Testosterone differentially regulates targets of lipid and glucose metabolism in liver, muscle and adipose tissues of the testicular feminised mouse

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glucose metabolism in liver, muscle and adipose tissues of

the testicular feminised mouse

KELLY, Daniel <http://orcid.org/0000-0002-7463-0692>, AKHTAR, Samia,

SELLERS, Donna, MURALEEDHARAN, Vakkat, CHANNER, Kevin S. and

JONES, Thomas Hugh

Available from Sheffield Hallam University Research Archive (SHURA) at:

http://shura.shu.ac.uk/12966/

This document is the author deposited version. You are advised to consult the

publisher's version if you wish to cite from it.

Published version

KELLY, Daniel, AKHTAR, Samia, SELLERS, Donna, MURALEEDHARAN, Vakkat,

CHANNER, Kevin S. and JONES, Thomas Hugh (2016). Testosterone differentially

regulates targets of lipid and glucose metabolism in liver, muscle and adipose

tissues of the testicular feminised mouse. Endocrine, 54 (2), 504-515.

Copyright and re-use policy

See

http://shura.shu.ac.uk/information.html

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: M K O ;[ : ;_;a 9 : K =A 9 =K < L < S ;: M KJ < L K ;O OR < O ` A KL < 9KG < : K ? > æê ée – 9 O O ;G ;=9 L =A S < a L < 9 O < S ;: G R O a =< 5 c j

˜^ ú w ?w v | ™ 9 : S F H Y ˜^ ú w ?w v{ ™ M _ Y _G G ;a < U < L OR O – ;=S —KA ^ < G ;a < E – ;KJ : M < _ _< aK M _ K < OK M OK < L M : < 5 c k

KL < 9 KG < : K ? Z < ^ 9 K ;a > f ? å g – 9 O < =< U 9 K < S ;: Y _G G ;a < a M G ^ 9 L < S K M x š G ;a < ˜^ h w ?w w v ™ 9 : S K < OK M OK < L M : < 5 f l

i j k l mn o pq r k st l r u spk v n w x y z { y | { } ~ 

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KL < 9 KG < : K OJ M – < S 9 KL < : S K M L < S R a ;: [ KJ ;O <~ ^ L < O O ;M : ;: Y _G G ;a < ˜^ ú w ?w |  ™ ? H == M KJ < L [ < : < K 9L [ < K O 5 f 5

– < L < : M K 9 =K < L < S ` < K– < < : < ~ ^ < L ;G <: K 9 = [ L M R ^ O ;: KJ < K ;O OR < O ;: U < OK ;[ 9 K < S ? 5 f 7

u L M K < ;: < ~ ^ L < O O ;M : M _ u i @ ;: KJ < =;U < L M _ <~ ^ < L ;G <: K 9 = 9: ;G 9 =O G 9 K a J < S [ < : < < ~ ^ L < O O ;M : S 9 K 9 – ;KJ 5 f X

L < S R a < S =< U < =O ;: Y _G ^ =9 a < ` M G ;a < a M G ^ 9 L < S K M – ;=S —KA ^ < ˜^ ú w ?w w | ™ 9 : S : M < _ _< a K M _ KL < 9 KG < : K – ;KJ 5 f \

K < OK M OK < L M : < ˜i ;[ R L < q ™ ? > R O a =< ^ L M K < ;: <~ ^ L < O O ;M : M _ u i @ – 9 O L < S R a < S ;: Y _G G ;a < ˜^ ú w ?w v ² ™ – ;KJ 9 5 f ]

O ;[ : ;_;a 9 : K ;: a L < 9 O < ;: < ~ ^ L < O O ;M : _M ==M – ;: [ Y e Y ˜^ ú w ?w v ™ ? Z < ^ 9K ;a ‚ V @ ^ L M K < ;: – 9 O 9 =O M L < S R a < S ;: 5 f c

Y _G G ;a < L < a < ;U ;: [ ^ =9 a < ` M ˜^ ú w ?w w v ™ 9 O S < G M : OKL 9 K < S 9 K KJ < [ < : < =<U < =E J M – < U < L K < OK M OK < L M : < 5 f f

9 S G ;: ;OKL 9 K ;M : J 9 S : M < __ < a K OJ M – ;: [ S ;O a L < ^ 9 : a A ` < K– < < : [ < : < 9 : S ^ L M K < ;: < ~ ^ L < O O ;M : ? Z @ q ;: 5 f j

G R O a =< – 9 O 9 =O M L < S R a < S 9 K KJ < ^ L M K < ;: =< U < = ;: Y _G G ;a < ˜^ ú w ?w q { ™ E ` R K KJ < L < – 9 O : M < __ < a K S R < K M 5 f k

KL < 9 KG < : K ? > R O a =< ‚ ž b Y { – 9 O S < a L < 9 O < S ;: Y _G G ;a < a M G ^ 9 L < S K M – ;=S KA ^ < ˜^ ú w ?w r ± ™ 9 : S Y e Y 5 j l

S < G M : OKL 9 K < S 9 KL < : S K M – 9 L S O ;: a L < 9 O ;: [ KJ ;O < ~ ^ L < O O ;M : ˜^ ú w ?w | r ™ ? œ < – < L < R : 9 ` =< K M S < K < a K ‚  u 8 5 j 5

^ L M K < ;: < ~ ^ L < O O ;M : ;: KJ < =;U < L M _ < ~ ^ < L ;G < : K 9 = 9: ;G 9 =O ? 5 j 7

5 j X

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9™ V J M =< OK < L M = G < K 9 ` M =;OG 5 j ]

g ~ ^ L < O O ;M : M _ a J M =< OK <L M = KL 9 : O^ M L K < L O E ‹ ? [ ] 9: S ‹ è b ä Œ E – < L < L < S R a < S ;: KJ < =;U < L M _ Y _G G ;a < 5 j c

a M G ^ 9 L < S K M =;KK < LG 9 K < O ˜^ ú w ?w w s E ^ ú w ?w w q ™ ? Y L < 9 KG <: K – ;KJ K < OK M OK < L M : < O ;[ : ;_ ;a 9 : K =A ;: a L < 9 O < S KJ ;O 5 j f

<~ ^ L < O O ;M : ˜^ ú w ?w q ± E ^ ú w ?w q ™ E O ;G ;=9 L K M – ;=S —KA ^ < =<U < =O ˜Y 9 ` =< r ™ ? F ;G ;=9 L =A E ‹ ? [ ] – 9 O S < a L < 9 O < S ;: 5 j j

F H Y M _ Y _G G ;a < ˜^ ú w ?w v™ E 9 : < __ < a K KJ 9 K – 9 O 9 ` M =;OJ < S ` A K < OKM OK < L M : < 9 S G ;: ;OKL 9 K ;M : ˜^ ú w ?w v | U O 5 j k

Y _G u ™ ?  Z ] èd Œ 9 : S  Z ] èd = < ~ ^ L < O O ;M : – 9 O O ;[ : ;_;a 9 : K =A =M – < L ;: KJ < F H Y M _ Y _G G ;a < U < L OR O x š 5 k l

=;KK < LG 9 K < O ˜^ ú w ?w w q E ^ ú w ?w w r ™ ? Y L < 9 KG < : K – ;KJ K < OK M OK < L M : < < =< U 9 K < S KJ < O < < ~ ^ L < O O ;M : =< U < =O M _  Z ] èd Œ 5 k 5

˜^ ú w ?w v | ™ O ;G ;=9 L K M KJ M O < S < G M : OKL 9 K < S ;: – ;=S —KA ^ < G ;a < 9 =KJ M R [ J : M K O ;[ : ;_;a 9 : K =A O M _ M L  Z ] èd = 5 k 7

– ;KJ M : =A 9 KL < : S K M – 9L S O ;: a L < 9 O < S <~ ^ L < O O ;M : M ` O < LU < S ˜^ ú w ?w | r ™ ? ¯: a M : KL 9 OK K M [ < : < < ~ ^ L < O O ;M : 5 k X

S 9 K 9 E H d V H v ^ L M K < ;: – 9 O : M K O ;[ : ;_ ;a 9 : K =A 9 =K < L < S ` < K– < < : KL < 9 KG < : K [ L M R ^ O 9 =KJ M R [ J G < 9 : 5 k \

<~ ^ L < O O ;M : 9 ^ ^ < 9 L < S K M S < G M : OKL 9 K < 9 O ;G ;=9 L ^ 9 KK < L: ˜i ;[ R L < q ™ ? H u Í g ^ L M K < ;: <~ ^ L < O O ;M : G 9 K a J < S 5 k ]

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[ < : < < ~ ^ L < O O ;M : S 9 K 9 – ;KJ O ;[ : ;_ ;a 9 : K =A =M – <L =< U < =O ;: ^ =9 a < ` M —KL < 9 K < S Y _G G ;a < a M G ^ 9 L < S K M x š 5 k c

=;KK < LG 9 K < O 9 : S K < OK M OK < L M : < —KL < 9 K < S Y _G G ;a < ˜^ ú w ?w v v E ^ ú w ?w w ± L < O^ < a K ;U < =A ™ ? 5 k f

` ™ i 9KKA 9 a ;S G < K 9 ` M =;OG 5 k j

Q ;O a < L 9 = 9 S ;^ M O <  b å Œ < ~ ^ L < O O ;M : – 9 O O ;[ : ;_;a 9 : K =A J ;[ J < L ;: Y _G G ;a < L < a < ;U ;: [ ^ =9 a < ` M KJ 9 : ;: x š 5 k k

=;KK < LG 9 K < O 9 =O M L < a < ;U ;: [ ^ =9 a < ` M ;: Ì < aK ;M : O ˜^ ú w ?w r { E Y 9 ` =< r ™ ? Y < OK M OK < L M : < KL < 9 KG < : K M _ Y _G G ;a < 7 l l

L < KR L: < S < ~ ^ L < O O ;M : =< U < =O K M KJ M O < M _ x š G ;a < – ;KJ 9 O ;[ : ;_;a 9 : K L < S R a K;M : a M G ^ 9 L < S K M ^ =9 a < ` M — 7 l 5

KL < 9 K < S Y _G G ;a < ˜^ ú w ?w q ± ™ ? Y K < OK 9 : 9 =A O ;O O ;G ;=9 L =A L <U < 9 =< S 9 : ;: aL < 9 O < ;: J < ^ 9K ;a  b å Œ < ~ ^ L < O O ;M : 7 l 7

;: Y _G ^ =9 a < ` M G ;a < 9 =KJ M R [ J : M K OK 9 K ;OK ;a 9 ==A O ;[ : ;_;a 9 : K ˜^ ú w ?w ² ™ ? 8 < a L < 9 O < S Š ? æ < ~ ^ L < O O ;M : – 9 O 7 l X

M ` O < LU < S ;: F H Y _L M G Y _G G ;a < a M G ^ 9 L < S K M – ;=S KA ^ < ˜^ ú w ?w v  ™ 9 =KJ M R [ J K < OK M OK < L M : < 7 l \

9 S G ;: ;OKL 9 K ;M : K M Y _G 9 : ;G 9 =O J 9 S : M < __ < a K M : KJ ;O ? Z < ^ 9 K ;a [ < : < < ~ ^ L < O O ;M : M _ ˜ ä _ ™ 9 : S ‹ b ä b ä E KJ < 7 l

]

I < A L < [ R =9 K M L A < : Ÿ A G < O ;: S < : M U M =;^ M [ < : < O ;O E – < L < O ;[ : ;_ ;a 9 : K =A ;: a L < 9 O < S ;: Y _G G ;a < L < a < ;U ;: [ 7 l c

^ =9 a < ` M ;: Ì < a K ;M : O a M G ^ 9 L < S K M – ;=S —KA ^ < =;KK < LG 9 K < O ˜^ ú w ?w { s E ^ ú w ?w { q L < O^ < a K ;U < =A š™ ? Y < OK M OK < L M : < 7 l

f

KL < 9 KG < : K S < a L < 9 O < S KJ ;O < ~ ^ L < O O ;M : ` R K : M K O ;[ : ;_ ;a 9 : K =A ? ‚ < : < < ~ ^ L < O O ;M : M _ 9 == M KJ < L =;^ ;S 7 l j

G < K 9 ` M =;OG K 9 L [ < K O ;: =;U < L 9 : S 9 S ;^ M O < K ;O OR < – < L < : M K O ;[ : ;_;a 9 : K =A S ;__ < L < : K ` < K– < < : 9 : ;G 9 = [ L M R ^ O ? 7 l k

œ < OK < L : ` =M KK ;: [ OJ M – < S J < ^ 9 K ;a ^ L M K < ;: < ~ ^ L < O O ;M : M _ i H F h 9 : S H V H V H K M ` < ;: a L < 9 O < S ;: Y _G 7 5 l

G ;a < a M : _ ;L G ;: [ [ < : < < ~ ^ L < O O ;M : _;: S ;: [ O š ? Y < OK M OK < L M : < KL < 9 KG < : K O ;[ : ;_ ;a 9 : K =A L < S R a < S KJ < ^ L M K < ;: 7 5 5

<~ ^ L < O O ;M : M _ KJ < O < < : Ÿ A G < O U < L OR O ^ =9 a < ` M KL < 9 K < S Y _G G ;a < K M O ;G ;=9 L =< U < =O 9 O x š =;KK <L G 9 K < O ? 7 5 7

h M K 9L [ < K O M _ _9 K G < K 9 ` M =;OG 9 : S a J M =< OK < L M = J M G < M OK 9 O ;O S ;O^ =9 A < S 9 =K < L < S [ < : < < ~ ^ L < O O ;M : ;: G R O a =< 7 5 X

K ;O OR < _L M G KJ < S ;__ <L < : K <~ ^ < L ;G < : K 9 = [ L M R ^ O ? 7 5 \

7 5 ]

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‚ < : < <~ ^ L < O O ;M : M _ Š g Z – 9 O O ;[ : ;_;a 9 : K =A L < S R a < S ;: Y _G ^ =9 a < ` M G ;a < ;: 9 == K ;O OR < O M KJ < L KJ 9 : 7 5 f

U ;O a < L 9 = 9 S ;^ M O < ˜G R O a =< ^ ú w ?w r q E =;U < L ^ h w ?w w v E F H Y ^ ú w ?w w r ™ E 9 : S K < OK M OK < L M : < 9 S G ;: ;OKL 9 K ;M : 7 5 j

;: a L < 9 O < S < ~ ^ L < O O ;M : O ;[ : ;_;a 9 : K =A 9 : S ` 9 a I K M – ;=S —KA ^ < =< U < =O ;: KJ < O < K ;O OR < O ˜G R O a =< ^ ú w ?w w ² E =;U < L 7 5 k

 j k l mn o pq r k st l r u spk v n w x y ž 

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^ ú w ?w q { E F H Y ^ ú w ?w r ™ ? c ? ä Z ä 9 : S c ? ä Zœ – <L < O ;[ : ;_;a 9 : K =A L < S R a < S ;: F H Y M _ Y _G G ;a < L < a < ;U ;: [ 7 7 l

^ =9 a < ` M U < L OR O x š =;KK < L G 9 K < a M : KL M =O ˜^ ú w ?w v E ^ ú w ?w q L < O^ < a K ;U < =A ™ ? u ^ 9L [ – 9 O 9 =O M L < S R a < S ;: 7 7 5

U ;O a < L 9 = 9 S ;^ M O < K ;O OR < M _ Y _G G ;a < ˜^ ú w ?w w v ™ ? Y < OK M OK < L M : < KL < 9 KG < : K J 9 S : M < _ _< a K M : KJ < 9 =K < L < S 7 7 7

<~ ^ L < O O ;M : M _ c ? ä Z_ – J < : a M G ^ 9 L < S K M ^ =9 a < ` M KL < 9 K < S Y _G G ;a < ˜O < < Y 9 ` =< r ™ ? 7 7 X

ž x e ^ L M K < ;: <~ ^ L < O O ;M : ;: =;U < L 9 : S G R O a =< S < G M : OKL 9K < S KJ < O 9 G < ^ 9 KK < L : ;: S ;a 9K < S ` A [ < : < 7 7 \

<~ ^ L < O O ;M : 9 : 9 =A O ;O – ;KJ 9 L < S R a K ;M : ;: Y _G ^ =9 a < ` M G ;a < a M G ^ 9 L < S K M – ;=S —KA ^ < =;KK <L G 9 K < O ˜^ ú w ?w w v E 7 7 ]

^ ú w ?w v ™ ? Y L < 9KG < : K – ;KJ K < OK M OK < L M : < < =< U 9 K < S ž x e =<U < =O O ;[ : ;_;a 9 : K =A ;: =;U < L 9 : S G R O a =< ˜^ ú w ?w q { E 7 7 c

^ ú w ?w q q ™ E K M O ;G ;=9 L =< U < =O O < < : ;: ^ =9 a < ` M —KL < 9 K < S Y _G G ;a < ˜i ;[ R L < q ™ ? 7 7 f

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g ~ ^ =M L 9K M L A < U ;S < : a < _L M G KJ ;O OKR S A OR [ [ < OK O KJ 9 K K < OK M OK < L M : < J 9 O K ;O O R < —O^ < a ;_;a G < K 9 ` M =;a < __ < a K O ;: 7 7 k

KJ < L < [ R =9K ;M : M _ [ < : < K 9L [ < K O – J ;a J a M : KL M = [ =R a M O < R K ;=;O 9 K;M : ;: =;U < L E F H Y 9 : S OI < =< K 9 = G R O a =< E 9 : S 7 X l

=;^ ;S G <K 9 ` M =;OG ;: =;U < L 9 : S F H Y ? F M G < M _ KJ < O < < _ _ < a K O 9 L < E 9 K =< 9 OK ;: ^ 9 L KE 9 : S L M [ < : L < a < ^ K M L — 7 X 5

;: S < ^ < : S < : K 9 : S G 9 A ^ M K < : K ;9 ==A < ~ ^ =9 ;: O M G < M _ KJ < M ` O < L U < S a =;: ;a 9 = ` < : < _;K M _ K < OK M OK < L M : < ;: G < : 7 X 7

– ;KJ Y q 8 9 : S > < K F ? 7 X X

7 X \

ª ]_ é[ _ é] Z [ ™ ] « dd ] b é_ [ ™ « g ? Z ] _ _ ç[ ™ [d ª ä Zœ ] é_ [d > æê b [ _ ] ^ ] éä è [ æç_ ë 7 X ]

‚ ž b Y { < ~ ^ L < O O ;M : ;O I : M – : K M a M LL < =9 K < ^ M O ;K ;U < =A – ;KJ ;: OR =;: L < O^ M : O ;U < : < O O 9 : S S < _< a K O ;: 7 X c

<~ ^ L < O O ;M : M _ ‚ ž b Y { J 9 U < ` < < : M ` O < LU < S ;: ^ 9 K ;< : K O – ;KJ Y q 8 ­q { ®? œ < J 9 U < OJ M – : KJ 9 K KJ <L < ;O 7 X f

S < a L < 9 O < S < ~ ^ L < O O ;M : M _ ‚ ž b Y { ;: G R O a =< 9 : S F H Y ;: KJ < K < OK M OK < L M : < S < _ ;a ;< : K Y _G G M R O < ? 7 X j

Y < OK M OK < L M : < J 9 O ^ L < U ;M R O =A ` < < : OJ M – : K M ;: a L < 9 O < KJ < < ~ ^ L < O O ;M : M _ ‚ ž b Y { ;: a R =KR L < S OI < =< K 9 = 7 X k

G R O a =< a < ==O E J < ^ 9 K M a A K < O 9 : S 9 S ;^ M aA K< O ­q | —q ± ® 9 O – < == 9 O 9 R [ G < : K ;: [ G < G ` L 9 : < KL 9 : O =M a 9 K ;M : 9 : S 7 \ l

^ L M G M K ;: [ [ =R a M O < R ^ K 9 I < ;: 9 S ;^ M O < 9 : S OI < =< K 9 = G R O a =< K ;O OR < ­q ± ®? @ < A < : Ÿ A G < O ;: U M =U < S ;: 7 \ 5

[ =A a M =A O ;O E u i @ 9 : S Z @ E – < L < O ;[ : ;_ ;a 9 : K =A L < S R a < S ;: G R O a =< E =;U < L 9 : S F H Y M _ Y _G G ;a < ? Y J ;O 7 \ 7

OR ^ ^ M LK O ^ L < U ;M R O OKR S ;< O – J ;a J J 9 U < S < G M : OKL 9 K < S 9 : ;: aL < 9 O < ;: KJ < 9 a K ;U ;KA M _ u i @ 9 : S Z @ ;: 7 \ X

aR =KR L < S L 9 K OI < =< K 9 = G R O a =< a < ==O 9 : S ;: a L < 9 O < S J < ~ M I ;: 9 O < 9 a K;U ;KA ;: G R O a =< K ;O OR < M _ a 9 OKL 9 K < S L 9 K O 7 \ \

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(14)

_M ==M – ;: [ K < OK M OK < L M : < KL < 9 KG < : K KJ R O S ;G ;: ;OJ ;: [ KJ < L 9 ;O < S ` =M M S =< U < =O M _ [ =R a M O < O < < : ;: R : KL < 9 K < S 7 \ ]

a M : KL M = L 9 K O ­q ± —q s ® ? ¯G ^ L M U < S [ =R a M O < R K ;=;O 9 K ;M : ;: G R O a =< E =;U < L 9 : S F H Y ` A K < OK M OK < L M : < G 9 A 7 \ c

L < S R a < KJ < a M : U < L O ;M : M _ [ =R a M O < K M _9 K ;: K ;G < O M _ < ~ a < O O 9 : S ;G ^ L M U < ;: OR =;: O <: O ;K ;U ;KA KJ R O 7 \ f

L < S R a ;: [ =;^ ;S 9 a a R G R =9K ;M : ;: KJ < O < 9 : S M KJ < L K ;O OR < O ? Y J ;O a =;: ;a 9 ==A – M R =S ` < U < LA ;G ^ M L K 9 : K ;: 7 \ j

G R O a =< 9 O KJ ;O K ;O OR < 9 a a M R : K O _M L 9 ^ ^ L M ~ ;G 9 K < =A ± | Î M _ – J M =< —` M S A ;: OR =;: —OK ;G R =9 K < S [ =R a M O < 7 \ k

R ^ K 9 I < ­r w E r v ® ? 7 ] l

œ < J 9 U < 9 =O M S < G M : OKL 9 K < S ;: KJ ;O O KR S A KJ 9 K KJ < G e h H < ~ ^ L < O O ;M : M _ ‚ =R a M O < — —^ J M O^ J 9 K < 7 ] 5

S < J A S L M [ < : 9 O < ˜ > f? å ™ E KJ < [ 9 K < – 9 A < : Ÿ A G < ;: KJ < ^ < : K M O < ^ J M O ^ J 9K < OJ R : K ^ 9 KJ – 9 A E ;O < =< U 9 K < S ;: 7 ] 7

KJ < =;U < L M _ Y _G G ;a < OR [ [ < OK ;: [ KJ 9 K [ =R a M O < G 9 A 9 =O M ` < R K ;=;O < S S M – : KJ ;O L M R K < S R L ;: [ K < OK M OK < L M : < 7 ] X

S < _ ;a ;< : a A ? h H 8 u Z ;O ^ L M S R a < S ` A ‚  u 8 ;: KJ < ^ < : K M O < ^ J M O^ J 9 K < ^ 9 KJ – 9 A OR ^ ^ =A ;: [ L < S R a ;: [ 7 ] \

^ M – < L K M a M : KL ;` R K < K M _9 KKA 9 a ;S OA : KJ < O ;O ­ r q ® ? H : 9 ` < LL 9 : K ;: a L < 9 O < M _ ‚  u 8 < ~ ^ L < O O ;M : ;O ^ L < O < : K ;: 7 ] ]

M ` < O < 9 : S S ;9 ` < K ;a OR ` Ì < a K O E 9: S M U < L < ~ ^ L < O O ;M : M _ ‚  u 8 9 =K < L O =;^ ;S G < K 9 ` M =;OG E ;G ^ 9 ;L O ;: OR =;: 7 ] c

O ;[ : 9 ==;: [ 9 : S OR ^ ^ L < O O < O ;: OR =;: —S < ^ < : S < : K [ =R a M O < R ^ K 9 I < ;: G M R O < 9 S ;^ M a A K < O ­r q ®? Z M – < U < L E KJ < 7 ] f

<~ 9 aK L M =< M _ J < ^ 9 K ;a ‚  u 8 ;: G < K 9 ` M =;a _R : a K ;M : ;O R : I : M – : ?

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¯: KJ < ^ L < O < : K OKR S A – < S < G M : OKL 9 K < KJ 9 K K < OK M OK < L M : < S < _;a ;< : a A : < [ 9 K ;U < =A 9 =K < L O KJ < < ~ ^ L < O O ;M : M _ 7 c 5

K 9 L [ < K O M _ =;^ ;S G < K 9 ` M =;OG ^ L ;G 9 L ;=A ;: =;U < L 9 : S F H Y ` R K J 9 S =;KK =< < __ < a K ;: Q H Y ? 8 < a L < 9 O < S Š ? æ ;: 7 c 7

Y _G G ;a < – ;KJ =M – K < OK M OK <L M : < G 9 A =;G ;K KJ < J A S L M =A O ;O M _ =;^ M ^ L M K < ;: O 9 : S KJ < OR ` O <  R <: K R ^ K 9 I < M _ 7 c X

i i H ;: K M F H Y ? H ^ L < U ;M R O OKR S A E J M – < U < L E J 9 O OJ M – : KJ < <~ ^ L < O O ;M : M _ J M L G M : < O < : O ;K ;U < =;^ 9 O < 7 c \

˜Z F ž ™ 9 : S ž u ž K M ` < < =< U 9 K < S ;: F H Y M _ G 9 =< G ;a < – ;KJ 9 O < =< a K ;U < 9 S ;^ M a A K < H e I : M a I S M – : 7 c ]

˜_H e @ Í ™ ­r r ®? Y J < O < G ;a < – < L < _ < S 9 : M L G 9 = a J M – S ;< K 9 : S KJ <L < _M L < ž u ž ;: aL < 9 O < ;: KJ < 9 ` O < : a < M _ 7 c c

K < OK M OK < L M : < 9 a K ;U 9 K < S H e O ;[ : 9 ==;: [ G 9 A L < _ =< a K < =< U 9 K < S OR ` a R K 9 : < M R O =;^ ;S OK M L 9 [ < 9 : S S < aL < 9 O < S 7 c f

KL ;[ =A a < L ;S < R O 9 [ < 9 O 9 : < : < L [ A O M R L a < ;: M KJ < L K ;O OR < O ;: K ;G < O M _ =M – _9 K ;: K 9 I < ? Y L < 9KG < : K M _ 7 c j

J A ^ M [ M : 9 S 9 = G < : – ;KJ Y e Y _ M L : ;: < G M : KJ O L < OR =K < S ;: 9 G 9 L I < S S < a L < 9 O < ;: ` M KJ ž u ž 9 a K ;U ;KA 9 : S 7 c k

KL ;[ =A a < L ;S < R ^ K 9 I < ;: 9 ` S M G ;: 9 = 9 S ;^ M O < K ;O OR < ­r { ®? i M ==M – ;: [ _R LKJ < L ;: U < OK ;[ 9K ;M : E 9 =KJ M R [ J ž u ž 7 f l

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<~ ^ L < O O ;M : M L 9 a K ;U ;KA – 9 O : M K L < ^ M L K < S E KJ < ;: J ;` ;K ;M : M _ =;^ ;S R ^ K 9I < 9 _K < L K < OK M OK < L M : < 9 S G ;: ;OKL 9 K ;M : 7 f 5

– 9 O 9 ^ ^ 9 L < : K ;: U ;O a < L 9 = ˜M G < : K 9 = ^ =R O G < O < : K <L ;a ™ 9 : S L < KL M ^ < L ;K M : < 9 = ` R K ;: a L < 9 O < S ;: 9 ` S M G ;: 9 = 7 f 7

F H Y OR [ [ < OK ;: [ KJ 9 K ;: J ;` ;K ;M : M _ KL ;[ =A a < L ;S < 9 O O ;G ;=9 K ;M : G 9 A S ;L < a K =;^ ;S K M OR ` a R K 9 : < M R O _ 9K ;: 7 f X

Y e Y —KL < 9K < S G < : 9 : S G 9 A KJ < L < _ M L < ;: U M =U < 9 =K < L < S =;^ 9 O < 9 a K ;U ;KA M L <~ ^ L < O O ;M : ;: O^ < a ;_;a K ;O OR < O 7 f \

­r | ®E 9 O OR [ [ < OK < S ;: KJ < ^ L < O < : K OKR S A ? 7 f ]

Z R G 9 : F V 8 v ;O 9 a L ;K ;a 9 = a M : KL M = ^ M ;: K M _ =;^ ;S ^ 9 L K ;K ;M : ;: [ – ;KJ J ;[ J F V 8 9 a K;U ;KA _9 U M R L ;: [ _ 9 K 7 f c

OK M L 9 [ < 9 : S OR ^ ^ L < O O ;M : M _ KJ < < : Ÿ A G < 9 a K ;U 9 K ;: [ G < K 9` M =;a ^ 9 KJ – 9 A O KJ 9 K ^ L M G M K < KJ < ` R L: ;: [ M _ _ 9K 7 f f

9: S S < a L < 9 O < =;^ ;S OA : KJ < O ;O ­ r  ® ? > ;a < – ;KJ 9 K 9L [ < K < S S ;OLR ^ K ;M : M _ KJ <  b å Œ [ < : < J 9 U < U < L A =M – 7 f j

=< U < =O M _ Q ž 8 ž 9 : S ;G ^ 9 ;L < S KL ;[ =A a < L ;S < 9 : S a J M =< OK < L M = < O K< L ` ;M OA : KJ < O ;O 9 O – < == 9 O G 9 L I < S =A 7 f k

L < S R a < S 9 S ;^ M O ;KA 9 : S S < a L < 9 O < S J < ^ 9 K ;a OK < 9K M O ;O M : ` M KJ =< 9 : 9 : S M ` ü M ` ` 9 a I [ L M R : S S < O^ ;K < J ;[ J < L 7 j l

_M M S ;: K 9 I < ­ r ± E r ² ® ? ¯ : KJ < ^ L < O < : K OKR S A – < S < G M : OKL 9 K < O ;[ : ;_ ;a 9 : K =A ;: a L < 9 O < S  b å Œ < ~ ^ L < O O ;M : ;: 7 j 5

Q H Y M _ Y _G G ;a < 9 : S 9 KL <: S K M – 9 L S O ;: aL < 9 O < S < ~ ^ L < O O ;M : ;: KJ < =;U < L ? d <A M : S ;K O L M =< ;: _ 9 KKA 9 a ;S 7 j 7

` ;M OA : KJ < O ;O E F V 8 v ;O 9 : ;G ^ M LK 9 : K _9 aK M L ;: KJ < ^ 9 KJ M [ < : < O ;O M _ =;^ ;S —;: S R a < S ;: OR =;: L < O ;OK 9 : a < – ;KJ 7 j X

F V 8 v S < _ ;a ;< : a A R ^ —L < [ R =9K ;: [ ;: OR =;: —O ;[ : 9 ==;: [ a M G ^ M : < : K O 9 : S [ =A a M [ < : G < K 9 ` M =;OG ;: ;: OR =;: — 7 j \

O < : O ;K ;U < K ;O OR < O ­r ² ® ? Y J ;O OR [ [ < OK O KJ 9K K < OK M OK < L M : < J 9 O KJ < ^ M K < : K ;9 = K M ;G ^ L M U < ` M KJ =;^ ;S 9 : S 7 j ]

[ =R a M O < G < K 9 ` M =;OG U ;9 L < S R a ;: [ F a S v < ~ ^ L < O O ;M : ;: Q H Y 9 : S KJ < =;U < L M _ Y _G G ;a < ? 7 j c

ž M – < L OR ` aR K 9 : < M R O ‹ ? [ ] <~ ^ L < O O ;M : ;: K < OK M O K< L M : < S < _;a ;< : K Y _G G ;a < G 9 A ` < ;: S ;a 9 K ;U < M _ 7 j f

S < a L < 9 O < S L < U <L O < a J M =< OK < L M = KL 9 : O^ M LK S < =;U < L A M _ =;^ M ^ L M K < ;: O 9 : S a J M =< OK < L M = _L M G F H Y K M KJ < =;U < L 7 j j

_M L a =< 9 L 9 : a < ? Y J ;O S ;__ <L <: a < – 9 O : M K 9 ^ ^ 9L < : K ;: Q H Y OR ^ ^ M LK ;: [ 9 : ;G ^ M L K 9 : K S <^ M K —O^ < a ;_ ;a L M =< M _ 7 j k

H u Í g ;: 9 S ;^ M O < K ;O OR < OR ` OKL 9 K < _=R ~ 9 : S 9 a a R G R =9 K ;M : M _ KL ;[ =A a < L ;S < ;: KJ < O < S <^ M K O ­ r s ® ? 7 k l

H S S ;K ;M : 9 ==A E ;: KJ < ^ L < O < : K OKR S A – < S < G M : OKL 9 K< KJ 9 K G e h H < ~ ^ L < O O ;M : M _  Z ] èd Œ 9 : S  Z ] èd = E I < A 7 k 5

KL 9 : O a L ;^ K ;M : _9 a K M L O 9 : S G 9 OK < L L < [ R =9 K M L O M _ =;^ M [ < : < O ;O ­{ w ® E – < L < O ;[ : ;_ ;a 9 : K =A S < a L < 9 O < S ;: F H Y M _ 7 k 7

Y _G G ;a < a M G ^ 9 L < S K M K < OK M O K <L M : < KL < 9 K < S 9 : ;G 9 =O 9 : S – ;=S —KA ^ < a M : KL M =O ? F ;G ;=9 L =A E M L a J ;S < a K M G A 7 k X

O ;[ : ;_;a 9 : K =A L < S R a < S J < ^ 9 K ;a F e g d u — v < ~ ^ L < O O ;M : ;: G ;a < _ < S 9 J ;[ J _ 9K S ;<K M L : M L G 9 = a J M – E 9 : 7 k \

< __ < a K KJ 9 K – 9 O 9 G < =;M L 9 K < S ` A K < OK M OK <L M : < KL < 9 KG < : K ;: J ;[ J _9 K S ;< K a M : S ;K ;M : O ­{ v ® ? H O F e g d u O 9L < 7 k

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I : M – : K M S ;L < a K =A ;: S R a < KL 9 : O a L ;^ K ;M : M _ G 9 : A [ < : < O : < < S < S _M L R ^ K 9 I < 9 : S OA : KJ < O ;O M _ a J M =< OK < L M =E 7 k c

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(16)

_9 KKA 9 a ;S O E KL ;[ =A a < L ;S < O E 9 : S ^ J M O^ J M =;^ ;S O ­{ q ® E K 9 I < : K M [ < KJ < L E KJ < O < S 9 K 9 =< 9 S R O K M J A ^ M KJ < O ;O < KJ 9 K 7 k f

K < OK M OK < L M : < S < _;a ;< : a A G 9 A S ;G ;: ;OJ F H Y G < K 9 ` M =;a _R : a K ;M : 9 : S L < S R a < =;^ ;S OKM L 9 [ < a 9^ 9 a ;KA ? 7 k j

¯: a L < 9 O < S =;U < L _ 9 K ;: Y _G G ;a < _L M G KJ < ^ L < O < : K OKR S A ;O a M : O ;S < L < S ^ 9L K =A S R < K M ;: a L < 9 O < S S < : M U M 7 k k

=;^ M [ <: < O ;O 9 : S KJ < < ~ ^ L < O O ;M : M _ i H F h 9 : S H V H V H ­ v ± ® E – J ;a J OR ^ ^ M LK < S < 9 L =;< L OKR S ;< O ;: S ;a 9 K ;: [ X l l

KJ 9 K 9 =9 a I M _ K < OK M OK < L M : < 9 a K ;M : L < OR =K O ;: J < ^ 9 K ;a =;^ ;S 9 a a R G R =9 K ;M : ­{ v —{ r ®? Y J < ^ L < O < : K OKR S A X l 5

9 S S ;K ;M : 9 ==A ;: S ;a 9 K < O KJ 9 K H d V H v 9 : S H u Í g E ;: U M =U < S ;: a J M =< OK < L M = 9 : S =;^ M ^ L M K < ;: < _ _=R ~ E 9 L < X l 7

L < S R a < S ;: KJ < K < OK M OK < L M : < S < _ ;a ;< : K OK 9 K < ;: KJ < =;U < L M _ Y _G G ;a < ? Y J < M U <L < ~ ^ L < O O ;M : M _ J < ^ 9 K ;a X l X

‹ è b ä Œ ;: KL 9 : O [ < : ;a G ;a < L < O R =K O ;: 9 G 9 L I < S ;: a L < 9 O < ;: Z 8 ž L < =< 9 O < E S < aL < 9 O < S ž 8 ž E 9 : S X l \

O ;[ : ;_;a 9 : K =A L < S R a < S 9 KJ <L M O a =< L M O ;O – J < : a M G ^ 9 L < S – ;KJ a M : KL M = G ;a < ­{ { ® ? i R LKJ < L G M L < E ;: a L < 9 O < S X l ]

J < ^ 9 K ;a a J M =< OK < L M = a M : K < : K – 9 O L <^ M LK < S ;: KJ < O < G ;a < 9 O KJ < =< U < = M _ < ~ ^ L < O O ;M : M _ KJ < H d V H v X l c

KL 9 : O^ M LK < L S < a L < 9 O < S ­{ | ® ? ¯ : S < < S E Y _G G ;a < _L M G KJ < ^ L < O < : K OKR S A J 9 U < < =< U 9 K < S K M K9 = a J M =< OK < L M = X l f

9: S ž 8 ž a M G ^ 9 L < S K M – ;=S —KA ^ < G ;a < ­ v ² ® ? Y J < L < _ M L < E KJ < ;: a L < 9 O < S J < ^ 9K ;a =;^ ;S 9 a a R G R =9 K;M : ;: M R L X l j

Y _G G ;a < G 9 A 9 S S ;K ;M : 9 ==A L < OR =K _L M G 9 ` O < : a < M _ ` < : < _;a ;9 = K < OKM OK < L M : < < __ < a K O M : =;^ ;S KL 9 : O^ M LK ? X l k

ª ]_ é[ _ é] Z [ ™ ] « dd ] b é_ [ ™ ^ ä _ é] Z › ] œ ê æä é[ Z_ [d Š ç? çå ä ™ å > æê b [ _ ] ^ ] éä è [ æç_ ë X 5 l

Y < OK M OK < L M : < 9 =K < L < S KJ < < ~ ^ L < O O ;M : M _ G 9 OK < L G <K 9` M =;a L < [ R =9 K M L O 9 O 9 ^ M K< : K ;9 = O ;[ : 9 ==;: [ G M S < M _ X 5 5

9 a K ;M : KM ;: _=R < : a < =;^ ;S 9 : S [ =R a M O < L < [ R =9 K ;M : ? e < S R a < S < ~ ^ L < O O ;M : M _ KJ < : R a =< 9L L < a < ^ K M L E =;U < L x X 5 7

L < a < ^ K M L ˜ž x e ™ E ;: G R O a =< E =;U < L 9 : S F H Y M _ Y _G G ;a < a M G ^ 9 L < S K M K < OK M OK < L M : < L < ^ =< K < 9 : ;G 9 =O X 5 X

– J < KJ < L – ;KJ M L – ;KJ M R K H e _R : aK ;M : =< 9 S O K M KJ < J A ^ M KJ < O ;O KJ 9K K < OK M OK <L M : < G 9 A ;: a L < 9 O < ž x e X 5 \

O ;[ : 9 ==;: [ K M < ~ < L K O M G < M _ ;K O ^ L M K < a K ;U < G < K 9 ` M =;a < __< a K O M : ? ž x e O 9 L < I <A KL 9 : O a L ;^ K ;M : 9 = L < [ R =9 K M L O X 5 ]

M _ =;^ ;S 9 : S a 9 L` M J A S L 9K < G < K 9 ` M =;OG I : M – : K M a M : KL M = G M =< a R =9 L ^ 9 KJ – 9 A O ;: a =R S ;: [ a J M =< OK < L M = X 5 c

< __ =R ~ E [ =R a M O < L < [ R =9K ;M : E _9 KKA 9 a ;S OA : KJ < O ;O 9: S ;: _ =9 G G 9 K ;M : ­{  ® ? ¯: ^ 9 L 9 ==< = – ;KJ K < OK M OK < L M : < X 5 f

9 O O M a ;9 K < S aJ 9 : [ < O ;: ž x e < ~ ^ L < O O ;M : ;: KJ < ^ L < O < : K OKR S A E – < O 9 – 9 =K < L 9K ;M : O ;: I : M – : ž x e K 9L [ < K X 5 j

[ < : < O } ˜ ä _ ™ E ‹ ? [ ] E ‹ è b ä Œ E Š ? æE  Z ] è? d Œ ? e 9KJ < L KJ 9 : ;: S R a ;: [ J < ^ 9 K ;a OK < 9 K M O ;O 9 O – ;KJ G 9 : A ž x e X 5 k

9 [ M : ;OK O E K < OK M OK < L M : < 9 S S ;K ;M : 9 ==A ^ L M K < a K O 9 [ 9 ;: OK S ;<K —;: S R a < S J <^ 9 K ;a =;^ ;S 9 a a R G R =9 K ;M : ;: KJ ;O X 7 l

G M S < = ­ v ± ® ? Y _G G ;a < 9 =O M J 9 S L < S R a < S F H Y 9 : S Q H Y < ~ ^ L < O O ;M : M _ c ? ä Zœ G e h H ;: S ;a 9 K ;: [ 9 X 7 5

^ M K < : K ;9 = G < a J 9 : ;OG ` A – J ;a J K < OK M OK < L M : < S < _ ;a ;< : a A G 9 A =< 9 S K M G < K 9 ` M =;a S A OL < [ R =9 K ;M : 9 : S X 7 7

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(17)

9 S U < L O < _9 K S ;OKL ;` R K ;M : ? H S S ;K ;M : 9 ==A E Y _G G ;a < S ;O^ =9 A < S =M – < L F H Y c ? ä Z ä Ô9 G 9 OK < L L < [ R =9 K M L M _ X 7 X

_9 KKA 9 a ;S M ~ ;S 9 K ;M : ™ < ~ ^ L < O O ;M : OR [ [ < OK ;: [ KJ 9 K K < O K M OK <L M : < S < _;a ;< : aA G 9 A _R L KJ < L ;: J ;` ;K =;^ ;S X 7 \

L < [ R =9K ;M : ? X 7 ]

Y J < ^ L < O < : K OKR S A ;: S ;a 9 K < O KJ 9 K K < OK M OK <L M : < G 9 A O ;[ : 9 =E 9 K =< 9 OK ;: ^ 9 LK E ` < A M : S ;K O a =9 O O ;a 9 = : R a =< 9 L X 7 c

9: S L M [ < : L < a < ^ K M L ˜H e ™ K M G M S R =9 K < K 9 L [ < K O M _ =;^ ;S 9 : S [ =R a M O < G < K 9 ` M =;OG 9 : S KJ 9 K KJ < O < 9 aK ;M : O 9 L < X 7 f

_R LKJ < L S ;_ _< L < : K ;9 ==A S < ^ < : S < : K M : KJ < K 9 L [ < K K ;O OR < ? œ J <KJ < L KJ < H e — ;: S < ^ < : S < : K < __ < a K O ;: KJ ;O OKR S A X 7 j

9L < U ;9 a M : U < L O ;M : K M < OKL 9 S ;M = 9 : S OR ` O <  R < : K 9 a K ;U 9 K ;M : M _ KJ < M < OKL M [ < : L < a < ^ K M L ˜g e ™ – 9 O : M K X 7 k

9 S S L < O O < S ? œ < J 9 U < ^ L < U ;M R O =A OJ M – : E J M – < U < L E KJ 9 K K < OK M OK < L M : < J 9 O 9 S S ;K ;M : 9 = 9 a K ;M : O M : J < ^ 9 K ;a X X l

9: S 9 M LK ;a =;^ ;S 9 a a R G R =9 K ;M : ;: Y _G G ;a < < U < : – ;KJ 9 L M G 9K 9 O < ;: J ;` ;K ;M : 9 : S g e ` =M a I 9 S < ­ v  E v ± ®? X X 5

i R LKJ < L ;: U < OK ;[ 9 K ;M : ;O L <  R ;L < S K M < =R a ;S 9 K < KJ < H e —;: S < ^ < : S < : K O ;[ : 9 ==;: [ G < a J 9 : ;OG O M _ X X 7

K < OK M OK < L M : < 9 a K ;M : ? X X X

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Y J < ^ L < O < : K < ~ ^ =M L 9 K M LA OKR S A ;O =;G ;K < S K M K 9L [ < K < ~ ^ L < O O ;M : S 9 K 9 9 : S – J ;=< ;K ;: S ;a 9 K < O ^ M K < : K ;9 = X X c

G < K 9 ` M =;a < __ < a K O M _ K < OK M OK< L M : < E ;K S M < O : M K S ;L < a K =A 9 O O < O O G < K 9 ` M =;a _R : aK ;M : ? ž 9 a I M _ K ;O OR < X X f

^ L < U < : K < S ^ L M K < ;: 9: 9 =A O ;O M _ F H Y 9 : S Q H Y S R < K M KJ < L < S R a < S 9 G M R : K O M _ ^ L M K < ;: L < a M U < L 9 ` =< _L M G X X j

9U 9 ;=9 ` =< 9 S ;^ M O < K ;O OR < ? ¯: 9 S S ;K ;M : E KJ < Y _G G M R O < ;O 9 G M S < = M _ [ =M ` 9 = H e S A O _R : a K ;M : 9: S O < U < L < =A X X k

L < S R a < S K < OK M OK < L M : < =< U < =O _L M G ` ;LKJ E KJ < L < _M L < – < a 9 : : M K L R =< M R K 9 : A S < U < =M ^ G < : K 9 = < _ _< aK O M _ KJ < O < X \ l

_9 a K M L O M : K ;O OR < O – J ;a J G 9 A ;: _=R < : a < KJ < ^ 9 KJ M [ < : < O ;O M _ G < K 9 ` M =;a S ;O M L S < L O ? œ J ;=OK KJ < X \ 5

K < OK M OK < L M : < ;: Ì < a K ;M : O ^ L M S R a < =< U < =O – ;KJ ;: KJ < : M L G 9 = L 9 : [ < E S ;R L : 9 = ^ 9 KK <L : O 9 L < 9 ` O < : K 9 : S X \ 7

OR ^ L 9 ^ J A O ;M =M [ ;a =< U < =O ;: KJ < _ ;L OK _< – S 9 A O 9 L < 9 ^ ^ 9 L < : K – ;KJ : < 9 L —;: _L 9 ^ J A O ;M =M [ ;a =< U < =O K M – 9 L S O X \ X

KJ < < : S M _ KJ < ;: K < L U 9 = ­ v  ® ? F R a J 9 S G ;: ;OKL 9 K;M : G 9 A < ~ ^ =9 ;: KJ < ;: _=R < : a < M _ K < OK M OK < L M : < KL < 9KG < : K X \ \

M : [ < : < <~ ^ L < O O ;M : 9 ` M U < 9: S ` < A M : S KJ 9 K M ` O < LU < S ;: – ;=S —KA ^ < a M : KL M =O ? H : 9 S S ;K ;M : 9 = X \ ]

M L a J ;S < a K M G ;O < S x š =;KK < LG 9 K < [ L M R ^ L < a < ;U ;: [ K < OK M OK < L M : < KL < 9 KG <: K – M R =S 9 =O M 9 ==M – R O K M a M : KL M = X \ c

_M L ^ J 9 L G 9 a M =M [ ;a 9 = 9 : S S M O ;: [ < _ _< aK O ;: 9 : ;G 9 =O – ;KJ _R ==A _R : a K ;M : 9 = H e ? Y J < O < ;O OR < O OJ M R =S ` < X \

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