• Aucun résultat trouvé

Influence de Toxoplasma Gondii dans la régulation d'UHRF1 via la voie NF-KB

N/A
N/A
Protected

Academic year: 2021

Partager "Influence de Toxoplasma Gondii dans la régulation d'UHRF1 via la voie NF-KB"

Copied!
155
0
0

Texte intégral

(1)

HAL Id: tel-01251696

https://tel.archives-ouvertes.fr/tel-01251696

Submitted on 6 Jan 2016

HAL is a multi-disciplinary open access archive for the deposit and dissemination of sci-entific research documents, whether they are pub-lished or not. The documents may come from teaching and research institutions in France or abroad, or from public or private research centers.

L’archive ouverte pluridisciplinaire HAL, est destinée au dépôt et à la diffusion de documents scientifiques de niveau recherche, publiés ou non, émanant des établissements d’enseignement et de recherche français ou étrangers, des laboratoires publics ou privés.

Influence de Toxoplasma Gondii dans la régulation

d’UHRF1 via la voie NF-KB

Ghaidaa Kanjo

To cite this version:

Ghaidaa Kanjo. Influence de Toxoplasma Gondii dans la régulation d’UHRF1 via la voie NF-KB. Parasitologie. Université de Strasbourg, 2014. Français. �NNT : 2014STRAJ068�. �tel-01251696�

(2)

ϭ

ÉCOLE DOCTORALE des Sciences de la Vie et de la Santé

!"#$%&'()*(+)%",(-#.,%/"+)01,(+)2#,0/34"(5)67)89:9)

;<=>6

)

)

!"#$%&'#% !(" )

?0#%*##)@7ABC)

*+,'%&,% -% ) DE)>(2,($F-()9EGH ) ) .+," +/'%&0" -% 1"(2% 2% )

/.,('-)*()IJ'"%K(-+%,L)*()>,-#+F/'-3

30$40!-0&%

) *40%&4%$ 2, 505(&' 6 $!#40(-0'# ) .("($0'+-+10%

MANOP6AQ6 ) 6)TOXOPLASMA)GONDII)

7A>)O7)R6?PO7;MCA) JP<RNG)VIA)

O7)SCM6)AN5κ

κ

κT)

κ

) ) ) ;<=>6) MRM?66)U7R)V) U-W)Q7A CONM)6-$#""/ %-(.,('-)*(),04+(X)'"%K(-+%,L)*()>,-#+F/'-3) R7UUCR;6PR>)V) U-W))SMOO6A7)M+#F(II())))))))))))))))))))))))R#22/-,('-)6Y,(-"(X)P"%K(-+%,L)*()R(%$+ -W)7BZ6AT6R?) #"%(I) R#22/-,('-)6Y,(-"(X)P"%K(-+%,L)*()O%$/3(+

6Y#$%"#,('-)M",(-"()V))

U-W)>;COO5@6OO6R)N-#"[/%+( 6Y#$%"#,('-)M",(-"(X)P"%K(-+%,L)*()>,-#+F/'-3)

(3)

Ϯ

R6]6RQM6]6A;>

)

7 -80$$,% 2% -( "#2(4'0+& 2% 4% '"(5(0-9 :% '0%&$ ; %<!"0=%" =%$ "%=%"40%=%&'$ ; '+,'%$ -%$ !%"$+&&%$ >,0 =?+&' (02# !+," "#(-0$%" 4% '"(5(0- 2% '@A$% %' =?+&' !%"=0$ 2% !"+1"%$$%" %' 2% B0&0" 4%''% "%4@%"4@%C

D% '0%&$ '+,' 2?(/+"2 ; "%=%"40%" =+& 20"%4'%," 2% '@A$% ."C E"=(&&+ F7G3HIJK9 !+," =8(5+0" B(0' 4+&B0(&4% =(-1"# -%$ 4+&&(0$$(&4%$ !-,'L' -#1A"%$ >,% :8(5(0$ %& (""05(&' 2(&$ $+& #>,0!% 2% "%4@%"4@%9 !,0$ !+," =8(5+0" 1,02#9 %&4+,"(1#9 4+&$%0--#9 B(0' /%(,4+,! 2?%BB+"' !%&2(&' -( 2,"#% 2% =( '@A$% !+," '%"=0&%" 4% '"(5(0-C D8(0=%"(0$ #1(-%=%&' -,0 20"% ; >,%- !+0&' :?(0 (!!"#40# $%$ >,(-0'#$ @,=(0&%$ 28#4+,'% %' 2% 4+=!"#@%&$0+& '+,' (, -+&1 2% 4%''% '@A$%C

D% '0%&$ ; %<!"0=%" =%$ !-,$ 50B$ "%=%"40%=%&'$ (, 3"C D,-0% MNOGEP !+," =?(5+0" %&4(2"# !%&2(&' -( 2,"#% 2% =( '@A$%C *( 4+=!#'%&4%9 $( "01,%," $40%&'0B0>,% %' $( 4-(0"5+Q(&4% =?+&' /%(,4+,! (!!"0$C

D?%<!"0=% '+,$ =%$ "%=%"40%=%&'$ ; -?%&$%=/-% 2%$ =%=/"%$ 2% =+& :,"Q 2% '@A$%)

U-W)S%II("#)M+#F(II(X)>,0 =8( B(0' -8@+&&%," 2?%<(=0&%" 4%''% '@A$%9 %' :% -,0 %& "%=%"40%9 2% =R=% >,% !+," $( !("'040!('0+& (, D,"QC)

U-W)>,/II5@(II(-)N-#"[/%+(9 >,0 ( (44%!'# 2% :,1%" 4%''% '@A$%9 %' :% -,0 %& "%=%"40%9 2%

=R=% >,% !+," $( !("'040!('0+& (, D,"QC)

-W)7^_("F(-3) #"%(IX)>,0 ( (44%!'# 2% :,1%" 4%''% '@A$%9 %' :% -,0 %& "%=%"40%9 2% =R=% >,% !+," $( !("'040!('0+& (, D,"QC))

(4)

ϯ D?(2"%$$% '+,'% =( 1"('0',2% ; '+,$ =%$ (=0S%T$ %' ; '+,'%$ -%$ !%"$+&&%$ >,0 =?+&' (02# 2(&$ -( "#(-0$('0+& 2% 4% '"(5(0-C D% !%&$% !("'04,-0A"%=%&' ; 7-%<(&2%"9 E$'#"0&(9 IQ20(9 F("+-0&%9 F#40-%9 E-0$%9 *Q-50%9 N+--(&2 %' M",&+ !+," -%,"$ 4+&$%0-$ %' -%," 4+--(/+"('0+& (0&$0 >,% -%," 20$!+&0/0-0'# ; -?#1("2 2% =%$ '"(5(,<C D% '0%&$ ; "%=%"40%" #1(-%=%&' E-0$(/%'@9 U("0% >,0 =?+&' (44+=!(1&# 2,"(&' -%$ 2%,< !"%=0A"%$ (&&#%$ 2% =( '@A$% !+," -%," 1%&'0--%$$% %' /+& @,=%,"C

D?(2"%$$% '+,'% =( 1"('0',2% ; '+,$ =%$ (=0S%T$ V W+=(=9 N(=%X9 N+,/(9 Y+, 7-B(Z("9 7-09 7/0"9 [($$%" %' \(-(( !+," -%," $+,'0%& %' -%," #4+,'%C

E&B0&9 >,?0- =% $+0' !%"=0$ 2% 2#20%" 4% =#=+0"% (,< =%=/"%$ 2% =( B(=0--% %' ; =%$ (=0$ >,0 =?+&' $+,'%&,% 2(&$ -( "#(-0$('0+& 2% 4%''% '@A$%C D% '0%&$ ; "%=%"40%" '+,' !("'04,-0A"%=%&' =%$ !("%&'$9 =%$ B"A"% SK=(29 U+2(" %' W+,$$(=T9 =%$ $],"$ SK-@(= %' I(&(T9 =%$ &0A4%$ S\(B((9 *+,X(&9 30(&( %' U(0$T9 =%$ &%5%,< S^("(=9 W($$(& %' _@(0'@T !+," -%," $+,'0%& %' -%," #4+,'%C

` =+& !(Q$ -( *Q"0%9 =(-1"# -( 1,%""%9 -( =+"' %' -( $+,BB"(&4%9 ', "%$'%"($ -( !-,$ /%--% "+$% 2(&$ -% =+&2%C

(5)
(6)

ϱ

>C]]7MR6)

>C]]7MR6)

a

7TR6SM7;MCA>))

b

OM>;6) 6>)NM?PR6>))

cc

MA;RC PQ;MCA)

cd M) Q<7UM;R6)M)V)TOXOPLASMA GONDII)6;)O7);C`CUO7>]C>6) ce ) G)

Toxoplasma gondii))

ce cCc

I%$ 20BB#"%&'$ $'(2%$ !("($0'(0"%$

cf cCcCc P(4@QX+g'%$ ca cCcCd M"(2QX+g'%$ ca cCcCe *!+"+X+g'% ch cCd

H"1(&0$('0+& =+-#4,-(0"%

ch cCdCc I%$ =04"+&A=%$ ci cCdCd I%$ "@+!'"0%$ ci

cCdCe I%$ 1"(&,-%$ 2%&$%$ ci

cCe _#&+'Q!%$ %' 50",-%&4%

dj

cCeCc _#&+'Q!% K dc

cCeCd _#&+'Q!% KK dc

(7)

ϲ

cCeCf _#&+'Q!%$ "%4+=/0&(&'$ +, (5%4 (--A-%$ ('Q!0>,%$ dd

cCeCa J(4'%,"$ !("($0'(0"%$ -0#$ ; -( 50",-%&4% da

cCf FQ4-% /0+-+10>,% 2% !" #$%&''

dk cCfCc FQ4-% ($%<,# dk cCfCd FQ4-% $%<,# dk

cCaC F+&'(=0&('0+&

dh ) 9)

O#),/Y/2I#+$/+()

di dCc

E!02#=0+-+10%

di dCd

I( '+<+!-($=+$% @,=(0&%

ec

dCe P"(0'%=%&'$

ed MM)

Q0#2%,-()MMV)T. gondii (,)I()a#.,('-)*(),-#"+.-%2,%/")AN5κ

κ

κ

κT)

ee G)

O()a#.,('-)*(),-#"+.-%2,%/")AN5κ

κ

κ

κT))

ee 9)

>,-'.,'-()*())I#)a#$%II()AN5κ

κ

κ

κTbR(I)

eh D)

O#)a#$%II()*(+)2-/,L%"(+)Mκ

κ

κ

κT))

fj H)

M",(-#.,%/")ANκ

κ

κ

κTb)Mκ

κ

κ

κT))

fc c)

O()./$2I(Y()M@@)dMκ

κ

κT)e%"#+(f)

κ

fe g)

RL3'I#,%/")*()IJ(Y2-(++%/")*(+)34"(+)2#-)AN5κ

κ

κ

κT))

fa

(8)

ϳ

8)

O#)K/%()*()+%3"#I%+#,%/")*()AN5κ

κ

κ

κT)

fh

h)

R1I()*()AN5κ

κ

κ

κT)*#"+)Ii%"aI#$$#,%/"))

ac

:)

O#)K/%()AN5κ

κ

κT)(,))IJ%"a(.,%/")2#-)T.gondii)

κ

ae

GE)

O#)-L3'I#,%/")*%aaL-(",%(II()*()AN5κ

κ

κT)(")a/".,%/")*()I#)K%-'I(".()*(+)

κ

+/'.0(+)*()T.gondii

ai GG)

M",(-#.,%/"+)*()*%aaL-(",+)a#.,('-+)2#-#+%,#%-(+)#K(.)I#)K/%()*()

+%3"#I%+#,%/")*()AN5κ

κ

κ

κT)

ke MMM

Q0#2%,-()MMM)V)Toxoplasma gondii)(,)I()a#.,('-)*(),-#"+.-%2,%/")

P<RNG

kh )G)

O()a#.,('-)*(),-#"+.-%2,%/")P<RNG)

kh )9)

N/".,%/"+)*JP<RNG)

ki dCc RL3'I#,%/")*').!.I().(II'I#%-( ki dCdC 62%3L"L,%&'(

)

hc dCe Q#".(-+

)

hd dCf OJ%"a(.,%/")2#-)Toxoplasma gondii) hf

CTB6Q;MN>)6;)]Cj6A>)

hk

7-,%.I(V

 ij ) )

>!",04+()(")a-#"[#%+)*()IJ#-,%.I()k)+/'$(,,-())

ij

(9)

ϴ

7-,%.I()

ia

QCAQOP>MCA)6;)U6R>U6Q;MS6>))

cca

TMTOMC?R7U<M6

cdb

(10)

ϵ

OM>;6) 6>)7TR6SM7;MCA>)

7 AV 7402% 3#$+<Q"0/+G,4-#0>,%

7N>>7)V 71%&4% J"(&l(0$% 2% *#4,"0'# *(&0'(0"% 2%$

7-0=%&'$

QQRc)V I% "#4%!'%," ; FmF 4@0=0+Z0&% 2% 'Q!% a Q*eV FQ4-0&% 2%!%&2(&'% Z0&($%$

Q0MUV F@"+=('0&% K==,&+!"%40!0'('0+& .5M7UV F%--,-(" K&@0/0'+"$ +B 7!+!'+$0$ Q@MV F2Z K&@0/0'+"

QjO V FQ-0&2"+=('+$0$ S',"/(& ',=+" $Q&2"+=%T OV 3+$% I#'(-%

A];GV 3G7 =#'@Q-'"(&$B#"($% c 6 ;7)V E'@Q-A&% 30(=0&% P#'"( 74#'0>,% <D@:];V @0$'+&% We -Q$0&% b =%'@Q-'"(&$B%"($% <7;V W0$'+&% 74%'Q- P"(&$B%"($% < 7QV W0$'+&%$ 3%$(4#'Q-($%$ < 7QGV @0$'+&% 2#$(4#'Q-($% c <MNGV WQ!+<0( K&2,4'0/-% J(4'+" c M7U+V K&@0/0'+" +B (!+!'+$0$ MQ7]5GV 0&'"(4%--,-(" 4%-- (2@%$0+& =+-%4,-% c MNA5γ:γ:γ:γ: 0&'%"B%"+& 1(==( M@@V KκM ^0&($% MO5GV K&'%"-%,Z0&%c MO5G9V K&'%"-%,Z0&%cd MO5gV K&'%"-%,Z0&%k MO5hV K&'%"-%,Z0&% i M]QV K&&%" U%=/"(&% F+=!-%< MRN5GV K&'%"B%"+& "%1,-('+"Q B(4'+" c MR?+V KJGȖm0&2,40/-% K==,&0'QmN%-('%2 _P.($%$ κκTV K&@0/0'+" +B GJmκM κ e #V Z0-+2(-'+& @CV ^&+4Z +,' OU>V I0!+!+-Qm$(44@("02%$ O;FV IQ=!@+'+<0& / N%4%!'+" O'.)V I,40B%"($%

]QU5G)V U+&+4Q'% 4@%=+(''"(4'(&' !"+'%0& +, FFId ]??) U(Qm_",&n(-2m_0%=$(

]SUV U%=/"(&% 2% -( o(4,+-% .("($0'+!@+"%

]! hhV UQ%-+02 20BB%"%&'0('0+& !"0=("Q "%$!+&$% 1%&% SiiT A6]CV GJmκM E$$%&'0(- U+2,-('+"

AN7MUDV -87dj 2#$,/0>,0'0&('0+& %&XQ=% AN5κΒκΒκΒκΒV G,4-%(" B(4'+"−κΒ

AM@V GJmκMm0&2,40&1 Z0&($% A@V G(',"(- ^0--%"

AO>V *01&(- 2% -+4(-0$('0+& &,4-#(0"% AC)V U+&+<Q2% 2?(X+'%

U7]UV =+'0B$ =+-#4,-(0"%$ 0$$,$ 2% =04"+m+"1(&0$=%$

!('@+1A&%$

(11)

ϭϬ

UQA7V ."+-0B%"('0&1 F%-- G,4-%(" 7&'01%& U<V !+'%&'0%- @Q2"+1A&%

U< V .-(&' W+=%+ 3+=(0& URTV ."+'%0&% +B N%'0&+/-($'+=(

.NN) "#4%!'%,"$ 2% "%4+&&(0$$(&4% 2% -( $,"B(4% 4%--,-(0"%

RNOUV N%$'"04'0+& J"(1=%&'$ I%&1@' .+-Q=+"!@0$= R< V N%- @+=+-+1Q 2+=(0&

RMA?V N%(-Q K&'%"%$'0&1 G%n _%&% >AUV $0&1-%m&,4-#+'02% !+-Q=+"!@0$=%TC >R7V *%' (&2 N0&1 7$$+40('%2

>;7;V *01&(- P"(&$2,4%"$ (&2 74'05('+"$ +B P"(&$4"0!'0+& ;7 V I% 2+=(0&% 2% '"(&$(4'05('0+& 2% -( '"(&$4"0!'0+& ;0GV P @%-!%" 2% 'Q!% c ;ORV P+--m-0Z% "%4%!'+" ;AN5ĮV P,=+" G%4"+$0$ J(4'+" (-!@( ;R7NG)(,)9V PGJm"%4%!'+"m($$+40('%2 B(4'+"$ c (&2 d ;RMNV PKNm2+=(0&m4+&'(0&0&1 (2(!'%"m0&2,40&1 0&'%"B%"+&mȕ ;; )V ,& 2+=(0&% P(2%= P,2+" ;SAV "#$%(,< ',/,-+m5#$04,-(0"%

P<RNGV O/0>,0'0&m-0Z% 4+&'(0&0&1 .W3 (&2 N0&1

J0&1%" 2+=(0&$9 c

SM<)V o0",$ 2% -?K==,&+2#B040%&4% W,=(0&% SMR)V o0",-%&' -+40

SU)V o(4,+-% .("($0'+!@+"% SU)V o(4,+-% .("($0'+!@+"%)

(12)

ϭϭ

OM>;6) 6>)NM?PR6>)6;);7TO67P`

N%3'-()G)V) ()*%$&+,)-./0 #/%&' ce

N%3'-()9)V) .+$0'0+& '(<+&+=0>,% 2% !" #$%&'' cf

N%3'-()D)V) *'",4',"% 2?,& '(4@QX+g'% 2% !" #$%&'' ck

N%3'-()H)V) ^Q$'% '0$$,-(0"% "%&B%"=(&' 2%$ /"(2QX+g'%$ 2% !" #$%&'' ck

N%3'-()c)V) H+4Q$'%$ 2% !" #$%&'' ch

N%3'-()g)V) ."#$%&'('0+& $4@#=('0>,% 2%$ 4-(2%$ 2% !"#$%&'' df

N%3'-()8)V) FQ4-% !("($0'(0"% 2% !" #$%&'' di

N%3'-()h)V) ."#5(-%&4% 2% -( '+<+!-($=+$% %& J"(&4% ej

N%3'-():)V) U%=/"% 2% -( B(=0--% GJmκM ei

N%3'-()GE)V) U%=/"% 2, F-($$% K 2% -( B(=0--% GJmκM eb

N%3'-()GG)V) U%=/"% 2% -( B(=0--% KκM fc

N%3'-()G9)V) O& =+2A-% 2% -( 2#1"(2('0+& 2%$ 5+0%$ 2% "#1,-('0+& KκMα 2(&$ -%$ 4%--,-%$ /($(-%$ %' $'0=,-#%$

fd

N%3'-()GD)V) o+0% 2% $01&(-0$('0+& 2% GJκM %' -?(4'05('0+& 2, 4+=!-%<% K^^ fa

N%3'-()GH)V) *'0=,-0 %' 1A&%$ 40/-%$ 2% GJmκM fk

N%3'-()Gc )) o+0% 4-($$0>,% %' 5+0% (-'%"&('05% 2% GJmκM fi

N%3'-()Gg)V) U+2A-% "%!"#$%&'(&' -( =+2,-('0+& 2% -?%<!"%$$0+& 2% GJmκM 2(&$ -%$ 4%--,-%$ 0&B%4'#%$ ah

N%3'-()G8)V))))NL-% B+&4'0+&&%- 2%$ 2+=(0&%$ 2? OWNJc kh

N%3'-()Gh)V) N#1,-('0+& 2, 4Q4-% 4%--,-(0"% hj

N%3'-()G:) ."#$%&'('0+& $@#=('0>,% 2%-( !@($% !"#4+4% %' '("205% 2(&$ -%$ 4%--,-%$ 0&B%4'#%$ (5%4 NW %' .",

cdk

N%3'-()9E) ."#$%&'('0+& $@#=('0>,% 2%-( !@($% !"#4+4% %' '("205% 2(&$ -%$ 4%--,-%$ 0&B%4'#%$ (5%4 IEJ cdi

(13)

ϭϮ

)

)

)

MA;RC PQ;MCA)

)

) )

(14)

ϭϯ

Q<7UM;R6)M)V)TOXOPLASMA GONDII)6;)O7);C`CUO7>]C>6)

GW))Toxoplasma gondii))

!$1$2.+03+ #$%&'' %$' ,& !("($0'% 0&'"(4%--,-(0"% +/-01('+0"% >,0 (!!("'0%&' (, !@Q-,= 2%$

7!04+=!-%<(C F% !@Q-,= "%1"+,!% 2% &+=/"%,< (1%&'$ !('@+1A&%$ 2?0&402%&4% =(:%,"% $," -% !-(& =#204(- %' 5#'#"0&(0"% 4+==% -%$ 4.+03$&'/3 "%$!+&$(/-% 2, !(-,20$=%9 -%$

5'3*6'+ "%$!+&$(/-%$ 2% 4+44020+$%$ (50(0"%$C !" #$%&'' ( #'# 0$+-# !+," -( !"%=0A"% B+0$

%& cbji !(" G04+--% %' U(&4%(,< 4@%X ,& "+&1%," 7B"04(0&9 -% 1+&20 S()*%$&+,)-./0

#/%&'TC

)

)

7'#/6* 8 9 ()*%$&+,)-./0 #/%&' :;$)<3+%%= 8>>?@

!" #$%&'' %$' 4(!(/-% 2?0&B%4'%" ,& '"A$ 1"(&2 &+=/"% 2?(&0=(,< ; $(&1 4@(,2 Q 4+=!"0$

(15)

ϭϰ )           7'#/6* A 9 4$0')'$% )+1$%$3'B/* &* !" #$%&'' GWG O(+)*%aaL-(",+)+,#*(+)2#-#+%,#%-(+)) !$1$2.+03+ #$%&'' %<0$'% $+,$ '"+0$ $'(2%$ #5+-,'0B$ 20BB#"%&'$ ) 6$F-#".0($(",) ."+'+X+( U0!I'$) 7!04+=!-%<( QI#++() *!+"+X+( >/'+5.I#++() F+44020( C-*-() E,4+4402(% >/'+5/-*-() E0=%"0+"0&( N#$%II() *("4+4Q$'02(% >/'+5a#$%II() P+<+!-($=('02(% ?("-() P+<+!-($=( 6+24.() 1+&200

(16)

ϭϱ

GWGWGW ;#.0!_/l,(+)

J+"=% 0&'"(4%--,-(0"% 5#1#'('05%9 0- $% !"#$%&'% $+,$ B+"=% 2?,& 4"+0$$(&' 2% k ; i ȝ= 2% -+&1 $," e ; f ȝ= 2% -("1%C K- 0&B%4'% '+,'%$ -%$ 4%--,-%$ &,4-##%$ 2% -?@L'% %' $?Q =,-'0!-0% (, $%0& 2?,&% 5(4,+-% !("($0'+!@+"%C

I( !("'0% (&'#"0%,"% !"#$%&'% ,& 4+=!-%<% (!04(- >,0 4+=!+"'% -% 4+&+g2% %' 2%$ +"1(&%--%$ $#4"#'+0"%$ S=04"+&A=%$9 "@+!'"0%$ %' 1"(&,-%$ 2%&$%$TC

GWGW9W T-#*!_/l,(+))

I%$ /"(2QX+g'%$ "#$,-'%&' 2% -( '"(&$B+"=('0+& 2, $'(2% '(4@QX+g'% $+,$ -?0&B-,%&4% 2% B(4'%,"$ 0==,&$ SKJGm1(==(9 GHT +, =#'(/+-0>,%$ S!WTC K-$ $+&' 4("(4'#"0$'0>,%$ 2% -() !@($% 4@"+&0>,% 2% -?0&B%4'0+&C U+"!@+-+10>,%=%&' '"A$ !"+4@% 2%$ '(4@QX+g'%$9 0-$ $?%& 20$'0&1,%&' !(" ,& =#'(/+-0$=% %' ,&% "#!-04('0+& "(-%&'0$C I%$ /"(2QX+g'%$ $+&' "%1"+,!#$ (, $%0& 2% ZQ$'%$ +p 0-$ $+&' 0&(44%$$0/-%$ (,< 2#B%&$%$ 0==,&0'(0"%$ %' (,< '"(0'%=%&'$ (4',%-$C K-$ $0A1%&' 2(&$ 2%$ +"1(&%$ 40/-%$ ) =,$4-%$ %' -% $Q$'A=% &%"5%,< Q 4+=!"0$ -?]0-C I%$ 4%--,-%$ 0&B%4'#%$ $+&' !"0&40!(-%=%&' -%$ &%,"+&%$9 -%$ ($'"+4Q'%$ %' -%$ 4%--,-%$ =,$4,-(0"%$C

(17)

ϭϲ F+&+g2% 7&&%(, !+-(0"% U04"+&A=% N@+!'"0% _"(&,-% 2%&$% U04"+',/,-%$ U%=/"(&% 0&'%"&% U0'+4@+&2"0% 7!04+!-($'% G+Q(, N#'04,-,= %&2+!-($=0>,% U%=/"(&% !-($=0>,% .L-% !+$'#"0%," F+&+g2% 7&&%(, !+-(0"% U04"+&A=% N@+!'"0% _"(&,-% 2%&$% U04"+',/,-%$ U%=/"(&% 0&'%"&% U0'+4@+&2"0% 7!04+!-($'% N#'04,-,= %&2+!-($=0>,% U%=/"(&% !-($=0>,% .L-% !+$'#"0%," G+Q(,

7'#/6* C 9 D)6/,)/6* &E/% )+,<-F$G)* &* !" #$%&'' :&E+26H0 I+/3 *) +."= AJJ?@

7'#/6* K 9 L-0)* )'00/.+'6* 6*%M*63+%) &*0 N6+&-F$G)*0 &* !" #$%&'' :O6 P"Q" R*'00= S.N*6) 5'%0)*'% ($..*#* $M Q*&','%*"@

(18)

ϭϳ

GWGWDW >2/-/_/l,())

I% $!+"+X+g'% %$' -% "#$,-'(' 2% -( "%!"+2,4'0+& $%<,#% >,0 ( -0%, 2(&$ -%$ 4%--,-%$ 2% -?#!0'@#-0,= 0&'%$'0&(- 2% -?@L'% 2#B0&0'0BC U+"!@+-+10>,%=%&' !%, 20BB#"%&' 2%$ (,'"%$ $'(2%$ 0&B%4'0%,<9 0- %$' 4+&'%&, 2(&$ 2%$ ++4Q$'%$ $!+",-#$ >,0 !%,5%&' $,"505"% $," -% $+- !-,$ 2?,& (& 2(&$ ,& 4-0=(' @,=02%C F?%$' -% $'(2% 2% 20$$#=0&('0+& '%--,"0>,%C

)



7'#/6* T 9 $$,-0)*0 &* !" #$%&'' :<))29UUVVV"&2&",&,"#$WU&2&1U<)3.U'3+#*X.'N6+6-"<)3@



GW9)C-3#"%+#,%/")$/IL.'I#%-())

!$1$2.+03+ #$%&'' %$' ,& !("($0'% %,4("Q+'% ,&04%--,-(0"% @(!-+g2%9 >,0 !"#$%&'% 4%"'(0&%$

4("(4'#"0$'0>,%$ 4+&$%"5#%$ 4@%X '+,$ -%$ %,4("Q+'%$9 =(0$ #1(-%=%&' 2%$ +"1(&0'%$ $!#40B0>,%$ 2% $+& 1"+,!%9 -%$ 7-5%+-('(9 +, 2% $+& !@Q-,=9 -%$ 7!04+=!-%<(C

H,'"% -%$ +"1(&0'%$ 4+==,&$ (,< (,'"%$ 4%--,-%$ %,4("Q+'%$ S&+Q(,9 =0'+4@+&2"0%9 "#'04,-,= %&2+!-($=0>,%qT9 !" #$%&'' !+$$A2% 2%$ +"1(&0'%$ 0=!-0>,#$ 2(&$ -% !"+4%$$,$ 2?0&5($0+& 4%--,-(0"% 4+==,&$ (,< 7!04+=!-%<(% ) -%$ =04"+&A=%$9 -%$ "@+!'"0%$ %' -%$ 1"(&,-%$ 2%&$%$C

(19)

ϭϴ

GW9WG)O(+)$%.-/"4$(+)

F% $+&' -%$ +"1(&0'%$ (!04(,< -%$ !-,$ !%'0'$9 %& B+"=% 2% /r'+&&%'$ =%$,"(&' j9f < j9c s=C K-$ (!!("(0$$%&' $+,5%&' -#1A"%=%&' (">,#$9 !"+/(/-%=%&' 2, B(0' 2?,&% ($$+40('0+& (5%4 -( B(4% 4Q'+!-($=0>,% 2% -?KUF SK&&%" U%=/"(&% F+=!-%<TC K- %<0$'% cd =04"+&A=%$ (!!%-#%$ UKF SUKF c ; cdT9 >,0 $+&' $+0' $+-,/-%$ $+0' '"(&$=%=/"(&(0"%$C E--%$ $+&' -0/#"#%$ -+"$ 2, 4+&'(4' 0&0'0(- 2, !("($0'% (5%4 -( 4%--,-%m@L'% %' !("'040!%&' ; -?(''(4@%=%&' 2, !("($0'% ; -( =%=/"(&% 2% -( 4%--,-%m@L'% S*@%0&%" *) +."= djcjTC

GW9W9)O(+)-0/2,-%(+) ))

I%$ "@+!'"0%$9 %& B+"=% 2% 4"+0$$(&' (--+&1#9 =%$,"%&' 2% c ; f s= 2% -+&1C H& '"+,5% k ; cd "@+!'"0%$ (, !L-% (!04(- 2% 4@(>,% !("($0'%C E--%$ $+&' 2#-0=0'#%$ !(" ,&% =%=/"(&%C I%$ "@+!'"0%$ 4+&'0%&&%&' ,& 1"(&2 &+=/"% 2%$ =+-#4,-%$ 4-#$ >,0 !%"=%''%&' (, !("($0'% 2% !#&#'"%" 2(&$ -%$ 4%--,-%$ @L'%$ %' 2% $,/5%"'0" -%$ 5+0%$ 2% $01&(-0$('0+& 2% -( 4%--,-% @L'% %& B(5%," 2, !("($0'% SM++'@"+Q2 %' 3,/"%=%'XC9 djji9 *(%0: *) +."9 djjk9 P(Q-+" *)

+."9 djjhTC I%$ "@+!'"0%$ 4+&'0%&&%&' 2% &+=/"%,$%$ !"+'#0&%$ S2+&' -%$ !"+'#0&%$ NH.T

(0&$0 >,% 2%$ -0!02%$C I?(&(-Q$% !"+'#+=0>,% 2?,&% B"(4'0+& !,"0B0#% 2% "@+!'"0%$ ( !%"=0$ 2% 2#4+,5"0" ei &+,5%--%$ !"+'#0&%$ 2% "@+!'"0%$ !,'('05%$9 2+&' 4%"'(0&%$ $+&' /0%& -+4(-0$#%$ 2(&$ -%$ "@+!'"0%$ SM"(2-%Q *) +."9 djjaTC I%$ NH. $+&' "%-("1,#%$ 2,"(&' -?0&5($0+& %' =01"%&' 5%"$ e -+4(-0$('0+&$ !"0&40!(-%$ ) -( -,=0A"% 2% -( 5(4,+-% !("($0'+!@+"% &(0$$(&'%9 -( =%=/"(&% 2% -( 5(4,+-% !("($0'+!@+"% +, -?0&'#"0%," 2% -( 4%--,-%m@L'% S&+Q(,TC E--%$ $+&' &+'(==%&' 0=!-0>,#%$ 2(&$ -( "#1,-('0+& 2%$ 5+0%$ 2% $01&(-0$('0+& 2% -( 4%--,-%m@L'%C

(20)

ϭϵ

GW9WD)O(+)3-#"'I(+)*("+(+))

I%$ 1"(&,-%$ 2%&$%$ $+&' 2%$ $'",4',"%$ $!@#"0>,%$ 2% djj &= 2% 20(=A'"%9 >,0 (!!("(0$$%&' 2%&$%$ (,< #-%4'"+&$ %& =04"+$4+!0% #-%4'"+&0>,%C H& %& 4+=!'% %&50"+& ca !(" '(4@QX+g'%9 20$!%"$#$ 2(&$ -?%&$%=/-% 2, 4Q'+!-($=%C I% 4+&'%&, 2%$ 1"(&,-%$ 2%&$%$ $% 2#5%"$% 2(&$ -( 5(4,+-% !("($0'+!@+"% -+"$ 2% -?0&5($0+& 2% -( 4%--,-% @L'% S7-%<(&2%"

*) +."9 djja9 I0= *) +.C9 djcdTC K-$ :+,%&' ,& "L-% =(:%," 2(&$ -%$ =+20B04('0+&$

$'",4',"(-%$ 2% -( 5(4,+-% !("($0'+!@+"%C E--%$ $?($$+40%&' ; -( 5(4,+-% !("($0'+!@+"% %' (, "#$%(, ',/,-(0"% 2% -( 4%--,-%m@L'%9 0&'%"5%&(&' 2% 4% B(0' 2(&$ -% "%4",'%=%&' 2%$ &,'"0=%&'$ %' 2(&$ -?0&@0/0'0+& 2%$ 5+0%$ (!+!'+'0>,%$ 2% -( 4%--,-%m@L'% SU%"40%" *) +."= djjaTC

(21)

ϮϬ

ͳǤ͵?L"/,!2(+)(,)K%-'I(".(

M0%& >,% 4+""%$!+&2(&' ; ,&% $%,-% %$!A4% %' =(-1"# bit 2?02%&'0'# 1#&#'0>,%9 -%$ $+,4@%$ 2% !" #$%&'' 20BBA"%&' 1"(&2%=%&' %& =('0A"% 2% 50",-%&4% S*0/-%Q %' M++'@"+Q29 cbbdTC I( !('@+1#&040'# 2%$ $+,4@%$ %$' 2#B0&0% !(" -?#',2% 2% -%," 50",-%&4% 4@%X -( $+,"0$ S2#'%"=0&('0+& 2% -( 3I aj %' 3I cjj9 2+$%$ =0&0=(-%$ 2% !("($0'%$ %&'"(u&(&' -( =+"' 2% ajt ; cjjt 2%$ $+,"0$ 0&B%4'#%$TC

I( 4-($$0B04('0+& (&40%&&% +!!+$(0'9 2?,&% !("' -%$ $+,4@%$ 2?E,"+!% %' 2?7=#"0>,% 2, G+"29 2?#=%"1%&4% "#4%&'% %' (!!("'%&(&' ; '"+0$ -01&#%$ 4-+&(-%$ =(:+"0'(0"%$ K9 KK %' KKK (Q(&' 2%$ 2%1"#$ 2% 50",-%&4% 5("0#$ S*, *) +.C9 djjd9 *(%0: *) +.C9 djjaT V 2?(,'"% !("' -%$ $+,4@%$ ('Q!0>,%$ 0$+-#%$ %& 7=#"0>,% 2, *,29 /%(,4+,! !-,$ (&4%$'"(-%$9 (Q(&' ,& @(,' 2%1"# 2% !+-Q=+"!@0$=% %' ,&% 50",-%&4% #-%5#% S*, *) +.C9 djjdTC F%$ $+,4@%$ $+&' 4("(4'#"0$#%$ !(" -( !"#$%&4% 2?(--A-%$ ,&0>,%$ &+& "%'"+,5#$ 2(&$ -%$ '"+0$ 'Q!%$ !"0&40!(,<C

I%$ 20BB#"%&4%$ %&'"% -%$ !"0&40!(,< 1#&+'Q!%$ 2?0$+-('$ 4+&4%"&%&' )

cC -( 50",-%&4% !+," -( $+,"0$ S,&% $+,4@% %$' 4+&$02#"#% 4+==% 50",-%&'% !+," -( $+,"0$ -+"$>,% -( 2+$% -#'(-% 3I cjj %$' 2% c '(4@QX+g'% 0&+4,-# !(" 5+0% 0&'"(!#"0'+&#(-%9 %&'"(u&(&' -( =+"' 2% -( $+,"0$ %& =+0&$ 2% cj :+,"$TC

dC -( 50'%$$% 2% =,-'0!-04('0+& %& 4,-',"% 4%--,-(0"% %' -( !+$$0/0-0'# 2% '"(&$B+"=('0+& '% W')6$ 2%$ '(4@QX+g'%$ %& /"(2QX+g'%$ (/+,'0$$(&' ; -( B+"=('0+& 2% ZQ$'%$C

(22)

Ϯϭ eC -%$ 4(!(40'#$ 2% =01"('0+& %' 2% '"(&$=01"('0+& ; '"(5%"$ -%$ /(""0A"%$

/0+-+10>,%$C

GWDWG)?L"/,!2()M))

K- %$' '"A$ "("%=%&' 0$+-# S&% "%!"#$%&'% >,% cjt 2%$ $+,4@%$ %& E,"+!% %' (,< E'('$m O&0$9 0- %$' !"0&40!(-%=%&' 0$+-# 4@%X -?@+==%TC F?%$' ,& 1#&+'Q!% '"A$ 50",-%&' !+," -( $+,"0$C .(" %<%=!-%9 -( $+,4@% NW ( ,&% 3Icjj 2% c '(4@QX+g'% 4@%X -( $+,"0$C I%$ '(4@QX+g'%$ $% =,-'0!-0%&' "(!02%=%&' '% W')6$9 0-$ $% '"(&$B+"=%&' !%, %& /"(2QX+g'%$ %' %& 4+&$#>,%&4%9 !%, 2% ZQ$'%$ $+&' B+"=#$ S4%$ $+,4@%$ !%, ZQ$'+1A&%$ $+&' !%, 0&B%4'(&'%$ !+," -% 4@('TC I%," 4(!(40'# 2% =01"('0+& %' 2% '"(&$=01"('0+& ; '"(5%"$ -%$ /(""0A"%$ /0+-+10>,%$ *1 W'W$ %$' $,!#"0%,"% ; 4%--% 2%$ 'Q!%$ KK %' KKKC I+"$ 2% '+<+!-($=+$%$ @,=(0&%$ 4(,$#%$ !(" 4%$ $+,4@%$9 -%$ (''%0&'%$ 4+&1#&0'(-%$ $+&' $+,5%&' $#5A"%$C

GWDW9)?L"/,!2()MM))

F?%$' -% 1#&+'Q!% -% !-,$ B"#>,%&' 0$+-# 4@%X -?@+==% Sijt 2%$ $+,4@%$ @,=(0&%$ %& E,"+!% %' (,< E'('$mO&0$T9 -% =+,'+&9 -% !+"4CCC K- %$' &+& 50",-%&' !+," -( $+,"0$ S3I cjj•cje '(4@QX+g'%$T9 =(0$ 4%$ $+,4@%$ $+&' ZQ$'+1A&%$ 2+&4 0&B%4'(&'%$ !+," -% 4@(' %' 1#&#"('"04%$ 2?++4Q$'%$C F@%X -( $+,"0$9 -( $#4"#'0+& 2?KJGm Ȗ %$' 4+&'"L-#% %' !"+'%4'"04%9 0- Q ( !(" (0--%,"$ 0&2,4'0+& !"#4+4% 2% -( $#4"#'0+& 2?KImcd !(" -%$ =(4"+!@(1%$C 3%$ #',2%$ +&' =+&'"# >,% -%$ $+,4@%$ 2% 'Q!% KK #'(0%&' !-,$

(23)

ϮϮ B"#>,%==%&' "%'"+,5#%$ 4@%X -%$ !('0%&'$ oKW %' -+"$ 2?0&B%4'0+&$ 4+&1#&0'(-%$C I+"$ 2% '+<+!-($=+$% @,=(0&% ) -%$ (''%0&'%$ 4+&1#&0'(-%$ !%,5%&' R'"% ($Q=!'+=('0>,%$ ; =+"'%--%$9 2%$ "#(4'05('0+&$ $+&' !+$$0/-%$ (, 4+,"$ 2%$ #'('$ 0==,&+2#B040%&'$ S4% $+&' -%$ $+,4@%$ !"#2+=0&(&'%$ 4@%X -%$ $02#%&$T9 4@%X -% !('0%&' 0==,&+4+=!#'%&'9 +& +/$%"5% 2%$ B+"=%$ -Q=!@(2#&+!('@0>,%$ %' 2%$ B+"=%$ ($Q=!'+=('0>,%$C

K- %$' "%$!+&$(/-% 2% iiCkt 2%$ 4($ 2% '+<+!-($=+$%$ 4+&1#&0'(-%$ (5%4 -%$ =R=%$ 4("(4'#"0$'0>,%$ >,% 2?@(/0',2%9 2% bj9bt 2%$ 4($ 2% '+<+!-($=+$%$ +4,-(0"%$ S0==,&+2#!"0=#$ +, 0==,&+4+=!#'%&'$T9 2% ift 2%$ '+<+!-($=+$%$ $Q$'#=0>,%$ 2%$ !('0%&'$ 0==,&+2#!"0=#$C SN(!!+"' FGN9 djcfTC

GWDWD)?L"/,!2()MMM))

K- %$' -% !-,$ "("% S1#&+'Q!% $,"'+,' 4+&B0&# ; -?(&0=(-TC *( 50",-%&4% !+," -( $+,"0$ %$' 0&'%"=#20(0"%C F+&4%"&(&' -( '+<+!-($=+$% @,=(0&%9 -%$ (''%0&'%$ 4+&1#&0'(-%$ $+&' !+$$0/-%$ =(0$ !%, B"#>,%&'%$C

I% 'Q!% KKK "%$'% !%, B"#>,%&' !("=0 -%$ 4($ djce Set 5$ f9dt $," -?%&$%=/-% 2%$ cdie !"#-A5%=%&'$ 1#&+'Q!#$ (, FGNT %' &% !"#$%&'% !($ ,& 4("(4'A"% 2% !('@+1#&040'# !-,$ =(">,# >,% -% 'Q!% KKC SN(!!+"' FGN9 djcfTC

ͳǤ͵ǤͶǤ±‘–›’‡•”‡…‘„‹ƒ–•‘—ƒ˜‡…ƒŽŽ°Ž‡•ƒ–›’‹“—‡•

I%$ 1#&+'Q!%$ ('Q!0>,%$ %' 6 +, "%4+=/0&(&' $% "%'"+,5%&' !-,$ B"#>,%==%&' 4@%X -%$ 0$+-('$ 28%$!A4%$ %<+'0>,%$ 28(44,%0-9 4+==% !(" %<%=!-% -%$ 0$+-('$ !"+5%&(&' 2% -( _,Q(&%9 2%$ X+&%$ 1#+1"(!@0>,%=%&' #-+01&#%$9 %' 4@%X -%$ !('0%&'$ (''%0&'$ 2% =(-(20%

(24)

Ϯϯ 1"(5% S7:X%&/%"1 *) +.C9 djjfTC F%$ 1#&+'Q!%$ +&' #'# ($$+40#$ ; -( '+<+!-($=+$% $#5A"% (4>,0$% 4@%X -%$ !('0%&'$ 0==,&+4+=!#'%&'$ %& _,Q(&% B"(&l(0$% SF("=% *) +.C9 djjdT9 -( '+<+!-($=+$% @,=(0&% +4,-(0"% S_"011 *) +.C9 djjcT9 %' -%$ B+"=%$ 1"(5%$ 2% '+<+!-($=+$% 4+&1#&0'(-% S7:X%&/%"1 *) +.C9 djjdTC K-$ $+&' !-,$ $+,5%&' 0$+-#$ 2(&$ 2%$ 40"4+&$'(&4%$ #!02#=0+-+10>,%$ !("'04,-0A"%$ S(&0=(,<9 /0+'+!%$ $(,5(1%$T %' $+&' !-,$ 50",-%&'$ 4@%X -( $+,"0$ >,% -% 'Q!% KKC I+"$ 2?0&B%4'0+& 4@%X -?@+==%9 +& !%,' +/$%"5%" 2%$ (''%0&'%$ $#5A"%$ 20$$#=0&#%$ +, +4,-(0"%$ 4@%X 2%$ $,:%'$ 0==,&+4+=!#'%&'$ S_"011

*) +.C9 djjcTC 3%$ 0$+-('$ ('Q!0>,%$ 2% _,Q(&% $+&' "%$!+&$(/-%$ 2% '+<+!-($=+$%$

(4>,0$%$ (5%4 2%$ B+"=%$ =,-'050$4#"(-%$9 !("B+0$ =+"'%--%$9 4@%X -%$ !%"$+&&%$ 0==,&+4+=!#'%&'%$ S3%=(" *) +.C9 djcdT

E& J"(&4%9 0- %<0$'% ,&% !"#2+=0&(&4% '"A$ &%''% 2%$ $+,4@%$ 2% 'Q!% KK SbatT (-+"$ >,% -%$ $+,4@%$ 2% 'Q!% K &% "%!"#$%&'%&' >,% dt %' 4%--%$ 2% 'Q!% KKK et 2%$ $+,4@%$ SN(!!+"' FGN*C9 djcf V W+&+"# *) +.C9 djjjTC .-,$0%,"$ #',2%$ +&' 0&5%$'01,# -%$ 4+""#-('0+&$ !+$$0/-%$ %&'"% -%$ 'Q!%$ 2% $+,4@%$ %' -%$ =(&0B%$'('0+&$ 4-0&0>,%$ 4@%X -?W+==% SM++'@"+Q2 *) +.C9 djjdTC

I( '%4@&0>,% 2% .FN =,-'0!-%< ca =04"+$('%--0'%$ ( #'# 5(-02#% %& djcj S7:X%&/%"1 *) +." 9 djcjTC E--% ( B(0' -?+/:%' %& djcd 2?,&% 4+=!("(0$+& (5%4 -%$ "#$,-'('$ 2%$ '%4@&0>,%$ 2% .FNmNJI. Scc =(">,%,"$T %' 2% $#>,%&l(1% Sf 0&'"+&$T ,'0-0$#%$ !(" 2?(,'"%$ #>,0!%$ S*,

*) +.= .G7*9 djcdTC F%''% 4+=!("(0$+& >,0 ( !+"'# $," cei $+,4@%$ ( =+&'"# -% !-,$ 1"(&2

!+,5+0" 20$4"0=0&(&' 2%$ =04"+$('%--0'%$9 /0%& (2(!'# (,< #',2%$ #!02#=0+-+10>,%$C E--% ( (,$$0 !%"=0$ ,& "%1"+,!%=%&' 2%$ 1#&+'Q!%$ 2(&$ 2%$ @(!-+1"+,!%$ !-,$ -("1%$C 7, $%0& 2, FGN !+," -( P+<+!-($=+$%9 -?(&(-Q$% 1#&#'0>,% 2%$ "#$,-'('$ +/'%&,$ !+," -%$ ca =04"+$('%--0'%$ !%"=%' 2% 2#B0&0" -%$ $+,4@%$ 4+==% (!!("'%&(&' $+0' ; ,&% 2%$ e -01&#%$

(25)

Ϯϰ 4-+&(-%$ 0&0'0(-%=%&' 2#4"0'% S'Q!% K9 KK +, KKK9 (5%4 5("0(&' -+"$>,?,& $%,- 2%$ =04"+$('%--0'%$ 2#B0&0$$(&' -% 'Q!% %$' 20BB#"%&'T9 $+0' ; 2?(,'"%$ 1"+,!%$ 1#&#'0>,%$ 4("(4'#"0$#$ !(" -%," 20$'"0/,'0+& 1#+1"(!@0>,% S7B"04(& c9 d9 e V F("0//%(& c9 d9 e V 7=(X+&0(&qTC 3%$ 1#&+'Q!%$ &?(!!("'%&(&' !($ ; 4%$ 1"+,!%$ $+&' >,(-0B0#$ 2?('Q!0>,%$ +, 2% "%4+=/0&(&'$C SN(!!+"' FGNC9 djcfTC

I%$ =04"+$('%--0'%$ !%"=%''%&' #1(-%=%&' !(" -%," B+"' !+,5+0" 20$4"0=0&(&' ,& '"(l(1% #!02#=0+-+10>,% 2%$ 4($ 1"+,!#$C

I( 4+=/0&(0$+& 2% '%4@&0>,%$ 2% NJI. SN%$'"04'0+& J"(1=%&'$ I%&1@' .+-Q=+"!@0$=T9 =04"+$('%--0'%$ %' $#>,%&l(1% 2% =(">,%,"$ $!#40B0>,%$ +&' "#4%==%&' !%"=0$9 $," ,&% 4+@+"'% 2% bak 0$+-('$9 2% 2#B0&0" cei 1#&+'Q!%$9 +"1(&0$#$ %& ca @(!-+1"+,!%$ 2#B0&0$$(&'$ k 4-(2%$ =(:%,"%$ S J01,"% hT S*, *) +.C9 djcdTC

(26)

Ϯϱ 3(&$ -% 4(2"% 2% 4% '"(5(0-9 -%$ $+,4@%$ NW S@(!-+1"+,!% c9 4-(2% 7 V (&40%&&%=%&' 'Q!% K9 0$+-#% 2?,&% %&4#!@(-0'% @,=(0&% (01,v %& cbfcT9 .",1&0(,2 S.",9 @(!-+1"+,!% d9 4-(2% 39 (&40%&&%=%&' 'Q!% KK9 0$+-#% 2?,&% '+<+!-($=+$% 4+&1#&0'(-% $#5A"% %& cbkfT %' IEJ S!"+/(/-%=%&' @(!-+1"+,!% i9 4-(2% M9 (&40%&&%=%&' 4-($$#% !("=0 -%$ $+,4@%$ ('Q!0>,%$9 0$+-#% 2?,&% '+<+!-($=+$% 4+&1#&0'(-% 4@+"0+"#'0&+1A&% %& cbbfT +&' #'# ,'0-0$#%$C

ͳǤ͵ǤͷǤ ƒ…–‡—”•’ƒ”ƒ•‹–ƒ‹”‡•Ž‹±•Žƒ˜‹”—Ž‡…‡)

I%$ 1A&%$ "%$!+&$(/-%$ 2% -( 50",-%&4% 2%$ $+,4@%$ +&' #'# #5(-,#$ !(" %<!#"0%&4%$ 2% 4"+0$%=%&' %&'"% 2%$ $+,4@%$ K %' KKK +, KK %' KKK9 0&'"+2,0'%$ %&$,0'% 4@%X -( $+,"0$C F0&> oKN So0",-%&' -+40T +&' #'# =0$ %& #502%&4% %' '"+0$ !"+'#0&%$ 2% "@+!'"0%$ $+&' 0&4"0=0&#%$ )

m NH.ci9 ,&% !"+'#0&%m$#"0&%m'@"#+&0&% Z0&($% >,0 !@+$!@+"Q-% %' 0&(4'05% -%$ KN_$ SKJGȖm0&2,40/-% K==,&0'QmN%-('%2 _P.($%$T 2% -?@L'% SJ%&'"%$$ *) +.C9 djcj9 *'%0&B%-2' *)

+.C9 djcjT V 

m NH.ck9 ,&% !"+'#0&%m'Q"+$0&% Z0&($% !+-Q=+"!@0>,% >,0 !@+$!@+"Q-% -( B(=0--% 2%$ !"+'#0&%$ *P7P S*01&(- P"(&$2,4%"$ (&2 74'05('+"$ +B P"(&$4"0!'0+&T 2% -?@L'% S[(=(=+'+ *) +.C9 djjb9 H&1 *) +.C9 djcjTC I+"$ 2% $+& 0&:%4'0+& 2(&$ -( 4%--,-% @L'%9 %--% !@+$!@+"Q-% 20"%4'%=%&' *P7Pe %' *P7Pk9 %' 2(&$ -%$ 4%--,-%$ 0&B%4'#%$ !(" -%$ $+,4@%$ 2% 'Q!%$ K %' KKK9 =(0$ !($ 2(&$ -%$ 4%--,-%$ 0&B%4'#%$ !(" 2%$ $+,4@%$ 2% 'Q!% KK9 4%' %BB%' $% =(0&'0%&' S*(%0: *) +.C9 djjh9 [(=(=+'+ *) +.C9 djjb9 H&1 *) +.C9 djcjT V

(27)

Ϯϲ m ,& '"+0$0A=% -+4,$9 02%&'0B0# 4+==% (Q(&' -% !-,$ 1"(&2 0=!(4' $," -( 50",-%&4% 2?,&% $+,4@% 2% 'Q!% K 4@%X -( $+,"0$ %$' "%!"#$%&'# !(" NH.a9 >,0 4+2% !+," ,& 4-,$'%" 2% !$%,2+Z0&($%$ !+-Q=+"!@0>,%$ $#4"#'#%$ S*(%0: *) +.C9 djjk9 N%%$% *) +.C9 djccTC

.-,$ "#4%==%&'9 ,& &+,5%(, B(4'%," 2% 50",-%&4%9 _N7da ( #'# 02%&'0B0#C _N7da %$' ,&% !@+$!@+!"+'#0&% $#4"#'#% !(" -%$ 1"(&,-%$ 2%&$%$ %' "%'"+,5#% 2(&$ -( 5(4,+-% !("($0'+!@+"%C _N7da :+,%"(0' ,& "L-% 4-# 2(&$ -( "#1,-('0+& 2% -( "#!+&$% 0==,&0'(0"% ; -?0&B%4'0+& !(" !" #$%&'' S*@($'"0 *) +.C9 djcfTC

GWH)Q!.I()F%/I/3%&'()*()T. gondii)

I% 4Q4-% #5+-,'0B 4+=!"%&2 ,&% !@($% 2% =,-'0!-04('0+& $%<,#% >,0 ( -0%, 4@%X -% 4@(' %' ,&% !@($% 2% =,-'0!-04('0+& ($%<,#% >,0 ( -0%, 4@%X -%$ 5%"'#/"#$ ; $(&1 4@(,2C

GWHWG)Q!.I()#+(Y'L))

K- !%,' $% 2#"+,-%" 4@%X '+,$ -%$ (&0=(,< ; $(&1 4@(,2C 7!"A$ 0&1%$'0+& 2%$ ++4Q$'%$9 -%," !("+0 $% "+=!' 2(&$ -?0&'%$'0& V 0-$ !#&A'"%&' 2(&$ -%$ 4%--,-%$ #!0'@#-0(-%$ 0&'%$'0&(-%$ %' $% '"(&$B+"=%&' %& '(4@QX+g'%$ >,0 20$$#=0&%&' "(!02%=%&' 2(&$ -?+"1(&0$=%C F%''% 20$$#=0&('0+& 4+""%$!+&2 ; -( !@($% (01w% 2% -?0&B%4'0+&C *+,$ -?0&B-,%&4% 2% -( "#!+&$% 0==,&0'(0"%9 -( 1"(&2% =(:+"0'# 2%$ '(4@QX+g'%$ %$' #-0=0&#%C F%,< >,0 &% $+&' !($ -Q$#$ =01"%&' %' $% '"(&$B+"=%&' %& /"(2QX+g'%$ (, $%0& 2% 4%"'(0&$ '0$$,$ V 0- %& "#$,-'% -( B+"=('0+& 2% ZQ$'%$ !"0&40!(-%=%&' -+4(-0$#$ 2(&$ '+,' -% $Q$'A=% &%"5%,< 4%&'"(-9 -?]0-9 %' -%$ =,$4-%$ 4("20(>,%$ %' $>,%-%''0>,%$ S3,/%Q9 djjiTC I%$ ZQ$'%$ '0$$,-(0"%$ !%,5%&' !%"$0$'%" '+,'% -( 50% 2% -?@L'% %& -?(/$%&4% 2?(-'#"('0+& 2% -?0==,&0'# $(&$ %&'"(0&%" 2% =(&0B%$'('0+&$ 2#-#'A"%$C

(28)

Ϯϳ

GWHW9)Q!.I()+(Y'L))

K- ( -0%, 4@%X -?@L'% 2#B0&0'0B S4@('T >,0 $% 4+&'(=0&% !(" 0&1%$'0+& 2% ZQ$'%$ 4+&'%&,$ 2(&$ -%," !"+0% +, !(" -?0&'%"=#20(0"% 2?++4Q$'%$ $+,0--(&' -?%(, +, -%$ 5#1#'(,<C I%$ /"(2QX+g'%$ +, -%$ $!+"+X+g'%$ -0/#"#$ $% =,-'0!-0%&' 2(&$ -%$ 4%--,-%$ #!0'@#-0(-%$ 2, 4@(' !(" $4@0X+1+&0%C 7!"A$ !-,$0%,"$ 1#&#"('0+&$9 ,&% !@($% 2% 1(=+1+&0% $?0&0'0%C I%$ B+"=%$ ($%<,#%$ $% '"(&$B+"=%&' %& 1(=#'+4Q'%$ =r-%$ %' B%=%--%$C I( B#4+&2('0+& 4+&2,0' ; -( B+"=('0+& 2?++4Q$'%$ $!+",-#$9 #-0=0&#$ 2(&$ -% =0-0%, %<'#"0%," (5%4 -%$ BA4%$ 2, 4@('C K-$ 2%50%&&%&' 0&B%4'0%,< %& c ; a :+,"$C I%$ ++4Q$'%$ $!+",-#$ $+&' '"A$ "#$0$'(&'$ %' "%$'%&' 0&B%4'(&'$ !-,$0%,"$ =+0$ 2(&$ ,& $+- @,=02%C

GWc)Q/",#$%"#,%/"))

I( 4+&'(=0&('0+& !%,' $% B(0"% )

¾ @+"0X+&'(-%=%&' !(" 0&1%$'0+& +"(-% 2%$ ++4Q$'%$ 0&B%4'0%,< 2% -?%&50"+&&%=%&' ) 5#1#'(,< +, %(, 2% /+0$$+& $+,0--#$ !(" 2%$ $%--%$ 2% B#-0&$C

¾ @+"0X+&'(-%=%&' !(" 0&1%$'0+& +"(-% 2%$ ZQ$'%$ 4+&'%&,$ 2(&$ -( 50(&2% 4",% +, 0&$,BB0$(==%&' 4,0'%C

¾ 5%"'04(-%=%&' !(" '"(&$=0$$0+& '"(&$!-(4%&'(0"% 2%$ '(4@QX+g'%$ -+"$ 2?,&% !@($% 2% !("($0'#=0% =('%"&%--%C

¾ '"(&$=0$$0+& !-,$ %<4%!'0+&&%--% !(" 1"%BB+& -+"$ 2?,&% '"(&$!-(&'('0+& +, !(" 0&+4,-('0+& (4402%&'%--%C

(29)

Ϯϴ

7'#/6* > 9 (-,.* 2+6+0')+'6* &* !$1$2.+03+ #$%&'' :&E+26H0;$N*6)Z[+%#%*/1*) O+6&Y= AJ8A@

9W)O#),/Y/2I#+$/+()

9WG)62%*L$%/I/3%()

I( '+<+!-($=+$% 2,% ; !$1$2.+03+ #$%&'' %$' ,&% 0&B%4'0+& 4+$=+!+-0'% 2+&' -( !"#5(-%&4% 5("0% 2?,& !(Q$ ; -?(,'"%C

I( '+<+!-($=+$% %$' -?,&% 2%$ 0&B%4'0+&$ -%$ !-,$ !"#5(-%&'%$ %& J"(&4% (5%4 2%$ 5(-%,"$ 2% $#"+!"#5(-%&4% 4@%X -?(2,-'% 4+=!"0$%$ %&'"% dj %' aat9 5("0(/-%$ $,05(&' -?r1%9 -( "#10+& 1#+1"(!@0>,% %' -( 4('#1+"0% $+40+!"+B%$$0+&&%--%C I( $#"+!"#5(-%&4% %& J"(&4% (

(30)

Ϯϵ 5("0# 4+&$02#"(/-%=%&' 2%!,0$ ej (&$ (5%4 ,&% /(0$$% 2% !"#5(-%&4% "#1,-0A"%9 %$'0=#% (4',%--%=%&' ; eht SN#$,-'('$ EG.9 djcjTC

_#&#"(-%=%&' /#&01&% !+," -?@+==%9 4%''% =(-(20% !%,' 4%!%&2(&' R'"% 1"(5% 2(&$ 4%"'(0&%$ 40"4+&$'(&4%$ )

cT F@%X -( B%==% %&4%0&'%9 +p ,&% !"0=+m0&B%4'0+& '+<+!-($=0>,% !%,' R'"% '"(&$=0$% (, B+%',$ (''%0&' 2% '+<+!-($=+$% 4+&1#&0'(-% SPFT !+,5(&' %&'"(u&%" 2% 1"(5%$ $#>,%--%$C I%$ B+"=%$ 0&(!!("%&'%$ $+&' -%$ !-,$ B"#>,%&'%$ ; -?@%,"% (4',%--% =(0$ -%," !"+&+$'04 #5+-,'0B 2%=%,"% 0&4%"'(0& S"0$>,% 2% 4@+"0+"#'0&0'%$ ,-'#"0%,"TC

E& djce9 djf 4($ 2% '+<+!-($=+$%$ 4+&1#&0'(-%$ +&' #'# 20(1&+$'0>,#$ %& J"(&4% V -( !"#5(-%&4% 1-+/(-% 2% -( '+<+!-($=+$% 4+&1#&0'(-% +/$%"5#% %& J"(&4% %$' (,'+," 2% d9k !+," cj jjj &(0$$(&4%$) %' -( !"#5(-%&4% 2%$ 4($ 2% '+<+!-($=+$% 4+&1#&0'(-% 20(1&+$'0>,#$ ; -( &(0$$(&4% %$' 2% c9ea !+," cj jjj &(0$$(&4%$C I( -#'(-0'# -0#% ; -( '+<+!-($=+$% 4+&1#&0'(-% %$' 2% k9ft S$'(/0-0'# 2%!,0$ djjhT %' -( =+"/020'# 1-+/(-% "%!"#$%&'% i9a t S%& 20=0&,'0+& !(" "(!!+"' ; -?(&&#% djcc9 =(0$ $'(/-% !(" "(!!+"' (,< (&&#%$ !"#4#2%&'%$TC

dT F@%X -%$ =(-(2%$ 0==,&+2#!"0=#$9 -( '+<+!-($=+$% %$' !("'04,-0A"%=%&' $#5A"%9 =+"'%--% $(&$ '"(0'%=%&'C E--% 4+""%$!+&2 -% !-,$ $+,5%&' ; -( "#(4'05('0+& 2?,&% 0&B%4'0+& (&'#"0%,"%=%&' (4>,0$%C

I( B+"'% !"#5(-%&4% 2% -( '+<+!-($=+$% %& J"(&4% 2(&$ -( !+!,-('0+& 1#&#"(-% %<!-0>,(0' -( '"A$ B+"'% 0&402%&4% 2% -( '+<+!-($=+$% +/$%"5#% 2(&$ -%$ (&&#%$ cbijmbj (, 4+,"$ 2, *K37C 3%!,0$ -%$ (&&#%$ djjj9 (5%4 -?0&'"+2,4'0+& 2% &+,5%(,< (&'0m"#'"+50"(,< %& 4+=/0&(0$+&9 ,&% 20=0&,'0+& 2%$ 4($ %$' +/$%"5#% S(5%4 $+,5%&' 2#4+,5%"'% 2, $'(','

(31)

ϯϬ oKW %& =R=% '%=!$ >,?%$' !+$# -% 20(1&+$'04 2% -( '+<+!-($=+$% 20'% +!!+"',&0$'%TC F%!%&2(&'9 4% "0$>,% 2%=%,"% !+," 2?(,'"%$ 'Q!%$ 2?0==,&+2#!"%$$0+& %' '%&2 =R=% ; (,1=%&'%" $(&$ >,% 2%$ 2+&&#%$ 4+&$+-02#%$ &% $+0%&' %&4+"% 20$!+&0/-%$ S!('0%&'$ (''%0&'$ 2% 4(&4%"9 !('0%&'$ 1"%BB#$T

eT F@%X -% !('0%&' 0==,&+4+=!#'%&'9 2%$ B+"=%$ 1"(5%$ (5%4 2#'"%$$% "%$!0"('+0"% (01,v9 5+0"% 2%$ B+"=%$ =+"'%--%$ +&' #'# 2#4"0'%$ &+'(==%&' %& _,Q(&% J"(&l(0$%C 3%$ B+"=%$ 1"(5%$ +&' (,$$0 #'# 2#4"0'%$9 /0%& >,% !-,$ %<4%!'0+&&%--%=%&'9 $," -% '%""0'+0"% =#'"+!+-0'(0&C I%$ $+,4@%$ ; -?+"010&% 2% 4%$ 4($ $#5A"%$ $% $+&' "#5#-#%$ 1#&#'0>,%=%&' 20BB#"%&'%$ 2% 4%--%$ 40"4,-(&' @(/0',%--%=%&' %& J"(&4% =#'"+!+-0'(0&%9 -( $,"5%0--(&4% 2% -?#=%"1%&4% 2% 1#&+'Q!%$ ('Q!0>,%$ %$' -?,&% 2%$ =0$$0+&$ 2, I(/+"('+0"% 7$$+40# .L-% *+,4@%$C

7'#/6* \ 9 46YW+.*%,* &* .+ )$1$2.+03$0* *% 76+%,* :;+22$6) S7DDS= AJJT@"

(32)

ϯϭ

9W9)O#),/Y/2I#+$/+()0'$#%"()

W(/0',%--%=%&' /#&01&% 4@%X -?R'"% @,=(0&9 %--% %$' ($Q=!'+=('0>,% 2(&$ bjt 2%$ 4($ 2?0&B%4'0+& (01,v SU+&'+Q( %' I0%$%&B%-29 djjfTC F%!%&2(&' 2(&$ 4%"'(0&% $0',('0+&$ S1"+$$%$$%9 0==,&+2#!"%$$0+&$qT9 -?0&B%4'0+& !%,' $?(5#"%" 1"(5%C

O#) ,/Y/2I#+$/+() *() IJ%$$'"/*L2-%$L SoKW9 (--+1"%BB% 2% =+%--% +$$%,$%9 1"%BB%

2?+"1(&%9 4@0=0+'@#"(!0% +, '"(0'%=%&'$ 0==,&+$,!!"%$$%,"$T "#$,-'% 2?,&% !"0=+m 0&B%4'0+& +, 2?,&% "#(4'05('0+& 2% ZQ$'%$ !"#$%&'$ 4@%X ,& $,:%' (&'#"0%,"%=%&' 0&B%4'#C 3(&$ -% 4($ 2% '"(&$!-(&'('0+&$ 2?+"1(&%$9 -( "#(4'05('0+& !%,' $% B(0"% ; !("'0" 2, 1"%BB+& $0 4%-,0m40 %$' 0&B%4'#C I( !"0&40!(-% 4+=!-04('0+& %$' -?%&4#!@(-0'% '+<+!-($=0>,% /0%& >,% 2?(,'"%$ +"1(&%$ !%,5%&' R'"% '+,4@#$ 4+==% -%$ !+,=+&$ +, -?]0-C

O#),/Y/2I#+$/+()./"3L"%,#I( "#$,-'% 2?,&% !"0=+m0&B%4'0+& (4>,0$% !(" -( =A"% (, 4+,"$

2% -( 1"+$$%$$%C E--% ( -0%, !(" !($$(1% '"(&$!-(4%&'(0"% -+"$ 2% -( !@($% 2% !("($0'#=0%C I( !"+/(/0-0'# 2% '"(&$=0$$0+& 2, '+<+!-($=% %' -( 1"(50'# 2% -?(''%0&'% B]'(-% #5+-,%&' 0&5%"$%=%&' (, 4+,"$ 2% -( 1"+$$%$$% SU+&'+Q( %' I0%$%&B%-29 djjfTC K- &?Q ( !($ 2% '"(&$=0$$0+& 4@%X -( B%==% 0==,&0$#% !(" ,&% 0&B%4'0+& (&'#"0%,"% ; -( 1"+$$%$$% 4(" -( "#!+&$% 0==,&0'(0"% !"#$%&'% %$' !"+'%4'"04% %' 4+&'"L-% -( 20$$#=0&('0+& !("($0'(0"%C I( !('@+1#&0% !%,' (--%" 2% -?0&B%4'0+& ($Q=!'+=('0>,% ; 2% 1"(5%$ $Q=!'L=%$ S@Q2"+4#!@(-0%9 %&4#!@(-0'%T +, ; 2%$ $#>,%--%$ S"%'("2 !$Q4@+=+'%,"9 4@+"0+"#'0&0'%$TC I% "0$>,% 2?0&B%4'0+& B]'(-% -,0 %$' 20"%4'%=%&' 4+""#-# ; -?#'(' 2?(5(&4%=%&' 2% -( 1"+$$%$$%C K- !($$% 2% cjt (, !"%=0%" '"0=%$'"% ; !-,$ 2% kjt %& B0& 2% 1"+$$%$$%C

O#) ,/Y/2I#+$/+() /.'I#%-( %$' ,&% 0&B%4'0+& "%2+,'(/-%9 4(" !+'%&'0%--%=%&' 4#40'(&'%C

(33)

ϯϮ 0==,&0'(0"% 2% -?@L'%C I( 4@+"0+"#'0&0'% '+<+!-($=0>,% %$' -% !-,$ $+,5%&' -% "#$,-'(' 2?,&% "#(4'05('0+& ZQ$'0>,% -+4(-% 2(&$ -( "#'0&%9 >,% 4%$ ZQ$'%$ !"+50%&&%&' 2?,&% 0&B%4'0+& 4+&1#&0'(-% S4($ -% !-,$ B"#>,%&'T +, &+&C

K- %<0$'% 2% !-,$ ,&% 5("0(/0-0'# 4-0&0>,% (, &05%(, =+&20(-% %&'"% -?E,"+!% %' -?7=#"0>,% 2, $,2 &+'(==%&'C I% &+=/"% %$'0=# 2% '+<+!-($=+$%$ 4+&1#&0'(-%$ (, M"#$0- !(" %<%=!-% %$' a ; ca B+0$ !-,$ #-%5# >,?%& E,"+!% +, %& 7=#"0>,% 2, G+"2C I( =(-(20% %$' &+& $%,-%=%&' !-,$ B"#>,%&'% %& 7=#"0>,% 2, $,2 =(0$ #1(-%=%&' !-,$ $#5A"% >,?%& E,"+!%C E& %BB%'9 2%$ #',2%$ =+&'"%&' >,% !"A$ 2% ajt 2%$ %&B(&'$ $,2m(=#"04(0&$ 5+&' 2#5%-+!!%" 2%$ -#$0+&$ +4,-(0"%$ (, 4+,"$ 2% -( !"%=0A"% (&&#% 2% 50% $,0'% ; ,&% '+<+!-($=+$% 4+&1#&0'(-%C 3% !-,$9 -%$ -#$0+&$ $+&' !-,$ 1"(&2%$9 !-,$ &+=/"%,$%$ %' -% !+,"4%&'(1% 2% "#4,""%&4% %$' #1(-%=%&' !-,$ #-%5# S.B(BB 7\ *) +.C9 djcfTC E& F+-+=/0%9 -% '(,< 2% -#'(-0'# 4@%X -%$ %&B(&'$ 4+&1#&0'(-%=%&' 0&B%4'#$ %& -8(/$%&4% 2% '"(0'%=%&' !"#&('(- %$' 2?%&50"+& dat S_+=%XmU("0& *) +.C9 djccTC



9WD);-#%,($(",+))

K- %<0$'% ,& &+=/"% -0=0'# 2% =#204(=%&'$ "%1"+,!#$ %& 2%,< 1"(&2%$ B(=0--%$ ) -%$ 0&@0/0'%,"$ 2% -( $Q&'@A$% 2% -?(402% B+-0>,% %' -%$ =(4"+-02%$ S3%"+,0&9 djjcTC F%$ =#204(=%&'$ &% $+&' (4'0B$ >,% $," -%$ '(4@QX+g'%$ %' &+& $," -%$ ZQ$'%$C K- &?%<0$'% ; -?@%,"% (4',%--% (,4,& 5(440& !+," -?@+==%C

I( !"%=0A"% !"#5%&'0+& 4@%X -( B%==% %&4%0&'% "%$'% -?+/$%"5(&4% 2?,&% $#"0% 2% "A1-%$ @Q10#&+m20#'#'0>,%$C O&% !"0$% %& 4@("1% '@#"(!%,'0>,% %$' !+$$0/-%C E& J"(&4%9 -%

(34)

ϯϯ '"(0'%=%&' 2% -( =A"% !(" $!0"(=Q40&% %$' ,'0-0$# %& 4($ 2?0&B%4'0+& 2,"(&' -( 1"+$$%$$%C E& 4($ 2?0&B%4'0+& B]'(-% !"+,5#%9 -?($$+40('0+& !Q"0=#'@(=0&%m$,-B(=02% $%"( !"%$4"0'%C .+," -%$ !('0%&'$ 0==,&+2#!"0=#$9 -%$ =R=%$ "%4+==(&2('0+&$ (!!-0>,#%$ 4@%X -( B%==% %&4%0&'% 50$(&' ; #50'%" -( 4+&'(=0&('0+& $%"+&' !"#4+&0$#%$C O&% 4@0=0+!"+!@Q-(<0% !(" '"0=#'@+!"0=%m$,-B(=#'@+<(X+-% !+,""( R'"% 0&$'(,"#% %& 4($ 2% B+"'% 0==,&+2#!"%$$0+&C

)

(35)

ϯϰ

Q0#2%,-()MMV)T. gondii (,)I()a#.,('-)*(),-#"+.-%2,%/")AN5κ

κ

κ

κT)

)

GW)O()a#.,('-)*(),-#"+.-%2,%/")AN5κ

κ

κ

κT)

I% &,4-%(" B(4'+"mκM SGJmκ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djjdTC .(" 4+&$#>,%&'9 -( =+2,-('0+& 2% -8(4'05('0+& 2% GJmκM %' $( 5+0% 2% $01&(-0$('0+& +BB"%&' ,&% $'"('#10% '@#"(!%,'0>,% %<4%!'0+&&%--% >,0 !%"=%''"(0' 2% 40/-%" 4%''% 5+0% 2% $01&(-0$('0+& 2% -( 4%--,-% @L'% So0'0%--+ *) +.C 9 djcdTC

I%$ "#!+&$%$ 0==,&0'(0"%$ (,< 0&B%4'0+&$ /(4'#"0%&&%$ +, 50"(-%$ &#4%$$0'%&' -8(4'05('0+& 2%$ 5+0%$ 2% $01&(-0$('0+& $!#40B0>,%$ >,0 4+&2,0$%&' -%$ $01&(,< %<'"(4%--,-(0"%$ 2% -( $,"B(4% 2% -( 4%--,-% @L'% 5%"$ -% &+Q(, S^("0& *) +.C9 cbbaT9 &+Q(, 2(&$ -%>,%- -( -0(0$+& +, -( =+2,-('0+& 2% -8(4'050'# 2%$ B(4'%,"$ 2% '"(&$4"0!'0+& :+,%&' ,& "L-% %$$%&'0%- 2(&$ -( "#1,-('0+& 2% -8%<!"%$$0+& 2%$ 1A&%$C

GJmκM %$' %<!"0=#% 2(&$ 2% &+=/"%,< 'Q!%$ 4%--,-(0"%$ %' 4+&'"L-% -8%<!"%$$0+& 2% !-,$0%,"$ 1A&%$ !"+m0&B-(==('+0"%$ %' 0==,&+m"#1,-('%,"$ >,0 2#'%"=0&%&' -80$$,% 2% -( "#!+&$% 0==,&0'(0"% 0&&#% 2% -8@L'% 50$m;m50$ 2%$ !('@+1A&%$ =04"+/0%&$ %' 4+&'"0/,%&' ; -8(4'05('0+& 2% !"%$>,% '+,$ -%$ 'Q!%$ 2% 4%--,-%$ 0=!-0>,#%$ 2(&$ -( "#!+&$% 0==,&0'(0"%)

(36)

ϯϱ -%$ &%,'"+!@0-%$9 -%$ =(4"+!@(1%$9 -%$ -Q=!@+4Q'%$9 -%$ 4%--,-%$ %&2+'@#-0(-%$ %' #!0'@#-0(-%$C

P+,'%$ -%$ =+-#4,-%$ 2% GJmκM $+&' (4'05#%$ !(" 20BB#"%&'$ 'Q!%$ 2% $'0=,-0 0&B%4'0%,<9 =(0$ 0- !%"$0$'% %&4+"% 2%$ >,%$'0+&$ 4+&4%"&(&' -%$ "L-%$ $!#40B0>,%$ 2%$ =%=/"%$ 2% -( B(=0--% GJmκM (, 4+,"$ 28,&% "#!+&$% 0==,&0'(0"% -+"$ 2?,&% 0&B%4'0+&C 30BB#"%&' !("($0'%$ 4+==% !$1$2.+03+ #$%&''9 P*'0<3+%'+ &$%$W+%'9 P*'0<3+%'+ 3+]$6 %'

!6',</6'0 3/6'0 $+&' 4(!(/-%$ 2?(4'05%" -( 4($4(2% 2% $01&(-0$('0+& 2% GJmκMC 3% !-,$ 2%$ =+-#4,-%$ !("($0'(0"%$ !%,5%&' 0&'%"B#"%" (5%4 -%$ 20BB#"%&'%$ 5+0%$ >,0 =A&%&' ; -8(4'05('0+& 2% GJmκMC 70&$09 /0%& >,% -?(4'05('0+& 2% -( 5+0% GJmκM "%!"#$%&'% ,&% !("'0% 2% -( "#1,-('0+& !@Q$0+-+10>,% &+"=(-%9 ,&% (4'05('0+& !%"$0$'(&'% %' %<4%$$05% 2, GJmκM 4+&2,0' ; -( !"+2,4'0+& 2%$ 4Q'+Z0&%$ !"+m0&B-(==('+0"%$9 -( $,"!"+2,4'0+& 2%$ 4@0=0+Z0&%$ %' ; ,&% 0&B-(==('0+& 4@"+&0>,% 4("(4'#"0$'0>,% 2%$ !("($0'+$%$ -('%&'%$C .-,$0%,"$ #',2%$ "#4%&'%$ $,11A"%&' >,% -8@Q!%"(4'05('0+& 2% -( 5+0% 2% $01&(-0$('0+& 2% GJmκM %$' ($$+40#% ; -8#-(/+"('0+& %' ; -( !"+1"%$$0+& 2% &+=/"%,$%$ =(-(20%$9 4+==% -%$ '"+,/-%$ (,'+m0==,&$9 -% 4(&4%"9 -%$ '"+,/-%$ 4("20+m5($4,-(0"%$ %' -%$ '"+,/-%$ &%,"+2#1#&#"('0B$C

I8(4'05('0+& 2%$ !"+'#0&%$ 2% -( B(=0--% GJmκM %' -%," 20=#"0$('0+& $+&' -%$ 4+&20'0+&$ !"#(-(/-%$ &#4%$$(0"%$ ; -?0&2,4'0+& 2% -( '"(&$4"0!'0+& 2%$ 1A&%$ >,0 4+2%&' !+," 20BB#"%&'%$ 4Q'+Z0&%$ !"+m0&B-(==('+0"%$9 2%$ 4@0=0+Z0&%$9 2%$ 4+m$'0=,-('%,"$ (0&$0 >,% 2%$ =+-#4,-%$ 28(2@#$0+& SF((=(&+ %' W,&'%"9 djjd9 M+&0XX0 %' ^("0&9 djjfTC K- Q ( 40&> =%=/"%$ 2% -( B(=0--% GJmκM ) N%-7 S!kaT9 4mN%-9 N%-M9 GJmκMc S!aj6!cjaT %' GJmκMd S!ad6!cjjTC 3%,< 28%&'"% %--%$9 GJmκMc %' GJmκMd $+&' '+,' 2?(/+"2 !"+2,0'%$ $+,$ -(

(37)

ϯϲ B+"=% 2% !"#4,"$%,"$C I% 4-05(1% %&XQ=('0>,% 2% !cja %' !cjj "#$,-'% 2(&$ -( B+"=('0+& 2%$ $+,$m,&0'#$ =(',"%$ !aj %' !ad9 "%$!%4'05%=%&' SM%0&Z% %' I%Q9 djjfTC I%$ !"+'#0&%$ 4mN%- %' N%-M $+&' 20"%4'%=%&' "%$!+&$(/-%$ 2% -80&2,4'0+& 2%$ 1A&%$ 40/-%$9 4(" 0-$ !+$$A2%&' ,& 2+=(0&% 2% '"(&$(4'05('0+&C P(&20$ >,% -%$ $+,$m,&0'#$ !aj %' !ad !"#$%&'%&' ,& $0'% 2% -0(0$+& ; -873G9 %' ,&% 4(!(40'# -0=0'#% !+," (4'05%" -( '"(&$4"0!'0+& 2%$ 1A&%$9 %--%$ &#4%$$0'%&' 2+&4 2?R'"% 20=#"0$%" (5%4 ,&% !"+'#0&% 2, 4+=!-%<% !+$$#2(&' ,&% 2+=(0&% 2% '"(&$(4'05('0+& SW(Q2%& %' _@+$@9 djjfTC I( 4(!(40'# 2% "%4+&&(0$$(&4% 2%$ 73G 40/-%$ !(" -%$ 20BB#"%&'$ =%=/"%$ 2% -( B(=0--% 5("0%&' (,$$0C F%40 !%,' R'"% 2x ; -( '(0--% 2%$ $#>,%&4%$ &,4-#+'020>,%$ "%4+&&,%$ !(" -% B(4'%," 2% '"(&$4"0!'0+& 4+==% !(" %<%=!-% 4mN%- >,0 "%4+&&(0' 2%$ $#>,%&4%$ 2?73G 40/-%$ !-,$ -("1%$ >,% N%-7 +, !ajC F%''% !"+!"0#'# !+,""(0' !%"=%''"% ; GJmκM 2% "#1,-%" 2%$ $+,$m %&$%=/-%$ $!#40B0>,%$ 2%$ 1A&%$ 40/-%$9 4+==% !(" %<%=!-% GJmκMc 0&'%"5%&(&' 2(&$ -?+&'+1#&A$% 2%$ '0$$,$ -Q=!+g2%$ +, GJmκMd 0&'%"5%&(&' 2(&$ -( "#!+&$% 0==,&0'(0"% %' 0&B-(==('+0"% S_0-=+"% %' _%"+&2(Z0$9 djccTC 3(&$ -%$ 4%--,-%$ &+& $'0=,-#%$9 GJmκM %<0$'% $+,$ -( B+"=% 2?@+=+m+, 2?@#'#"+20=A"%$ -0#$ (,< !"+'#0&%$ 0&@0/0'"04%$ KκM >,0 "%'0%&&%&' -%$ =+-#4,-%$ 2, 4+=!-%<% GJmκM 2(&$ -% 4Q'+!-($=%C 30BB#"%&'$ $'0=,-0 %<'"(4%--,-(0"%$ '%-$ >,% -%$ !"+2,0'$ =04"+/0%&$ %' 2%$ 4Q'+Z0&%$ !"+m0&B-(==('+0"%$ $+&' 4(!(/-%$ 2% 2#4-%&4@%" -?,/0>,0'0&('0+& %' -( 2#1"(2('0+& 2?KκM !(" -%$ !"+'%($+=%$C F%40 -0/A"% -%$ =%=/"%$ 2, 4+=!-%<% GJmκM9 !%"=%' -( B+"=('0+& 2%$ 20=A"%$ (4'0B$ 2% GJm κM9 -%," '"(&$-+4('0+& 2(&$ -% &+Q(, %' -?(4'05('0+& 2% -( '"(&$4"0!'0+& 2%$ 1A&%$ 40/-%$ S.($!("(Z0$ djjb9 *,& %' I%Q9 djjiTC

(38)

ϯϳ .-,$0%,"$ #',2%$ +&' "(!!+"'# >,% P*'0<3+%'+ %' 2?(,'"%$ !("($0'%$ '%-$ >,% !" #$%&'' !%,5%&' 0&'%"B#"%" (5%4 -8(4'05('0+& 2%$ 5+0%$ 2% $01&(-0$('0+& 2, GJ−κΒC I%$ =(4"+!@(1%$ 0&B%4'#$ !(" !" #$%&'' $+&' 4(!(/-%$ 280&2,0"% -% 4-05(1% 2?KκMV '+,'%B+0$9 -( 2#1"(2('0+& 2% 4%''% !"+'#0&% 0&@0/0'"04% &8( !($ 4+&2,0' ; -( '"(&$-+4('0+& &,4-#(0"% 2% GJmκM S*@(!0"( *) +.C9 djjd9 djjfTC .-,$ "#4%==%&'9 0- ( #'# 2#=+&'"# >,% -80&B%4'0+& 2%$ =(4"+!@(1%$ @,=(0&$ (5%4 P*'0<3+%'+ 3+]$6 ( 4+&2,0' ; ,& /-+4(1% 2(&$ -80&2,4'0+& 2%$ $+,$m,&0'#$ (4'05%$ !ka6!aj9 '(&20$ >,% -( '"(&$-+4('0+& &,4-#(0"% 2% 4+=!-%<%$ 4mN%- 4+&'%&(&' &8#'(0' !($ '+,4@#% S_,0X(&0mP(//(&% *) +.C9 djjfTC F%!%&2(&'9 =(-1"# -%$ &+=/"%,$%$ "%4@%"4@%$ $," -80=!-04('0+& 2%$ "#$%(,< 2% "#1,-('0+& GJmκM 2(&$ -%$ 0&B%4'0+&$ !("($0'(0"%9 2% &+=/"%,< ($!%4'$ 2% -%," "L-% 2(&$ -( "#$0$'(&4% ; 4%$ (1%&'$ !('@+1A&%$ &% $+&' !($ %&4+"% %&'0A"%=%&' 4+=!"0$C

9W)>,-'.,'-()*())I#)a#$%II()AN5κ

κ

κ

κTbR(I)

GJmκM %$' -% =%=/"% -% !-,$ -("1%=%&' "%&4+&'"# 2(&$ -( B(=0--% 2%$ B(4'%,"$ 2% '"(&$4"0!'0+& 4+&'%&(&' 2%$ 20=A"%$ 2% -( !"+'#0&% N%-C I%$ =%=/"%$ 2% -( B(=0--% N%- 4+&'0%&&%&' ,& =+'0B 2% $'",4',"% $0=0-(0"%9 -( "#10+& 28@+=+-+10% N%- SNW39 +, NWTC I% NW3 %$' 0=!-0>,# 2(&$ -( -0(0$+& ; -873G !,0$ $( 20=#"0$('0+& (,'+," 2% -873G9 %' 2(&$ -%$ 0&'%"(4'0+&$ (5%4 -%$ !"+'#0&%$ 0&@0/0'"04%$ 2(&$ -% 4Q'+!-($=%C P+,'%$ -%$ $+,$m,&0'#$ 2% GJmκM $+&' 4("(4'#"0$#%$ !(" ,& 2+=(0&% Gm'%"=0&(- 4+&$%"5# 2?%&50"+& ejj (402%$ (=0&#$ S.%"Z0&$ *) +."= djjkTC I%$ !"+'#0&%$ N%- $+&' +/$%"5#%$ 4@%X 2% &+=/"%,< +"1(&0$=%$ %,4("Q+'%$ %' -% 2+=(0&% NW3 %$' -("1%=%&' 4+&$%"5#C P+,'%$ -%$ !"+'#0&%$

(39)

ϯϴ N%- !("'(1% 2%$ $'",4',"%$ $0=0-(0"%$ %' !%,5%&' R'"% 2050$#$ %& 2%,< 4-($$%$ /($#%$ $," -( $#>,%&4% 2% Fm'%"=0&(- 2, 2+=(0&% NW3 SJ01,"% cjTC

)

7'#/6* ^9 3*3N6*0 &* .+ M+3'..* _7ZκI= &E+26H0 :`+-&*% *) +."=AJJ\@

I%$ 20=A"%$ 2% GJmκM $% -0%&' ; 2%$ $0'%$ κM $," -%$ !"+=+'%,"$ 6 (4'05('%,"$ 2%$ 1A&%$ 40/-%$ %' "#1,-%&' (0&$0 -( '"(&$4"0!'0+& W'+ -% "%4",'%=%&' 2% 4+m(4'05('%,"$ +, 2% 4+m "#!"%$$%,"$CI% 2+=(0&% 2% '"(&$(4'05('0+& 2% -( '"(&$4"0!'0+& SP73T %$' &#4%$$(0"% ; -( "#1,-('0+& !+$0'05% 2%$ 1A&%$C K- %$' !"#$%&' $%,-%=%&' 4@%X !ka9 4mN%-9 %' N%-M9 =(0$ !($ 4@%X !aj %' !ad SW(Q2%& *) +."= djjiTC

P+,'%$ -%$ !"+'#0&%$ N%- !("'(1%&' -% N%- W+=+-+1Q 3+=(0& SNW3T9 4+&'%&(&' ,& 2+=(0&% 2% -0(0$+& ; -?73G9 ,& 2+=(0&% 2% 20=#"0$('0+&9 ,& $01&(- 2% -+4(-0$('0+&

(40)

ϯϵ &,4-#(0"% %' ,& 2+=(0&% 280&'%"(4'0+& (5%4 -( !"+'#0&% 0&@0/0'"04% KțMC E--%$ !%,5%&' R'"% 2050$#%$ %& 2%,< 4-($$%$ /($#%$ $," -( $#>,%&4% 2% Fm'%"=0&(- 2, 2+=(0&% NW3

I( !"%=0A"% 4-($$% S.cja9 .cjj %' 3"+$+!@0-( N%-0$@T ( 2%$ 2+=(0&%$ Fm'%"=0&(,< -+&1$ >,0 4+&'0%&&%&' 2%$ =+'0B$ "#!#'#$ 2?,& 2+=(0&% (&ZQ"0&%9 >,0 4+&'"0/,%&' ; 0&@0/%" 4%$ =+-#4,-%$ :7'#/6* 88@C I+"$ 2% $+& (4'05('0+&9 4%''% 4@(u&% 2% =+'0B$ "#!#'#$ %$' 4-05#% !(" !"+'#+-Q$% %& !aj +, !ad9 -%$ "%&2(&' (0&$0 (4'05%$C

I%$ @+=+20=A"%$ !aj6!aj %' !ad6!ad $+&' '"(&$4"0!'0+&&%--%=%&' 0&(4'0B$ %' !%,5%&' =R=% R'"% "#!"%$$%,"$ %& %=!R4@(&'9 !(" 4+=!#'0'0+&9 2%$ 20=A"%$ '"(&$4"0!'0+&&%--%=%&' (4'0B$ S!aj6!ka9 !aj64mN%-9 !ka6!ka9 !ka64mN%- +, !ad6N%-MT 2% $% -0%" ; -%," $0'% țM $," -?73G SW(Q2%& *) +."= djjfTC F%$ 2%,< 20=A"%$ !%,5%&' #1(-%=%&' +44,!%" 4+&$'0','05%=%&' 4%"'(0&$ $0'%$ κM '+,' %& "%$!%4'(&' (0&$0 ,& $%,0- 28(4'05('0+& 2% 4%"'(0& 1A&%$ 40/-%$ 2% GJmκM SW(Q2%& *) +."= djcdT

(41)

ϰϬ I( 2%,<0A=% 4-($$% 2% !"+'#0&%$ GJmκM %$' -( B(=0--% 2%$ !"+'#0&%$ N%- >,0 4+&'0%&' ,&% "#10+& P73 $," -% 5%"$(&' Fm'%"=0&(-% 2% -( NW3 SJ01,"% ccTC F% 2+=(0&% %$' "%$!+&$(/-% 2% -%," 4(!(40'# ; (4'05%" -( '"(&$4"0!'0+& 2%$ 1A&%$ 40/-%$9 %' 4+&'0%&' -% $01&(- 2% -+4(-0$('0+& &,4-#(0"% SGI*T >,0 %$' "%$!+&$(/-% 2% -( -0(0$+& ; -?73G9 2% -( 20=#"0$('0+& %' 2% -?($$+40('0+& (5%4 KțMC

F%''% B(=0--% 2%$ !"+'#0&%$ 4+=!"%&2 4mN%- S%' $+& @+=+-+1,% "#'"+50"(- 5mN%-T -%$ !"+'#0&%$9 N%-M9 N%-7 S!kaT9 %' 3"+$+!@0-( 2+"$(-%$ %' 30B S_0-=+"%= AJ8ATC

I%$ =%=/"%$ 2% -( B(=0--% N%-6GJmκM $% -0%&' ; 2%$ $0'%$ 2% -0(0$+& ; -?73G 4+=!+$#%$ 2% b ; cj !(0"%$ 2% /($%$9 (!!%-#$ $0'% κM) a?m___NGG[[FFme? +p N %$' ,&% !,"0&%9 [ ,&% !Q"0=020&% %' G &?0=!+"'% >,%--% (,'"% /($% >,0 !%,' 5("0%" -#1A"%=%&'C F% >,0 %<!-0>,% -( $!#40B040'# 2% -( "#!+&$% 2?,& 1A&% ; -?(,'"% $%-+& -% $0'% 2% -0(0$+& !"#B#"%&'0%--% 2% 4%"'(0&$ 20=A"%$ (5%4 4%"'(0&$ $0'%$ 2% GJmțM SW(Q2%& *) +."= djjf9 ^("0& *) +." = djjjTC

P+,'%$ -%$ !"+'#0&%$ N%- !%,5%&' B+"=%" 2%$ @+=+20=A"%$ +, 2%$ @#'#"+20=A"%$9 ; -8%<4%!'0+& 2% N%-M9 >,0 &% !%,' !($ B+"=%" 2%$ @#'#"+20=A"%$C

)

DW)O#)a#$%II()*(+)2-/,L%"(+)Mκ

κ

κT))

κ

F+==% (5%4 -%$ !"+'#0&%$ N%-6GJmκM9 0- %<0$'% !-,$0%,"$ !"+'#0&%$ KκM9 >,0 +&' 2%$ (BB0&0'#$ 20BB#"%&'%$ !+," 4@(>,% 4+=!-%<%$ N%-6GJmκMC E--%$ $+&' "#1,-#%$ 2% B(l+& -#1A"%=%&' 20BB#"%&'%9 %' $+&' %<!"0=#E$ 2(&$ ,& '0$$, 2% =(&0A"% $!#40B0>,%C I( B(=0--% 2%$ !"+'#0&%$ KκM 4+=!"%&2 ) !cja9 !cjj9 ΙκΒα, ΙκΒβ, ΙκΒε, ΙκΒζ, M4-m3 %' -( !"+'#0&%

(42)

ϰϭ 2% 3"+$+!@0-% F(4',$ SJ01,"% cdTC E--%$ $+&' 4("(4'#"0$#%$ !(" -( !"#$%&4% 2% =+'0B$ (&ZQ"0&% "%$!+&$(/-%$ 2% -?0&'%"(4'0+& (5%4 -% NW3 2, GJmțMC

I% =%=/"% 2% -( B(=0--% -% !-,$ #',20# %$' KκMĮC K- %$' "(!02%=%&' 2#1"(2# 2,"(&' -?(4'05('0+& 2% -( 5+0% 2% $01&(-0$('0+& GJmκM 4+&2,0$(&' ; -( -0/#"('0+& 2% =,-'0!-%$ 20=A"%$ 2% GJmκMC 3% !-,$9 -?@#'#"+20=A"% !ka6!aj %$' ,&% 2%$ !"%=0A"%$ 40/-%$ 2?KκMĮC

7'#/6* 88 9 Q*3N6*0 &* .+ M+3'..* aκI= &E+26H0 :`+-&*% *) +." AJJ\@

HW)M",(-#.,%/")ANκ

κ

κ

κTb)Mκ

κ

κ

κT)V

I?(4'050'# 2% GJmκM %$' "#1,-#% !"0&40!(-%=%&' !(" -?0&'%"(4'0+& (5%4 -%$ !"+'#0&%$ KκMC I?0&'%"(4'0+& -( =0%,< #',20#% %$' 4%--% 2% ΙκΒα (5%4 -% 20=A"% GJmκΒ !aj6N%-7C F%''% 0&'%"(4'0+& /-+>,% -( 4(!(40'# 2% GJmκM ; $% -0%" ; -?73G 4(" -% 4+=!-%<% GJmκM6 ΙκM !%"$0$'% 2(&$ -% 4Q'+!-($=% %& "(0$+& 2?,& $01&(- 2?%<!+"'('0+& &,4-#(0"% B+"' 2, ; ΙκΒα.

(43)

ϰϮ 7,'"%=%&' 20'9 -% 4+=!-%<% GJ−κΒ/ΙκΒα %BB%4',% 4+&'0&,%--%=%&' -( &(5%''% %&'"% -% &+Q(, %' -% 4Q'+!-($=%9 =(0$ $+& '(,< 2?%<!+"'('0+& &,4-#(0"% 2#!($$% $+& '(,< 2?0=!+"'('0+& %' 2+&4 -% 4+=!-%<% "%$'% 1#&#"(-%=%&' 4Q'+!-($=0>,%C ΙκΒα %&'"% %& 4+&'(4' B"#>,%==%&' (5%4 GJmκM %' 4%''% 0&'%"(4'0+& 4(4@% 2%$ $#>,%&4%$ 2% GJmκM >,0 $+&' 0=!+"'(&'%$ !+," -( -0(0$+& ; -?73GC 3% =R=%9 -+"$>,% ΙκΒβ 0&'%"(10' (5%4 -% 4+=!-%<% GJmκM9 -% 4+=!-%<% %$' "%'%&, 2(&$ -% 4Q'+!-($=%C :M'#/6* 8C@

7'#/6* 8A 9 c% 3$&H.* &* .+ &Y#6+&+)'$% &*0 W$'*0 &* 6Y#/.+)'$% aκIα &+%0 .*0 ,*../.*0 N+0+.*0 *) 0)'3/.Y*0 :[<$0< *) +."= AJ8A@

G+'"% 4+&&(0$$(&4% (!!"+B+&20% 2% -( 5+0% 2% $01&(-0$('0+& N%-6 GJmκM =%' %& -,=0A"% #1(-%=%&' &+$ -(4,&%$C G+,$ (5+&$ %&4+"% '"A$ !%, 2% 4+=!"#@%&$0+& 2% 4%''% 5+0% 2% $01&(-0$('0+& '% W'W$ (, "%1("2 2%$ 4+&20'0+&$ 2% B+&4'0+&&%=%&' 2?,& =+2% 2% B+&4'0+&&%=%&' !@Q$0+-+10>,% %' &+& !('@+-+10>,%C I( 4,-',"% 2% 4%--,-%$ %& -01&#% +,

Références

Documents relatifs

To test whether the vesicular pool of Atat1 promotes the acetyl- ation of -tubulin in MTs, we isolated subcellular fractions from newborn mouse cortices and then assessed

Néanmoins, la dualité des acides (Lewis et Bronsted) est un système dispendieux, dont le recyclage est une opération complexe et par conséquent difficilement applicable à

Cette mutation familiale du gène MME est une substitution d’une base guanine par une base adenine sur le chromosome 3q25.2, ce qui induit un remplacement d’un acide aminé cystéine

En ouvrant cette page avec Netscape composer, vous verrez que le cadre prévu pour accueillir le panoramique a une taille déterminée, choisie par les concepteurs des hyperpaysages

Chaque séance durera deux heures, mais dans la seconde, seule la première heure sera consacrée à l'expérimentation décrite ici ; durant la seconde, les élèves travailleront sur

A time-varying respiratory elastance model is developed with a negative elastic component (E demand ), to describe the driving pressure generated during a patient initiated

The aim of this study was to assess, in three experimental fields representative of the various topoclimatological zones of Luxembourg, the impact of timing of fungicide

Attention to a relation ontology [...] refocuses security discourses to better reflect and appreciate three forms of interconnection that are not sufficiently attended to