e opraHH3Me
cJJeJJ,yIOl.l\asi [Varchoan R. et al.,
1989]
THMH,ll.[!Jl8TKH
A3T-f\H·
m1pHMHf\HH
A3T-rpmjJocqiar
Ha3a
---
(p0C(p3T
HYKJie03Hf\·
f\HCpOC(p8T·
KHH838
UHTHJ\Hl!M0H0-
J\J\U-!\mpoc
!IHpHMH,!(HH·
MU-rpHqiocqiar
cp0CQl8TKHH833
qiaT
HYKJIC03llf\
-·-
f\ll(jJ0C(jJaT·
KIIH838
----- -·········--
)lHJJ.eOKCHUJITH,!]JIH 11 JUIJ COKCHHH03HH 60.nee 9qiqJCKTHBHbl, lfCM 33H)l.OTIIMil,!UIH, OIHI OJ{33LIBaJOT MOil HOC 3HTI1BHpycuoe )l,ClfCTBHC KaK B T-JIHMcpo1urrax, T3K H 13 MOHOJUITax. 9nI npe
napaTbl H ;J)l)l.COKCI!a)l.eH03HH npoXO)l.HT KJiliHI!4CCKHe HClll,t· TaIIIl5l [Perno C. F. et al., 1988; Jarchoan R. et al., 1989;
Merigan T. ct al., 1989]. EmihlllHM npenMymecTBOM nperra paTOB 5IBJIHCTCH I!X 3<p<fieKTHBHOCTh npH nepopaJJhlIOM l3BC)l.C Hl1H. )lpyrIIMH aHOM3JibliblMH HYKJIC03IUl,aMH, npHMCHHCMhlMII npa Jiel !eHHH CTII1JJ,, 5IBJIHIOTCH a11,HKJIOBHp II pn6annpn11 (Ta6.1. 21). PH6aBHpHH-aHTHBHpyCHblH areiiT urnpoKoro CITCKTpa )l.Cl1CTBIIH, npHMCHHCTC5I B BH)l.e a::ip030JI5I ,ll_Jl5I Jiel!e HH5I BHpycHI)IX pccrnipaTopHblX fIHqJeKlUifl. OH 110)],aBJIHCT pe IIJIHK311,HJ{) BI1l I n oKa3LIBaeT anT1mnpyc11bl?i 9cpcflewr B opra lili3MC 60J1LHLIX CTII1)l. Ero JWCTOHHCTBOM 5IB,1 u1eTc5I npo HHKHOBenHe qepe3 reMaTO:rnll,ecpaJIH'ICCJUll"l6apbep. 0)l.H3KO OH Mcnee 9qicpeKnrneH, 4CM J].H,ll_COKCHHyK.1C03H)l.bl, H Bbl3blB3CT no601 rnue 9(p(p€KTbl (am:MH5I, 6eccOHHIHl,a, fOJ!OBHble 6om1, p33,ll_p,LiKiiT€J1bllOCTb)
T a 6 JI H II. a 21. cJ>apMaKOKHHeTHKa HCK0T0pblX aHa,1oroe HyKJJe0311)],0B npH nepopaJJbH0M npHMetteHHH npH JJe"leJIHII Cn 1-f)l
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AnTH1311pyCHUM }leiicTBHeM MoryT o6JrnJI)ITL npenapaTbI, n11rn611py1outr1e (!>YHKILJIIO PHKa3bI I-I u Hapymaiouuie JJ.erpa JJ,a11mo PBK B PHK - ,JJ,I IK-rn6pHJLe, a T3K2Ke npenapaThI, 11Hrn611py1011\l1C <!)YHKI\IIIO IIIITCrpa3h! H 6.ilOKHpy10uu1e HIITC rpaumo 11poBI1pyCHoii ,[]JIK B xpoMocoMy. T,lKHe npenapaTbl H0Ka T0JII,I{() C03Jl_aJ0TCH.
TTepcneKTHBHOii MHIIlCHh!0 JJ_JJH XIIMH0TCpanerHH'.JeCK0ro BMewaTeJll,eTBa 51B,i lf!J0TCH Tp3HCaKTHB3T0p Tat H ,upyrne TpattcaI<THBaTopb1 np0Mo1opa BI1 1 I. Tpanca1mrna1op Tat npC,llCTclBJJ}lCT co6oii MaJ1e1IbKH!1 6CJIOI{, COJlCp}IGHUHii 86 aM11H0KIIC,II0TllhIX 0CT3Tli0B. MoJICKy.na C0,D,epnrnT 06.nacTb 0CTaT K0B 0CII0BHblX aMHH0KHCJI0T, 06ycJIOB.mJBaIOII1,HXTpancnopT 6em<a B 5-lilp0, II 6oran,1h0CTaTIUlMII 101CTCHHa Y'JaCT0K, Ba)l{ Hb!H )l,}151 o6pa30B3HHH )UIMepcrn. ,ll.r rH IIHI'H6IIWlll TJ)3HC3KTJI· B3I]J!II no.11yt1eHbl CllllT('Tll'ICCK!IC TICllTIIJ(bl, C00TBCTCTBY!OlllllC
6oraToMy 0CT3TK3MII J IICTCH!ia ytJaCTKy. I lcnTHJlh] 6JIOKHpo B3.JIH 3KTHBH0CTh HToCIaCp0oKeC B pe3y.llLTaTe o6pa3onamrn. re
TCJ)0]UIMCJ)0B C flaT!IBHblM 6CJIKOM, 0JJ,IIaK0 ,11,eiJCTBHC HX 6bWO
HecneruHpw1ecK11M, II nenTIIAI>I 1rnru611pona,n11 Ta1OKe rrp0Mo1opb1 p:.iHHl!X remm OB40 [Frankel A. ct al., 1989].
TTpenapaTLI, K0IIKY[Hipy10IIJ,He C TJ)3IICaKTHB3T0p0M rev, 6y/lYT 6.JIOKHposaTb 1pa11cnopT B IlHT0!IJJa3MY MPH!\ Jl,TIH BH pyc11ux c1py1nypm,1x 6eJIKOB. TaKIIM )KC ffy'TCM MOryT 6bITb 11aii,D,eHbI npenapan,1 JJ.Jrn 6.110K11ponamrn {l)YHKHHH ,n.pyrnx pe ry.i!HT0J)HbIX reli0B. TTpn 3T0M C.TI02KHOCTb rCHCTH'ICCKOl1 CTpyK TYJ)bl II perymnurn BHLJ npn ycJIOBHH pacm11q)pOBKH ee Mexa• HH3M0B M0)l(CT o6paTIJThCH IIJ)0TIIB caMoro Bnpyca H nprrneCTII
K ycnem110My BMC'JU3TC'JlhCT l3Y B
lfll(j)CKUIIOIIIII,Ij:\
npouecc.
C 1 a ;1, II H T p a H c .11 H IJ, H 11.
PeTpom1pyc1,1
HCIT0Jib3YIOT
He06WJHblC MCXaHH3Mbl ,ll,/!5! 9Kcnpeccrrn HCK0T0pblX reH0B. TaK, Olm MoryT CHHTC3Hp0BaTb CAHHblH J1HI1011J)0TCHA gag-pol c ,D,BYM.5I 0T,UCJILHLIMH paMKaMH C'mn,rnamrn MPHK. )1,Jrn Kop peKTHoro cqHTbIBaHHH Tpe6yeTCH, lJT06hI pH60COMbl C0Bepurn J1IJ C)].Blff C 0,11.HOii paMKH Ha JJ.pyry10 - npo ecc, He06hl'lHbll1 /lJIH 9yKapH0TI1'1CCKIIX KJ1CT0K. TeopeTHqCCKH B03M0}l(H() II0JIY11HTh cneruHpI!lleCKHe XHMH0npenapaTbl H.'lH aHTH6HOTHKO no;1.06HLie areHTbI, K0T0Jlbie MoryT 3a6.no1rnponan, npouecc pn6oCOMHOro CJlBiff3 paMKII II 1rnpy11mTh 9Kenpeccmo BHpyc HblX reHoB. AreHTbl, 1mTepq>epupy10m,11e c MPHK B11pyca B 33p3,KCHHb!X KJICTK3X, T3K)l{C 6y;1,yT 6.iJOKilj)OB3Th peIWHKa11,HI0 BHpyca. Bo3MO)KHblM aHTHBHpycHblM npenapaT0M, ll.CH CTBYJOW.HM Ha 3TY MHUICHb, HB.llHCTCH pn6arrnpm1 (BHp330JI), amJJior rym1031rna. A1rnmnpyc1-1btMII cnoi'icrnaMn MoryT 06J1a ,11.aTb K0J)0TKIIC <ppiffMCHTbl ,UHK, [{01\,ffl,'lCMCHTapHLIC )K!I3IICH H0 133)1{!11,!M Y'l!lCTK3M BnpycHoro reII0\1:l. flyT(''' CRR3Ll!HlH!l5{
c HHMH cpparMeHTLI )lHK 6yAYT no)l.aBJUITL 11x Q)YHK11mo HJIH
MaCKIIpcrnaTb yqaCTIUI
CBfl3bJBaHJU:I perymrropHb!X 6eJIKOB.
K Q)parMeHTaM MoryT
6LITb rrpHBH3aHbI CHJibHOJJ.CHCTsyrouvie
XIIMH'lCCKHe rpynrrbI, Hapywarom0e CTPYKTYPY Mo.ncKyJibI HYK·
.'ICHHOBoi1 KIICJIOTbl [Goodchild et al., oCll0el AHTHCCHC·OJlllfO)l.COKCHHy1,JieOTH,ll.bl ll HX aHaJiorn MoryT
l!Cl10.1b30B3ThCH }])!fl IlO,ll.aBJICHIIH pen.JrnKaJ Hll BHl J, 6JIOIUI· ponamrn crr.naticnnra II TpaHc.nm1,11H MPHK, peryJrn11,1m 3KC· npeccm1 reHon. I1oKa3ano 1rnrn6ttpy1ou ee ,LI,CHCTBHe pHJJ,a
()J!IffO)lCOKCI!HYJ{.JJCOTl!Jl,OB II llX cjioccpopocyJlbQ)l!f)()I3aHHblX aHa-
Jioron Helpenmrnaw110 c3Jl pbiYpflnXl!Blylal., oClCplMat
sukura M. ct al., oClC6Il
3 a !( JI IO lJ II T e JI b H I,[ e
C T a )l II II
H H (fl e K II II O H H O r 0
II H K JI a
B H p y C a.
0Hll CBH3aHI,I C MO)l!HfHIK311HCii 6CJ1KOB,
c6opKOH
Bnpyca II
Bb!XO)].OM
II3 KJJCTl{I{
rryTeM HO'!K0B3HH½.
Ha 3THX cTa)l1rnx T3KJKe MoryT )leHCTBonaTb cne1vHjrn'lCCKHC 1rnrn611TopL1. Ee.nKH BHlJ, KaK y,Ke OTMe•ra.110c1,, 06pa3y10-rcH B pC3YJibT3TC uape3aHHfl rrpeJJ,rneCTBCIIHHKOB npOTCOJlllTll'IC CKHMH cpcpMCHT3MH - KaK BIIpyccHCIUHj)ll'lCCirnMH, TaK H 1rne TQqHb!MII. B 3apa1KCHHblX BHpycoM KJICTKax HaX0,/1.HTCH Bnpyc Cl1Cllll(fllJIICCKaH rrpcJTea3a, KOTOpaH 33KO,ll.HpoBaHa B TOM ,KC re11e, •rro 11 06paT11att TpaucKpnnTa3a, 11 Hape3aeT npenrnec-r
BCHHHKH gag 11 pol. KaK 11 11p0Tca3LI npyrnx peTpOBHpycos
[Pearl
Taylor TaHO nlClM3al OHa OTHOCHTCfl K acrrapnmOBblM
rrpOTC33aM.
f1poTCa3a HBJ!flCTCfl llOTCHJ IHIJlbHOl I MHlliCHblO
JJ,JIH JI.Cl'ICTBHH CIICUHQJH'ICCKHX HHrn61nopon, HO)l06m,rx rren CTanrny [Katoh Hl ct al., oClM3Tl o,IIHaKo KOHUeHTpaurrn rren
c-ranma A, Heo6xon11MaR ,/I.JUI noJJ,aB.nemrn aKTHBHOCTH C3JOl Ha HCCKOJlbKO llOjnIJLKOB BhIWe, qeM J{()HUCHTJ)311llfl, IIOJJ,3B
Jlfl!OUJ,aH aKTHBHOCTh rreuc1rna H pemrna [Krausslich
H. G.
et al., oCll3Cl pH,IIe ,1a6opa-ropHii
ll,/1.CT aKTHBHbll'I
IIOHC!(
1:rnrn6HTOJ)OB acnapTI1HOBOI1 lIJ)OTC33bl
rC1JKl 06.na)la1omHx aH
THirnpycHOl1 aKTHBHOCThIO. AKTllBHbl!f IICHTp lll-IrH61nopoB npeJJ,CT3B.TTCH ClllITCTHqecKHM IICIITH)l.OM; )Urn 33LQHTbl OT KJlC TO'lHb!X 11pOTCHHa3 TICIITH)l. KOWh!OrHJ)YIOT C yrJICBOJ].Hb!Mll IIC· noqKaMH; )l.Jlfl rrpOHHKHOBCHH}l B 3apa)KCHHhIC CIHl KJH:TKII Ilj)HCOC)l.HHHIOT y<JaCTKH, Y3Harom11e BllPYCHb!C 6elIKH env Ha IlJia3MaTHYCCKOfl MeM6paHe; B OCHOBHOM IICCJ1CJJ,OB3HIIH BC· Jl.YTCH Ha 611on.'XHOJIOrHqecKOM 1rnrrpaBJICHHH.
06napy)!{CHO, l!TO anrnrpII6KOBbiiI 3HTH6HOT!lK 11epy.ne1rnu 06.'la)l,aeT aHTHilpOTC33HOH aKTl!BHOCTh!O ll 1mru6npye-r pe llJ11IKa1111JO HCKOTOpb!X peTposnpycoB. A11n16tIOTim 6.HOKHpyeT pe11JIHK ll(IIJO B111-f rryTeM 11apyrncmrn ll!)O]lCCCIIHl'8 6C,1KOB. )l.rrn HHr11611ponaHIIH npclTea:rnoii <IKTI!l3HOCTII cyni,eeTBCITlfOC :.Hralfe[llIC l!MCCT 3IlOKCU)IHOC KO,ilhll,0 B MO:IcKyJlC:' aHTl!OllOTlllrn.
Puc. Pc.u ,UeilCTBHC JJHnOCOM ua npotmKHOBCHl!c n KJieTKY Bv!L J
(ilaHHhlC TO'IC1 JIIOii
rn6ptt,!l.H3a
UHB)
[TCHUOD IO.
IO. H J\P-,
1989].
K
JIHM<pOH)I.HOll BHpycnpO/.\ YU.HPY!O·
w;el\
JIH!JllH H9/I I lB ).(0 6aBJI8JtH Jill·
II0C0MLI HS 5lH'-IH0ro JIClI.,HTHHa 8 KOHU.e11Tpa1lmI JO MKr/MJt no JtHIJII· ;r.y; Ha 3-7-e CYTKH onpc;r.eJIRJIII KOJt!l'!eCTllO HYKJICHHOBb!X KHCJI01 MCT0)l0M Toqe•rnoli l'H6pH)lH3,Ul,Hll,
f I - K0HTp0,% - KJieTKH 6e3 .n.06a11- J(eHHll JIHUOCOM; !I - JIHllOCOMbl ,11.0· 6aaJ1eHbl K K. eTK3M B 11epHO)I. C 3-.x no 7-e cyTKH 1111Ky6au,HH (coorneT· CTBCHHO 1 5); a-r-TpexKJlaTHbl': paane,u.e1-11u1 HYKJieHHODblX KHCJIOT III - KOlrl"J)OJtb '!YllCTllHTCJlbHOCTH MeTO)l.a c HatteceHHeM I, 5, 10 u 100 nr nHpyccnc.u.11<1nJ< 1ecKoii .D.HK
TioJiytieHbI CHHTen1t1ec1rne rrenTH)l.bl, HHrH6Hpy10m.He npoTe0.1JHTHqecKy10 3KTHB H0CTb B114-2 H o6Jia)l.aIO ,une MCHbmerr T0KCHH !H0-
CTb10, 11eM uepyJ1emrn [Blu menstein J. et aI., e0T0HHa
3TU l-1HrH61nopblHe BJIHH· IOT Ha cpH3HOJIOrHqecKH B3)KHb[e JI,JIH opraHH3Ma acnapTHH0Bb!e npoTea3hI (B qacnrocn1, Ha pemrn) H npe,n.cTaB.TISHOT co6011 KJiacc areHT0B nponm Bl1l!-11mj.>eK1I,HH. MHpttcTHJIHpo baHHe nomrnp0TCH,ll.a-rrpeJJ.mecTBCHHHJ(a gag T3!OKC MO)KeT 6b!Tb MHmeHbI0 )I.JIH XHMH0TepaneBT}ll !CCI(0ro BMemaTeJlbCTBa, npliB0LLHW.ero K 6.110KHp0B3HHIO rrpou:ecCHHra 6eJIKOB. PeTpO· BHpycbI Bb!X0,TlHT H3 KJieTKH rryTeM IT0'lK0BaHI-rn, K0T0poe MO· 2KCT 6bITb Il0}laBJICH0 HHTepcpepOHOM H ero HHll,YKTopaMH H an TlffeHaMH, B33HMO,[\ettcrny1om.HMH C KJieT0'lHb!MH JIHITl-i,TlaMH.
fJTHK0I1p0TCH,ll.bl BHpyca Hape3aIOTCH KJICT0'lHbIMH TpHTICH· HOllOJI,06Hb!MH npoTea33MH, H cneu:i1cpHqecKHe HHrH6HTOphl 110)],06Hb!X rrpoTeas - anp0THHHHbl - MoryT 110)1.aBJISITb HX aK TliBH0CTh. Mb! Il0Ka3a.1H, H IT0 penpo.n.yKl..(HH Bttpyca 110)1.3BJH!· eTCSI npH A06aBJICHHH rop)l.0KCa H K0HTpHKaJia K BHpycnpo jW 1J,HPYJOW.et1 JIHI-llrn JlHMcpOH,IJ,HbIX KJieT0K I-I9/IIIB. Arrporn- 1-111Hb1 npHB0.ll.HJIH K yBeJIH•{eHHJO qHcJia )KH3Hecnoco6HblX KJleT0K H YMCHbllleHHIO e !HCJJa KJ1eT0K, co.11.ep.lKalll.HX BHpyc crreuttcpHqecKHfl aHTHreH llJ)H HerrpHMOH HMMYHOcpmoopec1J,eH lI,HH (pHc. THHp C. 208). MetteeeHT aKTHBHb!M HHf116!1TOpOM 0K333· Jiacb AKK [ByKpttttcKa5l A. AP-, 1989].
OcG>6eHHOCTbIO rJHIKonp0TCHJI,0B Bl-14 HBJIHeTCSI HX HHTeH rnnnoe fJIHK03HJIHpona1111<:. 13eJIOK gp T0ep C0JI.ep:t!{IIT 0ap IT0TCI-l·
eHT0
unaJibHbIX yt-IaCTKa rJI!iK03l!JmpoBaHIHI H II0l!0BHHa ero M0Jie
KYJIHpHOH MaCCbI npHX0AHTCH Ha yrneao,rl,1,1 [Ratner rrB1 et al., oCcI6H1 0 poJJH yrJieB0A0B CBH,LI,eTem,cTByeT Ii TOT cpaKT, 'ITO
cao6oAHLIH gpl20, scpcpeKTHBHO KOHKypHpy1ow.Hii sa peuemop C HaTIIBHblM 6CJIKOM B C0CT3BC BHpyca, TepHeT 3T0 CBOI°icTBO rrpH oTcyTcTBHH caxapoB B ero MoJ1eKy.r1e [Putney HHa et al., oCcMl1 Matthews ya1 et al., oCbO6p1 I1HTCHCHBHO rJIHK03HJIHpoBaH Ii 6eJIOK gp4l. B CB513H C 3THM Hapywemie rJlliK03H.'IHp0Ba llHH M0il(CT 6JIOKHpoBaTb cpy11KI..1,HIO 6eJIKOB H HapymaTb npo uecc rrpOH!lKH0BCHHH BHpyca B KJieTKy. l13BCCTHI,I no KpaitttefJ Mepe 5 rpynn MyTal.l,HH, K0T0J)h!C HapyruaIOT cnoco6HOCTb rJIH· rwnpoTeHA0B K o6pascmamuo omu11THH [Kowalski M. et al., oCcO61 Ii CBHJleTCJlhCTBYIOT 0 CJI0)l(H0M MCX3Hll3MC 3T0f0 npo uecca, BKJIIO'I3IOII.l,CM rrpaBHJlbH0e rJJHK03HJ1Hp0B3HHe env 6eJI K0B. Tio,ll,aBJJCHl!C rJIHK031iJIHpoBaHHH M01KeT np!!B0)]JiTh K Ha pyrneHHJO cpyHKI.l,HH H TpaHCII0pTa 6eJIKOB K IIJ133MaTifl!eCKOH MeM6paHe, a nperrapaTbI, HHrn6Hpy10mne rm1K031!JI1Ip0Bam1c, 6YAYT oKa3bIBaTb aHTHBHpycnoe JJ.eikTBHe. TaKHe 1rnrn60.To pbr rJIHK03HJIHp0BaHIIH y)l(e orrnca111,1 B JHITepaType, K HHM 0T H0CHTCH N-6yTHJl)leOKCHHO)l(HpHMHI..1,llH, N·MeTHJl,ll,e0KCHH01Kl1•
pHMHI..1,HH, N-3THJI)leOKCHHO)l(HpHMHI.l,HH, l,4-mmeoKCH-1,4-HMH· Ho-1-apa6mrnToJI n AP- [Karpas 3e1 et al., oCbb601
AHrnm1pyc11bii1 c.>(jJlj>CKT rrpn ;1.06anJ1emm K Bnpycrrp0AYUII· py101udi .1ll lll!H JrnM<jJOH,llllblX KJICT0K np0HBJUieT H IIHfH0HTOp
6nocmne3a 1KllpHblX KHCJIOT uepyJICI-IHH, KOTOpbl11 II0)l3Bll51.CT MHpHCTHJ!Hp0BaHHe IT0JIHIIpOTeH)la gag [Pal HHa et al., oCbb6a1
O)1,HaKO KJIHHHqecKoe rrpHMCHCHHe nperrapaT0B C YK333HHWM MexaH!I3M0M ,ll,el!CTBHH orpaHH'IeH0 HX B.'IHHHHCM Ha 6R0CIIHTC3 KJICT0qHblX M0.'ICKyJI.
Ilpoxo)J.HT KJIHHil'ICCKHC HCilbITaHHH anbcpa-HHTepcpepOH H ero peKOM0HIIaHTHb!e aHaJiorn. I1perrapaT ACifCTBYCT Ha He CK0J!bKHX CTa,ll,HHX penmrnal.l,H0HH0ro I.l,IiKJia, BKJll0'Ia51 TpaHC·
.'IHI.l,HIO ll Bb!X0A 113 KJ1CTKH. Onnca1-1 HOBbill MexaIIH3M )leiicr BHH aJibcp3-HHTepq>epoHa - HHrH0HpOBaIIIIC Il0tJK0BaHJrn BHpy
ca c KJieTo,rnoii noBepxttocTH [Hartshorn HH et al., oCbCH1 Yarchoan R. et al., oCbJrT1
I1ocKOJibKY TtXH1 OTJIHqaeTcH ymrnaJJbHhIMH qepTaMH, He o6bl'IHbIM rraToreHC30M, II0AX0)lbl K JIClJCHliI0 ero TaK)l(C )l.0.'l1K· HhI 0b!Tb 1-1ecTaH)lapTHblMH. OHH )lQJI)KHbl np0B0)lHTbCH a Tex H0BbIX HanpaBneHHHX, K0T0pbie 0CI-IOBal-!bl Ha H3Y,ICIUIH M0Jie KYJIHpHb!X rrpoueccoB rrpH perrp0AYKUHH BHpycoa. yK1 M. )K)la H0B O0bC,ll,HHHJI I-IX II0,'l na3BaHHeM «He1'p8JlHUH0HH3H XHMH0TeparrHH BHpycHb!X HHcpeKI.l,HH».
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