ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΠΎΠ»ΠΈ dCTCF Π² ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΠΈ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ bithorax-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ° Drosophila melanogaster
Π£ΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ Π² ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ ΠΠ₯-Π‘, ΠΏΠΎΠΌΠΈΠΌΠΎ dCTCF, Π²ΠΎΠ²Π»Π΅ΡΠ΅Π½Ρ ΠΏΠΎΠΊΠ° Π½Π΅ ΠΈΠ΄Π΅Π½ΡΠΈΡΠΈΡΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ. Π¦Π΅Π»ΡΡ ΡΠ°Π±ΠΎΡΡ ΡΡΠ°Π»ΠΎ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΠΎΠ»ΠΈ dCTCF Π² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ Π³ΡΠ°Π½ΠΈΡ Π»ΠΎΠΊΡΡΠ° bithorax Drosophila melanogaster. ΠΡ ΠΏΡΠΎΡΠ΅ΡΡΠΈΡΠΎΠ²Π°Π»ΠΈ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΡ Π³ΡΠ°Π½ΠΈΡ. ΠΡΡΠ²Π»Π΅Π½Π° ΡΠ΅Π»Π΅ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ ΠΏΡΠ΅Π΄ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ Π³Π΅Π½Π° Abd-B Ρ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ Π³ΡΠ°Π½ΠΈΡ ΡΠ²ΠΎΠ΅ΠΉ… 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΅
- Π‘ΠΠΠ‘ΠΠ Π‘ΠΠΠ ΠΠ©ΠΠΠΠ
- 1. ΠΠΠΠΠ ΠΠΠ’ΠΠ ΠΠ’Π£Π Π«
- 1. 1. Π Π΅Π³ΡΠ»ΡΡΠΈΡ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΈ Π² biihorax-KommQK.ce Drosophila melanogaster
- 1. 1. 1. Π‘ΡΡΡΠΊΡΡΡΠ° ΠΠ₯-Π‘
- 1. 1. 2. Π€Π°Π·Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΠΠ₯-Π‘
- 1. 1. 3. ΠΡΠ°Π½ΠΈΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ ΠΠ₯-Π‘
- 1. 1. 3. 1. Fab
- 1. 1. 3. 2. Fab
- 1. 1. 3. 3. ΠΠ΅Ρ
- 1. 1. 3. 4. Fab
- 1. 1. 4. ΠΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΠ΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ ΠΠ₯-Π‘
- 1. 1. 4. 1. ΠΠ»Π΅ΠΌΠ΅Π½Ρ PTS (Promotor Targeting Sequence)
- 1. 1. 4. 2. ΠΠ»Π΅ΠΌΠ΅Π½Ρ PTE (Promoter Tetherig Element)
- 1. 1. 4. 3. ΠΠΎΠΌΠΌΡΠ½ΠΈΠΊΠ°ΡΠΎΡΠ½Π°Ρ ΡΠΎΠ»Ρ ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΠΎΠ²
- 1. 1. Π Π΅Π³ΡΠ»ΡΡΠΈΡ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΈ Π² biihorax-KommQK.ce Drosophila melanogaster
- 1. 2. ΠΠ΅Π»ΠΊΠΈ, ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΠ΅ Π² ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΠΈ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΡ
Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ
- 1. 2. 1. CTCF
- 1. 2. 1. 1. CTCF ΠΊΠ°ΠΊ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΎΠ½Π½ΡΠΉ ΡΠ°ΠΊΡΠΎΡ
- 1. 2. 1. 2. ΠΠ½ΡΡΠ»ΡΡΠΎΡΠ½Π°Ρ ΡΠΎΠ»Ρ CTCF
- 1. 2. 1. 3. CTCF ΠΈ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΠ΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ
- 1. 2. 2. ΠΠΎΠΌΠΎΠ»ΠΎΠ³ CTCF Ρ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ
- 1. 2. 1. CTCF
- 2. 1. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ
- 2. 1. 1. ΠΠΈΠ½ΠΈΠΈ Drosophila melanogaster
- 2. 1. 2. Π’ΡΠ°Π½ΡΡΠΎΡΠΌΠ°ΡΠΈΡ ΡΠΌΠ±ΡΠΈΠΎΠ½ΠΎΠ² Drosophila melanogaster ΠΈ ΠΏΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΡΡΠ°Π½ΡΠ³Π΅Π½Π½ΡΡ Π»ΠΈΠ½ΠΈΠΉ
- 2. 1. 3. Π€Π΅Π½ΠΎΡΠΈΠΏΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½ΠΎΠ² yellow ΠΈ white Π² ΡΡΠ°Π½ΡΠ³Π΅Π½Π½ΡΡ Π»ΠΈΠ½ΠΈΡΡ
- 2. 1. 4. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠΊΡΠ΅ΡΠΈΠ²Π°Π½ΠΈΡ
- 2. 2. ΠΠΈΠΎΡ
ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ
- 2. 2. 1. ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ ΠΠΠ ΠΈΠ· Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ
- 2. 2. 2. Π‘Π°ΡΠ·Π΅ΡΠ½-Π±Π»ΠΎΡ-Π°Π½Π°Π»ΠΈΠ·
- 2. 2. 3. ΠΠ΅ΡΠΎΠ΄ ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π½ΠΎΠΉ ΡΠ΅ΠΏΠ½ΠΎΠΉ ΡΠ΅Π°ΠΊΡΠΈΠΈ (ΠΠ¦Π )
- 2. 2. 4. ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎΠ΅ ΠΊΠ»ΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅
- 2. 2. 5. Π’ΡΠ°Π½ΡΡΠΎΡΠΌΠ°ΡΠΈΡ Π±Π°ΠΊΡΠ΅ΡΠΈΠ°Π»ΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ ΠΏΠ»Π°Π·ΠΌΠΈΠ΄Π°ΠΌΠΈ
- 2. 2. 6. ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ ΠΠΠ ΠΏΠ»Π°Π·ΠΌΠΈΠ΄ ΠΌΠ΅ΡΠΎΠ΄ΠΎΠΌ ΡΠ΅Π»ΠΎΡΠ½ΠΎΠ³ΠΎ Π»ΠΈΠ·ΠΈΡΠ°
- 2. 2. 7. ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΠΎΠ² ΠΠΠ ΠΈΠ· Π°Π³Π°ΡΠΎΠ·Π½ΠΎΠ³ΠΎ Π³Π΅Π»Ρ
- 2. 2. 8. Π‘ΠΎΠ·Π΄Π°Π½ΠΈΠ΅ ΠΊΠΎΠ½ΡΡΡΡΠΊΡΠΈΠΉ
- 2. 2. 9. ΠΠ΅ΡΠΎΠ΄ ΡΠΎΡΠΌΠΎΠΆΠ΅Π½ΠΈΡ ΠΠΠ-Π±Π΅Π»ΠΊΠΎΠ²ΡΡ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠΎΠ² Π² Π³Π΅Π»Π΅ (EMSA)
- 2. 2. 10. ΠΠΌΠΌΡΠ½ΠΎΠΏΡΠ΅ΡΠΈΠΏΠΈΡΠ°ΡΠΈΡ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π°
- 3. 1. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π³ΡΠ°Π½ΠΈΡ Fab-3, Fab-4 ΠΈ Fab-6 Π½Π° ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ Π±Π»ΠΎΠΊΠΈΡΠΎΠ²Π°ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ½Ρ Π°Π½ΡΠ΅ΡΠ°ΠΌΠΈ ΠΈ ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ°ΠΌΠΈ
- 3. 2. Fab-3, Fab-4 ΠΈ Fab-6 ΡΠΏΠΎΡΠΎΠ±Π½Ρ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠΈΠ²Π°ΡΡ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΠ΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ
- 3. 3. Π‘ΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ Π³ΡΠ°Π½ΠΈΡ ΠΠ΅Ρ, Fab-6, PTS/F8 ΠΊ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΠΌ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡΠΌ Π½Π΅ Π·Π°Π²ΠΈΡΠΈΡ ΠΎΡ dCTCF
- 3. 4. ΠΠ·ΡΡΠ΅Π½ΠΈΠ΅ ΡΠΎΠ»ΠΈ dCTCF Π²ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡΡ ΠΌΠ΅ΠΆΠ΄Ρ ΠΏΡΠ΅Π΄ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΡΡ remAbd-B ΠΈ Π³ΡΠ°Π½ΠΈΡΠ°ΠΌΠΈ PTS/Fab-8 ΠΈ Fab
- 3. 5. Π£ΡΠ°ΡΡΠΎΠΊ ΠΏΡΠ΅Π΄ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ Π³Π΅Π½Π° Abd-B ΡΠ΅Π»Π΅ΠΊΡΠΈΠ²Π½ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΠ΅Ρ Ρ Π³ΡΠ°Π½ΠΈΡΠ°ΠΌΠΈ ΡΠ²ΠΎΠ΅ΠΉ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ
- 4. 1. Π‘Π²ΠΎΠΉΡΡΠ²Π° Π½ΠΎΠ²ΡΡ Π³ΡΠ°Π½ΠΈΡ Fab-3, Fab-4 ΠΈ Fab
- 4. 2. Π ΠΎΠ»Ρ dCTCF Π² ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΠΈ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ Π² 6 zY/zorax-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ΅ Drosophila melanogaster
- 4. 3. Π‘Π΅Π»Π΅ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ Π² bithorax-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ΅
ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΠΎΠ»ΠΈ dCTCF Π² ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΠΈ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ bithorax-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ° Drosophila melanogaster (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
ΠΠ»Ρ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΡ ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ ΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΡΠ°Π·Π½ΠΎΠΎΠ±ΡΠ°Π·ΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ, ΡΠΎΡΠΌΠΈΡΡΡΡΠ΅Π³ΠΎΡΡ Π² Ρ ΠΎΠ΄Π΅, ΡΠ°Π·Π²ΠΈΡΠΈΡ ΠΌΠ½ΠΎΠ³ΠΎΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠ°, Π½Π΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΡ ΡΠΎΡΠ½ΡΠ΅ ΠΈ Π²ΡΡΠΎΠΊΠΎΡΡΡΠ΅ΠΊΡΠΈΠ²Π½ΡΠ΅ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ Π³Π΅Π½ΠΎΠ². ΠΠ΅Π½ΠΎΠΌΡ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΠΌΠΎΠ³ΡΡ ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΡ Π΄Π΅ΡΡΡΠΊΠΈ ΡΡΡΡΡ Π³Π΅Π½ΠΎΠ², Π΄Π»Ρ ΠΊΠ°ΠΆΠ΄ΠΎΠ³ΠΎ ΠΈΠ· ΠΊΠΎΡΠΎΡΡΡ ΡΡΡΠ΅ΡΡΠ²ΡΠ΅Ρ ΠΏΡΠΎΠ³ΡΠ°ΠΌΠΌΠ° ΡΠ΅Π°Π»ΠΈΠ·Π°ΡΠΈΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠΉ ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΈ, Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΠ·ΡΡΡΠ°ΡΡΡ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΠΌ Π²ΡΠ΅ΠΌΠ΅Π½Π΅ΠΌ, ΠΌΠ΅ΡΡΠΎΠΌ ΠΈ ΡΡΠΎΠ²Π½Π΅ΠΌ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ. ΠΠ΄Π½ΠΈΠΌ ΠΈΠ· Π²Π°ΠΆΠ½Π΅ΠΉΡΠΈΡ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠΎΠ², ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΠΈΡ ΠΏΡΠ°Π²ΠΈΠ»ΡΠ½ΡΡ ΡΠ°Π±ΠΎΡΡ Π³Π΅Π½ΠΎΠ², ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΈ. ΠΠ° ΡΡΠ΅Ρ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Π±Π΅Π»ΠΊΠΎΠ²ΡΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ² Ρ ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ°ΠΌΠΈ Π³Π΅Π½ΠΎΠ² ΠΈ Π΄ΡΡΠ³ΠΈΠΌΠΈ Π³/ΠΌΡ-ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ ΠΠΠ (ΡΠ½Ρ Π°Π½ΡΠ΅ΡΠ°ΠΌΠΈ ΠΈΠ»ΠΈ ΡΠ°ΠΉΠ»Π΅Π½ΡΠ΅ΡΠ°ΠΌΠΈ) ΠΏΡΠΎΠΈΡΡ ΠΎΠ΄ΠΈΡ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΡ ΠΈΠ»ΠΈ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΡ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΈ, ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΠΎ.
Π‘ΡΠΈΡΠ°Π΅ΡΡΡ, ΡΡΠΎ ΡΠΊΠΎΠΎΡΠ΄ΠΈΠ½ΠΈΡΠΎΠ²Π°Π½Π½Π°Ρ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΡ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°Π΅ΡΡΡ, Π² ΡΠ°ΡΡΠ½ΠΎΡΡΠΈ, ΠΏΡΡΠ΅ΠΌ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΠΈ Π³Π΅Π½ΠΎΠ² Π² Π½Π΅Π·Π°Π²ΠΈΡΠΈΠΌΡΠ΅ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ΠΎΠ²ΡΠ΅ Π΄ΠΎΠΌΠ΅Π½Ρ Ρ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΠΌΠΈ ΠΏΠ°ΡΡΠ΅ΡΠ½Π°ΠΌΠΈ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ (Dillon & Sabbattini, 2000, Spellman & Rubin, 2002). Π‘ΡΡΠ΅ΡΡΠ²ΡΠ΅Ρ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠ΅Π½ΠΈΠ΅ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ Π² ΡΡΠΎΠΌ ΠΏΡΠΎΡΠ΅ΡΡΠ΅ Π²Π°ΠΆΠ½ΡΡ ΡΠΎΠ»Ρ ΠΈΠ³ΡΠ°ΡΡ ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΡ — z/wc-ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠ΅ ΡΠ»Π΅ΠΌΠ΅Π½ΡΡ, ΠΊΠΎΡΠΎΡΡΠ΅ ΠΎΠ±Π»Π°Π΄Π°ΡΡ Π΄Π²ΡΠΌΡ ΠΎΡΠ½ΠΎΠ²Π½ΡΠΌΠΈ ΡΠ²ΠΎΠΉΡΡΠ²Π°ΠΌΠΈ: (1) ΠΎΠ½ΠΈ Π±Π»ΠΎΠΊΠΈΡΡΡΡ Π°ΠΊΡΠΈΠ²ΠΈΡΡΡΡΠ΅Π΅ Π΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ ΡΠ½Ρ Π°Π½ΡΠ΅ΡΠ°, Π΅ΡΠ»ΠΈ ΡΠ°ΡΠΏΠΎΠ»ΠΎΠΆΠ΅Π½Ρ ΠΌΠ΅ΠΆΠ΄Ρ Π½ΠΈΠΌ ΠΈ ΠΏΡΠΎΠΌΠΎΡΠΎΡΠΎΠΌ, Π° ΡΠ°ΠΊΠΆΠ΅ (2) ΡΠ²Π»ΡΡΡΡΡ Π±Π°ΡΡΠ΅ΡΠ°ΠΌΠΈ, ΠΏΡΠ΅Π΄ΠΎΡΠ²ΡΠ°ΡΠ°ΡΡΠΈΠΌΠΈ ΡΠ°ΡΠΏΡΠΎΡΡΡΠ°Π½Π΅Π½ΠΈΠ΅ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΠΎΠ½Π½ΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π° (Cai & Levine, 1995, Gerasimova & Corees, 1996, Kuhn & Geyer, 2003). Π ΠΏΠΎΡΠ»Π΅Π΄Π½Π΅Π΅ Π²ΡΠ΅ΠΌΡ ΠΏΠΎΡΠ²Π»ΡΠ΅ΡΡΡ Π²ΡΠ΅ Π±ΠΎΠ»ΡΡΠ΅ Π΄Π°Π½Π½ΡΡ , ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΡΡΠΈΡ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ ΡΠ½Ρ Π°Π½ΡΠ΅ΡΡ ΠΈ ΠΏΡΠΎΠΌΠΎΡΠΎΡΡ ΡΠ±Π»ΠΈΠΆΠ΅Π½Ρ Π² ΠΏΡΠΎΡΡΡΠ°Π½ΡΡΠ²Π΅, Π° ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΡ ΠΈΠ³ΡΠ°ΡΡ ΠΊΠ»ΡΡΠ΅Π²ΡΡ ΡΠΎΠ»Ρ Π² ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠΈ ΡΡΠΈΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ (Cleard et al., 2006).
Π£Π΄ΠΎΠ±Π½ΠΎΠΉ ΠΌΠΎΠ΄Π΅Π»ΡΡ Π΄Π»Ρ ΠΈΠ·ΡΡΠ΅Π½ΠΈΡ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΈ ΡΠ²Π»ΡΠ΅ΡΡΡ bithorax-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ (ΠΠ₯-Π‘) Drosophila melanogaster. ΠΠ΅Π½Ρ ΠΠ₯-Π‘: Ultrabithorax (Ubx), abdominal, Π (abd-A) ΠΈ Abdominal Π (.Abd-B), — ΠΎΡΠ²Π΅ΡΠ°ΡΡ Π·Π° ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠ°ΡΡΠΈ ΡΡΠ΅ΡΡΠ΅Π³ΠΎ Π³ΡΡΠ΄Π½ΠΎΠ³ΠΎ ΠΈ Π²ΡΠ΅Ρ Π±ΡΡΡΠ½ΡΡ ΡΠ΅Π³ΠΌΠ΅Π½ΡΠΎΠ². ΠΡΠΈ Π³Π΅Π½Ρ ΡΠ΅Π³ΡΠ»ΠΈΡΡΡΡΡΡ ΡΠΊΠ°Π½Π΅ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ½ΡΠΌΠΈ ΡΠ½Ρ Π°Π½ΡΠ΅ΡΠ°ΠΌΠΈ (abx/bx, bxd/pbx, iab-2 — iab-8), ΡΠ°ΡΠΏΠΎΠ»ΠΎΠΆΠ΅Π½Π½ΡΠΌΠΈ Π² ΠΏΠΎΡΡΠ΄ΠΊΠ΅ ΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΡΠ΅Π³ΠΌΠ΅Π½ΡΠΎΠ², ΠΊΠΎΡΠΎΡΡΠ΅ ΠΎΠ½ΠΈ ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡ. ΠΠ½Ρ Π°Π½ΡΠ΅ΡΡ ΠΎΡΠ΄Π΅Π»Π΅Π½Ρ Π΄ΡΡΠ³ ΠΎΡ Π΄ΡΡΠ³Π° ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ Π³ΡΠ°Π½ΠΈΡΠ°ΠΌΠΈ. Π ΡΠΎΡΡΠ°Π²Π΅ ΡΡΠ΅Ρ ΠΈΠ·ΡΡΠ΅Π½Π½ΡΡ ΡΠ°Π½Π΅Π΅ Π³ΡΠ°Π½ΠΈΡ (ΠΠ΅Ρ, Fab-7 ΠΈ PTS/Fab-8) Π±ΡΠ»ΠΈ Π½Π°ΠΉΠ΄Π΅Π½Ρ ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΡ. ΠΡΠ»ΠΎ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΠ½Π°Ρ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Π³ΡΠ°Π½ΠΈΡΡ PTS/Fab-8 Π·Π°Π²ΠΈΡΠΈΡ ΠΎΡ Π½Π°Π»ΠΈΡΠΈΡ ΡΠ°ΠΉΡΠΎΠ² ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ Π±Π΅Π»ΠΊΠ° dCTCF, ΠΊΠΎΡΠΎΡΡΠΉ ΡΠ²Π»ΡΠ΅ΡΡΡ Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΎΠΌ ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΠ½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° ΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ — CTCF (Moon et al., 2005). ΠΠ΅Π΄Π°Π²Π½ΠΎ ΡΠ°ΠΉΡΡ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ dCTCF Π±ΡΠ»ΠΈ ΠΊΠ°ΡΡΠΈΡΠΎΠ²Π°Π½Ρ ΠΌΠ΅ΠΆΠ΄Ρ /ΠΎΠ±-Π΄ΠΎΠΌΠ΅Π½Π°ΠΌΠΈ bithorax-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ° (Holohan et al., 2007, Smith et al., 2009). ΠΡΠΎ ΠΏΠΎΠ·Π²ΠΎΠ»ΠΈΠ»ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ ΡΡΡΠ΅ΡΡΠ²ΠΎΠ²Π°Π½ΠΈΠ΅ Π½ΠΎΠ²ΡΡ ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΠΎΠ² Π² bithorax-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ΅ (Fab-3, Fab-4 ΠΈ Fab-6).
ΠΠ»Ρ CTCF ΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ Π±ΡΠ»Π° ΠΏΠΎΠΊΠ°Π·Π°Π½Π° ΡΠΎΠ»Ρ Π² ΡΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΈΠΈ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΄Π°Π»Π΅Π½Π½ΡΠΌΠΈ ΡΡΠ°ΡΡΠΊΠ°ΠΌΠΈ Π³Π΅Π½ΠΎΠΌΠ° (Kurukuti et al., 2006, Murrell et al., 2004, Splinter et al., 2006). Π£ΡΠ°ΡΡΠΈΠ΅ dCTCF Π² Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡΡ Π±ΡΠ»ΠΎ ΠΏΡΠΎΠ΄Π΅ΠΌΠΎΠ½ΡΡΡΠΈΡΠΎΠ²Π°Π½ΠΎ ΡΠ°Π½Π΅Π΅ Π² Π½Π°ΡΠ΅ΠΉ Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ (Kyrchanova et al., 2008). ΠΠΎΠΆΠ½ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ dCTCF ΠΈΠ³ΡΠ°Π΅Ρ Π²Π°ΠΆΠ½ΡΡ ΡΠΎΠ»Ρ Π² ΡΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΈΠΈ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ Π² Π»ΠΎΠΊΡΡΠ΅ ΠΠ₯-Π‘. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π΄Π°Π½Π½ΠΎΠ³ΠΎ Π²ΠΎΠΏΡΠΎΡΠ° ΡΠ²Π»ΡΠ΅ΡΡΡ Π²Π°ΠΆΠ½ΡΠΌ Π΄Π»Ρ ΠΏΠΎΠ½ΠΈΠΌΠ°Π½ΠΈΡ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΈ Π³Π΅Π½ΠΎΠ².
Π¦Π΅Π»ΡΡ ΡΠ°Π±ΠΎΡΡ ΡΡΠ°Π»ΠΎ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΠΎΠ»ΠΈ dCTCF Π² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ Π³ΡΠ°Π½ΠΈΡ Π»ΠΎΠΊΡΡΠ° bithorax Drosophila melanogaster. ΠΡ ΠΏΡΠΎΡΠ΅ΡΡΠΈΡΠΎΠ²Π°Π»ΠΈ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΡ Π³ΡΠ°Π½ΠΈΡ.
Π Π°Π¬-Π, Π Π°Π¬-4 ΠΈ Π Π°Π¬-6, ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΠ΅ ΡΠ°ΠΉΡΡ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ <1Π‘Π’Π‘Π , Π½Π° ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ ΠΊ ΠΈΠ½ΡΡΠ»ΡΡΠΈΠΈ. ΠΡΠΎΠΌΠ΅ ΡΡΠΎΠ³ΠΎ, ΠΌΡ ΠΈΠ·ΡΡΠΈΠ»ΠΈ ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΡ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠΎΠ² ΠΠ₯-Π‘ ΠΊ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΡ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΈ ΡΠΎΠ»Ρ Ρ1Π‘Π’Π‘Π Π² ΡΡΠΎΠΌ ΠΏΡΠΎΡΠ΅ΡΡΠ΅.
Π²ΡΠ²ΠΎΠ΄Ρ.
1. ΠΠΏΠ΅ΡΠ²ΡΠ΅ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ Π² ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΠ΅ Π½Π° ΠΎΡΠ½ΠΎΠ²Π΅ Π³Π΅Π½ΠΎΠ² yellow ΠΈ white ΡΡΠ°Π³ΠΌΠ΅Π½ΡΡ Π³ΡΠ°Π½ΠΈΡ Fab-3, Fab-4 ΠΈ Fab-6, ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΠ΅ ΡΠ°ΠΉΡΡ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ dCTCF, Π½Π΅ ΠΏΡΠΎΡΠ²Π»ΡΡΡ ΡΠ²ΠΎΠΉΡΡΠ² ΠΈΠ½ΡΡΠ»ΡΡΠΎΡΠΎΠ², a Fab-4 ΠΈ Fab-Π± ΠΎΠ±Π»Π°Π΄Π°ΡΡ ΡΠ²ΠΎΠΉΡΡΠ²Π°ΠΌΠΈ ΡΠ°ΠΉΠ»Π΅Π½ΡΠ΅ΡΠΎΠ².
2. ΠΡΠΎΠ΄Π΅ΠΌΠΎΠ½ΡΡΡΠΈΡΠΎΠ²Π°Π½ΠΎ, ΡΡΠΎ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΡ Π³ΡΠ°Π½ΠΈΡ Fab-3, Fab-4 ΠΈ Fab-6 ΡΠΏΠΎΡΠΎΠ±Π½Ρ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠΈΠ²Π°ΡΡ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΠ΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ.
3. Π£ΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ Π³ΡΠ°Π½ΠΈΡ ΠΠ΅Ρ, Fab-6, PTS/F8 ΠΊ Π΄ΠΈΡΡΠ°Π½ΡΠΈΠΎΠ½Π½ΡΠΌ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡΠΌ Π½Π΅ Π·Π°Π²ΠΈΡΠΈΡ ΠΎΡ dCTCF.
4. ΠΡΡΠ²Π»Π΅Π½Π° ΡΠ΅Π»Π΅ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ ΠΏΡΠ΅Π΄ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ Π³Π΅Π½Π° Abd-B Ρ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ Π³ΡΠ°Π½ΠΈΡ ΡΠ²ΠΎΠ΅ΠΉ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ.
5. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ dCTCF Π½Π΅ ΡΡΠ°ΡΡΠ²ΡΠ΅Ρ Π² ΡΠ΅Π»Π΅ΠΊΡΠΈΠ²Π½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡΡ ΠΏΡΠ΅Π΄ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ ΡΠ΅ΠΏΠ°. Abd-B Ρ Π³ΡΠ°Π½ΠΈΡΠ°ΠΌΠΈ Fab-6, Fab-7 ΠΈ PTS/F8.
6. Π£ΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ Π² ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠ½ΡΠΌΠΈ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠ°ΠΌΠΈ ΠΠ₯-Π‘, ΠΏΠΎΠΌΠΈΠΌΠΎ dCTCF, Π²ΠΎΠ²Π»Π΅ΡΠ΅Π½Ρ ΠΏΠΎΠΊΠ° Π½Π΅ ΠΈΠ΄Π΅Π½ΡΠΈΡΠΈΡΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ.
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