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[[File:DNA Transposon.png|thumb|(A) [[Transposable element|Transposable elements]] are flanked by inverted [[Tandem repeat|tandem repeats]] (TIRs). (B) [[Transposase|Transposases]] cleave the transposable element at the TIRs. The free transposable element inserts into another part of the [[genome]].|alt=|260x260px]]

The '''mobilome''' is the entire set of [[mobile genetic elements]] in a [[genome]]. Mobilomes are found in [[Eukaryote|eukaryotes]],<ref>{{cite journal | vauthors = Hurst GD, Werren JH | title = The role of selfish genetic elements in eukaryotic evolution | journal = Nature Reviews. Genetics | volume = 2 | issue = 8 | pages = 597–606 | date = August 2001 | pmid = 11483984 | doi = 10.1038/35084545 | s2cid = 2715605 }}</ref> [[Prokaryote|prokaryotes]],<ref>{{cite journal | vauthors = Toussaint A, Merlin C | title = Mobile elements as a combination of functional modules | journal = Plasmid | volume = 47 | issue = 1 | pages = 26–35 | date = January 2002 | pmid = 11798283 | doi = 10.1006/plas.2001.1552 }}</ref> and [[Virus|viruses]].<ref>{{cite journal | vauthors = Miller DW, Miller LK | title = A virus mutant with an insertion of a copia-like transposable element | journal = Nature | volume = 299 | issue = 5883 | pages = 562–4 | date = October 1982 | pmid = 6289125 | doi = 10.1038/299562a0 | bibcode = 1982Natur.299..562M | s2cid = 4275018 }}</ref> The compositions of mobilomes differ among lineages of life, with [[Transposable element|transposable elements]] being the major mobile elements in eukaryotes, and [[Plasmid|plasmids]] and [[prophages]] being the major types in prokaryotes.<ref name=":0">{{cite book | vauthors = Siefert JL | chapter = Defining the Mobilome | series = Methods in Molecular Biology | volume = 532 | pages = 13–27 | date = 2009 | pmid = 19271177 | doi = 10.1007/978-1-60327-853-9_2 | publisher = Humana Press | isbn = 978-1-60327-853-9 | title = Horizontal Gene Transfer: Genomes in Flux | editor-first = Maria Boekels | editor-last = Gogarten | editor2-first = Johann Peter | editor2-last = Gogarten | editor3-first = Lorraine C. | editor3-last = Olendzenski }}</ref> [[Virophage|Virophages]] contribute to the viral mobilome.<ref name=":3">{{cite journal | vauthors = Bekliz M, Colson P, La Scola B | title = The Expanding Family of Virophages | journal = Viruses | volume = 8 | issue = 11 | page = 317 | date = November 2016 | pmid = 27886075 | pmc = 5127031 | doi = 10.3390/v8110317 | doi-access = free }}</ref>

== Mobilome in eukaryotes ==
[[Transposable element]]s are elements that can move about or propagate within the genome, and are the major constituents of the [[eukaryote|eukaryotic]] mobilome.<ref name=":0" /> Transposable elements can be regarded as genetic [[Parasitism|parasites]] because they exploit the host [[Cell (biology)|cell's]] [[Transcription (biology)|transcription]] and [[Translation (biology)|translation]] mechanisms to extract and insert themselves in different parts of the genome, regardless of the [[Phenotype|phenotypic]] effect on the host.<ref>{{cite journal | vauthors = Wallau GL, Ortiz MF, Loreto EL | title = Horizontal transposon transfer in eukarya: detection, bias, and perspectives | journal = Genome Biology and Evolution | volume = 4 | issue = 8 | pages = 689–99 | date = 2012 | pmid = 22798449 | pmc = 3516303 | doi = 10.1093/gbe/evs055 }}</ref>

Eukaryotic transposable elements were first discovered in [[maize]] (''Zea mays'') in which [[Kernel (seed)|kernels]] showed a dotted color pattern.<ref>{{cite journal | vauthors = Coe EH | title = The origins of maize genetics | journal = Nature Reviews. Genetics | volume = 2 | issue = 11 | pages = 898–905 | date = November 2001 | pmid = 11715045 | doi = 10.1038/35098524 | s2cid = 5498836 }}</ref> [[Barbara McClintock]] described the maize [[Ac/Ds transposable controlling elements|Ac/Ds system]] in which the Ac [[Locus (genetics)|locus]] promotes the excision of the Ds locus from the genome, and excised Ds elements can [[Mutation|mutate]] [[Gene|genes]] responsible for [[Biological pigment|pigment]] production by [[Insertional mutagenesis|inserting]] into their [[Coding region|coding regions]].<ref>{{cite journal | vauthors = McClintock B | title = The origin and behavior of mutable loci in maize | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 36 | issue = 6 | pages = 344–55 | date = June 1950 | pmid = 15430309 | pmc = 1063197 | doi = 10.1073/pnas.36.6.344 | bibcode = 1950PNAS...36..344M | doi-access = free }}</ref>

Other examples of transposable elements include: [[Saccharomyces cerevisiae|yeast]] (''Saccharomyces cerevisiae'') [[Ty5 retrotransposon|Ty elements]], a [[retrotransposon]] which encodes a [[reverse transcriptase]] to convert its [[Transcription (biology)|mRNA transcript]] into DNA which can then insert into other parts of the genome;<ref>{{cite journal | vauthors = Mellor J, Malim MH, Gull K, Tuite MF, McCready S, Dibbayawan T, Kingsman SM, Kingsman AJ | display-authors = 6 | title = Reverse transcriptase activity and Ty RNA are associated with virus-like particles in yeast | journal = Nature | volume = 318 | issue = 6046 | pages = 583–6 | date = December 1985 | pmid = 2415827 | doi = 10.1038/318583a0 | bibcode = 1985Natur.318..583M | s2cid = 4314282 }}</ref><ref>{{cite journal | vauthors = Garfinkel DJ, Boeke JD, Fink GR | title = Ty element transposition: reverse transcriptase and virus-like particles | journal = Cell | volume = 42 | issue = 2 | pages = 507–17 | date = September 1985 | pmid = 2411424 | doi = 10.1016/0092-8674(85)90108-4 | s2cid = 35750065 }}</ref> and [[Drosophila melanogaster|fruit fly]] (''Drosophila melanogaster'') [[P element|P-elements]], which randomly inserts into the genome to cause mutations in [[Germ cell|germ line cells]], but not in [[somatic cells]].<ref>{{cite journal | vauthors = Laski FA, Rio DC, Rubin GM | title = Tissue specificity of Drosophila P element transposition is regulated at the level of mRNA splicing | journal = Cell | volume = 44 | issue = 1 | pages = 7–19 | date = January 1986 | pmid = 3000622 | doi = 10.1016/0092-8674(86)90480-0 | s2cid = 18364777 }}</ref>

== Mobilome in prokaryotes ==
[[File:Conjugation.svg|left|thumb|alt=|[[Bacterial conjugation]]. (1) Production of [[pilus]]. (2) Pilus connects two [[bacteria]]. (3) One strand of [[plasmid]] [[DNA]] moves into the recipient. (4) Both bacteria contain identical plasmids.]]
Plasmids were discovered in the 1940s as genetic materials outside of [[Bacteria|bacterial]] [[Chromosome|chromosomes]].<ref>{{cite journal | vauthors = Sonneborn TM | title = The cytoplasm in heredity | journal = Heredity | volume = 4 | issue = 1 | pages = 11–36 | date = April 1950 | pmid = 15415003 | doi = 10.1038/hdy.1950.2 | doi-access = free }}</ref> Prophages are genomes of [[bacteriophage]]s (a type of virus) that are inserted into bacterial chromosomes; prophages can then be spread to other bacteria through the [[lytic cycle]] and [[lysogenic cycle]] of [[viral replication]].<ref name=":1">{{cite journal | vauthors = Bertani G | title = Lysogenic versus lytic cycle of phage multiplication | journal = Cold Spring Harbor Symposia on Quantitative Biology | volume = 18 | pages = 65–70 | date = 1953-01-01 | pmid = 13168970 | doi = 10.1101/SQB.1953.018.01.014 }}</ref>

While transposable elements are also found in prokaryotic genomes,<ref>{{cite journal | vauthors = Campbell A, Berg DE, Botstein D, Lederberg EM, Novick RP, Starlinger P, Szybalski W | title = Nomenclature of transposable elements in prokaryotes | journal = Gene | volume = 5 | issue = 3 | pages = 197–206 | date = March 1979 | pmid = 467979 | doi = 10.1016/0378-1119(79)90078-7 }}</ref> the most common mobile genetic elements in the prokaryotic genome are [[Plasmid|plasmids]] and [[prophages]].<ref name=":0" />

Plasmids and prophages can move between genomes through [[bacterial conjugation]], allowing [[horizontal gene transfer]].<ref>{{cite journal | vauthors = Juhas M | title = Horizontal gene transfer in human pathogens | journal = Critical Reviews in Microbiology | volume = 41 | issue = 1 | pages = 101–8 | date = February 2015 | pmid = 23862575 | doi = 10.3109/1040841X.2013.804031 | s2cid = 5193869 | url = https://www.repository.cam.ac.uk/bitstream/1810/244728/1/Juhas%20embargo%20text.pdf }}</ref> Plasmids often carry genes that are responsible for bacterial [[Antimicrobial resistance|antibiotic resistance]]; as these plasmids replicate and pass from one genome to another, the whole bacterial [[Population genetics|population]] can quickly [[Adaptation|adapt]] to the [[antibiotic]].<ref>{{cite journal | vauthors = Harrison E, Brockhurst MA | title = Plasmid-mediated horizontal gene transfer is a coevolutionary process | journal = Trends in Microbiology | volume = 20 | issue = 6 | pages = 262–7 | date = June 2012 | pmid = 22564249 | doi = 10.1016/j.tim.2012.04.003 | url = http://eprints.whiterose.ac.uk/75385/1/Harrison_Brockhurst_Postprint.pdf }}</ref><ref>{{cite journal | vauthors = Gillings MR | title = Evolutionary consequences of antibiotic use for the resistome, mobilome and microbial pangenome | journal = Frontiers in Microbiology | volume = 4 | page = 4 | date = 2013 | pmid = 23386843 | pmc = 3560386 | doi = 10.3389/fmicb.2013.00004 | doi-access = free }}</ref> Prophages can loop out of bacterial chromosomes to produce bacteriophages that go on to infect other bacteria with the prophages; this allows prophages to propagate quickly among the bacterial population, to the harm of the bacterial host.<ref name=":1" />

== Mobilome in viruses ==
Discovered in 2008 in a strain of ''Acanthamoeba castellanii [[Mimiviridae|mimivirus]]'',<ref name=":2">{{cite journal | vauthors = La Scola B, Desnues C, Pagnier I, Robert C, Barrassi L, Fournous G, Merchat M, Suzan-Monti M, Forterre P, Koonin E, Raoult D | display-authors = 6 | title = The virophage as a unique parasite of the giant mimivirus | journal = Nature | volume = 455 | issue = 7209 | pages = 100–4 | date = September 2008 | pmid = 18690211 | doi = 10.1038/nature07218 | bibcode = 2008Natur.455..100L | s2cid = 4422249 }}</ref> [[Virophage|virophages]] are an element of the virus mobilome.<ref name=":3" /> Virophages are viruses that [[Viral replication|replicate]] only when host cells are co-infected with [[Helper virus|helper viruses]].<ref name=":4">{{cite journal | vauthors = Claverie JM, Abergel C | title = Mimivirus and its virophage | journal = Annual Review of Genetics | volume = 43 | issue = 1 | pages = 49–66 | date = 2009 | pmid = 19653859 | doi = 10.1146/annurev-genet-102108-134255 }}</ref> Following co-infection, helper viruses exploit the host cell's transcription/translation machinery to produce their own machinery; virophages replicate through the machinery of either the host cell or the viruses.<ref name=":4" /> The replication of virophages can negatively impact the replication of helper viruses.<ref name=":2" /><ref>{{cite journal | vauthors = Duponchel S, Fischer MG | title = Viva lavidaviruses! Five features of virophages that parasitize giant DNA viruses | journal = PLOS Pathogens | volume = 15 | issue = 3 | article-number = e1007592 | date = March 2019 | pmid = 30897185 | pmc = 6428243 | doi = 10.1371/journal.ppat.1007592 | doi-access = free }}</ref>

[[Sputnik virophage|Sputnik]]<ref name=":2" /><ref>{{cite journal | vauthors = Sun S, La Scola B, Bowman VD, Ryan CM, Whitelegge JP, Raoult D, Rossmann MG | title = Structural studies of the Sputnik virophage | journal = Journal of Virology | volume = 84 | issue = 2 | pages = 894–7 | date = January 2010 | pmid = 19889775 | pmc = 2798384 | doi = 10.1128/JVI.01957-09 }}</ref> and [[mavirus]]<ref>{{cite journal | vauthors = Fischer MG, Hackl T | title = Host genome integration and giant virus-induced reactivation of the virophage mavirus | journal = Nature | volume = 540 | issue = 7632 | pages = 288–291 | date = December 2016 | pmid = 27929021 | doi = 10.1038/nature20593 | bibcode = 2016Natur.540..288F | s2cid = 4458402 | url = https://www.biorxiv.org/content/biorxiv/early/2016/10/18/068312.full.pdf }}</ref> are examples of virophages.

== References ==
{{reflist}}

{{Genetic recombination}}
{{Self-replicating organic structures}}

[[Category:Genetics]]

Mobilome - История изменений

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