See discussions, stats, and author profiles for this publication at: https://www.researchgate.

net/publication/6415717

Paul Broca's historic cases: High resolution MR imaging of the brains of

Leborgne and Lelong

Article  in  Brain · June 2007

DOI: 10.1093/brain/awm042 · Source: PubMed

CITATIONS READS
427 6,183

4 authors, including:

Nina Dronkers Odile Plaisant

VA Northern California Health Care System & University of California, Davis Paris Descartes, CPSC
168 PUBLICATIONS   21,359 CITATIONS    99 PUBLICATIONS   2,084 CITATIONS   

SEE PROFILE SEE PROFILE

Emmanuel Alain Cabanis

Académie Nationale dee Médecine, Paris, France, Pierre and Marie Curie University - …
377 PUBLICATIONS   9,566 CITATIONS   

SEE PROFILE

Some of the authors of this publication are also working on these related projects:

The Teaching of the Anatomical Sciences View project

_Personality assessment / Personality and adaptation_ View project

All content following this page was uploaded by Nina Dronkers on 19 June 2014.

The user has requested enhancement of the downloaded file.

 Brain Advance Access published April 2, 2007

doi:10.1093/brain/awm042 Brain (2007) Page 1 of 10

OCC ASIONAL PAPER

Paul Broca’s historic cases: high resolution MR
imaging of the brains of Leborgne and Lelong
N. F. Dronkers,1 O. Plaisant,2 M. T. Iba-Zizen3 and E. A. Cabanis4
1
VA Northern California Health Care System, University of California, Davis and University of California, San Diego, CA,
USA, 2Faculte¤ de Me¤decine, Universite¤ Paris-Descartes, Paris V and APHP GH Pitie¤-Salpe“trie're, Epileptology Department,
Paris, France, 3Universite¤ Pierre et Marie Curie Paris VI, Faculte¤ Pierre et Marie Curie and Centre Hospitalier National
d’Ophtalmologie des Quinze-Vingts, Neuroimaging Department, Paris, France and 4Universite¤ Pierre et Marie Curie Paris VI,
Faculte¤ Pierre et Marie Curie, Centre Hospitalier National d’Ophtalmologie des Quinze-Vingts, Neuroimaging Department
and Acade¤mie Nationale de Me¤decine, Paris, France

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Correspondence to: Nina F. Dronkers, PhD, Center for Aphasia and Related Disorders, VA Northern California Health Care
System, 150 Muir Road (126s), Martinez, CA 94553, USA
E-mail: [email protected]

In 1861, the French surgeon, Pierre Paul Broca, described two patients who had lost the ability to speak after
injury to the posterior inferior frontal gyrus of the brain. Since that time, an infinite number of clinical and
functional imaging studies have relied on this brain^behaviour relationship as their anchor for the localization
of speech functions. Clinical studies of Broca’s aphasia often assume that the deficits in these patients are due
entirely to dysfunction in Broca’s area, thereby attributing all aspects of the disorder to this one brain region.
Moreover, functional imaging studies often rely on activation in Broca’s area as verification that tasks have
successfully tapped speech centres. Despite these strong assumptions, the range of locations ascribed to
Broca’s area varies broadly across studies. In addition, recent findings with language-impaired patients have
suggested that other regions also play a role in speech production, some of which are medial to the area origin-
ally described by Broca on the lateral surface of the brain. Given the historical significance of Broca’s original
patients and the increasing reliance on Broca’s area as a major speech centre, we thought it important
to re-inspect these brains to determine the precise location of their lesions as well as other possible areas of
damage. Here we describe the results of high resolution magnetic resonance imaging of the preserved brains
of Broca’s two historic patients. We found that both patients’ lesions extended significantly into medial regions
of the brain, in addition to the surface lesions observed by Broca. Results also indicate inconsistencies
between the area originally identified by Broca and what is now called Broca’s area, a finding with significant
ramifications for both lesion and functional neuroimaging studies of this well-known brain area.

Keywords: aphasia; Broca; history; language; magnetic resonance imaging

Abbreviations: MRI ¼ magnetic resonance imaging; CT ¼ computerized tomography; S(RH) ¼ sagittal image of the right
hemisphere; S(LH) ¼ sagittal image of the left hemisphere; SLF ¼ superior longitudinal fasciculus
Received June 30, 2006. Revised December 19, 2006. Accepted February 13, 2007

Introduction
Most neuroscientists would agree that the foundations of Clarke and O’Malley, 1968; Schiller, 1992; Monod-Broca,
modern neuropsychology and cognitive neuroscience were 2005). Amidst these discussions, Broca was consulted about
laid by the French surgeon, anatomist and anthropologist, a 51-year-old patient by the name of Leborgne with
Paul Broca, in the 1860s. At that time, Broca and his multiple neurological problems, who had been without
colleagues in Paris were discussing a lingering claim of the any productive speech for many years. Every time Leborgne
phrenologists that language functions were located in the attempted to utter a phrase or respond to a question,
frontal lobes of the brain (Gall and Spurtzheim, 1809; he could only produce a single repetitive syllable, ‘tan’.

ß The Author (2007). Published by Oxford University Press on behalf of the Guarantors of Brain. All rights reserved. For Permissions, please email: [email protected]
Page 2 of 10 Brain (2007) N. F. Dronkers et al.

He could vary the intonation of the sound but was not able
to produce any recognizable words or phrases. Broca saw
Leborgne’s lack of speech as a test case for the question of
language localization in the frontal lobes, since the patient
clearly had no productive language. Leborgne died of his
ailments several days later and, at autopsy, a lesion was
found on the surface of the left frontal lobe as Broca had
suspected. He presented his finding to the Anthropological
Society (Broca, 1861b) where some of the earlier discussions
had taken place, and to the more established Anatomical
Society of Paris (Broca, 1861c) several months later. The
finding was met with enthusiasm and taken as support for
the premise that cognitive functions could be localized to
specific convolutions of the brain.
A few months later, Broca encountered a second patient,
Lelong, who also exhibited reduced productive speech as
the result of a stroke 1 year before. This 84-year-old patient

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Fig. 1 Three-dimensional MRI reconstruction of the lateral left
could say only five words, ‘oui’ (‘yes’), ‘non’ (‘no’), ‘tois’ hemisphere of a normal in vivo brain. The location now considered
(a mispronunciation of ‘trois’ (‘three’) which he used to Broca’s area includes the pars opercularis (Brodmann’s area 44,
represent any number), ‘toujours’ (‘always’) and ‘Lelo’ anterior to the precentral sulcus) and the pars triangularis
(Brodmann’s area 45, between the ascending and horizontal limbs
(a mispronunciation of his own name). At autopsy, Lelong
of the sylvian fissure) in the posterior inferior frontal gyrus.
was also found to have a lesion in approximately the
same region of the lateral frontal lobe as the first case, and
Broca reported it to the Anatomical Society as an important fluency, articulation, word-finding, repetition and producing
case, confirming the localization of speech to this area. and comprehending complex grammatical structures, both
He wrote, orally and in writing.
A subset of patients with Broca’s aphasia have a more
The integrity of the third frontal convolution (and perhaps of the severe form of the disorder. These patients are not able to
second) seems indispensable to the exercise of the faculty of produce much in the way of meaningful words or phrases,
articulate language . . . I found that in my second patient, but typically can only articulate the same recurring sounds,
the lesion occupied exactly the same seat as with the first - words or phrases that are uttered each time they attempt
immediately behind the middle third, opposite the insula and to speak. It is apparent that Broca’s two patients also
precisely on the same side (translation ours, Broca, 1861a). suffered from this same severe form of aphasia. The first
patient, Leborgne, could only produce the jargon syllable
Broca was subsequently presented with other cases of
‘tan’. Broca himself wrote,
speech disturbance with lesions encompassing the third
convolution of the frontal lobe and, within 4 years, had He could no longer produce but a single syllable, which he
realized that most of the cases were lesioned on the left side usually repeated twice in succession; regardless of the question
of the brain (Broca, 1865). Though Marc Dax may have asked him, he always responded: tan, tan, combined with varied
suggested the same relationship earlier (Joynt and Benton, expressive gestures. This is why, throughout the hospital, he is
1964), Broca’s careful documentation was pivotal in known only by the name Tan (Broca, 1861c).
establishing the important connection between speech and
the inferior frontal gyrus of the left cerebral hemisphere. Broca’s second patient, Lelong, was able to produce only
the same five words. Broca believed that these patients
understood what was said to them and that their language
Broca’s area and Broca’s aphasia was therefore intact. He called this disorder ‘aphemia’
Since Broca’s time, the approximate region he identified has referring to the absence of speech and believed it to be
become known as Broca’s area, and the deficit in language independent of language dysfunction. Later, Trousseau
production as Broca’s aphasia. Broca’s area is now typically coined the term ‘aphasia’ (Broca, 1864), and over time,
defined in terms of the pars opercularis and pars triangularis the term ‘Broca’s aphasia’ evolved and began to include the
of the inferior frontal gyrus, represented in Brodmann’s many different symptoms of Broca’s aphasia known today.
(1909) cytoarchitectonic map as areas 44 and 45 (Fig. 1). The The relationship between Broca’s aphasia and Broca’s
disorder is considered to be a complex of several symptoms area continued to be studied by numerous researchers and
that, together, contribute to the syndrome of Broca’s aphasia clinicians. Pierre Marie noted early on that the type of
(Goodglass, 1993; Benson and Ardila, 1996; Dronkers and disorder described by Broca was not always related to
Ogar, 2003). These symptoms can include problems with lesions in Broca’s area, but rather to lesions involving more
Neuroimaging of Broca’s historic cases Brain (2007) Page 3 of 10

medial structures, including the insula and basal ganglia of aphasic patients had not been made, in particular
(Marie, 1906). Others also questioned Broca’s conclusions with regard to these deep lesions. The brain of Broca’s
regarding the localization of speech functions (e.g. Brown- second patient, Lelong, was never scanned and the
Sequard, 1877; Bramwell, 1898; Moutier, 1908). Modern extent of the damage in this second case was never
lesion studies (e.g. Mohr, 1976; Mohr et al., 1978; Naeser determined.
and Hayward, 1978; Kertesz et al., 1979; Schiff et al., 1983; We had the unique opportunity to scan the brains of both
Basso et al., 1985; Murdoch et al., 1986; Alexander et al., Leborgne and Lelong using high-resolution volumetric MRI.
1990) have found that Broca’s aphasia is caused by This allowed us to view the brains in three dimensions and to
large lesions encompassing not necessarily Broca’s area, examine the extent of both cortical and subcortical lesions in
but surrounding frontal cortex, underlying white matter, close detail. In particular, we were interested in the exact
the insula, basal ganglia and parts of the anterior superior location of the lesion in the frontal lobe in relation to what is
temporal gyrus. This implies that other brain regions also now called Broca’s area and the extent of subcortical
participate in speech production besides Broca’s area. involvement. The relation of these lesions to the deficits
Considering that the syndrome of Broca’s aphasia affects observed in these two historic cases is discussed later.
so many different components of language, it is not
surprising that other brain areas would also be involved.
More specific roles for some of these deeper areas have Gross anatomy of the brains

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

been discussed for parts of the insula (Dronkers, 1996),
The brains were removed from the Musee Dupuytren in
basal ganglia (e.g. Crosson, 1985; Radanovic and Scaff,
Paris, France by the museum director at the time, Prof. de
2003), medial subcallosal fasciculus and periventricular
Saint Maur. They were transported under his supervision to
white matter (Naeser et al., 1989) and the arcuate/superior
the Neuroradiology Service of the Centre Hospitalier
longitudinal fasiculus (Dronkers, 1993).
It is noteworthy that Broca provided minimal discussion National d’Ophtalmologie des Quinze-Vingts in Paris
of such medial lesions in his original cases. This is because where they were photographed and scanned.
he made the decision not to dissect the brains of his two Figure 3 (panels A and B) shows lateral views of the
patients, but to preserve them in alcohol and place them in brains of the two patients as photographed. A portion of
a Paris museum for posterity. Thus, Broca focused on Leborgne’s lesion was clearly visible in the inferior frontal
observable lesions to the cortical surface of the brain. gyrus, most noticeably in its middle third with additional
However, Broca did perform a provisional analysis of the atrophy in the posterior third, just above the sylvian fissure.
brain of Leborgne, and confirmed that the lesion included Furthermore, modest atrophy was observed in the middle
‘the small convolutions of the insula, and the corpus frontal gyrus and the anterior superior temporal gyrus.
striatum’. . . ‘As to the deep parts . . . I could half-way Upon close inspection, deformed and necrotic gyri were
examine the inner surface of the anterior horn of the also noted in the anterior, inferior parietal lobe. The right
lateral ventricle . . .’ [translated by Schiller (1992), p. 185]. hemisphere, not seen in this figure, was intact. These
Thus, he was aware that the lesion included more medial findings are largely consistent with Broca’s original
structures but could not determine the precise extent of the description of the lesion at autopsy (Broca, 1861c).
damage without dissecting the brain. Importantly, the most significant area of damage in the
Since the advent of neuroimaging technology, the brain frontal lobe was in the middle third of the inferior frontal
of Leborgne has been scanned on two occasions, both in gyrus, not in the posterior third of the gyrus, now typically
the Radiology Service of E. A. Cabanis and M. T. Iba-Zizen. designated as Broca’s area. The posterior third clearly shows
The first was a computerized tomography (CT) study abnormalities but is not the most extensively damaged.
described by Castaigne and colleagues (Castaigne et al., Broca considered the posterior half of the inferior frontal
1980) with further descriptions provided by Signoret gyrus lesion to be related to the articulatory deficit. Thus,
(Signoret et al., 1984). Even with the comparatively low the modern view of the location of Broca’s area, restricted
resolution of these early-generation CT scans, it was clear to the posterior third of the inferior frontal gyrus, is not
that Leborgne’s lesion was much larger than originally entirely consistent with Broca’s own determination that
reported by Broca (Fig. 2). It extended medially, affecting included far more of this gyrus. This has ramifications for
the left basal ganglia as well as the entire insula. In 1994, both lesion and functional neuroimaging studies ascribing
Leborgne’s brain was scanned with magnetic resonance specific functions to Broca’s area, as interpretations of its
imaging (MRI) to demonstrate the use of this technology exact location may have changed over time.
in the dissection of conserved brains (Cabanis et al., 1994). Lateral views of Lelong’s brain are shown in Fig. 3
These higher resolution images showed the extent of (panels C and D). The cortex is severely atrophied. The
the deep lesion much more clearly, and that critical sylvian fissure has become so widened that the insula is,
brain areas other than Broca’s area were also affected in abnormally, almost completely exposed. Broca’s writings
this patient. Still, a direct comparison to modern indicate that Lelong had resided at the hospital for the 8
interpretations of Broca’s area and to current lesion studies years previous to his stroke because of dementia. The report
Page 4 of 10 Brain (2007) N. F. Dronkers et al.

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Fig. 2 Rows 1^ 4: Samples of the first neuroradiological images of Leborgne’s brain (1978 ^79) with diagrams. These CT images were the
first radiographic scans obtained on this historic brain and, though not of high resolution, gave an indication of the medial extension of the
lesion. The CT slices are compared with Dejerine’s diagrams in the three planes. Row 5: MRI sagittal slices (1999) discriminating the cortex
and grey and white matter with higher resolution and demonstrating the use of MR imaging in preserved specimens.
of a dementing disorder is consistent with the finding of lesion, Lelong’s is also inconsistent with the location of
atrophy in this brain of an 84-year-old. Broca’s area as it is defined today. In this case, the lesion
Lelong’s brain also shows evidence of a stroke that actually spares the anterior portion of modern Broca’s area.
affected half of the pars opercularis in the posterior, inferior This is a significant finding as it implies that this second
frontal gyrus, sparing the pars triangularis. Like Leborgne’s brain on which current theories of localization are based
Neuroimaging of Broca’s historic cases Brain (2007) Page 5 of 10

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Fig. 3 Photographs of the brains of Leborgne and Lelong, Paul Broca’s first two aphasic patients. (A) Lateral view of the brain of the
first patient, Leborgne. The external lesion is clearly visible in the inferior frontal lobe. The softening in the area superior and posterior
to the lesion suggests further cortical and subcortical involvement. (B) Close-up of the visible lesion in Leborgne’s brain. (C) Lateral
view of Broca’s second patient, Lelong. The frontal, temporal and parietal lobes have retracted due to severe atrophy, exposing the
insula. (D) Close-up of the visible lesion in Lelong’s brain. Note that only the most posterior part of what is currently called
Broca’s area is infarcted; the anterior portion is completely spared.

does not have a lesion encompassing the entire area we now rendering of the brain. The images demonstrate significant
call Broca’s area. Thus, gross re-examination of these two damage throughout the left hemisphere, both cortically and
important brains has revealed that the area defined by subcortically. The left hemisphere is clearly smaller and
Broca as critical for articulation is not necessarily the same distorted due to the destruction of cortex and white matter
as the area currently described. throughout the hemisphere. Sagittal, axial and coronal slices
through the brain reveal lesions in the left inferior
MRI findings frontal gyrus (slices A2, C1, S1), deep inferior parietal
lobe (slices A4, C4, S1–3) and anterior superior temporal
Leborgne and Lelong’s brains were imaged with a 1.5 tesla
lobe (slices A2, C1–2, S1). In addition, there is extensive
MRI scanner (GE Signa Echospeed HDX LCC Magnet
8.2.5). Several sequences were conducted, including a fast subcortical involvement including the claustrum, putamen,
spin echo series (512
512 matrix, zip 1024). This series globus pallidus, head of the caudate nucleus and internal
normally appears T1 weighted, but, in these cases had to be and external capsules (slices A2–3, C2–3, S2–3). The insula
scanned differently from a living brain because of contrast is completely destroyed (slices A3, C2–3, S2). The entire
differences caused by the solution in which the brains were length of the superior longitudinal fasciculus is also
preserved. obliterated (slices A4, C2–5, S2–3), along with other
Representative MRI images of Leborgne’s brain are frontal-parietal periventricular white matter. The medial
shown in Fig. 4. Coloured markings illustrate the major subcallosal fasciculus is also affected (slices A2, C2).
sulci of the brain to highlight the gyri and key structures The right hemisphere is unaffected (in particular, see
and clarify the extent of the damage. Sulcal locations were slices S5–8) and serves as an excellent comparison to the
determined by two neuroanatomists, independently, and damaged left hemisphere of this preserved brain.
transferred to individual slices by matching coordinates The extent of the damage in the left hemisphere of
from tracings on the lateral surface of a 3D computerized Leborgne’s brain is most obvious when comparing the
Page 6 of 10 Brain (2007) N. F. Dronkers et al.

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Fig. 4 High-resolution MRI of the preserved brain of Leborgne with representative slices throughout the brain. The first row shows
photographs of the lateral and superior surfaces of the brain, with lines indicating the slices shown below. Row A shows axial slices, Row C
coronal slices, and Row S sagittal slices through the left and intact right hemisphere for comparison with each other. In the axial and
coronal planes, the left hemisphere appears on the left side of the images. The following structures are delineated: interhemispheric/long-
itudinal fissure (orange), central sulcus/Rolandic fissure (dark blue), sylvian/lateral fissure (aqua), inferior frontal sulcus (red), superior frontal
sulcus (yellow), frontomarginal sulcus (pink), superior temporal sulcus (light green) and inferior temporal sulcus (brown). Sagittal slices S3
and S4 show the superior portion of the right hemisphere crossing over the midline due to extensive damage in the left hemisphere.
Neuroimaging of Broca’s historic cases Brain (2007) Page 7 of 10

sizes of the two hemispheres on the MRI images. The left sides of Leborgne’s brain can also be seen in the axial (A)
hemisphere, as measured from the midline to the lateral and coronal (C) slice in which both hemispheres are
surface, is up to 50% smaller than the right hemisphere. represented. In contrast, the cerebellum, occipital lobes,
The coronal slices, in particular, exhibit extensive damage inferior temporal lobes, superior frontal and parietal lobes
in the left frontal lobe when examining the distance from and hippocampi are intact in both hemispheres.
the interhemispheric fissure (in orange) to the Sylvian Figure 8 highlights the lesions in the superior long-
fissure (in aqua) (slices C2–3). Posterior damage is itudinal fasciculus in the brain of Lelong. Sagittal, coronal
apparent in the reduced distance from the interhemispheric and axial slices through the critical areas are shown, along
fissure to the superior temporal sulcus (slices C4–5). Several with a 3D computer reconstruction with crosshairs to
of the inferior parietal gyri and deep structures are missing. demonstrate the positions of the coronal and axial slices.
It is difficult to tell from examining the 3D images or even Damage to the superior longitudinal fasciculus is evident on
the brain itself whether the supramarginal and angular gyri all slices.
are affected due to the amount of damage and effacement
of identifying landmarks.
MR images of the left hemisphere of Lelong’s brain, as Conclusions
well as 3D reconstructions illustrating the lateral and Re-examination of the brains of Paul Broca’s two historic
superior surfaces are shown in Fig. 5. The right hemisphere cases has yielded several interesting findings. First, high-

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

was not preserved with the left hemisphere specimen. resolution MRI images showed that the lesions in these two
Images were computed from 60 sagittal slices at 512
512 important patients extended far deeper than Broca was able
resolution, and coloured markings again denote major sulci. to report and suggests that other areas besides Broca’s area
Sagittal, axial and coronal slices confirm the severe atrophy may also have contributed to these patients’ speech deficits.
noted earlier and the lesion in the posterior part of Broca’s Both cases had lesions extending into the superior long-
area on the inferior frontal gyrus (slices A6–7, C4, S1). The itudinal fasciculus (SLF), a large intrahemispheric fibre tract
lesion involves the posterior part of the pars opercularis, (Cabanis et al., 2004) that connects posterior and anterior
while the anterior half of this structure and the entire pars language regions (Geschwind, 1972).
triangularis are completely spared. This can be seen in Though lesions to Broca’s area alone may cause temporary
closer detail in the white square of Fig. 6. In addition, small speech disruption, they do not result in severe and persisting
but distinct lesions are present in the superior longitudinal speech arrest (Penfield and Roberts, 1959; Mohr et al., 1978).
fasciculus above the insula and lateral to the anterior horn Therefore, it is possible that the aphemia characterized by
of the left lateral ventricle (slices A7, C4–5, S3). The insula, Broca as an absence of productive speech was also influenced
though severely atrophied, is not specifically lesioned in this by the lesions in the region of the superior longitudinal
case nor are other deep structures, including the medial fasciculus. Damage to Broca’s area in both cases may, in
subcallosal fasciculus. There are also abnormalities in the isolation, have resulted in milder speech deficits, but would
white matter pathways in the left temporal lobe (slices A3, not likely have caused the complete and persisting disruption
C7, S2–3) that may have been caused by small strokes. of productive speech in these cases.
Though the current findings provide additional anato-
mical information, they by no means detract from Broca’s
Commonalities between the cases phenomenal discovery. Because he elected not to slice the
Of interest is the fact that both patients had what Broca brains, Broca could not have known the extent of
described as ‘aphemia’, a disruption in the patients’ underlying damage in his patients and the role it might
voluntary speech production ability. A comparison of the play in their speech disorders. Broca understood the lesion
anatomy of the two patients’ lesions reveals that both also extended subcortically in Leborgne, but could not deter-
had lesions in an important white matter pathway, the mine how medially or posteriorly it extended. Broca wrote
arcuate/superior longitudinal fasciculus, the long fibre tract that Leborgne’s disease was progressive with the aphemia
that connects posterior and anterior brain areas and being the most persistent deficit. Since the most apparent
includes fibres of the arcuate fasciculus, previously damage was in the inferior frontal convolution, he
implicated in language processing (Geschwind, 1972; concluded that this area was the first affected and thus
Cabanis et al., 2004). The destruction of this region in the cause of the articulation deficit. This conclusion was
the case of Leborgne can be seen most clearly in Fig. 7 consistent with neurological theory of his time, and Broca
where the sagittal image of the right hemisphere [S(RH)] had no reason to consider other areas as the source of the
can be compared to the matched sagittal slice through the speech disorder.
left hemisphere [S(LH)]. The white matter tracts seen so Though it may not play as extensive a role as once thought,
clearly in the right hemisphere are entirely absent in the Broca’s area is certainly involved in the execution of
left. There is complete destruction of the deep frontal and articulatory movements. Patients with newly acquired lesions
parietal areas through which the superior longitudinal restricted to Broca’s area tend to be mute or exhibit speech
fasciculus passes. A striking comparison between the two distortions for a few weeks after the injury (Penfield and
Page 8 of 10 Brain (2007) N. F. Dronkers et al.

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Fig. 5 High-resolution magnetic resonance images of the preserved left hemisphere of the brain of Lelong with representative slices
throughout the brain. The first row shows computerized 3D reconstructions of the lateral and superior surfaces of the brain with lines
indicating the locations of the slices below. The widened sulci are easily visible and indicate severe atrophy. Row A depicts axial slices,
Row C coronal slices and Row S sagittal slices through the left hemisphere. In these images, the colours have been reversed to enhance the
contrast; cortex appears white and white matter appears dark. They have been flipped horizontally so that the lateral cortex of the left
hemisphere is on the left side of the slice. Coloured lines again show the major sulci of the brain (see Fig. 4 for color codes).
Neuroimaging of Broca’s historic cases Brain (2007) Page 9 of 10

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Fig. 7 Views of the damage to the superior longitudinal fasciculus
Fig. 6 Views of the damage to Broca’s area in Lelong’s brain. Slices in the brain of Leborgne. The affected areas can be seen in sagittal
through the affected area can be seen in sagittal slice S, in coronal slice S (LH) through the left hemisphere, in coronal slice (C) and in
slice (C), and axial slice (A).Three-dimensional reconstruction of the axial slice (A). Sagittal slice S (RH) through the intact right hemi-
left hemisphere is also shown, computed from 60 sagittal slices at sphere illustrates the complete sparing of the superior longitudinal
512
512 resolution.The widened sulci are easily visible and indicate fasciculus on this side of the brain. Areas of comparison between
severe atrophy.The lesion in Broca’s area occupies only the posterior the left and right hemispheres are boxed in white. Crosshairs on
portion of the pars opercularis and is boxed in white.The anterior slice S (LH) indicate the positioning of the coronal and axial slices.
parts of Broca’s area (pars triangularis and the anterior half of the
pars opercularis) are still intact.Crosshairs on the 3D reconstruc-
tion indicate the orientation of the coronal and axial slices.

Roberts, 1959). These deficits tend to resolve quickly but

nevertheless suggest that the area supports some end-stage
articulatory function, probably assisting in control over the
muscles of articulation. Functional neuroimaging studies also
consistently observe activation in Broca’s area with tasks that
involve articulation (Cabeza and Nyberg, 2000).
This current study causes us to re-evaluate the nomen-
clature used to describe the location of Broca’s area on the
inferior frontal gyrus. Inspection of these historic brains
indicates that the lesion viewed by Broca and considered by
him to be critical for speech is not precisely the same
region now termed Broca’s area. Leborgne’s lesion did
involve the posterior inferior frontal gyrus, but was even
more extensive in the region anterior to it. Lelong’s lesion
occupied only the posterior third of what is now called
Broca’s area. It is important to note that Broca referred to
the ‘posterior inferior frontal gyrus’ as the general area
important for speech. The lesions of these two cases do
indeed fall within this broad region. It is since his time that
Broca’s area was redefined and currently differs from Fig. 8 Views of the damage to the superior longitudinal fasciculus
Broca’s original description. Future lesion and functional in the brain of Lelong. Slices through the affected portions of this
tract can be seen in sagittal slice S, in coronal slice (C) and in axial
neuroimaging studies should be advised that the current slice (A). A sagittal slice through the 3D reconstruction of the left
designation of Broca’s area is not what Broca originally hemisphere is also shown, with crosshairs indicating the orienta-
intended and that defining regions of interest that specify tion of the coronal and axial slices.
 Page 10 of 10 Brain (2007) N. F. Dronkers et al.

Broca’s area may not be including the inferior frontal area Brodmann K. Vergleichende Lokalisationlehre der Grosshirnrinde in ihren
Broca originally described. Prinzipien dargestellt auf Grund des Zellenbaues. Leipzig: Johann
Ambrosius Barth; 1909.
Paul Broca began the study of the localization of function Brown-Sequard CE. Aphasia as an effect of brain-disease. J Med Sci 1877;
that has matured into the fields of neuropsychology, 63: 209–25.
speech–language pathology, neurolinguistics and cognitive Cabanis EA, Iba-Zizen MT, Abelanet R, Monod-Broca P, Signoret JL.
neuroscience. We now know that language and cognition ‘‘Tan-Tan’’ the first Paul Broca’s patient with ‘‘Aphemia’’ (1861): CT
are far more complicated than once thought and involve (1979), & MRI (1994) of the brain. In: Picard L, Salamon G, editors. 4th
refresher course of the ESNR: language and the aphasias. Nancy:
large networks of brain regions and connecting fibres.
European Society of Neuroradiology; 1994. p. 9–22.
During Broca’s time, though some may have proposed Cabanis EA, Iba-Zizen MT, Nguyen TH, Bellinger L, Stievenart JL,
networks to support cognition, scientists could not have Yoshida M, et al. Les voies de la vision, de l’IRM anatomique a la
imagined the complexity of such systems, nor could Broca physiologie (IRM(f)) et IRM en tenseur de diffusion (IRMTD) ou
have supposed that his discovery would launch such tractographie. Bulletin de l’Academie Nationale de Médecine 2004; 188:
remarkable explorations into the functions of the human 1153–72.
Cabeza R, Nyberg L. Imaging cognition II: an empirical review of 275 PET
brain. Fortunately, Broca had great foresight in preserving and fMRI studies. J Cogn Neurosci 2000; 12: 1–47.
these historic brains and in some ways, Leborgne and Castaigne P, Lhermitte F, Signoret JL, Abelanet R. Description et étude
Lelong can speak to us more eloquently now than they scannographique du cerveau de Leborgne. Rev Neurol (Paris) 1980; 136:
could over 140 years ago. 563–83.

Downloaded from http://brain.oxfordjournals.org/ by guest on June 7, 2013

Clarke E, O’Malley CD. The human brain and spinal cord. Berkeley:
University of California Press; 1968.
Acknowledgements Crosson B. Subcortical functions in language: a working model. Brain
We are particularly grateful to the late Professor Philippe Lang 1985; 25: 257–92.
Monod-Broca, the great-grandson of Paul Broca, and his wife Dronkers NF. Neuroanatomical correlates of production deficits in
aphasia. Telerounds. Tucson, Arizona: National Center for Neurogenic
and daughter for their time and interest in this work.
Communication Disorders; 1993.
Professor Monod-Broca’s recent book is the most compre- Dronkers NF. A new brain region for coordinating speech articulation.
hensive biography concerning the contributions of Paul Nature 1996; 384: 159–61.
Broca, and his support of new work related to his great- Dronkers NF, Ogar J. Aphasia. In: Aminoff MJ, Daroff RB, editors.
grandfather’s discoveries was inspiring to us. We also thank Encyclopedia of the neurological sciences. San Diego, CA: Academic
Professor Paul Prudhomme de St Maur, Director of the Press; 2003.
Gall FJ, Spurtzheim G. Recherches sur le système nerveux en général et sur
Musee Dupuytren, for his generous assistance in obtaining celui de cerveau en particulier. Paris: F. Schoell; 1809.
and transporting the brains and in supporting this project. Geschwind N. Language and the brain. Sci Am 1972; 226: 76–83.
Appreciation is also extended to Benjamin Russell, Michael Goodglass H. Understanding aphasia. San Diego: Academic Press; 1993.
DeRiesthal, Jelena Jovanovic, David Payet, Bruno Delamain, Joynt RJ, Benton AL. The memoir of Marc Dax on aphasia. Neurology
Juliana Baldo, David Wilkins, Brenda Redfern, Carl Ludy, 1964; 14: 851–4.
Marie P. Revision de la question de l’aphasie: la troisieme circonvolution
Darren Husted, Helen Ettlinger, Art Shimamura and the UC
frontale gauche ne joue aucun role special dans la fonction du langage.
Berkeley Institute for Personality and Social Research for their Semaine Medicale 1906; 26: 241–7.
contributions. This work was supported by the Department Mohr JP. Broca’s area and Broca’s aphasia. In: Whitaker H and
of Veterans Affairs Medical Research, the National Institute Whitaker H, editors. Studies in Neurolinguistics. New York Academic
of Neurological Disorders and Stroke (NS040813), the Press; 1976: 1: 201–233.
National Institute on Deafness and other Communication Mohr JP, Pessin MS, Finkelstein S, Funkenstein HH, Duncan GW,
Davis KR. Broca aphasia: pathologic and clinical. Neurology 1978; 28:
Disorders (DC00216) and the Centre Hospitalier National 311–24.
d’Ophtalmologie des Quinze-Vingts, Paris. Monod-Broca P. Paul Broca, un géant du XIXe siècle. Collection hommes
de science, homme de vérité. Paris: Vulbert; 2005.
References Moutier F. L’aphasie de Broca. Paris: Steinheil; 1908.
Benson DF, Ardila A. Aphasia: a clinical perspective. New York: Oxford Naeser MA, Hayward RW. Lesion localization in aphasia with cranial
University Press; 1996. computed tomography and the Boston Diagnostic Aphasia Exam.
Bramwell B. A remarkable case of aphasia. Brain 1898; 21: 343–73. Neurology 1978; 28: 545–551.
Broca P. Nouvelle observation d’aphémie produite par une lésion de la Naeser MA, Palumbo CL, Helm-Estabrooks N, Stiassny-Eder D,
troisième circonvolution frontale. Bulletins de la Société d’anatomie Albert ML. Severe nonfluency in aphasia. Role of the medial subcallosal
(Paris), 2e serie 1861a; 6: 398–407. fasciculus and other white matter pathways in recovery of spontaneous
Broca P. Perte de la parole: ramollissement chronique et destruction speech. Brain 1989; 112: 1–38.
partielle du lobe anterieur gauche du cerveau. Bulletins de la Societe Penfield W, Roberts L. Speech and brain mechanisms. Princeton, NJ:
d’anthropologie, 1re serie 1861b; 2: 235–8. Princeton University Press; 1959.
Broca P. Remarques sur le siège de la faculté du langage articulé, suivies Radanovic M, Scaff M. Speech and language disturbances due to
d’une observation d’aphémie (perte de la parole). Bulletins de la Société subcortical lesions. Brain Lang 2003; 84: 337–52.
d’anatomie (Paris), 2e serie 1861c; 6: 330–57. Schiller F. Paul Broca: explorer of the brain. Oxford: Oxford University
Broca P. Sur les mots aphemie, aphasie et aphrasie; Lettre a M. Press; 1992.
le Professeur Trousseau. Gazette des hopitaux 1864; 23. Signoret J, Castaigne P, Lehrmitte F, Abelanet R, Lavorel P. Rediscovery of
Broca P. Sur le siege de la faculte du langage articule. Bulletin de la Societe Leborgne’s brain: anatomical description with CT scan. Brain Lang
d’anthropologie 1865; 6: 337–93. 1984; 22: 303–19.

View publication stats