Медицинская вирусология / Букринская А.Г., Жданов В.М. Молекулярные основы патогенности вирусов

41

JleHhI peuenTOpm,re ceMeHCTBa, BKJIIO'la10m11e pasHbie cepoTH· llbI BHpycoB, KOTop1,1e ne HMeIOT o6mero aHTlffeHa.

I.no.m1omrpyc1,1 T1111a 1-3.

2.BIIpyc1,1 KoKcaKH rpy11nb1 B, TIIn 1-6.

3.BIIpycu KoKcaKH rpynnb1 A, THIILI 2; 5; 13; 15 11 18.

4.BHpychI Echo, nm 1-6.

5.PHHOBHPYCbl l(CJ10BC!<a, TH!Ibl IA; 2; 3; 5; 9; 12; 14; 15; 22;

32; 36; 39; 41; 44; 49; 51; 58-60; 66; 67 II 89. BHpyc KoKcaKH rpymm A, nm 21.

.L Kap H0Bnpyc1>r.

Cneu,Il(pH'lecKoe .n.Jrn mrnopnam1pycos ,mcJ10 peue11Top0B :1a IlOBepXHOCTH O,lI.HOH KJ1CTKH o6b[lIHQ KOJ1e6J1eTCH B npe,ll.e.nax

'106.

CT O JI lI O B lI p y C hl. C IIOMOlll.blO JIYtJ:CBOH HpHCTaJ1J10rpa­ qJIIII ycTaHOBJICHbl TpexMep11aH CTPYKTypa IIOJ1HOBHpyca I1 B33· HMopacnoJ1o)KeHHe 4 Bttpycm,rx crpy1nypm,1x 6eJIKOB (VPl, VP2, VP3 Il VP4). Ha nosepxnocrn BHpyrnoii tJ:aCTIIll,bI 3KC· rroHHpoBaHhI 3 6eJ1Ka VP1, VP2 II VP3, caMbIH MaJ1CHbKHH VP4 3aHHMaeT BHyTpeHHee IIOJ10)KCHHe, HO KOHTaKTHpyer C OCTa.lJbHbIMH 6eJIKaMH. TiepBb!C 3 6eJIKa 5lBJrnIOTC5I OCHOBOH KarrCH,ll.HOH o60J!Ol!KH, Ha)J. KOTopoii BblCTYllaIOT IICTJ11I aMHHO· KHCJ10T II yrneBO,lI.OB. TaKIIe neTJ1lI H Bb!CTYllbl cpopMHpyIOT Hapy)KHYIO nosepXHOCTb BHpHOHa H ero 3HTHreHHhie Y'JaCTKH. B BIIpHoHe o6'be,lI.ttHHIOTCH 5 KOIIHii VP1, o6pasyH 11HTHrpaH11y10 ocesy10 crpyKrypy. Ihrn noii CTJ)YKTyp1,1 oKpy)KeH yrny6

Fab-cpparMCHT aHTIITCJI, Jl.liaMeTp KOTOporo OKOJIO 3,5 HM, He MO)KeT ,ll,OCTH% ,lI.Ha KaJlbOHa. TIO-BH,Ll,HMOMY, Bb!COKHH KOHcep­ B3TH3M Y3H3!0lll.lIX peuenTOpbl yqacTKOB np1rnpenHTeJibHblX 6eJIKOB o6bHCH5ICTCH HMCHHO 3THM o6CTOHTCJibCTBOM - OTCYT· CTBHCM HMMYHHOI'O rrpecca. TeM C3MbIM rroJ1yiraeT 06'bHCHCHHe HaJ1HtJ:He o6mHx peuenTopoB y I1IlKOpHam1pycoB, He HMC!OW.HX 06m11x aHTHreHOB.

BbICOKHH KOHCepBaTI:I3MIIOCJIC,lJ,OBaTeo1bHOCTefi 6eJ1KOB, y3-

42

HaI0ll!JIX K.TJCT0llHb!C peuernopbl, II3BCCTeH H ,UJUI ,upyrnx BH­ pycoB H MO}KCT o6ecnelfHBaTbCH a,ueKBaTIIblMH MCXaHH3M3MII. TaK, «per(e1uopm,1ii KapMan» BIIpyca rpmma 11 yrny6J1em1e Ha Il0BepxH0CTil rJIHK0llpoTCH,IJ.a gp120 B114 HBJI5IIOTC5I o6pa30BamrnMH, aH3J10flll{HblMH KaHb0HY Ill!K0pHaBHpycoB. BbrnB­ JieHHC BblC0K0K0HCepBaTHBHblX yirncTK0B B 06.JiaCTH ysnaIOI.I.l,HX pe1(eIIT0J)bl IIOCJIC,UOBaTeJlbHOCTeH OTKpbrnaeT llllIJ)0KIIe B03MQ)KIIQCTH ,U,JI5I XIIMil0Tepamm BnpycHbIX 1rncpeKll)Il'i. MoryT 6b!Tb Il0JiytJeHU Xl!MH0IIperrapaTbl, crroco611ble 6JIOKHp0B3Tb 3TH IIOCJlC,U0BaTeJibH0CTH H HapyrnaTb H3ll3JlbHble CT3,UHH pe­ rrpo,uyKUHH BHpycoB. O,uuH H3 HIIX C0CTOHJI ll3 ru,upocpo6HOH fOJ!OBKH - cpeIIOKC330Jia, 7 3J1HqJaTII4CCKHX uenot:JeK H H30ECa30JlOBOro XBOCTa; XB0CT rrp!IKpCIIJlHJlCH K rn,upocpo6HbIM rro­ CJle,UoBaTCJlbHOCT5IM BHYTPCHHCH noBepxHOCTH -cTBoJia VP1,

pa3y10T II0BCJ)XllOCTb BHpycnoro KaHCH/l.a, B TO BJ)CMH KaK VP4 Haxo,uHTCH Me2K,uy Karrcn,noM II PHK, HO KOHTaKTHpyeT c VPI 11 VP2. Ha 1v10,neJm p11HoB11pyca nma 14 6b1J10 noKa3a­ no, llT0 6eJIKH o6pa3yIOT 1rnn1rpannyIO CTPYKTYPY, rJiaBJ-IaH tJaCTb Kornpofi cciiop1v111poBaHa cy61,c,umrnuaM11 BToHOcrpyn­ rrnpoBaHHbIMH oKo,10 nenTaMcp11oft OCH. BoKpyr aT011 LJaCTH 115ITllrpaHHIIK3 npOXO}(IlT rny6oKHH K3Hb0H (rny61rna OKOJIO 2,5 HM). Ero «!0)KH3H» CTCHKa orpaHIJlJCHa yrJICB0,UHbIMH ocTaTKaMn MOJieKyJibI BpoaO«ceBepnan»- yr.neB0JlHbIMH ocTaT­ IGJMH VP3, ,uHo o6pa3oBaHo MOJ1CKYJiaMH VP MB3Ha ,une K3Hb01ia H3XO,UHTCH Bb!COKOKOHCepBaTHBHbIC ,/l,JIH Bcero CCMeikrna ll!IK0ptraBHpycoB IIOCJlC,IJ.OB3TCJlbHOCTII, Y3H3IOULIJe KJICTO'IHbIH jlCILCIITOp, a Y43CTI{Il CB513b!B3HHH nei'npaJIIl3YIOU(HX aHTHTCJl

JI0JlylIHJ1ll MOJ-IOKJIOH3.Ilbllb!C aHTHTCJia, cnoco6Hble 6JIOKHpO­ llaTI, peuenT0pHbIH yqaCTOK 11yBCTBI!TCJlbHb!X KJICT0K. TaKHe aIITllTCJ[a 6JIOKHJ)OBaJIH H penp0)I,yKUlll0 HCK0T0phIX BHpycoB KoKcaKH A, Bnpyca Be3HKyJrnpnoro cTOMaTHTa, 11apaMHKC01111pyecrn, 1mpyca rpnnrra A, a,ueH01rnpycon H ,up.

aHT

HHa

Bel!KOM, Y3HaIOll]J1M peuenTop, HBJUieTCH MHHOpHb!H 6eJIOK crl, KOTO{)biii B BII,ll,e ;'I,HMepa uaxo,n,HTC5l B Hapy:>KH0M Kanc11,n,e [Yeung M. et al., 1987]. 3TOT 6eJIOK 0TBeTCTBeH 3a CBH3bl­ Bami:e C KJieTK011, TKaHeBb!H H KJieTQllHblH Tp0ilH3M. 3TOT :>Ke 6eJioK HB.1HeTcH BHpycHbIM reMarrnIOTHHHH0M. KJ1eT01rnhIM pe11errropoM, yaHaBaeMblM 9THM 6Cl!K0M Ha II0BepXH0CTH 9pHT­ poumoB 4C.10BCK3, CiIY)KIIT rmrno¢opHH. OIJHIUeHHhIH rJl!IK0- <pop1rn 9(plj)CKTIIBHO rro,naBJIHCT reMarrJIIOTHHaUHIO, a TaK)KC CB.H3b!BailHC BHpyca C MblIIUIHblMII L-¢116po6JiacraMH. L11IIKO­ ¢op1rn, o6pa6oramrnfi Heil:paMHIHIJ{a30il, TCp5IeT peUCIIT0PHY!O aKTHBII0CTb. TaKHM o6pa30M, cyu CCTBCHHOH IJaCTb!O peuenro­

ocyn1ecTBJIHeTC5l rroc.1e,noBaTeJibll0CT5IMII al C C-KOHIIa M0Jie­ KYJll,l, K0T0()b!C HaX0,L(HTCH Ila Hapy)KHOII noeepXH0CTH BHpH0Ha, B TO BpeM5l KaK K0HCU: 33MaCK!Ip0B3H H He 9KCll01IHp0B3H Ha noeepxH0CT!L Y3HaIOrunH peuenrop y•rncT0K HMeeT CJIO)K- 11y10 CTpy1uypy, B C0CTaBe KOTopofi ecTb cerMCHTbl, HMelOIUHC 60J]bll1Ce 3ll3'IellllC lVl5I reMarUIIOTIIH3Ulflf, 1JeM ; JI}l np11Kpen­ J1CHirn: K KJieTKe [Nagata S. et al., 1987].

B H p y c r e n a T H T a B. KJieT01rn1,rii penerrrop Ha II0Bcpx11ocTH renaT0llllT0B Y3HaeTC5l noBepxH0CTl!b!M aHTlifCH0M

m1pyca renanna B - HBs-aurnreHOM. HBs-anT11ren co,n,ep­ ,Kl!T 3 0CH0BHblX 6eu'IKa: 6CWbUIOH, cpe.ll,Illlfl H M3m,Iff S-aHTH·

0611.ribHO Ha noBepXH0CTH BHpycHbIX IJ3CTHU npe,LicTaBJiefl 60.16llI0H. S-aHTHreH (p24). EeJIOK p30 BK,1I0IJ3CT B ce6H TI0JJH0C'rb!0 6CJIOK p24 II ,ll,0ll0JJHHTe.1bHYIO noc:rre,n,onaTCJibH0CTh 113 55 alvrn­ H0KHCJIOTHblX 0CTaTK0B N-KOHI(a. BeJIOK p39 co,n,ep)KHT 6eJIOK p30 II 1'I,0TI0JlHHTCJibHhlC 117 3MHH0KHCJI0THblX 0CTilTK0B Y N-KoHua M0JieKy.rrbl.

B 6oJiblllOM H cpemreM S-anrurene HaX0JJ.HTCR nerrTH,n,Han nocJICJ(0BaTeJibH0CTb pre-S2, B 6oJib1HOM S-aHTHreHe - neII­ TIIJJ.HaH noc,ne;i,osaTe,11bH0CTb prc-S1. Ehn 11ocJie,Li0BaTe.1JbH0CTH

45

TOUHTOB 6b!JIH Bb!HBJieHbl ,ll,Ba pa3JIH'!Hb!X peuenTopa, 0)1,!iH H3 KoTopbIX CBH3hIBaJI 6e,10K pre-SI, Jipyroii cBH3bIBa,1 pre-S2 npH yqacnm TIOJIHM€pH30BaHHOro CblBOPOT01IHOro aJib6yMHHa. Ilo-BH,ll,HMOMY, npHKpcmneJibHblM 6eJIKOM BHpyca renaT11Ta B HBJIHCTCH pre-S1, KOTOpb!H npHMO CBH3bIBaeTCH C peuerTTopoM CBOHM N-KOHUOM. TTo,'IHMepH30BaHHbiit cbmopoTOlfHbIH aJ1b6y­ MHH HBJIHeTCH rrpoMe)KYTOlJHblM peue1nopoM, npHKpeITJIHIO­ ll(HMCH K ITJia3MaTJ!TieCKOH MeM6pane rcnaTOUl!TOB H Y3HalO­

mttMCH 6eJIKOM prc-S2 B coCTaBe 6oJibworo u cpe.n,J-Iero S-aH­

P. et al., 1989].

S-aHTHreHa, B03M0)1{HO, 3a crreT KOHcepBanrnnoii aHTHreHHOH

.n,eTepMnnaHTbl «a» [Peeples M. ct al., 1987), t1acTHUbl peKOM- 61rnattTI-IOro HBs attn1rena, co.n,ep)K31ll.He TOJibKO MaJibiii S-6eJIOK, TaK)Ke 6u,111 cnoco6m,I nplIKperrJIHTbCH K KJieTKaM

COB rpnrma HBJIHeTCH 60Jibl!JaH cy61,e)Umnua MOJieKyJJbT re­ MarrJJIOTHHHH3, a cneunqrnqecKHMH peuenTOp8MH 11a KJlCTOt{­ HOH IIOBCpXHOCTH - coemme,rn.H , co.n.ep1I<a11I.He neiipaMHHOBYJO KHCJIOTY H npe,IJ,CT3BJ1RIOmne co6oi:i rJIHKOIIpoTeH,11.bl HJIH cua­ J!OrJIHKOJIHilli,ll.I-,I (raHrJIH03H)lbl). I(oJIH'IeCTBOKJieTO'!HbIX pe11,emopoB Ha O)J,HOH KJieTKe onpe.n,eJlHJJH npn 3apa)Kemrn KJie­ TOK CMeCb!O OlJHlll.eHHOro HeMe11em1oro H MelJeHIJOrO [3H]-ypn­ ,ll,!HIOM nupyca (wTaMM BHpyca KYPHHOii 'IYMLI)' a KOJIHtteCTBO CBH3aBwerocH nnpyca - no yMeHbWemno co.n,ep)KaHI-IH reMar­ rJlfOTHH!IHOB H pa.n.110aKTHBHOCTH B )KHJlKOH cpe,rr.e rroc.r1e HHKy6aunn C K.'IeTKaMH rrpH 4 H 20 °C. 0Ka3aJIOCb, 'ITO HeMetteHbIH

06pa6oTKa K.JJeTOK HeiipaMIHIH,ll,330H pe3KO CHH}K3eT 3)l­ cop61m10, o,rr.HaKo OHa BOCCTaHaBmrnaeTCH )J,O HCXO)J.HOro ypon­ HH npn BCTpattna1um 3K30feHHblX peuenTOpOB, B qacTHOCTH CO,llep)K311I.IIX cnaJioByro KHCJIOTY rJIHKOJIHrIH,ll,OB. TaKHe KJJeTKH C pa3pyrneHHblMH npHp0,11.HblMH penenTopaMH npe.n,cTaBJI5110T co6oi1 yno6uy10 Mo,neJib .n,M1 1-13yqemrn penenTopHbIX cpyHKUJif1 aK orem1h1xpeuernopoB.

ricpBifI1Ha.H cTpyKTypa MOJICKYJibI reMarrJIIOTHH!ma H ee Mo,urnp11Ka1mu, Be,IJ,yu ne K H3MeHefnno TKauenoro Tpon113Ma

46

lrnpyca rpimna, II3ycreHbJ aua,urreJJbHO nom1ee, 1ICM y ,l1,pyrn:{ m1pycon.

Ya1rnrowJ1ii peuenTopb1 yiiacTOK rcMarrJJIOTHmrna pacnoJia­ raeTC.H B perienropHoM KapMaHe; .rtno KapMaHa o6paaonano q:>eHOJJOKCiffeHOM Tl!p03HHa 98 H rn,11.pocpo6HOII nonepxHOCTb!O TPHUToq:>airn 153, 3a,[lmrn CTCHKa BblCTynoM n!CTli,LlIIHa I 83, COC,11.I!HCHHOfO BO,llOpOJ(HOfi CBH3bJO C THp03HHOM 98. B nepx­ Hefi qacTH 33)],HCH CTCHKII IIaXO,llHTC51 Y,.JaCTKH rJiyTaMnHa 190 H JICihuma 194, JJCBYIO CTOpOHy o6paaymT aMIIHOKHCJJOTHbIC OCT3TKH 226-228 n JJCBOM yrJJy H 133--137 n nepC,11.HCM yr­ JJY, npanaH cTopo1rn 06pa3onaHa TpeoHHHOM 155. N-au_eTHJI­ HeiipaMHHOBa.H KHCJJOTa B COCT3BC peu_cnTopoB rmru_epHHOBOH Il.Cllb!O MO:.>I<:eT B33HMO,llCHCTBOB3Tb C THp03HHOM 98, rJJyTaMH­ HOM 90 n cepHHOM 228. MeTHJIOBbIC rpyIIIIbI N-aU,CTl!JIOBoro y•iacTKa MOryT HMCTb rH,npocpo6HyIO CBH3b C TpIIIITOqlaHOM 153 H JJCHil.HHOM 194, a KHCJJ0!)0,11. a30T11CTblX 11 yrJieBO,ll.HbIX yqa• CTKOB KOHT3KTHpyer C llOJIHpHbIMH aroMaMH 3MIIHOKHCJIOTHbIX OCTaTKOB B llOJIO}KCHJrnX 134 rr 135.

KoucepBaTHBHbIC IlOC.'Ie)],0B3TCJibHOCTII aMHHOKHCJIOTHbIX OCTaTKOB pacnoJIO}KCHbI Ha )],!IC peu_enTopHoro KapMaHa 11 OKpy}KeHbl BapbnpyromnMH J'IaCTKaMH. TI03TOMY CJie,11.yer O)KH­ JlaTb, LJTO aHnneJia nponrn npnKpenHTCJibHOro yqacrKa reM­ arrJl!OTHHHHa He 6yJWT 9q>cpeKTHBHO 33ll\HII.l3Tb OT HmpeKil.lUI, nocKOJihKY OHH cKopee 6YllYT peanrpoBaTb c Bapb11pyromHM11 aMl!HOKHC.10THblMH OCT3TK3MH, '-ICM C MaJI0,11,0CTYIIHblMH (B CII­

.r1y TOnorpaq:>HlJCCKHX oco6eHHOCTC!i) KOHcepBaTHBHblMH yira­ CTKaMH MOJJCKYv1bI IIpHK!)CIIHTCJibHOfO 6CJIKa. B03MO:lKHOe BMC­ IIIaTCJlbCTBO B npouccc CB.H3bIB3Hlf5I BrrpycaC pCil.CTITOpOM MO:lKCT 661Tb 0CHOB3HO Ha 6JIOKHpoBaHHH y3na10mcro peucIITO·

N-Komia MOJieKyJILI 6enKa HA2.

HMeIOTCJI ,[laHIIbIC, CBH)lCTCJlbCTByiom11e O 11pe3BbIIrafmo Bbl­ COKO!i Cl1CllflqJJ11mocTH Y3H3!0II.lHXpcu.ernopbI yqaCTKOB reMar­ fJIIOTHHHHa. .:ha cneunq:>wmoCTb OII!)C)lCJI5ICT KJICTOqHbHI TpO­ flI!3M BHpycon rpimna, Kpyr 'IYBCTBIITCJibHbIX X03.HCB, ce ll3MC­ uenne .'IC}Kl1T B OCHOBC a)laI1TaIJ.Hl1 n11pycaK JlpyroMy X03mmy. TaK, aJJ,amaIJ.H.H K KYPHHbIM 3M6pnouaM Bnpyca rpnnna TH­ na B, BbI)lCJICHHOfO OT qe.TJOBCKa, np11B0,IJ.HJia K yTpaTc aM11HO­ KHCJIOTHbIX OCTaTKOB 197-199 n MOJieKy.'Ie HAI, KOTOpbre .HB­

.!J.HIOTC51 llOTCHil.H3.'IbHblM yqaCTKOM fJIHK0311JI11pOBamrn. A)lan­ Tail.!1.H nnpyca rpmma rnna A (HlNl) 6wia CBH3aHa c 3a­ Mcnoii aMHHOKHCJIOTHbIX OCTaTKOB 138, 163, 187, 189, 190 11 225, a aJJ,amau1rn n11pyca rpmma THna A (H3N2)- c 3aMe­ HOH aMHHOKHC.'JOTHbIX OCTaTKOB 137, 138, ]56, 193, 216, 246

47

H 248 B Mo.neKy.1Je HAI. lJacTb 3THX 3aMe11 rrpHBOJJJIT K rroTepe Y'IaCTKOB fJ1IIKO311JIHpOBaJrn5!. Bee 3TH 3aMellbl JIOKaJIH3OBaHbI B a1n11reIIHh!X Y'WCTKclX MOjJCKy.Hbl reMaITJl!OTHH11IIa p5!,ll.OM C pen.e1nopI1b!M KapMallOM. ,Ua»,e lI3MCHCll!IC O,l]JlOfO OCTaTKa B IlOJI02KeHim 218 (fJIH-+- fJry/Apr) conpOBO)K,ll_aeTC5! Hapy­ rneHHCM aHTHI'CHHOfr cneumjrn•IHOCTH, peuell'ropHOH aKTHBHOCTH II TpOITI13Ma BHpyca.

Cpauuemre nepuH'!HOi'I CTPYKTYPLI MOJICKYJibI reMarrmOTH· mrna BHpyca rp1mrra Tima A (H3N2), a,11_amHpoBa1-rnoro K KY· plU!bIM 3M6pIIOJlaM II llOlleamblM KJJeTKaM rrepeBHBaeMOH JIHHHH co6aKH M,UCI(, noKa3aJIO pa3JIHarne 110 5-6 aMHHOKHCJJOTHLIM OCTaTKaM. llpH 3TOM y BCCX uapnaHTOB 6bIJia o6uapy2KeHa sa­ MeHa OCTaTKa B 11OJimKeHHH 156, Haxo,11_51u eroc51 Ha KOHUe MO·

.JJeKyJibl reMaSfJIIOTHHlilla y Y3H3IOlUCfO peuernop yqacTKa.

TT3MCHeHH51 rrepBHIJHOii CTPYKTypw KoppeJIHpoBaJIH c 0apHa­ UH.HMH aHTHfCHHbfX H 6HOJIOrH11ecKHX CBOlkTB a,n_arrTHpoBaH­ HbIX Bap1rnHTOB [l(atz Mpoet al., 1987].

I1pH cpaBHeHHH reMarr.lIIOTHIIHH3 H3 BHpycoB rpunna IfCJIO· se1rn, ITTilU 11 JlOIIIa,n_eii 6LIJIH Bb151BJJeHbl ,LI.Ba TllTia Y3HaJOIIUIX peuerITopLI CTPYKTyp: u11pycL1 rpmma 'IeJ1oueKa y3HaBaJiu CB5!3b eapT2,6 Gal, a B11pych1 rpimna IITHU 11 JI01IIaAef1 CB5l3b1B3JIHCL c KJieTKaMH, COAep2KaI.UHMH CB5!3b eOHT2,3 Gal [Rogers G., Paul IHao1983; Rogers G. et a aIA1983].

Pa3JIHlJH5f u pe1 eHTOpH011 aKTHBHOCTH BHpycou rpmma 6L1JIII 0611apy2KeHL1 11 y 1IITaMMOB o;u1oro HOAT1rna (H3N I) B npo1iecce ,L!.peiiqrn. 9TH 113MeHeHHH 6LIJIH o6yc.IIOBJICHbl 3aMeHOH aMHHOKHCJIOTHLIX OCTaTKOB 155 l:Wll 188, KOTOpb!e pacnoJIO­ )KeIIbI y peuernopHoro KapMaua MOJieKyJILI HAl [Higa H., Paulson I., 1985; Higa KOo al., 1985; Anders Teoet al., 1986).

CymecTBeHHhrM q>aKTopoM ,1J..1rn Y3HaBamrn K.neToqttoro pe­ u,eHTopa 5!BJ!51CTC51 yr.r1eBO,Ll.HblH KOMIIOHeHT MOJieKyJILI reMar­ fJI!OTHHl!Ha. ,UJJ51 CIHI3b!BaHirn C KJICTOl!HblM per enTOpOM HMe­ eT 3Ha'!eHHe He CTOJibKO H3JIH'!He OJillrocaxapII,L!.HOH II,CITO'IKH B onpe,11_e.rie111IOM ITOJIO)KCHHH, CKOJibKO KOH<pOpMaUH5! OJJllfOCa­ xapH11,a, o6ycJIOB.JJCHHa.H KJICTKOii-X035IHHOM, B KOTOpo!'inpottc xoJuura penpo,11_yKUH51 Bttpyca. CTpH a,1J.amau1111 m1pyca K ,L!.py­ roMy XO3mrny npOHCXO,ll_lIT IIOTep51 l!Jlll H3MeHeHne rJIIIKO3H·

48

3MopHOHa 0blJla Cl35!3aHa C H3MeHeHIWM HJIH notepe:fi rJHHl'.03Jl·

nrpaIOT ooJir,rnyto MCIHbB npoueccax, cBsl3aHHbIX C I(JICTO'IHO}i nonepxHOCTbJO.

LlHr.ll!I031!,LLhl o6m:1)],3IOT peuenTopnoii cj)yHKUHClI )],,'HI pH)l3

PHK-co,ll,ep1KaIUHX 1rnpyc0B: rrapaMHKC0B11pycon [Haywood BTO Cl0yyH3BHpycoB rpmma [Bukrinskaya ua3XeO3C3Hcr3Bergel­ son KabD., C3HCM311 6erneHCTBa [Superti F. et al., C3H6J3Con­

OHTnB3et al., C3HHiK3

KoJill'ICCTBO OCTaTKOB CHaJIOBOH KllCJJOTbl H HX IIOJJO)KeI!ITe 3naq11TCJibHO BJIH5IIOT Ha peuernopHyIO aKTHBHOCTb raHrJll103H­ JJ.0B. Ha1160J1ee aKTHBna,r cTpyKTypa oXuryCBH3bIBaHHH Bttpy­ COB - raHrJUI03HJl.bI C6H0yCy30ayKOTOpbie COAepmaT ocTaT­ RH CHaJIOBO!! KHCJIOTbI, pacnOJIO)KCHHhie Ha «BHyTpeHHCH» H «K0Hueno11» raJiaKTose. ITpH 3TOM )],Ba ocTaTKa Ha «BHyTpeH­

uet1» raJiaKT03C 3H3'IHTeJibllO YBCJIH1IHBaJOT 3q)q_JCKTHBHOCTb CBH3b!B3HHH BHpyca [Superti Yp3et al., C3H6yo3raHrJIH03H,LLbl C y)l,aJieHHblMH OCTaTK3MH CHaJ10BOfI KHCJIOTbl BHpyc HOaCBH· 3b!B3IOT.

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