ΠΠ·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° 1 ΠΈ Π±Π΅Π»ΠΊΠ° ΠΎΠ±ΠΎΠ»ΠΎΡΠΊΠΈ ΠΏΡΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΠΏΠΎΡΠ΅ΠΊΡΠ²ΠΈΡΡΡΠΎΠ²
ΠΠΎ Π½Π΅Π΄Π°Π²Π½Π΅Π³ΠΎ Π²ΡΠ΅ΠΌΠ΅Π½ΠΈ Π² Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΠ΅ Π±ΡΠ»ΠΈ ΠΎΠΏΠΈΡΠ°Π½Ρ Π΄Π²Π΅ ΠΎΡΠ½ΠΎΠ²Π½ΡΠ΅ ΠΌΠΎΠ΄Π΅Π»ΠΈ Π±Π»ΠΈΠΆΠ½Π΅Π³ΠΎ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ° ΠΏΠΎΡΠ΅ΠΊΡΠ²ΠΈΡΡΡΠΎΠ². Π‘ΠΎΠ³Π»Π°ΡΠ½ΠΎ ΠΏΠ΅ΡΠ²ΠΎΠΉ, Π²ΠΈΡΡΡΠ½Π°Ρ Π ΠΠ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠΈΡΡΠ΅ΡΡΡ Π² ΡΠΎΡΠ΅Π΄Π½ΡΡ ΠΊΠ»Π΅ΡΠΊΡ Π² ΡΠΎΡΡΠ°Π²Π΅ Π²ΠΈΡΠΈΠΎΠ½Π° (Chapman et al., 1992; Oparka et al., 1996), Π° ΡΠΎΠ³Π»Π°ΡΠ½ΠΎ Π²ΡΠΎΡΠΎΠΉ — Π² ΡΠΎΡΡΠ°Π²Π΅ ΡΠΈΠ±ΠΎΠ½ΡΠΊΠ»Π΅ΠΎΠΏΡΠΎΡΠ΅ΠΈΠ΄Π° Π½Π΅Π²ΠΈΡΠΈΠΎΠ½Π½ΠΎΠΉ ΠΏΡΠΈΡΠΎΠ΄Ρ, ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½Π½ΠΎΠ³ΠΎ Π ΠΠ, Π±Π΅Π»ΠΊΠΎΠΌ ΠΎΠ±ΠΎΠ»ΠΎΡΠΊΠΈ (ΠΠ) ΠΈ ΠΏΠ΅ΡΠ²ΡΠΌ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΡΠΌ Π±Π΅Π»ΠΊΠΎΠΌ (Π’Π1) (Lough et al., 1998… Π§ΠΈΡΠ°ΡΡ Π΅ΡΡ >
- Π‘ΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅
- ΠΡΠ΄Π΅ΡΠΆΠΊΠ°
- ΠΠΈΡΠ΅ΡΠ°ΡΡΡΠ°
- ΠΡΡΠ³ΠΈΠ΅ ΡΠ°Π±ΠΎΡΡ
- ΠΠΎΠΌΠΎΡΡ Π² Π½Π°ΠΏΠΈΡΠ°Π½ΠΈΠΈ
Π‘ΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅
- ΠΠΠΠΠ ΠΠΠ’ΠΠ ΠΠ’Π£Π Π«
- 1. Π‘ΡΡΡΠΊΡΡΡΠ° Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ ΠΈ Π΅Π΅ ΠΊΠΎΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΎΠ½Π½ΡΠ΅ ΠΏΠ΅ΡΠ΅ΡΡΡΠΎΠΉΠΊΠΈ
- 1. 1. Π‘ΡΡΡΠΊΡΡΡΠ° ΠΠ, Π²Ρ ΠΎΠ΄ΡΡΠ΅Π³ΠΎ Π² ΡΠΎΡΡΠ°Π² Π²ΠΈΡΠΈΠΎΠ½Π° Π₯ΠΠ
- 1. 2. ΠΠ»ΠΈΠΊΠΎΠ·ΠΈΠ»ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΠΠ₯ΠΠ
- 1. 3. ΠΠΊΡΠΈΠ²Π°ΡΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ
- 1. 3. 1. ΠΠΊΡΠΈΠ²Π°ΡΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ ΠΏΡΠΈ ΡΠΎΡΡΠΎΡΠΈΠ»ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΠΠΠ₯ΠΠ
- 1. 3. 2. ΠΠΊΡΠΈΠ²Π°ΡΠΈΡ Π’Π1 ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ
- 1. 4. Π Π°Π·Π±ΠΎΡΠΊΠ° Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ Π₯ΠΠ
- 1. 5. Π‘ΡΡΡΠΊΡΡΡΠ° ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΠΎΠΉ ΡΠΎΡΠΌΡ Π₯ΠΠ
- 2. Π€ΡΠ½ΠΊΡΠΈΠΈ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ°
- 2. 1. Π‘Π²ΠΎΠΉΡΡΠ²Π° Π’Π1 in vitro
- 2. 1. 1. Π ΠΠ-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠ°Ρ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Π’Π
- 2. 1. 2. Π₯Π΅Π»ΠΈΠΊΠ°Π·Π½Π°Ρ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Π’Π
- 2. 1. 3. ΠΠ·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ Π’Π1 Ρ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΠ΅ΠΉ
- 2. 2. ΡΡΠ½ΠΊ1 (ΠΈΠΈ ΡΠ± 1 in vivo
- 2. 2. 1. Π ΠΎΠ»Ρ Π’Π1 Π² ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΠΌΠ΅ΠΆΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ° Π²ΠΈΡΡΡΠ°
- 2. 2. 2. Π’Π1 — ΠΈΠ½Π³ΠΈΠ±ΠΈΡΠΎΡ ΡΠΌΠΎΠ»ΠΊΠ°Π½ΠΈΡ Π³Π΅Π½ΠΎΠ²
- 2. 1. Π‘Π²ΠΎΠΉΡΡΠ²Π° Π’Π1 in vitro
- 8. 1. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΠΊΠΎΠ½ΡΡΡΡΠΊΡΠΈΠΉ, ΡΠΎΠ΄Π΅ΡΡ/ΡΠ°ΡΠΈΡ Π³Π΅Π½Ρ ΠΌΡΡΠ°Π½ΡΠΎΠ² Π’Π
- 8. 2. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΠΊΠΎΠ½ΡΡΡΡΠΊΡΠΈΠΉ, ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΡ Π³Π΅Π½Ρ ΠΠ ΠΈ Π΅Π³ΠΎ Π΄Π΅Π»Π΅ΡΠΈΠΎΠ½Π½ΡΡ ΠΌΡΡΠ°Π½ΡΠΎΠ²
ΠΠ·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° 1 ΠΈ Π±Π΅Π»ΠΊΠ° ΠΎΠ±ΠΎΠ»ΠΎΡΠΊΠΈ ΠΏΡΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΠΏΠΎΡΠ΅ΠΊΡΠ²ΠΈΡΡΡΠΎΠ² (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
ΠΠ·ΡΡΠ΅Π½ΠΈΠ΅ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΡ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΡΠ°ΡΠΏΡΠΎΡΡΡΠ°Π½Π΅Π½ΠΈΡ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΠΈΠ½ΡΠ΅ΠΊΡΠΈΠΈ Π² ΡΠ°ΡΡΠ΅Π½ΠΈΠΈ ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΎΠ΄Π½ΠΈΠΌ ΠΈΠ· Π²Π°ΠΆΠ½Π΅ΠΉΡΠΈΡ Π²ΠΎΠΏΡΠΎΡΠΎΠ² ΡΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎΠΉ ΡΠΈΡΠΎΠ²ΠΈΡΡΡΠΎΠ»ΠΎΠ³ΠΈΠΈ. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ° Π²ΠΈΡΡΡΠ½ΠΎΠ³ΠΎ Π³Π΅Π½ΠΎΠΌΠ° ΡΠ»ΡΠΆΠΈΡ ΠΎΡΠ½ΠΎΠ²ΠΎΠΉ Π΄Π»Ρ ΠΏΠΎΠ½ΠΈΠΌΠ°Π½ΠΈΡ ΠΏΡΠΈΠ½ΡΠΈΠΏΠΎΠ² Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Π²ΠΈΡΡΡΠΎΠ² Ρ Π·Π°ΡΠ°ΠΆΠ΅Π½Π½ΠΎΠΉ ΠΊΠ»Π΅ΡΠΊΠΎΠΉ.
ΠΠ΅ΠΆΠΊΠ»Π΅ΡΠΎΡΠ½ΡΠΉ ΡΡΠ°Π½ΡΠΏΠΎΡΡ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΠΈΠ½ΡΠ΅ΠΊΡΠΈΠΈ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΠ΅Ρ ΡΠΎΠ±ΠΎΠΉ ΠΏΠ΅ΡΠ΅Π΄Π²ΠΈΠΆΠ΅Π½ΠΈΠ΅ Π²ΠΈΡΡΡΠ½ΠΎΠ³ΠΎ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ ΠΌΠ°ΡΠ΅ΡΠΈΠ°Π»Π° ΠΈΠ· Π·Π°ΡΠ°ΠΆΠ΅Π½Π½ΠΎΠΉ Π² ΡΠΎΡΠ΅Π΄Π½ΡΡ, Π·Π΄ΠΎΡΠΎΠ²ΡΡ ΠΊΠ»Π΅ΡΠΊΡ ΡΠ΅ΡΠ΅Π· ΠΏΠ»Π°Π·ΠΌΠΎΠ΄Π΅ΡΠΌΡ.
ΠΠ±ΡΠ΅ΠΊΡΠΎΠΌ Π½Π°ΡΠ΅Π³ΠΎ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΡΠ²Π»ΡΠ»ΡΡ X Π²ΠΈΡΡΡ ΠΊΠ°ΡΡΠΎΡΠ΅Π»Ρ (Π₯ΠΠ) -ΡΠΈΠΏΠΎΠ²ΠΎΠΉ ΠΏΡΠ΅Π΄ΡΡΠ°Π²ΠΈΡΠ΅Π»Ρ ΡΠΎΠ΄Π° Potexvirus, ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΠΉ Π³Π΅Π½ΠΎΠΌΠ½ΡΡ ΠΎΠ΄Π½ΠΎΡΠ΅ΠΏΠΎΡΠ΅ΡΠ½ΡΡ ΠΏΠ»ΡΡ Π ΠΠ.
ΠΠΎ Π½Π΅Π΄Π°Π²Π½Π΅Π³ΠΎ Π²ΡΠ΅ΠΌΠ΅Π½ΠΈ Π² Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΠ΅ Π±ΡΠ»ΠΈ ΠΎΠΏΠΈΡΠ°Π½Ρ Π΄Π²Π΅ ΠΎΡΠ½ΠΎΠ²Π½ΡΠ΅ ΠΌΠΎΠ΄Π΅Π»ΠΈ Π±Π»ΠΈΠΆΠ½Π΅Π³ΠΎ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ° ΠΏΠΎΡΠ΅ΠΊΡΠ²ΠΈΡΡΡΠΎΠ². Π‘ΠΎΠ³Π»Π°ΡΠ½ΠΎ ΠΏΠ΅ΡΠ²ΠΎΠΉ, Π²ΠΈΡΡΡΠ½Π°Ρ Π ΠΠ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠΈΡΡΠ΅ΡΡΡ Π² ΡΠΎΡΠ΅Π΄Π½ΡΡ ΠΊΠ»Π΅ΡΠΊΡ Π² ΡΠΎΡΡΠ°Π²Π΅ Π²ΠΈΡΠΈΠΎΠ½Π° (Chapman et al., 1992; Oparka et al., 1996), Π° ΡΠΎΠ³Π»Π°ΡΠ½ΠΎ Π²ΡΠΎΡΠΎΠΉ — Π² ΡΠΎΡΡΠ°Π²Π΅ ΡΠΈΠ±ΠΎΠ½ΡΠΊΠ»Π΅ΠΎΠΏΡΠΎΡΠ΅ΠΈΠ΄Π° Π½Π΅Π²ΠΈΡΠΈΠΎΠ½Π½ΠΎΠΉ ΠΏΡΠΈΡΠΎΠ΄Ρ, ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½Π½ΠΎΠ³ΠΎ Π ΠΠ, Π±Π΅Π»ΠΊΠΎΠΌ ΠΎΠ±ΠΎΠ»ΠΎΡΠΊΠΈ (ΠΠ) ΠΈ ΠΏΠ΅ΡΠ²ΡΠΌ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΡΠΌ Π±Π΅Π»ΠΊΠΎΠΌ (Π’Π1) (Lough et al., 1998, 2000). Π Π½Π°ΡΠ΅ΠΉ Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΏΡΠΈ ΠΈΠ½ΠΊΡΠ±Π°ΡΠΈΠΈ Π ΠΠ Π₯ΠΠ Ρ ΠΠ, Π² ΠΏΡΠΈΡΡΡΡΡΠ²ΠΈΠΈ ΠΈ Π² ΠΎΡΡΡΡΡΡΠ²ΠΈΠΈ Π’Π1, in vitro ΠΎΠ±ΡΠ°Π·ΡΠ΅ΡΡΡ Π²ΠΈΡΡΡΠ½ΡΠΉ ΡΠΈΠ±ΠΎΠ½ΡΠΊΠ»Π΅ΠΎΠΏΡΠΎΡΠ΅ΠΈΠ΄ (Π²Π ΠΠ), Π½Π°Π·Π²Π°Π½Π½ΡΠΉ «ΠΎΠ΄Π½ΠΎΡ Π²ΠΎΡΡΠΎΠΉ ΡΠ°ΡΡΠΈΡΠ΅ΠΉ» (single tailed particle — STP) ΠΈ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΡΡΠΈΠΉ ΡΠΎΠ±ΠΎΠΉ Π½Π΅ΠΏΠΎΠ»Π½ΠΎΡΡΡΡ ΡΠΎΠ±ΡΠ°Π½Π½ΡΠΉ Π²ΠΈΡΠΈΠΎΠ½ (Karpova et al., 2006). Π ΠΠ Π₯ΠΠ, Π½Π°Ρ ΠΎΠ΄ΡΡΠ°ΡΡΡ Π² ΡΠΎΡΡΠ°Π²Π΅ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ ΠΈΠ»ΠΈ Π²Π ΠΠ, Π½Π΅Π΄ΠΎΡΡΡΠΏΠ½Π° Π΄Π»Ρ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ in vitro. Π‘Π²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Π’Π1 Π₯ΠΠ Ρ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΠ΅ΠΉ ΠΈΠ»ΠΈ Π²Π ΠΠ Π°ΠΊΡΠΈΠ²ΠΈΡΡΠ΅Ρ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΡ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ (Atabekov et al., 2000, Karpova et al., 2006). ΠΡ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»Π°Π³Π°Π΅ΠΌ, ΡΡΠΎ Π ΠΠ Π₯ΠΠ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠΈΡΡΠ΅ΡΡΡ Π² ΡΠΎΡΠ΅Π΄Π½ΡΡ ΠΊΠ»Π΅ΡΠΊΡ Π² ΡΠΎΡΡΠ°Π²Π΅ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠΎΠ² Π’Π1 Ρ Π²Π ΠΠ ΠΈΠ»ΠΈ Π²ΠΈΡΠΈΠΎΠ½ΠΎΠΌ Π₯ΠΠ.
ΠΠ°ΡΡΠΎΡΡΠ°Ρ ΡΠ°Π±ΠΎΡΠ° ΠΏΠΎΡΠ²ΡΡΠ΅Π½Π° Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΡΡΠ°ΡΡΠΊΠΎΠ² ΠΌΠΎΠ»Π΅ΠΊΡΠ» ΠΠ ΠΈ Π’Π1, ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π²ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΈ Π’Π1 Ρ Π²Π ΠΠ ΠΈ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΠ΅ΠΉ Π₯ΠΠ, ΠΏΡΠΈΠ²ΠΎΠ΄ΡΡΠ΅ΠΌΡ ΠΊ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ.
ΠΠ±Π·ΠΎΡ Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΡ.
Π₯-Π²ΠΈΡΡΡ ΠΊΠ°ΡΡΠΎΡΠ΅Π»Ρ (Π₯ΠΠ) — ΡΠΈΠΏΠΎΠ²ΠΎΠΉ ΠΏΡΠ΅Π΄ΡΡΠ°Π²ΠΈΡΠ΅Π»Ρ ΡΠΎΠ΄Π° Potexvirus, ΠΎΡΠ½ΠΎΡΡΡΠ΅Π³ΠΎΡΡ ΠΊ ΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Ρ Flexiviridae. ΠΠ΅Π½ΠΎΠΌΠ½Π°Ρ Π ΠΠ Π₯ΠΠ Π΄Π»ΠΈΠ½Π½ΠΎΠΉ 6345 ΠΏΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄ΠΎΠ², ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠ°Ρ «ΠΊΡΠΏ» ΡΡΡΡΠΊΡΡΡΡ Π½Π° 5' ΠΊΠΎΠ½ΡΠ΅ ΠΈ ΠΏΠΎΠ»ΠΈ, Π ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΡ Π½Π° 3' ΠΊΠΎΠ½ΡΠ΅, ΠΊΠΎΠ΄ΠΈΡΡΠ΅Ρ ΠΏΡΡΡ Π±Π΅Π»ΠΊΠΎΠ². Π Π΅ΠΏΠ»ΠΈΠΊΠ°Π·Π° (165 Π) ΡΡΠ°Π½ΡΠ»ΠΈΡΡΠ΅ΡΡΡ Π½Π΅ΠΏΠΎΡΡΠ΅Π΄ΡΡΠ²Π΅Π½Π½ΠΎ Ρ Π³Π΅Π½ΠΎΠΌΠ½ΠΎΠΉ Π ΠΠ, Π° ΡΡΠΈ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΡΡ Π±Π΅Π»ΠΊΠ° (Π’Π1, Π’Π2, Π’ΠΠ — ΠΏΡΠΎΠ΄ΡΠΊΡΡ «ΡΡΠΎΠΉΠ½ΠΎΠ³ΠΎ Π±Π»ΠΎΠΊΠ° Π³Π΅Π½ΠΎΠ²») ΠΈ Π±Π΅Π»ΠΎΠΊ ΠΎΠ±ΠΎΠ»ΠΎΡΠΊΠΈ (ΠΠ) Ρ ΡΡΠ±Π³Π΅Π½ΠΎΠΌΡΡ Π ΠΠ.
ΠΡΠ²ΠΎΠ΄Ρ.
1. 18 Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ² Π±Π΅Π»ΠΊΠ° ΠΎΠ±ΠΎΠ»ΠΎΡΠΊΠΈ (ΠΠ) Π½Π΅ ΡΡΠ΅Π±ΡΡΡΡΡ Π΄Π»Ρ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ Π²ΠΈΡΡΡΠ½ΠΎΠΉ Π ΠΠ ΠΈ ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½ΠΈΡ Π²ΠΈΡΡΡΠ½ΠΎΠ³ΠΎ ΡΠΈΠ±ΠΎΠ½ΡΠΊΠ»Π΅ΠΎΠΏΡΠΎΡΠ΅ΠΈΠ΄Π° (Π²Π ΠΠ). ΠΠ΅Π»Π΅Π½ΠΈΡ Π±ΠΎΠ»Π΅Π΅ ΡΠ΅ΠΌ 18 Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ² ΠΠ Π±Π»ΠΎΠΊΠΈΡΡΠ΅Ρ Π΅Π³ΠΎ ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ ΡΠ²ΡΠ·ΡΠ²Π°ΡΡ Π ΠΠ Π₯ΠΠ.
2. 10 Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ² ΠΠ ΠΊΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈ Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΡ Π΄Π»Ρ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° 1 (Π’Π1) Ρ Π²Π ΠΠ.
3. Π£ΡΠ°ΡΡΠΎΠΊ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° 1 (Π’Π1) Π₯ΠΠ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΠΈΠΉ Π΅Π³ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ Ρ ΡΠΎΡΡΠ΅Π²ΡΠΌΠΈ ΡΡΠ±ΡΠ΅Π΄ΠΈΠ½ΠΈΡΠ°ΠΌΠΈ ΡΠΏΠΈΡΠ°Π»ΡΠ½ΠΎ ΡΠΏΠ°ΠΊΠΎΠ²Π°Π½Π½ΠΎΠ³ΠΎ ΠΠ Π²ΠΈΡΠΈΠΎΠ½Π°, Π½Π°Ρ ΠΎΠ΄ΠΈΡΡΡ ΠΌΠ΅ΠΆΠ΄Ρ 112 ΠΈ 173 Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΠΌ ΠΎΡΡΠ°ΡΠΊΠΎΠΌ Π’Π1.
4. Π‘Π²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Π’Π1 Π₯ΠΠ Ρ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΠ΅ΠΉ Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΠΎ, Π½ΠΎ Π½Π΅Π΄ΠΎΡΡΠ°ΡΠΎΡΠ½ΠΎ Π΄Π»Ρ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ.
5. Π Π΅ΠΊΠΎΠΌΠ±ΠΈΠ½Π°Π½ΡΠ½ΡΠΉ ΠΠ Π₯ΠΠ Ρ ΡΠ΅ΡΡΡΡ ΠΎΡΡΠ°ΡΠΊΠ°ΠΌΠΈ Π³ΠΈΡΡΠΈΠ΄ΠΈΠ½Π° Π½Π° N-ΠΊΠΎΠ½ΡΠ΅ Π½Π΅ ΡΠΏΠΎΡΠΎΠ±Π΅Π½ ΠΎΠ±ΡΠ°Π·ΠΎΠ²ΡΠ²Π°ΡΡ Π²ΠΈΡΡΡΠ½ΡΠΉ ΡΠΈΠ±ΠΎΠ½ΡΠΊΠ»Π΅ΠΎΠΏΡΠΎΡΠ΅ΠΈΠ΄ in vitro. Π£Π΄Π°Π»Π΅Π½ΠΈΠ΅ ΠΎΡΡΠ°ΡΠΊΠΎΠ² Π³ΠΈΡΡΠΈΠ΄ΠΈΠ½Π° ΠΏΠΎΠ»Π½ΠΎΡΡΡΡ Π²ΠΎΡΡΡΠ°Π½Π°Π²Π»ΠΈΠ²Π°Π΅Ρ ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ ΠΠ ΡΠ²ΡΠ·ΡΠ²Π°ΡΡ Π²ΠΈΡΡΡΠ½ΡΡ Π ΠΠ Ρ ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ Π²Π ΠΠ.
ΠΠ°ΠΊΠ»ΡΡΠ΅Π½ΠΈΠ΅
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ΠΠΌΠ΅ΡΡΠΈΠ΅ΡΡ Π½Π° ΡΠ΅Π³ΠΎΠ΄Π½ΡΡΠ½ΠΈΠΉ Π΄Π΅Π½Ρ Π΄Π°Π½Π½ΡΠ΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ ΡΡΠ²Π΅ΡΠΆΠ΄Π°ΡΡ, ΡΡΠΎ Π’Π1 ΡΠ²ΡΠ·ΡΠ²Π°Π΅ΡΡΡ Ρ Π²Π ΠΠ, ΡΠΊΠΎΡΠ΅Π΅ Π²ΡΠ΅Π³ΠΎ, Π·Π° ΡΡΠ΅Ρ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ ΠΠ. ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π² ΡΡΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ ΡΠ΄Π΅Π»Π°ΡΡ Π²ΡΠ²ΠΎΠ΄, ΡΡΠΎ Π’Π1 Π½Π΅ ΡΠ²ΡΠ·ΡΠ²Π°Π΅ΡΡΡ Ρ Π²Π ΠΠ, ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½Π½ΡΠΌ Π²ΠΈΡΡΡΠ½ΠΎΠΉ Π ΠΠ ΠΈ ΠΠ, Ρ ΠΊΠΎΡΠΎΡΠΎΠ³ΠΎ Π΄Π΅Π»Π΅Π³ΠΈΡΠΎΠ²Π°Π½Ρ Π΄Π΅ΡΡΡΡ Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ².
Π ΡΠ΅ΠΌ ΠΏΡΠΈΡΠΈΠ½Π° ΡΠ°ΠΊΠΎΠ³ΠΎ ΡΡΡΠ΅ΠΊΡΠ°? Π‘ΡΡΠ΅ΡΡΠ²ΡΠ΅Ρ Π΄Π²Π° Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΡΡ ΠΎΠ±ΡΡΡΠ½Π΅Π½ΠΈΡ:
1) Π’Π1 ΡΠ²ΡΠ·ΡΠ²Π°Π΅ΡΡΡ Ρ Π²Π ΠΠ Π·Π° ΡΡΠ΅Ρ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ ΠΊΠ°ΠΊΠΈΠΌΠΈ-ΡΠΎ ΠΈΠ· Π΄Π΅ΡΡΡΠΈ Π‘ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ² ΠΠ;
2) Π’Π1 Π½Π΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΠ΅Ρ Ρ Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΠΌΠΈ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΠΌΠΈ ΠΎΡΡΠ°ΡΠΊΠ°ΠΌΠΈ ΠΠ. ΠΠ΄Π½Π°ΠΊΠΎ Π΄Π΅Π»Π΅ΡΠΈΡ Π΄Π΅ΡΡΡΠΈ Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ² ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΡ ΡΡΡΡΠΊΡΡΡΡ ΠΠ, Π΄Π΅Π»Π°ΡΡΠ΅ΠΌΡ Π½Π΅Π΄ΠΎΡΡΡΠΏΠ½ΡΠΌ ΡΡΠ°ΡΡΠΎΠΊ, Ρ ΠΊΠΎΡΠΎΡΡΠΌ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΎΠ²Π°Π» Π’Π1 ΠΏΡΠΈ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠΈ Ρ Π²Π ΠΠ.
ΠΠ°ΠΊΠΎΠ΅ ΠΈΠ· ΡΡΠΈΡ ΠΎΠ±ΡΡΡΠ½Π΅Π½ΠΈΠΉ ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²ΡΠ΅Ρ Π΄Π΅ΠΉΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ? Π ΡΠ°Π±ΠΎΡΠ΅ Baratova et al. (1992) ΠΏΠΎΠ»ΡΡΠ΅Π½Ρ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ, ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΡΡΠΈΠ΅ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ Π‘-ΠΊΠΎΠ½Π΅Ρ Π±ΠΎΠ»ΡΡΠΈΠ½ΡΡΠ²Π° ΠΌΠΎΠ»Π΅ΠΊΡΠ» ΠΠ, ΠΎΠ±ΡΠ°Π·ΡΡΡΠΈΡ Π²ΠΈΡΡΡΠ½ΡΡ ΡΠ°ΡΡΠΈΡΡ, Π½Π°Ρ ΠΎΠ΄ΠΈΡΡΡ Π² Π³Π»ΡΠ±ΠΈΠ½Π΅ Π΅Π΅ ΡΡΡΡΠΊΡΡΡΡ, ΡΡΠΎ ΡΡΠ°Π²ΠΈΡ ΠΏΠΎΠ΄ ΡΠΎΠΌΠ½Π΅Π½ΠΈΠ΅ Π½Π°ΡΠ΅ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠ΅Π½ΠΈΠ΅ ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΈ Π΄Π°Π½Π½ΠΎΠ³ΠΎ ΡΡΠ°ΡΡΠΊΠ° ΠΠ Ρ Π’Π1. ΠΠ΄Π½Π°ΠΊΠΎ, Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΠΎ ΠΎΡΠΌΠ΅ΡΠΈΡΡ, ΡΡΠΎ ΠΌΠ΅ΡΠΎΠ΄ ΡΡΠΈΡΠΈΠ΅Π²ΠΎΠΉ ΠΏΠ»Π°Π½ΠΈΠ³ΡΠ°ΡΠΈΠΈ, ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½Π½ΡΠΉ Π² ΡΠ°Π±ΠΎΡΠ΅ Baratova et al. (1992), Π΄Π°Π΅Ρ ΡΡΡΠ΅Π΄Π½Π΅Π½Π½ΡΡ ΠΊΠ°ΡΡΠΈΠ½Ρ Π΄Π»Ρ Π²ΡΠ΅Ρ ΠΠ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ. Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, Π΄Π°Π½Π½ΡΠ΅, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Ρ ΠΏΠΎΠΌΠΎΡΡΡ ΡΡΠΎΠ³ΠΎ ΠΌΠ΅ΡΠΎΠ΄Π°, ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ, Π½Π°ΠΌ ΡΠ΄Π΅Π»Π°ΡΡ Π·Π°ΠΊΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΠΎ ΡΠΎΠΌ, ΠΊΠ°ΠΊΠΈΠ΅ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΠ΅ ΠΎΡΡΠ°ΡΠΊΠΈ Π½Π°Ρ ΠΎΠ΄ΡΡΡΡ Π½Π° ΠΏΠΎΠ²Π΅ΡΡ Π½ΠΎΡΡΠΈ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ Ρ Π±ΠΎΠ»ΡΡΠΈΠ½ΡΡΠ²Π° Π²Ρ ΠΎΠ΄ΡΡΠΈΡ Π² Π΅Ρ ΡΠΎΡΡΠ°Π² ΡΡΠ±ΡΠ΅Π΄ΠΈΠ½ΠΈΡ ΠΠ. Π£ ΠΌΠΎΠ»Π΅ΠΊΡΠ» ΠΠ, ΠΎΠ±ΡΠ°Π·ΡΡΡΠΈΡ ΡΠΎΡΡΡ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ (ΠΏΠ΅ΡΠ²ΡΠΉ Π²ΠΈΡΠΎΠΊ ΡΠΏΠΈΡΠ°Π»ΠΈ Ρ ΠΊΠ°ΠΆΠ΄ΠΎΠ³ΠΎ ΠΊΠΎΠ½ΡΠ°), Π²ΠΎ Π²Π½Π΅ΡΠ½ΡΡ ΡΡΠ΅Π΄Ρ ΡΠΊΡΠΏΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΎ Π±ΠΎΠ»ΡΡΠ΅ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ², ΡΠ΅ΠΌ Ρ ΠΎΡΡΠ°Π»ΡΠ½ΡΡ ΡΡΠ±ΡΠ΅Π΄ΠΈΠ½ΠΈΡ ΠΠ.
ΠΠ΅ΡΠ²ΡΠΉ Π²ΠΈΡΠΎΠΊ ΡΠΏΠΈΡΠ°Π»ΠΈ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½ 8,9 ΠΌΠΎΠ»Π΅ΠΊΡΠ» ΠΠ Ρ ΠΊΠ°ΠΆΠ΄ΠΎΠ³ΠΎ ΠΊΠΎΠ½ΡΠ° (ΡΡΠΎ 0,6% ΠΎΡ ΠΎΠ±ΡΠ΅Π³ΠΎ ΡΠΈΡΠ»Π° ΡΡΠ±ΡΠ΅Π΄ΠΈΠ½ΠΈΡ ΠΠ, ΠΎΠ±ΡΠ°Π·ΡΡΡΠΈΡ Π²ΠΈΡΠΈΠΎΠ½ Π₯ΠΠ). Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΠΏΠΎΠ²Π΅ΡΡ Π½ΠΎΡΡΠΈ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ ΠΌΠ΅ΡΠΎΠ΄ΠΎΠΌ ΡΡΠΈΡΠΈΠ΅Π²ΠΎΠΉ ΠΏΠ»Π°Π½ΠΈΠ³ΡΠ°ΡΠΈΠΈ Π½Π΅ Π΄Π°Π΅Ρ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»Π΅Π½ΠΈΡ ΠΎ ΡΠΎΠΌ, ΠΊΠ°ΠΊΠΈΠ΅ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΠ΅ ΠΎΡΡΠ°ΡΠΊΠΈ ΡΡΠ±ΡΠ΅Π΄ΠΈΠ½ΠΈΡ ΠΠ, ΠΎΠ±ΡΠ°Π·ΡΡΡΠΈΡ ΡΠΎΡΡΡ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ, ΡΠΊΡΠΏΠΎΠ½ΠΈΡΠΎΠ²Π°Π½Ρ Π²ΠΎ Π²Π½Π΅ΡΠ½ΡΡ ΡΡΠ΅Π΄Ρ.
ΠΡΠΈ ΡΡΠΎΠΌ, ΠΊΠ°ΠΊ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ Π² ΡΠ°Π±ΠΎΡΠ°Ρ Atabekov et al. (2000), Rodionova et al. (2003), Karpova et al. (2006), Π’Π1 ΡΠ²ΡΠ·ΡΠ²Π°Π΅ΡΡΡ Ρ ΡΠΎΡΡΠΎΠΌ Π³ΠΎΠ»ΠΎΠ²ΠΊΠΈ Π²Π ΠΠ ΠΈΠ»ΠΈ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ, Ρ. Π΅., Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎ, ΠΈΠΌΠ΅Π½Π½ΠΎ Ρ ΡΠ΅ΠΌΠΈ ΠΌΠΎΠ»Π΅ΠΊΡΠ»Π°ΠΌΠΈ ΠΠ, ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΈ ΠΎ ΠΊΠΎΡΠΎΡΡΡ , Π½Π΅ Π΄Π°ΡΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ ΠΌΠ΅ΡΠΎΠ΄ΠΎΠΌ ΡΡΠΈΡΠΈΠ΅Π²ΠΎΠΉ ΠΏΠ»Π°Π½ΠΈΠ³ΡΠ°ΡΠΈΠΈ, ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½Π½ΡΠ΅ Π² ΡΠ°Π±ΠΎΡΠ΅ Baratova et al. (1992).
Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΠΌΡ ΠΌΠΎΠΆΠ΅ΠΌ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ Ρ ΠΌΠΎΠ»Π΅ΠΊΡΠ» ΠΠ, ΠΎΠ±ΡΠ°Π·ΡΡΡΠΈΡ ΡΠΎΡΠ΅Ρ Π³ΠΎΠ»ΠΎΠ²ΠΊΠΈ Π²Π ΠΠ ΠΈΠ»ΠΈ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ, Π² ΠΊΠΎΡΠΎΡΠΎΠΌ ΡΠΏΠ°ΠΊΠΎΠ²Π°Π½ 5' ΠΊΠΎΠ½Π΅Ρ Π²ΠΈΡΡΡΠ½ΠΎΠΉ Π ΠΠ, Π΄Π΅ΡΡΡΡ Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ² ΡΠΊΡΠΏΠΎΠ½ΠΈΡΠΎΠ²Π°Π½Ρ Π½Π° ΠΏΠΎΠ²Π΅ΡΡ Π½ΠΎΡΡΡ ΠΈ Π΄ΠΎΡΡΡΠΏΠ½Ρ Π΄Π»Ρ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ Π’Π1. ΠΠ΄Π½Π°ΠΊΠΎ, Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΠΎ ΠΎΡΠΌΠ΅ΡΠΈΡΡ, ΡΡΠΎ ΠΌΡ Π½Π΅ ΠΈΠΌΠ΅Π΅ΠΌ ΡΠΊΡΠΏΠ΅ΡΠΈΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΡΡ ΡΠ°ΠΊΡΠΎΠ² Π΄Π»Ρ ΠΏΠΎΠ΄ΡΠ²Π΅ΡΠΆΠ΄Π΅Π½ΠΈΡ ΡΡΠΎΠΉ Π³ΠΈΠΏΠΎΡΠ΅Π·Ρ.
Π‘ Π΄ΡΡΠ³ΠΎΠΉ ΡΡΠΎΡΠΎΠ½Ρ, Π°Π²ΡΠΎΡΡ ΡΠ°Π±ΠΎΡΡ Nemykh et al., (2008) Π²ΡΠ΄Π²ΠΈΠ³Π°ΡΡ Π³ΠΈΠΏΠΎΡΠ΅Π·Ρ, ΡΡΠΎ ΠΏΡΠΈ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠΈ Ρ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΠ΅ΠΉ Π’Π1 Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΠ΅Ρ Ρ ΡΡΠ°ΡΡΠΊΠ°ΠΌΠΈ, Π»Π΅ΠΆΠ°ΡΠΈΠΌΠΈ ΠΌΠ΅ΠΆΠ΄Ρ 35−38 ΠΈ 136−144 Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΠΌΠΈ ΠΎΡΡΠ°ΡΠΊΠ°ΠΌΠΈ ΠΠ. ΠΠ½Π°Π»ΠΈΠ·ΠΈΡΡΡ ΠΏΡΠ΅Π΄Π»ΠΎΠΆΠ΅Π½Π½ΡΡ Π² ΡΡΠΎΠΉ ΡΡΠ°ΡΡΠ΅ ΠΌΠΎΠ΄Π΅Π»Ρ ΡΡΠ΅ΡΠΈΡΠ½ΠΎΠΉ ΡΡΡΡΠΊΡΡΡΡ ΠΠ, Π½Π°Ρ ΠΎΠ΄ΡΡΠ΅Π³ΠΎΡΡ Π² ΡΠΎΡΡΠ°Π²Π΅ Π²ΠΈΡΡΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈΡΡ, ΠΌΡ Π²ΠΈΠ΄ΠΈΠΌ, ΡΡΠΎ Π΄Π΅Π»Π΅ΡΠΈΡ Π΄Π΅ΡΡΡΠΈ Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡ ΠΌΠΎΠΆΠ΅Ρ Π½Π°ΡΡΡΠ°ΡΡ ΡΡΡΡΠΊΡΡΡΡ ΠΠ ΡΠ°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΡΡΠΎ ΡΡΠ°ΡΡΠΊΠΈ 35−38 ΠΈ 136−144 Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ² ΡΡΠ°Π½ΡΡ Π½Π΅Π΄ΠΎΡΡΡΠΏΠ½ΡΠΌΠΈ Π΄Π»Ρ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ Π’Π1.
ΠΡΠ°ΠΊ, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π΄Π°Π½Π½ΡΠ΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ Π½Π°ΠΌ ΡΠ΄Π΅Π»Π°ΡΡ Π²ΡΠ²ΠΎΠ΄, ΡΡΠΎ Π’Π1 Π½Π΅ ΡΠ²ΡΠ·ΡΠ²Π°Π΅ΡΡΡ Ρ Π²Π ΠΠ, ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½Π½ΡΠΌ Π²ΠΈΡΡΡΠ½ΠΎΠΉ Π ΠΠ ΠΈ ΠΠ, Ρ Π‘-ΠΊΠΎΠ½ΡΠ° ΠΊΠΎΡΠΎΡΠΎΠ³ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½Ρ Π΄Π΅ΡΡΡΡ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°ΡΠΊΠΎΠ². ΠΠΌΠ΅ΡΡΠΈΠ΅ΡΡ Π½Π° ΡΠ΅Π³ΠΎΠ΄Π½ΡΡΠ½ΠΈΠΉ Π΄Π΅Π½Ρ Π΄Π°Π½Π½ΡΠ΅ Π½Π΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ Π½Π°ΠΌ ΡΠ΄Π΅Π»Π°ΡΡ Π²ΡΠ²ΠΎΠ΄ ΠΎ ΠΏΡΠΈΡΠΈΠ½Π°Ρ ΡΠ°ΠΊΠΎΠ³ΠΎ ΡΡΡΠ΅ΠΊΡΠ°.
ΠΠ° Π²ΡΠΎΡΠΎΠΌ ΡΡΠ°ΠΏΠ΅ ΡΠ°Π±ΠΎΡΡ, Π½Π°ΡΠ΅ΠΉ Π·Π°Π΄Π°ΡΠ΅ΠΉ Π±ΡΠ»Π° Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ ΡΡΠ°ΡΡΠΊΠ° Π’Π1, ΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΠΎΠ³ΠΎ Π·Π° ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Ρ Π²Π ΠΠ ΠΈ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ.
ΠΠ°ΠΊ Π±ΡΠ»ΠΎ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ Π² ΡΠ°Π±ΠΎΡΠ΅ Gorbalenya and Koonin (1993), Π’Π1 Π₯ΠΠ ΡΠΎΠ΄Π΅ΡΠΆΠΈΡ ΡΠ΅ΠΌΡ ΠΊΠΎΠ½ΡΠ΅ΡΠ²Π°ΡΠΈΠ²Π½ΡΡ ΠΌΠΎΡΠΈΠ²ΠΎΠ², Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠ½ΡΡ Π΄Π»Ρ Ρ Π΅Π»ΠΈΠΊΠ°Π· ΡΡΠΏΠ΅ΡΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Π° 1,2. ΠΠ° Π΄Π°Π½Π½ΡΠΉ ΠΌΠΎΠΌΠ΅Π½Ρ ΠΈΠ·ΡΡΠ΅Π½Π° ΡΡΡΡΠΊΡΡΡΠ° Π½Π΅ΡΠΊΠΎΠ»ΡΠΊΠΈΡ Π±Π΅Π»ΠΊΠΎΠ², Π²Ρ ΠΎΠ΄ΡΡΠΈΡ Π² ΡΡΠΈ ΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Π° (Caruthers and McKay, 2002). Π‘ΡΠ°Π²Π½Π΅Π½ΠΈΠ΅ ΡΡΡΡΠΊΡΡΡ ΡΡΠΈΡ Π±Π΅Π»ΠΊΠΎΠ² ΠΏΠΎΠ·Π²ΠΎΠ»ΠΈΠ»ΠΎ Π²ΡΡΠ²ΠΈΡΡ ΡΡΠ΄ ΠΎΠ±ΡΠΈΡ ΡΠ΅ΡΡ. Π’Π°ΠΊ, Π²ΡΠ΅ Ρ Π΅Π»ΠΈΠΊΠ°Π·Π½ΡΠ΅ ΠΌΠΎΡΠΈΠ²Ρ ΠΊΠ°ΠΆΠ΄ΠΎΠ³ΠΎ ΠΈΠ· ΡΡΠΈΡ Π±Π΅Π»ΠΊΠΎΠ² Π²Ρ ΠΎΠ΄ΠΈΠ»ΠΈ Π² ΡΠΎΡΡΠ°Π² Π΄Π²ΡΡ Π΄ΠΎΠΌΠ΅Π½ΠΎΠ²: 1Π, 2Π. ΠΠΎΠ»ΡΡΠ°Ρ ΡΠ°ΡΡΡ ΡΡΡΡΠΊΡΡΡΡ ΡΡΠΈΡ Π΄ΠΎΠΌΠ΅Π½ΠΎΠ², Π° ΡΠ°ΠΊΠΆΠ΅ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ Ρ Π΅Π»ΠΈΠΊΠ°Π·Π½ΡΡ ΠΌΠΎΡΠΈΠ²ΠΎΠ² Π² ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²ΡΡΡΠΈΡ Π΄ΠΎΠΌΠ΅Π½Π°Ρ , Π±ΡΠ»Π° ΠΈΠ΄Π΅Π½ΡΠΈΡΠ½Π° Ρ Π²ΡΠ΅Ρ ΠΈΠ·ΡΡΠ΅Π½Π½ΡΡ Ρ Π΅Π»ΠΈΠΊΠ°Π· ΡΡΠΏΠ΅ΡΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Π° 1,2 (Caruthers and McKay, 2002).
Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, Π²Π΅ΡΠΎΡΡΠ½ΠΎ, ΠΎΠΏΠΈΡΠ°Π½Π½ΡΠ΅ ΡΠ»Π΅ΠΌΠ΅Π½ΡΡ ΡΡΡΡΠΊΡΡΡΡ Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠ½Ρ Π΄Π»Ρ Π²ΡΠ΅Ρ Π±Π΅Π»ΠΊΠΎΠ² ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΡ ΡΠ΅ΠΌΡ ΠΊΠΎΠ½ΡΠ΅ΡΠ²Π°ΡΠΈΠ²Π½ΡΡ Ρ Π΅Π»ΠΈΠΊΠ°Π·Π½ΡΡ ΠΌΠΎΡΠΈΠ²ΠΎΠ², Ρ. Π΅. ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΠΎ, ΠΈ Π΄Π»Ρ Π’Π1 Π₯ΠΠ. Π’ΠΎΠ³Π΄Π°, ΠΈΡΡ ΠΎΠ΄Ρ ΠΈΠ· ΡΠ°ΡΠΏΠΎΠ»ΠΎΠΆΠ΅Π½ΠΈΡ Π² ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ Π’Π1 Ρ Π΅Π»ΠΈΠΊΠ°Π·Π½ΡΡ ΠΌΠΎΡΠΈΠ²ΠΎΠ², ΠΌΠΎΠΆΠ½ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ ΠΏΠΎΡΡΠΈ Π²ΡΡ ΠΏΠΎΠ»ΠΈΠΏΠ΅ΠΏΡΠΈΠ΄Π½Π°Ρ ΡΠ΅ΠΏΡ Π’Π1 ΡΡΠ°ΡΡΠ²ΡΠ΅Ρ Π² ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½ΠΈΠΈ Π΄ΠΎΠΌΠ΅Π½ΠΎΠ² 1Π, 2Π, Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠ½ΡΡ Π΄Π»Ρ Ρ Π΅Π»ΠΈΠΊΠ°Π· ΡΡΠΏΠ΅ΡΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Π° 1 (Kalinina et ai, 2002).
ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π² ΡΡΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ Π΄Π»Ρ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ ΡΡΠ΅Π±ΡΠ΅ΡΡΡ ΡΠ΅Π»ΠΎΡΡΠ½ΠΎΡΡΡ ΠΊΠΎΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΈ Π’Π1. Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΠΌΠΎΠΆΠ½ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ Π΄Π»Ρ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΠ° ΡΠ΅Π»ΠΎΡΡΠ½ΠΎΡΡΡ Π΄ΠΎΠΌΠ΅Π½ΠΎΠ² 1Π ΠΈ 2Π Π’Π1. ΠΠ° Π΄Π°Π½Π½ΠΎΠΌ ΡΡΠ°ΠΏΠ΅ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ Π½Π΅Π»ΡΠ·Ρ ΡΠ΄Π΅Π»Π°ΡΡ Π²ΡΠ²ΠΎΠ΄, Π½ΡΠΆΠ½Π° Π»ΠΈ Ρ Π΅Π»ΠΈΠΊΠ°Π·Π½Π°Ρ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Π’Π1 Π΄Π»Ρ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ. ΠΠΏΠΎΠ»Π½Π΅ Π²Π΅ΡΠΎΡΡΠ½ΠΎ, ΡΡΠΎ Π’Π1 Π°ΠΊΡΠΈΠ²ΠΈΡΡΠ΅Ρ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΡ ΠΈΠ½ΠΊΠ°ΠΏΡΠΈΠ΄ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Π ΠΠ Π·Π° ΡΡΠ΅Ρ ΠΊΠ°ΠΊΠΎΠ³ΠΎ-ΡΠΎ Π΄ΡΡΠ³ΠΎΠ³ΠΎ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ°, ΡΡΠ΅Π±ΡΡΡΠ΅Π³ΠΎ Π΄Π°Π»ΡΠ½Π΅ΠΉΡΠΈΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ.
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