8
INVES(IGA(IONS)R ARJG) A(ION(RAD)C(IONNE EPENDAN( A RJPONSEIMM)NI(AIREVJGJ(A EIND)I(EPAR ESPRO(JINESNB- RR
p0A
oD8B-V0;4nC8n MNC486n84A
C7OB4 pANB4nCN4 0D DNp0AC4m4nC 34 18o;o684 4n ED4 34 ;Xo1C4nC8on 3D 6A034 34 3o2C4DA OB B284n24B (P7.D.)
FAC) (E DES SCIENCES
)NIVERSI(E DE SHERBROOKE
88
4 23 FNE84A 2015
le jury a accepté la thèse de M nsieur L uis-Valentin Meteignier dans sa versi n finale.
M4m1A4B 3D 9DAH
PAo54BB4DA P4C4A Mo554CC D8A42C4DA 34 A4274A274 DNp0AC4m4nC 34 18o;o684
PAo54BB4DA J40n-B4no8C C70AAon JE0;D0C4DA 4GC4An4 )n8E4AB8CN M2G8;; PAo54BB4DA SN10BC84n Ro3A86D4 JE0;D0C4DA 8nC4An4 DNp0AC4m4nC 34 18o;o684 PAo54BB4DA D2 G0D3A40D PANB834nC-A0ppoAC4DA DNp0AC4m4nC 34 18o;o684
888 SOMMAIRE
X8mmDn8CN EN6NC0;4 4BC 60A0nC84 p0A p;DB84DAB n8E40DG 3X02C8on 3onC ;X8nC4A2onn4G8on 4C ;4B Eo84B B86n0;NC8qD4B BonC p4D N;D283N4B. D4DG 34B CAo8B 2oD274B 34 3N54nB4B 8mmDn8C08A4B BonC 2onBC8CDN4B p0A ;4B pAoCN8n4B NB- RR (ND2;4oC834-B8n38n6 4D28n4-R827 R4p40C), 4n2o3N4B p0A ;4B 6On4B 34 RNB8BC0n24 (6On4B R), 4C ;X8nC4A5NA4n24 K ARN (8ARN). XNCD34 34 ;0 Eo84 34 B86n0;8B0C8on 8n3D8C4 p0A ;X02C8E0C8on 34B NB- RR 4n ANponB4 K ;0 A42onn08BB0n24 (38A42C4 oD 8n38A42C4) 34 pAoCN8n4B p0C7o6Nn8qD4B 4BC CAOB 2omp;8qDN4, 20A 24CC4 ANponB4 2D;m8n4, 30nB ;0 m094DA4 p0AC84 34B 20B, 4n Dn p7NnoCHp4 m02AoB2op8qD4 34 moAC 24;;D;08A4 3NnommN4 RNponB4 7Hp4AB4nB81;4 (HR), 4mpP270nC CoDC4 0n0;HB4 18o278m8qD4. RN24mm4nC, 8; 0 NCN monCAN 274I
Nic tiana benthamiana qD4 ;0 ANponB4 0nC8E8A0;4 3N2;4n27N4 p0A ;0 pAoCN8n4 NB- RR
N, qD8 nX8n3D8C p0B 34 HR ;oABqDX02C8EN4, 8mp;8qD4 ;0 ANpA4BB8on 34 ;0 CA03D2C8on 34 ;XARN E8A0;. D0nB 24 BHBCOm4, ;0 ANB8BC0n24 2on5NAN4 p0A N 4BC 3Np4n30nC4 34 ;0 pAoCN8n4 AGO4 3onC ;4 AR;4 30nB ;X8ARN 4BC 0BB4I 184n 3N58n8. En 4554C, 184n qD4 ;XARN E8A0; B4 mD;C8p;84 K ;X8nCNA84DA 34 ;0 24;;D;4 7RC4, 8; nX4BC p0B 0BBo28N K ;0 m0278n4A84 34 CA03D2C8on 4C n4 pAo3D8C 3on2 p;DB 34 E8A8onB. C4p4n30nC, noDB n4 2onn08BBonB p0B K 24 9oDA ;4B NEOn4m4nCB 34 AN6D;0C8on 34 ;X4GpA4BB8on 6Nn8qD4 A4BponB01;4B 34 24 mN20n8Bm4 34 3N54nB4. 4B NCD34B 3D CA0nB2A8pCom4 8mmDn8C08A4 274I Arabid psis
thaliana nXonC ANEN;N qD4 p4D 34 20n3830CB m094DAB 8mp;8qDNB 30nB ;0 ANponB4
RR, 0;oAB qD4 24AC08n4B NCD34B m4CC4nC 4n ;Dm8OA4 ;X8mp;820C8on 34 ;0 AN6D;0C8on 34 ;X4GpA4BB8on 6Nn8qD4, 0D n8E40D CA03D2C8onn4;, 30nB ;0 ANponB4 8mmDn8C08A4. En BX0ppDH0nC BDA ;4B CA0E0DG 34 B70CC0270A944 et al. (2009), noDB 0EonB 4nCA4pA8B 34 20A02CNA8B4A, 0D n8E40D 24;;D;08A4, ;0 ANpA4BB8on 34 ;0 CA03D2C8on 34 ;XARN E8A0; p4n30nC ;0 ANponB4 8n3D8C4 p0A N. NoDB o1B4AEonB, 30nB Dn BHBCOm4 8n3D2C81;4 p4Am4CC0nC 34 3N2;4n274A ;0 ANponB4 8mmDn8C08A4 0pAOB ;XNC01;8BB4m4nC 34 ;X8n542C8on E8A0;4, Dn4
8E
;0A64 5oAm0C8on 34 6A0nD;4B K ARN 0pp4;NB PBB (PAo24BB8n6 Bo384B), 4n 2onBNqD4n24 34 ;X8n7818C8on BpN2858qD4 34 ;0 CA03D2C8on 34 ;XARN E8A0;. 4 mN20n8Bm4 45542C4DA 34 24CC4 ANpA4BB8on 4BC 3855NA4nC 34B mN20n8Bm4B 34 ANpA4BB8on CA03D2C8onn4;;4 m8B 4n p;024 4n ANponB4 K ;0 p4A24pC8on 34 BCA4BB C4; qD4 ;4B )VB, oD 34 ;X8n3D2C8on 34B mN20n8Bm4B 34 ANpA4BB8on CA03D2C8onn4;;4 p0A ;X8ARN. D4 p;DB, 34B p;0nC4B 3XArabid psis mDC0nC4B poDA Dn4 pAoCN8n4 5on2C8onn0nC 30nB ;4B PBB BonC p;DB ANB8BC0nC4B K 34B 102CNA84B p7HCop0C7o6On4B. DXDn4 50Mon 8nCNA4BB0nC4, on 3NC42C4 Dn n8E40D 3X4GpA4BB8on 0D6m4nCN 34 24AC08nB 6On4B 34 3N54nB4 30nB 24 mDC0nC, 34 m0n8OA4 B8m8;08A4 K 34B mDC0nCB 3XArabid psis 2ompAom8B 30nB ;XDn4 34B Eo84B 34 3N6A030C8on 34B ARNm. En p0A0;;O;4, noDB 0EonB 3NE4;oppN 34B oDC8;B CA0nB6Nn8qD4B poDA 3NC4Am8n4A B8 ;0 AN6D;0C8on 34 ;0 CA03D2C8on 34B ARNm 34 ;X7RC4, K ;XN274;;4 6Nnom8qD4, 9oD4 Dn AR;4 30nB ;XNC01;8BB4m4nC 34 ;X8mmDn8CN. NoDB 0EonB 6NnNAN Dn4 ;86nN4 3XArabids psis 4GpA8m0nC BoDB ;4 2onCAR;4 3XDn pAomoC4DA 8n3D2C81;4 ;4 502C4DA 3XAE8AD;4n24 (AEA) AEARpm1, 3onC ;0 pANB4n24 4BC A42onnD4 p0A ;0 pAoCN8n4 NB- RR RPM1; 4n 2om18n08Bon 0E42 Dn4 pAoCN8n4 A81oBom0;4 NC8qD4CN4 noDB p4Am4CC0nC 3X8mmDnopDA8584A ;4B A81oBom4B 0E42 ;4B ARNm 4n 2oDAB 34 CA03D2C8on. 4 BNqD4nM064 K 70DC-3N18C 34B ARN CoC0DG 4C 34B ARNm 4n606NB 30nB ;0 CA03D2C8on 0 ANEN;N qD4 28nq 5o8B p;DB 34 6On4B BonC AN6D;NB 0D n8E40D CA03D2C8onn4; p0A A0ppoAC 0D n8E40D CA0nB2A8pC8onn4;, 0pAOB ;X8n3D2C8on 34 ;0 ANponB4 NB- RR. )n4 0n0;HB4 2omp0AN4 0E42 ;4B 2onn08BB0n24B pAN2N34nC4B 0 ANEN;N qD4 ;0 ANponB4 RR 8n3D8C ;X4GpA4BB8on 34 20% 34B 6On4B NB- RR CoC0DG, qD8 BonC 0DBB8 2onBC8CDC8E4m4nC 4GpA8mNB 30nB 34B mDC0nCB 2ompAom8B 30nB 24AC08nB pAo24BBDB 34 3N6A030C8onB 34B ARNm. C4p4n30nC, ;0 ANponB4 NB- RR 8n3D8C ;X4GpA4BB8on 34 24nC08n4B 34 6On4B pAN2N34mm4nC 8mp;8qDNB 30nB ;4B 3N54nB4B, 24 qD8 nX4BC p0B ;4 20B 30nB ;4B mDC0nCB 34 3N6A030C8on 3XARNm, BoD;86n0nC 08nB8 ;0 BpN285828CN 34 ;0 ANponB4 NB- RR. )n4 0n0;HB4 18o-8n5oAm0C8qD4 0 3NC4Am8nN qDX4nE8Aon 28nq 24nC 6On4B BonC AN6D;NB 0D n8E40D CA03D2C8onn4; 34 m0n8OA4 8n3Np4n30nC4 34 ;4DA 01on30n24 CoC0;4, 5oDAn8BB0nC Dn4 ;8BC4 34 noDE40DG 20n3830CB poC4nC84;;4m4nC
E
8mp;8qDNB 30nB ;X8mmDn8CN NB- RR 4C BpN2858qD4m4nC AN6D;NB 0D n8E40D CA03D2C8onn4;. )n4 0n0;HB4 6NnNC8qD4 34 24AC08nB mDC0nCB 34 24B 6On4B 20n3830CB 30nB ;4B 3N54nB4B 0 NCN 4nCA4pA8B4, 4C 0 ANEN;N qD4 34B p;0nC4B 3XArabid psis (OR-3N58284nC4B oD mDC0nC4B poDA ;4B 6On4B CIPK5, CCT2 4C BIG poBBO34nC Dn4 ANB8BC0n24 0;CNAN4 poB8C8E4m4nC poDA ;4B (OR-3N58284nC4B oD nN60C8E4m4nC poDA ;4B 0DCA4B, p0A A0ppoAC K Dn4 p;0nC4 B0DE064. D0nB ;4DA 4nB4m1;4, 24B ANBD;C0CB pAoDE4nC Dn4 6A0n34 8mp;820C8on 34 ;0 AN6D;0C8on 34 ;0 CA03D2C8on 30nB ;0 m8B4 4n p;024 34 ;X8mmDn8CN EN6NC0;4.
E8
REMERCIEMEN(S
J4 C84nB K CA0nBm4CCA4 ;X4GpA4BB8on ;0 p;DB B8n2OA4 34 mon 8n58n84 6A0C8CD34 K P4C4A Mo554CC, ;X)n8E4AB8CN 34 S74A1Aoo:4 4C CoDC oA60n8Bm4 34 58n0n24m4nC poDA 0Eo8A 58n0n2N m4B CA0E0DG 34 3o2CoA0C. DXDn4 m0n8OA4 2AD280;4, 94 C84nB K A4m4A284A CoDB 24B 2o;;O6D4B 5on30m4nC0;4m4nC 7Dm08nB, 0D 6A0n3 2WDA 4n U oA m0BB85 V : (7NAOB4 W0;;on, Mo70m43 E; O8A38 4C C70nC0; BAoBB40D; m4A28 m4B 0m8B, 94 m4 BoDE84n3A08 34 EoDB CoDC4 m0 E84. DXDn po8nC 34 ED4 B284nC858qD4, 94 A4m4A284 34 noDE40D B8n2OA4m4nC P4C4A Mo554CC, 4C 0DBB8 ;4B pAo54BB4DAB SN10BC84n Ro3A86D4 4C P84AA4-JC84nn4 J02qD4B poDA ;X8mm4nB4 4Gp4AC8B4 qDX8;B onC 0ppoACN4 0D pAo94C 34 6Nnom8qD4 qD4 9X08 4D ;0 270n24 34 2on3D8A4. F8n0;4m4nC, m4A28 K M034mo8B4;;4 0DA4 BNA0A3 qD8 mX0 CoD9oDAB BoDC4nD 30nB m4B pA8B4B 34 3N28B8onB.
E88
(AB E DES MA(IERES
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7(f(7( x an cp rx an1 y1a i 1a 1an n ue an e d i(((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((88
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1a u a an((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((k4
7(f(f( 1c a i1i 1a x y i1 i 1 pi pcs n! n 1a x y n1d n1ca((((((((((((((((((((((k7
7(f(I( ! s .cp1ny c a ley cps i eca i nc n! is ee i1q e ii c i(((((((((((((((((((((((((((kI
7(f(?( c csuca ani a! a i 3 3s 1 n p i1in a 1a (((((((((k?
7(f(8( ! sxn an i!cli a! a p i1in a nc (((((((((kk
Cl1l e .ee mollllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllll’’
ClIl omc S o(e lllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllA1
G
7(8(6( e is1 cainpx n1ca(((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((àI
7(8(7( a p n1ca c np ai a1 ue ani((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((à?
7(8(C( pc n p1xs3s 1 n np ai1 an np ai cps n1ca (((((((((((((((((((((((((((((((((((((((((((((((((((((((((((à8
7(8(f( x pci xi!1ca ca anp n1ca c p1 cics i a ix pci p 1 an p n1ca n1ca(((((((((((((à8
7(8(I( n n1ca c 7 (((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((àk 7(8(?( 1pxi3 a x a 1e a 1a a u n!c a 1a n1cai((((((((((((((((((((((((((((((((((((((((((((((((((((((àà 7(8(8( x an1n n1d 3 ((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((àà 7(8(k( ! u ee icxp c (((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((644 Cl’l r o elllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgg: a (r hllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgC’
hl r oe (mS o te e m o( e o c Samr( o( mo(r .(mre (m è è
S ( SS.o (t lllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgCA
hlgl e(r (lllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgCA hlCl o(rm . ( mo llllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgCA
hlhl ( r e o S ( m ellllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllghC
C(C(6( e an pcln! ca 1n1cai a ((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6C7
C(C(7( 1 cics a onp n1ca(((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6C7
C(C(C( ssxa 3 ecnn1a a i1ed p in 1a1a (((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6CC
C(C(f( x pci ca anp n1ca c p1 cics i a ix pci p 1 an p n1ca n1ca ((((((((((((((((((((((((6CC
C(C(I( i r e1 p py cainpx n1ca((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6Cf
C(C(?( 1 ! n!pcx !uxn i rx a 1a (((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6Cf
C(C(8( a oup ii1ca a eyi1i ((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6Cf
C(C(k( a eyi1i ((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6CI C(C(à( n s1a1a (((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6CI hlpl e. (elllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllghI
C(f(6( a e i n! uxp1 1 n1ca c !1 ! rx e1ny np aie n s i xp1a ((((((((((((((((((((6C?
C(f(7( cssca a canp in1a nxp i nl a n! 1ssxa np aie ncs a np ai p1uncs
(((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6Ck
G8
C(f(f( a i csscaey en p y a 3 i1 a e1a ((((((((((((((((((((((((((((((((((((((((((((((((((6f7
C(f(I( an1 1 n1ca c 1 p an1 eey np aie n a i I iF 1a x y 6 i1 a e1a ((((((((6ff
C(f(?( d pe u c d1pxe an u n!c a3m !yuco1 a 631a x i((((((((((((((((((((((((((((((((((6f8
C(f(8( e1 n1ca c a 1 n a i(((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6fk
hl1l e .ee mollllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllg1f
hlIl omc S o(elllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllg1h
hl:l .aamr( o ( r o ( m e lllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllg1h C(8(6( 3 ou p1s an((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((((6IC hl’l r o elllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllg1p a (r pllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgIf pl e .ee mo o r ( mo .e mo llllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgIf oo n elllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllg:f oo n g L d ar e T Cfgpé lllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllg:f oo n C b .r e ( ( .n e.aaa S o( r e . a (r Cllllllllllllllllllllllllllllllllllllllllllllllllg:g oo n h b .r e ( ( .n e.aaa S o( r e . a (r hllllllllllllllllllllllllllllllllllllllllllllllllg’f llllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllllgAh
G88
IS(E DES ABBREVIA(IONS
4E-BP: 4E-B8n38n6 PAoC48n
ADN: A2834 DNBoGHA81onD2;NQqD4 AGO: AA6on0DC4
ARC: Ap05-1, R pAoC48n, CED-4 ARN: A2834 R81onD2;NQqD4 ARN31: ARN 3oD1;4 1A8n ARNA : ARN A81oBom0DG ARNC: ARN 34 CA0nB54AC
BAK1: BA0BB8noBC4Ao83-A2C8E0C43 K8n0B4 B8P: 8mmDno6;o1D;8n B8n38n6 PAoC48n BMV: BAom4 MoB082 V8ADB
CC: Co8;43-Co8;
CDPK: C0;28Dm-D4p4n34nC PAoC48n K8n0B4
CIPK5: C0;28n4DA8n B-InC4A02C8n6 PAoC48n K8n0B4 5 C;-VV: Cl ver yell w vein virus
CP: Co0C PAoC48n DC : D824A- 8:4
DCP: D420pp8n6 PAoC48n
DRB4: DoD1;4-BCA0n343 RNA B8n38n6 4 4IF: 4D:0AHoC82 In8C80C8on F02CoA
E(I: E5542CoA-(A8664A43 ImmDn8CH E((: E5542C4DA 34 (Hp4 (Ao8B F S2: F;064;;8n S4nB8n6 2
G888 GRP7: G;H28n-R827 PAoC48n 7
HBI1: Homo;o6 o5 B442 8nC4A02C8n6 F8C7 IBH1 HCV: H4p0C8C8B C V8ADB
HR: HHp4AB4nB8C8E4 R4BponB4 HRI: H4m4-A46D;0C43 4IF2α :8n0B4 8ARN: 8nC4A5NA4n24 K ARN
IRE1: InoB8Co;-R4qD8A8n6 EnIHm4 1 IRES: InC4An0; R81oBom4 EnCAH S8C4 KO: Kno2:-ODC
MAMP: M82Ao14-ABBo280C43 Mo;42D;0A P0CC4An MAPK: M8Co64n-A2C8E0C43 PAoC48n K8n0B4 M4C: MNC78on8n4
m8ARN : m82Ao ARN MID: M833;4
mRNP: m4BB4n64A R81onD2;4opAoC48n M(I: MAMP-(A8664A43 ImmDn8CH
m(OR: m0m0;;80n (0A64C o5 R0p0mH28n
n0CB8ARN: n0CDA0; 0nC8-B4nB B7oAC 8nC4A54A8n6 RNA NB- RR: ND2;4oC834 B8n38n6, 4D28n4-R827 R4p40C NMD: NonB4nB4-M4380C43 mRNA D420H
p70S6K: p70 S6 K8n0B4 PA.: P8F8/AA6on0DC4/.F8;;4
PABP: Po;H034nH;0C8on B8n38n6 PAoC48n
PAMP: P0C7o64n-ABBo280C43 Mo;42D;0A P0CC4An PB: PAo24BB8n6 Bo3H
PERK: PKR-;8:4 En3op;0Bm82 R4C82D;Dm K8n0B4
PGN: P4nC0CA82op4pC834 A4p40C pAoC48n 5oA G4Am8n0C8on on N0C; PKR: PAoC48n K8n0B4 RNA-34p4n34nC
G8E PR: P0C7o64n4B8B-R4;0C43
PRR: P0CC4An R42o6n8C8on R424pCoA P(I: PAMP-(A8664A43 ImmDn8CH
P(GS: PoBC-(A0nB2A8pC8on0; G4n4 S8;4n28n6 PV,: PoC0Co V8ADB ,
PV,ΔCP: PoC0Co V8ADB , B0nB ;XORF 34 Co0C PAoC48n R : RNB8BC0n24
RBP: RNA-B8n38n6 PAoC48n
R3RP: RNA-34p4n34nC RNA Po;Hm4A0B4 RE: RNC82D;Dm En3op;0Bm8qD4
RIN4: RPM1-InC4A02C8n6 pAoC48n 4 RIPK: RPM1-In3D243 PAoC48n K8n0B4 RISC: RNA-In3D243 S8;4n28n6 Comp;4G
RPM1: R4B8BC0n24 Co PB4D3omon0B p0C7oE0A M02D;82o;0 RPS2: R4B8BC0n24 Co PB4D3omon0B SHA8n604
BARN: B7oAC ARN
B8ARN: B7oAC 8nC4A54A8n6 ARN SG: SCA4BB GA0nD;4
SS(( : SHBCOm4 34 SN2ANC8on 34 (Hp4 (Ao8B (AS: (A0nB A2C8E0C8n6 BRNA
(BSV: (om0Co BDB7H SCDnC V8ADB (C: (4An0AH Comp;4G
(C(P: (A0nB;0C8on0;;H-ConCAo;;43 (DmoA PAoC48n (CV: (DAn8p CA8n:;4 V8ADB
(EV: T bacc etch virus
(IA-1: ( C4;; InCA024;;D;0A AnC864n 1
(IA-R: ( C4;; InCA024;;D;0A AnC864n 1-A4;0C43 (IR: (o;; 0n3 InC4A;4D:8n 1 R424pCoA 7omo;o6H
GE (MV: (o1022o MoB082 V8ADB
(DMV: Turnip M saic virus (.F: (0n34m .8n2 F8n64A
DORF: DpBCA40m Op4n R4038n6 FA0m4 )PR: )n5o;343 PAoC48n R4BponB4 )(R: )nCA0nB;0C43 R468on
)V: );CA0E8o;4CB
VSR: V8A0; SDpA4BBoA o5 RNA B8;4n28n6 ,RN: 4GoAH1omD2;40B4
GE8
IS(E DES (AB EA),
1. (01;40D 34B 3855NA4nCB 2ompoB0nCB 34 PAo24BB8n6 Bo384B, ;4DA oA60n8Bm4 34 AN5NA4n24 4C ;X4554C 34 ;4DA mDC0C8on. ... 49
GE88
IS(E DES FIG)RES
1. SCAD2CDA4 6NnNA0;4 34 ;XARNm. ... 3
2. 4 A81oBom4 4D20AHoC8qD4 (030pCN 34 R01; et al., 2011 ; R0m0:A8B7n0n, 2011). ... 4
3. 4 po;HBom4 (030pCN 34 Son4n14A6 4C D4E4A, 2003). ... 8
4. X8n8C80C8on 34 ;0 CA03D2C8on (030pCN 34 J02:Bon et al., 2010). ... 10
5. RN6D;0C8on 34 ;X8n8C80C8on 34 ;0 CA03D2C8on 274I ;4B 0n8m0DG (030pCN 3XHo;I et al., 2005). ... 13
6. 4B BHBCOm4B 4GCA0 4C 8nCA0-24;;D;08A4B 34 A42onn08BB0n24 3D p0C7o6On4 (030pCN 34 Do33B 4C R0C794n, 2010). Vo8A C4GC4 poDA 3NC08;B. ... 19
7. AA278C42CDA4 6NnNA0;4 3XDn4 NB- RR. ... 22
8. S27Nm0 34B 8nC4A02C8onB mo;N2D;08A4B 4nCA4 ;4B 45542C4DAB AEAB, AEARpm1 oD AEARpC2 0E42 ;4B 02C4DAB 34 ;X8mmDn8CN EN6NC0;4. Vo8A C4GC4 poDA 3NC08;B. ... 26
9. 4B 3855NA4nCB CHp4B 3X8nC4A02C8onB AEA-R. ... 29
10. 4B NC0p4B 34 ;X02C8E0C8on 3XDn4 NB- RR (030pCN 34 Co;;84A 4C Mo554CC, 2009). .... 32
11. VD4 2A8BC0;;o6A0p78qD4 6NnNA0;4 3XDn4 AA6on0DC4 (030pCN 34 Son6 et al., 2004 ; J8n4: 4C DoD3n0 et al., 2009). ... 38
12. VD4 2A8BC0;;o6A0p78qD4 34B 3om08n4B 5on2C8onn4;B 3XDn4 AA6on0DC4 (030pCN 34 J8n4: 4C DoD3n0, 2009). ... 41
13. X8ARN 2omm4 mN20n8Bm4 0nC8E8A0; (030pCN 34 D8n6, 2010). ... 44
14. 4B Eo84B 4n3o6On4B 34 pAo3D2C8on 34 B8ARN. ... 46
15. 4B Eo84B 34 3N6A030C8onB 34 ;XARNm (030pCN 34 D42:4A 4C P0A:4A, 2012). ... 53
16. 4 2H2;4 34 ;XARN. ... 55
17. PAo58;B 34 po;HBom4B CHp8qD4B 0pAOB 2on24nCA0C8on p0A D;CA024nCA85D60C8on oD 8mmDnopDA85820C8on (030pCN 34 .0n4CC8 et al., 2005). ... 61 1
1 CHAPI(RE 1
1. IN(ROD)C(ION GENERA E
AD 2onCA08A4 34B oA60n8Bm4B 0n8m0DG, ;4B 24;;D;4B EN6NC0;4B n4 poBBO34nC p0B 34 BHBCOm4 28A2D;0Co8A4 p4Am4CC0nC ;0 m8B4 4n p;024 3XDn BHBCOm4 8mmDn8C08A4 030pC0C85 C4; qDXon ;4 2onn08C 274I ;X7Dm08n. C4p4n30nC, 0D n8E40D mo;N2D;08A4, Dn 346AN B86n85820C85 34 B8m8;0A8CN 4G8BC4 0E42 ;4 BHBCOm4 8mmDn8C08A4 0n8m0;, 2omm4 ;0 BDAE48;;0n24 8nCA0-24;;D;08A4 34 ;0 pANB4n24 34 ;X4nE078BB4DA p0A ;4B pAoCN8n4B RRB (M04:0F0 et al., 2011), oD p0A ;0 m0278n4A84 3X8ARN 30nB ;4 20B 34B E8ADB (M08;;0A3 et al., 2013; R08A30n 0n3 Mo554CC, 2007). C4p4n30nC, ;4B 20B2034B 34 B86n0;8B0C8on 8n3D8C4B p0A ;4B NB- RRB 4C m4n0nC K ;X8mmDn8CN n4 BonC qD4 CAOB p4D N;D283N4B. D4 nom1A4DB4B NCD34B 20A02CNA8B0nC ;4 CA0nB2A8pCom4 8mmDn8C08A4 onC 5oDAn8 CAOB p4D 34 6On4B 20n3830CB K ;0 m8B4 4n p;024 34 ;X8mmDn8CN 3Np4n30nC4 34B NB- RRB. X0n0;HB4 4n pAo5on34DA 34 ;0 AN6D;0C8on 34 ;0 CA03D2C8on p4n30nC ;0 ANponB4 NB- RR, 08nB8 qD4 ;4B 2onn4G8onB 4nCA4 ;0 Eo84 34B NB- RR 4C 24;;4 34 ;X8ARN 4BC 3on2 2AD280;4 poDA Dn4 2ompAN74nB8on 8nCN6A0C8E4 34 ;X8mmDn8CN EN6NC0;4. C4CC4 8nCAo3D2C8on 34 C7OB4 0DA0 3on2 2omm4 o1942C85 34 3N58n8A ;4B 2on24pCB nN24BB08A4B K ;0 2ompAN74nB8on 34B NCD34B pD1;8N4B oD 4n Eo84 34 ;XPCA4 5oAm0nC ;4B 34DG8Om4 4C CAo8B8Om4 270p8CA4B 34 24CC4 C7OB4 : 1) 4B NC0p4B 20non8qD4B 34 ;0 CA03D2C8on; 2) 0 AN6D;0C8on 34 ;0 CA03D2C8on; 3) 4 BHBCOm4 8mmDn8C08A4 EN6NC0;; 4) 4 BHBCOm4 3X8ARN; 5) 4 mNC01o;8Bm4 34 ;XARN 4C 58n0;4m4nC 6) ;0 6Nnom8qD4 3D BHBCOm4 8mmDn8C08A4 EN6NC0; ; 4n 60A30nC CoD9oDAB 2omm4 o1942C85 34 2onn42C4A CoDB 24B 2on24pCB.
2 1.1. La traduction : de l’ARNm à la protéine.
1.1.1. Les acteurs principaux de la traduction eucaryotique.
On 38BC8n6D4 p;DB84DAB pAoC06on8BC4B 30nB ;0 CA03D2C8on qD8 BonC ;XARNm, ;4 A81oBom4, ;4B 502C4DAB 3X8n8C80C8on 4D20AHoC8qD4B, ;4B 502C4DAB 3XN;on60C8on 4C ;4B 502C4DAB 34 C4Am8n08Bon. X8n8C80C8on 34 ;0 CA03D2C8on 4BC ;XNC0p4 m094DA4 34 AN6D;0C8on 34 ;0 CA03D2C8on, 4C 34 24 508C, noDB n4 3NC08;;4AonB p0B ;0 AN6D;0C8on p0A ;4B 502C4DAB 3XN;on60C8on n8 34 C4Am8n08Bon. XARNm 2onC84nC ;X8n5oAm0C8on 6NnNC8qD4 BoDB 5oAm4 3X02834 A81onD2;N8qD4 4C 4BC oA60n8BN 4n 2o3onB. X8n8C80C8on 34 ;0 CA03D2C8on 2oAA4Bpon3 K ;X0BB4m1;064 3XDn A81oBom4 80S 2ompNC4nC poDA ;XN;on60C8on p4pC838qD4 0D 2o3on 34 3Np0AC 34 ;0 CA03D2C8on 34 ;XARNm, 6AL24 K ;X0834 34B 502C4DAB 3X8n8C80C8on 4D20AHoC8qD4 (4IF).
1.1.1.1. L’ARN messager (ARNm).
XARNm 4BC 8BBD 3D mN20n8Bm4 34 CA0nB2A8pC8on. I; 2onC84nC ;X8n5oAm0C8on 6NnNC8qD4 3onnN4 p0A ;XADN 4n ED4 34 pAo3D8A4 Dn4 pAoCN8n4 BpN2858qD4, 08nB8 qD4 34B BNqD4n24B 05542C0nC ;X4GpA4BB8on 3D 6On4 0D n8E40D CA03D2C8onn4;. ApAOB ;0 po;HmNA8B0C8on 3XDn 1A8n 3XARN 0E42 Dn 1A8n 3XADN m0CA824 6AL24 K ;XARN po;HmNA0B4 II, ;XARNm 4BC BD224BB8E4m4nC 2o855N 4n 5X, B4B 8nCAonB BonC Np8BBNB 34 50Mon K pAo3D8A4 Dn 203A4 oDE4AC 34 ;42CDA4 (ORF), 4C ;0 qD4D4 po;H(A) 4BC 09oDCN4. C4B NC0p4B BonC CoDC4B nD2;N08A4B 4C nN24BB08A4B K ;XARN poDA p0BB4A ;4B NC0p4B 34 2onCAR;4 34 qD0;8CN. V80 ;X8nC4A02C8on 0E42 34B pAoCN8n4B ;80nC ;XARN (RNA-18n38n6
3
pBoD59nC ; RBPC), lZARNm 5CD 5HpoBDN 4E noI1E 1E 3IDoCol poEB PDB5 DB14E9D. L5C CDBE3DEB5C 16653D1nD l1 DB14E3D9on ConD 4N3B9D5C 39-45CCoEC (697. 1).
Figure 1. Structure généra.e de .’ARNm.
L1 CDBE3DEB5 3o9665 m7GpppN (oT N B5pBNC5nD5 nZ9mpoBD5 AE5l nE3lNoD945), 5D l1 AE5E5 polI (A)H (oT H 5CD l5 nom2B5 4Z14NnoC9n5) ConD 45C CDBE3DEB5C 6oBD5m5nD 9mpl9AEN5C 41nC l1 pBomoD9on 45 lZ9n9D91D9on 45 l1 DB14E3D9on. EnDB5 l1 3o9665 5D l5 3o4on 4Z9n9D91D9on 45 l1 DB14E3D9on, 5D l5 3o4on CDop 5D l1 AE5E5 polI(A) C5 DBoEF5nD l5C *)RC. C5C CNAE5n35C n5 ConD p1C DB14E9D5C 5D p5EF5nD PDB5 9mpl9AEN5C 41nC l1 BN7El1D9on 45 l1 DB14E3D9on. L5C IRE( p5Bm5DD5nD l5 B53BED5m5nD 45C B92oCom5C (5n BoE75), l5C CDBE3DEB5C 5n Np9n7l5C K 385F5EH (D975-2oE3l5 5n no9B) ConD CoEF5nD l9N5C p1B 45C RBPC, AE9 p5EF5nD 1ECC9 l95B 4Z1EDB5C CNAE5n35C (oF1l5C 2l5E5C). D1nC 35BD19nC ARNm, En5 BN79on 5n 1monD 4E 314B5 45 l53DEB5 (E%RF) 5CD 9mpl9AEN5 41nC lZ9n8929D9on 45 l1 DB14E3D9on 45 lZ%RF pB9n39p1l.
1.1.1.2. Le ribosome.
C5C B92onE3lNopBoDN9n5C ConD 45C CDBE3DEB5C 3ompoCN5C K 40% 45 pBoDN9n5 5D K 60% 4ZARN B92oCom1EH (ARNB). P1B B1ppoBD 1EH pBo31BIoD5C, l1 CDBE3DEB5 B92oCom1l5 5E31BIoD5 5CD C97n96931D9F5m5nD plEC 9mpoC1nD5, 1F53 45EH ARNB 5D 45 pBoDN9n5C CEpplNm5nD19B5C, 1E7m5nD1nD l5 po94C molN3El19B5 45 1700 ;D1 (2600 ;D1 3onDB5
5
1.1.1.3. Les eIFs et autres protéines régulatrices.
%n 49CD9n7E5 DBo9C 7B1n45C 31DN7oB95C 45 pBoDN9n5C AE9 BN7El5nD lZ9n9D91D9on 45 l1 DB14E3D9on, 4onD 9) l5C 5IF2, 9mpl9AEN5C 41nC l1 4Nl9FB1n35 45 lZARND 9n9D91D5EB, 99) l5C 5IF4, 9mpl9AEN5C 41nC lZ13D9F1D9on 45 lZARNm 1F53 999) l5C pBoDN9n5C l91nD l5C C9D5C polI(A) (PABP). L5C 5IF2 5D 4 ConD 1ECC9 45C 392l5C 9mpoBD1nD5C poEB l5 3onDBSl5 45 lZ9n9D91D9on l1 DB14E3D9on.
1.1.1.3.1. Fonctions des eIF2.
PB5m9OB5m5nD, 5IF2 5CD 3ompoCN5 45 DBo9C CoEC-En9DNC, α β 5D γ, 5D 5CD 9mpl9AEN5 41nC l1 4Nl9FB1n35 45 lZARND 9n9D91D5EB M5D-ARND9M5D 41nC l5 C9D5 P 45 l1 p5D9D5 CoEC-En9DN B92oCom1l5. L1 l919Con 4ZEn5 molN3El5 45 G)P 5CD 9n49Cp5nC12l5 K l1 CD129l9C1D9on 45 l1 6oBm1D9on 4E 3ompl5H5 D5Bn19B5 ()C) 5IF2*G)P*M5D-ARND9M5D. ApBOC l5 381B75m5nD 4E )C 41nC l5 B92oCom5 5D Con 4Npl135m5nD 41nC En5 49B53D9on 5Z 3Z, lZ1pp1B95m5nD 5nDB5 l5 3o4on A*G 5D lZ1nD93o4on 45 lZARND 9n9D91D5EB pBoFoAE5 lZ8I4BolIC5 4E G)P 5n GDP. C5DD5 BN13D9on nN35CC9D5 5IF5. 5IF2*GDP 5CD 5nCE9D5 B53I3lN5 p1B 5IF2B 5n 5IF2*G)P. C5DD5 ND1p5 45 B53I3l175, 4on3 l1 49Cpon929l9DN 5n 3ompl5H5 D5Bn19B5, 5CD lZEn5 45C 45EH 392l5C m1:5EB5C poEB l5 3onDBSl5 45 l1 DB14E3D9on. NoD1mm5nD, l1 p8oCp8oBIl1D9on 4E BNC94E (5B51 3onF5BD9D 5IF2 5n 9n8929D5EB 3ompND9D96 45 5IF2B plEDSD AEZ5n CE2CDB1D ((on5n25B7 1n4 D5F5B, 2003).
6
1.1.1.3.2. Fonctions des eIF4.
D5EH9Om5m5nD, 9nDNB5CConC-noEC 1EH 5IF4 p5Bm5DD1nD l5 B53BED5m5nD 4E B92oCom5 CEB lZARNm. %n 4Nnom2B5 5IF4A, 5IF4B, 5IF4E, 5IF4H 5D 5IF4G. 5IF4A, 4E 5D 4G 6oBm5nD En 3ompl5H5 D5Bn19B5 1pp5lN 5IF4F poEF1nD l95B l1 CDBE3DEB5 3o9665 5n 5Z F91 l1 CoEC-En9DN 5IF4E. 5IF4A 1CCo39N K 5IF4B poCCO45 En5 13D9F9DN 8Nl931C5 K ARN 9mpl9AEN5 41nC lZ1pl1n9CC5m5nD 45C CDBE3DEB5C C53on419B5C poEB p5Bm5DDB5 1E B92oCom5 45 pBo7B5CC5B l5 lon7 45 lZARN. 5IF4G 5CD En5 CoEC-En9DN 3BE391l5 pE9CAEZ5ll5 9nD5B179D 1F53 l5C pBoDN9n5C pBNC5nD5C CEB l1 AE5E5 polI(A), 5D p5Bm5D 19nC9 l1 6oBm1D9on 45 l1 2oE3l5 45 lZARNm. L5C 5IF4 ConD 1ECC9 45C 392l5C poD5nD95ll5C poEB BN7El5B l1 DB14E3D9on, p1B p8oCp8oBIl1D9on oE p1B CNAE5CDB1D9on 7BL35 1EH pBoDN9n5C l91nD l5C 5IF4E (4E-BP), AEZon n5 3onn1îD p1C 385J l5C FN7ND1EH (G9n7B1C et al., 1999). C85J l5C FN7ND1EH, on o2C5BF5 En5 3ompl5H96931D9on 45C 5IF4. En 5665D, 5IF4E 5D 5IF4G poCCO45nD 45C 9Co6oBm5C 1pp5lN5C B5Cp53D9F5m5nD 5IF(9Co)4E 5D 5IF(9Co)4G1 5D (9Co)4G2, poEF1nD 6oBm5B 1F53 5IF4A l5 3ompl5H5 5IF(9Co)4F. L5C 45EH 9Co6oBm5C 45 5IF4E p1BD175nD 50% 4Z945nD9DN 41nC l5EB CNAE5n35 4Z13945C 1m9nNC 5D onD l5 mPm5 po94C molN3El19B5, 1loBC AE5 5IF4G 5D 5IF(9Co)4G pOC5nD B5Cp53D9F5m5nD 180 ;D1 5D 86 ;D1. Comp1BN5 K l1 p1BD95 N-D5Bm9n1l5 45 l1 7B1n45 9Co6oBm5, l1 p5D9D5 5CD 4NpoEBFE5 45 plEC95EBC 4om19n5C, 35 AE9 CE77OB5 45C 6on3D9onC 5D/oE 45C mN31n9Cm5C 45 BN7El1D9on 4966NB5nDC (BBoGn9n7, 2004). L1 C97n96931D9on 6on3D9onn5ll5 45 lZ5H9CD5n35 45 45EH 3ompl5H5C 5IF4F 385J l5C pl1nD5C 5CD 5n3oB5 5m2BIonn19B5. %n C19D p1B 5H5mpl5 AE5 5IF(9Co)4F 5CD plEC 12on41nD AE5 5IF4F 1E CD145 5m2BIonn19B5 4E 2lN, 41nC lZ1p5H B139n19B5 45 m1RC 5D 41nC l5C 9n6loB5C35n35C 4ZArabidopsis. D5 plEC, 5IF4F 5CD plEC 5669395nD 41nC l1 DB14E3D9on 45C ARNm 3onD5n1nD 45C BN79onC 5Z*)R 3ompl5H5C (5H. 3onD5n1nD 45C IRE() 1loBC AE5 5IF(9Co)4F 5CD 4Np5n41nD 45 l1 3o9665 (K1G17E389 1n4 B19l5I-(5BB5C, 2002). RN35mm5nD, 45EH noEF5ll5C 9Co6oBm5C 45 5IF4E, 5IF4E12 5D 13, onD NDN 4N3oEF5BD5C
7
5D 31B13DNB9CN5C in vitro m19C l5EB 6on3D9on in vivo B5CD5 9n5Hpl9AEN5 (P1DB93; et al., 2014).
1.1.1.4. Fonction de la protéine liant les sites poly(A).
En69n, CoEl97nonC lZ9mpoBD1n35 45C pBoDN9n5C pBNC5nD5C K lZ5HDBNm9DN 3Z 45 lZARNm. Il 1 9n9D91l5m5nD NDN pBoEFN AE5 l1 pBNC5n35 4ZEn5 AE5E5 polI(A) ND19D nN35CC19B5 K l1 DB14E3D9on 45 lZARNm 385J l1 l5FEB5, l1 4BoCop89l5 5D l5C pl1nD5C. Il 5H9CD5 plEC95EBC pBoDN9n5C 3onnE5C poEB 9nD5B179B 1F53 l5C C9D5C 45 polI14NnIl1D9on. NoD1mm5nD, l1 AE5E5 polI(A) 5CD 8129llN5 4ZEn5 pBoDN9n5 45 70;D1 poEF1nD l95B l5C C9D5C polI14NnIlNC (PABP). L1 pB5m9OB5 NF945n35 6on3D9onn5ll5 1 NDN 4Zo2C5BF5B lZ5665D CIn5B79AE5 45 l1 3o9665 5D 45 l1 AE5E5 polI(A) CEB l1 CD9mEl1D9on 45 l1 DB14E3D9on (G1ll95 1n4 )1n7E1I, 1994). Il 1 p1B l1 CE9D5 NDN monDBN AE5 l1 PABP p5Bm5DD19D K l1 AE5E5 polI(A) 4Z9nD5B179B 1F53 l1 3o9665 F91 5IF4G (Im1D1;1 et al., 1998). A9nC9, 35DD5 9nD5B13D9on p5Bm5D l1 6oBm1D9on 45 l1 2oE3l5 45 lZARNm AE9 5CD nN35CC19B5 K lZ9n9D91D9on 45 l1 DB14E3D9on. LZEn 45C mN31n9Cm5C CE77NBNC 5CD AE5 lZ9nD5B13D9on 5nDB5 l1 3o9665 5D l1 AE5E5 polI(A) p5Bm5D 1EH B92oCom5C 45 49B53D5m5nD B5DoEBn5B K lZ5HDBNm9DN 5Z 1pBOC 1Fo9B pBo7B5CCN l5 lon7 45 lZARNm (697. 3). *n ARNm 9n9D9N 45 plEC95EBC B92oCom5C 5CD 1pp5lN l5 polICom5 5D 1 4N:K NDN o2C5BFN 5n m93BoC3op95 Nl53DBon9AE5 ((on5n25B7 1n4 D5F5B, 2003). *n polICom5 5CD 3onC94NBN 3omm5 DB14E3D9onn5ll5m5nD 13D96, 295n AE5 35BD19nC 5H5mpl5C 9n49AE5nD, 41nC 35BD19nC 31C DBOC CpN3969AE5C, AEZEn ARNm p5ED PDB5 41nC l5C polICom5C, 41nC l5CAE5lC l5 moEF5m5nD 45C B92oCom5C 5CD 2loAEN p1B 35BD19n5C pBoDN9n5C CpN3969AE5C (D1Bn5ll et
9
DB14E3D9on 619D 9nD5BF5n9B En5 F9n7D19n5 45 polIp5pD945C 4onD En5 4oEJ19n5 45 613D5EBC 4Z9n9D91D9on 5E31BIoD9AE5 (5IF), l1 plEp1BD nZ1I1nD p1C 4Z8omolo7E5 pBo31BIoD9AE5 (G521E5B 1n4 H5nDJ5, 2004). A D9DB5 45 3omp1B19Con, l5C pBoDN9n5C 9mpl9AEN5C 41nC l1 p81C5 4ZNlon71D9on 5D 45 D5Bm9n19Con ConD 3onC5BFN5C K DB1F5BC l5C BO7n5C. LZ9n9D91D9on 45 l1 DB14E3D9on 5CD l5 pBo35CCEC 4Z1CC5m2l175 45 B92oCom5C 80( 3ompND5nDC poEB l1 p81C5 4ZNlon71D9on p5pD949AE5, 41nC l5CAE5lC l5 3o4on 4Z9n9D91D9on 5CD 1pp1B9N 1F53 l1 2oE3l5 45 lZ1nD93o4on 45 lZARN 45 DB1nC65BD 9n9D91D5EB (M5D-DRNAM5D). Ell5 5CD 49F9CN5 5n 45EH ND1p5C 4onD 9) l1 6oBm1D9on 4E 3ompl5H5 4Z9n9D91D9on 48( F91 l1 B53onn19CC1n35, 41nC En5 49B53D9on 5Z 3Z, 4E 3o4on 4Z9n9D91D9on 45 lZARNm p5Bm5DD1nD 4ZND12l9B En 1pp1B95m5nD 1F53 lZ1nD93o4on 45 lZARND 5D 99) lZ1CCo391D9on 45 l1 CoEC-En9DN 48( 1F53 l1 CoEC-En9DN B92oCom1l5 60( 6oBm1nD l5 3ompl5H5 4Z9n9D91D9on 80( (697. 4). (EB l1 plEp1BD 45C ARNm 35llEl19B5C, 35 pBo35CCEC CZ56653DE5 p1B En mN31n9Cm5 45 W C31nn9n7 X, oT l5 3ompl5H5 45 pBN9n9D91D9on 43( (3ompB5n1nD En5 CoEC-En9DN 40(, l5 3ompl5H5 D5Bn19B5 5D 45C 5IFC 14F5nD9F5C) CZ1DD1385 K lZ5HDBNm9DN 5Z 45 l1 CNAE5n35, pBN1l12l5m5nD 1pl1n95 p1B l5 3ompl5H5 5IF4F :ECAEZK l1 B5n3onDB5 4ZEn A*G (697. 4). L5 4Npl135m5nD 4E B92oCom5 5CD A)P-4Np5n41nD. LZ1pp1B95m5nD 5D l1 6oBm1D9on 4E 3ompl5H5 48( 4N3l5n385 lZ8I4BolIC5 4E G)P 1CCo39N K 5IF2 p1B 5IF5, l5 4Npl135m5nD 45C 5IFC 5D l5 B53BED5m5nD 45 l1 CoEC-En9DN 60( ( 13;Con et al., 2010). ApBOC lZ1:oED 45 l1 mND89on9n5 45 4Np1BD, CE9D lZND1p5 4ZNlon71D9on, oT l1 CInD8OC5 4ZEn ponD p5pD949AE5 (C9D5 P) p1B l5 B92oCom5 5CD 4Np5n41nD5 45 lZ1CCo391D9on 5nDB5 l5 3o4on 5D lZARND 1m9no-13IlN 3oBB5Cpon41nD (C9D5 A). C5DD5 BoED9n5 CZ1BBPD5 loBC 45 l1 B5n3onDB5 4ZEn 3o4on CDop. C5p5n41nD, noEC 3onn19CConC m19nD5n1nD 45C mN31n9Cm5C 45 DB14E3D9on 4onD lZ9n9D91D9on 5CD 9n4Np5n41nD5 45 l1 3o9665. C5 mo45 45 DB14E3D9on 5CD noD1mm5nD ED9l9CN p1B 35BD19nC F9BEC 3onD5n1nD 45C CNAE5n35C 9nD5Bn5C 4Z5nDBN5 45 B92oCom5 (IRE( ; 9nD5Bn1l B92oCom5 5nDBI C9D5C) 41nC l5EB 5HDBNm9DN 5Z.
11
1.2. Régu.ati1n traducti1nne..e en c1nditi1ns de stress.
GNnNB1l5m5nD, on 49CD9n7E5 45EH DIp5C 45 CDB5CC. C5EH 31ECNC p1B 45C 613D5EBC 5nF9Bonn5m5nD1EH 5D 35EH 31ECNC p1B l5C oB71n9Cm5C F9F1nDC, B5Cp53D9F5m5nD 1pp5lNC CDB5CC 129oD9AE5C 5D 29oD9AE5C. NoEC n5 4ND19ll5BonC AE5 lZ9mp13D 45C CDB5CC 29oD9AE5C CEB l1 DB14E3D9on. %n 49CD9n7E5 45EH 392l5C m1:5EB5C poEB l1 BN7El1D9on 45 lZ9n9D91D9on 45 l1 DB14E3D9on AE5 ConD 5IF2α 5D 5IF4E. LZ13D9F9DN 45 5IF2α 5CD BN7ElN5 p1B Con ND1D 45 p8oCp8oBIl1D9on. PlEC95EBC ;9n1C5C ConD 9mpl9AEN5C 41nC l1 p8oCp8oBIl1D9on 45 5IF2α m19C ConD 4966NB5nD95ll5m5nD 13D9FN5C 5n 6on3D9on 4E CDB5CC p5BME. L1 pBoDN9n5 5IF4E 5CD 1ECC9 BN7ElN5 p1B p8oCp8oBIl1D9on (1pBOC l1 6oBm1D9on 4E 3ompl5H5 43() m19C lZ5CD pB9n39p1l5m5nD F91 En mN31n9Cm5 4Z9nD5B13D9on 1F53 En 9n8929D5EB 3ompND9D96 : l5C 4E-BP (1E CD145 4Z9n9D91D9on). LZ9nD5B13D9on 5nDB5 35C pBoDN9n5C 5CD 3onDBSlN5 p1B l1 p8oCp8oBIl1D9on 45C 4E-BP F91 l1 Fo95 m)%R/p70(6K (697. 5). P1B B1ppoBD 1EH 35llEl5C 1n9m1l5C, 35C 45EH Fo95C B5CD5nD o2C3EB5C 385J l5C pl1nD5C, pE9CAE5 lZon n5 DBoEF5 p1C 4ZoBD8olo7E5 45C 4E-BP n9 45 35BD19n5C ;9n1C5C K 5IF2α DBoEFN5C 385J l5C m1mm96OB5C.
1.2.1. La fami..e des kinases à eIF2.
D1nC l1 CEp5B61m9ll5 45C pBoDN9n5C ;9n1C5C, on 49CD9n7E5 l1 61m9ll5 45C ;9n1C5C 5IF2α 3ompoCN5 45 AE1DB5 m5m2B5C 385J l5C m1mm96OB5C. En 6on3D9on 4E CDB5CC p5BME, 3813En5 45C ;9n1C5C p8oCp8oBIl5 5IF2α CEB l5 BNC94E CNB9n5 51. L1 p8oCp8oBIl1D9on CEB 35 BNC94E 5mpP385 l5 613D5EB 4ZN381n75 5IF2B 45 B53I3l5B 5IF2*GDP 5n 3ompl5H5 D5Bn19B5 G)P*5IF2*M5D-ARND9M5D, BNCElD1nD 5n En5 5HD9n3D9on 7NnNB1l5 45 l1 DB14E3D9on (697. 5B). C5C ;9n1C5C nZonD p1C 3omm5 392l5 5H3lEC9F5 5IF2α, m19C
12
p5EF5nD 1ECC9 p8oCp8oBIl5B 45C 613D5EBC 45 DB1nC3B9pD9on (R1F5n 1n4 KoBom9l1C, 2008). C5DD5 61m9ll5 3ompB5n4 l1 pBoDN9n5 ;9n1C5 13D9FN5 p1B lZARN 4oE2l5-2B9n (PKR ; PBoD59n K9n1C5 RNA-45p5n45nD), :oE1nD En BSl5 9mpoBD1nD 41nC l1 BNponC5 35llEl19B5 1nD9F9B1l5, l1 pBoDN9n5 45 DIp5 PKR 4E BND93ElEm 5n4opl1Cm9AE5 (PERK ; PKR-l9;5 En4opl1Cm93 R5D93ElEm) 13D9FN5 p1B lZ133EmEl1D9on 45 pBoDN9n5C non 3on6oBm5C 41nC 35DD5 oB71n9D5, l1 pBoDN9n5 G5n5B1l ConDBol Non-45B5pB5CC92l5 2 (GCN2) 13D9FN5 p1B l5 m1nAE5 4Z13945 1m9nN 5D En5 45Bn9OB5 9mpl9AEN5 41nC l1 4ND53D9on 45C 31B5n35C 5n noI1E 8Om5 (HRI ; 85m5-B57El1D54 5IF2α ;9n1C5). L5 C5El 8omolo7E5 FN7ND1l 3onnE 5CD GCN2. A 35 CE:5D, En5 13D9F9DN 45 DIp5 PKR 1 NDN 4ND53DN5 41nC 45C 5HDB19DC 3IDopl1Cm9AE5C FN7ND1EH m19C C1nC B5monD5B K l1 CNAE5n35 7Nnom9AE5 (L1n7l1n4 et
al., 1996). (EB 35DD5 21C5 6B912l5 1 NDN 4N3oEF5BD En 9n8929D5EB 35llEl19B5 FN7ND1l 45 l1
CEppoCN5 PKR, P58IPK (B9l79n et al., 2003). C85J l5C m1mm96OB5C, l1 4ND53D9on 45 lZARN 4oE2l5 2B9n p1B l1 PKR 5nDB19n5 CE335CC9F5m5nD l1 6oBm1D9on 4ZEn 49mOB5 pE9C C1 p8oCp8oBIl1D9on, 5nDB19n1nD 19nC9 Con 13D9F1D9on (G9l 1n4 M1B91no, 2000). LZ5H9CD5n35 4ZEn5 PERK FN7ND1l5 5CD 1ECC9 9n3onnE5. C85J l5C m1mm96OB5C, l1 p5B35pD9on 4ZEn C97n1l 45 CDB5CC lo31l9CN 41nC l5 RE (8omNoCD1C95 4E C12+, pBoDN9n5C 4Npl9N5C) pBoFoAE5 l1 49CCo391D9on 4E 3ompl5H5 PERK-B9P, 5D 4ZEn5 m1n9OB5 C9m9l19B5 K l1 PKR, lZ8omo49mNB9C1D9on PERK-PERK p5Bm5DD1nD Con 13D9F1D9on p1B p8oCp8oBIl1D9on (Ron 1n4 ,1lD5B, 2007). C5p5n41nD, 35BD19nC 1BD93l5C 4NmonDB5nD lZ5H9CD5n35 45 l1 Fo95 13D9FN5 p1B l5 CDB5CC 4E BND93ElEm 5n4opl1Cm9AE5, 1pp5lN5 BNponC5 1EH pBoDN9n5C 4Npl9N5C (*PR ; *n6ol454 PBoD59n R5CponC5). C85J l5C pl1nD5C 5D l5C 1n9m1EH, l1 Fo95 IRE1 (InoC9Dol B5CponC9F5 5l5m5nD) 3onDBSl5 lZ5HpB5CC9on 45C 7On5C nN35CC19B5C K lZ*PR, mPm5 C9 lZ8omolo7E5 FN7ND1l 45 IRE1 nZ1 p1C 5n3oB5 NDN 945nD969N (C85n 1n4 BB1n49JJ9, 2013; M9C89211 et al., 2013).
14 1.2.2. La v1ie OR.
C85J l5C 5E31BIoD5C, l1 )1B75D o6 R1p1mI39n5 ()%R) 5CD 3o4N5 p1B En En9AE5 lo3EC. C5DD5 pBoDN9n5 5CD lZEn 45C po9nDC 45 3onDBSl5C 35llEl19B5C m1:5EBC pE9CAEZ5ll5 K l1 31p139DN 4Z9n8925B oE 45 CD9mEl5B lZ9n9D91D9on 45 l1 DB14E3D9on CEppoCNm5nD 4ZEn5 m1n9OB5 7NnNB1l5. L5C 392l5C pB9n39p1l5C 45 )%R ConD l5C (6 ;9n1C5C ((K1 5D (K2) 5D l5C 4E-BP. L1 p8oCp8oBIl1D9on 45C 4E-BP 5nDB19n5 l5EB 49CCo391D9on 45C 5IF4E, p5Bm5DD1nD l1 6oBm1D9on 4E 3ompl5H5 5IF4F. En 35 AE9 3on35Bn5 l5C 392l5C 45C (6K, 9l CZ179D pB9n39p1l5m5nD 45 l1 pBoDN9n5 B92oCom1l5 (6 19nC9 AE5 45 5IF4B AE9 ConD 13D9FN5C p1B p8oCp8oBIl1D9on (697. 5A) (HolJ et al., 2005). C85J l5C pl1nD5C, 35DD5 Fo95 C5m2l5 3onC5BFN5. En 5665D, Arabidopsis thaliana poCCO45 En5 En9AE5 3op95 4E 7On5 )%R, AD)%R (M5n1n4 et al., 2002). AECC9, l5C 392l5C 5n 1F1l C5m2l5nD PDB5 3onC5BFN5C, D5ll5 l1 (6 ;9n1C5, 295n AEZ1E3En 8omolo7E5 45 4E-BP n5 Co9D 3onnE. D5 plEC, 3onDB19B5m5nD 1EH CICDOm5C 1n9m1EH, AD)%R 5CD 9nC5nC92l5 K l1 B1p1mI39n5. %B C9 on 3omplNm5nD5 Arabidopsis 1F53 l1 pBoDN9n5 FKBP12 45 lZ8Em19n oE 35ll5 45 l1 l5FEB5, l1 C5nC929l9DN 45 AD)%R K l1 B1p1mI39n5 5CD BND12l95 ((oBm1n9 et al., 2007). D5 35 619D, lZ9nC5nC929l9DN 4ZAD)%R K l1 B1p1mI39n5 C5B19D 4E5 K lZ9n31p139DN 4ZADFKBP12 45 l95B l1 B1p1mI39n5. Il 5H9CD5 35p5n41nD 45C pl1nD5C DB1nC7Nn9AE5C oT lZ5HpB5CC9on 45 )%R 5CD ND59nD5 p1B lZ5HpB5CC9on CD12l5 4ZEn5 3onCDBE3D9on 5n Np9n7l5 K 385F5EH 392l1nD l5 DB1nC3B9D )%R, 5D 45C l97nN5C l1 CEB5HpB9m1nD (D5pBoCD et al., 2007). (EB l1 21C5 7NnND9AE5 45 lZ9nC5nC929l9DN 45 )%R K l1 B1p1mI39n5 1 NDN 3on4E9D5 En5 NDE45 CZ9nDNB5CC1nD 1E DB1nC3B9pDom5 45C pl1nD5C 5HpB9m1nD FKBP12 45 l5FEB5, 1EDoB9C1nD 19nC9 lZ9n8929D9on 45 )%R F91 35D 9n8929D5EB p81Bm13olo79AE5. In8925B l1 Fo95 )%R BNCElD5 5n En5 BN4E3D9on 7NnNB1l5 45 l1 3Bo9CC1n35 F91 En5 BNpB5CC9on DB1nC3B9pD9onn5ll5 45 nom2B5EH 7On5C 9mpl9AENC 41nC 45C pBo35CCEC 1n12ol9AE5C 3omm5 l1 p8oDoCInD8OC5. Il 5CD 1ECC9 DBOC 9nDNB5CC1nD 4Zo2C5BF5B AEZ5n p1B1llOl5 45 Con 9mpl931D9on 1DD5n4E5 41nC l5 mND12ol9Cm5 pB9m19B5, lZ9n8929D9on 45 )%R pBoFoAE5
15
lZ9n4E3D9on 45 7On5C 3onDBSl1nD l5 31D12ol9Cm5, l1 pBo4E3D9on 45 mND12ol9D5C C53on419B5C 5D 45 pBoDN9n5C 1pp1BD5n1nD K l1 3l1CC5 45C 4N65nC9n5C, 35C 45EH 45Bn9OB5C 9mpl9AEN5C 41nC l1 BNponC5 1E CDB5CC. DZEn5 m1n9OB5 7NnNB1l5, 35DD5 NDE45 monDB5 AE5 lZ9n8929D9on 45 )%R 9n8925 l5C pBo35CCEC 1n12ol9AE5C, 3onComm1nD 45 lZNn5B795 poEB pBo4E9B5 45 l1 29om1CC5, D5l l1 p8oDoCInD8OC5, 5D 13D9F5 l5C pBo35CCEC 31D12ol9AE5 pBo4E3D5EBC 4ZNn5B795, 3omm5 l1 7lI3olIC5 (R5n et al., 2012). DZ1EDB5C NDE45C m5DD5nD 5n B5l956 l5C mo496931D9onC DB1nC3B9pD9onn5ll5C 9n4E9D5 p1B l1 Fo95 )%R F91 Con 13D9F1D9on p1B l5 7lE3oC5 (-9on7 et al., 2013; -9on7 1n4 (855n, 2012). C5p5n41nD, 35 CICDOm5 o2C5BF5 4ZEn5 m1n9OB5 7NnNB1l5 lZ9nF5BC5 45C mo496931D9onC DB1nC3B9pD9onn5ll5C o2C5BFN5C p1B R5n et al. (2012). NoD1mm5nD, lZ13D9F1D9on 45 )%R p1B lZ1:oED 45 7lE3oC5 pBoFoAE5 939 En5 9n4E3D9on DB1nC3B9pD9onn5ll5 45 7On5C 9mpl9AENC 41nC 45C pBo35CCEC 31D12ol9AE5C 3omm5 l1 7lI3olIC5 (-9on7 et al., 2013). %n p5ED CEppoC5B AE5 lZ13D9F1D9on 45 )%R p1B l5 7lE3oC5 5CD 251E3oEp mo9nC CpN3969AE5 AE5 lZ9n8929D9on 45 35ll5-39 p1B l1 B1p1mI39n5 loBCAE5 l5 CICDOm5 l5 p5Bm5D. D5 plEC, R5n et
al. (2012) nZo2C5BF5nD 1E3En 381n75m5nD DB1nC3B9pD9onn5l oE p8NnoDIp9AE5 41nC l5C
pl1nD5C ,) DB19DN5C K l1 B1p1mI39n5, mPm5 K DBOC 6oBD5 3on35nDB1D9on, 1loBC AE5 35D 9n8929D5EB C5m2l5 6on3D9onn5B C1nC l1 pBNC5n35 4ZFKBP12 41nC l5 CICDOm5 45 -9on7
et al. (2013).
1.2.3. Exem2.es de stratégies infectieuses basées sur .a mani2u.ati1n de .’initiati1n de .a traducti1n.
L5 619D AE5 35BD19nC p1D8o7On5C 195nD 3omm5 CDB1DN795 4Z9n653D9on 45 m1n9pEl5B l1 m1389n5B95 4Z9n9D91D9on 45 l1 DB14E3D9on CoEl97n5 lZ9mpoBD1n35 45 35 mN31n9Cm5 41nC lZ141pD1D9on molN3El19B5 1E CDB5CC. Il 1 p1B 5H5mpl5 NDN monDBN AE5 l5 pBoDoJo19B5 5D p1B1C9D5 o2l971Do9B5 45C m13Bop8175C 45 mo5ll5 Np9n9OB5, Leishmania major, 9n4E9C19D
16
En5 5HD9n3D9on 7NnNB1l5 45 l1 DB14E3D9on F91 l5 3l9F175 45 m)%R poEB p5Bm5DDB5 C1 CEBF95 5D C1 mElD9pl931D9on ( 1B1m9llo et al., 2011). En 35 AE9 3on35Bn5 En 5H5mpl5 213DNB95n, 9l 5CD 3onnE AE5 Pseudomonas entomophila 9n8925 1ECC9 l1 Fo95 )%R poEB C1 CEBF95 (Mo8B 1n4 (on5n25B7, 2012). L5 31C 45C F9BEC 5CD pBo212l5m5nD lZEn 45C plEC 4o3Em5nDN, 31B nZND1nD p1C 45C 35llEl5C, 9lC ConD 9nDB9nCOAE5m5nD 4Np5n41nDC 45 l1 m1389n5B95 45 DB14E3D9on 45 l1 35llEl5 8SD5. D1nC l5 31C 4E F9BEC 45 lZ8Np1D9D5 C (HC+), l1 4ND53D9on 45 lZARN 4oE2l5-2B9n 7NnNBN p5n41nD l5 3I3l5 45 BNpl931D9on F9B1l, p1B l1 PKR 9n4E9D l1 p8oCp8oBIl1D9on 45 5IF2α, 5D ND59nD 45 m1n9OB5 7NnNB1l5 l1 DB14E3D9on 45C ARNm 8SD5C. %B, on DBoEF5 En5 CNAE5n35 IRE( 41nC l5 5Z*)R 45 lZARNm F9B1l, B53BED1nD 49B53D5m5nD l1 pBoDN9n5 5IF2A AE9 31D1lIC5 lZ1n3B175 45 lZARND 9n9D91D5EB CEB l1 p5D9D5 CoEC-En9DN 4ZEn5 m1n9OB5 A*G-4Np5n41nD5. L1 DB14E3D9on 45 lZARNm F9B1l 5CD 41nC 35 31C 9n4Np5n41nD5 45 l1 49Cpon929l9DN 5n 59F2α 13D96 (K9m et
al., 2011). L1 l9DDNB1DEB5 CEB l1 F9Bolo795 FN7ND1l5 6oEBm9ll5 4Z5H5mpl5C 4NmonDB1nD
lZ9mpl931D9on 45 l1 DB14E3D9on 3omm5 613D5EB m1:5EB 9n6lE5nM1nD l5 CE33OC 45 lZ9mmEn9DN. *n nom2B5 7B1n49CC1nD 45 7On5C 45 BNC9CD1n35 BN35CC96C, oE 7On5C 45 CEC35pD929l9DN (7On5C () ConD 945nD969NC. D5 p1B l5 619D AE5 45C p1B1C9D5C o2l971Do9B5C, D5lC l5C F9BEC, C5 B5poC5nD 5n DoD1l9DN CEB l1 m1389n5B95 45 DB14E3D9on 45 lZ8SD5 poEB 5HpB9m5B l5EB nom2B5 l9m9DN 45 pBoDN9n5C, 9l 1pp1B1îD lo79AE5 AEZEn5 7B1n45 p1BD95 45C 7On5C ( 3o45nD poEB 5IF4E 5D 4G, oE l5EBC 9Co6oBm5C (F1n (3895 1n4 )1;;5n, 2014; ,1n7 1n4 KB9C8n1CG1mI, 2012). L5C 31C 45C Pot viruses 3omm5 l5 Tobacco etch
virus ()E+), l5 Turnip mosaic virus ()EM+) 5D l5 Clover ellow vein virus (Cl.++) ConD
pBo212l5m5nD l5C plEC 4o3Em5nDNC CEB l5 CE:5D. L5C F9BEC 45 35DD5 61m9ll5 5n3o45nD l1 pBoDN9n5 F9B1l5 +P7 (+9B1l PBoD59n 7) 4N3oEF5BD5 385J l5 )E+, AE9 l95 45 m1n9OB5 3oF1l5nD5 lZARN F9B1l 7Nnom9AE5 (MEBp8I et al., 1990). Il p1B l1 CE9D5 NDN monDBN AE5 l1 pBoDN9n5 +P7 45 )EM+ 9nD5B179D 1F53 5IF(9Co)4E 5D AE5 35DD5 9nD5B13D9on 5CD nN35CC19B5 K l1 mElD9pl931D9on 4E F9BEC 385J Arabidopsis (DEpB1D et al., 2002; L5on1B4
et al., 2000). En 3onDB1CD5, l5 F9BEC Cl.++ 1 25Co9n 45 5IF4E m19C p1C 45 (9Co)4E
17
35BD19nC 5IF 45 lZ8SD5 p1B l5 +P7 45 F9BEC 4966NB5nDC. D5 plEC, )EM+ n5 p5ED plEC 9n653D5B 45C Arabidopsis mED1nD5C poEB l5C 45EH 9Co6oBm5C 45 5IF(9Co)4G 1loBC AE5 lZ9n653D9on p1B Cl.++ 4Np5n4 45 5IF4G (N9319C5 et al., 2007). A9nC9, 1loBC AE5 )EM+ 1pp1B1îD 3omm5 4Np5n41nD 4E 3ompl5H5 5IF(9Co)4F poEB C1 mElD9pl931D9on, Cl.++ 1pp1B1îD 1E 3onDB19B5 3omm5 4Np5n41nD 4E 3ompl5H5 5IF4F.
1.3. Les s:stèmes immunitaires basés sur .es mécanismes 2r1téi3ues.
L5 CICDOm5 9mmEn9D19B5 45C pl1nD5C 5CD 7NnNB1l5m5nD CE249F9CN 5n 45EH 3oE385C m1:5EB5C : En5 5HDB135llEl19B5 1CCEBN5 p1B 45C BN35pD5EBC K B53onn19CC1n35 45 moD96 (PRRC ; P1DD5Bn B53o7n9D9on B535pDoBC) B53onn19CC1nD l5C moD96C molN3El19B5C 1CCo39NC 1EH p1D8o7On5C (PAMPC ; P1D8o75n-1CCo391D54 mol53El1B p1DD5BnC), 5D En5 9nDB135llEl19B5 1CCEBN5 p1B l5C pBoDN9n5C NB-LRR B53onn19CC1nD l5C pBoDN9n5C 9n:53DN5C oE pBo4E9D5C p1B l5 p1D8o7On5 41nC l5 3IDopl1Cm5, 1pp5lN5C 56653D5EBC. LZ13D9F1D9on 45C PRRC 9n4E9D lZ9mmEn9DN 4N3l5n38N5 p1B l5C PAMPC (P)I ; PAMP-DB9775B54 9mmEn9DI) 1loBC AE5 lZ13D9F1D9on 45C NB-LRR 9n4E9D lZ9mmEn9DN 4N3l5n38N5 p1B l5C 56653D5EBC (E)I ; E6653DoB-DB9775B54 9mmEn9DI) (Do44C 1n4 R1D8:5n, 2010).
1.3.1. Descri2ti1n intégrative du s:stème immunitaire végéta..
En BNponC5 K l1 4ND53D9on 4ZEn p1D8o7On5, 381AE5 35llEl5 FN7ND1l5 oB385CDB5 C1 pBopB5 4N65nC5 1F1nD 45 m5DDB5 5n pl135 En 491lo7E5 1F53 C5C Fo9C9n5C. L1 m9C5 5n pl135 45C 4N65nC5C 9mmEn9D19B5C o2l975 l1 35llEl5 K 5HpB9m5B En NF5nD19l 45 7On5C 1CCo39NC K lZ141pD1D9on 4E p1D8o7On5 CpN3969AE5m5nD 4ND53DN. A 35 CE:5D, l5C pl1nD5C
18
n5 49CpoC5nD p1C 45 CICDOm5 45 CEBF59ll1n35 mo29l5 3omm5 l5C m1mm96OB5C, m19C plEDSD 4ZEn CICDOm5 45 4ND53D9on 45 DIp5 35llEl5 1EDonom5. C813En5 45C 45EH 3oE385C 4ND53D5 45C C97n1EH p1BD93El95BC. L1 plEC 31B13DNB9CN5 K 35 :oEB 3on35Bn5 l1 B53onn19CC1n35 45C moD96C molN3El19B5C 3onC5BFNC 5D 1CCo39NC 1EH m93Bo25C (5H. 389D9n5, 6l175ll9n5, LP( : PAMPC oE MAMPC). L1 B53onn19CC1n35 45 35C moD96C 5CD 1CCEBN5 p1B l5C pBoDN9n5C PRR AE9 ConD 45C BN35pD5EBC 45 DIp5 ;9n1C5 p5Bm5DD1nD lZ13D9F1D9on 45C 4N65nC5C 4N3l5n38N5C p1B l5C PAMPC : P)I (oE M)I). L1 4ND53D9on 45 nZ9mpoBD5 AE5l DIp5 45 p1D8o7On5 p1B l1 35llEl5 FN7ND1l5 p1CC5 p1B 35 mN31n9Cm5, 5D 5CD 7NnNB1l5m5nD CE669C1nD5 K l1 B5CDB93D9on 45 l1 3Bo9CC1n35 m93Bo295nn5. %B, l5C p1D8o7On5C 141pDNC K l5EB(C) 8SD5(C) 1E 3oEBC 45 lZNFolED9on onD 14opDN 45C moI5nC 45 CEBp1CC5B 35DD5 pB5m9OB5 21BB9OB5 F91 l1 4Nl9FB1n35, K lZ9nDNB95EB 45 l1 35llEl5 8SD5, 45 pBoDN9n5C 56653DB935C 1I1nD poEB 6on3D9on 4Z1E7m5nD5B l1 49CCNm9n1D9on 4E m93Bo25 41nC lZoB71n9Cm5 5n CEppB9m1nD noD1mm5nD l1 P)I (/81n7 et al., 2012). L5C 56653D5EBC m93Bo295nC poBD5nD plEC95EBC nomC CE9F1nD l5 3onD5HD5, 3omm5 pBoDN9n5 4Z1F9BEl5n35 (AFB), 56653D5EB 45 DIp5 DBo9C (E))) oE DoED C9mpl5m5nD 56653D5EB. L5C plEC 31B13DNB9CNC 3on35Bn5nD Pseudomonas s ringae pF. Tomato (Pst DC3000), 31EC1nD 45C CImpDSm5C 45 nN3BoC5C CEB l5C pl1nD5C mo4Ol5C Arabidopsis 5D l1 Dom1D5. Comm5 En 7B1n4 nom2B5 45 213DNB95C p8IDop1D8o7On5C, 35DD5 CoE385 ED9l9C5 En (ICDOm5 45 (N3BND9on 45 DIp5 III ((())) 45 n1DEB5 mElD9pBoDN9AE5 poEB 4Nl9FB5B 1E DB1F5BC 45 l1 m5m2B1n5 FN7ND1l5 l5C 56653D5EBC 213DNB95nC, 41nC 35 31C, 45 DIp5 III. L1 pl1nD5 poCCO45 45C 7On5C 45 RNC9CD1n35 5n3o41nD 45C pBoDN9n5C R NB-LRR 31p12l5C 45 B53onn1îDB5 l1 pBNC5n35 4ZEn 56653D5EB 4onnN 5D 45 4N3l5n385B 45C Fo95C 45 C97n1l9C1D9on m5n1nD K lZ9mmEn9DN. (38Nm1D9AE5m5nD, En5 45C 3oE385C 5CD 381B7N5 45 B53onn19DB5 l1 n1DEB5 45 lZ5nF189CC5EB (P)I) p5n41nD AE5 lZ1EDB5 4ND53D5 C5C 1Bm5C (Do44C 1n4 R1D8:5n, 2010) (697. 6). En BNponC5 K 35C mN31n9Cm5C 45 B53onn19CC1n35, l1 35llEl5 poCCO45 plEC95EBC opD9onC CEB l5 38o9H 4E n9F51E oT 4o9D CZopNB5B l1 BN7El1D9on. L1 m95EH 31B13DNB9CN5 K 35 :oEB 5CD 35BD19n5m5nD l1 BN7El1D9on DB1nC3B9pD9onn5ll5. C5p5n41nD, 45C NF945n35C 29o389m9AE5C CE77OB5nD AE5 l1
20
p1D8o7On5 F91 l1 B53onn19CC1n35 45C MAMPC p1B l5C PRR. DZEn5 m1n9OB5 7NnNB1l5, l5C PRR poCCO45nD En 4om19n5 5HDB135llEl19B5 45 DIp5 L5E39n5-R938 R5p51D 5D En 4om19n5 9nDB135llEl19B5 45 DIp5 CNB9n5/D8BNon9n5 ;9n1C5. L1 B53onn19CC1n35 4E MAMPC p1B l1 PRR 1ppBopB9N5 5nDB19n5 CoEF5nD Con 1CCo391D9on 1F53 En BN7El1D5EB 35nDB1l 45 l1 BNponC5 9mmEn9D19B5, BAK1 (BB1CC9noCD5Bo94-A3D9F1D54 K9n1C5). C5DD5 9nD5B13D9on 13D9F5 p1B p8oCp8oBIl1D9on l5C 45EH BN35pD5EBC, BNCElD1nD 5n lZ13D9F1D9on 45 31C3145 45 ;9n1C5C CD9mEl1nD 1E n9F51E DB1nC3B9pD9onn5l 45C 7On5C 1CCo39NC K l1 pBo4E3D9on 45 2oE38onC 45 31lloC5 F5n1nD B5n6oB35B l5C m5m2B1n5C 5D 3olm1D5B l5 3onD9nEEm CImpl1Cm9AE5, 45 pBoDN9n5C 45 4N65nC5 21C1l5 3omm5 45C 389D9n1C5C, 5n4o7lE31n1C5C, 45 molN3El5C 1nD9m93Bo295nn5C 3omm5 l5C p8IDo1l5H9n5C AE9 ConD 45C mND12ol9D5C C53on419B5C. C5p5n41nD, l5C p1D8o7On5C 141pDNC K l5EBC 8SD5C onD 4NF5loppN 45C CDB1DN795C 45 4Nl9FB1n35 9nDB135llEl19B5 45 molN3El5C 1pp5lN5C 56653D5EBC, poEF1nD 3oEBD-39B3E9D5B l1 M)I (5D 4Z1EDB5C pBo35CCEC p8IC9olo79AE5C). L5 mo45 45 4Nl9FB1n35 45C 56653D5EBC 5CD 4Np5n41nD 4E p1D8o7On5, l5C 381mp97nonC FonD p1B 5H5mpl5 7NnNB5B En 81ECDoB9Em F5n1nD 619B5 pB5CC9on 3onDB5 l1 35llEl5 5D pBoFoAE5B lZ9nF179n1D9on 45 l1 m5m2B1n5 C1nC :1m19C pNnNDB5B lZ5Cp135 9nDB135llEl19B5. L5C 213DNB95C 45 pl1nD5C onD, AE1nD K 5ll5C, m9C 5n pl135 En CICDOm5 4Z W 9n:53D9on X DB1nCm5m2B1n19B5 45C 56653D5EBC. D1nC 35C 45EH 31C, l1 B53onn19CC1n35 49B53D5 oE 9n49B53D5 45 lZ56653D5EB p1B l1 pBoDN9n5 45 BNC9CD1n35 NB-LRR 3oBB5Cpon41nD5 F1 p5Bm5DDB5 l1 m9C5 5n pl135 45 lZE)I poEF1nD 3Elm9n5B CoEC l1 6oBm5 4ZEn5 moBD 35llEl19B5 pBo7B1mmN5 5D lo31l9CN5 1E po9nD 4Z9n653D9on : l1 BN13D9on 8Ip5BC5nC92l5 (Do44C 1n4 R1D8:5n, 2010).
1.3.2. La 2remière .igne de défense : .’immunité déc.enchée 2ar .es m1tifs m1.écu.aires ass1ciés au 2ath1gène.
21
L1 pB5m9OB5 3oE385, oE 9mmEn9DN 4N3l5n38N5 p1B l5C PAMPC (P)I), 5CD 381B7N5 45 B53onn1îDB5 l5C moD96C molN3El19B5C 1CCo39NC 1EH m93Bo25C p1B l5C BN35pD5EBC K B53onn19CC1n35 45 moD96C (PRR). C5DD5 B53onn19CC1n35 5CD l1 pB5m9OB5 ND1p5 45 4ND53D9on 4E p1D8o7On5 pE9CAEZ5ll5 C5 4N3l5n385 1E 3onD13D 45 l1 35llEl5, 1pBOC l1 pNnNDB1D9on 45 35lE9-39 F91 45C 2l5CCEB5C oE l5C CDom1D5C. L5C PRR 3onnE5C 6onD p1BD95C 45 45EH 3l1CC5C : 35ll5 45C BN35pD5EBC DB1nCm5m2B1n19B5C K 13D9F9DN ;9n1C5 oE 35ll5 45C pBoDN9n5C DB1nCm5m2B1n19B5C 45 DIp5 BN35pD5EB. L5 mo4Ol5 l5 m95EH 31B13DNB9CN 385J l5C pl1nD5C 5CD 35BD19n5m5nD l1 B53onn19CC1n35 4E p5pD945 213DNB95n 6l175ll9n5 22 (6l722) p1B l1 PRR Fl175ll9n C5nC9n7 2 (FL(2). L5 BN35pD5EB 9nD5B179D 49B53D5m5nD 1F53 l5 p5pD945 4N3l5n381nD l1 6oBm1D9on 4ZEn 3ompl5H5 45 C97n1l9C1D9on 3onD5n1nD l1 pBoDN9n5 ;9n1C5 BB1CC9noCD5Bo94-ACCo391D54 K9n1C5 1 (BAK1). LZ9nD5B13D9on FL(2-BAK1 p5Bm5D l5EB p8oCp8oBIl1D9on B5Cp53D9F5 pE9C l5EB 13D9F1D9on. LZ13D9F9DN ;9n1C5 45 35C 45EH pBoDN9n5C p5Bm5D lZ13D9F1D9on 45 plEC95EBC Fo95C D5ll5C AE5 l5C Fo95C MAPK (M9Do75n-ACCo391D54 PBoD59n K9n1C5) oE CDPK (C1l39Em-D5p5n41nD PBoD59n K9n1C5) (M5n7 1n4 /81n7, 2013). En 1F1l 45 35C Fo95C ;9n1C5C ConD BN7ElNC 45C 613D5EBC 45 DB1nC3B9pD9on poEF1nD CD9mEl5B lZ5HpB5CC9on 45 7On5C B5l9NC K l1 p1D8o7NnOC5 (7On5C PR) 5D 3o41nD poEB 45C pBoDN9n5C 45 4N65nC5 21C1l5 D5ll5C 45C 389D9n1C5C, 45C 5n4o7lE31n1C5C, oE CD9mEl5B l1 29oCInD8OC5 45 molN3El5C 1nD9m93Bo295nn5C D5ll5C l5C p8IDo1l5H9n5C (Do44C 1n4 R1D8:5n, 2010). D1nC l1 m1:oB9DN 45C 31C, l1 P)I 5CD CE669C1nD5 poEB 3onDB5B DoED5 D5nD1D9F5 4Z9nF1C9on p1B En m93BooB71n9Cm5 non p1D8o7On5 CEB lZ8SD5 38o9C9. En 35 C5nC, 5ll5 5CD 1pp5lN5 4N65nC5 21C1l5. C5p5n41nD, l5C 213DNB95C p1D8o7On5C 141pDNC K l5EBC 8SD5C onD 4NF5loppN 45C CDB1DN795C 45 4Nl9FB1n35 9nDB135llEl19B5 45 molN3El5C 56653DB935C (l1 m1:oB9DN ConD 45C pBoDN9n5C), poEF1nD 9nD5B6NB5B 1F53 l1 P)I 5D p5Bm5DDB5 l1 m9C5 5n pl135 45C 3on49D9onC 61FoB12l5C K l1 mElD9pl931D9on 5D K l1 49Cp5BC9on 4E p1D8o7On5 41nC lZoB71n9Cm5 8SD5. *n 45 35C 56653D5EBC 5CD Hop*1 5D 392l5 45C RBPC 3omm5 GRP7. GRP7 9nD5B179D 3onCD9DED9F5m5nD 1F53 5IF4E 5D lZARNm 45 FL(2, 5D 5CD nN35CC19B5 K l1 P)I. L5 392l175 45 GRP7 p1B Hop*1 2loAE5 lZ9nD5B13D9on 45 35ll5-39
23
O? 5:DE:?8F6 56FI 5om2:?6D AC:?4:A2FI : l2 3oSE6 56 l:2:Do? 2G64 l6D ?F4lPoE:56D, 7oCmP6 A2C l6D 5om2:?6D %B (%F4l6oE:56 B:?5:?8) 6E A(C (AA27-1, ( ACoE6:?, CED-4) 1 6E 2 ; l6 5om2:?6 L(( (L6F4:?6-(:49 (6A62E) :?E6CG6?2?E 52?D l6D :?E6C24E:o?D :?EC2 6E :?E6CmolP4Fl2:C6D. L6 5om2:?6 1) 6DE ECèD 4o?D6CGP E2?5:D BF6 l6 5om2:?6 2) 6DE G2C:23l6. AFDD:, l’o? 5:DE:?8F6 56FI 8C2?56D 4l2DD6D 56 %B-L(( DF:G2?E l2 ?2EFC6 56 l6FC 5om2:?6 ACPD6?E 6? %-E6Cm:?2l. L6 AC6m:6C EJA6 AoDDè56 F? 5om2:?6 T ( (Toll 2?5 ?E6Cl6Fk:?-1 (646AEoC 9omolo8J) 2loCD BF6 l6 D64o?5 AoDDè56 F? 5om2:?6 ACP5:E AoFC 7oCm6C F? 5om2:?6 4o:l65-4o:l (CC) :mAl:BFP 52?D l2 D:8?2l:D2E:o? G:2 l6D :?E6C24E:o?D ACoEP:?6D-ACoEP:?6D. CFC:6FD6m6?E, l6D ETT 6E EoFD l6D 67764E6FCD o?E lo?8F6m6?E 6E 8P?PC2l6m6?E PEP 2AA6lPD l6D ACoEP:?6D 5’2G:CFl6?46 (AGC). E? 6776E, 6ll6D Do?E PEF5:P6D AlFETE AoFC l6D Go:6D 56 5P76?D6D BF’6ll6D Pl:4:E6?E BF6 AoFC l6FC CP6ll6 7o?4E:o? 56 G:CFl6?46. AFDD:, 56D 6I6mAl6D 5’:?E6C24E:o?D 5:C64E6D 6?EC6 AGC 6E %B-L(( o?E PEP 5P4C:ED M 46 ;oFC. D6 ?om3C6FI mo5èl6D C6l2E6?E AlFETE l2 A6C46AE:o?, A2C l6D %B-L((, 56 l2 mo5:7:42E:o? 56D 4:3l6D 56D AGC. D6 46 72:E, l6 CTl6 56 46CE2:?6D 4:3l6D 56D ETT 6DE 4o??F 6E 3:6? 42C24EPC:DP 2F ?:G62F molP4Fl2:C6 52?D l2 C64o??2:DD2?46 A2C l6D %B-L((, m2:D l6FC CP6ll6 7o?4E:o? 2F ?:G62F 56 l2 A9JD:olo8:6 56 l2 46llFl6-9TE6 C6DE6 M ACoFG6C.
D2?D l2 m2;oC:EP 56D 42D, l2 A6C46AE:o? 5’F? AGC A2C l2 %B-L(( 4oCC6DAo?52?E6 4o?5F:E M l2 CPAo?D6 9JA6CD6?D:3l6 (H(), F?6 moCE 46llFl2:C6 ACo8C2mmP6 6E lo42l:DP6 2F Ao:?E 5’:?764E:o?. (B6C?oFI et al., 2011). C6A6?52?E, l‘2AA2C:E:o? 56 l2 H( ?’6DE A2D EoF;oFCD 2DDo4:P6 M l’24E:G2E:o? 5’F?6 %B-L(( 4o?5F:D2?E M l’:mmF?:EP. C’6DE DoFG6?E l6 42D 56D G:CFD (Mo776EE, 2009).
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+? 56D 6I6mAl6D 4o??FD 6DE l2 C64o??2:DD2?46 56D AGCL 5F 492mA:8?o? 56 l2 CoF:ll6 5F l:?, A2C l6D %B-L(( L. D2?D F? 2CE:4l6, 5oFK6 G2C:2?ED 5F lo4FD AGCL567 56 D:I DoF496D 56 CoF:ll6 5:77PC6?E6D Do?E :DolPD 6E mo?EC6?E F? 46CE2:? 568CP 56 AolJmoCA9:Dm6, 2F m2I:mFm 20% 56 5:77PC6?46 6? 24:56D 2m:?PD. L2 CoF:ll6 5F l:? PE2?E F? A2C2D:E6 o3l:82Eo:C6 ?’:?764E2?E BF6 46EE6 Al2?E6, 46 92FE 568CP 56 AolJmoCA9:Dm6 6DE DFAAoDP AC:moC5:2l AoFC l6 m2:?E:6? 56 l2 G:CFl6?46 DFC 46ll6-4:, 3:6? BF’2F4F?6 7o?4E:o? 56 G:CFl6?46 ?6 Do:E 4o??F6 AoFC AGCL567. DF Ao:?E 56 GF6 GP8PE2l, l2 mRm6 49oD6 6DE GC2:6, AF:DBF6 l2 AC6DD:o? 56 DPl64E:o? G2 72GoC:D6C l2 4o?D6CG2E:o? 56 ACoEP:?6D ( 42A23l6D 56 C64o??2SEC6 DAP4:7:BF6m6?E F? G2C:2?E 5’F? 2FEC6. E? 6776E, l6 l:? 6?4o56 AlFD:6FCD 2llèl6D 56 l2 %B-L(( L, ?oE2mm6?E L5, L6 6E L7. L6D CPDFlE2ED mo?EC6?E BF6 D6Fl6m6?E D6AE 56D 5oFK6 AGCL ACo5F:D6?E 56D DJmAETm6D 56 H( loCDBF6 DFC6IAC:mPD 52?D F? 8P?oEJA6 4o?E6?2?E l2 %B-L(( L 4oCC6DAo?52?E6, 46 BF: G2 52?D l6 D6?D 5’F?6 DAP4:7:42E:o? 56 46CE2:?6D %B-L(( L AoFC F? G2C:2?E A2CE:4Fl:6C. AFDD:, 46 8CoFA6 ACoFG6 l’:?E6C24E:o? 5:C64E6 6?EC6 AGCL567 6E L5/6 A2C F? DJDEèm6 56 5oF3l6 9J3C:56 52?D l2 l6GFC6. L6D 2FE6FCD ?‘6I4lF6?E A2D l’:?E6CG6?E:o? 5’2FEC6D ACoEP:?6D 52?D 46EE6 :?E6C24E:o? in planta, m2:D l2 4o?D:5èC6?E ?o? 6DD6?E:6ll6. E? CPDFmP, l6D 2FE6FCD ACPD6?E6?E F? mo5èl6 AlFETE PlP82?E 42C 5PA6?52?E 5F mo56 5’:?764E:o? 56 l2 CoF:ll6 (A2C2D:E6 o3l:82Eo:C6), 52?D l6BF6l l2 AC6DD:o? 56 DPl64E:o? 72GoC:D6 l6D mFE2E:o?D A6Cm6EE2?E l’:?E6C24E:o? 5:C64E 6?EC6 ( 6E AGC D2?D AC6?5C6 6? 4omAE6 l2 7o?4E:o? 56 G:CFl6?46 56 l’67764E6FC (Do55D et al., 2006).
1.3.3.2. Exemple ’in-erac-ion in irec-e en-re A/r e- NB-LRR.
E? 72:E, l2 m2;oC:EP 56D 4oFAl6D AGC/%B-L(( 42C24EPC:DPD ?P46DD:E6?E l’:?E6CG6?E:o? 5’F? ECo:D:èm6 A2CE6?2:C6 : l6 4o724E6FC. L’6I6mAl6 l6 m:6FI 5o4Fm6?EP 24EF6ll6m6?E
25
4o?46C?6 l6 4:3l286 5’F? D6Fl 6E mRm6 4o724E6FC, ( %4 ((PM1- ?E6C24E:?8 ACoE6:?-4), A2C ECo:D 67764E6FCD 5:77PC6?ED : AGCB, AGC(AE2 oF AGC(Am1 :DDFD 56 P t DC3000. D6FI %B-L((D Do?E :mAl:BFP6D 52?D l2 DFCG6:ll2?46 5F DE2EFE 56 ( %4. ( %4 6DE F?6 A6E:E6 ACoEP:?6 56 211 24:56D 2m:?PD BF:, 2G64 l6D %B-L(( (PM1 ((6D:DE2?46 Eo P.
yringae AG. Maculicola) 6E (P)2 ((6D:DE2?46 Eo P. yringae), 6DE lo42l:DP6 M l2
m6m3C2?6 (G2o et al., 2011; M24k6J et al., 2003; M24k6J et al., 2002). L’:?E6C24E:o? 56 ( %4 2G64 (PM1 6E (P)2 6DE 5Pmo?ECP6 M 56 ?om3C6FD6D C6AC:D6D G:2 56D 6IAPC:6?46D 56 5oF3l6 9J3C:56 496K l2 l6GFC6 oF 56 4o:mmF?oACP4:A:E2E:o?. PC6m:èC6m6?E, (PM1 6DE 24E:GP6 A2C l’:?o4Fl2E:o? 56 324EPC:6D 6IAC:m2?E l6D ETT AGCB oF AGC(Am1. LoCDBF’:?ECo5F:ED 52?D l2 46llFl6-9TE6, 46D AGC :?E6C28:DD6?E 2G64 56D ACoEP:?6D k:?2D6D 4omm6 ( PK oF MAPK4, :lD Do?E 24PEJlPD AoFC A6Cm6EEC6 l6FC 25C6DD286 M l2 m6m3C2?6, 6E :?5F:D6?E l2 A9oDA9oCJl2E:o? 56 ( %4 (CF: et al., 2010; L:F et al., 2011). ( %4 6DE 4:3lP A2C F? ECo:D:èm6 ETT M 24E:G:EP 4JDEP:?6 ACoEP2D6, AGC(AE2. L’:?o4Fl2E:o? 56 324EPC:6D 6IAC:m2?E AvrRpt2 4o?5F:E C2A:56m6?E 2F 4l:G286 56 ( %4 6E CPDFlE6 6? l’24E:G2E:o? 56 (P)2 (AIE6ll et al., 2001; D2J et al., 2005; K:m et
al., 20052). A:?D:, l6 DE2EFE 56 ( %4 6DE m2?:AFlP A2C ECo:D 67764E6FCD 5:77PC6?ED 6E 46
DE2EFE 6DE 82C5P A2C 56FI %B-L(( 5:77PC6?E6D AoFG2?E 5P4l6?496C l2 ET (7:8. 8). L’6?D6m3l6 56 46D EC2G2FI DoFl:8?6 2FDD: l6 72:E BF6 l6D mFE2?ED KO AoFC ( %4 Do?E lPE2FI, 46 BF: 6DE lo8:BF6 D: l’o? 4o?D:5èC6 BF6 l2 Go:6 (P)2 6DE 4o?DE2mm6?E 24E:G6. E? 6776E, :l 2 FlEPC:6FC6m6?E PEP o3D6CGP 52?D F?6 l:8?P6 DFC6IAC:m2?E ( %4, BF6 l6D 5PATED 56 42lloD6 PE2:6?E ECèD CP5F:ED 2ACèD EC2:E6m6?E 2G64 56D MAMPD, A2C C2AAoCE 2F EJA6 D2FG286. AFDD:, l2 DFC6IAC6DD:o? 56 ( %4 A6Cm6E M F?6 DoF496 mFE2?E6 56 DC3000 5P7:4:6?E6 52?D Do? ))TT 56 D6 mFlE:Al:6C 100 7o:D AlFD BF6 DFC 56D Al2?E6D 56 EJA6 D2FG286 (K:m et al., 20053). E? 5PA:E 56 Do? :mAoCE2?46 52?D l’PE23l:DD6m6?E 56D 5P76?D6D, ( %4 ?’6DE A2D :?5:DA6?D23l6 2FI 7o?4E:o?D 56 G:CFl6?46 6I6C4P6D A2C AGCB, AGC(PM1 6E AGC(AE2 52?D 56D Al2?E6D DFD46AE:3l6D KO AoFC rin4 rpm1 rp 2, DF88PC2?E l’6I:DE6?46 5’2FEC6D 4:3l6D (B6lk925:C et al., 2004). L2 AC6m:èC6 9JAoE9èD6 56 46EE6 :?E6C24E:o? EC:A2CE:E6 2 PEP BF6 l6D %B-L(( 82C56?E 24E:G6m6?E l6D 4:3l6D 56
27
( %4 CPAC:m6 4o?DE:EFE:G6m6?E l2 MT . L2 D6Fl6 6IAl:42E:o? 4o??F6 M 46 ;oFC D6C2:E BF6 ( %4 CP8Fl6 l’oFG6CEFC6 DEom2E2l6 : 56D Al2?E6D KO AoFC ( %4 ?6 A6FG6?E A2D oFGC:C l6FCD DEom2E6D 6? CPAo?D6 M P t (L:F et al., 2009). D’F? Ao:?E 56 GF6 :mmF?:E2:C6, ( %4 7oCm6 F? 4omAl6I6 4o?DE:EFE:7 56 5P76?D6 2G64 l6D %B-L(( (PM1 6E (P)2. (PM1 6DE 24E:GP6 6? CPAo?D6 M l2 A9oDA9oCJl2E:o? :?5F:E6 A2C AGCB 6E AGC(PM1 E2?5:D BF6 (P)2 6DE 24E:GP6 6? CPAo?D6 2F 4l:G286 56 ( %4 A2C AGC(AE2. L2 A9oDA9oCJl2E:o? :?5F:E6 A2C AGCB ?P46DD:E6 ( PK in vivo 2loCD BF6 l2 A9oDA9oCJl2E:o? 56 ( %4 A2C MAPK4 ?’2 PEP 5Pmo?ECP6 BF’in vitro. L’24E:G2E:o? 56D 46D 56FI Go:6D 4o?5F:E M l’ET 6E M l2 CP24E:o? 56 H(.
1.3.3.2.1. 2mergence . mo 4le . le.rre.
L6 m2?BF6 56 AC6FG6D DFC l2 7o?4E:o? 56 ?om3C6FI 4o724E6FCD 2 A6Cm:D l’Pm6C86?46 5’F? ?oFG62F 4o?46AE 5’:?E6C24E:o? :?5:C64E6 AGC/%B-L((. +? 3o? 6I6mAl6 6DE l6 42D 56 PEo, F?6 DPC:?6/E9CPo?:?6 k:?2D6, BF: 2G64 l2 %B-L(( PC7, 4o?7èC6 l’:mmF?:EP M
P eudomona yringae 4o?E6?2?E l’ETT AGCPEo, 496K l2 Eom2E6. AF ?:G62F
molP4Fl2:C6, PEo 7oCm6 2G64 PC7 F? 4omAl6I6 4o?DE:EFE:7 2J2?E 4omm6 7o?4E:o? l2 C64o??2:DD2?46 56 l’:?9:3:E:o? 56 l’24E:G:EP k:?2D6 6I6C4P6 A2C AGCPEo DFC PEo. L2 A6CEFC32E:o? 5F 4omAl6I6 PEo-PC7 :?5F:E F?6 42D4256 56 D:8?2l:D2E:o?D m6?2?E M l2 H( 6E M F?6 5:m:?FE:o? 56 l2 mFlE:Al:42E:o? 324EPC:6??6 (MF4J? et al., 2006). +? 2? AlFD E2C5, F? C2AAoCE 5Pmo?EC6 BF6 AGCPEo :?E6C28:E 5:C64E6m6?E 2G64, 6E :?9:36 FL)2 6E EF(, 56FI P(( M 24E:G:EP DPC:?6/E9CPo?:?6 k:?2D6 AC:moC5:2l6 52?D l6 5P4l6?496m6?E 56 l2 MT (.:2?8 et al., 2008). AGCPEo :?E6C28:E 5o?4 2G64 ECo:D DPC:?6/E9CPo?:?6 k:?2D6D 5o?E D6Fl6m6?E 56FI Do?E 4o??F6D AoFC 2Go:C F?6 :mAoCE2?46 A9JD:olo8:BF6. M2:D 2loCD, PEo ?’6DE-6ll6 BF’F? l6FCC6 ? (2E:o??6ll6m6?E ?o?, o? ?6 A6FE A2D 6I4lFC6 BF’6ll6 2:E F?6 7o?4E:o? D:8?2lPE:BF6 2F ?:G62F :mmF?.
28
(P46mm6?E, F?6 PEF56 mo?EC6 DF44:?4E6m6?E BF’2ACèD l2 5PE64E:o? 5’AGCPEo A2C l6 4omAl6I6 PEo-PC7, PEo 56G:6?E 9JA6CA9oDA9oCJlP6 6E 464: 6DE ?P46DD2:C6 M l2 m:D6 6? Al246 56 l’ET . PEo D6C2:E AoE6?E:6ll6m6?E A9oDA9oCJlP6 6? tran A2C l’ol:8omPC:D2E:o? 2G64 F? 2FEC6 4omAl6I6 PEo-PC7 6E l6 5om2:?6 k:?2D6 56 PEo 6DE 7o?4E:o??6l in vivo (%EoFk2k:D et al., 2013). A:?D:, F? 4o724E6FC A6FE ECèD 3:6? C6ACPD6?E6C F? l6FCC6 4omm6 52?D l6 42D 56 PEo, BF: AoDDè56 F?6 8C2?56 9omolo8:6 56 DPBF6?46 2G64 l6D P(( 4:3lP6 A2C AGCPEo, 6E 6? mRm6 E6mAD AoDDè56 F?6 7o?4E:o? ?P46DD2:C6 M l’ET .
1.3.3.2.2. S1n-h4se es concep-s mo ernes s.r les in-erac-ions A/r/NB-LRR.
)F:G2?E l6 A2E9oDJDEèm6, o? A6FE DFAAoD6C BF6 5:77PC6?E6D AC6DD:o?D 56 DPl64E:o?D o?E 23oFE: M l’6I:DE6?46 56 AlFD:6FCD mo5èl6D 56 C64o??2:DD2?46 56D AGC A2C l6D %B-L((. AFEC6m6?E 5:E, o? A6FE DFAAoD6C BF6 l6D 5:77PC6?ED mP42?:Dm6D 5’:?E6C24E:o?D 6?EC6 46D 56FI 4l2DD6D 56 ACoEP:?6D (7:8. 9) CPDFlE6?E 56 mP42?:Dm6D 4oPGolFE:7D 5:77PC6?ED 6? 7o?4E:o? 5F A2E9o8è?6 6E 56 l’9TE6. AF 4o?EC2:C6 56D :?E6C24E:o?D 5:C64E6D, l6D mo5èl6D 5’:?E6C24E:o?D :?5:C64E6D m6EE6?E 6? C6l:67 l6D AC6DD:o?D 56 DPl64E:o? 6I6C4P6D DFC 56D A2E9oDJDEèm6D oU l6 DA64EC6 5’:?764E:o? 6E 5’9TE6 6DE l2C86. D2?D 46 86?C6 56 42D, o? 2DD:DE6 M F?6 GPC:E23l6 4oFCD6 M l’2Cm6m6?E 56 l2 Al2?E6 6E 5F A2E9o8è?6 : 1) l6D A2E9o8è?6D 252AEPD M l6FC(D) 9TE6(D) A6FG6?E DFAAC:m6C l6D 5P76?D6D 32D2l6D 6E 4o?E:?F6C M D6 mFlE:Al:6C ; 2) 6? CPAo?D6 M l2 DFAAC6DD:o? 56 l2 MT , l6D Al2?E6D o?E 24BF:D 2F 4oFCD 56 l’PGolFE:o? 56D mP42?:Dm6D 56 C64o??2:DD2?46 56 l2 mo5:7:42E:o? 56D 4:3l6D 6E 7:?2l6m6?E 3) o?E 25oAEP l2 m:D6 6? Al246 56 l6FCC6D ACPD6?E2?E 56 7oCE6D 9omolo8:6D 56 DPBF6?46 2G64 l6D 4:3l6D 56 G:CFl6?46 56D AGC.
30
1.3.3.3. Diff3rences en-re ETI e- HR.
L2 H( 2 lo?8E6mAD PEP A6?DP6 4omm6 :?5:DA6?D23l6 M l2 m:D6 6? Al246 56 l2 CPD:DE2?46. C’6DE 4omACP96?D:3l6 42C 6ll6 4o?DE:EF6 l6 D6Fl A9P?oEJA6 m24CoD4oA:BF6 5’:mmF?:EP GP8PE2l6 6E :l 6DE ECèD DP5F:D2?E 56 A6?D6C BF’F? mP42?:Dm6 :mmF?:E2:C6 :mAl:BF2?E F? 92C2-k:C: 46llFl2:C6 6DE 6DD6?E:6l AoFC D6 5P76?5C6 56D A2E9o8è?6D 3:oECoA96D BF: o?E 36Do:? 56 5PG:6C l6 mPE23ol:Dm6 56 l’9TE6 AlFETE BF6 5’:?5F:C6 D2 moCE. Comm6 52?D 46CE2:?6D 7oCm6D 56 moCED 46llFl2:C6D 2?:m2l6D, l2 H( GP8PE2l6 C6BF:6CE l’:mAl:42E:o? 56 mPE242DA2D6D. M2:D, 52?D F?6 Al2?E6 mFE2?E6 AoFC 46D mPE242DA2D6D, l2 H( 6DE 4omACom:D6 m2:D A2D l2 CPD:DE2?46 (Coll et al., 2010). AF 4o?EC2:C6, l2 ET ?P46DD:E6 AlFETE l’:mAl:42E:o? 56D ACoEP:?6D E6ll6D ED)1 (F2lk et al., 1999) oF 6?4oC6 %D(1 (C6?EFCJ et al., 1995). D’F?6 m2?:èC6 :?EPC6DD2?E6, 52?D l6 42D 56 l2 %B-L(( (PM1, Do? 24E:G2E:o? 4o?5F:E M l2 CPD:DE2?46 D2?D H( 6? 23D6?46 56 l2 mPE242DA2D6 1, 6E 4o?5F:E M l2 H( D2?D CPD:DE2?46 6? l’23D6?46 56 %D(1. O? A6FE 5o?4 4o?4lFC6 BF6 l6D %B-L((D 4o?ECTl6?E 56FI Go:6D 56 D:8?2l:D2E:o? 5:77PC6?E6D 2F ?:G62F 8P?PE:BF6. D’2FEC6D 42D 56 CPD:DE2?46 D2?D H( Do?E DoFG6?E 7oFC?:D A2C l6D %B-L(( 4o?EC6 l6D G:CFD 4omm6 (I (B6?529m2?6 et al., 1999), BF: :?5F:E l’6IECRm6 CPD:DE2?46.
1.3.3.4. Vers .ne -h3orie .nifica-rice es in-erac-ions en-re A/r e- NB-LRR ?
L6D 5:77PC6?ED 6I6mAl6D 5’:?E6C24E:o? m6?E:o??PD 4:-56DDFD ?’6IAl:BF6?E A2D, 2F ?:G62F molP4Fl2:C6, 56 BF6ll6 72Oo? F?6 %B-L(( 6DE 24E:GP6 ?: 4omm6?E 6ll6 EC2?Dm6E l6 D:8?2l 56 l2 A6C46AE:o? 5’F? AGC G6CD 46D 4:3l6D 6? 2G2l. D6 AlFD, l2 ?2EFC6 56 46 D:8?2l 6DE :?4o??F6. PoFC l’:?DE2?E, EoFD l6D CPDFlE2ED 5Pmo?EC6?E BF6 :) l6
