Endosymbiosis and self-organization
Resumen
From the beginning, symbiosis has been viewed as an association of two specifically distinct organisms living together. Firstly, symbiosis may suggest an idea of mutual benefit, but partners become dependent on symbiosis. Such dynamical phenomenon between two organisms in physiological interdependence is liable to evolve by creating a new life unit. This notion implies a reorganization of genomes and seems especially adapted to describe endosymbiosis. Putting forward a hypothesis of interactions leads us to consider the organism as a consequence of a self-organization process. In this paper, we refer to Kauffman’s works and to various cases of endosymbiosis. Considering the symbiotic genome, some biological results tend to indicate that models with (C=1 or C=2, K=1 or K=2) are not unrealistic and should be tested. But different epistemological questions prompt us to reflect further on the role of natural selection, the reality of the modeled interactions, the nature of the process of mutual adjustment. Moreover, regular interactions cannot exist in constant numbers for each gene. In spite of all these caveats regarding the applicability of Kauffman’s model concerning the establishment of a symbiosis in amoebae, simulation and experimentation yield results included in the same order of magnitude.
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S.-A. Kauffman, (note 37), p. 177. According to Weisbuch (1989), attractorsrepresent an ensemble of states of the system forming a loop. If each cellular type corresponds to a cycle of attractors, this means that differentiation results in reproducing indefinitely all the cellular types of the organism, the different cycles being regulated and, in some way, showing “solidarity” among themselves. Certain cell types are prone to differentiation, others are rather final results. As B. Feltz points out, the boolean model can help in interpreting the phenomenon of cellular differentiation. See: G. Weisbuch, Dynamique des systèmes complexes, Une introduction aux réseaux d’automates, Paris: Editions du CNRS, 1989. B. Feltz, “Auto-organisation, sélection et émergence dans les théories de l’évolution”. In: B. Feltz, M. Crommelinck, P. Goujon (ed), Auto-organisation et émergence dans les sciences de la vie, in printing, Bruxelles: Ousia, p. 332.
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The epistemologist must analyze biologists’ understanding of this newstructure and associated characters. According to R. N. Brandon (in printing, 265-279), one can apply either the selected effect (ES) model or the causal role (RC) model. The selected effect (ES) model will give:
(1) A bacterium has begun to live in intracellular symbiosis with the cerealweevil.
(2) This new genetic heritage represents something of a «gene kit» whichcan be transmitted to descendants.
(3) Selection acts on the physiological advantage brought by the symbiont(in the case of the cereal weevil, the supply of 5 vitamins).
(4) The «selected» interrelation between host and symbiont results in aninterdependence, where the symbiont can no longer survive outside the host. The RC model can be described as follows: The eukaryotic cell functions through the ability to integrate a bacterium in the physiological mechanisms of growth and reproduction of the host (s). The functioning of this model is relative to an analytic explanation (A) (here showing how the eukaryotic cell receives new genetic aptitudes). A is relative to the capacity of s to accomplish G (to transmit the endosymbiont to descendants). The eukaryotic cell functions in that manner because A adequately elucidates the aptitude of s to accomplish G, drawing on the aptitude of the eukaryotic cell to effect the integration of the endosymbiotic bacterium.
These two models are borrowed from Brandon, who himself borrowed the second one from Armundson and Lauder (1994). The first model involves a teleological view. The second is totally void of the idea of finality and aims only to analyze a complex system as such. It can be interpreted only in terms of capacity, functionality and the resulting structure: the integration of the endosymbiotic bacterium in the eukaryotic cell. See: R. Armundson, G.V. Lauder, “Function without purpose : the uses of causal role function in evolutionary biology”, Biology and Philosophy, 9 (1994), 443-469. R.N. Brandon, “La téléologie dans les systèmes auto-organisés”. In: B. Feltz, M. Crommelinck, P. Goujon (ed), Auto-organisation et émergence dans les sciences de la vie, in printing, Bruxelles: Ousia.
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The limits to reasoning coming obviously from the extreme simplification ofbiological data by use of the values of «0» or «1».
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S.-A. Kauffman, “Investigations, The nature of autonomous agents and theworld they mutually create”, Lecture 1, Santa-Fe Institute, 1996.
S.-A. Kauffman, (note 43), pp. 215-223.
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A computer simulation was made for N = 30, K =2, C = 1, and seems to show the establishment of cycles after several hundred repetitions based on random connectivity for interactions of the types «K» or «C». Indeed, in simulating an induction on the part of the host on the symbiont, then two intra-genomic interactions, and a retro-action of the symbiont on the host (reciprocity of interaction of type «C»), one obtains at first a chaotic result (from t to t+1, the number of activated sites varies up to 100% without there being really any order). When the simulation runs about 250 to 300 times, one gets a certain regularity in the sequences with periodicity of 5 to 6. Cycle length on the order of 5 to 6 corresponds well to Kauffman’s forecast (1993) which gives the root result (N) for the number of cycles as well as for their length. It appears that a certain equilibrium is reached before the value of 700 generations, the maximum forecast by Kauffman. In any case, one finds the same order of magnitude.
P. Nardon, “Symbiosis and evolution : the weevil model (Sitophilus Oryzae,Coleoptera, Curculionidae)”. In: R. Lesel (ed.), Microbiology in Poecilotherms, London: Elsevier Science Publishers, 1990.
See: K.W. Jeon, (note 26b).
See: K.W. Jeon, (note 22a).
See: S.-A. Kauffman, (note 37) et (note 70).
See: H. Charles, G. Condemine, C. Nardon, P. Nardon, (note 19). 105 See: R. Von Sternberg, (note 94).
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