microbiology_konspekt_lysak_2008

ɪɢɣ ɞɥɹ ɚɫɫɢɦɢɥɹɰɢɢ ɭɝɥɟɤɢɫɥɨɝɨ ɝɚɡɚ (ɮɨɬɨɚɜɬɨɬɪɨɮɵ) ɢ ɞɪɭɝɢɯ ɫɨɟɞɢɧɟɧɢɣ (ɮɨɬɨɝɟɬɟɪɨɬɪɨɮɵ).

9.1.2. ɉɭɪɩɭɪɧɵɟ ɛɚɤɬɟɪɢɢ

ɉɭɪɩɭɪɧɵɟ ɛɚɤɬɟɪɢɢ ɨɫɭɳɟɫɬɜɥɹɸɬ ɚɧɨɤɫɢɝɟɧɧɵɣ ɮɨɬɨɫɢɧɬɟɡ ɢ ɨɬɧɨɫɹɬɫɹ ɤ ɤɥɚɫɫɭ Anoxyphotobacteria. Ɉɛɳɢɦ ɞɥɹ ɜɫɟɯ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɩɭɪɩɭɪɧɵɯ ɛɚɤɬɟɪɢɣ ɹɜɥɹɟɬɫɹ ɬɨ, ɱɬɨ ɤɨɦɩɨɧɟɧɬɵ ɮɨɬɨɫɢɧɬɟɬɢɱɟɫɤɨɝɨ ɚɩɩɚɪɚɬɚ ɧɚɯɨɞɹɬɫɹ ɜ ɬɢɥɚɤɨɢɞɚɯ. ȼ ɛɨɥɶɲɢɧɫɬɜɟ ɫɥɭɱɚɟɜ ɬɢɩɢɱɧɵɦ ɞɥɹ ɞɚɧɧɨɣ ɝɪɭɩɩɵ ɛɚɤɬɟɪɢɣ ɯɥɨɪɨɮɢɥɥɨɦ ɹɜɥɹɟɬɫɹ ɛɚɤɬɟɪɢɨɯɥɨɪɨɮɢɥɥ ɚ. ȼɫɟ ɷɬɢ ɛɚɤɬɟɪɢɢ ɫɩɨɫɨɛɧɵ ɮɢɤɫɢɪɨɜɚɬɶ CO2 ɜ ɰɢɤɥɟ Ʉɚɥɶɜɢɧɚ. Ɇɧɨɝɢɟ ɩɭɪɩɭɪɧɵɟ ɛɚɤɬɟɪɢɢ ɩɪɨɹɜɥɹɸɬ ɫɩɨɫɨɛɧɨɫɬɶ ɤ ɚɡɨɬɮɢɤɫɚɰɢɢ.

Ƚɪɭɩɩɚ ɩɭɪɩɭɪɧɵɯ ɛɚɤɬɟɪɢɣ ɜ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɧɚɫɱɢɬɵɜɚɟɬ ɛɨɥɟɟ 50 ɜɢɞɨɜ. ȼɫɟ ɩɭɪɩɭɪɧɵɟ ɛɚɤɬɟɪɢɢ – ɨɞɧɨɤɥɟɬɨɱɧɵɟ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ ɪɚɡɧɨɣ ɦɨɪɮɨɥɨɝɢɢ (ɪɢɫ. 88). Ɋɚɡɦɟɪɵ ɢɯ ɤɨɥɟɛɥɸɬɫɹ ɨɬ 1 ɞɨ 20 ɦɤɦ ɜ ɞɥɢɧɭ ɢ ɨɬ 0,3 ɞɨ 6 ɦɤɦ ɜ ɲɢɪɢɧɭ. ɋɪɟɞɢ ɩɭɪɩɭɪɧɵɯ ɛɚɤɬɟɪɢɣ ɟɫɬɶ ɧɟɩɨɞɜɢɠɧɵɟ ɢ ɩɨɞɜɢɠɧɵɟ ɮɨɪɦɵ. Ⱦɜɢɠɟɧɢɟ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɫ ɩɨɦɨɳɶɸ ɨɞɧɨɝɨ ɢɥɢ ɩɭɱɤɚ ɠɝɭɬɢɤɨɜ, ɪɚɫɩɨɥɨɠɟɧɧɵɯ ɨɛɵɱɧɨ ɩɨɥɹɪɧɨ. Ȼɨɥɶɲɢɧɫɬɜɨ ɩɭɪɩɭɪɧɵɯ ɛɚɤɬɟɪɢɣ ɪɚɡɦɧɨɠɚɟɬɫɹ ɛɢɧɚɪɧɵɦ ɞɟɥɟɧɢɟɦ, ɧɟɤɨɬɨɪɵɟ ɜɢɞɵ – ɩɨɱɤɨɜɚɧɢɟɦ. ȼɫɟ ɩɭɪɩɭɪɧɵɟ ɛɚɤɬɟɪɢɢ ɨɤɪɚɲɢɜɚɸɬɫɹ ɨɬɪɢɰɚɬɟɥɶɧɨ ɩɨ Ƚɪɚɦɭ.

Ɋɢɫ. 88. Ɉɫɧɨɜɧɵɟ ɦɨɪɮɨɥɨɝɢɱɟɫɤɢɟ ɬɢɩɵ ɩɭɪɩɭɪɧɵɯ ɛɚɤɬɟɪɢɣ:

Ⱥ – ɧɟɫɟɪɧɵɟ ɩɭɪɩɭɪɧɵɟ ɛɚɤɬɟɪɢɢ: 1 – Rhodospirillum; 2 – Rhodomicrobium; 3 – Rhodobacter sphaeroides; 4 – Rhodocyclus; 5 – Rhodopseudomonas palustris; Ȼ – ɩɭɪɩɭɪɧɵɟ ɫɟɪɨɛɚɤɬɟɪɢɢ: 6 – Ectothiorhodospira; 7 – Thiospirillum; 8 – Chromatium; 9 – Thiocystis; 10 – Thiodictyon; 11 – Thiocapsa (ɩɨ ȼ. Ɇ. Ƚɨɪɥɟɧɤɨ, Ƚ. Ⱥ. Ⱦɭɛɢɧɢɧɨɣ, ɋ. ɂ. Ʉɭɡɧɟɰɨɜɭ, 1977)

298

ɉɨ ɫɩɨɫɨɛɧɨɫɬɢ ɢɫɩɨɥɶɡɨɜɚɬɶ ɜ ɤɚɱɟɫɬɜɟ ɞɨɧɨɪɨɜ ɷɥɟɤɬɪɨɧɨɜ ɷɥɟɦɟɧɬɚɪɧɭɸ ɫɟɪɭ ɜ ɝɪɭɩɩɟ ɩɭɪɩɭɪɧɵɯ ɛɚɤɬɟɪɢɣ ɜɵɞɟɥɹɸɬ ɞɜɚ ɫɟɦɟɣɫɬɜɚ: ɩɭɪɩɭɪɧɵɟ ɫɟɪɧɵɟ (Chromatiaceae) ɢ ɩɭɪɩɭɪɧɵɟ ɧɟɫɟɪɧɵɟ (Rhodospirillaceae) ɛɚɤɬɟɪɢɢ.

ȼɫɟ ɩɪɟɞɫɬɚɜɢɬɟɥɢ ɩɭɪɩɭɪɧɵɯ ɫɟɪɧɵɯ ɛɚɤɬɟɪɢɣ ɦɨɝɭɬ ɪɚɫɬɢ ɩɪɢ ɨɫɜɟɳɟɧɢɢ ɜ ɚɧɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ ɧɚ ɦɢɧɢɦɚɥɶɧɨɣ ɫɪɟɞɟ, ɫɨɞɟɪɠɚɳɟɣ ɜ ɤɚɱɟɫɬɜɟ ɟɞɢɧɫɬɜɟɧɧɨɝɨ ɢɫɬɨɱɧɢɤɚ ɭɝɥɟɪɨɞɚ ɋɈ2, ɢɫɩɨɥɶɡɭɹ H2S ɜ ɤɚɱɟɫɬɜɟ ɞɨɧɨɪɚ ɷɥɟɤɬɪɨɧɨɜ. Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɞɥɹ ɫɟɪɧɵɯ ɛɚɤɬɟɪɢɣ ɨɫɧɨɜɧɨɣ ɫɩɨɫɨɛ ɫɭɳɟɫɬɜɨɜɚɧɢɹ – ɮɨɬɨɥɢɬɨɚɜɬɨɬɪɨɮɢɹ. Ɇɧɨɝɢɟ ɜɢɞɵ ɢɫɩɨɥɶɡɭɸɬ ɞɥɹ ɷɬɨɣ ɰɟɥɢ ɦɨɥɟɤɭɥɹɪɧɭɸ ɫɟɪɭ (S0), ɫɭɥɶɮɢɬ (SO32 ), ɬɢɨɫɭɥɶɮɚɬ

(S2O32 ), ɦɨɥɟɤɭɥɹɪɧɵɣ ɜɨɞɨɪɨɞ. ɋɭɥɶɮɢɞ ɨɤɢɫɥɹɟɬɫɹ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨ ɞɨ ɦɨɥɟɤɭɥɹɪɧɨɣ ɫɟɪɵ, ɞɚɥɟɟ ɞɨ ɫɭɥɶɮɚɬɚ (SO24 ), ɩɪɢ ɷɬɨɦ ɤɚɩɥɢ ɫɟɪɵ, ɨɤɪɭɠɟɧɧɵɟ ɛɟɥɤɨɜɨɣ ɦɟɦɛɪɚɧɨɣ, ɜɪɟɦɟɧɧɨ ɨɬɤɥɚɞɵɜɚɸɬɫɹ ɜ ɩɟɪɢɩɥɚɡɦɚɬɢɱɟɫɤɨɦ ɩɪɨɫɬɪɚɧɫɬɜɟ. ɗɬɨ ɩɪɨɢɫɯɨɞɢɬ ɜ ɪɟɡɭɥɶɬɚɬɟ ɬɨɝɨ, ɱɬɨ ɨɤɢɫɥɟɧɢɟ H2S ɞɨ S0 ɩɪɟɜɨɫɯɨɞɢɬ ɫɤɨɪɨɫɬɶ ɩɨɫɥɟɞɭɸɳɟɝɨ ɨɤɢɫɥɟɧɢɹ S0 ɞɨ

SO24 .

Ⱦɥɹ ɛɨɥɶɲɢɧɫɬɜɚ ɩɭɪɩɭɪɧɵɯ ɧɟɫɟɪɧɵɯ ɛɚɤɬɟɪɢɣ ɯɚɪɚɤɬɟɪɟɧ ɮɨɬɨɨɪɝɚɧɨɝɟɬɟɪɨɬɪɨɮɧɵɣ ɨɛɪɚɡ ɠɢɡɧɢ. Ⱦɨɧɨɪɚɦɢ ɷɥɟɤɬɪɨɧɨɜ ɢ ɢɫɬɨɱɧɢɤɚɦɢ ɭɝɥɟɪɨɞɚ ɜ ɩɪɨɰɟɫɫɟ ɮɨɬɨɫɢɧɬɟɡɚ ɹɜɥɹɸɬɫɹ ɠɢɪɧɵɟ ɤɢɫɥɨɬɵ, ɫɩɢɪɬɵ, ɭɝɥɟɜɨɞɵ, ɚɦɢɧɨɤɢɫɥɨɬɵ.

ɇɟɤɨɬɨɪɵɟ ɩɭɪɩɭɪɧɵɟ ɧɟɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ ɪɚɫɬɭɬ ɩɪɢ ɨɫɜɟɳɟɧɢɢ ɧɚ ɦɢɧɟɪɚɥɶɧɨɣ ɫɪɟɞɟ, ɢɫɩɨɥɶɡɭɹ ɜ ɤɚɱɟɫɬɜɟ ɞɨɧɨɪɚ ɷɥɟɤɬɪɨɧɨɜ H2S, S2O32 ,

ɢɥɢ S0. ȼ ɛɨɥɶɲɢɧɫɬɜɟ ɫɥɭɱɚɟɜ ɫɭɥɶɮɢɞ ɨɤɢɫɥɹɟɬɫɹ ɬɨɥɶɤɨ ɞɨ ɦɨɥɟɤɭɥɹɪɧɨɣ ɫɟɪɵ, ɧɢɤɨɝɞɚ ɧɟ ɨɬɤɥɚɞɵɜɚɸɳɟɣɫɹ ɜ ɤɥɟɬɤɟ, ɧɨ ɜ ɨɬɞɟɥɶɧɵɯ ɫɥɭɱɚɹɯ ɜɨɡɦɨɠɧɨ ɩɨɫɥɟɞɭɸɳɟɟ ɨɤɢɫɥɟɧɢɟ S0 ɞɨ ɫɭɥɶɮɚɬɚ.

ȼ ɝɪɭɩɩɟ ɩɭɪɩɭɪɧɵɯ ɧɟɫɟɪɧɵɯ ɛɚɤɬɟɪɢɣ ɨɛɧɚɪɭɠɟɧɨ ɛɨɥɶɲɨɟ ɪɚɡɧɨɨɛɪɚɡɢɟ ɦɟɬɚɛɨɥɢɱɟɫɤɢɯ ɩɭɬɟɣ, ɫɜɹɡɚɧɧɵɯ ɫ ɩɨɥɭɱɟɧɢɟɦ ɷɧɟɪɝɢɢ. Ɇɧɨɝɢɟ ɩɪɟɞɫɬɚɜɢɬɟɥɢ ɷɬɨɣ ɝɪɭɩɩɵ ɫɩɨɫɨɛɧɵ ɪɚɫɬɢ ɜ ɬɟɦɧɨɬɟ ɜ ɦɢɤɪɨɚɷɪɨɮɢɥɶɧɵɯ ɢɥɢ ɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ, ɩɨɥɭɱɚɹ ɷɧɟɪɝɢɸ ɜ ɩɪɨɰɟɫɫɟ ɞɵɯɚɧɢɹ. ɍ ɧɢɯ ɚɤɬɢɜɧɨ ɮɭɧɤɰɢɨɧɢɪɭɟɬ ɡɚɦɤɧɭɬɵɣ ɰɢɤɥ Ʉɪɟɛɫɚ, ɝɥɢɤɨɥɢɬɢɱɟɫɤɢɣ ɩɭɬɶ ɢ ɞɪɭɝɢɟ ɩɭɬɢ ɤɚɬɚɛɨɥɢɡɦɚ ɨɪɝɚɧɢɱɟɫɤɢɯ ɫɨɟɞɢɧɟɧɢɣ. ɉɪɟɞɫɬɚɜɢɬɟɥɢ ɪɨɞɚ Rhodobacter ɫɩɨɫɨɛɧɵ ɤ ɯɟɦɨɚɜɬɨɬɪɨɮɢɢ. Ɉɧɢ ɪɚɫɬɭɬ ɧɚ ɦɢɧɟɪɚɥɶɧɨɣ ɫɪɟɞɟ ɜ ɬɟɦɧɨɬɟ ɩɪɢ ɩɨɧɢɠɟɧɧɨɣ ɤɨɧɰɟɧɬɪɚɰɢɢ Ɉ2, ɢɫɩɨɥɶɡɭɹ ɷɧɟɪɝɢɸ, ɩɨɥɭɱɚɟɦɭɸ ɩɪɢ ɨɤɢɫɥɟɧɢɢ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɜɨɞɨɪɨɞɚ, ɞɥɹ ɚɫɫɢɦɢɥɹɰɢɢ ɋɈ2. ɍ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɪɨɞɚ Rhodobacter ɨɛɧɚɪɭɠɟɧɚ ɬɚɤɠɟ ɫɩɨɫɨɛɧɨɫɬɶ ɤ ɪɨɫɬɭ ɜ ɚɧɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ ɡɚ ɫɱɟɬ ɨɤɢɫɥɟɧɢɹ ɨɪɝɚɧɢɱɟɫɤɢɯ ɫɨɟɞɢɧɟɧɢɣ ɜ ɩɪɨɰɟɫɫɟ ɧɢɬɪɚɬɧɨɝɨ ɞɵɯɚɧɢɹ. Ʉɪɨɦɟ ɬɨɝɨ, ɞɥɹ ɪɹɞɚ ɩɭɪɩɭɪɧɵɯ ɧɟɫɟɪɧɵɯ ɛɚɤɬɟɪɢɣ ɩɨɤɚɡɚɧɚ ɫɩɨɫɨɛɧɨɫɬɶ ɪɚɫɬɢ ɚɧɚɷɪɨɛɧɨ ɜ

299

ɬɟɦɧɨɬɟ, ɨɫɭɳɟɫɬɜɥɹɹ ɫɛɪɚɠɢɜɚɧɢɟ ɨɪɝɚɧɢɱɟɫɤɢɯ ɫɭɛɫɬɪɚɬɨɜ (ɭɝɥɟɜɨɞɵ, ɩɢɪɭɜɚɬ ɢ ɞɪ.).

Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɦɨɠɧɨ ɡɚɤɥɸɱɢɬɶ, ɱɬɨ ɞɥɹ ɩɭɪɩɭɪɧɵɯ ɧɟɫɟɪɧɵɯ ɛɚɤɬɟɪɢɣ ɢɫɬɨɱɧɢɤɚɦɢ ɷɧɟɪɝɢɢ ɦɨɝɭɬ ɛɵɬɶ ɮɨɬɨɫɢɧɬɟɡ, ɚɷɪɨɛɧɨɟ ɞɵɯɚɧɢɟ, ɚɧɚɷɪɨɛɧɨɟ ɞɵɯɚɧɢɟ ɢ ɛɪɨɠɟɧɢɟ.

9.1.3. Ɂɟɥɟɧɵɟ ɛɚɤɬɟɪɢɢ

Ɂɟɥɟɧɵɟ ɛɚɤɬɟɪɢɢ ɨɫɭɳɟɫɬɜɥɹɸɬ ɚɧɨɤɫɢɝɟɧɧɵɣ ɮɨɬɨɫɢɧɬɟɡ ɢ ɨɬɧɨɫɹɬɫɹ ɤ ɤɥɚɫɫɭ Anoxyphotobacteria. ȼ ɝɪɭɩɩɟ ɡɟɥɟɧɵɯ ɛɚɤɬɟɪɢɣ ɜɵɞɟɥɹɸɬ ɡɟɥɟɧɵɟ ɫɟɪɧɵɟ (Chlorobiaceae) ɢ ɡɟɥɟɧɵɟ ɧɢɬɱɚɬɵɟ (Chloroflexaceae).

ȼɫɟ ɡɟɥɟɧɵɟ ɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ – ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɨɞɧɨɤɥɟɬɨɱɧɵɟ ɧɟɩɨɞɜɢɠɧɵɟ ɦɢɤɪɨɨɪɝɚɧɢɡɦɵ (ɪɢɫ. 89). Ʉɥɟɬɤɢ ɦɟɥɤɢɟ (0,3–1,2 ɯ 0,5– 2,7 ɦɤɦ), ɩɚɥɨɱɤɨɜɢɞɧɵɟ, ɹɣɰɟɨɛɪɚɡɧɵɟ ɢɥɢ ɫɥɟɝɤɚ ɢɡɨɝɧɭɬɵɟ. Ɋɚɡɦɧɨɠɟɧɢɟ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɛɢɧɚɪɧɵɦ ɞɟɥɟɧɢɟɦ. ȼ ɤɚɱɟɫɬɜɟ ɡɚɩɚɫɧɨɝɨ ɜɟɳɟɫɬɜɚ ɡɟɥɟɧɵɟ ɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ ɧɚɤɚɩɥɢɜɚɸɬ ɝɥɢɤɨɝɟɧɨɩɨɞɨɛɧɵɣ ɩɨɥɢɫɚɯɚɪɢɞ. ɉɨɥɢ-ȕ-ɝɢɞɪɨɤɫɢɦɚɫɥɹɧɚɹ ɤɢɫɥɨɬɚ ɧɟ ɨɛɧɚɪɭɠɟɧɚ.

Ɋɢɫ. 89. Ɉɫɧɨɜɧɵɟ ɦɨɪɮɨɥɨɝɢɱɟɫɤɢɟ ɮɨɪɦɵ ɡɟɥɟɧɵɯ ɫɟɪɨɛɚɤɬɟɪɢɣ:

1 – Chlorobium limicola; 2 – Chlorobium vibrioforme; 3 – Prosthecochloris aestuarii; 4 – Pelodictyon lutecium; 5 – Pelodictyon clathratiforme; 6 – Clathrochloris sulfurica;

7 – Ancalochloris perfilievii (ɩɨ ȼ. Ɇ. Ƚɨɪɥɟɧɤɨ, Ƚ. Ⱥ. Ⱦɭɛɢɧɢɧɨɣ, ɋ. ɂ. Ʉɭɡɧɟɰɨɜɭ, 1977)

Ɂɟɥɟɧɵɟ ɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ ɹɜɥɹɸɬɫɹ ɫɬɪɨɝɨ ɚɧɚɷɪɨɛɧɵɦɢ ɮɨɬɨɥɢɬɨɚɜɬɨɬɪɨɮɚɦɢ. ȼ ɤɚɱɟɫɬɜɟ ɞɨɧɨɪɨɜ ɷɥɟɤɬɪɨɧɨɜ ɨɧɢ ɢɫɩɨɥɶɡɭɸɬ H2S ɢɥɢ ɞɪɭɝɢɟ ɜɨɫɫɬɚɧɨɜɥɟɧɧɵɟ ɧɟɨɪɝɚɧɢɱɟɫɤɢɟ ɫɨɟɞɢɧɟɧɢɹ ɫɟɪɵ, ɚ ɬɚɤɠɟ ɦɨɥɟɤɭɥɹɪɧɵɣ ɜɨɞɨɪɨɞ. ɉɨɥɭɱɚɸɳɚɹɫɹ ɩɪɢ ɨɤɢɫɥɟɧɢɢ H2S ɷɥɟɦɟɧɬɚɪɧɚɹ ɫɟɪɚ ɞɨ ɨɤɢɫɥɟɧɢɹ ɟɟ ɞɨ ɫɭɥɶɮɚɬɚ ɨɬɤɥɚɞɵɜɚɟɬɫɹ ɜɧɟ ɤɥɟɬɤɢ.

Ɇɧɨɝɢɟ ɢɡ ɧɢɯ ɧɭɠɞɚɸɬɫɹ ɜ ɜɢɬɚɦɢɧɟ ȼ12. Ʉɚɤ ɩɪɚɜɢɥɨ, ɡɟɥɟɧɵɟ ɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ ɫɩɨɫɨɛɧɵ ɮɢɤɫɢɪɨɜɚɬɶ ɦɨɥɟɤɭɥɹɪɧɵɣ ɚɡɨɬ.

ɉɨ ɜɫɟɦ ɜɵɲɟɩɟɪɟɱɢɫɥɟɧɧɵɦ ɫɜɨɣɫɬɜɚɦ ɨɧɢ ɩɨɯɨɠɢ ɧɚ ɩɭɪɩɭɪɧɵɟ ɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ. Ⱦɟɣɫɬɜɢɬɟɥɶɧɨ, ɩɭɪɩɭɪɧɵɟ ɢ ɡɟɥɟɧɵɟ ɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ

300

ɨɛɵɱɧɨ ɫɨɫɭɳɟɫɬɜɭɸɬ ɜ ɨɫɜɟɳɟɧɧɨɣ, ɛɨɝɚɬɨɣ ɫɭɥɶɮɢɞɚɦɢ ɚɧɚɷɪɨɛɧɨɣ ɜɨɞɧɨɣ ɫɪɟɞɟ ɢ ɨɛɥɚɫɬɢ ɢɯ ɪɚɫɩɪɨɫɬɪɚɧɟɧɢɹ ɜ ɡɧɚɱɢɬɟɥɶɧɨɣ ɫɬɟɩɟɧɢ ɩɟɪɟɤɪɵɜɚɸɬɫɹ. Ɉɞɧɚɤɨ ɷɬɢ ɞɜɟ ɝɪɭɩɩɵ ɡɚɦɟɬɧɨ ɪɚɡɥɢɱɚɸɬɫɹ ɩɨ ɢɫɩɨɥɶɡɭɟɦɵɦ ɢɦɢ ɢɫɬɨɱɧɢɤɚɦ ɭɝɥɟɪɨɞɚ. ɇɢ ɨɞɢɧ ɢɡ ɜɢɞɨɜ ɡɟɥɟɧɵɯ ɫɟɪɧɵɯ ɛɚɤɬɟɪɢɣ ɧɟ ɦɨɠɟɬ ɪɚɫɬɢ ɮɨɬɨɝɟɬɟɪɨɬɪɨɮɧɨ, ɢɦɟɹ ɜ ɤɚɱɟɫɬɜɟ ɟɞɢɧɫɬɜɟɧɧɨɝɨ ɢ ɨɫɧɨɜɧɨɝɨ ɢɫɬɨɱɧɢɤɚ ɭɝɥɟɪɨɞɚ ɨɪɝɚɧɢɱɟɫɤɢɟ ɫɨɟɞɢɧɟɧɢɹ, ɟɫɥɢ ɜ ɫɪɟɞɟ ɨɬɫɭɬɫɬɜɭɸɬ ɧɟɨɪɝɚɧɢɱɟɫɤɢɟ ɜɨɫɫɬɚɧɨɜɢɬɟɥɢ (ɩɭɪɩɭɪɧɵɟ ɫɟɪɧɵɟ ɦɨɝɭɬ ɢɫɩɨɥɶɡɨɜɚɬɶ ɢ ɨɪɝɚɧɢɱɟɫɤɢɟ ɢɫɬɨɱɧɢɤɢ ɭɝɥɟɪɨɞɚ). Ɂɟɥɟɧɵɟ ɫɟɪɧɵɟ ɛɚɤɬɟɪɢɢ ɧɟ ɢɦɟɸɬ ɪɢɛɭɥɨɡɨ-1,5-ɞɢɮɨɫɮɚɬɤɚɪɛɨɤɫɢɥɚɡɵ, ɩɨɷɬɨɦɭ ɨɧɢ ɧɟ ɦɨɝɭɬ ɮɢɤɫɢɪɨɜɚɬɶ ɋɈ2 ɜ ɰɢɤɥɟ Ʉɚɥɶɜɢɧɚ. Ɉɧɢ ɮɢɤɫɢɪɭɸɬ ɋɈ2 ɜ ɰɢɤɥɟ Ⱥɪɧɨɧɚ, ɜ ɤɨɬɨɪɨɦ ɋɈ2 ɮɢɤɫɢɪɭɟɬɫɹ ɧɚ ɨɪɝɚɧɢɱɟɫɤɢɯ ɤɢɫɥɨɬɚɯ ɫ ɨɛɪɚɡɨɜɚɧɢɟɦ ɤɨɧɟɱɧɨɝɨ ɩɪɨɞɭɤɬɚ – ɳɚɜɟɥɟɜɨɭɤɫɭɫɧɨɣ ɤɢɫɥɨɬɵ (ɞɥɹ ɫɪɚɜɧɟɧɢɹ, ɤɨɧɟɱɧɵɣ ɩɪɨɞɭɤɬ ɰɢɤɥɚ Ʉɚɥɶɜɢɧɚ – ɝɥɸɤɨɡɚ, ɝɥɢɤɨɝɟɧ ɢ ɞɪɭɝɢɟ ɭɝɥɟɜɨɞɵ).

Ɂɟɥɟɧɵɟ ɧɢɬɱɚɬɵɟ ɛɚɤɬɟɪɢɢ ɩɪɟɞɫɬɚɜɥɹɸɬ ɫɨɛɨɣ ɩɟɪɟɞɜɢɝɚɸɳɢɟɫɹ ɩɭɬɟɦ ɫɤɨɥɶɠɟɧɢɹ ɨɪɝɚɧɢɡɦɵ, ɫɨɫɬɨɹɳɢɟ ɢɡ ɦɧɨɠɟɫɬɜɚ ɩɚɥɨɱɤɨɜɢɞɧɵɯ ɤɥɟɬɨɤ, ɤɨɬɨɪɵɟ ɧɚɡɵɜɚɸɬɫɹ ɬɪɢɯɨɦɚɦɢ ɢ ɞɨɫɬɢɝɚɸɬ ɜ ɞɥɢɧɭ 100– 300 ɦɤɦ (ɪɢɫ. 90). ɍ ɧɟɤɨɬɨɪɵɯ ɜɢɞɨɜ ɬɪɢɯɨɦɵ ɨɤɪɭɠɟɧɵ ɫɥɢɡɢɫɬɵɦ ɱɟɯɥɨɦ. ȼɫɟ ɨɩɢɫɚɧɧɵɟ ɩɪɟɞɫɬɚɜɢɬɟɥɢ ɫɟɦɟɣɫɬɜɚ Chloroflexaceae ɢɦɟɸɬ ɬɢɩɢɱɧɭɸ ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɭɸ ɤɥɟɬɨɱɧɭɸ ɫɬɟɧɤɭ, ɧɨ ɧɟ ɪɢɝɢɞɧɭɸ, ɚ ɝɢɛɤɭɸ, ɨɛɟɫɩɟɱɢɜɚɸɳɭɸ ɫɤɨɥɶɡɹɳɟɟ ɞɜɢɠɟɧɢɟ ɫɨ ɫɤɨɪɨɫɬɶɸ 0,1–0,4 ɦɤɦ/ɫ. Ʉɥɟɬɤɢ ɪɚɡɦɧɨɠɚɸɬɫɹ ɩɨɩɟɪɟɱɧɵɦ ɛɢɧɚɪɧɵɦ ɞɟɥɟɧɢɟɦ. Ʉɪɨɦɟ ɬɨɝɨ, ɤɚɤ ɢ ɜɫɟ ɧɢɬɱɚɬɵɟ ɮɨɪɦɵ, ɡɟɥɟɧɵɟ ɫɤɨɥɶɡɹɳɢɟ ɛɚɤɬɟɪɢɢ ɪɚɡɦɧɨɠɚɸɬɫɹ ɩɭɬɟɦ ɨɬɞɟɥɟɧɢɹ ɧɟɛɨɥɶɲɨɣ ɱɚɫɬɢ ɬɪɢɯɨɦɚ.

Ɋɢɫ. 90. Ɉɫɧɨɜɧɵɟ ɦɨɪɮɨɥɨɝɢɱɟɫɤɢɟ ɮɨɪɦɵ ɡɟɥɟɧɵɯ ɫɤɨɥɶɡɹɳɢɯ ɛɚɤɬɟɪɢɣ:

1 – Chloroflexus aurantiacus; 2 – Chloronema giganteum; 3 – Oscillochloris chrysea

(ɩɨ ȼ. Ɇ. Ƚɨɪɥɟɧɤɨ, Ƚ. Ⱥ. Ⱦɭɛɢɧɢɧɨɣ, ɋ. ɂ. Ʉɭɡɧɟɰɨɜɭ, 1977)

Ɏɢɡɢɨɥɨɝɨ-ɛɢɨɯɢɦɢɱɟɫɤɚɹ ɯɚɪɚɤɬɟɪɢɫɬɢɤɚ ɡɟɥɟɧɵɯ ɧɢɬɱɚɬɵɯ ɛɚɤɬɟɪɢɣ ɨɫɧɨɜɚɧɚ ɧɚ ɞɚɧɧɵɯ, ɩɨɥɭɱɟɧɧɵɯ ɞɥɹ ɨɞɧɨɝɨ ɢɡ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɷɬɨɣ ɝɪɭɩɩɵ – Chloroflexus aurantiacus, ɩɨɫɤɨɥɶɤɭ ɨɫɬɚɥɶɧɵɟ ɧɟ ɩɨɥɭɱɟɧɵ ɤ ɧɚɫɬɨɹɳɟɦɭ ɜɪɟɦɟɧɢ ɜ ɱɢɫɬɨɣ ɤɭɥɶɬɭɪɟ. ɇɢɬɱɚɬɵɟ ɡɟɥɟɧɵɟ ɛɚɤɬɟɪɢɢ

301

C. aurantiacus ɹɜɥɹɸɬɫɹ ɮɚɤɭɥɶɬɚɬɢɜɧɵɦɢ ɚɧɚɷɪɨɛɚɦɢ ɢ ɮɨɬɨɬɪɨɮɚɦɢ. ɇɚ ɫɜɟɬɭ ɨɧɢ ɪɚɫɬɭɬ ɜ ɚɷɪɨɛɧɵɯ ɢ ɚɧɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ ɜ ɩɪɢɫɭɬɫɬɜɢɢ ɪɚɡɧɨɨɛɪɚɡɧɵɯ ɨɪɝɚɧɢɱɟɫɤɢɯ ɫɨɟɞɢɧɟɧɢɣ: ɭɝɥɟɜɨɞɨɜ, ɫɩɢɪɬɨɜ, ɨɪɝɚɧɢɱɟɫɤɢɯ ɤɢɫɥɨɬ ɢ ɚɦɢɧɨɤɢɫɥɨɬ. ȼ ɬɟɦɧɨɬɟ ɪɨɫɬ ɜɨɡɦɨɠɟɧ ɬɨɥɶɤɨ ɜ ɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ. Ɍɚɤɢɦ ɨɛɪɚɡɨɦ, ɨɪɝɚɧɢɱɟɫɤɢɟ ɫɨɟɞɢɧɟɧɢɹ ɢɫɩɨɥɶɡɭɸɬɫɹ ɷɬɢɦ ɨɪɝɚɧɢɡɦɨɦ ɧɟɫɤɨɥɶɤɢɦɢ ɩɭɬɹɦɢ: ɜ ɤɚɱɟɫɬɜɟ ɢɫɬɨɱɧɢɤɨɜ ɭɝɥɟɪɨɞɚ, ɢɫɬɨɱɧɢɤɨɜ ɷɧɟɪɝɢɢ ɢ ɞɨɧɨɪɨɜ ɷɥɟɤɬɪɨɧɨɜ.

Ɂɟɥɟɧɵɟ ɧɢɬɱɚɬɵɟ ɛɚɤɬɟɪɢɢ ɩɪɚɤɬɢɱɟɫɤɢ ɧɟ ɫɩɨɫɨɛɧɵ ɤ ɮɨɬɨɬɪɨɮɧɨɦɭ ɪɨɫɬɭ ɧɚ ɋɈ2 ɢ ɇ2 ɢ, ɫɥɟɞɨɜɚɬɟɥɶɧɨ, ɷɬɢ ɛɚɤɬɟɪɢɢ ɹɜɥɹɸɬɫɹ, ɫɤɨɪɟɟ, ɮɨɬɨɝɟɬɟɪɨɬɪɨɮɚɦɢ. Ɏɨɬɨɚɫɫɢɦɢɥɹɰɢɹ ɋɈ2 ɭ Chloroflexus aurantiacus ɩɪɨɢɫɯɨɞɢɬ ɜ ɰɢɤɥɟ Ʉɚɥɶɜɢɧɚ, ɧɚ ɱɬɨ ɭɤɚɡɵɜɚɟɬ ɨɛɧɚɪɭɠɟɧɧɵɣ ɭ ɧɢɯ ɚɤɬɢɜɧɵɣ ɮɟɪɦɟɧɬ ɪɢɛɭɥɨɡɨ-1,5-ɞɢɮɨɫɮɚɬɤɚɪɛɨɤɫɢɥɚɡɚ. Ɇɟɬɚɛɨɥɢɡɦ ɨɪɝɚɧɢɱɟɫɤɢɯ ɭɝɥɟɪɨɞɧɵɯ ɫɨɟɞɢɧɟɧɢɣ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɜ ɪɟɡɭɥɶɬɚɬɟ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ɩɨɥɧɨɝɨ ɰɢɤɥɚ ɬɪɢɤɚɪɛɨɧɨɜɵɯ ɤɢɫɥɨɬ ɢ ɝɥɢɨɤɫɢɥɚɬɧɨɝɨ ɰɢɤɥɚ. ɉɨɩɵɬɤɢ ɨɛɧɚɪɭɠɢɬɶ ɭ ɞɚɧɧɵɯ ɛɚɤɬɟɪɢɣ ɫɩɨɫɨɛɧɨɫɬɶ ɤ ɮɢɤɫɚɰɢɢ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɚɡɨɬɚ ɩɨɤɚ ɧɟ ɞɚɥɢ ɩɨɥɨɠɢɬɟɥɶɧɵɯ ɪɟɡɭɥɶɬɚɬɨɜ.

9.1.4. Ƚɟɥɢɨɛɚɤɬɟɪɢɢ

Ƚɟɥɢɨɛɚɤɬɟɪɢɢ ɨɫɭɳɟɫɬɜɥɹɸɬ ɚɧɨɤɫɢɝɟɧɧɵɣ ɮɨɬɨɫɢɧɬɟɡ ɛɥɚɝɨɞɚɪɹ ɧɚɥɢɱɢɸ ɜ ɤɥɟɬɤɚɯ ɟɞɢɧɫɬɜɟɧɧɨɝɨ ɛɚɤɬɟɪɢɨɯɥɨɪɨɮɢɥɥɚ g, ɤɨɬɨɪɵɣ ɧɟ ɨɛɧɚɪɭɠɟɧ ɭ ɞɪɭɝɢɯ ɛɚɤɬɟɪɢɣ ɫ ɛɟɫɤɢɫɥɨɪɨɞɧɵɦ ɬɢɩɨɦ ɮɨɬɨɫɢɧɬɟɡɚ.

ȼ ɤɥɟɬɤɚɯ ɝɟɥɢɨɛɚɤɬɟɪɢɣ, ɤɪɨɦɟ ɛɚɤɬɟɪɢɨɯɥɨɪɨɮɢɥɥɚ g, ɢɦɟɟɬɫɹ ɧɟɛɨɥɶɲɨɟ ɤɨɥɢɱɟɫɬɜɨ ɤɚɪɨɬɢɧɨɢɞɨɜ.

Ƚɟɥɢɨɛɚɤɬɟɪɢɢ ɹɜɥɹɸɬɫɹ ɟɞɢɧɫɬɜɟɧɧɵɦɢ ɝɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɦɢ ɮɨɬɨɬɪɨɮɚɦɢ, ɫɩɨɫɨɛɧɵɦɢ ɤ ɨɛɪɚɡɨɜɚɧɢɸ ɧɚɫɬɨɹɳɢɯ ɷɧɞɨɫɩɨɪ. Ɉɩɢɫɚɧɵ ɞɜɚ ɜɢɞɚ, ɪɚɡɥɢɱɚɸɳɢɟɫɹ ɦɨɪɮɨɥɨɝɢɱɟɫɤɢ: Heliobacterium chlorum – ɨɞɢɧɨɱɧɵɟ ɞɥɢɧɧɵɟ ɩɚɥɨɱɤɢ (1,0 ɯ 7,0–10 ɦɤɦ), ɫɩɨɫɨɛɧɵɟ ɩɟɪɟɞɜɢɝɚɬɶɫɹ ɫɤɨɥɶɠɟɧɢɟɦ, ɢ Heliobacillus mobilis – ɤɨɪɨɬɤɢɟ ɩɚɥɨɱɤɨɜɢɞɧɵɟ ɮɨɪɦɵ ɫ ɩɟɪɢɬɪɢɯɢɚɥɶɧɨ ɪɚɫɩɨɥɨɠɟɧɧɵɦɢ ɠɝɭɬɢɤɚɦɢ.

Ƚɟɥɢɨɛɚɤɬɟɪɢɢ – ɨɛɥɢɝɚɬɧɵɟ ɮɨɬɨɬɪɨɮɵ. Ɋɨɫɬ ɜɨɡɦɨɠɟɧ ɬɨɥɶɤɨ ɧɚ ɫɜɟɬɭ ɜ ɚɧɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ. ɂɫɬɨɱɧɢɤɚɦɢ ɭɝɥɟɪɨɞɚ ɦɨɝɭɬ ɫɥɭɠɢɬɶ ɧɟɤɨɬɨɪɵɟ ɨɪɝɚɧɢɱɟɫɤɢɟ ɤɢɫɥɨɬɵ (ɭɤɫɭɫɧɚɹ, ɦɨɥɨɱɧɚɹ, ɦɚɫɥɹɧɚɹ, ɩɢɪɨɜɢɧɨɝɪɚɞɧɚɹ). Ɏɢɤɫɚɰɢɹ ɋɈ2 ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɜ ɰɢɤɥɟ Ʉɚɥɶɜɢɧɚ. Ⱦɵɯɚɬɟɥɶɧɵɣ ɦɟɬɚɛɨɥɢɡɦ ɨɬɫɭɬɫɬɜɭɟɬ. Ƚɟɥɢɨɛɚɤɬɟɪɢɢ – ɚɤɬɢɜɧɵɟ ɚɡɨɬɮɢɤɫɚɬɨɪɵ. Ɉɧɢ ɨɛɢɬɚɸɬ ɜ ɩɨɱɜɚɯ ɢ ɫɨɞɨɜɵɯ ɨɡɟɪɚɯ.

9.1.5. ɐɢɚɧɨɛɚɤɬɟɪɢɢ

302

ɐɢɚɧɨɛɚɤɬɟɪɢɢ ɟɳɟ ɧɟɞɚɜɧɨ ɪɚɫɫɦɚɬɪɢɜɚɥɢ ɤɚɤ ɫɢɧɟɡɟɥɟɧɵɟ ɜɨɞɨɪɨɫɥɢ – ɨɫɨɛɵɣ ɬɢɩ ɧɢɡɲɢɯ ɪɚɫɬɟɧɢɣ. ɋɯɨɞɫɬɜɨ ɫɢɧɟɡɟɥɟɧɵɯ ɜɨɞɨɪɨɫɥɟɣ ɫ ɛɚɤɬɟɪɢɹɦɢ ɛɵɥɨ ɨɬɦɟɱɟɧɨ ɞɚɜɧɨ. ȼ 1875 ɝ. Ɏ. Ʉɨɧ ɨɛɴɟɞɢɧɢɥ ɢɯ ɫ ɛɚɤɬɟɪɢɹɦɢ ɜ ɨɬɞɟɥ Schyzophyta – ɪɚɫɬɟɧɢɹ-ɞɪɨɛɹɧɤɢ, ɬ. ɟ. ɪɚɫɬɟɧɢɹ, ɪɚɡɦɧɨɠɚɸɳɢɟɫɹ ɛɢɧɚɪɧɵɦ ɞɟɥɟɧɢɟɦ.

ȼ 1978 ɝ. ɇ. Ƚɢɛɛɨɧɫ ɢ Ɋ. Ɇɟɪɪɟɣ, ɨɫɧɨɜɵɜɚɹɫɶ ɧɚ ɪɹɞɟ ɫɜɨɣɫɬɜ, ɨɬɧɟɫɥɢ ɫɢɧɟɡɟɥɟɧɵɟ ɜɨɞɨɪɨɫɥɢ ɤ ɩɪɨɤɚɪɢɨɬɚɦ, ɚ Ɋ. ɋɬɟɣɧɢɟɪ ɩɪɟɞɥɨɠɢɥ ɨɬɤɚɡɚɬɶɫɹ ɨɬ ɧɚɡɜɚɧɢɹ «ɫɢɧɟɡɟɥɟɧɵɟ ɜɨɞɨɪɨɫɥɢ» ɢ ɜɜɟɫɬɢ ɬɟɪɦɢɧ «ɰɢɚɧɨɛɚɤɬɟɪɢɢ», ɱɬɨ ɨɛɭɫɥɨɜɥɟɧɨ ɪɹɞɨɦ ɫɩɟɰɢɮɢɱɟɫɤɢɯ ɫɜɨɣɫɬɜ:

x ɧɚɥɢɱɢɟɦ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ, ɫɨɫɬɨɹɳɟɣ ɢɡ ɩɟɩɬɢɞɨɝɥɢɤɚɧɚ ɦɭɪɟɢ-

ɧɚ;

xɨɬɫɭɬɫɬɜɢɟɦ ɦɟɦɛɪɚɧɵ, ɨɬɞɟɥɹɸɳɟɣ ɧɚɫɥɟɞɫɬɜɟɧɧɵɣ ɦɚɬɟɪɢɚɥ ɨɬ ɰɢɬɨɩɥɚɡɦɵ;

xɧɚɥɢɱɢɟɦ ɪɢɛɨɫɨɦ 70S-ɬɢɩɚ;

xɫɩɨɫɨɛɧɨɫɬɶɸ ɪɚɡɜɢɜɚɬɶɫɹ ɩɪɢ ɬɟɦɩɟɪɚɬɭɪɟ ɜɵɲɟ 55 ºɋ;

xɫɩɨɫɨɛɧɨɫɬɶɸ ɮɢɤɫɢɪɨɜɚɬɶ ɦɨɥɟɤɭɥɹɪɧɵɣ ɚɡɨɬ;

xɧɚɥɢɱɢɟɦ ɜ ɪɹɞɟ ɫɥɭɱɚɟɜ ɚɷɪɨɫɨɦ ɢ ɤɚɪɛɨɤɫɢɫɨɦ – ɰɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɢɯ ɜɤɥɸɱɟɧɢɣ, ɜɫɬɪɟɱɚɸɳɢɯɫɹ ɬɨɥɶɤɨ ɭ ɩɪɨɤɚɪɢɨɬ;

xɫɯɨɞɫɬɜɨɦ ɫ ɞɪɭɝɢɦɢ ɷɭɛɚɤɬɟɪɢɹɦɢ ɩɨ ɫɩɟɤɬɪɚɦ ɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɢ

ɤɚɧɬɢɛɢɨɬɢɤɚɦ.

ɐɢɚɧɨɛɚɤɬɟɪɢɢ – ɦɨɪɮɨɥɨɝɢɱɟɫɤɢ ɪɚɡɧɨɨɛɪɚɡɧɚɹ ɝɪɭɩɩɚ ɝɪɚɦɨɬɢɰɚɬɟɥɶɧɵɯ ɩɪɨɤɚɪɢɨɬ, ɜɤɥɸɱɚɸɳɚɹ ɨɞɧɨɤɥɟɬɨɱɧɵɟ, ɤɨɥɨɧɢɚɥɶɧɵɟ ɢ ɦɧɨɝɨɤɥɟɬɨɱɧɵɟ ɮɨɪɦɵ. Ʉɥɟɬɤɢ ɦɨɝɭɬ ɛɵɬɶ ɫɮɟɪɢɱɟɫɤɢɦɢ, ɩɚɥɨɱɤɨɜɢɞɧɵɦɢ ɢɥɢ ɢɡɨɝɧɭɬɵɦɢ, ɨɞɢɧɨɱɧɵɦɢ ɢɥɢ ɨɛɪɚɡɭɸɳɢɦɢ ɫɤɨɩɥɟɧɢɹ, ɭɞɟɪɠɢɜɚɟɦɵɟ ɫɨɜɦɟɫɬɧɨ ɩɨɫɥɟ ɪɹɞɚ ɞɟɥɟɧɢɣ ɫ ɩɨɦɨɳɶɸ ɨɤɪɭɠɚɸɳɟɝɨ ɢɯ ɨɛɳɟɝɨ ɱɟɯɥɚ. Ɇɧɨɝɨɤɥɟɬɨɱɧɵɟ ɮɨɪɦɵ ɢɦɟɸɬ ɧɢɬɟɜɢɞɧɨɟ ɫɬɪɨɟɧɢɟ. ɇɢɬɢ (ɬɪɢɯɨɦɵ ɢɥɢ ɮɢɥɚɦɟɧɬɵ) ɛɵɜɚɸɬ ɩɪɨɫɬɵɟ ɢɥɢ ɜɟɬɜɹɳɢɟɫɹ. ɉɪɨɫɬɵɟ ɧɢɬɢ ɫɨɫɬɨɹɬ ɢɡ ɨɞɧɨɝɨ ɪɹɞɚ ɤɥɟɬɨɤ, ɢɦɟɸɳɢɯ ɨɞɢɧɚɤɨɜɵɟ ɪɚɡɦɟɪɵ, ɮɨɪɦɭ ɢ ɫɬɪɨɟɧɢɟ, ɢɥɢ ɤɥɟɬɨɤ, ɪɚɡɥɢɱɚɸɳɢɯɫɹ ɩɨ ɷɬɢɦ ɩɚɪɚɦɟɬɪɚɦ. ȼɟɬɜɹɳɢɟɫɹ ɬɪɢɯɨɦɵ ɜɨɡɧɢɤɚɸɬ ɜ ɪɟɡɭɥɶɬɚɬɟ ɧɟɫɤɨɥɶɤɢɯ ɦɟɯɚɧɢɡɦɨɜ, ɜ ɫɜɹɡɢ ɫ ɱɟɦ ɪɚɡɥɢɱɚɸɬ ɥɨɠɧɨɟ ɢ ɢɫɬɢɧɧɨɟ ɜɟɬɜɥɟɧɢɟ. Ʉ ɢɫɬɢɧɧɨɦɭ ɜɟɬɜɥɟɧɢɸ ɩɪɢɜɨɞɢɬ ɫɩɨɫɨɛɧɨɫɬɶ ɤɥɟɬɨɤ ɬɪɢɯɨɦɚ ɞɟɥɢɬɶɫɹ ɜ ɪɚɡɧɵɯ ɩɥɨɫɤɨɫɬɹɯ, ɜ ɪɟɡɭɥɶɬɚɬɟ ɜɨɡɧɢɤɚɸɬ ɦɧɨɝɨɪɹɞɧɵɟ ɬɪɢɯɨɦɵ ɢɥɢ ɨɞɧɨɪɹɞɧɵɟ ɧɢɬɢ ɫ ɨɞɧɨɪɹɞɧɵɦɢ ɛɨɤɨɜɵɦɢ ɜɟɬɜɹɦɢ. Ʌɨɠɧɨɟ ɜɟɬɜɥɟɧɢɟ ɬɪɢɯɨɦɨɜ ɧɟ ɫɜɹɡɚɧɨ ɫ ɨɫɨɛɟɧɧɨɫɬɹɦɢ ɞɟɥɟɧɢɹ ɤɥɟɬɨɤ ɜɧɭɬɪɢ ɧɢɬɢ, ɚ ɟɫɬɶ ɪɟɡɭɥɶɬɚɬ ɩɪɢɤɪɟɩɥɟɧɢɹ ɢɥɢ ɫɨɟɞɢɧɟɧɢɹ ɧɢɬɟɣ ɩɨɞ ɭɝɥɨɦ ɞɪɭɝ ɤ ɞɪɭɝɭ.

Ⱦɥɹ ɪɚɡɧɵɯ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɷɬɨɣ ɝɪɭɩɩɵ ɩɪɨɤɚɪɢɨɬ ɯɚɪɚɤɬɟɪɧɚ ɫɩɨɫɨɛɧɨɫɬɶ ɤ ɫɤɨɥɶɡɹɳɟɦɭ ɞɜɢɠɟɧɢɸ, ɨɫɭɳɟɫɬɜɥɹɸɳɟɦɭɫɹ ɩɨ ɬɜɟɪɞɨɦɭ ɫɭɛɫɬɪɚɬɭ ɛɟɡ ɩɨɦɨɳɢ ɠɝɭɬɢɤɨɜ.

303

Ɉɫɧɨɜɧɨɣ ɫɩɨɫɨɛ ɪɚɡɦɧɨɠɟɧɢɹ ɰɢɚɧɨɛɚɤɬɟɪɢɣ – ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɟ ɛɢɧɚɪɧɨɟ ɞɟɥɟɧɢɟ. Ɉɞɧɚɤɨ ɩɪɢ ɨɩɪɟɞɟɥɟɧɧɵɯ ɭɫɥɨɜɢɹɯ ɜɟɝɟɬɚɬɢɜɧɚɹ ɤɥɟɬɤɚ ɝɟɧɟɬɢɱɟɫɤɢ ɩɪɟɞɪɚɫɩɨɥɨɠɟɧɚ ɤ ɞɢɮɮɟɪɟɧɰɢɚɰɢɢ ɜ ɫɩɟɰɢɚɥɢɡɢɪɨɜɚɧɧɭɸ ɪɟɩɪɨɞɭɤɬɢɜɧɭɸ ɫɬɪɭɤɬɭɪɭ. ɗɬɨ ɧɚɧɧɨɰɢɬɵ, ɛɚɟɨɰɢɬɵ ɢ ɷɤɡɨɫɩɨɪɵ ɭ ɨɞɧɨɤɥɟɬɨɱɧɵɯ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɢ ɝɨɪɦɨɝɨɧɢɢ ɢ ɝɨɪɦɨɰɢɫɬɵ ɭ ɧɢɬɱɚɬɵɯ ɰɢɚɧɨɛɚɤɬɟɪɢɣ.

ɇɚɧɧɨɰɢɬɵ ɨɛɪɚɡɭɸɬɫɹ ɜ ɪɟɡɭɥɶɬɚɬɟ ɪɹɞɚ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɵɯ ɞɟɥɟɧɢɣ ɦɚɬɟɪɢɧɫɤɨɣ ɤɥɟɬɤɢ, ɧɟ ɫɨɩɪɨɜɨɠɞɚɸɳɢɯɫɹ ɪɨɫɬɨɦ ɞɨɱɟɪɧɢɯ ɤɥɟɬɨɤ.

Ȼɚɟɨɰɢɬɵ ɨɛɪɚɡɭɸɬɫɹ ɜ ɪɟɡɭɥɶɬɚɬɟ ɦɧɨɝɨɤɪɚɬɧɨɝɨ ɞɟɥɟɧɢɹ ɤɥɟɬɨɤ ɜɧɭɬɪɢ ɨɛɳɟɣ ɞɨɩɨɥɧɢɬɟɥɶɧɨɣ ɨɛɨɥɨɱɤɢ-ɱɟɯɥɚ, ɜ ɪɟɡɭɥɶɬɚɬɟ ɤɨɬɨɪɨɝɨ ɨɛɪɚɡɭɟɬɫɹ ɫɩɨɪɚɧɝɢɣ ɫ ɦɟɥɤɢɦɢ ɛɚɟɨɰɢɬɚɦɢ. ɉɨɫɥɟɞɧɢɟ ɨɫɜɨɛɨɠɞɚɸɬɫɹ ɩɨɫɥɟ ɪɚɡɪɵɜɚ ɨɛɨɥɨɱɤɢ ɫɩɨɪɚɧɝɢɹ. Ɉɫɜɨɛɨɠɞɚɸɳɢɟɫɹ ɛɚɟɨɰɢɬɵ ɨɤɪɭɠɟɧɵ ɥɢɛɨ ɬɨɥɶɤɨ ɨɛɵɱɧɨɣ ɞɥɹ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɨɣ ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɨɝɨ ɬɢɩɚ, ɥɢɛɨ ɢɦɟɸɬ ɞɨɩɨɥɧɢɬɟɥɶɧɵɣ ɜɧɟɲɧɢɣ ɱɟɯɨɥ. Ȼɚɟɨɰɢɬɵ ɦɨɝɭɬ ɛɵɬɶ ɩɨɞɜɢɠɧɵɦɢ ɢ ɧɟɩɨɞɜɢɠɧɵɦɢ.

Ɉɛɪɚɡɨɜɚɧɢɟ ɷɤɡɨɫɩɨɪ ɩɪɨɢɫɯɨɞɢɬ ɬɚɤɠɟ ɩɭɬɟɦ ɞɟɥɟɧɢɹ ɤɥɟɬɤɢ ɜɧɭɬɪɢ ɱɟɯɥɚ. ɉɪɢ ɷɬɨɦ ɧɚɛɥɸɞɚɟɬɫɹ ɧɟɩɪɟɪɵɜɧɨɟ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɟ ɨɬɱɥɟɧɟɧɢɟ ɫɩɨɪ ɨɬ ɚɩɢɤɚɥɶɧɨɣ ɱɚɫɬɢ ɦɚɬɟɪɢɧɫɤɨɣ ɤɥɟɬɤɢ. Ȼɚɡɚɥɶɧɚɹ ɱɚɫɬɶ ɩɪɨɞɨɥɠɚɟɬ ɪɚɫɬɢ ɢ, ɞɨɫɬɢɝɧɭɜ ɢɫɯɨɞɧɵɯ ɪɚɡɦɟɪɨɜ, ɞɟɥɢɬɫɹ ɫ ɨɛɪɚɡɨɜɚɧɢɟɦ ɷɤɡɨɫɩɨɪɵ.

Ƚɨɪɦɨɝɨɧɢɢ ɩɪɟɞɫɬɚɜɥɹɸɬ ɫɨɛɨɣ ɮɪɚɝɦɟɧɬɵ ɬɪɢɯɨɦɚ, ɨɬɞɟɥɢɜɲɢɟɫɹ ɨɬ ɦɚɬɟɪɢɧɫɤɨɣ ɧɢɬɢ ɢ ɨɛɥɚɞɚɸɳɢɟ ɩɨɞɜɢɠɧɨɫɬɶɸ.

Ƚɨɪɦɨɰɢɫɬɵ, ɢɥɢ ɝɨɪɦɨɫɩɨɪɵ, – ɤɨɪɨɬɤɢɟ ɪɹɞɵ ɝɪɚɧɭɪɨɜɚɧɧɵɯ ɤɥɟɬɨɤ, ɨɤɪɭɠɟɧɧɵɟ ɫɚɦɨɫɬɨɹɬɟɥɶɧɵɦ ɩɥɨɬɧɵɦ ɬɨɥɫɬɵɦ ɫɥɢɡɢɫɬɵɦ ɱɟɯɥɨɦ.

ȼ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɰɢɚɧɨɛɚɤɬɟɪɢɢ ɪɚɡɞɟɥɟɧɵ ɧɚ ɩɹɬɶ ɩɨɪɹɞɤɨɜ, ɪɚɡɥɢɱɚɸɳɢɯɫɹ ɦɨɪɮɨɥɨɝɢɱɟɫɤɢɦɢ ɩɪɢɡɧɚɤɚɦɢ: Chroococcales, Pleurocapsales (ɨɞɢɧɨɱɧɵɟ ɤɥɟɬɤɢ ɢɥɢ ɤɨɥɨɧɢɢ), Oscillatoriales, Nostocales ɢ Stigonematales (ɦɧɨɝɨɤɥɟɬɨɱɧɵɟ ɧɢɬɱɚɬɵɟ ɮɨɪɦɵ).

Ɉɞɧɨɤɥɟɬɨɱɧɵɟ ɮɨɪɦɵ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɨɬɥɢɱɚɸɬɫɹ ɞɪɭɝ ɨɬ ɞɪɭɝɚ ɫɩɨɫɨɛɨɦ ɪɚɡɦɧɨɠɟɧɢɹ. Ɋɚɡɦɧɨɠɟɧɢɟ ɛɢɧɚɪɧɵɦ ɞɟɥɟɧɢɟɦ ɜ ɨɞɧɨɣ ɢɥɢ ɛɨɥɟɟ ɩɥɨɫɤɨɫɬɹɯ ɢɥɢ ɩɨɱɤɨɜɚɧɢɟɦ ɯɚɪɚɤɬɟɪɧɨ ɞɥɹ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɩɨɪɹɞɤɚ Chroococcales. ɐɢɚɧɨɛɚɤɬɟɪɢɢ, ɜɯɨɞɹɳɢɟ ɜ ɩɨɪɹɞɨɤ Pleurocapsales, ɪɚɡɦɧɨɠɚɸɬɫɹ ɦɧɨɠɟɫɬɜɟɧɧɵɦ ɞɟɥɟɧɢɟɦ ɢɥɢ ɱɟɪɟɞɨɜɚɧɢɟɦ ɛɢɧɚɪɧɨɝɨ ɢ ɦɧɨɠɟɫɬɜɟɧɧɨɝɨ ɞɟɥɟɧɢɹ (ɪɢɫ. 91).

Ⱥ

304

Ȼ

Ɋɢɫ. 91. Ɇɢɤɪɨɮɨɬɨɝɪɚɮɢɢ ɨɞɧɨɤɥɟɬɨɱɧɵɯ ɰɢɚɧɨɛɚɤɬɟɪɢɣ:

Ⱥ – Chroococcales; Ȼ – Pleurocapsales (ɢɡ «The Japanese Fresh-water Algae, 1977»; http://protist.i.hosei.ac.jp/PDB/Images/Prokaryotes/Chroococcaceae/)

305

ȼ ɨɫɧɨɜɟ ɞɟɥɟɧɢɹ ɧɚ ɩɨɪɹɞɤɢ ɦɧɨɝɨɤɥɟɬɨɱɧɵɯ ɮɨɪɦ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɥɟɠɢɬ ɫɩɨɫɨɛɧɨɫɬɶ ɧɢɬɟɣ ɤ ɜɟɬɜɥɟɧɢɸ, ɚ ɬɚɤɠɟ ɦɨɪɮɨɥɨɝɢɹ ɢ ɫɬɪɨɟɧɢɟ ɤɥɟɬɨɤ, ɢɡ ɤɨɬɨɪɵɯ ɨɧɢ ɨɛɪɚɡɨɜɚɧɵ. ɐɢɚɧɨɛɚɤɬɟɪɢɢ, ɜɯɨɞɹɳɢɟ ɜ ɩɨɪɹɞɨɤ Oscillatoriales, ɢɦɟɸɬ ɧɟɜɟɬɜɹɳɢɟɫɹ ɬɪɢɯɨɦɵ, ɫɨɫɬɨɹɳɢɟ ɢɡ ɨɞɧɨɝɨ ɪɹɞɚ ɬɨɥɶɤɨ ɜɟɝɟɬɚɬɢɜɧɵɯ ɤɥɟɬɨɤ. Ɋɨɫɬ ɬɪɢɯɨɦɚ ɨɫɭɳɟɫɬɜɥɹɟɬɫɹ ɞɟɥɟɧɢɟɦ ɤɥɟɬɨɤ ɜ ɨɞɧɨɣ ɩɥɨɫɤɨɫɬɢ. ɇɟɜɟɬɜɹɳɢɟɫɹ ɬɪɢɯɨɦɵ ɯɚɪɚɤɬɟɪɧɵ ɬɚɤɠɟ ɞɥɹ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɩɨɪɹɞɤɚ Nostocales. Ɉɞɧɚɤɨ ɢɯ ɬɪɢɯɨɦɵ, ɩɨɦɢɦɨ ɜɟɝɟɬɚɬɢɜɧɵɯ ɤɥɟɬɨɤ, ɫɨɞɟɪɠɚɬ ɝɟɬɟɪɨɰɢɫɬɵ ɢ ɢɧɨɝɞɚ ɚɤɢɧɟɬɵ. Ɉɬɥɢɱɢɬɟɥɶɧɵɦ ɩɪɢɡɧɚɤɨɦ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɩɨɪɹɞɤɚ Stigonematales ɹɜɥɹɟɬɫɹ ɫɩɨɫɨɛɧɨɫɬɶ ɜɟɝɟɬɚɬɢɜɧɵɯ ɤɥɟɬɨɤ ɬɪɢɯɨɦɚ ɤ ɞɟɥɟɧɢɸ ɛɨɥɟɟ ɱɟɦ ɜ ɨɞɧɨɣ ɩɥɨɫɤɨɫɬɢ, ɩɪɢɜɨɞɹɳɟɦɭ ɤ ɩɨɹɜɥɟɧɢɸ ɦɧɨɝɨɪɹɞɧɵɯ ɬɪɢɯɨɦɨɜ ɢɥɢ ɬɪɢɯɨɦɨɜ ɫ ɢɫɬɢɧɧɵɦ ɜɟɬɜɥɟɧɢɟɦ (ɪɢɫ. 92).

Ɋɢɫ. 92. Ɇɢɤɪɨɮɨɬɨɝɪɚɮɢɢ ɧɢɬɱɚɬɵɯ ɰɢɚɧɨɛɚɤɬɟɪɢɣ:

Ⱥ – Oscillatoriales; Ȼ – Nostocales; ȼ – Stigonematales (ɢɡ «The Japanese Fresh-water Algae, 1977»; http://protist.i.hosei.ac.jp/PDB/Images/Prokaryotes/Chroococcaceae/)

ɉɨɞɚɜɥɹɸɳɟɟ ɛɨɥɶɲɢɧɫɬɜɨ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɹɜɥɹɟɬɫɹ ɨɛɥɢɝɚɬɧɵɦɢ ɮɨɬɨɬɪɨɮɚɦɢ, ɬ. ɟ. ɦɨɝɭɬ ɪɚɫɬɢ ɬɨɥɶɤɨ ɡɚ ɫɱɟɬ ɷɧɟɪɝɢɢ ɫɨɥɧɟɱɧɨɝɨ ɫɜɟɬɚ. ɉɪɢ ɷɬɨɦ ɞɥɹ ɧɢɯ ɯɚɪɚɤɬɟɪɟɧ ɮɨɬɨɫɢɧɬɟɡ, ɨɫɧɨɜɚɧɧɵɣ ɧɚ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɢ ɞɜɭɯ ɮɨɬɨɫɢɫɬɟɦ ɫ ɢɫɩɨɥɶɡɨɜɚɧɢɟɦ ɇ2Ɉ ɜ ɤɚɱɟɫɬɜɟ ɞɨɧɨɪɚ ɷɥɟɤɬɪɨɧɨɜ ɢ ɫɨɩɪɨɜɨɠɞɚɸɳɢɣɫɹ ɜɵɞɟɥɟɧɢɟɦ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɤɢɫɥɨɪɨɞɚ. Ɉɛɪɚɡɭɸɳɢɟɫɹ ɜ ɩɪɨɰɟɫɫɟ ɮɨɬɨɫɢɧɬɟɡɚ ȺɌɎ ɢ ɇȺȾɎ · ɇ2 ɢɫɩɨɥɶɡɭɸɬɫɹ ɞɚɥɟɟ ɜ ɬɟɦɧɨɜɵɯ ɪɟɚɤɰɢɹɯ ɞɥɹ ɮɢɤɫɚɰɢɢ ɋɈ2 ɜ ɰɢɤɥɟ Ʉɚɥɶɜɢɧɚ. ȼ ɤɚɱɟɫɬɜɟ ɩɟɪɜɨɝɨ ɫɬɚɛɢɥɶɧɨɝɨ ɩɪɨɞɭɤɬɚ ɷɬɨɝɨ ɰɢɤɥɚ ɢɞɟɧɬɢɮɢɰɢɪɨɜɚɧ ɝɥɢɤɨɝɟɧ. ɉɨɦɢɦɨ ɷɬɨɝɨ ɩɭɬɢ, ɭ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɨɛɧɚɪɭɠɟɧɚ ɚɤɬɢɜɧɨɫɬɶ ɮɨɫɮɨɟɧɨɥɩɢɪɭɜɚɬɤɚɪɛɨɤɫɢɥɚɡɵ, ɱɬɨ ɩɨɡɜɨɥɹɟɬ ɢɦ ɚɫɫɢɦɢɥɢɪɨɜɚɬɶ ɋɈ2 ɩɭɬɟɦ ɤɚɪɛɨɤɫɢɥɢɪɨɜɚɧɢɹ ɮɨɫɮɨɟɧɨɥɩɢɪɨɜɢɧɨɝɪɚɞɧɨɣ ɤɢɫɥɨɬɨɣ.

ȼ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɭ ɦɧɨɝɢɯ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɨɛɧɚɪɭɠɟɧɚ ɫɩɨɫɨɛɧɨɫɬɶ ɤ ɛɟɫɤɢɫɥɨɪɨɞɧɨɦɭ ɮɨɬɨɫɢɧɬɟɡɭ, ɤɨɝɞɚ ɮɭɧɤɰɢɨɧɢɪɭɟɬ ɬɨɥɶɤɨ ɮɨɬɨɫɢɫɬɟɦɚ I. Ⱥɤɬɢɜɧɨɫɬɶ ɚɫɫɢɦɢɥɹɰɢɢ CO2 ɡɚ ɫɱɟɬ ɷɬɨɝɨ ɩɪɨɰɟɫɫɚ ɧɢɡɤɚ ɢ ɫɨɫɬɚɜɥɹɟɬ ɧɟɫɤɨɥɶɤɨ ɩɪɨɰɟɧɬɨɜ ɨɬ ɫɤɨɪɨɫɬɢ ɚɫɫɢɦɢɥɹɰɢɢ ɋɈ2 ɜ ɭɫɥɨɜɢɹɯ ɮɭɧɤɰɢɨɧɢɪɨɜɚɧɢɹ ɨɛɟɢɯ ɮɨɬɨɫɢɫɬɟɦ.

306

Ɉɛɧɚɪɭɠɟɧɢɟ ɭ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɛɟɫɤɢɫɥɨɪɨɞɧɨɝɨ ɮɨɬɨɫɢɧɬɟɡɚ ɩɨɡɜɨɥɹɟɬ ɥɢɤɜɢɞɢɪɨɜɚɬɶ «ɪɚɡɪɵɜ», ɫɭɳɟɫɬɜɭɸɳɢɣ ɦɟɠɞɭ ɮɨɬɨɫɢɧɬɟɡɨɦ ɩɭɪɩɭɪɧɵɯ ɢ ɡɟɥɟɧɵɯ ɛɚɤɬɟɪɢɣ, ɝɟɥɢɨɛɚɤɬɟɪɢɣ ɢ ɤɢɫɥɨɪɨɞɧɵɦ ɮɨɬɨɫɢɧɬɟɡɨɦ ɰɢɚɧɨɛɚɤɬɟɪɢɣ, ɩɪɨɯɥɨɪɨɮɢɬ ɢ ɷɭɤɚɪɢɨɬɢɱɟɫɤɢɯ ɨɪɝɚɧɢɡɦɨɜ. ɋɩɨɫɨɛɧɨɫɬɶ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɩɟɪɟɤɥɸɱɚɬɶɫɹ ɩɪɢ ɢɡɦɟɧɟɧɢɢ ɭɫɥɨɜɢɣ ɫ ɨɞɧɨɝɨ ɬɢɩɚ ɮɨɬɨɫɢɧɬɟɡɚ ɧɚ ɞɪɭɝɨɣ ɫɥɭɠɢɬ ɢɥɥɸɫɬɪɚɰɢɟɣ ɝɢɛɤɨɫɬɢ ɢɯ ɫɜɟɬɨɜɨɝɨ ɦɟɬɚɛɨɥɢɡɦɚ. ɗɬɨ ɢɦɟɟɬ ɜɚɠɧɨɟ ɷɤɨɥɨɝɢɱɟɫɤɨɟ ɡɧɚɱɟɧɢɟ ɞɥɹ ɰɢɚɧɨɛɚɤɬɟɪɢɣ.

ɏɨɬɹ ɩɨɞɚɜɥɹɸɳɟɟ ɛɨɥɶɲɢɧɫɬɜɨ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɦɨɝɭɬ ɪɚɫɬɢ, ɢɫɩɨɥɶɡɭɹ ɬɨɥɶɤɨ ɷɧɟɪɝɢɸ ɫɜɟɬɚ, ɬ. ɟ. ɹɜɥɹɸɬɫɹ ɨɛɥɢɝɚɬɧɵɦɢ ɮɨɬɨɬɪɨɮɚɦɢ, ɜ ɩɪɢɪɨɞɟ ɨɧɢ ɱɚɫɬɨ ɧɚɯɨɞɹɬɫɹ ɜ ɭɫɥɨɜɢɹɯ ɬɟɦɧɨɬɵ. ȼ ɬɟɦɧɨɬɟ ɭ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɨɛɧɚɪɭɠɟɧ ɚɤɬɢɜɧɵɣ ɷɧɞɨɝɟɧɧɵɣ ɦɟɬɚɛɨɥɢɡɦ, ɷɧɟɪɝɟɬɢɱɟɫɤɢɦ ɫɭɛɫɬɪɚɬɨɦ ɤɨɬɨɪɨɝɨ ɫɥɭɠɢɬ ɡɚɩɚɫɟɧɧɵɣ ɧɚ ɫɜɟɬɭ ɝɥɢɤɨɝɟɧ. ȼ ɤɚɱɟɫɬɜɟ ɨɫɧɨɜɧɨɝɨ ɩɭɬɢ ɤɚɬɚɛɨɥɢɡɢɪɨɜɚɧɢɹ ɩɨɫɥɟɞɧɟɝɨ ɢɞɟɧɬɢɮɢɰɢɪɨɜɚɧ ɨɤɢɫɥɢɬɟɥɶɧɵɣ ɩɟɧɬɨɡɨɮɨɫɮɚɬɧɵɣ ɩɭɬɶ. Ⱦɪɭɝɨɣ ɜɨɡɦɨɠɧɵɣ ɩɭɬɶ ɩɨɥɭɱɟɧɢɹ ɰɢɚɧɨɛɚɤɬɟɪɢɹɦɢ ɜ ɬɟɦɧɨɬɟ ɷɧɟɪɝɢɢ – ɝɥɢɤɨɥɢɡ.

ɐɢɚɧɨɛɚɤɬɟɪɢɢ, ɨɫɭɳɟɫɬɜɥɹɸɳɢɟ ɛɟɫɤɢɫɥɨɪɨɞɧɵɣ ɮɨɬɨɫɢɧɬɟɡ, ɦɨɝɭɬ ɩɨɥɭɱɚɬɶ ɷɧɟɪɝɢɸ ɜ ɬɟɦɧɨɬɟ ɜ ɩɪɨɰɟɫɫɟ ɚɧɚɷɪɨɛɧɨɝɨ ɞɵɯɚɧɢɹ ɩɪɢ ɧɚɥɢɱɢɢ ɜ ɫɪɟɞɟ ɫɟɪɵ. ɉɪɢ ɷɬɨɦ ɩɪɨɢɫɯɨɞɢɬ ɩɟɪɟɧɨɫ ɷɥɟɤɬɪɨɧɨɜ ɧɚ ɦɨɥɟɤɭɥɹɪɧɭɸ ɫɟɪɭ, ɜɨɫɫɬɚɧɚɜɥɢɜɚɹ ɟɟ ɞɨ ɫɭɥɶɮɢɞɚ. ɍɫɬɚɧɨɜɥɟɧɨ, ɱɬɨ ɭ ɬɚɤɢɯ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɰɢɤɥ Ʉɪɟɛɫɚ «ɧɟ ɡɚɦɤɧɭɬ» ɢɡ-ɡɚ ɨɬɫɭɬɫɬɜɢɹ Į-ɤɟɬɨɝɥɭɬɚ- ɪɚɬɞɟɝɢɞɪɨɝɟɧɚɡɵ.

ɋɪɟɞɢ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɲɢɪɨɤɨ ɪɚɫɩɪɨɫɬɪɚɧɟɧɚ ɫɩɨɫɨɛɧɨɫɬɶ ɤ ɚɡɨɬɮɢɤɫɚɰɢɢ. ɗɬɨ ɨɛɟɫɩɟɱɢɜɚɟɬɫɹ ɧɚɥɢɱɢɟɦ ɧɢɬɪɨɝɟɧɚɡɧɨɣ ɚɤɬɢɜɧɨɫɬɢ, ɤɨɬɨɪɚɹ ɜ ɫɜɨɸ ɨɱɟɪɟɞɶ ɡɚɜɢɫɢɬ ɨɬ ɫɨɞɟɪɠɚɧɢɹ ɜ ɫɪɟɞɟ ɫɜɹɡɚɧɧɨɝɨ ɚɡɨɬɚ ɢ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɤɢɫɥɨɪɨɞɚ. ɋɜɹɡɚɧɧɵɣ ɚɡɨɬ ɪɟɩɪɟɫɫɢɪɭɟɬ ɫɢɧɬɟɡ ɢ ɢɧɝɢɛɢɪɭɟɬ ɚɤɬɢɜɧɨɫɬɶ ɧɢɬɪɨɝɟɧɚɡɵ, ɦɨɥɟɤɭɥɹɪɧɵɣ ɤɢɫɥɨɪɨɞ ɛɵɫɬɪɨ ɢɧɚɤɬɢɜɢɪɭɟɬ ɮɟɪɦɟɧɬ.

ɉɪɨɛɥɟɦɚ ɮɢɤɫɚɰɢɢ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɚɡɨɬɚ ɜ ɚɷɪɨɛɧɵɯ ɭɫɥɨɜɢɹɯ ɭ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɪɟɲɟɧɚ ɩɭɬɟɦ ɮɨɪɦɢɪɨɜɚɧɢɹ ɞɢɮɮɟɪɟɧɰɢɪɨɜɚɧɧɵɯ ɤɥɟɬɨɤ – ɝɟɬɟɪɨɰɢɫɬ, ɜ ɤɨɬɨɪɵɯ ɱɭɜɫɬɜɢɬɟɥɶɧɵɣ ɤ ɦɨɥɟɤɭɥɹɪɧɨɦɭ ɤɢɫɥɨɪɨɞɭ ɚɩɩɚɪɚɬ ɮɢɤɫɚɰɢɢ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɚɡɨɬɚ ɨɬɞɟɥɟɧ ɨɬ ɤɢɫɥɨɪɨɞɨɜɵɞɟɥɹɸɳɟɝɨ ɮɨɬɨɫɢɧɬɟɬɢɱɟɫɤɨɝɨ ɚɩɩɚɪɚɬɚ ɫ ɩɨɦɨɳɶɸ ɨɩɪɟɞɟɥɟɧɧɵɯ ɭɥɶɬɪɚɫɬɪɭɤɬɭɪɧɵɯ ɢ ɛɢɨɯɢɦɢɱɟɫɤɢɯ ɩɟɪɟɫɬɪɨɟɤ.

ɉɪɢ ɨɬɫɭɬɫɬɜɢɢ ɜ ɫɪɟɞɟ ɫɜɹɡɚɧɧɨɝɨ ɚɡɨɬɚ ɧɟɤɨɬɨɪɵɟ ɤɥɟɬɤɢ ɧɢɬɱɚɬɵɯ ɰɢɚɧɨɛɚɤɬɟɪɢɣ ɩɪɟɜɪɚɳɚɸɬɫɹ ɜ ɝɟɬɟɪɨɰɢɫɬɵ. Ɉɛɪɚɡɨɜɚɧɢɟ ɝɟɬɟɪɨɰɢɫɬ ɩɪɨɢɫɯɨɞɢɬ ɡɚ 24 ɱ ɢ ɦɨɠɟɬ ɛɵɬɶ ɪɚɡɞɟɥɟɧɨ ɧɚ ɞɜɚ ɷɬɚɩɚ:

1. Ɏɨɪɦɢɪɭɸɳɢɟɫɹ ɧɚ ɩɟɪɜɨɦ ɷɬɚɩɟ ɩɪɨɝɟɬɟɪɨɰɢɫɬɵ ɧɟ ɫɩɨɫɨɛɧɵ ɡɚɳɢɬɢɬɶ ɧɢɬɪɨɝɟɧɚɡɭ ɨɬ ɢɧɚɤɬɢɜɢɪɭɸɳɟɝɨ ɞɟɣɫɬɜɢɹ Ɉ2. ɉɪɨɰɟɫɫ ɞɢɮɮɟɪɟɧɰɢɪɨɜɤɢ ɧɚ ɷɬɨɣ ɫɬɚɞɢɢ ɨɛɪɚɬɢɦ: ɩɪɢ ɜɧɟɫɟɧɢɢ ɜ ɫɪɟɞɭ ɚɦɦɨɧɢɣ-

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