підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

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3. MEHRJIATU:MKJIA3HAH CMCTEMA

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rauHqecK11i1 cpoc¢aT (P.) OT)lem1eTCH OT a- cy6beJlHHHUhI, a KOMITJie~c [a-f)J,<I>] coxpaHH-

eTcH; CKOpOCTb rHJlPOJIH3a OITpe)l,eJIHeT BpeMH npoBe,ll,eHHH cwruana;

•o6pa30Bauwe B aKTl1BHOM ueHTpe a-cy6be)ll1- Hl1Ubl MOJieKYJihI f)J,<I> CHH)K:aeT ero CPOJlCTBO K a).l,eHMnaTUHKJia3e, HO yBeJil1'lHBaeTcpOJlCTBO K py-cy6beAHHHUaM. G,-6enoK B03BpamaeTCH K HeaKTl1BHOH ¢opMe;

•ecJIH peuernop CBH3aH c aKTHBaTopoM, HaITpHMep ropMOHOM, UHKJI <i>YHKUHOHttpoBamrn G, 6eJIKa IIOBTOpHeTCH.

AlcrHBallHH npoTeUHKHHa3hI A (TIKA)

•MoJieKYJihI uAM<I> Moryr 06paT11Mo coeJJMHHThcH c peryAATOpHbIMH cy6beJlttH11uaMH IIKA.

•IlpHCOe)l,HHeHHe uAM<I> K peryJIHTOPHbIM cy6be)l,11HHUaM (R) Bbl3bIBaeT JlHCCOU:HaUHJO KOM-

ITJieKCa C2~ Ha KOMTIJieKC uAM<l>4 ~MC + C.

• AKTHBHaH ITpOTel1HK11Ha3a A ¢oc¢opHJT11pyeT cITeUHWM'leCK11e6eJIKM ITO cepwuy 11 TpeOHHHY, B pe3ynhTaTe H3MeHHIOTCH KottcpopMaum.1. H aKTMBHOCTb ¢oc¢opHJTHPOBaHHhIX 6eJIKOB, a 3TO npHBO,[J,11T K H3MeHeHl1IO CKOpOCTM 11 HarrpaBJieHMH peryJittpyeMbIX HMH ITpoueccoB B KJJeTKe.

• KouueHTpauHH uAM<I> B KJ1eTKe MO)K:eT pefYJIHPOBaTbCH, Otta 3aBl1CHT OT COOTHOWeHl1H aKTl1BHOCTeii ¢epMeHTOB a).l,eH11JiaTIJ,MKJia3bl 11 ¢oc- ¢oJll13CTepa3hl.

oOJibWYIO pOJib B peryJIHUllll BHYTPl1KJieToqHOH c11ruanbHOM CHCTeMhI ttrpaeT 6eJioK AKAP . <<3m:1KopetttthIM» 6eJioK AKAPs yqacrnyeT B c6opK~ ¢epMeHTHbIX KOMITJieKCOB, BKJIJOqaJOIUl1X He TOJib-

KO npOTeHHKHHa3y A, HO 11 ¢oc¢0.u113cTepa3y 11

¢oc¢onp0Te11H¢oc¢arn3y.

KacKa,ruthtu Mexauu3M ycHJieHJUI n noL1,aBJ1eum1 cnrHaJia. nepe,UatJa Cl1rHaJia OT MCM6paHHOfO peue11TOpa Yepe3 G-6e110K Ha ¢epMeHT an.eH1111aTu11KJia3y CJ1j')Kl1Tnp11MepoM KaCKMHOM Cl1CTeMbl yc1111eH!15! 3Toro c11rHaJia. OuHa MOJieKy11a, aKTHB11py10iuaH peuenTop, MO)KeT «BKJIIOYaTh» HecKOJlhKO G-6en- KOB, 11 3aTeM Ka>KJib/M aKTl1Bl1pyeT HeCKOJlbKO MO-·

JleKyJI a.ueHHJiaTU!1KJia3bl c o6pa30BaHl1CM TblC51lJ MOJICKYJI uAM<P. Ha 3TOM 3Tane c11rna.JI ycl1J111BaeTcs1 B 102-103 pa3. 06pa3y10w11i1c51 uAM cD «BKJIIOtJaIOT» .upyroi1 ¢epMeHT - npoTeHHKHHa3y A, yc11mrna51 Cl1rHa'lewe B 1000 pa3. <PoccflOpl1JJ11pOBaHl1e ¢epMeHTOB npoTeHHKHHaJoti A eiue 6011bwe yc111111BaeT cJ..ffHari, B pe3yJihTaTe cyMMapHoe ycl1JleH11e paRHO 106-107 pa3. TaKHM o6pa30M, no MexaHH3MY KacK<lllHOro yc1111eH11H o.uHa MOJieKYJJa perym1rnpa

cnoco6Ha H3MeHl1Th aKTl1BHOCTh MHJIJIHOHOB LlPYfHX MOJieKYJI.

Ho MH mo6oti H3 cHcTeM TpattcMeM6pattttoti nepe,ll,al!l1 CHfHaJia KJieTKa HMeeT LlPYryIO Cl1CTeMy, no,ll,aBJIHIOJUYIO 3TOT c11rnan. K(l)K)J,hIH 113 3TaTIOB B <PepMeHTHOM KaCKa)l.e HaXO)l,11TCH no)], KOHTPOJleM cneIJ,11aJihHhIX TIO)l.aBJIHIOIIl,HX 3TOT Cl1fHaJI MexaHH3MOB. Hanp11Mep, )].JIHTeJihHoe )l.eticrn11e ropMoHa npHBO)l.HT K )l,eCeHCl16llJUf3aIJ,Hl1 MeM6paHHhIX peuenTopoB: OHll Jil160 ttHaKTHBttpyIOTCH, Jlll60 BMeCTe c ropMOHOM norp)')KaIOTCH B KJieTKY nocpe,ll,CTBOM 3H,ll,OUMT03a. B pe3yJihTaTe )l.ecettc11611n113au,u11 peuenrnpoB cTenetth aKTttBau1rn a)l.eHttJiaTUMKJia3HOH CHCTeMhl CHIDKaeTCH. Ecm1 B KJieTKe )l,Jll1TeJibHOe BpeMH TIOBhIIIIeHa KOHIJ,eHTpaU,ttH uAM<I> (nOBhIWeHa aKTllBHOCTh npoTellHKllHa3hI A), MO)KeT npoucxo)l.ttTh <Poc<PopllJlttpoBaHtte KaJihIJ,tteBhIX KaHaJIOB, qTO npHBO,ll,llT K IIOBhIIIIeHl1IO KOHu,ettTpau11u Ca2+ B KJieTKe. Kanhu,11ti aKTttBttpyeT Ca2+-3aBttCl1MYIO <Poc<Po)l.113cTepa3y, KaTM113npy10my10 rrpeBpamettne uAM<I> B AM <I>.

B pe3yJihTaTe ttHaKTMBau,u11 rrpoTel1HKMHa3hI A (R2C2) CHtt)KaeTCH CKOPOCTh <Poc<PopllJlttpOBaHl1H crreun<P11qecKHX <PepMeHTOB. 3aBepIIIaeT «BhIKJIIOlfeHtte» cttcTeMhI <Poc<PonpOTentt<Poc<Parn3a, ,ll,e- <Poc<PopllJlttPYIOmaH <Poc<PonpoTettHhI.

BJmsrnue 6aKTepuaJibHblX TOKCHHOB ua aKTHBuocTb a,l],eHHJiaTIJ,HK.JUl3bl (A,Il;<l>-pu6o3HRHpOBa- HHe G-6eRKOB)

Ll,JIH 113yqettnH cPYHKUl10H11poBatt11H G-6eJIKOB a,n:eHHJiaTIJ,llKJla3HOH CllCTeMbI 6bmll llCTIOJlh30BaHhI 3K30reHHhie 6aKTep11aJihHhie H)l.hl - XOJiepHhIH H KOKJilOIIIHhIB TOKCllHhl. ToKCllHhl B 3KCrrepllMeHTaJihHhIX ycnoBHHX noBhIIIIa10T aKTttBHOCTh a,n:eHHJlaTUHKJia3hI IlpaKTll'-leCKllBO Bcex KJieTKax opraHl13Ma; TaK, XOJiepHhIH TOKCHH MO)KeT CTUMYJlllpoBaTb ceKpeu,1110 Tttpeott)l,HhIX ropMOHOB KJieTKaMll IUl1TOBH)],HOH )KeJle3hl, CTepol1)].HhJX ropMOHOB KJieTKaMH Ha)l.noqeqHllKOB, pacrra)l. )KttpoB B )K}IPOBhIX KJieTKax. PeaKUHH pa3HbIX KJieTOK Ha XOJiepHblli TOKCllH BhI3BaHa nOBhIIIIeHtteM ypoBHH uAM<I> B 3THX KJTeTKax.

XoJiepHhIH TOKCl1H - on11roMepHhIH 6enoK. 0)].Ha 113 cy6be)l.11H11U - <PepMeHT NJ,<1>-pll603llJITPaHC- <Pepa3a; npOHl1KaH B KJieTKy, OHa KaTaJil1311pyeT rrp11COe)l,l1HeH11e ALl,<l>-p11603hl K a,-cy6oe)l.llHl1IJ,e KOMilJleKca [a,-IT<l>][AUJ (3Tan aKT11BaIJ,ttl1 a)l.eHttJiaTIJ,llKJia3hl).

NAO++ [a s-IT<l>][AU] ~ [ALl,<l>-p11603l1Jl-a,- IT<l>][AU] + HllKOT11HaMl1)]. + H+.

NJ,<l>-p11603l1JlttpoBaH11e ttttrn6ttpyeT rrpoHBJieHtte IT<l>-<Poc<Parn3HOH aKT11BHOCTtt a,-cy6'he)l.tt- H11UhI, He npo11CXO)l,HT )l,e<Poc<PopttJil1POBaHtte IT<I>. UnKJI cPYHKUHOH11poBaH11H G,-6eJIKa ocrnHaBJinBaeTcH Ha 3Talle aKTHBaIJ,1111 <PepMeHTa a)l.e- Hl1JlaTIJ,llKJla3hI, OTBetiaIOmero 3a o6pa3oBaHtte uAM <I> tt3 AT<I>. <l>epMeHT a,ll,eH11naTUHKJia3a coxpattHeT nOBhIIIIeHHYIO aKTllBHOCTh B TelJeHtte rurnTeJihHOfO BpeMeH11.

Cy6'he)l.11Httu,a KOKJilOIIIHoro TOKC11Ha, npoH11KaH B KJieTKY, KaTaJil1311pyeT ALl,<l>-p1160311Jil1PO- BaH11e CX;-Cy6'he)l.11HttUhIaKT11BHpoBaHHoro G;-6enKa (a;py-fT<I>).

NAD++ [a;py-IT<I>] ~ [ALl,<l>-p11603ttJI- a;PYIT<I>] + Hl1KOTttHaMl1)]. + H+.

M o)l.11<P11uupoBaHHaH a;-cy6'he)l.11H11uacoxpaHH- eT BhICOKOe cpO,ll,CTBO K py-cy6'he)l,HHHUaM, T.e. G;-6eJioK TepHeT crroco6HocTh )l.11ccou1111poBaTh Ha a;-IT<I> 11 py-cy6'he)l.llHttUhI.TaKttM o6pa3oM, HHrn6ttpy10m11ti Cl1fHaJI ( ai-IT<l>) He )l.OCT11raeT a,l],e- HHJlaTIJ,llKJia3hI, 3HalJl1T B 3TOM CJiyqae B03MO)!(Ha TOJlhKO ee aKTllBaIJ,ttH rrp11 CBH3hIBaHl111 c a,-IT<I>. Ll,ettCTBtte KOKJllOWHOfO TOKCttHa Ha KJieTKH TKaHeH Bcema np11BO)l,11T K IIOBhIWeHl110 ypoBHH uAM<l>.

CttMIITOMhI xonephI 11 KOKJIIOwa pa3B11Ba10TcH B pe3yJlhTaTe ,l],eHCTBl15l TOKCl1HOB, Bhipa6aThJBaeMhIX COOTBeTCTBYIOI11,11Ml1 Ml1KpoopraHl13MaMl1.

If. lfH03HTOJI<I>OC<I>ATHASI CHCTEMA

<l>yHKUHOHHPOB<lHtte HH03HTOJlcPOCcPaTHOH CHCTeMhI TpaHCMeM6paHHOH nepe,ll,alJl1 Cl1fHaJia (p11c.

5-42) o6ecnel!HMIOT: R (peuemop), <Poc<Ponttna3a C, Gplc - 6eJIOK, aKT11BttpYIQIIl,11H <Poc<PoJIHTia3Y C, 6eJIKll 11 <PepMeHThI MeM6paH 11 UHT030J1H.

IloCRe)l.OBaTMbHOCTb co6blTHH, DpHBO,ll,HUUIX K aKTHBaIJ,HH <1>oc<1>0Runa1h1 C:

• CBH3bIBaHHe Cl1fHaJihHOH MOJieKYJihI, Hanp11-

Mep ropMOHa c peuenTOpOM (R), Bhl3hIBaeT

l13MetteH11e KOHcPOPMaIJ,1111 H yBem1lJeHtte cpo,ll,-

CTBa K Gp1c-6eJIKY.

•o6pa30Batt11e KOMnJieKca [f][R][Gp1c-fLl,<l>] rrp11BO)l,HT K CHIDKeHl1IO cpO,ll,CTBa a-npOTOMe-

pa Gplc-6eJIKa K fLl,<l> tt YBeJil1lJett1110 cponcrna K IT<l>. f.ll,<I> 3aMem1eTCH Ha IT<l>.

•3TO Bhl3bIBaeT )l,11CCOIJ,11aIJ,1110 KOMilJleKca; OT- )l,eJ111BlllaHCH a-cy6'be)].l1H11Ua,CBH3aHHaH c MO:-

JieKYJIOH IT<I>, nptto6perneT cpo,ll,cTBo K <Poc- <Ponttna3e C.

c~ca

C~a

KanbM(),£1YTllAH 4Ca2+

-

PMC. 5-42. lllH03MTon<1>oc<1>aTH8SI CMCTeMa.

B3al1MO)J.eilcTByeT c <Poc<PommaJoi1: C 11 aKTIIBl1pyer ee. no)]. )J.eHCTBl1eM ¢oc<}lomma- 3bI-C rrpol1CXO)J.HT fH)lpom13 .JIID1l1,Ila MeM6paHbI <Poc<Parn.ru1mrno311TOJI-4,5-611c¢oc¢aTa (<l>M<l>2).

• B XO)J.e f11)J.pOJil13a 06pa3yeTC5I 11 Bb!XO)J.l1T B Ul1T030Jlb fH)J.POcPHJibHOe Bemecrno l1H0311TOJI- 1,4,5-Tp11<}loc¢aT (M<l>). )1,pyroH: rrpo)J.)'KTpeaKu1111 )J.11aUl1JifJil1UepoJI ()1,Af) OCTaeTC51 B MeM6patte 11 yqacrnyeT B aKT11Bau1111 <PepMeH-

Ta rrpoTel1HKHHa3hI c

• l1H0311TOJI-1,4,5-Tpl1<}loc<}laT (H<1>3) CB513hIBaeTC51 cneuJI¢wrecKHMH uempaMJI Cai+-Katta-

4>oc¢on111naaa C

AKTWBHblH cpepMeHT

Jia MeM6paHbl 3P, 3TO npJIBO)J.JIT K l13MeHeHl1IO KOH<}lOpMaUJIJI 6eJIKa JI OTKpbITJIIO KaHaJia - Ca2+ nocTynaeT B UJIT030Jih. B oTcyrCTBJie B UHT030Jie 11<1>3 KaHaJI 3aKpbIT.

AKTuBal.UUI npoTeHHirnHa3hl C

• noBhillleHJie KOHileHTpauJIJI Ca2+ B Ul1T030Jie KJieTKH YBeJIWI11BaeT CKopoCTh B3aHMO)J.ettcTBmr Ca2+ c tteaKTJIBHhIM UJIT030JihHhIM <PepMeHTOM npoTel1HKHHa30H c (IlKC) JI 6eJIKOM KaJibMOJl)'Jil1HOM, TaKHM o6pa30M CJifHaJI, rrpMH51TbIH peuerrTopoM KJieTKH, pa3,llBal1Baerc51.

•Cmt3hIBam1e npoTeHHKHHa3hI C c HOHaMH Kllilh-

11,IDI n03BOJUieT ¢epMeHTY BC'rynaTbB KaJlhll,HMonocpe.n:oBaHHOe B3aHMOL(eHCTBHe c MOJieK)'- JiaMII «KIICJIOro» ¢oc¢0JI1In1I.n:a MeM6paHhI, ¢oc¢aTttnHJicepHHa ( <l>C). Ll,1Ia11,HJirn1I11,epoJI, 3aHHMaH cneuttQ:>wrecKJie ueHTphI B npOTeHHKIIHa3e C, eme 6oJiee yileJilllIIIBaeT ee cpo.n:CTBO K HOHaM Kllilhll,IDI.

•Ha BHyrpeHHei1 crnpoHe MeM6paHhI o6pa3yeTCH

¢epMeHTaTHBHbIM KOMilJieKC - [ nKC] [Ca2+l

[Ll,Af][<l>C] - aKTIIBHaH npoTe1IHK1IHa3a C, ¢oc¢opHJIIIPYJOmaH crreu1I¢wrecK1Ie ¢epMeHThI ITO cepHHY H TpeOHHtty.

Y11acme 6eJIKa KaJibMO,rzyJIHHa B HH03HTOJ](l>occt>aTHoii nepe,l1,a11e cumaJia

B KJieTKax MHOrHX TKaHei1 rrplICYTCTByeT 6eJIOK KllilhMOnyJIHH, KOTOpbIM cPYHKll,HOHHpyeT KaK BttyTPIIKJieTOlJHhlH peuerrTop Ca2+, OH HMeeT 4 ueHTpa nJUI CB513hrnamrn Ca2+. KoMITJieKc [KllilhMOD:YJIIIH]- [4 Ca2+] He o6JianaeT ¢epMeHTaTIIBHoi1 aKTIIBHOCThIO, HO B3aHMOL(eHCTBHe KOMilJieKca c pa3JUi'l- HhIMII 6eJIKaMH II ¢epMeHTaMII rrplIBOL(HT K HX aKTHBaUHII.

CaMoperymm,us cucTeMbI

KaK H 6oJibWIIHCTBO CIICTeM TpaHcMeM6paHHoi1

rrepe.n:alJII CIIfH3JIOB, IIH031ITOJIQ:>oc¢aTHM CHCTeMa HMeeT He TOJibKO MexaHH3M ycHJieHH51, HO H MexaHH3M rro.n:aBJieHHJI CHfHllila. IlpHCYTCTBYJOill,He B UIIT030Jie IIH031ITOJI- l ,4,5-TplI¢oc¢aT (11<1>3) II L(1IaU11JifJIHUepoJI (Ll,Ar) B MeM6paHe MOfYT B pe3YJihTaTe cepHH peaKll,HM OIIJITh rrpeBpamaTbCJI B ¢oc¢aTtt.n:wmrno31ITOJI-4,5-6wc¢oc¢aT (<l>H<l>2).

<l>epMeHThI, KaTaJIII31Ipy10m1Ie BOCCTaHOBJieHIIe ¢oc¢oJIHTIH.LJ.a, aKTIIBHPYJOTCJI ¢oc¢op11JIIIPOBaHHeM npOTeHHKHHa30ll c.

KoHUeHTpauHJI Ca2+ B KJieTKe CHIDKaeTCJI .n:o HCxonttoro ypoBH51 nplI ,nei1cTB11H Ca2+-AT<l>-m UJITOTIJia3MaTH'lecKoi1MeM6paHb1H3P,a TaIOKe Na+/Ca2+ - H H+/CaH-TpaHCJIOKa3 (aKTHBHhIH aHTHIIOPT) KJie-

TOlJHOi1 II MHTOXOH,IlpHaIIbHOM MeM6paH.

<l>yHKIJHOH1IpoBaH1Ie TpattcJioKa3 CaH w Ca2+ -

AT<l>-a3 MO)!(eT aKTIIBHpoBaThcH:

• npOTeHHKHHa30M c (¢oc¢opHJIHPOBaHHeM). I10HH)!(ett11e KOHUeHTpau,HH Ca2+ B KJieTKe H .n:1IauHJirn1IuepoJia B MeM6paHe nplIBO.IllIT K II3MeHeHHIO KoH¢opMau11H rrpoTeHHKMHa3hI C, CHII)!(e-

HHIO ee CpOL(CTBa K Q:>ocQ:>aTHL(HJICepHHy, Q:>epMeHT ,IlHCCOll,HHpyeT B 11,HT030Jlh (HeaKTHBHM ¢opMa).

<l>oc¢opHJIHpOBaHHbie rrpOTeHHKHHa30H c ¢epMeHThl II 6eJIKM rro.n: .n:ei1cTB1IeM ¢oc¢orrpoTeHH- ¢oc¢arn3h1 rrepexo,nHT B .ne¢oc¢op11JIHpoBaHHy10 Q:>opMy.

K. IIEPE,lJ,AqA ClffHAJIA c IIOMOIIIJ>IO

BHYTPHK.JIETQqffhIX PEUEIITOPOB

nepe.n:aqa CHrHarta JIHTIIIL(OpaCTBOplIMbIX CTepo- 1-0lHhIX ropMOHOB H THPOKCHHa B03MO)Klia TOJihKO rrpH rrpoxo~eHHH 3THX ropMOHOB qepe3 ITJia3MaTH'leCK)'IOMeM6paHy KJieTOK-MHWeHeM (p11c. 5-43).

Peuemoph1 ropMOHOB MOfYT Haxo.n:HThCJI B UHT030Jie HJIH B Mpe. ll,HT030JihHhie peuemophr CB513aHhI c 6eJIKOM-warrepOHOM (qacTO 3TO rpyrrrra 6eJIKOBwarrepoHOB). 51.n:epHhie II UIIT030JibHhre peuemophr CTepOH.LJ.HhIX H THpeOIIL(HbIX ropMOHOB co.n:ep)!(aT Ll,HK-CBJI3hIBaIOI11,HM L(OMeH, xapaKTeplI3YIOI11,HMC51 HaJIH'IHeM.n:Byx crpyKzyp «UHHKOBbIX Illlilbll,eB».

IloCJie,lJ,OBaTeJILHOCTh co6LITHil, npuso)UlllUIX K aKTHBal.UIH TpaHCKpHOl.UIH:

•ropMOH rrpoxo.n:HT qepe3 L(BOMHOM JIHII.H.llHhlM CJIOM KJieTO'IHOMMeM6paHhI.

•B3aHMOL(eMCTBHe ropMOHa c peuenTOpOM (R)

IIPIIBO,Ll,IIT K H3MeHeHIIIO KOHQ:>opMaumt peuenTOpa w CHH)!(eHHIO cpo.ncrna K 6eJIKaM-warre- poHaM, OT,Ll,eJIJIIOill,HMCJI OT KOMITJieKca ropMoHpeuemop.

• KOMTIJieKC ropMOH-peuerrTOP npoxo.n:HT B MPO, B3aHMO,Ilei1crnyeT c peryJIJITOPHOM HYKJieOTH.LJ.- HOM IIOCJie.n:oBaTeJibHOCTbIO B Ll,HK - 3HXaHcepoM HJIH cai1JieHcepoM.

•yBeJUtlJIIBaeTC51 (rrpH B3alIMO,Ll,eMCTBHH c 3HxaHcepoM) IIJIH YMeHhwaeTCH (rrplI B3aHMo.n:etiCTBIIII c cai1JieHcepoM) ,Ll,OCTynHOCTh rrpOMOTOpa .LJ.JIJI PHK-rroJIHMepa3hI.

•cooTBeTCTBeHHO YBeJIIIlJHBaeTCJI HJIH yMeHhwaeTCH cKopocTh TpaHcKpHrruHII CTPYKTYPHhIX reHOB.

•yBeJIH'IHBaeTCJI IIJIH yMeHbwaeTCJI CKOpOCTh TpaHCJIJIUHH.

•II3MeHHeTC51 KOJIII'leCTBO 6eJIKOB, KOTOPhie MOfYT BJIHJITh Ha MeTa60JIH3M H cPYHKIJHOH3JlbHOe COCT051HIIe KJieTKII.

3¢¢eKThl ropMOHOB, KOTOpbie rrepe.n:a10T CIIfHaJI qepe3 BHYTPIIKJieTOlJHhie peuerrrnphI, HeJih3JI Ha6JI10.n:aTh cpa3y, TaK KaK Ha rrpoTeKaH1Ie MaTplI'IHhIXrrpoueccoB (TpaHCKplIIIUHIO II TpaHCJI51UHIO) Tpe6yioTCJI 'laCbl.

P111c. 5-43. nepeAa•ta CIArHana ~la BHyTpmmeT04Hble pe1..1enTOpbl.

JI. CrrEU,M(()lflfHOCTh CHrHAJUI3AUHH

Ll;rn !1CCJJe).lOUaTCJ1eti, 11MelOW,11X npe).lCTaBJJeH11e 0 KOJJMl.Jecrne C11fHaJJbllbIX MOJJeKyJI, 0 COOTBeTCTBYIOW,eM KOJ1111.Jecrne peuenrnpou, o TpaHcMeM6paHHhIX Cl1CTel\1aX nepe,lla1.J11 Cl1rHaJIOB, BTOpH'IHbJXno-

cpe,llH!1KaX, OCTaeTC51 33ral!KOH, K3K npOTeHHKl1Ha3bl Bb!6Hp3!0T COOTBCTCTBYIOlUH~i cpepMCHT MeTa60JIHlJeCKOfO nyT11 .IL151 cj:>occj:Jopvtm1ponaH11s:i. f!lccJieao-

BaTen11 !JJJ5l o6b51CHCHl151 3TOfO 51BJICHl151 npeJJ.naraIOT «rnnOTe3y M11weH11» (OT aHrn. targeting hypothesis).

no 3TOH rnnoTe3e CilCIJl1cP114HOCTb 11pOTel1HKl1Ha3 11 cj:Joccj:>onp0Te11Hcj:>occj:>aTa3 ,r1,ocrnranc51 nyreM 06-

pa30BaHl151 KOMnapnvteHTOB Ha MeM6paHe, u COCTaU KOTOpblX BXO,ll51T He TOJJbKO C3Ml1 npoTel1HKl1Ha3bl 11 cj:Jocqmnp0Te11Hcj:Joccj:>arn3h1, HO 11 cneu11cj:Jw1ecK11e 6eJIK11-cy6crpaTb1. Ha;111411e ocrnTKa M11p11cT11Houol1 11JIH naJ1bMMT11Houotf Kl1CJIOThJ n cTpyKType 6eJIKOBcy6cTpmon -- ycJIOBl1e 11X «3a51KOpi1B3Hl151» B COOTBCTCTBYIOW,CM MeM6paHHOM KOMnapTMCllTC.

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