In evolutionary biology, the GARD (Graded Autocatalysis Replication Domain) model is a general kinetic model for homeostatic-growth and fission of compositional-assemblies, with specific application towards lipids.[1]

In the context of abiogenesis, the lipid-world[2] suggests assemblies of simple molecules, such as lipids, can store and propagate information, thus undergo evolution.

These 'compositional assemblies' have been suggested to play a role in the origin of life. The idea is the information being transferred throughout the generations is compositional information – the different types and quantities of molecules within an assembly. This is different from the information encoded in RNA or DNA, which is the specific sequence of bases in such molecule. Thus, the model is viewed as an alternative or an ancestor to the RNA world hypothesis.

The model

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The composition vector of an assembly is written as:  . Where   are the molecular counts of lipid type i within the assembly, and NG is how many different lipid types exist (repertoire size).

The change in the count of molecule type i is described by:

 

  and   are the basal forward (joining) and backward (leaving) rate constants, βij is a non-negative rate enhancement exerted by molecule type j within the assembly on type i from the environment, and ρ is the environmental concentration of each molecule type. β is viewed as a directed, weighted, complex network.

The assembly current size is  . The system is kept away from equilibrium by imposing a fission action once the assembly reaches a maximal size, Nmax, usually in the order of NG. This splitting action produces two progeny of same size, and one of which is grown again.

The model is subjected to a Monte Carlo algorithm based simulations, using Gillespie algorithm.

Selection

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In 2010, Eors Szathmary and collaborators chose GARD as an archetypal metabolism-first realization. They introduced into the model a selection coefficient which increases or decreases the growth rate of assemblies depending on how similar or dissimilar they are to a given target. They found that the ranking of the assemblies is unaffected by the selection pressure and concluded that GARD does not exhibit Darwinian evolution.[3]

In 2012, Doron Lancet and Omer Markovitch disputed this. Two major drawbacks of the 2010 paper were: (1) the authors focused on a general assembly and not on a composome or compotype (faithfully replicating and quasispecies, respectively); (2) they performed only a single, random simulation to test the selectability.[4]

Quasispecies

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The quasispecies model describes a population of replicators that replicate with relatively high mutations. Due to mutations and back mutations the population eventually centres around a master-replicator (master sequence). GARD's populations were shown to form a quasispecies around a master-compotype and to exhibit an error catastrophe, similarly to classical quasispecies such as RNA viruses.[5]

See also

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References

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  1. ^ Segré, Daniel; Ben-Eli, Dafna; Lancet, Doron (11 April 2000). "Compositional genomes: Prebiotic information transfer in mutually catalytic noncovalent assemblies". Proceedings of the National Academy of Sciences of the United States of America. 97 (8): 4112–4117. Bibcode:2000PNAS...97.4112S. doi:10.1073/pnas.97.8.4112. PMC 18166. PMID 10760281.
  2. ^ Segre, D.; Ben-Eli, D.; Deamer, D.; Lancet, D. (2001). "The lipid world". Orig Life Evol Biosph. 31 (1–2): 119–145. Bibcode:2001OLEB...31..119S. doi:10.1023/A:1006746807104. PMID 11296516. S2CID 10959497.
  3. ^ Vasas, Vera; Szathmáry, Eörs; Santos, Mauro (26 January 2010). "Lack of evolvability in self-sustaining autocatalytic networks constraints metabolism-first scenarios for the origin of life". Proceedings of the National Academy of Sciences of the United States of America. 107 (4): 1470–1475. Bibcode:2010PNAS..107.1470V. doi:10.1073/pnas.0912628107. PMC 2824406. PMID 20080693.
  4. ^ Markovitch, O.; Lancet, D. (2012). "Excess Mutual Catalysis Is Required for Effective Evolvability". Artificial Life. 18 (3): 243–266. doi:10.1162/artl_a_00064. PMID 22662913. S2CID 5236043.
  5. ^ Gross, Renan; Fouxon, Itzhak; Lancet, Doron; Markovitch, Omer (30 December 2014). "Quasispecies in population of compositional assemblies". BMC Evolutionary Biology. 14 (1): 265. Bibcode:2014BMCEE..14..265G. doi:10.1186/s12862-014-0265-1. PMC 4357159. PMID 25547629.
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