Molecular characterisation of the recombinant Vesicular Stomatitis Virus- ZEBOV-GP virus, prototype vaccine against Ebola virus
Texte intégral
(2) N° d’ordre NNT : 2019LYSE1009. . THÈSE de DOCTORAT DE L'UNIVERSITÉ DE LYON opérée au sein de. l’Université Claude Bernard Lyon 1 . Ecole Doctorale N° 340 Biologie Moléculaire Intégrative et Cellulaire . Spécialité de doctorat : Virologie Discipline : Biologie moléculaire Soutenue publiquement le 01/02/2019, par : Nicolas DANET . Molecular characterisation of the recombinant Vesicular Stomatitis VirusZEBOV-GP virus, prototype vaccine against Ebola virus Devant le jury composé de : . Jamin Marc, Professeur, Institut de Biologie Structurale - CEA Grenoble, Rapporteur et Président du jury. Blondel Danielle, Directeur de Recherche CNRS, Institut de Biologie Intégrative de la Cellule Université Paris-Saclay, Rapporteuse. Gonzales-Dunia Daniel, Directeur de Recherche CNRS, Centre de Physiopathologie Toulouse-Purpan - Université de Toulouse, Examinateur.. . Manet Evelyne, Directrice de Recherche CNRS, Centre International de Recherche en Infectiologie - Université Claude Bernard Lyon 1, Examinatrice. . . Gilbert Christophe, Maître de Conférences UCBL, Centre International de Recherche en Infectiologie - Université Claude Bernard Lyon 1, Examinateur. . Volchkov Viktor, Professeur UCBL, Centre International de Recherche en Infectiologie, Directeur de Thèse. . . . . g.
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(30) • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •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kLFH=B?B>=!:@E>k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n.
(31) • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •. . #,<EE>:O>=#, #/#>G>/M:KM #/#>G>LM:KM $//$:GDLk;:E:G<>=L:EMLHENMBHG $>2$>G=K:OBKNL $BL$BLMB=BG>M:@ $%2$NF:GBFFNGH=>?B<B>G<ROBKNL $'$B@AFHE><NE:KP>B@AMDBGBGH@>G $( .$NF:GE>NDH<RM>:GMB@>G .BLHMRI> $,%2$NF:GI:K:BG?EN>GS:OBKNLMRI> $. $>IM:=>K>@BHGn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o.
(32) • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •. . )%, o q):<KHIA:@>BG?E:FF:MHKRIKHM>BGo q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c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gf.
(33) • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • •. . K)(K><HF;BG:GM)( .*.B;HGN<E>HMB=><B= K*,<.><HF;BG:GMG>F:MH=>:GMB<H:@NE:GMIKHM>BG< .*,.B;HGN<E>H<:ILB= K.*.B;HLHF:E.* .0.>MKHMK:GL<KBIMBHG K2/2 (/2#,K2/2 (:LL:OBKNL@ER<HIKHM>BG K2/2 #, $/K2/2 #,$B@ALA>=FNM:GM K2/2 #, (/K2/2 #,(HPLA>=FNM:GM K2/2 6!+2K><HF;BG:GM2/2 6!+2 /><</!AHFHEH@ L#,/HEN;E>#, L%( ./HEN;E>BGM>KE>NDBG>K><>IMHK LB.*LF:EEBGM>K?>KBG@.* /*,/BG@E>*N<E>HMB=>,HERFHKIABLF /,*0/NI>KG:M:GM LL#,/NI>KLF:EE#, L0*" ./HEN;E>0NFHKG><KHLBL?:<MHKK><>IMHK /1 2/N=:G>;HE:OBKNL 0!0*"o <HGO>KMBG@>GSRF> 0"20:Y"HK>LM>;HE:OBKNL 0% 0BLLN><NEMNK>BG?><MBO>=HL> 0"0BLLN>":<MHK 0@L0NFHKLNL<>IMB;BEBMR@>G> 0%),0BLLN>BGAB;BMHKH?F>M:EEHIKHM>BG:L> 0(.0HEE EBD>K><>IMHK 0)0K:GLF>F;K:G>=HF:BG 0*"o0NFHKG><KHLBL?:<MHKo 0.%(0*" K>E:M>=:IHIMHLBL BG=N<BG@EB@:G= 0.,0AKHF;BGK><>IMHK:<MBO:MHKI>IMB=> 11EMK:<>GMKB?N@>=LNI>KG:M:GM b@)B<KH@K:FL 10.1GMK:GLE:M>=K>@BHGL 2) 2:L<NE:K<>EE:=A>LBHGIKHM>BG 2!!22>G>SN>E:G>JNBG>>G<>IA:EBMBLOBKNL 2>KH!2>K=:K>GH! 2(,2BKNL EBD>I:KMB<E> 2,2BK:EIKHM>BG 2,*,,2,*,BG=BG@,>IMB=> O,),2BK:EI:MAH@>G:LLH<B:M>=FHE><NE:KI:MM>KG 2.,2>G>SN>E:G>JNBG>>G<>IA:EBMBLOBKNLK>IEB<HG 2/22>LB<NE:KLMHF:MBMBLOBKNL 3:LA>=,(03:LA>=IE:M>E>ML 303BE= MRI> 33,33=H:BG<HGM:BGBG@!N;BJNBMBGIKHM>BGEB@:L> 50RKHLBG> oEIA: q>M: p >EM: gg.
(34) !;HE:OBKNL!+2>LMNG?BEHOBKNLK>LIHGL:;E>=>?BWOK>LAVFHKK:@BJN>LOBK:E>LLVOWK>L<A>S EkANF:BG JNB I>NO>GM XMK> EVM:E>L =:GL =>L <:L
(35) (k:<MN>EE> VIB=VFB> >G .VIN;EBJN> VFH<K:MBJN>=NHG@H>MEk:FIE>NK=VF>LNKV>=>EVIB=VFB>=> >G?KBJN>=> Ek+N>LM JNB:<:NLVE:FHKM=>IENL=>I>KLHGG>L HGMIHNLLVE>L:@>G<>LL:GBM:BK>L BGM>KG:MBHG:E>L U M>LM>K IENLB>NKL :IIKH<A>L MAVK:I>NMBJN>L :?BG =k>LL:R>K =k>G=B@N>K K:IB=>F>GME:IKHI:@:MBHGOBK:E>>M=>EBFBM>KE:FHKM:EBMVEBV>:NOBKNLEHKL=>?NMNK>LVIB=VFB>L
(36) ,:KFBMHNM>LE>LLMK:MV@B>LM>LMV>L E>OBKNLK><HF;BG:GMKVIEB<:MB?K2/2 6!+2JNB>QIKBF> E:@ER<HIKHMVBG>=>LNK?:<>=k!+2 L>F;E>H??KBKE:F>BEE>NK>IKHM><MBHG :NLLB;B>G>GFH=WE> :GBF:EB>KL JN> LNK E> M>KK:BG
(37) O:GM =kXMK> M>LMV <A>S EkANF:BG => GHF;K>NL>L VMN=>L HGM I>KFBL => F>MMK> >G VOB=>G<> Ek>??B<:<BMV >M EkBGGH<NBMV => <> O:<<BG IKHMHMRI>
(38) ,HNKM:GM >M F:E@KV E> ?:BM JN> => GHF;K>NL>L VMN=>L HGM =VFHGMKV EkBFIHKM:G<> >M E> KZE> => E: @ER<HIKHMVBG>#,=:GLEk>??B<:<BMV=>LO:<<BGL<HGMK><>OBKNL :N<NG>VMN=>Gk:>G<HK>VMV KV:EBLVLNKE:G:MNK>=>L@ER<HIKHMVBG>LOBK:E>LLRGMAVMBLV>LI:KE>@WG>#,=k!+2BGLVKV =:GLE>@VGHF>=NOBKNL2/2
(39) BGLB E>L<:K:<MVKBL:MBHGLFHEV<NE:BK>L=>LIKHMVBG>LOBK:E>L IKH=NBM>LEHKL=>EkBG?><MBHGI:KE>OBKNLK2/2 #,=V<KBM>L=:GL<>LMK:O:NQ=>MAWL>H??K>GM=> GHNO>EE>LI>KLI><MBO>LIHNK<HFIK>G=K>E>LN<<WL=><>O:<<BGF:BL:NLLBEkHKB@BG>OBK:E>L =:GL E>L >??>ML L><HG=:BK>L LVOWK>L H;L>KOVL EHKL => E: O:<<BG:MBHG >M IHNKKHGM :B=>K U =VO>EHII>K NG O:<<BG IENL L[K JNB Gk>LM :<MN>EE>F>GM I:L NMBEBL:;E> <A>S E>L I>KLHGG>L BFFNGH=VIKBFV>L
(40) )HML<E>?L!($'% K2/2 6!+2 O:<<BG @ER<HLRE:MBHGL )"&'% $'$($'%
(41) . . . gh.
(42) 0A>?BEHOBKNL!;HE:OBKNL!+2BLMA><:NL:MBO>:@>GMH?L>O>K>OBK:EA:>FHKKA:@B<?>O>KLBG ANF:GLMA:M<:G;>E>MA:EBGH?<:L>L
(43) 0A><NKK>GMHNM;K>:DBGMA> >FH<K:MB<.>IN;EB< H?HG@H:G=MA>>QMK:HK=BG:KRL<:E>H?MA> HNM;K>:DBG3>LM?KB<: MA:M<:NL>= MA>=>:MAH?FHK>MA:G=BL>:L>OB<MBFL E>:=MA>BGM>KG:MBHG:EIN;EB<A>:EMA:@>G<B>LMH M>LML>O>K:EMA>K:I>NMB<:IIKH:<A>LMHEBFBMOBK:ELIK>:=BG@:G=FHKM:EBMR
(44) FHG@LMMAHL> MA> K><HF;BG:GM K>IEB<:MBHG <HFI>M>GM K2/2 6!+2 OBKNL MA:M >QIK>LL>= !+2 #, @ER<HIKHM>BG :II>:KLMHH??>KMA>;>LMIKHM><MBHGBG:GBF:EFH=>EL:G=HNM;K>:DL>MMBG@L
(45) 3ABE> BML >??><MBO>G>LL :G= L:?>MR A:O> ;>>G PB=>ER BGO>LMB@:M>= ;>?HK> ANF:G MKB:EL :G= =>LIBM> GNF>KHNL LMN=B>L MA:M LAHP>= MA> BFIHKM:G<> MA> G:MNK> H? MA> @ER<HIKHM>BGL PAB<A :K> IKH=N<>==NKBG@MA>BG?><MBHG?KHFMA>!+2#,@>G>MA:MA:L;>>GBGL>KM>=BG2/2@>GHF> :K> NGDGHPG
(46) %G MABL K>LI><M MA> FHE><NE:K <A:K:<M>KBL:MBHG H? MA> OBK:E @ER<HIKHM>BGL LRGMA>LBL>= =NKBG@ K2/2 #, IK>L>GM>= BG MABL MA>LBL H??>K G>P BGLB@AML PBMA PAB<A MH NG=>KLM:G= MA> LN<<>LL H? MA> K2/2 #, O:<<BG> ;NM :ELH MA> IHM>GMB:E OBK:E HKB@BGL H? MA> L>O>K>:=O>KL>LB=>>??><MLH;L>KO>==NKBG@O:<<BG:MBHG:G=<HNE=A>EIBG=>O>EHIBG@:L:?>K O:<<BG> PAB<A<NKK>GMER<:GGHM;>NL>=BG:GBFFNGH<HFIKHFBL>=IHINE:MBHG
(47) '>RPHK=L!($'% K2/2 6!+2 O:<<BG> @ER<HLRE:MBHGL )"&'% $'$($'%. . gi.
(48) L J
(49) !;HE:OBKNLBL:G>GO>EHI>= GHG L>@F>GM>=LBG@E> LMK:G=>=G>@:MBO> .*OBKNLMA:MP:L?BKLM BLHE:M>=BG?HEEHPBG@HNM;K>:DLBG/HNMA/N=:G:G=6:BK>GHPMA> >FH<K:MB<.>IN;EB< H?HG@H78
(50) %MBL:SHHGHMB<OBKNL:G=MA><:NL:MBO>:@>GMH?!;HE:OBKNL=BL>:L>!2 : L>O>K>=BL>:L>BGANF:GL:G=GHG ANF:GIKBF:M>L*$,PBMA:?:M:EBMRK:M>MA:M<:GK>:<A BG LHF> HNM;K>:DL
(51) L MA>K> :K> <NKK>GMER GH O:<<BG> HK MA>K:I>NMB< :IIKH:<A>L :IIKHO>= !;HE:OBKNLBL:F:CHKIN;EB<A>:EMA<HG<>KG
(52) !;HE:OBKNLBLI:KMH?MA>!($'%@>G>K: PAB<A:EHG@LB=> $'$($'%:G='(($'% <HFIHL>MA>"BEHOBKB=:>?:FBER"B@
(53)
(54) 0A>R:K><NKK>GMER?BO>K><H@GBL>=@>G>MB<:EER=BLMBG<M LI><B>L H? !;HE:OBKNL $ !($'% !+2 ' !($'% /1 2 * !$%& !($'% 0"2 ' ')! !($'% 1 2 :G= %&! !($'% .!/02
(55) ):K;NK@OBKNL BL <HFIHL>= H? : LBG@E> LI><B>L $'$ $'$($'% PBMA MPH F>F;>KL ):K;NK@ OBKNL ).2 :G= .:OG OBKNL .22
(56) "BG:EER =BL<HO>K>= BG N>O:OBKNL BL ?HKF>=H?:NGBJN>LI><B>L
(57) !(''(($'%((+27 8
(58) . =5
(59)
(60) 5 0%*! ".+) Zo[M $5(+#!*!0% 0.!! .!,.!/!*0%*# 0$! 0$.!! )&+. %(+2%.1/ #!*!.L +(2%.1/J .1.#2%.1/ * 1!22%.1/M +(2%.1/ %/ +),+/! +" "%2! /,!%!/L %.! !+(2%.1/ XYJ 1 * !+(2%.1/XYJ%+.!/0!+(2%.1/XYJ1* %1*#5+!+(2%.1/XY* !/0+*!+(2%.1/ XYM.1.#).1.#2%.1/XY* 2*2%.1/XY.!"+.)%*#0$!.1.#2%.1/#!*1/* (+2%12%.1/J0$!1!22%.1/#!*1/M$!/(!..!,.!/!*0/!2+(10%+*.5 %/0*!!03!!*!$*+ !M . . gj.
(61) /BG<>MA>?BKLM?BEHOBKNLHNM;K>:DBG <:NL>=;R).2 ?HKMR=B??>K>GMHNM;K>:DLA:O>;>>G K><HK=>= PBMA MA>BK F:CHKBMR ;>BG@ <:NL>= ;R HG> H? MA> ?BO> LI><B>L H? !;HE:OBKNL
(62) /H ?:K ((+2A:LGHM<:NL>=:GRHNM;K>:DBGANF:GL;NMP:L=>M><M>=HGERBG=>:=BGL><MBOHKHNL ;:ML 0:;E>
(63) 0A> F:CHKBMR H? MA> HNM;K>:DL H<<NKK>= BG >GMK:E :G= /HNMA ?KB<: PAB<A LN@@>LML MA:M ?BEHOBKNL>L :K> >G=>FB< H? MA>L> K>@BHGL
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(65) 7f8
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