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åéãÖäìãüêçÄü ÅàéãéÉàü, 2009, ÚÓÏ 43, ‹ 5, Ò. 795–806 795 * äéêéíäàÖ êÖíêéèéáéçõ ê‡ÒÒÂflÌÌ˚ ÔÓ „ÂÌÓÏÛ ÔÓ‚ÚÓfl˛˘ËÂÒfl ÔÓÒΉÓ-‚‡ÚÂθÌÓÒÚË Ó˜Â̸ ÏÌÓ„Ó˜ËÒÎÂÌÌ˚ Ë ÏÓ„ÛÚ ÒÓÒÚ‡‚-ÎflÚ¸ ‰Ó ÔÓÎÓ‚ËÌ˚ „ÂÌÓχ ˝Û͇ËÓÚ˘ÂÒÍËı Ó„‡-ÌËÁÏÓ‚. èÓ‰‡‚Îfl˛˘Â ·Óθ¯ËÌÒÚ‚Ó Ú‡ÍËı ÔÓ‚ÚÓ-fl˛˘ËıÒfl ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ Ô‰ÒÚ‡‚ÎÂÌÓ ÏÓ·ËθÌ˚ÏË „ÂÌÂÚ˘ÂÒÍËÏË ˝ÎÂÏÂÌÚ‡ÏË, ÍÓÚÓ˚ ‰ÂÎflÚÒfl ̇ ‰‚‡ ·Óθ¯Ëı Í·ÒÒ‡: Ñçä-Ú‡ÌÒÔÓÁÓÌ˚ Ë ÂÚÓÚ‡ÌÒÔÓÁÓÌ˚. Ñçä-Ú‡ÌÒÔÓÁÓÌ˚ ÍÓ‰ËÛ˛Ú ÙÂÏÂÌÚ Ú‡ÌÒÔÓ-Á‡ÁÛ. èÓfl‚ÎÂÌË ÍÓÔËÈ Ñçä-Ú‡ÌÒÔÓÁÓÌÓ‚ ‚ ÌÓ‚˚ı ÏÂÒÚ‡ı „ÂÌÓχ ÔÓËÒıÓ‰ËÚ ÔÓ ÏÂı‡ÌËÁÏÛ ‚˚ÂÁ‡ÌËfl (cutandpaste), Ú.Â. Ú‡ÌÒÔÓÁ‡Á‡ ‚˚ÂÁ‡ÂÚ Ú‡ÌÒÔÓÁÓÌ ËÁ „ÂÌÓχ, ‡ Á‡ÚÂÏ Í‡Ú‡ÎËÁËÛÂÚ Â„Ó ‚ÒÚ‡Ë‚‡ÌË ‚ ÌÓ‚Ó ÏÂÒÚÓ. êÂÚÓÚ‡ÌÒÔÓÁÓÌ˚ ËÒÔÓθÁÛ˛Ú ÒÓ‚Â¯ÂÌÌÓ ËÌÓÈ ÏÂı‡ÌËÁÏ ‚ÒÚ‡Ë‚‡ÌËfl, ÔË ÍÓÚÓÓÏ ÒÚ‡˚ ÍÓÔËË Ì ۉ‡Îfl˛ÚÒfl, ‡ ÍÓÔËË ‚ ÌÓ‚˚ı ÏÂÒÚ‡ı ‚ÓÁÌË-͇˛Ú ·Î‡„Ó‰‡fl ÔÓˆÂÒÒÛ Ó·‡ÚÌÓÈ Ú‡ÌÒÍËÔˆËË (ÏÂı‡ÌËÁÏ ÍÓÔËÓ‚‡ÌËfl, copy and paste). ä ˝ÚÓÏÛ Í·ÒÒÛ ˝ÎÂÏÂÌÚÓ‚, ‚Ó-ÔÂ‚˚ı, ÓÚÌÓÒflÚÒfl ÂÚÓ-Ú‡ÌÒÔÓÁÓÌ˚, ÒÓ‰Âʇ˘Ë ‰ÎËÌÌ˚ (200–400 Ô.Ì.) ÍÓ̈‚˚ ÔÓ‚ÚÓ˚ (longterminalrepeat, LTR) [1]. í‡-ÍË ÂÚÓÚ‡ÌÒÔÓÁÓÌ˚ ‰ÎËÌÓÈ ‰Ó 8 Ú.Ô.Ì. ËÏÂ˛Ú ‰‚ ËÎË ÚË ÓÚÍ˚Ú˚ ‡ÏÍË Ò˜ËÚ˚‚‡ÌËfl, Ӊ̇ ËÁ ÍÓÚÓ-˚ı ÍÓ‰ËÛÂÚ Ó·‡ÚÌÛ˛ Ú‡ÌÒÍËÔÚ‡ÁÛ (‚ÂÚ‡ÁÛ). * ùÎ. ÔÓ˜Ú‡: [email protected] åÌÓ„Ë ËÁ LTR-ÂÚÓÚ‡ÌÒÔÓÁÓÌÓ‚ – ˝ÚÓ ÔÓ ÒÛÚË ˝Ì-‰Ó„ÂÌÌ˚ ÂÚÓ‚ËÛÒ˚, ÔÓÚÂfl‚¯Ë ÒÔÓÒÓ·ÌÓÒÚ¸ Í ËÌÙˈËÓ‚‡Ì˲ ÍÎÂÚÓÍ, ÌÓ ÒÔÓÒÓ·Ì˚ ӷ‡ÁÓ‚˚-‚‡Ú¸ ÌÓ‚˚ ÔÓ‚ËÛÒÌ˚ ÍÓÔËË ‚ „ÂÌÓÏÂ. êÂÚÓÚ‡ÌÒÔÓÁÓÌ˚ ‰Û„Ó„Ó ÚËÔ‡ ̇Á˚‚‡˛Ú ‰ÎËÌ-Ì˚ÏË ÂÚÓÔÓÁÓ̇ÏË, ËÎË LINE (longinterspersed el-ements). ùÎÂÏÂÌÚ˚ ˝ÚÓ„Ó ÚËÔ‡ ËÏÂ˛Ú ‰ÎËÌÛ ÓÚ 3 ‰Ó 7Ú.Ô.Ì. Ë Ì ÒÓ‰ÂÊ‡Ú ‰ÎËÌÌ˚ı ÍÓ̈‚˚ı ÔÓ‚ÚÓÓ‚. LINE Ó·˚˜ÌÓ ÒÓ‰ÂÊ‡Ú Ó‰ÌÛ ËÎË ‰‚ ÓÚÍ˚Ú˚ ‡Ï-ÍË Ò˜ËÚ˚‚‡ÌËfl, ÍÓ‰ËÛ˛˘Ë ӷ‡ÚÌÛ˛ Ú‡ÌÒÍËÔ-Ú‡ÁÛ Ë êçä-Ò‚flÁ˚‚‡˛˘ËÈ ·ÂÎÓÍ (‡Ì‡ÎÓ„ ÂÚÓ‚Ë-ÛÒÌÓ„Ó Gag). ùÚË ·ÂÎÍË ÌÂÓ·ıÓ‰ËÏ˚ ‰Îfl ÂÚÓÔÓ-ÁˈËË ‰‡ÌÌ˚ı ˝ÎÂÏÂÌÚÓ‚, Ú.Â. ‰Îfl ÔÓˆÂÒÒ‡ Ó·‡ÚÌÓÈ Ú‡ÌÒÍËÔˆËË Ëı êçä, ÔË‚Ó‰fl˘ÂÈ Í Ó·-‡ÁÓ‚‡Ì˲ ÌÓ‚˚ı ÍÓÔËÈ ‚ „ÂÌÓÏÂ. ä ̇ÒÚÓfl˘ÂÏÛ ‚ÂÏÂÌË ÓÔËÒ‡ÌÓ ÌÂÒÍÓθÍÓ ‰ÂÒflÚÍÓ‚ ÒÂÏÂÈÒÚ‚ ‡Á-Ì˚ı LINE, ÍÓÚÓ˚ ӷ˙‰ËÌfl˛Ú ‚ 15 Í·‰ [2–4]. LINE ӷ̇ÛÊÂÌ˚ ‚ „ÂÌÓχı Ò‡Ï˚ı ‡ÁÌ˚ı ˝Û͇Ë-ÓÚ – ˝Û͇Ë-ÓÚ ·ÂÒÔÓÁ‚ÓÌÓ˜Ì˚ı ‰Ó ˜ÂÎÓ‚Â͇; ‰Îfl Ó‰ÌÓÍÎÂ-ÚÓ˜Ì˚ı ˝Û͇ËÓÚ ÓÌË ÏÂÌ ı‡‡ÍÚÂÌ˚ [5]. é‰ËÌ ËÁ ̇˷ÓΠËÁÛ˜ÂÌÌ˚ı ‰ÎËÌÌ˚ı ÂÚÓÔÓÁÓÌÓ‚ – ˝ÎÂÏÂÌÚ L1 [6]. éÌ ÔËÒÛÚÒÚ‚ÛÂÚ ‚ „ÂÌÓχı ‚ÒÂı ÏÎÂ-ÍÓÔËÚ‡˛˘Ëı, Ë ˜ËÒÎÓ Â„Ó ÍÓÔËÈ ‚ „ÂÌÓÏ ËÒ˜ËÒÎflÂÚ-Òfl ‰ÂËÒ˜ËÒÎflÂÚ-ÒflÚ͇ÏË Ú˚ËÒ˜ËÒÎflÂÚ-Òfl˜. çÛÊÌÓ ÓÚÏÂÚËÚ¸, ˜ÚÓ ÔÓ‰‡‚Îfl-˛˘Â ˜ËÒÎÓ ÍÓÔËÈ L1 ÌÂÔÓÎÌÓ‡ÁÏÂÌ˚ Ë ÒËθÌÓ ÛÍÓÓ˜ÂÌ˚ Ò 5'-ÍÓ̈‡. í‡ÍË ÍÓÔËË Ì ÒÔÓÒÓ·Ì˚ Ò‡-ÏÓÒÚÓflÚÂθÌÓ ‡ÁÏÌÓʇڸÒfl, Ú‡Í Í‡Í Ì ÍÓ‰ËÛ˛Ú ÔÓÎÌÓˆÂÌÌÛ˛ Ó·‡ÚÌÛ˛ Ú‡ÌÒÍËÔÚ‡ÁÛ. ÑÎfl Ëı

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© 2009 „. Ñ. Ä. ä‡ÏÂÓ‚*, ç. ë. LJÒˆÍËÈ

äβ˜Â‚˚ ÒÎÓ‚‡: ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚, SINE, ˝‚ÓβˆËfl, ÙËÎÓ„ÂÌËfl.

SHORTINTERSPERSEDREPETITIVESEQUENCES (SINEs) ANDTHEIRUSEASA PHYLOGENET-ICTOOL, byD. A. Kramerov*, N. S. Vassetzky (EngelhardtInstituteofMolecularBiology, RussianAcademy ofSciences, Moscow, 119991 Russia; *e-mail: [email protected]). Thedataononeofthemostcommon re-petitiveelementsofeukaryoticgenomes, shortinterspersedelements (SINEs), arereviewed. Theirstructure, origin, andfunctioninginthegenomearediscussed. Thevariationandabundanceoftheseneutralgenomic markersmakesthemaconvenientandreliabletool for phylogenetic analysis. The main methods of such anal-ysis are presented, and the potential and limitations of this approach are discussed using specific examples. Key words: short interspersed elements, SINEs, evolution, phylogeny.

ÏÌÓÊÂÌËfl Ú·ÛÂÚÒfl Ó·‡Ú̇fl Ú‡ÌÒÍËÔÚ‡Á‡, Á‡ÍÓ-‰ËÓ‚‡Ì̇fl ‚ ÌÂÏÌÓ„Ëı ÔÓÎÌÓˆÂÌÌ˚ı ÍÓÔËflı L1. èÓÎÌÓˆÂÌÌ˚ ÍÓÔËË Ì‡Á˚‚‡˛Ú ‡‚ÚÓÌÓÏÌ˚ÏË ˝ÎÂ-ÏÂÌÚ‡ÏË, ÚÓ„‰‡ Í‡Í ÍÓÔËË, ÔÓÚÂfl‚¯Ë ÒÔÓÒÓ·ÌÓÒÚ¸ ÍÓ‰ËÓ‚‡Ú¸ ÌÓχθÌ˚ ÔÓÎËÔÂÔÚˉ˚, ̇Á˚‚‡˛ÚÒfl ̇‚ÚÓÌÓÏÌ˚ÏË. í‡Í‡fl ÒËÚÛ‡ˆËfl ̇·Î˛‰‡ÂÚÒfl ‚Ó ‚ÒÂı ÒÂÏÂÈÒÚ‚‡ı ‰ÎËÌÌ˚ı ÂÚÓÔÓÁÓÌÓ‚. ç‡ÍÓ̈, ÔÓÒΉÌËÈ ÚËÔ ÂÚÓÚ‡ÌÒÔÓÁÓÌÓ‚ – ÍÓ-ÓÚÍË ÂÚÓÔÓÁÓÌ˚, ËÎË ÍÓÍÓ-ÓÚÍË ‡ÒÒÂflÌÌ˚ ÔÓ-‚ÚÓ˚ (SINE, short interspersed elements). äÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ ËÏÂ˛Ú ‰ÎËÌÛ ÓÚ 100 ‰Ó 500 Ô.Ì. óËÒÎÓ ÍÓÔËÈ ‡ÁÌ˚ı SINE ‚ „ÂÌÓÏ ÏÓÊÂÚ ËÒ˜ËÒÎflÚ¸Òfl ‰Â-ÒflÚ͇ÏË Ë ÒÓÚÌflÏË Ú˚Òfl˜. äÓÌÍÂÚÌ˚ ÍÓÔËË Ó·˚˜-ÌÓ ÓÚ΢‡˛ÚÒfl Á‡ÏÂ̇ÏË 10–30% ÌÛÍÎÂÓÚˉӂ, ‚ÓÁ-ÌËͯËÏË Á‡ ‚ÂÏfl ÒÛ˘ÂÒÚ‚Ó‚‡ÌËfl SINE ‚ „ÂÌÓÏÂ. äÓÔËË SINE, ӷ·‰‡˛˘Ë Á̇˜ËÚÂθÌ˚Ï ÒıÓ‰ÒÚ‚ÓÏ ÏÂÊ‰Û ÒÓ·ÓÈ Ë Ò‚flÁ‡ÌÌ˚ ӷ˘ËÏ ÔÓËÒıÓʉÂÌËÂÏ, Ó·‡ÁÛ˛Ú ÒÂÏÂÈÒÚ‚‡. Ç „ÂÌÓχı ÏÎÂÍÓÔËÚ‡˛˘Ëı Ú‡-ÍËı ÒÂÏÂÈÒÚ‚ Ó·˚˜ÌÓ ‰‚‡–˜ÂÚ˚ [7]. SINE ‚Ó ÏÌÓ-ÊÂÒÚ‚Â ‚ÒÚ˜‡˛ÚÒfl ̇ Û˜‡ÒÚ͇ı, ‡ÒÔÓÎÓÊÂÌÌ˚ı ÏÂÊ‰Û „Â̇ÏË, ‡ Ú‡ÍÊ ‚ ËÌÚÓ̇ı, ÂÊ ‚ ˝ÍÁÓ̇ı (Ó·˚˜ÌÓ ‚ Ëı ÌÂÚ‡ÌÒÎËÛÂÏ˚ı ‡ÈÓ̇ı). SINE Ì ÍÓ‰ËÛ˛Ú ÔÓÎËÔÂÔÚˉ˚ Ë ‰Îfl ‡ÁÏÌÓÊÂÌËfl ÌÛʉ‡-˛ÚÒfl ‚ Ó·‡ÚÌÓÈ Ú‡ÌÒÍËÔÚ‡ÁÂ, ÍÓ‰ËÛÂÏÓÈ LINE (ËÒ. 1) [8, 9]. í‡ÍËÏ Ó·‡ÁÓÏ, ‚Ò SINE Ô‰ÒÚ‡‚Îfl˛Ú ÒÓ·ÓÈ Ì‡‚ÚÓÌÓÏÌ˚ ÏÓ·ËθÌ˚ ˝ÎÂÏÂÌÚ˚. èÂ‚˚ ËÁ‚ÂÒÚÌ˚ ÒÂÏÂÈÒÚ‚‡ ÍÓÓÚÍËı ÂÚÓÔÓ-ÁÓÌÓ‚ – Ç1 Ë B2, ̇ȉÂÌÌ˚ ‚ „ÂÌÓÏ „˚ÁÛÌÓ‚ [10– 13], Ë Alu ÔËχÚÓ‚ [14] – Ô‰ÒÚ‡‚Îfl˛Ú ‰‚‡ ÓÒÌÓ‚-Ì˚ı Í·ÒÒ‡ SINE, ÓÚ΢‡˛˘ËıÒfl ÚËÔÓÏ ÌÛÍÎÂÓÚˉ-Ì˚ı ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ, ËÁ ÍÓÚÓ˚ı ÓÌË ÔÓËÁÓ-¯ÎË. çÛÍÎÂÓÚˉÌ˚ ÔÓÒΉӂ‡ÚÂθÌÓÒÚË B1 Ë Alu ÒıÓ‰Ì˚ Ò 7SL êçä, ÍÓÏÔÓÌÂÌÚÓÏ êçè-˜‡ÒÚˈ, ÛÁ̇-˛˘Ëı Ò˄̇θÌ˚È ÔÂÔÚˉ Ë Û˜‡ÒÚ‚ÛÛÁ̇-˛˘Ëı ‚ Ú‡ÌÒÎfl-ˆËË ÒÂÍÂÚËÛÂÏ˚ı Ë ÏÂÏ·‡ÌÌ˚ı ·ÂÎÍÓ‚ [15]. èË ‚ÓÁÌËÍÌÓ‚ÂÌËË Ç1 ÔÓËÁӯ· ‰ÂΈËfl Û˜‡ÒÚ͇ ‰ÎË-ÌÓÈ 180 Ì. 7SL êçä, ‰ÓÔÓÎÌËÚÂθÌ˚ Ì·Óθ¯Ë ‰Â-ΈËË Ë ‰ÛÔÎË͇ˆËË, ‡ Ú‡ÍÊ ÏÌÓ„Ó˜ËÒÎÂÌÌ˚ ÌÛÍ-ÎÂÓÚˉÌ˚ Á‡ÏÂÌ˚ (ËÒ. 1‡). ç‡ ÍÓ̈ Ç1 ËÏÂÂÚÒfl Ä-·Ó„‡Ú˚È Û˜‡ÒÚÓÍ, Ó·˚˜ÌÓ Ì‡Á˚‚‡ÂÏ˚È “ı‚ÓÒÚÓÏ”. Alu ÔËχÚÓ‚ ÒÓÒÚÓËÚ ËÁ ‰‚Ûı ÒıÓ‰Ì˚ı Ò Ç1 ÌÛÍÎÂÓ-ÚˉÌ˚ı ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ – ÎÂ‚Ó„Ó Ë Ô‡‚Ó„Ó ÏÓ-ÌÓÏÂ‡. í‡ÍËÏ Ó·‡ÁÓÏ, Alu fl‚ÎflÂÚÒfl ‰ËÏÂÌ˚Ï SINE. óËÒÎÓ ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚, ÔÓ-ËÁӯ‰¯Ëı ÓÚ 7SL êçä, Ì‚ÂÎËÍÓ. ùÎÂÏÂÌÚ B2 ÔÓËÒıÓ‰ËÚ ÓÚ Úêçä, ‡ ËÏÂÌÌÓ, ÓÚ ‡Î‡ÌËÌÓ‚ÓÈ Úêçä. èÂ‰ÌËÈ ÍÓ̈ Ç2 ӷ·‰‡ÂÚ ÒıÓ‰ÒÚ‚ÓÏ Ò ˝ÚÓÈ Úêçä, ‰‡Î ÒΉÛÂÚ ‡ÈÓÌ ÌÂËÁ-‚ÂÒÚÌÓ„Ó ÔÓËÒıÓʉÂÌËfl, ‡ ̇ 3'-ÍÓ̈ ڇÍÊ ‡ÒÔÓ-·„‡ÂÚÒfl Ä-·Ó„‡Ú˚È ı‚ÓÒÚ. ÅÓθ¯ËÌÒÚ‚Ó ËÁ‚ÂÒÚÌ˚ı Í Ì‡ÒÚÓfl˘ÂÏÛ ÏÓÏÂÌÚÛ SINE ËÏÂ˛Ú ‡Ì‡Îӄ˘Ì˚È ÔÎ‡Ì ÒÚÓÂÌËfl (ËÒ. 1·), ıÓÚfl Í‡Í Úêçä-Ô‡Ó‰ËÚÂ-ÎË, Ú‡Í Ë Úêçä-ÌÂÓ‰ÒÚ‚ÂÌÌ˚ ‡ÈÓÌ˚ ˜‡˘Â ‚ÒÂ„Ó ‡Á΢‡˛ÚÒfl. ùÚÓÚ Í·ÒÒ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ Ò‡Ï˚È ‡ÒÔÓÒÚ‡ÌÂÌÌ˚È [7]. ç‡ÍÓ̈, ÒÛ˘ÂÒÚ‚ÛÂÚ ÚÂÚËÈ Í·ÒÒ ÍÓÓÚÍËı Â-ÚÓÔÓÁÓÌÓ‚, ÔÓËÒıÓ‰fl˘Ëı ÓÚ 5S êçä. ÇÔÂ‚˚ ÓÌ ·˚Î ÓÔËÒ‡Ì ‡ÏÂË͇ÌÒÍËÏË ËÒÒΉӂ‡ÚÂÎflÏË Û ˚·˚ Danio rerio [16]. Ç ÔÂ‰ÌÂÈ ˜‡ÒÚË ˝ÚÓ„Ó ÍÓ-ÓÚÍÓ„Ó ÂÚÓÔÓÁÓ̇ ‡ÒÔÓ·„‡ÂÚÒfl ÔÓÒΉӂ‡ÚÂθ-ÌÓÒÚ¸, Ó‰ÒÚ‚ÂÌ̇fl 5S êçä, Á‡ÚÂÏ ÒΉÛÂÚ ÚÂÎÓ SINE Ë Â„Ó ı‚ÓÒÚ. 片‚ÌÓ Û ˚· [17] Ë ÏÎÂÍÓÔËÚ‡˛-˘Ëı [18] ·˚ÎË ÓÚÍ˚Ú˚ ‰Û„Ë Ô‰ÒÚ‡‚ËÚÂÎË ˝ÚÓ-„Ó Í·ÒÒ‡ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚. ‡ · 7SL PHK (A)_n B1 Ï˚¯Ë LINE íËÔ˘Ì˚È SINE Úêçä êËÒ. 1. ëÚÛÍÚÛ‡ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚: B1 Ï˚¯Ë (‡) Ë ÚËÔ˘Ì˚È Úêçä SINE (·). ëÚÂÎÍË Ó·ÓÁ̇˜‡˛Ú ÔÓ‚ÚÓ˚; ‰Û„Ë ÔÓflÒÌÂÌËfl ‚ ÚÂÍÒÚÂ.

åéãÖäìãüêçÄü ÅàéãéÉàü      ÚÓÏ 43      ‹ 5      2009 ÇÒ ÚË ÚËÔ‡ êçä, ÓÚ ÍÓÚÓ˚ı ‚‰ÛÚ Ò‚Ó ÔÓËÒ-ıÓʉÂÌË ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚, ÒËÌÚÂÁËÛ˛ÚÒfl êçä-ÔÓÎËÏÂ‡ÁÓÈ III, ‡ Ì êçä-ÔÓÎËÏÂ‡ÁÓÈ II, ÍÓ-ÚÓ‡fl Ú‡ÌÒÍË·ËÛÂÚ „ÂÌ˚, ÍÓ‰ËÛ˛˘Ë ·ÂÎÍË, ‚ ÚÓÏ ˜ËÒΠ„ÂÌ˚ LINE. ë‡ÏË ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ ÚÓÊ Ú‡ÌÒÍË·ËÛ˛ÚÒfl êçä-ÔÓÎËÏÂ‡ÁÓÈ III, ‚ Â-ÁÛθڇÚ ˜Â„Ó Ó·‡ÁÛ˛ÚÒfl Ì·Óθ¯Ë ÔÓ ‰ÎËÌ êçä [19, 20]. íÓ, ˜ÚÓ SINE Ú‡ÌÒÍË·ËÛ˛ÚÒfl ËÏÂÌÌÓ ˝ÚËÏ ÙÂÏÂÌÚÓÏ, ÌÂÒÎÛ˜‡ÈÌÓ. Ç ÓÚ΢ˠÓÚ ÔÓÏÓ-ÚÓ‡ êçä-ÔÓÎËÏÂ‡Á˚ II, ÔÓÏÓÚÓ êçä-ÔÓÎËÏÂ‡-Á˚ III, ̇Ô‡‚Îfl˛˘ÂÈ Ú‡ÌÒÍËÔˆË˛ „ÂÌÓ‚ Úêçä, 5S êçä, 7SL êçä, ‡ Ú‡ÍÊ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚, ‡ÒÔÓ·„‡ÂÚÒfl ‚ÌÛÚË Ú‡ÌÒÍË·ËÛÂÏÓÈ Ñçä [7, 21]. í‡ÍËÏ Ó·‡ÁÓÏ, ÔÓÒΠӷ‡ÚÌÓÈ Ú‡ÌÒÍËÔˆËË êçä SINE ÌÓ‚‡fl „ÂÌÓÏ̇fl ÍÓÔËfl ËÏÂÂÚ ÔÓÎÌÓˆÂÌ-Ì˚È ÔÓÏÓÚÓ, ˜ÚÓ ‰Â·ÂÚ Â ÒÔÓÒÓ·ÌÓÈ Í ‰‡Î¸ÌÂÈ-¯ËÏ ˆËÍÎ‡Ï Ú‡ÌÒÍËÔˆËË. Ç ÓÚ΢ˠÓÚ SINE Â-ÚÓÔÒ‚‰Ó„ÂÌ˚, ‚ÓÁÌËͯË ËÁ ·ÂÎÓÍÍÓ‰ËÛ˛˘Ëı „Â-ÌÓ‚, ÚÂfl˛Ú ÔÓÏÓÚÓ˚ Ë Ì Ú‡ÌÒÍË·ËÛ˛ÚÒfl, ‡ ÔÓÚÓÏÛ Ì ÒÔÓÒÓ·Ì˚ Í ‰‡Î¸ÌÂȯÂÈ ÂÚÓÔÓÁˈËË Ë ‡ÁÏÌÓÊÂÌ˲ ‚ „ÂÌÓÏÂ. çÂÒÍÓθÍÓ ÒÎÓ‚ Ó ÒÚÛÍÚÛ ÔÓÏÓÚÓ‡ êçä-ÔÓ-ÎËÏÂ‡Á˚ III. Ç SINE, ÔÓËÁӯ‰¯Ëı ËÁ Úêçä ËÎË 7SL êçä, ÔÓÏÓÚÓ ÒÓÒÚÓËÚ ËÁ ‰‚Ûı Û˜‡ÒÚÍÓ‚ ‰ÎËÌÓÈ ÔÓ 11 Ô.Ì., ̇Á˚‚‡ÂÏ˚ı ·ÓÍÒ‡ÏË Ä Ë Ç, ÍÓÚÓ˚ ‡Á-‰ÂÎÂÌ˚ 30–35 Ô.Ì. Ç SINE, ‚ÓÁÌËͯËı ËÁ 5S êçä, ÔÓÏÓÚÓ ÒÓÒÚÓËÚ ËÁ ÚÂı ·ÓÍÒÓ‚ – Ä, IE Ë ë. ÅÓÍÒ˚ ÔÓÏÓÚÓ‡ ÓÚÌÓÒflÚÒfl Í Ì‡Ë·ÓΠÍÓÌÒÂ‚‡ÚË‚Ì˚Ï ‡ÈÓÌ‡Ï ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚. ë͇Á‡‚ Ó ÔÓÏÓÚÓÂ, ÒΉÛÂÚ ÛÔÓÏflÌÛÚ¸ Ë Ó ÚÂ-ÏË̇ÚÓ êçä-ÔÓÎËÏÂ‡Á˚ III. àÁ‚ÂÒÚÌÓ, ˜ÚÓ Ú‡Ì-ÒÍËÔˆËfl, ÓÒÛ˘ÂÒÚ‚ÎflÂχfl ˝ÚÓÈ êçä-ÔÓÎËÏÂ‡ÁÓÈ, ÚÂÏËÌËÛÂÚÒfl ̇ Û˜‡ÒÚ͇ı ËÁ ˜ÂÚ˚Âı ËÎË ·ÓΠí [19, 21]. àÌÚÂÂÒÌÓ, ˜ÚÓ ·Óθ¯ËÌÒÚ‚Ó ÒÂÏÂÈÒÚ‚ SINE Ì ËÏÂÂÚ ÒÓ·ÒÚ‚ÂÌÌÓ„Ó ÚÂÏË̇ÚÓ‡. ùÚÓ ÓÁ̇˜‡ÂÚ, ˜ÚÓ Ú‡ÌÒÍËÔˆËfl Ú‡ÍËı ˝ÎÂÏÂÌÚÓ‚ ÔÓ‰ÓÎʇÂÚÒfl Á‡ Ëı Ô‰ÂÎ˚ Ë ÓÒڇ̇‚ÎË‚‡ÂÚÒfl ̇ ÒÎÛ˜‡ÈÌ˚ı í-·ÎÓ-͇ı ‚ ÔËÎÂʇ˘Ëı Û˜‡ÒÚí-·ÎÓ-͇ı Ñçä. 3'-ÍÓ̈‚ÓÈ ‡ÈÓÌ ÔÓ‰‡‚Îfl˛˘Â„Ó ·Óθ¯ËÌÒÚ‚‡ SINE ÏÎÂÍÓÔËÚ‡˛˘Ëı Ô‰ÒÚ‡‚ÎflÂÚ ÒÓ·ÓÈ ·Ó„‡-ÚÛ˛ ÔÓÒΉӂ‡ÚÂθÌÓÒÚ¸. èÓ ÒÚÛÍÚÛ ڇÍÓ„Ó Ä-·Ó„‡ÚÓ„Ó ı‚ÓÒÚ‡ ‚Ò ÒÂÏÂÈÒÚ‚‡ ÍÓÓÚÍËı ÂÚÓÔÓÁÓ-ÌÓ‚ ÏÎÂÍÓÔËÚ‡˛˘Ëı ÏÓÊÌÓ ‡Á‰ÂÎËÚ¸ ̇ ‰‚‡ Í·ÒÒ‡ [22]. ì Í·ÒÒ‡, ‚ ÍÓÚÓ˚È ‚ıÓ‰ËÚ Ç2-˝ÎÂÏÂÌÚ „˚ÁÛ-ÌÓ‚, ‚ ÒÓÒÚ‡‚ ı‚ÓÒÚ‡ ‚ıÓ‰ËÚ ÚÂÏË̇ÚÓ Ú‡ÌÒÍËÔ-ˆËË, ‡ Ú‡ÍÊ „ÂÍÒ‡ÌÛÍÎÂÓÚˉ˚ ÄÄíÄÄÄ. í‡ÍË SINE ÓÚÌÓÒflÚÒfl Í Í·ÒÒÛ í+_{, ÚÓ„‰‡ Í‡Í SINE ·ÂÁ} ÚÂ-ÏË̇ÚÓ‡ Ë ÄÄíÄÄÄ – Í Í·ÒÒÛ í–_{. í‡ÌÒÍËÔˆËfl} B2 SINE Á‡Í‡Ì˜Ë‚‡ÂÚÒfl ̇ í-·ÎÓÍÂ, Ë Ì‡ 3'-ÍÓ̈ ӷ-‡ÁÓ‚‡‚¯ÂÈÒfl χÎÓÈ êçä ÒËÌÚÂÁËÛÂÚÒfl ÔÓÎË(Ä) [23]. ÑÎfl Ú‡ÍÓ„Ó ÔÓΡ‰ÂÌËÎËÓ‚‡ÌËfl ÌÂÓ·ıÓ‰Ëχ ÔÓÒΉӂ‡ÚÂθÌÓÒÚ¸ ÄÄíÄÄÄ. ùÚÓ ÛÌË͇θÌ˚È ÔËÏÂ ÔÓΡ‰ÂÌËÎËÓ‚‡ÌËfl Ú‡ÌÒÍËÔÚÓ‚, ÒËÌÚÂ-ÁËÓ‚‡ÌÌ˚ı êçä-ÔÓÎËÏÂ‡ÁÓÈ III. èÓÎË(Ä), ‡ÒÔÓ-ÎÓÊÂÌ̇fl ̇ ÍÓ̈ Ú‡ÌÒÍËÔÚÓ‚ Ú‡ÍËı SINE, ÒÛ˘Â-ÒÚ‚ÂÌÌ˚Ï Ó·‡ÁÓÏ Û‚Â΢˂‡ÂÚ ‚ÂÏfl ÊËÁÌË êçä ‚ ÍÎÂÚÍÂ Ë Ë„‡ÂÚ ‚‡ÊÌÛ˛ Óθ ‚ ÔÓˆÂÒÒ ‚ÓÁÌËÍÌÓ-‚ÂÌËfl Ëı ÌÓ‚˚ı ÍÓÔËÈ ‚ „ÂÌÓÏÂ. “ï‚ÓÒÚ˚” SINE Û ÂÔÚËÎËÈ Ë ˚· (Ë ÌÂÍÓÚÓ˚ SINE ÏÎÂÍÓÔËÚ‡˛˘Ëı) ÛÒÚÓÂÌ˚ ÌÂÒÍÓθÍÓ Ë̇˜Â: ÓÌË ÍÓÓ˜Â Ë ÒÓÒÚÓflÚ ËÁ ڇ̉ÂÏÌ˚ı ÔÓ‚ÚÓÓ‚ ‰ÎËÌÓÈ 3–5 Ì. (ËÒ. 2). Ö˘Â Ó‰Ì‡ ÓÒÓ·ÂÌÌÓÒÚ¸ Ú‡ÍËı SINE ÒÓ-ÒÚÓËÚ ‚ ÚÓÏ, ˜ÚÓ ÔÂ‰ ı‚ÓÒÚÓÏ ‚ ÌËı ‡ÒÔÓÎÓÊÂÌ Û˜‡ÒÚÓÍ ‰ÎËÌÓÈ 30–100 Ì., „ÓÏÓÎӄ˘Ì˚È 3'-ÍÓ̈‚Ó-ÏÛ ‡ÈÓÌÛ Ó‰ÌÓ„Ó ËÁ LINE (ËÒ. 2) [24], ̇ÔËÏÂ Bov-B, CR-1, L2, RSg-1 Ë ÌÂÍÓÚÓ˚ı ‰Û„Ëı. ìÒÚ‡-ÌÓ‚ÎÂÌÓ, ˜ÚÓ ‰‡ÌÌ˚È Û˜‡ÒÚÓÍ ‚ êçä SINE ÛÁ̇ÂÚÒfl ‚ÂÚ‡ÁÓÈ ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘Â„Ó LINE, ˜ÚÓ ÔË‚Ó‰ËÚ Í Ó·‡ÚÌÓÈ Ú‡ÌÒÍËÔˆËË ˝ÚÓÈ êçä Ë Ó·‡ÁÓ‚‡Ì˲ ÌÓ‚ÓÈ ÍÓÔËË SINE. Ä Û ÏÎÂÍÓÔËÚ‡˛˘Ëı ·Óθ¯ËÌ-ÒÚ‚Ó ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚, ÌÂÒÛ˘Ëı Ä-·Ó„‡Ú˚È ı‚ÓÒÚ, Ì ËÏÂÂÚ ‡ÈÓ̇ ÒıÓ‰ÒÚ‚‡ ÒÓ Ò‚ÓËÏ LINE-Ô‡ÚÌÂÓÏ, ‡ ËÏÂÌÌÓ L1. ë˜ËÚ‡ÂÚÒfl, ˜ÚÓ ÔÓ-ÎË(Ä) ËÎË Ä-·Ó„‡Ú˚È ı‚ÓÒÚ ‚ ÒÓÒÚ‡‚ êçä Ú‡ÍËı SINE ‚˚ÒÚÛÔ‡ÂÚ ‚ ͇˜ÂÒÚ‚Â ÔÓÒΉӂ‡ÚÂθÌÓÒÚË, ÛÁ̇‚‡ÂÏÓÈ ‚ÂÚ‡ÁÓÈ [25, 26]. èÓ ÒÓ‚ÂÏÂÌÌ˚Ï Ô‰ÒÚ‡‚ÎÂÌËflÏ Ó·‡ÁÓ‚‡ÌË ÌÓ‚˚ı ÍÓÔËÈ SINE ÔÓËÒıÓ‰ËÚ ÔÓ ÒÎÂ‰Û˛˘ÂÏÛ ÏÂı‡-ÌËÁÏÛ [27, 28]. äÓÏÔÎÂÍÒ êçä SINE Ë ‚ÂÚ‡Á˚ LINE Ú‡ÌÒÔÓÚËÛÂÚÒfl ‚ fl‰Ó, „‰Â ‚ÂÚ‡Á‡, ӷ·-‰‡˛˘‡fl Ú‡ÍÊ ˝Ì‰ÓÌÛÍ·ÁÌÓÈ ‡ÍÚË‚ÌÓÒÚ¸˛, ‡ÁÂ-Á‡ÂÚ Ó‰ÌÛ ˆÂÔ¸ „ÂÌÓÏÌÓÈ Ñçä ̇ Û˜‡ÒÚÍÂ, „ÓÏÓÎÓ-„˘ÌÓÏ ı‚ÓÒÚÛ SINE. ùÚÓ ÔË‚Ó‰ËÚ Í ÚÓÏÛ, ˜ÚÓ ı‚Ó-ÒÚÓ‚‡fl ˜‡ÒÚ¸ êçä ÒÔ‡Ë‚‡ÂÚÒfl Ò ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘ÂÈ ˆÂÔ¸˛ Ñçä, 3'-ÍÓ̈ ÍÓÚÓÓÈ ÒÎÛÊËÚ Ô‡ÈÏÂÓÏ ‚ Ó·‡ÚÌÓÈ Ú‡ÌÒÍËÔˆËË. èÓ Á‡‚Â¯ÂÌ˲ ˝ÚÓÈ ‡Í-ˆËË Ë ‡ÁÂÁ‡ÌËfl ‚ÚÓÓÈ ˆÂÔË „ÂÌÓÏÌÓÈ Ñçä ÔÓËÒ-ıÓ‰ËÚ ‰ÓÒÚÓÈ͇ ‚ÚÓÓÈ ˆÂÔË Ë Á‡ÔÓÎÌÂÌË ·¯ÂÈ ÍÎÂÚÓ˜Ì˚ÏË Ñçä-ÔÓÎËÏÂ‡Á‡ÏË. Ç ÂÁÛθڇÚ ӷ-‡ÁÛÂÚÒfl ÌÓ‚‡fl ÍÓÔËfl SINE, ÓÍÛÊÂÌ̇fl ÔflÏ˚ÏË ÔÓ‚ÚÓ‡ÏË ‰ÎËÌÓÈ 8–16 Ô.Ì. (target site duplication, TSD). ÇÚÓ‡fl ÍÓÔËfl TSD Ó·‡ÁÛÂÚÒfl ‚ ÂÁÛθڇÚ ‡ÒËÏÏÂÚ˘ÌÓ„Ó ‡ÁÂÁ‡ÌËfl „ÂÌÓÏÌÓÈ Ñçä Ë Á‡ÔÓÎ-ÌÂÌËfl ·¯ÂÈ. èÓ˜ÚË ‚Ò ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ ÓÍÛÊÂÌ˚ TSD, Ë ÓÌË ÒÎÛÊ‡Ú Û‰Ó·Ì˚ÏË Ï‡ÍÂ‡ÏË ÍÓ̈ӂ ÍÓÔËÈ SINE. çÂÒÏÓÚfl ̇ Ó„ÓÏÌÓ ˜ËÒÎÓ SINE ‚ „ÂÌÓχı, Î˯¸ ‰ËÌ˘Ì˚ Ëı ÍÓÔËË ‡ÍÚË‚Ì˚ Ë ÒÔÓÒÓ·Ì˚ Í ‡ÁÏÌÓÊÂÌ˲. èÓ‰‡‚Îfl˛˘Â ˜ËÒÎÓ Ëı ÔÓËÁ‚Ó‰Ì˚ı ̇ÍÚË‚Ì˚, ÌÓ ÓÌË ÒÓı‡Ìfl˛ÚÒfl ‚ „ÂÌÓÏÂ Ë ÔÓÒÚÂ-ÔÂÌÌÓ ‰Â„‡‰ËÛ˛Ú. ë ‰Û„ÓÈ ÒÚÓÓÌ˚, ‡ÍÚË‚ÌÓÒÚ¸ ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ Ú‡ÍÊ ÏÂÌflÂÚÒfl ‚Ó ‚ÂÏÂÌË. ç‡ÔËÏÂ, Ther-1 SINE ·˚Î ‡ÍÚË‚ÂÌ Û Ô‰ÍÓ‚ ÏÎÂÍÓÔËÚ‡˛˘Ëı, ‡ ÒÂȘ‡Ò ˜ÂÎӂ˜ÂÒÍËÈ Ther-1 ̇ÍÚË‚ÂÌ, ‚ ÚÓ ‚ÂÏfl Í‡Í Alu ÔÓ‰ÓÎʇÂÚ ‡ÁÏÌÓʇڸÒfl ‚ „ÂÌÓÏ ˜ÂÎÓ‚Â͇ [29]. ÅÓΠÚÓ„Ó, ÔÓÌflÚË “‚ÂÏÂÌË ÊËÁÌË” ÓÚÌÓÒËÚÒfl Ë Í ÔÓ‰ÒÂÏÂÈ-ÒÚ‚‡Ï ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚: Ú‡Í, Ò‡Ï˚ ÒÚ‡˚ ÔÓ‰ÒÂÏÂÈÒÚ‚‡ Alu ·ÓΠ̠‡ÁÏÌÓʇ˛ÚÒfl; ÓÌË ·˚ÎË ‡ÍÚË‚Ì˚ ‚ ‡ÁÌ˚ ÔÂËÓ‰˚ ‚ÂÏÂÌË, Ë ˜ËÒÎÓ ÍÓÔËÈ SINE, ÓÚÌÓÒfl˘ËıÒfl Í ‡ÁÌ˚Ï ÔÓ‰ÒÂÏÂÈÒÚ‚‡Ï, ‚ „ÂÌÓ-Ï ڇÍÊ Á̇˜ËÚÂθÌÓ ‡Á΢‡ÂÚÒfl [30]. àëèéãúáéÇÄçàÖ äéêéíäàï êÖíêéèéáéçéÇ Ç îàãéÉÖçÖíàóÖëäàï àëëãÖÑéÇÄçàüï ä ̇˜‡ÎÛ 90-ı „Ó‰Ó‚ ÔÓ¯ÎÓ„Ó ‚Â͇ ¯ËÓÍÓ ‡Ò-ÔÓÒÚ‡ÌÂÌË ÔÓÎÛ˜ËÎË ‡·ÓÚ˚ ÔÓ ÔÓÒÚÓÂÌ˲ ÙË-ÎÓ„ÂÌÂÚ˘ÂÒÍËı ‰Â‚¸Â‚ ̇ ÓÒÌÓ‚Â ÌÛÍÎÂÓÚˉÌ˚ı

ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ „ÂÌÓ‚ ËÎË ‡ÏËÌÓÍËÒÎÓÚÌ˚ı ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ ÍÓ‰ËÛÂÏ˚ı ËÏË ·ÂÎÍÓ‚. Ç ˝ÚËı ‡·ÓÚ‡ı ËÒÔÓθÁÓ‚‡ÎË „·‚Ì˚Ï Ó·‡ÁÓÏ ÏËÚÓ-ıÓ̉ˇθÌÛ˛ Ñçä. ç‡‰Ó Ò͇Á‡Ú¸, ˜ÚÓ ˜‡ÒÚÓ ˝ÚË ËÒ-ÒΉӂ‡ÌËfl ÔË‚Ó‰ËÎË Í Ô‡‡‰ÓÍ҇θÌ˚Ï ‚˚‚Ó‰‡Ï. ç‡ÔËÏÂ, Òӄ·ÒÌÓ Ó‰ÌÓÈ ËÁ ÔÛ·ÎË͇ˆËÈ ‚ ÊÛ̇Π“Nature”, ÏÓÒ͇fl Ò‚ËÌ͇ Ì ÏÓÊÂÚ ·˚Ú¸ ÓÚÌÂÒÂ̇ Í „˚ÁÛÌ‡Ï [31]. Ç ÔÓÒΉÌË „Ó‰˚ Ò ÔÂÂÌÓÒÓÏ ‡ÍˆÂÌ-Ú‡ ̇ fl‰ÂÌ˚ „ÂÌ˚, Û‚Â΢ÂÌËÂÏ ˜ËÒ· ‡Ì‡ÎËÁËÛÂ-Ï˚ı Ó„‡ÌËÁÏÓ‚ Ë, ̇ÍÓ̈, Ò ‡Á‚ËÚËÂÏ Ï‡ÚÂχÚË-˜ÂÒÍËı ÏÂÚÓ‰Ó‚ ÔÓÒÚÓÂÌËfl ÙËÎÓ„ÂÌÂÚËχÚÂχÚË-˜ÂÒÍËı ‰ÂÂ-‚¸Â‚ ‰ÓÒÚË„ÌÛÚÓ Á̇˜ËÚÂθÌÓ ÎÛ˜¯Â ÒÓÓÚ‚ÂÚÒÚ‚Ë ÂÁÛθڇÚÓ‚ ‡ÁÌ˚ı ‡·ÓÚ. é‰Ì‡ÍÓ ÒΉÛÂÚ Ò͇Á‡Ú¸, ˜ÚÓ ÏÌÓÊÂÒÚ‚Ó ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ‚˚‚Ó‰Ó‚, ҉·Ì-Ì˚ı Ò ÔÓÏÓ˘¸˛ ÏÓÎÂÍÛÎfl҉·Ì-Ì˚ı ÏÂÚÓ‰Ó‚, Ì Òӄ·ÒÛ-˛ÚÒfl Ò ÔÓÒÚÓÂÌËflÏË, ÓÒÌÓ‚‡ÌÌ˚ÏË Ì‡ ËÁÛ˜ÂÌËË ÏÓÙÓÎӄ˘ÂÒÍËı ÔËÁ̇ÍÓ‚. ÑÛ„ËÏË ÒÎÓ‚‡ÏË, ‚ ÔÓÒΉÌË „Ó‰˚ ÏÓÎÂÍÛÎflÌ˚ ÒËÒÚÂχÚËÍË ÒÛ˘Â-ÒÚ‚ÂÌÌÓ ÔÂÂÒÏÓÚÂÎË Ú‡‰ËˆËÓÌÌ˚ ‚Á„Îfl‰˚ ̇ ÙËÎÓ„ÂÌ˲ ÚÂı ËÎË ËÌ˚ı „ÛÔÔ Ó„‡ÌËÁÏÓ‚ [32–34]. à ‚Ò Ê ˝‚ÓβˆËfl „Â̇ Ë Ó„‡ÌËÁχ ÏÓ„ÛÚ Ì ÒÓ‚Ô‡-‰‡Ú¸, Ë ÙËÎÓ„ÂÌÂÚ˘ÂÒÍË ÔÓÒÚÓÂÌËfl, ÓÒÌÓ‚‡ÌÌ˚ ̇ ‡Ì‡ÎËÁ „ÂÌÓ‚, ÏÓ„ÛÚ ·˚Ú¸ ӯ˷ӘÌ˚ÏË. ê¯ËÚ¸ ˝ÚÛ ÔÓ·ÎÂÏÛ ÏÓÊÌÓ Á̇˜ËÚÂθÌÓ Û‚Â΢˂ ˜ËÒÎÓ ‡Ì‡ÎËÁËÛÂÏ˚ı „ÂÌÓ‚. é‰Ì‡ÍÓ ‰Îfl ÏÌÓ„Ëı ‚ˉӂ ÔÓ-‰Ó·Ì˚ ‰‡ÌÌ˚ Ì‰ÓÒÚÛÔÌ˚, ‡ Ëı ÔÓÎÛ˜ÂÌË ÓÒÚ‡ÂÚÒfl ‚ÂҸχ ÚÛ‰ÓÂÏÍËÏ. ùÚÓ ‰Â·ÂÚ ‚ÓÒÚ·ӂ‡ÌÌ˚ÏË ‰Û„Ë ÏÂÚÓ‰˚, ËÒÔÓθÁÛ˛˘Ë ÌÂÁ‡‚ËÒËÏ˚ ËÒÚÓ˜-ÌËÍË ÙËÎÓ„ÂÌÂÚ˘ÂÒÍÓÈ ËÌÙÓχˆËË. åÂÚÓ‰ ÒÂÏÂÈÒÚ‚ ֢ ‚ ̇˜‡Î 90-ı Ï˚ Ô‰ÎÓÊËÎË ËÒÔÓθÁÓ‚‡Ú¸ SINE Í‡Í ËÒÚÓ˜ÌËÍ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍÓÈ ËÌÙÓχˆËË [35–37]. 燯 ÔÓ‰ıÓ‰ ÒÓÒÚÓflÎ ‚ ÚÓÏ, ˜ÚÓ ÌÛÊÌÓ ÛÒÚ‡-ÌÓ‚ËÚ¸, „ÂÌÓÏ˚ ͇ÍËı ËÒÒΉÛÂÏ˚ı ‚ˉӂ ÒÓ‰ÂÊ‡Ú ‰‡ÌÌÓ ÒÂÏÂÈÒÚ‚Ó SINE, ‡ Û Í‡ÍËı ‚ˉӂ Â„Ó ÌÂÚ. ùÚÓ ÏÓÊÌÓ Ò‰Â·ڸ Ò ÔÓÏÓ˘¸˛ ‰ÓÚ-„Ë·ˉËÁ‡ˆËË Ë ÔÓ-ÎËÏÂ‡ÁÌÓÈ ˆÂÔÌÓÈ ‡͈ËË (èñê). ÇÒ ‚ˉ˚, ÌÂÒÛ-˘Ë ‚ Ò‚ÓËı „ÂÌÓχı SINE ‰‡ÌÌÓ„Ó ÒÂÏÂÈÒÚ‚‡, ‡Ò-ÒχÚË‚‡˛ÚÒfl Í‡Í ·ÓΠ·ÎËÁÍÓ Ó‰ÒÚ‚ÂÌÌ˚ ‰Û„ ‰Û„Û, ˜ÂÏ ‚ˉ‡Ï, ‚ ÍÓÚÓ˚ı ˝ÚÓ ÒÂÏÂÈÒÚ‚Ó ÓÚÒÛÚ-ÒÚ‚ÛÂÚ. ç‡ ËÒ. 3 ‚ˉ˚ 1, 2 Ë 3 ÒÓ‰ÂÊ‡Ú ‚ Ò‚ÓËı „ÂÌÓ-χı SINE ÒÂÏÂÈÒÚ‚‡ Ä, ‡ ‚ˉ 4 Â„Ó Ì ÒÓ‰ÂÊËÚ. Ç ÚÓ Ê ‚ÂÏfl, SINE ÒÂÏÂÈÒÚ‚‡ Å ÂÒÚ¸ Û ‚ˉӂ 1 Ë 2, ÌÓ ÌÂ Û ‚ˉӂ 3 Ë 4. ùÚÓ ÔÓÁ‚ÓÎflÂÚ ÔÓÒÚÓËÚ¸ ‰‚Ó, ËÁÓ·-‡ÊÂÌÌÓ ̇ ËÒ. 3. àÒÔÓθÁÓ‚‡ÌË ˝ÚÓ„Ó ÔÓ‰ıÓ‰‡ Ú·ÛÂÚ, ˜ÚÓ·˚ ‚ „ÂÌÓχı Ó„‡ÌËÁÏÓ‚ ËÁÛ˜‡ÂÏÓÈ Ú‡ÍÒÓÌÓÏ˘ÂÒÍÓÈ „ÛÔÔ˚ ·˚ÎË Ë‰ÂÌÚËÙˈËÓ‚‡Ì˚ ͇ÍËÂ-ÎË·Ó ÍÓ-ÓÚÍË ÂÚÓÔÓÁÓÌ˚. ÖÒÎË ÓÌË Ì ·˚ÎË Óı‡‡ÍÚÂË-ÁÓ‚‡Ì˚ ‡ÌÂÂ, ÚÓ ÏÓÊÌÓ ‚ÓÒÔÓθÁÓ‚‡Ú¸Òfl ÒÎÂ‰Û˛-˘ËÏ Ò‡‚ÌËÚÂθÌÓ ÛÌË‚Â҇θÌ˚Ï ÏÂÚÓ‰ÓÏ, Ô‰-ÎÓÊÂÌÌ˚Ï Ì‡ÏË ‚ 1999 „. [38]. ë̇˜‡Î‡ ÔÓ‚Ó‰flÚ èñê Ò ËÒÔÓθÁÓ‚‡ÌËÂÏ Ó˜Â̸ Ì·Óθ¯Ëı ÍÓ΢ÂÒÚ‚ „ÂÌÓÏÌÓÈ Ñçä Ë ‚˚ÓʉÂÌÌ˚ı Ô‡ÈÏÂÓ‚, ÍÓÏÔÎÂ-ÏÂÌÚ‡Ì˚ı ·ÓÍÒ‡Ï Ä Ë Ç ÔÓÏÓÚÓ‡ êçä-ÔÓÎËÏÂ-‡Á˚ III. Ç ÂÁÛθڇÚ ̇ ÏÌÓ„Ó˜ËÒÎÂÌÌ˚ı ÍÓÔËflı SINE, Ô‰ÔÓÎÓÊËÚÂθÌÓ Ô‰ÒÚ‡‚ÎÂÌÌ˚ı ‚ ‰‡ÌÌÓÏ „ÂÌÓÏÂ, Ó·‡ÁÛÂÚÒfl èñê-ÔÓ‰ÛÍÚ, ÍÓÚÓ˚È ÏÓÊÌÓ ËÒÔÓθÁÓ‚‡Ú¸ ‚ ͇˜ÂÒÚ‚Â „Ë·ˉËÁ‡ˆËÓÌÌÓ„Ó ÁÓ̉‡ ÔË ÒÍËÌËÌ„Â „ÂÌÓÏÌÓÈ ·Ë·ÎËÓÚÂÍË ‰‡ÌÌÓ„Ó ËÎË Ó‰ÒÚ‚ÂÌÌÓ„Ó ‚ˉ‡. ëÂÍ‚ÂÌËÓ‚‡ÌË ÔÓÎÛ˜ÂÌÌ˚ı ÍÎÓÌÓ‚ Ë ‡Ì‡ÎËÁ Ëı ÌÛÍÎÂÓÚˉÌ˚ı ÔÓÒΉӂ‡ÚÂθÌÓ-ÒÚÂÈ ÔÓÁ‚ÓÎflÂÚ Ë‰ÂÌÚËÙˈËÓ‚‡Ú¸ ÌÓ‚˚ ÒÂÏÂÈÒÚ‚‡ SINE. ç‡ÍÓ̈, Á̇fl ÍÓÌÒÂÌÒÛÒ ÌÛÍÎÂÓÚˉÌÓÈ ÔÓÒÎÂ-‰Ó‚‡ÚÂθÌÓÒÚË SINE, ÏÓÊÌÓ ‚˚·‡Ú¸ ÒÔˆËÙ˘Â-ÒÍË ‰Îfl ˝ÚÓÈ ÔÓÒΉӂ‡ÚÂθÌÓÒÚË Ô‡ÈÏÂ˚ Ë ÔÓ-LINE OPC1 OPC2 PçK LINE í‡ÌÒÍËÔˆËfl êçä-ÔÓÎËÏÂ‡ÁÓÈ II í‡ÌÒÎflˆËfl é·‡Ú̇fl Ú‡ÌÒÍËÔÚ‡Á‡ LINE Ñçä ÄÍÚË‚Ì˚È SINE çÓ‚‡fl ÍÓÔËfl SINE é·‡Ú̇fl Ú‡ÌÒÍËÔˆËfl Ë ËÌÚ„‡ˆËfl í‡ÌÒÍËÔˆËfl êçä-ÔÓÎËÏÂ‡ÁÓÈ III êËÒ. 2. ëıÂχ ÂÔÎË͇ˆËË ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚. éêë – ÓÚÍ˚Ú‡fl ‡Ï͇ Ò˜ËÚ˚‚‡ÌËfl („ÂÌ); ‰Û„Ë ÔÓflÒÌÂÌËfl ‚ ÚÂÍÒÚÂ.

åéãÖäìãüêçÄü ÅàéãéÉàü      ÚÓÏ 43      ‹ 5      2009 ‚ÂÒÚË èñê Ò „ÂÌÓÏÌ˚ÏË Ñçä ˆÂÎÓ„Ó fl‰‡ ‚ˉӂ. èÓ‰Ó·Ì˚È ‡Ì‡ÎËÁ ÏÓÊÂÚ ·˚Ú¸ ÔÓ‰ÂÎ‡Ì Ë ÍÓÏÔ¸˛-ÚÂÌ˚ÏË ÏÂÚÓ‰‡ÏË, ÂÒÎË ‚ ·‡ÌÍË ‰‡ÌÌ˚ı ‰ÂÔÓÌËÓ-‚‡ÌÓ ‰ÓÒÚ‡ÚÓ˜ÌÓ ˜ËÒÎÓ ÌÛÍÎÂÓÚˉÌ˚ı ÔÓÒΉӂ‡-ÚÂθÌÓÒÚÂÈ ËÁÛ˜‡ÂÏ˚ı Ú‡ÍÒÓÌÓ‚. êËÒ. 4 ËÎβÒÚËÛÂÚ ÔËÏÂÌÂÌË ˝ÚÓ„Ó ÏÂÚÓ‰‡ ‰Îfl ÓÚfl‰‡ „˚ÁÛÌÓ‚. ç‡ ˝ÚÓÏ “Í·ÒÒ˘ÂÒÍÓÏ” ‰‚Â, ‚ÁflÚÓÏ ËÁ ÏÓÌÓ„‡ÙËË Romer [39], ÔÓ͇Á‡-ÌÓ ‡ÒÔ‰ÂÎÂÌË ÌÂÒÍÓθÍËı ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ Û „˚ÁÛÌÓ‚. ê‡ÒÔ‰ÂÎÂÌË ӉÌÓ„Ó ËÁ ÌËı, ̇Á‚‡ÌÌÓ„Ó Ì‡ÏË B1-dID, Ì ÔÓÎÌÓÒÚ¸˛ ÒÓ-ÓÚ‚ÂÚÒÚ‚Ó‚‡ÎÓ Ô‰ÒÚ‡‚ÎÂÌËflÏ Ó ÙËÎÓ„ÂÌËË „˚-ÁÛÌÓ‚. B1-dID ̇ȉÂÌ Û Ô‰ÒÚ‡‚ËÚÂÎÂÈ ÚÂı ÒÂ-ÏÂÈÒÚ‚: ·ÂÎÓÍ (Sciuridae), ‡ÔÎÓ‰ÓÌÚËÈ (Aplodon-tidae) Ë ÒÓ̸ (Gliridae). ç‡ ‰‚ Romer ÒÓÌË ‡ÒÔÓ·„‡ÎËÒ¸ ‚ ËÌÓÏ ÏÂÒÚ – ÒÂ‰Ë ÏËÓÏÓÙÌ˚ı (Ï˚¯ÂÔÓ‰Ó·Ì˚ı) „˚ÁÛÌÓ‚ (̇ ËÒ. 4 ÓÚϘÂÌÓ Á‚ÂÁ‰Ó˜ÍÓÈ). àÁÛ˜ÂÌË ‡ÒÔ‰ÂÎÂÌËfl B1-dID ÔÓÁ‚ÓÎËÎÓ ÔÓÏÂÒÚËÚ¸ ÒÓ̸ ‚ ‰Û„Ó ÏÂÒÚÓ Ì‡ ‰Â-‚ fl‰ÓÏ Ò ·ÂÎ͇ÏË Ë ‡ÔÎÓ‰ÓÌÚËflÏË [40]. èÓÁ‰Ì Ó‰ÒÚ‚Ó ˝ÚËı ÚÂı ÒÂÏÂÈÒÚ‚ „˚ÁÛÌÓ‚ ÔÓ‰Ú‚Â‰Ë-ÎË ‰Û„Ë ‡‚ÚÓ˚ ÔË ‡Ì‡ÔÓ‰Ú‚Â‰Ë-ÎËÁ fl‰ÂÌ˚ı „ÂÌÓ‚ [41]. Ç ‰Û„ÓÏ ÒÎÛ˜‡Â ‚Ò ̇˜‡ÎÓÒ¸ Ò ÙËÎÓ„ÂÌÂÚ˘Â-ÒÍÓ„Ó ‡Ì‡ÎËÁ‡ „ÂÌÓ‚, ‚˚‰ÂÎË‚¯Â„Ó ÌÓ‚˚È Ì‡‰ÓÚfl‰ ÏÎÂÍÓÔËÚ‡˛˘Ëı Afrotheria [33]. äÓÏ ıÓ·ÓÚÌ˚ı, ‰‡Ï‡ÌÓ‚ Ë ÒËÂÌ, ‚ ÌÂ„Ó ‚ıÓ‰flÚ ÚÛ·ÍÓÁÛ·˚, Ô˚„ÛÌ-˜ËÍË Ë ÚÂÌÂÍË ÒÓ Á·ÚÓÍÓÚ‡ÏË, ÍÓÚÓ˚ı ‡Ì ÌË-ÍÓ„‰‡ Ì ӷ˙‰ËÌflÎË. ÉÛÔÔ‡ flÔÓÌÒÍËı ËÒÒΉӂ‡ÚÂ-ÎÂÈ Ó·Ì‡ÛÊË· ‚ „ÂÌÓχı ‚ÒÂı ˝ÚËı ÏÎÂÍÓÔËÚ‡˛-˘Ëı ÒÂÏÂÈÒÚ‚Ó ÍÓÓÚÍÓ„Ó ÂÚÓÔÓÁÓ̇ AfroSINE [42]. ùÚÓ ÔÓÒÎÛÊËÎÓ Ì‡‰ÂÊÌ˚Ï ÔÓ‰Ú‚ÂʉÂÌËÂÏ Ô‡‚ËθÌÓÒÚË ‚˚‰ÂÎÂÌËfl Í·‰˚ Afrotheria, Í ÍÓÚÓ-ÓÈ ÒËÒÚÂχÚËÍË-ÏÓÙÓÎÓ„Ë ÓÚÌÓÒËÎËÒ¸ Ò ·Óθ¯ËÏ ÒÍÂÔÒËÒÓÏ. ç‡ÍÓ̈, ÏÓÊÌÓ ÛÔÓÏflÌÛÚ¸ ¢ ÌÓ‚˚È Ì‡‰ÓÚfl‰ Supraprimates (ËÎË Euarchontoglires) [33]. Ö„Ó ÒÛ˘Â-1 2 3 4 Çˉ˚ ÑÓÚ-„Ë·ˉËÁ‡ˆËfl èÓÎËÏÂ‡Á̇fl ˆÂÔ̇fl ‡͈Ëfl ëÂÏÂÈÒÚ‚Ó SINE Çˉ˚ áÓ̉ Ä áÓ̉ Å è‡ÈÏÂ˚ Ä Ë Ä' è‡ÈÏÂ˚ Å Ë Å' Ä Å Å Ä 1 2 3 4 êËÒ. 3. àÒÔÓθÁÓ‚‡ÌË ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ‚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ËÒÒΉӂ‡ÌËflı. åÂÚÓ‰ ÒÂÏÂÈÒÚ‚. áÓ̉˚ Ä Ë Å Ë Ô‡ÈÏÂ˚ Ä/Ä' Ë Å/Å' ÒÔˆËÙ˘Ì˚ ‰Îfl ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ Ä Ë Å ÒÓÓÚ‚ÂÚÒÚ‚ÂÌÌÓ. ëÚÂÎ͇ÏË Ì‡ ‰‚ ÔÓ͇Á‡Ì˚ ÏÓÏÂÌÚ˚ ÔÓfl‚ÎÂÌËfl ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚.

B1 ID B4 4.5SH 4.5S1 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, _4.5S1 B4, 4.5SH IDL-Geo B1-dID êËÒ. 4. ù‚ÓβˆËÓÌÌÓ ‰Â‚Ó „˚ÁÛÌÓ‚. èβÒ˚ Ë ÏËÌÛÒ˚ – ÔËÒÛÚÒÚ‚Ë ËÎË ÓÚÒÛÚÒÚ‚Ë ÚÓ„Ó ËÎË ËÌÓ„Ó ÍÓÓÚÍÓ„Ó ÂÚÓÔÓÁÓ̇ ‚ „ÂÌÓχı Ô‰ÒÚ ‡‚ËÚÂÎÂÈ ÒÓÓÚ-‚ÂÚÒÚ‚Û˛˘Ëı ÒÂÏÂÈÒÚ‚ „˚ÁÛÌÓ‚ [37, 40, 46, 72–74]. ëÚÂÎ͇ÏË Ì‡ ‰‚ ÔÓ͇Á‡Ì˚ ÏÓÏÂÌÚ˚ ÔÓfl‚ÎÂÌËfl ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ . á‚ÂÁ‰Ó˜ÍÓÈ ÓÚϘÂÌÓ ÏÂÒÚÓ ÓÚ‚ÂÚ‚ÎÂÌËfl Í·‰˚ Gliridae ÔÓ Romer [39].

åéãÖäìãüêçÄü ÅàéãéÉàü      ÚÓÏ 43      ‹ 5      2009 ÒÚ‚Ó‚‡ÌË ÔÓ‰Ú‚ÂʉÂÌÓ ÌÂÁ‡‚ËÒËÏÓ ‚ ‡·ÓÚ‡ı flÔÓÌÒÍËı ËÒÒΉӂ‡ÚÂÎÂÈ Ë Ì‡¯ÂÈ Î‡·Ó‡ÚÓËË [43, 44]. é͇Á‡ÎÓÒ¸, ˜ÚÓ ÒÂ‰Ë ‚ÒÂı ÏÎÂÍÓÔËÚ‡˛˘Ëı ÚÓθÍÓ Ô‰ÒÚ‡‚ËÚÂÎË ˝ÚÓ„Ó Ì‡‰ÓÚfl‰‡, ‡ ËÏÂÌÌÓ „˚ÁÛÌ˚, ÔËχÚ˚ Ë ÚÛÔ‡Ë, ÒÓ‰ÂÊ‡Ú ‚ „ÂÌÓÏ ÍÓ-ÓÚÍË ÂÚÓÔÓÁÓÌ˚, ÔÓËÁӯ‰¯Ë ÓÚ 7SL êçä. å˚, Í‡Í Ë ‰Û„Ë ‡‚ÚÓ˚ [45], ÔÓ·„‡ÂÏ, ˜ÚÓ Û Ó·˘Â-„Ó Ô‰͇ ˝ÚÓÈ „ÛÔÔ˚ ÏÎÂÍÓÔËÚ‡˛˘Ëı, ·Î‡Ó·˘Â-„Ó‰‡fl ÔÓÚflÊÂÌÌÓÈ ‚ÌÛÚÂÌÌÂÈ ‰ÂΈËË, ËÁ ÔÓÒΉӂ‡-ÚÂθÌÓÒÚË 7SL êçä ‚ÓÁÌËÍ ÂÚÓ˝ÎÂÏÂÌÚ, ÍÓÚÓ˚È ˝‚ÓβˆËÓÌËÓ‚‡Î ‚ Ó˜Â̸ ÛÒÔ¯Ì˚ SINE – Ç1 Û „˚ÁÛÌÓ‚ Ë Alu Û ÔËχÚÓ‚. Ç Í‡˜ÂÒÚ‚Â ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı χÍÂÓ‚ ÏÓÊÌÓ ËÒÔÓθÁÓ‚‡Ú¸ Ì ÚÓθÍÓ ÒÂÏÂÈÒÚ‚‡ SINE, ÌÓ Ë Ëı ÔÓ‰-ÒÂÏÂÈÒÚ‚‡. í‡Í, Ï˚ ÍÎÓÌËÓ‚‡ÎË Ë ÒÂÍ‚ÂÌËÓ‚‡ÎË ÍÓÔËË Ç1 ËÁ „ÂÌÓÏÓ‚ Ô‰ÒÚ‡‚ËÚÂÎÂÈ ·Óθ¯ËÌÒÚ‚‡ ÒÂÏÂÈÒÚ‚ „˚ÁÛÌÓ‚. äÓÌÒÂÌÒÛÒÌ˚ ÔÓÒΉӂ‡ÚÂθ-ÌÓÒÚË Ç1 Û „˚ÁÛÌÓ‚ ‡ÁÌ˚ı ÒÂÏÂÈÒÚ‚ ÓÚ΢‡˛ÚÒfl ‰Û„ ÓÚ ‰Û„‡ Á‡ÏÂ̇ÏË ÌÛÍÎÂÓÚˉӂ, ‰ÂΈËflÏË Ë ‰ÛÔÎË͇ˆËflÏË. Ä̇ÎËÁ ‡ÒÔ‰ÂÎÂÌËfl ˝ÚËı ÔËÁ̇-ÍÓ‚ ÔÓ‰Ú‚Âʉ‡ÂÚ Ó‰ÌÛ ËÁ ÒÓ‚ÂÏÂÌÌ˚ı ‚ÂÒËÈ ÙË-ÎÓ„ÂÌÂÚ˘ÂÒÍÓ„Ó ‰‚‡ „˚ÁÛÌÓ‚ [46]. åÂÚÓ‰ ÍÓÔËÈ ÅÓΠ‡ÒÔÓÒÚ‡ÌÂÌÌ˚È Ë ‡Á‡·ÓÚ‡ÌÌ˚È ÔÓ‰-ıÓ‰ Í ËÁÛ˜ÂÌ˲ ÙËÎÓ„ÂÌËË, ÓÒÌÓ‚‡ÌÌ˚È Ì‡ ËÒÔÓθ-ÁÓ‚‡ÌËË ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚, ÒÓÒÚÓËÚ ‚ ÔÓ‚ÂÍ ÔËÒÛÚÒÚ‚Ëfl ÍÓÔËË SINE ‚ ÍÓÌÍÂÚÌÓÏ ÎÓÍÛÒ „ÂÌÓ-χ Û ‡ÁÌ˚ı ‚ˉӂ [47]. ùÚÓ ÏÓÊÌÓ Ò‰Â·ڸ Ò ÔÓÏÓ-˘¸˛ èñê Ò ËÒÔÓθÁÓ‚‡ÌËÂÏ Ô‡ÈÏÂÓ‚, ÍÓÏÔÎÂÏÂÌ-Ú‡Ì˚ı ÌÛÍÎÂÓÚˉÌ˚Ï ÔÓÒΉӂ‡ÚÂθÌÓÒÚflÏ, ÎÂʇ-˘ËÏ Á‡ Ô‰Â·ÏË ‰‡ÌÌÓÈ ÍÓÔËË SINE (ËÒ. 5). ÄÏÔÎËÙˈËÓ‚‡ÌÌ˚È Ù‡„ÏÂÌÚ Ñçä ·Û‰ÂÚ ÍÓÓ˜Â, ÂÒÎË ÓÌ Ì ÒÓ‰ÂÊËÚ SINE. ÑÎfl ÔÓÒÚÓÂÌËfl ‰‚‡ ÌÂ-Ó·ıÓ‰ËÏÓ ËÒÒΉӂ‡Ú¸ ÔÓ Í‡ÈÌÂÈ ÏÂ ÌÂÒÍÓθÍÓ ÎÓÍÛÒÓ‚, ÌÂÒÛ˘Ëı SINE. Çˉ˚, ÒÓ‰Âʇ˘Ë ÍÓÔ˲ SINE ‚ ‰‡ÌÌÓÏ ÎÓÍÛÒÂ, ÙËÎÓ„ÂÌÂÚ˘ÂÒÍË ·ÎËÊ ‰Û„ Í ‰Û„Û, ˜ÂÏ Í ‚ˉ‡Ï, Û ÍÓÚÓ˚ı ÌÂÚ ˝ÚÓÈ ÍÓÔËË. ÖÒÎË ‰ÓÒÚÛÔÌ˚ ÔÓÎÌ˚ ËÎË ˜‡ÒÚ˘Ì˚ ÌÛÍÎÂÓÚˉÌ˚ ÔÓ-ÒΉӂ‡ÚÂθÌÓÒÚË „ÂÌÓÏÓ‚, ÚÓ Ú‡ÍÓÈ ‡Ì‡ÎËÁ ÏÓÊÌÓ ÔÓ‚ÂÒÚË Ò ÔÓÏÓ˘¸˛ ÚÓθÍÓ ÍÓÏÔ¸˛ÚÂÌ˚ı ÏÂÚÓ-‰Ó‚ ËÎË Ò ËÒÔÓθÁÓ‚‡ÌËÂÏ ÍÓÏ·Ë̇ˆËË ÍÓÏÔ¸˛ÚÂ-Ì˚ı Ë ÏÓÎÂÍÛÎflÌÓ-·ËÓÎӄ˘ÂÒÍËı ÏÂÚÓ‰Ó‚. ùÚÓÚ ÏÂÚÓ‰ ‚ÔÂ‚˚ ÔËÏÂÌËÎË ‰Îfl ËÁÛ˜ÂÌËfl ÙË-ÎÓ„ÂÌËË ÔÓÔÛÎflˆËÓÌÌ˚ı „ÛÔÔ ˜ÂÎÓ‚Â͇ [48] Ë ÎÓÒÓ-Ò‚˚ı [49]. Ç ‰‡Î¸ÌÂȯÂÏ ÔÓÎËÏÓÙËÁÏ (̇ÎË-˜ËÂ/ÓÚÒÛÚÒÚ‚ËÂ) ÓÚ‰ÂθÌ˚ı ÍÓÔËÈ SINE Alu ¯ËÓÍÓ ËÒÔÓθÁÓ‚‡ÎË ‰Îfl ËÁÛ˜ÂÌËfl ÙËÎÓ„ÂÌËË ÔËχÚÓ‚, ÔÓËÒıÓʉÂÌËfl ˜ÂÎÓ‚Â͇ Ë ‚ÓÔÓÒÓ‚ ‰ÂÏÓ„‡ÙËË (ÒÏ. Ó·ÁÓ [30] Ë ÒÒ˚ÎÍË ‚ ÌÂÏ). ÇÔ˜‡ÚÎfl˛˘ËÏ Ë Û‰Ë‚ËÚÂθÌ˚Ï ÓÚÍ˚ÚËÂÏ, ҉·ÌÌ˚Ï Ò ÔÓÏÓ˘¸˛ ‡ÒÒÏÓÚÂÌÌÓ„Ó ÔÓ‰ıÓ‰‡, ·˚ÎÓ ÚÓ, ˜ÚÓ ÓÚfl‰ ÍËÚÓ-Ó·‡ÁÌ˚ı (Cetacea) Ô‰ÒÚ‡‚ÎflÂÚ ÒÓ·ÓÈ ‚ÌÛÚÂÌÌ˛˛ ‚ÂÚ‚¸ ‰‚‡ ÓÚfl‰‡ Ô‡ÌÓÍÓÔ˚ÚÌ˚ı (Artiodactyla). í‡ÍËÏ Ó·‡ÁÓÏ, „ËÔÔÓÔÓÚ‡Ï˚ Ó͇Á˚‚‡˛ÚÒfl ·ÎËÊÂ Í ÍËÚ‡Ï, ˜ÂÏ Í Ê‚‡˜Ì˚Ï ËÎË Ò‚Ë̸flÏ [50, 51]. èÓÁ‰Ì ڇÍÛ˛ ÙËÎÓ„ÂÌ˲ Ô‡ÌÓÍÓÔ˚ÚÌ˚ı Ë ÍËÚÓÓ·‡ÁÌ˚ı ÔÓ‰Ú‚Â‰ËÎË Ë ‰Û„ËÏË ·ÓΠÚ‡‰ËˆËÓÌÌ˚ÏË ÏÓÎÂ-ÍÛÎflÌ˚ÏË ÏÂÚÓ‰‡ÏË [32]. àÌÚÂ-SINE-èñê ëÛ˘ÂÒÚ‚ÛÂÚ Â˘Â Ó‰ËÌ ÏÂÚÓ‰ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎËÁ‡, ‚ ÍÓÚÓÓÏ ËÒÔÓθÁÛ˛ÚÒfl ÍÓÓÚÍË ÂÚÓ-ÔÓÁÓÌ˚. ùÚÓÚ ÏÂÚÓ‰, ̇Á˚‚‡ÂÏ˚È ËÌÚÂ- SINE-èñê, Á‡Íβ˜‡ÂÚÒfl ‚ SINE-èñê, ÔÓ‚Ó‰ËÏÓÈ Ì‡ „ÂÌÓÏ-ÌÓÈ Ñçä ‚ ͇˜ÂÒڂ χÚˈ˚ Ò Ï˜ÂÌ˚ÏË Ô‡ÈÏÂ-‡ÏË, ÍÓÏÔÎÂÏÂÌÚ‡Ì˚ÏË Û˜‡ÒÚÍ‡Ï SINE. è‡ÈÏÂ-˚ ÔÓ‰·Ë‡˛Ú Ú‡ÍËÏ Ó·‡ÁÓÏ, ˜ÚÓ·˚ ‡ÏÔÎËÙˈË-Ó‚‡Ú¸ Û˜‡ÒÚÍË „ÂÌÓχ, ‡ÒÔÓÎÓÊÂÌÌ˚ ÏÂÊ‰Û ÒÓÒ‰ÌËÏË ÍÓÔËflÏË SINE (ËÒ. 6). Ç ÂÁÛθڇÚ ‡Á-‰ÂÎÂÌËfl ÔÓ‰ÛÍÚÓ‚ èñê Ó·‡ÁÛÂÚÒfl ̇·Ó Ó·˘Ëı Ë ÛÌË͇θÌ˚ı ÔÓÎÓÒ, Ë Ëı ÍÓÏÔ¸˛ÚÂÌ˚È ‡Ì‡ÎËÁ ÔÓÁ-‚ÓÎflÂÚ ÒÚÓËÚ¸ ˝‚ÓβˆËÓÌÌ˚ ‰Â‚¸fl. ùÚÓÚ ÏÂÚÓ‰ ·˚Î ‚ÔÂ‚˚ ËÒÔÓθÁÓ‚‡Ì ‰Îfl ‡Ì‡ÎËÁ‡ ˝‚ÓβˆËË Ô‡ÌÓÍÓÔ˚ÚÌ˚ı [52]. ë ÚÂı ÔÓ Â„Ó Ò ÛÒÔÂıÓÏ ÔË-ÏÂÌflÎË ‚ ÙËÎÓ„ÂÌËË ÛÍÓÍ˚Î˚ı [53], ̇ÒÂÍÓÏÓfl‰-Ì˚ı [54–56], fl˘Âˈ [57] Ë ‚˚Ò¯Ëı ÔËχÚÓ‚ [58]. ÑÓÒÚÓËÌÒÚ‚‡ Ë ÔÓ·ÎÂÏ˚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ÏÂÚÓ‰Ó‚, ËÒÔÓθÁÛ˛˘Ëı ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ ïÓÚfl ÓÚ‰ÂθÌ˚ ÍÓÔËË ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ÏÓ„ÛÚ ·˚Ú¸ ‚‡ÊÌ˚ ‰Îfl ‡·ÓÚ˚ „ÂÌÓ‚, ÔÓ‰‡‚Îfl˛˘Â Ëı ·Óθ¯ËÌÒÚ‚Ó ÌÂÙÛÌ͈ËÓ̇θÌÓ, Ë ÓÌË ÔÓ ÒÛÚË Ô‰ÒÚ‡‚Îfl˛Ú ÒÓ·ÓÈ “„ÂÌÓÏÌ˚ı Ô‡‡ÁËÚÓ‚”. í‡ÍËÏ Ó·‡ÁÓÏ, ·Û‰Û˜Ë ÌÂÈÚ‡Î¸Ì˚ÏË ÔËÁ͇̇ÏË, ËÁ ‚ÒÂı ˝‚ÓβˆËÓÌÌ˚ı ÔÓˆÂÒÒÓ‚ ÔÓ‰‚ÂÊÂÌÌ˚ÏË ÚÓθÍÓ Ï‰ÎÂÌÌÓÈ ‰Â„‡‰‡ˆËË (Á‡ Ò˜ÂÚ ÒÎÛ˜‡ÈÌ˚ı ÏÛÚ‡ˆËÈ), ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ ÏÓ„ÛÚ ÒÎÛÊËÚ¸ Ó˜Â̸ Û‰Ó·-Ì˚ÏË Ï‡ÍÂ‡ÏË ˝‚ÓβˆËË ÌÂÒÛ˘Ëı Ëı Ó„‡ÌËÁÏÓ‚. éÚÒÛÚÒÚ‚Ë ÙÛÌ͈ËË (̇ ÛÓ‚Ì ÍÎÂÚÍË Ë Ó„‡ÌËÁχ) Û ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ‚ ˆÂÎÓÏ Ë ·Óθ¯ËÌÒÚ‚‡ Ëı ÍÓÔËÈ ‚ „ÂÌÓÏ ‰Â·ÂÚ Ëı ÔËÁ͇̇ÏË, Ì ÔÓ‰‚Â-ÊÂÌÌ˚ÏË ÍÓÌ‚Â„ÂÌÚÌÓÈ ˝‚ÓβˆËË. ùÚÓ ‚˚„Ó‰ÌÓ ÓÚ΢‡ÂÚ Ëı Í‡Í ÓÚ ÏÓÙÓÎӄ˘ÂÒÍËı ÔËÁ̇ÍÓ‚, Ú‡Í Ë ÓÚ ÌÛÍÎÂÓÚˉÌ˚ı ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ „ÂÌÓ‚. èÓ ÒÛÚË, ‡Ì‡ÎËÁ ̇ ÓÒÌÓ‚Â ÍÓÓÚÍËı ÂÚÓÔÓÁÓ-ÌÓ‚ – ˝ÚÓ ˜‡ÒÚÌ˚È ÒÎÛ˜‡È ÙËÎÓ„ÂÌÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎË-Á‡, ËÒÔÓθÁÛ˛˘Â„Ó ‰ÍË „ÂÌÓÏÌ˚ ËÁÏÂÌÂÌËfl (rare genomic changes), ÍÓÚÓ˚ ‚Íβ˜‡˛Ú Ú‡ÍÊ ‚ÒÚ‡‚ÍË Ë ‰ÂΈËË ‚ ÌÂÍÓ‰ËÛ˛˘Ëı ÔÓÒΉӂ‡ÚÂθÌÓÒÚflı, ËÌÚ„‡ˆËË LINE Ë ÌÂÍÓÚÓ˚ı ‰Û„Ëı ÏÓ·ËθÌ˚ı ˝ÎÂÏÂÌÚÓ‚, ËÁÏÂÌÂÌË ÔÓfl‰Í‡ „ÂÌÓ‚ Ë Ëı ‰ÛÔÎË͇-ˆËË, ËÁÏÂÌÂÌË „ÂÌÂÚ˘ÂÒÍÓ„Ó ÍÓ‰‡ Ë Ú.Ô. [59]. ÇÏÂ-ÒÚÂ Ò ÚÂÏ, ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ Ò‡‚ÌËÚÂθÌÓ ÌÂ-·Óθ¯ËÂ Ë Ó˜Â̸ ÏÌÓ„ÓÍÓÔËÈÌ˚Â, Ëı ΄ÍÓ ‰ÂÚÂÍÚË-Ó‚‡Ú¸. äÓÏ ÚÓ„Ó, ÒÂÏÂÈÒÚ‚‡ SINE Ó˜Â̸ ‡ÁÌÓÓ·‡ÁÌ˚, ÓÌË ÏÌÓ„Ó ‡Á ÌÂÁ‡‚ËÒËÏÓ ‚ÓÁÌË͇ÎË ‚ ˝‚ÓβˆËË (˜ÚÓ ÔË̈ËÔˇθÌÓ ‰Îfl ÏÂÚÓ‰‡ ÒÂ-ÏÂÈÒÚ‚). ÇÒ ÔÂ˜ËÒÎÂÌÌ˚ ҂ÓÈÒÚ‚‡ ‰Â·˛Ú Ëı ÓÒÓ-·ÂÌÌÓ Û‰Ó·Ì˚ÏË Ë Ì‡Ë·ÓΠËÒÔÓθÁÛÂÏ˚ÏË ÒÂ‰Ë ‰ÍËı „ÂÌÂÚ˘ÂÒÍËı ËÁÏÂÌÂÌËÈ ‚ ÙËÎÓ„ÂÌÂÚ˘Â-ÒÍËı ËÒÒΉӂ‡ÌËflı. ë ÙÓχθÌÓÈ ÚÓ˜ÍË ÁÂÌËfl, ÓÔËÒ‡ÌÌ˚ ÔÓ‰ıÓ‰˚ Í ‡Ì‡ÎËÁÛ ˝‚ÓβˆËË Ó„‡ÌËÁÏÓ‚ ̇ ÓÒÌÓ‚Â ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ÓÒÌÓ‚‡Ì˚ ̇ ÒÎÂ‰Û˛˘Ëı ‰ÓÔÛ˘ÂÌËflı. 1. ÇÓÁÌËÍÌÛ‚ ‚ „ÂÌÓÏ ͇ÍÓ„Ó-ÚÓ ‚ˉ‡, ÒÂÏÂÈÒÚ‚Ó SINE ÔÂ‰‡ÂÚÒfl ‚ÒÂÏ ‚ˉ‡Ï, Ó·‡ÁÓ‚‡‚¯ËÏÒfl ËÁ 4

Ì„Ó. ÇÒÚÓË‚¯ËÒ¸ ‚ ÌÂÍËÈ ÎÓÍÛÒ „ÂÌÓχ, ÍÓÔËfl SINE ÒÓı‡ÌflÂÚÒfl ‚ ÌÂÏ ÌÂÓÔ‰ÂÎÂÌÌÓ ‰ÓÎ„Ó Ë ÔÂ-‰‡ÂÚÒfl ‚ÒÂÏ ‚ˉ‡Ï, ÔÓ‰ÓÎʇ˛˘ËÏ ‰‡ÌÌÛ˛ ˝‚Ó-βˆËÓÌÌÛ˛ ÎËÌ˲. 2. çÂÁ‡‚ËÒËÏÓ ‚ÓÁÌËÍÌÓ‚ÂÌË ӉË̇ÍÓ‚˚ı ÒÂ-ÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ Û ‡ÁÌ˚ı ‚ˉӂ − ÒÓ·˚ÚË Í‡ÈÌ χÎÓ‚ÂÓflÚÌÓÂ. ÇÂÓflÚÌÓÒÚ¸ ÌÂÁ‡-‚ËÒËÏÓÈ ËÌÚ„‡ˆËË SINE ‚ ÚÓ˜ÌÓ ÚÓÚ Ê ҇ÈÚ „ÂÌÓ-χ ‡ÁÌ˚ı ‚ˉӂ Ó˜Â̸ „ÂÌÓ-χ·. 3. ëÂÏÂÈÒÚ‚‡ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ Ì ÔÂ‰‡-˛ÚÒfl „ÓËÁÓÌڇθÌÓ, Ú.Â. ÓÚ Ó‰ÌËı ‚ˉӂ Í ‰Û„ËÏ, ÙËÎÓ„ÂÌÂÚ˘ÂÒÍË ËÏ ÌÂÓ‰ÒÚ‚ÂÌÌ˚Ï. чÌÌ˚ ӷ ˝‚ÓβˆËË SINE ҂ˉÂÚÂθÒÚ‚Û˛Ú Ó ÒÔ‡‚‰ÎË‚ÓÒÚË ˝ÚËı ‰ÓÔÛ˘ÂÌËÈ (Ò ÌÂÍÓÚÓ˚ÏË Ó„Ó-‚Ó͇ÏË). í‡Í, ıÓÚfl ‚ÒÚ‡Ë‚‡ÌË ÍÓÓÚÍËı ÂÚÓÔÓ-ÁÓÌÓ‚ ‚ „ÂÌÓÏ ‚ ˆÂÎÓÏ ÌÓÒËÚ ÒÎÛ˜‡ÈÌ˚È ı‡‡ÍÚÂ, ˝Ì‰ÓÌÛÍ·Á˚ ‰ÎËÌÌ˚ı ÂÚÓÔÓÁÓÌÓ‚ ÔÓfl‚Îfl˛Ú ÌÂÍÓÚÓÛ˛ ÒÔˆËÙ˘ÌÓÒÚ¸, ˜ÚÓ Û‚Â΢˂‡ÂÚ ‚ÂÓflÚ-ÌÓÒÚ¸ ÌÂÁ‡‚ËÒËÏÓÈ ËÌÚ„‡ˆËË SINE ÚÓ˜ÌÓ ‚ ÚÛ Ê ÚÓ˜ÍÛ „ÂÌÓχ. ÑÂÈÒÚ‚ËÚÂθÌÓ, ÓÔËÒ‡ÌÓ ÌÂÒÍÓθÍÓ ÒÎÛ˜‡Â‚ ÌÂÁ‡‚ËÒËÏÓ„Ó ‚ÒÚ‡Ë‚‡ÌËfl ÍÓÓÚÍËı ÂÚÓ-ÔÓÁÓÌÓ‚ ÔËÏÂÌÓ ‚ ÚÓÚ Ê ‡ÈÓÌ [60] Ë Ó‰ËÌ ÒÎÛ˜‡È – ÚÓ˜ÌÓ ‚ ÚÛ Ê ÚÓ˜ÍÛ „ÂÌÓχ [61], Ӊ̇ÍÓ Ú‡ÍË ÒÓ·˚-ÚËfl ÌÓÒflÚ ËÒÍβ˜ËÚÂθÌ˚È ı‡‡ÍÚÂ, Ë Ëı ‚ÎËflÌËÂÏ Ì‡ ÂÁÛÎ¸Ú‡Ú ÙËÎÓ„ÂÌÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎËÁ‡ ÏÓÊÌÓ ÔÂÌ·˜¸. íÓ˜ÌÓ Ú‡Í Ê ÔË̈ËÔˇθÌÓ ‚ÓÁÏÓÊ̇ ˝ÎËÏË-̇ˆËfl ÍÓÌÍÂÚÌ˚ı ÍÓÔËÈ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ËÁ „ÂÌÓχ. í‡Í‡fl ˝ÎËÏË̇ˆËfl ÏÓÊÂÚ ÔÓËÒıÓ‰ËÚ¸ ÌÂ-ÒÔˆËÙ˘ÂÒÍË, Ú.Â. Á‡ı‚‡Ú˚‚‡Ú¸ ‡ÈÓÌ˚, ÔË΄‡˛-˘ËÂ Í Ò‡ÈÚÛ ‚ÒÚ‡Ë‚‡ÌËfl (‚ ˝ÚÓÏ ÒÎÛ˜‡Â Ó̇ Ì ‚ÎËfl-ÂÚ Ì‡ ÂÁÛÎ¸Ú‡Ú ÙËÎÓ„ÂÌ‚ÎËfl-ÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎËÁ‡, Ú‡Í Í‡Í ÉÂÌÓÏ ãÓÍÛÒ Ä ãÓÍÛÒ Å Çˉ˚ 1 2 3 4 Çˉ˚ èñê ãÓÍÛÒ Ä ãÓÍÛÒ Å SINE ‚ ÎÓÍÛÒÂ Ä Å 1 2 3 4 Ä Å 1 2 3 4 1 2 3 4 Çˉ˚ êËÒ. 5. àÒÔÓθÁÓ‚‡ÌË ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ‚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ËÒÒΉӂ‡ÌËflı. åÂÚÓ‰ ÍÓÔËÈ. ëÚÂÎÍË ‚ ‚ÂıÌÂÈ ˜‡ÒÚË ËÒÛÌ͇ Ó·ÓÁ̇˜‡˛Ú Ô‡ÈÏÂ˚, ÒÔˆËÙ˘Ì˚ ‰Îfl „ÂÌÓÏÌ˚ı ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ ÎÓÍÛÒÓ‚ Ä Ë Å. ëÚÂÎ͇ÏË Ì‡ ‰‚ ÔÓ͇Á‡Ì˚ ÏÓÏÂÌÚ˚ ËÌÚ„‡ˆËË ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ‚ ÎÓÍÛÒ˚ Ä Ë Å.

åéãÖäìãüêçÄü ÅàéãéÉàü      ÚÓÏ 43      ‹ 5      2009  ΄ÍÓ Ó·Ì‡ÛÊËÚ¸), Ë ÒÔˆËÙ˘ÂÒÍË, ÔÓ ‚ÒÂÈ ‚ˉË-ÏÓÒÚË, Á‡ Ò˜ÂÚ ÂÍÓÏ·Ë̇ˆËË ÏÂÊ‰Û TSD. èÓ ÔË-·ÎËÁËÚÂθÌ˚Ï ÓˆÂÌÍ‡Ï 0.5–1% ‚ÒÂı ÎÓÍÛÒÓ‚ ÔËχ-ÚÓ‚, ÓÚ΢‡˛˘ËıÒfl Û ‡ÁÌ˚ı ‚ˉӂ ̇΢ËÂÏ Alu, Ò‚flÁ‡ÌÓ ËÏÂÌÌÓ Ò Ú‡ÍÓÈ ˝ÎËÏË̇ˆËÂÈ, ‡ ÌÂ Ò ËÌÚ„‡-ˆËÂÈ ÍÓÔËË SINE [62]. é·Â ˝ÚË ÔÓ·ÎÂÏ˚ ÒÛ˘ÂÒÚ‚ÂÌ-Ì˚ ÚÓθÍÓ ‰Îfl ÏÂÚÓ‰‡ ÍÓÔËÈ, ‚ÔÓ˜ÂÏ, Ë ‚ ˝ÚÓÏ ÒÎÛ-˜‡Â ÓÌË Î„ÍÓ ÛÒÚ‡Ìfl˛ÚÒfl ÔË Û‚Â΢ÂÌËË ˜ËÒ· ‡Ì‡ÎËÁËÛÂÏ˚ı ÎÓÍÛÒÓ‚. äÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ ÔÓËÒıÓ‰flÚ ËÁ ÔÒ‚‰Ó„Â-ÌÓ‚ ÍÎÂÚÓ˜Ì˚ı êçä Ë, ˜‡˘Â ‚Ò„Ó, 3'-ÍÓ̈‚ÓÈ Ó·-·ÒÚË LINE, ˜¸˛ ‚ÂÚ‡ÁÛ ËÒÔÓθÁÛÂÚ ‰‡ÌÌ˚È SINE. é‰Ì‡ÍÓ Ëı ‚ÓÁÌËÍÌÓ‚ÂÌË ‚Íβ˜‡ÂÚ ÏÌÓÊÂÒÚ‚Ó ‰Û-„Ëı ÔÂÂÒÚÓÂÍ, Ú‡ÍËı Í‡Í ‚ÌÛÚÂÌÌË ‰ÂΈËË Ë ‰Û-ÔÎË͇ˆËË. éÒÓ·Ó ÒΉÛÂÚ ÓÚÏÂÚËÚ¸ ÔÓfl‚ÎÂÌË ÒÔÂ-ˆËÙ˘ÂÒÍËı ÌÛÍÎÂÓÚˉÌ˚ı ÔÓÒΉӂ‡ÚÂθÌÓÒÚÂÈ ÌÂ-ËÁ‚ÂÒÚÌÓ„Ó ÔÓËÒıÓʉÂÌËfl Û ·Óθ¯ËÌÒÚ‚‡ ÒÂÏÂÈÒÚ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚, ‡ÒÔÓÎÓÊÂÌÌ˚ı ÏÂÊ‰Û Úêçä-Ó‰ÒÚ‚ÂÌÌ˚Ï Û˜‡ÒÚÍÓÏ Ë ı‚ÓÒÚÓÏ. ÇÒ ÔÂÂ-˜ËÒÎÂÌÌÓ ‰Â·ÂÚ Ô‡ÍÚ˘ÂÒÍË Ì‚ÓÁÏÓÊÌ˚Ï ÌÂÁ‡-‚ËÒËÏÓ ‚ÓÁÌËÍÌÓ‚ÂÌË ‰‚Ûı Ó‰Ë̇ÍÓ‚˚ı ËÎË Ó˜Â̸ ÔÓıÓÊËı SINE, Á‡ ËÒÍβ˜ÂÌËÂÏ Ú‡Í Ì‡Á˚‚‡ÂÏ˚ı ÔÓÒÚ˚ı ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚, ÍÓÚÓ˚ ÒÓÒÚÓflÚ ËÁ ÔÓÒΉӂ‡ÚÂθÌÓÒÚË, ÔÓËÒıÓ‰fl˘ÂÈ ËÁ ÍÎÂÚÓ˜ÌÓÈ êçä, Ë Ä-·Ó„‡ÚÓ„Ó ı‚ÓÒÚ‡ [63]. í‡Í, ÍÓÓÚÍË ÂÚÓ-ÔÓÁÓÌ˚ ID „˚ÁÛÌÓ‚, Vic-1 ‚Â·Î˛‰Ó‚ Ë Das-1 ·ÓÌÂ-ÌÓÒˆ‡, ÔÓËÒıÓ‰fl˘Ë ÓÚ ‡Î‡ÌËÌÓ‚ÓÈ Úêçä, ËÏÂ˛Ú Ó˜Â̸ ·Óθ¯Ó ÒıÓ‰ÒÚ‚Ó Ë ÔË ˝ÚÓÏ ÌÂÁ‡‚ËÒËÏÓ ÔÓËÒıÓʉÂÌËÂ. í‡ÍËÏ Ó·‡ÁÓÏ, ÔÓÒÚ˚ ÍÓÓÚÍË ÂÚÓÔÓÁÓÌ˚ Ì ÒΉÛÂÚ ËÒÔÓθÁÓ‚‡Ú¸ ‚ ÏÂÚӉ ÒÂ-ÏÂÈÒÚ‚ (Í ‰‚ÛÏ ‰Û„ËÏ ÏÂÚÓ‰‡Ï ˝ÚÓ Ì ÓÚÌÓÒËÚÒfl). ÇÓÔÓÒ Ó „ÓËÁÓÌڇθÌÓÏ ÔÂÂÌÓÒ ÂÚÓÚ‡ÌÒ-ÔÓÁÓÌÓ‚ „ÛÎflÌÓ Ó·ÒÛʉ‡ÂÚÒfl ‚ ̇ۘÌÓÈ ÎËÚÂ‡ÚÛ-Â. ÉÓËÁÓÌڇθÌ˚È ÔÂÂÌÓÒ ÏÓ·ËθÌ˚ı ˝ÎÂÏÂÌÚÓ‚ – Ó·˚˜ÌÓ fl‚ÎÂÌË ‚ ÒÎÛ˜‡Â Ñçä-Ú‡ÌÒÔÓÁÓÌÓ‚ Ë LTR-ÂÚÓÚ‡ÌÒÔÓÁÓÌÓ‚ [64]. ÉÓËÁÓÌڇθÌ˚È ÔÂ-ÂÌÓÒ ‰ÎËÌÌ˚ı ÂÚÓÔÓÁÓÌÓ‚ Ú‡ÍÊ ‚ÓÁÏÓÊÂÌ, ıÓÚfl, ÔÓ ‚ÒÂÈ ‚ˉËÏÓÒÚË, ÔËÒÛ˘ Ó˜Â̸ ÌÂÏÌÓ„ËÏ ÒÂÏÂÈ-ÒÚ‚‡Ï LINE (̇ÔËÏÂ, LINE Í·‰˚ Bov-B [18, 65]). ÖÒÚ¸ ÌÂÒÍÓθÍÓ ÔÛ·ÎË͇ˆËÈ, ‰ÂÍ·ËÛ˛˘Ëı „ÓË-ÁÓÌڇθÌ˚È ÔÂÂÌÓÒ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ (‚ Ó‰-ÌÓÈ ËÁ ÌËı “„ÓËÁÓÌڇθÌ˚È ÔÂÂÌÓÒ SINE” ‰‡Ê ÙË„ÛËÛÂÚ ‚ Á‡„ÓÎÓ‚Í [65]). ç‡ Ò‡ÏÓÏ ‰ÂÎÂ, ‚ ·Óθ-¯ËÌÒÚ‚Â ÒÎÛ˜‡Â‚ ˜¸ ˉÂÚ Ó ÔÂÂÌÓҠ̇‚ÚÓÌÓÏ-Ì˚ı LINE, ÍÓÚÓ˚ ‡Ì¸¯Â Ô˘ËÒÎflÎË Í ÍÓÓÚÍËÏ ÂÚÓÔÓÁÓ̇Ï. ÉÓËÁÓÌڇθÌ˚È ÔÂÂÌÓÒ SINE Ú‡Í-Ê ÔË‚ÎÂ͇ÎË Í ËÌÚÂÔÂÚ‡ˆËË ‰‡ÌÌ˚ı ÔÓ ÍÓÓÚ-ÍËÏ ÂÚÓÔÓÁÓÌ‡Ï ‚ ÒÎÛ˜‡Â ˝‚ÓβˆËË ÎÓÒÓÒ‚˚ı [66]. äÓÓÚÍËÈ ÂÚÓÔÓÁÓÌ SmaI ‡ÒÔÓÒÚ‡ÌÂÌ Ë Ó˜Â̸ ÔÓıÓÊ ‚ „ÂÌÓχı ‰‚Ûı ÓÚ‰‡ÎÂÌÌ˚ı ‚ÂÚ‚ÂÈ ˝ÚËı ˚· – ÒË„Ó‚˚ı, Ò Ó‰ÌÓÈ ÒÚÓÓÌ˚, ÍÂÚ˚ Ë „Ó·Û¯Ë, Ò ‰Û„ÓÈ, ‚ ÚÓ ‚ÂÏfl Í‡Í ‚ „ÂÌÓχı ‰Û„Ëı ‚ˉӂ Â„Ó ÍÓ-ÔËË ÌÂÏÌÓ„Ó˜ËÒÎÂÌÌ˚ Ë ÏÂÌ ÒıÓ‰Ì˚. ùÚÓ ÔÓÁ‚ÓÎË-ÎÓ Ô‰ÔÓÁ‚ÓÎË-ÎÓÊËÚ¸ [66] ‚ÓÁÏÓÊÌÓÒÚ¸ Ëı „ÓËÁÓÌڇθÌÓ-„Ó ÔÂÂÌÓÒ‡. ÇÏÂÒÚÂ Ò ÚÂÏ, ̇ ̇¯ ‚Á„Îfl‰, ÒÛ˘ÂÒÚ‚ÛÂÚ ·ÓΠÔÓÒÚÓ ӷ˙flÒÌÂÌËÂ: ˝ÚÓÚ ÍÓÓÚÍËÈ ÂÚÓÔÓ-ÁÓÌ ·˚Î ‡ÍÚË‚ÂÌ Û Ô‰ÍÓ‚ ‚ÒÂı ÎÓÒÓÒ‚˚ı Ë ÔÓÒÚÂ-ÔÂÌÌÓ ÛÚ‡ÚËÎ ‡ÍÚË‚ÌÓÒÚ¸ Û ‚ÒÂı ÌÂÒË„Ó‚˚ı ˚·, ÍÓÏ ÍÂÚ˚ Ë „Ó·Û¯Ë, ‡ “ÌÂÔÓıÓÊÂÒÚ¸” Ô‰ÍÓ‚˚ı ÍÓÔËÈ Ó·˙flÒÌflÂÚÒfl ÏÛÚ‡ˆËÓÌÌÓÈ ‰Â„‡‰‡ˆËÂÈ, ˉÛ-˘ÂÈ Ò ÏÓÏÂÌÚ‡ Ëı ‚ÓÁÌËÍÌÓ‚ÂÌËfl Û ‰‚ÌËı ÎÓÒÓÒÂ-‚˚ı. ÇÔÓ˜ÂÏ, Ú‡ Ê „ÛÔÔ‡ ËÒÒΉӂ‡ÚÂÎÂÈ Ô‰ÎÓ-ÊË· Ë ‰Û„Ó ӷ˙flÒÌÂÌË ˝ÚËÏ ‰‡ÌÌ˚Ï, ‡ ËÏÂÌÌÓ, ÉÂÌÓÏ Çˉ˚ ÍÓÔËfl α 1 2 3 4 èñê ÍÓÔËfl β ÍÓÔËfl γ ÍÓÔËfl δ 1 2 3 4 1 2 3 4 Çˉ˚ Çˉ˚ α δ êËÒ. 6. àÒÔÓθÁÓ‚‡ÌË ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ‚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ËÒÒΉӂ‡ÌËflı. àÌÚÂ-SINE-èñê. ëÚÂÎÍË ‚ ‚Âı-ÌÂÈ ˜‡ÒÚË ËÒÛÌ͇ Ó·ÓÁ̇˜‡˛Ú Ô‡ÈÏÂ˚, ÒÔˆËÙ˘Ì˚Â Í SINE. ëÚÂÎ͇ÏË Ì‡ ‰‚ ÔÓ͇Á‡Ì˚ ÏÓÏÂÌÚ˚ ËÌÚ„‡ˆËË ÍÓÔËÈ α Ë δ ‰‡ÌÌÓ„Ó ÒÂÏÂÈÒÚ‚‡ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚. 4*

ËÌÚÓ„ÂÒÒ˲ [67]. ç‡ÍÓ̈, ÓÔËÒ‡Ì „ÓËÁÓÌڇθ-Ì˚È ÔÂÂÌÓÒ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ Ë ˆÂÎÓ„Ó fl‰‡ ‰Û„Ëı Ñçä ÎÓÒÓÒÂÈ ‚ „ÂÌÓÏ Ô‡‡ÁËÚ‡, ‰Ë„ÂÌÂÚ˘Â-ÒÍÓ„Ó ÒÓ҇θ˘Ë͇ Schistosoma japonicum [68, 69]. éÔËÒ‡Ì Ë ÔÂÂÌÓÒ ÍÓÓÚÍÓ„Ó ÂÚÓÔÓÁÓ̇ ÂÔÚËÎËÈ ‚ „ÂÌÓÏ Ëı ÔÓÍÒ‚ËÛÒ‡, Ú‡ÍÊ (Ì ‚ÔÓÎÌ ÍÓÂÍÚÌÓ) ̇Á‚‡ÌÌ˚È “„ÓËÁÓÌڇθÌ˚Ï ÔÂÂÌÓÒÓÏ” [70]. ÄÍ-ÚË‚ÌÓÒÚ¸ Ú‡ÍËı ÔÂÂÌÂÒÂÌÌ˚ı ÍÓÔËÈ ÍÓÓÚÍËı Â-ÚÓÔÓÁÓÌÓ‚ Ì ÛÒÚ‡ÌÓ‚ÎÂ̇ Ë Ô‰ÒÚ‡‚ÎflÂÚÒfl Í‡ÈÌ χÎÓ‚ÂÓflÚÌÓÈ, ÔÓÒÍÓθÍÛ SINE – ̇‚ÚÌÓÏÌ˚ ÏÓ-·ËθÌ˚ ˝ÎÂÏÂÌÚ˚, Ë Ëı ‡ÁÏÌÓÊÂÌË Ú·ÛÂÚ ‡Í-ÚË‚ÌÓÒÚË ÒÓÓÚ‚ÂÚÒÚ‚Û˛˘Ëı ‡‚ÚÓÌÓÏÌ˚ı ÂÚÓ-Ú‡ÌÒÔÓÁÓÌÓ‚ – LINE. í‡ÍËÏ Ó·‡ÁÓÏ, ÓÔËÒ‡ÌÌ˚ ÒÎÛ˜‡Ë ÔÂÂÌÓÒ‡ Ñçä SINE Ì ÏÓ„ÛÚ ·˚Ú¸ ̇Á‚‡Ì˚ „ÓËÁÓÌڇθÌ˚Ï ÔÂÂÌÓÒÓÏ ÍÓÓÚÍÓ„Ó ÂÚÓÔÓÁÓ̇ (ÔÓ͇ Ì ·Û‰ÂÚ ‰Ó͇Á‡ÌÓ Â„Ó ‡ÁÏÌÓÊÂÌË ‚ „ÂÌÓÏ ˆËÔËÂÌÚ‡). Ç Î˛·ÓÏ ÒÎÛ˜‡Â, ÔÓ‰Ó·Ì˚È ÔÂÂÌÓÒ Ñçä SINE ÌËÍ‡Í Ì ÏÓÊÂÚ ÔÓ‚ÎËflÚ¸ ̇ ÂÁÛθڇÚ˚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎËÁ‡, Ôӂ‰ÂÌÌÓ„Ó Ò ËÒÔÓθ-ÁÓ‚‡ÌËÂÏ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚. é‰Ì‡ÍÓ ÒÛ˘ÂÒÚ‚ÛÂÚ ÒËÚÛ‡ˆËfl, ‚ ÍÓÚÓÓÈ ÂÁÛθ-Ú‡Ú˚ ‡Ì‡ÎËÁ‡ ̇ ÓÒÌÓ‚Â SINE ÏÓ„ÛÚ Ì ÒÓÓÚ‚ÂÚÒÚ‚Ó-‚‡Ú¸ ̇ÒÚÓfl˘ÂÏÛ ‰Â‚Û ‚ˉӂ. ꘸ ˉÂÚ Ó fl‚ÎÂÌËË, ̇Á˚‚‡ÂÏÓÏ ÌÂÔÓÎÌÓÈ ÒÓÚËÓ‚ÍÓÈ ÎËÌËÈ: ÔË Ó˜Â̸ ·˚ÒÚÓÏ ‚ˉÓÓ·‡ÁÓ‚‡ÌËË ËÒıÓ‰Ì˚È ÔÓÎË-ÏÓÙËÁÏ ÔËÁ̇ÍÓ‚ (̇ÔËÏÂ, ÎÓ͇ÎËÁ‡ˆËfl SINE ‚ ÍÓÌÍÂÚÌ˚ı ÎÓÍÛÒ‡ı) ÒÓı‡ÌflÂÚÒfl ÌÂÍÓÚÓÓ ‚ÂÏfl ÔÓÒΠ‡Á‰ÂÎÂÌËfl ‚ˉӂ Ë Ëı Á‡ÍÂÔÎÂÌË ÔÓËÒıÓ-‰ËÚ ÔÓÁÊ (ËÒ. 7). ë Ú‡ÍÓÈ ÒËÚÛ‡ˆËÂÈ ÒÚÓÎÍÌÛÎËÒ¸ flÔÓÌÒÍË ËÒÒΉӂ‡ÚÂÎË, ËÁÛ˜‡‚¯Ë ˝‚ÓβˆË˛ ˆËı-Îˉ – ˚· ‡ÙË͇ÌÒÍËı ÓÁÂ [71]. èÓ‰Ó·Ì˚ ÂÁÛθ-Ú‡Ú˚ ÏÓÊÌÓ ÓÊˉ‡Ú¸ ‚ ÒÎÛ˜‡Â ÏÂʂˉӂÓÈ „Ë·ˉË-Á‡ˆËË. ëÚÓËÚ Î˯¸ ÓÚÏÂÚËÚ¸, ˜ÚÓ ÔÓ‰Ó·Ì˚ ÒÎÛ˜‡Ë ËÒÍβ˜ËÚÂθÌ˚ Ë Ô‰ÒÚ‡‚Îfl˛Ú ·Óθ¯Û˛ ÔÓ·ÎÂ-ÏÛ ‰Îfl ‚ÒÂı ÏÂÚÓ‰Ó‚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍÓ„Ó ‡Ì‡ÎËÁ‡. ֢ ӉÌÓ Ó„‡Ì˘ÂÌË ÔËÏÂÌÂÌËfl ÍÓÓÚÍËı Â-ÚÓÔÓÁÓÌÓ‚ ‚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ËÒÒΉӂ‡ÌËflı ÒÓ-ÒÚÓËÚ ‚ ÚÓÏ, ˜ÚÓ Ò Ëı ÔÓÏÓ˘¸˛ ÏÓÊÌÓ ËÁÛ˜‡Ú¸ ̇·Ó ÚÓθÍÓ ÚÂı ‚ˉӂ, „‰Â ‡ÒÔÓÒÚ‡ÌÂÌÓ ‰‡ÌÌÓ ÒÂÏÂÈ-ÒÚ‚Ó SINE. åÓÊÌÓ Ò͇Á‡Ú¸, ˜ÚÓ „·‚ÌÓ ÔËÏÂÌÂÌË ˝ÚËı ÏÂÚÓ‰Ó‚ Ò‚flÁ‡ÌÓ Ò ÔÓ‚ÂÍÓÈ ÒÔÓÌ˚ı ˝‚Óβ-ˆËÓÌÌ˚ı „ËÔÓÚÂÁ, ıÓÚfl ˝ÚÓ Ì ËÒÍβ˜‡ÂÚ Ëı ËÒÔÓθ-ÁÓ‚‡ÌËfl Ë ‰Îfl ÔÓÒÚÓÂÌËfl ÔÓÎÌÓˆÂÌÌÓÈ ÙËÎÓ„ÂÌËË ÌÂÍÓÚÓ˚ı Ú‡ÍÒÓÌÓ‚. çÂÒÏÓÚfl ̇ ˝ÚË Ó„‡Ì˘ÂÌËfl, ÓÔËÒ‡ÌÌ˚ ÏÂÚÓ-‰˚ Ô‰ÒÚ‡‚Îfl˛Ú ÒÓ·ÓÈ Û‰Ó·Ì˚È Ë ‚ÂҸχ ̇‰ÂÊÌ˚È ËÌÒÚÛÏÂÌÚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ËÒÒΉӂ‡ÌËÈ. ᇠÒ‡‚ÌËÚÂθÌÓ ÍÓÓÚÍÓ ‚ÂÏfl ÒÛ˘ÂÒÚ‚Ó‚‡ÌËfl ˝ÚÓ„Ó ÔÓ‰ıÓ‰‡ ÓÌ ÒÛÏÂÎ ÔÓÎÛ˜ËÚ¸ ÔËÁ̇ÌË ÒÔˆˇÎËÒÚÓ‚ ‚ ӷ·ÒÚË ˝‚ÓβˆËÓÌÌÓÈ ·ËÓÎÓ„ËË, Ú‡‰ËˆËÓÌÌÓ ÒÍÂÔÚ˘ÂÒÍË ÓÚÌÓÒfl˘ËıÒfl Í ÌÓ‚˚Ï ÏÂÚÓ‰‡Ï. ÅÓΠÚÓ„Ó, ÓÌ ÔÓÁ‚ÓÎËÎ ‚ÌÂÒÚË ÌÂÒÍÓθÍÓ ÔË̈ËÔˇθ-Ì˚ı ËÒÔ‡‚ÎÂÌËÈ ‚ Ú‡‰ËˆËÓÌÌÛ˛ ÙËÎÓ„ÂÌ˲ ÏÎÂ-ÍÓÔËÚ‡˛˘Ëı. ÄÍÚ˂̇fl ‡ÎËÁ‡ˆËfl ÔÓÂÍÚÓ‚ ÒÂÍ‚ÂÌËÓ‚‡ÌËfl „ÂÌÓÏÓ‚ ÓÚÍ˚‚‡ÂÚ ·Óθ¯Ë ÔÂÒÔÂÍÚË‚˚ ‰Îfl ËÒ-ÔÓθÁÓ‚‡ÌËfl SINE ‚ ÙËÎÓ„ÂÌÂÚ˘ÂÒÍËı ËÒÒΉӂ‡-ÌËflı. ëÂÍ‚ÂÌËÓ‚‡ÌË ‰‡Ê ӉÌÓ„Ó „ÂÌÓχ ‚ fl‰Û ‡ÒÒχÚË‚‡ÂÏ˚ı ‚ˉӂ Ô‰ÓÒÚ‡‚ÎflÂÚ Ó„ÓÏÌ˚È ‚˚·Ó ÎÓÍÛÒÓ‚ Ò ÍÓÓÚÍËÏË ÂÚÓÔÓÁÓ̇ÏË ‰Îfl ‡Ì‡ÎËÁ‡ ÏÂÚÓ‰ÓÏ ÍÓÔËÈ. ïÓÚfl ‰Ó ÒËı ÔÓ ˝ÚÓÚ ÔÓ‰-ıÓ‰ ËÒÔÓθÁÓ‚‡ÎË, „·‚Ì˚Ï Ó·‡ÁÓÏ, ‰Îfl ËÁÛ˜ÂÌËfl ˝‚ÓβˆËË ÏÎÂÍÓÔËÚ‡˛˘Ëı, ÏÓÊÌÓ ÓÊˉ‡Ú¸, ˜ÚÓ ÓÌ Ì‡È‰ÂÚ ÔËÏÂÌÂÌËÂ Ë ‚ ËÒÒΉӂ‡ÌËflı ˝‚ÓβˆËË ‰Û„Ëı ÔÓÁ‚ÓÌÓ˜Ì˚ı, ‡ Ú‡ÍÊ ·ÂÒÔÓÁ‚ÓÌÓ˜Ì˚ı Ë ‡ÒÚÂÌËÈ. àÌÚ„‡ˆËfl SINE îËÍÒ‡ˆËfl ‘–’ ÎÓÍÛÒ‡ îËÍÒ‡ˆËfl ‘+’ ÎÓÍÛÒ‡ îËÍÒ‡ˆËfl ‘+’ ÎÓÍÛÒ‡ êËÒ. 7. ÇÓÁÏÓÊÌ˚È ÒˆÂ̇ËÈ ÔË ·˚ÒÚÓÏ ‚ˉÓÓ·‡ÁÓ‚‡ÌËË, ÔË ÍÓÚÓÓÏ ‡Ì‡ÎËÁ ÎÓÍÛÒÓ‚ ÍÓÓÚÍËı ÂÚÓÔÓÁÓÌÓ‚ ÏÓ-ÊÂÚ ‰‡‚‡Ú¸ Ì‚ÂÌ˚È ÂÁÛθڇÚ. èβÒ˚ Ë ÏËÌÛÒ˚ – ÔËÒÛÚÒÚ‚Ë ËÎË ÓÚÒÛÚÒÚ‚Ë ÍÓÔËË SINE ‚ ÓÔ‰ÂÎÂÌÌÓÏ ÎÓÍÛÒ „ÂÌÓχ ‚ ÔÓÔÛÎflˆËË.

åéãÖäìãüêçÄü ÅàéãéÉàü      ÚÓÏ 43      ‹ 5      2009 ëèàëéä ãàíÖêÄíìêõ

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