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Autopoiesis 40 years Later. A Review and a Reformulation

Origins of Life and Evolution of Biospheres volume 42, pages 543–567 (2012)Cite this article

Abstract

The concept of autopoiesis was proposed 40 years ago as a definition of a living being, with the aim of providing a unifying concept for biology. The concept has also been extended to the theory of knowledge and to different areas of the social and behavioral sciences. Given some ambiguities of the original definitions of autopoiesis, the concept has been criticized and has been interpreted in diverse and even contradictory ways, which has prevented its integration into the biological sciences where it originated. Here I present a critical review and conceptual analysis of the definition of autopoiesis, and propose a new definition that is more precise, clear, and concise than the original ones. I argue that the difficulty in understanding the term lies in its refined conceptual subtlety and not, as has been claimed by some authors, because it is a vacuous, trivial or very complex concept. I also relate the concept of autopoiesis to the concepts of closed systems, boundaries, homeostasis, self-reproduction, causal circularity, organization and multicellularity. I show that under my proposed definition the concept of a molecular autopoietic system is a good demarcation criterion of a living being, allowing its general integration into the biological sciences and enhancing its interdisciplinary use.

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Notes

  1. See Ruiz-Mirazo et al. 2004 for a criticism of this kind of descriptive, “check-out list” definition

  2. In Maturana and Varela (1980, p. 79) the definition was: ‘An autopoietic machine is a machine organized (defined as a unity) as a network of processes of production (transformation and destruction) of components that produces the components which:

    1. (i).

      through their interactions and transformations continuously regenerate and realize the network of processes (relations) that produced them; and

    2. (ii).

      constitute it (the machine) as a concrete unity in the space in which they (the components) exist by specifying the topological domain of its realization as such a network’.

  3. To give a few examples (italics mine): ‘All components of an autopoietic network are produced by other components in the network’ (Capra 1996, p. 167). ‘In this way the entire network continually “makes itself”. It is produced by its components and in turn produces those components’ (Capra 1996, p. 98). ‘In an Autopoietic system, the result of any given process is the production of components that eventually would be transformed by other processes in the network into the components of the first process.’ (Letelier et al. 2003, p. 266). ‘An autopoietic unit is a system that is capable of self-sustaining owing to an inner network of reactions that re-generate all the system’s components’ (Luisi 2003, p. 51). Note that, unfortunately, these definitions are not clearly differentiated from the descriptions of closed systems. There are also more ambiguous notions as: ‘autopoiesis… means “self-renewing”’ (Skyttner 2005, p. 60)

  4. To my knowledge, no organisms have been reported to date that directly obtain a functional enzyme from its environment, but the requirement of cofactors and coenzymes have been widely described, which are necessary to catalysis processes. Anyway, it seems evident that if we discover that a kind of bacteria directly obtains some functional enzymes from its environment, that would not negate that they are living beings. Indeed, the bacterial production of enzymes was discovered much later than the discovering of bacteria, which were considered living beings from the very beginning (Porter 1976). Note that in the model of Varela et al. (1974) there are catalysts that are not produced by any reaction in the network. Note also that if consider membranes as catalytic structures in the sense that they ‘emerge unchanged from the catalyzed reaction, while drastically increasing the reaction rate’ (McMullin 2000) and that the membrane of the first living beings probably self-assembled from simple lipids synthesized in the environment (Hargreaves and Deamer 1978; Hanczyz et al. 2003; Budin and Szostak 2011), then the first living beings on Earth would be examples of living systems with catalysts coming from the environment.

  5. Maturana and Varela usually differentiated between autopoiesis understood as the ‘organizing principle’ of living beings, and living beings as the ‘physical realization’ of autopoiesis (see Fleischacker 1988).

  6. It is perhaps worth mentioning that the border of viruses is formed by proteins; however, since there is no dynamic processes of production (metabolism) within this border, viruses cannot be considered to be autopoietic systems.

  7. I owe this last observation to Carlos Medina.

  8. Another advantage of not including the production of a boundary in the definition of autopoiesis is that non-molecular autopoietic systems, as society, may be more clearly characterized as autopoietic systems. In fact, one of the major objections to consider society autopoietic is the absence of a production of a boundary which makes sense: ‘These ideas are based, in my opinion, on an abuse of language. In autopoiesis, the notion of boundary has a more or less precise meaning. When, however, the net of processes is transformed into an “interaction among people”, and the cellular membrane is transformed into the limit of a human group, one falls into an abuse, as I expressly said.’ (Varela 2000, see also Luisi 2003). Note that medieval cities were defined by their boundaries (a closed defensive wall), however this kind of definition is not applicable in modern societies.

  9. The symbol (M,R) comes from ‘metabolism-repair systems’, also called ‘metabolism-replacement systems’ by Letelier et al. (2006). Essential to any materialization of a (M,R) system is the decay of the catalysts, which is counterbalanced by a group of replacement reactions which re-synthesizes the catalysts (Cornish-Bowden and Cárdenas 2007, 2008). By contrast, in an autocatalytic network the catalysts do not necessarily decay (or their half-life is much greater than the life of the organisms). I thank Carlos Medina for bringing this point to my attention.

  10. The authors make reference to the circularity of autopoiesis in most of their works. Here I take only two examples (from Maturana and Varela 1980): ‘…living systems as systems defined as unities through the basic circularity of their production of their components’ (xiv); ‘Living systems… are…all organized in a closed causal circular process’ (p. 9, italics are mine).

  11. According to Fleischacker (1988), the term ‘organizational closure’ was replaced by ‘operational closure’ by Varela since 1982.

  12. See Note 8. According to Robert Rosen ‘organisms are different from machines because they are closed to efficient causes.’ (Quoted in Letelier et al. 2003, p.265). Letelier et al. (2003, 2006) identified this claim with the circularity of autopoiesis subsumed by the term “metabolic closure”.

  13. Note that contemporary organizational approaches to function (e.g., Moreno and Ruiz-Mirazo 2009) do not ascribe functionality in any component of a living system created externally, which contrasts with the wide intuition that vitamins and cofactors are functional molecules which are not internally produced but obtained from the environment. Consistent with the present analysis the function of these molecules is guaranteed because they have the effect of allowing the maintainance of the autopoiesis, not matter whether they come from external or internal sources. Furthermore, many functional parts of multicellular living beings are microorganisms of other species, kingdoms and domains which are not internally produced but obtained from the environment (Pepper and Rosenfeld 2012; Costello et al. 2012).

  14. It was recently argued that in a network of biochemical processes the distinction between ‘enzyme’ and ‘metabolite’ is arbitrary and unnecessary (Cornish-Bowden and Cárdenas 2007). This reinforces the fact that it is not necessary for the understanding of autopoietic organization to know what types of elements enter into the system or what role they play in the network of chemical reactions (e.g., whether they participate in ‘peripheral’, ‘intermediate’ or ‘central’ reactions of the network).

  15. Any autocatalytic set ‘must be confined to a sufficiently small volume’ (Kauffman 1993, p.298).

  16. A review of the notion of probabilistic causality and its application to evolutionary biology may be found in Razeto-Barry and Frick (2011).

  17. According to Maturana and Varela (1973) “An autopoietic machine is a homeostatic system which has its own organization as the variable which it maintains constant” ([1994], p. 69, in this article all quotations from publications in Spanish are my translations). I criticize this affirmation in the next section.

  18. Note that this definition of ‘individual identity’ (or ‘quantitative’ or ‘numerical’) is only applicable in the context of systems in physical space. To apply this analysis to other spaces or domains (social, linguistic, psychological, etc.), the concept of ‘individual identity’ (and its differentiation from ‘qualitative identity’) must be clearly specified.

  19. This property may be related to thermodynamics as the maintenance of low entropy (see note 20).

  20. From this viewpoint, I dare to propose that the thermodynamic characteristic recognized in living beings as being systems permanently away from thermodynamic equilibrium (Schneider and Kay 1994) may be considered as a consequence of the definition of autopoiesis applied to systems of concatenated chemical reactions. That is, the mere requirement that a system maintain its individual identity in spite of consisting of concatenated chemical reactions requires that the system be in permanent thermodynamic disequilibrium. This affirmation is not immediately obvious and must be demonstrated. However, it is plausible from the fact that systems far from equilibrium may tend to form local zones of order (Schneider and Kay 1994), and that the spatio-temporal congruence proper to the components of a unit with a continuous existence in space and time constitutes a source of order in itself (as happens with tornadoes).

  21. A classical example of this is the woman who says “you’re not the same man I fell in love with”. Obviously she is not denying the “individual identity” of the man, she says it to him (e.g., Peter) and not to someone else. In fact, if she were talking to another person and not to Peter, we would either think she was crazy or had confused Peter with someone else. She means that he (the same individual, Peter) doesn´t have the same qualities he had when she fell in love with him; that is, she claims that his “qualitative identity” changed (see Tugendhat 2002 [1992]). The individual identity is not lost when properties change; what is lost is the qualitative identity.

  22. According to the author: ‘The relations between components that define a composite unity (system) as a composite unity of a particular kind, constitute its organization.’ He defines ‘structure’ as ‘The actual components (all their properties included) and the actual relations holding between them that concretely realize a system as a particular member of the class (kind) of composite unities to which it belongs by its organization, constitute its structure.’ (Maturana 1980, “Introduction”, pp. xix-xxx, in Maturana and Varela 1980). Maturana (1978, p. 32) affirms that organization ‘refers to the relations between components that define and specify a system as a composite unity of a particular class, and determine its properties as such a unity’, while structure ‘refers to the actual components and the actual relations that these must satisfy in their participation in the constitution of a given composite unity’. I thank Aldo Mascareño for the last reference.

  23. This also runs counter to studies of autopoiesis such as that of Cornish-Bowden and Cárdenas (2008), in which the maintenance of individuality is an accident, an abstract imposition which does not result from the dynamics of the system. The same can be said about the majority of computational models of autocatalytic reaction networks (see McMullin 2000).

  24. It is possible that what made so attractive the introduction of the notion of ‘circularity’ in cybernetics due to the ‘circular causality’ (of feedback systems), is that we erroneously tend to think that if the causality of tokens is never backward (that is, there is never ‘backward causation’, see Faye 2010), then neither is the causality of types. The causal circularity of feedback is precisely a circularity of types of events, not of particular events.

  25. ‘What is common to all metacellulars in the five kingdoms is that they include cells as components of their structure. That is why we say that metacellulars are second-order autopoietic systems.’ (Maturana and Varela 1992, p. 87)

  26. Note that the formation of external and internal boundaries in multicellular organisms is a derived evolved property, posibly increasing autonomy (Rosslenbroich 2009), analogously to the formation of external and internal membranes in the first cells (Hargreaves and Deamer 1978; Hanczyz et al. 2003; Budin and Szostak 2011) and in firsts unicellular eukaria (Kutschera and Niklas 2008), respectively. The idea of a living being as a “bag” containing liquid should be thus given up.

  27. ‘These [colonies of social insects] are sometimes referred to as superorganisms, but why not just call them organisms? From an adaptive perspective, a colony of social insects is analogous to the colony of cells that comprise a human’ (West and Kiers 2009). Following Queller and Strassman (2009), we may say that an autopoietic system is a system with high level of cooperation among parts based on “strong reciprocity” (see West et al. 2011 for a critical review), such that a typical homeostatic autopoietic system is a system with high “indirect reciprocity” among types of molecules (autocatalytic network) and some level of “punishment” (degradation of molecules).

  28. I owe this point to Carlos Medina (see note 7).

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Acknowledgements

I thank Carlos Medina, Roberto Torretti, Luis Razeto, Lafayette Eaton, Gabriel Razeto and Leonardo Sepúlveda for useful comments on previous versions of this manuscript. I especially thank Carlos Medina for a number of ideas in this article (see notes 7, 9, and 28). I thank an anonymous reviewer for useful suggestions.

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  1. Instituto de Filosofía y Ciencias de la Complejidad, IFICC, Los Alerces 3024, Santiago, Chile

    Pablo Razeto-Barry

  2. Universidad Diego Portales, Vicerrectoría Académica, Manuel Rodríguez Sur 415, Santiago, Chile

    Pablo Razeto-Barry

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  1. Pablo Razeto-Barry
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Correspondence to Pablo Razeto-Barry.

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Razeto-Barry, P. Autopoiesis 40 years Later. A Review and a Reformulation. Orig Life Evol Biosph 42, 543–567 (2012). https://doi.org/10.1007/s11084-012-9297-y

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Keywords

  • Autopoiesis
  • Definition of life
  • Individual identity
  • Physical proximity
  • Self-reproduction
  • Circular causality