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підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

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144

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145

proteins).
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(HMG-6emrn -

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f. CTPYKTYPA PMEOHYKJIEMHOBhlX KMCJJOT

(PHK)

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cytb E

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146

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KO)J,OHa, 06b1qHo 3TO TpmmeT 3a KO)J,11-

PYIOml1M yqacTKOM cJie)J.yeT O)J.HH 113 TepMHH11py10m11x KO)J.OHOB - -UGA-, -UUA-, -UAG-. Ha

3'-KOHUe 6oJihllll1HCTBa MPHK rrp11CYTCTByeT nocJie)J,oBaTeJihHOCTh HyKJieOTl1)],0B 113 100-200 a)J,e-

H0311HMOHO$occJ:>aTHbIX OCTaTKOB.

PuoocoMaJibHble PHK (pPHK)

P116ocoMaJihHhie PHK HMeIOT MHoroq11c11eHHbie crr11pan1130BaHHhie yqacTKH. Pa31IH'laJOTpPHK -

5S, 5,8S, 28S H 18S (S - Ko3cJ:>c):>uu11em ce)l11MeHrnu1111). Pn6ocoManhHhie PHK co)J.ep)f(aT ttecKoJih-

147

4-13).
4-12.

7-Men111ryaH111H

nces.i:1oypa1..1111n

PMC. 4-11. MMHOpHble OCHOBBHMR rPHK.

KO MO)IH<PHIIHPOBaHHhIX H)'K.JleOTil,!lOB,qam:e BCero 3TO MenvmpoBaHHhie rrpOH3BO)IHhie a30THCThlX OCHOBaHHH HJIH pH603hl (2'-MenrnpH6o3a). pPHK o6pa3yIOT KOMIIJieKChl c 6eJIKaMH, KOTOpbie Ha3hlBaIOT pH60COMaMH. KIDK,JJ,aSI pH60COMa COCTOHT H3 JJ;Byx Cy6be)IHHHU - MaJIOH (40S) H 60JihIIIOH (60S).

Cy6be)IHHHUhI pH6ocoM paJJIIfqaIOTCSI He TOJihKO Ha6opoM pPHK, HO H KOJIIfqecTBOM H CTPYKTYPOH 6eJIKOB (pHC. 4-12).

,ll;. fHBPH)J,H3AI.JJI.H HYK.JIEHHOBblX KHCJIOT

Brop11qHasi: CTPYKTYPa HYKJieHHOBhIX KHCJIOT 06pa3yeTcsi: 3a cqeT CJia6hIX B3aHMO)IeHCTBHH - B0- ,!1,0p0,!1,HhIX H rH,!1,p0<po6HhIX. Il03TOMY ecJIH BO)I- HhIH pacTBop .UHK HarpeTh )IO 100 °C, TO CBH3H, y;:i:ep:>1arna10m:He )IBe u:errH )IBOHHOH CIIHpaJIH BMeCTe, pa3pYIIIaIOTCSI. B pe3yJihTaTe pa3phrna BO,ll,O- po.11.HhIX H rn;:i:poq>06HhIX CBSI3ett uerrH .UHK pacXO,ll,SITCSI. 3'roTrrp011ecc Ha3hIBaIOT «)IeH8TYP8IUUI». 0.uHaKO eCJIH paCTBOp, co.uep)K:am:HH .ueHaTypHpOBaHHyIO .UHK, oqeHh Me)IJieHHO OXJia)K:,ll,aTh, TO MOryT rronyqttThCSI ;:i:ByxcrrnpaJihHhie CTPYKTYPhI, H)IeHTifqHhie HCXO)IHhIM. TaKOH rrpouecc IIOJiyqHJI Ha3BaHHe «peH8THB8IUUI».

Ha SIBJieHHH .11.ettazypaI.1,HH H pettaTHBall,HH ocHoBaH MeTO)J., Ha3hIBaeMhIH «MOJieicyJUIJHlaJI m6p~38-

IJ,HJI». Ilpouecc rn6pH,!1,HJmHH MO)K:eT ocym:eCTB.JUIThcsi: Me)K:.11.y .11.BYMSI JIJ06hIMH ueIISIMH H)'K.JleHHOBhlX KHCJIOT ()J;HK-.UHK, .UHK-PHK) rrpH yCJIOBHH, lfI'O

OHM co;:i:ep)K:aT KOMIIJieMemapHhie rrocne;:i:oBaTeJihHOCTH ttyKJieOTil,!lOB. TaKHe rn6plf)IHh1e CTPYK'fYPhI MO)[(HO Bhl)IeJIHTh ueHTPH<l>yrnpoBaHHeM B rpa,LI,H- eHTe IIJIOTHOCTH caxapo3hl HJIH Ha6JII0,!1,aTh B 3JieKTPOHHOM MHKpoCKOrre (pHC.

EcJIH pacTBop, co.uep)K:am:Hif o6pa3UhI .UHK I

H 2, BhI)IeJieHHhle H3 opraHH3MOB pa3Hh1X BH)IOB,

.ueHaTypHpOBaTh, a 3aTeM rrpoBeCTH peHaTHBaIIHIO, TO 06pa3y10Tcsi: .11.ByxcrrHpanhHhle cTpyKTYphI. Ho HapSl,ll,y c HCXO)J.HhIMH )],HK 1 H ,llHK 2

npoKaplAOTbl

 

705

/

~

Cy6be.Q111H1114b1

505 0 305

l

~58

pPHK ~235 + 34 6e11Ka

pPHK ~ 165 + 21 6enoK

3yKapll10Tbl

P1116ocoMa

808

 

/

~

 

Cy6be.Q111H1111..1b1

608 •

40$

1

~55

5,8$ 0.

pPHK ~285 + 50 6eJ1KOB

pPHK ~ 188 + 33 6enKa

P111c. CrpoeHMe 3yicap111or1111.1ecK111x 111 npoicap111or1111.1ecK111x pMCiOCOM. BenW-IMHa s xapaKTepM3Y0T CKOpocTb ocei:iaHMSI 4aCTMLI npM ynbTpa4eHTpMcj>yrnpoeaHMM M nponop4MOHanbHa MX MOneKYnSlpHOH Macce. PM6ocOMa npoKapMOTOB (70S) COCTOMT M3 50S M30S cy6beJ:IMHMl.I, syKapMOTOB (SOS) - COCTOMT M3 cy6b0J:IMHMLI SOS M40S. PM6ocoMbl syKapMOTOB MnpoKapMOTOB pa3nM4alOTCSI no MoneKynsipHOH Macce cy6b0AMHMLI, KOnM40CTBY MOneKyn pPHK, Macce pPHK, KOnM40CTBY M pa3H006pa3MIO 6enKOB, cnoCOOHblX CBSl3blBaTb cne41o1cj>1o11.1ecKMe nMraHJ:lbl.

148

o6pa3yioTCH m:6pH)J,Hhie ,D;BOHHbie crr:upa;m, co-

.n;epxam:ue uenh D:HK o6pa3ua 1 :u uerrh D:HK

06pa3ua 2, r,D;e rrp:ucyrcTByIOT KaK CIIMPaJIM30BaHHhie, TaK M Hecrr:upaJIM30BaHHhie yqaCTKM.

B Hecn:upaJIM30BaHHhIX yqacTKax <}>parMeHThI ueneti: D:HK He KOMIIJieMeHTapHhl, T.e. B xo.n;e r:u6- p:U,D;M3aU:UM IIOJiyqaIOTCH HeCOBeprneHHhie r:u6p:u- ,!l;hI. MeTo,n;oM MoJieKyJIHpHoti: r:u6p:u,D;:u3au:u:u MOXHO ycTaHOB:UTh:

CXO,Ll;CTBO M pa3JI:uq:ue rrepB}fqHOM CTpYKTYPhI pa3HhIX o6pa3UOB ttyKJie:UHOBhIX KMCJioT;

paJJI:uq:ue D:HK, Bhl,lJ;eJieHHhIX M3 opraH:U3MOB pa3HhIX BH)J,OB;

• H)J,eHT:uqHOCTh D:HK Bcex opraHOB M TKaHeti: O,!l;HOfO opraHM3Ma.

Ilp11 npoBe,D;eH:u11 r:u6p11,D;11Jau1111 D:HK-PHK

6hIJIH Bhl,lJ;eJieHhl r116pH)J,Hhie MOJieKYJihl, CO,D;epxa- m:ue o,D;Hy uenh D:HK :u O,D;HY uerrh PHK. EcJI11

,D;JIH 3Kcnep11MeHTa 6hIJIM B3HThl D:HK" PHK (nepBMqHhIM TpaHCKPMIIT), BhI,D;eJieHHhie M3 O,!l;HOfO opraHM3Ma, TO o6pa30BhIBaJIMCh COBeprneHHhie r:u6pH)J,hl, noTOMY qTo MOJieKYJia PHK KOMIIJieMeHrnptta uen:u JJ:HK. f :u6p11.n;113al(:ueti: LJ:HK- PH K

6hmo BnepBhie ycTaHOBJieHo, qro Bee BH)J,hI PHK

KJieTKM MMeIOT Ha MOJieKYJie JJ:HK KOMruieMeHTapHhie yqacTKM.

+

AHK

AHK

.QeHarypHpOBaHHbleAHK r1o16pHA AHK-AHK

 

 

06pa3e41

06pa3e4 2

 

 

 

A

+

,QHK

PHK

QeHaryp1o1poeaHHb1e AHK 1o1 PHK

r1116p1o1A AHK-PHK

 

 

6

 

PMC. 4-13. rM6pMAM38~MR HyKneMHOBblX KMCnOT. A - r116p11i:1113al.111~ .QHK-.QHK; 6 -

r116p11i:111aaL111~ .QHK-PHK.

149

SSB-6e~KH

II. PEnnlllKAUlllSI

)l(n]3b1e opraHM3MhI B TeqeHtte S-cPaJhl KJieToqHoro UMKJia, KOTOprui. npe,ll,rnecrnyeT ,!l;enettmo KJieTKM, y,!l;BaMBaIOT CO,!l;epJKaHMe LI.HK TaKMM o6paJOM, ~o KaJK,!l;ruJ. ,!J;Oqepmui: KJieTKa rrocne ,!l;enemra rronyqaeT Ha6op xpOMOCOM, M,!l;eHTMqHbIM pO,!l;MTeJibCKOH KJieTKe. I1pouecc y,!l;Boettm1 xpoMocoM Ha3hlBaIOT peIIJittKautteii (pe,!J;yIIJIMKautteii).

XpoMocoMa conepJKMT onHY tterrpephrnHyIO ,!l;ByxuenoqeqHyIO MOJieKyny LI.HK. I1p11 perrJIMKaUMM KaJK,!l;aH uenh ponttTeJihCKoii nByxuerroqeqHOH LI.HK CJIYJKMT MaTpttueii ,!l;JI5I CMHTe3a HOBOH KOMIIJieMeHrnpttoii uerrtt. BttoBh o6pa3oBaHHaH ,!l;BOMHaH CIIMpaJib MMeeT OllHY MCXOllHYIO (poaMTeJibCKYIO) M OllHY BHOBb CMHTe3MpoBaHHYIO (,!1;0- qepHIOIO) uerrh. TaKoii MexaHM3M YllBOeHMH LI.HK IIOJiyqHJI Ha3BaHMe «UOJiyKOHCepBaTHBHaH pen~u­ KaIUIH» (pttc. 4-14). I1epBMqHa51 CTpyKTypa ,!l;oqepHeM uerrtt orrpeaeIDieTcH nepBMqHoii CTPYKTypoi1 POllMTeJibCKOM uenM, IIOTOMY qTo B OCHOBe ee o6pa30BaH1rn Jie)J(JfT rrpMHUMII KOMJIJieMeHTapHOCTM OCHOBaHMM (G =: C M A = T).

<l>epMeHThI M 6eJIKM, yqacrnyiow:11e B peIIJI11KaUMM, llOJIJKHbl pa60TaTb 6bICTPO M TOqHo. 3TM ycJIOBMH BblTIOJIH51IOTC51 c TIOMOW:bIO oco6oro MYJibTM¢epMeHTHOf0 KOMIIJieKca.

PeIIJittKaUMIO MOJKHo proaenttTh Ha 4 ::irnrra: 06prooBaHtte peIIJIMKaTMBHOH BMJIKM (11HMU11aUH51), CMHTe3 HOBhIX uenei1 (3JIOHraUHH), MCKJIIOqem1e rrpaiiMepoB, JaBeprneHtte CMHTeJa aByx aoqepHMX uenett LI.HK (TepMMHaUHH).

A. HHHUHAII.HH PEIIJIHKAUHH

CMHTeJ LI.HK y 3YKaPMOTOB rrpottcxoaMT B S-¢roy KJiewqHoro u11KJia. MHMUMaumo peIIJIMKaUMM pefYJIMPYIOT crreutt<t>lfqeCKMe CMfHaJibHhie 6eJIKOBbie MOJieKyJihI - ct>aKTOpb1 pocra. <l>aKTophI poem CBH-

3hIBaIOTC51 peuemopaMM MeM6paH KJieTOK, KOTOpbie nepe.n:aIOT Cl1fHaJI, II06YJK,!J;aIOW:MH KJieTKY K HaqaJiy peIIJIMKaUMM (CM. paJ,!l;eJI 11).

C11HTe3 HOBhIX O,!l;HouenoqeqHhIX MOJieKYJI ,ll,HK MO)K_eT IIPOM30MTM TOJibKO IIPM pacxoJK,!l;eHMl1 pollHTeJihCKMX ueneii. B orrpeneneHHOM caiiTe (ro'l- Ka na'l~a pen~HKa....u) npo11cxo.n:ttT JIOKaJihHaH ,!l;eHaTypaUM51 LI.HK, uerrM paCXOL(51TC51 M o6pa3yIOTC51 ,!l;Be pen~HKaTHBHbie BH~KH, ,!l;BIDKYIImec51 B npOTMBOIIOJIOJKHblX HanpaBJieHHHX.

B o6prooBaHMM peIIJIMKaTMBHoii Bl1JIKM npttHMMaeT yqacT11e PM 6eJIKOB M ¢epMeHTOB. TaK, ce- Meiicrno ,ll,HK-Torrott30Mepa3 (I, II M Ill), o6naarui. ttyKJieaJHOH aKTHBHOCThIO, yqacrnyeT B peryJUIUMM cynepcn11pan113au11tt LI.HK. Hanp11Mep, ,ll;HK-ro- nou3oMepua I pa3phrnaeT ¢oc<t>m<t>11pHyio CB513h B 0,UHOH 113 ueneii llBOHHOH cn11pan11 M KOBa.JieHTHO rrpttcoe.n:ttH51eTCH K 5'-KoHuy B ToqKe propbrna (pttc. 4-15). f1o OKOHqamrn ¢opMHpOBaHM51 peIIJIMKaTMBHOH Bl1JIKH ¢epMeHT n11KBM,UttpyeT pa3phrn B uerrtt M OT,!l;eJUieTC51 OT LI.HK.

Pa3pbrn BO,!l;opo,!l;HhIX CBHJetl: B ,!l;Byxuerroqeqttoii MOJieKyJie LI.HK ocyw:ecTBJI51eT ,Il;HK-xe~HKala. <l>epMeHT ,ll,HK-xeJIMKa3a MCIIOJih3YeT 3HepmIO AT<l> rorn pacIIJieTeHM51 ,n:BOHHOH cnttpantt LI.HK.

B peJyJihTaTe rrpottcxo.n:ttT pacKpyqttBaHue yqaCTKa cynepcnttpa.TI1130BaHHOH MOJieKyJihl LI.HK. B rro.n:aepJKaHMM 3Toro yqacTKa LI.HK B pacKpyqeHHOM cocTOHHMM yqacTByIOT

(oT aHrn. single strand binding proteins, T.e. 6eJIKM, CB513bIBaIOW:MeC51 c OllHOUenoqeqHbIMM HMTHMM LI.HK). SSB-6enKu, He JaKphrnaH aJOTMCThIX ocHOBaH11tt, CB513hIBaJOTC51 c O.UHOUerroqeqHOM LI.HK no BCeM llJIMHe pa3,!l;eJIMBUJMXC51 ueneii 11 TaK11M o6pa30M npe,!l;OTBpaw:aJOT MX KOMIIJieMeHTapHoe cKpyq11BaH11e 11 o6pa3oBaHtte «UJIIMJieK». OHM 06naaaroT 60JiblIIMM cpo.n:CTBOM K OllHOUerroqeqHhIM yqacTKaM ,ll,H K, HeJaBMCMMO OT rrepBMqHott CTPYKTYPhI ueneii.

3' C-A-G-A-C-C 5'/

' '

I I I I

''

5' G-T-C-T-G-G 3'

~

C-A-G-A-C-C ---- 3' y-f;\-~-1;\-<?-<?

5'

'

' '

'

!

:

 

5' G-T-C-T-G-G 3'

3' c;;-l;\-~-1;\- <?-<?

5'

 

I

I

I

I

 

I

I

 

'I

'I

3'

G-T-C-T-G-G----. 5' G-T-C-T-G-G

PMC. 4-14. nonyKOHC8pB8TMBH8R pennMKB~MR.

150

 

AHK

1

AHK-ronm130Mepa3a I

T1-1p-OH

H

,QHK

T1-1p-OH

AHK-xen1-1Ka3a

PMC. 4-15. YLIBCTMe .QHK·TOnOM30Mepa3blI B o6p830B8HMM peMMKBTMBHOM BMnKlt. 1 - cfJepMeHT pacU1enni:1er C>,QfiY 4enb llHK; Me)l(,Qy OCTaTKOM rnpo311Ha MOneKYnbl cfJepMeHTa 11 cpocqJopHblM OCTaTKOM 4en11 o6paayerci:1 KOBaneHTHal'!CBl'l3b;2 - npOHCXC>,QITT noKanbHOO pacKpyY11eaH11e i:1eoli\Holi\ cn11pan11 np11 yYacrn11 QHK-xen11Kaab1; QHK-rono11a0Mepaaa I eoccraHaenHeaer cpocqiosqw1pHy10 cel'!3b.

B. 3J10HrAU,1rn

PerrJIHKaUIDI .UHK ocymecTBJIJieTcH .UHK-Jamr- CHMhIMH .UH K-rro1mMepaJaM11 (p11c. 4-16). Cy6cTpaTaM11 11 HCTO'-IHHKaM11 3Hepr1111 ,IIJlJI Cl1HTe3a npo.uyKTa CJI)')J(aT4 MaKp03Pfl1'-IeCK11Xcoe)ll1He- Hl1Jl - .ueJ0Kcttptt6ottyKJieoJMTP11<Pocq>aTbI llAT<I>,

.uIT<l>, .ull,T<l> l1 .uTT<J>, ,IIJlH aKTHBaJJ;HH KOTOphIX Heo6XO.UHMhl HOHhl Marnm1. HetfrpaJI113Y5l OTpMUaTeJihHhIH 3apH)l ttyKJieOTMOB, OHM TIOBhllllaIOT l1X peaKUHOHttyIO cnoco6HOCTh. <l>epMeHThl rrpm!BJIHIOT KaTaJIHTWieCKYJO aKTHBHOCTh TOJihKO B npHCYTCTB11H npe)lBapttTeJihHO pacKpyqeHHOH MaTPWIHOH

.usyxuerro'-le<£Hoii.UHK. Cttme3 uerre:H .UHK rrpoHCXO)lHT B HanpaBJieH11H 5'~3' paCTYllleH uerrtt, T.e. O'-lepe)lHOHttyKJieOTM np11coe)ll1H5leTC5l K CB06o.u- HOMY 3'-0H-KOHUYnpemnecrnyiolllero HyKJieOTMHoro ocTaTKa. CttttTeJttpyeMM uerrh Bcer.ua aHTHrrapaJIJieJihHa MaTPWIHOH uerrH. B xo.ue perrm1xauH11 o6paJyIOTcJI 2 .uoqeptt11e uerm, rrpe.ucTaBJI5IIOllll1e co6o:H KOIIHH Ma'fPH'!HhIXuerre:H.

B Cl1HTe3e 3yKap110Tw1ecKMx .UHK rrpHHHMaIOT yqacTHe 5 ,I(HK-noJI11Mepa3 (a, p, y, o, £). ,I(HK- DOJIHMepa:iw pa3JIW-IalOT no '-IHCJIYcy6be,Ul1H11U, MOJieKYAAPHOH Macce, accouttaUH11 c pa3HhIMH BCIIOMOraTeJThHhIMH 6eJIKaMH, ycKOPJIIOIUHMH rrpouecc 6HOCHHTe3a LI.HK, H cPYHKUMOHaJThHOMY Ha3Ha-

qeH1110. .UHK-IIOJIHMepa3hI a (aJihcpa), p (6eTa), () (.UeJihTa), £ (3IICHJIOH) yqacrny10T B Cl1HTe3e .UHK B JI,Upe KJieTOK, .UHK-IIOJIHMepa3a y (raMMa) - B peIIJIHKaum-1 MllTOXOH,UpHaJihHOH .UHK.

.UHK-IIOJIHMepaJbl p, 0, £ He MOfYT HHHUimpoBaTh o6paJOBaHtte .uoqepHMX uene:H, TaK KaK He HMeIOT cpo.ucrna K 0.11,HHO'-IHOHHllTII .UHK l1HHUlfHPYeT perrJI11Kau1110 .UHK-noJIHMepa3a a, KOTopaJI KoMIIJieMeHTaptta onpe.ueneHHOMY caifTY o.uttouenoqeqHoii LI.HK. 0pHCOe.l1,HH5l5ICh K HeMJ, .IlHK-rro- JIHMepa3a a C11HTe311pyeT He60JihllIOH cJ;>parMeHT PHK - rrpa:HMep, cocro5lllll1H HJ 8-10 p116oHyKJie- OTMOB. .UHK-rrOJIHMepaJa a COCTOHT H3 qen.1pex cy6be.l1,11HHII. KaxolaJI H3 cy6be.l1,l1R11D cJ;>epMeHTa BhlTIOJIHJieT onpe.ueJieHHYIO cPYHKUHIO: «Y3HaBaHHe» caH:Ta peTIJIHKauirn, cttHTe3 rrpa:HMepa (8-10 p116oHYKJieorn.uoB), CHHTe3 ¢parMeHTa u;erm ,LI.HK, OKOJIO 50 .ue3oKcttptt6oHyKJieQTMoB. TaKHM o6paJOM,

.UHK-IIOJil1Mepa:3a a CHHTe3ttpyeT OJI11fOttyKJieQTM, co.uep)!(aIQ11H rrpHMepHO 60 ttyKJieOTMHhIX OCTaTKOB; nepBhie 8-10 rrpe.ucrn.BJieHhI ptt6ottyKJieorn.uaMl1 (npattMep), a OCTaJlhHbie - ,Ue30KClfpl16oHyKJ1e- OTMaM11.

,I(HK-noJIHMepa3a o

OJ1ttrottyKJ1eOTM, CMHTe3ttpoBaHHhitt )l.HK-no- JIHMepa30H: a 11 o6pa3YJ0llllfi1 He6oJihIIIOtt .usyxue-

151

ttyK-
0Ka3a-

PHK-npal1Mep

5'

flMAl1PYIOIJ.t8A

1..1enb

 

PHK-npalliMep YAJ111HAerci:i

 

SSB-6enK111

 

AHK-no111o1Mepa30H & AO scrpe~111

/

AHK-xe.nMKa3a

 

c APYn.tM npaHMepoM

 

~§WC~

 

 

 

 

 

 

 

 

PHK-npal-!Mep ablpe3aerci:i

 

 

 

 

 

 

J:IHK-nonHMepa3ol1 fl

 

 

 

 

 

 

 

 

 

0AH04000'-l~Hb1e

 

pa3pb1Bbl C06,Qf/IHAeT

PHK-npail!Mep

AHK-n11ra3a

 

~-~

 

 

.QHK-no111t1Mepaaa &

3'

 

 

AHK-non11Mepa3a ~

PMc. 4-16. PennMK81.1MR.

rroqeqHhIH <PparneHT c MaTp11u;eM:, rroJBOJHieT rrp11coeJJ:HHHThCH )J,HK-rroJIHMepa3e 0 H rrpo;:i:OJDKHTb CHHTe3 HOBOH uerrH B HarrpaBJieHHH OT 5'- K 3'- KOHUY no xo.n:y pacKpyq11BaHHH peilJIHKaTHBHoif

Bl1JIKM.

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153

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