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едеге е е ецедедеце, 2009, 43, 5, 795806 ег 57721 LTR-appleappleapple - apple applefl apple, appleappleapple е appleapple apple - LINE (long interspersed elements) е 3 7 apple apple LINE apple apple applefl, apple apple appleapple- е -fl ( appleappleapple Gag) е fl appleapple-, fl apple apple appleapple е, applefl - apple fl apple fl apple- LINE, apple fl 15 [24] LINE apple apple apple- ; fl - apple appleapple [5] ед appleapple L1 [6] ед apple -, flfl fl fl, fl- L1 appleapple apple 5'- еф - fl applefl, apple apple appleapple fl appleеде едефец е ефе едееедебед ец ец ецеееде ебедец ецедефеце«ец ецедеце 2009 appleapple*, ец flapple е apple е,, 119991 ее apple 30122008 ееapplefl 19022009 apple apple fl appleappleapple appleflfl - apple apple appleapple (short interspersed elements, SINE) е apple apple appleapple apple, appleapple appleapple apple - ед, apple appleapple fl - applefl : apple SINE, fl, fl SHORT INTERSPERSED REPETITIVE SEQUENCES (SINEs) AND THEIR USE AS A PHYLOGENET- IC TOOL, by D A Kramerov*, N S Vassetzky (Engelhardt Institute of Molecular Biology, Russian Academy of Sciences, Moscow, 119991 Russia; *e-mail: kramerov@eimbru) The data on one of the most common repetitive elements of eukaryotic genomes, short interspersed elements (SINEs), are reviewed Their structure, origin, and functioning in the genome are discussed The variation and abundance of these neutral genomic markers makes them a convenient and reliable tool for phylogenetic analysis The main methods of such analysis are presented, and the potential and limitations of this approach are discussed using specific examples Key words: short interspersed elements, SINEs, evolution, phylogeny * еде едефец е ефе едееедебед е fl appleflfl - - fl apple apple- ееfl appleflfl apple, apple flfl : -apple appleappleapple -apple apple apple apple- ееfl -apple apple applefl (cut and paste), apple apple apple, apple apple е appleapple apple applefl, apple apple apple flfl, - applefl apple apple appleapple ( applefl, copy and paste), -apple, flfl appleappleapple, apple (200400 ) apple (long terminal repeat, LTR) [1] еф- appleappleapple 8 apple apple apple fl, apple apple apple appleapple (appleapple) * е : kramerov@eimbru 795

796 7SL PHK (A) n B1 LINE еф SINE е е 1 appleapple apple appleapple: B1 () е SINE () apple apple; apple flfl fl applefl applefl - applefl fl L1 ее -,, applefl apple apple, fl ефfl fl fl appleapple, appleappleapple apple apple applefl - apple (SINE, short interspersed elements) apple appleapple 100 500 е«apple SINE flfl - fl fl fl apple - fl 1030%, - applefl fl SINE SINE, fl apple, apple - apple [7] SINE - applefl, apple, apple, apple ( appleapple apple) SINE apple fl applefl - fl apple apple LINE (apple 1) [8, 9] еф apple, SINE applefl ееapple apple appleapple- 1 B2, apple [10 13], Alu apple [14] applefl - SINE, fl -, apple apple- B1 Alu 7SL е, е ее-, - applefl- appleapple apple [15] ееapple 1 apple fl - 180 7SL е, -, - (apple 1) 1 fl -, А А Alu apple 1 - apple - apple еф apple, Alu flflfl apple SINE е«apple apple- 7SL е, е B2 apple е,, е ееapple 2 е, apple - applefl, 3'- applefl - fl SINE applefl (apple 1), fl е -appleapple-, е -apple apple applefl е apple appleapple appleappleapple [7], apple apple appleapple, applefl 5S appleе apple apple fl apple Danio rerio [16] apple apple appleapple applefl -, applefl 5S appleе, SINE apple [17] - [18] apple apple apple - apple appleapple

еде едефец е ефе едееедебед ец ец ецеееде ебедец 797 apple е, apple apple apple applefl е -apple III, е -apple II, - applefl appleappleapple, apple, LINE apple appleapple appleappleapplefl е -apple III, appleapplefl е [19, 20] еф, SINE appleappleapplefl apple, appleapple е -apple II, appleapple е -apple- III, applefl appleapple е, 5S appleе, 7SL е, apple applefl apple appleappleapple [7, 21] еф apple, apple appleapple е SINE fl fl fl - - appleapple SINE appleapple, apple -, applefl appleapple appleappleapplefl, appleapple apple appleapple appleapple е -- apple III SINE, apple е 7SL е, appleapple 11,, apple apple- 3035 SINE, 5S appleе, appleapple apple, IE appleapple flfl apple apple apple appleapple fl appleapple е -apple III ец, appleapplefl, flfl е -apple, appleapplefl apple еф [19, 21] ецapple, SINE appleapple е, appleapplefl applefl apple fl еф-- apple 3'- apple fl SINE applefl -- ее appleapple - apple appleappleapple [22] ег, apple 2- apple-, appleapple appleapple-, еф еф SINE flfl еф +, SINE appleapple еф еф ефappleapplefl B2 SINE fl еф-, 3'- - applefl е applefl () [23] fl applefl еф е appleapple applefl - apple е -apple III ее(), applefl appleapple SINE, - apple applefl е apple apple apple - fl А А SINE apple apple ( apple SINE ) apple : apple fl apple 35 (apple 2) SINE -, apple apple 30100, 3'-- apple LINE (apple 2) [24], appleapple Bov-B, CR-1, L2, RSg-1 apple apple ег-, е SINE fl appleapple LINE, apple apple appleapple е apple SINE - apple -, apple LINE- L1 fl, - () - е SINE, appleapple [25, 26] ее apple applefl apple SINE apple - [27, 28] е SINE appleapple LINE appleappleapplefl flapple, - fl, appleapple-, - SINE е apple, - fl е applefl, 3'- apple appleapple apple appleapple ее apple appleappleapplefl apple apple apple apple apple -apple apple - applefl fl fl SINE, applefl applefl apple 816 (target site duplication, TSD) applefl fl TSD applefl apple apple appleapplefl - fl apple ее apple appleapple apple TSD, appleapple SINE applefl apple SINE, apple ееfl apple, appleflfl - appleapple apple apple, apple appleapple flfl apple Ther-1 SINE apple, Ther-1, applefl Alu apple applefl [29], fl А apple А fl - apple appleapple:, apple Alu applefl; apple apple apple, SINE, flfl apple, - applefl [30] ецеееде ебедец еде едефец е ефе едееедебедед ецедефеце«ец ецедеце 90- apple apple appleappleapple apple apple - apple

798 OPC1 LINE OPC2 SINE fl fl SINE ефappleapplefl е -apple II ефappleapplefl е -apple III PK LINE ефappleflfl едapplefl appleapplefl applefl едapplefl appleapple LINE е 2 apple apple appleapple еде applefl apple fl (); apple flfl apple apple apple apple, - fl apple apple apple А NatureА, applefl apple [31] apple - flapple, appleapple,, apple applefl appleapple apple apple ед,, - flapple, - fl applefl, apple apple apple, flapple - appleapple apple fl apple apple [3234] ец fl apple -, applefl,, е apple apple ед fl -, fl apple е apple apple, - apple 90- apple SINE apple [3537] fl, -, apple SINE, е -apple - apple apple (еее ), - SINE, appleapplefl apple apple apple,, apple - apple 3 1, 2 3 apple - SINE, 4 apple applefl, SINE 1 2, 3 4 е fl apple apple, - apple apple 3 ец apple, apple apple apple - apple appleapple appleappleapple, fl - apple apple, apple- 1999 [38] applefl еее apple - apple appleapple е -apple III apple fl SINE, apple apple, applefl еее -apple, apple apple apple apple apple apple - fl apple SINE, fl - SINE, apple - fl appleapple apple-

еде едефец е ефе едееедебед ец ец ецеееде ебедец 799 1 2 3 4 apple appleapplefl fl [40] ее apple apple apple appleapple apple apple flapple [41] apple -, applefl Afrotheria [33] apple, apple, fl apple, apple apple apple, apple apple - fl apple fl - apple - apple appleapple AfroSINE [42] е apple apple fl Afrotheria, apple -apple, fl applefl Supraprimates ( Euarchontoglires) [33] - - applefl еб еб ееapplefl fl applefl ееappleapple ' ееappleapple ' SINE 1 2 3 4 е 3 ец apple appleapple fl еб appleapple /' /' fl apple appleapple apple apple flfl apple appleapple еее applefl ее apple - apple, apple- - е 4 appleapple apple fl applefl apple А А apple, fl apple Romer [39], - appleapple apple appleapple apple е apple, B1-dID, - applefl apple- B1-dID apple apple - : (Sciuridae), (Aplodontidae) (Gliridae) apple Romer apple apple apple () apple ( apple 4 ) ец appleapplefl B1-dID

800 B1 ID B4 45SH 45S1 B2 B1-dID IDL-Geo DIP Cricetidae Muridae Spalacidae Rhizomyidae Dipodidae Zapodidae Geomyidae Heteromyidae Gliridae Aplodontidae Sciuridae Castoridae Hystricidae Anomaluridae Hydrochoeridae Caviidae Dasyproctidae DIP B2, 45S1 B4, 45SH IDL-Geo B1-dID е 4 е apple apple ее apple apple appleapple apple - apple [37, 40, 46, 7274] apple apple flfl apple appleapple еб fl Gliridae Romer [39]

еде едефец е ефе едееедебед ец ец ецеееде ебедец 801 apple apple fl appleapple [43, 44] ед, apple apple applefl, apple, apple, apple apple apple 7SL е, apple apple [45],, - apple apple, applefl applefl apple, - 7SL е apple apple SINE 1 apple Alu apple appleapple SINE, - еф, apple apple 1 apple apple - 1 apple apple fl apple apple, fl fl appleapplefl appleapple apple apple - apple apple [46] appleappleapple appleapple -, - apple appleapple applefl SINE apple - apple [47] е - еее - apple fl, - apple SINE (apple 5) apple apple apple, apple SINE fl applefl apple - apple apple, SINE, apple SINE, apple apple,, apple -, apple apple - appleflapple- е apple apple fl fl - fl apple [48] - [49] apple (/) SINE Alu apple fl fl apple, applefl apple apple ( apple [30] ) fl apple, appleapple,, applefl - apple (Cetacea) applefl apple apple applefl apple (Artiodactyla) еф apple, fl, fl [50, 51] ее apple apple apple apple apple - flapple [32] ецapple-sine-еее, apple fl apple appleapple- е, apple-sineеее, fl еее, apple - apple apple SINE ееappleapple apple apple, apple, apple fl SINE (apple 6) apple applefl apple еее applefl apple, apple - fl apple applefl е apple fl apple [52] apple applefl appleapple [53], fl- [5456], flapple [57] apple [58] apple, apple appleapple fl apple appleapple fl apple, fl, applefl А appleа еф apple, apple apple, apple apple apple ( ), apple appleapple - appleapple apple ед ( apple apple) apple appleapple apple, appleapple е apple apple, ее, apple appleapple- -, apple fl (rare genomic changes), apple apple fl, apple LINE apple apple, applefl -, [59] -, apple appleapple apple -, apple apple, SINE apple ( apple fl - ) apple - apple apple - fl apple applefl, apple apple appleapple fl 1 -, SINE applefl, applefl 4

802 1 2 3 4 1 2 3 4 1 2 3 4 еее SINE 1 2 3 4 е 5 ец apple appleapple fl apple apple apple fl apple apple apple apple appleapple apple, fl SINE appleflfl apple applefl, apple - 2 - apple appleapple apple apple applefl applefl - apple SINE - apple 3 apple appleapple applefl apple, apple, apple SINE apple ( apple - apple) еф, fl apple apple appleapple- appleapple appleflfl apple, appleflapple SINE, applefl apple appleapple- appleapple apple [60] [61], - fl fl fl apple appleapple еф apple - fl apple apple appleapple ефfl fl apple -, apple, appleapplefl ( flapple,

еде едефец е ефе едееедебед ец ец ецеееде ебедец 803 1 2 3 4 fl д fl д fl д fl д 1 2 3 4 1 еее д д 2 3 4 е 6 ец apple appleapple fl ецapple-sine-еее apple appleapple SINE apple apple apple д д apple appleapple apple),, -, apple TSD ее apple- 051% apple-, fl apple Alu, fl, apple- SINE [62] ед apple - fl, apple, appleflfl apple apple apple appleapple applefl - е,, 3'- - LINE, appleapple SINE ед appleapple - ед fl - - applefl apple apple е -apple apple apple - SINE, apple apple apple fl, applefl е, - [63] еф, apple appleapple- ID apple, Vic-1 apple Das-1 apple-, applefl е, apple apple еф apple, apple apple appleapple - ( apple fl) apple apple apple appleappleapple- appleflapple fl appleapple apple apple fl -apple LTR-appleappleapple [64] apple apple appleapple, fl,, apple - LINE ( LINE Bov-B [18, 65]), appleapple appleapple apple appleapple ( - А apple apple SINEА appleapple [65]), - apple apple - LINE, apple apple applefl apple appleapple apple apple SINE - apple appleapple appleappleapple [66] apple appleapple SmaI appleappleapple apple, apple, apple, apple, applefl apple - е - apple [66] appleapple, fl, apple fl: apple appleappleapple - apple apple, apple apple, А А apple flflfl apple, - fl apple - apple, apple appleapple fl,, 4*

804 fl А«А» ецapplefl SINE fl А«+А» fl А«+А» е 7 apple apple apple apple, apple apple apple appleapple - apple apple ее apple SINE apple fl appleapple [67], appleapple apple appleapple applefl apple apple, - Schistosoma japonicum [68, 69] ед apple apple appleapple apple apple, ( appleapple) А apple appleа [70] - apple apple appleapple appleflfl apple applefl, SINE -, apple apple - appleappleapple LINE еф apple, apple SINE apple apple apple appleapple ( apple apple), apple SINE fl apple, apple - apple appleapple ед fl, apple apple- SINE - fl apple е fl, appleapple : apple apple apple - apple apple ( fl SINE apple ) appleflfl apple applefl applefl apple apple- (apple 7) fl, - apple apple apple [71] ее appleapple-, applefl applefl apple applefl apple appleapple fl -, apple, appleappleapple - SINE, apple fl appleapple apple -, fl - fl fl applefl apple applefl applefl, - applefl apple еб apple apple applefl fl apple, apple flfl, appleapple apple - fl applefl apple applefl apple apple fl - fl SINE - fl apple applefl appleapple applefl apple apple apple appleapple fl fl apple -, apple, fl fl,, apple fl apple, apple

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