View source for Mobile genetic elements

{{short description|DNA sequence whose position in the genome is variable}}
[[File:Components of the Human Genome.jpg|thumb|upright=1.15|DNA transposons, LTR retrotransposons, SINEs, and LINEs make up a majority of the human genome.]]
'''Mobile genetic elements''' ('''MGEs'''), sometimes called '''selfish genetic elements''',<ref name="Moreira_2009" /> are a type of [[genetic material]] that can move around within a genome, or that can be transferred from one species or replicon to another. MGEs are found in all organisms. In humans, approximately 50% of the genome are thought to be MGEs.<ref>{{cite book | vauthors = Mu X, Ahmad S, Hur S | title = Endogenous Retroelements and the Host Innate Immune Sensors | series = Advances in Immunology | volume = 132 | pages = 47–69 | year = 2016 | pmid = 27769507 | pmc = 5135014 | doi = 10.1016/bs.ai.2016.07.001 | isbn = 9780128047972 }}</ref> MGEs play a distinct role in evolution. Gene duplication events can also happen through the mechanism of MGEs. MGEs can also cause mutations in protein coding regions, which alters the protein functions. These mechanisms can also rearrange genes in the host genome generating variation. These mechanisms can increase fitness by gaining new or additional functions. An example of MGEs in evolutionary context are that virulence factors and [[antimicrobial resistance|antibiotic resistance]] genes of MGEs can be transported to share genetic code with neighboring bacteria. However, MGEs can also decrease fitness by introducing disease-causing alleles or mutations.<ref>{{cite journal | vauthors = Singh PK, Bourque G, Craig NL, Dubnau JT, Feschotte C, Flasch DA, Gunderson KL, Malik HS, Moran JV, Peters JE, Slotkin RK, Levin HL | display-authors = 6 | title = Mobile genetic elements and genome evolution 2014 | journal = Mobile DNA | volume = 5 | pages = 26 | date = 2014-11-18 | pmid = 30117500 | pmc = 4363357 | doi = 10.1186/1759-8753-5-26 | doi-access = free }}</ref> The set of MGEs in an organism is called a [[mobilome]], which is composed of a large number of [[plasmids]], [[transposons]] and [[viruses]].<ref name="wolf" />[[File:PBLU plasmid map.jpg|thumb|pBLU is a 5437bp vector plasmid. This vector contains the origin of replication sequence, the restriction enzyme cut site, lacZ gene, and an ampicillin resistance gene.]]

==Types==
* '''[[Plasmids]]''': These are generally circular extrachromosomal [[DNA]] molecules that replicate and are transmitted independently from chromosomal DNA. These molecules are present in [[prokaryotes]] ([[bacteria]] and [[archaea]]) and sometimes in eukaryotic organisms such as [[yeast]]. Fitness of a plasmid is determined by its mobility. The first factor of plasmid fitness is its ability to replicate DNA. The second fitness factor is a plasmid's ability to horizontally transfer. Plasmids during their cycle carry genes from one organism to another through a process called [[Bacterial conjugation|conjugation]]. Plasmids usually contain a set of mobility genes that are necessary for conjugation. Some plasmids employ membrane associated mating pair formation (MPF). A plasmid containing its own MPF genes is considered to be self transmissible or conjugative.<ref name="Smillie-2010">{{cite journal | vauthors = Smillie C, Garcillán-Barcia MP, Francia MV, Rocha EP, de la Cruz F | title = Mobility of plasmids | journal = Microbiology and Molecular Biology Reviews | volume = 74 | issue = 3 | pages = 434–452 | date = September 2010 | pmid = 20805406 | pmc = 2937521 | doi = 10.1128/MMBR.00020-10 }}</ref> Plasmids can be further divided into mobilizable and non-mobilizable classes. Plasmids that use other genetic element MFPs in the cell are mobilizable. Plasmids that are not mobilizable but spread by transduction or transformation are termed non-mobilizable.<ref name="Smillie-2010" /> Plasmids can often inject genes that make bacteria [[Antimicrobial resistance|resistant to antibiotics]].<ref>{{cite book |chapter-url= https://books.google.com/books?id=a4lrPKQWjtAC&pg=PA21 |title= The Biology of Plasmids |publisher= Wiley-Blackwell|edition= First  |year= 1996 | vauthors = Summers D |chapter= Chapter 1 – The Function and Organization of Plasmids |pages= 21–22 |isbn= 978-0632034369}}</ref><ref name="Smillie-2010" />

* [[Cloning vectors]]: These are types of hybrid plasmids with [[bacteriophages]], used to transfer and replicate DNA . Fragments of DNA can be inserted by [[recombinant DNA]] techniques. A viable vector must be able to replicate together with the DNA fragments it carries. These vectors can contain desired genes for insertion into an organism's genome.  Examples are [[cosmids]] and [[phagemid]]s.<ref>{{cite book |vauthors=Glick BR, Pasternak JJ |year=2005 |title=Molecular Biotechnology Principles and Applications of Recombinant DNA |edition=3rd |publisher=ASM Press |url=https://books.google.com/books?id=Wz3CtTBe9aUC&pg=PA86 |isbn=9781555816124}}</ref>
[[File:Bacterial mobile elements.svg|thumb|upright=2 |Examples of mobile genetic elements in the cell (left) and the ways they can be acquired (right)|center]]
[[File:Cut and Paste mechanism of transposition.svg|thumb|upright=1.5|Transposition of target sequence into recombination site in DNA by Transposase. Replication of the transposable sequence starts to occur when transposase cuts single strands on opposite sides of the dsDNA. The replication is completed in the transposon complex and excised to target sequence for recombination.]]
* [[Transposons]]: These are [[DNA sequences]] that can move and replicate in different parts of a cell's [[genome]]. Also called "jumping genes", they can be transferred horizontally between organisms that live in [[symbiosis]]. Transposons are present in all [[Organism|living things]] and in [[giant viruses]].<ref>{{cite book |vauthors=Makałowski W, Gotea V, Pande A, Makałowska I |chapter=Transposable Elements: Classification, Identification, and Their Use as a Tool for Comparative Genomics |title=Evolutionary Genomics |veditors=Anisimova M |series=Methods in Molecular Biology |place=New York, NY |publisher=Humana |volume=1910 |pages=185–186 |date=2019 |pmid=31278665 |doi=10.1007/978-1-4939-9074-0_6 |isbn=978-1-4939-9074-0 |doi-access=free |s2cid=195814061}}</ref>

* [[DNA transposons]]: These are transposons that move directly from one position to another in the genome using a [[transposase]] to cut and stick at another [[Locus (genetics)|locus]].<ref>{{cite journal |vauthors=Muñoz-López M, García-Pérez JL |title=DNA transposons: nature and applications in genomics |journal=Current Genomics |volume=11 |issue=2 |pages=115–128 |date=April 2010 |pmid=20885819 |pmc=2874221 |doi=10.2174/138920210790886871}}</ref> These genetic elements are cleaved at four single stranded sites in DNA by transposase. In order to achieve max stability of the intermediate transposon, one single strand cleavage at the target DNA occurs. Simultaneously the donor strand is ligated to the target strand after cleavage leaving a single strand overhang on either end of the target sequence. These sites usually contain a 5 to 9 base pair overhang that can create a cohesive end.<ref name="Shapiro-1979" /> Transposase then holds the sequence in a crossed formation and ligates the donor strand to the target strand. The structure formed by the duplex of DNA and transposase in replicative transposons is known as the Shapiro Intermediate.<ref>{{Cite book | vauthors = Bushman F |url=https://www.worldcat.org/oclc/47283049 |title=Lateral DNA transfer : mechanisms and consequences |date=2002 |publisher=Cold Spring Harbor Laboratory Press |isbn=0-87969-603-6 |location=Cold Spring Harbor, NY |oclc=47283049}}</ref> The 5 to 9 base pair overhang is left on either side of the target sequence allowing it to join to its target sequence in either orientation. The sequence of these overhangs can determine joining orientation.<ref name="Shapiro-1979">{{cite journal |vauthors=Shapiro JA |title=Molecular model for the transposition and replication of bacteriophage Mu and other transposable elements |journal=Proceedings of the National Academy of Sciences of the United States of America |volume=76 |issue=4 |pages=1933–1937 |date=April 1979 |pmid=287033 |doi=10.1073/pnas.76.4.1933 |pmc=383507 |doi-access=free |bibcode=1979PNAS...76.1933S}}</ref> Before site specific recombination can occur, the oligonucleotide ends must be filled. The ligation of these ends generates a replication fork at each end of the transposable element. The single strand displacement causes synthesis from the un-ligated 3' hydroxyl group to form long single stranded sections adjacent to the 5' end. Therefore, the opposite strand is sequenced discontinuously as both replication forks approach the center of the transposable element.  This results in two recombinant duplexes containing the semi conserved transposable element flanked by the previous 5 to 9 base pair overhang. Site specific reciprocal recombination takes place between the two transposable elements facilitated by proteins. This reciprocal replication overlaps in time and occurs between duplicated segments of the replication element before replication is completed.<ref name="Shapiro-1979" /> The target molecule as a result contains the inserted element flanked by the 5 to 9 base pair sequences. Transposition of these elements duplicates the transposition element leaving a transposition element in its original location and a new transposon at the reciprocal replication site. In doing so, organisms total base pairs in their genomes are increased. Transposition occurrences increase over time and as organisms age.

[[File:Retrotransposons.png|thumb|upright=1.35|Retrotransposon mechanism that uses reverse transcriptase to change mRNA transposon back into DNA for integration.]]

* [[Retrotransposons]]: These are transposons that move in the genome, being transcribed into [[RNA]] and later into [[DNA]] by [[reverse transcriptase]]. Many retrotransposons also exhibit [[replicative transposition]]. Retrotransposons are present exclusively in [[eukaryotes]].<ref>{{cite journal |vauthors=Richardson SR, Doucet AJ, Kopera HC, Moldovan JB, Garcia-Perez JL, Moran JV |title=The Influence of LINE-1 and SINE Retrotransposons on Mammalian Genomes |journal=Microbiology Spectrum |volume=3 |issue=2 |pages=1165–1208 |date=April 2015 |pmid=26104698 |pmc=4498412 |doi=10.1128/microbiolspec.mdna3-0061-2014 |isbn=9781555819200}}</ref> Retrotransposons consist of two major types, [[long terminal repeat]]s (LTRs) and Non-LTR transposons. Non-LTR transposons can be further classified into [[Long interspersed nuclear element]] (LINEs) and [[Short interspersed nuclear element]] (SINEs).<ref>{{cite book |vauthors=Makałowski W, Gotea V, Pande A, Makałowska I |chapter=Transposable Elements: Classification, Identification, and Their Use as a Tool for Comparative Genomics |title=Evolutionary Genomics |date=2019 |series=Methods in Molecular Biology |volume=1910 |pages=177–207 |veditors=Anisimova M |place=New York, NY |publisher=Humana |language=en |doi=10.1007/978-1-4939-9074-0_6 |pmid=31278665 |isbn=978-1-4939-9074-0 |s2cid=195814061}}</ref> These retrotransposons are regulated by a family of short non-coding RNAs termed as PIWI [P-element induced wimpy testis]-interacting RNAs (piRNAs).<ref name="Monga">{{cite journal |vauthors=Monga I, Banerjee I |title=Computational Identification of piRNAs Using Features Based on RNA Sequence, Structure, Thermodynamic and Physicochemical Properties |journal=Current Genomics |volume=20 |issue=7 |pages=508–518 |date=November 2019 |doi=10.2174/1389202920666191129112705 |pmid=32655289 |pmc=7327968}}</ref> piRNA is a recently discovered class of ncRNAs, which are in the length range of ~24-32 nucleotides. Initially, piRNAs were described as repeat-associated siRNAs (rasiRNAs) because of their origin from the repetitive elements such as transposable sequences of the genome. However, later it was identified that they acted via PIWI-protein. In addition to having a role in the suppression of genomic transposons, various roles of piRNAs have been recently reported like regulation of 3’ UTR of protein-coding genes via RNAi,  transgenerational epigenetic inheritance to convey a memory of past transposon  activity, and RNA-induced epigenetic silencing.<ref name="Monga" />
* [[Integrons]]: These are [[gene cassette]]s that usually carry antibiotic resistance genes to bacterial plasmids and transposons.<ref>{{cite journal |doi=10.1023/A:1015361704475 |year=2002 |vauthors=Kovalevskaya NP |journal=Molecular Biology |volume=36 |issue=2 |pages=196–201 |title=Mobile Gene Cassettes and Integrons |s2cid=2078235}}</ref>
* [[Introns]]: [[Group I catalytic intron|Group I]] and [[Group II intron|II introns]] are [[nucleotide]] sequences with catalytic activity that are part of host transcripts and act as ribozymes that can invade genes that encode [[tRNA]], [[rRNA]], and [[proteins]]. They are present in all cellular organisms and viruses.<ref>{{cite journal |vauthors=Hausner G, Hafez M, Edgell DR |title=Bacterial group I introns: mobile RNA catalysts |journal=Mobile DNA |volume=5 |issue=1 |pages=8 |date=March 2014 |pmid=24612670 |pmc=3984707 |doi=10.1186/1759-8753-5-8 |doi-access=free }}</ref> 
* Introners: Sequences similar to transposons that can jump in the genome leaving new introns where they were, they have been pointed as a possible mechanism of intron gain in the evolution of eukaryotes where they are present in at least 5% of all species, specially in the aquatic taxa due possibly to [[Horizontal gene transfer in evolution|horizontal gene transfer]] that occurs more frequently in these animals.<ref>{{cite journal | vauthors = Gozashti L, Roy SW, Thornlow B, Kramer A, Ares M, Corbett-Detig R | title = Transposable elements drive intron gain in diverse eukaryotes | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 119 | issue = 48 | pages = e2209766119 | date = November 2022 | pmid = 36417430 | pmc = 9860276 | doi = 10.1073/pnas.2209766119 | doi-access = free | bibcode = 2022PNAS..11909766G }}</ref><ref>{{Cite web | vauthors = Buehler J |date=2023-03-30 |title=How a DNA 'Parasite' May Have Fragmented Our Genes |url=https://www.quantamagazine.org/how-a-dna-parasite-may-have-fragmented-our-genes-20230330/ |access-date=2023-03-31 |website=Quanta Magazine |language=en}}</ref> They were first described in 2009 in the unicellular green algae [[micromonas]].<ref>{{cite journal | vauthors = Worden AZ, Lee JH, Mock T, Rouzé P, Simmons MP, Aerts AL, Allen AE, Cuvelier ML, Derelle E, Everett MV, Foulon E, Grimwood J, Gundlach H, Henrissat B, Napoli C, McDonald SM, Parker MS, Rombauts S, Salamov A, Von Dassow P, Badger JH, Coutinho PM, Demir E, Dubchak I, Gentemann C, Eikrem W, Gready JE, John U, Lanier W, Lindquist EA, Lucas S, Mayer KF, Moreau H, Not F, Otillar R, Panaud O, Pangilinan J, Paulsen I, Piegu B, Poliakov A, Robbens S, Schmutz J, Toulza E, Wyss T, Zelensky A, Zhou K, Armbrust EV, Bhattacharya D, Goodenough UW, Van de Peer Y, Grigoriev IV | display-authors = 6 | title = Green evolution and dynamic adaptations revealed by genomes of the marine picoeukaryotes Micromonas | journal = Science | volume = 324 | issue = 5924 | pages = 268–272 | date = April 2009 | pmid = 19359590 | doi = 10.1126/science.1167222 | bibcode = 2009Sci...324..268W | s2cid = 206516961 | url = https://escholarship.org/uc/item/7c83j3jr }}</ref> 
* [[Acellular life|Viral agents]]: These are mostly infective [[acellular]] agents that replicate in cellular hosts. During their infective cycle they can carry genes from one host to another. They can also carry genes from one organism to another in case that viral agent infects more than two different species. Traditionally they are considered separate entities, but the truth is that many researchers who study their characteristics and evolution refer to them as mobile genetic elements. This is based on the fact that viral agents are simple particles or molecules that replicate and are transferred between various hosts like the remaining non-viral mobile genetic elements. According to this point of view, viruses and other viral agents should not be considered [[living beings]] and should be better conceived as mobile genetic elements. Viral agents are evolutionarily connected with various mobile genetic elements.<ref name="ictv">{{cite report | vauthors = Kuhn JH, Dolja VV, Krupovic M, Adriaenssens EM, Di Serio F, Dutilh BE, Flores R, Harrach BM, Owens B, Randles J, Rubino L | display-authors = 6 | title = Expand, amend, and emend the International Code of Virus Classification and Nomenclature (ICVCN;"the Code") and the Statutes to clearly define the remit of the ICTV | date = 2020 | doi = 10.13140/RG.2.2.26202.26565 }}</ref> These viral agents are thought to have arisen from secreted or ejected plasmids of other organisms. Transposons also provide insight into how these elements may have originally started. This theory is known as the vagrancy hypothesis proposed by [[Barbara McClintock]] in 1950.<ref name="Koonin_2021">{{cite journal | vauthors = Koonin EV, Dolja VV, Krupovic M, Kuhn JH | title = Viruses Defined by the Position of the Virosphere within the Replicator Space | journal = Microbiology and Molecular Biology Reviews | volume = 85 | issue = 4 | pages = e0019320 | date = December 2021 | pmid = 34468181 | pmc = 8483706 | doi = 10.1128/MMBR.00193-20 }}</ref><ref name="Moreira_2009">{{cite journal | vauthors = Moreira D, López-García P | title = Ten reasons to exclude viruses from the tree of life | journal = Nature Reviews. Microbiology | volume = 7 | issue = 4 | pages = 306–311 | date = April 2009 | pmid = 19270719 | doi = 10.1038/nrmicro2108 | s2cid = 3907750 }}</ref><ref>{{cite journal | vauthors = Koonin EV, Dolja VV | title = Virus world as an evolutionary network of viruses and capsidless selfish elements | journal = Microbiology and Molecular Biology Reviews | volume = 78 | issue = 2 | pages = 278–303 | date = June 2014 | pmid = 24847023 | pmc = 4054253 | doi = 10.1128/MMBR.00049-13 }}</ref><ref name="wolf">{{cite journal | vauthors = Koonin EV, Wolf YI | title = Genomics of bacteria and archaea: the emerging dynamic view of the prokaryotic world | journal = Nucleic Acids Research | volume = 36 | issue = 21 | pages = 6688–6719 | date = December 2008 | pmid = 18948295 | pmc = 2588523 | doi = 10.1093/nar/gkn668 }}</ref><ref>{{cite journal | vauthors = Rankin DJ, Rocha EP, Brown SP | title = What traits are carried on mobile genetic elements, and why? | journal = Heredity | volume = 106 | issue = 1 | pages = 1–10 | date = January 2011 | pmid = 20332804 | pmc = 3183850 | doi = 10.1038/hdy.2010.24 }}</ref>
** [[Viruses]]: These are viral agents composed of a molecule of [[genetic material]] (DNA or RNA) and with the ability to form complex particles called [[virion]]s to be able to move easily between their hosts. Viruses are present in all living things. Viral particles are manufactured by the host's replicative machinery for horizontal transfer.<ref name="ictv" /><ref name="Koonin_2021" /><ref>{{cite book |vauthors=Crawford D |year=2011 |title=Viruses: A Very Short Introduction |page=[https://archive.org/details/virusesveryshort0000craw/page/4 4] |publisher=Oxford University Press |location=New York |url=https://archive.org/details/virusesveryshort0000craw |isbn=978-0199574858}}</ref>
** [[Satellite (biology)|Satellite nucleic acids]]: These are DNA or RNA molecules, which are encapsulated as a stowaway in the virions of certain helper viruses and which depend on these to be able to replicate. Although they are sometimes considered genetic elements of their helper viruses, they are not always found within their helper viruses.<ref name="ictv" /><ref name="Koonin_2021" /><ref>{{cite web | vauthors = Briddon RW, Ghabrial S, Lin NS, Palukaitis P, Scholthof KB, Vetten HJ | title = 3 - Satellites and Other Virus-dependent Nucleic Acids - Subviral Agents - Subviral Agents (2011) |url=https://ictv.global/report_9th/subviral/Satellites-introduction |archive-url=https://web.archive.org/web/20190113062904/https://talk.ictvonline.org/ictv-reports/ictv_9th_report/sub-viral-agents-2011/w/sub_viruses/302/3-satellites-and-other-virus-dependent-nucleic-acids |archive-date=13 January 2019 |website=International Committee on Taxonomy of Viruses (ICTV) |language=en}}</ref>
** [[Viroids]]: These are viral agents that consist of small circular RNA molecules that infect and replicate in [[plants]]. These mobile genetic elements do not have a protective protein coating. Specifically, these mobile genetic elements are found in [[angiosperms]].<ref name="ictv" /><ref name="Koonin_2021" /><ref>{{cite web |vauthors=Di Serio F, Owens RA, Li SF, Matoušek J, Pallás V, Randles JW, Sano T, Verhoeven JT, Vidalakis G, Flores R |display-authors=6 |title=Viroids |date=November 2020 |publisher=International Committee on Taxonomy of Viruses (ICTV) |url=https://talk.ictvonline.org/ictv-reports/ictv_online_report/subviral-agents/w/viroids |archive-url=https://web.archive.org/web/20201202061614/https://talk.ictvonline.org/ictv-reports/ictv_online_report/subviral-agents/w/viroids |archive-date=2 December 2020}}</ref>
** [[Endogenous viral element]]: These are viral nucleic acids integrated into the genome of a cell. They can move and replicate multiple times in the host cell without causing disease or mutation. They are considered autonomous forms of transposons. Examples are [[proviruses]] and [[endogenous retroviruses]].<ref>{{cite journal |vauthors=Feschotte C, Gilbert C |title=Endogenous viruses: insights into viral evolution and impact on host biology |journal=Nature Reviews. Genetics |volume=13 |issue=4 |pages=283–296 |date=March 2012 |pmid=22421730 |s2cid=205485232 |url=https://hal.archives-ouvertes.fr/hal-00679842/file/Feschotte_2012_endegenousvirusesNRG.pdf |doi=10.1038/nrg3199}}</ref>

==Research examples==

[[CRISPR|CRISPR-Cas]] systems in bacteria and [[archaea]] are adaptive immune systems to protect against deadly consequences from MGEs. Using comparative genomic and phylogenetic analysis, researchers found that CRISPR-Cas variants are associated with distinct types of MGEs such as transposable elements. In [[CRISPR-associated transposons]], CRISPR-Cas controls transposable elements for their propagation.<ref>{{cite journal | vauthors = Peters JE, Makarova KS, Shmakov S, Koonin EV | title = Recruitment of CRISPR-Cas systems by Tn7-like transposons | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 114 | issue = 35 | pages = E7358–E7366 | date = August 2017 | pmid = 28811374 | pmc = 5584455 | doi = 10.1073/pnas.1709035114 | doi-access = free | bibcode = 2017PNAS..114E7358P }}</ref>

MGEs such as plasmids by a horizontal transmission are generally beneficial to an organism. The ability of transferring plasmids (sharing) is important in an evolutionary perspective. Tazzyman and Bonhoeffer found that fixation (receiving) of the transferred plasmids in a new organism is just as important as the ability to transfer them.<ref>{{cite journal | vauthors = Tazzyman SJ, Bonhoeffer S | title = Fixation probability of mobile genetic elements such as plasmids | journal = Theoretical Population Biology | volume = 90 | pages = 49–55 | date = December 2013 | pmid = 24080312 | doi = 10.1016/j.tpb.2013.09.012 | doi-access = free | hdl = 20.500.11850/72638 | hdl-access = free }}</ref> Beneficial rare and transferable plasmids have a higher fixation probability, whereas deleterious transferable genetic elements have a lower fixation probability because they are lethal to the host organisms.

One type of MGEs, namely the Integrative Conjugative Elements (ICEs) are central to horizontal gene transfer shaping the genomes of prokaryotes enabling rapid acquisition of novel adaptive traits.<ref>{{cite journal | vauthors = Guglielmini J, Quintais L, Garcillán-Barcia MP, de la Cruz F, Rocha EP | title = The repertoire of ICE in prokaryotes underscores the unity, diversity, and ubiquity of conjugation | journal = PLOS Genetics | volume = 7 | issue = 8 | pages = e1002222 | date = August 2011 | pmid = 21876676 | pmc = 3158045 | doi = 10.1371/journal.pgen.1002222 | doi-access = free }}</ref><ref>{{cite journal | vauthors = Wozniak RA, Waldor MK | title = Integrative and conjugative elements: mosaic mobile genetic elements enabling dynamic lateral gene flow | journal = Nature Reviews. Microbiology | volume = 8 | issue = 8 | pages = 552–563 | date = August 2010 | pmid = 20601965 | doi = 10.1038/nrmicro2382 | s2cid = 21460836 }}</ref>

As a representative example of ICEs, the ICE''Bs1'' is well-characterized for its role in the global DNA damage SOS response of [[Bacillus subtilis]]<ref>{{cite journal | vauthors = Auchtung JM, Lee CA, Garrison KL, Grossman AD | title = Identification and characterization of the immunity repressor (ImmR) that controls the mobile genetic element ICEBs1 of Bacillus subtilis | journal = Molecular Microbiology | volume = 64 | issue = 6 | pages = 1515–1528 | date = June 2007 | pmid = 17511812 | pmc = 3320793 | doi = 10.1111/j.1365-2958.2007.05748.x }}</ref> and also its potential link to the radiation and desiccation resistance of ''Bacillus pumilus'' SAFR-032 spores,<ref>{{cite journal | vauthors = Tirumalai MR, Fox GE | title = An ICEBs1-like element may be associated with the extreme radiation and desiccation resistance of Bacillus pumilus SAFR-032 spores | journal = Extremophiles | volume = 17 | issue = 5 | pages = 767–774 | date = September 2013 | pmid = 23812891 | doi = 10.1007/s00792-013-0559-z | s2cid = 8675124 }}</ref> isolated from spacecraft cleanroom facilities.<ref>{{cite journal | vauthors = Link L, Sawyer J, Venkateswaran K, Nicholson W | title = Extreme spore UV resistance of Bacillus pumilus isolates obtained from an ultraclean Spacecraft Assembly Facility | journal = Microbial Ecology | volume = 47 | issue = 2 | pages = 159–163 | date = February 2004 | pmid = 14502417 | doi = 10.1007/s00248-003-1029-4 | bibcode = 2004MicEc..47..159L | s2cid = 13416635 }}</ref><ref>{{cite journal | vauthors = Newcombe DA, Schuerger AC, Benardini JN, Dickinson D, Tanner R, Venkateswaran K | title = Survival of spacecraft-associated microorganisms under simulated martian UV irradiation | journal = Applied and Environmental Microbiology | volume = 71 | issue = 12 | pages = 8147–8156 | date = December 2005 | pmid = 16332797 | pmc = 1317311 | doi = 10.1128/AEM.71.12.8147-8156.2005 | bibcode = 2005ApEnM..71.8147N }}</ref><ref>{{cite journal | vauthors = Kempf MJ, Chen F, Kern R, Venkateswaran K | title = Recurrent isolation of hydrogen peroxide-resistant spores of Bacillus pumilus from a spacecraft assembly facility | journal = Astrobiology | volume = 5 | issue = 3 | pages = 391–405 | date = June 2005 | pmid = 15941382 | doi = 10.1089/ast.2005.5.391 | bibcode = 2005AsBio...5..391K }}</ref>

Transposition by transposable elements is mutagenic. Thus, organisms have evolved to repress the transposition events, and failure to repress the events causes cancers in somatic cells. Cecco et al. found that during early age transcription of retrotransposable elements are minimal in mice, but in advanced age the transcription level increases.<ref>{{cite journal | vauthors = De Cecco M, Criscione SW, Peterson AL, Neretti N, Sedivy JM, Kreiling JA | title = Transposable elements become active and mobile in the genomes of aging mammalian somatic tissues | journal = Aging | volume = 5 | issue = 12 | pages = 867–883 | date = December 2013 | pmid = 24323947 | pmc = 3883704 | doi = 10.18632/aging.100621 }}</ref> This age-dependent expression level of transposable elements is reduced by calorie restriction diet. Replication of transposable elements often results in repeated sequences being added into the genome. These sequences are often non coding but can interfere with coding sequences of DNA. Though mutagenetic by nature, transposons increase the genome of an organism that they transpose into. More research should be conducted into how these elements may serve as a rapid adaptation tool employed by organisms to generate variability. Many transposition elements are dormant or require activation. should also be noted that current values for coding sequences of DNA would be higher if transposition elements that code for their own transposition machinery were considered  as coding sequences.

Some others researched examples include [[Maverick (genetics)|Mavericks]],<ref>{{cite journal | vauthors = Barreat JG, Katzourakis A | title = Phylogenomics of the Maverick Virus-Like Mobile Genetic Elements of Vertebrates | journal = Molecular Biology and Evolution | volume = 38 | issue = 5 | pages = 1731–1743 | date = May 2021 | pmid = 33481003 | pmc = 8097293 | doi = 10.1093/molbev/msaa291 | veditors = Battistuzzi FU }}</ref><ref>{{cite journal | vauthors = Widen SA, Bes IC, Koreshova A, Pliota P, Krogull D, Burga A | title = Virus-like transposons cross the species barrier and drive the evolution of genetic incompatibilities | journal = Science | volume = 380 | issue = 6652 | pages = eade0705 | date = June 2023 | pmid = 37384706 | doi = 10.1126/science.ade0705 | s2cid = 250645873 }}</ref><ref name="Bolakhe_2023">{{Cite web | vauthors = Bolakhe S |date=2023-08-03 |title=Selfish, Virus-Like DNA Can Carry Genes Between Species |url=https://www.quantamagazine.org/selfish-virus-like-dna-can-carry-genes-between-species-20230803/ |access-date=2023-08-06 |website=Quanta Magazine |language=en}}</ref> [[Starship (genetics)|Starships]]<ref>{{cite journal | vauthors = Gluck-Thaler E, Ralston T, Konkel Z, Ocampos CG, Ganeshan VD, Dorrance AE, Niblack TL, Wood CW, Slot JC, Lopez-Nicora HD, Vogan AA | display-authors = 6 | title = Giant Starship Elements Mobilize Accessory Genes in Fungal Genomes | journal = Molecular Biology and Evolution | volume = 39 | issue = 5 | date = May 2022 | pmid = 35588244 | pmc = 9156397 | doi = 10.1093/molbev/msac109 | veditors = Larracuente A }}</ref><ref name="Bolakhe_2023" /> and Space invaders (or SPINs)<ref>{{cite journal | vauthors = Pace JK, Gilbert C, Clark MS, Feschotte C | title = Repeated horizontal transfer of a DNA transposon in mammals and other tetrapods | journal = Proceedings of the National Academy of Sciences of the United States of America | volume = 105 | issue = 44 | pages = 17023–17028 | date = November 2008 | pmid = 18936483 | pmc = 2579371 | doi = 10.1073/pnas.0806548105 | doi-access = free }}</ref><ref>{{Cite web |date=2008-11-03 |title=Space Invader DNA jumped across mammalian genomes |url=https://www.nationalgeographic.com/science/article/space-invader-dna-jumped-across-mammalian-genomes |archive-url=https://web.archive.org/web/20210504082210/https://www.nationalgeographic.com/science/article/space-invader-dna-jumped-across-mammalian-genomes |url-status=dead |archive-date=May 4, 2021 |access-date=2023-08-06 |website=Science |language=en}}</ref>

==Diseases==

The consequence of mobile genetic elements can alter the transcriptional patterns, which frequently leads to genetic disorders such as immune disorders, breast cancer, multiple sclerosis, and amyotrophic lateral sclerosis. In humans, stress can lead to transactional activation of MGEs such as [[endogenous retrovirus]]es, and this activation has been linked to [[neurodegeneration]].<ref>{{cite journal | vauthors = Antony JM, van Marle G, Opii W, Butterfield DA, Mallet F, Yong VW, Wallace JL, Deacon RM, Warren K, Power C | display-authors = 6 | title = Human endogenous retrovirus glycoprotein-mediated induction of redox reactants causes oligodendrocyte death and demyelination | journal = Nature Neuroscience | volume = 7 | issue = 10 | pages = 1088–1095 | date = October 2004 | pmid = 15452578 | doi = 10.1038/nn1319 | s2cid = 9882712 }}</ref>

==Other notes==
The total of all mobile genetic elements in a genome may be referred to as the [[mobilome]].

[[Barbara McClintock]] was awarded the [[list of Nobel laureates in Physiology or Medicine#1983|1983]] [[Nobel Prize in Physiology or Medicine]] "for her discovery of mobile genetic elements" ([[transposable element]]s).<ref name="nobel med 1983">{{cite web |title=The Nobel Prize in Physiology or Medicine 1983 |url=http://nobelprize.org/nobel_prizes/medicine/laureates/1983/ |publisher=nobelprize.org |access-date=14 July 2010}}</ref>

Mobile genetic elements play a critical role in the spread of virulence factors, such as [[exotoxin]]s and [[exoenzyme]]s, among bacteria. Strategies to combat certain bacterial infections by targeting these specific virulence factors and mobile genetic elements have been proposed.<ref>{{cite journal | vauthors = Keen EC | title = Paradigms of pathogenesis: targeting the mobile genetic elements of disease | journal = Frontiers in Cellular and Infection Microbiology | volume = 2 | pages = 161 | date = December 2012 | pmid = 23248780 | pmc = 3522046 | doi = 10.3389/fcimb.2012.00161 | doi-access = free }}</ref>

== See also ==
* [[ACLAME]] (The CLAssification of Mobile genetic Elements) database
* [[De novo gene birth]]
* [[Exon shuffling]]
* [[Fusion gene|Gene fusion]]
* [[Gene duplication]]
* [[Horizontal gene transfer]]
* [[Virulence factor]]s
* [[Miniature Inverted-repeat Transposable Elements]] (MITEs)
* [[Starship (genetics)|Starship]]

== References ==
{{Reflist}}

==Bibliography==
* {{cite book | veditors = Miller WJ, Capy P |year=2004 |title=Mobile genetic elements: protocols and genomic applications |publisher=Humana Press |isbn=978-1-58829-007-6 |url-access=registration |url=https://archive.org/details/mobilegeneticele00wolf}}
* {{cite book | veditors = Shapiro JA |editor-link=James A. Shapiro |year=1983 |title=Mobile genetic elements |publisher=Academic Press |isbn=978-0-12-638680-6 |url-access=registration |url=https://archive.org/details/mobilegeneticele00elem}}

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[[Category:Mobile genetic elements| ]]