Lifelong endocrine fluctuations and related cognitive disorders.
Texte intégral
(2) HAL author manuscript Current Pharmaceutical Design 2005;11(32):4229-52.
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(9) IHKJ9 LDE(3#5MN(O,"&# "0!P Q R 6 HAL author manuscript. FG(S?0T# #2"&VU$W $X(S?0 &@A Y,Z;0(9 :# * \[G#T(S;0(S@ #LD:$].F^0UU X$&UX`_a"93DA 8#2# b,. * 7ced$[A E(*13 fb #:g&K`UKXXh.U dh$+ij[A(Sckg&h`UKXX`.UdKlX$i_m(3T#Eg&(*1* *# T no!"0SDpkq T
(10) ^q rE %'&)(+*,.-(. inserm-00243083, version 1. fN; `(3Ts"&rNDE; 5 R ? t .D"u
(11) c+(3!/ YDp; K : *#T(8Dv "& ; wCP xDvth * m "&1* :/ hr#w@ 1
(12) D@ (DE "&u(*+e19"&% LDE/? hr@+ 1
(13) DE2"& T%Gq$y # RDE;+"j (h"&r9¨t¦¥@¨»§l£}«K(* ¨l¦¥@¨t¥@«e ED@+j2 6 I;(*? z !"&{ EDv :(8D *Dv; 5 7@ "3+ :"*D: *1
(14) DvT? 5 :"#2 "&rb * EDE "&% .(S D; *T \TK (31
(15) D|"& D; }
(16) @+ x"*DE (* !LD D 8 ' ED ! `TK{#T 1*(DE *CT~13"0% LDE "&Gq4 13 3D; "& !"&(3#N R 7{#T(913 ! 3DbDv; :(S'2Hh4fb\DE :T(3# zTC "& ! jD *OA"0 vD:! "9 (S@{ R(9#th"0! @+%% * ED!(mDE K+"j R(9 E') " RLDE/? a *rEr *1
(17) Dep "*DE(*P #w'"&? 8 vP (3#z! !"& '0:(* rE"9@. : 13 3DI!
(18) DE(j(S0(9#w'$ 7 * 19"&? E% 5D"^ x@ %%+ 6 EDI(K "
(19) RwPA# z{ "6Dv *1
(20) DET? K *rEr *1
(21) D.Tu R 8#T(8Dv "&!D:"^yh#T*; *T! 8 z 7 8(&
(22) K : *&@+1*T% : KP+'!W X`DE"!U$U
(23) q9HK"&th ? x+&(8Dv(hrE "&D; `"&#T'#(9 E% z :(*"0!9 *!19"&3DE :"#2# *DE :T(9#+@PA# ; *hD"!&(DE 6&Dv; "&! { Hh (j#LD;E+ Dv(DET? o_O !"j 'BjD@+3' z "*D1*"&r D; *
(24) O"9P{ 7 8 v?0(DE "& C(S );(9? O ? @%%+ 6 ED *(*T1* R 8(& R OT ! jDT( : rE"& t`"&! @{ R/%sHh4f1*"&` (3 *Dv"19"&3DE :"&# *qoB ?+ 8 (3#O! RDE;+""&# "&% T1*(9#)TrEr: R 019 3 PA xDt` * * "9P{
(25) E?(DE "&u ED@ 2 6 (*+!13"0SDv :"# #2 *hDE (9# t Dv;e %(3 :`D:"> (DET 3D\% :"9@{$D'&{ *DE/!T%e(S 9@ :(DET"&u"jr|Hh4fZ1*"3%$ LDET?. ! 8(& @ R * \(*h(S(3#T' 7 6
(26) $(3 R1@A
(27)
(28) *5DE"K 7cA#(9TD; *
(29) 5 R1* R 7 (*132 6 *q9f|;0 (j
(30) "j19T(8DE "0hPA xDpth * !; "& v!"&(3#A
(31) E@+# ? *# (*m13"0%j+ DT? >r@ 1
(32) D "&T%m R V(&T `19"&3DE :"&?+ 8 R RT(3#. @ %$% 6 vD:T&%o;0/%;aT3D: x/ T? 9@ (9#\?j@A#T :(SP T# LDE'a/ / xq_O"& : *"&? x : *
(33) (3 1;~"&eDv; >Th+(31
(34) D"r5 "&1* :T+ Kr@ 1
(35) D "&T%m"&~19"&%LD: "09@ $%eDE;+ Kr: K(j#2 K x 7 "9@ 1
(36) D:/? K1*'13#2 ! @ %$% 6 vDx z#Tr 3 # "&&%mrE#@A1
(37) D@ (DET"& /~?&@ #T+ 8 (*P #TLDv'qGDE "&# "&%+1*(9#N!"$ *# DE(*T%CT3D:"u(91*13"9@3D\DE; >wSD :(31
(38) Dv "&m"jr13#TT T1*(9#/ : 7 :"9@A1
(39) DET? * (*`! { EDv E@ (9#A
(40) ? 3D: Dv; :"9@+%; "9@jD# Tr: V(9'` :"&?+ (z!" ?(9#2K(* "&(31;KTh@+ x vDE(* &T$%Dv; 5 *rEr: *1
(41) D: I"&r vD: :"20 I"& D; `P (3w~(S ~wm RD: !T T%m @P{2% R"9@{ z(8DI;0 */%;3D: *C : GqG,I"9% LDET? >
(42) "& 8 R /eD; *# :#/'~(3 K!"& R K# T *#T'P{ : #T(8D *DE" 1@+`@{#T(8Dv * # Tr: *# "&$%
(43) cA" @ R -Dv" ED :"
(44) :(
(45) Dp; 8 oD;(*D:"¡( *1*2rE 1 7c{" @+ : -D"¡(% T? ED 8 R"& Aq _m@ #LDETrE(91
(46) DE"& :T(&#\!" *# zP (j 7 *m"9a(*a 7c ;(S@{ EDET?+ !?+ t"r5(3#T#\; "& !"&(3#. ? 3D: 5Dv; :"9@+%; "9@jD5 : 7 "9@ 1
(47) D:/? # Tr: KDE"&% RDv; 8 thLD;e"*D;+ 8 \ : !rE(91
(48) DE"& x .+"6Dv(*P #T'K%+ xDT1 : !rE(91
(49) DE"& x \ R #(*DE *5D"! * EDE :"9% ! R 13 7jD"j {"&#T'!"& v ; } b ; "9@A# PA 7cA# "& : * DE"`1*#(9 :TrE'D; :"&# 5"&r; "& !"90(&#+ : KrE(31
(50) DE"& R
(51) "& "6Dv 81
(52) DE/? rE(91
(53) DE"& x \rE"& N1*"&%+ DvT? 5&' Rr@+01
(54) Dv "&K(* ` e 3DT( q !¢£¥¤V¦§*¨A©*ª 19"&%& LD:2"& «yh#T*; *T! 8 ¬ 7 8(& R * * EDE :"0%
(55) r: K(&# s : 7 :"9@A1
(56) DET? s1S' 1*# *! "&+(*@{ R *«; "& E!"0{ : 7 #T(319 ! jDAD; :(S' 1@+h@+#(D: *`#T r: *#2"&%K
(57) cA" @ bD"V EDv 8 R"& *q.
(58) Ã 45&)( 6.7 %$%$*98;:94<,=(>456.?9&. HAL author manuscript inserm-00243083, version 1. yegSyh# ; */e 8 \ 7 8(&
(59) y_aFAyg3(9KT "Kz;' & :"9c+'}dz! RDE;$' # R"9c (39"# U` :"9 T"& (DE y5{"&5g3(S "&# w{"9 "6D TV yFGFbg9(*K'+# "T+ : *1@ R R"& :"*DE T y β: (*`' # " β ]<K[\g*P (3wK 8 T? *` 7@ "6Dv :"9 ;+15rE(91
(60) DE"& ]._!gSP "&+ !/ 8 (3#+ { RLDv' ,.Ng31*"&:@$%0(D: *K 9@{/ 5 6 EDv :"&% . ,N<Bg31* 3DE (3#$ E? "9@{ . ' ED ,."B f|y4hg& ED@ 9'!"&rADE(*!"9cATr: K(* ` :(9#"9c Tr G19"&% LDE2"& ,.4N]g31y_oFu : * "& 7T? *E *# 3DP T /%h+ :"*DE *T ,.4Fbg3,o : (31
(61) Dv?+ . "6D *T 5yeg9"9 (9!T. .Bg3& RDE;$' # EDE #wPA 6 EDv :"# N4hg9 * EDE :"0% V 819 jDE"& N4f5g9 6 EDv :"9% K : 7{#T(319 ! 3DAD; :(*$' [NB$Heg3rE"&# #219# Y vDET`@{#T(8DvT%`; "& :!"0 6 yh]Iyeg (*!/ "9P@jD' :T15(31*2 4.Heg9%+γ"&(3"*DE "3+TV *# (j RT$%h; "& !"&. H
(62) MgS; T%$;V 7LDE'K# w "9 R"*D: T Hh4hgS;(3*(9 :` E(8DET" Hh4f5gS;0"& K"& R 7 #T(319 ! jDAD; :(S' M Mg9# "&t 7 Dv'K# w{"9 "6D T MHeg9#w@jD: */02*T%K; "& E!"0 6 _my
(63) >g9!"& "&(*!T+ Y"9c 2j(&
(64) _myF>g9! D"&% K(31
(65) Dv?(D: *z :"*DE */h /0(& R KE_a_«B g&!"$r *&!T p! jDE(3#A EDE(D@{ |
(66) c+(*!T(8Dv "& _aFy}g9! *& R"9c'& R"9% 3 ED 8 "& (913 RDE(8Dv <`yegS "& :(9j (3# T+ < [\g* E?+ % "&tDE;KrE(31
(67) DE"& <K_yeg8<>! RDE;'+#/`(j (9 D(8Dv ,g9"& :(9# 1*"&3DE (31* 7jDET?+ 54hg9"` :(
(68) Dv" FGf5g* " RLDE "&! !
(69) R "0DE"0!"&% :(*$;' FG:>g*;+"0 x;(8DE 0' #TT "
(70) LDE"$#{pT(j 7 FG4NFAy4g*
(71) ?+ SDw%`{" EDv! "&(*@A R(3#$! !"& :'K# "
(72) \(S y`#S; *T! * \
(73) (j 7 .thLD;K R
(74) +#(913 ! jD| 6 vDE :"9% . 4Y,\f5g3 E(S"&!T3 *`13"&3DE "# # *5DE (9# B N4_-g& R # *1
(75) DET? 5 * EDE "&% K : 13 jDE"& |!"&3@A#T(8DE"& B Fb,\fYg& Rw&% # I; "*D:"0! !/ 7
(76) T"&K13"&K$@jD: 5D:"&!"0%0 (*;&' Hh:_oRBkg9th"&! |; 8(9#LDE;KT LDET(8DvT? Yp! *!"& ' vD@ &' RB5_-gSth"&! k \/SD E(8DE2"&(3#A ED@ &'K"r# "&&%!9@ (8DET"&K" * EDE :"0% V(9r D: ! "9 (S@{ 7 d2Hhf5gj 8 "6Dv"& T.
(77) Å B?=(+*=6.C'D=-(;436.?. f|;0 u 7T% rE 1*(S3DE#T'a;+T%; + : ?(3# +13 u"&rzyh#T*; *T! * K 7 8(& R ~py zhTmA" ED:p! "9 (*@{ (3#\th"&! )"9P{ 7 8 v? *a2 !" ED|th 6 vD: !13"9@ 3DE * \;(j I# *D:">w 1* : (j R/%eT3D 8 6 vDbwKD; 5 :"#2 "&rN% "00(9&(3#A;+"j !"&+ 6
(78) +(S!TK (9 DET1@ #T(9 b * EDE "3%+ $+2 1*"&% LDE/? r@+ 1
(79) DE "&+T&%q T? KDE;(Db * EDE "9% e R 13 jDE"& R (9 E h 7cjD
(80) T?+ *#T'! 6 7 3D\T!P (3wVDv/ 7 @{ *A(`t"& :#20pth ` 7P{(DE HAL author manuscript. 1@ :
(81) 3DE#/'m@ $ 8 t(*'a19"&13 E T$%~D; >{"*D 3D:T(3#\?(9#@A u"rz; "& E!"00(9#. R
(82) #T(913 ! jDD; :(*$' 2Hh4f\wa R 89@A1*T%a(*%+ *T% : #T(8D *!13"0% LDET? `1*;(S% *
(83) A(S *#T(*' T$%$ r\ "*D| : ?+ SDT% $D; `"&
(84) RD."r. ! &DET( q4 *13 3D\r:T+j/% rE R"&DE; `# (9 E% 6 vD DE"!&(D: (*"&! 7 *KA#T(313 7P ">13"&3DE "# # *}Hh4faD (9#Gw13#w@AT&%u"&? .U T}t"&! u;(*? ` @ %$% 6 vD: *+;+"&th ? x$D;(8DH`4.f V(9'u;(S? K(>;0(9 EKr@{#| *rEr: *1
(85) D"&~13 R
(86) P+ :"?(j 1@ #T(3 Y x'$ vD: e 5(Su13"&%$ LDET? Kr@+1
(87) D: "9 /% G(S K(9' ? CT 1* : (j 7 ` : C"r. inserm-00243083, version 1. ! jDT( q+fN; *
(88) KrE/ T% ;(*? Kr"j 13 *>@{ .Dv"u : ? 2 t : ?+ "9@{ ?+ 13 Kr"j (K "*DE *1
(89) DET? > 8rr: 1
(90) D.(9!DE"u ?+ *# "&m!"& : :"9P@{ vD
(91) 2 ! "# "0% T1*(3#+!" *# \"r / xq aDE; ? tth C(3 v%&@{ DE;(DD; C!" ED! 8 "9@{ u ; "& D:1*"&!T%"jr`+ x ? "9@{ ED@ 2 6 ^;(j >P * aD;+ * >r:"&1@{ "$ ; "& !"&(3#51*;(*$% 6 ^(3r DE hD;+ C! "& (S@{ R *.t` Dv; "9@jDDv(*T%)T3D" (91*13"9@3D>+T? 9@ (9#5# Tr: 3TDET! o
(92) c${" @ eD"o; "& E!"&(&# ?(3 :T(SP T# LD' A(*+VDp;0(D\K(3$'^"&rNDv; `T13"& 7/ vD: 7 132 6 r:"9@+ >P{ RDvth * ~ ED@ 2 6 K(9'P{ `9@{ 5D:"`DE;+ hrE(91
(93) DkDp;0(D\th"&! C"> "*D (3 E ?+ !(8D! "9 (*@A
(94) >thLDv;a 3@+(3#. : O"&r13"9% LDw? u *1*# / "j ` 9@ (3#. @{ 13 jDEwPA #TLDE'uD:"Dv; u rEr: *1
(95) D: }"jrHh4f5q VDp;j@A : * 7 SD(` @KK(3 E'K"r1@ : 7SD "&t`# *&% 13"&13 w%DE; ! *1S;0(9 Re \"$r{Dp;
(96) @+ :"9 R"*D: 1
(97) DET? (31
(98) DT"&!"&rN 6 vDE :"&%+ { \(* DET!T$%!"jr; "& :!"00(9#A c " @ R \DE; :"9@+%;"9@jDADE;+ Yr V(&# 5# Tr 31*'13# *$(* KLD: | "9P (*P #2 8rEr *1
(99) DN"&!1*"0% LD:w? 5r@+1
(100) D: "00/%Gq8 D; K R ?02 t ED@Aj2 6 Z@ $ 8 DE(* hDE">j(DE 5"&DE; 519"&%LD:T?+ 8rEr *1
(101) D: ."&rN 7cA"&% "9@{ Z; "& :!"00(9#A *rEr: *1
(102) D"& x
(103) 0"6Dv(*P #T'`Hh4fZ$(* rET(3# #/'} @$%% * ED(*KwSD % :(DEw? !" *#A"&r : `P (j R *K"0D;+ 6 7 rT w% DE"`%&@ $ r@jD@+ : 6 7 (3 :1S;Gq HG)I8J-;K,?945&#LM&6.7N=6.?,C,9OK6.*9L6.?P8Q8;7>7R8J-($&6P?S-'6TN.?945(>436.?. ].
(104)
(105)
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(107) T?+ *#T'a : ? 2 th *R8E(* m1*(SmP{ u @+KK(3 :23 *m(j }rE"&# # "&th *qf|t"uDE'&A 6 >"&r 6 vDE :"&%+ e R 819
(108) &D:"& x 5N4N;(S? 5PA * ^2 3DE rE *uN45 α (S !|4Y β|t; 1S;e! *$(DE `TrEr: x 3D\r@+01
(109) DT"& *qf|; *
(110) z : *13 &DE"& x (3 Y 19" *`P+'hDEth">& rEr: R SD% 3 I# "&1*(8D *!"9e
(111) 7 (9 :(8Dv Y1; "&!" 7"&! 6 ZP@jD\ ;(3 :/%!13"&K!"0^ EDE v@ 1
(112) D@ E(9#ArE 8(D@ 6
(113) $t` Dv; D; : * z :/019w (3#"&K(&T }Dv; `(9!+" TD: V/0(9#N"& \y p]"0V(&w Dv;
(114) <Kyh2PAT w%~"& ,"0V(&T~E ; "0thT%!"& R YD;(9 Xd ;"&!"# "&%$'uP{ RDvth * . α. (*+ β:(*!Dp; !#TT%(9 2P T $T&%"& 5-&"&K(&TEdX$s; "0!"# "&%'
(115) q+y¡r:"9@ Dp;~"&K(&T[G . T?"#T?+ *uT~ EDvT%&@{ ;T%P{ RDvth * * EDE "&% ~(S% "& ED 5(*(9SDv(*% "9 ED: Sq M|T K"*DE;+ 8 ! KPA 8 R "&rNDv; ! ED 8 "j2u : *19
(116) jDv"j @ :rE(9K #T' .|4 (* : u# /%0(S (31
(117) DE/?0(D: *CDE :(S R1* :wjDET"&OrE(31DE"& x *q|b ED :"9% >1*(*) 7c{ D5D; *T > 8rEr *1
(118) D: }@{ RT%mD; !S13#T(j 7 RT1*(3#.
(119) % "&!21m *#T(*' *o (D;t(*'^P 'oP{T+j/%D"OT3DE :(91* *##w@A#T(3 K : 13 7jDE"& R hDp;0(D`TD@+ E)# (3mDE"o&@{1*# (3 Dv :(* R#T"&1*(8Dv "& \D; T 9@ 1
(120) DE2"&"&r% + aDE :(S R1* :wjDET"&(S )DE (* R#T(8DET"&"$r :"*DE */$ D;(8D1*(*!"9@A#(DE O 7@ "&(3#! @ v? w?(j#K/ ;(*19T% 7@ :"*DE "3+; T1a @$ "& D:5 @$0 6 7 RT%)(*+"3&DE"
(121) "& !(9rErE 81
(122) DT%O 7@ R"9(j# EDv E@ 1
(123) D@ jq\b ED "3%+ 1*(9)(3# R"O(91
(124) D`? E') :( 2j#2' D; :"9@+%;"&% "&!21O! *1*;(9/ e mw3D 8 (31
(125) DET$%jT R 1
(126) D:#T't` Dp;?0(9 : "9@{ C/SDv :(31* *#2#@+#(9 ~ Rw%(3#2T% (8D;$t(S' mT13#/@ w% HAL author manuscript. !LD:"&%+ (91
(127) DET?(8DE * "6D *w T(j _oyF: 1yh_aF. 6 +"&
(128) w? 3v *# ! jD¥P T+T% :"*DE */ E,I4N].: (*. ; " ;(DE &' #TT+"0 7LD:"&#p T(j ^F::D;+ 8 R 7P 'a(9rEr: 1
(129) DET%aT+T : *1
(130) DE#T'~Dv; KDE (* R1* :wjDv"0 "&rh 7 ? 8 (3#."*Dv; 8 5DE(* v%0 RDz% * *q b EDE "9% e1*(9e(91
(131) DET?(8Dv V 7 *# 1
(132) D: *T3DE (31* *# #w@ #T(3 T%(3#TT%>{(D;&t(*' 5 A &T%u"&!D; ` R 819
(133) &D:"& @P&D'&{ 5PA"9@ +{q0y "*Dv; 8 "& 13#T(&
(134) RT1*(3#+!"$ "rN * EDE "&% K(31
(135) DE "0V(DAD; 5! KP (* ;(j \(9# R"PA * K "3A"
(136) *K/? "&#T? T$%Dv; .PAT &T$%!"rN * EDE :"0% D". inserm-00243083, version 1. ! *KP (* 3(j 7 R"&13T(8DE *` : 13 &DE"& x
(137) 0 "
(138) RwP #T'K(8DADp;0 1*(*? *"$#(9 5# ? *#Eq ]."*DE;¡N4 @PjD'& * «13"9@{#TPA T?+"#T? */ * EDE "&% 7@ "3+ :"*D: *1
(139) Dv "& (j @%&% * ED *P'Dp; :"*D 81
(140) D "&¡"r ; w "1*(9` (3# 7@ :"& ZP'!(9%0"0 ED: .r"& b *LD; \N4 N4. α. (S0. β. "&1S "9@jD\!T13 *AN4 α++"6D\N4. β,. α. "& b|4 β $(*%(9 ED\ 7c 13LDE"*D"9c 13E/ 9@ 13 *>j(9K(*% 5(9 K (8Dv;!TU
(141) q 7T%. t(& 5 ;"&t!DE"KPA 3@AT : *>r:"& ND; 5+ :"*D: 1
(142) DT? ` *rEr: *1
(143) D: "&r. 6 vDE :(9j2"#G(S%0(9T ED. R1*; !T(&ET 9@ 19 *KT @ ' d
(144) qDE;+ Y"*Dv; ;(* 9D; .P+ :(3/$ \"$rk|4. β. & "&1*+"3@&D!T13 5 c+;0wPALD: *`(*!T1* R (j R 5TK(* "0jDE"*DET1. 13 # # `TeDv; K?+ SDv : 1@ #T(9 Y9"& * `"&rbDv; !19 8 R 7P :(9#N13"& Dv
(145) cuDE;+(8D : * @ #LD: CT(SP "& V(9# LDE * `TuP (3Tm ? *#2"3+! jD
(146) qN4. β. % R v@jDET"&Ct(j z(3#
(147) "r:"9@ +!D:"CTK (9T Y"9jDTK(3#. (DET(3#\# 8(9 ET%mTmr: K(&# >!19 6| @ %$% 6 vD:T&%uD; > :"# "&r\4. w. β. ! *T(8DE/%V#2 8(9 E T$%V(S0`! S!"& :'K 7c{ D *P$'! 6 EDv :(3& "&# l
(148) q DE; ;&@+V(9Kr:"& R
(149) P (3w &P+"6Dv;KN4- @P&D'&{ * \(3 . *"&!T(*3D#T'! 7c : *
(150) 7 *KT!#TTKPA13 R *#T(8Dv *h(9 : (j
(151) $(3#LD;"9@ %$;D;+ *' ; "&t j EDw 1
(152) DK EDv :wP@jDE2"&O (8D Dv 8 v$ Sqbf|; ~|4. α. K4b<}y
(153) c$ R 6
(154) "&)(*+ 8(9 x KDE"O"0!T(8D ~TaDv; C;'&A"6Dp;0(9#T(3`@G (*. (*K'%+j(9#T(jAw $1*(DET%KD;(8DD; αE @$PjDE'&A !/%;3D\!"9@{#T(DE +"6D\"&+#T'
(155) @+ :"&(3#13 *# #{"9@+#(DE "& T?+"#T? *T(*@jDE"9 "0! 1 (*!
(156) + :"9@ 1
(157) DT? z 7@ :" +"&1* :T zr@ 1
(158) Dv "& \P@&DN(3# 7"!(9 : (j ./hA#T 1*(8DE *>Tu !"*D: "00(9#GT3D 8 v : RDE(DET"&e(S ` :"$13 *
(159)
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(161) c$ R 6
(162) "&!"&rN4. β. β. β. &"&!T(93DN(9 : (j
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(164) 7$%$z 91*#(9 :(DEw? o! !"j '(* !"*D"& ur@1
(165) D:2"&T% R
(166) qM| D2DE# O"& ". I xD *1
(167) D *hT!
(168) + :"9@ 1
(169) DT? \D:
(170) @A * . @ 1;K(j D; @jD E@{
(171) 0t5; x N4. R 7c@ (3#\1*;(9 :(31
(172) Dv 8 / vDET13 `"r * EDE :"9% a *rEr *1
(173) D:. α. ;(j |P+ * e R;"&thDv"`! *T(6Dv bDv;{.
(174) q+LD;m!T /V(9#NA 8 w ; :(9#Z 8rr: 1
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(176) |N4. β. R # *1
(177) DE/? !(9%"& ED: `1*"9@A# . D;&@{ |P 3 RT%+ *YD"+ :"&!"*D ! !"& 'Kr@ 1
(178) DE "0V(S + :"*DE *1
(179) DE "0V(S%0(9T ED 7@ :"$ % :(DE "&K(j
(180) "j1*T(D: `thLD;Ky!q .
(181)
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(183) T13#w@{$T%e19"&% LDE/?{ T` (3T ! 3Dh(S a$ e 3D:T(&\1*(3 :& "p?0(& R1@ #T(3 z (D;"# "&% &
(184) \"& `!"Tr:T1*(8Dv "& "&r5D; C *rEr: *1
(185) D: >"&r` c{"9% "0@{ > 6 EDv :"&% )"&.
(186) #TA # ? *# 5T~"&# th"&! Gq&LD;u R %(3 VDv"e19"9% LD:/? KT` (3T ! 3D.(* ! jD(&+D; hN45 α KBbEXlf p,2F+?&@ E:: KBb8Eduy p,sP (9!TT3DE :"&¡85(*aDp; C :"0!"6D e % "&-f|y : 7A 8(DK "$#'$!"j v ;+/ ! ;0(S? m R 819 *T? *aDp; m!" D (8D Dv
(187) 3D"0q&f|; *
(188) R(9! A"#w'!"& E+; R! I;0(S? 5(&#/ "`P{ * TK # T1*(8D *KT!1*(9 : "9?(j 1@{#T(3 |(8D;"#2"&% 6 |D:"0% xDv; Nt` Dv;!KB\8 HAL author manuscript. U Of 2, wT3DE "&8>,\y x 7A 8(D:K(S0( "&? *#}{"$#T'!"& E+;/ T :"&!"*D 8 m] \ 8X j Xf 1*r
(189) q X E
(190) qzfN; ODth" "#w'!"& E+; } !" ED.rE R 9@{ 3DE#T'u(&
(191) R"&13T(8Dv *thLD;~1*"&%$ LD:T?+ hTK+(3T ! SDI(* u$ ! jDET(!(9 : `FA?&@ E.(*+zP (3
(192) g+r:"& I!"0 vD 1*(j R 13"&3DE "#A ED@ 2 6 T13#w@{$T% j 5DE"`WW K1*(& R * R:(` RT% rET1*(93DT1* x (j 7 /Dv; 5rE : 9@A 13 5"&rADE; * 7 ZDpth"A"#T'$!"j v ;+/ ! t(j "9P{ 7 8 v? *OTO (8Dv 3D: >thLD;- A"& :(3&T1my R 8
(193) q #/' "0 vD@ &'-& "*Dh ? (j#(*')(&
(194) R"&13T(8Dv "9 R 7U
(195) q\fNt" ED@ & * `;(9? }P{ * a1*(9 : :T *m"9@jDz"&F(3 v0/$ 7"&G K& 7 8(&
(196) >thLD;O ! jDET( g|"9 u8m1*(& R 6 ; "&t` *m(*m(j
(197) R"&19(DE "&mt` Dv;. inserm-00243083, version 1. FA?&@ E\P@&D\ "*DP (9\ "#T'$!"j v ; R :t;+ 8 R (j .D;+ K"3D; YUNz1*(j R * R ; "&th *+"e(&
(198) R"&13T(DE "&ut` Dp;^F?&@ E R 8d 8 qf|;. "& #w'a+ :" A 81
(199) DT? u19"9; "& Dh vD@ &'O"jr`W &W+dm+"3+ e 3D *at`"&! )"&? Vd=' (3 R K1*(9 E :T *a"9@jDD:"oj(D: (9#/ 7"a @+%% * :D K(* (j
(200) "j1*T(DE "&`P RDEth * KF?&@ E(* P (9G+"j#/'!"& E; } \(S `13"&%& LDT? $ *13#TT R 8
(201) q y RT%+rT1*(33D./SD :(31
(202) Dv"0u;(j 5(9#/ "!P+ * ~"9P{
(203) ? *>P{ RDvth * !DE; * 7 YDvth"> "#T'$!"j v; e
(204) FA?&@ E."& P+(3:A(SVDv;. (* "&# w "9 R"*D 8T! εU!(9# # *# y5A"&b εU|ThDE;+ Y R!"r| ?{ *#2"3AT&%erE(9K # T(3#y 54¡3qLK1*"0h (9 : *D"! @P 7 *1
(205) D: \thwD; "9@jD D; *
(206) % +"6Dv'&{ * .(S K ? >; w%; brE"& \ 8(9 :#T'!"& 7 xD 4)WW+ R l
(207) qB$@ 1;!(*!/SD :(31
(208) Dv"0K;0(& \(3# 7"PA * !r:"9@+ hr"& b#T(8Dv Y"0$ 7 RD "& :(9&1` ! jDv(&$t Dv;K(3!/01* R (j R P'elq $r:"&# KT>;0"0e"&*'%+"3@A \y5{"&b εUw? 23@+(3# \thLDv;Dv; *
(209) 5N4 R8
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(211) D\(j \(` :"*DE 1
(212) DT? rE(91
(213) DE"& .T!y 54 mAqLdd& R Rq&y "*DE 3D:T(9#GT3D 8 (31
(214) DET"&>P{ RDEt` * VDv; 6 7 `N4 FA?&@ E.(S
(215) P (3E.(S N4. β. α. EA 5fb4 "&#T'!"& E$; R by`#T@ E\ "#/'!"& E; } (3KDv; h R~"&r ? *# "9A$%uyElz! *. (*m
(216) W}th"&! T 1*#w@{$ *K(j .1*(& R * R;(j I(3# R"`PA * ! : 7{"& Dv * R 8X P@&DNThD;+/ I1*(j R 0">T 7A 3DN(&
(217) R"&13T(8Dv"0!13"9@{#2`P{ rE"9@ Aq`HK 13 *KD; c ED 13 "jro13 DE(9T "$#T'!"& E;+/ ! "raN4 1*"9@A# 13"0SDv :wP@jDE )D"( Rw% Tr1*(93DE#T' w 1* : *(& R * @{ R1* 7jDwP # LD'a"rth"&! ~D"eDv; KPA *r19T(3#.E"& 5;(3 Kr@{# z 8rEr *1
(218) D: K"&r * EDE "&%
(219) |( "
(220) 7P+ # LD'atz;021*;m;(j z "*D' xDP *
(221) c${# "& *YD;+"j R"9@%;+#T'Gq B.
(222)
(223)
(224) . b EDv :"9% 1*(3 !"$3@A#T(8D
(225) @+ :"*DE :(S R!LD DE aD@+ E E"&?+ 8 «P'
(226) ? :(9#C! *1*;(9/ e
(227) T? "$#?+T% 7@ R"*DE :(S$ x! D2D: '3D; * 7/ 7 8@&DE(* *$ : *# (j 7 6$ : 13 7jD"& N RLDE' $ : 13 7jD"j |3D 8 (31
(228) DE2"& *'$V(D:T15 : %&@A#(DET"& "& %+ 5 c : *
(229) 7 "9Gq&_a"& : 13Tr:T1*(&#T#T' 6 vDE :"9% ¡1*(9 @${ : %&@A#(DE 1S; "#TT % 1(31
(230) Dv?LDv' P+' / 9@ 1
(231) D:T"&"&r1*;+"# w (913 RDE'+# Dv :(* Rr 8 (j (S (31* RDE'# 1S; "&# T K * ED 8 (j R Kw!Dv; P+(j R(9#krE"& R 7P :(9 (*u/01* R *(& Rw%!D; `; T%$;^(9rErET LD'~1;+"# T z@jDv(* `t; 1S;u/ (! : 7%&@{#2(8DE"& E' :(
(232) DE «# T!LD:w% ED
(233) T(313 RD' #T1S; "# T+ - '3Dv; 6 7 T W KW|
(234) q b ED :"9% 1*(*(3#
(235) "T1* R 8(&
(236) aDp; ) :(DE )"&rm % :(9&(8DET"&"r.
(237) !"& "&(9!T "9c &(j 7 )_ay
(238) 5:Vt; T1*;"1*1@ R mT-Dvth" R"&r:"& }
(239) Vy(* ]`(* !
(240) DE(*P+"# T3 * o1*(8DE 1*; "$#(9!T+ 6
(241) Q¤ Â R :"*D:"0 TEd2H`fb:&"9 (9!T y:+"j (3& : (9# T !2<KyhIthLDE;m& rEr: R 3DY @$P{ EDE E(DE ` : r: R 13 * TWW
(242) qAeDv; }V(913( 9@A 6 "& >!"&3Dv;a"jrz 6 EDv :"&% m : 7A#(913 ! jD5(9#T"& *1* : (j 7 *D;
(243) c$ R 6
(244) "&a"&r5_my pyhP@&D0"*D5]|TuDE; > :(S; !&@A1*# 7@{
(245) t; R 8(&
(246) TC;'&{"*D;(9#(9!T1&@ 19# *PA"6Dv;~_oy
(247) y (*+~]K4b<}y th 8 R > *1* : (j 7 * T W WU
(248) qAv"& R R(3#N (*;+ 6N x; "& D: HAL author manuscript. !"&3DE;:{P@jD|"6D\# "0% bEd>!"0SDp;$ 7.TDE 8 6 vDE :"&%+ ^ 7cA" @ : z *1* : (j 7 6 _=y zy+ :"*D: */e 7c R 6 7 R "& A(SK "&% * ED: R"9{ @+# ! 3DE(8DE2"&Ot(& >tLDE; "9@jD} 8rEr *1
(249) D
(250) qNv 13"0SDv :(j vDpI_my
(251) ] :"*D *TOt(j }@(3rEr *1
(252) D: *=P ') * EDE :"0% (9# "& *\P@&DhLD`t(& T1* R 8(& R *KthLDv;e ; "& DN"& \# "9%{TD 8 :"0% 6 vD: :"&+ x@A# ! jDE(DET"& T WU
(253) q&fN; Trr: x 3DET(9#G 7c R 6 7 R "&"jrD; ` % :(9&T% *'$! 6 /Dv; 6 7 13 C"&rV vD: :"20 @$%% * ED: >(m1*"&KA#
(254) c)19"&KP T(8Dv "&)"&r`% DE :(S$ 1* :wjDET"& \+"0 vD:TDE :(S R1* :wjDE2"&(3#. inserm-00243083, version 1. :"&19 6
(255) T% (* @PA EDE :(DE sr: * *9P (91! *1*;(9 } t; T1S;K(3'+#(9'( :"&# sT 7 8 R"*DE"&+T"& 1*(DE *1*; "&# (S!T. 7@ :"*DE (* R!
(256) "&K(* z; 19 6$K""`(* `13"9%+ D "9!T`;@+V(S$ Sq D; ! &P (3w!
(257) :"*DE"&+ 8 % T1 ' ED }+ * EDE "&% ! \T`{# T19(8DE KTD; 5w 9@{1
(258) DE2"&V"$r{Dv E'&jDE"9+;0(Sh;$' & :"9c '+#(& R *(S YDp;{ : %&@ #T(8Dv "&!"&rNd2Hhf~Dv E(* "& Dv 8 R \(* Kd2Hhfo : 13 7jD:"& b x@PjD2'&{ * T WU
(259) q9b EDE "9% ! : %&@A#(DE * D@ E+"&? N"&r zyTK;'&A"*D; (3#T(S! 1 &@ 13# *\(*+~Tm+T% :" EDE (DE(3#\(*C! *
(260) "$#/KP{T1 y '$ vD: ! `(90m1; R"& 1u * EDE "9% (&&!/ EDE :(DET"9a *1* : (j 7 6 K(* zW : 13 7jD"j 51*"&19 SDv :(8Dv"0$ SqAb EDE "9% ~(3# 7"urE(31*2# LDE(8DE * z(*` ; RDE(9!T *"& Y(S "&!"& E$;0/ 3E EDETK@A#T(8DE * y *# (j 7 jqB$' EDE K(8D 1 : ? tV "rND; K *rEr *1
(261) D: 5"r 6 EDv :"&% ;0(S? K13"9 1*#w@A *KDE;(D. 6 vDE :"&%+ $ z ;(*+13 `(3& R 8 v% T1K(31
(262) DE/? LDE'P'~/01* R (j R/%!D' :"
(263) /{ ;' & :"9c+'#(&
(264) C(31
(265) DvT? LDE' rE(31*2# LDE(8DEw%O<Ky R 8#2 8(&
(266) *(* )/019 : (j Rw%%(3#T(*+T)K4b<}y T R"0! m @ "& tz; T1*;V(S'-(3rEr 81D "& :(9j E% T1.D:"0 .P'K R *9@ 13T&%<Ky R
(267) @$jDE(*+ jq9[A@ Dp;0 !"& x *$ 6 EDv :"&% DE 8(DE! 3DADE (* RT 3D:#T'K"&tz$ R %&@A#(DE * yh]Iy (S ]. :(9 ` 8 T? *<>
(268) @ R"*DE "3$; T1[G(31
(269) Dv"j a]<K[GC(31
(270) Dw? LDE'TT;+P+ D"& E'T3D 8 v @ :"0$ SqLDv; T Dv; «% #w@jDE(9K(8DE : *19
(271) jD"& rE(*! #/'$9<!p! xDv;'#h(& (3 Dv(
(272) DE YT<K_ y : 13 &DE"& x
(273) 0y_aFAy : 13 7jD"& x \(*z(3T LD *13 7jDE"& x .(& |t *#2# (& . Rw%V(z : *13 7jDE"& R R ;(*? (3#
(274) "zP * ! !"0{ EDE (8DE YDv"PA Y 7
(275) 7LDET? \D:"K EDE :"2!"9@A#T(8DE "0 T W d
(276) q T?+ =Dp; Cth C :(S&% m"&r! r:r: 1
(277) D: "$r! EDE :" ;0"& !"& * "3«
(278) @+ :"*DE :(S R!LD D 8 ' EDE e
(279) (S0O(9#/ ")"&O 7@ :"&(&# "& E%(* T*(DE "&(S '(*jDv"3%+ 3
(280)
(281) eh@A#LDEw{# @ "3+ :"*D: *1
(282) DvT? *rEr *1
(283) D: «1*"3@A# PA 7cA *1
(284) D: *¡"&19"&%LD:/? r@ 1DET"&. 2 "*DE(*P+#T'~(8D2DE 3DE "&(3#|! *1*;(9 }
(285) # 8(9 E /%~(*+u! S!"j ':|#"$13"&!"*D"j 5(91
(286) DEw? LDE' (9w
(287) 7LD:w? LDv'$(* ?&@ #T :(SP # LD'uDE"
(288) A # 7{ ' E 7 * !r"& Y ? th T WW W W l E
(289) qG,."9% LDE/? !r@+1
(290) D: "00/%~(9Cy ¡K(9'uPA !!"$3@A#T(8DE *>DE; :"9@+%;~(913 RDE'# 1S; "#TT. (*C D 5A" RLD:T?+ K :"# "9a '(*jDv"&% 3 R `TDE; }; wA"j1*(SK@{
(291) kt; T1*;C;(j z(!+T?"6Dv(3#N R"&# Tm 819#T(3 :(DEw? K! S!"& :'TW j W X
(292) q5]\'!"9@ #T(8DET$%-D; O(31
(293) DE/? LDE'"&r!Dv; O13 R
(294) PA *#T#@+ (*)D; O+T% " EDE (9#K(*! 3 7"&# TKP 21)"9 (9!T E% T1- x'$ DE e
(295) 6 vDE :"&%+ $ Z;(*? 5 *rEr: *1
(296) D \"&K "& V(9# (S h(SP "& V(9# #2"j19"&!"&DE"& N(31
(297) DEw? LD'GqSHhT%;!#2 ? *# ."&rN * EDE :"0% !
(298) c+(313 EP+(8DE 5 'KjDE"0} I"r FG(* v0/$ "&k 5&
(299) (j 7 *Atz; 8 R (j # "&t# ?+ *#/ rE(913 #TLDE(8Dv Kj"9 (9!T+ 8 % T1`r@ 1
(300) Dv "&Gq[T(3#T#T' +P+'!"&@ #T(8DvT%VDv; h 7
(301) LDE/? D'u"&r W 819
(302) jD"& x \wD; .P+ :(3w + * EDE :"9% \(S (3 GDE"z;(*? TKA"j DE(*3D 7@ "j#2 jDET1j# / 5 *rEr: 1
(303) D
(304) q.
(305) KDE; `"*DE;+ 8 I;0(S + "&% * ED 8 "& (91
(306) D: (j (`th (3(3# #2"0 vDE * :T1`!"9@A#(DE"& ."&r y].y (SP T# LDE'KD:"T1* 8(&
(307) `1; # "& :2. 1*"& 9@ 1
(308) DE(S 13 `D; R"3@$%;^Dv; y].y^ R 13 jDv"j z1*"&K{#2
(309) cot; T1*;m13"&r 8 R `"&~+ :"&% * ED 8 "& >(* mLD: V! xDE(SP "#TLD: * `(*m(31
(310) Dv "& RT! #T(9 !DE"OD;(8D"&r>P{ 3"$T(33 7 T+ 6 C(9 «P (3 vP LD@ E(DE 6 C(SV(9'13"&3DE P$@jD: CDE"LD: C(*3DE13"&?&@ # R(S3D> :"9A 8 DE *
(311) q FG "3%+ 6 ED :"& o(* :"&%+ 6 EDT1*"&KA"3@+ (3# "&+ & W " w13"&KP /0(DE "&thLDv; * EDE "&% >(3 : o(3# 7")1*(* (SP # o"r HAL author manuscript. ;(* 1*/%Dp; zy ' ED ^q .
(312)
(313)
(314) . b EDv :"9% );(j ^(m
(315) @+ :"9 :"*D *1
(316) DET? m *rEr *1
(317) DK"&-"*Dv; e *1*2rE1=! *1*;(9/ e *#(DE *mD:"O19"&%LD:T?+ m&' Rr@ 1
(318) DT"& .P+'. inserm-00243083, version 1. P # "&1*+$%=Dp; ~(91
(319) DE "&)"&rK+
(320) @ "*D:"9c 1C(*%+ SD "& !T&; wPA DvT%Dv; *T `% :(DE "&Gq| Dh1*(9O "*DE(SP #w' + :"*D: 1
(321) DK1@ #LD@+ : *O @+ :"9. (*%(3w EDDv; +
(322) @ "*D:"9cA1K *rEr *1
(323) D: "rD; K 7c 13LDE(DE"& E' 7@ R"6Dv :(* VLD D .Mb% #w@jDv(*K(8D &U . "& ."rDE;+ h *
(324) 7 SDT(3#13"& EDELD@A 3D."&r R #2 5{#T( 3@A * 5TyhD;+ h(9K'# " . β. y β|A
(325) jDv *Atz; 1;u r"& :! *KD; "3@+%;KDE; `T 9@ 1
(326) D "9~"&r."9c &(8DvT? K EDE : *
(327) $ d
(328) q. fN; C(*P 2# LD'-"&rm8l β E * EDE :(9&"$#ZD"OT R 81
(329) DE#/'« 1*(3?+ % C; T%$; #T') : (31
(330) DE/? mrE : * C :(9T1*(3#
(331) (*mDE;&@{ ; *#
(332) @+ :"& (S%0(9T ED
(333) cA"&% +"3@A \"3cA &(8DvT? /$ x@{#LD: \(S0z 7@ R"90(&#+(* "9jDv"0 / 7 \9@{ \DE"5D; .+ x * 7 019 5"jrAD;+ .+; "#T 1y :T%
(334) q f|;0 ^% # T(3#.19 *# # !"&rDp; P (3wa(3 VDv(3 E% RD: Kr"& V EDv 8 " m;+"j v!"& 3
(335) \(Sm! *T(8D uTrEr 8 R 3Dz; "& :!"00(9# rEr: *1
(336) D: }T~Dv;. : %&@ #T(8Dv "&u"&rkDv; ` ? *# "9+! jD\(* > #T(j vDET13LDE'u"&r. { 13Tr 1+ @ :"&(9#AA"3$@ #T(8DE "0$ (S TVDv; `1*"&3DE :"#"&r. "$1* :T ` ? 3DR E R * r:"& ? t T WE
(337) q # T(3#.19 *#T#/ } c 6
(338) `(^j@+KPA 8 "$r5 :"*DE */$ 5Dp;0(D51*(*CP{ %j@A#T(8DE *P+'a 6 EDv :"&% }T13#w@{$$%ay5{"&b t; T1*; a/hA# T1*(8DE *T! SKP :(S Or:"& V(DE "&(S EDE v@ 1
(339) D@ E(9#` #T(j EDv19LD' `(* % # T(3#KrP+ : #T#T(3 v'(31* T1« :"*D 8/tz; "
(340)
(341) c$ R 6
(342) "&uT1* R 8(&
(343) * thLD;P (3we(9%T%Gq\ EDE :"0% 5T1* R *(j R 19 8 R 7P :(9#% #w@ 13" 7 5@jD #T *(8DET"& {{"
(344) RwPA#T'!P+'uT 9@A1*T%u% #w@ 19"
(345) DE E(* +"j DE muw=Dp; m "*Dv; *#T(
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(346) @ "TK(*% /%} vD@ * \;(S? (9#/ 7"K !"& EDv :(8Dv *YD;+ :"$# 5"jr| 6 EDv :"&% !/!T1* : (j RT+% P "*D; R % "&(9#!13 R
(347) P+ :(3#}P #T""r:# "&t (*% #w@ 13" 7 )! RD(*P "&# . lEX
(348) q5y. *13 3DC : ? t "r 7@ :"/V(9% T$% ED@+j2 3. 1*"& 1*#w@A *~D;(Dh * EDE "&% O : 7{#T(319 ! jDDv; :(S' N4f\5K(9' *1* R (j R P+ :(3wot; LD: ^V(D D: K# *
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