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Organizational invariance and metabolic closure: Analysis in terms of (M,R)
systems
Juan-Carlos Leteliera,
,
, Jorge Soto-Andradeb,
Flavio Guíñez Abarzúab,
Athel Cornish-Bowdenc
and María Luz Cárdenasc
aDepartamento de Biología, Facultad
de Ciencias, Universidad de Chile, Casilla 653, Santiago, Chile
bDepartamento de Matemáticas, Facultad de Ciencias,
Universidad de Chile, Casilla 653, Santiago, Chile
cBioénergétique et Ingénierie des Protéines, Institut
de Biologie Structurale et Microbiologie, Centre National de la Recherche
Scientifique, 31 chemin Joseph-Aiguier, 13402 Marseille Cedex 20, France
Received 21 February 2005; revised 5 July 2005; accepted 7
July 2005. Available online 24 August 2005.
This article analyses the work of Robert Rosen on an interpretation of metabolic networks that he called (M,R) systems. His main contribution was an attempt to prove that metabolic closure (or metabolic circularity) could be explained in purely formal terms, but his work remains very obscure and we try to clarify his line of thought. In particular, we clarify the algebraic formulation of (M,R) systems in terms of mappings and sets of mappings, which is grounded in the metaphor of metabolism as a mathematical mapping. We define Rosen's central result as the mathematical _expression_ in which metabolism appears as a mapping f that is the solution to a fixed-point functional equation. Crucially, our analysis reveals the nature of the mapping, and shows that to have a solution the set of admissible functions representing a metabolism must be drastically smaller than Rosen's own analysis suggested that it needed to be. For the first time, we provide a mathematical example of an (M,R) system with organizational invariance, and we analyse a minimal (three-step) autocatalytic set in the context of (M,R) systems. In addition, by extending Rosen's construction, we show how one might generate self-referential objects f with the remarkable property f(f)=f, where f acts in turn as function, argument and result. We conclude that Rosen's insight, although not yet in an easily workable form, represents a valuable tool for understanding metabolic networks.
Keywords: (M,R)
systems; Metabolic network; Metabolic closure; Infinite regress; Systems biology