
Медицинская вирусология / Букринская А.Г., Жданов В.М. Молекулярные основы патогенности вирусов
.pdf
rnaea 8 nPOTEOnHTM"IECl<ASI Al<TMBAijMSI BMPYCOB
)lmr 0K0H'!aTeJibH0ro qJOf)MHJ)OBaIIIrn m1pycHblX 6eJIKOB, a TaK>Ke npHo6peTCIIIIH BHpycoM mHf)Cl<I IIOHHOH aKTHBH0CTH II BIIJlYJieHTH0CTI! HC06XO)]J!Mbl ll0CTTpa11cJllUUI0HHhle M0AH(pIIKa UHH 6eJ1K0B, cpem1 K0T0J)blX cylll,CCTBCHHblM II 065I33TCJibHbIM npaKTiflieCKH J Jl5I ncex BHpycoB Ilj)0LJ,CCC0M 5IBJI5IeTC5I rrpoTe0- \.JIHTWICCK0e Hape3aHHC BHpycm,1x OCJIKO!l. PoJib 3T0ro rrpo qecca AMI HIHpCKIIHOll llOro IIIIKJia lrnpyca II0AqepKHBaCTC5I 1'epMHH0M «IIJ)0TC'0JIIITll'leCKaH aKTJmanm1». II0;1. TIJ)0TC0JIIITH qecKOH 3KTHBaJ]J'Ieii II0H!IMalOT 1rnpesam1e nepBHlJHbIX TIJ)0AYK T0B TpaHCJl51UHII - 6eJIKOB·npe;uneCTBeHHIIKOB C o6pa30B3HHeM HHJU-IBH/1,YaJJl,HblX c!JYHKLJ,IIOHa,/lbHblX BHpycHblX 6eJIK0B, B pe3YJibTaTe K0Toporo BHpyc np1106peTaeT IIHcpeKUIIOHHYIO aK TIIBII0CTb. TaKHM o6pa30M, npoTe0JIIITIIqecKa5I aKT11Bau1rn: HBJ1HCTCH 33KJII0llllTCJ]bHhIM aKT0M MOpcporeHesa.
IIpoTCOJIIITJ1lICCKOC Hape3aHHe M0)KeT Jie)KaTh B 0CH0Be qiopMI1J)OBaHH5I Bcex BHpycHblX 6eJ1KOB, o6pa3yl0lll,IIXCH II3 Ok Horo 11.nII 2-3 fl0JIH!Ip0TCHH0B - npCALUCCTBCHHl1K0B, KaK 3TO lIMCCT MCCT0 y 1111KOj)H31lHpycoB, ToraBnpycoB, peTp0BHpycoB, JlH60 )KC njl0TC0Jl llT[l'ICCK,HI aKTHBaIIII5I MO)KCT 3aK.'JlOlJaTbCHB Hapesamm o;u1oro IIJIH ABYX lJ,llLJ,C Bcero Il0BepXH0CTHblX 6eJI K0B. 3THM 6CJ1KOM y 060JroqeqHhlX n11pyc0B o6blI IH0 HBJIHeTCH l'JllIK0IIp0TCHA - 6eJIOK CJIH5IIIHH. Hape3aHHC np0HCX0AIIT rry TCM T0lJClJH0ro (orpanuqeHHoro) npoTC0JlH3a C o6pa30B3HIIeM 2 cpparMeHTOB, BII0Bb o6pa30B3HHblH K0HCLJ, 0AHOJ'0 113 HIIX (N-KoHeu) C0JJ.Cp2KHT fl0CJJeA0BaTeJlbH0CTh H3 0CT3TK0B rHA pocpo6HblX 3MIIH0I(lICJIOT, K0Topan B33IIMO/\CHCTByeT C JllII1Hk Hb!M 6IIC.!JOCM KJICT01IHOH MCM6paHbl.
)lJI5I KoppeKTH0ro Hape3amrn BHpycHhIX 6eJIKOB Tpe6y10TCH np0Tea31,1 onpeJ1,eJiettttoi1 cne11.HcpHamocT11. Bee npnpoAHhie npo-
BKeHoT0oTiepeKpecrm,ie peamurn MC}K.D.Y M-6eJllrnMH napaMHKco1rnpycos, BbJJIBJICIHlble MC'l'Ot{OM HMMYH06,10TTl!Hra C HMMYHHb!MH CblllOpOTKaMH KJlOBH 1rnJJoseKa H )KHBOTHbtX [UJapoaa H. K. n .n.p., 1989].
1 |
) |
H.iITXMMnrnb. uo |
|
|
msseJCOsCYMJ!irn J |
|
T[bCM]M |
C8IJIMJI |
|
acJJ9 |
|
t{l acJel |
|||||||||
c |
6enKaMn |
|
napaMHKCOBHpycoa; |
I, |
2 - BHpyc |
naparp1mna ITTHI.\ |
|
4; |
|
3, 4 - |
|||||||||||
DHPYC |
Ce.H,n.afi; |
6 - |
MCHI1KBJahp3nA- |
|
b3lIHIp6oyppJI- |
|
MO{)CKoH |
|
|
|
npOTHB |
||||||||||
m-1pyca naparp1rnna |
'leJIOBCKa |
TJHia J |
C O('l!KUMII |
napaMl!KCOHKJJYCOB: |
I -· Bl!pyc |
||||||||||||||||
Cett/\ail; 2 - Bnrt:I-1; 3- 1mpyc |
rraparpttnrra ITTHI.\ THna 4; 4 - BIIpyc napo• |
||||||||||||||||||||
lpH(HO- |
5 - BCTflJ-3 |
D |
c |
3c4cnKTYcIqp5Ia- |
KnaynIi)qrrrrpaal- |
I |
BTyOJX1- |
XnIeaJ- |
)l,!rr- |
||||||||||||
THB |
aupyca |
Bnrq.2 |
|
tieJIKaMH |
rrapaMnKco,mpycon: |
Bnrq"2; |
2 |
|
nnpyc |
||||||||||||
napoTHTa; |
3 |
anpyc |
|
naparpnrrna |
nTHU. THrra |
4; 4 |
sHpyc |
Ce11.n.ni'1; |
r |
|
B3aHMO |
||||||||||
,ReilcTBHe CblllOpOTKH .n.eTel!, rrepe!loJICBllllfX naparpnnHOM Tl!Jla 3, c OCJJKaMll |
|||||||||||||||||||||
uapaMllKCOBllpycos: |
I BHP.-l-3; |
2 |
1mpyc |
uapornrn; 3 - BIIpyc |
uaparpnnna |
||||||||||||||||
IITllIJ.; |
4 |
Bllpyc |
CCIJ)\afr. |
|
|
|
|
|
|
|
|
|
|
|
|
H0HT

"" |
|
|
|
|
|
|
|
|
Tao JI Hu, a 17. BHpyrnble |
npoTea3bl |
|
|
|
|
|
|
|
|||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Ce H fICTBO |
I |
|
|
I |
|
T;m nporea3h! |
I |
OCTaT0h'. aKTHB- |
I |
|
|
Y4aCT0K Hape3aHHH |
I |
<1>.rnHKHpy10ume |
|||||||
|
|
|
|
|
|
|||||||||||||||||
|
|
|
|
Horo catna. ero |
|
|
|
|
|
aMHHOKHCJIOTHbre |
||||||||||||
|
|
|
|
no HHrn61nopy |
|
|
-1 |
I |
+1 |
|||||||||||||
|
|
|
|
|
TI030tKeHHe |
|
|
|
OCTaTKH |
|
|
|||||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|||||||||||
|
|
|
|
|
|
|
|
|
|
|
||||||||||||
PerpoBnpy- |
|
p15 BHpyca cap- |
|
|
AcrrapTHHO· |
|
Acrr-30 |
|
- |
|
|
- |
|
Tipo, |
A,ia, |
Ba.11, |
||||||
Cb! |
|
K0Mhl Payca |
|
|
Ball - |
|
Acrr-95 |
|
|
|
|
|
|
|
Jleii, |
l1,1efr |
|
|
||||
|
|
|
|
pl3 Bnpyca ,1eii- |
|
|
|
|
|
Acrr-30 |
|
THp |
|
|
Ilpo |
|
Cep, |
Tprr, |
AJJa, |
|||
|
|
|
|
KeMIIH Mbllllelf |
|
|
|
|
|
Acrr-95 |
|
Jleil |
|
|
AJia |
|
,'lea |
|
|
|
||
|
|
|
|
|
|
|
|
|
|
|
|
|
Cep |
|
|
- |
|
|
|
|
|
|
|
|
|
|
pl I Bl1q |
|
|
|
|
|
|
Acrr-25 |
|
|
|
Tpu |
|
Tpn, |
Cep |
fmi |
|||
|
|
|
|
|
|
|
|
|
|
|
[JIH |
|
|
|
|
|||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
Acn |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Tnp |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Tipo |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
l1.11e |
|
|
|
|
|
|
|
|
|
ToraB11pycbl |
|
P2 |
|
|
|
CepHnoBaH |
|
- |
|
Ba.11 |
|
|
AJJa |
|
A,1a, |
Ba.'l, |
|
|
||||
|
|
|
|
|
|
A,1a |
|
|
|
|
|
|||||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|||||||||||
|
|
|
|
|
|
|
|
|
|
|
|
|
L1n |
|
|
f.1u |
|
J,faeii |
|
|
||
|
|
|
|
Karrcn.n |
|
|
|
|
|
|
fHC-}40 |
|
UHc |
|
|
Tup |
|
fJ!H |
|
|
|
|
|
|
|
|
|
|
|
? |
|
|
Tpn |
|
|
Cep |
|
fJJy |
|
|
|
||||
|
|
|
|
|
|
|
|
|
|
|
Acrr-145 |
|
|
|
|
|
|
|
|
Lrn |
|
|
<i>.1!3BHBHpy- |
|
? |
|
|
|
CepnnoBall |
|
Cep-215 |
|
|
|
|
|
|
|
|
|
|
|
|||
|
|
|
|
|
|
|
Apr |
|
|
[JIH |
|
.llns, |
Apr |
|
|
|||||||
Chi |
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Cep |
|
|
|
|
|
||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
Tpll |
|
|
|
|
|
|
|
|
|
2A |
|
|
|
|
|
|
|
|
|
|
|
|
A.1a |
|
|
|
|
|
TinKopna- |
|
|
|
|
? |
|
Acrr-40 |
|
Tnp |
|
|
[JIH |
|
J1eil, |
I1.1e11 |
|
|
|||||
BHPYCb! |
|
|
|
|
|
|
|
|
UHc-110 |
|
Tpu |
|
|
|
|
Mer |
|
|
|
|||
|
|
|
|
3C |
|
|
|
|
|
|
fHc-118 |
|
A,ia |
|
|
L1H |
|
A.11a, |
Ba.11 |
|
|
|
|
|
|
|
|
|
|
UncTeHno- |
|
Acn-80 |
|
[JIH |
|
|
|
|
|
||||||
|
|
|
|
|
|
|
|
B3ll |
|
Uuc-150 |
|
r.w |
|
|
Cep |
|
Jlefl, |
l1,1el1 |
|
|
||
BnpycbI |
|
YqacToK |
re11rnr- |
|
|
Cepl!HOBllll |
|
[Hc-165 |
|
|
|
|
|
|
|
<l>eH |
[JJH, |
Acn, |
||||
|
|
|
|
Cep-71 |
|
|
|
|
|
|
|
<!>en, |
||||||||||
rpHrrna C |
|
rJIIOTllHHHa |
|
|
(3cTepa3a) |
|
|
|
|
|
|
|
|
|
Cep |
|
|
|
||||
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|
|

TCa3bl B 3aBHCHMOCTH OT qJYHKUHOIIaJ!btlbIX rpynn B aKTHBH0M ILCHTpe !IX MOJICKYJI 0TH0CHTCH K 4 KJiaccaM: CCj)HII0Bb!C, BKJTI0- 'Ia!0l.l.]JIC 110,ll.KJiaCCbl TpHIICllHOBhlX H cy6THJIH3HHOl3hIX nporea3, ILHCTCHH013b!C, acnapnrnOBblC H ll,llHKC0,D,ep>KamHe. ITpoTea3bl, 1iape3aiomne BHpycHhIC 6eJIKl1, MoryT 6hITb KaK KJieroqHoro npoHCXO.iK,ll.CHIHI, HlK H BHpyccnelJ,mj.JH'leCKHMll, 3aK0,ll,HponaH HhlMH B m1pycHOM reHOMC. HanpHMep, npoTe0JillTll'lCCKa5I aKnmalJ,rrn BHpycon rpmrna H napaMHKC0BHpycon ocymeCT n.irnerc5I KJICT0'lHbIMII Tj)IHICHHOIIOJI.06HUMH npoTeasaMH, TIH K0pHaa11pycoB m1pyccnelJ,m.j)H'lCCKIIMIt lJ,!ICTCllHOBh!Mll npo TCa3aMH, roranHpycon H qrnan01rnpycoa - n11pyccne11,mpH1Ie CKHMH cepHH0Bh!MH Ilj)OTCa3aMII, ))CTp0Bll!JYC0B - acnapnrno
BLIMH rrporeasaMH (Ta6J1. 17) [Wcllink 3eHhvon Kammcn A., 0ICCnhBazan J., Fletterick Ipah0ICCyhWeber rohet al., 0ICIhaHe
O6Hapy2KeHa BHpyccnelJ,m.j)HlJCCKa5I 3CTepasa B C0CTaBe BHpy ca rpmma THna C.. AKTHBHhIH cailr ee npe).l.CTaBJieH K0Hcepaa THBK't'li1 ,n:,:rn cepm10BbIX rHApoJias TpHaAoii Cep Acn fuc, 0,IJ,IWKO CTCI1CHh r0M0J!0rHll C APYfHMH cepHHOBblMH rHAPOJia33MH HCBhlCOKa5I, H <pepMCHT npe,D,CTaB.115ICT co6oii: H0BOC ce MCHCTBO cep!Ul0BblX npoTea3 [Vlasak Haet al., 0ICIlTh3cTe pasa <pyHKILHOHHpyeT Ilj)Il npomIKHOBemrn Bttpyca B KJieTKy.
CxoJJ,HaH 3CTepa3Ha51 aKT!!BH0CTh o6HapymeHa y KOpoHaBI-1py C0B [Vlasak paTet al., 0ICClKhChnpoTeoJ111THHrecKOH aKTHBaIJ,HH
peTpom1pycon Y'IaCTDYIOT KilK KJICT0'IHUC cepHH0Bbie, TaK H B11pyccneILlHjrn11ecKHe acnapTllBOBb!C 11porea3bl. IIpoTCOJIHTHqe. cKoe Hape3am1e ocymecrnJrneTcH B annapare foJib,IJ,:iKH JIH6o npH H0CJICJI.YIOI.l.LeM TpaHCIIOpTe 6eJIKOB, JIH6o JJ,a.iKe noc.r1e OT· 110lJK0BaHH5I BHpycHOH qacTHil,bl OT KJICTK!L
IIpoTCOJIHTl:l'lCCKa5I 3KT!IB3lJ,ll5I BIIpycon saBHCHT OT p5IJI.a ycJIOBHH: HaJIH'lH5I cne1LH<pw1ecKoil nporea31,1 B sapa2KeHHOH 1rneTKe IIJIH OMhmammei\ KJieTKll )K!l,ll,K0CTH, nepeHqHoH crpyK Typbl C0C,IJ,HHHTeJihH0ro nenTH,ll,a B yqacrKe napesaHHH, JJ,0CTYil H0CTll yqacTKa Hape3aHH5I ,IJ,JI5I nporea3bl. CymecTBCHHOe 3Ha tteHHC HMeIOT KOHqJOj)Ma1vrn MOJlCKYJlbl 6eJIKa, HaJIIPIHC 0OKO Bh!X 11,eno11eK 0Jllff0CaxapHA0B, K0T0pb!C MOryT MaCKHp0BaTb y11acT0K Hape38Hll5I.
MHorne rJIHKOI1pOTCH,ll,bl o60JIO'lel!Hb!X BHpycoB IlO,IJ,Bepra IOTCH nocTTpaHCJIHUH0HHOMy npoTe0JIHTH'ICCKOMY HapesaHHIO no ocTaTKY aprnHHHa. ,UJIH rmrnonpoTeHA0B, co,1J,ep2Kam11x ap rIIHHHOBhie HJIH .'1H311110Bble 0CTaTKll B yqacrKax Hape3amrn, nporea3h! HMCIOTC5I BO BCCX KJICTKax; ,IJ,JIHHa C0C)J.HHHTe.nbHOro nenTH,ll,a IT0JIHIIp0TCHJJ,a navbHpyer OT 2 OCT3TK0B 0CH0BHliX aMHHOKHCJIOT (n cnyqae, Hanp11Mep, BHpyca J1acca) JI,O 6 ap-
rHHHH0BbIX H .7Hl3HH0BhIX OCTaTK0B B reMarrJIIOTHHHHe B!Ipyca rpHnna [Klenk bBhMcahRott peHT0ICCrOh
8 3aKa3 N• 1293 |
1233 |

MyTair.tm B yqacTKe nape3amrn Be)l.yT K 6JIOKHpOB3HHIO rrpOTeOJlll33. TiocJICJUICC o6bilJlIO He npernITCTByeT BHYTPHKJie TOlJHOMY TpaHcrropTy Bnpycuux 6enKoe H c6opKe BHpycHoA ll3CTliULI, OJ1.H3KO rrpt!BO)]JIT K rrpo,11_yKUIHI HeirnqJeKUHOHHOrO TIOTOMCTB8.
TaKHM o6pa30M, npoTCOJIHT!-flICCK35I 3KTHB3UH5I BHpycoB HMCCT ymrnepcaJII,HbIA xapaKTep, HrpaeT cyw.ecTBeHHYIO pOJ1b B rrpHo6peTel!HH 1mcjJeKUHOlIHOH aKTHBHOCTH H HBJIHCTCH KpH Tlll!CCKHM co6bITHeM, o6ycJJOB.'1!1B3lOW.HM naTOfCHHOCTb BHpyca. B H3CT05!.lllee BpCM51 H3YlJCHHe npoTeOJIHTlf!IeCKOA 3KTHB3II.HH BupycoB np1106peTaeT oco6oe 3H3lJCHHe B CB5!3H C TeM, 'ITO 3TOT npouecc npeJ1.cTaBJ1HeT co6oi\ yHsBnMy10 MHIIJeHh J1..irn 1111ru61nop0B npOTCOJ!H33, H B03M02KHOCTb HC110Jlb30B3HH51 no CJIC,lUUlX B Ka'leCTBe aHTHBHpycHblX npenapaTOB JJeLKHT B OCHO
Be IIOBOro HanpaBJICIIl:Ul XHMlIOTepanHH BHPYCHb!X HH(peKJ.I,HH. Hau60.r1ee xopowo nsy11eHa npoTeOJIHTII"ICCK3H 3KTHB3llH51 y Bnpycos rp1mrra, napaMl!KCOBupycoo, peTpom1pycoB, nHKOpHa supycoB, Torasnpycoo. TirrnBJUllOTCH ,ll,3HHble O CYW.HOCTII 3TO· ro rrpouecca H y ,npyrnx BHpycoe.
Bupycbl rpmma. BHpyCbl rpHnna l!C,10BeKa H JKHBOT!lbIX rnna A HMeIOT 13 rro.nnmon reMarrnIOTHHHHa (HA). Cpem1 BH pycoo rpmma ITTHU eCTb BbICOKOITaToreHHh!C sapHaHTbl, Bhl3hl "B3!011lHe J1eTaJ11,ny10 HHtj)cxumo; OHH coAep)KaT HA noJ1.THHOB H5 H H7, XOT51 HMCIOTC5I II HenaTOrCHHblC BapaaHTbl, BK,1IOl!a IOW.HC TC }KC rIOJJ,THITblYcraHOB,1CHO,. 'ITO pema10w.ee3Ha<JCIIHC ,Ll_,lj}I naToreHI!OCTH HMeeT 3cp<peKTIIBHOCTb 1rnpe3aII!I51 npe.n rneCTBeHHl!Ka HA (MoJ1exym1pHaH Macca oTKaaaOKua 60JiblliYIO n MaJ1y10 cy61,c,1urnnuu, HAI H HA2 c MOJ1exyJ1HpHoii Macco0 TaKaaaKH 25 aaaeKcoeJJ,1rneHHb1e 11.HCY/Ihcjrn,n,nofi CBH3b!O. Hco6XOJ1.HMOCTb ttapesamrn CBH3aHa C TeM, '!TO C ITOMOlll.blO BHOBb o6pa3osairnoro N-KOHIJ,a HA2 ocy11LeCTBJISieTCH CJIHHHHe BH· pyCHOH o60JIOlJKH C KJICTO'lHOH MCM6pauoi-i H TCM C3MhlM o6ec nelJHBaeTC51 npoIIHKHOBCHHC B UHT030Jlb BHYTPCHHero KOMDO ueHTa BHpyca. B cocTase Y'IaCTKa, rrpttJieraion{ero K N-KOHUY HA2, Bbl51BJleHa BbICOKOKOHCepBaTHBHaH o6JJaCTb 113 IO aMH· HOirnCJIOTHb!X OCTaTKOB, fOMOJ!OfH'lH351 N-KOHIIY F-6CJIKa BHpy ca CeH,llai-i. 9TOT 6eJJOK T3K}KC ocyw.eCTBJI51CT CJIHHI!He BHpyc HOH H KJlCTO•rnofr MeM6paH npH npOllllKHOBCHHll BHpyca 13 KJICTKy.
TaKHM o6pa30M, 1rncpeKilHOHIIa51 aKTHBHOCTb BHpycoB rpHn rra II rrapaMHKCOBHpycoB B 3H3'IHTC,1bHOH CTeneHH saBHCHT OT CJHlB3IOIUdi cpyHKI]JIH BHpycHbIX 6eJIKOB CJIHHII!f51. HOBHpycoB
rpmma 3TOT rrenTH)], CKpbIT B TpexMepHOI1 CTPYKType M0,1C· KyJrbl HA, O,ll,HaKo rrpH IIH3KHX aHatJeHHHx pH HTHaOpKtteo6xo
.UHMbIX )]JH! CJIIUTIIH51, □pOHCXO)l.HT KOH(pOpM3UHOHHOe H3MCHC· HOeH

1-rne M0,1CKYJibl, H rrerrTH,ll. 3KCilOHHpyeTCH Ha mrnepxHOCTH MO JICKYJlbl. I!ape3aHne3aBHCHT He TOJlbKO OT 11aJll-14IH1 K.TICTO'lHb!X cueu,mpttYeCJ{HX TpHTICHHOllO)l.061-Ib!Xq1epMeHTOI3, HO 11 nepmll[ HMI CTpyr<Typbl B y•iaCTKe Hape38HI-IH,B ITCITTH}le, CBH3hll3810ll.J,eM HAI u HA2. 8h1coK0B11pyJrcHTIH,rf1 BHpyc KJiaccw1ecKoi-i 11yMhl nTI-JIJ, (BK1-II1), K0Top1,1il Bbl8bIBaeT J1cTaJ11,Hy10 IHH JeK u:1110 y UbmJI»T, MC)KJW C-KOHllOM HAI H N-KOHUOM HA2 HMe cT nocJ1eJJ,0BaTCJ1bHOCTb JlH3 •···- Jlm -Apr - I'JIJ1-- Jhn - Apr, 13 TO Bpc;,,rn KaK BHpyCbl rpmnrn 1ICJIOBCKa, 8BHPYJICHTHblC JVHI llblIIJrnT, H HCIT3TOfCIJill,JC IITll'll,11 B11pyc1.,1 rp1rn11a Toro .>KC rro,1rr1ma, 'ITO H BKYI1 (H7), coAC)l)KaT B 3Totl 06J1acT11 TOJ!b KO OJl,HH ocTaTOK apr11111111a [Webster I( d al., 1982]. CpaBHH TeJ11,Hb1ii 8HclJIU3 IIeJ)BU'JIIOii CT[)YKTypu IIUTOI'CHll1,IX II IICTT3TO reHHblX lllT3MMOB IITilllbHX BHpycoB rpmrna, co.uep)Kall.lJ1X H7, IIOK338JI, 4TO HA Hena-roreHHblX BllpycoB HMCeT MCHb lile aMn HOKHCJIOTHblX OCT8Tl<OB B CBH3b1B8IOJ..UeM rrenTH,1.(e, 'ICM l18TO rettHble BapHaHTbl.
)lJJSJ 1-1ape3a1rn» o65I3aTeJICH ,1.(HIICITTH}T, J1113 - Apr B COCTa Be C .ll.HHIITe.!fbJ-IOrn nenTH)l,3. I1pn OTCYTCTB!Il-1 ;umerrTH,1.(8 8 npe.uirrecTBCHHHKe HA n11pyca rp1mna 11apc3an11e HA nponcxo
AHT rom,KO B orpaHHIJCHHOM lJHCJle KJ1CTOK-X035!CD, l1 DHOBJ:, o6pa3ol3aHHI,lli mrpyc 00.llcl]ltlCT HII3!{0[f lllltilCKUHOHHOH 3KTHB
HOCTbJO. :XOT5l 3,lMCll/1 ilMll!IOIOIC,IJOT!IhlX OCTclTKOB B coeJU!HH T€JlbHOM IICIITll}l.C l!C lJaPYIJJ;]CT TIHllICflOj)T rJll!KOllpOTCH,1.(a rrpe}I.WeC'rneHHHKa K KJJCT011HOH IIOBCJ)XIIOCTII 1-1 ero 3KCITOH!1- p0B8HIIC Ha f!J183MaTHlJCCKOfl MeM6pa1-1e.
51pKHM npHMepOM pOJIII npOTeOJ!H3a rrpe.umecTBeH!mKa I-IA Bttpyca rpunna CJ1Y}KHT 3IlH,UCMHH rpnrrna cpeAH ,ll,OM3UIHHX rrTHU B llJTaTe TTeHCHJILnamrn (CLUA) n 1983 r. B anpeJJe BH pyc rpurrna THna A (H5N2) o6JJa}l,aJJ HIBKOH n11pyJ1eHTHOCTblO, He Bbl3h!B3JI JICTclJlbHOH I!l-lCpeKI.1,JIH y JWM3111HHX llTI!U. O.unaKo ym:e B OHTH6pe 1mpyc nprro6peJJ BbICOKYIO BHpyJJeHTHOCTb 11 Bbl3B8JI T5I}l{CJIYIO 311H)].CMmO, B pe3y,/1bT8TC KOTOpofi norn6.110 60J1ee 17 MJIII ,UOM3ll.lHHX ITTIIU [Webster R. et al., 1986]. BH
PYJJCHTHbIH HIT3MM OT.Tlf-1'18J!CflOT aBnpyJJCHTHOfO MHOfHMH CI30HCTB8MH. On Bbl3bJBaJJ o6pa30BclHHC 6JISIHICK IIO)]. arapOBblM 110Kp1>n1-1eM 6e3 ;\06a13.11emrn: Tp1-mc1ma, D ero coCTanc He 061rn PY:>KHBaJ1rn npe,n,wecTBCHHHK HA, a JlHllJb npOAYKTbl ero rrpo Teo.;11n114ec1<oro tiape3aHHH HA I H HA2. Y an11pyJ1eHTHoro llIT8MM8 rrpe)I.WCCTBCHIHIK 6b!JI Bb!HBJ!CH B 3H8'IHTCJ!bHOM KOMl 'ICCTBe. 3n1 ,U3HHble CBH)].CTC.1bCTBYIOT 06 l!HTCHCHBHOM 1-1ape- 33Hllll HA nupyJreHTHoro UJT3MMa.
ITp11 cpanncmrn rrepm1111rnii crpynypbr reHon anupyJ1cHT Horo H BHpy.rieHTHOfO 1UT8MMOB 6bi.'lll 131,151BJICHhl pa3JIIJLIJISI B
06,nacn1 COC/1,llHIJTeJ/J,Hl,I.X |
IlCf!T!l)lOB B y11acnrnx npoTe0.1U'l'll- |
8* |
115 |

Ta 6 JI !In a 18. CpaBHeHHe coe/1,HH!ITeJJbH0f0 nenrH11.a H cp.UHK11py10ru.Hx aMIIH0KHC.rnTttblX IT0CJJe)l,0BareJJbH0CTeli HA BHpycoe rpHnna A/l.lbllIJ1eHoK/ne11cHJJbBaHH!1/83, BHpyJJeHTHblX H aeHpy.qeHTHblX wraMM0B
|
B11pyc |
I B11py ,1eur- \ |
|
|
|
|
|
HOCTb |
|
|
|
|
|
|
UbmJJeHoK/Illor/59(H5) |
+ |
TIpo |
||
Tern/SA/61 (H5) |
+ |
ITpo |
||
|
l-1H.ll!OK/OHr/66(H6) |
+ |
Tipo |
|
l.l.hlTIJieHoK/Tiett/83(H5) |
+ |
ITpo |
||
|
HH}l.lOK/!1pJI/83 (I-15) |
+ |
TIpo |
|
Y-rKa/I-1:p,1/83 (H5) |
+ |
TIpo |
||
FPV/34(H7) |
+ |
Tipo |
||
l.(wn,1eHOK/I1ett/83 (H5) |
- |
Tipo |
||
|
||||
lltt""IOK/GA/72 (HS) |
- |
flpo |
||
|
||||
;rn:a/Ml/80(H5) |
- |
Tipo |
||
|
||||
J'TKa/Tiett/84(H5) |
- |
flpo |
||
|
L1u
r.m |
Apr |
||
fJIH |
|
- |
|
|
|
|
|
rm, |
- |
|
|
|
|
|
|
r.111 |
- |
|
|
Lntt |
- |
|
|
|
|
|
|
L1n |
|
flpo |
|
r. H |
- |
|
|
f.lH |
- |
|
|
|
|
|
|
r.w |
- |
|
|
|
|
|
|
rJl!I |
- |
|
|
|
|
|
noc,1e.aoaaTC,lbHOCTb• ocTaTKOO
- |
- |
|
- |
|
|
,\pr |
- |
|||
|
|
|
|
|
|
|
|
J1H3 |
||
f.1m |
TpH |
|
Apr |
|
Apr |
f,nH |
||||
|
|
|
|
|
|
|
|
|
|
|
- |
- |
|
Apr |
|
..\pr |
|
- - |
|||
|
|
|
|
|
J1H3 |
|||||
- |
- |
|
- |
|
|
,1H3 |
.'1H3 |
|||
|
|
|
|
|
|
|
|
|||
- |
Apr |
|
J1H3 |
|
,\pr |
-- |
||||
|
|
|
||||||||
|
|
|
J1H3 |
|||||||
|
|
|
|
|
|
|
|
|
|
|
- |
Apr |
-- |
|
Apr |
J!H3 |
|||||
|
|
J1H3 |
|
|||||||
|
|
|
|
|
|
|
|
|
||
Cep |
|
Jl113 |
--J11n |
|
Apr |
|
r.m |
|||
- |
- |
|
- |
|
.11!3 |
.7H3 |
||||
|
|
|
|
|
|
|
||||
- |
- |
|
- |
|
|
Apr |
r..111 |
|||
|
|
|
|
|
|
|
|
|||
|
|
|
|
|
|
|
|
|
||
- |
- |
|
- |
|
Apr |
f;m |
||||
|
|
|
|
|
|
|
||||
|
|
|
|
|
|
|
|
L1H |
||
- |
- |
|
- |
|
|
Apr |
|
|||
|
|
|
|
|
|
|
|
Hl
JlH3
,JIH3--
.:1u3
.1,n
--J!H3
jltt3
JT!l3
J1H3
flpo
Tpa
Tptt
Ii2
. .\pr Lin
,\pr Lm Apr r.m
Apr r.111
Apr r.111
.-\pr fm1
Apr r. 11
Apr r.11-1
.-\pr r. !I
Apr fmt
Apr fJIH
TT pH Me tJ. a HI{ e. TIOJl.'l.epKHyn,1 ocraTKH 0CH0BHblX aMHH0KHCJ!0T.

1ICCK0ro 1-1apC33H!UJ: aB11pyJ1eHTHblH IlJTaMM HMeJI ,LI,Ba acrrapa rHH0BLIX 0CT3TK3, K0T0pb!C 0TCYTCTB0B3JIH y BHpyJieHTH0f0 mTaMMa (Ta6JJ. 18) [Kawaoka Y. et al., 1987]. IIo-BH,LI,HMO·
MY, B npouecce urrpKyJrnpoBamrn nrrpyca rrpmrnomJrn e,LI,HHH'l· Hble MyTaUHH, K0T0phle npHBeJIII K HHTCHCHBH0MY Hape3aHHEO HA c o6pa3oBaHHeM CJIH13a10niero MeM6pa11br N-KOHUeBoro
nennr,LI,a Ma.noi'.i cy61,e,LI,HHilUhI. |
choro OI<a3aJiocb ,n:ocTaT0'IH0, |
||||
1IT06LI BHpy.r1eI-1TII0CTb BHpyca |
pe3K0 B03p0CJJa. |
|
|||
|
flem11,11, B Y43CTKe ll3j1e381Ilrn M02KeT 6L!Tb 33M3CKHp0BaH |
||||
uerr0l!l{aMII |
OJIIIroeaXaj)IIJJ:0B, II IIX 0Tlll,CHJICHIIC o6ycJIOBJIHB3· |
||||
JIO |
pe3Koe |
B03pac1a1-111e 1rnrnn'IIH0CTII |
Bap11aHT0B [Rott |
R. |
|
et |
al., 1984]. O/1,!UlKO 11:lll('CTHLI CJ1Y'l31!, |
K0r,ll,a, HCCM0TpH |
Ha |
H3JIH4IIC OJIHrocaxap11)(0B, BllpycbI rpnnna TITHI.l, coxpaHHJIH naTorettH0CTb 6Jiaro,LI,apH O,ll,IIOBpeMeHHOMY yBe.TIH'lCHHIO '!HC.'13
OCTaTKOB 0CHOBHb!X aMHH0K!IC.10T B coe,ll,HHHTCJILH0M IICITTH,ll,e [phuchi M. et al., 1989].
Cylll,CCTBCHHOe 3Ha1ICHHe ,l],JIH npon'.OJIHTH4CCKOH aKTHBa UHH BHpyca rpmma HMeeT CTerreHb JJ:OCTYIIH0CTH y'!aCTKa Ha pe3aHHH ,ll,.151 npOTea3bL O pa3.lllfl!H0!{ CTCIJCHH CBH,ll,CTCJlbCTBY eT, ttarrpHMep, TOT cpaKT, 1rro Bee no,n:THIIbI HA aKTHBHPYIOTCfl Tp11rrc11110M 11 JIHllib HCK0Tophle IIJTa3MIUIOM [Tashiro M. et al., 1987]. OcTaTOK npomrna, 11axo,r1,Hlll,HikH B M0JieKy,ne HA B IIOJIO)Ke,ll!J[ 320, enoco6en yBe.llII11HTb Jl0CTYllHOCTL yqacTKa Hape3amrn JJ.Jrn npcneas1,1 fSchwartz T., 1986]. Ta1rnH )Ke poJib
ero 61,wa 06Hapy)1<eHa H ,ll,mr F-6CJ1Ka n11pyca Cen,ll,afl [Tashi ro M. et al., 1988].
11ocKOJibKY Ilp0TC0JIHTH9eCKaH aKTHBaUHH BHpyca rpnrrrra 06ycJJOBJ1e1w aape3aH11eM HA 11 onpeJJ.E'JIHeT TKaHeBbIH Tpo TIH3M H rraToreHH0CTb B3pH3HT0I3, CTaH0BHTCH HCCOMHCIIHOrl poJib HA B ITaToreHHOCTII nnpyca.
B 1iaTorcHe3e rpm111mHoil. 1111cj)eKnm1 na*IIYIQ po.m, Mory1 HrpaTb cep11H0BblC np0TC33hl, ceKpen1pyeMhlC HCKOTOpblt,Hl 6aKTCPilflMH, HarrpHMCJ) 30,710THCTbIM CTaqJIIJIOKOKKOM, KOTOj)bIH <IaCTO BLl,ll,CJJHIOT OT 60JibilblX C rpHIIII03HOII ITIICBMOHHeii. Ha JIH'IHC 6aKTepnaJJLHoro rt)epMe11Ta, ocylll,eCTBJJHIOlll.ero npoTeo JJHTHIJecKy10 aKTHB3U:IllO rrnpyca, cnoco6CTBYCT MIIO)KCCTBeH HblM UHKJiaM penJIIJK3UIIII BHpyca B opraHH3MC ;-1.a2Ke npn OTCYTCTBHH crreumjrn11ecKOii llj)()TCa3bl. 3apa,Ke1me MbllllClf 0,1], H0BpCMCIIHO C !3HpycoM rp1rnrra 30JIOTHCTblM CTacjrnJJOKOKKOM npHBO,QIIJ!O K j)C3KOMY YBCJllfllCllll!0 THTpa !3Hpyca B JierKIIX II rn6em1 MblillC!f OT rpHilll03HOH HHeBMOHHH. l1Hrn61nop1,1 npo
TC33, Tairne K3K c:-aMJIHOKanpo110BaH KHCJIOTa H anpOTllHHHbl [)K11pHoB 0. II. I! AP-, 1984], JwynenTIJH H ,n:p. [Tashiro M.
et aI., 1987], rrpHB0JUIJrn K 6.110Frnp0B,rnmo nape311H11H HA KJ1e T0'l!lhlM11 n GaKTcp11a.nhHbJM11 np<nea3aM!I LTashiro M. ct al.,
117

1987] H npe,Q0TBpamaJIH rn6eJib Mbllllei'i. OJI.HOBpeMeHIIOe 3a pa1KCHHC Mbm1e11 BupycoM rpHmrn I I 6aKTepuerr, npo,11.ynnpy10meii 3K30npoTea3y, Serratia rnarcescens, npIIB0JJ.H.r10 K pe3K0· MY ycHJICHJ:l!0 .neTa.!JbI!0r0 }l,eii:cTBIIH BHpyca, B 0CH0Be K0T0poro
.TielliaJia aKTHB311IIH nJia3MI!HoreHa Jl.0 IlJ133MHH3 6aKTepHaJlb HOH npoTea3ott; nJia3MHH Hape3aJl BHpycm,n'i HA H cnoco6CT· B0BaJI rcttepamrnaIJ,HH HHCpeK11,m1 [Akaikc T. et al., 1989]. B3aIIMOJl.CHCTBHe MC,!{Jl.Y BHpycoM rpmma H 6aKTCpHHMH II tta6JlIOJI.aCMb!H CHIIeprnqecKHH 3qHpCKT 061,HCHHIOT qaCT0C OC.f!Oli<HCHHe rprmrra 6aKTCpHaJibHbIMH rrnq>CKl(HHMH H YBCJ!H qem1e JICT3JlbH0CTH cpe,ru1 60.ribHbIX xpo!Ul'ICCKHMH pecmrpa T0pHblMH HIHj>eKLUI5!MH BO BpeMH rpHTITI03HblX 3TIH,1l,CMHH.
napaMHKCOBHpycbl. y napaMHKC0BHpycon npoTe0.rIHT!IlJC CKaH 8KTHB81[IHI CBH33Ha C Hape33HHCM npe,runeCTBCHHHKa 6e.riKa CJlHHHHH F0 Ha q.iynK11110HaJibHhIH 6eJioK F1 H 6eJIOK F2, KOTOpbll'i, 110-BH,l[I!M0MY, He HrpaeT p0JIJI B rrp0HHKH0Bemrn BH pyca B K.'lCTKy. Hape3aHHe ocyni:ecTB.r!HCTC5!, KaK H B c.riyqae m1pyca rp1mna, K.11eT0tIHblMH TpHITCHHOTJO,l[06HbIMH cpepMCHTa MH. 3TH cpepMCHTbI B }l,0cTaTO'IHOM KO.!l!-Jl!CCTBe06yc.rioB,TIIIBaIOT TPOilII3M BHpycoB K orrpe,11.elICHHblM TK3HHM If opraHaM, a T3K· )Ke CTCITeHh BHpy.rieHTl!OCTII, CB5133HHYIO C re11epaJIH3aUHeH BII pyca. HanpIIMep, B ocHoBc TpomI3Ma BHpyca Ce1u1af1 K J1e ro1rnott TKaHH MblllleH .r!C)KI!T BbIC0I<aH K0IIIl,CHTpal(llH B TKaHH TpHrrcHHOITOJL06HbIX qJepMeHTOB [Tashiro M., Homma M., 1983]. flpu 33MCHC 3Ml!II0KHCJ10THblX 0CT3TK0B 13 y1IaC'I'KC Hape3aHTIH F-6e.riKa B rrponecce Hacc11poBaHHH Bnpyca B npII· cyTCTBHH XllM0TpHITCIIHa 6hIJIII ITOJ1Y1ICHbl MyTaHThI, K0TOpble aKTJIBHpOBaJlHCb He TPIIl1Cl1H0M, a XHMl!OTpimCHHOM. Y T8KIIX MyTa!ITOB 061-iapy)JU!BaJIIICb l.J.BC 33MeHhl B C0C)LHHHTCJlbH0M IlCITTlIAC [Hsu M. et al., 1987]. MyTaHTbI 6bI.T!Il cnoco6Hbl T0.rlbKO K 0)].1-IOMY ll,IlKJiy penp0JJ.YKl( liH B .!lCrKIIX MblIIIeH, r,r e He 6bL10 q.iepMCHTOB JJJ!Sl ero aKT1maurm. I1mjlCKI[l1H He npHB0· AII.ria K nopa)l{€'HHIO ,'lerKIIX. 3aMeHa O)LHOfI aMIIH0KHC.rIOTbl n
yqacTKC nape3amrn F-6eJIKa mipyca Ce11Aaii npusomrna K no HBJ1e1-11110 ycT011,mBocTH K Tp1rncIIHY [ Itoh M. et al., 1987].
MyTaHTbI nnpyca CeH,l!at1 n NDV rro F-6eJrny 06.riana.riH pa3Hof1 cTeneHbI0 IIHTonaTIITICCK0ro 3cl>q.ieKTa u pa3.rIHlJaJmcb no BIIpyJICHTH0CTII B 33BIICIIM0CTH OT cmrnaionieH aKTHBHOCTII F-6eJIKa, KOT0paH B CBOIO oqepe)J.h onpe,1l,CJI5IJJaCb npoTC0JIII·
TIJllCCKHM Hape33IIHCM 6e.11Ka Fo Kl!CTO'!Hb!Mll npoTeasaMH [Toyoda T. et al., 1987; Tashiro M. et al., 1988]. Cxo,11.nb1e
JWHHhie 6bIJlH noJiyqerrn n ,rvrn mipyca rraponna: F-6eJJOK mipyJieIITHOI'0 1UT[lMM8 0TJHlll3JlC5! OT F-6eJIK3 HCB1IpyJ1eHTIIOro
ruTaMMa 14 aMHH0KHCJIOTHhIMII ocTaTirnMH [Elango N. et al.,
1989].
118

llape3am1c F-6eJJ1rn 1rnpyca naparpmrna •rc•JioBcKa THna 3 l3 pa3Hh!X KJWro'!HL!X KYJlbTypax [fiyKjlHHCKaH A. f,. 1987].
Bnpyc BL!pait\en: l - n uepm,,1- noii KYJJhTypc; 2 - n Il pemrnae MhIX KYJlhTYpax IlO'l<'K o6C3bHH Vero H 3 - 4G/47; ·1 il 11\lli CyTCTllHH Hnn16wrorw Uf)OTCOJlll 3a rop;,oKca (50 Cll/MJij. u<:mrn 01:IHH.l_eUHOf'O BHpyca 110;:J.BC.pt'aJIH sJJeKTpo<jJopeay B 7,5% nAAL
Y BHpyca naparpHnna O1eJ10BeKa nma OH06Hapy}Ke11 cTpo rni% napa.TIJICJHI3M Me:IBJJ.Y uapc33HlleM F-6eJ1Ka 11 npOJJ.YKU.Hdi HHq:>eKilHOIJIIOro TJOTOMCTI 3a KJIC'1'KaM11 Ky.nr,Typbl TKaHII. 0 CTe IlCHll H3 C33IHUI F-6cJIJrn cy) IIJIII !10 3JICK'rpmpopeTH'!eCKOH Il0,l1.l3!LtKHOCTll 6CJIKOB QLIHIJJ.eHHOI'O 1rnpyca n I]AA IA]F-6eJJOK Ob!Jl BHJI.eH B BH,ll.C ,ll.BOHHOH riOJ!OCbl, o6a KOMIIOHCHT3 KOTOpOH IrnpbHpOB3Jll! no IIO/I.BHlKHOCTH B 33BHCHMOCTH OT KJleTKH-XO· 3HIIH3. Hatt60J1blllCH IIO)].BH}KHOCTb!O 06J18JI,:'1JIH 6eJJKH BHpyca, I1<WY,ICHHOro B KJICTK3X nepBH<IHO.H KyJibTypu flO'fCK o6C3bf!Hbl OaTeHBKxapaKTepH3YIOll.lJIXC51 M3KCHMaJibHOH 'IYBCTBIITCJ1b· HOCTblO K BHpycy, MHHIIM3JlhHOM - B H3HMeHee 11yBCTBH TeJibHbIX KJieTKax KJICTKax Vero. ITpoMe)KyTOtIHOe IIOJIO)Ke• HHe 33HHM3JI F-6eJIOK BHpyca, Bb!palllCHHOfO B KJ1CTKax nepe mrnaeMblX IJO'-ICK o6e3Millbl pbTCnOlKOTOpb!e npOAYU.HpOB3Jlll yMeperrnoe KOJllflICCTBO BHpyca (pnc, 9). CTeneHb reMOJIII33 TaK}Ke KOppeJrnpoBaJ1a c flO,ll,BHIKHOCTblO F-6eJIKa (:':lKCTHHKIUUI
npH K3Il HM) H COCT3BJIRJra Jl,Jlfl IHtpycoB, npOJJ,YllHpyeMb!X KJJeTKaMH MKC, 3ba3olH Vero, cooTBCTcrneirno IHcClIBICl H
IeIHKlIlo-BH,ll.HMOMY, pa3Jll1'fIUI B ITO,ll.BIDKHOCTH F-6eJ1Ka CBH33· Hbl m160 C era HeKoppeKTHbIM Hape33HlICM, r6M1lC TCM, 'fTO
Hape3am1e He A.OCTHfJJOKOHe4HOfi CT3,l1.IIH. B TOM CJJyqae, KOr· JLa BHpyc KYJibTHBHponamr B KJieTKax Ml(C B npHcyTCTBHII HJirH6HTOpOB npOTeOJJH3a rop,U.OKCa, KOHTpHK3JI3 Ii e-aMHHO· KanpoHOBOii KHCJIOTbl, o6a KOMIIOHCHT3 F-6eJIKa MHrp11pOB3JIH MC}J_J1CHHee, 1IeM KOMDOHeHTbI, H3XO)J.flIJJ.HCCH B COCT3BC CTaH-
OHBe

,11ap'l'HOro Bllpyca, JI T3KOI[ BHpyc OQJ!all,aJI pe3KO CHIUKeHHOH |
|
111Hj)eKu110HIIOCTbJO [Bukrinskaya |
G., 0lIC6ec)lo6anJ1e1rnc |
K KYJibTypc MKC ropJ1.0Kca (50cJ1./MJ1), 1rnH'rp11KaJia (5 eJ1./r,rn) HJ!H s-aMHHOKarrpoHOBOM KHCJIOTbl aH3Mr/r.rn) npllBO)],l!JlO K llO)],aBJiemno JJJiTOITaTll'clCCKOfO3q>cpeKTa BHpyca 11 CHIDKCHlIIO lIH(pCKUllOHHOH aKTlfBHOCTll Bllpyc,mro IlOTOMCTBa. YMeHbIIIa• JIOCb KOJII14CCTBO OaeB KYJlbTypaJibHO!i )Kl!JLKOCTH, CHIDK3JI3Cb fCMO.iH!TllllCCK35l aKTHBHOCTb BHpyca.
113 3THX )],3HHblX CJie)],yeT, l!TO Hlffll6HTOpbl npoTeOJIU3a MoryT C ycrrexoM HCIIOJJb30B3TbC.H B KJIIIHIIKC )LJ15I llO)],aBJJeHJrn nape3aHirn 6CJJK0B CJ!ll.HIIH.H napaMHKCOBIIpycOB ll 6JIOKHPO· B3HHH umpeKJJ.l!l!, YHHBepcaJibHOCTb cj)eHOMCHa rrpoTeOJIHTHllC· CKOII aKTllBarvm II cro YH3BHMOCTb ,ll,JI.H JI.CHCTBHH HHrH6HTOp0l3 npoTeOJJll33 Il03B0JJH!OT paccMaTpHBaTb ero KaK y,uo6Hyf0 MO· JI.CJib Jl,JIH XHMHOTepamrn p5IJI.3 m1pycHb!X HHq)CKI.!Jlll.
ToraBnpycb1. Y aJib<!rn-roraBnpycon nee CTpyKrypHbie H He crpyKrypttbIC 6eJIKH, raK iKe KaK II y IlliKOpHaBHpycoB II peT
pOBHpycoB, o6pa3yJOTC5l nyTeM npoTCOJJH1'll'JCCKOro ttape3aHIHI |
|
npCJI.lllCCTBCHHHKOB B |
K3aKTOB npOTCOJIH3a. llporeo |
JIH3 ocymecTBJIHeTCH B npouecce TpaHCJJ5lILHI1, ,Il.O OKOHtiaIUlH CHHTC3a nomrnpOTCHJI.a-npe,ullleCTBCHHHKa. flepBbIM OT npeJI. I.IICCTBCHHHKa OTpe3aCTC5l KancHJI.llbli1 C-6CJJOK. aOpoTCOJlH3 npOIICXOJI.HT B ytJaCTl{e C XHMOTpHIICHHOBOH CIICUilq>HtIHOCTblO
(rJ1y - Tp11 Cep) 11 ocymecTBJJHercH caMHM C-6eJIKOM no |
|
Trrny ayTOnporeomrna [Hahn bTcct cbeO3KMIyncMelancon 1pacGa |
|
rofi I-1., HcICBHc3aTeM npOHCXOJl,HT |
Hapesmrne 6eJJKOB 1HCc |
H Cn3no OCTaTKY 3JiaHHHa, KOTopoe ocymecTBJlHCT KJICT0l!Ha5l |
|
npon:asa, JIOKaJJH30BaHHaH B MeM6pauax. TiocJie H3JI.aJICHHH |
|
PE2 110HBJrnercH HOBhIH NHrKoneu |
rrenrnJI. c MOJieKyJIHP· |
HOh Maccoi:i: 6000, BbIIlOJJH.HIOlllHH qJyHKl.1_1110 CHrHaJJbHOfO nerr |
||||
THJI.a JI)IH I'JlflKOilpOTCHAa |
3HOcKOTOpbUaI BIIOCJICJI.CTBHH TaK)KC |
|||
0Tpc3aeTC5l |
H |
He 06Hapym:1rnaeTCH B |
3pCJIOM I'JIHKOTlpOTeHJJ,e |
|
E 1. HaKoneit, |
3aKJJI04JITe.fJbttoe nporeoJmTJiqecKoe Hape3att11e |
|||
IoT3Ha p3rJIHKonp(ne11J1,a |
BH311 yK3npoHcXOJI.HT yJKe rrocJJe |
|||
TpattcnopTa |
MCH3B nJJasMaT1it1ecKy10 MeM6pany H no BpeMem1 |
|||
COBlla,JJ,aer |
C |
Il0 IKOBaHHeM BHpycHblX |
!aCTHU. aOporeomrn B |
|
|
|
a |
|
O |
9TOM CJJy,rae ocymeCTB,'15ICTCH KJieTO'lHOI{ rp1rnc1mono,IJ.06HOH |
||||
3H,Il.onenTIJJI.a30IJ, accou_1rnponaHHOH C MCM6paHaMH 3HJl.0IlJJa3- |
||||
Manil!CCKOro peTHKYJlyMa |
(CHfHaJibHaH rreIITHJI.a3a) B ytJaCTKe |
|||
Apr -- Apr |
|
Cep. |
|
|
O;THOBpCMCIIHO C Hape3aHHeM npoHCXOJI.HT KOHCjJOpMaJJ,HOH· Il35! nepecTpOHKa I'JlllKOilf)OTeH)\0B B IlJJa3MaTll'ICCKOH MeM6paHe, 5IBJIHIOUJ,,lHC.H, 110-BHJU!MOMY, IIYC.KOBblM ClffH3JlOM AJIH f!O'IKOBaHIHI BHpycHbIX 1IaCTHU. 3aKJl!Ol!HTCJJbHblH npoTCOJlHTH· l!eCKHn a1<r - uapeaam1e rrpeJI.mecrBeHHHKa 1HocHa BcrJIHKO·
120