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In [[evolutionary biology]], the 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.<ref>{{cite journal |last1=Segre |first1=D. |last2=Ben-Eli |first2=D. |last3=Lancet |first3=D. |year=2001 |title=Compositional genomes: Prebiotic information transfer in mutually catalytic noncovalent assemblies |journal=Proc Natl Acad Sci U S A |volume= |issue= |pages=219–230 |publisher=Elsevier |doi=10.1073/pnas.97.8.4112  |url=http://www.pnas.org/content/97/8/4112.abstract }}</ref>


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In the context of [[abiogenesis]], the [[Abiogenesis#Lipid world|lipid-world]] <ref>{{cite journal |last1=Segre |first1=D. |last2=Ben-Eli |first2=D. |last3=Deamer |first3=D. |last4=Lancet |first4=D. |year=2001 |title=The lipid world |journal=Orig Life Evol Biosph |volume=31 |pages=119–145 |doi=10.1023/A:1006746807104 |url=http://www.springerlink.com/content/rq23036351237250/ |pmid=11296516}}</ref> 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 of an [[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 GARD model==
 
The composition vector of an assembly is written as: <math>v=n_1\cdots n_{N_G}</math>. Where <math> n_1\cdots n_{N_G} </math> are the molecular counts of lipid type ''i'' within the assembly, and NG is how many different lipid types exist (''repertiore size'').
 
The change in the count of molecule type ''i'' is described by:
 
: <math> \frac{dn_i}{dt} = (k_f \rho_i N-k_b n_i) \left(1+\sum_{j=1}^{N_G}\beta_{ij} \frac{n_j}{N}\right) </math>
 
<math>k_f</math> and <math>k_b</math> are the basel forward (joining) and backward (leaving) rate constants, ''β''<sub>''ij''</sub> 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.
 
The assembly current size is <math>N=\sum_{i=1}^{N_G}n_i</math>. 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.<br>
The model is subjected to a [[Monte Carlo algorithm]] based simulations, using [[Gillespie algorithm]].
 
==Selection in GARD==
In 2010, [[Eors Szathmary]] and collaborators have chosen GARD as an archetypal [[metabolism-first]] realization.<ref>{{cite journal |last1=Vasas |first1=V. |last2=Szathmary |first2=E. |last3=Santos |first3=M. |year=2010 |title=Lack of evolvability in self-sustaining autocatalytic networks constraints metabolism-first scenarios for the origin of life |journal=Proc Natl Acad Sci U S A |volume=107 |issue=4 |pages=1470–1475 |doi=10.1073/pnas.0912628107 |url=http://www.pnas.org/content/107/4/1470.abstract}}</ref> They have introduced selection coefficient into the model, which increase or decrease the growth rate of assemblies, depending on how similar or dis-similar they are to a given target. They found that the ranking of the assemblies are un-affected by the selection pressure, and concluded that GARD does not exhibit Darwinian evolution.<br>
In 2012 it was shown that this criticism is errorneous and was refuted.<ref>{{cite journal |last1=Markovitch |first1=O. |last2=Lancet |first2=D. |year=2012 |title=Excess Mutual Catalysis Is Required for Effective Evolvability |journal=Artificial Life |volume=18 |issue=3 |pages=243–266 |doi=10.1162/artl_a_00064 |url=http://www.mitpressjournals.org/doi/abs/10.1162/artl_a_00064 |pmid=22662913}}</ref> Two major drawbacks of the 2010 paper were: (1) they have focused on a general assembly and not on a composome or compotype (faithfuly replicating and [[quasispecies]], respectively); (2) they have performed only a single, random, simulation to test the selectability.
 
==External links==
*Doron Lancet [http://www.weizmann.ac.il/molgen/members/lancet.html homepage] at [[Weizmann Institute of Science]], who is the inventor of GARD.
*Origin of life ([http://ool.weizmann.ac.il OOL]) at the Weizmann Institute.
 
==References==
{{Reflist}}
 
[[Category:Evolutionary biology]]
[[Category:Origin of life]]
[[Category:Stochastic simulation]]

Revision as of 10:51, 22 May 2013

In evolutionary biology, the 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 of an 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 GARD model

The composition vector of an assembly is written as: v=n1nNG. Where n1nNG are the molecular counts of lipid type i within the assembly, and NG is how many different lipid types exist (repertiore size).

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

dnidt=(kfρiNkbni)(1+j=1NGβijnjN)

kf and kb are the basel 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.

The assembly current size is N=i=1NGni. 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 in GARD

In 2010, Eors Szathmary and collaborators have chosen GARD as an archetypal metabolism-first realization.[3] They have introduced selection coefficient into the model, which increase or decrease the growth rate of assemblies, depending on how similar or dis-similar they are to a given target. They found that the ranking of the assemblies are un-affected by the selection pressure, and concluded that GARD does not exhibit Darwinian evolution.
In 2012 it was shown that this criticism is errorneous and was refuted.[4] Two major drawbacks of the 2010 paper were: (1) they have focused on a general assembly and not on a composome or compotype (faithfuly replicating and quasispecies, respectively); (2) they have performed only a single, random, simulation to test the selectability.

External links

References

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