Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20

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Studies in History and Philosophy of Biological and

Biomedical Sciences
journal homepage: www.elsevier.com/locate/shpsc

Vital forces and organization: Philosophy of nature and biology in Karl

Friedrich Kielmeyer
Andrea Gambarotto
IHPST, 13, Rue du four, 75006 Paris, France

a r t i c l e i n f o a b s t r a c t

Article history: The historical literature on German life science at the end of the 18th century has tried to rehabilitate
Received 4 April 2013 eighteenth century vitalism by stressing its difference from Naturphilosophie. Focusing on the work of
Received in revised form Karl Friedrich Kielmeyer this paper argues that these positions are based on a historiographical bias and
17 July 2014
that the clear-cut boundary between German vitalism and Naturphilosophie is historically unattested. On
Available online
the contrary, they both belong to the process of conceptual genealogy that contributed to the project of a
general biology. The latter emerged as the science concerned with the laws that regulate the organization
Keywords:
of living nature as a whole. The focus on organization was, at least partially, the result of the debate
Romantic life sciences
Philosophy of nature
surrounding the notion of “vital force”, which originated in the mid-eighteenth century and caused a
Vital forces shift from a regulative to a constitutive understanding of teleology.
Karl Friedrich Kielmeyer Ó 2014 Elsevier Ltd. All rights reserved.
Friedrich Wilhelm Joseph Schelling
Gottfried Reinhold Treviranus

When citing this paper, please use the full journal title Studies in History and Philosophy of Biological and Biomedical Sciences

In the course of the last thirty years a considerable body of
scholarship has examined the life sciences that arose in Germany
at the end of the eighteenth century. This literature has shown
that previous dismissals of this tradition, assumed to be infected
with a pathological imagination, were unwarranted. Yet the in-
terpretations of the period have not always been consistent with
each other, and have often been characterized by vagueness.
Generally speaking, the scholarly debate has focused on the his-
torical and conceptual relationship between three elements: (1)
Kant’s philosophy of biology, (2) the biological vitalism developed
at the Göttingen medical school by Blumenbach and his students
Kielmeyer, Link, Treviranus, and Reil, and (3) the Naturphilosophie
of Schelling, Oken and Carus.
In his pioneering studies Timothy Lenoir (1978, 1980, 1981,
1982) argues that, although the life sciences developed in Ger-
many in the late eighteenth century have been dismissed as an era
dominated by empty speculation, they were in fact the result of
a coherent research program. This program was developed in
E-mail address: andrea.gambarotto@gmail.com.
Göttingen by a well-connected group of biologists after receiving its
first formulation in Kant’s Critique of the Power of Judgment in 1790.
In the second part of this work, Kant sees teleology as a necessary
tool to understand fundamental features of living beings such as
functions and development. He also considers it as a mere heuristic
principle, not as a constitutive character of organized bodies. Lenoir
claims that Blumenbach was the first naturalist who accepted the
Kantian understanding of teleological principles and organized it as
a structured research program. This program was first developed by
his most distinguished students Karl Friedrich Kielmeyer, Alex-
ander von Humboldt and Gottfried Reinhold Treviranus. This group
of naturalists is thereby addressed as “Göttingen School.” The
distinctive approach practiced at Göttingen derived from ideas
fashioned principally by Blumenbach during the 1780s and 1790s.
He synthesized some of the best elements of Enlightenment
thought on biology, particularly Buffon, Linnaeus and Haller, in
terms of a view of biological organization found in the writings of
Kant (Lenoir, 1981, 115). The disregard of this “Kantian” tradition in
life sciences has, for Lenoir, both theoretical and historical grounds.
The main issue is the assumption that only reductionist models are
capable of generating a quantitative account of natural phenomena.

http://dx.doi.org/10.1016/j.shpsc.2014.07.007
1369-8486/Ó 2014 Elsevier Ltd. All rights reserved.
 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20
Nevertheless, the idea that biological organization is not reducible
to the laws of physics and chemistry is fully compatible with the
fidelity to quantitative rigor as a touchstone of scientific explana-
tion. According to Lenoir, the “vital-materialism” of the Göttingen
School accepted this challenge and developed a “teleo-mechanical”
research program based on the Kantian distinction between
constitutive and regulative understandings of teleology. On the
other hand, ascribing constitutive character to teleology, i.e.
considering it as a real feature of living bodies, the Natur-
philosophen exceeded the boundaries of science in the direction of
empty speculation. From the historical point of view, the main
reason for disregarding the Göttingen tradition is the assumption
that it was just another example of speculative philosophy of na-
ture. Lenoir believes this assumption is proven wrong by textual
evidence, since both Kielmeyer and Treviranus openly criticize
Naturphilosophie. Thirty years later, Hans Peter Reill (2005) unifies
the vital-materialism of the Göttingen School with the Montpellier
tradition of Bordeu, Barthez and Ménuret under the general label of
“Enlightenment vitalism.” Once again, the pivotal point of the
argument is the distinction of this tradition from Romantic Natur-
philosophie. Both accounts share a common argumentative pattern.
Their fundamental aim is to rehabilitate eighteenth century
vitalism by showing that its research program can be considered in
“naturalized” terms. Their most important concern is then to mark
the difference of this program from Naturphilosophie, which is
considered the metaphysical and anti-naturalist program par
excellence. In what follows, I argue that these positions are based on
a historiographical bias, using the work of Karl Friedrich Kielmeyer
as a basis for my argument.
Scholars have already argued that the alleged agreement
between Kant and Blumenbach was based on a substantial
misunderstanding of the respective conception of teleology:
Blumenbach ignores the Kantian distinction between constitutive
and regulative principles and conceives of the Bildungstrieb as a
goal-directed drive proper to all organized beings (Richards,
2000). For this reason the Lenoir thesis can no longer serve as
point of departure for the reconstruction of the German life sci-
ences of this period (Zammito, 2012). Resting upon these studies I
will develop my argument by showing that the clear-cut boundary
between the vital-materialism advocated by Lenoir and Natur-
philosophie is historically unattested. I will thereby position
myself in accordance with Richards (2002), but use a different
argumentative strategy. I will argue that Naturphilosophie is part
of the same process of conceptual genealogy that contributed to
the emergence of a general biology as a unified science. The latter
became possible only after the determination of its proper object
had reached completion. The object in question is “organization”
as a specific property of living nature. Biology emerged as the
science that deals with the laws regulating organization, both of
single natural bodies and of living nature as a whole. I will try to
show that the focus on the concept of organization was at least
partially the result of the debate on the notion of “vital force”
originated in mid-eighteenth century Germany. This debate led to
a functional interpretation of the scala naturae according to which
higher levels of organization display a greater number of vital
functions than lower ones. I will develop my argument as follows.
I will first provide a sketch of the different theories of vital forces
formulated in Germany before Kielmeyer, with a particular
attention to Haller, Wolff and Blumenbach. I will then take into
detailed account the lecture on organic forces held by Kielmeyer
in 1793 to verify the meaning of the claim of it being the first
systematic program of a general biology. Finally, I will show how
the framework put forward by Kielmeyer was coherently devel-
oped by Schelling and Treviranus.
13
1. Theories of vital forces before Kielmeyer: from Haller and
Wolff to the Göttingen School
It is a striking fact that the great majority of scholarly works
dedicated to the vital-materialism of the Göttingen School dealt
with the issue using the vocabulary of Lakatos. The idea of a Kante
Blumenach “teleo-mechanical” program for biology, formulated for
the first time by Lenoir, is still endorsed in recent studies (Bach,
2001; Dupont, 2007; Schmitt, 2006). This notion, however, is
inadequate for understanding the transformations that led to the
birth of biology at the beginning of the nineteenth century. The idea
that the Critique of the Power of Judgment provides a research pro-
gram for biology can in fact be criticized not only by emphasizing
the divergence of Kant and Blumenbach, but also by showing that
Kant’s attention to biological issues was not the result (at least not
primarily) of scientific concerns, but of strictly transcendental
questions (Huneman, 2012; Zuckert, 2007). On the other hand,
however, it is equally true that Kant dealt with biological matters in
at least three different respects: (1) the relationship connecting the
notion of Naturzweck to modern epigenesis, since the process of
embryogenesis seems to presuppose its result (the adult organism)
and be directed towards its realization (Huneman, 2007; Zammito,
2007); (2) the problem of functions, which cannot be explained
without referring to final causes (the structure of a bird, for
example, is apparently grounded in the purpose of allowing flight:
Illetterati, 2008); and (3) the difference between Naturbeschreibung
and Naturgeschichte, i.e. the discussion of the epistemological status
of natural history as descriptive cataloguing or causal explanation
of varieties (Fischer, 2007; Sloan, 1979, 2006).
This discussion is not the result of a research program but of a
conceptual shift taking place in the last decades of the eighteenth
century. It has been pointed out (Cheung, 2006) that during the
eighteenth century the word “organism” generally refers to a spe-
cific form of order that could apply to different kinds of entities
such as plants and animals, but also artifacts. At the end of the
eighteenth century, the term became a generic name for individual
living entities, by around 1830 it became a recurrent technical term
in the emerging biological disciplines. Theories and models of
living beings developed until the late eighteenth century must
therefore be defined as pre-biological. With regard to this frame-
work, even Kant makes no exception: although the critique of
teleological judgment has in different ways been regarded as an
endeavor towards the scientific foundation of biology (Löw, 1980;
Marcucci, 1972; Philonenko, 1982; Zumbach, 1984), Kant’s posi-
tion on this issue is actually much more cautious (Ginsborg, 2001,
2006; Goy & Watkins, 2014; Guyer, 2005; Illetterati, 2010;
McLaughlin, 1989, 1990; Zammito, 1992). According to Kant,
biology may indeed never be regarded as a proper science, as the
consideration of living beings implies reference to teleological
principles, and the use of these principles has only regulative
character. On the other hand, from the late eighteenth century, the
term “biology” began to appear in the works of several naturalists,
the most important of which is the monumental Biologie, oder
Philosophie der lebenden Natur für Naturforscher und Ärzte (1802e
1822) by Gottfried Reinhold Treviranus. What happened between
1790, the year of publication of the third Critique, and 1802, when
Treviranus used the term “biology” as the title of a scientific work
concerned with living nature as a whole? I wish to argue that the
answer lies in the shift involving the modern semantics of orga-
nization. The birth of biology as a field is better understood in
genealogical terms, considering the way through which “organi-
zation” became a specific object of natural science. In what follows I
will argue that this process is intertwined with the conceptual
history of the notion of “vital force.”
14 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20
The undisputed father of the notion of vital force is Albrecht von
Haller. He joined the Göttingen faculty in 1737 and from 1747
directed the Göttingische Anzeigen der gelehrten Sachen, a Journal
founded in 1739 which hosted important scientific debates in
embryology and physiology. Eighteenth century physiology pays
tribute to Newton’s natural philosophy. It assumes the notion of
“irreducible properties” of matter whose laws of expression had to
be formulated without speculating on the cause of their origin
(Wolfe, 2014). This concept is combined with the notion of fiber,
used since the end of the seventeenth century by Francis Glisson for
his study on the contraction of muscles. Especially in his 1672 work
De natura substantiae energetica, Glisson was the first to bind the
notion of fiber to the specific property of irritability. Living fiber is
flexible, extensible, resistant, elastic and irritable, i.e. it has a radical
force activated by external or internal stimuli, and causes the
change of shape and all remaining vital phenomena. For Glisson,
the irritable fiber provides a general overview of a character of the
living body, not a concept defining a specific property. As Boer-
haave’s student, in the general context of a mechanistic physio-
logical framework, Haller gave specific attention to fibers. The first
appearance of irritability can be found in a note on Haller’s own
edition of Boerhaave’s Praeselectiones academica, where he remarks
that the heart keeps beating after it has been extracted from the
animal’s body. It is however in the dissertation De partibus corporis
humani sensilibus et irritabilibus (1752) that he further develops the
notion. Haller claims to have proven by animal experiments that
only the muscular fiber possesses the ability of contraction, which
he defines as “irritability”, which was responsible for movement.
From this property he clearly distinguishes “sensibility,” which is
responsible for sensual impressions and inherent only to nerves
and parts furnished with them. He thereby challenges the tradi-
tional mechanical (mainly Boerhaavian) model on three main
points. First, he postulates a force inherent in the muscular fiber
and independent of the nerves and the soul. Second, he concep-
tually and physically separates the two properties of movement
and sense perception. Third, he establishes a strict correlation be-
tween structure and function not on the level of elementary par-
ticles but of compounding structures, i.e. muscular and nervous
fibers (Monti, 1990). The explicit connection made between struc-
ture and function poses a question concerning the purposiveness of
the structure. Haller never doubted that heart, muscles, intestines
etc. had the structure needed to perform their function. Even the
deformed organs of the so-called “monsters” were arranged in such
a way as to most easily fulfill their duty. This teleological postula-
tion underlays Haller’s physiological arguments and allows
the correlation of structure and function. Haller establishes the
correlation of muscle-irritability and nerve-sensibility on what he
considers a sufficient experimental basis. However, as a prefor-
mationist, the teleological arguments, for Haller, are based on the
assumption that objects are formed in accord with intelligent
design. It was not nature itself, but God who arranged structures,
laws and functions. As shown by his embryological positions, Haller
envisages nature as a machine that runs according to prescribed
laws of mechanics. For this reason, Haller’s physiology does not
break with Boerhaavian mechanism, but is rather a full acknowl-
edgment and elaboration of non-reductionist elements in the late
conception of his teacher (Steinke, 2005, 94e96). The integration of
this physiological theory on vital properties into a framework of
self-organization was a consequence of modern epigenesis.
Georges Canguilhem has argued that research concerned with
the origin of organic models in biology would have to begin with
the detailed consideration of Caspar Friedrich Wolff’s doctoral
dissertation Theoria Generationis (1759) (Canguilhem, 1962, 13). A
copy of the dissertation was sent to Haller, triggering his review in
the Göttingische Anzeigen von gelehrten Sachen, which started the
famous debate between them (Roe, 1981). The second edition of the
work, Theorie von der Generation (1764), is not merely a German
translation and restatement of the theory, but it also entails a
detailed attack on the preformationist views of Haller and Bonnet.
It was followed by a major response in Haller’s Elementa physi-
ologiae corporis humani (1766). The pivotal concept of the work is
the notion of “essential force” (vis essentialis). According to Wolff,
generation cannot be conceived without the existence of a peculiar
force inherent to epigenetic development. Plants and animals arise
from raw material, an amorphous and undifferentiated substance.
Within it, vesicles (vesicuale) and vessels (vasae) are formed as
spherical cavities filled with liquid. Vesicles then become organs
that appear one after the other. According to this model, the
streaming of liquid and its tendency to coagulate were sufficient to
explain the formation of plants and animals without reference to
preexistent structures. The tendency of vegetable and animal em-
bryos to coagulate is always caused by the vis essentialis. Each part
secretes the other after its own formation in a gradual process of
self-organization. As each part begins to solidify, it becomes orga-
nized and acquires vessels and vesicles produced by the movement
of fluids into the new parts. In higher animals, the first structure to
solidify is the spinal chord, from which all other organs are sub-
sequently produced. These processes are caused by the essential
force, which Wolff deems possible to postulate on the basis of
observational evidence: “the essential force along with the ability
of nutrient fluid to solidify constitutes the sufficient principle of all
vegetation [i.e., development] both in plants and in animals” (Wolff,
1759, x 242).
French vitalism developed only marginally with respect to the
problem of epigenesis and more directly on Haller’s physiological
model. The works of Bordeu on glands, for example, led him to
attempt a direct characterization of functional factors. Based on the
idea of the sensibility of parts, he developed a model of organiza-
tion grounded on the integration of elementary organs. The inte-
gration of complex structures is the emergent effect of elementary
processes that can be reduced to phenomena of tonicity and reac-
tivity. Such a notion of the organic machine was criticized by Bar-
thez, who instead tried to establish that the phenomenon of vital
mobility, like those of sensibility and of “sympathies,” translates the
regulative activity of a vital principle heterogeneous from structural
and mechanical conditions (Duchesneau, 1982, 361e430). In Ger-
many, on the other hand, the groundbreaking relevance of Haller’s
physiology was measured up to Wolff’s compelling arguments. The
idea of a structuring order operating in the epigenetic process
produced a vitalist modification of the Hallerian model in which the
idea of self-organization played a seminal role.
Johan Friedrich Blumenbach (1752e1840) studied in Göttingen
and graduated in 1775. He was appointed extraordinary professor
of medicine and inspector of the museum of natural history in
Göttingen in 1776 and ordinary professor in 1778. Like Haller he
was in the habit of illustrating his theory by a careful comparison of
the animal functions of man with those of other animals. Rejecting
Haller’s preformationism while simultaneously proposing a teleo-
logical agent as sufficient reason for the resulting order, Blu-
menbach’s physiology operates as a significant variation on the
Hallerian model. In his Institutiones physiologicae (1787), Blu-
menbach uses the notions of “force” and “drive” to indicate five
fundamental properties of living matter: (1) formative drive (nisus
formativus), which is responsible for developmental organization
and remains active in the organized body throughout its life; (2)
contractility (vis cellularis), which is responsible for the contraction
of mucosa; (3) irritability (vis muscolaris), which is responsible for
the contraction of muscles; (4) vitae propriae, which is responsible
for the contraction of specific organs such as the iris or the fallopian
tube; and (5) sensibility (vis nervea), which allows perceptive
 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20
functions (Blumenbach, 1828, 27e33). The peculiarity of this
schema is the presence of the formative drive (nisus, Trieb) as
somehow opposed to the other forces (vis, Kraft). While in fact
Blumenbach associates the notion of force to a blind agent, he
considers the nisus as a goal-directed drive responsible for the
specific form of the single organism. In the famous paper Über den
Bildungstrieb (1791) he argues that in the unformed matter of an
organized body is active “a peculiar lifelong drive” which “pushes it
towards its determined shape.” This drive is “clearly different from
all the other vital forces of living bodies (contractility, irritability,
sensibility etc.) and from all the universal forces of bodies in gen-
eral.” It is “the first important force for every generation, nutrition
and reproduction, and that can be called formative drive (Bil-
dungstrieb, nisus formativus) in order to distinguish it from other
vital forces” (Blumenbach, 1791, 32e33). Here Blumenbach refers to
“a perspicacious physiologist, Caspar Wolff,” who “assumed
another force for the growth of animal and plants, that he calls vis
essentialis. If one knows it only by hearsay, he could confuse it with
my nisus formativus.” According to Blumenbach, however, the dif-
ference is clear as soon as one reads the definition of the vis
essentialis given in the Theoria generationis. For Wolff the vis
essentialis is merely the force by mean of which the nutritive ma-
terial is driven through the plant or the animal. This is for Blu-
menbach only one aspect of the formative drive. The essential force
can be observed among less formed outgrows (like tree strains)
where no definite formative drive is to be found. On the other hand,
the essential force can be very weak in poorly nourished organic
bodies, whose formative drive is unscathed (Blumenbach, 1791,
39e41). An explicit discussion between Wolff and Blumenbach
took place in 1789 in a publication titled Zwo Abhandlungen über die
Nutritionskraft. The volume includes a long paper, Von der eigen-
tümlichen und wesentlichen Kraft der vegetabilischen sowohl als auch
der animalischen Substanz. This treatise was written as a response to
and published along with two essays on similar topics by Blu-
menbach and Carl Friedrich Born. Both being selected for first-place
honors in a competition held by the St. Petersburg Academy of
Sciences for the best essay on the nature of the “nutritive force.” In
this sense, Blumenbach’s vitalist physiology represents, as it were,
the point of intersection between Haller’s physiological model and
Wolff’s theory of generation. He multiplies the vital properties of
the former calling them “forces” and includes the vis essentialis of
the latter under the name of “formative drive.” The latter displays a
teleological character that was extraneous to Wolff. The Bildung-
strieb does not only represent the vital principle of organic self-
construction, but also the sufficient reason of the resulting vital
forces that exercise their action in the different organs. From this
point of view, the Bildungstrieb supplies an explanation for the
different functions present in the living body.
In conclusion, Haller’s “animated anatomy” opened a new field
of investigation concerned with the vital properties of fibers. He
thereby adopted a conception of organic functions that relies on the
specific properties of elementary structures. Blumenbach modifies
this framework on the basis of Wolffian epigenesis, elaborating an
organic model characterized by a multiplication of those proper-
ties. Like Haller, Blumenbach understood the vital forces as cor-
relatives of specific physiological properties, but their reciprocal
relation was understood as forming a functionally integrated
whole. The main result of the notion of Bildungstrieb is the
conceptualization of an agent of organic formation modeled on
Wolff’s vis essentialis. In Blumenbach’s physiological formulation
this was a model for the distribution of the different vital forces in
the individual living body. This model was employed by Kielmeyer
to account for the distribution of vital functions in the animal
kingdom. This gave rise to a peculiar line of inquiry in animal
taxonomy based on the different vital forces displayed by various
15
species. Kielmeyer’s lecture Über die Verhältnisse der organischen
Kräfte untereinander in der Reihe der verschiedenen Organisationen,
die Gesetze und Folgen dieser Verhältnisse (1793) is the first and most
important example of this program, which has been properly
defined as “physiological systematics” (Schmitt, 2006).
2. Vital forces and organization: Kielmeyers’s lecture as a
program for a general biology
Karl Friedrich Kielmeyer (1765e1844) began his studies at the
Karlsschule in Stuttgart and furthered his education at the Univer-
sity of Göttingen between 1786 and 1788 under Blumenbach’s su-
pervision. He returned afterward to Stuttgart where in 1792 he was
appointed professor of chemistry and zoology. He published little in
his lifetime, but on February 11th 1793 he delivered a famous
plenary lecture in honor of the sixty-fifth birthday of Duke Carl
Eugen of Württemberg. In this lecture Kielmeyer attempted to
demonstrate, on the basis of inductive evidence, that peculiar
teleological laws regulate the distribution of vital functions
throughout the animal kingdom. Concerned with a general theory
of animal organization, Kielmeyer’s lecture has been regarded as
the first Systemprogramm of biology (Bach, 2001; Kanz, 1994).
In his lectures on comparative zoology, held in Stuttgart be-
tween 1790 and 1793, Kielmeyer defines this program as Physik der
Tierreichs (physics of the animal kingdom). This discipline had to
account for: (a) the number of organs in the machine of the animal
kingdom, the number of animal forms, and the laws according to
which they are divided into different groups Additionally, the
discipline must consider the causes, consequences or purposes of
these classification (Zwecke); (b) the relative position of the organs
in the machine of the animal kingdom, the division of the animal
kingdom into groups upon the earth (geography) according to
different characters and laws, causes and effects of the differences
according to different groups; (c) the interrelated formation of or-
gans in the animal kingdom, the gradation of animals and affinities
in their formation generally as well as according to groups, and the
laws, causes, and effects of this gradation; (d) changes in the animal
kingdom and its groups, and the developmental history of the animal
kingdom in relation to the epochs of the earth and those probable
for our solar system, which are symbolized by the parabola; (e)
changes which the animal kingdom and its groups undergo
repeatedly throughout all epochs, and the life of the machine of the
animal kingdom or its physiology, which is symbolized by the circle
(Kielmeyer, 1938, 28e29).
This program never resulted in an extensive work, but the Rede
held in 1793 constitutes an incisive outline. Like Blumenbach,
Kielmeyer considers natural purposiveness as a proper feature of
living bodies, which he explicitly refers to as “organizations”: “let
us grant that nature had no intention in establishing this artful
juxtaposition of appearances in time, that effects and their conse-
quences were to form no goals that she had wished to achieve,”
nonetheless “we still must confess that the chain of effects and
causes in most cases seems like a chain of means and ends to us and
that we would find it advantageous for our reason to assume such a
chain” (Kielmeyer, 1793, 6). As stressed by Richards, Kielmeyer’s
remarks are here grammatically cast as counterfactual sub-
junctives. They grant that nature might not have intrinsic purposes
and that the search for higher goals might appear to be a mere
fantasy. These concessions, however, display a direct reference to a
Kantian argument. In the third Critique, Kant maintains that any
inductions focused on organic processes would lead to poetic
phantasizing (dichterisch zu schwärmen), as one conjures up tele-
ological principles supposedly governing those processes. Kiel-
meyer’s point is rather that such inductions nonetheless convince
us, and properly so, that nature is teleologically structured
16 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20
(Richards, 2002, 242). Organic forces must therefore be regarded as
teleological principles which make organisms different from
nonliving matter. On these premises, the questions Kielmeyer
wished to address were: Which forces gathered in most in-
dividuals? What are the reciprocal relationships between these
forces in different kinds of organizations? And according to which
laws are these relationships modified in the series of different or-
ganizations? On the basis of Blumenbach’s schema, Kielmeyer
distinguishes five forces: (1) sensibility, or the ability of the nerves
to retain representation; (2) irritability, or the ability of muscles and
other organs to respond to stimulation through contraction; (3)
reproductive force, or the ability of organization to restore injured
parts of a creature or to produce a new individual of like kind; (4)
secretive force, or the ability to deliver different fluids to the right
places; and (5) propulsive force, or the ability to move fluids through
vessels. The original Hallerian forces, sensibility and irritability,
remain unaltered, while the formative drive is considered as a force
among the others (not as principle of organization) corresponding
to reproductive functions. The vis cellularis and the vitae propriae
are replaced by secretive and propulsive force. This difference is
irrelevant since it involves the least important forces of both
schemas. The latter are, in fact, not even taken into account in
Kielmeyer’s lecture, while several examples are provided with re-
gard to the others. First, Kielmeyer deemed necessary the estab-
lishment of the criterion according to which the magnitude of
various forces had to be measured and compared. Frequency of
effects at equal times, plurality and magnitude of the resistance to
other forces, or the permanence of the effect under equal condi-
tions were considered particularly suitable for the task.
Sensibility is the first organic force he takes into account.
Empirical evidence, argues Kielmeyer, shows that the faculty to
receive different sensations decreases across the series of organi-
zations from higher mammals to lower classes (Kielmeyer, 1793,
12e13). Among quadrupeds, birds, snakes and fishes, all sense or-
gans still display high perfection; however, among insects, the
auditory and olfactory organs are largely gone. Among worms, the
brain and most of the nervous system found in other animals is not
present, and a single organ collects all sensory stimuli. In plants,
this receptivity to impressions is present only in very obscure
traces. It is also evident, however, that when a sense organ is lost,
which effectively decreases the multiplicity of sensation, the
remaining senses are finer and more elaborate. Insects and worms,
mostly deprived of eyes and ears, have a significantly more devel-
oped sense of touch than monkeys and humans. The malfunc-
tioning eyes of moles seem to be compensated for by the
development of finer paws and noses. Other animals’ ocular
insensitivity similarly results in the sharpening of auditory and
olfactory organs. From these observations, Kielmeyer draws a law:
“the multiplicity of possible sensations decreases in the series of
organizations as much as the ease and finesse of remaining
sensation increases” (Kielmeyer, 1793, 19). The lack of multiple
kinds of sensations among lower animals is thus balanced by the
sophistication of those remaining.
After his discussion of sensibility, Kielmeyer provides empirical
observations on irritability. Nature has placed major differences not
only in relation to the multiplicity of its manifestations, but also to
the duration and frequency of their effects under the same condi-
tions. Among warm-blooded quadrupeds and birds, all traces of
irritability are extinguished as a consequence of the separation of
the body from the sensory apparatus or the division of individual
limbs from the trunk. By contrast, the manifestations of this force
are almost indestructible in cold-blooded animals: frogs continue
hopping after decapitation, and turtles continue moving for days
after experiencing fatal heart trauma and/or amputation of the
head. Similar phenomena are observed among fish and insects: the
feet of some spiders, for example, have been observed moving for
more than seven days after being removed from the body. On the
other hand, the majority of animals displaying such enduring irri-
tability tend to be slower in at least some of their movements. The
movements of most amphibians are weak and their hearts beat
more slowly than those in warm-blooded animals. Finally, Kiel-
meyer points out that most animals with long-lasting irritability
are precisely those in which the multiplicity of sensations has been
greatly limited. The opposite is the case for warm-blooded animals,
whose irritability is utterly temporary, but whose muscles are far
more articulated. Based on these observations, Kielmeyer formu-
lates a second law: “irritability, estimated according to the
permanence of its manifestations, increases as much as speed,
frequency and variety of these same manifestations, and the variety
of sensations decreases” (Kielmeyer, 1793, 23). The lack of duration
is thereby compensated for by frequency, variety, speed and better
sensibility.
The reproductive force, claims Kielmeyer, is the most universal
and distributed force among organizations, and could be defined as
the force that distinguishes them from other products of nature.
Since it is the most universal among organizations, a greater variety
of manifestation is to be expected. However, for Kielmeyer, this
amazing variety fits under a few simple laws. The number of
offspring produced at any one time by warm-blooded quadrupeds
is between one and fifteen. This number increases among birds, and
more so among amphibians and fishes. Insects, worms and plants
generate with greater frequency than all the previous classes. This
pattern suggests a general decrease of reproductive faculties from
complex mammals to simpler organizations. Higher organizations
generate a small amount of larger individuals in a longer time than
lower ones. A third law is also formulated: “the reproductive force,
the number of new individuals that will be created in a given place,
increases as much as the magnitude of what is to be produced; or
more generally, the dimension of individual product, as it appears
after birth, decreases” (Kielmeyer, 1793, 28). Animals with lower
fertility are also those of larger size and are more extensively
developed at birth. They are also those for which generation takes
more time. To form an elephant, for example, nature strains for two
years, while it only takes a few weeks to generate a rat. Kielmeyer
thus formulates a more specific and universal version of the law:
the more the reproductive force is manifested in a certain place by
the number of new individuals, the smaller the size of the body of
such individuals; the simpler is the constitution of their bodies, the
lower is the time it takes to form them in the body of the parents
(Kielmeyer, 1793, 30).
These are, for Kielmeyer, the laws that regulate the distribution
of the vital forces in the animal kingdom, which he generalizes as
follows: “the faculty of sensation is gradually replaced in the series
of organizations from irritability and reproductive force and finally
decreases the irritability of the latter [organizations]; the more the
first increases the less is the other. Furthermore, there is little
accord between sensitivity and reproductive force; the greater one
of these forces is at the time of initial development, the more the
other is neglected” (Kielmeyer, 1793, 35). In general, then, the fac-
ulties of sensation, widely developed in higher animals, gradually
decrease in lower classes and are endowed with great irritability.
Equally, in lower classes (insects, worms), irritability is gradually
replaced with reproductive force. The simplicity of these laws
becomes clearer after the realization that “the forces divided
among the various organizations are also those under which the
division of forces took place among the different individuals of the
same species, even in the same individual at different phases of
development” (Kielmeyer, 1793, 36). At the beginning of their
development, humans and birds are both similar to plants. They are
mainly endowed with reproductive force, only in the following
 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20 17

developmental phases does their irritability increase. Thereafter, which I refer the reader to the lecture of prof. Kielmeyer on this
senses hatch sequentially, according to the order in which they subject that already appeared in 1793, a lecture from which in
appeared in the series of organizations. The faculty of irritability the years to come a new epoch of natural history is to be ex-
thereby replaced by the faculty of representation, or at least in its pected” (Schelling, 2000, 252e253).
most immediate material organ. These far-reaching conclusions
have been interpreted as the first formulation of the principle of
The concrete realization of this program is found in the Erster
recapitulation, which will have extraordinary implications for the
Entwurf eines System der Naturphilosophie (1799), where different
theory of species evolution (Coleman, 1973, 1977; Richards, 1992).
animal classes are conceived of as the result of different relations of
This will be indeed one of the most appealing and long-lived doc-
the various vital forces. They differ from each other not only in their
trines of Naturphilosophie and transcendental morphology. The
material composition, but in the relation and proposition of their
core of this principle stresses the fact that many animals, as they
constitutive forces among themselves. According to Schelling, a
develop embryologically, temporarily manifest structures which
species is the expression of a certain proportion of primary func-
closely resemble those exhibited by the adult forms of species
tions. The continuity of organic functions throughout the animal
lower down in the scale of animal organization. The detailed
kingdom forms a universal organism. Schelling establishes a
explication of the law was carried out by Johan Friedrich Meckel
comparative physiology of organic functions, which endeavors to
(1781e1833) in Germany and by Étienne Serres (1786e1868) in
establish the various degrees and proportions of essential forces
France. It has often been referred to as the MeckeleSerres Law of
belonging to different organisms. In fact, every organism displays a
parallelism, in order to distinguish it from the later biogenetic law
specific proportion of reproductive force, irritability and sensibility.
formulated by Ernst Haeckel (1835e1919). In my view, however,
It is primarily defined not by its external form, but by the particular
this is neither the most important aspect nor the actual contribu-
proportion of forces active within it. Its form and organs will follow
tion of the lecture. Its meaning can be properly understood only
from the disposition of these forces, each one prominent in a
within the genealogical process connecting the various theories of
different class. Plants, for example, have a preponderance of
vital forces in the second half of the eighteenth century. Blu-
reproductive force while displaying no sensibility. Mammals have a
menbach’s contribution was the unification of Wolff’s vis essentialis
preponderance of sensibility but can generate few offspring and
with Haller’s physiological notions of sensibility and irritability,
they do not display regenerative features which are frequent in
which created a model of animal economy characterized by mul-
amphibians. Each organism is itself nothing else than “the collec-
tiple vital forces. The latter were responsible for different functions
tive expression for a multiplicity of actions (Aktionen), which
in the living body conceived as an organized whole. Kielmeyer
mutually limit themselves to a determinate sphere.” This conflict of
made use of a similar model to explain the distribution of vital
actions will express itself in certain functions. The variety of the
forces, not however in the individual body, but rather in the entire
animal kingdom results from the variation in proportion of these
living nature. Animal organization is thereby understood as the
functions. The latter have inverse relationships to one another, such
result of the organization of the “great machine of the organic
that as the one increases the other necessarily decreases. Functions
world.” In this sense, Kielmeyer’s Rede addresses the problem of a
must be reciprocally opposed to maintain balance, “which in itself
general biology, a unified scientific field concerned with the laws
already corresponds with the concept of an organism.” In a single
that regulate the organization of living nature as a whole through
organism any one of these functions could be dominant while the
its internal relations.
rest are present in a lower degree. On account of this opposition, it
is impossible for those forces to unify in the same part. The or-
3. Philosophy of nature and biology after Kielmeyer:
ganism in which they are unified must be divided into many single
Schelling and Treviranus
individual parts performing different functions. Through their
collective actions, these parts must produce the single organism:
The pivotal role of Kielmeyer’s Rede for the elaboration of the
“they would relate simultaneously to the whole organism as cause
conceptual framework of Schelling’s Naturphilosophie has been
and effect of its activity. That which so relates itself to the organism
already addressed by scholars (Bach, 2001; Schmitt, 2006, 2007).
(as a whole) is called an organ.” Functions must relate to different
The first work with biological relevance among Schelling’s writings
organs, such that “the more the multiplicity of the functions in-
is Von der Weltseele (1798), whose subtitle is “A hypothesis of higher
creases in the organic domain of nature, the more complexly the
physics for the explanation of the universal organism.” This venture
system of the organs develops” (Schelling, 2001, 114e115). This
is a development of the physiological framework put forward by
model conceives “the continuity of organic functions as principle of
Kielmeyer, with a peculiar emphasis on the functional lexicon. The
organization” (Schelling, 2001, 116, emphasis added). These pas-
animal kingdom is thereby understood as a balanced system of vital
sages are almost quotes from Kielmeyer’s Rede.
functions corresponding to different animal classes:
This Schellinghian interpretation of Kielmeyer is adopted by
“Since it is undeniable that in living beings takes place a series of Gottfried Reinhold Treviranus (1766e1837), who also studied with
functions (Stufenfolge der Funktionen), as nature opposed irri- Blumenbach in Göttingen, where he got his doctorate degree in
tability to the animal process, and sensibility to irritability, and 1796. In the first volume of his Biologie, oder Philosophie der lebenden
so arranged an antagonism of forces that keep each other Natuur für Naturforscher und Ärzte (1802e1822), Treviranus con-
reciprocally into balance, as while one increases, the other de- ceives of nature as a “universal organism” where no change can
creases and vice versa, so one will be led to the thought that all occur to a part without influencing the whole. He thereby criticizes
the functions are only branches of the same force, and that for Schelling for the assumption of a “world soul” as explanatory ground
example a natural principle, that we must assume as the cause of for the ongoing activity of the universe (Treviranus, 1802, 33). This
life, emerges in them only as in its own phenomena, just as without fact may lead to endorse Lenoir’s position. This critique, however,
doubt one and the same universally spread principle manifests must be read in context. Not many pages before criticizing Schelling,
itself in light, electricity etc. Since great naturalists reached the Treviranus addresses in fact Alexander von Humboldtdhis senior at
same result, it is possible to boldly trust this idea. In particular, it Göttingen and also Blumebach’s studentdwhose definition of the
is confirmed through the consideration of the progressing vital force he considers “even more useless” than that provided by
development of organic forces in the series of organizations, about Stahl (Treviranus, 1802, 19). Shortly after he criticizes Kant for his
18 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20
mechanistic conception of matter (Treviranus, 1802, 27, 39) and
explicitly agrees with Schelling that the reduction of fundamental
forces to “attraction” and “repulsion” makes it impossible to explain
the “specific qualities and forms” of living nature (Treviranus, 1802,
54). The fact that Treviranus criticizes Schelling’s assumption of a
world soul while accepting his idea of nature as a universal organism
proves that he considered him as much a reliable discussant as all
other major figures of late eighteenth-century science.
Other passages of the Biologie provide further evidence for this
claim. In the fourth volume Treviranus maintains that life is “the
product of the interaction between excitable substances and the
external exciting powers.” It is vain to look for an explanation of life
if we do not assume that its emergence “is grounded on a principle
to which must be ascribe a certain degree of independence from
external influences, of self-determination to effectiveness, an
analogous of spontaneity” (Treviranus, 1814, 626). This principle is
relevant with respect to the procreation of species, which was hard
to explain for most biological systems. The unexplainable aspect
lies in the fact that this phenomenon displays teleological features
which appear irreducible to a mechanistic framework. Plants that
vegetate under unfavorable conditions rush in producing flowers
and fruits before they pass away, so that from their seed germinates
again a healthy progeny. Vegetation behaves as if it was the product
of a spiritual principle: “precisely this similarity between the action
of a spiritual being and the effects of the vital principle points to
some kind of spontaneity of the latter.” Monsters are another class
of phenomena in which this effect of life is expressed. The internal
parts of these bodies are “purposefully organized only as much as
the degree of external deformity allows, in all of them is expressed
a tendency of the formative drive (Bildungstrieb) to produce a
possibly complete organism also under unfavorable conditions”
(Treviranus, 1814, 627). These phenomena involve both the onto-
genesis of single organisms and “the formative levels (Bildungs-
stufen) climbed by living nature as a whole.” The kind, as the
individual, has its periods of formation, bloom and death, “the
whole as the single is in a state of eternal transformation”
(Treviranus, 1814, 628). Treviranus maintains that these trans-
formations cannot be reduced to a mere modification of the effect
exerted by cosmical influences and must rather have their ground
in the laws that regulate life: “the vital force (Lebenskraft) of every
individual, inasmuch it expresses itself as formative force (Bil-
dungskraft) is the outflow of the common fundamental force
(Grundkraft) that, as the light broken with a prism, splits itself into
countless rays and produces the plurality of kinds and individuals
of the kingdom of living organisms” (Treviranus, 1814, 628).
The idea that all vital forces are branches of the same funda-
mental force and that this force is a general excitability of an or-
ganism in relation to the external environment is first formulated in
Schelling’s Erster Entwurf. Here excitability is considered as the
general principle from which all other forces could be deduced.
Formative drive, irritability and sensibility were conceived of as
further determinations of this fundamental force. In the Biologie
Treviranus maintains that the vital principle (Lebesprinzip) of every
organism depends from a communal fundamental force (Grungk-
raft). For this reason “every living being takes part to the modifi-
cation of the original source of life and thus living nature displays
phenomena whose cause lies way higher than the effect of me-
chanical or chemical powers” (Treviranus, 1814, 629). Teleology is
envisaged by Treviranus as the result of a higher level of organiza-
tion than mechanism and chemical affinity. Accordingly, the
formative principle (Bildungsprincip) of living bodies is ascribed a
certain degree of independence from external influences. Trevir-
anus maintains that this peculiar independence and spontaneity
had been previously defined in different ways. In ancient times
perspicacious thinkers assumed a purposeful principle independent
from external stimuli to be the origin of life. Such principle was the
world soul (Weltseele) of the platonists, Cudworth’s plastic nature,
or Blumenbach’s Bildungstrieb. Chemical principles can explain the
elements of which living bodies are composed. One can also inquire
“all the traces of electricity, magnetism and all further physical
forces in the living body and pursue them as far as possible. The
result of this investigations will always be that those agents in the
animal and vegetable organism are as much active as outside of it,”
but that “the actual secret of living nature will not be disclosed.” The
nourishment of living bodies is in fact regulated by specific laws of
the formative drive that cannot be reduced to the laws of physics or
chemistry (Treviranus, 1814, 631). The realm of living organisms is
different from those of mechanism and chemical affinities and re-
quires a reference to teleological principles.
There are two main tenets behind these passages. The first is
that all vital forces (sensibility, irritability, reproductive force) are
branches of the same fundamental force. This fundamental force is
the excitability of a living organism in relation to the external
environment. The second is that this relation is characterized by a
certain degree of spontaneity which is peculiar to living beings. The
latter must be understood as a higher level of organization with
regard to mechanism and chemical affinity and can be explained
only in reference to the principle of purposiveness. Both of these
tenets are drawn from Schelling’s Naturphilosophie.
Lenoir considers Treviranus to be the most paradigmatic reali-
zation of the Göttingen program. With the Biologie the “transcen-
dental biology” of the Göttingen School had concluded its
formative period. According to Lenoir, Treviranus succeeded in
pulling together various aspects of the program that had been
under intense discussion beginning in 1750 with Haller’s trans-
lation of Buffon’s Histoire naturelle. Treviranus synthesizes these
conceptual elements into what he describes as a dynamic theory of
organic nature, which he attempts to ground in an encyclopedic
overview of biological research from mid-eighteenth century on-
ward. Lenoir does not disregard the relationship between this
conceptual framework and Naturphilosophie. Through careful
reading of Schelling’s early philosophy of nature in conjunction
with the critical edition of his correspondence he deems it possible
to show that while in Leipzig, in the period during which he
devoted himself almost exclusively to acquiring background in
natural science, Schelling concentrated on the works of the Göt-
tingen School, particularly Lichtenberg, Blumenbach and Kiel-
meyer. Through direct personal contact with Pfaff and Eschenmaier,
Schelling gained also an in depth knowledge of Kielmeyer’s Physik
der Tierreichs. In addition to these personal debts Lenoir identifies
other intellectual ties between transcendental Naturphilosophie
and speculative theories of nature. The concepts of Einheit, Stu-
fenfolge, Polarität, Metamorphose, Urtyp and Analogie have been
described as distinguishing elements of this approach to nature.
The notion of ideal types was central to the Göttingen thought on
comparative anatomy. Similarly, in the work of Kielmeyer the dy-
namic interaction of vital forces bears strong similarity to Schel-
ling’s notion of polarity, as well as the use made of that notion in
the work of Nees von Essenbeck and Oken. Goethe’s notion of
metamorphosis is closely parallel to the modification of an original
ground plan developed by the Göttingen biologists (Lenoir, 1981,
192e193). Surprisingly enough, after stressing the proximity be-
tween vital-materialism and Naturphilosophie Lenoir maintains
that beside the strong similarities in key concepts of these ap-
proaches, there are still major differences in both their interpre-
tation and significance. Most importantly, vital-materialism
worked hard at remaining consistent with Kant’s philosophy of
organic nature, while Naturphilosophie worked towards its over-
coming. This distinction is historically unattested and unsupported
by textual evidence.
 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20
4. Conclusions
Lenoir has argued that Kielmeyer’s project of a Physik der Tier-
reichs is developed in accordance with the guidelines of the Kante
Blumenbach vital-materialist program and in opposition to Natur-
philosophie. A letter Kielmeyer wrote to Cuvier (Kielmeyer, 1837)
seems to endorse this claim. Cuvier had been Kielmeyer’s student
in Stuttgart and kept epistolary contacts with him throughout his
Parisian carrier. In 1807 he wrote to his former teacher asking of his
opinion on the recent development of German philosophy of na-
ture. Kielmeyer’s answer is long and detailed. He criticizes Natur-
philosophie, argues against the endeavor of “deducing the
multiplicity of nature from our own I” and positions himself in
proximity to Kant’s Metaphysische Anfangsgründe. He also says,
however, that “through the recent philosophical systems it has
become more usual to consider nature in its entirety as a living
organism in all its aspects, and to conceive of single organization as
individualized exponents of the great nature.” More than twenty
years later, in the course of a lecture held at the Muséum in 1832,
Cuvier expresses his own position on the matter, defining Kiel-
meyer as “the father of Naturphilosophie who did not want to
recognize his daughter” (Bach, 1994). This statement provides a
balanced understanding of the issue at hand: Naturphilosophie
draws indeed some metaphysical conclusions that are unaccept-
able for the empirical scientist, and in this sense Kielmeyer had
right reasons to refuse its assumptions; on the other hand, how-
ever, Schelling’s philosophy of nature develops the framework laid
down in Kielmeyer’s Rede and formulates conceptual models later
employed in Treviranus’ Biologie.
I agree with Zammito that the vital-materialism as it was
developed at Göttingen is not quite the Kantian “transcendental
philosophy of nature” that Lenoir wants it to have been. On the
contrary, we find the Göttingen School far closer to the Natur-
philosophen than Lenoir would like. Instead of viewing the close-
ness of the Göttingen School to Naturphilosophie as contamination,
we might view it as historical evidence that something essential to
the character of biology as a special science was at stake (Zammito,
2012, 130). I hope to have shown that what was at stake was the
determination of “organization” as the specific property of living
nature and that this process implied a shift from a regulative to a
constitutive understanding of teleology. Biology emerged as the
general science dealing with the teleological laws that regulate the
organization of living nature as a whole. In this scenario, Kant’s
views on teleology constituted more of an hindrance than an aid.
On the other hand, if the Göttingen vitalism cannot be excluded
from this process, neither can Naturphilosophie.
References
Bach, T. (1994). “Kielmeyer als »Vater der Naturphilosophie« ? Anmerkungen zu
seiner Rezeption in deutschen Idealismus”. In K. T. Kanz (Ed.), Philosophie des
Organischen in der Goethezeit: Studien zu Werk und Wirkung des Naturforschers
Carl Friedrich Kielmeyer (1765e1844) (pp. 232e251). Stuttgart: Steiner.
Bach, T. (2001). Biologie und Philosophie bei C.F. Kielmeyer und F.W.J. Schelling.
Stuttgart, Bad Cannstatt: Frommann-holzboog.
Blumenbach, J. F. (1787). Institutiones physiologicae. Göttingen: Dietrich.
Canguilhem, G. (1962). Du développement à l’évolution au XIXe siècle. Paris: Puf.
Cheung, T. (2006). From the organism of a body to the body of an organism:
Occurrence and meaning of the word ‘organism’ from the seventeenth to
the eighteenth centuries. The British Journal for the History of Science, 39(3),
319e339.
Coleman, W. (1973). Limits of the recapitulation theory. Carl Friedrich Kielmeyer’s
critique of the presumed parallelism of earth history, ontogeny, and the present
order of organisms. Isis, 63(3), 341e350.
Coleman, W. (1977). Biology in the nineteenth century. Problems of form, function, and
transformation. Cambridge: Cambridge University Press.
Duchesneau, F. (1982). La physiologie des Lumières. Empirisme, modèles et théories. La
Hague, Boston, London: Nijhoff.
Dupont, J. C. (2007). Pre-Kantian revival of epigenesis: Caspar Friedrich Wolff’s De
Formatione Intestinorum (1768e69). In P. Huneman (Ed.), North American Kant
19
Society studies in philosophy: 8. Understanding purpose. Kant and the philosophy
of biology (pp. 37e50). Rochester: University of Rochester Press.
Fischer, M. (2007). Kant’s explanatory natural history: Generation and classification
of organisms in Kant’s natural philosophy. In P. Huneman (Ed.), North American
Kant Society studies in philosophy: 8. Understanding purpose. Kant and the phi-
losophy of biology (pp. 101e122). Rochester: University of Rochester Press.
Ginsborg, H. (2001). Kant on understanding organisms as natural purposes. In
E. Watkins (Ed.), Kant and the sciences (pp. 231e258). Oxford: Oxford University
Press.
Ginsborg, H. (2006). Kant’s biological teleology and its philosophical significance. In
G. Bird (Ed.), The Blackwell companion to Kant (pp. 455e469). Oxford: Blackwell
Publishing.
Goy, I., & Watkins, E. (Eds.). (2014). Kant’s theory of biology. Berlin, New York: De
Gruyter.
Guyer, P. (2005). Purpose in nature: What is living, and what is dead in Kant’s
teleology? In P. Guyer (Ed.), Kant’s system of nature and freedom (pp. 343e372)
Oxford: Oxford University Press.
Huneman, P. (2007). Reflexive judgment and Wolffian embriology: Kant’s shift
between the first and the third critique. In P. Huneman (Ed.), North American
Kant Society studies in philosophy: 8. Understanding purpose. Kant and the phi-
losophy of biology (pp. 75e100). Rochester: University of Rochester Press.
Huneman, P. (2012). Métaphysique et biologie. Kant et la constitution du concept
d’organisme. Paris: Kimé.
Illetterati, L. (2008). Being for. Purposes and functions in artifacts and living beings.
In L. Illetterati, & F. Michelini (Eds.), Purposiveness: Teleology between nature and
mind (pp. 135e161). Frankfurt: Ontos.
Illetterati, L. (2010). Biologia. In S. Besoli, C. La Rocca, & Martinelli (a cura di) (Eds.),
L’universo kantiano. Filosofia, scienze, sapere (pp. 53e96). Macerata: Quodlibet.
Kielmeyer, K. F. (1938). Natur und Kraft. Gesammelte Schriften (von F.-H. Holler unter
Mitwirkung von J. Schuster, nach den Handschriften zum erstenmal veröffen-
tlicht). Berlin: Keiper
Lenoir, T. (1978). Generational factors in the origin of Romantische Natur-
philosophie. Journal of the History of Biology, 11(1), 57e108.
Lenoir, T. (1980). Kant Blumenbach and the vital materialism in German biology.
Isis, 70, 77e108.
Lenoir, T. (1981). The Göttingen School and the development of transcendental
Naturphilosophie in the Romantic Era. Studies in the History of Biology, 5, 111e205.
Lenoir, T. (1982). The strategy of life: Teleology and mechanics in nineteenth-century
German biology. Dordrecht: Reidel.
Löw, R. (1980). Philosophie des Lebendigen. Der Begriff des Organischen bei Kant. Sein
Grund und seine Aktualität. Frankfurt: Suhrkamp.
Marcucci, S. (1972). Aspetti epistemologici della finalità in Kant. Firenze: Le Monnier.
McLaughlin, P. (1989). Kants Kritik der teleologischen Urteilskraft. Bonn: Bouvier.
McLaughlin, P. (1990). Kant’s critique of teleology in biological explanation: Antinomy
and teleology. New York: Mellem Lewinston.
Monti, M. T. (1990). Congettura ed Esperienza nella Fisiologia di Haller: La Riforma
dell’Anatomia Animata e il Sistema della Generazione. Firenze: Olschki.
Philonenko, A. (1982). Kant et la philosophie biologique. In L’Heritage de Kant.
Melanges philosophiques offerts au P. Marcel Regnier (pp. 63e79). Paris:
Beauchesne.
Reill, H. P. (2005). Vitalizing nature in the enlightenment. Berkeley: California Uni-
versity Press.
Richards, R. J. (1992). The meaning of evolution. The morphological construction and
ideological reconstruction of Darwin’s theory. Chicago: University of Chicago
Press.
Richards, R. J. (2000). Kant and Blumenbach on the Bildungstrieb: A historical
misunderstanding. Studies in History and Philosophy of Biological and Biomedical
Sciences, 31(1), 11e32.
Richards, R. J. (2002). The romantic conception of life. Science and philosophy in the
Age of Goethe. Chicago: University of Chicago Press.
Roe, S. A. (1981). Matter, life, generation. Eighteenth-century embryology and the
Haller-Wolff debate. Cambridge: Cambridge University Press.
Schelling, F. W. J. (2000). Von der Weltseele. Eine Hypothese der höhern Physik zur
Erklärung des allgemeinen Organismus (Von J. Jantzen unter mitwirkung von T.
Kisser). Stuttgart: Frommann-Holzboog
Schelling, F. W. J. (2001). Erster Entwurf eines Systems der Naturphilosophie (von W.G.
Jacobs und P. Ziche). Stuttgart: Frommann-Holzboog
Schmitt, S. (2006). Les forces vitales et leur distribution dans la nature. Un essai de
“systématique physiologique” (Texte de C. F. Kielmeyer (1765e1844), H. F. Link
(1767e1851) et L. Oken (1779e1851) rassemblés, introduits et annotés par S.
Schmitt). Turnhout: Brepols
Schmitt, S. (2007). Succession of functions and classification in post-Kantian
Naturphilosophie around 1800. In P. Huneman (Ed.), North American Kant So-
ciety studies in philosophy: 8. Understanding purpose. Kant and the philosophy of
biology (pp. 123e136). Rochester: University of Rochester Press.
Sloan, P. (1979). Buffon, German biology and the historical interpretation of
biological species. The British Journal for the History of Science, 12(2), 109e
153.
Sloan, P. (2006). Kant on the history of nature: The ambiguous heritage of the
critical philosophy for natural history. Studies in the History and Philosophy of
Biological and Biomedical Sciences, 37(4), 627e648.
Steinke, H. (2005). Irritating experiments: Haller’s concept and the European contro-
versy on irritability and sensibility, 1750e90. Amsterdam, New York: Rudopi.
Treviranus, G. R. (1802). Biologie oder Philosophie der lebenden Natur für Naturforcher
und Ärzte, Band I. Göttingen: Röwer.
20 A. Gambarotto / Studies in History and Philosophy of Biological and Biomedical Sciences 48 (2014) 12e20
Treviranus, G. R. (1814). Biologie, oder Philosophie der lebenden Natur für Natur-
forcher und Ärzte, Band IV. Göttingen: Röwer.
Wolfe, C. T. (2014). On Newtonian analogies in eighteenth-century life science:
Vitalism and provisionally inexplicable explicative devices. In Newton and
empiricism (pp. 223e261). Oxford: Oxford University Press.
Wolff, C. F. (1759). Theoria generationis. Halle: Hendel.
Zammito, J. (1992). The genesis of Kant’s critique of judgment. Chicago: University of
Chicago Press.
Zammito, J. (2007). Kant’s persistent ambivalence toward epigenesis 1764e90. In
P. Huneman (Ed.), North American Kant Society studies in philosophy: 8.
Understanding purpose. Kant and the philosophy of biology (pp. 51e74).
Rochester: University of Rochester Press.
Zammito, J. (2012). The Lonoir thesis revisited: Blumenbach and Kant. Studies in
History and Philosophy of Science, Part C: Studies in History and Philosophy of
Biological and Biomedical Sciences, 43(1), 120e132.
Zuckert, R. (2007). Kant on beauty and biology. An interpretation of the critique of
judgement. Cambridge: Cambridge University Press.
Zumbach, C. (1984). The transcendent science. Kant’s conception of biological meth-
odology. The Hague: Nijhoff.