2 !

" # $ % & ' ( ) *

+ , ' - . ) / 0 # + - 1 2 ) 3 ! " , 4 5 ( % , 6 " 5 7 ! / & 5 8 # + , 4 9 " % # + , ( & ) * :
! / " # $ % & ' ( ) * +
! , - ( . / 0 1 2 3 4 ( 5
6 7 ( " 8 9 : 2 ; < = , > 1 ? @ = ' 0 2 ; A . 1 9 B C 1 2 - D = E F G +



1 . 1 2 H 4 B ) ( I 1 ? $ J K 6 ; & " ( L ( M 1 2 H N > 1 M ) 1 ? H B ! O 348 " H P " Q ( ! O
1 ? D = R M (
S H ; T 1 ? H B ! O 348 " H P " Q ( ! O 1 ? D H = R M " 3 H U 1 2 H % # U 18 - 327 2 - 3 H A E H A
1 2 4 ( 1 6 H H H A 1 ? H H H $ J 4 + 6 ; & " ( L ( M 1 2 H H H N > 1 M " - & V F ; K 1 W " H H H = ' H H H B 2 H H H - 3 X G Y
1 2 H A 1 Z H ; 0 [ \ H ] ^ 1 2 4 ( 1 6 H A 1 2 - 0 _ Z H 7 F ` B 1 ? D H = M ? H J A _ 9 H ; $ B I 1 a . 1 ! O
' # ( . O _ 9 H H H B 9 H H H G + [ / 0 S H H H b _ 2 H H H > c \ H H H G d + 1 2 H H H - 3 X G Y ' - $ J G K 1 2 4 ( 1 6 H H H A
1 ? ( L ( 6 H H H H G + / 0 1 2 H H H H 4 B ) ( I 1 2 H H H H ] e V H H H H = f _ ) H H H H g 1 W , H H H H h = 1 V H H H H # B i ' H H H H B 2 H H H H N > 1 M [
R 6 H H ; G ` B / 0 j H H < G k 1 W S ( 1 l _ I V H H = 1 6 0 m H H = R n 1 2 H H F 4 A " ( L ( M 1 2 H H N > 1 M
R V H H o Z H H ] ' H H F b 1 i 6 H H - J B . 1 W 6 H H = 1 p 1 2 H H - * B . E H H + R 1 2 H H F B ! 1 C R " H H B j H H = f 6 H H ; G g
1 2 H H H H - F B " H H H H U ' H H H H b 1 2 H H H H $ = / H H H H b [ R q r 1 1 V % K s H H H H = / H H H H B 1 2 H H H H - 3 X G Y t H H H H = 1 S H H H H + 6 ; &
V % ( E H u \ H G d + 1 2 H - 3 X G Y t H % + R 9 H $ B v ' H B 2 H - = 1 L 0 [ V H % # U 1 2 H ` G d + / 0
1 2 H H H H - F B " H H H H U ' H H H H b 1 2 H H H H $ = / H H H H b [ R q r 1 1 V % K s H H H H = / H H H H B 1 2 H H H H - 3 X G Y t H H H H = 1 S H H H H + 6 ; &
V % ( E H u \ H G d + 1 2 H - 3 X G Y t H % + R 9 H $ B v ' H B 2 H - = 1 L 0 [ V H % # U 1 2 H ` G d + / 0
\ H H H H H H H ] ^ 1 2 H H H H H H H 3 B 2 H H H H H H H + / 0 " ( L ( M 1 2 H H H H H H H N > 1 M ' H H H H H H H N B v 6 ; & w ( 1 6 H H H H H H H A 1 2 H H H H H H H F A 1 2 H H H H H H H +
R 1 a ) H # B p ! R I 1 2 H - 4 G A ' H B 2 H 4 B ) ( I / 0 S H B 2 H + 6 H A c 1 V H % B 5 1 i s H = 1 p 6 ; &
1 2 4 ( 1 6 A 1 2 4 B ) ( ) G + 1 2 ( 1 j J + 1 2 - $ J G K 6 ; & " ( L ( M 1 2 N > 1 M .
R q r 1 , H H H B ) H H H T \ H H H B V H H H g 1 ? H H H 4 - 7 G B C V H H H - < B m & " k s H H H J G F + 1 2 F 4 ( ! 1 ? H H H A ) H H H G +
R 1 2 H H - * B . E H H + R 1 2 - 0 V * H H A 9 H H N A \ H H B / 0 1 2 H H F 4 A m H H = E H H F + 1 ? H H - F B w H H A E H H P [ / H H B I
\ H H H H H B V H H H H H g 1 ? H H H H H 4 - 7 G B C i V H H H H H - < B m & " k s H H H H H J G F + 1 2 H H H H H F 4 A 1 a ! 1 . e [ ' H H H H H U _ I
1 2 3 H A E H x 6 H P 1 2 H F 4 A 1 a ! 1 . e \ H N B i E H F X : " 7 < ( ) H g R r 2 H y 1 ) H $ z w H B " H P
V { H H H H H ; | m H H H H H = s H H H H H b _ 9 H H H H H B 9 H H H H H G b R \ H H H H H < B 1 2 H H H H H 4 B L 0 1 a ! 1 . e R " H H H H H 4 - 7 G B C
1 2 4 B ) ( I 1 2 F B c 1 2 - 0 V 3 A ! s J G F + \ ] 1 1 2 F 4 A .
/ H H H } r 1 " H H H B 9 H H H ; < N B ' H H H } " H H H X B ) H H H G + 1 9 H H H - J A 1 l 1 2 H H H 4 B L 0 1 a ! 1 . e ' H H H B 2 H H H 4 B L 0
1 2 H { B ~ 1 2 H ] e \ ( 1 ? H 3 X Y [ / H B ) H g i E H < X P 1 2 - Q H ; G Y ' H B 9 H - J A 1 l 1 2 H 4 B 6 H A O
1 2 H 4 B ) ( ) H G + ' 4 ( 1 6 H A 1 2 H F A l R 1 a ) H # B p [ r 2 H y _ I 1 2 H - > 1 c 1 a ! 1 . O ' H B 2 H 4 B 6 H A O
1 2 H H 4 B ) ( ) H H G + \ ( " 3 H H A ! _ 9 H H B 9 0 2 H H A R 2 H H + 1 2 H H 4 B ) ( I R 1 ? 8 9 H H Q B C R E H H < - A
\ H H H ] 1 1 ? 3 H H H A ! 2 H H H G D < U j H H H < G k _ m H H H X B l ) H H H D B n 1 a ! 1 . O 9 ( 1 v V F ; K 1 W " H H H =
' B 2 4 = 1 . 1 C 1 a ! 1 . E + _ R 1 2 F 4 ( ! 1 a ! 1 . E + .
R 1 ) H H H $ z w H H H B " H H H < B 9 H H H ; | E H H H * q 6 H H H B ! O 1 2 H H H N = / 0 t H H H G B H H H + " H H H 4 - 7 G B C
1 ? H H H B ! O 348 " H H H P " Q ( ! O 1 ? D H H H = R M R 1 9 H H H - h N B v 1 2 H H H - 3 X G Y / 0 1 ? H H H N B 6 H H H +
1 a ! 1 . E H H + " H H P S H H = E H H + 1 Z H H - G B . 1 2 4 ( 1 6 H H A 1 ? H H $ J 4 + 6 ; & " ( L ( M 1 2 H H N > 1 M R
, ] 2 y 1 9 - J F B ! q " X B ) G + 1 2 - 3 X G Y ' B 2 # ; / G ` B .
1 . 1 2 $ F P ' } 6 B ! O 1 2 N = ) 1 ? B ! O 364 " P 1 ? Q ( ! O (
S H ; T 1 ? H B ! O 364 " 3 H U 1 2 H % # U 34 - 327 R w H 7 T ' H B I 1 2 H 3 X Y 1 2 H # B ! .
6 H P 1 2 H ` G d + 1 2 H - 3 X G < G + E H < X P 1 I E X ( I " ( L ( M q 6 H B ! O 1 2 H N = - s H J 4 B
2 ; D H = R n 1 ? H 4 = . O 2 ` H ] 1 1 2 N ( M " H P 1 2 $ F ( I - _ " H B c 1 ? H 3 X < + 1 2 - 0 , H B ) H T
9 - N = / 0 1 2 4 7 G + / 0 S B 2 + 6 A c R j ( ! 1 V % B 5 1 2 - 3 X G Y .
R , H H H H B I S H H H H = E H H H H B ' ( 1 L F 0 1 ? H H H H B ! O 364 " H H H H P 1 ? Q ( ! O 1 2 H H H H - N # G 6 ; &
R , H H H H B I S H H H H = E H H H H B ' ( 1 L F 0 1 ? H H H H B ! O 364 " H H H H P 1 ? Q ( ! O 1 2 H H H H - N # G 6 ; &
1 " H X B ) H G + 1 2 H $ F P ' H B 6 H B ! O 1 2 H N = 1 " H B c 1 2 H ` G d + 1 2 H - 3 X G < G + " H B ! 1 " H T 1 ) H ` B
\ 0 1 ? H H H H H H H # A . O 2 H H H H H H H ; < 4 = . 1 2 - 3 X G < 0 [ " k / - 1 " H H H H H H H X B ) H H H H H H H G + 1 2 ; * ( v 1 2 &
1 2 4 7 B v 1 2 = 9 < 0 ' F A V F ] . 1 2 $ F P 1 " B c 1 2 ` G d + 1 2 - 3 X G < G + .
1 . " A 2 ( l 1 2 - 3 X G Y 1 2 A R 2 0 ) 1 ? B ! O 370 " P 1 ? Q ( ! O (
V H J N T 1 ? H B ! O 370 " H P 1 ? Q ( ! O " H 4 - 7 G B C 1 2 H % # U 40 - 327 " H P 5 c c R
S H B / H T 6 ; & " % ` ( c 1 2 H - 3 X G Y 1 2 H A R 2 0 1 ) H $ z w H B " H P 1 ? H > ' H b 1 ? H F G B .
1 i w - # B ! e R 1 ? F G B . 1 2 * @ = 1 / 0 [ / 0 S b _ I 1 ? F G B . 1 i w - # B ! e
E H F - J = _ , H h = 1 ? H F B E H G = 1 ? H F - < A O " H P w H J U 1 2 - D H = E H F B C 1 2 3 H A E H h + R 1 2 H % 4 g
R 1 2 4 7 B v 1 ? 4 B . ) b .
R 1 ) H $ z w H B " H P r 2 H y [ / H } ) H N B ) H = f ' H 7 = R . O q 6 H B ! O 1 2 H N = / 0 " 7 < ( I
1 ? H H H H H B ! O 370 " H H H H H P 1 ? Q ( ! O [ R 1 6 H H H H H - J B . _ I 1 2 H H H H H - 3 X G Y E X ( I ! R 2 H H H H H G B " - &
V F ; K ' H H H < # B 2 V H H H * B . E H H H + ! R 2 H H H G + ! R I _ Z H H H ] " H H H X B I 1 2 H H H - 3 X G Y _ R w H H H B ) ( I
q j = 1 v 1 C 1 ? 3 B , < + 1 ? $ J K _ R j N Q G + 1 W s = 1 p ' F b 1 i 6 - J B . .
1 . 1 a w H H H H H H H H H = 1 . ' < % ` ( c 1 2 H H H H H H H H H F 4 A 1 a ! 1 . e 1 2 H H H H H H H H H A R 2 0 ) 1 ? H H H H H H H H H B ! O 376 " H H H H H H H H H P
1 ? Q ( ! O (
E H H H H F - J = " % ` ( c 1 2 H H H H F 4 A 1 a ! 1 . e 1 2 H H H H A R 2 0 " H H H H P 1 ? H H H H % B \ H H H H G Y 1 2 3 H H H H A E H H H H h + / 0
1 2 H H H 4 B ) ( I 1 a ! 1 . e [ / H H H } 2 & 6 ` H H H A H H H G = ' H H H F G A , H H H B I " % ` ( c 1 2 H H H F 4 A 1 a ! 1 . e
E - 3 H H H A ! / 0 1 2 H H H F 4 A 1 a ! 1 . e 1 2 H H H A 1 Z H H H ; 0 / H H H 4 [ H H H G = _ ) H H H g R ' H H H B 2 H H H N = q 2 &
1 2 - $ ( . 1 2 H H ] e 6 H H = / H H - g 1 2 H H - A Z H H z C 1 i w H H - # B ! E H H + 2 H H ; A R l 6 H H J = 1 2 3 H H A R ! V H H Y
1 a w = 1 . ' ( j ( ! 1 2 F 4 A 1 a ! 1 . e 1 2 A R 2 0 / 4 ` B R w 7 B v 1 .
R 2 H H H H F U " H H H H P S H H H H Q N B C \ H H H H B V H H H H g 1 ? Q ( ! O _ I 1 ? H H H H B ! O 376 " H H H H N ` B - R 1 2 - 0
S H ; T " 3 H U 1 2 H % # U 46 - 327 " H P 5 c c - V H 7 < N T 1 L H B / H + / H 4 = O 1 Z H G = O
V H H H H H H H Y ' < ( j H H H H H H H J ` B 1 9 H H H H H H H N B ! 1 i Z H H H H H H H - # B ~ 2 H H H H H H H ; 4 7 B v 1 i ! 1 . e 2 H H H H H H H ; J T / 0
1 i 6 H H - = 1 p R " N 1 2 H H # G @ + 1 2 H H - N % G ] E H H + 2 H H ; < 4 = . 1 C 1 2 H H - 3 X G < G + 1 2 H H A R 2 H H G +
1 2 # B ! . O / 0 1 ? N B 6 B C 1 ? - F ; 4 + ' B 2 F 4 ( ! 1 i ! 1 . E + 1 2 A R 2 G + .
1 i 6 H H - = 1 p R " N 1 2 H H # G @ + 1 2 H H - N % G ] E H H + 2 H H ; < 4 = . 1 C 1 2 H H - 3 X G < G + 1 2 H H A R 2 H H G +
1 2 # B ! . O / 0 1 ? N B 6 B C 1 ? - F ; 4 + ' B 2 F 4 ( ! 1 i ! 1 . E + 1 2 A R 2 G + .
/ H H H H H H < 4 - 7 G B C 1 2 H H H H H H % # U 46 - 327 " H H H H H H P 5 c c , H H H H H H B ) H H H H H H T w H H H H H H A Z H H H H H H ; 4 T 6 H H H H H H A O
1 m H H H H H X B i C w H H H H H B ) ( ) H H H H H G + R r 2 H H H H H y 1 ) H H H H H $ z w H H H H H B " H H H H H P _ I 1 2 H H H H H 4 = 1 v O 1 2 H H H H H B \ H H H H H = E H H H H H +
? H H H H 4 - 7 G B C 1 2 H H H H % # U 310 " H H H H P 5 c c V H H H H % G A ' H H H H B I \ H H H H B V H H H H g 1 ? H H H H 4 - 7 G B C
V - F ; K ' H H H B 2 H H H - 3 X G Y 1 2 H H H A 1 Z H H H ; 0 / 0 1 2 F 4 ( ! 1 a ! 1 . E H H H + / H H H 4 [ _ " H H H B / H H H G < B
E - F ; K ' H H B 2 H H - 3 X G Y 1 2 H H A R 2 0 / H H B ? D H H = M 1 ? H H @ = ' 0 2 H H Y E H H < G > 1 2 H H F 4 A 1 a ! 1 . e
6 H H H H P ' H H H H B w 0 1 2 F 4 ( ! 1 2 H H H H - * B . E H H H H + R 2 H H H H Y E 3 H H H H A ! _ e w H H H H G A " H H H H P 1 2 4 G ( ! 1 2 - 0
S H A ! \ H B ' { # ( ~ 1 2 H - 3 X G Y 1 2 H A 1 Z H ; 0 / 0 1 2 F 4 ( ! 1 a ! 1 . E H + [ , H < B _ ) H g
_ w # & 1 2 H H H H H 4 7 B v 1 a ! 1 . e " H H H H H P " H H H H H * B l V H H H H H ] E H H H H H G U 1 W S H H H H H X B c 1 2 H H H H H - 3 X G < G +
' H H H B 2 H H H # G @ + 1 2 H H H - N % G ] E H H H + " - & V F ; K 1 i " H H H = ' H H H B 2 H H H - 3 X G Y 1 2 H H H A R 2 0 [ R _ 9 H H H N A
1 W " = q 2 & 1 2 4 7 B v 1 2 - * B . e .
R ' H H H H B 2 H H H H = H H H H Y " H H H H P t H H H H G B H H H H + 1 2 H H H H % # U 46 - 327 " H H H H P 5 c c [ / H H H H B I 1 2 H H H H 4 7 B v
1 a ! 1 . e 1 ? H H H H @ = ' 0 , H H H H B I w H H H H A 1 6 H H H H - J = R 6 H H H H P S K - ' H H H H o ) H H H H g R q r 1 1 2 H H H H 3 X Y
1 2 - 3 X G < 0 " ( L ( M s H H ; 1 2 H H - ] E H H G U ' H H B 2 H H # G @ + 1 2 H H - N % G ] E H H + t H H B ! . / 0
q s H H B . ) H H > 1 M ) H H B m H H 6 H H P V H H N % G ] 6 H H 4 A q ! 1 . e / H H B I 1 i Z H H - # B ~ E F ( ! q 2 &
1 2 H H H H 4 7 B v 1 a ! 1 . e [ " H H H H B ! 1 c _ ) H H H H g E H H H H = j k 1 Z H H H H - # B ~ 1 2 H H H H N = / 0 s H H H H ;
V H H H H ] E H H H H G U 1 2 H H H H 3 X Y 1 2 - 3 X G < 0 1 2 H H H H # B ! . / 0 q s H H H H B . 1 2 H H H H N > 1 6 H H H H B C 1 ? H H H H - F ; 4 +
' H H H H B 2 F 4 ( ! 1 2 - 0 V H H H H J = " H H H H ` B 1 2 H H H H A R 2 H H H H + q 2 & 1 ? H H H H 3 X < + 1 a ! 1 . E H H H H + 1 2 - 0 9 H H H H G - Y
V H H H H H H N % G ] 1 2 H H H H H H 3 X Y 1 2 - 3 X G < 0 / 0 ! 1 H H H H H H = V H H H H H H ` B _ R q 2 & 1 ? H H H H H H 3 X < + 1 a ! 1 . E H H H H H H +
' H H H H H B 2 H H H H H = ' H H H H H B n 6 H H H H H N A " H H H H H B E X ( I V H H H H H N % G ] 1 2 H H H H H 3 X Y E H H H H H D < U " * H H H H H < ( M 1 2 H H H H H - = 1
1 2 ( s N 0 ) 1 W " H = 1 2 H # B ! . 6 H P 1 2 H Q G A . H G : 1 ? H 3 X < + 1 2 H - * B . E H + ' H B 2 H = ' H B n [
. w Y 670 [ ' - B . E 18/6/2012 [ / 0 1 ? ; | . w Y 414/3/2012 ( .
R 1 V H * B ^ 1 2 H 4 7 B v 1 ? H @ = ' 0 \ H ] 1 [ 2 H Y E H X P ) H D B 1 [ ' H U _ ) H g j H B v " H Q B E H = 1
' H H ] 2 H H y ) ` 1 2 H H 4 7 B v 1 2 H H % = ) Q 0 / 0 w H H 7 G + " L'INSERM" R 1 2 - 0
m H H X ; T ) H H 4 ; + ) ( 6 H H G + / H H G < B E H H { 1 i Z H H - # B ~ / 0 1 2 H H - 3 X G Y 1 2 H H A R 2 0
/ 0 1 2 F 4 ( ! 1 a ! 1 . E H H + R , H H ] 2 H H y 1 2 H H 4 7 B v 1 ? H H # = e / 0 w H H 7 G + " m H H = , H H +
" B 2 G X ( . 2 G < - A " .
/ 0 1 2 F 4 ( ! 1 a ! 1 . E H H + R , H H ] 2 H H y 1 2 H H 4 7 B v 1 ? H H # = e / 0 w H H 7 G + " m H H = , H H +
" B 2 G X ( . 2 G < - A " .