A substantial number of “retrogenes” that are derived from the mRNA of various intron-containing genes have been reported. A class of mammalian retroposons, long interspersed element-1 (LINE1, L1), has been shown to be involved in the reverse transcription of retrogenes (or processed pseudogenes) and non-autonomous short interspersed elements (SINEs). The -end sequences of various SINEs originated from a corresponding LINE. As the -untranslated regions of several LINEs are essential for retroposition, these LINEs presumably require “stringent” recognition of the -end sequence of the RNA template. However, the -ends of mammalian L1s do not exhibit any similarity to SINEs, except for the presence of -poly(A) repeats. Since the -poly(A) repeats of L1 and Alu SINE are critical for their retroposition, L1 probably recognizes the poly(A) repeats, thereby mobilizing not only Alu SINE but also cytosolic mRNA. Many flowering plants only harbor L1-clade LINEs and a significant number of SINEs with poly(A) repeats, but no homology to the LINEs. Moreover, processed pseudogenes have also been found in flowering plants. I propose that the ancestral L1-clade LINE in the common ancestor of green plants may have recognized a specific RNA template, with stringent recognition then becoming relaxed during the course of plant evolution. 1. RNA-Mediated Gene Duplication and Retroposons 1.1. Retrogenes and Processed Pseudogenes Gene duplication is a fundamental process of gene evolution [1]. There are two types of gene duplication: direct duplication of genomic DNA and retropositional events [2–4]. Processed pseudogenes (PPs) are reverse-transcribed intronless cDNA copies of mRNA that have been reinserted into the genome (Figure 1) [5, 6]; they are especially abundant in mammalian genomes [7, 8]. PPs are not usually transcribed because they lack an external promoter; therefore, they have long been viewed as evolutionary dead ends with little biological relevance. However, recent studies have unveiled a substantial number of “processed genes” or “retrogenes” with novel functions that are derived from the mRNA of various intron-containing genes [9–12]. Molecular biological studies showed that a class of mammalian retroposons, long interspersed element-1 (LINE1, L1), has been involved in the reverse transcription of nonautonomous retroposons, such as PPs (retrogenes) and short interspersed elements (SINEs) [13]. Figure 1: Schematic representation of the formation of a processed pseudogene. 1.2. Retroposons Eukaryotic genomes generally contain an extraordinary number of
References
[1]
S. Ohno, Evolution by Gene Duplication, Springer, New York, NY, USA, 1970.
[2]
M. Long, E. Betrán, K. Thornton, and W. Wang, “The origin of new genes: glimpses from the young and old,” Nature Reviews Genetics, vol. 4, no. 11, pp. 865–875, 2003.
[3]
D. V. Babushok, E. M. Ostertag, and H. H. Kazazian Jr., “Current topics in genome evolution: molecular mechanisms of new gene formation,” Cellular and Molecular Life Sciences, vol. 64, no. 5, pp. 542–554, 2007.
[4]
C. Charon, Q. Bruggeman, V. Thareau, and Y. Henry, “Gene duplication within the green lineage: the case of TEL genes,” Journal of Experimental Botany, vol. 63, no. 14, pp. 5061–5077, 2012.
[5]
E. F. Vanin, “Processed pseudogenes: characteristics and evolution,” Annual Review of Genetics, vol. 19, pp. 253–272, 1985.
[6]
A. M. Weiner, P. L. Deininger, and A. Efstratiadis, “Nonviral retroposons: genes, pseudogenes, and transposable elements generated by the reverse flow of genetic information,” Annual Review of Biochemistry, vol. 55, pp. 631–661, 1986.
[7]
Z. Yu, D. Morais, M. Ivanga, and P. M. Harrison, “Analysis of the role of retrotransposition in gene evolution in vertebrates,” BMC Bioinformatics, vol. 8, article 308, 2007.
[8]
Y.-J. Liu, D. Zheng, S. Balasubramanian et al., “Comprehensive analysis of the pseudogenes of glycolytic enzymes in vertebrates: the anomalously high number of GAPDH pseudogenes highlights a recent burst of retrotrans-positional activity,” BMC Genomics, vol. 10, article 480, 2009.
[9]
J. Brosius, “RNAs from all categories generate retrosequences that may be exapted as novel genes or regulatory elements,” Gene, vol. 238, no. 1, pp. 115–134, 1999.
[10]
H. Kaessmann, N. Vinckenbosch, and M. Long, “RNA-based gene duplication: mechanistic and evolutionary insights,” Nature Reviews Genetics, vol. 10, no. 1, pp. 19–31, 2009.
[11]
H. Sakai, H. Mizuno, Y. Kawahara et al., “Retrogenes in rice (Oryza sativa L. ssp. japonica) exhibit correlated expression with their source genes,” Genome Biology and Evolution, vol. 3, no. 1, pp. 1357–1368, 2011.
[12]
J. Ciomborowska, W. Rosikiewicz, D. Szklarczyk, W. Maka?owski, and I. Maka?owska, ““Orphan” retrogenes in the human genome,” Molecular Biology and Evolution, vol. 30, no. 2, pp. 384–396, 2013.
[13]
H. H. Kazazian Jr., “Mobile elements: drivers of genome evolution,” Science, vol. 303, no. 5664, pp. 1626–1632, 2004.
[14]
J. Brosius, “Retroposons—seeds of evolution,” Science, vol. 251, no. 4995, p. 753, 1991.
[15]
J. Jurka, V. V. Kapitonov, A. Pavlicek, P. Klonowski, O. Kohany, and J. Walichiewicz, “Repbase Update, a database of eukaryotic repetitive elements,” Cytogenetic and Genome Research, vol. 110, no. 1–4, pp. 462–467, 2005.
[16]
D. D. Luan, M. H. Korman, J. L. Jakubczak, and T. H. Eickbush, “Reverse transcription of R2Bm RNA is primed by a nick at the chromosomal target site: a mechanism for non-LTR retrotransposition,” Cell, vol. 72, no. 4, pp. 595–605, 1993.
[17]
D. D. Luan and T. H. Eickbush, “RNA template requirements for target DNA-primed reverse transcription by the R2 retrotransposable element,” Molecular and Cellular Biology, vol. 15, no. 7, pp. 3882–3891, 1995.
[18]
J. L. Goodier and H. H. Kazazian Jr., “Retrotransposons revisited: the restraint and rehabilitation of parasites,” Cell, vol. 135, no. 1, pp. 23–35, 2008.
[19]
G. J. Cost, Q. Feng, A. Jacquier, and J. D. Boeke, “Human L1 element target-primed reverse transcription in vitro,” The EMBO Journal, vol. 21, no. 21, pp. 5899–5910, 2002.
[20]
N. Okada, “SINEs: short interspersed repeated elements of the eukaryotic genome,” Trends in Ecology and Evolution, vol. 6, no. 11, pp. 358–361, 1991.
[21]
V. V. Kapitonov and J. Jurka, “A novel class of SINE elements derived from 5S rRNA,” Molecular Biology and Evolution, vol. 20, no. 5, pp. 694–702, 2003.
[22]
N. S. Vassetzky and D. A. Kramerov, “SINEBase: a database and tool for SINE analysis,” Nucleic Acids Research, vol. 41, pp. D83–D89, 2013.
[23]
K. Ohshima and N. Okada, “SINEs and LINEs: symbionts of eukaryotic genomes with a common tail,” Cytogenetic and Genome Research, vol. 110, no. 1–4, pp. 475–490, 2005.
[24]
J. Schmitz, A. Zemann, G. Churakov et al., “Retroposed SNOfall—a mammalian-wide comparison of platypus snoRNAs,” Genome Research, vol. 18, no. 6, pp. 1005–1010, 2008.
[25]
H. S. Malik, W. D. Burke, and T. H. Eickbush, “The age and evolution of non-LTR retrotransposable elements,” Molecular Biology and Evolution, vol. 16, no. 6, pp. 793–805, 1999.
[26]
V. V. Kapitonov, S. Tempel, and J. Jurka, “Simple and fast classification of non-LTR retrotransposons based on phylogeny of their RT domain protein sequences,” Gene, vol. 448, no. 2, pp. 207–213, 2009.
[27]
A. V. Furano, D. D. Duvernell, and S. Boissinot, “L1 (LINE-1) retrotransposon diversity differs dramatically between mammals and fish,” Trends in Genetics, vol. 20, no. 1, pp. 9–14, 2004.
[28]
D. Kordi?, N. Lov?in, and F. Guben?ek, “Phylogenomic analysis of the L1 retrotransposons in Deuterostomia,” Systematic Biology, vol. 55, no. 6, pp. 886–901, 2006.
[29]
K. Ichiyanagi, H. Nishihara, D. D. Duvernell, and N. Okada, “Acquisition of endonuclease specificity during evolution of L1 retrotransposon,” Molecular Biology and Evolution, vol. 24, no. 9, pp. 2009–2015, 2007.
[30]
K. K. Kojima and H. Fujiwara, “Cross-genome screening of novel sequence-specific Non-LTR retrotransposons: various multicopy RNA genes and microsatellites are selected as targets,” Molecular Biology and Evolution, vol. 21, no. 2, pp. 207–217, 2004.
[31]
K. Ohshima, M. Hamada, Y. Terai, and N. Okada, “The 3′ ends of tRNA-derived short interspersed repetitive elements are derived from the 3′ ends of long interspersed repetitive elements,” Molecular and Cellular Biology, vol. 16, no. 7, pp. 3756–3764, 1996.
[32]
N. Okada, M. Hamada, I. Ogiwara, and K. Ohshima, “SINEs and LINEs share common 3′ sequences: a review,” Gene, vol. 205, no. 1-2, pp. 229–243, 1997.
[33]
A. M. Weiner, “SINEs and LINEs: the art of biting the hand that feeds you,” Current Opinion in Cell Biology, vol. 14, no. 3, pp. 343–350, 2002.
[34]
K. Ohshima, “Parallel relaxation of stringent RNA recognition in plant and mammalian L1 retrotransposons,” Molecular Biology and Evolution, vol. 29, no. 11, pp. 3255–3259, 2012.
[35]
Y. Yoshioka, S. Matsumoto, S. Kojima, K. Ohshima, N. Okada, and Y. Machida, “Molecular characterization of a short interspersed repetitive element from tobacco that exhibits sequence homology to specific tRNAs,” Proceedings of the National Academy of Sciences of the United States of America, vol. 90, no. 14, pp. 6562–6566, 1993.
[36]
M. Kajikawa and N. Okada, “LINEs mobilize SINEs in the eel through a shared 3′ sequence,” Cell, vol. 111, no. 3, pp. 433–444, 2002.
[37]
H. Takahashi and H. Fujiwara, “Transplantation of target site specificity by swapping the endonuclease domains of two LINEs,” The EMBO Journal, vol. 21, no. 3, pp. 408–417, 2002.
[38]
M. Osanai, H. Takahashi, K. K. Kojima, M. Hamada, and H. Fujiwara, “Essential motifs in the 3′ untranslated region required for retrotransposition and the precise start of reverse transcription in non-long-terminal-repeat retrotransposon SART1,” Molecular and Cellular Biology, vol. 24, no. 18, pp. 7902–7913, 2004.
[39]
T. Anzai, M. Osanai, M. Hamada, and H. Fujiwara, “Functional roles of 3′-terminal structures of template RNA during in vivo retrotransposition of non-LTR retrotransposon, R1Bm,” Nucleic Acids Research, vol. 33, no. 6, pp. 1993–2002, 2005.
[40]
Q. Feng, J. V. Moran, H. H. Kazazian Jr., and J. D. Boeke, “Human L1 retrotransposon encodes a conserved endonuclease required for retrotransposition,” Cell, vol. 87, no. 5, pp. 905–916, 1996.
[41]
J. Jurka, “Sequence patterns indicate an enzymatic involvement in integration of mammalian retroposons,” Proceedings of the National Academy of Sciences of the United States of America, vol. 94, no. 5, pp. 1872–1877, 1997.
[42]
A. Pavlí?ek, J. Pa?es, D. Elleder, and J. Hejnar, “Processed pseudogenes of human endogenous retroviruses generated by LINEs: their integration, stability, and distribution,” Genome Research, vol. 12, no. 3, pp. 391–399, 2002.
[43]
C. Esnault, J. Maestre, and T. Heidmann, “Human LINE retrotransposons generate processed pseudogenes,” Nature Genetics, vol. 24, no. 4, pp. 363–367, 2000.
[44]
W. Wei, N. Gilbert, S. L. Ooi et al., “Human L1 retrotransposition: cis preference versus trans complementation,” Molecular and Cellular Biology, vol. 21, no. 4, pp. 1429–1439, 2001.
[45]
A. M. Roy-Engel, A.-H. Salem, O. O. Oyeniran et al., “Active Alu element “A-tails”: size does matter,” Genome Research, vol. 12, no. 9, pp. 1333–1344, 2002.
[46]
M. Dewannieux, C. Esnault, and T. Heidmann, “LINE-mediated retrotransposition of marked Alu sequences,” Nature Genetics, vol. 35, no. 1, pp. 41–48, 2003.
[47]
E. N. Kroutter, V. P. Belancio, B. J. Wagstaff, and A. M. Roy-Engel, “The RNA polymerase dictates ORF1 requirement and timing of LINE and SINE retrotransposition,” PLoS Genetics, vol. 5, no. 4, Article ID e1000458, 2009.
[48]
J. V. Moran, S. E. Holmes, T. P. Naas, R. J. DeBerardinis, J. D. Boeke, and H. H. Kazazian Jr., “High frequency retrotransposition in cultured mammalian cells,” Cell, vol. 87, no. 5, pp. 917–927, 1996.
[49]
J. V. Moran, R. J. DeBerardinis, and H. H. Kazazian Jr., “Exon shuffling by L1 retrotransposition,” Science, vol. 283, no. 5407, pp. 1530–1534, 1999.
[50]
K. Ohshima, M. Hattori, T. Yada, T. Gojobori, Y. Sakaki, and N. Okada, “Whole-genome screening indicates a possible burst of formation of processed pseudogenes and Alu repeats by particular L1 subfamilies in ancestral primates,” Genome Biology, vol. 4, no. 11, article R74, 2003.
[51]
J. D. Boeke, “LINEs and Alus—the polyA connection,” Nature genetics, vol. 16, no. 1, pp. 6–7, 1997.
[52]
J. Schmitz, G. Churakov, H. Zischler, and J. Brosius, “A novel class of mammalian-specific tailless retropseudogenes,” Genome Research, vol. 14, no. 10a, pp. 1911–1915, 2004.
[53]
Z. Zhang, P. M. Harrison, Y. Liu, and M. Gerstein, “Millions of years of evolution preserved: a comprehensive catalog of the processed pseudogenes in the human genome,” Genome Research, vol. 13, no. 12, pp. 2541–2558, 2003.
[54]
A. C. Marques, I. Dupanloup, N. Vinckenbosch, A. Reymond, and H. Kaessmann, “Emergence of young human genes after a burst of retroposition in primates,” PLoS Biology, vol. 3, no. 11, article e357, 2005.
[55]
H. Sakai, K. O. Koyanagi, T. Imanishi, T. Itoh, and T. Gojobori, “Frequent emergence and functional resurrection of processed pseudogenes in the human and mouse genomes,” Gene, vol. 389, no. 2, pp. 196–203, 2007.
[56]
B. J. Wagstaff, E. N. Kroutter, R. S. Derbes, V. P. Belancio, and A. M. Roy-Engel, “Molecular reconstruction of extinct LINE-1 elements and their interaction with nonautonomous elements,” Molecular Biology and Evolution, vol. 30, no. 1, pp. 88–99, 2013.
[57]
F. Burki and H. Kaessmann, “Birth and adaptive evolution of a hominoid gene that supports high neurotransmitter flux,” Nature Genetics, vol. 36, no. 10, pp. 1061–1063, 2004.
[58]
L. Rosso, A. C. Marques, M. Weier et al., “Birth and rapid subcellular adaptation of a hominoid-specific CDC14 protein,” PLoS Biology, vol. 6, no. 6, article e140, 2008.
[59]
D. V. Babushok, K. Ohshima, E. M. Ostertag et al., “A novel testis ubiquitin-binding protein gene arose by exon shuffling in hominoids,” Genome Research, vol. 17, no. 8, pp. 1129–1138, 2007.
[60]
Y. Zhang, S. Lu, S. Zhao, X. Zheng, M. Long, and L. Wei, “Positive selection for themale functionality of a co-retroposed gene in the hominoids,” BMC Evolutionary Biology, vol. 9, article 252, 2009.
[61]
K. Ohshima and K. Igarashi, “Inference for the initial stage of domain shuffling: tracing the evolutionary fate of the PIPSL retrogene in hominoids,” Molecular Biology and Evolution, vol. 27, no. 11, pp. 2522–2533, 2010.
[62]
P. M. Harrison, D. Zheng, Z. Zhang, N. Carriero, and M. Gerstein, “Transcribed processed pseudogenes in the human genome: an intermediate form of expressed retrosequence lacking protein-coding ability,” Nucleic Acids Research, vol. 33, no. 8, pp. 2374–2383, 2005.
[63]
N. Vinckenbosch, I. Dupanloup, and H. Kaessmann, “Evolutionary fate of retroposed gene copies in the human genome,” Proceedings of the National Academy of Sciences of the United States of America, vol. 103, no. 9, pp. 3220–3225, 2006.
[64]
R. Baertsch, M. Diekhans, W. J. Kent, D. Haussler, and J. Brosius, “Retrocopy contributions to the evolution of the human genome,” BMC Genomics, vol. 9, article 466, 2008.
[65]
K. K. Kojima and N. Okada, “mRNA retrotransposition coupled with 5′ inversion as a possible source of new genes,” Molecular Biology and Evolution, vol. 26, no. 6, pp. 1405–1420, 2009.
[66]
R. S. Baucom, J. C. Estill, C. Chaparro et al., “Exceptional diversity, non-random distribution, and rapid evolution of retroelements in the B73 maize genome,” PLoS Genetics, vol. 5, no. 11, Article ID e1000732, 2009.
[67]
K. Noma, H. Ohtsubo, and E. Ohtsubo, “ATLN elements, LINEs from Arabidopsis thaliana: identification and characterization,” DNA Research, vol. 7, no. 5, pp. 291–303, 2000.
[68]
A. Lenoir, L. Lavie, J.-L. Prieto et al., “The evolutionary origin and genomic organization of SINEs in Arabidopsis thaliana,” Molecular Biology and Evolution, vol. 18, no. 12, pp. 2315–2322, 2001.
[69]
F. Myouga, S. Tsuchimoto, K. Noma, H. Ohtsubo, and E. Ohtsubo, “Identification and structural analysis of SINE elements in the Arabidopsis thaliana genome,” Genes and Genetic Systems, vol. 76, no. 3, pp. 169–179, 2001.
[70]
J. Faris, A. Sirikhachornkit, R. Haselkorn, B. Gill, and P. Gornicki, “Chromosome mapping and phylogenetic analysis of the cytosolic acetyl-CoA carboxylase loci in wheat,” Molecular Biology and Evolution, vol. 18, no. 9, pp. 1720–1733, 2001.
[71]
Y. Zhang, Y. Wu, Y. Liu, and B. Han, “Computational identification of 69 retroposons in Arabidopsis,” Plant Physiology, vol. 138, no. 2, pp. 935–948, 2005.
[72]
D. Benovoy and G. Drouin, “Processed pseudogenes, processed genes, and spontaneous mutations in the Arabidopsis genome,” Journal of Molecular Evolution, vol. 62, no. 5, pp. 511–522, 2006.
[73]
N. Nurhayati, D. Gondé, and D. Ober, “Evolution of pyrrolizidine alkaloids in Phalaenopsis orchids and other monocotyledons: identification of deoxyhypusine synthase, homospermidine synthase and related pseudogenes,” Phytochemistry, vol. 70, no. 4, pp. 508–516, 2009.
[74]
K. Mochizuki, M. Umeda, H. Ohtsubo, and E. Ohtsubo, “Characterization of a plant SINE, p-SINE1, in rice genomes,” Japanese Journal of Genetics, vol. 67, no. 2, pp. 155–166, 1992.
[75]
J.-H. Xu, I. Osawa, S. Tsuchimoto, E. Ohtsubo, and H. Ohtsubo, “Two new SINE elements, p-SINE2 and p-SINE3, from rice,” Genes and Genetic Systems, vol. 80, no. 3, pp. 161–171, 2005.
[76]
Y. Yasui, S. Nasuda, Y. Matsuoka, and T. Kawahara, “The Au family, a novel short interspersed element (SINE) from Aegilops umbellulata,” Theoretical and Applied Genetics, vol. 102, no. 4, pp. 463–470, 2001.
[77]
J. A. Fawcett, T. Kawahara, H. Watanabe, and Y. Yasui, “A SINE family widely distributed in the plant kingdom and its evolutionary history,” Plant Molecular Biology, vol. 61, no. 3, pp. 505–514, 2006.
[78]
E. Yagi, T. Akita, and T. Kawahara, “A novel Au SINE sequence found in a gymnosperm,” Genes and Genetic Systems, vol. 86, no. 1, pp. 19–25, 2011.
[79]
Y. Shu, Y. Li, X. Bai et al., “Identification and characterization of a new member of the SINE Au retroposon family (GmAu1) in the soybean, Glycine max (L.) Merr., genome and its potential application,” Plant Cell Reports, vol. 30, no. 12, pp. 2207–2213, 2011.
[80]
M. J. Moore, C. D. Bell, P. S. Soltis, and D. E. Soltis, “Using plastid genome-scale data to resolve enigmatic relationships among basal angiosperms,” Proceedings of the National Academy of Sciences of the United States of America, vol. 104, no. 49, pp. 19363–19368, 2007.
[81]
D. H. Mathews, A. R. Banerjee, D. D. Luan, T. H. Eickbush, and D. H. Turner, “Secondary structure model of the RNA recognized by the reverse transcriptase from the R2 retrotransposable element,” RNA, vol. 3, no. 1, pp. 1–16, 1997.
[82]
Y. Nomura, M. Kajikawa, S. Baba et al., “Solution structure and functional importance of a conserved RNA hairpin of eel LINE UnaL2,” Nucleic Acids Research, vol. 34, no. 18, pp. 5184–5193, 2006.
[83]
V. Cognat, J.-M. Deragon, E. Vinogradova, T. Salinas, C. Remacle, and L. Maréchal-Drouard, “On the evolution and expression of Chlamydomonas reinhardtii nucleus-encoded transfer RNA genes,” Genetics, vol. 179, no. 1, pp. 113–123, 2008.
[84]
K. G. Karol, R. M. McCourt, M. T. Cimino, and C. F. Delwiche, “The closest living relatives of land plants,” Science, vol. 294, no. 5550, pp. 2351–2353, 2001.
[85]
M. G. Kidwell and D. Lisch, “Transposable elements as sources of variation in animals and plants,” Proceedings of the National Academy of Sciences of the United States of America, vol. 94, no. 15, pp. 7704–7711, 1997.
[86]
D. Kordi? and F. Guben?ek, “Unusual horizontal transfer of a long interspersed nuclear element between distant vertebrate classes,” Proceedings of the National Academy of Sciences of the United States of America, vol. 95, no. 18, pp. 10704–10709, 1998.
[87]
A. M. Walsh, R. D. Kortschak, M. G. Gardner, T. Bertozzi, and D. L. Adelson, “Widespread horizontal transfer of retrotransposons,” Proceedings of the National Academy of Sciences of the United States of America, vol. 110, no. 3, pp. 1012–1016, 2013.
[88]
S. Chambeyron, A. Bucheton, and I. Busseau, “Tandem UAA repeats at the 3′-end of the transcript are essential for the precise initiation of reverse transcription of the I factor in Drosophila melanogaster,” Journal of Biological Chemistry, vol. 277, no. 20, pp. 17877–17882, 2002.
[89]
M. Komatsu, K. Shimamoto, and J. Kyozuka, “Two-step regulation and continuous retrotransposition of the rice LINE-type retrotransposon Karma,” Plant Cell, vol. 15, no. 8, pp. 1934–1944, 2003.
[90]
X. Zhang and S. R. Wessler, “Genome-wide comparative analysis of the transposable elements in the related species Arabidopsis thaliana and Brassica oleracea,” Proceedings of the National Academy of Sciences of the United States of America, vol. 101, no. 15, pp. 5589–5594, 2004.
[91]
H. Yamashita and M. Tahara, “A LINE-type retrotransposon active in meristem stem cells causes heritable transpositions in the sweet potato genome,” Plant Molecular Biology, vol. 61, no. 1-2, pp. 79–94, 2006.
[92]
T. Heitkam and T. Schmidt, “BNR—a LINE family from Beta vulgaris—contains a RRM domain in open reading frame 1 and defines a L1 sub-clade present in diverse plant genomes,” Plant Journal, vol. 59, no. 6, pp. 872–882, 2009.
[93]
J. D. Hollister, L. M. Smith, Y.-L. Guo, F. Ott, D. Weigel, and B. S. Gaut, “Transposable elements and small RNAs contribute to gene expression divergence between Arabidopsis thaliana and Arabidopsis lyrata,” Proceedings of the National Academy of Sciences of the United States of America, vol. 108, no. 6, pp. 2322–2327, 2011.
[94]
E. Khazina, V. Truffault, R. Büttner, S. Schmidt, M. Coles, and O. Weichenrieder, “Trimeric structure and flexibility of the L1ORF1 protein in human L1 retrotransposition,” Nature Structural and Molecular Biology, vol. 18, no. 9, pp. 1006–1014, 2011.
[95]
S. J. Smerdon, J. J?ger, J. Wang et al., “Structure of the binding site for nonnucleoside inhibitors of the reverse transcriptase of human immunodeficiency virus type 1,” Proceedings of the National Academy of Sciences of the United States of America, vol. 91, no. 9, pp. 3911–3915, 1994.
[96]
M. Zuker, “Mfold web server for nucleic acid folding and hybridization prediction,” Nucleic Acids Research, vol. 31, no. 13, pp. 3406–3415, 2003.
[97]
A. F. A. Smit and A. D. Riggs, “MIRs are classic, tRNA-derived SINEs that amplified before the mammalian radiation,” Nucleic Acids Research, vol. 23, no. 1, pp. 98–102, 1995.
[98]
J. Jurka, E. Zietkiewicz, and D. Labuda, “Ubiquitous mammalian-wide interspersed repeats (MIRs) are molecular fossils from the mesozoic era,” Nucleic Acids Research, vol. 23, no. 1, pp. 170–175, 1995.
[99]
N. Gilbert and D. Labuda, “Evolutionary inventions and continuity of CORE-SINEs in mammals,” Journal of Molecular Biology, vol. 298, no. 3, pp. 365–377, 2000.
[100]
A. F. A. Smit, “The origin of interspersed repeats in the human genome,” Current Opinion in Genetics and Development, vol. 6, no. 6, pp. 743–748, 1996.
[101]
V. V. Kapitonov and J. Jurka, “The esterase and PHD domains in CR1-like non-LTR retrotransposons,” Molecular Biology and Evolution, vol. 20, no. 1, pp. 38–46, 2003.
[102]
N. Gilbert and D. Labuda, “CORE-SINEs: eukaryotic short interspersed retroposing elements with common sequence motifs,” Proceedings of the National Academy of Sciences of the United States of America, vol. 96, no. 6, pp. 2869–2874, 1999.
[103]
K. P. Gogolevsky, N. S. Vassetzky, and D. A. Kramerov, “Bov-B-mobilized SINEs in vertebrate genomes,” Gene, vol. 407, no. 1-2, pp. 75–85, 2008.
[104]
N. Okada and M. Hamada, “The 3′ ends of tRNA-derived SINEs originated from the 3′ ends of LINEs: a new example from the bovine genome,” Journal of Molecular Evolution, vol. 44, supplement 1, pp. S52–S56, 1997.
[105]
J. A. Lenstra, J. A. van Boxtel, K. A. Zwaagstra, and M. Schwerin, “Short interspersed nuclear element (SINE) sequences of the Bovidae,” Animal Genetics, vol. 24, no. 1, pp. 33–39, 1993.
[106]
J. Szemraj, G. P?ucienniczak, J. Jaworski, and A. P?ucienniczak, “Bovine Alu-like sequences mediate transposition of a new site-specific retroelement,” Gene, vol. 152, no. 2, pp. 261–264, 1995.
[107]
M. Nikaido, H. Nishihara, Y. Hukumoto, and N. Okada, “Ancient SINEs from African endemic mammals,” Molecular Biology and Evolution, vol. 20, no. 4, pp. 522–527, 2003.
[108]
C. Gilbert, J. K. Pace II, and P. D. Waters, “Target site analysis of RTE1_LA and its AfroSINE partner in the elephant genome,” Gene, vol. 425, no. 1-2, pp. 1–8, 2008.
[109]
H. Endoh and N. Okada, “Total DNA transcription in vitro: a procedure to detect highly repetitive and transcribable sequences with tRNA-like structures,” Proceedings of the National Academy of Sciences of the United States of America, vol. 83, no. 2, pp. 251–255, 1986.
[110]
H. Endoh, S. Nagahashi, and N. Okada, “A highly repetitive and transcribable sequence in the tortoise genome is probably a retroposon,” European Journal of Biochemistry, vol. 189, no. 1, pp. 25–31, 1990.
[111]
T. Sasaki, K. Takahashi, M. Nikaido, S. Miura, Y. Yasukawa, and N. Okada, “First application of the SINE (Short Interspersed Repetitive Element) method to infer phylogenetic relationships in reptiles: an example from the turtle superfamily testudinoidea,” Molecular Biology and Evolution, vol. 21, no. 4, pp. 705–715, 2004.
[112]
M. Kajikawa, K. Ohshima, and N. Okada, “Determination of the entire sequence of turtle CR1: the first open reading frame of the turtle CR1 element encodes a protein with a novel zinc finger motif,” Molecular Biology and Evolution, vol. 14, no. 12, pp. 1206–1217, 1997.
[113]
T. L. Vandergon and M. Reitman, “Evolution of chicken repeat 1 (CR1) elements: evidence for ancient subfamilies and multiple progenitors,” Molecular Biology and Evolution, vol. 11, no. 6, pp. 886–898, 1994.
[114]
O. Piskurek, C. C. Austin, and N. Okada, “Sauria SINEs: novel short interspersed retroposable elements that are widespread in reptile genomes,” Journal of Molecular Evolution, vol. 62, no. 5, pp. 630–644, 2006.
[115]
O. Piskurek, H. Nishihara, and N. Okada, “The evolution of two partner LINE/SINE families and a full-length chromodomain-containing Ty3/Gypsy LTR element in the first reptilian genome of Anolis carolinensis,” Gene, vol. 441, no. 1-2, pp. 111–118, 2009.
[116]
K. K. Kojima, V. V. Kapitonov, and J. Jurka, “Recent expansion of a new Ingi-related clade of Vingi non-LTR retrotransposons in hedgehogs,” Molecular Biology and Evolution, vol. 28, no. 1, pp. 17–20, 2011.
[117]
K.-I. Matsumoto, K. Murakami, and N. Okada, “Gene for lysine tRNA1 may be a progenitor of the highly repetitive and transcribable sequences present in the salmon genome,” Proceedings of the National Academy of Sciences of the United States of America, vol. 83, no. 10, pp. 3156–3160, 1986.
[118]
Y. Kido, M. Aono, T. Yamaki et al., “Shaping and reshaping of salmonid genomes by amplification of tRNA-derived retroposons during evolution,” Proceedings of the National Academy of Sciences of the United States of America, vol. 88, no. 6, pp. 2326–2330, 1991.
[119]
V. Matveev, H. Nishihara, and N. Okada, “Novel SINE families from salmons validate Parahucho (Salmonidae) as a distinct genus and give evidence that SINEs can incorporate LINE-related 3′-tails of other SINEs,” Molecular Biology and Evolution, vol. 24, no. 8, pp. 1656–1666, 2007.
[120]
R. J. Winkfein, R. D. Moir, S. A. Krawetz, J. Blanco, J. C. States, and G. H. Dixon, “A new family of repetitive, retroposon-like sequences in the genome of the rainbow trout,” European Journal of Biochemistry, vol. 176, no. 2, pp. 255–264, 1988.
[121]
Y. Terai, K. Takahashi, and N. Okada, “SINE cousins: the 3′-end tails of the two oldest and distantly related families of SINEs are descended from the 3′ ends of LINEs with the same genealogical origin,” Molecular Biology and Evolution, vol. 15, no. 11, pp. 1460–1471, 1998.
[122]
K. Takahashi, Y. Terai, M. Nishida, and N. Okada, “A novel family of short interspersed repetitive elements (SINEs) from cichlids: the patterns of insertion of SINEs at orthologous loci support the proposed monophyly of four major groups of cichlid fishes in Lake Tanganyika,” Molecular Biology and Evolution, vol. 15, no. 4, pp. 391–407, 1998.
[123]
M. Kajikawa, K. Ichiyanagi, N. Tanaka, and N. Okada, “Isolation and characterization of active LINE and SINEs from the eel,” Molecular Biology and Evolution, vol. 22, no. 3, pp. 673–682, 2005.
[124]
C. Tong, B. Guo, and S. He, “Bead-probe complex capture a couple of SINE and LINE family from genomes of two closely related species of East Asian cyprinid directly using magnetic separation,” BMC Genomics, vol. 10, article 83, 2009.
[125]
H. Nishihara, A. F. A. Smit, and N. Okada, “Functional noncoding sequences derived from SINEs in the mammalian genome,” Genome Research, vol. 16, no. 7, pp. 864–874, 2006.
[126]
I. Ogiwara, M. Miya, K. Ohshima, and N. Okada, “Retropositional parasitism of SINEs on LINEs: identification of SINEs and LINEs in elasmobranchs,” Molecular Biology and Evolution, vol. 16, no. 9, pp. 1238–1250, 1999.
[127]
I. Ogiwara, M. Miya, K. Ohshima, and N. Okada, “V-SINEs: a new superfamily of vertebrate SINEs that are widespread in vertebrate genomes and retain a strongly conserved segment within each repetitive unit,” Genome Research, vol. 12, no. 2, pp. 316–324, 2002.
[128]
Z. Izsvák, Z. Ivics, D. Garcia-Estefania, S. C. Fahrenkrug, and P. B. Hackett, “DANA elements: a family of composite, tRNA-derived short interspersed DNA elements associated with mutational activities in zebrafish (Danio rerio),” Proceedings of the National Academy of Sciences of the United States of America, vol. 93, no. 3, pp. 1077–1081, 1996.
[129]
N. Shimoda, M. Chevrette, M. Ekker, Y. Kikuchi, Y. Hotta, and H. Okamoto, “Mermaid, a family of short interspersed repetitive elements, is useful for zebrafish genome mapping,” Biochemical and Biophysical Research Communications, vol. 220, no. 1, pp. 233–237, 1996.
[130]
B. Venkatesh, E. F. Kirkness, Y.-H. Loh et al., “Survey sequencing and comparative analysis of the elephant shark (Callorhinchus milii) genome,” PLoS Biology, vol. 5, no. 4, article e101, 2007.
[131]
P. E. Nisson, R. J. Hickey, M. F. Boshar, and W. R. Crain Jr., “Identification of a repeated sequence in the genome of the sea urchin which is traescribed by RNA polymerase III and contains the features of a retroposon,” Nucleic Acids Research, vol. 16, no. 4, pp. 1431–1452, 1988.
[132]
Z. Tu, S. Li, and C. Mao, “The changing tails of a novel short interspersed element in Aedes aegypti: genomic evidence for slippage retrotransposition and the relationship between 3′ tandem repeats and the poly(dA) tail,” Genetics, vol. 168, no. 4, pp. 2037–2047, 2004.
[133]
Z. Tu and J. J. Hill, “MosquI, a novel family of mosquito retrotransposons distantly related to the Drosophila I factors, may consist of elements of more than one origin,” Molecular Biology and Evolution, vol. 16, no. 12, pp. 1675–1686, 1999.
[134]
O. Piskurek and D. J. Jackson, “Tracking the ancestry of a deeply conserved eumetazoan SINE domain,” Molecular Biology and Evolution, vol. 28, no. 10, pp. 2727–2730, 2011.
[135]
P. Kachroo, S. A. Leong, and B. B. Chattoo, “Mg-SINE: a short interspersed nuclear element from the rice blast fungus, Magnaporthe grisea,” Proceedings of the National Academy of Sciences of the United States of America, vol. 92, no. 24, pp. 11125–11129, 1995.
[136]
A. M. Shire and J. P. Ackers, “SINE elements of Entamoeba dispar,” Molecular and Biochemical Parasitology, vol. 152, no. 1, pp. 47–52, 2007.
[137]
K. Van Dellen, J. Field, Z. Wang, B. Loftus, and J. Samuelson, “LINEs and SINE-like elements of the protist Entamoeba histolytica,” Gene, vol. 297, no. 1-2, pp. 229–239, 2002.
[138]
U. Willhoeft, H. Bu?, and E. Tannich, “The abundant polyadenylated transcript 2 DNA sequence of the pathogenic protozoan parasite Entamoeba histolytica represents a nonautonomous non-long-terminal-repeat retrotransposon-like element which is absent in the closely related nonpathogenic species Entamoeba dispar,” Infection and Immunity, vol. 70, no. 12, pp. 6798–6804, 2002.
[139]
K. K. Kojima and H. Fujiwara, “An extraordinary retrotransposon family encoding dual endonucleases,” Genome Research, vol. 15, no. 8, pp. 1106–1117, 2005.
[140]
S. Tsuchimoto, Y. Hirao, E. Ohtsubo, and H. Ohtsubo, “New SINE families from rice, OsSN, with poly(A) at the 3′ ends,” Genes and Genetic Systems, vol. 83, no. 3, pp. 227–236, 2008.
[141]
J.-M. Deragon, B. S. Landry, T. Pélissier, S. Tutois, S. Tourmente, and G. Picard, “An analysis of retroposition in plants based on a family of SINEs from Brassica napus,” Journal of Molecular Evolution, vol. 39, no. 4, pp. 378–386, 1994.
[142]
A. Lenoir, T. Pélissier, C. Bousquet-Antonelli, and J. M. Deragon, “Comparative evolution history of SINEs in Arabidopsis thaliana and Brassica oleracea: evidence for a high rate of SINE loss,” Cytogenetic and Genome Research, vol. 110, no. 1–4, pp. 441–447, 2005.
[143]
X. Zhang and S. R. Wessler, “BoS: a large and diverse family of short interspersed elements (SINEs) in Brassica oleracea,” Journal of Molecular Evolution, vol. 60, no. 5, pp. 677–687, 2005.
[144]
J.-M. Deragon and X. Zhang, “Short interspersed elements (SINEs) in plants: origin, classification, and use as phylogenetic markers,” Systematic Biology, vol. 55, no. 6, pp. 949–956, 2006.
[145]
M. Gadzalski and T. Sakowicz, “Novel SINEs families in Medicago truncatula and Lotus japonicus: bioinformatic analysis,” Gene, vol. 480, no. 1-2, pp. 21–27, 2011.