Ataxia telangiectasia and Rad3 related kinase drives both the early and the late DNA damage response to the monofunctional antitumor alkylator S23906: S23906 is a potent inducer of replication stress
Texte intégral
(2) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d ce Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t.
(3)
(4)
(5)
(6)
(7)
(8)
(9) &'( &'(
(10) !"! #$!% )*+!,, %,!"" ) -" !. #//&'() !,,*#&'(0) .#1#!$ 2*!%033)!,,!$%!&'()+"!4+$!%/$5*! "&'(
(11)
(12) !! " # $ % & '
(13) ($ %)*+" , - . & '&$/01" 2 3 4 56
(14) $ % " 7 8 99 - $ %
(15) :
(16) :8 ;<0/=* ; 3) : >) >
(17) ?@ AB 1C !! :" )>4;: )>D4) ? E; 3. 4 3. @
(18) @ 3. . Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. .
(19) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma pte d ce Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. .4 % 4 4 3.) 4 F @ 3. 4@ 3.4 F %4 :4 %%4 ) @ @ %4 F % F 4 3. F ) A F 8 3.4 4 % % <0/=* 4 4 F 4 % 3.)<0/=*:4 4
(20) @ % )- '. ('!=.+ ' ( : 4 + <0/=*) - (' 30 + % A<G
(21) @ F
(22) @<)'% 4 @ @ H3 *< 4 % F <0/=*) 3. 4 <0/=* %4 4 <0/=* 4 F @ ). Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. <.
(23) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t.
(24) >@ %% 4 44 3. ) 4I 3. 4 F 0@ @ @ (-??+ : 4 ('+ ' 30 (+ J<K)' 3. @(3+ 4 '!=. ('.+ 4: % F
(25) 4 4 . J0CK) ' F : % 4 % /1 4 %4 4J0K)- 4 4 4 3. J!K) 4 @ % % 4 4 J* K) (-+ (+ 3. (3.+ 4 % @ J1K) 3 - 3.L 4 -4: 4 J=<K) " % L ( 4 + @ F /4:J0 K) ' % 44 @A<G ) " :4 ' % @
(26) @< F
(27) @ @ J1/K) % @ F F F % F F % L % ) " :4 % 3. 4 $M : (A$+ F ' % J<=<K)- 4 ' % ' % :J<<K)
(28) % % F 8 (-+ F ' '. J<0<CK) 3. 4 4 L % -??) " :4 4 0< 3.? J<!<*K) 4 0< F $M A$ : F F @ L ' '. J< 0=K) % 3.4 4 4 4 3.) ' 4 : 3 4 J<=000K)
(29) F @
(30) < F 3. F J0CK) - F 3. 4 % ) $ 4 F4 : @ % @ J0!K) AF% F @ @ %4 J0*K) H3 *< %
(31) @6< F 4@ % 4 % 4 Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. 0.
(32) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. @ 3 J0 01K) @ "=C 0* F
(33) @ : % 4 F % J0/C=K) - 4 F F @ J0 0/K) 4 F %%4 F 4 J<=CC0K) % @ @ F 4 4 ) - 3. F @ 3. 4@ 3.4 F )":4 F @ 3. 4
(34) .$ (0(< + + : JCC0K) '% F 4 %%
(35) @ F 4 JC0K) 3. 4 L 4: F 4 @ ) 4 '3.?JCCK) - 8 % 6
(36) @ F F % '6
(37) @< JC!C K) %% 3.4 % F 8 F ) - F 8 3. 4 <0/=* % % F % ) <0/=* 4 @ 4 F : 4 4 % 3. : JC1C/K)- 4 4 <0/=* : F 4 3. 4 3 F @ F J0<!=K) - <0/=* 3. 4 F @ J!K A<G J!<K)N F 4 A<G F <0/=* : F 3. F %)- 4 % A<G
(38) @ F
(39) @<) '% 4 % F<0/=*) 3.4 <0/=* %4 4 <0/=* 4 F @ )
(40)
(41) . 6!7 /"% % %<0/=*(< : 0C 00C 4 * 4 : A 8(+ (0<+ + F 4 - % (
(42) " + F .01 ( = : 4 + % 4 4 F 4 (GO
(43) +) 3: $
(44) .= F 4 4 ( P "% " +) H3 *< F 4 : '4 (7 . +) F 43 (' 474 +) Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. C.
(45) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. !""/*",*$! A F F 3Q 4 >Q '4 (- M
(46) " + 4 F =R 4 ( S" +)E (7'=0*! +@ (7'10* TFU+ : (+ 4 (7'=01/ ' + F F '-*C= 44 (- M + 4 F !R 4( S" +)E 7'10* TFU % 4 @ 4 F 4 (<=7+ F : J!0K) @: F 47'1C MC= 4 F 3. @ % : % 4 J!CK F 4 '>' (- M + 4 F =R 4 < 446 4 (> $ " +) " %: @ % @: F F ! V64 : C1 :4 ) 4 @: F @ % " (- 7 % MI " +) 4 F 4 F == 64 ==V64 4 ( 4"S" + <! V644 ( 8% " +) , 8#+ !% *1
(47) @<(W <**+
(48) @< (
(49) < W 0CC=+ 0
(50) @ (W <0CC+ 0C!
(51) @ (W <0C+
(52) @ ( <73! W <0*=+ /1' ( =A)>< W C!<*+ ' ( 3<>< W<1 0+ 0< ( C>C W <<=1+ = (W <<* + F 4
(53) (N8 4 P X% " +) (W *1 C+ (
(54) ! W !CC<+ F 4 3 (
(55) : " + 4 % ) A<G(W= *< + YA<G(W=!*0*+ F 4 '(@.X+)'( <3 +!=( 00+ . F 4 .% (- 4 ' " +)" 4 ( '<=W*< +( >CW<0 0*+
(56) <!( "*W 01/+ F 4
(57) 8 ( X% " +)A I F 4 Z@ -44 ( A'>+) . !"" 9/"!"9% % " :4 F % J!!K) AF: V' <0/=* F * 4) E F F == ' $
(58) .= V' <0/=* * == ' $
(59) .=) @: F ! V64 : C1 ) @ F : V' <0/=* F < 44)
(60) :4 F F <0/=*L V' F 4 -
(61) != % <0/=* 4 ' )
(62) F F :F =R )
(63) F 0=4 4 4 !V643-(4 Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. !.
(64) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. + ==V64.(4+)4F 8F -- F 4 3-M F(3 " +) 77*#:"*#$!%/!/!+7 /$#%/#;9
(65) F F <0/=* 0 [
(66) 4)" : 4(+!V'<0/=*F % % 3. 4 A 4 @ )E F < (" ! !
(67) + F < V' <0/=* % J!<K)-44 :4 F % J!*K) E %FF F
(68) ? (!= 4' .
(69) 0 4' '
(70) < R G== != 4' A>> A )C 0= 4' + @ C[
(71) !4 : CR 4 (> ' A $))+) F % 4 ) 3. F 3-) F >$(! . <Q: + (
(72) @\ >$ : "] C11 -4 ? W
(73) =00 - M
(74) " + 0= 4 ) >$ F 4 O ) -4 F G* 4 " 4 F (N 4+) " F4 ' ' F($ %-4
(75) 3F F +) @ % F ) == F 8 4) M % F :4 ) 77*#8"#,, 5
(76) F F <0/=* 0 [
(77) 4 F 8 0= 4 C[
(78) 3 4 ) % FF F
(79) ? (!= 4' .
(80) 0 4' '
(81) < R G== != 4' A>>A )C 0= 4' + @ C[
(82) !4 ) F 4 !== !4 )" : 4(+!V'<0/=* F % % 3.4 ) E F <C ("07 !+ <0/=*F <V' % J!<K) F % 3 4 (!!R+ 44 (+) '4 F 4@ (!= 4'A1)=!=4'.
(83) =)!RF <= !R @444@+ F % 44 ) F 44 @ (4 +)
(84) !;"!, ##: #$89%7"" ,!$:!$ 5 4 . (.+ F 4 4 Q F A (P
(85) " + = 46 ). : <! F 4- M ). L !Q
(86) $$
(87) 777$$7
(88) $$$
(89) $$$$$0Q). L Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. *.
(90) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. ' !Q $7
(91) 7$
(92) $$7$
(93) $7
(94) $0Q) 4 . ( .\ . . %
(95) N 7
(96) + F 4 - M ) 4: F 4% * F F 4 44 C1 4 F <0/=*)
(97) F 44 44 %) 8 " ,9%%9% F % <0/=* F 4 ' (4 8 844+% % J! K)%% :4 ) @: F ! V64 : C1 % ) @ : F : <0/=* F ! 4 44 : ) % @%: F : <0/=* != 'H3 *<) F F ! 4 != ' H3 *<) 5"!/!""5!"!"!/,$#;6#$!% %</#7!,%%9= 4 F : <0/=* 0 [
(98) @ I @ J00K)-4 F 4 ?4 !)! F(? -4 . 4$?+) ==F 8 ) %;!/ : /""9 +*/!%!$"9
(99) +75!$!%;#%! $!;" /, 5/!""% F A<G A F F >$ 4 4 F <0/=*) E A<G % <0/=* : F 4 4 A<G ) -4 4I F % A<GF % >$F A<G F% % ! V'(" 4 " +)" % <0/=* A<G F F 3. F % (" + 3. 4 4 % 3. ) <0/=* : F 4 4 ) - : <0/=* ( V'+ F * 4 F 4 4@ 4 ("
(100) 4 04 + 4 % : <0/=* FF ) F 4 $
(101) .=
(102) @
(103) @<@ @ ("
(104) 4 0 C +) - <0/=* F4 $
(105) . = F== ')
(106) @F %%
(107) @< L $
(108) .= J!1K) 4 <0/=* Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited.
(109) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. F @ 3. 4 %8 A<G 4 ) +#!%#,;$, / ;,! ,6!!$"9
(110) +75!$!%;#%!,# @ 3. 4 <0/=* F 4 % ') -44 4 F 4 % ' 4 ' /1 : !V'<0/=* <== '.01("<+) F <0/=* 4 ' 4 F.01F % ' (" <+) - <0/=* F ' 4 J!/K ' /1 4 F <0/=*:("< F 4 " <+) - 4 F ' <0/=* 4
(111) @< *1 ("<4 < 4 + 4 F .01: ("<04 +)% % ' F <0/=*:) 4 4 F 4 44 44 ) F <0/=* % '
(112) @< 4 F .01 (" <
(113) +) " 4 <0/=* : F 4 4 != '
(114) ? : F F% : .01("<3+ $M J*=K) F 4 . 8 <0/=*:("<>+) 3.4 <0/=* 4 %''.) +*/!% $!/$* ,7!, #: ,# /, >!"9 $!;" /, 5
(115) $!5 #% + /, >,!%,6! ? 6@A;,6B9 3. (3.+ 4 % @) ( + <0/=* : F 4 4 4 =6 (" 0+ F F % (" 0 4 " 0+) 4 F = F L % 0<6<("0+)<0/=* F F % 3. (" 0+ <0/=* 4 % ) - 4 F <0/=* : F 4 0< F F% .01 (" 0
(116) + @ <0/=* .01 ) 3. (3.+ 4 - 4: 4 J1K F L 4 4 % J0! K) <0/=*: F4 4 F 44 4 (" 03 F+) F 4 Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. 1.
(117) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. % 4
(118) @ 0 0C! (" 0 > "+ F F % % (4 " C !+) %
(119) @ F 4
(120) <!F 4
(121) @ 4 (" 07+) F 4 % <0/=*:) 6!!$"9
(122) +75!$!%;#%!,# % C+!;!+!, " 8 4 3. 4 <0/=* F 4) " <0/=* A<G
(123) @ F 4 % 4 ;7'=0*! ( 4 +7'10* (@ TFU+ 7'=01/ ( : ' +) <0/=*: 4 ' F4 4 4% A<GF A<G F% @ (" C C+)
(124) @ <0/=* F 4 @ 4 4 ' ("C+)-4 @ A<G
(125) @ @F 3. 4 <0/=* : 4 % 3.4 0 4 @ 4 ("C
(126) +) - 4 %: @ 4 (@+ 7'1C 4 (" C3 + 4 <0/=* A<G F F% .01 (" C>+) - % 4 4 = F <0/=* : (" C"+) F 4% 3. 4 L A<G ) 4 3.4 <0/=*. :4 F A) C1 . % ' F @ @F <=R %("C7+)-4 F : F@ @F
(127) @ 0 F 4@ F <0/=* 4 (" C7+) - ' @ @F
(128) @ ("C7+)'%@ @F '@ @F A<G 4 F <0/=* 4 (" CA+) @ F 3. <0/=*L@ % ') :"*!/!#: #,6!+!"9!+$!%;#%!,# <0/=* : F 4 4 A<G F % F (" !+) - F F 4 A<G < * A<G F % 4 % F F % (" !+) Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. /.
(129) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t.
(130) @ F F % % ("!+ F @ A<G(" !+) - % 0< (" !+) <0/=* F % A<G
(131) @)L 4 3 % F A<G F 0<) *1
(132) @< F <0/=* 4 F 4 @ A<G (" ! !
(133) 4+) - %
(134) @ A<G F *1
(135) @< (" ! !
(136) +) 4 %
(137) @
(138) @< %% J/K) 4 4 4 %
(139) @
(140) @< 7'1C %: @ F : <0/=* F < 4 F 4 ) F % F <0/=* (" !3 4 < C +)- F 4 <0 % <0/=* (" !>+) @ <0/=*) D
(141) EE)#>!"/6!/@;# ,8$#5,#$;#,!, ,!% /, > ,9 >% F % 3.4 % % @
(142) @
(143) @< (" ! !
(144) +) @ @ :4 @ ()) '+ 4 % 4% : % <0/=* % ) <0/=*F 4 F % @ H3 *< %
(145) @
(146) @< J0 K) F " !" H3 *< : % <0/=* :4 ! ) 4 H3 *< F % A (4 " *+) - H3 *< <0/=*F4F H3 *< .01 4 F % (4 " +)
(147) <0/=* 4@ 4 3. F JC/K) $ % 3 F % @J0<!=K) <C<0/=*: F A<G 4 F J0<K 4 F % 4 3. J<=000K) - @ F <0/=*) - 8 A<G 4 F 3 F F @ ) 4 % 6
(148) @ F 8 @ 4 3. J!K) % 4 Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. =.
(149) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. L 4 3) @ F % F <0/=* % 4 <0/=* 4 F 3.4 ) AF 4 A<G F <0/=*: F A<G F 3. ) 4 4 3. 4 @@ 4 % )
(150) F 4 3. 3.4 :% ) - '. ' % 4% ())!V'+) AF% F 4 % A<G % 4 % F % ) % A<G : F :%4)" 34 4 3 4 @ J*K % ' 3.? $M 3. J<*K) - 4 F 4 % %
(151) @
(152) @<@ F : <0/=* 4 4 F % F @ @ ) % F 4 % F @ F 4 J<=K F @ % 4 A<G) '. A<G F ' L %
(153) @< J<=K) - 4 F F 4 F F %
(154) @< % F <0/=* :) - F 4 $M % J<<<C*K '6
(155) @< F % % F <0/=*:)0< 4 4 4 8 J<!<*0K 4 4 F @ 3. 4 4 J*<K) 44 3.4 3. 4 % 4 L 4 4 % ) -4 % <0/=* 6
(156) @ F F ( %: @ + 4 (F $
(157) .=+ F 4 )% 4 %
(158) @ 4 F 4 F 3. J*0K)
(159) @ : % 43.4 4 F 76 @ J0 0/K) '% F 6
(160) @ F 0 A$ % J*CK) 4 <0/=* F @ 4 : % ) - 4 FF F . Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. .
(161) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. sc rip. nu. Ma. pte d. ce. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. t. @ @ H3 *<$
(162) .= : <0/=*) - 4 F @ ) - %
(163) @6< JC<C0K) 4 :
(164) @ 4 F 4 %% % F J0*K) - @ 4 4: <0/=*)- @ @ 3. @ F @F 3. 4 3.4 ) 8 L 3. 4 @ %4 @ @ ) - 4
(165) @6< @ H3 *< % 3 4 4@ F 3 % 4 H3 *< 4 J01K)- F F 3 F % 4 ) 4 @ F : F 8 % % @ F L 4 3. 4 F ) 3 7) 44 4 % ) 4 4 ) 44 44 )
(166) & Z) 4 ) ' 44 % ) Z^ ) ) A L ) ?) % ) ? 4 ): >)> % ) : >)> 4 )
(167) F@F
(168) 33% - %
(169) "
(170) >6
(171) N">
(172) $(" 8 % .!106= +)3 7)F
(173) (
(174) +MI " ) ' F
(175) _^ _4 .5% (
(176) >+8) X) (<==0+ ' @ ; 4 ). )%)
(177) !!*1). Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. <.
(178) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. . / = < 0 C ! * . t. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. 1. sc rip. *. nu. !. Ma. C. pte d. 0. 4 ) ) (<==C+ - 0@ @ ; OO F )3.(4 +11011 ) @@
(179) ) Z) ? ') ) (<==0+ 3. 4 % ' 4 4 ) . FAC//!=*) "@ Z)
(180) Z) Z@ ) ) (<==!+
(181) % 4 4 ' 3.? 3. 4) . FF*=!*)
(182) 4 ?) )
(183) 8 3) (<==1+ ; 4 ). )%)')
(184) ))***< ) . ?) ) ' ) Z) 4
(185) ) E) E )) (<==<+ F4 4;3.4 @ ) )%)7 )* *!*) 3)
(186) 8 M) 7 Z) (<==C+ ' 4 3. ). )
(187) ))*C1*!!)
(188) 8 3) 7 @ ) P Z) > )Z) (<==+ -; @ ) F 0 *)
(189) 8M) 3))Z)
(190) 4?))7 4 ') 7 Z) (<==0+
(191) 3. 4 @ 3. )')
(192) AA<=0<0) H ) > ) Z) (<==0+ 3. 4 - 3.4:) !C<!C1) ))@')X')
(193) )E Z)
(194) )
(195) 4?))(<==!+ " '
(196) ' 3. 4 % % @ )7 3%)A=C==!<) A))' Z))
(197) 83)(<==!+- 3. 4 - 8
(198) @ )'))
(199) A<0 <<01) ?4Z)>)'% ))4 3)-)
(200) Z)(<==!+A4 4 4)')
(201) ))G=/= =/!) ?4 ) Z) X A) X) 3 E) 7) (<==*+ % -4:)
(202) AF/C0/!!) )@@Z )' .)@
(203) ) @Z) @Z) (<==*+
(204) F 4 F
(205) @ % )')
(206) ))*=!**=*C) 3:)E Z)')? ')X44 ?) ? 8) ') (<== + /A (/+ 4 % @ %)7 3%)AC <C ) ' 3) ) 7@ 7) 7) H )
(207) 8 3) (<==1+ % - -?? 4 ) 7 3%) C 1C1/) Z8 ) "@ Z) @
(208) ) @ Z) 4 7)
(209) ) @ Z) Z@ )) (<==*+ ' 3. @). )
(210) )H0 C!) A)
(211) ) X '))(<==/+? 3. 4;
(212) @
(213) @< '?<)
(214) ) N )
(215) ) A<C!<!!) " )@4A)H)X) 7)Z)
(216) ) @% N))3))@>))
(217) )
(218) A)). 8F. ce. <. 1. /. <=. Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. 0.
(219) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. <*. < . <1 </ 0=. nu. Ma. <!. pte d. <C. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. <0. ce. <<. sc rip. t. <. )I4 ')-) E)') 44X)(<==0+ A<G % ' != . 3. @ 444 3. 4 - % 4:) Z) )
(220) 4) EH <=0=0 <=0<) ) E@ ) )
(221) ?) 78 ) ) 4 >)
(222) ) NO3 ') Z ) ) (<==*+ % ' $M 4 @ )>'NZ)G! !! 1<) @4 A) M) ) N) " ) ' H) A) ' ) H A) 44X)(<==*+3 %' %
(223) @< : 4 4 3. @) Z) )
(224) 4)HA0=1C<0) 7 ') ?)
(225) ) I` ) "@ Z) A ?) % ?) @ Z) H ) ) @ Z) ? ?) ?) (<==0+ : 4 ('+ .
(226) @ 0 8 ) Z) )
(227) 4) EH C1=*C1) ' Z) )
(228) 8 3) (<==*+ % 8 L' ')Z))
(229) 4)HA/0C*/0!=) )7)
(230)
(231) )G)H A)? ?)E)E')) 44 X) (///+ 4 3. 4 4 3. @ ;3.?4:)>'NZ)AH0/ C=*) @ ) ?) -I4 X) N@ .) ) ?) @A)(<==*+3 %%4 0@ @ < 4 3. @) 7
(232) AA<0 <C*) N@ 7)7))-)XZ) '))X @))H @ ?@ ') ? ?) ?)% ') ")
(233) ') ) 3: ?)(<==+ $M 3. : ' )'))
(234) A//<0) Z) 7) > Z) 3: ?) N@ 7) 7) (<==C+ '!=. 4: 8 @)Z))
(235) 4)E0C1=<0C1=) N >) .%
(236) ) Z) ?4% M) " >) E G) (<==*+ < F 4 @ )Z))
(237) 4)HA0/! 0/!00) ' ?)
(238) ) N ) 7 8 Z) 7) 34 % 3) > Z) N@ 7)7) (<== + . 4 F @ ) Z)
(239) ) )AC<<C<</) Z) )? )) 4 ) ) ' ) (<==0+ - 3. 4 8 % @)')
(240) )CC ) & ) ? ) 7 ) ') 3%
(241) ') A)
(242) ) : ) M) ) ?) & ). 0 0<. Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. C.
(243) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. 01. 0/. C= . C. t. sc rip. nu. 0 . Ma. 0*. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. 0!. pte d. 0C. ce. 00. 7 @ %.< 7@ <0/=*)(<==*+
(244) ) <=0 <=) 3)7)>)>) M) )74 ) 3) A L Z) ) )?) (<== + 4 4 3. @ 4 4@ C0)). ))) $AF0=*<0=* ) " )
(245) ) 3) )X ) ) )E )'% @ ') ' 4 ) F A) ) NO
(246) ') Z) F )
(247) 4 Z) ? ) ) Z) A) Z) ) (<==/+ - (3+ 4 4 4
(248) 4 ).)> )Z)')A)<00C) " >) A) @ ?) 4 ?) ? .) ? A) ?F ) ? ) 4 X) (///+ 4@ % $
(249) .= 4 )
(250) )G =!C=*=) E Z) ') ? 4 ) A) (<==+ )4 000C) H ) 3) 3
(251) ) 7 ') ) ) ') F )
(252) )) 7 ) A A) ) A
(253) ) ) Z @ Z) E) 3) ' >) ) Z) ) P%
(254) ) F 8 7) ?) ?)Z)).)E@7)>) E )')(<==1+H3 *< % @ @ % @ 3. )')
(255) )E</!!</**) '. )
(256)
(257) ) @ ) A) H ) 44 X) F 8 7) ) (<==+
(258) @ % % @ 4 ' 3. @ )
(259)
(260) //!==C) ) A Z)
(261) >) ') >) ? % >) Z) 7 ). @%)
(262) ).)NO
(263) ) ?) (<==1+ 3. 4 @ % "==C 0* % 44 @ @ )')
(264) )E)<0/C <C=C) .)' 7))
(265) )E )7)E >)E ?)3) )F 87)?)7Z) ).)(<==+- 4 (7>'+ 4 F "==C 0* (" 0*+ %
(266) A?@ )
(267) N' <==H0=*<)
(268) ) Z ) ) @ .) N4 ") 3) 3) "4 A) ) (<==/+ 3. @ 0(< + %
(269) @ F 44 F )')4)EG0!*0*0) F?)) ?)?) >))% E))? 4 )A)
(270) E) ) (<==/+ : 0 @ %% : ) ')
(271) )H1!!1*0) E Z) ') ? 8 ) ') (<==/+
(272) 3. 4 % @ 4 4%%)')4)E<=1<C). C<. C0. Licenced copy. Copying is not permitted, except with prior permission and as allowed by law. © 2011 The Authors Journal compilation © 2011 Portland Press Limited. !.
(273) Biochemical Journal Immediate Publication. Published on 07 Apr 2011 as manuscript BJ20101770. C/. !=. !. !<. !0. t. sc rip. Ac. THIS IS NOT THE VERSION OF RECORD - see doi:10.1042/BJ20101770. C1. nu. C . Ma. C*. pte d. C!. )
(274) ) ' ) -) 4 ) ) (<==!+ A4 3. @ (44+A<G 44 ; %%4 0@ @ 3. @ 4%4 )')
(275) )F) !1! Z) ) ? ) ) ' ) (<==0+
(276) 4 @ 3. 4 % 3. 4 )' ))G)<!<* E Z)E )E X)'@
(277) )@Z)
(278) 4
(279) )M >))E )Z) >) E E) (<==C+ '
(280) A?< % % )Z))
(281) 4)E) 0/!1C/<) E X)E ) Z) >) E E) (<== + - %
(282) A?< % !0 ) 4)4)EC*/1=) 3%
(283) ') A) E) : ) ? ') 7) &)A@4 Z))
(284) )(<==<+@ 3. <0/=* % 4 4 % 4 @ )4 FA////<=) 3%
(285) ') A) E) : ) ")
(286) ) F>)')L ")?')A@4 Z))&)
(287) )(<==!+
(288) % 4 8 3. : 4 3.; L L % 3. 4 4) ')
(289) )F 1=)
(290) )Z) M)3)7)N ?)>) ) 7& >)74 )A LZ))>)>) )?)(<==+.> G
(291)
(292) >
(293)
(294) % %3. <0/=*;4 4@ )4)4)H00!0C0 & ) &8 M) 4 ) 3) L ") ' ) ? ') ) 7) A@4 Z) ) & ) (<==+ - > 3. % % %<0/=* % 4 )')4)0100/)
(295) 8.) & )'?)M -)E ?) ) & ) A@4 Z) ) & ) 7 ) ') (<==+ 3. 4 <0/=* 4 4 )
(296) )H 11 ) NO3')88M))E
(297) )7)Z)) 7 Z))(<==0+ 4 : : 0 (+ @ 4). )7 ) C/ !=)
(298) E) )
(299) ) Z)
(300) 4 ?) )
(301) ) ') 4 Z) ) ) ) " ) A) (//1+ N%: @ % % 3.4 @ )>'NZ)AE)!/*/) @ F@)
Documents relatifs
An environmental study of a certain community suggests that t years from now the level of carbon monoxide in the air will be changing at the rate of 0.1t + 0.1 parts per million
Chez les poissons, l’effet Root observé pour les hémoglobines anodiques permet la sécrétion d’oxygène dans la vessie gazeuse (qui régule la flottabilité) et au niveau de la
Dans cet article, nous avons présenté le cadre méthodologique adopté pour construire l’ontologie modulaire de la cuisine numérique en justifiant le choix de la
نيدمتعم هوقبس ءاهقفو ءاملع دنع نًسافتلاو ليواقلأا هذى تدرو ثيح ،اديدج يسماغلدا فقوم نكي لم ونوك يميىاربلإا رودلا تاءارقل رظنلا ةرورض مزلي ام ،ةينآرقلا
:( ﺎـﻬﺘﻘﻴﻘﺤو ،اّدـﺠ ةدودـﺤﻤ تﺎّﻴـﺼﺨّﺸﻝﺎﺒ يواّرـﻝا ﺔـﻓرﻌﻤ نوـﻜﺘ ﺔـﻴؤرﻝا نـﻤ عوـﻨﻝا اذﻫ ﻲﻓ ﺔــﻴؤرﻝا ﻩذــﻫو ،نﺎــﻴﻌﻠﻝ رﻫﺎــظ وــﻫ ﺎــﻤ ّﻻإ
Previous experiments have shown the effective restitution coefficient to be a function of a single parameter, namely the Stokes number which compares the inertia of the solid
Objectif : Identifier les impacts des antécédents d’abus sexuels sur le déroulement, le vécu et la peur de l’accouchement, dans le but d’améliorer la prise en soin de ces
l’énigme 1 (les voies de l’open access : voie dorée et voie verte) dans l’ordre des couleurs qui apparaît à l’écran (les articles ont un ou deux carré(s) de couleur dans