підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

NH

II

H:zN-C-NH

OH

CTpenTOM1114111H

PMc. 4-45. HeKOTOpble 8HTM6MOTMKM-MHrM6MTOpbl CMHTeaa 6enKOB y npoKapMOTOB.

CTbIO H cpaBHHTeJihHO MaJIO TOKCMtlHbI wrn ''feJIOBeKa. 3TO o6bHCHHeTCH pa3JIHqJ1HMH B CTpyicrype PHK-noJIHMepa3, PHK u 6eJIKOB pu6ocoM B 3yKaPHOTHqecKHx M npoKapttOTHqecKHX KJieTKax.

B. BHPYChl 11 TOKCHHhl - HHfllBHTOPhl

MATPWIHhIX CHHTE30B B 3YKAPHOTWIECKHX

KJIETKAX

Bupycbl

f eHeTWieCKHM MaTepHaJI BHpycoB npe,11,CTaBJieH MOJieKyJIOH ,[I.HK HJIH PHK. Ott, KaK npaBHJIO, HeBeJIHK ll co,11,ep)l(IIT HH<l:>OpMalUfIO JIIDIIb 0 HeKOTOPhIX cneuuct>wiecKHX 6eJIKaX H <l>epMeHTax, Heo6xo,11,HMhIX JlJIH penpo;zy1a.urn supyca (HanpttMep, supycos ocnhl, rpttnna, rroJIHOMHeJIHTa, renaTHTa). BcKope nocne 3apIDKeHHH c BhICOKo:U cKopocThIO Haq1rnaeTCH CHHTe3 BHPYCHhIX ,[I.HK, PHK H 6eJIKOB c HCTIOJib30BaHHeM ¢>epMeHTOB 11 6eJIKOB, cy6-

CTPaTOB H HCToqHHKOB 3Heprtttt KJieTKH x03H11Ha. Ilpll 3TOM B 11HcPHUllpOBaHHhIX KJieTKax npeKpaIUaeTCH CHHTe3 HyKJieHHOBblX KMCJIOT M 6eJIKOB, cso:UcTBeHHhIX opraH113MY X03HHHa. Penpo,11,yKUHH BttpycHhIX qacTHU Mfa BTIJIOTh JJ,o rn6entt 3apIDKeHHoli KJieTKH.

T0Kcnn&1

IlpttqllHOM m6eJIH JII0,11,ett nptt OTpaBJieHMl1 6ne,11,HOH noraHKOH Amanita phalloides HBJIHeTCH TOKCMH - a.-aMaHHTHH, KOTOpblH co,11,ep:xrnTCH B TeJie rptt6a M Bbl3hlBaeT Heo6paTMMYJO ,11,HCcPYHKUHIO neqeHH H noqeK. BhICOKaH TOKcttqHOCTh 3TOro coeJJ,HHeHHH JJ,JIH qeJIOBeKa CBH3aHa c TeM, qTO OHO ttHrn6upyeT 3YKap110TuqecKHe PHK-noJIHMe- pa3hI. Hatt6oJihllIYJO qyscrnttTeJihHOCTh K MY 06ttap)')KHBaeT PHK-rroJIHMepa3a II, KaTaJil13ttpYJOlllaH CHHTe3 MPHK. ,[(ml a.-aMaHttTHHa LD50 (,11,03a per OS, npll KOTOpOH norn6aeT 50% JIHU, nonyqHBumx TOKCHH) COCTaBJIHeT 0,1 Mr/Kr MaCCbl TeJia.

qpe3Bhlqai1:HO TOKCHqeH 6eJIOK PHIUfll, Bbl)],eJieHHblH tt3 KJieIUeBMHhI o6hIKHOBeHHOH. OH npeJJ,cTaBAAeT co6oi1: N-rnHK03HJia3y, KOTOpaH y,11,aJIHeT OJJ,HH OCTaTOK a,11,eHMHa ll3 28S pPHK 60JiblllOH cy6oe- )1,HHHl.J,bl ptt60COMhl ll HHrH6HpyeT CMHTe3 6eJIKa y

3YKapttoToB. PttuHH - 6eJIKOBhIH KOMIToHeHT Ka-

cToposoro Macna, HHoma ucnoJih3yeMoro B Ka'-Ie- crne CJia6HTeJibHOrO cpeJJ,CTBa. lfa-3a BbICOKOH TOKClfqHOCTH pttuttHa neqeHtte KacTopoBhlM MacnoM npoBOMT KopoTKHMM KYPCaMH, TaK KaK JlJIHTeJihHOe ynoTpe6JieHHe MmKeT Bhl3BaTb HerrpeKpaIUalOllJHHCH IlOHOC, Hapyrnett11e pa60Tbl KHlllelJHMKa

ll ,11,IDKe rn6eJib 60JibHOfO.

y qeJIOBeKa pa3BMTMe HeKOTOpbIX 6aKTepttaJih-

HblX HHcPeKl.J,MH corrpOBO)KJJ,aeTCH MHrtt6ttposaHMeM MaTp11'-IHhIX CMHTe30B. Hatt6onee M3yqeHHhlH npMMep - HHrM6MpOBaH11e CHHTe3a 6eJIKOB

noro naToreHHOro MMKpoopraH113Ma rronyqaIOT reH TOKCHHa OT 6aKTepHaJihHOro BMpyca, Ha3hIBaeMOro ~-ct>aroM, KOTOpblH HHcPMU,11pyeT 6aKTepmo tt 11H,11,Yll,HpyeT CHHTe3 TOKCMHa - 0,11,HOUerroqeqHoro 6enKa c MoneKyAApttoM: Maccoi1 60 K,U. B u11TOTIJia3Me KJieTOK X03HHHa TIO}], BJIHHHMeM rrpoTeOJIMTlfqeCKHX ¢>epMeHTOB TOKCHH pacIUeIIJIHeTCH Ha

2 cpparMeHTa, OJJ,11H 113 KOTOpbIX HBAAeTCH cpepMeHTOM A,Ll;<l>-ptt603HJITPaHc¢>epa30H. 3ToT cpepMeHT KaTaJIM3MpyeT A)J,<l>-ptt60311Jil1POBaHHe M

1maKTHBau1uo q>aKTopa 3JIOttrauHH EF-2 rro peaKUHH:

EF-2 + NAD+ -t N1,Cl>-ptt603HJI-EF-2 +

HHKOTHHaMH)J. + H+.

B ycJIOBIDIX in vitro 3Ta peaKUIDI o6paTHMa, HO rrpH pH H KOHUeHTpaUHH HHKOTHHaMH)J.a, KOTOphle cyw:eCTBYJOT B KJieTKax, Otta CTaHOBHTCH tteo6paTHMOM. Mo)J,HQJHKaUIDI ¢anopa EF-2 ttapyrnaeT TPattcJioKaumo ptt6ocoM, Be)J,eT K rrpeKpamettH10 6HOCHHTe3a 6eJIKOB B HHQJHUHPOBaHHbIX KJieTKax H K HX rn6eJIH. c )J,eMCTBHeM TOKCHHa CBH3aHbl ocHOBHbie CHMITTOMbl )J,HQJTepHH.

OrrttcattbI H ,ll,pyrtte TOKCHHhl 6aKTepHaJihHOro H paCTHTeJihHOfO rrpoHCXO)!()J,eHHH, HHrH6HPYIOmHe CHHTe3 H QJYHKUHOHaJibHYlO aKTHBHOCTb 6eJIKOB rryTeM N1,Cl>-pu6o3HJIHPOBaHIDI HJIH MO)J,HQJHKaUHH pPHK.

r. IIHTEP<l>EPOHb1

l1ttTep¢epotthI - tte60JihIIme 6eJIKH (rnHKorrpoTeHHhI), COCTO~e rrpHMepHO H3 160 aMHHOKHCJIOTHblX ocrnTKOB. OHM ceKpeTHpYIQTCH HeKOTOphlMH KJleTKaMH I103BOHOqHblX B OTBeT Ha 3apIDKeHHe BHpycaMH u rrpeITHTCTBYJOT pacrrpoc'rpaHeHHlOBHpycHoif HHQJeKUHH. 3TOT KJiacc 6eJIKOB CHHTe3HpyeTCH B HCKJllOqlfTeJibHO MaJibIX KOJIH'leCTBaX:OT HaHOrpaMMOB (10-9 r) )J,0 ITHKOrpaMMOB (10- 12 r), HO HB.JIBeTCH oqeHh aI<THBHhIM ttecrreutt¢H'lecKHMrrpoTHBOBHpycHbIM areHTOM (106-109 e)J,HHHU aHTHBHpycHOM aKTHBHOCTH Ha 1 Mr 6eJIKa). 3To COOTBeTCTBYeT crroco6HOCTH O)J,HOH MOJieKYJibl HHTep¢epoHa JaJJ.IHmaTb OT HHQJeKUHH 0)1,Hy KJieTKY.

HeK0Topb1e KOMrroHeHThI BHPYCHhIX qacTHU (HarrpHMep, )J,Byxuerro1-ie'rnaHPHK) HH)J,yUHPYIDT CHHTe3 no KpaifHeM Mepe 3 THITOB HHTep¢epoHOB. y 'IeJIOBeKa HMelOTCH 14 reHOB, KO)J,Hpy10mHx a-HHTep¢epOHbl, KOTOpbie rrpO,ll,YUHPYlOTCH B-JIHMQJOUHTaMH H MaKpo¢araMH, 5 rettoB ~-HH­

Tep¢epoHOB, 06ecrreq1rna10mttx o6pa3oBaHHe cooTBeTCTBYIOmHx 6eJIKOB QJH6po6JiaCTaMH, H 1 reH y-HHTep¢epoHa, 3KcrrpeccHH KOToporo H)J,i:lr B T-JIHMQJOUHTax.

CBH3hIBaHCh c peuerrTopaMH Ha rma3MaTH'IecKOi1: MeM6patte 3apIDKeHHblX KJieTOK, 3TH 6eJIKH, I10)J,06HO 6eJIKOBbIM ropMOHaM, CTHMYJIHPYIOT CHHTe3 ¢epMeHTOB, crroco6HbIX pa3pyrnaTb MPHK BHPYCOB H npeKpamaTb CHHTe3 6eJIKOB Ha pH6oCOMax, rrpeITHTCTBYH TeM caMblM 3KCrrpecCHH BHPYCHhIX reHoB B KJieTKax 3yKapHOTOB.

MccJie)J,oBattHe MexaHH3Ma )J,ei1:CTBHH HHTep¢e- poHOB rroKa3aJIO, 'ITOOHH:

•HHrH6HPYIOT CHHTe3 6eJIKOB, He06XO)J,HMbIX )J,JlH peIDIHKaUHH BHpycoB;

•CTHMYJIHPYIOT CHHTe3 QJepMeHTa OJDII'OuyKJieo-

Tll!UIOJIHMep83bl, KaTaJIH3HPYIOmero o6paJOBaHHe tte60JlbllIHX KOJIH'IeCTB KOpoTKHX OJIHroa)J,eHHJiaTOB: 2',5'-OJIHfO (A). 3TH OJIHfOHYKJieOTH)J,bl HBJUIIOTCH aKTHBaTOPaMH pH60HYKJiea3hl - ¢epMeHTa, pacmeIDIH10mero Ma'fPH'IHhle ptt6ocoMHb1e PHK;

• CTHMYJIHPYIOT CHHTe3 rrpoTeHHKHHa3bl, KOTOpag ¢oc¢opHJIHpyeT H, TeM caMbIM, HHaKTHBHpyeT ¢aKTop HHHUHaUHH eIF2:

elF2 + AT<I> -t eIF2-0P03H 2 +NJ.Cl>.

B peJyJihTaTe CHHTe3 ocex 6eJIKOB B HHQJHUHPOBaHHhIX KJieTKax rrpeKpamaeTcH. KneTKH rrortt6aIOT' HO BMeCTe c HHMH OCTaHaBJIHBaeTCH pa3MHO- )KeHHe BHpycoB, H Ha'IHHaeTcH BhIJ)J,opoMeHHe.

TaKHM o6pa30M, )KeprnyH tte6oJihllIHM KOJIH'leCTBOM KJieToK, opraHHJM JamHmaeT ce6H OT 6oJie3HH.

B HacToHmee BpeWI 11mep¢epotth1, ITOJIY'leHHhie rrpOMbilllJleHHhIM rryTeM c HCI10Jlb30BaHHeM TeXHHKH KJIOHHPOBaHHH reHOB, llIHPOKO HCI10Jlb3YIQT rrpH Jie'leHHHo6bI'IHOMrrpOCTY)J,bI, rpHITIIa, ITOJIHOMHeJIHTa, Be'rpHHOMOCITbl, reprreca, BHpyca rerraTHTa H ,ll,pyrttx HHQJeKUHH. Xoporn11e pe3yJihTaThI ITOKa3blBaeT HCITOJib30Bam1e HHTep¢epOHOB B TeparrHH HeKOTOpbIX BH)J,08 3JIOKa1.JeCTBeHHblX orryxoJieM, rnaBHbIM o6pa30M, reM06JiaCT030B (CM. pa3)J,eJI 15), xorn HX pOJib B XHMHOTeparrHH orryxoJieM )J,0 HaCTOHmero BpeMeHH OCTafacg MaJIOITOHHTHOM.

VII. PErYfls:IU"1s:I 3KCnPECC"1"1

rEHOB y nPO- "1 3YKAP"10TOB

OprattHJMhI a)J,arrTHPYIOTCH K MeHHIOI.llHMCH ycJIOBHHM OKp)l)Ka10meM: cpe)J,bl IlYTeM H3MeHeHHH 3KCrrpeCCHH (CKOpOCTH 'rpaHCKPHITUHH)feHOB. 3TOT rrpouecc, B )J,eTMHX HJyqeHHhlH Ha 6aKTepHHx H BHpycax, BKJilO'laeTB3aHMO)J,eMCTBHe crreUH¢H'leCKHX 6eJIKOB c yqaCTKaMH ,ll;HK B Herrocpe)J,CTBeHHOM 6JIH30CTH OT CTapTOBOfO yqacTKa 'rpaHCKpHITllHH. IlpH 3TOM MO)KeT rrpoHCXO)J,HTb BKJilO'IeHHe IDIH BhIKJI10'lett11eTPattCKPHITIIHH. 3yKap110TH'lecKHe KJieTKH HCITOJib3YIOT TOT )Ke CaMblH rrpHHUHII, XOTH B peryJIHUHH peaJIH3YJOTCH H HeKOTOpble ,ll,pyrHe 6oJiee CJIO)KHble MexaHHJMhl.

A. PErYJrn:u;m1 AKTMBHocm rEHOB

Y TIPOKAPllOTOB. TEOPllB OTIEPOHA

11eeJie.n;onamrn: Ha KJieTKax E. coli no3BOJIHJm yeTaHOBHTb, '-ITOy 6aKTepMH: eyrueeTB)'KlTQ:>epMeHTbI

KOHCTHTYfllBHbie, npMeYTeTByIOIUMe B KJieTKax B noeTOSIHHbIX KOJIMqeeTBaX He3aBMeMMO OT MeTa6om1qeeKOro eoeTOSIHHSI opraHM3Ma (Hanp11-

Mep, Q:>epMeHTbI fJIMKOJIH3a);

uu,n,yu,upyeMbie, MX KOHIJ,eHTpall,MSI B 06bJqHbIX yeJIOBMHX MaJia, HO MO)!(eT B03paeTaTh B

pa3 11 6oJiee, eeJIM, ttanpMMep, B epeny KYJihTMBMpoBaHHSI KJieTOK no6aBMTb ey6eTpaT TaKOro Q:>epMeHTa;

penpeccupyeMble, T.e. Q:>epMeHThI MeTa60JIMqeeKHX IIYTeH:, eMHTe3 KOTOPhIX npeKparuaeTeH npM no6aBJieHMM B epeny BbipaIUMBaHMSI KOHeqHo-

ro npOLJ:YKTa 3TMX IIYTeH:.

1. Teopua onepoua

Ha oeHOBaHMM reHeTMqeeKHX 11eeJieLJ:OBaHMH MH-

LJ:YKll,MM P-raJiaKT03HLJ:a3hI, yqaeTByrorueH: B KJieTKax E.coli, B rHLJ:pOJIMTMqeeKOM paerueTIJieHMM JiaKT03hI (pMe. <I>paHeya )l(aKo6 M )l(aK MOHO B

1961 r. eQ:>opMyJIMpOBaJIM rnnOTe3y onepotta, KOTopaH 06bS1eHS1Jia MexaHM3M KOHTpOJISI eMHTe3a 6eJIKOB y npoKapMOTOB.

B 3KenepMMemax rMnoTe3a onepotta nonyqHJia TIOJIHOe TIOLJ:TBep)l(,ll,eHMe, a npeLJ:JIO)!(eHHbIH B HeH: TMTI peryJIHUMM eTaJIM Ha3bIBaTb KOHTpOJieM e11HTe3a 6eJIKa Ha ypoBHe TpaHeKp11Illl,MM, TaK KaK B 3TOM enyqae M3MeHeHMe eKopoeTM eMHTe3a 6eJIKOB oeyrueeTBJIHeTeSI 3a elleT 113MeHeHMSI eKopoeTM TpaHeKpMilll,MM reHOB, T.e. Ha eTa.IlMM o6pa30BaHMSI MPHK.

y E. coli, KaK M y LJ:PYfMX npoKap110TOB, )1,HK He OTLJ:eJieHa OT 11,MTOTIJia3MhI Sl,UepHOH o6oJioqKoH:.

B npoueeee TpatteKPMIIIUrn o6pa3yroTeH nepB~hie TpatteKPlliIThI, He eonep)!(aI1U1e HHTpOHOB, a MPHK JIHIIIeHbI «K3na» M noJIM-A-Kottua. CMHTe3 6eJIKa

Haq11HaeTeS1 ).1,0 TOfO, KaK 3aKaHlJHBaeTeH eMHTe3 ero MaTpHll,bl, T.e. TpaHeKPHilUHSI M TpaHeJISIIJ,HH npOTe-

KaIOT IlOlfflf OLJ:HOBpeMeHHO. 11eXOM M3 pa3Mepa reHOMa (4xl06 nap HyKJieOTMLJ:OB), Ka)l(,ll,aSI KJieTKa E. coli eonep)l(IIT HHQ:>OpMaUHIO 0 HeeKOJibKHX ThieHqax 6eJIKoB. Ho npM HOpMaJibHhIX yeJIOBHHX poeTa

Otta eHHTe3HpyeT OKOJIO pa31ImHhIX 6eJIKOB, a 3TO 03HalJaeT, lJTO MHOrMe reHbI He TpaHeKpM6MpYIOTeH, T.e. HeaKTHBHbl. feHhI 6eJIKOB, Q:>ytt-

K11,H11 KOTOpbIX B Mera6onmeeKHX npoueeeax Teetto

eBSl3aHbl, qaeTO B rettoMe rpynnMp)'KlTeSIBMeCTe B eTPYKTYPHhJe en11tt1111,b1 (onepoHb1). Cornaetto TeopMM )l(aKo6a H Motto, onepottaMM Ha3hIBaIOT yqaeTKH MOJieKYJibl )1,HK, KOTOphre eonep)!(aT HHQ:>opMaU11IO 0 rpynne Q:>yttKUHOHaJibHO B3aMMOeBSl3aHHbIX eTpYKTYPHbIX 6eJIKOB, M peryJIHTOPHYIO 30tty, KOHTpoII11PYKlIUYIO TpaHeKpMnUMIO 3THX rettoB. CTpYKTYPHhie reHbI orrepoHa 3KerrpeeeMpyroTeH eornaeoBaHHO, JIM6o Bee OHM TpaHeKpM6HPYIOTCSI, 11 TOrLJ:a orrepoH aKTHBeH, JIM60 HM OLJ:MH 113 reHOB He «npolJHTbIBaeTeSI», M Torna orrepoH HeaKT11BeH. Korna onepoH aKTHBeH M Bee ero reHbI TpaHeKpM6MPYIOTeS1, TO e11HTe311pyereS1 noJIMUMeTpOHHaH MPHK, eny- )!(aIUaH MaTpm1eH: LJ:JIH e11HTe3a Beex 6eJIKOB 3TOro orrepoHa. TpaHeKpMIIUHH eTpyKfYPHhIX rettoB 3aBMeHT OT enoeo6ttoeT11 PHK-rroJIMMepa3hI rrp11eoeLJ:H- HHTbeS1 K npoMOTOpy, paenOJIO)!(eHHOMY Ha 5'-KOH- ue onepotta rrepen eTpYKTYPHhIMM rettaM11.

CBH3hIBaHMe PHK-noJIHMepa3bI e npoMornpoM 3aBMeMT OT np11eyTeTBHSI 6eJIKa-perrpeeeopa Ha eMe)!(HOM e IIPOMOTOPOM yqaeTKe, KOTOpbIH Ha3blBaIOT «OnepaTOp». EenoK-perrpeeeop eMHTe3MpyeTeH B KJieTKe e rroeTOSIHHOH eKopoeTbIO M MMeeT eponerno K orreparnpttoMy yqaeTKY. CTPYKTYPHO yqaeTKH rrpoMoTopa M oneparnpa qaeTMllHO rrepeKPhlBaIOTeH, rro3TOMY npMeoeLJ:MttettMe 6eJIKa-perr- peeeopa K orrepaTopy eo3naeT eTep11qeeKoe rrpeIIHTeTB11e MH rrpMeoen11ttett11S1 PHK-rroJI11Mepa3bI.

EoJibIIIMHeTBO MexaHH3MOB peryMUHM eMHTe3a 6eJIKOB HarrpaBJieHO Ha 113MeHeHMe eKopoeTl1 eBSl3hIBaHHSI PHK-rroJI11Mepa3hI e npoMoTopoM, BJIHHH TaKHM o6pa30M Ha 3Tan HHHUMaUHM TpaHeKpMIIUMM. feHbI, ocyrueeTBAAIOIUHe eMHTe3 peryJISITOpHbIX 6eJIKOB, MOfYT 6hITb YLJ:aJieHhI OT orrepoHa, TpaHeKpMIIUHIO KOTOporo OHM KOHTpOJIMPYIOT.

2. HuiJyKU,UH cuume3a 6e.aKo8. Lac-onepou

Teop11S1 onepotta 6bIJia rrpeLJ:JIO)!(eHa Ha oettoBaHMM )laHHbIX, IIOJiyqeHHbIX npM M3yqeHMl1 eBOHeTB JiaKT03ttoro onepotta (lac-onepotta) E. coli, T.e. orrepotta, B KOTOPOM 3aKOLJ:MPOBaHbI 6eJIKH, yqaeTByIOIUMe B yeBOeHHH JiaKT03bI.

KneTKH E. coli o6billHO paeJYI Ha epene, MeIIOJih3YH B Kaqeerne MeToqHMKa yrJiepona fJIIOK03y. Een11 B epeJie KYJihTMBMpoBaHHSI fJIIOK03Y 3aMeHMTh Ha LJ:MeaxapHLJ: JiaKT03y, TO ITO npoIIIeeTBMM HeeKOJibKHX MMHYT KJieTKH a,uarrT11pyroTeS1 K 113MeHMBIIIMMeSI yeJIOBHSIM. OHM HalJMHaIOT rrpoJIYu11poBaTb 3 6eJIKa, o6eenellMBaIOIUHX YTHJIM3aUMIO JiaKT03bI. 0LJ:MH M3 3TMX 6eJIKOB - Q:>epMeHT P-ranaKT03HLJ:a3a, KaTaJIM3Mpy10ru11H: mJipOIIMTM-

+Hp

OH

OH naKT038

PMC. 4-46. rM,qponM3 naKT03bl p-ranaKT03MA830M.

qecKoe pacrueTIJieHHe naKT03hI )J.O rmoK03hI H ranaKT03hI.

B rrpncyrcTBHll rmoK03hI KJieTKH E. coli co)J.ep)KaT MeHee 10 MOJieI<yJI 3THX <f>epMeHTOB Ha KJieTI<y. IlepeHOC KJieTOK Ha cpe;zy, CO)J.ep)KaJIIYIO JiaKT03y, Bbl3hIBaeT HHJzyKIIHIO - ysennqeHtte KOJinqecTBa MOnekyJI KaJKJJ,Oro H3 <f>epMeHTOB )J,0 5000 (pttc.

TeopIDI orrepoHa o6'h51CH51eT3TO 51BJieHHe cne- )J.YIOIUHM o6pa30M. B oTcyrcrntte HH;zyKTOpa (naKT03hl) 6enoK-perrpeccop CB513aH c orrepaTopoM.

A rrocKOJihI<y yqacTKH orrepaTopa H rrpoMoTopa rrepeKphrna10TC5I, TO rrp11coe)J.11HeH11e perrpeccopa K orrepaTopy rrpen51TCTByeT CB513hIBaH1110 PHK-rro- n11Mepa3hI c rrpoMOTOpOM, 11 TpaHcKp11nu1151 CTPYKTYPHhIX reHoB orrepoHa He H)J.eT. Kor)J,a B cpe)J.e Il051BJI5IeTC51 11HJ:zyKTOp, T.e. JiaKT03a, TO OH npnco- e)J,llH51eTC51 K 6enKy-perrpeccopy, 113MeH51eT ero KOH<f>opMaUllIO ll CHll)KaeT cpO)J.CTBO K onepaTopy. PHK-rronnMepa3a CB513hrnaeTC51 c rrpoMOTopoM n TpaHcKptt6npyeT CTPYKTYPHhie retthI.

3. PenpeccUH cunme1a 6eAK06. Tpunmot}Jan06btU u lUCfflUOUH06blU onepOHbl

CHll)KeHne KOHuempau1m <t>epMema B 6aKTepnaJibHOM KJieTKe MO)KeT OCYIIJ:eCTBJI51ThC51 rryreM penpeCCl1.ll cHHTe3a <f>epMeHTOB. CYIIJ:HOCTh 3TOro MexaHH3Ma peryAAUHll 3aKJI10qaeTC5I B CJie)J.yIOrueM: KOr)J,a KJieTKH E. coli pacTyr Ha cpe)J.e, co)J.ep~ruei1 B KaqeCTBe emrncTBeHHoro HCTOqH11Ka a30Ta COJih aMMOHll51, TO HM rrpHXO)J,MTC51 CllHTe311POBaTb BCe a30TCO- )J.ep)KaIIJ:l1.e BeruecTBa. TaKHe KJieTKH, B qacTHOCTll, )J,OJDKHhI CO)J.epJKaTb BCe <t>epMeHThl, He06XO)J,HMhie )J,JI5I CHHTeJa 20 pa3llttqHhIX aMl1HOKl1CJIOT. 0)1.HaKO ecJIH )J.06aBMTh B cpe;zy I<yJibTHBllpOBaHH51 0)1.HY H3 aMHHOKHCJIOT, HanpHMep TPHITTo<t>aH HJll1 IBCTH)J.HH, TO KJieTKa rrepecrnHeT Bhrpa6aThIBaTh Bech Ha6op

~ +

OH

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