microbiology_konspekt_lysak_2008

Micrococcus, Actinomyces, Bifidobacterium, Propionibacterium, Actinoplanes, Nocardia, Rhodococcus, Corynebacterium, Mycobacterium. Ƚɪɭɩɩɚ

Firmicutes («ɤɥɨɫɬɪɢɞɢɚɥɶɧɚɹ ɜɟɬɜɶ» – ɝɥɚɜɧɵɦ ɨɛɪɚɡɨɦ ɝɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɟ ɛɚɤɬɟɪɢɢ ɫ ɧɢɡɤɢɦ ɫɨɞɟɪɠɚɧɢɟɦ Ƚɐ-ɩɚɪ ɜ ȾɇɄ) ɫɨɫɬɨɢɬ ɢɡ ɫɥɟɞɭɸɳɢɯ ɪɨɞɨɜ: Clostridium, Lactococcus, Pediococcus, Streptococcus, Enterococcus, Leuconostoc, Listeria, Caryophanon, Staphylococcus, Sarcina, Sporosɚrcina, Bacillus, Desulfotomaculum, Heliobacterium, Mycoplasma, Ureaplasma ɢ ɞɪ.

Ɍɚɛɥɢɰɚ 2

Ƚɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɛɚɤɬɟɪɢɢ ɮɢɥɨɝɟɧɟɬɢɱɟɫɤɨɣ ɝɪɭɩɩɵ

Proteobacteria (ɉɪɨɬɟɨɛɚɤɬɟɪɢɢ)

	Ɉɫɧɨɜɧɵɟ ɮɟɧɨɬɢɩɢɱɟɫɤɢɟ	ɇɚɢɛɨɥɟɟ ɪɚɫɩɪɨɫɬɪɚɧɟɧɧɵɟ ɪɨɞɵ
	ɝɪɭɩɩɵ
	Ɏɟɪɦɟɧɬɢɪɭɸɳɢɟ ɩɚɥɨɱɤɢ	ɗɧɬɟɪɨɛɚɤɬɟɪɢɢ, Vibrio, Photobacterium, Aeromo-
	ɢ ɜɢɛɪɢɨɧɵ	nas, Zymomonas
	ɉɚɥɨɱɤɢ ɢ ɤɨɤɤɢ, ɨɛɥɚɞɚɸɳɢɟ	Pseudomonas, Zoogloea, Azotobacter, Beijerinckia,
	ɚɷɪɨɛɧɵɦ ɞɵɯɚɧɢɟɦ	Azomonas, Rhizobium, Bradyrhizobium, Agrobacte-
		rium, Acetobacter, Gluconobacter, Legionella, Neis-
		seria, Acinetobacter, Rickettsia
	Ȼɚɤɬɟɪɢɢ, ɨɛɪɚɡɭɸɳɢɟ ɱɟɯɥɵ	Sphaerotilus, Leptothrix, Crenothrix
	Ȼɚɤɬɟɪɢɢ, ɨɛɪɚɡɭɸɳɢɟ ɩɪɨɫɬɟɤɢ	Caulobacter, Hyphomicrobium
	ɉɚɪɚɡɢɬɵ ɛɚɤɬɟɪɢɣ	Bdellovibrio
	ɋɩɢɪɢɥɥɵ ɢ ɦɚɝɧɢɬɨɫɩɢɪɢɥɥɵ	Spirillum, Aquaspirillum, Magnetospirillum,
		Campylobacter, Helicobacter
	Ɇɢɤɫɨɛɚɤɬɟɪɢɢ	Polyangium, Myxococcus
	Ȼɚɤɬɟɪɢɢ, ɜɨɫɫɬɚɧɚɜɥɢ-	Desulfovibrio, Desulfococcus, Desulfosarcina, Desul-
	ɜɚɸɳɢɟ ɫɭɥɶɮɚɬɵ ɢ ɫɟɪɭ	furomonas
	ɇɢɬɪɢɮɢɤɚɬɨɪɵ	Nitrosomonas, Nitrosospira, Nitrosococcus, Nitro-
		bacter, Nitrococcus
	Ȼɚɤɬɟɪɢɢ, ɨɤɢɫɥɹɸɳɢɟ ɫɟɪɭ	Thiobacillus, Thiomicrospira, Thermothrix, Beggiatoa,
	ɢ ɠɟɥɟɡɨ	Thiothrix, Gallionella
	Ȼɚɤɬɟɪɢɢ, ɨɤɢɫɥɹɸɳɢɟ ɜɨɞɨɪɨɞ	Alcaligenes, Ancylobacter, Paracoccus, Rhizobium,
		Pseudomonas, Spirillum
	Ɇɟɬɢɥɨɬɪɨɮɧɵɟɛɚɤɬɟɪɢɢ	Methylomonas, Methylocystis, Methylobacter, Methy-
		lococcus
	Ɏɨɬɨɫɢɧɬɟɡɢɪɭɸɳɢɟ ɩɭɪɩɭɪ-	ɋɟɪɧɵɟ: Chromatium, Thiospirillum, Thiocapsa;
	ɧɵɟ ɛɚɤɬɟɪɢɢ	ɧɟɫɟɪɧɵɟ: Rhodobacter, Rhodopseudomonas,
		Rhodospirillum, Rhodocyclus

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ȼɫɨɫɬɚɜɟ ɚɪɯɟɛɚɤɬɟɪɢɣ ɜɵɞɟɥɹɸɬ ɬɪɢ ɮɢɥɨɝɟɧɟɬɢɱɟɫɤɢɟ ɝɪɭɩɩɵ:

Crenarchaeota, Euryarchaeota ɢ Korarchaeota. Ƚɪɭɩɩɚ Crenarchaeota ɫɨ-

ɫɬɨɢɬ ɢɡ ɷɤɫɬɪɟɦɚɥɶɧɨ ɬɟɪɦɨɮɢɥɶɧɵɯ ɛɚɤɬɟɪɢɣ, ɛɨɥɶɲɢɧɫɬɜɨ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɤɨɬɨɪɵɯ ɨɫɭɳɟɫɬɜɥɹɸɬ ɦɟɬɚɛɨɥɢɡɦ ɫɟɪɵ, ɧɟɤɨɬɨɪɵɟ ɜɨɫɫɬɚɧɚɜɥɢɜɚɸɬ ɢɨɧɵ ɠɟɥɟɡɚ ɢ ɦɨɥɢɛɞɟɧɚ. ȼ ɝɪɭɩɩɭ Euryarchaeota ɜɯɨɞɹɬ ɨɛɥɢɝɚɬɧɨ ɚɧɚɷɪɨɛɧɵɟ ɦɟɬɚɧɨɝɟɧɧɵɟ ɚɪɯɟɛɚɤɬɟɪɢɢ, ɚ ɬɚɤɠɟ ɷɤɫɬɪɟɦɚɥɶɧɵɟ ɬɟɪɦɨɮɢɥɵ ɢ ɝɚɥɨɮɢɥɵ. Ƚɪɭɩɩɚ Korarchaeota ɨɛɪɚɡɨɜɚɧɚ ɚɪɯɟɛɚɤɬɟɪɢɹɦɢ, ɨɛɢɬɚɸɳɢɦɢ ɜ ɝɨɪɹɱɢɯ ɫɟɪɧɵɯ ɢɫɬɨɱɧɢɤɚɯ. Ⱦɨ ɧɚɫɬɨɹɳɟɝɨ ɜɪɟɦɟɧɢ ɧɢ ɨɞɢɧ ɢɡ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɷɬɨɣ ɝɪɭɩɩɵ (ɨɛɥɚɞɚɸɳɢɯ ɫɯɨɞɧɨɣ 16S-ɪɊɇɄ) ɧɟ ɜɵɞɟɥɟɧ ɜ ɜɢɞɟ ɱɢɫɬɨɣ ɤɭɥɶɬɭɪɵ, ɩɨɷɬɨɦɭ ɢɯ ɮɟɧɨɬɢɩɢɱɟɫɤɢɟ ɩɪɢɡɧɚɤɢ ɢɡɭɱɟɧɵ ɧɟɞɨɫɬɚɬɨɱɧɨ.

Ɂɚɤɚɧɱɢɜɚɹ ɪɚɫɫɦɨɬɪɟɧɢɟ ɮɢɥɨɝɟɧɟɬɢɱɟɫɤɢɯ ɜɟɬɜɟɣ ɩɪɨɤɚɪɢɨɬ, ɫɥɟɞɭɟɬ ɨɬɦɟɬɢɬɶ, ɱɬɨ ɩɪɟɞɥɨɠɟɧɧɚɹ ɮɢɥɨɝɟɧɟɬɢɱɟɫɤɚɹ ɫɢɫɬɟɦɚ, ɨɫɧɨɜɚɧɧɚɹ ɧɚ ɢɫɫɥɟɞɨɜɚɧɢɢ ɧɭɤɥɟɨɬɢɞɧɵɯ ɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɟɣ ɬɨɥɶɤɨ ɨɞɧɨɝɨ ɝɟɧɚ ɪɢɛɨɫɨɦɧɨɣ ɊɇɄ – ɧɟ ɛɨɥɟɟ ɱɟɦ ɨɞɧɚ ɢɡ ɬɟɯɧɢɱɟɫɤɢ ɭɞɨɛɧɵɯ ɢ ɪɚɡɪɚɛɨɬɚɧɧɵɯ ɫɢɫɬɟɦ ɭɩɨɪɹɞɨɱɟɧɢɹ ɦɧɨɝɨɱɢɫɥɟɧɧɵɯ ɨɪɝɚɧɢɡɦɨɜ ɜ ɰɟɥɹɯ ɢɯ ɢɞɟɧɬɢɮɢɤɚɰɢɢ, ɩɨɷɬɨɦɭ ɩɨɫɬɪɨɢɬɶ ɥɨɝɢɱɟɫɤɢ ɜɟɪɧɭɸ ɬɚɤɫɨɧɨɦɢɸ ɛɚɤɬɟɪɢɣ ɬɨɥɶɤɨ ɫ ɭɱɟɬɨɦ ɷɬɨɝɨ ɩɪɢɡɧɚɤɚ ɧɟ ɩɪɟɞɫɬɚɜɥɹɟɬɫɹ ɜɨɡɦɨɠɧɵɦ.

ɇɚɢɛɨɥɟɟ ɩɪɢɡɧɚɧɧɨɣ ɢ ɢɫɩɨɥɶɡɭɟɦɨɣ ɮɟɧɨɬɢɩɢɱɟɫɤɨɣ ɤɥɚɫɫɢɮɢɤɚɰɢɟɣ ɛɚɤɬɟɪɢɣ ɹɜɥɹɟɬɫɹ ɤɥɚɫɫɢɮɢɤɚɰɢɹ, ɩɪɟɞɫɬɚɜɥɟɧɧɚɹ ɜ ɞɟɜɹɬɨɦ ɢɡɞɚɧɢɢ Ɉɩɪɟɞɟɥɢɬɟɥɹ ɛɚɤɬɟɪɢɣ Ȼɟɪɞɠɢ. ȼ ɷɬɨɦ ɢɡɞɚɧɢɢ ɛɚɤɬɟɪɢɢ ɧɚ ɨɫɧɨɜɚɧɢɢ ɫɬɪɨɟɧɢɹ ɩɨɝɪɚɧɢɱɧɨɝɨ ɫɥɨɹ ɤɥɟɬɤɢ ɪɚɡɞɟɥɟɧɵ ɧɚ ɱɟɬɵɪɟ ɨɫɧɨɜɧɵɟ ɤɚɬɟɝɨɪɢɢ (ɨɬɞɟɥɚ): 1) Gracilicutes (ɨɬ ɥɚɬ. cutes – ɤɨɠɚ, gracilis – ɬɨɧɤɢɣ) – ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɷɭɛɚɤɬɟɪɢɢ, ɢɦɟɸɳɢɟ ɤɥɟɬɨɱɧɵɟ ɫɬɟɧɤɢ;

2)Firmicutes (ɨɬ ɥɚɬ. firmus – ɩɪɨɱɧɵɣ) – ɝɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɟ ɷɭɛɚɤɬɟɪɢɢ, ɢɦɟɸɳɢɟ ɤɥɟɬɨɱɧɵɟ ɫɬɟɧɤɢ; 3) Tenericutes (ɨɬ ɥɚɬ. tener – ɦɹɝɤɢɣ, ɧɟɠɧɵɣ) – ɷɭɛɚɤɬɟɪɢɢ, ɥɢɲɟɧɧɵɟ ɤɥɟɬɨɱɧɵɯ ɫɬɟɧɨɤ; 4) Mendosicutes (ɨɬ ɥɚɬ. mendosus – ɨɲɢɛɨɱɧɵɣ) – ɚɪɯɟɛɚɤɬɟɪɢɢ, ɤɥɟɬɨɱɧɵɟ ɫɬɟɧɤɢ ɤɨɬɨɪɵɯ ɨɬɥɢɱɚɸɬɫɹ ɨɬ ɚɧɚɥɨɝɢɱɧɵɯ ɫɬɪɭɤɬɭɪ ɞɪɭɝɢɯ ɩɪɨɤɚɪɢɨɬ.

ȼɨɬɞɟɥ Gracilicutes ɜɯɨɞɹɬ ɛɚɤɬɟɪɢɢ ɪɚɡɥɢɱɧɨɣ ɦɨɪɮɨɥɨɝɢɢ ɫ ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɨɣ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɨɣ. Ɋɚɡɦɧɨɠɟɧɢɟ ɩɪɨɢɫɯɨɞɢɬ ɜ ɨɫɧɨɜɧɨɦ ɛɢɧɚɪɧɵɦ ɞɟɥɟɧɢɟɦ, ɧɟɤɨɬɨɪɵɟ ɛɚɤɬɟɪɢɢ ɪɚɡɦɧɨɠɚɸɬɫɹ ɩɨɱɤɨɜɚɧɢɟɦ. ɗɧɞɨɫɩɨɪ ɧɟ ɨɛɪɚɡɭɸɬ. Ȼɨɥɶɲɢɧɫɬɜɨ ɩɨɞɜɢɠɧɵ: ɜɫɬɪɟɱɚɸɬɫɹ ɜɫɟ ɬɢɩɵ ɩɟɪɟɞɜɢɠɟɧɢɹ ɛɚɤɬɟɪɢɣ – ɫ ɩɨɦɨɳɶɸ ɠɝɭɬɢɤɨɜ, ɫɤɨɥɶɠɟɧɢɟɦ, ɢɡɝɢɛɚɧɢɟɦ. Ɉɬɞɟɥ ɜɤɥɸɱɚɟɬ ɚɷɪɨɛɧɵɟ, ɚɧɚɷɪɨɛɧɵɟ ɢ ɮɚɤɭɥɶɬɚɬɢɜɧɵɟ ɚɧɚɷɪɨɛɧɵɟ ɛɚɤɬɟɪɢɢ; ɮɨɬɨɬɪɨɮɧɵɟ ɢ ɯɟɦɨɬɪɨɮɧɵɟ ɛɚɤɬɟɪɢɢ. Ɉɬɞɟɥ ɩɨɞ-

ɪɚɡɞɟɥɹɸɬ ɧɚ ɬɪɢ ɤɥɚɫɫɚ: Scotobacteria, Oxyphotobacteria, Anoxyphotobacteria. ȼ ɤɥɚɫɫ Scotobacteria ɜɯɨɞɹɬ ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɛɚɤɬɟɪɢɢ, ɧɟ ɢɫɩɨɥɶɡɭɸɳɢɟ ɫɜɟɬɨɜɭɸ ɷɧɟɪɝɢɸ ɞɥɹ ɰɟɥɟɣ ɦɟɬɚɛɨɥɢɡɦɚ, ɚ ɩɨɥɭɱɚɸɳɢɟ ɟɟ

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ɬɨɥɶɤɨ ɜ ɪɟɡɭɥɶɬɚɬɟ ɨɤɢɫɥɢɬɟɥɶɧɨ-ɜɨɫɫɬɚɧɨɜɢɬɟɥɶɧɵɯ ɪɟɚɤɰɢɣ. ɇɚɡɜɚɧɢɟ ɤɥɚɫɫɚ ɩɪɨɢɫɯɨɞɢɬ ɨɬ ɝɪɟɱ. sɫotos – ɬɟɦɧɨɬɚ. ɗɬɨ ɫɚɦɵɣ ɤɪɭɩɧɵɣ ɤɥɚɫɫ ɛɚɤɬɟɪɢɣ. ȼ ɤɥɚɫɫ Anoxyphotobacteria ɜɯɨɞɹɬ ɩɭɪɩɭɪɧɵɟ ɛɚɤɬɟɪɢɢ, ɡɟɥɟɧɵɟ ɛɚɤɬɟɪɢɢ ɢ ɝɟɥɢɨɛɚɤɬɟɪɢɢ, ɨɫɭɳɟɫɬɜɥɹɸɳɢɟ ɚɧɨɤɫɢɝɟɧɧɵɣ ɮɨɬɨɫɢɧɬɟɡ (ɛɟɡ ɜɵɞɟɥɟɧɢɹ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɤɢɫɥɨɪɨɞɚ). Ʉɥɚɫɫ Oxyphotobacteria ɩɪɟɞɫɬɚɜɥɟɧ ɰɢɚɧɨɛɚɤɬɟɪɢɹɦɢ ɢ ɩɪɨɯɥɨɪɨɮɢɬɚɦɢ, ɨɫɭɳɟɫɬɜɥɹɸɳɢɦɢ ɨɤɫɢɝɟɧɧɵɣ ɮɨɬɨɫɢɧɬɟɡ (ɫ ɜɵɞɟɥɟɧɢɟɦ ɦɨɥɟɤɭɥɹɪɧɨɝɨ ɤɢɫɥɨɪɨɞɚ). ɗɬɨɬ ɬɢɩ ɮɨɬɨɫɢɧɬɟɡɚ ɚɧɚɥɨɝɢɱɟɧ ɮɨɬɨɫɢɧɬɟɡɭ, ɩɪɨɬɟɤɚɸɳɟɦɭ ɜ ɪɚɫɬɟɧɢɹɯ.

ȼ ɨɬɞɟɥ Firmicutes ɜɤɥɸɱɟɧɵ ɛɚɤɬɟɪɢɢ ɫ ɝɪɚɦɩɨɥɨɠɢɬɟɥɶɧɨɣ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɨɣ. Ʉɥɟɬɤɢ ɦɨɝɭɬ ɢɦɟɬɶ ɪɚɡɧɭɸ ɮɨɪɦɭ: ɩɚɥɨɱɤɢ, ɤɨɤɤɢ, ɧɢɬɟɜɢɞɧɵɟ, ɜɟɬɜɹɳɢɟɫɹ. ɇɟɤɨɬɨɪɵɟ ɩɪɟɞɫɬɚɜɢɬɟɥɢ ɨɛɪɚɡɭɸɬ ɷɧɞɨɫɩɨɪɵ. Ȼɨɥɶɲɢɧɫɬɜɨ ɢɡ ɧɢɯ ɧɟɩɨɞɜɢɠɧɵ; ɩɨɞɜɢɠɧɵɟ ɮɨɪɦɵ ɢɦɟɸɬ ɩɟɪɢɬɪɢɯɢɚɥɶɧɨɟ ɠɝɭɬɢɤɨɜɚɧɢɟ. ȼ ɫɨɫɬɚɜ ɨɬɞɟɥɚ ɜɯɨɞɹɬ ɚɷɪɨɛɧɵɟ, ɚɧɚɷɪɨɛɧɵɟ ɢ ɮɚɤɭɥɶɬɚɬɢɜɧɨ ɚɧɚɷɪɨɛɧɵɟ ɛɚɤɬɟɪɢɢ. Ɉɬɞɟɥ ɫɨɫɬɨɢɬ ɢɡ ɞɜɭɯ ɤɥɚɫɫɨɜ:

Firmibacteria, Thallobacteria. Ʉɥɚɫɫ Firmibacteria ɜɤɥɸɱɚɟɬ ɛɨɥɶɲɨɟ ɤɨ-

ɥɢɱɟɫɬɜɨ «ɧɟɜɟɬɜɹɳɢɯɫɹ» ɝɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɯ ɛɚɤɬɟɪɢɣ. Ʉɥɚɫɫ Thallobacteria ɜɤɥɸɱɚɟɬ ɛɚɤɬɟɪɢɢ, ɤɥɟɬɤɢ ɤɨɬɨɪɵɯ ɫɩɨɫɨɛɧɵ «ɜɟɬɜɢɬɶɫɹ».

Ɉɬɞɟɥ Tenericutes ɩɪɟɞɫɬɚɜɥɟɧ ɛɚɤɬɟɪɢɹɦɢ, ɧɟ ɢɦɟɸɳɢɦɢ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ. ȼ ɫɜɹɡɢ ɫ ɨɬɫɭɬɫɬɜɢɟɦ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ ɮɨɪɦɚ ɤɥɟɬɨɤ ɧɟɩɨɫɬɨɹɧɧɚ: ɜ ɱɢɫɬɨɣ ɤɭɥɶɬɭɪɟ ɨɞɧɨɝɨ ɜɢɞɚ ɨɞɧɨɜɪɟɦɟɧɧɨ ɩɪɢɫɭɬɫɬɜɭɸɬ ɤɨɤɤɨɜɢɞɧɵɟ, ɩɚɥɨɱɤɨɜɢɞɧɵɟ, ɧɢɬɟɜɢɞɧɵɟ, ɝɪɭɲɟɜɢɞɧɵɟ, ɞɢɫɤɨɜɢɞɧɵɟ ɢ ɞɪɭɝɢɟ ɤɥɟɬɤɢ. Ɋɚɡɦɧɨɠɟɧɢɟ ɛɚɤɬɟɪɢɣ, ɜɯɨɞɹɳɢɯ ɜ ɷɬɨɬ ɨɬɞɟɥ, ɩɪɨɢɫɯɨɞɢɬ ɛɢɧɚɪɧɵɦ ɞɟɥɟɧɢɟɦ, ɩɨɱɤɨɜɚɧɢɟɦ. Ɉɤɪɚɲɢɜɚɧɢɟ ɩɨ Ƚɪɚɦɭ ɨɬɪɢɰɚɬɟɥɶɧɨɟ. ɏɚɪɚɤɬɟɪɧɨ ɨɛɪɚɡɨɜɚɧɢɟ ɦɟɥɤɢɯ, ɜɪɚɫɬɚɸɳɢɯ ɜ ɚɝɚɪ ɤɨɥɨɧɢɣ. Ɇɨɝɭɬ ɛɵɬɶ ɫɚɩɪɨɮɢɬɧɵɦɢ, ɩɚɪɚɡɢɬɚɦɢ ɢɥɢ ɩɚɬɨɝɟɧɚɦɢ. Ɉɬɞɟɥ ɫɨɫɬɨɢɬ ɢɡ ɨɞɧɨɝɨ ɤɥɚɫɫɚ Mollicutes (ɦɢɤɨɩɥɚɡɦɵ).

Ɉɬɞɟɥ Mendosicutes ɨɛɪɚɡɨɜɚɧ ɛɚɤɬɟɪɢɹɦɢ ɫ ɪɢɝɢɞɧɨɣ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɨɣ, ɧɨ ɧɟ ɫɨɞɟɪɠɚɳɟɣ ɩɟɩɬɢɞɨɝɥɢɤɚɧɚ ɦɭɪɟɢɧɚ. Ȼɨɥɶɲɢɧɫɬɜɨ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ – ɫɬɪɨɝɢɟ ɚɧɚɷɪɨɛɵ, ɦɧɨɝɢɟ ɢɡ ɤɨɬɨɪɵɯ ɢɦɟɸɬ ɠɝɭɬɢɤɢ. ȼɢɞɵ ɯɚɪɚɤɬɟɪɢɡɭɸɬɫɹ ɷɤɨɥɨɝɢɱɟɫɤɢɦ ɢ ɦɟɬɚɛɨɥɢɱɟɫɤɢɦ ɪɚɡɧɨɨɛɪɚɡɢɟɦ, ɫɩɨɫɨɛɧɨɫɬɶɸ ɠɢɬɶ ɜ ɷɤɫɬɪɟɦɚɥɶɧɵɯ ɭɫɥɨɜɢɹɯ. Ɉɬɞɟɥ ɫɨɫɬɨɢɬ ɢɡ ɨɞɧɨɝɨ ɤɥɚɫɫɚ – Archaebacteria.

ȼ ɫɨɫɬɚɜɟ ɱɟɬɵɪɟɯ ɨɬɞɟɥɨɜ (ɨɫɧɨɜɧɵɯ ɤɚɬɟɝɨɪɢɣ) ɜɵɞɟɥɟɧɨ 35 ɝɪɭɩɩ (ɢɥɢ ɫɟɤɰɢɣ) ɛɚɤɬɟɪɢɣ, ɤɨɬɨɪɵɟ ɜ ɛɨɥɶɲɟɣ ɢɥɢ ɦɟɧɶɲɟɣ ɫɬɟɩɟɧɢ ɛɭɞɭɬ ɨɯɚɪɚɤɬɟɪɢɡɨɜɚɧɵ ɜ ɩɨɫɥɟɞɭɸɳɢɯ ɝɥɚɜɚɯ.

Ʉ ɨɬɞɟɥɭ Gracilicutes ɩɪɢɧɚɞɥɟɠɚɬ ɫɥɟɞɭɸɳɢɟ ɝɪɭɩɩɵ. Ƚɪɭɩɩɚ 1. ɋɩɢɪɨɯɟɬɵ.

Ƚɪɭɩɩɚ 2. Ⱥɷɪɨɛɧɵɟ (ɢɥɢ ɦɢɤɪɨɚɷɪɨɮɢɥɶɧɵɟ), ɩɨɞɜɢɠɧɵɟ, ɫɩɢɪɚɥɟɜɢɞɧɵɟ (ɢɥɢ ɜɢɛɪɢɨɢɞɧɵɟ) ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɛɚɤɬɟɪɢɢ.

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Ƚɪɭɩɩɚ 3. ɇɟɩɨɞɜɢɠɧɵɟ ɢɥɢ ɪɟɞɤɨ ɩɨɞɜɢɠɧɵɟ ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɢɡɨɝɧɭɬɵɟ ɛɚɤɬɟɪɢɢ.

Ƚɪɭɩɩɚ 4. Ƚɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɚɷɪɨɛɧɵɟ (ɢɥɢ ɦɢɤɪɨɚɷɪɨɮɢɥɶɧɵɟ) ɩɚɥɨɱɤɢ ɢ ɤɨɤɤɢ.

Ƚɪɭɩɩɚ 5. Ɏɚɤɭɥɶɬɚɬɢɜɧɨ ɚɷɪɨɛɧɵɟ ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɩɚɥɨɱɤɢ. Ƚɪɭɩɩɚ 6. Ƚɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɚɧɚɷɪɨɛɧɵɟ ɩɪɹɦɵɟ, ɢɡɨɝɧɭɬɵɟ ɢɥɢ

ɫɩɢɪɚɥɟɜɢɞɧɵɟ ɩɚɥɨɱɤɢ.

Ƚɪɭɩɩɚ 7. Ȼɚɤɬɟɪɢɢ, ɨɫɭɳɟɫɬɜɥɹɸɳɢɟ ɞɢɫɫɢɦɢɥɹɰɢɨɧɧɨɟ ɜɨɫɫɬɚɧɨɜɥɟɧɢɟ ɫɟɪɵ ɢɥɢ ɫɭɥɶɮɚɬɚ.

Ƚɪɭɩɩɚ 8. Ⱥɧɚɷɪɨɛɧɵɟ ɝɪɚɦɨɬɪɢɰɚɬɟɥɶɧɵɟ ɤɨɤɤɢ. Ƚɪɭɩɩɚ 9. Ɋɢɤɤɟɬɫɢɢ ɢ ɯɥɚɦɢɞɢɢ.

Ƚɪɭɩɩɚ 10. Ⱥɧɨɤɫɢɝɟɧɧɵɟ ɮɨɬɨɬɪɨɮɧɵɟ ɛɚɤɬɟɪɢɢ. Ƚɪɭɩɩɚ 11. Ɉɤɫɢɝɟɧɧɵɟ ɮɨɬɨɬɪɨɮɧɵɟ ɛɚɤɬɟɪɢɢ.

Ƚɪɭɩɩɚ 12. Ⱥɷɪɨɛɧɵɟ ɯɟɦɨɥɢɬɨɬɪɨɮɧɵɟ ɛɚɤɬɟɪɢɢ ɢ ɛɥɢɡɤɢɟ ɨɪɝɚɧɢɡɦɵ.

Ƚɪɭɩɩɚ 13. ɉɨɱɤɭɸɳɢɟɫɹ ɢ (ɢɥɢ) ɨɛɪɚɡɭɸɳɢɟ ɜɵɪɨɫɬɵ ɛɚɤɬɟɪɢɢ. Ƚɪɭɩɩɚ 14. Ȼɚɤɬɟɪɢɢ, ɢɦɟɸɳɢɟ ɱɟɯɥɵ.

Ƚɪɭɩɩɚ 15. ɇɟɮɨɬɨɫɢɧɬɟɡɢɪɭɸɳɢɟ ɫɤɨɥɶɡɹɳɢɟ ɛɚɤɬɟɪɢɢ, ɧɟ ɨɛɪɚɡɭɸɳɢɟ ɩɥɨɞɨɜɵɯ ɬɟɥ.

Ƚɪɭɩɩɚ 16. ɋɤɨɥɶɡɹɳɢɟ ɛɚɤɬɟɪɢɢ, ɨɛɪɚɡɭɸɳɢɟ ɩɥɨɞɨɜɵɟ ɬɟɥɚ.

ȼ ɨɬɞɟɥ Firmicutes ɜɯɨɞɹɬ:

Ƚɪɭɩɩɚ 17. Ƚɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɟ ɤɨɤɤɢ.

Ƚɪɭɩɩɚ 18. Ƚɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɟ ɩɚɥɨɱɤɢ ɢ ɤɨɤɤɢ, ɨɛɪɚɡɭɸɳɢɟ ɷɧɞɨɫɩɨɪɵ.

Ƚɪɭɩɩɚ 19. Ƚɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɟ ɩɚɥɨɱɤɢ ɩɪɚɜɢɥɶɧɨɣ ɮɨɪɦɵ, ɧɟ ɨɛɪɚɡɭɸɳɢɟ ɫɩɨɪ.

Ƚɪɭɩɩɚ 20. Ƚɪɚɦɩɨɥɨɠɢɬɟɥɶɧɵɟ ɩɚɥɨɱɤɢ ɧɟɩɪɚɜɢɥɶɧɨɣ ɮɨɪɦɵ, ɧɟ ɨɛɪɚɡɭɸɳɢɟ ɫɩɨɪ.

Ƚɪɭɩɩɚ 21. Ɇɢɤɨɛɚɤɬɟɪɢɢ. Ƚɪɭɩɩɵ 22–29. Ⱥɤɬɢɧɨɦɢɰɟɬɵ.

Ʉ ɨɬɞɟɥɭ Tenericutes ɩɪɢɧɚɞɥɟɠɢɬ: Ƚɪɭɩɩɚ 30. Ɇɢɤɨɩɥɚɡɦɵ.

Ɉɬɞɟɥ Mendosicutes ɜɤɥɸɱɚɟɬ:

Ƚɪɭɩɩɚ 31. Ɇɟɬɚɧɨɝɟɧɵ.

Ƚɪɭɩɩɚ 32. ɋɭɥɶɮɚɬɪɟɞɭɰɢɪɭɸɳɢɟ ɚɪɯɟɛɚɤɬɟɪɢɢ.

Ƚɪɭɩɩɚ 33. ɗɤɫɬɪɟɦɚɥɶɧɨ ɝɚɥɨɮɢɥɶɧɵɟ ɚɪɯɟɛɚɤɬɟɪɢɢ (ɝɚɥɨɛɚɤɬɟɪɢɢ). Ƚɪɭɩɩɚ 34. Ⱥɪɯɟɛɚɤɬɟɪɢɢ, ɥɢɲɟɧɧɵɟ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ.

Ƚɪɭɩɩɚ 35. ɗɤɫɬɪɟɦɚɥɶɧɨ ɬɟɪɦɨɮɢɥɶɧɵɟ ɢ ɝɢɩɟɪɬɟɪɦɨɮɢɥɶɧɵɟ ɚɪɯɟɛɚɤɬɟɪɢɢ, ɦɟɬɚɛɨɥɢɡɢɪɭɸɳɢɟ ɫɟɪɭ.

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ȼ ɡɚɤɥɸɱɟɧɢɟ ɫɥɟɞɭɟɬ ɩɨɞɱɟɪɤɧɭɬɶ, ɱɬɨ ɛɨɥɶɲɢɧɫɬɜɨ ɦɢɤɪɨɨɪɝɚɧɢɡɦɨɜ, ɫɭɳɟɫɬɜɭɸɳɢɯ ɜ ɩɪɢɪɨɞɧɵɯ ɫɨɨɛɳɟɫɬɜɚɯ, ɟɳɟ ɞɨɥɠɧɨ ɛɵɬɶ ɜɵɞɟɥɟɧɨ ɜ ɱɢɫɬɵɟ ɤɭɥɶɬɭɪɵ. ɋɱɢɬɚɟɬɫɹ, ɱɬɨ ɜ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɤɭɥɶɬɢɜɢɪɨɜɚɬɶ ɦɨɠɧɨ ɬɨɥɶɤɨ 0,1 % ɜɫɟɝɨ ɦɢɤɪɨɛɧɨɝɨ ɪɚɡɧɨɨɛɪɚɡɢɹ, ɚ ɨɫɬɚɥɶɧɵɯ ɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɛɚɤɬɟɪɢɣ ɜɵɪɚɫɬɢɬɶ ɢ ɢɞɟɧɬɢɮɢɰɢɪɨɜɚɬɶ ɧɟ ɭɞɚɟɬɫɹ, ɯɨɬɹ ɭɠɟ ɜ ɱɢɫɬɭɸ ɤɭɥɶɬɭɪɭ ɜɵɞɟɥɟɧɵ ɢ ɨɩɢɫɚɧɵ ɨɤɨɥɨ 5 ɬɵɫ. ɜɢɞɨɜ ɩɪɨɤɚɪɢɨɬ.

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Ƚɥɚɜɚ 2. ɆɈɊɎɈɅɈȽɂə ɂ ɋɌɊɍɄɌɍɊɇȺə ɈɊȽȺɇɂɁȺɐɂə ȻȺɄɌȿɊɂȺɅɖɇɈɃ ɄɅȿɌɄɂ

2.1. Ɇɨɪɮɨɥɨɝɢɹ ɛɚɤɬɟɪɢɣ

Ⱦɥɹ ɛɚɤɬɟɪɢɣ ɯɚɪɚɤɬɟɪɧɵ ɤɥɟɬɤɢ ɬɪɟɯ ɨɫɧɨɜɧɵɯ ɮɨɪɦ: ɲɚɪɨɜɢɞɧɚɹ (ɫɮɟɪɢɱɟɫɤɚɹ), ɢɥɢ ɤɨɤɤɨɜɢɞɧɚɹ (ɨɬ ɝɪɟɱ. kokkos – ɡɟɪɧɨ), ɰɢɥɢɧɞɪɢɱɟɫɤɚɹ (ɩɚɥɨɱɤɨɜɢɞɧɚɹ) ɢ ɢɡɜɢɬɚɹ (ɪɢɫ. 3).

Ⱥ	Ȼ	ȼ
	Ɋɢɫ. 3. Ɇɢɤɪɨɮɨɬɨɝɪɚɮɢɢ ɤɥɟɬɨɤ ɛɚɤɬɟɪɢɣ:
	Ⱥ – ɫɮɟɪɢɱɟɫɤɢɯ; Ȼ – ɢɡɜɢɬɵɯ; ȼ – ɩɚɥɨɱɤɨɜɢɞɧɵɯ

Ʉɨɤɤɨɜɢɞɧɵɟ ɛɚɤɬɟɪɢɢ ɨɛɵɱɧɨ ɢɦɟɸɬ ɮɨɪɦɭ ɩɪɚɜɢɥɶɧɨɝɨ ɲɚɪɚ ɞɢɚɦɟɬɪɨɦ 1,0–2,0 ɦɤɦ, ɧɨ ɦɨɝɭɬ ɛɵɬɶ ɨɜɚɥɶɧɵɦɢ, ɷɥɥɢɩɫɨɢɞɧɵɦɢ, ɛɨɛɨɜɢɞɧɵɦɢ. Ʉɨɤɤɨɜɢɞɧɵɟ ɛɚɤɬɟɪɢɢ ɫɩɨɫɨɛɧɵ ɞɟɥɢɬɶɫɹ ɜ ɧɟɫɤɨɥɶɤɢɯ ɩɥɨɫɤɨɫɬɹɯ, ɩɪɢ ɷɬɨɦ ɩɨɫɥɟ ɞɟɥɟɧɢɹ ɤɥɟɬɤɢ ɦɨɝɭɬ ɧɟ ɪɚɫɯɨɞɢɬɶɫɹ ɢ ɮɨɪɦɢɪɨɜɚɬɶ ɪɚɡɥɢɱɧɨɝɨ ɜɢɞɚ ɫɤɨɩɥɟɧɢɹ (ɪɢɫ. 4).

ȿɫɥɢ ɞɟɥɟɧɢɟ ɤɨɤɤɨɜ ɩɪɨɢɫɯɨɞɢɬ ɜ ɨɞɧɨɣ ɩɥɨɫɤɨɫɬɢ, ɬɨ ɦɨɝɭɬ ɨɛɪɚɡɨɜɵɜɚɬɶɫɹ ɩɚɪɵ ɤɥɟɬɨɤ – ɞɢɩɥɨɤɨɤɤɢ (ɨɬ ɥɚɬ. diplos – ɞɜɨɣɧɨɣ) ɢ ɰɟɩɨɱɤɢ ɤɥɟɬɨɤ ɪɚɡɧɨɣ ɞɥɢɧɵ – ɫɬɪɟɩɬɨɤɨɤɤɢ (ɨɬ ɝɪɟɱ. streptos – ɰɟɩɨɱɤɚ). Ʉɨɤɤɢ, ɞɟɥɹɳɢɟɫɹ ɜ ɞɜɭɯ ɜɡɚɢɦɧɨ ɩɟɪɩɟɧɞɢɤɭɥɹɪɧɵɯ ɩɥɨɫɤɨɫɬɹɯ ɢ ɧɟ ɪɚɫɯɨɞɹɳɢɟɫɹ ɩɨɫɥɟ ɷɬɨɝɨ, ɨɛɪɚɡɭɸɬ ɬɟɬɪɚɞɵ ɤɨɤɤɨɜ (ɬɟɬɪɚɤɨɤɤɢ) (ɨɬ ɥɚɬ. tetra – ɱɟɬɵɪɟ). Ʉɨɝɞɚ ɞɟɥɟɧɢɟ ɤɥɟɬɨɤ ɩɪɨɢɫɯɨɞɢɬ ɜ ɬɪɟɯ ɜɡɚɢɦɧɨ ɩɟɪɩɟɧɞɢɤɭɥɹɪɧɵɯ ɩɥɨɫɤɨɫɬɹɯ, ɨɛɪɚɡɭɸɬɫɹ ɩɚɤɟɬɵ ɢɡ ɜɨɫɶɦɢ ɤɨɤɤɨɜ ɜ ɜɢɞɟ ɬɸɤɨɜ ɤɭɛɢɱɟɫɤɨɣ ɮɨɪɦɵ (ɫɚɪɰɢɧɵ) (ɨɬ ɥɚɬ. sarcina – ɫɜɹɡɤɚ, ɬɸɤ). ɍ ɧɟɤɨɬɨɪɵɯ ɜɢɞɨɜ ɛɚɤɬɟɪɢɣ ɩɪɢ ɞɟɥɟɧɢɢ ɤɨɤɤɨɜ ɜ ɧɟɫɤɨɥɶɤɢɯ ɩɥɨɫɤɨ-

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ɫɬɹɯ ɦɨɝɭɬ ɨɛɪɚɡɨɜɵɜɚɬɶɫɹ ɧɟɩɪɚɜɢɥɶɧɵɟ ɩɨ ɮɨɪɦɟ ɫɤɨɩɥɟɧɢɹ, ɧɚɩɨɦɢɧɚɸɳɢɟ ɝɪɨɡɞɶɹ ɜɢɧɨɝɪɚɞɚ – ɫɬɚɮɢɥɨɤɨɤɤɢ (ɨɬ ɥɚɬ. staphyle – ɝɪɨɡɞɶ ɜɢɧɨɝɪɚɞɚ).

ȺȻ

ȼ

Ƚ Ⱦ

Ɋɢɫ. 4. Ɍɢɩɵ ɫɤɨɩɥɟɧɢɣ ɤɨɤɤɨɜɢɞɧɵɯ ɤɥɟɬɨɤ:

Ⱥ – ɞɢɩɥɨɤɨɤɤɢ; Ȼ – ɫɬɪɟɩɬɨɤɨɤɤɢ; ȼ – ɬɟɬɪɚɤɨɤɤɢ; Ƚ – ɫɬɚɮɢɥɨɤɨɤɤɢ; Ⱦ – ɫɚɪɰɢɧɵ

ɉɚɥɨɱɤɨɜɢɞɧɵɟ (ɰɢɥɢɧɞɪɢɱɟɫɤɢɟ) ɤɥɟɬɤɢ ɫɢɥɶɧɨ ɪɚɡɥɢɱɚɸɬɫɹ ɩɨ ɨɬɧɨɲɟɧɢɸ ɞɥɢɧɵ ɤɥɟɬɤɢ ɤ ɟɟ ɩɨɩɟɪɟɱɧɢɤɭ. Ɉɧɢ ɞɟɥɹɬɫɹ ɬɨɥɶɤɨ ɜ ɨɞɧɨɣ ɩɥɨɫɤɨɫɬɢ – ɩɟɪɩɟɧɞɢɤɭɥɹɪɧɨ ɨɫɢ ɰɢɥɢɧɞɪɚ. Ʉɥɟɬɤɢ ɩɪɢ ɷɬɨɦ ɦɨɝɭɬ ɪɚɫɩɨɥɚɝɚɬɶɫɹ ɩɨɨɞɢɧɨɱɤɟ (ɦɨɧɨɛɚɤɬɟɪɢɢ), ɨɛɪɚɡɨɜɵɜɚɬɶ ɩɚɪɵ (ɞɢɩɥɨɛɚɤ-

ɬɟɪɢɢ) ɢɥɢ ɰɟɩɨɱɤɢ (ɫɬɪɟɩɬɨɛɚɤɬɟɪɢɢ) (ɪɢɫ. 5).

Ⱥ

Ȼ

ȼ

Ɋɢɫ. 5. Ɍɢɩɵ ɪɚɫɩɨɥɨɠɟɧɢɹ ɩɚɥɨɱɤɨɜɢɞɧɵɯ ɤɥɟɬɨɤ:

Ⱥ – ɦɨɧɨɛɚɤɬɟɪɢɢ; Ȼ – ɞɢɩɥɨɛɚɤɬɟɪɢɢ; ȼ – ɫɬɪɟɩɬɨɛɚɤɬɟɪɢɢ

ɂɡɜɢɬɵɟ ɤɥɟɬɤɢ ɦɨɝɭɬ ɢɦɟɬɶ ɪɚɡɧɨɟ ɱɢɫɥɨ ɡɚɜɢɬɤɨɜ. ȼ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɮɨɪɦɵ ɢ ɤɨɥɢɱɟɫɬɜɚ ɡɚɜɢɬɤɨɜ ɪɚɡɥɢɱɚɸɬ ɬɪɢ ɬɢɩɚ ɤɥɟɬɨɤ: ɜɢɛɪɢɨɧɵ (ɨɬ ɝɪɟɱ. vibrio – ɢɡɜɢɜɚɸɫɶ, ɢɡɝɢɛɚɸɫɶ) ɢɦɟɸɬ ɨɞɢɧ ɡɚɜɢɬɨɤ, ɧɟ ɩɪɟɜɵɲɚɸɳɢɣ ɱɟɬɜɟɪɬɢ ɨɛɨɪɨɬɚ ɫɩɢɪɚɥɢ (ɢɡɨɝɧɭɬɵɟ ɤɥɟɬɤɢ ɧɚɩɨɞɨɛɢɟ ɡɚɩɹɬɨɣ); ɫɩɢɪɢɥɥɵ (ɨɬ ɝɪɟɱ. speira – ɫɩɢɪɚɥɶ) ɢɦɟɸɬ 3–5 ɤɪɭɩɧɵɯ ɡɚɜɢɬɤɨɜ ɢ ɫɩɢɪɨɯɟɬɵ – ɛɨɥɶɲɨɟ ɤɨɥɢɱɟɫɬɜɨ ɦɟɥɤɢɯ ɡɚɜɢɬɤɨɜ (ɪɢɫ. 6).

36

Ⱥ

Ȼ

ȼ

Ɋɢɫ. 6. Ɍɢɩɵ ɢɡɜɢɬɵɯ ɤɥɟɬɨɤ:

Ⱥ – ɜɢɛɪɢɨɧɵ; Ȼ – ɫɩɢɪɢɥɥɵ; ȼ – ɫɩɢɪɨɯɟɬɵ

Ʉɪɨɦɟ ɨɩɢɫɚɧɧɵɯ ɮɨɪɦ, ɤɨɬɨɪɵɟ ɩɪɟɨɛɥɚɞɚɸɬ ɫɪɟɞɢ ɛɚɤɬɟɪɢɣ, ɢɡɜɟɫɬɧɵ ɢ ɤɥɟɬɤɢ ɢɧɵɯ ɬɢɩɨɜ: ɤɥɟɬɤɢ ɫ ɜɵɪɨɫɬɚɦɢ; ɛɭɥɚɜɨɜɢɞɧɨɣ ɮɨɪɦɵ; ɜɟɪɟɬɟɧɨɨɛɪɚɡɧɵɟ; ɥɚɧɰɟɬɨɜɢɞɧɵɟ; ɪɚɡɜɟɬɜɥɟɧɧɵɟ ɢ ɧɟɪɚɡɜɟɬɜɥɟɧɧɵɟ ɧɢɬɱɚɬɵɟ; ɢɦɟɸɳɢɟ ɜɢɞ ɤɨɥɶɰɚ, ɡɚɦɤɧɭɬɨɝɨ ɢɥɢ ɪɚɡɨɦɤɧɭɬɨɝɨ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɫɬɚɞɢɢ ɪɨɫɬɚ; ɜɟɬɜɹɳɢɟɫɹ; ɧɚɩɨɦɢɧɚɸɳɢɟ ɲɟɫɬɢɭɝɨɥɶɧɭɸ ɡɜɟɡɞɭ; ɩɥɚɫɬɢɧɤɨɨɛɪɚɡɧɵɟ; ɜ ɜɢɞɟ ɤɭɫɨɱɤɨɜ ɛɢɬɨɝɨ ɫɬɟɤɥɚ, ɤɜɚɞɪɚɬɢɤɨɜ ɢ ɬ. ɞ.

Ɏɨɪɦɚ ɤɥɟɬɨɤ ɛɨɥɶɲɢɧɫɬɜɚ ɛɚɤɬɟɪɢɣ ɹɜɥɹɟɬɫɹ ɭɫɬɨɣɱɢɜɵɦ ɜɢɞɨɜɵɦ ɩɪɢɡɧɚɤɨɦ. Ɉɞɧɚɤɨ ɫɭɳɟɫɬɜɭɸɬ ɛɚɤɬɟɪɢɢ, ɨɛɥɚɞɚɸɳɢɟ ɦɨɪɮɨɥɨɝɢɱɟɫɤɨɣ ɢɡɦɟɧɱɢɜɨɫɬɶɸ (ɩɥɟɨɦɨɪɮɢɡɦɨɦ), ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɭɫɥɨɜɢɣ ɢɦɟɸɳɢɟ ɜɢɞ ɩɚɥɨɱɟɤ, ɤɨɤɤɨɜ ɢɥɢ ɨɛɧɚɪɭɠɢɜɚɸɳɢɟ ɫɥɚɛɨɟ ɜɟɬɜɥɟɧɢɟ. ɍ ɧɟɤɨɬɨɪɵɯ ɜɢɞɨɜ ɛɚɤɬɟɪɢɣ ɩɪɢ ɩɪɨɯɨɠɞɟɧɢɢ ɰɢɤɥɚ ɪɚɡɜɢɬɢɹ ɬɚɤɠɟ ɧɚɛɥɸɞɚɟɬɫɹ ɢɡɦɟɧɟɧɢɟ ɮɨɪɦɵ ɤɥɟɬɨɤ.

2.2. ɋɬɪɭɤɬɭɪɧɚɹ ɨɪɝɚɧɢɡɚɰɢɹ ɛɚɤɬɟɪɢɚɥɶɧɨɣ ɤɥɟɬɤɢ

Ʉɥɟɬɤɚ ɩɪɨɤɚɪɢɨɬ, ɧɟɫɦɨɬɪɹ ɧɚ ɨɬɧɨɫɢɬɟɥɶɧɨ ɦɚɥɵɟ ɪɚɡɦɟɪɵ, ɢɦɟɟɬ ɜɫɟ ɨɫɧɨɜɧɵɟ ɫɬɪɭɤɬɭɪɧɵɟ ɤɨɦɩɨɧɟɧɬɵ, ɧɟɨɛɯɨɞɢɦɵɟ ɞɥɹ ɨɫɭɳɟɫɬɜɥɟɧɢɹ ɨɛɦɟɧɚ ɜɟɳɟɫɬɜ (ɪɢɫ. 7).

Ʉɚɤ ɢ ɥɸɛɚɹ ɞɪɭɝɚɹ, ɩɪɨɤɚɪɢɨɬɢɱɟɫɤɚɹ ɤɥɟɬɤɚ ɢɦɟɟɬ ɰɢɬɨɩɥɚɡɦɭ, ɤɨɬɨɪɚɹ ɨɤɪɭɠɟɧɚ ɰɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɨɣ ɦɟɦɛɪɚɧɨɣ. ɐɢɬɨɩɥɚɡɦɚ ɢ ɰɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɚɹ ɦɟɦɛɪɚɧɚ ɫɨɫɬɚɜɥɹɸɬ ɩɪɨɬɨɩɥɚɫɬ, ɫɧɚɪɭɠɢ ɨɬ ɧɟɝɨ ɪɚɫɩɨɥɨɠɟɧɵ ɩɨɜɟɪɯɧɨɫɬɧɵɟ ɫɬɪɭɤɬɭɪɵ. Ʉ ɢɯ ɱɢɫɥɭ ɨɬɧɨɫɹɬɫɹ ɤɥɟɬɨɱɧɚɹ ɫɬɟɧɤɚ, ɤɚɩɫɭɥɵ, ɱɟɯɥɵ, ɫɥɢɡɢɫɬɵɟ ɫɥɨɢ, ɠɝɭɬɢɤɢ, ɜɨɪɫɢɧɤɢ ɢ ɬ. ɞ.

2.2.1. Ʉɥɟɬɨɱɧɚɹ ɫɬɟɧɤɚ

Ʉɥɟɬɨɱɧɚɹ ɫɬɟɧɤɚ ɹɜɥɹɟɬɫɹ ɨɛɹɡɚɬɟɥɶɧɵɦ ɫɬɪɭɤɬɭɪɧɵɦ ɷɥɟɦɟɧɬɨɦ ɛɚɤɬɟɪɢɚɥɶɧɨɣ ɤɥɟɬɤɢ, ɢɫɤɥɸɱɟɧɢɟ ɫɨɫɬɚɜɥɹɸɬ ɦɢɤɨɩɥɚɡɦɵ ɢ L-ɮɨɪɦɵ. ɇɚ ɞɨɥɸ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ ɩɪɢɯɨɞɢɬɫɹ ɨɬ 5 ɞɨ 50 % ɫɭɯɢɯ ɜɟɳɟɫɬɜ ɤɥɟɬɤɢ.

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ȼɧɭɬɪɢɩɥɚɡɦɚɬɢɱɟɫɤɢɟ ɜɤɥɸɱɟɧɢɹ

Ɋɢɛɨɫɨɦɵ

ɀɝɭɬɢɤ

ɇɭɤɥɟɨɢɞ

Ʉɥɟɬɨɱɧɚɹ ɫɬɟɧɤɚ /

ɜɧɟɲɧɹɹ ɦɟɦɛɪɚɧɚ

ɐɢɬɨɩɥɚɡɦɚɬɢɱɟɫɤɚɹ

ɦɟɦɛɪɚɧɚ

ɐɢɬɨɩɥɚɡɦɚ

Ʉɚɩɫɭɥɚ

ȼɨɪɫɢɧɤɢ

Ɇɟɡɨɫɨɦɚ

Ɋɢɫ. 7. ɋɯɟɦɚɬɢɱɟɫɤɨɟ ɫɬɪɨɟɧɢɟ ɛɚɤɬɟɪɢɚɥɶɧɨɣ ɤɥɟɬɤɢ

ɉɨ ɫɬɪɨɟɧɢɸ ɢ ɯɢɦɢɱɟɫɤɨɦɭ ɫɨɫɬɚɜɭ ɤɥɟɬɨɱɧɚɹ ɫɬɟɧɤɚ ɩɪɨɤɚɪɢɨɬ ɨɬɥɢɱɚɟɬɫɹ ɨɬ ɬɚɤɨɜɨɣ ɷɭɤɚɪɢɨɬɢɱɟɫɤɢɯ ɨɪɝɚɧɢɡɦɨɜ. Ɉɫɧɨɜɧɵɦ ɤɨɦɩɨɧɟɧɬɨɦ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ ɛɨɥɶɲɢɧɫɬɜɚ ɛɚɤɬɟɪɢɣ ɹɜɥɹɟɬɫɹ ɦɭɪɟɢɧ, ɨɬɧɨɫɹɳɢɣɫɹ ɤ ɤɥɚɫɫɭ ɩɟɩɬɢɞɨɝɥɢɤɚɧɨɜ. Ɇɭɪɟɢɧ – ɝɟɬɟɪɨɩɨɥɢɦɟɪ, ɩɨɫɬɪɨɟɧɧɵɣ ɢɡ ɰɟɩɨɱɟɤ, ɜ ɤɨɬɨɪɵɯ ɱɟɪɟɞɭɸɬɫɹ ɨɫɬɚɬɤɢ N-ɚɰɟɬɢɥɝɥɸɤɨɡɚɦɢɧɚ ɢ N-ɚɰɟɬɢɥɦɭɪɚɦɨɜɨɣ ɤɢɫɥɨɬɵ, ɫɨɟɞɢɧɟɧɧɵɟ ɦɟɠɞɭ ɫɨɛɨɣ ȕ-1,4-ɝɥɢɤɨɡɢɞ- ɧɵɦɢ ɫɜɹɡɹɦɢ (ɪɢɫ. 8).

Ɍɚɤɢɟ ɧɟɪɚɡɜɟɬɜɥɟɧɧɵɟ ɝɟɬɟɪɨɩɨɥɢɦɟɪɧɵɟ ɰɟɩɢ ɨɛɪɚɡɭɸɬ ɨɫɧɨɜɭ ɦɭɪɟɢɧɚ. Ɉɫɬɚɬɤɢ N-ɚɰɟɬɢɥɦɭɪɚɦɨɜɨɣ ɤɢɫɥɨɬɵ ɱɟɪɟɡ ɥɚɤɬɢɥɶɧɵɟ ɝɪɭɩɩɵ ɫɨɟɞɢɧɟɧɵ ɩɟɩɬɢɞɧɨɣ ɫɜɹɡɶɸ ɫ ɚɦɢɧɨɤɢɫɥɨɬɚɦɢ. Ʉ ɬɢɩɢɱɧɵɦ ɚɦɢɧɨɤɢɫɥɨɬɚɦ, ɨɛɧɚɪɭɠɟɧɧɵɦ ɜ ɫɨɫɬɚɜɟ ɦɭɪɟɢɧɚ, ɨɬɧɨɫɹɬɫɹ L-ɚɥɚɧɢɧ, D-ɝɥɭɬɚ- ɦɢɧɨɜɚɹ ɤɢɫɥɨɬɚ, ɦɟɡɨ-ɞɢɚɦɢɧɨɩɢɦɟɥɢɧɨɜɚɹ ɤɢɫɥɨɬɚ ɢ D-ɚɥɚɧɢɧ.

Ⱦɢɚɦɢɧɨɩɢɦɟɥɢɧɨɜɚɹ ɤɢɫɥɨɬɚ ɨɬɧɨɫɢɬɫɹ ɤ ɞɢɚɦɢɧɨɤɢɫɥɨɬɚɦ ɢ ɧɚɯɨɞɢɬɫɹ ɜ ɦɭɪɟɢɧɟ ɜ ɦɟɡɨɮɨɪɦɟ:

HOOC	COOH
CH	CH2 CH2 CH2 HC
H2N	NH2

ɍ ɧɟɤɨɬɨɪɵɯ ɛɚɤɬɟɪɢɣ ɜɦɟɫɬɨ ɦɟɡɨ-ɞɢɚɦɢɧɨɩɢɦɟɥɢɧɨɜɨɣ ɤɢɫɥɨɬɵ ɜɫɬɪɟɱɚɸɬɫɹ L-ɥɢɡɢɧ, ɥɢɛɨ L- ɢɥɢ D-ɨɪɧɢɬɢɧ, ɥɢɛɨ 2,4-ɞɢɚɦɢɧɨɦɚɫɥɹ- ɧɚɹ ɤɢɫɥɨɬɚ, ɥɢɛɨ ɝɨɦɨɫɟɪɢɧ, ɥɢɛɨ ɝɢɞɪɨɤɫɢɥɢɡɢɧ.

38

N-ɚɰɟɬɢɥɝɥɸɤɨɡɚɦɢɧ	N-ɚɰɟɬɢɥɦɭɪɚɦɨɜɚɹ ɤɢɫɥɨɬɚ
CH2OH	CH2OH
O			O		O
OH	O

O

HNCOCH3

Ɉɫɬɚɬɨɤ ɦɨɥɨɱɧɨɣ ɤɢɫɥɨɬɵ

L-ɚɥɚɧɢɧ

D-ɝɥɭɬɚɦɢɧɨɜɚɹ ɤɢɫɥɨɬɚ

ɦɟɡɨ-Ⱦɢɚɦɢɧɨɩɢɦɟɥɢɧɨɜɚɹ ɤɢɫɥɨɬɚ

D-ɚɥɚɧɢɧ

O	HNCOCH3

H3C – CH – CO

«

NH

«

HC – CH3

«

CO

«

NH

«

HC – COOH

«

CH2

«

CH2

«

CO

«

NH NH2

««

HC– (CH2)3 – CH – COOH

«

CO

«

NH

«

HC – CH3

«

COOH

Ɋɢɫ. 8. ɋɬɪɭɤɬɭɪɚ ɫɭɛɴɟɞɢɧɢɰɵ ɦɭɪɟɢɧɚ ɤɥɟɬɨɱɧɨɣ ɫɬɟɧɤɢ ɛɚɤɬɟɪɢɣ

Escherichia coli (ɩɨ Ƚ. ɒɥɟɝɟɥɸ, 1987)

Ɇɟɡɨ-ɞɢɚɦɢɧɨɩɢɦɟɥɢɧɨɜɚɹ ɤɢɫɥɨɬɚ, L-ɥɢɡɢɧ ɢɥɢ ɞɪɭɝɢɟ ɞɢɚɦɢɧɨɤɢɫɥɨɬɵ ɢɝɪɚɸɬ ɛɨɥɶɲɭɸ ɪɨɥɶ ɜ ɮɨɪɦɢɪɨɜɚɧɢɢ ɦɟɠɦɨɥɟɤɭɥɹɪɧɵɯ ɫɲɢɜɨɤ,

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