підручники. біохімія / Uchebnik_po_Biokhimii_E_S_Severin

CTeneHh roMoJiomH Me)l(,.ll,y 3THMH no,U- KJiaccaMH oqeHh BhICOKa (OKOJIO

lgG He TOJibKO 3¢¢eKTHBHO CB.H3hIBaIOT H HHaKTHBHPYIOT 'l)')KepOil.HbieMOJieKYJThI H KJieTKH, IIOITaBume B opraHH3M, HO TaIOKe o6JierqaIOT HX ,llaJihHeti:- rnee yttWITO::>KeHtte. Kott¢opMaI1HOHHb1e H3MeHeHHH B <<XBOCTOBOH» o6JiaCTH lgG IIOCJie ero B3aHMO,lleHCTBH.H c aHTRreHOM IIPHBO,ll.HT K CB.H3bIBaHHIO H aKTHBaUHH 6eJIKOB CHCTeMhl KOMITJieMeHTa. KpoMe rnro, C-KOHUeBa.H o6JiaCTh lgG crroco6Ha B3aHMO- )l.ettCTBOBaTh co cneutt¢ttqecKHMH peueITTOpaMH MaKpo¢aroB H HeHTpOcPHJIOB, ~o rrpHBO)l.HT K ct>arouHT03Y KOMITJieKCOB aHTHreH-aHTHTeJIO H pa3pyrneHHIO HX B ct>aroCOMax (PHC.

IgG - e)l.HHcTBeHHhIH KJiacc aHTHTeJI, crroco6Hhitt rrpOHHKaTh qepe3 ITJiaueHTapHbIH 6apbep H 06ecnetI1rnaTh BHYTPHYTp06HyIO 3aIUHTY ITJIO)l.a OT HHcPeKUHH.

lIMMYHOrJ106yJIHHb1 A. OcttOBHOtt KJiacc aHTH-

TeJI, rrpHCYTCTBYIOIUHfi B ceKpernx )J(eJie3 opraHH3Ma ( CJIIOHhI' MOJIOKa, m1meBapHTeJibHOfO COKa, ceKpeTOB )l.hIXaTeJihHhIX flYTett). B ChIBopoTKe KPOBH ero co)l.ep::>KaHtte He npeBhirnaeT 10-15% OT o6mero KOJitttiecrna HMMyttorno6yJIHHOB. MoHOMepHa.H ct>opMa no CTpOeHmo HaIIOMHHaeT lgG. 0)].HaKO B ceKpernx lgA HaXO)l.HTC.H B OCHOBHOM. B ct>opMe )l.HMepa, f,lle MOHOMepbl coe)l,HHeHhI )l,OIIOJIHHTeJibHOH rreIITH)].HOll uerrhIO J (pHC. 1-49).

Ha 6a3aJThHOtt rroBepXHOCTH 3IIHTeJIHaJihHhlX KJieTOK ,llHMep lgA crreuwct>WieCKH B3aHMO)l.ettcTByeT c 6eJIKaMH KJieTO'lHOH IlOBepXHOCTH, Ha3bIBaeMbIMH ceKpeTOpHbIM KOMIIOHeHTOM. 06pa3YJOIUMHCH KOMIlJieKC nocpe)l.CTBOM 3H)l.OUHT03a IIOfJIOIUaeTCH BHyTpb KJieTKH H nepeMemaeTC.H K aITHKaJihHOH qaCTH. 3)].eCb KOMIIJieKC IIO)l,BepraeTC.H )l,eHCTBHIO npoTeomuHtieCKHX ct>epMeHTOB, Jf CB060,llHhIH )l,HMep

BhICB060)l(,.ll,aeTC.H BO BHeKJieTOqHoe npocTpaHCTBO (pttc.

06pa3yIOIUMHCH nptt B3aHMO,llettCTBHH lgA c aHTHfeHOM KOMIIJieKC He B3attMO)l,eHCTBYeT c 6eJIKaMH

CHCTeMbI KOMIIJieMeHTa Jf ct>arOUHTHPYIOIUHMH KJieT· KaMJf, HO rrpelliITCTByeT rrpHKpeIIJieHmo aHTMreHOB K IIOBepXHOCTH 3IIMTeJIHaJibHbIX KJieTOK Jf npOHMKHOBeHHIO HX B oprattH3M.

Co)l.ep::>KaHHe 3Toro KJiacca HMMyttOfJIOOYJIHHOB B KpoBH KpatiHe MaJIO. MOHOMepbI, HO, B OTmNHe OT IgG, HX TH::>KeJThie uenH e CO,llep::>KaT He 3, a 4 KOHCTaHTHbIX ,llOMeHa. Ilocne CHHTe3a H ceKpeUHH B KPOBb B-JIHMcPoUHTaMH lgE

MOHOMep

PMC. 1-49. C'rpoeHMeAMMePHoM Monei<yni:.1 MMMyHorno6ynMH8 A.

CD 0

0 Ill

:c 1-

:T ()

0 :c

I- tll

CD C.

!2 b

CD 0

:c c.

Ill c:

lgA

npOTOKa

>1<ene3bl

6HOJIOrHqecKH aKTHBHhIX BemecTB ( cepoTomrna, rHcTaMHHa), xpaH51IUHXC51 B ceKpeTopHhIX ny3hlphKax. Bhl6poc 3THX BemecTB B 3HaqttTCJihHO:i1 Mepe OTBCTCTBCHCH 3a pa3BHTHe BOCIIaJIHTCJihHOH peaKUHH, a TaIOKe TaKHX aJIJiepr11qecKHX peaKUHW, KaK 6pOHXHaJihHa51 aCTMa, KparrHBHHUa, CCHHa51 JIHxopaaKa. YBeJIHqeHHe KOJIHqecTBa lgE MO)!(eT rrpe)llIICCTBOBaTh pa3BHTHIO aJIJieprnqecKHX peaKUHif.•

PMc.1-50. TpaHcnopt MMMyHorno6yn111Hoe A 'Llepeaan111ten111anbHb1e KneTKM B npoTOKM )l(efle3.

CB513hIBaIOTC51 CBOHMH C-KOHUeBhIMH yqacTKaMH c COOTBeTCTBYJOIUHMH peuemopaMH Ha IIOBepXHOCTH Tyq:HhIX KJieTOK w 6a30QJJUIOB. B pe3yJihTaTe OHM cTaHOB51TC51 peuerrropaMH aHTHreHOB Ha IIOBepXHOCTH

.11,aHHhIX KJieTOK (pwc.

IlOCJie IIpllCOe)lttHeHH51 aHTHreHa XOT51 6bl K LJ:BYM aHTttreHCB513bIBaIOIUHM yqacTKaM LJ:Byx coCe)lHHX IgE KJieTKa rroJiyqaeT cwrHaJI K ceKpeuHw

• '.

AHrnreH

4. Ce.Meiicmno T-K.llemo'tHblX anmuzenpacno:Jnaww,ux peu,enmopon

EcJIH aHTHTeJia, Bhipa6aThrnaeMhre B-JIHMcPOUH-

TaMH, CB513hIBaIOT aHTHrCHhI B )l(ll,IU(OCT51X opraHH3Ma (TaK Ha3hIBaeMbIB ryMopaJibHhIM HMMyttHTeT), TO T-JIHMcPOUMThI B3attMoaewcTBYJOT c aHTttreHaMw Ha IIOBepxHOCTH 3apa)!(eHHbIX BllpycaMll lI ll3MeHeHHhIX B pe3yJihTaTe orryxoJieBoif TPaHc¢opMaUHH co6crneHHhIX KJICTOK opraHH3Ma (KJieToqHhIM lIMMYHMTeT). T - JIHMcPOUMThI y3HaroT aHTttreHhI TOJihKO B KOMrrJieKce c MOJICKYJiaMtt MHC 11 KJiacca, TaK)!(e

rrpHCYTCTBYJOIUHMll Ha KJieToqHotf IIOBepXHOCTH.

reTepoawMephI, T.e. cocT051T tt3 a- H ~-uerreif. Kmr<LJ:a51 uenh ttMeeT )lBa

HMMYHOrno6ymrnorrono6HhIX Ll.OMeHa: sap11a6eJibHhlli

(V) 11 KOHCTaHTHhIH (C) (p11c. 1-52). C-KOHUeBbie yqaCTKH KIDl<,LJ.OH u:eITH BCTpOeHhI B IlJUl3MaTWieCKYJO MeM6patty. E.r\11HCTBeHHbIH aHT11reHCBH3bIBaIOIIl,11H yqacroK pacrrOJiaraeTCH Me)!(}l)'Ll.BYMH rop11a6eJibHhIMH .!J,OMeHaM11 Va 11 vp. KOJIH'leCTBOpeu:errropoB T-JI11McpoUHTOB c pa.3HhIMl1 am11reHCBH3hIBaIOrn,11MH yqaCTKaMl1 corrocraBHMO c pa.3H006pa.311eM 11MMYHOrJI06yJIHHOB.

CeMeH:crso 6eJDCoB masuoro KOMIIJieKca rucTocosMecTHMOCTH

BeJIKH rnaBHoro KOMilJieKca r11cTOCOBMecT11MocT11 6bIJil1 OTKPbITbl rrp11 113y<IeHl111 BOilpOCOB BttyTp11B11.!J,OBOH rrepecanKH TKatteil:, OTKYLl.a 11 rrpo1130IIIJIO 11x Ha3BaH11e. Hx Ha3bIBaIOT TaJOKe 6eJIKaM11

MHC (CM. BhIIIIe), 11J111 6eJIKaM11 HLA (OT aHrn.

human lymphocyte antigen - qeJI0BeqecK11e Jil1M-

<t>ou:11TapHh1e aHT11reHbI), TaK KaK BrrepBbie OHl1 6hIJil1 o6HapyxeHbl Ha JI11M<t>ou:11rnx qeJioBeKa.

CymecTByeT LI.Ba ocHoBHhIX KJiacca MOJieKyJI MHC: I 11 II. MoJieKYJibI MHC KJiacca I pacrroJio- )l(eHbI Ha ITOBepXHOCTl1 rrpaKT11qecK11 Bcex KJieTOK opraHl13Ma qeJioBeKa, a 6eJIKl1 MHC KJiacca II

TOJibKO Ha orrpe.!1,eJieHHbIX KJieTKax 11MMYHHOH Cl1-

CTeMbl, Ha3hlBaeMhlX aHTl1feHrrpe.!1,CTaBJIHIOIIl,11Ml1 KJieTKaMl1. K Hl1M, B rrepByIO oqepe.!1,b, OTHOCHT MaKpo¢>ar11 11 B-JI11M<t>ou:11ThI, KOHTaKT11poBaBIIIMe c aHTl1feHOM.

MoJJeKYJihI MHC KJiacca I - reTepon11Mepb1.

0Hl1 HMeIOT OLl.HY IlOJIMITeITT11.!1,HYIO a-u:errb, CBH3aHHYJO HeKOBaJieHTHbIMl1 CBH3HMl1 c He60JiblIIMM BHeKJieToqHhIM 6eJIKOM p2-M11Kporno6yJI11HoM. Ilo1rnrrerrTl1Ll.HaH a-u:errh 11MeeT Tp11 BHeKJieToqHhIX rno6yJIHpHbIX .!J,OMeHa (al'al, a3), TpaHCMeM6paHHblH yqacTOK 11 Kap60KCl1JlbHbIH KOHeu:, JIOKaJil1-

30BaHHbIH B U:11TOilJia3Me (p11c. a 3-)lo- MeH 11 P2-M11Kporno6y1rnH 11MeIOT KoH¢>opMau:11IO, HarroM11HaIOmYIO CTPYKTYPY 11MMyttorno6yJI11HoB. )loMeHbI a 1 11 a 2 conep)l(aT Bap11a6eJihHbie yqacTKH, crroco6Hble CBH3bIBaTh «pa3BepttyTbIH» aHTl1reH (qarn,e Bcero rrerrTl1.!J,HhIH ¢>parMeHT qyxeponHoro 6eJIKa), pacrroJIO)l(eHHbIH Ha ITOBepXHOCTM KJieTOK.

MOJieK)'JlhIMHC KJiacca D - TaIOKe rerepo.!1,11Mepb1. OHH cocrom 113 .!J,Byx rroJ111I1eI1Tl1.!J,HhlX u:erreil: - a 11 p, HMeIOIIl,HX no OLl.HOMY KOHCepBaTHBHOMY 11MMYHOfJI06yJ111HI10.!J,06HOMY .!J,OMetty 11 no OLl.HOMY Bap11a6eJihHOMY Ll.OMeHy Ha N-KOHU:eBhIX yqacTKax. CBH3bIBaHHe aHTHreHOB rrpo11cxo.!J,11T B o6JiaCTH Bap11a6eJibHhlX .!J,OMeHOB a- 11 P-u:erreil: (p11c. 1-53, B).

MeM6paHa

PMc. 1-52. CTpoeHMe pei.iemopa T-nMM$01.1MTOB.

PMC. 1-53. CTpoeHMe 6enKOB rnaeHoro KOMMEU<CB rMCTOCOBMecTMMOCTM: MHC Knacca I (A) MMHC Knacca II (6).

qy)l(epo.!1,Hbie 6eJIK11 B KJieTKe qeJioBeKa (Harrp11Mep, 6eJIKH B11pycHbIX qacT11u:), B Jil130COMax rro;11,- BepraIOTCH orpaHH'leHHOMYrrpoTeOJil13Y, 11 He60JibIII11e ¢>parMeHTbI 3TMX 6eJIKOB BMeCTe c 6eJIKaM11 MHC KJiacca l 11J111 II 3KcnoH11pyIOTCH Ha rr0BepXHOCT11 KJieToqHOH MeM6paHbI.

KoMilJieKChI rreI1T11.!J,-6eJIOK MHC:y3HaIOTCH pe- u:emopaM11 T-JI11M<t>ou:HT0B. B pe3yJihTare rrpo11cXOL1.HT crreu:11¢>11'1eCKOeB3al1MO)J,eHCTBl1e (p11c. aKTMBaU:MH T-JI11M¢>ou:11rn 11 pa3B11T11e MMMYHHOH peaKU1111. TaK, B3a11Mo;11,ei1cTB11e u:11TOTOKc11qecKoro T-JI11M<t>ou:HTa c KOMilJieKCOM am11reH-MHC I

Ha rroBepXHocT11 3apa)l(eHHOH B11pycoM KJieTKH rrp11BOL1.HT K BhlCB060)K)J.eHl1IO J111M<l>OUHTOM crreu:11aJib-

KneTKa-MlllWeHb,

Harrp11Mep, 1111TOxpoM c - MHTOXOHJJ.pH.aJibHbIH 6eJIOK, yqacrnyIOIUHH B 611.oJiormecKOM OKHCJieHHH, rrp11cyrcrnyeT y MHornx BHJJ.OB )l(HBOTHbIX. l.J,HToxpoMbI C KYPHUhI 11 YTKH oTJIH':laIOTCHJIHlllb JJ.BYMH aMHHOKHCJIOTHhIMH OCTaTKaMH B nepBH':IHOH CTPYKType, BbIIIOJIHHIOT OJJ.H.HaKOBYIO <l>YHKIIHIO, HO TaK KaK OHH npH.Ha)J;Jle)l(aT pa3HbIM BHJJ.aM, TO HX OTHOCHT K fOMOJIOfH':IHblM6eJIKaM.

K 11306eJIKaM OTHOCHT TaK)l(e MHO)l(eCTBO 1130<l>opM CTPYKTYPHOfO 6eJIKa KOJIJiareHa (CM. pa3)1.eJI 15).

MHorne <l>epMeHThI HMeIOT HeCKOJibKO H.3o<l>OPM 11 HOCHT Ha3BaHH.e H3o<l>epMeHTOB (CM. pa3JJ.eJI 2).

T111po3111HKlllH03a

ltMTOTOKClll\.f8CK14H

T-n111Mcpo4111T

PMC. 1-54. Cne~'ft!CKoe esaMMQAeic:TBMe pewHTTopa ~o­

TOKCM'ff!CKoroT·nMMci>ol.IMT8 c KOMM8KCOM 8HTMreH-MHC I

6enKOM.

HhIX 6eJIKOB, Bbl3blBaIOIUHX IlOBpe)l()leHHe H rn6eJib 3apIDK.eHHOH KJHITKH.

E. 1130<l>YHKI.J,HOHAJibHhlE BEJIKH

lfao4>)'HKIUloHaJlhHhle6emrn - ceMell:crna 6eJI-

KOB, BhITIOJIHHIOIUHX IlO':ITHOJJ.HHaKOBYJO HJIH 6JIH3-

Kyio <l>YHKIIHIO, HO He60JibllIHe oco6eHHOCTH CTIJOeHHH H <l>YHKUHOHHPOBaHHH HeKOTOphIX ':IJieHOB 3TOfO ceMeHCTBa MOryr H.MeTb BIDKHOe <l>H3HOJIO-

fH':leCKOe3Ha<ieHHe. IlpHMep TaKHX 6eJIKOB - H30- <PopMbl reMorno611Ha 'leJioBeKa:HbA, Hb~, HbF 11 JJ.pyr11e, paccMOTIJeHHbie Bhillle. Bee OHH npeJJ.- crarurn10T co6oll: TeTIJaMepbI, HO COCTOHT H3 pa3HOfO Ha6opa IlpOTOMepOB a, j3, y, 0. reMOfJI06HHbl BbITIOJIHHIOT OJJ.H.HaKOBYIO <l>YHKIIHIO - npHCOeJJ.H- IDllOT 02 H nepeHOCHT era B TKaHH. 0JJ.HaKO KIDK- )J;blH H.3 HHX o6Jia;::i;aeT <l>YHKIIHOHaJibHbIMH oco6eHHOCTHMH. TaK, reMorno6HH F HMeeT 6oJihlllee cpOJJ.CTBO K 0 2, ':leMHbA, 11 6JiaronapH 3TOMY o6ecne':IHBaeTJJ.H<l><l>Y3HIO 02 OT HbA H.3 KpOBH MaTepH K Hbf B KPOBH IlJIOJJ.a.

lfao6eJIKH - MHO)l(eCTBeHHble <PopMbl 6eJIKa, 06HapyxttBaeMbJe B opraHH3Max OJJ.Horo BHJJ.a. EeJIKH, BbIIlOJIHHIOIUHe OJJ.H.HaKOBbie <l>YHKIIHH B opraHH3Max pa3HhIX 6H.OJIOfH':leCKHX BHJJ.OB, HOCHT

VII. <1>1113111KO-Xll1Mll1LIECKll1E CBO~CTBA 6E11KOB 111 METO.llbl

111X BblllE11EHll1SI

Ba)l(Hoe MecTo B 6HOXHMH':lecKHX11.ccJieJJ.oBaHHHX 3aHH.MaeT BhIJJ.eJieHH.e HHJJ.HBHJJ.YaJihHblX 6eJIKOB H3 opraHOB H TKaHeH. 0'lHIUeHHbieHH,llHBHJJ.YaJibHbie 6eJIKH tty)l(Hbl JJ.llil H3yqeHH.H HX nepBH.':IHOH CTPYKTYPhI, rroJiyqeHHH KpHCTaJIJIOB 6eJIKOB c ueJihIO HCCJieJJ.OBaHHH HX npOCTIJaHCTBeHHOH CTIJYKTyphl MeTOJJ.OM peHTfeHOCTpyKTypHOfO aHaJIH.3a, ycTaHOBJieHHH B3aHMOCBH3H Me)l(JJ.y nepBH.':IHOH, rrpocTpaHCTBeHHOH CTIJYKTYPOH 6eJIKa Hera <PYHKUHeH.

HeKOTOpb1e O':IHIUeHHbieHHJJ.HBHJJ.YaJihHhie 6eJIKH HCilOJib3yioT B MeJJ.HUH.He KaK JieKapcTBeHHbie rrperrapaThI, Hanp11Mep ropMOH HHCYJIHH npHMeHHIOT JJ.JIH Jie':leHHHcaxapHoro n11a6eTa, a n11meBap11TeJihHbie <PepMeHTbl IIOJJ.)l(eJiyJJ.O':IHOH )l(eJie3bl Ha3Ha':laIOTnpH HapYllleHH.H ee <l>YHKIIHH B Ka':leCTBe 3aMeCTHTeJibHOH Tepan11H. KpoMe Toro, O':IHmeHHhie <PepMeHTbI ':laCTO HCilOJib3YJOT B 611.oxHMH.':leCKHXHCCJieJJ.OBaHHHX B Ka':leCTBeXHMH':leCKHX peaKTHBOB JJ.llil orrpeneJieHHH BemecTB B 6HOJIOfH- ':leCKHX)l(HJJ.KOCTHX.

EoJihllIHHCTBO MeTOJJ.OB, 11crroJih3yeMhIX JJ.JIH O':IHCTKHHHJJ.HBHJJ.YaJihHbIX 6eJIKOB, OCHOBaHO Ha pa3JIH':IHHX HX <l>H3HKO-XHMH':leCKHX CBOHCTB, a TaKXe B03MO)l{HOCTH crreu11<PH':IHOCBH3bJBaTbCH c JIHfaHJJ.OM.

A. <l>H3HKO-XHMWIECKHE CBO:RCTBA BEJIKOB

HHJJ.HBHJJ.YaJibHhie 6eJIKH paJJIH':laIOTCH no cooHM <l>H3HKO-XJfMH':leCKHM CBOHCTBaM: <PopMe MOJieKyJI, MOJieKyJIHPHOH Macce, CYMMapHOMY 3apHJJ.Y

67

MOJieicyJihI, COOTHOIIIemuo ITOIDIPHhIX H HeITOIDlpHhIX rpyrrIT Ha ITOBepxHOCTH HaTHBHOH MOJieicyJihI 6eJIKa, paCTBOpHMOCTH 6eJIKOB, a TaIOKe CTeITeHH ycTOM:qHBOCTH K 803)],eHCTBHIO )l,eHaTYPHPYIOIIIHX areHTOB.

1. P(l3Jlu11ua iie.ttK06 no <}JopMe MOAeKYA

KaK yxe rosopHJIOCh BhIIIIe, ITO <t>opMe MOJieKYJI 6emrn Jl,eJUIT Ha rno6yJUipHhie H <Ptt6pHJIJUipHhie. f no6ym1pHbie 6eJIKH HMeIOT 6onee KOMITaKTHYJO CTpYKTYPY, HX TIIJ],po<}:>o6Hbie PMHKaJihl B 60JihllIHHCTBe CBOeM CITpHTaHbl B rHJ],po<t>o6Hoe H,Upo, H OHH 3HaqwreJibHO nyqIIIe pacTBOPHMhl B )IQf)],KOCTIDC opraHH3Ma, qeM <Ptt6pHJIJUipHhie 6eJIKH (HCKJIJOqeHMe COCTaBJUllOT MeM6paHHbie 6eJIKM).

2. P(l3JlU'IUR iie.ttK06 no MOJ1eKYAJ1PHoii Macce

EenKM - BhICOKOMoneicyJUipHhie coeJl,MHemm:, Ho MOryT CIDihHO OTJIMqaTbCH ITO MOJieicyAApHOH Macce, KOTOpaH KOJie6JieTCH OT 6000 )],0 I 000 000 ,z:i: M Bhillle. MoneKYJUIPHaH Macca 6enKa Jasncm OT KoJIMqecrna aMHHOKMCJIOTHhlX OCTaTKOB B ITOJIMIIerrTH)],HOH 11errM, a Jl,JIH OJIMrOMepHbIX 6eJIKOB - M OT KOJIMqeCTBa BXOJl.HIIIHX B Hero IIPOTOMepoB (IDIM cy6De)l,MHMII).

3. CyMMapHblii 1apaiJ iie.ttK06

EenKM MMeIOT B csoeM cocrnBe PMMKaJihI JIM3MHa, aprMHMHa, rMCTMJl,MHa, rnyTaMHHOBOll M acrraparnHOBOll KMCJIOT, co.uep)l(all!Me <l:>YHKUHOHaJihHhie rpyITIIhI, crroco6Hhie K MOHM3aIIHH (HOHoreHHhie rpyrrnb1). KpoMe Toro, Ha N- w C-KoH11ax rronwrremMJl,HhlX 11eITeH HMeIOTCH a-aMMHO- H a-Kap6oK- CMJihHaH rpyrrllhI, TaIOKe cnoco6Hbie K HOHM3aIIMM. CYMMapHhIH 3apHJ1. 6eJIKOBOH MOJieicyJihI JaBHCMT OT COOTHOIIIeHMH MOHM3MpOBaHHhIX aHHOHHhIX PMHKaJIOB f ny 11 Acrr 11 KaTHOHHhlX PMHKaJIOB JlM3, Apr w fnc.

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

Ha ITOBepXHOCTH 6oJihIIIHHCTBa BHYTPMKJieToqHhIX 6eJIKOB ITpeo6JiaJl,aIOT rroJIHpHhie PMMKaJihI, 0)1,HaKO COOTHOllleHMe rromi:pHbIX H HeIIOJIHPHhIX rpyrrrr OTJIHqHo Jl,JIH pa3HhlX MH)],HBMJl.YaJihHhIX 6eJIKOB. TaK, ITPOTOMephI OJIMroMepHhIX 6eJIKOB B 06JiacTM KOHTaKTOB Jl.Pyr c Jl.PYfOM qacTO COJ],ep)l(aT rMJl,po<}:>o6Hhie PMMKaJihI. I1oBepxHoCTM 6eJIKOB, <l:>YHKllMOHMPYIOIIIMX B COCTaBe MeM6paH MJIM npMKpeIIJIHIOIIIHeCH K HMM B ITpo11ecce <l:>YHKIIHOHMpoBaHMH, rnIOKe o6orameHhI rMJl,po<t>o6HhIMM paJl.MKanaMtt. TaKMe 6eJIKM JiyqIIIe pacTBOPMMhI B JIMITMJl,aX, qeM B BO)l,e.

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

Ilonyqemi:e HHLl,HBHLl,yaJihHblX 6eJIKOB H3 6ttoJIOfHIJ:eCKOfO MaTepnana (TKaHeil, opraHOB, KJieTOqHbIX KyJlbTYP) Tpe6yeT rrpoBe,ll,eHIDI IIOCJie,ll,O- BaTeJibHbIX orrepauttil, BKJIIOqaIOIUHX:

•L1,p06JieHHe 6HOJIOfJfqecKOfO MaTepHaJia H pa3pyrneHHe KJieTOlJ:HbIX MeM6paH;

•¢paKUHOHHpoBaHHe opraHeJIJI, COLl,ep)l(aIUHX Te HJIH HHhie 6eJIKH;

•3KCTpaKUHIO 6eJIKOB (rrepeBOLI, 11X B pacrnopeHHOe COCTOHHHe);

•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.

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

Herro,UBH)l(HOH tt 110,UBH)l(HOll ¢a3aMtt. XpoMaTor-

pa¢ttqecKaH KOJIOHKa 3aTIOJIHHeTCH rpaHyJiaMH

nopttcToro BemecTBa (ce¢a.ueKc, arapo3a H ,Up.). B CTpYKTYPe rronttcaxapH,Ua o6pa3yIOTCH nonepeqHhie CBH3H H ¢opMHPYIOTCH rpaHyJThl c «nopaMH», qepe3 KOTOphie JiefKO npOXO,UHT BO,Ua H HH3KOMOJieKYJIHpHble Be1UecTBa. B 3aBHCHMOCTH OT ycnoBHH MO)l(HO ¢opMttpoBaTh rpaHyJihI c pa3H0ll BeJIHqHHOll <<IIOp».

Heno,UBH)l(HaH ¢a3a - )l(H,ll,KOCTh BHYTPH rpa-

HYJI, B KOTOPYIO cnoco6HhI rrpoHttKaTh HH3KOMOJieKYJIHPHhie BemecTBa H 6enKH c He60JihlllOH MOJieKYJIHPHOH Maccoll. CMech 6eJIKOB, HaHeceHttyIO

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-

3aHo HH)l(e).

YJTLTpaQeHTPHtl>yntpoeauue

MeTo,U paJ,11,eneHHH TaIOKe ocHOBaH Ha pa31Itt-

qJ1H B MOJieKYnHpHhIX Maccax 6eJTKOB. CKopoCTh ce,UttMeHTaUHH Be1UeCTB B npouecce Bpa1UeHHH B ynhTpaueHTptt$yre, r.ue ueHTpo6e)l(Hoe ycKopeHtte ,UOCTttraeT I00 000-500 000 g, nponoputtoHaJihHO HX MoneKYnHpHoll Macce. Ha rroBepXHOCTh 6y¢epttoro paCTBOpa, ITOMemettHOfO B KIOBery'HaHOCHT TOHKHll CJIOll CMeCH 6eJIKOB. KloBery noMe1UaIOT B poTOp ynhTpaueHTptt¢yrtt. Ilptt Bpa1UeHHH poTO-

pa B TeqeHtte q 6onee KpynHhie MOJieKYJihI

( c 60JihllleH MOJieKYJIHpHOll Maccoll) oce,UaIOT B 6y$epHOM pacTBope c 60Jihlllell CKOPOCThlO. B pe3YJihTaTe B KIOBeTe npOHCXO,UHT paccJIOeHHe CMeCH 6eJIKOB Ha OT,UeJihHhie $paKUHH c pa3HOll MOJieKY-

JIHPHOll Maccoll (pttc. Ilocne paccnoeHHH

6eJIKOBhlX ¢paKUHll ,UHO KIOBeThl rrpoKaJihIBaIOT HrJIOll H no KaIIJUIM co6ttpaIOT CO,Uep)l(HMOe He60JihlllHMH nopUHHMH B npo6ttpKH.

KpynHb1e Monei<ynbl 6enKa BHyrpb rpaHyn He nonaAalOT

0

H1113KOM011ei<ym1pHbte 6e11K111 npoH111Ka10T e rpaHynbl ce¢a-

.QeKca

~r--+--- Bb1COKOM011eKy11S1pHb1e 6enKH npoxo.QsiT Me)f(,[ly rpaHy11aM111

PaJAeneHHe 6enKoe Ha cppaKL\"1H

6eI1KOBble

cppaKL\HH

PMc. 1·56. KioeeTa, sanonHeHHaR 6ycj>epHb1M peCTeopoM c paSA&neHHblMM 6enKOBblMM ctJpaK~MAMM.

3JieKTPocl>ope3 6e.JJ:KOB

MeTo.JJ. ocHoBaH Ha TOM, qTo rrptt orrpe.JJ.eJieH-

HOM 3Ha'IeHHHpH H HOHHOH CHJibI pacTBopa 6eJI-

KH )1.BMralOTCH B 3JieKTPM'IeCKOMIIOJie co CKOpocTblO, rrporropUHOHaJibHOH HX CyMMapHOMy 3apH.JJ.Y. EeJIKM, MMe10mtte cyMMaptthIH oTpttuaTeJihHhIH 3apH)1., )1.BHralOTCH K aHOJJ.Y (+), a IIOJIO)l(HTeJibHO 3apIDKeHHhie 6eJIKH - K KaTO.JJ.Y (-).

3JieKTpo<t>ope3 rrpOBOMT Ha pa3JIHqHblX HOCHTeJIHX: 6yMare, KpaXMaJibHOM rene, IIOJIHaKpHJiaMH)J,HOM rene H .JJ.p. B OTJIH'IHeOT 3JieKTPo<t>ope3a Ha 6yMare, f)].e CKOpOCTb )J,BH:)((eHHH 6eJIKOB rrpoIIOpUHOHaJihHa TOJibKO HX cyMMapHOMy 3apH)1.y, B IIOJIHaKpHJiaMH)J,HOM rene CKOpOCTb )J,BH)l(eHH51 6eJIKOB rrporropuHOHaJibHa HX MOJieKyJIHPHbIM MaccaM.

Pa3peillalOll.l,a51 CIIOCOOHOCTb 3JieKfPOCPopeJa B rroJIHaKpHJiaMM)J,HOM rene Bh1rne, qeM Ha 6yMare. TaK,

npH 3Jlel<Tpocpope3e 6eJIKOB CbIBOPOTKH KPOBM qeJIOBeKa Ha 6)'Mareo6Hap)l)KHBaIOT TOJlbKO 5 rJiaBHblX

cppmcu:Hit aJib6YMMHhI, a 1-rno6yfillHhI, a2-rno6y1rn- Hb1, ~-rno6yJIMHbl M y-rno6yJIHHbl (pMC. 1-57). 3JieK-

TpOcpope3 TeX )l(e 6eJIKOB B IIOJIHaKpHJiaMH,llHOM reJie

n03BOJ1HeT rronyqHTu )lO 18 pa3.JIHqfillX <l>PaKu:HM. ,lJ,JuI o6Hap~eHHH 6eJIKOBblX <l>PaKu:MH IIOJIOCKH 6)'Marn

HJIH crnn6HKH reJIH o6pa6aThIBa.IOT K})acmeneM (qaiu:e Bcero 6poMcpeHOJIOBbIM CHHHM HJIH aMH)lOBbIM qepHbIM). OKparneHHhIH KOMIIJleKc 6eJIKoB c KpacHTeJieM BbIHRJIHeT paCIIOJIO)l(eHHe paJJimHbIX <l>PaKu:MH Ha HOCHTeJie.

lfoH006MeHHaJI xpOMaTOrpa<l>m1

TaK )Ke KaK 11 3JieKTpocpope3, MeTOJ.J. ocHOBaH Ha pa3)leJieHl1M 6eJIKOB, pa3JI11qaIOll1l1XCH C)'MMapHbIM 3apH)lOM rrpM onpeJ.J.eJieHHbIX 3HaqeH11H:x pH M MOHHOH CHJihI pacrnopa. Ilp11 rrponycKaH1111 pacrnopa 6eJIKOB qepe3 xpoMaTorpacp11qecKYJO KOJIOHKY, 3aIIOJIHeHHYJO rnep)lblM nop11CTbIM 3apH)l(eHHbIM MaTep11aJIOM, qacTb 6eJIKOB 3a)lep)l(l1BaeTCH Ha HeM B pe3yJihTaTe 3JieKTpOCTaT11qecK11X B3al1MO)leHCTB11H.

B Ka'-lecrne HenOJ.J.Bl1)1(HOH cpa3bI 11cnoJib3YJOT MOH006MeHHMKM - IIOJl11MepHbie opraH11qecK11e Bemecrna, coJ.J.ep)l(am11e 3apH:)l(eHHbie cpyttKU:110HaJibHb1e rpynrrbI.

Pa3J1HqaJOT noJIO)l(l1TeJihHO 3apH)l(eHHb1e att110Hoo6MeHHl1KM, cpe)lM KOTOpbIX HaM60Jiee qacTO HCITOJ1b3YJOT )lM3THJiaMl1H03THJIU:eJIJIIOJI03Y ()l,3A3- u:eJIJIIOJI03y), CO)lep)l(alllYIO KaTl10HHbie rpynnbI, 11 OTPMUaTeJibHO 3apH:)l(eHHb1e KaTMOHoo6MeHHMKM, HarrpHMep Kap60KCMMeTHJIUeJIJIIOJ103Y (KM-uen- JIIOJI03y), CO)lep)l(alllYIO aHHOHHbie rpynrrbl.

,(\113Tl1I18M11H3Tl1Ill.leI1I1IOI1038 Kap60Kc11MeT11114e1111101103a

Bb16op MOHoo6MeHHMKa onpe)leJIHeTcH: 3apH)lOM

BhIJ.J.eJIHeMoro 6eJIKa. TaK, JlJIH Bbl)leJieHHH OTpHu:a-

y-r1106ym1Hb1

CTapT

PMC. 1-57. 3neKTpo4>0pes 6enKOB CbIBOpOTKM KpOBM 3AOPOeoro 1.1enoeeKa Ha 6yMare.

TeJlhHO 3apH)l(eHHOro 6eJIKa MCilOJib3YJOT aHMOH006MeHHMK. IlpM nponycKaHMH pacrnopa 6eJIKa qepe3 KOJIOHKY npOqttOCTb CBH3hIBaHHH 6eJIKa c aHMOHOo6MeHHl1KOM 3aBl1CHT OT KOJimecrna OTpHUaTeJibHO 3apH)l(eHHhIX Kap60KCHJlhHbIX rpynn B MOJieKYJie. 6eJIKM, a)lCOp6MpOBaHHbie Ha aHMOH006MeHHl1Ke, MO)l(HO CMbITb (3JIIOl1POBaTb) 6ycpepHblMl1 pacrnoPa.MH c pa3JIHqHOH KOHUempauMeil COJIH, qaiu:e Bcero NaCl, M pa3HbIM11 3HaqeHMHMM pH. HoHbI XJiopa CBH3bIBaIOTCH c IlOJIO)l(lfTeJihHO 3apH)l(eHHbIMM <l>YHKU:110HaJibHhlM11 rpynnaMl1 aHl10H006MeHHl1Ka 11 BhITeCHHIOT Kap60KCMJlbHbie rpynilhI 6eJIKOB. Ilp11 HM3Kl1X KOHUempau11HX COJIM 3JIIOMPYJOTCH 6eJIKM, cna6o CBH:3aHHhie c aH110Hoo6MeHH11KOM. ITocTeneHHOe )'BeJIJfqeH11eKOHUeHTpauMH COJIH HJIM M3MeHeHMe pH, ~o MeHHeT 3aPHJl 6eJIKOBOH MOJieKyJ1bl, IIPMBO)lHT K Bbl)leJiemtIO 6eJIKOBblX <l>PaKU11H, B OJ.I.HOM 113 KOTOpbIX HaxO)lMTCH MCKOMbIH 6eJIOK.

Acl>tl>HHHaJI xpoMaTOrpact>m1,

HJIH xpoMaTOrpatl>HJI no cpO)lCTBY

3To Ha116onee cneu11cp11qHbIH MeTOJ.J. Bhl)leJieHHH: 11H)ll1BH)lYaJihHbIX 6eJIKOB, OCHOBaHHhIH Ha 113611paTeJibHOM B3aMMO)leHCTBMM 6eJIKOB c JIMfaH)laMM, np11KpenJieHHbIMH (HMM0611JIH311POBaHHbIM11) K rnepJ.J.OMY HOCMTeJIIO. B Kaqecrne J111faH)la MO)l(eT 6bITb 11CITOJib30BaH cy6cTpaT HJIM KOcpepMeHT, eCJil1 Bhl)leJIH:IOT KaKOM-JIH6o cpepMeHT, aHTHreHbl )lJIH Bbl)leJieHHH aHTHTeJI M T.Jl. l:lepe3 KOJIOHKy, 3aITOJ1HeHHYJO l1MM06MJil130BaHHblM JIHraH)lOM, nponycKaIOT pacrnop, coJ.J.ep)KalllMH cMecb 6eJIKOB. K JIHraHJ.J.Y npMCOe)lHHHeTCH TOJibKO 6eJIOK, cneu11cpmHO B3al1MO)leMCTBYIOIUl1M c HMM; BCe OCTaJibHbie 6eJIKl1 BbIXO)lHT c 3JIIOaTOM (pHC. 1-58). 6eJIOK, an.- cop611poBaHHblH Ha KOJIOHKe, MO)l(HO CHH:Th, rrpoMbIB ee paCTBOPOM C l13MeHeHHhIM 3HaqeHMeM pH MJIM H3MeHeHHoil MOHHoil cMJioil. B HeK0Toph1x CJiy'-laHX11CilOJih3YIOT pacrnop )leTepreHTa, pa3pbIBaIOIUHH rH)lpocpo6Hbie CBH3H Me)l(Jly 6eJIKOM l1 JlHfaH)lOM.

AcpcpMHHaH xpoMaTOrpacpHH ornlfqaeTCH BhICOKOH H36MpaTeJibHOCThIO M noMoraeT oq11cTMTb BhI- J.J.eJIHeMhIH 6eJIOK B TbIC5Jql1 pa3.

3. Dllucnuca Oe.!IK06 om HU3KOMO.rleK)l.11.RpHblX npUMeceu

)],ml YJ.J.aJieHMH HM3KOMOJieKyJIHpHhIX COeJ.J.11HeHl1M, B qacTHOCTM cyJibcparn aMMOHHH nocne BbICaJIMBaHMH, npMMeHH:IOT J.J.MaJIM3. MeTOJ.J. ocHoBaH Ha TOM, qTO qepe3 nonynpoHMU:aeMyio MeM6patty,

He rrpoXO)UIT 6eJIK11, 11MeIOIIJ,11e 6onee BbICOKYIO MOJieKYJUIPHYIO Macey. B CTaKaH 60JibllIOH eMKOCTM (OKOJIO 1 JI) c 6y<PepHbIM pacrnopoM ITOMeIIJ,a- IOT rronyrrpoH11uaeMb1:H MeIIIoqeK, 3arronHeHHhIH pacrnopoM 6enKa c COJihIO.

CKopoCTh BhIXO,D,a con11 113 MeIIIoqKa B 6y<PepHhIH pacrnop rrporropu110HaJibtta rpa,D,11eHTY ero KOHUeHTpaU11H ITO o6e CTOpOHbI OT MeM6paHbI. flo Mepe BbIXO,D,a COJI11 113 MeIIIoqKa 6y<PepHbIH pacrnop B CTaKaHe MemIIOT.

L(IDr oq11cTKH 6eJIKOB OT H113KOMOJieKYJI51PHhIX rrp11Mece:H 11CITOJib3YIOT TaIOKe MeTO,D, reJib-cPMJibT- pau1111 (CM. BbIIIIe).

L(IDr orrpe,D,enemrn qacTOThI (roMoreHHOCT11) BhI- ,D,eJieHttoro 6eJIKa rrp11MemIIOT MeTO)J,bl c BbICOKOH pa3peIIIaIOIUetl. crroco6HocThIO, Harrp11Mep :ueKTpo<t>ope3 B IIOJI11aKpJ1JiaMH,D,HOM rene, BbICOK03cP- <PeKT11BHaH xpoMaTorpa<t>11H BbICOKOro ,D,aBJieH1151. OT q11cTOThI neKapcrneHHoro 6enKosoro rrperraparn 3aB11CHT ero 611onorw1eCKaH 3<P<PeKT11BHOCTb 11 aJIJiepreHHOCTb (T.e. crroco6HOCTb Bbl3bIBaTb aJIJiepmqecK11e peaKu1m). 1IeM KaqecrneHHee oq11IIJ,eH rrperrapaT'TeM MeHhllle BepOHTHOCTb ocnm1rneH11M rrp11 ero np11MeHeH1111.

VIII. Vl3MEHEHVISI 5EflKOBOro

COCTABA OPrAHVl3MA

neJIKOBbIH COCTaB opraH113Ma 3,D,OpOBOfO B3pOCJIOfO qeJIOBeKa OTHOC11TeJibHO ITOCTOHHeH, XOTH

B03MO)l(Hbl 113MeHeH1151 KOJI11qeCTBa OT,D,eJihHbIX 6eJIKOB B opraHax 11 TKaHHX B 3aB11C11MOCT11 OT coCTaBa IT11IU11 11 pex11Ma mrraHMH, OT cP113110JIOf11qecKOH aKT11BHOCT11 qeJIOBeKa, 611onor11qecK11X p11TMOB.

A. U3MEHEHIIE BF.JIKOBOfO COCTABA K.JIETOK

B IIPOQECCE lfX )J;lf<l><l>EPEHQIIPOBKII

B rrpouecce pa3B11TMH MHOroKJieToqHoro opraH113Ma, oco6eHHO Ha CTa,D,MHX ,D,11<P<t>epeHU11POBK11 KJieTOK, 6eJIKOBbIH COCTaB 3Haq11TeJihHO 113MeH5IeTC51. L(JIH Ka)J()J,oro T11ITa crreu11an11311poBaHHbIX KJieTOK xapaKTepHo rroHBJieH11e crreu11<t>wrecK11x 6eJIKOB, KOTOpbie orrpe,D,eJIHIOT oco6eHHOCT11 11X 611onomqecKMX <PYHKU11M. TaK, TOJibKO B 3p11Tpou11rnx ecTh reMorno611H, ocyiuecrnnHIOIU11ti rpattcrropT KHCJIOpO,D,a K TKaHHM, a B KJieTKax ceTqaTK11 rna3a - 6eJIOK po,D,OITC11H, He06XO)J,11MblM ,[(JUI yJiaBJI11BaH1151 <PoTOHOB cBern. KpoMe Toro, crreu11ami311poBaHHbie KJieTKH OTJiffqaIOTCH 11 KOJI11qeCTBOM 6eJIKOB, np11cyrCTBYIOIU11X npaKT11qecKH BO Bcex MJI11 BO MHOfMX KJieTKaX opraH113Ma.

Tip11 pa3JI11qHbIX 3a6oJieBaH1151X npo11CXO)J,11T 113MeHeH11e 6eJIKOBOfO COCTaBa TKaHeM. 3T11 113MeHeH1151 Ha3hIBaIOTCH rrpoTe11HorraT11HM11. Pa3JI11qaIOT Hacne,D,crneHHbie 11 np1106peTeHHhie rrpoTe11HonaT1111.

B. IIACJIE,l(CTBEHHhIE IIPOTEIIHOIIATlflf

Hacne,D,CTBeHHhie npoTe11HorraT1111 pa3s11BaIOTCH B pe3yJibTaTe rrospe)J()J,eH11H B reHeT11qeCKOM annapaTe ,D,aHHoro 11H,D,11B11,D,yyMa. KaKotl-n1160 6eJIOK He C11HTe311pyeTCH BOBCe 11JIM C11HTe311pyeTCH, HO ero rrepB114HaH CTPYKTypa 113Mettetta. Op11Mepb1 Hacne,D,CTBeHHbTX rrpoTe11HonaT11tl - reMorno611HonaT1111, paccMoTpeHHhie BhIIIIe. B 3aB11c11MocT11 OT POJI11 ,D,e<PeKTHOro 6eJIKa B )((113He- ,D,eHTeJibHOCT11 opraH113Ma, OT CTerreH11 HapYIIIeH1151 KOHcPOpMau1111 11 cPYHKUlUI 6eJIKOB, OT fOMO11JI11 reTep0311fOTHOCT11 11H)J,11BH,D,YYMa no 3TOMy 6eJIKY HaCJie,D,CTBeHHhie npoTe11HOITaT1111 MOryT BbI3bIBaTb