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<span id="openzim-page-title" class="mw-page-title-main"><span class="mw-page-title-main">Transposable element</span></span>

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</style><div role="note" class="hatnote navigation-not-searchable">"Mobile DNA" redirects here. For the academic journal, see <a href="Mobile_DNA_(journal)" title="Mobile DNA (journal)">Mobile DNA (journal)</a>.</div>

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<p>A <b>transposable element</b> (<b>TE</b>), also <b>transposon</b>, or <b>jumping gene</b>, is a type of <a href="Mobile_genetic_element" class="mw-redirect" title="Mobile genetic element">mobile genetic element</a>, a nucleic acid sequence in DNA that can change its position within a genome.

</p><p>The discovery of mobile genetic elements earned <a href="Barbara_McClintock" title="Barbara McClintock">Barbara McClintock</a> a Nobel Prize in 1983.<sup id="cite_ref-1" class="reference"><a href="#cite_note-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup>

</p><p>TEs are very common in nature, especially in <a href="Plant" title="Plant">plants</a> and <a href="Animal" title="Animal">animals</a>. About 50% of the <a href="Maize" title="Maize">maize</a> genome, for instance, is made up by TEs.<sup id="cite_ref-:0_2-0" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>

</p><p>There are at least two classes of TEs: Class I TEs or <a href="Retrotransposon" title="Retrotransposon">retrotransposons</a> generally function via <a href="Reverse_transcription" class="mw-redirect" title="Reverse transcription">reverse transcription</a>, while Class II TEs or <a href="DNA_transposon" title="DNA transposon">DNA transposons</a> encode the protein <a href="Transposase" title="Transposase">transposase</a>, which they require for insertion and excision, and some of these TEs also encode other proteins.<sup id="cite_ref-prayla_3-0" class="reference"><a href="#cite_note-prayla-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading2"><h2 id="Discovery_by_Barbara_McClintock">Discovery by Barbara McClintock</h2></div>

<p><a href="Barbara_McClintock" title="Barbara McClintock">Barbara McClintock</a> discovered the first TEs in <a href="Maize" title="Maize">maize</a> (<i>Zea mays</i>) at the <a href="Cold_Spring_Harbor_Laboratory" title="Cold Spring Harbor Laboratory">Cold Spring Harbor Laboratory</a> in New York. McClintock was experimenting with maize plants that had broken chromosomes.<sup id="cite_ref-McGrayne165_4-0" class="reference"><a href="#cite_note-McGrayne165-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup>

</p><p>In the winter of 1944–1945, McClintock planted corn kernels that were self-pollinated, meaning that the silk (<a href="Style_(botany)" title="Style (botany)">style</a>) of the flower received pollen from its own <a href="Anther" class="mw-redirect" title="Anther">anther</a>.<sup id="cite_ref-McGrayne165_4-1" class="reference"><a href="#cite_note-McGrayne165-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> These kernels came from a long line of plants that had been self-pollinated, causing broken arms on the end of their ninth chromosomes.<sup id="cite_ref-McGrayne165_4-2" class="reference"><a href="#cite_note-McGrayne165-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> As the maize plants began to grow, McClintock noted unusual color patterns on the leaves.<sup id="cite_ref-McGrayne165_4-3" class="reference"><a href="#cite_note-McGrayne165-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> For example, one leaf had two albino patches of almost identical size, located side by side on the leaf.<sup id="cite_ref-McGrayne165_4-4" class="reference"><a href="#cite_note-McGrayne165-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> McClintock hypothesized that during cell division certain cells lost genetic material, while others gained what they had lost.<sup id="cite_ref-McGrayne166_5-0" class="reference"><a href="#cite_note-McGrayne166-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> However, when comparing the chromosomes of the current generation of plants with the parent generation, she found certain parts of the chromosome had switched position.<sup id="cite_ref-McGrayne166_5-1" class="reference"><a href="#cite_note-McGrayne166-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup> This refuted the popular genetic theory of the time that genes were fixed in their position on a chromosome. McClintock found that genes could not only move but they could also be turned on or off due to certain environmental conditions or during different stages of cell development.<sup id="cite_ref-McGrayne166_5-2" class="reference"><a href="#cite_note-McGrayne166-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>

</p><p>McClintock also showed that gene mutations could be reversed.<sup id="cite_ref-McGrayne167_6-0" class="reference"><a href="#cite_note-McGrayne167-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> She presented her report on her findings in 1951, and published an article on her discoveries in <i>Genetics</i> in November 1953 entitled "Induction of Instability at Selected Loci in Maize".<sup id="cite_ref-7" class="reference"><a href="#cite_note-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>

</p><p>At the 1951 Cold Spring Harbor Symposium where she first publicized her findings, her talk was met with silence.<sup id="cite_ref-8" class="reference"><a href="#cite_note-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> Her work was largely dismissed and ignored until the late 1960s–1970s when, after TEs were found in bacteria, it was rediscovered.<sup id="cite_ref-9" class="reference"><a href="#cite_note-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> She was awarded a <a href="List_of_Nobel_laureates_in_Physiology_or_Medicine" title="List of Nobel laureates in Physiology or Medicine">Nobel Prize in Physiology or Medicine</a> in 1983 for her discovery of TEs, more than thirty years after her initial research.<sup id="cite_ref-10" class="reference"><a href="#cite_note-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading2"><h2 id="Classification">Classification</h2></div>

<p>Transposable elements represent one of several types of <a href="Mobile_genetic_elements" title="Mobile genetic elements">mobile genetic elements</a>. TEs are assigned to one of two classes according to their mechanism of transposition, which can be described as either <i>copy and paste</i> (Class I TEs) or <i>cut and paste</i> (Class II TEs).<sup id="cite_ref-11" class="reference"><a href="#cite_note-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading3"><h3 id="Class_I:_Retrotransposon">Class I: Retrotransposon</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="Retrotransposon" title="Retrotransposon">Retrotransposon</a></div>

<p>Class I TEs are copied in two stages: first, they are <a href="Transcription_(genetics)" class="mw-redirect" title="Transcription (genetics)">transcribed</a> from DNA to <a href="RNA" title="RNA">RNA</a>, and the RNA produced is then <a href="Reverse_transcription" class="mw-redirect" title="Reverse transcription">reverse transcribed</a> to DNA. This <a href="CDNA" class="mw-redirect" title="CDNA">copied DNA</a> is then inserted back into the genome at a new position. The reverse transcription step is catalyzed by a <a href="Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a>, which is often encoded by the TE itself. The characteristics of retrotransposons are similar to <a href="Retrovirus" title="Retrovirus">retroviruses</a>, such as <a href="HIV" title="HIV">HIV</a>.

</p><p>Despite the potential negative effects of retrotransposons, like inserting itself into the middle of a necessary DNA sequence, which can render important genes unusable, they are still essential to keep a species' <a href="Ribosomal_DNA" title="Ribosomal DNA">ribosomal DNA</a> intact over the generations, preventing infertility.<sup id="cite_ref-12" class="reference"><a href="#cite_note-12"><span class="cite-bracket">[</span>12<span class="cite-bracket">]</span></a></sup>

</p><p>Retrotransposons are commonly grouped into three main orders:

</p>

<ul><li>Retrotransposons, with <a href="Long_terminal_repeat" title="Long terminal repeat">long terminal repeats</a> (LTRs), which encode reverse transcriptase, similar to retroviruses</li>

<li>Retroposons, <a href="LINEs" class="mw-redirect" title="LINEs">long interspersed nuclear elements</a> (LINEs, LINE-1s, or L1s), which encode reverse transcriptase but lack LTRs, and are transcribed by <a href="RNA_polymerase_II" title="RNA polymerase II">RNA polymerase II</a></li>

<li><a href="Short_interspersed_nuclear_element" title="Short interspersed nuclear element">Short interspersed nuclear elements</a> (SINEs) do not encode reverse transcriptase and are transcribed by <a href="RNA_polymerase_III" title="RNA polymerase III">RNA polymerase III</a></li></ul>

<p>Retroviruses can also be considered TEs. For example, after the conversion of retroviral RNA into DNA inside a <a href="Host_(biology)" title="Host (biology)">host</a> cell, the newly produced retroviral DNA is integrated into the <a href="Genome" title="Genome">genome</a> of the host cell. These integrated DNAs are termed <i><a href="Provirus" title="Provirus">proviruses</a></i>. The provirus is a specialized form of <a href="Eukaryotic" class="mw-redirect" title="Eukaryotic">eukaryotic</a> retrotransposon, which can produce RNA intermediates that may leave the host cell and infect other cells. The transposition cycle of retroviruses has similarities to that of <a href="Prokaryotic" class="mw-redirect" title="Prokaryotic">prokaryotic</a> TEs, suggesting a distant relationship between the two.

</p>

<div class="mw-heading mw-heading3"><h3 id="Class_II:_DNA_transposons">Class II: DNA transposons</h3></div>

<div role="note" class="hatnote navigation-not-searchable">Main article: <a href="DNA_transposon" title="DNA transposon">DNA transposon</a></div>

<p>The cut-and-paste transposition mechanism of class II TEs does not involve an RNA intermediate. The transpositions are catalyzed by several <a href="Transposase" title="Transposase">transposase</a> enzymes. Some transposases non-specifically bind to any target site in DNA, whereas others bind to specific target sequences. The transposase makes a staggered cut at the target site producing <a href="Sticky_ends" class="mw-redirect" title="Sticky ends">sticky ends</a>, cuts out the DNA transposon and ligates it into the target site. A <a href="DNA_polymerase" title="DNA polymerase">DNA polymerase</a> fills in the resulting gaps from the sticky ends and <a href="DNA_ligase" title="DNA ligase">DNA ligase</a> closes the sugar-phosphate backbone. This results in target site duplication and the insertion sites of DNA transposons may be identified by short direct repeats (a staggered cut in the target DNA filled by DNA polymerase) followed by <a href="Inverted_repeat" title="Inverted repeat">inverted repeats</a> (which are important for the TE <a href="DNA_repair" title="DNA repair">excision</a> by <a href="Transposase" title="Transposase">transposase</a>).

</p><p>Cut-and-paste TEs may be duplicated if their transposition takes place during <a href="S_phase" title="S phase">S phase</a> of the <a href="Cell_cycle" title="Cell cycle">cell cycle</a>, when a donor site has already been replicated but a target site has not yet been replicated. Such duplications at the target site can result in <a href="Gene_duplication" title="Gene duplication">gene duplication</a>, which plays an important role in genomic <a href="Evolution" title="Evolution">evolution</a>.<sup id="cite_ref-Brock_14-0" class="reference"><a href="#cite_note-Brock-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup><sup class="reference nowrap"><span title="Page / location: 284">: 284 </span></sup>

</p><p>Not all DNA transposons transpose through the cut-and-paste mechanism. In some cases, a <a href="Replicative_transposition" title="Replicative transposition">replicative transposition</a> is observed in which a transposon replicates itself to a new target site (e.g. <a href="Helitron_(biology)" title="Helitron (biology)">helitron</a>).

</p><p>Class II TEs comprise less than 2% of the human genome, making the rest Class I.<sup id="cite_ref-The_impact_of_L1_retrotransposons_o_15-0" class="reference"><a href="#cite_note-The_impact_of_L1_retrotransposons_o-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading3"><h3 id="Autonomous_and_non-autonomous">Autonomous and non-autonomous</h3></div>

<p>Transposition can be classified as either "autonomous" or "non-autonomous" in both Class I and Class II TEs. Autonomous TEs can move by themselves, whereas non-autonomous TEs require the presence of another TE to move. This is often because dependent TEs lack transposase (for Class II) or reverse transcriptase (for Class I).

</p><p>Activator element (<i>Ac</i>) is an example of an autonomous TE, and dissociation elements (<i>Ds</i>) is an example of a non-autonomous TE. Without <i>Ac,</i> <i>Ds</i> is not able to transpose.

</p>

<div class="mw-heading mw-heading3"><h3 id="Class_III">Class III</h3></div>

<p>Some researchers also identify a third class of transposable elements,<sup id="cite_ref-16" class="reference"><a href="#cite_note-16"><span class="cite-bracket">[</span>16<span class="cite-bracket">]</span></a></sup> which has been described as "a grab-bag consisting of transposons that don't clearly fit into the other two categories".<sup id="cite_ref-17" class="reference"><a href="#cite_note-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup> Examples of such TEs are the Foldback (FB) elements of <i>Drosophila melanogaster</i>, the TU elements of <i><a href="Strongylocentrotus_purpuratus" title="Strongylocentrotus purpuratus">Strongylocentrotus purpuratus</a></i>, and <a href="Miniature_Inverted-repeat_Transposable_Elements" title="Miniature Inverted-repeat Transposable Elements">Miniature Inverted-repeat Transposable Elements</a>.<sup id="cite_ref-18" class="reference"><a href="#cite_note-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-19" class="reference"><a href="#cite_note-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading2"><h2 id="Locations_in_genomes">Locations in genomes</h2></div>

<p>Transposable elements can be all over a genome, as in the case of maize, in which TEs make up 50% of the genome.<sup id="cite_ref-:0_2-1" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> In yeast (which has 5 classes of retrotransposons, Ty1-Ty5), over 90% of the Ty1 through T4 elements are located within 750 bp upstream of genes transcribed by <a href="RNA_polymerase_III" title="RNA polymerase III">RNA polymerase III</a>, particularly <a href="Transfer_RNA" title="Transfer RNA">tRNA</a> genes. The Ty5 elements are all located at the <a href="Telomere" title="Telomere">telomeres</a> or regions with telomeric <a href="Chromatin" title="Chromatin">chromatin</a>.<sup id="cite_ref-:0_2-2" class="reference"><a href="#cite_note-:0-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading2"><h2 id="Negative_effects">Negative effects</h2></div>

<p>Transposons can damage the genome of their host cell in different ways:

</p>

<ul><li>A transposon can insert into a functional gene and disable that gene.</li>

<li>After a DNA transposon is excised, the resulting gap may not be repaired correctly.</li></ul>

<ul><li>Many TEs contain promoters that drive transcription of their own genes. These promoters can cause aberrant expression of linked genes.</li></ul>

<div class="mw-heading mw-heading2"><h2 id="Diseases">Diseases</h2></div>

<p>Diseases often caused by TEs include

</p>

<ul><li><a href="Hemophilia" class="mw-redirect" title="Hemophilia">Hemophilia</a> A and B

<ul><li><a href="LINE1" title="LINE1">LINE1</a> (L1) TEs that land on the human Factor VIII have been shown to cause haemophilia<sup id="cite_ref-:6_20-0" class="reference"><a href="#cite_note-:6-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup></li></ul></li>

<li><a href="Severe_combined_immunodeficiency" title="Severe combined immunodeficiency">Severe combined immunodeficiency</a>

<ul><li>Insertion of L1 into the APC gene causes colon cancer, confirming that TEs play an important role in disease development.<sup id="cite_ref-21" class="reference"><a href="#cite_note-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup></li></ul></li>

<li><a href="Porphyria" title="Porphyria">Porphyria</a>

<ul><li>Insertion of <a href="Alu_element" title="Alu element">Alu element</a> into the PBGD gene leads to interference with the coding region and leads to acute intermittent porphyria<sup id="cite_ref-22" class="reference"><a href="#cite_note-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> (AIP).</li></ul></li>

<li>Predisposition to <a href="Cancer" title="Cancer">cancer</a>

<ul><li>LINE1(L1) TE's and other retrotransposons have been linked to cancer because they cause genomic instability.<sup id="cite_ref-:6_20-1" class="reference"><a href="#cite_note-:6-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup></li></ul></li>

<li><a href="Duchenne_muscular_dystrophy" title="Duchenne muscular dystrophy">Duchenne muscular dystrophy</a>.<sup id="cite_ref-23" class="reference"><a href="#cite_note-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-24" class="reference"><a href="#cite_note-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup>

<ul><li>Caused by SVA transposable element insertion in the <a href="Fukutin" title="Fukutin">fukutin</a> (FKTN) gene which renders the gene inactive.<sup id="cite_ref-:6_20-2" class="reference"><a href="#cite_note-:6-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup></li></ul></li>

<li>Alzheimer's Disease and other Tauopathies

<ul><li>Transposable element dysregulation can cause neuronal death, leading to neurodegenerative disorders<sup id="cite_ref-25" class="reference"><a href="#cite_note-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup></li></ul></li></ul>

<div class="mw-heading mw-heading2"><h2 id="Rate_of_transposition,_induction_and_defense">Rate of transposition, induction and defense</h2></div>

<p>One study estimated the rate of transposition of a particular retrotransposon, the Ty1 element in <i><a href="Saccharomyces_cerevisiae" title="Saccharomyces cerevisiae">Saccharomyces cerevisiae</a></i>. Using several assumptions, the rate of successful transposition event per single Ty1 element came out to be about once every few months to once every few years.<sup id="cite_ref-26" class="reference"><a href="#cite_note-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> Some TEs contain <a href="Heat_shock_protein" title="Heat shock protein">heat-shock like</a> promoters and their rate of transposition increases if the cell is subjected to stress,<sup id="cite_ref-27" class="reference"><a href="#cite_note-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> thus increasing the mutation rate under these conditions, which might be beneficial to the cell.

</p><p>Cells defend against the proliferation of TEs in a number of ways. These include <a href="Piwi-interacting_RNA" title="Piwi-interacting RNA">piRNAs</a> and <a href="SiRNA" class="mw-redirect" title="SiRNA">siRNAs</a>,<sup id="cite_ref-28" class="reference"><a href="#cite_note-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> which <a href="Gene_silencing" title="Gene silencing">silence</a> TEs after they have been transcribed.

</p><p>If organisms are mostly composed of TEs, one might assume that disease caused by misplaced TEs is very common, but in most cases TEs are silenced through <a href="Epigenetics" title="Epigenetics">epigenetic</a> mechanisms like <a href="DNA_methylation" title="DNA methylation">DNA methylation</a>, chromatin remodeling and piRNA, such that little to no phenotypic effects nor movements of TEs occur as in some wild-type plant TEs. Certain mutated plants have been found to have defects in methylation-related enzymes (methyl transferase) which cause the transcription of TEs, thus affecting the phenotype.<sup id="cite_ref-prayla_3-1" class="reference"><a href="#cite_note-prayla-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Mobilization_of_transposons_by_a_mu_29-0" class="reference"><a href="#cite_note-Mobilization_of_transposons_by_a_mu-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup>

</p><p>One hypothesis suggests that only approximately 100 LINE1 related sequences are active, despite their sequences making up 17% of the human genome. In human cells, silencing of LINE1 sequences is triggered by an <a href="RNA_interference" title="RNA interference">RNA interference</a> (RNAi) mechanism. Surprisingly, the RNAi sequences are derived from the 5′ untranslated region (UTR) of the LINE1, a long terminal which repeats itself. Supposedly, the 5′ LINE1 UTR that codes for the sense promoter for LINE1 transcription also encodes the antisense promoter for the <a href="MiRNA" class="mw-redirect" title="MiRNA">miRNA</a> that becomes the substrate for siRNA production. Inhibition of the RNAi silencing mechanism in this region showed an increase in LINE1 transcription.<sup id="cite_ref-prayla_3-2" class="reference"><a href="#cite_note-prayla-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-30" class="reference"><a href="#cite_note-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading2"><h2 id="Evolution">Evolution</h2></div>

<p>TEs are found in almost all life forms, and the scientific community is still exploring their evolution and their effect on genome evolution. It is unclear whether TEs originated in the <a href="Last_universal_common_ancestor" title="Last universal common ancestor">last universal common ancestor</a>, arose independently multiple times, or arose once and then spread to other kingdoms by <a href="Horizontal_gene_transfer" title="Horizontal gene transfer">horizontal gene transfer</a>.<sup id="cite_ref-31" class="reference"><a href="#cite_note-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup>

</p><p>Because excessive TE activity can damage <a href="Exon" title="Exon">exons</a>, many organisms have acquired mechanisms to inhibit their activity. Bacteria may undergo high rates of <a href="Gene_deletion" class="mw-redirect" title="Gene deletion">gene deletion</a> as part of a mechanism to remove TEs and viruses from their genomes, while <a href="Eukaryote" title="Eukaryote">eukaryotic</a> organisms typically use <a href="RNA_interference" title="RNA interference">RNA interference</a> to inhibit TE activity. Nevertheless, some TEs generate large families often associated with <a href="Speciation" title="Speciation">speciation</a> events.<sup id="cite_ref-32" class="reference"><a href="#cite_note-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup> Evolution often deactivates DNA transposons, leaving them as <a href="Intron" title="Intron">introns</a> (inactive gene sequences). In vertebrate animal cells, nearly all 100,000+ DNA transposons per genome have genes that encode inactive transposase polypeptides.<sup id="cite_ref-33" class="reference"><a href="#cite_note-33"><span class="cite-bracket">[</span>33<span class="cite-bracket">]</span></a></sup>

</p><p><b>Sleeping Beauty/Tc1/mariner</b>. The first synthetic transposon designed for use in vertebrate (including human) cells, the <a href="Sleeping_Beauty_transposon_system" title="Sleeping Beauty transposon system">Sleeping Beauty transposon system</a>, is a Tc1/mariner-like transposon. Its dead ("fossil") versions are spread widely in the salmonid genome and a functional version was engineered by comparing those versions.<sup id="cite_ref-34" class="reference"><a href="#cite_note-34"><span class="cite-bracket">[</span>34<span class="cite-bracket">]</span></a></sup> Human Tc1-like transposons are divided into Hsmar1 and Hsmar2 subfamilies. Although both types are inactive, one copy of Hsmar1 found in the <a href="SETMAR" title="SETMAR">SETMAR</a> gene is under selection as it provides DNA-binding for the histone-modifying protein.<sup id="cite_ref-35" class="reference"><a href="#cite_note-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup> Many other human genes are similarly derived from transposons.<sup id="cite_ref-36" class="reference"><a href="#cite_note-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> Hsmar2 has been reconstructed multiple times from the fossil sequences.<sup id="cite_ref-37" class="reference"><a href="#cite_note-37"><span class="cite-bracket">[</span>37<span class="cite-bracket">]</span></a></sup>

</p><p><b>Selective advantages</b>. TEs may affect gene regulatory networks and thus have evolutionary advantages.<sup id="cite_ref-38" class="reference"><a href="#cite_note-38"><span class="cite-bracket">[</span>38<span class="cite-bracket">]</span></a></sup> <a href="Interspersed_repeat" title="Interspersed repeat">Interspersed repeats</a> are created by transposition; since they can inhibit <a href="Gene_conversion" title="Gene conversion">gene conversion</a>, they protect novel gene sequences from being overwritten by similar gene sequences and thereby facilitate the development of new genes. TEs may also have been co-opted by the <a href="Adaptive_immune_system" title="Adaptive immune system">vertebrate immune system</a> as a means of producing antibody diversity. The <a href="V(D)J_recombination" title="V(D)J recombination">V(D)J recombination</a> system operates by a mechanism similar to that of some TEs. TEs also serve to generate repeating sequences that can form <a href="DsRNA" class="mw-redirect" title="DsRNA">dsRNA</a> to act as a substrate for the action of <a href="ADAR" title="ADAR">ADAR</a> in RNA editing.<sup id="cite_ref-39" class="reference"><a href="#cite_note-39"><span class="cite-bracket">[</span>39<span class="cite-bracket">]</span></a></sup>

</p><p>TEs can contain many types of genes, including those conferring antibiotic resistance and the ability to transpose to conjugative plasmids. Some TEs also contain <a href="Integron" title="Integron">integrons</a>, genetic elements that can capture and express genes from other sources. These contain <a href="Integrase" title="Integrase">integrase</a>, which can integrate <a href="Gene_cassette" title="Gene cassette">gene cassettes</a>. There are over 40 antibiotic resistance genes identified on cassettes, as well as virulence genes.

</p><p><b>Novel genes and exon shuffling</b>. Transposons do not always excise their elements precisely, sometimes removing the adjacent base pairs; this can lead to merged exons in a process called <a href="Exon_shuffling" title="Exon shuffling">exon shuffling</a>. Shuffling two unrelated exons can create a novel gene product or, more likely, an intron.<sup id="cite_ref-40" class="reference"><a href="#cite_note-40"><span class="cite-bracket">[</span>40<span class="cite-bracket">]</span></a></sup>

</p><p>Some non-autonomous DNA TEs found in plants can capture coding DNA from genes and shuffle them across the genome.<sup id="cite_ref-41" class="reference"><a href="#cite_note-41"><span class="cite-bracket">[</span>41<span class="cite-bracket">]</span></a></sup> This process can duplicate genes in the genome (a phenomenon called transduplication), and can contribute to generate novel genes by exon shuffling.<sup id="cite_ref-42" class="reference"><a href="#cite_note-42"><span class="cite-bracket">[</span>42<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading3"><h3 id="Evolutionary_drive_for_TEs_on_the_genomic_context">Evolutionary drive for TEs on the genomic context</h3></div>

<p>There is a hypothesis that states that TEs might provide a ready source of DNA that could be co-opted by the cell to help regulate gene expression. Research showed that many diverse modes of TEs co-evolution along with some transcription factors targeting TE-associated genomic elements and chromatin are evolving from TE sequences. Most of the time, these particular modes do not follow the simple model of TEs and regulating host gene expression.<sup id="cite_ref-Zhou_19359–19366_43-0" class="reference"><a href="#cite_note-Zhou_19359–19366-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup>

</p>

<div class="mw-heading mw-heading2"><h2 id="De_novo_repeat_identification"><i>De novo</i> repeat identification</h2></div>

<p><i>De novo</i> repeat identification is an initial scan of sequence data that seeks to find the repetitive regions of the genome, and to classify these repeats. Many computer programs exist to perform <i>de novo</i> repeat identification, all operating under the same general principles.<sup id="cite_ref-Wojciech_Makałowski_p._337-359_44-0" class="reference"><a href="#cite_note-Wojciech_Makałowski_p._337-359-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> As short tandem repeats are generally 1–6 base pairs in length and are often consecutive, their identification is relatively simple.<sup id="cite_ref-:1_45-0" class="reference"><a href="#cite_note-:1-45"><span class="cite-bracket">[</span>45<span class="cite-bracket">]</span></a></sup> Dispersed repetitive elements, on the other hand, are more challenging to identify, due to the fact that they are longer and have often acquired mutations. However, it is important to identify these repeats as they are often found to be transposable elements (TEs).<sup id="cite_ref-Wojciech_Makałowski_p._337-359_44-1" class="reference"><a href="#cite_note-Wojciech_Makałowski_p._337-359-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup>

</p><p><i>De novo</i> identification of transposons involves three steps: 1) find all repeats within the genome, 2) build a <a href="Consensus_sequence" title="Consensus sequence">consensus</a> of each family of sequences, and 3) classify these repeats. There are three groups of algorithms for the first step. One group is referred to as the <a href="K-mer" title="K-mer">k-mer</a> approach, where a k-mer is a sequence of length k. In this approach, the genome is scanned for overrepresented k-mers; that is, k-mers that occur more often than is likely based on probability alone. The length k is determined by the type of transposon being searched for. The k-mer approach also allows mismatches, the number of which is determined by the analyst. Some k-mer approach programs use the k-mer as a base, and extend both ends of each repeated k-mer until there is no more similarity between them, indicating the ends of the repeats.<sup id="cite_ref-Wojciech_Makałowski_p._337-359_44-2" class="reference"><a href="#cite_note-Wojciech_Makałowski_p._337-359-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Another group of algorithms employs a method called sequence self-comparison. Sequence self-comparison programs use databases such as <a href="BLAST_(biotechnology)" title="BLAST (biotechnology)">AB-BLAST</a> to conduct an initial <a href="Sequence_alignment" title="Sequence alignment">sequence alignment</a>. As these programs find groups of elements that partially overlap, they are useful for finding highly diverged transposons, or transposons with only a small region copied into other parts of the genome.<sup id="cite_ref-:2_46-0" class="reference"><a href="#cite_note-:2-46"><span class="cite-bracket">[</span>46<span class="cite-bracket">]</span></a></sup> Another group of algorithms follows the periodicity approach. These algorithms perform a <a href="Fourier_transformation" class="mw-redirect" title="Fourier transformation">Fourier transformation</a> on the sequence data, identifying periodicities, regions that are repeated periodically, and are able to use peaks in the resultant spectrum to find candidate repetitive elements. This method works best for tandem repeats, but can be used for dispersed repeats as well. However, it is a slow process, making it an unlikely choice for genome-scale analysis.<sup id="cite_ref-Wojciech_Makałowski_p._337-359_44-3" class="reference"><a href="#cite_note-Wojciech_Makałowski_p._337-359-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup>

</p><p>The second step of <i>de novo</i> repeat identification involves building a consensus of each family of sequences. A <a href="Consensus_sequence" title="Consensus sequence">consensus sequence</a> is a sequence that is created based on the repeats that comprise a TE family. A base pair in a consensus is the one that occurred most often in the sequences being compared to make the consensus. For example, in a family of 50 repeats where 42 have a T base pair in the same position, the consensus sequence would have a T at this position as well, as the base pair is representative of the family as a whole at that particular position, and is most likely the base pair found in the family's ancestor at that position.<sup id="cite_ref-Wojciech_Makałowski_p._337-359_44-4" class="reference"><a href="#cite_note-Wojciech_Makałowski_p._337-359-44"><span class="cite-bracket">[</span>44<span class="cite-bracket">]</span></a></sup> Once a consensus sequence has been made for each family, it is then possible to move on to further analysis, such as TE classification and genome masking in order to quantify the overall TE content of the genome.

</p>

<div class="mw-heading mw-heading2"><h2 id="Adaptive_TEs">Adaptive TEs</h2></div>

<p>Transposable elements have been recognized as good candidates for stimulating gene adaptation, through their ability to regulate the expression levels of nearby genes.<sup id="cite_ref-:3_47-0" class="reference"><a href="#cite_note-:3-47"><span class="cite-bracket">[</span>47<span class="cite-bracket">]</span></a></sup> Combined with their "mobility", transposable elements can be relocated adjacent to their targeted genes, and control the expression levels of the gene, dependent upon the circumstances.

</p><p>The study conducted in 2008, "High Rate of Recent Transposable Element–Induced Adaptation in Drosophila melanogaster", used <i>D. melanogaster</i> that had recently migrated from Africa to other parts of the world, as a basis for studying adaptations caused by transposable elements. Although most of the TEs were located on introns, the experiment showed a significant difference in gene expressions between the population in Africa and other parts of the world. The four TEs that caused the selective sweep were more prevalent in <i>D. melanogaster</i> from temperate climates, leading the researchers to conclude that the selective pressures of the climate prompted genetic adaptation.<sup id="cite_ref-:4_48-0" class="reference"><a href="#cite_note-:4-48"><span class="cite-bracket">[</span>48<span class="cite-bracket">]</span></a></sup> From this experiment, it has been confirmed that adaptive TEs are prevalent in nature, by enabling organisms to adapt gene expression as a result of new selective pressures.

</p><p>However, not all effects of adaptive TEs are beneficial to the population. In the research conducted in 2009, "A Recent Adaptive Transposable Element Insertion Near Highly Conserved Developmental Loci in Drosophila melanogaster", a TE, inserted between Jheh 2 and Jheh 3, revealed a downgrade in the expression level of both of the genes. Downregulation of such genes has caused <i>Drosophila</i> to exhibit extended developmental time and reduced egg to adult viability. Although this adaptation was observed in high frequency in all non-African populations, it was not fixed in any of them.<sup id="cite_ref-:5_49-0" class="reference"><a href="#cite_note-:5-49"><span class="cite-bracket">[</span>49<span class="cite-bracket">]</span></a></sup> This is not hard to believe, since it is logical for a population to favor higher egg to adult viability, therefore trying to purge the trait caused by this specific TE adaptation.

</p><p>At the same time, there have been several reports showing the advantageous adaptation caused by TEs. In the research done with silkworms, "An Adaptive Transposable Element insertion in the Regulatory Region of the EO Gene in the Domesticated Silkworm", a TE insertion was observed in the cis-regulatory region of the EO gene, which regulates molting hormone 20E, and enhanced expression was recorded. While populations without the TE insert are often unable to effectively regulate hormone 20E under starvation conditions, those with the insert had a more stable development, which resulted in higher developmental uniformity.<sup id="cite_ref-50" class="reference"><a href="#cite_note-50"><span class="cite-bracket">[</span>50<span class="cite-bracket">]</span></a></sup>

</p><p>These three experiments all demonstrated different ways in which TE insertions can be advantageous or disadvantageous, through means of regulating the expression level of adjacent genes. The field of adaptive TE research is still under development and more findings can be expected in the future.

</p>

<div class="mw-heading mw-heading2"><h2 id="TEs_participates_in_gene_control_networks">TEs participates in gene control networks</h2></div>

<p>Recent studies have confirmed that TEs can contribute to the generation of transcription factors. However, how this process of contribution can have an impact on the participation of genome control networks. TEs are more common in many regions of the DNA and it makes up 45% of total human DNA. Also, TEs contributed to 16% of transcription factor binding sites. A larger number of motifs are also found in non-TE-derived DNA, and the number is larger than TE-derived DNA. All these factors correlate to the direct participation of TEs in many ways of gene control networks.<sup id="cite_ref-Zhou_19359–19366_43-1" class="reference"><a href="#cite_note-Zhou_19359–19366-43"><span class="cite-bracket">[</span>43<span class="cite-bracket">]</span></a></sup>

</p>

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<ul><li><a href="Decrease_in_DNA_Methylation_I_(DDM1)" class="mw-redirect" title="Decrease in DNA Methylation I (DDM1)">Decrease in DNA Methylation I (DDM1)</a></li>

<li><a href="Epigenetic_regulation_of_transposable_elements_in_the_plant_kingdom" title="Epigenetic regulation of transposable elements in the plant kingdom">Epigenetic regulation of transposable elements in the plant kingdom</a></li>

<li><a href="Evolution_of_sexual_reproduction" title="Evolution of sexual reproduction">Evolution of sexual reproduction</a></li>

<li><a href="Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li>

<li><a href="Intragenomic_conflict" title="Intragenomic conflict">Intragenomic conflict</a></li>

<li><a href="PiggyBac_transposon_system" title="PiggyBac transposon system">PiggyBac transposon system</a></li>

<li><a href="Polinton" title="Polinton">Polinton</a></li>

<li><a href="Tn3_transposon" title="Tn3 transposon">Tn3 transposon</a></li>

<li><a href="Transpogene" title="Transpogene">Transpogene</a></li>

<li><a href="Transposon_tagging" title="Transposon tagging">Transposon tagging</a></li></ul></div>

<div class="mw-heading mw-heading2"><h2 id="Notes">Notes</h2></div>

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</style><cite id="CITEREFKidwell2005" class="citation book cs1">Kidwell MG (2005). "Transposable elements". In T.R. Gregory (ed.). <i>The Evolution of the Genome</i>. San Diego: Elsevier. pp. <span class="nowrap">165–</span>221. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-123-01463-4</bdi>.</cite></li>

<li><cite id="CITEREFCraigCraigieGellertand_Lambowitz2002" class="citation book cs1">Craig NL, Craigie R, Gellert M, and Lambowitz AM, eds. (2002). <i>Mobile DNA II</i>. Washington, DC: ASM Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-1-555-81209-6</bdi>.</cite></li>

<li><cite id="CITEREFLewin2000" class="citation book cs1">Lewin B (2000). <span class="id-lock-registration" title="Free registration required"><a rel="nofollow" class="external text" href="https://archive.org/details/genesvii00lewi"><i>Genes VII</i></a></span>. Oxford University Press. <a href="ISBN_(identifier)" class="mw-redirect" title="ISBN (identifier)">ISBN</a> <bdi>978-0-198-79276-5</bdi>.</cite></li></ul>

<div class="mw-heading mw-heading3"><h3 id="References">References</h3></div>

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<li id="cite_note-prayla-3"><span class="mw-cite-backlink">^ <a href="#cite_ref-prayla_3-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-prayla_3-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-prayla_3-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><cite id="CITEREFPray2008" class="citation journal cs1">Pray LA (2008). <a rel="nofollow" class="external text" href="http://www.nature.com/scitable/topicpage/transposons-the-jumping-genes-518">"Transposons: The jumping genes"</a>. <i>Nature Education</i>. <b>1</b> (1): 204.</cite></span>

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<li id="cite_note-McGrayne166-5"><span class="mw-cite-backlink">^ <a href="#cite_ref-McGrayne166_5-0"><sup><i><b>a</b></i></sup></a> <a href="#cite_ref-McGrayne166_5-1"><sup><i><b>b</b></i></sup></a> <a href="#cite_ref-McGrayne166_5-2"><sup><i><b>c</b></i></sup></a></span> <span class="reference-text"><a href="#CITEREFMcGrayne1998">McGrayne 1998</a>, p. 166</span>

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<li id="cite_note-:5-49"><span class="mw-cite-backlink"><b><a href="#cite_ref-:5_49-0">^</a></b></span> <span class="reference-text"><cite id="CITEREFGonzálezMacphersonPetrov2009" class="citation journal cs1">González J, Macpherson JM, Petrov DA (September 2009). <a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2734154">"A recent adaptive transposable element insertion near highly conserved developmental loci in Drosophila melanogaster"</a>. <i>Molecular Biology and Evolution</i>. <b>26</b> (9): <span class="nowrap">1949–</span>61. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fmolbev%2Fmsp107">10.1093/molbev/msp107</a>. <a href="PMC_(identifier)" class="mw-redirect" title="PMC (identifier)">PMC</a> <span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/pmc/articles/PMC2734154">2734154</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/19458110">19458110</a>.</cite></span>

</li>

<li id="cite_note-50"><span class="mw-cite-backlink"><b><a href="#cite_ref-50">^</a></b></span> <span class="reference-text"><cite id="CITEREFSunShenHanCao2014" class="citation journal cs1">Sun W, Shen YH, Han MJ, Cao YF, Zhang Z (December 2014). <a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fmolbev%2Fmsu261">"An adaptive transposable element insertion in the regulatory region of the EO gene in the domesticated silkworm, Bombyx mori"</a>. <i>Molecular Biology and Evolution</i>. <b>31</b> (12): <span class="nowrap">3302–</span>13. <a href="Doi_(identifier)" class="mw-redirect" title="Doi (identifier)">doi</a>:<span class="id-lock-free" title="Freely accessible"><a rel="nofollow" class="external text" href="https://doi.org/10.1093%2Fmolbev%2Fmsu261">10.1093/molbev/msu261</a></span>. <a href="PMID_(identifier)" class="mw-redirect" title="PMID (identifier)">PMID</a> <a rel="nofollow" class="external text" href="https://pubmed.ncbi.nlm.nih.gov/25213334">25213334</a>.</cite></span>

</li>

</ol></div></div>

<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>

<ul><li><cite class="citation journal cs1"><a rel="nofollow" class="external text" href="https://www.newscientist.com/article/mg19025565.500">"An immune system so versatile it might kill you"</a>. <i>New Scientist</i> (2556). 21 June 2006.</cite> – A possible connection between aberrant reinsertions and lymphoma.</li>

<li><a rel="nofollow" class="external text" href="http://www.girinst.org/">Repbase</a> – a database of transposable element sequences</li>

<li><a rel="nofollow" class="external text" href="http://www.dfam.org/">Dfam</a> - a database of transposable element families, multiple sequence alignments, and sequence models</li>

<li><a rel="nofollow" class="external text" href="http://www.repeatmasker.org/">RepeatMasker</a> – a computer program used by computational biologists to <a href="Annotation#Computational_biology" title="Annotation">annotate</a> transposons in DNA sequences</li>

<li><a rel="nofollow" class="external text" href="http://cshprotocols.cshlp.org/cgi/content/full/2009/8/pdb.prot5270">Use of the Sleeping Beauty Transposon System for Stable Gene Expression in Mouse Embryonic Stem Cells</a></li>

<li><a rel="nofollow" class="external text" href="https://www.youtube.com/watch?time_continue=53&v=eSD_tbjfSlA&feature=emb_logo">Introduction to Transposons, 2018 YouTube video</a></li></ul>

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</style><div id="Genetics:_repeated_sequence,_transposon,_gene_duplication239" style="font-size:114%;margin:0 4em"><a href="Genetics" title="Genetics">Genetics</a>: <a href="Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">repeated sequence</a>, , <a href="Gene_duplication" title="Gene duplication">gene duplication</a></div></th></tr><tr><td class="navbox-abovebelow" colspan="2"><div><a href="Repeatome" title="Repeatome">Repeatome</a></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequence</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Tandem_repeat" title="Tandem repeat">Tandem repeats</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Satellite_DNA" title="Satellite DNA">Satellite DNA</a></li>

<li><a href="Variable_number_tandem_repeat" title="Variable number tandem repeat">Variable number tandem repeat</a>/<a href="Minisatellite" title="Minisatellite">Minisatellite</a></li>

<li><a href="Microsatellite" title="Microsatellite">Short tandem repeat/Microsatellite</a> (<a href="Trinucleotide_repeat_disorder" title="Trinucleotide repeat disorder">Trinucleotide repeat disorders</a>)</li>

<li><a href="Macrosatellite" title="Macrosatellite">Macrosatellite</a><br></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Interspersed_repeat" title="Interspersed repeat">Interspersed<br>repeat</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Gene_conversion" title="Gene conversion">Gene conversion</a></li>

<li><a href="Retrotransposon" title="Retrotransposon">Retrotransposon</a></li>

<li>DNA transposon

<ul><li><a href="Polinton" title="Polinton">Polinton</a></li>

<li><a href="Helitron_(biology)" title="Helitron (biology)">Helitron</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Inverted_repeat" title="Inverted repeat">Inverted repeat</a></li>

<li><a href="Direct_repeat" title="Direct repeat">Direct repeat</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Retrotransposon" title="Retrotransposon">Retrotransposon</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Short_interspersed_nuclear_element" title="Short interspersed nuclear element">SINEs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Alu_element" title="Alu element">Alu sequence</a></li>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Long_interspersed_nuclear_element" title="Long interspersed nuclear element">LINEs</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="LTR_retrotransposon" title="LTR retrotransposon">LTRs</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<li><a href="Retroposon" title="Retroposon">retroposon</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="DNA_transposon" title="DNA transposon">DNA transposon</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li>Academ</li>

<li>Crypton</li>

<li>EnSpm/CACTA</li>

<li>Ginger1</li>

<li>Ginger2</li>

<li>Harbinger</li>

<li><a href="Helitron_(biology)" title="Helitron (biology)">Helitron</a></li>

<li>Kolobok</li>

<li><a href="Tc1/mariner" title="Tc1/mariner">Tc1/mariner</a></li>

<li>Merlin</li>

<li>Novosib</li>

<li><a href="P_element" title="P element">P element</a></li>

<li><a href="PiggyBac_transposon_system" title="PiggyBac transposon system">PiggyBac</a></li>

<li><a href="Polinton" title="Polinton">Polinton</a></li>

<li><a href="Transib" title="Transib">Transib</a></li>

<li>Zator</li>

<li>Zisupton</li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Gene_duplication" title="Gene duplication">Gene duplication</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Gene_amplification" title="Gene amplification">Gene amplification</a></li>

<li><a href="Tandemly_arrayed_genes" title="Tandemly arrayed genes">Tandemly arrayed genes</a>

<ul><li><a href="Ribosomal_DNA" title="Ribosomal DNA">Ribosomal DNA</a></li></ul></li>

<li><a href="Gene_family" title="Gene family">Gene family</a>

<ul><li><a href="Gene_cluster" title="Gene cluster">Gene cluster</a></li></ul></li>

<li><a href="Pseudogene" title="Pseudogene">Pseudogene</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Genomic_island" title="Genomic island">Genomic island</a>

<ul><li><a href="Pathogenicity_island" title="Pathogenicity island">Pathogenicity island</a></li>

<li>Symbiosis island</li></ul></li>

<li><a href="Low_copy_repeats" title="Low copy repeats">Low copy repeats</a></li>

<li><a href="CRISPR" title="CRISPR">CRISPR</a></li>

<li><a href="Telomere" title="Telomere">Telomere</a></li>

<li><a href="Protein_tandem_repeats" title="Protein tandem repeats">Protein tandem repeats</a></li></ul>

</div></td></tr></tbody></table></div>

<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Genetics:_homologous_recombination_/_mobile_genetic_elements147" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Genetics:_homologous_recombination_/_mobile_genetic_elements147" style="font-size:114%;margin:0 4em"><a href="Genetics" title="Genetics">Genetics</a>: <a href="Homologous_recombination" title="Homologous recombination">homologous recombination</a> / <a href="Mobile_genetic_elements" title="Mobile genetic elements">mobile genetic elements</a></div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Primarily <a href="Prokaryote" title="Prokaryote">prokaryotic</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Bacterial_conjugation" title="Bacterial conjugation">Conjugation</a></li>

<li><a href="Transduction_(genetics)" title="Transduction (genetics)">Transduction</a></li>

<li><a href="Transformation_(genetics)" class="mw-redirect" title="Transformation (genetics)">Transformation</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Occurs in <a href="Eukaryote" title="Eukaryote">eukaryotes</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Transfection" title="Transfection">Transfection</a></li>

<li><a href="Chromosomal_crossover" title="Chromosomal crossover">Chromosomal crossover</a></li>

<li><a href="Gene_conversion" title="Gene conversion">Gene conversion</a></li>

<li><a href="Fusion_gene" title="Fusion gene">Fusion gene</a></li>

<li><a href="Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li>

<li><a href="Sister_chromatid_exchange" title="Sister chromatid exchange">Sister chromatid exchange</a></li>

</ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Virus" title="Virus">Viral</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Antigenic_shift" title="Antigenic shift">Antigenic shift</a></li>

<li><a href="Reassortment" title="Reassortment">Reassortment</a></li>

<li><a href="Viral_shift" class="mw-redirect" title="Viral shift">Viral shift</a></li></ul>

</div></td></tr></tbody></table></div>

<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Self-replicating_organic_structures216" style="padding:3px"><table class="nowraplinks mw-collapsible autocollapse navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2" style="text-align: center;"><div id="Self-replicating_organic_structures216" style="font-size:114%;margin:0 4em"><a href="Self-replication" title="Self-replication">Self-replicating</a> organic structures</div></th></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Life" title="Life">Cellular life</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Bacteria" title="Bacteria">Bacteria</a></li>

<li><a href="Archaea" title="Archaea">Archaea</a></li>

<li><a href="Eukaryote" title="Eukaryote">Eukaryota</a>

<ul><li><a href="Animal" title="Animal">Animalia</a></li>

<li><a href="Fungus" title="Fungus">Fungi</a></li>

<li><a href="Plant" title="Plant">Plantae</a></li>

<li><a href="Protist" title="Protist">Protista</a></li></ul></li>

<li><i><a href="Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i>

<ul><li><i><a href="Parakaryon" title="Parakaryon">Parakaryon</a></i></li>

<li><a href="Biological_dark_matter" title="Biological dark matter">Biological dark matter</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="Virus" title="Virus">Virus</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="DNA_virus#Group_I:_dsDNA_viruses" title="DNA virus">dsDNA virus</a>

<ul><li><a href="Giant_virus" title="Giant virus">Giant virus</a></li></ul></li>

<li><a href="DNA_virus#Group_II:_ssDNA_viruses" title="DNA virus">ssDNA virus</a></li>

<li><a href="Double-stranded_RNA_viruses" title="Double-stranded RNA viruses">dsRNA virus</a></li>

<li><a href="RNA_virus#Group_IV—positive-sense_ssRNA_viruses" title="RNA virus">(+)ssRNA virus</a></li>

<li><a href="RNA_virus#Group_V—negative-sense_ssRNA_viruses" title="RNA virus">(−)ssRNA virus</a></li>

<li><a href="Retrovirus" title="Retrovirus">ssRNA-RT virus</a></li>

<li><a href="DsDNA-RT_virus" class="mw-redirect" title="DsDNA-RT virus">dsDNA-RT virus</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%;background:#CEDAF2;"><a href="Subviral_agents" class="mw-redirect" title="Subviral agents">Subviral<br>agents</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="Viroid" title="Viroid">Viroid</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li>

<li><i><a href="Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;;background:#CEDAF2;"><a href="Helper_virus" title="Helper virus">Helper-virus<br>dependent</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;"><a href="Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li>ssRNA satellite virus</li>

<li>dsDNA satellite virus (<a href="Virophage" title="Virophage">Virophage</a>)</li>

<li>ssDNA satellite virus</li>

<li>ssDNA satellite</li>

<li>dsRNA satellite</li>

<li>ssRNA satellite (<a href="Virusoid" title="Virusoid">Virusoid</a>)</li>

<li>Satellite-like nucleic acids

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%;text-align: center;;background:#CEDAF2;">Other</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="Prion" title="Prion">Prion</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Prion" title="Prion">Mammalian prion</a></li>

<li><a href="Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Nucleic_acid" title="Nucleic acid">Nucleic acid</a><br>self-replication</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;"><a href="Mobile_genetic_elements" title="Mobile genetic elements">Mobile genetic<br>elements</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Mobilome" title="Mobilome">Mobilome</a>

<ul><li><a href="Horizontal_gene_transfer" title="Horizontal gene transfer">Horizontal gene transfer</a></li>

<li><a href="Genomic_island" title="Genomic island">Genomic island</a></li></ul></li>

<li>

<ul><li><a href="Retrotransposon" title="Retrotransposon">Class I or retrotransposon</a></li>

<li><a href="DNA_transposon" title="DNA transposon">Class II or DNA transposon</a></li></ul></li>

<li><a href="Plasmid" title="Plasmid">Plasmid</a>

<ul><li><a href="Fertility_factor_(bacteria)" class="mw-redirect" title="Fertility factor (bacteria)">Fertility</a></li>

<li><a href="R-factor" class="mw-redirect" title="R-factor">Resistance</a></li>

<li>Degradative</li>

<li><a href="Virulence_factor" title="Virulence factor">Virulence</a>/<a href="Ti_plasmid" title="Ti plasmid">Ti</a></li>

<li>Cryptic</li></ul></li>

<li><a href="Cosmid" title="Cosmid">Cosmid</a>

<ul><li><a href="Fosmid" title="Fosmid">Fosmid</a></li></ul></li>

<li><a href="Phagemid" title="Phagemid">Phagemid</a></li>

<li><a href="Group_I_catalytic_intron" title="Group I catalytic intron">Group I intron</a></li>

<li><a href="Group_II_intron" title="Group II intron">Group II intron</a></li>

<li><a href="Retrozyme" title="Retrozyme">Retrozyme</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:7em;text-align: center;">Other aspects</th><td class="navbox-list-with-group navbox-list navbox-even" style="padding:0"><div style="padding:0 0.25em">

<ul><li><a href="DNA_replication" title="DNA replication">DNA replication</a>

<ul><li><a href="RNA-dependent_RNA_polymerase" title="RNA-dependent RNA polymerase">RNA replication</a></li></ul></li>

<li><a href="Chromosome" title="Chromosome">Chromosome</a>

<ul><li><a href="Linear_chromosome" title="Linear chromosome">Linear</a></li>

<li><a href="Circular_chromosome" title="Circular chromosome">Circular</a></li>

<li><a href="Extrachromosomal_DNA" title="Extrachromosomal DNA">Extrachromosomal DNA</a></li>

<li><a href="Secondary_chromosome" title="Secondary chromosome">Secondary chromosome</a></li></ul></li>

<li><a href="Genome" title="Genome">Genome</a>

<li><a href="Gene_duplication" title="Gene duplication">Gene duplication</a></li>

<li><a href="Non-coding_DNA" title="Non-coding DNA">Non-coding DNA</a></li></ul></li>

<li><a href="Origin_of_replication" title="Origin of replication">Origin of replication</a>

<ul><li><a href="Replicon_(genetics)" title="Replicon (genetics)">Replicon</a></li></ul></li>

<li><a href="Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a>

<ul><li><a href="Provirus" title="Provirus">Provirus</a></li>

<li><a href="Prophage" title="Prophage">Prophage</a></li>

<li><a href="Endogenous_retrovirus" title="Endogenous retrovirus">Endogenous retrovirus</a></li>

<li><a href="Transpoviron" title="Transpoviron">Transpoviron</a></li></ul></li>

<li><a href="Repeated_sequence_(DNA)" title="Repeated sequence (DNA)">Repeated sequences in DNA</a>

<ul><li><a href="Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li>

<li><a href="Interspersed_repeat" title="Interspersed repeat">Interspersed repeat</a></li></ul></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Endosymbiont" title="Endosymbiont">Endosymbiosis</a></th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Mitochondrion" title="Mitochondrion">Mitochondrion</a>

<ul><li><a href="Mitosome" title="Mitosome">Mitosome</a></li>

<li><a href="Hydrogenosome" title="Hydrogenosome">Hydrogenosome</a></li></ul></li>

<li><a href="Plastid" title="Plastid">Plastid</a>

<ul><li><a href="Chloroplast" title="Chloroplast">Chloroplast</a></li>

<li><a href="Chromoplast" title="Chromoplast">Chromoplast</a></li>

<li><a href="Gerontoplast" title="Gerontoplast">Gerontoplast</a></li>

<li><a href="Leucoplast" title="Leucoplast">Leucoplast</a></li>

<li><a href="Apicoplast" title="Apicoplast">Apicoplast</a></li></ul></li>

<li><a href="Kappa_organism" title="Kappa organism">Kappa organism</a></li>

<li>Organs

<ul><li><a href="Bacteriome" title="Bacteriome">Bacteriome</a></li>

<li><a href="Trophosome" title="Trophosome">Trophosome</a></li></ul></li>

<li><a href="Nitroplast" title="Nitroplast">Nitroplast</a></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%"><a href="Abiogenesis" title="Abiogenesis">Abiogenesis</a></th><td class="navbox-list-with-group navbox-list navbox-even hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Last_universal_common_ancestor" title="Last universal common ancestor">Last universal common ancestor</a></li>

<li><a href="Earliest_known_life_forms" title="Earliest known life forms">Earliest known life forms</a></li>

<ul><li><a href="Ribozyme" title="Ribozyme">Ribozyme</a></li></ul></li>

<li>†<a href="Protocell" title="Protocell">Protocell</a></li>

<li><a href="Coacervate" title="Coacervate">Coacervate</a></li>

<li><a href="Proteinoid" title="Proteinoid">Proteinoid</a></li>

<li><a href="Sulphobes" title="Sulphobes">Sulphobe</a></li>

<li>Research

<ul><li><a href="Model_lipid_bilayer" title="Model lipid bilayer">Model lipid bilayer</a></li>

<li><a href="Jeewanu" title="Jeewanu">Jeewanu</a></li></ul></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="text-align: center;;width:1%">See also</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Organism" title="Organism">Organism</a></li>

<ul><li><a href="Cell_division" title="Cell division">Cell division</a></li>

<li><a href="Artificial_cell" title="Artificial cell">Artificial cell</a></li></ul></li>

<li><a href="Non-cellular_life" title="Non-cellular life">Non-cellular life</a></li>

<li><a href="Synthetic_virology" title="Synthetic virology">Synthetic virus</a>

<ul><li><a href="Viral_vector" title="Viral vector">Viral vector</a></li>

<li><a href="Helper_dependent_virus" title="Helper dependent virus">Helper dependent virus</a></li></ul></li>

<li>?<a href="Nanobacterium" title="Nanobacterium">Nanobacterium</a></li>

<li>?<a href="Nanobe" title="Nanobe">Nanobe</a></li>

<li><a href="Cancer_cell" title="Cancer cell">Cancer cell</a>

<li><a href="Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li>

<li><a href="Virome" title="Virome">Virome</a></li></ul>

</div></td></tr></tbody></table></div>

<div class="navbox-styles"></div><div role="navigation" class="navbox" aria-labelledby="Life,_non-cellular_life,_and_comparable_structures112" style="padding:3px"><table class="nowraplinks mw-collapsible mw-collapsed navbox-inner" style="border-spacing:0;background:transparent;color:inherit"><tbody><tr><th scope="col" class="navbox-title" colspan="2"><div id="Life,_non-cellular_life,_and_comparable_structures112" style="font-size:114%;margin:0 4em"><a href="Life" title="Life">Life</a>, <a href="Non-cellular_life" title="Non-cellular life">non-cellular life</a>, and comparable structures</div></th></tr><tr><th scope="row" class="navbox-group" style="width:1%">Cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">"<a href="Prokaryote" title="Prokaryote">Prokaryota</a>"</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Archaea" title="Archaea">Archaea</a></li>

<li><a href="Bacteria" title="Bacteria">Bacteria</a>

<ul><li><a href="Mitochondrion" title="Mitochondrion">Mitochondrion</a></li>

<li><a href="Plastid" title="Plastid">Plastid</a></li></ul></li>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Eukaryote" title="Eukaryote">Eukaryota</a></th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Animal" title="Animal">Animalia</a></li>

<li><a href="Fungus" title="Fungus">Fungi</a></li>

<li><a href="Plant" title="Plant">Plantae</a></li>

<li>"<a href="Protist" title="Protist">Protista</a>"</li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%"><i><a href="Incertae_sedis" title="Incertae sedis">Incertae sedis</a></i></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Parakaryon" title="Parakaryon">Parakaryon</a></i></li>

<li><a href="Microbial_dark_matter#Microbes_with_highly_unusual_DNA" title="Microbial dark matter">Microbes with highly unusual DNA</a> (?)</li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Non-cellular life</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%"><a href="Virus" title="Virus">Virus</a>

<hr><a href="Viroid" title="Viroid">Viroid</a>

<hr><a href="Satellite_(biology)" title="Satellite (biology)">Satellite</a></th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Realms</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Adnaviria" title="Adnaviria">Adnaviria</a></i></li>

<li><i><a href="Duplodnaviria" title="Duplodnaviria">Duplodnaviria</a></i></li>

<li><i><a href="Monodnaviria" title="Monodnaviria">Monodnaviria</a></i></li>

<li><i><a href="Riboviria" title="Riboviria">Riboviria</a></i></li>

<li><i><a href="Ribozyviria" title="Ribozyviria">Ribozyviria</a></i></li>

<li><i><a href="Varidnaviria" title="Varidnaviria">Varidnaviria</a></i></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Unassigned</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"></div><table class="nowraplinks navbox-subgroup" style="border-spacing:0"><tbody><tr><th scope="row" class="navbox-group" style="width:1%">Classes</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em"><i><a href="Naldaviricetes" title="Naldaviricetes">Naldaviricetes</a></i></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Families</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Alphasatellitidae" class="mw-redirect" title="Alphasatellitidae">Alphasatellitidae</a></i></li>

<li><i><a href="Ampullaviridae" title="Ampullaviridae">Ampullaviridae</a></i></li>

<li><i><a href="Anelloviridae" title="Anelloviridae">Anelloviridae</a></i></li>

<li><i><a href="Avsunviroidae" title="Avsunviroidae">Avsunviroidae</a></i></li>

<li><i><a href="Bicaudaviridae" title="Bicaudaviridae">Bicaudaviridae</a></i></li>

<li><i><a href="Clavaviridae" title="Clavaviridae">Clavaviridae</a></i></li>

<li><i><a href="Finnlakeviridae" title="Finnlakeviridae">Finnlakeviridae</a></i></li>

<li><i><a href="Fuselloviridae" title="Fuselloviridae">Fuselloviridae</a></i></li>

<li><i><a href="Globuloviridae" title="Globuloviridae">Globuloviridae</a></i></li>

<li><i><a href="Guttaviridae" title="Guttaviridae">Guttaviridae</a></i></li>

<li><i><a href="Halspiviridae" title="Halspiviridae">Halspiviridae</a></i></li>

<li><i><a href="Ovaliviridae" title="Ovaliviridae">Ovaliviridae</a></i></li>

<li><i><a href="Plasmaviridae" title="Plasmaviridae">Plasmaviridae</a></i></li>

<li><i><a href="Polydnaviridae" class="mw-redirect" title="Polydnaviridae">Polydnaviridae</a></i></li>

<li><i><a href="Portogloboviridae" title="Portogloboviridae">Portogloboviridae</a></i></li>

<li><i><a href="Pospiviroidae" title="Pospiviroidae">Pospiviroidae</a></i></li>

<li><i><a href="Spiraviridae" title="Spiraviridae">Spiraviridae</a></i></li>

<li><i><a href="Thaspiviridae" title="Thaspiviridae">Thaspiviridae</a></i></li>

<li><i><a href="Tolecusatellitidae" title="Tolecusatellitidae">Tolecusatellitidae</a></i></li></ul>

</div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Genera</th><td class="navbox-list-with-group navbox-list navbox-odd" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><i><a href="Dinodnavirus" title="Dinodnavirus">Dinodnavirus</a></i></li>

<li><i><a href="Rhizidiovirus" title="Rhizidiovirus">Rhizidiovirus</a></i></li></ul>

</div></td></tr></tbody></table><div></div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Other</th><td class="navbox-list-with-group navbox-list navbox-even" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Nanobe" title="Nanobe">Nanobe</a> (?)</li>

<li><a href="Obelisk_(biology)" title="Obelisk (biology)">Obelisk</a></li></ul>

</div></td></tr></tbody></table><div></div></td></tr><tr><th scope="row" class="navbox-group" style="width:1%">Comparable<br>structures</th><td class="navbox-list-with-group navbox-list navbox-odd hlist" style="width:100%;padding:0"><div style="padding:0 0.25em">

<ul><li><a href="Bio-like_structure" title="Bio-like structure">Bio-like structure</a></li>

<li><a href="Cancer_cell" title="Cancer cell">Cancer cell</a>

<li><a href="Clonally_transmissible_cancer" title="Clonally transmissible cancer">Clonally transmissible cancer</a></li></ul></li>

<li><a href="Cosmid" title="Cosmid">Cosmid</a></li>

<li><a href="Defective_interfering_particle" title="Defective interfering particle">Defective interfering particle</a></li>

<li><a href="Endogenous_viral_element" title="Endogenous viral element">Endogenous viral element</a></li>

<li><a href="Fosmid" title="Fosmid">Fosmid</a></li>

<li><a href="Integrative_and_conjugative_element" title="Integrative and conjugative element">Integrative and conjugative element</a></li>

<li><a href="Jeewanu" title="Jeewanu">Jeewanu</a></li>

<li>"<i><a href="Nanobacterium" title="Nanobacterium">Nanobacterium</a></i>"</li>

<li><a href="Phagemid" title="Phagemid">Phagemid</a></li>

<li><a href="Plasmid" title="Plasmid">Plasmid</a></li>

<li><a href="Prion" title="Prion">Prion</a>

<ul><li><a href="Fungal_prion" title="Fungal prion">Fungal prion</a></li></ul></li>

<li><a href="Proteinoid" title="Proteinoid">Proteinoid</a> <a href="Microparticle#Biological_protocells" title="Microparticle">microsphere</a></li>

<li>Retroelements not elsewhere classified

<ul><li><a href="Retron" title="Retron">Retron</a></li>

<li><a href="Diversity-generating_retroelement" title="Diversity-generating retroelement">Diversity-generating retroelement</a></li>

<li><a href="Telomerase_reverse_transcriptase" title="Telomerase reverse transcriptase">Telomerase reverse transcriptase</a></li>

<li><a href="Reverse_transcriptase-related_cellular_gene" class="mw-redirect" title="Reverse transcriptase-related cellular gene">Reverse transcriptase-related cellular gene</a></li></ul></li>

<li><a href="Ribozyme" title="Ribozyme">Ribozyme</a></li>

<li><a href="Spiegelman's_Monster" title="Spiegelman's Monster">Spiegelman's Monster</a></li>

<li><a href="Tandem_repeat" title="Tandem repeat">Tandem repeat</a></li>

<li>

<ul><li><a href="Retroposon" title="Retroposon">Retroposon</a></li></ul></li>

<li><a href="Transpoviron" title="Transpoviron">Transpoviron</a></li>

<li><a href="Xenobot" title="Xenobot">Xenobot</a></li></ul>

</div></td></tr></tbody></table></div>

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