ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· Π»ΠΎΠΊΡΡΠ° Delta Ρ Drosophila virilis
ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π½Π΅ΠΉΡΠΎΠ³Π΅Π½Π½ΡΡ Π³Π΅Π½ΠΎΠ² ΠΌΠΎΠΆΠ΅Ρ ΠΏΠΎΠΌΠΎΡΡ Π² ΡΠ΅ΡΠ΅Π½ΠΈΠΈ ΠΏΡΠ°ΠΊΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ΅Π΄ΠΈΡΠΈΠ½ΡΠΊΠΈΡ ΠΏΡΠΎΠ±Π»Π΅ΠΌ. ΠΠ±Π½Π°ΡΡΠΆΠ΅Π½ΠΎ, ΡΡΠΎ Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ ΠΏΠ°ΡΠΎΠ»ΠΎΠ³ΠΈΠΈ Ρ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° (Π±ΠΎΠ»Π΅Π·Π½Ρ ΠΠ»ΡΡΠ³Π΅ΠΉΠΌΠ΅ΡΠ°, ΡΠΈΠ½Π΄ΡΠΎΠΌ CADASIL, Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ ΡΠ°ΠΊΠΎΠ²ΡΠ΅ ΠΎΠΏΡΡ ΠΎΠ»ΠΈ) ΠΈ Π΄ΡΡΠ³ΠΈΡ ΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ ΡΠ²ΡΠ·Π°Π½Ρ Ρ ΠΌΡΡΠ°ΡΠΈΡΠΌΠΈ Π² Π³Π΅Π½Π°Ρ , Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ½ΡΡ Delta ΠΈ Notch, ΠΈΠ»ΠΈ Ρ Π΄ΡΡΠ³ΠΈΠΌΠΈ Π΄Π΅ΡΠ΅ΠΊΡΠ°ΠΌΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ Delta-Notch-cnmajihtioro ΠΏΡΡΠΈ (Ellisen et al., 1991; Wong, 1997; Struhl et al… 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. ΠΡΠ½ΠΎΠ²Π½ΡΠ΅ ΡΡΠ°ΠΏΡ ΡΠ°Π½Π½Π΅Π³ΠΎ Π½Π΅ΠΉΡΠΎΠ³Π΅Π½Π΅Π·Π° Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ
- 1. 1. Π€ΠΎΡΠΌΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ Π½Π΅ΠΉΡΠΎΠ³Π΅Π½Π½ΠΎΠΉ Π·Π°ΠΊΠ»Π°Π΄ΠΊΠΈ
- 1. 2. ΠΠΈΡΡΠ΅ΡΠ΅Π½ΡΠΈΡΠΎΠ²ΠΊΠ° Π½Π΅ΠΉΡΠΎΠ±Π»Π°ΡΡΠΎΠ²
- 1. 3. ΠΠΈΡΡΠ΅ΡΠ΅Π½ΡΠΈΡΠΎΠ²ΠΊΠ° ΠΊΠ»Π΅ΡΠΎΠΊ — ΠΏΡΠ΅Π΄ΡΠ΅ΡΡΠ²Π΅Π½Π½ΠΈΠΊΠΎΠ² ΡΠ΅Π½ΡΠΎΡΠ½ΡΡ ΠΎΡΠ³Π°Π½ΠΎΠ²
- 2. ΠΠ΅Π½Ρ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΠ΅ ΡΠ°Π·Π²ΠΈΡΠΈΠ΅ Π½Π΅ΡΠ²Π½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ
- 3. ΠΡΠ½ΠΎΠ²Π½ΡΠ΅ Π³Π΅Π½Ρ Delta-Notch ΡΠΈΠ³Π½Π°Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ D. melanogaster
- 3. 1. ΠΠΎΠΊΡΡ Notch
- 3. 2. ΠΠΎΠΊΡΡ Delta
- 3. 3. ΠΠΎΠΊΡΡ Serrate
- 4. Π£ΡΠ°ΡΡΠΈΠ΅ Delta-Notch ΡΠΈΠ³Π½Π°Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π² ΠΆΠΈΠ·Π½Π΅Π΄Π΅ΡΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ²
- 4. 1. ΠΠ»Π΅ΠΉΠΎΡΡΠΎΠΏΠ½ΠΎΠ΅ Π΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ Π³Π΅Π½ΠΎΠ² Delta ΠΈ Notch Ρ D. melanogaster
- 4. 2. ΠΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈ Π³Π΅Π½ΠΎΠ² Delta-Notch ΡΠΈΡΡΠ΅ΠΌΡ Ρ Π΄ΡΡΠ³ΠΈΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ²
- 4. 3. Π€ΡΠ½ΠΊΡΠΈΠΈ Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΎΠ² Π³Π΅Π½ΠΎΠ² Delta ΠΈ Notch
- 5. ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΠ΅ ΠΌΠ΅Ρ
Π°Π½ΠΈΠ·ΠΌΡ Delta-Notch ΡΠΈΠ³Π½Π°Π»ΠΈΠ·Π°ΡΠΈΠΈ
- 5. 1. ΠΠ·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ Π»ΠΈΠ³Π°Π½Π΄Π° ΠΈ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠ°
- 5. 2. ΠΠ΅ΡΠ΅Π΄Π°ΡΠ° Delta-Notch ΡΠΈΠ³Π½Π°Π»Π° Π² ΠΊΠ»Π΅ΡΠΊΡ
- 5. 3. ΠΠ΅ΡΠ΅Π΄Π°ΡΠ° Delta-Notch ΡΠΈΠ³Π½Π°Π»Π° ΠΈΠ· ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΡ Π² ΡΠ΄ΡΠΎ
- 6. Π Π΅Π³ΡΠ»ΡΡΠΈΡ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ Delta-Notch ΡΠΈΠ³Π½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΏΡΡΠΈ
- 6. 1. Π Π΅Π³ΡΠ»ΡΡΠΈΡ ΡΠ½Π°ΡΡΠΆΠΈ ΠΊΠ»Π΅ΡΠΊΠΈ
- 6. 2. ΠΠΎΡΡΡΡΠ°Π½ΡΠ»ΡΡΠΈΠΎΠ½Π½ΡΠ΅ ΠΌΠΎΠ΄ΠΈΡΠΈΠΊΠ°ΡΠΈΠΈ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠ° ΠΈ Π»ΠΈΠ³Π°Π½Π΄Π°
- 6. 3. Π¦ΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠ°ΠΊΡΠΎΡΡ, Π²Π»ΠΉΡΡΡΠΈΠ΅ Π½Π° Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ Delta-Notch ΡΠΈΠ³Π½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡΠ°
- 6. 4. Π―Π΄Π΅ΡΠ½ΡΠ΅ ΡΡΡΠ΅ΠΊΡΠΎΡΡ
- 7. Π€ΠΈΠ»ΠΎΠ³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· DSL Π»ΠΈΠ³Π°Π½Π΄ΠΎΠ²
- 8. ΠΠ½Π°ΡΠ΅Π½ΠΈΠ΅ Π³ΡΡΠΏΠΏΡ Drosophila virilis Π΄Π»Ρ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎΠ³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ
- ΠΠΠ’ΠΠ ΠΠΠΠ« Π ΠΠΠ’ΠΠΠ«
- Π ΠΠΠ£ΠΠ¬Π’ΠΠ’Π« Π ΠΠΠ‘Π£ΠΠΠΠΠΠ
- 1. ΠΠ»ΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ Π³Π΅Π½Π° Delta Drosophila virilis
- 2. Π¦ΠΈΡΠΎΠ³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· Π»ΠΎΠΊΡΡΠ° Dl D. virilis
- 3. ΠΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ Π³Π΅Π½Π° DI Π½Π° ΠΏΠΎΠ»ΠΈΡΠ΅Π½Π½ΡΡ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ°Ρ D. virilis
- 4. Π‘ΡΠ°Π²Π½Π΅Π½ΠΈΠ΅ Π½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄Π½ΡΡ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠ΅ΠΉ Π³Π΅Π½Π° DI Ρ D. virilis ΠΈ D. melanogaster
- 5. Π‘ΡΠ°Π²Π½Π΅Π½ΠΈΠ΅ Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡΠ½ΡΡ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠ΅ΠΉ Π³Π΅Π½Π° DI Ρ D. virilis ΠΈ D. melanogaster
- 6. ΠΠΎΠ½ΡΠ΅ΡΠ²Π°ΡΠΈΠ²Π½ΡΠ΅ ΠΈ Π΄ΠΈΠ²Π΅ΡΠ³ΠΈΡΠΎΠ²Π°Π½Π½ΡΠ΅ Π΄ΠΎΠΌΠ΅Π½Ρ Π½Π΅ΠΊΠΎΡΠΎΡΡΡ Π±Π΅Π»ΠΊΠΎΠ², Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ½ΡΡ Π±Π΅Π»ΠΊΡ Dl D. melanogaster
- 7. ΠΠΎΠΈΡΠΊ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»Π°Π³Π°Π΅ΠΌΡΡ ΡΠ°ΠΉΡΠΎΠ² ΠΌΠΎΠ΄ΠΈΡΠΈΡΠΈΡΠΎΠ²Π°Π½ΠΈΡ Π±Π΅Π»ΠΊΠ° Delta D. virilis
- 8. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΈΠΎΠ½Π½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ Π»ΠΎΠΊΡΡΠ° Delta D. virilis
- 9. ΠΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ ΠΌΠ ΠΠ Π³Π΅Π½Π° Delta D. virilis Π² ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Ρ Π½Π° ΡΠ°Π·Π½ΡΡ ΡΡΠ°Π΄ΠΈΡΡ ΡΠ°Π·Π²ΠΈΡΠΈΡ
- 10. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ?)Π΅//Π°-ΠΌΡΡΠ°Π½ΡΠ½ΡΡ Π»ΠΈΠ½ΠΈΠΉ D. virilis
- ΠΠ«ΠΠΠΠ«
ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· Π»ΠΎΠΊΡΡΠ° Delta Ρ Drosophila virilis (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
Π Π°Π·Π²ΠΈΡΠΈΠ΅ ΡΠ»ΠΎΠΆΠ½ΠΎΠ³ΠΎ ΠΌΠ½ΠΎΠ³ΠΎΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠ° ΠΈΠ· ΠΎΠ΄Π½ΠΎΠΉ ΠΊΠ»Π΅ΡΠΊΠΈ — ΡΠ΅Π·ΡΠ»ΡΡΠ°Ρ ΠΊΠΎΠΎΡΠ΄ΠΈΠ½ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠ³ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ ΠΌΠ½ΠΎΠ³ΠΈΡ ΡΠΈΡΡΠ΅ΠΌ Π³Π΅Π½ΠΎΠ². Π¦Π΅Π½ΡΡΠ°Π»ΡΠ½ΡΠΌ Π²ΠΎΠΏΡΠΎΡΠΎΠΌ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΡΠ²Π»ΡΠ΅ΡΡΡ Π²ΡΡΡΠ½Π΅Π½ΠΈΠ΅ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ° Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Π² ΠΎΠ½ΡΠΎΠ³Π΅Π½Π΅Π·Π΅ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ Π²Π½ΡΡΡΠ΅Π½Π½ΠΈΡ ΠΈ Π²Π½Π΅ΡΠ½ΠΈΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ², ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΡΡΠΈΡ ΡΡΠ΄ΡΠ±Ρ ΠΊΠ»Π΅ΡΠΎΠΊ. ΠΠ°ΠΆΠ΄Π°Ρ ΠΊΠ»Π΅ΡΠΊΠ° Π½Π΅ΠΎΠ΄Π½ΠΎΠΊΡΠ°ΡΠ½ΠΎ ΡΠ΅ΡΠ°Π΅Ρ Π·Π°Π΄Π°ΡΡ Π²ΡΠ±ΠΎΡΠ° Π΄Π°Π»ΡΠ½Π΅ΠΉΡΠ΅Π³ΠΎ ΠΏΡΡΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ Π½Π° ΡΠ°Π·Π½ΡΡ ΡΡΠ°ΠΏΠ°Ρ ΠΆΠΈΠ·Π½ΠΈ, ΡΠ°ΠΊΠΈΡ ΠΊΠ°ΠΊ ΠΏΡΠΎΠ»ΠΈΡΠ΅ΡΠ°ΡΠΈΡ, ΠΌΠΈΠ³ΡΠ°ΡΠΈΡ, ΡΠΎΡΡ, Π΄ΠΈΡΡΠ΅ΡΠ΅Π½ΡΠΈΡΠΎΠ²ΠΊΠ° ΠΈ ΠΊΠ»Π΅ΡΠΎΡΠ½Π°Ρ ΡΠΌΠ΅ΡΡΡ. Π Π°Π·Π»ΠΈΡΠ½ΡΠ΅ Π°Π²ΡΠΎΠ½ΠΎΠΌΠ½ΡΠ΅ ΠΈ Π½Π΅Π°Π²ΡΠΎΠ½ΠΎΠΌΠ½ΡΠ΅ ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΠ΅ ΡΠ°ΠΊΡΠΎΡΡ, ΠΌΠ΅ΠΆΠΊΠ»Π΅ΡΠΎΡΠ½ΡΠ΅ ΡΠΈΠ³Π½Π°Π»Ρ Ρ ΡΠ°Π·Π½ΡΠΌ Π΄ΠΈΠ°ΠΏΠ°Π·ΠΎΠ½ΠΎΠΌ Π΄Π΅ΠΉΡΡΠ²ΠΈΡ Π½Π°ΠΏΡΠ°Π²Π»ΡΡΡ ΡΠ°Π·Π²ΠΈΡΠΈΠ΅ ΠΊΠ»Π΅ΡΠΊΠΈ Π² ΡΠΎΠΌ ΠΈΠ»ΠΈ ΠΈΠ½ΠΎΠΌ Π½Π°ΠΏΡΠ°Π²Π»Π΅Π½ΠΈΠΈ. Π§Π°ΡΡΠΎ Π΄Π»Ρ Π΄ΠΎΡΡΠΈΠΆΠ΅Π½ΠΈΡ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΡ ΡΠ΅Π»Π΅ΠΉ Π² ΡΠ°Π·Π½ΡΡ Π°ΡΠΏΠ΅ΠΊΡΠ°Ρ ΠΆΠΈΠ·Π½ΠΈ ΠΊΠ»Π΅ΡΠΊΠΈ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌ ΠΈΡΠΏΠΎΠ»ΡΠ·ΡΠ΅Ρ ΠΎΠ΄Π½Ρ ΠΈ ΡΡ ΠΆΠ΅ ΡΠΈΠ³Π½Π°Π»ΡΠ½ΡΡ ΡΠΈΡΡΠ΅ΠΌΡ, ΠΈ Π½Π°ΠΎΠ±ΠΎΡΠΎΡ, ΠΎΠ΄Π½Π° ΠΏΡΠΎΠ³ΡΠ°ΠΌΠΌΠ° ΡΠ°Π·Π²ΠΈΡΠΈΡ ΠΌΠΎΠΆΠ΅Ρ ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°ΡΡΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ΠΌ ΡΠ°Π·Π½ΡΡ ΡΠΈΠ³Π½Π°Π»ΡΠ½ΡΡ ΠΏΡΡΠ΅ΠΉ.
Π¦Π΅Π½ΡΡΠ°Π»ΡΠ½ΡΡ ΡΠΎΠ»Ρ Π² ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠΈ ΡΡΠ΄ΡΠ±Ρ ΠΊΠ»Π΅ΡΠΎΠΊ Π² Π»ΠΎΠΊΠ°Π»ΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡΡ ΠΈΠ³ΡΠ°Π΅Ρ Delta-Notch-ΡΠΌΡΠ½Π°Π»ΡΠΏΠ°Ρ ΡΠΈΡΡΠ΅ΠΌΠ° (ΠΈΠ»ΠΈ ΡΠΈΠ³Π½Π°Π»ΡΠ½ΡΠΉ ΠΏΡΡΡ). Π’Π°ΠΊ, ΠΌΠ½ΠΎΠ³ΠΎΡΠΈΡΠ»Π΅Π½Π½ΡΠ΅ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΏΠΎ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π½Π΅ΠΉΡΠΎΠ³Π΅Π½Π΅Π·Π° Ρ Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ ΠΏΠΎΠΊΠ°Π·Π°Π»ΠΈ, ΡΡΠΎ ΠΏΡΠΎΠ΄ΡΠΊΡΡ Π½Π΅ΠΉΡΠΎΠ³Π΅Π½Π½ΡΡ (Notch, Delta, Suppressor of Hairless ΠΈ Π΄Ρ.) ΠΈ ΠΏΡΠΎΠ½Π΅ΠΉΡΠ°Π»ΡΠ½ΡΡ (achaete-scute ΠΊΠΎΠΌΠΏΠ»Π΅ΠΊΡ ΠΈ Π΄Ρ.) Π³Π΅Π½ΠΎΠ², ΡΠ²Π»ΡΡΡΠΈΠ΅ΡΡ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΠ°ΠΌΠΈ Delta-Notch-ΡΠΈΠ³Π½Π°Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ, ΠΎΡΡΡΠ΅ΡΡΠ²Π»ΡΡΡ ΠΈΠ½Π΄ΡΠΊΡΠΈΡ ΠΈ ΠΏΠ΅ΡΠ΅Π΄Π°ΡΡ ΡΠΈΠ³Π½Π°Π»Π° Π»Π°ΡΠ΅ΡΠ°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΈΠ½Π³ΠΈΠ±ΠΈΡΠΎΠ²Π°Π½ΠΈΡ (Muskavitch et al., 1994; Artavanis-Tsakonas et al., 1995). Π ΠΏΡΠΎΡΠ΅ΡΡΠ΅ Π»Π°ΡΠ΅ΡΠ°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΈΠ½Π³ΠΈΠ±ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΏΡΠ΅Π΄ΡΠ΅ΡΡΠ²Π΅Π½Π½ΠΈΠΊ Π½Π΅ΡΠ²Π½ΠΎΠΉ ΠΊΠ»Π΅ΡΠΊΠΈ ΠΏΠΎΠ΄Π°Π²Π»ΡΠ΅Ρ Π½Π΅ΠΉΡΠ°Π»ΡΠ½ΡΠΉ ΠΏΠΎΡΠ΅Π½ΡΠΈΠ°Π» ΡΠΎΡΠ΅Π΄Π½ΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ, ΠΊΠΎΡΠΎΡΡΠ΅ Π² ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠ΅ Π΄ΠΈΡΡΠ΅ΡΠ΅Π½ΡΠΈΡΡΡΡΡΡ Π² ΡΠΏΠΈΠ΄Π΅ΡΠΌΠΎΠ±Π»Π°ΡΡΡ (ΠΏΡΠ΅Π΄ΡΠ΅ΡΡΠ²Π΅Π½Π½ΠΈΠΊΠΈ ΡΠΏΠΈΠ΄Π΅ΡΠΌΠ°Π»ΡΠ½ΡΡ ΠΊΠ»Π΅ΡΠΎΠΊ). ΠΠΏΠΎΡΠ»Π΅Π΄ΡΡΠ²ΠΈΠΈ Π²ΡΡΡΠ½ΠΈΠ»ΠΎΡΡ, ΡΡΠΎ Delta-Notch-ΡΠΈΠ³Π½Π°Π»ΡΠ½Π°Ρ ΡΠΈΡΡΠ΅ΠΌΠ° ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΠ΅Ρ Π²ΡΠ±ΠΎΡ ΠΊΠ»Π΅ΡΠΊΠ°ΠΌΠΈ ΠΈΡ ΠΏΡΡΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ Π½Π΅ ΡΠΎΠ»ΡΠΊΠΎ Π² Π½Π΅ΠΉΡΠΎΠ³Π΅Π½Π΅Π·Π΅, Π½ΠΎ ΠΈ Π²ΠΎ ΠΌΠ½ΠΎΠ³ΠΈΡ Π΄ΡΡΠ³ΠΈΡ ΠΏΡΠΎΡΠ΅ΡΡΠ°Ρ Π½Π° ΡΠ°Π·Π½ΡΡ ΡΡΠ°ΠΏΠ°Ρ ΡΠΌΠ±ΡΠΈΠΎΠ³Π΅Π½Π΅Π·Π° ΠΈ ΠΌΠ΅ΡΠ°ΠΌΠΎΡΡΠΎΠ·Π° Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ (Artavanis-Tsakonas et al., 1999).
Π ΠΏΠΎΡΠ»Π΅Π΄Π½ΠΈΠ΅ Π³ΠΎΠ΄Ρ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ, ΠΊΠ°ΡΠ°ΡΡΠΈΠ΅ΡΡ ^//Ρ-Π^Ρ/Π³-ΡΠΈΠ³Π½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΏΡΡΠΈ, Π²ΡΠ·ΡΠ²Π°ΡΡ ΠΎΠ³ΡΠΎΠΌΠ½ΡΠΉ ΠΈΠ½ΡΠ΅ΡΠ΅Ρ, ΠΏΠΎΡΠΊΠΎΠ»ΡΠΊΡ ΠΎΡΠ° ΡΠΈΡΡΠ΅ΠΌΠ° ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΠ΅Ρ ΡΡΠ΅Π·Π²ΡΡΠ°ΠΉΠ½ΠΎ ΡΠΈΡΠΎΠΊΠΈΠΉ ΡΠΏΠ΅ΠΊΡΡ ΠΊΠ»Π΅ΡΠΎΡΠ½ΡΡ ΠΏΡΠΎΡΠ΅ΡΡΠΎΠ², ΡΠ°ΠΊΠΈΡ ΠΊΠ°ΠΊ Π΄ΠΈΡΡΠ΅ΡΠ΅Π½ΡΠΈΡΠΎΠ²ΠΊΠ°, ΠΏΡΠΎΠ»ΠΈΡΠ΅ΡΠ°ΡΠΈΡ, Π°ΠΏΠΎΠΏΡΠΎΠ· ΠΈ Π΄Ρ., Ρ ΡΠ°Π·Π½ΡΡ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠ². ΠΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈ Π³Π΅Π½ΠΎΠ² Delta ΠΈ Notch Π½Π°ΠΉΠ΄Π΅Π½Ρ Ρ ΡΠ°Π·Π½ΡΡ Π²ΠΈΠ΄ΠΎΠ² ΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ ΠΈ Π±Π΅ΡΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ : Ρ C. elegans (Tax et al., 1994), Ρ Π»ΡΠ³ΡΡΠ΅ΠΊ Xenopus (Chitnis et al., 1995), ΠΊΡΡ (Henrique et al., 1995,) ΠΌΡΡΠ΅ΠΉ (Dunwoodie et al., 1997) ΠΈ Π΄Ρ., Π° ΡΠ°ΠΊΠΆΠ΅ Ρ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° (Laborda et al, 1993). ΠΠ°Π»ΠΈΡΠΈΠ΅ Π³Π΅Π½ΠΎΠ², Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ½ΡΡ Delta ΠΈ Notch, Ρ ΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ , Π²ΠΊΠ»ΡΡΠ°Ρ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΈ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ°, ΠΏΠΎΠΊΠ°Π·ΡΠ²Π°Π΅Ρ, ΡΡΠΎ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠΈΡ Π³Π΅Π½ΠΎΠ² ΠΈΠΌΠ΅Π΅Ρ ΡΠ½ΠΈΠ²Π΅ΡΡΠ°Π»ΡΠ½ΠΎΠ΅ Π·Π½Π°ΡΠ΅Π½ΠΈΠ΅ Π² ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠΈ Π²ΡΠ΅ΠΎΠ±ΡΠΈΡ Π·Π°ΠΊΠΎΠ½ΠΎΠΌΠ΅ΡΠ½ΠΎΡΡΠ΅ΠΉ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΠ°Π·Π²ΠΈΡΠΈΡ ΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΡΡΠΊΠ°ΡΠΈΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΊΠ»Π΅ΡΠΎΠΊ.
ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π½Π΅ΠΉΡΠΎΠ³Π΅Π½Π½ΡΡ Π³Π΅Π½ΠΎΠ² ΠΌΠΎΠΆΠ΅Ρ ΠΏΠΎΠΌΠΎΡΡ Π² ΡΠ΅ΡΠ΅Π½ΠΈΠΈ ΠΏΡΠ°ΠΊΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ΅Π΄ΠΈΡΠΈΠ½ΡΠΊΠΈΡ ΠΏΡΠΎΠ±Π»Π΅ΠΌ. ΠΠ±Π½Π°ΡΡΠΆΠ΅Π½ΠΎ, ΡΡΠΎ Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ ΠΏΠ°ΡΠΎΠ»ΠΎΠ³ΠΈΠΈ Ρ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° (Π±ΠΎΠ»Π΅Π·Π½Ρ ΠΠ»ΡΡΠ³Π΅ΠΉΠΌΠ΅ΡΠ°, ΡΠΈΠ½Π΄ΡΠΎΠΌ CADASIL, Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ ΡΠ°ΠΊΠΎΠ²ΡΠ΅ ΠΎΠΏΡΡ ΠΎΠ»ΠΈ) ΠΈ Π΄ΡΡΠ³ΠΈΡ ΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ ΡΠ²ΡΠ·Π°Π½Ρ Ρ ΠΌΡΡΠ°ΡΠΈΡΠΌΠΈ Π² Π³Π΅Π½Π°Ρ , Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ½ΡΡ Delta ΠΈ Notch, ΠΈΠ»ΠΈ Ρ Π΄ΡΡΠ³ΠΈΠΌΠΈ Π΄Π΅ΡΠ΅ΠΊΡΠ°ΠΌΠΈ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ Delta-Notch-cnmajihtioro ΠΏΡΡΠΈ (Ellisen et al., 1991; Wong, 1997; Struhl et al., 1999).
Π Π½Π°ΡΡΠΎΡΡΠ΅Π΅ Π²ΡΠ΅ΠΌΡ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π° ΠΎΠ±ΡΠ°Ρ ΡΡ Π΅ΠΌΠ° Delta-Notch ΡΠΈΠ³Π½Π°Π»ΠΈΠ·Π°ΡΠΈΠΈ, Π²ΡΡΠ²Π»Π΅Π½Ρ ΠΎΡΠ½ΠΎΠ²Π½ΡΠ΅ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΡ ΡΡΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ, Π° ΡΠ°ΠΊΠΆΠ΅ ΠΏΠΎΡΡΠΎΡΠ½Π½ΠΎ ΠΏΠΎΠΏΠΎΠ»Π½ΡΡΡΠΈΠΉΡΡ Π½Π°Π±ΠΎΡ Π²Π½Π΅ΡΠ½ΠΈΡ ΠΈ Π²Π½ΡΡΡΠΈΠΊΠ»Π΅ΡΠΎΡΠ½ΡΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ², ΡΠ΅Π³ΡΠ»ΠΈΡΡΡΡΠΈΡ Π΅Π΅ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅. Π’Π΅ΠΌ Π½Π΅ ΠΌΠ΅Π½Π΅Π΅, ΠΎΡΡΠ°Π΅ΡΡΡ Π΅ΡΠ΅ ΠΌΠ½ΠΎΠ³ΠΎ Π²ΠΎΠΏΡΠΎΡΠΎΠ², ΠΊΠ°ΡΠ°ΡΡΠΈΡ ΡΡ ΡΡΡΡΠΊΡΡΡΡ ΠΈ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Π²ΡΠ΅Ρ ΡΡΠΈΡ ΠΊΠΎΠΌΠΏΠΎΠ½Π΅Π½ΡΠΎΠ². Π ΡΠ°ΡΡΠ½ΠΎΡΡΠΈ, Π½Π΅Π΄ΠΎΡΡΠ°ΡΠΎΡΠ½ΠΎ ΠΏΠΎΠ½ΡΡΠ΅Π½ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Π»ΠΈΠ³Π°Π½Π΄Π° (Delta) ΠΈ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠ° (Notch), Π² ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠ΅ ΠΊΠΎΡΠΎΡΠΎΠ³ΠΎ ΠΏΠΎΡΠ»Π΅Π΄Π½ΠΈΠΉ Π°ΠΊΡΠΈΠ²ΠΈΡΡΠ΅ΡΡΡ ΠΈ ΠΏΠ΅ΡΠ΅Π΄Π°Π΅Ρ ΡΠΈΠ³Π½Π°Π» Π²Π½ΡΡΡΡ ΠΊΠ»Π΅ΡΠΊΠΈ. ΠΠ΅ΠΊΠΎΡΠΎΡΡΠ΅ Π΄Π΅ΡΠ°Π»ΠΈ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Π²Π½ΡΡΡΠΈΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠΉ ΡΠ°ΡΡΠΈ Π±Π΅Π»ΠΊΠ° Notch, ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈ ΡΠ΄Π΅ΡΠ½ΡΡ Π±Π΅Π»ΠΊΠΎΠ² — ΡΡΠ°ΡΡΠ½ΠΈΠΊΠΎΠ² Delta-Notch ΡΠΈΠ³Π½Π°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΡΠ°ΠΊΠΆΠ΅ Π½Π΅ ΡΡΠ½Ρ. ΠΡΠ΅Π½Ρ ΠΌΠ°Π»ΠΎ ΠΈΠ·Π²Π΅ΡΡΠ½ΠΎ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ ΠΏΡΠΎΠΈΡΡ ΠΎΠ΄ΠΈΡ Ρ Π»ΠΈΠ³Π°Π½Π΄ΠΎΠΌ Π΄ΠΎ ΠΈ ΠΏΠΎΡΠ»Π΅ Π΅Π³ΠΎ Π½Π΅ΠΏΠΎΡΡΠ΅Π΄ΡΡΠ²Π΅Π½Π½ΠΎΠ³ΠΎ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠΎΠΌ Π½Π° ΠΏΠΎΠ²Π΅ΡΡ Π½ΠΎΡΡΠΈ ΠΊΠ»Π΅ΡΠΊΠΈ.
ΠΠΎΠΊΠ° Π½ΠΈΡΠ΅Π³ΠΎ Π½Π΅ ΠΈΠ·Π²Π΅ΡΡΠ½ΠΎ ΡΠ°ΠΊΠΆΠ΅ ΠΎ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ°Ρ ΡΠ°Π±ΠΎΡΡ Π²Π½ΡΡΡΠ΅Π½Π½Π΅ΠΉ, Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ Π΄ΠΈΠ²Π΅ΡΠ³ΠΈΡΠΎΠ²Π°Π½Π½ΠΎΠΉ, ΡΠ°ΡΡΠΈ Π±Π΅Π»ΠΊΠ° Delta, Ρ ΠΎΡΡ Π΅ΡΡΡ Π΄Π°Π½Π½ΡΠ΅ ΠΎ ΡΠΎΠΌ, ΡΡΠΎ ΡΡΠΎΡ ΡΠΈΡΠΎΠΏΠ»Π°Π·ΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ Π΄ΠΎΠΌΠ΅Π½ Π²Π°ΠΆΠ΅Π½ Π΄Π»Ρ ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ Delta-ΠΏΠΎΠ΄ΠΎΠ±Π½ΡΡ Π»ΠΈΠ³Π°Π½Π΄ΠΎΠ² (Chitnis et al., 1995; Henderson et. al., 1997; Lissemore and Starmer, 1999).
ΠΠ»Ρ ΡΠ΅ΡΠ΅Π½ΠΈΡ Π²ΠΎΠΏΡΠΎΡΠΎΠ² ΡΡΠ½ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ Π³Π΅Π½ΠΎΠ² ΠΈ ΠΈΡ ΠΏΡΠΎΠ΄ΡΠΊΡΠΎΠ² Π²Π΅ΡΡΠΌΠ° ΠΏΠΎΠ»Π΅Π·Π½ΠΎ ΡΡΠ°Π²Π½Π΅Π½ΠΈΠ΅ ΡΡΡΡΠΊΡΡΡΡ ΡΡΠΈΡ Π³Π΅Π½ΠΎΠ², Π° ΡΠ°ΠΊΠΆΠ΅ ΠΎΡΠΎΠ±Π΅Π½Π½ΠΎΡΡΠ΅ΠΉ ΠΈΡ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ ΠΈ ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Ρ ΡΠ²ΠΎΠ»ΡΡΠΈΠΎΠ½Π½ΠΎ ΠΎΡΠ΄Π°Π»Π΅Π½Π½ΡΡ Π²ΠΈΠ΄ΠΎΠ².
ΠΠΎΡΡΠΎΠΌΡ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΠ»ΠΎΡΡ ΠΈΠ½ΡΠ΅ΡΠ΅ΡΠ½ΡΠΌ ΡΡΠ°Π²Π½ΠΈΡΡ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΡ ΠΈ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΡ Π³Π΅Π½Π° Delta Ρ Π΄Π²ΡΡ Π²ΠΈΠ΄ΠΎΠ² Π΄ΡΠΎΠ·ΠΎΡΠΈΠ»Ρ — D. melanogaster ΠΈ D. virilis, Π΄ΠΈΠ²Π΅ΡΠ³ΠΈΡΠΎΠ²Π°Π²ΡΠΈΡ ΠΎΠΊΠΎΠ»ΠΎ 60 ΠΌΠ»Π½. Π»Π΅Ρ Π½Π°Π·Π°Π΄ ΠΈ ΠΎΡΠ½ΠΎΡΡΡΠΈΡ ΡΡ ΠΊ ΡΠ°Π·Π½ΡΠΌ ΠΏΠΎΠ΄ΡΠΎΠ΄Π°ΠΌ ΡΠΎΠ΄Π° Drosophila (Beverly et al., 1984). ΠΠ²Π΅Π½Π°Π΄ΡΠ°ΡΡ Π²ΠΈΠ΄ΠΎΠ² Π³ΡΡΠΏΠΏΡ virilis ΠΈΠ·Π΄Π°Π²Π½Π° ΡΠΈΡΠΎΠΊΠΎ ΠΈΡΠΏΠΎΠ»ΡΠ·ΡΡΡΡΡ ΠΊΠ°ΠΊ ΡΠ΄ΠΎΠ±Π½ΡΠΉ ΡΠΊΡΠΏΠ΅ΡΠΈΠΌΠ΅Π½ΡΠ°Π»ΡΠ½ΡΠΉ ΠΎΠ±ΡΠ΅ΠΊΡ Π² ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ ΡΠ²ΠΎΠ»ΡΡΠΈΠΎΠ½Π½ΡΡ , ΡΠΈΡΠΎΠ³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ , ΠΏΠΎΠΏΡΠ»ΡΡΠΈΠΎΠ½Π½ΡΡ ΠΈ Π±ΠΈΠΎΡ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡΡ . ΠΠΈΠ΄ Drosophila virilis Π½Π° ΠΎΡΠ½ΠΎΠ²Π°Π½ΠΈΠΈ Π°Π½Π°Π»ΠΈΠ·Π° Ρ ΡΠΎΠΌΠΎΡΠΎΠΌ ΠΈ ΡΡΠ΄Π° Π΄ΡΡΠ³ΠΈΡ ΠΌΠΎΡΡΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΡ ΠΏΡΠΈΠ·Π½Π°ΠΊΠΎΠ² ΡΠ°ΡΡΠΌΠ°ΡΡΠΈΠ²Π°Π΅ΡΡΡ ΠΊΠ°ΠΊ Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ ΠΏΡΠΈΠΌΠΈΡΠΈΠ²Π½Π°Ρ, ΠΏΠΎ-Π²ΠΈΠ΄ΠΈΠΌΠΎΠΌΡ, ΠΏΡΠ΅Π΄ΠΊΠΎΠ²Π°Ρ ΡΠΎΡΠΌΠ° Π²ΡΠ΅ΠΉ Π³ΡΡΠΏΠΏΡ virilis (Throckmorton, 1982).
ΠΡΠ° Π³ΡΡΠΏΠΏΠ° ΠΏΡΠΈΠ²Π»Π΅ΠΊΠ»Π° Π½Π°ΡΠ΅ Π²Π½ΠΈΠΌΠ°Π½ΠΈΠ΅ Π΅ΡΠ΅ ΠΈ ΠΏΠΎΡΠΎΠΌΡ, ΡΡΠΎ Π² Π½Π°ΡΠ΅ΠΌ ΡΠ°ΡΠΏΠΎΡΡΠΆΠ΅Π½ΠΈΠΈ Π½Π°Ρ ΠΎΠ΄ΠΈΠ»Π°ΡΡ ΡΠ½ΠΈΠΊΠ°Π»ΡΠ½Π°Ρ ΠΊΠΎΠ»Π»Π΅ΠΊΡΠΈΡ ΠΌΡΡΠ°ΡΠΈΠΉ Π² Π»ΠΎΠΊΡΡΠ΅ Delta, ΠΊΠΎΡΠΎΡΡΠΉ Ρ Drosophila virilis ΡΠ²Π»ΡΠ΅ΡΡΡ Π³ΠΎΡΡΡΠ΅ΠΉ ΡΠΎΡΠΊΠΎΠΉ (hot spot) Π΄Π»Ρ ΡΡΠ°Π½ΡΠΏΠΎΠ·ΠΈΡΠΈΠΈ ΠΌΠΎΠ±ΠΈΠ»ΡΠ½ΡΡ ΡΠ»Π΅ΠΌΠ΅Π½ΡΠΎΠ² (Evgen'ev et al., 1997). ΠΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠΎΠΉ ΠΊΠΎΠ»Π»Π΅ΠΊΡΠΈΠΈ ΠΎΡΠΊΡΡΠ²Π°Π΅Ρ ΡΠΈΡΠΎΠΊΠΈΠ΅ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡΠΈ Π΄Π»Ρ Π΄Π°Π»ΡΠ½Π΅ΠΉΡΠ΅Π³ΠΎ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²Π° Π½Π΅ΠΉΡΠΎΠ³Π΅Π½ΠΎΠ².
ΠΠΠΠΠ ΠΠΠ’ΠΠ ΠΠ’Π£Π Π«.
Π²ΡΠ²ΠΎΠ΄Ρ.
1. ΠΠ»ΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ ΠΈ ΠΊΠ°ΡΡΠΈΡΠΎΠ²Π°Π½ ΡΡΠ°ΡΡΠΎΠΊ ΠΠΠ ΡΠ°Π·ΠΌΠ΅ΡΠΎΠΌ 14 Ρ.ΠΏ.Π½., ΡΠΎΠ΄Π΅ΡΠΆΠ°ΡΠΈΠΉ 3' ΠΎΠ±Π»Π°ΡΡΡ Π»ΠΎΠΊΡΡΠ° Delta D.virilis.
2. ΠΠ΅ΡΠΎΠ΄ΠΎΠΌ in situ Π³ΠΈΠ±ΡΠΈΠ΄ΠΈΠ·Π°ΡΠΈΠΈ Π³Π΅Π½ Delta Π±ΡΠ» Π»ΠΎΠΊΠ°Π»ΠΈΠ·ΠΎΠ²Π°Π½ Π² ΡΠ°ΠΉΠΎΠ½Π΅ 21 °F Π²ΠΎ 2 Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ D.virilis. ΠΡΠΎΠ²Π΅Π΄Π΅Π½Π½Π°Ρ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ ΠΏΠΎΠ΄ΡΠ²Π΅ΡΠ΄ΠΈΠ»Π° ΠΈ ΡΡΠΎΡΠ½ΠΈΠ»Π° ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ ΡΠΈΡΠΎΠ³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π° Π»ΠΎΠΊΡΡΠ° Delta D. virilis, Π²ΡΠΏΠΎΠ»Π½Π΅Π½Π½ΠΎΠ³ΠΎ Ρ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ½ΡΡ ΠΏΠ΅ΡΠ΅ΡΡΡΠΎΠ΅ΠΊ.
3. Π‘Π΅ΠΊΠ²Π΅Π½ΠΈΡΠΎΠ²Π°Π½ ΡΡΠ°Π³ΠΌΠ΅Π½Ρ Π»ΠΎΠΊΡΡΠ° Delta D. virilis, Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ½ΡΠΉ ΠΏΠΎΡΠ»Π΅Π΄Π½Π΅ΠΌΡ (ΡΠ΅ΡΡΠΎΠΌΡ) ΡΠΊΠ·ΠΎΠ½Ρ Π»ΠΎΠΊΡΡΠ° Dl D.melanogaster.
4. ΠΠΎΠΌΠΏΡΡΡΠ΅ΡΠ½ΡΠΉ Π°Π½Π°Π»ΠΈΠ· ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΠΠ ΠΈΡΡΠ»Π΅Π΄ΡΠ΅ΠΌΠΎΠ³ΠΎ ΡΡΠ°Π³ΠΌΠ΅Π½ΡΠ° ΠΏΠΎΠΊΠ°Π·Π°Π», ΡΡΠΎ ΠΎΠ½ ΠΊΠΎΠ΄ΠΈΡΡΠ΅Ρ ΠΏΡΠΈΠΌΠ΅ΡΠ½ΠΎ ΠΏΠΎΠ»ΠΎΠ²ΠΈΠ½Ρ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»Π°Π³Π°Π΅ΠΌΠΎΠ³ΠΎ Π±Π΅Π»ΠΊΠ° Delta D. virilis, Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ½ΠΎΠ³ΠΎ Π½Π° 83% Π±Π΅Π»ΠΊΡ Delta D. melanogaster, ΠΏΡΠΈΡΠ΅ΠΌ ΠΈΡΡΠ»Π΅Π΄ΡΠ΅ΠΌΠ°Ρ ΡΠ°ΡΡΡ Π±Π΅Π»ΠΊΠ° Dl D. virilis Π½Π° 35 Π°ΠΌΠΈΠ½ΠΎΠΊΠΈΡΠ»ΠΎΡ Π΄Π»ΠΈΠ½Π½Π΅Π΅, ΡΠ΅ΠΌ Ρ Π±Π΅Π»ΠΊΠ° D1 D.melanogaster.
5. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Ρ ΡΠ°Π·ΠΌΠ΅ΡΡ ΠΈ ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²ΠΎ ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΎΠ² Π»ΠΎΠΊΡΡΠ° Delta D.virilis. Π£ΡΡΠ°Π½ΠΎΠ²Π»Π΅Π½ΠΎ, ΡΡΠΎ Π»ΠΎΠΊΡΡ Delta D. virilis ΠΊΠΎΠ΄ΠΈΡΡΠ΅Ρ Π΄Π²Π° Π·ΠΈΠ³ΠΎΡΠΈΡΠ΅ΡΠΊΠΈΡ (5,7 ΠΈ 4,8 Ρ.Π½.) ΠΈ Π΄Π²Π° ΠΌΠ°ΡΠ΅ΡΠΈΠ½ΡΠΊΠΈΡ (3,9 ΠΈ 2,8 Ρ.Π½.) ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠ°, Π² ΠΎΡΠ»ΠΈΡΠΈΠ΅ ΠΎΡ ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²ΡΡΡΠ΅Π³ΠΎ Π»ΠΎΠΊΡΡΠ° D. melanogaster, Π³Π΄Π΅ ΠΈΠ· ΡΡΠ΅Ρ ΠΌΠ°ΠΆΠΎΡΠ½ΡΡ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΠΎΠ² ΠΎΠ΄ΠΈΠ½ (5.4 Ρ.Π½.) ΡΡΠΈΡΠ°Π΅ΡΡΡ Π·ΠΈΠ³ΠΎΡΠΈΡΠ΅ΡΠΊΠΈΠΌ, Π° Π΄Π²Π° Π΄ΡΡΠ³ΠΈΡ , ΡΠ°Π·ΠΌΠ΅ΡΠΎΠΌ 4.6 ΠΈ 3.6 Ρ.Π½., ΠΌΠ°ΡΠ΅ΡΠΈΠ½ΡΠΊΠΈΠΌΠΈ.
6. ΠΠ½Π°Π»ΠΈΠ· ΡΠ°ΡΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΡ ΠΌΠ ΠΠ Delta Π² ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Ρ ΠΌΠ΅ΡΠΎΠ΄ΠΎΠΌ Π½Π΅ΡΠ°Π΄ΠΈΠΎΠ°ΠΊΡΠΈΠ²Π½ΠΎΠΉ Π³ΠΈΠ±ΡΠΈΠ΄ΠΈΠ·Π°ΡΠΈΠΈ ΠΏΠΎΠΊΠ°Π·Π°Π», ΡΡΠΎ ΡΡΠ°Π½ΡΠΊΡΠΈΠΏΡΡ Π³Π΅Π½Π° Delta D. virilis Π»ΠΎΠΊΠ°Π»ΠΈΠ·ΡΡΡΡΡ Π² ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Ρ Π² ΡΠ΅Ρ ΠΆΠ΅ ΠΎΡΠ³Π°Π½Π°Ρ ΠΈ ΡΠΊΠ°Π½ΡΡ ΠΈ Π½Π° ΡΠ΅Ρ ΠΆΠ΅ ΡΡΠ°Π΄ΠΈΡΡ ΡΠ°Π·Π²ΠΈΡΠΈΡ, ΡΡΠΎ ΠΈ Ρ D.melanogaster.
Π‘ΠΏΠΈΡΠΎΠΊ Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΡ
- ΠΠ»Π±Π΅ΡΡΡ Π., ΠΡΠ΅ΠΉ Π., ΠΡΡΠΈΡ ΠΠΆ., Π Π΅ΡΡ Π., Π ΠΎΠ±Π΅ΡΡΠ΅ Π., Π£ΠΎΡΡΠΎΠ½ ΠΠΆ., ΠΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½Π°Ρ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΡ ΠΊΠ»Π΅ΡΠΊΠΈ. // Π.: ΠΠΈΡ, 1994−1996, Ρ. 1 -3.
- ΠΡΠ±Π΅Π½ΠΊΠΎ Π.Π‘., ΠΠ°ΡΠΈΡΠ΅Π²Π° Π. Π. ΠΠΎΠ΄ΠΈΡΠΈΠΊΠ°ΡΠΈΡ ΡΠΊΡΠΏΡΠ΅ΡΡΠΈΠΈ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ° Delta Ρ Drosophila virilis. ΠΡΡΠ΅ΠΊΡ ΠΏΠΎΠ»ΠΎΠΆΠ΅Π½ΠΈΡ ΠΏΡΠΈ ΡΡΠ°Π½ΡΠ»ΠΎΠΊΠ°ΡΠΈΡΡ , Π΄Π΅ΡΠΈΡΠ΅Π½ΡΠΈ ΠΈ Π΄ΡΠΏΠ»ΠΈΠΊΠ°ΡΠΈΡΡ . // ΠΠ΅Π½Π΅ΡΠΈΠΊΠ°. 1982. Π’.28. № 3. Π‘.441−453.
- ΠΡΠ±Π΅Π½ΠΊΠΎ Π.Π‘., Π‘ΡΠ±Π±ΠΎΡΠ° Π . Π. ΠΡΠΎΠ±Π΅Π½Π½ΠΎΡΡΠΈ ΠΏΠΎΡΠΎΠΌΡΡΠ²Π° ΠΎΠ±Π»ΡΡΠ΅Π½Π½ΡΡ ΡΠ°ΠΌΡΠΎΠ² ΠΈΠ· Π΄Π²ΡΡ Π»ΠΈΠ½ΠΈΠΉ Drosophila virilis, ΡΠ°Π·Π»ΠΈΡΠ°ΡΡΠΈΡ ΡΡ ΠΏΠΎ ΡΡΠΎΠ²Π½Ρ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π½Π΅ΡΡΠ°Π±ΠΈΠ»ΡΠ½ΠΎΡΡΠΈ. //ΠΠΎΠΊΠ»Π°Π΄Ρ ΠΠΊΠ°Π΄Π΅ΠΌΠΈΠΈ ΠΠ°ΡΠΊ, 1990. Ρ. 312. № 6. Ρ. 1501−1504.
- ΠΡΠ±Π΅Π½ΠΊΠΎ Π.Π‘., Π ΡΠ±ΡΠΎΠ²Π° Π. Π., ΠΠ΅Π»Π΅Π½ΡΠΎΠ²Π° Π. Π‘., ΠΠ΅Π·ΠΈΠ½ Π. Π’., ΠΠΎΡΠΎΡΠΊΠΈΠ½ Π. Π., ΠΠ²Π³Π΅Π½ΡΠ΅Π² Π. Π. ΠΠΎΠΊΡΡ Delta Ρ Drosophila virilis: ΠΊΠ»ΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΈ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ½Π°Ρ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΡ. //ΠΠΎΠΊΠ»Π°Π΄Ρ ΠΠΊΠ°Π΄Π΅ΠΌΠΈΠΈ ΠΠ°ΡΠΊ, 1998. Ρ.358, № 4 Ρ.567−569.
- ΠΠΎΡΠΎΡΠΊΠΈΠ½ Π.Π. ΠΠ²Π΅Π΄Π΅Π½ΠΈΠ΅ Π² Π³Π΅Π½Π΅ΡΠΈΠΊΡ ΡΠ°Π·Π²ΠΈΡΠΈΡ. //ΠΠΎΡΠΊΠ²Π°. ΠΠ°ΡΠΊΠ°,. 1999. 235 Π‘.
- Ahmad, I., Zagouras, P., and Artavanis-Tsakonas S. Involvement of Notch-1 in mammalian retinal neurogenesis-association of Notch-1 activity with both immature and terminally differentiated cells.// Mech. Development, 1995. V. 53, P .73−85.
- Alexander M.L. The Genetics and Biology of Drosophila. /(M.Ashburner, E. Novitski Eds.).// Academic Press. New York, 1976. V.lc. P.1365−1427.
- Artavanis-Tsakonas S., Rand M., Lake R. Notch signaling: Cell fate control and signal integration in development. // Science, 1999. V. 284. P. 770−776.
- Artavanis-Tsakonas S., Matsuno K., Fortini M.E. Notch signalling.// Science, 1995. V.268. P. 225−232.
- Ashburner M. Drosophila. A laboratory handbook. //Cold Spring Harbor Laboratory Press. Cold Spring Harbor, 1989. NY. 256 P.
- Axelrod JD, Matsuno K, Artavanis-Tsakonas S, Perrimon N. Interaction between Wingless and Notch signaling pathways mediated by disheveled.// Science. 1996, V.271 P. 1826−1832.
- Bang AG, Bailey AM, Posakony JW. Hairless promotes stable commitment to the sensory organ precursor cell fate by negatively regulating the activity of the Notch signaling pathway.// Dev Biol., 1995. V.172 P. 479−494.
- Bettenhausen Π., Hrabe de Angelis, M., Simon, D., Guenet, J.L. and Gossler, A. Transient and restricted expression during mouse embryogenesis of Dill, a murine gene closely related to Drosophila Delta. //Development, 1995, V.121, P.2407−2418.
- Beverley S.M. Wilson A.C. Molecular evalution in Drosophila and the higher Diptera II. A time scale for fly evolution. //J. Mol. Evol., 1984. V. 21. P. l-13.
- Bier E. Anti-neural-inhibition: a conserved mechanism for neural induction.// Cell. Review, 1997 V. 89 P.681−684.
- Bishop SA, Klein T, Arias AM, Couso JP. Composite signalling from Serrate and Delta establishes leg segments in Drosophila through Notch.// Development, 1999 Jul, V.126, P.2993−3003.
- Blanchet-Tournier M.F., Tricoire H., Busson D., Lamour-Isnard C. The segment-polarity gene fused is highly conserved in DrosophilaJ/GensA995.v. 19.p. 157−162.
- Blaumueller C. M, Qi H.L., Zagouras P., and Artavanis-Tsakonas S. Intercellular cleavage of Notch leads to a heterodimeric receptor on the plasma membane.// Cell, 1997. V.90, P.281−291.
- Bray S. Notch affair. //Cell, 1998. V. 93. P. 499−503.
- Brennan K, Tateson R, Lewis K, Arias AM. A functional analysis of Notch mutations in Drosophila. //Genetics, 1997, SeP. V.147 P.177−188.
- Burris P.A., Zhang Y., Rusconi J.C., Corbin V. The pore-forming and cytoplasmic domains of the neurogenic gene product, BIG BRAIN, are conserved between Drosophila virilis and Drosophila melanogaster. Gene. 1998. v. 206. p. 69−76.
- Cabrera C.V. Lateral inhibition and cell fate during neurogenesis in Drosophila: the interactions between Scute, Notch and Delta.// Development, 1990. V.109, P.733−742.
- Campos-Ortega J.A. Genetic mechanisms of early neurogenesis in Drosophila melanogaster. //Mol Neurobiol. 1995, V.10 № 2 P. 75−89.
- Campos-Ortega J.A. Celluar interactions during early neurogenesis of Drosophila melanogaster. IITINS, V. ll, No.9,1988, P.245−150.
- Campos-Ortega J.A. Genetics of neurogenesis in Drosophila melanogaster.//TINS, V.8, No.6,1985, P.245−250.
- Campos-Ortega, J. A. and Hartenstein, V. The embryonic development of Drosophila melanogaste.//. Springer-Verlag: Berlin, 1985.
- Campuzano S, Modolell J. Patterning of the Drosophila nervous system: the Achaete-scute gene complex. Trends //Genet., 1992 Jun.V.8 P. 202−208. Review.
- Casey P. J. Protein lipidation in cell signalling.// Science. 1995. V.268. P. 221−225.
- Doherty, D., Feager G., Younger Sheperd S Jan L.Y. and Yan Y.N. Delta is a ventral dorsal signal complementary to Serrate, another Notch ligand, in Drosophila wing formation.// Genes and Dev., 1996. V. 10 P. 421−434.
- Dominguez M, de Celis JF. A dorsal/ventral boundary established by Notch controls growth and polarity in the Drosophila eye. //Nature, 1998, Nov 19, V.396 P.276−278.
- Donoviel, D.B., Hadjantonakis A., Ikeda M., Zheng H., Hyslop P., Bernstein A. Mice lacking both presenilin genes exhibit early embrionic patterning defects.//Genes and Development, 1999. V.13. P. 2801−2810.
- Dorsky RI, Chang WS, Rapaport DH, Harris WA. Regulation of neuronal diversity in the Xenopus retina by Delta signalling. //Nature, 1997, V.385 P.67−70.
- Dunwoodie S.L., Henrique D, Harrison S.M., Beddington R.S. Mouse D113: a novel divergent Delta gene which may complement the function of other Delta homologues during early pattern formation in the mouse embryo. //Development, 1997. V.124 P.3065−3076.
- Ellisen L.W., Bird J., West D.C., Soreng A.L., Reynolds T.C., Smith S.D. and Sklar J. TAN-l, the human homolog of the Drosophila Notch gene, is broken by chromosomal translocations in T lymphoblastic neoplasms// Cell, 1991 V.66 № 4, P.649−661.
- Fanto M, Mlodzik M. Asymmetric Notch activation specifies photoreceptors R3 and R4 and planar polarity in the Drosophila eye. //Nature. 1999 V.397 P.523−6.
- Fehon RG, Johansen K, Rebay I, Artavanis-Tsakonas S. Complex cellular and subcellular regulation of Notch expression during embryonic and imaginal development of Drosophila: implications for Notch function.//J Cell Biol., 1991. V.113 P.657−669.
- Fehon, R.G., Kooh, P.J., Rebay, I., Regan, C.L., Xu, Π’., Muskavitch, M.A. and
- Artavanis-Tsakonas, S. Molecular interactions between the protein products of the neurogenic loci Notch and Delta, two EGF-homologous genes in Drosophila. //Cell, 1990. Π£. 61, P.523−534.
- Fleming RJ, Gu Y, Hukriede NA. Serrate-mediated activation of Notch is specifically blocked by the product of the gene fringe in the dorsal compartment of the Drosophila wing imaginal disc.//Development, 1997 № 15 P.2973−81.
- Fleming RJ, Scottgale TN, Diederich RJ, Artavanis-Tsakonas S. The gene Serrate encodes a putative EGF-like transmembrane protein essential for proper ectodermal development in Drosophila melanogaster J'/Genes Dev. 1990 № 12A P.2188−2201.
- Fortini, M.E. and Artavanis-Tsakonas, S. The suppressor of Hairless protein participates in Notch receptor signaling. //Cell, 1994. V.79 P.273−282
- Fortini, M.E. and Artavanis-Tsakonas, S. Notch-. Neurogenesis is only part of the picture. //Cell, 1993. 75 P. 1245−1247.
- Fortini ME, Rebay I, Caron LA, Artavanis-Tsakonas S. An activated Notch receptor blocks cell-fate commitment in the developing Drosophila eye. //Nature, 1993 V.365 № 6446 P.555−557.
- Fransson L. Structure and function of cell associated proteoglican.// Trends Biochem. Sci., 1987, V.12p 406−411.
- Gho M, Lecourtois M, Geraud G, Posakony JW, Schweisguth F. Subcellular localization of Suppressor of Hairless in Drosophila sense organ cells during Notch signalling. //Development. 1996, Y.122 P.1673−1682.
- Ghysen A, Dambly-Chaudiere C. Genesis of the Drosophila peripheral nervous system. //Trends Genet., 1989 V.5 P.251−5. Review.
- Go, M. J. and Artavanis-Tsakonas, S. A genetic screen for novel components of the Notch signaling pathway during Drosophila bristle development. //Genetics, 1998. V.150. № 1 P.211−220.
- Greenwald I. LN-HNotch signaling: lessons from worms and flies.//Genes Dev. 1998i1. N. 12, Π .1751−1762.
- Gu, Y., Hukriede, N. A. and Fleming, R. J. Serrate expression can functionally replace Delta activity during neuroblast segregation in the Drosophila embryo.// Development, 1995. V.121: 855−865.
- Gubenko, I. S, Evgen’ev M.B. Cytological and linkage maps of Drosophila virilis chromosomes. //Genetica. 1984. 65, P. 127−139.
- Gubenko I.S., Subbota R.P., Semeshin V.F. Unusual Drosophila virilis stress-puff at 20CD: cytological localization of a heat sensitive locus and some peculiarities of the heat shock response. //Hereditas. 1991. V. 115. P. 283−290.
- Hackel P.O., Zwick E., Prenzel N., Ullrich A. Epidermal growth factor receptors: critical mediators of multiple receptor pathways. //Curr. Opin. Cell Biol. 1999, V. 11. P. 184−189.
- Haenlin M, Kunisch M, Kramatschek B, Campos-Ortega J. A .Genomic regions regulating early embryonic expression of the Drosophila neurogenic gene Delta.// Mech Dev., 1994, V. P.199−110
- Haenlin M., Kramatschek Π., Campos-Ortega J.A. The pattern of transcription of the neurogenic gene Delta, of Drosophila melanogaster. Development. 1990. No. 110, P.905−914.
- Hardy J. and Israel A. In search of y-secretase.// Nature. 1999. V. 398. P. 466−467.
- Hartenstein V, Younossi-Hartenstein A, Lekven A. Delamination and division in the Drosophila neurectoderm: spatiotemporal pattern, cytoskeletal dynamics, and common control by neurogenic and segment polarity genes. //Dev Biol., 1994, V. l65 P.480−499.
- Hartenstein V. Atlas of of Drosophila Development. //Cold Spring Harbor Laboratory1. Press, 1993. 57 P.
- Hartenstein, A.Y., Rugendorff, A., Tepass, U. and Hartenstein, V. The function of the neurogenic genes during epithelial development in the Drosophila embryo.// Development, 1992. V. l 16 P.1203−1220
- Heitzler P, Simpson P. Altered epidermal growth factor -like sequences provide evidence for a role of Notch as a receptor in cellfate decisions.// Development. 1993. V. 117. P. 1113−1123.
- Heitzler P, Simpson P. The choice of cell fate in the epidermis of Drosophila./ICe 11., 1991 Mar 22. V.64 P.1083−1092.
- Heitzler P, Bourouis M, Ruel L, Carteret C, Simpson P. Genes of the Enhancer of split and achaete-scute complexes are required for a regulatory loop between Notch and Delta during lateral signalling in Drosophila. //Development, 1996 V 122 P. l61 -171.
- Henderson S., Gao D. Christensen S., E., Kimble J. Functional domains of lag-2, a putative signaling ligand for LIN-12 and GLP-1 receptors in Caenorhabditis elegands. //Mol.Biol.Cell, 1997, V.8 P.1751−1762.
- Henrique D, Hirsinger E, Adam J, Le Roux I, Pourquie O, Ish-Horowicz D, Lewis J. Maintenance of neuroepithelial progenitor cells by Delta-Notch signalling in the embryonic chick retina.// Curr Biol, 1997. V.7 P.661−670.
- Hing H.K., Sun X., Artavanis-Tsakonas S. //Mech. Dev. 1994. V. 47. P. 261.
- Hinz, U., Giebel, B. and Campos-Ortega, J.A. The basic-helix-loop-helix domain of Drosophila lethal of scute protein is sufficient for proneural function and activates neurogenic genes. //Cell, 1994. V.76 P. 77−87
- Hirschberg C. and Snider M., Topography of glycosilation in the rough endoplasmic reticulum and Golgi apparatus.//Annu. Rev. Biochem. 1987. V.56. P.63−88.
- Hukriede NA, Gu Y, Fleming RJ. A dominant-negative form of Serrate acts as a general antagonist of Notch activation. //Development. 1997. V. 124 P.3427−37.
- Huppert SS, Jacobsen TL, Muskavitch MA. Feedback regulation is central to Delta
- Notch signalling required for Drosophila wing vein morphogenesis. //Development. 1997. V.124P.3283−3291.
- Jacobsen T.L., Brennan K, Arias A.M., Muskavitch M.A. Cis-interactions between Delta and Notch modulate neurogenic signalling in Drosophila J7 Development, 1998 Nov. V.125 P.4531−4540.
- Jehn BM, Bielke W, Pear WS, Osborne BA. Cutting edge: protective effects of Notch-1 on TCR-induced apoptosis.// J Immunol. 1999.V. 162 P.635−8.
- Jen W. C, Wettstein D., Turner D., Chitnis A., Kintner C. the Notch ligand, X-Delta-2, mediates segmentation of the paraxial mesoderm in Xenopus embryos.// Development. 1997. V. 124. P. 1169−1178.
- Johnston S.H., Rauskolb C, Wilson R, Prabhakaran B, Irvine K.D., Vogt T.F. A family of mammalian Fringe genes implicated in boundary determination and the Notch pathway. Development. 1997 Jun, V.124 P.2245−2254.
- Jonsson, F. and Knust, E. Distinct functions of the Drosophila genes Serrate and Delta revealed by ectopic expression during wing development.// Dev. Genes Evol., 1996. 206 P. 91−101
- Joutel A., Corpechot C., Ducros A. NotchZ mutations in CADASIL, a hereditary adult-onset condition causing stroke and dementia.// Nature. 1996. V. 383. P. 707−710.
- Kerszberg M., Changeux J.P. A simple molecular model of neurulation. //Bioessays, 1998 SeP.V.20 P.758−770.
- Kidd S., Kelley M. R. and Young M. W. Sequence of the Notch locus of Drosophila melanogaster: relationship of the encoded protein to mammalian clotting and growth factors.// Mol. Cell. Biol. 1986. V.6 P. 3094−3108.
- Kidd S., Lieber Π’., Young M.W. Ligand-iduced cleavage and regulation of nuclear enry of Notch in Drosophila melanogaster embryos. //Genes Dev. 1998.12 P. 3728−3740.
- Kim J., Irvine K. D., Carroll S. B. Cell recognition, signal induction, and symmetrical gene activation at the dorsal-ventral boundary of the developing Drosophila wing.
- Cell, 1995.V.82 P.795−802.
- Kimble J, Simpson P. The LIN-12/Notch signaling pathway and its regulation.//Annu Rev Cell Dev Biol. 1997- N.13 P.333−361.
- Klambt C, Jacobs JR, Goodman CS. The midline of the Drosophila central nervous system: a model for the genetic analysis of cell fate, cell migration, and growth cone guidance. //Cell, 1991 Feb 22. V.64 P.801−815.
- Klein, T. and Arias, A. M. Interactions among Delta, Serrate and Fringe modulate Notch activity during Drosophila wing development. //Development, 1998. V.125 P. 2951−2962.
- Klueg, Π. M., Parody, T. R. and Muskavitch, M. A. T. Complex proteolytic processing acts on Delta, a transmembrane ligand for Notch, during Drosophila development. //Mol. Biol. Cell, 1998. V.9 P.1709−1723.
- Knust E., Schorans H., Grawe F., Campos-Ortega J. A. Seven genes of Enhancer of split complex of Drosophila melanogaster encode helix-loop-helix proteins.// Genetics. 1992. V. 132. P. 505−518.
- Kopan R., Schroeter E.H., Weintraub H., Nye J.S.Signal transduction by activated mNotch: impotance of proteolytic processing and its regulation by the extracellular domain. //Proc. Natl. Acad. Sci. USA. 1996. V. 93. P. 1683−1688.
- Kopan R., Turner D.L. The Notch pathway: democracy and aristocracy in the selection of cell fate. //Curr. Opin. Neurobiol. 1996. V.6 P. 594−601.
- Kopczynski C.C., Muskavitch M.A.T. Complex spatio-temporal accumulation of alternative transcripts from the neurogenic gene Delta during Drosophila embriogenesis.// Development, 107,1989, P. 623−636.
- Kunisch M, Haenlin M, Campos-Ortega JA Lateral inhibition mediated by the Drosophila neurogenic gene Delta is enhanced by proneural proteins.// Proc Natl Acad Sci USA, 1994, V.11,N.91 P.10 139−10 143.
- Laborda J., Sausville E.A., Hoffman Π’., Notario V. dlk, a putative mammalian homeotic gene differentially expressed in small cell lung carcinoma and neuroendocrine tumor cell line.// J. Biol. Chem, 1993. V.268 P. 3817−3820.
- Lardelli, M., Dahlstrand, J. and Lendahl, U. The novel Notch homologue mouse Notch 3 lacks specific epidermal growth factor-repeats and is expressed in proliferating neuroepithelium //Mech. Dev. 1994. V.46, P. 123−136.
- Lawrence P. The making of a fly. The genetics of animal design. //Oxford, Blackwell, 1992.
- Lecourtois M, Schweisguth F Indirect evidence for Delta-dependent intracellular processing of Notch in Drosophila embryos. Curr Biol 1998 V.13 P.771−774
- Lehmann R., Dietrich U., Jimenez F., Campos-Ortega J. A. et al. Mutations of early neurogenesis in Drosophila. Roux Arch. dev. Biol., 1981. V. l90 P. 226−229.
- Lehmann R., Jimenez F., Dietrich U., Campos-Ortega J.A. On the phenotype and development of mutants of early neurogenesis in Drosophila melanogaster. Roux’s Arch. Dev. Biol. 192,1983, P. 62−74
- Lecourtois M, Schweisguth F. The neurogenic suppressor of hairless DNA-binding protein mediates the transcriptional activation of the enhancer of split complex genes triggered by Notch signaling. Genes Dev. 1995-V.9 P.2598−2608.
- Lendahl U. A growing family ofNotch ligands. BioEssays, 1998, v. 20. P. 103−107.
- Lim, J.K. In situ hybridization with biotinylated DNA. 11 Dros. Inf. Serv. 1993, V.72,73−77.
- Lindsley D.L., Zimm G. Genetic variations of Drosophila melanogaster. II Carnegie Inst. Wash. Publ., 1968. P.23−29.
- Lindsell CE, Shawber CJ, Boulter J, Weinmaster G. Jagged: a mammalian ligand that activates Notchl. // Cell, 1995 Mar 24 V.80 P.909−917.
- Lindsell CE, Boulter J, diSibio G, Gossler A, Weinmaster G. Expression patterns of Jagged, Deltal, Notchl, Notch2, and Notch3 genes identify ligand-receptor pairs that may function in neural development. Mol Cell Neurosci., 1996. V. 8 P. 14−27.
- Lissemore JL, Starmer WT Phylogenetic analysis of vertebrate and invertebrate Delta/Serrate/LAG-2 (DSL) proteins. Mol Phylogenet Evol 1999 V.2 P.308−319
- Long AD, Lyman RF, Langley CH, Mackay TF. Two sites in the Delta gene region contribute to naturally occurring variation in bristle number in Drosophila melanogaster. II Genetics, 1998. V.149 P. 999−1017.
- Lukowitz W., Schroder C., Glaser G., Hulskamp M., Tautz D. Regulatory and coding regions of the segmentation gene hunchback are functionally concerved between Drosophila virilis ΠΈ Drosophila melanogaster. II Mech. Dev. 1994. v. 45. p. 105−115.
- Maine E.M., Lissemore J.L., Starmer W.T. A Phylogenetic analysis of vertebrate and invertebrate Notch-related genes. //Mol Phylogenet Evol. 1995. V. 4. P. 139−149.
- Martin-Bermudo, M. D., Carmena A. and Jimenez, F. Neurogenic genes control gene expression at the transcriptional level in early neurogenesis and in mesectoderm specification. //Development, 1995. 121 P. 219−224
- Matsuno K., Go M.J., Sun X., Eastman S., Artavanis-Tsakonas S. Suppressor of Hairless-independent events in Notch signaling imply novel pathway elements. // Development, 1997. V.124. P. 4265−4273.
- Michael M.W., Bowtell D., Rubin J.M. Comparison of the sevenless genes of Drosophila virilis ΠΈ Drosophila melanogaster. //Proc. Natl. Acad. Sci. USA. 1990. v.87. p. 5351−5353.
- Modolell J., Campuzano S. The achaete-scute complex as an integrating device. Int. II J. Dev. Biol. 1998,42, P.275−282.
- Moghal N., Sternberg P.W. Multiple positive and negative regulators of signaling by the EGF-receptor. //Curr. Opin. Cell Biol. 1999, V. 11. P. 190−196.
- Murrel, J., Farlow M., Ghetti Π., Benson M. A mutation in the amyloid precursor protein assosiated with hereditary Alzheimer’s disease.// Science. 1991. V.254. P.97−99.
- Muskavitch M.A.T. Delta-Notch Signaling and Drosophila II Cell Fate Choice. (Review) Develop.Biol. No.166,1994, P. 415−430.
- Newfeld S.J., Smoller D.A., Yedvobnick B. Interspecific comparison of the unusually repetitive Drosophila locus mastermind. //J.Mol.Evol. 1991.V.32. p.415−420.
- Oellers N, Dehio M, Knust E. bHLH proteins encoded by the Enhancer of split complex of Drosophila negatively interfere with transcriptional activation mediated by proneural genes. //Mol Gen Genet, 1994 Sep 1. V.244 P.465−473.
- Pan D., Valentine S.A., Courey A.J. The bipartite Drosophila melanogaster twist promoter is reorganized in Drosophila virilis. // Mech Dev. 1994. v. 46. p. 41−53.
- Pan, D. and Rubin, G. M. Kuzbanian controls proteolytic processing of Notch and mediates lateral inhibition during Drosophila and vertebrate neurogenesis. //Cell, 1997 V.90P. 271−280.
- Panin VM, Papayannopoulos V, Wilson R, Irvine KD. Fringe modulates Notch-ligand interactions. //Nature, 1997 Jun 26. V. 387 P.908−912.
- Parks, A. L., Klueg Π. M., Stout J. R. and Muskavitch, M. A. Ligand endocytosis drives receptor dissotiation and activation in the Notch pathway. //Development. 2000. V. 127. P. 1373−1385.
- Parks, A. L., Huppert, S. S. and Muskavitch, M. A. The dynamics of neurogenic signalling underlying bristle development in Drosophila melanogaster. //Mech. Dev., 1997. V. 63 P. 61−74
- Parody, Π’. and Muskavitch, M.A. The pleiotropic function of Delta during postembryonic development of Drosophila melanogaster. //Genetics, 1993.V.135 P.527−539.
- Paulson D.F. Chromosomal deficiensis and embrionic development of Drosophila melanogaster. //Proc. Natl. Acad. Sci., V.23, 1937, P.133−137.
- Pear WS, Aster JC, Scott ML, Hasserjian RP, Soffer B, Sklar J, Baltimore D. Exclusive development of T cell neoplasms in mice transplanted with bone marrow expressing activated Notch alleles.// J Exp Med., 1996 -183 P.2283−2291.
- Qi H, Rand MD, Wu X, Sestan N, Wang W, Rakic P, Xu T, Artavanis-Tsakonas S. Processing of the Notch ligand Delta by the metalloprotease Kuzbanian.// Science, 1999. V.283 P.91−94.
- Rauskolb C, Correia T, Irvine K.D. Fringe-dependent separation of dorsal and ventral cells in the Drosophila wing. //Nature. 1999 V.401 P.476−480.
- Rebay I., Fleming R.J., Fehon R.G., Cherbas L., Cherbas P., Artavanis-Tsakonas S. Specific EGF repeats of Notch mediate interactions with Delta and Serrate: implications for Notch as a multifunctional receptor. /Cell. 1991. V. 67. P. 687−699.
- Robey E. Notch in vertebrates. //Curr. Opin. Genet. Dev. 1997. V. 7. P. 551 -557.
- Rooke J., Xu Π’., Positive and negative signals between interacting cells for establishing neural fate. //BioEssays 1998, V.20 P.209−214.
- Rooke J, Pan D, Xu T, Rubin GM. KUZ, a conserved metalloprotease-disintegrin protein with two roles in Drosophila neurogenesis.// Science. 1996 V.273 P. 1227−1231.
- Sambrook J., Fritsch E.F., Maniatis T. Molecular cloning. A laboratory manual. N.Y.: Cold Spring Harbor Lab. Press, 1989.
- Sanger F., Nicklen S. Coulson A.K. DNA sequencing with chain-termination ingibitors// Proc. Nat.Acad.Sci. USA., 1977 V.74.P.5463−5467.
- Seugnet L., Simpson P., Haenlin M. Transcriptional regulation of Notch and Delta: requiment for neuroblast segregation in Drosophila. Development. 1997. v. 124 P. 2015−2025.
- Shelly LL, Fuchs C, Miele L. Notch-1 inhibits apoptosis in murine erythroleukemia cells and is necessary for differentiation induced by hybrid polar compounds. J Cell Biochem, 1999. V.73 P.164−175.
- Simpson P., Carteret C. 1990a. Proneural clusters: equivalence groups in the epithelium of Drosophila. //Development 110, P.927−932.
- Simpson P. Lateral inhibition and the development of the sensory bristles of the adult peripheral nervous system of Drosophila. il Development, 1990b.V.l09 P.509−519.
- Siren M. and Portin P. Interaction of Hairless, Delta, Enhancer of split and Notch genes of Drosophila melanogaster as expressed in adult morphology. //Genet. Res., 1989. V. 53 P. 23−26.
- Skeath JB, Carroll SB. The achaete-scute complex: generation of cellular pattern and fate within the Drosophila nervous system.// FASEB J., 1994 Jul. V.8 P.714−721. Review.
- Sun, X. and Artavanis-Tsakonas, S. Secreted forms of Delta and Serrate define antagonists of Notch signaling in Drosophila. //Development, 1997. V. l24 P. 34 393 448.
- Sun, X., and Artavanis-Tsakonas, S., The intercellular deletions of Delta and Serrate define dominant negative forms of the Drosophila Notch ligands. //Development, 1996, V.122, p2465−2474.
- Struhl G, Greenwald I. Presenilin is required for activity and nuclear access of Notch in Drosophila. //Nature, 1999. V.398 P.522−525.
- Struhl G, Adachi A. Nuclear access and action of Notch in vivo. //Cell, 1998.V. 93 P.649−660.
- Struhl G, Basler K. Organizing activity of wingless protein in Drosophila.!1 Cell, 1993. Feb 26.V. 72 P.527−540.
- Struhl G, Fitzgerald K, Greenwald I. Intrinsic activity of the Lin-12 and Notch intracellular domains in vivo. //Cell, 1993.V.74 P.331−345.
- Tautz D., Hulskamp M., Sommer R.J. Whole mount in situ hybridization in Drosophila. In: In Situ Hybridization. A Practical Approach. /Ed. D.G. Wilkinson.1994. P.61−73.
- Tax, F.E., Yeargers, J.J. and Thomas, J.H. Sequence of C. elegans lag-2 reveals a cell signalling domain shared with Delta and Serrate of Drosophila. //Nature, 1994. V.368 P. 150−154
- Technau G.M., Campos-Ortega J.A. Cell autonomy of expression of neurogenic genes of Drosophila melanogaster.// Proc. Natl. Acad. Sci., V.84,1987, P.4500−4504.
- Throckmorton L.H. The virilis species grouP. In The genetics and biology of Drosophila.// Ashburner M., ed., Acad. Press, 1982, V. 36, P. 227−296.
- Thomas, U., Speicher, S. A. and Knust, E. The Drosophila gene Serrate encodes an EGF-like transmembrane protein with a complex expression pattern in embryos and wing discs.// Development, 1991. V. 111 P.749−61.
- Vassin H., Bremer K.A., Knust E., Campos-Ortega J.A. The neurogenic locus Delta, of Drosophila melanogaster, is expressed in neurogenic territories and encodes a putative transmembrane protein with EGF-like repeats.// EMBO J. No.6, 1987, P. 3431−3440.
- Vassin H, Vielmetter J, Campos-Ortega J A. Genetic interactions in early neurogenesis of Drosophila melanogaster. J Neurogenet. 1985 Nov, V.2 P.291−308.
- Vervoort M., Dambly-Chaudiere C., Ghysen A.// Curr. Opin. Neur. 1997. V.7. P. 21−28.
- Wadsworth, S.C., Vincent, W.S. III. and Bilodeau-Wentworth, D. A Drosophila genomic sequence with homology to human epidermal growth factor receptor.// Nature, 1985.V. 314, P. 178−180.
- Welshons W. J., Keppy T. The recombinational analysis of aberrations and the position of the Notch locus on the polytene chromosome of Drosophila. //Molec. gen. Genet., 1981. V. 181 P.319−324.
- Wong, P.C., Zheng H., Chen H., Becher M., Sisinathsinghji D., Trumbauer M., Chen H. Y., Price L., Van der Ploeg L., Sisodia S. Presenilin 1 is required for Notch 1 and Dll 1 expression in the paraxial mesoderm. //Nature. 1997. V. 387. P. 288−292.
- Yochem, J., Weston, K. and Greenwald, I. The Caenorhabditis elegans I in-12 gene encodes a transmembrane protein with overall similarity to Drosophila Notch. //Nature, 1988.V. 335,547−550.
- Zhou L., Boulianne G.L. Comparison of the neuralized genes of Drosophila virilis and D. melanogaster. Genome. 1994. v. 37. p. 840−847.