
Медицинская вирусология / Букринская А.Г., Жданов В.М. Молекулярные основы патогенности вирусов
.pdf
Puc. 6. f 1.1Cl)o60IB](r11J-Jr |
6uma |
|||||||||
C('PAl(CIHIJlbl |
(p24) |
11pH |
o6paCioTKC |
|||||||
BI·Jll |
A. |
; cn'prc11rnirn |
|
[oyKp1111- |
||||||
rnaa |
r., |
Waposa H. |
K., |
1989). |
||||||
B11pyc |
o6pa6aTb1Ba/IH |
0,5% |
)J.e30KCIIX0· |
|||||||
/lUTOM HaTpRI! (AOPOlKKll 1; 2) |
H lo,\, |
|||||||||
NP40 (,topo:H<1rn :J---5) a Teqenm, |
JO MIIH |
|||||||||
npH |
20 ·c. |
3aTeM |
cy6s11pyCHble |
CTPY1'· |
||||||
rypw |
|
oca:>K.n,am, |
IJ,CHTpllq)YrHPOBa1111cM |
|||||||
up!{ |
40 000 |
Is, |
a |
6e,1K» |
H3 |
Ba)J.0C3A0'1· |
||||
Holl JKl{Jl,K0CTH - aIJ,CT0H0M, BeJIKH oCa/l.· |
||||||||||
KOil (AOpOlKKll 2; 4) 11 na11,oca;\O'IH<Jii |
||||||||||
)KJl)J.KOC'fl{ |
|
(AOPOJ!{K!I |
I; |
3) |
pas)J.emrnH |
|||||
C HOMOIIJ,b!O 3JICKTpoq,opesa B nAAr H |
||||||||||
HJJ.CRTH<!>IIU.IIPOBaJm |
C |
ll0M0ll.l,b!0 |
HMMY• |
|||||||
1106J10TTHHra, |
HCIJ0Jl"3Y |
ChlB0p0TKY KpO |
||||||||
l!H |
cepono3HT1Hnwro |
Juu1,a. |
5 - Ucmo-1 |
|||||||
l!CXOA!iOfO |
|
BHpyca. |
|
|
|
|
|
2 3 4
•
110HllblMH CB513}1MH, TO C JIH· nonpoTeHJI,HoA MeM6pattoA on accouttHpyeTC51 6naroAapH CBOHM n!,L!.pOOOq)HbIM CBOlkT DaM, Eho ttecneu1-Hpw1ecKa>-1 CBH3b, H npH CMernaHHOH llH cpeKlOH:I K.TleTKH JI.BYM51 BHpy- caMH MOiyr o6pa30BaTbC51 nceB,Ll.OTlU[bl, CO,Ll_ep>Kauu1c HYKJ1e0Karrc11;\ H M-6CJJOK O/l-
Horo BHpyca, a J'J1IIIWllpOTCIIJ.IJ,l - ;.1,pyroro. Tal,HM o6pa30M, ,UJUl. c6opKH BHpycnoH 1JaCTHUhl HeT Heo6XOJ]JIMOCTH s cneuH cp1111ecKOM COOTBeTCTBHH BHyTpeHl!HX 6eJlKOB rJil1KOnpoTeHJl,3M Ha Hapy)l{HOH TIOBepXHOCTH KJleTKH. B TO :>Ke BpeMR B33HMO ,!l,eHCTBJ1e M-6eJIKa c HYKJleOKancH,!l,OM cneuHq:JlllleCKOe, no cIrnnbKY HeT HX CMell!HBclHIUI npH 3apa,1,eHHH K,leTKH p33Hbl· MH BHpycaMH,
PoJib MaTpHKCHOI'O 6enKa s c6opKe BHpyca MO)l(eT 6bITb MHoroo6pa311011. 0;1,Hoii 1:13 B3)1<1I1,Ix cpyHKUHH M-6e,1Ka 51BJrn erc51 H3MeneIrnc KompopMal.l_HH HyKneoK311CHAa npH B3aHMO· ,LJ.eikTBTJH M-6enKa C HHM - H3MCHeHJ{C, Heo6XOJ]J JMOC ,LI.JUI BKnIO L[eHH51 HyKneoKancn,1.1,a B COCT3B TI01!KH H onpe.uem:.11on1.ee, 6y;1,eT Jlli HYKJICOKaTICHJJ, CJlY)½{HTb MaTpHl.l_eH J].Jl51 TpaHcKp1m UHH reHoMa HJlli BOHAeT B Cocran BHpycHOH 113CTHJJ,bl. J,faBeCT HO, 11To M-6enoK noJJ,annsrer TpaHCKp1rnumo Ha Mwrpane HYK·
JJeoKancnAa s11pycon Bea1rnyJ1HJ)Horo CToMa·nna [Le Y. d al., 1989], rpI,rnna [3so1-rnpesA., re11J1,0H IO. 3., 1980] H napa
rpmrna 11cJ1oseKa rnna 3 [KpaCbKO A. r., EyKpHHCKa51 A. r., 1986], B TIOCJie,UHCM cny1rne M-oeJlOK 6blJI 9cpcpeKTHBeH rrpH
cooTHorneHHH NP :M= l :0,2, r. e. B ropa3.ll,O 60J1ee HH3KHX OTHOCHTeJlbHblX KOHUCHTpaurrnx, l[CM B BHpHOHC. Bo3MO)i{l:10, M-6eJJOK B3aHMO.lleHcrnyeT c HYKJieoKancn;1,oM y2Ke B 1 11ro-
101

I1Jla3Me; eCTb ,ll,aHHJ,IC O TOM, lJTO Ha TIOBepx11ocn1 UlfTOJ1JJa3· MaTJJlJeCKHX HYKJJeoKal!CH,ll,OB rrnpyca Kop11 C IIOMOlilblO HMMY· H0.9,TTCKTpOUHOH MHKpoc1<0mrn MO.'lKHO 0611apy)KHTb MOJICKy.'Ibl M-6e.r1Ka.
BKJJJ011eHne M-6eJIKa B IIJJa3Man111ecKy10 MeM6paHy K.11eTKH rrpHBO,ll,IIT K rrepepacrrpe,ll,eJJeHIIIO rJlHKOllpOTeH,ll,OB Ha napy)E· HOil noaepx11onn KJICTKU II !IX a,ll.eKBaTHOMY ,ll.Jl5I ll0lJKOBaHl]51 pacrrOJJ02KCHIIlO. B BX TpaHc,noKaivrn yqacTBy10T He CB060,ll,Hb!C MOJieKyJibl Ha M-6eJJKe, a ero KOMITJICKC c HYKJieoKallCH,ll,OM: xapaKTepHb!C lIJHllbl Ha Ke'ICT011HOH rroaepxHOCTI1 q>OpMHpy10TC5! J!HUib B TOM CJ1ytJae, KOT,l],a IIO)l MO)lH!pHl(HpoBaHHblMH ytJaCT
KaMH l(JieTOl!HOH MeM6paHbl BblCTpaliBaJOTC51 |
HYK.JleOKaIICIIJ(bl |
B KOMn.'IeKce c M-6eJIKOM [Heggeness M. et |
al., 1982]. |
BKJUO'leHHe M-6e.rIKa B IIJJa3MaTw1ecKym MeM6pany HBJI5I· CTCR JJHMHTHpyIDWJJM co6blTHCM, onpenemuomHM B03M0)]{HQCTb IJOl!KOBaHH5I BHpycHblX qaCTHil. B COOTBeTCTBHH C 3TOfl tj)yHK IJ)fCH M-6eJIOK )la)Ke npH nyJihCOBOH MeTKe npaKTH11ecKtt He o6Hapy.'lKHBaCTC5! B 3apa)](eHHOfi K.'ICTKC H ero paJJ,noaKTJIB Hbie MOJJeKyJibl 6bICTpee npyrnx CTpy1nypHb!X 6e.'IKOB BKJIIO· 11/:llOTOl B BHpIIOH.
CHHTC3 M-6eJ1Ka MCCTKO perym1pyeTcH KaK m1pyccneun (IHnecKHMH, TaK H KJieTO'IHblMll MexaHH3M3Mll. 9KcnpeccHH re Ha M-6e.DKa HatJHHaeTcn 1103me .9Kcnpeccm1 npyn1x re11on, u M-6eJIOK Clfl!Te:mpyeTCH Ha caMbIX Il03,ll,HHX ,3Tauax penpoJJ,yK UHH. fIOCjfe CHHTC3a M-6e..noK BHpyca rpmma H napaMHKCO BHpycon 06J.1apy)KHBaeTcB n B,n,pax 3apa:i1,emrux K.rreTOK [Pat terson S. et al., 1988; Peeples M., 1988]. TpancnopT B HJJ.po nponcxo,l!,HT B OTcyTcTBHe CHHTe3a npyrnx BHpycHblX 6eJIKOB [Bucher D. et al., 1989], cJieJIOBaTeJJbHO, M-6eJ10K conep):KHT HYKJieoqrnm,11uii cnrnaJ1. B flHTon.ria3Me M-6eJIOK cBH3a11 c BOJIOKHaMH aKnrna, 3TO MO.'lKCT !IMCTb 60.fJbIIJOe 3Ha11e11He ,l],JIH c6opKH Bllpyca H 1m1rn:0BaHHn c KJJCTo,mol'i: nonepx11ocTH, Ko
.rmqecTBo M-6e.11Ka n 3apa)Kem!hlX K,!JeTJrnX: pery.nHpyeTcB KJieT01lHbIM tj)aKTOj)OM HeH3BCCTHOH npHpO,ll,bl H BO MHOroM onpe,11,eJHWT oco6eHHOCTH peTipo,n,yKil,HH B11pyca B )faHHOH K}lC TOlJHOH cHcTeMe. A6eppaHTHbli"I CHHTC3 M-6eJIKa II ero riapy lIICHHhIH BHY'fPHKJJeTO'IHLIH TpaHCIIOPT CJIY)KfiT oJJ,Hoti H3 iia cTux npH'IHH a6opnIBHLIX H nepcHCTCIITHhlX BHpycHI,lX 1IHQ)CK· uuii. TaK, MHOrlle aBTOpbl IlOKa3aJIH, 11TO rrp11 a6opTIIBllblX 1rncpeKUll5!X, Bbl3Bl:lHI!blX BHpycoM rpmma, B KJlCTKaX pe3KO cmrn(e1m rKOJJIP1ecTBO., MPHK ,vrn M-6e.1rna ficM. EyKpuH cKa5l A. 1985]. l1HTepeCH:o npe,ll.CTOJJO)](CHIIC O TOM, 11TO IIJ)Hlll!HOH a6opnrnHOH rrnqieKIJJJH HBJ15ICTC5! Bb!IIIC)J,IIllll1 If3-ITO,ll, I<J!CTO'll-Wro KOHTponn ycw1eHHW1 cnJiaikimr MPHK 7-ro reHa m1pyca rpmrna, n pe3yJJbTaTe 11ero 06pa3y10Tc5l ll36b!T01 rnb1e
102

KOJIIP,ecrna MPHK JJ.JIH Broporo npopyKTa 7-ro reHa M2"6eJ1Ka [Inglis S., 1984].. B HeKoropbrx a6opnrnHhrx CHCTeMax o6Ha PY)KeHo Hapymemre BuyrpHKJJeT01moro rpaHcnopTa M-6e.r:1Ka. HanpHMep, 3apa)f{eHHhie BHpycoM rpimrra KJJeTKH acuHTH011 KapUHHOMbl 3pmrxa rrpoJLyUHponaJJH HCHH(pCKUHOHHble BHpyc Hhle qaCTHHbl co CHH)KeHHhlM KOJIII11CCTBOM M1-6eJ1Ka, KOTO pb!H K TOMY Me 6bUI ,ll.eq:>CKTC!I 110 rnoefr CTPYKType. CraH papTI!bll1 M1-6e,TJOK npH 9JICKTporf>ope3e B flAAr BbIHBJ!}lCTC51 B BHJJ,e nnyx paCTIOJIO:iKCHHhIX PHJWM TIOJIOC (MOJJeKyJJHpHa51 Macca OKOJIO 27 000 H 24 000) IICJIC/ CTBIIC pa3110!1 CTenemr qioc¢opHJJHpoBaHHR MOJJCKYJI. M 1-6c.110K 11c1u1THoro ,rnpyca He copeplliaJJ BTOporo KOMI1011e11T,l. I lp11 :i11:1J1ll:K· Gemu1 MCTOJJ,OM OTllClfaTKOB naJlhll.CB 6bIJIO 0611apy)KCHO OTCYTCTBHC no KpaJ'i.Hei1 Mepe J{Byx omironenTHJ(OB. M1-6e.10K Bbl5!BJUJJIC5! B ITJJa3M3TH1!CCKOll MCM6paae II naKclJJJIHBaJJCSI B OKOJJOHJI.CPHOii 06-
.riaCTH [Bukrinskaya A. G. ct aI., l981]. CJICJJ.OBhle KOJJHlfe CTBa J(CqJCKTHOI'O M-6eJJKa, KOTOpble BCC-TaKH TpaHcnopTHpoBa JJHCb J( IlJia3MaTIF!CCKOH MCM6pane, HC MOl'Jlll o6ecnetIHTb KoppeKTHOii c6opKH BIIJJYCilblX 1I3CTIUJ., II OT KJJCTOK OTITO'IKO Bb!BaJJlICb HemupCKUHOIIHhle IJ3CTIIII,hl, CO/l,Cp:iKaUH:!C, ITOMl1MO BHpyccHCIJ)IQ)llqecKHX 6eJIKOB, 3Ha1IJITCJ1hHOe KOJJHtJCCTBO KJJC TO'lllblX 6e.1KOB. M1-6eJJOK B Ti!KIIX 1JaCTHI1,aX JJH6o HC o6Hapy
lliI!BaJJCH, ,!JJJOO Bb!HBJl5VIC5l |
B pe3KO |
CHIIllieHHhlX OTHOCH |
|
TCJlbHhIX K0jllJ1ICCTBaX (CM. |
lJyI<pIUICKaH A. |
f., 1985]. fipH |
|
nepCHCTCIITHhIX HI!(j)CKUIU!X B |
Kym,Type |
TKaHH, Bhl3BaHUh!X na |
paMHKCOBHpycaMH, TaK)KC 06Hapy:>KCH JJ.CqJCKTHblH BblCOKOJJa611Jlbllbiii M-6CJ1<)K, ero CHHTC3 pC3KO CHII.lKeH H JJIHIIb He6oJJb moe KOJJII'ICCTBO nnpycm,Ix lJaCTllH npoJJyu11pyeTC51 KJJCTKO!i B OKpy)Ka10111.y10 cpe,n,y. AHOMaJJIIH B CHHTC3C H rrpoueccirnre M-6e.TJKa Bb!HBJICHbl rrpH rrcpCIICTCHTHblX IIH(pCKUH51X, Bb!3BaH HblX nupycaMn CeH/tnil. u Kopn (Roux L., Waldogel F., 1982]. flpH ITO,ll.OCTJ)OM CKJJCp031IJ)YJOll(CM rraH3HIJ,eq:>amrre B KJJCTKaX UHC BhIHBJICH Jletj)eKTIIbIH M-6eJJOK BcJJeJJ,CTBHC JIH6o ero npe pnamwro CIIHTe3a, JJH6o OTCYTCTBIIH CTa6HJJbHOCTH; M-6eJJOK 6blJ] J!HlJICH qJyHKHHOHaJJbHOli aKTHBHOCTH H HC MOr 06ecrret!HTb c6opKy BHpycm,Ix '!3CTHU [Enami M. ct al., 1989).
flpwrnttbl OTCYTCTBHH M-6CJ1K3 HJl lI ero CHIDKe1rnot1 9KC npeccm, B KJICTKax CB5133Hbl, nO-BHJJ,HMOMY, C KJICT()t)HblM cµaK TopoM, o6ycJJOBmrnaJOIIJ,HM nenepMHCCHBHOCTb K.llCTOK: H3BeCT HO, l/TO 9KC1lpecc1rn rena M 3H3TJHTCJlbHO nap1i11pyeT B KJICTKax pa3HOro IIJ)OllCXO)KJJ,eHirn. TaKHM o6pa30M, CB5I3h MClliJJ,Y a6ep paHTHhIM CIIHTC30M M-6e.1ma II nepcncreHTHhlM II a6opnIBHhlM TIITiaMH nmpe1rnJIH J{a)!(CTC51 O'ICIHl;']JIOH.
Cy.n.h6a MaTpHKCHOro 6eJ1Ka npu pa3.ll,enam-m nupycoB rpun na H napaMHKConupycoa. flocKOa1bKY M-6e.r10K nnpycoB rpHrr-
103

|
Puc. |
7. |
|
CrrnttorpaMMa |
6eJIKoo |
|||||
|
cy6B11pycm,IX |
•iacrnu, |
llOl!BJil!IO· |
|||||||
|
11\HXCH npH pa3.u_ena11ttH Bnpyca |
|||||||||
|
rptmna n UHT0nJia3Me aapa)KeJI- |
|||||||||
|
11wx |
KJICTOK |
(5yKpUHCKM! A. f'. |
|||||||
|
H ,u_p., J 982]. |
|
|
|
|
|
|
|||
|
J(11erKH |
aCl\llTHoi\ |
Kapa.HR0Mbl |
::Jp. |
||||||
|
JIHxa sapaJKaJIH UHpycoM rp,mna A |
|||||||||
|
(UJTaMM KJIHCC"'iecKo!l 'IYMbl llTHU.). |
|||||||||
|
MC':'l[('Jlll!,lM l-1C·rtM)llt0KlfCJIOT3MH, '·IC |
|||||||||
|
(10,1 2 1r |
|
tHK'Jf(' |
:wpntKcrmn |
noJIY |
|||||
|
•1a1111 |
U.HT0rIJl3:hl8Tl!'lecKHl1 3KCrJ)3KT |
||||||||
|
H a.esrpnq,yrnponann era II rpaJJ.H· |
|||||||||
|
erne x11opn.lla 1 e3m1. Ee11KH pa)lno |
|||||||||
|
a1:eTHDHblX nHKOB aH3J!H3HPOB3J1J-I C |
|||||||||
|
ll0M0U\blO |
|
SJJeKTJ)0,j,ope3a B |
IJAAr: |
||||||
|
a - 6e11KH nHpyca rpnnna (nnor |
|||||||||
|
KOCTb |
B |
XJIOPlrJl.e |
qeaH51 |
1,22 r/MJI); |
|||||
|
6-e - 6eJIKH |
cy6nHpycHblX |
'13CTllll |
|||||||
|
C |
TIJIOTIIOCTbJO |
|
COOTBCTCTBeRH0 |
||||||
|
1.35 ((6), |
1,32 |
(n), |
1,30 |
(r), |
1,29 |
(n) |
|||
|
H 1,28 (e) |
r/Mn. |
|
|
|
|
|
|||
|
na |
H |
|
napaMHKCOBHpycoe |
||||||
|
B3aHMo,neiicTByeT |
KaK |
C |
|||||||
|
HYKJICOK3I1Cll.ll.OM, T8K ll Jlli |
|||||||||
|
IlO!lpOTeH,ll,HOH |
060J1011KOH, |
||||||||
|
BO3Hl1K3CT sonpoc O TOM, C |
|||||||||
|
KaKHMlf CTPYKTypaMH oH co |
|||||||||
|
xpa,rneT |
CB513b |
npH pa3,ne |
|||||||
|
namrn |
BnpyCHoii |
tracTHUbl. |
|||||||
A |
011CBHJlHbl |
|
;rne |
aJibTep- |
||||||
HaTJIBlll,!C |
|
13O3MO2KHOCTH: |
||||||||
|
l) |
M-6cjlOK |
|
coxpam1eT |
||||||
|
CflH3h |
C J]Ill10flJ)OTeH,nHm1 |
||||||||
|
0O0,H)ql{OH II npH ee CJllUl |
|||||||||
|
HllH |
C |
MCM6paH8MH |
3H,llO |
||||||
|
IU!T3pHblX |
BaKyo.11eti |
BMe |
|||||||
|
CTe |
C |
|
r.'IlIKOnpoTCH,ll,aMH |
||||||
|
OK33blBaeTCH |
|
13 |
COCT3Be |
||||||
|
K.!ICT01JHLJX |
|
MeM6paH, |
B |
.':)TOM CJiyYae HIHjlCl<UHOHHa51 CTpyKTypa nnpycou, HYKJJeonpo• TeHJI,, BblCB000)K;{1:1eTCH B UHTOITJia3My; 2) M-OCJ!OK COXJ)8HHCT CBH3h C HyKJICOK3TICH)lOM l1 B KOMTI.llCKCC C !HIM BbICBOOOlli· ,nacTCH B UUTOIJ.i!a3MY TIOCJJe CJ11rn1urn B!Ipy,Hoti OOOJlO'IKH C KJ1eT01I11oir MeM6paHOH. B noM cj1yqae npouecc pa3,nesaInrn ycJIO)KH.HeTCH, nocKOJJbKY M-6eJioK /lOJI)l<CH 61>JTh y/laJieH c no n0pxHOCTH llYKJ1eOK3ITCH.ll.OB.
1'lccJ1eJ(oBamrn, npoBe,neHHhie n Hamci1 Jia6opaTOp1rn, ITOk TBcp;u1.rm nrnpy10 Bepcn,o: rrpll :rnpa>Kemm KJJeTOK supycaMH rpnnna n rraparpnnna '!CJIOBeKa nma 3, MecieHHbJMH aMHHOKHC J10T,IMif, o u,rf'fonJTa3Me oom1pyi1-:11BaJrnc1, cTpyr<Typ1.1, CO/lCJJ)I<a-
104

rr.u1e 6eJtKH NP H M (pHc. 7). 3nr CTPYKTYPhI OTJrnirnJIHCh ot HYKJleQl«lIICl!JJ,OB 66JibllllIMH pa3McpaMII (OfHI OLICTpee CCJJ,H MCHTl!pOBam1 B rpa,ll.HCHT3X KOIIIJ,CIITJ)aIJ,HH I'JUIU,epmia) lI Me!IbllJCH f!JIOTHOCTblO B XJIOPHJI.e llC3HH B CBH3H C H3Jllfl!HCM M-6eJIKa (p 1,27-1,29 r/MJI). I1oMHMO HHX, BWIBJI5!.llHCh TI1- ]UJl1HblC BHpycHhie 11yK.rreoKancHJJ,bI. Bepo.HTHO, cy6BHpycm,1e CTPYKTYPhI HBJIHlOTCH IIpoMerKyT011HLIMH B pa3,11,eBa1rnH BHpy c<rn; Ha Il0CJ1CJI.YIDIIJ,e{1 CTcl,/1,lIH M-6CJIOK ,ll.l!CCOUHHpyeT C rro BepXHOCTH HYKJJCOKanc11,na, H cuo60JtHLIJI HYKJleOK3TICHJJ: rrpH o6peTaeT B03MO}KHOCTb Bbl3BHTh 11mj>CKUHOHHLlH npouecc [6yK pHHCKaH A. r., lllapoBa H. K, l98t!; Bukrinskaya A. G. et al., 1982]. TaKHM o6pa30M, JHl3JlCirnllIIC BHpycoB, COJiep,KaIIJ,HX M-6eJlOK, JlOJrnrno npeJI.C"raBJIHTb co6oil JJ.BYXCTa;1,11HHbIH npo uecc: Ha nepBOH CTa,n:1rn pa3JICBal!H.H rrpO!ICXO)lHT y,uaJICH!IC
.rumonpoTeHJV!OH 060JI011KH H BhlXOJI. B UHTOnJia3MY cy6mrpyc HLJX L!3CTIIIJ,, a Ha BTopoi'r CTaJlHH - yJtaJJeHIIe M-6e.1ma H Bbl CB0601KJ1eH11e HYKJ1e01rnncn,na. IIo-BHJl.HMoMy, o6e CT3Jl.HH n K.rieTKe «TeppIITOpHa.libHO» pa306IUC!lbl. BbICOKHe KOHUCHTpa
l]JiH M-6e.11Ka 6hIJlH 0611apy}KeHhl B COCTaBe rJiaJlKHX MeM6paH |
|
: apa,KeHHhIX BHpycoM |
rpmma K.IleTOK [6yKpHHCK3H A. r. |
H Jtp., 1982]. Bo3MOiKHO, |
.n.nccou,HaUH5! M-6eJIKa c noBepxHOCTH |
HYKJleOKancn}la Ilj)OllCXO)lHT B pe3yJ1bTaTe B33liMOJJ:eil:cTBH5! 3TOro 6eJ1Ka C KJie'ro•mI,IMll MCMopaIHJMII, o6yc.nOBJICHHOfO .IIll HOcjHI.TthHl,IMH CllOl0ICT!:laMH 6eJIKiL B:iat JMOJieiicTBHC M-oeJIKa C
MCM6paHaMH 6hIJIO 110Ka3aJIO B |
3KCTICpIIMCIITC rrp11 HCTIOJlb3(,. |
||
Ba!llrn 6HC.110l1HblX MCMCipaH |
lI |
.II!IIJOCOM [BopKyHoBa r. |
K |
11 .n.p., 1983; KpaBUOB U. B. |
H JI.p., 1984]. |
|
|
nepn1flmaSI CTPYKTypa MaTpHKcuoro 6eJ1Ka nupyca rpunna. |
|||
Y BHpycoB rpIIrrna MaTpHKC!lblH 6e.IJOK (Mi ) KO,LI.HpyeTC5! |
7-M |
reHOM. 3ToT 1Ke reH, KpoMe M1 |
-6e,1Ka, Ko.n.npyeT HCCTPYKTyp |
HblH Mr6CJIOK: KO)J.llpy10111.crn |
IlOCJIC)J.OBaTeJihHOCTb llMeeT |
TOT }Ke llHIIUHHPY!OlUIIH KOJJ:OH, |
HO B pe3y.IJhT3Te CilJia1k1rnra |
yJlamICTC.H 689 llYKJICOTIIJ}OB, II HYKJJCOTll,ll. 51 npHMbiKaeT K HYKJieOTHJl.Y 740. CIIJiaiICttHr co11e1aeTC5! co c,nB11roM paMKH. 6e,1KH M1 H M2 nepeKpbIB3KlTCH Ha 14 aMiiHOKHC.'lOTHhlX OCT3T KOB co CJlBIIraMH paMKH Ha O)l.lIH HYKJICOTH,lJ., B pe3y.IlhT3TC
qero 6eJIKH HC HMCIOT o6IUHX TIOC.IJeJ}OBaTCJJhHOCTei'r aMIUIO |
||
KIICJIOTHblX ocTaTKOB [Lamb |
R, Lai C.-J., 1981]. |
|
MoJ1eKyJia M1-6e.1KoB n11pyc0B rpHnna THTIOB A 11 B co,ll.ep |
||
lKHT COOTBCTCTBemw 252 |
H |
248 aMHIIOKHC.IIOTHblX OCTaTKOB |
[Briedis D. et al., 1982], M-6e.1Ka BHpyca naparpHnna 11eJ10- |
||
BeKa rnna 3 - 354 ocTaTKa |
[Galinski M. et al., 1987] II |
|
M-6e.nKa Bnpyca CenJiaH |
347 ocTaTKOB. MoneKyJia 6eJJKa |
BK.IIIOl\aeT HeCKOJlhKO nenpepbIB3IOIUHXCH TIOCJIC.lJ.OBaTeJlhl!O· CTeil. OCT3TKOB HeHTpaJJbHhlX II rHJipOq)06HblX aMHHOKHCJIOT,
105

KOTOpble o6yCJIOBJIHB3lOT JIHTIO lfJlbHOCTb 6e.'1Ka,Bb!C0KYIO cte
neHb r11,11,pocpo6HOCTH, pacrnopHM0CTb B opraHW!eCKHX pacTB0· |
||
p.11Tem1x. B M-6em,e |
1rnpyca Cett,ll.aii KoJ111<1ecrno aprmmrrn |
|
H JIH3HUa C0CTal3JI51CT |
13% MOJJCKYJ1HJ)HOli M3CCbl 6eJIKa, 6eJIOK |
|
C0JJ.CpiKIIT T0Jlbl{0 11 % u-crmpaJICH H 20% |
r,-npyrnyp, npH |
|
3T0M 18 H3 32 aMHH0KHCJI0THblX 0CT3TK0B |
13 o6JUICTH crrnpa |
JJH33UHH H3X0JJ.51TC5! B Or/J,HOH a.-cnHp3Jlll [Blumberg B. et al., 1984]. '.:ha 06.:JaCTb, no-BHJJ.HM0My, 5113JIHCTC5l B32KHOH )leTep
MIUiaHTOH, onpe,ne.rrn10J..UeH BTop1:11my10 crpyKrypy M0JJeKyJiu CeJIKa. 13.r:iaro,n:apHBbJco1<011 cTeneHH rn,ll,poqio6HocTH M-6eJJOK,
e,n:HHCTBCHHbIH H3 CTPYKTypHblX 6eJIK0B BHpycoB rpmrna, MO· )KeT 661Tb 3KCTparupoaaH H3 BHpH0H0B KHC-llblM XJJOpoqiop MOM - MeTaHoJroM [Gregoriades A., I973].
CyJ..UecrneHHaH 11 CJI0:>KHaH cpyHKUHH M-6eJrKa B Mopcpore nese Bttpycoa ,n,o.'DKHa onpeJJ.eJieHHUM o6pa3oM perym1poBaTb C51 B 3apa:>KeHI-IOH !(JICTKC. OJJ,HOH H3 T8KHX ¢opM peryJrm.um
51B,15JeTcH rpoccpopHJIHp0BaJ-JHe.KaK oKa3aJiocb, M1-6e.rIOK B co CTaBe BHPH0Ha rp1-mna H B 3apaJKeHH0H KJJeTKe cyJ..UeCTByeT B ABYX cj)opMax, 0TJIH9310ll(IIXC5! no fI0JJ,BHJKH0CTH B flAAr. Ehn (popMbI oKa3aJIHCb cpoccpopHJrnpoBaHbI: B coCTane Bepx• HCH IT0J!0Cbl - 1 10 cep1rny, HH:>KHCH - no cepHHY H Tpeom-my [Gregoriades A. et al., 1984]. <Poc¢op1rnttpoBaHHC M-6eJJKa 6b!JIO IT0IW33H0 11 y JJ.pyrnx BHpyc<JB - napaMHKC0BHpycoB H pa6,ll,OBHpycoB [cM. EyKpHHCKafl A. r., 1985]. <PoccpopHJ!HpO· 13aHHe M0)KCT 6b!Tb TCCH0 CB5133H0 C perymur,uefi ero cpy11Kll_Hl1 npn pa36opKe H c6opr<e n11pyctto11 tJaCTHUbl, K0HTpoJ1»pyj1, ua npHMep, B3aHMO)],CHCTBHe C JlHTIH)],aM!I H HYKJ!C0K3ITCl1)],aMH H onpe/.1,eJIRH JI.3JlbHCllllIYIO cyJI.h6y HYKJ!C0K3TICHJJ.3 - J!l160 KaK MaTpHll_LI JJ,Jlfl TpaHCK!)HTIUflll reH0M3, JJII6o K8K c-rpyKTypHoro
1<0MnoHeHTa noqKyromeiicH BHpycuofi 1JaCTHI b1. KoucepsaTHBHOCTb MaTpHKCHoro 6eJ1Ka. <l>yHKUHl-1 M-6eJ1-
1rn, KpHTHL!eCKH B3)KHf.lC J.(JlSl MopcjJOreHC3a, C03peBaHI:fSl BH· p»0H0B JI BLIX0,Qa HX H3 KJieTKH, npeJJ.TI0JraraI0T BLIC0KYI0 K0H cepnaTJIBH0CTb ero nepBHlIH0H CTPYKTypb!. ,UettCTBHTCJlbHO, M-6eJJOK 0l<333JlCH HaH60,1ee K0HcepBaTHBHbfM 6eJIK0M y BHPY· COD rpnnna H napaMHKC0BHpycoB, npeB0CX0JJ,51 no K0HCepBa THBH0CTH JI.pyron BHyTpeHHHH 6eJJoK NP. Y B11pycoB rpanna A roMoJiorirn 7-ro retta ,uocT11raeT 91-92%, a caMoro 6eJ1Ka - JJ,O 97 % . 3rn THnocneumpw.1ecK11 i attrnreH, 06rn:i1ii ,DJrn mIpy c0B nma A, B H C. I-fa 744 HyK,1eoTH)l,oB, K0JJ.HpyIom11x MaT·
pHKC11bJH 6eJIOK, 289 RBJHIIOTCfl K0HcepBaTIIBHblMH, |
a H3 252 |
||
!I |
248 |
3MHHOKHC.'!OTHblX OCTaTKOB BHpycoB rpunna |
THD0B A |
ll |
B 63 |
0CTaTJ<a r0M0JJOrl1111ff.l. O)lH<ll<O f0Mo,;1or1I1 mb1e 0CTaTI<H |
pacnpe,ueJICHbl B pa3HblX ytJaCTKax 6eJJKOBOil M0JleKyJlbl H KO· JUl'!CCTB0 noc.rre)l0BaTeJibHbIX roMOJ1orrrqHblX aMHH0KHCJI0T He
106

npeBhrrnaeT 3 [;Briedis D. et al., 1982]. Ilp11 HCII0Jib30Bamrn MOIIOCIICUHqHl4CCKIIX aHTl!CbIB0p0TOK rrponrn M-6CJ1Ka C rro M0JUb!O HMMYlIOCjJepMCHTIIOfO a11amrna (I1¢A) H HMMy1rn6J10TH!ffa 6blJIH BblflBJICHI,J rrepeKpecnH,lC peaKUllII MC)l{AY MaT pHKCHblMH 6e.rmaMH m1pycoB rpHnna Tima A H B, anT11MaT
p11KCHble Cb!B0J)0TKH |
B3aHMOlldicT130133.IJH |
C reTep0J10fH4Hb!M |
||
M-6CJ1l{OM IIJ)H I1¢A |
,ff,0 |
¼- 1/s Tl!Tpa |
[EyKpHHCKaH |
A. r. |
H AP-, 1984; Bukrinskaya |
A. G. et al., 1985]. CJ1ell,yeT |
0TMe |
THTb, 41'0 K0HCepBaTH3M0M o6Ml/l310T 11 J].pyrne BHyTpeHHHe 6eJIKH s1-1pycos rpmma, P 11 NP, BbrnomrnromHe BaJKHbJC (py1m-
1urn npn TJ)3HCKJ)HIIUHH ll pe11mmmurn l'CH0Ma. TaK, y 6eJIKOB |
||
PB 1 |
BIIpycoB rpmma Tlllta A II B roMo.rronJ1 m1>1e aMHH0KHCJIOT· |
|
Hhle |
0CTaTKll C0CTaBJHll0T 61 % [Kcmdirim s. |
et al., 1986], |
a y 6cJmos PA o6uapy)l{eIIo 38% roM0JI0rIIH |
[Akoto-Aman |
|
fu E. ct al., 1987). |
|
BbIC0KaH CTCIICHb fOM0JI0fHII 3MliH0KHCJ10T 6bIJ13 061rnpy )l(CHa y M-6CJ1KOB napaMHKC0BHpycoB. IlpH cpaBHCIIHH Jl0CJ1e )1JJBaTCJlbHOCTCH aMHH0KHCJ10THblX 0CT3TK0B B M0J!eKyJiaX oc H0BHblX cTpyKrypm,1x 6e.11K0B N, P/C, M, HN u F BHpycoB
3 po,noB caMaH BbJC0KaH r0Mo.11onrn 6uJ1a o6napy)l{eHa y M-6eJrKOB [,Galinski M. et al., 1987]. rip11 cpasnemm nepBH4·
Hofi CTJ)YKTyp1>1 M0JieKy:1 6e,'lKOB NP n M BHpycoB Ce11J].aii 11 naparpmrna lJCJI0BeKa T1ma 3 oKa3aJl0Ch, 4T0 B 6e.r1Ke NP roM0JI0flP!llblMH 61,rJIH 70% H:l 387 aMHH0KHCJ10THb[X 0CTaTK0B C N-Konua M0JleKy.1161, 13 TO BpCMH KaK HH3Kcl5l CTCIICl!b f0M0· JlOl'HH 6bJJ1a o6Hapy.tKeHa C C-KOHUa M0JleKyJibl. foMOJIOl'H4-
Hb[C aMHH0KHCJl0THhle 0CTaTKH C N-KOHUa nax0}l,HJIHCb B TI0J!O· )l{emm 1- 133 11 160-386 [Galinski M. et al., 1986]. IlpII
cpas1rnTeJ1bH0M aHaJ1H3e 6eJJ1<0B NP snpyca naparpmma 11e J10BeKa nma 3 H BHpycos K0PH II 1IYMbl co6aK BblHBJlCHa He3Ha'IHTCJlbllaH r0M0JI0fHH. O,nnaKO neoOJib lllHe rpyrnibl, BKJII04a!OJUHe 0K0JI0 7 0CTaTK0B B o6JJaCTH 166-383, 0Ka3aJIHCb K0HCCpBaTHBHblMH. Y pecn11paTopH0·CHHUHTHa,11,uoro BHpyca roMoJrornH B 6e.rrKe NP o6napy.tKeHo 1-1e 6bJJIO.
flpH cp3B!ICHHI1 aMHH0KHCJ10THblX IIOCJ!CJ].OBaTCJlbllOCTCH M-6eJma BHpyca naparpnnna 11e.rroseKa nrna 3 c T3K0BbIMH ,npyr11x napaMHKC0BHpycoB 0Ka33J10Cb, lJT0 HaH60J1bl113H CTe lICHb fOM0JIOrIIH 6bIJ1a C m1pycoM Cen,naii (61 % ) H HclHMeHI, lllaH - c BHpycaMH K0pH H llYMbl co6aK (35%). KoncepBaTHB· lib[ HCK0T0pble aMHH0EHCJ10THble II0CJlC)l0BclTCJlhH0CTH, T3KHe K3K 21/24 f.fJHUHII, 17/19 npoJlHH, 4/5 UHCTCHH, yqaCTBY!OIUHC B KOH<popMa1 1m Mo.rreKyJ1b1 M-6CJ1Ka. Hc:rna111ne.,r1,11aH roMo J1or1151 0blJia o6Hapy)KeHa C pcc11npaT0p110-cmrn)ITIIaJll,HJ,IM Bii pycoM [,Galinsky M. ct al., 1987]. l1Mc10TcH H rrpOTHBorroJJO)!< Hble ; amlble - 06 aHTHreHHoii m1,nns1v yam,II0CTll M-6eJJKoB
107

0 |
Ta (j JI HI.I, a 16. OepeKpecrnble |
peaKl.l,HH MeJK,11,y |
napaMHKCOBHpycaMH B HMMYH00,10THHre |
|
|
||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
B11pyc |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
||
|
AHTHChlBOpOTKa |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Bllf'-1-1 |
I |
Bf1fll-2 |
I Bllf'l-3 |
|
I |
naponna |
I |
CeH,lall |
I !Wp!! |
|
|
|
|
|
|
|
|||||||||
|
|
|
|
|
|
|
BTif4-l |
(Mop• |
Bee |
NP, M |
NP, |
M |
M |
Jl |
||
ClWil eBHHKH) |
|
6eJIKH |
|
|
|
|
|
|
||
ripoTHB BTIP.J-2 (Mop- |
NP, |
l,1 |
Bee |
|
NP, |
M |
M |
M |
||
CEOH eBHHKH) |
|
|
|
6e.rIKH |
|
|
|
|
||
|
BTif4-3 |
('le- |
NP, |
M |
NP, |
Ni |
Bee |
M |
M |
|
|
|
|
|
|
|
|
OC'1KH |
|
|
|
ITpoTHB |
B11pyca |
napa- |
M |
|
M |
|
M |
|
Bee |
M |
rpimna |
TITHU |
|
|
|
|
|
|
|
6e.1Kl1 |
|
|
BHpyea |
napo- |
NP, |
M. |
NP, |
M |
NP, |
M. |
M |
Bee |
THTa (tJeJioBeKa) |
|
|
|
|
|
|
|
|
6eJIKH |
|
fipOTHB |
Bttpyca |
CeH- |
M |
|
M |
|
M |
|
NP, M |
NP, lv1 |
,ll.att |
|
|
|
|
|
|
|
|
|
|
Bee 6eJJKI!
NP, M |
0 |
,\1 |
|
Bee |
|
OeJJKH |
|

napaMHKCOBHpycoB 4CJ10BCKa H 2KHBOTHblX [Ito Y. et al.,
aKBpCHe
Mu a3yqaJ1H nepeKpCCTHblC peaKIVIH MC2K,11.y napaMHKCO
BHpycaMH, HCIIOJ1h3YH IIMMYH06J10THHf. 13eJIKH QlIHI.UCHHblX BH |
|
pycoB naparp1mna 'lCJJOBeKa (Biff4) rttnoB |
I, 2 II 3, napa |
, rp:ttnna IITHU:, napoTirra H CeH,naA pa3Jl.CJIHJIH |
nyTeM 3JICKTpo |
.tj)opeaa B 7,5% TTAAr, nepCl!OCHJIH Ha IIHTPOUCJIJ1!0J103Hb!H (J)HJlbTp C IJOMOI.Ub!O 3JICKTpHl!CCKOro TOKa H MaTepHaJI o6pa6aTbIB8JIH 8HTHCbIBOpOTK3MH nponrn rrapaMIIKCOBHpycoB, rro- 6!ylJ.eHHblMH OT peKOHB8JICCI(CHTOB H npli HMMYHH331].HII MOp CKHX CBHHOK H KpomIKOB ('ra6.11. TpOKpnc. BHaKKaK BIIJJ.IIO H3 Ta6J1. 16, nee 1mpyc1,1 1rnparp111ma TJCJIOBCKa HMCI<lT 06m1Ie aHTHrc1rn1,1c ,neTcpMIIIJaHThI B cocTanc 6e.r1KOB NP II M: aHTH· CbIBOpOTKH ncpeKpecTHO B3a!IMOJl.ClfCTBYIOT C o60HMH 6e.rrKaMH. AnrncbrnopoTKa npoTirn BTTf4- HOB3aHMOJl.CHCTByeT co BCCMH 6eJ1KaMH BIIpyca CeHJ1.aH, II SITO noJ1.TBep2KJI.aeT rrpmrn1'l.JIC}K• HOCTb o6crnx BHpycoB K OJl.HOMY Timy. EeJIKH NP H M BHpycoB
rraparpmma nTHU: u |
Cen;iati |
nepeKpecTHO B3aHMOACHCTBYIOT |
C aHTHCb1BOpOTK8MH |
npOTIIB |
o60HX BHpycoB, aHTHCblBOpOTKH |
TlpOTHB napaMHKCOBHpycoB 4CJ10BCKa H )KHBOTHblX nepeKpCCT· HO B:-HUIM0,11.CHCTBYl()T C M-6CJ1K3MII ncex HCil0Jlb30BaHHblX na paMHKCOBHpycoB, HO HC B3aHMO)'.l,CHCTBy10T C M-6eJIKOM BHpyca KOpH.
BbICOKHJ'.\ KOHCCpBi1Tll3M M-6C.'1Ka 1103BOJIHCT npCJl.TIOJIO )KHTb, '!TO 3BOJJIOIUIOHHaH mrncprcmurn cro WJia C HHbIMH 33KOHOMCj)HOCTHMH no cpairnenmo c TaKOBOH JJ.pyrHX 6eJIKOB. TTo-BII)lHMOMV, SITO 11a1IMCHCC TIJHJCTHH1HblH 6e.lIOK, HCC!lOC06HblH K coxpaHCHHIO CBOHX cj)yHKll,l!H Jl.3}KC npn HC3IrnO1lfTCJlbHblX MYT8UHHX. KoncepsaTll3M 3MHHOKHCJIOTHbIX fIOCJICJI,OBaTCJlbHO· CTCH y JJ.BYX pOJl.0B CCMei\crna YKJI3Jl.bIBaCTC51 B Ilj)C,II.CTaBJlCHHe 0 KJIIOlJCBOfl j)OJIII M-6eJJKa B C03penaHHH H c6opKe BHpycoB. TTospe2KJJ,CHHH B npe,11.eJJaX SITOro reHa TCCHO conpH2KCHbl C B03Hl1KHOBCHIICM a6opT!IBHb!X Ii nepcHCTCIITHblX H!!(pCKIJ,Hli.
BbICOKllH KOHcepBaTH3M MaTpHKCIIOf'O 6CJ1Ka !I 3Ha•rnTCJib· HblC ero KOJJ!]lJCCTBa B B llpHOHC OTKpbIBaIOT nepcneKnmy ;'l,,'lH l1CllOJlb30BaHHH ero C ll,CJlbIO AH8fHOCTilKH rpllTITI03HblX H napa MllKCOBHpycHb!X IIH(pCKll,HH. HaMll pa3pa60TaHa BblCOKOl!YB CTBHTC.lJbHaH TCCT-cHcTeMa H<I>A JVI51 Bb151BJICHHH M-6e.rrna BH·
pyca rpmrna B |
IIOCOI'JIOT04Hb!X CCKPCTax 60JibllblX, OCIIOBaHH35l |
|
Ha KOIIKypeHTHOM sapHaHTe [BopKyHoBa |
OHaIf ,11.p., aKBTCOe |
|
TecT-CHCTCMa |
H03BOJIHCT 061rnpy2KHTb |
Jl.O I Iff M-6eJ1Ka |
C aKKaer. TCCT·CHCTeMa rrpOH3B0,i{HTC5! K3K KOMMCj)lJCCKHJH! rrpe napaT.
109

4
l
j'\\ ···1(
M J