ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ Π½ΠΎΠ²ΠΎΠΉ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΡΠΈΠΏΠ° ΠΈΠ· ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΎΠΊ ΠΊΠΎΡΡΠΈΡΡΠΎΠΉ ΡΡΠ±Ρ ΠΡΡΠ½ (Misgurnus fossilis L.)
Π Π΅ΡΠ΅Π½ΠΈΠ΅ Π²ΠΎΠΏΡΠΎΡΠ° ΠΎ ΡΠΎΠ»ΠΈ ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½Π½ΠΎΠΉ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ ΡΡΠ΅Π±ΡΠ΅Ρ Π΄Π΅ΡΠ°Π»ΡΠ½ΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π° ΡΡΡΠΎΠ΅Π½ΠΈΡ ΡΠ΅ΡΠΌΠ΅Π½ΡΠ°, ΠΈΠ΄Π΅Π½ΡΠΈΡΠΈΠΊΠ°ΡΠΈΠΈ ΠΊΠ°ΡΠ°Π»ΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ ΡΡΠ±ΡΠ΅Π΄ΠΈΠ½ΠΈΡΡ ΠΈ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ° Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΠΌΠΈ ΠΠΠ-ΡΡΠ±ΡΡΡΠ°ΡΠ°ΠΌΠΈ. Π€ΠΈΠ·ΠΈΠΊΠΎ-Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠΉ ΠΈ ΡΠ½Π·ΠΈΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π‘, ΠΌΠΎΠΆΠ΅Ρ Π²ΡΡΠ²ΠΈΡΡ Π½ΠΎΠ²ΡΠ΅ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΡ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠΉ ΠΠΠ ΠΈ ΠΈΡ ΡΠΏΠ΅ΡΠΈΡΠΈΠΊΡ Π² ΡΠ°Π½Π½Π΅ΠΌ ΡΠ°Π·Π²ΠΈΡΠΈΠΈ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ . Π‘ ΠΏΠΎΠΌΠΎΡΡΡ Π½Π°ΡΠΈΠ²Π½ΠΎΠ³ΠΎ… 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‘ΠΏΠΈΡΠΎΠΊ ΡΠΎΠΊΡΠ°ΡΠ΅Π½ΠΈΠΉ
- 1. ΠΠΠΠΠΠΠΠ
- 2. ΠΠΠΠΠ ΠΠΠ’ΠΠ ΠΠ’Π£Π Π«
- 2. 1. ΠΠΎΠ²ΡΠ΅ΠΆΠ΄Π΅Π½ΠΈΡ Π² ΠΠΠ ΠΈ ΡΠ°ΠΊΡΠΎΡΡ, ΠΈΡ Π²ΡΠ·ΡΠ²Π°ΡΡΠΈΠ΅
- 2. 2. ΠΠ΅Ρ
Π°Π½ΠΈΠ·ΠΌΡ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΠΠ
- 2. 2. 1. ΠΠΊΡΡΠΈΠ·ΠΈΠΎΠ½Π½Π°Ρ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΡ ΠΎΡΠ½ΠΎΠ²Π°Π½ΠΈΡ
- 2. 2. 2. ΠΠΊΡΡΠΈΠ·ΠΈΠΎΠ½Π½Π°Ρ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΡ Π½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄Π½ΠΎΠ³ΠΎ ΠΎΡΡΠ°ΡΠΊΠ°
- 2. 3. Π Π΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΡΠ΅ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ
- 2. 3. 1. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π° Ρ
- 2. 3. 2. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ 5 ΠΈ Π³
- 2. 3. 3. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π° Π°
- 2. 3. 4. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π°Π‘
- 2. 4. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π² ΡΠ°Π½Π½Π΅ΠΌ ΡΠ°Π·Π²ΠΈΡΠΈΠΈ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ
- 2. 5. ΠΠΊΠΈΡΠ»ΠΈΡΠ΅Π»ΡΠ½ΡΠΉ ΡΡΡΠ΅ΡΡ ΠΈ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΡ ΠΠΠ
- 3. ΠΠΠ’ΠΠ ΠΠΠΠ« Π ΠΠΠ’ΠΠΠ« ΠΠ‘Π‘ΠΠΠΠΠΠΠΠΠ―
- 3. 1. ΠΡΡΠΎΡΠ½ΠΈΠΊ ΠΈ ΠΊΠ²Π°Π»ΠΈΡΠΈΠΊΠ°ΡΠΈΡ ΠΎΡΠ½ΠΎΠ²Π½ΡΡ ΡΠ΅Π°ΠΊΡΠΈΠ²ΠΎΠ² ΠΈ ΡΠ΅ΡΠΌΠ΅Π½ΡΠΎΠ²
- 3. 2. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ Π±ΠΈΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΎΠ³ΠΎ ΠΌΠ°ΡΠ΅ΡΠΈΠ°Π»Π°
- 3. 3. Π€ΡΠ°ΠΊΡΠΈΠΎΠ½ΠΈΡΠΎΠ²Π°Π½ΠΈΠ΅ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·
- 3. 3. 1. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΠΏΡΠ΅ΠΏΠ°ΡΠ°ΡΠΎΠ² ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΡΡ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· Π΄Π»Ρ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ Π΄ΠΈΠ½Π°ΠΌΠΈΠΊΠΈ ΠΈΡ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ Π² ΡΠΌΠ±ΡΠΈΠΎΠ³Π΅Π½Π΅Π·Π΅
- 3. 3. 2. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΠΎΡΠΈΡΠ΅Π½Π½ΡΡ ΠΏΡΠ΅ΠΏΠ°ΡΠ°ΡΠΎΠ² ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· Ρ ΠΈ Π‘, ΠΈΠ· ΠΈΠΊΡΡ Π²ΡΡΠ½Π° Π΄Π»Ρ ΡΠ΅Π΄ΠΈΠΌΠ΅Π½ΡΠ°ΡΠΈΠΎΠ½Π½ΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π°
- 3. 4. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΠΠΠ-ΡΡΠ±ΡΡΡΠ°ΡΠΎΠ²
- 3. 4. 1. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΡΡΠ±ΡΡΡΠ°ΡΠΎΠ² ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Ρ, ΠΌΠ΅ΡΠ΅Π½Π½ΡΡ 3Π Ρ Π'-ΠΊΠΎΠ½ΡΠ°
- 3. 4. 2. ΠΠΎΠ»ΡΡΠ΅Π½ΠΈΠ΅ ΡΡΠ±ΡΡΡΠ°ΡΠΎΠ² ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Ρ, ΠΌΠ΅ΡΠ΅Π½Π½ΡΡ 32Π Ρ 5'-ΠΊΠΎΠ½ΡΠ°.8 -—""^3.5. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ ΡΠ΅ΡΠΌΠ΅Π½ΡΠΎΠ²
- 3. 5. 1. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·
- 3. 5. 2. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ 3'-«5' ΠΈ 5'→3'-ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Ρ
- 3. 6. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΠΏΡΠΎΡΠ΅ΡΡΠΈΠ²Π½ΠΎΡΡΠΈ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·
- 3. 7. ΠΡΡΠ³ΠΈΠ΅ ΠΌΠ΅ΡΠΎΠ΄Ρ
- 3. 7. 1. ΠΠ»Π΅ΠΊΡΡΠΎΡΠΎΡΠ΅ΡΠΈΡΠ΅ΡΠΊΠΎΠ΅ ΡΠ°Π·Π΄Π΅Π»Π΅Π½ΠΈΠ΅ Π±Π΅Π»ΠΊΠΎΠ² ΠΈ Π½ΡΠΊΠ»Π΅ΠΈΠ½ΠΎΠ²ΡΡ ΠΊΠΈΡΠ»ΠΎΡ
- 3. 7. 2. ΠΠ»Π΅ΠΊΡΡΠΎΡΠΎΡΠ΅Π· ΡΠ΅ΡΠΌΠ΅Π½ΡΠΎΠ² Π² Π½Π΅Π΄Π΅Π½Π°ΡΡΡΠΈΡΡΡΡΠ΅ΠΌ ΠΏΠΎΠ»ΠΈΠ°ΠΊΡΠΈΠ»Π°ΠΌΠΈΠ΄Π½ΠΎΠΌ Π³Π΅Π»Π΅
- 3. 7. 3. ΠΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ ΠΊΠΎΠ½ΡΠ΅Π½ΡΡΠ°ΡΠΈΠΈ Π±Π΅Π»ΠΊΠ° ΠΈ ΡΠΎΠ»ΠΈ
- 4. 1. ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΡΡ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· ΠΈΠ· Π·ΡΠ΅Π»ΡΡ ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΎΠΊ Π²ΡΡΠ½Π°
- 4. 2. ΠΠΎΠ»ΠΈΠΏΠ΅ΠΏΡΠΈΠ΄Π½ΡΠΉ ΡΠΎΡΡΠ°Π² ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ (
- 4. 3. ΠΠΎΠ»ΠΈΠΏΠ΅ΠΏΡΠΈΠ΄Π½ΡΠΉ ΡΠΎΡΡΠ°Π² ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π‘
- 4. 4. ΠΠ½Π·ΠΈΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠ°Ρ Ρ
Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΡΡΠΈΠΊΠ° ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π‘
- 4. 4. 1. ΠΠ°ΡΡΠΈΡΠ½ΡΠ΅ ΡΠ²ΠΎΠΉΡΡΠ²Π°
- 4. 4. 2. ΠΡΠΎΡΠ΅ΡΡΠΈΠ²Π½ΠΎΡΡΡ.®
- 4. 4. 3. 3'→5'-ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Π½Π°Ρ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ. ' '
- 4. 4. 4. ΠΠ½Π³ΠΈΠ±ΠΈΡΠΎΡΠ½ΡΠΉ Π°Π½Π°Π»ΠΈΠ·
- 4. 5. ΠΠ·ΠΌΠ΅Π½Π΅Π½ΠΈΠ΅ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· Ρ ΠΈ Π‘ Π² ΡΠ°Π½Π½Π΅ΠΌ ΡΠ°Π·Π²ΠΈΡΠΈΠΈ Π²ΡΡΠ½Π°
- 4. 6. ΠΠ»ΠΈΡΠ½ΠΈΠ΅ ΠΎΠΊΠΈΡΠ»ΠΈΡΠ΅Π»ΡΠ½ΠΎΠ³ΠΎ ΡΡΡΠ΅ΡΡΠ° Π½Π° ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π² ΡΠΌΠ±ΡΠΈΠΎΠ³Π΅Π½Π΅Π·Π΅ Π²ΡΡΠ½Π°
- 5. 1. Π Π΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΡΠ΅ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π² ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠΌ ΡΠ°Π·Π²ΠΈΡΠΈΠΈ
- 5. 2. Π₯Π°ΡΠ°ΠΊΡΠ΅ΡΠΈΡΡΠΈΠΊΠ° ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π‘ ΠΈΠ· Π·ΡΠ΅Π»ΡΡ ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΎΠΊ Π²ΡΡΠ½Π°
ΠΡΠ΄Π΅Π»Π΅Π½ΠΈΠ΅ Π½ΠΎΠ²ΠΎΠΉ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΡΠΈΠΏΠ° ΠΈΠ· ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΎΠΊ ΠΊΠΎΡΡΠΈΡΡΠΎΠΉ ΡΡΠ±Ρ ΠΡΡΠ½ (Misgurnus fossilis L.) (ΡΠ΅ΡΠ΅ΡΠ°Ρ, ΠΊΡΡΡΠΎΠ²Π°Ρ, Π΄ΠΈΠΏΠ»ΠΎΠΌ, ΠΊΠΎΠ½ΡΡΠΎΠ»ΡΠ½Π°Ρ)
Π Π΅ΠΏΠ°ΡΠ°ΡΠΈΡ ΠΠΠ ΡΠ²Π»ΡΠ΅ΡΡΡ Π²Π°ΠΆΠ½Π΅ΠΉΡΠ΅ΠΉ Π³Π΅Π½Π΅ΡΠΈΡΠ΅ΡΠΊΠΎΠΉ ΡΡΠ½ΠΊΡΠΈΠ΅ΠΉ, Π½Π°ΠΏΡΠ°Π²Π»Π΅Π½Π½ΠΎΠΉ Π½Π° ΡΠΎΡ ΡΠ°Π½Π΅Π½ΠΈΠ΅ ΠΈΠ½ΡΠ°ΠΊΡΠ½ΠΎΠΉ ΡΡΡΡΠΊΡΡΡΡ ΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠ³ΠΎ Π³Π΅Π½ΠΎΠΌΠ°. Π Π°Π·Π»ΠΈΡΠ½ΡΠ΅ ΡΠΈΠ·ΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΠΈ Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠ΅ ΡΠ°ΠΊΡΠΎΡΡ Π²ΡΠ·ΡΠ²Π°ΡΡ ΡΠ°Π·Π»ΠΈΡΠ½ΡΠ΅ ΠΏΠΎΠ²ΡΠ΅ΠΆΠ΄Π΅Π½ΠΈΡ Π² ΠΠΠ, ΡΡΠΎ ΠΎΠΏΡΠ΅Π΄Π΅Π»ΡΠ΅Ρ ΠΌΠ½ΠΎΠ³ΠΎΠΎΠ±ΡΠ°Π·ΠΈΠ΅ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠΎΠ² ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΡΡΠΈΡ ΠΏΠΎΠ²ΡΠ΅ΠΆΠ΄Π΅Π½ΠΈΠΉ. ΠΡΠΎΠ±Π»Π΅ΠΌΠ° ΠΊΠΎΡΡΠ΅ΠΊΡΠΈΠΈ ΡΡΡΡΠΊΡΡΡΠ½ΡΡ Π°Π½ΠΎΠΌΠ°Π»ΠΈΠΉ, Π²ΠΎΠ·Π½ΠΈΠΊΠ°ΡΡΠΈΡ Π² ΠΠΠ, ΠΎΡΠΎΠ±Π΅Π½Π½ΠΎ Π°ΠΊΡΡΠ°Π»ΡΠ½Π° Π½Π° ΡΠ°Π½Π½ΠΈΡ ΡΡΠ°ΠΏΠ°Ρ ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Π»ΡΠ½ΠΎΠ³ΠΎ ΡΠ°Π·Π²ΠΈΡΠΈΡ, ΡΠ°ΠΊ ΠΊΠ°ΠΊ Π΄Π΅ΡΠ΅ΠΊΡ Π² ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΠΎΠΉ ΡΠΈΡΡΠ΅ΠΌΠ΅ ΠΊΠ»Π΅ΡΠΎΠΊ ΡΠΌΠ±ΡΠΈΠΎΠ½Π° Π² ΡΡΠ»ΠΎΠ²ΠΈΡΡ ΠΈΠ½ΡΠ΅Π½ΡΠΈΠ²Π½ΠΎΠΉ ΠΏΡΠΎΠ»ΠΈΡΠ΅ΡΠ°ΡΠΈΠΈ ΠΏΡΠΈΠ²ΠΎΠ΄ΠΈΡ ΠΊ ΠΌΠ½ΠΎΠ³ΠΎΠΊΡΠ°ΡΠ½ΠΎΠΌΡ ΠΏΠΎΠ²ΡΠΎΡΠ΅Π½ΠΈΡ ΠΎΡΠΈΠ±ΠΊΠΈ. Π Π΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΡΠΉ ΡΠΈΠ½ΡΠ΅Π· ΠΠΠ Π² Ρ ΠΎΠ΄Π΅ ΠΏΠ΅ΡΠ²ΡΡ Π΄Π΅Π»Π΅Π½ΠΈΠΉ ΠΊΠ»Π΅ΡΠΎΠΊ ΡΠΌΠ±ΡΠΈΠΎΠ½Π° ΠΎΠ±Π΅ΡΠΏΠ΅ΡΠΈΠ²Π°Π΅ΡΡΡ Π·Π°ΠΏΠ°ΡΠ΅Π½Π½ΡΠΌΠΈ Π² Π·ΡΠ΅Π»ΠΎΠΌ ΠΎΠΎΡΠΈΡΠ΅ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΡΠΌΠΈ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π°ΠΌΠΈ. ΠΠΈΠ·ΠΊΠΎΠ΅ ΡΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ? Π² Π·ΡΠ΅Π»ΡΡ ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΊΠ°Ρ Π²ΡΡΠ½Π° Π² ΡΡΠ°Π²Π½Π΅Π½ΠΈΠΈ Ρ Π΅Π΅ ΡΠΎΠ΄Π΅ΡΠΆΠ°Π½ΠΈΠ΅ΠΌ Π² ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΊΠ°Ρ Π΄ΡΡΠ³ΠΈΡ Π²ΠΈΠ΄ΠΎΠ² (Mikhailov and Gulyamov, 1983; Carre et al., 1981, Matsumoto et al., 1994) ΡΡΠ°Π²ΠΈΡ Π²ΠΎΠΏΡΠΎΡ ΠΎ Π²ΠΎΠ·ΠΌΠΎΠΆΠ½ΠΎΠΌ ΡΡΠ°ΡΡΠΈΠΈ Π΄ΡΡΠ³ΠΈΡ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· Π² ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΏΠΎΠ²ΡΠ΅ΠΆΠ΄Π΅Π½ΠΈΠΉ ΠΠΠ Π½Π° ΡΠ°Π½Π½ΠΈΡ ΡΡΠ°ΠΏΠ°Ρ ΡΠΌΠ±ΡΠΈΠΎΠ³Π΅Π½Π΅Π·Π°.
Π‘ΠΎΠ³Π»Π°ΡΠ½ΠΎ ΡΠΎΠ²ΡΠ΅ΠΌΠ΅Π½Π½ΠΎΠΉ ΠΊΠ»Π°ΡΡΠΈΡΠΈΠΊΠ°ΡΠΈΠΈ, ΡΡΡΠ΅ΡΡΠ²ΡΡΡ ΠΏΡΡΡ ΡΠΈΠΏΠΎΠ² ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· ΡΡΠΊΠ°ΡΠΈΠΎΡ: Π°, ?, Ρ, 8 ΠΈ s (Burgers et al., 1990), ΠΊΠΎΡΠΎΡΡΠ΅ ΡΡΠ°ΡΡΠ²ΡΡΡ Π² ΡΠ΅ΠΏΠ»ΠΈΠΊΠ°ΡΠΈΠΈ ΠΈ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΠΠ. ΠΠ΅Π΄Π°Π²Π½ΠΎ ΠΎΠΏΡΠ±Π»ΠΈΠΊΠΎΠ²Π°Π½Ρ Π΄Π°Π½Π½ΡΠ΅ ΠΎ Π½ΠΎΠ²ΠΎΠΉ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π΅, Π½Π°Π·Π²Π°Π½Π½ΠΎΠΉ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·ΠΎΠΉ Π‘, Π²ΡΠ΄Π΅Π»Π΅Π½Π½ΠΎΠΉ ΠΈΠ· ΡΠΈΠΌΡΡΠ° ΡΠ΅Π»Π΅Π½ΠΊΠ° (Bialek and Grosse, 1993), Π·ΡΠ΅Π»ΡΡ ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΎΠΊ Π²ΡΡΠ½Π° (Π¨Π°ΡΠΎΠ²Π° ΠΈ Π΄Ρ., 1994) ΠΈ ΠΊΠ»Π΅ΡΠΎΠΊ Π΄ΡΠΎΠΆΠΆΠ΅ΠΉ (Nelson et al., 1996). ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π° Π‘, ΡΡΠ°ΡΡΠ²ΡΠ΅Ρ, ΠΏΠΎ-Π²ΠΈΠ΄ΠΈΠΌΠΎΠΌΡ, Π² ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΠΠ. ΠΡΠ»ΠΈ ΠΎ ΡΡΡΡΠΊΡΡΡΠ΅, ΡΠΈΠ·ΠΈΠΊΠΎ-Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΡ ΠΈ ΡΠ½Π·ΠΈΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΡ ΡΠ²ΠΎΠΉΡΡΠ²Π°Ρ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· Π°, ?, Ρ, 5 ΠΈ Π² ΡΡΡΠ΅ΡΡΠ²ΡΡΡ ΠΌΠ½ΠΎΠ³ΠΎΡΠΈΡΠ»Π΅Π½Π½ΡΠ΅ Π΄Π°Π½Π½ΡΠ΅, ΡΠΎ ΠΎ Π½ΠΎΠ²ΠΎΠΌ ΡΠ΅ΡΠΌΠ΅Π½ΡΠ΅ ΠΈΠ·Π²Π΅ΡΡΠ½ΠΎ ΠΊΡΠ°ΠΉΠ½Π΅ ΠΌΠ°Π»ΠΎ.
Π Π΅ΡΠ΅Π½ΠΈΠ΅ Π²ΠΎΠΏΡΠΎΡΠ° ΠΎ ΡΠΎΠ»ΠΈ ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½Π½ΠΎΠΉ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ ΡΡΠ΅Π±ΡΠ΅Ρ Π΄Π΅ΡΠ°Π»ΡΠ½ΠΎΠ³ΠΎ Π°Π½Π°Π»ΠΈΠ·Π° ΡΡΡΠΎΠ΅Π½ΠΈΡ ΡΠ΅ΡΠΌΠ΅Π½ΡΠ°, ΠΈΠ΄Π΅Π½ΡΠΈΡΠΈΠΊΠ°ΡΠΈΠΈ ΠΊΠ°ΡΠ°Π»ΠΈΡΠΈΡΠ΅ΡΠΊΠΎΠΉ ΡΡΠ±ΡΠ΅Π΄ΠΈΠ½ΠΈΡΡ ΠΈ ΠΈΡΡΠ»Π΅Π΄ΠΎΠ²Π°Π½ΠΈΡ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΠ° Π²Π·Π°ΠΈΠΌΠΎΠ΄Π΅ΠΉΡΡΠ²ΠΈΡ Ρ ΠΌΠΎΠ΄Π΅Π»ΡΠ½ΡΠΌΠΈ ΠΠΠ-ΡΡΠ±ΡΡΡΠ°ΡΠ°ΠΌΠΈ. Π€ΠΈΠ·ΠΈΠΊΠΎ-Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠΉ ΠΈ ΡΠ½Π·ΠΈΠΌΠΎΠ»ΠΎΠ³ΠΈΡΠ΅ΡΠΊΠΈΠΉ Π°Π½Π°Π»ΠΈΠ· ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π‘, ΠΌΠΎΠΆΠ΅Ρ Π²ΡΡΠ²ΠΈΡΡ Π½ΠΎΠ²ΡΠ΅ ΠΌΠ΅Ρ Π°Π½ΠΈΠ·ΠΌΡ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠΈ ΠΊΠ»Π΅ΡΠΎΡΠ½ΠΎΠΉ ΠΠΠ ΠΈ ΠΈΡ ΡΠΏΠ΅ΡΠΈΡΠΈΠΊΡ Π² ΡΠ°Π½Π½Π΅ΠΌ ΡΠ°Π·Π²ΠΈΡΠΈΠΈ ΠΆΠΈΠ²ΠΎΡΠ½ΡΡ .
2. ΠΠΠΠΠ ΠΠΠ’ΠΠ ΠΠ’Π£Π Π«.
Π²ΡΠ²ΠΎΠ΄Ρ.
1. Π ΡΠΉΡΠ΅ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΠΈ ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Ρ Π²ΡΡΠ½Π° ΠΎΠ±Π½Π°ΡΡΠΆΠ΅Π½Π° ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π° Π½ΠΎΠ²ΠΎΠ³ΠΎ ΡΠΈΠΏΠ°, ΠΎΠ±ΠΎΠ·Π½Π°ΡΠ΅Π½Π½Π°Ρ ΠΊΠ°ΠΊ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π°? (Π΄Π·Π΅ΡΠ°), ΠΊΠΎΡΠΎΡΠ°Ρ ΠΎΡΠ»ΠΈΡΠ°Π΅ΡΡΡ ΠΏΠΎ ΡΠΈΠ·ΠΈΠΊΠΎ-Ρ ΠΈΠΌΠΈΡΠ΅ΡΠΊΠΈΠΌ ΠΈ ΡΠ½Π·ΠΈΠΌΠ°ΡΠΈΡΠ΅ΡΠΊΠΈΠΌ ΡΠ²ΠΎΠΉΡΡΠ²Π°ΠΌ ΠΎΡ ΠΎΠΏΠΈΡΠ°Π½Π½ΡΡ ΡΠ°Π½Π΅Π΅ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· Π°, Π , Ρ, 6 ΠΈ Π΅ ΡΡΠΊΠ°ΡΠΈΠΎΡ.
2. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π°? Π²ΡΡΠ½Π° ΠΎΡΡΡΠ΅ΡΡΠ²Π»ΡΠ΅Ρ Π΄ΠΈΡΡΡΠΈΠ±ΡΡΠΈΠ²Π½ΡΠΉ ΡΠΈΠ½ΡΠ΅Π· ΠΠΠ Π½Π° ΠΌΠ°ΡΡΠΈΡΠ΅ ΡΠΎ[Ρ (Ρ1Π)*Ρ1Π’1ΠΎ ΠΊΠ°ΠΊ Π² ΠΎΡΡΡΡΡΡΠ²ΠΈΠ΅, ΡΠ°ΠΊ ΠΈ Π² ΠΏΡΠΈΡΡΡΡΡΠ²ΠΈΠΈ ΡΠ΅ΠΏΠ»ΠΈΠΊΠ°ΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΡΠ°ΠΊΡΠΎΡΠ° Π Π‘ΠΠ.
3. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π° Π²ΡΡΠ½Π° ΠΎΠ±Π»Π°Π΄Π°Π΅Ρ Π²ΡΡΠΎΠΊΠΎΠΉ 3'->5'-ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Π½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡΡ. ΠΡΠ½ΠΎΡΠ΅Π½ΠΈΠ΅ 3'->5'-ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Π½ΠΎΠΉ ΠΊ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ Π² 20 ΡΠ°Π· Π²ΡΡΠ΅ ΠΏΠΎΠ΄ΠΎΠ±Π½ΠΎΠ³ΠΎ ΡΠΎΠΎΡΠ½ΠΎΡΠ΅Π½ΠΈΡ Ρ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π΅ ΡΠ΅Π»ΠΎΠ²Π΅ΠΊΠ°.
4. 3'-«5'-ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Π° ΡΠ΅ΡΠΌΠ΅Π½ΡΠ° Π³ΠΈΠ΄ΡΠΎΠ»ΠΈΠ·ΡΠ΅Ρ ΠΏΡΠ΅Π΄ΠΏΠΎΡΡΠΈΡΠ΅Π»ΡΠ½ΠΎ ΠΎΠ΄Π½ΠΎΠ½ΠΈΡΠ΅Π²ΡΠ΅ ΠΎΠ»ΠΈΠ³ΠΎΠ½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄Ρ ΠΈ Π½Π΅ΡΠΏΠ°ΡΠ΅Π½Π½ΡΠ΅ Π'-ΠΊΠΎΠ½ΡΠ΅Π²ΡΠ΅ Π½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄Ρ Π΄Π²ΡΡ Π½ΠΈΡΠ΅Π²ΡΡ ΡΡΠ±ΡΡΡΠ°ΡΠΎΠ². ΠΠΈΠ΄ΡΠΎΠ»ΠΈΠ· ΠΎΠ΄Π½ΠΎΠ½ΠΈΡΠ΅Π²ΡΡ ΠΎΠ»ΠΈΠ³ΠΎΠΈ ΠΏΠΎΠ»ΠΈΠ½ΡΠΊΠ»Π΅ΠΎΡΠΈΠ΄ΠΎΠ² ΠΎΡΡΡΠ΅ΡΡΠ²Π»ΡΠ΅ΡΡΡ Π΄ΠΈΡΡΡΠΈΠ±ΡΡΠΈΠ²Π½ΡΠΌ ΠΎΠ±ΡΠ°Π·ΠΎΠΌ.
5. ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π½Π°Ρ ΠΈ 3'-«5'-ΡΠΊΠ·ΠΎΠ½ΡΠΊΠ»Π΅Π°Π·Π½Π°Ρ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ ΡΠΎΠΎΡΠ°ΠΆΠ΄Π°ΡΡΡΡ ΡΠΎΠ²ΠΌΠ΅ΡΡΠ½ΠΎ ΠΏΡΠΈ ΡΠ»ΡΡΡΠ°ΡΠ΅Π½ΡΡΠΈΡΡΠ³ΠΈΡΠΎΠ²Π°Π½ΠΈΠΈ Π² Π³ΡΠ°Π΄ΠΈΠ΅Π½ΡΠ΅ 15−30%-Π½ΠΎΠ³ΠΎ Π³Π»ΠΈΡΠ΅ΡΠΈΠ½Π° Ρ ΠΊΠΎΡΡΡΠΈΡΠΈΠ΅Π½ΡΠΎΠΌ ΡΠ΅Π΄ΠΈΠΌΠ΅Π½ΡΠ°ΡΠΈΠΈ 6 Π, ΠΏΡΠΈ ΡΡΠΎΠΌ ΠΎΠ±Π΅ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΠΈ ΠΊΠΎΡΡΠ΅Π»ΠΈΡΡΡΡ Ρ ΡΠ°ΡΠΏΡΠ΅Π΄Π΅Π»Π΅Π½ΠΈΠ΅ΠΌ ΠΏΠΎΠ»ΠΈΠΏΠ΅ΠΏΡΠΈΠ΄Π° 68 ΠΊΠΠ Π² Π³ΡΠ°Π΄ΠΈΠ΅Π½ΡΠ΅.
6. Π‘ ΠΏΠΎΠΌΠΎΡΡΡ Π½Π°ΡΠΈΠ²Π½ΠΎΠ³ΠΎ ΡΠ»Π΅ΠΊΡΡΠΎΡΠΎΡΠ΅Π·Π° Ρ ΠΏΠΎΡΠ»Π΅Π΄ΡΡΡΠΈΠΌ ΡΠ»Π΅ΠΊΡΡΠΎΡΠΎΡΠ΅Π·ΠΎΠΌ Π² Π΄Π΅Π½Π°ΡΡΡΠΈΡΡΡΡΠΈΡ ΡΡΠ»ΠΎΠ²ΠΈΡΡ ΠΏΠΎΠΊΠ°Π·Π°Π½ΠΎ, ΡΡΠΎ Π² ΠΏΡΠ΅ΠΏΠ°ΡΠ°ΡΠ΅ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ? Π²ΡΡΠ½Π° ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π½ΠΎΠΉ Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡΡ ΠΎΠ±Π»Π°Π΄Π°Π΅Ρ ΠΏΠΎΠ»ΠΈΠΏΠ΅ΠΏΡΠΈΠ΄ Ρ ΠΊΠ°ΠΆΡΡΠ΅ΠΉΡΡ ΠΌΠΎΠ»Π΅ΠΊΡΠ»ΡΡΠ½ΠΎΠΉ ΠΌΠ°ΡΡΠΎΠΉ 68 ΠΊΠΠ°. ΠΠΎΠ»ΠΈΠΏΠ΅ΠΏΡΠΈΠ΄ 68 ΠΊΠΠ° ΡΠ²Π»ΡΠ΅ΡΡΡ ΡΠ»Π°Π±ΠΎΠΊΠΈΡΠ»ΡΠΌ Π±Π΅Π»ΠΊΠΎΠΌ Ρ ΠΈΠ·ΠΎΡΠ»Π΅ΠΊΡΡΠΈΡΠ΅ΡΠΊΠΎΠΉ ΡΠΎΡΠΊΠΎΠΉ Ρ! 6,3.
7. ΠΠΊΡΠΈΠ²Π½ΡΠ΅ ΡΠΎΡΠΌΡ ΠΊΠΈΡΠ»ΠΎΡΠΎΠ΄Π°, Π³Π΅Π½Π΅ΡΠΈΡΡΠ΅ΠΌΡΠ΅ ΡΠΈΡΡΠ΅ΠΌΠΎΠΉ ΠΊΡΠ°Π½ΡΠΈΠ½Π° Ρ ΠΊΡΠ°Π½ΡΠΈΠ½ΠΎΠΊΡΠΈΠ΄Π°Π·ΠΎΠΉ Π² ΡΠΌΠ±ΡΠΈΠΎΠ½Π°Ρ Π²ΡΡΠ½Π°, ΠΏΠΎΠ²ΡΡΠ°ΡΡ Π² Π½Π΅ΡΠΊΠΎΠ»ΡΠΊΠΎ ΡΠ°Π· Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π· Ρ ΠΈ? Π½Π΅ Π²Π»ΠΈΡΡ Π½Π° Π°ΠΊΡΠΈΠ²Π½ΠΎΡΡΡ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ Π°.
8. Π‘Π²ΠΎΠΉΡΡΠ²Π° ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Ρ? ΡΠ²ΠΈΠ΄Π΅ΡΠ΅Π»ΡΡΡΠ²ΡΡΡ ΠΎ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΠ²Π½ΠΎΠΉ ΡΡΠ½ΠΊΡΠΈΠΈ ΡΡΠΎΠ³ΠΎ ΡΠ΅ΡΠΌΠ΅Π½ΡΠ°. ΠΡΡΠΊΠ°Π·Π°Π½ΠΎ ΠΏΡΠ΅Π΄ΠΏΠΎΠ»ΠΎΠΆΠ΅Π½ΠΈΠ΅, ΡΡΠΎ ΠΠΠ-ΠΏΠΎΠ»ΠΈΠΌΠ΅ΡΠ°Π·Π° ^ ΠΎΡΡΡΠ΅ΡΡΠ²Π»ΡΠ΅Ρ ΡΠ΅ΠΏΠ°ΡΠ°ΡΠΈΡ ΠΠΠ Π² ΠΊΠ»Π΅ΡΠΊΠ°Ρ ΡΠΌΠ±ΡΠΈΠΎΠ½ΠΎΠ² ΠΊΠΎΡΡΠΈΡΡΡΡ ΡΡΠ±.
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