ArticlePDF Available

Abstract and Figures

A cytoembryological study of the seed formation in Orchis purpurea Huds. ssp. purpurea from two populations of Krasnodar region, which are characterized by low real seed productivity, is carried out. It is established that the seeds become nonviable up to the moment of dissemination because of the lethal anomalies in embryo and suspensor development, that are connected with the manifestation of genetic instability in the division and differentiation of cells during embryogenesis.
No caption available
… 
Content may be subject to copyright.
ɋɟɥɶɫɤɨɯɨɡɹɣɫɬɜɟɧɧɵɟɧɚɭɤɢ ,661ɇɚɭɤɚɄɭɛɚɧɢʋ 13
ɍȾɄ
ȿȼȺɧɞɪɨɧɨɜɚȿȿȿɜɞɨɤɢɦɨɜɚȺȼɋɟɦɺɧɨɜ
ɉɊɂɑɂɇɕɎɈɊɆɂɊɈȼȺɇɂəɇȿɀɂɁɇȿɋɉɈɋɈȻɇɕɏ
ȾɂȺɋɉɈɊɍ25&+,6385385($68%63&$8&$6,&$25&+,'$&($(
Ɏɟɞɟɪɚɥɶɧɨɟɝɨɫɭɞɚɪɫɬɜɟɧɧɨɟɛɸɞɠɟɬɧɨɟɭɱɪɟɠɞɟɧɢɟɧɚɭɤɢ
ȻɨɬɚɧɢɱɟɫɤɢɣɢɧɫɬɢɬɭɬɢɦȼɅɄɨɦɚɪɨɜɚɊɨɫɫɢɣɫɤɨɣɚɤɚɞɟɦɢɢɧɚɭɤ
ɎɢɥɢɚɥɎɟɞɟɪɚɥɶɧɨɝɨɛɸɞɠɟɬɧɨɝɨɭɱɪɟɠɞɟɧɢɹ
©Ɋɨɫɫɢɣɫɤɢɣɰɟɧɬɪɡɚɳɢɬɵɥɟɫɚª
©ɐɟɧɬɪɡɚɳɢɬɵɥɟɫɚɄɪɚɫɧɨɞɚɪɫɤɨɝɨɤɪɚɹª
ɉɪɨɜɟɞɟɧɨɰɢɬɨɷɦɛɪɢɨɥɨɝɢɱɟɫɤɨɟɢɫɫɥɟɞɨɜɚɧɢɟ ɮɨɪɦɢɪɨɜɚɧɢɹ ɫɟɦɹɧ ɭOrchis purpurea+XGV
VVSSXUSXUHDɢɡɞɜɭɯɩɨɩɭɥɹɰɢɣɄɪɚɫɧɨɞɚɪɫɤɨɝɨɤɪɚɹɯɚɪɚɤɬɟɪɢɡɭɸɳɢɯɫɹɧɢɡɤɨɣɪɟɚɥɶɧɨɣɫɟ-
ɦɟɧɧɨɣ ɩɪɨɞɭɤɬɢɜɧɨɫɬɶɸ ɍɫɬɚɧɨɜɥɟɧɨ ɱɬɨɫ ɟɦɟɧɚ ɫɬɚɧɨɜɹɬɫɹ ɧɟɠɢɡɧɟɫɩɨɫɨɛɧɵɦɢ ɤ ɦɨɦɟɧ-
ɬɭ ɞɢɫɫɟɦɢɧɚɰɢɢ ɢɡɡɚ ɥɟɬɚɥɶɧɵɯ ɚɧɨɦɚɥɢɣ ɫɬɪɨɟɧɢɹ ɡɚɪɨɞɵɲɚ ɢ ɫɭɫɩɟɧɡɨɪɚ ɤɨɬɨɪɵɟ ɫɜɹɡɚ-
ɧɵɫɩɪɨɹɜɥɟɧɢɟɦɝɟɧɟɬɢɱɟɫɤɨɣɧɟɫɬɚɛɢɥɶɧɨɫɬɢɩɪɢɞɟɥɟɧɢɢɢɞɢɮɮɟɪɟɧɰɢɚɰɢɢ ɤɥɟɬɨɤɜɯɨɞɟ
ɷɦɛɪɢɨɝɟɧɟɡɚ
ɄɥɸɱɟɜɵɟɫɥɨɜɚɪɚɫɬɢɬɟɥɶɧɵɟɛɢɨɪɟɫɭɪɫɵɧɨɜɵɟɢɧɟɬɪɚɞɢɰɢɨɧɧɵɟɤɭɥɶɬɭɪɵɪɚɫɬɟɧɢɣOrchis
SXUSXUHDɷɦɛɪɢɨɝɟɧɟɡɚɧɨɦɚɥɢɢɪɚɡɜɢɬɢɹɡɚɪɨɞɵɲɚɢɫɭɫɩɟɧɡɨɪɚɧɟɠɢɡɧɟɫɩɨɫɨɛɧɨɫɬɶɫɟɦɹɧ
ɝɟɧɟɬɢɱɟɫɤɚɹɧɟɫɬɚɛɢɥɶɧɨɫɬɶ
ȼɜɟɞɟɧɢɟ ȼɫɟ ɞɢɤɨɪɚɫɬɭɳɢɟ ɜɢɞɵ
ɪɚɫɬɟɧɢɣ ɨɬɧɨɫɹɬɫɹ ɤɛɢɨɥɨɝɢɱɟɫɤɢɦ ɪɟ-
ɫɭɪɫɚɦ ɢɩɪɟɞɫɬɚɜɥɹɸɬ ɫɨɛɨɣ ɜɚɠɧɭɸ ɱɚɫɬɶ
ɫɵɪɶɟɜɨɝɨ ɩɨɬɟɧɰɢɚɥɚ Ɋɨɫɫɢɢ Ɉɞɧɚɤɨ ɦɧɨ-
ɝɢɟ ɢɡ ɧɢɯ ɹɜɥɹɸɬɫɹ ɪɟɞɤɢɦɢ ɢɫɱɟɡɚɸɳɢɦɢ
ɪɚɫɬɟɧɢɹɦɢ ɢɢɯ ɩɪɚɤɬɢɱɟɫɤɨɟ ɢɫɩɨɥɶɡɨɜɚ-
ɧɢɟ ɨɝɪɚɧɢɱɢɜɚɟɬɫɹ ɷɬɢɦ ɮɚɤɬɨɦ ɇɚ ɫɨɜɪɟ-
ɦɟɧɧɨɦ ɷɬɚɩɟ ɨɱɟɜɢɞɧɚ ɧɟɨɛɯɨɞɢɦɨɫɬɶ ɪɚɫ-
ɲɢɪɟɧɢɹ ɛɢɨɪɟɫɭɪɫɧɨɣ ɛɚɡɵ Ɋɨɫɫɢɢ ɡɚ ɫɱɟɬ
ɜɜɟɞɟɧɢɹɜɤɭɥɶɬɭɪɭɧɨɜɵɯɧɟɬɪɚɞɢɰɢɨɧɧɵɯ
ɪɚɫɬɟɧɢɣɫɯɨɡɹɣɫɬɜɟɧɧɨɰɟɧɧɵɦɢɩɪɢɡɧɚɤɚ-
ɦɢɈɞɧɨɣɢɡ ɡɚɞɚɱɜɞɟɥɟɫɨɯɪɚɧɟɧɢɹɢɪɚɫ-
ɲɢɪɟɧɢɹ ɛɢɨɪɟɫɭɪɫɧɨɣ ɛɚɡɵ ɧɚɲɟɣ ɫɬɪɚɧɵ
ɹɜɥɹɟɬɫɹɪɚɡɜɢɬɢɟ ɬɟɯɧɨɥɨɝɢɣ ɫɨɡɞɚɧɢɹ ɛɢɨ-
ɤɭɥɶɬɭɪ ɪɟɫɭɪɫɧɵɯ ɜɢɞɨɜ ɢɢɫɤɭɫɫɬɜɟɧɧɵɯ
ɷɤɨɫɢɫɬɟɦɞɥɹɩɟɪɟɯɨɞɚɨɬ ɢɡɴɹɬɢɹ ɛɢɨɥɨɝɢ-
ɱɟɫɤɢɯ ɨɛɴɟɤɬɨɜ ɢɡ ɩɪɢɪɨɞɧɵɯ ɩɨɩɭɥɹɰɢɣ
ɤɢɯɤɭɥɶɬɢɜɢɪɨɜɚɧɢɸ>@
Orchis purpurea ɹɬɪɵɲɧɢɤ ɩɭɪɩɭɪ-
ɧɵɣ ɢɡ ɫɟɦɟɣɫɬɜɚ 2UFKLGDFHDH ɹɜɥɹɟɬɫɹ
ɨɞɧɢɦ ɢɡ ɪɟɫɭɪɫɧɵɯ ɪɚɫɬɟɧɢɣ ȼɦɟɞɢɰɢɧɟ
ɢɩɢɳɟɜɨɣ ɩɪɨɦɵɲɥɟɧɧɨɫɬɢ ɢɫɩɨɥɶɡɭɸɬ
ɩɨɞɡɟɦɧɵɣɨɪɝɚɧɷɬɨɝɨɪɚɫɬɟɧɢɹ² ɬɭɛɟɪɨ-
ɢɞɤɨɪɧɟɜɨɣɤɥɭɛɟɧɶɗɬɨɬɜɢɞɢɦɟɟɬɟɜɪɨ
ɩɟɣɫɤɨɦɚɥɨɚɡɢɚɬɫɤɢɣ ɚɪɟɚɥ ɪɚɫɩɪɨɫɬɪɚɧɟ-
ɧɢɹɜɊɨɫɫɢɢɧɚɯɨɞɢɬɫɹɧɚɫɟɜɟɪɨɜɨɫɬɨɱɧɨɦ
ɩɪɟɞɟɥɟ ɩɪɨɢɡɪɚɫɬɚɧɢɹ ɜɧɟɫɟɧ ɜɄɪɚɫɧɵɟ
ɤɧɢɝɢ Ɋɨɫɫɢɣɫɤɨɣ Ɏɟɞɟɪɚɰɢɢ Ʉɪɚɫɧɨɞɚɪ
ɫɤɨɝɨ ɢɋɬɚɜɪɨɩɨɥɶɫɤɨɝɨ ɤɪɚɟɜ ɪɟɫɩɭɛɥɢɤ
Ⱥɞɵɝɟɹ Ⱦɚɝɟɫɬɚɧ ɂɧɝɭɲɟɬɢɹ Ʉɚɛɚɪɞɢɧɨ
Ȼɚɥɤɚɪɫɤɚɹ Ʉɚɪɚɱɚɟɜɨɑɟɪɤɟɫɢɹ ɋɟɜɟɪɧɚɹ
ȿȼȺɧɞɪɨɧɨɜɚȿȿȿɜɞɨɤɢɦɨɜɚȺȼɋɟɦɟɧɨɜ
Ⱥɧɞɪɨɧɨɜɚ ȿɥɟɧɚ ȼɚɥɟɧɬɢɧɨɜɧɚ² ɤɚɧɞ ɛɢɨɥ
ɧɚɭɤ ɫɬ ɧɚɭɱ ɫɨɬɪɭɞɧɢɤ ɥɚɛɨɪɚɬɨɪɢɢ ɗɦɛɪɢɨɥɨɝɢɢ
ɢɪɟɩɪɨɞɭɤɬɢɜɧɨɣɛɢɨɥɨɝɢɢȻɂɇɊȺɇɝɋɚɧɤɬɉɟɬɟɪɛɭɪɝ
HPDLOHOHQDDQGURQRYD#PDLOUX
ȿɜɞɨɤɢɦɨɜɚ ȿɥɟɧɚ ȿɜɝɟɧɶɟɜɧɚ² ɦɥ ɧɚɭɱ ɫɨɬɪɭɞɧɢɤ
ɥɚɛɨɪɚɬɨɪɢɢ ɗɦɛɪɢɨɥɨɝɢɢ ɢɪɟɩɪɨɞɭɤɬɢɜɧɨɣ ɛɢɨɥɨɝɢɢ
ȻɂɇɊȺɇɝɋɚɧɤɬɉɟɬɟɪɛɭɪɝ
ɋɟɦɺɧɨɜȺɥɟɤɫɚɧɞɪȼɥɚɞɢɦɢɪɨɜɢɱ²ɦɥɧɚɭɱɫɨɬɪɭɞ-
ɧɢɤɮɢɥɢɚɥɚ Ɋɨɫɫɢɣɫɤɨɝɨɰɟɧɬɪɚ ɡɚɳɢɬɵɥɟɫɚ ©ɐɟɧɬɪɡɚ-
ɳɢɬɵɥɟɫɚɄɪɚɫɧɨɞɚɪɫɤɨɝɨɤɪɚɹªɝɄɪɚɫɧɨɞɚɪ
Ɋɚɛɨɬɚ ɜɵɩɨɥɧɟɧɚ ɩɨ ɝɨɫɭɞɚɪɫɬɜɟɧɧɨɦɭ ɡɚɞɚɧɢɸ
ʋ ɢɩɨɞɞɟɪɠɚɧɚ ɝɪɚɧɬɨɦ ʋ 
ɪBɸɝBɚ Ɋɨɫɫɢɣɫɤɨɝɨɮɨɧɞɚ ɮɭɧɞɚɦɟɧɬɚɥɶɧɵɯ ɢɫɫɥɟɞɨ
ɜɚɧɢɣɢɚɞɦɢɧɢɫɬɪɚɰɢɢɄɪɚɫɧɨɞɚɪɫɤɨɝɨɤɪɚɹ
ɈɊɂȽɂɇȺɅɖɇɕȿɋɌȺɌɖɂ,661ɇɚɭɤɚɄɭɛɚɧɢʋ
14
Ɉɫɟɬɢɹ ɑɟɱɟɧɫɤɚɹ ɢɝɨɪɨɞɚ ɋɨɱɢ >@ Ⱦɥɹ
ɟɫɬɟɫɬɜɟɧɧɨɝɨ ɜɨɡɨɛɧɨɜɥɟɧɢɹ ɛɢɨɪɟɫɭɪɫɚ
ɹɬɪɵɲɧɢɤɚ ɩɭɪɩɭɪɧɨɝɨ ɧɟɨɛɯɨɞɢɦɨ ɛɨɥɟɟ
ɥɟɬȼɧɚɫɬɨɹɳɟɟɜɪɟɦɹɩɨɥɭɱɟɧɢɟɛɢɨɫɵ-
ɪɶɹɢɡɟɫɬɟɫɬɜɟɧɧɵɯɩɨɩɭɥɹɰɢɣ2SXUSXUHD
ɩɪɨɢɡɪɚɫɬɚɸɳɢɯɜɊɨɫɫɢɢɧɟɜɨɡɦɨɠɧɨɬɚɤ
ɤɚɤ ɜɢɞ ɨɬɧɨɫɢɬɫɹ ɤɪɟɞɤɢɦ ɢɫɱɟɡɚɸɳɢɦ
ɪɚɫɬɟɧɢɹɦȻɢɨɫɵɪɶɺɹɬɪɵɲɧɢɤɚɩɭɪɩɭɪɧɨ-
ɝɨ ɦɨɠ ɧɨ ɩɨɥɭ ɱɚɬɶ ɜɪɟɡɭɥɶɬɚɬɟɪɚɡɪɚɛɨɬɤɢ
ɢɧɧɨɜɚɰɢɨɧɧɵɯ ɛɢɨɬɟɯɧɨɥɨɝɢɣ ɪɚɡɦɧɨɠɟ-
ɧɢɹ ɢɩɨɫɥɟɞɭɸɳɟɝɨ ɤɭɥɶɬɢɜɢɪɨɜɚɧɢɹ ɪɚɫ-
ɬɟɧɢɣɷɬɨɝɨɜɢɞɚɜɤɚɱɟɫɬɜɟɧɨɜɨɣɧɟɬɪɚɞɢ-
ɰɢɨɧɧɨɣɤɭɥɶɬɭɪɵ
2SXUSXUHDɨɬɥɢɱɚɟɬɫɹɢɡɦɟɧɱɢɜɨɫɬɶɸ
ɨɛɳɢɯɪɚɡɦɟɪɨɜɚɬɚɤɠɟɨɤɪɚɫɤɢɢɫɬɟɩɟɧɢ
ɪɚɫɫɟɱɟɧɧɨɫɬɢ ɝɭɛɵ Ɉɫɨɛɢ ɜɢɞɚ ɝɢɛɪɢɞɢ-
ɡɢɪɭɸɬ ɫɞɪɭɝɢɦɢ ɩɪɟɞɫɬɚɜɢɬɟɥɹɦɢ ɪɨɞɚ
ȼɫɜɹɡɢ ɫɩɪɨɹɜɥɟɧɢɟɦ ɜɵɫɨɤɨɝɨ ɩɨɥɢɦɨɪ-
ɮɢɡɦɚ ɜɧɺɦ ɜɵɞɟɥɹɸɬɫɹ ɤɚɤ ɦɢɧɢɦɭɦ ɞɜɚ
ɩɨɞɜɢɞɚ² 2 SXUSXUHD +XGV VVS purpurea
ɢ2 SXUSXUHD +XGV VVS caucasica 5HJHO
%+%DXPDQQ/RUHQ]3HWHU>@ɢɛɨɥɶɲɨɟ
ɱɢɫɥɨɮɨɪɦ
Ⱦɥɹ 2 SXUSXUHD ɯɚɪɚɤɬɟɪɟɧ ɩɪɟɢɦɭ-
ɳɟɫɬɜɟɧɧɨ ɫɟɦɟɧɧɨɣ ɫɩɨɫɨɛ ɪɚɡɦɧɨɠɟɧɢɹ
ɋɨɝɥɚɫɧɨ ɢɦɟɸɳɢɦɫɹ ɫɜɟɞɟɧɢɹɦ ɩɪɨɰɟɧɬ
ɡɚɜɹɡɵɜɚɧɢɹ ɩɥɨɞɨɜ ɭɨɫɨɛɟɣ ɞɚɧɧɨɝɨ ɜɢɞɚ
ɨɱɟɧɶɧɢɡɤɢɣɜɸɠɧɨɣȺɧɝɥɢɢ²ɜɄɪɵ-
ɦɭ²ɜɬɟɧɢɫɬɵɯɥɟɫɚɯ²>@ɉɨ
ɞɚɧɧɵɦɡɚ± ɝɝɩɪɨɰɟɧɬɡɚɜɹɡɵɜɚ-
ɧɢɹɩɥɨɞɨɜɭ2SXUSXUHDVVScaucasicaɜɇɨ-
ɜɨɪɨɫɫɢɣɫɤɨɦɢȺɛɢɧɫɤɨɦ ɪɚɣɨɧɚɯ Ʉɪɚɫɧɨ
ɞɚɪɫɤɨɝɨɤɪɚɹɫɨɫɬɚɜɥɹɥɜɫɟɝɨ±>@
Ⱦɥɹ 2 SXUSXUHD ɨɬɦɟɱɚɟɬɫɹ ɜɵɫɨɤɚɹ
ɫɟɦɟɧɧɚɹ ɩɪɨɞɭɤɬɢɜɧɨɫɬɶ ɧɨ ɫɪɚɜɧɢɬɟɥɶɧɨ
ɧɢɡɤɚɹɪɟɚɥɶɧɚɹɫɟɦɟɧɧɚɹɩɪɨɞɭɤɬɢɜɧɨɫɬɶ²
ɤɚɤ ɡɚ ɫɱɟɬ ɜɵɫɨɤɨɝɨ ɩɪɨɰɟɧɬɚ ɚɛɨɪɬɢɜɧɵɯ
ɫɟɦɹɧ > @ ɬɚɤ ɢɢɡɡɚ ɜɵɫɨɤɨɣ ɞɨɥɢ ɛɟɡ-
ɡɚɪɨɞɵɲɟɜɵɯ ɧɟɠɢɡɧɟɫɩɨɫɨɛɧɵɯ ɞɢɚɫɩɨɪ
ɜɡɪɟɥɵɯɩɥɨɞɚɯɧɚɩɪɢɦɟɪɭ2SXUSXUHDVVS
caucasicaɨɧɚɜɫɪɟɞɧɟɦɫɨɫɬɚɜɥɹɟɬ>@
Ɇɨɠɧɨɩɪɟɞɩɨɥɨɠɢɬɶɧɟɫɤɨɥɶɤɨɮɚɤ-
ɬɨɪɨɜɜɥɢɹɸɳɢɯɧɚɩɥɨɞɨɧɨɲɟɧɢɟɢɧɚɤɚ-
ɱɟɫɬɜɨ ɫɟɦɹɧ ɭ2 SXUSXUHD VVS FDXFDVLFD
ɗɬɢ ɮɚɤɬɨɪɵ ɦɨɠɧɨ ɪɚɡɞɟɥɢɬɶ ɧɚ ɝɪɭɩ-
ɩɵ ɩɨ ɢɯ ɜɥɢɹɧɢɸ ɧɚ ɩɪɨɰɟɫɫ ɪɚɡɜɢɬɢɹ
ɞɢɚɫɩɨɪ
ɇɢɡɤɢɣ ɩɪɨɰɟɧɬ ɡɚɜɹɡɵɜɚɧɢɹ ɩɥɨɞɨɜ
ɢɫɧɢɠɟɧɢɟɤɚɱɟɫɬɜɚɫɟɦɹɧɦɨɝɭɬɛɵɬɶɫɜɹ-
ɡɚɧɵ ɫɧɟɨɩɬɢɦɚɥɶɧɵɦɢ ɭɫɥɨɜɢɹɦɢ ɦɟɫɬɚ
ɩɪɨɢɡɪɚɫɬɚɧɢɹ ɨɫɨɛɟɣ ɧɚ ɝɪɚɧɢɰɟ ɚɪɟɚɥɚ
Ʉɪɨɦɟ ɬɨɝɨɧɟ ɢɫɤɥɸɱɟɧɨ ɱɬɨ ɜɝɨɞɵ ɩ ɪɨ-
ɜɟɞɟɧɢɹɢɫɫɥɟɞɨɜɚɧɢɣɩɨɝɨɞɧɵɟɭɫɥɨɜɢɹ ɧɟ
ɛɵɥɢ ɛɥɚɝɨɩɪɢɹɬɧɵɦɢ ɞɥɹ ɨɩɵɥɟɧɢɹ ɢɞɥɹ
ɪɨɫɬɚ ɪɚɫɬɟɧɢɣ ɱɬɨ ɩɪɢɜɟɥɨ ɤɧɟɞɨɨɩɵ-
ɥɟɧɢɸ ɰɜɟɬɤɨɜ ɢ ɤɚɤ ɫɥɟɞɫɬɜɢɟ ɤɨɬɦɢɪɚ-
ɧɢɸ ɧɟɨɩɥɨɞɨɬɜɨɪɟɧɧɵɯ ɫɟɦɹɡɚɱɚɬɤɨɜ ɢɥɢ
ɤɨɬɫɭɬɫɬɜɢɸ ɞɨɫɬɚɬɨɱɧɨɝɨ ɤɨɥɢɱɟɫɬɜɚ ɩɢ-
ɬɚɬɟɥɶɧɵɯɜɟɳɟɫɬɜɧɟɨɛɯɨɞɢɦɵɯɞɥɹɪɚɡɜɢ-
ɬɢɹɜɫɟɯɨɛɪɚɡɨɜɚɜɲɢɯɫɹɞɢɚɫɩɨɪ ɢɜɵɛɪɚ-
ɤɨɜɤɟ ɧɟɤɨɬɨɪɨɣ ɱɚɫɬɢ ɫɟɦɹɧ Ɉɞɧɚɤɨ ɟɫɥɢ
ɡɚɪɨɞɵɲɨɛɪɚɡɭɟɬɫɹɬɨɟɝɨɪɚɡɜɢɬɢɟɞɨɥɠɧɨ
ɩɪɨɢɫɯɨɞɢɬɶ ɜɫɨɨɬɜɟɬɫɬɜɢɢ ɫɬɚɤɫɨɧɫɩɟɰɢ-
ɮɢɱɧɵɦɢ ɡɚɤɨɧɨɦɟɪɧɨɫɬɹɦɢ ɮɨɪɦɢɪɨɜɚɧɢɹ
ɤɥɟɬɨɱɧɵɯɝɟɧɟɪɚɰɢɣɢɥɢɡɚɤɨɧɚɦɢɷɦɛɪɢɨ-
ɝɟɧɢɢ>@
ȼɫɜɨɸɨɱɟɪɟɞɶɜɵɫɨɤɚɹɞɨɥɹɩɨɥɧɨɫ
ɬɶɸ ɧɟɠɢɡɧɟɫɩɨɫɨɛɧɵɯ ɫɟɦɹɧ ɤɦɨɦɟɧɬɭ
ɞɢɫɫɟɦɢɧɚɰɢɢ ɩɨɜɢɞɢɦɨɦɭ ɦɨɠɟɬ ɛɵɬɶ
ɫɥɟɞɫɬɜɢɟɦ ɦɟɠɜɢɞɨɜɨɣ ɝɢɛɪɢɞɢɡɚɰɢɢ
ɢɢɥɢɢɧɛɪɢɞɢɧɝɚȾɟɣɫɬɜɢɹ ɜɬɨɪɨɣɝɪɭɩɩɵ
ɮɚɤɬɨɪɨɜɫɨɩɪɨɜɨɠɞɚɸɬɫɹɝɟɧɟɬɢɱɟɫɤɨɣɧɟ-
ɫɬɚɛɢɥɶɧɨɫɬɶɸɱɬɨɩɪɢɜɨɞɢɬɤɚɧɨɦɚɥɶɧɨɦɭ
ɪɚɡɜɢɬɢɸɡɚɪɨɞɵɲɚɬɨɟɫɬɶɤɪɚɡɥɢɱɧɵɦɨɬ-
ɤɥɨɧɟɧɢɹɦɨɬɧɨɪɦɵɩɨɞɨɛɧɨɬɨɦɭɤɚɤɷɬɨ
ɨɩɢɫɚɧɨɭɷɦɛɪɢɨɥɟɬɚɥɶɧɵɯɦɭɬɚɧɬɨɜɚɪɚɛɢ-
ɞɨɩɫɢɫɚ >@ ȼɫɥɭɱɚɟ ɨɪɯɢɞɧɵɯ ɥɟɬɚɥɶɧɵɟ
ɚɧɨɦɚɥɢɢɪɚɡɜɢɬɢɹɡɚɪɨɞɵɲɚɨɩɢɫɚɧɵɭɨɫɨ-
ɛɢDactylorhiza fucshiiɢɡɦɚɥɨɱɢɫɥɟɧɧɨɣɩɨ-
ɩɭɥɹɰɢɢɜɫɟɝɨɨɫɨɛɢɭɤɨɬɨɪɨɣɜɡɪɟɥɵɯ
ɩɥɨɞɚɯɬɨɥɶɤɨɫɟɦɹɧɢɦɟɥɢɡɚɪɨɞɵɲ>@
Ɏɚɤɬɨɪɵɜɥɢɹɸɳɢɟɧɚɤɚɱɟɫɬɜɨɫɟɦɹɧ
ɭɪɟɞɤɢɯɜɢɞɨɜ ɨɪɯɢɞɧɵɯ ɜɧɚɫɬɨɹɳɟɟɜɪɟ-
ɦɹɢɡɭɱɟɧɵɧɟɞɨɫɬɚɬɨɱɧɨɩɨɞɪɨɛɧɨɇɢɡɤɨɟ
ɤɚɱɟɫɬɜɨ ɫɟɦɹɧ ɦɨɠɟɬ ɛɵɬɶ ɨɞɧɢɦ ɢɡ ɥɢ-
ɦɢɬɢɪɭɸɳɢɯ ɮɚɤɬɨɪɨɜ ɜɥɢɹɸɳɢɯ ɧɚ ɱɢɫ
ɥɟɧɧɨɫɬɶɧɟɤɨɬɨɪɵɯɪɟɞɤɢɯɜɢɞɨɜɪɚɫɬɟɧɢɣ
ɜɬɨɦɱɢɫɥɟɨɪɯɢɞɧɵɯɢɧɚɢɯɟɫɬɟɫɬɜɟɧɧɨɟ
ɜɨɡɨɛɧɨɜɥɟɧɢɟ ɢɪɚɫɩɪɨɫɬɪɚɧɟɧɢɟ Ɉɰɟɧ-
ɤɚ ɤɚɱɟɫɬɜɚ ɫɟɦɹɧ ɜɚɠɧɚ ɢɞɥɹ ɪɚɡɪɚɛɨɬɤɢ
ɢɧɧɨɜɚɰɢɨɧɧɵɯ ɛɢɨɬɟɯɧɨɥɨɝɢɣ ɩɨɥɭɱɟɧɢɹ
ɩɨɫɚɞɨɱɧɨɝɨ ɦɚɬɟɪɢɚɥɚ 2 SXUSXUHD ɤɚɤ ɩɨ-
ɬɟɧɰɢɚɥɶɧɨ ɧɨɜɨɝɨ ɧɟɬɪɚɞɢɰɢɨɧɧɨɝɨ ɤɭɥɶ-
ɬɭɪɧɨɝɨɪɚɫɬɟɧɢɹ
ɋɟɥɶɫɤɨɯɨɡɹɣɫɬɜɟɧɧɵɟɧɚɭɤɢ ,661ɇɚɭɤɚɄɭɛɚɧɢʋ 15
ɐɟɥɶɸɞɚɧɧɨɣɪɚɛɨɬɵɹɜɥɹɥɨɫɶɩɪɨɜɟ-
ɞɟɧɢɟɰɢɬɨɷɦɛɪɢɨɥɨɝɢɱɟɫɤɨɝɨɢɫɫɥɟɞɨɜɚɧɢɹ
ɮɨɪɦɢɪɨɜɚɧɢɹɫɟɦɹɧɭ 2SXUSXUHD+XGVVVS
purpureaɢɡɞɜɭɯɩɨɩɭɥɹɰɢɣɄɪɚɫɧɨɞɚɪɫɤɨɝɨ
ɤɪɚɹ ɯɚɪɚɤɬɟɪɢɡɭɸɳɢɯɫɹ ɧɢɡɤɨɣ ɪɟɚɥɶɧɨɣ
ɫɟɦɟɧɧɨɣɩɪɨɞɭɤɬɢɜɧɨɫɬɶɸ
Ɇɚɬɟɪɢɚɥɵ ɢɦɟɬɨɞɵ ɂɫɫɥɟɞɨɜɚɧɢɹ
ɜɵɩɨɥɧɟɧɵ ɜɥɚɛɨɪɚɬɨɪɢɢ ɗɦɛɪɢɨɥɨɝɢɢ
ɢɪɟɩɪɨɞɭɤɬɢɜɧɨɣ ɛɢɨɥɨɝɢɢ Ȼɨɬɚɧɢɱɟɫɤɨ-
ɝɨ ɢɧɫɬɢɬɭɬɚ ɢɦȼɅɄɨɦɚɪɨɜɚ ɊȺɇ Ȼɂɇ
ɊȺɇ ɉɨɫɤɨɥɶɤɭ ɭɨɫɨɛɟɣ ɞɚɧɧɨɝɨ ɜɢɞɚ
ɜɦɟɫɬɚɯ ɢɯ ɟɫɬɟɫɬɜɟɧɧɨɝɨ ɩɪɨɢɡɪɚɫɬɚɧɢɹ
ɜɇɨɜɨɪɨɫɫɢɣɫɤɨɦ ɢȺɛɢɧɫɤɨɦ ɪɚɣɨɧɚɯ
Ʉɪɚɫɧɨɞɚɪɫɤɨɝɨ ɤɪɚɹ ɩɪɨɰɟɧɬ ɡɚɜɹɡɵɜɚɧɢɹ
ɩɥɨɞɨɜ ɨɱɟɧɶ ɧɢɡɤɢɣ ɬɨ ɞɥɹ ɰɢɬɨɷɦɛɪɢɨ-
ɥɨɝɢɱɟɫɤɨɝɨ ɢɫɫɥɟɞɨɜɚɧɢɹ ɫɟɦɹɧ ɧɚ ɪɚɡɧɵɯ
ɫɬɚɞɢɹɯɢɯɪɚɡɜɢɬɢɹɩɪɨɜɨɞɢɥɢɢɫɤɭɫɫɬɜɟɧ-
ɧɨɟ ɨɩɵɥɟɧɢɟ ɰɜɟɬɤɨɜ ɭ ɪɚɫɬɟɧɢɣ ɞɚɬɚ
ɨɩɵɥɟɧɢɹ  Ɉɩɵɥɟɧɧɵɟ ɰɜɟɬɤɢ
ɦɚɪɤɢɪɨɜɚɥɢ ɧɢɬɤɨɣ ɫɷɬɢɤɟɬɤɨɣ ȼɪɚɛɨɬɟ
ɢɫɩɨɥɶɡɨɜɚɥɢɬɚɤɠɟɩɥɨɞɵɨɫɨɛɟɣɰɜɟɬɤɢ
ɤɨɬɨɪɵɯɧɟɨɩɵɥɹɥɢ
ȼɫɟ ɪɚɫɬɟɧɢɹ ɡɚɞɟɣɫɬɜɨɜɚɧɧɵɟ ɜɷɤɫ-
ɩɟɪɢɦɟɧɬɟ ɛɵɥɢ ɦɚɪɤɢɪɨɜɚɧɵ ɷɬɢɤɟɬɤɚɦɢ
ɫɧɨɦɟɪɨɦɋɤɚɠɞɨɝɨɢɡɧɢɯɛɵɥɢɫɨɛɪɚɧɵ
ɰɜɟɬɤɢɞɥɹɦɨɪɮɨɥɨɝɢɱɟɫɤɨɝɨɚɧɚɥɢɡɚɢɞɥɹ
ɭɬɨɱɧɟɧɢɹɜɨɡɦɨɠɧɨɫɬɢɢɯɝɢɛɪɢɞɧɨɝɨɩɪɨ-
ɢɫɯɨɠɞɟɧɢɹ
ɋɨɰɜɟɬɢɹ ɫɧɟɡɪɟɥɵɦɢ ɩɥɨɞɚɦɢ ɫɨ-
ɛɢɪɚɥɢ  ɬɪɚɧɫɩɨɪɬɢɪɨɜɚɥɢ ɧɟ
ɫɤɨɥɶɤɨ ɫɭɬɨɤ ɞɨ ɥɚɛɨɪɚɬɨɪɢɢ ɢɯɪɚɧɢɥɢ
ɜɯɨɥɨɞɢɥɶɧɢɤɟ ɩɪɢ ɬɟɦɩɟɪɚɬɭɪɟ  ɋɞɨ
 ɡɚɬɟɦ ɮɢɤɫɢɪɨɜɚɥɢ ɜɧɨɦ
ɷɬɚɧɨɥɟ Ɍɟɦ ɫɚɦɵɦ ɮɢɤɫɚɰɢɹ ɫɟɦɹɧ ɛɵɥɚ
ɩɪɨɜɟɞɟɧɚɱɟɪɟɡ±ɞɧɟɣɨɬɞɚɬɵɨɩɵɥɟ-
ɧɢɹɰɜɟɬɤɨɜɌɚɤɚɹɫɯɟɦɚɪɚɛɨɬɵɫɦɚɬɟɪɢɚ-
ɥɨɦɫɜɹɡɚɧɚɫɬɟɦɱɬɨɱɚɫɬɶɫɟɦɹɧɢɫɩɨɥɶɡɨ-
ɜɚɥɢɬɚɤɠɟɞɥɹɩɪɨɜɟɞɟɧɢɹɪɨɫɬɨɜɨɝɨɬɟɫɬɚ
ɫɩɪɢɦɟɧɟɧɢɟɦ ɦɟɬɨɞɢɤɢ ɩɪɨɪɚɳɢɜɚɧɢɹ ɢɯ
ɧɚɩɢɬɚɬɟɥɶɧɨɣɫɪɟɞɟLQYLWURɋɟɦɟɧɚɧɚɦɨ-
ɦɟɧɬ ɮɢɤɫɚɰɢɢ ɛɵɥɢ ɧɟɡɪɟɥɵɟ ɦɟɥɤɢɟ ɛɟ-
ɥɨɝɨɰɜɟɬɚɨɧɢɩɥɨɬɧɨɫɢɞɟɥɢɧɚɩɥɚɰɟɧɬɟ
Ɏɢɤɫɢɪɨɜɚɧɧɵɣ ɦɚɬɟɪɢɚɥ ɩɟɪɟɞ ɩɪɨ-
ɜɨɞɤɨɣ ɨɬɦɵɜɚɥɢ ɧɵɦ ɫɩɢɪɬɨɦ ɡɚɬɟɦ
ɨɛɟɡɜɨɠɢɜɚɥɢ ɩɪɨɜɨɞɹ ɱɟɪɟɡ ɫɟɪɢɸ ɫɩɢɪ-
ɬɨɜ ɪɚɡɧɨɣ ɤɨɧɰɟɧɬɪɚɰɢɢ   
 ɫɦɟɫɢ ɫɩɢɪɬ ɧɵɣ ɯɥɨɪɨɮɨɪɦ
ɜɫɨɨɬɧɨɲɟɧɢɢɢɱɢɫɬɵɣɯɥɨɪɨ-
ɮɨɪɦȼɫɨɫɭɞɵɫɦɚɬɟɪɢɚɥɨɦɜɯɥɨɪɨɮɨɪɦɟ
ɧɚɥɢɜɚɥɢ ɪɚɫɩɥɚɜɥɟɧɧɵɣ ɩɚɪɚɮɢɧ ɩɨɫɥɟ
ɱɟɝɨ ɢɯ ɫɬɚɜɢɥɢ ɜɬɟɪɦɨɫɬɚɬ ± żɋ ɞɨ
ɩɨɥɧɨɝɨɢɫɩɚɪɟɧɢɹɯɥɨɪɨɮɨɪɦɚ
Ɂɚɤɥɸɱɟɧɧɵɣ ɜɩɚɪɚɮɢɧ ɦɚɬɟɪɢɚɥ ɪɟ-
ɡɚɥɢɧɚɦɢɤɪɨɬɨɦɟ0LFURP&DUO=HLVV
ɬɨɥɳɢɧɚ ɫɪɟɡɨɜ ɫɨɫɬɚɜɥɹɥɚ  ɦɤɦ ɋɪɟɡɵ
ɦɨɧɬɢɪɨɜɚɥɢɧɚɩɪɟɞɦɟɬɧɵɟɫɬɺɤɥɚɞɟɩɚɪɚ-
ɮɢɧɢɡɢɪɨɜɚɥɢɩɪɨɜɨɞɹɢɯɱɟɪɟɡɫɟɪɢɸɤɫɢ-
ɥɨɥɨɜ ɉɪɟɩɚɪɚɬɵ ɨɤɪɚɲɢɜɚɥɢ ɝɟɦɚɬɨɤɫɢ-
ɥɢɧɨɦɩɨɗɪɥɢɯɭɫɩɨɞɤɪɚɫɤɨɣɚɥɰɢɚɧɨɜɵɦ
ɫɢɧɢɦɋɪɟɡɵɡɚɤɥɸɱɚɥɢɜɦɨɜɢɨɥɢɩɨɤɪɵ-
ɜɚɥɢ ɩɨɤɪɨɜɧɵɦ ɫɬɟɤɥɨɦ ɉɪɨɫɦɨɬɪ ɩɨɫɬɨ-
ɹɧɧɵɯɩɪɟɩɚɪɚɬɨɜɩɪɨɜɨɞɢɥɢɧɚɦɢɤɪɨɫɤɨɩɟ
$[LRSODQHPRW&DUO=HLVVɞɥɹɞɨɤɭɦɟɧɬɢ-
ɪɨɜɚɧɢɹ ɨɛɴɟɤɬɨɜ ɢɫɩɨɥɶɡɨɜɚɥɢ ɜɢɞɟɨɤɚɦɟ-
ɪɭ $[LR&DP 95F ɢɩɪɨɝɪɚɦɦɭ $[LR9LVLRQ
4.8&DUO=HLVV
Ɋɟɡɭɥɶɬɚɬɵ ɢɨɛɫɭɠɞɟɧɢɟȾɥɹɜɵɹɜ-
ɥɟɧɢɹɩɪɢɱɢɧɮɨɪɦɢɪɨɜɚɧɢɹɛɨɥɶɲɨɝɨɱɢɫ-
ɥɚɧɟɠɢɡɧɟɫɩɨɫɨɛɧɵɯɞɢɚɫɩɨɪɭ2SXUSXUHD
VVS caucasica ɩɪɨɢɡɪɚɫɬɚɸɳɢɯ ɜɇɨɜɨɪɨɫ-
ɫɢɣɫɤɨɦɢȺɛɢɧɫɤɨɦɪɚɣɨɧɚɯɄɪɚɫɧɨɞɚɪɫɤɨ-
ɝɨɤɪɚɹɛɵɥɨɩɪɨɜɟɞɟɧɨɰɢɬɨɷɦɛɪɢɨɥɨɝɢɱɟ-
ɫɤɨɟɢɫɫɥɟɞɨɜɚɧɢɟɫɟɦɹɧɧɚɪɚɡɧɵɯɫɬɚɞɢɹɯ
ɢɯɪɚɡɜɢɬɢɹ
Ɋɚɡɛɨɪ ɫɨɰɜɟɬɢɣ ɫɩɥɨɞɚɦɢ ɩɨɤɚɡɚɥ
ɱɬɨ ɱɚɫɬɶ ɩɥɨɞɨɜ ɨɛɪɚɡɨɜɚɥɚɫɶ ɜɪɟɡɭɥɶɬɚ-
ɬɟ ɫɜɨɛɨɞɧɨɝɨ ɨɩɵɥɟɧɢɹ ɍɛɨɥɶɲɢɧɫɬɜɚ
ɠɟɪɚɫɬɟɧɢɣɭɱɚɫɬɜɭɸɳɢɯɜɷɤɫɩɟɪɢɦɟɧɬɟ
ɩɥɨɞɵ ɡɚɜɹɡɚɥɢɫɶ ɬɨɥɶɤɨ ɜɪɟɡɭɥɶɬɚɬɟ ɢɫ
ɤɭɫɫɬɜɟɧɧɨɝɨɨɩɵɥɟɧɢɹɗɬɨɫɜɢɞɟɬɟɥɶɫɬɜɭ-
ɟɬ ɨɬɨɦ ɱɬɨ ɧɢɡɤɢɣ ɩɪɨɰɟɧɬ ɡɚɜɹɡɵɜɚɧɢɹ
ɩɥɨɞɨɜ ɫɜɹɡɚɧ ɫɨɬɫɭɬɫɬɜɢɟɦ ɧɚɫɟɤɨɦɵɯ
ɨɩɵɥɢɬɟɥɟɣɢɥɢɛɨɥɟɟɜɟɪɨɹɬɧɨɫɢɯɧɢɡɤɨɣ
ɚɤɬɢɜɧɨɫɬɶɸɜɫɜɹɡɢɫɧɟɨɩɬɢɦɚɥɶɧɵɦɢɩɨ
ɝɨɞɧɵɦɢɭɫɥɨɜɢɹɦɢɜɩɟɪɢɨɞɰɜɟɬɟɧɢɹ
ɐɢɬɨɷɦɛɪɢɨɥɨɝɢɱɟɫɤɨɟ ɢɫɫɥɟɞɨɜɚ-
ɧɢɟɧɟɡɪɟɥɵɯ ɫɟɦɹɧ ɩɨɤɚɡɚɥɨɱɬɨɡɚɪɨɞɵɲ
ɜɛɨɥɶɲɢɧɫɬɜɟ ɢɡ ɧɢɯ ɩɪɢɫɭɬɫɬɜɭɟɬ Ɉɞɧɚ-
ɤɨ ɤɦɨɦɟɧɬɭ ɞɢɫɫɟɦɢɧɚɰɢɢ ɛyɥɶɲɚɹ ɱɚɫɬɶ
ɡɪɟɥɵɯ ɫɟɦɹɧ ɨɤɚɡɵɜɚɟɬɫɹ ɛɟɡ ɡɚɪɨɞɵɲɚ
ɂɡɷɬɨɝɨ ɫɥɟɞɭɟɬ ɱɬɨ ɨɩɥɨɞɨɬɜɨɪɟɧɢɟ ɩɪɨ-
ɢɫɯɨɞɢɬɜɛɨɥɶɲɟɣɱɚɫɬɢɫɟɦɹɡɚɱɚɬɤɨɜɢɱɬɨ
ɡɚɪɨɞɵɲɜɧɢɯɪɚɡɜɢɜɚɟɬɫɹɁɚɬɟɦɧɚɤɚɤɨɦ
ɬɨɷɬɚɩɟɫɟɦɟɧɧɨɝɨɪɚɡɜɢɬɢɹɩɪɨɢɫɯɨɞɢɬɞɟ-
ɈɊɂȽɂɇȺɅɖɇɕȿɋɌȺɌɖɂ,661ɇɚɭɤɚɄɭɛɚɧɢʋ
16
ɝɟɧɟɪɚɰɢɹɡɚɪɨɞɵɲɚɢɨɧɫɨɯɪɚɧɹɟɬɫɹɬɨɥɶ-
ɤɨɜɧɟɛɨɥɶɲɨɣɱɚɫɬɢɫɟɦɹɧ
ɐɢɬɨɷɦɛɪɢɨɥɨɝɢɱɟɫɤɨɟ ɢɡɭɱɟɧɢɟ ɡɚ-
ɪɨɞɵɲɟɣ ɧɟɡɪɟɥɵɯ ɫɟɦɹɧ 2 SXUSXUHD VVS
caucasica ɜɵɹɜɢɥɨ ɜɢɯ ɫɬɪɨɟɧɢɢ ɧɚɥɢɱɢɟ
ɦɧɨɝɨɱɢɫɥɟɧɧɵɯ ɨɬɤɥɨɧɟɧɢɣ ɨɬ ɧɨɪɦɵ
ɪɢɫ ± ȼɧɨɪɦɟ ɩɟɪɟɝɨɪɨɞɤɢ ɩɪɢ ɞɟɥɟ-
ɧɢɢɤɥɟɬɨɤɬɟɥɚ ɡɚɪɨɞɵɲɚɢɫɭɫɩɟɧɡɨɪɚɡɚ-
ɤɥɚɞɵɜɚɸɬɫɹ ɨɩɪɟɞɟɥɟɧɧɵɦ ɨɛɪɚɡɨɦ ɱɬɨ
ɨɩɪɟɞɟɥɹɟɬ ɪɟɝɭɥɹɪɧɵɣ ɬɢɩ ɷɦɛɪɢɨɝɟɧɟɡɚ
ɢɧɨɪɦɚɥɶɧɵɣ ɯɨɞ ɷɦɛɪɢɨɧɚɥɶɧɨɝɨ ɪɚɡ-
ɜɢɬɢɹ ɂɡɦɟɧɟɧɢɹ ɜɧɚɩɪɚɜɥɟɧɢɢ ɡɚɤɥɚɞɤɢ
ɩɟɪɟɝɨɪɨɞɨɤɩɪɢ ɤɥɟɬɨɱɧɵɯɞɟɥɟɧɢɹɯ ɢɧɚ-
ɪɭɲɟɧɢɹɜɩɨɫɥɟɞɨɜɚɬɟɥɶɧɨɫɬɢɢɬɟɦɩɚɯɞɟ-
ɥɟɧɢɹ ɤɥɟɬɨɤ ɬɟɥɚ ɡɚɪɨɞɵɲɚ ɢɫɭɫɩɟɧɡɨɪɚ
ɫɜɢɞɟɬɟɥɶɫɬɜɭɸɬ ɨɛ ɚɧɨɦɚɥɶɧɨɦ ɪɚɡɜɢɬɢɢ
ɡɚɪɨɞɵɲɚɢɨɛɚɧɨɦɚɥɶɧɨɦɬɟɱɟɧɢɢɩɪɨɰɟɫ-
ɫɚɷɦɛɪɢɨɝɟɧɟɡɚ
ȼɧɨɪɦɟ ɪɚɡɜɢɬɢɟ ɡɚɪɨɞɵɲɚ ɞɚɧɧɨɝɨ
ɬɚɤɫɨɧɚ ɞɨɥɠɧɨ ɩɪɨɢɫɯɨɞɢɬɶ ɫɥɟɞɭɸɳɢɦ
ɨɛɪɚɡɨɦ ɉɨɫɥɟ ɨɩɥɨɞɨɬɜɨɪɟɧɢɹ ɡɢɝɨɬɚ ɞɟ-
ɥɢɬɫɹ ɩɨɩɟɪɟɱɧɨɣ ɩɟɪɟɝɨɪɨɞɤɨɣ ɩɪɢ ɷɬɨɦ
ɨɛɪɚɡɭɸɬɫɹɬɟɪɦɢɧɚɥɶɧɚɹɚɩɢɤɚɥɶɧɚɹɢɛɚ-
ɡɚɥɶɧɚɹ ɤɥɟɬɤɢ ɡɚɪɨɞɵɲɚ ɂɡ ɩɪɨɢɡɜɨɞɧɵɯ
ɚɩɢɤɚɥɶɧɨɣ ɤɥɟɬɤɢ ɮɨɪɦɢɪɭɟɬɫɹ ɬɟɥɨ ɡɚ-
ɪɨɞɵɲɚ ɬɨɝɞɚ ɤɚɤ ɢɡ ɞɟɪɢɜɚɬɨɜ ɛɚɡɚɥɶɧɨɣ
ɤɥɟɬɤɢ²ɫɩɟɰɢɚɥɶɧɵɣɨɪɝɚɧɫɭɫɩɟɧɡɨɪ
Ɋɨɥɶɫɭɫɩɟɧɡɨɪɚɡɚɤɥɸɱɚɟɬɫɹɜɨɛɟɫɩɟ-
ɱɟɧɢɢ ɩɨɫɬɭɩɥɟɧɢɹ ɩɢɬɚɬɟɥɶɧɵɯ ɜɟɳɟɫɬɜ
ɢɡ ɬɤɚɧɟɣ ɦɚɬɟɪɢɧɫɤɨɝɨ ɨɪɝɚɧɢɡɦɚ ɤɪɚɡɜɢ-
ɜɚɸɳɟɦɭɫɹɡɚɪɨɞɵɲɭɋɬɪɨɟɧɢɟɫɭɫɩɟɧɡɨɪɚ
ɹɜɥɹɟɬɫɹ ɯɚɪɚɤɬɟɪɧɵɦ ɩɪɢɡɧɚɤɨɦ ɬɚɤɫɨɧɨɜ
ɪɚɡɧɨɝɨɪɚɧɝɚ²ɤɚɤɰɟɥɨɝɨɩɨɪɹɞɤɚɢɥɢɨɬ-
ɞɟɥɶɧɨɝɨɫɟɦɟɣɫɬɜɚɬɚɤɢɪɨɞɚɢɜɢɞɚ>@
ȼɧɨɪɦɟ ɩɪɢ ɨɛɪɚɡɨɜɚɧɢɢ ɫɭɫɩɟɧɡɨɪɚ
ɜɫɥɭɱɚɟɩɪɟɞɫɬɚɜɢɬɟɥɟɣɪɨɞɚOrchisɤɥɟɬɤɢ
ɞɨɥɠɧɵɞɟɥɢɬɶɫɹɬɨɥɶɤɨɩɨɩɟɪɟɱɧɵɦɢɩɟɪɟ-
ɝɨɪɨɞɤɚɦɢ ɢɨɧ ɞɨɥɠɟɧɫɨɫɬɨɹɬɶ ɢɡ ɨɞɧɨɝɨ
ɪɹɞɚɤɥɟɬɨɤ>@ɋɮɨɪɦɢɪɨɜɚɧɧɵɣɫɭɫɩɟɧ-
ɡɨɪ ɭɩɪɟɞɫɬɚɜɢɬɟɥɟɣ ɪɨɞɚ Orchis ɧɢɬɟɜɢɞ-
ɧɵɣ ɫɨɫɬɨɢɬ ɢɡ ɨɞɧɨɪɨɞɧɵɯ ɜɚɤɭɨɥɢɡɢɪɨ-
ɜɚɧɧɵɯɜɵɬɹɧɭɬɵɯɩɨɚɩɢɤɚɥɶɧɨɛɚɡɚɥɶɧɨɣ
ɨɫɢ ɨɞɧɨɹɞɟɪɧɵɯ ɤɥɟɬɨɤ >@ Ɍɚɤɨɣ ɠɟ ɬɢɩ
ɫɭɫɩɟɧɡɨɪɚ ɯɚɪɚɤɬɟɪɟɧ ɧɟ ɬɨɥɶɤɨ ɞɥɹ ɩɪɟɞ-
ɫɬɚɜɢɬɟɥɟɣɪɨɞɚOrchisɧɨɢɞɥɹɩɪɟɞɫɬɚɜɢ-
ɬɟɥɟɣɪɨɞɚDactylorhiza>@ɢɞɪɭɝɢɯɬɚɤɫɨɧɨɜ
ɩɨɞɬɪɢɛɵOrchidoidea>@ɍO purpureaɫɭ-
ɫɩɟɧɡɨɪɫɨɫɬɨɢɬɤɚɤɩɪɚɜɢɥɨɢɡ±ɤɥɟɬɨɤ
Ɉɧ ɜɵɯɨɞɢɬ ɱɟɪɟɡ ɦɢɤɪɨɩɢɥɟ ɢɜɧɟɞɪɹɟɬɫɹ
ɜɬɤɚɧɢɩɥɚɰɟɧɬɵɡɚɜɹɡɢ>11].
ɋɬɪɨɟɧɢɟ ɡɚɪɨɞɵɲɚ ɬɚɤɠɟ ɹɜɥɹɟɬɫɹ
ɷɜɨɥɸɰɢɨɧɧɨɡɧɚɱɢɦɵɦɢɬɚɤɫɨɧɫɩɟɰɢɮɢɱ-
ɧɵɦɩɪɢɡɧɚɤɨɦ>@
ɉɟɪɜɨɟ ɞɟɥɟɧɢɟ ɬɟɪɦɢɧɚɥɶɧɨɣ ɤɥɟɬ-
ɤɢ ɞɜɭɤɥɟɬɨɱɧɨɝɨ ɡɚɪɨɞɵɲɚ ɭɩɪɟɞɫɬɚɜɢɬɟ-
ɥɟɣ ɪɨɞɚ Orchis ɤɚɤ ɩɪɚɜɢɥɨ ɩɪɨɞɨɥɶɧɨɟ
ɩɨ ɨɬɧɨɲɟɧɢɸ ɤɚɩɢɤɚɥɶɧɨɛɚɡɚɥɶɧɨɣ ɨɫɢ
ɡɚɪɨɞɵɲɚ ȼɧɨɪɦɟ ɷɬɨ ɞɟɥɟɧɢɟ ɩɪɢɜɨɞɢɬ
ɤɨɛɪɚɡɨɜɚɧɢɸɞɜɭɯɤɥɟɬɨɤɨɞɢɧɚɤɨɜɨɝɨɪɚɡ-
ɦɟɪɚ ɉɨɫɥɟɞɭɸɳɟɟ ɞɟɥɟɧɢɟ ɩɪɨɢɫɯɨɞɢɬ
ɤɚɤ ɩɪɚɜɢɥɨ ɫɮɨɪɦɢɪɨɜɚɧɢɟɦ ɩɨɩɟɪɟɱɧɨɣ
ɩɟɪɟɝɨɪɨɞɤɢɉɨɦɟɪɟɞɟɥɟɧɢɹɤɥɟɬɨɤɨɛɪɚ-
ɡɭɟɬɫɹ ɝɥɨɛɭɥɹɪɧɵɣ ɡɚɪɨɞɵɲ Ⱦɚɥɟɟ ɤɪɨɦɟ
ɩɪɨɞɨɥɠɚɸɳɢɯɫɹɞɟɥɟɧɢɣɤɥɟɬɨɤɧɚɛɥɸɞɚ-
ɟɬɫɹ ɝɢɫɬɨɥɨɝɢɱɟɫɤɚɹ ɞɢɮɮɟɪɟɧɰɢɚɰɢɹ ɬɤɚ-
ɧɟɣ² ɜɵɱɥɟɧɟɧɢɟ ɩɪɨɬɨɞɟɪɦɵ ɩɨɤɪɨɜɧɚɹ
ɬɤɚɧɶɢɤɥɟɬɨɤɜɧɭɬɪɟɧɧɟɣɩɚɪɟɧɯɢɦɵ ɡɚ-
ɩɚɫɚɸɳɚɹ ɬɤɚɧɶ ɨɛɪɚɡɭɟɬɫɹ ɷɦɛɪɢɨɧɚɥɶ-
ɧɵɣ ɚɩɟɤɫ ɩɨɛɟɝɚ ɛɟɡ ɩɪɢɡɧɚɤɨɜ ɡɚɱɚɬɤɨɜ
ɥɚɬɟɪɚɥɶɧɵɯɨɪɝɚɧɨɜ
Ⱥɧɨɦɚɥɢɢ ɩɪɢ ɪɚɡɜɢɬɢɢ ɡɚɪɨɞɵɲɚ
ɭ2 SXUSXUHD VVS caucasica ɩɪɨɹɜɥɹɥɢɫɶ
ɧɚɫɚɦɵɯ ɪɚɧɧɢɯ ɷɬɚɩɚɯ ɟɝɨ ɪɚɡɜɢɬɢɹ ɉɪɟ-
ɠɞɟ ɜɫɟɝɨ ɧɚɛɥɸɞɚɥɢɫɶ ɡɚɞɟɪɠɤɢ ɞɟɥɟɧɢɣ
ɧɟɤɨɬɨɪɵɯ ɤɥɟɬɨɤ ɫɨɛɫɬɜɟɧɧɨ ɬɟɥɚ ɡɚɪɨɞɵ-
ɲɚɪɢɫɗɬɨɜɩɨɫɥɟɞɫɬɜɢɢɩɪɢɜɨɞɢɥɨ
ɤɪɚɡɞɟɥɟɧɢɸ ɬɟɥɚ ɡɚɪɨɞɵɲɚ ɧɚ ɞɟɥɹɳɢɟɫɹ
ɢɥɢ ɧɟɞɟɥɹɳɢɟɫɹ ɤɥɚɫɬɟɪɵ ɤɥɟɬɨɤ ɪɢɫ 
Ⱥɧɨɦɚɥɶɧɵɦ ɬɚɤɠɟ ɹɜɥɹɥɨɫɶ ɞɟɥɟɧɢɟ ɬɟɪ-
ɦɢɧɚɥɶɧɨɣ ɤɥɟɬɤɢ ɩɪɨɷɦɛɪɢɨ ɧɚɤɥɨɧɧɨɣ
ɩɟɪɟɝɨɪɨɞɤɨɣɱɬɨɩɪɢɜɨɞɢɥɨɤɨɛɪɚɡɨɜɚɧɢɸ
ɞɜɭɯɤɥɟɬɨɤɧɟɨɞɢɧɚɤɨɜɨɝɨɪɚɡɦɟɪɚɪɢɫ
Ⱥɧɨɦɚɥɢɢ ɩɪɢ ɮɨɪɦɢɪɨɜɚɧɢɢ ɫɭɫɩɟɧ-
ɡɨɪɚ ɩɪɨɹɜɥɹɥɢɫɶ ɨɞɧɨɜɪɟɦɟɧɧɨ ɫɧɚɪɭ-
ɲɟɧɢɹɦɢ ɜɵɹɜɥɹɟɦɵɦɢ ɩɪɢ ɨɛɪɚɡɨɜɚɧɢɢ
ɫɨɛɫɬɜɟɧɧɨɬɟɥɚɡɚɪɨɞɵɲɚɢɥɢɠɟɧɟɛɵɥɢ
ɫɜɹɡɚɧɵɫɧɢɦɢɈɧɢɨɛɧɚɪɭɠɢɜɚɥɢɫɶɫɪɚɜ
ɧɢɬɟɥɶɧɨɩɨɡɞɧɨɜɷɦɛɪɢɨɝɟɧɟɡɟɇɚɩɪɢɦɟɪ
ɩɪɢ ɜɧɟɲɧɟ ɧɨɪɦɚɥɶɧɨɣ ɝɢɫɬɨɥɨɝɢɱɟɫɤɨɣ
ɞɢɮɮɟɪɟɧɰɢɚɰɢɢ ɬɟɥɚ ɦɧɨɝɨɤɥɟɬɨɱɧɨɝɨ ɡɚ-
ɪɨɞɵɲɚ ɧɚɛɥɸɞɚɥɨ ɫɶ ɚɧɨɦɚɥɶɧɨɟ ɪɚɡɪɚɫɬɚ-
ɧɢɟ ɨɬɞɟɥɶɧɵɯ ɤɥɟɬɨɤ ɫɭɫɩɟɧɡɨɪɚ ɮɨɪɦɢ-
ɪɨɜɚɧɢɟ ɬɚɤ ɧɚɡɵɜɚɟɦɵɯ ©ɩɭɡɵɪɟɜɢɞɧɵɯª
ɤɥɟɬɨɤɪɢɫ
ɋɟɥɶɫɤɨɯɨɡɹɣɫɬɜɟɧɧɵɟɧɚɭɤɢ ,661ɇɚɭɤɚɄɭɛɚɧɢʋ 17
Ɋɢɫɋɬɪɨɟɧɢɟɡɚɪɨɞɵɲɚ2UFKLVSXUSXUHDVXEVSFDXFDVLFD2UFKLGDFHDH
ɧɚɩɪɨɞɨɥɶɧɵɯɫɪɟɡɚɯ
Ɉɤɪɚɫɤɚɝɟɦɚɬɨɤɫɢɥɢɧɨɦɩɨɗɪɥɢɯɭɫɩɨɞɤɪɚɫɤɨɣɚɥɰɢɚɧɨɜɵɦɫɢɧɢɦ
Ɇɚɫɲɬɚɛɧɵɟɥɢɧɟɣɤɢɦɤɦɨɞɢɧɚɤɨɜɵɟɦɚɫɲɬɚɛɧɵɟɥɢɧɟɣɤɢ
ɞɥɹɞɟɬɚɥɟɣ±ɢɞɥɹɞɟɬɚɥɟɣ
ɫɭɫ²ɫɭɫɩɟɧɡɨɪɩɤɫɭɫ²©ɩɭɡɵɪɟɜɢɞɧɵɟɤɥɟɬɤɢɫɭɫɩɟɧɡɨɪɚ
Ⱥɧɨɦɚɥɶɧɵɟ ɡɚɪɨɞɵɲɢ ɯɚɪɚɤɬɟɪɢɡɨ-
ɜɚɥɢɫɶ ɡɚɦɟɞɥɟɧɧɵɦɢ ɬɟɦɩɚɦɢ ɪɚɡɜɢɬɢɹ
ɩɨ ɫɪɚɜɧɟɧɢɸ ɫɧɨɪɦɚɥɶɧɵɦɢ ɡɚɪɨɞɵɲɚ-
ɦɢɇɚɫɬɚɞɢɢɫɮɨɪɦɢɪɨɜɚɧɧɵɯɫɟɦɹɧɦɟɥ-
ɤɢɟ ɡɚɪɨɞɵɲɢ ɢɦɟɥɢ ɩɪɢɡɧɚɤɢ ɞɟɝɟɧɟɪɚ-
ɰɢɢ ɚɤɦɨɦɟɧɬɭ ɞɢɫɫɟɦɢɧɚɰɢɢ ɨɧɢ ɩɨɱɬɢ
ɩɨɥɧɨɫɬɶɸ ɞɟɝɟɧɟɪɢɪɨɜɚɥɢ Ɏɨɪɦɢɪɨɜɚɧɢɟ
ɫɟɦɟɧɧɨɣ ɤɨɠɭɪɵɩɨɜɢɞɢɦɨɦɭ ɨɫɭɳɟɫɬɜ
ɥɹɥɨɫɶ ɛɟɡ ɫɭɳɟɫɬɜɟɧɧɵɯ ɨɬɤɥɨɧɟɧɢɣ ɨɬ
ɧɨɪɦɵ ɬɟɦ ɫɚɦɵɦ ɫɟɦɟɧɚ ɢɫɡɚɪɨɞɵɲɟɦ
ɢɛɟɡɡɚɪɨɞɵɲɚɩɪɢɨɛɪɟɬɚɥɢɪɚɡɦɟɪɵɢɮɨɪ-
ɦɭ ɯɚɪɚɤɬɟɪɧɭɸ ɞɥɹ ɫɟɦɹɧ ɡɪɟɥɵɯ ɩɥɨɞɨɜ
ɢɡɭɱɟɧɧɨɝɨ ɜɢɞɚ ɨɪɯɢɞɧɵɯ ɉɨ ɜɫɟɣ ɜɢɞɢ-
ɦɨɫɬɢ ɚɧɨɦɚɥɶɧɵɟ ɡɚɪɨɞɵɲɢ ɨɫɬɚɜɚɥɢɫɶ
ɠɢɡɧɟɫɩɨɫɨɛɧɵɦɢɞɨɩɟɪɟɯɨɞɚɫɟɦɹɧɜɫɬɚ-
ɞɢɸ ɫɨɡɪɟɜɚɧɢɹ ɬɨɟɫɬɶ ɩɨɤɚ ɨɧɢɨɛɟɫɩɟ
ɱɢɜɚɥɢɫɶɩɢɬɚɬɟɥɶɧɵɦɢɜɟɳɟɫɬɜɚɦɢ ɨɬɦɚ-
ɬɟɪɢɧɫɤɨɝɨɨɪɝɚɧɢɡɦɚȼɩɪɨɰɟɫɫɟɮɨɪɦɢɪɨ-
ɜɚɧɢɹɫɟɦɟɧɧɨɣɤɨɠɭɪɵɢɫɨɡɪɟɜɚɧɢɹɫɟɦɹɧ
ɚɧɨɦɚɥɶɧɵɟ ɡɚɪɨɞɵɲɢ ɧɟ ɛɵɥɢ ɫɩɨɫɨɛɧɵ
ɈɊɂȽɂɇȺɅɖɇɕȿɋɌȺɌɖɂ,661ɇɚɭɤɚɄɭɛɚɧɢʋ
18
ɤɫɚɦɨɫɬɨɹɬɟɥɶɧɨɦɭ ɫɭɳɟɫɬɜɨɜɚɧɢɸ ɢɩɨ-
ɝɢɛɚɥɢ
ɉɪɢɱɢɧɵ ɩɪɢɜɨɞɹɳɢɟ ɤɮɨɪɦɢɪɨɜɚ-
ɧɢɸ ɧɟɠɢɡɧɟɫɩɨɫɨɛɧɵɯ ɫɟɦɹɧ ɦɨɝɭɬ ɛɵɬɶ
ɪɚɡɥɢɱɧɵɦɢ Ʉɚɤ ɫɱɢɬɚɥɚ Ɋȿ Ʌɟɜɢɧɚ >@
ɨɫɧɨɜɧɵɟ ɮɚɤɬɨɪɵ ɨɩɪɟɞɟɥɹɸɳɢɟ ɧɢɡɤɨɟ
ɤɚɱɟɫɬɜɨ ɫɟɦɹɧ ɦɨɝɭɬ ɛɵɬɶ ɨɛɴɟɞɢɧɟɧɵ
ɜ3ɝɪɭɩɩɵɧɟɞɨɫɬɚɬɨɱɧɨɫɬɶ ɨɩɵɥɟɧɢɹ ɧɟ-
ɯɜɚɬɤɚ ɪɟɫɭɪɫɨɜ ɩɨɜɪɟɠɞɟɧɢɟ ɫɟɦɹɧ ɮɢɬɨ-
ɮɚɝɚɦɢ ɢɛɨɥɟɡɧɹɦɢ ȼɫɟ ɝɪɭɩɩɵ ɮɚɤɬɨɪɨɜ
ɧɚɯɨɞɹɬɫɹɜɫɥɨɠɧɨɣɩɪɹɦɨɣɢɨɩɨɫɪɟɞɨɜɚɧ-
ɧɨɣɡɚɜɢɫɢɦɨɫɬɢɨɬɚɛɢɨɬɢɱɟɫɤɢɯɬɟɦɩɟɪɚ-
ɬɭɪɚ ɜɥɚɠɧɨɫɬɶ ɢɛɢɨɬɢɱɟɫɤɢɯ ɮɚɤɬɨɪɨɜ
ɚɤɬɢɜɧɨɫɬɶ ɢɤɨɥɟɛɚɧɢɟɱɢɫɥɟɧɧɨɫɬɢ ɧɚɫɟ-
ɤɨɦɵɯɬɢɩɨɩɵɥɟɧɢɹ>@
Ɏɟɧɨɦɟɧ ɝɟɧɟɬɢɱɟɫɤɨɣ ɧɟɫɬɚɛɢɥɶɧɨɫ
ɬɢ ɜɵɹɜɥɹɟɦɵɣ ɩɪɢ ɪɚɡɜɢɬɢɢ ɡɚɪɨɞɵɲɚ
ɢɩɪɢɜɨɞɹɳɢɣɤɧɟɠɢɡɧɟɫɩɨɫɨɛɧɨɫɬɢɫɟɦɹɧ
ɯɨɪɨɲɨɢɡɜɟɫɬɟɧɢɡɪɟɡɭɥɶɬɚɬɨɜɷɤɫɩɟɪɢɦɟɧ-
ɬɚɥɶɧɨɣ ɪɚɛɨɬɵ ɩɨ ɩɨɥɭɱɟɧɢɸ ɢɫɤɭɫɫɬɜɟɧ-
ɧɵɯ ɦɟɠɜɢɞɨɜɵɯ ɝɢɛɪɢɞɨɜ ɢɬɪɚɧɫɝɟɧɧɵɯ
ɪɚɫɬɟɧɢɣɚɬɚɤɠɟɭɝɚɩɥɨɢɞɨɜɈɞɧɚɤɨɝɟɧɟ-
ɬɢɱɟɫɤɚɹɧɟɫɬɚɛɢɥɶɧɨɫɬɶɦɨɠɟɬɩɪɨɹɜɥɹɬɶɫɹ
ɢɜɩɪɢɪɨɞɧɵɯ ɩɨɩɭɥɹɰɢɹɯ ɪɚɫɬɟɧɢɣ ɉɪɨ-
ɜɟɞɟɧɧɨɟ ɧɚɦɢ ɰɢɬɨɷɦɛɪɢɨɥɨɝɢɱɟɫɤɨɟ ɢɡ-
ɭɱɟɧɢɟ ɫɟɦɹɧ 2 SXUSXUHD VVS caucasicaɧɚ
ɪɚɡɧɵɯ ɫɬɚɞɢɹɯ ɢɯ ɪɚɡɜɢɬɢɹ ɩɨɡɜɨɥɹɟɬ ɫɞɟ-
ɥɚɬɶɜɵɜɨɞɨɬɨɦɱɬɨɜɵɹɜɥɟɧɧɵɟɚɧɨɦɚɥɢɢ
ɪɚɡɜɢɬɢɹ ɢɫɬɪɨɟɧɢɹ ɡɚɪɨɞɵɲɟɣ ɹɜɥɹɸɬɫɹ
ɥɟɬɚɥɶɧɵɦɢɢɢɦɟɧɧɨɨɧɢɩɪɢɜɨɞɹɬɤɬɨɦɭ
ɱɬɨɛyɥɶɲɚɹɱɚɫɬɶɫɟɦɹɧɫɬɚɧɨɜɢɬɫɹɧɟɠɢɡ-
ɧɟɫɩɨɫɨɛɧɵɦɢ ɤɦɨɦɟɧɬɭ ɞɢɫɫɟɦɢɧɚɰɢɢ
Ɍɚɤɢɦ ɨɛɪɚɡɨɦ ɧɟɩɨɥɧɨɰɟɧɧɨɫɬɶ ɛɨɥɶɲɟɣ
ɱɚɫɬɢɫɟɦɹɧɭɢɡɭɱɟɧɧɵɯɨɫɨɛɟɣ2SXUSXUHD
VVSFDXFDVLFDɛɵɥɚɜɵɡɜɚɧɚɚɧɨɦɚɥɢɹɦɢɪɚɡ-
ɜɢɬɢɹɡɚɪɨɞɵɲɚɚɧɟɧɟɞɨɨɩɵɥɟɧɢɟɦɰɜɟɬ-
ɤɨɜ ɋɬɪɭɤɬɭɪɧɵɟ ɢɡɦɟɧɟɧɢɹ ɧɚɛɥɸɞɚɟɦɵɟ
ɭɚɧɨɦɚɥɶɧɵɯɡɚɪɨɞɵɲɟɣ²ɨɬɫɭɬɫɬɜɢɟ ɪɟ-
ɝɭɥɹɪɧɨɫɬɢɞɟɥɟɧɢɹ ɤɥɟɬɨɤɩɪɢ ɮɨɪɦɢɪɨɜɚ-
ɧɢɢɬɟɥɚɡɚɪɨɞɵɲɚɢɯɚɨɬɢɱɧɨɟɪɚɫɩɨɥɨɠɟ-
ɧɢɟɩɟɪɟɝɨɪɨɞɨɤɩɪɢɢɯɞɟɥɟɧɢɢɢɡɦɟɧɟɧɢɟ
ɪɚɡɦɟɪɨɜ ɢɮɨɪɦɵ ɤɥɟɬɨɤ ɫɭɫɩɟɧɡɨɪɚ ɡɚ-
ɞɟɪɠɤɚ ɞɟɥɟɧɢɣ ɨɬɞɟɥɶɧɵɯ ɤɥɟɬɨɤ² ɫɜɢ-
ɞɟɬɟɥɶɫɬɜɭɸɬ ɨɝɟɧɟɬɢɱɟɫɤɢɯ ɧɚɪɭɲɟɧɢɹɯ
ɧɨɪɦɚɥɶɧɨɝɨɯɨɞɚɷɦɛɪɢɨɧɚɥɶɧɨɝɨɪɚɡɜɢɬɢɹ
ɚɧɟɨɧɟɯɜɚɬɤɟɪɟɫɭɪɫɨɜɞɥɹɪɚɡɜɢɬɢɹɦɨɥɨ-
ɞɨɝɨɫɩɨɪɨɮɢɬɚ
Ʉɜɵɲɟɭɤɚɡɚɧɧɵɦ ɩɪɢɱɢɧɚɦ ɩɪɢɜɨ-
ɞɹɳɢɦ ɤɪɚɡɧɨɤɚɱɟɫɬɜɟɧɧɨɫɬɢ ɫɟɦɹɧ ɭɤɚ-
ɡɚɧɧɵɦɜɪɚɛɨɬɟɊȿɅɟɜɢɧɨɣ>@ɤɨɬɨɪɵɟ
ɩɨɜɢɞɢɦɨɦɭ ɜɫɟɝɞɚɢɦɟɸɬɦɟɫɬɨɭɥɸɛɨɝɨ
ɰɜɟɬɤɨɜɨɝɨ ɪɚɫɬɟɧɢɹ ɫɥɟɞɭɟɬ ɞɨɛɚɜɢɬɶ ɟɳɟ
ɢɬɟɤɨɬɨɪɵɟɩɪɢɜɨɞɹɬɤɩɪɨɹɜɥɟɧɢɸɝɟɧɟ-
ɬɢɱɟɫɤɨɣ ɧɟɫɬɚɛɢɥɶɧɨɫɬɢ ɧɚ ɪɚɧɧɢɯ ɷɬɚɩɚɯ
ɪɚɡɜɢɬɢɹ ɦɨɥɨɞɨɝɨ ɫɩɨɪɨɮɢɬɚ ɜɩɟɪɢɨɞ ɟɝɨ
ɫɟɦɟɧɧɨɝɨɪɚɡɜɢɬɢɹɄɚɤɫɥɟɞɭɟɬɢɡɪɟɡɭɥɶɬɚ-
ɬɨɜɩɨɥɭɱɟɧɧɵɯɜɨɛɥɚɫɬɢɷɤɫɩɟɪɢɦɟɧɬɚɥɶ-
ɧɨɣ ɝɟɧɟɬɢɤɢ ɷɦɛɪɢɨɥɨɝɢɢ ɢɛɢɨɬɟɯɧɨɥɨ-
ɝɢɢɩɪɢɡɧɚɤɢɝɟɧɟɬɢɱɟɫɤɨɣɧɟɫɬɚɛɢɥɶɧɨɫɬɢ
ɦɨɝɭɬ ɛɵɬɶ ɫɥɟɞɫɬɜɢɟɦ ɧɟɫɤɨɥɶɤɢɯ ɪɚɡɧɵɯ
ɦɟɯɚɧɢɡɦɨɜ
 ɂɡɡɚ ɧɟɫɛɚɥɚɧɫɢɪɨɜɚɧɧɨɫɬɢ ɝɟɧɨ-
ɬɢɩɚɭɦɟɠɜɢɞɨɜɵɯɝɢɛɪɢɞɨɜɢɥɢɭɧɟɤɨɬɨ-
ɪɵɯɨɪɝɚɧɢɡɦɨɜɫɢɡɦɟɧɟɧɧɨɣɩɥɨɢɞɧɨɫɬɶɸ
ɇɚɩɪɢɦɟɪɭɦɟɠɜɢɞɨɜɵɯɝɢɛɪɢɞɨɜɫɧɚɱɚɥɚ
ɧɚɛɥɸɞɚɟɬɫɹ ɡɚɞɟɪɠɤɚ ɪɚɡɜɢɬɢɹ ɡɚɪɨɞɵ-
ɲɚ ɧɚ ɪɚɧɧɢɯ ɫɬɚɞɢɹɯ ɟɝɨ ɪɚɡɜɢɬɢɹ ɚɡɚ-
ɬɟɦ² ɩɨɥɧɚɹ ɨɫɬɚɧɨɜɤɚ ɪɨɫɬɚ ɢɪɚɡɜɢɬɢɹ
ɧɚ ɝɥɨɛɭɥɹɪɧɨɣ ɫɬɚɞɢɢ Ʉɦɨɦɟɧɬɭ ɞɢɫɫɟ-
ɦɢɧɚɰɢɢɝɢɛɪɢɞɧɵɣɡɚɪɨɞɵɲɤɚɤɩɪɚɜɢɥɨ
ɞɟɝɟɧɟɪɢɪɭɟɬ Ⱦɥɹ ɩɨɥɭɱɟɧɢɹ ɦɟɠɜɢɞɨɜɵɯ
ɝɢɛɪɢɞɨɜɢɫɩɨɥɶɡɭɸɬɩɪɢɟɦɤɭɥɶɬɢɜɢɪɨɜɚ-
ɧɢɹɢɡɨɥɢɪɨɜɚɧɧɵɯɡɚɪɨɞɵɲɟɣɢɡɧɟɡɪɟɥɵɯ
ɫɟɦɹɧɧɚɢɫɤɭɫɫɬɜɟɧɧɨɣɩɢɬɚɬɟɥɶɧɨɣɫɪɟɞɟ
in vitro
 ɉɪɢɡɧɚɤɢ ɝɟɧɟɬɢɱɟɫɤɨɣ ɧɟɫɬɚɛɢɥɶ-
ɧɨɫɬɢ ɩɪɨɹɜɥɹɸɬɫɹ ɜɫɥɭɱɚɟ ɥɟɬɚɥɶɧɵɯ
ɚɧɨɦɚɥɢɣ ɫɬɪɨɟɧɢɹ ɢɪɚɡɜɢɬɢɹ ɡɚɪɨɞɵɲɚ
ɧɚɩɪɢɦɟɪɭɷɦɛɪɢɨɥɟɬɚɥɶɧɵɯɦɭɬɚɧɬɨɜɚɪɚ-
ɛɢɞɨɩɫɢɫɚ>@ɩɨɥɭɱɟɧɧɵɯɷɤɫɩɟɪɢɦɟɧɬɚɥɶ-
ɧɵɦ ɩɭɬɟɦ ɗɬɨ ɹɜɥɟɧɢɟ ɬɚɤɠɟ ɨɛɧɚɪɭɠɢ-
ɜɚɟɬɫɹɢɜɥɢɧɢɹɯɩɨɞɞɟɪɠɢɜɚɟɦɵɯɡɚɫɱɟɬ
ɢɫɤɭɫɫɬɜɟɧɧɨɝɨɫɚɦɨɨɩɵɥɟɧɢɹȼɩɨɫɥɟɞɧɟɦ
ɫɥɭɱɚɟ ɢɦɟɸɬ ɦɟɫɬɨ ɢɧɛɪɢɞɢɧɝ ɢ ɤɚɤ ɟɝɨ
ɫɥɟɞɫɬɜɢɟ  ɩɨɜɵɲɟɧɢɟ ɝɨɦɨɡɢɝɨɬɧɨɫɬɢ
ɩɪɨɹɜɥɟɧɢɟɪɟɰɟɫɫɢɜɧɵɯɚɥɥɟɥɟɣ  ɩɪɢ
ɨɛɵɱɧɨ ɨɬɪɢɰɚɬɟɥɶɧɨɦ ɷɮɮɟɤɬɟ ɪɟɰɟɫɫɢɜ-
ɧɵɯ ɚɥɥɟɥɟɣ ɢɧɛɪɢɞɢɧɝ ɜɥɟɱɟɬ ɡɚ ɫɨɛɨɣ
ɨɫɥɚɛɥɟɧɢɟ ɨɫɨɛɟɣ ɢɧɛɪɟɞɧɚɹɞɟɩɪɟɫɫɢɹ
ɩɨɜɵɲɟɧɢɟɮɟɧɨɬɢɩɢɱɟɫɤɨɣɢɡɦɟɧɱɢɜɨɫ
ɬɢ ɜɫɥɟɞɫɬɜɢɟ ɜɵɯɨɞɚ ɜɝɨɦɨɡɢɝɨɬɭ ɦɧɨɝɢɯ
ɚɥɥɟɥɟɣ>@
ɉɪɨɜɟɞɟɧɧɵɣ ɧɚɦɢ ɰɢɬɨɷɦɛɪɢɨɥɨɝɢ-
ɱɟɫɤɢɣ ɚɧɚɥɢɡ ɪɚɡɜɢɬɢɹ ɡɚɪɨɞɵɲɚ ɜɫɟɦɟ-
ɋɟɥɶɫɤɨɯɨɡɹɣɫɬɜɟɧɧɵɟɧɚɭɤɢ ,661ɇɚɭɤɚɄɭɛɚɧɢʋ 19
ɧɚɯ 2 SXUSXUHD VVS caucasica ɩɨɞɬɜɟɪɠɞɚ-
ɟɬ ɧɚɥɢɱɢɟ ɝɟɧɟɬɢɱɟɫɤɨɣ ɧɟɫɬɚɛɢɥɶɧɨɫɬɢ
ɩɪɢ ɮɨɪɦɢɪɨɜɚɧɢɢ ɫɟɦɹɧ ɗɬɨ ɦɨɠɟɬ ɛɵɬɶ
ɫɥɟɞɫɬɜɢɟɦ ɤɚɤ ɦɟɠɜɢɞɨɜɨɣ ɝɢɛɪɢɞɢɡɚɰɢɢ
ɬɚɤ ɢɛɥɢɡɤɨɪɨɞɫɬɜɟɧɧɵɯ ɫɤɪɟɳɢɜɚɧɢɣ ɬɨ
ɟɫɬɶɢɧɛɪɢɞɢɧɝɚȼɨɡɦɨɠɧɨɫɬɶɢɦɛɪɢɞɢɧɝɚ
ɭ2 SXUSXUHD ɩɨɞɬɜɟɪɠɞɚɟɬɫɹ ɧɚɛɥɸɞɟɧɢɹ-
ɦɢɋȺɅɢɬɜɢɧɫɤɨɣɉɨɟɟɫɜɟɞɟɧɢɹɦɪɚɫ-
ɬɟɧɢɹɞɚɧɧɨɝɨɬɚɤɫɨɧɚɪɚɫɬɭɬɟɞɢɧɢɱɧɨɢɥɢ
ɜɜɢɞɟɧɟɛɨɥɶɲɢɯɝɪɭɩɩ>@
Ɇɟɯɚɧɢɡɦɵɩɪɢɜɨɞɹɳɢɟɤɚɧɨɦɚɥɶɧɨ-
ɦɭ ɪɚɡɜɢɬɢɸ ɡɚɪɨɞɵɲɚ ɯɨɬɹ ɢɪɚɡɥɢɱɚɸɬ-
ɫɹɧɨ ɨɛɚɮɚɤɬɨɪɚɜɥɨɤɚɥɶɧɵɯɩɨɩɭɥɹɰɢɹɯ
ɦɨɝɭɬ ɞɟɣɫɬɜɨɜɚɬɶ ɨɞɧɨɜɪɟɦɟɧɧɨ ɬɚɤ ɤɚɤ
ɜɪɟɡɭɥɶɬɚɬɟɨɞɢɧɮɚɤɬɨɪɡɚɩɭɫɤɚɟɬɞɟɣɫɬɜɢɟ
ɞɪɭɝɨɝɨ ɇɚɩɪɢɦɟɪ ɩɨɬɨɦɫɬɜɨɦɟɠɜɢɞɨɜɵɯ
ɝɢɛɪɢɞɨɜ ɦɨɠɟɬ ɛɵɬɶ ɧɟɠɢɡɧɟɫɩɨɫɨɛɧɵɦ
ɢɥɢ ɢɦɟɬɶ ɫɧɢɠɟɧɧɭɸ ɠɢɡɧɟɫɩɨɫɨɛɧɨɫɬɶ
ɉɪɢ ɷɬɨɦ ɜɨɡɨɛɧɨɜɥɟɧɢɟ ɜɩɨɩɭɥɹɰɢɢ ɨɫɭ-
ɳɟɫɬɜɥɹɟɬɫɹ ɡɚ ɫɱɟɬ ɫɟɦɟɧɧɨɝɨ ɩɨɬɨɦɫɬɜɚ
ɟɞɢɧɢɱɧɵɯ ɨɫɨɛɟɣ ɬɨ ɟɫɬɶ ɢɦɟɟɬ ɦɟɫɬɨ
ɫɤɪɵɬɵɣɢɧɛɪɢɞɢɧɝ
Ɂɚɤɥɸɱɟɧɢɟ ɉɪɨɜɟɞɟɧɧɨɟ ɧɚɦɢ ɰɢ-
ɬɨɷɦɛɪɢɨɥɨɝɢɱɟɫɤɨɟ ɢɫɫɥɟɞɨɜɚɧɢɟ ɫɟɦɹɧ
OrchispurpureaVVScaucasicaɧɚɪɚɡɧɵɯɫɬɚ-
ɞɢɹɯɢɯ ɮɨɪɦɢɪɨɜɚɧɢɹɩɨɤɚɡɚɥɨɱɬɨɪɚɡɜɢ-
ɬɢɟɡɚɪɨɞɵɲɚɜɛɨɥɶɲɢɧɫɬɜɟɫɥɭɱɚɟɜɫɨɩɪɨ-
ɜɨɠɞɚɟɬɫɹɦɧɨɝɨɱɢɫɥɟɧɧɵɦɢɨɬɤɥɨɧɟɧɢɹɦɢ
ɨɬɧɨɪɦɵɉɪɢ ɮɨɪɦɢɪɨɜɚɧɢɢ ɬɟɥɚ ɡɚɪɨɞɵ-
ɲɚ ɢɫɭɫɩɟɧɡɨɪɚ ɧɚɛɥɸɞɚɥɢɫɶ ɢɡɦɟɧɟɧɢɹ
ɜɧɚɩɪɚɜɥɟɧɢɢɡɚɥɨɠɟɧɢɹɩɟɪɟɝɨɪɨɞɨɤɢɬɟɦ-
ɩɚɯɞɟɥɟɧɢɹɤɥɟɬɨɤɩɨɫɪɚɜɧɟɧɢɸɫɧɨɪɦɨɣ
Ⱥɧɨɦɚɥɶɧɵɟ ɡɚɪɨɞɵɲɢ ɯɚɪɚɤɬɟɪɢɡɨɜɚɥɢɫɶ
ɡɚɦɟɞɥɟɧɧɵɦɢ ɬɟɦɩɚɦɢ ɪɚɡɜɢɬɢɹ ɩɨ ɫɪɚɜ-
ɧɟɧɢɸ ɫɡɚɪɨɞɵɲɚɦɢ ɜɧɨɪɦɟ ɚɤɦɨɦɟɧɬɭ
ɞɢɫɫɟɦɢɧɚɰɢɢɨɧɢɞɟɝɟɧɟɪɢɪɨɜɚɥɢɉɨɜɫɟɣ
ɜɢɞɢɦɨɫɬɢ ɚɧɨɦɚɥɶɧɵɟ ɡɚɪɨɞɵɲɢ ɨɫɬɚɜɚ-
ɥɢɫɶ ɠɢɡɧɟɫɩɨɫɨɛɧɵɦɢ ɞɨ ɩɟɪɟɯɨɞɚ ɫɟɦɹɧ
ɤɫɨɡɪɟɜɚɧɢɸɬɨɟɫɬɶɩɨɤɚɨɧɢɨɛɟɫɩɟɱɢɜɚ-
ɥɢɫɶɩɢɬɚɬɟɥɶɧɵɦɢɜɟɳɟɫɬɜɚɦɢɫɨ ɫɬɨɪɨɧɵ
ɦɚɬɟɪɢɧɫɤɨɝɨɨɪɝɚɧɢɡɦɚɇɚɨɫɧɨɜɚɧɢɢɩɪɨ-
ɜɟɞɟɧɧɨɝɨ ɢɫɫɥɟɞɨɜɚɧɢɹ ɛɵɥ ɫɞɟɥɚɧ ɜɵɜɨɞ
ɨɬɨɦ ɱɬɨ ɜɵɹɜɥɟɧɧɵɟ ɚɧɨɦɚɥɢɢ ɪɚɡɜɢɬɢɹ
ɡɚɪɨɞɵɲɟɣɹɜɥɹɸɬɫɹɥɟɬɚɥɶɧɵɦɢɢɢɦɟɧɧɨ
ɨɧɢɩɪɢɜɨɞɹɬɤɬɨɦɭɱɬɨɛyɥɶɲɚɹ ɱɚɫɬɶ ɫɟ-
ɦɹɧɫɬɚɧɨɜɢɬɫɹɧɟɠɢɡɧɟɫɩɨɫɨɛɧɨɣɤɦɨɦɟɧ-
ɬɭɞɢɫɫɟɦɢɧɚɰɢɢ
Ɍɚɤɢɦɨɛɪɚɡɨɦɛɨɥɶɲɨɟɱɢɫɥɨ ɧɟɩɨɥ-
ɧɨɰɟɧɧɵɯɫɟɦɹɧɭɢɡɭɱɟɧɧɵɯɨɫɨɛɟɣOrchis
purpurea VVS caucasica ɇɨɜɨɪɨɫɫɢɣɫɤɢɣ
ɢȺɛɢɧɫɤɢɣ ɪɚɣɨɧɵ Ʉɪɚɫɧɨɞɚɪɫɤɨɝɨ ɤɪɚɹ
ɮɨɪɦɢɪɭɟɬɫɹ ɜɫɥɟɞɫɬɜɢɟ ɝɟɧɟɬɢɱɟɫɤɢɯ ɧɚ-
ɪɭɲɟɧɢɣɧɨɪɦɚɥɶɧɨɝɨɯɨɞɚɷɦɛɪɢɨɧɚɥɶɧɨɝɨ
ɪɚɡɜɢɬɢɹ ɚɧɟ ɜɪɟɡɭɥɶɬɚɬɟ ɧɟɞɨɨɩɵɥɟɧɢɹ
ɰɜɟɬɤɨɜ ɢɥɢ ɢɡɡɚ ɧɟɯɜɚɬɤɢ ɪɟɫɭɪɫɨɜ ɞɥɹ
ɪɚɡɜɢɬɢɹɦɨɥɨɞɨɝɨɢɧɞɢɜɢɞɚ
ɋɉɂɋɈɄɅɂɌȿɊȺɌɍɊɕ
Ȼɢɨɥɨɝɢɱɟɫɤɢɟ ɪɟɫɭɪɫɵɊɨɫɫɢɣɫɤɨɣɎɟ-
ɞɟɪɚɰɢɢ >ɗɥɟɤɬɪɨɧɧɵɣ ɪɟɫɭɪɫ@ Ɋɟɠɢɦ ɞɨɫɬɭɩɚ
KWWSZZZVHYLQUXELRUHVUXV
ȼɚɯɪɚɦɟɟɜɚɆȽɈɪɯɢɞɧɵɟɊɨɫɫɢɢɛɢɨ-
ɥɨɝɢɹ ɷɤɨɥɨɝɢɹ ɢ ɨɯɪɚɧɚ  ɆȽ ȼɚɯɪɚɦɟɟɜɚ
ɂȼȼɚɪɥɵɝɢɧɚɂȼɌɚɬɚɪɟɧɤɨ²ɆɌɨɜɚɪɢ-
ɳɟɫɬɜɨɧɚɭɱɧɵɯɢɡɞɚɧɢɣɄɆɄ²ɫ
 .UHW]VFKPDU + 7KH RUFKLG JHQHUD
$QDFDPSWLV 2UFKLV DQG 1HRWLQHD SK\ORJHQ\
WD[RQRP\ PRUSKRORJ\ ELRORJ\ GLVWULEXWLRQ
HFRORJ\ DQG K\EULGL]DWLRQ  +.UHW]VFKPDU
:(FFDULXV+'LHWULFK ²%UJHO(FKLQR0HGLD
²S
 Ⱥɧɞɪɨɧɨɜɚ ȿȼ ɉɥɨɞɨɧɨɲɟɧɢɟ ɝɟɬɟ-
ɪɨɫɩɟɪɦɢɹ ɢ ɤɚɱɟɫɬɜɨ ɫɟɦɹɧ ɭ 2UFKLV SXUSXUHD
VXEVS FDXFDVLFD  ȿȼȺɧɞɪɨɧɨɜɚ ȿȿ ȿɜɞɨɤɢ-
ɦɨɜɚ Ⱥȼɋɟɦɺɧɨɜ  Ȼɨɬɚɧɢɱɟɫɤɢɣ ɠɭɪɧɚɥ ²
²Ɍ²ʋ²ɋ±
ɉɟɪɟɛɨɪɚȿȺ Ɉɪɯɢɞɧɵɟ ɋɟɜɟɪɨɁɚɩɚɞ-
ɧɨɝɨɄɚɜɤɚɡɚ²Ɇɇɚɭɤɚ²ɫ
 ɇɚɡɚɪɨɜ ȼȼ Ɋɟɩɪɨɞɭɤɬɢɜɧɚɹ ɛɢɨɥɨɝɢɹ
ɨɪɯɢɞɧɵɯ Ʉɪɵɦɚ ɞɢɫɫ « ɤɚɧɞ ɛɢɨɥ ɧɚɭɤ ²
ɋɉɛ²ɫ
ɗɦɛɪɢɨɥɨɝɢɹɰɜɟɬɤɨɜɵɯɪɚɫɬɟɧɢɣɌɟɪ-
ɦɢɧɨɥɨɝɢɹ ɢ ɤɨɧɰɟɩɰɢɢ Ɍ  ɋɟɦɹ  ɩɨɞ ɪɟɞ
ɌȻȻɚɬɵɝɢɧɨɣ²ɋɉɛɆɢɪ ɢɫɟɦɶɹ²
ɫ
 0HLQNH ': (PEU\ROHWKDO PXWDQWV RI
$UDELGRSVLV WKDOLDQD D PRGHO V\VWHP IRU JHQHWLF
DQDO\VLVRISODQWHPEU\RGHYHORSPHQW':0HLQNH
,06XVVH['HYHO %LRO ²  ²9RO  ²
3±
 Ⱥɧɞɪɨɧɨɜɚ ȿȼ Ʌɟɬɚɥɶɧɵɟ ɚɧɨɦɚɥɢɢ
ɫɬɪɨɟɧɢɹ ɢ ɪɚɡɜɢɬɢɹ ɡɚɪɨɞɵɲɚ ɭ 'DFW\ORUKL]D
IXFKVLL2UFKLGDFHDHȻɨɬɚɧɢɱɟɫɤɢɣ ɠɭɪɧɚɥ ²
²Ɍʋ²ɋ±
ɈɊɂȽɂɇȺɅɖɇɕȿɋɌȺɌɖɂ,661ɇɚɭɤɚɄɭɛɚɧɢʋ
20
 &OHPHQWV 0$ (PEU\RORJ\  *HQHUD
2UFKLGDFHDUXP 9RO  *HQHUDO ,QWURGXFWLRQ
$SRVWDVLRLGHDH &\SULSHGLRLGHDH  (GV $03ULG
JHRQ 3- &ULEE 0: &KDVH )1 5DVPXVVHQ ² 
(QJODQG 2[IRUG 2[IRUG 8QLY3UHVV ²  ²
3±
 Ʌɚɝɭɬɨɜɚ Ɉɂɂɫɫɥɟɞɨɜɚɧɢɟɷɦɛɪɢ-
ɨɝɟɧɟɡɚ 2UFKLV SXUSXUHD ɢ 2UFKLV SURYLQFLDOLV
2UFKLGDFHDH  Ɉɂ Ʌɚɝɭɬɨɜɚ ȺȺ ɑɟɛɨɬɚɪɶ 
Ȼɨɬɚɧɢɱɟɫɤɢɣɠɭɪɧɚɥ²²Ɍʋ²
ɋ±
 Ʌɟɜɢɧɚ Ɋȿ Ɋɟɩɪɨɞɭɤɬɢɜɧɚɹ ɛɢɨɥɨɝɢɹ
ɫɟɦɟɧɧɵɯɪɚɫɬɟɧɢɣ²Ɇɇɚɭɤɚ²ɫ
 Ɂɚɭɝɨɥɶɧɨɜɚ ɅȻ ɐɟɧɨɩɨɩɭɥɹɰɢɢ
ɪɚɫɬɟɧɢɣ ɨɱɟɪɤɢ ɩɨɩɭɥɹɰɢɨɧɧɨɣ ɛɢɨɥɨɝɢɢ 
ɅȻɁɚɭɝɨɥɶɧɨɜɚ ȺȺɀɭɤɨɜɚ ȺɋɄɨɦɚɪɨɜɚ
Ɉȼɋɦɢɪɧɨɜɚ²Ɇɇɚɭɤɚ²F
ɂɧɝɟȼɟɱɬɨɦɨɜɋȽȽɟɧɟɬɢɤɚɫ ɨɫɧɨɜɚ-
ɦɢɫɟɥɟɤɰɢɢ²Ɇȼɵɫɲɚɹɲɤɨɥɚ²ɫ
ɅɢɬɜɢɧɫɤɚɹɋȺɋɟɦɟɣɫɬɜɨ2UFKLGDFHDH
ɧɚ ɋɟɜɟɪɨɁɚɩɚɞɧɨɦ Ʉɚɜɤɚɡɟ ɝɟɨɝɪɚɮɢɹ ɷɤɨɥɨ-
ɝɢɹɨɯɪɚɧɚɈɯɪɚɧɚɢɤɭɥɶɬɢɜɢɪɨɜɚɧɢɟɨɪɯɢɞɟɣ
Ɇɚɬɟɪɢɚɥɵ,;Ɇɟɠɞɭɧɚɪɨɞɧɨɣɤɨɧɮ±ɫɟɧ-
ɬɹɛɪɹɝ²ɋɉɛ²ɋ±
ɉɨɥɭɱɟɧɨ
(9$QGURQRYD(((YGRNLPRYD$96HPHQRY
5($62162)7+(819,$%/(',$6325(6)250$7,21
,125&+,6385385($68%63&$8&$6,&$25&+,'$&($(
,QVWLWXWLRQRIWKH5XVVLDQ$FDGHP\RI6FLHQFH.RPDURY%RWDQLFDO
,QVWLWXWHRI5$66DQNW3HWHUVEXUJHPDLOHOHQDDQGURQRYD#PDLOUX
)HGHUDO)RUHVWU\$JHQF\RIWKH)HGHUDO%XGJHW,QVWLWXWLRQ
©5XVVLDQ&HQWUHRI)RUHVW+HDOWKªEUDQFK©&HQWUHRI)RUHVW+HDOWK
RI.UDVQRGDU5HJLRQª.UDVQRGDU
$F\WRHPEU\RORJLFDO VWXG\ RI WKH VHHG IRUPDWLRQ LQ 2UFKLV SXUSXUHD +XGV VVS SXUSXUHD IURP WZR
SRSXODWLRQVRI.UDVQRGDUUHJLRQZKLFKDUHFKDUDFWHUL]HGE\ORZUHDOVHHGSURGXFWLYLW\LVFDUULHGRXW
,WLVHVWDEOLVKHGWKDWWKH VHHGV EHFRPH QRQYLDEOH XS WR WKH PRPHQW RI GLVVHPLQDWLRQ EHFDXVH RIWKH
OHWKDODQRPDOLHV LQHPEU\RDQGVXVSHQVRU GHYHORSPHQWWKDWDUHFRQQHFWHG ZLWKWKHPDQLIHVWDWLRQRI
JHQHWLFLQVWDELOLW\LQWKHGLYLVLRQDQGGL൵HUHQWLDWLRQRIFHOOVGXULQJHPEU\RJHQHVLV
.H\ZRUGVSODQWELRUHVRXUFHVQHZDQGQRQWUDGLWLRQDOSODQWFXOWXUHV2UFKLVSXUSXUHDHPEU\RJHQHVLV
HPEU\RDQGVXVSHQVRUGHYHORSPHQWDQRPDOLHVVHHGXQYLDELOLW\JHQHWLFLQVWDELOLW\
ResearchGate has not been able to resolve any citations for this publication.
ResearchGate has not been able to resolve any references for this publication.