EeJIKOBLie TOKCHHbl (3K30TOKCHHbl)

6eJIKOBbie T0KCHHI,I qame npo.zzyu;11pyJOTcjJ rpaMllOJI0)KIITCJII,HI,IMH 6at<TepHHMH, HO TaK)Ke BCTpe JafOTCj{ y rpaM0Tpnu:aTCJII,HI,IX 6a1<Tepnu KaK a3po6HbIX, TaK H aHa 3p O0 HI,l X.

 

/(llR 6eJlK06blX moKCU1l06 xapa«mepno:

Bbipa)KeHHaH T0KCH'fH0CTh, TO eCTb noBpe)KJJ:alOI.Qee)J,CHCTBHe Ha 3yKapnoTnqecKHe KJICTKH B MHHHMaJihHbIX K0Hll;CHTpau;HHx; cneumpH<iHOCTb.ueHCTBffj{ - pa3JIHqHI,IC T0KCHHbl, KaK npaBHJI0, BI,I- 3bIBalOT pa3BHTHe l<JlHHM'JeCKH 0TJIHqaJOI.QHXCH CHHJJ:p0M0B (K0MllJleKC xapaKTepHbIX KJIHHH'ieCKHX CHMIITOMOB);

HaJIH<JHe q>epMeHTaTHBHOH aKTHBH0CTH, CBOHCTBeHHOH MHOrHM T0K­

ClfHaM, qTo o6ycnasmrnaeT cneu:uq:m<JHOCTh MX.nei1cTBHH; npeHMYI.QeCTBCHHaH TepMorra6HJibHOCTh, HO BCTpelfafOTC.ll H TepM0- CTa0HJibHble TOKCHHhI;

Bbipa)KCHHaH HMMyttoreHHOCTb - CITOCO0HOCTh Bhl3bIBaTh 06pa30- BaHHe cneu.m:pHqecKHX aHTHT0KCHlfeCKHX aHTHTCJI (aHTIITOKCHH0B),

scrynatol.QHX c ToKc1rnaMH B peaKUHH ttewrparrn3au:1m; CilOC06HOCTI, HCK0T0pbIX 0CJIKOBbIX T0KCHH0B nepeX0.UHTh B aHaTOKCH­

HhI,.UJIH K0T0pbIX xapal<TepHa Il0TepH T0KCH'IH0CTH npH coxpaHeHHH HMMYH0r eHH0CTH.

 

ITo cmenenu C6R3U C oaKmepUOllbHOU KllemKOU 6eJIKOBbie T0KCHHbl

.nemTT Ha TpH KJiacca:

K. Jtacc A - moKCUHbl, ceKpemupye.lwble eo eHeu1:n1010 cpeoy. IlpHMe­ poM.IIBJI.IICTCH.D:Hq>TepHHHbIH T0KCHH, KOTOpblH npH pocTe 6aKTepHH

0TUIHypoBbIBaeTCH OT nosepXH0CTH KJieT0K B BH)le ny3bipbK0B H

nocrynaeT B 0KPY*aIOI.Qyto cpe.zzy.

 

45

 

KJ,acc B - 11acmwmo ceKpemupyeMble moKCUH bl. EoJJbllJa}l 'l:aCTh TOKCHHa 0CTaeTC}l BHYTPH 6aKTepHaJihHOM KJieTKH, a qacTh T0KCHHa BbT)].eJrneTC}l H npoHHKaeT B KJieTKH MaKpoopraHH3Ma, HanpHMep 60-

'I)'JIHHH'leCKHM T0KCHH n CTOJI6H}l'l:HblM TeTaH0CTia3MHH.

K;1acc C - HeceKpemupye.Mble moKCUHbl, K0T0pb1e ocso6o)J()].aIOTc}l

T0JibK0 nocne rn6emr " pacna)].a KneTKH-npo)].yUeHTa, HanpHMep lllwra-TOKCHH H urnraII0)].06HI,Ie T0KCHHbl.

 

B 3Q6UCUMOcmu om MOJleKJ!JIRpHOZO cmpoeHUR 6eJIKOBhle T0KCHHbT 6h1BalOT np0CThIMH H CJI0)l(HblMH. IlpoCTbTe T0KCHHbl npe)].CTaBJI5IIOT co6ow 0)].H)' nommerrrH)].H)'IO uenb Cno:>KHbie T0KCHHhl TI0CTpoeHbl H3 CB}13aHHbIX Me)J()].y co6oii HeCK0JTbKMX nommenTH)].HbIX uenew, HeK0T0pble BKJIIO'lalOT

)].0TIOJTHHTeJibHbie K0MTI0HeHTbl.

llpocmbte moKCUHbl Bbipa6aTbIBalOTCSI 6aKTepHSIMH s BH)].e HeaK­ THBHbIX npoTOKCHH0B. OHH aKTHBHpyroTC}l (npno6peTalOT TOKCHqHOCTb) 6aKTepHam,HbIMH npoTea3aMH mm npoTea3aMH)KKT, KJieToK H TKaHeii

MaKpoopraHH3Ma. B pe3yJibTaTe M51rK0ro (orpan11qett1-1oro) npoTeonma IIp0HCX0)].HT pacm:ennemte TIOJIHIIeTITii)].HOM uenH T0KCHHa Ha)].Ba cppar­ MeHTa, 0CTalOIJ.IJIXC51C0e)].HHeHHI,IMH Me)J()].y co6oii)].HCYJib<pH)].HOH CBll3blO.

O6pa3yeTC}l 6mpyHKUHOHaJibHa51 AB-CHCTeMa, C0CTO}lW:ruI H3 cpparMeHTa AH cpparMeHTa B.

<l>parMeHT A (active) - o6Jia,[(aeT CBOHCTBaMH cpepMeHTa H HMeeT

KaTaJIHTH<{eCKHH u eHTp. 3TOT cpparMeHT-aKTHBaTop 0TBeTCTBeHeH 3a cneuHcpHqecKoe TOKCH<{eCK0e,[(eHCTBHe Ha KJieTKH MaKp o opr aHH3Ma.

<l>parMeHT B (binding) - o6na;::i:aeT CB513bIBarom:HMH H TpattcnopT­

HbIMH cpyHKUH51MH, a TaIOKe HMM)'H0reHHblMH CBOHCTBaMH. AB-CHCTeMa

npmcpenmieTC}l K qyBCTBHTeJibHOH 3yKapHOTH'leCKOM KJieTKe C Il0M0W:blO JIHraH;::i:-peuerrropttoro B3aHMo;::i:eiicTBIDl Me)J()].y peuernopoM cpparMeHTa B H C0OTBeTCTBYlOW:HMH peQernopaMH KJTeTirn, qam:e Bcero raHrJIH03H,UaMH HJIH JIHTI0IIpOTeHHaMH, rJIHKOJIHilH)].aMH H.D:p y r HMH.

CB}13aBWHCb c JyKapHoTJNecKott KJieTKOM, cpparMeHT B cnoco6cTByeT OT)].eJieHHlO cpparMeHTa A H ero TpaHcnopTHp0BKe BttyTpb I<JieTKH qepe3

o6p83yeMbIH  Kattan  B KJleTO<{HOM  MeM6paHe HJIH nyreM 3H)0l UHT03a. Ha­

X0)].}lCb BH)'TpH KJieTKH, cpparMeHT A (aKTHBaTOp) nopIDKaeT (KaTaJIH3HpyeT) Te wrn: HHbie KJieT0<{Hbie MHIJieHH, HapyrnaeT)KH3HeHH0 BIDKHbie cpyHKUHH KITeTI<H " '18CTO npHB0JJ.HT ee K r n 6 eJIH.

CJlO:HCHble Ol!Jll<.06ble mOKCUHbl C0CT051T He MeHee qeM H3 JJ:BYX B3aHM0CB}138HHblX Il0JIHTieTITH)].HblX uenei-i - cpparMeHTa A H cpparMeH-

46

 

Ta B, H npe,II.cTaBJUIIOT co6oA roTOBy1:0 6mpyHK[(HOHaJTbHyJO AB-cHcTeMy.

<I>parMeHT B (mn, KOMTIJieKc, o6bI'lfHO COCTO.Slll(HH H3 5 cpparMeHTOB - B-neHTaMep) npHcoe,nHHlfBUIHCb K raHrJU103H)laM HJIH.npyrHM pe[(eTITOpaM qyncTBHTem,HoH 3yKapHOTHqecKott KJieTKH, pacm.enmJeT cpparMeHT A Ha

.nse cy61>e.!1,HHHllhl A I H A2. Cy61,e,n:HHH[(a A2 cnoco6crnyeT npoHHKHo­

Bemuo cy61,e,n:HHH[(hl A1 B KJieTicy, Ha MHIIIeHH KOTOpOH OHa OKa3bIBaeT TOKCH'-IeCKOe.netic TBHe.

TaKHM o6pa30M, MeXG1-IU3M 63aUMOOeucm6Ufl IlpOCTbIX H CJIO)l(HhlX TOKCHHOB C 3yKapHOTH"-leCKOH KJieTKOH BO MHOfOM cxo.n;eH H.sIBJI.sieTC.sI YHHBepcaJibHblM)lJI.sJ MHOnlX 6eJIKOBbIX TOKCHHOB.

CneljUq_JU'-IHOCmb oeucm6Ufl TOKCHHOB o6ycJIOBJieHa opraHoTpon­ HOCTbJO <pparMeHTa B, npHcoe,rurn.sirom.eroc.si K pa3JIH'-IHhTM TKaH.sIM Ma­ KpoopraHmMa, a TaIOKe cne[(HqlW-JHOCTI,fO <pepMeHTaTHBHoro.n:eiicTBH.sI cpparMeHTa A, aTa1cy10m.ero y pa3HbIX TOKCHHOB pa:moo6pa3Hhie KJieTOtIHble MHUJeHH

Cne,nyeT OTMeTHTh, ttTo KpoMe AB-ToKCHHOB, cym.ecTBYFOT TOKCHHhI C.n:pyrHM CTpoem1eM (cynepaHTHreHbI H HeKOTOpbie MeM6paHOTOKCHHbI).

 

 

K.rlaccmj,urca1-HH TOKCHHOB no MexaHHlMY,!leHCTBHB

EeJIKOBble TOKCHHhl o6na,naJOT pa3JIHT.fHbIM MexaHH3MOM,n:eHCTBH.sI, ITO 3TOMY Kpmepmo Hx no.n:pa3,n:en.sIIOT Ha 5 rpynIT (rn6n. 5):

TOKCHHhI, HHnT6HTOpbI CHHTe3a 6enKa - [(HTOTOKCHHhI;

TOKCHHhI, TIOBpe)l(,n:aJOllJ,He KJieToqffI,Ie MeM6paHbl - MeM6paHo­ TOKCHHhI;

TOKCHHhl, Hapyrna101.I.(He nepe,n:aqy CHrHanOB - cpyHKllHOHaJJbHbie 6JIOKaTOpbI;

TOKCHHI,I-ITpOTea3bI - cpyHK[(HOHaJibHble 6rroKaTOpbI; TOKCHHbI-CyITepaHTHreHbI.

 

To1<.CUHbt, u1uu6upy10w.ue cuHme:J oeJ11<.a. Ilo,n:aBJJ.sIJOT cHHTe3 6en1<a Ha pH6ocoMax,.n:ettcTBYH Ha pa3JIHT.fHhie 3Tam,1 3Toro npo[(ecca. HanpHMep,

.n;HcpTepHHHbIH U:HTOTOKCHH.sIBJIHeTC.sI aHTH3JIOHraTopoM. 3ToT npocTOH ITO

CTpoemuo AB-TOKCHH B3aHMo,n:eHCTByeT C 3TIHTeJIHOll}ffaMH HJIH.n:pyrHMH 3yKapHOTHt-leCKHMH KJleTKaMH B COOTBeTCTBHH C MexaHH3MOM, paccMOTpeH­

HbIM BbIIIIe. BKJieTey npoHHKaeT aKTHBHpOBaHHbIH cpparMeHT A, o6na.n:ruo-

1.I.(HH A,[{<l>-pH603HJITpattccpepa3HOH aKTHBHOCTblO, KOTOpbIH nepeHOCHT A,[{<I>-pu603y oT HA,Z:(+ na MHIIIeHI, - cpaKTop 3JIOHraUHH 2 (EF-2), 'ITO

47

 

npHBO,!:UIT K ero HHaKTHBaUIU{ H npeKpam:emno HapaW:HBaHH.SI nennl,IJ;HblX ueneif Ha pH6ocoMax, a B nanhHetHueM H K rw6eJIH KJieTKH. Tio TaKOMY

:>Ke npHHUHny neAcTByeT T0KCHH A CHHerHOHHOH narro<JKH (Pseudomonas aeruginosa).

.r(pyron npHMep - llfara-TOKCHH (STL), 06pa3yeMbIH HeK0TOpblMH

B036yAHTemrMH,IJ;H3eHTepHH (Shigella dysenteriae), a TaIOKe urnranono6- HI,IR TOKCHH, npo.nyQHpyeMbIH urnrenJJaMH H HeKOTOpbIMH,IJ;HapeereHHbIMH 3w epHXHHMH. O6a TOKCHHa - CJIO)l(Hbie H HMeIOT AB5-cTpyKzypy. lllHra­ TOKCHH o6nanaeT 6onee Bblpa)l(eHHbIMHTOKCli'-ICCKHMH CBOHCTBaMH (U.HTO­ TOKCH'ICCKHMH, 3HTCpOTOKCHtieCKHMH H HeHpOTOKCH<JeCKHMH) H qacTH<JHO OTJUNaeTCj(no aHTHreHHbIM CBOHCTBaM OT WHranO,!I06Horo TOKCHHa. O6a

TOKCHHa, T0<JHee HX cyfrhe,D.HHHUbI A 1, Bbl3bIBaIOT q>epMeHTaTHBHOe no­

Bpe:>KAeHHe 28S pH6ocoMaJJhHOH PHK (60S) KOJIOH0U.HTOB ToncToro KH­

wet.J:HHKa, npeKpam:ag q>yHKIJ.HOHHpOBaHHe pH60COM, KOTOpbie CTaHOBHTCH He cnoco6HbIMH B3aHMo,ne»cTsosaTh c qia1nopaMH 3JIOHrauim EF-1 H EF-2. B pe3yJihTaTe npeKpam:aeTcH cHHTe3 6enKa H KJieTKa norH6aeT.

 

To1<.cunb1, n0Bpe:m:oa10u,,ue KJ1emo'IHb1e., tt e.M.opan bt. HapyrneHHe uenoCTHOCTH KJieT0'IHblX MeM6paH 3y1<apHOTli'-ICCKHX KJieTOK npoHCXO,I{HT, 6om,wei1qacTblO, B pe3yJibTaTe qiopMHpOBaHHH nop HJIH cpepMeHTaTHBHOro

nmpomna, Bbl3b1BaeMoro pg,noM TOKCHHOB. 3TH TOKCHHbl 0TJIJNalOTCj(npyr

OT npyra no CB0eMy CTpoeHHIO H npHKpeTTJlj[IOTCj[ K peuemopaM KJICTOK. Tiopoo6pa3y10m:HM,nencTBHeM o6nanaioT neHKOU.HAHHhI H reMonu-

3HHbI. OHH cnoco6Hbl nospe)l(AaTb MeM6paHbl H mnuposaTb He TOJibKO

JICHKOUHTbl HJlH 3plnpoUHTbl, HO MHOfHe THnbl KJICTOK (MOHOUHTbl, TpOM60QHTbI, JIHMq>OUHTbl, 3H,!I.OTCJIH0UHTbl "np.). HeKOTOpbie  TOKCHHbl

,neHCTBYJOT TOJibK0 Ha KJleTI<H OAHOro THna, npyrHe - HaHCCK0JlbKO THflOB KJieTOK (KaK HanpHMep, a-T9KCHH Staphylococcus aureus).

,ll;pyron MexaHH3M noBpe:>KAeHIDI - HapyweHHe U.CJIOCTHOCTH MeM6paH c noMom:1:,10 q>epMeHTaTHBHoro rnnponH3a qiocq>onnntt,noB. TaKoe neifcTBHe oKa3bIBaeT a-TOKCIIB Clostridium perfringens, o6na,naJOW:HH cpoc¢onHna3- HOH aKTHBH0CTblO.

HapymeHHe np0HHUaeM0CTH H OCMOTH<JeCKOH uenocTH0CTH K.Jle­ TO"lfHbIX MeM6paH npHBOJJ.HT KJTCTKY K rn6enn, 061:,prno BCJTe,I{CTBHe UHTOJIH3a.

 

ToKCUHbl, napyma10u,,ue npou,ecc nepeoa11u cuznar,oB. 3TH TOKCHHbl He BbI3bIBaJOT npHMOro nospe:>KAeffHj(3yKapHOTH<JeCKHX KJICTOK, a JJ.CHCTBY-

48

 

IDT onocpe.n:oBaHH0, IT0CbUia.sJ ClffHaJlbl K pa3Hb1M q_)epMeHTaM, K0T0pble aKTHBHpYJOT MOJieKYJlbl nocpe,UHHK0B- BT0pWIHhlX MecceH,n:)KepoB (HanpH­ Mep, lll·U<JlHqecKHH a,n:eH03HHMOHOQ)OCq_)aT- u;AM<l>, HH03HTOJI-TpHcpocq>aT H,n:p.). MecceH,U)l(epbI - Me,D.I-taT0pbI, no.n:.uep)l(HBaIDmHe pa3Jil[qHI,Ie cpyHKD;HH KJieT0K. TaKHM o6pa30M, T0KCHHbl, B3aHMO,n:e»cTBy.sr C KJieTKOH,

3anycKalOT u;eJibTH KaCKa,n: peaKD;HH, npHBo,n:.srmHX K HapymeHHIO HJIH HC­ KIDKeHHIO KJieToqHI,IX Q)YHKUHH' qTo, B K0HeqHoM HT0re, M0)l(eT npHBeCTH

K pa3BIITHID naTonorWiecKHX npoueccoB Ha ypoBHe oprattH3Ma.

Mttorne T0KCHHbl 3TOH rpynnbI Bbl3bIBruoT ycroiiqHBYJO aKTHBaumo a.n:eH1-rnan m<Jia3HOH CHCTeMbl, H, KaK CJie.[(CTBHe, HapymeHHe B0.D:H0- 3Jiel<Tp0JIHTH0ro o6Mett a. B pe3yJibTaTe B03HHKaeT ceKpeT0pHa.sr,n:Hape51 (npH xonepe, KOJIH-HH<peKD;HH), OTeKH TKaHeH (npH CH6HpCKOH 513Be), H36blTOqHa.sr

CeKpeUH51 B5f3KOH MOKp0ThI (npH KOKJIIOUie) H T.n.

K TaKHM T0KCHHaM 0TH0CHTCH TepMona6HJibHbIH T0KCHH Vibrio

cholerae, tta3bIBaeMbIM xoJ1epo2e11.oM. 3To   cJIO)l(HbIH ABS-ToKCHH, co­ CT051IIJ.HH H3 cy6'be,n:1rnHD;bl A, OKpy)l(eHHOM K0MITJieKC0M H3 5 cy6'be.D:HHHU

B B BH,n:e Kon1,ua. IleHTOMep B npHcoe,uHH51eTC51 K rattrnH03H.[(y Gm 1 Ha Il0BepXH0CTH 3HTepou;HTa, 06pa3y.s: KaHaJI, no K0TopoMy cy6'be,UHHIU a Al npoHHKaeT B KJieTKy. AKTHBaTOp Al rH,n:ponH3YeT KJieToqHbIH HA,[(

.n:o HHK0THHaMH,n:a H A,[(<I>-pH603bl H pH60JIH3HpyeT G-6eJIOK (ryaHH3HH­

TpHcpoccpaTCB5f3bIBaIOmHH 6enoK). IloBpe)l(Jl:ettue G-6enKa npHBo,n:HT K 6noKa,n:e peryJIHUHH aKTHBH0CTH a.n:eHHJiaTUHKJla3bl, K0T0paH yqacTByeT s npespaur.ettHH A,[(<I> B uAM<t>. YcToj:j:q1ma51 aKTHBaUHH a,n:eHHJiaTUHKJia3bI Bbl3bIBaeT 3HaqHTeJibH0e HaK0JIJieHHe uAM<I>. 3To HapymaeT TpaHCil0pT 3JieKTp0JIHT0B B TOHK0M KHrueqm1Ke H, B qacTH0CTH, B 3HTepoUHTaX, B K0-

3

T0pblX IT0BbIUiaeTCH CeKpeUIDI H0H0B xnopa, HaTpIDI, Kap6oHaTOB (Na+, Cl·, CQ-) H Il0.[(aBJIHeTCH Il0CTYTIJieHHe B KJieTKY H0HOB KaJIHH. OCMOTJflleCI<Oe

,n:aBJieHHe B Il0JI0CTH TOHKOH KHIIIKH CTaH0BHTCH BblIIIe, qeM B KJieTKax, H3

HHX-CeKp eTHpy eTCH Bo,n:a. Pa3BHBaeTCH xapaKTepHM wrn xonepbl npocpY3HM ·

.n:uapeH C qacTblM H o6HJlbHblM cryJI0M B BH,n:e «pHC0B0ro 0TBapa», 60JibHOH M0)l(eT norH6ttyTh OT o6e3BO)l(HBaHH51.

Cxo.L{HbIM 110 cTpoem110 H MexaHH3MY.n:encTBH51 HBJIHeTc51 TepMo­ na6HJibHbIH 3HTep0T0KCHH (LT) 3HTepoT0KCHreHHblX E.coli, o6na,n:aIOIIJ.HH

MeHblllei-i TOKcHreHH0CThIO, qeM xoneporeH. TepMocTa6HJibHbIH 3HTeporoK­ CHH (ST) E.coli 110 TaKOM)')Ke THny H C TeM)Ke pe3yJibTaT0M B03)leHcTByeT Ha ryaHHJTaTu;HKJia3HYJO CHCTeM)' 3TIHTeJIHOUHTOB, 11pH 3T0M HaKailJIHBaeTCR

UHKJIHqecKHH ryaHH3HHMOHOcpoc<paT - urM<I>.

 

 

49

 

1tJICCllHbl - 11pometnbl. 8 31)' rpynny T0KCHH0B BX0,!lRT HetipOTOKCH­ Hl.1-TeTaHocna'lMHH, npo.!lym-1pyeM1.1tt Clostridium tetani H 6orynOTOKCHHI.I p8.3J111'-1HblX THn o s, 06pa3yeMb1e Clostridium botulinum. O6a T0KCHHa RBJl.ll­

lOTCR U,HHK0BblMH 3H.11.0npoTea3aMH. 3To caMbte CHJlbHble 6aKTepmU1bHble TOKCHHbl. 1-13 HHX Ha nepB0M MeCTe - 601yJ10TOKCHHbl, JleTaJtbHaR)l03a K0TOpblX.llJlR tfeJI0BeKa C0CTaBJUleT I 0·8 r.

Tem a 11ocna 3_.wu,1.   Ott HMeeT  ocH0BHoe  1Ha4eHHe  s  naTorette1e

CTOJ16HRKa, HaKanJlHBaRCb e paHe, HH$HUHpOBaHHOH cnopaMH C.tetani. KOT0pble npopacTatOT B serenrrHBHble KJleTKH, npo.11.yu,HpytotUHe 4aCTH4H0 ceKpempyeMblH npoCTOH T0KCHH. Ero noJrnnenTH.D,Ha.R uenb (npoTOKCHH) npeo6pa3yeTcR s 61-1cpyHKU.HOHaJtbHyto A B - c HcTeMy n)"TeM npoTeonHTH­ ttecKoro pacmennettn.11 Ha nerK)llO uenb (cpparMeHT A) H TR)Kenyto u,enb

(<l>parMeHT B), coe,mmeHHble,L1Hcy11bqm.11.t-roit cBR3bto 11 TeTaHocna1MHH

npno6peTaeT CBOHCTBa HeHp0TOKCHHa.

<t>parMeHT B CB.113bIBaeTCH C MeM6paHOH nepmpepw-iecKoro M0T0- ttenp0Ha H cnoco6cTByeT np0HHKH0BeHHIO B KJleTK)' nyTeM 3H.D,0U.HT03a

<l>parMeHTa A, KOTOpblH c noM0lllbto BHyTpHaKc0HHoro pe-rporpa.n.ttoro

TpaHCn0PTa nocrynaeT B T0pM03Hble BCTaB0l.fHble HeHpOHbl cnHHH0f0 M03- l a. 3nt HeHpOHbl peryirnpytoT.11.e11TeJlbH0CTb)lBHfaTeJlbHblX MOTOHeHpOHOB nocpe,nCTB0M 6eJ1Ka CHHanTo6peBHHa, 0TBeTCTBeHH0f0 3a <t>YHKU.H0HHpO­

baHHc CHHaTIC0b.:.:)roT 6enoK Heo6xo,nHM.llJlR coe.nHHeHltH CHHanTHtteCKHX llY'.3blpbK0B C 11pt:CHHatlTH'it:CKHMH Mt:MUpaHaMH H BblX0.Ua T0pM03HblX HdipoMe,llHaTopos - y-aMHH0MaCJIHH0H KHCJl0Tbl H rJIHU,HHa. <I>parMeHT A,

o6na,n,aIOtUHH npoTea3HOH aKTHBH0CTbto, pa3pywaer CHHanTo6peBHH H, TeM CBMLIM, 6JIOKHpyeT CHCTeMy T0pM0)KeHH.sr, t.jT0 npHB0)lHT K noCT0RHH0M)'

nocrynneHHIO HMnyJlbC0B Ha cnHHH0M03r0BblC.llBHraTCJlbHblC HCHp0Hbl, libl:ShlliaH .I-{.'( Ht:pt:Hu 6y)K)leHHe, H 06yc11aBJ1HBaeT.llJlHTeJibH0e cnaCTHYe­ CII.0C: co11.-pamemte MblUIU - cnaCTHl.feCKHe napaJJHl.fH. K0HTpaKrypbl, a TalOKe

TeTaHHYeCKHe cy.11.oporn. "I lepBblMl-1 803HHK8IOT T0HH'ieCKHe  C0KpameHHR

:>KeBaTCJlbHblX H MHMHt.jCCKHX Mbl WU, '.3UTCM cna'.3M MblWU '.3UTblJIKa, Cnl·IHbl,

Mt:)Kpclit!pHblX MblWU. CMt:prb MWKt:r Hac1y1n11b UT ac4J11KCl1H H 11upi0Kt:HH>I

.1Kl13HettH0 BWKHblX ueH fp0B;

b'OTt l)!llOfn O KCU/l bl. '.:h'H T0KCHHbl nona.nat0T B nHI.J..{eBapHTCJlbHblH TpaKT C nHuteBblMH npo.nyKTaMH, HHqmuHpoaaHHblMH cnopaMH C.botulinum. 4ame BCero 3T0 He)],0CTaT0YH0 npocTep1-1n1-130BaHHble.noMaWHHe K0Hcep­

l:lbl, 1:1 11.or-opblX C03.[18JlHCb 611aronpmlTHble (aHa3p06Hbte) ycnoBHR.nm, HaT<onncHHa T0T<CHH0B. CymecTsyeT ceMb THnos oorynoToKCHHOB -A,  B,

 

50

 

C, 0, I::, F, U, pa1JJH'-IaJOUti-1xcsi no aHTHreHHbtM csoikrnaM, HeKOTOpbte H3 HHX HMelOT OTJIH<msi B MexaHH3Me.ri;eHCTBHSI.

DO"fYJlOTOKCHH nocTpoeH no THTIY npocToro TOKCHHa " HaxO,DJ-lTCSI B KOl\mJJeKce C HeTOKCWJeCKHMH 6eJJKOBblMH KOMHOHeHTaMH, KOTOpble 3al.I{H­

UlaJOT ero OT,lleHCTBHSI nHII{eBapwreJTbHbfX cpepMeHTOB _J,f COJTSIHOH KHCJlOTbl.

vb sepXHHX OT,[(eJIOB)KKT TOKCHH npOHHJ{aei B JIHMq><tTH'-leCl<yIO cHcTeMy H 1aTeMB Kpos.srnoe pycno, MO)KeT npeo.nonesaTb reMaT0-3H'QecpanWiecKHH

6apbep. nocJJe npoTeom-tTINecI<Oro pacmenneHHSI npoTOKCHH npuo6perneT

CBOHCTBa HeHpOTOKCHHa. TSl)KeJJasr nommernH,ll.HaSI a;enb TOKCHHa- q>par­ MeHT B - cBK3bIBaeTcsi co cneumf>wtteCKHMH pea;emopaMn npec1manTH­

ttec1<HX MeM6paH MOTOHeHpOHOB (raHrJJH03H.ll.HOH HJIH.npyrou npHpO.ll.bl). Jlernasi llenh - <pparMeHT A - TpaHCTIOpTHpyeTCH BnyTpb KJleTKH nyTeM onocpe.nosaHHOrO peu:enTopaMH 3H,ll.OQHT03a. BttyTpH MOTOHenpoHa A-ah.,HBaTOp,.nettCTBY" KaK 3H,[(onpoTea3a, pa1pyrnaeT 6eJIKH-MHlileHH,

•no npuso.nHT 1< unrn6uposamno ceKpeuJrn: au:eTHJJxomrna H npephIBaHHJO HepBHblX HMnyJTbCOB, HJlYUlHX OT MOTOHeHpOHOB K MblllIQaM. B pe3yJibTaTe

6JTOKa,nbl MOTOHeHpOHOB npe1<pamaJOTCSI MblIIJet{Hhie COKpameHHH, pa3- BHBalOTCSI BSIJlbie napaJIWIJf.

DO"fYJJOTOKCHH MO)KeT nopa)J(aTh MOTOHeHpOHhI cnHHa..'lbHbJX MOTOp­

HblX uenTpOB, Kllep npo.uonrosaToro M03ra H nepH<}}epHt.leCKOH HepBHOH CHCTeMbl. qame Bcero nopa)KaJOTC1l Mhlllie<fHbJe TKaHH, HHHepsnpyeMbJe

.!1,1.lepHblMH ttepBHblMH ueHTpaMH. Hacry11aeT paccTpOHCTBO aKKOMO,l],aQIUI, nopaiKeHHe rna3HhlX MblIIJQ,,llBOeHHe B rna3ax, 3aTpy.llHeHHOe ID,IXaHHe, acpoHHll'. TTpn Tll)l{eJJhlX cpopMax 6oryJJmMa CMepn, ttacrynaeT OT napanlflfa

,llhIXaHHll, HHor.na OT BHe3anHOM OCTaHOBKH cepm a.

 

T01<e1utbt - cynepa11muienb1, iJeilc1111Jy10114ue na UMMynny10 cucnteMJt

B ·3ry rpynny BXO,USIT 6eJJKOBbJe TOKCHHbl, cnoco6Hbie Hecneu:mfmtte­ CKH aKTHBHpOBaTh MHOiKeCTBO KJIOHOB T-JIHM<}}ou:wroa, T. e. Bbl3hJBaThHX

OOJIHl<JlOH8JlbH)'fO 8KTHB8QHfO, CTHM)'JIHpoBaTb MaccOBytO npom1cf>epaa;mo 11 1·1111t:p11po,L{)'Klll1IO Mt:.U,HaTopoa aoc11a.11eHI1>t H HMM}'}UIOl'l) OTBeTa - UH­ TOKHHOB. Pa3BHBaeTC,t HHTOKCHKaUH1l, onocpenosaHHaH UITTOKHHaMH H,

oco6etttto, HHTepneHKHHOM 2 (HJl-2 u HJI1), KOTopasi nposreJJsieTcK a BM.Ile

ITOBb!WeHHSI TeMneparypb!, CHHiKeHHSI apTepHaJJbHOro,ll.aBJJeHIDI (A.[() H

.npyntX CHMTITOMOB.)J;anee ttacrynaeT rH6eJTh aKTHBHpoaaHHhlX KJJOHOB T-JJHM<pOUHTOB, 803MOiKHO pa3BHTHe T-KJTeTOt{HOro HMMyno.necf>HUHTa.

 

51

 

CynepaHTHreH CB513bIBaeTC51 C peu:enTOpOMH rnaBHOro KOMTIJieKca rncTocoaMecTHMOCTH TI KJiacca (MHC TI) Ha noaepxttoCTH MaKpoq>aroB H

,IleH,IlpHTHhIX KJieToK H c TCR (T-KJieTO'-IHhIH peu:enTOp) T-JIHMq>au:moB. Ho cynepaHTHreH npo11coe,IlHHSieTc»: K 3THM peu;enTopaM, «Henpa­

BHJibHO» - BHe HX Af-pacn03HaJOIUHX H Af-CB513bIBaJOIUHX yqacTKOB.

Tio3TOMY cneu;H<j)Ht.IecKHHHMMYHHLIH OTBeT He pa3BHBaeTc»:, a rrpowcxo,IlHT IlOJIHKJIOHaJlbHa»: aICTHBaU:H»: T-KJieTOK (pHC. 1).

EaI<TepHaJibHble TOKCHHbJ-cyrrepru-rrnreHbJ B OCHOBHOM CIDITe3HpYJOTC51 CTaqlHJIOKOKKaMHH CTPe ITTOKOK.KaMH. S.aureus o6pa3yeT 3HTepoTOKCHHbl, 3KC­ qlOJIHailIBHbleTOKCHHbl H TOKCHH, Bbl3bJBalOIUHH CHH,IlpOM TOKCH'-leCKOro lIIOKa

(TTIII); S.py ogen es - 3pJ.fTPOreHHbie TOKCHHLI, Bbl3bIBaJOI.D;He 3K3aHTeMY.

PHCYHOK l

CxeMa a1<muBaliuu T-llUMq>ou,umoB aHmuzeHoM u cynepaHmuzeHoM

AKTIIBA[(JUI AHTllfEHOM	AKTHBA[(JUI CYTIEPAHTHfEHOM
MHC II Kll. TCR •lr' AKTHBauHll aHTHreH-cneuvqrnqecKoro KJIOHa T-mfM(pOI.IHTOB Pa3BHTHe a,nanTHBHoro HMMYHHTeTa	AKTHBallHll MHOrHX KJIOHOB T-JJHMq>Ol.{HTOB Tipo.nyKl.{Hll H ceKpel.{Hll UHTOKHHOB Pa3BJITHe cHHnpoMa 06111eli RHTOKCHKal.{HH fH6em, aKTHBHpOBaHHhJX T-JJHM(j:,OUHTOB qepe3 anonTOJ BoJMOll<HOe pa3BHTHe RMMYffOne(jlHUHTa

llpUMe'laHue:

ATIK - aHTHreHnpeJeHTHpy10mall KJJeTKa

MHC II 1rnacca • MOJJCKYJlbl (aHTHreHLI) IT KJJacca rnaBHOro KOMJJeKca rnCTOCOBMeCTHMOCTH

Ar- aHTHreH

CAr - cynepaHrnreH

TCR -T-cneuH(jlH'!ecKHR peQenTOp

 

52