Π ΠΎΠ»Ρ ΠΌΠ°ΠΆΠΎΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° ΠΌΠ ΠΠ Ρ50 Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΈ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ
![ΠΠΈΡΡΠ΅ΡΡΠ°ΡΠΈΡ: Π ΠΎΠ»Ρ ΠΌΠ°ΠΆΠΎΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° ΠΌΠ ΠΠ Ρ50 Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΈ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ
ΡΡΠΊΠ°ΡΠΈΠΎΡ](https://niscu.ru/work/3726573/cover.png)
Π Π½Π°ΡΡΠΎΡΡΠ΅Π΅Π΅ Π²ΡΠ΅ΠΌΡ Π½Π΅ Π²ΡΠ·ΡΠ²Π°Π΅Ρ ΡΠΎΠΌΠ½Π΅Π½ΠΈΡ ΡΠΎΡ ΡΠ°ΠΊΡ, ΡΡΠΎ Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ Π±Π΅Π»ΠΊΠΈ, Π°ΡΡΠΎΡΠΈΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΠΌΠΈ ΠΌΠ ΠΠ, ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΡΡ ΡΡΠ½Π΄Π°ΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΠΎ ΡΠ°Π·Π»ΠΈΡΠ½ΡΠ΅ ΠΏΡΡΠΈ ΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ ΡΡΠ΄ΡΠ±Ρ. ΠΠ΅ΡΠΌΠΎΡΡΡ Π½Π° ΠΎΡΠΊΡΡΡΠΈΠ΅ ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²Π° Π±Π΅Π»ΠΊΠΎΠ², ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ½ΡΡ ΠΊ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΠΌ ΡΠΈΠΏΠ°ΠΌ ΠΌΠ ΠΠ ΠΈ ΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΡΡ Π·Π° ΡΠ΅Π»Π΅ΠΊΡΠΈΠ²Π½ΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, Π±ΡΠ»ΠΈ Π½Π°ΠΉΠ΄Π΅Π½Ρ ΡΠΎΠ»ΡΠΊΠΎ Π΄Π²Π° ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ Π±Π΅Π»ΠΊΠ°, Π°ΡΡΠΎΡΠΈΠΈΡΠΎΠ²Π°Π½Π½ΡΡ Ρ Π±ΠΎΠ»ΡΡΠΈΠ½ΡΡΠ²ΠΎΠΌ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ … 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΅
- ΠΠ±Π·ΠΎΡ Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΡ
- Π€Π£ΠΠΠ¦ΠΠ ΠΠΠΠΠ ΠΠ«Π₯ ΠΠΠΠΠ, ΠΠ‘Π‘ΠΠ¦ΠΠΠ ΠΠΠΠΠΠ«Π₯ Π‘ ΠΌΠ ΠΠ Π ΠΠΠΠ’ΠΠΠ₯ ΠΠ£ ΠΠΠ ΠΠΠ’
- 1. Π ΠΎΠ»Ρ ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ
Π±Π΅Π»ΠΊΠΎΠ² Π² ΡΡΡΡΠΊΡΡΡΠ½ΠΎΠΉ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ
- 1. 1. Π ΠΠΠ ΠΈ Π΅Π³ΠΎ ΡΠΎΠ»Ρ Π² ΡΡΡΡΠΊΡΡΡΠ½ΠΎΠΉ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ
- 1. 3. Π‘ΡΡΡΠΊΡΡΡΠ½ΡΠ΅ ΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΡΠ΅ ΡΠ²ΠΎΠΉΡΡΠ²Π° Ρ50 ΠΈ ΡΠΎΠ΄ΡΡΠ²Π΅Π½Π½ΡΡ YB Π±Π΅Π»ΠΊΠΎΠ²
- 1. 4. Ρ5 0 ΠΊΠ°ΠΊ ΡΡΡΡΠΊΡΡΡΠ½ΡΠΉ Π±Π΅Π»ΠΎΠΊ ΠΌΠ ΠΠ
- 2. Π ΠΎΠ»Ρ Π ΠΠΠ ΠΈ Ρ50 Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ
- 2. 1. Π ΠΠΠ ΠΊΠ°ΠΊ Π½Π΅ΠΊΠ°Π½ΠΎΠ½ΠΈΡΠ΅ΡΠΊΠΈΠΉ ΡΠ°ΠΊΡΠΎΡ ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ
- 2. 2. Ρ50 ΠΈ ΡΠΎΠ΄ΡΡΠ²Π΅Π½Π½ΡΠ΅ YB Π±Π΅Π»ΠΊΠΈ ΠΊΠ°ΠΊ ΡΠ΅ΠΏΡΠ΅ΡΡΠΎΡΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ
- 2. 3. Ρ50 ΠΊΠ°ΠΊ Π½Π΅ΠΊΠ°Π½ΠΎΠ½ΠΈΡΠ΅ΡΠΊΠΈΠΉ ΡΠ°ΠΊΡΠΎΡ ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ
- 2. 4. ΠΠΎΠ·ΠΌΠΎΠΆΠ½Π°Ρ ΠΌΠΎΠ΄Π΅Π»Ρ ΡΡΠ°ΡΡΠΈΡ Π ΠΠΠ ΠΈ Ρ50 Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΌΠ ΠΠ
- 3. ΠΠ΅Ρ
Π°Π½ΠΈΠ·ΠΌΡ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΡ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ
ΡΡΠΊΠ°ΡΠΈΠΎΡ
- 3. 1. ΠΠΎΠ»ΠΈ (Π)-Ρ Π²ΠΎΡΡ ΠΈ Π ΠΠΠ — ΠΎΠ±ΡΠΈΠ΅ Π΄Π΅ΡΠ΅ΡΠΌΠΈΠ½Π°Π½ΡΡ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΌΠ ΠΠ
- 3. 2. Π‘ΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ ΠΈ Π΄Π΅ΡΠ΅ΡΠΌΠΈΠ½Π°Π½ΡΡ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΌΠ ΠΠ
- 3. 3. YB Π±Π΅Π»ΠΊΠΈ — ΠΊΠ»ΡΡΠ΅Π²ΠΎΠΉ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ ΡΠΈΡΡΠ΅ΠΌΡ ΠΌΠ°ΡΠΊΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ΅
- 4. Π’ΡΠ°Π½ΡΠ»ΡΡΠΈΡ ΠΈ ΡΠΈΡΠΎΡΠΊΠ΅Π»Π΅Ρ
Π ΠΎΠ»Ρ ΠΌΠ°ΠΆΠΎΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° ΠΌΠ ΠΠ Ρ50 Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΈ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
Π Π½Π°ΡΡΠΎΡΡΠ΅Π΅Π΅ Π²ΡΠ΅ΠΌΡ Π½Π΅ Π²ΡΠ·ΡΠ²Π°Π΅Ρ ΡΠΎΠΌΠ½Π΅Π½ΠΈΡ ΡΠΎΡ ΡΠ°ΠΊΡ, ΡΡΠΎ Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ Π±Π΅Π»ΠΊΠΈ, Π°ΡΡΠΎΡΠΈΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΠΌΠΈ ΠΌΠ ΠΠ, ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΡΡ ΡΡΠ½Π΄Π°ΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΠΎ ΡΠ°Π·Π»ΠΈΡΠ½ΡΠ΅ ΠΏΡΡΠΈ ΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ ΡΡΠ΄ΡΠ±Ρ. ΠΠ΅ΡΠΌΠΎΡΡΡ Π½Π° ΠΎΡΠΊΡΡΡΠΈΠ΅ ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²Π° Π±Π΅Π»ΠΊΠΎΠ², ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ½ΡΡ ΠΊ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΠΌ ΡΠΈΠΏΠ°ΠΌ ΠΌΠ ΠΠ ΠΈ ΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΡΡ Π·Π° ΡΠ΅Π»Π΅ΠΊΡΠΈΠ²Π½ΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, Π±ΡΠ»ΠΈ Π½Π°ΠΉΠ΄Π΅Π½Ρ ΡΠΎΠ»ΡΠΊΠΎ Π΄Π²Π° ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ Π±Π΅Π»ΠΊΠ°, Π°ΡΡΠΎΡΠΈΠΈΡΠΎΠ²Π°Π½Π½ΡΡ Ρ Π±ΠΎΠ»ΡΡΠΈΠ½ΡΡΠ²ΠΎΠΌ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ ΠΈ ΠΎΡΡΡΠ΅ΡΡΠ²Π»ΡΡΡΠΈΡ ΠΎΠ±ΡΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ. ΠΠ΄ΠΈΠ½ ΠΈΠ· Π½ΠΈΡ — Π±Π΅Π»ΠΎΠΊ Ρ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΠΌ Π²Π΅ΡΠΎΠΌ 70 ΠΊΠΠ°, Ρ ΠΎΡΠΎΡΠΎ ΠΈΠ·Π²Π΅ΡΡΠ΅Π½ ΠΊΠ°ΠΊ ΠΏΠΎΠ»ΠΈ (Π)-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΉ Π±Π΅Π»ΠΎΠΊ (poly (A)-binding protein, Π ΠΠΠ ), ΠΎΠ±Π»Π°Π΄Π°ΡΡΠΈΠΉ ΠΏΡΠ΅ΠΈΠΌΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΠΎΠΉ Π°ΡΡΠΈΠ½Π½ΠΎΡΡΡΡ ΠΊ 3'-ΠΊΠΎΠ½ΡΠ΅Π²ΡΠΌ ΠΏΠΎΠ»ΠΈ (Π) — ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΡΠΌ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ ΠΠ. Π Π½Π°ΡΡΠΎΡΡΠ΅ΠΌΡ Π²ΡΠ΅ΠΌΠ΅Π½ΠΈ Π ΠΠΠ ΡΠ²Π»ΡΠ΅ΡΡΡ Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ Ρ ΠΎΡΠΎΡΠΎ ΠΈΠ·ΡΡΠ΅Π½Π½ΡΠΌ Π±Π΅Π»ΠΊΠΎΠΌ ΠΌΠ ΠΠΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ Π΅Π³ΠΎ ΡΡΠ°ΡΡΠΈΠ΅ Π² ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΌΠ ΠΠ ΠΈ ΠΆΠΈΠ·Π½Π΅ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΠΊΠ»Π΅ΡΠΊΠΈ Π² ΡΠ΅Π»ΠΎΠΌ (Sachs and Davis, 1989; Jackson and Standart, 1990; Munroe and Jacobson, 1990; Gallie, 1991; Tarun and Sachs, 1995; Sachs et al., 1997; Tarun et al., 1997). ΠΡΡΠ³ΠΎΠΉ Π±Π΅Π»ΠΎΠΊ, p50, ΠΈΠΌΠ΅ΡΡΠΈΠΉ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΠΉ Π²Π΅Ρ ΠΎΠΊΠΎΠ»ΠΎ 35 ΠΊΠΠ°, Π½ΠΎ Π΄Π²ΠΈΠΆΡΡΠΈΠΉΡΡ ΠΏΡΠΈ SDS-ΡΠ»Π΅ΠΊΡΡΠΎΡΠΎΡΠ΅Π·Π΅ ΠΊΠ°ΠΊ Π±Π΅Π»ΠΎΠΊ Ρ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΠΌ Π²Π΅ΡΠΎΠΌ 50 ΠΊΠΠ°, Π±ΡΠ» ΠΎΠΏΠΈΡΠ°Π½ Π±ΠΎΠ»Π΅Π΅ 20 Π»Π΅Ρ Π½Π°Π·Π°Π΄ ΠΊΠ°ΠΊ Π΄ΠΎΠΌΠΈΠ½Π°Π½ΡΠ½ΡΠΉ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ ΠΠ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ , ΠΎΠ΄Π½Π°ΠΊΠΎ Π΄ΠΎ Π½Π°ΡΡΠΎΡΡΠ΅Π³ΠΎ Π²ΡΠ΅ΠΌΠ΅Π½ΠΈ ΡΡΠ½ΠΊΡΠΈΠΈ Ρ50 ΠΎΡΡΠ°ΡΡΡΡ Π½Π΅ΡΡΠ½ΡΠΌΠΈ. ΠΠ½ΠΎΠΆΠ΅ΡΡΠ²ΠΎ ΡΠ°ΠΊΡΠΎΠ² Π³ΠΎΠ²ΠΎΡΠΈΡ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ ΡΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ ΡΡΠΈΡ Π΄Π²ΡΡ Π±Π΅Π»ΠΊΠΎΠ² Π² ΡΠΎΡΡΠ°Π²Π΅ ΠΌΠ ΠΠ Π²Π°ΡΡΠΈΡΡΠ΅Ρ Π² Π·Π°Π²ΠΈΡΠΈΠΌΠΎΡΡΠΈ ΠΎΡ ΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΡΠΎΡΡΠΎΡΠ½ΠΈΡ ΠΌΠ ΠΠ Π’Π°ΠΊ, Π ΠΠΠ Π±ΡΠ» Π½Π°ΠΉΠ΄Π΅Π½ ΠΏΡΠ΅ΠΈΠΌΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΠΎ Π² Π°ΠΊΡΠΈΠ²Π½ΡΡ ΠΏΠΎΠ»ΠΈΡΠΈΠ±ΠΎΡΠΎΠΌΠ½ΡΡ ΠΌΠ ΠΠ (Van Venrooij et al., 1977; Vincent et al, 1981; Minich and Ovchinnikov, 1992). Π ΠΏΡΠΎΡΠΈΠ²ΠΎΠΏΠΎΠ»ΠΎΠΆΠ½ΠΎΡΡΡ Π ΠΠΠ , Ρ50 ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΠΌ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΠΎΠΌ ΠΊΠ°ΠΊ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎ Π½Π΅Π°ΠΊΡΠΈΠ²Π½ΡΡ , ΡΠ°ΠΊ ΠΈ Π°ΠΊΡΠΈΠ²Π½ΡΡ ΠΏΠΎΠ»ΠΈΡΠΈΠ±ΠΎΡΠΎΠΌΠ½ΡΡ Π³Π»ΠΎΠ±ΠΈΠ½ΠΎΠ²ΡΡ ΠΌΠ ΠΠ, ΠΏΡΠΈΡΠ΅ΠΌ ΡΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ Ρ50 Π½Π° ΠΌΠΎΠ»Ρ ΠΌΠ ΠΠ Π² ΠΏΠΎΠ»ΠΈΡΠΈΠ±ΠΎΡΠΎΠΌΠ½ΡΡ ΠΌΠ ΠΠ Π²Π΄Π²ΠΎΠ΅ Π½ΠΈΠΆΠ΅, ΡΠ΅ΠΌ Π² ΡΠ²ΠΎΠ±ΠΎΠ΄Π½ΡΡ Π½Π΅Π°ΠΊΡΠΈΠ²Π½ΡΡ ΠΌΠ ΠΠ. ΠΠΎΠ»Π΅Π΅ ΡΠΎΠ³ΠΎ, ΠΎΠΊΠ°Π·Π°Π»ΠΎΡΡ ΡΠ°ΠΊΠΆΠ΅, ΡΡΠΎ ΠΏΡΠΈ Π½Π΅Π²ΡΡΠΎΠΊΠΎΠΌ ΡΠΎΠΎΡΠ½ΠΎΡΠ΅Π½ΠΈΠΈ Ρ50: ΠΌΠ ΠΠ, ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²ΡΡΡΠ΅ΠΌ ΡΠ°ΠΊΠΎΠ²ΠΎΠΌΡ Π² ΠΏΠΎΠ»ΠΈΡΠΈΠ±ΠΎΡΠΎΠΌΠ½ΡΡ ΠΌΠ ΠΠ, Ρ50 ΡΡΠΈΠΌΡΠ»ΠΈΡΡΠ΅Ρ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΡ, ΡΠΎΠ³Π΄Π° ΠΊΠ°ΠΊ ΡΠ²Π΅Π»ΠΈΡΠ΅Π½ΠΈΠ΅ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²Π° Ρ50 ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΠΏΠΎΠ»Π½ΠΎΠΉ ΠΎΡΡΠ°Π½ΠΎΠ²ΠΊΠ΅ Π±Π΅Π»ΠΊΠΎΠ²ΠΎΠ³ΠΎ ΡΠΈΠ½ΡΠ΅Π·Π°. Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΠΏΠ΅ΡΠ΅Ρ ΠΎΠ΄ ΠΌΠ ΠΠ ΠΈΠ· Π½Π΅Π°ΠΊΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΡΠΎΡΡΠΎΡΠ½ΠΈΡ Π² Π°ΠΊΡΠΈΠ²Π½ΠΎΠ΅ ΡΠΎΠΏΡΠΎΠ²ΠΎΠΆΠ΄Π°Π΅ΡΡΡ ΡΠΌΠ΅Π½ΡΡΠ΅Π½ΠΈΠ΅ΠΌ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²Π° Ρ50, ΡΠ²ΡΠ·Π°Π½Π½ΠΎΠ³ΠΎ Ρ ΠΌΠ ΠΠ, ΠΈ ΠΏΠΎΡΠ²Π»Π΅Π½ΠΈΠ΅ΠΌ Π ΠΠΠ , ΠΈ Π½Π°ΠΎΠ±ΠΎΡΠΎΡ, ΠΈΠ½Π°ΠΊΡΠΈΠ²Π°ΡΠΈΡ ΠΌΠ ΠΠ ΡΠΎΠΏΡΡΠΆΠ΅Π½Π° Ρ ΡΠ²Π΅Π»ΠΈΡΠ΅Π½ΠΈΠ΅ΠΌ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²Π° Ρ50 ΠΈ Π²ΡΡΠ΅ΡΠ½Π΅Π½ΠΈΠ΅ΠΌ Π ΠΠΠ ΠΈΠ· ΡΠΎΡΡΠ°Π²Π° ΠΌΠ ΠΠ. ΠΡΠ½ΠΎΠ²ΡΠ²Π°ΡΡΡ Π½Π° ΡΡΠΈΡ Π΄Π°Π½Π½ΡΡ ΠΌΡ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΠ»ΠΈ, ΡΡΠΎ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²Π° ΡΡΠΈΡ Π΄Π²ΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ², ΡΠ²ΡΠ·Π°Π½Π½ΡΡ Ρ ΠΌΠ ΠΠ, ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΊΠ»ΡΡΠ΅Π²ΡΠΌ ΡΠ°ΠΊΡΠΎΡΠΎΠΌ ΠΎΠ±ΡΠ΅ΠΉ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ ΠΌΠ ΠΠ. Π Π΅ΡΠ΅Π½ΠΈΡ ΡΡΠΎΠ³ΠΎ Π²ΠΎΠΏΡΠΎΡΠ° ΠΈ Π±ΡΠ»Π° ΠΏΠΎΡΠ²ΡΡΠ΅Π½Π° ΠΏΠ΅ΡΠ²Π°Ρ ΡΠ°ΡΡΡ Π½Π°ΡΡΠΎΡΡΠ΅ΠΉ ΡΠ°Π±ΠΎΡΡ.
ΠΡΠΎΡΡΠΌ Π²Π°ΠΆΠ½ΡΠΌ Π°ΡΠΏΠ΅ΠΊΡΠΎΠΌ ΡΡΠΎΠΉ ΡΠ°Π±ΠΎΡΡ ΡΠ²ΠΈΠ»ΠΎΡΡ ΠΈΠ·ΡΡΠ΅Π½ΠΈΠ΅ Π²Π»ΠΈΡΠ½ΠΈΡ Ρ50 Π½Π° ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΡ ΠΌΠ ΠΠ. ΠΠ΅ΠΉΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎ, ΠΈΠ·Π²Π΅ΡΡΠ½ΠΎ, ΡΡΠΎ Ρ50 ΠΈ Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ ΡΠ²Π»ΡΡΡΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΠΌΠΈ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΠ°ΠΌΠΈ Π½Π΅Π°ΠΊΡΠΈΠ²Π½ΡΡ ΠΌΠ ΠΠ, ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΡ ΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎ Π°ΠΊΡΠΈΠ²Π½ΡΡ ΠΌΠ ΠΠ Π² Π·Π°ΠΏΠ°ΡΠ΅Π½Π½ΠΎΠΌ ΡΠΎΡΡΠΎΡΠ½ΠΈΠΈ. ΠΡΡΡΠ΄Π° Π±ΡΠ»ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠ΅Π½ΠΎ, ΡΡΠΎ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ ΠΈΠ·Π±ΡΡΠΊΠ° Ρ50 Ρ ΠΌΠ ΠΠ ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ Π½Π΅ ΠΏΡΠΎΡΡΠΎ ΠΊ ΠΈΠ½Π³ΠΈΠ±ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, Π½ΠΎ ΡΠ°ΠΊΠΆΠ΅ ΠΈ ΠΊ ΡΡΠ°Π±ΠΈΠ»ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ Π² ΡΠΎΡΠΌΠ΅ Π½Π΅Π°ΠΊΡΠΈΠ²Π½ΡΡ ΠΌΠ ΠΠ.
Π’ΡΠ΅ΡΡΠΈΠΌ Π½Π°ΠΏΡΠ°Π²Π»Π΅Π½ΠΈΠ΅ΠΌ Π½Π°ΡΠΈΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ ΡΠ²ΠΈΠ»ΡΡ ΠΏΠΎΠΈΡΠΊ Π±Π΅Π»ΠΊΠΎΠ², Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΡΡΠΈΡ Ρ Ρ50. ΠΠ΅Π΄Π°Π²Π½ΠΈΠ΅ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ, ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½Π½ΡΠ΅ Π² Π½Π°ΡΠ΅ΠΉ Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ, ΠΏΠΎΠΊΠ°Π·Π°Π»ΠΈ, ΡΡΠΎ Ρ50 ΠΏΡΠΈΠ½Π°Π΄Π»Π΅ΠΆΠΈΡ ΠΊ ΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Ρ ΠΌΡΠ»ΡΡΠΈΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ², Π½Π°ΠΉΠ΄Π΅Π½Π½ΡΡ ΠΏΠΎΡΡΠΈ Π²ΠΎ Π²ΡΠ΅Ρ ΠΆΠΈΠ²ΡΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠ°Ρ , ΠΎΡ Π±Π°ΠΊΡΠ΅ΡΠΈΠΉ Π΄ΠΎ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ , ΠΈ ΡΠ΅Π³ΡΠ»ΠΈΡΡΡΡΠΈΡ ΡΠ°Π·Π»ΠΈΡΠ½ΡΠ΅ Π°ΡΠΏΠ΅ΠΊΡΡ Π±ΠΈΠΎΠ³Π΅Π½Π΅Π·Π° ΠΌΠ ΠΠ, ΡΠ΅ΠΏΠ»ΠΈΠΊΠ°ΡΠΈΡ ΠΈ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΡ ΠΠΠ, ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΠΉ ΡΠΎΡΡ ΠΈ ΠΏΡΠΎΠ»ΠΈΡΠ΅ΡΠ°ΡΠΈΡ, Π° ΡΠ°ΠΊΠΆΠ΅ ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ°Ρ ΠΏΠ΅ΡΠ΅Π΄Π°ΡΠΈ ΡΡΡΠ΅ΡΡΠΎΠ²ΡΡ ΡΠΈΠ³Π½Π°Π»ΠΎΠ² Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ (Grant and Deeley, 1993; Lee et al, 1994; Wolffe, 1994; Ladomery and Sommerville, 1995; Graumann and Marahiel, 1996bOhga et al., 1996; Swamynathan et al., 1998). Π ΡΠ²ΡΠ·ΠΈ Ρ ΡΡΠΈΠΌ ΠΏΠΎΠΈΡΠΊ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΡ ΠΏΠ°ΡΡΠ½Π΅ΡΠΎΠ² Ρ50 ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΠ΅Ρ ΠΎΡΠΎΠ±ΡΠΉ ΠΈΠ½ΡΠ΅ΡΠ΅Ρ, ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡΠΈΠΉ ΠΏΡΠΎΠ»ΠΈΡΡ ΡΠ²Π΅Ρ Π½Π° ΡΠ°Π·Π»ΠΈΡΠ½ΡΠ΅ ΡΡΠΎΡΠΎΠ½Ρ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ Ρ50 ΠΈ Π½Π° Π΅Π³ΠΎ ΠΌΠ΅ΡΡΠΎ Π² ΠΆΠΈΠ·Π½Π΅ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΠΊΠ»Π΅ΡΠΊΠΈ Π² ΡΠ΅Π»ΠΎΠΌ.
Π ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠ΅ ΠΏΡΠΎΠ΄Π΅Π»Π°Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΡ ΠΎΠΊΠ°Π·Π°Π»ΠΎΡΡ, ΡΡΠΎ Ρ50 ΠΎΠ±Π»Π°Π΄Π°Π΅Ρ ΡΠ°ΠΊΠΎΠΉ ΠΆΠ΅ Π°ΡΡΠΈΠ½Π½ΠΎΡΡΡΡ ΠΊ ΠΏΠΎΠ»ΠΈ (Π)-ΠΏΠΎΠ»ΠΈΠ½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄Π°ΠΌ, ΠΊΠ°ΠΊ Π ΠΠΠ ΠΈ, ΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎ, ΠΌΠΎΠΆΠ΅Ρ ΠΊΠΎΠ½ΠΊΡΡΠΈΡΠΎΠ²Π°ΡΡ Ρ Π ΠΠΠ Π·Π° ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Ρ ΠΏΠΎΠ»ΠΈ (Π)-Ρ Π²ΠΎΡΡΠΎΠΌ ΠΌΠ ΠΠ. ΠΡΠΏΠΎΠ»ΡΠ·ΡΡ Π±Π΅ΡΠΊΠ»Π΅ΡΠΎΡΠ½ΡΠ΅ ΡΠΈΡΡΠ΅ΠΌΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈ ΠΌΠ ΠΠ, ΠΌΠ΅ΡΠ΅Π½Π½ΡΡ ΠΏΠΎ ΠΏΠΎΠ»ΠΈ (Π)-Ρ Π²ΠΎΡΡΡ, ΠΌΡ ΠΏΠΎΠΊΠ°Π·Π°Π»ΠΈ, ΡΡΠΎ Π΄ΠΎΠ±Π°Π²Π»Π΅Π½ΠΈΠ΅ Ρ50 ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ Π²ΡΡΠ΅ΡΠ½Π΅Π½ΠΈΡ Π ΠΠΠ ΠΈ ΠΈΠ½Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΠΌΠ ΠΠ, ΠΈ, Π² ΡΠΎ ΠΆΠ΅ Π²ΡΠ΅ΠΌΡ, Π ΠΠΠ ΠΌΠΎΠΆΠ΅Ρ Π²ΠΎΡΡΡΠ°Π½Π°Π²Π»ΠΈΠ²Π°ΡΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΡ ΠΏΡΡΡΠΌ Π²ΡΡΠ΅ΡΠ½Π΅Π½ΠΈΡ Ρ50. Π‘Π»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎ, ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²ΠΎ Ρ50 ΠΈ Π ΠΠΠ , ΡΠ²ΡΠ·Π°Π½Π½ΡΡ Ρ ΠΌΠ ΠΠ, ΡΠ²Π»ΡΠ΅ΡΡΡ Π²Π°ΠΆΠ½ΡΠΌ ΡΠ°ΠΊΡΠΎΡΠΎΠΌ, ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΡΡΠΈΠΌ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΡΡ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ ΠΌΠ ΠΠ ΠΈ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΠΈΠΌ ΠΏΠ΅ΡΠ΅Ρ ΠΎΠ΄ ΠΌΠ ΠΠ ΠΈΠ· Π½Π΅Π°ΠΊΡΠΈΠ²Π½ΠΎΠ³ΠΎ Π² Π°ΠΊΡΠΈΠ²Π½ΠΎΠ΅ ΡΠΎΡΡΠΎΡΠ½ΠΈΠ΅ ΠΈ Π½Π°ΠΎΠ±ΠΎΡΠΎΡ.
ΠΡΠ»ΠΎ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ ΡΠ°ΠΊΠΆΠ΅, ΡΡΠΎ Ρ50 ΡΠ΄Π»ΠΈΠ½ΡΠ΅Ρ Π²ΡΠ΅ΠΌΡ ΠΆΠΈΠ·Π½ΠΈ ΠΌΠ ΠΠ Π² 5−10 ΡΠ°Π·, ΠΏΠΎΠ»Π½ΠΎΡΡΡΡ ΠΏΡΠ΅Π΄ΠΎΡΠ²ΡΠ°ΡΠ°Ρ Π΅Ρ Π΄Π΅Π³ΡΠ°Π΄Π°ΡΠΈΡ. ΠΠΎΠ»Π΅Π΅ ΡΠΎΠ³ΠΎ, ΡΠ΄Π°Π»Π΅Π½ΠΈΠ΅ Ρ50 ΠΈΠ· ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΡ ΡΠΊΡΡΡΠ°ΠΊΡΠΎΠ² ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΠ»ΠΎ ΠΊ ΡΡΠΈΠ»Π΅Π½ΠΈΡ Π΄Π΅Π³ΡΠ°Π΄Π°ΡΠΈΠΈ ΠΌΠ ΠΠ, ΡΠΎΠ³Π΄Π° ΠΊΠ°ΠΊ Π΄ΠΎΠ±Π°Π²Π»Π΅Π½Π½ΡΠΉ Ρ50 ΠΏΠΎΠ»Π½ΠΎΡΡΡΡ Π²ΠΎΡΡΡΠ°Π½Π°Π²Π»ΠΈΠ²Π°Π» ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΡ ΠΌΠ ΠΠ. ΠΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Ρ50 Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΠΠ΅Π¬Π° ΠΏΡΠΈΠ²Π΅Π»Π° ΠΊ ΠΈΠ½Π³ΠΈΠ±ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈ ΠΏΠΎΠ»Π½ΠΎΠΉ ΡΡΠ°Π±ΠΈΠ»ΠΈΠ·Π°ΡΠΈΠΈ ΡΠ΅ΠΏΠΎΡΡΠ΅ΡΠ½ΠΎΠΉ ΠΌΠ ΠΠ. ΠΠ°ΠΆΠ½ΠΎ ΠΎΡΠΌΠ΅ΡΠΈΡΡ ΡΠ°ΠΊΠΆΠ΅, ΡΡΠΎ Ρ50 ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°Π΅Ρ Π·Π°ΡΠΈΡΡ Π»ΡΠ±ΡΡ ΠΌΠ ΠΠ, Π½Π΅Π·Π°Π²ΠΈΡΠΈΠΌΠΎ ΠΎΡ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΈ Π½Π°Π»ΠΈΡΠΈΡ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠΎΠ² Π½Π΅ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ. Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, Ρ50 ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠ½ΠΈΠ²Π΅ΡΡΠ°Π»ΡΠ½ΡΠΌ Π±Π΅Π»ΠΊΠΎΠΌ, ΠΎΡΠ²Π΅ΡΠ°ΡΡΠΈΠΌ Π·Π° ΡΡΠ°Π±ΠΈΠ»ΠΈΠ·Π°ΡΠΈΡ ΠΈ Ρ ΡΠ°Π½Π΅Π½ΠΈΠ΅ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ ΠΠ.
Π‘ΠΊΡΠΈΠ½ΠΈΠ½Π³ Π΄Π²ΡΡ Π³ΠΈΠ±ΡΠΈΠ΄Π½ΠΎΠΉ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡΡΠ΅ΠΌΠΎΠΉ Π±ΠΈΠ±Π»ΠΈΠΎΡΠ΅ΠΊΠΈ ΠΊΠΠΠ ΠΈΠ· ΠΌΡΡΡ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° ΠΏΡΠΈΠ²ΡΠ» ΠΊ ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½ΠΈΡ Π°ΠΊΡΠΈΠ½Π° ΠΊΠ°ΠΊ Π±Π΅Π»ΠΊΠ°, Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΡΡΠ΅Π³ΠΎ Ρ Ρ50. ΠΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ Π΄Π΅Π»Π΅ΡΠΈΠΎΠ½Π½ΡΡ ΠΌΡΡΠ°Π½ΡΠΎΠ² Ρ50 ΠΏΠΎΠ·Π²ΠΎΠ»ΠΈΠ»ΠΎ Π»ΠΎΠΊΠ°Π»ΠΈΠ·ΠΎΠ²Π°ΡΡ ΡΡΠ°ΡΡΠΎΠΊ, ΠΎΡΠ²Π΅ΡΠ°ΡΡΠΈΠΉ Π·Π° ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Ρ Π°ΠΊΡΠΈΠ½ΠΎΠΌ Π² 1Π-ΠΊΠΎΠ½ΡΠ΅Π²ΠΎΠΌ Π΄ΠΎΠΌΠ΅Π½Π΅ Ρ50, Π±ΠΎΠ³Π°ΡΠΎΠΌ Π°Π»Π°Π½ΠΈΠ½ΠΎΠΌ ΠΈ ΠΏΡΠΎΠ»ΠΈΠ½ΠΎΠΌ ΠΈ ΠΎΠ±Π½Π°ΡΡΠΆΠΈΡΡ Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠ½ΡΠΉ Π°ΠΊΡΠΈΠ½-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΉ ΠΊΠΎΠ½ΡΠ΅Π½ΡΡΡ. ΠΠ°Π»ΡΠ½Π΅ΠΉΡΠΈΠ΅ ΡΠΊΡΠΏΠ΅ΡΠΈΠΌΠ΅Π½ΡΡ ΠΏΠΎΠΊΠ°Π·Π°Π»ΠΈ, ΡΡΠΎ Ρ50 Π»ΠΎΠΊΠ°Π»ΠΈΠ·ΡΠ΅ΡΡΡ ΠΏΡΠ΅ΠΈΠΌΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΠΎ Π²Π΄ΠΎΠ»Ρ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΠΌΠΈΠΊΡΠΎΡΠΈΠ±ΡΠΈΠ»Π» Π² βΠ ΠΠ’Π ΡΠΈΠ±ΡΠΎΠ±Π»Π°ΡΡΠ°Ρ ΠΌΡΡΠΈ. ΠΠΊΠ°Π·Π°Π»ΠΎΡΡ ΡΠ°ΠΊΠΆΠ΅, ΡΡΠΎ ΠΊΠΎΠ½ΡΡΠ°Π½ΡΠ° Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ50 Ρ Π°ΠΊΡΠΈΠ½ΠΎΠΌ ΡΡΠ°Π²Π½ΠΈΠΌΠ° Ρ ΡΠ°ΠΊΠΎΠ²ΡΠΌΠΈ Π΄Π»Ρ Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ ΠΏΡΠΎΡΠ½ΠΎ Π°ΠΊΡΠΈΠ½-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΡ Π±Π΅Π»ΠΊΠΎΠ². ΠΡΠΎΠΌΠ΅ ΡΠΎΠ³ΠΎ, ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Ρ50 Ρ Π°ΠΊΡΠΈΠ½ΠΎΠΌ ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΡ ΡΡΡΡΠΊΡΡΡΡ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΡΠΈΠ±ΡΠΈΠ»Π», ΡΡΠΎ ΡΠΊΠ°Π·ΡΠ²Π°Π΅Ρ Π½Π° Π°ΠΊΡΠΈΠ²Π½ΡΠΉ Ρ Π°ΡΠ°ΠΊΡΠ΅Ρ ΡΡΠΎΠ³ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ. ΠΠΎΡΠΊΠΎΠ»ΡΠΊΡ Ρ50 ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΠΌ Π±Π΅Π»ΠΊΠΎΠΌ ΠΌΠ ΠΠ ΠΈ, ΠΎΠ΄Π½ΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎ, Π°ΠΊΡΠΈΠ½-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΌ Π±Π΅Π»ΠΊΠΎΠΌ, ΠΌΠΎΠΆΠ½ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ Ρ50 ΠΈΠ³ΡΠ°Π΅Ρ Π²Π°ΠΆΠ½ΡΡ ΡΠΎΠ»Ρ Π² Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π΄Π°Π½Π½ΡΠ΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ ΠΏΠΎ-Π½ΠΎΠ²ΠΎΠΌΡ ΠΎΡΠΌΡΡΠ»ΠΈΡΡ ΡΠΎΠ»Ρ Ρ50 Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΈ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ ΠΠ. Π ΡΠ΅Π»ΠΎΠΌ, ΠΎΠΏΠΈΡΠ°Π½Π½ΡΠ΅ Π² ΡΡΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ Π½Π΅ ΠΈΠΌΠ΅ΡΡ Π°Π½Π°Π»ΠΎΠ³ΠΎΠ² Π² Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΠ΅ ΠΈ ΡΠ²Π»ΡΡΡΡΡ ΠΎΡΠ΅ΡΠ΅Π΄Π½ΡΠΌ ΡΠ°Π³ΠΎΠΌ Π² ΠΈΠ·ΡΡΠ΅Π½ΠΈΠΈ ΡΡΠ½ΠΊΡΠΈΠΉ Ρ50 Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
ΠΠ±Π·ΠΎΡ Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΡ.
Π€Π£ΠΠΠ¦ΠΠ ΠΠΠΠΠ ΠΠ«Π₯ ΠΠΠΠΠ, ΠΠ‘Π‘ΠΠ¦ΠΠΠ ΠΠΠΠΠΠ«Π₯ Π‘ ΠΌΠ ΠΠ Π ΠΠΠΠ’ΠΠΠ₯ ΠΠ£ΠΠΠ ΠΠΠ’.
ΠΠ΄Π½ΠΈΠΌ ΠΈΠ· ΠΊΠ»ΡΡΠ΅Π²ΡΡ Π·Π²Π΅Π½ΡΠ΅Π² ΡΠ΅Π°Π»ΠΈΠ·Π°ΡΠΈΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠΉ ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΈ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΏΠΎΡΡ-ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΎΠ½Π½Π°Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½ΠΎΠ². Π’Π°ΠΊ, ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²ΠΎ Π²Π°ΠΆΠ½ΡΡ ΡΠΎΠ±ΡΡΠΈΠΉ, ΠΏΡΠΎΠΈΡΡ ΠΎΠ΄ΡΡΠΈΡ ΠΏΡΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΠΈ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠ°, Π΄ΠΈΡΡΠ΅ΡΠ΅Π½ΡΠΈΡΠΎΠ²ΠΊΠ΅ ΠΊΠ»Π΅ΡΠΎΠΊ, Π΄Π° ΠΈ Π² ΠΆΠΈΠ·Π½ΠΈ ΠΊΠ»Π΅ΡΠΊΠΈ Π²ΠΎΠΎΠ±ΡΠ΅, ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΠ΅ΡΡΡ ΠΏΠΎΡΡ-ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΎΠ½Π½ΡΠΌΠΈ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ°ΠΌΠΈ, ΡΠ΅Π³ΡΠ»ΠΈΡΡΡΡΠΈΠΌΠΈ ΠΏΡΠΎΡΠ΅ΡΡΠΈΠ½Π³ ΠΌΠ ΠΠ Π² ΡΠ΄ΡΠ΅, Π° ΡΠ°ΠΊΠΆΠ΅ Π΅Ρ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΡ, Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ ΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΡ Π² ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ΅.
ΠΠ΄Π½ΠΈΠΌ ΠΈΠ· Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ Ρ ΠΎΡΠΎΡΠΎ ΠΈΠ·ΡΡΠ΅Π½Π½ΡΡ ΠΌΠΎΠΌΠ΅Π½ΡΠΎΠ² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΌΠ ΠΠ ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ Π±Π΅Π»ΠΊΠΎΠ²ΠΎΠ³ΠΎ ΡΠΈΠ½ΡΠ΅Π·Π° (ΡΠΌ. ΠΎΠ±Π·ΠΎΡ Pain, 1996; Day and Tuite, 1998; Gray and Wickens, 1998; Preiss and Hentze, 1999). Π ΡΠ°ΡΡΠ½ΠΎΡΡΠΈ, ΠΏΠΎΠΊΠ°Π·Π°Π½Π° Π²Π°ΠΆΠ½Π°Ρ ΡΠΎΠ»Ρ Π² ΡΡΠΎΠΌ ΠΏΡΠΎΡΠ΅ΡΡΠ΅ ΡΠ°ΠΊΡΠΎΡΠ° eIF4G, ΠΊΠΎΡΠΎΡΡΠΉ ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΏΠ»Π°ΡΡΠΎΡΠΌΠΎΠΉ Π΄Π»Ρ ΡΠ±ΠΎΡΠΊΠΈ 48 S ΠΏΡΠ΅ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ° Π½Π° ΠΌΠ ΠΠ. eIF4G ΠΌΠΎΠΆΠ΅Ρ ΡΠ²ΡΠ·ΡΠ²Π°ΡΡ ΠΊΡΠΏ-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΉ Π±Π΅Π»ΠΎΠΊ eIF4E, Π ΠΠ-Ρ Π΅Π»ΠΈΠΊΠ°Π·Ρ eIF4A ΠΈ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠΈΠ²Π°ΡΡ ΡΠ΅ΠΌ ΡΠ°ΠΌΡΠΌ ΠΌΡΠ»ΡΡΠΈΠΌΠ΅ΡΠ½ΡΠΉ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ Π½Π°Π·Π²Π°Π½Π½ΡΠΉ eIF4 °F. ΠΡΠΎΡ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌ Π΄Π»Ρ ΡΠ·Π½Π°Π²Π°Π½ΠΈΡ ΠΊΡΠΏ-ΡΡΡΡΠΊΡΡΡΡ ΠΈ ΡΠΊΠ°Π½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΌΠ ΠΠ Π΄ΠΎ ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΠΊΠΎΠ΄ΠΎΠ½Π°. ΠΡΠΎΠΌΠ΅ ΡΠΎΠ³ΠΎ eIF4G ΠΌΠΎΠΆΠ΅Ρ ΡΠ²ΡΠ·ΡΠ²Π°ΡΡ eEF3 ΠΈ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡ ΠΏΠΎΡΠ°Π΄ΠΊΡ 43 S ΠΏΡΠ΅ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ° Π½Π° ΠΌΠ ΠΠ (ΡΠΌ. ΠΎΠ±Π·ΠΎΡ Pain, 1996; Day and Tuite, 1998; Gray and Wickens, 1998; Preiss and Hentze, 1999). Π‘Π°ΠΌΡΠΌ Π½Π΅ΠΎΠΆΠΈΠ΄Π°Π½Π½ΡΠΌ ΠΎΡΠΊΡΡΡΠΈΠ΅ΠΌ ΡΠ²ΠΈΠ»ΠΎΡΡ ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½ΠΈΠ΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ eIF4G Ρ Π ΠΠΠ , ΠΏΡΠΈΠ²ΠΎΠ΄ΡΡΠ΅Π΅ ΠΊ ΡΠΈΡΠΊΡΠ»ΡΡΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ ΠΈ ΠΎΠ±Π»Π΅Π³ΡΠ΅Π½ΠΈΡ ΡΠ΅ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ (Gallie and Tanguay, 1994; Wells et al., 1998). Π‘ΠΎΠ²Π΅ΡΡΠ΅Π½Π½ΠΎ Π½ΠΎΠ²ΡΠΉ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌ Π²Π½ΡΡΡΠ΅Π½Π½Π΅ΠΉ ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ, ΡΠ°Π½Π΅Π΅ ΠΎΠΏΠΈΡΠ°Π½Π½ΡΠΉ Ρ ΠΏΠΈΠΊΠΎΡΠ½Π°Π²ΠΈΡΡΡΠ½ΡΡ ΠΌΠ ΠΠ, Π±ΡΠ» ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½ Ρ ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²Π° ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ ΠΠ, ΠΊΠΎΠ΄ΠΈΡΡΡΡΠΈΡ , Π² ΠΎΡΠ½ΠΎΠ²Π½ΠΎΠΌ, ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ (Nanbru et al., 1997; Pyronnet et al., 2000). ΠΡΠ»ΠΈ ΡΠ°ΠΊΠΆΠ΅ Π΄Π΅ΡΠ°Π»ΡΠ½ΠΎ ΠΈΠ·ΡΡΠ΅Π½Ρ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΡ Π³Π»ΠΎΠ±Π°Π»ΡΠ½ΠΎΠΉ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΏΡΡΠ΅ΠΌ ΡΠΎΡΡΠΎΡΠΈΠ»ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΊΠ»ΡΡΠ΅Π²ΡΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ² ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ eIF2a, eIF4E ΠΈ 4Π-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΡ Π±Π΅Π»ΠΊΠΎΠ², Π° ΡΠ°ΠΊΠΆΠ΅ ΡΠ°ΡΡΠ΅ΠΏΠ»Π΅Π½ΠΈΡ eIF4GΠΏΠΎΠΊΠ°Π·Π°Π½Π° ΡΠΎΠ»Ρ ΡΡΠΈΡ ΠΌΠΎΠ΄ΠΈΡΠΈΠΊΠ°ΡΠΈΠΉ Π² ΠΌΠΎΠ΄ΡΠ»ΡΡΠΈΠΈ ΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ²Π΅ΡΠ° Π½Π° ΡΠ°Π·Π»ΠΈΡΠ½ΡΠ΅ ΡΠΈΠ·ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΡΠΈΠΌΡΠ»Ρ ΠΈ ΡΡΡΠ΅ΡΡΠΎΠ²ΡΠ΅ ΡΡΠ»ΠΎΠ²ΠΈΡ (ΡΠΌ. ΠΎΠ±Π·ΠΎΡ Pain, 1996; Day and Tuite, 1998; Gray and Wickens, 1998; Gingras et al., 1999; Preiss and Hentze, 1999).
ΠΡΠΎΠΌΠ΅ ΡΠΎΠ³ΠΎ, ΠΏΠΎΡΠ²ΠΈΠ»ΠΎΡΡ ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²ΠΎ ΠΏΡΠΈΠΌΠ΅ΡΠΎΠ² ΡΠΎΠ³ΠΎ, ΡΡΠΎ c/s-ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠ΅ ΡΠΈΠ³Π½Π°Π»Ρ, ΡΠΎΡΡΠ΅Π΄ΠΎΡΠΎΡΠ΅Π½Π½ΡΠ΅ Π² 5'- ΠΈ 3'- Π½Π΅ΡΡΠ°Π½ΡΠ»ΠΈΡΡΠ΅ΠΌΡΡ ΠΎΠ±Π»Π°ΡΡΡΡ , ΠΈ ΡΠ·Π½Π°ΡΡΠΈΠ΅ ΠΈΡ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ ΠΌΠΎΠ³ΡΡ ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΡΡ ΠΎΡΠΎΠ±Π΅Π½Π½ΠΎΡΡΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΠ½Π½ΡΡ ΠΊΠ»Π°ΡΡΠΎΠ² ΠΌΠ ΠΠ. ΠΡΠ»Π° ΠΏΠΎΠΊΠ°Π·Π°Π½Π° Π²Π°ΠΆΠ½ΠΎΡΡΡ ΡΡΠΈΡ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ Π ΠΠ-Π±Π΅Π»ΠΊΠΎΠ²ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠΈΠ²ΠΉ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ, ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ ΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²Π° ΠΌΠ ΠΠ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ², Π° ΡΠ°ΠΊΠΆΠ΅ ΠΌΠ ΠΠ, Π°ΡΡΠΎΡΠΈΠΈΡΠΎΠ²Π°Π½Π½ΡΡ Ρ ΡΠΎΡΡΠΎΠΌ ΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΠ΅ΠΌ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠ° (ΡΠΌ. ΠΎΠ±Π·ΠΎΡ Spirin, 1994; Gray and Wickens, 1998). ΠΠΌΠ΅ΡΡΠ΅ Ρ ΡΠ΅ΠΌ, ΠΏΠΎΡΠ²Π»ΡΠ΅ΡΡΡ Π²ΡΡ Π±ΠΎΠ»ΡΡΠ΅ Π΄Π°Π½Π½ΡΡ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ ΡΡΡΠ΅ΡΡΠ²ΡΠ΅Ρ Π³ΡΡΠΏΠΏΠ° Π±Π΅Π»ΠΊΠΎΠ², ΡΠΎΠΏΡΠΎΠ²ΠΎΠΆΠ΄Π°ΡΡΠΈΡ ΠΌΠ ΠΠ Π½Π° Π²ΡΠ΅Ρ ΡΡΠ°ΠΏΠ°Ρ Π΅Ρ Π±ΠΈΠΎΠ³Π΅Π½Π΅Π·Π° ΠΈ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΠΈΡ ΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΡΡ ΡΠ²ΡΠ·Ρ ΠΌΠ΅ΠΆΠ΄Ρ «ΡΠ΄Π΅ΡΠ½ΠΎΠΉ ΠΈΡΡΠΎΡΠΈΠ΅ΠΉ» ΠΌΠ ΠΠ ΠΈ Π΅Ρ Π΄Π°Π»ΡΠ½Π΅ΠΉΡΠ΅ΠΉ ΡΡΠ΄ΡΠ±ΠΎΠΉ Π² ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ΅. Π’ΡΠΈ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ ΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Π° Π±Π΅Π»ΠΊΠΎΠ²: Π±Π΅Π»ΠΊΠΈ Π³Π΅ΡΠ΅ΡΠΎΠ³Π΅Π½Π½ΡΡ ΡΠ΄Π΅ΡΠ½ΡΡ Π ΠΠ (Π³ΡΠ ΠΠ), Y-box ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠ΅ Π±Π΅Π»ΠΊΠΈ (YB Π±Π΅Π»ΠΊΠΈ) ΠΈ Π ΠΠΠ , Π±ΡΠ»ΠΈ ΠΎΠΏΠΈΡΠ°Π½Ρ ΠΊΠ°ΠΊ ΠΊΠ»ΡΡΠ΅Π²ΡΠ΅ ΡΠ°ΠΊΡΠΎΡΡ ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΠ΅ ΠΏΡΠΎΡΠ΅ΡΡΠΈΠ½Π³ ΠΌΠ ΠΠ Π² ΡΠ΄ΡΠ΅, Π° ΡΠ°ΠΊΠΆΠ΅ ΡΡΠ°Π½ΡΠΏΠΎΡΡ ΠΈ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ, ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΡΡ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ ΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΡ Π±ΠΎΠ»ΡΡΠΈΠ½ΡΡΠ²Π° ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ ΠΠ (ΡΠΌ. ΠΎΠ±Π·ΠΎΡ Caponigro and Parker, 1995; Sommerville and Ladomery, 1996bSpirin, 1996; Matsumoto and Wolffe, 1998; Sommerville, 1999; Shyu and Wilkinson, 2000). ΠΡΠΈ Π±Π΅Π»ΠΊΠΈ ΡΠ²Π»ΡΡΡΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΠΌΠΈ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΠ°ΠΌΠΈ ΠΌΠ ΠΠ ΠΈ ΠΎΠ±Π»Π°Π΄Π°ΡΡ ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡΡ ΠΈΠ·ΠΌΠ΅Π½ΡΡΡ ΠΏΡΠΎΡΡΡΠ°Π½ΡΡΠ²Π΅Π½Π½ΡΡ ΡΡΡΡΠΊΡΡΡΡ ΠΌΠ ΠΠ. ΠΠΎ-Π²ΠΈΠ΄ΠΈΠΌΠΎΠΌΡ, ΠΈΠΌΠ΅Π½Π½ΠΎ ΡΡΠΎ ΡΠ²ΠΎΠΉΡΡΠ²ΠΎ Π»Π΅ΠΆΠΈΡ Π² ΠΎΡΠ½ΠΎΠ²Π΅ ΠΈΡ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ.
Π ΡΡΠΎΠΌ ΠΎΠ±Π·ΠΎΡΠ΅ ΠΌΡ ΠΎΠ³ΡΠ°Π½ΠΈΡΠΈΠ»ΠΈΡΡ ΠΊΡΠ°ΡΠΊΠΈΠΌ ΠΏΠ΅ΡΠ΅ΡΠΈΡΠ»Π΅Π½ΠΈΠ΅ΠΌ Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ Π²Π°ΠΆΠ½ΡΡ Π½Π°ΠΏΡΠ°Π²Π»Π΅Π½ΠΈΠΉ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ ΠΏΡΡΠ΅ΠΉ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΠΌΠ΅ΡΠ°Π±ΠΎΠ»ΠΈΠ·ΠΌΠ° ΠΌΠ ΠΠ ΠΈ ΡΠΊΠΎΠ½ΡΠ΅Π½ΡΡΠΈΡΠΎΠ²Π°Π»ΠΈΡΡ Π² ΠΎΡΠ½ΠΎΠ²Π½ΠΎΠΌ Π½Π° ΡΡΠ½ΠΊΡΠΈΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ² ΠΌΠ ΠΠ, ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΡΡΠΈΡ ΡΡΡΡΠΊΡΡΡΠ½ΡΡ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΡ, ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΡΡ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ ΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΡ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
ΠΡΠ²ΠΎΠ΄Ρ.
1. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ Ρ50 ΠΈ Π ΠΠΠ ΠΎΠ±Π»Π°Π΄Π°ΡΡ ΠΎΠ΄ΠΈΠ½Π°ΠΊΠΎΠ²ΡΠΌ ΡΡΠΎΠ΄ΡΡΠ²ΠΎΠΌ ΠΊ ΠΏΠΎΠ»ΠΈ (Π) ΠΈ ΠΊΠΎΠ½ΠΊΡΡΠΈΡΡΡΡ Π·Π° ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Ρ ΠΏΠΎΠ»ΠΈ (Π)-Ρ Π²ΠΎΡΡΠΎΠΌ ΠΌΠ ΠΠ. Π‘Π²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ Ρ50 Ρ ΠΏΠΎΠ»ΠΈ (Π)-Ρ Π²ΠΎΡΡΠΎΠΌ ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ ΠΈΠ½Π°ΠΊΡΠΈΠ²Π°ΡΠΈΡ ΠΌΠ ΠΠ. Π ΡΠ²ΠΎΡ ΠΎΡΠ΅ΡΠ΅Π΄Ρ, Π ΠΠΠ ΠΌΠΎΠΆΠ΅Ρ Π²ΡΡΠ΅ΡΠ½ΡΡΡ Ρ50 ΠΈ Π²ΠΎΡΡΡΠ°Π½Π°Π²Π»ΠΈΠ²Π°ΡΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΡ ΠΏΠΎΠ»ΠΈΠ°Π΄Π΅Π½ΠΈΠ»ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ ΠΌΠ ΠΠ. ΠΠΎΠ»Π΅Π΅ ΡΠΎΠ³ΠΎ, ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΡ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ ΡΠΎΠΏΡΠΎΠ²ΠΎΠΆΠ΄Π°Π΅ΡΡΡ ΡΠ²Π΅Π»ΠΈΡΠ΅Π½ΠΈΠ΅ΠΌ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²Π° Ρ50 Π² ΡΠΎΡΡΠ°Π²Π΅ ΠΌΠ ΠΠ. Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΡΠΎΠΎΡΠ½ΠΎΡΠ΅Π½ΠΈΡ Π΄Π²ΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ², Ρ50 ΠΈ Π ΠΠΠ , Π² ΡΠΎΡΡΠ°Π²Π΅ ΠΌΠ ΠΠ ΠΌΠΎΠΆΠ΅Ρ ΡΠ²Π»ΡΡΡΡΡ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠΎΠΌ Π³Π»ΠΎΠ±Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠ³ΠΎ ΠΊΠΎΠ½ΡΡΠΎΠ»Ρ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
2. ΠΠ±Π½Π°ΡΡΠΆΠ΅Π½ΠΎ, ΡΡΠΎ Ρ50 ΡΠ΄Π»ΠΈΠ½ΡΠ΅Ρ Π²ΡΠ΅ΠΌΡ ΠΆΠΈΠ·Π½ΠΈ ΠΌΠ ΠΠ in vitro ΠΈ in vivo Π±ΠΎΠ»Π΅Π΅, ΡΠ΅ΠΌ Π² 100 ΡΠ°Π·, ΡΠΎΠ³Π΄Π° ΠΊΠ°ΠΊ Π΅Π³ΠΎ ΡΠ΄Π°Π»Π΅Π½ΠΈΠ΅ ΠΈΠ· ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΡ ΡΠΊΡΡΡΠ°ΠΊΡΠΎΠ² ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ Π΄Π΅ΡΡΠ°Π±ΠΈΠ»ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ Π΄Π²Π΅ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ, ΡΡΠ°Π±ΠΈΠ»ΠΈΠ·Π°ΡΠΈΡ ΠΌΠ ΠΠ ΠΈ ΠΈΠ½Π³ΠΈΠ±ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ, Π»ΠΎΠΊΠ°Π»ΠΈΠ·ΠΎΠ²Π°Π½Ρ Π² ΡΠ°Π·Π½ΡΡ Π΄ΠΎΠΌΠ΅Π½Π°Ρ Ρ50: Π΄ΠΎΠΌΠ΅Π½ Ρ ΠΎΠ»ΠΎΠ΄ΠΎΠ²ΠΎΠ³ΠΎ ΡΠΎΠΊΠ° ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°Π΅Ρ ΡΡΠ°Π±ΠΈΠ»ΠΈΠ·Π°ΡΠΈΡ ΠΌΠ ΠΠ, Π² ΡΠΎ Π²ΡΠ΅ΠΌΡ ΠΊΠ°ΠΊ Π‘-ΠΊΠΎΠ½ΡΠ΅Π²ΠΎΠΉ Π΄ΠΎΠΌΠ΅Π½ ΠΎΡΠ²Π΅ΡΠ°Π΅Ρ Π·Π° ΠΈΠ½Π³ΠΈΠ±ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΈ. Ρ50 ΠΈΠ·Π±ΠΈΡΠ°ΡΠ΅Π»ΡΠ½ΠΎ Π·Π°ΡΠΈΡΠ°Π΅Ρ ΠΏΠΎΠ»ΠΈ (Π)-Ρ Π²ΠΎΡΡ, Π½ΠΎ Π½Π΅ «ΡΠ΅Π»ΠΎ» ΠΌΠ ΠΠ ΠΎΡ Π΄Π΅ΠΉΡΡΠ²ΠΈΡ ΡΠ½Π΄ΠΎΡΠΈΠ±ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°Ρ, ΠΏΠΎ-Π²ΠΈΠ΄ΠΈΠΌΠΎΠΌΡ, Π·Π°ΡΠΈΡΡ ΠΌΠ ΠΠ ΠΎΡ Π΄Π΅Π°Π΄Π΅Π½ΠΈΠ»ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΈ ΠΏΠΎΡΠ»Π΅Π΄ΡΡΡΠ΅ΠΉ Π΄Π΅Π³ΡΠ°Π΄Π°ΡΠΈΠΈ. Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, Ρ50 ΠΌΠΎΠΆΠ΅Ρ ΡΠ²Π»ΡΡΡΡΡ ΡΠ½ΠΈΠ²Π΅ΡΡΠ°Π»ΡΠ½ΡΠΌ ΡΠ°ΠΊΡΠΎΡΠΎΠΌ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΠΈΠΌ ΠΎΠ±ΡΡΡ ΡΡΠ°Π±ΠΈΠ»ΠΈΠ·Π°ΡΠΈΡ ΠΈ Ρ ΡΠ°Π½Π΅Π½ΠΈΠ΅ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
3. ΠΠΊΡΠΈΠ½ Π±ΡΠ» ΠΈΠ΄Π΅Π½ΡΠΈΡΠΈΡΠΈΡΠΎΠ²Π°Π½ ΠΈ ΠΎΡ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΠ·ΠΎΠ²Π°Π½ ΠΊΠ°ΠΊ Π±Π΅Π»ΠΎΠΊ, Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΡΡΠΈΠΉ Ρ Ρ50. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ Ρ50 ΡΠΎΠ΄Π΅ΡΠΆΠΈΡ Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠ½Π½ΡΠΉ Π°ΠΊΡΠΈΠ½-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΉ ΠΌΠΎΡΠΈΠ² ΠΈ Π»ΠΎΠΊΠ°Π»ΠΈΠ·ΡΠ΅ΡΡΡ Π² βΠΠ£ΠΠ’Π ΡΠΈΠ±ΡΠΎΠ±Π»Π°ΡΡΠ°Ρ Π²Π΄ΠΎΠ»Ρ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΠΌΠΈΠΊΡΠΎΡΠΈΠ»Π°ΠΌΠ΅Π½ΡΠΎΠ². Ρ50 ΠΎΠ±Π»Π°Π΄Π°Π΅Ρ Π²ΡΡΠΎΠΊΠΈΠΌ ΡΡΠΎΠ΄ΡΡΠ²ΠΎΠΌ ΠΊ ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ½ΠΎΠΌΡ Π°ΠΊΡΠΈΠ½Ρ ΠΈ Π΅Π³ΠΎ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΠ΅ ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ Π·Π½Π°ΡΠΈΡΠ΅Π»ΡΠ½ΠΎΠΌΡ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΡ ΡΡΡΡΠΊΡΡΡΡ Π°ΠΊΡΠΈΠ½Π°. ΠΡΠΈΠ½ΠΈΠΌΠ°Ρ Π²ΠΎ Π²Π½ΠΈΠΌΠ°Π½ΠΈΠ΅, ΡΡΠΎ Ρ50 ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΌΠ°ΠΆΠΎΡΠ½ΡΠΌ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΠΎΠΌ ΠΌΠ ΠΠ ΠΈ, ΠΎΠ΄Π½ΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎ, Π°ΠΊΡΠΈΠ½-ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΌ Π±Π΅Π»ΠΊΠΎΠΌ, ΠΌΠΎΠΆΠ½ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ Ρ50 ΠΈΠ³ΡΠ°Π΅Ρ Π²Π°ΠΆΠ½ΡΡ ΡΠΎΠ»Ρ Π² Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΠΌΠ ΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
ΠΠ»Π°Π³ΠΎΠ΄Π°ΡΠ½ΠΎΡΡΠΈ.
ΠΡΠ° ΡΠ°Π±ΠΎΡΠ° Π±ΡΠ»Π° ΡΠ΄Π΅Π»Π°Π½Π° Ρ ΠΏΠΎΠΌΠΎΡΡΡ ΠΌΠ½ΠΎΠ³ΠΈΡ Π»ΡΠ΄Π΅ΠΉ, ΠΈΡ ΡΠ½ΡΡΠ·ΠΈΠ°Π·ΠΌ, ΠΈΠ½ΡΠ΅ΡΠ΅Ρ, ΠΈΠ΄Π΅ΠΈ Π»Π΅ΠΆΠ°Ρ Π² ΠΎΡΠ½ΠΎΠ²Π΅ ΡΡΠΎΠΉ Π΄ΠΈΡΡΠ΅ΡΡΠ°ΡΠΈΠΈ.
ΠΡΠ΅ΠΆΠ΄Π΅ Π²ΡΠ΅Π³ΠΎ, Ρ Π±Π»Π°Π³ΠΎΠ΄Π°ΡΠ΅Π½ ΡΠ²ΠΎΠ΅ΠΉ Π»ΡΠ±ΠΈΠΌΠΎΠΉ ΠΆΠ΅Π½Π΅ ΠΈ Π·Π°ΠΌΠ΅ΡΠ°ΡΠ΅Π»ΡΠ½ΠΎΠΌΡ ΠΊΠΎΠ»Π»Π΅Π³Π΅ ΠΠ°Π»Π΅ ΠΠ²Π΄ΠΎΠΊΠΈΠΌΠΎΠ²ΠΎΠΉ, ΡΠΎΠ²ΠΌΠ΅ΡΠ½ΠΎ Ρ ΠΊΠΎΡΠΎΡΠΎΠΉ Π±ΡΠ»ΠΈ ΡΠ΄Π΅Π»Π°Π½Ρ ΠΏΡΠ°ΠΊΡΠΈΡΠ΅ΡΠΊΠΈ Π²ΡΠ΅ ΡΠΊΡΠΏΠ΅ΡΠΈΠΌΠ΅Π½ΡΡ, ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ ΠΊΠΎΡΠΎΡΡΡ ΠΎΠΏΠΈΡΠ°Π½Ρ Π² ΡΡΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅. Π― ΠΌΠΎΠ³Ρ Ρ ΡΠ²Π΅ΡΠ΅Π½Π½ΠΎΡΡΡΡ ΡΠΊΠ°Π·Π°ΡΡ, ΡΡΠΎ Π±Π΅Π· Π΅Ρ ΠΏΠΎΠΌΠΎΡΠΈ ΡΡΠ° ΡΠ°Π±ΠΎΡΠ° Π½ΠΈΠΊΠΎΠ³Π΄Π° Π±Ρ Π½Π΅ Π±ΡΠ»Π° ΡΠ΄Π΅Π»Π°Π½Π° ΠΈ Π½Π°ΠΏΠΈΡΠ°Π½Π°, ΠΏΠΎ ΠΊΡΠ°ΠΉΠ½Π΅ΠΉ ΠΌΠ΅ΡΠ΅ — Π² ΡΠΎΠΌ Π²ΠΈΠ΄Π΅, Π² ΠΊΠ°ΠΊΠΎΠΌ ΠΎΠ½Π° ΡΠ΅ΠΉΡΠ°Ρ Π½Π°Ρ ΠΎΠ΄ΠΈΡΡΡ.
Π― ΡΠ°ΠΊΠΆΠ΅ Π±Π»Π°Π³ΠΎΠ΄Π°ΡΠ΅Π½ ΠΌΠΎΠ΅ΠΌΡ ΡΡΠΊΠΎΠ²ΠΎΠ΄ΠΈΡΠ΅Π»Ρ, ΠΡΠ²Ρ ΠΠ°Π²Π»ΠΎΠ²ΠΈΡΡ ΠΠ²ΡΠΈΠ½Π½ΠΈΠΊΠΎΠ²Ρ, Π·Π° ΡΠΊΠΎΠ»Ρ Π½Π°ΡΡΠ½ΡΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ ΠΈ ΡΠΊΠΎΠ»Ρ ΠΆΠΈΠ·Π½ΠΈ. ΠΠ½ ΡΠΎΠ·Π΄Π°Π» Π² Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ Π°ΡΠΌΠΎΡΡΠ΅ΡΡ Π²Π·Π°ΠΈΠΌΠ½ΠΎΠ³ΠΎ Π²Π½ΠΈΠΌΠ°Π½ΠΈΡ, ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠΊΠΈ ΠΈ ΡΠ²ΠΎΠ±ΠΎΠ΄Ρ ΡΠ²ΠΎΡΡΠ΅ΡΡΠ²Π°, Π±Π΅Π· ΡΠ΅Π³ΠΎ Π²ΡΠΏΠΎΠ»Π½Π΅Π½ΠΈΠ΅ ΡΠ°Π±ΠΎΡΡ Π±ΡΠ»ΠΎ Π±Ρ Π½Π΅Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΡΠΌ. Π― ΡΠ°ΠΊΠΆΠ΅ Π±Π»Π°Π³ΠΎΠ΄Π°ΡΠ΅Π½ Π²ΡΠ΅ΠΌΡ ΠΊΠΎΠ»Π»Π΅ΠΊΡΠΈΠ²Ρ Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ ΠΡΠ²Π° ΠΠ°Π²Π»ΠΎΠ²ΠΈΡΠ°.
Π’Π°ΠΊΠΆΠ΅ Ρ ΠΎΡΡ ΠΏΠΎΠ±Π»Π°Π³ΠΎΠ΄Π°ΡΠΈΡΡ ΡΠ²ΠΎΠ΅Π³ΠΎ Π±ΡΠ°ΡΠ°, ΠΠ°ΠΊΡΠΈΠΌΠ° Π ΡΠ·Π°Π½ΠΎΠ²Π° Π·Π° Π½Π΅ΠΎΡΠ΅Π½ΠΈΠΌΡΡ ΠΏΠΎΠΌΠΎΡΡ Π² ΠΏΠΎΠ΄Π³ΠΎΡΠΎΠ²ΠΊΠ΅ ΠΊ ΠΏΠ΅ΡΠ°ΡΠΈ Π°Π²ΡΠΎΡΠ΅ΡΠ΅ΡΠ°ΡΠ° ΠΌΠΎΠ΅ΠΉ Π΄ΠΈΡΡΠ΅ΡΡΠ°ΡΠΈΠΎΠ½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅ ΠΈ Π·Π° ΠΏΠΎΠΌΠΎΡΡ Π² ΠΎΡΠΎΡΠΌΠ»Π΅Π½ΠΈΠΈ ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²Π° Π±ΡΠΌΠ°Π³, Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΡΡ Π΄Π»Ρ ΠΏΡΠΎΡΠ΅ΡΡΠ° Π·Π°ΡΠΈΡΡ.
Π§Π°ΡΡΡ ΡΡΠΎΠΉ ΡΠ°Π±ΠΎΡΡ Π±ΡΠ»Π° Π²ΡΠΏΠΎΠ»Π½Π΅Π½Π° Π² Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ ΠΠ°ΡΠΌΠ° ΠΠΎΠ½Π΅Π½Π±Π΅ΡΠ³Π° Π² ΠΌΠΎΠ½ΡΠ΅Π°Π»ΡΡΠΊΠΎΠΌ ΡΠ½ΠΈΠ²Π΅ΡΡΠΈΡΠ΅ΡΠ΅ ΠΡ (ΠΠ³Π¨. Π― ΠΏΡΠΈΠ·Π½Π°ΡΠ΅Π»Π΅Π½ Π΅ΠΌΡ Π·Π° ΠΏΡΠ΅Π΄ΠΎΡΡΠ°Π²Π»Π΅Π½Π½ΡΡ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡΡ ΡΠ°Π±ΠΎΡΠ°ΡΡ Π² Π΅Π³ΠΎ Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ. ΠΠ»Π°Π³ΠΎΠ΄Π°ΡΡ ΡΡΡΠ΄Π΅Π½ΡΠΎΠ² Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ ΠΠΎΠ½Π΅Π½Π±Π΅ΡΠ³Π°, Π·Π° ΠΊΠΎΠ½ΡΡΠ»ΡΡΠ°ΡΠΈΠΈ ΠΈ ΠΏΠΎΠΌΠΎΡΡ Π² ΡΠ°Π±ΠΎΡΠ΅.
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