Protein fragment complementation assay for studying viral protein-protein interactions

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6 ; 1 20 , * , @ -! -! % ! ! ! - $ = ! < ! ? -! $ @ =& ! " $ % ! -$ 6 < ? ! $ % < ? ! " =& " =& $ % <- ? - % ! ! =&$ = ! - -! ! ! - ! $ 1! ! % ! % ! ! ! ! -! < > 0 ! %'((7?$ 7$2$9$2$ 1! =& ! ! ! $ 1! - - %! -<E =? ! ! <#:? ! - <6= ?$ 1! ! ! - ! % ! ! $ 1! ! - 2(( ! " - ! ! ! ! - ! ! $ 1! ! ! ! G G $

6 2( & ! ! - " ! % - - ! - $ 6 ! - ! > ! < > @? < $%'(('?$ ! %- ! < :? < I" !% - ! - ?$ 1 ! % - ! 79 72 < 9? ! $ - ! ! α ! @%40@ 2%40@ ' #4@ 2% ! β ! > @ 4> @ ! γ ! # @$ & - ! ! - $ 1! ! - % ! <=#>?% ! ! G ! < - $%'((9J ! $% '(('J 4 ! % '((9J= - $% '(('J: $% '((2JK ! $%'((2?$ 1! 79 - 66 > ! - % ! ! ! ! #: ! ! ! - - $ 1! 72 - ! ! - ! - = " <= 0? < I" ! $%'(()?$ 1! - ! ! - ! ! " ! ! - $ ! % ! - - 79 72 40@ 2 < %'((A?$ 9 ! ! - - -! 79 72 - ! % ! - $ 1! - ! 79 72 - ! ! γ ! 4@)B < ! )B?$ 1! - ! ! ! - ! % ! ! $ > -! ! 79 ! - B(L ! $ . ! ! 72 ! ! )(L % ! $

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6 2B 7$'$2$7$ 1 ! & ! - ! ! %! ! $ ! =& ! ! ! ! ! ! - ! - $ ! ! ! ! ! - $ & ! ! =& $ 1! ! - % ! . &K 2A(27B% ! - ! ) ! 40@ 2%- =& ! <= - %'(('J , D $%'(((?$ 1! ! % ! % ! - - " " !

! =& ! =& =&

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6 '' <1& ? ! G ! ! -< $%'((2J: $%2;;;?$ 7$7$'$ ! ! - %! < >&? ! $ 6 ! K'4 % ! >& - ! " ! $ 1! >& - -! ! $ 6 >&% $ 1! - ! - ! ! $ 4 % ! -! ! % ! < 3?$ & % " $ 1! >&% 2;;9 - M ! @ ! % = > ! - G $ 6 ! -! - G ! ! - G - - G $ & - ! - ! " ! >&% ! ! ! - < $% 2;;;? ! β β <: $% 2;;3J. ! $%'(('?J ! 7$7$7?$ 1! ! " ! - G -$ 1! - ! ! - ! ! - ! $ 1! >& >% ! - % - ! F K <F! ! $%'(((J4 $%'(('?$ 6 - -< $%'(('? ! % - ! -! ! - %! ! - $ >& - ! ! % % % $ <= $% '((AJ0 - $%'((2?$ 1! >& -% ! ! ! - - ! F >& - % =& - <: ! %'((9?$ 4

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6 ') ! - % A(% ! - - A7% ! >&$ 1! ! - -! A( ! ! ! ! - - ! A( A7 ! ! ! > @ $ 7$9$'$ 0 ! 79C 72 ! E - ! ! > @ A( A7% ! ! >& ! ! ! $ & ! - - ! % - @ 40@ 2 α ! %4> @ > @ ! β ! -# @ 4@)B ! γ ! - $ ! " ! 79C 72 %! ! $ 1! - ! $ 0 ! - ! ! - $ ! % ! ! >& ! - ! ! - - ! $ 7$9$7$ # - ! ! ! ! ! ! >& 1! - ! >& $ A( A7%! A( A7 ! 4> @% - ! - ! - ! $ ! 4> @ - ! ! >&% ! $ 0 ! - ! - ! A(C A7 $

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'B F : !%F F : !%F 4 - - & ! %F 4 #> & ! ! %F 6 " : !%F 6 1F : !%F 6 %F + 0 %F F 6 %F ! < >' )4'E? %F β ! 0 %F ! : !%F ! 0 %F - # %F # < # ? 6 %F = > %F = - <=>0? 6 %F ='R =1& 5 %F = E %F 0 < ! ! - ? 6 %F 0 6 %F ! C>! : !%F <A %2( %'A ? 0 %F % %F : 0 %F : - %F : " = # - <=# ? :=& % =& # %F : ! F 7( : !%F : 6 2)9( 6 %F 0 : !%F 0 5 %F 19 = # - <=# ? 1# # 0 %F 1 : !%F 1 C# 1& 6 %F 1 %F 1 '( 0 %F %F . ! ! X = %F K # %F , 6 %F 9$'$2$ + #> . - 0 & ! ! %F # 4 ! >: : ! %F F Y >:J =& F - + & ! ! %F F Y + & ! ! %F = - >2(( ! = %F

'; -9$7$2$ > - - ! > '( >! . %2;;A E :)+ %2;BB >2; =# % 0& :>9:42# & % 0& >9#= 2 & % 0& 'B =# % 0& E) 6#1 : $%'((A E :)+ " - $%'((9 E :)+ " A( - $%'((9 E :)+ " A( A'% - $%'((9 E :)+ " A( 229 - $%'((9 E :)+ " A7 I" ! $%'(() > =&9C1E 6 %F 0 7 2) :123 . $%2;;; 0 7 Z A( - $%'((9 0 7 Z A7 I" ! $%'(() #1 '9- = % 0& - #1 'B- &$ ! 9$7$'$ ! D $ 9$7$'$2$ ,49 , 1 ! % 23 C <02C0' 07C09% ! 22$7$2? D - Y- 6 - <F $%'(('?$ 1! =& ! 'B <=# ? - 66 & 66$ ,496 8 1! = 1# 2β < J#>*7$A$'$)? >: >2; - ! = = $ 1! >: - #6 & 66 ! 0 $ ,496 . 1! > >: >2; ! > > $ 1! >: - 46 & 6 ! 0 $ ,490)9 ,

1! 0 ! #6 & 66 ! 4&2 4&'

7( ,490)98 1! G ! = >: >2; <=# ? ! = = $ 1! = ! < '9 2;9? 4& 0 - #6C=! 6 4& =$ ,490)9. 1! G ! > < 2;) 'B)? >: >2; <=# ? ! > C > $ 1! !

4& 0 - & 66C #6 4& >$ ,49.9

1! 0 ! = 6 46 ! 012 01'

% ! 0 % 0 01$ 1! >

< 2;) 'B)? 4& > - =! 6 #6

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

A(84 A782 #1'B A(4 C A782

A(84 A78' #1'B A(4 C A78'

A( E :)+ " A(

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77 ,;$ 989 "#' >,;$ 989 "#" >,;$ 989 "# 1! E: A( % A( A' A( 229 - ! A( A( ! E :)+ " A( % E :)+ " A( A' E :)+ " A( 229$ 1! >: -- 0 6 & 6 ! G 4& =$ ! 'B - & 66 #6 % ! E)1 - ! # :@ & 66 $ 1!

E)1 = A( % E)1 = A( A' E)1 = A( 229%

$ ,;$ 9.9 " 1 ! > A7 %! A7 E: E :)+ " A7 < I" ! $%'(()? ! A7 A7 $ 1! 4& > - ! 46 & 6 $ ! 'B - & 66 #6 % ! E)1 - ! # :@ & 66 $ ,;$ 9 " 9. 1! A7 E: E :)+ " A7 < I" ! $%'(()?

! A7& A70 % - & 66 0 6 !

& 66C0 6 >$ ! ! A7 E: > 01 - 46 & 66% A7 > 01$ ! ! 'B - - ! E)1 % A7 > 01 - 0 6 & 66 ! & 66 # :@ E)1$ ,;$ 98934 ?0 5 79 40@ 2 ! = $ 1! E: >: &> =& - ! 40@ 79 40@ 79 $ 1! 4& = - ! Y! 6 & 6 $ 1!

E)1 - & 6 =! 6% E)1 = 79<40@?$ ,;$ 98934 ?- 5>,;$ 98934"#>,;$ 98961%1 >,;$ 989; $!

79 @% A( 4> @%E )3 4@)B : 2 # @ !

= $ E: >: =&% &> =&

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79 @ C 79 @ % A( C A( % E: )3 CE: )3 : 2 C

: 2 % $ 1! 4& = - !

46 & 6 $ 1! ! E)1 - & 6

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79

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A$2$9$ &> =&

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! 9' A( \ C - =&$ 1! - ! <E ')(CE 'B(? - 2%B '$ " ) '8) A$'$2$ : " %Aµ ! % (%A 2µ =&% $ 1 '(µ% 'µ ! - %'µ 2( 0& < ?% ' ! G " <2µ? - 2 ' ! ! $ 1! &> =& 3(µ % 3µ ! 2( - %3 µ 2( 0&% 2( ! " <2%A µ ?

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! 97 <& ! ? Q $ 1 @ =& % @ ! " ! ! $ A$'$9$ ! ! =& 1 " % " =&% ! AQ =& ! ! $ =& ! ! !

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- $ 6 - %) \ =& - ! 'A( \ 'A( > >'$ 1!

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! A2 "" 7 + . A$A$2$ F - &> # 42( %- > @ &> '( < ! - ? ! A( B( =& ! ) " $ - %- 2 ! 7( P>$ 1! %2A( µ > C, ! 97 P> $ $ 1! % &> =& ! $ A$A$'$ > @

1 " &> % &> =& #

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! A7 G $ & A % G - -! ! - ! G * @ < C ?] < G ?C $ <(%9 ? 4 "/ / 7%3A - ! 7BB 4'E -'A >0 A( 2( # A '%A =#&0 < ? A C0 <(%)L < C? $%2%7L < C? 0 ? '9%3 = 4>E7 <3%AL? "$ 6 :, A$)$2$ # ! 1! - % - G 42( $ 2(( =& '2 < #7? :6 <0 ? - $ $ $ 73 P> <+ C> ?$ 1! %A( 2(( ! A( $ & A ) ! 73 P>%2( ! 2 ! $ . ! ! E )(( ! ! (%)% 2A $ 1! ! ! - 6 1F% (%A ! - '( P> $ $ 1! ! -9 P> 7A(( 7( <0 J0 > )(((?$ 1! ! 7A - % " G B( P>$ A$)$'$ -1! - ! - ! 73 P> <0 9A(J ? - 2 '( % - $ 1! - % =&% - - - 2 !

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