ΠΠ·ΡΡΠ΅Π½ΠΈΠ΅ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ², ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Ρ Drosophila melanogaster
Π‘ΠΎΠ·Π΄Π°Π½Ρ ΠΎΡΠΈΠ³ΠΈΠ½Π°Π»ΡΠ½ΡΠ΅ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΠ΅ ΡΠΈΡΡΠ΅ΠΌΡ, ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡΠΈΠ΅ Π²ΡΡΠ²ΠΈΡΡ Π½ΠΎΠ²ΡΠ΅ ΡΠ²ΠΎΠΉΡΡΠ²Π° ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠΏΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π° ΠΈ ΡΡΠ°Π²Π½ΠΈΡΡ ΡΠ²ΠΎΠΉΡΡΠ²Π° Π½Π΅Π΄Π°Π²Π½ΠΎ ΠΎΡΠΊΡΡΡΡΡ ΠΌΡΡΠ°ΡΠΈΠΉ Enhancer of terminal gene conversion ΠΈ Telomere elongation, Π²Π»ΠΈΡΡΡΠΈΡ Π½Π° ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Ρ Drosophila melanogaster.2. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ Π½Π΅Π³Π°ΡΠΈΠ²Π½ΠΎ Π²Π»ΠΈΡΠ΅Ρ Π½Π° ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΠΎΠ½Π½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠΎΠ²ΠΎΠ³ΠΎ PcG-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ°, ΡΡΠΎ ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΠ΅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΅
- Π¦ΠΠΠ¬ Π ΠΠΠΠΠ§Π ΠΠ‘Π‘ΠΠΠΠΠΠΠΠΠ―
- Π‘ΠΠΠ‘ΠΠ Π‘ΠΠΠ ΠΠ©ΠΠΠΠ
- ΠΠΠΠΠ I. ΠΠΠΠΠ ΠΠΠ’ΠΠ ΠΠ’Π£Π Π«
- 1. 1. Π’Π΅Π»ΠΎΠΌΠ΅ΡΡ, ΠΈΡ ΡΡΠ½ΠΊΡΠΈΠΈ ΠΈ ΡΠΈΠΏΡ
- 1. 2. Π‘ΡΡΡΠΊΡΡΡΠ° ΡΠ΅Π»ΠΎΠΌΠ΅Ρ
- 1. 2. 1. ΠΡΠΏΠΏΠΈΠ½Π³ΠΎΠ²ΡΠΉ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ
- 1. 2. 2. Π’Π΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Π³Π΅ΡΠ΅ΡΠΎΡ ΡΠΎΠΌΠ°ΡΠΈΠ½
- 1. 2. 3. Π’Π΅Π»ΠΎΠΌΠ΅Ρ-Π°ΡΡΠΎΡΠΈΠΈΡΠΎΠ²Π°Π½ΡΠ΅ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ
- 1. 2. 4. ΠΠ΅Ρ Π°Π½ΠΈΠ·ΠΌ ΡΠ΄Π»ΠΈΠ½Π΅Π½ΠΈΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ ΠΈ ΠΊΠΎΠ½ΡΡΠΎΠ»Ρ ΠΈΡ Π΄Π»ΠΈΠ½Ρ
- 1. 3. ΠΠΈΠΏΠΎΡΠ΅Π·Ρ ΠΎΠ± ΡΠ²ΠΎΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ ΡΠ²ΡΠ·ΠΈ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°ΠΌΠΈ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΠΈ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°ΠΌΠΈ, ΠΈΡΠΏΠΎΠ»ΡΠ·ΡΡΡΠΈΠΌΠΈ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°Π·Ρ
- ΠΠΠΠΠ II. ΠΠΠ’ΠΠ ΠΠΠΠ« Π ΠΠΠ’ΠΠΠ«
- 2. 1. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ
- 2. 1. 1. ΠΡΡΠ°ΡΠΈΠΈ ΠΈ Π»ΠΈΠ½ΠΈΠΈ Drosophila melanogaster, ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½Π½ΡΠ΅ Π² ΡΠ°Π±ΠΎΡΠ΅
- 2. 1. 2. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠΊΡΠ΅ΡΠΈΠ²Π°Π½ΠΈΡ
- 2. 2. ΠΠΈΠΎΡ
ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ
- 2. 2. 1. ΠΡΠΈΠ³ΠΎΡΠΎΠ²Π»Π΅Π½ΠΈΠ΅ ΠΊΠΎΠΌΠΏΠ΅ΡΠ΅Π½ΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ Π΄Π»Ρ ΡΡΠ°Π½ΡΡΠΎΡΠΌΠ°ΡΠΈΠΈ ΠΏΠ»Π°Π·ΠΌΠΈΠ΄Π½ΠΎΠΉ ΠΠΠ
- 2. 2. 2. Π’ΡΠ°Π½ΡΡΠΎΡΠΌΠ°ΡΠΈΡ ΠΊΠΎΠΌΠΏΠ΅ΡΠ΅Π½ΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ ΠΏΠ»Π°Π·ΠΌΠΈΠ΄Π°ΠΌΠΈ
- 2. 2. 3. ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ ΠΠΠ ΠΏΠ»Π°Π·ΠΌΠΈΠ΄ ΠΌΠ΅ΡΠΎΠ΄ΠΎΠΌ ΡΠ΅Π»ΠΎΡΠ½ΠΎΠ³ΠΎ Π»ΠΈΠ·ΠΈΡΠ°
- 2. 2. 4. ΠΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π½Π°Ρ ΡΠ΅ΠΏΠ½Π½Π°Ρ ΡΠ΅Π°ΠΊΡΠΈΡ
- 2. 2. 5. Π Π°ΡΡΠ΅ΠΏΠ»Π΅Π½ΠΈΠ΅ ΠΠΠ ΡΠ½Π΄ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Π°ΠΌΠΈ ΡΠ΅ΡΡΡΠΈΠΊΡΠΈΠΈ
- 2. 2. 6. ΠΠ³Π°ΡΠΎΠ·Π½ΡΠΉ Π³Π΅Π»Ρ-ΡΠ»Π΅ΠΊΡΡΠΎΡΠΎΡΠ΅Π·
- 2. 2. 7. ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΠΎΠ² ΠΠΠ ΠΈΠ· Π³Π΅Π»Ρ ΠΈ ΠΎΡΠΈΡΡΠΊΠ° ΠΠΠ ΠΎΡ ΠΏΡΠΎΠ΄ΡΠΊΡΠΎΠ² ΡΠ΅ΡΠΌΠ΅Π½ΡΠ°ΡΠΈΠ²Π½ΡΡ ΡΠ΅Π°ΠΊΡΠΈΠΉ
- 2. 2. 8. ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ Π³Π΅Π½ΠΎΠΌΠ½ΠΎΠΉ ΠΠΠ Drosophila melanogaster
- 2. 2. 9. Π‘Π°ΡΠ·Π΅ΡΠ½-Π±Π»ΠΎΡ Π°Π½Π°Π»ΠΈΠ·
- 2. 2. 10. Π‘ΠΈΠΊΠ²Π΅Π½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅
- 2. 1. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ
- 3. 1. ΠΠ·ΡΡΠ΅Π½ΠΈΠ΅ Π²Π»ΠΈΡΠ½ΠΈΡ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎΠ³ΠΎ Ρ
ΡΠΎΠΌΠ°ΡΠΈΠ½Π° Π½Π° Polycomb— Π·Π°Π²ΠΈΡΠΈΠΌΡΡ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΡ
- 3. 1. 1. ΠΡΠ±ΠΎΡ Π±Π°Π·ΠΎΠ²ΠΎΠΉ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π΄Π»Ρ ΠΈΠ·ΡΡΠ΅Π½ΠΈΡ ΡΠ²ΠΎΠΉΡΡΠ² Polycomb-Π·Π°Π²ΠΈΡΠΈΠΌΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π°
- 3. 1. 2. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½ΠΏΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ, ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΡ Π -ΡΠ»Π΅ΠΌΠ΅Π½Ρ Π² ΠΏΡΠ΅Π΄ΠΏΡΠΎΠΌΠΎΡΠΎΡΠ½ΠΎΠΉ ΠΎΠ±Π»Π°ΡΡΠΈ Π³Π΅Π½Π° yellow
- 3. 1. 3. ΠΠ½Π°Π»ΠΈΠ· /?/Π³ΡΠ£-Π·Π°Π²ΠΈΡΠΈΠΌΠΎΠΉ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΠΈ Π² ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΡ ΠΏΡΠΎΠΈΠ·Π²ΠΎΠ΄Π½ΡΡ Π»ΠΈΠ½ΠΈΡΡ Drosophila melanogaster
- 3. 2. Π€ΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠ΅ ΡΠ°Π·Π΄Π΅Π»Π΅Π½ΠΈΠ΅ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ
ΡΠ°ΠΊΡΠΎΡΠΎΠ² Telomere elongation (Tel) ΠΈ Enhancer of terminal gene conversion (E (tc))
- 3. 2. 1. ΠΡΠ±ΠΎΡ Π±Π°Π·ΠΎΠ²ΠΎΠΉ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π΄Π»Ρ ΡΡΠ°Π²Π½Π΅Π½ΠΈΡ ΡΠ²ΠΎΠΉΡΡΠ² ΠΌΡΡΠ°ΡΠΈΠΉ Telomere elongation (Π’el) ΠΈ Enhancer of terminal gene conversion (E (tc))
- 3. 2. 2. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ Π»ΠΈΠ½ΠΈΠΉ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ, Π½Π΅ΡΡΡΠΈΡ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ ΠΈ Π°ΡΡΠΎΡΠΎΠΌΡ ΠΈΠ· Π»ΠΈΠ½ΠΈΠΈ Gaiano
- 3. 2. 3. ΠΠ½Π°Π»ΠΈΠ· ΠΏΡΠΈΡΠΎΠ΄Ρ ΠΏΡΠΎΠΈΠ·Π²ΠΎΠ΄Π½ΡΡ Π»ΠΈΠ½ΠΈΠΉ, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΡ Π½Π° ΡΠΎΠ½Π΅ ΠΌΡΡΠ°ΡΠΈΠΈ Tel
- 3. 2. 4. ΠΠ½Π°Π»ΠΈΠ· ΡΠΎΠΎΡΠ½ΠΎΡΠ΅Π½ΠΈΡ ΡΠ°ΡΡΠΎΡΡ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎΠΉ Π³Π΅Π½Π½ΠΎΠΉ ΠΊΠΎΠ½Π²Π΅ΡΡΠΈΠΈ ΠΈ ΠΏΡΠΈΡΠΎΠ΅Π΄ΠΈΠ½Π΅Π½ΠΈΠΉ ΠΌΠΎΠ±ΠΈΠ»ΡΠ½ΡΡ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠΎΠ² ΠΊ ΠΊΠΎΠ½ΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ Π² ΡΠΎΠ·Π΄Π°Π½Π½ΠΎΠΉ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΠ΅
- 4. 1. ΠΠ½ΡΠΎΠ³ΠΎΠ½ΠΈΠ·ΠΌ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΌ ΠΈ Polycomb — Π·Π°Π²ΠΈΡΠΈΠΌΡΠΌ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ΠΎΠΌ Π½Π° ΠΊΠΎΠ½ΡΠ΅ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½Π½ΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ Ρ Drosophila melanogaster
- 4. 2. Π€ΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠ΅ ΡΠ°Π·Π΄Π΅Π»Π΅Π½ΠΈΠ΅ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ² Telomere elongation (Π’el) ΠΈ Enhancer of terminal gene conversion (E (tc)), ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Ρ Drosophila melanogaster
ΠΠ·ΡΡΠ΅Π½ΠΈΠ΅ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ², ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Ρ Drosophila melanogaster (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
Π’Π΅Π»ΠΎΠΌΠ΅ΡΡ — ΡΡΠΎ ΡΠΏΠ΅ΡΠΈΠ°Π»ΠΈΠ·ΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ ΠΠΠ-Π±Π΅Π»ΠΊΠΎΠ²ΡΠ΅ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΡ, Π½Π°Ρ ΠΎΠ΄ΡΡΠΈΠ΅ΡΡ Π½Π° ΠΊΠΎΠ½ΡΠ°Ρ Π»ΠΈΠ½Π΅ΠΉΠ½ΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ. Π’Π΅Π»ΠΎΠΌΠ΅ΡΡ ΠΏΡΠ΅Π΄ΠΎΡ ΡΠ°Π½ΡΡΡ ΠΊΠΎΠ½ΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ ΠΎΡ ΡΠ»ΠΈΠΏΠ°Π½ΠΈΡ, Π΄Π΅Π³ΡΠ°Π΄Π°ΡΠΈΠΈ, ΡΠ·Π½Π°Π²Π°Π½ΠΈΡ ΡΠΈΡΡΠ΅ΠΌΠΎΠΉ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΠΠ (Blackburn, 2001; Cech, 2004). Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, ΠΎΡΠ½ΠΎΠ²Π½ΠΎΠΉ ΡΡΠ½ΠΊΡΠΈΠ΅ΠΉ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠ΅Π½ΠΈΠ΅ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ Π³Π΅Π½ΠΎΠΌΠ°. Π Π½Π°ΡΡΠΎΡΡΠ΅Π΅ Π²ΡΠ΅ΠΌΡ Π΄ΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ ΡΡΡΠΎΠ³ΠΎ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΠΎΠ³ΠΎ ΡΠ°Π·ΠΌΠ΅ΡΠ° ΡΠ΅Π»ΠΎΠΌΠ΅Ρ ΠΊΡΠΈΡΠΈΡΠ½ΠΎ Π΄Π»Ρ ΠΆΠΈΠ·Π½Π΅Π΄Π΅ΡΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠ°. ΠΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ ΡΠ΅ΡΠ½ΠΎ ΡΠ²ΡΠ·Π°Π½ΠΎ Ρ ΠΎΠΏΡΡ ΠΎΠ»Π΅ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ ΠΈ ΡΡΠ°ΡΠ΅Π½ΠΈΠ΅ΠΌ ΠΊΠ»Π΅ΡΠΊΠΈ.
Π£ Π±ΠΎΠ»ΡΡΠΈΠ½ΡΡΠ²Π° Π²ΡΡΡΠΈΡ ΡΡΠΊΠ°ΡΠΈΠΎΡ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΡ ΡΠΎΡΡΠΎΡΡ ΠΈΠ· ΠΏΡΠΎΡΡΡΡ ΠΏΠΎΠ²ΡΠΎΡΠΎΠ² G-Π±ΠΎΠ³Π°ΡΠΎΠΉ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ, Π° ΠΈΡ ΡΠ΄Π»ΠΈΠ½Π΅Π½ΠΈΠ΅ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°Π΅ΡΡΡ ΡΠΏΠ΅ΡΠΈΠ°Π»ΡΠ½ΡΠΌ ΡΠ΅ΡΠΌΠ΅Π½ΡΠΎΠΌ — ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°Π·ΠΎΠΉ (Blackburn, 2001; Cech, 2004). Π£ Drosophila melanogaster ΡΠ΅Π»ΠΎΠΌΠ΅ΡΡ ΡΠΎΡΡΠΎΡΡ ΠΈΠ· ΠΌΠΎΠ±ΠΈΠ»ΡΠ½ΡΡ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠΎΠ² ΡΠΈΠΏΠ° LINE, ΠΎΡΠΈΠ΅Π½ΡΠΈΡΠΎΠ²Π°Π½Π½ΡΡ «Π³ΠΎΠ»ΠΎΠ²Π° ΠΊ Ρ Π²ΠΎΡΡΡ» — ΠΠ΅Π’-Π, TART ΠΈ I.
TAHRE (Biessmann & Mason, 2003; Casacuberta & Pardue, 2003aCasacuberta & Pardue, 2003bPardue & DeBaryshe, 2003; Abad et al., 2004aAbad et al, 2004b).
ΠΡΠ½ΠΎΠ²Π½ΡΠΌΠΈ ΡΡΡΡΠΊΡΡΡΠ½ΡΠΌΠΈ Π΅Π΄ΠΈΠ½ΠΈΡΠ°ΠΌΠΈ Π½ΠΎΡΠΌΠ°Π»ΡΠ½ΠΎΠΉ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΡ ΡΠ²Π»ΡΡΡΡΡ: 1) Π’Π΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΡΠΉ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ — ΡΡΠΎ Π±Π΅Π»ΠΊΠΎΠ²ΡΠΉ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ, ΡΠΎΡΠΌΠΈΡΡΡΡΠΈΠΉΡΡ Π½Π° ΠΊΠΎΠ½ΡΠ΅ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ ΠΈ Π·Π°ΡΠΈΡΠ°ΡΡΠΈΠΉ Π΅Π³ΠΎ ΠΎΡ ΡΠ΅ΡΠΌΠ΅Π½ΡΠΎΠ² ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ- 2) Π’Π΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½, ΠΊΠΎΡΠΎΡΡΠΉ ΡΠΎΡΠΌΠΈΡΡΠ΅ΡΡΡ Π½Π° ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΡΡ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΠΎΠΉ ΠΠΠ. ΠΠ°Π½Π½ΡΠ΅ ΡΡΡΡΠΊΡΡΡΡ ΠΈΠ³ΡΠ°ΡΡ ΠΎΡΠ½ΠΎΠ²Π½ΡΡ ΡΠΎΠ»Ρ Π² ΠΏΡΠΎΡΠ΅ΡΡΠ΅ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π΄Π»ΠΈΠ½Ρ ΠΈ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ. ΠΠ°ΠΆΠ½ΠΎΠΉ ΠΎΡΠΎΠ±Π΅Π½Π½ΠΎΡΡΡΡ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ, ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠΎ, ΡΡΠΎ Ρ Π½Π΅Π΅ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ ΠΌΠΎΠΆΠ΅Ρ ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°ΡΡΡΡ Π½Π° Π»ΡΠ±ΠΎΠΉ Π½Π΅ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ½ΠΎΠΉ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΠΠ, Π² ΡΠΎ Π²ΡΠ΅ΠΌΡ ΠΊΠ°ΠΊ Ρ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°Π·ΠΎΠ·Π°Π²ΠΈΡΠΈΠΌΡΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½Π°Ρ ΡΡΡΡΠΊΡΡΡΠ° ΡΠΎΡΠΌΠΈΡΡΠ΅ΡΡΡ ΡΠΎΠ»ΡΠΊΠΎ ΠΏΡΠΈ Π½Π°Π»ΠΈΡΠΈΠΈ ΡΡΡΠΎΠ³ΠΎ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΡ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠ΅ΠΉ, ΠΊΠΎΡΠΎΡΡΠ΅ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΡΡ ΡΠΎΠ±ΠΎΠΉ ΡΠ°ΠΉΡΡ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ Π΄Π»Ρ Π±Π΅Π»ΠΊΠΎΠ², ΡΠΎΡΠΌΠΈΡΡΡΡΠΈΡ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½.
ΠΠ΅ΡΠΊΠΎΠ»ΡΠΊΠΎ Π»Π΅Ρ Π½Π°Π·Π°Π΄ Π±ΡΠ»ΠΈ ΠΏΠΎΠ»ΡΡΠ΅Π½Ρ Π»ΠΈΠ½ΠΈΠΈ Drosophila melanogaster, ΠΊΠΎΡΠΎΡΡΠ΅ Π½Π΅ΡΠ»ΠΈ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΡΠ΅ Π΄Π΅Π»Π΅ΡΠΈΠΈ (Biessmann and Mason 1988; Biessmann et.al. 1990aGolubovsky et.al. 2001; Levis 1989; Mason et.al. 1984; Traverse and Pardue 1988). ΠΡΠ»ΠΎ ΡΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ 4 ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΡΠ°ΠΊΠΆΠ΅ ΠΊΠ°ΠΊ ΠΈ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ Ρ Π½ΠΎΡΠΌΠ°Π»ΡΠ½ΡΠΌΠΈ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°ΠΌΠΈ ΡΠΏΠΎΡΠΎΠ±Π½Ρ ΡΡΡΠΎΠΉΡΠΈΠ²ΠΎ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠΈΠ²Π°ΡΡΡΡ Π² ΡΠ΅ΡΠ΅Π½ΠΈΠ΅ ΠΌΠ½ΠΎΠ³ΠΈΡ ΠΏΠΎΠΊΠΎΠ»Π΅Π½ΠΈΠΉ (Biessmann et.al. 1990Π°Levis 1989). ΠΡΠΎΠΌΠ΅ ΡΠΎΠ³ΠΎ, Π±ΡΠ»ΠΎ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΠ΅Π’-Π ΠΈ TART ΡΠ»Π΅ΠΌΠ΅Π½ΡΡ ΡΠΏΠΎΡΠΎΠ±Π½Ρ ΠΏΡΠΈΡΠΎΠ΅Π΄ΠΈΠ½ΡΡΡΡΡ ΠΊ ΠΊΠΎΠ½ΡΠ°ΠΌ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½ΠΏΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ (Biessmann et.al. 1990b- 1992; Sheen and Levis 1994; Traverse and Pardue 1988). ΠΡΠΈ Π΄Π°Π½Π½ΡΠ΅ ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΡΡ ΡΡΠΎ Ρ Drosophila melanogaster Π½Π° ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΡΡ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½Π½ΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ ΡΠΎΡΠΌΠΈΡΡΠ΅ΡΡΡ Π½ΠΎΡΠΌΠ°Π»ΡΠ½ΡΠΉ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ ΠΈ ΡΠΎΠ±ΠΈΡΠ°Π΅ΡΡΡ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΡΠΉ Π±Π΅Π»ΠΊΠΎΠ²ΡΠΉ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ. ΠΠΎΡΡΠΎΠΌΡ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅ΡΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΠΌΠΎΠ³ΡΡ ΡΠ»ΡΠΆΠΈΡΡ ΡΠ΄ΠΎΠ±Π½ΠΎΠΉ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΠΎΠΉ Π΄Π»Ρ ΠΈΠ·ΡΡΠ΅Π½ΠΈΡ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΡ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΠΉ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ ΠΈ ΠΏΠΎΠΈΡΠΊΠ° ΡΠΎΡΡΠ°Π²Π»ΡΡΡΠΈΡ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΠΎΠ² ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ΅Π»ΠΎΠΌΡΡΠ½ΡΡ ΡΡΡΡΠΊΡΡΡ.
ΠΠ΅ΠΎΠ±Ρ ΠΎΠ΄ΠΈΠΌΠΎ ΠΎΡΠΌΠ΅ΡΠΈΡΡ, ΡΡΠΎ Π½Π΅ΡΠΌΠΎΡΡΡ Π½Π° ΡΠ°Π·Π»ΠΈΡΠΈΡ Π² ΡΡΡΡΠΊΡΡΡΠ΅ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ, Ρ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°Π·ΠΎΠ·Π°Π²ΠΈΠΌΡΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΠΈ Ρ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΡΡΡΠ΅ΡΡΠ²ΡΡΡ ΠΎΠ±ΡΠΈΠ΅ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΡ, ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΠΈΠ΅ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΠ³ΠΎ ΡΠ°Π·ΠΌΠ΅ΡΠ° ΡΠ΅Π»ΠΎΠΌΠ΅Ρ. ΠΠΎ-ΠΏΠ΅ΡΠ²ΡΡ , ΡΠΊΡΠΏΠ΅ΡΠΈΠΌΠ΅Π½ΡΡ, ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½Π½ΡΠ΅ Π½Π° Π΄ΡΠΎΠΆΠΆΠ°Ρ ΠΈ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ , Π΄ΠΎΠΊΠ°Π·Π°Π»ΠΈ, ΡΡΠΎ ΠΏΡΠΈ ΠΈΠ½Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ°Π·Ρ Ρ Π΄Π°Π½Π½ΡΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΠΈΠ½Π΄ΡΡΠΈΡΡΠ΅ΡΡΡ Π°Π»ΡΡΠ΅ΡΠ½Π°ΡΠΈΠ²Π½ΡΠΉ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌ ΡΠ΄Π»ΠΈΠ½Π΅Π½ΠΈΡ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ, ΡΠ²ΡΠ·Π°Π½Π½ΡΠΉ Ρ ΠΏΡΠΎΡΠ΅ΡΡΠ°ΠΌΠΈ ΠΊΠΎΠ½Π²Π΅ΡΡΠΈΠΈΡΠ΅ΠΊΠΎΠΌΠ±ΠΈΠ½Π°ΡΠΈΠΈ. Π ΡΠΎΠΌ ΡΠΈΡΠ»Π΅, Π°Π»ΡΡΠ΅ΡΠ½Π°ΡΠΈΠ²Π½ΡΠ΅ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΡ ΡΠ΄Π»ΠΈΠ½Π΅Π½ΠΈΡ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ ΡΠ°ΡΡΠΎ ΡΠ΅Π°Π»ΠΈΠ·ΡΡΡΡΡ Π² ΡΠ°ΠΊΠΎΠ²ΡΡ ΠΊΠ»Π΅ΡΠΊΠ°Ρ . ΠΠΎ-Π²ΡΠΎΡΡΡ , Π² ΠΏΠΎΡΠ»Π΅Π΄Π½Π΅Π΅ Π²ΡΠ΅ΠΌΡ Π±ΡΠ»ΠΎ Π½Π°ΠΉΠ΄Π΅Π½ΠΎ, ΡΡΠΎ Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ ΠΊΠΎΠ½ΡΠ΅ΡΠ²Π°ΡΠΈΠ²Π½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ, ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΠ΅ Π² ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΠΠ, ΡΠ°ΠΊΠΆΠ΅ ΠΏΡΠΈΠ½ΠΈΠΌΠ°ΡΡ ΡΡΠ°ΡΡΠΈΠ΅ Π² ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ°, ΠΊΠ°ΠΊ Ρ Π΄ΡΠΎΠΆΠΆΠ΅ΠΉ ΠΈ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ , ΡΠ°ΠΊ ΠΈ Ρ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ. ΠΠΎΡΡΠΎΠΌΡ Π΄Π°Π½Π½ΡΠ΅, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ ΠΏΡΠΈ ΠΈΠ·ΡΡΠ΅Π½ΠΈΠΈ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΠΏΠΎΠΌΠΎΠ³ΡΡ ΠΏΠΎΠ½ΡΡΡ Π·Π°ΠΊΠΎΠ½ΠΎΠΌΠ΅ΡΠ½ΠΎΡΡΠΈ ΠΏΡΠΎΡΠ΅ΡΡΠ° ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Ρ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΠΈ Π²ΡΡΠ²ΠΈΡΡ ΠΎΡΠ½ΠΎΠ²Π½ΡΠ΅ Π±Π΅Π»ΠΊΠΈ, ΡΠΎΡΠΌΠΈΡΡΡΡΠΈΠ΅ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ. Π Π½Π°ΡΡΠΎΡΡΠ΅Π΅ Π²ΡΠ΅ΠΌΡ Π½Π°ΠΉΠ΄Π΅Π½ΠΎ ΠΈ ΠΈΠ·ΡΡΠ΅Π½ΠΎ ΡΠΎΠ»ΡΠΊΠΎ Π½Π΅ΡΠΊΠΎΠ»ΡΠΊΠΎ ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ², Π²Ρ ΠΎΠ΄ΡΡΠΈΡ Π² ΡΠΎΡΡΠ°Π² ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΠΎΠ³ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ° Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ: ΡΡΠΎ.
ΠΠ 1 (ΠΎΡΠ½ΠΎΠ²Π½ΠΎΠΉ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½Ρ Π³Π΅ΡΠ΅ΡΠΎΡ ΡΠΎΠΌΠ°ΡΠΈΠ½Π°), ΠΠ ΠΠ (Π±Π΅Π»ΠΎΠΊ, Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΡΡΡΠΈΠΉ Ρ ΠΠ 1), Mrel 1/Rad50 (Π±Π΅Π»ΠΊΠΈ ΡΠΈΡΡΠ΅ΠΌ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΈ ΡΠ΅ΠΊΠΎΠΌΠ±ΠΈΠ½Π°ΡΠΈΠΈ).
ΠΠ°Π½Π½ΡΠ΅, ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π² Π½Π°ΡΠ΅ΠΉ Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΠΈ Π² ΡΠ΅ΡΠ΅Π½ΠΈΠ΅ Π½Π΅ΡΠΊΠΎΠ»ΡΠΊΠΈΡ ΠΏΠΎΡΠ»Π΅Π΄Π½ΠΈΡ Π»Π΅Ρ, ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΡΡ, ΡΡΠΎ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½, ΡΠΎΡΠΌΠΈΡΡΡΡΠΈΠΉΡΡ Ρ Drosophila melanogaster Π½Π° ΠΊΠΎΠ½ΡΠ΅Π²ΡΡ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΡΡ ΠΠΠ Π΄Π»ΠΈΠ½ΠΎΠΉ 4−5 Ρ.ΠΏ.Π½. ΠΎΠ±Π»Π°Π΄Π°Π΅Ρ ΠΎΡΠΎΠ±ΡΠΌΠΈ ΡΠ²ΠΎΠΉΡΡΠ²Π°ΠΌΠΈ (Savitsky et al., 2003; Melnikova et al., 2008). Π Π½Π°ΡΡΠΎΡΡΠ΅ΠΉ ΡΠ°Π±ΠΎΡΠ΅ ΠΌΡ ΠΈΠ·ΡΡΠΈΠ»ΠΈ, ΠΊΠ°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ Π²Π»ΠΈΡΠ΅Ρ Π½Π° ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΡ, Π²ΡΠ·ΡΠ²Π°Π΅ΠΌΡΡ Π±Π΅Π»ΠΊΠ°ΠΌΠΈ Π³ΡΡΠΏΠΏΡ Π½Π° Polycomb. ΠΠ·Π²Π΅ΡΡΠ½ΠΎ, ΡΡΠΎ ΠΌΠ½ΠΎΠ³ΠΈΠ΅ Π±Π΅Π»ΠΊΠΈ Π³ΡΡΠΏΠΏΡ Polycomb ΡΡΠ°ΡΡΠ²ΡΡΡ Π² ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ ΡΡΠ±ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π° ΠΈ ΠΏΠΎΠ΄Π°Π²Π»ΡΡΡ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Π²ΡΡΡΠΎΠ΅Π½Π½ΠΎΠ³ΠΎ Π² Π½Π΅Π³ΠΎ ΡΡΠ°Π½ΡΠ³Π΅Π½Π°. ΠΠ΄Π½Π°ΠΊΠΎ Π²ΠΎΠΏΡΠΎΡ ΠΎ ΡΠΎΠΌ, ΠΊΠ°ΠΊ — ΠΏΠΎΠ·ΠΈΡΠΈΠ²Π½ΠΎ ΠΈΠ»ΠΈ Π½Π΅Π³Π°ΡΠΈΠ²Π½ΠΎ — Π²Π»ΠΈΡΠ΅Ρ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ Π½Π° ΡΠ±ΠΎΡΠΊΡ ΡΡΠ±ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ²ΡΡ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠΎΠ², Π΄ΠΎ ΡΠΈΡ ΠΏΠΎΡ ΠΎΡΡΠ°Π΅ΡΡΡ ΠΎΡΠΊΡΡΡΡΠΌ. ΠΡΠΏΠΎΠ»ΡΠ·ΡΡ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΡ ΡΠΈΡΡΠ΅ΠΌΡ, Π² ΠΊΠΎΡΠΎΡΠΎΠΉ Π -ΡΠ»Π΅ΠΌΠ΅Π½Ρ, Π²ΡΡΡΠΎΠ΅Π½Π½ΡΠΉ ΠΏΠ΅ΡΠ΅Π΄ ΠΏΡΠΎΠΌΠΎΡΠΎΡΠΎΠΌ Π³Π΅Π½Π° yellow ΠΈ Π½Π°Ρ ΠΎΠ΄ΡΡΠΈΠΉΡΡ Π½Π° ΠΊΠΎΠ½ΡΠ΅ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎ Π΄Π΅Π»Π΅Π³ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ, Π² ΠΏΡΠΈΡΡΡΡΡΠ²ΠΈΠΈ ΠΌΡΡΠ°Π½ΡΠ½ΠΎΠ³ΠΎ Π°Π»Π»Π΅Π»Ρ phP1 ΠΌΠΎΠΆΠ΅Ρ ΡΠ»ΡΠΆΠΈΡΡ ΠΌΠ΅ΡΡΠΎΠΌ ΡΠ²ΡΠ·ΡΠ²Π°Π½ΠΈΡ Π±Π΅Π»ΠΊΠΎΠ² PcG, ΠΌΡ ΠΏΠΎΠΊΠ°Π·Π°Π»ΠΈ, ΡΡΠΎ ΠΎΡΠΎΠ±Π°Ρ ΡΡΡΡΠΊΡΡΡΠ° ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π° Π½Π΅Π³Π°ΡΠΈΠ²Π½ΠΎ Π²Π»ΠΈΡΠ΅Ρ Π½Π° ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΠΎΠ½Π½ΠΎΠ³ΠΎ Π ΠΎ1ΡΡΠΎΡΠ¬-Π·Π°Π²ΠΈΡΠΈΠΌΠΎΠ³ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ°. ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΡ, ΡΡΠΎ ΡΡΡΠ΅ΡΡΠ²ΡΠ΅Ρ Π°Π½ΡΠΎΠ³ΠΎΠ½ΠΈΠ·ΠΌ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΌ ΠΈ ΡΡΠ±ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΌ (PcG-Π·Π°Π²ΠΈΡΠΈΠΌΡΠΌ) Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ΠΎΠΌ.
ΠΠΎΠ»ΡΠ½Π΅Π½ΡΡΠ²ΠΎ Π³Π΅Π½ΠΎΠ², ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΠ΅ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ, Π² Π½Π°ΡΡΠΎΡΡΠ΅Π΅ Π²ΡΠ΅ΠΌΡ Π½Π΅ΠΈΠ·Π²Π΅ΡΡΠ½Ρ. ΠΠ΅ΡΠΊΠΎΠ»ΡΠΊΠΎ Π»Π΅Ρ Π½Π°Π·Π°Π΄ Ρ D. melanogaster Π±ΡΠ»ΠΈ Π½Π°ΠΉΠ΄Π΅Π½Ρ Π΄Π²Π° Π΄ΠΎΠΌΠΈΠ½Π°Π½ΡΠ½ΡΡ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠΎΡΠ°, Π²Π»ΠΈΡΡΡΠΈΠ΅ Π½Π° ΡΠ΄Π»ΠΈΠ½Π΅Π½ΠΈΠ΅ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ — ΡΡΠΎ Telomere elongation (Π’el) (Siriaco et al., 2002) ΠΈ Enhancer of terminal gene conversion (E (lc)) (Melnikova & Georgiev, 2002). ΠΠΎΡΡΠΈ ΠΎΠ΄Π½ΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎ ΡΡΠΈ ΠΌΡΡΠ°ΡΠΈΠΈ Π½Π΅Π·Π°Π²ΠΈΡΠΈΠΌΠΎ Π±ΡΠ»ΠΈ ΠΏΠΎΠ»ΡΡΠ΅Π½Ρ Π² ΡΠ°Π·Π½ΡΡ Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠΈΡΡ .
Π ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»Π΅Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΠ΅ Ρ ΠΏΠΎΠΌΠΎΡΡΡ ΠΎΠ΄Π½ΠΎΠΉ ΠΈ ΡΠΎΠΉ ΠΆΠ΅ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π±ΡΠ»ΠΎ ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½ΠΎ ΡΡΠ°Π²Π½Π΅Π½ΠΈΠ΅ ΡΠ²ΠΎΠΉΡΡΠ² Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ² Tel ΠΈ E (tc). ΠΡΠ»ΠΎ ΡΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ Telomere elongation ΠΈ Enhancer of terminal gene conversion ΡΠ²Π»ΡΡΡΡΡ ΡΠ°Π·Π½ΡΠΌΠΈ ΠΌΡΡΠ°ΡΠΈΡΠΌΠΈ.
Π¦ΠΠΠ¬ Π ΠΠΠΠΠ§Π ΠΠ‘Π‘ΠΠΠΠΠΠΠΠΠ―
.
ΠΡΠ½ΠΎΠ²Π½ΡΠΌΠΈ ΡΠ΅Π»ΡΠΌΠΈ Π΄Π°Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΡ ΡΠ²Π»ΡΠ»ΠΈΡΡ: 1) ΠΈΠ·ΡΡΠ΅Π½ΠΈΠ΅ Π²Π»ΠΈΡΠ½ΠΈΡ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π° Π½Π° ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠ±ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ²ΡΡ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠΎΠ²- 2) ΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠ΅ ΡΡΠ°Π²Π½Π΅Π½ΠΈΠ΅ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ² Tel ΠΈ E (tc), Π²Π»ΠΈΡΡΡΠΈΡ Π½Π° ΡΠ΄Π»ΠΈΠ½Π΅Π½ΠΈΠ΅ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Drosophila melanogaster.
Π ΡΠ°Π±ΠΎΡΠ΅ Π±ΡΠ»ΠΈ ΠΏΠΎΡΡΠ°Π²Π»Π΅Π½Ρ ΡΠ»Π΅Π΄ΡΡΡΠΈΠ΅ Π·Π°Π΄Π°ΡΠΈ:
1. Π‘ΠΎΠ·Π΄Π°ΡΡ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΠ΅ ΡΠΈΡΡΠ΅ΠΌΡ, ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡΠΈΠ΅ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΡ ΡΠ²ΠΎΠΉΡΡΠ²Π° ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π° ΠΈ ΡΡΠ°Π²Π½ΠΈΡΡ Π½Π΅Π΄Π°Π²Π½ΠΎ ΠΎΡΠΊΡΡΡΡΠ΅ ΠΌΡΡΠ°ΡΠΈΠΈ Enhancer of terminal gene conversion ΠΈ Telomere elongation.
2. ΠΡΡΡΠ½ΠΈΡΡ ΠΊΠ°ΠΊ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ Π²Π»ΠΈΡΠ΅Ρ Π½Π° ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΠΎΠ½Π½ΠΎΠ³ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ° Π±Π΅Π»ΠΊΠΎΠ² Π³ΡΡΠΏΠΏΡ Polycomb.
3. ΠΠΏΡΠ΅Π΄Π΅Π»ΠΈΡΡ ΡΡΠ°ΡΡΠ½ΠΈΠΊΠΎΠΌ ΠΊΠ°ΠΊΠΎΠ³ΠΎ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ° ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½ΠΈΡ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ (ΠΏΡΠΈΡΠΎΠ΅Π΄ΠΈΠ½Π΅Π½ΠΈΡ ΠΌΠΎΠ±ΠΈΠ»ΡΠ½ΡΡ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠΎΠ² ΠΊ ΠΊΠΎΠ½ΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΡ ΠΈΠ»ΠΈ ΠΊΠΎΠ½Π²Π΅ΡΡΠΈΠΈ/ΡΠ΅ΠΊΠΎΠΌΠ±ΠΈΠ½Π°ΡΠΈΠΈ) ΡΠ²Π»ΡΠ΅ΡΡΡ Π΄ΠΎΠΌΠΈΠ½Π°Π½ΡΠ½ΡΠΉ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ ΡΠ°ΠΊΡΠΎΡ Telomere elongation.
Π‘ΠΠΠ‘ΠΠ Π‘ΠΠΠ ΠΠ©ΠΠΠΠ.
1. ΠΠ¦Π — ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π½Π°Ρ ΡΠ΅ΠΏΠ½Π°Ρ ΡΠ΅Π°ΠΊΡΠΈΡ.
2. Ρ.ΠΏ.Π½. — ΡΡΡΡΡ ΠΏΠ°Ρ Π½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ½ΠΎΠ².
3. ΠΏ.Π½. — ΠΏΠ°Ρ Π½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄ΠΎΠ².
4. TAS — ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Π°ΡΡΠΎΡΠΈΠΈΡΠΎΠ²Π°Π½Π½Π°Ρ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΡ (Telomere associated sequence).
5. PRE — ΡΠ»Π΅ΠΌΠ΅Π½Ρ, ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΉ Π±Π΅Π»ΠΊΠΈ Π³ΡΡΠΏΠΏΡ Polycomb (Polycomb response element).
6. P-Ph — Ρ ΠΈΠΌΠ΅ΡΠ½ΡΠΉ Π±Π΅Π»ΠΎΠΊ P-Polyhomeotic.
7. PcG — Π±Π΅Π»ΠΊΠΈ Π³ΡΡΠΏΠΏΡ Polycomb.
8. BIR — Π½Π΅ΡΠ΅ΡΠΈΠΏΡΠΎΠΊΡΠ½Π°Ρ ΡΠ΅ΠΊΠΎΠΌΠ±ΠΈΠ½Π°ΡΠΈΠΎΠ½Π½ΠΎ-Π·Π°Π²ΠΈΡΠΈΠΌΠ°Ρ ΡΠ΅ΠΏΠ»ΠΈΠΊΠ°ΡΠΈΡ (Breakinduced replication).
9. E (tc) — ΡΠ½Ρ Π°Π½ΡΠ΅Ρ ΡΠ΅ΡΠΌΠΈΠ½Π°Π»ΡΠ½ΠΎΠΉ Π³Π΅Π½Π½ΠΎΠΉ ΠΊΠΎΠ½Π²Π΅ΡΡΠΈΠΈ {Enhancer of terminal gene conversion).
10. Tel — Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ ΡΠ°ΠΊΡΠΎΡ, ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ (!Telomere elongation).
1. Π‘ΠΎΠ·Π΄Π°Π½Ρ ΠΎΡΠΈΠ³ΠΈΠ½Π°Π»ΡΠ½ΡΠ΅ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΠ΅ ΡΠΈΡΡΠ΅ΠΌΡ, ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡΠΈΠ΅ Π²ΡΡΠ²ΠΈΡΡ Π½ΠΎΠ²ΡΠ΅ ΡΠ²ΠΎΠΉΡΡΠ²Π° ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠΏΠΎΠ³ΠΎ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½Π° ΠΈ ΡΡΠ°Π²Π½ΠΈΡΡ ΡΠ²ΠΎΠΉΡΡΠ²Π° Π½Π΅Π΄Π°Π²Π½ΠΎ ΠΎΡΠΊΡΡΡΡΡ ΠΌΡΡΠ°ΡΠΈΠΉ Enhancer of terminal gene conversion ΠΈ Telomere elongation, Π²Π»ΠΈΡΡΡΠΈΡ Π½Π° ΡΠ΅Π³ΡΠ»ΡΡΠΈΡ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ Ρ Drosophila melanogaster.2. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΉ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ Π½Π΅Π³Π°ΡΠΈΠ²Π½ΠΎ Π²Π»ΠΈΡΠ΅Ρ Π½Π° ΡΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠ΅ΠΏΡΠ΅ΡΡΠΈΠΎΠ½Π½ΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠΎΠ²ΠΎΠ³ΠΎ PcG-ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ°, ΡΡΠΎ ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΠ΅Ρ ΠΎ Π½Π°Π»ΠΈΡΠΈΠΈ Π°Π½ΡΠ°Π³ΠΎΠ½ΠΈΠ·ΠΌΠ° ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΌ ΠΈ ΡΡΠ±ΡΠ΅Π»ΠΎΠΌΠ΅ΡΠ½ΡΠΌ Polycomb-Π·Π°Π²ΠΈΡΠΈΠΌΡΠΌ Ρ ΡΠΎΠΌΠ°ΡΠΈΠ½ΠΎΠΌ.3. ΠΡΠΎΠ΄Π΅ΠΌΠΎΠ½ΡΡΡΠΈΡΠΎΠ²Π°Π½ΠΎ, ΡΡΠΎ Π΄ΠΎΠΌΠΈΠ½Π°Π½ΡΠ½ΡΠ΅ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠ°ΠΊΡΠΎΡΡ Enhancer of terminal gene conversion ΠΈ Telomere elongation Π½Π΅ ΡΠ²Π»ΡΡΡΡΡ ΠΎΠ΄Π½ΠΎΠΉ ΠΈ ΡΠΎΠΉ ΠΆΠ΅ ΠΌΡΡΠ°ΡΠΈΠ΅ΠΉ.4. ΠΠΏΠ΅ΡΠ²ΡΠ΅ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ Π²ΡΠΎΡΠ°Ρ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ° Drosophila melanogaster ΠΈΠ· Π»ΠΈΠ½ΠΈΠΈ Gaiano ΡΠΎΠ΄Π΅ΡΠΆΠΈΡ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ ΡΠ°ΠΊΡΠΎΡ, ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΠΉ Π² ΠΊΠΎΠ½ΡΡΠΎΠ»Π΅ Π΄Π»ΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠΌΠ΅Ρ.
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