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In
evolutionary biology Evolutionary biology is the subfield of biology that studies the evolutionary processes (natural selection, common descent, speciation) that produced the diversity of life on Earth. It is also defined as the study of the history of life fo ...
, the GARD (Graded Autocatalysis Replication Domain) model is a general kinetic model for homeostatic-growth and fission of compositional-assemblies, with specific application towards
lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
s. In the context of
abiogenesis In biology, abiogenesis (from a- 'not' + Greek bios 'life' + genesis 'origin') or the origin of life is the natural process by which life has arisen from non-living matter, such as simple organic compounds. The prevailing scientific hypothes ...
, the lipid-world suggests assemblies of simple molecules, such as
lipids Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids include ...
, can store and propagate information, thus undergo
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
. 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 Ribonucleic acid (RNA) is a polymeric molecule essential in various biological roles in coding, decoding, regulation and expression of genes. RNA and deoxyribonucleic acid ( DNA) are nucleic acids. Along with lipids, proteins, and carbohydra ...
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

The composition vector of an assembly is written as: v=n_1\cdots n_. Where n_1\cdots n_ 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: : \frac = (k_f \rho_i N-k_b n_i) \left(1+\sum_^\beta_ \frac\right) k_f and k_b are the basel forward (joining) and backward (leaving) rate constants, ''β''''ij'' 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 In the context of network theory, a complex network is a graph (network) with non-trivial topological features—features that do not occur in simple networks such as lattices or random graphs but often occur in networks representing real s ...
. The assembly current size is N=\sum_^n_i. 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 In computing, a Monte Carlo algorithm is a randomized algorithm whose output may be incorrect with a certain (typically small) probability. Two examples of such algorithms are Karger–Stein algorithm and Monte Carlo algorithm for minimum Feedba ...
based simulations, using
Gillespie algorithm In probability theory, the Gillespie algorithm (or the Doob-Gillespie algorithm or ''Stochastic Simulation Algorithm'', the SSA) generates a statistically correct trajectory (possible solution) of a stochastic equation system for which the reaction ...
.


Selection

In 2010, Eors Szathmary and collaborators have chosen GARD as an archetypal metabolism-first realization. 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 erroneous and was refuted. Two major drawbacks of the 2010 paper were: (1) they have focused on a general assembly and not on a composome or compotype (faithfully replicating and
quasispecies The quasispecies model is a description of the process of the Darwinian evolution of certain self-replicating entities within the framework of physical chemistry. A quasispecies is a large group or "cloud" of related genotypes that exist in an en ...
, respectively); (2) they have performed only a single, random, simulation to test the selectability.


Quasispecies

The
quasispecies model The quasispecies model is a description of the process of the Darwinian evolution of certain self-replicating entities within the framework of physical chemistry. A quasispecies is a large group or "cloud" of related genotypes that exist in an env ...
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 Error catastrophe refers to the cumulative loss of genetic information in a lineage of organisms due to high mutation rates. The mutation rate above which error catastrophe occurs is called the error threshold. Both terms were coined by Manfred ...
, similarly to classical quasispecies such as RNA viruses.


See also

*
Abiogenesis In biology, abiogenesis (from a- 'not' + Greek bios 'life' + genesis 'origin') or the origin of life is the natural process by which life has arisen from non-living matter, such as simple organic compounds. The prevailing scientific hypothes ...
*
Protocell A protocell (or protobiont) is a self-organized, endogenously ordered, spherical collection of lipids proposed as a stepping-stone toward the origin of life. A central question in evolution is how simple protocells first arose and how they could ...


References


External links

*GARD10 MATLAB code (see Markovitch and Lancet, 2012): https://github.com/ModelingOriginsofLife/GARD *Doron Lance
homepage
at
Weizmann Institute of Science The Weizmann Institute of Science ( he, מכון ויצמן למדע ''Machon Vaitzman LeMada'') is a public research university in Rehovot, Israel, established in 1934, 14 years before the State of Israel. It differs from other Israeli unive ...
, who is the inventor of GARD. *Origin of life
OOL
{{Webarchive, url=https://web.archive.org/web/20070710121329/http://ool.weizmann.ac.il/ , date=2007-07-10 ) at the Weizmann Institute. Evolutionary biology Origin of life Stochastic simulation