Review Neurosciences and History 2024; 12(4): 240-251

Functional organisation of the cerebral cortex: from Gall to Lashley

A. García-Molina1,2,3, J. Peña-Casanova4,5
1
Institut Guttmann, Institut Universitari de Neurorehabilitació (UAB), Badalona, Barcelona, Spain.
2
Fundació Institut d’Investigació en Ciències de la Salut Germans Trias i Pujol, Badalona, Barcelona, Spain.
3
Universitat Autònoma de Barcelona, Bellaterra, Barcelona, Spain.
4
Fundació Institut Mar d’Investigacions Médiques (FIMIM), Barcelona, Spain.
5
Psychiatric and Legal Medicine Department. Universitat Autònoma de Barcelona, Bellaterra, Barcelona, Spain.

ABSTRACT

Introduction. In the late 18th century, Franz Joseph Gall questioned the concept of the functional unity of the
cerebral cortex, foreshadowing the advent of a new era in the study of cortical function.
Development. This review aims to provide a general view of the understanding of the functional organisation of
the cerebral cortex in the 19th and early 20th centuries. This historical period is characterised by the dialectical
confrontation between two factions: supporters of the parcellation of the cerebral cortex, and proponents of the
functional unity of the cerebral cortex. The latter position, supported by Marie-Jean-Pierre Flourens’ doctrine
of cortical equipotentiality, dominated scientific thought in the first half of the 19th century. In the 1860s, it was
supplanted by the theory of cortical parcellation, thanks to the contributions of Paul Pierre Broca, Eduard Hitzig,
and Gustav Fritsch. During the same period, Carl Wernicke established the basis for cortical connectionism, a
doctrine that conceived the cortex as a mosaic of interconnected functional centres. The early years of the 20th
century saw an expansion of the anti-localisationist current, led by Pierre Marie, Henry Head, and Shepherd Ivory
Franz; this current reached its greatest height with Karl Lashley.
Conclusions. In the 19th century and the first decades of the 20th century, the study of the functional organisation
of the brain pivoted between two opposing epistemological positions: the localisationist/connectionist doctrine
and the anti-localisationist current.

KEYWORDS
Cerebral cortex, localisationism, connectionism, antilocalisationism, equipotentiality, neuroanatomy

Introduction years. The Renaissance marked the beginning of the end

of the ventricular doctrine, although the transition from
Medieval understanding of the nervous system was classical/medieval to modern medical thought did not
based on the teachings of Galen of Pergamon (130- take place until the 17th and 18th centuries. The meta-
220 AD) and their interpretation by the first fathers of physical, theological, and philosophical debates of me-
the Eastern Christian Church (ca. fourth century AD). dieval thought gave way to medical argument based on
Galenic theories about the brain included the ventricular concepts that were better aligned with anatomical and
doctrineA and the theory of animal spirits,B dogmas that physiological reality.1
guided the study of the nervous system for over 1500
The collapse of the ventricular doctrine gave rise to
the need to identify a new biological substrate for psy-
A
The ventricular doctrine asserted that the cerebral ventricles were the chic life. In 1664, in his work Cerebri anatome, Thomas
biological substrate of intellectual faculties.
B
According to the theory of animal spirits, brain function is the product of a
Willis (1621-1675)2 theorised that the cerebral cortex is
substance (the animal spirits) stored in the cerebral ventricles. responsible for memory, imagination, and volition. In

Corresponding author: Dr Alberto García-Molina Received: 30 January 2024 / Accepted: 25 March 2024
E-mail: [email protected] © 2024 Sociedad Española de Neurología. Open Access CC BY-NC-ND 4.0.

240
 Functional organisation of the cerebral cortex: from Gall to Lashley

the first half of the 18th century, Emanuel Swedenborg Most philosophers find the opinion ridiculous that
(1688-1772) published Oeconomia regni animalis the various psychic faculties and notions have their
(1740)3. He employed similar arguments to Willis’, and seats in different places of the brain. But if this is
incorporated the hypothesis that the cortex is made up ridiculous, it is also ridiculous that the different
of specialised regions, which are responsible for different senses are placed in different parts of the body.6(p197),D
functions. Soon after, in 1779, Georg Procháska (1749- He dedicated several years to gathering evidence, and
1820) argued that the cortex contained numerous organs in 1798 he wrote a letter to Joseph Friedrich von Retzer
that acted in concert. The theories of Swedenborg and (an official at the censor’ s office in Vienna) explaining
Procháska were the prelude to a new era in the study of the basic postulates of a new “science”: Schädellehre (the
cortical function. “skull doctrine”).E
For centuries, the anatomical structure of the cerebral Schädellehre, which later came to be known as phrenol-
ventricles had been extensively researched due to the ogy, was based on the observation that the skull presents
significant interest in their function. It was only in the pronounced bumps in certain areas; from these, a series
19th century, when the cerebral cortex became pre-emi- of specific individual talents could be derived, which
nent as the seat of psychic life, that a detailed topography were the result of uneven development of different areas
of the cortex was developed. Until then, the lateral sul- of the brain. Gall proposed that, just as the body contains
cus (or Sylvian fissure) was one of the few cortical struc- organs associated with specific physiological functions,
tures to have been identified. In 1663, Franciscus Sylvius
the brain is also made up of mental organs, each of which
(1614-1672)4 wrote:
is dedicated to a specific task.6 Through external analy-
Particularly noticeable is the deep fissure or hiatus sis of the skull (cranioscopy), Gall concluded that there
which begins at the roots of the eyes […] it runs were 27 mental organs or faculties, localised bilaterally
posteriorly above the temples as far as the roots across both cerebral hemispheres (Figure 1).7
of the brain stem. It divides the cerebrum into an
upper, larger part and a lower, smaller part. Schädellehre was based on the following assumptions:
Johann Christian Reil (1759-1813) described the insula — The brain is the organ of the mind.
in 1809, and Luigi Rolando (1773-1831) described the — The brain comprises a set of organs or mental faculties.
fissure that bears his name in 1829. The terms frontal, — The organs or mental faculties making up the brain
temporal, parietal, and occipital (now used to divide are located in different brain areas, each of which has a
the surface of the cortex) were introduced by Friedrich specific function.
Arnold (1803-1890) in 1838.C In the 1850s, Louis Pierre — As the skull ossifies over the brain during its forma-
Gratiolet (1815‐1865) identified nearly all the gyri as tion, external analysis of the cranium (cranioscopy) is a
we know them today. This work was completed in the method for diagnosing the state of the organs or mental
following decade by William Turner (1832-1916) and faculties.
Alexander Ecker (1816-1887).5 In the early 19th century, phrenology enjoyed great
This review aims to provide a general view of the under- scientific/academic prestige; this status was gradual-
standing of the functional organisation of the cerebral ly lost, until it was reduced to a pseudoscience. One of
cortex in the 19th and early 20th centuries. This histori- its main critics was Marie-Jean-Pierre Flourens (1794-
cal period is characterised by the dialectic confrontation 1867) (Figure 2).8,9 Flourens accepted that intellectual
between two opposing epistemological positions: the
supporters of the atomisation of the cortex, and those
who believed it to exist as a functional unit. C
In 1807, François Chaussier (1746-1828) divided the cerebral cortex into
three sections: the anterior or frontal, middle or temporal, and posterior or
Gall, precursor of cortical localisationism occipital lobes.
D
Translator’ s note: the English translation of this excerpt is taken from:
Eling P, Finger S, Whitaker H. On the origins of organology: Franz Joseph
In 1871, Franz Joseph Gall (1758-1828) intuited that dif- Gall and a girl named Bianchi. Cortex. 2017;86:123-31.
ferent mental faculties were located in different regions E
The letter was subsequently published in the journal Der Neue Teutsche
Merkur (Gall FJ. Schreiben über seinen bereits geendigten Prodromus über
of the brain; however, he possessed no evidence support- die Verichtungen des Gehirns der Menschen und der Thiere, an Herrn Jos.
ing this hypothesis: Fr. von Retzer. Der neue Teutsche Merkur. 1798;27:311-32).

241
 A. García-Molina, J. Peña-Casanova

Figure 1. Left: Franz Joseph Gall (1758-1828). Centre/right: approximate locations of the mental faculties, according to Gall (Anatomie
et physiologie du système nerveux en général et du cerveau en particulier, 1810).7

Figure 2. Left: Marie-Jean-Pierre Flourens (1794-1867). Right: title page of Examen de la phrénologie (second edition; 1845).9

Figure 3. Left: the brain of Monsieur Leborgne, showing a lesion to the left third frontal gyrus. Right: Paul Pierre Broca (1824-1880).

242
 Functional organisation of the cerebral cortex: from Gall to Lashley

functions are the domain of the cerebral cortex, but re- the first half of the 19th century. However, some research-
jected the notion that the cortex was made up of a fed- ers, continuing along the path laid out by Gall, argued that
eration of organs, each of which had a specific function: the cerebral cortex was not a homogeneous structure, but
Hence it appears, that the cerebral hemispheres rather was made up of multiple functional regions.
concur, by their whole mass, in the full and entire Jean-Baptiste Bouillaud (1796-1881), a dedicated follow-
exercise of the intelligence. In fine, as soon as one er of Gall,G considered cranioscopy not to be the most
sensation is lost, all sensation is lost; when one suitable method for studying brain function. Rather than
faculty disappears, all the faculties disappear. There inferring the functional role of brain regions by examin-
are not, therefore, different seats for the different ing the bumps on a person’ s skull, he used the anatomo-
faculties, nor for the different sensations. The faculty
pathological method, establishing correlations between
of feeling, of judging, of willing any thing, resides
neurological signs and structural brain lesions.11 In May
in the same place as the faculty of feeling, judging,
or willing any other thing, and consequently this
1825, he gave a lecture at the Académie de Médecine
faculty, essentially a unit, resides essentially in a entitled “Recherches cliniques propres à démontrer que
single organ. The understanding is, therefore, a la perte de la parole correspond à la lésion des lobules
unit.9(p28-9),F antérieurs du cerveau, et à confirmer l’ opinion de M.
Gall sur le siège de l’ organe du langage articulé” (Clinical
This French physician also harshly criticised Gall’ s research aimed at demonstrating that loss of speech cor-
methodology, noting that it was based on the topography responds to a lesion to the anterior lobes of the brain,
of the skull, rather than that of the brain. “But as to the and at corroborating Gall’ s opinion on the location of
pretended organs of the brain, are they really situated at the organ of articulate language),12 in which he suggest-
the surface of the brain, as Gall asserts? In plain terms, ed that the organ of articulate language was located in
is the surface of the brain the only active part of the or- the anterior lobes of the brain, an idea he defended for
gan?”9(p78-9) In another passage, he explains how: over 50 years.13-15 His contributions were fundamental
The cranium, especially the external surface of it, in the transition from Gall’ s theoretical speculation to-
represents the superficial configuration of the brain wards the scientific studies that supported the relation-
but very imperfectly. Gall knows it. “I was the first,” ship between the cerebral cortex and specific cognitive
says he, “to maintain that it is impossible for us processes.
to determine with exactitude the development of
certain circumvolutions, by the inspection of the On 18 April 1861, Paul Pierre Broca (1824-1880) pre-
external surface of the cranium.” […] Gall is aware sented to the Société Anthropologique de Paris the case
of all this, and nevertheless he inscribes his twenty- of a patient who had lost the ability to speak (Monsieur
seven faculties upon the skulls.9(p83) Leborgne, also known as Monsieur Tan-Tan).16 The post
mortem examination revealed a lesion in the left third
Gabriel Andral (1797-1876), a contemporary of Flourens,
frontal gyrus (Figure 3). The same year, he presented a
notes that:
second case with similar characteristics, in which a le-
In the point where a lesion is discovered, the direct sion was detected in the same location.17 In 1865, after
cause of the effects which it produces does not always analysing other cases, he concluded that the left third
reside […] If then it happened that we succeeded frontal gyrus was the neuroanatomical substrate of artic-
in discovering in the encephalon a certain number ulate language.18 Broca’ s observations constitute the first
of parts, the lesions of which always occasioned the
documented empirical evidence of the correspondence
disturbance of the same cerebral act, it would not,
between a cognitive process and a specific region of the
in our opinion, be fair to object to the doctrine of
cerebral cortex, contravening Flourens’ dogma of corti-
localisation, that there are also other cases where
this same functional disturbance is reproduced, cal equipotentiality.19
though the lesion might be elsewhere.10(p734-5)

Cortical localisationism F
Flourens’ assertions anticipated the concepts of equipotentiality and mass
action, developed in the 1920s by Karl Lashley (1890-1958).
Flourens’ conception of the cerebral cortex was broadly G
Bouillaud was a founding member of the Société Phrénologique de Paris
accepted by his contemporaries, becoming dogma during (the inaugural meeting of the society was held on 14 January 1831).

243
 A. García-Molina, J. Peña-Casanova

Broca’ s discovery sparked interest among the scientific (1839-1912)25 demonstrated that destruction of the oc-
community in identifying cortical functional regions, cipital lobes caused vision loss, and Leonardo Bianchi
leading to the development of an explanatory model of (1848-1927)26 observed that surgical lesions to the frontal
the neurobiological basis of mind based on the “one re- lobes caused dramatic changes in complex behaviours.
gion, one function” axiom. Jean-Martin Charcot (1825- In the latter third of the 19th century, experiments were
1893) argued in 1875 that “there are areas in the brain in also performed in humans (from today’ s perspective,
which a lesion inescapably leads to the same symptoms. these were ethically reprehensible). In 1874, Roberts
Beyond this law, all else is confusion.”20(p400) He also ar- Bartholow (1831-1904) replicated the study by Hitzig
gued that: and Fritsch, performing the first documented demon-
The brain is not a homogenous, single organ, but stration of the excitability of the human cerebral cortex.
rather a group, or if you wish, a confederation, To conduct the study, he applied direct electrical current
composed of a number of different organs. To each to the left postcentral region of the brain of Mary Rafferty
of these, distinct properties, function and faculties (which was exposed due to erosion of the scalp and skull
are physiologically attached. Once the physiologic by a cancerous ulcer).28 Soon after, Ezio Sciamanna
properties of each of these parts is known, it should (1850-1905) conducted a series of experiments in which
be possible to deduce the pathologic situation, he electrically stimulated the surface of the brain in a pa-
since this would only represent a modification, tient who had undergone trepanation due to a traumatic
mild or marked, of the normal state, without any
brain injury. In 1883, Alberto Alberti (1856-1913) rep-
intervention of new laws.21(p4)
licated Bartholow’ s study in a woman with an eroding
In 1883, Charcot and Albert Pitres (1848-1928) pub- tumour of the skull, allowing access to the surface of the
lished a study describing a large series of patients with underlying dura mater.29
cortical lesions. Faithful to his convictions, he consid-
ered evidence against the localisationist doctrine to pres- Cortical connectionism
ent “defects of form,” diminishing its credibility.22 In 1868, Theodor Meynert (1833-1892) proposed an
Animal experimentation provided new evidence in sup- associationist brain model based on the premise that
port of cortical localisationism. In 1870, Eduard Hitzig psychic functions resulted from the interaction between
(1838-1907) and Gustav Fritsch (1838-1927) observed different parts of the brain. In other words, associations
that application of galvanic current to the posterior fron- created complex structures that cannot be located in a
tal lobe in a dog caused movement in the hemibody con- specific brain region.30 Carl Wernicke (1848-1905) rec-
tralateral to the cortical region stimulated.23 Specifically, ognised the importance of this model, and applied it in
they identified five motor centres, whose stimulation his analysis and interpretation of cognitive deficits sec-
provoked muscle contractions in the neck, abduction ondary to neurological lesions (Figure 5). In 1874, he
of the forelimbs, flexion of the hindlimbs, movement of published his work Der aphasische Symptomencomplex:
the forelimbs, and facial contractions (Figure 4). These eine psychologische Studie auf anatomischer Basis (“The
findings strengthened localisationist positions, and cast aphasia symptom-complex: a psychological study on
doubt on the idea of cortical insensibility.H As noted by an anatomical basis”),31 in which he proposed a corti-
Hitzig and Fritsch themselves, “with the results of our cal associationist model based on two principles: 1) the
research, many of the conclusions about the basic prop- cerebral cortex contains centres responsible for specific
erties of the brain are substantially changed.”23(p308) Their functions; and 2) these centres are connected with one
findings brought about a cascade of experimental stud- another through association pathways. Thus, a particular
ies that helped to consolidate the pre-eminence of cor- pathological behaviour may be predicted according to
understanding of how the flow of information between
tical localisationism: David Ferrier (1843-1928)24 iden-
centres has been interrupted, or how these centres have
tified several cortical motor centres, Hermann Munk
been destroyed. To differentiate his theories from local-
isationism (and by extension, phrenology), Wernicke
argued that “Any higher psychic process, […] could not
H
In the 1750s, Albrecht von Haller (1708-1777) had concluded that the
external surface of the cerebral cortex was insensitive to mechanical, electric,
[…] be localized, but rested on the mutual interaction
or chemical stimulation. This theory was endorsed by Flourens in the 1820s. of these fundamental psychic elements mediated by

244
 Functional organisation of the cerebral cortex: from Gall to Lashley

Figure 4. Left: Eduard Hitzig (1838-1907). Centre: drawing of the excitable area of the dog cerebral cortex.27 Δ Contraction of the neck
muscles. + Abduction of the forelimbs. +’ Flexion of the forelimbs. # Movement of the forelimbs. ο Facial contraction. Right: Gustav
Fritsch (1838-1927).

Figure 5. Left: a diagram by Wernicke depicting the sensory and motor language areas and their connections.31 Right: Carl Wernicke
(1848-1905).

Figure 6. Left: a diagram by Magnan (1881). Centre: a diagram by Lichtheim (1885). Right: Charcot’ s bell diagram (1885). Source:
Moutier, 1908.36

245
 A. García-Molina, J. Peña-Casanova

means of their manifold connections via the association destroying speech and to locate speech were two different
fibers.”32(p824) Although Wernicke’ s proposal is an adapta- things.37 Similarly, Veniamin Mikhailovich Tarnovsky
tion of Meynert’ s associationist model, it constitutes the (1837-1906) argued that it is impossible to conclude, as
starting point for a new model of cortical physiology: the many do, that post mortem evidence of the destruction
connectionist doctrine. of the left frontal gyrus in an aphasic patient means that
said lesion is the sole cause of the aphasia and that, as a
The connectionist doctrine offered a working method-
result, the ability to speak is located in that brain region.38
ology that was useful both in research and in clinical
practice. The elegant system of centres and connection Although Hughlings Jackson’ s critiques were eclipsed
pathways brought order to the chaos of symptoms ob- by the success of the connectionist approach, his hy-
served in patients, helped to classify symptoms within potheses played an important role in the development
a limited set of syndromes, and enabled individualised of alternative explanations about cortical functional
neuroanatomical examination of each patient’ s deficits. organisation. The turn of the century saw an increased
Connectionism was adopted, adapted, and disseminated emphasis on theses that questioned the clinical-anatom-
by various disciples of Wernicke’ s.33 Other authors who ical associations defended by localisationists and con-
embraced this doctrine include Henry Charlton Bastian nectionists. Friedrich Goltz (1834-1902), Jacques Loeb
(1837-1915), Charles K. Mills (1845-1931), Joseph (1859-1924), and Karl Lashley (1890-1958), who sub-
Grasset (1849-1918), and Jules Dejerine (1849-1917), scribed to the theory of equipotentialism, alleged that
among others. all cortical areas were equivalent from a functional per-
spective (Figure 7). The holists, in turn, believed that the
Henry Head (1861-1940) was a harsh critic of the con-
brain acted as an integrated functional unit. They argued
nectionist doctrine, and particularly of authors who
that mental processes did not result from the independ-
created graphical representations of their functional
ent activity of individual parts of the brain, but rather
architectures (Figure 6), whom he dismissed as “dia-
from the interdependent activity of the brain as a whole.
gram-makers.” He believed that physicians ran the risk of
This school of thought included Pierre Marie (1853-
considering the diagram to be more important than the
1940), Henry Head (1861-1940), Charles Sherrington
patient. Clinical reality, argued Head, was richer, more
(1857-1952), Shepherd Ivory Franz (1874-1933), and
varied, and more complex than diagrams, and did not
Kurt Goldstein (1878-1965).I Outside the field of med-
fit the categories generated on the basis of a framework
icine, we should also note the organicist ideas of Karl
of centres and connections.34 Similarly to Head, Pierre Ludwig von Bertalanffy (1901-1972).J This Austrian biol-
Marie (1853-1940) believed these graphical representa- ogist declared that organisms were open systems, which
tions to be of no value as they oversimplified the clinical were in constant interchange with other, nearby systems
reality and illustrated the prejudices and obsessions of by means of complex interactions. With respect to the
their authors.35 brain, Bertalanffy questioned the premise that it was a
The anti-localisationist current set of centres, and proposed that focal dysfunction results
in a general alteration to the functioning of the brain.39
The localisationist-connectionist doctrine became the
The term “anti-localisationist current” encompasses
frame of reference for neurology and neurophysiology in
highly varied propositions; one of the most unique was
the latter third of the 19th century and the early years of
the 20th century. However, many authors believed that it
offered only a crude, ingenuous explanation of cortical I
Analysis of Goldstein’ s work reveals a transformation in his thinking: his
physiology. In this context, the anti-localisationist cur- early texts (written in his youth) are based on essentially localisationist-
connectionist ideas, whereas his later works present holistic positions.
rent comprised a heterogeneous group of physicians and This conceptual evolution took place in line with the growth of his clinical
researchers who rejected the principles of the localisa- experience.
J
The functional system concept proposed by Alexandr Romanovich Luria
tionist-connectionist doctrine. (1902-1977) is based on Karl Ludwig von Bertalanffy’ s (1901-1972)
biological systems theory, Alexei Alexeievich Ukhtomsky’ s (1875-1942)
In 1874, John Hughlings Jackson (1835-1911) warned dominant system principle, and Pyotr Kuzmich Anokhin’ s (1898-1974)
that localisationists committed the error of equating functional systems theory. Luria established that mental functions were
organised in systems of brain areas, with each area playing a specific role
the localisation of symptoms with the localisation of within the system. A lesion to one area would alter the function of the
functions. Specifically, he noted that to locate the lesion system as a whole (but with specific characteristics).

246
 Functional organisation of the cerebral cortex: from Gall to Lashley

the theory set forth by Constantin von Monakow (1853- lobes without destroying their functions complete-
1930) in his 1914 work Die Lokalisation im Grosshirn ly. However, it is even more than that. The lobe can
und der Abbau der Funktion durch kortikale Herde (The recover these functions in their entirety after having
localization in the cerebrum and the degradation of lost them completely.”8(p101) In another passage he
the function by cortical foci).40 The author argues that notes that:
the brain is organised in constellations of synchronous As long as not too much of the lobes is removed,
networks, producing chronogenic localisations. The dy- they may regain in due time the exercise of their
namic-functional concept of chronogenic localisation functions. […] if one sensation comes back, all
emphasises the importance of the temporal dimension come back. If one faculty reappears, they all
in understanding brain physiology, transcending the reappear.8(p102)
topographic/spatial visions of the localisationists and
A century later, Lashley reported that the functional
anti-localisationists of the day. Historically, the temporal
impairment observed in experimental animals sub-
properties and chronotopic distribution of cerebral pro-
jected to cortical ablation was related to the amount
cesses have played a marginal role in research into the
of cortex removed, not to its location (Figure 8).
functional organisation of the brain.K
This finding led him to formulate the general prin-
One of the many arguments employed by the anti-local- ciples of cortical equipotentiality and mass action.41
isationists against localisationism-connectionism is the According to Lashley, all brain regions were func-
so-called problem of recovery (evidence of the brain’ s tionally equivalent; however, this equipotentiality
capacity to regain lost or impaired functions). The lo- was not absolute, but rather was subject to the prin-
calisationist-connectionist doctrine gave the impression ciple of mass action (the efficiency of functional
that the cerebral cortex is a rigid, non-malleable structure performance is determined by the extension, rath-
lacking the capacity for functional reorganisation after er than the localisation, of the cortical lesion). The
an injury. Thus, from this perspective, the structural al- mass action principle is alluded to by Flourens8 in
teration of a brain region (or its connections) would nec- his Recherches expérimentales, in which he explained
essarily lead to loss of the associated function. According that it was possible for the cerebral lobes to lose a
to proponents of the holistic doctrine, the fact that many certain portion of their mass, from the front, back,
patients were able (to an extent) to compensate for lost top, or side, without losing their functions, although
brain functions indicated that one brain region was able function was lost if the amount of substance re-
to take on the function of another. This suggested that moved exceeded a certain level.8
the cortex presented a dynamic adaptability, or plasticity.
At the 7th International Medical Congress, held in
This view collides head-on with the mechanistic concep-
tion of the nervous system, in addition to undermining London in 1881, Goltz launched a ferocious attack
the principle of strict localisation, an essential pillar of against the localisationist doctrine. If there is no re-
the connectionist theories popularised by Wernicke and newal of the extirpated tissue, he argued, then the
his followers. functional recovery observed after the ablation re-
sults from the activity of undamaged regions, con-
The concept of dynamic cortical adaptability had pre- stituting reliable evidence that the brain is equipo-
viously been hinted at by Flourens. In Recherches ex- tential: “If a centre that has a specific function is able
périmentales sur les propriétés et les fonctions du systéme to assume the function of another, destroyed centre
nerveux dans les animaux vertébrés (Experimental re- with a different function, then the same section of
searches on the properties and functions of the nerv- the brain performed different functions at the same
ous system in the vertebrate animal),8 he wrote that “it time.”43(p220) Subsequently, he ridicules Hermann
is possible to remove a certain portion of the cerebral Munk’ s (1839-1912) hypothesis explaining post-le-
sion recovery:
K
In the 1860s, Franz Cornelius Donders (1818-1889) proposed that reaction [Munk] supposes that each centre with a
times could be used to measure “the speed of mental processes.” (Donders specific function is to some extent surrounded
FC. Over de snelheid van psychische processen. Onderzoekingen gedaan
in het Physiologisch Laboratorium der Utrechtsche Hoogeschool, Tweede by a fallow field made up of virgin cortical
reeks. 1868;2:92-120). matter, which only begins functioning when the

247
 A. García-Molina, J. Peña-Casanova

Figure 7. Left: diagrams showing the lesion localisation and extension used in different groups of rats in Lashley’s experiments.42 Right:
Karl Lashley (1890-1958).

Figure 8. Left: Friedrich Goltz (1834-1902). Centre: drawings showing the amount of cortical matter removed from the dog presented
by Goltz at the 7th International Medical Congress (1881). Right: illustration of the brain of said dog. The centimetre scale shows the
size of the openings cut in the bone.44

Figure 9. Left: diagram of the localisation of various cortical processes identified by Foerster (1931).50 Right: diagram of the localisation
of functions in the cerebral cortex, according to Kleist (1934).51

248
 Functional organisation of the cerebral cortex: from Gall to Lashley

normally occupied centre is accidentally destroyed. and Karl Kleist (1879-1960) (Figure 9), anti-localisation-
According to this strange doctrine, we would have ist thought became the frame of reference for under-
an extraordinary amount of excess brain matter standing cortical functional organisation in the first half
available that would be spared in the event of of the 20th century.
mutilation in the brain.43(p220)
On 28 December 1911, Franz presented his ideas on the Epilogue
localisationist-connectionist doctrine at a meeting of the Lashley’ s ideas had a profound impact on the under-
Southern Society for Philosophy and Psychology.45 In his standing of the precepts governing behaviour, favour-
presentation, he questioned the evidence from architec- ing the consolidation of American behaviourism thanks
tural neuroanatomy in support of the localisationist-con- to Edward Chace Tolman (1886-1959), Clark Leonard
nectionist doctrine. While he conceded that Korbinian Hull (1884-1952), and Burrhus Frederic Skinner (1904-
Brodmann’ s (1868-1918) cytoarchitectonic studies had 1990), among others. It was not until the 1940s that var-
revealed that the left third frontal gyrus differed struc- ious experimental researchers began to disprove many
turally from the surrounding areas, he maintained that: of Lashley’ s anti-localisationist assertions. This group
The criticisms of von Monakow and of Marie are, included a follower of Lashley at the Yerkes Laboratories
however, too trenchant to be disregarded, and the for Primate Biology, Roger Wolcott Sperry (1913-1994).
negative cases which they have cited are sufficient
evidence that neither the mental processes Sperry conducted various experiments to study the reor-
connected with motor speech nor the supposed ganisation of the motor and sensory nerves, as well as the
cortical speech mechanisms are definitely located in sensory organs (particularly the eye). He concluded that
the part of the brain to which they were assigned by nerve fibres were not interchangeable and that, contrary
Broca, by Wernicke and by their followers.45(p326) to popular belief (and Lashley’ s hypotheses), neural cir-
In 1917, Franz wrote a short article entitled “Cerebral ad- cuits were established very early in the course of deve-
aptation vs. cerebral organology.”46 Citing the research of lopment, and appeared to lack any capacity for modifi-
Vitzou,47,48 he explained that: cation.52 Later, he studied the functional repercussions
of interhemispheric disconnection by surgical section
Assuming that the destructions [performed
of the corpus callosum (the “split-brain experiments”),
by Vitzou] were complete, it is not possible to
understand these recoveries from the standpoint
reporting that the experimental animals presented be-
of cerebral organology, for the cortex is looked at havioural changes.53,54 These findings contradicted those
as a locus of certain physiological processes which conducted by Andrew Akelaitis (1904-1955) in human
give rise to or which are coincident with the mental subjects,L as well as the results of his old mentor at the
states.46(p138) Yerkes Laboratories for Primate Biology. Lashley argued
After describing various cases that cast doubt on the that the corpus callosum was a merely skeletal structure,
straightforward relationship between region and func- since he “could find no function for them.”55(p132)
tion, and supporting the possibility of post-injury func- Sperry’ s split-brain experiments sparked the interest of
tional recovery, he concluded: the American neurologist Norman Geschwind (1926-
The destruction of a part of the cerebrum which is 1984).M Geschwind was very critical of the clinical value
followed by an obvious defect does not mean that of anti-localisationist approaches, and considered the
that part of the cerebrum is solely concerned with possibility that the disconnection syndromes observed
that function […]. When these facts are admitted, in animals could be extrapolated to humans (see his
as they must be admitted, the whole structure of
cerebral organology breaks down. The histological
localization of function which has been in vogue
L
In 1939, the neurosurgeon William Perrine Van Wagenen (1897-1961)
conducted the first corpus callosotomy procedures to treat drug-resistant
takes its true place as a histological differentiation epilepsy. Van Wagenen’s colleague, the psychiatrist Akelaitis, observed no
of an anatomical nature, without the functional remarkable cognitive changes after the procedures, concluding that surgical
section of this set of nerve fibres could be performed without fear of adverse
implications which have been assumed.46(p140) effects.
M
Geschwind was the master and mentor of brilliant neurologists including
Despite the obstinacy of such authors as Salomon Marsel Mesulam, António Damásio, Frank Benson, François Boller, and
Henschen (1847-1930),49 Otfrid Foerster (1873-1941),50 Albert Galaburda.

249
 A. García-Molina, J. Peña-Casanova

description of the patient PJK56). In 1965, he published an 7. Gall FJ, Spurzheim JG. Anatomie et physiologie du système
influential article in two parts, entitled “Disconnection nerveux en général et du cerveau en particulier. Paris: Chez
F. Schoell; 1810.
syndromes in animals and man.” In the first part, he pro- 8. Flourens P. Recherches expérimentales sur les propriétés et
posed that aphasias, apraxias, and agnosias be reinter- les fonctions du systéme nerveux dans les animaux vertébrés.
preted in terms of anatomical disconnections.57 In the Paris: Chez Crevot, Librare-Éditeur; 1824. [Flourens P.
Experimental researches on the properties and functions of
second, he argued that knowledge of anatomical-clini- the nervous system in the vertebrate animal. Kann J, transl.
cal correlations was essential to understanding cortical In: Readings in the history of psychology. Dennis W, ed. New
physiology, and defended the localisationist-connec- York: Appleton-Century-Crofts, Inc.; 1948. p. 129-139].
tionist doctrine: 9. Flourens P. Examen de la phrénologie. Paris: Paulin; 1845.
[Flourens P. Phrenology examined. De Lucena Meigs C,
For the past forty years there have been schools transl. Philadelphia (PA): Hogan & Thompson; 1846].
of thought which have stressed the importance 10. Andral G. Clinique médicale. Paris: Deville Cavellin; 1833.
[Andral G. Medical clinic. Spillan D, transl. Philadelphia
of thinking of the patient as a whole, of seeing
(PA): Barrington & Haswell; 1843].
his responses as those of an integrated unitary 11. García-Molina A, Peña-Casanova, J. Jean-Baptiste
structure […]. I have attempted to show that many Bouillaud y el dogma de la tercera circunvolución frontal
disturbances of the higher functions of the nervous izquierda. Neurosci Hist. 2023;11:33-43.
system, such as the aphasias, apraxias, and agnosias 12. Bouillaud JB. Recherches cliniques propres à démontrer
que la perte de la parole correspond à la lésion des lobules
may be most fruitfully studied as disturbances
antérieurs du cerveau, et à confirmer l’ opinion de M. Gall
produced by anatomical disconnexion of primary sur le siège de l’ organe du langage articulé. Arch Gen Med.
receptive and motor areas from one another.58(p637-40) 1825;8:25-45.
13. Bouillaud JB. Exposition de nouveaux faits à l’ appui de
This marked the birth of the so-called neo-associationist l’ opinion qui localise dans les lobules antérieurs du cerveau
school, a theoretical school of thought that recovered le principe législateur de la parole; précédé de l’ examen des
(and updated) the connectionist doctrine of Wernicke objections dont cette opinion a été le sujet. Bull Acad Med.
1839;4:282-328.
and constitutes the foundations of contemporary
14. Bouillaud JB. Recherches cliniques propres à démontrer
behavioural neurology and neuropsychology. que le sens du langage articulé est le principe coordinateur
des mouvements de la parole résident dans les lobules
Conflicts of interest antérieurs du cerveau. Memoire lu à l’ Académie nationale
de Médecine, le 22 février et 7 mars 1848. Paris: Chez J.-B.
The authors have no conflicts of interest to declare. This Baillière; 1848.
study has received no public or private funding. 15. Bouillaud JB. Discussion sur la faculté du langage articulé.
Bull Acad Imp Med. 1864-65;30:575-600, 604-38, 724-80.
References 16. Broca P. Perte de la parole; ramollissement chronique et
destruction partielle du lobe antérieure gauche du cerveau.
1. García-Molina A, Peña-Casanova J. Fundamentos Bull Mem Soc Anthropol Paris. 1861;2:235-8.
17. Broca P. Nouvelle observation d’ aphémie produite par une
históricos de la neuropsicología y la neurología de la
lésion de la moitié postérieure des deuxième et troisième
conducta. Teià (ES): Test Barcelona Services; 2022.
circonvolutions frontales. Bull Soc Anat Paris. 1861;6:398-
2. Feindel W, ed. The anatomy of the brain and nerves. 407.
Montreal (CA): McGill University Press; 1965. 18. Broca P. Sur le siège de la faculté du language articulé. Bull
3. Swedenborg E, Tafel RL. The brain, considered anatomically, Mem Soc Anthropol Paris. 1865;6:377-93.
physiologically, and philosophically. Volume I. London: 19. Hécaen H, Lanteri-Laura G. Evolution des connaissances et
Speirs; 1882. des doctrines sur les localisations cérébrales. Paris: Desclée
4. Clarke E, O’ Malley CD. The human brain and spinal cord. de Brouwer; 1977.
2nd ed. San Francisco: Norman; 1996. 20. Compte rendu des séances de la Société de Biologie. Paris:
Chez Adrien Delehaye; 1875.
5. Lazar JW. Nineteenth- and twentieth-century brain maps
21. Charcot JM. Oeuvres complètes: leçons sur les localisations
relating to locations and constructions of brain functions. J
dans les maladies du cerveau et de la moelle épinaire. Vol.
Hist Neurosci. 2022;31:368-93. 4. Paris: Bureaux du Progrès Médical; 1875. [Charcot JM.
6. Gall FJ. Philosophisch-medicinische Untersuchungen über Lectures on the localization of cerebral and spinal diseases:
Natur und Kunst im kranken und gesunden Zustande des delivered at the Faculty of Medicine of Paris. Hadden WB,
Menschen. Vienna (AT); Rudolph Gräffer; 1791. transl. London: New Sydenham Society; 1883].

250
 Functional organisation of the cerebral cortex: from Gall to Lashley

22. Charcot JM, Pitres A. Étude critique et clinique de la 41. Lashley KS. Brain mechanisms and intelligence: a
doctrine des localisations motrices dans l’ écorce des quantitative study of injuries to the brain. Chicago:
hémisphères cérébraux de l’ homme. Paris: Félix Alcan; University of Chicago Press; 1929.
1883. 42. Lashley KS, Wiley LE. Studies of cerebral function in
23. Fritsch G, Hitzig E. Über die elektrische Erregbarkeit des learning IX. Mass action in relation to the number of
Grosshirns. Arch für Anat Physiol und Wissenschaftliche elements in the problem to be learned. J Comp Neurol.
Med. 1870;36:30032. 1933;57:3-55.
24. Ferrier D. The functions of the brain. London: Elder Smith; 43. Mac Cormack W, Makins GH. Transactions of the
1878. International Medical Congress. Vol. 1. London: J.
25. Munk H. Über die Functionen der Grosshirnrinde: Kolckmann; 1881.
gesammelte Mittheilungen. Berlin: August Hirschwald; 44. Gowers WR, Klein E, Schaefer AE, Langley JN. Preliminary
1881. report. In: Transactions of the International Medical
26. Bianchi L. The functions of the frontal lobes. Brain. Congress. Vol 1. Mac Cormack W, ed. London: J. W.
1895;18:497522. Kolckmann; 1881. p.242a-242d.
27. Hitzig E. Untersuchungen über das Gehirn: Abhandlungen 45. Franz SI. New phrenology. Science. 1912;35:321-8.
physiologischen und pathologischen Inhalts. Berlin: Verlag 46. Franz SI. Discussion: cerebral adaptation vs cerebral
Von August Hirschwald; 1874. organology. Psychol Bull. 1917;14:137-40.
28. Bartholow R. Experimental investigations into the functions 47. Vitzou AN. Sur les effets de l’ ablation totale en un temps
of the human brain. Am J Med Sci. 1874;134:305-13. d’ un hémisphère cerebral chez le chien. Arch Physiol Norm
29. Zago S, Ferrucci R, Fregni F, Priori A. Bartholow, Sciamanna, Pathol. 1893;2:265-78.
Alberti: pioneers in the electrical stimulation of the exposed 48. Vitzou AN. Récupération de la vue perdue à la suite d’ une
human cerebral cortex. Neuroscientist. 2008;14:521-8. première ablation totale des lobes occipitaux chez les
30. Meynert T. Der Bau der Großhirnrinde und seine singes. J Physiol. 1899(Suppl);23:40-74.
örtlichen Verschiedenheiten nebst einem pathologisch- 49. Henschen S. Uber Sprach, Musik, und Rechenmechanismen
anatomischen Korollarium. Neuwied (DE): J.H. Heuser; und ihre Lokalisationen im Grosshirn. Zeitschrift für die
1868. gesamte Neurologie und Psychiatr. 1919;52:273-98.
31. Wernicke C. Der aphasische Symptomencomplex: eine 50. Foerster O. The cerebral cortex in man. Lancet.
psychologiische Studie auf anatomisher Basis. Breslau 1931;218:309-12.
(DE): Cohn & Weigert; 1874. 51. Kleist K. Gehirnpathologie vornehmlich auf Grund der
32. Wernicke C. Einige neuere Arbeiten über Aphasie. Kriegserfahrungen. Leipzig (DE): Verlag von Johann
Fortschr Med. 1885;3:824-30. [Wernicke C. Recent Works Ambrosius Barth; 1934.
on Aphasia. In: Philosophy and neurosciences: a reader. 52. Puente A. Roger W Sperry, from neuroscience to
Bechtel W, Mandik P, Mundal J, Stufflebeam R, eds. Oxford: neurophilosophy. In: Stringer AY, Cooley EL, Christensen
Blackwell; 2001. p.100-111]. AL, eds. Pathways to prominence in neuropsychology:
33. García-Molina A, Peña-Casanova J. La escuela de Wernicke. reflections of twentieth century pioneers. New York:
Neurosci Hist. 2022;10:178-85. Psychology Press; 2002. p. 63-75.
34. Head H. Aphasia and kindred disorders of speech. 53. Sperry RW. Cerebral organization and behavior: the
Cambridge: Cambridge University Press; 1926. split brain behaves in many respects like two separate
35. Marie P. Existetil chez l’ homme, des centres préformés brains, providing new research possibilities. Science.
ou innés du langage? In: Wilson SAK, ed. Questions 1961;133:1749-57.
neurologiques d’ actualité. Paris: Masson et Cie; 1922. p. 54. Volz LJ, Gazzaniga MS. Interaction in isolation: 50 years of
527551. insights from split-brain research. Brain. 2017;140:2051-60.
36. Moutier F. L’ aphasie de Broca. Paris: Steinheil; 1908. 55. Lashley, K. S. The problem of serial order in behavior.
37. Jackson JH. On the nature of the duality of the brain. In: Jeffress LA, ed. Cerebral mechanisms in behavior; the
Medical Press and Circular. 1874;17:19-21, 41-44, 63-65. Hixon Symposium. New York: Wiley; 1951. p.112-46.
38. Tarnovsky V. Some forms of the syphilitic aphasia. Voenno- 56. Geschwind N, Kaplan E. A human cerebral
Meditsynsky Jurnal. 1867;100:225-96. deconnection syndrome: a preliminary report.
39. Von Bertalanffy, L. General theory of systems: application Neurology. 1962;12:675-85.
to psychology. Soc Sci Inf. 1967;6:125-36. 57. Geschwind N. Disconnection syndromes in animals and
40. Von Monakow C. Die Lokalisation im Grosshirn und der man. I. Brain. 1965;88:237-94.
Abbau der Funktion durch kortikale Herde. Wiesbaden 58. Geschwind N. Disconnection syndromes in animals and
(DE): Bergmann; 1914. man. II. Brain. 1965;88:585-644.

251