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Evolution of
The origin of species through living
system differentiation autopoietic
systems
The evolution of living autopoietic systems
Jörg Räwel 2365
Independent Researcher, Zürich, Switzerland
Received 7 March 2019
Revised 11 June 2019
2 September 2019
Abstract Accepted 10 September 2019

Purpose – Maturana and Mpodozis (2000) developed a theory of evolution that is based on the concept of
autopoiesis and differs paradigmatically from the conventional theory derived from Darwin (1859). The
present study aims to show that the authors have not exhausted the explanatory potential that the concept of
autopoiesis can offer for the theory of evolution. Based on the critique of Maturana and Mpodozis, a system
theoretic-oriented concept for the origin of species will be developed.
Design/methodology/approach – To render the explanatory potential of the concept of autopoiesis
more fruitful for the theory of evolution, the proposition is made that the application of this concept is not
limited to the molecular, or organismal level, as propounded by Maturana and Mpodozis, but should be also
related to populations and species. By exempting the design of Maturana and Mpodozis from the rudiments of
methodological individualism, a new field of application for the concept of autopoiesis is explored.
Findings – The proposed system theoretic concept of evolution theory makes it possible to shed new,
constructive light on fundamental problems in the conventional biology of evolution. For example, with
regard to the significance of the emergence of sexuality, or how phases of accelerated change in the course of
evolution (e.g. the Cambrian explosion) are possible, or regarding the problem of the units of selection.
Originality/value – Although there have been attempts in the social sciences to interpret populations as
autopoietic systems (for example by Niklas Luhmann), the proposed approach to evolutionary biology is new.
Also original is a system theoretic conception of the evolutionary theory, in a strict renunciation of
methodological individualism. This renunciation permits systems theories of evolution in social science and
biology to be compared across disciplines.
Keywords Adaptation, Ecology, Systems theory, Evolution, Autopoiesis, Self-referential systems
Paper type Conceptual paper

1. Introduction
Maturana and Varela (1980) developed the concept of autopoiesis with reference to the
molecular operations of living systems (cells, in the first instance), i.e. systems characterized
by self-preservation and the maintenance of their own organization. Autopoiesis thus
distinguishes what characterizes living organisms as actually alive. As autopoiesis specifies
what different living organisms are, it does not, in this respect, differentiate between them.
If we consider autopoiesis in connection with the evolution of living organisms (“The
origin of species by means of natural drift”; Maturana and Mpodozis, 2000), then
the conventional (neo-Darwinian) theory of evolution has to be radically overhauled. We are
here referring to the theory deriving from Darwin (1859) and its subsequent Modern
Synthesis, which integrates (Darwin still unknown) knowledge of genetics, population

Kybernetes
Vol. 49 No. 10, 2020
The author thanks anonymous reviewers for their valuable hints and helpful critical comments, pp. 2365-2383
which certainly contributed to the improvement of the paper. The author thanks Stan Jones for © Emerald Publishing Limited
0368-492X
editing the English language of this paper. DOI 10.1108/K-03-2019-0141
K biology, palaeontology, zoology, botany and systematics. Modern Synthesis was especially
49,10 advanced and established, above all, by the evolutionary biologists Dobzhansky (1937),
Mayr (1942) and Huxley (1942). The present paper explores the possibilities for a
paradigmatic revision of the neo-Darwinian theory of evolution, a revision that is appearing
increasingly necessary (Margulis, 1997a; West-Eberhard, 2003; Jablonka and Lamb, 2005;
Callebaut et al., 2007; Hoffmeyer, 2008; Pigliucci and Müller, 2010; Koonin, 2011; Shapiro,
2366 2011; Noble, 2015; Kull, 2016).
Taking the concept of autopoiesis into account makes it imperative to question the
everyday observation of the apparently abundant variety of adaptations that organisms
have to their specific environments. It seems evident that the differentiation of life forms in
water from those on land, for example, shows up precisely in their different specific
adaptations to their various habitats. However, the concept of autopoiesis demands that we
understand adaptation counter-intuitively as a constant. Adaptation in the maintaining of
autopoiesis is precisely what remains constant amid all the differences between living
systems[1]. Hence, it is not surprising that organisms can adapt to differing environments.
What is surprising is rather the variety of forms in which living systems are adapted to
different environments.
Maturana and Mpodozis (2000) demonstrate the necessary radical reconfiguration of the
conventional evolutionary theory under the condition of presupposed autopoiesis and
adaptation of living systems. In Section 2 of the present analysis, the authors’ theoretical
concept will be briefly sketched by way of recapitulation. Nevertheless, we intend to show in
the present analysis that the authors do not fully exploit the explanatory potential ensuing
from the thesis of the assumed autopoiesis of living systems (as posited) as well as the
concomitant reconfiguration of the theory of evolution. By way of explanation, we subject
their proposition to a critical examination in Section 3.
What then follows, Section 4, focuses on developing an alternative proposition for a
theory of the evolution of autopoietic systems. The crucial point here is the attempt to make
so-called “population thinking” (Mayr, 1959) productive for the evolution of autopoietic
systems to counter the weaknesses of the proposition by Maturana and Mpodozis. The
authors of “The origin of species by means of natural drift” ascribe autopoiesis exclusively
to systems of molecular operation, i.e. to cells and individual organisms. We will argue for
the theoretical productivity of also understanding populations and species as autopoietic
systems. The aim is not to counteract the insights from Modern Synthesis with those from
the application of the concept of autopoiesis to evolution, rather it is a matter of integrating
these insights into a unified body of knowledge; in other words, the development of a “post-
modern” synthesis[2].
In this way, the meaning and consequences of the development of sexuality, for instance,
can be determined more plausibly than is possible with the conventional theory of evolution
(“twofold cost of sex”, or respectively, “paradox of sex”). By the same token, the difficulty of
the conventional evolutionary theory to determine which units are targeted by natural
selection (for instance: phenotype, genotype or groups?) will turn out to be a pseudo problem
in the study at hand (on this problem, Mayr, 1997).
The analysis will conclude by briefly recapitulating the productivity and the problem-
solving potential of the proposition presented.

2. “The origin of species by means of natural drift” by Maturana and Mpodozis

The present discussion only has sufficient space to sketch the authors’ theoretical
proposition, and the ensuing consequences, along the lines of their results. Nevertheless, our
sketch should be understood also in its setting, which is necessarily incomprehensive. This
is because the authors derive theoretical consequences directly from their definition of the Evolution of
autopoiesis of living systems as a premise for their proposition. That means: living
[. . .] based on the understanding that living systems as autopoietic systems (Maturana and autopoietic
Varela, 1972) are structure determined systems, and that they exist as such only as long as in systems
their operation and their interactions they conserve their organization (autopoiesis) and their
operational congruence with the circumstances in which they live (adaptation). (Maturana and
Mpodozis, 2000, p. 263)[3].
2367
The theoretical concept further assumes that autopoiesis should always refer to individual
entities operating at the molecular level. Therefore, the authors are
[. . .] claiming that all biological phenomena, including those that take place in supra-individual
relational domains, have to be explained and understood by taking into account what happens to
living beings during the process of their realization as individuals that maintain their
organization and adaptation as a condition of their existence. (Maturana and Mpodozis, 2000,
p. 263)
As Maturana and Mpodozis derive their theoretical conclusions directly from these
premises, that is, in particular from the posited autopoiesis of all living systems, it suffices
here to present these, together with an explanatory reference to the concept of autopoiesis, as
set out in the following sections.

2.1 Autopoiesis
An autopoietic system is one:
[. . .] organized (defined as a unity) as a network of processes of production (transformation and
destruction) of components which (i) through their interactions and transformations continuously
regenerate and realize the network of processes (relations) that produced them; and (ii) constitute
it [the system, J.R.] as a concrete unity in space in which they (the components) exist by specifying
the topological domain of its realization as such a network. (Maturana and Varela, 1980, p. 78)
Hence, autopoietic systems are simultaneously producers and products of themselves and,
to that extent, determined by their own structures[4].
From this perspective positing the complete, internal structural determination of
autopoietic systems, environmental influences can only be understood as irritations or
perturbations – not as information or instructions from outside – that can be processed only
in the internal, structure-determined logic of these systems. The regenerative nature of
living organizations can only be altered to the extent that these perturbations affect the
reproduction conditions of these organizations. If the reproductive conditions of autopoietic
systems – operations on a molecular basis with respect to cells and organisms; (sexual)
reproduction with respect to populations and species – are completely disrupted, the death
(extinction) of these systems occurs (Maturana, 1988; Section 6.5).

2.2 Maturana and Mpodozis’ theoretical concept

The definition of autopoiesis immediately implies what is fundamentally capable of change
in the course of evolution and what – despite all evolutionary changes – is to be kept stable.
Evolution refers to the change of living systems and is, hence, only possible at all as long
as – despite all changes – the autopoietic organization and adaptation of living systems
continue to be preserved constant in a “medium”, or respectively, to an environment. The
point of reference of evolutionary changes are intrinsically, on the one hand, the structures
of autopoietic systems, and extrinsically, on the other, the medium (the environment) in
maintaining the autopoiesis and adaptation of living systems:
K [W]e claim that it is the conservation of adaptation and organization while living system and
medium are in continuous change, what defines moment after moment the courses that the
49,10 continuous change of the living system and its circumstances follow, and what makes that those
changes remain dynamically congruent in the conservation of the living of the living system
while this lives. (Maturana and Mpodozis, 2000, p. 276)
It should be noted that continuous change is a matter of course for the structures of a living
2368 system and for its environment. The continuously occurring operations – potentially
changing structures – are ensuring the stability of living systems and the maintenance of
autopoietic organization and adaptation. To this extent, evolutionary change is not in need
of explanation:
We claim that change occurs continuously as a spontaneous feature of the molecular existence of
living systems (and of all molecular systems), and that as an intrinsic condition of the existence of
living systems it must not be explained. (Maturana and Mpodozis, 2000, p. 276, emphasis added)
This is especially the case as the authors limit autopoiesis (with a high probability) to
individual systems operating at a molecular level: “[. . .] as molecular systems living systems
exist as the kind of entities that are perhaps the only ones that are autopoietic” (Maturana
and Mpodozis, 2000, p. 305).
Hence, we can state that all living things are subject to an individual “natural drift” in
maintaining their autopoiesis and adaptation. This includes, on the one hand – referring to
the lifetimes of individual organisms – “ontogenic structural drift,” understood as the “flow
of epigenic structural changes of a system in its domain of existence along its ontogeny”
(Maturana and Mpodozis, 2000, p. 307), and, on the other hand – with reference to
generations of living systems – “natural phylogenic drift,” understood as:
[. . .] a history of conservation of ontogenic phenotypes and of variation of the ontogenic
phenotypes conserved through systemic reproduction, in a process in which the organisms and
the medium change together in the conservation of some form of organism/medium relation.
(Maturana and Mpodozis, 2000, p. 279)
What is systemically maintained as a “lineage” is “a particular ontogenic phenotype/
ontogenic niche relation through systemic reproduction” (Maturana and Mpodozis, 2000,
p. 307). Hence, the maintenance of specific genetic structures through “natural selection” is
not characteristic for evolution; however, it is nothing more than an epiphenomenon of the
autopoietic maintenance and adaptation of living systems: “A lineage [. . .] is in fact
the result of the systemic reproductive conservation of a phenotype [. . .] and not of the
conservation of a genotype” (Maturana and Mpodozis, 2000, p. 307, emphasis added).
If autopoiesis and adaptation can be maintained as an ontogenic phenotype/ontogenic
niche relation, change or variation is basically possible regarding the system or the
environment. This means change or variation occurs with regard to the genetic structures of
an autopoietic system or with regard to its niche, or respectively, its medium:
[S]ince an ontogenic phenotype can be realized through many different total genotypes, the
conservation of an ontogenic phenotype through systemic reproduction allows the total genotype
to vary in the course of the generations as long as these variations do not interfere with
the realization of the ontogenic phenotype that is conserved. Similarly, since an ontogenic
phenotype can be realized under many different configurations of the medium as long as this
allows for the realization of the niche of the ontogenic phenotype, the conservation of an ontogenic
phenotype through its systemic reproduction allows for medium to change around the
conservation of the conditions of realization of its niche. (Maturana and Mpodozis, 2000,
pp. 276-277)
However, these possibilities of evolutionary change can ultimately only be stated in a Evolution of
descriptive way by observers. This is precisely why evolutionary change should be living
understood as “drift”. Maturana and Mpodozis focus theoretically on the maintenance and autopoietic
adaptation of individual autopoietic systems, or respectively, their intergenerational
maintenance as lineages. In this theoretical perspective, evolutionary transformation cannot
systems
be recognized in any other way than as being determined by the structures of individual
systems. 2369
The identification of other (external) factors would basically require a supra-individual
perspective. Ultimately, it is because of their perspective limited exclusively to individuals
that the authors can only provide a description but not an explanation for what happens in
the course of the evolution of living systems:
[T]he process of phylogenic drift occurs through lineages formed by systemically reproducing
individual living systems, even when these depend for their existence on the conservation of the
populations that they integrate. Furthermore, this is so even when some populations may
themselves, as a result of the particular manners of relating (such as sexuality or symbiosis) of the
individual living systems that compose them, constitute unities that exist as such in some other
relational domain. (Maturana and Mpodozis, 2000, p. 278, emphasis added)
In this respect, it is theoretically consistent that the authors do not provide any specific
determinants as regards the development of new lineages but are only be able to point to
randomness and spontaneity:
[T]he history of diversification of living systems, as well as the configuration of the biosphere, is
the result of the spontaneous generation and diversification of lineages in the natural phylogenic
drift that began with the systemic reproduction of living systems. (Maturana and Mpodozis, 2000,
p. 279, emphasis added)
Focusing on the autopoiesis of individual living systems, the evolutionary development of
sexuality appears of secondary relevance for the authors:
Certainly, sexuality modifies the phenomenon of genetic drift by establishing closed networks of
gene flow, and through that adds a new dimension to the dynamics of conservation of ontogenic
phenotypes and manners of living. [. . .]. Yet, none of these phenomena that modulate the course of
the phylogenic drift of the sexual organisms, alters the systemic nature of reproduction and
heredity, or the nature of the phylogenic drift in the terms that we have presented. (Maturana and
Mpodozis, 2000, p. 288)
As a consequence, the authors are not able to connect their perspective with the biological
species concept linked to sexuality. Rather, when assessing species, their perspective
depends on an arbitrary factor given by taxonomists:
[W]e consider that an appropriate characterization of the species as a taxonomic category is: A
species is a taxonomic category that corresponds to the distinction of a lineage defined by the
conservation of an ontogenic phenotype/ontogenic niche relation that may or may not include
sexuality, and which a taxonomist considers that has a nodal character in the dynamics of
diversification of lineages because the new lineages that may arise from it will have the same
nodal character in the course of the natural phylogenic drift. (Maturana and Mpodozis, 2000,
p. 291, emphasis added)

3. A critique of the origin of species by means of natural drift

The authors themselves provide a vital critique of their own notion:
K In this essay we claim that change and adaptation are not what we have to explain because living
constitutively takes place under continuous structural change in the conservation of autopoiesis
49,10 and adaptation. (Maturana and Mpodozis, 2000, p. 302, emphasis added)
Hence, evolutionary change is, like autopoiesis and adaption, simply assumed. It is, to a
certain extent, by definition, a self-evident and essential characteristic of the effective
operation of autopoiesis of (and in) living systems and therefore not in need of explanation.
2370 However, for a theory of evolution, a theory which, even if autopoiesis is always assumed,
seeks to explain the variety of living things, this conclusion seems to us to be extremely
unsatisfactory[5].
Accordingly, the concept of Maturana and Mpodozis leaves observers uncertain as to
how new lineages emerge. The authors mask this lack of clarity by asserting a spontaneous
generation of lineages, and yet more generally, by claiming that there is no need to explain
evolutionary change. When describing the variety of lineages (or respectively, species) by
taxonomists, this uncertainty – one, according to our notion, caused by their theoretical
concept – then reappears.
It is the authors’ strict focus on the individuality of organisms as a presupposed
“methodological individualism,”[6] making it impossible to find an explanation for the
evolution of living systems. This means an explanation, which does not depend on a merely
undifferentiated assertion of a change of living systems as an assumption of their (in turn
assumed) autopoiesis. Similarly, the authors, when discussing lineages, assume
metaphorically individual organisms, only taking into account the dimension of time.
However, we can only understand evolution if we abandon this perspective, precisely
because it does not go beyond the consideration of individuals.
If we can object to conventional theory of evolution for its focus on the variety of
living things without developing a concept (“autopoiesis”) of what remains the same
despite all the differences, then the complementary objection applies to Maturana and
Mpodozis. The authors’ strict focus on the autopoiesis of individual organisms
(autopoiesis exclusively on the basis of molecular operations) makes it impossible to see
how the distinction between living beings comes about or can be explained[7]. If the
focus is absolutely and strictly oriented on the autopoiesis of individual living
organisms, then a comparative consideration between (different) living organisms –
autopoiesis on the basis of the operation of reproduction – must indeed appear to be an
inappropriate abstraction. This makes it impossible for the authors to understand
explanations for differentiated forms of change in living systems such as the
acceleration of change in the sense of the “Cambrian explosion” (see, for instance,
Zhuravlev and Riding, 2001).
The authors’ methodological fixation on individuals is because they limit the
application of the – albeit conceptually abstract – autopoiesis theory to cells or
organisms, i.e. to systems that operate at a molecular level: “[. . .] as molecular systems
living systems exist as the kind of entities that are perhaps the only ones that are
autopoietic” (Maturana and Mpodozis, 2000, p. 305)[8]. Consequently the authors’
approach attracts Mayr’s (1959) critique, which opposes “population thinking” – without
which evolution cannot be understood – to an essentialist (typological) way of thinking.
The fundamental and exclusive consideration of individual organisms – taking a
methodical individualism as a basis – is in danger of absolutizing autopoiesis in an
essentialist manner.
In what follows, we will develop a supra-individual perspective on the autopoiesis of
living systems. For this purpose, we will argue that also populations and species have to be
understood as autopoietic systems. The aim is to link Modern Synthesis with the concept of
autopoiesis. By making “population thinking” (Mayr) accessible to the concept of Evolution of
autopoiesis, or, in the case of Modern Synthesis, by abandoning its merely statistical living
interpretation (population genetics), we propose a, so to speak, “post-modern” synthesis.
autopoietic
4. The origin of species through the differentiation of systems
systems
The order of sections undertaken here is not free from a certain arbitrariness because
evolution refers to circular events, and hence, it is not to be treated in a rigidly causal,
chronological sequence with matching of the serial logic of texts. Accordingly, we will now
2371
go on to explain how populations and species should be understood as autopoietic systems,
although an understanding of sexuality, which only made the evolutionary emergence of
populations and species possible, is presupposed. Sexuality will, however, come to be dealt
with in a later section. By the same token, we assume an understanding of an ecosystem –
which will only come to be dealt with in a later section – when we discuss species and
populations as autopoietic systems. A retrospective overview of the individual sections will,
however, make clear that species, or respectively, populations emerge through system
differentiation.

4.1 Populations and species as autopoietic systems

If populations and species are to be understood as autopoietic systems, then we must
initially check whether they satisfy the definition of autopoiesis (Section 2.1). In fact, it is
possible to consider individual organisms within populations and species as “nodes” of a
network, which replicate exactly that network through the self-preserving operation of
reproduction. Hence, species and populations are, in a strict sense, products and producers
of themselves (for an approach in the social sciences to understand populations as
autopoietic systems, cf. Luhmann, 2012, p. 273).
Can we, however, also assume that species and populations realize “a concrete unity in
space”? Do they “exist by specifying the topological domain of its realization as such a
network”? (Maturana and Varela, 1980, p. 78). In this context, the question arises as to
whether it is not essential that cells (as paradigms of autopoietic systems) possess a
membrane that populations and species obviously do not have.
It should be noted that the boundary between an autopoietic system and its environment
is not to be understood as a material entity but that it results from the operations of
autopoietic systems. All operations of an autopoietic system, which serve to promote these
actual operations, should be ascribed to the “inside” of this system while any other event
should be assigned to the “outside” of the system. The boundary (or “membrane”) thus
exists only temporally, i.e. it only exists as long as an autopoietic system is able to maintain
its (self-preserving) operations. Seen abstractly (or only spatially, without consideration of
the dimension of time), the “boundary” of an autopoietic systems presents a paradox (“the
unity of the difference between inside and outside”) that must be temporally resolved (“de-
paradoxed”) for the system to survive. Assuming that cells possess a membrane which
populations and species do not presents an inappropriate reification of the “boundary” of
autopoietic systems. The argument ignores the fact that the boundaries of autopoietic
systems “materialize” through their operations, i.e. through the circular and regenerative
production of those products that enable the production of exactly those products that allow
their own production (in the truest sense of the word autopoiesis). Autopoietic systems are,
therefore, closed operationally – also with regard to membranes – and not closed by a
stagnation of material entities.
Autopoietic systems are, to that extent, determined by their structures (“closed”), because
only those influences from their environment that impinge on the operations of the system
K can show any effect at all. Moreover, these influences show an effect only in as far as they
49,10 are processed within the system’s own logic, as determined by their structures.
Environmental influences, therefore, always have an irritating or disturbing effect on
autopoietic systems and never a causal or instructive one.
It is via the operation of reproduction that populations and species become autopoietic
systems. What matters is that, by reproduction, not (only) individual organisms are
2372 reproduced, but the reproducibility itself. Thus, reproduction in populations and species
is precisely that operation – in a network made up of organisms as structural
characteristics of these systems – which reproduces precisely this operation itself and the
boundaries of these systems. The reproduction of a specific species or population is only
possible within the boundary of this population, or species, never outside. Thus, the
boundaries between different populations and species are continuously reproduced or
maintained by the operation of reproduction itself. In this respect, we can make a direct
connection to the biological definition of species: “Species are groups of actually or
potentially interbreeding populations, which are reproductively isolated from other such
groups” (Mayr, 1942, p. 120).

4.2 The ecosystem

We can assume that populations and species are not to be understood as a precondition of
evolution, but only as evolutionary developments themselves. It is, therefore, not just
speculation but a necessary theoretical postulate when we reflect on the state of evolution
before the evolutionary emergence of sexuality (see Section 4.3) and consequently that of
species and populations.
What we want to define as the ecosystem is that comprehensive, autopoietic system,
which reproduces itself via the operation of reproduction and has its origin in the
appearance of the first autopoietic, self-reproducing living things (“viruses?”; Haldane,
1928), presumably some 3.8 billion years ago. Subsequently, it came to include populations
and species too after the evolutionary emergence of sexuality about a billion years ago
(Margulis and Sagan, 1990, p. 204ff.). Hence, the environment of the ecosystem is inanimate.
This is in contrast with populations and species, whose environment, as the internal
environment of the ecosystem, can be in addition animate, namely, through other
populations and species. In this perspective, the ecosystem must be understood in the
singular and is a corollary of the quasi “digital” definition of life by autopoiesis. Life
manifests itself in autopoietic organization, or not.
The operations of the ecosystem thus reproduce exclusively the boundary between
itself as a living system and the inanimate environment, while populations and species
do, in addition, as the internal environment of the ecosystem, operatively reproduce the
boundary between those living things capable of sexual reproduction and those not. To
this extent, the operation of reproduction (here the capacity for sexual reproduction)
should be regarded, when referring to species or populations, as contingent. If the
operation of reproducing a species or a population cannot be maintained, then this
simply leads to the end of that specific autopoietic system (for instance, a species “dies
out”), not of life altogether. However, this would be the case as regards the ecosystem if
the operation of reproducing could not be maintained here. Thus, the ecosystem (as the
whole world ecosystem) exists as a singularity; the operation of reproducing is here not
contingent. If reproduction cannot be reproduced here, then life and evolution (on earth)
are finished altogether[9].
4.3 Sexuality Evolution of
In the world of the prokaryotes (that is with regard to cells without nuclei), reproduction living
usually takes place through cloning. The reproduction of bacteria happens asexually
through the division of cells, among other things, through simple fission, budding and the
autopoietic
formation of spores (see Margulis and Sagan (1997) for a comprehensive account). This systems
process does not produce organisms with their own particular genetic attributes but
identical clones. Not until the evolutionary development of eukaryotes, of multi-celled
entities, and finally of meiotic sexuality, does the systematic reproduction of variants become 2373
possible. “The enormous power of the process of genetic recombination by sexual
reproduction becomes evident if we remember that in sexually reproducing species no two
individuals are genetically identical” (Mayr, 1976, p. 18).
However sexuality may have developed via evolution (on this, in common speculation,
Margulis, 1997b): it is, in any case, crucial that prior to the development of sexuality, only
unsystematic and sporadic reproduction of variants by mutation or through horizontal
transfer of genes (Mayr, 2001, p. 44) was possible. With the evolutionary emergence of
sexuality, systematic reproduction of variants was possible henceforth. If we bear in mind
that autopoietic systems can only change in as far as the conditions of their reproduction
change (environmental changes only have any influences to this extent), then we can assume
a revolutionary innovation. Variants were now not only generated randomly by mutational
reproduction any more – the operation of reproduction itself underwent, rather more, a
“mutation” and made systematic production of variants possible.
In relation to the ecosystem, we can assume that this evolutionary innovation
represents a destabilizing structural contradiction. The systematic production of variants
meant a risk for the conventional reproduction of the ecosystem (as an autopoietic
system) to the extent that, in their multiplicity, variants were now highly likely to differ
in their ability to reproduce as well, or respectively, could turn out to be competitors,
something that could, at least, impede the reproduction of the ecosystem as had been
usual so far. Reproduction – not only through cloning – no longer reproduced only (the
ability of) reproduction, but could now, through systematic generation of variants, also
reproduce an inability of reproduction systematically. The differentiation of reproductive
capacity and incapacity, fundamentally enabled through the systematic generation of
variants, basically made species and populations possible, subsystems of the ecosystem
that are stabilizing themselves operationally around exactly this difference or
respectively, at this boundary. Thus, it is precisely those networks of organisms capable
of sexual reproduction, which reproduce exactly that network and operationally maintain
the boundary to other organisms (populations and species) that are, as regards this
network, incapable of sexual reproduction.
This perspective refers not only to different reproduction rates, which are, in relation to
any form of reproduction, always also conditioned by random variations. Therefore,
“cloning” in this context cannot mean that, at least in the “world of prokaryotes”, error-free
information transmission would be possible. This is impossible because of the laws of
thermodynamics (Eigen, 1971). What is decisive here is, rather, that only the systematic
reproduction of variants by sexuality makes it possible that a system forming distinction
between reproductive capacity and incapacity arises.
Because of the evolutionary development of sexuality, the operation of reproduction itself
became contingent. Reproduction is, as the case of the prokaryotes shows, possible in a
particular way, but not exclusively in this form. Reproduction itself is, therefore, brought
into question by sexuality. Sexual reproduction should be understood as an evolutionary
reflection (variation) of what was previously the only form of reproduction (cloning);
K therefore, sexuality enabled a differentiation (formation of sub-systems) of the ecosystem by
49,10 the distinction thereby generated (reproductive capacity and incapacity).

4.4 Competition
The dominant individual-based perspective of the conventional theory of evolution suggests
a preference for selecting variants of organisms, which assert themselves through egoistical
2374 competitiveness against organisms from their own population or against organisms of other
populations (“struggle for existence”). From the present perspective, oriented not according
to individual organisms but according to autopoietic systems, the mechanism of
competition, understood as one of the fundamental structuring elements of conventional
theory, has no validity.
Individual organisms are, from the present perspective, to be understood as structural
elements of autopoietic systems. They represent “nodes” of a network, which operationally
maintains, through reproduction, precisely that network itself in its organization. That is,
just in the same way, as with a view to individual organisms, the genetic configuration
belongs to the structural elements of these organisms in maintaining their autopoiesis
(ontogenesis) and should not be understood as that crucial structure, which is maintained by
evolution through the comparatively better adaptation of its associated organisms
(phenotypes) as the “survival of the fittest”. It is hence not the selective maintenance
(“natural selection”) of specific individual organisms, their specific genetic configuration or
the selection of groups of organisms (Wilson, 1975) that are decisive for evolution, but the
maintenance of autopoietic systems (the ecosystem as well as species and populations).
Epigenetics has therefore, when related to species and populations, the same meaning as the
genetics related to individual organisms.
Just as it is meaningless, from the current perspective, to talk about competition between
individual organisms – as structural elements of autopoietic systems – so it is meaningless,
where the condition of the autopoiesis of populations and species applies, to assume
competition between populations and species. The maintenance, or respectively, the
reproductive capacity of autopoietic systems is structurally determined. There exists no
outwardly aligned agent that could realize a competitive mechanism. It makes no difference
to populations and species as autopoietic systems, whether irritative environmental
influences occur through other living systems or simply materially, for instance, by a
change in the climatic conditions. They can carry on reproducing themselves under these
(changing) conditions or, in dying out, not carry on. The fact that autopoiesis, if it is not to
fail, is only possible where adaptation is continuously sustained, requires us to assume,
rather more, a preference for co-evolution and symbiosis as something necessary for
evolution overall. This is a finding supported by empirical evidence (Margulis and Fester,
1991; Margulis, 1997a, 1999).
If the disputable mechanism of “competition” is to be retained for methodological
reasons, it must seem surprising from the neo-Darwinian theoretical point of view on
evolution that, in the present perspective, evolution is characterized through the restriction
of competition. As regards the ecosystem, the emergence of sexuality initially allowed
something of an “unruly” production of probably also competing variants. The
differentiation of the ecosystem constitutes a restriction of competition to the extent that the
sexual production of variants is now limited to populations and species. This means it is
limited to autopoietic systems where adaptation (formation of niches) is always a
precondition and hence a “competitive expansion” is precisely not what can be assumed.
The differentiation of systems, and with that the emergence of new species, thus comes
about through the restriction of destabilizing competition, which is being forced by Evolution of
increasing variability, i.e. the emergence of sexuality. living
In this context, the question arises as to how the tenacious focus of conventional
evolutionary theory on individual organisms, or respectively and as a consequence, on the
autopoietic
mechanism of competition, is to be explained. We assume that it involves systems
anthropomorphism: an inappropriate transfer of occurrences, enrooted in social evolution
based on communication, onto the circumstances of biotic evolution. This applies especially
because the observation depends on the context. Competition and cooperation can be 2375
observed with ease if the focus is on the behaviour of individual organisms. However, it
would be pointless to observe competition and cooperation in the context of system-internal
events of populations and species, understood as autopoietic systems.
To go any further in explaining social evolution would exceed the limits of the present
discussion (more detail on this issue: Luhmann, 2012, p. 251ff.). At this point, it can only be
stated that the concept of the “individual” has itself developed in social evolution (Luhmann,
1995a, p. 121ff.). It should be noted that communication itself solidifies around the
differences (as themselves, in turn, communicative constructed categories) such as “persons”
or “individuals” as “communicators” (Luhmann, 1995b, p. 137ff.). Therefore, it is entirely
plausible to assume that Darwin was, when developing his theory, guided by the social
circumstance of his time, for instance, by the so-called “Manchester Capitalism” (Weikart,
1998).

4.5 Adaptation
The conventional theory of evolution makes an obvious categorical mistake when it asserts
that individual organisms were maintained in their existence by a comparatively better
capacity for adaptation and reproduction. It is much rather the ecosystem or species and
populations as systems, which maintained their existence by maintaining their organization
and adaptation autopoietically.
With reference to individual organisms, it is varying diversity, in the sense of
structural flexibility, which is needed, much rather, to ensure the reproduction and
adaptation of an autopoietic system under changing (environmental) conditions. It is only
in this way, that at least a few organisms are able to reproduce nevertheless, despite, for
example, massive changes in temperature in the course of climate change, that the
autopoietic organization and adaptation of a given population or species is maintained.
Hence, it is the relative “non-adaptivity” of the structures (organisms) of autopoietic
systems, which permits their adaptation, i.e. the capacity to continued reproduction (of
their reproduction) despite changing environmental circumstances. To that extent, the
present study constructively takes up an influential critique of “adaptionism” (Gould and
Lewontin, 1979).
Adaptation, the capacity of autopoietic systems to maintain their existence through the
reproduction of their own reproduction is, in contrast to the view of the conventional
evolutionary theory, not to be understood as a gradual process. Autopoiesis is possible, or it
is not. Life and death are clearly separate categories.

4.6 Evolutionary mechanisms

4.6.1 Variation. In principle, we have to understand variations as reproductions
diverging from given structures in autopoietic systems: e.g. through mutations,
horizontal gene transfer genetic recombination or, in line with new research results,
through epigenesis (Pigliucci and Müller, 2010). The structures of individual organisms
– autopoietic systems on the basis of molecular reproduction – are determined by their
K genetic configuration. When referring to the ecosystem – or to species and populations
49,10 respectively as autopoietic systems, which reproduce themselves via the sexual
reproduction of organisms – “epigenetically” given individual organisms themselves
are to be understood again as structural elements[10]. Systematic variations through
genetic recombination (meiotic sexuality) occur exclusively in species and populations,
while in the ecosystem both unsystematic (resulting from mutations and horizontal
2376 gene transfer) and systematic variations take place after the evolutionary emergence of
sexual reproduction.
4.6.2 Natural selection. We understand natural selection as the difference between, on the
one hand, reproduction permitting survival of organisms (positive selection), and, on the
other hand, the premature demise of organisms rendering reproduction impossible (negative
selection). The term “selection” is misleading because positive selection is not about
selection, adaptation and maintenance of individual organisms, as is asserted by
conventional evolutionary theory, but simply about being an operation in maintaining an
autopoietic system, and thus, a distinction (survival/demise of organisms), which leads to a
further distinction (reproduction/non-reproduction of organisms).
Both the survival of organisms (positive selection) and their demise (negative
selection) belong to the productive operations that serve the maintenance of autopoietic
systems (see above with regard to the definition of autopoiesis: “transformation and
destruction” belong to the processes of autopoietic production). This is immediately
apparent: a population or species, as an autopoietic system, would relatively quickly
collapse into overpopulation if “positive” selection would be effectively preferred. In
this respect, “natural selection” is not an accomplishment of selection at all, but simply
has to be understood as the factum of the reproduction permitting survival, or
respectively, the premature demise of organisms, making reproduction impossible, in
autopoietic systems[11]. It would thus be more accurate to talk here of natural
production or natural construction. After all, it is an operation internal to the system.
The term “selection”, however, suggests a performance that goes beyond systems and
refers to only positive selection (the survival of organisms). It should be noted that the
demise of organisms (negative selection) is also to be understood as a productive
operation that provides for the maintenance of autopoietic systems.
Natural selection can be understood as natural drift in the sense proposed by Maturana
and Mpodozis (2000). This finding is not surprising because we follow these authors in
actually having to focus on the individual autopoiesis of organisms as regards the
evolutionary mechanism of natural selection. From our present perspective, these authors
have not dealt with the emergence of species in their study but have described the
evolutionary mechanism of natural selection (or more precisely: natural production) in
extreme detail.
4.6.3 Restabilization. While the evolutionary mechanism of variation concerns the
operations (reproduction) of autopoietic systems, the mechanism of natural selection refers
to their structures. The evolutionary mechanism of restabilization concerns the level of
systems. The issue here concerns the processing of resultant variations and selections in
such a way as to maintain autopoietic systems (see also the concept of “retention,” Campbell,
1974). Thus, for instance, the resultant systemic differentiation in species and populations in
the course of emergence of sexuality can be understood as accomplishing the restabilization
of the ecosystem.
In the conventional theory of evolution, the mechanism of restabilization remains
underexposed by the focus on individual organisms, which should, according to our
approach, be understood as structures of autopoietic systems.
4.7 System differentiation Evolution of
Evolution cannot, of course, be reconstructed in detail as a development involving the living
emergence and disappearance of untold millions of living systems (species and populations)
over billions of years. Nevertheless, the present theoretical perspective clarifies what
autopoietic
fundamentally promotes the development of species, or prevents it. systems
The differentiation of the ecosystem into species and populations was made possible
through the evolutionary emergence of sexuality and promoted a vibrant inner
environment within the ecosystem. Unlike the case of the ecosystem, the environment of
2377
species and populations is not just inanimate, it is also animated by other populations
and species.
Therefore, where species and population do exist, comparatively complex animate
environmental conditions are to be anticipated, and they, in turn, involve higher demands on
their capacity to maintain and adapt. Under variable, unsettled and unstable environmental
conditions, it is to be assumed that, within the varying spectrum of the structures of a
species or a population (i.e. with regard to individual organisms), only extreme forms are
able to continue to reproduce nevertheless. This circumstance forces the rapid change of a
species, or respectively, the emergence of new species by the system differentiation of the
original species. This happens by systemically extreme forms of organisms being
reproductively favoured from generation to generation, so that, in the course of time, their
differentiation from the original species is advanced to the extent that they are
reproductively isolated from it.
It should be emphasized that, from our perspective, this is not a case of the adaptation of
a species or population to extreme, volatile environmental conditions. Rather, extreme
structures (individual organisms) are reproductively favoured to continue maintaining the
adaptation of a (new) species or population despite changing and demanding environmental
conditions.
Therefore, a volatile environment essentially increases evolutionary change, a change
which, in turn, renders the environment of species and populations – as influenced by other
species and populations – more complex and challenging and so itself accelerates
evolutionary change [. . .]. Thus, we can assume recursive and self-reinforcing effects in the
emergence of species through the differentiation of systems[12]. This demonstrates precisely
the reason for phases of accelerated change in evolution (by way of example, the “Cambrian
explosion”; see, for instance, Zhuravlev and Riding, 2001). This theoretical perspective also
explains the emergence of so-called “hot spots” of high biodiversity (Myers, 1988), for
instance, in rain forests.
Yet, it is not just an accelerated change in evolution that requires explanation but a
surprising evolutionary stability as well. How can we explain the relative evolutionary
“uneventfulness” over more than 2 billion years? It occurs, namely, in the time between the
appearance of the first prokaryotes 3.8 billion years ago and the emergence of sexual
reproduction about 1 billion years ago (on the dates, Margulis and Sagan, 1990, p. 204ff.)[13].
It must be stated, first of all, that the environment of the ecosystem, as opposed to species
and populations as autopoietic systems, is entirely inanimate (cf. Section 4.2.). Rarely
changing environmental conditions can be assumed, making it scarcely necessary to change
the structures (prokaryotes) and with them the reproductive conditions, in order to maintain
the autopoiesis and adaptation of the ecosystem. This obtains despite the fact that, in the
long term (in the history of the earth), effects of feedback do occur also in relation to the
ecosystem: the alteration of the earth’s atmosphere, for example, as mainly attributable to
biogenic factors, into a (for most of the established organisms: deadly) one rich in oxygen
K certainly resulted in a wide-reaching restructuring of the organisms’ development and
49,10 genesis (Margulis and Sagan, 1997, p. 99ff.)
Despite massive changes in the environment, in the course of the Earth’s history, the
ecosystem’s autopoiesis and adaptation is obviously maintained because of its specific
structures: “They [prokaryotes, J.R.] reproduce asexually, they have very large populations,
and they are able to live under highly variable and often extreme environmental conditions”
2378 (Mayr, 2001, p. 47). Evidently, under these conditions, the structural diversity of the
ecosystem (Margulis and Sagan, 1997), which can be achieved by random and sporadic
mutative reproduction, or by horizontal gene transfer, is sufficient to ensure the continued
autopoiesis of the ecosystem (the maintenance of its organization and adaptation) over what
is now about 3.8 billion years[14].
It was, therefore, not extreme environmental influences which were the catalyst for the
emergence of species and populations, in order to restabilize the ecosystem. Rather, it was
what we could call a “mutation” in the reproductive conditions of the ecosystem itself, which
promoted the emergence of species and populations. In the ecosystem, sexuality enabled the
systematic variation of reproductions that was previously possible only in an unsystematic
way (randomly and sporadically).
In its autopoiesis the ecosystem is, therefore, far more stable than species and
populations. This more so, as the failure of restabilizations, related to populations and
species, manifests itself quasi-systematically as the so-called background extinction
(Lawton and May, 1995). Nevertheless, from the perspective of the Earth’s long history, we
have to assume the finitude of the Sun’s energy and, of course, the ultimate collapse of the
Earth’s ecosystem and the end of its evolution.

5. Conclusion
The proposal we outline here for a wider theory that integrates autopoiesis and evolution
diverges substantially from the theory outlined by Maturana and Mpodozis. The crux of the
difference between the two theories lies in their respective understandings of organismic
reproduction. In their focus on autopoiesis of individual organisms over the course of time,
Maturana and Mpodozis essentially maintain that the form of organismic reproduction is
practically meaningless in terms of evolution: “[W]e think that [. . .] a reference [to sexuality,
J.R.] does not introduce a substantial modification to the phenomenon of phylogenic drift”
(Maturana and Mpodozis, 2000, p. 288).
By contrast, we understand reproduction, as it refers to the ecosystem, precisely as
that form of operation that is responsible for maintaining it as an autopoietic system.
From our viewpoint, a change in the form of reproduction has, therefore, to have
significant consequences. Variants could now be produced systematically through
sexuality (genetic recombination), where previously prokaryotes were only varied
randomly and sporadically in the usual form of reproduction through cloning. As we
have shown, it is only in this way that system differentiation was possible. It became
necessary for the ecosystem to differentiate itself systematically, and in a self-
reinforcing manner in species, and populations. Indeed, from a functionalist
perspective, the evolutionary “invention” of sexual reproduction did pose a problem
for the ecosystem of that time, so that system differentiation, the emergence of
populations and species became the solution. The form of reproduction, as it is related
to prokaryotes, allowed, with mutations and horizontal gene transfers, merely the
increasing of the structural diversity of the ecosystem.
Nevertheless, it should be noted that Maturana and Mpodozis have indeed set out a
specific circumstance precisely. The point of reference is, however, not, as the authors
mistakenly state, the emergence of species, but the evolutionary mechanism of natural Evolution of
selection. The authors describe the factual occurrence of the differing survival, or living
respectively, demise of individual organisms in the course of evolution. They describe the
structural changes of the ecosystem, as well as of species and populations as autopoietic
autopoietic
systems. By focusing on lineages – individual organisms over generations in the course of systems
time – it is, in actual fact, only possible to assert changes. Here, it is not only not necessary,
as the authors note, but impossible to explain evolutionary change.
What is required is the supra-individual perspective we have presented, a perspective
2379
which abstracts from individual organisms to explain the emergence of species, or at least
makes it possible to understand in principle how differentiated forms of evolutionary
change, such as the “Cambrian explosion” or “hot spots” of high biodiversity, became
possible.
The perspective we have presented allows a better understanding of the evolutionary
emergence of sexuality and the problem of the units of selection. Sexuality is to be equated
with a radical change of the existing form of reproduction, which only allowed a system
differentiation of the existing ecosystem into species and populations. In this respect, this
approach supports Mayr’s biological species concept.
In the system theoretic perspective presented here, the problem of the units of selection
turns out to be a pseudo problem, one occurring in the context of methodological
individualism, which dominates the conventional evolutionary theory.
With regard to future research, it would be interesting to investigate whether
what we have presented as a system theoretic conceptualization of the origin of species
and populations can be reconstructed via computer simulation (although there are
doubts as to whether the concept of autopoiesis can be formalized at all, Razeto-Barry,
2012, p. 546). In this way, specific research results from population biology could be
linked with a theory of evolution itself oriented in accordance with systems theory
(Table I).

Maturana and
Topic Mpodozis Räwel

Methodology Methodological Functional method

individualism
Interdisciplinarity Erratic approach Connection with knowledge from the Modern
Synthesis, e.g. the “biological species concept”;
compatibility with theories of social evolution
Autopoiesis Limited to operations Also autopoiesis based on the operation of
on the molecular reproduction: populations and species
level: cells
Sexual reproduction Irrelevant for Decisive for evolution: to be understood as a
evolution radical modification of the conditions of
reproduction
Table I.
Differentiated forms of evolution, such Remains Explanation by mechanisms of feedback in Summarized
as Cambrian Explosion or “hot spots” unexplained the system differentiation of the ecosystem overview of the
of biodiversity comparison of the
Natural selection in the meaning of the Natural drift Natural production or construction theoretical
Modern Synthesis approaches
K Notes
49,10 1. In evolutionary biology, adaptation as a constant must be understood just as counter-intuitively
as, in physics, a speed, namely that of light, is understood. The development of the special
theory of relativity became possible because Einstein takes the contra-intuitive (but empirically
confirmed) idea of the speed of light as a constant seriously. The special theory of relativity
develops the physical consequences concomitant on abstracting from the concrete, plausible
everyday observation that in a classical (Newtonian) perspective speeds always cumulate. The
2380 dimensions of time and space now varied dependent on differing systems of reference, if
the speed of light has to be accepted as a constant. (Einstein, 1905, pp. 895 ff.). The radicality of
the change required in evolutionary theory can thus be compared with the paradigm shift
between classical physics and Einstein’s theories of relativity.
2. In the scientific literature, the concept of autopoiesis, at least in the form of the “self-organization”
of systems, can be regarded as established. In this respect, it is astonishing that Maturana’s and
Mpodozis’ approach to evolutionary theory, which takes up this concept, has so far received
relatively little response. In our opinion, the main reason for this is the erratic approach of these
authors, who hardly refer to any established research.
3. In this context, it is, therefore, not unreasonable to critically interrogate the status of Maturana’s
and Mpodozis’s proposition as a “theory” (as does Podgorski, 2010, p. 85).
4. See Razeto-Barry, 2012 for an overview of the currently existing variety of interpretations of the
original definition of autopoiesis. We refer to the original definition in what follows simply
because of our essential reference to Maturana and Mpodozis, 2000.
5. This form of theoretical proposition does, at least, once more justify our incomprehensive
presentation of its consequences. These are not derived from a complex theoretical construction,
but are linked straightforwardly (in fact, in the sense of an assumption) to the meaning of the
concept of autopoiesis.
6. See Dupré (1994) and Hodgson (2007) for a critical discussion of methodological individualism.
However, our approach, oriented as it is on systems-theory, generates knowledge on the basis of
the functional method (Luhmann, 1970; Räwel, 2007; Besio and Pronzini, 2008; or Knudsen, 2010).
Accordingly, our research findings are generated on the basis of a functional comparison
between the approach of Maturana and Mpodozis and our own proposal.
7. It is a logical requirement that a distinction – qua distinction – can only be distinguished with
reference to an undifferentiated “unmarked space” (Spencer-Brown, 1969). Recognizing the
dissimilarity of living systems assumes their similarity, namely that they are, without exception,
alive. Accordingly, given that we are talking about evolution, if variant differentiation is to be
recognized as differentiation at all, then it is not just a matter of identifying what is different, but
also what remains the same amid all the differentiation.
8. In this respect it is only consistent that Maturana strictly rejects the application of the concept of
autopoiesis to other operative domains (such as communication: cf. Luhmann, 1995b): “Thank
you for having made me famous in Germany,’ I said to Niklas Luhmann, but I disagree with the
way in which you are using my ideas.” (Maturana in Poerksen, 2004, p. 78) See instead Räwel
2017 for an application of the concept of autopoiesis in both social and biotic domains.
9. The notion of modern molecular biology, “that the prokaryote world is a single connected gene pool”
(Koonin, 2011, p. 141), can be well understood as an empirical indication of the concept of the
ecosystem as a comprehensive, singular autopoietic system. Similarly, the finding that “viruses are
the dominant entities in the biosphere, in both physical and genetic terms.” (Koonin, 2011, p. 325)
Viruses can be, as well as prokaryotes, regarded as structural elements of the ecosystem.
10. If species and populations are to be understood as sub-systems of the ecosystem, then individual
organisms, which are, again, differentiated sub-systematically into cells, can be considered once
again as sub-systems of species, or, of populations. Living systems are, therefore, organized in
 fractal manner (see Mandelbrot, 1979). Darwin’s evolutionary tree of life thus represents a special Evolution of
case of the generally fractal nature of evolution.
living
11. “One of the most basic questions of evolutionary biology is what objects are being selected in the autopoietic
process of natural selection? Lloyd (1992, Keywords in evolutionary biology, eds. Keller EF, Lloyd,
EA, Cambridge, 334–340) found nearly 200 references to books and papers by biologists and systems
philosophers, beginning with Darwin, that treated this question, “and these represent just a fraction
of the literature on the topic’, she reports. Indeed, in the recent literature the answer to this question
has been argued each year by at least a half dozen authors.” (Mayr, 1997, p. 2091) From the present 2381
perspective, the question as to the “objects of selection” (individual organisms? genes? groups?) is
revealed as a pseudo problem. The conventional theory of evolution fails to recognize that natural
selection is an operation, a difference producing a difference. It is, therefore, a matter of processing
information, not of selecting specific “entities”. Here, we adhere to a famous definition of information
from Bateson (1972, p. 315): “Information is definable as a difference which makes a difference”.
12. Only to refer to the events of singular populations or species, to explain the origin of (genetic)
complexity (“non-adaptive theory”, see Lynch and Conery, 2003), appears inappropriate from the
present perspective.
13. “Uneventfulness” is here understood as the remarkable stability of the prokaryotes: “About a
third of the early fossil species of prokaryotes are morphologically indistinguishable from still
living species and nearly all of them can be placed in modern genera”. (Mayr, 2001, p. 47).
14. The difference between the generation of structural diversity in the “world of prokaryotes” and
the production of a variety of species in the “world of eukaryotes” is reflected in the following
statement: “‘Microbes certainly evolve, but their evolution is quite different from the narrative of
Modern Synthesis.’ (Doolittle, 1999; Woese and Goldenfeld, 2009). The key insight is that
prokaryote genomes do not behave as if they were coherent, faithfully inherited repositories of
the genetic information of an organism (species)” (Koonin, 2011, p. 107).

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Further reading
Keller, E.F. and Lloyd, E.A. (1992), Keywords in Evolutionary Biology, Harvard University Press,
Cambridge, MA.

Corresponding author
Jörg Räwel can be contacted at: [email protected]

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