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Short interspersed nuclear element

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452:. SINEs however should not be mistaken as RNA pseudogenes. In general, pseudogenes are generated when processed mRNAs of protein-coding genes are reverse-transcribed and incorporated back into the genome (RNA pseudogenes are reverse transcribed RNA genes). Pseudogenes are generally functionless as they descend from processed RNAs independent of their evolutionary-context which includes introns and different regulatory elements which enable transcription and processing. These pseudogenes, though non-functional may in some cases still possess promoters, CpG islands, and other features which enable transcription; they thus can still be transcribed and may possess a role in the regulation of gene expression (like SINEs and other non-coding elements). Pseudogenes thus differ from SINEs in that they are derived from transcribed- functional RNA whereas SINEs are DNA elements which retrotranspose by co-opting RNA genes transcriptional machinery. However, there are studies which suggest that retro-transposable elements such as short-interspersed nuclear elements are not only capable of copying themselves in alternate regions in the genome but are also able to do so for random genes too. Thus SINEs can be playing a vital role in the generation of pseudogenes, which themselves are known to be involved in regulatory networks. This is perhaps another means by which SINEs have been able to influence and contribute to gene-regulation. 316:
element silencing in fact occurred before L1 long-interspersed nuclear element extinction; this is due to the fact that B1 SINEs are silenced in the genus most-closely related to the genus which does not contain active L1 LINEs (though the genus with B1 SINE silencing still contains active L1 LINEs). Another genus was also found which similarly contained active L1 long-interspersed nuclear elements but did not contain B1 short-interspersed nuclear elements; the opposite scenario, in which active B1 SINEs were present in a genus which did not possess active L1 LINEs was not found. This result was expected and strongly supports the theory that SINEs have evolved to co-opt the RNA-binding proteins, endonucleases, and reverse-transcriptases coded by LINEs. In taxa which do not actively transcribe and translate long-interspersed nuclear elements protein-products, SINEs do not have the theoretical foundation by which to retrotranspose within the genome. The results obtained in Rinehart et al. are thus very supportive of the current model of SINE retrotransposition.
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that had high hybridization E-values were genes particularly involved in metabolic and signaling pathways. Almost all miRNAs identified to have a strong ability to hybridize to putative V-SINE sequence motifs in genes have been identified (in mammals) to have regulatory roles. These results which establish a correlation between short-interspersed nuclear elements and different regulatory microRNAs strongly suggest that V-SINEs have a significant role in attenuating responses to different signals and stimuli related to metabolism, proliferation and differentiation. Many other studies must be undertaken to establish the validity and extent of short-interspersed nuclear element retrotransposons' role in regulatory gene-expression networks. In conclusion, though not much is known about the role and mechanism by which SINEs generate miRNA gene loci it is generally understood that SINEs have played a significant evolutionary role in the creation of "RNA-genes", this is also touched upon above in SINEs and pseudogenes.
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regulation can occur in different ways: the RNA transcript can directly bind to the transcription factor as a co-regulator; also, the RNA can regulate and modify the ability of co-regulators to associate with the transcription factor. For example, Evf-2, a certain long non-coding RNA, has been known to function as a co-activator for certain homeobox transcription factors which are critical to nervous system development and organization. Furthermore, RNA transcripts can interfere with the functionality of the transcriptional complex by interacting or associating with RNA polymerases during the transcription or loading processes. Moreover, non-coding RNAs like SINEs can bind or interact directly with the DNA duplex coding the gene and thus prevent its transcription.
440:, a tumor suppressor implicated in multiple forms of cancer, namely breast cancer. Furthermore, studies have established a strong correlation between transcriptional mobilization of SINEs and certain cancers and conditions such as hypoxia; this can be due to the genomic instability caused by SINE activity as well as more direct-downstream effects. SINEs have also been implicated in countless other diseases. In essence, short-interspersed nuclear elements have become deeply integrated in countless regulatory, metabolic and signaling pathways and thus play an inevitable role in causing disease. Much is still to be known about these genomic parasites but it is clear they play a significant role within eukaryotic organisms. 420:
gene expression. A microRNA is a non-coding RNA generally 22 nucleotides in length. This non-protein coding oligonucleotide is itself coded by longer nuclear DNA sequence usually transcribed by RNA polymerase II which is also responsible for the transcription of most mRNAs and snRNAs in eukaryotes. However, some research suggests that some microRNAs that possess upstream short-interspersed nuclear elements are transcribed by RNA polymerase III which is widely implicated in ribosomal RNA and tRNA, two transcripts vital to mRNA translation. This provides an alternate mechanism by which short-interspersed nuclear elements could be interacting with or mediating gene-regulatory networks involving microRNAs.
351:, short-interspersed nuclear element of about 300 nucleotides, are the most common SINE in humans, with >1,000,000 copies throughout the genome, which is over 10 percent of the total genome; this is not uncommon among other species. Alu element copy number differences can be used to distinguish between and construct phylogenies of primate species. Canines differ primarily in their abundance of SINEC_Cf repeats throughout the genome, rather than other gene or allele level mutations. These dog-specific SINEs may code for a splice acceptor site, altering the sequences that appear as exons or introns in each species. 218:, and other associated proteins to different degrees. Furthermore, the shape and density of certain areas of a chromosome can affect the shape and density of neighboring (or even distant regions) on the chromosome through interaction facilitated by different proteins and elements. Non-coding RNAs such as short-interspersed nuclear elements, which have been known to associate with and contribute to chromatin structure, can thus play huge role in regulating gene expression. Short-interspersed-nuclear-elements similarly can be involved in gene regulation by modifying genomic architecture. 312:
and primates show very strong homology at the insertion-site motif. Such evidence is a basis for the proposed mechanism in which integration of the SINE transcript can be co-opted with LINE-coded protein products. This is specifically demonstrated by a detailed analysis of over 20 rodent species profiled LINEs and SINEs, mainly L1s and B1s respectively; these are families of LINEs and SINEs found at high frequencies in rodents along with other mammals. The study sought to provide phylogenetic clarity within the context of LINE and SINE activity.
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remarkably successful at persisting and amplifying (through retrotransposition) within the genomes of eukaryotes. These "parasites" which have become ubiquitous in genomes can be very deleterious to organisms as discussed below. However, eukaryotes have been able to integrate short-interspersed nuclear elements into different signaling, metabolic and regulatory pathways and SINEs have become a great source of genetic variability. They seem to play a particularly important role in the regulation of
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double-stranded DNA breaks (rather than the endonuclease coded by related long-interspersed nuclear elements creating an insertion-site). These DNA breaks are utilized to prime reverse transcriptase, ultimately integrating the SINE transcript back into the genome. SINEs nonetheless depend on enzymes coded by other DNA elements and are thus known as non-autonomous retrotransposons as they depend on the machinery of LINEs, which are known as autonomous retrotransposons.<
189:. SINEs, like tRNAs and many small-nuclear RNAs possess an internal promoter and thus are transcribed differently than most protein-coding genes. In other words, short-interspersed nuclear elements have their key promoter elements within the transcribed region itself. Though transcribed by RNA polymerase III, SINEs and other genes possessing internal promoters, recruit different transcriptional machinery and factors than genes possessing upstream promoters. 424:
gene-expression. Furthermore, Scarpato et al. reveals (as discussed above) that genes predicted to possess short-interspersed nuclear elements (SINEs) through sequence analysis were targeted and hybridized by microRNAs significantly greater than other genes. This provides an evolutionarily path by which the parasitic SINEs were co-opted and utilized to form RNA-genes (such as microRNAs) which have evolved to play a role in complex gene-regulatory networks.
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networks (as discussed in SINEs as long non-coding RNAs) is crucial to beginning to understand the relationship between SINEs and certain diseases. Multiple studies have suggested that increased SINE activity is correlated with certain gene-expression profiles and post-transcription regulation of certain genes. In fact, Peterson et al. 2013 demonstrated that high SINE RNA expression correlates with post-transcriptional downregulation of
230:. The distribution of SINEs to genes was significantly more similar than that of other non-coding genetic elements and even differed significantly from the distribution of long-interspersed nuclear elements. This suggested that the SINE distribution was not a mere accident caused by LINE-mediated retrotransposition but rather that SINEs possessed a role in gene-regulation. Furthermore, SINEs frequently contain motifs for 17: 428:(DGCR8) which recruits and associates with the Drosha protein. This complex is responsible for cleaving some of the hair-pin structures from the pre-microRNA which is transported to the cytoplasm. The pre-miRNA is processed by the protein DICER into a double stranded 22 nucleotide. Thereafter, one of the strands is incorporated into a multi-protein 411:). The specific family of SINEs being examined was the Anamnia V-SINEs; this family of short interspersed nuclear elements is often found in the untranslated region of the 3' end of many genes and is present in vertebrate genomes. The study involved a computational analysis in which the genomic distribution and activity of the Anamnia V-SINEs in 239:(gene-silencing state). Thus, the analysis suggests that short-interspersed nuclear elements can function as a ‘signal-booster' in the polycomb-dependent silencing of gene-sets through chromatin re-organization. In essence, it is the cumulative effect of many types of interactions that leads to the difference between 254:
Also, many non-coding RNAs are distributed near protein-coding genes, often in the reverse direction. This is especially true for short-interspersed nuclear elements as seen in Usmanova et al. These non-coding RNAs, which lie adjacent to or overlap gene-sets provide a mechanism by which transcription
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With such evidence suggesting that short-interspersed nuclear elements have been evolutionary sources for microRNA loci generation it is important to further discuss the potential relationships between the two as well as the mechanism by which the microRNA regulates RNA degradation and more broadly,
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The theory that short-interspersed nuclear elements have evolved to utilize the retrotransposon machinery of long-interspersed nuclear elements is supported by studies which examine the presence and distribution of LINEs and SINEs in taxa of different species. For example, LINEs and SINEs in rodents
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SINEs are known to share sequence homology with LINES which gives a basis by which the LINE machinery can reverse transcribe and integrate SINE transcripts. Alternately, some SINEs are believed to use a much more complex system of integrating back into the genome; this system involves the use random
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In conclusion, non-coding RNAs such as SINEs are capable of affecting gene expression on a multitude of different levels and in different ways. Short-interspersed nuclear elements are believed to be deeply integrated into a complex regulatory network capable of fine-tuning gene expression across the
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zebrafish was examined; furthermore, these V-SINEs potential to generate novel microRNA loci was analyzed. It was found that genes which were predicted to possess V-SINEs were targeted by microRNAs with significantly higher hybridization E-values (relative to other areas in the genome). The genes
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transcriptional repressor is discussed above. Alternatively, it also provides a mechanism by which local gene expression can be curtailed and regulated because the transcriptional complexes can hinder or prevent nearby genes from being transcribed. There is research to suggest that this phenomenon
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Understanding the different ways in which microRNA regulates gene-expression, including mRNA-translation and degradation is key to understanding the potential evolutionary role of SINEs in gene-regulation and in the generation of microRNA loci. This, in addition to SINEs' direct role in regulatory
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of short-interspersed nuclear elements and is evolutionarily derived from an RNA synthesized by RNA Polymerase III such as ribosomal RNAs and tRNAs; the 5' head is indicative of which endogenous element that SINE was derived from and was able to parasitically utilize its transcriptional machinery.
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In essence, short interspersed nuclear elements are genetic parasites which have evolved very early in the history of eukaryotes to utilize protein machinery within the organism as well as to co-opt the machinery from similarly parasitic genomic elements. The simplicity of these elements make them
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The study arrived at a candidate taxa believed to be the first instance of L1 LINE extinction; it expectedly discovered that there was no evidence to suggest that B1 SINE activity occurred in species which did not have L1 LINE activity. Also, the study suggested that B1 short-interspersed nuclear
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In addition to directly affecting chromatin structure, there are a number of ways in which SINEs can potentially regulate gene expression. For example, long non-coding RNA can directly interact with transcriptional repressors and activators, attenuating or modifying their function. This type of
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The microRNAs are transcribed as part of longer RNA strands of generally about 80 nucleotides which through complementary base-pairing are able to form hairpin loop structures These structures are recognized and processed in the nucleus by the nuclear protein DiGeorge Syndrome Critical Region 8
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Apart from mammals, SINEs can reach high copy numbers in a range of species, including nonbony vertebrates (elephant shark) and some fish species (coelacanths). In plants, SINEs are often restricted to closely related species and have emerged, decayed, and vanished frequently during evolution.
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polycomb proteins. YY1 is a zinc-finger protein that acts as a transcriptional repressor for a wide-variety of genes essential for development and signaling. Polycomb protein YY1 is believed to mediate the activity of histone deacetylases and histone acetyltransferases to facilitate chromatin
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The regions coding miRNA can be independent RNA-genes often being anti-sense to neighboring protein-coding genes, or can be found within the introns of protein-coding genes. The co-localization of microRNA and protein-coding genes provides a mechanistic foundation by which microRNA regulates
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of SINEs is composed of short simple repeats of varying lengths; these simple repeats are sites where two (or more) short-interspersed nuclear elements can combine to form a dimeric SINE. Short-interspersed nuclear elements which only possess a head and tail are called simple SINEs whereas
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1 (ORF 1) encodes a protein which binds to RNA and acts as a chaperone to facilitate and maintain the LINE protein-RNA complex structure. Open reading frame 2 (ORF 2) codes a protein which possesses both endonuclease and reverse transcriptase activities. This enables the LINE mRNA to be
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origins in eukaryotic genomes. These SINEs have mutated and replicated themselves a large number of times on an evolutionary time-scale and thus form many different lineages. Their early evolutionary origin has caused them to be ubiquitous in many eukaryotic lineages.
122:. There are three types of SINEs common to vertebrates and invertebrates: CORE-SINEs, V-SINEs, and AmnSINEs. SINEs have 50-500 base pair internal regions which contain a tRNA-derived segment with A and B boxes that serve as an internal promoter for 280:(LINEs), as LINEs do in fact encode protein products which enable them to be reverse- transcribed and integrated back into the genome. SINEs are believed to have co-opted the proteins coded by LINEs which are contained in 2 reading frames. 72:
and remain highly conserved, suggesting positive pressure to preserve structure and function of SINEs. While SINEs are present in many species of vertebrates and invertebrates, SINEs are often lineage specific, making them useful markers of
148:, a sequence transcribed by RNA Polymerase III which codes for the RNA element of SRP, an abundant ribonucleoprotein. The body of SINEs possess an unknown origin but often share much homology with a corresponding 201:
primarily by affecting the accessibility of genes to transcriptional machinery. The chromosome has a very complex and hierarchical system of organizing the genome. This system of organization, which includes
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re-organization and the regulation of genomic architecture. The different lineages, mutations, and activities among eukaryotes make short-interspersed nuclear elements a useful tool in phylogenetic analysis.
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rearrangement and structure. The paper examined the global distribution of SINEs in mouse and human chromosomes and determined that this distribution was very similar to genomic distributions of genes and
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SINEs are characterized by their different modules, which are essentially a sectioning of their sequence. SINEs can, but do not necessarily have to possess a head, a body, and a tail. The head is at the
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The role of short-interspersed nuclear elements in gene regulation within cells has been supported by multiple studies. One such study examined the correlation between a certain family of SINEs with
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Scarpato M, Angelini C, Cocca E, Pallotta MM, Morescalchi MA, Capriglione T (September 2015). "Short interspersed DNA elements and miRNAs: a novel hidden gene regulation layer in zebrafish?".
332:. The transposition and recombination of SINEs and other active nuclear elements is thought to be one of the major contributions of genetic diversity between lineages during speciation. 432:(RISC). Among these proteins are proteins from the Argonaute family which are critical to the complex's ability to interact with and repress the translation of the target mRNA. 981:
Shi Y, Seto E, Chang LS, Shenk T (October 1991). "Transcriptional repression by YY1, a human GLI-Krüppel-related protein, and relief of repression by adenovirus E1A protein".
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Rinehart TA, Grahn RA, Wichman HA (2005). "SINE extinction preceded LINE extinction in sigmodontine rodents: implications for retrotranspositional dynamics and mechanisms".
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or changes to the regulatory region of the gene. Insertion of a SINE into the coding sequence of a gene can have deleterious effects and unregulated transposition can cause
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Seibt KM, Schmidt T, Heitkam T (February 2020). "The conserved 3' Angio-domain defines a superfamily of short interspersed nuclear elements (SINEs) in higher plants".
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There are >50 human diseases associated with SINEs. When inserted near or within the exon, SINEs can cause improper splicing, become coding regions, or change the
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Wicker T, Sabot F, Hua-Van A, Bennetzen JL, Capy P, Chalhoub B, et al. (December 2007). "A unified classification system for eukaryotic transposable elements".
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and during early development; as a result SINEs move around the genome most during these periods. SINE transcription is down-regulated by transcription factors in
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The activity of SINEs however has genetic vestiges which do not seem to play a significant role, positive or negative, and manifest themselves in the genome as
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factors and machinery can be recruited to increase or repress the transcription of local genes. The particular example of SINEs potentially recruiting the YY1
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Böhne A, Brunet F, Galiana-Arnoux D, Schultheis C, Volff JN (2008). "Transposable elements as drivers of genomic and biological diversity in vertebrates".
2342:"High cortisol in 5-year-old children causes loss of DNA methylation in SINE retrotransposons: a possible role for ZNF263 in stress-related diseases" 296:
after early development, though stress can cause up-regulation of normally silent SINEs. SINEs can be transferred between individuals or species via
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and inserted back into an alternate region in the genome. For this reason, short interspersed nuclear elements are believed to have co-evolved with
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Lau NC, Lim LP, Weinstein EG, Bartel DP (October 2001). "An abundant class of tiny RNAs with probable regulatory roles in Caenorhabditis elegans".
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Sun FJ, Fleurdépine S, Bousquet-Antonelli C, Caetano-Anollés G, Deragon JM (January 2007). "Common evolutionary trends for SINE RNA structures".
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Fawcett JA, Kawahara T, Watanabe H, Yasui Y (June 2006). "A SINE family widely distributed in the plant kingdom and its evolutionary history".
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Nevertheless, some SINE families such as the Au-SINEs and the Angio-SINEs are unusually widespread across many often unrelated plant species.
247:, which is tightly packed and generally not accessible to transcriptional machinery; SINEs seem to play an evolutionary role in this process. 285:
reverse-transcribed into DNA and integrated into the genome based on the sequence-motifs recognized by the protein's endonuclease domain.
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Lee Y, Ahn C, Han J, Choi H, Kim J, Yim J, et al. (September 2003). "The nuclear RNase III Drosha initiates microRNA processing".
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Kriegs JO, Churakov G, Jurka J, Brosius J, Schmitz J (April 2007). "Evolutionary history of 7SL RNA-derived SINEs in Supraprimates".
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based on differences in SINEs between species. SINEs are also implicated in certain types of genetic disease in humans and other
2396:"Aberrant methylation and associated transcriptional mobilization of Alu elements contributes to genomic instability in hypoxia" 2624: 161:
short-interspersed nuclear elements which also possess a body or are a combination of two or more SINEs are complex SINEs.
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groups, and a variety of proteins and RNAs allows different domains within a chromosome to be accessible to polymerases,
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The RNA coded by the short-interspersed nuclear element does not code for any protein product but is nonetheless
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Okada N, Hamada M, Ogiwara I, Ohshima K (December 1997). "SINEs and LINEs share common 3' sequences: a review".
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In fact Usmanova et al. 2008 suggested that short-interspersed nuclear elements can serve as direct signals in
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LINE-1 (L1) is transcribed and retrotransposed most frequently in the
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Insertion of a SINE upstream of a coding region may result in
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re-organization; this is often to facilitate the formation of
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Short-interspersed nuclear elements are believed to have
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Short-interspersed nuclear elements are transcribed by
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SINEs compose about 13% of the 2625: 2580:Current Opinion in Genetics & Development 2495: 2493: 1367: 1365: 1363: 611:Current Opinion in Genetics & Development 600: 598: 596: 68:The internal regions of SINEs originate from 8: 1621: 1619: 2389: 2387: 1724:Kramerov DA, Vassetzky NS (December 2011). 1318:Beauregard A, Curcio MJ, Belfort M (2008). 77:between species. Copy number variation and 2800: 2791: 2674: 2632: 2618: 2610: 2400:Journal of Cellular and Molecular Medicine 1270:Journal of Biomedicine & Biotechnology 1024:Yao YL, Yang WM, Seto E (September 2001). 560: 558: 556: 478:Vassetzky NS, Kramerov DA (January 2013). 197:Changes in chromosome structure influence 2473: 2463: 2419: 2367: 2357: 2316: 2306: 2265: 2224: 2132: 2032: 1983: 1749: 1700: 1651: 1599: 1488: 1343: 1291: 1281: 1240: 1230: 1189: 1148: 1049: 921: 911: 784: 630: 544: 503: 767:Deininger PL, Batzer MA (October 2002). 546:10.1146/annurev-cancerbio-030419-033525 461: 1463:Gogvadze E, Buzdin A (December 2009). 1336:10.1146/annurev.genet.42.110807.091549 1082:Nature Reviews. Molecular Cell Biology 1626:Wang W, Kirkness EF (December 2005). 1574:Cordaux R, Batzer MA (October 2009). 7: 1469:Cellular and Molecular Life Sciences 605:Hancks DC, Kazazian HH (June 2012). 2514:10.1146/annurev.ge.19.120185.001345 2295:The Journal of Biological Chemistry 1264:Mätlik K, Redik K, Speek M (2006). 29:Short interspersed nuclear elements 2701:Short tandem repeat/Microsatellite 2250:"Why do miRNAs live in the miRNP?" 2248:Schwarz DS, Zamore PD (May 2002). 14: 2289:Pratt AJ, MacRae IJ (July 2009). 2007:Faller M, Guo F (November 2008). 278:long interspersed nuclear element 2412:10.1111/j.1582-4934.2009.00792.x 1042:10.1128/mcb.21.17.5979-5991.2001 114:SINEs are classified as non-LTR 1533:Cytogenetic and Genome Research 533:Annual Review of Cancer Biology 46:in length. They are a class of 20:Genetic structure of human and 2705:Trinucleotide repeat disorders 1030:Molecular and Cellular Biology 1: 2692:Variable number tandem repeat 2226:10.1016/s0092-8674(04)00045-5 746:10.1016/s0378-1119(97)00409-5 430:RNA-induced silencing complex 320:Effects of SINE transposition 173:which is known to transcribe 101:. This regulation extends to 2025:10.1016/j.bbagrm.2008.08.005 1681:Genome Biology and Evolution 1434:10.1016/0092-8674(82)90194-5 995:10.1016/0092-8674(91)90189-6 913:10.1371/journal.pone.0001182 181:, two types of RNA vital to 120:long terminal repeats (LTRs) 118:because they do not contain 110:Classification and structure 140:For example, the 5' of the 40:transposable elements (TEs) 3157: 2207:Bartel DP (January 2004). 1191:10.1016/j.stem.2016.01.024 268:Propagation and regulation 193:Effects on gene expression 42:that are about 100 to 700 2659: 2592:10.1016/j.gde.2004.08.008 2502:Annual Review of Genetics 2359:10.1186/s13148-015-0123-z 1878:10.1007/s10577-015-9484-6 1787:10.1007/s11103-006-0026-7 1481:10.1007/s00018-009-0107-2 1386:10.1007/s10577-007-1202-6 1324:Annual Review of Genetics 769:"Mammalian retroelements" 711:10.1016/j.tig.2007.02.002 623:10.1016/j.gde.2012.02.006 579:10.1016/j.tig.2006.11.005 490:(Database issue): D83-9. 3126:Repetitive DNA sequences 1976:10.1038/sj.emboj.7600385 1580:Nature Reviews. Genetics 1232:10.1186/gb-2013-14-3-r22 810:Nature Reviews. Genetics 656:Nature Reviews. Genetics 3131:Mobile genetic elements 2254:Genes & Development 2078:10.1126/science.1065062 1775:Plant Molecular Biology 1129:Genes & Development 3100:Protein tandem repeats 3028:Tandemly arrayed genes 2308:10.1074/jbc.R900012200 1283:10.1155/JBB/2006/71753 853:Developmental Dynamics 484:Nucleic Acids Research 35:) are non-autonomous, 25: 444:SINEs and pseudogenes 216:transcription factors 19: 3073:Pathogenicity island 2465:10.3390/genes4020226 2346:Clinical Epigenetics 97:and the creation of 2178:10.1038/nature01957 2170:2003Natur.425..415L 2125:10.1261/rna.7135204 2070:2001Sci...294..858L 1929:10.1038/nature02871 1921:2004Natur.431..350A 1866:Chromosome Research 1742:10.1038/hdy.2011.43 1374:Chromosome Research 1141:10.1101/gad.1416106 904:2007PLoSO...2.1182R 496:10.1093/nar/gks1263 395:, and many others. 298:horizontal transfer 274:reverse-transcribed 264:eukaryotic genome. 75:divergent evolution 3023:Gene amplification 2267:10.1101/gad.992502 1693:10.1093/gbe/evv005 1644:10.1101/gr.3765505 865:10.1002/dvdy.21094 699:Trends in Genetics 567:Trends in Genetics 282:Open reading frame 171:RNA polymerase III 130:Internal structure 124:RNA polymerase III 26: 3121:Molecular biology 3108: 3107: 3009: 3008: 2877: 2876: 2781: 2780: 2670:Repeated sequence 2645:repeated sequence 2301:(27): 17897–901. 1830:10.1111/tpj.14567 1818:The Plant Journal 1545:10.1159/000084974 786:10.1101/gr.282402 393:neurofibromatosis 3148: 3085:Low copy repeats 3078:Symbiosis island 3015:Gene duplication 2801: 2792: 2675: 2653:gene duplication 2634: 2627: 2620: 2611: 2604: 2603: 2575: 2569: 2568: 2532: 2526: 2525: 2497: 2488: 2487: 2477: 2467: 2443: 2434: 2433: 2423: 2391: 2382: 2381: 2371: 2361: 2337: 2331: 2330: 2320: 2310: 2286: 2280: 2279: 2269: 2245: 2239: 2238: 2228: 2204: 2198: 2197: 2153: 2147: 2146: 2136: 2104: 2098: 2097: 2064:(5543): 858–62. 2053: 2047: 2046: 2036: 2004: 1998: 1997: 1987: 1964:The EMBO Journal 1955: 1949: 1948: 1904: 1898: 1897: 1861: 1842: 1841: 1813: 1807: 1806: 1770: 1764: 1763: 1753: 1721: 1715: 1714: 1704: 1672: 1666: 1665: 1655: 1638:(12): 1798–808. 1623: 1614: 1613: 1603: 1571: 1565: 1564: 1528: 1511: 1510: 1492: 1460: 1454: 1453: 1417: 1406: 1405: 1369: 1358: 1357: 1347: 1315: 1306: 1305: 1295: 1285: 1261: 1255: 1254: 1244: 1234: 1210: 1204: 1203: 1193: 1169: 1163: 1162: 1152: 1120: 1114: 1113: 1077: 1064: 1063: 1053: 1021: 1015: 1014: 978: 972: 971: 951: 936: 935: 925: 915: 883: 877: 876: 848: 842: 841: 805: 799: 798: 788: 764: 758: 757: 729: 723: 722: 694: 688: 687: 651: 645: 644: 634: 602: 591: 590: 562: 551: 550: 548: 524: 518: 517: 507: 475: 187:mRNA translation 144:is derived from 116:retrotransposons 57:, often through 48:retrotransposons 24:LINE1 and SINEs. 3156: 3155: 3151: 3150: 3149: 3147: 3146: 3145: 3141:Eukaryote genes 3111: 3110: 3109: 3104: 3056: 3005: 2873: 2845: 2822: 2796:Retrotransposon 2777: 2768:Inverted repeat 2756: 2741:DNA transposon 2737:Retrotransposon 2732:Gene conversion 2723: 2716: 2713: 2664: 2655: 2638: 2608: 2607: 2577: 2576: 2572: 2537:Nature Genetics 2534: 2533: 2529: 2499: 2498: 2491: 2445: 2444: 2437: 2406:(11): 2646–54. 2393: 2392: 2385: 2339: 2338: 2334: 2288: 2287: 2283: 2247: 2246: 2242: 2206: 2205: 2201: 2164:(6956): 415–9. 2155: 2154: 2150: 2119:(12): 1957–66. 2106: 2105: 2101: 2055: 2054: 2050: 2006: 2005: 2001: 1970:(20): 4051–60. 1957: 1956: 1952: 1915:(7006): 350–5. 1906: 1905: 1901: 1863: 1862: 1845: 1815: 1814: 1810: 1772: 1771: 1767: 1723: 1722: 1718: 1674: 1673: 1669: 1632:Genome Research 1625: 1624: 1617: 1592:10.1038/nrg2640 1586:(10): 691–703. 1573: 1572: 1568: 1539:(1–4): 416–25. 1530: 1529: 1514: 1475:(23): 3727–42. 1462: 1461: 1457: 1419: 1418: 1409: 1371: 1370: 1361: 1317: 1316: 1309: 1263: 1262: 1258: 1212: 1211: 1207: 1171: 1170: 1166: 1135:(11): 1470–84. 1122: 1121: 1117: 1094:10.1038/nrm1946 1079: 1078: 1067: 1036:(17): 5979–91. 1023: 1022: 1018: 980: 979: 975: 953: 952: 939: 885: 884: 880: 850: 849: 845: 822:10.1038/nrg3001 807: 806: 802: 779:(10): 1455–65. 773:Genome Research 766: 765: 761: 740:(1–2): 229–43. 731: 730: 726: 696: 695: 691: 668:10.1038/nrg2165 653: 652: 648: 604: 603: 594: 564: 563: 554: 526: 525: 521: 477: 476: 463: 458: 446: 401: 389:cystic fibrosis 361: 338: 330:genetic disease 322: 270: 245:heterochromatin 237:heterochromatin 199:gene expression 195: 167: 132: 112: 95:gene expression 12: 11: 5: 3154: 3152: 3144: 3143: 3138: 3136:Non-coding DNA 3133: 3128: 3123: 3113: 3112: 3106: 3105: 3103: 3102: 3097: 3092: 3087: 3082: 3081: 3080: 3075: 3068:Genomic island 3064: 3062: 3058: 3057: 3055: 3054: 3049: 3048: 3047: 3037: 3036: 3035: 3025: 3019: 3017: 3011: 3010: 3007: 3006: 3004: 3003: 2998: 2993: 2988: 2983: 2978: 2973: 2968: 2963: 2958: 2953: 2948: 2943: 2938: 2933: 2928: 2923: 2918: 2913: 2908: 2903: 2898: 2893: 2887: 2885: 2883:DNA transposon 2879: 2878: 2875: 2874: 2872: 2871: 2866: 2861: 2855: 2853: 2847: 2846: 2844: 2843: 2838: 2832: 2830: 2824: 2823: 2821: 2820: 2815: 2809: 2807: 2798: 2789: 2783: 2782: 2779: 2778: 2776: 2775: 2770: 2764: 2762: 2758: 2757: 2755: 2754: 2753: 2752: 2747: 2739: 2734: 2728: 2726: 2718: 2717: 2715: 2714: 2711:Macrosatellite 2708: 2698: 2689: 2683: 2681: 2679:Tandem repeats 2672: 2666: 2665: 2660: 2657: 2656: 2639: 2637: 2636: 2629: 2622: 2614: 2606: 2605: 2570: 2549:10.1038/ng1223 2527: 2489: 2435: 2383: 2332: 2281: 2260:(9): 1025–31. 2240: 2199: 2148: 2099: 2048: 1999: 1950: 1899: 1843: 1824:(3): 681–699. 1808: 1765: 1716: 1667: 1615: 1566: 1512: 1455: 1407: 1359: 1307: 1256: 1219:Genome Biology 1205: 1178:Cell Stem Cell 1164: 1115: 1065: 1016: 973: 958:(in Russian). 937: 878: 859:(4): 1144–56. 843: 800: 759: 724: 689: 662:(12): 973–82. 646: 617:(3): 191–203. 592: 552: 519: 460: 459: 457: 454: 445: 442: 400: 397: 385:Dent's disease 360: 357: 337: 334: 326:exon shuffling 321: 318: 269: 266: 194: 191: 166: 163: 131: 128: 111: 108: 13: 10: 9: 6: 4: 3: 2: 3153: 3142: 3139: 3137: 3134: 3132: 3129: 3127: 3124: 3122: 3119: 3118: 3116: 3101: 3098: 3096: 3093: 3091: 3088: 3086: 3083: 3079: 3076: 3074: 3071: 3070: 3069: 3066: 3065: 3063: 3059: 3053: 3050: 3046: 3043: 3042: 3041: 3038: 3034: 3033:Ribosomal DNA 3031: 3030: 3029: 3026: 3024: 3021: 3020: 3018: 3016: 3012: 3002: 2999: 2997: 2994: 2992: 2989: 2987: 2984: 2982: 2979: 2977: 2974: 2972: 2969: 2967: 2964: 2962: 2959: 2957: 2954: 2952: 2949: 2947: 2944: 2942: 2939: 2937: 2934: 2932: 2929: 2927: 2924: 2922: 2919: 2917: 2914: 2912: 2909: 2907: 2904: 2902: 2899: 2897: 2894: 2892: 2889: 2888: 2886: 2884: 2880: 2870: 2867: 2865: 2862: 2860: 2857: 2856: 2854: 2852: 2848: 2842: 2839: 2837: 2834: 2833: 2831: 2829: 2825: 2819: 2816: 2814: 2811: 2810: 2808: 2806: 2802: 2799: 2797: 2793: 2790: 2788: 2784: 2774: 2773:Direct repeat 2771: 2769: 2766: 2765: 2763: 2759: 2751: 2748: 2746: 2743: 2742: 2740: 2738: 2735: 2733: 2730: 2729: 2727: 2725: 2719: 2712: 2709: 2706: 2702: 2699: 2697: 2696:Minisatellite 2693: 2690: 2688: 2687:Satellite DNA 2685: 2684: 2682: 2680: 2676: 2673: 2671: 2667: 2663: 2658: 2654: 2650: 2646: 2642: 2635: 2630: 2628: 2623: 2621: 2616: 2615: 2612: 2601: 2597: 2593: 2589: 2585: 2581: 2574: 2571: 2566: 2562: 2558: 2554: 2550: 2546: 2542: 2538: 2531: 2528: 2523: 2519: 2515: 2511: 2507: 2503: 2496: 2494: 2490: 2485: 2481: 2476: 2471: 2466: 2461: 2458:(2): 226–43. 2457: 2453: 2449: 2442: 2440: 2436: 2431: 2427: 2422: 2417: 2413: 2409: 2405: 2401: 2397: 2390: 2388: 2384: 2379: 2375: 2370: 2365: 2360: 2355: 2351: 2347: 2343: 2336: 2333: 2328: 2324: 2319: 2314: 2309: 2304: 2300: 2296: 2292: 2285: 2282: 2277: 2273: 2268: 2263: 2259: 2255: 2251: 2244: 2241: 2236: 2232: 2227: 2222: 2219:(2): 281–97. 2218: 2214: 2210: 2203: 2200: 2195: 2191: 2187: 2183: 2179: 2175: 2171: 2167: 2163: 2159: 2152: 2149: 2144: 2140: 2135: 2130: 2126: 2122: 2118: 2114: 2110: 2103: 2100: 2095: 2091: 2087: 2083: 2079: 2075: 2071: 2067: 2063: 2059: 2052: 2049: 2044: 2040: 2035: 2030: 2026: 2022: 2019:(11): 663–7. 2018: 2014: 2010: 2003: 2000: 1995: 1991: 1986: 1981: 1977: 1973: 1969: 1965: 1961: 1954: 1951: 1946: 1942: 1938: 1934: 1930: 1926: 1922: 1918: 1914: 1910: 1903: 1900: 1895: 1891: 1887: 1883: 1879: 1875: 1872:(3): 533–44. 1871: 1867: 1860: 1858: 1856: 1854: 1852: 1850: 1848: 1844: 1839: 1835: 1831: 1827: 1823: 1819: 1812: 1809: 1804: 1800: 1796: 1792: 1788: 1784: 1781:(3): 505–14. 1780: 1776: 1769: 1766: 1761: 1757: 1752: 1747: 1743: 1739: 1736:(6): 487–95. 1735: 1731: 1727: 1720: 1717: 1712: 1708: 1703: 1698: 1694: 1690: 1687:(2): 567–80. 1686: 1682: 1678: 1671: 1668: 1663: 1659: 1654: 1649: 1645: 1641: 1637: 1633: 1629: 1622: 1620: 1616: 1611: 1607: 1602: 1597: 1593: 1589: 1585: 1581: 1577: 1570: 1567: 1562: 1558: 1554: 1550: 1546: 1542: 1538: 1534: 1527: 1525: 1523: 1521: 1519: 1517: 1513: 1508: 1504: 1500: 1496: 1491: 1486: 1482: 1478: 1474: 1470: 1466: 1459: 1456: 1451: 1447: 1443: 1439: 1435: 1431: 1427: 1423: 1416: 1414: 1412: 1408: 1403: 1399: 1395: 1391: 1387: 1383: 1380:(1): 203–15. 1379: 1375: 1368: 1366: 1364: 1360: 1355: 1351: 1346: 1341: 1337: 1333: 1329: 1325: 1321: 1314: 1312: 1308: 1303: 1299: 1294: 1289: 1284: 1279: 1275: 1271: 1267: 1260: 1257: 1252: 1248: 1243: 1238: 1233: 1228: 1224: 1220: 1216: 1209: 1206: 1201: 1197: 1192: 1187: 1184:(5): 637–52. 1183: 1179: 1175: 1168: 1165: 1160: 1156: 1151: 1146: 1142: 1138: 1134: 1130: 1126: 1119: 1116: 1111: 1107: 1103: 1099: 1095: 1091: 1087: 1083: 1076: 1074: 1072: 1070: 1066: 1061: 1057: 1052: 1047: 1043: 1039: 1035: 1031: 1027: 1020: 1017: 1012: 1008: 1004: 1000: 996: 992: 989:(2): 377–88. 988: 984: 977: 974: 969: 965: 962:(3): 256–60. 961: 957: 950: 948: 946: 944: 942: 938: 933: 929: 924: 919: 914: 909: 905: 901: 898:(11): e1182. 897: 893: 889: 882: 879: 874: 870: 866: 862: 858: 854: 847: 844: 839: 835: 831: 827: 823: 819: 816:(7): 459–63. 815: 811: 804: 801: 796: 792: 787: 782: 778: 774: 770: 763: 760: 755: 751: 747: 743: 739: 735: 728: 725: 720: 716: 712: 708: 705:(4): 158–61. 704: 700: 693: 690: 685: 681: 677: 673: 669: 665: 661: 657: 650: 647: 642: 638: 633: 628: 624: 620: 616: 612: 608: 601: 599: 597: 593: 588: 584: 580: 576: 572: 568: 561: 559: 557: 553: 547: 542: 538: 534: 530: 523: 520: 515: 511: 506: 501: 497: 493: 489: 485: 481: 474: 472: 470: 468: 466: 462: 455: 453: 451: 443: 441: 439: 433: 431: 425: 421: 417: 414: 410: 406: 398: 396: 394: 390: 386: 382: 378: 374: 370: 369:breast cancer 366: 365:reading frame 358: 356: 352: 350: 346: 343: 335: 333: 331: 327: 319: 317: 313: 309: 305: 303: 299: 295: 294:somatic cells 291: 286: 283: 279: 275: 267: 265: 261: 258: 252: 248: 246: 242: 238: 233: 229: 224: 219: 217: 213: 209: 205: 200: 192: 190: 188: 185:assembly and 184: 180: 176: 175:ribosomal RNA 172: 165:Transcription 164: 162: 159: 155: 154:endonucleases 151: 147: 143: 138: 129: 127: 125: 121: 117: 109: 107: 104: 100: 96: 90: 88: 84: 80: 76: 71: 66: 64: 60: 56: 53: 49: 45: 41: 38: 34: 30: 23: 18: 3045:Gene cluster 2813:Alu sequence 2804: 2722:Interspersed 2586:(6): 603–8. 2583: 2579: 2573: 2540: 2536: 2530: 2505: 2501: 2455: 2451: 2403: 2399: 2349: 2345: 2335: 2298: 2294: 2284: 2257: 2253: 2243: 2216: 2212: 2202: 2161: 2157: 2151: 2116: 2112: 2102: 2061: 2057: 2051: 2016: 2012: 2002: 1967: 1963: 1953: 1912: 1908: 1902: 1869: 1865: 1821: 1817: 1811: 1778: 1774: 1768: 1733: 1729: 1719: 1684: 1680: 1670: 1635: 1631: 1583: 1579: 1569: 1536: 1532: 1472: 1468: 1458: 1428:(3): 433–4. 1425: 1421: 1377: 1373: 1327: 1323: 1276:(1): 71753. 1273: 1269: 1259: 1222: 1218: 1208: 1181: 1177: 1167: 1132: 1128: 1118: 1088:(8): 612–6. 1085: 1081: 1033: 1029: 1019: 986: 982: 976: 959: 955: 895: 891: 881: 856: 852: 846: 813: 809: 803: 776: 772: 762: 737: 733: 727: 702: 698: 692: 659: 655: 649: 614: 610: 573:(1): 26–33. 570: 566: 536: 532: 522: 487: 483: 447: 434: 426: 422: 418: 412: 402: 373:colon cancer 362: 353: 349:Alu elements 347: 339: 336:Common SINEs 323: 314: 310: 306: 302:viral vector 287: 271: 262: 253: 249: 220: 196: 168: 133: 113: 91: 67: 32: 28: 27: 3040:Gene family 2951:Tc1/mariner 2906:EnSpm/CACTA 2543:(1): 41–8. 1330:: 587–617. 956:Tsitologiia 539:: 159–176. 450:pseudogenes 413:Danio rerio 241:euchromatin 83:phylogenies 3115:Categories 3052:Pseudogene 2869:retroposon 2787:Transposon 2649:transposon 2508:: 253–72. 1225:(3): R22. 456:References 381:hemophilia 300:through a 228:CpG motifs 87:eukaryotes 52:eukaryotic 44:base pairs 37:non-coding 2971:P element 2921:Harbinger 2662:Repeatome 2352:(1): 91. 1945:205210153 409:zebrafish 405:microRNAs 399:microRNAs 342:parasitic 290:germ-line 223:chromatin 183:ribosomal 103:chromatin 99:RNA genes 79:mutations 63:mammalian 3095:Telomere 3061:See also 3001:Zisupton 2981:Polinton 2976:PiggyBac 2931:Helitron 2750:Helitron 2745:Polinton 2641:Genetics 2600:15531153 2565:32151696 2557:12897783 2484:24705161 2430:19508390 2378:26339299 2327:19342379 2276:12000786 2235:14744438 2186:14508493 2143:15525708 2094:43262684 2086:11679671 2043:18778799 1994:15372072 1937:15372042 1894:16759020 1886:26363800 1838:31610059 1795:16830182 1760:21673742 1730:Heredity 1711:25577199 1662:16339378 1610:19763152 1561:36518754 1553:16093694 1507:23872541 1499:19649766 1490:11115525 1450:22129236 1402:10510149 1394:18293113 1354:18680436 1302:16877819 1251:23497673 1200:26996597 1159:16705037 1110:22274894 1102:16723972 1060:11486036 1011:19399858 968:18664128 932:18000552 892:PLOS ONE 873:17304537 838:21123216 830:21540878 795:12368238 719:17307271 684:32132898 676:17984973 641:22406018 587:17126948 514:23203982 377:leukemia 359:Diseases 257:polycomb 210:groups, 204:histones 142:Alu sine 65:genome. 2991:Transib 2966:Novosib 2946:Kolobok 2916:Ginger2 2911:Ginger1 2896:Crypton 2522:3909943 2475:3899967 2421:4373486 2369:4559301 2318:2709356 2194:4421030 2166:Bibcode 2134:1370684 2066:Bibcode 2058:Science 2034:2633599 1917:Bibcode 1803:7840648 1751:3242629 1702:4350176 1653:1356118 1601:2884099 1442:6280868 1345:2665727 1293:1559930 1242:3663115 1150:1475760 1003:1655281 923:2063516 900:Bibcode 754:9461397 632:3376660 505:3531059 158:3′ tail 146:7SL RNA 55:genomes 3090:CRISPR 2956:Merlin 2941:ISL2EU 2891:Academ 2724:repeat 2598:  2563:  2555:  2520:  2482:  2472:  2428:  2418:  2376:  2366:  2325:  2315:  2274:  2233:  2192:  2184:  2158:Nature 2141:  2131:  2092:  2084:  2041:  2031:  1992:  1985:524334 1982:  1943:  1935:  1909:Nature 1892:  1884:  1836:  1801:  1793:  1758:  1748:  1709:  1699:  1660:  1650:  1608:  1598:  1559:  1551:  1505:  1497:  1487:  1448:  1440:  1400:  1392:  1352:  1342:  1300:  1290:  1249:  1239:  1198:  1157:  1147:  1108:  1100:  1058:  1048:  1009:  1001:  966:  930:  920:  871:  836:  828:  793:  752:  717:  682:  674:  639:  629:  585:  512:  502:  212:acetyl 208:methyl 137:5' end 22:murine 2996:Zator 2936:IS3EU 2841:LINE2 2836:LINE1 2828:LINEs 2805:SINEs 2761:Other 2561:S2CID 2452:Genes 2190:S2CID 2090:S2CID 1941:S2CID 1890:S2CID 1799:S2CID 1557:S2CID 1503:S2CID 1446:S2CID 1398:S2CID 1106:S2CID 1051:87316 1007:S2CID 834:S2CID 680:S2CID 438:BRCA1 33:SINEs 2986:Sola 2961:MuDR 2901:Dada 2864:MER4 2859:HERV 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Index


murine
non-coding
transposable elements (TEs)
base pairs
retrotransposons
eukaryotic
genomes
RNA
mammalian
tRNA
divergent evolution
mutations
phylogenies
eukaryotes
gene expression
RNA genes
chromatin
retrotransposons
long terminal repeats (LTRs)
RNA polymerase III
5' end
Alu sine
7SL RNA
LINE
endonucleases
3′ tail
RNA polymerase III
ribosomal RNA
tRNA

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