Биосинтетическое получение аналогов бактериородопсина

ɆɈɋɄɈȼɋɄȺə ȽɈɋɍȾȺɊɋɌȼȿɇɇȺə ȺɄȺȾȿɆɂə ɌɈɇɄɈɃ ɏɂɆɂɑȿɋɄɈɃ ɌȿɏɇɈɅɈȽɂɂ ɢɦ. Ɇ.ȼ. ɅɈɆɈɇɈɋɈȼȺ ɇɚ ɩɪɚɜɚɯ ɪɭɤɨɩɢɫɢ ɆɂɊɈɇɈȼȺ ȿɄȺɌȿ...

Author: Миронова Е.В.

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ɆɈɋɄɈȼɋɄȺə ȽɈɋɍȾȺɊɋɌȼȿɇɇȺə ȺɄȺȾȿɆɂə ɌɈɇɄɈɃ ɏɂɆɂɑȿɋɄɈɃ ɌȿɏɇɈɅɈȽɂɂ ɢɦ. Ɇ.ȼ. ɅɈɆɈɇɈɋɈȼȺ

ɇɚ ɩɪɚɜɚɯ ɪɭɤɨɩɢɫɢ

ɆɂɊɈɇɈȼȺ ȿɄȺɌȿɊɂɇȺ ȼȺɅȿɇɌɂɇɈȼɇȺ

ȻɂɈɋɂɇɌȿɌɂɑȿɋɄɈȿ ɉɈɅɍɑȿɇɂȿ ȺɇȺɅɈȽɈȼ ȻȺɄɌȿɊɂɈɊɈȾɈɉɋɂɇȺ

ɋɩɟɰɢɚɥɶɧɨɫɬɶ 03.00.23. - Ȼɢɨɬɟɯɧɨɥɨɝɢɹ. ȾɂɋɋȿɊɌȺɐɂə ɧɚ ɫɨɢɫɤɚɧɢɟ ɭɱɟɧɨɣ ɫɬɟɩɟɧɢ ɤɚɧɞɢɞɚɬɚ ɯɢɦɢɱɟɫɤɢɯ ɧɚɭɤ

ɇɚɭɱɧɵɟ ɪɭɤɨɜɨɞɢɬɟɥɢ: ɞɨɤɬɨɪ ɯɢɦɢɱɟɫɤɢɯ ɧɚɭɤ ɏɨɞɨɧɨɜ Ⱥ.Ⱥ. ɞɨɤɬɨɪ ɛɢɨɥɨɝɢɱɟɫɤɢɯ ɧɚɭɤ, ɫ.ɧ.ɫ. ɋɤɥɚɞɧɟɜ Ⱦ.Ⱥ.

Ɇɨɫɤɜɚ, 2002

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ɋɈȾȿɊɀȺɇɂȿ. ȼȼȿȾȿɇɂȿ. ............................................................................................................................4 ɈȻɁɈɊ ɅɂɌȿɊȺɌɍɊɕ. Ƚɚɥɨɮɢɥɶɧɵɟ ɛɚɤɬɟɪɢɢ ɢ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ ɜ ɫɨɜɪɟɦɟɧɧɨɣ ɛɢɨɬɟɯɧɨɥɨɝɢɢ. .......................................................................................6 1. Ƚɚɥɨɛɚɤɬɟɪɢɢ. .....................................................................................................................6 1.1. ɉɪɢɦɟɧɟɧɢɟ ɝɚɥɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ ɜ ɛɢɨɬɟɯɧɨɥɨɝɢɢ. .........................................6 1.2. Halobacterium salinarum. ...............................................................................................9 2. ɉɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ ɢ ɪɨɞɨɩɫɢɧɵ ɝɚɥɨɛɚɤɬɟɪɢɣ. .................................................10 3. Ȼɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ: ɫɬɪɭɤɬɭɪɚ ɢ ɮɭɧɤɰɢɢ. .................................................................14 4. Ɉɫɧɨɜɧɵɟ ɫɩɨɫɨɛɵ ɢɡɦɟɧɟɧɢɹ ɩɚɪɚɦɟɬɪɨɜ ɮɨɬɨɰɢɤɥɚ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. ....19 4.1. Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɜɨɡɞɟɣɫɬɜɢɹ. .............................................................................20 4.2. Ɂɚɦɟɧɚ ɯɪɨɦɨɮɨɪɚ. .......................................................................................................21 4.3. Ɍɨɱɟɱɧɵɟ ɡɚɦɟɧɵ ɚɦɢɧɨɤɢɫɥɨɬ. ................................................................................24 5. Ɍɟɯɧɢɱɟɫɤɨɟ ɩɪɢɦɟɧɟɧɢɟ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. .........................................................36 5.1. Ȼɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ ɤɚɤ ɮɨɬɨɯɪɨɦɧɵɣ ɦɨɥɟɤɭɥɹɪɧɵɣ ɦɚɬɟɪɢɚɥ. .......................37 5.2. ɋɨɡɞɚɧɢɟ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɯ ɢɡɨɛɪɚɠɟɧɢɣ ɧɚ ɩɥɟɧɤɚɯ, ɫɨɞɟɪɠɚɳɢɯ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ. ........................................................................................................39 5.3. Ƚɨɥɨɝɪɚɮɢɱɟɫɤɚɹ ɢɧɬɟɪɮɟɪɨɦɟɬɪɢɹ. ........................................................................40 5.4. Ɉɛɪɚɛɨɬɤɚ ɜɢɞɟɨɢɡɨɛɪɚɠɟɧɢɣ. ..................................................................................42 5.5. Ɇɨɥɟɤɭɥɹɪɧɚɹ ɩɚɦɹɬɶ ɧɚ ɨɫɧɨɜɟ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. .........................................44 5.6. Ɇɨɥɟɤɭɥɹɪɧɵɟ ɮɨɬɨɪɟɰɟɩɬɨɪɵ ɧɚ ɨɫɧɨɜɟ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. ........................47 ɌȿɈɊȿɌɂɑȿɋɄȺə ɑȺɋɌɖ ɂ ɈȻɋɍɀȾȿɇɂȿ ɊȿɁɍɅɖɌȺɌɈȼ .....................................49 1. ȼɵɛɨɪ ɨɛɴɟɤɬɨɜ ɢɫɫɥɟɞɨɜɚɧɢɹ ..........................................................................................50 2. Ɉɤɢɫɥɢɬɟɥɶɧɚɹ ɦɨɞɢɮɢɤɚɰɢɹ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɪɟɬɢɧɨɢɞɨɜ...........53 3. ɋɢɧɬɟɡ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ..................................................................................................58 4. ɏɪɨɦɨɮɨɪɧɚɹ ɩɨɥɨɫɬɶ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ......................................................................62 5. Ⱥɧɚɥɢɡ ɞɚɧɧɵɯ ɩɨ ɜɥɢɹɧɢɸ ɬɨɱɟɱɧɵɯ ɦɭɬɚɰɢɣ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ ɧɚ ɫɜɨɣɫɬɜɚ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ .................................................64 6. Ⱥɧɚɥɨɝɢ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ .............................................................................................74 6.1. Ʉɭɥɶɬɢɜɢɪɨɜɚɧɢɟ H. salinarum .......................................................................................74 6.2. ɉɨɥɭɱɟɧɢɟ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ ......................................................................77 6.3. ɋɬɚɛɢɥɶɧɨɫɬɶ ɩɪɟɩɚɪɚɬɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ ɩɪɢ ɞɥɢɬɟɥɶɧɨɦ ɯɪɚɧɟɧɢɢ................79 6.4. Ɉɫɨɛɟɧɧɨɫɬɢ ɮɨɬɨɰɢɤɥɚ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ ...............................................81

2 ɗɄɋɉȿɊɂɆȿɇɌȺɅɖɇȺə ɑȺɋɌɖ. ....................................................................................85 1. ɋɢɧɬɟɡ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ..................................................................................................87 2. ɉɨɥɭɱɟɧɢɟ ɩɪɟɩɚɪɚɬɨɜ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ....................................................................92 3. ɉɨɥɭɱɟɧɢɟ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. ........................................................................95 ȼɕȼɈȾɕ.................................................................................................................................97 ȻɅȺȽɈȾȺɊɇɈɋɌɂ................................................................................................................98 ɋɉɂɋɈɄ ɅɂɌȿɊȺɌɍɊɕ ......................................................................................................99 ȺɄɌ ɈȻ ɂɋɉɈɅɖɁɈȼȺɇɂɂ ɉɊȿɉȺɊȺɌɈȼ ȻȺɄɌȿɊɂɈɊɈȾɈɉɋɂɇȺ....................117

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ɋɉɂɋɈɄ ɉɊɂɇəɌɕɏ ɋɈɄɊȺɓȿɇɂɃ.1 CCD

– ɩɪɢɛɨɪ ɫ ɡɚɪɹɞɨɜɨɣ ɫɜɹɡɶɸ (charge-coupled device)

CHAPS

– 3-[(ɯɨɥɚɦɢɞɨɩɪɨɩɢɥ)ɞɢɦɟɬɢɥɚɦɦɨɧɢɨ]-1-ɩɪɨɩɚɧ ɫɭɥɶɮɨɧɚɬ

CTAB

– ɰɟɬɢɥɬɪɢɦɟɬɢɥɚɦɦɨɧɢɣ ɛɪɨɦɢɞ

DA ɢ LA – ɬɟɦɧɨɚɞɚɩɬɢɪɨɜɚɧɧɚɹ ɢ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɚɹ ɮɨɪɦɵ ȻɊ DMPC

– L-D-1,2-ɞɢɦɢɪɢɫɬɨɢɥɮɨɫɮɚɬɢɞɢɥɯɨɥɢɧ

ITO

– ɨɤɫɢɞ ɢɧɞɢɹ-ɬɢɬɚɧɚ, ɩɪɨɡɪɚɱɧɵɣ ɦɚɬɟɪɢɚɥ ɞɥɹ ɫɨɡɞɚɧɢɹ ɷɥɟɤɬɪɨɞɨɜ ɜ ɨɩɬɢɱɟɫɤɢɯ ɭɫɬɪɨɣɫɬɜɚɯ

1

LCD

– ɠɢɞɤɨɤɪɢɫɬɚɥɥɢɱɟɫɤɢɣ ɞɢɫɩɥɟɣ

NBS

– N-ɛɪɨɦɫɭɤɰɢɧɢɦɢɞ

OD

– ɨɩɬɢɱɟɫɤɚɹ ɩɥɨɬɧɨɫɬɶ

PCC

– ɯɥɨɪɨɯɪɨɦɚɬ ɩɢɪɢɞɢɧɢɹ

SB

– ɨɫɧɨɜɚɧɢɟ ɒɢɮɮɚ

SDS

– ɞɨɞɟɰɢɥɫɭɥɶɮɚɬ ɧɚɬɪɢɹ

SLM

– ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɵɣ ɫɜɟɬɨɦɨɞɭɥɹɬɨɪ

ȺȻɊ

– ɚɧɚɥɨɝɢ ȻɊ

ȻɈ

– ɛɚɤɬɟɪɢɨɨɩɫɢɧ

ȻɊ

– ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ

ȻɊWT

– ȻɊ ɫ ɩɪɢɪɨɞɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ (ɞɢɤɨɝɨ ɬɢɩɚ)

ȾɆɋɈ

– ɞɢɦɟɬɢɥɫɭɥɶɮɨɤɫɢɞ

ȾɆɎȺ

– ɞɢɦɟɬɢɥɮɨɪɦɚɦɢɞ

Ʉɀ

– ɤɭɥɶɬɭɪɚɥɶɧɚɹ ɠɢɞɤɨɫɬɶ

ɉɆ

– ɩɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ

ɉɗɅ

– ɩɟɬɪɨɥɟɣɧɵɣ ɷɮɢɪ

ɋɊ I, II

– ɫɟɧɫɨɪɧɵɟ ɪɨɞɨɩɫɢɧɵ I ɢ II

ɌȽɎ

– ɬɟɬɪɚɝɢɞɪɨɮɭɪɚɧ

ɉɪɢ ɨɛɨɡɧɚɱɟɧɢɢ ɦɭɬɚɧɬɧɵɯ ɛɟɥɤɨɜ ɢɫɩɨɥɶɡɨɜɚɥɢ ɨɛɳɟɩɪɢɧɹɬɭɸ ɫɢɫɬɟɦɭ: «ɢɫɯɨɞɧɵɣ ɨɫɬɚɬɨɤ ɚɦɢɧɨɤɢɫɥɨɬɵ – ɟɝɨ ɩɨɥɨɠɟɧɢɟ ɜ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ ȻɊ – ɚɦɢɧɨɤɢɫɥɨɬɧɵɣ ɨɫɬɚɬɨɤ, ɧɚ ɤɨɬɨɪɵɣ ɩɪɨɜɟɞɟɧɚ ɡɚɦɟɧɚ», ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ ɨɛɨɡɧɚɱɟɧɵ ɫɬɚɧɞɚɪɬɧɵɦɢ ɨɞɧɨɛɭɤɜɟɧɧɵɦɢ ɫɨɤɪɚɳɟɧɢɹɦɢ. ȼ ɨɫɬɚɥɶɧɵɯ ɫɥɭɱɚɹɯ, ɜ ɬɟɤɫɬɟ ɢ ɧɚ ɪɢɫɭɧɤɚɯ, ɢɫɩɨɥɶɡɨɜɚɥɢ ɫɬɚɧɞɚɪɬɧɵɟ ɬɪɟɯɛɭɤɜɟɧɧɵɟ ɨɛɨɡɧɚɱɟɧɢɹ ɚɦɢɧɨɤɢɫɥɨɬ.

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ȼȼȿȾȿɇɂȿ. ȼ ɩɪɢɪɨɞɟ ɨɛɧɚɪɭɠɟɧɨ ɞɨɫɬɚɬɨɱɧɨ ɦɚɥɨɟ ɤɨɥɢɱɟɫɬɜɨ ɛɟɥɤɨɜ, ɮɭɧɤɰɢɢ ɤɨɬɨɪɵɯ ɫɜɹɡɚɧɵ ɫ ɩɪɟɨɛɪɚɡɨɜɚɧɢɟɦ ɫɜɟɬɨɜɨɣ ɷɧɟɪɝɢɢ, ɨɞɧɚɤɨ ɢɫɤɥɸɱɢɬɟɥɶɧɨ ɜɚɠɧɚɹ ɪɨɥɶ, ɤɨɬɨɪɭɸ ɨɧɢ ɢɝɪɚɸɬ ɜ ɩɪɨɰɟɫɫɚɯ ɠɢɡɧɟɞɟɹɬɟɥɶɧɨɫɬɢ ɪɚɡɥɢɱɧɵɯ ɨɪɝɚɧɢɡɦɨɜ, ɫɬɚɥɚ ɩɪɢɱɢɧɨɣ ɩɨɜɵɲɟɧɧɨɝɨ ɢɧɬɟɪɟɫɚ ɭɱɟɧɵɯ ɤ ɩɪɨɛɥɟɦɟ ɢɯ ɫɬɪɨɟɧɢɹ ɢ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ. ɉɨ ɦɟɪɟ ɢɫɫɥɟɞɨɜɚɧɢɹ ɫɜɨɣɫɬɜ ɢ ɦɟɯɚɧɢɡɦɚ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ɷɬɢɯ ɛɟɥɤɨɜ, ɛɵɥ ɜɵɹɜɥɟɧ ɪɹɞ ɨɫɨɛɟɧɧɨɫɬɟɣ, ɩɨɡɜɨɥɹɸɳɢɯ ɢɫɩɨɥɶɡɨɜɚɬɶ ɧɟɤɨɬɨɪɵɟ ɢɡ ɧɢɯ ɜ ɬɟɯɧɢɱɟɫɤɢɯ ɰɟɥɹɯ, ɚ ɬɚɤ ɠɟ ɪɚɡɪɚɛɨɬɚɧ ɧɚɛɨɪ ɦɟɬɨɞɨɜ ɧɚɩɪɚɜɥɟɧɧɨɝɨ ɢɡɦɟɧɟɧɢɹ ɫɜɨɣɫɬɜ ɛɟɥɤɚ ɜ ɫɨɨɬɜɟɬɫɬɜɢɢ ɫ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɦɢ ɬɪɟɛɨɜɚɧɢɹɦɢ. Ɉɞɧɢɦ ɢɡ ɧɚɢɛɨɥɟɟ ɩɪɨɫɬɵɯ ɢ ɞɨɫɬɭɩɧɵɯ ɩɪɟɨɛɪɚɡɭɸɳɢɯ ɫɜɟɬ ɛɟɥɤɨɜ ɹɜɥɹɟɬɫɹ ɨɛɧɚɪɭɠɟɧɧɵɣ ɜ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧɚɯ (ɉɆ) ɝɚɥɨɛɚɤɬɟɪɢɣ Halobacterium salinarum (ɪɚɧɟɟ ɧɚɡɵɜɚɟɦɵɯ H. salinarium ɢ H. halobium) ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ (ȻɊ). Ɉɧ ɛɵɥ ɨɬɤɪɵɬ Oesterhelt D. ɢ Stoeckenius W. ɜ 1971 ɝɨɞɭ [1]. ɗɤɫɬɪɟɦɚɥɶɧɵɟ ɭɫɥɨɜɢɹ, ɜ ɤɨɬɨɪɵɯ ɫɭɳɟɫɬɜɭɟɬ ɜ ɩɪɢɪɨɞɟ ɞɚɧɧɵɣ ɜɢɞ ɛɚɤɬɟɪɢɣ (ɤɪɚɣɧɟ ɜɵɫɨɤɚɹ ɤɨɧɰɟɧɬɪɚɰɢɹ ɫɨɥɟɣ, ɩɨɜɵɲɟɧɧɵɟ ɬɟɦɩɟɪɚɬɭɪɵ ɢ ɢɧɬɟɧɫɢɜɧɨɟ ɫɨɥɧɟɱɧɨɟ ɨɛɥɭɱɟɧɢɟ), ɫɩɨɫɨɛɫɬɜɨɜɚɥɢ ɨɬɛɨɪɭ ɛɟɥɤɨɜ, ɨɛɥɚɞɚɸɳɢɯ ɜɵɫɨɤɨɣ ɭɫɬɨɣɱɢɜɨɫɬɶɸ ɤ ɬɚɤɢɦ ɞɟɧɚɬɭɪɢɪɭɸɳɢɦ ɜɧɟɲɧɢɦ ɜɨɡɞɟɣɫɬɜɢɹɦ. ɏɪɨɦɨɮɨɪɨɦ ȻɊ ɹɜɥɹɟɬɫɹ ɩɪɨɬɨɧɢɪɨɜɚɧɧɵɣ ɚɥɶɞɢɦɢɧ ɪɟɬɢɧɚɥɹ ɫ H– ɚɦɢɧɨɝɪɭɩɩɨɣ ɨɫɬɚɬɤɚ Lys–216. ɉɨɝɥɨɳɚɹ ɤɜɚɧɬ ɫɜɟɬɚ, ȻɊ ɩɪɟɬɟɪɩɟɜɚɟɬ ɰɢɤɥ ɫɩɟɤɬɪɚɥɶɧɵɯ ɩɟɪɟɯɨɞɨɜ (ɮɨɬɨɰɢɤɥ), ɫɨɩɪɨɜɨɠɞɚɸɳɢɯɫɹ ɢɡɨɦɟɪɢɡɚɰɢɟɣ ɛɟɥɤɚ, ɞɟɩɪɨɬɨɧɢɪɨɜɚɧɢɟɦ ɢ ɪɟɩɪɨɬɨɧɢɪɨɜɚɧɢɟɦ ɚɥɶɞɢɦɢɧɧɨɣ ɫɜɹɡɢ ɢ ɩɟɪɟɧɨɫɨɦ ɩɪɨɬɨɧɚ. Ɏɨɬɨɰɢɤɥ ȻɊ ɦɨɠɧɨ ɨɩɢɫɚɬɶ ɫɥɟɞɭɸɳɟɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ ɢɧɬɟɪɦɟɞɢɚɬɨɜ: B568 o J625 o K590 o L550 o M412 o N560 o O640 o B568. ɍɧɢɤɚɥɶɧɚɹ ɫɬɚɛɢɥɶɧɨɫɬɶ, ɜ ɫɨɱɟɬɚɧɢɢ ɫ ɧɚɥɢɱɢɟɦ ɮɨɬɨɯɪɨɦɧɵɯ ɫɜɨɣɫɬɜ ȻɊ, ɧɚɪɹɞɭ ɫ ɩɪɨɫɬɨɬɨɣ ɟɝɨ ɩɪɨɢɡɜɨɞɫɬɜɚ ɜ ɛɨɥɶɲɢɯ ɤɨɥɢɱɟɫɬɜɚɯ ɨɛɟɫɩɟɱɢɜɚɸɬ ɲɢɪɨɤɢɣ ɞɢɚɩɚɡɨɧ ɬɟɯɧɢɱɟɫɤɢɯ ɩɪɢɥɨɠɟɧɢɣ, ɜ ɤɨɬɨɪɵɯ ɨɧ ɦɨɠɟɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧ [2]. Ɉɞɧɚɤɨ ɧɚ ɫɟɝɨɞɧɹɲɧɢɣ ɞɟɧɶ ɨɫɭɳɟɫɬɜɢɦɵ ɬɨɥɶɤɨ ɨɩɬɢɱɟɫɤɢɟ ɭɫɬɪɨɣɫɬɜɚ ɧɚ ɨɫɧɨɜɟ ȻɊ, ɩɨɫɤɨɥɶɤɭ ɢɯ ɢɧɬɟɝɪɚɰɢɹ ɜ ɫɨɜɪɟɦɟɧɧɵɟ ɩɪɢɛɨɪɧɵɟ ɫɢɫɬɟɦɵ ɧɚɢɛɨɥɟɟ ɩɪɨɫɬɚ.

5

Ɉɩɬɢɱɟɫɤɢɟ ɩɪɢɥɨɠɟɧɢɹ ɨɫɧɨɜɚɧɵ ɧɚ ɩɪɢɦɟɧɟɧɢɢ ɩɥɟɧɨɤ ȻɊ – ɩɨɥɢɦɟɪɧɵɯ ɦɚɬɪɢɰ ɪɚɡɥɢɱɧɨɝɨ ɫɨɫɬɚɜɚ ɫ ɜɤɥɸɱɟɧɧɵɦɢ ɜ ɧɢɯ ɦɨɥɟɤɭɥɚɦɢ ɛɟɥɤɚ. ȼɩɟɪɜɵɟ ɜ ɦɢɪɟ ɬɚɤɢɟ ɩɥɟɧɤɢ, ɧɚɡɜɚɧɧɵɟ “Ȼɢɨɯɪɨɦ”, ɛɵɥɢ ɩɨɥɭɱɟɧɵ ɢ ɢɫɫɥɟɞɨɜɚɧɵ ɜ Ɋɨɫɫɢɢ ɜ ɪɚɦɤɚɯ ɩɪɨɟɤɬɚ “Ɋɨɞɨɩɫɢɧ” ɜ 1980-ɯ ɝɨɞɚɯ. Ȼɵɥɚ ɩɪɨɞɟɦɨɧɫɬɪɢɪɨɜɚɧɚ ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɢ ɩɟɪɫɩɟɤɬɢɜɧɨɫɬɶ ɢɯ ɩɪɢɦɟɧɟɧɢɹ ɜ ɤɚɱɟɫɬɜɟ ɮɨɬɨɯɪɨɦɧɵɯ ɦɚɬɟɪɢɚɥɨɜ ɢ ɫɪɟɞ ɞɥɹ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɣ ɡɚɩɢɫɢ [3]. ȼ ɩɨɫɥɟɞɧɟɟ ɞɟɫɹɬɢɥɟɬɢɟ ɭɫɢɥɢɹɦɢ ɪɹɞɚ ɢɫɫɥɟɞɨɜɚɬɟɥɶɫɤɢɯ ɤɨɥɥɟɤɬɢɜɨɜ ɞɨɫɬɢɝɧɭɬ ɨɩɪɟɞɟɥɟɧɧɵɣ ɩɪɨɪɵɜ ɜ ɢɫɩɨɥɶɡɨɜɚɧɢɢ ɩɪɟɩɚɪɚɬɨɜ ɩɪɢɪɨɞɧɨɝɨ ɢ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɨɝɨ ȻɊ ɞɥɹ ɫɨɡɞɚɧɢɹ ɪɚɡɥɢɱɧɵɯ ɭɫɬɪɨɣɫɬɜ ɧɚ ɢɯ ɨɫɧɨɜɟ. Ȼɵɥɢ ɩɪɟɞɥɨɠɟɧɵ ɩɪɨɬɨɬɢɩɵ ɬɪɟɯɦɟɪɧɨɣ ɨɩɬɨ-ɷɥɟɤɬɪɨɧɧɨɣ ɩɚɦɹɬɢ, ɮɨɬɨɯɪɨɦɧɵɯ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɯ ɫɪɟɞ, ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɵɯ ɦɨɞɭɥɹɬɨɪɨɜ ɨɩɬɢɱɟɫɤɨɝɨ ɫɢɝɧɚɥɚ ɢ ɛɢɨɫɟɧɫɨɪɨɜ. Ɉɞɧɢɦ ɢɡ ɩɪɚɤɬɢɱɟɫɤɢɯ ɪɟɡɭɥɶɬɚɬɨɜ ɩɪɨɜɨɞɢɦɵɯ ɜ ɷɬɨɣ ɨɛɥɚɫɬɢ ɢɫɫɥɟɞɨɜɚɧɢɣ ɫɬɚɥɨ ɬɨ, ɱɬɨ ɜ 2001 ɝɨɞɭ ɮɢɪɦɚ Munich Innovative Biomaterials, GmbH (MIB) ɩɪɟɞɫɬɚɜɢɥɚ ɧɚ ɪɵɧɨɤ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɣ ɢɧɬɟɪɮɟɪɨɦɟɬɪ FringeMaker-plus® ɧɚ ɨɫɧɨɜɟ ɩɥɟɧɤɢ ɦɭɬɚɧɬɧɨɝɨ ȻɊD96N. ɉɨɫɤɨɥɶɤɭ ɧɚ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɟ ȻɊ ɜɥɢɹɟɬ ɦɨɞɢɮɢɤɚɰɢɹ ɤɚɤ ɛɟɥɤɨɜɨɣ, ɬɚɤ ɢ ɯɪɨɦɨɮɨɪɧɨɣ ɱɚɫɬɟɣ, ɜɨɡɧɢɤɚɟɬ ɧɟɨɛɯɨɞɢɦɨɫɬɶ ɩɨɢɫɤɚ ɧɨɜɵɯ ɫɨɱɟɬɚɧɢɣ ɦɨɞɢɮɢɤɚɰɢɣ ɯɪɨɦɨɮɨɪ-ɛɟɥɨɤ, ɩɨɡɜɨɥɹɸɳɢɯ ɩɨɥɭɱɢɬɶ ɩɟɪɫɩɟɤɬɢɜɧɵɟ ɞɥɹ ɬɟɯɧɢɱɟɫɤɨɝɨ ɢɫɩɨɥɶɡɨɜɚɧɢɹ ɩɪɟɩɚɪɚɬɵ ȻɊ. ɇɚɫɬɨɹɳɚɹ ɪɚɛɨɬɚ ɧɚɩɪɚɜɥɟɧɚ ɧɚ ɪɟɚɥɢɡɚɰɢɸ ɷɬɨɣ ɡɚɞɚɱɢ, ɟɟ ɰɟɥɶɸ ɛɵɥɨ ɤɨɦɩɥɟɤɫɧɨɟ ɢɫɫɥɟɞɨɜɚɧɢɟ ɜɥɢɹɧɢɹ ɨɞɧɨɜɪɟɦɟɧɧɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɛɟɥɤɨɜɨɣ ɢ ɯɪɨɦɨɮɨɪɧɨɣ ɱɚɫɬɟɣ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ (ɦɚɤɫɢɦɭɦɵ ɩɨɝɥɨɳɟɧɢɹ, ɜɪɟɦɹ ɪɟɝɟɧɟɪɚɰɢɢ ɚɩɨɛɟɥɤɨɜ ɫ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ, ɨɫɨɛɟɧɧɨɫɬɢ ɤɢɧɟɬɢɤɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɚ ɢ ɫɬɚɛɢɥɶɧɨɫɬɶ ɩɪɟɩɚɪɚɬɨɜ). Ⱦɢɫɫɟɪɬɚɰɢɨɧɧɚɹ ɪɚɛɨɬɚ ɹɜɥɹɥɚɫɶ ɱɚɫɬɶɸ ɩɥɚɧɨɜɵɯ ɢɫɫɥɟɞɨɜɚɧɢɣ ɩɨ ɪɚɡɪɚɛɨɬɤɟ ɧɨɜɵɯ ɨɩɬɢɱɟɫɤɢɯ ɦɚɬɟɪɢɚɥɨɜ ɧɚ ɨɫɧɨɜɟ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ, ɩɪɨɜɨɞɢɦɵɯ ɧɚ ɤɚɮɟɞɪɟ ɛɢɨɬɟɯɧɨɥɨɝɢɢ ɆɂɌɏɌ ɢɦ. Ɇ.ȼ. Ʌɨɦɨɧɨɫɨɜɚ ɫɨɜɦɟɫɬɧɨ ɫ ɇɂɂ Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɨɣ ɛɢɨɥɨɝɢɢ ɢɦ. Ⱥ.ɇ. Ȼɟɥɨɡɟɪɫɤɨɝɨ ɆȽɍ, ɐɟɧɬɪɨɦ ɮɨɬɨɯɢɦɢɢ ɊȺɇ, ɂɧɫɬɢɬɭɬɨɦ ɛɢɨɯɢɦɢɱɟɫɤɨɣ ɮɢɡɢɤɢ ɢɦ. ɇ.Ɇ. ɗɦɚɧɭɷɥɹ ɊȺɇ, OAO ɐɇɂɌɂ “Ɍɟɯɧɨɦɚɲ” ɢ ɎȽɍɉ «Ƚɨɫɇɂɂɝɟɧɟɬɢɤɚ». Ɋɚɛɨɬɚ ɩɪɨɜɨɞɢɬɫɹ ɜ ɪɚɦɤɚɯ ɜɵɩɨɥɧɟɧɢɹ ɝɪɚɧɬɨɜ ɊɎɎɂ 0103-32078 ɢ 02-03-06555 ɢ ɝɪɚɧɬɚ ɂɇɌȺɋ-01-0267.

6

ɈȻɁɈɊ ɅɂɌȿɊȺɌɍɊɕ. Ƚɚɥɨɮɢɥɶɧɵɟ ɛɚɤɬɟɪɢɢ ɢ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ ɜ ɫɨɜɪɟɦɟɧɧɨɣ ɛɢɨɬɟɯɧɨɥɨɝɢɢ. 1. Ƚɚɥɨɛɚɤɬɟɪɢɢ. Ƚɚɥɨɛɚɤɬɟɪɢɢ ɨɬɧɨɫɹɬ ɤ ɚɪɯɟɛɚɤɬɟɪɢɹɦ, ɤɨɬɨɪɵɟ ɩɪɟɞɫɬɚɜɥɹɸɬ ɫɨɛɨɣ ɫɨɜɟɪɲɟɧɧɨ ɨɫɨɛɭɸ ɥɢɧɢɸ ɷɜɨɥɸɰɢɢ, ɨɬɥɢɱɚɸɳɭɸɫɹ ɤɚɤ ɨɬ ɩɪɨɤɚɪɢɨɬ, ɬɚɤ ɢ ɨɬ ɷɭɤɚɪɢɨɬ. ɗɬɢ ɛɚɤɬɟɪɢɢ ɪɚɫɩɪɨɫɬɪɚɧɟɧɵ ɬɚɦ, ɝɞɟ ɟɫɬɶ ɩɨɞɯɨɞɹɳɢɟ ɭɫɥɨɜɢɹ ɞɥɹ ɢɯ ɪɨɫɬɚ: ɜɵɫɨɤɨɟ ɫɨɞɟɪɠɚɧɢɟ NaCl ɢ ɞɪɭɝɢɯ ɧɟɨɛɯɨɞɢɦɵɯ ɢɨɧɨɜ, ɜ ɫɨɥɟɧɵɯ ɩɪɢɪɨɞɧɵɯ ɜɨɞɨɟɦɚɯ, ɛɚɫɫɟɣɧɚɯ ɞɥɹ ɜɵɩɚɪɢɜɚɧɢɹ ɫɨɥɢ, ɛɟɥɤɨɜɵɯ ɦɚɬɟɪɢɚɥɚɯ (ɪɵɛɚ, ɦɹɫɨ), ɤɨɧɫɟɪɜɢɪɭɟɦɵɯ ɫ ɩɨɦɨɳɶɸ ɫɨɥɢ [3]. ȼ 1960-ɯ ɝɨɞɚɯ ɧɚɱɚɥɢɫɶ ɩɟɪɜɵɟ ɷɤɫɩɟɪɢɦɟɧɬɵ ɩɨ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɸ ɢ ɜɵɞɟɥɟɧɢɸ ɝɚɥɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ [4-6]. ȼɫɟɝɨ ɡɚ ɩɪɨɲɟɞɲɢɣ ɩɟɪɢɨɞ ɛɵɥɨ ɜɵɞɟɥɟɧɨ ɨɤɨɥɨ 170 ɩɪɢɪɨɞɧɵɯ ɲɬɚɦɦɨɜ. ȼ ɨɫɧɨɜɧɨɦ ɷɬɨ ɛɵɥɢ ɭɦɟɪɟɧɧɨ ɝɚɥɨɮɢɥɶɧɵɟ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ, ɬɨɥɶɤɨ ɧɟɫɤɨɥɶɤɨ ɲɬɚɦɦɨɜ ɨɤɚɡɚɥɢɫɶ ɷɤɫɬɪɟɦɚɥɶɧɵɦɢ ɝɚɥɨɮɢɥɚɦɢ [7]. Ȼɚɤɬɟɪɢɢ, ɤɨɬɨɪɵɦ ɞɥɹ ɪɨɫɬɚ ɧɟɨɛɯɨɞɢɦɨ ɩɪɢɫɭɬɫɬɜɢɟ NaCl, ɩɨ ɫɬɟɩɟɧɢ ɝɚɥɨɮɢɥɶɧɨɫɬɢ ɞɟɥɹɬ ɧɚ ɧɟɫɤɨɥɶɤɨ ɬɢɩɨɜ. ɇɚɢɛɨɥɟɟ ɲɢɪɨɤɨ ɢɫɩɨɥɶɡɭɸɬ ɤɥɚɫɫɢɮɢɤɚɰɢɸ ɩɨ ɞɢɚɩɚɡɨɧɭ ɤɨɧɰɟɧɬɪɚɰɢɣ NaCl, ɜ ɤɨɬɨɪɨɦ ɦɨɝɭɬ ɪɚɫɬɢ ɞɚɧɧɵɟ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ, ɩɪɟɞɥɨɠɟɧɧɭɸ Kushner D.J. [8]: ɝɚɥɨɬɨɥɥɟɪɚɧɬɧɵɟ (ɨɬ 0 ɞɨ 5% NaCl), ɷɤɫɬɪɟɦɚɥɶɧɨ ɝɚɥɨɬɨɥɥɟɪɚɧɬɧɵɟ (0 ɞɨ 15% NaCl), ɥɟɝɤɢɟ ɝɚɥɨɮɢɥɵ (ɨɤɨɥɨ 3% NaCl), ɭɦɟɪɟɧɧɵɟ ɝɚɥɨɮɢɥɵ (ɨɬ 3 ɞɨ 15% NaCl), ɩɨɝɪɚɧɢɱɧɵɟ ɷɤɫɬɪɟɦɚɥɶɧɵɟ ɝɚɥɨɮɢɥɵ (>15% NaCl), ɷɤɫɬɪɟɦɚɥɶɧɵɟ ɝɚɥɨɮɢɥɵ (ɨɤɨɥɨ 25% NaCl). ȼ IX ɢɡɞɚɧɢɢ ɨɩɪɟɞɟɥɢɬɟɥɹ ɛɚɤɬɟɪɢɣ Ȼɟɪɞɠɢ [9] ɷɤɫɬɪɟɦɚɥɶɧɨ ɝɚɥɨɮɢɥɶɧɵɟ ɚɪɯɟɛɚɤɬɟɪɢɢ ɨɛɴɟɞɢɧɟɧɵ ɜ ɩɨɪɹɞɨɤ Halobacteriales, ɫɟɦɟɣɫɬɜɨ Halobacteriaceae, ɤɨɬɨɪɨɟ ɜɤɥɸɱɚɟɬ 6 ɪɨɞɨɜ ɢ ɛɨɥɟɟ 20 ɜɢɞɨɜ, ɪɚɡɥɢɱɚɸɳɢɯɫɹ ɮɨɪɦɨɣ ɤɥɟɬɨɤ, ɫɩɨɫɨɛɧɨɫɬɶɸ ɤ ɞɜɢɠɟɧɢɸ, ɨɬɧɨɲɟɧɢɟɦ ɤ ɤɢɫɥɨɬɧɨɫɬɢ ɫɪɟɞɵ, ɭɫɬɨɣɱɢɜɨɫɬɶɸ ɤ NaCl ɢ ɞɪɭɝɢɦɢ ɩɪɢɡɧɚɤɚɦɢ. ɂɫɫɥɟɞɨɜɚɧɢɟ ɛɢɨɯɢɦɢɢ ɝɚɥɨɛɚɤɬɟɪɢɣ ɩɨɡɜɨɥɢɥɨ ɩɪɟɞɥɨɠɢɬɶ ɰɟɥɵɣ ɪɹɞ ɜɚɪɢɚɧɬɨɜ ɩɪɢɦɟɧɟɧɢɹ ɷɬɢɯ ɦɢɤɪɨɨɪɝɚɧɢɡɦɨɜ ɜ ɛɢɨɬɟɯɧɨɥɨɝɢɢ.

1.1. ɉɪɢɦɟɧɟɧɢɟ ɝɚɥɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ ɜ ɛɢɨɬɟɯɧɨɥɨɝɢɢ. ɋɩɨɫɨɛɧɨɫɬɶ ɝɚɥɨɛɚɤɬɟɪɢɣ ɜɵɠɢɜɚɬɶ ɜ ɷɤɫɬɪɟɦɚɥɶɧɵɯ ɭɫɥɨɜɢɹɯ ɩɨɜɵɲɟɧɧɨɣ ɤɨɧɰɟɧɬɪɚɰɢɢ ɫɨɥɢ ɢ ɢɧɬɟɧɫɢɜɧɨɣ ɢɧɫɨɥɹɰɢɢ ɨɛɭɫɥɨɜɥɟɧɚ ɩɪɢɨɛɪɟɬɟɧɧɨɣ ɜ ɩɪɨɰɟɫɫɟ ɷɜɨɥɸɰɢɢ ɜɵɫɨɤɨɣ ɫɬɚɛɢɥɶɧɨɫɬɶɸ ɤɥɟɬɨɱɧɵɯ ɤɨɦɩɨɧɟɧɬ ɤ ɪɚɡɪɭɲɢɬɟɥɶɧɵɦ ɜɧɟɲɧɢɦ ɜɨɡɞɟɣɫɬɜɢɹɦ. ɗɬɨ ɨɛɭɫɥɚɜɥɢɜɚɟɬ ɡɧɚɱɢɬɟɥɶɧɵɣ ɩɨɬɟɧɰɢɚɥ ɝɚɥɨɛɚɤɬɟɪɢɣ ɞɥɹ ɛɢɨɬɟɯɧɨɥɨɝɢɢ [10]. ɂɯ ɛɨɥɶɲɢɦ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɦ ɩɪɟɢɦɭɳɟɫɬɜɨɦ ɬɚɤ ɠɟ ɹɜɥɹɟɬɫɹ

7 ɜɨɡɦɨɠɧɨɫɬɶ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɜ ɨɬɤɪɵɬɨɦ ɪɟɡɟɪɜɭɚɪɟ ɛɟɡ ɪɢɫɤɚ ɡɚɝɪɹɡɧɟɧɢɹ ɤɚɤ ɨɤɪɭɠɚɸɳɟɣ ɫɪɟɞɵ ɬɚɤ ɢ ɫɚɦɨɣ ɤɭɥɶɬɭɪɵ. ɇɢɠɟ ɩɪɢɜɟɞɟɧɵ ɨɫɧɨɜɧɵɟ ɧɚɩɪɚɜɥɟɧɢɹ ɢɫɩɨɥɶɡɨɜɚɧɢɹ ɝɚɥɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ ɢ ɫɢɧɬɟɡɢɪɭɟɦɵɯ ɢɦɢ ɜɟɳɟɫɬɜ ɜ ɛɢɨɬɟɯɧɨɥɨɝɢɢ. 1. Ȼɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ. ɏɨɬɹ ȻɊ ɨɛɧɚɪɭɠɟɧ ɜɨ ɦɧɨɝɢɯ ɲɬɚɦɦɚɯ ɷɤɫɬɪɟɦɚɥɶɧɨ ɝɚɥɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ, ɞɥɹ ɟɝɨ ɩɪɨɢɡɜɨɞɫɬɜɚ ɢɫɩɨɥɶɡɭɸɬ Halobacterium salinarum, ɫɨɞɟɪɠɚɳɢɣ ɡɧɚɱɢɬɟɥɶɧɵɟ ɤɨɥɢɱɟɫɬɜɚ ȻɊ. ɍɧɢɤɚɥɶɧɵɟ ɮɨɬɨɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɷɬɨɝɨ ɛɟɥɤɚ ɞɟɥɚɸɬ ɜɨɡɦɨɠɧɵɦ ɟɝɨ ɲɢɪɨɤɨɟ ɬɟɯɧɢɱɟɫɤɨɟ ɩɪɢɦɟɧɟɧɢɟ ɜ ɛɢɨɷɥɟɤɬɪɨɧɢɤɟ. ɉɨɞɪɨɛɧɨ ɷɬɨɬ ɜɨɩɪɨɫ ɛɭɞɟɬ ɪɚɫɫɦɨɬɪɟɧ ɜ 5 ɪɚɡɞɟɥɟ ɧɚɫɬɨɹɳɟɝɨ ɨɛɡɨɪɚ ɥɢɬɟɪɚɬɭɪɵ. 2. Ɉɫɦɨɬɢɱɟɫɤɢ ɚɤɬɢɜɧɵɟ ɜɟɳɟɫɬɜɚ (ɨɫɦɨɥɢɬɵ). Ɉɫɧɨɜɧɨɣ ɢɯ ɮɭɧɤɰɢɟɣ ɜ ɤɥɟɬɤɚɯ ɝɚɥɨɛɚɤɬɟɪɢɣ ɹɜɥɹɟɬɫɹ ɩɨɞɞɟɪɠɚɧɢɟ ɩɨɥɨɠɢɬɟɥɶɧɨɝɨ ɜɨɞɧɨɝɨ ɛɚɥɚɧɫɚ ɫ ɜɧɟɲɧɟɣ ɫɪɟɞɨɣ, ɤɨɬɨɪɚɹ ɨɛɥɚɞɚɟɬ ɛɨɥɟɟ ɜɵɫɨɤɨɣ ɢɨɧɧɨɣ ɫɢɥɨɣ. Ɍɚɤɢɦɢ ɜɟɳɟɫɬɜɚɦɢ (ɫɦ. ɪɢɫ. 1) ɫɥɭɠɚɬ ɜɨɞɨɪɚɫɬɜɨɪɢɦɵɟ ɫɚɯɚɪɚ, ɫɩɢɪɬɵ, ɚɦɢɧɨɤɢɫɥɨɬɵ ɢ ɢɯ ɩɪɨɢɡɜɨɞɧɵɟ. Ɉɧɢ ɧɚɤɚɩɥɢɜɚɸɬɫɹ ɜ ɤɥɟɬɤɚɯ ɜ ɡɧɚɱɢɬɟɥɶɧɵɯ ɤɨɥɢɱɟɫɬɜɚɯ, ɚ ɩɪɢ ɪɟɡɤɨɦ ɭɦɟɧɶɲɟɧɢɢ ɢɨɧɧɨɣ ɫɢɥɵ ɫɪɟɞɵ ɛɵɫɬɪɨ ɫɟɤɪɟɬɢɪɭɸɬɫɹ. ɗɬɨ ɞɚɟɬ ɜɨɡɦɨɠɧɨɫɬɶ ɥɟɝɤɨɝɨ ɪɚɡɞɟɥɟɧɢɹ ɠɢɜɵɯ ɤɥɟɬɨɤ ɛɚɤɬɟɪɢɣ ɢ ɫɢɧɬɟɡɢɪɭɟɦɵɯ ɢɦɢ ɨɫɦɨɬɢɱɟɫɤɢ ɚɤɬɢɜɧɵɯ ɜɟɳɟɫɬɜ ɩɪɢ ɩɨɦɨɳɢ ɩɪɨɫɬɨɣ ɩɪɨɰɟɞɭɪɵ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɹ, ɱɬɨ ɩɨɡɜɨɥɹɟɬ ɨɫɭɳɟɫɬɜɢɬɶ ɢɯ ɷɮɮɟɤɬɢɜɧɨɟ ɛɢɨɫɢɧɬɟɬɢɱɟɫɤɨɟ ɩɪɨɢɡɜɨɞɫɬɜɨ. CH2OH CH2OH CH2OH H

OH

HO HO

HN

CH 2OH ɝɥɢɰɟɪɢɧ

+ NH

-

COO

ɷɤɬɨɢɧ

HO HO

O OH

OH

CH2OH O

O

O

H

CH 2OH ɦɨɧɨɝɥɢɤɨɡɢɥɝɥɢɰɟɪɢɧ

HO ɬɪɟɝɚɥɨɡɚ

CH2OH OH

O OH

Ɋɢɫ. 1. Ɉɫɦɨɬɢɱɟɫɤɢ ɚɤɬɢɜɧɵɟ ɜɟɳɟɫɬɜɚ ɝɚɥɨɛɚɤɬɟɪɢɣ [11]. ɉɨɥɭɱɚɟɦɵɟ ɢɡ ɝɚɥɨɛɚɤɬɟɪɢɣ ɨɫɦɨɬɢɱɟɫɤɢ ɚɤɬɢɜɧɵɟ ɜɟɳɟɫɬɜɚ ɦɨɝɭɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧɵ ɜ ɤɚɱɟɫɬɜɟ ɫɬɚɛɢɥɢɡɚɬɨɪɨɜ ɤɥɟɬɨɤ ɢ ɛɢɨɦɨɥɟɤɭɥ (ɛɟɥɤɨɜ, ɧɭɤɥɟɢɧɨɜɵɯ ɤɢɫɥɨɬ, ɦɟɦɛɪɚɧ), ɜ ɤɨɫɦɟɬɢɱɟɫɤɨɣ ɩɪɨɦɵɲɥɟɧɧɨɫɬɢ, ɤɚɤ ɚɧɬɚɝɨɧɢɫɬɵ ɫɨɥɢ ɢ ɤɪɢɨɩɪɨɬɟɤɬɨɪɵ. Ɉɞɢɧ ɢɡ ɧɚɢɛɨɥɟɟ ɪɚɫɩɪɨɫɬɪɚɧɟɧɧɵɯ ɨɫɦɨɥɢɬɨɜ – ɷɤɬɨɢɧ (ɫɦ. ɪɢɫ. 1). Ɉɧ ɦɨɠɟɬ ɛɵɬɶ ɩɪɨɢɡɜɟɞɟɧ ɬɨɥɶɤɨ ɛɢɨɬɟɯɧɨɥɨɝɢɱɟɫɤɢɦɢ ɦɟɬɨɞɚɦɢ ɢɡ Halomonas elongata [12], ɩɪɢɱɟɦ ɪɚɡɪɚɛɨɬɚɧɧɵɣ ɩɪɨɰɟɫɫ ɩɨɡɜɨɥɹɟɬ ɩɨɥɭɱɚɬɶ ɞɨ 155 ɦɝ ɷɤɬɨɢɧɚ ɧɚ 1 ɝ ɫɭɯɨɝɨ ɜɟɫɚ ɛɢɨɦɚɫɫɵ ɛɚɤɬɟɪɢɣ. 3. Ȼɢɨɞɟɝɪɚɞɚɰɢɹ ɨɬɯɨɞɨɜ. ɋɩɨɫɨɛɧɨɫɬɶ ɧɟɤɨɬɨɪɵɯ ɝɚɥɨɮɢɥɶɧɵɯ ɦɢɤɪɨɨɪɝɚɧɢɡɦɨɜ ɨɤɢɫɥɹɬɶ ɭɝɥɟɜɨɞɨɪɨɞɵ ɢ ɪɚɫɬɢ ɜ ɫɨɥɟɧɨɣ ɫɪɟɞɟ ɞɚɟɬ ɜɨɡɦɨɠɧɨɫɬɶ

8 ɢɫɩɨɥɶɡɨɜɚɬɶ ɢɯ ɞɥɹ ɛɢɨɥɨɝɢɱɟɫɤɨɣ ɨɱɢɫɬɤɢ ɦɨɪɫɤɢɯ ɷɤɨɫɢɫɬɟɦ, ɡɚɝɪɹɡɧɟɧɧɵɯ ɧɟɮɬɟɩɪɨɞɭɤɬɚɦɢ [13]. 4. Ȼɢɨɩɨɥɢɦɟɪɵ ɝɚɥɨɛɚɤɬɟɪɢɣ. Ȼɢɨɫɭɪɮɚɤɬɚɧɬɵ ɢ ɷɤɡɨɩɨɥɢɫɚɯɚɪɢɞɵ, ɫɢɧɬɟɡɢɪɭɟɦɵɟ ɦɧɨɝɢɦɢ ɝɚɥɨɛɚɤɬɟɪɢɹɦɢ [14, 15], ɩɪɢ ɤɨɧɬɚɤɬɟ ɫ ɧɟɮɬɶɸ ɫɧɢɠɚɸɬ ɟɟ ɩɨɜɟɪɯɧɨɫɬɧɨɟ ɧɚɬɹɠɟɧɢɟ, ɱɬɨ ɦɨɠɟɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧɨ ɩɪɢ ɧɟɮɬɟɞɨɛɵɱɟ. ɏɨɬɹ ɬɚɤɢɟ ɩɪɟɩɚɪɚɬɵ ɞɨɪɨɝɢ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɫɢɧɬɟɬɢɱɟɫɤɢɦɢ ɚɧɚɥɨɝɚɦɢ, ɨɧɢ ɹɜɥɹɸɬɫɹ ɛɢɨɞɟɝɪɚɞɢɪɭɟɦɵɦɢ ɢ ɧɟ ɧɚɧɨɫɹɬ ɜɪɟɞɚ ɨɤɪɭɠɚɸɳɟɣ ɫɪɟɞɟ. Ƚɚɥɨɛɚɤɬɟɪɢɚɥɶɧɵɟ ɛɢɨɩɨɥɢɦɟɪɵ ɦɨɝɭɬ ɧɚɣɬɢ ɞɨɫɬɚɬɨɱɧɨ ɲɢɪɨɤɨɟ ɩɪɢɦɟɧɟɧɢɟ ɢ ɜ ɦɟɞɢɰɢɧɟ. Ɉɞɢɧ ɢɡ ɩɪɢɦɟɪɨɜ - ɫɭɥɶɮɚɬɢɪɨɜɚɧɧɵɟ ɷɤɡɨɩɨɥɢɫɚɯɚɪɢɞɵ, ɤɨɬɨɪɵɟ ɩɪɟɩɹɬɫɬɜɭɸɬ ɩɪɨɧɢɤɧɨɜɟɧɢɸ ɜɢɪɭɫɨɜ ɜ ɤɥɟɬɤɢ. Ȼɵɥ ɩɪɟɞɥɨɠɟɧ ɷɮɮɟɤɬɢɜɧɵɣ ɦɟɬɨɞ ɢɯ ɩɪɨɢɡɜɨɞɫɬɜɚ ɢɦɦɨɛɢɥɢɡɨɜɚɧɧɵɦɢ ɤɥɟɬɤɚɦɢ ɝɚɥɨɮɢɥɶɧɵɯ ɰɢɚɧɨɛɚɤɬɟɪɢɣ [16]. ȿɳɟ ɨɞɧɢɦ ɩɪɢɦɟɪɨɦ ɦɨɠɟɬ ɫɥɭɠɢɬɶ ɢɫɩɨɥɶɡɨɜɚɧɢɟ ɝɚɥɨɛɚɤɬɟɪɢɚɥɶɧɵɯ ɥɢɩɢɞɨɜ ɞɥɹ ɫɨɡɞɚɧɢɹ ɥɢɩɨɫɨɦ. ɉɨɫɤɨɥɶɤɭ ɥɢɩɢɞɵ ɝɚɥɨɛɚɤɬɟɪɢɣ ɫɨɞɟɪɠɚɬ ɞɢɝɢɞɪɨɮɢɬɨɥ, ɨɧɢ ɡɧɚɱɢɬɟɥɶɧɨ ɛɨɥɟɟ ɫɬɚɛɢɥɶɧɵ, ɱɟɦ ɥɢɩɢɞɵ ɚɰɢɥɶɧɨɝɨ ɬɢɩɚ, ɫɨɞɟɪɠɚɳɢɟ ɷɮɢɪɵ ɠɢɪɧɵɯ ɤɢɫɥɨɬ. ɇɚɩɪɢɦɟɪ, ɩɨɥɭɱɚɟɦɵɟ ɢɡ ɩɨɥɹɪɧɵɯ ɥɢɩɢɞɨɜ Halobacterium cutirubrum ɥɢɩɨɫɨɦɵ ɭɫɬɨɣɱɢɜɵ ɤ ɞɟɣɫɬɜɢɸ ɮɨɫɮɨɥɢɩɚɡ Ⱥ2 ɢ ȼ ɢ ɦɨɝɭɬ ɯɪɚɧɢɬɶɫɹ ɜ ɩɪɢɫɭɬɫɬɜɢɢ ɜɨɡɞɭɯɚ ɛɨɥɟɟ 60 ɞɧɟɣ [17]. ɉɨɥɢɝɢɞɪɨɤɫɢɚɥɤɚɧɨɚɬɵ – ɜɧɭɬɪɢɤɥɟɬɨɱɧɵɟ ɩɨɥɢɦɟɪɵ ɝɚɥɨɛɚɤɬɟɪɢɣ, ɩɨ ɫɜɨɢɦ ɫɜɨɣɫɬɜɚɦ ɧɟɤɨɬɨɪɵɟ ɢɡ ɧɢɯ ɧɚɩɨɦɢɧɚɸɬ ɩɨɥɢɷɬɢɥɟɧ [18]. ɗɬɢ ɛɢɨɞɟɝɪɚɞɢɪɭɟɦɵɟ ɩɨɥɢɦɟɪɵ ɜɨ ɦɧɨɝɢɯ ɫɥɭɱɚɹɯ ɦɨɝɭɬ ɫ ɭɫɩɟɯɨɦ ɡɚɦɟɧɢɬɶ ɭɝɥɟɜɨɞɨɪɨɞɧɵɟ ɚɧɚɥɨɝɢ, ɩɨɫɤɨɥɶɤɭ ɢɯ ɩɪɨɢɡɜɨɞɫɬɜɨ ɞɨɫɬɚɬɨɱɧɨ ɞɟɲɟɜɨ. ɇɚɩɪɢɦɟɪ, Haloferax mediterranei, ɤɭɥɶɬɢɜɢɪɭɟɦɵɟ ɧɚ ɤɪɚɯɦɚɥɨɫɨɞɟɪɠɚɳɟɣ ɫɪɟɞɟ, ɧɚɤɚɩɥɢɜɚɸɬ ɩɨɥɢ(Eɝɢɞɪɨɤɫɢɦɚɫɥɹɧɭɸ ɤɢɫɥɨɬɭ) ɜ ɤɨɥɢɱɟɫɬɜɟ ɞɨ 60% ɫɭɯɨɝɨ ɜɟɫɚ [19]. Ʌɟɝɤɨɫɬɶ ɜɵɞɟɥɟɧɢɹ ɢ ɜɵɫɨɤɚɹ ɷɤɨɥɨɝɢɱɟɫɤɚɹ ɱɢɫɬɨɬɚ ɩɪɨɢɡɜɨɞɫɬɜɚ ɷɬɨɝɨ ɩɨɥɢɦɟɪɚ ɨɛɭɫɥɨɜɥɟɧɵ ɬɟɦ, ɱɬɨ ɥɢɡɢɫ ɤɥɟɬɨɤ ɜɵɡɵɜɚɟɬɫɹ ɜɨɡɞɟɣɫɬɜɢɟɦ ɧɚ ɧɢɯ ɞɢɫɬɢɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ. 5. Ɏɟɪɦɟɧɬɵ. Ɋɹɞ ɝɚɥɨɛɚɤɬɟɪɢɚɥɶɧɵɯ ɝɢɞɪɨɥɚɡ ɦɨɝɭɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧɵ ɞɥɹ ɩɨɥɭɱɟɧɢɹ ɚɦɢɧɨɤɢɫɥɨɬ D-ɪɹɞɚ (ɹɜɥɹɸɳɢɯɫɹ ɩɨɥɭɩɪɨɞɭɤɬɚɦɢ ɜ ɫɢɧɬɟɡɟ ɦɧɨɝɢɯ ɚɧɬɢɛɢɨɬɢɤɨɜ, ɩɟɩɬɢɞɧɵɯ ɝɨɪɦɨɧɨɜ ɢ ɩɟɫɬɢɰɢɞɨɜ) ɢ ɜ ɩɟɩɬɢɞɧɨɦ ɫɢɧɬɟɡɟ [20-21]. Ɉɞɧɚɤɨ ɨɛɳɢɦ ɧɟɞɨɫɬɚɬɤɨɦ ɝɚɥɨɛɚɤɬɟɪɢɚɥɶɧɵɯ ɮɟɪɦɟɧɬɨɜ ɹɜɥɹɟɬɫɹ ɧɟɨɛɯɨɞɢɦɨɫɬɶ ɞɨɫɬɚɬɨɱɧɨ ɜɵɫɨɤɢɯ ɤɨɧɰɟɧɬɪɚɰɢɣ ɫɨɥɢ ɞɥɹ ɩɨɞɞɟɪɠɚɧɢɹ ɢɯ ɚɤɬɢɜɧɨɫɬɢ ɢ ɧɚɬɢɜɧɨɝɨ ɫɨɫɬɨɹɧɢɹ. 6. ɉɢɳɟɜɚɹ ɛɢɨɬɟɯɧɨɥɨɝɢɹ. Ƚɚɥɨɬɨɥɟɪɚɧɬɧɵɟ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ ɢɝɪɚɸɬ ɫɭɳɟɫɬɜɟɧɧɭɸ ɪɨɥɶ ɩɪɢ ɮɟɪɦɟɧɬɚɰɢɢ ɜ ɩɪɢɫɭɬɫɬɜɢɢ ɫɨɥɢ. ɉɪɨɢɫɯɨɞɹɳɟɟ ɫ ɢɯ ɩɨɦɨɳɶɸ ɛɪɨɠɟɧɢɟ ɩɪɢɞɚɟɬ ɩɪɨɞɭɤɬɚɦ ɯɚɪɚɤɬɟɪɧɵɣ ɜɤɭɫ ɢ ɡɚɩɚɯ, ɱɬɨ ɢɫɩɨɥɶɡɭɟɬɫɹ ɞɥɹ ɩɪɨɢɡɜɨɞɫɬɜɚ ɧɟɤɨɬɨɪɵɯ ɦɚɪɢɧɚɞɨɜ ɢ ɫɨɭɫɨɜ [22, 23].

9 Ɍɚɤ ɠɟ ɢɡ ɛɢɨɦɚɫɫɵ ɝɚɥɨɛɚɤɬɟɪɢɣ ɦɨɠɧɨ ɩɨɥɭɱɚɬɶ ɪɚɡɥɢɱɧɵɟ ɩɢɳɟɜɵɟ ɞɨɛɚɜɤɢ ɢ ɤɪɚɫɢɬɟɥɢ. ɇɚɩɪɢɦɟɪ, ɛɢɨɦɚɫɫɚ ɚɧɬɚɪɤɬɢɱɟɫɤɢɯ ɦɨɪɫɤɢɯ ɛɚɤɬɟɪɢɣ ɫɨɞɟɪɠɢɬ ɡɧɚɱɢɬɟɥɶɧɵɟ ɤɨɥɢɱɟɫɬɜɚ ɩɨɥɢɧɟɧɚɫɵɳɟɧɧɵɯ ɠɢɪɧɵɯ ɤɢɫɥɨɬ [24, 25] ɢ ɦɨɠɟɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧɚ ɞɥɹ ɢɯ ɩɪɨɢɡɜɨɞɫɬɜɚ, ɱɬɨ ɡɧɚɱɢɬɟɥɶɧɨ ɞɟɲɟɜɥɟ, ɱɟɦ ɪɟɚɥɢɡɭɟɦɨɟ ɜ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɩɪɨɢɡɜɨɞɫɬɜɨ ɧɚ ɨɫɧɨɜɟ ɪɵɛɧɨɝɨ ɫɵɪɶɹ. ȼ Ɋɨɫɫɢɢ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɩɪɨɢɡɜɨɞɫɬɜɨ ɤɨɦɩɥɟɤɫɧɨɣ ɩɢɳɟɜɨɣ ɞɨɛɚɜɤɢ ɧɚ ɨɫɧɨɜɟ ɛɢɨɦɚɫɫɵ ɝɚɥɨɛɚɤɬɟɪɢɣ Halobacterium salinarum [26]. ȼ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɨɬɟɱɟɫɬɜɟɧɧɵɟ ɩɪɟɩɚɪɚɬɵ, ɫɨɞɟɪɠɚɳɢɟ ɥɢɨɮɢɥɢɡɢɪɨɜɚɧɧɭɸ ɛɢɨɦɚɫɫɭ ɷɬɢɯ ɝɚɥɨɛɚɤɬɟɪɢɣ («Ȼɚɤɫɢɧ» ɢ «Ȼɚɤɫɢɧɨɜɚɹ ɦɚɡɶ»), ɩɪɨɲɥɢ ɤɥɢɧɢɱɟɫɤɢɟ ɢɫɩɵɬɚɧɢɹ ɧɚ ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɢ ɛɟɡɨɩɚɫɧɨɫɬɶ ɜ ɭɫɥɨɜɢɹɯ in vitro ɢ in vivo. Ⱦɨɤɚɡɚɧɨ ɚɧɬɢɨɤɫɢɞɚɧɬɧɨɟ ɢ ɪɚɞɢɨɩɪɨɬɟɤɬɨɪɧɨɟ ɞɟɣɫɬɜɢɟ ɩɪɟɩɚɪɚɬɚ «Ȼɚɤɫɢɧ» ɢ ɩɨɥɭɱɟɧɨ ɪɚɡɪɟɲɟɧɢɟ ɧɚ ɟɝɨ ɩɪɢɦɟɧɟɧɢɟ ɜ ɤɚɱɟɫɬɜɟ ɩɢɳɟɜɨɣ ɞɨɛɚɜɤɢ ɧɚ ɬɟɪɪɢɬɨɪɢɢ ɊɎ. ɇɚ ɨɫɧɨɜɚɧɢɢ ɤɥɢɧɢɱɟɫɤɢɯ ɢɫɩɵɬɚɧɢɣ «Ȼɚɤɫɢɧɨɜɚɹ ɦɚɡɶ» ɪɟɤɨɦɟɧɞɨɜɚɧɚ ɤ ɩɪɢɦɟɧɟɧɢɸ ɤɚɤ ɩɪɨɬɢɜɨɜɨɫɩɚɥɢɬɟɥɶɧɨɟ, ɪɟɝɟɧɟɪɢɪɭɸɳɟɟ ɢ ɤɟɪɚɬɨɩɥɚɫɬɢɱɟɫɤɨɟ ɫɪɟɞɫɬɜɨ ɜ ɤɨɦɩɥɟɤɫɧɨɣ ɬɟɪɚɩɢɢ ɪɹɞɚ ɯɪɨɧɢɱɟɫɤɢɯ ɡɚɛɨɥɟɜɚɧɢɣ ɤɨɠɢ ɢ ɜ ɯɢɪɭɪɝɢɱɟɫɤɨɣ ɩɪɚɤɬɢɤɟ.

1.2. Halobacterium salinarum. Ɋɚɡɥɢɱɧɵɟ ɲɬɚɦɦɵ Halobacterium salinarum ɜ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɢɫɩɨɥɶɡɭɸɬɫɹ ɞɥɹ ɩɪɨɢɡɜɨɞɫɬɜɚ ȻɊ. ɋɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ, ɱɬɨ ɫ ɦɨɦɟɧɬɚ ɨɛɧɚɪɭɠɟɧɢɹ ɷɬɢɯ ɛɚɤɬɟɪɢɣ ɧɟɫɤɨɥɶɤɨ ɪɚɡ ɦɟɧɹɥɢ ɢɯ ɧɚɡɜɚɧɢɹ. ɒɬɚɦɦ ɝɚɥɨɛɚɤɬɟɪɢɣ, ɩɟɪɜɨɧɚɱɚɥɶɧɨ ɢɫɩɨɥɶɡɨɜɚɧɧɵɣ ɞɥɹ ɜɵɞɟɥɟɧɢɹ ɢ ɢɡɭɱɟɧɢɹ ȻɊ, ɛɵɥ ɩɨɥɭɱɟɧ ɢɡ ɤɨɥɥɟɤɰɢɢ ɤɭɥɶɬɭɪ ɩɨɞ ɧɚɡɜɚɧɢɟɦ Halobacterium halobium. Ɉɞɧɚɤɨ ɩɨɡɞɧɟɟ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɨɧ ɧɟɨɬɥɢɱɢɦ ɨɬ ɞɪɭɝɢɯ ɲɬɚɦɦɨɜ, ɧɚɩɪɢɦɟɪ, H. salinarium ɢ H. cutirubrum [27]. ȼ 1996 ɝ. ɛɵɥɚ ɜɧɨɜɶ ɩɟɪɟɫɦɨɬɪɟɧɚ ɧɨɦɟɧɤɥɚɬɭɪɚ ɝɚɥɨɛɚɤɬɟɪɢɣ [28] ɢ ɨɮɢɰɢɚɥɶɧɵɦ ɧɚɡɜɚɧɢɟɦ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɷɬɢɯ ɲɬɚɦɦɨɜ ɫɟɦɟɣɫɬɜɚ Halobacteriacea ɫɬɚɥɨ Halobacterium salinarum. ɂɫɯɨɞɧɨɟ ɜɢɞɨɜɨɟ ɧɚɡɜɚɧɢɟ salinarium ɛɵɥɨ ɢɡɦɟɧɟɧɨ ɧɚ salinarum, ɹɜɥɹɸɳɟɟɫɹ ɪɨɞɢɬɟɥɶɧɵɦ ɩɚɞɟɠɨɦ ɥɚɬɢɧɫɤɨɝɨ ɧɚɡɜɚɧɢɹ ɫɨɥɹɧɵɯ ɤɨɩɟɣ «salinae». Ȼɚɤɬɟɪɢɢ H. salinarum ɩɪɟɞɫɬɚɜɥɹɸɬ ɫɨɛɨɣ ɩɨɞɜɢɠɧɵɟ ɩɚɥɨɱɤɨɜɢɞɧɵɟ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ ɪɚɡɦɟɪɨɦ 0,5u5 ɦɤɦ [29], ɨɛɢɬɚɸɳɢɟ ɜ ɜɵɫɨɤɨɤɨɧɰɟɧɬɪɢɪɨɜɚɧɧɵɯ ɢɥɢ ɧɚɫɵɳɟɧɧɵɯ ɫɨɥɟɜɵɯ ɫɪɟɞɚɯ. Ɉɧɢ ɪɚɡɦɧɨɠɚɸɬɫɹ ɞɟɥɟɧɢɟɦ ɩɨɩɨɥɚɦ ɢ ɞɜɢɠɭɬɫɹ ɩɪɢ ɩɨɦɨɳɢ ɩɨɥɹɪɧɵɯ ɠɝɭɬɢɤɨɜ [3]. Ɂɪɟɥɵɟ ɤɥɟɬɤɢ ɫɨɯɪɚɧɹɸɬ ɩɚɥɨɱɤɨɜɢɞɧɭɸ ɮɨɪɦɭ ɜ 3,5—4 Ɇ NaCl, ɩɪɢ ɛɨɥɟɟ ɧɢɡɤɢɯ ɤɨɧɰɟɧɬɪɚɰɢɹɯ ɩɨɹɜɥɹɸɬɫɹ ɩɨɥɢɦɨɪɮɧɵɟ ɤɥɟɬɤɢ, ɚ ɩɪɢ ɤɨɧɰɟɧɬɪɚɰɢɢ NaCl ɪɚɜɧɨɣ 1.5 Ɇ ɤɥɟɬɤɢ ɫɬɚɧɨɜɹɬɫɹ ɫɮɟɪɢɱɟɫɤɢɦɢ ɜ ɪɟɡɭɥɶɬɚɬɟ ɧɚɛɭɯɚɧɢɹ ɢ ɭɬɪɚɬɵ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ ɢ ɱɚɫɬɢɱɧɨ ɥɢɡɢɪɭɸɬ. Ɍɚɤ ɠɟ ɩɨɥɧɨɟ ɪɚɡɪɭɲɟɧɢɟ ɤɥɟɬɨɱɧɵɯ ɫɬɪɭɤɬɭɪ ɩɪɨɢɫɯɨɞɢɬ ɩɪɢ ɨɞɧɨɜɪɟɦɟɧɧɨɦ ɭɦɟɧɶɲɟɧɢɢ ɤɨɧɰɟɧɬɪɚɰɢɢ NaCl ɢ

10 MgCl2 ɧɢɠɟ 2 Ɇ ɢ 20 ɦɆ ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ [3, 30], ɩɪɢɱɟɦ ɷɬɨ ɹɜɥɟɧɢɟ ɜ ɨɱɟɧɶ ɦɚɥɨɣ ɫɬɟɩɟɧɢ ɫɜɹɡɚɧɨ ɫ ɨɫɦɨɬɢɱɟɫɤɢɦ ɲɨɤɨɦ. Ɍɚɤɨɟ ɜɵɫɨɤɨɟ ɫɪɨɞɫɬɜɨ ɝɚɥɨɮɢɥɨɜ ɤ ɫɨɥɹɦ ɫɜɹɡɚɧɨ ɫ ɬɟɦ, ɱɬɨ ɛɟɥɤɢ, ɜɯɨɞɹɳɢɟ ɜ ɫɨɫɬɚɜ ɢɯ ɤɥɟɬɨɤ, ɫɨɞɟɪɠɚɬ ɦɧɨɝɨ ɤɢɫɥɵɯ ɚɦɢɧɨɤɢɫɥɨɬ ɢ ɦɚɥɨ ɧɟɩɨɥɹɪɧɵɯ ɚɦɢɧɨɤɢɫɥɨɬ [30], ɩɨɷɬɨɦɭ ɞɥɹ ɩɨɞɞɟɪɠɚɧɢɹ ɢɯ ɧɚɬɢɜɧɨɝɨ ɫɨɫɬɨɹɧɢɹ ɧɟɨɛɯɨɞɢɦɚ ɜɵɫɨɤɚɹ ɤɨɧɰɟɧɬɪɚɰɢɹ ɫɨɥɢ, ɤɨɬɨɪɚɹ ɩɪɟɩɹɬɫɬɜɭɟɬ ɢɯ ɚɝɪɟɝɚɰɢɢ [3]. ɉɪɢɪɨɞɧɵɟ ɲɬɚɦɦɵ H. salinarum ɢɦɟɸɬ ɠɟɥɬɭɸ ɢɥɢ ɨɪɚɧɠɟɜɭɸ ɨɤɪɚɫɤɭ, ɱɬɨ ɫɜɹɡɚɧɨ ɫ ɜɵɫɨɤɢɦ ɫɨɞɟɪɠɚɧɢɟɦ ɜ ɧɢɯ ɤɚɪɨɬɢɧɨɢɞɨɜ. Ʉɚɪɨɬɢɧɨɢɞɵ, ɹɜɥɹɸɳɢɟɫɹ ɨɫɧɨɜɧɵɦɢ ɩɢɝɦɟɧɬɚɦɢ ɤɥɟɬɨɤ ɝɚɥɨɛɚɤɬɟɪɢɣ, ɡɚɳɢɳɚɸɬ ɢɯ ɨɬ ɢɧɬɟɧɫɢɜɧɨɣ ɤɨɪɨɬɤɨɜɨɥɧɨɜɨɣ ɪɚɞɢɚɰɢɢ, ɤɨɬɨɪɨɣ ɨɧɢ ɩɨɞɜɟɪɠɟɧɵ ɜ ɟɫɬɟɫɬɜɟɧɧɵɯ ɭɫɥɨɜɢɹɯ ɨɛɢɬɚɧɢɹ [31]. ɋɨɞɟɪɠɚɧɢɟ ɤɚɪɨɬɢɧɨɢɞɨɜ, ɨɫɧɨɜɧɵɦ ɢɡ ɤɨɬɨɪɵɯ ɹɜɥɹɟɬɫɹ ɛɚɤɬɟɪɢɨɪɭɛɟɪɢɧ (ɫɦ. ɪɢɫ. 2) [32, 33], ɜ ɩɪɢɪɨɞɧɵɯ ɲɬɚɦɦɚɯ H. salinarum ɫɨɫɬɚɜɥɹɟɬ 7-10% ɨɬ ɫɭɦɦɚɪɧɨɣ ɥɢɩɢɞɧɨɣ ɮɪɚɤɰɢɢ [30]. ɉɟɪɜɨɧɚɱɚɥɶɧɨ [30] ɢɫɫɥɟɞɨɜɚɬɟɥɢ ɢɫɩɨɥɶɡɨɜɚɥɢ ɲɬɚɦɦ H. halobium R1, ɹɜɥɹɸɳɢɣɫɹ ɫɩɨɧɬɚɧɧɵɦ ɦɭɬɚɧɬɨɦ H. halobium NRC34020 ɢ ɧɟ ɨɛɪɚɡɭɸɳɢɣ ɝɚɡɨɜɵɯ ɜɚɤɭɨɥɟɣ, ɤɨɬɨɪɵɟ, ɤɚɤ ɢ ɩɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ (ɉɆ), ɭɫɬɨɣɱɢɜɵ ɜ ɨɬɫɭɬɫɬɜɢɢ ɫɨɥɟɣ ɢ ɦɟɲɚɸɬ ɢɯ ɜɵɞɟɥɟɧɢɸ. ɂɡ ɷɬɨɝɨ ɲɬɚɦɦɚ ɩɭɬɟɦ ɦɭɬɚɝɟɧɟɡɚ ɛɵɥɢ ɩɨɥɭɱɟɧɵ ɪɚɡɥɢɱɧɵɟ ɩɪɨɢɡɜɨɞɧɵɟ, ɜ ɱɚɫɬɧɨɫɬɢ, ɧɟ ɫɢɧɬɟɡɢɪɭɸɳɢɟ ɨɫɧɨɜɧɵɯ ɤɚɪɨɬɢɧɨɢɞɨɜ ɢ ɪɟɬɢɧɚɥɹ ɢɥɢ ɢɦɟɸɳɢɟ ɩɨɜɵɲɟɧɧɨɟ ɫɨɞɟɪɠɚɧɢɟ ȻɊ.

2. ɉɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ ɢ ɪɨɞɨɩɫɢɧɵ ɝɚɥɨɛɚɤɬɟɪɢɣ. ȼ 1966 ɝ. ɜ ɩɨɢɫɤɚɯ ɞɨɤɚɡɚɬɟɥɶɫɬɜɚ ɬɨɝɨ, ɱɬɨ ɦɟɦɛɪɚɧɵ ɢɦɟɸɬ ɛɢɫɥɨɣɧɭɸ ɫɬɪɭɤɬɭɪɭ ɫ ɜɧɟɞɪɟɧɧɵɦɢ ɜ ɧɟɟ ɛɟɥɤɚɦɢ, Oesterhelt D. et. al. [27] ɨɛɧɚɪɭɠɢɥɢ ɭɱɚɫɬɤɢ ɩɨɜɟɪɯɧɨɫɬɢ ɤɥɟɬɨɤ ɝɚɥɨɛɚɤɬɟɪɢɣ H. salinarum, ɜɵɝɥɹɞɟɜɲɢɟ ɩɨɞ ɷɥɟɤɬɪɨɧɧɵɦ ɦɢɤɪɨɫɤɨɩɨɦ ɛɨɥɟɟ ɝɥɚɞɤɢɦɢ, ɱɟɦ ɫɨɫɟɞɧɢɟ ɭɱɚɫɬɤɢ ɢ ɨɛɵɱɧɵɟ ɦɟɦɛɪɚɧɵ. ȼ 1971 ɝ. ɛɵɥɨ ɨɩɪɟɞɟɥɟɧɨ, ɱɬɨ ɜɯɨɞɹɳɢɣ ɜ ɢɯ ɫɨɫɬɚɜ ɛɟɥɨɤ ɫɯɨɞɟɧ ɩɨ ɫɬɪɭɤɬɭɪɟ ɫ ɪɨɞɨɩɫɢɧɨɦ [1, 34], ɩɨ ɷɬɨɣ ɩɪɢɱɢɧɟ ɨɧ ɛɵɥ ɧɚɡɜɚɧ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ. ɍɱɚɫɬɤɢ ɦɟɦɛɪɚɧ, ɫɨɞɟɪɠɚɳɢɟ ɛɨɥɶɲɨɟ ɤɨɥɢɱɟɫɬɜɨ ȻɊ ɛɥɚɝɨɞɚɪɹ ɯɚɪɚɤɬɟɪɧɨɣ ɨɤɪɚɫɤɟ ɛɵɥɢ ɧɚɡɜɚɧɵ ɩɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ (ɉɆ). ɉɆ ɫɨɞɟɪɠɚɬ ɫɚɦ ɛɟɥɨɤ (ɨɤɨɥɨ 75% ɩɨ ɫɭɯɨɦɭ ɜɟɫɭ), ɧɟɤɨɬɨɪɵɟ ɥɢɩɢɞɵ, ɤɚɪɨɬɢɧɨɢɞɵ ɢ ɜɨɞɭ [3, 29, 35]. Ɉɛɵɱɧɨ ɨɧɢ ɡɚɧɢɦɚɸɬ ɞɨ 50% ɩɨɜɟɪɯɧɨɫɬɢ ɤɥɟɬɤɢ ɢ ɢɦɟɸɬ ɨɤɪɭɝɥɭɸ ɮɨɪɦɭ [29, 35] ɢ ɪɚɡɦɟɪ ɨɤɨɥɨ 0,5 ɦɤɦ. ɋɭɦɦɢɪɭɹ ɪɟɡɭɥɶɬɚɬɵ ɢɫɫɥɟɞɨɜɚɧɢɹ ɫɜɨɣɫɬɜ ɉɆ, ɩɪɨɜɨɞɢɦɵɯ ɜ ɬɟɱɟɧɢɟ ɩɨɫɥɟɞɧɢɯ ɬɪɟɯ ɞɟɫɹɬɢɥɟɬɢɣ, ɦɨɠɧɨ ɨɬɦɟɬɢɬɶ ɫɥɟɞɭɸɳɢɟ ɢɯ ɨɫɨɛɟɧɧɨɫɬɢ [2, 36] (ɫɦ. ɬɚɤɠɟ ɬɚɛɥ. 1): x ɋɨɫɬɨɹɬ ɢɡ ɥɢɩɢɞɨɜ ɢ ȻɊ ɜ ɦɨɥɹɪɧɨɦ ɫɨɨɬɧɨɲɟɧɢɢ 10:1 ɢ ɜɟɫɨɜɨɦ 1:3 [37, 38];

11 x Ɉɛɪɚɡɭɸɬ ɞɜɭɦɟɪɧɭɸ ɝɟɤɫɚɝɨɧɚɥɶɧɭɸ ɤɪɢɫɬɚɥɥɢɱɟɫɤɭɸ ɪɟɲɟɬɤɭ ɢɡ ɨɞɢɧɚɤɨɜɨ ɨɪɢɟɧɬɢɪɨɜɚɧɧɵɯ ɬɪɢɦɟɪɨɜ ȻɊ; x ɂɦɟɸɬ ɫɪɟɞɧɸɸ ɩɥɨɬɧɨɫɬɶ 1.18 ɝ/ɫɦ3 ɢ ɩɨɤɚɡɚɬɟɥɶ ɩɪɟɥɨɦɥɟɧɢɹ 1.45 – 1.55; x ɂɦɟɸɬ ɪɚɡɦɟɪ ɨɤɨɥɨ 0,5 ɦɤɦ ɢ ɬɨɥɳɢɧɭ ɨɤɨɥɨ 5 ɧɦ; x Ɉɛɥɚɞɚɸɬ ɩɨɜɵɲɟɧɧɨɣ ɭɫɬɨɣɱɢɜɨɫɬɶɸ: o ɤ ɞɟɣɫɬɜɢɸ ɫɨɥɧɟɱɧɨɝɨ ɫɜɟɬɚ ɜ ɩɪɢɫɭɬɫɬɜɢɢ ɤɢɫɥɨɪɨɞɚ (ɭɫɬɨɣɱɢɜɵ ɜ ɬɟɱɟɧɢɟ ɧɟɫɤɨɥɶɤɢɯ ɥɟɬ); o ɤ ɞɟɣɫɬɜɢɸ ɩɨɜɵɲɟɧɧɵɯ ɬɟɦɩɟɪɚɬɭɪ (ɞɨ 80qɋ ɜ ɜɨɞɧɨɣ ɫɭɫɩɟɧɡɢɢ ɢ ɞɨ 140qɋ ɜ ɫɭɯɢɯ ɩɥɟɧɤɚɯ) [39]; o ɩɪɚɤɬɢɱɟɫɤɢ ɩɪɢ ɥɸɛɨɦ ɡɧɚɱɟɧɢɢ ɪɇ (ɨɬ 0 ɞɨ 12); o ɩɪɢ ɜɵɫɨɤɨɣ ɢɨɧɧɨɣ ɫɢɥɟ ɪɚɫɬɜɨɪɚ (3Ɇ NaCl); o ɤ ɞɟɣɫɬɜɢɸ ɦɧɨɝɢɯ ɩɪɨɬɟɚɡ [40, 41]; o ɤ ɞɟɣɫɬɜɢɸ ɧɟɩɨɥɹɪɧɵɯ ɪɚɫɬɜɨɪɢɬɟɥɟɣ (ɧɚɩɪɢɦɟɪ, ɝɟɤɫɚɧɚ), ɧɨ ɧɟɭɫɬɨɣɱɢɜɵ ɩɨ ɨɬɧɨɲɟɧɢɸ ɤ ɩɨɥɹɪɧɵɦ ɪɚɫɬɜɨɪɢɬɟɥɹɦ ɜ ɫɦɟɫɢ ɫ ɜɨɞɨɣ (ɷɬɚɧɨɥ, ɚɰɟɬɨɧ) [42] x ɋɨɯɪɚɧɹɸɬ ɨɤɪɚɫɤɭ ɢ ɮɨɬɨɯɢɦɢɱɟɫɤɭɸ ɚɤɬɢɜɧɨɫɬɶ ɩɪɢ ɩɪɚɤɬɢɱɟɫɤɢ ɩɨɥɧɨɦ ɭɞɚɥɟɧɢɢ ɜɨɞɵ; x ɋɭɳɟɫɬɜɭɟɬ ɜɨɡɦɨɠɧɨɫɬɶ ɜɫɬɪɚɢɜɚɧɢɹ ɉɆ ɜ ɩɨɥɢɦɟɪɧɵɟ ɦɚɬɪɢɰɵ ɛɟɡ ɩɨɬɟɪɢ ɮɨɬɨɯɢɦɢɱɟɫɤɢɯ ɫɜɨɣɫɬɜ [36]; ɏɢɦɢɱɟɫɤɢɣ ɚɧɚɥɢɡ [37] ɩɨɤɚɡɚɥ, ɱɬɨ ɩɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ ɫɨɫɬɨɹɬ ɧɚ 75% (ɩɨ ɫɭɯɨɦɭ ɜɟɫɭ) ɢɡ ɛɟɥɤɚ (ȻɊ) ɢ ɧɚ 25% ɢɡ ɮɨɫɮɨ- ɢ ɝɥɢɤɨɥɢɩɢɞɨɜ (ɪɢɫ. 2), ɱɬɨ, ɤɚɤ ɨɬɦɟɱɟɧɨ ɜɵɲɟ, ɫɨɨɬɜɟɬɫɬɜɭɟɬ ɩɪɢɛɥɢɡɢɬɟɥɶɧɨ ɞɟɫɹɬɢ ɦɨɥɟɤɭɥɚɦ ɥɢɩɢɞɨɜ ɧɚ ɨɞɧɭ ɦɨɥɟɤɭɥɭ ɛɟɥɤɚ. Ȼɨɥɟɟ ɬɨɱɧɨ ɥɢɩɢɞɧɵɣ ɫɨɫɬɚɜ ɉɆ ɛɵɥ ɨɩɪɟɞɟɥɟɧ ɩɪɢ ɩɨɦɨɳɢ ɦɟɬɨɞɚ əɆɊ ɧɚ ɹɞɪɚɯ 1ɇ ɢ 31Ɋ ɜ ɫɨɱɟɬɚɧɢɢ ɫ ɦɚɫɫ-ɫɩɟɤɬɪɨɦɟɬɪɢɟɣ [38], ɯɨɬɹ ɫɥɟɞɭɟɬ ɭɱɢɬɵɜɚɬɶ, ɱɬɨ ɨɧ ɦɨɠɟɬ ɧɟɫɤɨɥɶɤɨ ɢɡɦɟɧɹɬɶɫɹ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɭɫɥɨɜɢɣ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɝɚɥɨɛɚɤɬɟɪɢɣ. Ʌɢɩɢɞɵ ɨɛɥɚɞɚɸɬ ɯɚɪɚɤɬɟɪɧɨɣ ɞɥɹ ɝɚɥɨɛɚɤɬɟɪɢɣ ɨɫɨɛɟɧɧɨɫɬɶɸ: ɨɧɢ ɫɨɞɟɪɠɚɬ ɨɫɬɚɬɤɢ ɞɢɝɢɞɪɨɮɢɬɨɥɚ, ɩɪɢɫɨɟɞɢɧɟɧɧɵɯ ɩɪɨɫɬɨɣ ɷɮɢɪɧɨɣ ɫɜɹɡɶɸ, ɱɬɨ ɡɧɚɱɢɬɟɥɶɧɨ ɩɨɜɵɲɚɟɬ ɢɯ ɭɫɬɨɣɱɢɜɨɫɬɶ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɥɢɩɢɞɚɦɢ ɚɰɢɥɶɧɨɝɨ ɬɢɩɚ, ɜɫɬɪɟɱɚɸɳɢɦɢɫɹ ɭ ɨɫɬɚɥɶɧɵɯ ɠɢɜɵɯ ɨɪɝɚɧɢɡɦɨɜ.

12 HO

Ȼɚɤɬɟɪɢɨɪɭɛɟɪɢɧ

OH

ɇɟɣɬɪɚɥɶɧɵɟ ɥɢɩɢɞɵ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ O H

ɋɤɜɚɥɟɧ

O

8

Ɇɟɧɚɯɢɧɨɧ-8 (ɆɄ8) ɉɨɥɹɪɧɵɟ ɥɢɩɢɞɵ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ H 2C O O CH2 O CH CH2

CH2 OR CH OH O O P O CH2 O

3

HC O O P O O

R=

3

CH2

2-

PO3 SO3H H

Ɋɢɫ. 2. Ʉɚɪɨɬɢɧɨɢɞɵ ɢ ɥɢɩɢɞɵ, ɜɯɨɞɹɳɢɟ ɜ ɫɨɫɬɚɜ ɦɟɦɛɪɚɧ ɝɚɥɨɛɚɤɬɟɪɢɣ Ʉɪɨɦɟ ɩɭɪɩɭɪɧɵɯ, ɤɥɟɬɤɢ ɩɪɢɪɨɞɧɵɯ ɲɬɚɦɦɨɜ ɝɚɥɨɛɚɤɬɟɪɢɣ ɫɨɞɟɪɠɚɬ ɟɳɟ ɞɜɚ ɬɢɩɚ ɦɟɦɛɪɚɧ, ɭɫɬɨɣɱɢɜɵɯ ɩɪɢ ɨɬɫɭɬɫɬɜɢɢ ɫɨɥɟɣ, ɢ ɤɨɬɨɪɵɟ ɜɨɡɦɨɠɧɨ ɪɚɡɞɟɥɢɬɶ ɩɨ ɩɥɨɬɧɨɫɬɢ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɟɦ ɜ ɝɪɚɞɢɟɧɬɟ ɫɚɯɚɪɨɡɵ [37]. Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɷɬɢɯ ɦɟɦɛɪɚɧ ɩɪɟɞɫɬɚɜɥɟɧɵ ɜ ɬɚɛɥɢɰɟ 1. ɉɨɤɚɡɚɧɨ, ɱɬɨ ɤɨɪɢɱɧɟɜɵɟ ɦɟɦɛɪɚɧɵ ɫɨɞɟɪɠɚɬ ɛɚɤɬɟɪɢɨɨɩɫɢɧ ɜ ɫɦɟɫɢ ɫ ȻɊ (ɤɨɬɨɪɵɟ ɧɟ ɨɛɪɚɡɭɸɬ ɤɪɢɫɬɚɥɥɢɱɟɫɤɨɣ ɪɟɲɟɬɤɢ), ɰɢɬɨɯɪɨɦ ɢ ɦɢɧɨɪɧɨɟ ɤɨɥɢɱɟɫɬɜɨ ɞɪɭɝɢɯ ɛɟɥɤɨɜ. Ɉɧɢ ɹɜɥɹɸɬɫɹ ɩɪɟɞɲɟɫɬɜɟɧɧɢɤɚɦɢ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ [43], ɩɪɟɜɪɚɳɚɹɫɶ ɜ ɧɢɯ ɩɪɢ ɜɫɬɪɚɢɜɚɧɢɢ ɪɟɬɢɧɚɥɹ. Ɍɚɛɥɢɰɚ 1. Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɦɟɦɛɪɚɧ ɩɪɢɪɨɞɧɵɯ ɲɬɚɦɦɨɜ ɝɚɥɨɛɚɤɬɟɪɢɣ [37]

560

ɉɥɨɬɧɨɫɬɶ, ɝ/ɫɦ3 1,18

ɋɨɞɟɪɠɚɧɢɟ ɛɟɥɤɚ, % 75

Ʉɪɚɫɧɵɟ ɦɟɦɛɪɚɧɵ

340

1,06

5

ɥɢɤɨɩɢɧ, ɝɟɦ

Ʉɨɪɢɱɧɟɜɵɟ ɦɟɦɛɪɚɧɵ

419, 560

1,14

–

ɪɟɬɢɧɚɥɶ, ɝɟɦ

Ɍɢɩ

Omax, ɧɦ

ɉɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ

ɏɪɨɦɨɮɨɪɵ ɪɟɬɢɧɚɥɶ

Ʉ ɧɚɫɬɨɹɳɟɦɭ ɦɨɦɟɧɬɭ ɨɩɪɟɞɟɥɟɧɨ, ɱɬɨ ɤɥɟɬɤɢ H. salinarum ɫɨɞɟɪɠɚɬ ɢ ɞɪɭɝɢɟ ɪɨɞɨɩɫɢɧɵ, ɨɛɥɚɞɚɸɳɢɟ ɫɯɨɞɧɨɣ ɫ ȻɊ ɫɬɪɭɤɬɭɪɨɣ, ɧɨ ɜɵɩɨɥɧɹɸɳɢɟ ɞɪɭɝɢɟ ɮɭɧɤɰɢɢ.

13 ɗɬɨ ɞɜɚ ɫɟɧɫɨɪɧɵɯ ɪɨɞɨɩɫɢɧɚ, ɨɛɨɡɧɚɱɟɧɧɵɟ P565 (ɋɊ I) ɢ P370 (ɋɊ II), ɤɨɬɨɪɵɟ ɨɛɟɫɩɟɱɢɜɚɸɬ ɩɨɥɨɠɢɬɟɥɶɧɵɣ ɢ ɨɬɪɢɰɚɬɟɥɶɧɵɣ ɮɨɬɨɬɚɤɫɢɫ ɢ ɝɚɥɨɪɨɞɨɩɫɢɧ Ɋ578 (ȽɊ) [44]. Ɏɢɡɢɨɥɨɝɢɱɟɫɤɚɹ ɪɨɥɶ ȻɊ ɜ ɤɥɟɬɤɚɯ ɡɚɤɥɸɱɚɟɬɫɹ ɜ ɬɨɦ, ɱɬɨɛɵ ɨɛɟɫɩɟɱɢɜɚɬɶ ɝɚɥɨɛɚɤɬɟɪɢɹɦ ɜɨɡɦɨɠɧɨɫɬɶ ɫɭɳɟɫɬɜɨɜɚɧɢɹ ɤɚɤ ɮɚɤɭɥɶɬɚɬɢɜɧɵɦ ɚɧɚɷɪɨɛɚɦ ɜ ɫɥɭɱɚɟ, ɤɨɝɞɚ ɩɚɪɰɢɚɥɶɧɨɟ ɞɚɜɥɟɧɢɟ ɤɢɫɥɨɪɨɞɚ ɜ ɨɤɪɭɠɚɸɳɟɣ ɫɪɟɞɟ ɦɚɥɨ. Ȼɟɥɨɤ ɮɭɧɤɰɢɨɧɢɪɭɟɬ ɤɚɤ ɫɜɟɬɨɡɚɜɢɫɢɦɵɣ ɩɪɨɬɨɧɧɵɣ ɧɚɫɨɫ, ɨɛɟɫɩɟɱɢɜɚɸɳɢɣ ɨɛɪɚɡɨɜɚɧɢɟ ɷɥɟɤɬɪɨɯɢɦɢɱɟɫɤɨɝɨ ɝɪɚɞɢɟɧɬɚ ɩɪɨɬɨɧɨɜ ɧɚ ɩɨɜɟɪɯɧɨɫɬɢ ɦɟɦɛɪɚɧɵ ɤɥɟɬɤɢ, ɤɨɬɨɪɵɣ ɜ ɫɜɨɸ ɨɱɟɪɟɞɶ ɫɥɭɠɢɬ ɞɥɹ ɚɤɤɭɦɭɥɢɪɨɜɚɧɢɹ ɷɧɟɪɝɢɢ. ɉɟɪɜɢɱɧɚɹ ɪɚɛɨɬɚ, ɩɪɨɢɡɜɨɞɢɦɚɹ ɝɪɚɞɢɟɧɬɨɦ, ɡɚɤɥɸɱɚɟɬɫɹ ɜ ɫɢɧɬɟɡɟ ȺɌɎ - ɭɧɢɜɟɪɫɚɥɶɧɨɝɨ ɢɫɬɨɱɧɢɤɚ ɷɧɟɪɝɢɢ ɜ ɩɪɨɰɟɫɫɚɯ ɠɢɡɧɟɞɟɹɬɟɥɶɧɨɫɬɢ ɤɥɟɬɤɢ. Ʉɨɝɞɚ ɨɫɜɟɳɟɧɢɟ ɨɬɫɭɬɫɬɜɭɟɬ, ɚ ɩɚɪɰɢɚɥɶɧɨɟ ɞɚɜɥɟɧɢɟ ɤɢɫɥɨɪɨɞɚ ɜɵɫɨɤɨ, ɛɚɤɬɟɪɢɢ ɜɨɡɜɪɚɳɚɸɬɫɹ ɤ ɚɷɪɨɛɧɨɦɭ ɨɤɢɫɥɢɬɟɥɶɧɨɦɭ ɮɨɫɮɨɪɢɥɢɪɨɜɚɧɢɸ [29]. ȼ ɨɬɥɢɱɢɢ ɨɬ ȻɊ, ɋɊ I ɢ ɋɊ II ɜɨɫɩɪɢɧɢɦɚɸɬ ɫɜɟɬ ɫ ɪɚɡɥɢɱɧɨɣ ɞɥɢɧɨɣ ɜɨɥɧɵ ɢ ɜɵɡɵɜɚɸɬ ɤɚɫɤɚɞ ɫɢɝɧɚɥɨɜ, ɭɩɪɚɜɥɹɸɳɢɯ ɞɜɢɠɟɧɢɹɦɢ ɠɝɭɬɢɤɨɜ ɛɚɤɬɟɪɢɢ. ɉɪɢ ɩɨɦɨɳɢ ɬɚɤɨɝɨ ɷɥɟɦɟɧɬɚɪɧɨɝɨ ɩɪɨɰɟɫɫɚ ɫɜɟɬɨɜɨɝɨ ɜɨɫɩɪɢɹɬɢɹ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ ɫɚɦɨɫɬɨɹɬɟɥɶɧɨ ɩɟɪɟɦɟɳɚɸɬɫɹ ɜ ɨɛɥɚɫɬɢ, ɨɫɜɟɳɟɧɧɵɟ ɫɜɟɬɨɦ ɩɨɞɯɨɞɹɳɟɝɨ ɫɩɟɤɬɪɚɥɶɧɨɝɨ ɫɨɫɬɚɜɚ. Ƚɚɥɨɪɨɞɨɩɫɢɧɚ ɜ ɤɥɟɬɤɚɯ ɝɚɥɨɛɚɤɬɟɪɢɣ ɩɪɢɛɥɢɡɢɬɟɥɶɧɨ ɜ 200 ɪɚɡ ɦɟɧɶɲɟ, ɱɟɦ ȻɊ, ɟɝɨ ɦɨɥɟɤɭɥɹɪɧɵɣ ɜɟɫ ɫɨɫɬɚɜɥɹɟɬɨɤɨɥɨ 25 ɤȾɚ. Ɉɧ ɪɚɫɩɨɥɨɠɟɧ ɜ ɦɟɫɬɚɯ ɦɟɦɛɪɚɧɵ, ɝɞɟ ɧɟɬ ȻɊ. Ɉɫɧɨɜɧɚɹ ɮɭɧɤɰɢɹ ɝɚɥɨɪɨɞɨɩɫɢɧɚ – ɩɟɪɟɧɨɫ ɜ ɤɥɟɬɤɭ ɢɨɧɨɜ ɯɥɨɪɚ, ɤɨɬɨɪɵɟ ɩɨɫɬɨɹɧɧɨ ɬɟɪɹɸɬɫɹ ɛɚɤɬɟɪɢɟɣ ɩɨɞ ɞɟɣɫɬɜɢɟɦ ɷɥɟɤɬɪɢɱɟɫɤɨɝɨ ɩɨɥɹ, ɫɨɡɞɚɜɚɟɦɨɝɨ ȻɊ. ɉɪɟɞɩɨɥɚɝɚɟɬɫɹ, ɱɬɨ ɢɨɧ Cl – ɫɜɹɡɵɜɚɟɬɫɹ ɫ ɩɨɥɨɠɢɬɟɥɶɧɨ ɡɚɪɹɠɟɧɧɵɦ ɚɡɨɬɨɦ ɩɪɨɬɨɧɢɪɨɜɚɧɧɨɝɨ ɒɢɮɮɨɜɚ ɨɫɧɨɜɚɧɢɹ (SB), ɚ ɢɡɨɦɟɪɢɡɚɰɢɹ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɢɡ all-E ɜ 13Z-ɮɨɪɦɭ ɜɵɡɵɜɚɟɬ ɟɝɨ ɩɟɪɟɦɟɳɟɧɢɟ ɨɬ ɜɯɨɞɧɨɝɨ ɤ ɜɵɯɨɞɧɨɦɭ Cl –-ɩɪɨɜɨɞɹɳɟɦɭ ɩɨɥɭɤɚɧɚɥɭ, ɩɪɢɱɟɦ ɜ ɯɨɞɟ ɮɨɬɨɰɢɤɥɚ ɝɚɥɨɪɨɞɨɩɫɢɧɚ ɧɟ ɩɪɨɢɫɯɨɞɢɬ ɞɟɩɪɨɬɨɧɢɪɨɜɚɧɢɹ ɚɡɨɬɚ ɒɢɮɮɨɜɚ ɨɫɧɨɜɚɧɢɹ ɪɟɬɢɧɚɥɹ [2, 3]. ɇɚ ɪɢɫ. 3 ɩɪɟɞɫɬɚɜɥɟɧɨ ɫɯɟɦɚɬɢɱɟɫɤɨɟ ɢɡɨɛɪɚɠɟɧɢɟ ɜɡɚɢɦɨɫɜɹɡɚɧɧɵɯ ɫɢɫɬɟɦ ɠɢɡɧɟɞɟɹɬɟɥɶɧɨɫɬɢ ɝɚɥɨɛɚɤɬɟɪɢɣ, ɨɫɧɨɜɚɧɧɵɯ ɧɚ ɪɚɡɥɢɱɧɵɯ ɬɢɩɚɯ ɛɚɤɬɟɪɢɚɥɶɧɵɯ ɪɨɞɨɩɫɢɧɨɜ [45]. Ʉɪɚɬɤɨ ɷɬɭ ɫɢɫɬɟɦɭ ɦɨɠɧɨ ɨɩɢɫɚɬɶ ɫɥɟɞɭɸɳɢɦ ɨɛɪɚɡɨɦ: ɫɨɡɞɚɜɚɟɦɚɹ ɞɵɯɚɬɟɥɶɧɨɣ ɫɢɫɬɟɦɨɣ (ɩɪɢ ɜɵɫɨɤɨɣ ɤɨɧɰɟɧɬɪɚɰɢɢ ɤɢɫɥɨɪɨɞɚ) ɢɥɢ ȻɊ (ɜ ɭɫɥɨɜɢɹɯ ɧɟɞɨɫɬɚɬɨɱɧɨɣ ɚɷɪɚɰɢɢ) ɪɚɡɧɨɫɬɶ ɩɨɬɟɧɰɢɚɥɨɜ ɧɚ ɦɟɦɛɪɚɧɟ ɢɫɩɨɥɶɡɭɟɬɫɹ ɞɥɹ ɫɢɧɬɟɡɚ ȺɌɎ, ɩɟɪɟɧɨɫɚ ɢɨɧɨɜ ɤɚɥɢɹ ɜ ɤɥɟɬɤɭ, ɚ ɢɨɧɨɜ ɧɚɬɪɢɹ – ɜɨ ɜɧɟɲɧɸɸ ɫɪɟɞɭ. ɋɨɡɞɚɜɚɟɦɵɣ ɬɚɤɢɦ ɨɛɪɚɡɨɦ ɝɪɚɞɢɟɧɬ ɢɨɧɨɜ ɧɚɬɪɢɹ ɢɫɩɨɥɶɡɭɟɬɫɹ ɜ ɫɜɨɸ ɨɱɟɪɟɞɶ ɞɥɹ ɩɟɪɟɧɨɫɚ ɚɦɢɧɨɤɢɫɥɨɬ ɢ ɢɨɧɨɜ ɯɥɨɪɚ ɜ ɤɥɟɬɤɭ (ɪɢɫ. 3Ⱥ). ɋɢɝɧɚɥɵ, ɫɨɡɞɚɜɚɟɦɵɟ ɋɊ I ɢ

14 ɋɊ II, ɩɟɪɟɞɚɸɬɫɹ ɧɚ ɠɝɭɬɢɤɢ, ɨɛɟɫɩɟɱɢɜɚɸɳɢɟ ɞɜɢɠɟɧɢɟ ɤɥɟɬɤɢ ɜ ɧɚɩɪɚɜɥɟɧɢɢ ɧɚɢɛɨɥɟɟ ɛɥɚɝɨɩɪɢɹɬɧɵɯ ɭɫɥɨɜɢɣ ɨɫɜɟɳɟɧɢɹ. ɉɪɢ ɷɬɨɦ ɞɜɢɠɟɧɢɟ ɠɝɭɬɢɤɨɜ, ɤɚɤ ɢ ɛɢɨɫɢɧɬɟɬɢɱɟɫɤɢɟ ɩɪɨɰɟɫɫɵ ɨɫɭɳɟɫɬɜɥɹɸɬɫɹ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ȺɌɎ (ɪɢɫ. 3ȼ). A)

B) hQ

hQ Cl

+ H

H+ ȺȾɎ

-

hQ

+ H

ɋɟɧɫɨɪɧɵɟ ɪɨɞɨɩɫɢɧɵ I ɢ II

H+ + K

H+

ɉɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ

ATP

ADP

ȺɌɎ Na+

Na+ Cl-

H+

ɠɝɭɬɢɤɢ ȺɌɎ-ɫɢɧɬɟɬɚɡɚ

H+

+ ȺɄ Na

Ɋɢɫ. 3. ɋɯɟɦɵ ɦɟɦɛɪɚɧɧɵɯ ɩɪɨɰɟɫɫɨɜ ɤɥɟɬɤɢ ɝɚɥɨɛɚɤɬɟɪɢɢ, ɫɜɹɡɚɧɧɵɯ ɫ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɟɦ ɪɨɞɨɩɫɢɧɨɜ [45]. A) Ɍɪɚɧɫɩɨɪɬɧɵɟ ɫɢɫɬɟɦɵ ɞɥɹ ɢɨɧɨɜ: 1-ɩɟɪɟɧɨɫ ɩɪɨɬɨɧɚ ɢɡ ɤɥɟɬɤɢ ɫ ɩɨɦɨɳɶɸ ɞɵɯɚɬɟɥɶɧɨɣ ɫɢɫɬɟɦɵ; 2- ȻɊ; 3- ȺɌɎ-ɫɢɧɬɟɬɚɡɚ; +

+

4 – Na /H ɩɟɪɟɧɨɫɱɢɤ (ɚɧɬɢɩɨɪɬɟɪ); 5 – ɬɪɚɧɫɩɨɪɬ ɚɦɢɧɨɤɢɫɥɨɬ, ɡɚɜɢɫɢɦɵɣ ɨɬ +

ɝɪɚɞɢɟɧɬɚ ɤɨɧɰɟɧɬɪɚɰɢɢ Na ; 6 – ɬɪɚɧɫɩɨɪɬ ɢɨɧɨɜ ɤɚɥɢɹ, ɡɚɜɢɫɢɦɵɣ ɨɬ ɦɟɦɛɪɚɧɧɨɝɨ +

ɩɨɬɟɧɰɢɚɥɚ; 7 – ɫɜɟɬɨɧɟɡɚɜɢɫɢɦɚɹ ɫɢɫɬɟɦɚ ɩɟɪɟɧɨɫɚ ɢɨɧɨɜ ɯɥɨɪɚ (ɫɨɜɦɟɫɬɧɨ ɫ Na ); 8 – ɝɚɥɨɪɨɞɨɩɫɢɧ. B) ɋɢɫɬɟɦɚ, ɨɛɟɫɩɟɱɢɜɚɸɳɚɹ ɞɜɢɠɟɧɢɟ ɝɚɥɨɛɚɤɬɟɪɢɣ. 3. Ȼɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ: ɫɬɪɭɤɬɭɪɚ ɢ ɮɭɧɤɰɢɢ. Ɋɨɞɨɩɫɢɧɵ - ɛɟɥɤɢ, ɫɨɞɟɪɠɚɳɢɟ ɨɫɬɚɬɨɤ ɪɟɬɢɧɚɥɹ. Ɉɧɢ ɩɪɟɞɫɬɚɜɥɹɸɬ ɢɧɬɟɪɟɫ ɞɥɹ ɢɫɫɥɟɞɨɜɚɬɟɥɟɣ, ɩɨɫɤɨɥɶɤɭ ɨɧɢ ɹɜɥɹɸɬɫɹ ɧɚɢɛɨɥɟɟ ɩɪɨɫɬɨ ɭɫɬɪɨɟɧɧɵɦɢ ɩɪɢɪɨɞɧɵɦɢ ɩɪɟɨɛɪɚɡɨɜɚɬɟɥɹɦɢ ɷɧɟɪɝɢɢ ɫɜɟɬɚ ɜ ɷɥɟɤɬɪɨɯɢɦɢɱɟɫɤɭɸ. ɏɪɨɧɨɥɨɝɢɱɟɫɤɢ ɩɟɪɜɵɦ ɛɵɥ ɨɩɢɫɚɧ ɡɪɢɬɟɥɶɧɵɣ ɪɨɞɨɩɫɢɧ ɜɯɨɞɹɳɢɣ ɜ ɫɨɫɬɚɜ ɩɚɥɨɱɟɤ ɫɟɬɱɚɬɤɢ ɝɥɚɡɚ. ɍɪɨɜɟɧɶ ɢɫɫɥɟɞɨɜɚɧɢɣ ȻɊ ɧɚ ɫɟɝɨɞɧɹ ɜɵɲɟ, ɱɟɦ ɡɪɢɬɟɥɶɧɨɝɨ ɪɨɞɨɩɫɢɧɚ, ɯɨɬɹ ɩɨɫɥɟɞɧɢɣ ɢɫɫɥɟɞɭɟɬɫɹ ɛɨɥɟɟ 100 ɥɟɬ ɚ ȻɊ – ɦɟɧɟɟ 40 ɥɟɬ, ɱɬɨ ɦɨɠɧɨ ɫɜɹɡɚɬɶ ɫ ɩɪɨɫɬɨɬɨɣ ɩɨɥɭɱɟɧɢɹ ȻɊ ɜ ɛɨɥɶɲɢɯ ɤɨɥɢɱɟɫɬɜɚɯ, ɚ ɬɚɤɠɟ ɜɨɡɧɢɤɧɨɜɟɧɢɟɦ ɩɟɪɫɩɟɤɬɢɜ ɟɝɨ ɬɟɯɧɨɥɨɝɢɱɟɫɤɨɝɨ ɢɫɩɨɥɶɡɨɜɚɧɢɹ. ȻɊ – ɪɟɬɢɧɚɥɶ-ɫɨɞɟɪɠɚɳɢɣ ɩɪɨɬɨɧɧɵɣ ɧɚɫɨɫ ɦɟɦɛɪɚɧ ɝɚɥɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ ɪɨɞɚ Halobacterium, ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɬɪɚɧɫɦɟɦɛɪɚɧɧɵɣ ɛɟɥɨɤ ɫ ɦɨɥɟɤɭɥɹɪɧɵɦ ɜɟɫɨɦ 26 ɤȾɚ, ɫɨɫɬɨɹɳɢɣ ɢɡ 248 ɚɦɢɧɨɤɢɫɥɨɬ ɢ ɩɪɨɧɢɡɵɜɚɸɳɢɣ ɦɟɦɛɪɚɧɭ ɜ ɜɢɞɟ ɫɟɦɢ D - ɫɩɢɪɚɥɶɧɵɯ ɬɹɠɟɣ. D-ɋɩɢɪɚɥɶɧɵɟ ɭɱɚɫɬɤɢ ɪɚɫɩɨɥɨɠɟɧɵ ɜ ɦɟɦɛɪɚɧɟ, ɚ ɧɟɫɩɢɪɚɥɢɡɨɜɚɧɧɵɟ ɜɵɫɬɭɩɚɸɬ ɡɚ ɟɟ ɩɪɟɞɟɥɵ, ɨɛɪɚɡɭɹ ɩɟɬɥɢ. N- ɢ C-ɤɨɧɰɵ ɩɨɥɢɩɟɩɬɢɞɧɨɣ ɰɟɩɢ ɧɚɯɨɞɹɬɫɹ ɩɨ ɪɚɡɧɵɟ ɫɬɨɪɨɧɵ ɰɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɨɣ ɦɟɦɛɪɚɧɵ: Nɤɨɧɟɰ ɨɛɪɚɳɟɧ ɜɨ ɜɧɟɲɧɸɸ ɫɪɟɞɭ, ɚ C-ɤɨɧɟɰ – ɜɧɭɬɪɶ ɤɥɟɬɤɢ (ɪɢɫ. 4) [46].

15 ɍɩɚɤɨɜɚɧɧɵɟ ɬɚɤɢɦ ɨɛɪɚɡɨɦ ɦɨɥɟɤɭɥɵ ȻɊ ɨɪɝɚɧɢɡɨɜɚɧɵ ɜ ɬɪɢɦɟɪɵ, ɤɨɬɨɪɵɟ ɞɚɥɟɟ ɨɛɪɚɡɭɸɬ ɤɪɢɫɬɚɥɥɢɱɟɫɤɭɸ ɝɟɤɫɚɝɨɧɚɥɶɧɭɸ ɫɬɪɭɤɬɭɪɭ. ɏɪɨɦɨɮɨɪɨɦ ȻɊ (ɫɦ. ɪɢɫ. 4) ɹɜɥɹɟɬɫɹ ɩɪɨɬɨɧɢɪɨɜɚɧɧɵɣ ɚɥɶɞɢɦɢɧ ɪɟɬɢɧɚɥɹ ɫ İɚɦɢɧɨɝɪɭɩɩɨɣ ɨɫɬɚɬɤɚ Lys-216, ɤɨɬɨɪɵɣ ɪɚɡɦɟɳɟɧ ɜ ɝɢɞɪɨɮɨɛɧɨɣ ɱɚɫɬɢ ɦɨɥɟɤɭɥɵ. ɉɨɫɥɟ ɩɨɝɥɨɳɟɧɢɹ ɤɜɚɧɬɚ ɫɜɟɬɚ ɩɪɨɢɫɯɨɞɢɬ ɢɡɨɦɟɪɢɡɚɰɢɹ ɪɟɬɢɧɚɥɹ ɢɡ all-E- ɜ 13Zɮɨɪɦɭ. Ȼɟɥɤɨɜɨɟ ɦɢɤɪɨɨɤɪɭɠɟɧɢɟ ɯɪɨɦɨɮɨɪɚ ɦɨɠɧɨ ɪɚɫɫɦɚɬɪɢɜɚɬɶ ɤɚɤ ɪɟɰɟɩɬɨɪ ɫ ɫɭɛɫɬɪɚɬɧɨɣ ɫɩɟɰɢɮɢɱɧɨɫɬɶɸ ɞɥɹ all-E- ɢ 13Z- ɢɡɨɦɟɪɨɜ ɪɟɬɢɧɚɥɹ, ɤɨɬɨɪɵɣ ɩɨɞɚɜɥɹɟɬ ɢɡɨɦɟɪɢɡɚɰɢɸ ɩɨ ɞɪɭɝɢɦ ɞɜɨɣɧɵɦ ɫɜɹɡɹɦ, ɧɚɩɪɢɦɟɪ ɨɬ all-E- ɤ 7Z-, 9Z- ɢɥɢ 11Zɢɡɨɦɟɪɚɦ. Ɉɫɬɚɥɶɧɚɹ ɱɚɫɬɶ ɩɨɥɢɩɟɩɬɢɞɧɨɣ ɰɟɩɢ ɨɛɟɫɩɟɱɢɜɚɟɬ ɤɚɧɚɥ ɩɪɨɬɨɧɧɨɝɨ ɬɪɚɧɫɩɨɪɬɚ ɢɥɢ ɷɤɪɚɧɢɪɭɟɬ ɯɪɨɦɨɮɨɪ ɨɬ ɜɥɢɹɧɢɣ ɜɧɟɲɧɟɣ ɫɪɟɞɵ.

Ɋɢɫ. 4. ɋɯɟɦɚɬɢɱɟɫɤɚɹ ɦɨɞɟɥɶ ɬɪɟɯɦɟɪɧɨɣ ɫɬɪɭɤɬɭɪɵ ȻɊ [27]. ɋɟɦɶ D-ɫɩɢɪɚɥɟɣ ɮɨɪɦɢɪɭɸɬ ɯɪɨɦɨɮɨɪɧɭɸ ɩɨɥɨɫɬɶ ɢ ɬɪɚɧɫɦɟɦɛɪɚɧɧɵɣ ɤɚɧɚɥ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ. ɉɨɤɚɡɚɧɨ ɩɨɥɨɠɟɧɢɟ ɨɫɬɚɬɤɨɜ Asp85, Asp96, Asp212 ɢ Arg82, ɤɨɬɨɪɵɟ ɩɪɢɧɢɦɚɸɬ ɭɱɚɫɬɢɟ ɜ ɩɪɨɰɟɫɫɟ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ. ɇɟɫɦɨɬɪɹ ɧɚ ɤɚɠɭɳɭɸɫɹ ɩɪɨɫɬɨɬɭ, ɨɱɟɜɢɞɧɨ, ɱɬɨ ȻɊ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ +

ɞɨɫɬɚɬɨɱɧɨ ɫɥɨɠɧɭɸ ɫɢɫɬɟɦɭ. ɉɪɟɠɞɟ ɜɫɟɝɨ, ɩɭɬɶ, ɤɨɬɨɪɵɣ ɞɨɥɠɟɧ ɩɪɨɣɬɢ ɇ , ɱɬɨɛɵ ɩɟɪɟɫɟɱɶ ɦɟɦɛɪɚɧɭ, ɫɨɫɬɚɜɥɹɟɬ ɧɟ ɦɟɧɟɟ 5 ɧɦ, ɬ. ɟ. ɡɧɚɱɢɬɟɥɶɧɨ ɩɪɟɜɵɲɚɟɬ ɪɚɫɫɬɨɹɧɢɟ, ɧɚ ɤɨɬɨɪɨɟ ɨɧ ɦɨɠɟɬ ɛɵɬɶ ɩɟɪɟɧɟɫɟɧ ɩɪɢ ɢɡɨɦɟɪɢɡɚɰɢɢ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ. ɗɬɨ ɨɡɧɚɱɚɟɬ, ɱɬɨ ɩɨɝɥɨɳɟɧɢɟ ɤɜɚɧɬɚ ɫɜɟɬɚ ɞɨɥɠɧɨ ɩɪɢɜɨɞɢɬɶ ɤ ɜɨɡɧɢɤɧɨɜɟɧɢɸ ɧɚɩɪɹɠɟɧɧɨɣ

16 ɤɨɧɮɨɪɦɚɰɢɢ ɜɫɟɣ ɦɨɥɟɤɭɥɵ ȻɊ, ɫɥɭɠɚɳɟɣ ɜ ɞɚɥɶɧɟɣɲɟɦ ɢɫɬɨɱɧɢɤɨɦ ɷɧɟɪɝɢɢ ɞɥɹ +

ɩɟɪɟɧɨɫɚ ɇ ɩɪɨɬɢɜ ɷɥɟɤɬɪɨɯɢɦɢɱɟɫɤɨɝɨ ɝɪɚɞɢɟɧɬɚ. ȼ ɨɪɝɚɧɢɡɚɰɢɢ ɬɚɤɨɝɨ ɩɟɪɟɧɨɫɚ ɩɪɢɧɢɦɚɸɬ ɭɱɚɫɬɢɟ D-ɫɩɢɪɚɥɶɧɵɟ ɬɹɠɢ, ɮɨɪɦɢɪɭɸɳɢɟ ɞɜɟ ɱɚɫɬɢ ɩɪɨɬɨɧɧɨɝɨ ɤɚɧɚɥɚ, ɞɟɬɚɥɶɧɵɣ ɦɟɯɚɧɢɡɦ ɞɟɣɫɬɜɢɹ ɤɨɬɨɪɵɯ ɩɨɤɚ ɧɟ ɢɡɜɟɫɬɟɧ. ɂɧɞɭɰɢɪɨɜɚɧɧɵɣ ɫɜɟɬɨɦ ɩɪɨɬɨɧɧɵɣ ɬɪɚɧɫɩɨɪɬ ɫɨɩɪɨɜɨɠɞɚɟɬɫɹ ɪɹɞɨɦ ɤɨɧɮɨɪɦɚɰɢɨɧɧɵɯ ɢɡɦɟɧɟɧɢɣ ɦɨɥɟɤɭɥɵ ȻɊ, ɱɬɨ ɩɪɢɜɨɞɢɬ ɤ ɰɢɤɥɢɱɟɫɤɢɦ ɫɩɟɤɬɪɚɥɶɧɵɦ ɢɡɦɟɧɟɧɢɹɦ ɩɨɝɥɨɳɟɧɢɹ ȻɊ ɜ ɜɢɞɢɦɨɣ ɢ ɍɎ ɨɛɥɚɫɬɹɯ ɫɩɟɤɬɪɚ, ɫɨɜɨɤɭɩɧɨɫɬɶ ɤɨɬɨɪɵɯ ɧɚɡɵɜɚɟɬɫɹ ɮɨɬɨɰɢɤɥɨɦ2 (ɪɢɫ. 5) [27, 47].

B-ɮɨɪɦɚ

ȻɊ568

0,5*10-12 c hQ

J625

8*10-3 c

O640

3*10-12 c

K590 10-6 c

-4

10 c

N560 10-4 c

L550

M-ɮɨɪɦɚ II

M412

10-4 c

I

M412

5*10-5 c

Ɋɢɫ. 5. ɉɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶ ɢɧɬɟɪɦɟɞɢɚɬɨɜ ɜ ɮɨɬɨɰɢɤɥɟ ȻɊ [48, 49]. Ɏɨɬɨɰɢɤɥ ȻɊ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶ ɢɧɬɟɪɦɟɞɢɚɬɨɜ (ɫɦ. ɪɢɫ. 5), ɤɨɬɨɪɵɟ ɛɵɥɢ ɨɛɧɚɪɭɠɟɧɵ ɩɭɬɟɦ ɢɯ ɫɬɚɛɢɥɢɡɚɰɢɢ ɩɪɢ ɩɨɧɢɠɟɧɧɨɣ ɬɟɦɩɟɪɚɬɭɪɟ ɢ ɦɨɝɭɬ ɛɵɬɶ ɨɯɚɪɚɤɬɟɪɢɡɨɜɚɧɵ ɤɚɤ ɦɚɤɫɢɦɭɦɚɦɢ ɩɨɝɥɨɳɟɧɢɹ, ɬɚɤ ɢ ɤɢɧɟɬɢɤɨɣ ɨɛɪɚɡɨɜɚɧɢɹ ɢ ɪɚɫɩɚɞɚ. ɋɱɢɬɚɟɬɫɹ, ɱɬɨ ɜ ɮɨɬɨɰɢɤɥɟ ȻɊ ɬɨɥɶɤɨ ɩɟɪɜɚɹ ɫɬɚɞɢɹ ɜɵɡɵɜɚɟɬɫɹ ɩɨɝɥɨɳɟɧɢɟɦ ɤɜɚɧɬɚ ɫɜɟɬɚ (ɩɟɪɟɯɨɞ BoJ), ɨɫɬɚɥɶɧɵɟ ɩɟɪɟɯɨɞɵ ɹɜɥɹɸɬɫɹ ɬɟɦɧɨɜɵɦɢ ɩɪɨɰɟɫɫɚɦɢ, ɩɪɢɱɟɦ ɧɟɬ ɟɞɢɧɨɝɨ ɦɧɟɧɢɹ ɨɛ ɢɯ ɨɛɪɚɬɢɦɨɫɬɢ. ɂɡɜɟɫɬɧɵɟ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɢɧɬɟɪɦɟɞɢɚɬɨɜ ɮɨɬɨɰɢɤɥɚ ȻɊ [2, 17, 47, 50, 51] ɩɪɢɜɟɞɟɧɵ ɜ ɬɚɛɥɢɰɟ 2. ɉɪɢ ɨɬɫɭɬɫɬɜɢɢ ɨɫɜɟɳɟɧɢɹ ȻɊ ɧɚɯɨɞɢɬɫɹ ɜ ɬɟɦɧɨɚɞɚɩɬɢɪɨɜɚɧɧɨɦ ɫɨɫɬɨɹɧɢɢ

2

Ʉɪɨɦɟ ɧɟɝɨ ɫɭɳɟɫɬɜɭɟɬ ɪɹɞ ɞɪɭɝɢɯ ɰɢɤɥɨɜ, ɧɚɩɪɢɦɟɪ, ɰɢɤɥ ɩɟɪɟɦɟɳɟɧɢɹ ɡɚɪɹɞɚ, ɫɬɪɭɤɬɭɪɧɵɣ ɰɢɤɥ ɢ ɞɪ., ɧɨ ɬɟɪɦɢɧ «ɮɨɬɨɰɢɤɥ» ɩɪɢɦɟɧɹɬ ɩɨ ɨɬɧɨɲɟɧɢɸ ɬɨɥɶɤɨ ɤ ɰɢɤɥɭ ɫɩɟɤɬɪɚɥɶɧɵɯ ɢɡɦɟɧɟɧɢɣ.

17 (DA) ɫ ɦɚɤɫɢɦɭɦɨɦ ɩɨɝɥɨɳɟɧɢɹ ɨɤɨɥɨ 558 ɧɦ, ɤɨɬɨɪɨɟ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɫɦɟɫɶ 66:34 ɮɨɪɦ ȻɊ ɫ ɤɨɧɮɨɪɦɚɰɢɟɣ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ all-E- (Omax 568 ɧɦ) ɢ 13Z(Omax 548 ɧɦ) [52, 53]. ɉɪɢ ɨɫɜɟɳɟɧɢɢ ɩɪɨɢɫɯɨɞɢɬ ɫɜɟɬɨɜɚɹ ɚɞɚɩɬɚɰɢɹ, ɬ.ɟ. ɩɟɪɟɯɨɞ >97% ɦɨɥɟɤɭɥ ȻɊ ɜ ɫɨɫɬɨɹɧɢɟ ɫ all-E- ɤɨɧɮɢɝɭɪɚɰɢɟɣ ɪɟɬɢɧɚɥɹ, ɱɬɨ ɫɨɩɪɨɜɨɠɞɚɟɬɫɹ ɫɞɜɢɝɨɦ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ɩɪɟɩɚɪɚɬɚ ɫ 558 ɞɨ 568 ɧɦ ɢ ɧɟɤɨɬɨɪɵɦ ɭɜɟɥɢɱɟɧɢɟɦ ɷɤɫɬɢɧɤɰɢɢ. ɋɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ, ɱɬɨ 13Z-ɮɨɪɦɚ ȻɊ ɬɚɤɠɟ ɜ ɨɬɜɟɬ ɧɚ ɩɨɝɥɨɳɟɧɢɟ ɤɜɚɧɬɚ ɫɜɟɬɚ ɩɪɟɬɟɪɩɟɜɚɟɬ ɮɨɬɨɰɢɤɥ, ɨɞɧɚɤɨ ɨɧ ɧɟ ɫɨɩɪɨɜɨɠɞɚɟɬɫɹ ɩɟɪɟɧɨɫɨɦ ɩɪɨɬɨɧɚ, ɚ ɩɪɢɜɨɞɢɬ ɤ ɜɨɡɜɪɚɳɟɧɢɸ ɜ ɢɫɯɨɞɧɨɟ ɫɨɫɬɨɹɧɢɟ ɢɥɢ ɨɛɪɚɡɨɜɚɧɢɸ all-E-ɮɨɪɦɵ ȻɊ. Ɍɚɛɥɢɰɚ 2. Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɢɧɬɟɪɦɟɞɢɚɬɨɜ ɮɨɬɨɰɢɤɥɚ ȻɊ [2, 17, 47, 50, 51]. ɉɪɨɬɨɧɢɪɨɜɚɧɢɟ ɤɥɸɱɟɜɵɯ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ SB Asp85 Asp212 Asp115 Tyr185 568 63000 H H — — H — all-E 590 52100 H H — — H H 13Z 550 48900 H H — — H H 13Z 412 48800 H — H H H — 13Z 560 46100 — H H H 13Z 640 61900 H H H all-E * Tɫɬ - ɬɟɦɩɟɪɚɬɭɪɚ, ɩɪɢ ɤɨɬɨɪɨɣ ɢɧɬɟɪɦɟɞɢɚɬ ɬɟɪɦɢɱɟɫɤɢ ɫɬɚɛɢɥɢɡɢɪɨɜɚɧ. Omax, ɧɦ

B K L M N O

H, Ɇ-1·ɫɦ-1

Ʉɨɧɮɢɝɭɪɚɰɢɹ ɪɟɬɢɧɚɥɹ Asp96

Tɫɬ*, 0 ɋ

-190 -100 -30 -60

ɋɭɦɦɢɪɭɹ ɩɪɨɜɟɞɟɧɧɵɟ ɢɫɫɥɟɞɨɜɚɧɢɹ [2, 47, 50, 51, 54], ɩɪɨɰɟɫɫ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ ȻɊ ɦɨɠɧɨ ɨɩɢɫɚɬɶ ɫɥɟɞɭɸɳɢɦ ɨɛɪɚɡɨɦ: ɜ ɢɫɯɨɞɧɨɦ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɨɦ ɫɨɫɬɨɹɧɢɢ (B) ɚɥɶɞɢɦɢɧɧɚɹ ɝɪɭɩɩɚ ɯɪɨɦɨɮɨɪɚ ȻɊ ɧɚɯɨɞɢɬɫɹ ɜ ɩɪɨɬɨɧɢɪɨɜɚɧɧɨɦ ɫɨɫɬɨɹɧɢɢ, ɪɹɞɨɦ ɫ ɧɟɣ ɪɚɫɩɨɥɨɠɟɧ ɤɨɦɩɥɟɤɫɧɵɣ ɩɪɨɬɢɜɨɢɨɧ, ɨɛɪɚɡɭɟɦɵɣ ɨɫɬɚɬɤɚɦɢ Asp85, Asp212 ɢ H2O. ɉɪɢ ɩɨɝɥɨɳɟɧɢɢ ɤɜɚɧɬɚ ɫɜɟɬɚ ɩɪɨɢɫɯɨɞɢɬ all-Eo13Z ɢɡɨɦɟɪɢɡɚɰɢɹ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ, ɤɨɬɨɪɚɹ ɩɟɪɟɦɟɳɚɟɬ ɚɡɨɬ ɚɥɶɞɢɦɢɧɧɨɣ ɫɜɹɡɢ ɩɨ ɧɚɩɪɚɜɥɟɧɢɸ ɤ ɢɨɧɢɡɨɜɚɧɧɵɦ ɨɫɬɚɬɤɚɦ Asp85 ɢ Asp212. Ɍɚɤɨɟ ɩɟɪɟɦɟɳɟɧɢɟ ɡɚɪɹɠɟɧɧɵɯ ɮɪɚɝɦɟɧɬɨɜ ɦɨɥɟɤɭɥɵ ɫɥɭɠɢɬ ɨɞɧɨɣ ɢɡ ɩɪɢɱɢɧ ɡɧɚɱɢɬɟɥɶɧɨɝɨ ɛɚɬɨɯɪɨɦɧɨɝɨ ɫɞɜɢɝɚ Omax ɢɧɬɟɪɦɟɞɢɚɬɨɜ J ɢ K (ɨɬ ȼ568 ɞɨ J625 ɢ Ʉ590). ɉɪɢ ɩɟɪɟɯɨɞɟ ɤ ɢɧɬɟɪɦɟɞɢɚɬɭ L ɩɪɨɢɫɯɨɞɹɬ ɧɟɛɨɥɶɲɢɟ ɤɨɧɮɨɪɦɚɰɢɨɧɧɵɟ ɢɡɦɟɧɟɧɢɹ ɛɟɥɤɨɜɨɣ ɱɚɫɬɢ ɫɧɚɱɚɥɚ ɜ ɪɚɣɨɧɟ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ, ɡɚɬɟɦ SB ɪɟɬɢɧɢɥɢɞɟɧɨɜɨɝɨ ɨɫɬɚɬɤɚ [55], ɩɪɢɜɨɞɹɳɢɟ ɤ ɞɨɩɨɥɧɢɬɟɥɶɧɨɦɭ ɭɫɢɥɟɧɢɸ ɜɨɞɨɪɨɞɧɨɣ ɫɜɹɡɢ, ɨɛɪɚɡɨɜɚɧɧɨɣ ɦɨɥɟɤɭɥɨɣ ɜɨɞɵ, SB ɢ Asp85 [56]. ȼ ɬɟɱɟɧɢɟ ɩɟɪɟɯɨɞɚ L o Ɇ, ɞɥɹɳɟɝɨɫɹ ɩɪɢɦɟɪɧɨ 50 ɦɤɫ, ɩɪɨɬɨɧ ɩɟɪɟɯɨɞɢɬ ɫ ɨɫɧɨɜɚɧɢɹ ɒɢɮɮɚ ɧɚ Asp85 [57]. ɋɱɢɬɚɟɬɫɹ, ɱɬɨ ɫɭɳɟɫɬɜɭɟɬ ɩɨ-ɤɪɚɣɧɟɣ ɦɟɪɟ ɞɜɟ ɮɨɪɦɵ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ (ɤɨɬɨɪɵɟ ɧɟɪɚɡɥɢɱɢɦɵ ɫɩɟɤɬɪɨɫɤɨɩɢɱɟɫɤɢ), ɚ ɩɟɪɟɯɨɞ ɨɬ MI ɤ

18 MII, ɹɜɥɹɟɬɫɹ ɨɫɧɨɜɧɵɦ ɧɟɨɛɪɚɬɢɦɵɦ ɲɚɝɨɦ ɜ ɮɨɬɨɰɢɤɥɟ [46]. ȼ ɬɟɱɟɧɢɟ ɷɬɨɝɨ ɩɟɪɟɯɨɞɚ ɚɡɨɬ ɒɢɮɮɨɜɚ ɨɫɧɨɜɚɧɢɹ ɫɬɚɧɨɜɢɬɫɹ ɧɟɞɨɫɬɭɩɧɵɦ ɞɥɹ ɜɧɟɤɥɟɬɨɱɧɨɣ ɱɚɫɬɢ ɩɪɨɬɨɧɧɨɝɨ ɤɚɧɚɥɚ [58], ɱɬɨ ɫɜɹɡɚɧɨ ɫ ɤɨɧɮɨɪɦɚɰɢɨɧɧɵɦɢ ɢɡɦɟɧɟɧɢɹɦɢ ɛɟɥɤɨɜɨɣ ɦɨɥɟɤɭɥɵ. ȼ ɬɟɱɟɧɢɟ ɩɟɪɟɯɨɞɚ Ɇ o N ɚɥɶɞɢɦɢɧɧɚɹ ɝɪɭɩɩɚ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɪɟɩɪɨɬɨɧɢɪɭɟɬɫɹ, ɩɪɢ ɷɬɨɦ ɜ ɤɚɱɟɫɬɜɟ ɞɨɧɨɪɚ ɩɪɨɬɨɧɨɜ ɜɵɫɬɭɩɚɟɬ ɨɫɬɚɬɨɤ Asp96, ɤɨɬɨɪɵɣ ɡɚɬɟɦ ɪɟɩɪɨɬɨɧɢɪɭɟɬɫɹ ɱɟɪɟɡ ɰɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɢɣ ɩɪɨɬɨɧɧɵɣ ɩɨɥɭɤɚɧɚɥ ɩɪɢ ɩɟɪɟɯɨɞɟ N-ɢɧɬɟɪɦɟɞɢɚɬɚ ɜ O. ɉɟɪɟɞɚɱɚ ɩɪɨɬɨɧɚ ɫ Asp85 ɜɨ ɜɧɟɲɧɸɸ ɫɪɟɞɭ ɩɪɨɢɫɯɨɞɢɬ ɜ ɩɪɨɰɟɫɫɟ ɜɨɡɜɪɚɳɟɧɢɹ ɦɨɥɟɤɭɥɵ ȻɊ ɜ ɢɫɯɨɞɧɨɟ ɫɨɫɬɨɹɧɢɟ ȼ.

Asp38

Asp96 Thr46

Val49

Leu93 Thr89

Asp212

Asp85

Arg82 Glu204

Glu194

Glu9

Glu74

Ɋɢɫ. 6. ɋɨɜɪɟɦɟɧɧɚɹ ɦɨɞɟɥɶ ɤɚɧɚɥɚ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ [56-59]. Ʉ ɧɚɫɬɨɹɳɟɦɭ ɦɨɦɟɧɬɭ ɫɱɢɬɚɟɬɫɹ, ɱɬɨ ɩɟɪɟɧɨɫ ɩɪɨɬɨɧɚ ɜɧɭɬɪɢ ɦɨɥɟɤɭɥɵ ȻɊ ɩɪɨɢɫɯɨɞɢɬ ɱɟɪɟɡ ɤɚɧɚɥ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ, ɨɛɪɚɡɭɟɦɨɝɨ ɫɟɬɶɸ ɧɚɯɨɞɹɳɢɯɫɹ ɜɧɭɬɪɢ ȻɊ ɦɨɥɟɤɭɥ ɜɨɞɵ, ɭɞɟɪɠɢɜɚɟɦɵɯ ɜɨɞɨɪɨɞɧɵɦɢ ɫɜɹɡɹɦɢ ɫ ɢɨɧɢɡɭɟɦɵɦɢ ɨɫɬɚɬɤɚɦɢ ɚɦɢɧɨɤɢɫɥɨɬ (Asp ɢ Glu) [56]. Ɋɚɡɜɢɬɢɟ ɝɟɧɟɬɢɱɟɫɤɨɣ ɢɧɠɟɧɟɪɢɢ ɩɨɡɜɨɥɢɥɨ ɪɚɫɲɢɪɢɬɶ ɡɧɚɧɢɹ ɨ ɦɟɯɚɧɢɡɦɟ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ȻɊ. Ɇɟɬɨɞɨɦ ɫɚɣɬ-ɫɩɟɰɢɮɢɱɟɫɤɨɝɨ ɦɭɬɚɝɟɧɟɡɚ ɢɞɟɧɬɢɮɢɰɢɪɨɜɚɧɵ ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɨɤɚɡɵɜɚɸɳɢɟ ɡɧɚɱɢɬɟɥɶɧɨɟ ɜɥɢɹɧɢɟ ɧɚ ɨɩɪɟɞɟɥɟɧɧɵɟ ɫɬɚɞɢɢ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ, ɤɨɬɨɪɵɟ, ɩɨ ɜɫɟɣ ɜɟɪɨɹɬɧɨɫɬɢ, ɩɪɢɧɢɦɚɸɬ ɧɟɩɨɫɪɟɞɫɬɜɟɧɧɨɟ ɭɱɚɫɬɢɟ ɜ ɷɬɨɦ ɩɪɨɰɟɫɫɟ (ɫɦ. ɪɢɫ. 6).

19 ɉɨ ɞɚɧɧɵɦ ɩɪɨɜɟɞɟɧɧɵɯ ɢɫɫɥɟɞɨɜɚɧɢɣ [56, 59], ɜɧɟɤɥɟɬɨɱɧɚɹ ɱɚɫɬɶ ɤɚɧɚɥɚ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɰɟɩɨɱɤɭ ɜɨɞɨɪɨɞɧɵɯ ɫɜɹɡɟɣ, ɫɨɟɞɢɧɹɸɳɢɯ SB ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɫ ɜɧɟɤɥɟɬɨɱɧɨɣ ɫɬɨɪɨɧɨɣ ȻɊ, ɤɨɬɨɪɚɹ ɨɛɪɚɡɨɜɚɧɚ ɧɟɫɤɨɥɶɤɢɦɢ ɦɨɥɟɤɭɥɚɦɢ ɜɨɞɵ, ɭɞɟɪɠɢɜɚɟɦɵɦɢ ɜɨɞɨɪɨɞɧɵɦɢ ɫɜɹɡɹɦɢ ɫ ɨɫɬɚɬɤɚɦɢ Thr89, Asp85, Asp212, Arg82, Glu204 ɢ Glu194. ȼɨ ɜɧɭɬɪɢɤɥɟɬɨɱɧɵɣ ɩɨɥɭɤɚɧɚɥ ɜɯɨɞɹɬ Asp38, Asp96 ɢ ɧɟɫɤɨɥɶɤɨ ɦɨɥɟɤɭɥ ɜɨɞɵ. Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɦɨɠɧɨ ɨɬɦɟɬɢɬɶ ɫɥɟɞɭɸɳɢɟ ɨɫɨɛɟɧɧɨɫɬɢ ȻɊ [2]: x Ɇɨɥɟɤɭɥɹɪɧɵɣ ɜɟɫ 26,7 ɤȾɚ; x ɂɡɨɷɥɟɤɬɪɢɱɟɫɤɚɹ ɬɨɱɤɚ ɩɪɢ ɪɇ 4 – 5; x Ɋɚɫɩɨɥɨɠɟɧɢɟ ɪɟɬɢɧɢɥɢɞɟɧɨɜɨɝɨ ɨɫɬɚɬɤɚ: o ɩɨɥɢɟɧɨɜɚɹ ɰɟɩɶ all-E-ɪɟɬɢɧɚɥɹ ɧɚɯɨɞɢɬɫɹ ɩɨɞ ɭɝɥɨɦ 20q ɤ ɩɥɨɫɤɨɫɬɢ ɦɟɦɛɪɚɧɵ; o ɰɢɤɥɨɝɟɤɫɟɧɨɜɨɟ ɤɨɥɶɰɨ ɨɪɢɟɧɬɢɪɨɜɚɧɨ ɩɟɪɩɟɧɞɢɤɭɥɹɪɧɨ ɩɥɨɫɤɨɫɬɢ ɦɟɦɛɪɚɧɵ; x Ɉɫɨɛɟɧɧɨɫɬɢ ɮɨɬɨɰɢɤɥɚ ȻɊ: o ɨɛɪɚɬɢɦɨɟ ɢɡɦɟɧɟɧɢɟ ɤɨɧɮɢɝɭɪɚɰɢɢ ɪɟɬɢɧɚɥɹ all-E-l13Z-; o ɨɛɪɚɬɢɦɨɟ ɩɪɨɬɨɧɢɪɨɜɚɧɢɟ ɨɫɧɨɜɚɧɢɹ ɒɢɮɮɚ, ɨɛɪɚɡɭɟɦɨɝɨ ɯɪɨɦɨɮɨɪɨɦ; o ɨɛɪɚɬɢɦɨɟ ɢɡɦɟɧɟɧɢɟ ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɨɣ ɫɬɪɭɤɬɭɪɵ ɛɟɥɤɚ, ɦɟɧɹɸɳɟɟ ɞɨɫɬɭɩɧɨɫɬɶ ɨɫɧɨɜɚɧɢɹ ɒɢɮɮɚ ɩɪɨɬɨɧɭ; x Ʉɜɚɧɬɨɜɚɹ ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɩɟɪɜɢɱɧɨɣ ɮɨɬɨɪɟɚɤɰɢɢ ɫɨɫɬɚɜɥɹɟɬ a64%. ȼ ɯɨɞɟ ɢɡɭɱɟɧɢɹ ȻɊ ɢ ɦɟɯɚɧɢɡɦɚ ɩɪɨɬɨɧɧɨɝɨ ɬɪɚɧɫɩɨɪɬɚ ɛɵɥ ɩɪɟɞɥɨɠɟɧ ɪɹɞ ɫɩɨɫɨɛɨɜ ɜɥɢɹɧɢɹ ɤɚɤ ɧɚ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ȻɊ, ɬɚɤ ɢ ɧɚ ɤɢɧɟɬɢɤɭ ɟɝɨ ɮɨɬɨɰɢɤɥɚ.

4. Ɉɫɧɨɜɧɵɟ ɫɩɨɫɨɛɵ ɢɡɦɟɧɟɧɢɹ ɩɚɪɚɦɟɬɪɨɜ ɮɨɬɨɰɢɤɥɚ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. ɉɨɫɤɨɥɶɤɭ ȻɊ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɧɚɢɛɨɥɟɟ ɩɪɨɫɬɨ ɭɫɬɪɨɟɧɧɵɣ ɩɪɟɨɛɪɚɡɨɜɚɬɟɥɶ ɫɜɟɬɨɜɨɣ ɷɧɟɪɝɢɢ, ɦɟɯɚɧɢɡɦ ɟɝɨ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ɚɤɬɢɜɧɨ ɢɫɫɥɟɞɨɜɚɥɫɹ ɪɹɞɨɦ ɧɚɭɱɧɵɯ ɤɨɥɥɟɤɬɢɜɨɜ ɜ Ɋɨɫɫɢɢ ɢ ɡɚ ɪɭɛɟɠɨɦ. ȼ ɷɬɢɯ ɢɫɫɥɟɞɨɜɚɧɢɹɯ ɦɨɠɧɨ ɜɵɞɟɥɢɬɶ ɧɟɫɤɨɥɶɤɨ ɷɬɚɩɨɜ, ɤɨɬɨɪɵɟ ɛɵɥɢ ɫɜɹɡɚɧɵ ɤɚɤ ɫ ɪɚɡɜɢɬɢɟɦ ɩɪɢɛɨɪɧɨɣ ɛɚɡɵ, ɬɚɤ ɢ ɫ ɩɨɹɜɥɟɧɢɟɦ ɧɨɜɵɯ ɦɟɬɨɞɨɥɨɝɢɱɟɫɤɢɯ ɩɨɞɯɨɞɨɜ: 1) ȼ 1970-ɟ – ɧɚɱɚɥɟ 80-ɯ ɝɨɞɨɜ ɩɪɨɜɨɞɢɥɢɫɶ ɛɢɨɯɢɦɢɱɟɫɤɢɟ ɢ ɛɢɨɮɢɡɢɱɟɫɤɢɟ ɢɫɫɥɟɞɨɜɚɧɢɹ ȻɊ. Ȼɵɥɢ ɨɩɪɟɞɟɥɟɧɵ ɮɭɧɤɰɢɢ ȻɊ ɜ ɤɥɟɬɤɟ, ɨɛɧɚɪɭɠɟɧ ɮɨɬɨɰɢɤɥ ɢ ɢɡɭɱɟɧɵ ɟɝɨ ɤɢɧɟɬɢɱɟɫɤɢɟ ɨɫɨɛɟɧɧɨɫɬɢ [60], ɨɩɪɟɞɟɥɟɧɚ ɚɦɢɧɨɤɢɫɥɨɬɧɚɹ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶ [61] ɢ ɩɪɟɞɥɨɠɟɧ ɪɹɞ ɝɢɩɨɬɟɡ ɨ ɦɟɯɚɧɢɡɦɟ ɩɪɨɬɨɧɧɨɝɨ ɬɪɚɧɫɩɨɪɬɚ. 2) ȼ ɤɨɧɰɟ 80-ɯ ɝɨɞɨɜ ɫɬɚɥ ɜɨɡɦɨɠɟɧ ɩɪɨɪɵɜ ɜ ɨɩɪɟɞɟɥɟɧɢɢ ɦɟɯɚɧɢɡɦɚ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ȻɊ ɛɥɚɝɨɞɚɪɹ ɢɫɩɨɥɶɡɨɜɚɧɢɸ ɫɚɣɬ-ɫɩɟɰɢɮɢɱɧɨɝɨ ɦɭɬɚɝɟɧɟɡɚ. ɂɫɫɥɟɞɨɜɚɧɢɟ ɫɜɨɣɫɬɜ ɝɟɧɟɬɢɱɟɫɤɢ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɯ ȻɊ ɩɨɡɜɨɥɢɥɨ

20 ɢɞɟɧɬɢɮɢɰɢɪɨɜɚɬɶ ɨɫɧɨɜɧɵɟ ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɭɱɚɫɬɜɭɸɳɢɟ ɜ ɩɟɪɟɧɨɫɟ ɩɪɨɬɨɧɚ ɢɥɢ ɜɯɨɞɹɳɢɟ ɜ ɛɟɥɤɨɜɨɟ ɦɢɤɪɨɨɤɪɭɠɟɧɢɟ ɯɪɨɦɨɮɨɪɚ [62-74]. Ʉ ɫɨɠɚɥɟɧɢɸ, ɧɟɞɨɫɬɚɬɤɨɦ ɫɚɣɬ-ɫɩɟɰɢɮɢɱɧɨɝɨ ɦɭɬɚɝɟɧɟɡɚ ɹɜɥɹɥɨɫɶ ɨɬɫɭɬɫɬɜɢɟ ɢɧɮɨɪɦɚɰɢɢ ɨ ɜɥɢɹɧɢɢ ɦɭɬɚɰɢɣ ɧɚ ɬɪɟɬɢɱɧɭɸ ɫɬɪɭɤɬɭɪɭ ɛɟɥɤɚ, ɱɬɨ ɦɨɝɥɨ ɩɪɢɜɨɞɢɬɶ ɤ ɧɟɨɞɧɨɡɧɚɱɧɨɫɬɢ ɢɧɬɟɪɩɪɟɬɚɰɢɢ ɩɨɥɭɱɟɧɧɵɯ ɪɟɡɭɥɶɬɚɬɨɜ. 3) Ɋɚɡɪɚɛɨɬɤɚ ɧɨɜɨɝɨ ɫɩɨɫɨɛɚ ɤɪɢɫɬɚɥɥɢɡɚɰɢɢ ȻɊ ɜ ɤɭɛɢɱɟɫɤɭɸ ɮɚɡɭ ɩɨɡɜɨɥɢɥɚ ɨɫɭɳɟɫɬɜɢɬɶ ɤɪɢɫɬɚɥɥɨɝɪɚɮɢɱɟɫɤɢɟ ɢɫɫɥɟɞɨɜɚɧɢɹ ɟɝɨ ɫɬɪɭɤɬɭɪɵ. Ȼɵɥɨ ɩɨɥɭɱɟɧɨ ɛɨɥɶɲɨɟ ɤɨɥɢɱɟɫɬɜɨ ɫɬɪɭɤɬɭɪ ȻɊ ɫ ɜɵɫɨɤɢɦ ɪɚɡɪɟɲɟɧɢɟɦ [75] ɞɨ 1,5 Å, ɱɬɨ ɩɨɡɜɨɥɹɟɬ ɨɩɪɟɞɟɥɢɬɶ ɪɚɫɩɨɥɨɠɟɧɢɟ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɜ ȻɊ ɢ ɟɝɨ ɫɬɚɛɢɥɶɧɵɯ ɢɧɬɟɪɦɟɞɢɚɬɚɯ. ȼ ɫɨɱɟɬɚɧɢɢ ɫ ɞɚɧɧɵɦɢ, ɩɨɥɭɱɟɧɧɵɦɢ ɬɨɱɟɱɧɵɦ ɦɭɬɚɝɟɧɟɡɨɦ, ɷɬɨɬ ɩɨɞɯɨɞ ɩɨɡɜɨɥɹɟɬ ɨɩɪɟɞɟɥɢɬɶ ɮɭɧɤɰɢɨɧɚɥɶɧɨ ɜɚɠɧɵɟ ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ ȻɊ. ȼ ɯɨɞɟ ɩɪɨɜɟɞɟɧɧɵɯ ɢɫɫɥɟɞɨɜɚɧɢɣ ɛɵɥ ɩɪɟɞɥɨɠɟɧ ɪɹɞ ɫɩɨɫɨɛɨɜ ɢɡɦɟɧɟɧɢɹ ɮɨɬɨɯɪɨɦɧɵɯ ɫɜɨɣɫɬɜ ȻɊ ɢ ɧɚɤɨɩɥɟɧ ɛɨɥɶɲɨɣ ɨɛɴɟɦ ɞɚɧɧɵɯ ɩɨ ɟɝɨ ɫɜɨɣɫɬɜɚɦ, ɱɬɨ ɩɨɡɞɧɟɟ ɫɬɚɥɨ ɲɢɪɨɤɨ ɢɫɩɨɥɶɡɨɜɚɬɶɫɹ ɩɪɢ ɪɚɡɪɚɛɨɬɤɟ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɯ ɦɚɬɟɪɢɚɥɨɜ ɧɚ ɨɫɧɨɜɟ ȻɊ. Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɦɨɠɧɨ ɜɵɞɟɥɢɬɶ ɧɟɫɤɨɥɶɤɨ ɨɫɧɨɜɧɵɯ ɩɨɞɯɨɞɨɜ ɤ ɢɡɦɟɧɟɧɢɸ ɮɨɬɨɯɪɨɦɧɵɯ ɫɜɨɣɫɬɜ ȻɊ: 1. ɢɫɩɨɥɶɡɨɜɚɧɢɟ ɯɢɦɢɱɟɫɤɢɯ ɜɟɳɟɫɬɜ ɢɥɢ ɮɢɡɢɱɟɫɤɢɯ ɜɨɡɞɟɣɫɬɜɢɣ ɪɚɡɥɢɱɧɨɝɨ ɪɨɞɚ (ɧɢɡɤɢɯ ɬɟɦɩɟɪɚɬɭɪ, ɷɥɟɤɬɪɢɱɟɫɤɢɯ ɩɨɥɟɣ, ɭɪɨɜɧɹ ɜɥɚɠɧɨɫɬɢ, ɪɇ ɢ ɞɪ.); 2. ɡɚɦɟɳɟɧɢɟ ɩɪɢɪɨɞɧɨɝɨ ɯɪɨɦɨɮɨɪɚ ɧɚ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɣ; 3. ɡɚɦɟɳɟɧɢɟ ɨɞɧɨɝɨ ɢɥɢ ɧɟɫɤɨɥɶɤɢɯ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɜ ɨɩɪɟɞɟɥɟɧɧɵɯ ɩɨɥɨɠɟɧɢɹɯ ɦɨɥɟɤɭɥɵ ȻɊ ɦɟɬɨɞɚɦɢ ɫɚɣɬ-ɫɩɟɰɢɮɢɱɧɨɝɨ ɦɭɬɚɝɟɧɟɡɚ ɢɥɢ ɝɟɧɟɬɢɱɟɫɤɨɣ ɢɧɠɟɧɟɪɢɢ; 4. ɫɨɱɟɬɚɧɢɟ ɧɟɫɤɨɥɶɤɢɯ ɢɡ ɩɟɪɟɱɢɫɥɟɧɧɵɯ ɜɵɲɟ ɦɟɬɨɞɨɜ. ɇɢɠɟ ɩɪɢɜɟɞɟɧ ɨɛɡɨɪ ɬɟɯ ɫɩɨɫɨɛɨɜ ɢɡɦɟɧɟɧɢɹ ɫɜɨɣɫɬɜ ȻɊ, ɤɨɬɨɪɵɟ ɦɨɝɭɬ ɢɦɟɬɶ ɡɧɚɱɟɧɢɟ ɩɪɢ ɪɚɡɪɚɛɨɬɤɟ ɜɚɪɢɚɧɬɨɜ ȻɊ, ɨɛɥɚɞɚɸɳɢɯ ɨɩɪɟɞɟɥɟɧɧɵɦɢ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɦɢ ɯɚɪɚɤɬɟɪɢɫɬɢɤɚɦɢ.

4.1. Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɜɨɡɞɟɣɫɬɜɢɹ. ȼ ɛɨɥɶɲɢɧɫɬɜɟ ɫɥɭɱɚɟɜ ɜɥɢɹɧɢɟ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɯ ɮɚɤɬɨɪɨɜ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ ɛɵɥɨ ɨɛɧɚɪɭɠɟɧɨ ɫɥɭɱɚɣɧɨ, ɩɨɫɤɨɥɶɤɭ ɢɡɧɚɱɚɥɶɧɨ ɧɟɜɨɡɦɨɠɧɨ ɩɪɟɞɫɤɚɡɚɬɶ, ɤɚɤ ɨɧɢ ɜɨɡɞɟɣɫɬɜɭɸɬ ɧɚ ɩɪɨɰɟɫɫɵ ɜ ɦɨɥɟɤɭɥɟ ȻɊ. ɂɫɫɥɟɞɨɜɚɬɟɥɢ ɩɟɪɟɛɢɪɚɥɢ ɜɫɟɜɨɡɦɨɠɧɵɟ ɩɨɞɯɨɞɵ, ɩɨɡɜɨɥɹɸɳɢɟ ɢɡɦɟɧɹɬɶ ɫɜɨɣɫɬɜɚ ȻɊ: ɬɟɦɩɟɪɚɬɭɪɚ, ɪɇ ɫɪɟɞɵ, ɜɥɚɠɧɨɫɬɶ, ɷɥɟɤɬɪɢɱɟɫɤɢɟ ɩɨɥɹ, ɜɤɥɸɱɟɧɢɟ ɜ ɩɨɥɢɦɟɪɧɭɸ ɦɚɬɪɢɰɭ ɢ ɪɚɡɥɢɱɧɵɟ ɯɢɦɢɱɟɫɤɢɟ ɜɨɡɞɟɣɫɬɜɢɹ.

21 ɍɦɟɧɶɲɟɧɢɟ ɪɇ ɫ 7 ɞɨ 2 ɜɵɡɵɜɚɟɬ ɩɟɪɟɯɨɞ ȻɊ ɜ ɤɢɫɥɭɸ ɫɢɧɸɸ ɮɨɪɦɭ (Omax ɫ 552 ɢɡɦɟɧɹɟɬɫɹ ɞɨ 605 ɧɦ ɜ ɬɟɦɧɨ-ɚɞɚɩɬɢɪɨɜɚɧɧɨɦ ȻɊ), ɤɨɬɨɪɨɟ ɫɜɹɡɵɜɚɸɬ ɫ ɩɪɨɬɨɧɢɪɨɜɚɧɢɟɦ Asp85 ɢ ɢɫɩɨɥɶɡɭɸɬ ɞɥɹ ɨɰɟɧɤɢ ɟɟ ɪɄɚ [76]. Ⱦɚɥɶɧɟɣɲɟɟ ɩɨɧɢɠɟɧɢɟ ɪɇ (ɞɨ ɤɨɧɰɟɧɬɪɚɰɢɢ 2Ɇ HCl) ɩɪɢɜɨɞɢɬ ɤ ɨɛɪɚɡɨɜɚɧɢɸ ɤɢɫɥɨɣ ɩɭɪɩɭɪɧɨɣ ɮɨɪɦɵ ȻɊ ɫ Omax 565 ɧɦ. ɉɨɜɵɲɟɧɢɟ ɪɇ ɩɪɢɜɨɞɢɬ ɤ ɩɟɪɟɯɨɞɭ ȻɊ ɜ ɧɟɚɤɬɢɜɧɭɸ ɳɟɥɨɱɧɭɸ ɮɨɪɦɭ ɫ Omax 480 ɧɦ, ɱɬɨ ɫɜɹɡɵɜɚɸɬ ɫ ɞɟɩɪɨɬɨɧɢɪɨɜɚɧɢɟɦ ɩɪɨɬɨɧ-ɜɵɞɟɥɹɸɳɟɣ ɝɪɭɩɩɵ ɛɟɥɤɚ (SB) ɢ ɢɫɩɨɥɶɡɭɸɬ ɞɥɹ ɨɰɟɧɤɢ ɟɟ ɪɄɚ [76], ɨɞɧɚɤɨ ɭɜɟɥɢɱɟɧɢɟ ɪɇ ɛɨɥɶɲɟ 12 ɜɵɡɵɜɚɟɬ ɧɟɨɛɪɚɬɢɦɭɸ ɞɟɧɚɬɭɪɚɰɢɸ ɛɟɥɤɚ. ɂɡ ɪɚɡɧɨɨɛɪɚɡɧɵɯ ɯɢɦɢɱɟɫɤɢɯ ɦɟɬɨɞɨɜ ɢɡɦɟɧɟɧɢɹ ɫɜɨɣɫɬɜ ȻɊ ɦɨɠɧɨ ɜɵɞɟɥɢɬɶ ɫɥɟɞɭɸɳɢɟ: ɧɟɤɨɬɨɪɵɟ ɩɟɪɜɢɱɧɵɟ ɚɦɢɧɵ ɩɪɢɜɨɞɹɬ ɤ ɭɜɟɥɢɱɟɧɢɸ ɫɪɨɤɚ ɠɢɡɧɢ ɢɧɬɟɪɦɟɞɢɚɬɚ Ɇ, ɚ ɚɡɢɞ ɧɚɬɪɢɹ ɩɨɤɚɡɵɜɚɟɬ ɩɪɨɬɢɜɨɩɨɥɨɠɧɵɣ ɷɮɮɟɤɬ [77]. Ⱦɟɣɫɬɜɢɟ ɧɟɢɨɧɧɵɯ «ɦɹɝɤɢɯ» ɞɟɬɟɪɝɟɧɬɨɜ (ɧɚɩɪɢɦɟɪ, Triton X-100) ɜɵɡɵɜɚɟɬ ɞɢɫɫɨɰɢɚɰɢɸ ɉɆ ɧɚ ɦɨɧɨɦɟɪɵ ɢ ɭɦɟɧɶɲɚɟɬ ɫɬɚɛɢɥɶɧɨɫɬɶ ȻɊ ɤ ɜɧɟɲɧɢɦ ɜɨɡɞɟɣɫɬɜɢɹɦ, ɨɞɧɚɤɨ ɨɧ ɜ ɷɬɢɯ ɭɫɥɨɜɢɹɯ ɫɨɯɪɚɧɹɟɬ ɫɩɨɫɨɛɧɨɫɬɶ ɤ ɩɟɪɟɧɨɫɭ ɩɪɨɬɨɧɚ, ɯɨɬɹ ɤɢɧɟɬɢɤɚ ɮɨɬɨɰɢɤɥɚ ɡɚɦɟɬɧɨ ɢɡɦɟɧɹɟɬɫɹ [30]. Ȼɨɥɟɟ «ɠɟɫɬɤɢɟ» ɞɟɬɟɪɝɟɧɬɵ, ɬɚɤɢɟ ɤɚɤ SDS ɢɥɢ CTAB, ɧɟ ɬɨɥɶɤɨ ɪɚɡɪɭɲɚɸɬ ɫɬɪɭɤɬɭɪɭ ɉɆ, ɧɨ ɢ ɜɵɡɵɜɚɸɬ ɞɟɧɚɬɭɪɚɰɢɸ ȻɊ. Ȼɨɥɶɲɢɧɫɬɜɨ ɩɨɥɹɪɧɵɯ ɪɚɫɬɜɨɪɢɬɟɥɟɣ ɞɟɣɫɬɜɭɸɬ ɧɚ ɉɆ ɬɚɤ ɠɟ, ɤɚɤ «ɠɟɫɬɤɢɟ» ɞɟɬɟɪɝɟɧɬɵ, ɧɟɨɛɪɚɬɢɦɨ ɪɚɡɪɭɲɚɹ ɛɟɥɨɤ ɫ ɞɢɫɫɨɰɢɚɰɢɟɣ SB, ɯɨɬɹ ɧɟɤɨɬɨɪɵɟ ɪɚɫɬɜɨɪɢɬɟɥɢ (ɬɚɤɢɟ, ɤɚɤ ɞɢɷɬɢɥɨɜɵɣ ɷɮɢɪ, ȾɆɋɈ ɢ ȾɆɎȺ) ɜ ɜɵɫɨɤɢɯ ɤɨɧɰɟɧɬɪɚɰɢɹɯ ɜɵɡɵɜɚɸɬ ɥɢɲɶ ɨɛɪɚɬɢɦɭɸ ɞɟɧɚɬɭɪɚɰɢɸ ɛɟɥɤɚ [42].

4.2. Ɂɚɦɟɧɚ ɯɪɨɦɨɮɨɪɚ. Ɉɛɧɚɪɭɠɟɧɢɟ ɜɨɡɦɨɠɧɨɫɬɢ ɩɨɥɭɱɟɧɢɹ ɛɚɤɬɟɪɢɨɨɩɫɢɧɚ ɫɜɟɬɨɡɚɜɢɫɢɦɵɦ ɝɢɞɪɨɤɫɢɥɚɦɢɧɨɥɢɡɨɦ ɩɨɡɜɨɥɢɥɨ ɩɨɥɭɱɚɬɶ ȺȻɊ, ɜ ɤɨɬɨɪɵɯ ɩɪɢɪɨɞɧɵɣ ɯɪɨɦɨɮɨɪ ɡɚɦɟɧɟɧ ɧɚ ɫɢɧɬɟɬɢɱɟɫɤɢɣ. Ɋɟɡɭɥɶɬɚɬɵ ɢɫɫɥɟɞɨɜɚɧɢɣ ɜ ɷɬɨɣ ɨɛɥɚɫɬɢ ɞɨɫɬɚɬɨɱɧɨ ɩɨɞɪɨɛɧɨ ɨɩɢɫɚɧɵ ɜ ɨɛɡɨɪɚɯ [78-80]. Ⱦɚɧɧɵɣ ɩɨɞɯɨɞ ɲɢɪɨɤɨ ɢɫɩɨɥɶɡɨɜɚɥɫɹ ɞɥɹ ɢɡɭɱɟɧɢɹ ɫɬɪɨɟɧɢɹ ɢ ɦɟɯɚɧɢɡɦɚ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ȻɊ, ɚ ɢɦɟɧɧɨ ɞɥɹ ɨɩɪɟɞɟɥɟɧɢɹ ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɵɯ ɨɝɪɚɧɢɱɟɧɢɣ ɢ ɮɨɪɦɵ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ȻɊ, ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ, ɪɚɫɩɨɥɨɠɟɧɧɵɯ ɨɤɨɥɨ ɪɟɬɢɧɚɥɹ, ɫɬɟɩɟɧɢ ɡɧɚɱɢɦɨɫɬɢ ɨɩɪɟɞɟɥɟɧɧɵɯ ɷɥɟɦɟɧɬɨɜ ɫɬɪɭɤɬɭɪɵ ɪɟɬɢɧɚɥɹ ɞɥɹ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ɛɟɥɤɚ. ȼ ɪɟɡɭɥɶɬɚɬɟ ɩɪɨɜɟɞɟɧɧɵɯ ɢɫɫɥɟɞɨɜɚɧɢɣ ɛɵɥɨ ɩɨɥɭɱɟɧɨ ɧɟɫɤɨɥɶɤɨ ɫɨɬɟɧ ɚɧɚɥɨɝɨɜ ȻɊ, ɤɨɬɨɪɵɟ ɦɨɠɧɨ ɩɨɞɪɚɡɞɟɥɢɬɶ ɧɚ ɧɟɫɤɨɥɶɤɨ ɤɚɬɟɝɨɪɢɣ ɩɨ ɬɢɩɭ ɦɨɞɢɮɢɤɚɰɢɢ ɫɬɪɭɤɬɭɪɵ ɪɟɬɢɧɚɥɹ (ɫɦ. ɪɢɫ. 7): x ȼɜɟɞɟɧɢɟ ɪɚɡɥɢɱɧɵɯ ɡɚɦɟɫɬɢɬɟɥɟɣ ɜɨ 2, 3 ɢ 4 ɩɨɥɨɠɟɧɢɹ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ;

22 x Ɂɚɦɟɧɚ ɤɨɥɶɰɚ ɧɚ ɚɪɨɦɚɬɢɱɟɫɤɢɟ ɢ ɝɟɬɟɪɨɰɢɤɥɢɱɟɫɤɢɟ ɮɪɚɝɦɟɧɬɵ ɢɥɢ ɟɝɨ ɭɞɚɥɟɧɢɟ; x ɂɡɦɟɧɟɧɢɟ ɩɪɢɪɨɞɵ ɬɟɪɦɢɧɚɥɶɧɨɣ ɩɨɥɹɪɧɨɣ ɝɪɭɩɩɵ (ɩɨɥɭɱɟɧɢɟ ɫɥɨɠɧɵɯ ɷɮɢɪɨɜ ɢɥɢ ɧɢɬɪɢɥɨɜ), ɱɬɨɛɵ ɩɨɤɚɡɚɬɶ, ɱɬɨ ɜ ɷɬɨɦ ɫɥɭɱɚɟ ɚɧɚɥɨɝ ɪɟɬɢɧɚɥɹ ɨɛɪɚɡɭɟɬ ɬɨɥɶɤɨ ɧɟɤɨɜɚɥɟɧɬɧɵɣ ɤɨɦɩɥɟɤɫ ɫ ɛɚɤɬɟɪɢɨɨɩɫɢɧɨɦ; x ɂɡɦɟɧɟɧɢɟ ɞɥɢɧɵ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ, ɡɚɦɟɧɚ ɞɜɨɣɧɵɯ ɫɜɹɡɟɣ ɧɚ ɨɞɢɧɚɪɧɵɟ ɢɥɢ ɬɪɨɣɧɵɟ, ɩɨɥɭɱɟɧɢɟ ɧɟɢɡɨɦɟɪɢɡɭɸɳɢɯɫɹ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɩɭɬɟɦ ɜɜɟɞɟɧɢɹ ɰɢɤɥɨɜ ɪɚɡɥɢɱɧɨɝɨ ɪɚɡɦɟɪɚ ɢ ɩɪɢɪɨɞɵ; x Ɂɚɦɟɧɚ ɦɟɬɢɥɶɧɵɯ ɝɪɭɩɩ ɩɪɢ ɋ9 ɢ ɋ13 ɡɚɦɟɫɬɢɬɟɥɹɦɢ ɪɚɡɥɢɱɧɨɣ ɩɪɢɪɨɞɵ; x Ɂɚɤɪɟɩɥɟɧɢɟ s-ɰɢɫ-ɤɨɧɮɢɝɭɪɚɰɢɢ ɤɨɥɶɰɨ-ɩɨɥɢɟɧɨɜɚɹ ɰɟɩɶ ɜɜɟɞɟɧɢɟɦ ɰɢɤɥɢɱɟɫɤɢɯ ɫɬɪɭɤɬɭɪ ɜ ɨɛɥɚɫɬɶ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ. ȼɜɟɞɟɧɢɟ ɡɚɦɟɫɬɢɬɟɥɟɣ, ɨɛɥɚɞɚɸɳɢɯ ɪɚɡɥɢɱɧɵɦɢ ɪɚɡɦɟɪɚɦɢ ɢ ɷɥɟɤɬɪɨɨɬɪɢɰɚɬɟɥɶɧɨɫɬɶɸ

Ɂɚɦɟɧɚ ɧɚ ɚɪɨɦɚɬɢɱɟɫɤɢɟ ɢ ɝɟɬɟɪɨɰɢɤɥɢɱɟɫɤɢɟ ɮɪɚɝɦɟɧɬɵ

ɂɡɦɟɧɟɧɢɟ ɩɪɢɪɨɞɵ ɬɟɪɦɢɧɚɥɶɧɨɣ ɩɨɥɹɪɧɨɣ ɝɪɭɩɩɵ 16

1

2

7

6

3 4

ȼɜɟɞɟɧɢɟ ɡɚɦɟɫɬɢɬɟɥɟɣ ɪɚɡɥɢɱɧɨɣ ɩɪɢɪɨɞɵ

20

19 17

5

11

9 8

10

15

13 12

14

O

18

Ɂɚɤɪɟɩɥɟɧɢɟ ɤɨɧɮɢɝɭɪɚɰɢɢ ɜ ɪɚɣɨɧɟ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ

ɂɡɦɟɧɟɧɢɟ ɞɥɢɧɵ ɢ ɤɨɧɮɢɝɭɪɚɰɢɢ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ, ɩɨɥɭɱɟɧɢɟ ɧɟɢɡɨɦɟɪɢɡɭɸɳɢɯɫɹ ɚɧɚɥɨɝɨɜ

Ɋɢɫ. 7. Ɉɫɧɨɜɧɵɟ ɧɚɩɪɚɜɥɟɧɢɹ ɦɨɞɢɮɢɤɚɰɢɢ ɫɬɪɭɤɬɭɪɵ ɪɟɬɢɧɚɥɹ. Ȼɵɥɨ ɢɫɫɥɟɞɨɜɚɧɨ ɜɥɢɹɧɢɟ ɩɟɪɟɱɢɫɥɟɧɧɵɯ ɜɵɲɟ ɦɨɞɢɮɢɤɚɰɢɣ ɧɚ ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ȻɊ, ɫɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ ɢ ɢɡɨɦɟɪɧɵɣ ɫɨɫɬɚɜ ɫɜɟɬɨ- ɢ ɬɟɦɧɨɚɞɚɩɬɢɪɨɜɚɧɧɨɣ ɮɨɪɦɵ, ɧɚɥɢɱɢɟ ɢ ɫɤɨɪɨɫɬɶ ɮɨɬɨɰɢɤɥɚ, ɩɪɨɬɨɧɧɵɣ ɬɪɚɧɫɩɨɪɬ ɢ ɭɫɬɨɣɱɢɜɨɫɬɶ ɩɢɝɦɟɧɬɚ ɤ ɞɟɣɫɬɜɢɸ ɝɢɞɪɨɤɫɢɥɚɦɢɧɚ (ɜ ɬɟɦɧɨɬɟ) ɢ ɪɟɬɢɧɚɥɹ. Ʉ ɫɨɠɚɥɟɧɢɸ, ɥɢɲɶ ɨɤɨɥɨ ɬɪɟɬɢ ɩɨɥɭɱɟɧɧɵɯ ɚɧɚɥɨɝɨɜ ȻɊ ɛɵɥɢ ɢɫɫɥɟɞɨɜɚɧɵ ɩɨ ɜɫɟɦ ɩɟɪɟɱɢɫɥɟɧɧɵɦ ɩɚɪɚɦɟɬɪɚɦ. ɑɚɫɬɨ ɚɜɬɨɪɵ ɨɝɪɚɧɢɱɢɜɚɥɢɫɶ ɥɢɲɶ ɨɩɪɟɞɟɥɟɧɢɟɦ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ɢ ɧɚɥɢɱɢɹ ɬɪɚɧɫɩɨɪɬɚ ɩɪɨɬɨɧɨɜ. Ɉɞɧɚɤɨ ɩɪɨɜɟɞɟɧɧɵɟ ɢɫɫɥɟɞɨɜɚɧɢɹ ɩɨɡɜɨɥɹɸɬ ɫɞɟɥɚɬɶ ɪɹɞ ɡɚɤɥɸɱɟɧɢɣ ɨ ɜɥɢɹɧɢɢ ɫɬɪɭɤɬɭɪɧɵɯ ɦɨɞɢɮɢɤɚɰɢɣ ɪɟɬɢɧɚɥɹ ɧɚ ɫɜɨɣɫɬɜɚ ɛɟɥɤɚ ɜ ɰɟɥɨɦ. ɚ) Ⱦɥɢɧɚ, ɤɨɧɮɢɝɭɪɚɰɢɹ ɢ ɫɬɟɩɟɧɶ ɧɟɧɚɫɵɳɟɧɧɨɫɬɢ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɯɪɨɦɨɮɨɪɚ. ȻɈ ɫɩɨɫɨɛɟɧ ɨɛɪɚɡɨɜɵɜɚɬɶ ɩɢɝɦɟɧɬɵ ɫ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ, ɢɦɟɸɳɢɦɢ

23 ɰɟɩɶ ɫɨɩɪɹɠɟɧɢɹ ɧɚ ɞɜɚ ɢɥɢ ɩɹɬɶ ɭɝɥɟɪɨɞɧɵɯ ɚɬɨɦɚ ɛɨɥɶɲɟ, ɱɟɦ ɩɪɢɪɨɞɧɵɣ ɪɟɬɢɧɚɥɶ [81-84]. ȿɫɥɢ ɭ ɋ22-ȻɊ ɫɨɯɪɚɧɹɟɬɫɹ ɮɨɬɨɰɢɤɥ ɢ ɫɥɚɛɵɣ ɩɪɨɬɨɧɧɵɣ ɬɪɚɧɫɩɨɪɬ, ɬɨ ɭ ɋ25ȻɊ ɮɨɬɨɰɢɤɥ ɩɨɱɬɢ ɧɟ ɜɵɪɚɠɟɧ. ȼ ɬɨ ɠɟ ɜɪɟɦɹ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɫ ɛɨɥɟɟ ɤɨɪɨɬɤɢɦɢ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ (ɋ15- ɢ ɋ17-ɚɥɶɞɟɝɢɞɵ) [83, 84] ȻɈ ɞɚɟɬ ɬɨɥɶɤɨ ɧɟɤɨɜɚɥɟɧɬɧɵɟ ɤɨɦɩɥɟɤɫɵ. ȿɳɟ ɛɨɥɟɟ ɫɢɥɶɧɨɟ ɜɥɢɹɧɢɟ ɨɤɚɡɵɜɚɟɬ ɢɡɦɟɧɟɧɢɟ ɩɨɪɹɞɤɚ ɱɟɪɟɞɨɜɚɧɢɹ ɤɪɚɬɧɵɯ ɫɜɹɡɟɣ ɭ ɚɧɚɥɨɝɚ ɋ21-ȻɊ [81] ɫ ɪɟɬɪɨɫɬɪɭɤɬɭɪɨɣ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɯɪɨɦɨɮɨɪɚ, ɭ ɤɨɬɨɪɨɝɨ ɩɨɥɧɨɫɬɶɸ ɨɬɫɭɬɫɬɜɭɟɬ ɮɨɬɨɰɢɤɥ. Ɇɟɧɟɟ ɡɚɦɟɬɧɨɟ ɜɥɢɹɧɢɟ ɨɤɚɡɵɜɚɟɬ ɜɚɪɶɢɪɨɜɚɧɢɟ ɫɬɟɩɟɧɢ ɧɟɧɚɫɵɳɟɧɧɨɫɬɢ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɯɪɨɦɨɮɨɪɚ. ɉɨɤɚɡɚɧɨ, ɱɬɨ ɞɥɹ ɧɚɥɢɱɢɹ ɮɨɬɨɰɢɤɥɚ ɨɧɚ ɞɨɥɠɧɚ ɫɨɞɟɪɠɚɬɶ ɧɟ ɦɟɧɟɟ ɬɪɟɯ ɫɨɩɪɹɠɟɧɧɵɯ ɞɜɨɣɧɵɯ ɫɜɹɡɟɣ. ɏɨɬɹ ȻɈ ɨɛɪɚɡɭɟɬ ɯɪɨɦɨɩɪɨɬɟɢɧɵ ɢ ɫ ɝɢɞɪɢɪɨɜɚɧɧɵɦɢ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ [85, 86], ɧɨ ɞɚɠɟ ɜ ɫɥɭɱɚɟ ɜɨɫɫɬɚɧɨɜɥɟɧɢɹ ɤɪɚɬɧɨɣ ɫɜɹɡɢ ɜ ɤɨɥɶɰɟ (5,6-ɞɢɝɢɞɪɨ-ȻɊ), ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɩɪɨɬɨɧɧɨɝɨ ɬɪɚɧɫɩɨɪɬɚ ɡɚɦɟɬɧɨ ɫɧɢɠɟɧɚ [87]. ɋ ɞɪɭɝɨɣ ɫɬɨɪɨɧɵ, ɜɜɟɞɟɧɢɟ ɬɪɨɣɧɨɣ ɫɜɹɡɢ ɤɚɤ ɜ ɋ7ɋ8- [88] ɢɥɢ ɋ11-ɋ12- [89-91] ɩɨɥɨɠɟɧɢɹ ɪɟɬɢɧɚɥɹ ɧɟ ɩɪɟɩɹɬɫɬɜɭɟɬ ɩɪɨɰɟɫɫɭ ɪɟɤɨɧɫɬɪɭɤɰɢɢ ɩɢɝɦɟɧɬɨɜ, ɮɨɬɨɰɢɤɥɭ ɢ ɬɪɚɧɫɩɨɪɬɭ ɩɪɨɬɨɧɨɜ. Ȼɨɥɶɲɨɟ ɜɥɢɹɧɢɟ ɧɚ ɪɟɡɭɥɶɬɚɬ ɜɡɚɢɦɨɞɟɣɫɬɜɢɹ ɩɨɥɢɟɧɚɥɹ ɫ ȻɈ ɨɤɚɡɵɜɚɟɬ ɤɨɧɮɢɝɭɪɚɰɢɹ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ. ɋɱɢɬɚɟɬɫɹ, ɱɬɨ ɚɥɶɞɢɦɢɧɧɭɸ ɫɜɹɡɶ ɫ H-ɚɦɢɧɨɝɪɭɩɩɨɣ ɥɢɡɢɧɚ-216 ɦɨɠɟɬ ɨɛɪɚɡɨɜɵɜɚɬɶ ɬɨɥɶɤɨ all-E- ɢ 13Z-ɢɡɨɦɟɪɵ ɪɟɬɢɧɚɥɹ ɢ ɟɝɨ ɚɧɚɥɨɝɨɜ. ɇɚ ɫɟɝɨɞɧɹɲɧɢɣ ɞɟɧɶ ɛɵɥɢ ɨɛɧɚɪɭɠɟɧɵ ɬɨɥɶɤɨ ɟɞɢɧɢɱɧɵɟ ɢɫɤɥɸɱɟɧɢɹ ɢɡ ɷɬɨɝɨ ɩɪɚɜɢɥɚ: ɷɬɨ 13-ɞɟɡɦɟɬɢɥɪɟɬɢɧɚɥɶ, ɭ ɤɨɬɨɪɨɝɨ ɜ ȻɈ ɬɚɤɠɟ ɜɫɬɪɚɢɜɚɸɬɫɹ 11Z- ɢ 7Zɢɡɨɦɟɪɵ [92-94], ɢ ɧɚɮɬɢɥɶɧɵɣ ɚɧɚɥɨɝ ɪɟɬɢɧɚɥɹ, ɭ ɤɨɬɨɪɨɝɨ 9Z-ɢɡɨɦɟɪ ɫɩɨɫɨɛɟɧ ɨɛɪɚɡɨɜɵɜɚɬɶ ɩɢɝɦɟɧɬ, ɚ ɭ 11,12-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɹ ɩɢɝɦɟɧɬ ɨɛɪɚɡɭɸɬ ɬɚɤ ɠɟ 9Z- ɢ 9Z,13Z-ɢɡɨɦɟɪɵ [90, 91]. ɛ) Ɇɨɞɢɮɢɤɚɰɢɢ ɪɚɣɨɧɚ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɯɪɨɦɨɮɨɪɚ. ȻɈ ɨɛɪɚɡɭɟɬ ɯɪɨɦɨɩɪɨɬɟɢɧɵ ɫ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ, ɭ ɤɨɬɨɪɵɯ ɤɨɥɶɰɨ ɦɨɠɟɬ ɛɵɬɶ ɦɨɞɢɮɢɰɢɪɨɜɚɧɨ ɩɨ ɋ2-, ɋ3-, ɋ4-, ɋ5-ɩɨɥɨɠɟɧɢɹɦ ɢ ɋ5-ɋ6-ɞɜɨɣɧɨɣ ɫɜɹɡɢ, ɭɞɚɥɟɧɨ ɜɨɨɛɳɟ ɢɥɢ ɡɚɦɟɧɟɧɨ ɧɚ ɡɚɦɟɳɟɧɧɵɟ ɚɪɨɦɚɬɢɱɟɫɤɢɟ ɢ ɝɟɬɟɪɨɰɢɤɥɢɱɟɫɤɢɟ ɮɪɚɝɦɟɧɬɵ, ɨɬɥɢɱɚɸɳɢɟɫɹ ɤɚɤ ɫɜɨɢɦ ɪɚɡɦɟɪɨɦ, ɬɚɤ ɢ ɷɥɟɤɬɪɨɧɧɨɣ ɩɪɢɪɨɞɨɣ ɡɚɦɟɫɬɢɬɟɥɟɣ. Ⱦɥɹ ɷɬɨɝɨ ɬɢɩɚ ɦɨɞɢɮɢɤɚɰɢɣ ɦɨɠɧɨ ɫɞɟɥɚɬɶ ɪɹɞ ɡɚɦɟɱɚɧɢɣ: 1) ɩɪɚɤɬɢɱɟɫɤɢ ɞɥɹ ɜɫɟɯ ɩɢɝɦɟɧɬɨɜ, ɫɨɞɟɪɠɚɳɢɯ ɞɨɫɬɚɬɨɱɧɨ ɨɛɴɟɦɧɵɟ ɡɚɦɟɫɬɢɬɟɥɢ ɩɪɢ ɋ4, ɯɚɪɚɤɬɟɪɟɧ ɫɢɥɶɧɵɣ ɝɢɩɫɨɯɪɨɦɧɵɣ ɫɞɜɢɝ Omax [95, 96]; 2) ɫɭɳɟɫɬɜɭɟɬ ɨɩɪɟɞɟɥɟɧɧɚɹ ɚɫɢɦɦɟɬɪɢɹ ɛɟɥɤɨɜɨɣ ɩɨɥɨɫɬɢ ɜ ɷɬɨɦ ɪɚɣɨɧɟ, ɩɪɨɹɜɥɹɸɳɚɹɫɹ ɜ ɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɢ ɤ ɫɬɟɪɟɨɯɢɦɢɢ ɡɚɦɟɫɬɢɬɟɥɟɣ ɭ ɋ4-, ɋ3-ɢɥɢ ɋ5-ɋ6-ɚɬɨɦɨɜ ɤɨɥɶɰɚ ɢ ɩɪɢɜɨɞɹɳɚɹ ɤ ɩɨɹɜɥɟɧɢɸ ɧɟɫɤɨɥɶɤɢɯ ɫɩɟɤɬɪɚɥɶɧɵɯ ɮɨɪɦ ɫ ɪɚɡɥɢɱɧɵɦɢ Omax ɢɥɢ "ɪɚɡɦɵɬɨɫɬɢ" ɨɫɧɨɜɧɨɣ ɩɨɥɨɫɵ ɩɨɝɥɨɳɟɧɢɹ ȻɊ; 3) ɷɥɟɤɬɪɨɧɧɵɟ ɷɮɮɟɤɬɵ ɡɚɦɟɫɬɢɬɟɥɟɣ ɩɪɢ ɋ5-ɚɬɨɦɟ ɦɨɝɭɬ ɢɦɟɬɶ

24 ɪɟɲɚɸɳɟɟ ɜɥɢɹɧɢɟ ɧɚ ɫɜɟɬɨɡɚɜɢɫɢɦɵɣ ɬɪɚɧɫɩɨɪɬ ɩɪɨɬɨɧɨɜ (ɡɚɦɟɧɚ ɦɟɬɢɥɶɧɨɣ ɝɪɭɩɩɵ ɩɪɢ ɋ5 ɧɚ ɬɪɢɮɬɨɪɦɟɬɢɥɶɧɭɸ ɜɵɡɵɜɚɟɬ ɩɨɥɧɭɸ ɛɥɨɤɢɪɨɜɤɭ ɷɬɨɝɨ ɩɪɨɰɟɫɫɚ [97]). ɜ) Ɋɚɣɨɧ ɋ13-ɋ15-ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɯɪɨɦɨɮɨɪɚ. ɗɬɚ ɨɛɥɚɫɬɶ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ȻɊ ɩɨɞɜɟɪɝɚɥɚɫɶ ɧɚɢɛɨɥɟɟ ɢɧɬɟɧɫɢɜɧɨɦɭ ɢɫɫɥɟɞɨɜɚɧɢɸ, ɬɚɤ ɤɚɤ ɧɟɨɛɯɨɞɢɦɨɫɬɶ ɢɡɨɦɟɪɢɡɚɰɢɢ ɞɜɨɣɧɨɣ ɫɜɹɡɢ ɋ13-ɋ14 ɞɥɹ ɮɨɬɨɯɢɦɢɱɟɫɤɨɝɨ ɰɢɤɥɚ ȻɊ ɬɜɟɪɞɨ ɭɫɬɚɧɨɜɥɟɧ. ȼɩɟɪɜɵɟ ɷɬɨ ɛɵɥɨ ɫɬɪɨɝɨ ɞɨɤɚɡɚɧɨ Nakanishi et. al. [98], ɩɪɢɱɟɦ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɧɟɢɡɨɦɟɪɢɡɭɸɳɢɟɫɹ ɚɧɚɥɨɝɢ ȻɊ ɤɚɤ ɫ all-E- , ɬɚɤ ɢ ɫ 13Zɤɨɧɮɢɝɭɪɚɰɢɟɣ ɧɟ ɩɪɨɹɜɥɹɸɬ ɮɨɬɨɰɢɤɥɚ [99-103]. ɂɡɭɱɟɧɢɟ ɛɨɥɶɲɨɣ ɫɟɪɢɢ ȺȻɊ, ɫɨɞɟɪɠɚɳɢɯ ɪɚɡɥɢɱɧɵɟ ɡɚɦɟɫɬɢɬɟɥɢ ɩɪɢ ɋ13ɚɬɨɦɟ ɪɟɬɢɧɚɥɹ [96, 103, 104] ɩɨɤɚɡɚɥɨ, ɱɬɨ ɦɟɬɢɥɶɧɚɹ ɝɪɭɩɩɚ ɩɪɢ ɜɫɬɪɚɢɜɚɧɢɢ ɜ ɯɪɨɦɨɮɨɪɧɭɸ ɩɨɥɨɫɬɶ ɫɩɨɫɨɛɫɬɜɭɟɬ ɩɪɚɜɢɥɶɧɨɣ ɨɪɢɟɧɬɚɰɢɢ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ. ɗɬɨ ɩɪɟɞɩɨɥɨɠɟɧɢɟ ɨɫɧɨɜɚɧɨ ɧɚ ɬɨɦ, ɱɬɨ all-E-ɢɡɨɦɟɪ 13-ɞɟɡɦɟɬɢɥɪɟɬɢɧɚɥɹ ɫ ȻɈ ɞɚɟɬ ɬɨɥɶɤɨ ɧɟɤɨɜɚɥɟɧɬɧɵɣ ɤɨɦɩɥɟɤɫ [105], ɭɫɬɨɣɱɢɜɵɣ ɜ ɬɟɦɧɨɬɟ ɩɪɢ 4°ɋ ɜ ɬɟɱɟɧɢɢ ɧɟɫɤɨɥɶɤɢɯ ɞɧɟɣ, ɧɨ ɩɪɢ ɩɨɜɵɲɟɧɢɢ ɬɟɦɩɟɪɚɬɭɪɵ ɢɥɢ ɩɪɢ ɨɫɜɟɳɟɧɢɢ ɩɪɨɢɫɯɨɞɢɬ all-E- o13Z- ɢɡɨɦɟɪɢɡɚɰɢɹ, ɫɨɩɪɨɜɨɠɞɚɟɦɚɹ ɨɛɪɚɡɨɜɚɧɢɟɦ ɩɢɝɦɟɧɬɚ (ɭ ɤɨɬɨɪɨɝɨ ɫɨɞɟɪɠɚɧɢɟ 13Z-ɮɨɪɦɵ ɪɚɜɧɨ 85%). Ɉɞɧɚɤɨ ɨɛɴɟɦɧɵɟ ɡɚɦɟɫɬɢɬɟɥɢ ɜ ɷɬɨɦ ɩɨɥɨɠɟɧɢɢ ɫɧɢɠɚɸɬ ɫɤɨɪɨɫɬɶ ɮɨɬɨɰɢɤɥɚ ɢ ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɩɪɨɬɨɧɧɨɝɨ ɬɪɚɧɫɩɨɪɬɚ.

4.3. Ɍɨɱɟɱɧɵɟ ɡɚɦɟɧɵ ɚɦɢɧɨɤɢɫɥɨɬ. ȼ ɬɟɱɟɧɢɟ ɩɨɫɥɟɞɧɢɯ 10-12 ɥɟɬ ɫ ɰɟɥɶɸ ɢɡɭɱɟɧɢɹ ɜɥɢɹɧɢɹ ɨɩɪɟɞɟɥɟɧɧɵɯ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ ɩɪɨɜɨɞɢɥɢɫɶ ɨɛɲɢɪɧɵɟ ɢɫɫɥɟɞɨɜɚɧɢɹ ɩɨ ɝɟɧɟɬɢɱɟɫɤɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɟɝɨ ɛɟɥɤɨɜɨɣ ɱɚɫɬɢ, ɜ ɪɟɡɭɥɶɬɚɬɟ ɱɟɝɨ ɛɵɥɢ ɩɨɥɭɱɟɧɵ ɢ ɨɯɚɪɚɤɬɟɪɢɡɨɜɚɧɵ ɛɨɥɟɟ ɫɨɬɧɢ ɬɨɱɟɱɧɵɯ ɦɭɬɚɧɬɨɜ ȻɊ (ɫɦ. ɨɛɡɨɪɵ [75, 106]). ɉɪɢ ɷɬɨɦ ɢɫɩɨɥɶɡɨɜɥɢɫɶ ɪɚɡɥɢɱɧɵɟ ɫɩɨɫɨɛɵ ɩɨɥɭɱɟɧɢɹ ɦɭɬɚɧɬɧɨɝɨ ɛɟɥɤɚ: ɯɢɦɢɱɟɫɤɢɦ ɦɭɬɚɝɟɧɟɡ H. salinarum [107], ɝɟɬɟɪɨɥɨɝɢɱɧɚɹ (ɜ E. coli) [108-110] ɢ ɝɨɦɨɥɨɝɢɱɧɚɹ (ɜ H. salinarum) [67, 111] ɷɤɫɩɪɟɫɫɢɹ ɝɟɧɧɨ-ɢɧɠɟɧɟɪɧɵɯ ɤɨɧɫɬɪɭɤɰɢɣ, ɧɟɫɭɳɢɯ ɝɟɧ ȻɊ ɫ ɬɨɱɟɱɧɨɣ ɦɭɬɚɰɢɟɣ ɩɨ ɨɩɪɟɞɟɥɟɧɧɨɣ ɚɦɢɧɨɤɢɫɥɨɬɟ. ɉɨɫɤɨɥɶɤɭ ɦɟɬɨɞɵ ɝɟɧɟɬɢɱɟɫɤɨɣ ɢɧɠɟɧɟɪɢɢ ɛɵɥɢ ɯɨɪɨɲɨ ɪɚɡɪɚɛɨɬɚɧɵ ɩɪɟɠɞɟ ɜɫɟɝɨ ɞɥɹ E. coli, ɫɧɚɱɚɥɚ ɩɪɨɜɨɞɢɥɢ ɢɫɫɥɟɞɨɜɚɧɢɟ ȻɊ, ɷɤɫɩɪɟɫɫɢɪɨɜɚɧɧɨɝɨ ɜ ɞɚɧɧɨɦ ɦɢɤɪɨɨɪɝɚɧɢɡɦɟ ɢ ɪɟɧɚɬɭɪɢɪɨɜɚɧɧɨɝɨ ɞɨɛɚɜɥɟɧɢɟɦ ɥɢɩɢɞɨɜ ɢ ɪɟɬɢɧɚɥɹ [108-110, 112]. Ɉɞɧɚɤɨ ɜ ɷɬɨɦ ɫɥɭɱɚɟ ɞɨɫɬɚɬɨɱɧɨ ɫɥɨɠɧɨ ɨɩɪɟɞɟɥɢɬɶ ɜɤɥɚɞ ɝɟɧɟɬɢɱɟɫɤɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɜ ɢɡɦɟɧɟɧɢɟ ɫɜɨɣɫɬɜ ȻɊ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɜɥɢɹɧɢɟɦ ɦɢɤɪɨɨɤɪɭɠɟɧɢɹ ɱɭɠɟɪɨɞɧɵɯ ɥɢɩɢɞɨɜ, ɢɫɩɨɥɶɡɨɜɚɧɧɵɯ ɩɪɢ ɪɟɧɚɬɭɪɚɰɢɢ. Ɍɚɤ, ɞɚɠɟ ȻɊ ɫ ɩɪɢɪɨɞɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ, ɷɤɫɩɪɟɫɫɢɪɨɜɚɧɧɵɣ ɬɚɤɢɦ ɫɩɨɫɨɛɨɦ, ɨɛɥɚɞɚɟɬ ɩɨɧɢɠɟɧɧɨɣ ɫɬɚɛɢɥɶɧɨɫɬɶɸ, ɟɝɨ Omax ɫɞɜɢɧɭɬ ɜ ɫɬɨɪɨɧɭ ɦɟɧɶɲɢɯ ɞɥɢɧ ɜɨɥɧ, ɧɟɫɤɨɥɶɤɨ ɢɡɦɟɧɹɸɬɫɹ

25 ɩɚɪɚɦɟɬɪɵ ɮɨɬɨɰɢɤɥɚ ɢ ɪɄɚ ɩɟɪɟɯɨɞɨɜ ɜ ɤɢɫɥɭɸ ɫɢɧɸɸ ɢɥɢ ɧɟɚɤɬɢɜɧɭɸ ɳɟɥɨɱɧɭɸ ɮɨɪɦɵ [113]. ɂɫɩɨɥɶɡɨɜɚɧɢɟ ɞɥɹ ɪɟɧɚɬɭɪɚɰɢɢ ȻɊ ɩɨɥɹɪɧɵɯ ɥɢɩɢɞɨɜ H. salinarum (ɞɚɠɟ ɞɥɹ ɫɥɭɱɚɹ ɫ ɞɟɥɢɩɢɞɢɪɨɜɚɧɧɵɦ ȻɊ ɢɡ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ H. salinarum) ɧɟ ɜɵɡɵɜɚɟɬ ɩɨɥɧɨɝɨ ɜɨɫɫɬɚɧɨɜɥɟɧɢɹ ɫɜɨɣɫɬɜ ȻɊ [69, 114]. ȼ ɫɜɹɡɢ ɫ ɷɬɢɦ ɫɬɚɥɢ ɢɫɩɨɥɶɡɨɜɚɬɶ ɦɟɬɨɞ ɝɨɦɨɥɨɝɢɱɧɨɣ ɷɤɫɩɪɟɫɫɢɢ ȻɊ ɜ H. salinarum [67], ɧɟ ɨɛɥɚɞɚɸɳɢɣ ɩɟɪɟɱɢɫɥɟɧɧɵɦɢ ɜɵɲɟ ɧɟɞɨɫɬɚɬɤɚɦɢ. ȼ ɪɟɡɭɥɶɬɚɬɟ ɛɵɥɨ ɩɨɥɭɱɟɧɨ ɢ ɢɫɫɥɟɞɨɜɚɧɨ ɛɨɥɟɟ ɫɨɬɧɢ ɛɟɥɤɨɜ, ɫɨɞɟɪɠɚɳɢɯ ɟɞɢɧɢɱɧɵɟ ɢɥɢ ɦɧɨɠɟɫɬɜɟɧɧɵɟ ɬɨɱɟɱɧɵɟ ɦɭɬɚɰɢɢ. ɉɟɪɜɨɧɚɱɚɥɶɧɨ ɢɡɭɱɚɥɢ ɜɥɢɹɧɢɟ ɬɚɤɢɯ ɦɨɞɢɮɢɤɚɰɢɣ ɧɚ ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ, ɫɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ ɢ ɫɤɨɪɨɫɬɶ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ. ɉɨɡɞɧɟɟ ɛɵɥɢ ɩɨɥɭɱɟɧɵ ɡɚɜɢɫɢɦɨɫɬɢ ɤɢɧɟɬɢɤɢ ɨɛɪɚɡɨɜɚɧɢɹ ɢ ɪɚɫɩɚɞɚ ɪɚɡɥɢɱɧɵɯ ɢɧɬɟɪɦɟɞɢɚɬɨɜ ɮɨɬɨɰɢɤɥɚ (ɜ ɨɫɧɨɜɧɨɦ Ɇ). ȼ ɬɚɛɥɢɰɟ 3 ɩɪɢɜɟɞɟɧɵ ɨɩɭɛɥɢɤɨɜɚɧɧɵɟ ɤ ɧɚɫɬɨɹɳɟɦɭ ɦɨɦɟɧɬɭ ɞɚɧɧɵɟ ɩɨ ɫɜɨɣɫɬɜɚɦ «ɦɭɬɚɧɬɧɵɯ» ȻɊ. ȼɥɢɹɧɢɟ ɝɟɧɟɬɢɱɟɫɤɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɧɚ Omax, ɫɤɨɪɨɫɬɶ ɮɨɬɨɰɢɤɥɚ, ɫɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ ɢ ɇ+-ɬɪɚɧɫɩɨɪɬ ɛɨɥɟɟ ɩɨɞɪɨɛɧɨ ɪɚɫɫɦɨɬɪɟɧɨ ɜ ɪɚɡɞɟɥɟ 4 ɬɟɨɪɟɬɢɱɟɫɤɨɣ ɱɚɫɬɢ ɧɚ ɫ. 64-73. ɂɧɬɟɪɟɫɧɨ ɨɬɦɟɬɢɬɶ, ɱɬɨ ɩɪɢɪɨɞɧɵɣ ȻɊ, ɜɵɞɟɥɟɧɧɵɣ ɢɡ ɝɚɥɨɛɚɤɬɟɪɢɣ ɧɟ ɜɫɬɭɩɚɟɬ ɜ ɪɟɚɤɰɢɸ ɫ ɝɢɞɪɨɤɫɢɥɚɦɢɧɨɦ ɜ ɬɟɦɧɨɬɟ, ɚ ɷɤɫɩɪɟɫɫɢɪɨɜɚɧɧɵɣ ɜ E. coli ȻɊWT ɨɛɟɫɰɜɟɱɢɜɚɟɬɫɹ ɫ W1/2 ɨɤɨɥɨ 9.4 ɱ (ɫɦ. ɬɚɛɥ. 3). ɋɪɟɞɢ ɷɤɫɩɪɟɫɫɢɪɨɜɚɧɧɵɯ ɜ E. coli ɦɭɬɚɧɬɧɵɯ ȻɊ ɨɫɧɨɜɧɚɹ ɱɚɫɬɶ ɨɛɟɫɰɜɟɱɢɜɚɟɬɫɹ ɛɵɫɬɪɟɟ ȻɊWT ɢɥɢ ɫɨ ɫɪɚɜɧɢɦɨɣ ɫɤɨɪɨɫɬɶɸ. Ɉɛɟɫɰɜɟɱɢɜɚɧɢɟ ɡɚɦɟɞɥɟɧɨ ɜ ɧɟɫɤɨɥɶɤɨ ɪɚɡ ɭ ȻɊD85N, ȻɊR82Q, ȻɊR82A ɢ ȻɊD212E, ɢ ɜɫɟ ɷɬɢ ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ (Asp85, Arg82 ɢ Asp212) ɭɱɚɫɬɜɭɸɬ ɜ ɨɛɪɚɡɨɜɚɧɢɢ ɜɧɟɤɥɟɬɨɱɧɨɝɨ ɩɨɥɭɤɚɧɚɥɚ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ. Ʉɢɧɟɬɢɤɚ ɢ ɫɬɟɩɟɧɶ ɜɫɬɪɚɢɜɚɧɢɹ ɪɟɬɢɧɚɥɹ, ɩɪɢ ɨɛɪɚɡɨɜɚɧɢɢ ȻɊ ɪɟɧɚɬɭɪɚɰɢɟɣ ɷɤɫɩɪɟɫɫɢɪɨɜɚɧɧɨɝɨ ɜ E. coli ɛɚɤɬɟɪɢɨɨɩɫɢɧɚ, ɬɚɤ ɠɟ ɡɚɜɢɫɹɬ ɨɬ ɩɨɥɨɠɟɧɢɹ ɢɡɦɟɧɟɧɧɨɝɨ ɚɦɢɧɨɤɢɫɥɨɬɧɨɝɨ ɨɫɬɚɬɤɚ ɜ ɟɝɨ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ. ȼ ɛɨɥɶɲɢɧɫɬɜɟ ɫɥɭɱɚɟɜ ɜɪɟɦɹ ɪɟɝɟɧɟɪɚɰɢɢ ɩɢɝɦɟɧɬɚ ɢɡɦɟɧɹɟɬɫɹ ɜ ɧɟɫɤɨɥɶɤɨ ɪɚɡ. ɇɚɢɛɨɥɶɲɟɟ ɡɚɦɟɞɥɟɧɢɟ ɩɪɨɰɟɫɫɚ ɜɫɬɪɚɢɜɚɧɢɹ, ɩɪɚɤɬɢɱɟɫɤɢ ɧɚ 2 ɩɨɪɹɞɤɚ, ɩɪɨɢɫɯɨɞɢɬ ɩɪɢ ɡɚɦɟɧɟ M118ȿ (ɷɬɨɬ ɨɫɬɚɬɨɤ ɪɚɫɩɨɥɨɠɟɧ ɨɤɨɥɨ ɤɨɥɶɰɚ ɪɟɬɢɧɚɥɹ, ɪɹɞɨɦ ɫ 5- ɢ 9ɋɇ3), ɜ ɬɨ ɠɟ ɜɪɟɦɹ ɡɚɦɟɧɚ Ɇ118Ⱥ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɦɟɧɹɟɬ ɫɤɨɪɨɫɬɢ ɪɟɝɟɧɟɪɚɰɢɢ.

WT

H. sal. H. sal. H. sal. H. sal. E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli

H. sal.

H. sal H. sal.

1 2.0 1.8 1.6 1.0 1.6 1.0 1.0

2.2

2.1 1.83 1.0

1.52 1.3 1.2

ɦɢɧ

84 80

84

84

84

75

ɪɟɝ.

Ɋɟɝɟɧɟɪɚɰɢɹ Ɇɢɤɪɨ- ȻɊ ɫ all-EɆɭɬɚɨɪɝɚ- ɪɟɬɢɧɚɥɟɦ ɰɢɹ ɧɢɡɦ W1/2, %

549 552 551 551 550 551 549 551 551 548 551 551 551 551 551

558 558 560 560 559 548 551

561 560

DA

561 559 561 561

559 557 561 561 557

556 561 568 559 559 561

568 568 570 568

570 569 568

LA

Omax, ɧɦ

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1

ɧɚɱ.

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1

1 1 1

ɩɨɫɬ

+ ɇ ɬɪɚɧɫɩɨɪɬ*2

11.3

11.3

2.6 3.2 13.3

ɪɄ

60/40

SB551/544

40/60

40/60 39/61

42/58

36/64

54/46

DA

98/2

97/3

97/3 94/6

94/6

96/4

LA

0.9

1 0.8

1.3

13

8

9 9.1

13

71

4.1

4.8 6.4

57,5

25 60

3.1

0.3

6.0

5

150

7.9

170

12

86.3

6.4

5.7 3.0

Wɛɵɫɬɪ Wɦɟɞɥ Wɫɪɟɞɧ Wɛɵɫɬɪ Wɦɟɞɥ Wɫɪɟɞɧ

ɋɨɨɬɧɨɲɟɧɢɟ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬ, ɫɪɟɞɧɟɟ ɜɪɟɦɹ*4 ɢɡɨɦɟɪɨɜ Ɉɛɪɚɡɨɜɚɧɢɟ, ɦɤɫ Ɋɚɫɩɚɞ, ɦɫ all-E/13Z

SB551/544

D85 D85568/605 SB568/460

ɩɟɪɟɯɨɞ

ɪɇ-ɡɚɜɢɫɢɦɵɟ ɩɟɪɟɯɨɞɵ*3

ɋɜɨɣɫɬɜɚ ɝɟɧɧɨ-ɢɧɠɟɧɟɪɧɵɯ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɨɜ, ɫɨɞɟɪɠɚɳɢɯ ɨɞɧɭ ɬɨɱɟɱɧɭɸ ɦɭɬɚɰɢɸ.*1

26

9.42

9.4

W1/2, ɱ

ɋ / 7.0 C/ 6.8 C / 7.0 C / 6.9 A / 6.0 A / 6.0 A / 6.0 ȼ/ 6.0 Ⱥ / 6.0 A / 6.0 A / 6.0 Ⱥ / 6.0 Ⱥ / 7.3 Ⱥ / 6.0 Ⱥ / 6.0 A / 6.0 A / 6.2. Ⱥ / 7.2 Ⱥ / 6.0 A / 6.0 A / 6.0 Ⱥ / 6.0 Ⱥ / 6.0

ɋ / 6.9 ɋ / 8.8 ɋ / 5.6

117 118 119 120 62 66 68 69 72 73 74 115 121 122 123 124 125 126 127 128 129 130 131

67 71 116

Ɋ-ɰɢɹ ɫ NH2OH, ɉɪɢɦɟ[115], ɱɚɧɢɹ*5 Ʌɢɬ.

Ɍɚɛɥɢɰɚ 3.

L52A A53G T55A

P50G

P50A

T47V L48A V49A V49L

T46V

R7Q W10C W10F W12F T17V M20A M20E T24V D36N D38N A44G I45A

E. coli H. sal. E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli H. sal. H. sal. E. coli E. coli E. coli H. sal. E. coli E. coli E. coli E. coli E. coli E. coli H. sal. E. coli H. sal.

48 44 53

1.3 1.2 2.6 3.1 1.6 1.0 1.3 1.3

8.4

0.1

0.1 2.6 31

5.6 2.3

71

3.7

72 45

41 60 72

ɪɟɝ.

ɦɢɧ


558 544 558

549 548 548 551 551 551 550 549 550 553 559 558 550 553 550 558 549 551 546 546 546

DA

552 554 567 548 567

559 559 567 549 556 551

558 562 558 561 560 561 556 559 560 563 569 567

LA

Omax, ɧɦ

1.2

0.5

1 0.9 0.6

0,8 1

0.8 0.7 1 0.8 1 1 1.1 1

1

ɧɚɱ.

1.2

1

1 0.6 0.5

0,7 1

0.8 0.7 1 0.7 1 1 1.1 1.1

0.8

ɩɨɫɬ


ɪɄ

ɩɟɪɟɯɨɞ


27

47/53

65/35 46/47

43/57

49/51 45/55

46/54

DA

76/15

71/18 89/7

90/10

87/13 83/17

91/9

LA

7.1

0.5 0.8

2.9

0.4

11.8

1.0 2.8

22.7

1.6

14.2

6.3

167.4 100.6 11.2 8.0

23

3.8

0.3

0.6

0.1

1.0

0.5

0.8 1.6

15.3

8.3

5.9

14

1700

16.8 15.6

12.3

6.6

4.7

2.8 7.5

85.5

13.6 9.6



4.10

W1/2, ɱ

Ⱥ / 6.0 ɋ / 7.0 A / 6.0 A / 6.0 Ⱥ / 6.0 Ⱥ / 6.2 Ⱥ / 6.2 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 C / 6.9 C / 6.9 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 C / 6.9 Ⱥ / 6.2 Ⱥ / 6.2 Ⱥ / 6.0 ȼ / 6.0 Ⱥ / 6.0 ȼ / 6.0 C / 6.9 Ⱥ / 6.2 C / 6.9

72 117 74 74 130 125 125 130 131 131 120 120 115 130 130 120 125 125 68 69 68 69 120 125 120


Ɍɚɛɥɢɰɚ 3. ɉɪɨɞɨɥɠɟɧɢɟ.

H. sal.

Y83N

(-)Ⱥ84 E. coli

H. sal.

E. coli E. coli E. coli H. sal. E. coli.

Y83F

R82Q

E. coli

H. sal.

R82H

R82G

R82A

E. coli H. sal. E. coli E. coli E. coli E. coli E. coli E. coli E. coli E. coli H. sal. H. sal. E. coli

T55V Y57N S59A G72C E74C N76C Y79C T80F

82

1.0 1.91 2.9 1.6 3.52 2.3

70

73

3.4

13

63

3.2

4.6

56

1.0

92

ɪɟɝ.

ɦɢɧ


566

563

555

567

580 580 580 550

584 553

0.1 0.6

0.3

0.1 0.4

566 597 565

568

0.7

1 1.1

1

ɧɚɱ.

558

560 560 560 559

LA

584 548 559 547 575 580 575 547

550 555 551 548 551 552 525 549 547 564 597 555 554 558

DA

Omax, ɧɦ

0

0.6

0.4 0

1 0.9 1 0.9 1 0.5 1.2

1

ɩɨɫɬ


SB590/540

ɩɟɪɟɯɨɞ

2.6 8.2 3.4 8.8 6.6

11.2

50/50

39/61

DA

72/22

79/21

75/25

LA

0.4

61

1.0

0.2

7

200

30

3

5 5

80

2.0

0.1

46

8.2 a100 153

0.7

20

25

100

1.2

60

11

17

5 a40 0.7


2.76

26.5

26.9

W1/2, ɱ

Ⱥ / 6.0

C/ 6.8

Ⱥ / 6.0 Ⱥ / 7.3 Ⱥ / 6.0 ɋ / 6.9 Ⱥ / 6.0 C/ 6.8

ȼ / 6.0

Ⱥ / 6.0 ɋ / 7.0 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 A / 6.0 Ⱥ / 7.3 Ⱥ / 6.4 ɋ / 6.5 ɋ / 8.8 ȼ / 7.9 ɋ / 6.8

124

118

66 121 128 67 72 118

64

108 117 108 73 127 127 127 74 121 123 67 71 132 76


Ɍɚɛɥɢɰɚ 3. ɉɪɨɞɨɥɠɟɧɢɟ. ɋɨɨɬɧɨɲɟɧɢɟ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬ, ɫɪɟɞɧɟɟ ɜɪɟɦɹ*4 ɢɡɨɦɟɪɨɜ Ɉɛɪɚɡɨɜɚɧɢɟ, ɦɤɫ Ɋɚɫɩɚɞ, ɦɫ all-E/13Z

D85547/568 SB548/480 D85555/584 SB555/480 SB566/395 84/16

SB575/365

7.2 SB585/554 582/523 a7.5 SB 1.0 D85558/622 9.4 SB568/572 7.0 SB584/548

6.4

ɪɄ


28

T89A

W86F

D85N

D85H

E. coli E. coli E. coli E. coli E. coli E. coli H. sal. E. coli E. coli E. coli

E. coli

53

85

1

38

84

34

75

57

76

ɪɟɝ.

41

20 170

(+)Ⱥ85 E. coli 11 H. sal. E. coli D85A E. coli E. coli E. coli 50.8 D85C E. coli 124.3 E. coli 100 70

0.4

17



D85620/568 82/18 SB568/475 D85566/620

555/440 SB560/365

SB582/510 SB

ɩɟɪɟɯɨɞ


9.86

8.25

1.41

34.0

W1/2, ɱ

ɩɨɫɤɨɥɶɤɭ ɚɛɫɨɥɸɬɧɵɟ ɞɚɧɧɵɟ ɡɚɜɢɫɹɬ ɨɬ ɭɫɥɨɜɢɣ ɩɪɨɜɟɞɟɧɢɹ ɷɤɫɩɟɪɢɦɟɧɬɚ.

Ⱦɥɹ ɦɭɬɚɧɬɧɵɯ ȻɊ ɩɪɢɜɟɞɟɧɨ ɨɬɧɨɫɢɬɟɥɶɧɨɟ ɡɧɚɱɟɧɢɟ ɚɤɬɢɜɧɨɫɬɢ (ɤɨɥɢɱɟɫɬɜɨ ɩɪɨɬɨɧɨɜ, ɩɟɪɟɧɨɫɢɦɵɯ WT-ȻɊ ɩɪɢɧɹɬɨ ɡɚ ɟɞɢɧɢɰɭ),

ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ), ɩɨɥɭɱɟɧɧɵɯ ɢ ɢɫɫɥɟɞɨɜɚɧɧɵɯ ɜ ɚɧɚɥɨɝɢɱɧɵɯ ɭɫɥɨɜɢɹɯ.

72 130 115 132 72

134

ɋ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 ȼ / 7.9 Ⱥ / 6.0

64 121 123 128 131 67 121 123 128 131 67 130 116

ȼ / 8.0 Ⱥ / 7.3 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 ɋ / 6.9 Ⱥ / 7.3 Ⱥ / 6.0 Ⱥ / 6.0 Ⱥ / 6.0 ɋ / 6.9 Ⱥ / 6.0 ɋ / 6.0



ȼ ɧɚɱɚɥɟ ɬɚɛɥɢɰɵ ɞɥɹ ɜɨɡɦɨɠɧɨɫɬɢ ɫɪɚɜɧɟɧɢɹ ɫɜɨɣɫɬɜ ɩɪɢɪɨɞɧɵɯ ɢ ɦɭɬɚɧɬɧɵɯ ȻɊ, ɩɪɢɜɟɞɟɧɵ ɫɜɨɣɫɬɜɚ WT - ȻɊ (ɫ ɩɪɢɪɨɞɧɨɣ

R225Q E. coli E. coli S226A E. coli R227G E. coli R227Q E. coli

K216G H. sal.

K216A H. sal.

E. coli E. coli E. coli D212E E. coli E. coli H. sal. E. coli E. coli D212N E. coli E. coli H. sal. S214A E. coli

ɦɢɧ


34

*5

*4

*3

A Bf e

k f t

Bs e k st , ɝɞɟ kf, ks – ɤɨɧɫɬɚɧɬɵ ɫɤɨɪɨɫɬɢ ɛɵɫɬɪɨɣ ɢ ɦɟɞɥɟɧɧɨɣ ɤɨɦɩɨɧɟɧɬ

ɥɢɩɨɫɨɦɵ ɢɡ ɮɨɫɮɨɥɢɩɢɞɨɜ ɫɨɢ.

ɋ – ɢɫɫɥɟɞɨɜɚɥɢ ɦɟɦɛɪɚɧɵ, ɜɵɞɟɥɟɧɵɟ ɢɡ ɝɟɧɧɨ-ɢɧɠɟɧɟɪɧɨɝɨ ɲɬɚɦɦɚ H. salinarum, ɩɟɪɟɧɨɫ ɩɪɨɬɨɧɚ ɢɡɦɟɪɹɥɢ ɩɨɫɥɟ ɩɟɪɟɜɟɞɟɧɢɹ ɢɯ ɜ

H. salinarum ɢ ɢɫɫɥɟɞɨɜɚɥɢ ɢɯ ɫɜɨɣɫɬɜɚ, ɩɟɪɟɧɨɫ ɩɪɨɬɨɧɚ ɢɡɦɟɪɹɥɢ ɩɨɫɥɟ ɩɟɪɟɜɟɞɟɧɢɹ ȻɊ ɜ ɥɢɩɨɫɨɦɵ ɢɡ ɫɨɟɜɵɯ ɥɢɩɢɞɨɜ [64, 132, 63],

131], ȼ – ȻɊ, ɩɨɥɭɱɚɟɦɵɣ ɪɟɝɟɧɟɪɚɰɢɟɣ ɷɤɫɩɪɟɫɫɢɪɨɜɚɧɧɨɝɨ ɜ E. coli ȻɈ ɫ ɪɟɬɢɧɚɥɟɦ, ɩɟɪɟɜɨɞɢɥɢ ɜ ɜɟɡɢɤɭɥɵ ɩɨɥɹɪɧɵɯ ɥɢɩɢɞɨɜ

ɨɤɬɢɥɝɥɸɤɨɡɢɞɧɵɦ ɦɟɬɨɞɨɦ ɜ ɥɢɩɨɫɨɦɵ ɢɡ ɫɨɟɜɵɯ ɥɢɩɢɞɨɜ [68, 73, 66, 122, 124, 126, 127] ɢɥɢ ɜ ɚɡɨɥɟɤɬɢɧɨɜɵɟ ɥɢɩɨɫɨɦɵ [125, 129,

ɢɫɫɥɟɞɨɜɚɧɢɹ ɩɪɨɜɨɞɢɥɢ ɜ ɥɢɩɨɫɨɦɚɯ 1% DMPC, 1% CHAPS 0.2 % SDS, ɚ ɩɟɪɟɧɨɫ ɩɪɨɬɨɧɚ ɨɩɪɟɞɟɥɹɥɢ ɩɨɫɥɟ ɩɟɪɟɜɨɞɚ ȻɊ

ɍɫɥɨɜɢɹ, ɜ ɤɨɬɨɪɵɯ ɩɪɨɜɨɞɢɥɢɫɶ ɢɡɦɟɪɟɧɢɹ: ɥɢɩɢɞɧɚɹ ɫɢɫɬɟɦɚ / ɪɇ. Ʌɢɩɢɞɧɵɟ ɫɢɫɬɟɦɵ: A – ɪɟɝɟɧɟɪɚɰɢɸ ȻɊ ɢ ɫɩɟɤɬɪɚɥɶɧɵɟ

Ɍɟɦɩɟɪɚɬɭɪɚ ɩɪɢ ɤɨɬɨɪɨɣ ɩɪɨɜɨɞɢɥɨɫɶ ɢɫɫɥɟɞɨɜɚɧɢɟ ɤɢɧɟɬɢɤɢ ɮɨɬɨɰɢɤɥɚ: 20qɋ – [76, 125, 130], ɧɟɬ ɞɚɧɧɵɯ – [69, 121, 118].

ɤɨɧɫɬɚɧɬɵ ɫɤɨɪɨɫɬɢ [128, 63, 69, 121, 76, 125, 126, 130, 118], ɫɪɟɞɧɟɟ ɜɪɟɦɹ ɩɪɨɰɟɫɫɨɜ ɛɵɥɨ ɪɚɫɫɱɢɬɚɧɨ ɩɨ ɩɪɢɜɟɞɟɧɧɨɣ ɮɨɪɦɭɥɟ.

Wɦɟɞɥ=1/ks, Wɫɪɟɞɧ=(Af/kf)+( As/ks). Ⱦɥɹ ɩɪɢɜɟɞɟɧɢɹ ɞɚɧɧɵɯ ɤ ɟɞɢɧɨɨɛɪɚɡɧɨɦɭ ɜɢɞɭ ɜ ɫɥɭɱɚɟ, ɟɫɥɢ ɜ ɢɫɬɨɱɧɢɤɟ ɩɪɢɜɟɞɟɧɵ ɬɨɥɶɤɨ

ɭɪɚɜɧɟɧɢɟɦ, ɫɪɟɞɧɟɟ ɜɪɟɦɹ ɩɪɨɰɟɫɫɚ ɨɛɪɚɡɨɜɚɧɢɹ ɢɥɢ ɪɚɫɩɚɞɚ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɛɵɥɨ ɪɚɫɫɱɢɬɚɧɨ ɫɥɟɞɭɸɳɢɦ ɨɛɪɚɡɨɦ: Wɛɵɫɬɪ=1/kf,

ɩɪɨɰɟɫɫɚ, ɫɜɹɡɚɧɧɵɟ ɫ ɢɯ ɨɬɧɨɫɢɬɟɥɶɧɵɦɢ ɚɦɩɥɢɬɭɞɚɦɢ ɡɚɜɢɫɢɦɨɫɬɶɸ Af=Bf/(Bf+Bs) ɢ As=Bs/(Bf+Bs). ȼ ɫɨɨɬɜɟɬɫɬɜɢɢ ɫ ɞɚɧɧɵɦ

ɩɪɨɰɟɫɫɚ, ɫɜɹɡɚɧɧɵɟ ɫɨ ɫɪɟɞɧɢɦ ɜɪɟɦɟɧɟɦ ɡɚɜɢɫɢɦɨɫɬɶɸ k=1/Wɫɪ, Bf, Bs – ɫɨɨɬɜɟɬɫɬɜɭɸɳɢɟ ɤɨɧɫɬɚɧɬɵ ɛɵɫɬɪɨɣ ɢ ɦɟɞɥɟɧɧɨɣ ɤɨɦɩɨɧɟɧɬ

ɨɩɢɫɚɬɶ ɫɥɟɞɭɸɳɢɦ ɭɪɚɜɧɟɧɢɟɦ [69]: A405 (t )

Ʉɢɧɟɬɢɤɭ ɨɛɪɚɡɨɜɚɧɢɹ ɢ ɪɚɫɩɚɞɚ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ, ɪɟɝɢɫɬɪɢɪɭɟɦɭɸ ɩɨ ɢɡɦɟɧɟɧɢɸ ɩɨɝɥɨɳɟɧɢɹ ɩɪɢ 400-410 ɧɦ, ɦɨɠɧɨ

ɢɫɩɨɥɶɡɭɸɬ ɞɥɹ ɨɰɟɧɤɢ ɟɟ ɪɄɚ [76].

ɧɟɚɤɬɢɜɧɭɸ ɳɟɥɨɱɧɭɸ ɮɨɪɦɭ ɫ Omax 480 ɧɦ, ɱɬɨ ɫɜɹɡɵɜɚɸɬ ɫ ɞɟɩɪɨɬɨɧɢɪɨɜɚɧɢɟɦ ɩɪɨɬɨɧ-ɜɵɞɟɥɹɸɳɟɣ ɝɪɭɩɩɵ ɛɟɥɤɚ (SB) ɢ

ȻɊWT), ɤɨɬɨɪɨɟ ɫɜɹɡɚɧɨ ɫ ɩɪɨɬɨɧɢɪɨɜɚɧɢɟɦ Asp-85 ɢ ɢɫɩɨɥɶɡɭɟɬɫɹ ɞɥɹ ɨɰɟɧɤɢ ɟɟ ɪɄɚ [76]. ɉɨɜɵɲɟɧɢɟ ɪɇ ɩɪɢɜɨɞɢɬ ɤ ɩɟɪɟɯɨɞɭ ȻɊ ɜ

ɍɦɟɧɶɲɟɧɢɟ ɪɇ ɫ 7 ɞɨ 2 ɜɵɡɵɜɚɟɬ ɩɟɪɟɯɨɞ ȻɊ ɜ ɤɢɫɥɨɬɧɭɸ ɫɢɧɸɸ ɮɨɪɦɭ (Omax ɫ 552 ɢɡɦɟɧɹɟɬɫɹ ɞɨ 605 ɧɦ ɜ ɬɟɦɧɨ-ɚɞɚɩɬɢɪɨɜɚɧɧɨɦ

35

36

5. Ɍɟɯɧɢɱɟɫɤɨɟ ɩɪɢɦɟɧɟɧɢɟ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. ɉɪɚɤɬɢɱɟɫɤɢ ɫ ɦɨɦɟɧɬɚ ɨɬɤɪɵɬɢɹ ȻɊ ɧɚɱɚɥɢɫɶ ɢɫɫɥɟɞɨɜɚɧɢɹ ɜɨɡɦɨɠɧɨɫɬɢ ɟɝɨ ɩɪɢɦɟɧɟɧɢɹ ɜ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɯ ɰɟɥɹɯ. ȼ ɪɟɡɭɥɶɬɚɬɟ ɛɵɥ ɩɪɟɞɥɨɠɟɧ ɪɹɞ ɩɨɞɯɨɞɨɜ, ɨɫɧɨɜɚɧɧɵɯ ɧɚ ɢɫɩɨɥɶɡɨɜɚɧɢɢ ɞɜɭɯ ɨɫɧɨɜɧɵɯ ɫɜɨɣɫɬɜ ȻɊ: xɩɟɪɟɧɨɫ ɩɪɨɬɨɧɚ - ɞɥɹ ɫɢɧɬɟɡɚ ȺɌɎ ɜ ɛɢɨɪɟɚɤɬɨɪɚɯ, ɨɛɟɫɫɨɥɢɜɚɧɢɹ ɜɨɞɵ, ɩɪɟɜɪɚɳɟɧɢɹ ɫɜɟɬɨɜɨɣ ɷɧɟɪɝɢɢ ɜ ɷɥɟɤɬɪɢɱɟɫɤɭɸ, ɭɥɶɬɪɚɛɵɫɬɪɨɣ ɞɟɬɟɤɰɢɢ ɫɜɟɬɚ ɜ ɛɢɨɫɟɧɫɨɪɧɵɯ ɭɫɬɪɨɣɫɬɜɚɯ xɮɨɬɨɯɪɨɦɧɵɟ ɫɜɨɣɫɬɜɚ ɛɟɥɤɚ – ɩɪɢ ɫɨɡɞɚɧɢɢ ɫɪɟɞ ɞɥɹ ɯɪɚɧɟɧɢɹ ɢɧɮɨɪɦɚɰɢɢ (ɜ ɰɢɮɪɨɜɨɣ ɢ ɚɧɚɥɨɝɨɜɨɣ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɣ ɮɨɪɦɟ) ɢ ɨɛɪɚɛɨɬɤɢ ɢɧɮɨɪɦɚɰɢɢ (ɤɨɧɞɢɰɢɨɧɢɪɨɜɚɧɢɟ ɫɢɝɧɚɥɨɜ, ɚɫɫɨɰɢɚɬɢɜɧɚɹ ɩɚɦɹɬɶ, ɪɚɫɩɨɡɧɚɜɚɧɢɟ ɢɡɨɛɪɚɠɟɧɢɣ, ɤɨɦɩɨɧɟɧɬɵ ɧɟɣɪɨɤɨɦɩɶɸɬɟɪɨɜ). Ɉɞɧɚɤɨ ɨɤɚɡɚɥɨɫɶ, ɱɬɨ ɢɫɩɨɥɶɡɨɜɚɧɢɸ ȻɊ ɜ ɤɚɱɟɫɬɜɟ ɩɪɟɨɛɪɚɡɨɜɚɬɟɥɹ ɫɜɟɬɨɜɨɣ ɷɧɟɪɝɢɢ ɩɪɟɩɹɬɫɬɜɭɟɬ ɪɹɞ ɮɚɤɬɨɪɨɜ, ɧɚɩɪɢɦɟɪ, ɬɟɯɧɢɱɟɫɤɚɹ ɧɟɜɨɡɦɨɠɧɨɫɬɶ ɫɨɡɞɚɜɚɬɶ ɫɬɚɛɢɥɶɧɵɟ ɩɥɟɧɤɢ ȻɊ ɛɨɥɶɲɢɯ ɪɚɡɦɟɪɨɜ ɫ ɪɚɜɧɨɦɟɪɧɵɦɢ ɫɜɨɣɫɬɜɚɦɢ ɩɨ ɩɨɜɟɪɯɧɨɫɬɢ, ɚ ɝɥɚɜɧɨɟ, ɧɢɡɤɨɣ ɷɮɮɟɤɬɢɜɧɨɫɬɶɸ ɪɚɡɪɚɛɨɬɚɧɧɵɯ ɫɢɫɬɟɦ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɭɠɟ ɢɦɟɸɳɢɦɢɫɹ ɭɫɬɪɨɣɫɬɜɚɦɢ, ɜɵɫɨɤɨɣ ɫɬɨɢɦɨɫɬɶɸ ɪɟɚɥɢɡɚɰɢɢ ɩɪɨɰɟɫɫɨɜ ɢ ɞɪ. ȼ ɫɜɹɡɢ ɫ ɷɬɢɦ ɨɫɧɨɜɧɨɣ ɨɛɥɚɫɬɶɸ ɞɥɹ ɬɟɯɧɢɱɟɫɤɨɝɨ ɩɪɢɦɟɧɟɧɢɹ ȻɊ, ɫɬɚɥɚ ɨɩɬɢɱɟɫɤɚɹ ɨɛɪɚɛɨɬɤɚ ɞɚɧɧɵɯ. Ⱦɥɹ ɷɬɢɯ ɰɟɥɟɣ ɭɫɬɪɨɣɫɬɜɚ ɧɚ ɨɫɧɨɜɟ ȻɊ ɨɛɥɚɞɚɸɬ ɪɹɞɨɦ ɩɪɟɢɦɭɳɟɫɬɜ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɭɠɟ ɢɡɜɟɫɬɧɵɦɢ ɚɧɚɥɨɝɚɦɢ. ɇɚɩɪɢɦɟɪ, ɩɪɢ ɨɛɪɚɛɨɬɤɟ ɢɡɨɛɪɚɠɟɧɢɹ ɜ ɪɟɚɥɶɧɨɦ ɜɪɟɦɟɧɢ ɬɪɟɛɭɟɬɫɹ ɩɪɢɛɥɢɡɢɬɟɥɶɧɨ 106 ɰɢɤɥɨɜ WRE (ɡɚɩɢɫɶɱɬɟɧɢɟ-ɫɬɢɪɚɧɢɟ) ɡɚ ɞɟɧɶ ɪɚɛɨɬɵ, ɱɬɨ ɜɧɟ ɜɨɡɦɨɠɧɨɫɬɟɣ ɛɨɥɶɲɢɧɫɬɜɚ ɫɢɧɬɟɬɢɱɟɫɤɢɯ ɨɪɝɚɧɢɱɟɫɤɢɯ ɮɨɬɨɯɪɨɦɧɵɯ ɦɚɬɟɪɢɚɥɨɜ. Ʉɪɨɦɟ ɬɨɝɨ, ɩɨɫɤɨɥɶɤɭ ɜ ɨɤɪɭɠɚɸɳɟɣ ɫɪɟɞɟ ɧɚɯɨɞɢɬɫɹ ɛɨɥɶɲɨɟ ɱɢɫɥɨ ɞɨɫɬɚɬɨɱɧɨ ɦɨɳɧɵɯ ɢɫɬɨɱɧɢɤɨɜ ɪɚɞɢɨɢɡɥɭɱɟɧɢɹ ɪɚɡɥɢɱɧɨɣ ɩɪɢɪɨɞɵ, ɜɫɬɚɥ ɜɨɩɪɨɫ ɨ ɧɚɞɟɠɧɨɫɬɢ ɪɚɛɨɬɵ ɤɨɦɩɶɸɬɟɪɧɵɯ ɫɢɫɬɟɦ, ɢɫɩɨɥɶɡɭɟɦɵɯ ɧɚ ɜɚɠɧɵɯ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɯ ɢ ɜɨɟɧɧɵɯ ɨɛɴɟɤɬɚɯ, ɢ ɨ ɧɟɨɛɯɨɞɢɦɨɫɬɢ ɢɯ ɡɚɳɢɬɵ ɨɬ ɜɧɟɲɧɢɯ ɜɨɡɞɟɣɫɬɜɢɣ. ȼ ɷɬɢɯ ɭɫɥɨɜɢɹɯ ɛɢɨɦɨɥɟɤɭɥɹɪɧɵɟ ɭɫɬɪɨɣɫɬɜɚ ɨɤɚɡɵɜɚɸɬɫɹ ɛɨɥɟɟ ɧɚɞɟɠɧɵɦɢ ɢ ɧɟ ɬɪɟɛɭɸɬ ɬɚɤɢɯ ɦɨɳɧɵɯ ɫɢɫɬɟɦ ɡɚɳɢɬɵ, ɤɚɤ ɩɨɥɭɩɪɨɜɨɞɧɢɤɨɜɵɟ, ɱɬɨ ɩɨɡɜɨɥɹɟɬ ɡɧɚɱɢɬɟɥɶɧɨ ɭɦɟɧɶɲɢɬɶ ɝɚɛɚɪɢɬɵ ɢ ɦɚɫɫɭ ɤɨɦɩɶɸɬɟɪɨɜ, ɢ ɢɦɟɟɬ ɛɨɥɶɲɨɟ ɡɧɚɱɟɧɢɟ, ɧɚɩɪɢɦɟɪ, ɞɥɹ ɤɨɫɦɢɱɟɫɤɢɯ ɬɟɯɧɨɥɨɝɢɣ. ɗɬɨ ɹɜɥɹɟɬɫɹ ɞɨɩɨɥɧɢɬɟɥɶɧɵɦ ɫɬɢɦɭɥɨɦ ɜ ɪɚɡɪɚɛɨɬɤɟ ɛɢɨɦɨɥɟɤɭɥɹɪɧɵɯ ɭɫɬɪɨɣɫɬɜ [135]. ȼ ɬɚɛɥ. 4 ɩɟɪɟɱɢɫɥɟɧɵ ɧɟɤɨɬɨɪɵɟ ɜɚɪɢɚɧɬɵ ɩɪɢɦɟɧɟɧɢɹ ȻɊ ɞɥɹ ɫɨɡɞɚɧɢɹ ɨɩɬɢɱɟɫɤɢɯ ɭɫɬɪɨɣɫɬɜ ɢ ɩɪɢɜɟɞɟɧɵ ɞɚɧɧɵɟ ɩɨ ɧɟɨɛɯɨɞɢɦɨɫɬɢ ɞɥɹ ɢɯ ɪɟɚɥɢɡɚɰɢɢ ɧɟɤɨɬɨɪɵɯ ɫɩɟɰɢɮɢɱɟɫɤɢɯ ɫɜɨɣɫɬɜ ȻɊ.

37 Ɍɚɛɥɢɰɚ 4. ɂɫɩɨɥɶɡɨɜɚɧɢɟ ɪɚɡɥɢɱɧɵɯ ɫɜɨɣɫɬɜ ȻɊ ɜ ɟɝɨ ɬɟ ɯɧɢɱɟɫɤɢɯ ɩɪɢɦɟɧɟɧɢɹɯ [2]* ɉɪɨɫɬɪɚɧɫɬɜɟɧɧɵɟ ɫɜɟɬɨɦɨɞɭɥɹɬɨɪɵ

Ɉɛɪɚɳɟɧɢɟ ɜɨɥɧɨɜɨɝɨ ɮɪɨɧɬɚ

Ɋɚɫɩɨɡɧɚ ɜɚɧɢɟ ɨɛɪɚɡɨɜ

ɂɧɬɟɪɮɟɪɨɦɟɬɪɢɹ

ɂɡɦɟɧɟɧɢɟ ɩɨɝɥɨɳɟɧɢɹ + í í ɂɡɦɟɧɟɧɢɟ ɩɨɤɚɡɚɬɟɥɹ í ++ + ɩɪɟɥɨɦɥɟɧɢɹ ɋɜɟɬɨɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɶ í í í ɇɟɥɢɧɟɣɧɨɫɬɶ í +í ++ ɨɩɬɢɱɟɫɤɢɯ ɫɜɨɣɫɬɜ Ɍɟɩɥɨɜɚɹ ɪɟɥɚɤɫɚɰɢɹ + + í Ⱦɜɚ ɮɨɬɨɚɤɬɢɜɧɵɯ ++ ++ í ɫɨɫɬɨɹɧɢɹ ȼɨɡɦɨɠɧɨɫɬɶ ɩɨɥɹɪɢí í ++ ɡɚɰɢɢ ɡɚɩɢɫɢ Ɉɛɪɚɬɢɦɨɫɬɶ ++ ++ ++ * «+ +» – ɧɟɨɛɯɨɞɢɦɨ, «+» – ɠɟɥɚɬɟɥɶɧɨ, «í» – ɧɟ ɢɦɟɟɬ ɡɧɚɱɟɧɢɹ.

ɇɟɥɢɧɟɣɧɚɹ ɮɢɥɶɬɪɚɰɢɹ

í

+

++

í

++

í

í

++

í

++

++

++

++

í

++

++

5.1. Ȼɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ ɤɚɤ ɮɨɬɨɯɪɨɦɧɵɣ ɦɨɥɟɤɭɥɹɪɧɵɣ ɦɚɬɟɪɢɚɥ. Ɉɫɧɨɜɧɵɟ ɬɪɟɛɨɜɚɧɢɹ ɤ ɦɚɬɟɪɢɚɥɚɦ ɞɥɹ ɦɨɥɟɤɭɥɹɪɧɨɣ ɷɥɟɤɬɪɨɧɢɤɢ ɫɨɫɬɨɹɬ ɜ ɬɨɦ, ɱɬɨ ɨɧɢ ɞɨɥɠɧɵ ɨɛɪɚɬɢɦɨ ɢɡɦɟɧɹɬɶ ɫɜɨɸ ɫɬɪɭɤɬɭɪɭ ɜ ɨɬɜɟɬ ɧɚ ɧɟɤɨɟ ɮɢɡɢɱɟɫɤɨɟ ɜɨɡɞɟɣɫɬɜɢɟ ɢ ɫɭɳɟɫɬɜɨɜɚɬɶ ɩɨ ɤɪɚɣɧɟɣ ɦɟɪɟ ɜ ɞɜɭɯ ɫɨɫɬɨɹɧɢɹɯ, ɤɨɬɨɪɵɟ ɡɚɦɟɬɧɨ ɪɚɡɥɢɱɚɸɬɫɹ ɩɨ ɥɟɝɤɨ ɢɡɦɟɪɹɟɦɵɦ ɮɢɡɢɱɟɫɤɢɦ ɯɚɪɚɤɬɟɪɢɫɬɢɤɚɦ (ɧɚɩɪɢɦɟɪ, ɩɨ ɫɩɟɤɬɪɚɥɶɧɵɦ ɢɥɢ ɨɩɬɢɱɟɫɤɢɦ ɫɜɨɣɫɬɜɚɦ). ɏɨɬɹ ɤ ɧɚɫɬɨɹɳɟɦɭ ɜɪɟɦɟɧɢ ɪɚɡɪɚɛɨɬɚɧ ɰɟɥɵɣ ɧɚɛɨɪ ɮɨɬɨɯɪɨɦɧɵɯ ɨɪɝɚɧɢɱɟɫɤɢɯ ɫɨɟɞɢɧɟɧɢɣ [136], ɨɞɧɚɤɨ ɨɧɢ ɧɟ ɨɛɥɚɞɚɸɬ ɧɟɨɛɯɨɞɢɦɵɦɢ ɞɥɹ ɢɫɩɨɥɶɡɨɜɚɧɢɹ ɜ ɬɟɯɧɢɤɟ ɫɜɨɣɫɬɜɚɦɢ – ɜɵɫɨɤɨɣ ɰɢɤɥɢɱɧɨɫɬɶɸ3 ɢ ɨɞɧɨɜɪɟɦɟɧɧɨ ɞɨɫɬɚɬɨɱɧɨ ɜɵɫɨɤɢɦ ɤɜɚɧɬɨɜɵɦ ɜɵɯɨɞɨɦ [137]. ɐɢɤɥɢɱɧɨɫɬɶ ɠɟ ɦɨɥɟɤɭɥ ȻɊ ɩɪɟɜɵɲɚɟɬ 106 [138], ɱɬɨ ɞɨɫɬɚɬɨɱɧɨ ɞɥɹ ɟɝɨ ɢɫɩɨɥɶɡɨɜɚɧɢɹ ɜ ɭɫɬɪɨɣɫɬɜɚɯ ɧɟ ɬɨɥɶɤɨ ɞɥɹ ɯɪɚɧɟɧɢɹ ɢɧɮɨɪɦɚɰɢɢ (ɧɚɩɪɢɦɟɪ, ɞɥɹ CD-RW ɧɟɨɛɯɨɞɢɦɚ ɰɢɤɥɢɱɧɨɫɬɶ 104 - 105), ɧɨ ɢ ɞɥɹ ɨɛɪɚɛɨɬɤɢ ɢɧɮɨɪɦɚɰɢɢ (ɧɚɩɪɢɦɟɪ, ɞɥɹ ɨɛɪɚɛɨɬɤɢ ɜɢɞɟɨ ɤɚɞɪɨɜ ɧɟɨɛɯɨɞɢɦɨ 106 ɰɢɤɥɨɜ ɡɚ ɞɟɧɶ ɪɚɛɨɬɵ). Ⱦɥɹ ɩɪɢɦɟɧɟɧɢɹ ȻɊ ɜ ɤɚɱɟɫɬɜɟ ɮɨɬɨɯɪɨɦɧɨɝɨ ɦɚɬɟɪɢɚɥɚ ɩɟɪɜɨɧɚɱɚɥɶɧɨ ɛɵɥɨ ɩɪɟɞɥɨɠɟɧɨ ɢɫɩɨɥɶɡɨɜɚɬɶ ɞɜɚ ɨɫɧɨɜɧɵɯ (ɨɛɥɚɞɚɸɳɢɯ ɧɚɢɛɨɥɶɲɟɣ ɷɧɟɪɝɟɬɢɱɟɫɤɨɣ ɡɚɫɟɥɟɧɧɨɫɬɶɸ) ɫɨɫɬɨɹɧɢɹ M ɢ B, ɩɨɫɤɨɥɶɤɭ ɨɧɢ ɨɛɥɚɞɚɸɬ ɡɧɚɱɢɬɟɥɶɧɨ

3

ɐɢɤɥɢɱɧɨɫɬɶɸ ɧɚɡɵɜɚɸɬ ɱɢɫɥɨ ɰɢɤɥɨɜ ɮɨɬɨɩɪɟɜɪɚɳɟɧɢɣ ɦɨɥɟɤɭɥ ɦɟɠɞɭ ɞɜɭɦɹ ɨɫɧɨɜɧɵɦɢ ɫɨɫɬɨɹɧɢɹɦɢ, ɞɨ ɪɚɡɪɭɲɟɧɢɹ ɢɥɢ ɞɟɧɚɬɭɪɚɰɢɢ 37% ɨɛɪɚɡɰɚ [138]

38 ɪɚɡɥɢɱɚɸɳɢɦɢɫɹ ɩɨɥɨɠɟɧɢɹɦɢ ɦɚɤɫɢɦɭɦɨɜ ɩɨɝɥɨɳɟɧɢɹ, ɤɨɬɨɪɵɟ, ɤ ɬɨɦɭ ɠɟ, ɫɨɜɩɚɞɚɸɬ ɫ ɞɥɢɧɚɦɢ ɜɨɥɧ ɢɡɥɭɱɟɧɢɹ ɤɪɢɩɬɨɧɨɜɵɯ ɥɚɡɟɪɨɜ - 568 ɢ 412 ɧɦ. Ɉɞɧɚɤɨ ɫɢɫɬɟɦɵ ɧɚ ɢɯ ɨɫɧɨɜɟ ɧɟ ɩɨɞɯɨɞɹɬ ɞɥɹ ɞɥɢɬɟɥɶɧɨɝɨ ɯɪɚɧɟɧɢɹ ɢɧɮɨɪɦɚɰɢɢ, ɩɨɫɤɨɥɶɤɭ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬ ɢɦɟɟɬ ɞɨɫɬɚɬɨɱɧɨ ɤɨɪɨɬɤɨɟ ɜɪɟɦɹ ɠɢɡɧɢ. ɉɨ ɷɬɨɣ ɩɪɢɱɢɧɟ ɩɪɨɜɨɞɢɥɫɹ ɩɨɢɫɤ ɞɪɭɝɢɯ ɢɧɬɟɪɦɟɞɢɚɬɨɜ, ɨɛɥɚɞɚɸɳɢɯ ɛɨɥɟɟ ɞɥɢɬɟɥɶɧɵɦ ɜɪɟɦɟɧɟɦ ɠɢɡɧɢ. ɂɡɜɟɫɬɧɨ, ɱɬɨ ɩɪɚɤɬɢɱɟɫɤɢ ɜɫɟ ɢɧɬɟɪɦɟɞɢɚɬɵ ɮɨɬɨɰɢɤɥɚ ȻɊ ɮɨɬɨɚɤɬɢɜɧɵ, ɬɨ ɟɫɬɶ ɩɪɢ ɩɨɝɥɨɳɟɧɢɢ ɫɜɟɬɚ ɨɧɢ ɦɨɝɭɬ ɩɟɪɟɯɨɞɢɬɶ ɜ ɞɪɭɝɢɟ ɢɧɬɟɪɦɟɞɢɚɬɵ, ɜɯɨɞɹɳɢɟ ɜ ɩɪɢɪɨɞɧɵɣ ɮɨɬɨɰɢɤɥ ȻɊ ɢɥɢ ɨɛɪɚɡɨɜɵɜɚɬɶ ɫɨɜɟɪɲɟɧɧɨ ɢɧɵɟ ɮɨɪɦɵ. ɉɪɢ ɢɫɫɥɟɞɨɜɚɧɢɢ ȻɊ ɜ ɩɥɟɧɤɚɯ ɛɵɥɨ ɨɛɧɚɪɭɠɟɧɨ ɨɬɜɟɬɜɥɟɧɢɟ ɮɨɬɨɰɢɤɥɚ ɫ ɢɧɬɟɪɦɟɞɢɚɬɚ Ɉ [139]: ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɩɪɢ ɞɟɣɫɬɜɢɢ ɤɜɚɧɬɚ ɫɜɟɬɚ ɧɚ ȻɊ ɜ ɫɨɫɬɨɹɧɢɢ Ɉ (all-E-ɪɟɬɢɧɚɥɶ, Asp85 ɩɪɨɬɨɧɢɪɨɜɚɧ) ɩɪɨɢɫɯɨɞɢɬ 9ȿ-o9Z- ɢɡɨɦɟɪɢɡɚɰɢɹ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ, ɩɨɥɭɱɚɟɦɵɣ ɜ ɪɟɡɭɥɶɬɚɬɟ ɢɧɬɟɪɦɟɞɢɚɬ ɛɵɥ ɧɚɡɜɚɧ P. Ɉɞɧɚɤɨ, 9Z-ɪɟɬɢɧɚɥɶ ɧɟ ɫɬɚɛɢɥɟɧ ɜ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ, ɩɨɷɬɨɦɭ ɩɪɨɢɫɯɨɞɢɬ ɩɨɫɬɟɩɟɧɧɵɣ ɝɢɞɪɨɥɢɡ ɚɥɶɞɢɦɢɧɧɨɣ ɫɜɹɡɢ ɫ ɨɛɪɚɡɨɜɚɧɢɟɦ ɢɧɬɟɪɦɟɞɢɚɬɚ Q, ɜ ɤɨɬɨɪɨɦ 9Z-ɪɟɬɢɧɚɥɶ ɭɞɟɪɠɢɜɚɟɬɫɹ ɜ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ɧɟ ɨɛɪɚɡɭɹ ɤɨɜɚɥɟɧɬɧɵɯ ɫɜɹɡɟɣ ɫ ȻɊ (ɬɚɤɨɣ ɜɵɜɨɞ ɛɵɥ ɫɞɟɥɚɧ ɚɜɬɨɪɚɦɢ ɪɚɛɨɬɵ [139] ɧɚ ɨɫɧɨɜɟ ɨɩɪɟɞɟɥɟɧɢɹ ɢɡɨɦɟɪɧɨɝɨ ɫɨɫɬɚɜɚ ɩɪɟɩɚɪɚɬɨɜ, ɢɡɭɱɟɧɢɹ ɢɯ ɫɩɟɤɬɪɚɥɶɧɵɯ ɫɜɨɣɫɬɜ ɢ ɫɪɚɜɧɟɧɢɹ ɩɨɥɭɱɟɧɧɵɯ ɪɟɡɭɥɶɬɚɬɨɜ ɢ ɥɢɬɟɪɚɬɭɪɧɵɦɢ ɞɚɧɧɵɦɢ). ɂɧɬɟɪɦɟɞɢɚɬ Q ɢɦɟɟɬ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟɨɝɪɚɧɢɱɟɧɧɨɟ ɜɪɟɦɹ ɠɢɡɧɢ (ɞɨ ɧɟɫɤɨɥɶɤɢɯ ɥɟɬ) ɢ ɩɟɪɟɯɨɞɢɬ ɜ ɨɫɧɨɜɧɨɟ ɫɨɫɬɨɹɧɢɟ ɬɚɤ ɠɟ ɩɨɞ ɞɟɣɫɬɜɢɟɦ ɫɜɟɬɚ ɨɩɪɟɞɟɥɟɧɧɨɣ ɞɥɢɧɧɵ ɜɨɥɧɵ, ɤɨɬɨɪɵɣ ɜɵɡɵɜɚɟɬ all-E-ɢɡɨɦɟɪɢɡɚɰɢɸ ɪɟɬɢɧɚɥɹ, ɩɨɫɥɟ ɱɟɝɨ ɩɪɨɢɫɯɨɞɢɬ ɫɚɦɨɩɪɨɢɡɜɨɥɶɧɚɹ ɪɟɝɟɧɟɪɚɰɢɹ ȻɊ [139].

Q380

hQ

ȻɊ568 B-ɮɨɪɦɚ

P490

hQ

O640

hQ

hQ

M-ɮɨɪɦɚ

M412 Ɋɢɫ. 8. ɂɧɬɟɪɦɟɞɢɚɬɵ ɮɨɬɨɰɢɤɥɚ ȻɊ, ɤɨɬɨɪɵɟ ɢɫɩɨɥɶɡɭɸɬɫɹ ɞɥɹ ɨɩɬɢɱɟɫɤɨɣ ɡɚɩɢɫɢ, ɯɪɚɧɟɧɢɹ ɢ ɫɱɢɬɵɜɚɧɢɹ ɢɧɮɨɪɦɚɰɢɢ [49].

39

5.2. ɋɨɡɞɚɧɢɟ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɯ ɢɡɨɛɪɚɠɟɧɢɣ ɧɚ ɩɥɟɧɤɚɯ, ɫɨɞɟɪɠɚɳɢɯ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ. ȼ 1980-ɯ ɝɨɞɚɯ ɜɩɟɪɜɵɟ ɛɵɥɚ ɩɨɤɚɡɚɧɚ ɜɨɡɦɨɠɧɨɫɬɶ ɢɫɩɨɥɶɡɨɜɚɧɢɹ ȻɊ ɞɥɹ ɫɨɡɞɚɧɢɹ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɯ ɢɡɨɛɪɚɠɟɧɢɣ [140]. Ɂɚɩɢɫɶ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɯ ɢɡɨɛɪɚɠɟɧɢɣ ɧɚ ɩɥɟɧɤɚɯ ȻɊ ɨɫɧɨɜɚɧɚ ɧɚ ɢɧɬɟɪɮɟɪɟɧɰɢɢ ɞɜɭɯ ɤɨɝɟɪɟɧɬɧɵɯ ɥɚɡɟɪɧɵɯ ɥɭɱɟɣ, ɤɨɬɨɪɵɟ ɨɛɪɚɡɭɸɬ ɝɨɥɨɝɪɚɮɢɱɟɫɤɭɸ ɪɟɲɟɬɤɭ (ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɨɟ ɪɚɫɩɪɟɞɟɥɟɧɢɟ Ɇ/B ɢɧɬɟɪɦɟɞɢɚɬɨɜ ȻɊ), ɤɨɬɨɪɚɹ ɜɨɡɧɢɤɚɟɬ ɛɥɚɝɨɞɚɪɹ ɢɡɦɟɧɟɧɢɸ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɢ ɩɨɤɚɡɚɬɟɥɹ ɩɪɟɥɨɦɥɟɧɢɹ ɩɥɟɧɤɢ ȻɊ ɩɪɢ ɩɨɝɥɨɳɟɧɢɢ ɤɜɚɧɬɚ ɫɜɟɬɚ. ɉɟɪɜɨɧɚɱɚɥɶɧɨ ɛɵɥɢ ɪɚɡɪɚɛɨɬɚɧɵ ɝɨɥɨɝɪɚɦɦɵ ȼ-ɬɢɩɚ, ɡɚɩɢɫɶ ɤɨɬɨɪɵɯ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɞɥɢɧɵ ɜɨɥɧɵ, ɜɵɡɵɜɚɸɳɟɣ ɩɟɪɟɯɨɞ ɢɧɬɟɪɦɟɞɢɚɬɨɜ ȼoɆ, ɚ ɞɥɹ ɫɱɢɬɵɜɚɧɢɹ ɢɫɩɨɥɶɡɭɟɬɫɹ ɬɨɬ ɠɟ ɩɟɪɟɯɨɞ, ɧɨ ɫ ɩɪɢɦɟɧɟɧɢɟɦ ɥɭɱɚ ɦɟɧɶɲɟɣ ɢɧɬɟɧɫɢɜɧɨɫɬɢ. ɉɨɫɤɨɥɶɤɭ ɞɥɹ ɫɱɢɬɵɜɚɧɢɹ ɢɫɩɨɥɶɡɭɟɬɫɹ ɬɨɬ ɠɟ ɩɟɪɟɯɨɞ ɢɧɬɟɪɦɟɞɢɚɬɨɜ ȼoɆ, ɧɟɞɨɫɬɚɬɤɨɦ ɝɨɥɨɝɪɚɦɦ ɷɬɨɝɨ ɬɢɩɚ ɹɜɥɹɟɬɫɹ ɩɨɧɢɠɟɧɢɟ ɤɨɧɬɪɚɫɬɚ ɜ ɩɪɨɰɟɫɫɟ ɫɱɢɬɵɜɚɧɢɹ, ɱɬɨ ɜ ɤɨɧɟɱɧɨɦ ɫɱɟɬɟ ɩɪɢɜɨɞɢɬ ɤ ɪɚɡɪɭɲɟɧɢɸ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɣ ɡɚɩɢɫɢ [36]. Ⱦɥɹ ɩɪɟɨɞɨɥɟɧɢɹ ɷɬɨɣ ɩɪɨɛɥɟɦɵ ɛɵɥɢ ɩɪɟɞɥɨɠɟɧɵ ɝɨɥɨɝɪɚɦɦɵ Ɇ-ɬɢɩɚ, ɜ ɤɨɬɨɪɵɯ ɞɥɹ ɡɚɩɢɫɢ ɢɫɩɨɥɶɡɭɸɬ ɩɟɪɟɯɨɞ Ɇoȼ. Ⱦɥɹ ɫɨɡɞɚɧɢɹ ɬɚɤɨɣ ɝɨɥɨɝɪɚɦɦɵ ɬɪɟɛɭɟɬɫɹ ɩɪɟɞɜɚɪɢɬɟɥɶɧɨɟ ɩɨɜɵɲɟɧɢɟ ɡɚɫɟɥɟɧɧɨɫɬɢ ɭɪɨɜɧɹ Ɇ ɨɫɜɟɳɟɧɢɟɦ ɧɚɫɵɳɚɸɳɢɦ ɥɭɱɟɦ (520-670 ɧɦ) [141], ɚ ɫɱɢɬɵɜɚɧɢɟ ɜɨɡɦɨɠɧɨ ɩɪɢ 630 ɧɦ, ɝɞɟ ȻɊ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɩɨɝɥɨɳɚɟɬ ɢ ɧɟ ɩɪɨɢɫɯɨɞɢɬ ɪɚɡɪɭɲɟɧɢɹ ɡɚɩɢɫɢ [36]. Ʌɚɡɟɪ (ɱɬɟɧɢɟ) Ɂɟɪɤɚɥɨ

Ʌɚɡɟɪ (ɡɚɩɢɫɶ) Ɂɟɪɤɚɥɨ

Ⱦɟɥɢɬɟɥɶ ɥɭɱɚ

Ɉɛɴɟɤɬ Ɂɟɪɤɚɥɨ

ɩɥɟɧɤɚ ȻɊ

Ɋɢɫ. 9. ɋɯɟɦɚ ɡɚɩɢɫɢ ɢ ɱɬɟɧɢɹ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɣ ɢɧɮɨɪɦɚɰɢɢ. ȼɚɠɧɵɦɢ ɫɜɨɣɫɬɜɚɦɢ ɮɨɬɨɯɪɨɦɧɨɝɨ ɦɚɬɟɪɢɚɥɚ, ɢɫɩɨɥɶɡɭɟɦɨɝɨ ɞɥɹ ɫɨɡɞɚɧɢɹ ɝɨɥɨɝɪɚɦɦ ɹɜɥɹɟɬɫɹ ɞɨɫɬɚɬɨɱɧɨ ɜɵɫɨɤɚɹ ɨɩɬɢɱɟɫɤɚɹ ɩɥɨɬɧɨɫɬɶ ɢ ɞɢɮɪɚɤɰɢɨɧɧɚɹ ɷɮɮɟɤɬɢɜɧɨɫɬɶ. ɉɪɢ ɨɩɬɢɦɢɡɚɰɢɢ ɫɜɨɣɫɬɜ ɩɥɟɧɨɤ ɧɚ ɨɫɧɨɜɟ ȻɊ ɞɥɹ ɢɯ ɢɫɩɨɥɶɡɨɜɚɧɢɹ

40 ɩɪɢ ɡɚɩɢɫɢ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɯ ɢɡɨɛɪɚɠɟɧɢɣ Ɇ-ɬɢɩɚ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɤɥɸɱɟɜɭɸ ɪɨɥɶ ɜ ɪɟɝɭɥɢɪɨɜɚɧɢɢ ɩɨɤɚɡɚɬɟɥɹ ɩɪɟɥɨɦɥɟɧɢɹ ɢ ɩɨɝɥɨɳɟɧɢɹ ɩɥɟɧɤɢ ȻɊ ɢɝɪɚɟɬ ɜɪɟɦɹ ɠɢɡɧɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ [48]. Ɉɤɚɡɚɥɨɫɶ, ɱɬɨ ɝɟɧɟɬɢɱɟɫɤɢ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɣ ȻɊD96N ɨɛɥɚɞɚɟɬ ɥɭɱɲɢɦɢ ɯɚɪɚɤɬɟɪɢɫɬɢɤɚɦɢ, ɱɟɦ ɩɪɢɪɨɞɧɵɣ, ɩɨɫɤɨɥɶɤɭ ɨɧ ɨɛɥɚɞɚɟɬ ɛɨɥɟɟ ɞɨɥɝɢɦ ɜɪɟɦɟɧɟɦ ɠɢɡɧɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ. Ɍɚɤ ɠɟ ɫɜɨɣɫɬɜɚ ɩɥɟɧɤɢ ȻɊ ɭɥɭɱɲɚɸɬɫɹ ɩɪɢ ɭɜɟɥɢɱɟɧɢɢ ɟɟ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɩɪɢ 570 ɧɦ ɞɨ 5-6 ɢ ɩɪɢ ɩɨɧɢɠɟɧɢɢ ɜɥɚɠɧɨɫɬɢ ɢ ɬɟɦɩɟɪɚɬɭɪɵ, ɚ ɞɢɮɪɚɤɰɢɨɧɧɚɹ ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɩɥɟɧɤɢ ȻɊ ɡɚɜɢɫɢɬ ɨɬ ɚɛɫɨɥɸɬɧɨɣ ɢɧɬɟɧɫɢɜɧɨɫɬɢ ɨɛɥɭɱɟɧɢɹ ɢ ɫɨɨɬɧɨɲɟɧɢɹ ɧɚɫɵɳɚɸɳɟɝɨ ɢ ɡɚɩɢɫɵɜɚɸɳɟɝɨ ɥɭɱɟɣ [141]. ɋɯɟɦɚ ɭɫɬɪɨɣɫɬɜɚ ɞɥɹ ɫɨɡɞɚɧɢɹ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɝɨ ɢɡɨɛɪɚɠɟɧɢɹ ɩɪɢɜɟɞɟɧɚ ɧɚ ɪɢɫ. 9. ɉɪɢɧɰɢɩ ɟɝɨ ɪɚɛɨɬɵ ɡɚɤɥɸɱɚɟɬɫɹ ɜ ɫɥɟɞɭɸɳɟɦ: ɩɟɪɟɞ ɧɚɱɚɥɨɦ ɡɚɩɢɫɢ ɩɪɨɜɨɞɹɬ ɚɤɬɢɜɚɰɢɸ ɩɥɟɧɤɢ (ɩɨɜɵɲɚɸɬ ɡɚɫɟɥɟɧɧɨɫɬɶ Ɇ-ɫɨɫɬɨɹɧɢɹ ȻɊ) ɧɚɫɵɳɚɸɳɢɦ ɥɭɱɟɦ ɫ ɞɥɢɧɨɣ ɜɨɥɧɵ 568 ɧɦ. ɉɨɫɥɟ ɷɬɨɝɨ ɞɥɹ ɡɚɩɢɫɢ ɢɫɩɨɥɶɡɭɸɬ ɤɪɢɩɬɨɧɨɜɵɣ ɥɚɡɟɪ (413 ɧɦ), ɥɭɱ ɤɨɬɨɪɨɝɨ ɞɟɥɢɬɫɹ ɧɚ ɨɛɴɟɤɬɧɵɣ (ɩɪɨɯɨɞɹɳɢɣ ɱɟɪɟɡ ɨɛɴɟɤɬ) ɢ ɨɩɨɪɧɵɣ, ɤɨɬɨɪɵɟ ɩɟɪɟɫɟɤɚɸɬɫɹ ɧɚ ɩɥɟɧɤɟ ȻɊ ɢ ɨɛɪɚɡɭɸɬ ɝɨɥɨɝɪɚɦɦɭ. ȼ ɤɚɱɟɫɬɜɟ ɫɱɢɬɵɜɚɸɳɟɝɨ ɢɫɩɨɥɶɡɭɟɬɫɹ ɬɨɬ ɠɟ ɧɚɫɵɳɚɸɳɢɣ ɥɭɱ, ɧɨ ɩɪɢ ɩɨɧɢɠɟɧɧɨɣ ɢɧɬɟɧɫɢɜɧɨɫɬɢ. Ɋɚɡɪɚɛɨɬɚɧɧɚɹ ɦɟɬɨɞɢɤɚ ɡɚɩɢɫɢ ɝɨɥɨɝɪɚɦɦ ɧɚ ɩɥɟɧɤɟ ȻɊ ɛɵɥɚ ɡɚɬɟɦ ɢɫɩɨɥɶɡɨɜɚɧɚ ɩɪɢ ɫɨɡɞɚɧɢɢ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɯ ɢɧɬɟɪɮɟɪɨɦɟɬɪɨɜ.

5.3. Ƚɨɥɨɝɪɚɮɢɱɟɫɤɚɹ ɢɧɬɟɪɮɟɪɨɦɟɬɪɢɹ. Ƚɨɥɨɝɪɚɮɢɱɟɫɤɚɹ ɢɧɬɟɪɮɟɪɨɦɟɬɪɢɹ - ɫɨɜɪɟɦɟɧɧɚɹ ɦɟɬɨɞɢɤɚ ɚɧɚɥɢɡɚ ɞɟɮɨɪɦɚɰɢɢ ɨɛɴɟɤɬɨɜ ɜ ɦɚɫɲɬɚɛɟ ɨɬ 1/10 ɞɨ 1/100 ɞɥɢɧɵ ɜɨɥɧɵ ɩɭɬɟɦ ɨɛɪɚɛɨɬɤɢ ɫɥɨɠɧɨɝɨ ɨɩɬɢɱɟɫɤɨɝɨ ɫɢɝɧɚɥɚ. ɇɟɨɛɯɨɞɢɦɨɫɬɶ ɬɚɤɢɯ ɢɡɦɟɪɟɧɢɣ ɜɨɡɧɢɤɚɟɬ ɩɪɢ ɨɩɪɟɞɟɥɟɧɢɢ ɚɦɩɥɢɬɭɞ ɜɢɛɪɚɰɢɢ ɩɨɜɟɪɯɧɨɫɬɟɣ, ɦɢɤɪɨɫɦɟɳɟɧɢɣ ɢ ɤɪɚɣɧɟ ɦɚɥɵɯ ɫɤɨɪɨɫɬɟɣ ɞɜɢɠɟɧɢɹ [142]. ȼ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɜ ɞɚɧɧɵɯ ɰɟɥɹɯ ɢɫɩɨɥɶɡɭɸɬ ɧɟɨɛɪɚɬɢɦɵɟ ɦɚɬɟɪɢɚɥɵ (ɧɚɩɪɢɦɟɪ, ɝɚɥɨɝɟɧɢɞɵ ɫɟɪɟɛɪɚ), ɨɞɧɚɤɨ ɨɧɢ ɧɟ ɩɨɡɜɨɥɹɸɬ ɨɬɫɥɟɠɢɜɚɬɶ ɞɢɧɚɦɢɤɭ ɩɪɨɰɟɫɫɨɜ [36], ɱɬɨ ɹɜɥɹɟɬɫɹ ɨɛɳɢɦ ɧɟɞɨɫɬɚɬɤɨɦ ɜɫɟɯ ɧɟɨɛɪɚɬɢɦɵɯ ɫɪɟɞ. ɂɫɩɨɥɶɡɨɜɚɧɢɟ ɜ ɤɚɱɟɫɬɜɟ ɫɪɟɞɵ ɞɥɹ ɡɚɩɢɫɢ ɩɥɟɧɤɢ ȻɊ, ɹɜɥɹɸɳɟɝɨɫɹ ɩɨɥɧɨɫɬɶɸ ɨɛɪɚɬɢɦɵɦ ɦɚɬɟɪɢɚɥɨɦ, ɩɨɡɜɨɥɹɟɬ ɪɚɫɲɢɪɢɬɶ ɜɨɡɦɨɠɧɨɫɬɢ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɣ ɢɧɬɟɪɮɟɪɨɦɟɬɪɢɢ. ɉɪɢɧɰɢɩ ɞɟɣɫɬɜɢɹ ɭɫɬɪɨɣɫɬɜɚ ɞɥɹ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɣ ɢɧɬɟɪɮɟɪɨɦɟɬɪɢɢ ɦɨɠɧɨ ɨɩɢɫɚɬɶ ɫɥɟɞɭɸɳɢɦ ɨɛɪɚɡɨɦ (ɫɦ. ɪɢɫ. 10): ɥɭɱ ɥɚɡɟɪɚ ɪɚɡɞɟɥɹɟɬɫɹ ɞɟɥɢɬɟɥɟɦ ɥɭɱɚ ɧɚ ɨɛɴɟɤɬɧɵɣ ɢ ɨɩɨɪɧɵɣ. ɂɧɬɟɧɫɢɜɧɨɫɬɶ ɨɛɴɟɤɬɧɨɝɨ ɥɭɱɚ ɪɟɝɭɥɢɪɭɟɬɫɹ ɩɨɥɹɪɢɡɚɬɨɪɨɦ. Ɋɚɫɲɢɪɢɬɟɥɢ ɨɫɭɳɟɫɬɜɥɹɸɬ ɪɚɫɮɨɤɭɫɢɪɨɜɤɭ ɥɭɱɚ, ɱɬɨɛɵ ɨɫɜɟɬɢɬɶ ɧɟɨɛɯɨɞɢɦɭɸ ɩɥɨɳɚɞɶ. Ɉɬ ɨɛɴɟɤɬɚ, ɪɚɫɩɨɥɨɠɟɧɧɨɝɨ ɦɟɠɞɭ ɪɚɫɲɢɪɢɬɟɥɟɦ ɢ ɩɥɟɧɤɨɣ ȻɊ ɨɬɪɚɠɚɟɬɫɹ ɛɟɫɩɨɪɹɞɨɱɧɚɹ ɫɜɟɬɨɜɚɹ ɜɨɥɧɚ, ɨɫɜɟɳɚɸɳɚɹ ɩɥɟɧɤɭ ȻɊ. Ɉɩɨɪɧɵɣ ɥɭɱ ɪɚɫɲɢɪɹɟɬɫɹ ɢ

41 ɧɚɤɥɚɞɵɜɚɟɬɫɹ ɧɚ ɩɪɟɞɦɟɬɧɭɸ ɜɨɥɧɭ ɜ ɩɥɨɫɤɨɫɬɢ ɩɥɟɧɤɢ ȻɊ, ɮɨɪɦɢɪɭɹ ɝɨɥɨɝɪɚɦɦɭ ɨɬɪɚɠɚɬɟɥɶɧɨɝɨ ɬɢɩɚ, ɤɨɬɨɪɚɹ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɢɧɬɟɪɮɟɪɟɧɰɢɨɧɧɭɸ ɤɚɪɬɢɧɭ ɜ ɜɢɞɟ ɧɚɛɨɪɚ ɜɚɪɢɚɰɢɣ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɢ ɩɨɤɚɡɚɬɟɥɹ ɩɪɟɥɨɦɥɟɧɢɹ. ɋɢɧɯɪɨɧɢɡɚɰɢɹ ɥɭɱɟɣ ɩɪɨɢɫɯɨɞɢɬ ɩɨɫɪɟɞɫɬɜɨɦ ɲɬɨɪɨɤ. Ɂɚɬɟɦ ɨɛɴɟɤɬ ɞɟɮɨɪɦɢɪɭɟɬɫɹ ɢ ɩɪɨɰɟɞɭɪɚ ɩɨɜɬɨɪɹɟɬɫɹ. ɉɨɫɥɟ ɷɬɨɝɨ, ɩɨ ɤɪɚɣɧɟɣ ɦɟɪɟ, ɞɜɟ ɝɨɥɨɝɪɚɦɦɵ ɫɨɫɭɳɟɫɬɜɭɸɬ ɜ ȻɊ ɩɥɟɧɤɟ. Ɂɟɪɤɚɥɨ

ɉɥɟɧɤɚ ȻɊ

ɱ ɣ ɥɭ

CCD ɤɚɦɟɪɚ ɉɨɥɹɪɢɡɚɬɨɪ Ɉɛɴɟɤɬ

Ɉɛɴɟɤ ɬɧ

Nd-ɥɚɡɟɪ, 532 ɧɦ

ɪɧɵ Ɉɩɨ

ɵɣ ɥɭɱ

Ɋɚɫɲɢɪɢɬɟɥɶ ɥɭɱɚ ɒɬɨɪɤɚ Ⱦɟɥɢɬɟɥɶ ɥɭɱɚ

Ɋɢɫ. 10. ɋɯɟɦɚ ɩɪɢɛɨɪɚ ɞɥɹ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɣ ɢɧɬɟɪɮɟɪɨɦɟɬɪɢɢ. Ⱦɥɹ ɪɟɤɨɧɫɬɪɭɤɰɢɢ ɨɛɟɢɯ ɝɨɥɨɝɪɚɦɦ ɨɛɴɟɤɬɧɵɣ ɥɭɱ ɜɵɤɥɸɱɚɟɬɫɹ, ɚ ɨɩɨɪɧɵɣ ɥɭɱ ɨɫɜɟɳɚɟɬ ɝɨɥɨɝɪɚɦɦɭ. ȼɨɫɫɬɚɧɨɜɥɟɧɧɵɟ ɝɨɥɨɝɪɚɦɦɵ ɫɦɟɲɢɜɚɸɬɫɹ, ɢ ɜɨɡɧɢɤɚɸɳɟɟ ɜ ɪɟɡɭɥɶɬɚɬɟ ɪɚɫɩɪɟɞɟɥɟɧɢɟ ɢɧɬɟɧɫɢɜɧɨɫɬɢ ɤɨɧɬɪɨɥɢɪɭɟɬɫɹ CCD-ɤɚɦɟɪɨɣ. ɉɨɥɹɪɢɡɚɬɨɪ ɩɟɪɟɞ ɤɚɦɟɪɨɣ ɧɟɨɛɯɨɞɢɦ, ɱɬɨɛɵ ɩɨɞɚɜɢɬɶ ɛɟɫɩɨɪɹɞɨɱɧɵɣ ɫɜɟɬ. ɉɨɫɤɨɥɶɤɭ ɝɨɥɨɝɪɚɮɢɱɟɫɤɚɹ ɢɧɬɟɪɮɟɪɨɦɟɬɪɢɹ ɜɨɡɦɨɠɧɚ ɬɨɥɶɤɨ ɩɪɢ ɧɢɡɤɨɣ ɢɧɬɟɧɫɢɜɧɨɫɬɢ ɫɜɟɬɚ, ɧɚɢɛɨɥɟɟ ɜɚɠɧɨɣ ɯɚɪɚɤɬɟɪɢɫɬɢɤɨɣ ɢɫɩɨɥɶɡɭɟɦɵɯ ɦɚɬɟɪɢɚɥɨɜ ɹɜɥɹɟɬɫɹ ɫɜɟɬɨɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɶ. ɂɡɦɟɧɟɧɢɟ ɫɜɟɬɨɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɢ ɢ ɤɨɧɬɪɚɫɬɧɨɫɬɢ ɩɥɟɧɤɢ ȻɊ ɞɨɫɬɢɝɚɟɬɫɹ ɜɚɪɢɚɰɢɟɣ ɜɪɟɦɟɧɢ ɠɢɡɧɢ ɢɧɬɟɪɦɟɞɢɚɬɚ Ɇ: ɟɝɨ ɭɜɟɥɢɱɟɧɢɟ ɜɵɡɵɜɚɟɬ ɩɪɨɩɨɪɰɢɨɧɚɥɶɧɨɟ ɭɦɟɧɶɲɟɧɢɟ ɫɜɟɬɨɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɢ [143]. Ⱦɚɧɧɵɟ ɭɫɬɪɨɣɫɬɜɚ ɧɟ ɬɨɥɶɤɨ ɧɚɯɨɞɹɬɫɹ ɧɚ ɫɬɚɞɢɢ ɪɚɡɪɚɛɨɬɤɢ ɢ ɩɪɨɟɤɬɢɪɨɜɚɧɢɹ. Ɉɩɪɟɞɟɥɟɧɧɵɦ ɭɫɩɟɯɨɦ ɦɨɠɧɨ ɫɱɢɬɚɬɶ ɬɨ, ɱɬɨ ɜ 2001 ɝɨɞɭ ɮɢɪɦɚ Munich Innovative Biomaterials, GmbH (MIB) ɜɵɩɭɫɬɢɥɚ ɧɚ ɪɵɧɨɤ ɝɨɥɨɝɪɚɮɢɱɟɫɤɢɣ ɢɧɬɟɪɮɟɪɨɦɟɬɪ FringeMaker-plus ɧɚ ɨɫɧɨɜɟ ɩɥɟɧɤɢ ɦɭɬɚɧɬɧɨɝɨ ȻɊD96N (http://www.mib-biotech.de). Ɉɧ ɦɨɠɟɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧ ɞɥɹ ɚɧɚɥɢɡɚ ɫɬɪɭɤɬɭɪɵ ɩɨɜɟɪɯɧɨɫɬɟɣ ɢ ɦɢɤɪɨɜɢɛɪɚɰɢɣ ɢ ɨɛɥɚɞɚɟɬ ɫɥɟɞɭɸɳɢɦɢ ɯɚɪɚɤɬɟɪɢɫɬɢɤɚɦɢ: 1) ɜɵɫɨɤɚɹ ɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɶ ɢɡɦɟɪɟɧɢɣ (5 ɧɦ), 2) ɜɵɫɨɤɚɹ ɩɪɨɢɡɜɨɞɢɬɟɥɶɧɨɫɬɶ, 3) ɧɟ ɬɪɟɛɭɟɬ ɪɚɫɯɨɞɧɵɯ ɦɚɬɟɪɢɚɥɨɜ, 4) ɧɟɨɝɪɚɧɢɱɟɧɧɚɹ ɰɢɤɥɢɱɧɨɫɬɶ, 5) ɧɟɛɨɥɶɲɢɟ ɪɚɡɦɟɪɵ ɢ ɩɪɨɫɬɨɬɚ ɜ ɭɩɪɚɜɥɟɧɢɢ.

42 ɇɚ ɪɢɫ. 11 ɩɪɟɞɫɬɚɜɥɟɧɵ ɝɨɥɨɝɪɚɦɦɵ, ɩɨɥɭɱɟɧɧɵɟ ɫ ɤɟɪɚɦɢɱɟɫɤɢɯ ɩɨɜɟɪɯɧɨɫɬɟɣ, ɩɨ ɤɨɬɨɪɵɦ ɦɨɠɧɨ ɥɟɝɤɨ ɨɩɪɟɞɟɥɢɬɶ ɧɚɥɢɱɢɟ ɦɢɤɪɨɬɪɟɳɢɧɵ (ɪɢɫ. 11ȼ), ɤɨɬɨɪɭɸ ɧɟɜɨɡɦɨɠɧɨ ɨɛɧɚɪɭɠɢɬɶ ɧɟɜɨɨɪɭɠɟɧɧɵɦ ɝɥɚɡɨɦ. Ⱥ)

ȼ)

Ɋɢɫ. 11. Ƚɨɥɨɝɪɚɦɦɵ, ɩɨɥɭɱɟɧɧɵɟ ɫ ɤɟɪɚɦɢɱɟɫɤɢɯ ɩɨɜɟɪɯɧɨɫɬɟɣ: Ⱥ) ɪɨɜɧɚɹ ɩɨɜɟɪɯɧɨɫɬɶ, ȼ) ɩɨɜɟɪɯɧɨɫɬɶ ɫ ɦɢɤɪɨɬɪɟɳɢɧɨɣ.

5.4. Ɉɛɪɚɛɨɬɤɚ ɜɢɞɟɨɢɡɨɛɪɚɠɟɧɢɣ. Ɉɩɬɢɱɟɫɤɚɹ ɨɛɪɚɛɨɬɤɚ ɢɡɨɛɪɚɠɟɧɢɣ ɤɨɪɟɧɧɵɦ ɨɛɪɚɡɨɦ ɨɬɥɢɱɚɟɬɫɹ ɨɬ ɰɢɮɪɨɜɨɣ. ȼ ɩɪɢɦɟɪ ɦɨɠɧɨ ɩɪɢɜɟɫɬɢ ɜɵɱɢɬɚɧɢɟ ɢɡɨɛɪɚɠɟɧɢɣ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɭɫɬɪɨɣɫɬɜɚ ɧɚ ɨɫɧɨɜɟ ɩɥɟɧɤɢ ȻɊ, ɩɪɟɞɥɨɠɟɧɧɨɝɨ Gu et al. [144]. ɉɪɢɧɰɢɩ ɪɚɛɨɬɵ ɩɪɢɛɨɪɚ ɩɪɟɞɫɬɚɜɥɟɧ ɧɚ ɪɢɫ. 12. ɋɜɟɬ ɫ ɞɥɢɧɧɨɣ ɜɨɥɧɵ 410 ɧɦ, ɩɪɨɯɨɞɹɳɢɣ ɱɟɪɟɡ ɮɢɝɭɪɭ Ⱥ ɬɚɤ ɠɟ ɩɪɨɯɨɞɢɬ ɢ ɱɟɪɟɡ ɩɥɟɧɤɭ ȻɊ, ɞɨɫɬɢɝɚɹ ɞɟɬɟɤɬɨɪɚ ɋ. ɋɜɟɬ ɫ ɞɥɢɧɧɨɣ ɜɨɥɧɵ 570 ɧɦ, ɩɪɨɯɨɞɹɳɢɣ ɱɟɪɟɡ ɮɢɝɭɪɭ ȼ, ɩɨɝɥɨɳɚɟɬɫɹ ɜ ɩɥɟɧɤɟ ȻɊ ɢ ɢɧɢɰɢɢɪɭɟɬ ɮɨɬɨɰɢɤɥ, ɩɨɷɬɨɦɭ ɜ ɷɬɢɯ ɭɱɚɫɬɤɚɯ ɩɥɟɧɤɢ ɨɩɪɟɞɟɥɟɧɧɚɹ ɱɚɫɬɶ ɦɨɥɟɤɭɥ ȻɊ ɞɨɫɬɢɝɚɟɬ ɫɨɫɬɨɹɧɢɹ Ɇ (ɫ ɦɚɤɫɢɦɭɦɨɦ ɩɨɝɥɨɳɟɧɢɹ 412 ɧɦ). ȼ ɪɟɡɭɥɶɬɚɬɟ ɜ ɦɟɫɬɚɯ ɩɥɟɧɤɢ ȻɊ, ɡɚɫɜɟɱɟɧɧɵɯ ɫɜɟɬɨɦ 570 ɧɦ, ɩɪɨɢɫɯɨɞɢɬ ɩɨɝɥɨɳɟɧɢɟ ɥɭɱɚ 410 ɧɦ ɢ ɞɨ ɞɟɬɟɤɬɨɪɚ ɞɨɯɨɞɢɬ ɥɢɲɶ ɱɚɫɬɶ ɢɡɨɛɪɚɠɟɧɢɹ Ⱥ. ɂɫɩɨɥɶɡɨɜɚɧɢɟ ɩɥɟɧɤɢ ȻɊ ɜ ɤɚɱɟɫɬɜɟ ɫɪɟɞɵ, ɩɪɢ ɩɨɦɨɳɢ ɤɨɬɨɪɨɣ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɜɵɱɢɬɚɧɢɟ ɨɩɬɢɱɟɫɤɢɯ ɢɡɨɛɪɚɠɟɧɢɣ ɩɨɡɜɨɥɹɟɬ ɨɫɭɳɟɫɬɜɢɬɶ ɷɬɭ ɨɩɟɪɚɰɢɸ ɩɪɢ ɩɨɦɨɳɢ ɩɪɟɞɟɥɶɧɨ ɩɪɨɫɬɨɝɨ ɭɫɬɪɨɣɫɬɜɚ.

A

Ʌɢɧɡɚ 1

ɉɥɟɧɤɚ ȻɊ

C

410 ɧɦ ɮɢɥɶɬɪ 410 ɧɦ

B Ʌɢɧɡɚ 2

570 ɧɦ Ɋɢɫ. 12. ɋɯɟɦɚ ɭɫɬɪɨɣɫɬɜɚ ɞɥɹ ɜɵɱɢɬɚɧɢɹ ɢɡɨɛɪɚɠɟɧɢɣ.

43 ȼ ɞɚɧɧɨɦ ɭɫɬɪɨɣɫɬɜɟ ɩɪɢ ɢɫɩɨɥɶɡɨɜɚɧɢɢ ɩɨɥɹɪɢɡɨɜɚɧɧɨɝɨ ɫɜɟɬɚ ɩɪɨɢɫɯɨɞɢɬ ɭɥɭɱɲɟɧɢɟ ɭɪɨɜɧɹ ɜɵɱɢɬɚɧɢɹ ɢɡɨɛɪɚɠɟɧɢɣ [49], ɩɨɫɤɨɥɶɤɭ ɩɥɟɧɤɚ ɢɡ ɨɪɢɟɧɬɢɪɨɜɚɧɧɵɯ ɦɨɥɟɤɭɥ ȻɊ ɨɛɥɚɞɚɟɬ ɜɵɫɨɤɨɣ ɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɶɸ ɤ ɩɨɥɹɪɢɡɚɰɢɢ ɨɛɥɭɱɚɸɳɟɝɨ ɫɜɟɬɚ, ɱɬɨ ɫɜɹɡɚɧɨ ɫɨ ɫɬɪɨɝɨ ɨɪɢɟɧɬɢɪɨɜɚɧɧɵɦ ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɵɦ ɪɚɫɩɨɥɨɠɟɧɢɟɦ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɜ ɦɨɥɟɤɭɥɟ ȻɊ. ȿɫɥɢ ɜ ɨɩɢɫɚɧɧɨɣ ɜɵɲɟ ɫɯɟɦɟ ɢɫɩɨɥɶɡɨɜɚɬɶ ɥɢɧɡɵ ɫ Ɏɭɪɶɟ-ɩɪɟɨɛɪɚɡɨɜɚɧɢɟɦ, ɷɬɨ ɩɨɡɜɨɥɹɟɬ ɡɧɚɱɢɬɟɥɶɧɨ ɪɚɫɲɢɪɢɬɶ ɜɨɡɦɨɠɧɨɫɬɢ ɩɪɢɦɟɧɟɧɢɹ ɞɚɧɧɨɝɨ ɭɫɬɪɨɣɫɬɜɚ. Ɉɫɨɛɟɧɧɨɫɬɹɦɢ Ɏɭɪɶɟ-ɩɪɟɨɛɪɚɡɨɜɚɧɢɹ ɨɩɬɢɱɟɫɤɨɝɨ ɢɡɨɛɪɚɠɟɧɢɹ ɹɜɥɹɟɬɫɹ [36] ɬɨ, ɱɬɨ ɮɨɪɦɚ Ɏɭɪɶɟ-ɫɩɟɤɬɪɚ ɨɛɴɟɤɬɚ ɧɟ ɡɚɜɢɫɢɬ ɨɬ ɤɨɨɪɞɢɧɚɬ ɟɝɨ ɪɚɫɩɨɥɨɠɟɧɢɹ (ɧɨ ɡɚɜɢɫɢɬ ɨɬ ɭɝɥɚ ɟɝɨ ɧɚɤɥɨɧɚ). əɪɤɨɫɬɶ ɨɛɴɟɤɬɚ ɨɩɪɟɞɟɥɹɟɬɫɹ ɰɟɧɬɪɚɥɶɧɨɣ ɬɨɱɤɨɣ (ɧɭɥɟɜɵɦ ɩɨɪɹɞɤɨɦ) Ɏɭɪɶɟ-ɫɩɟɤɬɪɚ, ɚ ɝɪɚɧɢɰɵ ɢɡɨɛɪɚɠɟɧɢɹ – ɨɫɬɚɥɶɧɵɦɢ ɟɝɨ ɬɨɱɤɚɦɢ (ɫ ɛɨɥɟɟ ɜɵɫɨɤɢɦ ɩɨɪɹɞɤɨɦ). Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɩɪɢ ɭɞɚɥɟɧɢɢ ɧɭɥɟɜɨɝɨ ɩɨɪɹɞɤɚ ɢɡ Ɏɭɪɶɟ-ɫɩɟɤɬɪɚ, ɢɡɨɛɪɚɠɟɧɢɟ ɧɚ ɜɵɯɨɞɟ ɫɨɯɪɚɧɹɟɬ ɫɜɨɢ ɝɪɚɧɢɰɵ, ɧɨ ɬɟɪɹɟɬ ɨɤɪɚɫɤɭ (ɫɦ. ɪɢɫ. 13), ɬɨ ɟɫɬɶ, ɷɬɚ ɩɪɨɫɬɚɹ ɨɩɟɪɚɰɢɹ ɩɨɡɜɨɥɹɟɬ ɜɵɩɨɥɧɹɬɶ ɪɚɫɩɨɡɧɚɜɚɧɢɟ ɝɪɚɧɢɰ ɢɡɨɛɪɚɠɟɧɢɹ.

Ɋɢɫ. 13. ɉɪɢɧɰɢɩ ɨɛɪɚɛɨɬɤɢ ɢɡɨɛɪɚɠɟɧɢɣ ɫ Ɏɭɪɶɟ-ɩɪɟɨɛɪɚɡɨɜɚɧɢɟɦ. ɇɚ ɷɬɨɦ ɠɟ ɩɪɢɧɰɢɩɟ ɨɫɧɨɜɚɧɵ ɭɫɬɪɨɣɫɬɜɚ ɞɥɹ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɝɨ ɪɚɫɩɨɡɧɚɜɚɧɢɹ ɨɛɪɚɡɨɜ, ɤɨɬɨɪɵɟ ɩɨɡɜɨɥɹɸɬ ɨɩɪɟɞɟɥɹɬɶ ɫɬɟɩɟɧɶ ɫɨɜɩɚɞɟɧɢɹ ɞɜɭɯ ɜɯɨɞɧɵɯ ɢɡɨɛɪɚɠɟɧɢɣ. Ⱦɥɹ ɩɨɹɫɧɟɧɢɹ ɦɨɠɧɨ ɩɪɢɜɟɫɬɢ ɫɥɟɞɭɸɳɢɣ ɩɪɢɦɟɪ: ɞɨɩɭɫɬɢɦ, ɬɪɟɛɭɟɬɫɹ ɧɚɣɬɢ ɫɩɟɰɢɮɢɱɟɫɤɨɟ ɫɥɨɜɨ ɜ ɬɟɤɫɬɟ. ȼ ɷɬɨɦ ɫɥɭɱɚɟ ɜɟɫɶ ɬɟɤɫɬ ɨɬɨɛɪɚɠɚɟɬɫɹ ɜ ɨɞɧɨɦ ɤɚɧɚɥɟ ɤɨɪɪɟɥɹɬɨɪɚ, ɚ ɬɪɟɛɭɟɦɨɟ ɫɥɨɜɨ – ɜ ɞɪɭɝɨɦ. Ɍɨɝɞɚ ɜ ɩɥɨɫɤɨɫɬɢ Ɏɭɪɶɟ-ɫɩɟɤɬɪɚ ɛɭɞɟɬ ɩɪɨɢɫɯɨɞɢɬɶ ɝɚɲɟɧɢɟ ɬɟɯ ɭɱɚɫɬɤɨɜ ɬɟɤɫɬɚ, ɤɨɬɨɪɵɟ ɫɨɜɩɚɞɚɸɬ ɫ ɜɜɟɞɟɧɧɵɦ ɫɥɨɜɨɦ. ȼ ɪɟɡɭɥɶɬɚɬɟ, ɧɚ ɜɵɯɨɞɟ ɜ ɢɡɨɛɪɚɠɟɧɢɢ ɬɟɤɫɬɚ ɩɪɨɩɚɞɟɬ ɢɫɤɨɦɨɟ ɫɥɨɜɨ, ɚ ɬɚɤɠɟ ɛɭɞɟɬ ɨɫɥɚɛɥɟɧ ɫɢɝɧɚɥ ɫɥɨɜ, ɤɨɬɨɪɵɟ ɱɚɫɬɢɱɧɨ ɫɨɜɩɚɞɚɸɬ ɫ ɢɫɤɨɦɵɦ (ɪɚɡɥɢɱɚɸɬɫɹ ɧɚ 1-2 ɛɭɤɜɵ). Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɞɚɧɧɨɟ ɭɫɬɪɨɣɫɬɜɨ ɧɟ ɬɨɥɶɤɨ ɭɤɚɡɵɜɚɟɬ ɦɟɫɬɨɧɚɯɨɠɞɟɧɢɟ

44 ɫɥɨɜɚ, ɧɨ ɬɚɤɠɟ ɞɚɟɬ ɦɟɪɭ ɫɬɟɩɟɧɢ ɩɨɞɨɛɢɹ. ȻɊ ɢɞɟɚɥɶɧɨ ɫɨɨɬɜɟɬɫɬɜɭɟɬ ɡɚɩɪɨɫɚɦ ɝɨɥɨɝɪɚɮɢɱɟɫɤɨɝɨ ɪɚɫɩɨɡɧɚɜɚɧɢɹ ɨɛɪɚɡɨɜ ɤ ɦɚɬɟɪɢɚɥɭ, ɢɫɩɨɥɶɡɭɟɦɨɦɭ ɞɥɹ ɡɚɩɢɫɢ. ɗɬɨɬ ɩɪɢɧɰɢɩ ɡɚɥɨɠɟɧ ɢ ɜ ɨɫɧɨɜɭ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ɭɫɬɪɨɣɫɬɜ ɚɫɫɨɰɢɚɬɢɜɧɨɣ ɩɚɦɹɬɢ. ȼ ɧɢɯ ɫɩɨɫɨɛ ɨɛɪɚɛɨɬɤɢ ɞɚɧɧɵɯ ɤɨɪɟɧɧɵɦ ɨɛɪɚɡɨɦ ɨɬɥɢɱɚɟɬɫɹ ɨɬ ɨɛɵɱɧɨ ɢɫɩɨɥɶɡɭɟɦɨɝɨ ɜ ɫɭɳɟɫɬɜɭɸɳɢɯ ɤɨɦɩɶɸɬɟɪɧɵɯ ɚɪɯɢɬɟɤɬɭɪɚɯ. Ⱦɚɧɧɨɟ ɭɫɬɪɨɣɫɬɜɨ ɛɟɪɟɬ ɮɪɚɝɦɟɧɬ ɜɯɨɞɧɵɯ ɞɚɧɧɵɯ (ɢɥɢ ɢɡɨɛɪɚɠɟɧɢɟ) ɢ "ɫɤɚɧɢɪɭɟɬ" ɫɪɚɡɭ ɜɫɸ ɩɚɦɹɬɶ ɧɚ ɧɚɥɢɱɢɟ ɛɥɨɤɚ ɞɚɧɧɵɯ, ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɝɨ ɜɯɨɞɧɨɦɭ, ɩɪɢ ɨɬɫɭɬɫɬɜɢɢ ɩɨɥɧɨɝɨ ɫɨɨɬɜɟɬɫɬɜɢɹ ɛɭɞɟɬ ɧɚɣɞɟɧ ɧɚɢɛɨɥɟɟ ɛɥɢɡɤɢɣ ɜɚɪɢɚɧɬ. Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɨɧɨ ɜɵɞɚɫɬ ɧɚ ɜɵɯɨɞɟ ɮɪɚɝɦɟɧɬ ɞɚɧɧɵɯ, ɭɞɨɜɥɟɬɜɨɪɹɸɳɢɣ ɤɪɢɬɟɪɢɸ ɫɨɨɬɜɟɬɫɬɜɢɹ. Ɍɚɤɨɣ ɩɨɞɯɨɞ ɤ ɨɛɪɚɛɨɬɤɟ ɢɧɮɨɪɦɚɰɢɢ ɩɨɡɜɨɥɹɟɬ ɪɟɡɤɨ ɩɨɜɵɫɢɬɶ ɩɪɨɢɡɜɨɞɢɬɟɥɶɧɨɫɬɶ ɭɫɬɪɨɣɫɬɜ. Ⱦɚɧɧɵɣ ɩɨɞɯɨɞ ɨɛɥɚɞɚɟɬ ɪɹɞɨɦ ɩɪɟɢɦɭɳɟɫɬɜ: ɤɪɨɦɟ ɬɨɝɨ, ɱɬɨ ɢɧɮɨɪɦɚɰɢɹ ɫɨɯɪɚɧɹɟɬɫɹ ɢ ɫɱɢɬɵɜɚɟɬɫɹ ɩɚɪɚɥɥɟɥɶɧɨ, ɦɨɠɧɨ ɞɨɫɬɢɱɶ ɨɱɟɧɶ ɜɵɫɨɤɨɣ ɫɤɨɪɨɫɬɢ ɩɟɪɟɞɚɱɢ ɞɚɧɧɵɯ. Ⱥ ɫɚɦɨɟ ɝɥɚɜɧɨɟ – ɜ ɬɚɤɢɯ ɭɫɬɪɨɣɫɬɜɚɯ ɩɪɚɤɬɢɱɟɫɤɢ ɨɬɫɭɬɫɬɜɭɸɬ ɦɟɯɚɧɢɱɟɫɤɢɟ ɤɨɦɩɨɧɟɧɬɵ, ɫɜɨɣɫɬɜɟɧɧɵɟ ɧɵɧɟɲɧɢɦ ɯɪɚɧɢɬɟɥɹɦ ɢɧɮɨɪɦɚɰɢɢ (ɧɚɩɪɢɦɟɪ, ɞɢɫɤɢ ɫ ɝɢɝɚɧɬɫɤɢɦ ɱɢɫɥɨɦ ɨɛɨɪɨɬɨɜ). ɗɬɨ ɝɚɪɚɧɬɢɪɭɟɬ ɧɟ ɬɨɥɶɤɨ ɦɝɧɨɜɟɧɧɵɣ ɞɨɫɬɭɩ ɤ ɞɚɧɧɵɦ, ɦɟɧɶɲɭɸ ɜɟɪɨɹɬɧɨɫɬɶ ɫɛɨɟɜ, ɧɨ ɢ ɛɨɥɟɟ ɧɢɡɤɨɟ ɩɨɬɪɟɛɥɟɧɢɟ ɷɥɟɤɬɪɨɷɧɟɪɝɢɢ, ɩɨɫɤɨɥɶɤɭ ɫɟɝɨɞɧɹ ɠɟɫɬɤɢɣ ɞɢɫɤ - ɨɞɢɧ ɢɡ ɧɚɢɛɨɥɟɟ ɷɧɟɪɝɨɟɦɤɢɯ ɤɨɦɩɨɧɟɧɬɨɜ ɤɨɦɩɶɸɬɟɪɚ. ɉɪɚɜɞɚ, ɜ ɫɜɹɡɢ ɫ ɬɟɦ, ɱɬɨ ɬɚɤɢɟ ɭɫɬɪɨɣɫɬɜɚ ɬɪɟɛɭɸɬ ɬɨɱɧɨɣ ɸɫɬɢɪɨɜɤɢ ɨɩɬɢɤɢ, ɧɚ ɧɚɫɬɨɹɳɟɦ ɭɪɨɜɧɟ ɪɚɡɪɚɛɨɬɤɢ ɷɬɢɯ ɩɪɢɛɨɪɨɜ, ɨɧɢ, ɜɟɪɨɹɬɧɨ, ɛɭɞɭɬ ɜɫɟ ɟɳɟ "ɛɨɹɬɶɫɹ" ɫɬɨɪɨɧɧɢɯ "ɦɟɯɚɧɢɱɟɫɤɢɯ ɜɨɡɞɟɣɫɬɜɢɣ". ɉɪɢɦɟɧɟɧɢɟ ɨɩɬɢɱɟɫɤɢɯ ɭɫɬɪɨɣɫɬɜ ɫ Ɏɭɪɶɟ-ɩɪɟɨɛɪɚɡɨɜɚɧɢɟɦ ɬɟɦ ɛɨɥɟɟ ɷɮɮɟɤɬɢɜɧɨ, ɱɬɨ ɢɡɜɟɫɬɧɨ, ɱɬɨ ɦɨɡɝ ɱɟɥɨɜɟɤɚ ɨɩɟɪɢɪɭɟɬ ɜ ɚɫɫɨɰɢɚɬɢɜɧɨɦ ɪɟɠɢɦɟ, ɩɨɷɬɨɦɭ ɨɩɬɢɱɟɫɤɚɹ ɚɫɫɨɰɢɚɬɢɜɧɚɹ ɩɚɦɹɬɶ ɧɚ ɨɫɧɨɜɟ ɩɪɟɨɛɪɚɡɨɜɚɧɢɹ Ɏɭɪɶɟ ɹɜɥɹɟɬɫɹ ɧɟɨɛɯɨɞɢɦɵɦ ɤɨɦɩɨɧɟɧɬɨɦ ɞɥɹ ɩɨɫɬɪɨɟɧɢɹ ɤɨɦɩɶɸɬɟɪɧɨɣ ɚɪɯɢɬɟɤɬɭɪɵ ɢɫɤɭɫɫɬɜɟɧɧɨɝɨ ɪɚɡɭɦɚ.

5.5. Ɇɨɥɟɤɭɥɹɪɧɚɹ ɩɚɦɹɬɶ ɧɚ ɨɫɧɨɜɟ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. Ʉɨɧɫɬɪɭɤɰɢɹ ɩɨɞɫɢɫɬɟɦɵ ɩɚɦɹɬɢ, ɨɫɧɨɜɚɧɧɨɣ ɧɚ ȻɊ, ɛɟɡɭɫɥɨɜɧɨ, ɢɦɟɟɬ ɨɩɪɟɞɟɥɟɧɧɵɟ ɩɪɟɢɦɭɳɟɫɬɜɚ ɩɟɪɟɞ ɬɪɚɞɢɰɢɨɧɧɨɣ ɩɨɥɭɩɪɨɜɨɞɧɢɤɨɜɨɣ ɩɚɦɹɬɶɸ. ȼɨɩɟɪɜɵɯ, ɨɧɚ ɨɫɧɨɜɚɧɚ ɧɚ ɛɟɥɤɟ, ɤɨɬɨɪɵɣ ɩɪɨɢɡɜɨɞɢɬɫɹ ɜ ɛɨɥɶɲɨɦ ɤɨɥɢɱɟɫɬɜɟ ɢ ɩɨ ɧɟɞɨɪɨɝɨɣ ɰɟɧɟ, ɱɟɦɭ ɫɩɨɫɨɛɫɬɜɭɸɬ ɞɨɫɬɢɠɟɧɢɹ ɛɢɨɬɟɯɧɨɥɨɝɢɢ ɢ ɝɟɧɟɬɢɱɟɫɤɨɣ ɢɧɠɟɧɟɪɢɢ. ȼɨ-ɜɬɨɪɵɯ, ɫɢɫɬɟɦɚ ɦɨɠɟɬ ɮɭɧɤɰɢɨɧɢɪɨɜɚɬɶ ɜ ɛɨɥɟɟ ɲɢɪɨɤɨɦ ɞɢɚɩɚɡɨɧɟ ɬɟɦɩɟɪɚɬɭɪ, ɱɟɦ ɩɨɥɭɩɪɨɜɨɞɧɢɤɨɜɚɹ ɩɚɦɹɬɶ. ȼ-ɬɪɟɬɶɢɯ, ɞɚɧɧɵɟ ɫɨɯɪɚɧɹɸɬɫɹ ɩɨɫɬɨɹɧɧɨ - ɞɚɠɟ ɟɫɥɢ ɜɵɤɥɸɱɢɬɶ ɩɢɬɚɧɢɟ ɫɢɫɬɟɦɵ ɩɚɦɹɬɢ, ɷɬɨ ɧɟ ɩɪɢɜɟɞɟɬ ɤ ɩɨɬɟɪɟ ɢɧɮɨɪɦɚɰɢɢ.

45 ȼ-ɱɟɬɜɟɪɬɵɯ ɨɩɬɢɱɟɫɤɢɣ ɧɨɫɢɬɟɥɶ ɢɧɮɨɪɦɚɰɢɢ ɩɨɡɜɨɥɹɟɬ ɨɫɭɳɟɫɬɜɥɹɬɶ ɡɚɩɢɫɶ ɩɨ ɜɫɟɦɭ ɨɛɴɟɦɭ, ɚ ɧɟ ɬɨɥɶɤɨ ɩɨ ɩɨɜɟɪɯɧɨɫɬɢ (ɤɚɤ ɷɬɨ ɪɟɚɥɢɡɭɟɬɫɹ ɧɚ ɦɚɝɧɢɬɧɵɯ ɧɨɫɢɬɟɥɹɯ), ɱɬɨ ɡɧɚɱɢɬɟɥɶɧɨ ɭɜɟɥɢɱɢɜɚɟɬ ɟɝɨ ɟɦɤɨɫɬɶ. ɂ, ɧɚɤɨɧɟɰ, ɤɭɛɢɤɢ ɫ ɞɚɧɧɵɦɢ, ɢɦɟɸɳɢɟ ɦɚɥɟɧɶɤɢɟ ɪɚɡɦɟɪɵ, ɧɨ ɫɨɞɟɪɠɚɳɢɟ ɝɢɝɚɛɚɣɬɵ ɢɧɮɨɪɦɚɰɢɢ, ɦɨɠɧɨ ɩɨɦɟɳɚɬɶ ɜ ɚɪɯɢɜ ɞɥɹ ɯɪɚɧɟɧɢɹ ɤɨɩɢɣ, ɬɚɤ ɤɚɤ ɤɭɛɢɤɢ ɧɟ ɫɨɞɟɪɠɚɬ ɞɜɢɠɭɳɢɯɫɹ ɱɚɫɬɟɣ, ɷɬɨ ɭɞɨɛɧɟɟ, ɱɟɦ ɢɫɩɨɥɶɡɨɜɚɧɢɟ ɩɨɪɬɚɬɢɜɧɵɯ ɠɟɫɬɤɢɯ ɞɢɫɤɨɜ. Ƚɪɭɩɩɨɣ ɢɫɫɥɟɞɨɜɚɬɟɥɟɣ ɰɟɧɬɪɚ “W. M. Keck Center for Molecular Electronic” ɩɨɞ ɪɭɤɨɜɨɞɫɬɜɨɦ ɩɪɨɮɟɫɫɨɪɚ Birge R.R. [145] ɪɚɡɪɚɛɨɬɚɧ ɩɪɨɬɨɬɢɩ ɩɨɞɫɢɫɬɟɦɵ ɩɚɦɹɬɢ, ɢɫɩɨɥɶɡɭɸɳɟɣ ɞɥɹ ɡɚɩɨɦɢɧɚɧɢɹ ɰɢɮɪɨɜɵɟ ɛɢɬɵ ɦɨɥɟɤɭɥɵ ȻɊ. ȼ ɪɚɡɪɚɛɨɬɚɧɧɨɣ ɫɯɟɦɟ ɢɫɩɨɥɶɡɭɸɬɫɹ ȼ ɢ Q ɫɨɫɬɨɹɧɢɹ ȻɊ, ɜ ɤɨɬɨɪɨɣ ȼ ɫɨɨɬɜɟɬɫɬɜɭɟɬ ɥɨɝɢɱɟɫɤɨɦɭ ɡɧɚɱɟɧɢɸ ɛɢɬɚ “0”, ɚ Q - ɡɧɚɱɟɧɢɸ “1”. ɉɨɫɤɨɥɶɤɭ ɷɬɢ ɞɜɚ ɫɨɫɬɨɹɧɢɹ ɢɦɟɸɬ ɡɚɦɟɬɧɨ ɨɬɥɢɱɚɸɳɢɟɫɹ ɫɩɟɤɬɪɵ ɩɨɝɥɨɳɟɧɢɹ, ɥɟɝɤɨ ɨɩɪɟɞɟɥɢɬɶ ɬɟɤɭɳɟɟ ɫɨɫɬɨɹɧɢɟ ɦɨɥɟɤɭɥɵ. ɉɨ ɨɰɟɧɤɚɦ ɢɫɫɥɟɞɨɜɚɬɟɥɟɣ, ɞɚɧɧɵɟ, ɡɚɩɢɫɚɧɧɵɟ ɧɚ ɬɚɤɨɦ ɡɚɩɨɦɢɧɚɸɳɟɦ ɭɫɬɪɨɣɫɬɜɟ, ɞɨɥɠɧɵ ɫɨɯɪɚɧɹɬɶɫɹ ɧɟ ɦɟɧɟɟ ɩɹɬɢ ɥɟɬ. Ȼɵɥ ɩɨɫɬɪɨɟɧ ɩɪɨɬɨɬɢɩ ɫɢɫɬɟɦɵ ɩɚɦɹɬɢ, ɜ ɤɨɬɨɪɨɦ ȻɊ ɡɚɩɨɦɢɧɚɟɬ ɞɚɧɧɵɟ ɜ ɬɪɟɯɦɟɪɧɨɣ ɦɚɬɪɢɰɟ. Ɍɚɤɚɹ ɦɚɬɪɢɰɚ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɤɸɜɟɬɭ ɪɚɡɦɟɪɨɦ 1x1x2 ɞɸɣɦɚ, ɡɚɩɨɥɧɟɧɧɭɸ ɩɨɥɢɚɤɪɢɥɚɦɢɞɧɵɦɞɧɵɦ ɝɟɥɟɦ, ɜ ɤɨɬɨɪɵɣ ɩɨɦɟɳɟɧ ɛɟɥɨɤ. Ɇɨɥɟɤɭɥɵ ȻɊ ɮɢɤɫɢɪɭɸɬɫɹ ɜ ɩɪɨɫɬɪɚɧɫɬɜɟ ɩɪɢ ɩɨɥɢɦɟɪɢɡɚɰɢɢ ɝɟɥɹ. Ʉɸɜɟɬɭ ɨɤɪɭɠɚɸɬ ɛɚɬɚɪɟɹ ɥɚɡɟɪɨɜ ɢ ɞɟɬɟɤɬɨɪɧɚɹ ɦɚɬɪɢɰɚ, ɩɨɫɬɪɨɟɧɧɚɹ ɧɚ ɛɚɡɟ ɩɪɢɛɨɪɚ, ɢɫɩɨɥɶɡɭɸɳɟɝɨ ɩɪɢɧɰɢɩ ɡɚɪɹɞɨɜɨɣ ɢɧɠɟɤɰɢɢ (CID - Charge Injection Device), ɤɨɬɨɪɵɟ ɫɥɭɠɚɬ ɞɥɹ ɡɚɩɢɫɢ ɢ ɱɬɟɧɢɹ ɞɚɧɧɵɯ. Ɂɚɩɢɫɶ ɞɚɧɧɵɯ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɞɜɭɫɬɚɞɢɣɧɵɣ ɩɪɨɰɟɫɫ (ɫɦ. ɪɢɫ. 14): ɫɧɚɱɚɥɚ ɠɟɥɬɵɣ ɫɬɪɚɧɢɱɧɵɣ ɥɚɡɟɪ ɢɧɢɰɢɢɪɭɟɬ ɮɨɬɨɰɢɤɥ ɜ ɩɥɨɫɤɨɫɬɢ ɤɸɜɟɬɵ, ɱɟɪɟɡ 2 ɦɤɫ (ɤɨɝɞɚ ɮɨɬɨɰɢɤɥ ȻɊ ɞɨɯɨɞɢɬ ɞɨ ɢɧɬɟɪɦɟɞɢɚɬɚ Ɉ) ɱɚɫɬɶ ɢɡ ɧɢɯ ɩɟɪɟɜɨɞɢɬɫɹ ɜ Qɫɨɫɬɨɹɧɢɟ ɞɟɣɫɬɜɢɟɦ ɡɚɩɢɫɵɜɚɸɳɟɝɨ ɤɪɚɫɧɨɝɨ ɥɚɡɟɪɚ, ɧɚɩɪɚɜɥɟɧɧɨɝɨ ɩɨɞ ɭɝɥɨɦ 90q ɤ ɩɥɨɫɤɨɫɬɢ ɫɬɪɚɧɢɱɧɨɝɨ ɥɚɡɟɪɚ. Ʌɭɱ ɡɚɩɢɫɵɜɚɸɳɟɝɨ ɥɚɡɟɪɚ ɩɪɨɯɨɞɢɬ ɱɟɪɟɡ ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɵɣ ɦɨɞɭɥɹɬɨɪ ɫɜɟɬɚ (SLM), ɤɨɬɨɪɵɣ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɠɢɞɤɨɤɪɢɫɬɚɥɥɢɱɟɫɤɭɸ ɦɚɬɪɢɰɭ, ɫɨɡɞɚɸɳɭɸ ɦɚɫɤɭ ɢɡ ɞɜɨɢɱɧɵɯ ɞɚɧɧɵɯ ɧɚ ɩɭɬɢ ɥɭɱɚ. Ɇɨɥɟɤɭɥɵ ȻɊ ɜ ɦɟɫɬɚɯ ɩɟɪɟɫɟɱɟɧɢɹ ɠɟɥɬɨɝɨ ɢ ɤɪɚɫɧɨɝɨ ɥɚɡɟɪɨɜ ɩɟɪɟɣɞɭɬ ɜ Qɫɨɫɬɨɹɧɢɟ ɢ ɛɭɞɭɬ ɩɪɟɞɫɬɚɜɥɹɬɶ ɞɜɨɢɱɧɵɟ ɟɞɢɧɢɰɵ, ɨɫɬɚɜɲɚɹɫɹ ɱɚɫɬɶ ɫɬɪɚɧɢɰɵ ɜɨɡɜɪɚɬɢɬɫɹ ɜ ɩɟɪɜɨɧɚɱɚɥɶɧɨɟ ɫɨɫɬɨɹɧɢɟ ȼ ɢ ɛɭɞɟɬ ɩɪɟɞɫɬɚɜɥɹɬɶ ɫɨɛɨɣ ɞɜɨɢɱɧɵɟ ɧɭɥɢ. ɉɪɨɰɟɫɫ ɱɬɟɧɢɹ ɞɚɧɧɵɯ ɩɪɚɤɬɢɱɟɫɤɢ ɚɧɚɥɨɝɢɱɟɧ ɢɯ ɡɚɩɢɫɢ ɫ ɬɨɣ ɪɚɡɧɢɰɟɣ, ɱɬɨ ɢɫɩɨɥɶɡɭɟɬɫɹ ɤɪɚɫɧɵɣ ɥɚɡɟɪ ɧɢɡɤɨɣ ɢɧɬɟɧɫɢɜɧɨɫɬɢ, ɱɬɨɛɵ ɩɪɟɞɭɩɪɟɞɢɬɶ “ɩɟɪɟɩɪɵɝɢɜɚɧɢɟ” ɦɨɥɟɤɭɥ ȻɊ ɜ Q-ɫɨɫɬɨɹɧɢɟ. ȼ ɷɬɨɦ ɫɥɭɱɚɟ ɜ ɦɟɫɬɚɯ ɪɚɫɩɨɥɨɠɟɧɢɹ ɦɨɥɟɤɭɥ ȻɊ, ɧɚɯɨɞɹɳɢɯɫɹ ɜ ɫɨɫɬɨɹɧɢɢ Q, ɧɟ ɩɪɨɢɫɯɨɞɢɬ ɩɨɝɥɨɳɟɧɢɹ ɤɪɚɫɧɨɝɨ ɫɜɟɬɚ ɢ

46 ɨɧ ɞɨɯɨɞɢɬ ɞɨ ɞɟɬɟɤɬɨɪɚ. ɗɬɨ ɫɨɡɞɚɟɬ “ɲɚɯɦɚɬɧɵɣ” ɪɢɫɭɧɨɤ ɢɡ ɫɜɟɬɥɵɯ ɢ ɬɟɦɧɵɯ ɩɹɬɟɧ ɧɚ ɋCD-ɦɚɬɪɢɰɟ, ɤɨɬɨɪɚɹ ɢ ɫɱɢɬɵɜɚɟɬ ɫɬɪɚɧɢɰɭ ɰɢɮɪɨɜɨɣ ɢɧɮɨɪɦɚɰɢɢ. Ⱦɥɹ ɫɬɢɪɚɧɢɹ ɞɚɧɧɵɯ ɞɨɫɬɚɬɨɱɧɨ ɤɨɪɨɬɤɨɝɨ ɢɦɩɭɥɶɫɚ ɫɢɧɟɝɨ ɥɚɡɟɪɚ, ɤɨɬɨɪɵɣ ɜɨɡɜɪɚɳɚɟɬ ɦɨɥɟɤɭɥɵ ȻɊ ɢɡ Q-ɫɨɫɬɨɹɧɢɹ ɜ ɨɫɧɨɜɧɨɟ ɫɨɫɬɨɹɧɢɟ, ɩɪɢ ɷɬɨɦ ɫɢɧɢɣ ɫɜɟɬ ɧɟ ɨɛɹɡɚɬɟɥɶɧɨ ɞɨɥɠɟɧ ɢɞɬɢ ɨɬ ɥɚɡɟɪɚ: ɦɨɠɧɨ ɫɬɟɪɟɬɶ ɜɫɸ ɤɸɜɟɬɭ ɫ ɩɨɦɨɳɶɸ ɨɛɵɤɧɨɜɟɧɧɨɣ ɭɥɶɬɪɚɮɢɨɥɟɬɨɜɨɣ ɥɚɦɩɵ.

Ɋɢɫ. 14. Ɂɚɩɢɫɶ ɢ ɱɬɟɧɢɟ ɞɚɧɧɵɯ ɧɚ ɬɪɟɯɦɟɪɧɨɦ ɧɨɫɢɬɟɥɟ ɢɧɮɨɪɦɚɰɢɢ ɧɚ ɨɫɧɨɜɟ ȻɊ. Ⱥ) ɚɤɬɢɜɚɰɢɹ ȼ) ɡɚɩɢɫɶ ɢɧɮɨɪɦɚɰɢɢ ɋ) ɱɬɟɧɢɟ ɢɧɮɨɪɦɚɰɢɢ. ɂɧɬɟɪɦɟɞɢɚɬɵ ȼ ɢ Q (ɨɛɪɚɡɭɸɳɢɣɫɹ ɩɪɢ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɦ ɩɨɝɥɨɳɟɧɢɢ ɞɜɭɯ ɮɨɬɨɧɨɜ ɦɨɥɟɤɭɥɨɣ ȻɊ), ɩɨɡɜɨɥɹɸɬ ɩɪɨɜɨɞɢɬɶ ɩɚɪɚɥɥɟɥɶɧɭɸ ɨɛɪɚɛɨɬɤɭ ɞɚɧɧɵɯ [145], ɟɫɥɢ ɢɫɩɨɥɶɡɨɜɚɬɶ ɧɟɫɤɨɥɶɤɨ ɤɨɦɩɥɟɤɬɨɜ ɥɚɡɟɪɧɵɯ ɢɡɥɭɱɚɬɟɥɟɣ. Ⱦɥɹ ɨɛɟɫɩɟɱɟɧɢɹ ɰɟɥɨɫɬɧɨɫɬɢ ɞɚɧɧɵɯ ɩɪɢ ɜɵɛɨɪɨɱɧɨɦ ɫɬɢɪɚɧɢɢ ɫɬɪɚɧɢɰ ɩɪɢɦɟɧɹɟɬɫɹ ɤɷɲɢɪɨɜɚɧɢɟ ɧɟɫɤɨɥɶɤɢɯ ɫɦɟɠɧɵɯ ɫɬɪɚɧɢɰ. ɉɪɢ ɨɩɟɪɚɰɢɹɯ ɱɬɟɧɢɹ ɢ ɡɚɩɢɫɢ ɬɚɤɠɟ ɢɫɩɨɥɶɡɭɸɬɫɹ ɞɜɚ ɞɨɩɨɥɧɢɬɟɥɶɧɵɯ ɛɢɬɚ ɱɟɬɧɨɫɬɢ, ɱɬɨɛɵ ɡɚɳɢɬɢɬɶɫɹ ɨɬ ɨɲɢɛɨɤ. ɋɬɪɚɧɢɰɚ ɞɚɧɧɵɯ ɦɨɠɟɬ ɛɵɬɶ ɩɪɨɱɢɬɚɧɚ ɛɟɡ ɪɚɡɪɭɲɟɧɢɹ ɞɨ 5 000 ɪɚɡ. Ʉɚɠɞɚɹ ɫɬɪɚɧɢɰɚ ɨɬɫɥɟɠɢɜɚɟɬɫɹ ɫɱɟɬɱɢɤɨɦ, ɢ ɟɫɥɢ ɩɪɨɢɫɯɨɞɢɬ 1024 ɱɬɟɧɢɹ, ɬɨ ɫɬɪɚɧɢɰɚ ɪɟɝɟɧɟɪɢɪɭɟɬɫɹ ɫ ɩɨɦɨɳɶɸ ɧɨɜɨɣ ɨɩɟɪɚɰɢɢ ɡɚɩɢɫɢ. ɍɱɢɬɵɜɚɹ ɫɤɨɪɨɫɬɶ ɮɨɬɨɰɢɤɥɚ, ɫɭɦɦɚɪɧɨɟ ɜɪɟɦɹ ɞɥɹ ɜɵɩɨɥɧɟɧɢɹ ɨɩɟɪɚɰɢɢ ɱɬɟɧɢɹ ɢɥɢ ɡɚɩɢɫɢ ɫɨɫɬɚɜɥɹɟɬ ɨɤɨɥɨ 10 ɦɫ. Ɉɞɧɚɤɨ, ɩɨɫɤɨɥɶɤɭ ɞɚɧɧɨɟ ɭɫɬɪɨɣɫɬɜɨ ɨɫɭɳɟɫɬɜɥɹɟɬ ɩɚɪɚɥɥɟɥɶɧɵɣ ɞɨɫɬɭɩ ɜ ɰɢɤɥɟ ɱɬɟɧɢɹ-ɡɚɩɢɫɢ, ɦɨɠɧɨ ɪɚɫɫɱɢɬɵɜɚɬɶ ɧɚ ɫɤɨɪɨɫɬɶ ɞɨ 10 Mb/ɩɫ. ɉɪɟɞɩɨɥɚɝɚɟɬɫɹ, ɱɬɨ, ɟɫɥɢ ɨɛɴɟɞɢɧɢɬɶ ɩɨ ɜɨɫɟɦɶ ɡɚɩɨɦɢɧɚɸɳɢɯ ɛɢɬɨɜɵɯ ɹɱɟɟɤ ɜ ɛɚɣɬ ɫ ɩɚɪɚɥɥɟɥɶɧɵɦ ɞɨɫɬɭɩɨɦ, ɦɨɠɧɨ ɞɨɫɬɢɝɧɭɬɶ ɫɤɨɪɨɫɬɢ 80 Mb/ɩɫ.

47 ɉɪɟɞɥɨɠɟɧɧɚɹ ɫɢɫɬɟɦɚ ɩɨ ɛɵɫɬɪɨɞɟɣɫɬɜɢɸ ɛɥɢɡɤɚ ɤ ɩɨɥɭɩɪɨɜɨɞɧɢɤɨɜɨɣ ɩɚɦɹɬɢ, ɩɨɤɚ ɧɟ ɜɫɬɪɟɬɢɬɫɹ ɫɬɪɚɧɢɱɧɵɣ ɞɟɮɟɤɬ. ɉɪɢ ɨɛɧɚɪɭɠɟɧɢɢ ɬɚɤɨɝɨ ɞɟɮɟɤɬɚ ɧɟɨɛɯɨɞɢɦɨ ɩɟɪɟɧɚɩɪɚɜɢɬɶ ɥɭɱ ɞɥɹ ɞɨɫɬɭɩɚ ɤ ɬɚɤɢɦ ɫɬɪɚɧɢɰɚɦ ɫ ɞɪɭɝɨɣ ɫɬɨɪɨɧɵ. Ɍɟɨɪɟɬɢɱɟɫɤɢ ɤɸɜɟɬɚ ɦɨɠɟɬ ɜɦɟɫɬɢɬɶ 1 Ɍb ɞɚɧɧɵɯ, ɨɝɪɚɧɢɱɟɧɢɹ ɧɚ ɟɦɤɨɫɬɶ ɫɜɹɡɚɧɵ, ɜ ɨɫɧɨɜɧɨɦ, ɫ ɞɟɮɟɤɬɚɦɢ ɥɢɧɡɨɜɨɣ ɫɢɫɬɟɦɵ ɢ ɱɢɫɬɨɬɨɣ ȻɊ.

5.6. Ɇɨɥɟɤɭɥɹɪɧɵɟ ɮɨɬɨɪɟɰɟɩɬɨɪɵ ɧɚ ɨɫɧɨɜɟ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. ȼ ɨɫɧɨɜɭ ɞɚɧɧɵɯ ɭɫɬɪɨɣɫɬɜ ɡɚɥɨɠɟɧɚ ɫɩɨɫɨɛɧɨɫɬɶ ɦɨɥɟɤɭɥɵ ȻɊ ɩɟɪɟɧɨɫɢɬɶ ɩɪɨɬɨɧ ɜ ɨɬɜɟɬ ɧɚ ɩɨɝɥɨɳɟɧɢɟ ɫɜɟɬɚ ɢ ɫɨɡɞɚɜɚɬɶ ɮɨɬɨɬɨɤ. ɇɚɩɪɢɦɟɪ, ɭɫɬɪɨɣɫɬɜɨ, ɩɪɟɞɥɨɠɟɧɧɨɟ Min et al. [146], ɩɪɟɞɫɬɚɜɥɹɸɳɟɟ ɫɨɛɨɣ 9ɛɢɬɨɜɭɸ ɮɨɬɨɹɱɟɣɤɭ (ɫɦ. ɪɢɫ. 15Ⱥ), ɝɟɧɟɪɢɪɭɟɬ ɫɢɝɧɚɥɵ ɪɚɡɥɢɱɧɨɣ ɮɨɪɦɵ ɜ ɨɬɜɟɬ ɧɚ ɪɚɡɥɢɱɧɵɟ ɢɡɨɛɪɚɠɟɧɢɹ, ɱɬɨ ɩɨɡɜɨɥɹɟɬ ɫɤɥɚɞɵɜɚɬɶ ɢ ɜɵɱɢɬɚɬɶ ɢɡɨɛɪɚɠɟɧɢɹ ɢ ɩɟɪɟɜɨɞɢɬɶ ɩɨɥɭɱɟɧɧɵɟ ɪɟɡɭɥɶɬɚɬɵ ɜ ɷɥɟɤɬɪɨɧɧɵɣ ɜɢɞ. ɉɨɫɤɨɥɶɤɭ ɜ ɧɟɦ ɩɥɟɧɤɚ ȻɊ ɢɦɟɟɬ ɢɡɜɟɫɬɧɨɟ, ɧɨ ɧɟɪɚɜɧɨɦɟɪɧɨɟ ɫɨɞɟɪɠɚɧɢɟ ȻɊ ɩɨ ɫɜɨɟɣ ɩɨɜɟɪɯɧɨɫɬɢ, ɨɧɚ ɫɨɡɞɚɟɬ ɪɚɡɥɢɱɧɵɟ ɫɢɝɧɚɥɵ ɜ ɨɬɜɟɬ ɧɚ ɨɫɜɟɳɟɧɢɟ ɪɚɡɥɢɱɧɵɯ ɭɱɚɫɬɤɨɜ ɩɥɟɧɤɢ ɥɭɱɟɦ ɫɜɟɬɚ ɨɞɢɧɚɤɨɜɨɣ ɩɥɨɳɚɞɢ ɢ ɢɧɬɟɧɫɢɜɧɨɫɬɢ. Ⱥ.

ɋɜɟɬ

ɉɥɟɧɤɚ ɫɩɢɪɨɩɢɪɚɧɚ

ȼ.

ɗɥɟɤɬɪɨɞ ITO Ʉ ɭɦɧɨɠɢɬɟɥɸ ɬɨɤɚ

ɉɥɟɧɤɚ ȻɊ ɋɜɟɬ

ɋɬɟɤɥɹɧɧɚɹ ɩɥɚɫɬɢɧɚ ɗɥɟɤɬɪɨɞ ITO

Pt ɷɥɟɤɬɪɨɞ ɉɥɟɧɤɚ ɮɥɚɜɢɧɚ ɉɥɟɧɤɚ ȻɊ

Ɋɢɫ. 15. ɍɫɬɪɨɣɫɬɜɨ ɮɨɬɨɪɟɰɟɩɬɨɪɨɜ ɧɚ ɨɫɧɨɜɟ ȻɊ. Ⱦɥɹ ɭɥɭɱɲɟɧɢɹ ɩɚɪɚɦɟɬɪɨɜ ɮɨɬɨɹɱɟɟɤ ɢɫɩɨɥɶɡɭɸɬɫɹ ɪɚɡɥɢɱɧɵɟ ɤɨɦɛɢɧɚɰɢɢ ɩɥɟɧɨɤ. ɇɚɩɪɢɦɟɪ, ɩɥɟɧɤɚ ɢɡ ɫɩɢɪɨɩɢɪɚɧɚ ɩɨɡɜɨɥɹɟɬ ɭɥɭɱɲɢɬɶ ɫɨɨɬɧɨɲɟɧɢɟ ɫɢɝɧɚɥɲɭɦ [146, 147], ɚ ɤɨɦɛɢɧɚɰɢɹ ȻɊ ɢ ɮɥɚɜɢɧɚ (ɫɦ. ɪɢɫ. 15ȼ) ɪɚɫɲɢɪɹɟɬ ɞɢɚɩɚɡɨɧ ɚɤɬɢɜɧɨɫɬɢ (ɜɨɡɧɢɤɧɨɜɟɧɢɟ ɮɨɬɨɨɬɜɟɬɚ) ɧɚ ɜɫɸ ɜɢɞɢɦɭɸ ɨɛɥɚɫɬɶ ɫɩɟɤɬɪɚ 400-650 ɧɦ [148]. Ʉɪɨɦɟ ɬɨɝɨ, ɛɵɥɨ ɩɪɨɞɟɦɨɧɫɬɪɢɪɨɜɚɧɨ ɭɫɩɟɲɧɨɟ ɪɚɫɩɨɡɧɚɜɚɧɢɟ ɰɜɟɬɨɜ (ɤɪɚɫɧɵɣ, ɡɟɥɟɧɵɣ, ɫɢɧɢɣ, ɛɟɥɵɣ ɢ ɱɟɪɧɵɣ) ɫ ɩɨɦɨɳɶɸ ɮɨɬɨɪɟɰɟɩɬɨɪɚ ɧɚ ɨɫɧɨɜɟ ɤɨɦɩɥɟɤɫɧɨɣ ɩɥɟɧɤɢ ɢɡ ɮɥɚɜɢɧɚ ɢ ȻɊ [149] ɩɨɫɪɟɞɫɬɜɨɦ ɚɧɚɥɢɡɚ ɮɨɬɨɷɥɟɤɬɪɢɱɟɫɤɢɯ ɨɬɜɟɬɨɜ ɭɫɬɪɨɣɫɬɜɚ ɧɚ ɫɜɟɬ ɪɚɡɥɢɱɧɨɣ ɞɥɢɧɵ ɜɨɥɧɵ.

48 Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɨɩɬɢɱɟɫɤɢɟ ɦɚɬɟɪɢɚɥɵ ɧɚ ɨɫɧɨɜɟ ȻɊ ɦɨɝɭɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧɵ ɞɥɹ ɫɨɡɞɚɧɢɹ ɰɟɥɨɝɨ ɧɚɛɨɪɚ ɬɟɯɧɢɱɟɫɤɢɯ ɭɫɬɪɨɣɫɬɜ, ɧɨ ɩɪɢ ɷɬɨɦ ɜɨɡɧɢɤɚɟɬ ɧɟɨɛɯɨɞɢɦɨɫɬɶ «ɩɨɞɨɝɧɚɬɶ» ɮɨɬɨɯɪɨɦɧɵɟ ɫɜɨɣɫɬɜɚ ɛɟɥɤɚ ɤ ɬɪɟɛɨɜɚɧɢɹɦ ɷɬɢɯ ɭɫɬɪɨɣɫɬɜ. ɂɡɦɟɧɟɧɢɟ ɮɨɬɨɯɪɨɦɧɵɯ ɫɜɨɣɫɬɜ ȻɊ ɦɨɠɟɬ ɛɵɬɶ ɞɨɫɬɢɝɧɭɬɨ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɬɪɟɯ ɩɨɞɯɨɞɨɜ: ɩɪɢɦɟɧɟɧɢɹ ɪɚɡɥɢɱɧɵɯ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɯ ɜɨɡɞɟɣɫɬɜɢɣ, ɡɚɦɟɧɵ ɯɪɨɦɨɮɨɪɚ ɢ ɝɟɧɟɬɢɱɟɫɤɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɛɟɥɤɚ ɢɥɢ ɢɯ ɫɨɱɟɬɚɧɢɟɦ, ɤɨɬɨɪɵɟ ɛɵɥɢ ɪɚɫɫɦɨɬɪɟɧɵ ɜɵɲɟ, ɜ ɝɥɚɜɟ 4 ɧɚ ɫ. 19-25. ȼ ɩɨɫɥɟɞɧɢɟ ɧɟɫɤɨɥɶɤɨ ɥɟɬ ɩɨɹɜɢɥɢɫɶ ɩɟɪɜɵɟ ɪɚɛɨɬɵ ɩɨ ɢɫɫɥɟɞɨɜɚɧɢɸ ɫɜɨɣɫɬɜ ɝɟɧɟɬɢɱɟɫɤɢ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɯ ȺȻɊ (D96N ɢ D85N), ɫɨɞɟɪɠɚɳɢɯ ɜɢɞɨɢɡɦɟɧɟɧɧɵɣ ɨɫɬɚɬɨɤ ɪɟɬɢɧɚɥɹ (4-ɨɤɫɨɪɟɬɢɧɚɥɶ) [150, 151]. ȼ ɷɬɢɯ ɪɚɛɨɬɚɯ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɬɚɤɨɣ ɩɨɞɯɨɞ ɧɚɢɛɨɥɟɟ ɩɟɪɫɩɟɤɬɢɜɟɧ ɞɥɹ ɨɩɬɢɦɢɡɚɰɢɢ ɫɜɨɣɫɬɜ ȻɊ ɞɥɹ ɟɝɨ ɩɨɫɥɟɞɭɸɳɟɝɨ ɬɟɯɧɢɱɟɫɤɨɝɨ ɩɪɢɦɟɧɟɧɢɹ. ȼ ɫɜɹɡɢ ɫ ɷɬɢɦ ɨɞɧɨɣ ɢɡ ɰɟɥɟɣ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɵ ɛɵɥɨ ɩɨɥɭɱɟɧɢɟ ɢ ɢɫɫɥɟɞɨɜɚɧɢɟ ɪɹɞɚ ȺȻɊ ɤɚɤ ɫ ɩɪɢɪɨɞɧɨɣ, ɬɚɤ ɢ ɫ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɨɣ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ, ɫɨɞɟɪɠɚɳɢɯ ɪɚɡɥɢɱɧɵɟ ɚɧɚɥɨɝɢ ɪɟɬɢɧɚɥɹ. Ɋɟɡɭɥɶɬɚɬɵ ɩɪɟɞɫɬɚɜɥɟɧɵ ɢ ɨɛɫɭɠɞɟɧɵ ɜ ɬɟɨɪɟɬɢɱɟɫɤɨɣ ɱɚɫɬɢ.

49

ɌȿɈɊȿɌɂɑȿɋɄȺə ɑȺɋɌɖ ɂ ɈȻɋɍɀȾȿɇɂȿ ɊȿɁɍɅɖɌȺɌɈȼ ȼ ɥɢɬɟɪɚɬɭɪɧɨɦ ɨɛɡɨɪɟ ɭɠɟ ɨɬɦɟɱɚɥɨɫɶ, ɱɬɨ ɨɞɧɢɦ ɢɡ ɚɤɬɢɜɧɨ ɪɚɡɜɢɜɚɸɳɢɯɫɹ ɧɚɩɪɚɜɥɟɧɢɣ ɫɨɜɪɟɦɟɧɧɨɣ ɛɢɨɷɥɟɤɬɪɨɧɢɤɢ ɹɜɥɹɟɬɫɹ ɢɫɩɨɥɶɡɨɜɚɧɢɟ ɮɨɬɨɱɭɜɫɬɜɢɬɟɥɶɧɵɯ ɛɟɥɤɨɜ, ɜ ɱɚɫɬɧɨɫɬɢ ȻɊ, ɜ ɤɚɱɟɫɬɜɟ ɪɚɛɨɱɟɣ ɫɪɟɞɵ ɜ ɩɪɨɬɨɬɢɩɚɯ ɪɚɡɥɢɱɧɵɯ ɫɟɧɫɨɪɨɜ ɢ ɭɫɬɪɨɣɫɬɜ ɞɥɹ ɯɪɚɧɟɧɢɹ ɢ ɨɛɪɚɛɨɬɤɢ ɢɧɮɨɪɦɚɰɢɢ. Ɍɚɤɠɟ ɛɵɥɢ ɪɚɫɫɦɨɬɪɟɧɵ ɨɫɧɨɜɧɵɟ ɪɟɡɭɥɶɬɚɬɵ, ɩɨɥɭɱɟɧɧɵɟ ɜ ɯɨɞɟ ɩɪɨɜɨɞɢɦɵɯ ɛɨɥɟɟ 30 ɥɟɬ ɢɫɫɥɟɞɨɜɚɧɢɣ ɦɟɯɚɧɢɡɦɚ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ȻɊ. Ɂɚ ɷɬɨɬ ɩɟɪɢɨɞ ɛɵɥ ɩɪɟɞɥɨɠɟɧ ɪɹɞ ɩɨɞɯɨɞɨɜ ɤ ɢɡɦɟɧɟɧɢɸ ɟɝɨ ɫɜɨɣɫɬɜ, ɬɚɤɢɯ ɤɚɤ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɜɨɡɞɟɣɫɬɜɢɹ, ɡɚɦɟɧɚ ɯɪɨɦɨɮɨɪɚ ɧɚ ɫɢɧɬɟɬɢɱɟɫɤɢɟ ɚɧɚɥɨɝɢ ɢ ɬɨɱɟɱɧɵɟ ɡɚɦɟɧɵ ɚɦɢɧɨɤɢɫɥɨɬ, ɩɪɢɱɟɦ ɫɨɱɟɬɚɧɢɟ ɞɜɭɯ ɩɨɫɥɟɞɧɢɯ ɩɨɞɯɨɞɨɜ ɩɨɡɜɨɥɹɟɬ ɜɚɪɶɢɪɨɜɚɬɶ ɜ ɡɧɚɱɢɬɟɥɶɧɵɯ ɩɪɟɞɟɥɚɯ ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ɢ ɫɤɨɪɨɫɬɶ ɮɨɬɨɰɢɤɥɚ ȺȻɊ. ȼ ɩɨɹɜɢɜɲɢɯɫɹ ɜ ɩɨɫɥɟɞɧɢɟ ɝɨɞɵ ɩɢɨɧɟɪɫɤɢɯ ɪɚɛɨɬɚɯ ɩɨ ɢɫɫɥɟɞɨɜɚɧɢɸ ȺȻɊ, ɫɨɞɟɪɠɚɳɢɯ ɨɞɧɨɜɪɟɦɟɧɧɨ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɣ ɯɪɨɦɨɮɨɪ ɢ ɦɭɬɚɰɢɸ ɩɨ ɨɩɪɟɞɟɥɟɧɧɨɦɭ ɨɫɬɚɬɤɭ ɚɦɢɧɨɤɢɫɥɨɬɵ [143], ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɞɥɹ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɯ ɰɟɥɟɣ ɬɚɤɢɟ ɩɪɟɩɚɪɚɬɵ ɨɛɥɚɞɚɸɬ ɥɭɱɲɢɦɢ ɯɚɪɚɤɬɟɪɢɫɬɢɤɚɦɢ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɢɡɜɟɫɬɧɵɦɢ ɪɚɧɟɟ ɜɚɪɢɚɧɬɚɦɢ. ɇɚɫɬɨɹɳɚɹ ɪɚɛɨɬɚ, ɧɚɩɪɚɜɥɟɧɧɚɹ ɧɚ ɩɨɢɫɤ ɩɟɪɫɩɟɤɬɢɜɧɵɯ ɜ ɬɟɯɧɢɱɟɫɤɨɦ ɩɥɚɧɟ ɫɨɱɟɬɚɧɢɣ ɦɨɞɢɮɢɤɚɰɢɣ ɯɪɨɦɨɮɨɪ – ɛɟɥɨɤ, ɹɜɥɹɟɬɫɹ ɱɚɫɬɶɸ ɩɥɚɧɨɜɵɯ ɢɫɫɥɟɞɨɜɚɧɢɣ ɩɨ ɪɚɡɪɚɛɨɬɤɟ ɧɨɜɵɯ ɦɚɬɟɪɢɚɥɨɜ ɧɚ ɨɫɧɨɜɟ ȺȻɊ, ɩɪɨɜɨɞɢɦɵɯ ɧɚ ɤɚɮɟɞɪɟ ɛɢɨɬɟɯɧɨɥɨɝɢɢ ɆɂɌɏɌ ɢɦ. Ɇ.ȼ. Ʌɨɦɨɧɨɫɨɜɚ ɫɨɜɦɟɫɬɧɨ ɫ ɇɂɂ Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɨɣ ɛɢɨɥɨɝɢɢ ɢɦ. Ⱥ.ɇ. Ȼɟɥɨɡɟɪɫɤɨɝɨ ɆȽɍ, ɐɟɧɬɪɨɦ ɮɨɬɨɯɢɦɢɢ ɊȺɇ, ɂɧɫɬɢɬɭɬɨɦ ɛɢɨɯɢɦɢɱɟɫɤɨɣ ɮɢɡɢɤɢ ɢɦ. ɇ.Ɇ. ɗɦɚɧɭɷɥɹ ɊȺɇ, OAO ɐɇɂɌɂ “Ɍɟɯɧɨɦɚɲ” ɢ ɎȽɍɉ «Ƚɨɫɇɂɂɝɟɧɟɬɢɤɚ». Ɋɚɛɨɬɚ ɩɪɨɜɨɞɢɬɫɹ ɜ ɪɚɦɤɚɯ ɜɵɩɨɥɧɟɧɢɹ ɝɪɚɧɬɨɜ ɊɎɎɂ 01-03-32078 ɢ 02-03-06555 ɢ ɝɪɚɧɬɚ ɂɇɌȺɋ-01-0267. ɐɟɥɶɸ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɵ ɛɵɥɨ ɤɨɦɩɥɟɤɫɧɨɟ ɢɫɫɥɟɞɨɜɚɧɢɟ ɜɥɢɹɧɢɹ ɨɞɧɨɜɪɟɦɟɧɧɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɛɟɥɤɨɜɨɣ ɢ ɯɪɨɦɨɮɨɪɧɨɣ ɱɚɫɬɟɣ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ, ɚ ɢɦɟɧɧɨ ɧɚ ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɨɜ ɩɨɝɥɨɳɟɧɢɹ, ɜɪɟɦɹ ɪɟɝɟɧɟɪɚɰɢɢ ɚɩɨɛɟɥɤɨɜ ɫ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ, ɨɫɨɛɟɧɧɨɫɬɢ ɤɢɧɟɬɢɤɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɚ ɢ ɫɬɚɛɢɥɶɧɨɫɬɶ ɩɪɟɩɚɪɚɬɨɜ. ɋɬɪɭɤɬɭɪɚ ɩɪɨɜɟɞɟɧɧɨɝɨ ɢɫɫɥɟɞɨɜɚɧɢɹ ɩɪɢɜɟɞɟɧɚ ɧɚ ɫɯɟɦɟ 1.

50

Ⱦɥɹ ɪɟɚɥɢɡɚɰɢɢ ɩɨɫɬɚɜɥɟɧɧɨɣ ɡɚɞɚɱɢ ɛɵɥɨ ɧɟɨɛɯɨɞɢɦɨ ɪɟɲɢɬɶ ɪɹɞ ɩɪɨɦɟɠɭɬɨɱɧɵɯ ɡɚɞɚɱ: 1) ɫɢɧɬɟɡɢɪɨɜɚɬɶ ɧɚɛɨɪ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ; 2) ɨɩɬɢɦɢɡɢɪɨɜɚɬɶ ɭɫɥɨɜɢɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɜɵɛɪɚɧɧɵɯ ɲɬɚɦɦɨɜ H. salinarum (ET1001, JW5 ɢ D96N) ɞɥɹ ɩɨɥɭɱɟɧɢɹ ɦɚɤɫɢɦɚɥɶɧɨɝɨ ɜɵɯɨɞɚ ȻɊ; 3) ɧɚɪɚɛɨɬɚɬɶ ɩɪɟɩɚɪɚɬɵ ȻɊ ɲɬɚɦɦɨɜ ET1001, JW5 ɢ D96N ɜ ɞɨɫɬɚɬɨɱɧɨɦ ɞɥɹ ɞɚɥɶɧɟɣɲɢɯ ɢɫɫɥɟɞɨɜɚɧɢɣ ɤɨɥɢɱɟɫɬɜɚɯ; 4) ɩɨɥɭɱɢɬɶ ɩɪɟɩɚɪɚɬɵ ɪɹɞɚ ȺȻɊ; 5) ɨɰɟɧɢɬɶ ɢɯ ɤɚɱɟɫɬɜɨ ɢ 6) ɢɫɫɥɟɞɨɜɚɬɶ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɢ ɨɫɨɛɟɧɧɨɫɬɢ ɤɢɧɟɬɢɤɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɨɜ. ɋɯɟɦɚ 1. ɋɬɪɭɤɬɭɪɚ ɢɫɫɥɟɞɨɜɚɧɢɹ. Ʉɭɥɶɬɢɜɢɪɨɜɚɧɢɟ H. salinarum ɲɬɚɦɦɨɜ JW5 ET1001 D96N ɋɢɧɬɟɡ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ

ȼɵɞɟɥɟɧɢɟ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ ET1001 D96N ɉɨɥɭɱɟɧɢɟ ɚɩɨɦɟɦɛɪɚɧ ET1001 D96N

ȼɵɞɟɥɟɧɢɟ "ɛɟɥɵɯ" ɦɟɦɛɪɚɧ JW5

Ɋɟɤɨɧɫɬɪɭɤɰɢɹ ȺȻɊ ɜɡɚɢɦɨɞɟɣɫɬɜɢɟɦ ȻɈ ɫ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ ET1001

D96N

JW5

ɂɫɫɥɟɞɨɜɚɧɢɟ ɛɵɫɬɪɨɣ ɤɢɧɟɬɢɤɢ ɨɩɬɢɱɟɫɤɢɯ ɢɡɦɟɧɟɧɢɣ ȺȻɊ ET1001

D96N

ɂɡɭɱɟɧɢɟ ɫɬɚɛɢɥɶɧɨɫɬɢ ɢ ɤɨɧɬɪɨɥɶ ɫɨɫɬɚɜɚ ɩɪɟɩɚɪɚɬɨɜ ȺȻɊ

JW5

Ɉɩɪɟɞɟɥɟɧɢɟ ɩɚɪɚɦɟɬɪɨɜ ȺȻɊ, ɜɚɠɧɵɯ ɞɥɹ ɨɰɟɧɤɢ ɜɨɡɦɨɠɧɨɫɬɢ ɢɯ ɬɟɯɧɢɱɟɫɤɨɝɨ ɩɪɢɦɟɧɟɧɢɹ

1. ȼɵɛɨɪ ɨɛɴɟɤɬɨɜ ɢɫɫɥɟɞɨɜɚɧɢɹ Ⱦɥɹ ɢɫɫɥɟɞɨɜɚɧɢɹ ɜɥɢɹɧɢɹ ɡɚɦɟɧɵ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɧɚ ɫɜɨɣɫɬɜɚ ɦɭɬɚɧɬɧɨɝɨ ȻɊ ɛɵɥɨ ɧɟɨɛɯɨɞɢɦɨ ɜɵɛɪɚɬɶ ɪɹɞ ɬɚɤɢɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ, ɤɨɬɨɪɵɟ ɨɛɪɚɡɭɸɬ ɫ ȻɈ ɞɢɤɨɝɨ ɬɢɩɚ ɯɪɨɦɨɩɪɨɬɟɢɧɵ, ɨɛɥɚɞɚɸɳɢɟ ɯɚɪɚɤɬɟɪɧɵɦɢ ɪɚɡɥɢɱɢɹɦɢ ɜ ɦɚɤɫɢɦɭɦɚɯ ɩɨɝɥɨɳɟɧɢɹ ɢ ɤɢɧɟɬɢɱɟɫɤɢɯ ɨɫɨɛɟɧɧɨɫɬɹɯ ɮɨɬɨɰɢɤɥɚ.

51 Ɋɚɧɟɟ ɧɚ ɤɚɮɟɞɪɟ ɛɢɨɬɟɯɧɨɥɨɝɢɢ ɆɂɌɏɌ ɢɦ. Ɇ.ȼ. Ʌɨɦɨɧɨɫɨɜɚ ɛɵɥɢ ɢɡɭɱɟɧɵ ɫɜɨɣɫɬɜɚ ɧɚɛɨɪɚ ȺȻɊ, ɩɨɥɭɱɟɧɧɵɯ ɢɡ ɚɩɨɦɟɦɛɪɚɧ H. salinarum ɲɬɚɦɦɚ 353ɉ [78, 105, 152-157]. ɇɚ ɨɫɧɨɜɟ ɷɬɢɯ ɞɚɧɧɵɯ ɞɥɹ ɜɵɩɨɥɧɟɧɢɹ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɵ ɛɵɥ ɜɵɛɪɚɧ ɪɹɞ ɩɪɨɢɡɜɨɞɧɵɯ ɪɟɬɢɧɚɥɹ (1), ɭ ɤɨɬɨɪɵɯ ɦɨɞɢɮɢɰɢɪɨɜɚɧɨ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɟ ɤɨɥɶɰɨ (2)-(5). ɋɜɨɣɫɬɜɚ ɫɨɨɬɜɟɬɫɬɜɭɸɳɢɯ ɢɦ ɯɪɨɦɨɩɪɨɬɟɢɧɨɜ ɩɪɢɜɟɞɟɧɵ ɜ ɬɚɛɥɢɰɟ 5. ɉɨɫɤɨɥɶɤɭ ɡɚɦɟɧɚ ɯɪɨɦɨɮɨɪɚ ɜ ɩɪɢɪɨɞɧɨɦ ȻɊ – ɞɨɫɬɚɬɨɱɧɨ ɩɨɞɪɨɛɧɨ ɢɫɫɥɟɞɨɜɚɧɧɚɹ ɬɟɦɚ ɢ ɨɯɚɪɚɤɬɟɪɢɡɨɜɚɧɨ ɧɟɫɤɨɥɶɤɨ ɫɨɬɟɧ ɚɧɚɥɨɝɨɜ ȻɊ, ɷɬɨɬ ɜɵɛɨɪ ɛɵɥ ɨɫɭɳɟɫɬɜɥɟɧ ɧɚ ɨɫɧɨɜɟ ɥɢɬɟɪɚɬɭɪɧɵɯ ɞɚɧɧɵɯ. Ʉɪɚɬɤɨɟ ɢɡɥɨɠɟɧɢɟ ɪɟɡɭɥɶɬɚɬɨɜ, ɨɩɭɛɥɢɤɨɜɚɧɧɵɯ ɩɨ ɷɬɨɦɭ ɧɚɩɪɚɜɥɟɧɢɸ ɪɚɛɨɬ, ɩɪɢɜɟɞɟɧɨ ɜ ɩɭɧɤɬɟ 4.2 ɨɛɡɨɪɚ ɥɢɬɟɪɚɬɭɪɵ ɧɚ ɫ. 21. ɇɚ ɨɫɧɨɜɟ ɩɨɥɭɱɟɧɧɵɯ ɞɚɧɧɵɯ ɦɨɠɧɨ ɫɞɟɥɚɬɶ ɡɚɤɥɸɱɟɧɢɟ, ɱɬɨ ɧɚɢɛɨɥɶɲɟɟ ɜɥɢɹɧɢɟ ɧɚ ɩɨɥɨɠɟɧɢɟ Omax ȺȻɊ ɨɤɚɡɵɜɚɸɬ ɞɥɢɧɚ ɰɟɩɢ ɫɨɩɪɹɠɟɧɧɵɯ ɞɜɨɣɧɵɯ ɫɜɹɡɟɣ ɢ ɦɨɞɢɮɢɤɚɰɢɹ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ. ɉɪɢ ɜɵɛɨɪɟ ɢɫɩɨɥɶɡɨɜɚɧɧɵɯ ɜ ɧɚɫɬɨɹɳɟɦ ɢɫɫɥɟɞɨɜɚɧɢɢ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ (2)(5) ɭɱɢɬɵɜɚɥɢ, ɱɬɨ ɞɥɹ ɨɞɧɨɡɧɚɱɧɨɣ ɢɧɬɟɪɩɪɟɬɚɰɢɢ ɪɟɡɭɥɶɬɚɬɨɜ ɢɫɫɥɟɞɨɜɚɧɢɹ ɤɢɧɟɬɢɤɢ ɮɨɬɨɰɢɤɥɚ ȺȻɊ, ɠɟɥɚɬɟɥɶɧɨ ɨɬɨɛɪɚɬɶ ɬɟ ɢɡ ɧɢɯ, ɤɨɬɨɪɵɟ ɨɛɥɚɞɚɸɬ ɞɨɫɬɚɬɨɱɧɨ ɜɵɫɨɤɨɣ ɷɮɮɟɤɬɢɜɧɨɫɬɶɸ ɮɨɬɨɰɢɤɥɚ. ɗɬɨ ɬɚɤɠɟ ɩɨɡɜɨɥɢɥɨ ɜ ɞɚɥɶɧɟɣɲɟɦ ɢɫɩɨɥɶɡɨɜɚɬɶ ɩɨɥɭɱɟɧɧɵɟ ɩɪɟɩɚɪɚɬɵ ɞɥɹ ɫɨɡɞɚɧɢɹ ɧɚ ɢɯ ɨɫɧɨɜɟ ɨɩɬɢɱɟɫɤɢɯ ɫɪɟɞ ɢ ɨɰɟɧɤɢ ɜɨɡɦɨɠɧɨɫɬɢ ɢɯ ɬɟɯɧɨɥɨɝɢɱɟɫɤɨɝɨ ɩɪɢɦɟɧɟɧɢɹ (ɚɤɬ ɨɛ ɢɫɫɥɟɞɨɜɚɧɢɢ ɢ ɢɫɩɨɥɶɡɨɜɚɧɢɢ ɧɚɪɚɛɨɬɚɧɧɵɯ ɩɪɟɩɚɪɚɬɨɜ ȻɊ ɩɪɢɥɨɠɟɧ ɜ ɤɨɧɰɟ ɞɢɫɫɟɪɬɚɰɢɢ, ɧɚ ɫ. 117). Ɍɚɛɥɢɰɚ 5. ɋɜɨɣɫɬɜɚ ȺȻɊ, ɩɨɥɭɱɟɧɧɵɯ ɢɡ ɚɩɨɦɟɦɛɪɚɧ ɲɬɚɦɦɚ 353ɉ [105, 107, 108]. Omax, ɧɦ N. ɋɬɪɭɤɬɭɪɚ ɯɪɨɦɨɮɨɪɚ

CHO

(1)

CHO

(2)

Omax, ɧɦ

all-E- 13Z- LA DA

Omax Mɢɧɬ., ɧɦ

568 548 568 558 a410 603

-

603 593 a425

527 504 506

-

ɋɯɨɞɧɚ ɫ ɩɪɢɪɨɞɧɵɦ ȻɊ

Ɋɟɥɚɤɫɚɰɢɹ ɞɜɭɯɮɚɡɧɚ,

CHO

(3)

Ɉɫɨɛɟɧɧɨɫɬɢ ɤɢɧɟɬɢɤɢ ɪɟɥɚɤɫɚɰɢɢ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɚ

a410 ɢɦɟɟɬ ɫɢɥɶɧɨ ɡɚɦɟɞɥɟɧɧɭɸ ɱɚɫɬɶ

O CHO

(4)

512 504 508

-

a395

Ɂɚɦɟɞɥɟɧɧɚɹ

-

Ɏɨɬɨɰɢɤɥ ɨɬɫɭɬɫɬɜɭɟɬ

CHO

(5)

O O

-

-

450 450

52 ɏɪɨɦɨɩɪɨɬɟɢɧɵ, ɨɛɪɚɡɭɟɦɵɟ ɨɬɨɛɪɚɧɧɵɦɢ ɚɧɚɥɨɝɚɦɢ ɪɟɬɢɧɚɥɹ, ɨɬɥɢɱɚɸɬɫɹ ɨɬ ɩɪɢɪɨɞɧɨɝɨ ȻɊ ɛɚɬɨɯɪɨɦɧɵɦ (2) ɢɥɢ ɝɢɩɫɨɯɪɨɦɧɵɦ (3), (4), (5) ɫɞɜɢɝɨɦ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ. Ʉɢɧɟɬɢɤɚ ɪɟɥɚɤɫɚɰɢɢ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ 3,4-ɞɢɞɟɝɢɞɪɨ-ȻɊ ɫɯɨɞɧɚ ɫ ɩɪɢɪɨɞɧɵɦ ȻɊ. 4-Ɉɤɫɨ-ȻɊ ɢɦɟɟɬ ɯɚɪɚɤɬɟɪɧɭɸ ɮɨɪɦɭ ɤɪɢɜɨɣ ɪɟɥɚɤɫɚɰɢɢ Ɇɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ, ɤɨɝɞɚ ɩɪɢ ɨɬɫɭɬɫɬɜɢɢ ɧɚɱɚɥɶɧɨɝɨ ɡɚɦɟɞɥɟɧɢɹ ɤɢɧɟɬɢɤɢ ɩɪɢɫɭɬɫɬɜɭɟɬ ɫɢɥɶɧɨ ɡɚɦɟɞɥɟɧɧɚɹ ɮɚɡɚ. Ɏɟɧɢɥɶɧɵɣ ȺȻɊ ɢɦɟɟɬ ɡɚɦɟɞɥɟɧɧɵɣ ɮɨɬɨɰɢɤɥ, ɚ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨ-ȻɊ ɧɟ ɨɛɥɚɞɚɟɬ ɜɵɪɚɠɟɧɧɵɦ ɮɨɬɨɰɢɤɥɨɦ (ɞɥɹ ɷɬɨɝɨ ȺȻɊ ɞɚɧɧɵɟ ɧɟ ɛɵɥɢ ɪɚɧɟɟ ɨɩɭɛɥɢɤɨɜɚɧɵ). Ɉɬɛɨɪ ɲɬɚɦɦɨɜ H. salinarum, ɢɫɩɨɥɶɡɨɜɚɧɧɵɯ ɞɥɹ ɩɨɥɭɱɟɧɢɹ ȺȻɊ, ɨɫɭɳɟɫɬɜɥɟɧ ɧɚ ɨɫɧɨɜɟ ɫɥɟɞɭɸɳɢɯ ɬɪɟɛɨɜɚɧɢɣ, ɨɛɭɫɥɨɜɥɟɧɧɵɯ ɬɟɯɧɨɥɨɝɢɱɟɫɤɨɣ ɧɚɩɪɚɜɥɟɧɧɨɫɬɶɸ ɢɫɫɥɟɞɨɜɚɧɢɹ: 1) ɉɪɨɫɬɨɬɚ ɜɵɞɟɥɟɧɢɹ ɩɪɟɩɚɪɚɬɨɜ ȻɊ ɧɟ ɫɨɞɟɪɠɚɳɢɯ ɤɚɪɨɬɢɧɨɢɞɨɜ, ɩɨɫɤɨɥɶɤɭ ɢɯ ɩɪɢɦɟɫɢ ɦɟɲɚɸɬ ɨɩɪɟɞɟɥɟɧɢɸ ɫɩɟɤɬɪɚɥɶɧɵɯ ɯɚɪɚɤɬɟɪɢɫɬɢɤ ɩɪɟɩɚɪɚɬɨɜ ȻɊ. ɗɬɨ ɫɜɹɡɚɧɨ ɫ ɬɟɦ, ɱɬɨ ɩɨɥɨɫɵ ɩɨɝɥɨɳɟɧɢɹ ɤɚɪɨɬɢɧɨɢɞɨɜ ɧɚɯɨɞɹɬɫɹ ɜ ɢɧɬɟɪɜɚɥɟ 350-550 ɧɦ ɢ ɱɚɫɬɢɱɧɨ ɩɟɪɟɤɪɵɜɚɸɬɫɹ ɫ ȻɊ. ɉɨɞ ɞɟɣɫɬɜɢɟɦ ɫɜɟɬɚ ɩɪɨɢɫɯɨɞɢɬ ɧɟɥɢɧɟɣɧɨɟ ɢɡɦɟɧɟɧɢɟ ɩɨɥɨɠɟɧɢɹ ɦɚɤɫɢɦɭɦɨɜ ɩɨɝɥɨɳɟɧɢɹ ɤɚɪɨɬɢɧɨɢɞɨɜ, ɱɬɨ ɭɫɥɨɠɧɹɟɬ ɢɧɬɟɪɩɪɟɬɚɰɢɸ ɪɟɡɭɥɶɬɚɬɨɜ ɢɫɫɥɟɞɨɜɚɧɢɹ ɫɩɟɤɬɪɚɥɶɧɵɯ ɯɚɪɚɤɬɟɪɢɫɬɢɤ ȻɊ ɢ ɞɟɥɚɟɬ ɦɚɥɨɜɟɪɨɹɬɧɵɦ ɩɪɢɦɟɧɟɧɢɟ ɬɚɤɢɯ ɩɪɟɩɚɪɚɬɨɜ ɜ ɛɢɨɷɥɟɤɬɪɨɧɢɤɟ; 2) ȼɨɡɦɨɠɧɨɫɬɶɸ ɭɦɟɧɶɲɟɧɢɹ ɱɢɫɥɚ ɫɬɚɞɢɣ ɩɪɨɰɟɫɫɚ ɩɨɥɭɱɟɧɢɹ ȺȻɊ, ɩɨɫɤɨɥɶɤɭ ɩɪɟɞɩɨɥɚɝɚɥɨɫɶ ɩɨɥɭɱɚɬɶ ɩɪɟɩɚɪɚɬɢɜɧɵɟ ɤɨɥɢɱɟɫɬɜɚ ȺȻɊ (a100 ɦɝ ɢ ɛɨɥɟɟ); 3) Ɂɧɚɱɢɬɟɥɶɧɨ ɡɚɦɟɞɥɟɧɧɚɹ ɤɢɧɟɬɢɤɚ ɪɟɥɚɤɫɚɰɢɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɚ, ɩɨɫɤɨɥɶɤɭ ɪɚɧɟɟ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɬɚɤɢɟ ɜɚɪɢɚɧɬɵ ȻɊ ɞɥɹ ɛɢɨɷɥɟɤɬɪɨɧɢɤɢ ɨɛɥɚɞɚɸɬ ɪɹɞɨɦ ɩɪɟɢɦɭɳɟɫɬɜ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɩɪɢɪɨɞɧɵɦ ȻɊ. ȼ ɪɟɡɭɥɶɬɚɬɟ, ɫ ɭɱɟɬɨɦ ɩɪɢɜɟɞɟɧɧɵɯ ɜɵɲɟ ɨɝɪɚɧɢɱɟɧɢɣ, ɢɡ ɧɚɛɨɪɚ ɞɨɫɬɭɩɧɵɯ ɞɥɹ ɧɚɫ ɲɬɚɦɦɨɜ ɛɵɥɢ ɜɵɛɪɚɧɵ ɬɪɢ: ET1001, JW5 ɢ D96N. ɂɯ ɤɪɚɬɤɚɹ ɯɚɪɚɤɬɟɪɢɫɬɢɤɚ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɢɫɩɨɥɶɡɭɟɦɵɦ ɪɚɧɟɟ ɲɬɚɦɦɨɦ 353ɉ ɞɚɧɚ ɜ ɬɚɛɥ. 6. Ɍɚɛɥɢɰɚ 6. ɏɚɪɚɤɬɟɪɢɫɬɢɤɚ ɲɬɚɦɦɨɜ H. salinarum. Ȼɚɤɬɟɪɢɨɨɩɫɢɧ

Ɋɟɬɢɧɚɥɶ

Ʉɚɪɨɬɢɧɨɢɞɵ

353ɉ

++

+

++

ɋɤɨɪɨɫɬɶ ɪɟɥɚɤɫɚɰɢɢ Ɇɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɚ ȻɊ Ȼɵɫɬɪɚɹ

ET1001

++

+

r

Ȼɵɫɬɪɚɹ

JW5

+

–

–

ȻɊ ɧɟ ɫɢɧɬɟɡɢɪɭɟɬɫɹ

D96N ++ + Ɇɟɞɥɟɧɧɚɹ r «++» - ɩɨɜɵɲɟɧɧɨɟ ɫɨɞɟɪɠɚɧɢɟ, «+» - ɫɬɚɧɞɚɪɬɧɵɣ ɭɪɨɜɟɧɶ ɲɬɚɦɦɨɜ ɞɢɤɨɝɨ ɬɢɩɚ, «r» - ɩɨɧɢɠɟɧɧɨɟ ɫɨɞɟɪɠɚɧɢɟ, «–» - ɧɟ ɫɢɧɬɟɡɢɪɭɟɬɫɹ

53 ɒɬɚɦɦɵ H. salinarum ȿɌ1001 ɢ D96N ɯɚɪɚɤɬɟɪɢɡɭɸɬɫɹ ɡɧɚɱɢɬɟɥɶɧɨ ɫɧɢɠɟɧɧɵɦ ɫɨɞɟɪɠɚɧɢɟɦ ɤɚɪɨɬɢɧɨɢɞɨɜ, ɩɨɷɬɨɦɭ ɩɪɢ ɜɵɞɟɥɟɧɢɢ ɢɡ ɧɢɯ ɉɆ, ɩɨɥɭɱɚɟɦɵɟ ɩɨ ɫɬɚɧɞɚɪɬɧɨɣ ɦɟɬɨɞɢɤɟ ɩɪɟɩɚɪɚɬɵ ɫɨɞɟɪɠɚɬ ɥɢɲɶ ɫɥɟɞɨɜɵɟ ɤɨɥɢɱɟɫɬɜɚ ɤɚɪɨɬɢɧɨɢɞɨɜ. Ʉɪɨɦɟ ɬɨɝɨ, ɷɬɢ ɞɜɚ ɲɬɚɦɦɚ ɯɚɪɚɤɬɟɪɢɡɭɸɬɫɹ ɜɵɫɨɤɢɦ ɫɨɞɟɪɠɚɧɢɟɦ ȻɊ, ɱɬɨ ɩɪɟɞɩɨɱɬɢɬɟɥɶɧɨ ɞɥɹ ɩɨɥɭɱɟɧɢɹ ɩɪɟɩɚɪɚɬɢɜɧɵɯ ɤɨɥɢɱɟɫɬɜ ɛɟɥɤɚ. ɋɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ ɬɚɤɠɟ, ɱɬɨ ɤ ɧɚɫɬɨɹɳɟɦɭ ɜɪɟɦɟɧɢ ɩɨɤɚɡɚɧɚ ɩɟɪɫɩɟɤɬɢɜɧɨɫɬɶ ɩɪɢɦɟɧɟɧɢɹ ɝɟɧɟɬɢɱɟɫɤɢ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɨɝɨ ȻɊD96N ɞɥɹ ɫɨɡɞɚɧɢɹ ɪɹɞɚ ɭɫɬɪɨɣɫɬɜ ɧɚ ɟɝɨ ɨɫɧɨɜɟ, ɨ ɱɟɦ ɛɵɥɨ ɭɩɨɦɹɧɭɬɨ ɜ ɪɚɡɞɟɥɟ 5 ɥɢɬɟɪɚɬɭɪɧɨɝɨ ɨɛɡɨɪɚ. ɏɨɬɹ ɫɨɞɟɪɠɚɧɢɟ ȻɈ ɜ ɤɥɟɬɤɚɯ H. salinarum ɲɬɚɦɦɚ JW5 ɧɢɠɟ, ɱɟɦ ȻɊ ɜ ȿɌ1001, ɟɝɨ ɜɚɠɧɵɦ ɩɪɟɢɦɭɳɟɫɬɜɨɦ ɹɜɥɹɟɬɫɹ ɜɨɡɦɨɠɧɨɫɬɶ ɛɢɨɫɢɧɬɟɬɢɱɟɫɤɨɝɨ ɩɨɥɭɱɟɧɢɹ ȺȻɊ, ɧɟ ɩɪɢɦɟɧɹɹ ɩɪɨɰɟɞɭɪɵ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ ɉɆ. ɗɬɨ ɩɨɡɜɨɥɹɟɬ ɡɧɚɱɢɬɟɥɶɧɨ ɭɩɪɨɫɬɢɬɶ ɩɪɨɰɟɫɫ ɩɨɥɭɱɟɧɢɹ ȺȻɊ. Ʉɪɨɦɟ ɬɨɝɨ, ɜ ɞɚɧɧɨɦ ɫɥɭɱɚɟ ɬɚɤɠɟ ɫɭɳɟɫɬɜɭɟɬ ɜɨɡɦɨɠɧɨɫɬɶ ɪɟɤɨɧɫɬɪɭɤɰɢɢ ȺȻɊ in vivo ɞɨɛɚɜɥɟɧɢɟɦ ɫɬɚɛɢɥɶɧɵɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɤ ɪɚɫɬɭɳɟɣ ɤɭɥɶɬɭɪɟ ɛɚɤɬɟɪɢɣ [158]. Ɍɚɤɨɣ ɩɨɞɯɨɞ ɹɜɥɹɟɬɫɹ ɨɞɧɢɦ ɢɡ ɧɚɢɛɨɥɟɟ ɩɟɪɫɩɟɤɬɢɜɧɵɯ ɞɥɹ ɫɨɡɞɚɧɢɹ ɩɪɟɩɚɪɚɬɨɜ ȺȻɊ ɜ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɯ ɰɟɥɹɯ, ɩɪɢɱɟɦ ɩɪɨɜɟɞɟɧɧɵɟ ɜ ɈȺɈ ɐɇɂɌɂ «Ɍɟɯɧɨɦɚɲ» ɢɫɫɥɟɞɨɜɚɧɢɹ (ɫɦ. ɚɤɬ ɨɛ ɢɫɩɨɥɶɡɨɜɚɧɢɢ, ɫ. 117) ɩɨɤɚɡɚɥɢ, ɱɬɨ ɩɪɟɩɚɪɚɬɵ ȺȻɊ ɧɚ ɨɫɧɨɜɟ ȻɈ ɢɡ ɲɬɚɦɦɚ JW5 ɩɪɨɞɟɦɨɧɫɬɪɢɪɨɜɚɥɢ ɨɞɧɢ ɢɡ ɥɭɱɲɢɯ ɬɟɯɧɢɱɟɫɤɢɯ ɩɨɤɚɡɚɬɟɥɟɣ.

2. Ɉɤɢɫɥɢɬɟɥɶɧɚɹ ɦɨɞɢɮɢɤɚɰɢɹ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɪɟɬɢɧɨɢɞɨɜ ɋɢɧɬɟɡ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɫ ɨɤɢɫɥɟɧɧɵɦ ɤɨɥɶɰɨɦ – ɞɨɫɬɚɬɨɱɧɨ ɯɨɪɨɲɨ ɢɡɭɱɟɧɧɨɟ ɧɚɩɪɚɜɥɟɧɢɟ ɜ ɯɢɦɢɢ ɪɟɬɢɧɨɢɞɨɜ. ȼ ɥɢɬɟɪɚɬɭɪɧɵɯ ɢɫɬɨɱɧɢɤɚɯ ɛɵɥɨ ɩɪɟɞɥɨɠɟɧɨ ɞɜɚ ɩɨɞɯɨɞɚ ɤ ɩɨɥɭɱɟɧɢɸ ɬɚɤɢɯ ɦɨɞɢɮɢɤɚɰɢɣ: «ɩɨɥɧɵɣ» ɫɢɧɬɟɡ ɩɭɬɟɦ ɫɨɱɟɬɚɧɢɹ ɞɜɭɯ ɢɥɢ ɛɨɥɟɟ ɫɢɧɬɨɧɨɜ ɫ ɨɛɪɚɡɨɜɚɧɢɟɦ ɦɨɥɟɤɭɥɵ ɪɟɬɢɧɨɢɞɚ [159-167] ɢɥɢ ɦɨɞɢɮɢɤɚɰɢɹ ɰɢɤɥɢɱɟɫɤɨɣ ɱɚɫɬɢ ɭɠɟ ɝɨɬɨɜɨɝɨ ɋ20-ɭɝɥɟɪɨɞɧɨɝɨ ɫɤɟɥɟɬɚ ɩɨɞɯɨɞɹɳɟɝɨ ɩɪɟɞɲɟɫɬɜɟɧɧɢɤɚ [95, 104,168-173]. Ʉɚɤ ɩɪɚɜɢɥɨ, ɢɡɭɱɟɧɧɵɟ ɪɚɧɟɟ ɫɢɫɬɟɦɵ ɨɤɢɫɥɢɬɟɥɟɣ ɜ ɪɟɚɤɰɢɢ ɫ ɪɟɬɢɧɚɥɟɦ (1) ɨɬɥɢɱɚɥɢɫɶ ɨɬɫɭɬɫɬɜɢɟɦ ɜɵɪɚɠɟɧɧɨɣ ɪɟɝɢɨɫɟɥɟɤɬɢɜɧɨɫɬɢ ɢ ɜɵɡɵɜɚɥɢ ɢɡɨɦɟɪɢɡɚɰɢɸ ɞɜɨɣɧɵɯ ɫɜɹɡɟɣ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ, ɱɬɨ ɩɪɢɜɨɞɢɥɨ ɤ ɫɥɨɠɧɨɣ ɫɦɟɫɢ ɩɪɨɞɭɤɬɨɜ [170, 172174]. Ⱦɥɹ ɦɨɞɢɮɢɤɚɰɢɢ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɩɪɟɩɚɪɚɬɢɜɧɨɟ ɡɧɚɱɟɧɢɟ ɩɨɥɭɱɢɥɢ ɥɢɲɶ ɩɪɟɩɚɪɚɬɵ «ɚɤɬɢɜɧɨɝɨ» ɞɢɨɤɫɢɞɚ ɦɚɪɝɚɧɰɚ [95, 168-171], ɩɪɢɝɨɬɨɜɥɟɧɧɵɟ ɩɨ ɪɚɡɥɢɱɧɵɦ ɦɟɬɨɞɢɤɚɦ. Ɋɚɧɟɟ ɜ ɧɚɲɟɣ ɥɚɛɨɪɚɬɨɪɢɢ ɞɥɹ ɩɨɥɭɱɟɧɢɹ all-E- ɢ 13Z-ɢɡɨɦɟɪɨɜ 4-ɨɤɫɨɪɟɬɢɧɚɥɹ (3) ɢɫɩɨɥɶɡɨɜɚɥɢ ɦɟɬɨɞ ɚɥɥɢɥɶɧɨɝɨ ɨɤɢɫɥɟɧɢɹ ɋ4-ɦɟɬɢɥɟɧɨɜɨɣ ɝɪɭɩɩɵ ɤɨɥɶɰɚ

54 ɫɨɨɬɜɟɬɫɬɜɭɸɳɢɯ ɢɡɨɦɟɪɨɜ ɪɟɬɢɧɚɥɹ (1), ɨɫɭɳɟɫɬɜɥɹɟɦɵɣ ɞɟɣɫɬɜɢɟɦ «ɤɢɫɥɨɣ» ɮɨɪɦɵ MnO2 [95, 169]. ɇɟɫɦɨɬɪɹ ɧɚ ɲɢɪɨɤɨɟ ɜɚɪɶɢɪɨɜɚɧɢɟ ɭɫɥɨɜɢɣ (ɫɩɨɫɨɛ ɩɪɢɝɨɬɨɜɥɟɧɢɹ ɪɟɚɝɟɧɬɚ ɢ ɭɫɥɨɜɢɹ ɩɪɨɰɟɫɫɚ: ɬɟɦɩɟɪɚɬɭɪɚ, ɪɚɫɬɜɨɪɢɬɟɥɶ, ɫɨɨɬɧɨɲɟɧɢɟ ɨɤɢɫɥɢɬɟɥɶ – ɫɭɛɫɬɪɚɬ), ɤɨɧɜɟɪɫɢɹ ɪɟɬɢɧɚɥɹ (1) ɧɟ ɩɪɟɜɵɲɚɥɚ 30-40%. ɏɨɬɹ MnO2 ɧɟ ɜɵɡɵɜɚɟɬ ɡɚɦɟɬɧɨɣ ɢɡɨɦɟɪɢɡɚɰɢɢ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ, ɟɝɨ ɨɫɧɨɜɧɵɦ ɧɟɞɨɫɬɚɬɤɨɦ ɹɜɥɹɟɬɫɹ ɬɪɭɞɧɨɫɬɶ ɩɨɥɭɱɟɧɢɹ ɩɪɟɩɚɪɚɬɚ ɫɨ ɫɬɚɧɞɚɪɬɧɨɣ ɨɤɢɫɥɢɬɟɥɶɧɨɣ ɫɩɨɫɨɛɧɨɫɬɶɸ. ɗɬɨ ɩɪɢɜɨɞɢɬ ɤ ɧɟɨɛɯɨɞɢɦɨɫɬɢ ɞɨɩɨɥɧɢɬɟɥɶɧɵɯ ɷɤɫɩɟɪɢɦɟɧɬɨɜ ɩɨ ɬɟɫɬɢɪɨɜɚɧɢɸ ɤɚɠɞɨɣ ɧɨɜɨɣ ɩɚɪɬɢɢ ɨɤɢɫɥɢɬɟɥɹ [95, 169]. Ɉɞɧɨɣ ɢɡ ɰɟɥɟɣ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɵ ɛɵɥ ɩɨɢɫɤ ɦɟɬɨɞɚ ɨɤɢɫɥɟɧɢɹ, ɩɨɡɜɨɥɹɸɳɟɝɨ ɫɢɧɬɟɡɢɪɨɜɚɬɶ ɫ ɞɨɫɬɚɬɨɱɧɨ ɜɵɫɨɤɢɦɢ ɜɵɯɨɞɚɦɢ ɧɚɛɨɪ ɩɪɨɢɡɜɨɞɧɵɯ ɪɟɬɢɧɚɥɹ (1) ɫ ɨɤɢɫɥɟɧɧɵɦ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɵɦ ɤɨɥɶɰɨɦ ɞɥɹ ɩɨɫɥɟɞɭɸɳɟɝɨ ɩɨɥɭɱɟɧɢɹ ɚɧɚɥɨɝɨɜ ȻɊ. Ȼɵɥɨ ɩɪɨɜɟɞɟɧɨ ɢɫɫɥɟɞɨɜɚɧɢɟ ɫɥɟɞɭɸɳɟɝɨ ɪɹɞɚ ɨɤɢɫɥɢɬɟɥɟɣ: ɚ) ɯɥɨɪɨɯɪɨɦɚɬ ɩɢɪɢɞɢɧɢɹ ɜ ɞɢɯɥɨɪɦɟɬɚɧɟ (PCC/CH2Cl2); ɛ) ɧɢɬɪɚɬ ɰɟɪɢɹ (IV) ɚɦɦɨɧɢɹ ɜ ɞɢɯɥɨɪɦɟɬɚɧɟ; ɜ) ɫɭɥɶɮɚɬ ɰɟɪɢɹ (IV) ɜ ɫɦɟɫɢ ɦɟɬɚɧɨɥ-ɜɨɞɚ; ɝ) ɩɟɪɦɚɧɝɚɧɚɬ ɤɚɥɢɹ ɜ ɛɟɧɡɨɥɟ ɜ ɩɪɢɫɭɬɫɬɜɢɢ 18-ɤɪɚɭɧ-6. ȼ ɭɫɥɨɜɢɹɯ ɨɤɢɫɥɟɧɢɹ ɩɟɪɦɚɧɝɚɧɚɬɨɦ ɤɚɥɢɹ ɜ ɛɟɧɡɨɥɟ, ɪɟɬɢɧɚɥɶ (1) ɨɤɚɡɚɥɫɹ ɭɫɬɨɣɱɢɜ (ɧɟ ɩɨɞɜɟɪɝɚɥɫɹ ɦɨɞɢɮɢɤɚɰɢɢ ɜ ɬɟɱɟɧɢɟ 2 ɧɟɞɟɥɶ), ɚ ɟɝɨ ɨɤɢɫɥɟɧɢɟ ɫɨɟɞɢɧɟɧɢɹɦɢ ɰɟɪɢɹ ɩɪɢɜɨɞɢɥɨ, ɜ ɨɫɧɨɜɧɨɦ, ɤ ɩɨɥɭɱɟɧɢɸ ɧɚɛɨɪɚ ɩɪɨɞɭɤɬɨɜ ɫ ɭɤɨɪɨɱɟɧɧɨɣ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɶɸ. ɇɚɢɛɨɥɟɟ ɢɧɬɟɪɟɫɧɵɟ ɪɟɡɭɥɶɬɚɬɵ, ɩɨɥɭɱɟɧɧɵɟ ɫ ɫɢɫɬɟɦɨɣ PCC / CH2Cl2, ɛɭɞɭɬ ɨɛɫɭɠɞɟɧɵ ɧɢɠɟ. ȼ ɥɢɬɟɪɚɬɭɪɟ ɨɩɢɫɚɧɨ ɩɪɢɦɟɧɟɧɢɟ ɫɨɟɞɢɧɟɧɢɣ ɯɪɨɦɚ (VI) ɞɥɹ ɨɤɢɫɥɢɬɟɥɶɧɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɪɟɬɢɧɨɢɞɨɜ. Ɍɚɤ, ɚɜɬɨɪɵ ɪɚɛɨɬɵ [173] ɨɤɢɫɥɹɥɢ ɪɟɬɢɧɨɟɜɭɸ ɤɢɫɥɨɬɭ (6) ɢ ɟɟ ɷɬɢɥɨɜɵɣ ɷɮɢɪ (8) ɩɪɢ ɩɨɦɨɳɢ ɪɟɚɤɬɢɜɚ Ⱦɠɨɧɫɚ (ɪɚɫɬɜɨɪ H2CrO4 ɜ ɚɰɟɬɨɧɟ). Ȼɵɥɨ ɨɩɪɟɞɟɥɟɧɨ, ɱɬɨ ɜ ɫɢɥɶɧɨɤɢɫɥɨɣ ɫɪɟɞɟ ɩɪɨɰɟɫɫ ɨɤɢɫɥɟɧɢɹ (ɫɦ. ɫɯɟɦɭ 2Ⱥ) ɢɞɟɬ ɱɟɪɟɡ ɩɟɪɟɝɪɭɩɩɢɪɨɜɤɭ 5,6-ɷɩɨɤɫɢɞɚ (9) ɜ 5,8-ɮɭɪɚɧɨɢɞɧɨɟ ɩɪɨɢɡɜɨɞɧɨɟ (10) ɫ ɩɨɫɥɟɞɭɸɳɢɦ ɪɚɫɳɟɩɥɟɧɢɟɦ ɫɜɹɡɢ ɋ7-ɋ8, ɷɬɨ ɩɪɢɜɨɞɢɬ ɤ ɩɨɥɭɱɟɧɢɸ ɫɦɟɫɢ ɩɪɨɞɭɤɬɨɜ ɫ ɧɢɡɤɢɦɢ ɜɵɯɨɞɚɦɢ (ɬɚɤ, ɜɵɯɨɞ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨ-ɪɟɬɢɧɨɟɜɨɣ ɤɢɫɥɨɬɵ (7) ɫɨɫɬɚɜɢɥ 0.8% , ɚ ɟɟ ɷɬɢɥɨɜɨɝɨ ɷɮɢɪɚ (11) – 12%). ɐɟɥɟɜɵɟ ɜɟɳɟɫɬɜɚ ɛɵɥɢ ɩɨɥɭɱɟɧɵ ɫ ɞɨɫɬɚɬɨɱɧɨ ɧɢɡɤɢɦɢ ɜɵɯɨɞɚɦɢ ɢ ɩɪɢ ɨɤɢɫɥɟɧɢɢ ɪɟɬɢɧɢɥɢɞɟɧɞɢɦɟɞɨɧɚ (14) [165] ɛɢɯɪɨɦɚɬɨɦ ɩɢɪɢɞɢɧɢɹ (ɫɦ. ɫɯɟɦɭ 2ȼ). ȼɵɯɨɞ 2-(5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɢɥɢɞɟɧ)ɞɢɦɟɞɨɧɚ (15) ɫɨɫɬɚɜɢɥ ɜ ɷɬɨɦ ɫɥɭɱɚɟ 34 %, ɢɡ ɧɟɝɨ ɛɵɥ ɩɨɥɭɱɟɧ ɫɨɨɬɜɟɬɫɬɜɭɸɳɢɣ ɚɥɶɞɟɝɢɞ (5) ɫ ɜɵɯɨɞɨɦ 12 % (ɜ ɩɟɪɟɫɱɟɬɟ ɧɚ ɪɟɬɢɧɢɥɢɞɟɧɞɢɦɟɞɨɧ (14)).

55 ɋɯɟɦɚ 2. Ɉɤɢɫɥɟɧɢɟ ɪɟɬɢɧɨɢɞɨɜ ɫɨɟɞɢɧɟɧɢɹɦɢ ɯɪɨɦɚ (VI).

A.

COOH

COOH

H2CrO4 O

(6)

(7)

O CO2Et (8)

H2CrO4

CO2Et

[H+]

CO2Et

O

O

(10)

(9)

H 2CrO4

H 2CrO4 CHO OH OOC

CO2Et O

(11)

CO2Et

+

(13)

(12)

O O

B. (14)

CO2Et

O

O Py2Cr2O7 O

O

(15)

O

O

CHO O

(5)

O

CHO PCC / CH 2Cl2

C. OH

(16)

CHO

O

(3)

ɂɧɬɟɪɟɫɧɨ ɨɬɦɟɬɢɬɶ, ɱɬɨ ɜ ɪɚɛɨɬɟ [166] ɞɥɹ ɫɢɧɬɟɡɚ 4-ɨɤɫɨɪɟɬɢɧɚɥɹ (3) ɢɫɩɨɥɶɡɨɜɚɥɢ ɜ ɤɚɱɟɫɬɜɟ ɨɤɢɫɥɢɬɟɥɹ ɫɢɫɬɟɦɭ PCC / CH2Cl2 (ɫɦ. ɫɯɟɦɭ 2ɋ). Ɉɞɧɚɤɨ ɜ ɷɬɨɦ ɫɥɭɱɚɟ ɢɫɯɨɞɧɵɦ ɫɨɟɞɢɧɟɧɢɟɦ ɛɵɥ 4-ɝɢɞɪɨɤɫɢɪɟɬɢɧɚɥɶ (16), ɚ ɜɵɯɨɞ 4ɨɤɫɨɪɟɬɢɧɚɥɹ (3) ɫɨɫɬɚɜɢɥ 50 %. Ɉɬɫɭɬɫɬɜɢɟ ɨɤɢɫɥɢɬɟɥɶɧɨɝɨ ɪɚɫɤɪɵɬɢɹ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɨɛɴɹɫɧɹɟɬɫɹ, ɩɨ ɜɫɟɣ ɜɟɪɨɹɬɧɨɫɬɢ, ɬɟɦ, ɱɬɨ ɢɫɯɨɞɧɵɦ ɛɵɥɨ ɩɪɟɞɜɚɪɢɬɟɥɶɧɨ ɨɤɢɫɥɟɧɧɨɟ ɩɨ ɱɟɬɜɟɪɬɨɦɭ ɩɨɥɨɠɟɧɢɸ ɩɪɨɢɡɜɨɞɧɨɟ ɪɟɬɢɧɚɥɹ (16).

56 ȼ ɩɪɟɞɥɨɠɟɧɧɨɦ ɜ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɟ ɜɚɪɢɚɧɬɟ (ɫɦ. ɫɯɟɦɭ 3), ɩɪɢ ɨɤɢɫɥɟɧɢɢ ɪɟɬɢɧɚɥɹ (1) ɜ ɫɢɫɬɟɦɟ PCC / CH2Cl2, ɜ ɨɫɧɨɜɧɨɦ, ɩɪɨɢɫɯɨɞɢɬ ɢɡɛɢɪɚɬɟɥɶɧɨɟ ɪɚɫɳɟɩɥɟɧɢɟ ɋ5-ɋ6-ɞɜɨɣɧɨɣ ɫɜɹɡɢ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɫ ɨɛɪɚɡɨɜɚɧɢɟɦ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɹ (5), ɚ ɬɚɤ ɠɟ ɨɛɪɚɡɨɜɚɧɢɟ 4-ɨɤɫɨ- (3) ɢ 5,6-ɞɢɝɢɞɪɨ-5,6ɷɩɨɤɫɢ-ɪɟɬɢɧɚɥɟɣ (17), ɜɵɯɨɞ ɫɨɫɬɚɜɥɹɟɬ 47%, 7% ɢ 19% ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ. ɋɯɟɦɚ 3. Ɉɤɢɫɥɟɧɢɟ ɪɟɬɢɧɚɥɹ (1) ɯɥɨɪɨɯɪɨɦɚɬɨɦ ɩɢɪɢɞɢɧɢɹ. CHO

CHO

PCC / CH2Cl2 O

(1)

(5)

+

O

CHO O

(17)

+ CHO

(3)

O

Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɩɪɟɞɥɨɠɟɧɧɵɣ ɦɟɬɨɞ ɫɢɧɬɟɡɚ ɩɨɡɜɨɥɹɟɬ ɩɨɥɭɱɢɬɶ 5,6-ɞɢɨɤɫɨ5,6-ɫɟɤɨ-ɩɪɨɢɡɜɨɞɧɨɟ ɪɟɬɢɧɚɥɹ ɫɨ ɡɧɚɱɢɬɟɥɶɧɨ ɛɨɥɟɟ ɜɵɫɨɤɢɦ ɜɵɯɨɞɨɦ, ɱɟɦ ɷɬɨ ɛɵɥɨ ɢɡɜɟɫɬɧɨ ɪɚɧɟɟ. 5,6-Ⱦɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɶ (5) ɹɜɥɹɟɬɫɹ ɩɟɪɫɩɟɤɬɢɜɧɵɦ ɢɫɯɨɞɧɵɦ ɫɨɟɞɢɧɟɧɢɟɦ ɞɥɹ ɜɜɟɞɟɧɢɹ ɪɚɡɥɢɱɧɨɝɨ ɪɨɞɚ ɦɟɬɨɤ ɢ ɪɟɩɨɪɬɟɪɧɵɯ ɝɪɭɩɩ ɜ ɦɨɥɟɤɭɥɵ ɪɟɬɢɧɨɢɞɨɜ, ɚ ɞɪɭɝɢɟ ɩɪɨɞɭɤɬɵ ɪɟɚɤɰɢɢ (4-ɨɤɫɨɪɟɬɢɧɚɥɶ (3) ɢ 5,6-ɞɢɝɢɞɪɨ-5,6-ɷɩɨɤɫɢɪɟɬɢɧɚɥɶ (17)) ɩɪɟɞɫɬɚɜɥɹɸɬ ɢɧɬɟɪɟɫ ɞɥɹ ɢɫɫɥɟɞɨɜɚɧɢɹ ȻɊ ɢ ɞɪɭɝɢɯ ɪɟɬɢɧɨɢɞ-ɛɟɥɤɨɜɵɯ ɤɨɦɩɥɟɤɫɨɜ. ɋɯɟɦɚ 4. ȼɨɡɦɨɠɧɵɣ ɦɟɯɚɧɢɡɦ ɪɟɚɤɰɢɢ ɨɤɢɫɥɟɧɢɹ ɪɟɬɢɧɚɥɹ (1) ɯɥɨɪɨɯɪɨɦɚɬɨɦ ɩɢɪɢɞɢɧɢɹ. CHO

PCC / CH2Cl2

CHO O

(17)

(1)

CHO O O

(5)

PCC / CH2Cl2

CHO OH OH

(18)

ɋɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ, ɱɬɨ ɜ ɞɚɧɧɨɦ ɫɥɭɱɚɟ (ɫɦ. ɫɯɟɦɭ 4) ɩɪɨɰɟɫɫ ɪɚɫɤɪɵɬɢɹ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɢɞɟɬ ɜ ɫɪɚɜɧɢɬɟɥɶɧɨ ɦɹɝɤɢɯ ɭɫɥɨɜɢɹɯ ɱɟɪɟɡ ɫɬɚɞɢɢ

57 ɨɛɪɚɡɨɜɚɧɢɹ ɩɪɨɦɟɠɭɬɨɱɧɵɯ 5,6-ɷɩɨɤɫɢɞɚ (17) ɢ ɝɥɢɤɨɥɹ (18), ɱɬɨ ɩɨɞɬɜɟɪɠɞɚɟɬɫɹ ɧɚɥɢɱɢɟɦ ɜ ɪɟɚɤɰɢɨɧɧɨɣ ɫɦɟɫɢ 5,6-ɞɢɝɢɞɪɨ-5,6-ɷɩɨɤɫɢɪɟɬɢɧɚɥɹ (17).

Ⱥ

7.0

12H

6.5

10H

6.0

8H 11H

B

A

5.5

14H

7H

7.0

6.5

6.0

5.5

2.0

1.5

1.0

0.5

1.0

0.5

1,1-CH3

9-CH3

13-CH3

2-CH2 5-CH3

4-CH2

B

2.0

3-CH2

1.5

1

Ɋɢɫ. 16. ɋɪɚɜɧɟɧɢɟ ɇ-əɆɊ ɫɩɟɤɬɪɨɜ Ⱥ) ɪɟɬɢɧɚɥɹ (1) ɢ ȼ) 5,6-ɞɢɨɤɫɨ-5,6ɫɟɤɨɪɟɬɢɧɚɥɹ (5). ɋɬɪɨɟɧɢɟ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɹ (5) ɛɵɥɨ ɞɨɤɚɡɚɧɨ ɫɩɟɤɬɪɚɥɶɧɵɦɢ ɦɟɬɨɞɚɦɢ. ȼ ɫɩɟɤɬɪɟ 13ɋ-əɆɊ (ɫɦ. ɷɤɫɩɟɪɢɦɟɧɬɚɥɶɧɭɸ ɱɚɫɬɶ) ɩɪɢɫɭɬɫɬɜɭɸɬ ɫɢɝɧɚɥɵ 20 ɚɬɨɦɨɜ ɭɝɥɟɪɨɞɚ, ɫɪɟɞɢ ɤɨɬɨɪɵɯ, ɧɚɪɹɞɭ ɫ ɫɢɝɧɚɥɨɦ ɚɥɶɞɟɝɢɞɧɨɣ ɝɪɭɩɩɵ (190.6 ɦ.ɞ.), ɩɪɢɫɭɬɫɬɜɭɸɬ ɞɜɚ ɞɨɩɨɥɧɢɬɟɥɶɧɵɯ ɫɢɝɧɚɥɚ ɤɚɪɛɨɧɢɥɶɧɨɣ ɝɪɭɩɩɵ (203.6 ɢ 208.2 ɦ.ɞ.), ɚ ɜ ɫɩɟɤɬɪɟ 1ɇ-əɆɊ ɨɬɦɟɱɟɧɨ ɧɚɥɢɱɢɟ ɫɢɝɧɚɥɨɜ ɩɪɨɬɨɧɨɜ ɜɫɟɯ ɮɪɚɝɦɟɧɬɨɜ ɫɬɪɭɤɬɭɪɵ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɹ (5), ɢ ɨɬɦɟɱɟɧɵ ɫɥɟɞɭɸɳɢɟ ɨɫɨɛɟɧɧɨɫɬɢ (ɫɦ. ɪɢɫ. 16),

58 ɩɨɞɬɜɟɪɠɞɚɸɳɢɟ ɫɬɪɭɤɬɭɪɭ (5), ɷɬɨ: ɡɧɚɱɢɬɟɥɶɧɵɣ ɫɞɜɢɝ ɜ ɨɛɥɚɫɬɶ ɫɥɚɛɨɝɨ ɩɨɥɹ ɫɢɝɧɚɥɨɜ ɩɪɨɬɨɧɨɜ 4ɋH2, 5-CH3, 7-H ɢ 8-H, all-E-ɤɨɧɮɢɝɭɪɚɰɢɹ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɩɨɞɬɜɟɪɠɞɚɟɬɫɹ ɯɚɪɚɤɬɟɪɧɨɣ ɞɥɹ ȿ-ɤɨɧɮɢɝɭɪɚɰɢɢ ɜɟɥɢɱɢɧɨɣ ɤɨɧɫɬɚɧɬ ɫɩɢɧ-ɫɩɢɧɨɜɨɝɨ ɜɡɚɢɦɨɞɟɣɫɬɜɢɹ J7-8 ɢ J11-12 15 Ƚɰ, J10-1111 Ƚɰ. Ⱦɨɩɨɥɧɢɬɟɥɶɧɵɦ ɩɨɞɬɜɟɪɠɞɟɧɢɟɦ ɭɦɟɧɶɲɟɧɢɹ ɤɨɥɢɱɟɫɬɜɚ ɫɨɩɪɹɠɟɧɧɵɯ ɞɜɨɣɧɵɯ ɫɜɹɡɟɣ ɋ=ɋ ɨɞɧɭ ɞɜɨɣɧɭɸ ɫɜɹɡɶ ɭ ɚɧɚɥɨɝɚ (5) ɫɥɭɠɚɬ ɷɥɟɤɬɪɨɧɧɵɟ ɫɩɟɤɬɪɵ ɜ ɦɟɬɚɧɨɥɟ: ɦɚɤɫɢɦɭɦ ɩɨɝɥɨɳɟɧɢɹ (364 ɧɦ) ɢɦɟɟɬ ɝɢɩɫɨɯɪɨɦɧɵɣ ɫɞɜɢɝ ɧɚ 16 ɧɦ ɨɬɧɨɫɢɬɟɥɶɧɨ ɪɟɬɢɧɚɥɹ (1). ɋɬɪɭɤɬɭɪɵ ɦɢɧɨɪɧɵɯ ɩɪɨɞɭɤɬɨɜ ɨɤɢɫɥɟɧɢɹ (3) ɢ (17) ɛɵɥɢ ɭɫɬɚɧɨɜɥɟɧɵ ɧɚ ɨɫɧɨɜɚɧɢɢ ɫɩɟɤɬɪɚɥɶɧɵɯ ɞɚɧɧɵɯ (ɬɚɛɥ. 7), ɚ ɬɚɤɠɟ ɩɭɬɟɦ ɫɪɚɜɧɟɧɢɹ ɢɯ ɮɢɡɢɤɨɯɢɦɢɱɟɫɤɢɯ ɫɜɨɣɫɬɜ (ɬ. ɩɥ. ɢ Rf) ɫɨ ɫɜɨɣɫɬɜɚɦɢ ɨɛɪɚɡɰɨɜ ɚɧɚɥɨɝɢɱɧɵɯ ɫɨɟɞɢɧɟɧɢɣ, ɫɢɧɬɟɡɢɪɨɜɚɧɧɵɯ ɪɚɧɟɟ ɜ ɧɚɲɟɣ ɥɚɛɨɪɚɬɨɪɢɢ ɞɪɭɝɢɦɢ ɦɟɬɨɞɚɦɢ (ɫɦ. ɬɚɛɥ. 7). Ɍɚɛɥɢɰɚ 7. Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɢɟ ɫɜɨɣɫɬɜɚ ɫɢɧɬɟɡɢɪɨɜɚɧɧɵɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ (2)-(5). ɍɎ-ɫɩɟɤɬɪ

N. CHO

(2)

Omax, ɧɦ

H, Ɇ-1ɫɦ-1

398

45700

Rf

tɩɥ., qɋ

Ʌɢɬ.

0.53

58-59

[154]

374

50500

0.39

113-115

[165] [169] [176]

280 391

13700 56700

0,39

121-124

[175]

364

58000

0.09

76-78

[165] [176]

362

53500

0.48

104-106

[165]

CHO

(3) O CHO

(4)

CHO

(5)

O O

(17)

CHO O

3. ɋɢɧɬɟɡ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɋɢɧɬɟɡ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ (2), (3) ɢ (5), ɧɟɨɛɯɨɞɢɦɵɯ ɞɥɹ ɪɟɝɟɧɟɪɚɰɢɢ ȺȻɊ (ɫɦ. ɫɯɟɦɭ 5) ɛɵɥ ɨɫɭɳɟɫɬɜɥɟɧ ɯɢɦɢɱɟɫɤɨɣ ɦɨɞɢɮɢɤɚɰɢɟɣ ɤɨɦɦɟɪɱɟɫɤɢ ɞɨɫɬɭɩɧɨɝɨ all-Eɪɟɬɢɧɚɥɹ (1), ɚ ɮɟɧɢɥɶɧɵɣ ɚɧɚɥɨɝ (4) ɛɵɥ ɩɨɥɭɱɟɧ ɩɨ ɪɚɡɪɚɛɨɬɚɧɧɨɣ ɜ ɧɚɲɟɣ ɥɚɛɨɪɚɬɨɪɢɢ ɪɚɧɟɟ ɫɯɟɦɟ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɋ10-ɫɢɧɬɨɧɚ (19) [152], ɤɨɬɨɪɚɹ ɛɵɥɚ ɩɪɟɞɥɨɠɟɧɚ ɞɥɹ ɩɪɢɝɨɬɨɜɥɟɧɢɹ ɪɹɞɚ ɚɪɨɦɚɬɢɱɟɫɤɢɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ. ɉɪɨɢɡɜɨɞɧɵɟ ɪɟɬɢɧɚɥɹ (2) ɢ (4) ɛɵɥɢ ɫɢɧɬɟɡɢɪɨɜɚɧɵ ɩɨ ɪɚɡɪɚɛɨɬɚɧɧɵɦ ɪɚɧɟɟ ɦɟɬɨɞɢɤɚɦ [105, 152], ɚ ɞɥɹ ɩɨɥɭɱɟɧɢɹ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨ- (5) ɢ 4-ɨɤɫɨɪɟɬɢɧɚɥɹ (3)

59 ɩɪɟɞɥɨɠɟɧ ɭɞɨɛɧɵɣ ɦɟɬɨɞ ɨɤɢɫɥɟɧɢɹ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɪɟɬɢɧɚɥɹ [176], ɤɨɬɨɪɵɣ ɩɨɞɪɨɛɧɨ ɪɚɫɫɦɨɬɪɟɧ ɜɵɲɟ (ɜ ɪɚɡɞɟɥɟ 2 ɬɟɨɪɟɬɢɱɟɫɤɨɣ ɱɚɫɬɢ). Ɉɥɟɮɢɧɢɪɨɜɚɧɢɟ ɤɚɪɛɨɧɢɥɶɧɵɯ ɫɨɟɞɢɧɟɧɢɣ ɩɨ ȼɢɬɬɢɝɭ ɢ ɪɹɞ ɟɝɨ ɦɨɞɢɮɢɤɚɰɢɣ ɨɬɧɨɫɹɬɫɹ ɤ ɨɞɧɨɦɭ ɢɡ ɨɫɧɨɜɧɵɯ ɫɩɨɫɨɛɨɜ ɩɨɥɭɱɟɧɢɹ ɩɨɥɢɟɧɨɜɵɯ ɫɨɟɞɢɧɟɧɢɣ [177179]. Ɋɚɧɟɟ ɞɥɹ ɫɢɧɬɟɡɚ ɚɪɨɦɚɬɢɱɟɫɤɢɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɜ ɧɚɲɟɣ ɥɚɛɨɪɚɬɨɪɢɢ ɛɵɥɨ ɩɪɟɞɥɨɠɟɧɨ ɫɨɡɞɚɜɚɬɶ ɭɝɥɟɪɨɞɧɵɣ ɫɤɟɥɟɬ ɨɞɧɨɤɪɚɬɧɵɦ ɧɚɪɚɳɢɜɚɧɢɟɦ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɩɪɢɫɨɟɞɢɧɟɧɢɟɦ ɋ10-ɮɪɚɝɦɟɧɬɚ (19) ɤ ɢɥɢɞɭ, ɩɨɥɭɱɟɧɧɨɦɭ ɢɡ ɚɪɢɥɬɪɢɮɟɧɢɥɮɨɫɮɨɧɢɣɝɚɥɨɝɟɧɢɞɚ [152]. ȼ ɷɬɨɦ ɫɥɭɱɚɟ ɞɥɹ ɝɟɧɟɪɢɪɨɜɚɧɢɹ ɢɥɢɞɚ ɦɨɝɭɬ ɛɵɬɶ ɢɫɩɨɥɶɡɨɜɚɧɵ [180, 181] ɤɚɤ ɫɢɥɶɧɵɟ ɨɫɧɨɜɚɧɢɹ (ɧ-ɛɭɬɢɥɥɢɬɢɣ, ɦɟɬɢɥɚɬ ɢɥɢ ɝɢɞɪɢɞ ɧɚɬɪɢɹ), ɬɚɤ ɢ ɞɨɫɬɚɬɨɱɧɨ ɫɥɚɛɵɟ (ɩɨɬɚɲ). Ɉɫɨɛɟɧɧɨɫɬɶɸ ɫɢɧɬɟɡɚ ɚɪɨɦɚɬɢɱɟɫɤɢɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɹɜɥɹɟɬɫɹ ɬɨ, ɱɬɨ ɪɟɚɤɰɢɹ ɨɥɟɮɢɧɢɪɨɜɚɧɢɹ ɩɪɨɬɟɤɚɟɬ ɫ ɪɚɡɥɢɱɧɨɣ ɫɬɟɩɟɧɶɸ ɫɬɟɪɟɨɫɟɥɟɤɬɢɜɧɨɫɬɢ [152], ɤɨɬɨɪɚɹ ɡɚɜɢɫɢɬ ɤɚɤ ɨɬ ɩɪɢɪɨɞɵ ɢ ɪɚɡɦɟɪɚ ɡɚɦɟɫɬɢɬɟɥɟɣ, ɧɚɯɨɞɹɳɢɯɫɹ ɜ ɚɪɨɦɚɬɢɱɟɫɤɨɦ ɮɪɚɝɦɟɧɬɟ, ɬɚɤ ɢ ɨɬ ɢɫɩɨɥɶɡɭɟɦɨɝɨ ɨɫɧɨɜɚɧɢɹ ɢ ɪɚɫɬɜɨɪɢɬɟɥɹ, ɚ ɤɚɬɚɥɢɬɢɱɟɫɤɨɟ ɝɢɞɪɢɪɨɜɚɧɢɟ ɬɪɨɣɧɨɣ ɫɜɹɡɢ ɬɚɤ ɠɟ ɬɟɪɹɟɬ ɫɬɟɪɟɨɫɟɥɟɤɬɢɜɧɨɫɬɶ. ȼ ɫɜɹɡɢ ɫ ɷɬɢɦ ɛɵɥɢ ɪɚɡɪɚɛɨɬɚɧɵ ɭɫɥɨɜɢɹ ɬɟɪɦɨɢɡɨɦɟɪɢɡɚɰɢɢ 11Z-ɢɡɨɦɟɪɚ4 ɜ 11E- ɧɚɝɪɟɜɚɧɢɟɦ ɪɚɫɬɜɨɪɚ ɪɟɬɢɧɨɢɞɚ ɩɪɢ 60qɋ ɜ ɬɟɱɟɧɢɟ 1-2 ɱ, ɩɪɢ ɷɬɨɦ ɩɨɥɭɱɚɟɬɫɹ ɫɦɟɫɶ ɢɡɨɦɟɪɨɜ 13Z- ɢ 13E- ɫ ɩɪɟɨɛɥɚɞɚɧɢɟɦ 13Z- [182]. Ⱦɥɹ ɩɨɥɭɱɟɧɢɹ all-E-ɩɪɨɢɡɜɨɞɧɵɯ ɩɪɢɦɟɧɹɸɬ ɛɨɥɟɟ ɠɟɫɬɤɢɟ ɭɫɥɨɜɢɹ ɬɟɪɦɨɢɡɨɦɟɪɢɡɚɰɢɢ ɜ ɩɪɢɫɭɬɫɬɜɢɢ ɫɥɟɞɨɜ ɢɨɞɚ, ɜ ɷɬɨɦ ɫɥɭɱɚɟ ɞɨɥɹ all-Eɢɡɨɦɟɪɚ ɫɨɫɬɚɜɥɹɟɬ ɨɤɨɥɨ 75% [152]. ȼ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɟ ɫɢɧɬɟɡ ɮɟɧɢɥɶɧɨɝɨ ɚɧɚɥɨɝɚ ɪɟɬɢɧɚɥɹ (4) (ɫɦ. ɫɯɟɦɭ 5) ɨɫɭɳɟɫɬɜɢɥɢ ɚɧɚɥɨɝɢɱɧɨ ɪɚɡɪɚɛɨɬɚɧɧɨɣ ɪɚɧɟɟ ɫɯɟɦɟ [152]. ȿɝɨ ɤɥɸɱɟɜɨɣ ɪɟɚɤɰɢɟɣ ɹɜɥɹɥɨɫɶ ɨɥɟɮɢɧɢɪɨɜɚɧɢɟ ɩɨ ȼɢɬɬɢɝɭ ɋ10-ɚɥɶɞɟɝɢɞɚ (19) ɢɥɢɞɨɦ, ɝɟɧɟɪɢɪɨɜɚɧɧɵɦ ɢɡ ɛɟɧɡɢɥɬɪɢɮɟɧɢɥɮɨɫɮɨɧɢɣ ɛɪɨɦɢɞɚ (20). Ⱦɚɧɧɚɹ ɪɟɚɤɰɢɹ ɩɪɨɬɟɤɚɟɬ ɫ ɞɨɫɬɚɬɨɱɧɨ ɧɢɡɤɨɣ ɫɬɟɩɟɧɶɸ ɫɬɟɪɟɨɫɟɥɟɤɬɢɜɧɨɫɬɢ ɢ ɩɪɢɜɨɞɢɬ ɤ ɩɨɥɭɱɟɧɢɸ ɫɦɟɫɢ ɢɡɨɦɟɪɨɜ ɩɨ ɫɜɹɡɢ ɋ7-ɋ8 4 ɜ ɫɨɨɬɧɨɲɟɧɢɢ (7E,13Z):(7Z,13Z) ɤɚɤ 2:3, ɱɬɨ ɛɵɥɨ ɨɩɪɟɞɟɥɟɧɨ ɩɨ ɫɨɨɬɧɨɲɟɧɢɸ ɩɥɨɳɚɞɟɣ ɩɨɞ ɫɢɝɧɚɥɚɦɢ 7ɇ (ɢɥɢ 8ɇ) ɜ ɫɩɟɤɬɪɟ 1ɇ-əɆɊ ɪɟɚɤɰɢɨɧɧɨɣ ɫɦɟɫɢ (ɞɥɹ 7Z-ɫɜɹɡɢ ɯɚɪɚɤɬɟɪɧɚ ɤɨɧɫɬɚɧɬɚ ɫɩɢɧ-ɫɩɢɧɨɜɨɝɨ ɜɡɚɢɦɨɞɟɣɫɬɜɢɹ ɦɟɠɞɭ 7ɇ ɢ 8ɇ ɪɚɜɧɚɹ 12 Ƚɰ, ɚ ɞɥɹ 7E – 16 Ƚɰ). ɉɨɫɥɟ ɫɧɹɬɢɹ ɬɪɢɮɟɧɢɥɫɢɥɢɥɶɧɨɣ ɡɚɳɢɬɵ ɢ ɜɵɞɟɥɟɧɢɹ 7ȿ,13Z-ɢɡɨɦɟɪɚ ɤɨɥɨɧɨɱɧɨɣ ɯɪɨɦɚɬɨɝɪɚɮɢɟɣ ɜɵɯɨɞ ɮɟɧɢɥɶɧɨɝɨ ɚɧɚɥɨɝɚ 11,12-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɨɥɚ (22) ɫɨɫɬɚɜɢɥ 34%. ȿɝɨ ɨɤɢɫɥɟɧɢɟ ɞɢɨɤɫɢɞɨɦ ɦɚɪɝɚɧɰɚ ɩɪɢɜɨɞɢɥɨ ɤ ɚɥɶɞɟɝɢɞɭ (23) ɫ 4

ɉɪɢ ɨɛɫɭɠɞɟɧɢɢ ɫɩɟɤɬɪɨɜ ɢ ɢɡɨɦɟɪɧɨɝɨ ɫɨɫɬɚɜɚ ɩɪɟɩɚɪɚɬɨɜ ɚɪɨɦɚɬɢɱɟɫɤɢɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɧɭɦɟɪɚɰɢɹ ɚɬɨɦɨɜ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ (ɜ ɬɨɦ ɱɢɫɥɟ ɭ ɚɧɚɥɨɝɚ (4) ɢ ɟɝɨ ɩɪɟɞɲɟɫɬɜɟɧɧɢɤɨɜ) ɩɪɢɧɹɬɚ ɤɚɤ ɭ ɩɪɢɪɨɞɧɨɝɨ ɪɟɬɢɧɚɥɹ (1)

60 ɜɵɯɨɞɨɦ 82%. ɉɨɫɥɟ ɜɨɫɫɬɚɧɨɜɥɟɧɢɹ ɬɪɨɣɧɨɣ ɫɜɹɡɢ ɤɚɬɚɥɢɬɢɱɟɫɤɢɦ ɝɢɞɪɢɪɨɜɚɧɢɟɦ ɢ ɩɨɫɥɟɞɭɸɳɟɣ ɬɟɪɦɨɢɡɨɦɟɪɢɡɚɰɢɢ ɤɢɩɹɱɟɧɢɟɦ ɜ ɝɟɤɫɚɧɟ ɜ ɩɪɢɫɭɬɫɬɜɢɢ ɫɥɟɞɨɜɵɯ ɤɨɥɢɱɟɫɬɜ ɢɨɞɚ ɜɵɯɨɞ all-E-ɢɡɨɦɟɪɚ ɮɟɧɢɥɶɧɨɝɨ ɚɧɚɥɨɝɚ ɪɟɬɢɧɚɥɹ (4) ɫɨɫɬɚɜɢɥ 73% (ɜ ɪɚɫɫɱɟɬɟ ɧɚ ɚɥɶɞɟɝɢɞ (23)). ɋɯɟɦɚ 5. ɋɯɟɦɚ ɫɢɧɬɟɡɚ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ (2)-(5). 1) PPh3 Br

O

(19)

OSiPh3

K2CO3, 18-ɤɪɚɭɧ-6

(21)

2) F

(22)

-

OH

MnO2

PPh3 Br

(23)

(20)

O

1) H2, Pd / CaCO3 / Pb(OAc)2 2) ', 80 oC

O

(4)

CHO

CHO

PCC / CH2Cl2 O

(1)

(5)

+

O

NBS

CHO O CHO

Br

(17)

+

(24)

CHO KF / ȾɆɎȺ O

(3)

CHO

(2)

Ɇɟɬɨɞ ɩɨɥɭɱɟɧɢɹ ɢɫɩɨɥɶɡɨɜɚɧɧɨɝɨ ɜ ɩɪɢɜɟɞɟɧɧɨɦ ɜɵɲɟ ɫɢɧɬɟɡɟ ɋ10-ɚɥɶɞɟɝɢɞɚ, 2E,6Z-8-ɬɪɢɮɟɧɢɥɫɢɥɢɥɨɤɫɢ-2,6-ɞɢɦɟɬɢɥɨɤɬɚ-2,6-ɞɢɟɧ-4-ɢɧ-1-ɚɥɹ (19) ɛɵɥ ɪɚɡɪɚɛɨɬɚɧ ɪɚɧɟɟ ɜ ɧɚɲɟɣ ɥɚɛɨɪɚɬɨɪɢɢ [183]. 3,4-Ⱦɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɶ (2) ɫ ɜɵɯɨɞɨɦ 50% ɩɨɥɭɱɢɥɢ ɚɥɥɢɥɶɧɵɦ ɛɪɨɦɢɪɨɜɚɧɢɟɦ ɪɟɬɢɧɚɥɹ (1) N-ɛɪɨɦɫɭɤɰɢɧɢɦɢɞɨɦ (NBS) ɫ ɩɨɫɥɟɞɭɸɳɢɦ ɞɟɝɢɞɪɨɛɪɨɦɢɪɨɜɚɧɢɟɦ ɩɪɨɦɟɠɭɬɨɱɧɨɝɨ 4-ɛɪɨɦɪɟɬɢɧɚɥɹ (24) ɞɟɣɫɬɜɢɟɦ ɫɭɫɩɟɧɡɢɢ ɮɬɨɪɢɞɚ ɤɚɥɢɹ ɜ ȾɆɎȺ.

61 4-Ɉɤɫɨ- (3) ɢ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɶ (5) ɩɨɥɭɱɚɥɢ ɩɪɟɞɥɨɠɟɧɧɵɦ ɜ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɟ ɦɟɬɨɞɨɦ [176] ɫ ɜɵɯɨɞɨɦ 7% ɢ 47% ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ (ɨɛɫɭɠɞɟɧɢɟ ɫɦ. ɜ ɪɚɡɞɟɥɟ 2 ɬɟɨɪɟɬɢɱɟɫɤɨɣ ɱɚɫɬɢ ɧɚ ɫ. 53). ɋɬɪɨɟɧɢɟ ɢ ɱɢɫɬɨɬɚ ɩɪɟɩɚɪɚɬɨɜ ɜɫɟɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ (2)-(5), (22), (23) ɛɵɥɨ ɩɨɞɬɜɟɪɠɞɟɧɨ ɦɟɬɨɞɚɦɢ ɍɎ- ɢ 1ɇ-əɆɊ ɫɩɟɤɬɪɨɫɤɨɩɢɢ ɢ ɫɪɚɜɧɟɧɢɟɦ ɢɯ ɮɢɡɢɤɨɯɢɦɢɱɟɫɤɢɯ ɫɜɨɣɫɬɜ ɫɨ ɫɜɨɣɫɬɜɚɦɢ ɨɛɪɚɡɰɨɜ ɚɧɚɥɨɝɢɱɧɵɯ ɫɨɟɞɢɧɟɧɢɣ, ɫɢɧɬɟɡɢɪɨɜɚɧɧɵɯ ɜ ɧɚɲɟɣ ɥɚɛɨɪɚɬɨɪɢɢ ɪɚɧɟɟ (ɞɚɧɧɵɟ ɩɪɢɜɟɞɟɧɵ ɜ ɬɚɛɥ. 7 ɧɚ ɫ. 58).

(2) 4

ɋɥɚɛɨɟ ɩɨɥɟ

(3)

'G, ɦ.ɞ.

3

(4) (5)

2

(17)

1

0

13-CH m3

9-CHl3

k 14H

12Hj

11Hi

h 10H

g 8H

7Hf

5-CHe3

4CHd2

3CHc2

2CHb2

1,1-CHa3

-1

Ƚɪɭɩɩɵ ɚɬɨɦɨɜ

Ɋɢɫ. 17. ɂɡɦɟɧɟɧɢɟ ɯɢɦɢɱɟɫɤɨɝɨ ɫɞɜɢɝɚ ɫɢɝɧɚɥɨɜ ɩɪɨɬɨɧɨɜ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ (2)(5) ɢ (17) ɨɬɧɨɫɢɬɟɥɶɧɨ ɪɟɬɢɧɚɥɹ (1). ɉɪɢ ɫɪɚɜɧɟɧɢɢ 1ɇ-əɆɊ-ɫɩɟɤɬɪɨɜ ɨɫɧɨɜɧɵɯ ɩɪɨɞɭɤɬɨɜ ɪɟɚɤɰɢɢ ɨɤɢɫɥɟɧɢɹ ɪɟɬɢɧɚɥɹ (1) ɯɥɨɪɨɯɪɨɦɚɬɨɦ ɩɢɪɢɞɢɧɢɹ (3), (5), (17) ɫ ɩɚɪɚɦɟɬɪɚɦɢ ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɝɨ ɫɩɟɤɬɪɚ ɢɫɯɨɞɧɨɝɨ ɪɟɬɢɧɚɥɹ (1) (ɞɚɧɧɵɟ ɩɪɟɞɫɬɚɜɥɟɧɵ ɧɚ ɪɢɫ. 17) ɦɨɠɧɨ ɜɵɞɟɥɢɬɶ ɫɥɟɞɭɸɳɢɟ ɬɟɧɞɟɧɰɢɢ: ɭ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɹ (5) ɫɢɝɧɚɥɵ 3CH2 ɢ 11-ɇ ɫɦɟɳɟɧɵ ɜ ɫɢɥɶɧɨɟ ɩɨɥɟ ('G-0.24 ɢ -0.18 ɦ.ɞ., ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ), ɚ ɫɢɝɧɚɥɵ 4CH2, 5-CH3, 8-ɇ, 10-ɇ ɢ 12-ɇ ɜ ɨɛɥɚɫɬɶ ɫɥɚɛɨɝɨ ɩɨɥɹ ('G0.24, 0.25, 1.03, ɢ 0.16 ɦ.ɞ. ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ). ȼ ɫɩɟɤɬɪɟ 5,6-ɞɢɝɢɞɪɨ-5,6-ɷɩɨɤɫɢɪɟɬɢɧɚɥɹ (17) ɯɚɪɚɤɬɟɪɧɵ ɫɥɚɛɨɩɨɥɶɧɵɟ ɫɞɜɢɝɢ ɫɢɝɧɚɥɨɜ 3-CH2 ('G 0.14 ɦ.ɞ.) ɢ ɫɢɥɶɧɨɩɨɥɶɧɵɟ – 4CH2, 5-CH3, 8-ɇ ('G -0.13, -0.78, -0.13 ɦ.ɞ.), ɚ ɬɚɤ ɠɟ ɨɬɦɟɱɟɧɚ ɧɟɷɤɜɢɜɚɥɟɧɬɧɨɫɬɶ ɦɟɬɢɥɶɧɵɯ ɝɪɭɩɩ 1,1-CH3 (ɨɧɢ ɞɚɸɬ ɫɢɝɧɚɥɵ ɫ G1.16 ɢ 1.11 ɦ.ɞ.), ɤɨɬɨɪɚɹ ɧɟ ɧɚɛɥɸɞɚɟɬɫɹ ɭ ɛɨɥɶɲɢɧɫɬɜɚ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ. Ⱦɥɹ 4-ɨɤɫɨɪɟɬɢɧɚɥɹ (3) ɨɬɦɟɱɟɧɵ ɫɥɚɛɨɩɨɥɶɧɵɟ ɫɞɜɢɝɢ ɫɢɝɧɚɥɨɜ 2CH2 ɢ 3CH2 ('G 0.37 ɢ 0.89 ɦ.ɞ.). Ⱦɥɹ 3,4-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɹ (2) ɯɚɪɚɤɬɟɪɟɧ

62 ɡɧɚɱɢɬɟɥɶɧɵɣ ɫɥɚɛɨɩɨɥɶɧɵɣ ɫɞɜɢɝ ɫɢɝɧɚɥɨɜ 3-ɇ ɢ 4-ɇ (ɧɚ 4,13 ɢ 3,81 ɦ.ɞ. ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ). Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɜ ɷɬɨɣ ɱɚɫɬɢ ɪɚɛɨɬɵ ɛɵɥ ɨɫɭɳɟɫɬɜɥɟɧ ɫɢɧɬɟɡ ɪɹɞɚ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɫ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɦ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɵɦ ɤɨɥɶɰɨɦ (2)-(5). ɉɪɢ ɷɬɨɦ ɛɵɥ ɩɪɟɞɥɨɠɟɧ ɧɨɜɵɣ ɦɟɬɨɞ ɨɤɢɫɥɟɧɢɹ ɤɨɥɶɰɚ ɪɟɬɢɧɚɥɹ (1), ɩɨɡɜɨɥɹɸɳɢɣ ɩɨɥɭɱɢɬɶ ɟɝɨ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨ-ɩɪɨɢɡɜɨɞɧɨɟ ɫɨ ɡɧɚɱɢɬɟɥɶɧɨ ɛɨɥɶɲɢɦ ɜɵɯɨɞɨɦ, ɱɟɦ ɷɬɨ ɛɵɥɨ ɢɡɜɟɫɬɧɨ ɪɚɧɟɟ, ɚ ɬɚɤ ɠɟ ɛɵɥɢ ɜɨɫɩɪɨɢɡɜɟɞɟɧɵ ɢɫɩɨɥɶɡɨɜɚɜɲɢɟɫɹ ɧɚ ɤɚɮɟɞɪɟ ɛɢɨɬɟɯɧɨɥɨɝɢɢ ɆɂɌɏɌ ɢɦ. Ɇ.ȼ. Ʌɨɦɨɧɨɫɨɜɚ ɦɟɬɨɞɢɤɢ ɩɨɥɭɱɟɧɢɹ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɦɨɞɢɮɢɤɚɰɢɟɣ ɫɚɦɨɝɨ ɪɟɬɢɧɚɥɹ (1) ɢɥɢ ɧɚɪɚɳɢɜɚɧɢɟɦ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɭ ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɝɨ ɰɢɤɥɢɱɟɫɤɨɝɨ ɮɪɚɝɦɟɧɬɚ ɫ ɩɨɦɨɳɶɸ ɨɥɟɮɢɧɢɪɨɜɚɧɢɹ ɜɨ ȼɢɬɬɢɝɭ.

4. ɏɪɨɦɨɮɨɪɧɚɹ ɩɨɥɨɫɬɶ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ Ɇɨɥɟɤɭɥɚ ȻɊ ɫɨɫɬɨɢɬ ɢɡ 248 ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ. ɋɪɟɞɢ ɧɢɯ ɦɨɠɧɨ ɜɵɞɟɥɢɬɶ ɬɟ, ɤɨɬɨɪɵɟ ɫ ɧɚɢɛɨɥɶɲɟɣ ɜɟɪɨɹɬɧɨɫɬɶɸ ɦɨɝɭɬ ɢɝɪɚɬɶ ɮɭɧɤɰɢɨɧɚɥɶɧɭɸ ɪɨɥɶ ɜ ɦɨɥɟɤɭɥɟ ȻɊ (ɜɫɟɝɨ ɢɯ 107): ɩɨɥɨɠɢɬɟɥɶɧɨ ɡɚɪɹɠɟɧɧɵɟ (Arg ɢ Lys), ɚɪɨɦɚɬɢɱɟɫɤɢɟ (Trp, Tyr , Phe), ɨɬɪɢɰɚɬɟɥɶɧɨ ɡɚɪɹɠɟɧɧɵɟ (Asp, Glu), ɫ ɡɚɤɪɟɩɥɟɧɧɨɣ ɤɨɥɶɰɟɜɨɣ ɫɬɪɭɤɬɭɪɨɣ (Pro) ɢ ɝɢɞɪɨɤɫɢɥ-ɫɨɞɟɪɠɚɳɢɟ (Tyr, Ser, Thr). Ʉɨɥɢɱɟɫɬɜɨ ɢ ɪɚɫɩɨɥɨɠɟɧɢɟ ɷɬɢɯ ɨɫɬɚɬɤɨɜ ɜ ɩɟɪɜɢɱɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ ȻɊ ɩɪɟɞɫɬɚɜɥɟɧɨ ɜ ɬɚɛɥɢɰɟ 8. Ɍɚɛɥɢɰɚ 8. ɋɨɞɟɪɠɚɧɢɟ ɢ ɪɚɫɩɨɥɨɠɟɧɢɟ ɜ ɩɟɪɜɢɱɧɨɣ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ ȻɊ ɚɪɨɦɚɬɢɱɟɫɤɢɯ, ɡɚɪɹɠɟɧɧɵɯ ɢ ɝɢɞɪɨɤɫɢɥɫɨɞɟɪɠɚɳɢɯ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɢ ɩɪɨɥɢɧɚ. ȺɄ, ɤɨɥ-ɜɨ ɜ ȻɊ Arg (7) 7

ɉɨɥɨɠɟɧɢɟ ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɝɨ ɨɫɬɚɬɤɚ ɚɦɢɧɨɤɢɫɥɨɬɵ ɜ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ ȻɊ 82 134 164 175 225 227

Lys (7) 30 40 41 129 159 172 216 Asp (9) 36 38 85 96 102 104 115 212 242 Glu (10) 1

9

74 161 166 194 204 232 234 237

Pro (11) 8

37 50 70 77 91 165 186 200 236 238

Trp (8) 10 12 80 86 137 138 182 189 Phe (13) 27 42 54 71 88 135 153 154 156 171 208 219 230 Tyr (11) 26 43 57 64 79 83 131 133 147 150 185 Ser (13) 35 59 132 141 158 162 169 183 193 214 226 239 248 Thr (18) 5

17 24 46 47 55 67 89 90 107 121 128 142 157 170 178 205 247

63 Ⱦɥɹ ɨɰɟɧɤɢ ɜɨɡɦɨɠɧɨɫɬɢ ɜɫɬɪɚɢɜɚɧɢɹ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɜ ȻɈ ɧɟɨɛɯɨɞɢɦɨ ɨɰɟɧɢɬɶ ɩɪɨɫɬɪɚɧɫɬɜɟɧɧɵɟ ɨɝɪɚɧɢɱɟɧɢɹ, ɧɚɤɥɚɞɵɜɚɟɦɵɟ ɪɚɡɦɟɪɚɦɢ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ȻɊ ɧɚ ɫɬɪɭɤɬɭɪɭ ɯɪɨɦɨɮɨɪɚ, ɱɬɨ ɦɨɠɧɨ ɨɫɭɳɟɫɬɜɢɬɶ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɢɦɟɸɳɢɯɫɹ ɤ ɧɚɫɬɨɹɳɟɦɭ ɦɨɦɟɧɬɭ ɬɪɟɯɦɟɪɧɵɯ ɫɬɪɭɤɬɭɪ ȻɊ [184-186], ɩɨɥɭɱɟɧɧɵɯ ɦɟɬɨɞɨɦ ɪɟɧɬɝɟɧɨ-ɫɬɪɭɤɬɭɪɧɨɝɨ ɚɧɚɥɢɡɚ. ɂɫɯɨɞɧɵɟ ɮɚɣɥɵ ɧɚɯɨɞɹɬɫɹ ɜ Brookhaven Protein Data Bank. ɉɪɨɜɟɞɟɧɧɵɣ ɜ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɟ ɚɧɚɥɢɡ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ɩɪɨɜɨɞɢɥɢ ɢɡɦɟɪɟɧɢɟɦ ɪɚɫɫɬɨɹɧɢɣ ɦɟɠɞɭ ɚɬɨɦɚɦɢ ɭɝɥɟɪɨɞɚ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ (1) ɢ ɛɥɢɠɚɣɲɢɦɢ ɚɬɨɦɚɦɢ (ɋ, N, O) ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ. ɍɱɢɬɵɜɚɹ, ɱɬɨ ɞɥɢɧɚ ɫɜɹɡɢ ɋ-ɋ ɫɨɫɬɚɜɥɹɟɬ ɨɤɨɥɨ 1,54 Å, ɪɚɫɫɦɚɬɪɢɜɚɥɢɫɶ ɬɨɥɶɤɨ ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɭɞɚɥɟɧɧɵɟ ɧɟ ɛɨɥɟɟ ɱɟɦ ɧɚ 6 Å ɨɬ ɪɟɬɢɧɚɥɹ (1). Ⱦɥɹ ɢɡɦɟɪɟɧɢɣ ɛɵɥɚ ɢɫɩɨɥɶɡɨɜɚɧɚ ɩɪɨɝɪɚɦɦɚ Chem3D Ultra ɢɡ ɩɚɤɟɬɚ ChemOffice 6.0 (CambridgeSoft Com, USA). ȼ ɪɟɡɭɥɶɬɚɬɟ ɛɵɥɢ ɩɨɥɭɱɟɧɵ ɫɥɟɞɭɸɳɢɟ ɞɚɧɧɵɟ (ɬɚɛɥ. 9). Ɉɰɟɧɤɚ ɩɪɨɢɡɜɨɞɢɥɚɫɶ ɩɨ ɫɥɟɞɭɸɳɢɦ ɬɨɱɤɚɦ (ɚɬɨɦɚɦ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ (1)): ɋ1-, ɋ2-, ɋ3- ɢ ɋ4– ɚɬɨɦɵ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ, ɋ18- (5-ɋɇ3), ɋ19- (9-ɋɇ3) ɢ ɋ20- (13-ɋɇ3) ɚɬɨɦɵ ɦɟɬɢɥɶɧɵɯ ɝɪɭɩɩ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɢ NH-ɚɬɨɦ ɨɫɧɨɜɚɧɢɹ ɒɢɮɮɚ Lys–216 ɯɪɨɦɨɮɨɪɧɨɣ ɝɪɭɩɩɵ ȻɊ. ȼ ɬɚɛɥ. 9 ɩɪɟɞɫɬɚɜɥɟɧɵ ɪɚɫɫɬɨɹɧɢɹ ɦɟɠɞɭ ɪɟɩɟɪɧɵɦɢ ɬɨɱɤɚɦɢ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɢ ɚɦɢɧɨɤɢɫɥɨɬɧɵɦɢ ɨɫɬɚɬɤɚɦɢ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ȻɊ, ɪɚɫɩɨɥɨɠɟɧɧɵɦɢ ɛɥɢɠɟ 6Å. Ⱥɧɚɥɢɡ ɞɚɧɧɵɯ (ɬɚɛɥ. 9) ɩɨɤɚɡɵɜɚɟɬ, ɱɬɨ ɛɥɢɡɥɟɠɚɳɢɟ ɤ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɦɭ ɤɨɥɶɰɭ ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ ɪɚɫɩɨɥɨɠɟɧɵ ɧɚ D– ɫɩɢɪɚɥɶɧɵɯ ɬɹɠɚɯ C, D ɢ ȿ. ɂɡ ɩɪɢɜɟɞɟɧɧɵɯ ɜ ɬɚɛɥ. 9 ɞɚɧɧɵɯ ɦɨɠɧɨ ɫɞɟɥɚɬɶ ɡɚɤɥɸɱɟɧɢɟ, ɱɬɨ ɜ ɪɚɣɨɧɟ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɯɪɨɦɨɮɨɪɧɭɸ ɩɨɥɨɫɬɶ ɮɨɪɦɢɪɭɸɬ ɫɥɟɞɭɸɳɢɟ ɨɫɬɚɬɤɢ ɚɦɢɧɨɤɢɫɥɨɬ: Tyr83, Trp86, Met118, Gly122, Trp138, Ser141, Thr142, Met145, Tyr185, Pro186, Trp189. ȼ ɪɚɣɨɧɟ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɜ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ɪɚɫɩɨɥɨɠɟɧɵ Asp85, Trp86, Thr89, Thr90, Ile119, Met145, Trp182, Tyr185, Pro186, Asp212. ɇɚɢɛɨɥɟɟ ɛɥɢɡɤɢɟ ɤɨɧɬɚɤɬɵ (ɦɟɧɟɟ 4Å) ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ȻɊ ɫ ɚɬɨɦɚɦɢ ɪɟɬɢɧɚɥɹ:

ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɟ ɤɨɥɶɰɨ – Trp138, Ser141, Thr142, Met145, Pro186 ɢ

Trp189, ɪɚɫɩɨɥɨɠɟɧɧɵɟ ɜ ɬɹɠɚɯ E ɢ F;

ɋ9, 9-ɋɇ3 – Met118, Trp182, Tyr185, ɪɚɫɩɨɥɨɠɟɧɧɵɟ ɜ ɬɹɠɚɯ D ɢ F;

ɋ13, 13-ɋɇ3 – Thr90, Trp182, Tyr185, ɪɚɫɩɨɥɨɠɟɧɧɵɟ ɜ ɬɹɠɚɯ ɋ ɢ F;

NH-(SB) – Asp85, Thr89, Asp212, ɪɚɫɩɨɥɨɠɟɧɧɵɟ ɜ ɬɹɠɚɯ ɋ ɢ G.

64 Ɍɚɛɥɢɰɚ 9. Ɋɚɫɫɬɨɹɧɢɹ ɦɟɠɞɭ ɨɬɞɟɥɶɧɵɦɢ ɚɬɨɦɚɦɢ ɮɭɧɤɰɢɨɧɚɥɶɧɵɯ ɚɦɢɧɨɤɢɫɥɨɬ, ɨɛɪɚɡɭɸɳɢɯ ɯɪɨɦɨɮɨɪɧɭɸ ɩɨɥɨɫɬɶ, ɢ ɚɬɨɦɚɦɢ ɯɪɨɦɨɮɨɪɧɨɣ ɝɪɭɩɩɵ. Ⱥɦɢɧɨɤɢɫɥɨɬɧɵɣ ɨɫɬɚɬɨɤ*, ɚɬɨɦ Tyr 83, -OH Asp 85, -COOH Trp 86, 6c-C Trp 86, N Thr 89, -OH Thr 90, N (ɝ.) Thr 90, CJ Met 118, -SCH3 Met 118, -SCH3 Ile 119, CG Gly 122, CD Trp 138, CD Trp 138, N (ɝ.) Ser 141, C=O Ser 141, -OH Thr 142, N (ɝ.) Thr 142, -OH Met 145, -SCH3 Met 145, CE Trp 182, C=O Trp 182, Nc Tyr 185, -OH Tyr 185, 1c-C Pro 186, CJ Pro 186, N (ɝ.) Trp 189, CE Trp 189, N (ɝ.) Leu211, D-NH2 Asp 212, C=O

ɋ1 5.6

Ɋɚɫɫɬɨɹɧɢɟ (Å) ɦɟɠɞɭ ɚɬɨɦɚɦɢ ɯɪɨɦɨɮɨɪɧɨɣ ɝɪɭɩɩɵ ɢ ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɣ ȺɄ ɋ2 ɋ3 ɋ4 5-ɋɇ3 ɋ9 9-ɋɇ3 ɋ13 13-ɋɇ3 NH-(SB) 3.7 ɢ 4.0

4.4

5.5

5.0 4.3 5.0 5.5 3.7

6.4 6.5 6.2 4.9

6.4

4.5 5.6

5.6 5.5 3.9

6.2

4.0 3.6 4.9

4.4

5.9

4.8

4.4

4.4

4.2 4.3 4.3 3.3 5.2 5.1

5.2 4.5 3.9 3.5 3.2 4.1 3.8

6.5 5.6 3.4 3.3 3.6 3.8 4.3 3.9 6.2

4.5 6.4

4.2 5.2 3.6 5.7

4.8 3.5

5.0 5.7 5.3 4.9 4.5 4.7

4.2 3.8

3.7 5.4 3.6 3.7

5.8 4.5 4.7

4.5 5.2 5.5 6.1

3.5 5.0

5.7

5.2 5.7

4.3 OH

,

5.8 3.7 ɢ 4.8

5.4

Trp1'

Tyr 1' * ɇɭɦɟɪɚɰɢɹ ɚɬɨɦɨɜ ɜ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɚɯ:

7.9

3.5 4.1 4.2

N

6'

.

5. Ⱥɧɚɥɢɡ ɞɚɧɧɵɯ ɩɨ ɜɥɢɹɧɢɸ ɬɨɱɟɱɧɵɯ ɦɭɬɚɰɢɣ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ ɧɚ ɫɜɨɣɫɬɜɚ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ Ʉɚɤ ɭɠɟ ɨɬɦɟɱɚɥɨɫɶ ɜ ɝɥɚɜɟ 4.3 ɥɢɬɟɪɚɬɭɪɧɨɝɨ ɨɛɡɨɪɚ, ɡɚ ɩɪɨɲɟɞɲɟɟ ɜɪɟɦɹ ɛɵɥ ɧɚɤɨɩɥɟɧ ɛɨɥɶɲɨɣ ɦɚɫɫɢɜ ɢɧɮɨɪɦɚɰɢɢ ɨ ɫɜɨɣɫɬɜɚɯ ɨɤɨɥɨ 150 ɜɚɪɢɚɧɬɨɜ (ɧɚ ɪɢɫ. 18 ɨɧɢ ɜɵɞɟɥɟɧɵ ɤɪɭɠɤɚɦɢ) ȻɊ ɫ ɟɞɢɧɢɱɧɵɦɢ ɢ ɦɧɨɠɟɫɬɜɟɧɧɵɦɢ ɡɚɦɟɧɚɦɢ ɚɦɢɧɨɤɢɫɥɨɬ.

65 Ɉɞɧɚɤɨ ɞɨ ɫɢɯ ɩɨɪ ɧɟ ɛɵɥɨ ɨɫɭɳɟɫɬɜɥɟɧɨ ɫɢɫɬɟɦɚɬɢɡɚɰɢɢ ɩɨɥɭɱɟɧɧɵɯ ɪɟɡɭɥɶɬɚɬɨɜ. ɉɨ ɷɬɨɣ ɩɪɢɱɢɧɟ ɛɵɥɨ ɧɟɨɛɯɨɞɢɦɨ ɩɪɨɜɟɫɬɢ ɫɪɚɜɧɢɬɟɥɶɧɵɣ ɚɧɚɥɢɡ ɨɩɭɛɥɢɤɨɜɚɧɧɵɯ ɜ ɥɢɬɟɪɚɬɭɪɟ ɫɜɟɞɟɧɢɣ ɩɨ ɜɥɢɹɧɢɸ ɬɨɱɟɱɧɵɯ ɡɚɦɟɧ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɜ ɦɨɥɟɤɭɥɟ ȻɊ ɧɚ ɟɝɨ ɫɜɨɣɫɬɜɚ. Ɍɚɤɨɣ ɚɧɚɥɢɡ ɩɨɡɜɨɥɢɥ ɜɵɞɟɥɢɬɶ ɨɛɥɚɫɬɢ ɫɬɪɭɤɬɭɪɵ ɛɟɥɤɚ, ɜ ɤɨɬɨɪɵɯ ɡɚɦɟɧɚ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ, ɜɵɡɵɜɚɟɬ ɡɧɚɱɢɬɟɥɶɧɵɟ ɢɡɦɟɧɟɧɢɹ ɜɚɠɧɟɣɲɢɯ ɯɚɪɚɤɬɟɪɢɫɬɢɤ ȻɊ: ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɚ ɟɝɨ ɩɨɝɥɨɳɟɧɢɹ, ɫɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ, ɤɢɧɟɬɢɤɭ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɚ ɢ ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɬɪɚɧɫɩɨɪɬɚ ɩɪɨɬɨɧɚ.

Ɋɢɫ. 18. Ɋɚɫɩɨɥɨɠɟɧɢɟ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ȻɊ ɜ D-ɫɩɢɪɚɥɶɧɵɯ ɬɹɠɚɯ. ɉɪɢ ɚɧɚɥɢɡɟ ɨɩɭɛɥɢɤɨɜɚɧɧɵɯ ɜ ɥɢɬɟɪɚɬɭɪɟ ɞɚɧɧɵɯ (ɫɦ. ɬɚɛɥ. 3 ɧɚ ɫ. 26 ɜ ɨɛɡɨɪɟ ɥɢɬɟɪɚɬɭɪɵ) ɫɥɟɞɭɟɬ ɭɱɢɬɵɜɚɬɶ, ɱɬɨ ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ɩɢɝɦɟɧɬɚ ɢ ɦɧɨɝɢɟ ɞɪɭɝɢɟ ɟɝɨ ɫɜɨɣɫɬɜɚ ɜ ɡɧɚɱɢɬɟɥɶɧɨɣ ɫɬɟɩɟɧɢ ɡɚɜɢɫɹɬ ɨɬ ɦɟɬɨɞɚ ɩɨɥɭɱɟɧɢɹ ɯɪɨɦɨɩɪɨɬɟɢɧɚ, ɟɝɨ ɥɢɩɢɞɧɨɝɨ ɦɢɤɪɨɨɤɪɭɠɟɧɢɹ, ɪɇ, ɬɟɦɩɟɪɚɬɭɪɵ, ɩɪɢɫɭɬɫɬɜɢɹ ɪɚɡɥɢɱɧɵɯ ɢɨɧɨɜ ɜ ɪɚɫɬɜɨɪɟ. ɉɨ ɷɬɨɣ ɩɪɢɱɢɧɟ ɩɪɟɞɫɬɚɜɥɹɟɬɫɹ ɜɨɡɦɨɠɧɵɦ ɫɪɚɜɧɢɜɚɬɶ ɫɜɨɣɫɬɜɚ ɬɨɥɶɤɨ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɨɝɨ ȻɊ ɢ ȻɊ ɫ ɩɪɢɪɨɞɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ, ɩɨɥɭɱɟɧɧɵɯ ɢ ɢɫɫɥɟɞɨɜɚɧɧɵɯ ɜ ɨɞɢɧɚɤɨɜɵɯ ɭɫɥɨɜɢɹɯ. ȼ ɬɚɛɥ. 10 ɩɪɢɜɟɞɟɧɵ ɪɟɡɭɥɶɬɚɬɵ ɚɧɚɥɢɡɚ ɢɡɦɟɧɟɧɢɹ ɫɜɨɣɫɬɜ ȻɊ, ɩɪɨɜɟɞɟɧɧɨɝɨ ɫ ɭɱɟɬɨɦ ɷɬɢɯ ɨɫɨɛɟɧɧɨɫɬɟɣ. C ɰɟɥɶɸ ɜɵɞɟɥɟɧɢɹ ɬɟɯ ɦɭɬɚɰɢɣ, ɤɨɬɨɪɵɟ ɧɚɢɛɨɥɟɟ ɫɢɥɶɧɨ ɢɡɦɟɧɹɸɬ ɫɜɨɣɫɬɜɚ ȻɊ, ɚɧɚɥɢɡ ɫɜɨɣɫɬɜ ɦɭɬɚɧɬɧɵɯ ȻɊ ɛɵɥ ɨɫɭɳɟɫɬɜɥɟɧ ɧɚ ɨɫɧɨɜɟ ɫɥɟɞɭɸɳɢɯ ɤɪɢɬɟɪɢɟɜ: 1) ɛɵɥɢ ɜɵɞɟɥɟɧɵ ɬɟ ɦɭɬɚɰɢɢ, ɤɨɬɨɪɵɟ ɢɡɦɟɧɹɸɬ Omax ɩɢɝɦɟɧɬɚ ɛɨɥɟɟ ɱɟɦ ɧɚ 10 ɧɦ. ɉɪɢ ɜɵɛɨɪɟ ɷɬɨɝɨ ɡɧɚɱɟɧɢɹ ɛɵɥɨ ɭɱɬɟɧɨ ɬɨ, ɱɬɨ ɜ ɨɛɥɚɫɬɢ 570 ɧɦ ȻɊ ɢɦɟɟɬ ɞɨɫɬɚɬɨɱɧɨ ɲɢɪɨɤɭɸ ɩɨɥɨɫɭ ɩɨɝɥɨɳɟɧɢɹ (ɫ ɩɨɥɭɲɢɪɢɧɨɣ ɨɤɨɥɨ 70 ɧɦ), ɜ ɫɜɹɡɢ ɫ ɷɬɢɦ

66 ɬɨɱɧɨɫɬɶ ɨɩɪɟɞɟɥɟɧɢɹ ɩɨɥɨɠɟɧɢɹ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ɫɨɫɬɚɜɥɹɟɬ a1-2 ɧɦ. Ʉɪɨɦɟ ɬɨɝɨ, ɫɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ, ɱɬɨ Omax ɩɢɝɦɟɧɬɚ ɢɡɦɟɧɹɟɬɫɹ ɩɨɱɬɢ ɩɪɢ ɜɫɟɯ ɡɚɦɟɧɚɯ ɚɦɢɧɨɤɢɫɥɨɬ ɜ ɪɟɡɭɥɶɬɚɬɟ ɧɟɛɨɥɶɲɢɯ ɢɡɦɟɧɟɧɢɣ, ɜɧɨɫɢɦɵɯ ɢɦɢ ɜ ɬɪɟɬɢɱɧɭɸ ɫɬɪɭɤɬɭɪɭ ɛɟɥɤɚ. ɉɨɷɬɨɦɭ ɞɥɹ ɨɬɛɨɪɚ ɞɨɫɬɚɬɨɱɧɨ ɭɡɤɨɝɨ ɪɹɞɚ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ, ɨɤɚɡɵɜɚɸɳɢɯ ɧɚɢɛɨɥɟɟ ɫɢɥɶɧɨɟ ɜɥɢɹɧɢɟ ɧɚ ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ȻɊ, ɢ ɩɪɢɧɹɬɨ ɡɧɚɱɟɧɢɟ 10 ɧɦ. 2) ɭɱɢɬɵɜɚɹ, ɱɬɨ ɩɪɢ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ WT-ȻɊ ɫɞɜɢɝ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ɫɨɫɬɚɜɥɹɟɬ 10 ɧɦ, ɞɥɹ ɚɧɚɥɢɡɚ ɛɵɥɨ ɩɪɢɧɹɬɨ ɫɱɢɬɚɬɶ, ɱɬɨ ɫɜɟɬɨɜɚɹ ɚɞɚɩɬɚɰɢɹ ɧɟ ɧɚɪɭɲɟɧɚ, ɟɫɥɢ ɩɪɨɢɫɯɨɞɢɬ ɫɞɜɢɝ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ɛɨɥɟɟ ɱɟɦ ɧɚ 7 ɧɦ. 3) ɜɵɞɟɥɟɧɵ ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɫɧɢɠɚɸɳɢɟ ɷɮɮɟɤɬɢɜɧɨɫɬɶ ɬɪɚɧɫɩɨɪɬɚ ɩɪɨɬɨɧɨɜ ɤɚɤ ɦɢɧɢɦɭɦ ɧɚ ɨɞɢɧ ɩɨɪɹɞɨɤ. 4) ɩɪɟɞɫɬɚɜɥɟɧɚ ɜɵɛɨɪɤɚ ɬɟɯ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ, ɡɚɦɟɧɚ ɤɨɬɨɪɵɯ ɢɡɦɟɧɹɟɬ ɤɢɧɟɬɢɤɭ ɨɛɪɚɡɨɜɚɧɢɹ ɢ ɪɚɫɩɚɞɚ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɛɨɥɟɟ ɱɟɦ ɜ 5 ɪɚɡ (ɬɨ ɟɫɬɶ ɟɫɥɢ ɫɪɟɞɧɟɟ ɜɪɟɦɹ ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɝɨ ɩɪɨɰɟɫɫɚ ɢɡɦɟɧɹɟɬɫɹ ɩɨ ɤɪɚɣɧɟɣ ɦɟɪɟ ɜ 5 ɪɚɡ). Ɍɚɛɥɢɰɚ 10. ɂɡɦɟɧɟɧɢɟ ɫɜɨɣɫɬɜ ȻɊ, ɜɵɡɜɚɧɧɨɟ ɬɨɱɟɱɧɵɦɢ ɦɭɬɚɰɢɹɦɢ ɜ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ. *1 – ɪɚɫɩɨɥɨɠɟɧɢɟ ɨɫɬɚɬɤɚ ȺɄ ɜ ɦɨɥɟɤɭɥɟ WT-ȻɊ, ɨɩɪɟɞɟɥɟɧɧɨɟ ɩɨ ɟɝɨ ɤɪɢɫɬɚɥɥɨɝɪɚɮɢɱɟɫɤɨɣ ɫɬɪɭɤɬɭɪɟ: ɩ - ɜ ɩɟɬɥɹɯ, ɜɵɫɬɭɩɚɸɳɢɯ ɢɡ ɦɟɦɛɪɚɧɵ, ɧ – ɧɚɩɪɚɜɥɟɧ ɧɚɪɭɠɭ ɦɨɥɟɤɭɥɵ, ɤ – ɨɤɨɥɨ ɤɨɥɶɰɚ ɪɟɬɢɧɚɥɹ, ɰ – ɨɤɨɥɨ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɪɟɬɢɧɚɥɹ, SB – ɨɤɨɥɨ ɨɫɬɚɬɤɚ Lys-216, ɫɜɹɡɚɧɧɨɝɨ ɫ ɪɟɬɢɧɚɥɟɦ. *2 – ɫɜɟɬɨɜɚɹ ɚɞɚɩɬɚɰɢɹ « + » - ɧɨɪɦɚɥɶɧɚɹ, « – » - ɨɬɫɭɬɫɬɜɭɟɬ, « r » - ɧɚɪɭɲɟɧɚ (ɜ ɫɤɨɛɤɚɯ ɭɤɚɡɚɧɨ ɧɚ ɫɤɨɥɶɤɨ ɢɡɦɟɧɹɟɬɫɹ Omaɯ ɩɪɢ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ) *3 – ɞɚɧɧɵɟ ɩɪɢɜɟɞɟɧɵ ɨɬɧɨɫɢɬɟɥɶɧɨ ȻɊ ɫ ɩɪɢɪɨɞɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ.

Ɇɭɬɚɰɢɹ

ɉɨɥɨɠɟɧɢɟ ɨɫɬɚɬɤɚ ȺɄ ɜ ɦɨɥɟɤɭɥɟ ȻɊ*1 D-ɫɩɢɪɚɥɶɧɵɣ ɬɹɠ

R7Q W10C W10F W12F T17V M20A M20E T24V D36N D38N

A A A A A A A

ɩ ɧ ɧ SB SB ɩ ɩ

ɋɜɟɬɨȻɊ Omut O , ɜɚɹ max max ɚɞɚɩɬɚɧɦ ɰɢɹ*2 0 –10 –3 0 0 0 –1 –2 –1 +2

+ + + + + + r (6) + + +

ɋɨɨɬɧɨɲɟɧɢɟ ɢɡɨɦɟɪɨɜ (LA)

+

ɇ ɬɪɚɧɫɩɨɪɬ*3

Ʉɢɧɟɬɢɤɚ Ɇɢɧɬɟɪɦɟɞɢɚɬɚ, t ɫɪmut / t ɫɪȻɊ Ɉɛɪɚɡɨɜɚɧɢɟ

Ɋɚɫɩɚɞ

0.2 0.6

2.1 1.5

0.8 91/9

87/13 94/6

0.8 0.7 1 0.7 1 1 1.1 1.1

67

Ɇɭɬɚɰɢɹ


A44G I45A T46V T47V L48A V49A V49L P50A P50G L52A A53G T55A T55V Y57N S59A G72C E74C N76C Y79C T80F R82A R82G R82H R82Q Y83F Y83N (-)Ⱥ84 (+)Ⱥ85 D85A D85C D85E D85H D85N W86F T89A T89D T89N T89V T90C T90V P91A

B B B B B B B B B B B B B B B

Ɍɚɛɥɢɰɚ 10. ɉɪɨɞɨɥɠɟɧɢɟ. Ʉɢɧɟɬɢɤɚ Ɇɋɨɨɬɧɨɋɜɟɬɨ+ ɢɧɬɟɪɦɟɞɢɚɬɚ, ɲɟɧɢɟ ɇ ȻɊ Omut ɜɚɹ max O max , t ɫɪmut / t ɫɪȻɊ ɢɡɨɦɟ- ɬɪɚɧɫɚɞɚɩɬɚ*3 ɧɦ ɪɨɜ ɩɨɪɬ ɈɛɪɚɡɨɊɚɫɰɢɹ*2 (LA) ɜɚɧɢɟ ɩɚɞ

C C

–1 –2 0 –1 –2 –2 0 –5 –5 –2 –7 –2 +7 –3 0 –4 0 +1 –26 –2 –5

C C C C C C C C C C C C C C C C C C C C

+2 0 +19 –11 –3 +16 +2 +50 +36 +52 +35 +56 –22 –6 +6 +3 –3 –10 –5 –5

ɧ

SB SB

SB ɧ ɧ ɩ ɩ ɩ ɩ

ɧ

ɰ ɰ ɰ ɰ ɰ ɰ ɰ ɧ

+ + r (6) + + – r (5) r (5) + + r (4) + + r (5) + +

91/10

0.7 1

2.9

1

71/18 89/7

1 0.6 0.5 1

20.8 1.7 0.1 0.25

0.4 1.2

76/15

1.1

0.3

1.9

0.15

0.1

1 1 0.3 1.8 3.2

1 1 0.3 3 1.9

3 3.7 0.1 107

70

0.8 2.2 6

0.9 7.6 1.1

1.5 0.1

0.9 0.75

1 75/25 1 0.9 1 0.9 0.9 0.5 0.4

+ + (H.sal) –(E.coli) + – r (3) + – + – + r (4) + + –– r (3) + –– r (4) r (5) r (5)

79/21

0.1

97/3 92/8

0 0.3 0 0 0.3 0 0 0.3 0.7 0.7

71/29 58/32 42/47 56/33 75/13

64/15

0.7 1 0.7 0.5

2 1

68

Ɇɭɬɚɰɢɹ


P91G L92C L93A L93T L93V D96A D96C D96E D96G D96N A98C D102N A103C D104N Q105C T107V L109A L109C G113L G113C G113I D115A D115E D115N G116C I117A I117C I117E M118A M118E G120C T121V G122C V124C L127C T128V K129C S132A R134Q W137C W137F W138C

C C C C C C C C C C C

D D D D D D D D D D D D D D D D D D D D D

ɧ

ɩ ɩ ɩ ɧ ɧ ɧ ɧ ɧ

ɧ

ɤ ɤ ɧ ɤ ɧ ɩ ɩ ɩ ɩ

E E E

ɤ

Ɍɚɛɥɢɰɚ 10. ɉɪɨɞɨɥɠɟɧɢɟ. Ʉɢɧɟɬɢɤɚ Ɇɋɨɨɬɧɨɋɜɟɬɨ+ ɢɧɬɟɪɦɟɞɢɚɬɚ, ɲɟɧɢɟ ɇ ȻɊ Omut ɜɚɹ max O max , t ɫɪmut / t ɫɪȻɊ ɢɡɨɦɟ- ɬɪɚɧɫɚɞɚɩɬɚɧɦ ɪɨɜ ɩɨɪɬ*3 Ɉɛɪɚɡɨ- Ɋɚɫɰɢɹ*2 (LA) ɜɚɧɢɟ ɩɚɞ –8 –3 –19 –20 –4 +1 0 +1 –1 +1 0 +2 +1 –1 0 0 –1 +2 +6 –1 0 –8 –10 –6 –1 –2 +7 –1 –77 +1 0 –3 –27 0 0 +1 +2 +2 0 –5 –19 –8

+ + – – r (6) r (5) + + +

84/14 84/16 83/6

95/5 96/4

+ + + +

0.7 0.1 0.8 1 1 0.1 0.2 0.6

0.1

0.85

3

41

2.3

69

0.1 1 1 0.9 0.9 0.8 1

15

58

0.4 10 5.4

0.2 1.5 1.8

2.2 5

27 5.8

0.7

1.2

0.8 + 0.9 + 1 0.3 0.8

r (2) –

1 + 1 + r (4) r (4)

45/51 51/44

r (6) –

76/18

+ + + r (4) + r (4)

0.3 0.9 1 1 1.1 1.1 1.1 1 0.9 1 0.4 1 0.1 0.9

69 ɉɨɥɨɠɟɧɢɟ ɨɫɬɚɬɤɚ ȺɄ ɜ ɦɨɥɟɤɭɥɟ ȻɊ*1

Ɇɭɬɚɰɢɹ

D-ɫɩɢɪɚɥɶɧɵɣ ɬɹɠ

W138F S141A S141C T142V M145A M145E T157V S158A R164Q S169C R175Q T178V W182F S183A Y185A Y185F P186A P186G P186L P186V W189F S193A E194Q T205V D212A D212E D212N S214A K216A K216G R225Q S226A R227G R227Q

E E E E E E

F F F F F F F F F F F F

ɤ ɤ ɤ ɤ ɤ ɤ ɩ ɩ ɩ ɩ

ɰ ɰ ɫɫ ɰ ɤ

ɤ ɩ

G G G G G G G G

ɰ ɰ ɰ ɧ

ɧ ɩ ɩ ɩ

Ɍɚɛɥɢɰɚ 10. ɉɪɨɞɨɥɠɟɧɢɟ. Ʉɢɧɟɬɢɤɚ Ɇɋɨɨɬɧɨɋɜɟɬɨ+ ɢɧɬɟɪɦɟɞɢɚɬɚ, ɲɟɧɢɟ ɇ ȻɊ Omut ɜɚɹ max O max , t ɫɪmut / t ɫɪȻɊ ɢɡɨɦɟ- ɬɪɚɧɫɚɞɚɩɬɚɧɦ ɪɨɜ ɩɨɪɬ*3 Ɉɛɪɚɡɨ- Ɋɚɫɰɢɹ*2 (LA) ɜɚɧɢɟ ɩɚɞ 0 –11 +17 –5 –76 –1 +1 +1 +2 –4 +1 +1 –104 0 +3 –10 +2 –79 –81 –81 –38 +2 –14 +1 –11 +14 +4 –1 +10 +15 +2 0 0 0

+ r (3) – + r (2) + + + + + + + + +

20/65 69/24

78/20

+ + –

86/14 36/64 76/24

– + +

86/14 77/23

+ ɧɟɭɫɬ. –– –– + ɧɟɭɫɬ. ɧɟɭɫɬ. + +

94/6

+

63/26 49/34

0.6 0.9 0.2 0.9 0.1 0.9 1.1 1 1 1.1 0.8 1.1 0.6 1

6.8

0.2

5.7 1.6

20 0.9

0.33 0.6 0.7 0.4 0.2 0.57 1.2 1 0.7

36/21

0.1 0.1 1.1 1

92/8

0.8 0.6

1.5

0.1

0.2 16.9

5 0.6

2.36

0.005 0.1

0.4

ɂɬɨɝɢ ɚɧɚɥɢɡɚ ɜɥɢɹɧɢɹ ɝɟɧɟɬɢɱɟɫɤɨɣ ɦɨɞɢɮɢɤɚɰɢɢ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ ɩɪɢɜɟɞɟɧɵ ɜ ɬɚɛɥɢɰɟ 11. ɋɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ, ɱɬɨ ɫɭɳɟɫɬɜɭɸɬ ɨɛɥɚɫɬɢ ȻɊ, ɡɚɦɟɧɚ ɚɦɢɧɨɤɢɫɥɨɬ ɜ ɤɨɬɨɪɵɯ ɜɵɡɵɜɚɟɬ ɧɟɡɧɚɱɢɬɟɥɶɧɵɟ ɢɡɦɟɧɟɧɢɹ ɫɜɨɣɫɬɜ ɛɟɥɤɚ. ɏɚɪɚɤɬɟɪɧɨ, ɱɬɨ ɜ ɨɫɧɨɜɧɨɦ ɨɧɢ ɪɚɫɩɨɥɨɠɟɧɵ ɩɨ ɩɟɪɢɮɟɪɢɢ ɦɨɥɟɤɭɥɵ ȻɊ (ɪɢɫ. 20ɋ).

70 ȼ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɫɭɳɟɫɬɜɭɟɬ ɜɨɡɦɨɠɧɨɫɬɶ ɞɨɫɬɚɬɨɱɧɨ ɬɨɱɧɨ ɨɩɢɫɚɬɶ ɪɚɫɩɨɥɨɠɟɧɢɟ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɜ ɦɨɥɟɤɭɥɟ ȻɊ – ɩɨ ɞɚɧɧɵɦ ɟɝɨ ɪɟɧɬɝɟɧɨɫɬɪɭɤɬɭɪɧɨɝɨ ɚɧɚɥɢɡɚ. ɋɨɜɦɟɳɟɧɢɟ ɪɟɧɬɝɟɧɨ-ɫɬɪɭɤɬɭɪɧɵɯ ɞɚɧɧɵɯ ɫ ɚɧɚɥɢɡɨɦ ɫɜɨɣɫɬɜ ɦɭɬɚɧɬɧɵɯ ȻɊ (ɩɪɢɜɟɞɟɧɧɵɯ ɜ ɬɚɛɥ.11) ɞɚɟɬ ɜɨɡɦɨɠɧɨɫɬɶ ɨɩɪɟɞɟɥɟɧɢɹ ɮɭɧɤɰɢɨɧɚɥɶɧɨ ɜɚɠɧɵɯ ɪɟɝɢɨɧɨɜ ɦɨɥɟɤɭɥɵ ȻɊ, ɡɚɦɟɧɚ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɜ ɤɨɬɨɪɵɯ ɧɚɢɛɨɥɟɟ ɫɢɥɶɧɨ ɜɥɢɹɟɬ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ (ɫɦ. ɪɢɫ. 19 ɢ 20). Ɍɚɛɥɢɰɚ 11. ȼɥɢɹɧɢɟ ɬɨɱɟɱɧɵɯ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɡɚɦɟɧ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ. Ɂɚɦɟɧɵ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɉɨɥɨɠɟɧɢɟ Omax ɩɢɝɦɟɧɬɚ W10C, Y79C, Y83F, W86F, T89V, T90C, L93A, L93T, D115E, ɝɢɩɫɨɯɪɨɦɧɵɣ ɫɞɜɢɝ M118A, G122C, W137F, S141A, M145A, W182F, Y185F, ɛɨɥɟɟ 10 ɧɦ (ɪɢɫ. 19ȼ) P186G, P186L, P186V, W189F, D212A. ɛɚɬɨɯɪɨɦɧɵɣ ɫɞɜɢɝ (-)A84, R82A, R82Q, D85C, D85A, D85E, D85H, D112E, ɛɨɥɟɟ 10 ɧɦ (ɪɢɫ. 19Ⱥ) S141C, K216A, K216G. ɇɚɪɭɲɟɧɢɟ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ ɢ ɢɡɨɦɟɪɧɨɝɨ ɫɨɫɬɚɜɚ ȻɊ: ɋɜɟɬɨɜɚɹ ɚɞɚɩɬɚɰɢɹ V49A, R82C, (-)A84, D85C, D85A, T89A, T89V, L93A, L93T, ɨɬɫɭɬɫɬɜɭɟɬ (ɪɢɫ. 19C) D115N, G122C, S141C, P186G, P186V, D212E, D212N. ɇɚɪɭɲɟɧɨ ɫɨɨɬɧɨɲɟV49A, A53G, Y57N, R82Q, D85E, D85N, T89A, T89V, T89D, ɧɢɟ ɢɡɨɦɟɪɨɜ ɜ LA- ȻɊ P91A, L93A, L93T, M118A, M118E, G122C, S141C, M145A, (ɪɢɫ. 19D) W182F, P186A, P186G, W189F, D212N, D212E, K216A. Ʉɢɧɟɬɢɤɚ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ ɛɨɥɟɟ ɱɟɦ ɜ 10: P50A, D85E, P91A, P91G, ɍɫɤɨɪɟɧɢɟ ɨɛɪɚɡɨɜɚɧɢɹ ɛɨɥɟɟ ɱɟɦ ɜ 5 ɪɚɡ - ɩɪɢ M20A, R82A, D212E. ɛɨɥɟɟ ɱɟɦ ɜ 10 ɪɚɡ: V49A, D85N, P91A, P91G, D96N, D115E, Ɂɚɦɟɞɥɟɧɢɟ ɨɛɪɚɡɨD212N. ɛɨɥɟɟ ɱɟɦ ɜ 5 ɪɚɡ: T89V, D115N, M118E, S141C, ɜɚɧɢɹ M145A. ɍɫɤɨɪɟɧɢɟ ɪɚɫɩɚɞɚ ɛɨɥɟɟ ɱɟɦ ɜ 10 ɪɚɡ: R82A, T205V, S226A, R227G. (ɪɢɫ. 20Ⱥ) ɛɨɥɟɟ ɱɟɦ ɜ 5 ɪɚɡ: D115A, S141C. ɛɨɥɟɟ ɱɟɦ ɜ 10 ɪɚɡ: D85A, D96A, D96E, D96N, M118A, Ɂɚɦɟɞɥɟɧɢɟ ɪɚɫɩɚɞɚ M145A, D227G (ɪɢɫ. 20A) ɛɨɥɟɟ ɱɟɦ ɜ 5 ɪɚɡ:T89D, M118E, D212E. ɗɮɮɟɤɬɢɜɧɨɫɬɶ ɇ+-ɬɪɚɧɫɩɨɪɬɚ + (-)A84, D85A, D85C, D85H, D85N ɇ -ɬɪɚɧɫɩɨɪɬ ɨɬɫɭɬɫɬɜɭɟɬ (ɪɢɫ. 20B) + ɇ -ɬɪɚɧɫɩɨɪɬ ɫɧɢɠɟɧ ɛɨɥɟɟ ɱɟɦ ɧɚ D96A, D96N, W137F, M145A, D212E ɩɨɪɹɞɨɤ (ɪɢɫ. 20B) Ɋɢɫ. 19 ɢ 20 ɫɨɡɞɚɧɵ ɩɪɢ ɩɨɦɨɳɢ ɩɪɨɝɪɚɦɦɵ Chem3D Ultra ɢɡ ɩɚɤɟɬɚ ChemOffice 6.0 (CambridgeSoft Com, USA) ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɮɚɣɥɚ 1cwq ɪɟɧɬɝɟɧɨɫɬɪɭɤɬɭɪɧɨɝɨ ɚɧɚɥɢɡɚ WT-ȻɊ ɜ ɨɫɧɨɜɧɨɦ ɫɨɫɬɨɹɧɢɢ ȼ (c all-E-ɢɡɨɦɟɪɨɦ ɯɪɨɦɨɮɨɪɚ) ɫ ɪɚɡɪɟɲɟɧɢɟɦ 1,55 Å, ɪɚɫɩɨɥɨɠɟɧɧɨɝɨ ɜ Brookhaven Protein Data Bank. ɇɚ ɪɢɫɭɧɤɚɯ ɢɡɨɛɪɚɠɟɧɚ ɩɨɥɢɩɟɩɬɢɞɧɚɹ ɰɟɩɶ ȻɊ (ɨɫɬɚɥɶɧɵɟ ɚɬɨɦɵ ɫɤɪɵɬɵ), ɪɚɫɩɨɥɨɠɟɧɧɚɹ ɬɚɤɢɦ ɨɛɪɚɡɨɦ, ɱɬɨ ɟɟ N-ɤɨɧɟɰ ɧɚɯɨɞɢɬɫɹ ɜɧɢɡɭ, ɚ ɩɥɨɫɤɨɫɬɶ ɦɟɦɛɪɚɧɵ ɩɟɪɩɟɧɞɢɤɭɥɹɪɧɚ ɩɥɨɫɤɨɫɬɢ ɪɢɫɭɧɤɚ. Ɉɫɬɚɬɨɤ ɪɟɬɢɧɚɥɹ (1) ɨɛɨɡɧɚɱɟɧ ɡɚɩɨɥɧɟɧɧɵɦɢ ɬɨɱɤɚɦɢ ɫɮɟɪɚɦɢ.

71 Ʉɪɨɦɟ ɩɨɥɢɩɟɩɬɢɞɧɨɣ ɰɟɩɢ ɢ ɨɫɬɚɬɤɚ ɪɟɬɢɧɚɥɹ ɧɚ ɪɢɫɭɧɤɚɯ ɨɫɬɚɜɥɟɧɵ ɛɨɤɨɜɵɟ ɪɚɞɢɤɚɥɵ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ (ɢɯ ɚɬɨɦɵ ɢɡɨɛɪɚɠɟɧɵ ɜ ɜɢɞɟ ɤɪɭɠɤɨɜ), ɪɚɫɩɨɥɨɠɟɧɢɟ ɤɨɬɨɪɵɯ ɜ ɦɨɥɟɤɭɥɟ ȻɊ ɧɟɨɛɯɨɞɢɦɨ ɨɯɚɪɚɤɬɟɪɢɡɨɜɚɬɶ. xɂɡɦɟɧɟɧɢɟ ɩɨɥɨɠɟɧɢɹ Omax ɩɢɝɦɟɧɬɚ: Ɉɫɬɚɬɤɢ ɚɦɢɧɨɤɢɫɥɨɬ, ɜɵɡɵɜɚɸɳɢɯ ɢɡɦɟɧɟɧɢɟ ɩɨɥɨɠɟɧɢɹ Omax ɛɨɥɟɟ ɱɟɦ ɧɚ 10 ɧɦ, ɪɚɫɩɨɥɨɠɟɧɵ ɜ ɰɟɩɹɯ Ⱥ, ȼ, ɋ ɢ D, ɜ ɨɫɧɨɜɧɨɦ ɜ ɨɛɥɚɫɬɢ ɯɪɨɦɨɮɨɪɧɨɣ ɩɨɥɨɫɬɢ ɪɟɬɢɧɚɥɹ (ɫɦ. ɪɢɫ.19Ⱥ, ȼ). ɂɫɤɥɸɱɟɧɢɟɦ ɹɜɥɹɸɬɫɹ ɨɫɬɚɬɤɢ Trp10, Tyr79, Trp137, ɧɚɯɨɞɹɳɢɟɫɹ ɜ ɦɟɫɬɚɯ ɩɟɪɟɯɨɞɚ D-ɫɩɢɪɚɥɶɧɵɯ ɬɹɠɟɣ ɜ ɩɟɬɥɢ ɫ ɜɧɟɤɥɟɬɨɱɧɨɣ ɫɬɨɪɨɧɵ ɫɬɨɪɨɧɵ ɦɟɦɛɪɚɧɵ, ɢ Tyr79 ɢ Asp212, ɪɚɫɩɨɥɨɠɟɧɧɵɟ ɜɧɭɬɪɢ ɦɨɥɟɤɭɥɵ ȻɊ. xɇɚɪɭɲɟɧɢɟ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ ɢ ɢɡɨɦɟɪɧɨɝɨ ɫɨɫɬɚɜɚ ȻɊ: ɋɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ, ɱɬɨ ɨɫɬɚɬɤɢ ɚɦɢɧɨɤɢɫɥɨɬ, ɜɵɡɵɜɚɸɳɢɟ ɨɬɫɭɬɫɬɜɢɟ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ ɢ ɧɚɪɭɲɟɧɢɟ ɢɡɨɦɟɪɧɨɝɨ ɫɨɫɬɚɜɚ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɨɣ ɮɨɪɦɵ ȻɊ ɪɚɫɩɨɥɚɝɚɸɬɫɹ ɨɤɨɥɨ ɭɱɚɫɬɤɚ ɫɜɹɡɵɜɚɧɢɹ ɪɟɬɢɧɚɥɹ (ɫɦ. ɪɢɫ. 19C, D). ɉɪɢɱɟɦ ɬɨɥɶɤɨ ɞɟɥɟɰɢɹ (-)A84, ɜɥɢɹɹ ɧɚ ɫɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ, ɧɟ ɜɵɡɵɜɚɟɬ ɧɚɪɭɲɟɧɢɹ ɢɡɨɦɟɪɧɨɝɨ ɫɨɫɬɚɜɚ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɨɣ ɮɨɪɦɵ ȻɊ (ɞɥɹ ȻɊD115N ɷɬɢ ɞɚɧɧɵɟ ɨɬɫɭɬɫɬɜɭɸɬ). Ʉɚɤ ɢ ɫɥɟɞɨɜɚɥɨ ɨɠɢɞɚɬɶ, ɜ ɛɨɥɶɲɢɧɫɬɜɟ ɫɥɭɱɚɟɜ ɧɚɪɭɲɟɧɢɟ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ ɫɜɹɡɚɧɨ ɫ ɢɡɦɟɧɟɧɢɟɦ ɫɨɨɬɧɨɲɟɧɢɹ ɢɡɨɦɟɪɨɜ ɜ ɬɟɦɧɨɜɨɣ ɢ ɫɜɟɬɨɜɨɣ ɮɨɪɦɚɯ ȻɊ. xɄɢɧɟɬɢɤɚ ɪɚɫɩɚɞɚ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ ɢ ɬɪɚɧɫɩɨɪɬ ɩɪɨɬɨɧɚ Ɍɪɚɧɫɩɨɪɬ ɩɪɨɬɨɧɚ ɨɬɫɭɬɫɬɜɭɟɬ ɬɨɥɶɤɨ ɩɪɢ ɡɚɦɟɧɟ ɨɫɬɚɬɤɚ Asp85 (ɢɥɢ ɨɬɫɭɬɫɬɜɢɢ 84 ɚɦɢɧɨɤɢɫɥɨɬɧɨɝɨ ɨɫɬɚɬɤɚ) ɧɚ ɧɟɡɚɪɹɠɟɧɧɵɟ ɨɫɬɚɬɤɢ, ɩɪɢ ɜɫɟɯ ɨɫɬɚɥɶɧɵɯ ɢɫɫɥɟɞɨɜɚɧɧɵɯ ɦɭɬɚɰɢɹɯ ɫɩɨɫɨɛɧɨɫɬɶ ȻɊ ɩɟɪɟɧɨɫɢɬɶ ɩɪɨɬɨɧɵ ɫɨɯɪɚɧɹɟɬɫɹ. ɋɢɥɶɧɨɟ ɫɧɢɠɟɧɢɟ ɬɪɚɧɫɩɨɪɬɚ ɩɪɨɬɨɧɚ ɩɪɢ ɬɨɱɟɱɧɨɣ ɦɭɬɚɰɢɢ ȻɊ ɹɜɥɹɟɬɫɹ ɤɪɢɬɟɪɢɟɦ ɬɨɝɨ, ɱɬɨ ɢɡɦɟɧɟɧɧɵɣ ɨɫɬɚɬɨɤ ȺɄ ɢɝɪɚɟɬ ɫɭɳɟɫɬɜɟɧɧɭɸ ɪɨɥɶ ɜ ɩɪɨɰɟɫɫɟ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ ɛɟɥɤɨɦ. Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɜ ɪɟɡɭɥɶɬɚɬɟ ɩɪɨɜɟɞɟɧɧɨɝɨ ɚɧɚɥɢɡɚ (ɫɦ. ɬɚɛɥ. 10, 11), ɦɨɠɧɨ ɜɵɞɟɥɢɬɶ ɪɹɞ ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ, ɡɚɦɟɧɚ ɤɨɬɨɪɵɯ ɜɵɡɵɜɚɟɬ ɪɟɡɤɨɟ ɡɚɦɟɞɥɟɧɢɟ ɮɨɬɨɰɢɤɥɚ, ɷɬɨ Asp85, Asp96, Met118, Met145 ɢ Asp227. ɉɪɢ ɷɬɨɦ ɨɩɢɫɚɧɧɵɟ ɜɚɪɢɚɧɬɵ ɦɭɬɚɰɢɣ ɩɨ Asp85 ɜɵɡɵɜɚɸɬ ɛɚɬɨɯɪɨɦɧɵɣ ɫɞɜɢɝ Omax ɧɚ 30-50 ɧɦ, ɩɨ Ɇet145 – ɝɢɩɫɨɯɪɨɦɧɵɣ ɫɞɜɢɝ ɞɨ 70 ɧɦ, ɚ Asp96, Met118 ɢ Asp227 ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɢɡɦɟɧɹɸɬ ɩɨɥɨɠɟɧɢɹ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ ȻɊ. ɂɡ ɩɪɢɜɟɞɟɧɧɨɝɨ ɪɹɞɚ ɩɪɨɢɡɜɨɞɧɵɯ ȻɊ ɞɥɹ ɩɨɥɭɱɟɧɢɹ ȺȻɊ ɜ ɞɚɧɧɨɣ ɪɚɛɨɬɟ ɛɵɥɚ ɜɵɛɪɚɧɚ ɬɨɱɟɱɧɚɹ ɦɭɬɚɰɢɹ D96N, ɤɨɬɨɪɚɹ ɩɨɡɜɨɥɹɟɬ ɩɨɥɭɱɢɬɶ ɩɢɝɦɟɧɬ ɫ ɡɚɦɟɞɥɟɧɧɵɦ ɮɨɬɨɰɢɤɥɨɦ ɢ ɧɟɢɡɦɟɧɧɵɦ ɩɨɥɨɠɟɧɢɟɦ ɦɚɤɫɢɦɭɦɚ ɩɨɝɥɨɳɟɧɢɹ.

C)

Ɋɢɫ. 19. Ⱥɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɨɤɚɡɵɜɚɸɳɢɟ ɡɧɚɱɢɬɟɥɶɧɨɟ ɜɥɢɹɧɢɟ ɧɚ ɩɨɥɨɠɟɧɢɟ ɦɚɤɫɢɦɭɦɚ

B)

(ɜɧɢɡɭ ɩɪɢɜɟɞɟɧɚ ɩɪɨɟɤɰɢɹ ɦɨɥɟɤɭɥɵ ȻɊ ɩɚɪɚɥɥɟɥɶɧɨ ɩɥɨɫɤɨɫɬɢ ɦɟɦɛɪɚɧɵ).

ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ; D) ɡɚɦɟɬɧɨɟ ɧɚɪɭɲɟɧɢɟ ɫɨɨɬɧɨɲɟɧɢɹ ɢɡɨɦɟɪɨɜ ɜ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɨɦ ɫɨɫɬɨɹɧɢɢ

Ⱥ) ɛɚɬɨɯɪɨɦɧɵɣ ɫɞɜɢɝ ɛɨɥɟɟ ɱɟɦ ɧɚ 10 ɧɦ; ȼ) ɝɢɩɫɨɯɪɨɦɧɵɣ ɫɞɜɢɝ ɛɨɥɟɟ ɱɟɦ ɧɚ 10 ɧɦ; C) ɨɬɫɭɬɫɬɜɢɟ

ɩɨɝɥɨɳɟɧɢɹ ȻɊ, ɫɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ ɢ ɫɨɨɬɧɨɲɟɧɢɟ ɢɡɨɦɟɪɨɜ.

A)

72

D)

B)

C)

ɨɬɫɭɬɫɬɜɢɟ (ɱɟɪɧɵɣ); ɋ) ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɦɚɥɨ ɜɥɢɹɸɳɢɟ ɧɚ ɫɜɨɣɫɬɜɚ ȻɊ.

ɡɚɦɟɞɥɟɧɢɟ ɛɨɥɟɟ ɱɟɦ ɜ 5 ɪɚɡ; ȼ) ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɜɵɡɵɜɚɸɳɢɟ 10-ɤɪɚɬɧɨɟ ɭɦɟɧɶɲɟɧɢɟ ɇ-ɬɪɚɧɫɩɨɪɬɚ (ɫɟɪɵɣ) ɢɥɢ ɟɝɨ

Ɋɢɫ. 20. A) ɚɦɢɧɨɤɢɫɥɨɬɧɵɟ ɨɫɬɚɬɤɢ, ɜɥɢɹɸɳɢɟ ɧɚ ɤɢɧɟɬɢɤɭ ɪɚɫɩɚɞɚ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɚ ȻɊ, ɫɟɪɵɦ – ɭɫɤɨɪɟɧɢɟ, ɱɟɪɧɵɦ –

A)

73

74

6. Ⱥɧɚɥɨɝɢ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ Ⱦɥɹ ɩɨɥɭɱɟɧɢɹ ȺȻɊ, ɤɪɨɦɟ ɫɢɧɬɟɡɚ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ, ɛɵɥɨ ɧɟɨɛɯɨɞɢɦɨ ɪɟɲɢɬɶ ɧɟɫɤɨɥɶɤɨ ɞɨɩɨɥɧɢɬɟɥɶɧɵɯ ɡɚɞɚɱ: ɨɩɬɢɦɢɡɢɪɨɜɚɬɶ ɭɫɥɨɜɢɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɜɵɛɪɚɧɧɵɯ ɲɬɚɦɦɨɜ H. salinarum ɫ ɰɟɥɶɸ ɩɨɥɭɱɟɧɢɹ ɦɚɤɫɢɦɚɥɶɧɨɝɨ ɜɵɯɨɞɚ ȻɊ, ɚ ɬɚɤ ɠɟ ɨɬɪɚɛɨɬɚɬɶ ɦɟɬɨɞɢɤɭ ɩɨɥɭɱɟɧɢɹ ɚɩɨɦɟɦɛɪɚɧ ɢ ɪɟɝɟɧɟɪɚɰɢɢ ɯɪɨɦɨɩɪɨɬɟɢɧɨɜ ɢ ɨɯɚɪɚɤɬɟɪɢɡɨɜɚɬɶ ɤɚɱɟɫɬɜɨ ɩɨɥɭɱɟɧɧɵɯ ɩɪɟɩɚɪɚɬɨɜ.

6.1. Ʉɭɥɶɬɢɜɢɪɨɜɚɧɢɟ H. salinarum Ɉɞɧɨɣ ɢɡ ɡɚɞɚɱ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɵ ɛɵɥ ɩɨɢɫɤ ɭɫɥɨɜɢɣ ɞɥɹ ɧɚɢɛɨɥɟɟ ɷɮɮɟɤɬɢɜɧɨɝɨ ɩɨɥɭɱɟɧɢɹ ɩɪɟɩɚɪɚɬɨɜ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ, ɢɫɩɨɥɶɡɭɟɦɵɯ ɞɥɹ ɞɚɥɶɧɟɣɲɟɝɨ ɢɫɫɥɟɞɨɜɚɧɢɹ. ɏɨɬɹ ɦɟɬɨɞɵ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ H. salinarum ɢ ɩɨɥɭɱɟɧɢɹ ɉɆ ɞɨɫɬɚɬɨɱɧɨ ɩɨɞɪɨɛɧɨ ɨɩɢɫɚɧɵ ɜ ɥɢɬɟɪɚɬɭɪɟ, ɧɚɦ ɩɪɢɲɥɨɫɶ «ɚɞɚɩɬɢɪɨɜɚɬɶ» ɢɯ ɤ ɪɟɚɥɶɧɵɦ ɷɤɫɩɟɪɢɦɟɧɬɚɥɶɧɵɦ ɭɫɥɨɜɢɹɦ. Ɋɟɡɭɥɶɬɚɬɨɦ ɷɬɨɣ ɱɚɫɬɢ ɷɤɫɩɟɪɢɦɟɧɬɨɜ ɹɜɥɹɸɬɫɹ ɧɟɫɤɨɥɶɤɨ ɩɪɨɦɟɠɭɬɨɱɧɵɯ ɧɚɛɥɸɞɟɧɢɣ ɢ ɜɵɜɨɞɨɜ, ɩɨɡɜɨɥɢɜɲɢɯ ɜ ɢɬɨɝɟ ɩɨɥɭɱɢɬɶ ɩɪɟɩɚɪɚɬɵ ɉɆ ɫ ɞɨɫɬɚɬɨɱɧɨ ɜɵɫɨɤɢɦ ɜɵɯɨɞɨɦ (ɞɨ 50 ɦɝ/ɥ ɤɭɥɶɬɭɪɵ). ɇɚ ɨɫɧɨɜɚɧɢɢ ɦɧɨɝɨɱɢɫɥɟɧɧɵɯ ɢɫɫɥɟɞɨɜɚɧɢɣ ɝɚɥɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ ɢɡɜɟɫɬɧɨ, ɱɬɨ ɛɨɥɶɲɢɧɫɬɜɨ ɢɯ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɫɩɨɫɨɛɧɵ ɪɚɫɬɢ ɧɚ ɩɢɬɚɬɟɥɶɧɵɯ ɫɪɟɞɚɯ ɫ ɛɟɥɤɨɜɵɦɢ ɝɢɞɪɨɥɢɡɚɬɚɦɢ ɢɥɢ ɚɦɢɧɨɤɢɫɥɨɬɚɦɢ ɜ ɤɚɱɟɫɬɜɟ ɟɞɢɧɫɬɜɟɧɧɨɝɨ ɢɫɬɨɱɧɢɤɚ ɭɝɥɟɪɨɞɚ [29, 186], ɚ ɜɚɪɶɢɪɨɜɚɧɢɟ ɫɨɥɟɜɨɝɨ ɫɨɫɬɚɜɚ ɩɨɥɧɨɰɟɧɧɵɯ ɫɪɟɞ ɩɪɢɜɨɞɢɥɨ ɤ ɢɡɦɟɧɟɧɢɸ ɭɪɨɜɧɹ ɧɚɤɨɩɥɟɧɢɹ ɛɢɨɦɚɫɫɵ ɢ ɤ ɢɡɦɟɧɟɧɢɹɦ ɮɨɪɦɵ ɤɥɟɬɨɤ ɝɚɥɨɛɚɤɬɟɪɢɣ [187]. ȼ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɡɚɞɚɱ ɷɤɫɩɟɪɢɦɟɧɬɚɬɨɪɨɜ, ɢɫɩɨɥɶɡɭɸɬɫɹ ɪɨɫɬɨɜɵɟ ɫɪɟɞɵ, ɪɚɡɧɨɨɛɪɚɡɧɵɟ ɩɨ ɫɨɫɬɚɜɭ ɢ ɫɨɞɟɪɠɚɧɢɸ ɤɨɦɩɨɧɟɧɬɨɜ, ɧɨ ɫ ɞɜɭɦɹ ɨɛɹɡɚɬɟɥɶɧɵɦɢ ɫɨɫɬɚɜɥɹɸɳɢɦɢ - ɯɥɨɪɢɞɨɦ ɧɚɬɪɢɹ ɜ ɜɵɫɨɤɨɣ ɤɨɧɰɟɧɬɪɚɰɢɢ ɢ ɬɟɦ ɢɥɢ ɢɧɵɦ ɢɫɬɨɱɧɢɤɨɦ ɚɦɢɧɨɤɢɫɥɨɬ. ɇɚɢɛɨɥɟɟ ɨɛɵɱɧɵɦ ɢɫɬɨɱɧɢɤɨɦ ɚɦɢɧɨɤɢɫɥɨɬ ɩɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɝɚɥɨɛɚɤɬɟɪɢɣ ɹɜɥɹɸɬɫɹ ɜɫɟɜɨɡɦɨɠɧɵɟ ɝɢɞɪɨɥɢɡɚɬɵ ɛɟɥɤɨɜ (ɢɥɢ ɛɢɨɦɚɫɫɵ) ɦɢɤɪɨɛɧɨɝɨ, ɪɚɫɬɢɬɟɥɶɧɨɝɨ ɢɥɢ ɠɢɜɨɬɧɨɝɨ ɩɪɨɢɫɯɨɠɞɟɧɢɹ. ɂɞɟɚɥɶɧɵɦ ɜɚɪɢɚɧɬɨɦ ɝɢɞɪɨɥɢɡɚɬɚ ɦɨɠɟɬ ɫɱɢɬɚɬɶɫɹ ɩɪɟɩɚɪɚɬ ɤɚɡɚɦɢɧɨɤɢɫɥɨɬ - ɩɪɨɞɭɤɬ ɩɨɥɧɨɝɨ ɤɢɫɥɨɬɧɨɝɨ ɝɢɞɪɨɥɢɡɚ ɤɚɡɟɢɧɚ. ȼ ɪɹɞɟ ɫɥɭɱɚɟɜ ɢɫɩɨɥɶɡɭɸɬ ɛɨɥɟɟ ɞɟɲɟɜɵɟ ɝɢɞɪɨɥɢɡɚɬɵ, ɡɚɱɚɫɬɭɸ ɫ ɧɟɤɨɧɬɪɨɥɢɪɭɟɦɵɦ ɫɨɫɬɚɜɨɦ, ɩɨɫɤɨɥɶɤɭ ɩɪɢɦɟɧɟɧɢɟ ɫɪɟɞ, ɩɪɢɝɨɬɨɜɥɟɧɧɵɯ ɧɚ ɬɚɤɢɯ ɪɨɫɬɨɜɵɯ ɫɭɛɫɬɪɚɬɚɯ, ɨɩɪɚɜɞɚɧɨ ɷɤɨɧɨɦɢɱɟɫɤɢ. ɂɡ ɥɢɬɟɪɚɬɭɪɧɵɯ ɞɚɧɧɵɯ ɩɨ ɢɫɫɥɟɞɨɜɚɧɢɸ ɜɨɡɦɨɠɧɨɫɬɢ ɫɬɢɦɭɥɢɪɨɜɚɧɢɹ ɢ ɢɧɝɢɛɢɪɨɜɚɧɢɹ ɛɢɨɫɢɧɬɟɡɚ ȻɊ ɢɡɜɟɫɬɧɨ [3, 37], ɱɬɨ ɢɧɬɟɧɫɢɜɧɨɟ ɨɫɜɟɳɟɧɢɟ ɩɪɢ ɧɟɞɨɫɬɚɬɨɱɧɨɣ ɚɷɪɚɰɢɢ ɤɭɥɶɬɭɪɵ ɩɪɢɜɨɞɢɬ ɤ ɩɨɜɵɲɟɧɢɸ ɫɨɞɟɪɠɚɧɢɹ ȻɊ ɜ ɤɥɟɬɤɚɯ, ɬɨɝɞɚ ɤɚɤ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɟ ɜ ɬɟɦɧɨɬɟ ɩɪɢ ɫɢɥɶɧɨɣ ɚɷɪɚɰɢɢ ɩɪɢɜɨɞɢɬ ɤ ɬɨɪɦɨɠɟɧɢɸ ɢɥɢ ɩɪɟɤɪɚɳɟɧɢɸ ɨɛɪɚɡɨɜɚɧɢɹ ȻɊ. ɉɨ ɷɬɨɣ ɩɪɢɱɢɧɟ ɦɵ ɩɪɨɜɨɞɢɥɢ ɩɨɞɛɨɪ ɭɫɥɨɜɢɣ

75 (ɨɫɜɟɳɟɧɢɟ ɢ ɚɷɪɚɰɢɹ) ɫ ɰɟɥɶɸ ɭɜɟɥɢɱɟɧɢɹ ɜɵɯɨɞɚ ȻɊ ɫ ɟɞɢɧɢɰɵ ɨɛɴɟɦɚ ɤɭɥɶɬɭɪɚɥɶɧɨɣ ɠɢɞɤɨɫɬɢ (Ʉɀ), ɬɚɤ ɤɚɤ ɭɫɥɨɜɢɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɜ ɮɟɪɦɟɧɬɟɪɟ ɫɭɳɟɫɬɜɟɧɧɨ ɨɬɥɢɱɚɸɬɫɹ ɨɬ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɜ ɤɨɥɛɚɯ ɢ ɩɪɟɞɫɬɚɜɥɹɸɬ ɫɨɛɨɣ ɫɚɦɨɫɬɨɹɬɟɥɶɧɵɣ ɨɛɴɟɤɬ ɢɫɫɥɟɞɨɜɚɧɢɹ. ɉɪɢ ɜɵɩɨɥɧɟɧɢɢ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɵ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɟ H. salinarum ɨɫɭɳɟɫɬɜɥɹɥɢ ɜ 4 ɥ ɤɨɦɩɥɟɤɫɧɨɣ ɫɪɟɞɵ, ɫɨɞɟɪɠɚɳɟɣ ɧɚ 1 ɥ 250 ɝ NaCl, 20 ɝ MgSO47 ɇ2Ɉ, 3 ɝ ɰɢɬɪɚɬɚ ɧɚɬɪɢɹ, 2 ɝ KCl, 2 ɝ ɞɪɨɠɠɟɜɨɝɨ ɷɤɫɬɪɚɤɬɚ, 5 ɝ ɩɟɩɬɨɧɚ, 1 ɦɥ ɝɥɢɰɟɪɢɧɚ [188, 189]. ɉɪɨɰɟɫɫ ɩɪɨɜɨɞɢɥɢ ɜ ɩɟɪɢɨɞɢɱɟɫɤɨɦ ɪɟɠɢɦɟ ɜ ɩɪɨɡɪɚɱɧɨɣ ɩɥɨɫɤɨɣ ɤɸɜɟɬɟ ɨɛɴɟɦɨɦ 6 ɥ, ɫɧɚɛɠɟɧɧɨɣ ɨɫɜɟɳɟɧɢɟɦ 4-ɦɹ ɥɚɦɩɚɦɢ ɞɧɟɜɧɨɝɨ ɫɜɟɬɚ ɅȾ-20. Ⱥɷɪɚɰɢɸ ɢ ɩɟɪɟɦɟɲɢɜɚɧɢɟ ɤɭɥɶɬɭɪɵ ɨɛɟɫɩɟɱɢɜɚɥɢ ɩɨɞɚɱɟɣ ɜɨɡɞɭɯɚ ɜ ɧɢɠɧɸɸ ɱɚɫɬɶ ɤɸɜɟɬɵ (ɫɦ. ɪɢɫ. 28 ɧɚ ɫ. 93 ɷɤɫɩɟɪɢɦɟɧɬɚɥɶɧɨɣ ɱɚɫɬɢ). Ⱦɥɹ ɩɨɞɞɟɪɠɚɧɢɹ ɬɟɦɩɟɪɚɬɭɪɧɨɝɨ ɪɟɠɢɦɚ ɤɸɜɟɬɭ ɩɨɦɟɳɚɥɢ ɜ ɬɟɪɦɨɫɬɚɬɢɪɭɟɦɭɸ ɤɨɦɧɚɬɭ. ȼɪɟɦɹ, ɧɟɨɛɯɨɞɢɦɨɟ ɞɥɹ ɩɪɨɜɟɞɟɧɢɹ ɩɟɪɢɨɞɢɱɟɫɤɨɣ ɮɟɪɦɟɧɬɚɰɢɢ ɜ ɧɚɲɟɦ ɫɥɭɱɚɟ ɫɨɫɬɚɜɥɹɥɨ ɨɤɨɥɨ ɞɜɭɯ ɧɟɞɟɥɶ, ɩɨɷɬɨɦɭ ɨɞɧɨ ɢɡ ɨɫɧɨɜɧɵɯ ɬɪɟɛɨɜɚɧɢɣ, ɤɨɬɨɪɨɟ ɩɪɟɞɴɹɜɥɹɥɨɫɶ ɤ ɮɟɪɦɟɧɬɟɪɭ – ɫɨɯɪɚɧɟɧɢɟ ɫɬɟɪɢɥɶɧɨɫɬɢ ɩɪɢ ɢɧɬɟɧɫɢɜɧɨɣ ɚɷɪɚɰɢɢ ɩɢɬɚɬɟɥɶɧɨɣ ɫɪɟɞɵ ɜɨ ɜɪɟɦɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ. Ⱦɥɹ ɷɬɨɝɨ ɧɚ ɥɢɧɢɢ ɜɯɨɞɚ ɜɨɡɞɭɯɚ ɭɫɬɚɧɨɜɥɟɧɵ ɫɬɟɪɢɥɶɧɵɟ ɮɢɥɶɬɪɵ, ɩɪɟɞɨɬɜɪɚɳɚɸɳɢɟ ɩɨɩɚɞɚɧɢɟ ɜ ɧɟɝɨ ɩɨɫɬɨɪɨɧɧɢɯ ɦɢɤɪɨɨɪɝɚɧɢɡɦɨɜ. ɑɬɨɛɵ ɨɯɚɪɚɤɬɟɪɢɡɨɜɚɬɶ ɪɨɫɬ ɛɢɨɦɚɫɫɵ ɝɚɥɨɛɚɤɬɟɪɢɣ ɛɵɥɢ ɩɨɫɬɪɨɟɧɵ ɤɪɢɜɵɟ ɪɨɫɬɚ. ɂɡɜɟɫɬɧɨ, ɱɬɨ ɨɩɬɢɱɟɫɤɚɹ ɩɥɨɬɧɨɫɬɶ (OD) – ɨɞɢɧ ɢɡ ɨɫɧɨɜɧɵɯ ɩɚɪɚɦɟɬɪɨɜ, ɯɚɪɚɤɬɟɪɢɡɭɸɳɢɯ ɪɨɫɬ ɤɭɥɶɬɭɪɵ. Ⱦɥɹ ɩɨɫɬɪɨɟɧɢɹ ɤɪɢɜɨɣ ɪɨɫɬɚ ɢɡ ɤɸɜɟɬɵ ɩɟɪɢɨɞɢɱɟɫɤɢ ɨɬɛɢɪɚɥɢ ɩɪɨɛɵ ɞɥɹ ɨɩɪɟɞɟɥɟɧɢɹ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɤɭɥɶɬɭɪɵ, ɢɡɦɟɧɟɧɢɟ ɤɨɬɨɪɨɣ ɜ ɯɨɞɟ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɩɪɟɞɫɬɚɜɥɟɧɨ ɧɚ ɪɢɫ. 21. Ɋɚɧɟɟ ɛɵɥɨ ɨɬɦɟɱɟɧɨ [37], ɱɬɨ ɭ ɧɟɤɨɬɨɪɵɯ ɲɬɚɦɦɨɜ H. salinarum ɜ ɫɬɚɰɢɨɧɚɪɧɨɣ ɮɚɡɟ ɪɨɫɬɚ ɩɪɨɢɫɯɨɞɢɬ ɞɨɩɨɥɧɢɬɟɥɶɧɨɟ ɧɚɤɨɩɥɟɧɢɟ ȻɊ. ȼ ɫɜɹɡɢ ɫ ɷɬɢɦ ɧɚɪɹɞɭ ɫ ɨɩɪɟɞɟɥɟɧɢɟɦ ɧɚɤɨɩɥɟɧɢɹ ɛɢɨɦɚɫɫɵ ɩɨ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ, ɦɵ ɨɩɪɟɞɟɥɹɥɢ ɞɢɧɚɦɢɤɭ ɧɚɤɨɩɥɟɧɢɹ ȻɊ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɢɱɟɫɤɢɦ ɦɟɬɨɞɨɦ, ɩɪɟɞɥɨɠɟɧɧɵɦ Oesterhelt D. et al. [188], ɤɨɬɨɪɵɣ ɦɨɠɟɬ ɩɪɢɦɟɧɹɬɶɫɹ ɞɥɹ ɲɬɚɦɦɨɜ ɝɚɥɨɛɚɤɬɟɪɢɣ, ɫɨɞɟɪɠɚɳɢɯ ɩɨɧɢɠɟɧɧɨɟ ɤɨɥɢɱɟɫɬɜɨ ɤɚɪɨɬɢɧɨɢɞɨɜ. ɉɨɫɤɨɥɶɤɭ ɜ ɛɚɤɬɟɪɢɹɯ ɲɬɚɦɦɚ JW5 ȻɊ ɧɟ ɨɛɪɚɡɭɟɬɫɹ, ɚ ɩɪɨɢɫɯɨɞɢɬ ɧɚɤɨɩɥɟɧɢɟ ȻɈ, ɞɥɹ ɟɝɨ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɢɱɟɫɤɨɝɨ ɨɩɪɟɞɟɥɟɧɢɹ ɛɵɥɨ ɧɟɨɛɯɨɞɢɦɨ ɩɪɟɞɜɚɪɢɬɟɥɶɧɨ ɞɨɛɚɜɥɹɬɶ ɤ ɢɫɫɥɟɞɭɟɦɨɣ ɩɪɨɛɟ ɢɡɛɵɬɨɤ all-E-ɪɟɬɢɧɚɥɹ (1), ɱɬɨ ɩɪɢɜɨɞɢɥɨ ɤ ɨɛɪɚɡɨɜɚɧɢɸ ȻɊ.

76 1

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0,8

0,8

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0,4

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4

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8

10

Ɋɢɫ. 21. ɋɪɚɜɧɟɧɢɟ ɤɪɢɜɵɯ ɪɨɫɬɚ H. salinarum ɲɬɚɦɦɨɜ ET1001 (37qɋ), JW5 (37qɋ), D96N (34qɋ). Ɉɛɨɡɧɚɱɟɧɢɹ: i- ɨɩɬɢɱɟɫɤɚɹ ɩɥɨɬɧɨɫɬɶ ɤɭɥɶɬɭɪɵ ɩɪɢ 650 ɧɦ, Ƒ ɧɚɤɨɩɥɟɧɢɟ ȻɊ ɩɨ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ5 ɫɭɩɟɪɧɚɬɚɧɬɚ ɥɢɡɢɪɨɜɚɧɧɵɯ ɤɥɟɬɨɤ ɩɪɢ 570 ɧɦ. ȼ ɪɟɡɭɥɶɬɚɬɟ ɩɪɨɜɟɞɟɧɧɵɯ ɷɤɫɩɟɪɢɦɟɧɬɨɜ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɭ ɜɫɟɯ ɬɪɟɯ ɲɬɚɦɦɨɜ H. salinarum (ET1001, JW5 ɢ D96N) ɩɪɨɢɫɯɨɞɢɬ ɞɨɩɨɥɧɢɬɟɥɶɧɵɣ ɛɢɨɫɢɧɬɟɡ ȻɊ (ɢɥɢ ȻɈ) ɧɚ ɫɬɚɰɢɨɧɚɪɧɨɣ ɮɚɡɟ ɪɨɫɬɚ (ɫɦ. ɪɢɫ. 21), ɩɪɢɱɟɦ ɜ ɧɚɢɛɨɥɟɟ ɡɧɚɱɢɬɟɥɶɧɨɣ ɫɬɟɩɟɧɢ – ɭ ɲɬɚɦɦɚ JW5. ȼ ɬɨ ɠɟ ɜɪɟɦɹ, ɩɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɛɨɥɟɟ 10 ɞɧɟɣ ɩɪɢ 37qɋ ɩɪɨɢɫɯɨɞɢɬ ɫɧɢɠɟɧɢɟ ɜɵɯɨɞɚ ȻɊ. Ȼɵɥɨ ɨɬɦɟɱɟɧɨ, ɱɬɨ ɞɜɭɤɪɚɬɧɨɟ ɫɧɢɠɟɧɢɟ ɨɫɜɟɳɟɧɧɨɫɬɢ (ɞɜɭɦɹ ɥɚɦɩɚɦɢ ɜɦɟɫɬɨ ɱɟɬɵɪɟɯ) ɜ ɩɪɨɰɟɫɫɟ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɲɬɚɦɦɚ ȿɌ1001 ɜɵɡɵɜɚɸɬ ɩɪɨɩɨɪɰɢɨɧɚɥɶɧɨɟ ɫɧɢɠɟɧɢɟ ɜɵɯɨɞɚ ȻɊ ɩɪɢ ɫɨɯɪɚɧɟɧɢɢ ɭɪɨɜɧɹ ɧɚɤɨɩɥɟɧɢɹ ɛɢɨɦɚɫɫɵ. ɉɨ ɷɬɨɣ ɩɪɢɱɢɧɟ ɩɪɢ ɩɪɨɜɟɞɟɧɢɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɞɥɹ ɨɫɜɟɳɟɧɢɹ ɢɫɩɨɥɶɡɨɜɚɥɢ ɦɚɤɫɢɦɚɥɶɧɨɟ ɤɨɥɢɱɟɫɬɜɨ ɥɚɦɩ, ɤɨɬɨɪɨɟ ɩɨɡɜɨɥɹɥɢ ɪɚɡɦɟɪɵ ɤɸɜɟɬɵ ɢ ɤɨɧɫɬɪɭɤɰɢɹ ɨɫɜɟɬɢɬɟɥɶɧɵɯ ɩɪɢɛɨɪɨɜ. ɉɪɨɜɟɞɟɧɧɵɟ ɷɤɫɩɟɪɢɦɟɧɬɵ ɩɨɤɚɡɚɥɢ, ɱɬɨ ɧɚ ɭɪɨɜɟɧɶ ɧɚɤɨɩɥɟɧɢɹ ɛɢɨɦɚɫɫɵ ɜɥɢɹɟɬ ɤɚɤ ɨɛɴɟɦ ɩɨɫɟɜɧɨɝɨ ɦɚɬɟɪɢɚɥɚ, ɬɚɤ ɢ ɟɝɨ ɜɨɡɪɚɫɬ. ȼ ɨɛɳɟɦ ɫɥɭɱɚɟ, ɪɟɤɨɦɟɧɞɭɟɦɵɣ ɨɛɴɟɦ ɩɨɫɟɜɧɨɝɨ ɦɚɬɟɪɢɚɥɚ ɫɨɫɬɚɜɥɹɟɬ 5% ɨɬ ɨɛɴɟɦɚ ɡɚɫɟɜɚɟɦɨɣ ɫɪɟɞɵ, ɨɞɧɚɤɨ ɦɵ ɨɛɧɚɪɭɠɢɥɢ, ɱɬɨ ɟɝɨ ɭɜɟɥɢɱɟɧɢɟ ɞɨ 10% ɜɵɡɵɜɚɟɬ ɩɨɜɵɲɟɧɢɟ ɜɵɯɨɞɚ ɛɢɨɦɚɫɫɵ ɜ 1,3 ɪɚɡɚ ɢ ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɟ ɩɨɜɵɲɟɧɢɟ ɜɵɯɨɞɚ ȻɊ. Ɉɩɬɢɦɚɥɶɧɵɦ ɹɜɥɹɟɬɫɹ ɜɨɡɪɚɫɬ ɩɨɫɟɜɧɨɝɨ ɦɚɬɟɪɢɚɥɚ – ɜ ɤɨɧɰɟ ɷɤɫɩɨɧɟɧɰɢɚɥɶɧɨɣ ɮɚɡɵ ɪɨɫɬɚ (ɜ ɧɚɲɟɦ ɫɥɭɱɚɟ ɩɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɜ ɤɨɥɛɚɯ ɩɪɢ 34qɋ ɨɧ ɫɨɫɬɚɜɥɹɟɬ 5 ɞɧɟɣ). Ⱦɥɹ ɲɬɚɦɦɚ JW5 ɨɬɦɟɱɟɧɨ, ɱɬɨ ɞɨɛɚɜɥɟɧɢɟ ɪɟɬɢɧɚɥɹ (1) ɢɥɢ ɟɝɨ ɚɧɚɥɨɝɚ ɜ ɩɪɨɰɟɫɫɟ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɩɪɢ ɨɫɜɟɳɟɧɢɢ ɩɨɜɵɲɚɟɬ ɜɵɯɨɞ ȻɊ ɧɚ 10%, ɚ ɩɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɛɟɡ ɞɨɛɚɜɥɟɧɢɹ ɪɟɬɢɧɨɢɞɨɜ ɜ ɤɥɟɬɤɚɯ ɩɪɢɫɭɬɫɬɜɭɟɬ ɨɤɨɥɨ 30% ɫɨɞɟɪɠɚɳɢɯ ɛɨɥɶɲɨɟ ɤɨɥɢɱɟɫɬɜɨ ȻɈ ɦɟɦɛɪɚɧ ɩɨɧɢɠɟɧɧɨɣ ɩɥɨɬɧɨɫɬɢ, ɤɨɬɨɪɵɟ ɧɟ ɨɫɚɠɞɚɸɬɫɹ ɩɪɢ ɫɬɚɧɞɚɪɬɧɨɣ ɩɪɨɰɟɞɭɪɟ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɹ. ɉɪɢ ɞɨɛɚɜɥɟɧɢɢ ɪɟɬɢɧɚɥɹ (1) ɨɧɢ ɫɚɦɨɩɪɨɢɡɜɨɥɶɧɨ ɨɛɪɚɡɭɸɬ ɩɭɪɩɭɪɧɵɟ ɦɟɦɛɪɚɧɵ ɢ ɥɟɝɤɨ ɜɵɞɟɥɹɸɬɫɹ 5

Ⱦɥɹ ɲɬɚɦɦɚ JW5 ɨɩɪɟɞɟɥɹɥɢ ɩɨɫɥɟ ɢɧɤɭɛɚɰɢɢ ɨɛɪɚɡɰɚ ɫ ɢɡɛɵɬɤɨɦ ɪɟɬɢɧɚɥɹ (1)

77 ɜ ɫɬɚɧɞɚɪɬɧɵɯ ɭɫɥɨɜɢɹɯ, ɩɪɢɱɟɦ ɩɨɥɭɱɚɟɦɵɟ ɩɪɟɩɚɪɚɬɵ ɩɨ ɫɩɟɤɬɪɚɥɶɧɵɦ ɫɜɨɣɫɬɜɚɦ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɨɬɥɢɱɚɸɬɫɹ ɨɬ ɚɧɚɥɨɝɢɱɧɵɯ ɩɪɟɩɚɪɚɬɨɜ, ɩɨɥɭɱɟɧɧɵɯ ɢɡ ɜɵɞɟɥɟɧɧɵɯ ɩɨ ɫɬɚɧɞɚɪɬɧɨɣ ɦɟɬɨɞɢɤɟ «ɛɟɥɵɯ» ɦɟɦɛɪɚɧ. Ɍɚɤ ɱɬɨ, ɜ ɞɚɧɧɨɦ ɫɥɭɱɚɟ ɫɭɳɟɫɬɜɭɟɬ ɞɨɩɨɥɧɢɬɟɥɶɧɚɹ ɜɨɡɦɨɠɧɨɫɬɶ ɩɨɜɵɲɟɧɢɹ ɜɵɯɨɞɚ ȻɊ, ɟɫɥɢ ɪɟɝɟɧɟɪɚɰɢɸ ɯɪɨɦɨɩɪɨɬɟɢɧɚ ɨɫɭɳɟɫɬɜɥɹɬɶ ɩɟɪɟɞ ɨɫɚɠɞɟɧɢɟɦ ɞɨɛɚɜɥɟɧɢɟɦ ɤ ɫɭɫɩɟɧɡɢɢ ɦɟɦɛɪɚɧ ɢɡ ɲɬɚɦɦɚ JW5 ɧɟɨɛɯɨɞɢɦɨɝɨ ɤɨɥɢɱɟɫɬɜɚ ɪɟɬɢɧɚɥɹ (1) ɢɥɢ ɟɝɨ ɚɧɚɥɨɝɚ. ȼ ɪɟɡɭɥɶɬɚɬɟ ɩɪɨɜɟɞɟɧɧɵɯ ɷɤɫɩɟɪɢɦɟɧɬɨɜ ɛɵɥɢ ɜɵɛɪɚɧɵ ɨɛɳɢɟ ɞɥɹ ɜɫɟɯ ɲɬɚɦɦɨɜ ɝɚɥɨɛɚɤɬɟɪɢɣ ɭɫɥɨɜɢɹ: ɨɛɴɟɦ ɢɧɨɤɭɥɹɬɚ (ɜɨɡɪɚɫɬ – ɤɨɧɟɰ ɷɤɫɩɨɧɟɧɰɢɚɥɶɧɨɣ ɮɚɡɵ ɪɨɫɬɚ) ɫɨɫɬɚɜɥɹɟɬ 10% ɨɛɴɟɦɚ Ʉɀ, ɨɫɜɟɳɟɧɢɟ 4-ɦɹ ɥɚɦɩɚɦɢ ɞɧɟɜɧɨɝɨ ɫɜɟɬɚ (ɩɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɲɬɚɦɦɚ JW5 ɛɟɡ ɞɨɛɚɜɥɟɧɢɹ ɪɟɬɢɧɨɢɞɨɜ ɨɫɜɟɳɟɧɢɟ ɧɟ ɬɪɟɛɭɟɬɫɹ) Ɍɚɛɥɢɰɚ 12. ɍɫɥɨɜɢɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ H. salinarum. ȼɵɯɨɞ ɫ 1 ɥ

ȼɪɟɦɹ,

Ⱥɷɪɚɰɢɹ

ɞɧ

(ɥ ɜɨɡɞ)/(ɥ ɫɪɟɞɵ)/ɦɢɧ

ɛɢɨɦɚɫɫɚ, ɝ

ȻɊ, ɦɝ

37

8

0.45

5

45

JW5

37

10

0.45

3,2

13 (15*)

D96N

34

10

0.45

5,5

47

ɒɬɚɦɦ

t, qɋ

ET1001

* ɩɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɞɨɛɚɜɥɹɥɢ 3,4-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɶ (2).

6.2. ɉɨɥɭɱɟɧɢɟ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ Ⱦɥɹ ɩɨɥɭɱɟɧɢɹ ȺȻɊ ɢɡ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ ɲɬɚɦɦɨɜ ET1001 ɢ D96N ɛɵɥɨ ɧɟɨɛɯɨɞɢɦɨ ɩɪɟɞɜɚɪɢɬɟɥɶɧɨ ɩɪɢɝɨɬɨɜɢɬɶ ɚɩɨɦɟɦɛɪɚɧɵ ɩɪɢ ɩɨɦɨɳɢ ɫɜɟɬɨɡɚɜɢɫɢɦɨɝɨ ɝɢɞɪɨɤɫɢɥɚɦɢɧɨɥɢɡɚ, ɚ ɡɚɬɟɦ ɨɫɭɳɟɫɬɜɢɬɶ ɪɟɤɨɧɫɬɪɭɤɰɢɸ ɯɪɨɦɨɩɪɨɬɟɢɧɨɜ [190]. Ɉɛɟɫɰɜɟɱɢɜɚɧɢɟ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ ɨɫɭɳɟɫɬɜɥɹɥɢ ɜ ɫɨɨɬɜɟɬɫɬɜɢɢ ɫɨ ɫɬɚɧɞɚɪɬɧɨɣ ɦɟɬɨɞɢɤɨɣ [190]: ɫɭɫɩɟɧɡɢɸ ɦɟɦɛɪɚɧ ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ ɛɟɥɤɚ ɨɤɨɥɨ 0,5 ɦɝ/ɦɥ ɜ 1Ɇ ɪɚɫɬɜɨɪɟ ɝɢɞɪɨɤɫɢɥɚɦɢɧɚ, ɪɇ 7.0, ɩɪɢ ɬɟɦɩɟɪɚɬɭɪɟ 7-12qɋ ɨɫɜɟɳɚɥɢ ɫɜɟɬɨɦ 2-ɯ ɝɚɥɨɝɟɧɨɜɵɯ ɥɚɦɩ ɦɨɳɧɨɫɬɶɸ ɩɨ 500 ȼɬ, ɩɪɨɩɭɳɟɧɧɵɦ ɱɟɪɟɡ ɬɟɩɥɨɜɨɣ ɮɢɥɶɬɪ (0,2% ɜɨɞɧɵɣ ɪɚɫɬɜɨɪ ɫɭɥɶɮɚɬɚ ɦɟɞɢ, ɜ ɤɪɭɝɥɨɞɨɧɧɨɣ ɤɨɥɛɟ ɨɛɴɟɦɨɦ 1 ɥ), ɤɨɬɨɪɵɣ ɨɞɧɨɜɪɟɦɟɧɧɨ ɨɛɟɫɩɟɱɢɜɚɟɬ ɮɨɤɭɫɢɪɨɜɤɭ ɥɭɱɚ ɫɜɟɬɚ ɧɚ ɹɱɟɣɤɟ, ɫɨɞɟɪɠɚɳɟɣ ɫɭɫɩɟɧɡɢɸ ɦɟɦɛɪɚɧ. ɉɪɨɯɨɠɞɟɧɢɟ ɩɪɨɰɟɫɫɚ ɤɨɧɬɪɨɥɢɪɨɜɚɥɢ ɩɨ ɭɦɟɧɶɲɟɧɢɸ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɩɪɢ 563 ɧɦ. ȼ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɤɨɧɰɟɧɬɪɚɰɢɢ ɛɟɥɤɚ ɜɪɟɦɹ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ ɫɨɫɬɚɜɥɹɥɨ ɨɬ 17 ɞɨ 26 ɱ. ɇɟɫɦɨɬɪɹ ɧɚ ɬɨ, ɱɬɨ ɭ ȻɊ-D96N ɮɨɬɨɰɢɤɥ ɡɧɚɱɢɬɟɥɶɧɨ ɡɚɦɟɞɥɟɧ, ɜɪɟɦɹ ɟɝɨ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɨɬɥɢɱɚɟɬɫɹ ɨɬ ȻɊ ɫ ɩɪɢɪɨɞɧɨɣ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ.

78 Ʉɨɧɰɟɧɬɪɚɰɢɸ «ɚɤɬɢɜɧɨɝɨ» ȻɈ ɜ ɫɨɫɬɚɜɟ ɚɩɨɦɟɦɛɪɚɧ ɨɩɪɟɞɟɥɹɥɢ ɩɨ ɩɨɝɥɨɳɟɧɢɸ ɩɪɢ 568 ɧɦ ɩɨɫɥɟ ɪɟɝɟɧɟɪɚɰɢɢ ɞɟɣɫɬɜɢɟɦ ɢɡɛɵɬɤɚ all-E-ɪɟɬɢɧɚɥɹ (1). Ⱥ

ȼ

A 0,8 0,7

Ⱥɩɨɦɟɦɛɪɚɧɵ Ɋɟɝɟɧɟɪɢɪɨɜɚɧɧɵɣ ȻɊ ɂɫɯɨɞɧɵɣ ȻɊ

0,6 0,5

0,4

'A563

0,3 0,2

0,4 0,3 0,2

0,1

0,1 0,0 300

400

500

600

ɧɦ 700

ɦɢɧ

0,0 0

10

20

30

Ɋɢɫ. 22. Ɋɟɤɨɧɫɬɪɭɤɰɢɹ ȻɊ. ɋɩɟɤɬɪɵ ɧɨɪɦɚɥɢɡɨɜɚɧɵ ɩɨ ɛɟɥɤɨɜɨɣ ɩɨɥɨɫɟ ɩɨɝɥɨɳɟɧɢɹ ɩɪɟɩɚɪɚɬɨɜ ɦɟɦɛɪɚɧ ɞɨ ɞɨɛɚɜɥɟɧɢɹ ɪɟɬɢɧɚɥɹ. Ⱥ) ɋɩɟɤɬɪɵ ɩɪɟɩɚɪɚɬɨɜ, ȼ) Ʉɪɢɜɵɟ ɜɫɬɪɚɢɜɚɧɢɹ all-E-ɪɟɬɢɧɚɥɹ (1) ɜ ɚɩɨɦɟɦɛɪɚɧɵ D96N, ɜɪɟɦɹ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ ɩɪɢ ɩɨɥɭɱɟɧɢɢ ɚɩɨɦɟɦɛɪɚɧ ɫɨɫɬɚɜɥɹɟɬ -i- 17 ɱ, -- 21 ɱ, -Ÿ- 26 ɱ. ɉɨɤɚɡɚɧɨ, ɱɬɨ ɩɪɢɝɨɬɨɜɥɟɧɧɵɟ ɜ ɞɚɧɧɵɯ ɭɫɥɨɜɢɹɯ ɩɪɟɩɚɪɚɬɵ ɚɩɨɦɟɦɛɪɚɧ ɨɛɥɚɞɚɥɢ ɞɨɫɬɚɬɨɱɧɨ ɜɨɫɩɪɨɢɡɜɨɞɢɦɵɦɢ ɫɜɨɣɫɬɜɚɦɢ. Ɍɚɤ, ɫɬɟɩɟɧɶ ɜɫɬɪɚɢɜɚɧɢɹ ɪɟɬɢɧɚɥɹ (1) ɛɵɥɚ ɩɪɚɤɬɢɱɟɫɤɢ ɨɞɢɧɚɤɨɜɚ ɞɥɹ ɚɩɨɦɟɦɛɪɚɧ ɜ ɧɟɡɚɜɢɫɢɦɵɯ ɷɤɫɩɟɪɢɦɟɧɬɚɯ, ɨɞɧɚɤɨ ɫɚɦ ɩɪɨɰɟɫɫ ɩɪɨɢɫɯɨɞɢɥ ɦɟɞɥɟɧɧɟɟ, ɟɫɥɢ ɭɜɟɥɢɱɢɜɚɥɨɫɶ ɜɪɟɦɹ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ ɉɆ (ɫɦ. ɪɢɫ. 22). ȺȻɊ ɢ ɤɨɧɬɪɨɥɶɧɵɟ ɨɛɪɚɡɰɵ ȻɊ ɩɨɥɭɱɚɥɢ ɞɨɛɚɜɥɟɧɢɟɦ ɤ ɫɭɫɩɟɧɡɢɢ ɚɩɨɦɟɦɛɪɚɧ ɜ ɫɬɚɧɞɚɪɬɧɨɦ ɛɭɮɟɪɟ (100ɦɆ NaCl, 5 ɦɆ MES, ɪɇ 6.0) ɫɬɟɯɢɨɦɟɬɪɢɱɟɫɤɢɯ ɤɨɥɢɱɟɫɬɜ all-E-ɪɟɬɢɧɚɥɹ (1) ɢ ɟɝɨ ɚɧɚɥɨɝɨɜ. Ⱦɥɹ ɷɤɫɩɟɪɢɦɟɧɬɨɜ ɢɫɩɨɥɶɡɨɜɚɧɵ all-E-ɢɡɨɦɟɪɵ 3,4ɞɢɞɟɝɢɞɪɨ- (2), 4-ɨɤɫɨ- (3), ɮɟɧɢɥ- (4) ɢ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨ-ɪɟɬɢɧɚɥɹ (5). ɋɩɟɤɬɪɚɥɶɧɵɟ ɫɜɨɣɫɬɜɚ ɩɪɟɩɚɪɚɬɨɜ ȺȻɊ ɩɪɢɜɟɞɟɧɵ ɜ ɬɚɛɥɢɰɟ 13. Ɍɚɤ ɠɟ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɢɫɩɨɥɶɡɨɜɚɧɧɚɹ ɩɪɨɰɟɞɭɪɚ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ – ɪɟɤɨɧɫɬɪɭɤɰɢɢ ɫɥɚɛɨ ɜɥɢɹɟɬ ɧɚ ɤɢɧɟɬɢɤɭ ɨɛɪɚɡɨɜɚɧɢɹ ɢ ɪɚɫɩɚɞɚ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ (ɫɦ. ɪɢɫ. 26 ɧɚ ɫ. 83), ɞɥɹ ɷɬɢɯ ɰɟɥɟɣ ɩɪɨɜɨɞɢɥɢ ɪɟɝɟɧɟɪɚɰɢɸ all-E-ɪɟɬɢɧɚɥɟɦ (1). ɋɥɟɞɨɜɚɬɟɥɶɧɨ, ɩɨɥɭɱɚɟɦɵɟ ɩɪɟɩɚɪɚɬɵ ɚɩɨɦɟɦɛɪɚɧ ɩɪɢɝɨɞɧɵ ɞɥɹ ɢɫɫɥɟɞɨɜɚɧɢɹ ɤɢɧɟɬɢɤɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ȺȻɊ. ɋɤɨɪɨɫɬɢ ɜɫɬɪɚɢɜɚɧɢɹ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɜ ɛɚɤɬɟɪɢɨɨɩɫɢɧ ɫ ɩɪɢɪɨɞɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ ɢ ɦɭɬɚɧɬɧɵɣ D96N ɨɱɟɧɶ ɛɥɢɡɤɢ, ɤɚɤ ɢ ɦɚɤɫɢɦɭɦɵ ɩɨɝɥɨɳɟɧɢɹ ɩɨɥɭɱɟɧɧɵɯ ɩɢɝɦɟɧɬɨɜ (ɫɦ. ɬɚɛɥ. 13). ȼɫɬɪɚɢɜɚɧɢɟ ɜɫɟɯ ɢɫɫɥɟɞɨɜɚɧɧɵɯ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɜ ȻɈ ɩɪɨɢɫɯɨɞɢɬ ɞɨɫɬɚɬɨɱɧɨ ɛɵɫɬɪɨ, ɛɨɥɟɟ 90% ȺȻɊ ɪɟɝɟɧɟɪɢɪɭɟɬɫɹ ɜ ɬɟɱɟɧɢɟ ɱɚɫɚ.

79 ɉɪɢ ɩɨɥɭɱɟɧɢɢ ȺȻɊ ɢɡ ɲɬɚɦɦɚ JW5 ɢɫɩɨɥɶɡɨɜɚɧɨ 2 ɩɨɞɯɨɞɚ: ɞɨɛɚɜɥɟɧɢɟ ɪɟɬɢɧɨɢɞɨɜ ɤ ɪɚɫɬɭɳɟɣ ɤɭɥɶɬɭɪɟ ɤɥɟɬɨɤ ɢɥɢ ɤ ɩɪɟɩɚɪɚɬɭ «ɛɟɥɵɯ» ɦɟɦɛɪɚɧ. ɉɨɤɚɡɚɧɨ, ɱɬɨ ɜ ɩɟɪɜɨɦ ɫɥɭɱɚɟ (ɞɥɹ ɪɟɬɢɧɚɥɹ (1) ɢɥɢ 3,4-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɹ (4)) ɦɚɤɫɢɦɭɦɵ ɩɨɥɭɱɚɟɦɵɯ ɩɢɝɦɟɧɬɨɜ ɛɥɢɡɤɢ ɤ ɦɚɤɫɢɦɭɦɚɦ ɫɨɨɬɜɟɬɫɬɜɭɸɳɢɯ ɚɧɚɥɨɝɨɜ ɢɡ ɲɬɚɦɦɨɜ ET1001 ɢ D96N, ɚ ɜɨ ɜɬɨɪɨɦ (ɚɧɚɥɨɝɢ ɪɟɬɢɧɚɥɹ (3) ɢ (5))– ɫ ɧɟɤɨɬɨɪɵɦ ɫɞɜɢɝɨɦ ɜ ɤɨɪɨɬɤɨɜɨɥɧɨɜɭɸ ɨɛɥɚɫɬɶ (ɫɦ. ɬɚɛɥ. 13). ȼ ɨɛɳɟɦ, ɫɨɞɟɪɠɚɧɢɟ «ɚɤɬɢɜɧɨɝɨ» ɛɟɥɤɚ ɜ «ɛɟɥɵɯ» ɦɟɦɛɪɚɧɚɯ ɲɬɚɦɦɚ JW5 ɦɟɧɶɲɟ, ɱɟɦ ɜ ɚɩɨɦɟɦɛɪɚɧɚɯ ɲɬɚɦɦɨɜ ET1001 ɢ D96N, ɨɞɧɚɤɨ ɜɨɡɦɨɠɧɨɫɬɶ ɩɨɥɭɱɟɧɢɹ ȺȻɊ, ɢɫɤɥɸɱɢɜ ɫɬɚɞɢɸ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ, ɹɜɥɹɟɬɫɹ ɜɚɠɧɵɦ ɩɪɟɢɦɭɳɟɫɬɜɨɦ ɞɥɹ ɩɨɥɭɱɟɧɢɹ ɩɪɟɩɚɪɚɬɢɜɧɵɯ ɤɨɥɢɱɟɫɬɜ ȺȻɊ. Ɍɚɛɥɢɰɚ 13. ɋɩɟɤɬɪɚɥɶɧɵɟ ɫɜɨɣɫɬɜɚ ȺȻɊ. ɒɬɚɦɦɢɫɬɨɱɧɢɤ ȻɈ

ɋɬɪɭɤɬɭɪɚ ɯɪɨɦɨɮɨɪɚ (1)

CHO

(2)

CHO

CHO

(3) O

(4)

O

CHO

(5)

JW5 ET1001 D96N JW5 ET1001 D96N JW5 ET1001 D96N

Ⱦɢɮɮɟɪɟɧɰɢɚɥɶɧɵɣ Omax, ɧɦ DA LA 560 566 593 589 597 493 527 496 525 -

Omax, ɧɦ* DA 558 559 560 593 589 580 490 526 527

LA 565 567 569 600 596 592 483 508 503

D96N

497

497

508

508

JW5

442

-

440

-

O

445 (ɩɥ) * ɞɥɹ ɩɪɟɩɚɪɚɬɚ ȺȻɊ, ɩɟɪɟɨɫɚɠɞɟɧɧɨɝɨ ɩɨɫɥɟ ɪɟɝɟɧɟɪɚɰɢɢ 4 ɪɚɡɚ ɢɡ ɜɨɞɵ.

O

D96N

441

-

6.3. ɋɬɚɛɢɥɶɧɨɫɬɶ ɩɪɟɩɚɪɚɬɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ ɩɪɢ ɞɥɢɬɟɥɶɧɨɦ ɯɪɚɧɟɧɢɢ Ⱦɥɹ ɬɟɯɧɢɱɟɫɤɨɝɨ ɩɪɢɦɟɧɟɧɢɹ ȺȻɊ ɜɚɠɧɨ ɨɰɟɧɢɬɶ ɢɯ ɤɚɱɟɫɬɜɨ ɢ ɫɬɚɛɢɥɶɧɨɫɬɶ ɩɪɢ ɯɪɚɧɟɧɢɢ ɜ ɬɟɱɟɧɢɟ ɞɥɢɬɟɥɶɧɨɝɨ ɜɪɟɦɟɧɢ. ɋ ɷɬɨɣ ɰɟɥɶɸ ɩɪɟɩɚɪɚɬɵ ȺȻɊ ɛɵɥɢ ɢɫɫɥɟɞɨɜɚɧɵ ɦɟɬɨɞɨɦ SDS-ɷɥɟɤɬɪɨɮɨɪɟɡɚ ɜ 15% ɩɨɥɢɚɤɪɢɥɚɦɢɞɧɨɦ ɝɟɥɟ ɱɟɪɟɡ ɪɚɡɥɢɱɧɵɟ ɩɪɨɦɟɠɭɬɤɢ ɜɪɟɦɟɧɢ ɩɨɫɥɟ ɢɯ ɩɨɥɭɱɟɧɢɹ (ɫɦ. ɪɢɫ. 23). ɉɪɟɩɚɪɚɬɵ ɉɆ ɢ ȺȻɊ ɯɪɚɧɢɥɢ ɩɪɢ 4qɋ ɜ ɜɢɞɟ ɫɭɫɩɟɧɡɢɢ ɜ ɛɢɞɢɫɬɢɥɢɪɨɜɚɧɧɨɣ ɜɨɞɟ ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ ɛɟɥɤɚ 5-10 ɦɝ/ɦɥ.

80 ɂɡɜɟɫɬɧɨ, ɱɬɨ ȻɊ ɨɛɥɚɞɚɟɬ ɩɨɜɵɲɟɧɧɨɣ ɫɬɚɛɢɥɶɧɨɫɬɶɸ ɤ ɞɟɣɫɬɜɢɸ ɩɪɨɬɟɨɥɢɬɢɱɟɫɤɢɯ ɮɟɪɦɟɧɬɨɜ, ɩɨɫɤɨɥɶɤɭ ɨɫɧɨɜɧɚɹ ɱɚɫɬɶ ɛɟɥɤɚ ɧɚɯɨɞɢɬɫɹ ɜ ɦɟɦɛɪɚɧɟ ɢ ɟɝɨ ɦɨɥɟɤɭɥɵ ɨɪɝɚɧɢɡɨɜɚɧɵ ɜ ɪɟɝɭɥɹɪɧɭɸ ɝɟɤɫɚɝɨɧɚɥɶɧɭɸ ɫɬɪɭɤɬɭɪɭ, ɫɨɫɬɨɹɳɭɸ ɢɡ ɬɪɢɦɟɪɨɜ ȻɊ. «ȼɵɫɬɭɩɚɸɳɢɟ» ɢɡ ɦɟɦɛɪɚɧɵ ɮɪɚɝɦɟɧɬɵ ȻɊ (ɫɦ. ɪɢɫ. 18 ɧɚ ɫ. 65) ɦɨɝɭɬ ɛɵɬɶ ɝɢɞɪɨɥɢɡɨɜɚɧɵ ɩɪɨɬɟɚɡɚɦɢ, ɩɪɢɱɟɦ ɩɨɞ ɞɟɣɫɬɜɢɟɦ ɛɨɥɶɲɢɧɫɬɜɚ ɢɫɫɥɟɞɨɜɚɧɧɵɯ ɩɪɨɬɟɚɡ (ɧɚɩɪɢɦɟɪ, ɤɚɪɛɨɤɫɢɩɟɩɬɢɞɚɡɵ A, ɩɪɨɧɚɡɵ ɢɥɢ ɩɚɩɚɢɧɚ) ɩɪɨɢɫɯɨɞɢɬ ɨɬɳɟɩɥɟɧɢɟ ɧɟ ɛɨɥɟɟ 17 ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɫ ɋ-ɤɨɧɰɚ [137]. Ɋɚɧɟɟ ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɷɬɨ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɨɤɚɡɵɜɚɟɬ ɜɥɢɹɧɢɹ ɧɚ ɤɜɚɧɬɨɜɵɣ ɜɵɯɨɞ ɢ ɤɢɧɟɬɢɤɭ ɪɚɫɩɚɞɚ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ [40]. ɉɪɨɜɟɞɟɧɧɨɟ ɢɫɫɥɟɞɨɜɚɧɢɟ ɩɨɤɚɡɚɥɨ, ɱɬɨ ɩɪɚɤɬɢɱɟɫɤɢ ɜɫɟ ɩɪɟɩɚɪɚɬɵ ȻɊ ɫɨɞɟɪɠɚɬ ɟɞɢɧɫɬɜɟɧɧɵɣ ɛɟɥɨɤ, ɩɪɢɱɟɦ ɩɪɢ ɯɪɚɧɟɧɢɢ ɜ ɬɟɱɟɧɢɟ 12 ɦɟɫɹɰɟɜ ɟɝɨ ɤɚɱɟɫɬɜɨ (ɡɚ ɢɫɤɥɸɱɟɧɢɟɦ ȻɊ ɢɡ ɲɬɚɦɦɚ JW5) ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɢɡɦɟɧɹɟɬɫɹ. Ɍɨ, ɱɬɨ ɩɪɢ ɢɫɫɥɟɞɨɜɚɧɢɢ ɦɟɬɨɞɨɦ ɷɥɟɤɬɪɨɮɨɪɟɡɚ ɜɢɞɢɦɚɹ ɦɨɥɟɤɭɥɹɪɧɚɹ ɦɚɫɫɚ ȻɊ ɧɟɫɤɨɥɶɤɨ ɦɟɧɶɲɟ ɥɢɬɟɪɚɬɭɪɧɵɯ ɞɚɧɧɵɯ ɦɨɠɧɨ ɨɛɴɹɫɧɢɬɶ ɧɟɩɨɥɧɨɣ ɞɟɧɚɬɭɪɚɰɢɟɣ ɛɟɥɤɚ ɩɨɞ ɞɟɣɫɬɜɢɟɦ SDS, ɩɨɫɤɨɥɶɤɭ ɩɨɥɧɭɸ ɞɟɧɚɬɭɪɚɰɢɸ ȻɊ ɦɨɠɧɨ ɞɨɫɬɢɱɶ ɬɨɥɶɤɨ ɜ ɠɟɫɬɤɢɯ ɭɫɥɨɜɢɹɯ ɞɟɣɫɬɜɢɟɦ ɦɭɪɚɜɶɢɧɨɣ ɤɢɫɥɨɬɵ ɜ ɫɨɱɟɬɚɧɢɢ ɫ SDS [191]. ȻɊ D96N

ET1001

1

2

3

Ⱥɧɚɥɨɝɢ ȻɊ

4

5

ET1001

6

7

8

9

D96N

10 11 12 13 14

66 ɤȾɚ 45 ɤȾɚ 36 ɤȾɚ 29 ɤȾɚ 24 ɤȾɚ 20 ɤȾɚ

Ɋɢɫ. 23. ɗɥɟɤɬɪɨɮɨɪɟɡ ɩɪɟɩɚɪɚɬɨɜ ȻɊ. ɂɫɯɨɞɧɵɟ ɩɪɟɩɚɪɚɬɵ ȻɊ: 1, 2, 3ɜɵɞɟɥɟɧɵ ɢɡ ɲɬɚɦɦɚ ȿɌ1001, ɯɪɚɧɢɥɢɫɶ 10 ɥɟɬ, 1 ɝɨɞ ɢ 1 ɦɟɫ. ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ; 4, 5, 6 – ɢɡ ɦɭɬɚɧɬɧɨɝɨ ɲɬɚɦɦɚ D96N, ɯɪɚɧɢɥɢɫɶ 1 ɝɨɞ, 3 ɦɟɫ ɢ 1 ɦɟɫ ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ; 7- JW5 ɪɟɝɟɧɟɪɢɪɨɜɚɧɧɵɣ ɪɟɬɢɧɚɥɟɦ (1), ɯɪɚɧɢɥɫɹ 1 ɦɟɫ. ȺȻɊ: 8, 9 – ɢɡ ɲɬɚɦɦɚ ȿɌ1001, ɪɟɤɨɧɫɬɪɭɢɪɨɜɚɧɧɵɟ ɪɟɬɢɧɚɥɟɦ (1) ɢ 4-ɨɤɫɨɪɟɬɢɧɚɥɟɦ (3); 10, 11, 12, 13 - ɢɡ ɦɭɬɚɧɬɧɨɝɨ ɲɬɚɦɦɚ D96N, ɪɟɝɟɧɟɪɢɪɨɜɚɧɧɵɟ ɪɟɬɢɧɚɥɟɦ (1), 4-ɨɤɫɨ- (3), ɮɟɧɢɥ- (4) ɢ 3,4ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɟɦ (2). 14 – ɫɬɚɧɞɚɪɬɵ ɦɨɥɟɤɭɥɹɪɧɵɯ ɜɟɫɨɜ.

81 Ɉɬɦɟɱɟɧɨ ɧɟɛɨɥɶɲɨɟ ɭɜɟɥɢɱɟɧɢɟ ɷɥɟɤɬɪɨɮɨɪɟɬɢɱɟɫɤɨɣ ɩɨɞɜɢɠɧɨɫɬɢ ɭ ɩɪɟɩɚɪɚɬɨɜ ȻɊ, ɯɪɚɧɢɜɲɢɯɫɹ ɛɨɥɟɟ ɨɞɧɨɝɨ ɦɟɫɹɰɚ (ɫɦ. ɪɢɫ. 23), ɱɬɨ, ɤɚɤ ɢɡɜɟɫɬɧɨ ɢɡ ɥɢɬɟɪɚɬɭɪɧɵɯ ɞɚɧɧɵɯ [40], ɫɜɹɡɚɧɨ ɫ ɨɬɳɟɩɥɟɧɢɟɦ ɧɟ ɛɨɥɟɟ 17 ɚɦɢɧɨɤɢɫɥɨɬɧɵɯ ɨɫɬɚɬɤɨɜ ɫ ɋ-ɤɨɧɰɚ. Ʉɪɨɦɟ ɬɨɝɨ, ɜ ɩɪɟɩɚɪɚɬɟ ȻɊ, ɩɨɥɭɱɟɧɧɨɦ ɜ ɧɚɲɟɣ ɥɚɛɨɪɚɬɨɪɢɢ ɪɚɧɟɟ, ɢ ɯɪɚɧɢɜɲɟɦɫɹ ɩɪɢ 4qɋ 10 ɥɟɬ ɨɬɦɟɱɟɧɚ ɱɚɫɬɢɱɧɚɹ ɚɝɪɟɝɚɰɢɹ ɛɟɥɤɚ. Ɍɟɦ ɧɟ ɦɟɧɟɟ ɩɪɢ ɷɬɨɦ ɫɨɯɪɚɧɹɟɬɫɹ ɜɨɡɦɨɠɧɨɫɬɶ ɩɨɥɭɱɟɧɢɹ ɨɩɬɢɱɟɫɤɢ ɢ ɷɥɟɤɬɪɨɮɨɪɟɬɢɱɟɫɤɢ ɨɞɧɨɪɨɞɧɨɝɨ ɩɪɟɩɚɪɚɬɚ ɩɨɫɥɟ ɨɛɪɚɛɨɬɤɢ ɭɥɶɬɪɚɡɭɤɨɦ. ɋɪɨɤɢ ɯɪɚɧɟɧɢɹ ɩɪɟɩɚɪɚɬɨɜ ȺȻɊ ɩɟɪɟɞ ɩɪɨɜɟɞɟɧɢɟɦ ɢɯ ɷɥɟɤɬɪɨɮɨɪɟɬɢɱɟɫɤɨɝɨ ɢɫɫɥɟɞɨɜɚɧɢɹ ɫɨɫɬɚɜɥɹɥɢ 3-6 ɦɟɫɹɰɟɜ. Ɇɨɠɧɨ ɤɨɧɫɬɚɬɢɪɨɜɚɬɶ, ɱɬɨ ȺȻɊ ɬɚɤ ɠɟ ɞɨɫɬɚɬɨɱɧɨ ɭɫɬɨɣɱɢɜɵ ɤ ɞɟɣɫɬɜɢɸ ɩɪɨɬɟɚɡ, ɯɨɬɹ ɢ ɨɛɥɚɞɚɸɬ ɧɟɫɤɨɥɶɤɨ ɦɟɧɶɲɟɣ ɫɬɚɛɢɥɶɧɨɫɬɶɸ, ɱɟɦ ȻɊ, ɫɨɞɟɪɠɚɳɢɣ ɩɪɢɪɨɞɧɵɣ ɯɪɨɦɨɮɨɪ. Ɍɚɤ, ɩɨɫɥɟ ɯɪɚɧɟɧɢɹ ɜ ɬɟɱɟɧɢɟ 5 ɦɟɫɹɰɟɜ, ɨɬɦɟɱɟɧɨ ɧɟɡɧɚɱɢɬɟɥɶɧɨɟ ɩɨɜɪɟɠɞɟɧɢɟ ɛɟɥɤɨɜɨɣ ɱɚɫɬɢ ɭ ɧɟɤɨɬɨɪɵɯ ȺȻɊ (ɧɚɩɪɢɦɟɪ, 4-ɨɤɫɨ-ȻɊD96N). ɉɪɢ ɞɥɢɬɟɥɶɧɨɦ ɯɪɚɧɟɧɢɢ ɨɬɦɟɱɟɧɚ ɱɚɫɬɢɱɧɚɹ ɚɝɪɟɝɚɰɢɹ ɛɟɥɤɚ ɢ ɫɟɞɢɦɟɧɬɚɰɢɹ ɦɟɦɛɪɚɧɧɵɯ ɱɚɫɬɢɰ ɢɡ ɫɭɫɩɟɧɡɢɢ. Ʉɪɨɦɟ ɬɨɝɨ, ɩɪɢ ɯɪɚɧɟɧɢɢ ȻɊ ɜ ɜɢɞɟ ɤɨɧɰɟɧɬɪɢɪɨɜɚɧɧɨɣ ɜɨɞɧɨɣ ɫɭɫɩɟɧɡɢɢ ɜ ɬɟɱɟɧɢɟ 1 ɝɨɞɚ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɢɡɦɟɧɹɸɬɫɹ ɢ ɤɢɧɟɬɢɱɟɫɤɢɟ ɩɚɪɚɦɟɬɪɵ ɟɝɨ ɮɨɬɨɰɢɤɥɚ, ɱɬɨ ɛɵɥɨ ɢɫɫɥɟɞɨɜɚɧɨ ɧɚ ɩɪɢɦɟɪɟ ɲɬɚɦɦɨɜ ET1001 ɢ D96N (ɫɦ. ɪɢɫ. 24). ȼ)

'A, ɨɬɧ. ɟɞ.

'A, ɨɬɧ. ɟɞ.

Ⱥ)

Ɋɢɫ. 24. ȼɨɫɩɪɨɢɡɜɨɞɢɦɨɫɬɶ ɤɢɧɟɬɢɤɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɩɪɢ ɯɪɚɧɟɧɢɢ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ. Ⱥ) ɲɬɚɦɦ ȿɌ1001, ȼ) ɲɬɚɦɦ D96N. ɏɪɚɧɟɧɢɟ 1 – 1 ɦɟɫɹɰ, 2 – 10 ɦɟɫɹɰɟɜ.

6.4. Ɉɫɨɛɟɧɧɨɫɬɢ ɮɨɬɨɰɢɤɥɚ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ6 Ʉɢɧɟɬɢɤɢ ɮɨɬɨɰɢɤɥɨɜ ɩɪɟɩɚɪɚɬɨɜ ȻɊ ɢ ɟɝɨ ɚɧɚɥɨɝɨɜ ɛɵɥɢ ɢɫɫɥɟɞɨɜɚɧɵ ɜ ɫɭɫɩɟɧɡɢɹɯ ɜ ɫɬɚɧɞɚɪɬɧɨɦ ɛɭɮɟɪɟ (100 ɦɆ NaCl, 5 ɦɆ MES, ɪɇ 6) ɩɪɢ 21qC ɧɚ 6

Ⱦɚɧɧɚɹ ɱɚɫɬɶ ɪɚɛɨɬɵ ɩɪɨɜɨɞɢɥɚɫɶ ɩɨɞ ɪɭɤɨɜɨɞɫɬɜɨɦ ɫ.ɧ.ɫ., ɤ.ɛ.ɧ. ɏɢɬɪɢɧɨɣ Ʌ.ȼ. ɧɚ ɭɫɬɚɧɨɜɤɟ ɢɦɩɭɥɶɫɧɨɣ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɢɢ ɜ ɥɚɛɨɪɚɬɨɪɢɢ ɮɨɬɨɯɢɦɢɢ ɛɢɨɦɟɦɛɪɚɧ ɇɂɂ Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɨɣ ɛɢɨɥɨɝɢɢ ɢɦ. Ⱥ.ɇ. Ȼɟɥɨɡɟɪɫɤɨɝɨ ɆȽɍ.

82 ɭɫɬɚɧɨɜɤɟ ɢɦɩɭɥɶɫɧɨɣ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɢɢ ɜ ɥɚɛɨɪɚɬɨɪɢɢ ɮɨɬɨɯɢɦɢɢ ɛɢɨɦɟɦɛɪɚɧ ɇɂɂ Ɏɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɨɣ ɛɢɨɥɨɝɢɢ ɢɦ. Ⱥ.ɇ. Ȼɟɥɨɡɟɪɫɤɨɝɨ ɆȽɍ. ɉɪɢɧɰɢɩ ɪɚɛɨɬɵ ɩɪɢɛɨɪɚ: ɢɡɦɟɪɹɸɳɢɣ ɥɭɱ ɨɬ ɤɜɚɪɰɟɜɨɣ ɝɚɥɨɝɟɧɨɜɨɣ ɥɚɦɩɵ ɄȽɆ (9ȼ, 75 ȼɬ) ɩɪɨɯɨɞɢɥ ɱɟɪɟɡ ɪɟɲɟɬɨɱɧɵɣ ɦɨɧɨɯɪɨɦɚɬɨɪ HL1 (ɮɢɪɦɵ Jobin-Yvon) ɫ ɲɢɪɢɧɨɣ ɳɟɥɢ 2-8 ɦɦ, ɬɟɪɦɨɫɬɚɬɢɪɭɟɦɭɸ ɤɸɜɟɬɭ (ɞɥɢɧɚ ɨɩɬɢɱɟɫɤɨɝɨ ɩɭɬɢ 10 ɦɦ) ɫ ɨɛɪɚɡɰɨɦ, ɜɬɨɪɨɣ ɦɨɧɨɯɪɨɦɚɬɨɪ ɢ ɩɨɩɚɞɚɥ ɧɚ ɮɨɬɨɭɦɧɨɠɢɬɟɥɶ Ɏɗɍ-128. ɗɥɟɤɬɪɢɱɟɫɤɢɣ ɫɢɝɧɚɥ ɫ Ɏɗɍ ɩɨɞɚɜɚɥɫɹ ɧɚ ɨɩɟɪɚɰɢɨɧɧɵɣ ɭɫɢɥɢɬɟɥɶ Burr Brown 3554ȼɆ, ɚ ɫ ɭɫɢɥɢɬɟɥɹ – ɧɚ ɜɫɬɪɚɢɜɚɟɦɵɣ ɜ ɤɨɦɩɶɸɬɟɪ ɚɧɚɥɨɝɨ-ɰɢɮɪɨɜɨɣ ɩɪɟɨɛɪɚɡɨɜɚɬɟɥɶ CompuScope 8012Ⱥ-2Ɇ (“Gage Applied, Inc.”). ȼ ɤɚɱɟɫɬɜɟ ɢɫɬɨɱɧɢɤɚ ɜɨɡɛɭɠɞɚɸɳɟɝɨ ɫɜɟɬɚ ɢɫɩɨɥɶɡɨɜɚɥɫɹ ɧɟɨɞɢɦɨɜɵɣ ɥɚɡɟɪ Nd:YAG ɮɢɪɦɵ “Quantel” (Ɏɪɚɧɰɢɹ) ɫ ɭɞɜɨɟɧɢɟɦ ɱɚɫɬɨɬɵ (O 532 ɧɦ, W 15 ɧɫ, ȿ 30 ɦȾɠ/ɫɦ2). Ⱦɥɹ ȻɊ ɲɬɚɦɦɨɜ ET1001 ɢ D96N ɛɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ ɜ ɫɪɚɜɧɟɧɢɢ ɫ ɢɫɯɨɞɧɵɦɢ ɉɆ ɫɤɨɪɨɫɬɢ ɨɛɪɚɡɨɜɚɧɢɹ ɢ ɪɟɥɚɤɫɚɰɢɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɨɜ ɜ ɪɟɝɟɧɟɪɢɪɨɜɚɧɧɵɯ all-Eɪɟɬɢɧɚɥɟɦ (1) ɚɩɨɦɟɦɛɪɚɧɚɯ ɢɡɦɟɧɢɥɢɫɶ ɨɱɟɧɶ ɦɚɥɨ. Ʉɨɧɬɪɨɥɶɧɵɟ ɩɪɟɩɚɪɚɬɵ ȻɊ, ɩɨɥɭɱɟɧɧɵɟ ɪɟɝɟɧɟɪɚɰɢɟɣ ɛɚɤɬɟɪɢɨɨɩɫɢɧɚ ɢ ɪɟɬɢɧɚɥɹ (1), ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ɢɫɯɨɞɧɵɦɢ ɨɛɪɚɡɰɚɦɢ ɛɟɥɤɨɜ ɩɨɤɚɡɚɥɢ ɧɟɛɨɥɶɲɨɟ ɡɚɦɟɞɥɟɧɢɟ ɪɚɫɩɚɞɚ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɞɥɹ ȻɈ ɢɡ ɲɬɚɦɦɚ ȿɌ1001 ɢ, ɧɚɨɛɨɪɨɬ, ɧɟɤɨɬɨɪɨɟ ɟɝɨ ɭɫɤɨɪɟɧɢɟ ɞɥɹ ȻɈD96N (ɪɢɫ. 26). Ʉɨɧɬɪɨɥɶɧɵɟ ɢɡɦɟɪɟɧɢɹ ɢɫɯɨɞɧɵɯ ɩɪɟɩɚɪɚɬɨɜ ȻɊ (ɞɨ ɨɛɟɫɰɜɟɱɢɜɚɧɢɹ) ɲɬɚɦɦɨɜ ET1001, JW5 ɢ D96N ɜɵɹɜɢɥɢ ɩɪɢɫɭɳɭɸ ɢɦ ɪɚɡɧɢɰɭ ɤɢɧɟɬɢɤ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ (ɫɦ. ɪɢɫ. 25). ɉɨ ɫɜɨɢɦ ɤɢɧɟɬɢɱɟɫɤɢɦ ɩɚɪɚɦɟɬɪɚɦ ɩɪɟɩɚɪɚɬɵ ȻɊȿɌ1001 ɢ ȻɊJW5 ɞɨɫɬɚɬɨɱɧɨ ɛɥɢɡɤɢ. ɍ ȻɊJW5 ɨɬɦɟɱɟɧɨ ɧɟɫɤɨɥɶɤɨ ɛɨɥɟɟ ɛɵɫɬɪɨɟ ɨɛɪɚɡɨɜɚɧɢɟ ɢ ɧɟɦɧɨɝɨ ɡɚɦɟɞɥɟɧɧɵɣ ɪɚɫɩɚɞ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɩɨ ɫɪɚɜɧɟɧɢɸ ɫ ȻɊȿɌ1001. ɍ ȻɊD96N ɡɧɚɱɢɬɟɥɶɧɨ ɡɚɦɟɞɥɟɧɵ ɤɢɧɟɬɢɤɢ ɤɚɤ ɨɛɪɚɡɨɜɚɧɢɹ, ɬɚɤ ɢ ɪɚɫɩɚɞɚ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ. 1 D96N

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0.8 0.6 0.4

D96N

0.2 0

ET1001

JW5 50 ɦɫ

10 c

Ɋɢɫ. 25. ɋɪɚɜɧɟɧɢɟ ɤɢɧɟɬɢɤɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ȻɊ ɲɬɚɦɦɨɜ ȿɌ1001, JW5 ɢ D96N.

83 Ⱦɥɹ ɜɫɟɯ ȺȻɊ ɧɟ ɧɚɛɥɸɞɚɥɨɫɶ ɫɭɳɟɫɬɜɟɧɧɨɝɨ ɢɡɦɟɧɟɧɢɹ ɷɮɮɟɤɬɢɜɧɨɫɬɢ ɮɨɬɨɰɢɤɥɚ ɩɪɢ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɡɚɦɟɧɟ ɜ ɩɨɥɢɩɟɩɬɢɞɧɨɣ ɰɟɩɢ. ɍ 3,4-ɞɢɞɟɝɢɞɪɨ-ȻɊ ɫɤɨɪɨɫɬɢ ɨɛɪɚɡɨɜɚɧɢɹ ɢ ɪɚɫɩɚɞɚ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ (ɫɦ. ɪɢɫ. 26) ɛɵɥɢ ɜɟɫɶɦɚ ɛɥɢɡɤɢ ɤ ɢɫɯɨɞɧɨɣ ɭ ɤɨɧɬɪɨɥɶɧɵɯ ɩɪɟɩɚɪɚɬɨɜ ɫɨɨɬɜɟɬɫɬɜɭɸɳɟɝɨ ɲɬɚɦɦɚ. ɉɨɤɚɡɚɧɨ ɧɚɥɢɱɢɟ ɞɨɩɨɥɧɢɬɟɥɶɧɨɝɨ ɡɚɦɟɞɥɟɧɢɹ ɪɚɫɩɚɞɚ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɡɚ ɫɱɟɬ ɯɪɨɦɨɮɨɪɚ ɭ ɮɟɧɢɥ-ȻɊ ɢ 4-ɨɤɫɨ-ȻɊ ɜ ɫɢɥɶɧɨ ɡɚɦɟɞɥɟɧɧɨɦ ɰɢɤɥɟ ɦɭɬɚɧɬɚ D96N (ɪɢɫ. 26ȼ). Ʉɪɨɦɟ ɬɨɝɨ, ɫɨɯɪɚɧɢɥɢɫɶ ɢ ɯɚɪɚɤɬɟɪɧɵɟ ɪɚɡɥɢɱɢɹ ɜ ɮɨɪɦɟ ɤɪɢɜɨɣ ɮɨɬɨɨɬɜɟɬɚ: ɤɪɢɜɵɟ ɪɟɥɚɤɫɚɰɢɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɭ 4-ɨɤɫɨ-ɚɧɚɥɨɝɨɜ ȻɊET1001 ɢ ȻɊD96N ɞɜɭɯɮɚɡɧɵ, ɩɪɢɱɟɦ ɫɢɥɶɧɨ ɡɚɦɟɞɥɟɧɧɚɹ ɱɚɫɬɶ ɫɨɫɬɚɜɢɥɚ ~ 10-15 % ɨɬ ɚɦɩɥɢɬɭɞɵ ɫɢɝɧɚɥɚ (ɫɦ. ɪɢɫ. 26). B)

1 0.8

4

3

2 1

0.6 1

0.4

2, 3

0.2 0

4

'A, ɨɬɧ. ɟɞ.

'A, ɨɬɧ. ɟɞ.

A)

1 0.8 0.6 0.4

1,2,3 5

4

0.2 0

50 ɦɫ

10 c

200 c

Ɋɢɫ. 26. Ʉɢɧɟɬɢɤɚ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ ɞɥɹ ȺȻɊ ɢɡ ɚɩɨɦɟɦɛɪɚɧ A) ɲɬɚɦɦɚ ȿɌ1001, B) ɲɬɚɦɦɚ D96N. ɂɡɦɟɧɟɧɢɟ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɜ ɦɚɤɫɢɦɭɦɟ ɩɨɝɥɨɳɟɧɢɹ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ. 1 – ɢɫɯɨɞɧɵɟ ɦɟɦɛɪɚɧɵ, 2 – ɪɟɝɟɧɟɪɢɪɨɜɚɧɧɵɟ ɪɟɬɢɧɚɥɟɦ (1), 3 – ɫ 3,4-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɟɦ (2), 4 – ɫ 4-ɨɤɫɨɪɟɬɢɧɚɥɟɦ (3), 5 – ɮɟɧɢɥɪɟɬɢɧɚɥɟɦ (4). Ɋɚɡɥɢɱɢɟ ɩɪɨɹɜɢɥɨɫɶ ɜ ɧɚɥɢɱɢɢ ɧɚɱɚɥɶɧɨɝɨ ɡɚɦɟɞɥɟɧɢɹ ɪɚɫɩɚɞɚ Ɇ-ɩɨɞɨɛɧɨɝɨ ɢɧɬɟɪɦɟɞɢɚɬɚ ɭ ɮɟɧɢɥ-ȻɊD96N ɢ 4-ɨɤɫɨ-ȻɊD96N (ɞɥɹ ɨɛɨɢɯ ɦɭɬɚɧɬɧɵɯ ɚɧɚɥɨɝɨɜ t1/2 ɡɚɦɟɬɧɨ ɭɜɟɥɢɱɟɧɨ), ɤɨɬɨɪɨɟ ɨɬɫɭɬɫɬɜɭɟɬ ɭ ɫɨɨɬɜɟɬɫɬɜɭɸɳɢɯ ȺȻɊ ɫ ɩɪɢɪɨɞɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ. Ȼɵɥɨ ɩɨɤɚɡɚɧɨ, ɱɬɨ 5,6-ɞɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɶ (5) ɨɛɪɚɡɭɟɬ ɩɢɝɦɟɧɬɵ ɫ ȻɈ (ɛɵɥɢ ɢɫɩɨɥɶɡɨɜɚɧɵ ɲɬɚɦɦɵ D96N ɢ JW5), ɢɦɟɸɳɢɟ Omax ɨɤɨɥɨ 450 ɧɦ, ɨɞɧɚɤɨ ɨɧɢ ɧɟ ɞɚɸɬ ɜɵɪɚɠɟɧɧɨɝɨ ɮɨɬɨɨɬɜɟɬɚ ɧɚ ɥɚɡɟɪɧɭɸ ɜɫɩɵɲɤɭ ɧɢ ɜ ɨɛɥɚɫɬɢ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ, ɧɢ ɜ ɦɚɤɫɢɦɭɦɟ ɩɨɝɥɨɳɟɧɢɹ ɯɪɨɦɨɮɨɪɚ. ɉɨɥɭɱɟɧɧɵɟ ɪɟɡɭɥɶɬɚɬɵ ɧɟ ɫɨɝɥɚɫɭɸɬɫɹ ɫ ɥɢɬɟɪɚɬɭɪɧɵɦɢ ɞɚɧɧɵɦɢ ɨ ɬɨɦ, ɱɬɨ ɧɚɥɢɱɢɟ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɝɨ ɤɨɥɶɰɚ ɧɟ ɹɜɥɹɟɬɫɹ ɧɟɨɛɯɨɞɢɦɵɦ ɞɥɹ ɫɭɳɟɫɬɜɨɜɚɧɢɹ ɮɨɬɨɰɢɤɥɚ ɢ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɚ [192]. Ɍɚɤ, ɩɨ

84 ɞɚɧɧɵɦ Crouch R.K. [193], ɩɨɥɢɟɧɚɥɶ, ɩɪɟɞɫɬɚɜɥɹɸɳɢɣ ɫɨɛɨɣ ɚɰɢɤɥɢɱɟɫɤɢɣ ɮɪɚɝɦɟɧɬ ɪɟɬɢɧɚɥɹ (25) (ɫɦ. ɪɢɫ. 27) ɨɛɪɚɡɭɟɬ ɫ ȻɈ ɩɢɝɦɟɧɬ ɫ Omax 466 ɧɦ, ɨɛɥɚɞɚɸɳɢɣ ɮɨɬɨɰɢɤɥɨɦ ɢ ɫɜɟɬɨɡɚɜɢɫɢɦɵɦ ɬɪɚɧɫɩɨɪɬɨɦ ɩɪɨɬɨɧɚ, ɚ ɩɨɥɢɟɧɚɥɶ, ɫɨɞɟɪɠɚɳɢɣ 5 ɞɜɨɣɧɵɯ ɫɜɹɡɟɣ (26) ɨɛɪɚɡɭɟɬ ɞɚɠɟ ɛɨɥɟɟ ɫɬɚɛɢɥɶɧɵɣ ɩɢɝɦɟɧɬ c Omax 519 ɧɦ [193]. O

O

O

O (5)

(25)

O

Ɋɢɫ. 27. Ⱥɰɢɤɥɢɱɟɫɤɢɟ ɚɧɚɥɨɝɢ ɪɟɬɢɧɚɥɹ.

(26)

85

ɗɄɋɉȿɊɂɆȿɇɌȺɅɖɇȺə ɑȺɋɌɖ. ȼ ɪɚɛɨɬɟ ɢɫɩɨɥɶɡɨɜɚɧɵ ɪɟɚɤɬɢɜɵ: ɯɥɨɪɨɯɪɨɦɚɬ ɩɢɪɢɞɢɧɢɹ (Aldrich, ɋɒȺ), ɬɪɢɮɟɧɢɥɮɨɫɮɢɧ “zur synthese” (Merck, Ƚɟɪɦɚɧɢɹ), ɚɤɬɢɜɧɵɣ ɞɢɨɤɫɢɞ ɦɚɪɝɚɧɰɚ “zur synthese” (Merck, Ƚɟɪɦɚɧɢɹ), ɚɝɚɪɨɡɚ (CHEMAPOL, ɑɟɯɢɹ), ɛɚɤɬɨɩɟɩɬɨɧ ɢ ɞɪɨɠɠɟɜɨɣ ɷɤɫɬɪɚɤɬ (Difco, ɋɒȺ), ɩɚɧɤɪɟɚɬɢɱɟɫɤɚɹ ɞɟɡɨɤɫɢɪɢɛɨɧɭɤɥɟɚɡɚ I ɬɟɥɟɧɤɚ (Fluka, ɒɜɟɣɰɚɪɢɹ), MES («Sigma», ɋɒȺ), ɝɢɞɪɨɤɫɢɥɚɦɢɧ («Merck»), ɚɤɪɢɥɚɦɢɞ ɢ ɤɭɦɚɫɫɢ ɛɪɢɥɥɢɚɧɬɨɜɵɣ ɫɢɧɢɣ G250 (Serva, Ƚɟɪɦɚɧɢɹ). Ⱦɥɹ ɫɢɧɬɟɡɚ ɢɫɩɨɥɶɡɨɜɚɥɢ ɫɨɥɢ ɢ ɪɚɫɬɜɨɪɢɬɟɥɢ ɦɚɪɤɢ “ɯ.ɱ.” ɩɪɨɢɡɜɨɞɫɬɜɚ Ɋɨɫɫɢɢ, ɞɥɹ ɪɚɛɨɬɵ ɫ ɛɢɨɥɨɝɢɱɟɫɤɢɦɢ ɨɛɴɟɤɬɚɦɢ ɢɫɩɨɥɶɡɨɜɚɥɢ ɫɨɥɢ ɦɚɪɤɢ “ɱ.ɞ.ɚ.” ɩɪɨɢɡɜɨɞɫɬɜɚ Ɋɨɫɫɢɢ. ɉɨɥɭɱɟɧɢɟ «ɫɭɯɨɝɨ» ɌȽɎ ɢ ɞɢɷɬɢɥɨɜɨɝɨ ɷɮɢɪɚ ɨɫɭɳɟɫɬɜɥɹɥɢ ɩɟɪɟɝɨɧɤɨɣ ɧɚɞ ɚɥɸɦɨɝɢɞɪɢɞɨɦ ɥɢɬɢɹ. Ⱦɢɯɥɨɪɦɟɬɚɧ ɨɱɢɳɚɥɢ ɩɟɪɟɝɨɧɤɨɣ ɧɚɞ ɩɨɬɚɲɨɦ. Ʉɚɱɟɫɬɜɟɧɧɵɣ ɫɨɫɬɚɜ ɪɟɚɤɰɢɨɧɧɵɯ ɫɦɟɫɟɣ ɢ ɢɧɞɢɜɢɞɭɚɥɶɧɨɫɬɶ ɫɨɟɞɢɧɟɧɢɣ ɤɨɧɬɪɨɥɢɪɨɜɚɥɢ ɦɟɬɨɞɨɦ Ɍɋɏ ɧɚ ɩɥɚɫɬɢɧɚɯ Kieselgel 60 F254 (Merck, Ƚɟɪɦɚɧɢɹ) ɢɥɢ Silufol UV-254 (Kavalier, ɑɟɯɢɹ) ɜ ɫɢɫɬɟɦɟ ɝɟɤɫɚɧ-ɞɢɷɬɢɥɨɜɵɣ ɷɮɢɪ, 1:1; ɨɛɧɚɪɭɠɟɧɢɟ ɩɹɬɟɧ - ɨɩɪɵɫɤɢɜɚɧɢɟɦ ɩɪɨɹɜɥɟɧɧɨɣ ɩɥɚɫɬɢɧɤɢ ɤɨɧɰɟɧɬɪɢɪɨɜɚɧɧɨɣ ɫɟɪɧɨɣ ɤɢɫɥɨɬɨɣ. Ⱥɞɫɨɪɛɰɢɨɧɧɭɸ ɤɨɥɨɧɨɱɧɭɸ ɯɪɨɦɚɬɨɝɪɚɮɢɸ ɩɪɨɜɨɞɢɥɢ ɧɚ ɫɢɥɢɤɚɝɟɥɟ Kieselgel 60 (0,063 ɦɤɦ) (Merck, Ƚɟɪɦɚɧɢɹ). ȼɫɟ ɨɩɟɪɚɰɢɢ ɫ ɪɟɚɝɟɧɬɚɦɢ, ɱɭɜɫɬɜɢɬɟɥɶɧɵɦɢ ɤ ɤɢɫɥɨɪɨɞɭ ɜɨɡɞɭɯɚ, ɩɪɨɜɨɞɢɥɢ ɜ ɚɬɦɨɫɮɟɪɟ ɫɭɯɨɝɨ ɚɪɝɨɧɚ. Ɋɚɫɬɜɨɪɢɬɟɥɢ ɭɞɚɥɹɥɢ ɧɚ ɜɚɤɭɭɦɧɨɦ ɪɨɬɨɪɧɨɦ ɢɫɩɚɪɢɬɟɥɟ ɩɪɢ ɬɟɦɩɟɪɚɬɭɪɟ ɧɟ ɜɵɲɟ 35qɋ. ɋ ɩɪɟɩɚɪɚɬɚɦɢ ȻɊ, ɟɝɨ ɚɧɚɥɨɝɨɜ ɢ ɚɩɨɦɟɦɛɪɚɧ ɪɚɛɨɬɚɥɢ ɩɪɢ 4-7qɋ, ɯɪɚɧɢɥɢ ɢɯ ɩɪɢ 4qɋ. ȼ ɪɚɛɨɬɟ ɢɫɩɨɥɶɡɨɜɚɥɢ ɞɢɚɥɢɡɧɵɟ ɦɟɲɤɢ Spectrapor (SMI INC., ɋɒȺ). ɂɡɦɟɪɟɧɢɹ ɪɇ ɨɫɭɳɟɫɬɜɥɹɥɢ ɩɪɢ ɩɪɢɝɨɬɨɜɥɟɧɢɢ Ʉɀ ɫ ɩɨɦɨɳɶɸ ɛɭɦɚɠɧɵɯ ɪɇɢɧɞɢɤɚɬɨɪɨɜ ɞɥɹ ɢɧɬɟɪɜɚɥɚ ɪɇ 6.5-9.0 (Merck, Ƚɟɪɦɚɧɢɹ), ɩɪɢ ɩɪɢɝɨɬɨɜɥɟɧɢɢ ɛɭɮɟɪɧɵɯ ɪɚɫɬɜɨɪɨɜ ɪɇ-ɦɟɬɪɨɦ Metrohm-632 (ɒɜɟɰɢɹ). ɍɎ-ɫɩɟɤɬɪɵ ɩɪɟɩɚɪɚɬɨɜ ɦɟɦɛɪɚɧ ɫɧɢɦɚɥɢ ɧɚ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɟ Hitachi -330 (əɩɨɧɢɹ), ɪɚɫɬɜɨɪɨɜ ɫɨɟɞɢɧɟɧɢɣ ɜ ɦɟɬɚɧɨɥɟ - ɧɚ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɟ Hitachi -3400 (əɩɨɧɢɹ), ɨɩɬɢɱɟɫɤɭɸ ɩɥɨɬɧɨɫɬɶ ɤɥɟɬɨɱɧɵɯ ɫɭɫɩɟɧɡɢɣ ɨɩɪɟɞɟɥɹɥɢ ɧɚ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɟ Philips PU 8750 UV/VIS (Ƚɟɪɦɚɧɢɹ). ɋɩɟɤɬɪɵ 1ɇ- ɢ 13ɋ- əɆɊ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɫɧɢɦɚɥɢ ɜ ɞɟɣɬɟɪɨɯɥɨɪɨɮɨɪɦɟ ɧɚ ɫɩɟɤɬɪɨɦɟɬɪɟ Bruker-MSL-200 (Ƚɟɪɦɚɧɢɹ) ɫ ɪɚɛɨɱɟɣ ɱɚɫɬɨɬɨɣ 200 ɆȽɰ ɢ 50,32 ɆȽɰ ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ. ȼ ɤɚɱɟɫɬɜɟ ɜɧɭɬɪɟɧɧɟɝɨ ɫɬɚɧɞɚɪɬɚ ɢɫɩɨɥɶɡɨɜɚɥɢ ɝɟɤɫɚɦɟɬɢɥɞɢɫɢɥɨɤɫɚɧ (1ɇ-əɆɊ G 0.055). ɏɢɦɢɱɟɫɤɢɟ ɫɞɜɢɝɢ ɩɪɢɜɟɞɟɧɵ ɜ ɦɢɥɥɢɨɧɧɵɯ ɞɨɥɹɯ, ɤɨɧɫɬɚɧɬɵ ɫɩɢɧ-ɫɩɢɧɨɜɨɝɨ ɜɡɚɢɦɨɞɟɣɫɬɜɢɹ – ɜ ɝɟɪɰɚɯ. ɉɪɢ ɨɩɢɫɚɧɢɢ ɫɩɟɤɬɪɨɜ ɢɫɩɨɥɶɡɨɜɚɧɵ ɫɥɟɞɭɸɳɢɟ ɫɨɤɪɚɳɟɧɢɹ: ɫ – ɫɢɧɝɥɟɬ, ɞ – ɞɭɛɥɟɬ, ɬ – ɬɪɢɩɥɟɬ, ɞɞ – ɞɭɛɥɟɬ ɞɭɛɥɟɬɨɜ, ɦ – ɦɭɥɶɬɢɩɥɟɬ, ɩɥ. – ɩɥɟɱɨ.

86 ɋɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ ɩɪɟɩɚɪɚɬɨɜ ȻɊ ɨɫɭɳɟɫɬɜɥɹɥɢ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɪɬɭɬɧɨɣ ɥɚɦɩɵ ɜɵɫɨɤɨɝɨ ɞɚɜɥɟɧɢɹ ȾɊɒ-500Ɇ, ɫɧɚɛɠɟɧɧɨɣ ɮɨɤɭɫɢɪɭɸɳɟɣ ɫɢɫɬɟɦɨɣ, ɬɟɩɥɨɜɵɦ ɮɢɥɶɬɪɨɦ ɋ3ɋ22 ɢ ɫɜɟɬɨɮɢɥɶɬɪɨɦ ɀɋ18. Ʉɢɧɟɬɢɤɭ Ɇ-ɢɧɬɟɪɦɟɞɢɚɬɚ ɮɨɬɨɰɢɤɥɨɜ ɚɧɚɥɨɝɨɜ ȻɊ ɪɟɝɢɫɬɪɢɪɨɜɚɥɢ ɧɚ ɭɫɬɚɧɨɜɤɟ ɢɦɩɭɥɶɫɧɨɣ ɫɩɟɤɬɪɨɦɟɬɪɢɢ ɜ ɥɚɛɨɪɚɬɨɪɢɢ ɮɨɬɨɯɢɦɢɢ ɛɢɨɦɟɦɛɪɚɧ ɇɂɂ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɨɣ ɛɢɨɥɨɝɢɢ ɢɦ. Ⱥ.ɇ. Ȼɟɥɨɡɟɪɫɤɨɝɨ ɆȽɍ. ɂɫɫɥɟɞɨɜɚɧɢɟ ɩɪɟɩɚɪɚɬɨɜ ȻɊ ɢ ɟɝɨ ɚɧɚɥɨɝɨɜ ɩɪɨɜɨɞɢɥɢ ɜ ɫɭɫɩɟɧɡɢɹɯ ɜ ɫɬɚɧɞɚɪɬɧɨɦ ɛɭɮɟɪɟ (100 ɦɆ NaCl, 5 ɦɆ MES, ɪɇ 6) ɩɪɢ 21qC. ɉɪɢɧɰɢɩ ɪɚɛɨɬɵ ɩɪɢɛɨɪɚ: ɢɡɦɟɪɹɸɳɢɣ ɥɭɱ ɨɬ ɤɜɚɪɰɟɜɨɣ ɝɚɥɨɝɟɧɨɜɨɣ ɥɚɦɩɵ ɄȽɆ (9ȼ, 75 ȼɬ) ɩɪɨɯɨɞɢɥ ɱɟɪɟɡ ɪɟɲɟɬɨɱɧɵɣ ɦɨɧɨɯɪɨɦɚɬɨɪ HL1 (ɮɢɪɦɵ Jobin-Yvon) ɫ ɲɢɪɢɧɨɣ ɳɟɥɢ 2-8 ɦɦ, ɬɟɪɦɨɫɬɚɬɢɪɭɟɦɭɸ ɤɸɜɟɬɭ (ɞɥɢɧɚ ɨɩɬɢɱɟɫɤɨɝɨ ɩɭɬɢ 10 ɦɦ) ɫ ɨɛɪɚɡɰɨɦ, ɜɬɨɪɨɣ ɦɨɧɨɯɪɨɦɚɬɨɪ ɢ ɩɨɩɚɞɚɥ ɧɚ ɮɨɬɨɭɦɧɨɠɢɬɟɥɶ Ɏɗɍ-128. ɗɥɟɤɬɪɢɱɟɫɤɢɣ ɫɢɝɧɚɥ ɫ Ɏɗɍ ɩɨɞɚɜɚɥɫɹ ɧɚ ɨɩɟɪɚɰɢɨɧɧɵɣ ɭɫɢɥɢɬɟɥɶ Burr Brown 3554ȼɆ, ɚ ɫ ɭɫɢɥɢɬɟɥɹ – ɧɚ ɜɫɬɪɚɢɜɚɟɦɵɣ ɜ ɤɨɦɩɶɸɬɟɪ ɚɧɚɥɨɝɨɰɢɮɪɨɜɨɣ ɩɪɟɨɛɪɚɡɨɜɚɬɟɥɶ CompuScope 8012Ⱥ-2Ɇ (“Gage Applied, Inc.”). ȼ ɤɚɱɟɫɬɜɟ ɢɫɬɨɱɧɢɤɚ ɜɨɡɛɭɠɞɚɸɳɟɝɨ ɫɜɟɬɚ ɢɫɩɨɥɶɡɨɜɚɥɫɹ ɧɟɨɞɢɦɨɜɵɣ ɥɚɡɟɪ Nd:YAG ɮɢɪɦɵ “Quantel” (Ɏɪɚɧɰɢɹ) ɫ ɭɞɜɨɟɧɢɟɦ ɱɚɫɬɨɬɵ (O 532 ɧɦ, W 15 ɧɫ, ȿ 30 ɦȾɠ/ɫɦ2). SDS-ɷɥɟɤɬɪɨɮɨɪɟɡ ɩɪɟɩɚɪɚɬɨɜ ȻɊ ɩɪɨɢɡɜɨɞɢɥɢ ɜ 15% ɩɨɥɢɚɤɪɢɥɚɦɢɞɧɨɦ ɝɟɥɟ ɪɚɡɦɟɪɨɦ 125u76 ɦɦ ɩɪɢ ɩɨɫɬɨɹɧɧɨɦ ɬɨɤɟ 90 ɦȺ ɫ ɢɫɬɨɱɧɢɤɨɦ ɩɢɬɚɧɢɹ PS 500XT DC (Hoeffer scientific instruments, ɋɒȺ), ɩɨɫɥɟ ɮɢɤɫɚɰɢɢ ɝɟɥɹ ɜ ɬɟɱɟɧɢɟ 10 ɦɢɧ (10% ɬɪɢɯɥɨɪɭɤɫɭɫɧɨɣ ɤɢɫɥɨɬɵ, 10% ɢɡɨɩɪɨɩɚɧɨɥɚ ɜ ɜɨɞɟ) ɩɪɨɜɨɞɢɥɢ ɟɝɨ ɨɤɪɚɫɤɭ ɜ ɬɟɱɟɧɢɟ 1 ɱ ɩɪɢ ɩɨɦɨɳɢ ɜɨɞɧɨɝɨ ɪɚɫɬɜɨɪɚ, ɫɨɞɟɪɠɚɳɟɝɨ 0,1% ɤɭɦɚɫɫɢ ɛɪɢɥɥɢɚɧɬɨɜɨɝɨ ɫɢɧɟɝɨ, 10% ɭɤɫɭɫɧɨɣ ɤɢɫɥɨɬɵ ɢ 10% ɢɡɨɩɪɨɩɚɧɨɥɚ. Ʉɭɥɶɬɢɜɢɪɨɜɚɧɢɟ ɛɚɤɬɟɪɢɣ ɩɪɨɜɨɞɢɥɢ ɧɚ ɜɪɚɳɚɬɟɥɶɧɨɣ ɤɚɱɚɥɤɟ (190 ɦɢɧ-1) ɫ ɬɟɪɦɨɫɬɚɬɢɪɨɜɚɧɢɟɦ "LP-125" (ȼɟɧɝɪɢɹ) ɢɥɢ ɤɚɱɚɥɤɟ ɪɨɫɫɢɣɫɤɨɝɨ ɩɪɨɢɡɜɨɞɫɬɜɚ, ɥɢɛɨ ɜ ɩɪɨɡɪɚɱɧɨɣ ɤɸɜɟɬɟ ɨɛɴɟɦɨɦ a6 ɥ, ɩɪɢ ɨɫɜɟɳɟɧɢɢ 4 ɥɚɦɩɚɦɢ ɞɧɟɜɧɨɝɨ ɫɜɟɬɚ ɅȾ-20 ɫ ɩɨɞɚɱɟɣ ɜɨɡɞɭɯɚ ɧɚɫɨɫɨɦ AIR PUMP AC-9601 (RESUN, ɄɇɊ). ɋɬɟɪɢɥɢɡɚɰɢɸ ɪɨɫɬɨɜɵɯ ɫɪɟɞ ɢ ɫɬɟɤɥɹɧɧɨɣ ɩɨɫɭɞɵ ɩɪɨɜɨɞɢɥɢ ɜ ɚɜɬɨɤɥɚɜɟ “ST174”, ȼɟɧɝɪɢɹ. ȼ ɪɚɛɨɬɟ ɢɫɩɨɥɶɡɨɜɚɧɵ ɝɚɥɨɮɢɥɶɧɵɟ ɛɚɤɬɟɪɢɢ Halobacterium salinarum ɢɡ ɤɨɥɥɟɤɰɢɢ ɤɭɥɶɬɭɪ ɦɢɤɪɨɨɪɝɚɧɢɡɦɨɜ ɂɧɫɬɢɬɭɬɚ ɛɢɨɮɢɡɢɤɢ ɤɥɟɬɤɢ ɊȺɇ (ɝ. ɉɭɳɢɧɨ) ɢ ɂɧɫɬɢɬɭɬɚ ɛɢɨɯɢɦɢɱɟɫɤɨɣ ɮɢɡɢɤɢ ɢɦ. ɇ.Ɇ. ɗɦɚɧɭɷɥɹ, ɊȺɇ (Ɇɨɫɤɜɚ): ɲɬɚɦɦɵ ET1001, JW5 ɢ D96N.

Ɍɚɛɥɢɰɚ 14.

87 ɍɫɥɨɜɢɹ ɨɫɚɠɞɟɧɢɹ ɛɚɤɬɟɪɢɚɥɶɧɵɯ ɤɥɟɬɨɤ ɢ ɩɪɟɩɚɪɚɬɨɜ ɦɟɦɛɪɚɧ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɟɦ.

Ɉɫɚɠɞɟɧɢɟ ɤɥɟɬɨɤ ɢɡ Ʉɀ

Ɍɢɩ ɰɟɧɬɪɢɮɭɝɢ

Ɋɨɬɨɪ

ɍɫɥɨɜɢɹ

Beckman J-218

JA-10, 6u500

7000g(6500 ɦɢɧ-1)u30 ɦɢɧ

Beckman J2-21

JA-14, 6u250

10000g(15300 ɦɢɧ-1)u15 ɦɢɧ

8 u 50

50000g(20000 ɦɢɧ-1)u1-2 ɱ

8 u 25

50000g(24000 ɦɢɧ-1)u1-2 ɱ

Ⱦɢɮɮɟɪɟɧɰɢɚɥɶɧɨɟ MSE High Speed 25 ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɟ ɩɪɟɩɚɪɚɬɨɜ MSE High Speed 25 ɦɟɦɛɪɚɧ

Ɇɨɞɟɥɢɪɨɜɚɧɢɟ ɬɪɟɯɦɟɪɧɨɣ ɫɬɪɭɤɬɭɪɵ ȻɊ ɨɫɭɳɟɫɬɜɥɹɥɢ ɧɚ ɤɨɦɩɶɸɬɟɪɟ Pentium III ɫ ɱɚɫɬɨɬɨɣ 700 ɆȽɰ ɢ ɨɩɟɪɚɬɢɜɧɨɣ ɩɚɦɹɬɶɸ 256 Ɇɛ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɩɪɨɝɪɚɦɦɵ Chem3D Ultra ɢɡ ɩɚɤɟɬɚ ChemOffice 6.0 (CambridgeSoft Com, USA).

1. ɋɢɧɬɟɡ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ. ɋɢɧɬɟɡ all-E-3,4-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɹ (2). Ʉ ɪɚɫɬɜɨɪɭ 216 ɦɝ (0.76 ɦɦɨɥɶ) all-E-ɪɟɬɢɧɚɥɹ (1) ɜ 10 ɦɥ «ɫɭɯɨɝɨ» ɌȽɎ ɩɪɢ ɩɟɪɟɦɟɲɢɜɚɧɢɢ ɢ 0 0ɋ ɜ ɬɟɦɧɨɬɟ ɢ ɚɬɦɨɫɮɟɪɟ ɚɪɝɨɧɚ ɩɪɢɛɚɜɥɹɥɢ ɩɨ ɤɚɩɥɹɦ ɪɚɫɬɜɨɪ 251 ɦɝ (1.41 ɦɦɨɥɶ) NBS ɜ 13 ɦɥ «ɫɭɯɨɝɨ» ɌȽɎ ɢ ɩɟɪɟɦɟɲɢɜɚɥɢ 12 ɱ. ɉɨɫɥɟ ɡɚɜɟɪɲɟɧɢɹ ɪɟɚɤɰɢɢ ɛɪɨɦɢɪɨɜɚɧɢɹ (ɩɨ ɞɚɧɧɵɦ Ɍɋɏ) ɞɨɛɚɜɥɹɥɢ 2 ɦɥ ȾɆɎȺ ɢ 200 ɦɝ KF ɢ ɩɟɪɟɦɟɲɢɜɚɥɢ 36 ɱ ɩɪɢ 20 0ɋ, ɩɨɫɥɟ ɱɟɝɨ ɜ ɪɟɚɤɰɢɨɧɧɭɸ ɫɦɟɫɶ ɞɨɛɚɜɥɹɥɢ 20 ɦɥ ɜɨɞɵ. ȼɟɳɟɫɬɜɨ ɷɤɫɬɪɚɝɢɪɨɜɚɥɢ ɞɢɷɬɢɥɨɜɵɦ ɷɮɢɪɨɦ (3u10 ɦɥ), ɨɛɴɟɞɢɧɟɧɧɵɣ ɷɤɫɬɪɚɤɬ ɩɪɨɦɵɜɚɥɢ ɜɨɞɨɣ ɞɨ ɪɇ 7, ɫɭɲɢɥɢ 40 ɦɢɧ. ɧɚɞ Na2SO4, ɪɚɫɬɜɨɪɢɬɟɥɶ ɭɞɚɥɹɥɢ. Ɉɫɬɚɬɨɤ ɯɪɨɦɚɬɨɝɪɚɮɢɪɨɜɚɥɢ ɧɚ ɤɨɥɨɧɤɟ, ɷɥɸɢɪɭɹ ɫɦɟɫɶɸ ɝɟɤɫɚɧ – ɷɮɢɪ ɫ ɝɪɚɞɢɟɧɬɨɦ ɩɨɫɥɟɞɧɟɝɨ ɨɬ 0 ɞɨ 5 %. Ɏɪɚɤɰɢɢ, ɫɨɞɟɪɠɚɳɢɟ 3,4-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɶ (2), ɨɛɴɟɞɢɧɹɥɢ, ɪɚɫɬɜɨɪɢɬɟɥɶ ɭɞɚɥɹɥɢ. ȼɵɯɨɞ 108 ɦɝ (0.38 ɦɦɨɥɶ, 50 %); Rf 0.53; tɩɥ 58-59qɋ; ɍɎɫɩɟɤɬɪ, Omax, ɧɦ (H): 398 (45700). 1ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15. Ɉɤɢɫɥɟɧɢɟ all-E-ɪɟɬɢɧɚɥɹ (1) ɯɥɨɪɨɯɪɨɦɚɬɨɦ ɩɢɪɢɞɢɧɢɹ. Ʉ ɫɭɫɩɟɧɡɢɢ 3 ɝ (13.92 ɦɦɨɥɶ) ɯɥɨɪɨɯɪɨɦɚɬɚ ɩɢɪɢɞɢɧɢɹ ɜ 5 ɦɥ ɞɢɯɥɨɪɦɟɬɚɧɚ ɩɪɢɥɢɜɚɥɢ ɪɚɫɬɜɨɪ 1 ɝ (3.51 ɦɦɨɥɶ) all-E-ɪɟɬɢɧɚɥɹ (1) ɜ 20 ɦɥ ɞɢɯɥɨɪɦɟɬɚɧɚ. Ɋɟɚɤɰɢɨɧɧɭɸ ɫɦɟɫɶ ɩɟɪɟɦɟɲɢɜɚɥɢ 1 ɱ, ɨɤɢɫɥɢɬɟɥɶ ɨɬɞɟɥɹɥɢ ɮɢɥɶɬɪɨɜɚɧɢɟɦ ɱɟɪɟɡ ɫɥɨɣ ɰɟɥɢɬɚ, ɚ ɨɫɚɞɨɤ ɩɪɨɦɵɜɚɥɢ 10 ɦɥ ɞɢɯɥɨɪɦɟɬɚɧɚ. Ɏɢɥɶɬɪɚɬ ɭɩɚɪɢɜɚɥɢ, ɪɚɫɬɜɨɪɹɥɢ ɜ 30 ɦɥ ɞɢɷɬɢɥɨɜɨɝɨ ɷɮɢɪɚ ɢ ɪɚɫɬɜɨɪ ɩɪɨɦɵɜɚɥɢ ɜɨɞɨɣ (3 u 15 ɦɥ), ɫɭɲɢɥɢ ɛɟɡɜɨɞɧɵɦ ɫɭɥɶɮɚɬɨɦ ɧɚɬɪɢɹ, ɪɚɫɬɜɨɪɢɬɟɥɶ ɭɩɚɪɢɜɚɥɢ. Ɉɫɬɚɬɨɤ ɯɪɨɦɚɬɨɝɪɚɮɢɪɨɜɚɥɢ ɧɚ ɤɨɥɨɧɤɟ ɫ ɫɢɥɢɤɚɝɟɥɟɦ, ɷɥɸɢɪɭɹ ɜɟɳɟɫɬɜɚ ɝɪɚɞɢɟɧɬɨɦ ɞɢɷɬɢɥɨɜɨɝɨ ɷɮɢɪɚ ɨɬ 0 ɞɨ 100% ɜ ɝɟɤɫɚɧɟ.

88 Ɏɪɚɤɰɢɢ, ɫɨɞɟɪɠɚɳɢɟ ɫɨɨɬɜɟɬɫɬɜɭɸɳɢɟ ɩɪɨɢɡɜɨɞɧɵɟ ɪɟɬɢɧɚɥɹ, ɨɛɴɟɞɢɧɹɥɢ, ɪɚɫɬɜɨɪɢɬɟɥɶ ɭɩɚɪɢɜɚɥɢ, ɚ ɨɫɬɚɬɨɤ ɫɭɲɢɥɢ 1 ɱ ɩɪɢ 0.1 ɦɦ ɪɬ. ɫɬ. 5,6-Ⱦɢɨɤɫɨ-5,6-ɫɟɤɨɪɟɬɢɧɚɥɶ (5). ȼɵɯɨɞ 525 ɦɝ (47 %); Rf 0.09; tɩɥ 76-78qɋ; ɍɎɫɩɟɤɬɪ, Omax, ɧɦ (H): 364 (58000); 1ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15; 13ɋ-əɆɊ: 13.0 (ɋ19, ɋ20), 19.0 (ɋ3), 24.2 (ɋ16, ɋ17), 29.8 (ɋ18), 39.1 (ɋ2), 43.7 (ɋ4), 46.4 (ɋ1), 121.5 (ɋ14), 130.4 (ɋ7), 131.0 (ɋ11), 137.7 (ɋ12), 138.7 (ɋ10), 146.1 (ɋ8), 138.4 (ɋ9), 153.4 (ɋ13), 190.6 (ɋ15), 203.6, 208.2 (ɋ6, ɋ5). 5,6-Ⱦɢɝɢɞɪɨ-5,6-ɷɩɨɤɫɢɪɟɬɢɧɚɥɶ (17). ȼɵɯɨɞ 204 ɦɝ (19 %); Rf 0.48; tɩɥ 104-106qɋ; ɍɎ-ɫɩɟɤɬɪ, Omax, ɧɦ (H): 362 (53500);1ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15. 4-Ɉɤɫɨɪɟɬɢɧɚɥɶ (3). ȼɵɯɨɞ 73 ɦɝ (7 %); Rf 0.39; tɩɥ 113-115qɋ; ɍɎ-ɫɩɟɤɬɪ, Omax, ɧɦ (H): 374 (50500); 1ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15. ɋɢɧɬɟɡ ɮɟɧɢɥɶɧɨɝɨ ɚɧɚɥɨɝɚ ɪɟɬɢɧɚɥɹ 7 {3,7-ɞɢɦɟɬɢɥ-9-ɮɟɧɢɥ-ɧɨɧɚ-2,4,6,8ɬɟɬɪɚɟɧ-1-ɚɥɹ} (4). Ȼɟɧɡɢɥɬɪɢɮɟɧɢɥɮɨɫɮɨɧɢɣ ɛɪɨɦɢɞ (21). ɋɦɟɫɶ 1 ɦɥ ɛɪɨɦɢɫɬɨɝɨ ɛɟɧɡɢɥɚ (20) ɢ 2.4 ɝ ɬɪɢɮɟɧɢɥɮɨɫɮɢɧɚ ɜ 20 ɦɥ «ɫɭɯɨɝɨ» ɬɨɥɭɨɥɚ ɧɚɝɪɟɜɚɥɢ ɩɪɢ 110qɋ 5 ɱ, ɨɯɥɚɠɞɚɥɢ ɞɨ ɤɨɦɧɚɬɧɨɣ ɬɟɦɩɟɪɚɬɭɪɵ, ɞɨɛɚɜɥɹɥɢ 30 ɦɥ «ɫɭɯɨɝɨ» ɷɮɢɪɚ, ɜɵɩɚɜɲɢɣ ɨɫɚɞɨɤ ɨɬɞɟɥɹɥɢ ɮɢɥɶɬɪɨɜɚɧɢɟɦ, ɩɪɨɦɵɜɚɥɢ ɷɮɢɪɨɦ, ɤɪɢɫɬɚɥɥɢɡɨɜɚɥɢ ɢɡ ɫɦɟɫɢ ɷɮɢɪ-ɦɟɬɚɧɨɥ 10:1. ȼɵɯɨɞ ɛɟɧɡɢɥɬɪɢɮɟɧɢɥɮɨɫɮɨɧɢɣ ɛɪɨɦɢɞɚ (21) 3.1 ɝ (80%), ɜ ɜɢɞɟ ɛɟɥɵɯ ɤɪɢɫɬɚɥɥɨɜ, tɩɥ 294qɋ. Ɏɟɧɢɥɶɧɵɣ ɚɧɚɥɨɝ 11,12-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɨɥɚ, 7E,13Z- (22) ɢ 7Z,13Z- (27) ɢɡɨɦɟɪɵ7 {ɫɨɨɬɜɟɬɫɬɜɟɧɧɨ (2Z,8E)- ɢ (2Z,8Z)-ɢɡɨɦɟɪɵ 3,7-ɞɢɦɟɬɢɥ-9-ɮɟɧɢɥ-ɧɨɧɚ-2,6,8ɬɪɢɟɧ-4-ɢɧ-1-ɨɥɚ}. 550 ɦɝ (1.3 ɦɦɨɥɶ) ɨɬɫɭɲɟɧɧɨɝɨ ɩɪɢ 0.2 ɦɦ.ɪɬ.ɫɬ. ɛɟɧɡɢɥɬɪɢɮɟɧɢɥɮɨɫɮɨɧɢɣ ɛɪɨɦɢɞɚ (21) ɢ 450 ɦɝ (1.1 ɦɦɨɥɶ) ɋ10-ɫɢɧɬɨɧɚ (19) ɪɚɫɬɜɨɪɹɥɢ 30 ɦɥ ɯɥɨɪɢɫɬɨɝɨ ɦɟɬɢɥɟɧɚ, ɞɨɛɚɜɥɹɥɢ 2 ɝ K2CO3 ɢ 10 ɦɝ 18-ɤɪɚɭɧ-6. ɋɭɫɩɟɧɡɢɸ ɩɟɪɟɦɟɲɢɜɚɥɢ 30 ɱ ɞɨ ɡɚɜɟɪɲɟɧɢɹ ɪɟɚɤɰɢɢ (ɩɨ ɞɚɧɧɵɦ Ɍɋɏ). Ɉɫɚɞɨɤ ɨɬɮɢɥɶɬɪɨɜɵɜɚɥɢ, ɪɚɫɬɜɨɪɢɬɟɥɶ ɭɩɚɪɢɜɚɥɢ. Ɉɫɬɚɬɨɤ ɪɚɫɬɜɨɪɹɥɢ ɜ 50 ɦɥ ɫɦɟɫɢ ɚɰɟɬɨɧ ɧɚɫɵɳɟɧɧɵɣ ɜɨɞɧɵɣ ɪɚɫɬɜɨɪ ɮɬɨɪɢɞɚ ɚɦɦɨɧɢɹ (3:1) ɢ ɢɧɬɟɧɫɢɜɧɨ ɩɟɪɟɦɟɲɢɜɚɥɢ 3 ɱ ɩɪɢ 20qɋ. ȼɟɳɟɫɬɜɨ ɷɤɫɬɪɚɝɢɪɨɜɚɥɢ ɯɥɨɪɨɮɨɪɦɨɦ, ɨɛɴɟɞɢɧɟɧɧɵɣ ɷɤɫɬɪɚɤɬ ɩɪɨɦɵɜɚɥɢ __________________________ ɫɦ. ɤɨɦɦɟɧɬɚɪɢɢ ɨ ɧɭɦɟɪɚɰɢɢ ɚɬɨɦɨɜ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɚɧɚɥɨɝɚ (4) ɢ ɟɝɨ ɩɪɟɞɲɟɫɬɜɟɧɧɢɤɨɜ (22), (23), ɚ ɬɚɤɠɟ (27), (28) ɜ ɫɧɨɫɤɟ ɧɚ ɫ. 59.

7

89 ɜɨɞɨɣ ɞɨ ɪɇ 7, ɫɭɲɢɥɢ ɧɚɞ ɫɭɥɶɮɚɬɨɦ ɧɚɬɪɢɹ, ɪɚɫɬɜɨɪɢɬɟɥɶ ɭɩɚɪɢɜɚɥɢ. Ɉɫɬɚɬɨɤ ɯɪɨɦɚɬɨɝɪɚɮɢɪɨɜɚɥɢ ɧɚ ɤɨɥɨɧɤɟ ɫ ɫɢɥɢɤɚɝɟɥɟɦ ɜ ɫɢɫɬɟɦɟ ɉɗɅ-ɷɮɢɪ ɫ ɝɪɚɞɢɟɧɬɨɦ ɩɨɫɥɟɞɧɟɝɨ ɨɬ 0 ɞɨ 50%.

ȼɵɯɨɞ 7ȿ,13Z-ɢɡɨɦɟɪɚ 8 (22) 103 ɦɝ (34%); Rf 0,39; tɩɥ 101-102qɋ; ɍɎ-ɫɩɟɤɬɪ, Omax, ɧɦ (H): 320 (40200), ɩɥ; 332 (47000), 346 (35000), ɩɥ.; 1ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15. 7Z,13Z-ɢɡɨɦɟɪ 8 (27): Rf 0,43; ɦɚɫɥɨ; ɍɎ-ɫɩɟɤɬɪ, Omax, ɧɦ (H): 257(11000), 313 (19800); 1

ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15. Ɏɟɧɢɥɶɧɵɣ ɚɧɚɥɨɝ 13Z-11,12-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɹ8 {(2Z,8E)-3,7-ɞɢɦɟɬɢɥ-9-

ɮɟɧɢɥ-ɧɨɧɚ-2,6,8-ɬɪɢɟɧ-4-ɢɧ-1-ɚɥɶ} (23). Ʉ ɪɚɫɬɜɨɪɭ 330 ɦɝ (1,38 ɦɦɨɥɶ) ɮɟɧɢɥɶɧɨɝɨ ɚɧɚɥɨɝɚ 11,12-ɞɟɝɢɞɪɨɪɟɬɢɧɨɥɚ (22) ɜ 50 ɦɥ «ɫɭɯɨɝɨ» ɛɟɧɡɨɥɚ ɞɨɛɚɜɥɹɥɢ 1,8 ɝ ɚɤɬɢɜɧɨɝɨ ɞɢɨɤɫɢɞɚ ɦɚɪɝɚɧɰɚ ɢ ɫɭɫɩɟɧɡɢɸ ɢɧɬɟɧɫɢɜɧɨ ɩɟɪɟɦɟɲɢɜɚɥɢ ɜ ɚɬɦɨɫɮɟɪɟ ɚɪɝɨɧɚ ɩɪɢ 20qɋ ɜ ɬɟɱɟɧɢɟ 5 ɱ. Ɉɤɢɫɥɢɬɟɥɶ ɨɬɮɢɥɶɬɪɨɜɵɜɚɥɢ ɱɟɪɟɡ ɫɥɨɣ ɰɟɥɢɬɚ, ɩɪɨɦɵɜɚɥɢ ɞɢɷɬɢɥɨɜɵɦ ɷɮɢɪɨɦ, ɮɢɥɶɬɪɚɬ ɭɩɚɪɢɜɚɥɢ, ɨɫɬɚɬɨɤ ɤɪɢɫɬɚɥɥɢɡɨɜɚɥɢ ɢɡ ɫɦɟɫɢ ɩɟɧɬɚɧɷɮɢɪ 10:1. ȼɵɯɨɞ 271 ɦɝ (82 %), Rf 0.63; tɩɥ 82-84qɋ; ɍɎ-ɫɩɟɤɬɪ, Omax, ɧɦ (H): 227 (16600), 283 (18700), 357 (38000); 1ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15. Ɏɟɧɢɥɶɧɵɣ ɚɧɚɥɨɝ ɪɟɬɢɧɚɥɹ8 {(2ȿ,8E)-3,7-ɞɢɦɟɬɢɥ-9-ɮɟɧɢɥ-ɧɨɧɚ-2,4,6,8ɬɟɬɪɚɟɧ-1-ɚɥɶ}(4). Ʉ ɪɚɫɬɜɨɪɭ 120 ɦɝ (0,5 ɦɦɨɥɶ) ɩɨɥɢɟɧɚɥɹ (23) ɜ 15 ɦɥ ɝɟɤɫɚɧɚ ɩɪɢɛɚɜɥɹɥɢ 150 ɦɤɥ 0,37 Ɇ ɪɚɫɬɜɨɪɚ ɯɢɧɨɥɢɧɚ ɜ ɝɟɤɫɚɧɟ ɢ 50 ɦɝ ɤɚɬɚɥɢɡɚɬɨɪɚ Ʌɢɧɞɥɚɪɚ, ɝɢɞɪɢɪɨɜɚɥɢ ɜ ɡɚɦɤɧɭɬɨɣ ɫɢɫɬɟɦɟ ɩɪɢ ɩɟɪɟɦɟɲɢɜɚɧɢɢ. ɉɨɫɥɟ ɩɨɝɥɨɳɟɧɢɹ 12 ɦɥ ɜɨɞɨɪɨɞɚ (ɪɚɜɧɨɝɨ 105% ɨɬ ɪɚɫɱɟɬɧɨɝɨ) ɨɬɞɟɥɹɥɢ ɤɚɬɚɥɢɡɚɬɨɪ, ɩɪɨɦɵɜɚɥɢ ɟɝɨ ɝɟɤɫɚɧɨɦ (2u10 ɦɥ), ɮɢɥɶɬɪɚɬɵ ɨɛɴɟɞɢɧɹɥɢ ɢ ɩɪɨɜɨɞɢɥɢ ɬɟɪɦɨɢɡɨɦɟɪɢɡɚɰɢɸ, ɞɨɛɚɜɢɜ 1-3 ɦɝ ɢɨɞɚ ɩɪɢ ɤɢɩɟɧɢɢ ɜ ɚɬɦɨɫɮɟɪɟ ɚɪɝɨɧɚ ɜ ɬɟɱɟɧɢɢ 2 ɱ. Ɋɚɫɬɜɨɪɢɬɟɥɶ ɭɞɚɥɹɥɢ, all-E-ɢɡɨɦɟɪ ɮɟɧɢɥɶɧɨɝɨ ɚɧɚɥɨɝɚ ɪɟɬɢɧɚɥɹ (4) ɜɵɞɟɥɹɥɢ ɤɨɥɨɧɨɱɧɨɣ ɯɪɨɦɚɬɨɝɪɚɮɢɟɣ ɜ ɫɢɫɬɟɦɟ ɉɗɅ-ɷɮɢɪ ɫ ɝɪɚɞɢɟɧɬɨɦ ɩɨɫɥɟɞɧɟɝɨ ɨɬ 0 ɞɨ 50%. ȼɵɯɨɞ all-E-ɢɡɨɦɟɪɚ (4) 87 ɦɝ (73%), Rf 0,39; tɩɥ 121-124qɋ; ɍɎ-ɫɩɟɤɬɪ, Omax, ɧɦ (H): 280 (13700), 391(56700); 1

ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15. 13Z-ɢɡɨɦɟɪ (28) Rf 0,44; tɩɥ 73-75qɋ; ɍɎ-ɫɩɟɤɬɪ, Omax, ɧɦ (H):

283 (15400), 385 (55400); 1ɇ-əɆɊ ɫɦ. ɬɚɛɥ. 15.

______________________________ 8

ɫɦ. ɤɨɦɦɟɧɬɚɪɢɢ ɨ ɧɭɦɟɪɚɰɢɢ ɚɬɨɦɨɜ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɚɧɚɥɨɝɚ (4) ɢ ɟɝɨ ɩɪɟɞɲɟɫɬɜɟɧɧɢɤɨɜ (22), (23), ɚ ɬɚɤɠɟ (27), (28) ɜ ɫɧɨɫɤɟ ɧɚ ɫ. 59.

O

O

O

Ɏɨɪɦɭɥɚ

9

ɦ 2,07

ɫ 1,02

ɜɢɞ

G, ɦɞ

1.17

G, ɦɞ

ɬ

1.97

ɫ

ɦ

ɦ

ɬ

6.0

ɫ

-

ɞ

15.0

6.39

ɞ

15.0

7.21

ɞ

16,0

6,72

ɞ

16.0

6.30

ɞ

16,0

6,26

ɞ

16.5

6.18

8H 7.15

11H

7,12

ɞɞ

7.15

ɞɞ

7,15

ɞɞ

6.97

ɞɞ

ɞ

ɞɞ

11.0 11.0/15.0

6.42

ɞ

11,5 11,5/15,0

6,36

ɞ

11.5 11.5/15.0

6.25

ɞ

11,5 11,5/15,0

6,20

ɞ

12.0 12.0/15.4

6.20

10H

ɞ

15.0

6.53

ɞ

15,0

6,44

ɞ

15.0

6.40

ɞ

15,0

6,37

ɞ

15.4

6.37

12H

ɞ

8.0

5.87

ɞ

8,4

6,00

ɞ

8.0

5.98

ɞ

8,5

5,96

ɞ

8.0

5.98

14H

ɞ

8.0

9.97

ɞ

8,4

10,12

ɞ

8.0

10.12

ɞ

8,5

10,09

ɞ

8.0

10.12

15H

ɫ

-

1.91

ɫ

-

2,11

c

-

2.05

ɞ

1,0

2,02

ɫ

-

2.03

9-CH3

c

-

2.20

ɞ

1,5

2,35

c

-

2.33

ɞ

1,5

2,31

ɫ

-

2.33

13-CH3

Ɍɚɛɥɢɰɚ 15.

ɫɦ. ɤɨɦɦɟɧɬɚɪɢɢ ɨ ɧɭɦɟɪɚɰɢɢ ɚɬɨɦɨɜ ɩɨɥɢɟɧɨɜɨɣ ɰɟɩɢ ɚɧɚɥɨɝɚ (4) ɢ ɟɝɨ ɩɪɟɞɲɟɫɬɜɟɧɧɢɤɨɜ (22), (23), ɚ ɬɚɤɠɟ (27), (28) ɜ ɫɧɨɫɤɟ ɧɚ ɫ. 59.

ɜɢɞ

J

1.37

ɞ

ɦ

6.90

ɞ

16.0

6.37

ɞ

ɜɢɞ 2.27

c

-

1.83

ɫ

16,0

6,35

ɞ

16.5

6.36

16.0

1.37

-

ɞɬ

ɋ6ɇ5-: 7.32 ɢ 7,44

ɬ

6.5

2.50

ɞɬ

-

1,86

ɫ

-

1.72

7H

-

1.02

c

5,84

ɬ

6.5

2.03

4-CH2 5-CH3

J

G, ɦɞ

ɜɢɞ

6.5

1.85

ɫ

ɜɢɞ

-

ɞɞ

-

J

J

5,74

ɦ

-

1.61

3-CH2

4,0/1,5 9,5/4,0 9,5/1,5

-

1.48

-

1.03

1,1-CH3 2-CH2

J

G, ɦɞ

CHO G, ɦɞ

CHO

CHO

CHO

CHO

______________________________________

(5)

(4)

(3)

(2)

(1)

N

ɉɚɪɚɦɟɬɪɵ 1ɇ-əɆɊ ɫɩɟɤɬɪɨɜ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ9.

90

(28)

(27)

(23)

(22)

(17)

N

O

Ɏɨɪɦɭɥɚ

OH

O

O

OH

CHO

ɦ

ɜɢɞ

ɦ

ɞ

16.0

6.90

ɋ6ɇ5-: 7.39

G, ɦɞ -

ɞ

ɦ

ɜɢɞ

J

12,0

6,54

ɋ6ɇ5-: 7.2-7,3

G, ɦɞ -

ɞ

ɦ

ɜɢɞ

J

16,0

-

6,89

ɋ6ɇ5-: 7,32 ɢ 7,47

G, ɦɞ

J

ɞ

ɦ

ɜɢɞ

6,87

ɞ

15.5

6.32

16,0

c

-

0.94

7H

-

ɦ

-

1.90

4-CH2 5-CH3

J

ɋ6ɇ5-: 7.2-7,4

ɦ

c

ɜɢɞ

G, ɦɞ

-

-

1.75

1.40

1.16; 1.11

J

G, ɦɞ

3-CH2

1,1-CH3 2-CH2

91

ɞ

16,0

6,75

ɞ

12,0

6,20

ɞ

16,0

6,74

ɞ

16,0

6,62

ɞ

15.5

6.05

8H 7.10

11H

7,06

-

-

-

ɞɞ

ɞ

ɞɞ

11,5 11,5/15,0

6,39

ɫ

-

5,71

ɫ

-

5,79

ɫ

-

5,73

ɞ

11.2 11.2/15.0

6.23

10H

ɞ

15,0

7,37

-

-

-

ɞ

15.0

6.39

12H

ɞ

8,0

5,87

ɬ

6,5

5,84

ɞ. ɤɜ

8,0/1,5

6,14

ɬ ɤɜ

6,0/1,2

5,86

ɞ

8.0

5.98

14H

ɞ

8,0

10,23

ɞ

6,5

4,32

ɞ

8,0

10,09

ɞ

6,0

4,36

ɞ

8.0

10.11

15H

ɫ

-

2,11

ɫ

-

1,89

ɫ

-

2,17

ɫ

-

1,94

ɞ

1.2

2.00

9-CH3

ɞ

1,5

2,16

ɞ

1,5

1,51

ɞ

1,5

2,20

ɞ

1,2

2,12

ɞ

1.5

2.32

13-CH3


92

2. ɉɨɥɭɱɟɧɢɟ ɩɪɟɩɚɪɚɬɨɜ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ. ɉɪɢɝɨɬɨɜɥɟɧɢɟ ɪɨɫɬɨɜɵɯ ɫɪɟɞ ɞɥɹ ɝɚɥɨɛɚɤɬɟɪɢɣ. ȼ ɧɚɫɬɨɹɳɟɣ ɪɚɛɨɬɟ ɞɥɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɝɚɥɨɛɚɤɬɟɪɢɣ ɢɫɩɨɥɶɡɨɜɚɥɢ ɫɬɚɧɞɚɪɬɧɭɸ ɪɨɫɬɨɜɭɸ ɫɪɟɞɭ [188, 189], ɜɤɥɸɱɚɸɳɭɸ ɫɥɟɞɭɸɳɢɟ ɤɨɦɩɨɧɟɧɬɵ ɧɚ 1 ɥ ɜɨɞɵ: NaCl – 250 ɝ, MgSO47 ɇ2Ɉ – 20 ɝ, ɰɢɬɪɚɬ ɧɚɬɪɢɹ ɬɪɟɯɡɚɦɟɳɟɧɧɵɣ – 3 ɝ, KCl – 2 ɝ, ɞɪɨɠɠɟɜɨɣ ɷɤɫɬɪɚɤɬ – 2 ɝ, ɩɟɩɬɨɧ – 5 ɝ, ɝɥɢɰɟɪɢɧ – 1 ɦɥ. ɉɨɫɥɟ ɪɚɫɬɜɨɪɟɧɢɹ ɫɨɥɟɣ, ɞɪɨɠɠɟɜɨɝɨ ɷɤɫɬɪɚɤɬɚ ɢ ɩɟɩɬɨɧɚ ɞɨɛɚɜɥɹɥɢ ɝɥɢɰɟɪɢɧ, ɪɇ ɞɨɜɨɞɢɥɢ ɞɨ 7.2-7.4 ɬɢɬɪɨɜɚɧɢɟɦ ɪɚɫɬɜɨɪɨɦ ɄɈɇ (10 ɝ/100 ɦɥ). ɋɪɟɞɭ ɫɬɟɪɢɥɢɡɨɜɚɥɢ ɚɜɬɨɤɥɚɜɢɪɨɜɚɧɢɟɦ ɩɪɢ 121qɋ. Ⱦɥɹ ɩɪɢɝɨɬɨɜɥɟɧɢɹ ɬɜɺɪɞɵɯ ɫɪɟɞ ɞɨɛɚɜɥɹɥɢ 1,8 % ɚɝɚɪɨɡɵ ɢ ɧɚɝɪɟɜɚɥɢ ɧɚ ɜɨɞɹɧɨɣ ɛɚɧɟ ɞɨ ɪɚɫɬɜɨɪɟɧɢɹ ɚɝɚɪɨɡɵ. ɉɨɫɟɜɧɨɣ ɦɚɬɟɪɢɚɥ (ɢɧɨɤɭɥɹɬ). ɇɟɛɨɥɶɲɨɟ ɤɨɥɢɱɟɫɬɜɨ ɤɥɟɬɨɤ H. salinarum ɫ ɱɚɲɤɢ ɉɟɬɪɢ ɜɧɨɫɢɥɢ ɜ ɩɪɨɛɢɪɤɢ ɨɛɴɟɦɨɦ 30 ɦɥ, ɫɨɞɟɪɠɚɳɢɟ 5 ɦɥ ɫɪɟɞɵ ɢ ɤɭɥɶɬɢɜɢɪɨɜɚɥɢ ɧɚ ɜɪɚɳɚɬɟɥɶɧɨɣ ɤɚɱɚɥɤɟ 5 ɞɧ. ɩɪɢ 370ɋ. ȼ ɤɨɥɛɵ ɗɪɥɟɧɦɟɣɟɪɚ ɨɛɴɟɦɨɦ 750 ɦɥ, ɫɨɞɟɪɠɚɳɢɟ 200 ɦɥ ɫɪɟɞɵ, ɜɧɨɫɢɥɢ 5 ɦɥ ɩɨɫɟɜɧɨɣ ɤɭɥɶɬɭɪɵ ɢ ɩɪɨɞɨɥɠɚɥɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɟ ɧɚ ɜɪɚɳɚɬɟɥɶɧɨɣ ɤɚɱɚɥɤɟ (190 ɦɢɧ-1) ɩɪɢ 370ɋ. Ʉɭɥɶɬɢɜɢɪɨɜɚɧɢɟ ɝɚɥɨɛɚɤɬɟɪɢɣ ɩɪɨɜɨɞɢɥɨɫɶ 5 ɞɧɟɣ (ɞɨ ɡɚɜɟɪɲɟɧɢɹ ɷɤɫɩɨɧɟɧɰɢɚɥɶɧɨɣ ɮɚɡɵ). Ʉɭɥɶɬɢɜɢɪɨɜɚɧɢɟ Halobacterium salinarum ɜ ɮɟɪɦɟɧɬɟɪɟ. Ʉɭɥɶɬɢɜɢɪɨɜɚɧɢɟ H. salinarum ɨɫɭɳɟɫɬɜɥɹɥɢ ɜ 4 ɥ ɤɨɦɩɥɟɤɫɧɨɣ ɫɪɟɞɵ ɩɪɢ 34qɋ ɢɥɢ 37qɋ ɫ ɪɟɝɭɥɢɪɨɜɚɧɢɟɦ ɩɨɞɚɱɢ ɜɨɡɞɭɯɚ ɜ ɤɭɥɶɬɢɜɚɬɨɪ. ɉɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɭɫɬɚɧɨɜɥɟɧɧɵɣ ɪɚɫɯɨɞ ɜɨɡɞɭɯɚ ɫɨɫɬɚɜɥɹɥ 1.8 ɥ/ɦɢɧ. ɉɪɨɰɟɫɫ ɩɪɨɜɨɞɢɥɢ ɩɪɢ ɨɫɜɟɳɟɧɢɢ 4-ɦɹ ɥɚɦɩɚɦɢ ɞɧɟɜɧɨɝɨ ɫɜɟɬɚ ɅȾ-20. Ʉɸɜɟɬɚ 1 (ɫɦ. ɪɢɫ. 28) ɨɛɴɟɦɨɦ ɨɤɨɥɨ 6 ɥ ɫɞɟɥɚɧɚ ɢɡ ɨɪɝɫɬɟɤɥɚ ɢ ɢɦɟɟɬ ɪɚɡɦɟɪɵ ɜɵɫɨɬɚ 41 ɫɦ, ɞɥɢɧɚ 40 ɫɦ ɫ ɬɨɥɳɢɧɨɣ ɫɥɨɹ ɠɢɞɤɨɫɬɢ 3.5 ɫɦ, ɬɨɥɳɢɧɚ ɫɬɟɧɨɤ 1.2 ɫɦ. Ⱦɥɹ ɩɨɞɞɟɪɠɚɧɢɹ ɬɟɦɩɟɪɚɬɭɪɧɨɝɨ ɪɟɠɢɦɚ ɨɧɚ ɪɚɫɩɨɥɨɠɟɧɚ ɜ ɬɟɪɦɨɫɬɚɬɢɪɨɜɚɧɧɨɣ ɤɨɦɧɚɬɟ, ɫɧɚɛɠɟɧɚ ɚɷɪɢɪɭɸɳɢɦ ɭɫɬɪɨɣɫɬɜɨɦ, ɨɞɧɨɜɪɟɦɟɧɧɨ ɨɛɟɫɩɟɱɢɜɚɸɳɢɦ ɩɟɪɟɦɟɲɢɜɚɧɢɟ (ɜɨɡɞɭɯ, ɧɚɝɧɟɬɚɟɦɵɣ ɧɚɫɨɫɨɦ 2, ɩɪɨɯɨɞɢɬ ɞɥɹ ɩɨɞɞɟɪɠɚɧɢɹ ɫɬɟɪɢɥɶɧɨɫɬɢ ɱɟɪɟɡ ɜɨɡɞɭɲɧɵɣ ɮɢɥɶɬɪ 3, ɪɨɬɚɦɟɬɪ 4 ɢ ɩɨɫɬɭɩɚɟɬ ɜ ɤɸɜɟɬɭ 1, ɝɞɟ ɛɚɪɛɨɬɢɪɭɟɬɫɹ ɱɟɪɟɡ ɨɬɜɟɪɫɬɢɹ ɜɨɡɞɭɯɨɩɨɞɚɸɳɟɣ ɬɪɭɛɤɢ) ɢ ɨɫɜɟɳɚɟɬɫɹ 4 ɥɚɦɩɚɦɢ ɞɧɟɜɧɨɝɨ ɫɜɟɬɚ ɅȾ-20 5. ɂɦɟɸɬɫɹ ɨɬɜɨɞɵ ɞɥɹ ɨɬɛɨɪɚ ɩɪɨɛ ɢ ɫɥɢɜɚ Ʉɀ.

93

Ɋɢɫ. 28. Ʉɸɜɟɬɚ ɞɥɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ H. salinarum. 1 – ɤɸɜɟɬɚ, 2 – ɜɨɡɞɭɲɧɵɣ ɧɚɫɨɫ, 3 – ɮɢɥɶɬɪ ɞɥɹ ɫɬɟɪɢɥɢɡɚɰɢɢ ɜɨɡɞɭɯɚ, 4 – ɪɨɬɚɦɟɬɪ, 5 – ɥɚɦɩɵ ɞɧɟɜɧɨɝɨ ɫɜɟɬɚ ɅȾ-20. ɉɟɪɟɞ ɡɚɩɨɥɧɟɧɢɟɦ ɤɸɜɟɬɵ ɩɢɬɚɬɟɥɶɧɨɣ ɫɪɟɞɨɣ ɟɟ ɦɵɥɢ ɢ ɫɬɟɪɢɥɢɡɨɜɚɥɢ ɨɛɪɚɛɨɬɤɨɣ 96% ɫɩɢɪɬɨɦ (ɫɬɟɪɢɥɢɡɚɰɢɹ ɜ ɚɜɬɨɤɥɚɜɟ ɧɟ ɜɨɡɦɨɠɧɚ ɢɡ-ɡɚ ɦɚɬɟɪɢɚɥɚ, ɢɡ ɤɨɬɨɪɨɝɨ ɫɞɟɥɚɧɚ ɤɸɜɟɬɚ). Ⱦɥɹ ɨɛɟɫɩɟɱɟɧɢɹ ɚɫɟɩɬɢɱɟɫɤɢɯ ɭɫɥɨɜɢɣ ɫɬɟɪɢɥɢɡɨɜɚɥɢ ɬɚɤ ɠɟ ɢ ɜɫɟ ɤɨɦɦɭɧɢɤɚɰɢɢ, ɫɜɹɡɵɜɚɸɳɢɟ ɤɸɜɟɬɭ ɫ ɞɪɭɝɢɦɢ ɚɩɩɚɪɚɬɚɦɢ. ɉɨɫɥɟ ɡɚɩɨɥɧɟɧɢɹ 4-ɦɹ ɥ ɫɬɟɪɢɥɶɧɨɣ ɪɨɫɬɨɜɨɣ ɫɪɟɞɵ (ɨɛɴɺɦ ɧɟ ɩɪɟɜɵɲɚɟɬ 70% ɨɛɳɟɝɨ ɨɛɴɺɦɚ ɤɸɜɟɬɵ), ɫɬɟɪɢɥɶɧɨ ɜɜɨɞɢɥɢ 400 ɦɥ (10% ɨɛ/ɨɛ) ɩɨɫɟɜɧɨɝɨ ɦɚɬɟɪɢɚɥɚ. ɉɟɪɟɞ ɜɧɟɫɟɧɢɟɦ ɢɧɨɤɭɥɹɬɚ ɬɟɦɩɟɪɚɬɭɪɭ ɢ ɪɇ ɩɢɬɚɬɟɥɶɧɨɣ ɫɪɟɞɵ ɞɨɜɨɞɢɥɢ ɞɨ ɨɩɬɢɦɚɥɶɧɵɯ ɡɧɚɱɟɧɢɣ ɞɥɹ H. salinarum. Ʉɭɥɶɬɢɜɢɪɨɜɚɥɢ ɩɪɢ ɩɨɫɬɨɹɧɧɨɦ ɨɫɜɟɳɟɧɢɢ ɞɨ ɞɨɫɬɢɠɟɧɢɹ ɩɨɡɞɧɟɣ ɫɬɚɰɢɨɧɚɪɧɨɣ ɮɚɡɵ, ɪɟɝɢɫɬɪɢɪɭɹ ɢɡɦɟɧɟɧɢɟ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɤɭɥɶɬɭɪɵ ɩɨ ɩɟɪɢɨɞɢɱɟɫɤɢ ɨɬɛɢɪɚɟɦɵɦ ɩɪɨɛɚɦ. ɍɫɥɨɜɢɹ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɢ ɜɵɯɨɞ ɛɢɨɦɚɫɫɵ ɢ ȻɊ ɫ ɥɢɬɪɚ Ʉɀ ɩɪɢɜɟɞɟɧɵ ɜ ɬɚɛɥ. 12 ɬɟɨɪɟɬɢɱɟɫɤɨɣ ɱɚɫɬɢ ɧɚ ɫ. 77. ɉɪɢ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɢ ɲɬɚɦɦɚ JW5 ɫ ɞɨɛɚɜɥɟɧɢɟɦ ɪɟɬɢɧɨɢɞɨɜ, ɪɟɬɢɧɚɥɶ (1) ɢɥɢ 3,4-ɞɢɞɟɝɢɞɪɨɪɟɬɢɧɚɥɶ (2) ɞɨɛɚɜɥɹɥɢ ɜ ɜɢɞɟ ɷɬɚɧɨɥɶɧɨɝɨ ɪɚɫɬɜɨɪɚ ɧɚ 7 (200 ɦɤɥ), 8 (200 ɦɤɥ), 9 (400 ɦɤɥ) ɢ 10 (400 ɦɤɥ) (ɡɚ 2-3 ɱ. ɞɨ ɨɫɚɠɞɟɧɢɹ ɛɢɨɦɚɫɫɵ) ɞɧɢ. ɂɫɩɨɥɶɡɨɜɚɥɢ ɪɚɫɬɜɨɪ ɪɟɬɢɧɨɢɞɨɜ ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ 0.016 ɦɦɨɥɶ/ɦɥ. Ȼɢɨɦɚɫɫɭ ɢ ɩɪɨɛɵ, ɨɬɛɢɪɚɟɦɵɟ ɞɥɹ ɩɨɫɬɪɨɟɧɢɹ ɤɪɢɜɵɯ ɪɨɫɬɚ, ɩɪɢ ɧɟɨɛɯɨɞɢɦɨɫɬɢ ɯɪɚɧɢɥɢ ɩɪɢ 4qC . Ɉɬɛɨɪ ɩɪɨɛ ɢɡ ɪɚɫɬɭɳɢɯ ɤɭɥɶɬɭɪ ɝɚɥɨɛɚɤɬɟɪɢɣ. Ɉɬɛɨɪ ɩɪɨɛ ɨɛɴɟɦɨɦ ɨɤɨɥɨ 12 ɦɥ ɩɪɨɜɨɞɢɥɢ ɧɟ ɱɚɳɟ 1 ɪɚɡɚ ɜ ɞɟɧɶ (ɧɟ ɦɟɧɟɟ 6 ɪɚɡ ɡɚ ɩɟɪɢɨɞ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ). ɉɨɫɥɟ ɨɩɪɟɞɟɥɟɧɢɹ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ, ɩɪɨɛɵ ɨɛɴɟɦɨɦ 10 ɦɥ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɥɢ ɞɥɹ ɨɫɚɠɞɟɧɢɹ ɤɥɟɬɨɤ. Ɉɫɚɞɨɤ ɢɫɩɨɥɶɡɨɜɚɥɢ ɞɥɹ ɨɩɪɟɞɟɥɟɧɢɹ ɫɨɞɟɪɠɚɧɢɹ ȻɊ.

94 Ɉɰɟɧɤɚ ɫɨɞɟɪɠɚɧɢɹ ȻɊ ɜ ɛɢɨɦɚɫɫɟ ɝɚɥɨɛɚɤɬɟɪɢɣ. Ɉɰɟɧɤɭ ɫɨɞɟɪɠɚɧɢɹ ȻɊ ɜ ɛɢɨɦɚɫɫɟ ɝɚɥɨɛɚɤɬɟɪɢɣ ɨɫɭɳɟɫɬɜɥɹɥɢ ɩɨ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɨɦɭ ɦɟɬɨɞɭ [188], ɩɪɟɞɥɨɠɟɧɧɨɦɭ ɞɥɹ ɨɩɪɟɞɟɥɟɧɢɹ ȻɊ ɜ ɞɟɮɟɤɬɧɵɯ ɩɨ ɫɢɧɬɟɡɭ ɤɚɪɨɬɢɧɨɢɞɨɜ ɲɬɚɦɦɚɯ H. salinarum. Ʉ ɤɥɟɬɤɚɦ, ɩɨɥɭɱɟɧɧɵɦ ɨɫɚɠɞɟɧɢɟɦ ɢɡ ɩɪɨɛɵ ɨɛɴɟɦɨɦ 10 ɦɥ, ɞɨɛɚɜɥɹɥɢ 3 ɦɥ ɛɢɞɢɫɬɢɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ, ɫɨɞɟɪɠɚɳɟɣ 1 ɦɝ ȾɇɄɚɡɵ ɧɚ 100 ɦɥ. Ɉɫɭɳɟɫɬɜɥɹɥɢ ɥɢɡɢɫ ɤɥɟɬɨɤ ɚɤɬɢɜɧɵɦ ɩɢɩɟɬɢɪɨɜɚɧɢɟɦ. ɑɟɪɟɡ 3 ɱ. (ɩɨɫɥɟ ɡɚɜɟɪɲɟɧɢɹ ɝɢɞɪɨɥɢɡɚ ȾɇɄ), ɨɫɚɠɞɚɥɢ ɤɪɭɩɧɵɟ ɤɥɟɬɨɱɧɵɟ ɮɪɚɝɦɟɧɬɵ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɟɦ ɩɪɢ 10000 g, 10 ɦɢɧ. Ɉɩɪɟɞɟɥɹɥɢ ɨɩɬɢɱɟɫɤɭɸ ɩɥɨɬɧɨɫɬɶ ɫɭɩɟɪɧɚɬɚɧɬɚ ɩɪɢ 570 ɧɦ. ɉɪɢ ɨɩɪɟɞɟɥɟɧɢɢ ɫɨɞɟɪɠɚɧɢɹ ɛɚɤɬɟɪɢɨɨɩɫɢɧɚ ɜ ɞɟɮɟɤɬɧɨɦ ɩɨ ɪɟɬɢɧɚɥɸ ɲɬɚɦɦɟ JW5, ɨɫɚɠɞɟɧɧɵɟ ɢɡ ɩɪɨɛɵ ɤɥɟɬɤɢ ɩɪɟɞɜɚɪɢɬɟɥɶɧɨ ɫɭɫɩɟɧɞɢɪɨɜɚɥɢ ɜ 10 ɦɥ ɫɨɥɟɜɨɣ ɫɪɟɞɵ (ɚɧɚɥɨɝ ɪɨɫɬɨɜɨɣ ɫɪɟɞɵ, ɧɨ ɫɨɞɟɪɠɢɬ ɬɨɥɶɤɨ ɫɨɥɢ), ɢ ɩɪɨɜɨɞɢɥɢ ɪɟɝɟɧɟɪɚɰɢɸ ȻɊ ɢɡ ɛɚɤɬɟɪɢɨɨɩɫɢɧɚ ɢ ɪɟɬɢɧɚɥɹ (1) in vivo, ɞɨɛɚɜɢɜ ɤ ɤɥɟɬɨɱɧɨɣ ɫɭɫɩɟɧɡɢɢ 10 ɦɤɥ ɪɚɫɬɜɨɪɚ ɪɟɬɢɧɚɥɹ (1) (ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ 2 ɦɝ/ɦɥ). ɑɟɪɟɡ 2 ɱ ɨɫɭɳɟɫɬɜɥɹɥɢ ɨɩɪɟɞɟɥɟɧɢɟ ȻɊ ɩɨ ɨɩɢɫɚɧɧɨɣ ɜɵɲɟ ɦɟɬɨɞɢɤɟ. ȼɵɞɟɥɟɧɢɟ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ [1, 35]. Ɉɫɚɞɨɤ ɤɥɟɬɨɤ, ɩɨɥɭɱɟɧɧɵɣ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɟɦ 4 ɥ Ʉɀ ɫɭɫɩɟɧɞɢɪɨɜɚɥɢ ɜ 100 ɦɥ ɛɢɞɢɫɬɢɥɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ, ɜ ɤɨɬɨɪɭɸ ɩɪɟɞɜɚɪɢɬɟɥɶɧɨ ɞɨɛɚɜɢɥɢ ȾɇɄɚɡɭ I (1 ɦɝ ɧɚ 100 ɦɥ). ɉɟɪɟɦɟɲɢɜɚɥɢ ɩɪɢ 4 0C ɜ ɬɟɱɟɧɢɟ 2 ɱ. Ɂɚɬɟɦ ɩɪɢ 4 0C ɞɢɚɥɢɡɨɜɚɥɢ ɩɨɥɭɱɟɧɧɭɸ ɫɭɫɩɟɧɡɢɸ 17 ɱ ɩɪɨɬɢɜ 5 ɥ ɛɢɞɢɫɬɢɥɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ. Ⱦɥɹ ɨɫɚɠɞɟɧɢɹ ɤɪɭɩɧɵɯ ɤɥɟɬɨɱɧɵɯ ɮɪɚɝɦɟɧɬɨɜ ɫɭɫɩɟɧɡɢɸ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɥɢ ɩɪɢ 4 0C ɩɪɢ 10000g 10 ɦɢɧ., ɨɫɚɞɨɤ ɨɬɛɪɚɫɵɜɚɥɢ. ɋɨɞɟɪɠɚɳɢɟɫɹ ɜ ɫɭɩɟɪɧɚɬɚɧɬɟ ɉɆ ɩɟɪɟɨɫɚɠɞɚɥɢ 6 ɪɚɡ ɢɡ ɛɢɞɢɫɬɢɥɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɟɦ ɩɪɢ 4 0C ɜ ɬɟɱɟɧɢɟ 70-120 ɦɢɧ. (ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɲɬɚɦɦɚ ɢ ɱɢɫɬɨɬɵ ɩɪɟɩɚɪɚɬɚ) ɩɪɢ 50000g. Ƚɨɬɨɜɵɟ ɩɪɟɩɚɪɚɬɵ ɦɟɦɛɪɚɧ ɯɪɚɧɢɥɢ ɩɪɢ 4 0C ɜ ɜɢɞɟ ɜɨɞɧɨɣ ɫɭɫɩɟɧɡɢɢ ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ ɨɤɨɥɨ 10 ɦɝ ɛɟɥɤɚ ɜ 1 ɦɥ. Ɉɩɪɟɞɟɥɟɧɢɟ ɦɚɫɫɵ ȻɊ ɩɨ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ. ɋɜɟɬɨɜɭɸ ɚɞɚɩɬɚɰɢɸ ɩɪɟɩɚɪɚɬɨɜ ȻɊ ɨɫɭɳɟɫɬɜɥɹɥɢ ɨɫɜɟɳɟɧɢɟɦ ɜ ɬɟɱɟɧɢɢ 2 ɦɢɧ. ɫɜɟɬɨɦ ɪɬɭɬɧɨɣ ɥɚɦɩɵ, ɩɪɨɩɭɳɟɧɧɵɦ ɱɟɪɟɡ ɬɟɩɥɨɜɨɣ ɮɢɥɶɬɪ ɢ ɫɜɟɬɨɮɢɥɶɬɪ ɀɋ18. ɂɡɦɟɪɹɥɢ ɨɩɬɢɱɟɫɤɭɸ ɩɥɨɬɧɨɫɬɶ ɜ ɦɚɤɫɢɦɭɦɟ ɩɨɝɥɨɳɟɧɢɹ ɯɪɨɦɨɮɨɪɚ ɜ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɨɣ ɮɨɪɦɟ ȻɊ ɢ ɪɚɫɫɱɢɬɵɜɚɥɢ ɟɝɨ ɦɚɫɫɭ ɩɨ ɫɥɟɞɭɸɳɟɣ ɮɨɪɦɭɥɟ:

95 m ɉɆ

D Mr V ɪ ɪɚ Vɤɸɜ H ȻɊ Vɩɪɨɛɵ

0,4206

D570 V ɪ ɪɚ Vɤɸɜ

ɢɥɢ = 0,2409

Vɩɪɨɛɵ

, ɦɝ

D280 V ɪ ɪɚ Vɤɸɜ Vɩɪɨɛɵ

, ɦɝ

ɝɞɟ D – ɨɩɬɢɱɟɫɤɚɹ ɩɥɨɬɧɨɫɬɶ ɪɚɫɬɜɨɪɚ ɜ ɦɚɤɫɢɦɭɦɟ ɩɨɝɥɨɳɟɧɢɹ ɩɢɝɦɟɧɬɚ; Mr – ɦɨɥɟɤɭɥɹɪɧɚɹ ɦɚɫɫɚ ȻɊ (26700); Vɪ-ɪɚ – ɨɛɴɟɦ ɫɭɫɩɟɧɡɢɢ ȻɊ, ɦɥ; Vɤɸɜ – ɨɛɴɟɦ ɪɚɫɬɜɨɪɚ ɜ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɢɱɟɫɤɨɣ ɤɸɜɟɬɟ, ɦɥ; Vɩɪɨɛɵ – ɨɛɴɟɦ ɩɪɨɛɵ, ɜɧɟɫɟɧɧɨɣ ɜ ɤɸɜɟɬɭ, ɦɥ; HȻɊ – ɤɨɷɮɮɢɰɢɟɧɬ ɦɨɥɹɪɧɨɝɨ ɩɨɝɚɲɟɧɢɹ ȻɊ, ɩɪɢɧɹɬ ɪɚɜɧɵɦ 63 000 Ɇ-1ɫɦ-1 ɩɪɢ 570 ɧɦ ɢ 110 000 Ɇ-1ɫɦ-1 ɩɪɢ 270 ɧɦ. ɉɪɢ ɪɚɫɱɟɬɟ ɦɚɫɫɵ ȻɊ ɩɨ ɛɟɥɤɨɜɨɣ ɩɨɥɨɫɟ ɩɨɝɥɨɳɟɧɢɹ (270 ɧɦ) ɫɥɟɞɭɟɬ ɭɱɢɬɵɜɚɬɶ, ɱɬɨ ɜ ɷɬɨɣ ɨɛɥɚɫɬɢ ɡɚɦɟɬɧɵɣ ɜɤɥɚɞ ɜ ɨɩɬɢɱɟɫɤɭɸ ɩɥɨɬɧɨɫɬɶ ɦɨɠɟɬ ɜɧɨɫɢɬɶ ɫɜɟɬɨɪɚɫɫɟɹɧɢɟ, ɱɬɨ ɩɪɢɜɨɞɢɬ ɤ ɩɨɥɭɱɟɧɢɸ ɧɟɫɤɨɥɶɤɨ ɡɚɜɵɲɟɧɧɨɝɨ ɡɧɚɱɟɧɢɹ.

3. ɉɨɥɭɱɟɧɢɟ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. ɉɪɢɝɨɬɨɜɥɟɧɢɟ ɛɭɮɟɪɧɵɯ ɪɚɫɬɜɨɪɨɜ ɞɥɹ ɢɫɫɥɟɞɨɜɚɧɢɹ ɉɆ Ȼɭɮɟɪ Ⱥ: 1 Ɇ NH2OH, ɪɇ 7.0. Ɋɚɫɬɜɨɪɹɥɢ 34,75 ɝ ɝɢɞɪɨɤɫɢɥɚɦɢɧɚ ɝɢɞɪɨɯɥɨɪɢɞɚ ɜ 300 ɦɥ ɛɢɞɢɫɬɢɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ, ɞɨɜɨɞɢɥɢ ɪɇ ɞɨ ɡɧɚɱɟɧɢɹ 7.0 ɩɨɫɬɟɩɟɧɧɵɦ ɩɪɢɛɚɜɥɟɧɢɟɦ 10% ɜɨɞɧɨɝɨ ɪɚɫɬɜɨɪɚ ɝɢɞɪɨɤɫɢɞɚ ɧɚɬɪɢɹ ɩɪɢ ɩɟɪɟɦɟɲɢɜɚɧɢɢ, ɡɚɬɟɦ ɞɨɜɨɞɢɥɢ ɨɛɴɟɦ ɪɚɫɬɜɨɪɚ ɞɨ 500 ɦɥ. Ȼɭɮɟɪ ȼ: 5 ɦɦɨɥɶ MES, 100 ɦɆ NaCl, ɪɇ 6.0. Ɋɚɫɬɜɨɪɹɥɢ 533 ɦɝ ɦɨɪɮɨɥɢɧɷɬɚɧɫɭɥɶɮɨɤɢɫɥɨɬɵ ɢ 2,92 ɝ ɯɥɨɪɢɞɚ ɧɚɬɪɢɹ ɜ 300 ɦɥ ɛɢɞɢɫɬɢɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ, ɞɨɜɨɞɢɥɢ ɪɇ ɞɨ ɡɧɚɱɟɧɢɹ 6.0 ɩɨɫɬɟɩɟɧɧɵɦ ɩɪɢɛɚɜɥɟɧɢɟɦ 1 Ɇ ɪɚɫɬɜɨɪɚ NaOH, ɡɚɬɟɦ ɞɨɜɨɞɢɥɢ ɨɛɴɟɦ ɪɚɫɬɜɨɪɚ ɞɨ 500 ɦɥ. Ɉɛɟɫɰɜɟɱɢɜɚɧɢɟ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ. 100 ɦɥ ɫɭɫɩɟɧɡɢɢ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ ɜ ɛɭɮɟɪɟ Ⱥ (1 Ɇ NH2OH, ɪɇ 7.0) ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ ɛɟɥɤɚ ɨɤɨɥɨ 0,5 ɦɝ/ɦɥ ɩɪɢ ɨɯɥɚɠɞɟɧɢɢ (7-12qɋ) ɢ ɦɟɞɥɟɧɧɨɦ ɩɟɪɟɦɟɲɢɜɚɧɢɢ ɨɫɜɟɳɚɥɢ ɜ ɬɟɱɟɧɢɟ |17-26 ɱ ɫɜɟɬɨɦ 2 ɝɚɥɨɝɟɧɨɜɵɯ ɥɚɦɩ ɦɨɳɧɨɫɬɶɸ 500 ȼɬ ɤɚɠɞɚɹ, ɩɪɨɩɭɳɟɧɧɵɦ ɱɟɪɟɡ ɬɟɩɥɨɜɨɣ ɮɢɥɶɬɪ (0,2% ɪɚɫɬɜɨɪ ɫɭɥɶɮɚɬɚ ɦɟɞɢ (II) ɜ ɜɨɞɟ, ɜ ɤɨɥɛɚɯ ɨɛɴɟɦɨɦ 1 ɥ). ɋɬɟɩɟɧɶ ɩɪɨɯɨɠɞɟɧɢɹ ɩɪɨɰɟɫɫɚ ɤɨɧɬɪɨɥɢɪɨɜɚɥɢ ɩɨ ɭɦɟɧɶɲɟɧɢɸ ɩɨɝɥɨɳɟɧɢɹ ɜ ɨɛɥɚɫɬɢ 563 ɧɦ. Ɉɛɟɫɰɜɟɱɟɧɧɵɟ ɦɟɦɛɪɚɧɵ ɨɫɚɠɞɚɥɢ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɟɦ ɩɪɢ 50000g, 20 ɦɢɧ, 4qɋ ɢ 4 ɪɚɡɚ ɩɟɪɟɨɫɚɠɞɚɥɢ ɢɡ ɛɢɞɢɫɬɢɥɥɢɪɨɜɚɧɧɨɣ ɜɨɞɵ. ɉɨɫɥɟ ɩɨɫɥɟɞɧɟɝɨ ɰɟɧɬɪɢɮɭɝɢɪɨɜɚɧɢɹ ɨɫɚɞɨɤ ɫɭɫɩɟɧɞɢɪɨɜɚɥɢ ɜ ɛɭɮɟɪɟ ȼ, ɱɬɨɛɵ ɤɨɧɟɱɧɚɹ ɤɨɧɰɟɧɬɪɚɰɢɹ ɫɨɫɬɚɜɥɹɥɚ ɨɤɨɥɨ 2,5 ɦɝ/ɦɥ.

96 Ɍɨɱɧɭɸ ɤɨɧɰɟɧɬɪɚɰɢɸ «ɚɤɬɢɜɧɨɝɨ» ȻɈ ɨɩɪɟɞɟɥɹɥɢ ɩɨɫɥɟ ɜɡɚɢɦɨɞɟɣɫɬɜɢɹ ɫ ɪɟɬɢɧɚɥɟɦ (1) ɩɨ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɪɟɝɟɧɟɪɢɪɨɜɚɧɧɨɝɨ ȻɊ (ɫɦ. ɮɨɪɦɭɥɭ ɧɚ ɫ. 95). ȼɡɚɢɦɨɞɟɣɫɬɜɢɟ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ ɫ ɚɩɨɦɟɦɛɪɚɧɚɦɢ. 1) Ɉɰɟɧɤɚ ɫɤɨɪɨɫɬɢ ɢ ɫɬɟɩɟɧɢ ɪɟɝɟɧɟɪɚɰɢɢ ȻɊ ɢɡ ɪɟɬɢɧɚɥɹ (1) ɢ ȻɈ. Ʉ 100 ɦɤɥ ɫɭɫɩɟɧɡɢɢ ɚɩɨɦɟɦɛɪɚɧ ɜ ɛɭɮɟɪɟ ȼ (ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ ɛɟɥɤɚ ɨɤɨɥɨ 2.5 ɦɝ/ɦɥ) ɞɨɛɚɜɥɹɥɢ 3 ɦɥ ɛɭɮɟɪɚ ȼ ɢ 1,6 ɦɤɥ ɪɚɫɬɜɨɪɚ ɪɟɬɢɧɚɥɹ (1) (ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ 2 ɦɝ/ɦɥ), ɷɧɟɪɝɢɱɧɨ ɩɟɪɟɦɟɲɢɜɚɥɢ ɢ ɪɟɝɢɫɬɪɢɪɨɜɚɥɢ ɫɩɟɤɬɪɵ ɩɨɝɥɨɳɟɧɢɹ ɜ ɢɧɬɟɪɜɚɥɟ 250-700 ɧɦ ɨɬɧɨɫɢɬɟɥɶɧɨ ɛɭɮɟɪɚ (ɞɥɹ ɨɩɪɟɞɟɥɟɧɢɹ ɤɨɧɰɟɧɬɪɚɰɢɢ ɚɤɬɢɜɧɨɝɨ ɛɟɥɤɚ ɢ ɫɬɟɩɟɧɢ ɪɟɝɟɧɟɪɚɰɢɢ) ɢ ɚɩɨɦɟɦɛɪɚɧ (ɞɢɮɮɟɪɟɧɰɢɚɥɶɧɵɣ ɫɩɟɤɬɪ) ɱɟɪɟɡ 1, 4, 8, 15, 30 ɢ 60 ɦɢɧ. ɩɨɫɥɟ ɞɨɛɚɜɥɟɧɢɹ ɪɟɬɢɧɚɥɹ (1), ɡɚɬɟɦ – ɩɨɫɥɟ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ ɩɪɟɩɚɪɚɬɚ (ɨɫɜɟɳɟɧɢɟɦ ɜ ɬɟɱɟɧɢɟ 120 ɫ). Ʉɨɧɰɟɧɬɪɚɰɢɸ ɚɤɬɢɜɧɨɝɨ ɛɟɥɤɚ ɨɩɪɟɞɟɥɹɥɢ ɩɨ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɨɝɨ ɩɪɟɩɚɪɚɬɚ ɜ ɦɚɤɫɢɦɭɦɟ ɩɨɝɥɨɳɟɧɢɹ ɯɪɨɦɨɮɨɪɚ (ɫɦ. ɮɨɪɦɭɥɭ ɧɚ ɫɬɪ. 95). ɋɬɟɩɟɧɶ ɪɟɝɟɧɟɪɚɰɢɢ: (D1/D2)*100%, ɝɞɟ D1 ɢ D2– ɩɨɝɥɨɳɟɧɢɟ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɵɯ ɮɨɪɦ ɪɟɝɟɧɟɪɢɪɨɜɚɧɧɨɝɨ ɪɟɬɢɧɚɥɟɦ (1) ɢ ɢɫɯɨɞɧɨɝɨ ȻɊ, ɩɪɢ ɭɫɥɨɜɢɢ, ɱɬɨ ɜ ɦɚɤɫɢɦɭɦɟ ɩɨɝɥɨɳɟɧɢɹ ɛɟɥɤɚ (276 ɧɦ) ɨɩɬɢɱɟɫɤɚɹ ɩɥɨɬɧɨɫɬɶ ɢɫɯɨɞɧɨɝɨ ȻɊ ɪɚɜɧɚ ɨɩɬɢɱɟɫɤɨɣ ɩɥɨɬɧɨɫɬɢ ɚɩɨɦɟɦɛɪɚɧ ɞɨ ɪɟɝɟɧɟɪɚɰɢɢ. 2) Ɉɩɪɟɞɟɥɟɧɢɟ ɤɨɥɢɱɟɫɬɜɚ ɪɟɬɢɧɚɥɹ (1), ɧɟɨɛɯɨɞɢɦɨɝɨ ɞɥɹ ɪɟɤɨɧɫɬɪɭɤɰɢɢ ȻɊ ɢɡ ɢɫɫɥɟɞɭɟɦɨɝɨ ɩɪɟɩɚɪɚɬɚ ɚɩɨɦɟɦɛɪɚɧ. Q ɪɟɬ

Q ȻɊ

D570 V ɪ ɪɚ Vɤɸɜ H ȻɊ Vɩɪɨɛɵ

1,6 10 5

D570 V ɪ ɪɚ Vɤɸɜ Vɩɪɨɛɵ

, ɦɦɨɥɶ

ɝɞɟ D570 – ɨɩɬɢɱɟɫɤɚɹ ɩɥɨɬɧɨɫɬɶ ɪɚɫɬɜɨɪɚ ɜ ɦɚɤɫɢɦɭɦɟ ɩɨɝɥɨɳɟɧɢɹ ɫɜɟɬɨɚɞɚɩɬɢɪɨɜɚɧɧɨɝɨ ȻɊ, ɩɨɥɭɱɟɧɧɨɝɨ ɪɟɤɨɧɫɬɪɭɤɰɢɟɣ ɢɡ ȻɈ ɢ ɪɟɬɢɧɚɥɹ (1); Vɪ-ɪɚ – ɨɛɴɟɦ ɫɭɫɩɟɧɡɢɢ ɚɩɨɦɟɦɛɪɚɧ, ɦɥ; Vɤɸɜ – ɨɛɴɟɦ ɪɚɫɬɜɨɪɚ ɜ ɫɩɟɤɬɪɨɮɨɬɨɦɟɬɪɢɱɟɫɤɨɣ ɤɸɜɟɬɟ, ɦɥ; Vɩɪɨɛɵ – ɨɛɴɟɦ ɩɪɨɛɵ, ɜɧɟɫɟɧɧɨɣ ɜ ɤɸɜɟɬɭ, ɦɥ; HȻɊ – ɤɨɷɮɮɢɰɢɟɧɬ ɦɨɥɹɪɧɨɝɨ ɩɨɝɚɲɟɧɢɹ ȻɊ, ɩɪɢɧɹɬ ɪɚɜɧɵɦ 63 000 Ɇ-1ɫɦ-1. 3) ɉɨɥɭɱɟɧɢɟ ɚɧɚɥɨɝɚ ȻɊ ɜɡɚɢɦɨɞɟɣɫɬɜɢɟɦ ɚɧɚɥɨɝɚ ɪɟɬɢɧɚɥɹ ɫ ȻɈ. Ⱦɥɹ ɪɟɝɟɧɟɪɚɰɢɢ ɢɫɩɨɥɶɡɨɜɚɥɢ ɦɟɬɚɧɨɥɶɧɵɣ ɪɚɫɬɜɨɪ ɚɧɚɥɨɝɨɜ ɪɟɬɢɧɚɥɹ (2)-(5) ɫ ɤɨɧɰɟɧɬɪɚɰɢɟɣ 4 ɦɝ/ɦɥ. Ʉ ɫɭɫɩɟɧɡɢɢ ɚɩɨɦɟɦɛɪɚɧ ɞɨɛɚɜɥɹɥɢ ɚɧɚɥɨɝ ɪɟɬɢɧɚɥɹ ɫ ɢɡɛɵɬɤɨɦ 10% ɨɬ ɪɚɫɫɱɢɬɚɧɧɨɝɨ (ɫɦ. ɩ. 2) ɢ ɪɟɝɢɫɬɪɢɪɨɜɚɥɢ ɫɩɟɤɬɪɵ ɩɨɝɥɨɳɟɧɢɹ ɨɬɧɨɫɢɬɟɥɶɧɨ ɛɭɮɟɪɚ ɢ ɨɬɧɨɫɢɬɟɥɶɧɨ ɫɭɫɩɟɧɡɢɢ ɚɩɨɦɟɦɛɪɚɧ ɜ ɢɧɬɟɪɜɚɥɟ 250-700 ɧɦ ɱɟɪɟɡ 1, 4, 8, 15, 30 ɢ 60 ɦɢɧ ɩɨɫɥɟ ɞɨɛɚɜɥɟɧɢɹ ɚɧɚɥɨɝɚ ɪɟɬɢɧɚɥɹ, ɡɚɬɟɦ – ɩɨɫɥɟ ɫɜɟɬɨɜɨɣ ɚɞɚɩɬɚɰɢɢ ɩɪɟɩɚɪɚɬɚ (ɨɫɜɟɳɟɧɢɟɦ ɜ ɬɟɱɟɧɢɢ 120 ɫ).

97

ȼɕȼɈȾɕ. 1.

ɉɪɟɞɥɨɠɟɧ ɭɞɨɛɧɵɣ ɦɟɬɨɞ ɨɞɧɨɜɪɟɦɟɧɧɨɝɨ ɫɢɧɬɟɡɚ 5,6-ɞɢɨɤɫɨ-5,6ɫɟɤɨɪɟɬɢɧɚɥɹ, 5,6-ɞɢɝɢɞɪɨ-5,6-ɷɩɨɤɫɢɪɟɬɢɧɚɥɹ ɢ 4-ɨɤɫɨɪɟɬɢɧɚɥɹ ɨɤɢɫɥɟɧɢɟɦ ɪɟɬɢɧɚɥɹ.

2.

ɉɨɥɭɱɟɧ ɪɹɞ ȺȻɊ ɫ ɦɨɞɢɮɢɤɚɰɢɟɣ ɤɚɤ ɯɪɨɦɨɮɨɪɚ ɬɚɤ ɢ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ.

3.

Ɉɫɭɳɟɫɬɜɥɟɧɨ ɤɨɦɩɥɟɤɫɧɨɟ ɢɫɫɥɟɞɨɜɚɧɢɟ ȺȻɊ (ɨɩɪɟɞɟɥɟɧɵ ɫɩɟɤɬɪɚɥɶɧɵɟ ɫɜɨɣɫɬɜɚ, ɭɫɬɨɣɱɢɜɨɫɬɶ ɩɪɢ ɞɥɢɬɟɥɶɧɨɦ ɯɪɚɧɟɧɢɢ, ɫɤɨɪɨɫɬɶ ɪɟɝɟɧɟɪɚɰɢɢ ɢ ɨɫɨɛɟɧɧɨɫɬɢ ɮɨɬɨɰɢɤɥɚ).

4.

ɉɪɨɜɟɞɟɧɚ ɨɩɬɢɦɢɡɚɰɢɹ ɭɫɥɨɜɢɣ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ H. salinarum ɲɬɚɦɦɨɜ ET1001, JW5 ɢ D96N ɫ ɰɟɥɶɸ ɭɜɟɥɢɱɟɧɢɹ ɜɵɯɨɞɚ ȻɊ.

5.

ɉɨɤɚɡɚɧɚ ɩɟɪɫɩɟɤɬɢɜɧɨɫɬɶ ɢɫɩɨɥɶɡɨɜɚɧɢɹ ɲɬɚɦɦɚ JW5 ɞɥɹ ɩɨɥɭɱɟɧɢɹ ȺȻɊ ɜ ɬɟɯɧɨɥɨɝɢɱɟɫɤɢɯ ɰɟɥɹɯ.

6.

ɉɭɬɟɦ ɫɪɚɜɧɟɧɢɹ ɫɜɨɣɫɬɜ ȺȻɊ ɫ ɩɪɢɪɨɞɧɨɣ ɚɦɢɧɨɤɢɫɥɨɬɧɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ ɢ ɫ ɡɚɦɟɧɨɣ ɚɫɩɚɪɚɝɢɧɨɜɨɣ ɤɢɫɥɨɬɵ-96 ɧɚ ɚɫɩɚɪɚɝɢɧ ɭɫɬɚɧɨɜɥɟɧɨ, ɱɬɨ ɡɚɦɟɧɚ ɯɪɨɦɨɮɨɪɚ ɢ ɞɚɧɧɚɹ ɬɨɱɟɱɧɚɹ ɦɭɬɚɰɢɹ ɨɤɚɡɵɜɚɸɬ ɚɞɞɢɬɢɜɧɨɟ ɜɥɢɹɧɢɟ ɧɚ ɤɢɧɟɬɢɤɭ ɮɨɬɨɰɢɤɥɚ ȻɊ.

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ȻɅȺȽɈȾȺɊɇɈɋɌɂ ȼɵɪɚɠɚɸ ɢɫɤɪɟɧɧɸɸ ɛɥɚɝɨɞɚɪɧɨɫɬɶ ɦɨɢɦ ɧɚɭɱɧɵɦ ɪɭɤɨɜɨɞɢɬɟɥɹɦ: ɩɪɨɮɟɫɫɨɪɭ, ɞ.ɯ.ɧ. ɏɨɞɨɧɨɜɭ Ⱥ.Ⱥ. ɢ ɫɬɚɪɲɟɦɭ ɧɚɭɱɧɨɦɭ ɫɨɬɪɭɞɧɢɤɭ ɞ.ɛ.ɧ. ɋɤɥɚɞɧɟɜɭ Ⱦ.Ⱥ. (ɎȽɍɉ «Ƚɨɫɇɂɂɝɟɧɟɬɢɤɚ») ɡɚ ɩɨɫɬɨɹɧɧɨɟ ɜɧɢɦɚɧɢɟ ɤ ɪɚɛɨɬɟ ɜ ɩɪɨɰɟɫɫɟ ɟɟ ɜɵɩɨɥɧɟɧɢɹ ɢ ɰɟɧɧɵɟ ɫɨɜɟɬɵ ɩɪɢ ɨɛɫɭɠɞɟɧɢɢ ɪɟɡɭɥɶɬɚɬɨɜ ɢ ɧɚɩɢɫɚɧɢɢ ɞɢɫɫɟɪɬɚɰɢɢ. ə ɢɫɤɪɟɧɧɟ ɩɪɢɡɧɚɬɟɥɶɧɚ ɫɬɚɪɲɟɦɭ ɧɚɭɱɧɨɦɭ ɫɨɬɪɭɞɧɢɤɭ, ɤ.ɛ.ɧ. ɏɢɬɪɢɧɨɣ Ʌ.ȼ. (ɇɂɂ ɮɢɡɢɤɨ-ɯɢɦɢɱɟɫɤɨɣ ɛɢɨɥɨɝɢɢ ɢɦ. Ⱥ.ɇ. Ȼɟɥɨɡɟɪɫɤɨɝɨ ɆȽɍ) ɡɚ ɩɪɚɤɬɢɱɟɫɤɢɟ ɪɟɤɨɦɟɧɞɚɰɢɢ ɩɨ ɩɨɥɭɱɟɧɢɸ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ ɢ ɟɝɨ ɚɧɚɥɨɝɨɜ, ɩɨɦɨɳɶ ɩɪɢ ɢɫɫɥɟɞɨɜɚɧɢɢ ɤɢɧɟɬɢɱɟɫɤɢɯ ɨɫɨɛɟɧɧɨɫɬɟɣ ɮɨɬɨɰɢɤɥɚ ȺȻɊ ɢ ɰɟɧɧɵɟ ɤɨɦɦɟɧɬɚɪɢɢ ɩɪɢ ɨɛɫɭɠɞɟɧɢɢ ɚɜɬɨɪɟɮɟɪɚɬɚ.

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ɋɉɂɋɈɄ ɅɂɌȿɊȺɌɍɊɕ 1. Oesterhelt D., Stoeckenius W. Rhodopsin-like protein from the purple membrane of Halobacterium halobium. // Nature (New Biol.) – 1971. – V. 233, N. 39. – P. 149-152. 2. Hampp N. Bacteriorhodopsin as a photochromic retinal protein for optical memories. // Chem. Rev. – 2000. – V. 100, N 5. – P. 1755-1776. 3. ɋɜɟɬɨɱɭɜɫɬɜɢɬɟɥɶɧɵɟ ɛɢɨɥɨɝɢɱɟɫɤɢɟ ɤɨɦɩɥɟɤɫɵ ɢ ɨɩɬɢɱɟɫɤɚɹ ɪɟɝɢɫɬɪɚɰɢɹ ɢɧɮɨɪɦɚɰɢɢ. // ɋɛɨɪɧɢɤ ɧɚɭɱɧɵɯ ɬɪɭɞɨɜ, Ɋɟɞ. ɂɜɚɧɢɰɤɢɣ Ƚ.Ɋ., ɉɭɳɢɧɨ – 1985. – 209 ɫ. 4. Raymond J.C., Sistrom W.R. The isolation and preliminary characterization of a halophilic photosynthetic bacterium. // Arch. Mikrobiol. – 1967. – V. 59, N. 1. – P. 255-268. 5. Gibbons N.E. Isolation, growth and requirements of halophilic bacteria. // Methods in Microbiology. B. – 1969. – V. 3. – P. 169-183. 6. Kushner D.J. Mass culture of red halophilic bacteria. // Biotechnol. Bioengineering. – 1966. – V. 8, N. 2. – P. 237-245. 7. Kushner D.J. Halophilic bacteria. // Adv. Appl. Microbiol. – 1968. – V. 10. – P.73-99. 8. Kushner D.J. Life in high salt and solute concentrations: halophilic bacteria. // In Microbial life in extreme environments. / Eds. Kushner D.J. London: Academic Press. – 1978. – P. 317-368. 9. Ɉɩɪɟɞɟɥɢɬɟɥɶ ɛɚɤɬɟɪɢɣ Ȼɟɪɞɠɢ. // 9-ɨɟ ɢɡɞɚɧɢɟ, Ɍ. 2. – Ɇ: Ɇɢɪ. – 1997 – ɋ. 754. 10. Margesin R., Schinner F. Potential of halotolerant and halophilic microorganisms for biotechnology. // Extremophiles. – 2001. – V. 5, N. 2. – P. 73-83. 11. Oren A. Bioenergetic aspects of halophilism. // Microbiol. Mol. Biol. Rev. – 1999. – V. 63, N. 2. – P. 334–348. 12. Sauer T., Galinski E.A. Bacterial milking: a novel bioprocess for production of compatible solutes. // Biotechnol. Bioeng. – 1998. – V. 57, N. 3. – P. 306-313. 13. Kulichevskaya I.S., Milekhina E.I., Borzenkov I.A., Zvyagintseva I.S., Belyaev S.S. Oxidation of petroleum hydrocarbons by extremely halophilic archaebacteria. // Microbiology – 1992. – V. 60, N. 3. – P. 596-601. 14. Yakimov M.M., Timmis K.N., Wray V., Fredrickson H.L. Characterization of a new lipopeptide surfactant produced by thermotolerant and halotolerant subsurface Bacillus licheniformis BAS50. // Appl. Environ. Microbiol. – 1995. – V. 61, N. 5. – P. 17061713.

100 15. Banat I.M., Makkar R.S., Cameotra S.S. Potential commercial applications of microbial surfactants. // Appl. Microbiol. Biotechnol. – 2000. – V. 53, N. 5. – P. 495508. 16. Matsunaga T., Sudo H., Takemasa H., Wachi Y. Sulfated extracellular polysaccharide production by the halophilic cyanobacterium Aphanocapsa halophytica immobilized on light-diffusing optical fibers. // Appl. Microbiol. Biotechnol. – 1996. – V. 45, N. 1. – P. 24–27. 17. Choquet CG, Patel GB, Choquet CG, Ekiel I, Sprott GD Formation of stable liposomes from lipid extracts of archaeobacteria. // ɉɚɬɟɧɬ ɋɒȺ US5989587. – ɩɪɢɨɪɢɬɟɬ ɨɬ 23.11.1999. 18. Steinbüchel A., Füchtenbusch B., Gorenflo V., Hein S., Jossek R., Langenbach S., Rehm B.H.A. Biosynthesis of polyesters in bacteria and recombinant organisms. // Polymer. Degrad. Stabil. – 1997. – V. 59, N. 2. – P. 177-182. 19. Rodriguez-Valera F., Lillo J.G. Halobacteria as producers of polyhydroxyalkanoates. // FEMS Microbiol. Rev. – 1992. – V. 103, N. 2. – P. 181-186. 20. Sudge S.S., Bastawde K.B., Gokhale D.B., Kalkote U.R., Ravindranathan T. Production of D-hydantoinase by halophilic Pseudomonas sp. NCIM 5109. // Appl. Microbiol. Biotechnol. – 1998. – V. 49, N. . – P. 594-599. 21. Ryu K., Kim J., Dordick J.S. Catalytic properties and potential of an extracellular protease from an extreme halophile. // Enzyme Microb. Technol. – 1994. – V. 16, N. 4. – P. 266-275. 22. Thongthai C., Suntinanalert P. Halophiles in Thai fish sauce (nam pla). // In: Rodriguez-Valera F (ed) General and applied aspects of halophilic microorganisms. Plenum Press, New York. – 1991. – pp 381–388 23. Röling W.F.M., Prasetyo A.B., Stouthamer A.H., van Verseveld H.W. Physiological aspects of the growth of the lactic acid bacterium Tetragenococcus halophila during Indonesian soy sauce (kecap) production. // J. Appl. Microbiol. – 1999. – V. 86, N. 2. – P. 348-352. 24. Nichols D.S., Nichols P.D., McMeekin T.A. Polyunsatured fatty acids in ɚntarctic bacteria. // Antarct. Sci. – 1993. – V. 5, N. 1. – P. 149-160. 25. Nichols D.S., Russell N.J. Polyunsaturated fatty acids in marine bacteria—a dogma rewritten. // Microbiology (UK). – 1999. – V. 145 (Pt. 4) – P. 767-779. 26. Ⱥɡɢɡɨɜɚ Ɉ. Ƚɚɥɨɮɢɥɶɧɵɟ ɛɚɤɬɟɪɢɢ – ɧɨɜɵɣ ɢɫɬɨɱɧɢɤ ɛɢɨɥɨɝɢɱɟɫɤɢ ɚɤɬɢɜɧɵɯ ɜɟɳɟɫɬɜ. // www.cmjournal.com/rus_page404.htm

101 27. Stoeckenius W. Bacterial rhodopsins: evolution of a mechanistic model for the ion pumps. // Protein Sci. – 1999. – V. 8, N. 2. – P. 447-459. 28. Ventosa A., Oren A. Halobacterium salinarum nom. corrig., a name to replace Halobacterium salinarium (Elazari-Volcani) and to include Halobacterium halobium and Halobacterium cutirubrum. // Int. J. Syst. Microbiol. – 1996. – V. 46. – P. 347. 29. Lanyi J.K. Light energy conversion in Halobacterium halobium. // Microbiol. Rev. – 1978. – V. 42, N. 4. – P. 682-706. 30. Stoeckenius W., Lozier R.H., Bogomolni R.A. Bacteriorhodopsin and purple membrane of halobacteria. // Biochim. Biophys. Acta – 1979. – V. 505, N. 3. – P. 215278. 31. Shahmohammadi H.R., Asgarani E., Terato H., Saito T., Ohyama Y., Gekko K., Yamamoto O., Ide H. Protective roles of bacterioruberin and intracellular KCl in the resistance of Halobacterium salinarium against DNA-damaging agents. // J. Radiat. Res. (Tokyo) – 1998. – V. 39, N. 4. – P. 251-262. 32. Kelly M., Jensen S.L. Bacterial carotenoids. XXVI. C50-carotenoids. 2. Bacterioruberin. // Acta Chem. Scand. – 1967 – V. 21, N. 9. – P. 2578–2580. 33. Kushwaha SC, Kramer JK, Kates M. Isolation and characterization of C50-carotenoid pigments and other polar isoprenoids from Halobacterium cutirubrum. // Biochim. Biophys. Acta – 1975. – V. 398, N. 2. – P. 303-314. 34. Blawrock A.E., Stoeckenius W. Structure of purple membrane. // Nature (New Biol.) – 1971. – V. 233, N. 39. – P. 152-155. 35. Oesterhelt D., Stoeckenius W. Isolation of the cell membrane of Halobacterium halobium and its fractionation into red and purple membrane. // Methods Enzymol. – 1974. – V. 31 (Pt A) – P. 667-678. 36. Oesterhelt D., Brauchle C., Hampp N. Bacteriorhodopsin: a biological material for information processing. // Quart. Rev. Biophys. – 1991. – V. 24, N 4. – P. 425-478. 37. Sumper M., Reitmeier H., Oesterhelt D. Biosynthesis of the purple membrane of halobacteria. // Angew. Chem. Int. Ed. Engl. – 1976. – V.15, N. 5. – P. 187 – 194. 38. Corcelli A., Lattanzio V.M., Mascolo G., Papadia P., Fanizzi F. Lipid-protein stoichiometries in a crystalline biological membrane. NMR quantitative analysis of the lipid extract of the purple membrane. // J. Lipid Res. – 2002. – V. 43, N. 1. – P. 132140. 39. Lukashev E.P., Robertson B. Bacteriorhodopsin retains its light-induced proton pumping function after being heated to 140oC. // Bioelectrochem. Bioenerg. – 1995. – V. 37, N. 2. – P. 157-160.

102 40. Govindjee R., Ohno K., Ebrey T.G. Effect of the removal of the COOH-terminal region of bacteriorhodopsin on its light-induced H+ changes. // Biophys. J. – 1982. – V. 38, N. 1. – P. 85-87. 41. Liao M.J., Khorana H.G. Removal of the carboxyl-terminal peptide does not affect refolding or function of bacteriorhodopsin as a light-dependent proton pump. // J. Biol. Chem. – 1984. – V. 259, N. 7. – P. 4194-4199. 42. Eisenbach M., Caplan S.R. Interaction of purple membrane with solvents. II. Mode of interaction. // Biochim. Biophys. Acta – 1979. – V. 554, N. 2. – P. 281-292. 43. Hiraki K., Hamanaka T., Mitsui T., Kito Y. Structural studies of brown membrane by X-ray diffraction, circular dichroism and absorption spectra. // Photochem. Photobiol. – 1981. – V. 33, N. 4. – P. 429-433. 44. Dencher N.A. The five retinal-protein pigments of halobacteria: bacteriorhodopsin, halorhodopsin, P 565, P 370, and slow-cycling rhodopsin. // Photochem. Photobiol. – 1983. – V. 38, N. 6. – P. 753-767. 45. Oren A. Bioenergetic aspects of halophilism. // Microbiol. Molecular Biol. Rev. – 1999. – V. 63, N. 2. – P. 334-348. 46. ɋɤɭɥɚɱɟɜ ȼ.ɉ. Ȼɢɨɷɧɟɪɝɟɬɢɤɚ. Ɇɟɦɛɪɚɧɧɵɟ ɩɪɟɨɛɪɚɡɨɜɚɬɟɥɢ ɷɧɟɪɝɢɢ. // Ɇ.: ȼɵɫɲɚɹ ɲɤɨɥɚ. – 1989. – ɋ. 102-117. 47. Birge R.R. Nature of the primary photochemical events in rhodopsin and bacteriorhodopsin. // Biochim. Biophys. Acta – 1990. – V. 1016, N. 3. – P. 293 – 327. 48. Hampp N., Popp A., Braeuchle C. Diffraction efficiency of bacteriorhodopsin films for holography containing bacteriorhodopsin wild type BRWT and its BRD85E and BRD96N. // J. Phys. Chem. – 1992. – V. 96, N. 11. – P. 4679-4685. 49. Okada-Shudo Y., Yamaguchi I., Tomioka H., Sasabe H. Real-time image processing using polarization discrimination of bacteriorhodopsin. // Synthetic Metals. – 1996. – V. 81, N. 2-3. – P. 147-149. 50. Bousche O., Sonar S., Krebs M.P., Khorana H.G., Rothschild K.J. Time-resolved Fourier transform infrared spectroscopy of the bacteriorhodopsin mutant Tyr-185 o Phe: Asp-96 reprotonates during O formation; Asp-85 and Asp-212 deprotonate during O decay. // Photochem. Photobiol. – 1992. – V. 56, N. 6. – P. 1085-1095. 51. Braitman M.S., Mogi T., Marti T., Stern L.J., Khorana H.G., Rothschild K.J. Vibrational spectroscopy of bacteriorhodopsin mutants: light-driven proton transport involves protonation changes of aspartic acid residues 85, 96, and 212. // Biochemistry. – 1988. – V. 27, N.23. – P. 8516 – 8520.

103 52. Logunov I., Schulten K. Quantum chemistry: molecular dynamics study of the darkadaptation process in bacteriorhodopsin. // J. Am. Chem. Soc. – 1996. – V. 118, N. 40. – P. 9727-9735 53. Scherrer P., Mathew M.K., Sperling W., Stoeckenius W. Retinal isomer ratio in darkadapted purple membrane and bacteriorhodopsin monomers. // Biochemistry. – 1989. – V. 28, N. 2. – P. 829-835. 54. Hatcher M.E., Hu J.G., Belenky M., Verdegem P., Lugtenburg J., Griffin R.G., Herzfeld J. Control of the pump cycle in Bacteriorhodopsin: mechanisms elucidated by solid-state NMR of the D85N mutant. // Biophys. J. – 2002. – V. 82, N. 2. – P. 1017–1029 55. Hage W., Kim M., Frei H., Mathies R.A. Protein dynamics in the bacteriorhodopsin photocycle: a nanosecond step-scan FTIR investigation of the KL to L transition. // J. Phys. Chem. – 1996. – V. 100, N. 39. – P. 16026-16033. 56. Haupts U., Tittor J., Oesterhelt D. Closing on bacteriorhodopsin: progress in understanding the molecule. // Annu. Rev. Biophys. Biomol. Struct – 1999. – V. 28. – P. 367–399. 57. Subramaniam S., Greenhalgh D.A., Khorana H.G. Aspartic acid 85 in bacteriorhodopsin functions both as proton acceptor and negative counterion to the Schiff base. // J. Biol. Chem. – 1992. – V. 267, N. 36. – P. 25730-25733. 58. Brown L.S., Dioumaev A.K., Needleman R., Lanyi J.K. Local-access model for proton transfer in bacteriorhodopsin. // Biochemistry. – 1998. – V. 37, N 7. – P. 3982– 3993. 59. Luecke H., Richter H.T., Lanyi J.K. Proton transfer pathways in bacteriorhodopsin at 2.3 angstrom resolution. // Science. – 1998. – V. 280, N. 5371. – P. 1934-1937. 60. Ovchinnikov Yu.A. Bioorganic chemistry of rhodopsins. // Pure Appl. Chem. – 1986. – V. 58, N. 5. – P. 725-736. 61. Ɉɜɱɢɧɧɢɤɨɜ ɘ.Ⱥ., Ⱥɛɞɭɥɚɟɜ ɇ.Ƚ., Ɏɟɣɝɢɧɚ Ɇ.ɘ., Ʉɢɫɟɥɟɜ Ⱥ.ȼ., Ʌɨɛɚɧɨɜ ɇ.Ⱥ., ɇɚɡɢɦɨɜ ɂ.ȼ. ɉɟɪɜɢɱɧɚɹ ɫɬɪɭɤɬɭɪɚ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1978. – Ɍ. 4, N. 11. – ɋ. 1573-1574. 62. Hacket N.R., Stern L.J., Chao B.H., Kronis K.A., Khorana H.G. Structure-function studies on bacteriorhodopsin. V. Effects of amino acid substitutions in the putative helix F. // J. Biol. Chem. – 1987. – V. 262, N. 19. – P. 9277-9284.

104 63. Jang D., El-Sayed M.A., Stern L.J., Mogi T., Khorana H.G. Effect of genetic modification of tyrosine-185 on the proton pump and the blue-to-purple transition in bacteriorhodopsin. // Proc. Natl. Acad. Sci. USA – 1990. – V. 87, N 11. – P. 41034107. 64. Dunach M., Marti T., Khorana H.G., Rothschild K.J. UV-visible spectroscopy of bacteriorhodopsin mutants: substitution of Arg-82, Asp-85, Tyr-185, and Asp-212 results in abnormal light-dark adaptation. // Proc. Natl. Acad. Sci. USA – 1990. – V. 87, N. 24. – P. 9873-9877. 65. Lin S.W., Fodor S.P.R., Miercke L.J.W., Schand R.F., Beatlach M.C., Stroud R.M., Mathies R.A. Resonance Raman spectra of bacteriorhodopsin mutants with substitutions at Asp-85, Asp-96 and Arg-82. // Photochem. Photobiol. – 1991. – V. 53, N. 3. – P. 341-346. 66. Miercke L.J.W., Betlach M.C., Mitra A.K., Shand R.F., Fong S.K., Stroud R.M. Wildtype and mutant bacteriorhodopsins D85N, D96N, and R82Q: purification to homogeneity, pH dependence of pumping, and electron diffraction. // Biochemistry. – 1991. – V. 30, N. 12. – P. 3088 – 3098. 67. Krebs M.P., Mollaaghababa, Khorana H.G. Gene replacement in Halobacterium halobium and expression of bacteriorhodopsin mutants. // Proc. Natl. Acad. Sci. USA – 1993. – V. 90, N 5. – P. 1987-1991. 68. Mogi T., Stern L.J., Chao B.H., Khorana H.G. Structure-function studies on bacteriorhodopsin. VIII. Substitutions of the membrane-embedded prolines 50, 91 and 186: the effects are determined by the substituting amino acids. // J. Biol. Chem. – 1989. – V.264, N. 24. – P. 14192-14196. 69. Zhang Y.N., El-sayed M.A., Marti T., Stern L.J., Mogi T., Khorana H.G. Effect of mutagenic substitution of prolines on the rate of deprotonation and deprotonation of Schiff base during the photocycle of bacteriorhodopsin. // Photochem. Photobiol. – 1993. – V. 57, N. 6. – P. 1027-1031. 70. Feng Y., Menick D.R., Katz B.M., Beischel C.J., Hazard E.S., Misra S., Ebrey T.G., Crouch R.K. Probing of the retinal binding site of bacteriorhodopsin by affinity labeling. // Biochemistry. – 1994. – V. 33, N. 38. – P. 11624 – 11630. 71. Balashov S.P., Govindjee R., Kono M., Imasheva E., Lukashev E., Ebrey T.G., Crouch R.K., Menick D.R., Feng Y. Effect of the arginine-82 to alanine mutation in bacteriorhodopsin on dark adaptation, proton release, and the photochemical cycle. // Biochemistry. – 1993. – V. 32, N. 39. – P. 10331 – 10343.

105 72. Stern L.J., Khorana H.G. Structure-function studies on bacteriorhodopsin. X. Individual substitutions of arginine residues by glutamine affect chromophore formation, photocycle and proton translocation. // J. Biol. Chem. – 1989. – V. 264, N. 24. – P. 14202-14208. 73. Flitsch S.L., Khorana H.G. Structural studies on transmembrane proteins. I. Model study using bacteriorhodopsin mutants containing single cysteine residues. // Biochemistry. – 1989. – V. 28, N. 19. – P. 7800 – 7805. 74. Mogi T., Marti T., Khorana H.G. Structure-function studies on bacteriorhodopsin. IX. Substitutions of tryptophan residues affect protein-retinal interaction in bacteriorhodopsin. // J. Biol. Chem. – 1989. – V. 264, N. 24. – P. 14197-14201. 75. Brown L.S. Reconciling crystallography and mutagenesis: a synthetic approach to the creation of a comprehensive model for proton pumping by bacteriorhodopsin. // Biochim. Biophys. Acta – 2000. – V. 1460, N. 1. – P. 49–59. 76. Imasheva E.S., Balashov S.P., Ebrey T.G., Chen N., Crouch R.K., Menick D.R. Two groups control light-induced Schiff base deprotonation and reprotonation affinity of Asp85 in the Arg82His mutant of bacteriorhodopsin. // Biophys. J. – 1999. – V. 77, N. 5. – P. 2750-2763 77. Tittor J., Soell C., Oesterhelt D., Butt H.J., Bamberg E. A defective proton pump, point-mutated bacteriorhodopsin Aspo96Asn is fully reactivated by azide. // EMBO J. – 1989. – V. 8, N 11. – P. 3477-3482. 78. Ɇɢɰɧɟɪ Ȼ.ɂ., ɏɨɞɨɧɨɜ Ⱥ.Ⱥ., Ɂɜɨɧɤɨɜɚ ȿ.ɇ., ȿɜɫɬɢɝɧɟɟɜɚ Ɋ.ɉ. Ⱥɧɚɥɨɝɢ ɪɟɬɢɧɚɥɹ: ɫɢɧɬɟɡ ɢ ɜɡɚɢɦɨɞɟɣɫɬɜɢɟ ɫ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɨɦ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1986. – Ɍ. 12, N. 1. – ɋ. 5-53. 79. The Retinoids. V. 1, 2. // Eds. Sporn M.B., Roberts A.B., Goodman D.S. Orlando: Acad. Press – 1984. – 424 p., 446 p. 80. Chemistry and Biology of Synthetic Retinoids. // Eds. Dawson M.I., Okamura W.H. Boca Raton: CRC Press – 1990. – 615 p. 81. Khodonov A.A., Kirillova Yu.G., Eremin S.V., Khitrina L.V. Synthesis and investigation of retinal and bacteriorhodopsin analogs with variable length or alternative bond position of chromophore polyene chain. // Abstracts of Vl-th International Conference on Retinal Proteins. Leiden, The Netherlands. – 1994. – P. 119. 82. Tokunaga F., Govinjee R., Ebrey T.G., Crouch R. Synthetic pigment analogues of the purple membrane protein. // Biophys. J. – 1977. – V. 19, N. 2. – P. 191-198.

106 83. Crouch R.K., Scott R., Ghent S. Govindjee R., Chang Ch-H., Ebrey T. Properties of synthetic bacteriorhodopsin pigments. Further probes of the chromophore binding site. // Photochem. Photobiol. – 1986. – V. 43, N. 3. – P. 297-303. 84. Oesterhelt D., Christoffel V. Reconstitution of a proton pump. // Biochem. Soc. Transact. – 1976. – V. 4, N. 4. – P. 556-559. 85. Balog-Nair V., Carriker J.D., Honig B., Kamat N., Motto M.G., Nakanishi K., Sen R., Sheves M., Tanis M.A., Tsujimoto K. The opsin shift in bacteriorhodopsin studies with artificial bacteriorhodopsins. // Photochem. Photobiol. – 1981. – V. 33, N. 4. – P. 483-488. 86. Nakanishi Ʉ., Balogh-Nair V., Arnaboldi N., Tsujimoto K., Honig ȼ. An external point-charge model for bacteriorhodopsin to account for its purple color // J. Am. Chem. Soc. – 1980. – V. 102, N. 27. – P. 7945-7947. 87. Ɇɚɨ ȼ., Govindjee R., Ebrey T.G., Arnaboldi M., Balogh-Nair V., Nakanishi K., Crouch R. Photochemical and functional properties of bacteriorhodopsins formed from 5,6-dihydro-and 5,6-dehydrodesmethylretinals. // Biochemistry. – 1981. – V. 20, N. 2. – P. 428-435. 88. Gartner W., Oesterhelt D., Seifer-Schiller E., Towner P., Hopf H., Bohm I. Acetylenic retinals form functional bacteriorhodopsins but not bovine rhodopsins. // J. Am. Chem. Soc. – 1983. – V. 106, N. 19. – P. 5654-5659. 89. Ⱦɚɧɲɢɧɚ ɋ.ȼ., Ⱦɪɚɱɟɜ Ⱥ.Ʌ., Ⱦɪɚɱɟɜ Ʌ.Ⱥ., Ʉɚɭɥɟɧ Ⱥ.Ⱦ., Ɇɢɰɧɟɪ Ȼ.ɂ., ɏɢɬɪɢɧɚ Ʌ.ȼ. ɏɨɞɨɧɨɜ Ⱥ.Ⱥ. ɐɢɫ- ɢ ɬɪɚɧɫ-ɢɡɨɦɟɪɵ 11,12-ɞɢɞɟɝɢɞɪɨɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1989. – Ɍ. 15, N. 3. – ɋ. 307-312. 90. ɏɢɬɪɢɧɚ Ʌ.ȼ., ȿɪɟɦɢɧ ɋ.ȼ., ɏɨɞɨɧɨɜ Ⱥ.Ⱥ., Ʉɚɭɥɟɧ Ⱥ.Ⱦ. ɏɪɨɦɨɮɨɪ-ɨɛɪɚɡɭɸɳɢɟ ɢɡɨɦɟɪɵ 11,12-ɞɢɞɟɝɢɞɪɨɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. // Ȼɢɨɥ. ɦɟɦɛɪɚɧɵ. – 1994. – Ɍ. 11, N. 5. – ɋ. 575-576. 91. ɏɢɬɪɢɧɚ Ʌ.ȼ., Ʉɚɭɥɟɧ Ⱥ.Ⱦ., ȿɪɟɦɢɧ ɋ.ȼ., ɏɨɞɨɧɨɜ Ⱥ.Ⱥ. Ȼɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧ. Ȼɟɥɤɨɜɵɟ ɨɝɪɚɧɢɱɟɧɢɹ ɢɡɨɦɟɪɢɡɚɰɢɢ ɯɪɨɦɨɮɨɪɚ. // Ɇɨɥɟɤ. ɛɢɨɥɨɝɢɹ. – 1995. – Ɍ. 29, N. 6. – ɋ. 1398-1407. 92. Ⱦɚɧɲɢɧɚ ɋ.ȼ., Ⱦɪɚɱɟɜ Ⱥ.Ʌ., Ⱦɪɚɱɟɜ Ʌ.Ⱥ., ȿɪɟɦɢɧ ɋ.ȼ., Ʉɚɭɥɟɧ Ⱥ.Ⱦ., Ɇɢɰɧɟɪ Ȼ.ɂ., ɏɢɬɪɢɧɚ Ʌ.ȼ., ɑɟɤɭɥɚɟɜɚ Ʌ.ɇ. Ɏɨɬɨɰɢɤɥ ɢ ɷɥɟɤɬɪɨɝɟɧɟɡ 13ɞɟɡɦɟɬɢɥɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. // Ȼɢɨɮɢɡɢɤɚ. – 1989. – Ɍ. 34, N. 4. – ɋ. 623-626.

107 93. Broek A.D., Muradin-Szweykowska M., Lugtenburg J., van der Bend R.L., van Dijke P.W.M. Bacteriorhodopsins with a chemically modified chromophore. The lightdriven proton pump action of [13-demethyl-ll,14-epoxy]-, [9-demethyl]-, [13demethyl]- and [9,13-bissdemethyl]-bacteriorhodopsin. // Rec. Trav. Chim. Pays-Bas. – 1983. – V. 102, N. 1. – P. 42-46. 94. Broek A.D., Muradin-Szweykowska M., Courtin J.M.L., Lugtenburg J. Preparation of 11,14-epoxy-bridged and isomeric chain-demethylated retinals. 13-Demethyl-ll,14epoxy-, 9-demethyl-, 13-demethyl- and 9,13-bisdemethyl-retinals. // Rec. Trav. Chim. Pays-Bas. – 1983. – V. 102, N.1. P. 46-51. 95. ɋɟɪɟɛɪɹɧɵɣ ȼ.Ⱥ., Ɇɢɰɧɟɪ Ȼ.ɂ., Ɂɚɤɢɫ ȼ.ɂ., ɐɟɬɥɢɧ ȼ.ɂ. Ⱥɧɚɥɨɝɢ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ ɧɚ ɨɫɧɨɜɟ 4-ɡɚɦɟɳɟɧɧɵɯ ɪɟɬɢɧɚɥɟɣ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1981. – Ɍ. 7, N 11. – ɋ. 1731-1733. 96. Shealy Y.F., Hosmer C.A., Riordan J.M. Modification of the intact retinoid structure in the cyclohexenyl region: alkylation of methyl 4-oxoretinoate. // Synthesis. – 1993. – N. 11. – P. 1095-1098. 97. Rao V.J., Derguini F., Nakanishi K., Tagushi Ɍ., Hosoda A., Hanzava Y., Kobayashi Y., Pande S.M., Callender R.H. 5-(Trifluoromethyl)bacteriorhodopsin does not translocate protons. // J. Am. Chem. Soc. – 1986. – V. 108, N 19. – P. 6077-6078. 98. Fang J.M., Carriker J.D., Balog-Nair V., Nakanishi K. Evidence for the necessity of double bond (13-ene)-isomerization in the proton pumping of bacteriorhodopsin. // J. Am. Chem. Soc. – 1983. – V. 105, N. 15. – P. 5162-5164. 99. Ʉɢɪɢɥɥɨɜɚ ɘ.Ƚ., ȿɪɟɦɢɧ ɋ.ȼ., ɏɢɬɪɢɧɚ Ʌ.ȼ., Ɇɢɰɧɟɪ Ȼ.ɂ. ɐɢɤɥɨɩɟɧɬɟɧɨɜɵɟ ɢ ɰɢɤɥɨɝɟɤɫɟɧɨɜɵɟ ɚɧɚɥɨɝɢ ɪɟɬɢɧɚɥɹ ɩɨ-ɪɚɡɧɨɦɭ ɜɡɚɢɦɨɞɟɣɫɬɜɭɸɬ ɫ ɛɚɤɬɟɪɢɨɨɩɫɢɧɨɦ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1993. – Ɍ. 19, N. 8. – ɋ. 825-835. 100. Ʉɢɪɢɥɥɨɜɚ ɘ.Ƚ., ɏɢɬɪɢɧɚ Ʌ.ȼ., ɏɨɞɨɧɨɜ Ⱥ.Ⱥ. ȼɡɚɢɦɨɞɟɣɫɬɜɢɟ ɦɟɬɢɥɰɢɤɥɨɩɟɧɬɟɧɨɜɵɯ ɚɧɚɥɨɝɨɜ 13-ɰɢɫ ɪɟɬɢɧɚɥɹ ɫ ɛɚɤɬɟɪɢɨɨɩɫɢɧɨɦ. // Ȼɢɨɥ. ɦɟɦɛɪɚɧɵ. – 1993. – Ɍ. 10, N. 4. – ɋ. 447-448. 101. Kirillova Yu.G., Eremin S.V., Khitrina L.V., Khodonov A.A. 13-Cis-retinal locked analogues interact with bacterioopsin in different manner. // Abstracts of Vlth International Conference on Retinal Proteins. Leiden, The Netherlands. – 1994. – P. 120. 102. Groesbeek M., Kirillova Yu.G., Boeff R., Lugtenburg J. Synthesis of six novel retinals and their interaction with bacterioopsin. // Rec. Trav. Chim. Pays-Bas. – 1994. – V. 113. – P. 45-52.

108 103. Khodonov A.A., Khitrina L.V. Synthesis of heavy atom labeled retinal and investigation of its interaction with bacterioopsin. // Abstracts of Vl-th International Conference on Retinal Proteins. Leiden, The Netherlands. – 1994. – P. 118. 104. Danshina S.V., Drachev A.L., Drachev L.A., Eremin S.V., Kaulen A.D., Khitrina L.V., Mitsner B.I. ɋ13-substituted bacteriorhodopsin analogs. // Archiv. Biochem. Biophys. – 1990. – V.279, N2. – P. 225-231. 105. ɏɨɞɨɧɨɜ A.A., ȿɪɟɦɢɧ ɋ.ȼ., Ʌɨɤɲɢɧ Ⱦɠ. Ʌ., ɒɜɟɰ ȼ.ɂ., Ⱦɟɦɢɧɚ Ɉ.ȼ., ɏɢɬɪɢɧɚ Ʌ.ȼ., Ʉɚɭɥɟɧ Ⱥ.Ⱦ. Ⱥɧɚɥɨɝɢ ɪɟɬɢɧɚɥɹ ɢ ɢɯ ɪɨɥɶ ɜ ɢɫɫɥɟɞɨɜɚɧɢɢ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1996. – Ɍ. 22, N. 10-11. – P. 745-776. 106. Ȼɪɨɭɧ Ʌ.ɋ. ɂɫɫɥɟɞɨɜɚɧɢɟ ɦɟɯɚɧɢɡɦɨɜ ɩɟɪɟɧɨɫɚ ɩɪɨɬɨɧɨɜ ɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɟ ɫ ɩɨɦɨɳɶɸ ɬɨɱɟɱɧɨɝɨ ɦɭɬɚɝɟɧɟɡɚ ɢ ɢɡɦɟɧɱɢɜɨɫɬɢ ɩɟɪɜɢɱɧɨɣ ɫɬɪɭɤɬɭɪɵ ɛɟɥɤɚ. // Ȼɢɨɯɢɦɢɹ. – 2001. – Ɍ. 66, N. 11. – ɋ. 1546-1554. 107. Soppa J., Oesterhelt D. Bacteriorhodopsin mutants of Halobacterium sp. GRB. // J. Biol. Chem. – 1989. – V. 264, N. 22. – P. 13043-13048. 108. Dunn R.J., Hacket N.R., McCoy J.M., Chao B.H., Kimura K., Khorana H.G. Structure-function studies on bacteriorhodopsin. I. Expression of the bacterio-opsin gene in Escherichia coli. // J. Biol. Chem. – 1987. – V. 262, N. 19. – P. 9246-9254. 109. Karnik S.S., Nassal M., Doi T., Jay E., Sgaramella V., Khorana H.G. Structurefunction studies on bacteriorhodopsin. II. Improved expression of the bacterio-opsin gene in Escherichia coli. // J. Biol. Chem. – 1987. – V. 262, N. 19. – P. 9255-9263. 110. Nassal M., Mogi T., Karnik S.S., Khorana H.G. Structure-function studies on bacteriorhodopsin. III. Total synthesis of a gene for bacterio-opsin and its expression in Escherichia coli. // J. Biol. Chem. – 1987. – V. 262, N. 19. – P. 9264-9270. 111. Ni B., Chang M., Duschl A., Lani J., Needleman R. An efficient system for the synthesis of bacteriorhodopsin in Halobacterium halobium. // Gene. – 1990. – V. 90, N. 1. – P. 169 – 172. 112. Braiman M.S., Stern L.J., Chao B.H., Khorana H.G. Structure-function studies on bacteriorhodopsin. IV. Purification and renaturation of bacterio-opsin polypeptide expressed in Escherichia coli. // J. Biol. Chem. – 1987. – V. 262, N. 19. – P. 92719276. 113. London E., Khorana H.G. Denaturation and renaturation of bacteriorhodopsin in detergents and lipid-detergent mixtures. // J. Biol. Chem. – 1982. – V. 257, N. 12. – P. 7003-7011.

109 114. ɉɨɥɨɜɧɢɤɨɜɚ Ɉ.ɘ., ɋɢɦɨɧɨɜɚ Ɍ.ɇ., Ɍɚɪɚɯɨɜɫɤɢɣ ɘ.ɋ., ɇɢɤɨɧɨɜɚ Ɍ.ɇ., ɋɭɯɚɧɨɜ ɋ.ȼ., Ⱦɭɛɚɱɟɜ Ƚ.ɗ., Ȼɨɪɨɜɹɝɢɧ ȼ.Ʌ., ȼɚɫɢɥɟɧɤɨ ɂ.Ⱥ., Ȼɚɪɫɭɤɨɜ Ʌ.ɂ. Ɋɟɤɨɧɫɬɪɭɤɰɢɹ ɩɭɪɩɭɪɧɵɯ ɦɟɦɛɪɚɧ ɢɡ ɞɟɥɢɩɢɞɢɪɨɜɚɧɧɨɝɨ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɷɧɞɨɝɟɧɧɵɯ ɢ ɷɤɡɨɝɟɧɧɵɯ ɥɢɩɢɞɨɜ. ȼɥɢɹɧɢɟ ɫɬɪɭɤɬɭɪɧɵɯ ɢ ɜɧɟɲɧɢɯ ɮɚɤɬɨɪɨɜ. // Ȼɢɨɥ. ɦɟɦɛɪɚɧɵ. – 1995. – Ɍ. 12, N. 3. – ɋ. 260-277. 115. Subramaniam S., Marti T., Roesselet S.J., Rothschild K.J., Khorana H.G. The reaction of hydroxylamine with bacteriorhodopsin studied with mutants that have altered photocycles: selective reactivity of different photointermediates. // Proc. Natl. Acad. Sci. USA – 1991. – V. 88, N. 6. – P. 2583-2587. 116. Schweiger U., Tittor J., Oesterhelt D. Bacteriorhodopsin can function without a covalent linkage between retinal and protein. // Biochemistry. – 1994 – V. 33, N. 2. – P. 535 – 541 117. Soppa J., Otomo J., Staub J., Tittor J., Meeben S., Oesterhelt D. Bacteriorhodopsin mutants of Halobacterium sp. GRB. II. Characterization of mutant. // J. Biol. Chem. – 1989. – V. 264, N. 22. – P. 13049–13056. 118. Imasheva E.S., Lu M., Balashov S.P., Ebrey T.G., Chen Y., Ablonczy Z., Menick D.R., Crouch R.K. Exploring the function of Tyr83 in bacteriorhodopsin: features of the Y83F and Y83N mutants. // Biochemistry. – 2001. – V. 40, N. 44. – P. 13320– 13330 119. Russell T.S., Coleman M., Rath P., Nilsson A., Rothschild K.J. Threonine-89 participates in the active site of bacteriorhodopsin: evidence for a role in color regulation and Schiff base proton transfer. // Biochemistry. – 1997. – V. 36, N. 24. – P. 7490–7497. 120. Isenbarger T.A., Krebs M.P. Role of helix-helix interactions in assembly of bacteriorhodopsin lattice. // Biochemistry. – 1999. – V. 38, N. 28. – P. 9023–9030. 121. Otto H., Marti T., Holz M., Mogi T., Stern L.J., Engel F., Khorana H.G., Heyn M.P. Substitution of amino acids Asp-85, Asp-212, and Arg-82 in bacteriorhodopsin affects the proton release phase of the pump and pK of Schiff base. // Proc. Natl. Acad. Sci. USA – 1990. – V. 87, N. 3. – P. 1018-1022. 122. Subramaniam S., Greenhalgh D.A., Rath P., Rothschild K.J., Khorana H.G. Replacement of leucine-93 by alanine or threonine slows down the decay of the N and O intermediates in the photocycle of bacteriorhodopsin: implications for proton uptake and 13-cis-retinal o all-trans-retinal reisomerisation. // Proc. Natl. Acad. Sci. USA – 1991. – V. 88, N. 15. – P. 6873-6877.

110 123. Subramaniam S., Marti T., Khorana H.G. Protonation state of Asp (Glu)-85 regulates the purple-to-blue transition in bacteriorhodopsin mutants Arg-82 o Ala and Asp-85 o Glu: the blue form is inactive in proton translocation. // Proc. Natl. Acad. Sci. USA – 1990. – V. 87, N. 3. – P. 1013-1017. 124. Marti T., Otto H., Roesselet S.J., Heyn M.P., Khorana G.H. Consequences of amino acid insertions and/or deletions in transmembrane helix C of bacteriorhodopsin. // Proc. Natl. Acad. Sci. USA – 1992. – V. 89, N. 4. – P. 1219-1223. 125. Greenhalgh D.A., Farrensl D.L., Subramaniamg S., Khorana H.G. Hydrophobic amino acids in the retinal-binding pocket of bacteriorhodopsin. // J. Biol. Chem. – 1993. – V. 268, N. 27. – P. 20305-20311. 126. Greenhalgh D.A., Subramaniam S., Alexiev U., Otto H., Heyng M.P., Khorana H.G. Effect of introducing different carboxylate-containing side chains at position 85 on chromophore formation and proton transport in bacteriorhodopsin. // J. Biol. Chem. – 1992 – V. 267, N. 36. – P. 25734-25738. 127. Greenhalgh D.A., Altenbach C., Hubbel W.L., Khorana H.G. Locations of Arg-82, Asp-85, and Asp-96 in helix C of bacteriorhodopsin relative to the aqueous boundaries. // Proc. Natl. Acad. Sci. USA – 1991. – V. 88, N. 19. – P. 8626-8630. 128. Marti T., Rosselet S.J., Otto H., Hey M.P., Khoranal H.G. The retinylidene Schiff base counterion in bacteriorhodopsin. // J. Biol. Chem. – 1991. – V. 266, N. 28. – P. 1867418683. 129. Altenbach C., Grenhalgh D.A., Khorana H.G., Hubbel W.L. A collision gradient method to determine the immersion depth of nitroxides in lipid bilayers: application to spin-labeled mutants of bacteriorhodopsin. // Proc. Natl. Acad. Sci. USA – 1994. – V. 91, N. 5. – P. 1667-1671. 130. Marti T., Otto H., Mogi T., Roeselet S.J., Heyn M.P., Khorana H.G. Bacteriorhodopsin mutants containing single substitutions of serine or threonine residues are all active in proton translocation. // J. Biol. Chem. – 1991. – V. 266, N. 11. – P. 6919-6927. 131. Mogi T., Stern L.J., Marti T., Chao B.H., Khorana H.G. Aspartic acid substitutions affect proton translocation by bacteriorhodopsin. // Proc. Natl. Acad. Sci. USA – 1988. – V. 85, N. 12. – P. 4148-4152. 132. Drachev L.A., Kaulen A.D., Khorana H.G., Mogi T., Postanogova N.V., Skulachev V.P., Stern L.J. The role of arginine 82 and 227 in the bacteriorhodopsin proton pump. // Photochem. Photobiol. – 1992. – V. 55, N. 5. – P. 741-744.

111 133. Sonar S., Krebs M.P., Khorana H.G., Rothschild K.J. Static and time-resolved absorbtion spectroscopy of the bacteriorhodopsin mutant Tyr-185oPhe: evidence for an equilibrium between bR570 and O-like species. // Biochemistry. – 1993. – V. 32, N. 9. – P. 2263 – 2271. 134. Friedman N., Druckmann S., Lanyi J., Needleman R., Lewis A., Ottolenghi M., Sheves M. A covalent link between the chromophore and the protein backbone of bacteriorhodopsin is not required for forming a photochemically active pigment analogous to the wild type. // Biochemistry. – 1994 – V. 33, N. 8. – P. 1971 – 1976 135. Opportunities in biotechnology for future army applications. // National Academy Press, Washington, D.C. – 2000. – 118 pp. 136. Bouas-Laurent H., Durr H. Organic photochromism (IUPAC technical report). // Pure Appl. Chem. – 2001. – V. 73, N. 4. – P. 639-665. 137. Brauchle C., Hampp N., Oesterhelt D. Optical applications of bacteriorhodopsin and its mutated variants. // Adv. Mater. – 1991. – V. 3, N 9. – P. 420-428. 138. Birge R.R., Gillespie N.B., Izaguierre E.W., Kusnetzow A., Lawrence A.F., Sinch D., Song Q.W., Schmidt E., Stuart A., Seetharaman S., Wise K.J. Biomolecular electronics: protein-based associative processors and volumetric memory. // J. Phys. Chem. – 1999. – V. 103 B, N.49. – P. 10746-10766. 139. Popp A., Wolperdinger M., Hampp N., Bruchle C., Oesterhelt D. Photochemical conversion of the O-intermediate to 9-cis-retinal- containing products in bacteriorhodopsin films // Biophys. J. – 1993. – V. 65, N. 4. – P. 1449-1459. 140. ȼɫɟɜɨɥɨɞɨɜ ɇ.ɇ., ɉɨɥɬɨɪɚɧɫɤɢɣ ȼ.Ⱥ. Ƚɨɥɨɝɪɚɦɦɵ ɧɚ «Ȼɢɨɯɪɨɦɟ», ɛɢɨɥɨɝɢɱɟɫɤɨɦ ɮɨɬɨɯɪɨɦɧɨɦ ɦɚɬɟɪɢɚɥɟ. // ɋɨɜ. ɮɢɡ. ɬɟɯ. ɮɢɡ. – 1985. – Ɍ. 30. – ɋ. 1235. 141. Hampp N., Brauchle C., Oesterhelt D. Bacteriorhodopsin wild type and variant aspartate-96oasparagine as reversible holographic media. // Biophys. J. – 1990. – V. 56, N. 6. – P. 83-93. 142. Zaitzev S.Y., Kozhevnikov N.M., Bapmenkov Y.O., Lipovskaya M.Y. Kinetics of dynamic hologram recording in polymer films with immobilized bacteriorhodopsin. // Photochem. Photobiol. – 1992. – V. 55, N. 6. – P. 851-856. 143. Seitz A., Hampp N. Kinetic optimization of bacteriorhodopsin films for holographic interferometry. // J. Phys. Chem. B – 2000. – V. 104, N. 30. – P. 7183-7192.

112 144. Gu L.-Q., Zhang C.-P., Niu A.-F., Li J., Zhang G.-Y., Wang Y.-M., Tong M.-R., Pan J.-L., Song Q.W., Parsons B., Birge R.R. Bacteriorhodopsin based photonic logic gate and its applications to gray level image subtraction. // Optics Commun. – 1996. – V. 131, N. 1. – P. 25-30. 145. Birge R.R. Protein based computers. // Scientific American. – 1995. – N 3. – P. 90-95. 146. Min J., Choi H.G., Oh B.-K., Lee W.H., Paek S.-H., Choi J.-W. Visual information processing using bacteriorhodopsin-based complex LB films. // Biosens. Bioelectron. – 2001. – V. 16, N 9-12. – P. 917-923. 147. Min J., Choi J.-W., Lee W.H., Kim U.R. Photoreceptor consisting of spiropyranbacteriorhodopsin films for photosignal enhancement. // Biosens. Bioelectron. – 1998. – V. 13, N. 11. – P. 1151-1155. 148. Choi H.-G., Min J., Choi J.-W. , Lee W. H. Molecular photoreceptor consisting of bacteriorhodopsin/flavin complex Langmuir–Blodgett films. // Biosens. Bioelectron. – 1998. – V. 13, N. 10. – P. 1069-1075. 149. Choi H.-G., Jung W.-C., Min J., Lee W.H., Choi J.-W. Color image detection by biomolecular photoreceptor using bacteriorhodopsin-based complex LB films. // Biosens. Bioelectron. – 2001. – V. 16, N. 9-12. – P. 925-935. 150. Kolodner P., Lukashev E.P., Ching Y.-C., Druzhko A.B. Electric-field and photochemical effects in D85N mutant bacteriorhodopsin substituted with 4-ketoretinal. // Thin Solid Films. – 1997. – V. 302, N. 2. – P. 231-234. 151. Druzhko A.B., Weetal H.H. Photoinduced transformation of wild-type and D96Nmutant 4-keto-bacteriorhodopsin gelatin films. // Thin Solid Films. – 1997. – V. 293, N. 3. – P. 281-284. 152. ɏɨɞɨɧɨɜ Ⱥ.Ⱥ., Ɇɢɰɧɟɪ Ȼ.ɂ., Ɂɜɨɧɤɨɜɚ ȿ.ɇ., ȿɜɫɬɢɝɧɟɟɜɚ Ɋ.ɉ. Ⱥɪɨɦɚɬɢɱɟɫɤɢɟ ɚɧɚɥɨɝɢ 13-ɰɢɫ ɢ ɩɨɥɧɨɫɬɶɸ-ɬɪɚɧɫ-ɪɟɬɢɧɚɥɟɣ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1987. – Ɍ. 13, N. 2. – ɋ. 238-251. 153. Khodonov A.A., Demina O.V., Khitrina L.V., Kaulen A.D., Silfsen P., Parkkinen S., Parkkinen J., Jaaskelainen T. Modified bacteriorhodopsins as a basis for new optical devices. // Sensors Actuators. B. – 1997. – V. 38-39, N. 1-3. – 218-221. 154. Drachev L.A., Drachev A.L., Chekulaeva L.N., Evstigneeva R.P., Kaulen A.D., Khitrina L.V., Khodonov A.A., Lazarova Z.R., Mitsner B.I. An investigation of the electrochemical cycle of bacteriorhodopsin analogs with modified ring. // Arch. Biochem. Biophys. – 1989. – V. 270, N 1. – P. 184-197.

113 155. Jaaskelainen T., Leppanen V.P., Parkkinen S., Parkkinen J.P.S., Khodonov A. The photochromic properties of 4–keto bacteriorhodopsin. // Optical Materials. – 1996. – V. 6, N. 4. – P. 339-345 156. Ⱦɪɚɱɟɜ Ʌ.Ⱥ., Ɂɨɪɢɧɚ ȼ.ȼ., Ɇɢɰɧɟɪ Ȼ.ɂ., ɏɢɬɪɢɧɚ Ʌ.ȼ., ɏɨɞɨɧɨɜ Ⱥ.Ⱥ., ɑɟɤɭɥɚɟɜɚ Ʌ.ɇ. Ɏɨɬɨɰɢɤɥ ɢ ɷɥɟɤɬɪɨɝɟɧɟɡɢɫ 13-ɰɢɫ ɢ ɩɨɥɧɨɫɬɶɸ-ɬɪɚɧɫ ɚɪɨɦɚɬɢɱɟɫɤɢɯ ɚɧɚɥɨɝɨɜ ɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. // Ȼɢɨɯɢɦɢɹ. – 1987. – Ɍ. 52, N. 9. – ɋ. 1559-1569. 157. ɏɢɬɪɢɧɚ Ʌ.ȼ., Ʌɚɡɚɪɟɜɚ ɐ.Ɋ. ɂɫɫɥɟɞɨɜɚɧɢɟ 13-ɰɢɫ ɢ ɩɨɥɧɨɫɬɶɸ-ɬɪɚɧɫ ɢɡɨɦɟɪɨɜ 4-ɤɟɬɨɛɚɤɬɟɪɢɨɪɨɞɨɩɫɢɧɚ. // Ȼɢɨɯɢɦɢɹ. – 1989. – Ɍ. 54, N. 1. – ɋ. 136-138. 158. Seiff F., Wallat I., Ermann P., Heyn M.P. A neutron diffraction study on the location of the polyene chain of retinal in bacteriorhodopsin. // Proc. Natl. Acad. Sci. USA. – 1985. – V. 82, N. 10. P. 3227-3231. 159. Dawson M.I., Hobbs P.D. The synthetic chemistry of retinoids. // In The Retinoids: Biology, Chemistry, and Medicine. / Eds. Sporn M.B., Roberts A.B., Goodman D.S. New York: Raven Press. – 1994. – V. 1. Chapter 2. – P. 5-178. 160. Dawson M.I., Hobbs P.D., Chan R.L.-S., Chao W.-R. Retinoic acid analogues with ring modification. Synthesis and pharmacological activity. // J. Med. Chem. – 1981. – V. 24, N. 10 – P. 1214-1223 161. Aig E., Focella A., Parrish D.R., Rosenberger M., Scott J.W., Zenchoff G.B. Synthesis of the major metabolites of (2E,4E,6E,8E)- and (2Z,4E,6E,8E)-3,7-dimethyl-9-(2,6,6trimethyl-1-cyclohexen-1-yl)-2,4,6,8-nonatetraenoic acid (retinoic acid and 13-cisretinoic acid). // Synth. Comm. – 1987. – V. 17, N. 4. – P. 419-429. 162. Rosenberger M. Retinoic acid metabolites. 1. Total synthesis of 4-hydroxy and 4oxoretinoic acid. // J. Org. Chem. – 1982. – V. 47. – P. 1698-1701. 163. Rosenberger M., Neucom C. Retinoic acid metabolites. 2. Total synthesis of rac(2E,4E,6E,8E)-3,7-dimethyl-9-(6-carboxy-2,6-dimethyl-3-oxo-1-cyclohexen-1-yl)2,4,6,8-nonatetraenoic acid and rac-(2E,4E,6E,8E)-3,7-dimethyl-9-(2,6-dimethyl-6(hydroxymethyl)3-oxo-1-cyclohexen-1-yl)-2,4,6,8-nonatetraenoic acid. // J. Org. Chem. – 1982. – V. 47, N. 9. – P. 1779-1782. 164. Rosenberger M., Neucom C. Retinoic acid metabolites. 3. Total synthesis of (2E,4E,6E,8E)-3,7-dimethyl-9-(6,6-dimethyl-2-(hydroxymethyl)-1-cyclohexen-1-yl)2,4,6,8-nonatetraenoic acid. // J. Org. Chem. – 1982. – V. 47, N. 9. – P. 1782-1785. 165. Acton N., Brossi A. Oxidation products of retinylidene dimedone. // Helv. Chim. Acta – 1980. – V. 63, F. 6, N. 145. – P. 1391-1395. 166. Singh A.K. Synthesis of oxygenated retinoids. // Synth. Commun. – 1983. – V. 13, N. 11. – P. 919-925.

114 167. Barua A.B., Olson J.A. Synthesis of 4,4-difluoro analogs of retinol and retinoic acid. // J. Lipid Res. – 1984. – V. 25, N. 3. – P. 304-309. 168. Henbest H.B., Jones E.R.H., Owen T.C. Oxidation of vitamin A1 and retinene1 by manganese dioxide. // J. Chem. Soc. – 1957. – P. 4902-4912. 169. ɋɨɤɨɥɨɜɚ ɇ.Ⱥ., Ɇɢɰɧɟɪ Ȼ.ɂ., Ɂɚɤɢɫ ȼ.ɂ. ɋɢɧɬɟɡ 4-ɤɟɬɨ- ɢ 4ɝɢɞɪɨɤɫɢɩɪɨɢɡɜɨɞɧɵɯ all-E- ɢ 13Z-ɪɟɬɢɧɚɥɟɣ ɢ ɢɯ ɜɡɚɢɦɨɞɟɣɫɬɜɢɟ ɫ ɛɚɤɬɟɪɢɨɨɩɫɢɧɨɦ. // Ȼɢɨɨɪɝɚɧ. ɯɢɦɢɹ. – 1979. – Ɍ. 5, N 7. – ɋ. 1053-1058. 170. Beischel C.J., Mani V., Govindjee R., Ebrey T.G., Knapp D.R., Crouch R.K. Ring oxidized retinals form unusual bacteriorhodopsin analogue pigments. // Photochem. Photobiol. – 1991. – V. 54, N.6. – P. 977-983. 171. Colmenares L.U., Liu R.S.H. An unexpected C-19 product from reaction of retinal with MnO2. Structure characterization and bacteriorhodopsin analogs. // Tetrahedron Lett. – 1991. – V. 32, N. 17. – P. 1933-1936. 172. Williams T.C., Mani V. Design of a helix-bundle cross-link: NMR and UV-visible spectroscopic analysis and molecular modeling of ring-oxidized retinals. // Biochemistry. – 1991. – V. 30, N. 11. – P. 2976–2988. 173. Schwieter U., Arnold W., Oberhaensli W.E., Rigassi N., Vetter W. 263. Synthesen in der Carotinoid-Reihe. Ein Beitrag zur Chromsaeure-Oxydation von Polyenen. // Helv. Chim. Acta – 1971. – V. 54, F. 8, N. 262–263. – P. 2447-2459. 174. Oyler A.R., Motto M.G., Naldi R.E., Facchine K.L., Hamburg P.F., Burinsky D.J., Dunphy R., Cotter M.L. Characterization of autoxidation products of retinoic acid. // Tetrahedron. – 1989. – V. 45, N. 24. – P. 7679-7694. 175. Karras M., Snider B.B. Opsin shift in bovine rhodopsin and bacteriorhodopsin. Comparison of two external point-charge models. // J. Am. Chem. Soc. – 1980. – V. 102, N 27. – P. 7947-7949. 176. Ɇɢɪɨɧɨɜɚ ȿ.ȼ., Ʌɟɨɧɬɶɟɜɚ ɋ.ȼ., ɒɟɜɹɤɨɜ ɋ.ȼ., Ⱥɥɟɤɫɟɟɜɚ ɋ.Ƚ., ɒɜɟɰ ȼ.ɂ., Ⱦɟɦɢɧɚ Ɉ.ȼ., Ʉɪɚɫɧɨɤɭɬɫɤɚɹ ɂ.ɋ., Ɏɢɧɤɟɥɶɲɬɟɣɧ ȿ.ɂ., ɏɨɞɨɧɨɜ Ⱥ.Ⱥ. ɍɞɨɛɧɵɣ ɫɩɨɫɨɛ ɩɨɥɭɱɟɧɢɹ ɩɪɨɢɡɜɨɞɧɵɯ ɪɟɬɢɧɚɥɹ, ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɯ ɩɨ ɬɪɢɦɟɬɢɥɰɢɤɥɨɝɟɤɫɟɧɨɜɨɦɭ ɤɨɥɶɰɭ. // Ȼɢɨɨɪɝ. ɯɢɦɢɹ. – 2002. – Ɍ. 28, N. 6. – ɋ. 535-542. 177. Maryanoff B.E., Reitz A.B. The Wittig olefination reaction and modifications involving phosphoryl-stabilized carbanions. Stereochemistry, mechanism, and selected synthetic aspects. // Chem. Rev. – 1989. – V. 89, N 4. – P. 863-927. 178. Bestmann H.J. Synthesis of polyenes via phosphonium ylides. // Pure Appl. Chem. – 1979. – V. 51, N 3. – P. 515-533.

115 179. Pommer H. Synthesen in der Vitamin A- Reihe. // Angew. Chem. – 1960. – B. 72, N 22. – S. 811-819. 180. Buddrus J. Dehydrohalogenierung von Phosphoniumhalogeniden durch Epoxide. Teil 1: Bildung der Ilide und Umsetzung mit Carbonylverbindungen. // Chem. Ber. – 1974. – B. 107, N 6. – S. 2050-2061. 181. Buddrus J., Kimpenhaus W. Dehydrohalogenierung von Phosphoniumhalogeniden durch Epoxide. Teil 2: Kinetik und Mechanismus. // Chem. Ber. – 1974. – B. 107, N 6. – S. 2062-2078. 182. ɏɨɞɨɧɨɜ Ⱥ.Ⱥ., ȿɪɟɦɢɧ ɋ.ȼ., Ɇɢɰɧɟɪ Ȼ.ɂ. ȼɥɢɹɧɢɟ ɩɪɢɪɨɞɵ ɡɚɦɟɫɬɢɬɟɥɟɣ ɧɚ ɫɬɟɪɟɨ ɢ ɪɟɝɢɨɫɟɥɟɤɬɢɜɧɨɫɬɶ ɩɪɨɰɟɫɫɚ ɝɢɞɪɢɪɨɜɚɧɢɹ ɩɨɥɢɟɧɢɧɨɜɵɯ ɫɨɟɞɢɧɟɧɢɣ ɧɚ ɩɚɥɥɚɞɢɟɜɵɯ ɤɚɬɚɥɢɡɚɬɨɪɚɯ. // Ɍɟɡ. ɞɨɤɥɚɞɨɜ VII ȼɫɟɫɨɸɡɧɨɣ ɤɨɧɮɟɪɟɧɰɢɢ “Ʉɚɬɚɥɢɬɢɱɟɫɤɢɟ ɪɟɚɤɰɢɢ ɜ ɠɢɞɤɨɣ ɮɚɡɟ”. – Ⱥɥɦɚ-ɚɬɚ. – 1988. – ɑ. 1. – ɋ. 114. 183. Ɇɢɰɧɟɪ Ȼ.ɂ., ɏɨɞɨɧɨɜ Ⱥ.Ⱥ., Ɂɜɨɧɤɨɜɚ ȿ.ɇ., ȿɜɫɬɢɝɧɟɟɜɚ Ɋ.ɉ. ɋɩɨɫɨɛ ɩɨɥɭɱɟɧɢɹ 6Z- ɢ 6ȿ-ɢɡɨɦɟɪɨɜ (2ȿ)-8-ɬɪɢɮɟɧɢɥɫɢɥɢɥɨɤɫɢ-2,6-ɞɢɦɟɬɢɥɨɤɬɚ-2,6-ɞɢɟɧ-4-ɢɧ-1ɚɥɹ. // Ⱥ.ɫ. N 1027168 (ɋɋɋɊ). – ɉɪɢɨɪɢɬɟɬ ɨɬ 30.03.1982. – Ȼɸɥɥ. ɢɡɨɛɪ. – 1983. – N 25. 184. Belrhali H., Nollert P., Royant A., Menzel C., Rosenbusch J.P., Landau E.M., PebayPeyroula E. Protein, lipid and water organization in bacteriorhodopsin crystals: a molecular view of the purple membrane at 1.9Å resolution. // Structure. – 1999. – V. 7, N. 8. – P. 909-917. 185. Luecke H., Schobert B., Richter H.T., Cartailler J.P., Lanyi J.K. Structural changes in bacteriorhodopsin during ion transport at 2 angstrom resolution. // Science. – 1999. – V. 286, N. 5438. – P. 255-260. 186. MacDonald R.E., Green R.V., Lanyi J.K. Light-activated amino acid transport systems in Halobacterium halobium envelope vesicles: role of chemical and electrical gradients. // Biochemistry. – 1977. – V. 16, N. 14. – P. 3227–3235. 187. Brown H.J., Gibbons N.E. The effect of magnesium, potassium, and iron on the growth and morphology of red halophilic bacteria. // Can. J. Microbiol. – 1955. – V. 1. – P. 486-494. 188. Oesterhelt D., Krippahl G. Phototrophic growth of halobacteria and its use for isolation of photosynthetically-deficient mutants. // Ann. Microbiol. (Inst. Pasteur) – 1983. – V. 134 B, N 1. – P.137 – 150. 189. Katznelson H., Lochhead A.G. Growth factors requirement of halophilic bacteria. // Can. J. Research. – 1952. – V. 64, N. 1. – P. 97-103.

116 190. Oesterhelt D. Reconstitution of the retinal proteins bacteriorhodopsin and halorhodopsin. // Meth. Enzymol. – 1982. – V. 88, Pt. 2 – P. 10-17. 191. Huang K.S., Bayley H., Liao M.J., London E., Khorana H.G. Refolding of an integral membrane protein. Denaturation, renaturation, and reconstitution of intact bacteriorhodopsin and two proteolytic fragments. // J. Biol. Chem. – 1981. – V. 256, N. 8. – P. 3802-3809. 192. Crouch R.K. Analog pigments of rhodopsin and bacteriorhodopsin: retinal binding site and role of the chromophore in the function of these pigments. // Chapter 6 in: Chemistry and Biology of Synthetic Retinoids. / Eds. Dawson M.I., Okamura W.H. Boca Raton: CRC Press – 1990. – P. 125-146. 193. Rao V.J., Zingoni J.P., Crouch R., Denny M., Lui R.S.H. Isomers of 3,7,11-trimethyldodeca-2,4,6,8,10-pentaenal (a linear analog of retinal) and lower homologues in their interaction with bovine opsin and bacteriorhodopsin. // Photochem. Photobiol. – 1985. – V. 41, N. 2. – P. 171-174.

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