
Медицинская вирусология / Букринская А.Г., Жданов В.М. Молекулярные основы патогенности вирусов
.pdf
TYP no MexaHH3MY caM0Y3Hanamu:r. BHpycm,re K0MTI0HeHThl HCII0J!b3YI0T pa3IIblC ny-m TpaHCI!0pTa, 1IT06LI Jl0CTIF!b JJOKy
C0B, B K0T0ph!X rpopl\rnpy10TCH lllIYTJ)CIIHiiC CTpyKrypu (cepA IJ,CIHIIIhl Ii HYKJICOKancrt}lbl) Ii 3pC.l!Lie lliipycnr,Ic lJ8CTHUbl.
Y H,r1,cpm,1x m1pycon 3Tanb1 MOptllorc11e3a ocy113.ecrnJimoTcH B HApC, KyAa Ii3 UHT0DJia3Mbl TpaHCD0J)T!Ipy10TcH BH0Bb CHHTC-
3HpoBaHHhle 6eJIK!i. y UIIT0I1Jia3MaTIJl!CCKIIX nupycoB penm1-
Kau1rn 1rnpycHhIX re110B H Bee 11poueccu, CllH3aum,rc c cospe BaHHeM BliPYCHhlX 1!8CT!IJJ,, Ilj)0llCX0AHT ll IJJIT0IIJia3MC. O6Ll'I
HblM MeCT0M c6opKII OUOJIO'IC'Illb!X rmpycon HBll5ICTCH II,l!a3M3Tlfl!CCK3H MCM6pm1a, Ila llllCIJJIIJ0I0 Jl0llCJ)XH0CTb l!C
TJ)clllCll0J)THPYI0TCH 6emrn 11apy21u11,1x 060JI01 JCK rJIHK0npo
TCHALI, CIIIITC311J)YI0IU!lCCH 11a pn6ocoMax, CJ3H3aHIU,IX C MCM·
6paHaMIL C mryTperrnel'1: CT0p0Hb! ll0,ll.X0AHT cep/lJ CBIIHbl t1
IIYKJIC0KallCHAL>I, BX0AHIUHC B C0C'l'3B 06pa3yIOIUCHC5I Jl()lfKH.
O,n.H3KO M0JreKyJmpaue MCXaHH3Mbl, Jle)KaIUIIC B 0CH013C TJ)3HC norna BHpycm,IX K0MII0l!CI!T0B K TIJia3MaTH'leCKOH MCM6pane, H3 I HT0mia3MU B H/ j)0 H 113 HAJ)a B I HT0IIJJa3MY, H3Yl!CHbl RB HO HeAocTaT0'!H0. KaK 11 Ha Apyrnx cTafiHHX H11cj1eKu1rn, cTpa
TerHH BHpyca uarrpas.11ena Ha lJCII0JJL30BaHJle KJJCT0tJHblX MC Xallll3M0B. TpaFICIIOpT K IIJia3MaTIFICCKOH MeM6paHe ceKpeTop llbIX 6eJIKOU C0llJHDKCII C K0TlJilIICJIHUII0IIHUM BCTpa1rnaHHCM
6CJIKa B MCM6paur,1 311/ 0IIJJa,lM:lTll1JCCK0J'0 |
6miro |
AapH uaJIIJllHI0 ua N-KOIIJJ,e TI0JIHIICIITI!)Ul |
CIIrHJJJL,IJOl1 lI0CJlC· |
A0H3TCJ!bH0CT!l. BHOBL, o6pa3oBamwfi 6eJI0K ) l3IDKCTC5l I{ an napny I'OJlb}l)KH Ii OT Hero - K DJ133M3TH'lCCKOii MCM6pa
He. B npouecce Tj)allCII0pTa ocyn3.eCTBJIHIDTCH pa3Jllfl!HUe K0Tpa11c.rrnuH0HHble II nocnpa!ICJl.HUH0HHblC MO,D.Hcj)I1Kaum1 (nrn K03HJIHponam1e, ar HJ!Hposa1rne). HaJIHllllC yrJICB0AHL>IX UCII0tJeK cyru.eCTBCIIH0 ;vrn Tpancuopn1 rJIIIK0ITj)0TCHA0B K ll,1a3Ma Tlfl!CCKOH MCM6pa11c, poJIJ, yrJICB0JOIOI'0 K0Mfl0IJCIITa 3aKJ]I0· •iaeTCH s npH,ll.:.lllllH 6eJII<OUOi'i M0JICKYJIC npaBIIJibHOii K0!Hpop Mau1rn [Pitta A. et al., 1989]. Bupycuue LilHKonpoTcHpu TpaHcnopTnpy10TCH H<l KJICT0'IHYIO rronepxH0CTb II0 MCM6pa HaM 311,ll.0ITJia3M3TH'lCCK0ro peTIIKYJiyMa T3K)KC 6JJaro;:i_apH na Jll1!HI0 Jil,ll.CJJHOH ll0CJICA0BaTC.lll,H0CTII aMHH0KHCJIOT rm N-KOH ue II0JlliIJeIITHAa. CllniaJibHbIIi nel!TIIA rJUIK0TIJ)0TCHA0B C0CT0- l!T ll3 0CT3TK0B rHAj)Ocjlo6HhlX aMHH0KHCJIOT, K0T0J)LIC 1rncT0 qrnaHKHpyIOTCH 0CTaTKaMJl 0CH0BHblX aMHIIOKHCJIOT. B nacT0H IUee BpCMH rrpocCKBClIIIJ)0l3aH0 60.r1ee 200 rrpo- H 3yKapII0TIFIC CKIIX C!IrHaJlbllblX IICIITHA0B. Om! 0Tpe3aI0TCH OT ll0JIJIHCDTH
Aa-rrpeAlllCCTBCHIU!Ka B M0MeHT TpaHCJ10K3l\IilI accoum1poBaH
HblMH C MeM6paHaMII nporea3aMH (c11rHa.1a3aMH).
O.n.HaKo A0JILKCH cyiuecTB0B3Tb eiue KaKOH-TO A0ll0JIHII
Te.nhHbIH MCX3HH3M TpaHcnopTa BIIpycHL>IX 6e,1KOB K llJia3Ma-
91

|
|
|
|
o1l |
Cll aOpl1l |
Hcel BHKTlTTM!l |
rT11o1orrp1rbT6Rl |
|
|
|
|
|
|
|
|
|
||||
|
|
|
|
aHeO |
|
OKHeaHp |
|
cBTrH HaCToeOpplKC |
|
HKOHeap |
|
|
|
|
||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
||||||||
BTif |
|
Pamme |
rem,1 |
ICPO, |
ICP4 |
flpOMOTOpbl |
|
co6cTBellllblX |
Ycn,1eHHe |
rpaHCKp!Inl\HH, |
||||||||||
|
|
|
|
|
|
|
reHoB, |
BHll |
H |
J.Lp. |
|
peaKTHB3UHH |
Bllpyca |
npu |
||||||
9mllTeiiHa- |
PaHHl!e |
reHbl |
EB!, |
R |
|
|
|
|
|
|
|
JJareHTHO)! l!HqJeKUIIH |
|
|
||||||
fipoMOTOpb! |
|
co6cTBeH!!blX |
AyroperyJ15!1JHll |
|
TpaHc |
|||||||||||||||
5app |
|
|
|
|
|
|
reHOB |
|
|
|
|
|
Kp11rrn1rn, rrepeBOJ.L |
:1areHT· |
||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
HOH l!H(j}eKIJHH |
B |
,111THtJe |
||||
ITceB)lo6eweH |
CBepxpanH11e reHhl |
|
|
|
|
|
|
|
|
|
CKYJO |
|
|
|
|
|
|
|||
CBepxpaHHHe 6en |
To ,Ke |
|
|
|
|
|
AyroperymllJ!!H, |
yc1rnemie |
||||||||||||
cTBa |
|
|
|
|
KII |
|
|
|
|
|
|
|
|
CBH3HBaH11H |
)],HK |
c |
1,.1e |
|||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
TO'-!H61M11 3KTHB3Topaz-m |
||||||
AJ:1eH0B11pycu |
Ela |
|
EIA 289R |
ITpoMOTOpb! |
|
COOCTBeHHblX |
Yc1rnem1e |
II |
perym1n1rn |
|||||||||||
|
|
|
|
|
|
|
II |
1rneT01lHbIX reHOB |
|
TpancKpHllljllH, |
TpaHcqiop |
|||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
M3Ullll |
HenepMHCCHBHo!X |
|||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
K,1eTOK |
|
|
|
|
|
|
fenanna |
B |
X |
|
X |
|
flpoMOTOpb! |
|
,ilpyrnx |
Bil- |
Ycu.1eHHe |
TpaHCKpHlIUHll, |
|||||||||
|
|
|
|
|
|
|
pycoB |
(B114) |
u |
K,leTKl! |
rrepeBOJ:( |
JiaTeHTHO!l |
IIH· |
|||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
cjJeKUllH B |
,1llTH'!eCKY!O |
|||||
Tiamrn,10Mb! 'le- |
Pamme |
reHbI |
E7 |
TipmrnTopb! |
|
co6cTBeHHbJX |
Yc11,1eHHe |
rpaHcKp1rnumr, |
||||||||||||
,lOBeKa |
|
|
|
|
|
|
ll |
K,leTO'!llbIX |
reHOB |
|
TpaHctjlopMaU!lll |
|
KJ1eTOK |
|||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
KO)KH |
|
|
|
|
|
|
OB40 |
|
Pamm11 |
reH |
Eo.1bwoit T-arnuren |
ORI mrpYc11oi"r JlHK, K,1e- |
AyTopery,1HUl!ll |
|
rpaHc· |
||||||||||||
|
|
|
|
|
|
|
TO'-!Hhie |
JJ,HK |
11 |
6e,1KH |
|
Kpnnu1rn |
II |
perr,111Kaum1, |
||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
TpaHccjlop:vial\HH |
KJ1eTOK |
|||||
51,1'-!beil |
Ilal!H,1· |
To )Ke |
|
E2 |
|
ITpOMOTOpbl |
|
co6crBeHHblX |
To JKe |
|
|
|
|
|
|
|||||
J10Mbl |
|
|
|
|
|
|
reHOB, reHOB I1JI2 n I1J13 |
|
|
|
|
|
|
|
||||||
HTLV-1 |
|
tax |
|
Tax |
|
TRE B |
LTR, |
npoMoTopr,1 |
YC!l,leHHe |
TpaHCKPHlllJHH, |
||||||||||
|
|
|
|
|
|
|
KJieTO'!HhIX |
reHOB |
|
rpancqiopM3llllH |
|
K.'!eTO'l |
||||||||
51,1'-!hell JJefiKe- |
tax |
|
p38 |
|
flpOMOTOpbl |
|
co6crnem11,1x |
llblX reHOB |
|
|
|
|
|
|||||||
|
|
|
1'CIIJ1eH!le |
rpancKpllllI lliI, |
||||||||||||||||
Mill! |
|
tat |
|
Tat |
|
H |
K. eTO'!HblX reHOB |
|
,TpaHCqJOpM31JllH |
K.'!eTOK |
||||||||||
B114 |
|
|
|
TAR B |
LTR |
|
|
|
Yc11.1e1me |
Tpa11cKp11rrnnH,, |
||||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
nepeao.u |
JiaTeHTHoll, |
HH· |
||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
cjJeKllHH B |
JIHTH'!ecKy!O |
|||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|

Tlfl !eC1<0f1 MeM6paHe np11 OTCYTCTBHII Clll'IH!JlblIOI'O IICHTH,L(a. |
|||||||||||
TaK, T-aHTHreH OB40 B oc1-10BHOM TpancnoprnpyeTCH B HJipo, |
|||||||||||
HO He6o.!Jbllia51 norryJrnrurn ero MO.ilCKYJJ JIOKaJIII3YCTC5I B IJ.1133- |
|||||||||||
M3TJFJCCKOiI MeM6pm1e. B coerane 3THX MOJJeKyJ1 oTcyTcTByer |
|||||||||||
CHfH3.'IbH3H HOCJIC)l.OBaTCJibHOCTb, OHH He 110,LJ,BepraioTCH rJlll |
|||||||||||
K03HJII1j)OBaHHIO, XOT51 CO,Ll,Cp}KaT yciacTKI! rJIHK03HJIHpOB3Hll51. |
|||||||||||
B TO )KC npel\rn XHMepHa51 MOJJCKYJia T-aHTmcna, co;iep1Ka |
|||||||||||
ma51 N-Kom eny10 cnrHaJihHyro IIOCJICJ\OBaTCJJbHOCTb reMarrmo |
|||||||||||
THmrna BHpyca rpmma, CB513LllHICTCJI |
C |
3H,L(OIIJJa3ManJlICCKHM |
|||||||||
penIKy.'IyMOM H IIO)l.BCpracTCH |
I'JIIII<03HJ1IpoBaHIIIO. DCJJOK CHH |
||||||||||
|
|
|
|
|
|
|
1 |
|
|
|
|
Te3npyeTC51 He Ha CB513fHHlLIX C MCM6paHaMH, a Ha cno60AHbIX |
|||||||||||
110Jrnpn6oc0Max, |
H cro r.1rnK03IJ.1rnpoBaH11c n |
TpaucnopT 1m |
|||||||||
KJICTOllHYIO TIOBCpXHOCTb HC ITOJI3BJUJCT M01IeHCHH - lIHI'H6H |
|||||||||||
TOp cpyHKJVIH anrrapaTa roJJbJl)!Uf. TaKHM o6pa30M, TpaHCITOpT |
|||||||||||
T-aHTHfCHa K HJJa3MaTHlJCCKOH MeM6pane npoHCXO,L(HT He no |
|||||||||||
MeXa!IIT3MY |
TpaHcrropTa 1rneT0 lllblX ceKpeTOj)HblX |
6eJIKOB, |
a C |
||||||||
|
|
|
|
|
1 |
|
|
|
|
|
|
BOBJJCl!CHIICM JlJ)ynrx, IIOKa HCH3BCCTl!b!X MCX3l-lH3MOB. |
|
||||||||||
IlpaBITJJb!Tb!I'i TpaHcnopT 6CJIKOB 51BJHICTC}I HC06XO/llfMh!M co- |
|||||||||||
6b[THCM ,L(JI51 ycTaHOBJJemrn IIJ)OJIYKTHBHOiI IIIHpCKI HH, B |
TO |
||||||||||
BpeM51 KaK napylliCHHe Tp3HCIIOj)T8 0/lHOro H3 6eJIKOB MO)KCT |
|||||||||||
llOCJJY}KHTb npH4HHOi.i npcpBaIIHOH, a6opnrnHOH, HHqJCKUHH. |
|||||||||||
HarrpHMep, |
npH |
3apa)1{cum1 |
KJICTOK |
acu11rnoii |
K3J)UHIJOMbl |
||||||
3p.1JJXa BIIJJYCOM rplllllla <jioprvmpycTCH a6opnmmrn HHqJCK |
|||||||||||
UlIH, |
conpoBO)KJiaJOUlaHCH flJ)OJJ,YKUIICH |
IICll lHj)eKIU!OHHbIX |
He |
||||||||
CT36H.JlbHb!X B11pyconoc11.06HblX |
IaCTIIJJ,, |
He CO,Il.CJ))KaIIUIX |
mm |
||||||||
|
|
|
|
|
1 |
|
|
|
|
|
|
COACJ)}KcllllHX pe3KO YMCHblliCHHOe KOJI!ILJCCTBO M-6CJ!Ka. 061rn |
|||||||||||
py}KeHHHil |
B COCTaBe |
1 |
M-6eJIOK JJMeeT )l.CqJCKT |
||||||||
T3KHX |
JaCTll!I. |
||||||||||
Hyro crpyKTYPY (CM. many 7). M-13eJ10K, KaK JJ3BCCTHO, HBJJH |
|||||||||||
eTcH MC,Ll,lic!TOpOM c6opKH m1pyrn1,1x qacrnu. Cpa3y )KC JJOCJJe |
|||||||||||
CllHTe3a OH Tpa1rcnopnrpyeTCH B IIJia3M3T!TlJCCKYIO MCM6paHy, |
|||||||||||
BCJie)l. 3a LJeM HCMe,IJ.JICHHO H3CTYII3IOT IlOLJKOBa1-rne H Bb!XO/I. |
|||||||||||
BHpyca ll3 KJJeTOK. Ilpn a6opTJIBHOH HHqJCK!J.IIII KJJCTOK acuHT |
|||||||||||
HOH KapuHHOMbl 3pm1xa Bbl5IBJ151CTCH HC06bFIH351 ,Il.JJH M-6CJ]K3 |
|||||||||||
BHYTPHKJieT0 IlI3H |
JJOK3Jll13aJI.IIH - Ol<OJJ051,L1,epna51 o6JJaCTb |
UH |
|||||||||
|
|
1 |
|
|
|
|
|
|
|
|
|
TOll.!Ja3Mbl, |
r,IJ.e, llO-BHAHMOMY, |
ocy111.ecTBJrneTC51 |
ero |
Clll1Te3, a |
|||||||
JI.CqJeJ<T B |
CTPYKType |
rromrnenn1,LJ,a rrp1rnoAHT |
K |
Hapyweunro |
|||||||
cro |
TpaHcrropTa |
K |
nJia3MaTJI4CCKoit |
MCM6pane |
[Bukrin |
||||||
skaya A. G. et al., 1981]. |
|
|
|
|
|
|
|
||||
TpaHCilOJ)T 6eJIKOB II3 H,II.pa B UHT0!1JJa3MY 11 H3 UHTOJJJJa3- |
|||||||||||
Mbl B HAPO CBH3b!B3CT reHeTH'ICCKYIO 1rnqiopMaIJ,HIO C 6IIOCHH |
|||||||||||
TC30M 6eJIKa II HBJJHCTCH KJllOlJCBL!M MOMCHTOM B |
)KH3IIC)J,eH |
||||||||||
Tem,HOCTIT KJJeTim H penJIHKaTHBHOM JTKJIC BHpyca. 5I;LepHaH |
|||||||||||
MCM6paHa IIMCCT CBOHCTBa MOJJeKyJrnprmro CHTa |
c |
pa.n.nycoM |
|||||||||
nop OKOJIO 500 HM, H Memrnc, II CJ)C)l.HIIC MO.!ICKYs1bl MoryT npo |
|||||||||||
HHKHYTb B |
51)].pO |
nocpeACTBOM flJ)OCTOH )l.Hq>cpy31u1. 0AH3KO |
93

pa3MCp MH0rHX cpyHKUJIOH3JlhHO B32KHhIX M0JJeKyJJ npeBhll.liaeT /lH3MC::'TJ) 51/lCPIIL>IX II0p, TaKHe M0JJeKyJJbl ,l!_0JJ)KHLI IIMCTb cne Ull3Jlblll..>IH ClffH3JJ, 06cc11canrnaJOW.HH HX rrp0HHKHOBem1e 13 511l po. IT pnpo,LI.a :noro cnrna;1a 6L1JJa Bbl5IBJJeHa cpasmITeJJLHO IICJJ.DIHI0 C Il0M0liJ,610 M0JJeKyJrnpno-reHeTil'ICCKHX MeT0JJ.OB. flepu1.,1c JJ,clHHLIC 6hIJJH Il0JiytJeHbl Ha BHpyCHOl1 M0JJ.em, - T-aH THreHe OB40 [,Kalclcron D. ct al., 1984]. DLIJJo rroKa3aHo, LJT0 TpattcrropT 13 H,ll.po 9Toro 6enKa o6ycnosnctt cncun¢H4CCKHMH IT0CJJC,LI.0l3aTeJJbH0CT5IMII aMIIH0KHCJJ0THb!X 0CTaTK0l3 B CaM0M IT0JJHITCIITH,ll.C. .3TOT KaJrnOtpHJJhHLIH CHrHaJJ npcJJ,cTal3JICII rrpe HMYW.CCTBCHH0 0CII0BHLIMH 0CTaTKaMH aMHH0KHCJJ0T B TI0JIO )KCHHH 126-132: PKKKRKV. HaJTHLJHe 3THX ocTaTK0B B co erase T-aHTHreHa OB40 OKa3aJJ0CL neo6xo,1u1MLIM 11 JJ.0CTaT0<I HbIM ,ll.J151 TpattCJI0Kauirn 6eJll<a B HJJ.p0. TaKHC IIOCJJCJJ.0B3TC.'lb H0CTH, COCT0HW.HC 113 6-8 3MHH0KHCJJ0THbIX 0CTaTK0I3, 6bIJIII 0611apy)KeHbl B COCT3BC p51)l8 KJJCT0LIIILIX H BIIPYCHLIX 6eJIKOB c HJJ.ep1rn11 JJ0KaJ1113auv.eA: 1<a11cnJlHLrx 6eJJKOI3 OB40 VP! 11 VP2, 60JJLU1oro T-aHTHreua 13Hpyca noJJH0MhI, NSl 6eJJKa BH pyca rp,mrra, DBP aJJ.eH0B11pyca, 6cJlKOB gag Bl14-l, B114-2, BI1O 11 ,np. [Cleghon V. ct al., 1989; Dechambrc M. ct al., 1989; Rihs H.-P., Peters H0aI1989]. B cocTaBe HCKOTopL1x rro mmc1rTHJL0B BL!5IBJICH0 ITO ,LI.Ba HYKJJCOlpHJlbHLIX CIIrH3Jia.
O,iJ,HHM H3 TI0,ll.X0/l0B K pcillemrro npo6JJCMbl 51BJI5ICTC51 K0H CTPYKUHH XHMCj)HbIX rcrron, C0JJ.Cp)K3llJ.HX o6JJ3CTH, K0T0pb!C K0JJ.Hpy10T HJJ.epH1.,1c H urnorrJia3MaTw1ccKHC 6cJrnH. Tar<HM ny TCM, HanpHMep, 6bIJI IIOJJYLJCJI rn6pHJJJII,I(I 6enoK, COJJ.Cp)l{allJ.HI1 M0J1CKyJ1y 1111pyoaTKIIHa3Ll 1<yp H 06J1aCTb 113 0CTaTI<0B 0CH0B HblX aMHHOKHCJI0T T-a1n11rena OB40 ITpo - Jfo3 - Jlrrn - J1113 -- Apr -- J1113 - Ba.TI. 3TOT 6cJlOK 9qicpcKTHBno rpattcrrop THpooaJ1c51 13 KJICTO'IlIOe 51,ll,pO. bOJJCC Toro, 6CJJKII, accou1rnpo Bclllllble C CHHTCTlll!CCKIIM neIITIIJl0M, IIMCIOIUHM TY )KC ll0CJie )L0BaTCJILHOCTb 113 7 3MHH0KHCJJ0T, T3I()KC H3KallJ[[Jil3JJHCb B KJieT0amoM 51;1,pe [Lanford 0aHct al., 1986].
TioK[J3aHo, tJTO 51JJ.epHLIH Tpa11cnopT 6cJIKOl3 ny1rneoKancIIJJ,a VP2 H VP3 33IlHCHT OT llpHCYTCTBHH B VP2 3MllllOKHCJlOTHblX 0CT3TK0B B ITOJI02KCH!IH 317-323. IloJ1ya1Clll,I rn6pH,ll,Hhl[1 6e
.'I0K, COJJ.Cp)l{3IUHII BMecTo VPl OB40 6cJ10K c T11nwrno11 UHTO 11Jl33M3TH'ICCKOH JIOKaJJH33UHei-i - K31ICHJJJ!blll 6C.'IOK BHpyca 110JJHOMHCJIHTa VP1, H o6a K3IICHJJ.HLIX 6eJIKa OB40 c HJJ.epttofi JJ01<an113auHe11 - VP2 H VP3. l'n6pHJLHLIH 6eJJ01<, co.nep)Ka mn11 HYKJJeocpHJJbHYJO 110CJie,nosaTeJJhHOCTb, 6LICTpO MHrpHpo ll3JI 13 KJJeT0a1HblC 5IJJ.pa, 13 TO IlpCMH KaK npH JJ,CJieuim 9T0ro y•iacTKa 6enoK ocTasancH B UHTorrna3Me [Wychowski C. et al., 1987]. TpattcrropT B HJJ.po rpex KancHJJ.HbIX 6eJJKOB OB40 o6yc JIOBJICH VP1 6JJaroJJ.apH HaJJH0m10 ttyKJJeoqrnJJLHoro cHrHaJJa
0aH

H3 8 aMIIH0KlICJI0THblX 0CTaTK0n Ha N-KOHU:e M0JieKyJibl. Ta )Ke nocJie.uonaTeJibHOCTb B 6eJIKe VP2, no-BH.UHM0MY, HrpaeT ncnoMoraTeJibHYIO p0Jib, B3aHMo.uciicrnyH c VP1, CB5!3b!Ba5! 6eJIKH C pCI].CIITOpaMH H.Uep11oii MCM6pam,I JII6o rrpH.uanaH 6eJIKaM COOTBeTCTnYIDIIlYIO KOHCJiopMailIIIO .UJI5! K0ppeKTHOro n3aHMO}leiicrn,rn c VPl. HecMoTpH na TO qrn HYKJieocjmJII,HbIH
CHrHaJI B C0CTaBe |
T-aHTIIrCHa OB40 rrpCJI,CTaBJICH 0CTaTKaMJI |
|
B IIOJIO)KCI!HII 126-132, |
06.JiaCTb, cjJJiaJIKHpyromaH .9T0T y,ra-• |
|
CTOK, 6blJia Ba)KHa |
)],JI5! |
TpaHCJIOK8JlJIII II 3cjicjleKTHBHO BJIHHJia |
Ha KHIICTHKY ncpeHoca, 06cc1TC1 IIIB85! BhlCOKyro CK0p0CTb H ce JieKTHBHOCTb. B 3TOM Y'I ICTKC M0.lICI<YJII,J H8XO}l5!TC5! caii:Tbl cjloccjlop1urnpona11nH. ¢occ]lop1umponarmc T-8IITIIreHa rrpoHc X0.UHT 2 p,na - BII8lJaJJC B lUITOTIJiaJMC, :l8TCM B HJlPC, H CCTb ocHonaHHH rrpc.urroJI:1ran, yHrncT11e cjloccjiopnm1ponamrn n TpaHcrropTe T-a11rnreHa B H.UflO [Rihs H.-P., Peter ITHa1989]. 5foepH1,1ii T-aHTHreH cjloccjiop1rn11ponaH n 3HaH111TeJibH0 60Jib meii CTCIICHH, qeM U:11T0IIJJa3MaTWICCKHH. OTCYTCTDHe 35 aMH H0KHCJIOTIILIX ocTaTKOB c C-Konua KancII.uHoro 6eJIKa OB40 VP3 TaIOKe rrpHBO.UHT K Hapymemno TpaHcrropTa n11pycHLIXTHI 6eJIKOB n KJieT0'IHbie H}l,pa [Gharakhanian E. et al., 1987
cha IIOCJICJJ,OBaTeJibH0CTh o6yCJIOBJ1IIBaCT 5!ll,CpHyIO JIOJ(3,JIJI3a UHIO VP3 II n oTcyTcrnnc npyrux mIpycm,Ix 6cJJKon.
Pammii 6c.TToK <1}lc11on11pyca ElA, pcrym1py1omIIii Tpanc Kp11mo110 CO0CTBCIIIIbIX H KJICT0T1111,IX rcII0B, JIOKaJIH3YCTC5! B KcTTeT0HIH0M 51}].pe. Ero IIYKJlCO(JlII.JILHI,Ifr CIIrH8Jl HaX0ll,IITC5! CPC.UII 5 aMJIH0KIIC.JI0TIILIX OCTaTK0D Ha C-KOHIJ,e M0JJeKyJJh[ [Lyons THIet al., 1987]. 3TOT )J(C 6CJIOK C0JlCp)KIIT ll,0II0JlHII TeJJLHbIH HYKcTTeocjJIIJibHbllI CIIrllilJ] MC2KJI,Y 140-M H 185-M aMll H0KHCJl0THblMII OCTaTI'3MII. 3TOT CllrHaJJ cjiyHKl].IIOHHpycT B oou11Tax, no nc n coM8TIIHIcrn11x IrneTK8X [Sbvicek THIct al., 1989]. Ta,rnM o6pa30M, nyKJJcocJrn.JibHI,Ic cnnram,I MoryT .ueii cTnonaTb CCJJCKTHBIIO n 38BIICHMOCTII OT nma 1-rneTKII-035!IIllaX. ,Upyro11 pammn 6cJJoK a}lenonnpyca DBP (6cJJOI\, CBH3L1Ba10m11ii:c51 c ,UHK) HBJ1HeTcH rrpoll,yKTOM pam,ero reHa E2A. 3rnT 6eJJoK o6napy)KJ-IBacTc5! rrpeHMymecrneHH0 n H.Upc. OH co.uep MIIT .una HYKJICOcjJIIJibHI,IX cnnia.TTa. 0mm 113 HHX, C 43-ro ITO 85-ii aMHHOKHCJIOTHblH OCTaTOK, co.uep)!<::JIT DbIC0K0K0lICCpna THBHYIO II0CJle,U0BaTeJII,HOCTb flII3 -- IHaeT .TTH3 - flII3 - flII3 - Apr - TTpo, KOTopaH cxonHa c HYKJ1cocJrnm,m,1M cIirHa· JIOM n coCTanc T-aHTHreHa OB40 ITpo - .TTII3 - JI,n - fl113 - Apr - fl113 - BaJJ. BTopaH HYKJ1eocjrnm,11a5! 06.JiacTh MCHce KOHCepnaTHnHa, 0,UHaKo BCCLMa cxo;uia C COOTBCTCTBYIDilleH IIOCJie.uonaTCJibHOCTb!O B T-aHTIIrCHC OB40 II COll,ep2KIIT II0CJJC ll,0BaTeJJLH0CTI, ITpo --- .TTn::i -- Jhn - - Apr -- .TTeii --- Apr - Apr -- Apr rc1cr hon V. cl al., 198D]. )LBci ll)'l<JlCOC]m11l,Ilb[X
THI

CHrnaJia 061rnpy1Kel-lbl T3l0Ke y 60JihU10fO T-aHTHrena BHpyca
rroJIHOMbI n 11ecTpyKTypHoro 6eJ1Ka TIoHoBHpyca rpmma
cHMHBKnbKB1CCpC6XCBlTOCIoaayTC
TaKIJM o6pa30M, HYKJICO(jHWbHble CHrHa.nhl, COCTOHlIJ,He H3 KOpOT!(OH !IOCJIC)WBaTeJibHOCTH OCTaTICOB OCIIOBllh[X aMill!OKHC JIOT o6Hapy)KCHbI B cocTaBe noJrnncnTHJJ.OB Mirnrnx PHK- H JJ,HK-co,nep:iKamnx BHpycoB H o6yc.noB.llHBaIOT H/l.CPHYIO JIOKa JIH3a1uuo 3THX 6c.TJKOB.
rnaea 7
C60PKA 06ono'fE'fHblX BMPYCOB M POnb
MATPMKCHOfO 6EnKA
061.l.l,He npHHI Hilhl coopKH. MoprjmreHC3 o60JI0OIC'IHhlX Bll• pycoB, 33BepmaionJ,IIHC5! c6opKofi BHpycHblX 'l3CTHIJ, II BbJXO ,ll,OM HX 113 KJieTOK, HBJISieTCH CJIO)KH!,IM npo11,eccoM, OC!-10B3H HhlM Ha 06rru1x np1rnnmrnx (jlopMH[JOB31IIl5! trnpycHhlX CTpyK TYp, H B TO )KC BpeMH CTporo CIICUiltj)Il'JHb!M )VJH KcllK/WI'i rpynrrbI BHpycoB. eC3HCMOprj>0rerre3 npOCTblX rrnpycoB OCHOB3H Ha MOJieI<YJUiprroi\ caMoc6opKC, TO JVropc\JOrCIIC3 C,10)KHblX o6o Jl011C1!HhlX rrnpycoB COCTOIIT H3 HCCKQ.JlhKIIX IIOCJlC,AOBaTCJihHbIX 3Tanon, BKJlJOlia!OlIUlX n ce6H CHHTC3 II c6opKy cy6mrpycm,Ix CTPYKTYP H Tp,rncnopT Bcex BHpyccne1uHpw1ec1rnx KOMnoHeH TOB K MCCT8M qmpMllpOBaIIIIH !3ll()llOHOB. OIi Tpe6yeT yqacnrn pH,Qa Moptj1ono3TH1ICCKIIX (!HlKTOj)OB KJICT0O1HOJ'O H BHpyc11oro npm1cxo;,i<;1,emrn:. C6op1rn 060JIOlfCl1HbIX BHpycon o6wrno npo11cxo;1,JIT Ila ITJia3M3TllOICCI<Oil MCM6pa11e, KYlta He3aBIICHMO Jtpyr OT Apyra TpaHcuopTIIpymTCH see s11pyccne11,mjm11ecKHe KO\HJOHCIITL>I n cy6B11pycm,1e crpyKTYPL>L aope,uBapHTeJir,1-10 Ha IIOBepxIIOCTh ll.133M8TH'ICCKOi'.i MCM6paHbl TpaHcnopn1py!0TCH 11apy1Kl-lb!C BHpycm,re 6CJ1Kll - rJIHKOI!j)OTC!IJJJ,I. 3TH 6eJIKH CHHTC3Hpy10TCH Ha pH60COMax, CBH3aIIHblX C MCM6pa1rnMll, H no MCM6pa1-1aM3H,.TI.OII.J[33M3Tll'ICCKOH CCTH qepe3 arrnapaT fOJlb}J)Kll, r,u.e npoHCX0,11,llT ynopHJWHll!BcllllIC yrneBO,AHblX 11,e lIO'!CK, BbfHOCHTCH na KJICTO'!HYIO llOBCJ)XIIOCTb Ii BCTJ)3HB3JOTCH B onpe,11,CJ!Cl!Hb!C yirncTKll n.na3M3TWICCKOH MCM6paH1,I. BHYT· peHHl!C KOMIIOHCHTbl BHpyca HYK.TI('OK3llCH}lbl II cepnu.eBH Hhl -«y.3H3IDT» M0,11,HqlHI(HpOBaH!lhlC yrracTKH MeM6paHbl H IIO,rl.XOJUIT K HIIM co CTOpOHbl II)ITOJIJJa3MhL Y3H3Bamie ocyme CTBJI5!CTCH C HOMOl.l.l,b!O JLBYX Mexa11113MOB: 1) ny1rneoKaIICH)( B3al!MO/J.eikrnyeT C yqacTKOM fJJHKCmpoTCHJJ.3, rrpOHH3LIB310- l! IJM KJJeTOlIHYIO MeMfipany H Bb!XO,'l,5!U(IJM na ee IU!TOIIJI33- i,;;,'J'lp)e('K)iKl l!OBCpXHOCTb; pOoB c6opKy BOB.1eKaeTCH euie O)J,HIJ
HOp

A |
6 |
B |
3
Puc. 5. YcJ10lil.11cm1c cTpyKrypnoii opra11113a1 mt oOoJIO•w•rnb1x nttpycon.
A- ToraaHPyc; B - BHpyc rpHnna; B - BHll; 1 - nnnonponm)I.Haa 06ono•.1Ka; 2 - MaTJ)HKCHblll 6M0K; 3 - 0CH0BH0ll 6eJJOK cep)I.U:CBHHbl.
liefJlHK03IIJIHpoBaHHblll BHpycHI,Iii. 6eJJOK, HBJIHIOIUHHCH Me;urn TOpOM c6op1rn,- M-6eJJOK. BcJien 3a o6pa3oBaHHeM CBH3H MC)KJW fJIHKOrrpoTeH)laMH H HYKJieOK3HCH)lOM H3'1!1HaeTCH Bbl HfP!HB3HHe KOi-IrJJOMepaTa qepe3 IIJ133M3TH'IecKy10 MeM6paHy' 33K3H1Il:!BaIOUJ,eecH orrop)KCHIIeM BHpycHoii lJ3CT!fU,bl OT HO BepXHOCTH K.lleTKH. 310T rrpor ecc ITOJJYcJIIJl H33B3HHe «IIOllKO Baime». B IlOJJHpH30B3Hllb!X 3IIHTC.llH3JlbllblX KJieTK3X IIO'lKO Bamie HpOHCXO;lHT JHI6o C anHK3JJbHOH, mr6o C 6a3o-mnepaJJb HOH rroBepxHocTei\ EJ1ern11 [Stephens E., Cornpans R., 1988].
f1o cJimK11ocT11 CTPYKTypBoii opramna1um 11 MopcporeHe3a 060Jroqe1m1,1e n11pycb1 MoryT 6bITb pa3)leJJCHbI Ha 3 rpyrr111,1. B 1-ro rpyrrny BXO TJ:HT OTHOCHTeJibllO npocTo opraHH30B3HHble (t.-ToraBHPYCbl, fJJlIKOnpoTeH)lbl KOTO!)blX, npoHH3hIBaH JlHIIH)l HbIH 6IIc.rJOH, B33HMo,rxeiicTBYIOT c HYKJJeo1<ancH,iIOM. 3To B3aH MOJLeHCTBHe HBJIHCTCH MopqJOTI03TIIl!eCK11M rpawropoM, onpene ,/1H!Oll(liM CHMMCT[)JllllIOe pacnpe)].e.11em1e rJlllKOIIpoTeH)lOB lI 6eJ1KoB Hy1<.r1e0Kancn,II,a [Fuller C., I 987]. IioJJee CJJOlliHo op raHH3oBaHHbre opTOMHKCOBHpycH, napaMHKCOlrnpyCbl l1 pa6- )lOBHpycbI llMCIOT llOIIOJJHHTCJlhHbli'r MaTpHKCHblH 6eJ10K, JJOK3JJl130BaHHblH Ha BHyTpeHHeH ITOBCJ)XHOCTH JllU!OIIpOTell)lllOli 060"101m11. OH B3aIIMoneHcrnyeT KaK c rn!lKOrrpoTeII)laMH 060JIO'IKH, TaK H 6eJIK3MI-! HYKJICOK3lICH)l3. PeTpOBt1pyc1,1 o6JJa)la IOT erue 60J1ee CJIO>KHoi'1 cTpyKTypHofl opraHH3au.IIeH. TioMnMo airn.nora M3TpHKCHOro 6eJJKa na BHyTpenHCH noaepXHOCTH JIH nonpoTeII)lHOi'r 060.no1rKH, OHH conep}K3T ,iIOHOJIHHTCJlbHhI i 6e.10K cepmi,eBIIHLI, ¢opMIIpy10IIu1H KancyJJy cep/IJJ,CB!lHbI. OH pacno.1omeu Me.tKJW MaTpHKCHhIM 6eJIKOM II HYKJieonpo TNIJlOM (pnc. 5).
f1p11 sapa}Kel-llrn K.'JCTOK KYJlb TYpbl TK3Hll a-Tora1111pycoM BHpycoM J]CCOB CeM,IJl!KII. - Ha IIJ133Man111ecKoii MeM6paHe 1<.:reToK IIOHBJIHIOTCH 3 rJIHKOil[WTCIIJla EI, E2 II E3 B 3KBil-
97

M0JJ5lpHblX K0JII1'1CCTB3X. E2 np0HH3b!B3eT llJia3M3TH4eCKYIO MeM6paHy, ero C-KOHU,CBOH y4aCT0K pa3MepoM 0K0."10 30 aMH H0KHCJIOTHLIX 0CTaTK0B HaX0,l]JITC51 C IJ.HTOI1.,'Ja3M3THlfeCKOt1 CT0pOHbl MeM6paH H y3H3eTC51 orrpe,u_eJJe!-IHb!MH Il0CJIC,ll_0BaTeJJb l-l0CT51MH HYKJJeoKarrcn,n_a [Simon K, Garoff H., 1980]. O6pa3osaHHe CB513H MC}K.ll.Y JI.BYM51 6e"1KaM1-1 CJTYlKHT cHrHaJJoM JI.JJf! Il0"IK0Bamrn; B TI0'IK0BaHHH T3K2Ke MoryT nrpaTb p0JJb JJ3TC paJibHblC B3aHMO)l.CHCTBHSl MC)K)].y rJIHKOrrpoTCH.ll.3Ml1. HyKJJCO
KaTTCH.ll.HhlH 6e.rioK cBH3bIBaeTCSJ c C-KOIII(OM MOJJeKyJibI E2 nyTCM 3JICKTpOCTaTil IeCK0ro B3aHMO,l.lCliCTBIUl. B o6JJaCTb CB513h1B3HH5I .ll.1HpcpyH,ll,HpyeT Bee UOJlblllCC K0JrnqecTB0 BHpycm,IX rJll1K0ilpoTeH;'l,0B, K0T0pb!e MMM06HJIH3yIOTCSl nyTCM CBH3b!B3- I-IHJI C ll0)J.Jle)KafI.J}IMH1-1yx.ryeoxanCH,l.l_aMH. ho npHBO)l.HT K Bhl JlSltJHB3HH!0 TI0tJKH Ha,l.l KJie"rO'-IHO[\Jl0BCpXH0CTLIO. Tec1-1oepac IT0JJO)l{CJIHC rJrnKonpOTCHJl0B Ha Hapy.11<HOH CT0poHe JIHITHtl,HOro 6HCJJOJI H HYKJIC0K3ITCHJl.0B Ha nHyTpeHHeA CT0pOHe npH yse JIHtICHIHI TeKyIJCCTH Jll!TIHtl,0B npHB0tl,HT K y,naJieHJII0 KJlCTOl.JHblX 6eJIKOB H3 o6JiaCTCii c6opKH. Hco6xo)]JfMhlM yCJ10UHeM }].Jl51 IIOlJK0BaHIISl, I303M01KH0, 51BJIHe"l'CH npoTe0JJHTHlJCCKoe Hape3a HHC pE2 c o6pa3onaHHCM E2 y s11pyca C1m,n_6Hc H p62 Ha E2 H E3 y supyca JJCC0B CeMJIHKH. rio-BHtl,HMOMY, npH 3TOM npo HCX0,ll_HT KO!HpopMaILHOHHble H3MeHeHHH lllJIITa rmmonpoTeH,u_a, 06;1erqaJ0u r1e 06pa3ouar-me JJaTepa.nbHhIX K0HTaKT0B MelK,.'1,Y HapylKl·lblMH6e.r1KaMH.
Y 060J1oqeqHbIX BHpycoB, co,u_ep2Kam11x M-6eJJOK, Kpwrw1e CKHM M0MCHT0M, onpe,ll._eJISl!Oll.lHM B03M0lKH0CTb Il(JlJK0B3HHSl, SJBJJHeTcsi BKJJI011emre M-6cJJKa B n.ria3MaTI-I'lecKyJO111e1116paHy KJ!eTJm co CTOp0HbI lVIT0llJJa3Mbl. K: T3KHM BHpycaM 0TH0CSITC5I BHpycbJ rpmrna, napaMHKC0MHpycbI, pa6J(OBHpycbl.
.5e3060J10lJC'IHhlC BHpyChJ C03peBalOT BHYTPH KJJeTKH 11 oc BO0O>l{)].3IOTC.H BO BHCWHIOJ0 cpeny o6bJlIHO II0CJIC ,u_ec-rpyKUHH KJJCTKH (BhlXO,LI_ HX KJJCTKH IT'YTCM «B3pbIB3»). O,u_HaKO CCTb HCKJII0tJCIIHH H3 3T0ro npaBHJla. Co3peBaHne pOTaBttpycoB BKmOtJaeT He06LJllHLIH Tllfl llQl[K0BaHJrn, npH K0T0pOM cep,nue BHHbl, cpopr.rnpy1om11ec51 B UHT0I1/l33MC, !10'-!KYJOTCSl B nporneT 3H,ll.OrJJJa3M3THlfCCKOro pCTllKYJiyMa, npuo6pe-raSJ BpCMCHHYI0 O0OJJOtJKy. 3ro ca1110 ITO ce6e IIC0Uhl'IHOe 51BJ1CHl·Ie cpe,ll,H BHpy C0B. B npouecce !10"1K0B3HHSI 060JIO"!]<a yTpaq1rnaeTC51, npOHC X0.ll.HT c6opKa HapyMHoro KancH.n.a, 3asepmamm.asi npo1iecc MOpcporeHC3a, fl BHpycHI,TC 1JaCTHUbl H3K3ilJIHB3l0TCR B Be3IIKY JJax 3I·!tl,0IIJJa3M3TH1leCKoro peTHKYJIYMa [CM. 13yKpHHCK35! A. r. H ,!1.p., 1989]. ,UpyrnM llCOULlllJ-lb!M cpeHOMCIIOM s c6opKe po-ra• BHpycoB 51BJJSieTC5! y•iacrne HeCTPYKTypHoro BHpyCHoro 6eJJKU NS28, K0Top1.,111 JIOKa.rm3yeTCH 13 Me1116pa11ax 3HJI..0IIJJa3Mi:JTHqe, CK0ro pcnIKy.rryMa 11 Bhl[10JIH5IeT cj)yHKUHIO peuenTopa tl,Jl5I BH•
98

j)yCflhIX cep!(uemrn [,Chan W. ct al., 1988; Au K. ct al., 1989; Meyer J. et al., 1989]. 5eJIOK co,11.ep.tKHT 175 aMHH0KIICJIOTHhlX 0CTaTK0B, H3 K0T0pb!X 131 0CT3T0K 3KC!10H1Ip0B3H Ha UHT0· IIJ133MaTH<ICCKOii CT0p0He BC31lKYJI, II!)CJ(CT3BJ!5UI co6oil C-KO· HCU MOJICKYJ!bl. 3TOT YtJ3CT0K c.11y>1rnT ,Ll,Jl5I Cl35I3b!B3HJUI miraH J(a BHpycHbIX cep,LJ,UCBIIH. Ha N -Ko11ue, HaXO!\RmeMcR B rrpo cBeTe BC3HKYJJ, pacn0JJ02KCH HCOTpC33HHblli Ciff!IaJlbHblH 11Cl1Tlf,Ll,.
EeJIOK coJI,ep:ann |
2 |
cafiTa |
rJrnKosIumponamrn y N-KoHua |
H 3 rIIJJ,poctJ06HI,IC |
06Jracn1, |
113 l\OT0pblX T0J[bK0 OJJ,Ha (2-R) |
|
saRKopimaeT 6eJ10K B |
KJICTO'lrrnii MCM6p:rnc [ Bergmann C. |
et al., 1989].
13eJIOK NS28 RB,,'!HC'TC51. 11p111v1cpoM Toii He60.r11,mo1i rpyrrn1,I MCM6paHill,!X UCJIKOB TIIIJa I1,HT0XpoMa, K0T0pb!C H3XOJ],5ITC5l B MeM6pmiax B 11eo6bllJHOii OpHCI-ITaumf,KOI'J],3 N-KOHeu pacno
.TIO}KCH Ha BHyTpe1rnefi 110Bepxuocn1 MCM6pa1Ibl, a C-KOHCU - Ha Impy2KHOTI ee II0BepXH0CTH (B CX0,11,HOH opHCHT3UHH naxo J(HTC5I TaK)Ke 6eJioK EI Kopo11aB11pycoB, J10Kam1s0Bam1ufi B MeM6pa1rnx arrnap,na foJJbJl,)KH) [Machamer C., Ross J., I 987]. IIo CBoeii qiy1mum1 CJlY}KHTb MeM6paHHbIM peuenrn poM J],Jl}I BHYTPeHHero KOMllOHeina BHpyc1wr1 lJ3CTHUbl rrpn
c6opKe |
NS28 6JJH30K K MaTpHKCH0My 6eJJKY o6oJJol!etJHhIX |
BHpycon. |
O,I]JlaKo cymeCTneH!IblM pa3JIIJlJHCM 5IBJrn:eTC5I BK.'llO· |
lJCHHe M-6CJ!Ka B C0CTaB rmpmrna, J'/],C 011 !IpOJL0JL>K.aeT Bbl· II0JlHRTb CBOIO ueMel!'rnpyIomy10 c!JYIIKI(HIO. IIo-BHJJ,HMOMY, 1-1aH- 60J1bwee CX0!\CTBO 6eJrna NS28 6yJJ,eT 06napy}Ke110 c MeM·
6paHHblM M2 |
6eJJKOM B11pyca |
rpnnna, KOTOpblfI ![3XOJ],HTC5I |
n rrJJasMaTw1ecKofi MeM6paHe |
sapa:iKeHHoi-i K.'ICTKH [Zebe |
|
dee S., Lamb |
R., l 989], xorn qiyHKUIIH ero noKa HeH:-iBecTHbL |
OB40 TaK}Ke BhIX0.!lHT H3 1rneT0K ,LJ,o HX JI,ecTpyKUHH. Bn pycm,Ie qacTHll,bl HaKaIIJ1HBaIOTC5I Ha anHK3JlhHOH rronepXH0CTif
KJleTKH II BO BIIYTPHKJJCT0'IHhJX BaKyoJJRX [Clayson E. et al., 1989].
<l>yHKU,HM MaTpHKCHOfO 6eJJKa o60JIOlJelJHblX BHpycoB. MaT pHKCHhlfl 6CJ10K 113XOJI_HTC5I B m1p11011e 110;1, mmorrpOTCHJJ,llOH o6oJJ04KOH, 0Kpy2Ka5I HyK.rieoKancHJ],. 9TO C3Mb!H Hll3K0M0JJe KYl15IpllblTl CTPYKTyp1-1b1i"16e.rIOK (3a HCKmotJeIrneM peTpon11py coB). MoJieKyJrnpHaR Macca M1-6eJma BHpycoB rpmma co cTanm:ieT 25 000, napaMIIKCOBHpycon - OT 30 000 JI_O 40 000. B TO :lKe npeM5I Ha JJ,0JlIO 3T0ro 6eJJKa rrpHX0J],HTCH OT 30 J],0 50% 6eJIKOBOH Macchi. 4HC.'IO MOJICKYJJ yKasa1111oro 6e.rIKa na BHJ)H0II BHpyca rpHrrrra C0CTaB.irneT 1500-2500, Ha BHpH0H BHpyca BC3IIKYJJ5IpHoro CT0MaTJJTa - 0K0JJ0 1600. ::horo K0.J:lH l!CCTBa !(0CT3T01IIIO J],J151. o6pasoBaHH5I CIIJJ0lllHOro 6eJIK0130fO
MOHOCJJOR rroJJ, mrnonpoTeHJJ,1-1011 060JJO'IKOf1 Bnpyca. TaKaR cn.rwurnaR HJIH JI,a:iKe npepbrnaIOmaRCR 6eJJKOBa5l MeM6paHa
99

CT36Il-..'IH311])YeT m1pyCHy10 qacnm,y, H yMeHbUlCHHe OTHOCH TCJlbHOH KOHUeirrpau.HH M-6em<a B COCTaBe BHpycoB rp1mna H napaMHKCOBHpycoB conpoBO)K,ll.3eTCH TIOHBJICIU!eM xpynKHX BH])YCI!h[X '13CTRU (1rnnpttMep, y XOJJO,ll.OBhfX MYT3HTOB s11pyca rp,mna, y BHpycHoro ITOTOMCTBa, rrpo.uyuupyeMOro KJleTK3MH acunrnoi1 KapuuHoMhI 8pm1xa) [cM. I3yKp11HcKaH A. L, 1985], y BHpyc11L1x 1 iacn1u KOpH B 1rneTKax M03ra npH no,n,
ocTpOM |
CKJJepo311py10u eM naH3HI e<lrn-1111Te [Hall W., Chop |
pin P .. |
1979]. |
y peTpOBHpycoB no,u JIH110TIJ)CHeHJ{HOH o60JJOLJKO{J, Heno cpe.ncrneHHO npttMhma,1 K cepAuesrrne, 11axo)l.HTCH 11erJ1HK0311- JrnpoBaH1-1b1e6eJIKH, 3KBIIB3JJeHTHble M-6eJIK3M BHpycoB rp1m na, napaMHKCOBHpycoB H pa6,nOBHpycoB. 9TO 6eJ10K pl5 y Mbl lliHHblX peTpOBHpycos, p 19 y TITH%HX peTpOBupycos, p 17-p 18 y Bl1l.J. BepOHTHO, oHH BblllOJJHn!OT Te )Ke Q)YHKUHH B c6opKe BHpycon, •no 11 M-6eJIOK.
Ha Mo,u.e.1111 Bl1LJ noKJ3aHo, 1no p 17 c!rnKcttpyeT rJJHKorrpo TeHlJ.bI, ttaxo,zurn HCC5J Ha JJOBepxHOCTH BHpycHOll 43CTHIJ.bl, Ii
06pa3yeT |
MOCT |
Me}K/J.Y |
l'J1HK011])0TCH)].3MH |
Ii HYKJTeOK30CH |
|||
,D,OM, cnoco6cTBYH H30MeTpHqecKOfi |
YKJJ3)].Ke |
HYKJJeOKaITCHAa |
|||||
[Marx |
P., |
Munn |
R., I 988; Ozel |
ct |
al., I 988]. C 3TOH TOliKII |
||
3pemrn |
p 17 npe,ncraBJJHeT |
co6oii |
MarpnKc, npo1H13bIBaiomati |
e1-1pyCHy10 tJacrnuy. EenoK p24, (fJOpMHpy1omnf:'1 KancyJJy cepk I eBHHbl ll rrpn,na1orn.Hii er1 KOHJ:ll!eCKYlO q>OpMy, B33HMO)le(rcT ByeT JJHUJh HeMHOrHMH MOJleKy.J1aMH C OKPY)K3!01.UJIM ero M3T pHKCHl.>IM 6eJJKOM. TTO-Bll,D,HMOMY, p24 HC BKJIIOlfeH B MaTpHKC H OTHOCHTeJJbHO CB060AHO pacnoJJaraeTCH 13 ueHTpe BttpycHoA 1 13CTHIJ.bl, TIOCKOJibKY pa3pyweHHe JIHTIOIJJ)OTCHJtHOH o6oJl04KH BHpyca HeHOHHblMH ,11,erepreHT3MII (NP40, OKTllJlr.!llOK03llll,) H
ll,C30KCHX0.'13TOM H3TpHH conp0130)KJ.I.HCTCH 131.,IXOJlOM p24 B |
|||
0Kpy,Ka1on1,y10 cpe;w (pHc. 6). [GyKp1111cKa51 |
A. |
r., |
LUapo |
sa I-t K., 1989; Bukrinskaya A. G., Scharova |
N. |
K., |
1990]. |
I-IaJl!f4HC CB060)],HOfO 6eJ1Ka p24 13 CblBOpOTl{C KpOBH 3apa :l'KCHHblX Bl14 H 60JJbHhlX MO!l<eT 6b1Tb o6ycJJOBJICHO uapywe1-rn eM ueJJOCTHOCTH mrrrorrpOTeH,11,HO}i o60JJO'lKH BHpyca H BblXO,!J.OM He CBH33HHblX C MaTpHKCOM MOJleKyJJ 6eJJKa B OKpy>KaKrnWJO cpe,ny. ::>Ta CHTyaUHH Heo6wrna ,!J.Jl5J JJ.pyrnx 060.,,oqetJHblX BH
pycoB, |
pa3pywe1me |
KOTO])b!X HCHOHHblMH 11 llOHHl.>IMH ,!J.erepreH |
||
THMH |
Il])HBO,UHT K |
paC'rnopeHIIIO |
J'JlHKOllpOTCH,!J.OB [Oxford J. |
|
et al., |
1981; Thompson S. et al., |
1988], a B npHCy'TCTBHII BhlCO |
||
KOll Ko1-1ueHTpau.rrtt COJlH n 6ycpepe (0,5-1,0 MOJlb |
NaCl) K |
|||
pacrnope1rn10 H MaTpHKCHOfO 6eJ1Ka [oyKpHHCK35! A |
r., I 985J. |
B BHpHonax BHpycon rpHmrn H napaMllKCOBHpycoB M-6e JIOK Tecno B3aHMO)leHCTByeT C ny,rneoKanCH}],OM H mmonpo TeIIJlHMI o6oJJOLJKOH. EcJJH c HYKJJeoKarrcHJJ.OM M-6eJ10K cBH3aH
100