2 курс / Нормальная физиология / Торшин 2014

Ƚɭɦɨɪɚɥɶɧɚɹ ɪɟɝɭɥɹɰɢɹ ɞɟɹɬɟɥɶɧɨɫɬɢ ɫɟɪɞɰɚ

ɇɚ ɪɚɛɨɬɭ ɫɟɪɞɰɚ ɩɪɟɠɞɟ ɜɫɟɝɨ ɜɥɢɹɸɬ ɦɟɞɢɚɬɨɪɵ ɚɰɟɬɢɥɯɨɥɢɧ, ɜɵɞɟɥɹɸɳɢɣɫɹ ɜ ɨɤɨɧɱɚɧɢɹɯ ɩɚɪɚɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ, ɨɧ ɬɨɪɦɨɡɢɬ ɞɟɹɬɟɥɶɧɨɫɬɶ ɫɟɪɞɰɚ, ɚ ɬɚɤɠɟ ɧɨɪɚɞɪɟɧɚɥɢɧ ɢ ɚɞɪɟɧɚɥɢɧ – ɦɟɞɢɚɬɨɪɵ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ, ɨɤɚɡɵɜɚɸɳɢɟ ɧɚ ɫɟɪɞɰɟ ɩɨɥɨɠɢɬɟɥɶɧɵɣ ɢɧɨ- ɢ ɯɪɨɧɨɬɪɨɩɧɵɣ ɷɮɮɟɤɬɵ (ɪɢɫ. 7.23). Ⱥɰɟɬɢɥɯɨɥɢɧ ɛɵɥ ɨɬɤɪɵɬ Ɉɬɬɨ Ʌɟɜɢ ɜ 1921 ɝ. ɧɚ ɢɡɨɥɢɪɨɜɚɧɧɵɯ ɫɟɪɞɰɚɯ ɥɹɝɭɲɟɤ. Ɋɚɡɞɪɚɠɚɹ ɜɚɝɨɫɢɦɩɚɬɢɱɟɫɤɢɣ ɫɬɜɨɥ ɨɞɧɨɝɨ ɫɟɪɞɰɚ, ɨɧ ɧɚɛɥɸɞɚɥ ɨɫɬɚɧɨɜɤɭ ɢ ɞɟɧɟɪɜɢɪɨɜɚɧɧɨɝɨ ɜɬɨɪɨɝɨ ɫɟɪɞɰɚ, ɫɜɹɡɚɧɧɨɝɨ ɫ ɩɟɪɜɵɦ ɨɛɳɢɦ ɩɟɪɮɭɡɢɪɭɸɳɢɦ ɪɚɫɬɜɨɪɨɦ, ɜ ɤɨɬɨɪɵɣ ɩɨɩɚɞɚɥɨ ɜɟɳɟɫɬɜɨ (vagus-stoff), ɜɵɞɟɥɟɧɧɨɟ ɜ ɨɤɨɧɱɚɧɢɹɯ ɫɬɢɦɭɥɢɪɭɟɦɨɝɨ ɛɥɭɠɞɚɸɳɟɝɨ ɧɟɪɜɚ ɩɟɪɜɨɝɨ ɫɟɪɞɰɚ (ɪɢɫ. 7.24).

Ɋɢɫ. 7.23. ȼɥɢɹɧɢɟ ɝɭɦɨɪɚɥɶɧɵɯ ɮɚɤɬɨɪɨɜ ɧɚ ɫɨɤɪɚɳɟɧɢɹ ɢɡɨɥɢɪɨɜɚɧɧɨɝɨ ɫɟɪɞɰɚ ɥɹɝɭɲɤɢ (ɩɨ ɒɬɪɚɭɛɭ):

ɚ– ɋɚCl2; ɛ – Ʉɋl; ɜ – ɚɞɪɟɧɚɥɢɧ; ɝ – ɚɰɟɬɢɥɯɨɥɢɧ; ɞ – ɨɬɦɟɬɤɚ ɜɪɟɦɟɧɢ

ɫɰɟɧɨɣ ɞɟɥɟɧɢɹ 2 ɫ. ɋɬɪɟɥɤɨɣ ɨɛɨɡɧɚɱɟɧ ɦɨɦɟɧɬ ɜɜɟɞɟɧɢɹ ɜɟɳɟɫɬɜɚ

ɜɤɚɧɸɥɸ ɫ ɪɚɫɬɜɨɪɨɦ Ɋɢɧɝɟɪɚ

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Ɋɢɫ. 7.24. Ɇɟɬɨɞɢɤɚ ɨɛɧɚɪɭɠɟɧɢɹ ɦɟɞɢɚɬɨɪɨɜ – ɨɩɵɬ Ɉɬɬɨ Ʌɟɜɢ (Ⱥ) ɢ ɨɩɵɬ Ⱥ.ȼ. Ʉɢɛɹɤɨɜɚ (Ȼ):

I– ɨɫɬɚɧɨɜɤɚ ɫɟɪɞɰɚ ɥɹɝɭɲɤɢ (Iɚ) ɩɪɢ ɪɚɡɞɪɚɠɟɧɢɢ ɛɥɭɠɞɚɸɳɟɝɨ ɧɟɪɜɚ; II – ɨɫɬɚɧɨɜɤɚ ɫɟɪɞɰɚ ɥɹɝɭɲɤɢ (II) ɛɟɡ ɪɚɡɞɪɚɠɟɧɢɹ ɛɥɭɠɞɚɸɳɟɝɨ ɧɟɪɜɚ

(II ɚ); 1 – ɞɜɭɪɨɝɚɹ ɤɚɧɸɥɹ; 2 – ɪɟɝɢɫɬɪɚɰɢɹ ɫɨɤɪɚɳɟɧɢɣ ɫɟɪɞɰɚ;

3 – ɫɬɢɦɭɥɹɬɨɪ; 4 – ɚɪɬɟɪɢɹ, ɨɬɞɚɸɳɚɹ ɜɟɬɨɱɤɢ ɜ ɫɢɦɩɚɬɢɱɟɫɤɢɣ ɝɚɧɝɥɢɣ; 5 – ɫɢɦɩɚɬɢɱɟɫɤɢɣ ɝɚɧɝɥɢɣ, 6 – ɜɟɧɚ, ɩɨ ɤɨɬɨɪɨɣ ɨɬɬɟɤɚɟɬ ɤɪɨɜɶ ɢɡ ɫɢɦɩɚɬɢɱɟɫɤɨɝɨ ɝɚɧɝɥɢɹ; ɫɬɢɦɭɥɹɬɨɪɨɦ ɪɚɡɞɪɚɠɚɟɬɫɹ ɫɢɦɩɚɬɢɱɟɫɤɢɣ ɫɬɜɨɥ;

7 – ɩɟɪɮɭɡɚɬ, ɜ ɤɨɬɨɪɨɦ ɫɨɞɟɪɠɢɬɫɹ ɦɟɞɢɚɬɨɪ

ȼ ɧɚɫɬɨɹɳɟɟ ɜɪɟɦɹ ɫɬɚɥɨ ɢɡɜɟɫɬɧɨ, ɱɬɨ ɩɪɢ ɪɚɡɞɪɚɠɟɧɢɢ ɧɟɪɜɨɜ, ɢɧɧɟɪɜɢɪɭɸɳɢɯ ɫɟɪɞɰɟ, ɜ ɫɢɧɚɩɬɢɱɟɫɤɭɸ ɳɟɥɶ, ɩɨɦɢɦɨ ɨɫɧɨɜɧɵɯ ɦɟɞɢɚɬɨɪɨɜ, ɜɵɞɟɥɹɸɬɫɹ ɢ ɞɪɭɝɢɟ ɛɢɨɥɨɝɢɱɟɫɤɢ ɚɤɬɢɜɧɵɟ ɜɟɳɟɫɬɜɚ, ɜ ɱɚɫɬɧɨɫɬɢ ɩɟɩɬɢɞɵ. Ɉɧɢ ɨɛɥɚɞɚɸɬ ɦɨɞɭɥɢɪɭɸɳɢɦ ɞɟɣɫɬɜɢɟɦ ɜ ɨɬɧɨɲɟɧɢɢ ɨɫɧɨɜɧɨɝɨ ɦɟɞɢɚɬɨɪɚ. Ɍɚɤ, ɨɩɢɨɢɞɧɵɟ ɩɟɩɬɢɞɵ (ɷɧɤɟɮɚɥɢɧɵ ɢ ɷɧɞɨɪɮɢɧɵ) ɭɝɧɟɬɚɸɬ ɷɮɮɟɤɬɵ ɪɚɡɞɪɚɠɟɧɢɹ ɛɥɭɠɞɚɸɳɟɝɨ ɧɟɪɜɚ, ɚ ɩɟɩɬɢɞ «ɞɟɥɶɬɚ–ɫɧɚ» ɭɫɢɥɢɜɚɟɬ ɜɚɝɭɫɧɭɸ ɛɪɚɞɢɤɚɪɞɢɸ. ɇɟɣɪɨɩɟɩɬɢɞ Y(NPY), ɜɵɞɟɥɹɸɳɢɣɫɹ ɜɦɟɫɬɟ ɫ ɧɨɪɚɞɪɟɧɚɥɢɧɨɦ ɜ ɨɤɨɧɱɚɧɢɹɯ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ, ɭɝɧɟɬɚɟɬ ɜɵɫɜɨɛɨɠɞɟɧɢɟ ɚɰɟɬɢɥɯɨɥɢɧɚ ɢɡ ɪɚɫɩɨɥɨɠɟɧɧɵɯ ɪɹɞɨɦ ɧɟɪɜɧɵɯ ɜɨɥɨɤɨɧ ɛɥɭɠɞɚɸɳɢɯ ɧɟɪɜɨɜ.

ɉɨɥɨɠɢɬɟɥɶɧɨɟ, ɩɨɞɨɛɧɨɟ ɚɞɪɟɧɚɥɢɧɭ, ɜɥɢɹɧɢɟ ɧɚ ɫɟɪɞɰɟ ɛɵɥɨ ɨɬɦɟɱɟɧɨ ɭ ɞɨɮɚɦɢɧɚ. Ʉɨɪɬɢɤɨɫɬɟɪɨɢɞɵ, ɚɧ-

ɝɢɨɬɟɧɡɢɧ, ɫɟɪɨɬɨɧɢɧ ɨɤɚɡɵɜɚɸɬ ɩɨɥɨɠɢɬɟɥɶɧɵɣ ɢɧɨɬɪɨɩɧɵɣ ɷɮɮɟɤɬ.

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Ƚɥɸɤɚɝɨɧ, ɚɤɬɢɜɢɪɭɹ ɚɞɟɧɢɥɚɬɰɢɤɥɚɡɭ, ɭɜɟɥɢɱɢɜɚɟɬ ɫɢɥɭ ɢ ɱɚɫɬɨɬɭ ɫɟɪɞɟɱɧɵɯ ɫɨɤɪɚɳɟɧɢɣ.

Ɍɢɪɨɤɫɢɧ ɢ ɬɪɢɣɨɞɬɢɪɨɧɢɧ ɨɤɚɡɵɜɚɸɬ ɩɪɟɢɦɭɳɟɫɬ-

ɜɟɧɧɨ ɩɨɥɨɠɢɬɟɥɶɧɵɣ ɯɪɨɧɨɬɪɨɩɧɵɣ ɷɮɮɟɤɬ, ɭɜɟɥɢɱɢɜɚɹ ɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɶ ɫɟɪɞɰɚ ɤ ɜɥɢɹɧɢɸ ɫɢɦɩɚɬɢɱɟɫɤɨɣ ɧɟɪɜɧɨɣ ɫɢɫɬɟɦɵ, ɜɵɡɵɜɚɹ ɬɚɯɢɤɚɪɞɢɸ ɢ ɪɚɡɥɢɱɧɨɝɨ ɬɢɩɚ ɚɪɢɬɦɢɢ.

Ⱥɞɟɧɨɡɢɧ ɪɚɫɲɢɪɹɟɬ ɤɨɪɨɧɚɪɧɵɟ ɫɨɫɭɞɵ, ɭɜɟɥɢɱɢɜɚɟɬ ɤɨɪɨɧɚɪɧɵɣ ɤɪɨɜɨɬɨɤ ɜ 6 ɪɚɡ, ɨɤɚɡɵɜɚɹ ɩɨɥɨɠɢɬɟɥɶɧɨɟ ɢɧɨɬɪɨɩɧɨɟ ɢ ɯɪɨɧɨɬɪɨɩɧɨɟ ɜɥɢɹɧɢɟ ɧɚ ɫɟɪɞɰɟ.

ɂɨɧɵ ɋɚ2+ ɭɜɟɥɢɱɢɜɚɸɬ ɫɢɥɭ ɫɨɤɪɚɳɟɧɢɣ ɢ ɩɨɜɵɲɚɸɬ ɜɨɡɛɭɞɢɦɨɫɬɶ ɫɟɪɞɟɱɧɨɣ ɦɵɲɰɵ ɡɚ ɫɱɟɬ ɚɤɬɢɜɚɰɢɢ ɮɨɫɮɨɪɢɥɚɡɵ. ɉɟɪɟɞɨɡɢɪɨɜɤɚ ɢɨɧɨɜ ɋɚ2+ ɜɵɡɵɜɚɟɬ ɨɫɬɚɧɨɜɤɭ ɫɟɪɞɰɚ ɜ ɫɢɫɬɨɥɟ.

ɇɨɪɦɚɥɶɧɚɹ ɤɨɧɰɟɧɬɪɚɰɢɹ ɢɨɧɨɜ Ʉ+ ɜ ɤɪɨɜɢ ɫɨɫɬɚɜɥɹɟɬ ɞɨ 4 ɦɦɨɥɶ/ɥ. ȿɫɥɢ ɭɜɟɥɢɱɢɬɶ ɤɨɧɰɟɧɬɪɚɰɢɸ Ʉ+ ɜ 2 ɪɚɡɚ, ɬɨ ɜɨɡɛɭɞɢɦɨɫɬɶ ɢ ɩɪɨɜɨɞɢɦɨɫɬɶ ɫɟɪɞɰɚ ɪɟɡɤɨ ɫɧɢɠɚɸɬɫɹ ɢ ɦɨɠɟɬ ɩɪɨɢɡɨɣɬɢ ɟɝɨ ɨɫɬɚɧɨɜɤɚ ɜ ɞɢɚɫɬɨɥɟ. ȿɫɥɢ ɢɨɧɨɜ Ʉ+ ɧɟɞɨɫɬɚɟɬ (ɝɢɩɨɤɚɥɢɟɦɢɹ), ɱɬɨ ɧɚɛɥɸɞɚɟɬɫɹ ɩɪɢ ɩɪɢɟɦɟ ɞɢɭɪɟɬɢɤɨɜ, ɤɨɬɨɪɵɟ ɜɵɜɨɞɹɬ ɜɦɟɫɬɟ ɫ ɜɨɞɨɣ ɢ Ʉ+, ɬɨ ɜɨɡɧɢɤɚɟɬ ɚɪɢɬɦɢɹ ɫɟɪɞɰɚ ɢ, ɜ ɱɚɫɬɧɨɫɬɢ, ɷɤɫɬɪɚɫɢɫɬɨɥɢɹ, ɩɨɷɬɨɦɭ ɨɞɧɨɜɪɟɦɟɧɧɨ ɫ ɞɢɭɪɟɬɢɤɚɦɢ ɧɟɨɛɯɨɞɢɦɨ ɩɪɢɧɢɦɚɬɶ ɩɪɟɩɚɪɚɬɵ, ɫɛɟɪɟɝɚɸɳɢɟ Ʉ+ (ɧɚɩɪɢɦɟɪ, ɩɚɧɚɧɝɢɧ).

ɉɪɟɞɫɟɪɞɢɹ ɜɵɪɚɛɚɬɵɜɚɸɬ ɚɬɪɢɨɩɟɩɬɢɞ, ɢɥɢ ɧɚɬɪɢɣɭɪɟɬɢɱɟɫɤɢɣ ɝɨɪɦɨɧ, ɜ ɨɬɜɟɬ ɧɚ ɪɚɫɬɹɠɟɧɢɟ ɢɯ ɫɬɟɧɨɤ. Ɉɧ ɪɚɫɫɥɚɛɥɹɟɬ ɝɥɚɞɤɨɦɵɲɟɱɧɵɟ ɤɥɟɬɤɢ ɦɟɥɤɢɯ ɫɨɫɭɞɨɜ, ɜɵɞɟɥɹɟɬ ɧɚɬɪɢɣ ɫ ɦɨɱɨɣ (ɧɚɬɪɢɣɭɪɟɡ) ɢ ɩɨɜɵɲɚɟɬ ɞɢɭɪɟɡ, ɭɦɟɧɶɲɚɟɬ ɨɛɴɟɦ ɰɢɪɤɭɥɢɪɭɸɳɟɣ ɤɪɨɜɢ, ɩɨɞɚɜɥɹɟɬ ɫɟɤɪɟɰɢɸ ɪɟɧɢɧɚ, ɬɨɪɦɨɡɢɬ ɷɮɮɟɤɬɵ ɚɧɝɢɨɬɟɧɡɢɧɚ II ɢ ɚɥɶɞɨɫɬɟɪɨɧɚ, ɫɧɢɠɚɟɬ ɚɪɬɟɪɢɚɥɶɧɨɟ ɞɚɜɥɟɧɢɟ.

Ɋɟɝɭɥɹɰɢɹ ɬɨɧɭɫɚ ɫɨɫɭɞɨɜ

Ɇɟɯɚɧɢɡɦɵ, ɪɟɝɭɥɢɪɭɸɳɢɟ ɫɨɫɭɞɢɫɬɵɣ ɬɨɧɭɫ, ɦɨɠɧɨ ɭɫɥɨɜɧɨ ɪɚɡɞɟɥɢɬɶ: 1) ɧɚ ɦɟɫɬɧɵɟ, ɩɟɪɢɮɟɪɢɱɟɫɤɢɟ, ɪɟɝɭɥɢɪɭɸɬ ɤɪɨɜɨɬɨɤ ɜ ɨɬɞɟɥɶɧɨɦ ɨɪɝɚɧɟ ɢɥɢ ɭɱɚɫɬɤɟ ɬɤɚɧɢ ɧɟɡɚɜɢɫɢɦɨ ɨɬ ɰɟɧɬɪɚɥɶɧɨɣ ɪɟɝɭɥɹɰɢɢ; 2) ɰɟɧɬɪɚɥɶɧɵɟ, ɩɨɞɞɟɪɠɢɜɚɸɳɢɟ ɭɪɨɜɟɧɶ ȺȾ ɢ ɫɢɫɬɟɦɧɨɟ ɤɪɨɜɨɨɛɪɚɳɟɧɢɟ; 3) ɝɭɦɨ-

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ɪɚɥɶɧɵɟ, ɪɟɝɭɥɢɪɭɸɳɢɟ ɩɪɨɫɜɟɬ ɫɨɫɭɞɚ ɡɚ ɫɱɟɬ ɯɢɦɢɱɟɫɤɢɯ, ɪɚɫɬɜɨɪɟɧɧɵɯ ɜ ɤɪɨɜɢ ɜɟɳɟɫɬɜ.

Ɇɟɫɬɧɵɟ ɪɟɝɭɥɹɬɨɪɧɵɟ ɦɟɯɚɧɢɡɦɵ

Ɉɧɢ ɪɟɚɥɢɡɭɸɬɫɹ ɭɠɟ ɧɚ ɭɪɨɜɧɟ ɷɧɞɨɬɟɥɢɹ ɫɨɫɭɞɨɜ, ɤɨɬɨɪɵɣ ɨɛɥɚɞɚɟɬ ɫɩɨɫɨɛɧɨɫɬɶɸ ɜɵɪɚɛɚɬɵɜɚɬɶ ɢ ɜɵɞɟɥɹɬɶ ɛɢɨɥɨɝɢɱɟɫɤɢ ɚɤɬɢɜɧɵɟ ɜɟɳɟɫɬɜɚ, ɜɵɡɵɜɚɸɳɢɟ ɪɚɫɫɥɚɛɥɟɧɢɟ ɢɥɢ ɫɨɤɪɚɳɟɧɢɟ ɝɥɚɞɤɢɯ ɦɵɲɰ ɫɨɫɭɞɨɜ ɜ ɨɬɜɟɬ ɧɚ ɩɨɜɵɲɟɧɢɟ ȺȾ, ɚ ɬɚɤɠɟ ɦɟɯɚɧɢɱɟɫɤɢɟ ɢɥɢ ɮɚɪɦɚɤɨɥɨɝɢɱɟɫɤɢɟ ɜɨɡɞɟɣɫɬɜɢɹ. ɋɯɟɦɚ ɪɚɛɨɬɵ ɦɟɫɬɧɵɯ ɦɟɯɚɧɢɡɦɨɜ ɪɟɝɭɥɹɰɢɢ ɫɨɫɭɞɢɫɬɨɝɨ ɬɨɧɭɫɚ ɩɪɟɞɫɬɚɜɥɟɧɚ ɧɚ ɪɢɫ. 7.25. ɗɧɞɨɬɟɥɢɣ ɫɨɫɭɞɚ ɪɚɫɫɦɚɬɪɢɜɚɟɬɫɹ ɤɚɤ ɷɧɞɨɤɪɢɧɧɚɹ ɠɟɥɟɡɚ, ɫɩɨɫɨɛɧɚɹ ɜɵɞɟɥɹɬɶ ɫɜɨɣ ɫɟɤɪɟɬ, ɤɨɬɨɪɵɣ ɡɚɬɟɦ ɞɟɣɫɬɜɭɟɬ ɧɚ ɝɥɚɞɤɭɸ ɦɵɲɰɭ ɫɟɪɞɰɚ ɢ ɢɡɦɟɧɹɟɬ ɟɟ ɬɨɧɭɫ. Ʉ ɜɟɳɟɫɬɜɚɦ, ɫɢɧɬɟɡɢɪɭɟɦɵɦ ɷɧɞɨɬɟɥɢɟɦ, ɨɬɧɨɫɢɬɫɹ ɜɚɡɨɚɤɬɢɜɧɵɣ ɷɧɞɨɬɟɥɢɚɥɶ-

ɧɵɣ ɮɚɤɬɨɪ ɪɚɫɫɥɚɛɥɟɧɢɹ (ȼɗɎɊ) – ɧɟɫɬɚɛɢɥɶɧɨɟ ɫɨɟɞɢɧɟ-

ɧɢɟ, ɨɞɧɢɦ ɢɡ ɤɨɬɨɪɵɯ ɦɨɠɟɬ ɛɵɬɶ ɨɤɫɢɞ ɚɡɨɬɚ (NO). ɉɪɢ ɞɟɣɫɬɜɢɢ ɧɚ ɷɧɞɨɬɟɥɢɣ ɫɨɫɭɞɨɜ ɬɚɤɢɯ ɝɭɦɨɪɚɥɶɧɵɯ ɮɚɤɬɨɪɨɜ, ɤɚɤ ɚɰɟɬɢɥɯɨɥɢɧ, ɛɪɚɞɢɤɢɧɢɧ, ɫɟɪɨɬɨɧɢɧ, ɜɟɳɟɫɬɜɨ Ɋ, ȺɌɎ ɩɪɨɞɭɰɢɪɭɟɬɫɹ NO ɤɨɬɨɪɵɣ, ɜ ɫɜɨɸ ɨɱɟɪɟɞɶ ɚɤɬɢɜɢɪɭɟɬ ɝɭɚɧɢɥɚɬɰɢɤɥɚɡɭ ɜ ɝɥɚɞɤɨɣ ɦɵɲɰɟ ɫɨɫɭɞɚ, ɩɪɢ ɷɬɨɦ ɭɜɟɥɢɱɢɜɚɟɬɫɹ ɤɨɧɰɟɧɬɪɚɰɢɹ ɰȽɆɎ, ɭɦɟɧɶɲɚɸɳɚɹ ɫɨɞɟɪɠɚɧɢɟ ɋɚ2+ ɜ ɰɢɬɨɡɨɥɟ ɢ ɜɵɡɵɜɚɸɳɚɹ ɜɚɡɨɞɢɥɹɬɚɰɢɸ.

Ⱦɪɭɝɨɟ ɜɟɳɟɫɬɜɨ – ɷɧɞɨɬɟɥɢɧ, ɜɚɡɨɤɨɧɫɬɪɢɤɬɨɪɧɵɣ ɩɟɩɬɢɞ, ɩɨɥɭɱɟɧɧɵɣ ɢɡ ɷɧɞɨɬɟɥɢɨɰɢɬɨɜ ɚɨɪɬɵ ɫɜɢɧɶɢ. Ɉɧ ɫɨɫɬɨɢɬ ɢɡ 21 ɚɦɢɧɨɤɢɫɥɨɬɧɨɝɨ ɨɫɬɚɬɤɚ, ɜɵɞɟɥɹɟɬɫɹ ɜ ɨɬɜɟɬ ɧɚ ɪɚɡɥɢɱɧɵɟ ɮɢɡɢɨɥɨɝɢɱɟɫɤɢɟ ɢ ɮɚɪɦɚɤɨɥɨɝɢɱɟɫɤɢɟ ɜɨɡɞɟɣɫɬɜɢɹ. ɍɜɟɥɢɱɟɧɢɟ ɫɢɧɬɟɡɚ ɷɧɞɨɬɟɥɢɧɚ ɦɨɠɟɬ ɩɪɢɜɟɫɬɢ ɤ ɬɚɤɢɦ ɩɚɬɨɥɨɝɢɱɟɫɤɢɦ ɫɨɫɬɨɹɧɢɹɦ, ɤɚɤ ɚɬɟɪɨɫɤɥɟɪɨɡ, ɫɟɪɞɟɱɧɚɹ ɢ ɩɨɱɟɱɧɚɹ ɧɟɞɨɫɬɚɬɨɱɧɨɫɬɶ.

ȿɫɥɢ ɩɨɥɧɨɫɬɶɸ ɞɟɧɟɪɜɢɪɨɜɚɬɶ ɫɨɫɭɞ, ɨɧ ɯɨɬɹ ɢ ɪɚɫɲɢɪɢɬɫɹ, ɧɨ ɛɭɞɟɬ ɫɨɯɪɚɧɹɬɶ ɧɟɤɨɬɨɪɨɟ ɧɚɩɪɹɠɟɧɢɟ ɫɜɨɟɣ ɫɬɟɧ-

ɤɢ ɡɚ ɫɱɟɬ ɛɚɡɚɥɶɧɨɝɨ, ɢɥɢ ɦɢɨɝɟɧɧɨɝɨ, ɬɨɧɭɫɚ ɝɥɚɞɤɢɯ ɦɵɲɰ. ɗɬɨɬ ɬɨɧɭɫ ɫɨɡɞɚɟɬɫɹ ɚɜɬɨɦɚɬɢɟɣ ɝɥɚɞɤɨɦɵɲɟɱɧɵɯ ɤɥɟɬɨɤ ɫɨɫɭɞɨɜ, ɤɨɬɨɪɵɟ ɢɦɟɸɬ ɧɟɫɬɚɛɢɥɶɧɨ ɩɨɥɹɪɢɡɨɜɚɧɧɭɸ ɦɟɦɛɪɚɧɭ, ɨɛɥɟɝɱɚɸɳɭɸ ɜɨɡɧɢɤɧɨɜɟɧɢɟ ɫɩɨɧɬɚɧɧɵɯ

408

ɉȾ ɜ ɷɬɢɯ ɤɥɟɬɤɚɯ. ɍɜɟɥɢɱɟɧɢɟ ȺȾ ɪɚɫɬɹɝɢɜɚɟɬ ɤɥɟɬɨɱɧɭɸ ɦɟɦɛɪɚɧɭ, ɱɬɨ ɭɜɟɥɢɱɢɜɚɟɬ ɫɩɨɧɬɚɧɧɭɸ ɚɤɬɢɜɧɨɫɬɶ ɝɥɚɞɤɢɯ ɦɵɲɰ ɢ ɩɪɢɜɨɞɢɬ ɤ ɭɜɟɥɢɱɟɧɢɸ ɢɯ ɬɨɧɭɫɚ. Ȼɚɡɚɥɶɧɵɣ ɬɨɧɭɫ ɨɫɨɛɟɧɧɨ ɜɵɪɚɠɟɧ ɜ ɫɨɫɭɞɚɯ ɦɢɤɪɨɰɢɪɤɭɥɹɬɨɪɧɨɝɨ ɪɭɫɥɚ, ɩɪɟɢɦɭɳɟɫɬɜɟɧɧɨ ɜ ɩɪɟɤɚɩɢɥɥɹɪɚɯ, ɨɛɥɚɞɚɸɳɢɯ ɚɜɬɨɦɚɬɢɟɣ. Ɉɧ ɩɨɞɞɟɪɠɢɜɚɟɬɫɹ ɤɚɤ ɡɚ ɫɱɟɬ ɯɢɦɢɱɟɫɤɨɣ ɢɧɮɨɪɦɚɰɢɢ ɨɬ ɷɧɞɨɬɟɥɢɹ ɫɨɫɭɞɢɫɬɨɣ ɫɬɟɧɤɢ ɩɪɢ ɟɟ ɪɚɫɬɹɠɟɧɢɢ, ɬɚɤ ɢ ɜ ɪɟɡɭɥɶɬɚɬɟ ɞɟɣɫɬɜɢɹ ɪɚɡɥɢɱɧɵɯ ɜɟɳɟɫɬɜ, ɪɚɫɬɜɨɪɟɧɧɵɯ ɜ ɤɪɨɜɢ. ɉɪɟɞɩɨɥɚɝɚɸɬ, ɱɬɨ ɪɚɫɬɹɠɟɧɢɟ ɫɨɫɭɞɢɫɬɨɣ ɫɬɟɧɤɢ ɩɪɢ ɭɜɟɥɢɱɟɧɢɢ ȺȾ ɩɨɜɵɲɚɟɬ ɤɨɧɰɟɧɬɪɚɰɢɸ ɜɧɭɬɪɢɤɥɟɬɨɱɧɨɝɨ ɋɚ2+, ɬɚɤ ɤɚɤ ɭɜɟɥɢɱɢɜɚɟɬɫɹ ɩɪɨɧɢɰɚɟɦɨɫɬɶ ɤɚɥɶɰɢɟɜɵɯ ɤɚɧɚɥɨɜ ɜ ɦɟɦɛɪɚɧɟ, ɚ ɷɬɨ ɜ ɫɜɨɸ ɨɱɟɪɟɞɶ ɩɪɢɜɨɞɢɬ ɤ ɫɨɤɪɚɳɟɧɢɸ ɫɨɫɭɞɚ.

Ɋɢɫ. 7.25. ɋɯɟɦɚ ɦɟɫɬɧɵɯ ɦɟɯɚɧɢɡɦɨɜ ɪɟɝɭɥɹɰɢɢ ɫɨɫɭɞɢɫɬɨɝɨ ɬɨɧɭɫɚ

Ʉɪɨɜɟɧɚɩɨɥɧɟɧɢɟ ɨɪɝɚɧɚ ɜ ɪɟɡɭɥɶɬɚɬɟ ɪɚɫɲɢɪɟɧɢɹ ɟɝɨ ɫɨɫɭɞɨɜ ɦɨɠɟɬ ɩɪɨɢɫɯɨɞɢɬɶ ɜ ɥɸɛɨɦ ɪɚɛɨɬɚɸɳɟɦ ɨɪɝɚɧɟ (ɩɨɩɟɪɟɱɧɨɩɨɥɨɫɚɬɨɣ ɦɵɲɰɟ, ɠɟɥɟɡɚɯ, ɫɟɪɞɰɟ, ɦɨɡɝɟ, ɝɥɚɞɤɢɯ ɦɵɲɰɚɯ ɠɟɥɭɞɨɱɧɨ-ɤɢɲɟɱɧɨɝɨ ɬɪɚɤɬɚ) ɜ ɪɟɡɭɥɶɬɚɬɟ ɧɚɤɨɩɥɟɧɢɹ ɩɪɨɞɭɤɬɨɜ ɦɟɬɚɛɨɥɢɡɦɚ (ɭɜɟɥɢɱɟɧɢɹ ɪɋɈ2, ɦɨɥɨɱɧɨɣ ɤɢɫɥɨɬɵ). Ɍɚɤɨɟ ɫɨɫɬɨɹɧɢɟ ɧɚɡɵɜɚɟɬɫɹ ɪɚɛɨɱɟɣ ɝɢɩɟɪɟ-

ɦɢɟɣ.

409

ɍɜɟɥɢɱɟɧɢɟ ɤɪɨɜɨɬɨɤɚ ɢ ɤɪɨɜɟɧɚɩɨɥɧɟɧɢɟ ɨɪɝɚɧɚ ɦɨɠɟɬ ɩɪɨɢɫɯɨɞɢɬɶ ɩɨɫɥɟ ɨɤɤɥɸɡɢɢ (ɩɟɪɟɠɚɬɢɹ) ɫɨɫɭɞɨɜ ɷɬɨɝɨ ɨɪɝɚɧɚ – ɷɬɨ ɪɟɚɤɬɢɜɧɚɹ ɝɢɩɟɪɟɦɢɹ. ɉɪɢ ɷɬɨɦ ɤɪɨɜɟɧɚɩɨɥɧɟɧɢɟ ɨɪɝɚɧɚ ɩɪɟɜɵɲɚɟɬ ɧɚ 50–70% ɬɨɬ, ɤɨɬɨɪɵɣ ɛɵɥ ɞɨ ɨɤɤɥɸɡɢɢ. ɇɚɤɨɩɥɟɧɢɟ ɩɪɨɞɭɤɬɨɜ ɦɟɬɚɛɨɥɢɡɦɚ ɜ ɩɟɪɟɠɚɬɨɦ ɫɨɫɭɞɟ ɢ ɟɝɨ ɡɧɚɱɢɬɟɥɶɧɨɟ ɪɚɫɲɢɪɟɧɢɟ ɩɨɫɥɟ ɫɧɹɬɢɹ ɨɤɤɥɸɡɢɢ ɫɜɢɞɟɬɟɥɶɫɬɜɭɸɬ ɨ ɫɭɳɟɫɬɜɨɜɚɧɢɢ ɦɟɫɬɧɵɯ ɦɟɬɚɛɨɥɢɱɟɫɤɢɯ ɦɟɯɚɧɢɡɦɨɜ ɪɟɝɭɥɹɰɢɢ ɤɪɨɜɨɬɨɤɚ ɜ ɬɤɚɧɹɯ.

ɐɟɧɬɪɚɥɶɧɵɟ ɦɟɯɚɧɢɡɦɵ ɪɟɝɭɥɹɰɢɢ

ɗɬɢ ɦɟɯɚɧɢɡɦɵ ɨɛɟɫɩɟɱɢɜɚɸɬɫɹ ɧɟɪɜɚɦɢ, ɢɧɧɟɪɜɢɪɭɸɳɢɦɢ ɫɨɫɭɞɢɫɬɭɸ ɫɬɟɧɤɭ, ɚ ɬɚɤɠɟ ɜɥɢɹɧɢɹɦɢ ɰɟɧɬɪɚɥɶɧɨɣ ɧɟɪɜɧɨɣ ɫɢɫɬɟɦɵ.

ȼɚɡɨɤɨɧɫɬɪɢɤɬɨɪɧɵɣ ɷɮɮɟɤɬ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ ɛɵɥ ɜɩɟɪɜɵɟ ɩɨɤɚɡɚɧ Ⱥ. ȼɚɥɶɬɟɪɨɦ (1842) ɧɚ ɩɥɚɜɚɬɟɥɶɧɨɣ ɩɟɪɟɩɨɧɤɟ ɥɹɝɭɲɤɢ, ɫɨɫɭɞɵ ɤɨɬɨɪɨɣ ɪɚɫɲɢɪɢɥɢɫɶ ɩɪɢ ɩɟɪɟɪɟɡɤɟ ɫɟɞɚɥɢɳɧɨɝɨ ɧɟɪɜɚ, ɫɨɞɟɪɠɚɳɟɝɨ ɜ ɫɟɛɟ ɫɢɦɩɚɬɢɱɟɫɤɢɟ ɜɨɥɨɤɧɚ, ɢ Ʉɥɨɞɨɦ Ȼɟɪɧɚɪɨɦ (1851), ɩɟɪɟɪɟɡɚɜɲɢɦ ɧɚ ɲɟɟ ɭ ɤɪɨɥɢɤɚ ɫ ɨɞɧɨɣ ɫɬɨɪɨɧɵ ɫɢɦɩɚɬɢɱɟɫɤɢɣ ɧɟɪɜ. ȼ ɪɟɡɭɥɶɬɚɬɟ ɫɨɫɭɞɵ ɭɯɚ ɧɚ ɫɬɨɪɨɧɟ ɩɟɪɟɪɟɡɤɢ ɧɟɪɜɚ ɪɚɫɲɢɪɢɥɢɫɶ, ɚ ɭɯɨ ɫɬɚɥɨ ɤɪɚɫɧɵɦ ɢ ɝɨɪɹɱɢɦ. Ɋɚɡɞɪɚɠɟɧɢɟ ɩɟɪɢɮɟɪɢɱɟɫɤɨɝɨ ɤɨɧɰɚ ɩɟɪɟɪɟɡɚɧɧɨɝɨ ɫɢɦɩɚɬɢɱɟɫɤɨɝɨ ɧɟɪɜɚ ɩɪɢɜɟɥɨ ɤ ɪɟɡɤɨɦɭ ɫɭɠɟɧɢɸ ɫɨɫɭɞɨɜ, ɚ ɭɯɨ ɫɬɚɥɨ ɛɥɟɞɧɵɦ ɢ ɯɨɥɨɞɧɵɦ.

ȼɚɡɨɤɨɧɫɬɪɢɤɬɨɪɧɵɟ ɜɨɥɨɤɧɚ ɫɢɦɩɚɬɢɱɟɫɤɨɣ ɧɟɪɜɧɨɣ ɫɢɫɬɟɦɵ ɢɧɧɟɪɜɢɪɭɸɬ ɜɫɟ ɚɪɬɟɪɢɢ, ɚɪɬɟɪɢɨɥɵ ɢ ɜɟɧɵ, ɤɪɨɦɟ ɤɚɩɢɥɥɹɪɨɜ. ɂɯ ɜɨɡɞɟɣɫɬɜɢɟ ɩɪɨɹɜɥɹɟɬɫɹ ɩɪɟɢɦɭɳɟɫɬɜɟɧɧɨ ɧɚ ɚɪɬɟɪɢɨɥɚɯ ɢ ɦɟɥɤɢɯ ɚɪɬɟɪɢɹɯ ɢ ɨɫɨɛɟɧɧɨ ɧɚ ɟɦɤɨɫɬɧɵɯ ɫɨɫɭɞɚɯ – ɜɟɧɚɯ, ɤɨɬɨɪɵɟ ɫɭɠɢɜɚɸɬɫɹ ɞɚɠɟ ɩɪɢ ɧɢɡɤɨɱɚɫɬɨɬɧɨɣ ɫɬɢɦɭɥɹɰɢɢ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ. ɉɪɢ ɪɚɡɞɪɚɠɟɧɢɢ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ ɩɪɨɢɫɯɨɞɢɬ ɡɧɚɱɢɬɟɥɶɧɨɟ ɫɭɠɟɧɢɟ ɫɨɫɭɞɨɜ ɤɨɠɢ, ɨɪɝɚɧɨɜ ɛɪɸɲɧɨɣ ɩɨɥɨɫɬɢ.

Ⱦɥɹ ɫɨɫɭɞɨɜ ɛɪɸɲɧɨɣ ɩɨɥɨɫɬɢ ɝɥɚɜɧɵɣ ɜɚɡɨɤɨɧɫɬɪɢɤɬɨɪ – ɷɬɨ ɱɪɟɜɧɵɣ ɧɟɪɜ, ɜ ɤɨɬɨɪɨɦ ɩɪɨɯɨɞɹɬ ɫɢɦɩɚɬɢɱɟɫɤɢɟ ɜɨɥɨɤɧɚ. Ɂɧɚɱɢɬ, ɫɢɦɩɚɬɢɱɟɫɤɢɣ ɧɟɪɜ – ɨɫɧɨɜɧɨɣ ɜɚɡɨɤɨɧɫɬɪɢɤɬɨɪ, ɩɨɞɞɟɪɠɢɜɚɸɳɢɣ ɬɨɧɭɫ ɫɨɫɭɞɨɜ ɧɚ ɬɨɦ ɢɥɢ ɢɧɨɦ ɭɪɨɜɧɟ ɜ ɡɚɜɢɫɢɦɨɫɬɢ ɨɬ ɤɨɥɢɱɟɫɬɜɚ ɢɦɩɭɥɶɫɨɜ, ɩɨɫɬɭɩɚɸ-

410

ɳɢɯ ɩɨ ɟɝɨ ɜɨɥɨɤɧɚɦ ɤ ɫɨɫɭɞɭ. ɋɜɨɟ ɜɥɢɹɧɢɟ ɧɚ ɫɨɫɭɞɵ ɫɢɦɩɚɬɢɱɟɫɤɢɣ ɧɟɪɜ ɨɤɚɡɵɜɚɟɬ ɱɟɪɟɡ ɧɨɪɚɞɪɟɧɚɥɢɧ, ɜɵɞɟɥɹɸɳɢɣɫɹ ɜ ɟɝɨ ɨɤɨɧɱɚɧɢɹɯ, ɢ ɚɥɶɮɚ-ɚɞɪɟɧɨɪɟɰɟɩɬɨɪɵ, ɪɚɫɩɨɥɨɠɟɧɧɵɟ ɜ ɫɨɫɭɞɢɫɬɵɯ ɫɬɟɧɤɚɯ, ɜ ɪɟɡɭɥɶɬɚɬɟ ɩɪɨɢɫɯɨɞɢɬ ɫɭɠɟɧɢɟ ɫɨɫɭɞɨɜ, ɜ ɤɨɬɨɪɵɯ ɩɪɟɨɛɥɚɞɚɸɬ Į1-ɚɞɪɟɧɨ- ɪɟɰɟɩɬɨɪɵ, ɪɟɚɝɢɪɭɸɳɢɟ ɜ ɨɫɧɨɜɧɨɦ ɫ ɧɨɪɚɞɪɟɧɚɥɢɧɨɦ, ɜ ɬɨ ɜɪɟɦɹ ɤɚɤ ɜ ɫɨɫɭɞɚɯ ɫɟɪɞɰɚ, ɥɟɝɤɢɯ, ɦɨɡɝɚ ɢ ɫɤɟɥɟɬɧɵɯ ɦɵɲɰ ɩɪɟɨɛɥɚɞɚɸɬ ɧɟ D, ɚ E-ɚɞɪɟɧɨɪɟɰɟɩɬɨɪɵ, ɤɨɧɬɚɤɬɢɪɭɸɳɢɟ ɜ ɨɫɧɨɜɧɨɦ ɫ ɚɞɪɟɧɚɥɢɧɨɦ. E-ɚɞɪɟɧɨɪɟɰɟɩɬɨɪɵ ɨɛɥɚɞɚɸɬ ɛɨɥɶɲɟɣ ɜɨɡɛɭɞɢɦɨɫɬɶɸ ɢ ɪɟɚɝɢɪɭɸɬ ɧɚ ɫɥɚɛɵɟ ɪɚɡɞɪɚɠɟɧɢɹ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ, ɜ ɪɟɡɭɥɶɬɚɬɟ ɩɪɨɢɫɯɨɞɢɬ ɪɚɫɲɢɪɟɧɢɟ ɫɨɫɭɞɨɜ.

ɑɬɨ ɤɚɫɚɟɬɫɹ ɜɚɡɨɞɢɥɹɬɚɰɢɢ (ɪɚɫɲɢɪɟɧɢɹ ɤɪɨɜɟɧɨɫɧɵɯ ɫɨɫɭɞɨɜ), ɬɨ ɨɧɚ ɦɨɠɟɬ ɩɪɨɢɫɯɨɞɢɬɶ ɩɪɟɠɞɟ ɜɫɟɝɨ ɡɚ ɫɱɟɬ ɫɧɢɠɟɧɢɹ ɬɨɧɭɫɚ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɜɚɡɨɤɨɧɫɬɪɢɤɬɨɪɧɵɯ ɧɟɪɜɧɵɯ ɜɨɥɨɤɨɧ – ɷɬɨ ɝɥɚɜɧɵɣ ɜɚɡɨɞɢɥɹɬɚɬɨɪɧɵɣ ɦɟɯɚɧɢɡɦ.

ȿɫɥɢ ɜɚɡɨɤɨɧɫɬɪɢɤɬɨɪɧɵɣ ɷɮɮɟɤɬ ɫɢɦɩɚɬɢɱɟɫɤɨɣ ɧɟɪɜɧɨɣ ɫɢɫɬɟɦɵ ɧɨɫɢɬ ɨɛɳɢɣ ɫɢɫɬɟɦɧɵɣ ɯɚɪɚɤɬɟɪ, ɬɨ ɜɚɡɨɞɢɥɹɬɚɬɨɪɧɵɣ ɹɜɥɹɟɬɫɹ ɱɚɳɟ ɦɟɫɬɧɨɣ ɪɟɚɤɰɢɟɣ. ɇɟɥɶɡɹ ɭɬɜɟɪɠɞɚɬɶ, ɱɬɨ ɩɚɪɚɫɢɦɩɚɬɢɱɟɫɤɚɹ ɧɟɪɜɧɚɹ ɫɢɫɬɟɦɚ ɪɚɫɲɢɪɹɟɬ ɜɫɟ ɫɨɫɭɞɵ. ɂɡɜɟɫɬɧɵ ɥɢɲɶ ɧɟɫɤɨɥɶɤɨ ɩɚɪɚɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ, ɪɚɫɲɢɪɹɸɳɢɯ ɫɨɫɭɞɵ ɬɨɥɶɤɨ ɬɟɯ ɨɪɝɚɧɨɜ, ɤɨɬɨɪɵɟ ɨɧɢ ɢɧɧɟɪɜɢɪɭɸɬ. Ɍɚɤ, ɪɚɡɞɪɚɠɟɧɢɟ ɛɚɪɚɛɚɧɧɨɣ ɫɬɪɭɧɵ – ɜɟɬɨɱɤɢ ɩɚɪɚɫɢɦɩɚɬɢɱɟɫɤɨɝɨ ɥɢɰɟɜɨɝɨ ɧɟɪɜɚ – ɪɚɫɲɢɪɹɟɬ ɫɨɫɭɞɵ ɩɨɞɱɟɥɸɫɬɧɨɣ ɠɟɥɟɡɵ ɢ ɭɜɟɥɢɱɢɜɚɟɬ ɜ ɧɟɣ ɤɪɨɜɨɬɨɤ.

ȼɚɡɨɞɢɥɹɬɚɬɨɪɧɵɣ ɷɮɮɟɤɬ ɛɵɥ ɩɨɥɭɱɟɧ ɩɪɢ ɪɚɡɞɪɚɠɟɧɢɢ ɞɪɭɝɢɯ ɩɚɪɚɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ: ɹɡɵɤɨɝɥɨɬɨɱɧɨɝɨ, ɪɚɫɲɢɪɹɸɳɟɝɨ ɫɨɫɭɞɵ ɦɢɧɞɚɥɢɧ, ɨɤɨɥɨɭɲɧɨɣ ɠɟɥɟɡɵ, ɡɚɞɧɟɣ ɬɪɟɬɢ ɹɡɵɤɚ; ɜɟɪɯɧɟɝɨɪɬɚɧɧɨɝɨ ɧɟɪɜɚ – ɜɟɬɨɱɤɢ ɛɥɭɠɞɚɸɳɟɝɨ ɧɟɪɜɚ, ɪɚɫɲɢɪɹɸɳɟɝɨ ɫɨɫɭɞɵ ɫɥɢɡɢɫɬɨɣ ɝɨɪɬɚɧɢ ɢ ɳɢɬɨɜɢɞɧɨɣ ɠɟɥɟɡɵ; ɬɚɡɨɜɨɝɨ ɧɟɪɜɚ, ɪɚɫɲɢɪɹɸɳɟɝɨ ɫɨɫɭɞɵ ɨɪɝɚɧɨɜ ɦɚɥɨɝɨ ɬɚɡɚ. ȼ ɨɤɨɧɱɚɧɢɹɯ ɜɵɲɟɩɟɪɟɱɢɫɥɟɧɧɵɯ ɧɟɪɜɨɜ ɜɵɞɟɥɹɥɫɹ ɦɟɞɢɚɬɨɪ ɚɰɟɬɢɥɯɨɥɢɧ (ɯɨɥɢɧɟɪɝɢɱɟɫɤɢɟ ɜɨɥɨɤɧɚ), ɤɨɬɨɪɵɣ ɤɨɧɬɚɤɬɢɪɨɜɚɥ ɫ Ɇ-ɯɨɥɢɧɨɪɟɰɟɩɬɨɪɚɦɢ ɢ ɜɵɡɵɜɚɥ ɪɚɫɲɢɪɟɧɢɟ ɫɨɫɭɞɨɜ.

ɋɪɟɞɢ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɜɨɥɨɤɨɧ ɟɫɬɶ ɯɨɥɢɧɟɪɝɢɱɟɫɤɢɟ, ɜ ɨɤɨɧɱɚɧɢɹɯ ɤɨɬɨɪɵɯ ɜɵɞɟɥɹɟɬɫɹ ɧɟ ɧɨɪɚɞɪɟɧɚɥɢɧ, ɚ ɚɰɟɬɢɥ-

411

ɯɨɥɢɧ, ɢɯ ɪɚɡɞɪɚɠɟɧɢɟ ɜɵɡɵɜɚɟɬ ɧɟ ɫɭɠɟɧɢɟ, ɚ ɪɚɫɲɢɪɟɧɢɟ ɫɨɫɭɞɨɜ ɬɚɤɢɯ ɨɪɝɚɧɨɜ, ɤɚɤ ɫɟɪɞɰɟ ɢ ɫɤɟɥɟɬɧɵɟ ɦɵɲɰɵ, ɷɮɮɟɤɬ ɨɬ ɪɚɡɞɪɚɠɟɧɢɹ ɛɥɨɤɢɪɭɟɬɫɹ ɚɬɪɨɩɢɧɨɦ.

Ɋɢɫ. 7.26. ɋɯɟɦɚ ɚɤɫɨɧ-ɪɟɮɥɟɤɫɚ:

1 – ɪɚɡɞɪɚɠɟɧɢɟ ɤɨɠɢ; 2 – ɪɚɫɲɢɪɟɧɢɟ ɫɨɫɭɞɨɜ; 3 – ɚɤɫɨɧ

ɋɬɢɦɭɥɹɰɢɹ ɡɚɞɧɢɯ ɤɨɪɟɲɤɨɜ ɫɩɢɧɧɨɦɨɡɝɨɜɵɯ ɧɟɪɜɨɜ

ɜɷɤɫɩɟɪɢɦɟɧɬɟ ɩɪɢɜɨɞɢɬ ɤ ɪɚɫɲɢɪɟɧɢɸ ɫɨɫɭɞɨɜ ɞɚɧɧɨɝɨ ɫɟɝɦɟɧɬɚ ɬɟɥɚ. Ɋɚɡɞɪɚɠɚɹ ɤɨɠɭ, ɧɚɩɪɢɦɟɪ, ɝɨɪɱɢɱɧɢɤɚɦɢ, ɦɨɠɧɨ ɩɨɥɭɱɢɬɶ ɦɟɫɬɧɨɟ ɪɚɫɲɢɪɟɧɢɟ ɫɨɫɭɞɨɜ ɢ ɩɨɤɪɚɫɧɟɧɢɟ ɞɚɧɧɨɝɨ ɭɱɚɫɬɤɚ ɤɨɠɢ ɩɨ ɬɢɩɭ ɚɤɫɨɧ-ɪɟɮɥɟɤɫɚ, ɪɟɚɥɢɡɭɟɦɨɝɨ

ɜɩɪɟɞɟɥɚɯ ɞɜɭɯ ɪɚɡɜɟɬɜɥɟɧɢɣ ɨɞɧɨɝɨ ɚɤɫɨɧɚ ɢ ɛɟɡ ɭɱɚɫɬɢɹ ɰɟɧɬɪɚɥɶɧɨɣ ɧɟɪɜɧɨɣ ɫɢɫɬɟɦɵ (ɪɢɫ. 7.26).

Ƚɭɦɨɪɚɥɶɧɚɹ ɪɟɝɭɥɹɰɢɹ ɫɨɫɭɞɢɫɬɨɝɨ ɬɨɧɭɫɚ

Ʉɜɟɳɟɫɬɜɚɦ, ɪɟɝɭɥɢɪɭɸɳɢɦ ɬɨɧɭɫ ɫɨɫɭɞɨɜ, ɨɬɧɨɫɹɬɫɹ

ɝɨɪɦɨɧɵ ɨɛɳɟɝɨ ɞɟɣɫɬɜɢɹ, ɦɟɫɬɧɵɟ ɝɨɪɦɨɧɵ, ɦɟɞɢɚɬɨɪɵ ɢ ɩɪɨɞɭɤɬɵ ɦɟɬɚɛɨɥɢɡɦɚ. ɂɯ ɦɨɠɧɨ ɪɚɡɞɟɥɢɬɶ ɧɚ ɞɜɟ ɝɪɭɩɩɵ:

ɫɨɫɭɞɨɫɭɠɢɜɚɸɳɢɟ ɢ ɫɨɫɭɞɨɪɚɫɲɢɪɹɸɳɢɟ.

Ʉɫɨɫɭɞɨɫɭɠɢɜɚɸɳɢɦ ɜɟɳɟɫɬɜɚɦ ɨɬɧɨɫɹɬɫɹ: ɝɨɪɦɨɧɵ ɦɨɡɝɨɜɨɝɨ ɫɥɨɹ ɧɚɞɩɨɱɟɱɧɢɤɨɜ – ɚɞɪɟɧɚɥɢɧ ɢ ɧɨɪɚɞɪɟɧɚɥɢɧ.

412

Ⱥɞɪɟɧɚɥɢɧ ɜ ɦɚɥɵɯ ɞɨɡɚɯ (1 u 10-7 ɝ/ɦɥ) ɩɨɜɵɲɚɟɬ ȺȾ, ɫɭɠɢɜɚɹ ɫɨɫɭɞɵ ɜɫɟɯ ɨɪɝɚɧɨɜ, ɤɪɨɦɟ ɫɨɫɭɞɨɜ ɫɟɪɞɰɚ, ɦɨɡɝɚ, ɩɨɩɟɪɟɱɧɨɩɨɥɨɫɚɬɨɣ ɦɭɫɤɭɥɚɬɭɪɵ, ɜ ɤɨɬɨɪɵɯ ɧɚɯɨɞɹɬɫɹ ɛɟɬɚɚɞɪɟɧɨɪɟɰɟɩɬɨɪɵ. ɇɨɪɚɞɪɟɧɚɥɢɧ – ɫɢɥɶɧɵɣ ɜɚɡɨɤɨɧɫɬɪɢɤɬɨɪ, ɜɡɚɢɦɨɞɟɣɫɬɜɭɸɳɢɣ ɫ ɚɥɶɮɚ-ɚɞɪɟɧɨɪɟɰɟɩɬɨɪɚɦɢ.

ȼɚɡɨɩɪɟɫɫɢɧ, ɢɥɢ ɚɧɬɢɞɢɭɪɟɬɢɱɟɫɤɢɣ ɝɨɪɦɨɧ, – ɝɨɪɦɨɧ ɡɚɞɧɟɣ ɞɨɥɢ ɝɢɩɨɮɢɡɚ, ɫɭɠɢɜɚɟɬ ɦɟɥɤɢɟ ɫɨɫɭɞɵ ɢ, ɜ ɱɚɫɬɧɨɫɬɢ, ɚɪɬɟɪɢɨɥɵ, ɨɫɨɛɟɧɧɨ ɩɪɢ ɡɧɚɱɢɬɟɥɶɧɨɦ ɩɚɞɟɧɢɢ ɚɪɬɟɪɢɚɥɶɧɨɝɨ ɞɚɜɥɟɧɢɹ.

Ⱥɥɶɞɨɫɬɟɪɨɧ – ɦɢɧɟɪɚɥɨɤɨɪɬɢɤɨɢɞ – ɝɨɪɦɨɧ ɤɨɪɵ ɧɚɞɩɨɱɟɱɧɢɤɨɜ, ɩɨɜɵɲɚɟɬ ɱɭɜɫɬɜɢɬɟɥɶɧɨɫɬɶ ɝɥɚɞɤɢɯ ɦɵɲɰ ɫɨɫɭɞɨɜ ɤ ɜɚɡɨɤɨɧɫɬɪɢɤɬɨɪɧɵɦ ɚɝɟɧɬɚɦ, ɭɫɢɥɢɜɚɟɬ ɩɪɟɫɫɨɪɧɨɟ ɞɟɣɫɬɜɢɟ ɚɧɝɢɨɬɟɧɡɢɧɚ II.

ɋɟɪɨɬɨɧɢɧ – ɨɛɪɚɡɭɟɬɫɹ ɜ ɫɥɢɡɢɫɬɨɣ ɤɢɲɟɱɧɢɤɚ ɢ ɜ ɧɟɤɨɬɨɪɵɯ ɨɬɞɟɥɚɯ ɝɨɥɨɜɧɨɝɨ ɦɨɡɝɚ, ɫɨɞɟɪɠɢɬɫɹ ɜ ɬɪɨɦɛɨɰɢɬɚɯ, ɫɭɠɢɜɚɟɬ ɩɨɜɪɟɠɞɟɧɧɵɣ ɫɨɫɭɞ ɢ ɩɪɟɩɹɬɫɬɜɭɟɬ ɤɪɨɜɨɬɟɱɟɧɢɸ. Ɉɧ ɨɤɚɡɵɜɚɟɬ ɦɨɳɧɨɟ ɫɨɫɭɞɨɫɭɠɢɜɚɸɳɟɟ ɜɥɢɹɧɢɟ ɧɚ ɚɪɬɟɪɢɢ ɦɹɝɤɨɣ ɦɨɡɝɨɜɨɣ ɨɛɨɥɨɱɤɢ ɢ ɦɨɠɟɬ ɢɝɪɚɬɶ ɪɨɥɶ ɜ ɜɨɡɧɢɤɧɨɜɟɧɢɢ ɫɩɚɡɦɨɜ (ɩɪɢɫɬɭɩɵ ɦɢɝɪɟɧɢ).

Ɋɟɧɢɧ – ɨɛɪɚɡɭɟɬɫɹ ɜ ɸɤɫɬɚɝɥɨɦɟɪɭɥɹɪɧɨɦ ɤɨɦɩɥɟɤɫɟ ɩɨɱɤɢ, ɨɫɨɛɟɧɧɨ ɜ ɛɨɥɶɲɨɦ ɤɨɥɢɱɟɫɬɜɟ ɩɪɢ ɟɟ ɢɲɟɦɢɢ. Ɉɧ ɪɚɫɳɟɩɥɹɟɬ ɚɥɶɮɚ-2-ɝɥɨɛɭɥɢɧ ɩɥɚɡɦɵ – ɚɧɝɢɨɬɟɧɡɢɧɨɝɟɧ ɢ ɩɪɟɜɪɚɳɚɟɬ ɟɝɨ ɜ ɦɚɥɨɚɤɬɢɜɧɵɣ ɞɟɤɚɩɟɩɬɢɞ – ɚɧɝɢɨɬɟɧɡɢɧ I, ɤɨɬɨɪɵɣ ɩɨɞ ɜɥɢɹɧɢɟɦ ɮɟɪɦɟɧɬɚ ɞɢɩɟɩɬɢɞɤɚɪɛɨɤɫɢɩɟɩɬɢɞɚɡɵ ɩɪɟɜɪɚɳɚɟɬɫɹ ɜ ɨɱɟɧɶ ɚɤɬɢɜɧɨɟ ɫɨɫɭɞɨɫɭɠɢɜɚɸɳɟɟ ɜɟɳɟɫɬɜɨ – ɚɧɝɢɨɬɟɧɡɢɧ II, ɩɨɜɵɲɚɸɳɟɟ ȺȾ (ɩɨɱɟɱɧɚɹ ɝɢɩɟɪɬɨɧɢɹ). Ⱥɧɝɢɨɬɟɧɡɢɧ II – ɦɨɳɧɵɣ ɫɬɢɦɭɥɹɬɨɪ ɜɵɪɚɛɨɬɤɢ ɚɥɶɞɨɫɬɟɪɨɧɚ, ɭɜɟɥɢɱɢɜɚɸɳɟɝɨ ɫɨɞɟɪɠɚɧɢɟ ɜ ɨɪɝɚɧɢɡɦɟ Nɚ+ ɢ ɜɧɟɤɥɟɬɨɱɧɨɣ ɠɢɞɤɨɫɬɢ. ȼ ɬɚɤɢɯ ɫɥɭɱɚɹɯ ɝɨɜɨɪɹɬ ɨ ɪɚɛɨɬɟ

ɪɟɧɢɧ–ɚɧɝɢɨɬɟɧɡɢɧ–ɚɥɶɞɨɫɬɟɪɨɧɨɜɨɣ ɫɢɫɬɟɦɵ ɢɥɢ ɦɟɯɚ-

ɧɢɡɦɚ. ɉɨɫɥɟɞɧɢɣ ɢɦɟɟɬ ɛɨɥɶɲɨɟ ɡɧɚɱɟɧɢɟ ɞɥɹ ɧɨɪɦɚɥɢɡɚɰɢɢ ɭɪɨɜɧɹ ɤɪɨɜɹɧɨɝɨ ɞɚɜɥɟɧɢɹ ɩɪɢ ɤɪɨɜɨɩɨɬɟɪɟ.

ɂɨɧɵ ɋɚ2+ ɫɭɠɢɜɚɸɬ ɫɨɫɭɞɵ.

Ʉ ɫɨɫɭɞɨɪɚɫɲɢɪɹɸɳɢɦ ɜɟɳɟɫɬɜɚɦ ɨɬɧɨɫɹɬɫɹ: ɦɟɞɢɚ-

ɬɨɪ ɚɰɟɬɢɥɯɨɥɢɧ, ɚ ɬɚɤɠɟ ɬɚɤ ɧɚɡɵɜɚɟɦɵɟ ɦɟɫɬɧɵɟ ɝɨɪɦɨɧɵ.

Ɉɞɢɧ ɢɡ ɧɢɯ – ɝɢɫɬɚɦɢɧ – ɨɛɪɚɡɭɟɬɫɹ ɜ ɫɥɢɡɢɫɬɨɣ ɨɛɨɥɨɱɤɟ ɠɟɥɭɞɤɚ ɢ ɤɢɲɟɱɧɢɤɚ, ɜ ɤɨɠɟ, ɫɤɟɥɟɬɧɨɣ ɦɭɫɤɭɥɚɬɭɪɟ (ɜɨ

413

ɜɪɟɦɹ ɪɚɛɨɬɵ) ɢ ɜ ɞɪɭɝɢɯ ɨɪɝɚɧɚɯ. ɋɨɞɟɪɠɢɬɫɹ ɜ ɛɚɡɨɮɢɥɚɯ ɢ ɬɭɱɧɵɯ ɤɥɟɬɤɚɯ ɩɨɜɪɟɠɞɟɧɧɵɯ ɬɤɚɧɟɣ ɢ ɜɵɞɟɥɹɟɬɫɹ ɩɪɢ ɪɟɚɤɰɢɹɯ ɚɧɬɢɝɟɧ–ɚɧɬɢɬɟɥɨ. Ɋɚɫɲɢɪɹɟɬ ɚɪɬɟɪɢɨɥɵ ɢ ɜɟɧɭɥɵ, ɭɜɟɥɢɱɢɜɚɟɬ ɩɪɨɧɢɰɚɟɦɨɫɬɶ ɤɚɩɢɥɥɹɪɨɜ.

Ȼɪɚɞɢɤɢɧɢɧ ɜɵɞɟɥɟɧ ɢɡ ɷɤɫɬɪɚɤɬɨɜ ɩɨɞɠɟɥɭɞɨɱɧɨɣ ɠɟɥɟɡɵ, ɥɟɝɤɢɯ. Ɉɧ ɪɚɫɲɢɪɹɟɬ ɫɨɫɭɞɵ ɫɤɟɥɟɬɧɵɯ ɦɵɲɰ, ɫɟɪɞɰɚ, ɫɩɢɧɧɨɝɨ ɢ ɝɨɥɨɜɧɨɝɨ ɦɨɡɝɚ, ɫɥɸɧɧɵɯ ɢ ɩɨɬɨɜɵɯ ɠɟɥɟɡ, ɭɜɟɥɢɱɢɜɚɟɬ ɩɪɨɧɢɰɚɟɦɨɫɬɶ ɤɚɩɢɥɥɹɪɨɜ.

ɉɪɨɫɬɚɝɥɚɧɞɢɧɵ, ɩɪɨɫɬɚɰɢɤɥɢɧɵ ɢ ɬɪɨɦɛɨɤɫɚɧ ɨɛɪɚ-

ɡɭɸɬɫɹ ɜɨ ɦɧɨɝɢɯ ɨɪɝɚɧɚɯ ɢ ɬɤɚɧɹɯ. Ɉɧɢ ɫɢɧɬɟɡɢɪɭɸɬɫɹ ɢɡ ɩɨɥɢɧɟɧɚɫɵɳɟɧɧɵɯ ɠɢɪɧɵɯ ɤɢɫɥɨɬ ɚɪɚɯɢɞɨɧɨɜɨɣ ɢ ɥɢɧɨɥɟɜɨɣ. ɉɪɨɫɬɚɝɥɚɧɞɢɧɵ (ɊG) – ɷɬɨ ɝɨɪɦɨɧɨɩɨɞɨɛɧɵɟ ɜɟɳɟɫɬɜɚ. Ɋɚɡɥɢɱɧɵɟ ɝɪɭɩɩɵ ɢ ɩɨɞɝɪɭɩɩɵ ɷɬɢɯ ɜɟɳɟɫɬɜ ɨɤɚɡɵɜɚɸɬ ɧɟɨɞɢɧɚɤɨɜɵɣ ɷɮɮɟɤɬ ɧɚ ɫɨɫɭɞɵ. Ɍɚɤ, PGA1 ɢ PGA2 ɜɵɡɵɜɚɸɬ ɪɚɫɲɢɪɟɧɢɟ ɚɪɬɟɪɢɣ ɱɪɟɜɧɨɣ ɨɛɥɚɫɬɢ. Ɇɟɞɭɥɥɢɧ (PGA2), ɜɵɞɟɥɟɧɧɵɣ ɢɡ ɦɨɡɝɨɜɨɝɨ ɜɟɳɟɫɬɜɚ ɩɨɱɟɤ, ɫɧɢɠɚɟɬ ȺȾ, ɭɜɟɥɢɱɢɜɚɟɬ ɩɨɱɟɱɧɵɣ ɤɪɨɜɨɬɨɤ, ɜɵɞɟɥɟɧɢɟ ɩɨɱɤɚɦɢ ɜɨɞɵ, Nɚ+ ɢ Ʉ+. ɉɪɨɫɬɚɝɥɚɧɞɢɧɵ PGE ɪɚɫɲɢɪɹɸɬ ɫɨɫɭɞɵ ɩɪɢ ɜɧɭɬɪɢɚɪɬɟɪɢɚɥɶɧɨɦ ɜɜɟɞɟɧɢɢ ɢ ɬɨɪɦɨɡɹɬ ɜɵɞɟɥɟɧɢɟ ɧɨɪɚɞɪɟɧɚɥɢɧɚ ɢɡ ɨɤɨɧɱɚɧɢɣ ɫɢɦɩɚɬɢɱɟɫɤɢɯ ɧɟɪɜɨɜ. PGF ɫɭɠɢɜɚɸɬ ɫɨɫɭɞɵ ɢ ɩɨɜɵɲɚɸɬ ȺȾ. Ɍɪɨɦɛɨɤɫɚɧ ɨɤɚɡɵɜɚɟɬ ɫɨɫɭɞɨɫɭɠɢɜɚɸɳɢɣ ɷɮɮɟɤɬ.

ɉɪɨɞɭɤɬɵ ɦɟɬɚɛɨɥɢɡɦɚ – ɦɨɥɨɱɧɚɹ ɢ ɩɢɪɨɜɢɧɨɝɪɚɞɧɚɹ ɤɢɫɥɨɬɵ ɨɤɚɡɵɜɚɸɬ ɦɟɫɬɧɵɣ ɜɚɡɨɞɢɥɹɬɚɬɨɪɧɵɣ ɷɮɮɟɤɬ.

CO2 ɪɚɫɲɢɪɹɟɬ ɫɨɫɭɞɵ ɦɨɡɝɚ, ɤɢɲɟɱɧɢɤɚ, ɫɤɟɥɟɬɧɨɣ ɦɭɫɤɭɥɚɬɭɪɵ.

Ⱥɞɟɧɨɡɢɧ ɢ NO (ɨɤɫɢɞ ɚɡɨɬɚ) ɪɚɫɲɢɪɹɸɬ ɤɨɪɨɧɚɪɧɵɟ ɫɨɫɭɞɵ.

ɂɨɧɵ Ʉ+ ɢ Nɚ+ ɪɚɫɲɢɪɹɸɬ ɫɨɫɭɞɵ.

ɐɟɧɬɪɵ ɤɪɨɜɨɨɛɪɚɳɟɧɢɹ

Ɉɛɪɚɡɨɜɚɧɢɹ, ɢɦɟɸɳɢɟ ɨɬɧɨɲɟɧɢɟ ɤ ɰɟɧɬɪɚɦ ɤɪɨɜɨɨɛɪɚɳɟɧɢɹ, ɪɚɫɩɨɥɚɝɚɸɬɫɹ ɧɚ ɪɚɡɧɵɯ ɭɪɨɜɧɹɯ ɰɟɧɬɪɚɥɶɧɨɣ ɧɟɪɜɧɨɣ ɫɢɫɬɟɦɵ (ɪɢɫ. 7.27).

ɋɩɢɧɚɥɶɧɵɣ ɭɪɨɜɟɧɶ ɪɟɝɭɥɹɰɢɢ ɫɟɪɞɟɱɧɨɣ ɞɟɹɬɟɥɶɧɨɫɬɢ ɧɚɯɨɞɢɬɫɹ ɜ ɛɨɤɨɜɵɯ ɪɨɝɚɯ (Ɍh1–Ɍh5) ɫɩɢɧɧɨɝɨ ɦɨɡɝɚ – ɷɬɨ

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