
one observes gradual transitions instead of distinct boundaries.
We also observed that these boundaries are in
principle independent of the cortical sulci; nonetheless, there exist certain relations between the
extension of some cytoarchitectonic areas and
the size and shape of the respective gyri that we
still cannot understand clearly today.
Due to the lack of a correspondence between
the boundaries of cytoarchitectonic areas on the
one hand and the sulci of the cortex on the other,
investigators prior to us thought that only cytoarchitectonics mattered, whereas the gyri, according to them, a purely random result, should
not in the least be taken into account.
Conversely, based on the relation already mentioned between the cytoarchitectonic structure
on the one hand and the configuration and course
of the gyri, as well as the three different segments

of each gyrus (dome, walls, valleys) on the other,
we arrived at the following conclusion: every anatomic difference corresponds, as it is well known,
to a difference in function; since the various segments of a gyrus have a different structure, we
must accept that they also have a different function; therefore, the gyrus must not be considered
as a random configuration, but as constituting a
uniform ensemble, an organ so to say, each segment of which has an individual structure and
function. Thus, e.g. at the dome, layers V and VI
(the inner main zone of Kaes [1907] or internal
fundamental layer of Jakob and Onelli [1911]) prevail, which investigators unanimously consider as
the origin of efferent pathways (i.e. motor), whereas at the walls of the gyri these layers become reduced, while at the same time layers I and II (the
outer main zone of Kaes [1907] or external fundamental layer of Jakob and Onelli [1911]) become
reinforced, being considered as the termination of
afferent (i.e. sensory) pathways.
Our theory is further supported by the fact
that our so-called koniocortex, to which everybody attributes a sensory function, is mainly
found at the walls of gyri.
The study of the cytoarchitectonics of the cerebral cortex can contribute to the solution of
problems of comparative anatomy, e.g. the definition of the correspondence among the brain areas of different animals (definition of homologous areas) and vice versa; as certain parts are
developed in certain animals, comparative cytoarchitectonics can elucidate some problems of
the physiology of the human brain.
In a separate section on the anatomic and
physiologic significance of the areas we express
the opinion that, with the continuation of these
researches, the differences in cytoarchitectonic
structure must be studied depending on age, gender, intelligence and overall mental gifts of various individuals, and finally depending on race,
taking into account, in that case, mental variations among the various races.
Appendix
Although the main aim of our work was to

study the cytoarchitectonic structure of the cerebral cortex, we nevertheless did not neglect the
physiologic aspect of the issue. It is indeed accepted in general, one could even say that it is obvious
in advance, that the anatomic and histologic differences correspond to physiologic differences,
otherwise they would not have any reason for occurring. For that reason, following the description of the various layers, we expose in our work,
as we have seen above, their possible differences
from the physiologic aspect as well. We do the
same also when speaking of the five different cortical structural types, which we defined ourselves.
It was only natural then, that we attempted to do
the same regarding our defined areas as well.
It is known that various investigators, based
on both the experimental and the neuropathologic evidence, localize in different areas of the
cortex various functions, thus defining so-called
centers. Such localizations still present, in some
points, a certain relative vagueness. Although
certain functions are localized by everyone to almost the same areas (e.g. the famous language
centers), the same does not happen with other
functions; on the contrary, one and the same
function is often localized by different investigators in different areas and, vice versa, various authors localize diverse functions in one and the
same area.
Regarding this point, we emphasize in our
work the difficulty inherent in the relatively
vague distinction of the various functions. Indeed, we cannot be certain a priori that an ensemble of phenomena that we name in general
language with one and the same term, e.g. memory, corresponds to a uniform ensemble from a
physiologic viewpoint as well. Nothing proves,
e.g., that the preservation of visual and auditory
impressions corresponds to the same histologic
structure. It is possible that these two modalities
of perception are registered in the brain in two
different ways and thus claim two different histologic structures.
223
Due to the vagueness regarding localization,

having taken into account the suggestions of different investigators, we compiled, for the sake of
our own work, an independent schematic drawing of the cerebral hemispheres, a functional map
so to say, which, within the realms of possibility,
depicts on the one hand more generally accepted
localizations (the average in a sense), and on the
other, variations according to different researchers (cf. fig. 18 in the Atlas [Economo and Koskinas, 2008]).
If one juxtaposes against this drawing our
main cytoarchitectonic area map (fig. 1ad), one
gets the impression of a relatively meaningful
correspondence of the two.
The main and clearest case of such an analogy,
on which we particularly insist in our work, is
that of the koniocortex. As already mentioned,
this type of cortex is mainly found in the following five regions: (1) the anterior wall of the postcentral gyrus; (2) the transverse gyrus of Heschl;
(3) the wall and lips of the calcarine sulcus; (4) the
retrosplenial area of the limbic gyrus, and (5) the
inner wall and posterior lip of the hippocampal
gyrus.
As is generally known, the sensations of touch,
hearing and vision were localized in the first
three of these regions, and the sensations of olfaction and taste, with lesser certainty, in the last
two. Based on these facts, which we examine in
our work in detail, we were led to the conclusion
that the koniocortex constitutes the highest special adaptation of the cerebral cortex for receiving sensory stimuli. Our conlusion is in a relative
agreement with the opinions expressed by
Meynert [1872a, b] and Flechsig [1895, 1897,
1920]. However, we cannot deny a priori that additional cortical areas, in which such a granulization which constitutes the koniocortex is not
observed, subserve a similar goal; as a matter of
fact, for certain cortical areas, immediately adjacent to koniocortex, this is fairly probable.
We emphasize again that, because each area
comprises different layers, to each of which, as we
224
have seen, a particular function has to be assigned, we may ascribe to each area a certain
function as primary and dominant, but we cannot exclude the possibility that the same area
might have other secondary functions, what is
more, to the extent that, as we have seen, each
segment of the same gyrus has a relatively separate physiologic significance. That is why we
must be very cautious every time we characterize
a certain area as the center for a specific function,
which is only approximately true.
We compare the conclusions on the physiology of the areas to the views expressed by Flechsig
[1894, 1920], who distinguished three types of areas, depending on the time period over which
myelinated fibers make their appearance. Flechsig, as already known, distinguished the primordial centers (Primordialzentren) that already
contain fibers during embryonic life, the intermediate fields (intermedire Gebiete), whose
white matter develops during the first month of
extrauterine life, and the terminal fields (Terminalgebiete), in which the myelin of nerve fibers
develops even later.
The first of these centers, which coincide with
our cortical structural types 1 and 5, he considered as subserving the reception of sensory impulses. In the second, he thinks that the mnemonic images of sensory impressions are stored.
Lastly, the third he considers as subserving the
association of various representations; Flechsig
[1920] subdivides those into three parts, to which
he ascribes various components of intelligence:
he considers the posteriorly located parietotemporal association field, interposed between the
sensory domains, as subserving the association
of sensory representations, i.e. as the seat of what
we call positive knowledge or spirit (Geist); the
insular association center as pertaining to speech
and language (Sprache); and the frontopolar association field as constituting the memory domain of conscious bodily perceptions, emotions
and will, i.e. mainly the center of the so-called
ego (Ich).
In chapter 4 of our work [Economo and Koskinas, 1925] we examine to what extent such a distinction is in agreement with the broader cytoarchitectonic findings. Many debated, at times
justly, and at times unfairly, those views of Flechsig. It is true that the existence of self-contained
association centers has an element of improbability, insofar as it is more likely that association is a
function which occurs in all parts of the brain.
Anyway, we must accept that those areas, which
both develop later than others from the ontogenetic viewpoint and have evolved later from the
phylogenetic viewpoint and, moreover, are mostly found exclusively in humans, are apparently
related to higher intellectual functions.
Another conclusion, drawn from our cytoarchitectonic and physiologic observations, is that,
on the one hand, the part of the brain lying in
front of the central sulcus is mostly of a motor
nature, and, on the other, the part lying behind
the central sulcus is, reversely, mostly sensory,
and in it the impressions from the senses are
stored.
Method
In chapter 6 [Economo and Koskinas, 1925], we

expose the method adopted for our work. That
method is not totally new, but consists of the
method of formalin, ethanol, paraffin and toluidine blue staining, to which we added certain
necessary modifications, for the sake of perfection.
We expose these methods in our work, so that
anyone wishing to undertake a similar project, by
following them, would obtain results absolutely
comparable to ours, owing to the unity of the
method. The modified method is as follows:
After the removal of the pia mater we place the
brain into a 5% formalin solution. After about
12 h, by appropriately rotating the brain in the
formalin, we photograph separately all of its surfaces. About 12 h later, we remove the brain from
Appendix
the formalin, we wrap it in cotton sopped in the

same formalin solution and dissect blocks of a
thickness of about 4 mm, through cuts perpendicular to the surface of the gyri, exposing every
time only the segment necessary for obtaining
sections. Sometimes, due to the configuration of
the gyrus, the cut, in order to be perpendicular
not only to the dome, but also to the walls and the
valleys of the gyrus, must not be flat, but curviform, observing the form of the gyrus. We compress the curviform blocks between two glass
slides wrapped in absorbent paper, so that they
would become flattened, and we strap the whole
preparation. Thus, each hemisphere is dissected
into about 250350 blocks. Each block is numbered and the number is marked on the corresponding position in the respective photograph.
The blocks, during their stepwise dissection, are
placed three at a time into wide-neck vials containing the same formalin solution. Such a dissection requires about 12 h per hemisphere.
Twelve hours after the dissection, we decant
the formalin from the vials and replace it with
H2O, which we change 3 times over a total period
of 6 h. After that period of time has passed, we
replace H2O with 50% ethanol. Every 24 h, we replace the ethanol with ascending concentrations
70, 80 and 95%. We leave the blocks in the last
solution for 72 h, but with changes every 24 h.
After that we place them in absolute ethanol for
24 h, with one change in between. We next replace the ethanol by xylene, leaving the blocks in
it for 5 h, such that they become transparent. The
xylene must also be changed once during that
time period. Afterwards, decanting the xylene,
we fill the vials with chemically pure fluid paraffin, in which the blocks can be preserved unaltered indefinitely.
Thus, all of the blocks were subjected to the
action of the active fluids for the same amount of
time, and having since remained in the liquid
paraffin unaltered, they may be used at any time.
Subsequently, we remove any number of such
blocks and place them into an oven under a tem-
225
perature of 54 C in paraffin of a melting point

successively of 42, 48, and 52 C for 2 h each.
Then, removing them, we embed them in the last
type of paraffin. From each of these blocks we
obtain in a microtome sections of 25 m in thickness. We do not mount the sections, as it was
done before us, on glass slides by means of albumin, but we place many of them together in a cup
containing xylene to remove the paraffin. After
about 1 h we place them successively in ethanols,
100, 90, 80, 70, 60, 50, 40, 30, 20 and 10% for 10
min each, and lastly in distilled H2O for 1 h. Then
we place them in a solution of 0.3 g toluidine blue
in 3,000 ml distilled H2O, where they remain for
12 h at room temperature.
We next place them for 10 min in distilled
H2O and then in a mixture of 10 parts aniline oil
and 90 parts of 95% ethanol for discoloration. By
frequently changing the solution, we watch the
discoloration of each section under the microscope at a low magnification, until the ground
substance becomes totally discolored, and within
it the nerve cells are discerned as vividly stained
blue. As soon as that result is achieved, we bring
the sections successively in 95% and absolute ethanol for about 2 min each, with repeated changes
in each, and then in Cajeput oil for 510 min. Finally, we mount each section on a glass slide, and
we cover them with cedar oil and a coverslip.
Through this method, although it takes long,
we achieve the optimal results, because all blocks
and sections are subjected to the action of the fluids for the same amount of time, which is necessary for the determination of the size of cells and
the thickness of the layers and the cortex. Further, by achieving such a uniform staining, it becomes possible to compare specimens from different parts of the cerebral cortex and to prepare
uniform microphotographs as well.
226
We selected the thickness of 25 m ourselves

after many trials, because we found it to be the
most appropriate, as at that thickness one can
most clearly discern the layer arrangement of
cells, without obstructing the observation of the
main details of their structure.
For microphotography, following repeated
trials, we selected the most appropriate light filters, the most appropriate Hauffs Flavin photographic plates, pyrogallic acid for the development, which precipitates the silver in very fine
grains, such that even the minutest details can be
discerned, and for printing we used the best paper available.
Special Part
A detailed analysis of the Special Part (chapters

714), consisting of 550 pages in great octavo
[Economo and Koskinas, 1925], is not possible
here, owing to its nature and size, as anyone can
understand. Indeed, this part includes a detailed
description of each area separately. Most of the
material displayed in it is a result of our personal
studies. That is why I herein content myself with
presenting the layout of the description of each
area.
For each area we sequentially mention: (1) its
macroscopic view; (2) its microscopic view; (3)
arithmetic relations of the various layers to each
other; (4) the description of each layer, i.e. sequentially, the molecular, external granular, pyramidal, internal granular, ganglionic and spindle cell layers; (5) the general picture, extent, limits and variations of the area; (6) its history and
its histologic appearance, and (7) the physiologic
observations on all its cellular elements.
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Authors note: The comprehensive

list of 120 references in our larger work
[Economo and Koskinas, 1925] rids me of
the necessity to repeat them. I only mention here important early reviews, as well
as newer publications, many of which
appeared after our larger work was written and which I had not cited before.
with the 120 references found in the larger Textband. It further comprises Economos individual and subsequent studies
in cytoarchitectonics, as well as additional pertinent references, bringing the
total number to over 300, with full citation information. An extra care was expended on tracing the original articles.
Reference style was modified to conform to current conventions, supplementing full and emended book or journal titles, volume numbers, and inclusive
pagination. Such an attempt falls within
the scope of science as a continuous process, whereby bibliographic resources

should not be artificially interrupted by
the mere technical limitations of the
means at hand: oftentimes, modern research tends to take into consideration
only work published within the last few
years or decades at the most. Thus, this
section constitutes an essential list, covering the most important works on cytoarchitectonics from the 19th century up
to 1931, not readily found in the current
digital databases.
Editors note: The presented bibliography combines the 80 references cited by Economo in the German, French,
Italian and English versions of the book
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List of Persons
Roman numbers refer to the text body or tables of the cited pages; italic numbers refer to the figures on
the cited pages.
Aeby, C.T. 149, 151, 227

Actuarius, J.Z. 195
Alcmaeon of Croton IX, XV
Alzheimer, A. XI, 179
Arnold, F. 156, 159, 227
Baillarger, J. 1, 13, 227
Brny, R. 112, 227
Bekhterew, V. 13
Bertrand, I. 29, 178, 227
Betz, W.A. 1, 2, 14, 14, 35, 36, 63, 75, 76, 80, 83, 178, 181,
199, 203, 221, 227, 228
Bogaert, L. van VIII, XII, XIII, XV, XVI, 29, 178, 227
229
Bolton, J.S. XX, 228
Bonin, G. von IX, XIV, XV
Brissaud, . 83, 228
Broca, P.-P. 24, 31, 37, 44, 53, 54, 54, 5557, 64, 66, 67,
133, 141, 147, 160, 185, 228
Brodmann, K. IX, X, 1, 13, 15, 16, 199, 228
Burdach, K.F. 133, 228
78, 80, 81, 83, 85, 87, 88, 90, 94, 96, 98, 100, 103, 106,
108, 108n., 109, 111, 112n., 113, 117, 119, 121, 123,
124, 126, 127, 129, 130, 132134, 139, 139, 140149,
151, 157, 158, 160163, 165, 167, 169173, 173n., 174,
175, 177, 182186, 189, 189n., 190, 191, 193, 193,
194n., 199201, 202, 203, 204, 205, 205208, 209,
210, 211, 212, 213, 214, 217220, 221, 222, 224226,
227n., 228231
Elliot Smith, G. XX, 1, 102, 112, 189, 199, 229, 230
Enderle, C.A. XV, 229
Erasistratus of Chios 195
Feuchtwanger, E. 185, 230
Flechsig, P.E. 69, 92, 123, 137, 139, 221, 224, 225, 230
Foville, A.-L. 133, 230
Franceschi, F. 29, 230
Freud, S. IX
Fritsch, G. 1, 199, 230
Danilewsky, B. 28, 228

Dotto, G. 29, 228
Galen of Pergamon 194, 195

Gall, F.J. 1, 195, 230
Gans, A. 179, 230
Garrison, F.H. XII, XVI
Gennari, F. 13, 14, 102, 108, 110112, 230
Gerdy, P.-N. 133, 230
Giacomini, C. 17, 151, 154, 158, 160, 164, 170, 171, 230
Goldstein, K. IX, XV
Goldstein, M. 2, 230, 232
Ecker, A. 106, 112, 133, 228

Economo, C. von II, IV, VIIIXX, 2, 2, 3, 4, 6, 10, 14,
16, 17, 18, 20, 21, 25, 26, 2830, 3537, 41, 44, 47, 49,
51, 53, 55, 57, 59, 60, 63, 64, 67, 67n., 69, 72, 74, 75
Hammarberg, C. 1, 14, 29, 137, 180, 199, 230

Head, H. 184, 230
Henneberg, R. 28, 230
Henschen, S.E. 69, 231
Calleja, C. 203, 228

Campbell, A.W. XX, 1, 9, 199, 221, 228
Cruchet, R. IX
235
Herv, G. 69, 231

Heschl, R.L. 24, 119, 120, 122, 123, 178, 183, 185, 224,
233
His, W. 26, 29, 213, 231
Hitzig, E. 1, 199, 230
Hochstetter, F. 213, 214, 231
Horn, L. X, XV, 10, 119, 121, 123, 124, 130, 175177, 185,
229, 231
Huntington, G. 29, 179
Jger, R. 201, 231
Jakob, A.M. IX, XV, 178, 179, 231
Jakob, C. IX, XV, 13, 29, 223, 231
Kaes, T. 13, 29, 223, 231
Kallius, E. XI, XVI
Kleist, K. 112, 231
Klliker, R.A. von 203, 231
Klpin, O. 29, 231
Korsakoff, S.S. 179
Koskinas, G.N. VIIIXIII, XV, 2, 2, 3, 3n., 4, 5, 6, 10, 14,
17, 18, 20, 21, 25, 28, 30, 35, 36, 37, 40, 41, 44, 47, 49,
51, 53, 55, 57, 60, 63, 64, 67, 67n., 69, 72, 75, 77, 78,
80, 81, 83, 85, 87, 88, 90, 94, 96, 98, 100, 103, 106,
108, 109, 111, 113, 117, 119, 121, 123, 124, 126, 127,
129, 130, 132, 134, 139143, 145, 146, 157, 158, 160
163, 165, 167, 169, 170, 172, 173, 174, 193, 194, 194n.,
199201, 202, 203, 204, 205, 205208, 209, 210, 211,
212, 213, 214, 217220, 221, 222, 224226, 227n., 229,
231
Lancisi, G.M. 139, 231
Lenz, G. 112, 231
Levi, G. 152, 231
Lewis, W.B. 1, 212, 231
Lewy, F.H. 29, 231
Marburg, O. IX, XV, 232
Marinesco, G. 2, 14, 139, 232
Meynert, T.H. 1, 2, 14, 54, 57, 108, 110, 111, 173, 199,
203, 215, 221, 224, 232
Mingazzini, G. VIII, IX, XV, 229
Mironesco, T. 2, 232
Monro, A. 213, 214
Morton, L.T. XII, XVI
Mott, F.W. 13, 232
236
Neubrger, K. XI, XVI

Nikitin, M.P. 14, 139, 232
Nissl, F. 11, 13, 13, 14, 29, 35, 76, 103, 107, 111, 170, 190,
215, 232
Onelli, C. 13, 29, 223, 231
Osborn, H.F. 189, 191, 232
Parker, S. IX, XV, XVI, 229
Pfeifer, R.A. 123, 232
Pick, A. 179
Pusateri, E. 29, 228
Ramn y Cajal, S. 9, 13, 14, 110, 139, 172, 199, 203, 211,
232
Reil, J.C. 19
Reiner, C. XIV, 174
Retzius, A.A. 148, 160, 161, 166, 171
Retzius, G.M. 9, 13, 148, 171, 211, 232, 233
Rolando, L. (see central sulcus and operculum)
Rose, M. IX, XV, 10, 158, 167, 233
Rossi, I. 29, 233
Roussy, G. 29, 233
Schwalbe, G. 130, 133, 233
Sterzi, G. 158, 233
Sylvius, F. de le Bo (see lateral fissure)
Talairach, J. X, XV
Tarin, P. 151, 233
Tilney, F. 189, 233
Tournoux, P. X, XV
Turner, W. 133, 233
Vicq dAzyr, F. 1, 13, 14, 102, 234
Villiger, E. 17, 234
Vogt (ne Mugnier), C. 2, 15, 199, 234
Vogt, M. 229
Vogt, O. 1, 2, 13, 15, 178, 199, 229, 234
Volkmann, U. 112, 234
Wagner, H. 28, 234
Wagner-Jauregg, J.R. von IV, V, IX
Weigert, C. 9, 13, 80
Zeiss, C. 200
Subject Index
Roman numbers refer to the text body or tables of the cited pages; italic numbers refer to the figures on
the cited pages.
Acalculia 92, 183

Acoustic radiation (Pfeifer) 123, 124
Adhesion border (margo reuniens) 213
Agnosia 197
Agraphia 92
Alexia 92, 183
Alveus 153, 166, 170
Alzheimer disease XI, 179
Amaurotic idiocy, familial 178
Ammons horn 15, 27, 151, 152, 153, 167, 170, 172, 178,
179
Amygdala, nucleus of (NA) 95, 132, 151, 156, 160, 164
Amyotrophic lateral sclerosis 29, 36, 178, 180
Anterior commissure 54, 55, 214
Aphasia 90
amnesic 94, 127
motor 187
sensory 183, 187
Apraxia 90, 92
Area, cortical (Brodmann) IX, X, 199
Area, cortical (Economo-Koskinas) 25, 68, 10, 11, 30
agranular anterior limbic LA [EK 3638] 134, 141,
142, 147
agranular anterior limbic, posterior cingulate
transition LAC 11, 168
agranular frontal FB [EK 4] 36, 37, 40, 41, 42, 44, 46,
64, 67, 81, 83, 85, 90, 142, 178, 184, 186
agranular hippocampotemporal TH [EK 89] 129,
130, 153, 163
agranular insular frontal FB I 10, 44
agranular intermediate frontal transition FBC 10,
63
agranular orbital FF [EK 21] 38, 51, 53, 59
agranular prefrontal FH [EK 21] 10, 51, 53

agranular retrosplenial LD [EK 44] 138, 141, 142,
144, 145, 147149, 162, 166, 168
agranular temporopolar TG [EK 91] 130, 152, 154,
155, 162, 163
angular PG [EK 69] 79, 88, 90, 91, 92, 187
anterior cingulate agranular anterior limbic LA2
[EK 37] 134, 135, 137, 139, 140, 147, 168
anterior peristriate OA2 [EK 73] 104
anterior superior temporal TA2 [EK 81] 118, 119
anterior ultracingulate LB1 [EK 39] 135, 136, 139,
141, 142, 144, 146148, 166, 168
basal temporooccipital parietal PH [EK 7072] 79,
88, 92, 93, 94, 114, 126, 127, 166, 187
basal (temporooccipital) parietal at occipital
entrance PH O [EK 72] 94
basal (temporooccipital) parietal at parietal
entrance PH P [EK 70] 94
basal (temporooccipital) parietal at temporal
entrance PH T [EK 71] 94
caudal postcentral PD [EK 60] 74, 78, 8385, 84, 90,
189
caudal postcentral transitions PDE and PFD 10, 84,
85
cingulate agranular anterior limbic limitans LA3
[EK 38] 134, 135, 137, 139, 147, 168
dentate HF [EK 107] 138, 146, 147, 150, 152, 153,
154, 157, 158, 164, 166, 171, 172
dorsal posterior cingulate LC1 [EK 41] 140, 142144,
168
dorsal precentral insular IA1 [EK 49] 96
frontal piriform FK [EK 29] 39, 53, 57, 59, 60, 61, 62,
96, 100, 101, 132
237
frontoinsular FJ [EK 28] 39, 53, 57, 58, 59, 59, 60, 61,
62, 95, 100, 101, 129, 137
frontopolar FE [EK 18] 38, 47, 48, 49, 51, 63, 72, 137,
186, 187
fusiform TF [EK 87] 94, 115, 118, 127, 128, 129, 154,
163
geniculate FM [EK 33] 39, 53, 54, 5557, 59, 61, 62,
137, 142
geniculate of olfactory triangle FMt [EK 34] 57
giant pyramidal intermediate postcentral PC 10, 83
giant pyramidal parastriate boundary OB [EK 77]
25, 107, 108, 109, 112, 113, 220
giant pyramidal postcentral PA1 [EK 55] 10, 74, 75,
75, 7678, 80, 83, 189
giant pyramidal posterior superior parietal PE [EK
64] 87
giant pyramidal postparacentral PA2 [EK 56] 74,
7678, 83, 189
giant pyramidal precentral FA [EK 2] 33, 34, 35, 36,
63, 75, 76, 178
glomerular presubicular granulosa HD3 [EK 102]
150, 151, 153, 154, 165, 167, 169
granular frontal FD [EK 11] 37, 38, 44, 45, 46, 47, 51,
67, 69, 137, 185, 186, 189
granular orbital FF [EK 20] 38, 47, 51, 53, 57, 59, 59,
72
granular rhinal limitans HCg 11, 163
hippocampotemporal TH [EK 88] 115, 129, 130, 152,
164, 166, 169
hippocampotemporal piriform transition TJ H [EK
88] 11, 151, 158, 160, 164
indusium griseum LB2 [EK 40] 135, 136, 138, 139,
141, 142, 144, 146150, 166, 168
inferior retrosplenial granulosa LE2 [EK 46] 138,
141, 145, 146, 148, 149, 150, 151, 166, 168
inferior temporal proper TE2 [EK 86] 126
intercalated supratemporal TD [EK 84] 11, 115, 117,
119, 121, 123, 124, 185
intermediate frontal FC [EK 6] 37, 41, 43, 44, 46, 64,
81, 137, 185, 186
intermediate insular frontal FC I [EK 9] 44
intermediate postcentral PC [EK 59] 74, 77, 78, 81,
82, 83, 189
internal straight FGi [EK 24] 49
limbic frontopolar FEL [EK 19] 72, 129, 137, 141
limbic granular frontal FD L [EK 15] 64, 69, 71, 129,
135, 137, 141
limbic intermediate frontal FC L [EK 7] 64, 69, 129,
137, 141, 142
limbic prefrontal FHL [EK 27] 72, 129, 137, 141
maculae granulosae of parastriate OB [EK 78] 113
magnocellular agranular intermediate frontal FCBm
[EK 8] 37, 44, 64, 66, 185
magnocellular peristriate OAm [EK 75] 10, 106, 107
238
magnocellular granular frontal FDm [EK 12] 63, 69

magnocellular granular frontal at beginning of
intermediate frontal FDm(C) 10, 63
magnocellular granular frontal at beginning of
frontopolar FDm(E) 10, 63
magnocellular superior parietal PEm [EK 62] 85
magnocellular superior temporal TAm (EconomoHorn) 10, 119
magnocellular supramarginal PFm 10, 90
magnocellular supratemporal simplex TB [EK 82]
10, 115, 117, 119, 120, 121, 123
marginal superior temporal TA2 (Economo-Horn)
10, 119
middle granular frontal FD [EK 16] 38, 69, 70, 187,
189
middle presubicular granulosa HD2 [EK 101] 150,
151, 153, 154, 165, 167, 169
middle temporal proper TE1 [EK 85] 125
opercular agranular frontal FBop [EK 5] 64, 65
opercular agranular intermediate frontal FBCop
[EK 5] 10, 64
opercular granular frontal FDop [EK 14] 72
opercular intermediate frontal FCop [EK 10] 72
opercular intermediate granular frontal FCDop 10,
72
opercular precentral FAop [EK 3] 63, 64
opercular supramarginal PFop [EK 67] 90
oral postcentral PB [EK 5758] 74, 77, 78, 80, 81, 83,
189
oral postcentral granulosa PB1 [EK 58] 80
oral postcentral simplex PB2 [EK 57] 80
orbito-insular IC [EK 53] 59, 96, 100, 101
parastriate OB [EK 76] 102, 103, 106, 107, 107, 109,
109, 112, 113, 166, 192
parauncinate HB [EK 9798] 130, 152, 154, 155,
156158, 161163, 164, 170, 172
parolfactory FL [EK 3032] 39, 53, 54, 54, 5557, 59,
61, 137, 141
parolfactory prefrontal FHL [EK 26] 54, 55, 141
parvicellular granular frontal FDp [EK 13] 63, 69
parvicellular superior parietal PEp [EK 63] 85
periamygdalar piriform TJ H14 (Economo) 11, 151,
160, 229
peristriate OA [EK 7375] 102104, 105, 106, 113,
166, 192
piriform insular ID [EK 54] 59, 96, 100, 101, 132
postcentral insular IB [EK 51] 96, 98, 99, 129
postcentral insular at temporal entrance IB T [EK
52] 100, 124
posterior cingulate LC [EK 4143] 138, 140, 142
145, 147, 166
posterior cingulate limitans LC3 [EK 43] 136, 140,
142144, 147, 168
posterior peristriate OA1 [EK 74] 104
posterior superior temporal TA1 [EK 80] 116, 118,

119
posterior ultracingulate LF1 [EK 47] 136, 138, 144,
146150, 152, 166, 168, 171
precentral FA [EK 1] 32, 3537, 41, 46, 184, 186
precentral insular IA [EK 4950] 96, 97, 98, 129
precentral insular, postcentral transition IAB 10, 98
precingulate agranular anterior limbic LA1 [EK 36]
134, 135, 137, 139, 140, 147, 168
precommissural FN [EK 35] 39, 54, 55, 57, 59, 60, 61,
61, 62, 132, 142
prefrontal FH [EK 25] 39, 51, 52, 53, 54, 54, 56, 57,
59, 72, 137
presubicular granulosa HD [EK 100102] 152, 153
155, 157, 158, 162, 163, 164, 165, 166, 167, 169172
presubicular granulosa limitans HD1 [EK 100] 150,
151, 153, 154, 165, 167, 169
presubicular granulosa pars anterioris HDa (Rose)
11, 167
presubicular granulosa pars posterioris HDp (Rose)
11, 167
pretriangular orbital FF [EK 22] 72, 220
primary parauncinate HB1 [EK 97] 162
primary parolfactory FL1 [EK 30] 54, 55, 56
primary rhinal limitans HC1 11
primary uncinate HA1 [EK 94] 158
pyramidal hippocampal HE [EK 103106] 138, 146,
147, 152, 157, 158, 164, 165, 166, 167, 169, 171
pyramidal of Ammons horn HE2 [EK 105] 150, 152,
153155, 164, 168171
pyramidal of digitate gyrus of uncus HE3 [EK 106]
154, 155, 164, 170, 171
quaternary pyramidal HE4 (Economo) 11, 171
quinary pyramidal HE5 (Economo) 11, 171
recta (see straight gyrus)
retrosplenial granulosa LE [EK 4546] 144147,
162, 165, 166, 167
rhinal limitans HC [EK 99] 150, 152, 153, 157, 158,
162, 163, 164, 165, 166, 167, 170
rhinal limitans, presubicular granulous transition
HCD 11, 154, 155, 163, 164, 165, 166
secondary parauncinate HB2 [EK 98] 162
secondary parolfactory FL2 [EK 31] 54, 55, 56
secondary uncinate HA2 [EK 95] 158
straight gyrus FG [EK 23] 39, 49, 50, 51, 53, 59, 72,
129, 186, 220
striate (granulosa) OC [EK 79] 102, 103, 105, 106,
108, 109, 109, 110, 111113, 123, 166, 192
subicular glomerular pyramidal HE1 [EK 103] 153,
154, 169
subicular pyramidal HE1 [EK 103104] 150, 152,
153, 155, 164, 168171
subicular pyramidal simplex HE1 [EK 104] 153,
154, 169
Subject Index
substantia perforata TK [EK 93] 59, 60, 61, 61, 100,

115, 132
superior parietal PE [EK 6164] 74, 79, 85, 86, 87,
142, 189
superior retrosplenial granulosa LE1 [EK 45] 138,
141, 145, 146, 148, 149, 151, 166, 168
superior temporal TA [EK 8081] 115, 116, 117119,
126, 130
superior temporal in supratemporal gyrus TA2
(Economo-Horn) 10, 119
supramarginal PF [EK 65] 10, 79, 85, 88, 89, 90, 92,
115, 118, 119, 187
supramarginal columnata PFc 10, 90
supramarginal-postcentral transition PFD 10, 90
supratemporal granulosa TC [EK 83] 11, 115, 117,
119, 121, 122, 124, 185
temporal piriform TJ [EK 92] 11, 59, 115, 132, 151,
160, 164
temporal proper TE [EK 8586] 115, 117, 124, 125,
126, 127
temporopolar TG [EK 90] 11, 115, 118, 127, 129, 130,
131, 156, 162, 163, 164, 169, 216
tenuicortical supramarginal PFt [EK 66] 90
tertiary parauncinate HB3 11, 162
tertiary parolfactory FL3 [EK 32] 54, 55, 56
tertiary uncinate HA3 [EK 96] 158
triangular granular frontal FD [EK 17] 38, 67, 68,
69
ultracingulate obtecta LF2 [EK 48] 136, 138, 144,
146150, 152, 166, 168, 171
uncinate HA [EK 9496] 130, 152, 155, 155, 156,
156, 157, 158, 159, 160162, 164, 170, 172
ventral posterior cingulate LC2 [EK 42] 136, 140,
142144, 168
ventral precentral insular IA2 [EK 50] 96
Asemantic defects 184
Association centers (Flechsig) 25, 69, 181, 190, 224, 225
Association fibers 25, 26
Ataxia, static 127
Ataxic disturbances 49
Attention 47, 121, 185, 187
Auditory domain 123, 124
Autonomic nervous system 101
Band of Giacomini 17, 151, 154, 158, 160, 164, 170, 171
Basedow disease 197
Betz giant cells 1, 2, 14, 14, 35, 36, 63, 75, 76, 80, 83, 178,
181, 201, 203, 206
Brocas area 37, 44, 64, 66, 67, 67n., 185
Brocas diagonal band 57, 59, 61
Brocas grand lobe limbique VIII, 31, 133
Brocas parolfactory field (carrefour olfactif) 24, 51, 53,
54, 54, 55, 56, 133, 141, 147, 160, 216
239
Cajal cells 9, 13, 201, 203, 211

Cajal-Retzius cells 9, 203, 211
Calleja cells 203
Callosal anlage 213, 214
Callosal radiation 150
Canine brain 36
Caudate nucleus 54, 59, 60, 61, 61, 62, 132, 214
Central sulcus (Rolando) 16, 17, 19, 24, 29, 32, 35, 36,
62, 63, 7375, 75, 76, 77, 80, 95, 142, 144, 178, 181,
182, 182, 183, 184, 202, 216
Choroid fissure 213, 214
Chronic alcoholism 179
Claustrum 60, 61, 62, 95, 96, 100
Cleavage (Zerklftung) 215
Cognitive disturbances 183
Commissural plate (Hochstetter) 213, 214
Conical ridge 137, 139
Conscious awareness 187, 197
Conscious perception 183
Consciousness 181, 197
Corona radiata 87
Corpus callosum 15, 17, 26, 31, 53, 55, 133, 134, 135,
136, 137, 139, 141151, 160, 166, 171
Corpus striatum 57, 95, 100, 179, 213, 214
Cortical cell mass 28, 209
Cortical cell number 28, 209
Cortical layer thickness 15, 28
Cortical layers 1, 3, 9, 12, 13, 212, 215
Cortical margin 26, 51, 53, 5557, 62, 63, 100, 132, 139,
151, 160, 171, 172, 214
Cortical plate 26, 27, 214
Cortical specific weight 28
Cortical structural types
agranular (type 1) 19, 21, 22, 23, 24, 36, 69, 95, 147,
178, 179, 218, 221
frontal (type 2) 21, 22, 23, 24, 69, 74, 83, 87, 96, 126,
218, 221
granulous (type 5) 21, 22, 23, 24, 80, 179, 219, 221
parietal (type 3) 21, 22, 23, 24, 69, 74, 87, 88, 92, 95,
98, 114, 124, 126, 127, 142, 187, 218, 221
polar (type 4) 21, 22, 23, 24, 47, 179, 218, 221
temporal 74, 114, 115, 119, 124, 129
Cortical subplate 211
Cortical surface 28
Cortical thickness 15, 16, 18, 28, 30, 204, 205, 218, 219
Cortical varieties
allocortex 15, 16, 24, 26, 27, 53, 56, 57, 60, 62, 129,
130, 132134, 141, 146, 152, 156, 158, 160, 163,
169, 212, 213, 216, 221, 222
granular 19, 44, 134, 145
heterotypic 17, 22, 27, 56, 57, 80, 96, 212, 216, 219,
221, 222
heterotypic agranular 32, 36, 53, 75, 100, 134, 139,
144, 145
240
heterotypic granulous 121, 123, 145

homotypic 17, 22, 27, 32, 44, 75, 81, 96, 129, 145, 212,
216, 219, 221, 222
isocortex 15, 16, 17, 22, 32, 53, 56, 57, 62, 75, 81, 129,
133, 139, 145, 146, 158, 169, 174, 212, 216, 219,
221, 222
koniocortex 21, 24, 80, 81, 83, 102, 108, 111, 112,
121, 123, 124, 145147, 151, 162, 163, 165, 167,
172, 181, 183, 203, 215, 216, 219, 221, 223, 224
primordial allocortex 221
rudimentary allocortex 221
striate allocortex 221
Cortical volume 28
Cortical weight 28
Cresyl violet 190
Crus fornicis (Fornixschenkel) 133, 213, 214
Cuneus 102, 105, 106, 107, 111, 112
Cytoarchitectonic areas
brain model 174
cortical maps 25, 19, 21
ground X, 6, 63, 64, 67, 222
modification X, XII, 68, 63, 72, 178, 222
thickness 30
transition X, XII, 1011, 27
variant X, 6, 63, 64, 67, 69, 222
Cytoarchitectonic neuropathology 29, 178, 179, 198
Deafness, congenital 176
Dementia praecox 179
Density coefficient (Dichtigkeitskoeffizient) 209
Diencephalon 81, 180, 195, 213
Dwarf cells 44
Dyschromatopsia 94
Economo-Koskinas corkscrew cells (Korkzieherzellen)
14, 60, 137, 139, 203, 208, 229
Economo-Koskinas rod cells (Stbchenzellen) 14, 60,
137, 139, 203, 208, 229
Economo-Koskinas sectioning method 193, 201, 225
Efferent fibers 25, 26
Ego 224
Embryonic development 17, 26, 27, 29, 156, 211, 212,
213, 214
Emotion 47, 181, 185
Encephalitis lethargica IX, XII, 179, 197, 228, 229
Engrams 181, 196
Eoanthropus dawsoni 189
Equilibrium functions 49, 53
Evolution 29, 67, 94, 147, 172, 189, 190
External fundamental layer (Jakob-Onelli) 13, 223
Fascia dentata (Tarin) 151, 154, 161, 164, 166, 168, 170
172
Fasciola cinerea 17, 26, 146, 171
Fimbria 31, 133, 134, 147, 149, 150, 150, 151, 152, 160,
161, 164, 166, 172
fMRI X
Fold (pli) of frontolimbic passage 71
Foot (pes) of F1 (pF1) 36
Foot (pes) of F3 (pF3) 44, 51, 57, 64, 66, 67, 69
Fornix 15, 17, 144, 150
Frontopontocerebellar pathways 49
Functional map 224
Fusiform transformation (Verspindelung) 59, 60
Ganglion basale (see nucleus basalis)
G/C coefficient 29
General paralysis 178180, 187
Germinal layer 211
Gifted and talented, neurobiology of 173n., 174, 190
Glomeruli, cellular 59, 157, 158, 167, 169
Granular cellular columns 114, 115, 127, 129
Granular transformation (Verkrnelung) 24, 27, 80,
108, 111, 121, 123, 124, 145, 146, 165, 181, 224
Granule cells 9, 11, 13, 20, 41, 44, 47, 60, 63, 67, 69, 72,
74, 75, 85, 145, 171, 172, 185, 201, 203, 207, 215
Gustatory domain 172
Guttiform cells 130, 203
Gyrus
ambient of uncus (Sterzi) 158, 160
angular 87, 88, 90, 91, 126, 184
annular 133
anterior arcuate 77
anterior insular 24
anterior limbic 208
antidiagonal 72
antitriangular 72
arcuate 85, 87
cingulate 31, 60, 133, 134, 135, 136, 137, 138, 139,
141147, 178, 179
dentate 15, 17, 26, 31, 134, 149172
descending (Ecker) 106, 112
digitate of uncus 170
falciform 95, 132, 133
fasciolar 15, 17, 26, 148, 151, 160, 161, 166, 171
first short insular 95
fornicate 31, 133
frontolimbic 72
frontolimbic transition 64, 69
fusiform temporal (T4) 24, 74, 88, 114, 124, 127, 128,
129, 130, 149, 179
geniculate 54, 55, 56
Heschl (transverse supratemporal) 24, 118, 119, 120,
122, 123, 178, 183, 185, 224
Heschl HI (first transverse supratemporal) 119, 121,
122, 123
Heschl HII (second transverse supratemporal) 119,
121
Subject Index
Heschl HIII (third transverse supratemporal) 120

hippocampal 24, 26, 31, 114, 129, 130, 134, 149172,
224
hippocampotemporal bridge 163
inferior frontal (F3) 19, 27, 35, 36, 41, 44, 46, 47, 51,
57, 59, 63, 64, 67, 68, 95, 178, 179, 185, 187
inferior temporal (T3) 24, 73, 74, 92, 114, 124, 126,
127, 130, 178, 179
internal olfactory 54
intralimbic (Anders Retzius) 15, 17, 26, 27, 134, 135,
148, 160, 161, 166, 171
lateral olfactory 15, 26, 53, 57, 59, 60, 61, 62, 96, 132,
160
limbic 24, 29, 69, 73, 74, 111, 129, 133, 145, 147149,
151, 161, 202, 216
lingual 102, 111
medial olfactory 15, 26, 53, 55, 59, 61
middle frontal (F2) 29, 36, 41, 44, 45, 47, 51, 69, 70,
184, 187, 202, 204
middle parietal (P2) 184
middle temporal (T2) 24, 73, 74, 92, 114, 124, 125,
126, 127, 130, 178, 179, 203
olfactory 27, 100, 133, 151
paraterminal 147
parieto-occipital 87
polar insular 132
postcentral 24, 74, 75, 75, 76, 77, 80, 81, 82, 83, 84,
84, 87, 178, 182, 184, 224
postcentral insular 99
posterior frontolimbic transition 142
precentral 22, 33, 35, 36, 41, 63, 64, 65, 74, 75, 80, 83,
182, 184, 184, 202
precentral insular 97
retrolimbic 160, 161, 166
rostral secondary temporal (Schwalbe) 130
semilunar 132, 151, 158, 160, 164
straight (see under area, cortical)
subcallosal 15, 17, 26, 54, 55, 61
superior frontal (F1) 29, 36, 41, 42, 43, 44, 45, 47, 48,
59, 77, 185, 187, 202, 204
superior temporal (T1) 19, 24, 74, 88, 90, 114, 116,
118, 119, 121, 124, 127, 130, 178, 179, 203
supramarginal 87, 88, 89, 90
transverse insular 53, 57, 58, 59, 59, 60, 61, 62, 95,
96, 132, 139, 208, 216
Gyrus segments
brink (edge) 25, 25, 26, 205
dome X, 15, 25, 25, 26, 28, 201, 205, 212, 222
valley (sulcus floor) X, 15, 25, 25, 26, 28, 201, 205,
212, 222
wall X, 25, 25, 26, 28, 201, 205, 219, 222
241
Head (caput) of F3 46, 67, 68, 69, 72

Hearing 16, 123
Hedgehog brain 15, 16
Hemiplegia 36
Hemispheric asymmetry 29
Hippocampal parolfactory stria 59
Hippocampus XI
Homo erectus 189n.
Homo rhodesiensis 189
Homo sapiens neanderthalensis 189, 189n.
Homo sapiens sapiens 189, 190
Horizontal lamination 129
Human brain 16, 17, 57, 62, 67, 94, 172
Huntington chorea 29, 179, 231, 232
H/W ratio 40
Hypothalamic border (His) (margo hypothalamicus)
213, 214
Ideomotor apraxia 92
Individual variations 29, 44, 64, 83, 163, 174, 183, 190,
201, 203, 221
Indusium griseum 15, 17, 26, 53, 55, 57, 134, 161, 166
Inferior choroidal point 149
Inferior ventricular horn 150, 152, 156, 164, 166
Infrachoroid layer (His) (lamina infrachorioidea) 213,
214
Inner main zone (Kaes) 13, 223
Insula (of Reil) 19, 129, 215
anterior 24, 95
function X, 101
posterior 95
Insular girdle (Inselgrtel) 96, 100
Intellectual functions 46
Intelligence 187, 189, 190, 223, 224
Intermediate fields (Flechsig) 224
Intermediate zone 26, 27, 211, 214
Internal capsule 213, 214
Internal fundamental layer (Jakob-Onelli) 13, 223
Interventricular foramen (Monro) 213
Isthmus 31, 111, 133, 134, 142, 144, 147149, 150, 151,
152, 162, 163, 165, 166, 167, 171, 216
Java man 189
Klliker cells 203
Korsakoff syndrome 179
Lamina affixa 150, 152, 172
Lamina commissuralis 53, 55, 62
Lamina dissecans (Rose) 130, 158, 162, 163, 165
Lamina rostralis 54, 55
Lamina terminalis 53, 54, 55, 59, 213, 214
Lancet cells 56, 139, 144, 146, 148, 167
Language 224
242
Lateral (Sylvian) fissure 1, 24, 30, 36, 44, 47, 73, 114,
115, 118, 119, 121, 123, 124, 126, 130, 175177, 184
Lenticular nucleus 214
Lissauer paralysis 179
Lobe, cerebral hemispheric
frontal 24, 29, 3272, 74, 81, 124, 133, 141, 142, 222
hippocampal (inferior limbic) 31, 133, 147, 149172,
222
insular 29, 30, 95101, 178, 179, 222
occipital 19, 24, 30, 73, 74, 87, 94, 102113, 133, 202,
222
parietal 19, 29, 7394, 133, 202, 222
superior limbic 31, 60, 64, 133148, 149n., 161, 222
temporal 19, 30, 87, 94, 114133, 222
Lobule
inferior parietal 24, 74, 85, 87, 89, 91, 92, 184
paracentral 34, 35, 36, 63, 73, 74, 76, 80, 81, 83
superior parietal 24, 74, 77, 85, 86, 87, 143
Locomotor complexes 41, 87
Macrosmatic animals 15, 146, 172
Macula lutea 187
Mapmakers problem X, XVI
Marginal veil (Randschleier) 214
Marginal zone 26, 27
Mastication 185
Matrix 26, 27, 211, 214
Medial geniculate body 123, 124
Medulla oblongata 83
Medullary margin (limbus medullaris) 214
Memory XI, 24, 113, 183, 221, 223, 224
Mental retardation 29, 176, 180, 196
Meynert stellate or solitary cells 14, 110, 111, 203
Microsmatic animals 172
Modalities, sensory 181, 221, 224
Motor cortex 36, 41, 44
Mouse brain 189
Muscle sensibility 81, 90
Music comprehension 119, 196
Myelination 123, 224
Myeloarchitectonics 9, 13, 180
Navicular cells 19, 100, 162
Neuroepithelium (ependymre Anlage) 211, 212
Nissl bodies 11, 13, 14, 35, 67, 76, 103, 107, 111, 201
Nissl stain 13, 168, 170, 190, 201
Nucleus basalis (Meynert) 54, 57, 59, 60, 61, 61,
62
Occipitotemporal intermediate zone 92
Ocular movements 44, 87, 127
Olfaction 16, 51, 101, 146, 147, 172
Olfactory bulb 213, 214
Olfactory nerve 62
Olfactory root 49, 53, 56, 57

external ramus 172
lateral 101, 132
medial 55, 101
Olfactory striae 59
Olfactory tract 53, 56, 57, 59, 60, 132, 155
Olfactory triangle 26, 59, 132
Olfactory tubercle 26, 59, 60
Oligophrenia 180
Operculum
frontal 2, 32, 41, 44, 46, 63, 64, 72, 95
occipital (Affenspalte) 112, 112n., 188, 189, 229
parietal 2, 74, 80, 81, 85, 95
Rolando 41, 64, 65, 90, 183, 184, 216, 229
Optic chiasma 59, 213, 214
Optic tract 57, 60, 61, 61, 62
Orangutan brain 67
Orbital margin 53
Organ pipe formation 115, 119, 121
Outer main zone (Kaes) 13, 223
Paracentral fossa 36
Parasensory zones 25, 83, 107, 108, 108n., 121, 124, 229
Parolfactory eminence 59, 60
Pars ascendens of pF3 67
Pars opercularis of F3 67
Pars orbitalis of F2 51
Pars orbitalis (foot) of F3 (pF3) 44, 51, 57, 64, 66, 67, 69
Pars pretriangularis of F3 51, 72
Pars triangularis (head) of F3 46, 67, 68, 69, 72
Pathoclisis (Pathoklise) 178, 179, 234
Pearl bead arrangement 123
Phonation 185
Phylogeny 67, 94, 172, 190, 191, 195, 223
Pick disease 179
Piltdown man 189
Pithecanthropus erectus 189
Planar lenses 200
Planum septale 60
Planum temporale 119
Pole, hemispheric
frontal 16, 19, 22, 29, 32, 41, 46, 47, 49, 50, 51, 59, 62,
69, 187, 204
insular 57, 95, 96, 100, 216
occipital (posterior) 16, 19, 22, 47, 74, 88, 102, 106,
108, 109, 111
temporal 5, 22, 59, 114, 118, 119, 129, 131, 132, 149,
178, 213, 216
Pons 215
Porencephaly 180
Posterior border (margo posterior) 213
Postural functions 49, 83
Praxic functions 44
Precuneus 73, 142, 144
Subject Index
Preoccipital incisure (notch) 30, 73

Presubiculum 15, 151, 152, 165, 167170, 179
Primates, nonhuman 94, 172, 189
Primordial centers (Flechsig) 221, 224
Prosencephalon 195
Progressive cerebration XII, 173n., 190
Psychoses 178180, 187, 197
Pyramidal cells 9, 11, 19, 32, 35, 40, 41, 44, 49, 63, 67,
69, 76, 81, 83, 167, 168, 178, 182, 201, 203, 206, 215,
220
Pyramidal tract 36
Pyramidal transformation (Pyramidisierung) 22, 27, 35,
40, 203
Rabbit brain 17
Radial striation 19, 26, 36, 41, 47, 67, 74, 85, 87, 88, 90,
92, 114, 115, 118, 119, 121, 126, 209, 210
Rain shower formation 121
Receptive fibers 26
Red nucleus 215
Region, cortical
anterior superior limbic 222
basal parietal 87, 88, 104, 126, 222
entorhinal 151, 156, 172
fusiform 222
inferior parietal 73, 88, 104, 115, 118, 187, 222
orbital (orbitomedial) frontal 222
postcentral (anterior parietal) 74, 222
posterior superior limbic 222
posteroinferior parietal 187
prefrontal (anterior frontal) 185, 187, 222
prerolandic 222
retrosplenial 24, 31, 74, 133, 138, 142, 144148, 150,
161, 166, 216, 222, 224, 229
superior parietal 74, 83, 222
supratemporal 222
temporal proper 222
temporo-occipital 73, 87
temporopolar 115, 130, 216, 222
Retina 1, 181, 187
Rhinencephalon XI, 14, 15, 17, 19, 27, 49, 51, 53, 59, 60,
64, 72, 101, 129, 133, 134, 142, 146, 147, 220
Rhodesian man 189
Rows of soldiers 96, 158
Satellite cells 35, 207
Schizophrenia 179, 180, 187
Senile dementia 178, 179
Septum pellucidum 55, 57, 151
Sexual dimorphism (gender differences) XIII, 223
Silver impregnation 11, 13, 201
Single movements 83
Slenderness (Schlankheit) 11, 40, 201, 203
Song center 69
243
Sound perception 124

Spastic pseudosclerosis 179
Spatial disorientation 94
Special cells 13, 201, 203
Speech areas, motor 67, 69
Spinal cord 81
Spindle (fusiform) cells 9, 11, 19, 32, 40, 46, 59, 60, 83,
162, 169, 201, 203, 207, 219
Spindle transformation (Verspindelung) 203
Splenium 111, 133, 142, 144, 150, 151, 160, 161
Spokes of wheel 170
Stellate cells 14, 19, 56, 57, 59, 60, 62, 98, 100, 110, 132,
145, 157, 160163, 169, 203
Stereoagnosia 183
Stereognosis 90
Stratum cellulare 170
Stratum oriens 170
Stratum radiatum 170
Stria of Gennari 13, 14, 102, 108, 110, 111, 112
Stria of Lancisi 139
Stria of Vicq dAzyr 1, 13, 14, 102
Stripe (band) of Baillarger 1, 13
Stripe of Kaes-Bekhterew 13
Stylum septi 57, 59, 61
Subiculum 15, 27, 151, 152, 161, 167172, 179
Substantia perforata 15, 17, 26, 27, 29, 51, 53, 55, 57, 59,
60, 61, 61, 62, 95, 96, 100, 101, 130, 132, 133, 164
Substantia reticulata (Arnold) 156, 159
Sulcus
acoustic 118
anterior insular 30
anterior occipital 73
anterior parolfactory 54, 54, 55
anterior subcentral 183
calcarine 1, 16, 17, 24, 27, 73, 92, 94, 102, 107109,
109, 110, 111, 112, 133, 144, 149, 160, 161, 166,
188, 215, 224
callosal 3, 24, 134, 135, 136, 137, 138, 139, 142144,
147, 151, 160, 161, 166
callosomarginal 17, 36, 41, 46, 47, 51, 64, 69, 73, 76,
77, 85, 142
central insular 95, 96
cingulate 135, 137
circular insular 44, 51, 95, 100
collateral 29, 30, 129, 130, 152, 161, 163
cruciate 36
diagonal 59, 62
external calcarine 188
fimbriodentate 150, 151
frontomarginal 47
hippocampal 3, 24, 134, 150, 151, 160, 163, 164, 165,
167
hypothalamic (Monro) 213
inferior accessory occipital 188
244
inferior frontal ( f2) 35, 36, 41

inferior occipital 188
inferior postcentral 183
inferior precentral 183
internal calcarine 188
interoccipital 29, 30, 73, 188
interparietal 85, 90, 188
lateral occipital 188
limbic 69
lingual 106
lunatus 112, 189
middle parolfactory 54, 55, 56
occipito-temporal 94, 114, 216
olfactory 49, 51, 56, 152, 220
opercular limitans 188
orbital 51
parieto-occipital 17, 29, 30, 73, 85, 102, 104, 106,
111, 133, 143, 144, 160, 188
parolfactory postremus 54, 55, 59, 62
perpendicular occipital (Bischoff) 188
postcentral 74, 84, 85
posterior insular 30
posterior parolfactory 54, 55, 56, 62
posterior subcentral 183
precentral 36, 41
primary occipital 106
rhinal 130, 152, 158, 161163
secondary occipital 106
semilunar 106, 112
subparietal 73, 85, 143, 144
superior accessory occipital 188
superior frontal ( f1) 36
superior insular 30
superior rostral 52
superior temporal 126
temporoinsular 114
temporo-occipital 152, 163
tertiary occipital 106
transverse occipital, lateral (ventral) ramus 188
transverse occipital, medial (dorsal) ramus
188
transverse (parietal) 106
upturned (Brissaud) 85
Superior temporal plane 187
Suprachoroid margin (His) (limbus suprachorioideus)
213, 214
Supraoptic ganglion 61
Sympathetic nervous system 101, 147
Tactile domain 81
Taenia (embryonic brain) 214
Taenia tecta 144, 146, 151, 161
Taste modalities 81, 172
Telencephalodiencephalic border 213
Temporo-occipital transition (intermediate) zone 29,

73, 87, 88, 93
Temporopontine fiber tracts 127
Terminal fields (Flechsig) 224
Thalamic border (margo thalamicus) 213
Thalamus 215
Thermesthesia 81
Toluidine blue 190, 193, 201, 225, 226
Tone perception 123, 183
Trapezoidal field (Trapezfeld) 213, 214
Trigonum olfactorium 53, 56, 57
Tufted cells (Quastenzellen) 14, 203
Uncus 15, 24, 26, 27, 31, 127, 130, 132134, 147, 149172
Uncus knee 161, 170
Uncus pole 156, 170, 172
Upright posture 41
Subject Index
Velum terminale (Aeby) 149, 151

Verbal audition 119
Verbal cognition 119
Vertebrates, lower 94, 187
Vertical cellular columns 64, 67, 119, 121, 124, 126
Vesicles, anterior cerebral (Vorderhirnblschen) 55
Vision 16, 102, 112, 113
Visual orientation disturbances 129
Visuomotor disturbances 92
Walking 41
Weigert method 9, 13, 80
Will (psychomotor) 47, 185, 187
Word meaning 119, 183
Word phonetics 119
Writing center 41
245

Cellular Structure of The Human Cerebral Cortex (2009)

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Cellular Structure of the Human Cerebral Cortex

Constantin von Economo

88 gures including 48 original microphotographs, 4 in color, 12 tables, and fold-out, 2009

Basel Freiburg Paris London New York Bangalore

New English edition

Frontispiece portrait information:

Library of Congress Cataloging-in-Publication Data

Copyright 2009 by S. Karger AG, P.O. Box, CH4009 Basel (Switzerland)

Vorwort Prface Prefazione Preface

Superior Limbic Lobe

Hippocampal (Inferior Limbic) Lobe

The German edition of the book (146 pp.) [1]

[12, 13]. Like the German original, it contained

specifically appearing in one or the other of the

descriptions of the brain including the insula.

It is worth reviewing some of the reviews that the

As a matter of fact, all of the microphotographs depicting

most likely refers to the English edition [4] of

plement to Cellular Structure. The Outline has

Nothing may possibly conclude a discussion

16 Parker S: The determination of insanity

30 Jakob A: Normale und pathologische

40 Schwartz EL, Shaw A, Wolfson E: A

47 Moscovitch M, Chein JM, Talmi D,

53 Notices of Recent Publications: Encephalitis Lethargica: Its Sequelae and

Authors Foreword to the 1927 German Edition

Authors Preface to the 1927 French Edition

plus approfondie, ils fournissent cependant un

Authors Preface to the 1928 Italian Edition

Numerosi medici e numerosissimi studenti, e pi

per pu tornare utile al ricercatore che voglia

Authors Preface to the 1929 English Edition

The desire for a handy and concise introduction

cient use of the larger work. The general view and

1903, 1906, 1910, 1911, 1913], Vogt and Vogt

Partition Scheme of the Cerebral Cortical

The first two discoveries mentioned namely,

entire subsequent knowledge on cortical cytoarchitectonic areas has emerged.

The Six Cortical Layers

Frontal lobe (F)

Anterior frontal (prefrontal) region

granular frontal area

Orbital (orbitomedial) region

granular orbital area

Superior limbic lobe (L)

precingulate agranular anterior limbic area

Posterior superior limbic region

dorsal posterior cingulate area

agranular retrosplenial area

Insular lobe (I)

dorsal precentral insular area

Parietal lobe (P)

giant pyramidal postcentral area

Superior parietal region

superior parietal area (transition postcentral)

Inferior parietal region

Basal parietal region

basal (temporooccipital) parietal area at parietal entrance

Occipital lobe (O)

anterior peristriate area

Temporal lobe (T)

posterior superior temporal area

Temporal region proper