Marcos Michel de Souza
!" #$ ! % &'() *+,#-!.%
! " #
São José do Rio Preto – SP
/001
.2 +
! % - *
" 3 4 + .2 .'56%789/001
:;7<=0
> 7*
7? *3 @2$'(
A BC#$ ! % !"
D3 / = 3 E % ! ;% -* F%-% A!2 B1%-%A!2 B * @2$ '(? *3 #$ ! % !" GH
#C;11=//
Gostaria de agradecer, somente a Deus.
Agradeço a Deus por todas as pessoas que ele colocou no meu caminho durante minha vida acadêmica.
Agradeço a Deus pela vida da minha orientadora de iniciação científica, Maria da Piedade Rezende da Costa, que me ensinou os primeiros passos da vida acadêmica e me apoiou a vinda para a Biofísica.
Piedade costuma dizer que o orientador é o pai acadêmico do aluno, e eu tenho muito a agradecer a Deus por essa grande família que ele me deu. Todos foram muito importantes na minha vida, minha orientadora Dra. Fernanda Canduri e meu co-orientador Dr.Walter Filgueira de Azevedo Júnior, e todos os meus irmãos acadêmicos que eu tanto amo: minha querida e sincera Alessandra Renata, o dedicado e inteligentíssimo Márcio Vinícios, meus camaradas Paulo Camilo e Rodrigo Galo, Janaína, Helen, e todos os demais.
Agradeço a Deus por ter colocado em minha vida professores muito especiais, aos quais hoje tenho o mais profundo afeto e admiração: Dr. José Ruggiero, meu colega do coral que tanto me fez sofrer na disciplina “Forças Intermoleculares”, Dr. Eslso Drigo Filho, com sua gargalhada inconfundível, Dr. Jorge Chaíne, o coordenador da pós, que sempre me atendeu prontamente todas as vezes que precisei de sua ajuda, e a todos os outros professores, do departamento.
Agradeço a Deus pelos amigos que fiz no Departamento e por todas as pessoas que me ajudaram, minha amiga Cristiane, a sempre simpática Dona Rose, a prestativa Ilva, o Paulinho e o Barbosa sempre atenciosos.
Graças a Deus pelas agências de fomento à pesquisa CAPES e FAPESP sem as quais esse trabalho não seria possível.
Sou grato a Deus meus amigos de fora da Unesp, que também sempre me incentivaram nesta carreira: José Precioso Júnior, Anderson Rodrigues Silva, Fabiano Ferreira Ângelo, e ao meu grande amigo Thiago Laurêncio Rizzo.
Dou graças a Deus também pela vida do meu pai e minha mãe, que sempre lutaram para que eu pudesse estudar e sempre me deram incentivo para seguir na carreira acadêmica.
Agradeço a Deus até mesmo pelas pessoas que me fizeram algum mal, pois até as dificuldades me fizeram crescer.
/
+ ,
I*3
IH
@$
64@
D DD
DD@HC 3 3 DD
D/ % C + $ 3 DE
D=@%-%C% -* D:
DE%-%3 J DK
D;@L //
DF@%-% %-% /;
/>M$ /:
= 5 /K
=D> /K
=/ 2 %-% /K
== 6 N EE
=E>% * E1
=;. E1
E6 EK
ED 2 6 N EK
E/. ;D
E=6 3L ;/
=
E;O ;1
EF@@ H ;K
E1>.@$ FD
E:>!% !L FF
; 1D
6 1=
F@7> * . 1K
E
+
*3 DDH " /00= DD
*3 D/A B6 $$AB +
%-% D;
*3 D= %-% DF
*3 DE%$$ % -* //
*3 D;%$3 /=
*3 DFC-$ /E
*3 D1 3 P %-% /F
*3 D:@+ L %-% Q%-% /F
*3 DK P Q%-%A@B %-%A B /1
*3 =D% $$ =D
*3 =/6 )L 3 =/
*3 ==6 )L 3 A+ 33B3
A3B3 +A R+B =E
*3 =E 3 2 =F
*3 =;C!" =:
*3 =FC6 $ =K
*3 =1C6 33 ED
*3 =:C*"3 E:
*3 EDC %-% EK
*3 E/C! %-%" ;=
;
*3 EEC ;;
*3 E;C $ ;F
*3 EFC 3 6 + ΦΨ ;1
*3 E1C P 5 ;K
*3 E:C@5 F0
*3 EKC.@$ @ FD
*3 ED0C. $ F/
*3 EDDC. %-% +" F=
*3 ED/C. $ %-%" Q FE
*3 ED=C. $ %-%" 2 +" F;
*3 EDEC. L FF
*3 ED;C. LA B F1
*3 EDFC. LA$ B F:
*3 ED1C. L $ FK
*3 ED:C. L $ Q%-% 10
F + -$
H DD %-%3 /0
H =D>J =:
H ED ! N
%-7 ;0
H E/ ! N
%-7 ;0
H E=G ;D
H E= ! 3L %-%
" @ ;/
1 . $ "
%-% % -*
%-% % -*
Q%-% % -* A+ B
% % SA B
@@ @
@% @
T% T
%
T "3
T% "3
T% "3
TH% "3
-% -
-% "
-H% "
T T
@H% @
%
G G +R
%E $ %MUEA%M VEB
%-%W%>E % -*
:
%-%W@XW%>E % -*
"@$
%-%WQXW%>E % -*
"Q"
%-%WW%>E % -*
"
K /
>. ;000
=; +J $ Y" $ @ QG 3 A6B 3 " 3 )
@ % - * A%-%B 5
$ $ 3 $)5L$ @ %-% $
> M$ + 5 $ %-% "
J 3 @ %-% 2 2 J " 3 A B S3 > N 2 A Y- 2 . - . 2B > 3 >.* 2 $5 3 .@ @
3 P %/D/D/ A ZDD:KF[ ZD=EF; [
ZEEE/[B> 2 5 $53 @6
> $ 6 6 /=:1\ D1;D\ $
L" D:F[ * $ 3 P ! $ L @ L %-% " + $ J $ 3 $
D0 $
3+?Q>+S;000$X X
=; R " /0 X H+ +3+ X+ X$+ + 3 A6B + $ RR " 33 +
H+ % - %++X
A%-%B 3R3 R+
X+ H+ +
%-% X +
H+M$+RS$+%-%"R+
X2+ R++3 %-%R X 2 3+" R+ +3 A BR X S3 H+ N X R 3X+ A Y- 2. - . 2B R + >.* 3 3 + R +3+ + 3 .@ @ 3 ++ 3
%/D/D/ A ZDD:KF[ ZD=EF; [ ZEEE/ [B H+ X R X
+3+3 @6
H+ + 6 6 /=:1\ D1;D\ $X
" D:F[ R $ 3 X + + 3+ +H+ R + X H+ X + %-% " R+ + 3 SR3 + $ $ + R 3 R +
DD 0 123
00-$ 4 '$
@ ;000 H)K:\
$$
$L$ @ " H
+ ) ) -)
$ L7
3 ++ * I 5 % )A3 DDBA?Q>T
* +/00;<-#-- /00;B> (
$$
3 ) $ ]
) 3 +$ 3
*3 DDH " /00=A-#-- /00;B
! 3 H 5
2 3J "
) 3 D\ ! /00= 3 ::
D/
// > D;// )
3 K0000 !/00E
H /00E/001
A?Q>T * +/00;%3 H! 6
' $C/00;B
@ H 5 P
A& ^+,B) J
J) ) A>H6@-DKKDB
!"$ H5+Y
( !3 3 ( 5
%+L5+ 3 $
Y 5 ) 3 $
J 2 @ 2
$ Y+ D:F:) G
2 ++ ]+ M
J 3 + A@G. DK1=<G._>
! >-H6> ! @ H#!6# >.! G! C .!.4.%
RRR 3$44<.>!@!6@. !6@!%-!#> >T@
!H.> >T@RRR3B
$
" ! 2 5
NG+ +3 5
` a ` 2 L
( 5NG@H
D= $ - D::/ 6 ^+ 3
^+
# $2 $L
+ Y A Y "
B *
Y 6 ^+ ) +3
$ A@G. DK1=< $
H G! C .!.4.% RRR 3$44< .>!@!
6@. !6@!%-!#> >T@!H.> >T@RRR3B
# $ 35 A B ]3
" L ) D0K >3 2
. (A>.B @ 3E/;000$
H 3 6 AH6B " H6
3 # .$5 + 5 DE\
$ H Y3 $
A?Q>T * +/00;B
@ .
@) A@.B]H
3 " 3
) H -$ 3
L J
A.Q /00D<-> @.>/00;B
DE
056 $ 4%"
>5 A"B
) ( L 3 D ( >
5 )
L 5 @ a 5
$ )) 5 2L&$,
3 L5AG>!HbG>!H/000B
% ) $$
3 J $
A@T@6?@ b %@6^. DK1/< >. .@-H>. /00EB @
2L L$ $ 3 3 -3 5 $
3 DKDD^ S DKE;3
D $2 5 ! $
$ 3 ) $ 3-3 $
Y 2L A*3 D/ B $5 +Y
Y^ S DKE1A%#T6!b!@ ^/00/<>..@-H>./00EB
@ 3 5L5>
L5 $$ LP 33 5
L > P )
$ +
$ 3 @
D; $D
AG>!HbG>!H/000<>..@-H>./00EB
-cQ/>ĺ c6
-cQ/>ĺ c6D%
DF @ ) L 5 3
L5@ $
2 3
)d 3 3 " >
L5 5]3
2 ) AG>!HbG>!H/000B
-
A@%B3 AT%B
A%B 5 $$ (
$ $5 $ %-%@@+" 3
A@G@6>^ !H @ DKKKB @ %-%
$ ]$"3 ATB3 AT B
("3 ATH%B) 2
6 )d -@ A*3 D=B
Ae>?.^@ /000<-> @.>/00;B
D1
-) +
+L 2 A%-%B
A@@B )$ Y" Y
@ J +
P $ J3 2
DE) A%-%B /0) A@@B AQ!6.Q*! b^6!QDK:K<>.
.@-H>./00EB
@ %-% + $ L 3
+ T $ Q%-%
3 ) $ $ 5
H 5 AT !HH!H@ DK1;>.
.@-H>./00EB
H ) Y
+ 5 $ -@! 5
) 2 5 $ 2L
"Y
5 Ae>?.^@
/000B@5 3 $ Y 3
2 3 $
+ ! 5 !" #$
!%$ L3
AE+" 28=E9 B 3 ) 5 Y"
+ 5 + 3 Ae>?.^@ /000B
@ % - *
D: 3 > 3 + 5
Y Y3 +
) 5
M (3 +
* 5 ) 3 +
J
$$ A@..> /00DB
> 2 %-% + $
+ $$ L $
2 ! $
2 +$)
2 +
07 4 8 ! %" $
@%-%
$ 5 A!@e!G!> /00=B@%-% $ 3
3 -3 "
3a A%B 3 3
A"BD A*3 D/BA@-#6 /00EB
> $$ + % - *
"3 )2 3
) ] %-% +
DK $ 3L %-% + AQ%-%B "
L3
+ " Q%-%
3 A@-#6 /00;B
@ 3L %-% + DKK0
=/[ ! 3L +
/:[ 2 +
%-% A! @e!G!> /00=B ! /00= $5
N %-% /=[
A! @e!G!> /00=B !
$$ 3
+
$ 5$
%-%A@-#6 /00;B
09 2:
%-% $ 3
$ $
- + ) J7
2RS A/000B %3
b! SA/00/B
2RS A/000B 2
AB Y3
/0 @ 2RS A/000B 2 %-% "
+5 ++" 5 %-%
)) +3 & ,
> 3 & , $2 $ 3
73%-%)+
+ * 2 33 %-%)
++"a +
+"a! 5 H DD
H DD %-%3 e>?.^@ A/000B
Grupo2
Grupo1
Inosina e Guanosina Variada, exceto homotrímeros.60 – 180kDa
Homo Sapiens E. Coli, Salmonella
Exemplos
Inosina, Guanosina e Adenosina Inosina e Guanosina Inosina, Guanosina e AdenosinaEspecificidade
(substratos
naturais)
Variada, exceto homo-hexâmeros Homotrímeros HomohexâmerosEstrutura
Quaternária
80 – 100kDa 110 – 160kDa
/D -) %-% 3
$2 ) +L 2 5 2L
) 3 !
+ A ) %-% "
B $ "
> 2 %3 b ! S A/00/B
) $ ) 2 )
) 3 3
$ @ $
$
@ - * A-%B 2
) L 5 +" 5 + $
Į4ȕ@3 -* A-%B
2 ) ) L
5 ) Į
//
*3 DEC%$$ % -* @ 5 A@B5 % -* & # #@ 5 AB5
% -* @ +" 5 AB5 % -* '# #
0; ( 6
@ L Q%-% ) $L 2
$ >
2 A 3 B)
L Y" $A$3 D;B! ADKK1B
3 )
) ) " %-%7D
/= )$ 5
J %-% D A>. .@-H>. /00E<
-> @.>/00;<!6>- DKK1B
*3 D;C%$3 J 5 A !6>- DKK1B
@5 L %-%+
+ $ Q $
3L "
$ $L a J A/00;B
P$ %-%)" $
L %-%+ > Y
3 3L @2$
A/00=B) Q%-% /=[
> A/00;B5 3 DF
/E DF@B> a ADB ) ) 3 3
5
L 3 $ 3 A3 DFB 3 3 $ 2
>AFB 2 L @/E= A-/B )
5 3 -3 ! +3
3 3 $ 3 3 ) (
>AFB
*3 DF-$ Q%-% A/00;B
/; 0<
# )d +Y3 ] %-% +
3 > 3 5
EEDD;/K 5=K/E3 +3
)d J ) ] " E0\
$ )
EE\ DF\ +
+ + 3 J
A-> @.>/00;<> ! DKK:B
. %-%
+""
/00D - /00E ! /00D + Y
+X %-%
Q - $
3L %-% 2 " X$
A-> @.> /00EB
@ %-% %-%
+5 M 3 P
L $ 3 α4β 2 β
+5α
A*3 D1BA-> @.>/00;B
/F
*3 D1C 3 A@BPAB %-%
A-> @.>/00;B
@ L %-%=E\ L
Q%-% A*3 D:B 5 L L ) L
" Y" A-> @.> /00EBA*3D:B
*3 D:C@+ L %-% Q%-%2 3 @A>6%!HDK::B@ $+5 " =E\
3 %-%
5 %-% > $ %-% Q%-%
AY3 %7D1=B
/1 $ 6. /;[ 5
) P@$ 6.
0K[A-> @.> /00EB
*3 DKC P Q%-%A@B %-%ABA-> @.> /00EB
/: 5$* "
> + M$ 2
) $ % - *
" A $ B
# $2) %-%+ $ L $
%-% " +
/K 76 6=
70$ 23
@ 3 % Y3 . 3 .
% 2 @3 3
3 A% #$ Y 6T .B%@3C
3 ] 6 %) H @ %-%
5 3" A@..> /00DB
@ "
P %-% /1 E=1 Y"
$ ) /1;1D
75!236
!23 =
2 5 ) 5
J3 Y# 2
)a ) Y
. ) )5
A!@e!G!>DKK1B
> 3 $ DK=E )
R2 N
5
2 A#6#NbT!TfDKK/<>..@-H>./00EB
G 2 5)%2
=0 5Y $
!DKF05
3L 3 )
3 2 2 DKF;A #-!
b'>Q-.>-DK1FB
> M$ 2 5 Y3
L5$5 5
N ! 5 $L
Y3! 2 5
5 Y3 > $$
5 Y3 $ P
3 =DA@-#6/00DB
! 2 5 2
) ) 2 5( a)$$
L 2 7 5
>) 2 2 5 Y3 5
( a $$ ) ) $$
) 5 5 )
%" 2 5
Y3 ) L " Q )
2 ) 5 5
Y3 5 " $ )
5 ) ] J "
=D
*3 =D% $$ 5 Y3 A@-#6/00DB
$ ) L 5 Q
P 5 $
J ) " @ 2
J 3 3 $2
5
2 L ! 3 5 ( 5
$@ )
) ) A!@e!G!>DKK1B
> 5 5 $
$ 5
2 " !
$ $+ 5 A! @e!G!>
DKK1B
Purific a ç ã o
Crist a liza ç ã o
Cole t a de da dos
De t e rm ina ç ã o da s fa se s
M a pa de de nsida de
=/ % 5
2 5 N 5 $
+ N(
$ L$$$ 2 53 A
Q 3 $B J
$ A #-! b
'>Q-.>-DK1F<@-#6/00D-> @.>/00;B
% 2 5 $5L
$ )5 L$
) +3 )
)@ *3 =/
*3 =/C6 )L 3 3J $ ) 3 2 5
A-> @.>/00;B
@$ $ 3J "
- 3 " 5 L 2 3
== 2 2 O 5
) 5
2
( A 33 B
( 3 2 L$
L AT!Tf b #6#N DKK/< -> @.> /00;B
A*3 =/B
-= !23 =$>
. 2 ) 5 2 5
Y3 A3T!Tfb#6#NDKK/<@-#6/00DB7L
$ +$
> L ))$
L$ @ $ $
) 5 53 ($> +
) 2
3
" ($ )
$
@ $ )
J $5 $ + @
L $ $L 5 ) ) J
M 3
@5 $ $ J 7 3
=E A*3 ==B @ 5 3 A+ 3 B 5 )
2 & , 3 5
2 3 2 $
)L $ Y 3
2 ) 3 >)
L3 A 3 $ YB$
$ 3 )d
3 2 A#6#NbT!TfDKK/B
% 2 "
$ $ C $ Y C
3 ) ) $
@ 3 5 Y3 2
3 $ 5 ) $ Y
3 )
3 A@-#6/00DB
=; . $ $ Y 5 3
3 )) )5 3
5 L3 ]
$ Y
!236
@%-% 2 3J$ A.Q
/00D< - /00E /00;B @ %-% ADμ ] /;3DB
; - Q/%>E 3 $ )
+ D00 H Q :0 /;\ %!T ==;0 /; 3 /
) D0 5 2
$ 2 3
23
@ 3 2 $5 5
@ 2 5 !23&" ;
5 3 5 A;7DB ) 5
3 ML" 2 % @ 2 5
) 3 ) 5
2 A2 3 B - + S3
3 3 )+ )3
$ Y+$L
) 5 ($ $ $ @ 5
5 ) + 5 $ 5 )
5 A#6#N b T!Tf
=F
A B AB
*3 =E 3 2 A B AB
?@1--.&0ABCD
# 5 MYL# 5
-
5 (L@
3L3
!35 $ <
# 5 L
$ $) 3 J P M
" 3 $ ) $
E 7
EFGG$G- ?0D
-( L>ME
ADB $ - >
=1 5 $ 5A 5 L B)$
"$57
. = × • A/B
@
-$L J # 5
") !")"
LLL) LL) J
/ʌ /ʌ4/ /ʌ4= /ʌ4E /ʌ4F ( J
! J3
D/=EF-") Y )
J/ʌ41 /ʌ4; > 5
J )) " $
!J3 DE
3 =F =D>3 5
> DE $ 3
$ 5 L 7
P 3 ( 3 +" 3 !
$ J " A "B
5 L $ "A$Ba3
AĮȕ ȖB 3 =; > Y
H
=:
H =D>J
I
J /
I$/
/ 2: = " & H
- D Z
ĮȕȖ
6 /
ĮZȖZK00ȕ
K$ E /0
ĮZȕZȖZK00
- / / 1
ĮZȕZȖZK00
J$ =
/ 0
11
ĮZȕZȖZK00
- D 2 11
ĮZȕZȖgD/00K0
L D 1
ĮZȕZK00
ȖZD/00
=K
E0
@
) ) ) M .
$ "
J 3" 5
aA#3B
6=
! $5 Y5
@
) L$] )
" J
J3 A^HH! DK1:B
@3 Y N5 Ȝ
ȯZ#ȣZ#4Ȝ#5 % S+ZFF=WD0/13ZFF/WD0=E'!
Y7ȜA[BZD/E4ȯASGBA^HH! DK1:B
> N3 5 $
L) " 5L $>(
$ $ + N5
L$PA^HH! DK1:B
? 33 " "
.+ ) M A a3
53 a3"B L
) ) >"
) "J$ L
2 $
ED
M 5 șș5
A^HH! DK1:B
*3 =16 )L 33A #-! b'>Q-.>-DK1F<-> @.>
/00;B
@ $ $ $
) +( Ȝ
)7
șȜ ?7D
@" 33 Ȝ/
$2 ) $ 2 @ 33
$ + 3
! ) + 255
L L A^DK1:<!@e!G!>DKK1B
O ) A 3
5 5 35B
$ ) ) HZ c$cR$R
E/ 5AB$ Y J
" )ACHBZABA^HH! DK1:B
. 5 5 Y
L 5 *
* 5 5
A^HH! DK1:<!@e!G!>DKK1B
* 2 L * J Y J
M$L )
( )
=∑
•" AEB
M $ JH)" $ >$L
5*$ ) 5*
5 ML$ $ a ) )
7
# 3
3 Y ) 5 3
O 3 )
>$ 89<
$ 89D@ 5
< 5 * @
$5 $ ) "
7 4 × • × =/π <
& × • × =/π <
× • ×
E=
. $$ @
$$ $53 $
" % @ 7
MZ/ʌ NMZ0 MZ0
M$Z0 NM$Z/ʌ M$Z0 AFB
MZ0 NMZ0 MZ/ʌ
O ) M L $ $
2 M @) )$
L 3 7LF# G3NG&A#3ZB5 $ >M$L $"J
N$ " 337
5OMLGL5FP ?BD
5 + L Y
L P 3
A^HH! DK1:B
. )M$L $
"J N 5 $ 2 * AB
) + A^HH! DK1:!@e!G!>DKK1B
@ + $ 5 ]
5 @ A O $ "
OE $ " 3B )
+ OE 5 3
$ "8AOOEBW9$2AB.A 5
5 $B@ + 5
EE
F?D"AǻOMB A:B
2*AB7
FL"8ALǻOMB9 AKB
O ǻOZL +
- 7
LF ?D"ALMBZ-*A+SB AD0B
*5 3
( Z0$5
. ) + L 5
$ J L $
2 Y J$ L37
( )
∑
(
)
=
• = 5
6
6 6 7
8 7
0
D
/
" π ADDB
@ 3 2 I $5ǻO
7723 /Q
> Y" $
) $ N % +
Y3 #-!
b'>Q-.>-ADKKFB<6!-HQADKKEB<!@e!G!>ADKK1B
* 2 Y" 5 " "
NM $Y" a
L )M $ L$ O "
N $ "5L
E; + A! @e!G!> DKK1< @-#6 /00D< #-! b
'>Q-.>-DKKF<6!-HQDKKEB
) 2
M + 7
+ H+ )
35 2 + ) $
2 A!@e!G!>DKK1<@-#6/00DB
- % -% 5 "
N ) 5 3 5
) . ) 3
5 5 35 )
23 A %
(BA!@e!G!>DKK1B
> % ) N 5
$ 3 )a
$ 3 Y5@
) 23 % )
A % (B$] 3 Y
5A!@e!G!>DKK1B
!" ) +
+ H+
+ @ A + H+B
< 3
)
EF
O + N 5
"+ ) > A
B 5 * P
$ ) P 5 *
A!@e!G!>DKK1B
6$ 5
P 7
( )
=∫
( )
(
•)
. 70 ρ "/π AD/B
A"/ʌAhW.BB 3 $
AGiZD4GB)7
( )
=∫
( )
(
•)
i i / . 6 7 7 0 πρ AD=B
$i5$
@ 3 Y $ ) *A.B 5
5 2
)
(
)
∑
∑
∑
∞( )
(
)
−∞ = ∞ −∞ = ∞ −∞ = + + − = 3 # 9 3 #: #3 0 . 9 : πρ D " / ADEB
*A+SB +
"J#3 P ȡA"X2B
"X2 L A!@e!G!>DKK16!-HQDKKEB
E1 79$
" NY3
5 % 5 $
$ ) L
> 5 L) )
A! @e!G!> DKK1< #-! b '>Q-.>- DK1F< @-#6 /00D >.
.@-H>./00EB
% $ 35
@ ") 5
7AB5 P ( )
L) +3 Y
< AB 5 ML )
$$ <AB Y ( <A$B
5 ) 2 ML $ 4
$ ML )L
7;I$236
@ $ )5 )d L +Y3
$ + 5
$
JL A6Q>!.
DKKK>..@-H>./00EB
@ 5 5 $ $
-5 + +Y3 5 M
E: " $ > M
2 J3 )
A!@e!G!>DKK1B
- $ +Y3
$ 5 3
;0\ +3 5 f L$
) $ $
3
A-> @.>/00;B
% 2 3 @6
3 %EA%EDKKEB!3 2
+3 35 3 @
3 =: "3 3 +
@@ +>..@-H>./00E!!@e!G!>DKK1
EK 9/ 3
90!23 23 /Q
> %-%" )"
$5J$ A.Q /00D<
-> @.>/00E/00;B J " 0;"
0F"01! N D:;[
) " 3 $ P
%/D/D/ a ZDD:K[ ZD=EF[ ZEEE[ a3
αZβZγZK0VA$ Y a2
3 B %-% ZD/1F[ZDEEE[
ZE1F[ %-%"
*3 ED %-%!A B%-%" AB 2
% %-%7 $ /:::=/"J ) ==E=F
"J 3 AH EDB
;0
H ED! N %-%7
@
A[B DD::D
A[B D=E;;
A[B EE;
T %/D/D/
-( 32 DF0
-("J ==E=F
2 A\B KK=
6iXA\B 1K
A[B DKF D:F
2 ] A\B K1F
6iX] A\B E1
% %-%" @ $ E=K1=D"J
) EK/KE "J 3
H E/! N %-%7
@
A[B D/1FEFF
A[B DEEE/K;
A[B E1FK=/
T %/D/D/
-( 32 DF0
-("J EK/KE
2 A\B KKD
6iXA\B F1
A[B /DE
2 ] A\B KEK
6iX] A\B //K
;D 95I$236 * @
2 M
%-% " +" % 2
3 @6 A-@G@e@ DKKEB 3 %E A $
%M U EB $ AB 6
E=
H E=CG
I$236 ?RD / ?RD α α α α?SD ββββ?SD γγγγ?SD -?TD
-U?TD
-!?TD
;/
97/ (
> 3L 2 2
3 6!*@;A#6.Q#>G DKK1B 3 %E
> 5 6!*@; 5 5 L"
G+ A%# : "3 # #B A*.Q!6 DK/D< -> @.>
/00;B
>$ 3L
EE7
H EEC! 3L %-%" @ %-%" @
6M
7B&PCT0&C;T
/ 0C&;5R
/ 57&CBR
S (? D BA0
S = (? D ;<P
6M
99&<BB45&095T
/ 0A&90R
/ 5;&<<R
S (? D BA0
S = (? D 9;9
;=
99 23 6
@ ) L %-% "
ML $
+5 M 3
%-% P L $
3 α4β2 β +5α
! 2 7
P $ 3
+ A3 E/B
@ Y 3 ] $5 3 J 5
L3 2 + 3 L )
;E
@ $ 3L %-%
$ $ + A$3
E=B! $ )7ADB %-% 2L
< A/B 2 $ " $
<A=B 5 5
3L $2 ) $ 3+
) 2 "
*3 E= P 5
;;
% )$ ) $ 5
2 3 3
$ 3)
$ > )
5 P
5 A$*3 EEE;B
*3 EE P 5
;F
*3 E; $%3 2 N GRA6!!DKKKB
% M3 A B
L ML )
3 @ ) L 5 $5
3+
;1 9;V
% $ ) ) 2
1 ,/ 6 / 6 L / ) J
L L$ a3ĭȌ
@ ) K=/\ 3J
$L$ 3 6 + ΦΨ A*3 EFB F=\
3J + 3J 3
A0;\B3J @ L
P A/** B $
$ $ $ ) 3 L -
M $ P
;:
> a 2 L ) ,
-/ > T* ) 5 >$ T*
a33 $ A @.^>?.^ DKK=B
> T* 3 5
5 T* >$ T
* $ 0; A>66. DK:FB > $ T*
j3H0/; T $ 0=:$ T
00/
* L 5 $5W/1X7A>?! DKKD< #HQk
DKK/<^@.Qb.@-!6DK:=B) )d
L > 5
)d L = 2
$ ) 0E; ) M
$>!H D=:FD! D/D/K
2DDE
@5 L 2 5 L /6I
a33 $53 N% >6 A6l-T!6DKK/B$ a33 $
" 0D[ D0 $ > $
3 00DF[a33 D1D
;K 9<
$ P
5 3 )
$5 ML @ 3 E1
P ML
*3 E1 P 5 T
3 N GRA6!!DKKKB
@5 23 J+3
) 5 L3 5 L3 $23
5 L3 4 L @ DEE
F0 @5 L3 53 5 L3
3 5 3 LY" @ DEE @
@3 DE; A$3 E:B
*3 E:@5 T 3 N GR
A6!!DKKKB
# $ " 3
$5 3 @ +L
L L )
5
FD
9B"
% $ $ %-% 5 @ /=D T D:K ML
$ $ 5 L3 > T D:K
J 5 L3
L33
@5 5 L3 3
$ $ @ D/0 ) 2 J
L a ==D[ @
*$ 5 $ 3 G
/0; a )=;[ A$*3
EKED0B
*3 EK $ @ a T 3 N GRA6!!DKKKB
F/
> $ a 3
Ja33 ML
A@B
AB
*3 ED0A@B $ AB $ T 3 N GRA6!!DKKKB
F=
> L @ D/0G /0;
5 L3 $ %-%
" +" ! 2
%-% " 2
2 + 3 ) ML $
- L$ a
5 L3 $ @ 3 EDD
*3 EDD %-%" +" " T 3 N GRA6!!
DKKKB
@5 5 $
%-% A%C
FE 2 +" AY3 D:0B Q
AY3 DT/0B
- L $
) 5 L3
G
) 5 L3 !
5 L3 L
3a 3 AG3 ED/ED=B $2 2
) " ML+ $
F;
*3 ED= $ %-%" 2 +" %3 2 N GR A6!!DKKKB
# $2 5 L3
L $ $ Q%-% A
% - * B a L
Q%-% 3 @ @ +
$ $ $
%-%
* %-%A" @ B
Q%-%" 3 A %C% S
Y3 DG/QB % 2 3 .O^@
3 %EA%EDKKEB@
Į L2 Q%-%Y$
%-%3 " )$ 2
FF G /// Q%-% %-% 2 "
/00 @ /0= G /0; G /0; 2
P @$ $6.0=0/[
9C (
# L $5 3
%XA! @->/00EB %-% a
$ 3 $
$ > ) 3
3 ) a 3
3 $ ) A3 EDEED;EDFB
F1
*3 ED;@ L$ ]3 $ 2 5 %3 2 %XA! @->/00EB
@ L3 3 3 $
"3 5 L3 $ 3
53 3 $$ $ 5
Y 3 ) 5
L3 ) L
F:
*3 EDF.% !LA$ B%3 2 %XA! @-> /00EB
@ L $ 5
3 5 L3 a 3
3 $ 3 $ " A*3 ED1B
FK
*3 ED1 L $%3 2 %XA! @-> /00EB
# L $ Q%-% 3
LA*3 ED:EDKB% 2 +
+ L$Y"
$ L %-%M"5$! $
" ) " %-% 5 )
10
*3 ED:% !LP Q%-%%3 2 %XA! @->
/00EB
1D
;3
> 2 Y 2 L
% - * + 2
)+M
> L
%-% $L$ 5 3
@%-%" + ]Q%-%
5 $ 3
+ 7%-% 2
@ @ ) 2 +
$ @ @
! 2
%-%3 2RS A/000B )
+ ++"a
# " $ 2
2 + L
@ %-% a
L$ )3 "
$53 $) Q%-% $
L 3 $
> $
1/ + 2
. 5 ) + $
$ +
! $2$
+ $ $ 2$
]H
1=
@T@6?@ 6%< %@6^. 6! '6< % ++X +
X+X G + $ +XX 3# N % & $
/EFDD=1F==1F:DK1D
@G@6>^ < .@ ! '< .< < 3 + 2
6@4.H+3X , &$/==
/FD/FKDKKK
@..> @<.@-H>..<.Q?< <%++X
X @ X + X 3
X N% U&$E0/::DKF:/0=/00D
#-! H < b '>Q-.>- -< U%U! 7
@ %DK1F
>?! ' # @ + X ) + SR
+ I $/;==E1=;/DK:1
e>?.^@ @< ^# ^>?.^@ !< .Q#T@6 < %
++X 7 %-% $::
==EKE/;/000
@-#6*< (
1E
@-#6 *< *@! G< @..> @< %@ @ .< ! @e!G!> ?*'
N% N%U/ $=/1FEFFEK/00;
@-#6*<>..@-H>.<. G@6T< <.+
++X " R+ N% N%U /
&"=D=EK01KDE/00E
$ %M - E H+ %E 7 3
X 3 +X U$;01F01F=DKKE
> ! .H< 6>.Q 6< %@6^Q '< < +3 + 3X
X + 3 )
&$=K=FF:;;=1;EEDKK:
>6%!H*<) 3R++ + 3
/ $DF//D0::DD0:K0DK::
>H6@-6.^#@6G6>-.. E6
' 7T ^3 <DKKD
@G. < # !> 6<!-.!- Q-< <6$ 4 1 2:
1; !@e!G!>?*'6<123Y$
) 5?@5D $ #.%.% DKK1
! @e!G!> ?* '6< @-#6 *< >. .@-H>. < < X
+ ++X /=@ N%
N%U/ &$=0:=;E;;;//00=
! @->? -% U6.6 ,%IU . 7@
./00E
G._> ! >-H6> ! @ H#!6# >.! G! C .!.4.% $ 7
RRR 3$44@M+/00F
>. .@-H>. < @-#6 *< %!6!6@ 'Q< < X
+ ++X "R+ $N% N%U
/ &"=0:=;;=;;K/00=
>. .@-H>. <
L#-!.%.% /00E
#6#N @< b T!Tf 6<U!
1F
!6>- < .H>!^ !6 '< T#@ ?< < %
++X / X + N% U $ =F =K DD1=;DD1E:
DKK1
*.Q!6 6 @< `> + * H+ . `
%%-% /UI U.5@///=0K=F: DK/D
Q!6.Q*! .< ^6!Q -< .@X+X
"6 N&$D//@E/DE/;DK:0
^@@.Q ? b .@-!6 X 7
3+X3 3 NU $//D/
/;11/F=1DK:=
^HH! 123YI>;6' 7T
DK1:
@.^>?.^ 6 @ < %6>Q!^ C @ 3 +S + +
) X# U$/F;01/:=/KDDKK=
6!! !< N GR4N @ $ 3 3
X#IND/;A/=B7D;FF;ADKKKB
11 >66. %! '6< >#6@ T@< + + 2X
++X + X % &$FKKE=
K;K/0006$R
#6.Q#>GT-<G@T-@@<>.>-!'<6
X+ "S++ UNU
$;=%=/E0/;;DKK1
-@G@e@ '< @67 S 3
U&$;0D;1DF=DKKE
-> @.>><@-#6*<%!6!6@'Q< <X 3 +
%-% DK[ N% N%U
/ =/E1:K1KE/00E
-> @.>><(
#-!.%.%/00;H
-#--%<? @.<* >k^<k!<! e-T@T<6@GT >-!<
H+ QG / " Z41U;:DK
:/F/00;
%6>T6@@ ! >-H6> ! ! H#!6# >.! > !.H@> > 6> !
'@-!6> $6' /00;
1: %#T6! '< !@ ^ .!< . X $ R
++X N% #D<=FDA%DB7D/;A/00/B6$R
6Q>!.TU%U6 U D0DD@ %DKKK
.@-H>. < L
#.%/00E
.Q ?< @..> @< .@-H>. .< < . %
-%++X "R+Q
%N% UE0:/0E:/D;/00D
.>!@! 6@. !6@ ! H.> >T@ $ RRR3
@M+/00F
G>!HG>!H'T%6@H? $)H
*-@T %@37@5 ./000
?Q>A?Q +>3 2 BT * +/00;
1K
< '
$ I$236
O M &$, 2 N
P 5 %
P ) * ! )
"A;ʌ!"B "5 3 4 PA + "
B 5 3 +
3 $ 7
( )
( ) (
)
< =
.
0
7
7
π
ρ
"
/
−
=
∫
" 5($ 0,#3>
A6!-HQ DKKEB @3 $ Y
H7
( )
0
( )
#3
( )
#3
(
#:
3
9
)
.
:9
#3
+
+
=
∑
α
π
ρ
D
"
"
/
m0A#3Bm 5 "JA#3B"MXM2M
L M5 L L + S
"ĮA#3B5a3
- ) " +Y
Y
N
. A#3B Į m0A#3Bmn m0A#3Bm
%5+ 5$a3
:0 ) ) " N Y 3
5 % 5$
$ )
L > 5 L ) )
A! @e!G!> DKK1< #-! b '>Q-.>- DK1F<
@-#6/00D>..@-H>./00EB
% $ 35
@ ") 5
7AB5 P ( )
L) +3 Y
< AB 5 ML )
$$ < AB ( < A$B
5 ) 2 ML $ 4
$ ML
I$236
@ $ )5 )d L +Y3
$ + 5
$
JL A6Q>!.
DKKK>..@-H>./00EB
@ 5 5 $ $
-5 + +Y3 5 M
L ) $ +
:D 2 J3 )
A!@e!G!>DKK1B
- $ +Y3
$ 5 3
;0\ +3 5 f L$
) $ $
3
A-> @.>/00;B
> 5
% $ %
)5 5 7
( )
( ) (
)
? .
∫
+
=
ρ
ρ
ZA$RB$@ % 5 $ a P
@ ) P
#57
( )
=
∑
∑
(
−
)
∫
[
−
(
+
)
]
? . # # # #:
#
#
#
0
0
.
o o
/
"
/
o
"
/
o
D
π
π
) #37%AB5
*0A#3Bn N
( )
0
( )
#3
(
#
3
-
)
.
#3
+
+
=
/
∑
//
π
> $ )
:/ NA)B $ +
3 )$ +S A)%OB * $ 5 $
5 " N
.- Y" . $
2 )$ $% 3 2 +
3 5 L3 * 5 3 8A%E
DKKEB A B /0[/
a L $> )$ 3
+ A)
A* B B )$ P A>. .@-H>.
/00EB
>$ a L"
A
( BA>..@-H>./00EB
> a $ 5 " L
% 6"J
+3 $ ) "J 2
J + 6"J "
L @ )
" ) 2
) ) L
$ " 5) YA-> @.>/00;B
@ a
% AB
:= 3 A B5 2
(
)
0 7
(
)
(
)
0 = D−
5 /
0 0 /5 7
(
)
(
)
(
)
[
]
⎭⎬⎫⎩⎨⎧[
−(
)
]
⎭⎬⎫ ⎩ ⎨ ⎧ − − =∑
∑
∑
∑
∑
∑
∑
/ 4 D / / / 4 D / / 4 4 4 5 5 5= 0 /= 0
(
)
/ Y M"J$ ' 2 $
' 2 >
2 3) 7
( )
( )
∑
= = 5 6 6 8 0 0 D / / / ε5$ 86 + Pε5
3 3 D $ M
"J 3 A.H>#H b '!-.!- DK:K
-> @.>/00;B
> AB L $ CD A $ B
0A BL"DA B@
2 0$ ] "2 M a
M )+ M L" 0 / 0 /
:E
. "3 L
5 M
L "X2% *A.B
) $ L L
% %AB
@ ]5 )7
A"X2BZ89A"X2B
A"X2B $ 2 @
% $ a L
L @ ]
% %AB5 % %AB
AB 6 a3 Įȕ Ȗ
] 2 +3 7
(
)
=
∫
( ) ( )
@
2
α
β
γ
) L $ "2 ) +$ 5
3 5
@ "2 6AĮȕȖB 2 %AB
N %AB
L $ 5
@Y $ Y" $ 5
$ 2 $ )$ Y"
:;
) $ 5 $ ?$D<
)$ %$) ?D%<
<$ $ a 5 $
A>..@-H>./00EB
- Y $
$)$ % 33
$ Y" 3 !+
L" % L
+ + > L"
% 5 a
A6l-T!6 DKK/-> @.>/00;B
2323
@ + L
5 5 $ >
5 5
$ A* B
$ A*B @ ) 5
$5L
> 2 7 6 A28B
AB
> 65 "7
∑
∑
−
=
#3 #3
0
0
3
0
2
:F S
% a $ 0\ 28 D00\
G 28 pE0\ p;0\ )
$
2 2 )$
$ )%O %$ a $2
3A>..@-H>./00EB
> L 3
)$
2 3 AB ! 5
J33L)P
$ L )%O
a A>..@-H>./00EB
- )$ & ( .
+ 5 )$
$$ ! ( )$ $
$A>..@-H>./00EB