ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠ°Ρ Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΡΡΠΈΠΊΠ° ΠΌΡΡΠ°ΡΠΈΠΉ Π³Π΅Π½Π° Trithorax-like ΠΈ ΠΈΡ Π²Π»ΠΈΡΠ½ΠΈΠ΅ Π½Π° ΠΎΠΎΠ³Π΅Π½Π΅Π· Drosophila melanogaster
ΠΠΏΠ΅ΡΠ²ΡΠ΅ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½Π° Trl ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΡ ΡΡΡΡΠΊΡΡΡΡ ΡΠΈΡΠ½ΠΈΠΊΠΎΠ² ΠΌΡΡ . ΠΠ°ΡΡΡΠ΅Π½ΠΈΡ Π²ΡΡΠ²Π»ΡΡΡΡΡ ΠΊΠ°ΠΊ Π² ΠΎΠ²Π°ΡΠΈΠΎΠ»Π°Ρ , ΡΠ°ΠΊ ΠΈ Π² ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΡ ΡΠΉΡΠ΅Π²ΡΡ ΠΊΠ°ΠΌΠ΅ΡΠ°Ρ . Π£ΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ 7Π³/-ΠΌΡΡΠ°ΡΠΈΠΈ Π²ΡΠ·ΡΠ²Π°ΡΡ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΡΠΈΡΠ»Π° ΠΏΠΎΠ»ΠΎΠ²ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ (ΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΈ ΠΎΠΎΡΠΈΡΠΎΠ²) Π² ΡΠΉΡΠ΅Π²ΡΡ ΠΊΠ°ΠΌΠ΅ΡΠ°Ρ ΠΏΠΎ ΡΡΠ°Π²Π½Π΅Π½ΠΈΡ Ρ Π½ΠΎΡΠΌΠΎΠΉ. ΠΠΏΠ΅ΡΠ²ΡΠ΅ Ρ 7Π³/-ΠΌΡΡΠ°Π½ΡΠΎΠ² Π²ΡΡΠ²Π»Π΅Π½Ρ Π΄Π΅ΡΠ΅ΠΊΡΡ Π² ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΡΠ°Π΄ΠΈΠΎΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ … 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 1. ΠΠΠΠΠ ΠΠΠ’ΠΠ ΠΠ’Π£Π Π«
- 1. 1. ΠΡΠ½ΠΎΠ²Π½ΡΠ΅ Π°ΡΠΏΠ΅ΠΊΡΡ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ ΡΠ°ΠΌΠΎΠΊ Drosophila melanogaster
- 1. 1. 1. Π€ΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ ΡΠ°ΠΌΠΎΠΊ Π² ΡΠΌΠ±ΡΠΈΠΎΠ³Π΅Π½Π΅Π·Π΅
- 1. 1. 2. Π Π°Π·Π²ΠΈΡΠΈΠ΅ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ ΡΠ°ΠΌΠΎΠΊ Ρ Π»ΠΈΡΠΈΠ½ΠΎΠΊ ΠΈ ΠΏΡΠ΅Π΄ΠΊΡΠΊΠΎΠ»ΠΎΠΊ
- 1. 1. 3. Π Π°Π·Π²ΠΈΡΠΈΠ΅ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Ρ Π²Π·ΡΠΎΡΠ»ΡΡ ΡΠ°ΠΌΠΎΠΊ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ
- 1. 1. 4. Π Π°Π·Π²ΠΈΡΠΈΠ΅ ΡΠΉΡΠ΅Π²ΠΎΠΉ ΠΊΠ°ΠΌΠ΅ΡΡ ΠΏΠΎΡΠ»Π΅ Π²ΡΡ ΠΎΠ΄Π° ΠΈΠ· Π³Π΅ΡΠΌΠ°ΡΠΈΡ
- 1. 1. 5. Π’ΡΠ°Π½ΡΠΏΠΎΡΡ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΡ ΠΏΠΈΡΠ°ΡΡΠΈΡ
ΠΊΠ»Π΅ΡΠΎΠΊ Π² ΠΎΠΎΡΠΈΡ (nurse cell — oocyte transport)
- 1. 1. 5. 1. Π‘Π΅Π»Π΅ΠΊΡΠΈΠ²Π½ΡΠΉ ΡΡΠ°Π½ΡΠΏΠΎΡΡ
- 1. 1. 5. 2. ΠΡΡΡΡΡΠΉ ΡΡΠ°Π½ΡΠΏΠΎΡΡ
- 1. 1. 5. 3. Π‘ΡΡΡΠΊΡΡΡΠ° Π°ΠΊΡΠΈΠ½ΠΎΠ²ΠΎΠ³ΠΎ ΡΠΈΡΠΎΡΠΊΠ΅Π»Π΅ΡΠ° Π² ΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΊΠ»Π΅ΡΠΊΠ°Ρ Π²ΠΎ Π²ΡΠ΅ΠΌΡ Π±ΡΡΡΡΠΎΠ³ΠΎ ΡΡΠ°Π½ΡΠΏΠΎΡΡΠ°
- 1. 1. 6. ΠΠ²ΠΎΠ»ΡΡΠΈΠΎΠ½Π½Π°Ρ ΠΊΠΎΠ½ΡΠ΅ΡΠ²Π°ΡΠΈΠ²Π½ΠΎΡΡΡ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π°
- 1. 2. ΠΠ΅Π½ Trithorax-like: Π΅Π³ΠΎ ΡΠΎΠ»Ρ ΠΈ Π€ΡΠ½ΠΊΡΠΈΠΈ
- 1. 2. 1. ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½Π°Ρ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΡ ΠΈ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Π³Π΅Π½Π° Til
- 1. 2. 2. Π‘ΡΡΡΠΊΡΡΡΠ° 5'-ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ Π³Π΅Π½Π° Π’Π³
- 1. 2. 3. ΠΠΈΠΏΠΎΠΌΠΎΡΡΠ½ΡΠ΅ ΠΌΡΡΠ°ΡΠΈΠΈ Π³Π΅Π½Π° Π’ (
- 1. 1. ΠΡΠ½ΠΎΠ²Π½ΡΠ΅ Π°ΡΠΏΠ΅ΠΊΡΡ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ ΡΠ°ΠΌΠΎΠΊ Drosophila melanogaster
ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠ°Ρ Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΡΡΠΈΠΊΠ° ΠΌΡΡΠ°ΡΠΈΠΉ Π³Π΅Π½Π° Trithorax-like ΠΈ ΠΈΡ Π²Π»ΠΈΡΠ½ΠΈΠ΅ Π½Π° ΠΎΠΎΠ³Π΅Π½Π΅Π· Drosophila melanogaster (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
ΠΠΎΠ³Π΅Π½Π΅Π· Π½Π°ΡΠ΅ΠΊΠΎΠΌΡΡ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΠ΅Ρ ΡΠΎΠ±ΠΎΠΉ ΡΠ»ΠΎΠΆΠ½ΡΠΉ, ΠΌΠ½ΠΎΠ³ΠΎΡΡΠ°ΠΏΠ½ΡΠΉ ΠΏΡΠΎΡΠ΅ΡΡ, ΠΊΠΎΡΠΎΡΡΠΉ ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΠ΅ΡΡΡ ΡΠΊΠΎΠΎΡΠ΄ΠΈΠ½ΠΈΡΠΎΠ²Π°Π½Π½ΡΠΌ Π΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ΠΌ Π±ΠΎΠ»ΡΡΠΎΠ³ΠΎ ΡΠΈΡΠ»Π° Π³Π΅Π½ΠΎΠ², ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠΎΠΌ ΠΊΠΎΡΠΎΡΠΎΠ³ΠΎ ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ Π·ΡΠ΅Π»ΠΎΠ³ΠΎ ΡΠΉΡΠ°. ΠΠ·ΡΡΠ΅Π½ΠΈΠ΅ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° — ΠΎΠ΄Π½Π° ΠΈΠ· ΠΏΡΠΈΠΎΡΠΈΡΠ΅ΡΠ½ΡΡ Π·Π°Π΄Π°Ρ ΡΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎΠΉ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ, ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΠΎΠΌ ΡΠ΅Π³ΠΎ ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΎΠ³ΡΠΎΠΌΠ½ΠΎΠ΅ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²ΠΎ ΠΏΡΠ±Π»ΠΈΠΊΡΠ΅ΠΌΡΡ Π½Π° ΡΡΡ ΡΠ΅ΠΌΡ ΡΠ°Π±ΠΎΡ. Π Π΄Π°Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅ Π΄Π»Ρ ΠΈΠ·ΡΡΠ΅Π½ΠΈΡ ΠΏΡΠΎΡΠ΅ΡΡΠ° ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° Π±ΡΠ»Π° Π²ΡΠ±ΡΠ°Π½Π° D. melanogaster. ΠΡΠΎΠ·ΠΎΡΠΈΠ»Π° ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΊΠ»Π°ΡΡΠΈΡΠ΅ΡΠΊΠΈΠΌ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΠΌ ΠΎΠ±ΡΠ΅ΠΊΡΠΎΠΌ Π΄Π»Ρ ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ ΠΏΠΎΠ΄ΠΎΠ±Π½ΠΎΠ³ΠΎ ΡΠΎΠ΄Π° ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ, ΠΏΠΎΡΠΊΠΎΠ»ΡΠΊΡ ΠΏΡΠ΅Π΄ΠΎΡΡΠ°Π²Π»ΡΠ΅Ρ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡΡ ΠΏΡΠΎΠ²ΠΎΠ΄ΠΈΡΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΊΠ°ΠΊ Π² ΡΠΈΡΡΠ΅ΠΌΠ΅ in vitro, ΡΠ°ΠΊ ΠΈ Π² ΡΠΈΡΡΠ΅ΠΌΠ΅ in vivo. Π¨ΠΈΡΠΎΡΠ°ΠΉΡΠΈΠΉ Π½Π°Π±ΠΎΡ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ ΠΌΡΡΠ°ΡΠΈΠΉ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ, Π²Π»ΠΈΡΡΡΠΈΡ Π½Π° ΡΠ°Π·Π»ΠΈΡΠ½ΡΠ΅ ΡΡΠ°ΠΏΡ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π°, ΠΏΠΎΠ·Π²ΠΎΠ»ΡΠ΅Ρ Π·Π½Π°ΡΠΈΡΠ΅Π»ΡΠ½ΠΎ Π΄ΠΎΠΏΠΎΠ»Π½ΠΈΡΡ ΠΈ ΡΠ°ΡΡΠΈΡΠΈΡΡ Π΄Π°Π½Π½ΡΠ΅, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π² ΡΠΈΡΡΠ΅ΠΌΠ΅ in vitro. Π‘ Π΄ΡΡΠ³ΠΎΠΉ ΡΡΠΎΡΠΎΠ½Ρ, Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Π° ΠΏΡΠ΅Π΄ΠΎΡΡΠ°Π²Π»ΡΠ΅Ρ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡΡ ΠΎΠ΄Π½ΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎΠ³ΠΎ ΠΏΡΠΈΠΌΠ΅Π½Π΅Π½ΠΈΡ ΡΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΡΡ ΡΠΈΡΠΎΠ³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ΅ΡΠΎΠ΄ΠΎΠ² ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ.
ΠΡΠΎΡΠ΅ΠΊΠ°Π½ΠΈΠ΅ ΠΏΡΠΎΡΠ΅ΡΡΠ° ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° ΡΠΈΠ»ΡΠ½ΠΎ ΠΎΡΠ»ΠΈΡΠ°Π΅ΡΡΡ Ρ ΡΠ°Π·Π½ΡΡ Π²ΠΈΠ΄ΠΎΠ², ΠΎΠ΄Π½Π°ΠΊΠΎ, ΠΈΠΌΠ΅Π΅ΡΡΡ ΡΡΠ΄ ΠΎΠ±ΡΠΈΡ ΡΠ΅ΡΡ, Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΠ·ΡΡΡΠΈΡ Π΄Π°Π½Π½ΡΠΉ ΠΏΡΠΎΡΠ΅ΡΡ Π΄Π°ΠΆΠ΅ Ρ ΡΠ²ΠΎΠ»ΡΡΠΈΠΎΠ½Π½ΠΎ ΠΎΡΠ΄Π°Π»Π΅Π½Π½ΡΡ Π²ΠΈΠ΄ΠΎΠ². ΠΠΎΡΡΠΎΠΌΡ Π΄Π°Π½Π½ΡΠ΅, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π½Π° D. melanogaster, ΠΈΠΌΠ΅ΡΡ Π±ΠΎΠ»ΡΡΠΎΠ΅ ΡΡΠ½Π΄Π°ΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΠΎΠ΅ Π·Π½Π°ΡΠ΅Π½ΠΈΠ΅ ΠΈ ΡΠΏΠΎΡΠΎΠ±ΡΡΠ²ΡΡΡ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΡΠ΅Π»ΠΎΡΡΠ½ΠΎΠΉ ΠΊΠ°ΡΡΠΈΠ½Ρ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΠΈ ΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° Ρ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
ΠΠΊΡΡΠ°Π»ΡΠ½ΠΎΡΡΡ ΠΏΡΠΎΠ±Π»Π΅ΠΌΡ. ΠΠΎΠ³Π΅Π½Π΅Π· ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΠ΅Ρ ΡΠΎΠ±ΠΎΠΉ ΡΠ»ΠΎΠΆΠ½ΡΠΉ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΠ΅ΠΌΡΠΉ ΡΠΊΠΎΠΎΡΠ΄ΠΈΠ½ΠΈΡΠΎΠ²Π°Π½Π½ΡΠΌ Π΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ΠΌ Π±ΠΎΠ»ΡΡΠΎΠ³ΠΎ ΡΠΈΡΠ»Π° Π³Π΅Π½ΠΎΠ², ΠΌΠ½ΠΎΠ³ΠΎΡΡΠ°ΠΏΠ½ΡΠΉ ΠΏΡΠΎΡΠ΅ΡΡ, ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠΎΠΌ ΠΊΠΎΡΠΎΡΠΎΠ³ΠΎ ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ Π·ΡΠ΅Π»ΠΎΠΉ ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΊΠΈ. ΠΠ·ΡΡΠ΅Π½ΠΈΠ΅ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° — ΠΎΠ΄Π½Π° ΠΈΠ· ΠΏΡΠΈΠΎΡΠΈΡΠ΅ΡΠ½ΡΡ Π·Π°Π΄Π°Ρ ΡΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎΠΉ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ ΠΌΠ΅Π΄ΠΈΡΠΈΠ½Ρ. ΠΠ΅ΡΠΌΠΎΡΡΡ Π½Π° ΡΠΎ, ΡΡΠΎ ΡΠΆΠ΅ ΠΈΠ·Π²Π΅ΡΡΠ½Ρ ΠΌΠ½ΠΎΠ³ΠΈΠ΅ Π³Π΅Π½Ρ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΠ΅ ΡΡΠΎΡ ΠΏΡΠΎΡΠ΅ΡΡ, Π² Π½Π°ΡΡΠΎΡΡΠ΅Π΅ Π²ΡΠ΅ΠΌΡ Π²Π΅Π΄ΡΡΡΡ ΠΈΠ½ΡΠ΅Π½ΡΠΈΠ²Π½ΡΠ΅ ΠΏΠΎΠΈΡΠΊΠΈ Π½ΠΎΠ²ΡΡ Π³Π΅Π½ΠΎΠ² ΠΈ ΠΎΡΠΎΠ±Π΅Π½Π½ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠΎΠ² Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΡΡΠΈΡ Π³Π΅Π½ΠΎΠ², ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² Π΅Π³ΠΎ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ. ΠΠ°Π½Π½Π°Ρ ΡΠ°Π±ΠΎΡΠ° ΠΏΠΎΡΠ²ΡΡΠ΅Π½Π° ΠΈΠ·ΡΡΠ΅Π½ΠΈΡ ΡΠΎΠ»ΠΈ Π³Π΅Π½Π° Trithorax-like {Trl) Π² ΠΎΠΎΠ³Π΅Π½Π΅Π·Π΅ Drosophiia melanogaster. ΠΠ΅Π½ Trl ΠΊΠΎΠ΄ΠΈΡΡΠ΅Ρ ΠΌΠ½ΠΎΠ³ΠΎΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΡΠΉ Π±Π΅Π»ΠΎΠΊ GAGA, ΠΎΠ΄Π½ΠΎΠΉ ΠΈΠ· ΠΎΡΠ½ΠΎΠ²Π½ΡΡ ΡΡΠ½ΠΊΡΠΈΠΉ ΠΊΠΎΡΠΎΡΠΎΠ³ΠΎ ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½ΠΎΠ². ΠΠ°ΠΈΠ±ΠΎΠ»Π΅Π΅ Ρ ΠΎΡΠΎΡΠΎ ΠΈΠ·ΡΡΠ΅Π½Π° ΡΠΎΠ»Ρ ΡΡΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° Π² Ρ ΠΎΠ΄Π΅ ΡΠΌΠ±ΡΠΈΠΎΠ³Π΅Π½Π΅Π·Π°. Π ΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠΉ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΠΈ Π±Π΅Π»ΠΊΠ° GAGA Π² Ρ ΠΎΠ΄Π΅ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΡΠ°Π½Π΅Π΅ ΠΌΠΎΠΆΠ½ΠΎ Π±ΡΠ»ΠΎ ΡΡΠ΄ΠΈΡΡ ΡΠΎΠ»ΡΠΊΠΎ Π½Π° ΠΎΡΠ½ΠΎΠ²Π°Π½ΠΈΠΈ ΠΊΠΎΡΠ²Π΅Π½Π½ΡΡ Π΄Π°Π½Π½ΡΡ . ΠΡΠ»ΠΎ ΠΈΠ·Π²Π΅ΡΡΠ½ΠΎ, ΡΡΠΎ ΠΏΡΠΎΠ΄ΡΠΊΡΡ Π³Π΅Π½Π° Trl, Π°ΠΊΡΠΈΠ²Π½ΠΎ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡΡΡΡΠ΅Π³ΠΎΡΡ Π² ΡΠΈΡΠ½ΠΈΠΊΠ°Ρ Π²Π·ΡΠΎΡΠ»ΡΡ ΠΌΡΡ , Π½Π°ΠΊΠ°ΠΏΠ»ΠΈΠ²Π°ΡΡΡΡ Π² ΡΠ°ΡΡΡΡΠ΅ΠΌ ΠΎΠΎΡΠΈΡΠ΅ ΠΈ Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΡ Π΄Π»Ρ ΡΠ°Π·Π²ΠΈΠ²Π°ΡΡΠ΅Π³ΠΎΡΡ ΡΠΌΠ±ΡΠΈΠΎΠ½Π°. ΠΠ΄Π½Π°ΠΊΠΎ ΠΏΡΠΈ Π°Π½Π°Π»ΠΈΠ·Π΅ ΡΠ°Π½Π΅Π΅ ΠΈΠ·Π²Π΅ΡΡΠ½ΡΡ ΠΠ³ΠΠΌΡΡΠ°Π½ΡΠΎΠ² ΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠΎΠ·Π°ΠΈΠΊΠΎΠ² Π½Π΅ Π±ΡΠ»ΠΎ Π²ΡΡΠ²Π»Π΅Π½ΠΎ Π½ΠΈΠΊΠ°ΠΊΠΈΡ ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ Π½Π°ΡΡΡΠ΅Π½ΠΈΠΉ Π² ΡΡΡΡΠΊΡΡΡΠ΅ ΡΠΈΡΠ½ΠΈΠΊΠΎΠ² ΠΈ ΡΠΉΡΠ΅Π²ΡΡ ΠΊΠ°ΠΌΠ΅Ρ. ΠΡΡΡΡΡΡΠ²ΠΈΠ΅ ΠΏΡΡΠΌΡΡ Π΄ΠΎΠΊΠ°Π·Π°ΡΠ΅Π»ΡΡΡΠ² ΡΠΎΠ³ΠΎ, ΡΡΠΎ Π³Π΅Π½ Trl Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌ Π΄Π»Ρ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ, ΡΠ²ΡΠ·Π°Π½ΠΎ Ρ ΡΠ΅ΠΌ, ΡΡΠΎ Π² ΠΌΠΈΡΠΎΠ²ΡΡ ΠΊΠΎΠ»Π»Π΅ΠΊΡΠΈΡΡ ΠΎΡΡΡΡΡΡΠ²ΠΎΠ²Π°Π»ΠΈ ΠΌΡΡΠ°ΡΠΈΠΈ, ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡΠΈΠ΅ ΠΏΡΠΎΠ²ΠΎΠ΄ΠΈΡΡ ΠΏΠΎΠ΄ΠΎΠ±Π½ΠΎΠ³ΠΎ ΡΠΎΠ΄Π° ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ. Π Π΄Π°Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅ Π±ΡΠ»ΠΈ ΠΏΠΎΠ»ΡΡΠ΅Π½Ρ ΠΈ ΠΎΡ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΠ·ΠΎΠ²Π°Π½Ρ Π½ΠΎΠ²ΡΠ΅ Π³ΠΈΠΏΠΎΠΌΠΎΡΡΠ½ΡΠ΅ ΠΌΡΡΠ°ΡΠΈΠΈ Π³Π΅Π½Π° Trl, ΠΊΠΎΡΠΎΡΡΠ΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΠΈΠ»ΠΈ ΠΏΡΠΎΠ²Π΅ΡΡΠΈ Π²ΡΠ΅ΡΡΠΎΡΠΎΠ½Π½ΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· Π²Π»ΠΈΡΠ½ΠΈΡ Π³Π΅Π½Π° Trl Π½Π° ΠΎΠΎΠ³Π΅Π½Π΅Π· Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ. ΠΠ΅Π»ΠΎΠΊ GAGA ΡΡΠ΅Π±ΡΠ΅ΡΡΡ Π΄Π»Ρ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΡ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ ΠΌΠ½ΠΎΠΆΠ΅ΡΡΠ²Π° Π³Π΅Π½ΠΎΠ² Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ, ΠΏΠΎΡΡΠΎΠΌΡ ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π² ΡΠ°Π±ΠΎΡΠ΅ Π΄Π°Π½Π½ΡΠ΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ Π²ΡΡΠ²ΠΈΡΡ Π½Π΅ ΡΠΎΠ»ΡΠΊΠΎ ΡΡΠ°ΡΡΠΈΠ΅ ΡΠ°ΠΌΠΎΠ³ΠΎ Π³Π΅Π½Π° Trl Π² Ρ ΠΎΠ΄Π΅ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π°, Π½ΠΎ ΠΈ ΡΡΠ΄ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»Π°Π³Π°Π΅ΠΌΡΡ , ΡΠ°Π½Π΅Π΅ Π½Π΅ ΠΎΠΏΠΈΡΠ°Π½Π½ΡΡ Π² Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΠ΅, Π³Π΅Π½ΠΎΠ²-ΠΌΠΈΡΠ΅Π½Π΅ΠΉ Π±Π΅Π»ΠΊΠ° GAGA, ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² ΠΊΠΎΠ½ΡΡΠΎΠ»Π΅ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π°. Π Π΅Π·ΡΠ»ΡΡΠ°ΡΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΠΈΠΌΠ΅ΡΡ ΡΡΠ½Π΄Π°ΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΠΎΠ΅ Π·Π½Π°ΡΠ΅Π½ΠΈΠ΅ Π΄Π»Ρ Π²ΡΡΡΠ½Π΅Π½ΠΈΡ ΠΏΡΠΈΡΠΈΠ½ ΡΡΠ΅ΡΠΈΠ»ΡΠ½ΠΎΡΡΠΈ Π½Π°ΡΠ΅ΠΊΠΎΠΌΡΡ , Π° Π΄Π°Π½Π½ΡΠ΅ ΠΎ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΡΠΈΠ»Π°ΠΌΠ΅Π½ΡΠΎΠ² ΠΈ ΠΌΠΈΠ³ΡΠ°ΡΠΈΠΈ ΠΊΠ»Π΅ΡΠΎΠΊ ΠΌΠΎΠ³ΡΡ Π±ΡΡΡ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½Ρ Π² ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡΡ , ΠΏΡΠΎΠ²ΠΎΠ΄ΠΈΠΌΡΡ Π½Π° Π΄ΡΡΠ³ΠΈΡ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΡ ΠΎΠ±ΡΠ΅ΠΊΡΠ°Ρ .
Π¦Π΅Π»ΠΈ ΠΈ Π·Π°Π΄Π°ΡΠΈ ΡΠ°Π±ΠΎΡΡ ΡΠΎΡΡΠΎΡΠ»ΠΈ Π² ΠΈΠ·ΡΡΠ΅Π½ΠΈΠΈ ΡΠΎΠ»ΠΈ Π³Π΅Π½Π° Trl ΠΈ ΠΊΠΎΠ΄ΠΈΡΡΠ΅ΠΌΠΎΠ³ΠΎ ΠΈΠΌ Π±Π΅Π»ΠΊΠ° GAGA Π² ΠΎΠΎΠ³Π΅Π½Π΅Π·Π΅ D. melanogaster. Π ΡΠ²ΡΠ·ΠΈ Ρ ΡΡΠΈΠΌ Π±ΡΠ»ΠΈ ΠΏΠΎΡΡΠ°Π²Π»Π΅Π½Ρ ΡΠ»Π΅Π΄ΡΡΡΠΈΠ΅ Π·Π°Π΄Π°ΡΠΈ:
1. ΠΠΎΠ»ΡΡΠΈΡΡ Π½ΠΎΠ²ΡΠ΅ Π³ΠΈΠΏΠΎΠΌΠΎΡΡΠ½ΡΠ΅ ΠΌΡΡΠ°ΡΠΈΠΈ Π³Π΅Π½Π° Trl, Π²Π»ΠΈΡΡΡΠΈΠ΅ Π½Π° ΠΎΠΎΠ³Π΅Π½Π΅Π· Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ, ΠΈ ΠΏΡΠΎΠ²Π΅ΡΡΠΈ ΠΈΡ ΠΊΠ°ΡΡΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅;
2. ΠΡΠΎΠ²Π΅ΡΡΠΈ Π°Π½Π°Π»ΠΈΠ· Π²Π»ΠΈΡΠ½ΠΈΡ Π²Π½ΠΎΠ²Ρ ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΡ ΠΌΡΡΠ°ΡΠΈΠΉ Π½Π° ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Π³Π΅Π½Π° Trl ΠΊΠ°ΠΊ Π½Π° ΡΡΠΎΠ²Π½Π΅ ΠΌΠ ΠΠ, ΡΠ°ΠΊ ΠΈ Π½Π° ΡΡΠΎΠ²Π½Π΅ Π±Π΅Π»ΠΊΠ°;
3. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΡ Π²Π»ΠΈΡΠ½ΠΈΠ΅ ΡΡΠΈΡ ΠΌΡΡΠ°ΡΠΈΠΉ Π½Π° ΠΎΠΎΠ³Π΅Π½Π΅Π· Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ;
4. ΠΠ·ΡΡΠΈΡΡ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΡΠ΅ΡΡΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΈ ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΠΈ ΡΠΈΡΠ½ΠΈΠΊΠΎΠ² ΠΌΡΡΠ°Π½ΡΠΎΠ² Π² Π·Π°Π²ΠΈΡΠΈΠΌΠΎΡΡΠΈ ΠΎΡ ΡΡΠΎΠ²Π½Ρ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½Π° Trl.
ΠΠ°ΡΡΠ½Π°Ρ Π½ΠΎΠ²ΠΈΠ·Π½Π° ΡΠ°Π±ΠΎΡΡ. ΠΠΎΠ»ΡΡΠ΅Π½Ρ ΠΈ ΠΎΡ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΠ·ΠΎΠ²Π°Π½Ρ Π½ΠΎΠ²ΡΠ΅ Π³ΠΈΠΏΠΎΠΌΠΎΡΡΠ½ΡΠ΅ ΠΌΡΡΠ°ΡΠΈΠΈ Π³Π΅Π½Π° Trl, Π² ΡΠ°Π·Π½ΠΎΠΉ ΡΡΠ΅ΠΏΠ΅Π½ΠΈ ΡΠ½ΠΈΠΆΠ°ΡΡΠΈΠ΅ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Π³Π΅Π½Π°. ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ ΠΌΡΡΠ°ΡΠΈΠΈ ΡΠ²ΠΈΠ»ΠΈΡΡ ΡΠ½ΠΈΠΊΠ°Π»ΡΠ½ΡΠΌ ΠΈΠ½ΡΡΡΡΠΌΠ΅Π½ΡΠΎΠΌ, ΠΏΠΎΠ·Π²ΠΎΠ»ΠΈΠ²ΡΠΈΠΌ Π²ΠΏΠ΅ΡΠ²ΡΠ΅ ΠΏΡΠΎΠ²Π΅ΡΡΠΈ Π΄Π΅ΡΠ°Π»ΡΠ½ΠΎΠ΅ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π²Π»ΠΈΡΠ½ΠΈΡ Π³Π΅Π½Π° Trl Π½Π° ΠΎΠΎΠ³Π΅Π½Π΅Π· Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ.
ΠΠΏΠ΅ΡΠ²ΡΠ΅ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½Π° Trl ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΡ ΡΡΡΡΠΊΡΡΡΡ ΡΠΈΡΠ½ΠΈΠΊΠΎΠ² ΠΌΡΡ . ΠΠ°ΡΡΡΠ΅Π½ΠΈΡ Π²ΡΡΠ²Π»ΡΡΡΡΡ ΠΊΠ°ΠΊ Π² ΠΎΠ²Π°ΡΠΈΠΎΠ»Π°Ρ , ΡΠ°ΠΊ ΠΈ Π² ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΡ ΡΠΉΡΠ΅Π²ΡΡ ΠΊΠ°ΠΌΠ΅ΡΠ°Ρ . Π£ΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ 7Π³/-ΠΌΡΡΠ°ΡΠΈΠΈ Π²ΡΠ·ΡΠ²Π°ΡΡ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΡΠΈΡΠ»Π° ΠΏΠΎΠ»ΠΎΠ²ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ (ΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΈ ΠΎΠΎΡΠΈΡΠΎΠ²) Π² ΡΠΉΡΠ΅Π²ΡΡ ΠΊΠ°ΠΌΠ΅ΡΠ°Ρ ΠΏΠΎ ΡΡΠ°Π²Π½Π΅Π½ΠΈΡ Ρ Π½ΠΎΡΠΌΠΎΠΉ. ΠΠΏΠ΅ΡΠ²ΡΠ΅ Ρ 7Π³/-ΠΌΡΡΠ°Π½ΡΠΎΠ² Π²ΡΡΠ²Π»Π΅Π½Ρ Π΄Π΅ΡΠ΅ΠΊΡΡ Π² ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΡΠ°Π΄ΠΈΠΎΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΡΠΈΠ»Π°ΠΌΠ΅Π½ΡΠΎΠ² Π² ΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΊΠ»Π΅ΡΠΊΠ°Ρ Π½Π° ΠΏΠΎΠ·Π΄Π½ΠΈΡ ΡΡΠ°Π΄ΠΈΡΡ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΡΠΉΡΠ΅Π²ΠΎΠΉ ΠΊΠ°ΠΌΠ΅ΡΡ. Π ΡΠ°Π±ΠΎΡΠ΅ Π²ΠΏΠ΅ΡΠ²ΡΠ΅ ΠΎΠΏΠΈΡΠ°Π½ΠΎ Π½Π°ΡΡΡΠ΅Π½ΠΈΠ΅ Π² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠΎΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ Ρ 777 ΠΌΡΡΠ°Π½ΡΠΎΠ². ΠΠ° ΠΏΠΎΠ·Π΄Π½ΠΈΡ ΡΡΠ°Π΄ΠΈΡΡ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΡΠΉΡΠ΅Π²ΠΎΠΉ ΠΊΠ°ΠΌΠ΅ΡΡ Ρ ΠΌΡΡΠ°Π½ΡΠΎΠ² Π½Π°ΡΡΡΠ°Π΅ΡΡΡ ΠΌΠΈΠ³ΡΠ°ΡΠΈΡ Π±ΠΎΡΠ΄ΡΡΠ½ΡΡ ΠΈ ΡΠ΅Π½ΡΡΠΈΠΏΠ΅ΡΠ°Π»ΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ. ΠΠ΅ΡΠ΅ΠΊΡΡ Π² ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΡΠΈΠ»Π°ΠΌΠ΅Π½ΡΠΎΠ² ΠΈ Π² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠΎΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ ΠΏΡΠΈΠ²ΠΎΠ΄ΡΡ ΠΊ Π½Π°ΡΡΡΠ΅Π½ΠΈΡ ΠΏΡΠΎΡΠ΅ΡΡΠ° ΠΏΠ΅ΡΠ΅Π½ΠΎΡΠ° ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΡ ΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ Π² ΠΎΠΎΡΠΈΡ ΠΈ, ΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎ, ΠΊ ΠΏΡΠ΅ΠΊΡΠ°ΡΠ΅Π½ΠΈΡ ΡΠΎΡΡΠ° ΠΎΠΎΡΠΈΡΠ°.
ΠΠ°ΡΡΠ½ΠΎ-ΠΏΡΠ°ΠΊΡΠΈΡΠ΅ΡΠΊΠ°Ρ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΡ ΡΠ°Π±ΠΎΡΡ. Π Π΅Π·ΡΠ»ΡΡΠ°ΡΡ Π΄Π°Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΡ ΡΠΏΠΎΡΠΎΠ±ΡΡΠ²ΡΡΡ ΡΠ°ΡΡΠΈΡΠ΅Π½ΠΈΡ ΡΡΠ½Π΄Π°ΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΡΡ Π·Π½Π°Π½ΠΈΠΉ ΠΎ ΠΏΡΠΎΡΠ΅ΡΡΠ΅ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° Π½Π°ΡΠ΅ΠΊΠΎΠΌΡΡ . ΠΠ°ΡΠ΅ΡΠΈΠ°Π»Ρ Π΄Π°Π½Π½ΠΎΠΉ Π΄ΠΈΡΡΠ΅ΡΡΠ°ΡΠΈΠΎΠ½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΡ ΠΌΠΎΠ³ΡΡ Π±ΡΡΡ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½Ρ Π² ΠΊΡΡΡΠ°Ρ Π»Π΅ΠΊΡΠΈΠΉ Π΄Π»Ρ ΡΡΡΠ΄Π΅Π½ΡΠΎΠ² Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈ ΠΌΠ΅Π΄ΠΈΡΠΈΠ½ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠ»ΡΡΠ΅ΡΠΎΠ².
ΠΠΏΡΠΎΠ±Π°ΡΠΈΡ ΡΠ°Π±ΠΎΡΡ. ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π² Ρ ΠΎΠ΄Π΅ ΡΠ°Π±ΠΎΡΡ Π΄Π°Π½Π½ΡΠ΅ Π±ΡΠ»ΠΈ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»Π΅Π½Ρ Π°Π²ΡΠΎΡΠΎΠΌ Π² Π²ΠΈΠ΄Π΅ ΡΠ΅Π·ΠΈΡΠΎΠ² Π½Π° ΡΠ»Π΅Π΄ΡΡΡΠΈΡ ΠΊΠΎΠ½ΡΠ΅ΡΠ΅Π½ΡΠΈΡΡ :
β’ Π£ΡΡΠ½ΡΠΉ Π΄ΠΎΠΊΠ»Π°Π΄ Π½Π° ΠΎΡΡΠ΅ΡΠ½ΠΎΠΉ ΡΠ΅ΡΡΠΈΠΈ ΠΠ¦ΠΈΠ Π‘Π Π ΠΠ 2003 Π³.
β’ XIV Π¨ΠΊΠΎΠ»Π΅ ΠΏΠΎ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ «ΠΠΊΡΡΠ°Π»ΡΠ½ΡΠ΅ ΠΏΡΠΎΠ±Π»Π΅ΠΌΡ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΠΈ Π±ΠΈΠΎΡΠ΅Ρ Π½ΠΎΠ»ΠΎΠ³ΠΈΠΈ», ΠΠ²Π΅Π½ΠΈΠ³ΠΎΡΠΎΠ΄, 2004.
β’ Π¨ΠΊΠΎΠ»Π΅-ΡΠ΅ΠΌΠΈΠ½Π°ΡΠ΅ «ΠΠ°Π·Π΅ΡΠ½Π°Ρ ΠΊΠΎΠ½ΡΠΎΠΊΠ°Π»ΡΠ½Π°Ρ ΠΌΠΈΠΊΡΠΎΡΠΊΠΎΠΏΠΈΡ Π² Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ ΠΌΠ΅Π΄ΠΈΡΠΈΠ½Π΅», ΠΠ²Π΅Π½ΠΈΠ³ΠΎΡΠΎΠ΄, 2005.
β’ 10-ΠΎΠΉ ΠΡΡΠΈΠ½ΡΠΊΠΎΠΉ Π¨ΠΊΠΎΠ»Π΅-ΠΊΠΎΠ½ΡΠ΅ΡΠ΅Π½ΡΠΈΠΈ ΠΌΠΎΠ»ΠΎΠ΄ΡΡ ΡΡΠ΅Π½ΡΡ , ΠΡΡΠΈΠ½ΠΎ, 2006.
β’ ΠΠ΅ΠΆΠ΄ΡΠ½Π°ΡΠΎΠ΄Π½ΠΎΠΉ ΠΊΠΎΠ½ΡΠ΅ΡΠ΅Π½ΡΠΈΠΈ «ΠΠ΅Π½Π΅ΡΠΈΠΊΠ° Π² Π ΠΎΡΡΠΈΠΈ ΠΈ ΠΌΠΈΡΠ΅», 2006.
ΠΠΊΠ»Π°Π΄ Π°Π²ΡΠΎΡΠ°. ΠΡΠ½ΠΎΠ²Π½Π°Ρ ΡΠ°ΡΡΡ ΡΠΊΡΠΏΠ΅ΡΠΈΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΠΎΠΉ ΡΠ°Π±ΠΎΡΡ ΠΈ ΠΎΠ±ΡΠ°Π±ΠΎΡΠΊΠ° ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΡ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠΎΠ² Π²ΡΠΏΠΎΠ»Π½Π΅Π½Π° Π°Π²ΡΠΎΡΠΎΠΌ Π»ΠΈΡΠ½ΠΎ, Π»ΠΈΠ±ΠΎ ΡΠΎΠ²ΠΌΠ΅ΡΡΠ½ΠΎ Ρ Π½Π°ΡΡΠ½ΡΠΌ ΡΡΠΊΠΎΠ²ΠΎΠ΄ΠΈΡΠ΅Π»Π΅ΠΌ. ΠΠ°ΡΠ΅ΡΠΈΠ°Π»Ρ, Π²ΠΎΡΠ΅Π΄ΡΠΈΠ΅ Π² Π΄Π°Π½Π½ΡΡ ΡΠ°Π±ΠΎΡΡ, ΠΎΠ±ΡΡΠΆΠ΄Π°Π»ΠΈΡΡ ΠΈ ΠΏΡΠ±Π»ΠΈΠΊΠΎΠ²Π°Π»ΠΈΡΡ ΡΠΎΠ²ΠΌΠ΅ΡΡΠ½ΠΎ Ρ ΡΠΎΠ°Π²ΡΠΎΡΠ°ΠΌΠΈ ΠΈ Π½Π°ΡΡΠ½ΡΠΌ ΡΡΠΊΠΎΠ²ΠΎΠ΄ΠΈΡΠ΅Π»Π΅ΠΌ. ΠΠ½Π°Π»ΠΈΠ· ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½Π° Trl Ρ ΠΌΡΡΠ°Π½ΡΠΎΠ², Π° ΡΠ°ΠΊΠΆΠ΅ Π°Π½Π°Π»ΠΈΠ· ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½ΠΎΠ² Actin 5, chic, cpb ΠΈ bif Π²ΡΠΏΠΎΠ»Π½Π΅Π½ ΠΏΡΠΈ ΡΡΠ°ΡΡΠΈΠΈ ΠΠ°ΡΠ°Π³ΠΎΠ΄ΠΈΠ½Π° Π. Π. (ΠΠ¦ΠΈΠ Π‘Π Π ΠΠ, ΠΠ°Π±ΠΎΡΠ°ΡΠΎΡΠΈΡ ΡΠ²ΠΎΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΊΠ»Π΅ΡΠΊΠΈ). ΠΠ»Π΅ΠΊΡΡΠΎΠ½Π½ΠΎ-ΠΌΠΈΠΊΡΠΎΡΠΊΠΎΠΏΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΡΠΈΠ»Π°ΠΌΠ΅Π½ΡΠΎΠ² Π² ΡΠΈΡΠ½ΠΈΠΊΠ°Ρ ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½ ΠΏΡΠΈ ΠΏΠΎΠΌΠΎΡΠΈ ΠΠΈΡΠ΅Π»Π΅Π²ΠΎΠΉ Π. Π ΠΈ ΠΡΠ±Π°Π½ΠΎΠ²ΠΎΠΉ Π. Π. (ΠΠ¦ΠΈΠ Π‘Π Π ΠΠ, ΠΠ°Π±ΠΎΡΠ°ΡΠΎΡΠΈΡ ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ ΡΡΠ½ΠΊΡΠΈΠΈ ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΡ ΡΡΡΡΠΊΡΡΡ). ΠΠ½Π°Π»ΠΈΠ· Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ½ΠΎΠΉ ΡΠ΅Π³ΡΠ΅Π³Π°ΡΠΈΠΈ Π² ΡΠ΅ΡΠ΅Π½ΠΈΠ΅ ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Π»ΡΠ½ΡΡ ΠΌΠΈΡΠΎΠ·ΠΎΠ² ΠΏΡΠΎΠ²ΠΎΠ΄ΠΈΠ»ΡΡ Ρ ΡΡΠ°ΡΡΠΈΠ΅ΠΌ Π€Π΅Π΄ΠΎΡΠΎΠ²ΠΎΠΉ Π‘. Π. (ΠΠ¦ΠΈΠ Π‘Π Π ΠΠ, ΠΠ°Π±ΠΎΡΠ°ΡΠΎΡΠΈΡ Π³Π΅Π½Π΅ΡΠΈΠΊΠΈ ΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΡΠΈΠΊΠ»Π°). ΠΠΈΠΊΡΠΎΡΠΊΠΎΠΏΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· Π½Π° ΠΊΠΎΠ½ΡΠΎΠΊΠ°Π»ΡΠ½ΠΎΠΌ ΠΌΠΈΠΊΡΠΎΡΠΊΠΎΠΏΠ΅ ΠΏΡΠΎΠ²ΠΎΠ΄ΠΈΠ»ΡΡ ΠΏΡΠΈ ΡΠ΅Ρ Π½ΠΈΡΠ΅ΡΠΊΠΎΠΉ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠΊΠ΅ Π ΡΠ±ΡΠΎΠ²Π° Π. Π. ΠΈ ΠΠ°ΠΉΠ±ΠΎΡΠΎΠ΄ΠΈΠ½Π° Π‘. Π. (ΠΠ¦ΠΈΠ Π‘Π Π ΠΠ, ΠΠ°Π±ΠΎΡΠ°ΡΠΎΡΠΈΡ ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ ΡΡΠ½ΠΊΡΠΈΠΈ ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΡ ΡΡΡΡΠΊΡΡΡ). ΠΡΡΠ°ΡΠΈΡ Tifn82 Π±ΡΠ»Π° Π²ΡΠ΄Π΅Π»Π΅Π½Π° ΠΠΎΠ»ΠΎΡΠΈΠ½ΠΎΠΉ Π.Π.
ΠΠ«ΠΠΠΠ«.
1. ΠΠΎΠ»ΡΡΠ΅Π½Ρ Π½ΠΎΠ²ΡΠ΅ Π³ΠΈΠΏΠΎΠΌΠΎΡΡΠ½ΡΠ΅ ΠΌΡΡΠ°ΡΠΈΠΈ Π³Π΅Π½Π° Trl, Π·Π°ΡΡΠ°Π³ΠΈΠ²Π°ΡΡΠΈΠ΅ 5'-ΠΎΠ±Π»Π°ΡΡΡ, ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½ΠΎ ΠΈΡ ΠΊΠ°ΡΡΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅. ΠΡΡΠ°ΡΠΈΡ Trl362 ΠΎΠ±ΡΡΠ»ΠΎΠ²Π»Π΅Π½Π° Π²ΡΡΡΠΎΠΉΠΊΠΎΠΉ Π -ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ° Π² Π±'-Π½Π΅ΠΊΠΎΠ΄ΠΈΡΡΡΡΡΡ ΠΎΠ±Π»Π°ΡΡΡ Π³Π΅Π½Π° (Π² ΠΏΠΎΠ·ΠΈΡΠΈΡ 2979) ΠΈ ΠΏΡΠΈΠ»Π΅Π³Π°ΡΡΡΡ ΠΊ Π½Π΅ΠΌΡ Π΄Π΅Π»Π΅ΡΠΈΡ ΡΠ°Π·ΠΌΠ΅ΡΠΎΠΌ 97 ΠΏ.Π½. (2979−3076). ΠΠ΅Π»Π΅ΡΠΈΡ ΡΠ΄Π°Π»ΡΠ΅Ρ Π΄Π²Π° ΠΏΠ΅ΡΠ²ΡΡ ΡΠ°ΠΉΡΠ° ΠΈΠ½ΠΈΡΠΈΠ°ΡΠΈΠΈ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΈ, ΠΈΡΠΏΠΎΠ»ΡΠ·ΡΠ΅ΠΌΡΡ Π² ΡΠΈΡΠ½ΠΈΠΊΠ°Ρ . ΠΡΡΠ°ΡΠΈΡ Trf62(ex) ΡΠΎΠ΄Π΅ΡΠΆΠΈΡ ΡΠ°ΠΊΡΡ ΠΆΠ΅ Π΄Π΅Π»Π΅ΡΠΈΡ Π² 5'-Π½Π΅ΠΊΠΎΠ΄ΠΈΡΡΡΡΠ΅ΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ Π³Π΅Π½Π°, ΠΊΠ°ΠΊ ΠΈ ΠΌΡΡΠ°ΡΠΈΡ Π/7362, Π½ΠΎ Π½Π΅ ΡΠΎΠ΄Π΅ΡΠΆΠΈΡ Π -ΡΠ»Π΅ΠΌΠ΅Π½Ρ.
2. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΌΡΡΠ°ΡΠΈΠΈ Trf62, Π’Π³Π 62(Π΅Ρ > ΠΈ Π’/Π’®" 82 ΠΏΡΠΈΠ²ΠΎΠ΄ΡΡ ΠΊ ΡΠ½ΠΈΠΆΠ΅Π½ΠΈΡ ΠΆΠΈΠ·Π½Π΅ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΠΈ ΠΈ ΡΡΠ΅ΡΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΡΠ°ΠΌΠΎΠΊ Π² Π³ΠΎΠΌΠΎΠ·ΠΈΠ³ΠΎΡΠ½ΠΎΠΌ ΡΠΎΡΡΠΎΡΠ½ΠΈΠΈ ΠΈ Π² ΠΊΠΎΠΌΠΏΠ°ΡΠ½Π΄Π΅ Ρ Π½ΡΠ»Ρ-Π°Π»Π»Π΅Π»Π΅ΠΌ.
3. Π£ΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ Π² ΡΠΈΡΠ½ΠΈΠΊΠ°Ρ ΡΠ°ΠΌΠΎΠΊ Trl362 ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΡ Π³Π΅Π½Π° Trl ΡΠ½ΠΈΠΆΠ΅Π½Π° ΠΊΠ°ΠΊ ΠΌΠΈΠ½ΠΈΠΌΡΠΌ Π² 10 ΡΠ°Π·, ΡΡΠΎ ΡΠΎΠΏΡΠΎΠ²ΠΎΠΆΠ΄Π°Π΅ΡΡΡ Π·Π½Π°ΡΠΈΡΠ΅Π»ΡΠ½ΡΠΌ ΡΠΌΠ΅Π½ΡΡΠ΅Π½ΠΈΠ΅ΠΌ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²Π° Π±Π΅Π»ΠΊΠ° GAGA Π² ΠΏΠΎΠ»ΠΎΠ²ΡΡ ΠΊΠ»Π΅ΡΠΊΠ°Ρ . Π’ΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΡ Π³Π΅Π½Π° Trl Π² ΡΠΈΡΠ½ΠΈΠΊΠ°Ρ Π³ΠΎΠΌΠΎΠ·ΠΈΠ³ΠΎΡΠ½ΡΡ Trl8″ 82-ΠΌΡΡΠ°Π½ΡΠ½ΡΡ ΡΠ°ΠΌΠΎΠΊ ΡΠ½ΠΈΠΆΠ΅Π½Π° ΠΏΡΠΈΠ±Π»ΠΈΠ·ΠΈΡΠ΅Π»ΡΠ½ΠΎ Π½Π° ΡΡΠ΅ΡΡ, ΡΠΎΠ³Π΄Π° ΠΊΠ°ΠΊ Π² ΡΠΈΡΠ½ΠΈΠΊΠ°Ρ Π’Π³1362(Π΅Ρ )-1Π»ΡΡΠ°Π½ΡΠΎΠ² ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Π³Π΅Π½Π° ΠΏΡΠ°ΠΊΡΠΈΡΠ΅ΡΠΊΠΈ Π½Π΅ ΠΎΡΠ»ΠΈΡΠ°Π΅ΡΡΡ ΠΎΡ Π½ΠΎΡΠΌΡ.
4. ΠΡΡΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ Ρ Π²ΡΠ΅Ρ Π°Π½Π°Π»ΠΈΠ·ΠΈΡΡΠ΅ΠΌΡΡ Trl ΠΌΡΡΠ°Π½ΡΠΎΠ² Π² ΡΠΎΠΉ ΠΈΠ»ΠΈ ΠΈΠ½ΠΎΠΉ ΡΡΠ΅ΠΏΠ΅Π½ΠΈ Π½Π°ΡΡΡΠ΅Π½Π° ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΡ ΠΎΠ²Π°ΡΠΈΠΎΠ» ΠΈ ΡΠΉΡΠ΅Π²ΡΡ ΠΊΠ°ΠΌΠ΅Ρ. Π―ΠΉΡΠ΅Π²ΡΠ΅ ΠΊΠ°ΠΌΠ΅ΡΡ ΠΌΡΡΠ°Π½ΡΠΎΠ² Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΠ·ΡΡΡΡΡ ΠΎΡΠ»ΠΈΡΠ½ΡΠΌ ΠΎΡ Π½ΠΎΡΠΌΡ ΡΠΈΡΠ»ΠΎΠΌ ΠΏΠΎΠ»ΠΎΠ²ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ. ΠΠ° ΠΏΠΎΠ·Π΄Π½ΠΈΡ ΡΡΠ°Π΄ΠΈΡΡ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΡΠΉΡΠ΅Π²ΠΎΠΉ ΠΊΠ°ΠΌΠ΅ΡΡ Ρ ΠΌΡΡΠ°Π½ΡΠΎΠ² Π²ΡΡΠ²Π»Π΅Π½Ρ Π½Π°ΡΡΡΠ΅Π½ΠΈΡ Π² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠΎΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ, Π° ΡΠ°ΠΊΠΆΠ΅ Π² ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΡΠ°Π΄ΠΈΠΎΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΡΠΈΠ»Π°ΠΌΠ΅Π½ΡΠΎΠ² Π² ΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΊΠ»Π΅ΡΠΊΠ°Ρ .
5.
ΠΠ²Π΅Π΄Π΅Π½ΠΈΠ΅
ΡΡΠ°Π½ΡΠ³Π΅Π½Π½ΡΡ ΠΊΠΎΠ½ΡΡΡΡΠΊΡΠΈΠΉ Π² Π³Π΅Π½ΠΎΠΌ ΠΌΡΡΠ°Π½ΡΠΎΠ², ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΠΈΡ Π΄ΠΎΠΏΠΎΠ»Π½ΠΈΡΠ΅Π»ΡΠ½ΡΡ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ ΠΊΠΠΠ Π³Π΅Π½Π° Trl, ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ Π²ΠΎΡΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΈΡ ΡΠ΅ΡΡΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΠΌΡΡΠ°Π½ΡΠ½ΡΡ ΡΠ°ΠΌΠΎΠΊ ΠΈ ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈΡ ΡΠΈΡΠ½ΠΈΠΊΠΎΠ². ΠΡΠΎ ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΠ΅Ρ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ ΠΈΠΌΠ΅Π½Π½ΠΎ ΠΈΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ Π² ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½Π° Trl ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΏΡΠΈΡΠΈΠ½ΠΎΠΉ Π²ΡΡΠ²Π»Π΅Π½Π½ΡΡ Π½Π°ΡΡΡΠ΅Π½ΠΈΠΉ Π² ΠΎΠΎΠ³Π΅Π½Π΅Π·Π΅ ΠΌΡΡΠ°Π½ΡΠΎΠ².
ΠΠ°ΠΊΠ»ΡΡΠ΅Π½ΠΈΠ΅
.
ΠΠ° ΠΎΡΠ½ΠΎΠ²Π΅ Π°Π½Π°Π»ΠΈΠ·Π° Π½ΠΎΠ²ΡΡ Π³ΠΈΠΏΠΎΠΌΠΎΡΡΠ½ΡΡ ΠΌΡΡΠ°ΡΠΈΠΉ Π³Π΅Π½Π° Trl Π±ΡΠ»ΠΎ ΡΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ Π²ΡΠ΅ ΡΡΠΈ ΠΌΡΡΠ°ΡΠΈΠΈ, Π² ΡΠ°Π·Π½ΠΎΠΉ ΡΡΠ΅ΠΏΠ΅Π½ΠΈ ΡΠ½ΠΈΠΆΠ°ΡΡΠΈΠ΅ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Π³Π΅Π½Π° Trl, ΠΏΡΠΈΠ²ΠΎΠ΄ΡΡ Π½Π΅ ΡΠΎΠ»ΡΠΊΠΎ ΠΊ Π½Π°ΡΡΡΠ΅Π½ΠΈΡΠΌ Π² ΡΠ°Π·Π²ΠΈΡΠΈΠΈ ΡΠΌΠ±ΡΠΈΠΎΠ½Π°, ΠΊΠ°ΠΊ ΡΡΠΎ Π±ΡΠ»ΠΎ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ ΡΠ°Π½Π΅Π΅, Π½ΠΎ ΡΠ°ΠΊΠΆΠ΅ Π΄ΡΠ°ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈ ΡΠΊΠ°Π·ΡΠ²Π°ΡΡΡΡ ΠΈ Π½Π° ΠΏΡΠΎΡΠ΅ΠΊΠ°Π½ΠΈΠΈ ΡΠ°ΠΌΠΎΠ³ΠΎ ΠΏΡΠΎΡΠ΅ΡΡΠ° ΠΎΠΎΠ³Π΅Π½Π΅Π·Π°. ΠΡΠΈ ΡΡΠΎΠΌ Ρ Trl ΠΌΡΡΠ°Π½ΡΠΎΠ² Π½Π°ΡΡΡΠ°Π΅ΡΡΡ ΡΠ°Π·Π²ΠΈΡΠΈΠ΅ ΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅, ΠΊΠ°ΠΊ ΠΏΠΎΠ»ΠΎΠ²ΡΡ , ΡΠ°ΠΊ ΠΈ ΡΠΎΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ ΡΠΈΡΠ½ΠΈΠΊΠΎΠ².
ΠΠ°ΠΊ ΡΠ»Π΅Π΄ΡΠ΅Ρ ΠΈΠ· ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠΎΠ², ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»Π΅Π½Π½ΡΡ Π² Π΄Π°Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅, Π±Π΅Π»ΠΎΠΊ GAGA ΡΡΠ΅Π±ΡΠ΅ΡΡΡ Π½Π° ΡΠ°Π·Π½ΡΡ ΡΡΠ°ΠΏΠ°Ρ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΡΠΉΡΠ΅Π²ΠΎΠΉ ΠΊΠ°ΠΌΠ΅ΡΡ. ΠΠ· Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΠ½ΡΡ ΠΈΡΡΠΎΡΠ½ΠΈΠΊΠΎΠ² ΠΈΠ·Π²Π΅ΡΡΠ΅Π½ ΡΡΠ΄ Π³Π΅Π½ΠΎΠ², ΠΌΡΡΠ°ΡΠΈΠΈ ΠΊΠΎΡΠΎΡΡΡ Π²ΡΠ·ΡΠ²Π°ΡΡ Π½Π°ΡΡΡΠ΅Π½ΠΈΡ, ΠΏΠΎΠ΄ΠΎΠ±Π½ΡΠ΅ ΡΠ΅ΠΌ, ΡΡΠΎ ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½Ρ Ρ Trl ΠΌΡΡΠ°Π½ΡΠΎΠ². Π’Π°ΠΊ, Π΄Π»Ρ ΠΌΡΡΠ°Π½ΡΠΎΠ² ΠΏΠΎ Π³Π΅Π½Π°ΠΌ brainiac ΠΈ egghead Π±ΡΠ»ΠΈ ΠΏΠΎΠΊΠ°Π·Π°Π½Ρ Π½Π°ΡΡΡΠ΅Π½ΠΈΡ Π² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠΎΠ»Π»ΠΈΠΊΡΠ»ΡΡΠ½ΠΎΠ³ΠΎ ΡΠΏΠΈΡΠ΅Π»ΠΈΡ, ΡΡΠΎ ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΊΠ°ΠΌΠ΅Ρ Ρ ΠΎΡΠ»ΠΈΡΠ½ΡΠΌ ΠΎΡ Π½ΠΎΡΠΌΡ ΡΠΈΡΠ»ΠΎΠΌ ΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ (Goode et al., 1996). Π£ ΠΌΡΡΠ°Π½ΡΠΎΠ² ΠΏΠΎ Π³Π΅Π½Π°ΠΌ singed, chickadee ΠΈ quail Π±ΡΠ»ΠΈ Π²ΡΡΠ²Π»Π΅Π½Ρ Π½Π°ΡΡΡΠ΅Π½ΠΈΡ Π² ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π°ΠΊΡΠΈΠ½ΠΎΠ²ΡΡ ΡΠΈΠ»Π°ΠΌΠ΅Π½ΡΠΎΠ² (Paterson and Π’ΠΠ°Π³Π΅, 1991; Cant et al., 1994; Cooley et al., 1992; Mahajan-Miklos and Cooley, 1994). Π£ ΠΌΡΡΠ°Π½ΡΠΎΠ² ΠΏΠΎ Π³Π΅Π½Ρ slbo Π½Π°ΡΡΡΠ΅Π½Π° ΠΌΠΈΠ³ΡΠ°ΡΠΈΡ Π±ΠΎΡΠ΄ΡΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ (Montell et al., 1992). ΠΠ΄Π½Π°ΠΊΠΎ ΡΠ»Π΅Π΄ΡΠ΅Ρ ΠΎΡΠΌΠ΅ΡΠΈΡΡ, ΡΡΠΎ ΠΌΡΡΠ°ΡΠΈΠΈ Π½ΠΈ ΠΎΠ΄Π½ΠΎΠ³ΠΎ ΠΈΠ· Π²ΡΡΠ΅ΠΏΠ΅ΡΠ΅ΡΠΈΡΠ»Π΅Π½Π½ΡΡ Π³Π΅Π½ΠΎΠ² Π½Π΅ ΠΏΡΠΈΠ²ΠΎΠ΄ΡΡ ΠΊ ΡΠ°ΠΊΠΎΠΌΡ ΡΠΈΡΠΎΠΊΠΎΠΌΡ ΡΠΏΠ΅ΠΊΡΡΡ Π΄Π΅ΡΠ΅ΠΊΡΠΎΠ², ΠΊΠΎΡΠΎΡΡΠ΅ Π½Π°Π±Π»ΡΠ΄Π°ΡΡΡΡ Ρ Trl ΠΌΡΡΠ°Π½ΡΠΎΠ².
ΠΡ ΠΏΠΎΠ»Π°Π³Π°Π΅ΠΌ, ΡΡΠΎ Π³Π΅Π½ Trl ΠΈΠ³ΡΠ°Π΅Ρ ΡΠ΅ΡΠ°ΡΡΡΡ ΡΠΎΠ»Ρ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΠΏΡΠΎΡΠ΅ΡΡΠ° ΠΎΠΎΠ³Π΅Π½Π΅Π·Π° Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ, ΠΎΠ½ ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΠ΅Ρ ΡΠ°Π·Π½ΡΠ΅ ΠΏΠΎ Π²ΡΠ΅ΠΌΠ΅Π½ΠΈ ΡΡΠ°ΠΏΡ ΠΎΠΎΠ³Π΅Π½Π΅Π·Π°, Π° Π½Π°ΡΡΡΠ΅Π½ΠΈΠ΅ Π² Π΅Π³ΠΎ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ ΠΎΡΡΠ°ΠΆΠ°Π΅ΡΡΡ Π½Π° ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠ°Π·Π½ΡΡ ΡΠΈΠΏΠΎΠ² ΠΊΠ»Π΅ΡΠΎΠΊ ΡΠΈΡΠ½ΠΈΠΊΠΎΠ².
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