The brain under the knife: serial sectioning
and the development of late
University of Oulu, Finland, Department of History, PO Box 1000, 90014 Oulun yliopisto, Finland
Received 7 May 2005; received in revised form 17 November 2005
Major changes took place during the last quarter of the nineteenth century in the ways that the
brain tissue was maintained, manipulated and studied, and, consequently, in the ways that its struc-
ture, functions and pathologies were seen and represented in neurological literature. The paper
exempliﬁes these changes by comparing German neuroanatomy in the 1860s and early 1870s (repre-
sented above all by Theodor Meynert) with the turn-of-the-century view of the brain (represented by
Constantin von Monakow and others). It argues for the crucial importance of a method—serial sec-
tioning—to the emergence of the new view of the brain. Serial sectioning in turn owes its existence to
the new techniques in staining and sectioning that were introduced in the 1870s and 1880s. In par-
ticular, the paper highlights the role of a cutting device, the microtome, in enabling serial sectioning
and in thereby contributing to the emergence of a new view of the brain.
2006 Elsevier Ltd. All rights reserved.
Keywords: Late nineteenth-century neuroanatomy; Late nineteenth-century neuropathology; Serial sectioning;
History of the microtome
In the early 1870s, the Viennese neuroanatomist Theodor Meynert (1833–1892) was, as
a postgraduate student of his put it, ‘the most distinguished person in Europe as regards
1369-8486/$ - see front matter 2006 Elsevier Ltd. All rights reserved.
E-mail address: Heini.Hakosalo@oulu.ﬁ (H. Hakosalo).
Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
Studies in History
and Philosophy of
psychiatry and brain anatomy’.
Meynert’s research skills were praised, and his descrip-
tions of cerebral pathways and the structure of the cortex were considered authoritative.
By mid-1880s, however, Meynert was being heavily criticised and, by the 1890s, he was
mainly used as a negative reference point. The leitmotiv of the criticism was the more or
less outspoken charge that Meynert’s claims lack factual support and his observations can-
not be replicated by less imaginative observers. Thus August Forel (1848–1931) remarked
that Meynert’s ‘imagination made leaps which exceeded mine tens times over’,
mund Freud (1956–1939) disparaged Meynert’s ‘far-reaching speculations on anatomical
Even Friedrich Jolly’s (1844–1904) obituary contained ambiguous references
to Meynert’s ‘lively artistic imagination’, his ‘naı
¨ve artistic outlook’ and his ‘peculiar sys-
tem’ so rich in ‘startling notions’.
This violent counter-reaction against the Meynertian view of the brain has been put
down to his controversial personality, to his somewhat uneasy position between univer-
sity and asylum psychiatry and/or to his convoluted manner of expression.
It might be
added that the uniﬁcation of Germany in 1871 decreased the relative weight of the
Vienna Medical School and its anatomo-pathological emphasis in comparison to Berlin
with its more heavily physiological research orientation. This paper oﬀers a further expla-
nation for Meynert’s brusque fall from grace: it focuses on the major changes that took
place in dissecting and preparation techniques in the 1870s and 1880s, particularly on the
emergence of serial sectioning, and argues that this silent revolution needs to be taken
into account if one wants to understand, ﬁrst, the demise of the Meynertian view of
the brain and, second, the wider neurological reorientation that took place around the
turn of the century. This reorientation is characterised above all by severe erosion of
I shall start by looking at Meynert’s view of the brain and his anatomical research
techniques, move on to describe the emergence of serial sectioning in the mid-1870s (I
shall focus on a relatively little studied instrument, the microtome), and lastly discuss
the theoretical and disciplinary consequences of the introduction of the new method.
There are two strands of historical research that are relevant to this enterprise: studies
which discuss nineteenth-century serial sectioning and microtomy in contexts
other than brain studies
and studies which focus on neurological ideas—on Meynert’s
Letter from Auguste Forel to Pauline Forel, Vienna, 4 December 1871. Published in Forel (1968), p. 84. All
translations are my own unless otherwise stated. See also Forel in Stockert-Meynert (1930), p. 257.
Forel (1935), pp. 64, 81.
Freud (1953), p. 46.
Jolly (1892), pp. iv, vi–vii. See also Phleps (1904–1905), p. 233.
Lesky (1976), p. 339.
This reorientation is often called the holistic turn. I avoid this term because I think that it is still an open
question whether there was a wholesale ‘holistic turn’ (scepticism regarding localisationist notions does not
automatically imply holism). I agree with Harwood (1998), pp. 489, 496–497 that we do not yet know how far and
how deep holism in fact went in late nineteenth and early twentieth-century neurosciences.
Brian Bracegirdle charts the development of nineteenth-century microtechnique, including the microtome, in
painstaking technical detail in his History of microtechnique (1986); Nick Hopwood illustrates the importance of
microtomy for nineteenth and early twentieth-century embryology (1999, 2000, 2002, 2004) and Soraya de
Chadarevian (1993) discusses the role of the microtome in nineteenth-century botany.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 173
or on the questions of localism and/or holism
—in late nineteenth and early
twentieth-century Europe. This paper complements them by discussing the role of serial
sectioning in neurosciences and, more speciﬁcally, by assigning it a causal role in the
anti-Meynertian and anti-localisationist reactions. Instead of relating neurological ideas
to the broader social and political context, I shall ask what happened in the restricted
but fundamentally important context of the neurological laboratory.
2. The brain according to Meynert
I would hesitate to say that Theodor Meynert is representative of his era in the sense of
being typical. However, his work does illustrate the opportunities that were open to an
ambitious and skilful student of the brain in the late 1860s and early 1870s. Although
Meynert remained active until the late 1880s, the main tenets of both his method and
his conception of the brain were securely in place by the mid-1870s.
Meynert spent his professional life in Vienna, where he held a chair of psychiatry and
directed a psychiatric clinic. Although psychiatry had been his stepping stone to a chair
(neurology as an independent academic specialty did not exist yet), Meynert was a neuro-
anatomist ﬁrst and a clinical psychiatrist second, and he consistently interpreted mental
illness in a neuroanatomical framework. Characteristically, his ﬁrst academic appointment
(in 1866) was as Prosektor at the Vienna Lunatic Asylum; the Prosektor performed autop-
sies and collected anatomical specimens, leaving clinical work to others.
which was located at the Vienna General Hospital, specialised in acute neurological and
psychiatric cases, while patients with chronic mental illnesses were more likely to be trea-
ted at the State Psychiatric Hospital. Meynert also had a neuroanatomical laboratory at
People who worked with him testify to his lack of interest in clinical med-
and it is also clear from his published works that therapy was not a major concern
for him. For instance, the ﬁrst part of Meynert’s textbook on psychiatry (1884) concen-
Meynert is discussed in Whitaker & Etlinger (1993), Seitelberg (1997), and Hagner (1997), pp. 268–272.
Whitaker, Etlinger and Seitelberg act as Meynert’s advocates, and their praise can perhaps be seen as a counter
reaction to the predominantly negative turn-of-the-century estimates of Meynert. For Seitelberg, Meynert is
nothing less than ‘the founder of scientiﬁc brain research’ (Seitelberg 1997, p. 264). Whitaker and Etlinger
attribute a set of groundbreaking ideas to him (even raising him to the rank of nobility in the process) (Whitaker
& Etlinger, 1993, p. 567). In Hagner’s more contextual account, Meynert’s view of the brain epitomises the
emergence of the modern ‘nerve man’, an organism governed by his cortex.
The question of holism (of diﬀusionism) and localisationism has been one of the organising dichotomies in the
history of modern neurology at least since Walther Riese’s History of neurology (1959), and it has raised the
interest of many historians of medicine and neuroscience since: Robert Young’s Mind, brain and adaptation (1970)
is a standard work on neurological localisation. Susan Leigh Star relates localisationism to the organization of
research and clinical work in her Regions of the mind (1989), but has little to say about neuroanatomical methods
and nothing at all about serial sectioning. Anne Harrington’s Medicine, mind and the double brain (1987) studies
hemispheric dominance in late nineteenth and early twentieth-century brain research. In her later works
(Harrington, 1990, 1996) she has concentrated on the so-called holistic turn. Steven Jacyna’s Lost words (2000) is
about aphasia studies, a topic that is intimately related to the question of cerebral localisation. None of these
works discusses serial sectioning, let alone gives it a causal role in the turn-of-the-century neurological
Miciotto (1978), p. 183.
¨ller (1991), pp. 144–145, 124; Lesky (1976), pp. 159, 224–235; Grois (1965), p. 116.
Schnitzler (1970), pp. 223–224; Forel (1935), p. 65.
174 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
trates on brain anatomy. The second part, which was supposed to cover clinical psychia-
try, never materialised.
Meynert’s main research interests were in mapping cerebral pathways and sensory cen-
tres, in describing cortical structure and in ﬁnding anatomical correlations for mental
functions and disturbances.
Both contemporary and latter-day commentators have regarded the distinction between
projection and association ﬁbres as one of Meynert’s chief anatomical contributions.
Meynert ﬁrst introduced the classiﬁcation in 1865, in a chapter that he wrote for Maximilian
Leidesdorf’s (1816–1889) psychiatric textbook.
The distinction is the cornerstone of his
conception of the brain, and he returns to it in all his major publications. Projection ﬁbres
are the radiating ﬁbres which ascend from the spinal cord to the hemispheres and descend
from the cerebral cortex through the medulla and white substance to the spinal cord. Meyn-
ert believed that the entire body is projected onto the cortex through these ﬁbres. Association
ﬁbres are the ‘archiform ﬁbres’ that both begin and end in the cortex and unite diﬀerent parts
of the same and contralateral cortex to each other.
Some of them are short, reaching only as
far as the next convolution, while others are long. Together, they make sure that every part of
the cortex is both anatomically and functionally related to the other parts.
The two concepts took ﬁrm root in nineteenth-century neurological discourse and sur-
vived the criticism that was later aimed at Meynert’s neuroanatomical views, although
Meynert’s more speciﬁc description of the projection system did not escape criticism.
Meynert also charted functionally specialised cerebral pathways and centres. In 1865, he
discussed the anatomy of the olfactory, the visual and the auditory system.
In tracing the
auditory pathway in 1866, he relied on pathological anatomy: he reported a case of ‘distur-
bance of psychical speech’ (aphasia), and concluded from the pathological lesions that he
found in the patient’s brain that the auditory pathway terminates in the temporal lobe wall
of the Sylvian fossa. He labelled this area ‘the sound ﬁeld’ (Klangfeld) and regarded it as ‘a
central organ of speech’.
In 1870, Meynert followed the optic pathway from the retina to
the calcarine cortex of the occipital lobe, thus conﬁrming Bartolomeo Panizza’s (1785–
1867) 1855 observations.
In 1874, Eduard Hitzig credited him for ‘following up the termini
of the sensory pathways, that is, the nerves of sight, hearing and taste, to the occipital and
It was this part of Meynert’s work that would receive the heaviest blows
during the following decades. Thus Pierre Marie (1853–1940) called Meynert’s conception of
the course of the auditory pathway ‘absolutely mistaken’,
David Ferrier (1843–1928) con-
tested Meynert’s view that the thalamus has a motor function,
Sigmund Freud and Liweri
Darkschewitsch (1858–1925) opposed Meynert’s interpretation of the relationship between
Jolly (1892), p. v; Whitaker & Etlinger (1993), p. 568.
Meynert (1865), p. 51. I shall refer to this chapter as Meynert (1865), although it is published under
See, for example, Meynert (1868), p. 83; (1884), pp. 39–40, 138; (1892), pp. 28, 86.
Meynert (1884), p. 36.
Gehirn (1888), p. 690.
Meynert (1865), pp. 58–63.
Seitelberg (1997), p. 267; Marshall & Magoun (1998), p. 98.
Hitzig (1874), p. 21.
Marie (1906a), p. 241.
Marshall & Magoun (1998), p. 213.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 175
the posterior bundle and the cerebellum,
and Forel contradicted Meynert’s interpretations
of the pathway connections of the tegmental region, proposing to replace Meynert’s ‘fantas-
tic constructions’ with ‘objective facts’.
Meynert’s most extensive study on the histology of the cerebral cortex is Der Bau der
Gross-Hirnrinde und seine o
¨rtlichen Verschiedenheiten, published as a series of articles in
1867–1868. In this work, he estimates the number of cortical nerve bodies (Nervenko
to at least 612 112 000 and divides the grey surface of the cortex into two areas, one which
contains ﬁve cortical layers and another, more extensive area, which consists of eight layers
Meynert describes the structure of each cortical layer in detail. The pioneering nat-
ure of Meynert’s work is reﬂected in the fact that the vocabulary he uses is non-technical and
not yet ﬁrmly established. He notes the presence of diﬀerently formed cellular elements and
ﬁbres, comments on their density and arrangement and compares them with the visual
appearance of the other layers. There are spindle-shaped, pyramidal, as well as regular
and irregular cells. Some have a ‘pustule-like’ nucleus while others are particularly rich in
The study also contains references to function. Thus Meynert says
that the sound ﬁeld displays some morphological unity: it is populated by spindle-formed
cells, which seem to relate functionally to the ‘higher senses’, that is, sight and hearing.
Besides, this cortical area is particularly well developed in humans, which is only what one
would expect from a central organ of language.
He notes that the hippocampal gyrus,
the cortical area which he thinks is most directly concerned with epileptic convulsions, an
‘eminently motor disturbance’, contains pyramidal cells only, while the most well established
of all sensory areas, the olfactory lobe, is inhabited by granular cells.
Meynert’s view of cor-
tical structures was corrected, complimented and criticised during the following decades. For
instance, Franz Nissl (1860–1919) pointed out that Meynert’s description of the microscopic
structure of the optical region was inadequate,
Forel criticised Meynert’s description of the
cell structure of the tegmental region
and Freud rejected Meynert’s view that there are
‘empty’, that is, functionless patches between the various cortical centres.
Although Meynert regarded the cortex not only as histologically but also as function-
and although he is sometimes regarded as the arch-localisationist,
never posited a speciﬁc ‘centre for thought’ or ‘centre for association’, but rather placed
higher mental functions in the whole forebrain or the whole cortex.
Consistent with this
Darkschewitsch & Freud (1886), p. 123.
Forel (1877); (1935), pp. 65, 80–81.
Meynert (1868), pp. 80, 111.
Meynert (1867), pp. 205–206.
Meynert (1868), pp. 109, 112. On the localisation of the ‘sound ﬁeld’, see also Meynert (1867), p. 93; (1884),
Meynert (1868), p. 111.
Nissl (1898), p. 150.
Forel (1877); (1935), pp. 65, 80–81.
Freud (1953), p. 54; Meynert (1865), p. 53.
See, for example, Meynert (1884), p. 126–129.
Solms & Saling (1990), p. 125.
For example, Meynert (1884), pp. 137, 155. With Vorderhirn, or forebrain, Meynert usually seems to
understand much the same as we do, that is, the cerebrum plus some other important structures (he includes the
caudate and lenticular nuclei but excludes the thalamus, which he places in the ‘inter-brain’ rather than the
forebrain) (ibid., pp. 2, 70, 149). At other times, however, he equates Vorderhirn either explicitly (e.g. ibid., p. III)
or implicitly (e.g. ibid., p. 137) with the cerebral cortex.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 177
view, Meynert deﬁnes mental diseases as diseases of the forebrain. Knowledge of these dis-
eases ‘should be obtained, as all sound clinical knowledge is acquired, by study of the
structure, the function, and the nutrition of each organ’. Although clinical observation
comes ﬁrst in practical terms, epistemological primacy in both neurology and psychiatry
belongs to pathological anatomy and histology.
In the 1867–1868 work Meynert
refers—in very general terms—to histological (and even molecular) changes that accom-
pany psychological trouble and are connected to ‘hyperaemic ﬂux’.
His later writings
accentuate the vasomotor aspect. Mania and melancholia, for instance, are ‘nutritional
disturbances of the forebrain’, and the same goes for functional neuroses.
For Meynert, cerebral anatomy and histology are the key to understanding not only
pathological phenomena but also normal psychological functions. He explains that the
cortex supports the two key elements of psychological life, the ﬂeeting sensory impressions
¨cken) and the more durable mnemic images (Erinnerungsbilder), which derive
from past sensory impressions.
Sensory impressions are produced by the sensory organs
and the cortex, while memory—or the ability to retain the modiﬁcation brought about by
the innervation which accompanies all sensory impressions—is a function of all nerve cells.
Only part of these mnemic images ever become conscious. According to Meynert, ‘the
association-bundles may be compared to a connecting thread, which enables one image
to lift the other, as it were, over the threshold of consciousness’.
are ﬁrmer than others, and ‘individuality implies the sum of ﬁrmest associations’.
Meynert calls the primary ego coincides with the strongest and most archaic sensations,
the ones that concern one’s own body and its basic needs. The secondary ego com-
prises associations related to the world outside, including things such as one’s ‘intimate
relationships, possessions, rank-related social skills (consolidated by countless repetitions),
well-exercised arts, science, the deep-reaching goals in life, convictions, fatherland, and
In a highly optimistic manner, Meynert believed that not only individual men-
tal acts but also memories, associations, personality and many features of human culture
can be explained on the basis of a few neuroanatomical building blocks. By the turn of the
century, many people considered his manner of freely superimposing psychological and
anatomical terms and his eﬀort to explain mental functions and disturbances in anatom-
ical (or vasomotor) terms illegitimate. For instance, Nissl (1898) and Weygant (1900)
thought that it was a serious error from Meynert to confound the association tracts (an
anatomical structure) with associative processes (psychological phenomena). Weygant
added that the Meynertian way of seeing things had been nearly fatal for the development
Meynert wanted to build a neurological system, which was based on neuroanatomy but
also extended to the ﬁelds of neuropathology, neurophysiology and psychology. In doing
this, he extrapolated physiological and psychological phenomena from neuroanatomical
Meynert (1885), p. vi.
Meynert (1867), pp. 215–216.
Meynert (1890), pp. 5, 184, 186.
Meynert (1868), p. 79.
Meynert (1884), p. 142.
Ibid., p. 155; (1892), p. 24.
Meynert (1884), p. 162; (1892), p. 35.
Nissl (1898), p. 151; Weygant (1900), p. 658.
178 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
ﬁndings. His eﬀorts as a system-builder were hampered by his avowed reluctance to write
the relative crudity of his anatomical methods and the fact that he did not do
physiological or psychological research himself. With all its shortcomings, Meynert’s sys-
tem nevertheless provided a highly useful reference point—ﬁrst positive and then predom-
inantly negative—for the young science of the brain. What Randall Collins says about
philosophers can be applied to Meynert, too: ‘Great intellectual work is that which creates
a large space on which followers can work. This implies that the imperfection of major
doctrines are the source of their appeal’.
3. Cutting up the brain
In the mid-1870s, the two standard methods in the anatomical study of the human
brain were deﬁbering and freehand sectioning. By the mid-1880s, the two had been largely
surpassed by serial sectioning, by the study of secondary degenerations
and by the mye-
Given that the latter two methods relied heavily on it, serial sectioning
can be regarded as the centrepiece of this methodological upsurge.
Meynert’s method of choice was Abfaserung, also called Hirnfaserung or Zerfaserung.
The American translator of Meynert’s Psychiatrie, Sacks, renders Abfaserung as ‘the cleav-
age-method’. The term ‘deﬁbrillation’ has also been used,
but the term is probably best
translated as deﬁbering. The method had been used in neuroanatomy for a long time to
chart the course of ﬁbre bundles.
This was usually done by gently scraping away the tis-
sue around the ﬁbres, for instance with a needle or a pair of tweezers. The success of def-
ibering greatly depended on the way that the brain had been prepared beforehand. The
best hardening method was one that made nervous elements hard but left connective tissue
soft and easily removable. In Meynert’s day, hardening was usually done by immersing the
brain in or injecting it with a solution containing alcohol and nitric acid, hydrochloric acid
or chromic acid. The specimen was either conserved in alcohol or treated with oil of cloves,
mounted in balsam and placed in a small bell-jar.
A standard neuroanatomical
Meynert (1884), p. IV.
Collins (1998), p. 32.
The method of secondary degeneration is based on the fact that individual parts of the nervous system are
trophically connected to each other: if one of them is destroyed—either experimentally or by disease—the ﬁbre
bundles which are connected to it will cease to function and, in due course, also display microscopic changes in
structure. By observing these changes, one can ascertain the course of a pathway. While many people, including
¨rck (1810–1868) and Meynert, had occasionally studied spontaneous pathological degenerations, it
was Bernhard von Gudden who pioneered the experimental study of secondary degenerations. For contemporary
descriptions of the method, see Gehirn (1888), p. 688; Obersteiner (1892), p. 132.
The myelogenetic method is attributed to Paul Flechsig (1847–1929), who, in 1876, discovered that the ﬁbres
of the central nervous system do not myelinate simultaneously, but rather in a series of successive steps during
foetal and infant life. He realised that it was possible to distinguish the already myelinated pathways and regions
from the unmyelinated ones if one used stains that dye the myelin sheath—such as the Weigert stain. Flechsig
stated that the ﬁbres which myelinate simultaneously are ‘equally important’, and distinguished between three
main ﬁelds (primordial ﬁelds, intermediate ﬁelds and terminal ﬁelds) and thirty-six regions in the brain.
Johan Hultkranz’s book Gehirnpra
¨paration mittels Zerfaserung (1929) was translated into English as Brain
preparations by means of deﬁbrillation (1935).
A similar method had been used for instance by Raymond Viussens (1641–1716) to study nerve ﬁbre tracts, by
Franz Josef Gall (1758–1828) to reveal deep-lying structures of the brain, and by Bartholomeo Panizza (1785–
1867) to trace the course of the optic pathway in 1855.
Obersteiner (1888), pp. 5, 4–5.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 179
textbook warns that ‘all methods of deﬁbering ::: can very easily give rise to
As the idiosyncratic term ‘cleavage-method’ indicates, there seems to be something
beside the standard, run-of-the-mill deﬁbering involved in Meynert’s Abfaserung. There
is indeed something artistic—original and intuitive—in the way that Meynert describes
both the procedure and its results. He ‘dissects away’ and ‘tears asunder’ various struc-
tures of the brain in order to reveal the course of a suspected ﬁbre tract or to view the
‘mountains and valleys of the cortical waves’ (Wellenberg and Wellenthal).
He also writes
that ‘the structure of the cerebral cortex, like that of a crystal, can be best studied from its
The cleavage-surface represented in (Fig. 1) is clearly neither a
straightforward cross-section nor an isolated ﬁbre bundle. The results of this method
are not easy to replicate: Meynert’s Abfaserung requires practice and insight, but even
two well practised neuroanatomists are unlikely to tear the brain asunder in exactly the
same way to produce two identical specimens. In 1884, Meynert regrets that the younger
generation does not have the patience to learn to master the laborious cleavage-tech-
and indeed, a medical handbook from 1888 regards the method obsolete, except
for purposes of demonstration.
Meynert also sectioned: he either cut the brain into two or more chunks to get cross-
sections of the selected planes or else cut thin, even transparently thin slices from planes
which optimally displayed the internal structures under study. Gross cross-sections were
studied macroscopically, while transparent sections could be studied either microscopi-
cally or macroscopically. Meynert sectioned freehand, usually with a knife and sometimes,
when handling smaller specimens, with a hand microtome.
He was proﬁcient with the
razor; he was one of the few who is said to have been able to section through an entire
dog brain. Sections have a prominent place in Meynert’s publications. For instance, he
produced ten ‘absolutely veracious sketches of the natural transverse sections’ from vari-
ous parts of the olfactory, auditory and optic pathways for Leidesdorf’s psychiatric text-
and transparent sections also constitute the material for Meynert’s 1867–1868
study on the structure of the cortex.
Nevertheless, Meynert makes it clear that he regards
deﬁbering as the more demanding, sophisticated and important of the two methods. As he
saw it, sectioning produces a fragmentary view of the brain, a ‘mosaic’ as it were. Deﬁber-
ing, on the other hand, ‘not only [provides] the point of departure from which to under-
stand this mosaic, but also enables us to extend our knowledge of the structure of the brain
beyond the level where brain sections cease to render us further information’.
Exact replication of results was not easy with the sectioning method, either, because
freehand sectioning was actually very diﬃcult. Thus Forel said that ‘A successful slice
through a whole dog brain ought to be considered a ﬁrst-class trick in itself’ and confessed
Meynert (1884), p. 39.
Ibid., p. 35; This quote is taken from Sacks’s translation, Meynert (1885), p. 37.
Meynert (1884), p. v.
Gehirn (1888), p. 688.
Stockert-Meynert (1930), p. 38; Schopfhagen in ibid., p. 48.
Meynert (1865), p. 45, footnote.
Meynert (1867), p. 198. They were prepared with Rudolf Berlin’s (1833–1897) method, that is, ﬁxed with
potassium dichromate, stained with carmine and cleared with turpentine.
Meynert (1884), p. V.
180 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
that he never managed to produce a whole section from anything bigger than a mole or a
Monakow also spoke about the diﬃculty of making technically adequate
sections with a free hand,
and Jolly regarded Meynert as a ‘virtuoso’ because he could
Bracegirdle aﬃrms that ‘Much practice is required to make sections
of adequate quality (thinness and evenness) freehand ::: Parts of sections may be of ade-
quate quality, but to obtain complete sections is impossible’. He adds that ‘in the earlier
part of the nineteenth century :::many eminent workers in histology prided themselves on
their skill with a freehand razor, decrying such instruments as section-cutting machines’.
In this sense, Meynert was a member of the old school.
Notwithstanding the stalwarts of freehand sectioning, serial sectioning with the help of
a microtome largely replaced freehand sectioning and deﬁbering from the mid-1870s
onwards. In serial sectioning, the brain, or parts of it, is cut into an unbroken series of very
thin slices, which can then be stained and inspected either macroscopically or microscop-
ically. This technique was ﬁrst made possible by the perfection of the light microscope, by
the development of eﬀective chemical reagents for hardening, ﬁxing and staining micro-
scopic specimens and by the introduction of the brain microtome. The 1870s saw impor-
tant advances in all these fronts,
but since the ﬁrst two developments have been well
and since they are not speciﬁc to serial sectioning, I shall concentrate on the
development of the microtome.
The microtome is a device that is used to cut very thin slices of various substances for
microscopic work. The purpose of the microtome is to give greater control to the cutting
and to minimize the mechanical damage that is necessarily involved in the procedure.
While a simple hand microtome is little more than a metal tube that holds the embedded
specimen and whose edges support the knife or the razor, the most sophisticated nine-
teenth-century microtomes were large, complex and expensive machines, which not only
produced extremely thin, even-surfaced slices of a wide variety of materials, but also
did so very rapidly or even automatically. In the 1870s, most anatomists and histologists
still sectioned like Meynert, that is, with a free hand and a razor or a knife (the most
widely used model being the Valentin knife). The routine use of the microtome—together
with standard methods for ﬁxing and staining—started in zoology, in embryology and in
the various ﬁelds of human anatomy and histology by 1880. By the end of the century,
microtomes were commercially available and some models were in series production.
Benedikt Stilling (1810–1879) is usually credited for introducing serial sectioning to neu-
roanatomy. In 1842, he sliced a frozen spinal cord into a continuous series of sections
Forel (1935), p. 65; Forel in Stockert-Meynert (1930), p. 258.
Monakow (1970), pp. 108–109.
Jolly (1892), pp. iii–iv.
Bracegirdle (1986), p. 117.
The visible light microscope had gone through major improvements at mid-century. Its theory was clariﬁed by
Ernst Abbe in 1873. By the 1880s, it was technically more or less completed and had assumed a central role in
laboratory training in life sciences and medicine (Golinski, 1998, pp. 76–77). New techniques for hardening, ﬁxing
and staining nervous tissue were eagerly looked for and adopted in the 1870s. See, for example, Finger (2000),
Breidbach (1993) and Dierig (1993).
On the development of the light microscope, see Hartley (1993). On the history of tissue preparation, see
Monakow (1970), p. 152; Hopwood (1999), p. 465, (2000), pp. 37–38, (2002), p. 41, (2004), p. 186.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 181
and was able to see, even with a low-power microscope and with no staining, the radiating
bundles of nerves and the central tracts.
Brain tissue made special demands on the microtome. Because the brain, as a leading
nineteenth-century neuroanatomist put it, was ‘extremely delicate, soft and labile’,
needed very careful preparation. Sectioning through the entire human brain was particu-
larly hard, because the size of the organ made it diﬃcult to embed and too large for ordin-
ary microtomes to hold and cut. These problems were ﬁrst overcome in Bernhard von
Gudden’s (1824–1886) laboratory in Munich in 1875. Gudden, a psychiatrist and neurol-
ogist, had been using a Welker microtome for sectioning smaller pieces, but he found it
inadequate when he wanted to section entire brains.
To accomplish this, Gudden, his
assistant August Forel and his instrument-maker Katsch designed and constructed a
new microtome, subsequently known as ‘the Gudden microtome’. The microtome con-
sisted of a piston within a vertical metal cylinder. The embedded brain was placed on
the piston; there were three knobs on the piston to keep the specimen from moving. On
the bottom of the piston, there was a large micrometer screw, which allowed the person
working the apparatus to move the specimen up micrometer by micrometer with little
eﬀort. The cylinder of Katsch’s ﬁrst prototype was only 16 cm wide, which made it too
narrow for cutting sagittal sections of the whole human brain. The knife was moved by
hand and supported by the edges of the cylinder. In addition, there was a rubber hose
for wetting the cylinder in order to make the piston and the embedded specimen move
After producing a host of rumpled and torn sections, the Gudden team
noticed that they could diminish friction by cutting under water, and fastened the cylinder
to the bottom of a shallow basin.
The freshly cut sections ﬂoated freely in the basin until
they were caught on plates or on paper. Gudden and the others also experimented with
diﬀerent knives in order to minimize wrinkling and slanting (Fig. 2).
There was more to serial sectioning that just cutting. The brain had to be ﬁxed and hard-
ened before cutting, and stained, dehydrated and mounted afterwards. In Gudden’s labora-
tory, the brain was ﬁrst immersed in alcohol and then injected with potassium dichromate to
ﬁx and harden it. Gudden warned that a prolonged alcohol-bath would change the way that
the tissue stained.
The brain was then embedded in a mixture that contained 15 parts ste-
arine, 12 parts fat and 1 part wax (Gudden’s recipe does not specify the nature of the fat
and the wax). When every nook, ﬁssure and cavity of the brain had been covered with this
mixture, a little opening was made at the base of the third ventricle and the ventricle was ﬁlled
with the mixture in order to keep some of the ﬁner structures adjacent to it from collapsing
during cutting. Just before cutting, some of the embedding material was removed from the
Obersteiner (1888), p. 5; Naderi, Tu
¨re, & Pait (2004), p. 4; Bracegirdle (1986), p. 135. Bracegirdle has
reservations about attributing the introduction of serial sectioning to neuroanatomy to Stilling (Bracegirdle, 1986,
Obersteiner (1888), p. 3.
Gudden (1875), p. 229.
Ibid., pp. 230–231.
Ibid., p. 231; Forel (1935), p. 74. In his autobiography, Forel gives himself the key role in the construction of the
Gudden microtome. For instance, he says that he came up with the idea of cutting under water. Gudden on the other
hand mentions Forel only in passing and leaves the reader with the impression that he and Katsch were the
protagonists of the story.
Gudden (1875) p. 231; Forel (1935), p. 74.
Gudden (1875), p. 232. See also Obersteiner (1888), p. 6.
182 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
plane about to be cut.
Even so, the knife had to be wiped clean after each cut, and it also
required frequent sharpening.
The cut sections were caught on plates under water, kept
in pure water for a couple of hours and then stained with ammonia carmine. Next, they were
washed with water containing some acetic acid. After another twelve hours in water the
Fig. 2. The brain microtome that Katsch constructed for Bernhard von Gudden (from Gudden, 1875). The knife
is not seen in the illustration.
Gudden (1875), p. 233; Forel (1935), p. 74.
Obersteiner (1888), p. 8.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 183
specimens were transferred to glass plates, identiﬁed with rubber-coated labels, and
immersed in an airtight container with nitreous alcohol (in order to dehydrate them). The
specimens were then carefully brushed with oil of cloves until they were transparent (in order
to clear the alcohol and to penetrate the tissue with a medium of high refractive index), and
ﬁnally mounted in linseed oil varnish and covered with glasses. They were ready for use when
the varnish on the edges of the slides had dried.
Because commercially available slides were
too small for large specimens, brain anatomists often had to have their glasses cut to order.
Following this procedure, Gudden and his assistants produced the ﬁrst ever continuous series
of sections of an entire human brain. Both Gudden and Forel took great pride in this achieve-
ment, although Forel attributed the feat to himself and Gudden to himself and his two assis-
Next, they sectioned a human hippocampus, an ape brain (into 810 sections) and
some rabbit brains.
As can be gathered from Gudden’s description, serial sectioning was a labour-intensive
method. Gudden and Forel do not specify the time it took them to produce their series,
but other contemporary sources throw light on this question. Monakow had a Dutch vis-
itor to his neuroanatomical laboratory prepare a ‘complete series of sections through the
brain stem of a pathological brain’. The industrious Dr Walsem ‘worked without break
from morning to night, and the series was ready in four weeks’.
The adoption of the tech-
nique also called for some changes in the neurological laboratory. Gudden tells how to
make the laboratory better suited for serial sectioning: First, one should construct a small
worktop where the slides can be kept in a horizontal position when they are inspected mac-
roscopically, preferably against a dark sub-surface, with a strong side-light. Second, the
tables that are usually employed in microscopy would have to be replaced with bigger
ones. Third, one needs a lot of storage room in order to keep the hundreds and thousands
of slides, some of them very big, in order and readily available. A laboratory that is ideally
suited for serial sectioning thus has plenty of space, purpose-built tables and cabins, as
well as assistants, students and technicians.
The method soon became known beyond the conﬁnes of the Munich laboratory. The
adoption of the method was of course facilitated by the fact that sectioning as such was a
familiar, much used technique, although it had not been successfully applied to entire human
brains before. Gudden published a paper on the apparatus and its use in the Archiv fu
chiatrie und Nervenkrankheiten, the leading German journal on neuroanatomy. Gudden’s paper
also served as an advertisement for Katsch’s instrument and his workshop, and it no doubt
brought Katsch new orders. There must have been a demand for the apparatus, because Katsch
made a slightly modiﬁed Gudden microtome commercially available in 1882.
themselves travelled far and wide. Forel says that requests came from all over Europe and that
he soon found himself producing series of sections from morning till night.
In addition, anat-
omists came to Gudden to learn the technique in person, much as they had gone to Meynert in
the late 1860s to learn the deﬁbering technique. News of the novel method also travelled by
Gudden (1875), p. 233. See also Obersteiner (1888), pp. 10–11.
See Bracegirdle (1986), p. 112, on standard slide sizes.
Gudden (1875), p. 231; Forel (1935), p. 74.
Gudden (1875), p. 232.
Monakow (1970), pp. 204–205.
Bracegirdle (1986), p. 251.
Forel (1935), p. 74.
184 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
means of various publications that contained written descriptions of the method and, perhaps
even more eﬃciently, representations of sections.
What were the advantages of the new technique? Most obviously, it made the inner
structures of the brain visible in a new way. Heinrich Obersteiner (1847–1922) wrote that
‘signiﬁcant progress in the understanding of the inner structure of the brain only became
possible when we learned to prepare transparent cross-sections’.
As we shall shortly
see, it brought unprecedented precision to Faseranatomie, to the histological study of
the brain and also to pathological anatomy of the brain. A medical handbook from
1888 states that the method has become irreplaceable in brain anatomy, but it also men-
tions a drawback: while it is easy enough to follow a (stained and/or degenerated) ﬁbre
bundle if it is transversal to the plane that is being sectioned, other ﬁbre bundles fall into
small pieces when they are sectioned, which makes it hard to reconstruct them.
three-dimensional reconstruction was the main diﬃculty with serial sectioning. Recon-
struction was mostly done in the head and therefore subject to error. Obersteiner was
acutely aware of this problem: ‘The plastic reconstruction of the images that arise from
a number of sections can be diﬃcult; and yet the pictures which the cross-sections pro-
vide us with can further our anatomical knowledge only when we are able to gain a cor-
poral view of the object’.
Obersteiner gave some practical advice to aid the mental
reconstruction of anatomical structures. For instance, he advised the student of the brain
ﬁrst to inspect sections macroscopically with a stereoscopic loop (which retains a degree
of three-dimensionality) and only then study the sections microscopically.
He also told
the reader of the ﬁfth chapter of his book—the chapter which oﬀers a topographical
overview of the brain and relies heavily on microscopic sections—to keep a well hard-
ened brain stem and a fresh, cross-sectioned brain at hand, so as ‘never to lose sight
of the plastic-topographic totality’.
These diﬃculties of reconstruction are reﬂected in both two and three-dimensional
modes of representation. Serial sectioning did not essentially change neuroanatomical
modes of representation, although the number of drawings and later micrographs of sec-
tions in neurological publications of course increased. A drawing of a section looks the
same regardless of whether the section is part of a series of thousands or one of its kind.
In this sense, representation of serial sections can be compared to the representation of
another late nineteenth-century innovation, the ﬁlm: ﬁve minutes of ﬁlm is made of thou-
sands of individual pictures, but no more than a fraction of them can possibly be repro-
duced in traditional print media. In late nineteenth-century neuroanatomy, the surplus
information that was produced by a continuous series of sections had to be conveyed by
writing rather than by pictures, and anatomical descriptions found in late nineteenth-cen-
tury neurological journals are indeed often extremely long and detailed. Diagrams were a
way around this diﬃculty, where naturalism and three-dimensionality were sacriﬁced for
the sake of accuracy and comprehensiveness. Diagrams were particularly useful in provid-
ing information about the course of cerebral pathways and centres (Fig. 3).
Obersteiner (1888), p. 5.
Gehirn (1888), p. 688.
Obersteiner (1888), p. 5.
Ibid., pp. 1–2.
Ibid., pp. 208–209.
See Jacyna (2000), pp. 106–114, on schematic representation of language centres.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 185
Obersteiner (1888) pioneered the use of schematic and also paired representations. The
book contains several images where one half of the section is depicted naturalistically and
the other half, the mirror image of the latter, is rendered schematically (Fig. 4).
Fig. 3. A diagram representing the course of cerebral patways (from Obersteiner, 1888).
See Lynch (1990), pp. 160–166, on sequential transformation in photo-diagram pairs.
186 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
As for three-dimensional representation, there was no standard method in nineteenth-
century neuroanatomy for reconstructing three-dimensional images from a series of two-
dimensional specimens without losing much essential information on the ﬁner structures
of the brain. Modelling therefore never assumed the importance in neuroanatomy that
it had in contemporary embryology.
Although three-dimensional models of the brain
were available, mainly for didactic purposes, leading neuroanatomical authorities such
as Obersteiner and Wilhelm His (1831–1904) considered even the best of them
4. Serial sectioning in Faseranatomie
In less than a generation, serial sectioning changed the face of Meynert’s two main
ﬁelds of interest—the anatomy of cerebral pathways (Faseranatomi) and the morpholog-
ical study of the cortex—and it also left its mark on pathological anatomy.
The study of cerebral pathways was the ﬁrst and most obvious beneﬁciary of the new
opportunities oﬀered by serial sectioning. Meynert was the foremost authority in this ﬁeld,
and it is therefore no wonder that his view of cerebral pathways came under criticism with
the appearance of the ﬁrst series of cerebral sections. Forel says that immediately after pre-
paring the ﬁrst series of sections of the human brain he ‘managed to clarify various inner
structures which had so far been only unclearly and indistinctively depicted. The Meyner-
tian schemes fared badly indeed’.
Another early example of the potential of serial sec-
tioning to recast Meynert’s anatomical descriptions is the way that Emmanuel Mendel
(1839–1907) discussed the anatomy of the superior cerebellar peduncle at the Medico-Psy-
chological society of Berlin in January 1878. One of the issues was whether the auditory
pathway was related to the superior cerebellar peduncle or not. Mendel proposed to throw
light on the question with the help of a new method: he had used ‘the Gudden apparatus’
to produce several continuous series of frontal, horizontal and sagittal microscopic sec-
tions of the brain of monkeys and men. He demonstrated some of the sections at the meet-
ing, where they were viewed macroscopically, with the help of electric light. Mendel’s
conclusions—most importantly, that the acoustic pathway was related to the superior cer-
ebellar peduncle on the level of the dentate nucleus—were critical of Meynert’s views.
Meynert was not present in the meeting, but Carl Wernicke (1848–1904), his most loyal
disciple, hastened to contest Mendel’s conclusions. Wernicke, sensing where the real
enemy lay, attacked Mendel’s method: he maintained that serial sectioning was by deﬁni-
tion unable to produce conclusive information on the anatomy of the superior cerebellar
peduncle. According to him, the issue could only be decided on if it were possible to follow
the ﬁbre bundles in question directly, that is, with Meynert’s deﬁbering method.
Nick Hopwood has shown that late nineteenth-century embryologists came to rely heavily on plastic
reconstruction not only for purposes of demonstration but also for purposes of research (e.g. Wilhelm His’s
mechanical theory of embryogenesis). Serial sectioning played an important role in embryological modelling
(Hopwood, 1999, pp. 471–473).
His says about the best and the most expensive model, the Aeby model, that ‘it seems quite clear and
transparent when you have it in front of you, but it leaves you in the lurch as soon as you turn away your eyes’
(quoted in Obersteiner, 1888, p. V).
Forel (1935), p. 74.
Wernicke in Mendel (1878), p. 403.
188 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
simply urged Wernicke to see the slides for himself. Indeed, one of the practical advantages
of mounted sections was their relatively easy demonstrability, which in this case gave Men-
del an upper hand in the polemical situation.
The Russian-born Swiss neurologist Constantin von Monakow challenged and success-
fully reverted many basic features of Meynert’s view of the brain. He was also one of the
ﬁrst to put the Gudden microtome into concentrated use, and the major ﬁndings of the
ﬁrst part of his career would not have been possible without the instrument.
Monakow visited Gudden’s laboratory in 1875. The professor described his experimen-
tal studies with secondary degeneration, and showed Monakow a series of recently made
cerebral sections, leaving his younger colleague ‘stiﬀ with admiration’.
went to work at the St. Pirmingsberg asylum in 1878, he was delighted to discover that the
institution possessed ‘a brand new, unused Gudden microtome’. The instrument gave him
‘a friendly wink’ and provided him with the ﬁrst real opportunity to train himself in brain
Monakow sat by the instrument day after day, and, as a way of practice, made
the ﬁrst continuous series of frontal sections of the human brain stem.
In 1884, Mona-
kow read a neuroanatomical paper at the Berlin Physiological Society. Although the lis-
teners were interested, Monakow gained the impression that most of them ‘lacked real
understanding for ﬁner brain anatomical facts’. Perhaps this was only to be expected,
Monakow adds, given that ‘only very few physiologists and anatomists had set their eyes
upon microtome sections and series of such sections of the brain’.
In other words,
acquaintance with the microtome and serial sectioning initiated Monakow into brain anat-
omy and gave him a distinct feeling of superiority when faced with older colleagues. It also
made him into one of the most formidable critics of the Meynertian brain.
Equipped with the methods of serial sectioning and secondary degeneration, Monakow
tackled the question of cerebral pathways and centres. In 1870, Gudden had demonstrated
that the destruction of speciﬁc areas of the cortex caused atrophy in certain thalamic
nuclei. Monakow (1892) explored the implications of this ﬁnding. The paper starts with
a report on a series of animal experiments and moves on to discuss three pathological
cases. Two of the three patients had suﬀered from bilateral hemianopsia.
In all three
cases, post-mortem study had revealed secondary degenerations not only in the occipital
lobe and the optic pathway, but also in parts of the thalamus.
Monakow had prepared
continuous series of sections of the relevant parts of the brain, usually the brain stem and
the occipital lobes, and stained them with carmine.
He makes two methodological
points: ﬁrst, one must carefully distinguish between primary and secondary degenerations
(he provides criteria for recognising the latter) and, second, one must employ serial sec-
tioning. According to him, truly reliable observations about cortical pathways and centres
can only be obtained by serial sectioning. Any anatomical or pathological study that is not
supported by a series of sections carries less weight from the outset.
Cf. Latour (1990), pp. 35–36.
Monakow (1970), pp. 128–129.
Ibid., p. 152.
Ibid., p. 153.
Ibid., p. 188.
Hemianopsia is blindness in one half of the visual ﬁeld.
Monakow (1892), pp. 613, 635, 640, 643.
Ibid., pp. 620, 613–614, 618, 637, 650.
Ibid., pp. 231, 241, 243, 245, 247, 249.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 189
Comparing the analysis of the pathological ﬁndings with the results of the animal
experiments, he came to the following conclusions. First, both the experimental and the
pathological evidence conﬁrm that the integrity of the optic pathway depends on the integ-
rity of the optic cortex.
Second, the results suggest that the visual cortex is more exten-
sive and less clearly circumscribed than commonly thought. It does not comprise the
ﬁssura calcarina alone, as Munk would have it, but also extends to the banks of the ﬁs-
Third, the visual cortex and the optic pathway are not the only parts of the brain
that are involved in vision. In both higher mammals and in man, the ﬁssura calcarina is
related to what Monakow calls primary optic centres and locates in the lateral geniculate
body, certain caudal parts of the pulvinar and the superior colliculus. He noted that the
role of these centres had so far been ignored or undervalued.
Monakow stated that
the relationship between the primary optic centres and the visual cortex was a rule rather
than an exception in the organization of the brain, an instance of ‘a law’ according to
which ‘every segment of the grey substance of the thalamus (excluding periaqueductal gray
and its immediate environment) matches a quite clearly circumscribed cortical zone’, and
that these two structures are mutually interdependent.
Monakow used his results to oppose strict localisationism. He trusted that studies such
as his were likely to decentralise the prevalent view of the brain because they ‘point
towards the assumption that even quite simple nervous acts, such as simple sensory per-
ceptions, call for close simultaneous co-operation of several diﬀerent groups of ganglionic
cells and ﬁbres, perhaps in quite scattered and diverse parts of the brain’.
Monakow made this point even more forcefully: given that even the relatively uncompli-
cated optical perceptions are based on ‘physiologically diﬀerent, albeit closely interlocked
individual acts’, it would be absurd to attempt to localise complex processes of thought.
He regarded this decentralisation, ﬁrst achieved in the case of vision with the help of serial
sectioning, as ‘a cornerstone of anatomic localisation’ and ‘the main gain that the doctrine
of localisation has gleaned from anatomy in the past few years’.
The picture of cortical
pathways and centres that Monakow produced with the help of serial sectioning was not
only more detailed than Meynert’s conception, it was also much less cortico-centric and
less clearly localisationist.
5. Serial sectioning in the histological study of the cortex
At the turn of the century, the cellular structure and arrangement of the cortex became
the subject of an ambitious new research programme, cytoarchitectonics. The pioneers of
the programme were Alfred W. Campbell (1868–1937) in Britain and Korbinian Brod-
mann (1868–1918) and, somewhat later, Oskar Vogt (1870–1959) and his wife Ce
(1875–1962) in Germany.
A look at Brodmann’s publications shows that cytoarchitectonics relied heavily on
serial sectioning. According to Brodmann, ‘any precise histological localisation on the
Ibid., pp. 626, 628.
Ibid., pp. 634, 234.
Ibid., pp. 159, 234, 251, 299–230.
Ibid., p. 260.
Monakow (1904), p. 116.
Monakow (1902), pp. 588, 593.
190 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
cerebrum presupposes two technical conditions: ﬁrst, the preparation of unbroken series
of sections, second, the production of comprehensive synoptic sections (U
of the whole organ or at least of the portion of the brain under study and its immediate
According to Brodmann, ‘properly histological’ sections should be no
more than 10–12 micrometers thick, but the broader the section, the more diﬃcult it is
to achieve this.
To make sections which are both ‘synoptic’ and thin, Brodmann ﬁrst
used a macrotome to cut the organ in neat, parallel chunks and then a speciﬁcally
designed, exceptionally stable double-sledge microtome (Doppelschlittenmikrotom) to pro-
duce ﬁne histological sections. With these instruments, he was able to make an ‘unbroken
series of perfectly even microtome sections 80 ·70 millimetres wide and 10 or even 5
micrometres thick’ of human frontal and occipital lobes.
Brodmann studied the cortex of humans and several animals from such sections and
published his results in 1908. He divided the human cortex into forty-seven areas, num-
bered them and described their minute cellular structure and arrangement. Although
Brodmann wanted primarily to establish the morphological identity of these areas, he
was optimistic about the possibility of correlating the morphological and the functional.
Given that ‘quite a few anatomically circumscribed regions are within physiologically dis-
tinct zones or almost coincide with them’, it is only logical to postulate that ‘any region
with speciﬁc histological structure must have a speciﬁc physiological integrity’.
bell’s research agenda was much the same as Brodmann’s, although their results diﬀered.
Campbell, too, used comparative material to describe the cellular and laminar structure of
the cortex, and he, too, wanted eventually to ‘establish a correlation between physiological
function and histological structure’.
The picture that Campbell and Brodmann drew of
the simian and human cortex was immensely more detailed that the picture that Meynert
had drawn in 1867–1868. It is obvious that this degree of detail would have been impos-
sible without serial sectioning.
6. Serial sectioning in neuropathology: the case of aphasia
A shift of emphasis is also apparent in the ﬁeld of pathological anatomy around the
turn of the century, and this shift is partly due to new research techniques such as serial
sectioning. The following examples deal with aphasia, the most thoroughly studied
brain-based pathology in the late nineteenth century.
The ‘classical’ theory of aphasia was based on the correlation between various clinical
manifestations on the one hand and the presence of localisable lesions on the other hand.
The two most commonly recognised variants of aphasia were motor aphasia, characterised
by ‘loss of articulate speech’ and sensory aphasia, characterised by diﬃculties in under-
standing spoken and/or written language plus paraphasia. The former had been referred
back to lesions in the third frontal convolution by Paul Broca (1824–1880) in 1861 and
the latter had been correlated with lesions in the ﬁrst left temporal convolution by Carl
Brodmann (1903–1904), p. 206.
Ibid., p. 210.
Brodmann (1908), pp. 244, 245.
Campbell (1904), p. 148.
Cf. Bracegirdle (1986), p. 27.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 191
Wernicke in 1874. In the great majority of cases, aphasic lesions were in the left hemi-
sphere. Wernicke’s view of the brain owed a great deal to his one-time teacher Meynert.
It was commonly believed that the area or areas damaged in aphasia normally supported
linguistic functions. Because of this physiological implication, aphasia had become the
most important anatomo-clinical touchstone for the theory of functional localisation, that
is, for the view that diﬀerent mental functions can be localised in clearly delimited areas of
the cerebrum (or, more speciﬁcally, the cortex). Thus, any research which seriously chal-
lenged the classical theory of aphasia also automatically cast doubts on the localisationist
view of the brain.
In 1906, F. Quensel, who worked under Paul Flechsig (1847–1929) at the time, published
an extensive article on aphasia. The paper discusses three cases. The clinical description of
the cases is relatively short but the post-mortem report can only be described as exhaustive.
Quensel had followed a similar procedure in all three cases: he had hardened one or both
hemispheres in chrome solution and sectioned them with the microtome. He had then cho-
sen every ﬁfth (in the ﬁrst and third case) or every seventh section for closer study, stained
these with the Weigert method
and inspected them microscopically. His sections were
approximately 50–70 micrometres thick and they added up to 400, 235 and 200 respec-
Quensel had numbered the sections, and used them to describe the exact location
and extent of the lesions. Quensel’s conclusions are critical of former schematic represen-
tations of aphasia (e.g. Wernicke’s and Henry Charleton Bastian’s [1837–1915] diagrams).
He states that while their schemes and diagrams may serve didactic purposes, they have lit-
tle to do with anatomical reality. They are based on theoretical considerations on the one
hand, and on ‘superﬁcial anatomo-clinical evidence’ on the other hand, while a scientiﬁc
conception of aphasia must be based on the study of actual cerebral connections by means
of serial sectioning.
Quensel’s article oﬀers no radical revision of the doctrine of aphasia,
but it does indicate that pathological anatomy had come to rely heavily on serial sectioning,
and is another instance where a student of the brain has come to doubt strictly localisation-
ist interpretations after adopting the method.
The classical conception of aphasia was radically revised in 1906, although not by
Quensel. In a series of three articles, the French neurologist Pierre Marie (1853–1940)
attacked the prevailing conception of aphasia, coming very close to calling ‘the classical
aphasia’ the great nineteenth-century neurological hoax. Marie wanted to redraw both
the clinical and the anatomical face of the classical aphasia. Marie’s main anatomo-path-
ological claims were the following: lesions in the third left frontal convolution are in no
way speciﬁc to motor aphasia;
aphasic lesions always involve the Wernicke area;
Carl Weigert (1845–1904) introduced the method that was to become known by his name in 1885, when he
applied haematoxylin (with potassium dichromate as a mordant) to the study of myelinated nerve ﬁbres. The
Weigert method colours normal myelin sheaths black (or blue in some variants), and the method became an
important tool in the study of myelination and the study of secondary degenerations, which were popular
methods in the 1880s and after. A much used modiﬁcation was the Pal-Weigert stain, ﬁrst described in 1887. This
technique heightened the contrast between myelinated ﬁbres and the tissue surrounding them.
Quensel (1906), pp. 37, 56, 177.
Ibid., p. 387.
Marie (1906a), p. 243; Marie (1906b), p. 500.
Marie (1906a), p. 244; Marie (1906b), p. 500; Marie in ‘3e discussion sur l’aphasia’ (1908), p. 1037. Marie’s
deﬁnition of the Wernicke area was more extensive than Wernicke’s own: Marie included the gyrus
supramarginalis, gyrus angularis and the roots of the two ﬁrst temporal convolutions to it.
192 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
aphasic lesions are seldom, if ever, restricted to the cortex;
anarthria, which is not to be
seen as a form of aphasia but which sometimes complicates true aphasia, leading to the so-
called motor aphasia, is caused by a lesion in the ‘lenticular zone’ in the corpus stria-
the so-called images of language as well as specialised centres of language (such
as the centre for acoustic images of language), which supposedly house these images,
are mere ﬁgures of speech and have no support in facts.
Marie also re-examined the
brains of the two patients—Leborgne and Lelong—on whom Broca had initially based
his theory of aphasia, in order to show that they had been badly observed and interpreted
and did not warrant Broca’s weighty conclusions.
Generally speaking, Marie thought
that the aphasiological discourse had all too often been guided by theoretical assumptions
rather than empirically observed facts.
Marie did not only criticise his former maı
ˆtre Broca but also his long-term rival Jules
who became the champion of classical aphasia in the ensuing
´rine answered Broca in writing in 1906 and debated the issue of aphasia with
him in a series of three meetings at the Neurological Society of Paris in June–July 1908.
´rine contradicted Marie point by point. De
´rine’s main anatomical and anatomo-
pathological claims were that Marie’s deﬁnition of the ‘lenticular zone’ was inconsistent,
vague and ridiculously extensive
and that aphasic lesions usually did involve the Broca
He was, however, willing to depart from the classical, localisationist aphasia the-
ory in one important respect: he conceded that, in motor aphasia, the lesions are typically
not restricted to the root of the third frontal convolution of the left hemisphere, but tend
to be more extensive and less clearly delimited.
It is therefore better to speak about an
extensive zone of language that comprises both the Broca and the Wernicke areas than
about two distinct centres of language.
It seems that De
´rine had been convinced of
the truth of this revisionary view above all by the 1900 thesis of his student Fernard Bern-
heim. Bernheim had studied a number of brains from patients who had suﬀered from
motor aphasia with the help of serial sectioning and the microscope and he had come
to the ﬁrm conclusion that even in clear cases of motor aphasia the lesions are seldom
if ever limited to the small area that Broca had regarded as the seat of the disease.
Marie’s revision has been much discussed in historical literature, but the fact that tech-
nical questions played a major role in the debate between Marie and De
´rine has been
overlooked. Both Marie and De
´rine were products of the Paris school of medicine
and swore by the anatomoclinical method,
but their attitude towards serial sectioning
Marie (1906a), pp. 244, 246; (1906b), p. 493.
Marie (1906a), pp. 243–244; (1906b), p. 500; Marie in ‘Discussion sur l’aphasie’ (1908), pp. 615, 628.
Marie (1906b), p. 494; Marie in ‘2e discussion sur l’aphasia’ (1908), p. 1004; Marie in ‘3e discussion sur
l’aphasia’ (1908), pp. 1032, 1034–1035, 1038, 1042.
Marie (1906c), pp. 565, 567.
Marie (1906a), pp. 241, 247.
Marie (1906b) concentrates on critisising Jules De
´rine. For instance, Marie calls De
between cortical and subcortical aphasia and his deﬁnition of pure word deafness ‘purely hypothetical’ (ibid., p.
´rine in ‘2e discussion sur l’aphasie’ (1908), pp. 975–976, 1012;A.De
´rine in ibid., p. 991.
´rine in ibid., pp. 1000–1001, 1008.
´rine (1906b), p. 453.
Ibid., p. 457.
Bernheim (1900), (1906), p. 529;J. De
´rine (1906b), p. 453.
Marie (1906a), (1906b), p. 493.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 193
diﬀered. Marie’s 1906 revision was based on macroscopic study of brains, and De
was not slow in recognising this as the weak point of Marie’s case. In 1906, De
branded Marie’s approach, that is, macroscopic inspection of individual transverse sec-
tions, as old-fashioned and inadequate. He proclaimed that the only method capable of
producing sound anatomopathological data on aphasia is the microscopic study of series
of continuous sections, that is, the method that he and his laboratory were associated with
In his response, Marie refused to admit that serial sectioning was imperative in neuro-
pathology (although he admitted that it was necessary in neuroanatomy), and added that
it did not take a microscopic examination to revoke Broca’s doctrine, which is ‘built solely
upon badly interpreted results of macroscopic examination’.
He dismissed serial micro-
sectioning—at least in the case of the so called motor aphasia—as ‘futile rites’ performed
‘around an erroneous doctrine’.
On the other hand, Marie hastened to add that his lab-
oratory, too, produced serial sections and subjected them to microscopic analysis.
in a medical meeting in the same year Marie and his student Franc¸ois Moutier (1881–1961)
even claimed that they had been the ﬁrst to cross-section aphasic brains that had not been
spoiled by a prolonged alcohol-bath, which explained why they had been able to see the
true nature of aphasia.
This was guaranteed to outrage De
´rine, who immediately
listed a host of studies that had relied on the study of either cross- or microscopic sections
and predated Marie’s and Moutier’s work. Besides, De
´rine added, ‘during the past ﬁf-
teen years, every anatomoclinical study on the nervous centres that has left my laboratory
has been based ::: on serial microsections’.
In the 1908 debate, De
´rine and his collaborators—his wife Augusta De
1927) his students Fernand Bernheim and Andre
´Thomas—continued to use serial section-
ing to undermine Marie’s claims. The De
´rines showered the society with sections (as well
as with photographs and drawings made from sections) to demonstrate that Marie had
misconstrued the relationship of the lenticular zone and the third frontal convolution.
They argued emphatically that the true extent of a cerebral lesion can only be judged from
continuous series of sections and dismissed all data which was based on mere macroscopic
surface examination or on individual sections.
´rine even suggested that the Society
form a committee to oversee the sectioning of Broca’s original two specimens.
ming up the proceedings, De
´rine’s said that the debate had proven beyond doubt what
he had been saying for years already, namely that ‘the method of serial microscopic sec-
tions is absolutely necessary for all study on cerebral localisation’.
Marie cleaved to
´rine (1906b), pp. 453, 454.
Marie (1906c), p. 571. In Broca’s day, there had been no reliable way to section the brain. He had settled for a
macroscopic surface inspection of the two brains and then deposited them at the Muse
´e Dupuytren in alcohol-
ﬁlled jars (Broca, 1861).
Marie (1906c), p. 571.
Marie & Moutier (1906), p. 744.
´rine (1906b), p. 455 n.1.
´rine in ‘2e discussion sur l’aphasia’ (1908), pp. 979, 992.
´rine in ibid., pp. 992–993, 1000–1001, 1008; A. De
´rine in ibid., pp. 991, 999; J. De
´rine in ‘3e
discussion sur l’aphasia’ (1908), p. 1027.
´rine in ‘2e discussion sur l’aphasia’ (1908), p. 1011.
´rine in ‘3e discussion sur l’aphasia’ (1908), p. 1047.
194 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
Fig. 5. Jules De
´rine was strongly associated with the technique of serial sectioning in France, as can be gathered
from this caricature (drawn by G. Villa and published in Chanteclair,105, June 1912). He is depicted in Charcot’s
chair (or tower), sectioning a spinal cord.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 195
his former revisionary claims about aphasia, but we no longer hear him denying the crucial
importance of serial sectioning and microscopic study, and he and his students did their
best to supply series of sections in support of his views.
Although it is diﬃcult to say
who won the debate,
´rine is right in considering serial sectioning as a winner.
Monakow, too, read an important paper on aphasia in 1906. Although he did not
accept Marie’s re-interpretation of the concept of aphasia, he welcomed Marie’s eﬀorts
to rethink the question.
One reason why he considered Marie’s speciﬁc claims less than
convincing is because he agreed with De
´rine that only ‘modern brain anatomical
research methods (serial sections)’ can supply truly new and reliable knowledge about
aphasia. Monakow himself had analysed approximately thirty cases of aphasia for the
paper, ﬁfteen of which were dissected and ten of which thoroughly sectioned. These cases,
together with an analysis of a set of negative cases reported by others, have convinced
Monakow that ‘the closer we analyse the clinical experiences of aphasia patients and
the deeper we penetrate in the focal anatomical changes in the cerebrum of such patients
(by means of serial sections), the more diﬃcult it is to construct ::: a satisfactory expla-
nation of the more intimate interrelation between the locality of the lesion and the aphasic
symptoms that have been observed intra vitam’.
The standard anatomical explanation—
that aphasia is caused by destruction of either a centre or a connective pathway between
centres—is far from suﬃcient.
What is needed is a new point of view and a new explan-
atory factor. The new point of view has to be functional rather than localisationist, and the
new explanatory factor is his notion of ‘diaschisis’. Diaschisis is a nervous shock, a partic-
ular type of action at a distance (Fernwirkung), which accounts for the transitory symp-
toms of aphasia.
It also proved useful in explaining the possibility of recovery in
These historical examples show that, by 1908, serial sectioning was considered a sine
qua non in cerebral pathology as well as in cerebral anatomy. Not even Pierre Marie,
who knew he could not match De
´rine in this respect, denied any longer its crucial impor-
tance. Monakow equated serial sectioning with modern neuroanatomical research meth-
ods; the method was instrumental in convincing him that a psychologically complex
phenomenon such as aphasia cannot be understood solely from the study of focal lesions.
´rine, a zealous advocate of serial sectioning, has often been cast as an old-fash-
ioned localisationist because he became the main opponent of Marie’s reinterpretation of
aphasia. However, De
´rine, too, was willing to break down the strict correlation between
motor aphasia and a clearly delimited lesion at the root of the third left frontal convolu-
tion and to renounce the theory of two distinct centres of language. Given that this cor-
relation was the cornerstone upon which the whole classical theory of aphasia had been
Moutier in ‘2e discussion sur l’aphasia’ (1908), p. 1019.
Neither Marie nor De
´rine gave any ground to the opponent during the debate. It is diﬃcult to say who the
majority of the Society sided with, because, according to the written record, they were mostly supported by their
own students and collaborators, Jules De
´rine by Augusta De
´rine, Fernand Bernheim and Andre
Pierre Marie by Franc¸ois Moutier and Alexandre Souques. Latter-day commentators have clearly preferred
Marie’s ‘modern’ views to De
´rine’s more traditional ones, but the fact is that Marie did not eﬀectively
overthrow the classical theory.
Monakow (1906), p. 1038.
Ibid., p. 1026.
Ibid., p. 1027.
Ibid.; see esp. p. 960.
196 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
´rine was in fact making a major concession to a more holistic interpretation of
aphasia and language.
7. Disciplinary consequences
New microtechniques aﬀected the contents of histological and neurological theories but
they also modiﬁed the disciplinary landscape. I shall lastly discuss some of these disciplin-
First, there were changes in the relationship of the various specialties concerned with
the study of the brain. Armed with the new microtechniques, microscopical anatomy lar-
gely supplanted macroscopical anatomy, while neurophysiologists increasingly relied on
anatomists to supply their experimental studies—experimental ablation and electrical
stimulation above all—with crucial anatomical support.
Second, serial sectioning added to the replicability of neuroanatomical ﬁndings and
thereby to their scientiﬁc credentials. While Meynert’s results could only be replicated
by another rare virtuoso, the advocates of new microtechniques liked to emphasise that
their methods allowed reliable results to be obtained quickly and eﬃciently, with little
or no prior training. Thus Gudden notes in the 1875 paper: ‘Cutting with a free hand must
be learned laboriously. Many people who have really tried have never been able to learn it.
On the contrary, sectioning with the instrument described above is so easy that most stu-
dents with no prior experience manage the ﬁrst section in the most perfect manner’.
Obersteiner agrees that ‘the production of [brain anatomical] sections, which often have
to be fairly large, previously required a skilled hand and much practice; currently this
problem is solved by the introduction of the microtome’.
Although the new microtech-
niques, too, required a great deal of tacit knowledge and manual skill, they were still nota-
bly easier to learn than Meynert’s deﬁbering and freehand sectioning. The fact that
Meynert was proud rather than embarrassed about the diﬃculty of his favourite method
and the poor replicability of his results seems to indicate a change not only in research
techniques but also in scientiﬁc ideals.
Third, compared with fresh brains, or with Meynert’s macrospecimens, microsections
functioned as very eﬃcient inscriptions in Bruno Latour’s (1990) sense. They were mobile:
once microtechniques became routinely employed, and especially after reagents and
instruments became commercially available, slides could be produced en masse and widely
distributed in space and time. They could be taken to meetings and conferences for dem-
onstration. They were immutable: unlike whole or cross-sectioned brains that are kept in
alcohol, balsam-mounted slides are practically permanent.
Their scale could be modi-
ﬁed, they could be superimposed upon each other for comparisons and they could be made
part of a written text. They also allowed for some degree of mathematization.
Walker (1957), p. 445.
Gudden (1875), p. 231; see also p. 232.
Obersteiner (1888), p. 7.
Fresh brains of course start to deteriorate immediately, but it was also diﬃcult to maintain Meynert’s crisp
cleavage-surfaces when the brain was kept in alcohol for a longer period of time. Specimens ‘preserved in the
commonest type of alcohol :::soon changed their outer forms into artefacts (Kunstprodukte)’ (Monakow, 1970,
pp. 108–109); see also (Bracegirdle, 1986, p. 91).
Latour (1990), pp. 44–46.
H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202 197
were no small advantages if we believe with Latour that Western science is essentially char-
acterised by ‘this simple drift from watching confusing three-dimensional objects, to
inspecting two-dimensional images which have been made less confusing’.
Lastly, the adoption of serial sectioning as a standard method increased the cost of
neuroanatomical and neuropathological study. The increased reliance of neuroanatomy
on chemical and instrumental high tech further consolidated the barrier separating med-
ical scientists from the rest of the medical profession. The more heavily neuroanatomical
research depended on costly machinery and tailor-made facilities, the less hope there was
for amateur scientists or medical men without secure academic base to establish them-
selves in the ﬁeld.
And the rising cost of research can also shape the contents of scientiﬁc discourse. Turn-
of-the-century aphasiology is a case in point. Since 1861, aphasiological discourse had
been conducted on predominantly anatomoclinical terms. That is, the discourse had
mostly been based on what were regarded as ‘complete’ anatomoclinical case reports, that
is, reports which included a detailed clinical description of the patient on the one hand and
a post-mortem report on the state of the diseased brain on the other. By the turn of the
century, the archive constituted by such reports was massive but just about manageable.
´rine introduced a new deﬁnition for a complete anatomoclinical case in the
1908 debate: he insisted that a case report is not complete unless the whole cerebral area
suspected of being damaged is cut into a continuous series of very thin sections and micro-
scopically examined. Since no one present at the meetings of the Neurological Society
explicitly opposed this deﬁnition and several members of the Society explicitly embraced
it, we can assume that it was commonly accepted by that time. In practical terms, the
introduction of the new criteria for a complete, not-to-be-dismissed case report meant that
the damaged part of the brain was ﬁxed and/or hardened, cut into tens, hundreds or thou-
sands of microsections all or some of which were stained, dehydrated, mounted, studied
with the microscope and usually stored for further reference. Some of the sections were
perhaps sent to other laboratories for re-examination, demonstrated in scientiﬁc meetings
or classes, or drawn or micrographed for the purposes of demonstration or publication.
The cost of presenting a strong anatomoclinical argument thereby rose immensely. Very
few practising physicians or even clinical professors could mobilize the human resources,
the skills, the apparatus and the room required for collecting complete anatomoclinical
cases. Those who could not were left with two possibilities: they could either renounce
hope of reaching the forefront of aphasia research or approach the question of aphasia
not from the traditional anatomoclinical point of view but from a radically diﬀerent angle.
This may well be one reason for the rising popularity of functional, psychological and lin-
guistic approaches in early twentieth-century aphasiology.
8. Concluding remarks
Serial sectioning was eﬀectively introduced to brain anatomy after the mid-1870s when
it ﬁrst became possible to section entire human brains. It rapidly changed the face of
Meynert’s three major ﬁelds of interest: anatomical study of cerebral pathways and cen-
Ibid., p. 39.
Cf. Chadarevian (1993), p. 562, and Hopwood (2000), p. 45.
198 H. Hakosalo / Stud. Hist. Phil. Biol. & Biomed. Sci. 37 (2006) 172–202
tres, histological study of the cortex and pathological anatomy of brain diseases. The
demise of Meynert’s conception of the brain is thus readily understandable. More gener-
ally, serial sectioning undermined the strictly localist view of the brain in at least three
ways. (1) It shook the belief in speciﬁc functional centres in the brain by emphasising
the complexity of pathways and the obscurity of the presumed functional centres. (2) It
made highly detailed and intensive study of the cortex possible, and this study failed to
provide direct and immediate support for localisation of function. (3) In many cases,
and aphasia is the most notable example of this, serial sectioning and microscopic exam-
ination of pathological brains underlined the protean nature of cerebral lesions. Serial sec-
tioning was also responsible for the turn-of-the-century neurological reorientation by a
more indirect way. By immensely raising the cost of presenting a plausible neuroanatomi-
cal or anatomy-based neuropathological argument, serial sectioning helped to eliminate
private researchers from the ﬁeld and forced some of the less well endowed academic
workers to look for novel approaches.
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