Π ΠΎΠ»Ρ Π»Π΅ΡΡΡΠΈΡ ΡΡΠ΅ΡΠΎΠΈΠ΄ΠΎΠ² (Π½Π° ΠΏΡΠΈΠΌΠ΅ΡΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°) Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ
Π’Π΅ΠΎΡΠ΅ΡΠΈΡΠ΅ΡΠΊΠ°Ρ ΠΈ ΠΏΡΠ°ΠΊΡΠΈΡΠ΅ΡΠΊΠ°Ρ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΡ. ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ ΡΠΏΠΎΡΠΎΠ±ΡΡΠ²ΡΡΡ Π½Π°ΠΊΠΎΠΏΠ»Π΅Π½ΠΈΡ ΠΈ ΠΈΠ½ΡΠ΅Π³ΡΠ°ΡΠΈΠΈ Π΄Π°Π½Π½ΡΡ ΠΎ ΡΠ΅Π½ΠΎΡΠΈΠΏΠΈΡΠ΅ΡΠΊΠΈΡ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ°Ρ Ρ ΠΈΠ½Π±ΡΠ΅Π΄Π½ΡΡ Π»ΠΈΠ½ΠΈΠΉ ΠΌΡΡΠ΅ΠΉ-ΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π»ΠΎΠΊΡΡΠ°Ρ , ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΡ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΡ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ (Π½Π° ΠΏΡΠΈΠΌΠ΅ΡΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°), Π° ΡΠ°ΠΊΠΆΠ΅ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΡΡ Π°Π³ΡΠ΅ΡΡΠΈΡ, ΡΠ°ΡΡΠΈΡΡΡ ΡΠ²Π΅Π΄Π΅Π½ΠΈΡ ΡΡΡΠ΅ΡΡΠ²ΡΡΡΠΈΡ Π±Π°Π· Π΄Π°Π½Π½ΡΡ . ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π΄Π°Π½Π½ΡΠ΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ ΠΏΠΎΠ΄ΠΎΠΉΡΠΈ Π±Π»ΠΈΠΆΠ΅… 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°Π²Π° 1. ΠΠ±Π·ΠΎΡ Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΡ
- 1. 1. ΠΠ±ΡΠ°Ρ ΠΌΠΎΡΡΠΎΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½Π°Ρ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΡ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π°ΡΠΎΡΠ° ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ
- 1. 1. 1. ΠΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½Π°Ρ Π²ΡΡΡΠΈΠ»ΠΊΠ°
- 1. 1. 2. ΠΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΠ΅ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΡ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ
- 1. 1. 3. Π’ΡΠ°Π½ΡΠ΄ΡΠΊΡΠΈΡ ΠΈ ΠΊΠΎΠ΄ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠΈΠ³Π½Π°Π»Π° Π² ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΡ ΡΠ΅Π½ΡΠΎΡΠ½ΡΡ Π½Π΅ΠΉΡΠΎΠ½Π°Ρ
- 1. 1. 4. ΠΡΠ½ΠΎΠ²Π½Π°Ρ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½Π°Ρ Π»ΡΠΊΠΎΠ²ΠΈΡΠ° (OOJ1)
- 1. 1. 5. ΠΡΠΎΠ΅ΠΊΡΠΈΠΈ ΠΎΡΠ½ΠΎΠ²Π½ΠΎΠΉ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ Π»ΡΠΊΠΎΠ²ΠΈΡΡ
- 1. 1. 6. ΠΠΎΠ΄ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠ³ΠΎ ΡΠΈΠ³Π½Π°Π»Π° Π² ΠΠΠ
- 1. 1. 7. ΠΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΡΠΉ ΠΎΡΠ³Π°Π½
- 1. 1. 8. ΠΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΡΠ΅ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΡ
- 1. 1. 9. ΠΠΎΠΏΠΎΠ»Π½ΠΈΡΠ΅Π»ΡΠ½Π°Ρ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½Π°Ρ Π»ΡΠΊΠΎΠ²ΠΈΡΠ°
- 1. 1. 10. ΠΠΎΠ½Π²Π΅ΡΠ³Π΅Π½ΡΠΈΡ ΠΈ ΡΠΈΠ½Π΅ΡΠ³ΠΈΠ·ΠΌ ΠΠΠ‘ ΠΈ ΠΠΠ‘
- 1. 1. 11. ΠΡΡΠ³ΠΈΠ΅ Ρ Π΅ΠΌΠΎΡΠ΅Π½ΡΠΎΡΠ½ΡΠ΅ ΠΎΠ±ΡΠ°Π·ΠΎΠ²Π°Π½ΠΈΡ Π½ΠΎΡΠΎΠ²ΠΎΠΉ ΠΏΠΎΠ»ΠΎΡΡΠΈ Ρ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ
- 1. 2. Π₯ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠΈΠ³Π½Π°Π»Ρ ΠΏΠΎΠ·Π²ΠΎΠ½ΠΎΡΠ½ΡΡ
- 1. 2. 1. ΠΠΈΠ΄Ρ Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠΈΠ³Π½Π°Π»ΠΎΠ²
- 1. 2. 2. ΠΠ½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½
- 1. 3. Π ΠΎΠ»Ρ Ρ
ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ
ΡΠΈΠ³Π½Π°Π»ΠΎΠ² Π² Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠΌ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠΈ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ
- 1. 3. 1. ΠΠ³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ΅ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠ΅
- 1. 3. 2. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ Π΄Π΅ΡΠ΅ΡΠΌΠΈΠ½Π°Π½ΡΡΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉΠ°Π³ΡΠ΅ΡΡΠΈΠΈ
- 1. 3. 3. Π£ΡΠ°ΡΡΠΈΠ΅ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ-Π°Π³ΡΠ΅ΡΡΠΈΠΈ
- 1. 3. 4. Π₯ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠΈΠ³Π½Π°Π»Ρ, Π²Π»ΠΈΡΡΡΠΈΠ΅ Π½Π° ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΡΡ Π°Π³ΡΠ΅ΡΡΠΈΡ
- 1. 1. ΠΠ±ΡΠ°Ρ ΠΌΠΎΡΡΠΎΡΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½Π°Ρ ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΡ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π°ΡΠΎΡΠ° ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ
- 2. 1. ΠΠ±ΡΠ°Ρ ΡΡ Π΅ΠΌΠ° ΡΠ°Π±ΠΎΡΡ
- 2. 2. ΠΠ΅ΡΠΎΠ΄Ρ ΡΠ΅ΡΡΠΈΡΠΎΠ²Π°Π½ΠΈΡ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ
- 2. 2. 1. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΠΏΠΎΡΠΎΠ³ΠΎΠ² ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ
- 2. 2. 2. ΠΡΠ΅Π½ΠΊΠ° ΡΡΠΎΠ²Π½Ρ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠΈ
- 2. 2. 3. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΠΏΡΠ΅Π΄ΠΏΠΎΡΡΠ΅Π½ΠΈΡ Π·Π°ΠΏΠ°Ρ ΠΎΠ² ΠΌΠΎΡΠΈ
- 2. 2. 4. Π’Π΅ΡΡ «ΠΎΡΠΊΡΡΡΠΎΠ΅ ΠΏΠΎΠ»Π΅»
- 2. 2. 5. Π‘ΡΠ°ΡΠΈΡΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ ΠΎΠ±ΡΠ°Π±ΠΎΡΠΊΠΈ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΡΠ΅ΡΠΊΠΈΡ ΡΠΊΡΠΏΠ΅ΡΠΈΠΌΠ΅Π½ΡΠΎΠ²
- 2. 3. Π₯ΠΈΡΡΡΠ³ΠΈΡΠ΅ΡΠΊΠΎΠ΅ ΡΠ΄Π°Π»Π΅Π½ΠΈΠ΅ Π²ΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ³Π°Π½Π°
- 2. 4. ΠΠ°ΡΠ΅ΡΠΈΠ°Π» ΠΈ ΠΌΠ΅ΡΠΎΠ΄Ρ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ Π»ΠΎΠΊΠ°Π»ΠΈΠ·Π°ΡΠΈΠΈ Π½Π΅ΠΉΡΠΎΠ½Π°Π»ΡΠ½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ Π² ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΠ΅
- 2. 4. 1. ΠΠΊΡΠΏΠΎΠ·ΠΈΡΠΈΠΈ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΈ ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½ΡΠΌ ΠΎΠ΄ΠΎΡΠ°Π½ΡΠ°ΠΌ
- 2. 4. 2. ΠΡΠΈΠ³ΠΎΡΠΎΠ²Π»Π΅Π½ΠΈΠ΅ ΠΈ ΠΎΠΊΡΠ°ΡΠΈΠ²Π°Π½ΠΈΠ΅ ΡΡΠ΅Π·ΠΎΠ²
- 2. 4. 3. ΠΠ½Π°Π»ΠΈΠ· Fos-ΠΈΠΌΠΌΡΠ½ΠΎΡΠ΅Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ
- 2. 5. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΡ Π³ΠΎΡΠΌΠΎΠ½ΠΎΠ² ΠΌΠ΅ΡΠΎΠ΄ΠΎΠΌ ΡΠ²Π΅ΡΠ΄ΠΎΡΠ°Π·Π½ΠΎΠ³ΠΎ ΠΈΠΌΠΌΡΠ½ΠΎΡΠ΅ΡΠΌΠ΅Π½ΡΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π°
- 2. 5. 1. ΠΠΌΠΌΡΠ½ΠΎΡ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΎΠ΅ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΡ ΡΠ΅ΡΡΠΎΡΡΠ΅ΡΠΎΠ½Π° ΠΈ ΠΊΠΎΡΡΠΈΠΊΠΎΡΡΠ΅ΡΠΎΠ½Π° Π² ΠΏΠ»Π°Π·ΠΌΠ΅ ΠΊΡΠΎΠ²ΠΈ ΠΏΠΎΡΠ»Π΅ ΡΠΊΡΠΏΠΎΠ·ΠΈΡΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ
- 2. 5. 2. Π‘ΡΠ°ΡΠΈΡΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ ΠΎΠ±ΡΠ°Π±ΠΎΡΠΊΠΈ
- 2. 6. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΈ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠΈ
- 2. 6. 1. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ Π³ΠΈΠ±ΡΠΈΠ΄ΠΎΠ²
- 2. 6. 2. Π€Π΅Π½ΠΎΡΠΈΠΏΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ
- 2. 5. 3. ΠΠ½Π°Π»ΠΈΠ· ΡΠ΅Π½ΠΎΡΠΈΠΏΠΎΠ²
- 2. 5. 4. ΠΠ΅Π½ΠΎΡΠΈΠΏΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ Ρ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ ΠΠΠ ΠΌΠ°ΡΠΊΠ΅ΡΠΎΠ²
- 2. 5. 5. ΠΠ½Π°Π»ΠΈΠ· Π°ΡΡΠΎΡΠΈΠ°ΡΠΈΠΉ ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ΅Π½ΠΎΡΠΈΠΏΠ°ΠΌΠΈ ΠΈ Π³Π΅Π½ΠΎΡΠΈΠΏΠ°ΠΌΠΈ
- 3. 1. ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· ΠΏΡΠΈΠ·Π½Π°ΠΊΠΎΠ² ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΈ* ΠΏΠ°ΡΠ°ΠΌΠ΅ΡΡΠΎΠ² ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠΈ
- 3. 1. 1. ΠΠ½Π°Π»ΠΈΠ· ΡΠ΅Π½ΠΎΡΠΈΠΏΠΎΠ² ΡΠΎΠ΄ΠΈΡΠ΅Π»ΡΡΠΊΠΈΡ Π»ΠΈΠ½ΠΈΠΉ ΠΈ Π³ΠΈΠ±ΡΠΈΠ΄ΠΎΠ² ΠΏΠ΅ΡΠ²ΠΎΠ³ΠΎ ΠΏΠΎΠΊΠΎΠ»Π΅Π½ΠΈΡ
- 3. 1. 2. ΠΠ½Π°Π»ΠΈΠ· ΡΠ΅Π½ΠΎΡΠΈΠΏΠΎΠ² Π²ΡΠΎΡΠΎΠ³ΠΎ ΠΏΠΎΠΊΠΎΠ»Π΅Π½ΠΈΡ Π³ΠΈΠ±ΡΠΈΠ΄ΠΎΠ²
- 3. 1. 3. ΠΠ½Π°Π»ΠΈΠ· Π°ΡΡΠΎΡΠΈΠ°ΡΠΈΠΉ ΡΠ΅Π½ΠΎΡΠΈΠΏΠΎΠ² ΠΈ Π³Π΅Π½ΠΎΡΠΈΠΏΠΎΠ² Π²ΡΠΎΡΠΎΠ³ΠΎ ΠΏΠΎΠΊΠΎΠ»Π΅Π½ΠΈΡ Π³ΠΈΠ±ΡΠΈΠ΄ΠΎΠ², ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΡ Ρ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ ΠΠΠ ΠΌΠ°ΡΠΊΠ΅ΡΠΎΠ²
- 3. 1. 4. ΠΠ±ΡΡΠΆΠ΄Π΅Π½ΠΈΠ΅
- 3. 2. Π£ΡΠ°ΡΡΠΈΠ΅ Π΄ΠΎΠΏΠΎΠ»Π½ΠΈΡΠ΅Π»ΡΠ½ΠΎΠΉ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π² ΡΠ΅ΡΠ΅ΠΏΡΠΈΠΈ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ
- 3. 2. 1. Π£Π΄Π°Π»Π΅Π½ΠΈΠ΅ Π²ΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ³Π°Π½Π°
- 3. 2. 2. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π½Π΅ΠΉΡΠΎΠ½Π°Π»ΡΠ½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π°ΡΠΈΠΈ Π² ΠΠΠ Π² ΠΎΡΠ²Π΅Ρ Π½Π° ΡΡΠΈΠΌΡΠ»ΡΡΠΈΡ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½ΠΎΠΌ
- 3. 2. 3. ΠΠ±ΡΡΠΆΠ΄Π΅Π½ΠΈΠ΅
- 3. 3. ΠΠ»ΠΈΡΠ½ΠΈΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° Π½Π° Π½Π΅ΠΊΠΎΡΠΎΡΡΠ΅ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΡΠ΅ΡΠΊΠΈΠ΅ ΠΈ Π³ΠΎΡΠΌΠΎΠ½Π°Π»ΡΠ½ΡΠ΅ ΠΏΠΎΠΊΠ°Π·Π°ΡΠ΅Π»ΠΈ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ
- 3. 3. 1. Π’Π΅ΡΡ Π½Π° ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΡΡ Π°Π³ΡΠ΅ΡΡΠΈΡ
- 3. 3. 2. ΠΠ»ΠΈΡΠ½ΠΈΠ΅ Π΄Π»ΠΈΡΠ΅Π»ΡΠ½ΡΡ ΡΠΊΡΠΏΠΎΠ·ΠΈΡΠΈΠΉ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ Π½Π° ΡΡΠ΄ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΡΠ΅ΡΠΊΠΈΡ ΠΏΠΎΠΊΠ°Π·Π°ΡΠ΅Π»Π΅ΠΉ
- 3. 3. 3. ΠΠ»ΠΈΡΠ½ΠΈΠ΅ ΡΠΊΡΠΏΠΎΠ·ΠΈΡΠΈΠΉ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° Π½Π° ΡΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ ΠΎΡΠ½ΠΎΠ²Π½ΡΡ ΡΡΠ΅ΡΠΎΠΈΠ΄Π½ΡΡ Π³ΠΎΡΠΌΠΎΠ½ΠΎΠ² Π² ΠΏΠ»Π°Π·ΠΌΠ΅ ΠΊΡΠΎΠ²ΠΈ
- 3. 3. 4. ΠΠ±ΡΡΠΆΠ΄Π΅Π½ΠΈΠ΅
Π ΠΎΠ»Ρ Π»Π΅ΡΡΡΠΈΡ ΡΡΠ΅ΡΠΎΠΈΠ΄ΠΎΠ² (Π½Π° ΠΏΡΠΈΠΌΠ΅ΡΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°) Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
ΠΠΊΡΡΠ°Π»ΡΠ½ΠΎΡΡΡ ΠΏΡΠΎΠ±Π»Π΅ΠΌΡ. ΠΠ³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ΅ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠ΅ ΠΈΠ³ΡΠ°Π΅Ρ Π²Π°ΠΆΠ½ΡΡ ΡΠΎΠ»Ρ Π² ΠΏΡΠΎΡΠ΅ΡΡΠ°Ρ Π°Π΄Π°ΠΏΡΠ°ΡΠΈΠΈ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ ΠΊ ΠΎΠΊΡΡΠΆΠ°ΡΡΠ΅ΠΉ ΡΡΠ΅Π΄Π΅. ΠΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²Π°Ρ Π°Π³ΡΠ΅ΡΡΠΈΡ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ Π²ΠΎ ΠΌΠ½ΠΎΠ³ΠΎΠΌ ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΠ΅Ρ Ρ Π°ΡΠ°ΠΊΡΠ΅Ρ ΠΎΡΠ½ΠΎΡΠ΅Π½ΠΈΠΉ Π΄ΠΎΠΌΠΈΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΡ-ΠΏΠΎΠ΄ΡΠΈΠ½Π΅Π½ΠΈΡ, ΠΎΡΠ³Π°Π½ΠΈΠ·Π°ΡΠΈΡ ΠΏΠΎΠΏΡΠ»ΡΡΠΈΠΎΠ½Π½ΠΎΠΉ ΡΡΡΡΠΊΡΡΡΡ ΠΈ Ρ Π°ΡΠ°ΠΊΡΠ΅Ρ ΠΈΠ·Π±ΠΈΡΠ°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΡΠΊΡΠ΅ΡΠΈΠ²Π°Π½ΠΈΡ, ΠΊΠ°ΠΊ Π² Π΅ΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΡΡ ΠΌΠ΅ΡΡΠΎΠΎΠ±ΠΈΡΠ°Π½ΠΈΡΡ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ, ΡΠ°ΠΊ ΠΈ Π² Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠ½ΡΡ ΠΏΠΎΠΏΡΠ»ΡΡΠΈΡΡ (Poole, Morgan, 1973, ΠΠΎΠ²ΠΈΠΊΠΎΠ², 1988). Π‘Ρ ΠΎΠ΄ΡΡΠ²ΠΎ ΡΡΠ΄Π° ΡΠ»Π΅ΠΌΠ΅Π½ΡΠΎΠ² Π°Π³ΠΎΠ½ΠΈΡΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° ΠΈ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΠ΅Ρ ΠΎΠ± ΠΈΡ ΠΎΠ±ΡΠΈΡ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ ΠΎΡΠ½ΠΎΠ²Π°Ρ .
Π£ΡΠ°ΡΡΠΈΠ΅ Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠΈΠ³Π½Π°Π»ΠΎΠ² Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ ΡΠΈΠΏΠΎΠ². ΡΠΎΡΠΈΠ°Π»ΡΠ½ΠΎΠ³ΠΎ, Π² ΡΠΎΠΌ ΡΠΈΡΠ»Π΅ ΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ, Π±ΡΠ»ΠΎ ΠΏΡΠΎΠ΄Π΅ΠΌΠΎΠ½ΡΡΡΠΈΡΠΎΠ²Π°Π½ΠΎ Π΄Π»Ρ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ Π²ΠΈΠ΄ΠΎΠ² ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ (Halpern, Martinez-Marcos, 2003, Brennan, Kendrick, 2006). Π ΡΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎΠΌ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»Π΅Π½ΠΈΠΈ, Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠΈΠ³Π½Π°Π»Ρ ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ , ΠΊ ΠΊΠΎΡΠΎΡΡΠΌ ΠΎΡΠ½ΠΎΡΡΡ ΠΈ ΡΠ΅ΡΠΎΠΌΠΎΠ½Ρ, ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΡΡ ΡΠΎΠ±ΠΎΠΉ ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΠ΅ Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠ΅ Π²Π΅ΡΠ΅ΡΡΠ²Π° ΠΈΠ»ΠΈ ΡΠΌΠ΅ΡΠΈ, Π²ΡΠ΄Π΅Π»ΡΠ΅ΠΌΡΠ΅ ΠΎΠ΄Π½ΠΎΠΉ ΠΎΡΠΎΠ±ΡΡ ΠΈ ΠΎΠΊΠ°Π·ΡΠ²Π°ΡΡΠΈΠ΅ Π²Π»ΠΈΡΠ½ΠΈΠ΅ Π½Π° Π΄ΡΡΠ³ΡΡ ΠΎΡΠΎΠ±Ρ ΡΠΎΠ³ΠΎ ΠΆΠ΅ Π²ΠΈΠ΄Π°, Ρ. Π΅. ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΠ΅ Π² ΠΊΠΎΠΌΠΌΡΠ½ΠΈΠΊΠ°ΡΠΈΠΈ (Johnston, 2000). ΠΠΎΡΠ° Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ ΡΠ²Π»ΡΠ΅ΡΡΡ Π΅ΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΡΠΌ ΠΈΡΡΠΎΡΠ½ΠΈΠΊΠΎΠΌ Ρ Π΅ΠΌΠΎΡΠΈΠ³Π½Π°Π»ΠΎΠ², ΠΎΠΊΠ°Π·ΡΠ²Π°ΡΡΠΈΡ Π²Π»ΠΈΡΠ½ΠΈΠ΅ Π½Π° Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ΅ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠ΅, ΠΏΡΠΈΡΠ΅ΠΌ ΡΠ΅ΡΠΎΠΌΠΎΠ½Π°Π»ΡΠ½ΡΠΉΡΠΈΠ³Π½Π°Π» ΠΈΠΌΠ΅Π΅Ρ Π°Π½Π΄ΡΠΎΠ³Π΅Π½-Π·Π°Π²ΠΈΡΠΈΠΌΡΠΉ Ρ Π°ΡΠ°ΠΊΡΠ΅Ρ (Mugford, Nowell, 1970, Novotny, et al., 1985, Novotny, et al., 1990, Novikov, 1993, Chamero, et al., 2007). ΠΠ΅ΡΠ΅ΡΡΠ²Π°-ΡΡΠ΅ΡΠΎΠΈΠ΄Π½ΠΎΠΉ ΠΏΡΠΈΡΠΎΠ΄Ρ, ΠΊΠ°ΠΊ ΠΌΠ΅ΡΠ°Π±ΠΎΠ»ΠΈΡΡ Π³ΠΎΡΠΌΠΎΠ½ΠΎΠ², ΡΠΈΡΠΊΡΠ»ΠΈΡΡΡΡΠΈΡ Π² ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠ΅ ΠΎΡΠΎΠ±ΠΈΠΌΠΎΠ³ΡΡ Π±ΡΡΡ ΡΠ°ΡΡΠΌΠΎΡΡΠ΅Π½Ρ ΠΊΠ°ΠΊ ΠΊΠ°Π½Π΄ΠΈΠ΄Π°ΡΡ Π½Π° ΡΠΎΠ»Ρ Π΅ΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΡΡ Ρ Π΅ΠΌΠΎΡΠΈΠ³Π½Π°Π»ΠΎΠ² Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ ΡΠΎΡΠΈΠ°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ (Zahavi, 1975, Ingersoli, Launay, 1986, Nodari, et al., 2008). ΠΠ° ΠΏΠΎΡΠ»Π΅Π΄Π½ΠΈΠ΅ Π΄Π΅ΡΡΡΠΈΠ»Π΅ΡΠΈΡ ΠΈΠ· ΠΌΠΎΡΠΈ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ Π±ΡΠ» Π²ΡΠ΄Π΅Π»Π΅Π½ ΡΡΠ΄ Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠΎΠ΅Π΄ΠΈΠ½Π΅Π½ΠΈΠΉ Π½Π΅ΡΡΠ΅ΡΠΎΠΈΠ΄Π½ΠΎΠΉ ΠΏΡΠΈΡΠΎΠ΄Ρ, ΡΡΠ°ΡΡΠ²ΡΡΡΠΈΡ Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ, ΠΎΠ΄Π½Π°ΠΊΠΎ Π½ΠΈ ΠΎΠ΄Π½ΠΎ ΠΈΠ· ΡΡΠΈΡ Π²Π΅ΡΠ΅ΡΡΠ² Π½Π΅ ΠΈΠΌΠΈΡΠΈΡΡΠ΅Ρ ΠΏΠΎΠ»Π½ΠΎΡΡΡΡ Ρ Π΅ΠΌΠΎΡΠΈΠ³Π½Π°Π» ΠΌΠΎΡΠΈ (Novotny, et al., 1985, Chamero, et al., 2007).
Π ΠΊΠ°ΡΠ΅ΡΡΠ²Π΅ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΠΎΠ³ΠΎ ΠΎΠ΄ΠΎΡΠ°Π½ΡΠ° Π² Π½Π°ΡΠΈΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡΡ Π±ΡΠ» ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ ΡΡΠ΅ΡΠΎΠΈΠ΄ Π³ΠΎΠ½Π°Π΄Π½ΠΎΠ³ΠΎ ΠΏΡΠΎΠΈΡΡ ΠΎΠΆΠ΄Π΅Π½ΠΈΡ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½ (5Π°-Π°Π½Π΄ΡΠΎΡΡ-16-Π΅Π½-3-ΠΎΠ½), Π½Π°ΠΈΠ±ΠΎΠ»Π΅Π΅ ΠΈΠ·Π²Π΅ΡΡΠ½ΡΠΉ ΠΊΠ°ΠΊ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠΉ ΡΠ΅ΡΠΎΠΌΠΎΠ½ Ρ ΡΡΠΊΠ°, Π²ΡΠ·ΡΠ²Π°ΡΡΠΈΠΉ Π½Π΅ΠΌΠ΅Π΄Π»Π΅Π½Π½ΡΠΉ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΡΠ΅ΡΠΊΠΈΠΉ ΠΎΡΠ²Π΅Ρ Ρ ΡΠ΅ΡΠ΅ΠΏΡΠΈΠ²Π½ΠΎΠΉ ΡΠ°ΠΌΠΊΠΈ ΡΠ²ΠΈΠ½ΡΠΈ (Reed, et al., 1974). ΠΠ½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½ ΡΠ²Π»ΡΠ΅ΡΡΡ ΠΈ ΠΊΠ°Π½Π΄ΠΈΠ΄Π°ΡΠΎΠΌ Π½Π° ΡΠΎΠ»Ρ ΠΌΠΎΠ΄ΡΠ»ΡΡΠΎΡΠ½ΠΎΠ³ΠΎ ΡΠ΅ΡΠΎΠΌΠΎΠ½Π° Ρ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° (Cowley, et al., 1977, Filsinger, et al., 1984, Pause, 2004). Π£ 30−50% Π² ΠΏΠΎΠΏΡΠ»ΡΡΠΈΠΈ Π»ΡΠ΄Π΅ΠΉ Π½Π°Π±Π»ΡΠ΄Π°Π΅ΡΡΡ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠ°Ρ Π°Π½ΠΎΡΠΌΠΈΡ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ, Ρ. Π΅. Π½Π΅ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡ ΡΠ°ΡΠΏΠΎΠ·Π½Π°Π²Π°ΡΡ ΠΈΠΌΠ΅Π½Π½ΠΎ ΡΡΠΎΡ Π·Π°ΠΏΠ°Ρ . Π½Π° ΡΠΎΠ½Π΅, Π½ΠΎΡΠΌΠ°Π»ΡΠ½ΠΎΠΉ-ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π΄ΡΡΠ³ΠΈΠΌ ΠΎΠ΄ΠΎΡΠ°Π½ΡΠ°ΠΌ" (Labows, Wysocki, 1984). Π€ΡΠ½ΠΊΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠ΅ Π·Π½Π°ΡΠ΅Π½ΠΈΠ΅ ΡΡΠΎΠΉ, ΠΎΠ΄Π½ΠΎΠΉΠΈΠ·^ ΡΠ°ΠΌΡΡ ΡΠ°ΡΠΏΡΠΎΡΡΡΠ°Π½Π΅Π½Π½ΡΡ Ρ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ°, Π°Π½ΠΎΡΠΌΠΈΠΈΠ½Π΅ΡΡΠ½ΠΎ. Π‘ΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ Ρ Π΅ΠΌΠΎΠ΄Π΅ΡΠΈΡΠΈΡΡ Π²ΡΠ·ΡΠ²Π°ΡΡ ΠΎΠ³ΡΠΎΠΌΠ½ΡΠΉ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΡΠΊΠΈΠΉ ΠΈΠ½ΡΠ΅ΡΠ΅Ρ, ΠΏΡΠ΅Π΄ΠΎΡΡΠ°Π²Π»ΡΡ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΡΡΡ ΡΠ²ΡΠ·Π°ΡΡ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΡ. ΡΡΠ½ΠΊΡΠΈΡ Ρ ΠΊΠΎΠ½ΠΊΡΠ΅ΡΠ½ΡΠΌΠΈ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠ°ΠΌΠΈ ΠΈ Π³Π΅Π½Π°ΠΌΠΈ (Griff, Reed, 1995, Keller, et al., 2007).
ΠΠ°Π±ΠΎΡΠ°ΡΠΎΡΠ½Π°Ρ ΠΌΡΡΡ ΡΠ²Π»ΡΠ΅ΡΡΡ, ΠΏΡΠ΅Π²ΠΎΡΡ ΠΎΠ΄Π½ΡΠΌ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΠΌ ΠΎΡΠ³Π°Π½ΠΈΠ·ΠΌΠΎΠΌ Π΄Π»Ρ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ Π² ΠΎΠ±Π»Π°ΡΡΠΈ Π³Π΅Π½Π΅ΡΠΈΠΊΠΈ ΠΈ Π±ΠΈΠΎΠΌΠ΅Π΄ΠΈΡΠΈΠ½Ρ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° (Silver, 1995). 90% Π³Π΅Π½ΠΎΠ², ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° ΠΈΠΌΠ΅Π΅Ρ ΡΠ²ΠΎΠΈ Π³ΠΎΠΌΠΎΠ»ΠΎΠ³ΠΈ Ρ ΠΌΡΡΠΈ, ΠΈ, ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΠΎΠ½Π°ΠΎΠ±ΠΎΡΠΎΡ. ΠΡΠΎΡΡΠ΅Π½ΠΈΠ΅ ΠΏΠ΅ΡΠ²ΠΈΡΠ½ΠΎΠΉ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ' Π³Π΅Π½ΠΎΠΌΠ° ΠΌΡΡΠΈ Π±ΡΠ»ΠΎ Π² ΡΠ΅ΡΠ½ΠΎΠ²ΠΎΠΌ* Π²Π°ΡΠΈΠ°Π½ΡΠ΅ Π·Π°Π²Π΅ΡΡΠ΅Π½ΠΎ Π² 2002 Π³ΠΎΠ΄Ρ (Waterston, et al, 2002), ΠΎΠ΄Π½Π°ΠΊΠΎ, Π΄ΠΎΠΏΠΎΠ»Π½Π΅Π½ΠΈΠ΅ ΠΈ Π°Π½Π½ΠΎΡΠ°ΡΠΈΡ Π½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄Π½ΡΡ ΠΏΠΎΡΠ»Π΅Π΄ΠΎΠ²Π°ΡΠ΅Π»ΡΠ½ΠΎΡΡΠ΅ΠΉ ΠΏΡΠΎΠ΄ΠΎΠ»ΠΆΠ°Π΅ΡΡΡ* ΠΈ ΠΏΠΎ ΡΠ΅ΠΉ Π΄Π΅Π½Ρ, ΡΡΠΎ ΠΎΡΡΠ°ΠΆΠ°Π΅ΡΡΡ" Π² ΠΏΠΎΡΡΠΎΡΠ½Π½ΠΎΠΌ' ΠΎΠ±Π½ΠΎΠ²Π»Π΅Π½ΠΈΠΈΡΠΏΠ΅ΡΠΈΠ°Π»ΠΈΠ·ΠΈΡΠΎΠ²Π°Π½Π½ΡΡ Π±Π°Π·. Π΄Π°Π½Π½ΡΡ (Π½Π°ΠΏΡΠΈΠΌΠ΅Ρ, MGI, NCBI). ΠΠΊΡΡΠ°Π»ΡΠ½ΠΎΡΡΡ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΉ Π½Π°Π»Π°Π±ΠΎΡΠ°ΡΠΎΡΠ½ΡΡ ΠΌΡΡΠ°Ρ ΡΠΎΠ»ΠΈ Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠΈΠ³Π½Π°Π»ΠΎΠ² Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Π½Π΅ Π²ΡΠ·ΡΠ²Π°Π΅Ρ ΡΠΎΠΌΠ½Π΅Π½ΠΈΠΉ. ΠΠ΅ΡΠΎΠ΄ ΠΊΠ°ΡΡΠΈΡΠΎΠ²Π°Π½ΠΈΡ Π»ΠΎΠΊΡΡΠΎΠ² ΠΊΠΎΠ»ΠΈΡΠ΅ΡΡΠ²Π΅Π½Π½ΡΡ * ΠΏΡΠΈΠ·Π½Π°ΠΊΠΎΠ², ΠΎΡΠ½ΠΎΠ²Π°Π½Π½ΡΠΉ Π½Π° Π°Π½Π°Π»ΠΈΠ·Π΅ Π°ΡΡΠΎΡΠΈΠ°ΡΠΈΠΉ: ΠΌΠ΅ΠΆΠ΄Ρ. ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΡΠΌΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠΌΠΈ-ΠΌΠ°ΡΠΊΠ΅ΡΠ°ΠΌΠΈ*ΠΈ ΡΠ΅Π½ΠΎΡΠΈΠΏΠΈΡΠ΅ΡΠΊΠΈΠΌΠΈ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ°ΠΌΠΈ, ΡΠ²Π»ΡΠ΅ΡΡΡ, Π²Π°ΠΆΠ½ΡΠΌ ΠΈΠ½ΡΡΡΡΠΌΠ΅Π½ΡΠΎΠΌ^ Π΄Π»Ρ ΡΠ΅Π°Π»ΠΈΠ·Π°ΡΠΈΠΈ ΠΎΠ±ΡΠ΅Π³ΠΎ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ ΠΏΠΎΠ΄Ρ ΠΎΠ΄Π° ΠΊ ΠΈΠ΄Π΅Π½ΡΠΈΡΠΈΠΊΠ°ΡΠΈΠΈ Π³Π΅Π½ΠΎΠ², ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΡ ΠΏΡΠΈΠ·Π½Π°ΠΊ, ΠΏΠΎ ΠΏΡΡΠΈ «ΠΎΡ ΡΠ΅Π½ΠΎΡΠΈΠΏΠ° ΠΊ Π³Π΅Π½Ρ». ΠΠ΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠ°Ρ ΠΌΠΎΠ΄Π΅Π»Ρ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ-(Π½Π° ΠΏΡΠΈΠΌΠ΅ΡΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°) Π±ΡΠ»Π° ΡΠ°Π·ΡΠ°Π±ΠΎΡΠ°Π½Π° ΠΏΡΠΈ ΡΠΎΡΡΡΠ΄Π½ΠΈΡΠ΅ΡΡΠ²Π΅, Π½Π°ΡΡΠ½ΡΡ , Π³ΡΡΠΏΠΏ Π§. ΠΠ°ΠΉΡΠΎΠΊΠΈ ΠΈ Π. Π. ΠΠΎΠ·Π½Π΅ΡΠ΅Π½ΡΠΊΠΎΠΉ Π½Π° ΠΎΡΠ½ΠΎΠ²Π΅ 2-Ρ ΠΈΠ½Π±ΡΠ΅Π΄Π½ΡΡ Π»ΠΈΠ½ΠΈΠΉ ΠΌΡΡΠ΅ΠΉ NZB/B1NJ* ΠΈ. CBA/J, Π²ΡΡΠΎΠΊΠΎΠΊΠΎΠ½ΡΡΠ°ΡΡΠ½ΡΡ ΠΏΠΎ ΡΡΠΎΠΌΡ ΠΏΡΠΈΠ·Π½Π°ΠΊΡ (Voznessenskaya, et al, 1995). ΠΠ°Π»ΠΎΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΡΠ΅ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΌΡΡΠΈ Π»ΠΈΠ½ΠΈΠΈ NZB/B1NJ ΠΈΠΌΠ΅ΡΡ ΠΎΡΠ΅Π½Ρ Π²ΡΡΠΎΠΊΠΈΠΉ ΡΡΠΎΠ²Π΅Π½Ρ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΡΡΠΈ ΠΏΠΎ ΡΡΠ°Π²Π½Π΅Π½ΠΈΡ ΡΠΎ. ΠΌΠ½ΠΎΠ³ΠΈΠΌΠΈ Π΄ΡΡΠ³ΠΈΠΌΠΈ ΠΈΠ½Π±ΡΠ΅Π΄Π½ΡΠΌΠΈ Π»ΠΈΠ½ΠΈΡΠΌΠΈ. (Roubertoux, Carlier, 1988; Roubertoux, et al., 2005) ΠΈ, Π² ΡΠ°ΡΡΠ½ΠΎΡΡΠΈ, Ρ CBA/J (Voznessenskaya, Wysocki, 2000). Π‘ΠΎΠ³Π»Π°ΡΠ½ΠΎ Π½Π°ΡΠ΅ΠΉ Π³ΠΈΠΏΠΎΡΠ΅Π·Π΅, Ρ Π΅ΠΌΠΎΠ΄Π΅ΡΠΈΡΠΈΡ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ ΠΌΠΎΠΆΠ΅Ρ ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡΡ ΠΊ ΠΎΡΠΈΠ±ΠΊΠ°ΠΌ Π² ΡΠ°ΡΠΏΠΎΠ·Π½Π°Π²Π°Π½ΠΈΠΈ Π·Π°ΠΏΠ°Ρ ΠΎΠ²ΠΎΠ³ΠΎ ΡΠΈΠ³Π½Π°Π»Π°Π½Π΅ΡΡΡΠ΅Π³ΠΎ ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΡ ΠΎ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠΉ ΠΏΡΠΈΠ½Π°Π΄Π»Π΅ΠΆΠ½ΠΎΡΡΠΈ ΠΈ ΡΠΎΡΠΈΠ°Π»ΡΠ½ΠΎΠΌ ΡΡΠ°ΡΡΡΠ΅, ΡΡΠΎ, Π² ΡΠ²ΠΎΡ ΠΎΡΠ΅ΡΠ΅Π΄Ρ, ΠΌΠΎΠΆΠ΅Ρ ΡΠ»ΡΠΆΠΈΡΡ ΠΎΠ΄Π½ΠΎΠΉ ΠΈΠ· ΠΏΡΠΈΡΠΈΠ½ ΠΏΡΠΎΡΠ²Π»Π΅Π½ΠΈΡ ΠΈΠ·Π±ΡΡΠΎΡΠ½ΠΎΠΉ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠΈ.
Π¦Π΅Π»Ρ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ. Π¦Π΅Π»ΡΡ Π΄Π°Π½Π½ΠΎΠΉ ΡΠ°Π±ΠΎΡΡ Π±ΡΠ»ΠΎ ΠΌΠΎΠ΄Π΅Π»ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ Π½Π° ΠΏΡΠΈΠΌΠ΅ΡΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° ΡΠ²ΡΠ·ΠΈ ΠΌΠ΅ΠΆΠ΄Ρ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡΡ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ ΠΈ ΡΠΎΡΠΈΠ°Π»ΡΠ½ΡΠΌ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠ΅ΠΌ-Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ, Π° ΡΠ°ΠΊΠΆΠ΅ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΠΈ Π»Π΅ΡΡΡΠΈΡ ΡΡΠ΅ΡΠΎΠΈΠ΄ΠΎΠ² Π² ΠΊΠ°ΡΠ΅ΡΡΠ²Π΅ Ρ Π΅ΠΌΠΎΡΠΈΠ³Π½Π°Π»ΠΎΠ².
ΠΠ°Π΄Π°ΡΠΈ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ. ΠΡΠ»ΠΈ ΡΡΠΎΡΠΌΡΠ»ΠΈΡΠΎΠ²Π°Π½Ρ ΡΠ»Π΅Π΄ΡΡΡΠΈΠ΅ Π·Π°Π΄Π°ΡΠΈ:
1. ΠΠ½Π°Π»ΠΈΠ· Π½Π°ΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ ΠΈ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠΈ Ρ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ, Π²ΠΊΠ»ΡΡΠ°Ρ ΠΊΠ°ΡΡΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΡ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π»ΠΎΠΊΡΡΠΎΠ² ΠΈ ΠΌΠΎΠ΄Π΅Π»ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΎΠ΄Π½ΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎΠ³ΠΎ ΡΡΠ°ΡΡΠΈΡ Π½Π΅ΡΠΊΠΎΠ»ΡΠΊΠΈΡ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ°ΠΊΡΠΎΡΠΎΠ², ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΡ ΡΡΠΈ ΠΏΡΠΈΠ·Π½Π°ΠΊΠΈ, Π° ΡΠ°ΠΊΠΆΠ΅ ΠΏΠΎΠΈΡΠΊ Π³Π΅Π½ΠΎΠ²-ΠΊΠ°Π½Π΄ΠΈΠ΄Π°ΡΠΎΠ² ΠΏΠΎ Π±Π°Π·Π°ΠΌ Π΄Π°Π½Π½ΡΡ .
2. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠ²ΡΠ·ΠΈ ΠΌΠ΅ΠΆΠ΄Ρ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡΡ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ ΠΈ ΡΠΏΠΎΡΠΎΠ±Π½ΠΎΡΡΡΡ ΡΠ°ΡΠΏΠΎΠ·Π½Π°Π²Π°ΡΡ ΠΏΠΎ Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠΌ ΡΠΈΠ³Π½Π°Π»Π°ΠΌ ΠΏΠΎΠ» ΠΈ ΡΠ΅ΠΏΡΠΎΠ΄ΡΠΊΡΠΈΠ²Π½ΡΠΉ ΡΡΠ°ΡΡΡ ΠΎΡΠΎΠ±Π΅ΠΉ ΡΠ²ΠΎΠ΅Π³ΠΎ Π²ΠΈΠ΄Π° Π½Π° ΡΡΠΎΠ²Π½Π΅ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Π½Π° ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ.
3. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΡΡΠ°ΡΡΠΈΡ Π΄ΠΎΠΏΠΎΠ»Π½ΠΈΡΠ΅Π»ΡΠ½ΠΎΠΉ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π² ΡΠ΅ΡΠ΅ΠΏΡΠΈΠΈ ΡΡΠΎΠ³ΠΎ ΠΎΠ΄ΠΎΡΠ°Π½ΡΠ° Π½Π° ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ.
4. ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ Π½Π° ΡΡΠΎΠ²Π½Π΅ Π³ΠΎΡΠΌΠΎΠ½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΈ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΡΠ΅ΡΠΊΠΎΠ³ΠΎ ΠΎΡΠ²Π΅ΡΠ° ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΎΠ½Π½ΠΎΠΉ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΠΈ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° ΠΊΠ°ΠΊ Ρ Π΅ΠΌΠΎΡΠΈΠ³Π½Π°Π»Π° Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ.
ΠΠ°ΡΡΠ½Π°Ρ Π½ΠΎΠ²ΠΈΠ·Π½Π°. ΠΠΏΠ΅ΡΠ²ΡΠ΅ Π½Π° ΠΎΡΠ½ΠΎΠ²Π΅ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π° Π³ΠΈΠ±ΡΠΈΠ΄ΠΎΠ² Π²ΡΠΎΡΠΎΠ³ΠΎ ΠΏΠΎΠΊΠΎΠ»Π΅Π½ΠΈΡ ΠΎΡ ΡΠΊΡΠ΅ΡΠΈΠ²Π°Π½ΠΈΡ ΠΈΠ½Π±ΡΠ΅Π΄Π½ΡΡ Π»ΠΈΠ½ΠΈΠΈ ΠΌΡΡΠ΅ΠΉ CBA/J ΠΈ NZB/B1NJ ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Ρ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ½ΡΠ΅ Π»ΠΎΠΊΡΡΡ, ΠΎΠΊΠ°Π·ΡΠ²Π°ΡΡΠΈΠ΅ Π²Π»ΠΈΡΠ½ΠΈΠ΅ Π½Π° ΡΠ°ΠΊΠΈΠ΅ ΠΏΡΠΈΠ·Π½Π°ΠΊΠΈ,* ΠΊΠ°ΠΊ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½Π°Ρ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡ ΠΊ Π»Π΅ΡΡΡΠ΅ΠΌΡ ΡΡΠ΅ΡΠΎΠΈΠ΄Ρ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΈ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²Π°Ρ Π°Π³ΡΠ΅ΡΡΡΠΈΡ. ΠΠΏΠ΅ΡΠ²ΡΠ΅ Π½Π° ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ ΠΎΡΡΡΡΡΡΠ²ΠΈΠ΅ ΠΏΡΠ΅Π΄ΠΏΠΎΡΡΠ΅Π½ΠΈΡ Π·Π°ΠΏΠ°Ρ Π° ΡΡΡΡΠ°Π»ΡΠ½ΠΎΠΉ ΡΠ°ΠΌΠΊΠΈ ΠΏΠΎ ΡΡΠ°Π²Π½Π΅Π½ΠΈΡ Ρ Π·Π°ΠΏΠ°Ρ ΠΎΠΌ ΡΠ°ΠΌΡΠ° Ρ ΡΠ°ΠΌΡΠΎΠ² ΠΌΡΡΠ΅ΠΉ Ρ Ρ Π΅ΠΌΠΎΠ΄Π΅ΡΠΈΡΠΈΡΠΎΠΌ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ. ΠΠΏΠ΅ΡΠ²ΡΠ΅ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ ΡΡΠ°ΡΡΠΈΠ΅ Π΄ΠΎΠΏΠΎΠ»Π½ΠΈΡΠ΅Π»ΡΠ½ΠΎΠΉ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΡ Π² Π΄Π΅ΡΠ΅ΠΊΡΠΈΠΈ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°. ΠΠΏΠ΅ΡΠ²ΡΠ΅ Π½Π° ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ Π±ΡΠ» ΠΏΠΎΠ»ΡΡΠ΅Π½ ΠΎΡΠ²Π΅Ρ Π½Π° ΡΡΠΈΠΌΡΠ»ΡΡΠΈΡ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½ΠΎΠΌ Π½Π° ΡΡΠΎΠ²Π½Π΅ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠ½ΠΎΠΉ ΡΠΊΠ°Π½ΠΈ Π²ΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ³Π°Π½Π°. ΠΠΏΠ΅ΡΠ²ΡΠ΅ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΏΡΠ΅Π΄ΡΡΠ²Π»Π΅Π½ΠΈΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° (0.1%, 30 ΠΌΠΈΠ½), Π²ΡΠ·ΡΠ²Π°Π΅Ρ ΡΠ½ΠΈΠΆΠ΅Π½ΠΈΠ΅ ΡΡΠΎΠ²Π½Ρ ΡΠ΅ΡΡΠΎΡΡΠ΅ΡΠΎΠ½Π° Π² ΠΏΠ»Π°Π·ΠΌΠ΅ ΠΊΡΠΎΠ²ΠΈ Ρ ΡΠ°ΠΌΡΠΎΠ² Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ.
Π’Π΅ΠΎΡΠ΅ΡΠΈΡΠ΅ΡΠΊΠ°Ρ ΠΈ ΠΏΡΠ°ΠΊΡΠΈΡΠ΅ΡΠΊΠ°Ρ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΡ. ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΡ ΡΠΏΠΎΡΠΎΠ±ΡΡΠ²ΡΡΡ Π½Π°ΠΊΠΎΠΏΠ»Π΅Π½ΠΈΡ ΠΈ ΠΈΠ½ΡΠ΅Π³ΡΠ°ΡΠΈΠΈ Π΄Π°Π½Π½ΡΡ ΠΎ ΡΠ΅Π½ΠΎΡΠΈΠΏΠΈΡΠ΅ΡΠΊΠΈΡ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ°Ρ Ρ ΠΈΠ½Π±ΡΠ΅Π΄Π½ΡΡ Π»ΠΈΠ½ΠΈΠΉ ΠΌΡΡΠ΅ΠΉ-ΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π»ΠΎΠΊΡΡΠ°Ρ , ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΡ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΡ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ (Π½Π° ΠΏΡΠΈΠΌΠ΅ΡΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°), Π° ΡΠ°ΠΊΠΆΠ΅ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΡΡ Π°Π³ΡΠ΅ΡΡΠΈΡ, ΡΠ°ΡΡΠΈΡΡΡ ΡΠ²Π΅Π΄Π΅Π½ΠΈΡ ΡΡΡΠ΅ΡΡΠ²ΡΡΡΠΈΡ Π±Π°Π· Π΄Π°Π½Π½ΡΡ . ΠΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π΄Π°Π½Π½ΡΠ΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΡΡ ΠΏΠΎΠ΄ΠΎΠΉΡΠΈ Π±Π»ΠΈΠΆΠ΅ ΠΊ ΠΏΠΎΠ½ΠΈΠΌΠ°Π½ΠΈΡ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ ΠΎΡΠ½ΠΎΠ² ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Ρ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ°. ΠΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½Π°Ρ ΡΡΠ½ΠΊΡΠΈΡ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ° ΡΠ°ΡΡΠΌΠ°ΡΡΠΈΠ²Π°Π΅ΡΡΡ ΠΊΠ°ΠΊ ΠΏΠ΅ΡΡΠΏΠ΅ΠΊΡΠΈΠ²Π½ΡΠΉ ΠΌΠ°ΡΠΊΠ΅Ρ ΡΡΠ΄Π° Π½Π΅ΠΉΡΠΎΠ΄Π΅Π³Π΅Π½Π΅ΡΠ°ΡΠΈΠ²Π½ΡΡ Π·Π°Π±ΠΎΠ»Π΅Π²Π°Π½ΠΈΠΉ, Π² ΡΠΎΠΌ ΡΠΈΡΠ»Π΅ Π±ΠΎΠ»Π΅Π·Π½ΠΈ ΠΠ»ΡΡΠ³Π΅ΠΉΠΌΠ΅ΡΠ°, ΠΏΠ°ΡΠΊΠΈΠ½ΡΠΎΠ½ΠΈΠ·ΠΌΠ° ΠΈ Π΄Ρ. (Doty et al, 2008, Atanasova et al, 2008).
Π Π΅ΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΠΎΠΉ ΡΡΠ΅Π΄Π΅ ΠΎΠ±ΠΈΡΠ°Π½ΠΈΡ Π΄ΠΎΠΌΠΎΠ²ΡΠ΅ ΠΌΡΡΠΈ Π½Π°Π½ΠΎΡΡΡ ΡΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΡΠΉ ΡΠΊΠΎΠ½ΠΎΠΌΠΈΡΠ΅ΡΠΊΠΈΠΉ Π²ΡΠ΅Π΄ Ρ ΠΎΠ·ΡΠΉΡΡΠ²Ρ ΠΈ ΡΠ²Π»ΡΡΡΡΡ ΠΏΠ΅ΡΠ΅Π½ΠΎΡΡΠΈΠΊΠ°ΠΌΠΈ ΠΎΠΏΠ°ΡΠ½ΡΡ ΠΈΠ½ΡΠ΅ΠΊΡΠΈΠΎΠ½Π½ΡΡ Π·Π°Π±ΠΎΠ»Π΅Π²Π°Π½ΠΈΠΉ (ΠΡΠ»ΠΈΠΊ, 1979). ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠ΅ ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΎΠ½Π½ΠΎΠΉ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΠΈ Π»Π΅ΡΡΡΠΈΡ ΡΡΠ΅ΡΠΎΠΈΠ΄ΠΎΠ² Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΈ ΠΏΠΎΠ»ΠΎΠ²ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Π³ΡΡΠ·ΡΠ½ΠΎΠ² ΠΏΠΎΠ·Π²ΠΎΠ»ΡΠ΅Ρ ΠΎΠΏΡΠΈΠΌΠΈΠ·ΠΈΡΠΎΠ²Π°ΡΡ ΠΏΡΠΈΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ ΡΠΈΠ³Π½Π°Π»ΡΠ½ΡΡ ΠΌΠΎΠ»Π΅ΠΊΡΠ» Π½Π° ΠΏΡΠ°ΠΊΡΠΈΠΊΠ΅ ΠΊΠ°ΠΊ Π΅ΡΡΠ΅ΡΡΠ²Π΅Π½Π½ΡΡ ΡΠ΅ΠΏΠ΅Π»Π»Π΅Π½ΡΠΎΠ² ΠΈ/ΠΈΠ»ΠΈ ΡΠ΅Π³ΡΠ»ΡΡΠΎΡΠΎΠ² ΡΠ΅ΠΏΡΠΎΠ΄ΡΠΊΡΠΈΠ²Π½ΠΎΠΉ ΡΡΠ½ΠΊΡΠΈΠΈ.
ΠΠ»Π°Π³ΠΎΠ΄Π°ΡΠ½ΠΎΡΡΠΈ. ΠΠ²ΡΠΎΡ Π²ΡΡΠ°ΠΆΠ°Π΅Ρ Π±Π»Π°Π³ΠΎΠ΄Π°ΡΠ½ΠΎΡΡΡ ΠΊΠ°Π½Π΄ΠΈΠ΄Π°ΡΡ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ Π½Π°ΡΠΊ, ΡΡΠ°ΡΡΠ΅ΠΌΡ Π½Π°ΡΡΠ½ΠΎΠΌΡ ΡΠΎΡΡΡΠ΄Π½ΠΈΠΊΡ ΠΠ½ΡΡΠΈΡΡΡΠ° ΠΏΡΠΎΠ±Π»Π΅ΠΌ ΡΠΊΠΎΠ»ΠΎΠ³ΠΈΠΈ ΠΈ ΡΠ²ΠΎΠ»ΡΡΠΈΠΈ ΠΈΠΌ. Π. Π. Π‘Π΅Π²Π΅ΡΡΠΎΠ²Π° Π ΠΠ Π. Π. ΠΠΎΠ·Π½Π΅ΡΠ΅Π½ΡΠΊΠΎΠΉ, ΠΏΠΎΠ΄ Π½Π°ΡΡΠ½ΡΠΌ ΡΡΠΊΠΎΠ²ΠΎΠ΄ΡΡΠ²ΠΎΠΌ ΠΊΠΎΡΠΎΡΠΎΠΉ Π±ΡΠ»Π° Π²ΡΠΏΠΎΠ»Π½Π΅Π½Π° ΡΡΠ° ΡΠ°Π±ΠΎΡΠ°.
ΠΠ²ΡΠΎΡ Π³Π»ΡΠ±ΠΎΠΊΠΎ ΠΏΡΠΈΠ·Π½Π°ΡΠ΅Π»Π΅Π½ ΡΠΎΡΡΡΠ΄Π½ΠΈΠΊΠ°ΠΌ ΠΠΎΠ½Π΅Π»Π»Π΅Π²ΡΠΊΠΎΠ³ΠΎ ΡΠ΅Π½ΡΡΠ° Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ ΡΡΠ²ΡΡΠ² (Π€ΠΈΠ»Π°Π΄Π΅Π»ΡΡΠΈΡ, Π‘Π¨Π) ΠΡ. Π. ΠΠ°ΡΠΌΠ°Π½ΠΎΠ²Ρ, ΠΡ. Π. ΠΠΎΡΠ°ΠΊ ΠΈ ΠΡ. Π¦. ΠΠΈ Π·Π° ΠΏΠΎΠΌΠΎΡΡ Π² ΠΎΡΠ²ΠΎΠ΅Π½ΠΈΠΈ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎ-Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈ Π±ΠΈΠΎΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΠΌΠ΅ΡΠΎΠ΄ΠΎΠ², Π° ΡΠ°ΠΊΠΆΠ΅ Π·Π° ΡΠ΅Π½Π½ΡΠ΅ Π·Π°ΠΌΠ΅ΡΠ°Π½ΠΈΡ ΠΈ ΡΠ΅ΠΊΠΎΠΌΠ΅Π½Π΄Π°ΡΠΈΠΈ ΠΏΠΎ ΡΠ°Π±ΠΎΡΠ΅.
ΠΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΡΠ°ΡΡΠΈΡΠ½ΠΎ ΠΏΠΎΠ΄Π΄Π΅ΡΠΆΠ°Π½Ρ Π Π€Π€Π 07−04−1 538 ΠΈ ΠΡΠΎΠ³ΡΠ°ΠΌΠΌΠΎΠΉ ΠΡΠ΅Π·ΠΈΠ΄ΠΈΡΠΌΠ° Π ΠΠ «ΠΠΈΠ½Π°ΠΌΠΈΠΊΠ° Π³Π΅Π½ΠΎΡΠΎΠ½Π΄ΠΎΠ² ΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠ΅ ΡΠ°Π·Π½ΠΎΠΎΠ±ΡΠ°Π·ΠΈΠ΅» .
ΠΡΠ²ΠΎΠ΄Ρ:
1. Π‘ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΎΠΏΠΈΡΠ°Π½ ΠΏΠΎΠ»ΠΈΠ³Π΅Π½Π½ΡΠΉ Ρ Π°ΡΠ°ΠΊΡΠ΅Ρ Π½Π°ΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ. ΠΠΏΠ΅ΡΠ²ΡΠ΅ Π²ΡΡΠ²Π»Π΅Π½Ρ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΠ΅Π»ΡΠ½ΡΠ΅ Π»ΠΎΠΊΡΡΡ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΠ΅ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ Π½Π° 2-ΠΎΠΉ (rs3023694), 12-ΠΎΠΉ (rs3684371) ΠΈ 17-ΠΎΠΉ (rs3675244) Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ°Ρ , Π°Π΄Π΄ΠΈΡΠΈΠ²Π½ΠΎ ΠΎΠ±ΡΡΡΠ½ΡΡΡΠΈΠ΅ 25% ΡΠ΅Π½ΠΎΡΠΈΠΏΠΈΡΠ΅ΡΠΊΠΎΠΉ Π²Π°ΡΠΈΠ°Π±Π΅Π»ΡΠ½ΠΎΡΡΠΈ. ΠΠ° ΠΎΡΠ½ΠΎΠ²Π΅ ΡΠ΅Π·ΡΠ»ΡΡΠ°ΡΠΎΠ² Π°Π½Π°Π»ΠΈΠ·Π° QTL, ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½Π½ΠΎΠ³ΠΎ ΠΏΠΎ ΠΎΡΠ΄Π΅Π»ΡΠ½ΠΎΡΡΠΈ Π΄Π»Ρ ΡΠ°ΠΌΡΠΎΠ² ΠΈ ΡΠ°ΠΌΠΎΠΊ ΠΏΡΠ΅Π΄Π»ΠΎΠΆΠ΅Π½Ρ ΠΌΠΎΠ΄Π΅Π»ΠΈ, ΠΎΠ±ΡΡΡΠ½ΡΡΡΠΈΠ΅, ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΠΎ 59% ΠΈ 63% ΡΠ΅Π½ΠΎΡΠΈΠΏΠΈΡΠ΅ΡΠΊΠΎΠΉ Π²Π°ΡΠΈΠ°Π±Π΅Π»ΡΠ½ΠΎΡΡΠΈ. Π ΠΎΠ±Π»Π°ΡΡΡΡ ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½Π½ΡΡ Π»ΠΎΠΊΡΡΠΎΠ² Π±ΡΠ» ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½ ΠΏΠΎΠΈΡΠΊ Π³Π΅Π½ΠΎΠ²-ΠΊΠ°Π½Π΄ΠΈΠ΄Π°ΡΠΎΠ².
2. ΠΠΏΠ΅ΡΠ²ΡΠ΅ Ρ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ Π΄Π»Ρ ΠΏΠΎΠΊΠ°Π·Π°ΡΠ΅Π»Π΅ΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Π²ΡΡΠ²Π»Π΅Π½Ρ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΠ΅Π»ΡΠ½ΡΠ΅ Π»ΠΎΠΊΡΡΡ Π½Π° 8-ΠΎΠΉ (D8Mit205, rs4227112), 10-ΠΎΠΉ (rs3693721) ΠΈ 1-ΠΎΠΉ (D3Mit217) Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ°Ρ , Π° ΡΠ°ΠΊΠΆΠ΅ ΠΎΠΏΠΈΡΠ°Π½ ΡΡΠ΄ ΡΠΏΠΈΡΡΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠΉ. ΠΠΎΠ²Π΅ΡΠΈΡΠ΅Π»ΡΠ½ΡΠ΅ ΠΈΠ½ΡΠ΅ΡΠ²Π°Π»Ρ Π΄Π»Ρ Π΄ΠΎΡΡΠΎΠ²Π΅ΡΠ½ΠΎΠ³ΠΎ Π»ΠΎΠΊΡΡΠ° Π½Π° 10-ΠΎΠΉ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅, ΡΠ°ΡΡΠΈΡΠ½ΠΎ ΠΎΠ±ΡΡΡΠ½ΡΡΡΠ΅Π³ΠΎ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ, ΠΈ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΠ΅Π»ΡΠ½ΠΎΠ³ΠΎ Π»ΠΎΠΊΡΡΠ°, Π²ΡΡΠ²Π»Π΅Π½Π½ΠΎΠ³ΠΎ Π΄Π»Ρ ΠΏΠΎΠΊΠ°Π·Π°ΡΠ΅Π»Π΅ΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΡΡΠΈ* Π½Π° ΡΠΎΠΉ ΠΆΠ΅ Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅, ΠΏΠ΅ΡΠ΅ΠΊΡΡΠ²Π°ΡΡΡΡ. ΠΠ»Ρ ΠΊΠ°ΠΆΠ΄ΠΎΠ³ΠΎ ΠΈΠ· ΠΏΠΎΠΊΠ°Π·Π°ΡΠ΅Π»Π΅ΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Π±ΡΠ»Π° ΠΏΡΠ΅Π΄Π»ΠΎΠΆΠ΅Π½Π° ΠΌΠΎΠ΄Π΅Π»Ρ, ΠΎΠ΄Π½ΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎ ΠΎΠΏΠΈΡΡΠ²Π°ΡΡΠ°Ρ Π½Π°ΠΉΠ΄Π΅Π½Π½ΡΠ΅ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΡΡΠ΅ΠΊΡΡ. Π ΠΎΠ±Π»Π°ΡΡΡΡ ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½Π½ΡΡ Π»ΠΎΠΊΡΡΠΎΠ² Π±ΡΠ» ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½ ΠΏΠΎΠΈΡΠΊ Π³Π΅Π½ΠΎΠ²-ΠΊΠ°Π½Π΄ΠΈΠ΄Π°ΡΠΎΠ².
3. ΠΠ° ΠΌΠΎΠ΄Π΅Π»ΠΈ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΏΡΠΎΠ΄Π΅ΠΌΠΎΠ½ΡΡΡΠΈΡΠΎΠ²Π°Π½ΠΎ ΠΎΡΡΡΡΡΡΠ²ΠΈΠ΅ ΠΏΡΠ΅Π΄ΠΏΠΎΡΡΠ΅Π½ΠΈΡ Π·Π°ΠΏΠ°Ρ Π° Π°ΡΡΡΠ°Π»ΡΠ½ΠΎΠΉ ΡΠ°ΠΌΠΊΠΈ Ρ ΡΠ°ΠΌΡΠΎΠ² ΠΌΡΡΠ΅ΠΉ Ρ Ρ Π΅ΠΌΠΎΠ΄ΠΈΡΠΈΡΠΈΡΠΎΠΌ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ.
4. ΠΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ ΠΊΠ°ΠΊ ΠΎΡΠ½ΠΎΠ²Π½Π°Ρ, ΡΠ°ΠΊ ΠΈ Π΄ΠΎΠΏΠΎΠ»Π½ΠΈΡΠ΅Π»ΡΠ½Π°Ρ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½Π°Ρ ΡΠΈΡΡΠ΅ΠΌΠ° ΡΡΠ°ΡΡΠ²ΡΡΡ Π² Π΄Π΅ΡΠ΅ΠΊΡΠΈΠΈ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ. Π£ ΠΌΡΡΠ΅ΠΉ Ρ Ρ Π΅ΠΌΠΎΠ΄Π΅ΡΠΈΡΠΈΡΠΎΠΌ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ Π½Π° ΡΡΠΎΠ²Π½Π΅ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠ½ΠΎΠΉ ΡΠΊΠ°Π½ΠΈ Π²ΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ³Π°Π½Π° Π½Π°Π±Π»ΡΠ΄Π°Π»ΠΈ ΠΎΡΠ»ΠΈΡΠΈΡ ΠΏΠΎ Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΡ Fos-ΠΈΠΌΠΌΡΠ½ΠΎΡΠ΅Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ.
5. ΠΡΠ΅Π΄ΡΡΠ²Π»Π΅Π½ΠΈΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° (0.1%, 30 ΠΌΠΈΠ½) Π²ΡΠ·ΡΠ²Π°Π»ΠΎ Π΄ΠΎΡΡΠΎΠ²Π΅ΡΠ½ΠΎΠ΅ (Ρ<0.05) ΡΠ½ΠΈΠΆΠ΅Π½ΠΈΠ΅ ΡΡΠΎΠ²Π½Ρ ΡΠ΅ΡΡΠΎΡΡΠ΅ΡΠΎΠ½Π° Π² ΠΏΠ»Π°Π·ΠΌΠ΅ ΠΊΡΠΎΠ²ΠΈ Ρ ΠΌΡΡΠ΅ΠΉ Ρ Π½ΠΈΠ·ΠΊΠΈΠΌ ΠΏΠΎΡΠΎΠ³ΠΎΠΌ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π»Π΅ΡΡΡΠΈΠΌ ΡΡΠ΅ΡΠΎΠΈΠ΄Π°ΠΌ. ΠΡΠΈ ΡΡΠΎΠΌ ΡΡΠΎΠ²Π΅Π½Ρ ΠΊΠΎΡΡΠΈΠΊΠΎΡΡΠ΅ΡΠΎΠ½Π° Π΄Π»Ρ ΡΠ΅Ρ ΠΆΠ΅ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ ΠΎΡΡΠ°Π²Π°Π»ΡΡ Π½Π΅ΠΈΠ·ΠΌΠ΅Π½Π½ΡΠΌ, ΡΡΠΎ ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΠ΅Ρ ΠΎ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΌ Ρ Π°ΡΠ°ΠΊΡΠ΅ΡΠ΅ Π²Π»ΠΈΡΠ½ΠΈΡ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°.
6. Π‘ΠΎΠ²ΠΎΠΊΡΠΏΠ½ΠΎΡΡΡ ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΡ Π΄Π°Π½Π½ΡΡ : Π΄ΠΎΡΡΠΎΠ²Π΅ΡΠ½ΡΠΉ Π³ΠΎΡΠΌΠΎΠ½Π°Π»ΡΠ½ΡΠΉ ΠΎΡΠ²Π΅Ρ Π½Π° ΡΠΊΡΠΏΠΎΠ·ΠΈΡΠΈΡ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°, ΡΠ΅Π½Π΄Π΅Π½ΡΠΈΡ ΠΊ ΡΠ½ΠΈΠΆΠ΅Π½ΠΈΡ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ ΠΏΠΎΠ΄ Π²Π»ΠΈΡΠ½ΠΈΠ΅ΠΌ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π°, Π° ΡΠ°ΠΊΠΆΠ΅ Π²ΠΎΠ²Π»Π΅ΡΠ΅Π½ΠΈΠ΅ Π²ΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΠΎΡΠ³Π°Π½Π° Π² Π΄Π΅ΡΠ΅ΠΊΡΠΈΡ ΡΡΠΎΠ³ΠΎ ΠΎΠ΄ΠΎΡΠ°Π½ΡΠ°, ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΠ΅Ρ ΠΎΠ± ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΎΠ½Π½ΠΎΠΉ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΠΈ Π»Π΅ΡΡΡΠΈΡ ΡΡΠ΅ΡΠΎΠΈΠ΄ΠΎΠ² ΠΊΠ°ΠΊ Ρ Π΅ΠΌΠΎΡΠΈΠ³Π½Π°Π»ΠΎΠ² Π² ΡΠ΅Π³ΡΠ»ΡΡΠΈΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ.
ΠΠ°ΠΊΠ»ΡΡΠ΅Π½ΠΈΠ΅
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Π ΡΠ°Π±ΠΎΡΠ΅ Π±ΡΠ»Π° ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½Π° ΠΈΠ½ΡΠΎΡΠΌΠ°ΡΠΈΠΎΠ½Π½Π°Ρ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΡ ΠΌΠΎΠ»Π΅ΠΊΡΠ»Ρ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° Π² Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΎΠΉ ΠΊΠΎΠΌΠΌΡΠ½ΠΈΠΊΠ°ΡΠΈΠΈ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ Ρ ΠΏΡΠΈΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ΠΌ Π½Π΅ΡΠΊΠΎΠ»ΡΠΊΠΈΡ ΠΎΡΠ½ΠΎΠ²Π½ΡΡ ΠΏΠΎΠ΄Ρ ΠΎΠ΄ΠΎΠ². Π‘ ΠΎΠ΄Π½ΠΎΠΉ ΡΡΠΎΡΠΎΠ½Ρ, ΠΌΡ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π»ΠΈ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΡΡ ΠΌΠΎΠ΄Π΅Π»Ρ, ΠΎΡΠ½ΠΎΠ²Π°Π½Π½ΡΡ Π½Π° ΠΈΠ½Π±ΡΠ΅Π΄Π½ΡΡ Π»ΠΈΠ½ΠΈΡΡ ΠΌΡΡΠ΅ΠΉ Π‘ΠΠ ΠΈ NZB, ΠΊΠΎΠ½ΡΡΠ°ΡΡΠ½ΡΡ ΠΏΠΎ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ°ΠΌ, ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΈ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠΈ ' (Voznessenskaya et al, 1995). ΠΡΠ΄Π΅Π»ΡΠ½ΡΠΉ ΠΈΠ½ΡΠ΅ΡΠ΅Ρ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΠ΅Ρ ΡΡΠ°ΡΡΠΈΠ΅ ΡΠ°Π·Π»ΠΈΡΠ½ΡΡ .
I ΠΎΡΠ΄Π΅Π»ΠΎΠ² ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π°ΡΠΎΡΠ° Π² Π΄Π΅ΡΠ΅ΠΊΡΠΈΠΈ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° Ρ ΡΡΠΈΡ Π»ΠΈΠ½ΠΈΠΉ. Π‘ Π΄ΡΡΠ³ΠΎΠΉ ΡΡΠΎΡΠΎΠ½Ρ, Π±ΡΠ»ΠΎ ΠΈΠ·ΡΡΠ΅Π½ΠΎ Π²Π»ΠΈΡΠ½ΠΈΠ΅ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Π° Π½Π° ΡΡΠ΄ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΡΠ΅ΡΠΊΠΈΡ ΠΈ ΡΠΈΠ·ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΡ ΠΏΠΎΠΊΠ°Π·Π°ΡΠ΅Π»Π΅ΠΉ Ρ Π΄ΠΎΠΌΠΎΠ²ΠΎΠΉ ΠΌΡΡΠΈ. ΠΠ° ΠΎΡΠ½ΠΎΠ²Π΅ ΠΊΠ»Π°ΡΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ ΡΠΊΡΠ΅ΡΠΈΠ²Π°Π½ΠΈΡ ΠΌΠ΅ΠΆΠ΄Ρ Π»ΠΈΠ½ΠΈΡΠΌΠΈ CBA/J ΠΈ NZB/B1NJ Π±ΡΠ» ΠΏΡΠΎΠ²Π΅Π΄Π΅Π½ Π°Π½Π°Π»ΠΈΠ· Π°ΡΡΠΎΡΠΈΠ°ΡΠΈΠΉ ΠΌeΠΆΠ΄y^ ΡΠ΅Π½ΠΎΡΠΈΠΏΠ°ΠΌΠΈ (ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡΡ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ ΠΈ ΠΏΠΎΠΊΠ°Π·Π°ΡΠ΅Π»ΡΠΌΠΈ ΠΌΠ΅ΠΆΡΠ°ΠΌΡΠΎΠ²ΠΎΠΉ Π°Π³ΡΠ΅ΡΡΠΈΠΈ)^ ΠΈ |Π³Π΅Π½ΠΎΡΠΈΠΏΠΎΠΌ, ΠΎΠΏΡΠ΅Π΄Π΅Π»Π΅Π½Π½ΡΠΌ Ρ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π½ΠΈΠ΅ΠΌ ΠΠΠ ΠΌΠ°ΡΠΊΠ΅ΡΠΎΠ² (99 ΠΌΠΈΠΊΡΠΎΡΠ°ΡΠ΅Π»Π»ΠΈΡΠ½ΡΡ , 41 SNP), Π²ΡΡΠ²Π»Π΅Π½Ρ ΡΠΏΠΈΡΡΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΠ΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ ΠΌΠ΅ΠΆΠ΄Ρ ΠΌΠ°ΡΠΊΠ΅ΡΠ°ΠΌΠΈ, ΠΎΠΊΠ°Π·ΡΠ²Π°ΡΡΠΈΠ΅ Π²Π»ΠΈΡΠ½ΠΈΠ΅ Π½Π° ΠΏΡΠΎΡΠ²Π»Π΅Π½ΠΈΠ΅ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ°. ΠΡΠΈ Π°Π½Π°Π»ΠΈΠ·Π΅ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ° ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ Ρ Π²ΡΠ΅ΠΉ Π²ΡΠ±ΠΎΡΠΊΠΈ Π³ΠΈΠ±ΡΠΈΠ΄ΠΎΠ² F2 (ΠΏ=119) ΠΏΡΠΈ ΠΏΡΠΎΡΡΠΎΠΌ ΡΠΊΠ°Π½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ Π³Π΅Π½ΠΎΠΌΠ° Π² ΠΌΠΎΠ΄Π΅Π»ΠΈΡ ΠΎΠ΄ΠΈΠ½ΠΎΡΠ½ΡΠΌ QTL Π±ΡΠ»ΠΈ Π²ΡΡΠ²Π»Π΅Π½Ρ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΠ΅Π»ΡΠ½ΡΠ΅- (Ρ<0.63) Π»ΠΎΠΊΡΡΡ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΠΈΡΡΡΡΠΈΠ΅ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΡ ΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ Π½Π° 2 (rs3023694), 12(rs3684371) ΠΈ 17(rs3675244) Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ°Ρ . ΠΡΠΈ 3 Π»ΠΎΠΊΡΡΠ° Π°Π΄Π΄ΠΈΡΠΈΠ²Π½ΠΎ ΠΎΠ±ΡΡΡΠ½ΡΡΡ 25% ΡΠ΅Π½ΠΎΡΠΈΠΏΠΈΡΠ΅ΡΠΊΠΎΠΉ Π²Π°ΡΠΈΠ°Π±Π΅Π»ΡΠ½ΠΎΡΡΠΈ.' ΠΡΠΈ Π°Π½Π°Π»ΠΈΠ·Π΅, Π²ΡΠΏΠΎΠ»Π½Π΅Π½Π½ΠΎΠΌ Π΄Π»Ρ ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΡ Π³ΡΡΠΏΠΏ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ ^ Ρ Π²ΠΊΠ»ΡΡΠ΅Π½ΠΈΠ΅ΠΌ ^ ΠΊΠΎΠ²Π°ΡΠΈΠ°Ρ, ΠΏΠΎΠΌΠΈΠΌΠΎ, ΡΡΠ΄Π° ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΠ΅Π»ΡΠ½ΡΡ Π»ΠΎΠΊΡΡΠΎΠ². (ΡΠΌ. ΡΠ°Π±Π»ΠΈΡΡ 21), Π±ΡΠ» ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½ Π΄ΠΎΡΡΠΎΠ²Π΅ΡΠ½ΡΠΉ (Ρ<0.05) Π»ΠΎΠΊΡΡ Π½Π° 10 Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (D10Mit14) Ρ ΡΠ°ΠΌΡΠΎΠ², ΠΎΠ±ΡΡΡΠ½ΡΡΡΠΈΠΉ ΡΠ°ΡΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΠΏΠΎΡΠΎΠ³ΠΎΠ² Π²Π½ΡΡΡΠΈ Π³ΡΡΠΏΠΏ ΠΌΠ°Π»ΠΎΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΡΡ ΠΈ Π²ΡΡΠΎΠΊΠΎΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΡΡ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ , Π° ΡΠ°ΠΊΠΆΠ΅ Π΄ΠΎΡΡΠΎΠ²Π΅ΡΠ½ΠΎΠ΅ ΡΠΏΠΈΡΡΠ°ΡΠΈΡΠ΅ΡΠΊΠΎΠ΅ Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΠ΅ ΠΌΠ΅ΠΆΠ΄Ρ ΠΌΠ°ΡΠΊΠ΅ΡΠ°ΠΌΠΈ Π½Π° 2 Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (D2Mit266) ΠΈ X Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (rs3723498) Ρ ΡΠ°ΠΌΠΎΠΊ (LODfull Ρ<0.05, LODint Ρ<0.05). ΠΠ½Π°Π»ΠΈΠ· ΡΠ°ΡΡΠ΅ΠΏΠ»Π΅Π½ΠΈΡ ΠΏΠΎ ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΠΎΠΉ ΡΡΠ²ΡΡΠ²ΠΈΡΠ΅Π»ΡΠ½ΠΎΡΡΠΈΠΊ Π°Π½Π΄ΡΠΎΡΡΠ΅Π½ΠΎΠ½Ρ Π²ΠΎ Π²ΡΠΎΡΠΎΠΌ ΠΏΠΎΠΊΠΎΠ»Π΅Π½ΠΈΠΈ, Π° ΡΠ°ΠΊΠΆΠ΅ ΡΠ°Π·Π½ΠΈΡΠ° ΠΏΠΎΡΠΎΠ³ΠΎΠ² ΠΌΠ΅ΠΆΠ΄Ρ ΡΠ°ΠΌΡΠ°ΠΌΠΈ, Π½ΠΎ Π½Π΅ ΡΠ°ΠΌΠΊΠ°ΠΌΠΈ ΡΠ΅ΡΠΈΠΏΡΠΎΠΊΠ½ΡΡ ΠΊΡΠΎΡΡΠΎΠ² Π² Fi ΡΠΊΠ°Π·ΡΠ²Π°ΡΡ Π½Π° Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΠ΅ ΡΡΠ΅ΠΏΠ»Π΅Π½ΠΈΠ΅ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ° Ρ X-Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠΎΠΉ. ΠΠ΄Π½Π°ΠΊΠΎ ΠΌΡ Π½Π΅ ΠΎΠ±Π½Π°ΡΡΠΆΠΈΠ»ΠΈ Π»ΠΎΠΊΡΡΠΎΠ² Π½Π° X Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΎ 3 ΠΌΠ°ΡΠΊΠ΅ΡΠ°) Π² ΠΏΠΎΠ»Π½ΠΎΠΉ ΠΌΠ΅ΡΠ΅ ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΡΡΠΈΡ Π΄Π°Π½Π½ΡΠΉ ΠΏΡΠΈΠ·Π½Π°ΠΊ. Π’Π°ΠΊΠΈΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ, Π½Π°ΡΠΈ Π΄Π°Π½Π½ΡΠ΅ ΡΠΊΠ°Π·ΡΠ²Π°ΡΡ Π½Π° ΡΠ»ΠΎΠΆΠ½ΡΠΉ Ρ Π°ΡΠ°ΠΊΡΠ΅Ρ Π½Π°ΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΏΡΠΈΠ·Π½Π°ΠΊΠ° ΠΏΠΎΠ΄ Π²Π»ΠΈΡΠ½ΠΈΠ΅ΠΌ Π½Π΅ΡΠΊΠΎΠ»ΡΠΊΠΈΡ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΈΡ Π»ΠΎΠΊΡΡΠΎΠ² (ΡΠΌ. ΡΠ°Π±Π»ΠΈΡΡ 21). Π‘ΡΠΏΠ΅ΡΡΠ΅ΠΌΠ΅ΠΉΡΡΠ²ΠΎ Π³Π΅Π½ΠΎΠ² ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΡ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠΎΠ² ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ ΠΎΠ΄Π½ΠΎ ΠΈΠ· ΡΠ°ΠΌΡΡ ΠΎΠ±ΡΠΈΡΠ½ΡΡ Π² Π³Π΅Π½ΠΎΠΌΠ΅ (Buck and Axel, 1991). ΠΠ΄Π½ΠΈΠΌ^ΠΈΠ· Π²Π΅ΡΠΎΡΡΠ½ΡΡ ΠΎΠ±ΡΡΡΠ½Π΅Π½ΠΈΠΉ Π°Π½ΠΎΡΠΌΠΈΠΈ ΠΏΡΠ΅Π΄ΡΡΠ°Π²Π»ΡΠ΅ΡΡΡ ΡΡΡΠ΅ΡΡΠ²ΠΎΠ²Π°Π½ΠΈΠ΅ ΡΠΏΠ΅ΡΠΈΡΠΈΡΠ΅ΡΠΊΠΈΡ Π²Π°ΡΠΈΠ°Π½ΡΠΎΠ² Π³Π΅Π½ΠΎΠ² ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΡ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠΎΠ². ΠΠΎΠΈΡΠΊ Π² ΠΎΠ±Π»Π°ΡΡΡΡ Π½Π°ΠΉΠ΄Π΅Π½Π½ΡΡ Π½Π°ΠΌΠΈ ΠΏΡΠΈ ΡΠΊΠ°Π½ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ Π³Π΅Π½ΠΎΠΌΠ° Π»ΠΎΠΊΡΡΠΎΠ² ΠΎΠ±Π½Π°ΡΡΠΆΠΈΠ» ΠΎΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΠ΅ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΡ Π½Π° 2, 11, 10, 1 ΠΈ X Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ°Ρ . ΠΠ±ΠΎΠ½ΡΡΠ΅Π»ΡΠ½ΡΠ΅ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΡ Π² Π³Π΅Π½ΠΎΠΌΠ΅ ΠΌΡΡΠΈ ΡΠ°ΡΠΏΠΎΠ»ΠΎΠΆΠ΅Π½Ρ ΠΊΠ»Π°ΡΡΠ΅ΡΠ°ΠΌΠΈ (Zhang and Firestein, 2002), ΡΡΠΎ Π½Π΅ ΠΏΠΎΠ·Π²ΠΎΠ»ΡΠ΅Ρ ΡΠΎΠΎΡΠ½Π΅ΡΡΠΈ ΠΏΠΎΠ»ΡΡΠ΅Π½Π½ΡΠ΅ Π½Π°ΠΌΠΈ ΡΡΠ΅ΠΏΠ»Π΅Π½ΠΈΡ Ρ ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΠΌΠΈ Π³Π΅Π½Π°ΠΌΠΈ. Π’Π°ΠΊΠΆΠ΅ Π² ΠΎΠ±Π»Π°ΡΡΡΡ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΠ΅Π»ΡΠ½ΡΡ Π»ΠΎΠΊΡΡΠΎΠ² Π½Π° Π± ΠΈ X Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ°Ρ Π»ΠΎΠΊΠ°Π»ΠΈΠ·ΠΎΠ²Π°Π½Ρ Π³Π΅Π½Ρ Π²ΠΎΠΌΠ΅ΡΠΎΠ½Π°Π·Π°Π»ΡΠ½ΡΡ ΡΠ΅ΡΠ΅ΠΏΡΠΎΡΠΎΠ². ΠΠΌΠ΅Π΅ΡΡΡ Π΄ΠΎΠ²ΠΎΠ»ΡΠ½ΠΎ ΠΌΠ½ΠΎΠ³ΠΎ Π»ΠΈΡΠ΅ΡΠ°ΡΡΡΠ½ΡΡ ΠΈΡΡΠΎΡΠ½ΠΈΠΊΠΎΠ², ΡΠ²ΡΠ·ΡΠ²Π°ΡΡΠΈΡ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ ΠΎΡΠ΄Π΅Π»ΡΠ½ΡΡ Π³Π΅Π½ΠΎΠ² ΠΌΠ»Π΅ΠΊΠΎΠΏΠΈΡΠ°ΡΡΠΈΡ Ρ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΡΠΌ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΠ΅ΠΌ (Π½Π°ΠΏΡΠΈΠΌΠ΅Ρ, ΠΎΠ±Π·ΠΎΡ Maxson and Canastar, 2003). ΠΠ΄Π½Π°ΠΊΠΎ ΡΠ°Π±ΠΎΡ Π² ΠΊΠΎΡΠΎΡΡΡ Π±Ρ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π»ΠΈΡΡ ΠΏΡΠ΅ΠΈΠΌΡΡΠ΅ΡΡΠ²Π° QTL-Π°Π½Π°Π»ΠΈΠ·Π° ΠΈ ΡΠΊΠ°Π½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ Π²ΡΠ΅Π³ΠΎ Π³Π΅Π½ΠΎΠΌΠ° Π½Π΅ ΡΠ°ΠΊ ΠΌΠ½ΠΎΠ³ΠΎ (Π²ΡΠ΅Π³ΠΎ 2), ΠΏΡΠΈΡΠ΅ΠΌ ΠΈΡΠΏΠΎΠ»ΡΠ·ΠΎΠ²Π°Π»ΠΈΡΡ ΠΎΡΠ»ΠΈΡΠ½ΡΠ΅ ΠΎΡ Π½Π°ΡΠΈΡ ΠΌΠ΅ΡΠΎΠ΄Ρ ΠΎΡΠ΅Π½ΠΊΠΈ Π°Π³ΡΠ΅ΡΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΠΏΠΎΠ²Π΅Π΄Π΅Π½ΠΈΡ ΠΈ Π΄ΡΡΠ³ΠΈΠ΅ ΠΈΠ½Π±ΡΠ΅Π΄Π½ΡΠ΅ Π»ΠΈΠ½ΠΈΠΈ. ΠΠ±Π½Π°ΡΡΠΆΠ΅Π½Π½ΡΠ΅ Π² Π½Π°ΡΠ΅ΠΌ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΠΈ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠΈΡΠ΅Π»ΡΠ½ΡΠ΅ (Ρ<0.63) Π»ΠΎΠΊΡΡΡ Π½Π° 8 Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (D8Mit205, rs4227112) ΠΈ. Π½Π° 10 Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (rs3693721) ΠΏΠΎΠ΄ΡΠ²Π΅ΡΠΆΠ΄Π°ΡΡ Π·Π½Π°ΡΠΈΠΌΠΎΡΡΡ ΡΠ°Π½Π΅Π΅ ΠΎΠΏΠΈΡΠ°Π½Π½ΡΡ QTL, ΡΠΎΠΎΡΠ²Π΅ΡΡΡΠ²Π΅Π½Π½ΠΎ. Imab2 Π½Π° 8 Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (D8Mit8, Roubertoux et al- 2005) ΠΈ Aggrl Π½Π° 10 Ρ ΡΠΎΠΌΠΎΡΠΎΠΌΠ΅ (D10Mit267, D10Mit103, Brodkin et al, 2002) (ΡΠΌ: ΡΠ°Π±Π»ΠΈΡΡ 22).
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