Interested Article - Обходные пути биосинтеза андрогенов у человека

???????? ???? ?????????? ?????????? ( ????. backdoor pathway of androgen biosynthesis ) — ???????? ?????????????? ?????, ? ??????? ????????????? ??????? ??????? ??????? ( ????????? ) ?? ????????? , ?????????? 21 ???? ???????? (C ₂₁ ), ????? ?? 5?-?????????????? ??? ??????? ???????????? ? ?????????????? ? ???????? ????????????? ?????????????? ????????? . ?????? ?? ???????? ????? ??? ?????? ? 2003 ???? ? ???????? ??? ??????? ???????, ???????????? ? 17?-???????????????????? ? ????????????? 5?-???????????????????? . ? 2004 ???? ??? ??????? ??????? ???? ???????????? ? ? ?????. ????? ?????, ? ????? ???? ?????????? ????????? ?????? ?????, ??????? ????? ? ????? ?????? ??????? ???????????? ? ?????????????? ???????? ? ??????? 11-?????????????? – ????????? ?????????? ?????????? , ?????? ?? ???? ??????????? ???????????? ??? 5?-??????????????????? . ??? ?????????????? ???? ???????????? ?? ?????? ? ????????, ?? ? ? ?????? ????????????? , ? ???? ??????? ? ?????, ???? ? ??????? .

???????? ???? ???????? ????????????? ???????? ????, ??????????? ??????????? ? ????????????? ? ???????? ???????? ??? ????????????? ????????? . ???????? ???? ????? ????? ??? ???????? ????????, ??? ? ??????? ????. ????????? ? ???????? ???? ????? ????????? ? ??????????????? ???? ( ?????????? ???????? ??????? ???????) ? ????????????? ??????? ? ?????????????? ???????? ??????? ??????? ? ????????????? ????????? . ? ???????? ????? ???????? ???? ????? ????????? ? ??????? ?????????? ? ? ????? ???????? ??? ?????????? ???????? ??? ??? ?????????????? ?????? .

???????? ???? ?????????? ?????????? ???????????? ????? ????????? , ????? ??? SRD5A1 / SRD5A2 , / ? CYP17A1 . ??????? ? ?????, ?????????? ??? ????????, ????? ???????? ? ????????? ???????????????? ????????? ????, ??? ?????? ????????? ??????????? ??????????? . ???????? ???? ?????????? ?????????? ???????? ???????? ???????? ????????????, ? ??? ????????? ????? ????? ???????????? ???????? ??? ?????????? ??????? ??????? ? ???????? ?????????, ????????? ? ????????????? ??? ?????????? ?????????? ?????????? .

???????

????????? ?????? ???????? ?? ??????? 21 (??????? 18 ? 19 ???????) ? ?????????????? ?????????? ????, ??? ??? ?????????? ????????????? ?????????

???????? ??????????? ?????????? ?? ???????? ??????? ?? ?????????? ??????? ???? ???????? ? ????????????? ??????????? , ?????????????? ? ???????? ??????. ????? ??????????, ????????? ???????????? ? ???? ?????? ? ???????????? ??????? ???????????? ????? ????? ?????????????? ? ????????-??????????????? ?????????? . ???? ???????? ?????? ???????? ???? ? ???????????? ??????? ?????????????? ???????? ? ?????????? ??????? ????????, ? ????? ?????????? ?????? ??????? ?????????, ?? ??????????? ????????? ? ?????, ???????? ????????????? ?????? ??? ???? ????? .

????????? ????????? ????????? ????????? ??????? ?? ???????????? ?????????, ??? ??? ??? ????? ?????? ??????? ?????????? ? ????????? ??????????. ??? ?????????? ????????? ??????????? ?????????? ?? ?????????, ??????? ????????? ??????? ??????? ?? ???????? ? ???????????????? ??????? ??????? ?????????, ? ??????? ?? ?????????? ?????????? ??????????, ??????? ????? ?????? ????????????? ?????????? ??? ?????? ???????????????? ? ?????? ???????? . ?????????? ?????????? ????????? ?? ????????? ????????????? ??????? ?? ???????? ? ???????????????? ??????? ??? ??????? ??????? ?????????, ?? ??? ????? ???? ?????????? ??????????? ????? ???????? ??????????, ????? ??? ???????????, ??? ?? ???????????? (????????????????? ? ?????????????? ????).

???????? ???? ?????????? ?????????? — ?????? ?????????????? ???????? , ??????? ????????? ? ???????????? ?????????? ???????? ?????????? ?? C ₂₁ ????????? ( ????????? , ????? ??? ??????????? ? 17?-??????????????????? (17OHP)) ??????????? ?? 5?-??????????????, ?? ???? ??????????? ????????????????? ?????????? ??????? , ?????????? ? ????????? ?????????????? ?????? , ??????????? ? ??????? 5? ??????????? ( ??????????? ) ???? ????????. ??? ???? ???? ? ????? ?????? ??????? ???????????? ? ?????????????? — ??????, ??????? ???????????? ? ??????? ???? ?????????? ?????????? . C ₂₁ -???????? ???????????? ??? ??????, ??? ?? ???????? ????? 21 ???? ???????? , ?? ???? ?????????? ??????? ?????????? ? «C ₂₁ », ???????? C ₂₁ H ₃₀ O ₂ — ??????? ???????????? .

? 2004 ???? 5?-?????????????? 17OHP ???? ??????? ? ??????????? ?????????? ??? ??????? ?????????????? ????, ??????? ? ????? ???????? ? ????????? 5?-??????????????????? (DHT) ? ???????? ??????. ?? ??? ?????? ??????????? XXI ???? ???? ?????????? ??? ????????? ?????, ??????? ????? ? ????? ?????? ??????? ???????????? ? ?????????????? ????????? ? ????????? 11-?????????????? , ??????? ????? ???????? ???????????? ???????????, ? ??? ?? ???????????? ?? ???????????? ?????????, ??? ? ??????????? ??? DHT.

??? ?????????????? ???? ???????????? ?? ?????? ? ????????, ?? ? ? ?????? ????????????? , ? ???? ??????? ? ?????, ???? ? ??????? . ??? ??????? ??????? ( ???. Notamacropus eugenii , ????. Tammar wallaby ) ???????? ????????? ???????? ????????? ??????? ?????????????? ? ???????? ???????? ? ????????? ?????????? ?????????? ????????? ????, ??? ??????? ?????????????? ? ????? ???? ?????????? ?????? ????? ????????: ????? ???????? ????????????? ????? ??? ???, ? ??????? ?????? ?? ??????? ???? ????? ????????. ??? ????????? ??????? ?? ?????? ??????? ???????? ?? ???????? ? ?????? ???????? ??????? ??????????????, ?? ? ???? ?????????? ???? ????????, ????? ??????? ??? ???????, ? ?? ????????? ? ???????? ??? ? ?????? ?????????????, ??? ?????????? ?????????? ? ??????????? .

???????? ????????? ???? ?????????? DHT ? ??????? ??????? ? 2003 ???? ????????? ????????? ????????? ????????? ?????????? ? ????? ? ?????? ????? ? ?????????? ??????? ??????? ? ???????? ?????????, ????????? ? ????????????? ??? ?????????? ?????????? ??????????. ????????? ????? ????????? ???????? ??????????? ???????? ??????? ??????? ? ????????????? ??????? ???, ????????, ????????????? ???????? ??????? ??????? ? ?????????. ????? ?????????? ????? ?????????? ??? ???? , ???????? , ???????? ? ???????????? ????????????, ????????? ?? ???????? ???? ? (?????????? ???? ????? ?? ???????? ???? ?? ???? ??????), ?????????? ???????? ? ??? ?????????????? ?????? . ??? ????????????? ????????? ??????? ? ??????????? ?????????? ? ??????????? ??????????? ?????????? ? ????????? .

????????

???????? ???? ?????????? ???????????????????

???????? ???? ?????????? 5?-??????????????????? ?? ???????????? ? 17-???????????????????? ? ????? ?????? ??????? ???????????? (??????? ???????), ??????? ???? ?????????? ?????????? (?????? ???????). ????? ???????? ???? ?????????? ???????????????? ????? ????????????????? ??????? ??????? ????????? CYP11B1/2 (????? ???????). ??? ??????? ??????? ??????????, 5?-?????????????????? ?????????????? ????? ?? ???????????? .

?????? ? ???????? ?????? ????????? ???? ?????????? ?????????? ???????? 5?-??????????????? ???????????? ??? 17?-???????????????????? (17OHP), ? ????? ??????? ??????? ??????????????, ?????????? ? 5?-??????????????????? (DHT) ?????????????? ?????, ????? ???????? ????????????? ?????????? ????????????? ? ??????????? .

???? ???? ???????????? ?? ????? ??????????? ??????????????? ???????? ? ???????? ? ?????? ??????? ??????? ?????????????? . ??????? ?? ??? ?????? ? ???????? , ? ????? ??????????? ? ????? . ??? ???????, ??? ? ???????? ???? ?????????? DHT ?????????? ??? ??????????? ???????? ??????? ??????? ??????? ? ????????, ??????? ??????? ????????? ???? ?? 17OHP ??? ???????????? ? DHT ???????? ? ?????????????? ???????? ??????? ??????? ? ????????? ???????? ???? , ????????? ???????????? ??????????? ???????? ???????????????? ??? ?????????? DHT ????? ???????? ???? .

??? ???????? 21-???????????? ??? ???????? ??????????????? ????????? ?450 , ???? ???? ????? ???? ??????????? ?????????? ?? ???????? ? ???? ???? ??? ?????????????? ????????? ?????? ?????????????? 17OHP .

? ?? ????? ??? ??????? 5?-?????????????? (????????????????? ?????????? ??????? ? ??????? 5? ???? ???????? ????????) ???????? ????????? ?????????????? ? ??????? ???? ?????????? ??????????, ??? ??????? ? ???????? ????? ? DHT ???????? ??????. ??? ???????? ???? ?????????? ? 5?-?????????????? ???? 17OHP, ???? ???????????? , ??????? ? ???????? ????? ???????????? ? DHT .

????, ???????????? ?? 17?-????????????????????

?????? ?????? ????? ???? ???????? 5?-?????????????? 17OHP ?? (17OHDHP), ??????? ????? ???????? ??? 17?-??????????????????????????. ??? ??????? ???????????? ??????? SRD5A1 .

17OHDHP ????? ????????????? ? (5?-Pdiol) ??????????? 3?-?????????????? ???????????? 3?-??????????????? ????????????? (AKR1C2 ? AKR1C4) ??? ????????? HSD17B6, ??????? ????? ???????? 3?-????????????????? ??????????? . 5?-Pdiol ????? ???????? ??? 17?-??????????????????????? ??? 17OH-??????????????? .

5?-Pdiol ????? ????????????? ? 5?-?????????-3?-??-17-??, ????? ????????? ??? ??????????? (AST), ?? ???? 17,20-??????? ?????????? ???????? CYP17A1, ??????? ????????? ??????? ???? (????? C17-C20) ?? ??????????? (???????????) ???? ????????, ????????? ??????? ? ₂₁ ( ??????? ) ? ??????? ? ₁₉ ( ????????? ??? ???????? ) .

?? ????????? ?????, AST 17?-????????????????? ?? 5?-?????????-3?,17?-????? (3?-diol) ? ??????? ???????? HSD17B3 ??? AKR1C3 .

????????? ?????? ???????? 3?-????????? 3?-diol ? ???????? ?????? ?? DHT, ?????????, ?????????? ??????????? 3?-???????????????????????, ????? ??? AKR1C2 , HSD17B6, HSD17B10, RDH16, RDH5 ? DHRS9 . ??? ????????? ?? ????????? ? ??????? ???? ?????????? DHT .

??? ??????? ????????? ?? ???? ???? ????? ?????????? ????????? ???????: 17OHP ? 17OHDHP ? 5?-Pdiol ? AST ? 3?-diol ? DHT .

????, ???????????? ?? ????????????

???? ?? ???????????? ? DHT ??????? ?????????? ???? ???? ?? 17OHP ? DHT, ?? ???????? ?????????? ??? 5?-????????? ????? ???????? ???????????, ? ?? 17OHP. ???????????? ??????????? ? ????? ???????? ???? ???????? ???????? ?????????? ??? ????????? ????, ??????? ????????? ?? ?????? ?????????? ?????? .

?????? ????? ?? ???? ???? ???????? 5?-?????????????? ???????????? ?? 5?-??????????????????? (5?-DHP) ? ??????? ???????? SRD5A1 .

????? 5?-DHP ???????????? ? ??????????????? (AlloP5) ??????????? 3?-?????????????? ? ??????? AKR1C2 ??? AKR1C4 .

????? AlloP5 ???????????? ? 5?-Pdiol ??? ????????? 17?-?????????????? ?????????? CYP17A1 .

????? ???? ???? ????????? ??? ??, ??? ? ????, ??????? ?????????? ? 17OHP, ? ??? ????? ?????????? ????????? ???????: ??????????? ? 5?-DHP ? AlloP5 ? 5?-Pdiol ? AST ? 3?-diol ? DHT .

11-?????????????

????????????

??????? «11-????» ????????????? ? ???????? ???????? ??????????? ????? ????????? ? ???????? ???? (=O) ?????????????? ??????, ?????????????? ? ????? ???????? ? ??????? 11 ???? ???????? .

? ????????? ??????????, ??????????? ?????? ? ?????? ?????????, ????? ????????? ??????? «11-????», ????????, «11-???????????» . ? ????? ??????? «????» ?????? ???????????? ??? ??????????? ???????????? ??????????. ?????? ???????????? ???????????? ???????? ?? ????????????? ???????????? 1989 ???? ?? ??????????? ???????????? ??????? «????» ??? ???????????? ?????????, ???????? ??? ????????????? ? ??????????? ?????????? ??????????. ????????????? ???????????? ????????????? ???????? «????», ????????, 11-???????????? ?????? 11-????????????. ?????? — ??? ????????, ? ????????? ??????? ???????????? ?????? ??????? ? ????? ??????????????? ?????????? (R ₁ -CO-R ₂ ). ????? ???????, ? ??????? ??????? ??????? ? ???????????? ??????, ? ? ????????? ????? ???????? ???????? ?????? ?????????? ???? ? ?????????? ????????. ????????? ???????????? ?????? ??? ???????? ???????, ??????? ???????? ????? ?????? ??????????? ????, ?? ???? ? ??? ?? ???? ????????, ???????? ????????? ?????????????, ?? ?????? ??????????? ?????? .

???????? ???? ?????????? 11-??????????????

??????????? ???? ? 11-?????????????? ? 11-?????????????????? ? ??????????????? (?????? ???????). ??? ?????? ????????? ?????????? ????????????? ? ??????? 17 ???? ?????????, ??? ??????????? ???????? ?????????? ???????????-?????????????????. ?????? ?????? — ?????????? ????????????, ?????? — 17-????????????????????. CYP17A1 ???????????? ???????? C ₂₁ (????????) ?? ????????? C ₁₉ (??????????). ????????? ??????????????, ????????????? ??????? ??????????????, ?? ??? ?? ????????, ???????? ??????????? ?????????. ????????? ???????, ?????????????? CYP17A1, ??????? ?????????????? ?????? 11-?????????????? ? 11-?????????????????? (????? ?????), ??????? ??? ???????. ???????? ? ⁵ , ??????? ???????????????? ? ???????? ? ⁴ , ????? ??????? ??? ??????? .

???? ??? ????????? 11-????????????? — 11-??????????????? (11KT) ? 11-?????????????????????? (11KDHT) — ??????? ????????? ? ?????????? ????????? ?????????? ?? ???? ??????????? ????? ??, ??? ? ???????????? ? DHT ?????????????? . ? ????????? ??????? ??????????????, ???, ???? 11-?????????????????, ????? ??? 11-??????????????????? (11OHT) ? 11-?????????????????????????? (11OHDT) ????? ?? ???????? ???????????? ??????????? ???????????, ??? ??? ?? ????? ???? ??????? ????????????????? ??????????? ??????? 11KT ? 11KDHT . ????????????? ???????? ?????????? ? ?? ??????? ?? ???????? ??????? ?? ?? ??????????, ?? ?? ???????????? ? ????? ? ? ??????, ? ?? ?? ????????????. ???? ???????????, ??? ???????? 11-????????????????????? (11OHA4) ? 11-????????????????? (11KA4) ???????? ??????????? ??????????? ??????????? , ?? ???????? ??????? ?????????? ? ???? ?????????, ????????? ??? ????????? ??? ??????????????? ????????????-???????? ?????????? .

???????? ???? ? 11-?????????????? ? 11-?????????????????? ????? ???? ? ????? ???????????? ????? ??????? ????? ? ⁴ ????????? (17OHP ? ???????????), ??????? ????? ???????????? ????? ?????????????????? ?? ???? ??????????????:

11?-????????????????? ? ??????? CYP11B1 ? ???? ?????????????,
5?-?????????????? ? ??????? SRD5A1 / SRD5A2 ,
????????? 3?-??????????????/????????? ?????????????? ?????? ?????? / ?????? ? ???????? ???????? ? ??????? ??? .

???? ? ???????? ????????

?????????? ??????????? ???? ?????????????

??? ?????????? ??????????? ???? ????????????? (????) ??-?? ???????? 21-???????????? ??? ???????? ???????? P450 ??????????????? (POR) , ????????? ? ???? ?????????? ??????? 17OHP ???????? ? ???????? ?????????? DHT ????? ???????? ????, ??????? ?????????? ? 5?-?????????????? 17OHP. ???? ???? ????? ???? ??????????? ?????????? ?? ???????? ? ???? , ? ? ???????? ???????? ???? ? ????? ????? ????????? ? ??????????? ???????? ??????? ??????? , ?? ???? ? ?????????????? ???? ????????? ? ????????????? ???????, ??? ???????? ? ??????????? ??????????? ???? ??? ???????? ????? ??????? . ????? ?????????? ??????? 17OHP, ????? ???? ???????? ? ?????? ????????????, ??? ???????? ? ??? ?? ????????????. ????? ????, ? 17OHP, ? ???????????, ????? ????? ??????? ??????????? ??? 11-?????????????? ? 11-?????????????????? ??? ????? , ????????? ????????? ????????.

?????????????? ??????? ???????? ??????? ??????? ? ????? ??-?? ?????? ??????? ?????????? ???????? ? ??????????? ???????? , ??? ???????, ????? ???????, ??? ??? ???? ?, ? ??????? ?? ????, ?? ???????? ??????????????? ??????????? .

11-????????????? ? 11-?????????????????

??? ? 1990-? ????? ????????, ??? ?????? ? ????? C ₂₁ 11-????????????????? (11-??????????????????? (11OHP4) ? 21-?????????????? (21dF)) ???????? ??? ??? ????????????, ??? ? ??? ?????????????? ???? , ? ??????? ??-??/?? , ?????????? ??? ????????, ???? ?????????? ??? ???????????? ?????????? . ? ????????? ? ???????????? ????, ?????????? ??????? ????????????????? , ?????? C ₁₉ 11-????????????? ? 11-????????????????? ? ????????? ????? ???? ???????? ? 3-4 ???? ?? ????????? ? ??????????? ???????, ?????????? ?? ???????? ????? . ? ??? ?? ???????????? ?????? C ₁₉ 11-?????????????? ???????????? ????????????? ? ???????? ??????????? (??????, ??? ???????????) ? ??????, ?? ???????????? ? ??????. ?????? 11KT ???? ? 4 ???? ???? ?? ????????? ? ????????????? ? ?????? ? ???? ????????????. ? ???????? ?????? ? ???? ??????????? ?????????? DHT, ??????????????????? ???????? ?????, ? ??????????, ??????????????????? ??????? ?????, ????????????? ?? ???? ????, ??? ?????????? ???????? ??????? ?????????? ? ??????? ??????? ????????????????? . ? ?????? ? ????????? ????????????? ???????? ??????? 11KT ????? ??????? . ? ????????? ? ???? ? ?????? ????????? ???????????, 11-????????????? ???????? ??????????? ??????, ??? 17OHP, ??? ?????? ?????? ??????????? ????????????? ???????? ??????????? . ? ?????? ? ???? ??????? 11-?????????????? ????? ????????? ?? ??????? ???????? ????? .

??? ???????, ??? ? ???????? ???? ?????????? DHT, ?????????? ??? ??????????? ???????? ??????? ??????? ??????? ???????? ?? ????? ??????????????? ???????? . ??????? ?????????????? ?????????? ????????? ? ???????? ???? ? DHT ?? 17OHP ??? ?? ???????????? ???????? ? ?????????????? ???????? ??????? ??????? ??????? ? ????? ???????? ???? , ????????? ???????????? ??????????? ???????? ???????????????? DHT ? ???????? ???? .

???? ????????? ?????????

AKR1C2/AKR1C4

??? ???????? ???? ????????? AKR1C2 ? AKR1C4, ?? ?? ???? ? ???????? ???? ?????????? ?????????? ??????? ?????????? ?? ??????????? ???????????? ???? ????????? ? ?????????? 46,XY (???????) ?? ???? ?????. ??? ???????????? ????????, ??? ????????? ??????? ??? ????????? ? ???? ?????? ??? ?????? ? ????????? ? ?????, ?????????? ???????? AKR1C2 ?/??? AKR1C4. ??? ???????? ????????? ????????????? ? ???????? ???? ? DHT, ?? ?? ? ??????? ???? ?????????? DHT ????? ???????????. ??????? ? AKR1C3 ? ?????, ??????????? ? ??????? ???? ?????????? DHT, ???? ????????? ??? ??????? ?????????? ???????? ????. ??? ???? ????????? ? ?????????? 46,XX (???????), ???????????? ??????? ?????????-??????, ??????? ?? ?? ???????, ???? ????????????? ??????????? ? ???????????. ???? ? AKR1C2, ? AKR1C4 ?????????? ??? ??????? DHT ? ???????? ????, ???????????? ????????, ??? ??????? ?????? ? AKR1C2 ?????????? ??? ????????? ???????????????? ????? ???? . ??????? ? ????? AKR1C2/AKR1C4, ??????? ? ?????????? ???????? ???????? (???), ?????, ? ?? ??? ??? ???? ???????? ?????? ? ???? ???? ??????. ??? ?? ?????, ? ?????? ???????????? ??????? ?????????? ???? AKR1C2/AKR1C4 ??? ?????????? ?????? ????????? ? ?????????????? ????? ?????????? ?????????? .

CYP17A1

????????????? ??????? ???????? 17,20-????? (ILD), ????????????? ????????? ? CYP17A1, ????????? b ₅ ? POR, ????? ????? ???????? ???????? ???? ?????????? DHT, ????????? ?????????? 17,20-????? CYP17A1 ?????????? ??? ??? ????????, ??? ? ??? ????????? ???? ?????????? DHT . ???????? (46,XY) ? ILD ?????? ???????????? ??? ???????? ??-?? ?????????? ????????????? ??????????????, ? ?? ????? ??? ??????? (46,XX) ?????? ???????? ??? ??????????? ??????? ???????? ?????? ?? ???????? ?????????? , ????? ? ??? ??????????? ????????? ???????? , ????????? ? ???????????????? ??????? ????? .

11-????????????? ? ????????? ???????? ????????

11-????????????? ????? ?????? ?????? ???? ? ??? . ????????? 11-?????????????? ? ????? ????? ????????? ? ????????? ??????? ????????? ????????? — ?? 8-9-?, 13-24-? ??????? ????????????, ? ????? ? 31-? ?????? ? ?????. ?? ???? ???????, ????????? ?????????? CYP17A1 ? CYP21A2 ???????? ? ?????????? ???? ??-?? ???????? ????????? ? ?????????? ?????????? ??? CYP11B1 ? ????? ??????????? 11-??????????????, ? ????? ???????? ?????????? ???????? ????? ???????? ???? .

??? ?????????????? ??????

???????? ???? ?????????? ?????????? ????? ?????? ???? ??? ???? ?????????????? ?????? . ??????????? ?????????? — ??? ??????????????? ?????? ? ???? ?????????????? ??????, ??????? ????? ???? ?????????? ????? ????????? ??? ?????????? ????????????, ??????? ??????????? ????? , ?? ????? ??????????????? ??????? ????? ????????? ? ????????????-???????????? ??? ?????????????? ?????? (?????). ???? ????????? ???????? ? ???????? ?????? ???????????? ? ????????? ????? ?? 90-95 %, ??????? ??????????????????? ? ?????????????? ?????? ????????? ?????? ?? 50 %, ??? ???????????? ????????????? ? ???, ??? ???????? ???????????? ??????????? ???????? ??? ????????? DHT ??? ??????? ???????????? ?? ????? .

???????

? 1987 ???? ?.??????? ? ???????? ???????????? ????????????? ????????? ????? ? ??????????? ???????????? ???????? ????????? ? ??????????????????, ??? 5?-?????????-3?,17?-???? ??????????????? ????????????? ?? 17?-???????????????????? (17OHP) .

? 2000 ???? ?. ??? ? ???????? ??????????????????, ??? ????????????? 3?-diol ?????????? ???????? ?????????????? ?????? ? ??????? ??????? ????? ??????????? ? DHT ? ???????? ??????. ???????? ??????? ?? ????????????? ???????? ?????????? ? ????????????? ??????? ? ? DHT ?? ????? ???????? ????????, ??? ????????? ????????????, ??? ?? ??? ???????????? ?????? ???????? ??????????????. ???? ?????????? ?????????? 3?-????? ? ???????????? ??????????? ?? ???? ???????? ????, ??? ? DHT, ???? ????????, ??? 3?-diol ????? ?????????? ??????? ?? DHT ??? ????????? ???? ???????????? .

? 2003 ???? ??. ?????? ? ???????? ???????????? ????????? ??????? ???????? ( ??????? ) ? ???????????? ??????? ?????????????, ????? ????????, ??? ?????????? 5?-????????? ? ???? ????? ???????????? ????? ???? ?? 17OHP ?? 3?-????? ??? ???????????? ? ???????? ?????????????? ????????: 17OHP ? 17OHDHP ? 5? -Pdiol ? AST ? 3?-diol .

? 2004 ???? ?. ???????? ? ???????? ??????????????????, ??? ? ?????????? ????? ????????? ????? ????, ? ???????? ??, ??? ???? ?????????????????? ? ??????? : ??????????? ? 5?-DHP ? AlloP5 ? 5?-Pdiol ? AST ? 3?-diol .

????? ? ??? ?? ???? ?. ???? ???? ?????? ???????? ???? (backdoor pathway ) ? ????????? ??? ??? ???? ? DHT, ???????: 1) ??????? ??????? ????????????? ?????????? ????????????? (?4) ? ?; 2) ???????? 5?-?????????????? ? ₂₁ -????????? (?????????) ? ??????????? ?? ?????????? ? ? ₁₉ -???????? (??????????); ? 3) ???????? 3?-????????? 3?-????? ?? DHT. ???????? ???? ????????, ??? ????????? ????????????? ??? ???????????? ?????????? ? ?????????????? ?????????? ? ?????, ????? ??????? ???? ?????????? DHT ?? ????? ????????? ????????? ??????????? ???????????. ????, ??????? ????????? ?. ????, ????????? DHT ? ????? ????, ??????? ????????? ??. ??????, ? ??????: 17OHP ? 17OHDHP ? 5?-Pdiol ? AST ? 3?-diol ? DHT .

??????????? ?????????? ???? ??????????? ???? ??????? ?????????????????? ? 2012 ????, ????? ?. ?????? ? ???????? ????????, ??? ??????????, ????????? ? ????? , ????????? ? ????? ? ??????????? ????????? ???? ?????????? ?????????? ?? 17OHP ? DHT ? ????????? ? ????????? 21-???????????? .

?. ??????? ? ???????? ? 2017 ???? ?????????????????? ?????????????? ???? ?? ????????? C ₂₁ ?? 11KDHT, ??????? ??????? A4 ? ???????????. ???????? ???? 11KDHT ?????????? ????????? ???? ? DHT. ??? ???? ? 11-?????????????? ????? ???? ??????? ??? ???????? ???? ??-?? ?????????? ????????, ? ???? ????????????? ???????????????? ? ??????????? ????????????? .

??. ?????

??????????

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