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Seeking the content in what is transient
Citation preview
ORI GIN AL PA PER
Seeking the constant in what is transient: Karl Ernstvon Baer’s vision of organic formation
Florence Vienne
Received: 8 July 2014 / Accepted: 16 November 2014 / Published online: 7 January 2015
� Springer International Publishing AG 2015
Abstract A well-established narrative in the history of science has it that the
years around 1800 saw the end of a purely descriptive, classificatory and static
natural history. The emergence of a temporal understanding of nature and the
new developmental-history approach, it is thought, permitted the formation of
modern biology. This paper questions that historical narrative by closely ana-
lysing the concepts of development, history and time set out in Karl Ernst von
Baer’s study of the mammalian egg (1827). I show that Baer’s research on
embryogenesis aimed not simply to explain temporal changes, but to inscribe the
formation of new individual organisms into a continuous, unending organic
process. I confront Baer’s views with other explanations of embryogenesis
arising in the 1820s and 1830s, especially those of Jean-Baptiste Dumas and
Jean-Louis Prevost and of Theodor Schwann. By highlighting divergences
between these scientists, especially as to their view of the role of gender dif-
ferences in reproduction, I argue that biology evolved not from a homogeneous
concept of developmental history but out of various, even opposing, views and
research programmes. Thus, the birth of biology did not imply the end of all
natural history’s thought models.
Keywords Developmental history � Embryology � Cell theory � Reproduction
In the late eighteenth century, a new configuration was to appear that would definitively
blur the old space of natural history for modern eyes.
Michel Foucault, The Order of Things, 162
F. Vienne (&)
Abteilung fur Geschichte der Naturwissenschaften mit Schwerpunkt Pharmaziegeschichte,
Beethovenstr. 55, 38106 Braunschweig, Germany
e-mail: [email protected]
123
HPLS (2015) 37(1):34–49
DOI 10.1007/s40656-014-0057-3
1 Introduction
To mark the opening of Konigsberg’s zoological museum in 1821, Karl Ernst von
Baer (1792–1876) gave a speech on the ‘‘current state of natural history.’’1 Baer
painted a rather disparaging portrait of natural history: the field, he complained,
remained too narrowly restricted to merely collecting, observing, describing and
classifying. What was there to be learnt by introducing more and more new species,
continually renaming them, and multiplying the names for a single object? The fact
that the number of mammals described in zoological works had trebled since
Linnaeus’s day was just one of the many examples Baer offered to send his
audience’s heads spinning and demonstrate the absurdity of a purely quantitative
proliferation of ‘‘catalogues’’ (von Baer 1821, p. 31). Baer’s words reflect the
enlargement of experience and knowledge in this period that has been described by
Wolf Lepenies (Lepenies 1976, p. 21). According to Lepenies, a constantly
accelerating quantitative expansion of knowledge in the last third of the eighteenth
century thrust natural history—with its descriptive, classificatory and spatial
orientation—into crisis, prompting the adoption of ways of thinking based on
history and, more narrowly, developmental history (Lepenies 1976, p. 45, 16). It
was this process of ‘‘temporalization’’ that, in Lepenies’s view, helped to overcome
the crisis, yet also brought about the ‘‘end of natural history’’ itself (Lepenies 1976,
p. 24). A decade earlier, in The Order of Things, Michel Foucault had portrayed the
transition from natural history to the history of nature in no less dramatic terms:
‘‘One day, towards the end of the eighteenth century, Cuvier was to topple the glass
jars of the Museum [of Natural History], smash them open and dissect all the forms
of animal visibility that the Classical age had preserved in them’’ (Foucault 1974,
pp. 137–138). In Foucault’s account, Cuvier’s gesture stands for ‘‘the end of
history’’ as it was understood by the likes of Linnaeus or Buffon, and the beginning
of an epoch in which a history of the living became thinkable (Foucault 1974,
p. 138, see also 263–279). This historiographical image, shaped by Foucault and
Lepenies, of the transition from a spatial and static ‘‘natural history’’ to a historical
‘‘biology’’ is what the present essay will critically address, starting from the work of
Baer.2 Only a few years after Baer formulated his criticism of natural history, he
published groundbreaking new studies in embryology and developmental history.
The case of Baer is therefore a fruitful one through which to examine the early
nineteenth-century emergence of new perspectives on organic nature, perspectives
based on history and specifically on developmental history.
1 Baer was born in Estonia. Between 1817 and 1834 he lived in Konigsberg, teaching and researching in
zoology, anatomy and anthropology. It was during this period that he carried out his renowned
embryological studies. In 1834, he moved to the Russian Academy of Sciences in St Petersburg, where he
remained until his retirement. On Baer’s life and work see, for example, von Baer (1866), Raikov (1968),
Breidbach in Baer (1999, pp. vii–ix), Breidbach and Ghiselin in Baer (2006, pp. v–xxvii), Schmuck
(2009, pp. 115–188) and Riha and Schmuck (2011). Here and throughout, all translations are my own
unless otherwise attributed.2 Francois Jacob also contributed to this picture, although in his account the transformation begins
earlier, in the eighteenth century (Jacob 1993, pp. 130–152).
Seeking the constant 35
123
The questions I raise here pick up on several studies that have explicitly or
implicitly queried the historiographical narrative of a ‘‘temporalization of nature’’
around 1800. Arno Seifert, for instance, has radically questioned the notion of an
atemporal premodern knowledge of nature. Seifert demonstrates that diachronic
histories of nature were in fact formulated in early modernity; he argues that
eighteenth-century naturalists and philosophers reverted to these earlier diachronic
histories but interpreted them in radically new ways. Rather than setting a
premodern, atemporal view of nature against a modern, temporal one, Seifert thus
distinguishes between various ways of envisioning the historicities and temporal-
ities of nature that arose throughout the early modern period (Seifert 1983; see also
Stockhorst 2006). Other scholars have worked on the case of Buffon to produce a
highly differentiated picture of the eighteenth-century concept of time and history
(Rheinberger 1990; Hoquet 2010). Hans-Jorg Rheinberger shows that Buffon, the
author of the Histoire naturelle (1749–1788), certainly did conceive of temporal
changes in organic nature, but did not regard the principle of change as inherent to
nature. For Buffon, Rheinberger concludes, the organic world was ‘‘subject to the
regime of time, but without history’’ (Rheinberger 1990, p. 221). Another far-
reaching critique of the ‘‘temporalization of nature’’ hypothesis is offered by Janina
Wellmann in her cultural history of embryology between 1760 and 1830. Wellmann
argues that the epistemological innovation around 1800 was not simply ‘‘time
bursting into the world of the organic;’’ in examples that include Baer’s description
of the chick embryo’s formation in the first volume of Entwicklungsgeschichte der
Thiere. Beobachtung und Reflexion (von Baer 1828/1999), she identifies the
emergence of a new structure and order of time that she describes as ‘‘rhythm’’
(Wellmann 2010, p. 373; see also Wellmann in this issue). The historians Dietrich
von Engelhardt and Peter Hanns Reill have contrasted the concept of history and
time that took shape around 1800 in the context of Naturphilosophie with the one
that prevailed during the Enlightenment (von Engelhardt 1979, 1988, 1990; Reill
2005). Reill argues that the Naturphilosophen differed from Enlightenment
naturalists in rejecting linear change and development. Instead, their visions of
organic nature were formed by ideas of circular processes and the return to origins.
Reill interprets the attempts of the Naturphilosophen ‘‘to eliminate or minimize the
temporal and epistemological uncertainties of a fully historicized world’’ as a
reaction against the French Revolution, which they experienced as a source of
uncertainty and chaos (Reill 2005, p. 220). He further shows that Naturphiloso-
phie’s call for the renewal of order and clarity transformed the epigenetic theories of
generation formulated by Enlightenment naturalists. Notions of ambiguity and
equality between the sexes gave way to those of gender polarity and hierarchy (see
Reill 2005, pp. 220–236; Lettow 2014; Vienne 2014).
In this historiographical context, it becomes important to ask after the concept of
developmental history underlying Baer’s early nineteenth-century research. With a
micro-analysis of his 1827 study of the mammalian egg, I begin by describing in
more detail Baer’s understanding of development, history, and time. Continuing
Reill’s line of thought, I also analyse the concept of gender difference upon which
Baer’s new vision of the process of organic formation rested. To what extent did
early nineteenth-century embryologists and physiologists continue the efforts of the
36 F. Vienne
123
Naturphilosophen to reinterpret generation in terms of a gender dichotomy? To
answer this question, I confront Baer’s study of the mammalian egg with other
explanations of embryogenesis that arose in the 1820s and 1830s, especially the
studies of fertilization carried out jointly by the French scholar Jean-Baptiste Dumas
(1800–1884) and the Swiss Jean-Louis Prevost (1790–1850) and the new
interpretation of generation by Theodor Schwann (1810–1884) based on cell
theory. As we will see, these researchers reached widely divergent conclusions on
the meaning of gender difference in generation, divergences that were associated
with different models of developmental history. My key hypothesis is that the 1820s
and 1830s saw the emergence of very different, and sometimes contradictory, forms
of thought and research in developmental history. In this sense, I contest Timothy
Lenoir’s assumption that biology issued from a unified ‘‘historical way of seeing’’
(Lenoir 1985, p. 100; see also Lenoir 1990) and a monolithic research programme
that he characterizes as ‘‘teleo-mechanist’’ (Lenoir 1982, p. 12). I would also dispute
the image presented by many historians of biology that a straight line of
development led from the developmental-history approaches of the late eighteenth
and early nineteenth century up to the formulation of cell theory (see Temkin 1950,
p. 237; Balan 1979, pp. 281–285; Duchesneau 1987, pp. 113–128; Lenoir 1982,
pp. 112–134; Jacyna 1990; Parnes 2000). This image largely accords with the
interpretation put forward by Foucault and Lepenies that, in the 1820s, ‘‘the
historical point of view’’ (Lepenies 1976, p. 16; my emphasis) came to prevail in
science once and for all, bringing with it the end of natural history.3 My own
conclusion is that the emergence of developmental-history perspectives in the first
third of the nineteenth century was far from putting an end to the whole range of the
static, classifying approaches and objects of natural history.
2 Change and continuity: Baer’s concept of developmental history
In his 1821 address on the ‘‘current state of natural history,’’ Baer contrasted his
sarcastic portrait of endlessly proliferating descriptions of nature with comparative
anatomy (especially the work of Cuvier) and Naturphilosophie. In Baer’s view,
these had both embarked upon a new way of looking at nature, the highest goal of
which was to investigate ‘‘the laws of organic formation’’ (von Baer 1821, p. 40).4
Baer intended the exploration of these ‘‘laws’’ to encompass not only processes of
becoming and passing away, but also—and especially—the eternal or unchanging
aspect of nature. This becomes very clear in another of his lectures, held in 1822:
‘‘The idea, or what is essential, what is living in animals is eternal, merely passing
through a series of different individuals in which it is always independently realized.
3 Lepenies borrows Foucault’s time frame, which locates the transition from natural history to biology in
the period between 1775 and 1825 (Foucault 1974, p. 221).4 In his memoirs, Baer claimed only to have ‘‘fully studied’’ Oken’s ‘‘natural philosophy,’’ having had
neither the time nor the interest for exhaustive consideration of the works of Schelling and other natural
philosophers (von Baer 1866, pp. 289–290, also 170–171). There are various interpretations of Baer’s
relationship with Naturphilosophie. See, among others, Riha and Schmuck (2011, pp. 230–233); Lenoir
(1982, pp. 85–86).
Seeking the constant 37
123
Thus, the law prevails in nature that the various forms of organic bodies are always
preserved whereas the individuals always perish’’ (Baer 1822, p. 63).5 Baer’s words
may be read as a programmatic statement prefiguring his famous embryological
studies of subsequent years. He argues that the research carried out hitherto—
whether based on preexistence theory or the theory of epigenesis—has been too
strongly oriented on the search for a ‘‘specific beginning, a sharp boundary between
being and nonbeing.’’ Yet, Baer continues, ‘‘nature shows us no such boundary
anywhere. At every moment she has something new, but, upon closer inspection,
what is new always reveals itself to be merely a transformation of something that
went before; nowhere does nature’s workshop show us a series of works that she has
begun from the very beginning. Ingenuity and imagination have therefore been
stretched to the limit in the attempt to detect this moment’’ (Baer 1822, p. 60). For
Baer, then, changing the trajectory of research on generation meant seeing the
formation of a new organism no longer simply as the beginning of an individual
existence, but as part of a larger, unending organic process. In this section I discuss
how, and with what epistemological implications, Baer put this programme into
practice with his embryological research.
In 1822, Baer offered his audience a developmental history of the bird’s egg that
was based neither on the assumption that the embryo was preformed nor that it took
shape suddenly out of unformed material. He picked up on a study of the formation
of the chick embryo published in 1817 by Christian Pander (1794–1865), a friend
from university days. Pander had ascribed the emergence of the embryo to ‘‘germ
layers’’ (Keimblatter) that formed in the first hours after incubation (Pander 1817),6
but Baer was not satisfied with this description of embryogenesis. He declared that
in future he would investigate earlier stages in the emergence of life: ‘‘Before the
embryo, the germ layer was in the egg. But the egg was laid by the hen. And a germ
was already present before fertilization. Fertilization is only the impetus for further
development of something that is already present; not a new beginning. Neither is
the egg a new beginning, since it develops in the hen’s ovary from other elements
also already present there. Thus, an absolute beginning is never demonstrated.
Always there is constant transformation; in their essence, the process of growth and
the process of increase are alike’’ (Baer 1822, p. 63). This objective of identifying
and describing the original elements of the process of organic formation in the ovum
found expression in Baer’s landmark study De ovi mammalium et hominis genesi of
1827.7
Baer’s treatise describes seven stages in the development of the canine ovum.
However, he begins his report not with the first of those stages, but with the
5 The only published form of this lecture is the summary by Boris Raikov (Raikov 1968), which is the
source of the following quotations.6 For detail on the context of Pander’s work and on his concept of the germ layer, see Wellmann (2010),
pp. 315–343; Schmitt (2005); Balan (1979), pp. 237–254.7 He wrote this text in Latin as an open letter to the St Petersburg Academy of Sciences, which had just
appointed him a corresponding member. The following quotations are based on the 1956 English
translation by Charles Donald O’Malley, ‘‘On the Genesis of the Ovum of Mammals and of Man’’ (von
Baer 1827/1956), or the 1927 German translation by Benno Ottow, Uber die Bildung des Eies der
Saugetiere und des Menschen (von Baer 1827/1927).
38 F. Vienne
123
condition of the ovum three weeks after copulation. At this point, he explains, all
the embryo’s organs can be recognized in rudimentary form. Baer traces the
development of the embryo backward from this point, but attaches a date to only
one other of the six preceding developmental stages: the ovum twelve days after
coitus (see von Baer 1827/1956, p. 151). No precise time statements are made
regarding the initial stages of development, which are the chief object of his study.
Baer’s attempt to find the original element in the process of organic formation
essentially consists in microscopic examination of the ova in their various organic
environments—in the uterus, in the tubes, and finally in the ovaries. Baer
ascertained their size, structure and component parts. Following this approach, he
‘‘opened’’ the ‘‘Graafian vesicles’’ in the ovary and ‘‘saw’’ there a structure that he
‘‘already recognized from the tubes’’ (von Baer 1827/1956, p. 132). From this
discovery, Baer concludes that alongside the Graafian vesicles (which he defines as
the ova of mammals) there is another ovum: ‘‘an ovum raised to the second power’’
(von Baer 1827/1956, p. 148), which—to avoid confusion—he also calls the
ovulum, or little egg (in the English translation ‘‘ovule,’’ in the German Eichen, von
Baer 1827/(1927), p. 15). However, in order to derive mammalian embryogenesis
from this organic entity, it would be necessary to establish whether the ovulum was
present before the Graafian vesicle. Baer admits that this question cannot be
answered by means of microscopic observation, but decides to accord priority to the
ovulum anyway (von Baer 1827/1956, p. 137). In Baer’s view, the embryo arises out
of this organized entity, and he therefore describes the ovulum as, ‘‘by reference to
the future foetus,’’ the ‘‘veritable ovum,’’ or ‘‘the foetal ovum in the maternal
ovum’’ (von Baer 1827/1956, p. 147–148).8 He triumphantly claims in the study’s
introduction to have been ‘‘fortunate enough to discover’’ the primordial beginning
of the ‘‘ovum of mammals and of man’’ (von Baer 1827/1956, p. 122). In the
developmental series portrayed in the upper part of his illustration (see Fig. 1), the
ovulum takes first place (von Baer 1827/1956, p. 150). It also features as a recurring
element at the later stages of development. Baer’s newly described organic entity
thus functions as an original—and constant—element in the process of organic
formation.
In the lower half of the plate are numerous depictions of the ovulum of various
mammals or the ‘‘vesicle of Purkinje’’ of other animals. In 1825, the Czech
physiologist Jan Evangelista Purkyne (1787–1869) had described this ‘‘vesicle’’ in
birds’ eggs as the ‘‘rudiments of the ova’’ (von Baer 1827/1956, 143). Baer
encountered Purkyne’s study in 1826, and was obviously impressed by it. As he
explained, it was a crucial concern of his own research to seek the ‘‘vesicles of
Purkinje’’ in the whole spectrum of animals, ultimately including mammals.9 His
demonstration that in mammals the process of organic formation also proceeded
from a previously organized ‘‘little body’’ (Korperchen) (von Baer 1866, p. 320; see
also von Baer 1866, pp. 310–320; von Baer 1827/1956, pp. 137, 142–148) broke
with the prevailing view that in mammals—unlike in other animals—the embryo
was formed from a female generative substance. For example, in his influential
8 In his memoirs, he also referred to the ‘‘primordial ovum’’ (von Baer 1866, p. 328).9 On Purkynes’s influence on Baer, see Knorre (1971) and Kruta (1971–1972).
Seeking the constant 39
123
reference work Die Physiologie als Erfahrungswissenschaft, physiologist Karl
Friedrich Burdach (1776–1847) wrote that what arose in the mammalian ovary was
not an ovum ‘‘but merely foetal substance that, after fertilization, provides only the
first rudiments of foetal membranes and foetus …. It is thus merely a fluid that,
without its own membrane, is contained in the cells of the ovary and stretches them
into vesicles’’ (Burdach 1828, pp. 74–75). Baer’s description of the ovulum as the
Fig. 1 Illustration from ‘‘On the Genesis of the Ovum of Mammals and of Man’’ (De ovi mammalium ethomnis genesis), in: Universitatsbibliothek Giessen, Nachlass Baer, Schriften, Bd. 22, Bl. 30r
40 F. Vienne
123
primordial beginning of the process of formation in mammals enabled him to
attribute embryogenesis in all animals to an analogous entity. In this respect, von
Baer (1827) treatise was far more than an attempt to reinterpret the process of
organic formation in mammals. It revealed correspondences in the formation
process among the different classes of animals, and led Baer to formulate a general
law of formation: ‘‘Every animal which springs from the coition of male and female
is developed from an ovum, and none from a simple, formative liquid’’ (von Baer
1827/1956, p. 149).
However, if Baer found similarities and a certain uniformity in the process of
organic formation among the different animals, this only applied to the origins and
early stages of that process (see von Baer 1828/1999, pp. 220–221). In the first
volume of his study Uber Entwicklungsgeschichte der Thiere. Beobachtung und
Reflexion, he postulated a general law of development according to which ‘‘the
homogeneous and shared gradually gives rise to the heterogeneous and specific’’
(von Baer 1828/1999, p. 153, see also 224). With this law, Baer was countering
recapitulation theory, the dominant explanation of embryonal development in the
1820s.10 He argued that embryos, at least in ‘‘higher’’ species, did not pass through
developmental stages during which they resembled ‘‘other,’’ lower animal forms,
but differentiated in a way specific to that particular animal (von Baer 1828/1999,
p. 224). Baer thus rejected the model of linear progression, what he called a ‘‘single-
line progression’’ (von Baer 1828/1999, p. 202). Rather than allocating species to
positions in a hierarchical scale, Baer divided the animal kingdom into four ‘‘types,’’
each based on a different scheme of formation.11 The embryogenesis of each animal
was laid down from the start by its ‘‘type’’ (von Baer 1828/1999, p. 220). The notion
of predetermined and purposeful development was fundamental to Baer’s thinking.
He explicitly stressed that embryogenesis was not determined solely by the
‘‘preceding state,’’ but also guided by the ‘‘essence’’ (Wesenheit) or ‘‘idea’’ of the
future organism (von Baer 1828/1999, p. 147, 148). As already discussed, this
‘‘idea’’ stood for what remained eternally the same—in contrast to individuals,
which existed only temporarily. Interestingly, Baer also regarded ‘‘species or
generative series’’ as passing phenomena (von Baer 1833–1834/2006, p. 61). He
believed it perfectly possible that individual species would die out or be transformed
over the course of history, though he rejected the notion that all animals had
developed by evolving out of one another (von Baer 1833–1834/2006, p. 56). For
Baer, then, in organic nature developments could not arise from living material
alone.12 Processes of becoming and passing away were only partially subject to
time—and thus also to history—because ultimately they were always determined by
a higher, ideal and stable order.
10 Baer was particularly critical of the recapitulation hypothesis as formulated by Etienne Serres
(1786–1868). See Schmuck (2009, pp. 200–213) and Meyer (1935).11 ‘‘The peripheral or radial type, the jointed or extended type, the massy or mollusc type, and the type of
the vertebrates’’ (von Baer 1828/1999, 209). On Baer’s typology and the teleological thinking associated
with it, see Lenoir (1982, p. 86), Lenoir (1985), Oppenheimer (1963) and Riha and Schmuck (2011).12 Later, in the 1860s, Baer turned against Darwin’s theory of common descent. On Baer’s view of
evolution, see Brauckmann (2012), Riha and Schmuck (2011, pp. 180–212), Holmes (1947) and
Breidbach and Ghiselin in Baer 1999.
Seeking the constant 41
123
Baer’s aspiration to discover what is continuous or ‘‘abiding’’ in the ‘‘transience’’
of organic forms (von Baer 1833–1834/2006, p. 40) shows similarities with the
concept of history and time that was moulded by the Naturphilosophen around
1800. As discussed in the introduction, they rejected linear development, sought
continuities and origins, and attributed change to ‘‘ideas’’ that remained eternally
constant (von Engelhardt 1979, pp. 111–142). However, significant differences can
be identified between Baer’s views on organic nature and those of the Naturphi-
losophen. If thinkers like Friedrich Wilhelm Schelling (1775–1854) and Lorenz
Oken (1779–1851) had redefined the eighteenth century’s epigenetic theories of
generation in line with gender dualities (see von Engelhardt 1985; Reill 2005,
pp. 220–236; Lettow 2014; Vienne 2014), I contend in the following that Baer’s
concept of development as an unending continuum and his search for general laws
of formation was accompanied by a turn away from the two-sex model. This
argument proposes a new way of reading Baer’s study of the mammalian ovum. In
the literature, no account has hitherto been taken of the fact that Baer’s ‘‘discovery
of the human ovum’’ (Schmuck 2009, p. 189) was also a response to the attempt by
two of his contemporaries to attribute embryogenesis to two different, sex-specific
generative substances.
3 Contested origins: Baer’s response to the work of Jean-Baptiste Dumasand Jean-Louis Prevost on sperm and fertilization
Immediately before Baer published his study on the mammalian egg, Dumas and
Prevost had presented their joint research on generation to the French Academy of
Sciences (see Prevost and Dumas 1821, 1824a, b, c, 1827).13 Although their work
on the physiology of generation and on embryology has attracted little attention in
the historiography of science,14 in the first two decades of the nineteenth century it
was received with interest and regarded as groundbreaking. Even in 1837, the
German physiologist Rudolf Wagner (1805–1864) still argued that it must be
‘‘regarded as the basis of all more recent investigations’’ (Wagner 1837, p. 402).
Baer knew the two men’s work. As we will see, his study on the mammalian ovum
firmly opposed their hypotheses. But what was the nature of the new perspective on
generation that Dumas and Prevost had opened up?
Unlike Baer, Dumas and Prevost did not dedicate their research exclusively to the
ovum, but also investigated seminal fluid. The first, and most significant, step in
their extensive studies on generation had been to examine the male reproductive
organs and ‘‘spermatic animals’’ of nineteen different animal species. Whereas the
13 While little is known about Prevost, Dumas entered the history books chiefly as one of the founders of
organic chemistry, alongside Justus Liebig (1803–1873). After the 1848 revolution in France, Dumas also
began a political career. He held high offices, including those of the Minister of Trade and Agriculture
and the master of the French mint. On Dumas’s life and work, see Chaigneau (1984) and Klosterman
(1985).14 Exceptions are Gasking (1967, pp. 137–147) and Farley (1982, pp. 37–42). Ilse Jahn counts Prevost
and Dumas among the founders of animal physiology, but does not mention their research on generation
(Jahn 1998, p. 350).
42 F. Vienne
123
majority of naturalists in the early nineteenth century believed sperm to be
microscopic animals and parasites, Dumas and Prevost established that sperm were
‘‘produced’’ in the testes (Prevost and Dumas 1821, p. 106). Their attempt to study
sperm as specific products of the male body was fundamentally new, as was their
attribution of embryogenesis to a process that had previously almost never been
analysed microscopically: fertilization. Early nineteenth-century physiologists and
embryologists equated fertilization with the act of copulation. It was new to describe
fertilization as a distinct physiological process and to ask about its relevance to
embryogenesis. Dumas and Prevost’s numerous experiments in the artificial
fertilization of frogs’ eggs showed that only those eggs exposed to seminal fluid
underwent the processes of transformation that culminated in the formation of an
embryo. To capture the significance of the changes that occurred immediately
after—and as a consequence of—fertilization, they observed the frogs’ eggs hour by
hour from the moment of fertilization onwards. They noted the changes taking place
between fertilization and the appearance of the organs, and inscribed those changes
into a developmental series that followed a chronological and linear structure.
Fertilization was the starting point, and the point of reference, for all the later
developmental stages of the ovum and embryo (see Prevost and Dumas 1824b,
pp. 107–121). In a further series of in vitro fertilization experiments, Dumas and
Prevost began to describe the process of fertilization more precisely: the spermatic
animals penetrated the mucus surrounding the ovum and thus came into ‘‘close
contact’’ with the ovum itself (Prevost and Dumas 1824c, p. 134).
Dumas and Prevost thus declared the instigator of generation to be an organic
entity—the spermatic animal—that was defined as male. Without its action, the
ovum could not develop into an embryo. As for the fortunes of the spermatic
animals after fertilization, Dumas and Prevost held different opinions. While
Prevost assumed that the spermatic animal disappeared after fertilization, Dumas
hypothesized that it went on to make a material contribution to the embryo’s
formation. For him, the spermatic animal was ‘‘nothing other than the rudiment of
the nervous system’’ (Prevost and Dumas 1827, p. 452; see also 443–454). Dumas
even succeeded in finding evidence for his claim. Studying the embryogenesis of
mammals, he had observed a ‘‘primitive line’’ twelve hours after copulation that
could not be explained in any other way than as ‘‘the rudiment of the nervous
system’’ (Prevost and Dumas 1824c, p. 132). The explanation for ‘‘inherited
similarities’’ between fathers and their offspring that Dumas had thus supplied
(Prevost and Dumas 1827, p. 452) laid the foundations for a highly consequential
scientific enterprise in the nineteenth century: the attempt to establish a material
continuity, on the level of their organic components, between the new life and both
of the sexes. As mentioned, Dumas and Prevost’s innovative results and hypotheses
were much discussed in the first decades of the century, though for the most part in
negative terms.
Baer was among the most rigorous critics of Dumas und Prevost’s theses. He and
Burdach both raised objections in Burdach’s Die Physiologie als Erfahrungswis-
senschaft. They declared it quite impossible that ‘‘the spermatic animals’’ were
‘‘originally present’’ and were produced by a ‘‘secretion’’ (Burdach 1826, p. 90, 93).
Particularly unacceptable was the notion that these animals played a role in
Seeking the constant 43
123
generation and might even form the basis for the development of another organism’s
nervous system (Burdach 1826, pp. 542–543; see also von Baer 1837/1999, p. 5). In
1827, Baer’s desire to distance himself from Dumas and Prevost’s theories induced
him not merely to defend the animal and parasitic character of sperm, but to redefine
their classification within the system of nature: he assigned them to the cercariae, at
that time considered a genus of snail parasites. Because they were not identical with
these, he described them as ‘‘a very low level of the cercaria type’’ and, to avoid
ambiguity, gave them a name of their own—‘‘spermatozoa’’ (von Baer 1827,
p. 640).15 Dumas and Prevost having elevated sperm to an entity indispensable to
generation, Baer thus demoted them to the lowest ranks of a parasitic genus. And
indeed, Dumas and Prevost’s findings contradicted Baer’s view that fertilization was
not a new beginning but the continued development of something already present.
Against this background, Baer’s study of the mammalian ovum may also be read
as a response to Dumas and Prevost’s hypotheses. If Dumas and Prevost accorded
priority—in both material and temporal terms—to fertilization and the elementary
particles of the two sexes’ generative substances in the process of organic
formation, Baer ascribed no more than a secondary function to the two sexual
generative substances and the process of fertilization. As discussed in the previous
section, Baer showed firstly that in mammals as in other animals, the process of
organic formation sprang not from a generative substance, but from a ‘‘previously
organized little body.’’ What distinguished Baer’s newly described ovulum or foetal
ovum from the ‘‘Graafian vesicle’’ was precisely that, unlike the vesicle, it did not
carry any feminine connotation; as I have pointed out, Baer emphasized that it was
present prior to the ‘‘maternal ovum.’’ As such, he attributed the process of organic
formation to an entity that was not sex-specific. But the ‘‘foetal ovum’’ also existed
before the paternal semen could drive on its development into an embryo. Baer saw
the role of the male generative substance in fertilization as being to ‘‘make the idea
of the animal complete and endow it with the possibility of development’’ (von Baer
1828/1999, p. 152, see also 147–152). In contrast, he ruled out any immediate and
material paternal share in the formation of the embryo at the moment of
fertilization. Baer rejected Dumas and Prevost’s observation that the sperm entered
the ovum during fertilization (von Baer 1827/1956, p. 145). Although he reproduced
Dumas and Prevost’s description and illustration of canine ova ‘‘of twelve days’’ in
order to complete his developmental series (von Baer 1827/1956, p. 128), he did not
mention that Dumas had interpreted the ‘‘line’’ observed at this stage as signalling a
material contribution by the sperm to the construction of the new embryo.
Accordingly, at the end of his study Baer noted as a basic principle that ‘‘the male
semen acts through the membrane of the ovum [cuticula ovi], which is pervious by
no foramen, and in the ovum it acts first on certain innate parts of the ovum’’ (von
Baer 1827/1956, p. 149).
Whereas Dumas’ reinterpretation of the ‘‘spermatic animals’’ was associated with
an attempt to apprehend a material continuity between offspring and both
progenitors, Baer wanted to assert an organic continuity that went beyond the
15 The literature on Baer makes virtually no mention of his natural-history approach to sperm and its
significance for his embryological research. An exception is Riha and Schmuck (2011, pp. 179–188).
44 F. Vienne
123
emergence and decay of individual organisms. For this reason, he attributed the
process of organic formation in all animals, and also in the human being, to a single
entity—the ovulum—that was regarded as independent of the female body. More
generally, Baer viewed generation not from a two-sex perspective but in terms of its
congruence with other physiological processes, especially those of ‘‘growth’’ and
‘‘nutrition.’’ For Baer, all these processes had to be understood as part of a unified
and continuous process of formation. Thus, ‘‘growth is nutrition with the formation
of new bodily mass—in fact, it is continued generation, and generation is nothing
other than the beginning of individual growth’’ (von Baer 1837/1999, p. 4). In the
late 1830s, Schwann pursued a similar point, ascribing all physiological processes in
plants and animals—including the formation of new organisms—to a unified and
universal basic unit of life: the cell. Schwann explicitly set out his concept of the
‘‘cell’’ in analogy to an ‘‘Individual, an independent Whole’’ (Schwann 1839/1847,
p. 2).16 According to him, each cell possessed ‘‘an independent life’’ and thus ‘‘a
power of its own’’ (Schwann 1839/1847, p. 192). Applied to generation, this view
conflicted with the prevailing assumption that the action of the semen was necessary
to trigger the egg’s development into an embryo. Schwann admitted that the ‘‘ova of
higher animals’’ may in fact receive something that was ‘‘more than nutrient
matter’’ through the act of fertilization. Many species, however—those comprising
‘‘female individuals only’’ and ‘‘lower plants’’ procreating without fertilization—
showed that in general each cell was an autonomous entity able to develop into a
new organism without an external stimulus (Schwann 1839/1847, p. 192). In
Schwann’s version of cell theory, thus, a male contribution was not a necessary
precondition for organic formation. While Baer accorded fertilization and the two
sexes’ generative substances a subordinate function, Schwann’s cell theory
implicitly abandoned the model of bisexual reproduction altogether.17
4 Conclusion: rethinking early nineteenth-century perspectiveson developmental history
As mentioned in this paper’s introduction, historians of biology have often
discussed how the emergence of cell theory was influenced by thinking and research
based on developmental history (see Temkin 1950, p. 237; Balan 1979,
pp. 281–285; Duchesneau 1987, pp. 113–128; Lenoir 1982, pp. 112–134; Jacyna
1990; Parnes 2000). In his history of heredity Francois Jacob argued that Baer’s
embryological work of the 1820s and the cell theory of the 1830s played a key role
in the formation of our understanding of reproduction. With these studies, according
to Jacob, procreation was no longer regarded as the production of a completely new
beings, but as the formation out of an already existing organic fragment—a basic
16 With his principle of cell autonomy, Schwann also opposed the teleological assumptions so central to
Baer’s work. There were thus also significant differences between Baer’s and Schwann’s understanding
of organic processes, which cannot be detailed here. It was Schwann’s notion of the autonomous action of
cells that Baer particularly criticized after the formulation of cell theory (von Baer 1866, pp. 380–386).17 Matthias Jacob Schleiden (1804–1881), commonly regarded as the co-founder of the late 1830s cell-
theory, held a similar view, see Farley (1982, pp. 48–51).
Seeking the constant 45
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unit of life, a ‘‘cell.’’ In his account, Jacob takes into consideration Dumas’ and
Prevost’s investigations of sperm and claims that their findings immediately and
‘‘definitively’’ established the necessity of the sperms in generation (Jacob 1993,
p. 121). My analysis shows, however, that neither Baer nor Schwann took up
Dumas’ hypothesis. More generally, the examples discussed in this paper highlight
that it is not enough to assume a single, homogeneous way of thinking based on
developmental history, or a straight line leading the way to the formulation of cell
theory and our understanding of reproduction. In the nineteenth century’s first
decades, very different forms of thinking and researching within a developmental
history frame were still arising. In particular, there were contradictory views on the
relevance of sexual difference for generation. In this period, the search for general
laws of formation and the search for a bisexual explanation of reproduction
constituted two different, and opposing, programmes of research.
What all these models of developmental history had in common, however, was
their strong focus on the ovum. Physiological studies of generation and embryo-
logical studies in the first third of the nineteenth century made the ovum the prime
category of developmental history. Regarding the sperm, in contrast, a natural-
history approach remained intact during that period. As I have shown, even in the
late 1820s Burdach and Baer were still describing the sperm using a way of thought
drawn from natural history. They were not alone in this. The dispute around the
pioneering theses of Dumas and Prevost, a debate that continued into the 1840s,
prompted their opponents to affirm and reaffirm the animal—in some cases even
parasitical—character of sperm.18 Even Dumas and Prevost’s new interpretation of
sperm as products of the male body and as reproductive units did not persuade them
to study these entities as the outcome of a process of organic formation that
unfolded step by step, or to equate them with other elementary organic units (for
example the ‘‘globules’’ of blood, which they had researched previously19). Instead,
a large portion of their research on the sperm of various animal species consisted in
describing the sperm’s shape, size, and manner of movement, as if they were
describing animals (see Prevost and Dumas 1821). Yet even the formulation of cell
theory in the late 1830s did not bring an abrupt end to spermatic animals as an
object of natural history. Although Schwann was interested in a cell-based
reinterpretation of the ovum’s structures and elements as described by Pander,
Purkyne and Baer (Schwann 1839/1847, pp. 46, 217–218), he saw no reason to
rethink the notion of spermatic animals in the light of his theory. This was because
the sperm were completely superfluous for Schwann’s law of formation, just as they
were for Baer’s explanation of embryogenesis. Research on ova and sperm in the
18 Leading German researchers in the physiology of generation in the 1830s were Rudolf Wagner
(1805–1864) and Carl Theodor von Siebold (1804–1885). They regarded sperms as the fertilizing factor
in semen, but nevertheless still as animals. See, for example, Wagner (1837), Siebold (1837). Among the
naturalists who described sperm as parasites were Carl Gustav Carus (1789–1869) (see Carus 1839) and
Gabriel Gustav Valentin (1810–1883) (see Valentin 1839).19 Dumas and Prevost’s studies of semen were preceded by research on the blood’s ‘‘globules,’’ and were
inspired by this to the extent that in both cases they were seeking the ‘‘active principle’’ of these organic
substances (Prevost and Dumas 1821, p. 196). In the historiography of biology, Dumas and Prevost have
featured as French representatives of ‘‘globule theory,’’ often considered a precursor to cell theory (Klein
1936; Pickstone 1973).
46 F. Vienne
123
first third of the nineteenth century, then, was influenced in very different ways by
developmental-history approaches.
The example of Baer shows with particular clarity that the advent of new
perspectives based on developmental history by no means presupposed the collapse
of all the previous static, classifying approaches and objects of natural history. In his
1821 lecture, Baer had derided contemporary natural history in its predilection for
renaming and proliferating the names of one and the same object. Yet he himself
made his own contribution to that tradition, by renaming the ‘‘spermatic animals’’
after having redefined their location within the system of nature. The longevity of
the ‘‘spermatic animals’’ as an object of natural history in the nineteenth century
thus also shows that the relationship between natural history and biology was not
solely one of discontinuity, but also embraced continuities. The image—propagated
most notably by Foucault and Lepenies—of a radical rupture between the static,
classificatory natural history of premodern times and the historical biology of
modernity may obscure our view of the continuities between these two forms of
knowledge about nature. The concept of sperm propounded by natural history, for
example, seems never to have faded completely; in an altered form it has
contributed to our modern understanding of the male germ cell. At the very least,
the label ‘‘spermatozoa’’ introduced by Baer has survived. At the time when he
coined this term, Baer mocked the idea that during fertilization ‘‘millions of such
spermatic animals battle murderously until the one that is left enters the vesicle of
the female ovary as a triumphant victor’’ (von Baer 1837/1999, p. 5). Yet it is that
very idea which we encounter again in today’s cellular understanding of
fertilization.
Translated by Kate Sturge
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