
Букринская А.Г., Жданов В.М. Молекулярные основы патогенности вирусов
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3,ll,eCb )Ke yMeCTH0 CK333Tb HeCKOJlbKO CJIOB O BHpyceMIIII II ee p0JJII B naroreHe3e BHpyCHblX IIHcpeKUIIH. PacnpocrpaHe HIIe 3TIIX umpeKUIIH MO)KeT npOHCXO,ll,IITb nyTeM KOHT3KT3 C 1rneTKaMH (raK pacnpocrpaHHeTC51 repnecBIIpyc Ha K02Ke II CJI113IICTb!X o60JJO'!Kax)' C BbI,ll,eJJeHII51Mll CJJH3IICTblX 060J10qeK (TaK pacnpocrpaII51IOTC51 pecnIIpaTopHbie II KHillelJHbie IIHcpeK u1rn), IIO HCpBaM (TaK pacnpocrpa,rneTC51 6ellleHCTBO, a y Mbl weiI - IIeiiporpomible peoBIIpycb1). O,ll,HaKo qame Bcero IIMeeT MeCTO BIIpyceMII51, a IIMeHHO 3TIIM nyTeM BIIPYCbl pa3HOC51TC51 IIO opraHII3MY II Hepe,ll,K0 np!!H!IM3IOT BT0pllqHyIO JI0K3Jlll33UHIO. KJiaccIIqecKIIM npIIMepoM TaKoiI JIOK3JJII33UIIII BIIpycnoiI lIHcpeKUIIH npII reMaToreHH0M nyTII pacrrpocrpaHeHH51 CJIY)KIIT IIOJJIIOMIICJJIIT. I1epBllllH351 JJ0K3JII133Ulf51 lIHqJCKUHII npoIICX0- ,ll,IIT B 3f!HTCJIII!I TOHKOH KlllllKII. 3,ll,eCb BHpyc pa3MHO)K3eTC51 rrepBII'IIIO II B f!O,l],aBJ151IOIUeM 60JJbllillHCTBe CJiytJaeB 3apa)Ke1111e IIMCHHO 3,ll,eCb 0K3HlJIIBaeTC51 HH<peKUHeH, TaK KaK Haq11HaeT ,ll,eHCTBOBaTb CHCTeMa mnepcpepoHa II p33BIIB3IOTC51 IIM MYHHbie peaKUHII - HIIcpeKUH51 JJOK3JJII3YCTC51. O,1I,H3KO M02KeT pa3BHTbC51 BIIpyceMH51 II TOr,ll,a BIIpyc MO)KeT BTOpIIqHo JI0Ka JJII30B3TbC51 B UHC - B KJieTKax nepe,ll,HIIX p51,1I,OB CIIIIHHoro M03ra, a T3K)Ke B npo,1I,OJirOB3TOM M03re. Bo3HIIK3IOT napaJIII
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27

'1'11Ta HO pOBhl3b!Ba10T nopa)KeH1u1 nmueBapHTCJ1bHoto tpaK• Ta. P5l)l, rrnpycoB Bbl3h!BaIOT CHCTCMHbie nopa2KeHJIH KpOBH (peTpoB11pycb1), ne<JeHH (B11p_ych1 renarnTa AH B), reHepaJIH30B3HHbie IIHlpCKUHH (6oJIL,ll1HHCTBO ap60B11pycoB, BHpychI KO pH, KpaCHyxH, BeTp5IHKJI, ocn1,1 11 J1.p.), UHC (011pyc1,1 TpaHc MttCCHBHblX 3HlLeq:>3J1HTOB, TIOJll-iOMHCJlHTa, B036Y.ll.HTCJ1H MC,ll.· JICHHblX HHcpeKUHH, BHlJ).
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pOHBorpaHH<JCHH51 OTHOC51TC5I H K Kpyry nopa)l'<:aCMbJX KJle TOI<. Tat{, unpyc1,1 TIOJIHOMHCJIJITa lI JJ,pyrne 3HTepoBHpycbI MO· ryT pa3MHO)KaTbC51 B 3TIHTeJIHH KlllllC'IHHKa, II04eK H B HepB HblX KJleTK3X. B11pycbl rpmma '-!CJ!OBCKa, nopa;,1rn51 B eCTeCT BellllbIX ycJJOBH5IX TOJlbKO JJIO)l.CH, MOryT pa3MHO)KaTbC51 B KYJlbTypax KJ1CTOK MHOfHX 6HOJlOf!JlJCCKHX BH)J.08 - lJCJIOBeKa, o6e3b5JH, co6aK, IITH'IbJJX 3M6pHOHOB. Mnorne ap60Bupyc1,1 pa3MJIO)l{aJOTC5I B KJICTKaX K3K Tell.7JOKpOBHLIX )KlIBOTHblX, TaK l1 •!JICIIHCTOHOfHX. B TO JKe npeM5I BHpyc 6emeHcTBa pa3MHO )KaCTC51 TOJlhKO B HCpBHbJX l{JJCTK3X, 3 Bl11I - B MaKpoqrnrax H T4-J1HMQ)OUHT3X, a TaKiKe B K.1eTK3X M03ra H HCHpOrJlHll.
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28

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