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підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

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8-10

5' ---------

- A - G - C - T - T - A -----------

3'

3'

T-C - G - A - A - T

AHK

5'

ATCl>~j

rTCI> ~ PHK-nom1Mepaaa

LITCI>~ ""'

Ko.1:1111py10ll.\a5! 4enb AHK

5'~

-A-G-C-T-T-A- ~

\ ____ 3'

 

P1<1c. 4-28. TpaHcKp1<1n~MR PHK Ha MaTpM"IHOK 1.1enM JlHK. C111HTea PHK acerAa npo111cxOAlllT B HanpaaneH111111 5'-t3'.

CrpoeH111e npoMoTopa 3YKap111oros, yaHaeaeMoro

---i

 

 

PHK-nom1Mepaaoi71 II

l

 

 

40 n.H.

~~---

 

 

 

 

25 n.H.

5'

 

 

~~~~~~~~~~~~~3'

.QHK

~~~~~~~~~~~~~s·

3'

 

 

 

 

i

 

 

 

 

 

 

crapTrpaHcKP111n4111111

P1<1c. 4-29. CTpoeHMe npoMoropa ayKapMOToe. npoMoTOpHble aneMeHTbl -

cne4111cj>1111.1ecK111e nocneAOBaTenbHOCTlll HyKneOTlllAOB,

xapaKTepHb1e AnR mo6oro npoMOTopa, ceRab1Ba10lJ1ero PHK-non111Mepaay. nepab1~ npoMOTOpHbl~ aneMeHT - nocneAoaaTenbHOCTb ATATAA- (TATA-6oKc) OTAeneH OT ca~ra Ha4ana TpaHcKp111n4111111 np1116n111a111TenbHO Ha 25 nap HYKnBOTlllAOB (n.H.). Ha paccTORHllllll np111MBpHo 40 (lllHOrAa AO 120) n.H. OT Hero pacnonaraeTCR nocneAOBaTenbHOCTb GGCCAATC- (CAAT-6oKc).

Ka, B KOTopo:H MaTpwua ,UOCTyrIHa )l)UI l1Hl1Ul1aUHl1 cwHTe3a uenw PHK (pwc. 4-30).

TiocJie Toro KaK Cl1HTe3HPOBaH OJillfOHyKJieOTM,[( l13 HYKJieOTH,n:HblX OCTaTKOB, cr-cy6be- ,[(MHMUa OT):(eJUieTcH OT PHK-noJIMMepa3hI, a BMeCTO Hee K MOJieryJie <i>epMeHTa npwcoe,[(l1H51IOTC51 HeCKOJibKO <i>aKTopoB 3JIOHrauuu.

3JiouraIUU1

<l>aKTOpbl 3JlOHrauww IIOBbIIIIaIOT aKTMBHOCTb PHK-nonwMepa3hI w o6nePiaIOT pacxo)K)J,eHwe uenei1 ,l],HK. CuHTe3 MoneryJihI PHK ll):(fa OT 5'- K 3'-KOHUY KOMIIJieMeHTapHO MaTpll'IHOHuenw ,lJ,HK. Ha cTa,n,wu 3JIOHrauuw, B o6JiaCTH TpattcKpurruu-

OHHOH BMJIKM, 0,[(HOBpeMeHHO pa3,[(eJieHbl rrpttMepHO 18 HYKJieOTH):(HbIX nap ,UHK. PacTyIIIui1 KOHeu uenw PHK 06pa3yeT speMeHHYIO rn6pH):(HYI0 crrwpaJib, OKOJIO 12 nap HyKJieOTH,n:HbIX OCTaTKOB, c MaTpl1qHoi1 uenhIO ,[(HK. no Mepe IIPOJ1.BH.lKeHH51 PHK-noJIMMepa3hI no MaTpuue B ttarrpaBJieHHM OT 3'-K 5'-KOHUYBnepe):(M Hee rrpOMCXO).l.HT pacxo.lK- ,neHtte, a Il033,JJ,l1 - BOCCTaHOBJieHMe )1.BOHHOH CIIHpaJIM )],HK.

TepMHHaIUUI

PacKpyq11Bam1e ,n:BOMHOH crrwpanw ,l],HK B o6JiaCTM cai1Ta TepMMHauHM ,[(eJiaeT ero ):(OCTYflHblM )])ISi <i>aKTopa TepMMHaUHM. 3aBepIIIaeTrn CHHTe3 PHK B

164

4-31 ).
5',5'-

npoMOTOp

Cai:ir repMV1H8l\V1V1

 

,D,HK

 

l.r T~-cpaKrnp

:::::><~~><=><:=::><:::><::..C>C: ,D,HK

2

 

,,----- PHK-nonviMepa3a

 

]

PHK-nonviMepa3a

,,----- cPaKTOpbl V1HV1l\V18l\V1V1

 

 

 

3

PHK-rpaHcKpvinr 5'

,D,HK

4 )__-- <l>a"op '"'"""a"""

Perym1pHb1e cpaKTOpbl :1~ PHK-nonviMepa3a npe-PHK

5'V\/VVV'-- 3'

,D,HK

P11c. 4-30. CTaAMM TpaHcKp11n41111. 1 - npvicoeAviHeHvie TATA-cjlaKropa K npoMoropy. Yro6b1 npoMorop 6b111 ysHaH PHK-no11viMepa- soili, Heo6xOAVIMO o6pasosaHvie rpaHcKpvim.ivioHHoro KoMn11eKca TATA-cjlaKTopffATA-6oKc (npoMorop). TATA-cjlaKTOP ocraercfl CBflsaHHblM c TATA-6oKCOM so speMf! rpaHcKpvim.ivivi, :;iro 0611erYaer vicnollb30BaHvie npoMoropa MHOrVIMVI Mo11eKy11aMV1 PHK-no11viMepa- 3b1; 2 - o6pa30BaHVl8 rpaHCKpVlnl.IVIOHHOili BVlllKVI; 3 - 31lOHraL.jVl5'1;4 - repMVIHaUVIS'l.

cTporo onpe.neneHHbIX y'rnCTKax MaTp11UbI - TepM1rnarnpax (cai1ThI TepMHHaUIDt). <DaKTop TepM11Harnu1 o6nefl!aeT orn.eJ1eH11e nepswrnoro TpaHcKp11nrn (npe-MPHK), KOMnJleMeHTapHoro MaTp11ue, l1 PHK-non11Mepa3bI OT MaTpttUbI. PHK-nom1Mepa3a MO)l(eT BCTYflHTb B rne.nyIOlllHH UHK.11 TpaHCKp11nUHl1 nocJle np11coe.nm1eH1rn cy6bellMHl1UbI CT.

E. KoBAJIEHTHAfl MO)]H<I>HMUM51

(rrPOUECCHHr) MATPHqttoi\ PHK

nepBl14Hble TpaHCKp11ITTbl MPHK, npe)[(J.J,e lJeM 6y.nyT 11cnOJih30BaHhI s xone c11HTeJa 6enKa, no.n- sepraIOTCH PH.UY KOBarieHTHblX M0Jl11cp11K3Ul1H. 3Tl1 M01ltt¢11Kau1111 HC06XO.Lll1Mbl llJIH ¢yHKUHOH11pOB3Hl151 MPH K B KalJecrne MaTpl1Ubl.

Monml>nKa~usi 5'-Kott~a

Mo.ntt¢11Ka1.1,1111 npe-MPHK HaLJ11Ha10TCJ1 Ha cTa.ntt11 3JlOHra1.1,1111. Korna nJT11Ha nepswrnoro TpaHCKp11nrn nocn1raeT np11MepHo 30 HyKneoTl1llHbIX OCT3TKOB, npo11cxon11T K3IlHPOBaHHe ero 5'-KOH ua. OcymecTBJTHeT K3n11posa1·111e ryaH 11JI11JTTpaHc¢epa3a. <DepMeHT rnnpoJ111JyeT MaKpo3prn4ecKy10 CB513b B MOJleKyJle fT<D 11 npttcoen11HJ1eT HyKJTeOTl1lllll1¢oc¢aTH blH OCTaTOK 5'-

¢oc¢arnol1 rpynnol1 K 5'-KOHUYcHHTeJttposaH- HOro ¢parMeHrn PH K c 06paJ0BaH11eM <Poc¢on113¢11pHoi1 CBJIJl1. CT OCJiellYIOl.l.Lee MeTl1JT11pOB3HHe OCTaTKa ryaHlrna B COCTaBe fT<D c o6pa3osaHtteM N 7-MenrnryaH03HHa JaBepwaeT ¢opM11posaH11e K3na (pnc.

165

splice -

0

~~3

5'

 

)

5'

 

 

 

 

 

 

 

 

0

0

 

 

 

 

 

 

 

0

 

 

 

 

HN

7>-

 

 

 

II

 

II

II

 

 

 

I,,

~

a H

 

r0-\-0-\-0-\-01

 

 

H2N~N

 

 

 

 

o-

0-

o-

 

 

 

 

~H2

 

 

 

 

 

5~~0BOHMO

 

 

 

OH

OH

 

 

 

 

 

0

OH

3'-KOH01..\

 

 

 

 

 

 

 

 

 

 

 

0 =~-0

-vvvvS~pApApApApApA· ·····pA- OH

 

 

 

 

 

 

 

 

 

 

 

o-

MPHK

 

7-MernmyaHoa111Hrp111cpoccpar (K3n)

nonviA-nOCI10AOBaT0IlbHOCTb

 

P11c. 4-31. KoeaneHTH8H MOAllQlllK81.111H KOHQeBblX HyKneOTMAHblX OCT8TKOB nepBll'·IHOroTpaHCKpllnTa MPHK.

MoJJ,11qmu11poBaHtthIH 5'-KoHeu o6ecrre'rnBaeT

HH.Huwaumo TpaHcJisruww, YMttmi:eT Bpewi: )!Gl3Htt MPHK, 3aw,muaSI ee OT JJ,eHCTBHSI 5'-3K30HYK- Jiea3 B UHTOIIJia3Me. K::irrwpoBaHwe Heo6xoJJ,ttMO ,llJISI HHHU:ttauww CHHTe3a 6eJIKa, TaK KaK .HH.HUHMPYIO- w,we TpmmeThI AUG, GUG pacrro3ttaIOTCSI pw6ocoMoif TOJihKO ecJitt rrpttCYTCTByeT K3II. HaJittqwe K3na TaK)!(e Heo6XO)J,HMO ,!l;JISI pa60Thl CJIO)!(HOH Q:>epMeHTHOH CHCTeMbl, o6ecrreq.HBaIOW,eH y,n:aJie- H.He HHTpOHOB.

Mo.uuct>nKaUIDI 3'-Kouua

3'-KoHeu 60JihlliHHCTBa TpaHCKp.HIITOB, CHH-

Te3.HpOBaHHhIX PHK-rroJI.HMepa3oif II, TaK)!(e

rro,n:BepraeTcH Mo,n:wQ:>wKauw.H, rrpH. KOTopoif crre-

UHaJihHhIM Q:>epMeHTOM IIOJIAA-IIOJI.HMepa30H Q:>op-

MHpyeTCSI IIOJrnA-rrocJie,n:oBaTeJibHOCTh (nOJIMA-

«XBOCT»)' COCTOSIW,aSI lf3 100-200 OCTaTKOB a,n:e-

H.HJIOBOH KHCJIOThl.

CHrHaJIOM K HaqaJiy rroJIH.a,n:eHHJIHpoBaHHSI srBJISieTcsr IlOCJie,n:oBaTeJibHOCTh -AAUAAA- Ha pacTyw,etl: uerr.H PHK. <l>epMeHT IIOJIHA-rroJIHMe- pa3a, rrpOSIBJISISI 3K30HyKJiea3HYIO aKTlfBHOCTh, pa3phIBaeT 3'-Q:>ocQ:>o3Q:>H.pHyIO CBSl3h IIOCJie IIOSIBJieHHSI B uerrH PHK crreuHQ:>HqecKoif rrocJie- ,n:oBaTeJihHOCTH -AAUAAA-. K 3'-KOHUY B rnqKe pa3phIBa IIOJIHA-IIOJIH.Mepa3a ttapaW,HBaeT IIO- JIHA-«XBOCT». HaJIH.qtte rroJIHA-nocJie,n:oBaTeJih- HOCTH Ha 3'-Kottueo6JierqaeT Bh1xo,n: MPHK H3 sr,n:pa H 3aMe,llJISieT ee flfll:POJI.H3 B UlfTOIIJia3Me.

<l>epMeHThl, ocyrn,eCTBJISllOW,.He K3II.HpOBaHH.e H IIOJIHa,n:eHHJittpOBaHHe, .H36H.paTeJibHO CBSl3bIBaIOTCSI c PHK-rroJIH.Mepa3oif II, H B OTCYTCTB.He IIOJIHMepa3hI HeaKT.HBHhl.

Cm1aucuur nepBH'IHbIXTpaucKpumoe MPHK

c rrmmrreH.HeM MeTO,!l;OB, II03BOJIHI01UHX lf3Y'!aTh

rrepBmttyIO CTpyicrypy MOJieKYJI MPHK B umorrml3Me If IIOCJie,!l;OBaTeJihHOCTh ttyKJieorn,n:OB KOll:lfPyIOW,eH ee reHOMHOH )lHK, 6hmO yCTaHOBJieHO, 'ITO OHM He KOMIIJieMemap8h1, a ,llJIH.Ha retta B HeCKOJihKO Pa3 6oJihllie «3peJIOH» MPHK. ITocJie,n:osaTeJihHOCTH ttyK-

Jieorn,n:oB, rrpHCYTCTByIOllJ,He B ,[J,HK, HO He BXOll:SlUJ,He B cocraB 3peJioif MPHK, 6h1JI11 Ha3BaHhI HeKoll:HPYIOllllfMH, HJIH Hlfl)JOHbl, a IIOCJie,n:OBaTeJihHOCTH,

np11cyrcTByIOllJ,11e B MPH K, - KOll:HPYIOllJ,11M.H, lfJIM

3K30HbI. TaK.HM o6pa30M, rrepBmHhlli TpaHCKpHIIT - CTporo KOMIIJieMeHTapHasr MaTpH.ue ttyKJieHHOBaSI K.HCJIOTa (rrpe-MPHK), co,n:ep)l{arn,asr KaK 3K30Hhl, TaK H HHTpOHbl. ,[l.rmtta HHTpOHOB Bapb.Hpyer OT 80 AO I000 ttyKJieorn,n:oB. ITocJie,n:oBareJihHOCTH HHTpOHOB «Bblpe3aIOTCSI» H.3 rrepBmttoro TpaHCKpmrra, KOHUbl 3K30HOB coell:HHIDOTCSI JJ.PYf c ,n:pyroM. TacyIO MOll:lf- Q:>HKaUHIO PHK Ha3hIBaIOT «cwraH:cuur» (OT attrJI. to

cpalli,11BaTh). CIIJiaHCHHf rrpowcxoa.HT B Mpe, B UIITOIUia3MY rrocryrraer ~e «3peJiasr» MPHK.

f etthI ::iyKap11oroB co,n:ep)l{aT 6oJihllle HHTpOHOB, qeM 3K30HOB, II03TOMY oqeHb ,llJI.HHHhie MOJieKYJihI rrpe-MPHK (OKOJIO 5000 ttyKJieOTMOB) IIOCJie crmaifCH.Hra rrpeBpaw,aIOTCSI B 6oJiee KOpOTK.He MOJieKYJihl U.HTOIIJia3MaT.HqecKoif MPHK (oT 500 ,n:o 3000 HYKneoTMOB).

ITpouecc «Bhipe3aHHSI» HHTpOHOB rrpoTeKaeT rrpH. yqacTHH MaJihIX sr,n:epHbIX ptt6oHyKJieorrpoTeMHOB (MHPHIT). B cocraB MHPHIT BXOJJ.HT MaJiaSI MepHaSI PHK (MHPHK), ttyKJieorn,n:ttasr UeIIh KOTOpoif CBSI3atta c 6eJIKOBhIM OCTOBOM, COCTOSIW,HM H.3 HeCKOJihKHX rrpoToMepoB. B cIIJiatfcwttre rrpH.HHMaIOT yqacrne pa3JIMqHhie MHPHIT (p.Hc. 4-32).

166

HyKJieOTl1llHb1e rrocJieaoBaTeJibHOCTH HHTPOHOB cl:>YHKIUIOHa.JibHO HeaKTl1BHbI. Ho Ha 5'- H 3'-KoH-

uax OHl1 11MeIOT BbICOKOCrreu11¢w1ecK11e IIOCJieaoBaTeJibHOCTl1 - AGGU- 11 GAGG- coorneTcrneHHO (cattTbl CITJiaHCMHra), KOTOpbie o6ecne1-lMBaIOT 11x yaa.JieHMe 113 MOJieKyJibI rrpe-MPHK. lfaMeHeHMe CTPYKTyPbJ 3Tl1X nocJieaoBaTeJihHOCTeH: BJIM51eT Ha rrpouecc cITJia.ti:c11Hra.

Ha rrepBott CTMM11 rrpouecca M.s:rPHIT CBH3bIBaIOTCH co crreu11¢w1ecKMMM ITOCJie).l.OBaTeJibHOCTHMM rrepBJ1l!HOfO TpaHCKPl1IITa (cattTbl crrJiattCMHra), ).l.aJiee K HMM rrpMCOe).l.MHHIOTCH apynie MHPHTI. Tip11 ¢opM11poBaHMM CTPYKTYPhI crrJiaticocoMbI 3'-KOHeu 0)..1.HOfO 3K30Ha c6Jimt<:aeTC51 c

5' -KOHUOM cJieay10mero 3K30Ha. CrrJiaH:cocoMa KaTaJIM3MpyeT peaKUMIO pacmerrJieH11.s:r 3' ,5'-<l>oc- ¢oa113¢11pHott CBH3M Ha rpamrue 3K30Ha c HHTpoHoM. TiocJiellOBaTeJibHOCTb MHTpoHa yaan.s:reT- c.s:r, a )..I.Ba 3K30Ha coe).l.MHHIOTCH. 06pa30BaHHe 3',5'-¢oc¢oa113¢HpHOHCBH3M Me)K,Uy llBYMH 3K30HaMM KaTaJIM3MpyIOT M51PHK (MaJible .s:raepHbie PHK), BXOllHllJMe B CTPYKTYPY cnJiaH:cocoMhI.

B pe3yJihTaTe crrnaH:cirnra M3 rrepBM'-lHbIXTpattc- KpttrrToB M PHK o6pa3yIOTCH MOJieKyJihI «3penoti» MPHK.

All:bTepuaTHBHbIH CDJiaiiCHHf nepBH11HblX TpaHC-

KpHUTOB MPHK

)1,JIH HeKOTOpbIX reHOB OllMCaHbl aJibTepHaTMB-

Hble nyTM CITJiaHCMHra M ITOJIMa).l.eHMJIHPOBaHMH 0)..1.- HOfO M Toro )Ke TpaHCKPMIITa. 3K30H 0)..1.HOfO BapMaHTa CITJiaHCMHra MO:lKeT OKa3aTbCH MHTPOHOM B aJibTepHaTMBHOM rryTM, Il03TOMY MOJieKyJibl MPHK, o6pa3oBaHHbie B pe3yJihTaTe aJibTepHaTMBHoro CilJiaHCHHra, pa3llM1-laIOTCH Ha6opoM 3K30HOB. 3To rrpMBOllHT K o6pa3oBaHMIO pa3HbIX MPHK 11, COOTBeTCTBeHHO, pa3HbIX 6eJIKOB c 0)..1.HOfO rrep- BM'-lHOfOTpaHCKpMrrTa. TaK, B rrapa¢oJIJIMKyJIHpHbIX KJieTKax llJMTOBHllHOH :lKeJie3bl (pMC. 4-33) B XO,ll,e TpaHCKPMilUMM reHa ropMOHa KaJibUMTOHHHa (CM. pa3).l.eJI 11) 06pa3yeTCH nepBH'-lHbIH TpaHCKpMilT MPHK, KOTOpbIH cocTOHT 113 rnecTM 3K3oHOB. MaTpH1!Ha.s:r PHK KaJibUMTOHHHa 06pa3yeTc.s:r rryTeM CITJiaHCHHra nepBblX 1-leTbipex 3K30HOB

(1-4). TiocneaH11M: (qernepTb1tt) 3K30H coaep)Kl1T CHfHaJI IlOJIMaaeHMJIHPOBaHHH (nocJie).l.OBaTeJibHOCTb -AAUAAA-), Y3HaBaeMbIH nonHA-noJI11Me- pa3oti B napa¢0JIJI11KyJIHpHbIX KJieTKax llJHTOBMllHOH :lKeJie3hI. 3ToT )Ke rrepBl11!Hb1M: TpaHcKp11m B KJieTKaX fOJIOBHOfO M03ra B XO,ll,e llPYfOfO (aJibTep-

 

 

 

Ca!Ar cn11a1Ac111Hra

 

V1HrpoH

Caih cn11a!Ac111Hra

 

 

 

~~~~~'8K30H

------GAGG-~,--~8~K3_0_H~2~--~,

3'

5'I

1

 

1-AGGU-

 

I

/

 

 

 

 

4>opM111posaH1<1e

 

lr MS!PHn

 

 

 

 

 

 

cnnaillcocoMbl

 

 

 

 

 

 

 

 

 

 

 

 

 

 

lr MSIPHn

 

 

 

 

 

 

 

 

 

 

CnnaillcocoMa

 

 

 

5'

 

 

 

 

 

 

 

 

 

 

K30H 1

b----8-=-K3_0_H_2______,3'

 

 

' ------------ ..

 

 

 

 

 

 

 

 

 

 

 

«Bb1peJaHHbll'it» LolHTPOH

MSIPHn

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

BKJOH 1

 

 

8K30H 2

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

 

«3penasi»

MPHK

 

 

 

 

 

 

PMc. 4-32. CnnaMCMHr PHK. B npo1..1ecce cnnaiilc11ttra np11tt11Mat0r y'lacrnepaan11'lttb1eMsiPHn, KOTOpb1e cpopM11pyt0r cnnai:icocoMy. MsiPHn, saa11M0Aei:icrsysi c PHK 11 Apyr c ApyroM, cp11Kc11pyt0r 11 op11ettrnpytor peaK1..1110HHb1e rpynnb1 nepsH'lttororpattcKp11ma. Karan11TH'lecKasicpyHKl..IHSI cnnai:icocoM o6ycnosnetta PHK-cocrasnsit01411M11; TaKHe PHK ttaab1sat0r p116oa11MaM11.

167

4-35).
4-34).
14-16
-CCA,

8K30H

8K30H

8K30H

8K30H

8K30H

8K30H

reH

 

 

 

 

 

1-6 3K30Hbl

 

 

 

 

/

 

 

 

BU111Toa11~HOH >1<ene3e

 

8 K!leTKax ronoaHoro M03ra

 

 

 

 

 

 

 

 

./

4 r-- non11A

""

~

 

 

 

I

 

I 2 I 3 I 5 6 ~ non11A

2

3

 

PMc. 4-33. Ani.1epHBTMBH1>1iil cnnaiilCMHr reHa KBnbl.IMTOHMHa. B KllSTKax Ull1TOBl1AHOl1 >t<ene3bl cnnai1c11Hr nepew·iHororpaHCKp11ma np11eoA11T K 06pa3oeaH1110 Kanbl..ll1TOHl1Hoeoi1 MPHK, BK1110'"lat0U1ei14 SK30Ha 11 non11A-nocneAoearenbHOCTb, Kornpasi 06pa3yercsi nocne pacU1enneH11si rpaHcKp11ma e nepaoM y'"lacrKec11rHana non11ai:ieH11n11poeaH11S1. B KlleTKax Moara 06pa3yercsi MPHK, coAep>t<aU1asi: SK30Hbl 1, 2, 3, 5, 6 11 non11A-nocneAoearenbHOCTb, 06pa3oeaHHy10 nocne eroporo c11rHana non11ai:ieH11n11poeaH11si.

HaTHBHoro) rryn1 crrnail:cttHra rrpeBpalll,aeTcH B MPHK KaJihUHTOHHHITO)l06Horo 6enKa, OTBeqaIOlll,ero 3a BKYCOBoe BocrrpttHTtte. MaTPHqHa.H PHK 3TOro 6enKa cocTOITT H3 rrepBhIX TPex 3K30HOB, 06lll,HX c KaJihUHTOHHHOBoil: MPHK, HO BKJIIOqaeT )lOnoJIHHTeJihHO ITHThIH ll rneCTOH 3K30Hhl, He CBOHCTBeHHhie MPHK KanhUHTOHHHa. IllecToil: 3K30H TO)l(e llMeeT CllfHaJI IIOJllla)leHHJillpOBaHllH

-MUAAA-, Y3HaBaeMblll cpepMeHTOM nonllA-rro- JlllMepa30H B KJieTKax HepBHOii TKaHtt. Bh16op O)l- Horo H3 ITYTeii (anhTepHaTHBHhiii CI1Jlail:cttHr) H O)lHOro ll3 B03MO)l(HbIX cail:TOB I10Jllla)leHllJ1llpOBaHllH mpaeT BaJKHYIO POJih B TKaHecrreumt>HqeCKOH 3KCrrpeCCllll reHOB.

Pa3Hhie BapttaHThI CI1Jlail:CttHra MOI)'TrrpttBO)lllTh K o6pa30BaHHIO pa3HbIX mocpopM O)lHOro ll Toro xe 6enKa. HarrpttMep, reH TporroHttHa cocTOHT H3

18 3K30HOB ll KO)lllpyeT MHOroqttcJieHHhie ll30<popMhl 3Toro MhirneqHoro 6enKa. Pa3Hhie mocpopMhI TPOITOHHHa o6pa3YIQTCH B pa3HhIX TKaHHX Ha orrpe)leJieHHhIX CTa)lllHX llX pa3BllTHH.

B. IlPOI.J,ECCHHf IlEPBHqHhIX

TPAHCKPHilTOB PHBOCOMAJihHOii PHK

H TPAHCilOPTHOii PHK

feHhI, KO)lllpyIOlll,He 6oJihlliyIO qacTh CTPYKTYPHhIX PHK, TpaHcKptt6ttpyIOTCH PHK-rrom-i:Mepa- 3aMH I H III. HYKJie:irnoBhie KHCJIOThI - rrpe)lrneCTBeHHHKH pPHK H TPHK - ITO)lBepraIOTCH B H)lpe paclll,eITJleHHIO ll XllMllqecKOH MO)lllqmKaUllll (rrpouecCHHry).

IIocrrpaucK))HDIUIOHHbie MO)lH4>HKaIUIH nepeH11uoro T)JaHcKpurrra TPHK (npoueccuur TPHK)

IIepBHqHbIB TPaHCKPHIIT TPHK co,llep)l(llT oKono

100 HyKneoTH)lOB, a rrocne rrpoueccttHra -

70-90 ttyKJieOTll)lHhIX OCTaTKOB. IlOCTTPaHCKpllIIUllOHHhie MO)lllqmKau:im rrepBttqHblX TpaHCKPllITTOB TPHK rrpottcXOMT nptt yqacTllll PHK-a3 (pu6ouyK-

Jiea3). TaK, cJ:>opMttpoBaHtte 3'-KoHua TPHK KaTaJIH3HpyeT PHK-a3a, rrpe)lCTaBJIHIOlll,aH co6oil:

3'-3K30ttyKJiea3y, «OTPe3aIOlll,yIO» no O)lHOMY ttyK-

JieOTll)ly, IIOKa He )lOCTllrHeT ITOCJie)lOBaTeJibHOCTll O)lHHaKoBoil: )lJIH Bcex TPHK. ).lJIH HeKOTOphIX TPHK cpopMttpOBaHtte rrocne)loBaTeJihHOCTH -

CCA Ha 3'-KoHue (aKUeITTOpHbril: KOHeu) npottcxo-

)lllT B pe3yJlbTaTe ITOCJie)lOBaTeJlbHOro rrpttCOe)lHHeHHH 3TllX Tpex HyKJieOTH)lOB. Ilpe-TPHK CO)lep)((llT Bee- m O)lllH llHTpOH, COCTOHlll,llll ll3 HYKJieOTll- )lOB. Y)lanetttte HHTpOHa H cITJlail:cttttr rrpHBO)lHT K cpopMttpOBaHHIO CTpYKTYPhl, Ha3hIBaeMOH «aHTHKO- ,!lOH», - TpllI1JleTa ttyKJieOTll)lOB, o6ecrreqwBaIOmero B3al1MO)leil:CTBtte TPHK c KOMI1JleMeHTapHhIM KO)lOHOM MPHK B xo)le cHHTe3a 6eJIKOB (pttc.

IlOCTTpaHCKJ>HDIUIOHHble MO)lH4>HKaIUIH (npoueccuur) nepBH'IHOfOT)JaHCKpHDTa pPHK. <l>opMHpoeauue pu6ocoM

B KJieTKax qeJIOBeKa CO)lepXllTCH OKOJIO COTHll KOilllH reHa pPHK, JIOKaJIH30BaHHhlX rpynrraMll Ha ITHTll xpOMOCOMax. feHhl pPHK TpaHCKptt6llPYIOTCH PHK-rronttMepa3oil: I c o6pa30BaHHeM tt)leHTHqHhIX TpaHcKpttrrTOB. IIepBllqHbie TpaHcKPHITThI llMeIOT )lJlllHY OKOJIO 13 000 HyKJieOTll)l- HhIX OCTaTKOB (45S pPHK). Ilpe)!()le 'leMIIOKll-

HYTh H)lpo B COCTaBe ptt60COMHOH 'laCTHUhl, MOJieKyJia 45S pPHK IlO)lBepraeTCH rrpouecCHHry, B pe3yJihTaTe 06pa3yeTCH 28S pPHK (OKOJIO

5000 HyKJieOTll)lOB), 18S pPHK (OKOJIO 2000 HYKJieOTll)lOB) ll 5,8S pPHK (OKOJIO 160 HyKJieOTll- )lOB), KOTOpbie HBJIHIOTCH KOMITOHeHTaMll p1160-

COM (pttc. OcTaJihHaH qacTh TpattcKpttma

pa3pyrnaeTCH B Mpe.

168

L.l,1111on11aaMa

PMc. 4-34. npoL\ElCCMHr npe-TPHK. Onpe.c:ieneHHb1e aaornCTu1e OCHOBaH11R HYKn901l1AOB TPHK s xQ,Qe npo!.leca1Hra MeT11n11py10TCR noi:i i:iel1CTB11eM PHK-MeT11nasb1 11 npeBpalllalOTCR, Hanp11Mep, B 7-MernnryaH0311H 11 2-MernnryaH0311H (M11HOpHble OCHOBaH11R). B MoneKYne TPHK CQ,QeplKaTCR 11 i:ipyrne H900bl4Hbl9 OCHOB8Hl1Sl - nceB.QOYPl/1Al'IH,A11r11i:1poypl/1Al'IH,KOTOpble TaKlKe MOA11cf>111-111PYIOTCR BO BpeMR npo1-1ecc11Hra.

TIAepHafl ~I

MeM6paHa

U111onnaaua

flAepHaR nopa

PMC. 4-35. 06pa30BaHMe .. BblXOA M3 siiipa cy6beAMHMLI pM6ocoM. B pesynbTSTe npo1-1ecc11Hra 113 MOneKYnbl npe.QwecTBeHHl1Ka 45S pPHK o6pa3ylOTCSl Tp11 rnna pPHK: 18S, BX0Aflllj8Sl B COCTaB Manol1 cy6b0,Ql1Hl1l.lbl p116ocoM, a TaKlKe 28S 11 5,8S, noKanl13YIOllllll9CSl B 6onbWOl1 cy6b0,Q11Hl1l.19. Bee TPl1 pPHK o6pa3ylOTCSl B paBHblX KOnl14ecTBax, TaK KaK OHl1 npo11CXQ,QSlT 113 OAHOro 11 TOro )1(9 nepB114HOro TpaHCKpl1ITTa. 5S pPHK 6onbwol1 cy6b0,Ql'1Hl1l.lblp116ocoM TpaHcKp1161'1pyeTcs:ioTi:ienbHO OT nepB114HOro TpaHCKp11ma 45S pPHK. P116ocoManbHb1e PHK, o6paaosaHHb1e B xoi:ie nOCTTpaHCKp11n1-1110HHblX MOA11cf>11Kal.11111, CBSl3blBalOTCSl co cne411cp114eCKl1Ml1 6enKaMl1, 11 o6pasyeTCSl p116oc0Ma.

169

PH6ocoMa - opraHemrn KJienrn, yqacrny101uaH B 6HOCHHTe3e 6eJIKa. PH6ocoMa 3yKapHoTOB (SOS)

COCTOHT H3 )J,Byx, 60JibllIOM H MaJIOM, cy6oe)J,HHHU: 60S H 40S. Eemrn pH60COM BbIITOJIHHIOT CTpYKTYPttylO, peryJIHTOPHYIO H KaTaJIHTH'IeCKYJOcPYHKUHH.

V. 6lllOClllHTE3 6EJ1KOB (TPAHCJlSIUlllSI)

IlepeBO)J, HH¢>0pMaUHH, 3aKJIIO'leHHOMB TIOJIHHYKJieOTH)J,HOM nocnell,oBaTeJihHOCTH MPHK, B aM11HOKHCJIOTHYIO nocJiell,OBaTeJibHOCTb 6eJIKa Tpe6yeT onpe)J,eJieHHOfO cnoco6a KO)J,HpOBaHHH HJil1 lll11¢>poBamut, T.e. CYllJeCTBOBaHH51 onpe)J,eJieHHOro 3aKoHa, no KOTopoMy qepe,n:oBaHHe 'leTbipex ttyKJieOTH)J,OB B MPHK 3a,n:afu cneu,H¢>H'lecKYJOnocJie,UoBaTeJibHOCTh aMHHOKHCJIOT B 6eJIKe.

A. fEHETJlqECKllH KO.[( JI EfO CBOHCTBA

Heo6xo,n:HMOCTh KO,UHpoBaHAA CTPYKTYPhI 6enKOB B Jil1HeMHOM nocJie,UOBaTeJihHOCTH HYKJieOTH- )J,OB MPHK H LI.HK npo,n:HKTOBaHa TeM, 'ITOB xo,n:e TpaHCJI511J,11H:

• HeT COOTBeTCTBHJI Me)J().J,y 'IHCJIOMMOHOMepoB B MaTpHue MPHK H npo,UyKTe - CHHTe3HpyeMOM 6eJIKe;

OTCYTCTByeT CTPYKTYPHOe cxoncrno Me)J().J,y MoHoMepaMH PHK H 6eJIKa.

3To HCKJIIO'laeTKOMITJieMeHTapHoe B3aHMO,Uei1:- CTBHe Me)l()J,y MaTpHu,ei1: H npo,UyKTOM - npHH- u,11n, no KOTOpOMY OCYllJeCTBJI51eTC51 nocTpoeHHe HOBhIX MOJieKYJI LI.HK H PHK B xo,n:e peITJIHKaUH11 11 TpaHCKpHTIIJ,Hl1.

OrcIO)J,a CTaHOBHTCH HCHbTM, 'ITO)J,OJDKeH CYllJeCTBOBaTb «CJIOBapb», II03BOJI51IOIUHM BbIBCHHTb, KaKaH nocJie,n:oBaTeJihHOCTh ttyKJieOTH)J,OB MPHK o6ecne'IHBaeT BKJIIO'leHHe B 6eJIOK aMHHOKHCJIOT B 3a)J,aHHOH TIOCJie,UOBaTeJibHOCTl1. 3TOT «CJIOBapb» nonyqHJI Ha3BaHHe reHeTH'leCKOfO,6HOJIOrwieCKOro, HYKJieOTH)J,HOfO, HJIH aMl1HOKHCJIOTHOfO KO,Ua. OH no3BOJIHeT urn¢>poBaTh aMHHOKHCJIOThT, BXOMm11e B COCTaB 6eJIKOB, c nOMOIUblO onpe,n:eJieHHOH nocJie,n:oBaTeJibHOCTH ttyKJieOTH,UOB B LI.HK H MPHK. Ll,JIH Hero xapaKTepHhI onpe,n:eJieHHhie cBotl:crna.

TpHIIJieTHOCTb. 0,UH11M 113 OCHOBHbIX BOnpocoB npH Bbl51CHeHHH CBOMCTB KO,Ua 6bIJI BOnpoc 0 'IHCne HyKJieOTH,UOB, KOTOpoe ,UOJI)l(HO onpe,UeJIJITb BKJIIO'leH11eB 6eJIOK o,n:Hoti aMHHOKHCJIOThI. Cpa3y 6bIJIO nOHHTHO, 'ITO 3TO 'IHCJIO He MO)l(eT 6hITb

paBHblM 1 HJil1 2, TaK KaK B 3TOM CJiy'laeKOJIH'leCTBO KO,UHPYIOIUHX 3JieMeHTOB 6y,n:eT He,UOCTaTO'l- HblM ,UJIH llll1cPPOBaHHH 20 aMHHOKHCJIOT B 6eJIKax. qHCJIO KO)J,11PYIOLIJ,HX nocJie,UOBaTeJihHOCTeM 113 qeTblpex HYKJieOTH)J,OB no TPH paBHo 4 3=64, 'ITO60nee 'leMB 3 pa3a rrpeBbTIUaeT Ml1Hl1MaJibHOe KOJIH- 'leCTBO, KOTOpoe Heo6XO)J,11MO ,UJIH KO,Ul1POBaHl151

20 aM11HoKHcnoT. B ,n:anhttetlrueM 6hmo ycTaHOBneHo, 'ITOKO,UHPYJOIU11Ml1 3JieMeHTaMl1 B lll11¢>poBaH1111 aMHHOKHCJIOTHOM TIOCJie,UOBaTeJihHOCTH ,n:ei1:- CTBl1TeJihHO HBJIHIOTCH TpOMKl1 HyKJieOTH)J,OB, HJIH TJ)HilJICTbl, KOTOpb!e IlOJiyqHJil1 Ha3BaHl1e «KO)J,OHbl».

CMbICJI KO)J,OHOB

CMbICJI KO)J,OHOB CTaJI rromneH B 60-x r. XX

CTOJieTl1H, KOf)J,a, HCTIOJib3YH 6ecKJieTO'IHYIOCl1CTeMy CHHTe3a 6enKOB (Ta6JI. 4-3) 11 CMHTern:qecKMe noJIHptt60HyKJieoT11,UbI c 3a,UaHHoti rrocne,n:o- BaTeJibHOCTbIO HyKJieOTH,UOB B Ka11ecrne MaTpHU,bl,

M. HupeH6epr H r. MaTTett CHHTe3HPOBaJIM rroJIHrrerrTH,Ubl onpe,n:enemrnro cTpoeHHH. TaK, Ha MaTpttu,e nOJIH-Y' COCTOHLIJ,ett TOJibKO 113 OCTaTKOB YM<J>, 6bIJI rroJiy11eH noJIH¢>eHMJiaJiatt11H, a Ha MaTp1111e nOJIH-U - TIOJIHnpOJIHH. lfa 3T0f0 CJie,UOBaJIO, 'ITOTpHITJieT - uuu KO,UMpyeT <l>eH, a

Tp11ITJieT -CCC - Ilpo.

B nocJie,UYJOIUHX 3KcnepHMettrnx HcnoJib30Ba-

Jil1 CMelllaHHble Cl1HTeTWJ:eCKHe nOJIHpH60HyKJieOTH)J,bl c M3BecTHbIM cocTaBoM. B pe3yJibTaTe 3TOH pa60Tbl y,n:anOCb ycTaHOBl1Tb, 'ITO113 64 KO,UOHOB BKJIIO'leHHe aMHHOKHCJIOT B CHHTe3MpyIOLIJ,YIOCH nmmnenTH)l.Hy10 u,eflh mtt¢>pyeT 61 TpHITJieT, a 3 ocTaJibHbIX - UAA, UAG, VGA He Ko,n:11py10T BKJIIO'leHHeB 6eJIOK aM11HOKHCJIOT H rrepBOHa'laJibHO 6bUIH Ha3BaHbl 6eCCMblCJieHHblMl1, HJil1 HOH- ceHC-KO,UOHaM11. O,n:ttaKo B ,UaJibHetirneM 6hUio noKa3aHo, 'ITO 3Tl1 TpHnJieTbl CHrHaJIH311PYIOT 0 3aBeprneHl111 TpaHCJIHIJ,1111, H n03TOMY HX CTaJIH Ha3bIBaTb TepMHHHPYJOIIJ,MMl1, HJil1 CTOn-KO,UOHaMH.

Ko,n:OHbI MPHK 11 TPHITJieTbI HYKJieOTl1,UOB B KO-

.UHPYJOIUei1: HHTH LI.HK c HanpaBJieHHeM OT 5' K 3'- KOHIIY 11MeIOT o)J,11HaK0By10 nocJie)l.oBaTeJibHOCTb a30THCTblX OCHOBaHHM, 3a 11CKJIIO'leHHeMTOfO, 'ITO B LI.HK BMeCTO ypau,MJia (U), xapaKTepttoro ,UJIH MPHK, CTOl1T THMHH (T).

CneD.Hcl>H'IHOCTb

Ka)l()J,OMY KO)J,OHY COOTBeTCTByeT TOJibKO 0).1,Ha orrpe).l.eJieHHaH aMHHOKHCJIOTa. B 3TOM CMbICJie reHeTH'lecKHi1:KO)], CTporo O,UH03Ha'leH.

170

Ta611Ml.18 4-3. OcHOBHble KOMnOHeHTbl 6ellOKCMHTe3MPYIOLl.lelli CMCTeMbl

Heo6XO)J.HMbie KOMDOHeHThl

 

 

<J>yHKil,HH

 

 

 

 

 

 

 

 

1. AMHHOKHCJIOThI

Cy6cT)JaThr .L\ITH CHHTe3a 6errKOB

 

 

 

2. TPHK

TPHK BhmOrrHHIDT <liYHKI!HID a.r1anTopoB. OuH aKrrenrnpHhIM KOHI!OM

 

 

B3aHMO,L\eiicTBYIDT c aMHHOKHcrroTaMH, a auTHKO.L\OHOM ~ c KO,UOHOM MPHK.

3. AMHHOarrHrr-TPHK CHHTeTa3hI

KmK.L\aH aa-TPHK-CHHTeTa3a KaTarrH3HpyeT peaKI!HID crrerrH<iiHqecKoro

 

 

CBH3h!Baumr o,uuoii H3 20 aMHHOKHcrroT c cooTBeTCTBy10meii TPHK

 

 

4. MPHK

MaT)JHrra

co.r1ep)l(HT

rrHueiiuy10

rrocrre.r10BaTeJihHOCTh

Ko,uouoB,

 

 

orrpe.r1emr10mHx rrepBHquy10 CTPYKTYPY 6errKOB

 

 

5. PH6ocOMhr

PH6ouyKJieonpoTeHHOBhie cy6KrreToquhie CT)JYKTYPhI, JIBmr10mHec»

MecTOM

 

 

CHHTe3a 6errKOB

 

 

 

 

6. AT<D, fT<D

HcToquHKH 3ueprHH

 

 

 

 

7. EerrKOBhie qiaKTOphr

CnerrH<iiHqecKHe BuepH6ocoMHhre 6errKH, ueo6xo.L1HMhie .L\IT» nporrecca

HHHI!HaI!HH,3JIOHrarrHH,

T)JaHCJIHI!IfH (12 <tiaKTopoB HHHI!Harrim: elF; 2 <tiaKTOpa 3JIOHral!HH: eEFI,

TepMHHaI!HH

eEF2, H <tiaKTOphI TepMHHaI!HH: eRF)

 

 

 

8. lloHhl MarHHH

Ko<tiaKTop, crn6HrrH3Hpy10mHii CTJJYKTYPY pH6ocoM

 

 

npMMe'laHMH:eJF (eukaryotic initiation factors) - cjJaKTOpb1 1t1H1114111a4111111; eEF (eukaryotic elongation factors) - cpaKTOpb1

 

sn0Hra4111111;

eRF (eukaryotic releasing factors) - cpaKTOpb1 rnpM1t1Ha4111111.

 

 

 

 

BL1poQeuuocTb

B MPHK H ,1],HK HMeeT CMhICJI 61 TPHIIJieT, KmK-

.Uhiti: H3 KOTOphIX KO,UHpyeT BKJIIO'IeHHe B 6eJIOK O,UHOH H3 20 aMHHOKHCJIOT. lfa 3T0f0 CJie,n:yeT, 'ITO B HHcPOPMaI..l,HOHHhIX MOJieKYJiaX BKJIIO'IeHHeB 6eJIOK o,n:ttoti: H TOH )!(:e aMHHOKHCJIOThI orrpe.n:emnoT HeCKOJihKO KO,UOHOB. :3-ro CBOHCTBO 6HOJIOfH'IeCKOfO KO,Ua IIOJIY'IHJIOHa3BaHHe BhlpO){QleHHOCTH.

y qeJIOBeKa 0).1,HHM KO,UOHOM 3aIIIHcPPOBaHhl

TOJihKO 2 aMHHOKHCJIOThI

-

MeT H TpH, Tor.n:a

KaK neti:, Cep If Apr -

IIIeCThlO KO,UOHaMH, a

Arra, Barr, fm1, I1po, Tpe -

'IeThiphMHKO.UOHa-

MH (Ta6JI. 4-4).

J.136bITO'IHOCTbKO.UHPyIOIIIHX IIOCJie,n:oBaTeJihHOCTeti: - 1..1,eHHeti:IIIee CBOHCTBO KO,Ua, TaK KaK OHa IIOBhIIIIaeT ycTOH'IHBOCThHHcPOPMaIJ,HOHHOfO IIOTOKa K He6JiarorrpHHTHhIM B03,UeHCTBIDIM BHeIIIHeH H BttyTpeHHeti: cpe,Uhl. I1pH orrpe.n:eJieHHH IlpHPO.Uhl aMHHOKHCJIOThI, KOTopa5I .n:om1rna 6hITh BKJllO'IeHaB 6eJIOK, TpeTHH HYKJieOTH,U B KO,UOHe He HMeeT CTOJih B~oro 3Ha'IeHIDI, KaK rrepBble ,n:Ba. KaK BttnHo H3 Ta6JI. 4-4, )1,Jl5I MHornx aMHHoKHCJIOT 3aMeHa ttyKJieOTH.L(a B TPeTbeH II03HI..l,HH KO- ,UOHa He CKa3hIBaeTC5I Ha era CMhlCJie.

JluueimocTb 3aDHCH HHCj>OpMalJ,HH

B xo.n:e TpaHCAAIJ,HH KO.UOHhI MPHK «'IHTalOT-

CH» c qmKcupoBaHHOH CTapTOBOH TO'IKHIIOCJie,n:o- BaTeJihHO H He rrepeKphrnaioTcH. B 3arrHctt HHcPOP-

MaUHH OTCyTCTBYIOT CHfHa.Tihl, yKa3hIBalOIUHe Ha

KOHeIJ, O,UHOfO KO,UOHa H Ha'IaJIOCJie,UyIOmero.

Ko,n:oH AUG HBJI51eTcH HHHUHHPYIOIUHM H rrpo- 'IHThIBaeTCHKaK B Ha'IaJie,TaK H B .upyrHX yqaCTKax MPHK KaK MeT. CJienyiomHe 3a HHM TpHIIJieThI 'IHTalOTC51 IIOCJie,UoBaTeJibHO 6e3 KaKHX-JIH6o nponyCKOB BIIJIOTb no CTOil-KO.L(OHa, Ha KOTOPOM CHHTe3 IIOJIHITeIITH,UHOH uerrH 3aBepIIIaeTCH.

'YuuuepcaJibHOCTb

,1],o He,UaBHero BpeMeHH C'IHTaJIOCh,'ITOKO,U a6COJIIOTHO yttHBepcaJieH, T.e. CMhICJI KO,UOBblX CJIOB O,UHHaKOB )1,Jl51 Bcex H3YtfeHHhIX opraHH3MOB: BHpycoB, 6aKTepHti:, pacTeHHH, 3eMHOBO,UHhIX, MJieKOIIHTaIOll(HX, BKJIIO'IMqeJIOBeKa. 0.n:HaKO II03,UHee CTaJIO H3BeCTHO O,UHO HCKJllO'Iem1e,OKa3aJIOCh, 'ITO MHTOXOH.UPHaJihHaH MPHK co.n:ep)!(:HT 4 TPHIIJieTa, n:Me10mnx .upyroe 3Ha'IeHHe, 'IeMB MPHK H,UepHoro npoHcxo){QleHH51. TaK, B MPHK MHTOXOH.n:- pnrt TPHIIJieT UGA KO).mpyeT Tpn, AUA - MeT, a AGA n AGG rrpolfttThIBaIOTC51 KaK .n:orroJIHMTeJThHhie CTOII-KO,UOHbl.

KoJiuueapuocTh reua u npo):Q'K'fa

y rrpoKapMOTOB o6Hap~eHO JIHHeti:Hoe COOTBeTCTBHe IIOCJie.L(OBaTeJibHOCTM KO.L(OHOB reHa H IIOCJie.L(OBaTeJTbHOCTH aMHHOKMCJIOT B 6eJIKOBOM IIPO.UYKTe, MJIM, KaK roBOPHT, cymeCTByeT KOJIHHeapHOCTh reHa H IIPOLJ:YKTa.

171

Ta6m11..1a 4-4. reHen1~eCKl1M KOA

IlepBoe OCHOBaHHe

 

 

 

BTopoe OCHOBaHHe

 

 

 

 

 

 

 

 

 

 

 

 

 

u

 

c

 

A

 

G

u

 

UUU<DeH

 

UCUCep

 

UAU Tttp

 

UGU ll,ttc

 

 

UUC <DeH

 

UCCCep

 

UAC Tttp

 

UGC ll,ttc

 

 

 

 

 

 

 

UUA Jieii

 

UCACep

 

UAA*

 

UGA*

 

 

UUGJieii

 

UCGCep

 

UAG*

 

UGG Tptt

c

 

CUU Jleii

 

CCU Ilpo

 

CAU fttc

 

CGU Apr

 

 

CUC Jleii

 

CCCilpo

 

CAC fttc

 

CGC Apr

 

 

CUAJieii

 

CCAilpo

 

CAA fnH

 

CGAApr

 

 

CUG Jleii

 

CCG Ilpo

 

CAG fnH

 

CGGApr

A

 

AUUHne

 

ACU Tpe

 

AAU ACH

 

AGUCep

 

 

AUC Hne

 

ACC Tpe

 

AACAcH

 

AGCCep

 

 

AUAMeT

 

ACA Tpe

 

AAA JIHJ

 

AGAApr

 

 

AUGMeT

 

ACGTpe

 

AAG JIHJ

 

AGGApr

G

 

GUU Ban

 

GCU Ana

 

GAU Acrr

 

GGU fntt

 

 

GUC Ban

 

GCCAna

 

GAC Acrr

 

GGC fntt

 

 

GUABan

 

GCAAna

 

GAA fny

 

GGA fntt

 

 

GUG Ban

 

GCG Ana

 

GAG fny

 

GGG fntt

 

 

 

 

 

 

 

 

 

np11Me~aH11H: u - ypa1..1111n;

c - l..llilT03111H; A -

a,QeH111H; G - ryaHlilH; * -

repMlllHlllPYIOLlllil~ KOAOH.

 

 

y 3YKaPMOTOB IIOCJie.UOBaTeJibHOCTM OCHOBaHMll B reHe, KOJIMHeapHbie aMMHOKMCJIOTHOll nocne.uoBaTeJibHOCTl1 B 6enKe, npephIBaIOTCH MHTPOHaMtt. Il03TOMY B 3YKaPMOTWieCKl'.IXKJieTKax aMMHOKHCJIOTHaH IIOCJie.UOBaTeJihHOCTh 6eJIKa KOJIHHeapHa nocne.uoBaTeJibHOCTM 3K30HOB B reHe HJIM 3peJIOll MPHK rrocne rrocrrpaHcKpttm1,110HHoro y.uaneHHH MHTpOHOB.

Ii. 0CHOBHblE KOMTIOHEHTbl

l>F.JIOKCHHTE3HPYIOIIJ:EH CHCTEMbl

KaK BH.llHO tt3 rn6n. 4-3, .llJIH CMHTe3a rron11nenTH.llHOil uerr11 He06X0.1lMMO 60JibllIOe KOJIWieCTBO KOMIIOHeHTOB, COBMeCTHOe M cornaCOBaHHOe B3aMMO.LJ,eMCTBMe KOTOpbIX npttBO.llMT K o6pa30BaHMIO 6enKa.

AMHHOKHCJIOTbl

Bee 20 aMttHOKHCJIOT, BXOMIUMX B CTPYKTYPY 6eJIKOB opraHM3Ma 'IeJIOBeKa,.llOJI)l(HhI rrpMCYTCTBOBaTb B ,llOCTaTO'IHOMKOJIM'IeCTBe.3TO Tpe6oBaHtte rrpe)l(,lle Bcero OTHOCMTCH K He3aMeHMMhIM (T.e. He CMHTe3MPYJOIUMMCH B opraHM3Me) aMMHOKl'.ICJIOTaM, TaK KaK He.UOCTaTO'IHOe CHa6)l(eHMe KJieTKM XOTH 6hI 0.llHOll He3aMeHHMOll aMMHOKHCJIOTOM rrpttBO-

.llMT K CHIDKeHmo, a MHor.ua M IIOJIHOH OCTaHOBKe CMHTe3a 6eJIKa Ha KO.UOHe, Tpe6yioiueM BKJIIO'IeHMH 3TOH aMHHOKMCJIOTbl B 6eJIOK.

MPHK. Co.u,ep)l(MT ttmt>opMau1110 o CTPYKType Cl1HTe3MpyeMoro 6eJIKa 11 11CIIOJih3yeTCH B Ka'IeCTBe MaTpMUbl.

TPHK. Y 'IeJioBeKaOKOJIO 50 pa3JIH'IHhIXTPHK o6ecrre'IMBaIOTBKJIIO'IeHMeaMHHOKl'.ICJIOTB 6eJIOK. TPHK Ha3hIBaIDT «a,JJ,amopub1e MOJieK)'JibI», TaK KaK K aKUemopHOMY KOHUY 3THX MOJieKYJI MO)l(eT 6hITh rrpMCOe.UMHeHa onpe.ueneHHaH aMMHOKl'.ICJIOTa, a c IIOMOIUhl{) aHTMKO.UOHa OHM Y3HaIDT cneutt<t>wrecKl'.IllKO.UOH Ha MPHK. B npouecce cMHTe3a 6enKa Ha ptt6ocoMe CBH3hIBaH11e aHTMKO.UOHOB TPHK c K0.1lOHaMM MPHK npottcxo.nttT no rrpttHUMIIY KOMnneMeHTapHOCTH M aHTHnapanneJibHOCTM.

AHTIIKO~OH TPHKApr

(3' )-G-C-C-(5')

 

 

Ko~OHhl Apr:

(5')-C-G-G-(3')

0.UHaKO OKa3aJIOCh, 'ITO'IHCJIOTPH K .llJIH Ka)l(-

.uoii aMHHOKHCJIOTbl He COBna.uaeT c 'IMCJIOM K0-

.1lHPYIOIUMX ee KO.UOHOB B MPHK, 11, CJie).l,OBaTeJihHO, HeKOTOphie TPHK crroco6HhI CBH3hIBaTbCH 60JibWe 'IeMc 0).1,HHM KO.UOHOM.

172

11ccnenoBamie 3Toro Borrpoca rro3BOJIJUIO ycrnHOBHTh CJie)IyIOmee:

rrepBbie L(Ba OCHOBamrn KOL(OHa H ITOCJieL(HHe JJ;Ba OCHOBamrn aHTHKOL(OHa o6pa3YJOT 06bl'l- Hble rrpoqHbJe rrapbl (ryaHHH-UHT03HH H ane- HHH-ypauJUI) H BHOCHT HaH60JiblUHH BKJiaL( B cneumfmqHOCTb L(eKO,llHpOBaHHH;

CBH3bIBaHHe TpeThero OCHOBaHHH KOL(OHa c rrepBbIM OCHOBaHHeM aHTHKOL(OHa rrpoHCXOJJ:HT cna6ee, qeM c rrepBbIMH JJ:BYM51, H 3TO I103BOJI51eT HeKOTOpbIM TPHK rrpoqHThIBaTh 6oJihllle

qeM O,llHH KO,llOH.

f11rr0Te3a, 06bHCHHIOIUaH xapaKTep KO,llOH-aH- THKO,llOHOBOro B3aHMOL(eHCTBH51, IlOJiyqHJia Ha3BaHHe «fHUOTe3bl Ka'laHHH» (T.e. TpeThe OCHOBaHHe 60JibllIHHCTBa KO,llOHOB HMeeT onpe,lleJieHHYIO CTeTieHb CB060,llbl rrpH o6pa30BaHHH naphl c coOTBeTCTBYIOIUHM aHTHKO,llOHOM H KaK 6bI «KaqaeTCH» ).

TaK, ttanpHMep, O,llHa H3 aprnHHHOBbIX TPHK HMeeT aHTHKOL(OH 5'-I-C-G-3',KOTOpbIH MO)l(eT y3HaBaTh 3 pa3HhIX apmHHHOBbIX KOJJ:Otta:

AHTHKO,[IOH: (3')-G-C-1-(5') (3')-G-C-1- (3')-G-C-1-

Ko,n;OHhI: (5')-C-G-A-(3') (5')-C-G-U- (5')-C-G-C-

AMmloaD,HJI-TPHK CHHTeTa3bl

(a~muoauHJI-TPHK JIHra3bI)

B UHT030Jie KJieTOK 20 pa3JiffiHhIX aMHHOKHCJIOT rrpHCOeL(HHHIOTCH a-Kap60KCHJibHOH rpyrrrroi1 K 3'-rn.n.poKCHJihHOMy aKUeITTOpHOMy KOHuy COOTBeTCTBYJOIUHX TPHK c o6pa3oBaHHeM CJIO)l(Jf03qmpttoi1 CBH3H. 3T11 peaKUHH KaTaJIH3HpyeT ceMeHCTBO cl>epMeHTOB, HOC51IUee Ha3BaHHe aMJrnoauHJI-TPHK CHHTeTa3 (aa-TPHK-CHHTeTa3). Ka)l(,llblH qneH 3TOro ceMei1cTBa y3ttaeT TOJihKO o,11tty orrpe.n.eneHHYJO aMHHOKHCJIOTY 11 re TPHK, KOTOphie crroco6HhI CBH3bIBaTbCH c 3TOH aMHHOKHCJIOTOM. I-fa 3TOfO CJie- ,11yeT, 'ITOB rpyrrny TPHK CHHTeTa3 BXO.D.HT 20 paJJIJ1qHhIX cl>epMeHTOB. 0HH OCYllJeCTBJI5IIOT aKTHBaUHIO aMHHOKllCJIOT B 2 CTaL(HH: Ha rrepBOM CTaL(HH aMHHOKHCJIOTa rrp11coen11H5IeTCH K cl>epMeHry H pearnpyeT c AT<t> c o6paJOBaHHeM 6orarnro 3tteprneti rrpoMe)l(yroqttoro coe,llHHeHHH - aMHHOaUJUI-

AM<I>. Ha BTopoi1 CTaJJ:HH aM11ttoauHJibHhIM ocTaTOK aMHHOaUHJia,lleHHJiaTa, OCTaBaHCb CB513aHHhlM c cl>epMeHTOM, B3aHMO,lleticrnyeT c MOJieKYJIOM COOTBeTCTBYJOIUeti TPHK c o6paJoBaHHeM aMHHOaUJUITPHK (PHC. 4-36).

AT4>

aa-rPHK-ClllHTeT838 (E)

0

O-

Ii

I

NH -CH- c~o-P= 0

2 I I

R

0

 

 

 

I

 

 

 

~HO OH

xE

rPHK

 

 

 

 

 

 

AM4>

 

 

 

E

 

 

 

 

 

AO=OH

 

 

O

0 ~C-CH-NH2

 

HC

II

I

 

2 1

0

R

 

0

I

-0- P=O

I

0

I

c

¥I'

P111c. 4-36. 06paaoaaHMe aMMHoa1.1111n-TPHK. AMlllHOK111cnora ssa111MO.Qe~crsyer c AT¢ 111 aKT111s111pyercs:i, o6pasys:i aM111HoaL1111n- a.QeH111nar, KOTOpbl~, He OCB060)l(.Qa5'1Cb1113 CB5'13111 c Q>epMeHTOM (E), OT.Qaer aKTlllBlllpoB8HHYIO aMlllHOKlllCnory rPHK c o6paaosaHllleM aMlllHOal..lllln-rPHK (aa-rPHK).

CyMMapHYJO peaKu1110, KaTaJIH3HpyeMYJO aMH- HoauJUI-TPHK CHHTeTa3aMH B rrpHCyTCTBHH HOHOB Mg2+, MO)l(HO rrpe,llCTaBHTh cneJJ:YIOIUHM o6pa3oM:

AM11ttoKHcnorn +TPHK + AT<t> ~ aMHHoauHJI-

TPHK + AM<t> +PP.

I

173

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