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<span id="openzim-page-title" class="mw-page-title-main"><span class="mw-page-title-main">Alu element</span></span>
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<p>An <b>Alu element</b> is a short stretch of <a href="DNA" title="DNA">DNA</a> originally characterized by the action of the <i><a href="Arthrobacter_luteus" title="Arthrobacter luteus">Arthrobacter luteus</a> (Alu)</i> <a href="Restriction_endonuclease" class="mw-redirect" title="Restriction endonuclease">restriction endonuclease</a>.<sup id="cite_ref-pmid1052772_1-0" class="reference"><a href="#cite_note-pmid1052772-1"><span class="cite-bracket">[</span>1<span class="cite-bracket">]</span></a></sup> <i>Alu</i> elements are the most abundant <a href="Transposable_element" title="Transposable element">transposable elements</a> in the <a href="Human_genome" title="Human genome">human genome</a>, present in excess of one million copies.<sup id="cite_ref-pmid9694261_2-0" class="reference"><a href="#cite_note-pmid9694261-2"><span class="cite-bracket">[</span>2<span class="cite-bracket">]</span></a></sup> Most <i>Alu</i> elements are thought to be selfish or parasitic DNA. However, it has been suggested that at least some are likely to play a role in evolution and have been used as <a href="Genetic_marker" title="Genetic marker">genetic markers</a>.<sup id="cite_ref-pmid11263730_3-0" class="reference"><a href="#cite_note-pmid11263730-3"><span class="cite-bracket">[</span>3<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-Pray_4-0" class="reference"><a href="#cite_note-Pray-4"><span class="cite-bracket">[</span>4<span class="cite-bracket">]</span></a></sup> They are derived from the small cytoplasmic <a href="7SL_RNA" class="mw-redirect" title="7SL RNA">7SL RNA</a>, a component of the <a href="Signal_recognition_particle" title="Signal recognition particle">signal recognition particle</a>. <i>Alu</i> elements are not highly conserved within primate <a href="Genome" title="Genome">genomes</a>, as only a minority have retained activity, and originated in the genome of an ancestor of <a href="Supraprimates" class="mw-redirect" title="Supraprimates">Supraprimates</a>.<sup id="cite_ref-pmid17307271_5-0" class="reference"><a href="#cite_note-pmid17307271-5"><span class="cite-bracket">[</span>5<span class="cite-bracket">]</span></a></sup>
</p><p><i>Alu</i> insertions have been implicated in several inherited human diseases and in various forms of cancer.
</p><p>The study of Alu elements has also been important in elucidating human <a href="Population_genetics" title="Population genetics">population genetics</a> and the <a href="Evolution" title="Evolution">evolution</a> of <a href="Primate" title="Primate">primates</a>, including the <a href="Human_evolution" title="Human evolution">evolution of humans</a>.
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<div class="mw-heading mw-heading2"><h2 id="Alu_family">Alu family</h2></div>
<p>The Alu family is a family of repetitive elements in <a href="Primate" title="Primate">primate</a> genomes, including the <a href="Human" title="Human">human</a> <a href="Genome" title="Genome">genome</a>.<sup id="cite_ref-6" class="reference"><a href="#cite_note-6"><span class="cite-bracket">[</span>6<span class="cite-bracket">]</span></a></sup> Modern <i>Alu</i> elements are about 300 <a href="Base_pair" title="Base pair">base pairs</a> long and are therefore classified as <a href="Short_interspersed_nuclear_element" title="Short interspersed nuclear element">short interspersed nuclear elements</a> (SINEs) among the class of repetitive RNA elements. The typical structure is 5' - Part A - A5TACA6 - Part B - PolyA Tail - 3', where Part A and Part B (also known as "left arm" and "right arm") are similar nucleotide sequences. Expressed another way, it is believed modern <i>Alu</i> elements emerged from a head to tail fusion of two distinct FAMs (fossil antique monomers) over 100 million years ago, hence its dimeric structure of two similar, but distinct monomers (left and right arms) joined by an A-rich linker. Both monomers are thought to have evolved from 7SL, also known as <a href="SRP_RNA" class="mw-redirect" title="SRP RNA">SRP RNA</a>.<sup id="cite_ref-pmid17020921_7-0" class="reference"><a href="#cite_note-pmid17020921-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup> The length of the polyA tail varies between <i>Alu</i> families.
</p><p>There are over one million <i>Alu</i> elements interspersed throughout the human genome, and it is estimated that about 10.7% of the human genome consists of <i>Alu</i> sequences. However, less than 0.5% are <a href="Polymorphism_(biology)" title="Polymorphism (biology)">polymorphic</a> (i.e., occurring in more than one form or morph).<sup id="cite_ref-pmid11560904_8-0" class="reference"><a href="#cite_note-pmid11560904-8"><span class="cite-bracket">[</span>8<span class="cite-bracket">]</span></a></sup> In 1988, <a href="Jerzy_Jurka" title="Jerzy Jurka">Jerzy Jurka</a> and <a href="Temple_Smith" class="mw-redirect" title="Temple Smith">Temple Smith</a> discovered that <i>Alu</i> elements were split in two major subfamilies known as AluJ (named after Jurka) and AluS (named after Smith), and other Alu subfamilies were also independently discovered by several groups.<sup id="cite_ref-pmid3387438_9-0" class="reference"><a href="#cite_note-pmid3387438-9"><span class="cite-bracket">[</span>9<span class="cite-bracket">]</span></a></sup> Later on, a sub-subfamily of AluS which included active Alu elements was given the separate name AluY. Dating back 65 million years, the AluJ lineage is the oldest and least active in the human genome. The younger AluS lineage is about 30 million years old and still contains some active elements. Finally, the AluY elements are the youngest of the three and have the greatest disposition to move along the human genome.<sup id="cite_ref-pmid18836035_10-0" class="reference"><a href="#cite_note-pmid18836035-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup> The discovery of <i>Alu</i> subfamilies led to the hypothesis of master/source genes, and provided the definitive link between transposable elements (active elements) and interspersed repetitive DNA (mutated copies of active elements).<sup id="cite_ref-pmid1774786_11-0" class="reference"><a href="#cite_note-pmid1774786-11"><span class="cite-bracket">[</span>11<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading3"><h3 id="Related_elements">Related elements</h3></div>
<p>B1 elements in rats and mice are similar to Alus in that they also evolved from 7SL RNA, but they only have one left monomer arm. 95% percent of human Alus are also found in chimpanzees, and 50% of B elements in mice are also found in rats. These elements are mostly found in introns and upstream regulatory elements of genes.<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>The ancestral form of Alu and B1 is the fossil Alu monomer (FAM). Free-floating forms of the left and right arms exist, termed free left Alu monomers (FLAMs) and free right Alu monomers (FRAMs) respectively.<sup id="cite_ref-13" class="reference"><a href="#cite_note-13"><span class="cite-bracket">[</span>13<span class="cite-bracket">]</span></a></sup> A notable FLAM in primates is the <a href="BC200_lncRNA" title="BC200 lncRNA">BC200 lncRNA</a>.
</p>
<div class="mw-heading mw-heading2"><h2 id="Sequence_features">Sequence features</h2></div>
<p>Two main promoter "boxes" are found in Alu: a 5' A box with the consensus <style data-mw-deduplicate="TemplateStyles:r886049734">
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</style><span class="monospaced">TGGCTCACGCC</span>, and a 3' B box with the consensus <span class="monospaced">GTTCGAGAC</span> (IUPAC <a href="Nucleic_acid_notation" title="Nucleic acid notation">nucleic acid notation</a>). <a href="TRNA" class="mw-redirect" title="TRNA">tRNAs</a>, which are transcribed by <a href="RNA_polymerase_III" title="RNA polymerase III">RNA polymerase III</a>, have a similar but stronger promoter structure.<sup id="cite_ref-14" class="reference"><a href="#cite_note-14"><span class="cite-bracket">[</span>14<span class="cite-bracket">]</span></a></sup> Both boxes are located in the left arm.<sup id="cite_ref-pmid17020921_7-1" class="reference"><a href="#cite_note-pmid17020921-7"><span class="cite-bracket">[</span>7<span class="cite-bracket">]</span></a></sup>
</p><p>Alu elements contain four or fewer <a href="Retinoic_Acid" class="mw-redirect" title="Retinoic Acid">Retinoic Acid</a> response element hexamer sites in its internal <a href="Promoter_(biology)" class="mw-redirect" title="Promoter (biology)">promoter</a>, with the last one overlapping with the "B box".<sup id="cite_ref-pmid7667273_15-0" class="reference"><a href="#cite_note-pmid7667273-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup> In this 7SL (<a href="Signal_recognition_particle_RNA" title="Signal recognition particle RNA">SRP</a>) RNA example below, functional hexamers are underlined using a solid line, with the non-functional third hexamer denoted using a dotted line:
</p><p><span class="monospaced"><span style="line-break: anywhere">GCCGGGCGCGGTGGCGCGTGCCTGTAGTCCCAGCTACTCGGG<u>AGGCTG</u>AGGCTGGA<u>GGATCG</u>CTTG<u style="text-decoration-style: dotted;">AGTCCA</u>GG<b><u>AGTTCT</u>GGGCT</b>GTAGTGCGCTATGCCGATCGGAATAGCCACTGCACTCCAGCCTGGGCAACATAGCGAGACCCCGTCTC</span></span>.
</p><p>The recognition sequence of the <i><a href="Arthrobacter_luteus" title="Arthrobacter luteus">Alu I</a></i> endonuclease is 5' ag/ct 3'; that is, the enzyme cuts the DNA segment between the <a href="Guanine" title="Guanine">guanine</a> and <a href="Cytosine" title="Cytosine">cytosine</a> residues (in lowercase above).<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>
</p>
<div class="mw-heading mw-heading2"><h2 id="Alu_elements">Alu elements</h2></div>
<p>Some <i>Alu</i> elements are responsible for regulation of tissue-specific genes. Others are involved in the transcription of nearby genes and can sometimes change the way a gene is expressed.<sup id="cite_ref-pmid8790336_17-0" class="reference"><a href="#cite_note-pmid8790336-17"><span class="cite-bracket">[</span>17<span class="cite-bracket">]</span></a></sup>
</p><p><i>Alu</i> elements are <a href="Retrotransposon" title="Retrotransposon">retrotransposons</a> and look like DNA copies made from <a href="RNA_polymerase_III" title="RNA polymerase III">RNA polymerase III</a>-encoded RNAs. <i>Alu</i> elements do not encode for protein products. They are replicated as any other DNA sequence, but depend on <a href="Long_interspersed_nuclear_element" title="Long interspersed nuclear element">LINE</a> retrotransposons for generation of new elements, thus providing an easy explanation for their presence in large numbers in primate genomes.<sup id="cite_ref-pmid16344113_18-0" class="reference"><a href="#cite_note-pmid16344113-18"><span class="cite-bracket">[</span>18<span class="cite-bracket">]</span></a></sup>
</p><p><i>Alu</i> element replication and mobilization begins by interactions with <a href="Signal_recognition_particle" title="Signal recognition particle">signal recognition particles</a> (SRPs), which aid newly translated proteins to reach their final destinations.<sup id="cite_ref-pmid11089964_19-0" class="reference"><a href="#cite_note-pmid11089964-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> <i>Alu</i> RNA forms a specific RNA:protein complex with a protein heterodimer consisting of SRP9 and SRP14.<sup id="cite_ref-pmid11089964_19-1" class="reference"><a href="#cite_note-pmid11089964-19"><span class="cite-bracket">[</span>19<span class="cite-bracket">]</span></a></sup> SRP9/14 facilitates <i>Alu</i>'s attachment to ribosomes that capture nascent <a href="LINE1" title="LINE1">L1 proteins</a>. Thus, an <i>Alu</i> element can take control of the L1 protein's <a href="Reverse_transcriptase" title="Reverse transcriptase">reverse transcriptase</a>, ensuring that the <i>Alu</i>'s RNA sequence gets copied into the genome rather than the L1's mRNA.<sup id="cite_ref-pmid18836035_10-1" class="reference"><a href="#cite_note-pmid18836035-10"><span class="cite-bracket">[</span>10<span class="cite-bracket">]</span></a></sup>
</p><p><i>Alu</i> elements in primates form a fossil record that is relatively easy to decipher because <i>Alu</i> element insertion events have a characteristic signature that is both easy to read and faithfully recorded in the genome from generation to generation. The study of <i>Alu Y</i> elements (the more recently evolved) thus reveals details of ancestry because individuals will most likely only share a particular <i>Alu</i> element insertion if they have a common ancestor. This is because insertion of an Alu element occurs only 100 - 200 times per million years, and no known mechanism for the targeted deletion of one has been found. Therefore, individuals with an element likely descended from an ancestor with one—and vice versa, for those without. In genetics, the presence or lack thereof of a recently inserted <i>Alu</i> element may be a good property to consider when studying human evolution.<sup id="cite_ref-20" class="reference"><a href="#cite_note-20"><span class="cite-bracket">[</span>20<span class="cite-bracket">]</span></a></sup> Most human <i>Alu</i> element insertions can be found in the corresponding positions in the genomes of other primates, but about 7,000 <i>Alu</i> insertions are unique to humans.<sup id="cite_ref-pmid16136131_21-0" class="reference"><a href="#cite_note-pmid16136131-21"><span class="cite-bracket">[</span>21<span class="cite-bracket">]</span></a></sup>
</p>
<div class="mw-heading mw-heading2"><h2 id="Impact_in_humans">Impact in humans</h2></div>
<p>Some <i>Alu</i> elements have been proposed to affect <a href="Gene_expression" title="Gene expression">gene expression</a> and been found to contain functional <a href="Promoter_(genetics)" title="Promoter (genetics)">promoter</a> regions for <a href="Steroid_hormone_receptor" title="Steroid hormone receptor">steroid hormone receptors</a>.<sup id="cite_ref-pmid7667273_15-1" class="reference"><a href="#cite_note-pmid7667273-15"><span class="cite-bracket">[</span>15<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid7559405_22-0" class="reference"><a href="#cite_note-pmid7559405-22"><span class="cite-bracket">[</span>22<span class="cite-bracket">]</span></a></sup> Due to the abundant content of <a href="CpG_dinucleotides" class="mw-redirect" title="CpG dinucleotides">CpG dinucleotides</a> found in <i>Alu</i> elements, these regions can serve as a site of <a href="DNA_methylation" title="DNA methylation">methylation</a>, contributing to up to 30% of the methylation sites in the human genome.<sup id="cite_ref-pmid9753719_23-0" class="reference"><a href="#cite_note-pmid9753719-23"><span class="cite-bracket">[</span>23<span class="cite-bracket">]</span></a></sup> <i>Alu</i> elements are also a common source of mutations in humans; however, such mutations are often confined to non-coding regions of pre-mRNA (<a href="Introns" class="mw-redirect" title="Introns">introns</a>), where they have little discernible impact on the bearer.<sup id="cite_ref-pmid11237011_24-0" class="reference"><a href="#cite_note-pmid11237011-24"><span class="cite-bracket">[</span>24<span class="cite-bracket">]</span></a></sup> Mutations in the introns (or non-coding regions of RNA) have little or no effect on phenotype of an individual if the coding portion of individual's genome does not contain mutations. When Alu insertions occur in coding regions (<a href="Exons" class="mw-redirect" title="Exons">exons</a>), or into mRNA after the process of splicing, they're typically detrimental to the host organism.<sup id="cite_ref-pmid10381326_25-0" class="reference"><a href="#cite_note-pmid10381326-25"><span class="cite-bracket">[</span>25<span class="cite-bracket">]</span></a></sup>
</p><p>However, the variation generated can be used in studies of the movement and ancestry of human populations,<sup id="cite_ref-pmid11988762_26-0" class="reference"><a href="#cite_note-pmid11988762-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup> and the mutagenic effect of <i>Alu</i><sup id="cite_ref-pmid21282640_27-0" class="reference"><a href="#cite_note-pmid21282640-27"><span class="cite-bracket">[</span>27<span class="cite-bracket">]</span></a></sup> and retrotransposons in general<sup id="cite_ref-pmid19763152_28-0" class="reference"><a href="#cite_note-pmid19763152-28"><span class="cite-bracket">[</span>28<span class="cite-bracket">]</span></a></sup> has played a major role in the evolution of the human genome. There are also a number of cases where <i>Alu</i> insertions or deletions are associated with specific effects in humans:
</p>
<div class="mw-heading mw-heading3"><h3 id="Associations_with_human_disease">Associations with human disease</h3></div>
<p><i>Alu</i> insertions are sometimes disruptive and can result in inherited disorders. However, most <i>Alu</i> variation acts as markers that segregate with the disease so the presence of a particular <i>Alu</i> <a href="Allele" title="Allele">allele</a> does not mean that the carrier will definitely get the disease. The first report of <i>Alu</i>-mediated <a href="Genetic_recombination" title="Genetic recombination">recombination</a> causing a prevalent inherited predisposition to cancer was a 1995 report about <i>hereditary nonpolyposis <a href="Colorectal_cancer" title="Colorectal cancer">colorectal cancer</a></i>.<sup id="cite_ref-pmid7584997_29-0" class="reference"><a href="#cite_note-pmid7584997-29"><span class="cite-bracket">[</span>29<span class="cite-bracket">]</span></a></sup> In the human genome, the most recently active have been the 22 AluY and 6 AluS Transposon Element subfamilies due to their inherited activity to cause various cancers. Thus due to their major heritable damage it is important to understand the causes that affect their transpositional activity.<sup id="cite_ref-pmid29219079_30-0" class="reference"><a href="#cite_note-pmid29219079-30"><span class="cite-bracket">[</span>30<span class="cite-bracket">]</span></a></sup>
</p><p>The following human diseases have been linked with <i>Alu</i> insertions:<sup id="cite_ref-pmid11988762_26-1" class="reference"><a href="#cite_note-pmid11988762-26"><span class="cite-bracket">[</span>26<span class="cite-bracket">]</span></a></sup><sup id="cite_ref-pmid22204421_31-0" class="reference"><a href="#cite_note-pmid22204421-31"><span class="cite-bracket">[</span>31<span class="cite-bracket">]</span></a></sup>
</p>
<ul><li><a href="Alport_syndrome" title="Alport syndrome">Alport syndrome</a></li>
<li><a href="Breast_cancer" title="Breast cancer">Breast cancer</a></li>
<li>chorioretinal degeneration</li>
<li><a href="Diabetes_mellitus_type_II" class="mw-redirect" title="Diabetes mellitus type II">Diabetes mellitus type II</a></li>
<li><a href="Ewing's_sarcoma" class="mw-redirect" title="Ewing's sarcoma">Ewing's sarcoma</a></li>
<li><a href="Familial_hypercholesterolemia" title="Familial hypercholesterolemia">Familial hypercholesterolemia</a></li>
<li><a href="Hemophilia" class="mw-redirect" title="Hemophilia">Hemophilia</a></li>
<li><a href="Leigh_syndrome" title="Leigh syndrome">Leigh syndrome</a></li>
<li><a href="Mucopolysaccharidosis" title="Mucopolysaccharidosis">mucopolysaccharidosis</a> VII</li>
<li><a href="Neurofibromatosis" title="Neurofibromatosis">Neurofibromatosis</a></li>
<li><a href="Macular_degeneration" title="Macular degeneration">Macular degeneration</a><sup id="cite_ref-pnas_32-0" class="reference"><a href="#cite_note-pnas-32"><span class="cite-bracket">[</span>32<span class="cite-bracket">]</span></a></sup></li></ul>
<p>And the following diseases have been associated with <a href="Single-nucleotide_DNA_variation" class="mw-redirect" title="Single-nucleotide DNA variation">single-nucleotide DNA variations</a> in Alu elements affecting transcription levels:<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>
<ul><li><a href="Alzheimer's_disease" title="Alzheimer's disease">Alzheimer's disease</a></li>
<li><a href="Lung_cancer" title="Lung cancer">Lung cancer</a></li>
<li><a href="Gastric_cancer" class="mw-redirect" title="Gastric cancer">Gastric cancer</a></li></ul>
<p>The following disease have been associated with repeat expansion of AAGGG pentamere in Alu element :
</p>
<ul><li><a href="RFC1" title="RFC1">RFC1</a> mutation responsible of <a href="Cerebellar_ataxia%2C_neuropathy%2C_vestibular_areflexia_syndrome" title="Cerebellar ataxia, neuropathy, vestibular areflexia syndrome">CANVAS</a> (Cerebellar Ataxia, Neuropathy & Vestibular Areflexia Syndrome) <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></li></ul>
<div class="mw-heading mw-heading3"><h3 id="Associated_human_mutations">Associated human mutations</h3></div>
<ul><li>The <i>ACE</i> gene, encoding <a href="Angiotensin-converting_enzyme" title="Angiotensin-converting enzyme">angiotensin-converting enzyme</a>, has 2 common variants, one with an <i>Alu</i> insertion (<i>ACE</i>-I) and one with the <i>Alu</i> deleted (<i>ACE</i>-D). This variation has been linked to changes in sporting ability: the presence of the <i>Alu</i> element is associated with better performance in endurance-oriented events (e.g. triathlons), whereas its absence is associated with strength- and power-oriented performance.<sup id="cite_ref-pmid21615186_35-0" class="reference"><a href="#cite_note-pmid21615186-35"><span class="cite-bracket">[</span>35<span class="cite-bracket">]</span></a></sup></li>
<li>The <a href="Opsin" title="Opsin">opsin</a> <a href="Gene_duplication" title="Gene duplication">gene duplication</a> which <a href="Evolution_of_color_vision_in_primates" title="Evolution of color vision in primates">resulted</a> in the re-gaining of <a href="Trichromacy" title="Trichromacy">trichromacy</a> in <a href="Catarrhini" title="Catarrhini">Old World primates</a> (including humans) is flanked by an <i>Alu</i> element,<sup id="cite_ref-pmid10413401_36-0" class="reference"><a href="#cite_note-pmid10413401-36"><span class="cite-bracket">[</span>36<span class="cite-bracket">]</span></a></sup> implicating the role of <i>Alu</i> in the evolution of three colour vision.</li></ul>
<div class="mw-heading mw-heading2"><h2 id="References">References</h2></div>
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</ol></div>
<div class="mw-heading mw-heading2"><h2 id="External_links">External links</h2></div>
<ul><li><a rel="nofollow" class="external text" href="https://meshb.nlm.nih.gov/record/ui?name=Alu+Repetitive+Sequences">Alu+Repetitive+Sequences</a> at the U.S. National Library of Medicine <a href="Medical_Subject_Headings" title="Medical Subject Headings">Medical Subject Headings</a> (MeSH)</li>
<li><cite class="citation journal cs1"><a rel="nofollow" class="external text" href="https://www.ncbi.nlm.nih.gov/nuccore/NR_002715.1">"NCBI Genbank DNA encoding 7SL RNA"</a>. <i>National Center for Biotechnology Information</i>. 2018-05-12.</cite></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="Transposable_element" title="Transposable element">transposon</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%"><a href="Transposable_element" title="Transposable element">Transposon</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="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="Mammalian-wide_interspersed_repeat" title="Mammalian-wide interspersed repeat">MIR</a></li></ul>
</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">
<ul><li><a href="LINE1" title="LINE1">LINE1</a></li>
<li>LINE2</li></ul>
</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">
<ul><li><a href="Human_endogenous_retroviruses" class="mw-redirect" title="Human endogenous retroviruses">HERV</a></li>
<li>MER4</li>
<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>Dada</li>
<li>EnSpm/CACTA</li>
<li>Ginger1</li>
<li>Ginger2</li>
<li>Harbinger</li>
<li><a href="HAT_transposon" title="HAT transposon">hAT</a></li>
<li><a href="Helitron_(biology)" title="Helitron (biology)">Helitron</a></li>
<li>IS3EU</li>
<li>ISL2EU</li>
<li>Kolobok</li>
<li><a href="Tc1/mariner" title="Tc1/mariner">Tc1/mariner</a></li>
<li>Merlin</li>
<li>MuDR</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>Sola</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>
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