Progress in Biophysics and Molecular Biology 110 (2012) 137e149

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Progress in Biophysics and Molecular Biology

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Original research

Pure variation and organic stratificationq

Jérôme Rosanvallon
Université Paris 7 Diderot, France

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

Article history: The fundamental problem posed by Darwin distinguishes his theory from any transformism of the past
Available online 31 July 2012 as well as any evolutionism to come: since variation is inherent to the living, it is a question of explaining,
not at all why the living varies, but instead why the living does not vary in all directions to the point of
Keywords: constituting a continuum of forms varying ad infinitum. What limits and stabilizes this intrinsically
Variation unlimited variation, allowing certain forms to subsist and multiply to the detriment of others, is natural
Stratification
selection. This double principle of intrinsic variation/extrinsic selection constitutes a vector for the
Cosmogony
unification of reality that underlies Jean-Jacques Kupiec’s ontophylogenesis as well as Deleuze and
Symbiogenesis
Ontophylogenesis
Guattari’s global philosophy of Nature.
Body without organs Therefore, everything would potentially tend to incessantly vary. The work of Kupiec and others iden-
tifies an intrinsic random variation within ontogenesis itself. For Deleuze and Guattari, it is nothing but
the figure, already selected by the organic stratum, of a more fundamental or pure variation. But, in fact,
nothing really varies incessantly: everything undergoes a selective pressure according to which nothing
subsists as such except what manages to endure through invariance (physical stratum) or reproduction
(organic stratum). Thus, organic stratification only retains from variation what ensures and augments
this reproduction. In this sense, every organism stratifies, i.e. submits to its imperative of subsistence and
reproduction, a body without organs that varies in itself and always tends to escape the organism, for
better (intensifications of life) or worse (cancerous pathologies).
Ó 2012 Elsevier Ltd. All rights reserved.

1. Darwin: intrinsic variation and extrinsic selection
1.1. The two meanings of variation in the Origin of Species
In what sense can we really speak of a Darwinian revolution?
What is the comparative advantage of the Darwinian theory, which
has today still allowed it to extend and impose itself to the detri-
ment of all other theories, including those affiliated with it? Darwin
does not simply mean to break with creationism and to affirm
definitively that “each species had not been independently created,
but had descended, like varieties, from other species1”. More subtly
and with a certain prescience, he tends to break with theories of
transformism, whether past or contemporary (Lamarck, Owen,
etc.), while also breaking with their future evolutionist variants
that nevertheless ally with him (Th. Huxley, Spencer, etc.). In
a sense, all these theories are looking to explain the variation of the
living and are therefore led to suppose that the living tends to adapt
itself to its milieu, to complexify itself, to individualize itself, etc.,
supposing in this way some tendencies within individuals, species
or life itself that would constitute determined directions of varia-
tion. But Darwin simply posits axiomatically or in principle the fact
that there is, within both domesticated and natural living beings,
variation.2 Just as the order of the presentation of the Origin of
Species (OS) seems to suggest, every theory of the living must take
this fact as its point of departure and not its destination: every theory
must found itself entirely upon this fact rather than seeking to
found it, i.e. to completely explain it. Although Darwin did not
necessarily grasp its overall revolutionary capacity, this principle’s
position has been fully brought to light by one of his most faithful
correspondents, the botanist Joseph Dalton Hooker, who correctly
reproaches him for not having emphasized it in a more compelling
way in the OS and afterwards for too often neglecting e in his
ulterior hesitations e “to dwell on the facts of infinite incessant
variation3”.
Thus all the stakes of natural selection are deduced from this
primary fundamental fact: to explain that everything within the

q Translated from the French by Taylor Adkins. 2

Ibid., chapters 1 and 2.
3
E-mail address: jrosanva@orange.fr. Letter from Hooker to Darwin 26 November 1862, Correspondence of Charles
1
On the Origin of Species [OS], 1st ed., 1859; “Introduction”, p. 3. Darwin [CCD], X, p. 570.

0079-6107/$ e see front matter Ó 2012 Elsevier Ltd. All rights reserved.
http://dx.doi.org/10.1016/j.pbiomolbio.2012.06.002
138 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
living does not vary incessantly, that there are invariances e there is
stability, which is itself provisional, and there is reproduction of
identity, which is itself partial, i.e. ultimately that there are groups
of living beings that are distinct from and cross-fertile with others
and are thus reckoned to constitute a certain number of species e
second constitutive fact of the living and first observation. In this
sense, natural selection would designate nothing but the existence
of a condition of non-variation, the existence of a constraint that
limits this potentially infinite and incessant variation by conserving
only the favourable variations, which are then led to impose them-
selves to the detriment of others, thus laying the foundations for
specific divergences. We understand in a similar way that Hooker
could consider that a rigorous, axiomatic and pedagogical presen-
tation of the Darwinian theory sets out to “disentangle the two
phenomena of variation and Natural selection4”, the first and
always disguised fact of incessant variation and the second and
always partial fact of reproductive invariance. But the same text of
the OS seems to doubly resist this disentanglement that Darwin
never fully unravels. On the one hand, through several reiterations,
chapter four presents natural selection as having the capacity to
produce variations, to “act on and modify organic beings5”, which is
what Hooker explicitly bemoans: “Natural Selection is supposed to
make the varieties as well as to fix them”, whereas it is nothing but
the stabilizer or “fixer of these” varieties into species.6 On the other
hand, chapter five seems to claim to partially explain variation by
exposing its “laws”. Darwin’s difficulties in completely leaving
behind this pattern advocated by his predecessors and a number of
his successors, who believed it was necessary to explain the vari-
ations of the living and not its invariances, are further confirmed by
the way in which he responds to his interlocutor: “I wish I had done
what you suggest, started on the fundamental principle of variation
being an innate principle; and afterwards made a few remarks,
showing that hereafter perhaps this principle would be explicable7”.
The resistances Darwin seems to have do not at all contradict
the axiomatic framework redefined by Hooker but simply require
for new distinctions to operate within it. Concerning the “produc-
tion” of variation through natural selection, it is a question of an
abbreviated formula that does not support any ambiguity if one
takes care to distinguish, according to the celebrated “queer
diagram” presented in chapter four, between variation envisioned
“horizontally”, the spontaneous branching out with each generation,
and variation envisioned “vertically”, the progressive modification of
the same lineage. In the latter case, variation is nothing but that of
a subject, i.e. of a certain species taken as an arbitrary point of
departure: it simply measures the divergence between this point of
departure, “common descent”, and a destination that is just as
arbitrary, its so-called “modified” descent; this divergence in the
end is actually an effect induced by natural selection, which tends
to eliminate all the less advantageous, intermediary variations.
Nevertheless, in the first case variation is subjectless or has no
subject other than the living itself: it returns to a pure variability of
the living, “an inherent tendency to vary8” with each generation;
this variation, which remains without determined or perhaps
determinable cause, is instead intrinsic in the sense that natural
selection only intervenes in it secondarily, and thus extrinsically
(even if e we shall return to this e it also itself tends to conserve
and amplify this intrinsic variability of the living). If he often passes
from one to the other unexpectedly, Darwin acknowledges the
4
Letter from Hooker to Bates 2 February 1862, CCD, X, p. 128.
5
OS, 1st ed., 1859, IV, p. 86.
6 9
Letter from Hooker to Bates 2 February 1862, CCD, X, p. 129.
7
Letter from Darwin to Hooker 26 March 1862, CCD, X, p. 135 (my emphasis).
8 10
The variation of animals and plants under domestication, 2nd ed., 1875, p. 2.
importance of distinguishing between these two meanings of
variation, most notably when, in the final edited version of the last
chapter, he once again casts his theory as that of “descent with
modification through variation and natural selection9”.
It should be noted that the “laws of variation” are not the trans-
lation of some theory of variation that would attempt to explain the
primordial fact of variation by revealing its origin, reason or causes.
Concerned with the how and not the why like any other law, they only
seek to comparatively determine the “relative degrees of vari-
ability10”, i.e. the greater or lesser variability of a certain part of the
organism (according to its more or less developed character, the loss
of its usage rendering its elimination advantageous, its structural or
hereditary correlation with other parts, etc.) and of a certain pop-
ulation of living beings (according to whether it lives domestically or in
the wild, in an extensive or restrained habitat, etc.). Darwin’s question
is therefore never e for what reasons or according to which causes does
it vary? e but always e according to which differential speeds and rates
does it vary? e being given the greater or lesser selective constraint
that secondarily exerts itself upon this intrinsic variation.
1.2. The “Darwin-Hooker principle”
Having clarified this axiomatically, we can now call this prin-
ciple e according to which, on the one hand, what varies is
a primordial fact that is itself given without having to be explained
and, on the other hand, only what does not vary must be explained
as the effect of a constraint that extrinsically limits variation e
“Darwin’s principle”, or more precisely, in order to do justice to his
most prescient reader and interlocutor on this point, the “Darwin-
Hooker principle”. But could this principle be restricted to the
sphere of the living, since it is not simply valid within what we call
the “organic stratum”? Does it not necessarily call for a meta-
physical systematization that would take variation alone as the
point of departure and simply give itself the task of explaining on its
basis everything that does not vary, namely biological but also
cosmological and physico-chemical or even anthropological and
socio-historical invariants? The entire program of naturalist
metaphysics elaborated for a span of about twenty years by Deleuze
and Guattari could be taken back up in this way (from Anti-Oedipus
in 1972 up to What Is Philosophy? in 1991): this program implicitly
relies upon the “Darwin-Hooker principle”, which is the same as
reversing Leibniz’s fundamental ontological question by no longer
examining why something happens rather than nothing, but on the
contrary why sometimes nothing happens, i.e. why everything does
not incessantly vary. This fundamental reversal thus calls for
a radical generation of all forms of invariance, starting with those
that constitute the framework of the physical world (space-time,
constants, matter, things, etc.), which then cannot be an irreducible
given of nature but simply a secondary effect resulting from
a constraint exerted upon a primary variation.
The “Darwin-Hooker principle” seems to constitute a new
formulation, which is potentially valid for all the sciences and their
studied domains of reality, of the “principle of reason”: in effect,
this formulation is substituted point by point for the preceding
formulation that masterfully reigned over modern science and
philosophy and completely reverses it. This preceding formulation
can identically be called the “Newton-Galileo principle” and can be
defined in this way: every system tends to conserve its state and
only changes the latter if an external force is exerted upon it. This
fundamental principle of modern physics actually at the same time
OS, 6th ed., 1872, XV, p. 404 (my emphasis: “through variation” was only added
in the 6th and final edition).
OS, 1st ed., 1859, V, p. 151.
 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
underlies the kinematics of Galileo’s supposed inertial movements
and the dynamics of Newton’s supposed external forces: it is not
only translated by the principle of inertia but also by the principle of
relativity (which posits the equivalence of “immobile”, inertial and
e with its Einsteinian generalization e accelerated systems of
reference) and the various laws of conservation (energy, quantity of
movement, angular momentum, electric charge, etc.). Does it not
almost immediately become the object itself of a rigorous meta-
physical systematization through Spinoza? The entire Ethics in
a sense finds a logic of the conatus, i.e. the inertial perseverance of
every (physical, organic and human) thing in being, a perseverance
that only varies (augmenting or diminishing its capacity of being)
and only stops functioning due to external encounters alone. In this
sense, we can more easily grasp the fundamental reversal operated
by the “Darwin-Hooker principle”, which on the contrary affirms
that every system tends to vary in itself and only conserves its state
in accordance with an external constraint. As foundational for
modern biology, this principle subtends both phylogenesis e
following the Darwinian revolution e and ontogenesis e following
the extension of this revolution initiated by Kupiec: it combines
intrinsic variation (whose nature, as we shall see, is fundamentally
random) and extrinsic selection (which, in the case of the living,
does not with each generation restrain the variation and varieties it
generates without tending to conserve this variability that is
inherent11). It therefore remains for us to show how Deleuze and
Guattari’s philosophy constitutes, with the double idea of pure
variation and stratification, the most ambitious metaphysical
systematization of what remains after a century and a half a new
and revolutionary formulation of the “principle of reason”.
2. Everything potentially tends to vary incessantly: random
variation and pure variation
2.1. Decodings and deterritorializations: Neo-Darwinism according
to Deleuze and Guattari
On the Origin of Species therefore does not contain a theory of
variation but a theory of selection that limits variation, whatever the
scale, causes and nature of the latter might be. Darwinian theory in
this sense assigns a place to variation, taking it as a primary matter
without needing to determine it except by a variability upstream
that produces it and a variety downstream that it produces.
Darwinian theory must singularly determine, on one side, degrees
of variability, which are envisioned as relative to one another in
accordance with conditions of existence and, on the other, varieties
of population, which are envisioned as an array of nascent species.
As for the incessant variations produced in this way, they cannot
receive any supplementary determination e the theory does not
require it: this is not so merely by default, because, as we said, on
the contrary this is what focuses its entire force. Based on Darwin’s
own quite classic judgement, it therefore does not matter much
that the seemingly random character of all these variations is
nothing but the simple effect of our ignorance12: his theory will
constitute no less than the guiding thread for all those whose point
of departure is an intrinsic random variation upon which an
11
A recent reformulation of this principle e in contrast with that of Newton-
Galileo e has also been proposed by Brandon and McShea (2010). But is not
diversification a secondary consequence of intrinsic variation, and “complex-
ification” a useless evolutionary residue?
12
“I have hitherto sometimes spoken as if variations e so common and multiform
in organic beings under domestication, and to a lesser degree in those in a state of
nature e had been due to chance. This, of course, is a wholly incorrect expression,
but it serves to acknowledge plainly our ignorance of the cause of each particular
variation”, OS, 1st ed., 1859, V, p. 106.
139
extrinsic pressure of selection can be exerted, whatever the source
of variety and type of “population” concerned might be, whether
within the organic strata or beyond. Thus we have been able to
speak of molecular, cellular, immunitary and neuronal, as well as
social, economic and even mineral or cosmological Darwinism.13
The “synthetic theory of evolution” labelled as “Neo-
Darwinism” is obviously the first among all the others that has
given to the double Darwinian principle an irrefutable support, that
of the genetics of populations: on the one hand, genetic mutations
constitute the single a priori material support for transmissible and
thus selectable variations and, on the other hand, selection is
carried out a posteriori by the milieu, which eliminates unfav-
ourable mutations and conserves favourable ones insofar as the
latter allow the organisms to which they belong to acquire new
resources or utilize available resources better than other organisms.
The synthetic theory constitutes a primary authentic re-foundation
for Darwinism, insofar as it accentuates the divergence and estab-
lishes a single direction between the genotypic source of variability
and its selected phenotypic effects, thus confirming that variations
depend not on some internal finality of development or external
finality of evolution e whatever it might be e but solely on what
arrives in a stochastic fashion in the genetic code, which is exposed to
the constant bombardment of the milieu (oxidations, radiations)
and above all constrained to ongoing replications and recombina-
tions (cellular division and sexual reproduction). This fact alone,
however, does not prevent us from envisioning the replication of
the DNA complementary double helix as the most reliable process
imaginable, just as much as it does not prevent us from conse-
quently assimilating all these random variations with “accidents”,
“errors”, simple “noise” e which are all certainly decisive, since all
varieties and, in the end, all specific divergences result from them.
Such an interpretation illustrates the current attitude of the biol-
ogists who, lacking the ability to account for a source of chance
deterministically, at least tend to cast it in a simply perturbative
role. But a properly Darwinian perspective always implies taking
not invariance but variation as the norm and thus implies never
giving up on asking, including within DNA, not “how do variations
appear, but the other way around, how is it that they do not appear
more often!14”. In truth, a genome owes its stability to various
mechanisms of control and correction (notably the DNA mismatch
repair system15), which are the precocious products of selection
and which have been identified with multiple variants in (almost)
all uni- or multicellular prokaryotic and eukaryotic organisms.
Despite this reinforced fidelity of genomic replication, the
stability of genotypic forms on the geological scale is as provisional
as the species that express them phenotypically. These forms
therefore seem like “so many islands consolidated here and there in
the ocean of becoming16”, a scattering of little islands emerging in
time through selection and necessarily drifting within a perpetual
genetic whirlpool. Describing the organic stratum’s unity of
composition and logic of constitution in the third plateau of A
Thousand Plateaus titled “The Geology of Morals”, Deleuze and
Guattari specifically point out the fact that “a code is inseparable
from a process of decoding that is inherent to it”, a process of
decoding signifying for them neither the transduction nor the
translation of the genetic code alone on the scale of the individual,
13
For a complete panorama of all these Darwinisms, see Heams, 2011b; the work
of Gerald Edelman in neurobiology (see specifically Edelman, 1987) and that of Lee
Smolin in cosmology (see specifically Smolin, 1999) should also be included. We
shall take up the question of cosmological selection in our third part.
14
Thomas Heams, 2011a, p. 54e55.
15
See, historically, Meselson and Wildenberg, 1975, and, for a recent review,
Galles et al., 2009.
16
Bergson, The Creative Mind: An Introduction to Metaphysics, p. 64.
140 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
but the transformations and drifts of genomes in their entirety on
the scale of populations.
No genetics without ‘genetic drift’. The modern theory of
mutations has clearly demonstrated that a code, which neces-
sarily relates to population, has an essential margin of decoding:
not only does every code have supplements capable of free
variation, but a single segment may be copied twice, the second
left free for variation. In addition, fragments of code may be
transferred from the cells of one species to those of another [.]
by viruses or through other procedures.17
Deleuze and Guattari here offer a short and somewhat faithful,
general idea of the multiple mechanisms of stochastic genetic
variation discovered with the synthetic theory. These mechanisms
can be divided into two basic types: mutations that create variants
of code (alleles) and the recombinations that can combine favour-
able variants. But everything happens as if the bacterial prokaryotes
and the domain of uni- or multicellular eukaryotes form nothing
but two distinct exploratory paths of these two types of variation or
margins of decoding. On one side, prokaryotes slightly experience
more elevated and above all variable rates of mutation (SOS
response which thwarts the effects of the mismatch repair system
in critical moments18), but their mutations are essentially punctual
(single base substitutions) and quite rarely favourable (lesser size of
genomes). On the other side, the mutations of eukaryotes are
slower and more constant, but they essentially are produced by the
duplication of longer or shorter fragments of DNA, most notably
genes, including all the chromosomes during the course of repro-
duction (polyploidy); these duplications are such that they ballast
the genomes of an increasing non-coding part and therefore
augment the probability of neutral or favourable variations that
have been able to open the most diversified path, for then we shall
witness the “deterritorializable” path of multicellularity. From that
moment on, where recombinations are made simply in bacteria
through the horizontal transfer of one species to another (via plas-
mids, circular fragments of DNA or viruses), eukaryotes have
invented another path of genetic recombination, the sexual path,
which assures a greater variability and thus a greater adaptability
within the same species but also induces an inevitable genetic drift,
i.e. a random and non-optimizing selection of these variations.
Must we conclude that genotypic variation is the only source of
random variation of the living? It is important never to forget that this
perpetual genetic shuffling wouldn’t have more value than a simple
molecular disorder if there were not carried out in parallel, at the
other end of the spectrum, a systematic selection of favoured
phenotypes and thus a constant optimization of mutations: “milieus
always act through selection on entire organisms, whose forms
depend on the codes that these milieus indirectly sanction19”. The
reproductive success of each organism therefore will depend on its
adaptation to the milieu, i.e. on its capacity to effectively utilize the
available nutritive and energetic resources compared to other
organisms. The fact that this adaptation is the mechanical outcome of
selection does not prevent it from also bringing with it an important
margin of indetermination. The external milieu of organisms,
including their “organic and inorganic conditions of life20”, in fact
itself always tends to vary, whether concomitantly with or indepen-
dent of the actions of the latter: climatic variations, the arrival of new
predators, evolution of prey, scarcity of resources, etc. All these
17
Deleuze and Guattari, A Thousand Plateaus, p. 53.
18
See the whole work of Miroslav Radman (notably Radman, 1975; Radman et al.,
1995, 2003).
19
A Thousand Plateaus, p. 52.
20
OS, 1st ed., 1859, IV, p. 84.
variations intervene in the reproductive success of this or that indi-
vidual or population, and therefore no adaptation is acquired once
and for all. Adapting in fact always returns, according to Deleuze and
Guattari, to “being deterritorialized”: “it is populations that are
deterritorialized and reterritorialized” while “they are coded and
decoded”, and if “deterritorializations and reterritorializations do not
bring about the modifications” of code, they “strictly determine their
selection21”. Ecosystems can therefore be seen as the continually
changing territorial spinoffs of this great vector of deterritorialization
that has borne all the phylogenetic lineages and specifically assured
the maximal phenotypic diversification of multicellular organisms.
To deterrorialize consists in conquering, first and above all else, new
sources of energy through the extension of types of respiration (this is
specifically the main success of Archaea e between bacteria and
eukaryotes e which can only live in the most extreme milieus), but
also and correlatively new territories through the acquisition of a partial
or total mobility (this is the great success of multicellular eukaryotes),
since it is a question of the passive and aleatory dissemination of only
the reproductive elements (case of vegetal life) or the active, but also
aleatory, displacement of the entire organism (case of animal life): “[t]
he more internal milieus an organism has on its own stratum,
assuring its autonomy and bringing it into a set of aleatory relations
with the exterior, the more deterritorialized it is22”. Thus multicel-
lular organisms attain this maximal point of deterritorialization/
reterritorialization by becoming capable, solely based on chance
disseminations and migrations, of traversing, i.e. of occupying and/or
travelling throughout all the territories of the Earth favourable to life.
2.2. The third source of intrinsic variation: ontophylogenetic
unification
Between the margin of decoding inherent to the code and the
vector of deterritorialization inherent to organisms, is there not,
however, any other source of random variation within the organic
stratum? Wouldn’t variation simply intervene upstream from the
replicated codes or downstream from the produced forms?
Wouldn’t a phenotypic form be the programmed product e i.e.
simultaneously determined and finalized e of a given genotype,
through the intermediary of the proteins that the latter codes and
that compose and allow all the cells making up the organism to
function? Ontogenesis would then be the only stage of the living
that would result from a supposed genetic program and not from
the double Darwinian principle of intrinsic variation/extrinsic
selection. But, as Jean-Jacques Kupiec has already insisted, sources
of random variation have been discovered on every stage of onto-
genesis23: above all else, genes are expressed in a stochastic way
within each cell according to the frequency of encounters with
regulative proteins, the topological and temporal dynamics of
chromatin, etc.; moreover, molecular and proteinic interactions
form a veritable non-specific rhizomatic network24 where each
molecule can potentially participate in any transformation what-
soever; consequently, each cell follows paths of differentiation,
which are themselves stochastic, without obeying any inductive
molecular signal; and ultimately, all the cells of multicellular
organisms conserve a stochastic possibility of autonomous varia-
tion within the organism in its growth, maintenance and
21
A Thousand Plateaus, p. 54.
22
Ibid., p. 53e54.
23
Kupiec, 2008, pp. 47e66.
24
Rhizome is a concept that Deleuze and Guattari take up from botany mainly to
describe multiplicities that are not reducible to unity and more specifically to
describe non-arborescent networks, see “Introduction: Rhizome” in: A Thousand
Plateaus, pp. 3e25.
 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
reproduction in which they contribute to the unique fact that its
internal milieu itself contributes to their subsistence. Another fact
must be added to these variational facts that are irreducible to
simple “noise”: the apparent reproducible stability of the products
of ontogenesis. Kupiec concludes from this that only the Darwinian
paradigm can account for this double aspect and that it is thus
applied to both ontogenesis and phylogenesis, including all the
stages and scales of the living.
Before analysing the concrete consequences of this ontophylo-
genetic unification and showing that the logic of the living’s
stabilization that it envisions unites with the logic of organic
stratification described by Deleuze and Guattari, let us attempt to
further define the nature of random variation beyond and
throughout all the figures that it takes within the organic stratum.
As we mentioned, the “Darwin-Hooker principle” primarily affirms
the intrinsic nature of variation. But it is actually upon this intrinsic
nature that its random character depends, and not the other way
around. In effect, intrinsic variation must not simply be understood
in a negative sense, i.e. as everything that is not determined
extrinsically. The force of the Darwinian paradigm is precisely to
offer a positive definition of this mode of determination: an
intrinsic variation is not in itself without determination, it is simply
determined or caused by another mechanism than that which deter-
mines or limits it; in truth it is not undetermined but doubly deter-
mined. The limiting mechanism e selection e will therefore be
called extrinsic in relation to the causing mechanism e the source of
variability. It is this disproportion between cause and effects, this
irreducible gap between two determining orders e for example, the
molecular transformations of the genetic code and the adaptation
to the external milieu that its phenotypic expression requires, or
even the adaptability with which an organism is equipped and the
effective variations of the milieu into which it is plunged e that
leads us to qualify an intrinsic variation as random and makes us
wrongly believe that it would thus neither have causes nor condi-
tions. If random variation and causal indetermination are often
confused de facto, it is because of another major confusion that
manages to graft itself onto it and sustain it. Envisioned a priori or
abstractly, the variation intrinsic to the living is necessarily
unlimited and undetermined e or the “Darwin-Hooker principle”
would be violated and external limits or internal determinations to
the variation reintroduced, tending to explain it and restrain the
explanatory power of natural selection.25 But when we concretely
envision it within its various fields of intervention (genetic shuf-
fling, ontogenetic variability, phylogenetic deterritorialization),
intrinsic variation always seems determined by conditions
(genomes, cells and territories) and limited in its possibilities e
which fully open themselves for each new drawing according to the
preceding drawing, as Deleuze and Guattari describe them starting
with Anti-Oedipus by utilizing the image of Markov chains as
“partially dependent, aleatory phenomena” or a “succession of
partial re-linkings26”.
25
Darwin thus refuses any a priori limitation of variation: “the ordinary belief that
the amount of possible variation is a strictly limited quantity is likewise a simple
assumption”, OS, 3rd ed., 1861, IV, p. 89. Hoquet (2009) has clearly revealed the
stakes of this non-limitation/indetermination in order to preserve the integrity of
the explanatory power of natural selection: “On the one hand, if variation is neither
infinite nor unlimited, this reduces the scope of natural selection: it is revealed to be
confined within certain boundaries. On the other hand, if variation is not random
but follows determined directions, this reduces the latitude of natural selection and
seems to channel it along certain directions”, pp. 183e184.
26
Deleuze and Guattari, Anti-Oedipus, p. 289 and 343 and Deleuze, Foucault, p. 71
and 96. Markov chains are used today specifically in bio-informatics in order to
model the relations between the successive symbols of the genetic code and thus to
distinguish between coding and non-coding genes, specifically in eukaryotes.
141
2.3. Random variation, contingent variation and pure variation
However, taken as a guiding thread and a theoretical imperative,
the “Darwin-Hooker principle” prohibits us from remaining with
this sole intrinsic random determination of variation and obliges us
to extract it from the figure that it takes within the living or the
organic stratum: in effect, is it not, in this case, always already
constrained and selected, thus always already conditioned and
limited, in short always already partially explainable? Doesn’t
affirming that everything that is varies in itself, before receiving
some invariance or identity e whatever it be e lead to thinking
a variation outside all constraint and condition, i.e. absolutely
outside every stratum, a variation that precedes every real experi-
ence that there can be of it, so to speak a pure variation? Quentin
Meillassoux has recently shown the necessity for every speculative
enterprise seeking the absolute or the in-itself of things to take as
its point of departure a “principle of factuality” somewhat equiva-
lent to what we have been calling the “Darwin-Hooker principle”.
This principle of Humean obedience affirms that nothing can be
necessary except the single fact that there is contingency.27 The fact of
contingency therefore plays for Meillassoux the same absolute,
inexplicable and irreducible role as the fact of variation for Darwin
and Hooker or Deleuze and Guattari. The absolute primacy
implicitly granted by Darwinian theory to variation led us in a sense
to the threshold of such a speculative conclusion by inviting us to
consider as always necessary both the fact that it varies and the fact
that it does not vary in a necessary manner (i.e. explicably through
any sort of determinism or finality). But the fact that it is simply
necessary that there be the advent of variation without necessity
could signify either that the latter is random or that it is, more
simply and more radically, contingent. A random variation is in effect
nothing but a contingent variation already submitted to conditions, i.e.
able to be assimilated into a simple drawing among a defined set e
be it infinite e of combinational and spatio-temporal possibilities
that are equivalent or even into a fortuitous linkage of causal series
or orders of independent determinants. On the contrary, a radically
contingent variation governs every condition, establishing a regime
where “[t]here is no reason for anything to be or to remain the way
it is; everything must, without reason, be able not to be and/or be
able to be other than it is28”, no longer a multi-causal regime of
equivalent possibilities and fortuitous encounters, but instead an
acausal regime of unforeseeable possibilities and incessant
ruptures, which Meillassoux calls “hyper-chaos”.
Purifying variation of all its necessarily limiting empirical
baggage under which it always already presents itself leads us
therefore to think a more radical contingency, even more uncon-
ditional than chance alone. In their last work written together
What Is Philosophy?, Deleuze and Guattari carry out a comple-
mentary purification of the character e this time intrinsic e of
variation. The undoing of every extrinsic determinant boils down
to characterizing it according to three correlated components.
Primarily, a purely intrinsic variation does not predicate something;
it is neither the variation of a subject nor a substance, for it is the
sole subject and sole substantiality,29 the variation without begin-
ning or end of which all invariances and finite things are merely
ever provisional ends. Secondly, a purely intrinsic variation does not
unfurl itself into something; it is not reducible to a movement in
space or even to a duration in time, for it subtends all space-time,
27
Quentin Meillassoux, 2006, pp. 82e111.
28
Ibid., p. 60.
29
Bergson was the first to defend this unique substantiality of variation in “The
Perception of Change”, conference lecture reprinted in: The Creative Mind, pp.
107e132.
142 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
a variation in itself whose variable number of dimensions or
degrees of freedom, including the continuity of succession and
distance of separation, are nothing but secondary consequences30.
Thirdly e and this is the central point of the Deleuzo-Guattarian
perspective e a purely intrinsic variation is nothing but a single
rhythm; it is emphasized by no other regularity than the fact that
it never stops, an incessant variation comprised of all the rhythms
or intensive degrees of speed of variation, from the infinite speed
through which the authors define “chaos”31 in their own way,
down to the slowest speeds through which “stratification” vari-
ously operates. “Stratification is like the creation of the world from
chaos, a continual, renewed creation32”: we must now fully deploy
the logic of this ongoing creation that is differentiated by limita-
tion or constraint.
3. Nothing really varies incessantly: physical subsistence and
organic reproduction
3.1. Physical stratification: constants, invariances and material
durations
Governed by the principle of unreason e which rejects any
principle of reason, whether reformulated or not e (but also
governed by an inexplicable presupposed time of successive
contingency), hyper-chaos as defined by Meillassoux forces us to
ask de facto why everything and anything whatsoever does not take
place each instant. Characterized only by the infinite speed of
variation (in which each time of succession is merely one of the
derivations, a continuous and uniform derivative), chaos as
defined by Deleuze and Guattari first forces us to ask de facto how
anything whatsoever can subsist and have a more or less stable
duration without incessantly varying in all directions and thus
disappearing immediately as such. In fact, the infinite speed of
variation, the “infinite speed of birth and disappearance” of
everything that can be, can really only give rise to nothing or the
infinite void, i.e. an absolute instability or inconsistency: “chaos
makes chaotic and undoes every consistency in the infinite33”.
The fact that at least one universe, namely ours, would be created
on the basis of this “nothing” thus signifies that a relative
consistency is provisionally subtracted from this absolute incon-
sistency. This initial means of taking on consistency within chaos
that defines the object of physics, i.e. also the physical stratum,
calls for a selectionnist or expanded Darwinian view: both what
determines the genesis of the universe itself (cosmogony) and
that of which it is composed (cosmology) in effect benefit by
being seen as the result of a series of constraints exerted upon
and on behalf of pure variation. But since nothing escapes pure
30
The variation or movement of the infinite “does not refer to spatiotemporal
coordinates that define the successive positions of a moving object and the fixed
reference points in relation to which these positions vary”, Deleuze and Guattari,
What Is Philosophy?, p. 37. For a rigorous reconstruction of the mathematical models
(intrinsic and n-dimensional Riemannian geometry, phase space, etc.) that underlie
or at least inspire this aspect of their fundamental ontology already present in the
early Deleuze, see the now classic study by DeLanda (2002).
31
“Chaos is defined not so much by its disorder as by the infinite speed with
which every form taking shape in it vanishes. It is a void that is not a nothingness
but a virtual, containing all possible particles and drawing out all possible forms,
which spring up only to disappear immediately, without consistency or reference,
without consequence. Chaos is an infinite speed of birth and disappearance”,
What Is Philosophy?, p. 118. For a detailed analysis of this latter fundamental
ontological aspect, of which the principles of superposition and indetermination
in quantum physics give a general scientific idea, cf. our work, Rosanvallon, 2009,
pp. 93e105.
32
A Thousand Plateaus, p. 502.
33
What Is Philosophy?, p. 42.
variation, or in other words since nothing is exterior to it, how
could it initially be constrained by something besides itself?
These constraints are therefore produced solely from the fact that
pure variation, which is in itself infinitely infinite, can only
intersect itself: the infinite cutting the infinite creating some
breaks that will play the role of cutoffs or limits.34 These
constraints in fact produce three distinct limitations that tend to
cumulate and conjugate together: the infinity of variables or
degrees of freedom of variation, the infinity of values taken by
these variables or ultimately the infinity of speed according to
which these variables vary could successively be limited. This
triple limitation then makes possible the fact that the “finite”, as
partial as it may be, i.e. for whatever the infinity it conserves
(infinite space-time, perpetual chaotic inflation, superposition of
quantum states, space of mathematical possibilities35), manages
to take on consistency within chaos or pure variation, which is in
itself neither exhaustible nor limitable and never stops remaining
infinitely infinite.36 But such constraints do not carry out a selection
of universe (“cosmogonic” selection) without first carrying out
a selection of these constraints themselves (“physical” selection).
Since pure variation varies in a purely contingent way, the breaks,
which are its effect, are just as contingent; but what is not
contingent is the effect of these breaks, namely what, with them,
acquires not simply a form of consistency but also subsistence
amidst everything that comes to exist. This is how the breaks or
limits that produce a “physically” subsistent and not merely
“mathematically” consistent reality will be selected: in the same
way that what the organic stratum selects (and what forms it)
gauges its reproductive success, above all what the physical
stratum selects (and what forms it) gauges its degree of
subsistence.
But which forms of relative subsistence does this process of the
self-limitation of pure variation specifically produce on the basis of
its absolute inconsistency? And how to gauge this subsistence
without intervening with a prior time, which would come more or
less provisionally to occupy or fulfil a certain kind of consistency
(any sort of multiverse, universe or physical process whatsoever)?
From a Deleuzo-Guattarian perspective, time envisioned formally
does not at all constitute the prior form of any subsistence but
simply the belated product of a multiplicity of subsistences, i.e. the
simple offspring of a correlation of variables establishing both
a continuous and uniform variable in step with all the other
rhythms or durations of variation.37 The primordial forms of
subsistence therefore in no way constitute temporal forms but
simply forms of infinite non-variation, the partial cessation of
infinitely infinite variation. These partial cessations can be
regrouped into three broad overlapping categories. First, a defini-
tive subsistence is acquired by what no longer varies, i.e. the breaks
or limits productive of physical reality that therefore become
constants not only of a universe, but also inevitably of an infinite
34
The so-called technique of renormalization in quantum field theory,
which introduces an arbitrary cutoff between the infinity of field oscillations
in order to be able to subtract the infinity of the excited field (with particles)
from the thus determined infinity of the empty field (without particles,
unless they are virtual) so as to end up with finite values, can offer a general
idea of the genesis and functioning of what such an intersection and break of
the infinite by the infinite would be.
35
These products of pure variation are the four levels or sources of staggered
multiverses proposed by Tegmark (2007).
36
“It is not the limited thing that sets a limit to the infinite but the limit that
makes possible a limited thing”, What Is Philosophy?, op. cit., p. 120.
37
Rovelli (2008) has thus shown the necessity and possibility of formulating
a classical and quantum mechanics without time, which is nothing but the
dependent product of a correlation of variables and not the independent frame for
the evolution of variables.
 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
population of distinct universes.38 Second, a relative subsistence is
acquired by what varies within these limits that border the infinity of
variables and/or values, thus forming, when these variations can all
be constituted and invert themselves, groups of transformations
(whether discrete or continuous and/or global or local) that
compose a number of symmetry groups or groups of invariances. In
effect, each group defines a double invariance in relation to all its
variations or transformations: on the one hand, it renders them
equivalent either as descriptions or as possible states of that which
varies, following the example of the principle of relativity which in
truth is only the translation of a spatio-temporal or gravitational
symmetry39; on the other hand, it implies that they conserve or at
least leave invariant a quantity or an aspect of that which varies
(energy, charge, isospin, etc.). Third, a more or less provisional
subsistence is ultimately acquired by everything that varies more
slowly than at infinite speed, thus participating in this “primordial
slowing down” from whence all the superposed material durations
that compose a universe like ours arise: the expansion of space,
interaction of elementary particles and atoms, galaxies, stars,
planets and molecules that are basically compounds of the
elementary, stabilized and variable at the same time40. The
Deleuzo-Guattarian perspective prompts us to consider each of
these durations as the outcome of a stratification of continually
renewed chaotic variation. The entirety of what has already been
selected (constants, interactions and other persistent physical
processes) in fact plays the role of stratum that will constrain,
capture and give durable consistency to what never stops varying:
the variable duration of stars results, for example, only from the
combinational play of the four fundamental interactions.
The (so-called spontaneous) symmetry breakings described by
contemporary physics can be better understood as the crucial
results of such a logic of stratification. A primary symmetry
breaking would be the breaking of the perfect cosmological
principle into a simple cosmological principle that conserves the
38
The three so-called fundamental constants, since they are neither the product
of any theory nor any other constants (the Planck constant h or minimum of vari-
ation, the constant c or maximal speed and the constant of gravitation G), therefore
do not determine the physics of our universe and the physical theories that describe
it without also being at the heart of an infinity of other universes as a consequence
of these same theories (for example, the theory of eternal chaotic inflation
specifically has as a preliminary condition the structuration of space-time operated
by c e this limit-speed is in fact in no way violated by the inflation that does not
define a speed of propagation in space but a speed of expansion of space itself, i.e.
a speed of intrinsic variation which would indeed also be initially always already
infinite).
39
The generalized covariance induced by general relativity can in fact be inter-
preted in terms of a group of diffeomorphisms or the “supple group” (see the work
of Jean-Marie Souriau specifically Souriau, 2002) that includes the subset of the
Poincaré group of special relativity and determines the gravitational field, i.e. space-
time itself. Three other continuous symmetries e which are called gauge invari-
ances, this time internal e correspond with three fields of interaction or funda-
mental types of variation other than gravitation (weak, strong and electromagnetic
interaction): they do not make equivalent the spatiotemporal descriptions or
becomings of physical objects, which are already determined, but determine the
presence and the becoming itself of fundamental physical objects (bosons or
particles of interaction) as conditions of their invariance. Ultimately these
symmetries include the discrete symmetries of charge, parity and time (CPT) from
which, on the one hand, anti-matter and, on the other hand, the “perfect cosmo-
logical principle” (isotropy of space and time) are deduced.
40
“It is by slowing down that matter.actualizes itself”, What Is Philosophy?, p. 118
(“primordial slowing down” is employed by Guattari, 1992, p. 112). If cosmogenesis
is conveniently presented as the history of a linear chronology, it in fact more
profoundly constitutes the history of a temporal slowing down (and not simply
a spatial inflation), i.e. from a production and superposition of durations on the
order of 109 years (duration of the universe’s current life) and even 1030 years
(duration of the proton’s minimal life) to the primordial durations characteristic of
Planck time (10 43 s), which marks the frontier of contemporary physics and the
domain of quantum gravity still to be elaborated.
143
isotropy of space but implies the anisotropy of time, i.e. the
entropic rupture of symmetry between past and future (the arrow
of time): rather than deploying our universe on the basis of
a singularity and a time zero (Big Bang theory), the expansion of
observable space would instead be nothing but one anisotropic
episode e among an infinity of others e of an eternal, i.e.
perfectly isotropic, chaotic inflation.41 This dynamic of space
expansion was at first inflationary before slowing down and then
accelerating again after several billion years: described by the
standard model of cosmology (Lambda-Cold Dark Matter model)
but not theoretically understood, it also seems to result from
a play of symmetrical forces between an attractive force of
gravitation (matter and dark matter) and a repulsive force of anti-
gravitation (cosmological constant Lambda or dark energy) slowly
broken in favour of the latter42 in a way such that the universe
has a duration of expansion sufficiently long enough to allow for
the appearance of other durations, i.e. other processes of strati-
fication or instances of its slowing down. Besides the primordial
quantum fluctuations that explain the anisotropies of the cosmic
microwave background and their post-inflationary residue e
which are the great structures of the universe (galaxies clusters)
e and besides the violation of CP within CPT symmetry that
explains the disappearance of antimatter,43 this first breaking
entails a second series of symmetry breakings e this time internal
e that describe, albeit still without being able to explain, the
standard model of particles. If inflation allows for the appearance
of particles which are no longer virtual but now actual, i.e.
susceptible to interaction and composition,44 the expansion that
has slowed down, by cooling down the temperature of the
universe, in other words, by reducing the amplitudes and
41
See Aguirre and Gratton, 2008; the model of eternal chaotic inflation has been
furnished by Andréi Linde, who was the first to think inflation not as one episode of
the Big Bang among others but the Big Bang as one episode of inflation among
others (theory of bubble universes).
42
The symmetry between past and future and/or its breaking indeed seems to
play a central role in this regard. Two types of completely different works, which
are trying to deduce in a natural way e without the creation of an ad hoc field e the
effect or value of the cosmological constant, attest to it: on the one hand, when
temporal symmetry is fully integrated into general relativity, although Frédéric
Henry-Couannier proposes to reformulate it albeit sadly in a way that is not
background independent (Henry-Couannier, 2005), it naturally generates a flat and
accelerating universe (Henry-Couannier et al., 2006); on the other hand and more
profoundly, when on the contrary temporal dissymmetry introduced by material
durations, i.e. by the necessarily cumulative character of past cones of light or
causality, is formalized in a theory like that of causal sets (candidate for quantum
gravity), the cosmological constant is equipped with a non-null value that always
fluctuates in the midst of the value of material density, thus explaining this “cosmic
coincidence” in a natural way (Sorkin et al., 2002).
43
Weak interaction in fact violates CP symmetry and compensates this violation
with that of the symmetry or invariance through T in order to conserve CPT
symmetry. This violation of T is translated by a dissymmetry between the duration
of the life of certain particles (like neutral kaons, beautiful mesons, but also
neutrinos) and the shortest life of their corresponding antiparticles. This dissym-
metry creates a surplus of matter over antimatter that will thus be selected or, in
other words, the complete primordial annihilation of matter and antimatter will be
allowed to subsist as the residue from which all the matter of our world stems.
44
We call “virtual particles” the creations/annihilations of particles (or production
of pairs of particle/antiparticle or fluctuations) of which the quantum void is the
incessant site due to the relation of Heisenberg’s time/energy indetermination that
renders the conservation of energy violable (and thus, due to the Einsteinian
equivalence of mass and energy, creatable particles) on condition that this violation
lasts a sufficiently brief amount of time, in such a manner that the product of this
energetic indetermination through the interval of time must not be inferior to the
reduced Planck constant (or quantum of action). This specifically signifies that there
always subsist infinite energies in infinitely brief times. To make this energy
durably consistent, to transform it into an actual particle, external energy must
manage to repay this loan granted by nature: inflation would precisely constitute
such a source of external energy; concerning this point, see Gunzig, 2008, pp.
176e217.
144 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
possibilities of variation, crosses certain thresholds of energy that
tend to break and constrain symmetries, i.e. to successively
disjoin and render autonomous strong, weak and electromagnetic
interactions and thereby render particles of interaction e other
than photons e and particles of matter massive themselves
(chiral symmetry breaking and Higgs mechanism). Particles of
matter thus become able to be composed into neutral atoms
(primordial nucleosynthesis) which will themselves then be able
to be fused into heavy atoms (stellar nucleosynthesis) and
composed into metals or molecules (chemical bonds). Thus
symmetry breakings do not result necessarily from a determin-
istic hypothetical theory but contingently from the pressure
exerted by the “environmental” constraints that constitute the
residues, which are also contingent and productive, of pure
variation. What results from these breakings then in turn
constrains, i.e. conditions and selects the ever persistent possi-
bilities of variation. Therefore the expansion will have selected
a certain physics of low energies which has itself selected the
possibility of chemical bonds and reactions, opening them, among
other reactions, to the immense combinational and reactive
possibilities of carbon compounds (organic chemistry).
3.2. Organic stratification: dynamic kinetic stability, membranous
selection and natural selection
The phenomenon of the living is naturally grasped within this
frame of thought: it does not emerge mysteriously from the
physical world (theories of emergence), nor is there an ultimate
adjustment of its parameters that makes it miraculously possible
or at least “anthropically” probable (multiverse theories); it simply
marks the capture and continuation of pure variation by other means.
In effect, everything happens as if a new stratification would then
lodge itself within physical subsistences and would manage to
capture or subtract pure variation more than the cascade effects of
the causal subsistences that constitute physical stratification.
Everything happens as if stratification would give variation a more
direct and more immediate consistency than simple subsistence,
albeit without using other materials, other forces or energies than
those resulting from physical stratification and the conditions that
it has randomly established on a planet like ours. The constraints
of the physical world are also necessarily those of the living
world, but the latter has its own additional constraints that para-
doxically allow it to break free from the first constraints or rather
to provisionally suspend them by inventing another mode of
organization capturing intrinsic variation more directly. Before
analysing in detail this new mode of organization that
alone legitimizes speaking of a new stratum, let us insist upon
the specificity of the Deleuzo-Guattarian approach e a faithful
generalization of the Darwinian schema e in relation to
the “emergent” and/or “anthropic” frameworks currently utilized
e specifically due to the lack of other existing theoretical
frameworks.
In the first place, guarding against “any ridiculous cosmic
evolutionism” that would like to make one stratum emerge from
another, for example the organic stratum from the physical
stratum, or that would like to define a progression in complexity
between these strata, Deleuze and Guattari ask us to never “ques-
tion how something escapes from the strata, but instead how
things enter it45”. As a pure differential process of selection without
any directed evolution or presupposed progress, stratification
consists in sifting a variation that is presented as external to it: each
stratum captures, retains or subtracts whatever it can, thus
45
A Thousand Plateaus, p. 49 and 57.
transforming stratification into a vast array of relative movements
or limited variations within it; each stratum is distinguished by the
sieve that it operates in such a way that, from one to another, there
is a “change of organization, not augmentation46”; each stratum is
therefore intrinsic, not to one another from which it would then not
be distinguished, but to pure variation alone as the inexhaustible
ground exhausting all that there is. In this way A Thousand Plateaus
envisions the strata as “spinoffs, thickenings on a plane of consis-
tency that is everywhere, always primary and always immanent47”.
Stratification thus is resumed by a constraint, a selection or a sieve
carried out, secondarily, as extrinsic and polyrhythmic, on an
intrinsic variation e which is always primary e by nothing other
than its products or “residues”. But, in the second place, the
“anthropic” selection of variation allowed by the multiverse
depends upon a calculation of probabilities which is, despite
certain appearances, the exact opposite of such a Darwinian
schema: the source of variability is then conceived as exclusively
extrinsic to our universe, whereas the effect of selection by contrast
would be purely intrinsic to it due to the “anthropic principle” as the
imperative of observability of such a universe presumed to deter-
mine its conditions facing downstream.48 Only “natural cosmo-
logical selection” responds to the double Darwinian requirement;
this type of selection has been proposed by Lee Smolin for whom
the actual values of parameters would simply be those that maxi-
mize the production of black holes, i.e. “baby universes”, since each
birth within a black hole would be brought about after a slight
stochastic variation of values already selected from the parent
universe.49 But this selection also supposes that variation would
have taken place only upstream from a cosmogenesis that remains
deterministic in the same way as ontogenesis within Neo-
Darwinian optics e as if the distinct universe or individual
organism were both completely determined by a legal or genetic
program whose sole preliminary definition would be partially
stochastic. A theory of “grand Darwinian unification” can really be
nothing but a “cosmo-onto-phylogenesis” that introduces, on all the
stages of formation of the physical world or living world, simulta-
neously intrinsic and stochastic possibilities of variation and extrinsic
and necessary constraints of optimization. If the physical stratum is
constituted by eliminating possibilities that lack a durable causal
effect and by optimizing cascade effects of subsistence (the value of
each subsistence being gauged by the number of variations and
46
Ibid., p. 49.
47
Ibid., p. 70. “Plane of consistency” (which, as these two authors point out,
more properly speaking constitutes a “plane of inconsistency”) in A Thousand
Plateaus would be equivalent to the idea of pure variation, which will only be
implicitly theorized as such in What Is Philosophy? correlative with the idea of
chaos.
48
Aguirre (2007) perfectly shows that the “standard” framework of the mul-
tiverse has no other scientific stakes than that of calculating a distribution of
probabilities such that it “would explain”, i.e. would make less improbable, the
known values of the parameters of standard models, a number of which would
seem so suitably adjusted that their variation in the slightest would have
produced a universe unfit for life: so as to be calculated, these probabilities
should therefore be balanced or conditioned by an “anthropic principle”, and
this conditioning should be made typical or not improbable by a complemen-
tary “principle of mediocrity”. Aguirre demonstrates that reflection on the
unavoidable steps of such a calculation (for example, the determination of the
more or less fundamental character of the parameters and their order of
precedence) is often neglected on behalf of “shortcuts” that often lead to
making a masked evolutionist and even finalist usage of the anthropic principle,
which is supposed in order to then determine the direction taken by our
universe from its origin.
49
See specifically Smolin, 1999, along with his article (Smolin, 2007). Neverthe-
less, this theory has the major flaw of projecting onto the physical stratum the
principle of reproduction proper to the organic stratum without being able to
translate this modifiable memory into a hereditary material equivalent to the
genetic code.
 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
thus of potential subsistences that it conserves)50, how is the
organic stratum constituted, what will it eliminate and therefore
tend to optimize?
Although not without certain theoretical shortcuts, we can
consider that, within the physical stratum, potential energy is
always minimized, i.e. minimally “acquired” and maximally
“expended” (principle of least action): every system tends to attain
its state of maximal stability or, in not so rigorous terms derived
from statistical physics, of maximal kinetic “disorder” (increasing
entropy or second principle of thermodynamics). But biochemical
processes are called “far from equilibrium” because they doubly
manage to free themselves from this thermodynamic stability.
Primarily, it seems that they simply suspend this stability: the
carbon compounds that all these processes make intervene are
specifically characterized by their metastability, i.e. their intrinsi-
cally delayed expenditure of potential energy. It is in this sense that
Gilbert Simondon proposed to define the vital as “the physical in
suspense, slowed down in its process and indefinitely
expanded51”. This does not at all signify however that the living
varies more slowly because this slowing down together is specif-
ically obtained only due to the partial acceleration of the varia-
tions or transformations within it, i.e. due to the energetic
degradations (catabolisms) and material syntheses (anabolisms)
that the living carries out in order to conserve itself in time and
grow in space: the organic metabolism of the cell, the elementary
biochemical system for all other systems, is in fact itself founded
upon the action of catalysts (enzymes) that accelerate chemical
reactions by reducing their activation energy. Thus uni- or multi-
cellular organisms constitute fundamentally open systems that
use the energy of solar rays or terrestrial chemical reactions
(phototrophs or chemotrophs) and establish themselves on the
basis of inorganic or organic matter (autotrophs or heterotrophs)
in order to conserve a local negentropy within the global entropy
of the Sun-Earth system. But if the organic stratum then remains
far from thermodynamic equilibrium, this would be secondarily,
because it responds first to another equilibrium that would be
induced by the foundational element around which it is organized,
including the chemically acquired capacity e on the molecular scale
e no longer simply to subsist but to be reproduced in all senses of
the word and in every way possible. What in this case tends
toward equilibrium is the difference between the rates of repro-
duction of what is produced and the elimination rates of what is
thus variably reproduced, in such a manner that a dynamic kinetic
stability e according to what the chemist Addy Pross calls it e is
created, namely a stability of the population of entities (first
molecular, then cellular and multicellular) with the capacity to
50
Within this “cosmo-onto-phylogenetic” optics, the presumed goal of
adjustment of the parameters becomes tautological: since the parameters’ values
have been duly selected amongst all possibilities in virtue of the causal effects of
subsistence and variation in their core which they successively or correlatively
produce, there is no surprise that they seem so optimal for producing such
effects! In this sense, a parameter would be confused with its concrete instan-
tiation, with all of its effects, without preceding them anywhere else but in
theory.
51
Simondon, 1995, p. 150. This intuition is taken up again by Deleuze and
Guattari, for whom the plastic properties of carbon carry out a “bifurcation” that
will “search the infinite chaos of the virtual for new forms to actualize by carrying
out a sort of potentialization of matter” What Is Philosophy?, p. 123 (my emphasis).
But the first to inspire such an intuition is actually Bergson, for whom, after
encountering the caricatured interpretation which is often foisted upon the concept
of élan vital, life “has not the power to reverse the direction of physical changes”
determined by the increase in entropy: “Incapable of stopping the course of
material changes downwards, it succeeds in retarding it”, Creative Evolution, pp.
245e246 (my emphasis).
145
multiply, specifically due to the perpetual replenishment and
variation of these same entities that make up the population.52
This equilibrium translates a new stratification, a fundamental
change of organization according to which what subsists is not only
what tends to no longer react or react less (thermodynamic equi-
librium of the physico-chemical stratum with which every dead
organism also tends to unite), but what is all the more capable of
reacting and participating in reactions as well as reproducing it
and, in this sense, being reproduced (dynamic kinetic stability of
the chemico-organic stratum).
What does “to be reproduced” mean? Life fundamentally
derives its origin from a process of the reduction and canalization of
a physico-chemical variation that takes form both in the (essen-
tially cosmic) prebiotic soup of elementary organic building blocks
and in the (essentially geochemical) energetic cycles based on the
transfers of proton and electron: on the basis of only a part of all
these possible building blocks and cycles, little by little organic
stratification has constructed and developed the constitutive
materials and energetic transfers that belong only to the living. If
the imperative of reproduction has indeed played a role of selective
constraint carrying out this productive canalization, this imperative
is realizable in three irreducible and thus originally distinct ways: to
be reproduced and to multiply passes through the fact either of
being replicated (from RNA to DNA), or of reproducing its conditions
of production (from “proto-metabolism” to metabolism53), or of
growing to the point of being divided (from the permeable passive
membrane to active cellular division). These three forms of organic
reproducibility each concern a type of population whose mode of
operation has entailed the rise of a specific molecular path: the
populations of “replicators” (only ones able to be replicated iden-
tically) that function through the pairing of the bases of a linear
chain have opened the nucleic path; populations of “catalyzers”
(only ones able to reproduce conditions of production) that func-
tion through the foldings of a linear chain of amino acids have
opened the proteinic path; and populations of “selectors” (only ones
able to distinguish an outside filtered from a retained inside and
divide themselves) that function through the closure of a dual layer
of amphiphilic molecules have opened the phospholipidic path.54 It
is therefore not a question of asking which of these three pop-
ulations e which are by right reproducible and even reproductive e
would have initiated organic stratification by preceding, or indeed
generating the others, but to which encounters, correlative opera-
tions and durable couplings, leading each to their current molecular
form, the imperative of reproduction, upon which this stratification
depends, has constrained all three e thus designating a primary
52
Pross, 2011 (http://www.jsystchem.com/content/2/1/1#B50). As we shall see,
the capacity for molecular replication upon which Addy Pross explicitly applies his
reasoning seems to us a certainly fundamental, but merely particular case of a more
general logic of reproduction which is constitutive of the organic stratum.
53
The idea of “proto-metabolism” has been proposed and explored in detail by de
Duve, 2005, pp. 15e24. The reproduction of the conditions of production does not
signify that the living completely auto-produces itself (according to the idea of
“autopoiesis” developed by Maturana and Varela (1979)) since it cannot do so
without external materials or energy, but signifies that, in their presence, what it
produces manages to never stop reproducing itself.
54
“In actual cells, membranes are never born de novo; they grow by accretion, i.e.
by the insertion of new molecules in a pre-existent network. Thus membranes arise
from pre-existent membranes linked by an uninterrupted filiation with an ancestral
membrane which could harken back to the first days of life on the Earth. This genre
of development leaves to the imagination an evolutionary process in which the
original components of membranes would have disappeared after a long time so as
to be progressively replaced by more elaborate molecules to ultimately arrive at the
phospholipids and other complex constituents of today”, de Duve, 2005, pp.
131e132.
146 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
series of (molecular) synergies or symbiogenesis of which unicellular
living beings are the ultimate product.55
The renowned coupling between replicators and catalyzers
appears in all current living beings as inevitably circular: in a sense,
it gives rise to genetic code, i.e. the coded production in DNA of the
proteins that reproduce the conditions of production for the living,
and, in an opposite sense, to proteinic enzymes as necessary vectors
of the activation, acceleration and precision both of the transcrip-
tion and translation of coding replicators and already of their
replication itself. On the basis of chemically and thus randomly
produced chains of amino acids and nucleotides entering into
cross-catalytic56 networks, the optimization of such a reproductive
coupling has undoubtedly taken millions of years: if RNA is
evidently the precursor of DNA, through which the dissociation of
the function of replication and functions of transcription becomes
possible, it remains to identify the precursors of RNA itself. The fact
that the residues of the vectors of organic metabolism, i.e. its
fundamental energetic currency (ATP but also GTP, CTP, UTP) are
also the basic constituents of RNA (AMP, GMP, CMP, UMP) in each
case proves the ancient status of this first coupling. The coupling,
which is also completely fundamental and circular, between
productive catalyzers and membranous selectors, in a sense, for
actual translation has the metabolic synthesis of the lipidic
constituents of cellular membranes and, in an opposite sense, the
proton-motive force based on the electrochemical gradients
maintained by the membrane that allows for the permanent
recreation of ATP. Also, the extremely long selective history of this
coupling could be described as the passage from a passively ob-
tained reproduction of the necessarily selective conditions of
production of a reproducible cycle (proto-cellular proto-
metabolism) to the randomly active and progressively acquired
reproduction of each of these conditions, up to the function of
membranous selection itself, whose growth no longer becomes
possible but produced (cellular metabolism).57 However, this
passage can only be accomplished through the coupling e which is
itself optimized e of the first two, from which therefore the just as
55
Beyond the badly posed debate between advocates of the primacy of replica-
tion, metabolism or the membrane which thus obey the three logics of reproduc-
tion irreducible to one another, scientists are forced to identify entities that would
all alone from the start have been able to fulfil at least two of these three functions.
Thus, for example, is grasped the whole interest of the theory of mineral surfaces
(proposed by Günter Wächterhäuser in 1988, see Wächterhäuser, 1988) that
simultaneously combine selective functions (due to their bi-dimensionality) and
catalytic functions (due to their electric charge) or the idea of an RNA world
(proposed by Walter Gilbert in 1986, see Gilbert, 1986) that endorses replicative and
catalytic functions (according to the current model of ribozymes). However, the
idea that irreducibly distinct logics of reproduction would be secondarily linked and
that their functioning in symbiosis would have been itself selected among other
possibilities, and thus constantly optimized, is imposed by the inevitable coupling
of the three forms of reproduction that no molecular entity can assume alone.
56
On the random chemical production of peptides or “multimers”, see de Duve,
2005, pp. 15e24; on the cross-catalytic network or “hypercycle”, see Eigen and
Schuster, 1979 and Pross, 2011.
57
Through this movement of internalization of its conditions of production (the
first cellular membranes no doubt having survived only by inventing autotrophy
and no longer simply heterotrophy, i.e. the direct degradation of inorganic matter
once the organic components of the cosmic prebiotic soup have been exhausted),
the organic stratum then globally seems to auto-maintain itself, the majority of
conditions apparently external to membranes in truth remaining internal to it, for
they are completely transformed by its increasing reproduction: “the materials
furnished by the substrata are an exterior medium constituting the famous
prebiotic soup, and catalysts play the role of seed in the formation of interior
substantial elements or even compounds. These elements and compounds both
appropriate materials and exteriorize themselves through replication, even in the
conditions of the primordial soup itself. Once again, interior and exterior exchange
places, and both are interior to the organic stratum. The limit between them is the
membrane that regulates the exchanges and transformations in organization”, A
Thousand Plateaus, pp. 49e50 (my emphasis).
decisive coupling between replicators and selectors also results. The
importance of this latter coupling is translated by the fact that
a third of the genome of today’s uni- and multicellular organisms
would code for the proteins inserted into the dual lipidic layer that
makes the membrane multifunctional,58 and in an opposite sense,
that another membrane in eukaryotes surrounds DNA (cellular
nucleus), this time dissociating the site of transcription and the
sites of translation. Even so, during the lengthy course of primordial
phylogenesis, only the membranous containers that each time
receive a selective advantage for their replicative and productive
contents have been able to grow and divide in order to attain the
first great bifurcation between bacteria, Archaea and eukaryotes.
3.3. The organism and the body without organs
Once cellular reproduction takes on dynamic kinetic stability in
a selectively optimized way, two other great ensembles correlative
with selectable symbiotic possibilities arose during nearly three
billion years: “If evolution includes any veritable becomings, it is in
the domain of symbioses that bring into play beings of totally
different scales and kingdoms59”. A second series of (unicellular)
symbiogenesis traces the decisive dividing line between prokaryotes
and eukaryotes. In the first case, living in symbiosis is limited to
molecular diffusions (“quorum sensing”) and the random, instan-
taneous and genetic (and perhaps even proteinic) transfers
between growing populations of cells that increasingly reinforce
their autonomy: their phospholipidic membrane is doubled by
a peptidic cellular barrier and the characteristics of this membra-
nous foliation end up diverging, thus giving unicellular entities
a mesophilic tendency (bacteria) or extremophilic tendency
(Archaea). On the contrary, in the second case, living in symbiosis
opens into an “endosymbiosis in series60” between Archaean and
bacterial cells, of which eukaryotic cells would be the product that
is ultimately selected and thus optimized: methanogenic Archaea
would have benefited from the hydrogen and carbon dioxide
produced by bacteria to the point of completely surrounding them
to become their single nourishing milieu and to link up with their
genetic material and their “respiratory” capacities so as to trans-
form them into simple organelles delegated to the production and
supply of energy for the “host” cell (mitochondria and chloroplasts,
the latter opening the royal phototropic path for vegetal life).61
Even a unicellular eukaryote (protists) de facto already constitutes
in itself a veritable organism with differentiated functions, begin-
ning with the function of nutrition or molecular transfer made
possible by the membrane’s vesiculation (endocytosis and exocy-
tosis). A third series of (multicellular) symbiogenesis, which relies
upon these various forms of unicellular reproduction, inevitably
follows from this in a wide variety of eukaryotes (and several rare
bacteria) e since multicellular reproduction in a sense has been
invented on various occasions within distinct cellular lineages. But
this new reproduction cannot occur without somewhat reactivat-
ing and transforming each of these three logics of molecular
reproduction that will become the three inseparable poles for all
unicellular reproduction. First, the replicative pole gives rise, already
within unicellular eukaryotes, to a new path of reproduction e sexual
reproduction, where a cell no longer divides simply by replicating its
genetic material identically (mitosis and asexual reproduction) but
by also randomly dividing this material (meiosis and formation of
gametes) so as to then have the ability to recombine it with other
58
Arkin and Stevens, 2000.
59
A Thousand Plateaus, p. 238.
60
Margulis, 1981.
61
Martin and Müller, 1998.
 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
material (fertilization and formation of a zygote). Second, the
membranous pole opens what could be called a new path of cellular
adhesion, where, due to reasons simultaneously internal (genetic
mutations) and/or external (environmental constraints), cells that
are nevertheless divided aggregate together in a terrestrial milieu
or remain non-separated in an aquatic milieu so as to form a single
organism that can grow and reproduce e at least partly and ulti-
mately fully e and consequently be submitted to the optimizing
action of natural selection.62 Lastly, the metabolic pole on this basis
authorizes and even favours a new path of division of cellular labor
that involves the differentiation of the organism into multiple
cellular types that can ultimately form tissues and distinct organs.
It is with these two last poles that ontophylogenesis intervenes,
conceiving the differentiated production of organisms (ontogen-
esis) and the divergent production of phyla and species (phylo-
genesis) as the same unique process governed by natural selection.
In effect, no multicellular organism develops and grows by itself
without exterior nutritive supplies e whether they first be
partially stocked, notably in the zygote phase (seed, embryo, etc.).
Flows of matter and energy are indispensable in order for each
organism to develop and endure throughout the same process of
division, differentiation and thus cellular renewal on the basis of
(animal) stem cells or (vegetal) meristematic cells that themselves
issue from zygotes. Then how could these flows constitutive of the
exterior milieu not have a selective effect before the development
of the fully grown organism? How could the phenotype of an
organism be submitted to natural selection without the phenotype
of each of the cells that compose it and on the basis of which it
develops not be submitted as well? In fact, natural selection or
organic stratification presents itself as a two-faced Janus that
operates, upon an intrinsic variability, a double constraint that
produces each cell of the organism in a tendentially stabilized and
optimized way. On the one hand, a certain cellular phenotype is
selected, i.e. favoured, by the immediate environment to which each
cell must adapt itself. This cellular environment is primarily
determined by cells nearby, including the extra-cellular, tissual
milieu and ultimately the rest of the organism, forming so many
metabolic filters of the exterior milieu, which tend to produce
variously concentrated gradients of factors of growth or regula-
tion, i.e. “signal-foods63”. Whether the organism, i.e. all the
cellular microenvironments, remains e notably in plants e
dependent upon variations of the exterior milieu or it breaks free
from them by continually compensating them for enabling
constant conditions of cellular life, thus constituting e in warm
blooded animals e a veritable “interior milieu64”, it no less offers
environments that are variable according to its phases of life and
its constitutive parts and therefore tends to favour differentiation,
specialization and consequently the growing interdependence of
cellular tissues and organic systems (digestive, respiratory, circu-
62
Bonner, 1998.
63
Formulated by Atamas (1996), the idea of “signal-food” has been taken back up
by Jean-Jacques Kupiec in order to think the fact that, even present in small
quantities, all intercellular signals (including hormones) function through the
metabolic model, i.e. serve to transform or reproduce the conditions of production
of cellular life (see Kupiec, 2008; pp. 137e142).
64
The idea of “interior milieu” has been developed by Claude Bernard concerning
living beings that acquire a life that is no longer “latent” or “oscillating” but
“constant or free” (see Bernard, 1878; pp. 65e124). Despite conceiving the interior
milieu as a selective milieu, Bernard however never integrated this idea into the
Darwinian schema, although Wilhelm Roux will propose this shortly afterward (see
Roux, 1881) and today ontophylogenesis theorizes it (see Kupiec, 2008; pp.
118e123).
147
latory, nervous, etc.).65 But on the other hand and simultaneously,
a certain cellular phenotype is selected, i.e. rendered more prob-
able, by the distant history in which each multicellular being is
inscribed and which is inscribed in the genetic material of its cells.
In a sense, the genome resumes the reproductive success of an
evolutionary lineage. It constitutes a memory that does not serve
a deterministic program, but a probabilistic reservoir, factory and
architecture. The fact that the first multicellular lineages had no
genetic material of their own e which would not already be
present in their unicellular ancestors even in a different degree or
quantity66 e proves the original inadequation of the concept of
genetic program. Do not homeogenes, so-called “architect” or
“developmental” genes, then determine, through their stabilized
order in the genome, the plane of organization of all bilateria?
Ordered distributions of coding sequences within DNA, variable
distributions of sequences expressible within chromatin and
probabilistic diffusions of regulators within the nucleus veritably
constitute so many historically acquired and selectively optimized
factors that simply come to constrain and stabilize the frequency
of expression e which is by right purely stochastic e of genes and,
consequently, the cellular maturation and differentiation in all
multicellular beings.67
Nevertheless, how do we understand this intrinsic cellular
variability which is only doubly constrained secondarily? Whence
does this objective margin of flexibility come? How do we pass
from simple cellular indifferentiation, without differentiated tissues,
of the sponge or algae, to the veritable cellular dedifferentiation of
organisms with multiple organs and interlocking systems, such as
bilateria or triploblastic organisms (including the majority of the
contemporary animal world) and angiosperms or flowering plants
(including the majority of the contemporary vegetal world)? The
evolutionary success of the former group, which marks the
Cambrian explosion, like that of the second group, dominating dry
land from the end of the Cretaceous, would precisely seem to be
tied in both cases to an accrued capacity of dedifferentiation (and
consequently at the same time a capacity of ontogenetic differen-
tiation and phylogenetic divergence): in effect, bilateria are char-
acterized by the intercalation e between the other two embryonic
layers directly exposed to the exterior milieu and destined to
protect them from it and sense it (ectoderm) or provide respiration
and nourishment for it (endoderm) e of a third layer (mesoderm)
on the basis of which internal organs (locomotive, reproductive,
circulatory, etc.) of the body arise; in the same way, angiosperms
tend to accrue the dedifferentiation characteristic of the whole
vegetal world by intercalating a new meristematic layer that
specifically allows for a wide variability in the formation of tissues
(heteroxylated and no longer homoxylated wood) and organs
(leaves, flowers and fruits). Thus everything happens as if the pre-
existent cellular layers of the organism would serve as a membrane
making possible dedifferentiation or random variation accrued
from a more internal layer. Everything happens as if embryonic
65
“In order to make possible and more rigorously regulate cellular life, organs join
together with organs and systems with systems. The task imposed upon them is to
qualitatively and quantitatively unite the conditions of cellular life”. Thus there
exists an organic perfectibility that consists in a “more thoroughly marked differ-
entiation of preparatory labor in the constitution of the interior milieu”, Bernard, 1878,
pp. 358e59. In this sense, Deleuze and Guattari already fully situate themselves
within a Bernardian lineage: “Neither is the organic stratum separable from so-
called interior milieus that are interior elements in relation to exterior materials
but also exterior elements in relation to interior substances. These internal organic
milieus are known to regulate the degree of complexity or differentiation of the
parts of an organism” (A Thousand Plateaus, p. 50), i.e. primarily, in cellular differ-
entiation, according to Kupiec’s model (Kupiec, 1997).
66
Rokas, 2008.
67
See Kupiec, 2008, pp. 162e174.
148 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149

tissues would tend to be duplicated in a logic of dedifferentiation [W]e treat the BwO as the full egg before the extension of the
analogous to the logic of decoding inherent to the genetic code organism and the organization of the organs, before the
whose duplications offer so many possibilities of variation. formation of the strata; as the intense egg defined by axes and
Having been present in the living on every scale, membranes vectors, gradients and thresholds, by dynamic tendencies
veritably owe their importance to the fact that they do not carry out involving energy transformation and kinematic movements
a membranous selection without at the same time being the site of involving group displacement [.] the egg always designates
an intrinsic random variation that also submits them to natural this intensive reality, which is not undifferentiated, but is where
selection. The reduction of the physico-chemical possibilities of things and organs are distinguished solely by gradients, migra-
variation ultimately leads to the creation of a series of variations tions, zones of proximity. The egg is the BwO. The BwO is not
internal to the organic stratum which, in the first place, is thus ‘before’ the organism; it is adjacent to it and is continually in the
translated by the decoding inherent to the code. This decoding or process of constructing itself.69
this variation is simultaneously minimized to allow for the
The BwO therefore naturally returns to the multiple sources of
conservation of every selective advantage (with the selected
the intrinsic variation of the organic stratum, most notably to the
appearance of enzymes that augment the precision of replication,
intrinsically undifferentiated or rather dedifferentiated cellular
correcting errors and locating mutations), but is also reinforced and
multiplicity of every organism. In its theoretical as well as practical
maintained to perpetuate genetic variation, the indispensable
usage, it primarily designates our capacity for corporeal being, such
source of every adaptive advantage (notably with the selected
that it escapes from the organization of the organs that constitute
appearance of sexual reproduction in animals and plants that
the organism, but also, secondarily, it designates the variations
submits genetic material to a shuffling no less passively undergone
intrinsic to the organism, such that they spontaneously escape from
but actively produced). The double membranous redoubling of uni-
its control. The relation between the BwO and the organism is
and multicellular eukaryotes (cellular nucleus and organism in its
consequently the same as that between variation not yet stratified
entirety playing the role of membrane) then leads to the entry into
or destratified and the organic stratum:
this selective play of two other sources of intrinsic random varia-
tion which we catalogued earlier: the ontogenetic dedifferentiation For the BwO already exists in the strata as well as on the des-
of cells and the phylogenetic deterritorialization of organisms. By tratified plane of consistency, but in a totally different manner.
systematically eliminating the possibilities with null, meager or Take the organism as a stratum: there is indeed a BwO that
lesser rates of reproduction, the organic stratum thus tends to opposes the organization of the organs we call the organism, but
optimize its capacities for reproduction and tends to conserve all the there is also a BwO of the organism that belongs to the stratum.
variations that favour it, to begin with variability in all its forms: in Cancerous tissue: each instant, each second, a cell becomes
this way the organic stratum manages to capture intrinsic variation cancerous, mad, proliferates and loses its configuration, takes
more than the physico-chemical stratum by directly, and no longer over everything; the organism must resubmit it to its rule or
indirectly, selecting it. As an epistemic consequence, this fact restratify it, not only for its own survival, but also to make
explains that the ontological primacy of variation has indeed been possible an escape from the organism, the fabrication of the
discovered and identified within its inner core (Darwin-Hooker “other” BwO on the plane of consistency70.
principle) before being able to be generalized for all the strata. The
On the one hand, the BwO can be viewed as inherent to the
organic stratum thus only subsists by being reproduced, i.e. by
organism and as incessantly wanting to escape from it. The first
submitting to a dynamic kinetic stability that governs all uni- or
relation between the two passes through instruments of control
multicellular living beings and each multicellular organism that is
responsible for resubmitting the body without organs to the rule of
constrained to maintain its gradients of concentration and repro-
the organism. The concept of BwO could thus anticipate the onto-
duce itself as such. If every organism is therefore fundamentally
phylogenetic vision which considers that the organism is neither
subjugated by this imperative of reproduction, it is at the same time
a programmed nor autotelic entity but an arbitrary scale of the
intrinsically submitted to the force of variation from which the
living and that cells are not as such at the beck and call of the
organic stratum retains or captures whatever it can in order to
organism but must always be constrained from the exterior by the
subsist: this is what both translates the idea of ontophylogenesis as
selective pressure of the intercellular and tissual milieu in relation
well as the idea of body without organs (BwO) that Deleuze and
to the available supports. When this external control loosens its
Guattari take up from Artaud and meticulously conceptualize.68
grip, their own reproduction is no longer subordinated to that of
Should we understand the strange concept of body without
the organism, and they inevitably become cancerous.71 But, on the
organs as an echo of the undifferentiated elementary cell of the
other hand, the BwO also designates this possibility of blending
unicellular living being (even if, as we have seen, every eukaryotic
with the plane of consistency. The second relation between
cell already contains within itself organelles or functions that are
organism and body without organs thus passes through (practical
specialized on behalf of its reproduction) or rather as an echo of the
or conceptual) instruments of destratification, through crossing the
fertilized egg cell or zygote e which is not yet divided and orga-
strata and grappling with pure variation.72 This path of destratifi-
nized into differentiated tissues and organs e of the multicellular
cation, which includes the greatest dangers, can only be followed
living being?
with the most extreme caution:

68
In the transmission of his radio play from 1947, To Have Done with the Judgment
69
of God, Antonin Artaud thus asserts: “Man is sick because he is badly constructed/ A Thousand Plateaus, p. 153 and 164.
70
We must make up our minds to strip him bare in order to scrape off that animalcule Ibid., pp. 162e163.
71
that itches him mortally,/god,/and with god/his organs./For you can tie me up if you See Kupiec, 2008, pp. 155e158 and the work of Jean-Pascal Capp (notably Capp,
wish,/but there is nothing more useless than an organ./When you will have made 2012).
72
him a body without organs,/then you will have delivered him from all his automatic See “How Do You Make Yourself a Body without Organs?” in: A Thousand
reactions/and restored him to his true freedom./Then you will teach him again to Plateaus, pp. 149e166, which specifically describes in detail masochistic practices in
dance wrong side out/as in the frenzy of dance halls/and this wrong side out will be their corporeal, sometimes extreme, aspect as an example of the destratification or
his real place.”, pp. 571e572. construction of a body without organs.
 J. Rosanvallon / Progress in Biophysics and Molecular Biology 110 (2012) 137e149
This is because the BwO is always swinging between the
surfaces that stratify it and the plane that sets it free. If you free
it with too violent an action, if you blow apart the strata without
taking precautions, then instead of drawing the plane you will
be killed, plunged into a black hole, or even dragged toward
catastrophe [.] Dismantling the organism has never meant
killing yourself, but rather opening the body to connections that
presuppose an entire assemblage, circuits, conjunctions, levels
and thresholds, passages and distributions of intensity, and
territories and deterritorializations measured with the craft of
a surveyor.73
In sum, pure variation therefore constitutes the fundamental
given that does not require any explanation but is the basis upon
which all aspects of reality must be explained. Stratification is the
process through which this pure variation acquires a subsistence.
The strata tend to thicken pure variation and give it consistency,
i.e. forms but also and above all duration: it is due to the latter
that they incarnate finite degrees and various rhythms of varia-
tion that are characterized by various degrees of slowness. They
produce the reality of things by limiting and constraining varia-
tion from the exterior. The physical world is nothing but the first
of the solutions discovered by pure variation to the problem of
durable subsistence, i.e. its endurance as variation: if physico-
chemical stratification only conserves the subsistences or invari-
ances of variation, organic stratification transforms the former
into a genuine conservatory by maintaining and reproducing
variation itself, specifically its intrinsic and stochastic compo-
nents. Thus pure variation simultaneously constitutes a “non-
organic life” wider than the material world and a body without
organs wider than the organism, and we can coincide with pure
variation by loosening the stranglehold of the organism (more
intense “non-organic life”) without ever for all that dismantling it
completely (risk of organic death).
Acknowledgements
I would like to extend my most sincere thanks to Jean-Jacques
Kupiec and Olivier Gandrillon for their cordial invitation to
present these ideas in this special volume dedicated to the
colloquium «Chance at the heart of the cell» as well as for their
meticulous rereading and their infinite patience. I would also like
to warmly thank Quentin Meillassoux for his fundamentally
friendly and fruitful remarks. Lastly, I want to extend special
thanks to Taylor Adkins for his elegant and precise labour of
translation, his unwavering support and his consistent
availability.
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