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підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

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CH,

 

 

 

 

 

 

I

 

 

 

 

 

(CH, - CH = CH CH,), - H

 

 

 

 

I

 

 

 

 

 

CH,

?H,OH...

Yrneso,1:1opo.QH bl i1

 

 

 

pa,Q"1K8Jl

 

 

 

 

_ ..··

 

 

 

 

 

 

0 ·.·. 8 HC

~

CH 3

 

<l>opM"1llbH8ft );p _I;

qCH,

 

rpynna

H /

I

N-Fe-N

I

 

-coo-CH,-CH,

7 /:=Q:~~

4 CH=CH,

He- -cH

6 -- 5

CH, CH,

I

CH,

I coo-

PMc. 6-10. CTpOeHMe reMa A.

qacTbIO B TaK tta3brnaeMbIX CuA-uettTpax. Ilepettoc 3JieKTpOHOB KOMITJieKCOM a-a3 BKJIJOqaeT peaKUIUI:

Cu+ H Cu2+ + e,

Fe2+ H FeJ+ + e.

opraHH30BaHhI B 4 KOMITJieKCa, paCITOJIO)l(eHHhIX B orrpe,[(eJieHHOH ITOCJie,[(OBaTeJibHOCTH (BeKTOpHO). B 3TOH JlOCJie,[(OBaTeJihHOCTH HX CTaH,[(apTHbie OKHCJIHTeJibHO-BOCCTaHOBHTeJibHbie ITOTeHUHaJibI CTaHOBHTCH 6oJiee JlOJIO)l(HTeJibHbIMH no Mepe rrpvr6JII1)1(eHHH K KHCJIOpo):(y (rn6JI. 6-3, pHC.

6-11).

KoMTIJieKc u11ToxpoMoB a-a3 tterrocpe,[(cTBeHHO peampyeT c MOJieKym1pttbIM KHcnopo,[(oM. HeKoTOpb1e xapaKTepHCTHKH KOMITOHeHTOB Uil3 rrp11Be,[(ettb1 B rn6n. 6-4.

E. 0PrAHtt3AIJ,ttH ,IJ;bIXATF.JibHOH. UEIIH

B MHTOXOH,IJ;PHHX

OcttoBHhie rrepettOCl.[JIKH 3JieKTpottoB BCTpoeHbI BO BHYTPeHHIOIO MeM6patty MHTOXOH,[(pHH H

Ka)l(,[(oe 3Betto 3TOH uerr11 crreumpHl.[HO B oTttowettHH ,[(OHopa H aKuerrTopa 3JleKTPOHOB.

Ha rrepBoM 3Tarre ,[(erw):(poretta3bl KaTaJIH3Hpy10T OTW:eITJieHHe BO,[(Opo,[(a OT pa3JIH'IHhIXcy6CTpaTOB. EcJIH cy6cTpaTaMH cJiy)l(aT a-nr,[(pOKCH- KHCJIOThI MaJiaT, H30UHTpaT, 3-rH,[(pOKCH6yT11paT, BO,[(opo):( rrepettOCHTCH Ha NAD+. 06pa3oBasurnrtcH NADH B ,[(hIXaTeJihttotf uerr11, B CBOIO Ol.[e- pe):(b, OKHCJIHeTCH NADH-,[(ern,[(poretta3ort (KOMrrJieKc I).

Ta611M48 6-4. KoMnOHeHTbl MMTOXOHAPM81lbHOii 4enM nepeHoca 31leKTpOHOB

Ha1sauue KOMDOHeHTa

IlpocTeTuqecKaH rpynna

.ri;ouop e

A11:QenTop e

 

 

 

 

 

NADH-,IJ,ern,IJ,poreua3a,

FMN, FeS

NADH

KoQ

KOMIIJieKc I

 

 

 

 

 

 

 

 

 

Ko3H3HM Q, y6nxnuou

 

 

KoMnJieKc I

KOMIIJICKC III (bc1)

 

 

 

 

 

QHr,IJ,ern,IJ,poreuaJa,

FeS, reM b1 ( 562),

QH2

U)noxpoM c

KOMIIJICKC III

reM b2 (566). reM c 1

 

 

QIITOXpOM C

feM C

KoMnJieKc III

KoMIIJieKc IV

 

 

 

 

 

QHTOXpOMOKCII,IJ,a3a,

feM A

QHTOXpOM C

02

KOMIIJieKc IV

Cu2+

 

 

CYKJJ;HHaT,IJ,ern,IJ,poreuaJa,

FAD, FeS

CYKJJ;HHaT

KoQ

KOMrrneKc II

 

 

 

 

 

 

 

 

 

 

 

 

 

 

274

\1 ,,
1,5-2.5).
7-10).

1. AM1'1Il03a20%

P111c. 7-9. CTpoeH111e KpaxMana.

tJIVIKoreH npe.ucrns;rneT co6oi1 CTPYKTYPHbiii aHaJIOr KpaxMaJra, HO 111\tCeT 6oJJbWYIO CTeneHb sernnemrn: np1-1MepHo Ha Ka>Kllhre I0 ocTaTKOl:l rJJIOK03bI npHX01111TC51 011Ha a- l ,6-rnHK03MllHlHI CB513b.

II. nEPEBAPil1BAHlt1E

YrnEBO.QOB

3mne111-1a11httb1e KJieTKH KHWeY1mKa cnoco6HhI 13CaCbIBaTb TOJlbKO MOHOcaxapH1lbl. no3TOMY npoUCCC nepel:lap11satt1-15l JaKJJIOYaerrn l:l cj)epMettraTMBHOM r1-1J1pomne rJJl1K03M11HblX CB513ett B yrneBoLiax, HMCIOWHX 01mro111111 no;rncaxap1111ttoe cTpoe1-rne (p1-1c.

10 H H/ -- 0

\~_)L-\_)

r-H,O

lrn1K03V1Aa3b1

1 0\JOH

~HHO

P111c. 7-10. r111Apon1113 rn111K03111AHOH CB'.113111.

2. AML1rl0110KTl'1H80%

BoccrnHasn11s;;uo1.J.1i-1ii

KOHOLj

A. fIEPEBAPHBAlll1E YfJlEBOJJ,OB B POTOBOft

fJO.lOCTil

B poTOBOii nonocTM raHua ~ul\1e.:n»rncTc>i 1qw m> pc>KeBhJBaHml, C~f:l'IHBa51Ch npH HOM Cj!KJHOii.

CJJIOHa Ha 99% COCTOl1T HJ B0~1bl M o6bl'IHOM:\lCel pH 6,8. B UIOHC np11cyTcrnyer n111poJlHTw1ccKMi1 cj)epMCHT a-aMu.rmJa (a-1,4-rnttKOJHD,aJa). pacwen - J15llDillaH B KpaXMaJJe u- I ,4-r;rnK031-11lllble CB513H.

B poTmmw no110n11 11e MmKeT npm1cxo;unb noJIHoe pacwenJICHHe KpaX\1aJla, TaK KaK .u61cTGHC ttJcp- \1CJ-ITa 1-1a KpaxMafJ Kpmxol:lpcMeHHO. KpoMe Toro. ar-.111mna CJllOHhI He pacwenJUJCT a- I ,6-rJIHK0"31111-

Hh1e CB5l:H1 (CB5l3H ll \ICCTax paJuern;1em1w). mHTO-

MY KpaxM;rn

ncpenap11saeTC5l ;111111b LWCTMYHO c 06--

paJ0Bam10.1

KpynHblX q)parMCI !T013 - JlCKCTp111arn

11 1-1e60Jlbilloro KOJ!l1LieCTBa MaJlbT03bL C1e!lyCT orMeTMTh, LJTO aM11mna CJllDHbl HC rn11pomnycT L-ll1-

KOJH)lHhlC CBHJH l:l LlHCaxap1v.1,ax.

Jlei1cnrne aM11.1a3bI c:110HbI npeKpamancH 11 peJKO K~1CJIOi1 cpe11e CO!lep)I01\!0ro )KC,1YJlKU (pH On:HaKo rrnyrpM 1111LucBoro KOMKa aK-

TH BllOCT b a \1 lljJ a 3bl M O)K;'T JH~ KOTODOC I~ pc

coxpawnhcn., noKa pH He ;1 ;i11:.:x ,_; CTOpoHy. >KeJly!l(.)LJl-lblH COK He co,_1,ep)l(~IT ¢epMeH-

TOl3, pacwc11J1mmuMx yrnesoJ1h1. B )KeJiy110LJHO:\I co.uep>K11MOM B0'3MO)KeH Jll1Ulh HCJllaLJHTeJJbHblH KHCJJOTH1>111 rnnpomn rnHKOJl111HhIX cmnet1.

Ii. IIEPEBAPHBAHME YLTJEBO).l,OB

B Kl1lllE11HI1KE

nocJleLiytOWHe :nanbl ncprnap1mamrn Hepacwe11-- JieHHOr0 MJ1M lJ(ICTHYHO pacu.1.en.'leHHOroKpaxM<:Ula, a

305

II. OKlllCillllTEilbHOE

Cl>OCCl>OPlllillllPOBAHlllE A.Q<l>

TaK KaK 3JieKTpOHhI Bcer)J,a cTpeM>ITC51 ITepexo- )J,HTh OT 3JieKTpOOTpJ.IUaTeJihHbIX CHCTeM K 3JieKTPOITOJ10)!(HTeJihHbIM, 11x TpaHcITopT ITO UI13 K KHCJiopo,lJ,y COITPOBO)!()J,aeTC51 CHJ1)!(eHHeM CB060)J,HOM

3HeprH11.

I1p11 cpaBHeH1111 Be.JIWIHH 3JieKTPQXHM11qecKHX ITOTeHUHaJIOB nepeHocqMKOB 3JieKTpOHOB (Ta6JI. 6-3) Bl1)J,HO, qTQ CHJ1)!(ellfe CB060)J,HOM 3Hepfl1H npOMCXO)lHT Ha Ka)!()J,OM 3Tarre UI13, 11 3Hepr1151 3JieKTpo-

HOB Bhl)J,eAAeTC>I nopu1151MH.

BMeCTe c TeM B )J,hIXaTeJibHOM ueITH MO)!(HO BbI- )J,eJIHTb 3 yqacTKa, B KOTOpbIX ITepeHOC 3JieKTpoHOB COITPOBO)!()J,aeTC51 OTHOCHTeJihHO 60JibIIIHM CHM- )!(eHHeM CB060,!J,HOM 3HepmH (p11c. 6-11). 3TH 3TaIThI cnoco6HbI 06ecneq11Tb 3Hepr11e.H CHHTe3

A T<l>, TaK KaK KOJI11qecrno BhI,!J,eJimome.Hc>I cBo-

60,!J,HOM 3Hepr11M ITpM6JIM3MTeJihHO paBHO 3HeprHH, Heo6xo,!J,HMOM M>I CHHTe3a AT<l> 113 NJ,<l> 11 cpoc-

cparn. 3KcnepHMeHTaJibHO 6hIJIO ITO,!J,TBep)!()J,eHO,

qTo npouecc nepeHoca 3JieKTpOHOB no UI13 11 CHHTe3 AT<l> 3HepreTmecKH coITp5I)!(eHbI.

IlepBbIM ITpouecc - ITepeHOC 3JieKTpOHOB OT BOCCTaHOBJieHHhIX KocpepMemoB NADH 11 FADH2

qepe3 UI13 Ha KHCJiopo,!1,

-

3K3eproH11qecKMM.

HaITp11Mep:

 

 

 

 

NADH + H+ + 1/2 0 2

~ NAD+ + Hp +

(I)

52 KKaJI/MOJib(""220

K,[()!(/MOJib).

BTOpo.H ITpouecc -

cpoc<J>opMJittpoBaH11e A,I1.<l>,

HJIH c11HTe3 AT<l>, -

3H,!1,eproH11qecK11i1:

 

A,l],<l> + H3P04+7,3 KKaJI/MOJib (30,5 K)],)!(/MOJib) =

AT<l> +Hp.

(2)

CMHTe3 AT<l> M3 Nl<l> M H 3P04 3a cqeT 3Hepr1111 ITepeHoca 3JieKTpOHOB no UI13 Ha3hIBaIOT OKHCJIHTeJihHbIM <t>oc<J>opMJI11poBaH11eM.

A. MEXAHM3M COIIPIDKEHMH OKMCJIEHMH

M ~OC~OPRJIMPOBAHMH

KaKHM )!(e o6pa3oM ocymecrnm1eTc>I conp>I)!(eHHe 3THX ,!1,Byx npoueccoB? Ha1160Jiee o6ocHoBaHHhIM OTBeT Ha 3TOT BOnpoc )l,aeT xeMHOCMOTmeCKa>I TeoP1151 MHT'IeJIJia,npe)l,JIO)!(eHHa>I HM B 1961 r. OcHoBHhie noJIO)!(eH1151 6bIJIH IIO,!J,TBep)!()J,eHbl H pa3pa60TaHbl )l,eTaJibHO COBMeCTHbIMH YCHJIH5IMH MHOrHX HCCJie,!J,OBaTeJietJ B nocJie,!J,yIOIUHe fO)l,bl.

1. llpomOHHblU zpaiJuenm u 3.lleKmpOXUMUlleCKUU nomenu,ull.!l

IlepeHOC 3JieKTpOHOB no )l,b!XaTeJibHOM uerr11

OT NADH K KHCJiopo,!1,y conpoBo)!()J,aeTcsi BhIKaqMBaHHeM npOTOHOB 113 MaTpHKCa MHTOXOH)J,pHM qepe3 BHYTPeHHIOIO MeM6paHy B Me)!(MeM6paHHoe ITpocTpaHcrno. Ha 3TY pa6ow 3aTpaq11BaeTcsi qaCTb 3Hepr1111 3JieKTPOHOB, nepeHOCHMbIX ITO UI13.

IlpoTOHbl, nepeHeceHHbie 113 MaTpHKCa B Me)!(- MeM6paHHOe npocTpaHCTBO, He MOfYT BePHYTbCH o6paTHO B MaTpHKC, TaK KaK BHyTpeHHHH MeM6paHa HeITpoH11uaeMa )J,JI51 npoTOHOB. TaKHM

o6pa30M' C03)J,aeTC51 ITPOTOHHbIM rpa)J,MeHT' ITPH KOTOPOM KOHU,eHTpaumI ITpOTOHOB B Me)!(MeM6paHHOM npocTpaHCTBe 60JihIIIe, a pH MeHhWe, qeM B MaTp11Kce. KpoMe Toro, Ka)!()J,hIH npoTOH Hecfa ITOJIO)!(HTeJibHbIH 3ap5I)J,, 11 BCJle)J,CTBHe 3TOro IIOHBJI>IeTCH pa3HOCTb ITOTeHUHaJIOB no o6e

CTOpOHbl MeM6paHbI: OTpHuaTeJihHbIH 3apH)l

Ha BHyTpeHHeH CTOPOHe 11 noJIO)!(HTeJlhHblM -

Ha

BHeIIIHe'ti. B COBOKYIIHOCTH 3JieKTpMqecKHH

M

KOHI.J,eHTpau110HHbIM rpa,!1,HeHTbl COCTaBJIHIOT

3JieKTPOXl1MHqecKHM ITOTeHU,HaJI .:1µH+ - MCToq-

HHK 3HepmM M>I CMHTe3a AT<l>. TaK KaK HaM6oJiee aKTHBHhlM TpattcnopT ITPOTOHOB B Me)J(- MeM6paHHOe npocTpaHCTBO, He06XO)l,11MbIH )l,JIH o6pa3oBamrn .:1µH+, npo11cxo,!J,HT Ha y•-rncTKax UI13, cooTBeTcrny10m11x pacIToJIO)J(eH1110 KOMnJieKcoB I, III 11 IV, 3TH yqacTKH Ha3hIBaIOT ITYHKTaMH conpH)J(eHl151 )lbIXaHH51 H cpoccpopHJIHpOBaHMH (pMC. 6-11, 6-13).

MexaHH3M TpaHcIToprn npoTOHOB qepe3 MHTOXOH,!1,pHaJihHYIO MeM6paHy B nyHKTax COITP5I)J(e- Hl151 He)J,OCTaToqHo 51CeH. 0,!J,HaKO ycTaHOBJieHO, qTo Ba)J(HYIO poJib B 3TOM ITpouecce HrpaeT KoQ. Ha1160Jiee ,!J,eTaJihHO MexaHH3M nepeHoca npo10HOB np11 yqacT1111 KoQ 113yqeH Ha ypoBHe KOMITJieKca III (p11c. 6-14).

KoQ nepeHOCMT 3JieKTPOHhI OT KOMTIJieKca I K KOMITJieKcy III 11 npoTOHhI 113 MaTpHKca B Me)J(- MeM6paHHOe npocTpaHCTBO, COBepIIIa>I CBoeo6pa3Hble u,11KJI11qecK11e npeBpameHHH, Ha3hrnaeMhie Q- UMKJiaMM. ,l],OHOpOM 3JieKTpOHOB )],Jl51 KOMnJieKca

Ill CJIY)J(HT BOCCTaHOBJieHHbIH y6MXMHOH (QH2), a aKuenTopoM - u11ToxpoM c. U11ToxpoM c ttaxo- ,!l,HTCH c BHenrne.M CTopoHhI BHYTPeHHetJ MeM6paHhI MHTOXOH,!1,ptttJ; TaM )J(e pacnoJiaraeTCH aKTHBHblll ueHTP UMTOXPOMa cl'c KOToporo 3JieKTPOHbl nepeHOC>ITCH Ha UHTOXPOM c.

276

 

 

 

 

 

HAPY>KHAfl MEME>PAHA MV1TOXOH.QPV1V1

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

Me>KMeM6paHHoe npocrpaHcreo

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

:---------------i--------------i-----------,

 

 

 

 

 

 

 

I

 

 

I

 

I

:

 

 

 

 

 

 

 

nw

 

 

nw

 

nw

:

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

+

 

 

+

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

2e-

 

 

 

 

 

 

 

 

 

I

 

 

BHyrpeHHsu1

 

 

 

 

 

 

 

IV

 

 

I

 

 

 

 

I

 

 

Ill

 

 

 

 

I

 

 

MeM6paHa

 

 

 

 

 

 

a a,

 

 

I

 

 

 

 

FMN

 

 

be,

 

 

 

 

I

 

 

 

 

 

 

 

 

Cu..

 

 

 

 

MVITOXOH,Qpl!lH

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

NADH

 

 

NAO•

 

<l>yMapar

1/20,

H,O

 

I

 

 

 

 

 

 

 

 

I

 

 

>-<

CyKLIVl-

 

 

 

 

 

I

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

I

 

 

0 Kcanoa1.1eraT ManaT

H8T

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

A.Q<l>+H.PO.

 

 

 

 

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

\'>-<r:====:i

 

I

 

 

 

 

 

 

 

 

 

 

 

 

 

t- - l

 

 

 

 

 

 

 

 

 

MaTpVIKc

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

AT<l>

 

 

v

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

PMc. 6-13. ConpRll<eHMe AblXaHMSI M CMHTesa ATCI>

e MMTOXOHAPMSIX. I -

NADH-i:ier111i:iporeHa3a; II -

cyKL\lllHaT i:ier111i:iporeHa3a;

Ill - QH,-i:ier111i:iporeHa3a; IV -

L\lllTOxpoMOKCVIAa3a;

V - AT<i>-CVIHTa3a. 8Hepr111S1 npOTOHHOro noTeHL\V1ana (aneKTpOXVIMVl'-tecKoro

nOT9HL\Vlana L'lµH+)VICnOnb3Y9TCSl AflSl CVIHT93a ATC!>, ecnVI npoTOHbl B03BpaL11at0TCSl B MaTpVIKC Yepe3 VIOHHble KaHanbl AT<i>-CVIHTa3bl.

MarpviKc

FesL c,

KoMnneKc I KoMnneKc Ill KoMnneKc IV

PMc. 6-14. ConpsiJKeHMe nepeHoca aneKTpOHOB ..epes Ab1XaTenbHbli1 KOMnneKc Ill c rpaHcnopTOM H• ..epes MeM6paHy. BoccraHoaneHHbli1 y6VIXVIHOH (QH2) B3aVIMOAei1CTByeT c Fe3+ reMa b, VI, BOCCTaHaBnVIBaSl ero, ocao6o:>K,QaeT npOTOH B BOAHYIO cjJa3y, npespaL11allCb B C9MVIXVIHOH (HQ.). 8neKTpOH OT reMa b, nepeHOCVITCSl Ha Fe3+ reMa b2. Ha•oTi:iaeT BTOpoi1 aneKTpOH Ha FeS-LleHTp, pacnono- )K9HHbli1 6nVl)K9 K Hapy)KHOi'InosepXHOCTVI MeM6paHbl; npVI 3TOM BTopoi'InpOTOH OK83blB88TCSl B M8)KM8M6paHHOM npocTpaHCTBe; aneKTPOH nepei:iaeTCSl Ha L\VITOXpOM c,. ai:ianee Ha L\VITOXPOM c. 0KVICneHHbli1 Q AVlcjJcjJyHAVIPYST K BHyrpeHHei1 CTOpOHe M8M6paHbl, ri:ie nonyYaeT aneKTpoH OT reMa b2 VI npoTOH Vl3 MaTpVIKca, npeapaL11aSlCb a Ha•. Ha•-nonyYaeT aneKTpOH OT KOMnneKca I 111 npoTOH Vl3 MaTpVIKCa; B MeM6paHe o6pa3yeTCSl QH2, VI Beeb npOL\9CC nOBTOpS18TCSl CHaYana.

B MeM6patte cymecrnyeT crnu11ottapttn1H. 06m11H. <l>oH,ll Q/QH2 , H3 KOTOporo Ka)[(,ll;a5I MOJieKyJia QH2 B O,ll,HOM u11KJie 06ecrreq11BaeT rrepeHOC rrpoTOHOB 113 MaTp11KCa B Me)l(MeM6paHHOe rrpocTpaHCTBO 11 3JieKTpOHOB, KOTOpbie B KOHelJHOM 11TOre ITOCTyITaIOT Ha K11CJ10po.n.. Ha pa6oTy'

COBepurneMyIO rrp11 BhlKaq11BaH1111 rrpOTOHOB, pacXO,IJ,yeTC5I qacTb CB060,IJ,HOH 3HeprHl1, KOTopa5I OCB060)[(,ll;aeTC5I rrptt rrepettoce 3JleKTPOHOB ITO rpa,IJ,HeHTY pe,IJ,OKC-'IlOTeHII,HaJia. 3tteprm1 3JieKTpOXHMHqecKoro IIOTeHUHaJia (,1µH+) 11CITOJ1b3YeTC51 ,ll,Jl51 cttHTe3a AT<l>, ecn11. npoTOHhI B03Bpa-

277

rn;aIOTCH B MaTpHKC qepe3 ttOHHhie Kamurhr A T<l>- CHHTa3hr.

2. Cmpoe11ue AT<l>-cu11ma3bt u cu11me3 AT</>

AT<l>-rnHTa3a (H+-AT<l>-a3a) - ttHTerpanhtthIH

6enoK BttYTpetttteti MeM6paHhI MHToxott,n:pttti.. Ott

pacnoJTO)[(ett B ttenocpe,n:CTBeHttOH 6JTH30CTH K

A

ti:hIXaTenhHOti. uentt. AT<l>-cttHTa3a cocTOHT 113 2 6eJTKOBhIX KOMrmeKcoB, o6o3HaqaeMbIX KaK F0 11

F1 (pwc. 6-15).

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279

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280

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H20

 

PMC. 6-19. KBTa6onM3M OCHOBHblX nMtqeBblX eeiqecTB. 1-3 -

n11lljeBapeH11e; 4-8 - cne411¢11'-1eCK11enyr11 Kara6onl13Ma; 9-10 - 3aKnl0-

411TenbHblH (06L411H nyrb) KaTa6onl13Ma; 11 - Un3; 12 -

OKl1Cnl1TenbHQe ctiocctiop11n11posaH11e.

Ilpesparn:etttte nttpysarn B auenm-KoA npowcxonttT rrptt yqaCTHH Ha6opa ct>epMeHTOB, CTpyKTYPHO 06nen1-rneHHh1x B n:upyBaTnerttnporetta3Hhiit KOMilJieKC (II)l,K). AueTMJlhHhIH OCTaTOK - aueTHJI-KoA naJiee OKHCJUieTC51 B UMKJie JIMMOH-

HOH KHCJIOThI no co2 H H20. B 3THX peaKuM51X OKMCJieHH51 npHHHMaIOT y<racTHe NAD- M FAD-3aBHCHMhie nerIInporetta3hI, noCTaBJI51IOrn:tte 3JieKTpOHhI w rrpoTOHhI s UI13, no KOTOpott OHM nepenaIOTC51 Ha 0 2

282

A. OKIICJUITEJihHOE ll,EKAPBOKCHJIIIPOBAHHE

IUfPYBATA

OK11cmrreJTuHoe JieKap60KcMJI11posam1e n11pysarn rrpo11cxOJIHT B MaTpHKce MHTOXOHJip11i1. TpattcnopT rmpyBaTa B MHTOXOHJIPI-taJibHbIM MaTpHKC qepe3 BttyTpeHHIOIO MeM6patty MHTOXOHJIPHH ocyru:ecTBJUieTC51 npH yqacTHH cneu11aJibttoro 6eJIKa-nepettoc- 'IHKano MexaHmM)'CHMrroprn c H+ (pttc. 6-20).

OpeBparn:em1e n11pyBaTa B aueTHJI-KoA orrHChIBaIOT CJie)lylOW:HM cyMMapHbIM ypaBHeHHeM:

CH3-CO-COOH +NAO++ HSKoA ~ CH3-CO -SKoA + NADH + H+ + co2'

B xo)le 3TOH peaKU1m rrpo11cxo,LI,11T OKHCJIHTeJihHoe )leKap60KCHJIHpOBaHHe nHpysaTa, B pe3yJibTaTe KoToporo Kap6oKCMJihHa51 rpyrrrra YJiaJI51eTC51 B BH,LI,e C02 , a aueTHJihHa51 rpyrrrra BKJIIO•rneTrn B COCTaB aueTHJI-KoA. 0JIHH aTOM BOJIOPOJia OKa3bIBaeTC51 B cocrnBe NADH, a Jipyroi1 B BHJie H+ nocryrraeT B cpeJiy. PeaKUH51 Heo6paTHMa, rrocKOJih- KY i1G0'=-33,5 K,[1,)1(/MOJib.

B rr11pysaTJierm:1,poretta3Hhii1 KOMrrJieKc (Il,Ll.K) BXOJI51T 3 cpepMeHTa: n11pysaTJieKap60Kc1rna3a (E,), JIHftt:JIPOJIHITOHJJTpattcaueTHJia3a (E2) H JIHfHJIPOJIHTIOHJI)lerm1,poreHa3a (E3), a TaK)Ke 5 KO- ¢epMeHTOB: THaMHHil,HcpoccpaT (T,Ll.<D), JIHIIOeBa51 KHcnoTa, FAD, NAO+ 11 KoA. KpoMe Toro, B COCTaB KOMITJieKca BXOJJ,51T peryn51TOpHbie cy6be- ,Ll,HH HUbI: npOTeHHKHHa3a H cpocct>orrpoTeHHcPOC- ¢aTa3a (Ta6JI. 6-5).

Mell<MeM6paHHOe npocTpaHCTBO

CH,-c-coo-

11

0

BttyrpeHHflfl

MeM6paHa

Mil!TOXOHAPIAlil

cH.-c ·-coo-

11

1. CrpoeHHe mq>yeaTJieill.ll,poreuaJuoro KOMllJieKca

 

 

0

Opouecc OKHCJIHTenbttoro

JieKap60Kc11n11po-

 

 

MaTp111Kc

BaH115l rrnpyBaTa KaTaJIH3HpyeT

CJIO)l(HOOpraHH-

PMC. 6-20. TpaHcnopT nMpyeaTa 'lepe3MMTOXOHApManbHYIO

30BaHHbIM m1pyBaT)J,efHJiporeHa3HbIM KOMIIJieKC.

MeM6paHy.

 

OH

 

 

 

 

NAD•

 

 

 

 

 

 

 

 

I

 

 

 

 

 

 

HC-CH.

 

 

 

 

E.FADH,\V

co, xdA<> v-EJlK:::::!

\NADH+H•

 

 

 

 

 

 

 

 

I

A:~

Ill

 

COOH

E,TA<l>

E,nK

 

I

 

/\

 

 

 

 

 

 

 

 

 

C=O

0 = C-S

SH

 

 

 

I

 

6H.

 

 

 

 

 

CH.

 

 

 

 

 

 

 

HS-KoA

PMc. 6-21. nocn~oeaTenbHOCTb peaK4111i'i,KaTan111s111pyeMblX nJlK. I - E1 KaTan111a111pyer 11eKap60Kc111n111posaH111e m1pysara 111 nepeHoc c2cpparMeHra Ha TJJ,<!l; II - E2 Karan111a111pyeT oK111cneH111e r111i::ipoKCll13Tll1nbHOi1 rpynnbl 111 nepeHoc C,-cpparMeHra Ha n111noesyt0 Kll1Cnory (nK); Ill - a4eT111n111posaHHa5l J1111r111APOOll100111nTpaHca4ernnaaa B3all1MOJ:iei1crsyer c KoA c o6pa30BaHll1eM BOCCTaHosneHHOH cpopMbl n111noesoi1 KMcnoTbl 111 a4ernn-KoA; IV - oK111cneHHa5l cpopMa TpaHca4ernnaab1 pereHep111pyercR np111 y1.1acr111111 E3; V - oK111cneHHafl cpopMa E3 pereHep111pyerc5l np111 y1.1acrn111 NAO+.

283

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