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підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

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CTerreHh HOHttJauuu <l:>YHKIIHOHaJihHhIX rpyrrrr 3THX PMHKaJIOB JaBHCHT OT pH cpeJl.hI. I1pw pH pacrnopa oKono 7 Bee HOHoreHHhie rpyrrIThI 6enKa HaXOJl,HTCH B HOHH3HPOBaHHOM COCTOHHHH. B KHCJIOH cpe)l,e ysenwqeHHe KOHI1eHTpa1IHH ITPOTOHOB (H+) rrpHBOJl,HT K ITOJl,aBJieHHIO Jl,HCCOI1Ma1IMH Kap60KCHJihHhlX rpyrrIT H yMeHhIIIeHHIO OTpHIIaTeJibHOfO 3apHJ],a 6eJIKOB: -coo-+ H+ ~ -COOH.

B menoqHoM: cpe)l,e CBH3hIBaHtte tt36hnKa OH- c rrpoToHaMtt, o6pa3YJOIIIHMMCH ITPH Jl.HCco1111a111111 NH3+ c o6paJoBaHtteM BOJl.hI, ITPHBOJl.HT K yMeHhIIIeHMIO ITOJIO)J(HfeJihHOro 3apH,Ua 6eJIKOB:

-NH/+OH-~ -NH2 +Hp.

3HaqeHMe pH, IIPH KOTOPOM 6eJIOK ITp1106peTaeT CYMMapHhIH HyJieBoil 3apHJ1., Ha3hIBaIOT «H303JieKTp1111ecKaH TO'IKa» H 0603ttaqa10T KaK pl. B

M303JieKTpttqecKOll ToqKe KOJilfqecTBO ITOJIO)l(H- TeJlbHO H oTpttuaTeJihHO 3apH)l(eHHhIX rpyITIT 6enKa 0)1,HHaKOBO, T.e. 6eJIOK HaXOJl,MTCH B M303JleKTpwqecKOM COCTOHHHH.

TaK KaK 6onhIIIMHCTBO 6eJIKOB B KJieTKe MMeeT B CBOeM COCTaBe 60JlbIIIe aHMOHOreHHhIX rpyrrrr (-Coo-), TO H303JieKTpMqecKaH rnqKa 3TMX 6eJIKOB Jle)l(MT B cna60KHCJIOH cpe)l,e. l1303JieKTPMqecKaH TOqKa 6eJIKOB, B COCTaBe KOTOPhIX rrpeo6Jia)],aIOT KaTHOHOreHHhie rpyIIIThl, HaXOJl,HTCH B menoqHoil cpeJl.e. Han6onee HPKMH npnMep rnKMX BHYTPMKJieTOqHhIX 6eJIKOB, COJ],ep)l(aIIIHX MHOro aprMHHHa M JlM3MHa, - rHCTOHhl, BXO)l,H- IIIHe B COCTaB xpoMaTHHa.

EeJIKH, 11Me10m11e cyMMapHhIH ITOJIO)l(ttTenhHhIH MJIM OTPHIIaTeJihHhlll 3apHJ1., nyqllle paCTBOPHMhl, qeM 6eJIKH, HaxoaHIIIMecH B M303JieKTpttqecKoil ToqKe. CyMMapHhIH 3apHJ1. yBennqwBaeT KOJIHqeCTBO Jl,MJTOJieM: BO)l,hl, CIIOC06HblX CBH3hIBaTbCH c 6eJIKOBOH MOJieKyJIOH, H ITperrHTCTByeT KOHTaKTY 0)1,HOMMeHHO 3apH)l(eHHhIX MOJleKyJI, B pe3yJihTaTe pacTBOpMMOCTh 6eJIKOB yBeJiwq11BaeTCH. 3apH)l(eHHhie 6eJIKM MoryT Jl,BMraTbCH B 3JleKTpnqecKOM none: aHMOHHhie 6eJIKH, HMeIOIIIMe OTPMIIaTeJihHhlll 3apHJ1., 6YJl.YT Jl,BMraThCH K IIOJIO)l(MTeJihHO 3apH)l(eHHOMY aHOJl.Y ( + ), a KaTMOHHbie 6eJIKM - K OTPMIIaTeJihHO 3apH)l(eHHOMY KaTOJl.Y (-). EeJIKH, HaXOJl.HIIIMecH B M303JieKTpwqecKoM COCTOHHHH, He rrepeMemaIOTCH B 3JieKTpMqecKOM ITOJie.

4. Coomnowenue noAJ1pHbLX u nenoAJ1pHbLX zpynn na no6epxnocmu namu6Hbt.x MOAeKy.11 iJeAK06

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68

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5. Pacm6op11.Mocmb 6€.!IK06

PacrnopHMOCTb 6eJIKOB B BOLI.e JaBHCHT OT Bcex rrepeqncJieHHbIX BbIWe CBOHCTB 6eJIKOB: ¢opMbI, MOJieK)'JUIPHOMMaCCbI, BeJIJfqJfHbI 3ap5ILl,a, COOTHOWeHIDI IIOJUipHbIX H HeIIOilllpHbIX <l>YHKUHOHaJibHbIX rpyrrrr Ha rroBepXHOCTH 6eJIKa. KpoMe 3Toro, pacrnopHMOCTb 6eJIKa orrpe,ll,eJUieTCH COCTaBOM paCTBOpH-

TeJIH, T.e. HaJinqHeM B pacrnope npyrHX pacrnopeHHhlX Bemecrn. HarrpttMep, HeKOTOpbie 6eJIKH nerqe paCTBOpHIOTCH B CJia6oM COJieBOM pacrnope, IJ:eM B Ll,HCTHJIJIHpOBaHHOM BOLl,e. c L1,pyro:i1 CTOpOHbI, YBeJIJfqeHJfe KOHlleHTPaIJ,HH HeHTpaJibHblX COJietl: MO)l(eT crroco6crnoBaTh BhIIrnLI.eHHIO orrpeneneHHhIX 6eJIKOB B OCa,ll,OK. )leHaTYPHPYIOillHe areHTbl, rrpHCYTCTByIOlllHe B pacrnope, TaK)l(e CHI1)1(alOT paCTBOpHMOCTh 6eJIKOB.

Ii. METO,llhI Bhl,llEJIEHIDI tt oqttcTKtt BEJIKOB

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pa3L1,eJiem1e CMeCH 6eJIKOB Ha HHLl,HBHLl,yaJibHbJe 6eJIKH.

1.MemoiJbt p03pywenua mKaneu

u 3KCmpaK~UU 0€./IK06

)lJIH pa3pywemrn 611onorttqecKoro MaTepttana HCIIOJib3yIOT MeTOLl,bJ: fOMoreHH3aUHH TKaHH, MeTOLi, rrorrepeMeHHOfO 3aMOpa)l(HBaHIDI H orraHBaHIDI, a TaK)l(e o6pa6oTKY KJieToK YJibTpa3BYKOM.

f OMOreHH3aIJ,HB 6uonol'lflleCKOl'OMaTepuMa

TKaHb, HaxOMIUYIOCH B 6y¢epHOM pacrnope corrpeneJieHtthIM 3HaqeHJieM pH H KOHUeHTPaUHe:i1 coJietl:, IIOMeruaIOT B CTeKJIHHHbIB COCYLI. (roMOfeHH3aTOp) c rrecTHKOM. Bpama101.ll,11i1:c51 rrecTHK 113MeJih1:1:aeT H pacTnpaeT TKaHb o rrpttrepTu1e cTeHKH cocyna.

MeTo,11. JaMopruKHeaHHs u orraueaum1 TKauu

B pe3yJihTaTe rrorrepeMeHHoro 3aMOpa)l(HBamrn H OTTaHBaHH51 o6pa3ylOIUHeCH KpHCTaJIJibl Jlb,ll,a pa3pyrnaIOT o6oJIOlJ:KH KJieTOK.

IlocJie paJpyrneHHH TKaHH HepaCTBOpHMbie qaCTH OCa)l(Ll,alOT ueHTptt¢yrnpOBaHHeM. IlocJie,ll,ylOIUee ueHTPH<l>yrnpoBaHHe fOMOfeHaTa c pa3HOM CKOpOCTblO II03BOJIHeT IIOJiyqHTb OT,11.eJihHhie ¢paKUHH, COLl,ep)l(aIUHe KJieToqHbie 51L1,pa, MHTOXOHLI.PHH H ,11.pyrtte opratteJIJihI, a TaK)f(e Ha,11.oca,11.01:1:HyIO )f(Jf,11.- KOCTh, B KOTopotl: HaxOMTCH pacrnopHMbie 6eJIKH UHT030JIH KJieTKH. l1cKOMblll 6eJIOK 6y,11.eT co,11.ep- )f(aTbCH B 0,11.HOM H3 3THX ¢paKLI,HM.

3KCTpaKIJ,HB 6MKOB, CBB3aHHblX c MeM6pauaMu, u pa3pymeHHe onuroMepHhIX 6enKoB ua np0ToMeph1

ECJIH HCKOMbill 6eJIOK rrpOlJ:HO CBH3aH c KaKH- MM-JIH6o CTPYKTYPaMH KJieTKH, ero Heo6xo,11.HMO nepeBeCTH B pacrnop. TaK, LI.JIH pa3pyrneHHH rnnpo¢o6ttbIX B3aHM0,11.eilcTBMll Me)l(,11.y 6eJIKaMH 11 JIMIIHLl,aMH MeM6paH B pacrnop ,11.06aBJIHIOT Ll,eTepreHThI; qame Bcero ncnoJib3yIOT TPHTOH X-100 HJIH

,ll.0,11.eUHJICYJib<l>aT HaTpHH.

MexaHM3M ,11.e:HcrnIDI ,11.eTepreHTOB orrttcaH B pa3-

LI.eJie «lleHawpaunH 6eJIKOB» (CM. pttc. Ilptt

,11.eilCTBMH ,11.eTepreHTOB o6bilJ:HO pa3pyu.ra10TCH 11 fHL1,po¢06Hbie B3aHMOLl,etl:CTBIDI Me)l(,11.y npOTOMepaMH B OJIHfOMepHbIX 6eJIKax.

Y,11.Meuue u3 pacTBopa ue6enKOBhIX eem,ecTe

HYKJieHHOBbie KHCJIOThI, JIHIIHLI.hI H ,11.pyrne He6eJIKOBhie Bewecrna MO)KHO YLI.aJIHTh H3 pacTBopa, HCIIOJib3YH HX oco6eHHbie <l>tt3HKO-XHMHqecKtt:e cBotl:cTBa. TaK, mmtt,11.hI nerKo y,11.aJIHIOTCH H3 pacrnopa ,11.06aBJieH11eM oprattw-1ecKHx pacrnopttTeJie:H, HarrpHMep aueTOHa. 0,11.HaKO B03,ll,eilCTBHe ,11.0JI)l(HO 6bITb KpaTKOBpeMeHHbIM, TaK KaK aueTOH BhBbIBaeT ,11.eHaTYPaun10 HeKOTOpbIX 6eJIKOB. HYKneHHOBhie KHCJIOTbl OCa)KLl,alOT LI.06aBJieHHeM B paCTBOP CTperrTOMHUHtta.

2. MemoiJbt O'lucmKu fieAK06

Hatt6oJiee TPYLI.OeMKHtl: 3Tan nonyqettIDI HHLI.H- BHLI.YaJihHhIX 6eJIKOB - HX OlJ:HCTKa OT LI.PYfHX 6eJIKOB, HaXOLl,HIUHXC5I B paCTBOpe, IIOJiyqeHHOM M3 ,11.aHHOM TKaHH. qaCTO H3y1:1:aeMblM 6eJIOK rrptt:CYTCTByeT B tte60JibWHX KOJIHqecTBaX, COCTaBJUIIOIUMX LI.OJIH rrpoueHrn OT Bcex 6eJIKOB pacrnopa.

TaK KaK 6eJIKH 06Jia,11.a10T Kott¢opMaUHOHttoi1 Jia6HJibHOCTblO, npH pa6oTe c 6eJIKaMH CJie,ll,yeT H36eraTb ,IJ;eHaTypttpyIOIUHX B03,IJ;eHCTBMM, II03TO-

MY BbILI.eJietttte M OlJ:HCTKa 6eJIKOB npottcxoMT nptt: HH3KHX TeMnepawpax.

69

1-56).
10-12

Ha nepBhIX crn,n.1ui:x oqttcTKH 6eJIKOB uenecoo6pa3HO HCTIOJih30BaTh MeTO,Uhl, yqJ1Tb1BaI01UHe KaKYIQ-Jitt6o xapaKTepttyIO oco6eHHOCTh ,UaHHOfO 6enKa, HanpttMep TepMocrn6HJihHOCTh HJIH ycTOllqJIBOCTh B KHCJihIX paCTBOpax. TiepBhIMH MeTO,UaMH oqttcTKH Heo6xo,UHMO y,UaJIHTh tt3 pacTBopa OCHOBHYIO Macey 6aJIJiaCTHhlX 6eJIKOB, KOTOphie 3HaqttTeJihHO OTJIJlqaIOTCH OT BhI,UeJIHeMOfO 6eJIKa ¢tt3HKO-XHMJlqeCKHMH CBOllCTBaMH. Bnocne,11,CTBHH npttMeHHIOT Bee 6oJiee TOHKHe MeTO,Uhl oqHCTKH 6eJIKa.

()qucTKa 6eJTKOB H36HpaT~bHOH .ueuayYpaIU1eH

oOJihWHHCTBO 6eJIKOB ,UeHarypttpyeT H Bhma- ,11,aeT B OCa,UOK )')Ke nptt KpaTKOBpeMeHHOM HarpeBaHHH pacTBopa ,UO 50-70 °C HJIH TIO,UKHCJieHHH pacTBopa ,UO pH 5. Ecm1 BhI,UeJIHeMhlll 6enoK BhI,Uep:)!<HBaeT 3TH YCJIOBHH, TO c TIOMOIUhlO H36ttpaTeJibHOll .ueHaTypautttt MO)l(HO y,UaJIHTh 6onhIIIYIO qacTh TIOCTOPOHHHX 6eJIKOB, OTcPHJihTpoBaB BhITiaBIIIHe B OCa,UOK 6eJIKH, HJIH OCa,UHTh HX ueHTptt¢ympOBaHHeM.

BhlCMHBauue

MeTO,U oqttcTKH 6enKOB, ocHOBaHHhill Ha pa3JIJlqHHX B HX pacTBOpHMOCTH nptt pa3HOll KOHUeH - Tpautttt com1 s pacTBope. Contt 1UeJioqHh1x tt 1Ue- JIOqff0-3eMeJihHhIX MeTaJIJIOB Bhl3bIBaIOT o6paTHMOe OCa)l(,UeHHe 6eJIKOB, T.e. TIOCJie HX y,11,aJieHHH 6eJIKH BHOBh nptto6peTaIOT crroco6HOCTh paCTBOPHThCH, coxpaHIDI nptt 3TOM CBOH HaTHBHhie CBOllCTBa.

lfaIUe Bcero ,UJIH pa3,UeJieHHH 6eJIKOB MeTO,UOM BhICaJIHBaHHH HCIIOJih3YIQT pa3Hhle KOHUeHTpaUHH COJiell cynh¢arn aMMOHHH - (NH4)2S04. qeM Bhlllle paCTBOpHMOCTh 6eJIKa, TeM 60JihlliaH KOHUeHTpaUHH COJIH Heo6X0,11,HMa ,11,JIH ero BhICaJIHBaHHH.

fML-tl>HJibTpa.u,ml, ~ MeTO,U MMeKY.IUIJJHblX CHT

,UJuI pa3,11,eneHHH 6eJIKOB qaCTO HCllOJih3YIOT xpoMaTorpa¢ttqecKHe MeTO,Uhl, OCHOBaHHhie Ha pacrrpe,11,eJieHHH Be1UeCTB Me)l(,ll,y ,UBYMH ¢a3aMH, 0,UHa H3 KOTOphIX 110,11,BH)l(HaH, a .upyraH HellO,ll,BH)l(HaH.

B ocHOBY xpoMaTOrpa$ttqeCKHX MeTO,UOB 110JIO)l(e- HhI pa3Hhle llPHHUHllhI: reJib-cPHJibTpauttH, HOHHOfO o6MeHa, a.ucop6utttt, 6ttonorttqecKoro cpo,U- CTBa.

MeTo,11, pa3,11,enettttH 6enKoB c noMOIUhIO renhcPHJihTpauttOHHOll xpoMaTOrpa¢tttt OCHOBaH Ha TOM, qTo BeIUeCTBa, OTJIJ1qaI01UHeCH MOJieKYJIHpHOll Maccoll, llO-pa3HOMY pacnpe,UeJIHIOTCH Me)l(,Uy

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Ha xpoMaTorpa$ttqecKyIO KOJIOHKY' BhIMhIBaIOT

(3JIIOHPYIOT), npo11yCKaH qepe3 KOJIOHKY paCTBOpttTeJih. BMecTe c ¢poHTOM pacTBopHTeJIH .UB~­

CH H CaMbie KpynHhie MOJieKYJihl.

Bonee MeJIKHe MOJieKYJihI .UHcPcP)'H,UHPYIOTBHYTph rpattyn ce¢a.ueKca H Ha HeKOTopoe BpeMH nona.uaIOT B HeTIO,UBJl)l(HYIQ ¢a3y'B pe3yJihTaTe qero HX ,UBH)l(eHtte 3a,Uep)l(HBaeTCH. BeJIJlqttHa rrop orrpe,UeJIHeT pa3Mep MOJieKyn, cnoco6HhIX rrpOHHKaTh BHYTPh rpaHyn (pttc. 1-55).

TaK KaK reneBaH cTpyi<rypa ce¢a.ueKca nerKo ,Ue- ¢opMttpyeTCH no.u ,UaBJieHtteM, rentt CTantt 3aMeHHTh 6onee )l(eCTKHMH MaTpttUaMH (ce¢aKTHJI, TOllonepn), npe,11,cTaBJIHIOIUHMH c¢epttqeCKHe rpattynhI c pa3HhIMH pa3MepaMtt nop. Bh16op pa3MepoB nop B rpaHynax 3aBttCHT OT uenell xpoMaTOrpa¢tttt ( o ,Upymx xpoMaTOrpa¢ttqecKHX MeTO.uax 6y,UeT cKa-

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