????? ????????? ?? ?????? ????? ?? ?? ???? ????? ???? ????????? ?????, ??? ?? ???? ?? ????? ????? ?????? ????? ???? ???????? ?????? ????????? ??????? ?????? ????????. ??????? ?? ????? ?????, ??? ?????? ????? ??????? ?????, ??? ?????? ????? ??? ???????? ?? ?????? ??????. ?? ???????? ??????? ?-20 ?????? ????? ????? ???????? ???? ???? ??????? ?????? ????????? ???????. ????? 20 ?????? ????? ???? ???? ??? ???, ??? ?????? ?????? ?? ?????? ?? ??? ?????? ?????? ?. ??????? ???? 1820, ???? ?????? ???????? ?? ??? ??. ??? ???? 1935. ????? ?????? ?????? ???? ??? ?????? ???? ??????? ?? ?? ???? ?? ????? ???????? ?? ?????? ??????, ??? ?? ????? ?? ??????? ????? ???????? ????????? ????????? ????? ????. ?????????? ?? ?????? ????? ??? ????? ?????? ?? ????? ??????? ?? ???? ??????????. ???? ?? ????? ??? ???? ?????? ????? ??? ???????? ????????? ?????? ???? ????? ?? ???????. ?????????? ?? ?? ?????? ?????? ??????? ?? ??? ??????? ??????? ???????? ?????? ?????? ?? ??????? ???????? ??????? ???????. ????? ???? ????? ???????, ???? ???? ?? ?????????? ?? ?????? ????? ?-5 ??????: ?????? ??????, ???? ?????? (Glu), ??????? (Gln), ?????? (Pro) ???????? (Arg). ??????? ?? ?????? ????? ??? ?????? ???????, ??????? ???? ?????? ?????? ?????. ????? ??????? ????? ?????? (Asp), ???????? (Asn), ????? (Lys), ??????? (Thr), ??????? (Met) ??????????? (Ile). ?????? ?????? ?????? ?? ????? ?? ?????? ?????? ???????, ???? ?? ???? ?? ??????? ???????? ???????. ????? ?? ?????? ????? ????????, ???? ????????? (Phe), ??????? (Tyr) ????????? (Trp). ??????? ?? ?????? ????? ??? ?????? ?? ?????????-4-????? (E4P) ???????????????? (PEP), ??? ???????? ??? ?? ????? ?????? ?????? ???????? ????????. ????? ??????? ????? ???? (Ser), ?????? (Gly) ???????? (Cys). ?????? ?????? ?????? ?? ????? ?? ?????? ?? ????, ???? ????? ???????? ?? ??????? ??????????? ????. ????? ?????? ????? ????? (Ala), ????? (Val) ??????? (Leu). ????? ??????? ????? ??? ?????? ??????? ?????, ?? ??? ?????? ????? ????? ???????, ??????? ??? ???? ??? ??????? ?? ??? ???? ????? ???????.

??????????, ????? ???????, ????? ??? ?????? ??????? ?????, ??????? ?????? ????? ???????? ?? ??? ???? ?????. ???? ???? ???????? ??? ?????? ??????? ?? ????? ?????? ??????. ????????? ?? ????? ?????? ?????? ????????? ???? ?? ??? ?????????-4-P ?-PEP. ?????????? ?? ???????? ??? ??????, ?????? ????? ??? ????? ??????????? ????????? (PRPP). ??? C ?????? ?? PRPP ?????? ?????? ???? ?????????? ???? 5 ??????, ????? ?????? ?????? ????? ??? 3C. ?????????? ?? ?????? ????? ????? ????? ???? ?????? ????????. ?? ?? ?? ????? ????? ?????? ??????????? ??????? ???????, ??? ?? ???? ??? ???? ???? ?????? ??????? ????. ????? ?????? ????? ?? ??? ????? ??????????? ???? ????? ????? ???? ???????? ?????? ??? ????? ????? ??????, ???? ????? ??????? ????, ????, ?????? ??????? ?????.

?????, ?????????? ?? ?????? ????? ????? ?? ????? ????????? ???????? ??????? ????????, ?? ????? ????? ?????? ????? ???????? ?????????? ?? ??? ???????????????. ?? ?? ?? ???? ?? ?????? ??????, ??? ?? ???? ???????? ?????? ?????? ???????????? ????? ?????? ??? ???? ?? ???? ???????? ?? ?????? ?????????.