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Ɍɪɚɧɫɤɪɢɩɰɢɹ ɩɪɨɤɚɪɢɨɬ
ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ ɩɪɨɤɚɪɢɨɬ, ɟɟ ɫɭɛɴɟɞɢɧɢɱɧɵɣ ɫɨɫɬɚɜ
ɉɪɨɤɚɪɢɨɬɵ ɫɨɞɟɪɠɚɬ ɨɞɧɭ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɭ. ȿɟ ɫɭɛɴɟɞɢɧɢɱɧɵɣ ɫɨɫɬɚɜ ɢ ɚɫɫɨɰɢɢɪɨɜɚɧɧɵɟ ɫ ɧɟɣ ɮɚɤɬɨɪɵ ɩɪɟɞɫɬɚɜɥɟɧɵ ɧɚ ɪɢɫɭɧɤɟ 1. Ⱦɥɹ ɷɥɨɧɝɚɰɢɢ ɧɟɨɛɯɨɞɢɦ ɤɨɦɩɥɟɤɫ D EE’, ɚ ɞɥɹ ɢɧɢɰɢɚɰɢɢ ɤ ɧɟɦɭ ɞɨɥɠɧɚ ɛɵɬɶ ɩɪɢɫɨɟɞɢɧɟɧɚ V ɫɭɛɴɟɞɢɧɢɰɚ. V ɫɭɛɴɟɞɢɧɢɰɵ ɪɚɡɥɢɱɚɸɬɫɹ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɬɨɝɨ, ɤɚɤɭɸ ɝɪɭɩɩɭ ɩɪɨɦɨɬɨɪɨɜ ɞɨɥɠɧɚ ɭɡɧɚɜɚɬɶ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ. ɉɨɫɥɟ ɢɧɢɰɢɚɰɢɢ V ɫɭɛɴɟɞɢɧɢɰɚ ɞɢɫɫɨɰɢɢɪɭɟɬ ɢ ɤ ɷɥɨɧɝɢɪɭɸɳɟɣ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɟ ɦɨɝɭɬ ɩɪɢɫɨɟɞɢɧɹɬɶɫɹ ɚɫɫɨɰɢɢɪɨɜɚɧɧɵɟ ɮɚɤɬɨɪɵ. Ʉɨɝɞɚ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ ɞɨɯɨɞɢɬ ɞɨ ɬɟɪɦɢɧɚɬɨɪɚ ɬɪɚɧɫɤɪɢɩɰɢɢ, ɫɢɧɬɟɡ ɊɇɄ ɨɫɬɚɧɚɜɥɢɜɚɟɬɫɹ, ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ ɢ ɫɢɧɬɟɡɢɪɨɜɚɧɧɚɹ ɊɇɄ ɞɢɫɫɨɰɢɢɪɭɸɬ. Ɋɚɫɫɦɨɬɪɢɦ ɫɬɚɞɢɢ ɷɥɨɧɝɚɰɢɨɧɧɨɝɨ ɰɢɤɥɚ ɛɨɥɟɟ ɩɨɞɪɨɛɧɨ.
ɊɇɄ ɩɨɥɢɦɟɪɚɡɚ |
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Ɏɚɤɬɨɪɵ ɢɧɢɰɢɚɰɢɢ |
D EE’V |
V |
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70 kD ɨɫɧɨɜɧɚɹ |
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20 kD ɬɪɚɧɫɩɨɪɬ ɠɟɥɟɡɚ |
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Core-ɮɟɪɦɟɧɬ |
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28 kD ɮɥɚɝɟɥɥɚ |
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D 40kD |
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32 kD ɬɟɩɥɨɜɨɣ ɲɨɤ |
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E 155kD |
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54 kD ɦɟɬɚɛɨɥɢɡɦ ɚɡɨɬɚ |
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22 kD ɬɟɩɥɨɜɨɣ ɢ |
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E’ 160kD |
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ɨɤɢɫɥɢɬɟɥɶɧɵɣ ɲɨɤ |
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38 kD ? ɷɤɫɩɪɟɫɫɢɹ ɤɚɬɚɥɚɡɵ, |
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ɷɧɞɨɧɭɤɥɟɚɡɵ III |
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Ⱥɫɫɨɰɢɢɪɨɜɚɧɧɵɟ ɮɚɤɬɨɪɵ |
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NusA |
55kD |
ɮɚɤɬɨɪ ɷɥɨɧɝɚɰɢɢ, ɩɚɭɡɵ, |
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ɚɧɬɢɬɟɪɦɢɧɚɰɢɹ ON, ɚɬɬɟɧɸɚɰɢɹ, |
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ɤɨɬɪɚɧɫɤɪɢɩɰɢɨɧɧɨɟ ɫɜɨɪɚɱɢɜɚɧɢɟ |
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ɊɇɄ |
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NusG |
21kD |
ɮɚɤɬɨɪ ɷɥɨɧɝɚɰɢɢ, ɫɭɩɪɟɫɫɢɪɭɟɬ |
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ɩɚɭɡɵ, ɚɧɬɢɬɟɪɦɢɧɚɰɢɹ ON |
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GreA |
18kD |
ɪɚɡɪɟɡɚɧɢɟ ɬɪɚɧɫɤɪɢɩɬɚ 2-3ɧ. ɜ |
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“ɚɪɟɫɬɨɜɚɧɧɵɯ” ɤɨɦɩɥɟɤɫɚɯ |
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GreB |
19kD |
ɪɚɡɪɟɡɚɧɢɟ ɬɪɚɧɫɤɪɢɩɬɚ ɞɨ 9ɧ. ɜ |
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“ɚɪɟɫɬɨɜɚɧɧɵɯ” ɤɨɦɩɥɟɤɫɚɯ |
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Mfd |
130kD ɨɫɜɨɛɨɠɞɟɧɢɟ ɊɇɄ ɩɨɥɢɦɟɪɚɡɵ, |
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(TRCF) |
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ɛɥɨɤɢɪɨɜɚɧɧɨɣ ɬɢɦɢɧɨɜɵɦ ɞɢɦɟɪɨɦ, |
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ɚɤɬɢɜɚɰɢɹ UvrABC |
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Ɏɚɤɬɨɪ ɬɟɪɦɢɧɚɰɢɢ
U 47 kD, ɝɟɤɫɚɦɟɪ
Ɋɢɫ. 1. ɋɭɛɴɟɞɢɧɢɱɧɵɣ ɫɨɫɬɚɜ ɛɚɤɬɟɪɢɚɥɶɧɨɣ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɵ ɢ ɞɨɩɨɥɧɢɬɟɥɶɧɵɟ, ɚɫɫɨɰɢɢɪɨɜɚɧɧɵɟ ɫ ɧɟɣ ɮɚɤɬɨɪɵ.
ɂɧɢɰɢɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɪɨɤɚɪɢɨɬ
ɂɧɢɰɢɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɪɨɢɫɯɨɞɢɬ ɧɚ ɫɩɟɰɢɮɢɱɟɫɤɨɦ ɭɱɚɫɬɤɟ ȾɇɄ, ɧɚɡɵɜɚɟɦɨɦ ɩɪɨɦɨɬɨɪɨɦ. Ⱦɥɹ ɭɡɧɚɜɚɧɢɹ ɩɪɨɦɨɬɨɪɚ ɧɟɨɛɯɨɞɢɦ V ɮɚɤɬɨɪ, ɩɪɢɱɟɦ ɪɚɡɥɢɱɧɵɟ V ɮɚɤɬɨɪɵ ɨɬɜɟɱɚɸɬ ɡɚ ɭɡɧɚɜɚɧɢɟ ɪɚɡɥɢɱɧɵɯ ɤɥɚɫɫɨɜ ɩɪɨɦɨɬɨɪɨɜ. Ʉɚɤ ɩɪɚɜɢɥɨ ɭɡɧɚɸɬɫɹ ɛɥɨɤɢ, ɨɬɫɬɨɹɳɢɟ ɧɚ 10 ɢ 35 ɧɭɤɥɟɨɬɢɞɨɜ ɨɬ ɬɨɱɤɢ ɧɚɱɚɥɚ ɬɪɚɧɫɤɪɢɩɰɢɢ (ɬɚɛɥ. 1).
V |
-35 |
-10 |
E.coli V70 |
TTGACA |
TATAAT |
E.coli V32 |
TCTC-CCCTTGAA |
CCCCAT-TA |
E.coli V54 |
(-24)CTGG-A |
(-12)TTGCA |
B. sub VA |
TTGACA |
TATAAT |
B. sub V% |
AGGTTTAA |
GGGTAT |
B. sub VD |
CTAAA |
CCGATAT |
B. sub VE |
ATATT |
ATACA |
B. sub VK |
AC |
CATA---T |
B. sub VH |
CAGGA |
GAATT—T |
SPO1Vgp28 |
AGGAGA |
TTT-TTT |
T4Vgp55 |
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TATAAATA |
Ɍɚɛɥɢɰɚ 1. ɉɪɨɦɨɬɨɪɧɵɟ ɨɛɥɚɫɬɢ, ɭɡɧɚɸɳɢɟɫɹ ɪɚɡɥɢɱɧɵɦɢ V-ɮɚɤɬɨɪɚɦɢ.
-35 |
-10 |
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TTGACA ɩɪɨɦɨɬɨɪ TATAAT
V D EE’
D EE’
V
D EE’
V
D EE’
Ɋɢɫ. 2. ɉɪɨɦɨɬɨɪ ɛɚɤɬɟɪɢɚɥɶɧɨɝɨ ɝɟɧɚ ɢ ɫɯɟɦɚ ɢɧɢɰɢɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ ɛɚɤɬɟɪɢɚɥɶɧɨɣ ɊɇɄɩɨɥɢɦɟɪɚɡɨɣ.
Ɇɟɯɚɧɢɡɦ ɢɧɢɰɢɚɰɢɢ ɜɤɥɸɱɚɟɬ ɧɟɫɤɨɥɶɤɨ ɨɛɪɚɬɢɦɵɯ ɫɬɚɞɢɣ: ɨɛɪɚɡɨɜɚɧɢɟ «ɡɚɤɪɵɬɨɝɨ» ɤɨɦɩɥɟɤɫɚ (ɛɟɡ ɪɚɫɩɥɟɬɚɧɢɹ ɰɟɩɟɣ ȾɇɄ), ɨɛɪɚɡɨɜɚɧɢɹ «ɨɬɤɪɵɬɨɝɨ» ɤɨɦɩɥɟɤɫɚ (ɫ ɥɨɤɚɥɶɧɵɦ ɪɚɫɩɥɟɬɚɧɢɟɦ ȾɇɄ) ɢ ɫɢɧɬɟɡ ɤɨɪɨɬɤɢɯ ɊɇɄ (ɛɟɡ ɞɢɫɫɨɰɢɚɰɢɢ V- ɮɚɤɬɨɪɚ). ɉɨɫɥɟ ɬɨɝɨ, ɤɚɤ ɫɢɧɬɟɡɢɪɭɟɬɫɹ ɮɪɚɝɦɟɧɬ ɊɇɄ ɛɨɥɟɟ 9 ɧɭɤɥɟɨɬɢɞɨɜ, V-ɮɚɤɬɨɪ ɧɟɨɛɪɚɬɢɦɨ ɞɢɫɫɨɰɢɢɪɭɟɬ ɢ ɬɪɚɧɫɤɪɢɩɰɢɹ ɜɫɬɭɩɚɟɬ ɜ ɫɬɚɞɢɸ ɷɥɨɧɝɚɰɢɢ.
Ɇɟɯɚɧɢɡɦ ɢɧɢɰɢɚɰɢɢ ɨɬɥɢɱɚɟɬɫɹ ɜ ɫɥɭɱɚɟ V54. Ɂɚɤɪɵɬɵɣ ɤɨɦɩɥɟɤɫ V54 ɫ ɊɇɄɩɨɥɢɦɟɪɚɡɨɣ ɢ ɩɪɨɦɨɬɨɪɨɦ ɫɬɚɛɢɥɟɧ ɢ ɨɛɪɚɡɨɜɚɧɢɟ ɨɬɤɪɵɬɨɝɨ ɤɨɦɩɥɟɤɫɚ ɩɪɨɢɫɯɨɞɢɬ ɬɨɥɶɤɨ ɩɪɢ ɚɤɬɢɜɚɰɢɢ ɞɪɭɝɢɦ ɛɟɥɤɨɦ (ɫɦ. ɝɥɚɜɭ ɨ ɪɟɝɭɥɹɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ).
ɋɪɚɜɧɢɬɟɥɶɧɨ ɧɟɞɚɜɧɨ ɛɵɥ ɨɬɤɪɵɬ ɪɟɝɭɥɹɬɨɪɧɵɣ ɷɥɟɦɟɧɬ ȾɇɄ, ɬ.ɧ. UP, ɜɡɚɢɦɨɞɟɣɫɬɜɭɸɳɢɣ ɫ ɋ-ɤɨɧɰɟɜɵɦ ɞɨɦɟɧɨɦ D-ɫɭɛɴɟɞɢɧɢɰɵ ɊɇɄ ɩɨɥɢɦɟɪɚɡɵ. ɗɬɢɦ ɷɥɟɦɟɧɬɨɦ ɫɧɚɛɠɟɧɵ ɧɚɢɛɨɥɟɟ ɚɤɬɢɜɧɵɟ ɤɥɟɬɨɱɧɵɟ ɩɪɨɦɨɬɨɪɵ, ɬɚɤɢɟ, ɤɚɤ ɩɪɨɦɨɬɨɪ ɝɟɧɨɜ ɪɢɛɨɫɨɦɧɨɣ ɊɇɄ.
ɗɥɨɧɝɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɪɨɤɚɪɢɨɬ
ɉɨɫɥɟ ɬɨɝɨ, ɤɚɤ V-ɮɚɤɬɨɪ ɞɢɫɫɨɰɢɢɪɭɟɬ ɫ ɷɥɨɧɝɢɪɭɸɳɟɣ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɨɣ, ɫ ɧɟɣ ɧɚɱɢɧɚɸɬ ɜɡɚɢɦɨɞɟɣɫɬɜɨɜɚɬɶ ɧɟɫɤɨɥɶɤɨ ɛɟɥɤɨɜ – ɮɚɤɬɨɪɨɜ ɷɥɨɧɝɚɰɢɢ (ɪɢɫ. 3). ȼɫɟ ɞɨɩɨɥɧɢɬɟɥɶɧɵɟ ɮɚɤɬɨɪɵ ɧɟ ɹɜɥɹɸɬɫɹ ɧɟɨɛɯɨɞɢɦɵɦɢ ɞɥɹ ɮɟɪɦɟɧɬɚɬɢɜɧɨɣ ɚɤɬɢɜɧɨɫɬɢ ɫɚɦɨɣ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɵ.
Ȼɟɥɨɤ NusA, 56kD, ɫɩɨɫɨɛɫɬɜɭɟɬ ɩɚɭɡɚɦ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɵ ɧɚ ɧɟɤɨɬɨɪɵɯ ɭɱɚɫɬɤɚɯ ȾɇɄ, ɧɟɨɛɯɨɞɢɦɵɯ ɞɥɹ ɩɪɚɜɢɥɶɧɨɝɨ ɫɜɨɪɚɱɢɜɚɧɢɹ ɜɬɨɪɢɱɧɨɣ ɫɬɪɭɤɬɭɪɵ ɊɇɄ. Ɍɚɤɢɟ ɩɚɭɡɵ ɫɨɜɟɪɲɟɧɧɨ ɧɟɨɛɯɨɞɢɦɵ ɩɪɢ ɬɪɚɧɫɤɪɢɩɰɢɢ ɮɭɧɤɰɢɨɧɚɥɶɧɵɯ (ɧɚɩɪɢɦɟɪ ɪɢɛɨɫɨɦɧɵɯ) ɊɇɄ ɢ ɫɩɨɫɨɛɫɬɜɭɸɬ ɪɟɝɭɥɹɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ ɫ ɩɨɦɨɳɶɸ ɚɬɬɟɧɸɚɰɢɢ. NusA ɜɡɚɢɦɨɞɟɣɫɬɜɭɟɬ ɤɚɤ ɫ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɨɣ, ɬɚɤ ɢ ɫ ɊɇɄ ɢ U ɮɚɤɬɨɪɨɦ ɢ ɭɱɚɫɬɜɭɟɬ ɜ ɪɟɝɭɥɹɰɢɢ ɬɟɪɦɢɧɚɰɢɢ.
NusG ɬɚɤɠɟ ɮɚɤɬɨɪ, ɚɫɫɨɰɢɢɪɨɜɚɧɵɣ ɫ ɷɥɨɧɝɢɪɭɸɳɟɣ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɨɣ. ȼ ɨɬɥɢɱɢɢ ɨɬ NusA, ɨɧ ɧɚɨɛɨɪɨɬ, ɩɨɞɚɜɥɹɟɬ ɨɫɬɚɧɨɜɤɢ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɵ. Ɉɧ ɬɚɤɠɟ ɢɝɪɚɟɬ ɪɨɥɶ ɜ ɪɟɝɭɥɹɰɢɢ ɬɟɪɦɢɧɚɰɢɢ.
Ȼɟɥɤɢ GreA ɢ GreB ɭɡɧɚɸɬ ɬɚɤ ɧɚɡɵɜɚɟɦɵɟ «ɚɪɟɫɬɨɜɚɧɧɵɟ» ɤɨɦɩɥɟɤɫɵ. ɗɬɢ ɤɨɦɩɥɟɤɫɵ ɨɛɪɚɡɭɸɬɫɹ ɟɫɥɢ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ ɫɞɜɢɝɚɟɬɫɹ ɩɪɨɬɢɜ ɯɨɞɚ ɷɥɨɧɝɚɰɢɢ. ɉɪɢ ɷɬɨɦ ɜ ɚɤɬɢɜɧɨɦ ɰɟɧɬɪɟ ɨɤɚɡɵɜɚɟɬɫɹ ɧɟ 3’-ɤɨɧɟɰ ɫɢɧɬɟɡɢɪɭɟɦɨɣ ɊɇɄ, ɚ ɊɇɄ-ȾɇɄ ɞɭɩɥɟɤɫ. Ɍɚɤɢɟ ɤɨɦɩɥɟɤɫɵ ɫɬɚɛɢɥɶɧɵ ɢ ɧɟ ɦɨɝɭɬ ɧɢ ɜɟɪɧɭɬɶɫɹ ɤ ɷɥɨɧɝɚɰɢɢ ɧɢ ɞɢɫɫɨɰɢɢɪɨɜɚɬɶ ɫɚɦɨɫɬɨɹɬɟɥɶɧɨ. Ȼɟɥɤɢ Gre ɭɡɧɚɸɬ «ɚɪɟɫɬɨɜɚɧɧɵɟ» ɤɨɦɩɥɟɤɫɵ ɢ ɫɩɨɫɨɛɫɬɜɭɸɬ ɝɢɞɪɨɥɢɡɭ ɊɇɄ-ɬɪɚɧɫɤɪɢɩɬɚ ɊɇɄ-ɩɨɥɦɢɟɪɚɡɨɣ ɜ ɟɟ ɚɤɬɢɜɧɨɦ ɰɟɧɬɪɟ. ɉɨɫɥɟ ɷɬɨɝɨ ɷɥɨɧɝɚɰɢɹ ɦɨɠɟɬ ɛɵɬɶ ɩɪɨɞɨɥɠɟɧɚ.
NusG
ɗɅɈɇȽȺɐɂə NusA D EE’
NusG
NusA D EE’
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ȺɊȿɋɌ: |
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NusA D EE’ |
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ɝɢɞɪɨɥɢɡ ɬɪɚɧɫɤɪɢɩɬɚ |
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GreA/B |
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ɌɂɆɂɇɈȼɕɃ ȾɂɆȿɊ: |
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D EE’ |
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ɞɢɫɫɨɰɢɚɰɢɹ, ɪɟɩɚɪɚɰɢɹ |
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Mfd |
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TT
Ɋɢɫ.3. ɗɥɨɧɝɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɛɚɤɬɟɪɢɚɥɶɧɨɣ ɊɇɄɩɨɥɢɦɟɪɚɡɨɣ.Ɏɭɧɤɰɢɢ ɞɨɩɨɥɧɢɬɟɥɶɧɵɯ ɮɚɤɬɨɪɨɜ.
Ⱦɪɭɝɢɦ ɩɪɟɩɹɬɫɬɜɢɟɦ ɞɥɹ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɵ ɦɨɝɭɬ ɛɵɬɶ ɞɟɮɟɤɬɵ ȾɇɄ, ɬɚɤɢɟ, ɤɚɤ ɬɢɦɢɧɨɜɵɟ ɞɢɦɟɪɵ. ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ, ɨɫɬɚɧɨɜɢɜɲɚɹ ɬɪɚɧɫɤɪɢɩɰɢɸ ɜ ɦɟɫɬɚɯ ɩɨɜɪɟɠɞɟɧɢɹ ȾɇɄ, ɭɡɧɚɟɬɫɹ ɛɟɥɤɨɦ Mfd, ɤɨɬɨɪɵɣ ɫɩɨɫɨɛɫɬɜɭɟɬ ɞɢɫɫɨɰɢɚɰɢɢ ɩɨɥɢɦɟɪɚɡɧɨɝɨ ɤɨɦɩɥɟɤɫɚ ɢ ɩɪɢɫɨɟɞɢɧɟɧɢɸ ɛɟɥɤɨɜ ɪɟɩɚɪɚɰɢɢ uvrABC ɤ ȾɇɄ.
Ɍɟɪɦɢɧɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɪɨɤɚɪɢɨɬ
Ɍɟɪɦɢɧɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɪɨɤɚɪɢɨɬ ɦɨɠɟɬ ɩɪɨɢɫɯɨɞɢɬɶ ɞɜɭɦɹ ɫɩɨɫɨɛɚɦɢ: ɫ ɭɱɚɫɬɢɟɦ Uɮɚɤɬɨɪɚ ɢ ɛɟɡ ɧɟɝɨ (ɪɢɫ. 4). ɋɨɨɬɜɟɬɫɬɜɟɧɧɨ ɪɚɡɥɢɱɚɸɬ ɞɜɚ ɬɢɩɚ ɬ.ɧ. ɬɟɪɦɢɧɚɬɨɪɨɜ, ɬ.ɟ. ɧɭɤɥɟɨɬɢɞɧɵɯ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɟɣ ȾɇɄ, ɧɚ ɤɨɬɨɪɵɯ ɩɪɨɢɫɯɨɞɢɬ ɬɟɪɦɢɧɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ. Uɧɟɡɚɜɢɫɢɦɚɹ ɬɟɪɦɢɧɚɰɢɹ ɨɛɟɫɩɟɱɢɜɚɟɬɫɹ ɊɇɄ ɲɩɢɥɶɤɨɣ ɫ ɩɨɫɥɟɞɭɸɳɟɣ ɨɥɢɝɨɭɪɢɞɢɥɨɜɨɣ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶɸ. ɒɩɢɥɶɤɚ, ɩɨ-ɜɢɞɢɦɨɦɭ, ɩɪɢɜɨɞɢɬ ɤ ɩɚɭɡɟ ɜ ɬɪɚɧɫɤɪɢɩɰɢɢ, ɚ ɨɥɢɝɨɭɪɢɞɢɥ-ɨɥɢɝɨɚɞɟɧɢɥɨɜɵɣ ɞɭɩɥɟɤɫ, ɤɚɤ ɧɚɢɦɟɧɟɟ ɫɬɚɛɢɥɶɧɵɣ, ɞɢɫɫɨɰɢɢɪɭɟɬ ɜ ɯɨɞɟ ɩɚɭɡɵ. ɉɨ-ɞɪɭɝɨɦɭ ɨɛɫɬɨɢɬ ɞɟɥɨ ɜ ɫɥɭɱɚɟ Uɡɚɜɢɫɢɦɨɣ ɬɟɪɦɢɧɚɰɢɢ. ɇɚ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢ ɊɇɄ ɞɨɥɠɟɧ ɩɪɢɫɭɬɫɬɜɨɜɚɬɶ ɨɞɧɨɰɟɩɨɱɟɱɧɵɣ ɢ ɛɨɝɚɬɵɣ ɨɫɬɚɬɤɚɦɢ ɰɢɬɢɞɢɧɚ rut-ɭɱɚɫɬɨɤ, ɫɥɭɠɚɳɢɣ ɦɟɫɬɨɦ ɩɨɫɚɞɤɢ Uɮɚɤɬɨɪɚ. ɋɚɦ Uɮɚɤɬɨɪ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɝɟɤɫɚɦɟɪɧɭɸ NTPɚɡɭ, ɫɩɨɫɨɛɧɭɸ ɢɫɩɨɥɶɡɨɜɚɬɶ ɷɧɟɪɝɢɸ ɝɢɞɪɨɥɢɡɚ ɧɭɤɥɟɨɡɢɞɬɪɢɮɨɫɮɚɬɨɜ ɞɥɹ ɩɪɨɞɜɢɠɟɧɢɹ ɩɨ ɊɇɄ ɜ ɫɬɨɪɨɧɭ ɟɟ 3’-ɤɨɧɰɚ. Ʉɨɝɞɚ Uɮɚɤɬɨɪ “ɞɨɝɨɧɹɟɬ” ɪɚɛɨɬɚɸɳɭɸ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɭ, ɱɬɨ ɬɚɤɠɟ ɫɥɭɱɚɟɬɫɹ ɜ ɦɟɫɬɚɯ ɟɟ ɜɪɟɦɟɧɧɨɣ ɨɫɬɚɧɨɜɤɢ, ɩɪɨɢɫɯɨɞɢɬ ɬɟɪɦɢɧɚɰɢɹ. ȼ ɷɬɨɦ ɩɪɨɰɟɫɫɟ ɬɚɤɠɟ ɭɱɚɫɬɜɭɸɬ ɚɫɫɨɰɢɢɪɨɜɚɧɧɵɟ ɫ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɨɣ ɮɚɤɬɨɪɵ NusA ɢ NusG.
Uɡɚɜɢɫɢɦɚɹ
ɬɟɪɦɢɧɚɰɢɹ
NTP NDP
NusG
NusA
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ɩɨɫɚɞɤɚ U |
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D EE’ |
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ɩɚɭɡɚ |
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ɌȿɊɆɂɇȺɐɂə |
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Uɧɟɡɚɜɢɫɢɦɚɹ |
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ɬɟɪɦɢɧɚɰɢɹ |
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D EE’
Ɋɢɫ. 4. Ⱥɥɶɬɟɪɧɚɬɢɜɧɵɟ ɦɟɯɚɧɢɡɦɵ ɬɟɪɦɢɧɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɪɨɤɚɪɢɨɬ.
Ɋɟɝɭɥɹɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ
Ɍɪɚɧɫɤɪɢɩɰɢɹ ɦɨɠɟɬ ɪɟɝɭɥɢɪɨɜɚɬɶɫɹ ɧɚ ɥɸɛɨɣ ɫɬɚɞɢɢ, ɬ.ɟ. ɢɧɢɰɢɚɰɢɢ, ɷɥɨɧɝɚɰɢɢ ɢ ɬɟɪɦɢɧɚɰɢɢ. ȼɩɨɥɧɟ ɥɨɝɢɱɧɨ, ɜɩɪɨɱɟɦ, ɱɬɨ ɧɚɢɛɨɥɟɟ ɱɚɫɬɨ ɪɟɝɭɥɢɪɭɟɬɫɹ ɢɧɢɰɢɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ.
Ɋɟɝɭɥɹɰɢɹ ɢɧɢɰɢɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ
ɉɪɢ ɧɟɨɛɯɨɞɢɦɨɫɬɢ ɫɭɳɟɫɬɜɟɧɧɨ ɢɡɦɟɧɢɬɶ ɫɩɟɤɬɪ ɫɢɧɬɟɡɢɪɭɟɦɵɯ ɜ ɤɥɟɬɤɟ ɛɟɥɤɨɜ, ɱɚɫɬɨ ɡɚɦɟɳɚɟɬɫɹ ɫɢɝɦɚ ɮɚɤɬɨɪ. Ɍɚɤɢɦ ɫɩɨɫɨɛɨɦ, ɧɚɩɪɢɦɟɪ, ɪɟɝɭɥɢɪɭɟɬɫɹ ɤɚɬɚɛɨɥɢɡɦ ɚɡɨɬɚ (ɪɢɫ. 5). ɂɧɞɢɤɚɬɨɪɨɦ ɤɨɥɢɱɟɫɬɜɚ ɛɢɨɥɨɝɢɱɟɫɤɢ ɫɜɹɡɚɧɧɨɝɨ ɚɡɨɬɚ ɜ ɤɥɟɬɤɟ ɹɜɥɹɟɬɫɹ ɫɨɨɬɧɨɲɟɧɢɟ ɤɨɧɰɟɧɬɪɚɰɢɣ ɝɥɭɬɚɦɢɧɚ ɢ ɤɟɬɨɝɥɭɬɚɪɚɬɚ. ɉɪɢ ɧɟɞɨɫɬɚɬɤɟ ɚɡɨɬɚ ɜ ɤɥɟɬɤɟ ɧɚɱɢɧɚɟɬ ɧɚɤɚɩɥɢɜɚɬɶɫɹ ɤɟɬɨɝɥɭɬɚɪɚɬ, ɜɡɚɢɦɨɞɟɣɫɬɜɭɸɳɢɣ ɫ ɪɟɝɭɥɹɬɨɪɧɵɦ ɛɟɥɤɨɦ UTase/UR, ɫɩɨɫɨɛɧɵɦ ɜ ɩɪɢɫɭɬɫɬɜɢɢ ɤɟɬɨɝɥɭɬɚɪɚɬɚ ɭɪɢɞɢɥɢɪɨɜɚɬɶ PII ɛɟɥɨɤ. ȼ ɩɪɢɫɭɬɫɬɜɢɢ ɝɥɭɬɚɦɚɬɚ, ɧɚɨɛɨɪɨɬ, ɚɤɬɢɜɢɪɭɟɬɫɹ ɫɩɨɫɨɛɧɨɫɬɶ UTase/UR ɫɧɢɦɚɬɶ ɨɫɬɚɬɨɤ ɭɪɢɞɢɥɨɜɨɣ ɤɢɫɥɨɬɵ ɫ PII. ɂɧɮɨɪɦɚɰɢɹ ɨ ɛɚɥɚɧɫɟ ɚɡɨɬɚ ɩɟɪɟɞɚɟɬɫɹ ɛɟɥɤɭ PII. ȼ ɫɜɨɸ ɨɱɟɪɟɞɶ, ɧɟɭɪɢɞɢɥɢɪɨɜɚɧɧɚɹ ɮɨɪɦɚ ɛɟɥɤɚ PII ɫɩɨɫɨɛɧɚ ɫɜɹɡɵɜɚɬɶ ɪɟɝɭɥɹɬɨɪɧɵɣ ɛɟɥɨɤ NtrB ɢ ɚɤɬɢɜɢɪɨɜɚɬɶ ɟɝɨ ɮɨɫɮɚɬɚɡɧɭɸ ɚɤɬɢɜɧɨɫɬɶ. ɗɬɚ ɚɤɬɢɜɧɨɫɬɶ ɧɚɩɪɚɜɥɟɧɚ ɧɚ ȾɇɄɫɜɹɡɵɜɚɸɳɢɣ ɛɟɥɨɤ NtrC. ɉɪɢ ɢɡɛɵɬɤɟ ɚɡɨɬɚ ɜ ɤɥɟɬɤɟ ɢɧɮɨɪɦɚɰɢɹ, ɜ ɜɢɞɟ ɤɚɫɤɚɞɚ ɯɢɦɢɱɟɫɤɢɯ ɦɨɞɢɮɢɤɚɰɢɣ ɩɪɢɜɨɞɢɬ ɤ ɧɟɦɨɞɢɮɢɰɢɪɨɜɚɧɧɨɣ (ɞɟɮɨɫɮɨɪɢɥɢɪɨɜɚɧɧɨɣ) ɮɨɪɦɟ NtrC. ɉɪɢ ɧɟɞɨɫɬɚɬɤɟ ɚɡɨɬɚ, ɧɚɨɛɨɪɨɬ, ɫɜɨɛɨɞɧɵɣ (ɨɬ PII) NtrB ɩɟɪɟɧɨɫɢɬ ɮɨɫɮɚɬ ɫ ATP ɧɚ NtrC. Ɏɨɫɮɨɪɢɥɢɪɨɜɚɧɧɵɣ NtrC ɜɡɚɢɦɨɞɟɣɫɬɜɭɟɬ ɫ ɤɨɦɩɥɟɤɫɨɦ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɵ ɫ V, ɤɨɬɨɪɵɣ ɧɚɯɨɞɢɬɫɹ ɧɚ ɩɪɨɦɨɬɨɪɚɯ ɝɟɧɨɜ ɭɱɚɫɬɜɭɸɳɢɯ ɜ ɦɟɬɚɛɨɥɢɡɦɟ ɚɡɨɬɚ.
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2NH3+2[H] |
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HO |
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H2N |
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ɇȿȾɈɋɌȺɌɈɄ |
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ɂɁȻɕɌɈɄ |
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ɝɥɭɬɚɦɢɧ |
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UTase/UR |
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PII |
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UMP |
NtrB 
NtrB PII
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NtrC NtrC 
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ɊɇɄ |
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V54 ɩɨɥɢɦɟɪɚɡɚ |
Ɋɢɫ. 5. Ɋɟɝɭɥɹɰɢɹ ɦɟɬɚɛɨɥɢɡɦɚ ɚɡɨɬɢɫɬɵɯ ɫɨɟɞɢɧɟɧɢɣ ɫ ɩɨɦɨɳɶɸ ɚɤɬɢɜɚɰɢɢ V.
ɂɬɚɤ, ɩɪɢ ɧɟɞɨɫɬɚɬɤɟ ɚɡɨɬɚ ɩɪɨɢɫɯɨɞɢɬ ɚɤɬɢɜɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɝɟɧɨɜ, ɧɟɨɛɯɨɞɢɦɵɯ ɞɥɹ ɜɨɫɫɬɚɧɨɜɥɟɧɢɹ ɚɡɨɬɧɨɝɨ ɛɚɥɚɧɫɚ. Ⱦɚɧɧɵɣ ɩɪɢɦɟɪ ɩɨɤɚɡɵɜɚɟɬ ɞɜɚ ɦɟɯɚɧɢɡɦɚ, ɲɢɪɨɤɨ ɢɫɩɨɥɶɡɭɸɳɢɯɫɹ ɞɥɹ ɪɟɝɭɥɹɰɢɢ ɛɟɥɤɨɜɵɯ ɫɢɫɬɟɦ. ȼɨ-ɩɟɪɜɵɯ, ɢɫɩɨɥɶɡɨɜɚɧɢɟ ɤɨɜɚɥɟɧɬɧɨɣ ɦɨɞɢɮɢɤɚɰɢɢ, ɧɚɩɪɢɦɟɪ ɮɨɫɮɨɪɢɥɢɪɨɜɚɧɢɹ, ɞɥɹ ɢɡɦɟɧɟɧɢɹ ɚɤɬɢɜɧɨɫɬɢ ɛɟɥɤɨɜ. ȼɨ-ɜɬɨɪɵɯ, ɢɫɩɨɥɶɡɨɜɚɧɢɟ ɦɧɨɝɨɫɬɭɩɟɧɱɚɬɵɯ “ɤɚɫɤɚɞɨɜ” ɪɟɝɭɥɹɬɨɪɧɵɯ ɛɟɥɤɨɜ. Ɍɚɤɢɟ ɤɚɫɤɚɞɵ ɧɟɨɛɯɨɞɢɦɵ ɞɥɹ ɭɫɢɥɟɧɢɹ ɫɢɝɧɚɥɚ. Ɍ.ɤ. ɨɞɢɧ ɚɤɬɢɜɢɪɨɜɚɧɧɵɣ ɩɟɪɜɢɱɧɵɣ ɪɟɝɭɥɹɬɨɪɧɵɣ ɮɟɪɦɟɧɬ (ɜ ɞɚɧɧɨɦ ɫɥɭɱɚɟ UTase/UR) ɦɨɠɟɬ ɦɨɞɢɮɢɰɢɪɨɜɚɬɶ ɦɧɨɝɨ PII ɛɟɥɤɨɜ, ɚ ɜ ɫɜɨɸ ɨɱɟɪɟɞɶ ɤɚɠɞɵɣ ɦɨɞɢɮɢɰɢɪɨɜɚɧɧɵɣ PII ɜ ɤɨɦɩɥɟɤɫɟ ɫ NtrB ɚɤɬɢɜɢɪɭɟɬ ɦɧɨɝɨ NtrC. Ʉɪɨɦɟ ɬɨɝɨ, ɫɬɭɩɟɧɱɚɬɵɟ ɤɚɫɤɚɞɵ ɩɨɡɜɨɥɹɸɬ ɩɪɢɧɢɦɚɬɶ “ɫɢɝɧɚɥɵ” ɨɬ ɦɧɨɝɢɯ ɞɪɭɝɢɯ ɪɟɝɭɥɹɬɨɪɧɵɯ ɫɢɫɬɟɦ.
Ʉɥɚɫɫɢɱɟɫɤɢɣ ɩɪɢɦɟɪ ɪɟɝɭɥɢɪɭɟɦɨɣ ɢɧɢɰɢɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ, ɤɨɝɞɚ ɪɟɲɟɧɢɟ ɞɨɥɠɧɨ ɩɪɢɧɢɦɚɬɶɫɹ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɞɜɭɯ ɢɫɬɨɱɧɢɤɨɜ ɢɧɮɨɪɦɚɰɢɢ, - ɥɚɤɬɨɡɧɵɣ ɨɩɟɪɨɧ (ɪɢɫ. 6). Ɉɩɟɪɨɧ ɤɨɞɢɪɭɟɬ ɛɟɥɤɢ ɤɚɬɚɛɨɥɢɡɦɚ ɥɚɤɬɨɡɵ, ɦɟɧɟɟ ɩɪɟɞɩɨɱɬɢɬɟɥɶɧɨɝɨ ɩɨ
ɫɪɚɜɧɟɧɢɸ ɫ ɝɥɸɤɨɡɨɣ ɢɫɬɨɱɧɢɤɚ ɷɧɟɪɝɢɢ. ɋɥɟɞɨɜɚɬɟɥɶɧɨ, ɬɪɚɧɫɤɪɢɩɰɢɹ ɞɨɥɠɧɚ ɩɪɨɢɫɯɨɞɢɬɶ ɬɨɥɶɤɨ ɜ ɬɨɦ ɫɥɭɱɚɟ, ɟɫɥɢ ɧɟɬ ɝɥɸɤɨɡɵ ɢ ɟɫɬɶ ɥɚɤɬɨɡɚ. ɉɨɞɨɛɧɚɹ ɪɟɝɭɥɹɰɢɹ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɞɜɭɦɹ ɛɟɥɤɚɦɢ: LacI – ɪɟɩɪɟɫɫɨɪɨɦ ɢ ɛɟɥɤɨɦ ɚɤɬɢɜɚɬɨɪɨɦ ɤɚɬɚɛɨɥɢɡɦɚ CAP. LacI ɛɟɥɨɤ ɫɩɨɫɨɛɟɧ ɨɛɪɚɡɨɜɵɜɚɬɶ ɬɟɬɪɚɦɟɪɵ ɢ ɫɜɹɡɵɜɚɬɶɫɹ ɫ ȾɇɄ ɬɨɥɶɤɨ ɜ ɨɬɫɭɬɫɬɜɢɢ ɚɥɥɨ ɥɚɤɬɨɡɵ. Lac-ɪɟɩɪɟɫɫɨɪ ɦɨɠɟɬ ɫɜɹɡɵɜɚɟɬɫɹ ɬɨɥɶɤɨ ɫ ɭɱɚɫɬɤɨɦ ȾɇɄ, ɧɚɡɵɜɚɟɦɵɦ ɨɩɟɪɚɬɨɪɨɦ, ɬɪɢ ɤɨɩɢɢ ɤɨɬɨɪɨɝɨ ɧɚɯɨɞɹɬɫɹ ɜ ɪɚɣɨɧɟ lac-ɨɩɟɪɨɧɚ, ɜ ɬɨɦ ɱɢɫɥɟ ɢ ɜ ɩɪɨɦɨɬɨɪɧɨɣ ɨɛɥɚɫɬɢ. ȿɫɥɢ ɥɚɤɬɨɡɚ ɧɟɞɨɫɬɭɩɧɚ, LacI ɫɜɹɡɵɜɚɟɬɫɹ ɫ ȾɇɄ ɢ lac-ɨɩɟɪɨɧ ɧɟ ɬɪɚɧɫɤɪɢɛɢɪɭɟɬɫɹ.
ȿɫɬɶ ɝɥɸɤɨɡɚ, ɧɟɬ ɥɚɤɬɨɡɵ
O O
LacI LacI
P
ɇɟɬ ɝɥɸɤɨɡɵ, ɧɟɬ ɥɚɤɬɨɡɵ
O O
CAP
LacI LacI
P
ȿɫɬɶ ɝɥɸɤɨɡɚ, ɟɫɬɶ ɥɚɤɬɨɡɚ
P O LacZ O
ɇɟɬ ɝɥɸɤɨɡɵ, ɟɫɬɶ ɥɚɤɬɨɡɚ
CAP ɊɇɄ
ɩɨɥɢɦɟɪɚɡɚ
P O LacZ O
Ɋɢɫ. 6. Ɇɨɞɟɥɶ ɪɟɝɭɥɹɰɢɢ ɥɚɤɬɨɡɧɨɝɨ ɨɩɟɪɨɧɚ.
ȿɫɥɢ ɫɜɹɡɵɜɚɧɢɟ LacI ɫ ȾɇɄ ɩɪɟɩɹɬɫɬɜɭɟɬ ɬɪɚɧɫɤɪɢɩɰɢɢ, ɬɨ ɫɜɹɡɵɜɚɧɢɟ ɛɟɥɤɚɚɤɬɢɜɚɬɨɪɚ ɤɚɬɚɛɨɥɢɡɦɚ (CAP) ɧɚɨɛɨɪɨɬ, ɧɟɨɛɯɨɞɢɦɨ ɞɥɹ ɢɧɢɰɢɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ ɫ ɞɚɧɧɨɝɨ ɩɪɨɦɨɬɨɪɚ. ɉɪɢ ɧɟɞɨɫɬɚɬɤɟ ɝɥɸɤɨɡɵ ɜ ɤɥɟɬɤɟ ɧɚɱɢɧɚɟɬ ɧɚɤɚɩɥɢɜɚɬɶɫɹ ɰɢɤɥɢɱɟɫɤɢɣ AMP (cAMP), ɩɪɢ ɫɜɹɡɵɜɚɧɢɢ ɤɨɬɨɪɨɝɨ ɫ CAP, ɩɨɫɥɟɞɧɢɣ ɩɪɢɧɢɦɚɟɬ ɤɨɧɮɨɪɦɚɰɢɸ, ɧɟɨɛɯɨɞɢɦɭɸ ɞɥɹ ɫɜɹɡɵɜɚɧɢɹ ɨɩɪɟɞɟɥɟɧɧɵɯ ɭɱɚɫɬɤɨɜ ȾɇɄ. ɉɪɢ ɷɬɨɦ, CAP ɦɨɠɟɬ ɤɚɤ ɚɤɬɢɜɢɪɨɜɚɬɶ, ɬɚɤ ɢ ɩɨɞɚɜɥɹɬɶ ɬɪɚɧɫɤɪɢɩɰɢɸ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɩɨɥɨɠɟɧɢɹ, ɡɚɧɢɦɚɟɦɨɝɨ CAP ɨɬɧɨɫɢɬɟɥɶɧɨ ɩɪɨɦɨɬɨɪɚ. ɋɜɹɡɵɜɚɧɢɟ CAP ɜɤɥɸɱɚɟɬ ɬɪɚɧɫɤɪɢɩɰɢɸ ɥɚɤɬɨɡɧɨɝɨ ɨɩɟɪɨɧɚ. ɂɬɚɤ, ɬɪɚɧɫɤɪɢɩɰɢɹ lac-ɨɩɟɪɨɧɚ ɩɪɨɢɫɯɨɞɢɬ ɬɨɥɶɤɨ ɟɫɥɢ ɜ ɫɪɟɞɟ ɨɬɫɭɬɫɬɜɭɟɬ ɝɥɸɤɨɡɚ ɢ ɩɪɢɫɭɬɫɬɜɭɟɬ ɥɚɤɬɨɡɚ. ɉɨɞɨɛɧɚɹ ɪɟɝɭɥɹɰɢɹ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɫ ɩɨɦɨɳɶɸ ȾɇɄ-ɫɜɹɡɵɜɚɸɳɢɯ ɛɟɥɤɨɜ, ɨɞɢɧ ɢɡ ɤɨɬɨɪɵɯ ɚɤɬɢɜɢɪɭɟɬ, ɚ ɞɪɭɝɨɣ ɢɧɝɢɛɢɪɭɟɬ ɬɪɚɧɫɤɪɢɩɰɢɸ.
ɇȿȾɈɋɌȺɌɈɄ UTP
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ɦɊɇɄ |
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5’ AAUUUG |
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ɩɨɥɢɦɟɪɚɡɚ |
ɂɁȻɕɌɈɄ UTP
ɦɊɇɄ |
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ɊɇɄ |
5’ AAUUUUUUUUUUUUUU ɩɨɥɢɦɟɪɚɡɚ |
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Ɋɢɫ. 7. Ɋɟɝɭɥɹɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɢɪɢɦɢɞɢɧɨɜɨɝɨ ɨɩɟɪɨɧɚ: ɪɟɢɬɟɪɚɬɢɜɧɚɹ ɢɧɢɰɢɚɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ.
Ʉɪɨɦɟ ɡɚɦɟɧɵ V-ɮɚɤɬɨɪɨɜ ɢ ɢɫɩɨɥɶɡɨɜɚɧɢɹ ɛɟɥɤɨɜ-ɪɟɝɭɥɹɬɨɪɨɜ ɪɹɞ ɨɩɟɪɨɧɨɜ ɪɟɝɭɥɢɪɭɟɬɫɹ ɬɪɟɬɶɢɦ ɫɩɨɫɨɛɨɦ, ɪɟɢɬɟɪɚɬɢɜɧɨɣ ɬɪɚɧɫɤɪɢɩɰɢɟɣ. ɂɡɜɟɫɬɧɨ, ɱɬɨ ɩɪɢ ɢɧɢɰɢɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɟɪɜɵɟ ɧɟɫɤɨɥɶɤɨ ɩɨɥɢɦɟɪɢɡɨɜɚɧɧɵɯ ɧɭɤɥɟɨɬɢɞɨɜ ɩɥɨɯɨ ɫɜɹɡɚɧɵ ɜ ɬɪɚɧɫɤɪɢɩɰɢɨɧɧɨɦ ɤɨɦɩɥɟɤɫɟ. ɗɬɚ ɨɫɨɛɟɧɧɨɫɬɶ ɢɫɩɨɥɶɡɭɟɬɫɹ ɩɪɢ ɪɟɝɭɥɹɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɢɪɢɦɢɞɢɧɨɜɨɝɨ ɨɩɟɪɨɧɚ. ȿɫɥɢ ɤɨɧɰɟɧɬɪɚɰɢɹ UTP ɜ ɤɥɟɬɤɟ ɜɵɫɨɤɚ, ɧɟɬ ɧɭɠɞɵ ɬɪɚɧɫɤɪɢɛɢɪɨɜɚɬɶ ɷɬɨɬ ɨɩɟɪɨɧ. ȼ ɧɚɱɚɥɟ ɨɩɟɪɨɧɚ ɪɚɫɩɨɥɨɠɟɧ ɤɨɪɨɬɤɢɣ ɭɱɚɫɬɨɤ, ɤɨɞɢɪɭɸɳɢɣ ɨɥɢɝɨɭɪɢɞɢɥɨɜɭɸ ɤɢɫɥɨɬɭ. ɉɪɢ ɜɵɫɨɤɨɣ ɤɨɧɰɟɧɬɪɚɰɢɢ UTP ɨɥɢɝɨU ɮɪɚɝɦɟɧɬ ɧɚɱɢɧɚɟɬ ɩɪɨɫɤɚɥɶɡɵɜɚɬɶ ɩɨ ȾɇɄ, ɬɚɤ, ɱɬɨ ɫɢɧɬɟɡɢɪɭɟɬɫɹ ɞɥɢɧɧɵɟ ɨɥɢɝɨɭɪɢɞɢɥɨɜɵɟ ɮɪɚɝɦɟɧɬɵ, ɱɬɨ ɩɪɢɜɨɞɢɬ ɤ ɬɟɪɦɢɧɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ (ɪɢɫ. 7).
Ʉɪɨɦɟ ɩɢɪɢɦɢɞɢɧɨɜɨɝɨ ɨɩɟɪɨɧɚ, ɨɩɟɪɨɧɵ ɪɢɛɨɫɨɦɧɵɯ ɊɇɄ ɬɚɤɠɟ ɪɟɝɭɥɢɪɭɸɬɫɹ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɤɨɧɰɟɧɬɪɚɰɢɢ ɧɭɤɥɟɨɬɢɞɬɪɢɮɨɫɮɚɬɨɜ. ɉɪɨɦɨɬɨɪɵ ɝɟɧɨɜ ɪɊɇɄ ɨɤɚɡɵɜɚɸɬɫɹ ɨɱɟɧɶ ɱɭɜɫɬɜɢɬɟɥɶɧɵɦɢ ɤ ɭɪɨɜɧɸ ɢɧɢɰɢɢɪɭɸɳɢɯ ɧɭɤɥɟɨɬɢɞɨɜ – ATP ɢ GTP. Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɩɪɢ ɫɥɢɲɤɨɦ ɜɵɫɨɤɨɢɧɬɟɧɫɢɜɧɨɣ ɬɪɚɧɫɥɹɰɢɢ ɢɥɢ ɜ ɭɫɥɨɜɢɹɯ ɧɟɞɨɫɬɚɬɤɚ ɷɧɟɪɝɢɢ, ɤɨɝɞɚ ɫɭɳɟɫɬɜɟɧɧɨ ɩɨɧɢɠɚɟɬɫɹ ɭɪɨɜɟɧɶ ATP ɢ GTP, ɬɪɚɧɫɤɪɢɩɰɢɹ ɝɟɧɨɜ ɪɊɇɄ ɩɪɢɨɫɬɚɧɚɜɥɢɜɚɟɬɫɹ.
Ɋɟɝɭɥɹɰɢɹ ɷɥɨɧɝɚɰɢɢ ɢ ɬɟɪɦɢɧɚɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ
Ʉɪɨɦɟ ɫɬɚɞɢɢ ɢɧɢɰɢɚɰɢɢ, ɬɪɚɧɫɤɪɢɩɰɢɹ ɦɨɠɟɬ ɪɟɝɭɥɢɪɨɜɚɬɶɫɹ ɬɚɤɠɟ ɫ ɩɨɦɨɳɶɸ ɩɪɟɠɞɟɜɪɟɦɟɧɧɨɣ ɬɟɪɦɢɧɚɰɢɢ, ɬ.ɟ. ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɧɭɠɞ ɤɥɟɬɤɢ ɞɨɥɠɟɧ ɨɫɭɳɟɫɬɜɥɹɬɶɫɹ ɜɵɛɨɪ ɦɟɠɞɭ ɷɥɨɧɝɚɰɢɟɣ ɢ ɬɟɪɦɢɧɚɰɢɟɣ. ɇɚɢɛɨɥɟɟ ɩɪɨɫɬɨɣ ɫɩɨɫɨɛ ɨɫɬɚɧɨɜɢɬɶ ɬɪɚɧɫɤɪɢɩɰɢɸ – ɫɜɹɡɚɬɶ ɛɟɥɨɤ-ɪɟɝɭɥɹɬɨɪ ɧɚ ɩɭɬɢ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɵ. Ɍɚɤ ɩɪɨɢɫɯɨɞɢɬ ɜ ɫɥɭɱɚɟ ɨɩɟɪɨɧɚ, ɨɬɜɟɬɫɬɜɟɧɧɨɝɨ ɡɚ ɫɢɧɬɟɡ ɩɭɪɢɧɨɜɵɯ ɧɭɤɥɟɨɬɢɞɨɜ. Ⱦɜɚ ɛɨɥɟɟ ɫɥɨɠɧɵɯ ɦɟɯɚɧɢɡɦɚ ɪɟɝɭɥɢɪɭɟɦɨɣ ɬɟɪɦɢɧɚɰɢɢ ɷɬɨ ɚɧɬɢɬɟɪɦɢɧɚɰɢɹ ɢ ɚɬɬɟɧɸɚɰɢɹ.
Ⱥɬɬɟɧɸɚɰɢɹ – ɷɬɨ ɪɟɝɭɥɢɪɭɟɦɚɹ ɬɟɪɦɢɧɚɰɢɹ ɡɚɜɢɫɹɳɚɹ ɨɬ ɨɬɧɨɫɢɬɟɥɶɧɨɣ ɫɬɚɛɢɥɢɡɚɰɢɢ ɚɥɶɬɟɪɧɚɬɢɜɧɵɯ ɜɬɨɪɢɱɧɵɯ ɫɬɪɭɤɬɭɪ ɬɪɚɧɫɤɪɢɛɢɪɭɟɦɨɣ ɦɊɇɄ. Ʉɚɤ ɢɡɜɟɫɬɧɨ, Uɧɟɡɚɜɢɫɢɦɵɣ ɬɟɪɦɢɧɚɬɨɪ ɩɪɟɞɫɬɚɜɥɹɟɬ ɫɨɛɨɣ ɊɇɄ ɲɩɢɥɶɤɭ ɢ ɩɨɫɥɟɞɭɸɳɢɣ ɨɥɢɝɨɭɪɢɞɢɥɨɜɵɣ ɭɱɚɫɬɨɤ. ȿɫɥɢ ɛɭɞɟɬ ɫɬɚɛɢɥɢɡɢɪɨɜɚɧɚ ɚɥɶɬɟɪɧɚɬɢɜɧɚɹ ɜɬɨɪɢɱɧɚɹ ɫɬɪɭɤɬɭɪɚ, ɝɞɟ ɨɥɢɝɨɭɪɢɞɢɥɨɜɵɣ ɮɪɚɝɦɟɧɬ ɭɞɚɥɟɧ ɨɬ ɛɥɢɠɚɣɲɟɣ ɲɩɢɥɶɤɢ, ɬɟɪɦɢɧɚɰɢɹ ɧɟ ɛɭɞɟɬ ɩɪɨɢɫɯɨɞɢɬɶ. ȿɫɬɶ ɧɟɫɤɨɥɶɤɨ ɮɚɤɬɨɪɨɜ, ɫɬɚɛɢɥɢɡɢɪɭɸɳɢɯ ɨɞɧɭ ɢɡ ɚɥɶɬɟɪɧɚɬɢɜɧɵɯ ɜɬɨɪɢɱɧɵɯ ɫɬɪɭɤɬɭɪ ɊɇɄ ɜɨ ɜɪɟɦɹ ɬɪɚɧɫɤɪɢɩɰɢɢ.
Ɍɪɚɧɫɤɪɢɩɰɢɹ ɬɪɢɩɬɨɮɚɧɨɜɨɝɨ ɨɩɟɪɨɧɚ E. coli (ɪɢɫ. 8) ɞɨɥɠɧɚ ɩɪɨɢɫɯɨɞɢɬɶ ɬɨɥɶɤɨ ɤɨɝɞɚ ɤɨɧɰɟɧɬɪɚɰɢɹ ɬɪɢɩɬɨɮɚɧɚ ɜ ɤɥɟɬɤɟ ɦɚɥɚ. ȼ ɧɚɱɚɥɟ trp-ɨɩɟɪɨɧɚ ɡɚɤɨɞɢɪɨɜɚɧ ɤɨɪɨɬɤɢɣ ɩɟɩɬɢɞ, ɛɨɝɚɬɵɣ ɨɫɬɚɬɤɚɦɢ ɬɪɢɩɬɨɮɚɧɚ. ɉɨɫɤɨɥɶɤɭ ɬɪɚɧɫɥɹɰɢɹ ɦɊɇɄ ɭ ɤɢɲɟɱɧɨɣ ɩɚɥɨɱɤɢ ɩɪɨɢɫɯɨɞɢɬ ɤɨɬɪɚɧɫɤɪɢɩɰɢɨɧɧɨ, ɥɢɞɟɪɧɵɣ ɭɱɚɫɬɨɤ ɦɊɇɄ ɧɚɱɢɧɚɟɬ ɬɪɚɧɫɥɢɪɨɜɚɬɶɫɹ ɫɪɚɡɭ ɩɨɫɥɟ ɫɢɧɬɟɡɚ. Ⱦɥɹ ɥɢɞɟɪɧɨɝɨ ɭɱɚɫɬɤɚ ɦɊɇɄ ɫɭɳɟɫɬɜɭɸɬ ɞɜɟ ɚɥɶɬɟɪɧɚɬɢɜɧɵɟ ɜɬɨɪɢɱɧɵɟ ɫɬɪɭɤɬɭɪɵ, ɨɞɧɚ ɫɩɨɫɨɛɫɬɜɭɸɳɚɹ ɩɪɟɠɞɟɜɪɟɦɟɧɧɨɣ ɬɟɪɦɢɧɚɰɢɢ, ɞɪɭɝɚɹ ɧɟɬ.
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Ɋɢɫ. 8. Ɋɟɝɭɥɹɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ trp-ɨɩɟɪɨɧɚ E. coli.
ȿɫɥɢ ɪɢɛɨɫɨɦɚ ɛɵɫɬɪɨ ɬɪɚɧɫɥɢɪɭɟɬ ɥɢɞɟɪɧɵɣ ɭɱɚɫɬɨɤ ɦɊɇɄ, ɱɬɨ ɯɚɪɚɤɬɟɪɧɨ ɞɥɹ ɜɵɫɨɤɢɯ ɤɨɧɰɟɧɬɪɚɰɢɣ ɬɪɢɩɬɨɮɚɧɚ, ɨɧɚ ɪɚɫɩɥɚɜɥɹɟɬ ɷɥɟɦɟɧɬ ɜɬɨɪɢɱɧɨɣ ɫɬɪɭɤɬɭɪɵ, ɩɪɟɩɹɬɫɬɜɭɸɳɢɣ ɬɟɪɦɢɧɚɰɢɢ ɢ, ɬɚɤɢɦ ɨɛɪɚɡɨɦ, ɫɬɚɛɢɥɢɡɢɪɭɟɬ ɜɬɨɪɢɱɧɭɸ ɫɬɪɭɤɬɭɪɭ ɬɟɪɦɢɧɚɬɨɪɚ.
ɇȿȾɈɋɌȺɌɈɄ UTP
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Ɋɢɫ. 9. Ɋɟɝɭɥɹɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɩɢɪɢɦɢɞɢɧɨɜɨɝɨ ɨɩɟɪɨɧɚ ɫ ɩɨɦɨɳɶɸ ɚɬɬɟɧɸɚɰɢɢ.
ȿɫɥɢ ɜ ɪɟɝɭɥɹɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ trp-ɨɩɟɪɨɧɚ ɤɢɲɟɱɧɨɣ ɩɚɥɨɱɤɢ ɜɵɛɨɪ ɦɟɠɞɭ ɚɥɶɬɟɪɧɚɬɢɜɧɵɦɢ ɜɬɨɪɢɱɧɵɦɢ ɫɬɪɭɤɬɭɪɚɦɢ ɦɊɇɄ ɨɩɪɟɞɟɥɹɟɬɫɹ ɫɤɨɪɨɫɬɶɸ ɬɪɚɧɫɥɹɰɢɢ, ɬɨ ɞɥɹ ɩɢɪɢɦɢɞɢɧɨɜɨɝɨ ɨɩɟɪɨɧɚ, ɪɟɲɚɸɳɢɣ ɮɚɤɬɨɪ ɫɤɨɪɨɫɬɶ ɬɪɚɧɫɤɪɢɩɰɢɢ (ɪɢɫ. 9). ȿɫɥɢ ɭɪɨɜɟɧɶ UTP ɜɟɥɢɤ, ɬɨ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ ɛɵɫɬɪɨ ɩɪɨɱɢɬɵɜɚɟɬ ɨɥɢɝɨɭɪɢɞɢɥɨɜɭɸ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɶ ɪɟɝɭɥɢɪɭɟɦɨɝɨ ɬɟɪɦɢɧɚɬɨɪɚ ɢ ɬɪɚɧɫɥɢɪɭɸɳɚɹ ɦɊɇɄ
ɪɢɛɨɫɨɦɚ ɧɟ ɭɫɩɟɜɚɟɬ ɩɪɢɛɥɢɡɢɬɶɫɹ ɤ ɲɩɢɥɶɤɟ ɬɟɪɦɢɧɚɬɨɪɚ. ȿɫɥɢ ɤɨɧɰɟɧɬɪɚɰɢɹ UTP ɧɢɡɤɚ, ɬ.ɟ. ɜ ɭɫɥɨɜɢɹɯ, ɤɨɝɞɚ ɷɤɫɩɪɟɫɫɢɪɨɜɚɬɶ ɨɩɟɪɨɧ ɧɟɨɛɯɨɞɢɦɨ, ɊɇɄ-ɩɨɥɢɦɟɪɚɡɚ ɡɚɦɟɞɥɹɟɬɫɹ ɩɪɢ ɫɢɧɬɟɡɟ ɨɥɢɝɨɭɪɢɞɢɥɚɬɚ. ɉɪɢ ɷɬɨɦ ɬɪɚɧɫɥɢɪɭɸɳɚɹ ɪɢɛɨɫɨɦɚ ɞɨɝɨɧɹɟɬ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɭ ɢ ɩɪɟɩɹɬɫɬɜɭɟɬ ɨɛɪɚɡɨɜɚɧɢɸ ɲɩɢɥɶɤɢ ɬɟɪɦɢɧɚɬɨɪɚ.
ɇɚɤɨɧɟɰ, ɫɚɦɵɣ ɩɪɨɫɬɨɣ ɫɩɨɫɨɛ ɫɬɚɛɢɥɢɡɢɪɨɜɚɬɶ ɨɞɧɭ ɢɡ ɚɥɶɬɟɪɧɚɬɢɜɧɵɯ ɜɬɨɪɢɱɧɵɯ ɫɬɪɭɤɬɭɪ ɊɇɄ ɢɫɩɨɥɶɡɭɟɬɫɹ ɞɥɹ ɪɟɝɭɥɹɰɢɢ ɬɪɚɧɫɤɪɢɩɰɢɢ trp-ɨɩɟɪɨɧɚ B. subtilis (ɪɢɫ.10). ȼ ɩɪɢɫɭɬɫɬɜɢɢ ɛɨɥɶɲɢɯ ɤɨɧɰɟɧɬɪɚɰɢɣ ɬɪɢɩɬɨɮɚɧɚ ɬɪɢɩɬɨɮɚɧ-ɫɜɹɡɵɜɚɸɳɢɣ ɛɟɥɨɤ TRAP ɜɡɚɢɦɨɞɟɣɫɬɜɭɟɬ ɫ ɬɪɚɧɫɤɪɢɛɢɪɭɸɳɟɣɫɹ ɦɊɇɄ trp-ɨɩɟɪɨɧɚ ɢ ɫɬɚɛɢɥɢɡɢɪɭɟɬ ɤɨɧɮɨɪɦɚɰɢɸ, ɫɩɨɫɨɛɫɬɜɭɸɳɭɸ ɩɪɟɠɞɟɜɪɟɦɟɧɧɨɣ ɬɟɪɦɢɧɚɰɢɢ.
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Ɋɢɫ. 10. Ɋɟɝɭɥɹɰɢɹ ɬɪɚɧɫɤɪɢɩɰɢɢ ɬɪɢɩɬɨɮɚɧɨɜɨɝɨ ɨɩɟɪɨɧɚ B. subtilis ɫ ɩɨɦɨɳɶɸ ɚɬɬɟɧɸɚɰɢɢ.
Ʉɪɨɦɟ ɚɬɬɟɧɸɚɰɢɢ, ɪɟɝɭɥɹɰɢɹ ɬɟɪɦɢɧɚɰɢɢ ɬɚɤɠɟ ɩɪɨɢɫɯɨɞɢɬ ɩɪɢ ɬ.ɧ. ɚɧɬɢɬɟɪɦɢɧɚɰɢɢ. Ʌɭɱɲɟ ɜɫɟɝɨ ɩɪɨɰɟɫɫ ɚɧɬɢɬɟɪɦɢɧɚɰɢɢ ɢɡɭɱɟɧ ɞɥɹ ɛɚɤɬɟɪɢɨɮɚɝɚ O (ɪɢɫ.11). ɍ ɞɚɧɧɨɝɨ ɮɚɝɚ ɢɦɟɟɬɫɹ ɞɜɚ ɦɟɯɚɧɢɡɦɚ ɚɧɬɢɬɟɪɦɢɧɚɰɢɢ. Ɉɛɚ ɦɟɯɚɧɢɡɦɚ ɫɜɹɡɚɧɵ ɫ ɪɟɝɭɥɹɬɨɪɧɵɦɢ ɛɟɥɤɚɦɢ-ɚɧɬɢɬɟɪɦɢɧɚɬɨɪɚɦɢ. Ȼɟɥɨɤ N ɮɚɝɚ O ɫɜɹɡɵɜɚɟɬɫɹ ɫ ɨɩɪɟɞɟɥɟɧɧɨɣ ɫɬɪɭɤɬɭɪɨɣ ɧɚ ɬɪɚɧɫɤɪɢɛɢɪɭɟɦɨɣ ɊɇɄ – nut (ɨɬ N utilization). ɇɚ ɞɚɧɧɨɣ ɫɬɪɭɤɬɭɪɟ ɩɪɢ ɭɱɚɫɬɢɢ N ɫɨɛɢɪɚɟɬɫɹ ɊɇɄ-ɛɟɥɤɨɜɵɣ ɤɨɦɩɥɟɤɫ, ɜɤɥɸɱɚɸɳɢɣ ɛɟɥɤɢ S10, NusA, NusB ɢ NusG, ɜɡɚɢɦɨɞɟɣɫɬɜɭɸɳɢɣ ɫ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɨɣ. ɗɬɨɬ ɤɨɦɩɥɟɤɫ ɩɪɟɩɹɬɫɬɜɭɟɬ ɬɟɪɦɢɧɚɰɢɢ ɞɚɧɧɨɝɨ ɊɇɄ-ɩɨɥɢɦɟɪɚɡɧɨɝɨ ɤɨɦɩɥɟɤɫɚ ɧɚ U-ɡɚɜɢɫɢɦɵɯ ɢ ɧɟɤɨɬɨɪɵɯ U-ɧɟɡɚɜɢɫɢɦɵɯ ɬɟɪɦɢɧɚɬɨɪɚɯ.