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(2) AO 210 Ml'1H(3) pacnaA rn111KoreHa 111 rntoKOHeoreHes npaKrnYeCKl'1 B paBHOH creneHlll 06ecneY11BatoT KpOBb rntoK030H. 1 - COCT051Hille nOK051 (nocra6cop6Ti11BHblH nepillOA); 2, 3 -- cpi113111Y0C· Ka51 HarpysKa.

l1TaK, H3JlO)((eHHhle CBC,Ll,CHlUI IT03BOJ151IOT C,LJ,e- Jl3Tb BblBOll. 0 TOM, 'ITOKOOpD,HH3UH51 CKOpOCTeti: rJIHKOJltt3a, rnJOKotteoreHe3a, c11ttTe3a 11 pacrra-

D.a fJIHKOreHa c yqaCTMCM ropMOHOB o6ecnelJHBaeT:

• rrpenoTspameHtte qpe3Mepttoro noBhIIlleHHSI KOHUCHTpaUHH fJTIOK03hl B KPOBM ITOCJle nptteMa T1l1IUH;

357

3arracatt11e nmKoretta 11 ero 11crroJib30Batt11e B rrpoMe)K)'TKaxMe)K)zy rrp11eMaM11 rr11m11;

ctta6)!(ett11e rJIIOK030H MbillIU, I10Tpe6HOCTb KOTOpbIX B 3Hepr1111 6bICl]JO B03pacTaeT rrp11 MbIIIIe'IHOHpa6oTe;

CHa6)!(em1e rJIIOK030H KJieTOK, KOTOpbie rrp11 fOJIO)laH1111 B Ka'IeCTBe11CTO'IH11Ka3Hepnm: 11CI10Jlb3YIOT rrpe11MymecTBeHHO rJIIOK03y (HepBHbie KJieTK11, 3p111]JOU11Tbl, M03fOBOe Bern;eCTBO I10'leK, ceMeHH11K11).

XII. nEHT030<1>0C<l>ATHbl~

nYTb nPEBPAlUEH~s:I rntOK03bl

IleHT030¢ioc¢iaTHbIH rryTb, Ha3hIBaeMhIH TaK)!(e reKCOMOHo¢ioc¢iaTHbIM llIYHTOM, CJI~T aJibTepHaTHBHbIM rryTeM OKHCJieHHH fJIIOK030-6-¢ioc¢ia- Ta. IleHT03o¢ioc¢iaTHbIH IIYTb COCTOHT 113 2 ¢ia3

('rncTei1) - OKHCJIHTeJibHOH H HeOKHCJIHTeJibHOH. B OKHCJIHTeJibHoi1 ¢ia3e rJIIOK03o-6-¢ioc¢iaT tteo6paTHMO OKHCJUieTCH B rreHT03Y - p116yJI030-5- ¢ioc¢iaT, H 06pa3yeTCH BOCCTattoBJieHHhIH NADPH. B tteoKHCJIHTeJibHoi1 ¢ia3e p116ynmo-5-¢ioc¢iaT o6paTHMO npeapaIIIaeTCH B p116030-5-¢ioc¢iaT 11

MeTa60JIHTbl fJIHKOJIH3a.

IleHT03o¢ioc¢iaTHhii1 IIYTh 06ecrre'I11BaeTKJieTKH p11603oi1 ).J.JIH CHHTe3a nypHHOBbIX H I1HPHM11)lHHOBbIX HyKJieOTH)lOB 11 fH.lIPHPOBaHHbIM KO- ¢iepMeHTOM NADPH, KOTOph1i1 11crroJih3yeTcH B BOCCTaHOBH:TeJibHbIX rrpoueccax.

CyMMapttoe ypaBHettne neHT03o¢ioc¢iaTttoro fiYTH Bblpa)!(aeTCH cneeyIOW:HM o6pa3oM:

3 fJIIOK030-6-¢ioc¢iaT + 6 NADP+ ~ 3 co2 + 6 (NADPH + H+) + 2 <l>PYKT03o-6-¢ioc¢iaT + f JIHUepaJib,!IefH.lI-3-¢loc¢laT.

<I>epMeHThI neHT03o¢loc¢iaTHOro JIYT11, TaK )Ke, KaK H ¢iepMeHTbl rJI11KOJIH3a, JIOKaJI1130BaHhl B U11T030Jie.

Ha116onee aKT11BHO neHT03o¢ioc¢iaTHhii1 IIYTh npoTeKaeT B )JillpOBOH TKaH11, rre'leH11,Kope Ha,JJ,- I10'le'IHHKOB,3PHTPOUHTaX, MOJIO'IHOH)!(eJie3e B nepHO.lI JiaKTaUHH, CeMeHHHKax.

A. OKIICJIHTEJlbHblH 3TAII

B OKMCJIHTeJibHOi1 'IaCTH neHT03o¢ioc¢iaTttoro TIYTH fJIIOK030-6-¢ioc¢iaT I10.lIBepraeTC51 OKHCJIHTeJibHOMY Jl:eKap60KCHJIHPOBaH11IO, B pe3yJihTaTe KOToporo o6pa3YJOTC51 rreHT03bl. 3TOT 3Tan BKJIIOqaeT 2 peaKUH11 Jl:ern,Il.pHpOBaHHH.

IlepBaH peaKUHH ll:eflf.lIPHPOBaHHH - rrpeaparuettHe fJIIOK030-6-¢loc¢laTa B rJIIOKOHOJiaKTOH-6- ¢loc¢laT - KaTaJIH3HpyeTCH NADP+-3aB11c11Moi1 rnIOK03o-6-¢ioc¢iaT)J,ern,D,poreHaJoi1 11 corrpoBO)!(- ll:aeTcH OKHCJieHHeM aJibll:erHJl:HOH rpyrrrrbI y nepBOfO aTOMa yrnepOJl:a 11 o6pa30BaH11eM O,Ll,HOH MOJieKYJibl BOCcTaHOBJiettttoro Ko¢iepMeHTa NADPH.

)J,anee fJIIOKOHOJiaKTOH-6-¢ioc¢iaT 6bICl]JO rrpeBpamaeTCH B 6-¢ioc¢iormoKOHaT rrpn yqacTHH ct>epMettTa fJIIOKOHOJiaKTOHfH.lIPaTa3bl.

<I>epMeHT 6-ct>oc¢iornIOKOHaT,IJ,ern.u.porettaJa KaTanm11pyeT BTOpyIO peaKUHIO ,LJ,erH)J,pHpOBaHHH OKHCJIHTeJibHOH 'IaCT11,B xo.ue KOTopoi1 rrp011CXO)J,HT TaK- )i(e H )J,eKap6oKCHJI11POBaHJle. IlpH 3TOM yrnepO,!IHaH uerrh YKOPa'IHBaeTCHHa oa11tt aTOM yrnepo)J,a, o6pa3YeTCH p116yJI030-5-¢ioc¢iaT 11 BTOPaH MOJieKYJia fH)J,- pHpOBaHHOro NADPH (pnc. 7-62).

BocCTaHoBJieHHhIH NADPH HHrn6npyeT nepBbIH ¢iepMeHT OKHCJIHTeJihHOfO 3Tarra rreHT03o¢ioc¢iaTHOf0 IIYTH - rJIIOK03o-6-¢ioc¢iaTJl:ernaporetta3y. IlpeBpamett11e NADPH B oKHcnettttoe cocTOHHHe NADP+ rrp11BO.lIHT K ocna6nett1110 11ttrn611poBaHHH ¢iepMeHTa. Ilp11 3TOM CKOPOCTb COOTBeTCTBYIOW:ei1 peaKU11H B03pacTaeT, H o6pa3yeTCH 60JihIIIee KOJI11- 'IeCTBONADPH.

C)'MMapuoeypasueHHe OKHCJIHTem.uoro 3Tillla rrettTmoct>occt>aTttoro IIYT11 MO)l(HO rrpe.lICTaBHTb B BH)le:

fJIIOK030-6-¢ioc¢iaT + 2 NADP+ + H20 ~ P116y- J1030-5-¢ioc¢iaT + 2 (NADPH + H+) + COr

PeaKUHH oKHCJIHTeJibHoro 3Talla CJly)KllT ocHOBHbIM HCTO'IIDIKOMNADPH B KJieTKaX. fH)lp11po-BaHHb1e Ko¢iepMeHTbI CHa6)i(a!OT BO)J,Opo,ll,OM 611ocHHTeTH'IeCKHe rrpouecCbI, OKHCmueJibHO-BOCCTaHOBHTeJibHbie peaKUHH, BKJIIO'laIOIUHe3aIUHTY KJleTOK or aKTHBHbIX ct>opM KHCJIOpo)J,a. NADPH KaK Jl:OHop BO.lIOpoJl:a yqacTByeT B atta6on.H'IecKHXrrpou.eccax, Harrp11Mep B C11HTe3e xonecTepHHa. 3To HCTO'IHHKBOCCTaHOBHTeJibHbIX 3KBHBaJieHTOB )J,J1H U.HTOXpoMa p450' KaTaJIH311PyIOIUero o6pa30BaHMe rn.u.poKCHJlbHbIX rpynn np11 CHHTe3e CTePOH.lIHblX ropMOHOB, )l(fuNHbIX KHCJIOT, rrp11 KaTa60JI113Me JieKapCTBeHHbIX BemecTB 11

.lIPYfHX l!Y)i(epO)lHbIX coe)lHHeH11H (CM. pa3,ll,eJibI 8' 11, 12). BbICOKaH aKTHBHOCTb cl>epMema rJ110K030-6- ¢ioc¢iaTJl:ern.u.poreHa3hI o6ttapyxetta B ¢iarou.m11py10rn;HX nei1KoLI,HTaX, r)J,e NADPH-oKCH)J,aJa 11crroJib3yeT BoccrnttoBJieHHhrn NADPH )J,J1H o6pa30BaHHH cyrrepoKCH)J,HOfO 110Ha 113 MOJieKYJIHPHOfO KMCJIOpoJl:a. CyrrepoKClUUfhIB HOH rettepJiPyeT .lipyrne aKTHBHhie ¢iopMbI KHCJIOpoaa, IIO)J, Jl:eHCTB11eM KOTOpbIX H

358

1 -------- 1

 

 

 

 

 

 

 

I

H,

/,

Q

I

r JllOK030-6-¢oc¢aT-

 

 

 

I

 

I

 

 

 

I

 

Cy

 

I

A9rll1APOreHa3a

 

 

 

I----+-----I

 

 

 

 

H- C-OH

 

 

 

 

 

rrnoKOHOnaKTOH-

 

 

I

 

 

 

 

 

 

 

HO-C-H

 

 

NADPH+H+

 

rHApaTa3a

 

 

I

 

 

 

 

 

 

 

 

 

 

H-C-OH

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

Mg2+ 1-1n111 Ca2+

 

 

 

 

H-C-OH

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

H2C-OP03H2

 

 

 

 

 

 

r JllOK030-6-¢Joc¢JaT

 

rnt0KOHOnaKTOH-6-¢oc¢JaT

 

r------1

 

 

 

 

 

 

 

IQ

 

I

6-<1>oc¢ornlOKOHaT-

 

 

 

 

:

II

 

:

 

CH20H

 

IC-OH I

A9rll1APOreHa3a

 

'-+-----1

 

 

 

 

I

 

H-C-OH

 

 

 

C=O

 

 

I

 

 

 

NADPH+H'

I

 

HO-C-H

 

 

H-C-OH

 

H-C-OH

<(

 

 

H-C-OH

 

 

I

 

 

 

 

 

 

I

 

 

I

 

 

 

C02

I

 

H-C-OH

 

 

 

 

H2C-OPQ3H2

 

 

I

 

 

 

 

 

 

 

 

 

 

H2C-OP03H2

 

 

 

 

 

 

 

6-<1>oc¢ornt0KOHaT

 

 

 

P1-16ynoso-5-¢oc¢aT

P111c. 7-62. OK111cn111TenbHblM 3Tan neHTosoQ>ocQ>aTHoro nyT111.

noBpe)l(,!l,aJOTC51 MOJieJ<YJihI ,UHK, 6eJIKOB, JllUilf)lOB 6aKTepmUihHhIX KJieTOK. Cmrre3 )l(lfpHhrx KMCJIOT tt3 yrneBO)lOB B neIJeHM 5IBJUleTC51 OCHOBHhIM nyreM Y™- Jltt3aI..l,Hl1 NADPH tt 06ecneq1rnaeT pereHepaumo oKHcJieHttoti ¢opMhI NADP+. B neIJeHH rmoKo3o-6- <t>oc<t>aT)lerlf)lporettaJa, KaK tt KJIIOIJeBhle <l:>epMeHThI fJIMKOJil13a l1 6ttOCHHTe3a )l(lfpHhIX KllCJIOT, llH)lyI..l,ll- pyerc51 nptt yBeJIWieHllll COOTH01lleHH51 HHCYJIHH/rmoKaroH rroCJie rrptteMa 6oraToi1 yrneBo)laMH nttlll,ll.

HecMoTp51 Ha TO, '!TONADPH 06pa3yeTrn TaK- )[(e rrptt OKllCJieHllll MaJiaTa )lO rrttpyBaTa ll .UMOKClf)la yrnepo.ua (rrptt yqacTtttt NADP+-3aBttCttMoi1 MaJiaT.uertt.uporeHa3hI) tt )lertt.upttpoBaHtttt tt3outtTparn (rrptt y4acnt11 NADP+ -3aBllCHMOH l130- I..l,HTpaT.uertt.uporeHa3hl), B 60JihllIHHCTBe CJiyqaeB IIOTpe6HOCTll IUieTOK B BOCCTaHOBHTeJihHhlX 3KBllBaJieHTax y.uoBJieTBOp5IIOTC51 3a c'!eT nettTo3o- <t>oc<t>arnoro nyTtt.

PeaKutttt OKHCJittTeJihHoro nyTll rrpoTeKaIOT TOJihKO B TOM CJiyqae, eCJIH BOCCTaHOBJieHHhJH KO- <t>epMeHT NADPH B03BpamaeTC51 B ttCXO)lHoe OKttcneHHoe cocTOJIHtte NADP+ rrptt yqacTtttt NADPH-3aBttCHMhIX )lern)lporetta3 (T.e. rrptt ye-

JIOBHtt ttcrroJih30BaHttJI rn.upttpoBaHHOro NADPH B BoccrnttoBttTeJihHhIX npoueccax). Ecntt rroTpe6HOCTtt KJieTKll B NADPH tte3ttaIJttTeJihHhI, ptt6o- 30-5-<Poc<t>aT 06pa3yeTc5I B pe3yJihTaTe o6paTttMhIX peaKUHH HeOKllCJittTeJihHOro 3Tana rreHT03o<Poc- <t>aTHOro nyTtt, llCIIOJih3Y51 B Kaqecrne llCXO.UHhIX BemeCTB MeTa60JIHThl rJillKOJitt3a - rnttuepaJih-

.uern.u-3-<t>oc<t>aT tt <PPYKT03o-6-<t>oc<t>aT.

Ii. HEOKMCJIHTE.JihHhIH 3TAil

HeoKttCJittTeJihHhIH 3Tarr rrettT03o<Poc<t>aTHoro rryTll BKJIIOIJaeT cepllIO o6paTHMhlX peaKUHH, B pe3YJihTaTe KOTOphIX ptt6yJio3o-5-<Poc<t>aT npeBpamaeTcJI B ptt603o-5-<Poc<t>aT tt KCHJIYJI03o-5-<Poc<t>m, tt )laJiee 3a cqfa nepettoca yrnepo.utthIX <t>parMeHTOB B MeTa60JIMThI rJIMKOJill3a - <l>PYKT030-6-<t>oc<t>aT tt nmuepaJih.Uerlf)l-3-<t>oc<t>aT. B 3Tttx rrpeBpameHIDIX rrpttHHMaIOT yqaCTlle <t>epMeHThI: 3IIHMepa3a, ll30Mepa3a, TpattcKeToJia3a n TpaHCaJih.UOJia3a. TpaHcKernna3a B KaIJecrne Ko<t>epMeHTa ttcrronh3yeT TttaMttH.11.M<Poc<t>aT. HeoKMCJIITTeJihHhitt 3Tarr rrettT03o<Poc<t>arnoro rryTil He BKJIIOIJaeT peaKUtttt .ue-

359

).lJUI

rl1)J,pl1pOBamrn H 003TOMY l1CITOJlb3yeTCH TOJlbKO CHHTe3a neHT03.

P116yJI030-5-cjJoccjJaT C.lly)Kl1T cy6cTPaTOM )lJIH ll.Byx cjJepMeHTOB. <l>epMeHT p116yJI030-5-cjJoc<t>aT-3-3IUf- Mepa3a l13MeHHeT CTeXHOMeTPHYeCKOe IIOJIO)l(eH11e 011.ttoti OH-rpynnhI y TPeTbero aToMa yrnepo1J,a, npeBpamaH p116yJI03o-5-<Poc<PaT BKC11JIYJI03o-5-<Poc<t>aT. Ll,pyroti <PepMeHT - p116yJI03o-5-<Poc<t>aT-H30Mepa- 3a - KaTaJIH3ttpyeT npespameHHe p116yJJ030-5-<Poc- <Parn B p11603o-5-<Poc<PaT (p11c. 7-63). P11603o-5- <Poc<PaT, o6pa3yJOIUHHCH B HeOKHCJil1TeJJhHOH <Pa3e, o6ecnel.JHBaeT KJieTKH p116030H, Heo6XO)J,l1MOH ).lJUI CHHTe3a HyKJJeOTHJJ,OB, KOTOpbie CJI)')!(aTnpe)J,weCTBeHHHKaMH H CTPYKTYPHhIMl1 KOMITOHeHTaMH KO- <PepMeHTOB )J,erH)J,poreHa3 H HYKJieHHOBbIX KHCJIOT.

<l>epMeHTbI TPaHcKeToJJma 11 TPaHCaJibJJ.OJia3a KaTaJIH3HPYJOT nepeHOC ll.BYX- H nJexyrJiepoJJ,HbIX <t>parMeHTOB, COOTBeTCTBeHHO HCITOJJb3YH B Ka'-lecrne11,0- HOpa yrnepO)J,HbIX <t>parMeHTOB KeT03y, a aJib)J,03Y - B KaYecrne aKuenrnpa. 3TH peaKUHH rrpoTeKaJOT B 2 3Tana: cttaYaJia npo11cxo1J,ttT OTIUeIIJieH11e yrnePOll.HOro <PparMeHrn OT MOJJeKyJibI-JJ.OHopa, a 3aTeM - nepeHOC 3TOro <t>parMeHTa Ha MOJieKyJJy, BblIIOJIHHIOIUYIO pOJJb aKuenTopa. TpaHcKeToJia3a B HeOKHCJIHTeJJhHOH <Pme rreHT030cPOCcPaTHOro nyrn KaTaJJtt311pyeT 2 peaKUHl1. B nepsoti peaKu1111 (p11c. 7-64) TPaHcKeTOJia3a pacmerrm1eT CBH3b C-C Me)l(ll.y Kernrpynrrotf tt coce11,H11M aTOMOM yrJiepona B MOJJeKyJJe KCl1Jl)'J1030-5-<Poc<PaT,B pe3yJJhTaTe Yero KeTocaxap rrpespamaeTCH B aJihll.03y, r.JIMUepaJibll.e- rl1)J,-3-<Poc<PaT, conep)l(aIUYIO Ha 2 aToMa yrnepona MeHhWe. 06pa3yJOIUl1HC5I IIOCJie pacmeIIJieHttH ,llByxyrnepO,llHhlH $parMeHT OCTaeTCH KOBaJieHTHO CB5I3aHHbIM B KaTaJil1TH'-!eCKOMueHTPe <J:>epMeHTa c KO- <PepMeHTOM Tl1aMHH.Utt<Poc<t>aTOM. Ll,anee <PepMeHT nepeHOCl1T JJ,ByxyrJiepO)J,HbIH <J:>parMeHT Ha aJib)J,e- fl1)J,HYIO rpynrry aJib)J,OCaxapa, o6pa3yJO HOBYJO KeT03Y - cenoremyJio3o-7-<Poc<J:>aT.

TpattCaJJb.llOJia3a rrepeHoc11T Tpexyrnepoll.HhIM <PparMeHT OT ce1J,orenTyJI030-7-<Poc<Parn Ha rm1- uepanb,llern1J,-3-<Poc<PaT, 06pa3y51 3Pl1TP03o-4-<Poc- <PaT 11 cPP)'KT03o-6-<Poc<PaT(pttc. 7-65).

3Ta peaKUl15I IIO,ll06Ha peaKUlfH aJJb)J,OJlhHOro pacweIIJJeHl1H rnttKOJIHTwrecKoro IIYTH, 3a ttCKJJIOYetttteM TOro, '-!TOB ,llaHHOM CJIYYae Tpexyrnepo)J,- HblH <t>parMeHT, CO,llep)l(aIUHM KeTorpynrry, rrepeHOCHTCH Ha anb11,ocaxap rn11uepanb,llerl1)J,-3-<Poc<PaT, a B rJIHKOJll1Tl1'-!eCKOM nyTH KeTo<t>parMeHT BblCB060)l()J,aeTCH B Bl1ll.e ll.HrH11,poKcttauerntt<Poc<t>arn.

B cJJell.YIOlll.eti peaKUHH, KaTMl13l1pyeMoiI TPaHcKeTOJia3oi1, npOHCXO)J,HT nepeHOC ll.ByxyrJiepO,llHO-

ro <PparMeHTa OT KCHJIYJI030-5-<Poc<t>arn Ha 3pHT- p03o-4-<Poc<t>aT. flpOll.YKTaMH 3TOM peaKUl1H HBJIHIOTCH cPPYKT030-6-<Poc<t>aT H rnttuepaJJbJJ,em)J,- 3-<Poc<t>aT (pl1c. 7-66).

TaK KaK see peaKUHH HeoKHCJJttTeJJbHOro 3Tarra o6paTHMhr, 06pa3osatt11e p116030-5-<Poc<Parn MO)l(eT IIPOHCXO)J,HTb He TOJlbKO B pe3yJJbTaTe 1130MepHoro rrpespalll,eHl15I IIPO.llYKTa OKHCJJHTeJibHOM <Pa3bl rreHT030cPOCcPaTHOro IIyTH p116yJJ030-5-<Poc- <Parn B pl16o3o-5-<Poc<PaT no)], 1J,eticrnl1eM H30Mepa3bI, HO TaK)l(e l1 113 npoMe)l(yTO'-!HbIXrrpo11,yKTOB rn11KoJI113a - cPPYKT030-6-<Poc<Parn tt rn11uepanb- ll.erl1ll.-3-<Poc<Parn. flocJJe11,osaTeJJhHOCTh rrpespameHHM, npHBO)J,Hlll,HX K o6pa30BaHHIO p11603o-5- <t>oc<t>arn 113 TaKHX rrpO.llYKTOB rJIHKOJIHTlflieCKoro nyTH, MO)l(HO npe)J,CTaBHTb B BH)J,e:

2 <DpyKToJ0-6-<Poc<PaT + fnHUePaJibJJ.erl1)J,-3-<Poc<PaT ~ 2 KC11JIYJJ03o-5-<Poc<PaT + P116030-5-<Poc<PaT

2 KCHJIYJJOJo-5-<Poc<PaT ~ 2 P116yJI03o-5-<Poc<PaT 2 Ptt6yJJo3o-5-<Poc<J:>aT ~ 2 Ptt603o-5-<Poc<PaT.

CYMMapub1H: pe3yJibT3T Mern6oJJH3Ma 3 MoJieKyJJ ptt6yJI030-5-<J:>oc<Parn B HeOKHCJJHTeJibHOM <Pa3e rreHT030cPOC<i>aTHOro IlYTH - o6pa30BaHHe 2 MOJieKyJJ cPPYKT03o-6-<Poc<Parn 11 1 MOJJeKyJibI rn11ue- panhll.ellll.-3-<Poc<Parn. Ll,anee cPPYKT03o-6-<Poc<PaT 11 rm1uepanb)lerttll.-3-<Poc<PaT MOryT npespaTHTbCH B rJJIOK03y. c yyfaoM CTeXHOMeTPWieCKOro K03cPcPHUHeHTa, paBHOro 2, MH o6pa30BaHHH 5 MOJieKyJJ [JIJOK03hl (CO)J,ep)l(alll,HX 30 aTOMOB yrnepo,lla) IIOTPe6yJOTCH 4 MOJJeKyJibl cPPyKT030-6-<Poc<t>arn H 2 MoJieKyJibI rnttuepanhll.erH)J,-3-<Poc<Parn (B cyMMe co11,ep)l(alll,l1e TaK)l(e 30 aTOMOB yrnepo.ua) HJitt, coorneTcrneHHO, 6 MoJJeKyJJ p116yJI03o-5-<l>oc<Parn. TaKHM o6pa30M, HeOKHCJIHTeJJhHbIH IlyTb MO)l(HO rrpell.CTaBHTb KaK rrpouecc B03Bpaw,eHHH neHT03 B <J:>oH)l reKco3.

B. IIEHT030<l>OC<l>ATHhIH QMKJI

0KHCJllfTeJibHbIH 3TaII o6pa30BaHHH neHT03 H HeOKHCJll1TeJibHbIH 3TaII (IIyTb B03BpameHlfH rreHT03 B reKC03bI) COCTaBJJHIOT BMeCTe UHKJIH'-leCKHH rrpouecc.

TaKoit rrpouecc MO)l(HO om1caTb 06m11M ypasHeHlfeM:

6 fJIIOK030-6-<Poc<t>aT + 12 NADP+ + 2 Hp ~

5fJJJOK03o-6-<J:>oc<J:>aT + 12 NADPH +12 H+ +

6C02

3To 03Hal.JaeT, 'ITOH3 6 MOJJeKyJI rJJIOK03bl o6pa3YJOTCH 6 MOJieKyJI ptt6yJI030-5-<J:>oc<t>aT (rreHT03bI) H 6 MOJieKyn C02 . <l>epMeHTbI HeOKHCJJHTeJibHOH

360

7-67).

 

CH20H

 

I

 

C=O

 

I

 

HO-C-OH

 

I

 

H-C-OH

 

I

 

H2C-OP03H2

 

3m1Mepa3a

CH20H

Kc11ny.no3o-5-cj:Joccj:JaT

 

I

 

C=O

 

I

 

H-C-OH

 

I

 

H-C-OH

1113oMepa3a

I

HzC-OP03H2

CHO

 

 

I

P116yno3o-5-cj:Joccpar

H-C-OH

 

I

 

H-C-OH

 

I

 

H-C-OH

 

I

 

H2C-OP03H2

 

P116030-5-cj:Joccpar

Pl'IC.7-63. npespa~eHl'ISI pH6yn030·5-<l>oc¢1aTa.

 

, ___;_,.

 

_

'

 

 

 

I

 

 

/

 

"\...

 

 

 

CH20H

I

 

 

I

 

 

 

C=O

CH20H

H, /.0

 

 

 

I

:

 

 

 

I

1

I

 

1

c'

 

 

 

HO-C-OH

: C=O

 

I

I

 

 

 

I

c

-r- _____I

H - C- OH

TpaHcKerona3a

H-C-OH

HO-C-OH

+

I

+

I

H-C-OH

 

I

 

 

H-C-OH

 

 

 

H-C-OH

 

I

 

 

 

I

 

I

 

 

H-C-OH

 

TACI>

H-C-OH

H2C-OP03H2

I

 

 

 

I

 

 

 

 

H2C-OP03H2

 

 

 

H2C-OP03H2

Kc11nyno30-5-cpoccpar

P116030-5-cj:Joccpar

 

rn114epoanb.Qern,Q-3-cpoccpar

Ce.QorenTyno3o-7-cpoccj:Jar

PHc. 7-64. PeaKl,41'1SInepeHoca AByxyrnepoAHoro ¢iparMeHTa, KaTanl'l3HpyeMasiTpaHcKeTona301ii.

c}la3hI npeBpaw.a10T 6 MOJieKYJI p116yJI03o-5-c}locc}laT B 5 MOJTeKyJI rJIIOK03hl (reKC03bl). IlpH JlOCJie)J,OBaTeJibHOM rrpoBe)J,eHHH 3THX peaKUHH e)J,HHCTBeHHhIM noJie3HhIM rrpOJJ.YKTOM 5IRIDieTC5l NADPH, 06pa3y10lll,HHC5l B OKHCJIHTeJibHOM c}la3e neHT030cPOCcPaTHOro nyn1. TaKotl npouecc Ha3hTBaJOT neHT03oc}loc- <ParnhIM QMKJTOM (pHC.

TipoTeKaHHe neHT030cPOCcPaTHOro UHI<Jia TI03BOJI5leT KJTeTKaM rrponyuwposaTb NADPH, tteo6xo-

JJ.HMhIH )J,Jl5l C.11HTe3a )((.11p0B, He HaKaIIJTHBa5l neHT03hl.

3tteprw5l, BhIJJ.eJI5l10lll,a5IC5l np11 pacnane rmoK03hI, Tpattcc}lopM.H:pyeTC5l B 3HeprHIO BhICOK03HepreTH'IecKoro)lOttopa BO)lopona - NADPH. fwnpwpoBatttthIH NADPH cJT~MT wcToqttwKOM BO)lOpO)la )11151 BOCCTaHOBHTellhHbIX CHHTe30B, a ::itteprn51 NADPH npeo6pa3yeTC5l w coxpam1erc51 BO BHOBb CHHTe3ttpOBaHHhIX BeW,eCTBax, Hanpw-

361

)J,H K,

 

/

 

 

'

 

 

 

 

 

 

 

 

 

 

r----..L-----.,

 

''

 

 

 

 

 

 

 

 

 

I

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

'

 

 

 

 

 

 

 

 

 

I

CH20H

:

 

 

\

 

 

 

 

 

 

 

 

 

 

 

\

 

 

 

 

 

 

 

 

 

I

I

I

 

 

\

 

 

 

 

 

 

 

 

 

:

C=O

I

 

 

l.

 

 

 

 

 

 

H,c,,,,o

 

CH20H

 

 

H, ,,_,0

 

 

 

 

I

I

I

 

 

 

 

 

 

 

I

1 HO-C-H

I

 

 

c

 

 

 

 

 

 

I

 

C=O

I

J ______ ,I

 

 

 

TpaHcanbAonaJa

 

 

H-C-OH

 

+

 

1

 

H-C-OH

+

I

 

 

 

H-C-OH

 

 

 

I

HO-C-H

 

I

 

 

 

I

 

 

 

 

 

 

H-C-OH

 

I

 

H-C-OH

 

 

 

 

 

 

 

 

 

 

H-C-OH

 

 

 

 

H2C-OP03H2

 

 

 

I

 

 

I

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

H2C-OP03H2

 

H-C-OH

 

 

 

 

 

 

 

 

 

 

H--C-OH

 

I

 

 

 

 

 

 

 

 

 

 

 

 

I

 

H2C-OP03H2

 

 

 

 

 

 

 

 

 

 

 

HzC-OP03H2

CeAoremynoJo-7-c:j:>occ:j:>ar lmu-1epanbAen1A-3-c:j:>occ:j:>ar

 

 

3pl-1rpoao-4-c:j:>occ:j:>ar

<l>pyKToao-6-c:j:>occ:j:>ar

PMc. 7-65. PeaKL1MS1, KaTanM3MpyeMasi TpaHCanbAonaaolt.

 

 

 

 

 

 

 

/

 

 

 

'

 

 

 

 

 

 

 

 

 

 

../

 

, --'-----

 

 

 

 

 

 

 

 

H,

;;O

 

1

 

CH20H

I

 

 

 

 

 

CH20H

 

I

 

:

 

 

 

 

H,

;;O

 

 

c"

 

I

 

I

1

 

 

 

 

 

I

 

I

 

: C=O

I

 

 

 

c"

 

C=O

H-C-OH

 

•-i------' TpaHcKernnaaa

 

I

 

I

 

I

+

HO-C-OH

 

 

 

 

 

H-C-OH

+

HO-C-H

 

 

 

 

 

 

I

H-C-OH

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

I

 

H-C-OH

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

H2C-OP03H2

 

 

 

H-C-OH

 

T,Q<l>

 

 

 

I

HzC-OP03H2

 

 

 

I

 

 

 

 

 

 

H-C-OH

 

 

 

 

 

 

 

 

 

 

 

 

 

 

H2C-OP03H2

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

 

H2C-OP03H2

3pL-npoao-4-c:j:>occ:j:>ar

KcL-mynoao-5-c:j:>occ:j:>ar

lnl-11.1epoanbAen1A-3-c:j:>occ:j:>ar

<l>pyKToao-6-c:j:>occ:j:>ar

PMc. 7-66. PeaKL1MS1, KaTanMaMpyeMasi TpaHcKeTonaaolt.

Mep )f<I1:pHhIX KHCJIOTaX, BhICB06mK,naeTC5I rrpH HX

KaTa6omf3Me H HCIIOJih3yeTCH KJieTKaMH.

r. ,Il,E<l>EKT fJIIOK030-6-<1>oc<1>AT

)J,Efl1)1,POfEHA3hl B 3PllTPOll,lffAX

He<f>epMeHTaTHBHoe OKHCJieHHe reMorno611Ha (Fe2+) B MeTreMOfJI06HH (Fe3+) IIPHBO,lIHT K 0,lI-

H03JieKTpOHHOMY BOCCTaHOBJieHHIO KHCJIOpO,lia H IIOHBJieHHIO peaKIJ,HOHHO-CIIOC06Horo aHHOH-pa- ,lIHKaJia - cyrrepOKCH,lia 02-' KOTOphIH CJIY)f(HT npe,lirnecTBeHH11KOM ,lipyrHX aKTHBHhIX <f>opM KHCJiopo.na: nepoKcll.lia BO,liopo,lia H 20 2 11 rll,lipOKCHJihHoro PMMKaJia OH-. AKTHBHhie <f>opMhI KHCJiopo- ,lia 5IBJI5IIOTCH CHJihHe:HrnHMH OKHCJIHTeJI5IMH H II03TOMY crroco6Hhl Bhl3hIBaTh cephe3Hhie IIOBpe)f(- ,lieHHH MoJieKYJI 6eJIKOB, HeHachIUieHHhIX JIHIIH,lIOB.

B 3pHTpouttrnx, KaK u B 6oJihWttHcrne KJieTOK, rrp11cyrcrnyeT THOJICO,liep)f(aUIHH Tp11IIeIITH,lI - rnyTaTHOH (y-rnyraMHJI-uHcTeHHJI-rnttUttH). BoccrnHOBJieHttaH <f>opMa rnyraTHOHa (f-SH) CO,liep)f(llT SH-rpyrrny (pttc. 7-68), KOTopa51 MO)f{eT cJiy)f(llTh ,lIOHOpOM :rneKTpOHOB B peaKUH5IX BOCCTaHOBJieHHH. CTo.n ,lieHCTBHeM ¢>epMeHTa rnyraTHOHTTepOKCH,lia3hI BOCCTaHOBJieHHhlH myran-mtt npeBpamaeT MOJieKYJIY rrepOKCll,lia BO.UOpo.na B MOJieKYJIY BO,lihI, a caM nepeXO,lIHT B OKHCJieHHOe COCT051Hlle (f-SS-f). Perettepauu10 BOCCTaHOBJieHHOfO rnyraTHOHa o6ecneqttBaeT rnyTaTHOHpe,liyKTa3a, HCTTOJih3YH B KaqecTBe

.noHopa BO,liopo,na rll,lipttpoBaHHhIH NADPH. )],JIH

3pHTpOIJ,HTOB e,lIHHCTBeHHhIM HCTO'-IHHKOMrronyqe- HH5I NADPH CJI~ rreHT03o¢>ocq>aTHhIH flYTh, ,lIJIH

,lipyrn:x TKaHe:H cymecrnyeT aJihTepHaTHBHhIH cnoco6 - rrp11 yqacTttH NADH-3aBttCHMOH MaJiaT,lierll,liporeHa3hI (MaJIHK-<l>epMeHT).

362

 

 

 

 

 

(6) r moKoao-6-cpoccpar

 

 

 

(1)rntoKoao-

 

//~

 

h 12 NADPH+12W

6-cpoccpar

''-~~~:::Y/

 

 

~ 6C02

 

 

 

 

 

 

 

 

 

 

 

 

(5)

 

//////

 

(6) P1116ynoao-5-cpoccpar

 

 

 

rntoK030-

 

 

 

 

 

l8n1t1Mepa~I

~

1

~ l~1t1Mepaaal

-6-cpoccpaT

 

 

 

 

/

 

ltlaoMepaaa

 

~

 

t

 

 

 

 

 

 

 

 

 

 

 

 

(2)Kc111nynoao-

(2) P1116oao-5-cpoccpar

(2)Kc111nynoao-

 

 

 

5-cpoccpar

y IT:

Cs

 

5-cpoccpar

 

 

 

 

Cs

 

Cs

 

 

 

 

 

 

 

 

 

 

 

 

 

 

~aHe<erona'8.

 

 

 

I

 

(2) rn1114epa111>,Qer1t1,Q

(2) Ce,Qoremynoao-

 

 

I

 

 

3-cpoccpar

 

 

 

 

 

7-cpoccpar

 

 

 

I

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

I

 

 

ITpattca11b,Qo11aaa J

 

 

 

 

I

 

 

 

 

 

 

I

 

 

 

 

 

 

I

 

 

 

 

 

(2) 8p1t1lp030-

 

 

 

I

 

 

 

 

 

 

 

 

I

 

 

 

 

 

4-cpoccpar C4

 

 

 

I

 

 

 

 

 

 

TpaHcKeTonaaa

I

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

I

 

(2) cJ>pyKT030-

 

(2) cJ>pyKT030-

 

 

 

I

 

 

 

(2) rmu-1epallb,Qer1t1,Q

I

 

 

6-cpoccpaT

 

6-cpoccpaT

 

I

 

 

 

 

3-cpoccpar

 

 

 

C5

 

 

 

~I

 

 

I

 

 

 

 

1C3

I\

 

I

 

 

 

 

 

~'

 

I

 

 

 

 

 

 

11

---------

 

 

 

 

cJ>pyKT030-

! \

 

 

 

 

 

 

 

I

\

 

 

 

 

 

 

1,6-61t1ccpoccpaT

I

'

 

 

 

 

 

 

I

 

~------------------------------~

 

 

 

I

I

I I I

I

Cs

l

''

·

cJ>pyKT030-

6-61t1CcpoccpaT

 

 

C5

'MC.7-67. neHT030<j>oe<j>aTHblM l.IMKll B )l(MpOBOM TKaHM.

B3att:Mo.uetiCTBtt:e BoccrnttoRJieHttoro r.nyrarnotta : rrepoKctt:,ri:oM Bonopo.ua B 3pHTpou:irrax rrpe,UoxpaNeT UMCTeMHOBbie OCTaTI<ll: B rrpoTOMepax reMOrJIO- )tt:Ha OT OKHCJiemrn. Tip11 reHeTWieCKOM ,Ue<PeKTe :'JIIOK030-6-<Poc<Pani:er11,UpOreHa3blKOHUeHTPaUIDI :IOCCTaHOIDieHHoro Ko<PepMettrn NADPH YMettbllla- ~Tc51, B pe3yJihTaTe qero pe3KO CHIDKaeTC51 KOHUeHTJauH51 BOCCTaHOIDieHHOfO r.nyrarnoHa, a B KJieTKe, ;ooTBeTCTBeHHO, ~JIWil1BaeTC51 KOJIWieCTBO aKTMBtiblX <PopM KHCJiopo,Ua. B 3TOM c~ae OKHCJiett11e SH-rpynrr MoneKYn reMorno611tta B 3p11Tpou11Tax

rrptt:BO)ltt:T K o6pa30BaHMIO rrepeKpecTHblX lll1CYJib- <l>M.D:HblX CB513eti tt: arperautt:tt: rrpornMepoB reMorno611tta c <PopM11poBatt11eM Teneu Xatittua (CM. pa3-

.ueJI 14). B rrp11cYTCTBMM reneu Xatittua TIJiaCTWIHOCTb MeM6patthI ttapYlllaeTC51, 11 otta Tep51eT crroco6ttocTb K JJ:e<PopMau1111 rrp11 rrpoxo)l(,[(eHMH 3p11TpOUMTOB <Jepe3 KaI111JIJIHpbr. fun BhnhmaeT ttapYlllett11e ueJiocTHOCTl1 MeM6paHbI, 'ITOrrp11BO,UMT K reMomny 3PHTPOU11TOB. HeK0Toph1e JieKapcTBetttthre BeiuecTBa, ttarrp11Mep attTl1MaJI5Ip11tiHb1ti rrperrapm rrp11Max11H, cynh<PattMJiaMtt:,ri:hI, TaIOKe CHIDKaIOT crroco6ttocTb

363

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