Physiological and Biochemical Response of Tonoplast Vesicles Isolated from Broussonetia papyrifera to NaCl Stress-文献传递-植物通论文库

以构树幼苗根组织和叶组织为试材，测定不同浓度NaCl胁迫下液泡膜H⁺-ATPase的活性、液泡膜脂肪酸组成和膜的流动性。结果表明：在低浓度NaCl胁迫下，根组织中液泡膜H⁺-ATPase的活性有所增加，当胁迫浓度为150 mmol·L^-1时，活性又有所下降，但接近对照水平。而叶组织液泡膜H⁺-ATPase的活性在100 mmol·L^-1时降低，高浓度NaCl（150 mmol·L^-1）处理后又有所升高。液泡膜脂肪酸基本组成为C16： 0，C16： 1，C18： 0，C18： 1，C18： 2和C20： 0。盐胁迫后构树幼苗液泡膜脂肪酸组分相对含量发生变化。根组织液泡膜脂肪酸不饱和度下降，150 mmol·L^-1 NaCl处理后有所升高；而在叶组织中，液泡膜脂肪酸不饱和度在50 mmol·L^-1 NaCl处理后下降，随着盐胁迫浓度的增加，不饱和度又高于对照。NaCl胁迫下，构树液泡膜流动性和脂肪酸不饱和度的变化趋势相一致。脂肪酸不饱和度下降时，膜流动性下降，反之则膜流动性上升。研究结果为深入了解木本植物液泡膜H⁺-ATPase活性、脂肪酸组成和膜的流动性在盐胁迫下的适应及其3者的相互关系提供了参考。

The activity of H⁺-ATPase，fatty acid composition and membrane fluidity of tonoplast vesicles isolated from roots and leaves of Broussonetia papyrifera plantlets under salt stress with different concentrations of NaCl were investigated. The results showed that V-H⁺-ATPase activity in roots increased at low level of NaCl concentration (50 mmol·L^-1). When NaCl concentration exceeds 100 mmol·L^-1，the activity was similar with control. In leaves，V-H⁺-ATPase activity decreased at 100 mmol·L^-1 NaCl，but increased at higher levels of NaCl (150 mmol·L^-1). It was found that the fatty acid composition of tonoplast was consisted of C16： 0，C16： 1，C18： 0，C18： 1，C18： 2 and C20： 0. Alterations in relative content of tonoplast fatty acid composition under NaCl stress were observed. In roots the extent of unsaturation of tonoplast fatty acids decreased, and then it increased under 150 mmol·L^-1 NaCl. In leaves，the extent of unsaturation of tonoplast fatty acids decreased under 50 mmol·L^-1 NaCl treatment，however the extent increaseal with the increase of salinity,leaves and was higher than that in control. The changes in tonoplast fluidity coincided with those in unsaturation of fatty acids under NaCl stress. Tonoplast fluidity decreased with the decline of unsaturation of fatty acids. The objectives of this study were to gain better understand of the adaptive mechanism of H⁺-ATPase，fatty acid composition and membrane fluidity of tonoplast of woody plants and their relationship under salt stress.

全文：! !"" ! ##
$ % % &$ # %
& ’ ( )
’()*+,)- ’)./-* ’)+)(-*
/012!"*+02#
-345*
!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
$ % % &
+6(1[\8’mz{A%y6~6Br="
1 |7-$ RQ}$ O~!$ "uE$ #$%$ & Y8 ’()7
A72 YR&’T)W&lmi89)F YR $7%%89S$2 acE&’()DEF YR $777"8S
82 YRZ’T)6z())F6z()NOB7 YR $7%%&"Y
! "# e’mz{u&!zk&!uV4*UDno}G +6(1[\8A%y :; <-,=6>?1}<5A%yL{9
&*zy1rL<~!"#$%Z>}G +6(1[\8*u&!BA%y :; <-,=6>?1}<)GDg*h[\}G
u 7"% @@01A.A 7k*}<,)G85*ABC{UU<~Zk&!A%y :; <-,=6>?1}>*j}G +6(1A7"% @@01A.A 7Ye~$,)G|j~A%yL{9+|&*u (7B%%*(7B%7*(7#%%*(7#%7*
(7#%$z ($%%%~R[\$’mz{A%yL{9&?t{[K:6JB~u&!A%yL{9n0zG85*
7"% @@01A.A 7 +6(1e~$)G|jSZZk&!B*A%yL{9n0zGZ "% @@01A.A 7 +6(1e~$85*KO
R[\}G1Dg*n0zG,j,{U~+6(1[\8*’mA%yrLL{9n0zG85k*yrL<85*>,[yrL?
}<5L{9&*zy1rL$%&% ’mSR[\SA%ySL{9SyrL?
’()*+#’97#2!8 -./01#- -23+#7%%7 A 9!##4$%%&5%# A %%"% A %B
C?D?EF?G 0H I6JDK 7$*$%%& 5
L03HG6ME0H NJ0O?DM%,KE> J?>?6JDK P6> >3NN0JM?G QR +6ME0H61 ’DE?HD? S ,?DKH0104R =E116J =J04J6@ EH MK? *1?F?HMK LEF?$%%BT-U%8-%"%9Y5
(0JJ?>N0HGEH4 63MK0J%L6H4 V6H@EH45
!"#$%&’&(%)*’ *+, -%&)"./%)*’ 0.$1&+$. &2 3&+&1’*$4 5.$%)’.$ 6$&’*4.,
27&/ !"#$%%#&’()* +*+,")-’"* 4& 8*9’ :47.$$
WK6H4 IEH7*$ .E C0H4OEH$ :36H4 .EQEH$ XE V0H4K36$ U03 Y36HZEH$ YE6H I?H48 L6H4 V6H@EH47
A72 !"##$%$ "& ’"($)*(+ ,$)"-(.$) /01 2034("05$0* 6/0740% ’"($)*(+ 8043$()4*+ 6/0740% $7%%89S
$2 94/0%)- :./1$5+ "& ’"($)*(+ 6/0740% $777"8S82 ;4&$ <.4$0.$ ;/="(/*"(+ !$0*$(*6/0740% :%(4.-#*-(/# 8043$()4*+ 6/0740% $7%%&"Y
;<$47*)4% ,K? 6DMEFEMR 0[ :; <-,=6>?*[6MMR 6DEG D0@N0>EME0H 6HG @?@QJ6H? [13EGEMR 0[ M0H0N16>M F?>ED1?> E>016M?G [J0@ J00M> 6HG
1?6F?> 0[ >("-))"0$*4/ ?/?+(4&$(/ N16HM1?M> 3HG?J >61M >MJ?>> PEMK GE[[?J?HM D0HD?HMJ6ME0H> 0[ +6(1 P?J? EHF?>ME46M?G5 ,K? J?>31M>
>K0P?G MK6M /<:; <-,=6>? 6DMEFEMR EH J00M> EHDJ?6>?G 6M 10P 1?F?1 0[ +6(1 D0HD?HMJ6ME0HA"% @@01A.A 7Y5 \K?H +6(1
D0HD?HMJ6ME0H ?]D??G> 7%% @@01A.A 7*MK? 6DMEFEMR P6> >E@E16J PEMK D0HMJ01 5 )H 1?6F?>*/<:; <-,=6>? 6DMEFEMR G?DJ?6>?G 6M 7%%
@@01A.A 7 +6(1*Q3M EHDJ?6>?G 6M KE4K?J 1?F?1> 0[ +6(1A7"% @@01A.A 7Y5 )M P6> [03HG MK6M MK? [6MMR 6DEG D0@N0>EME0H 0[
M0H0N16>M P6> D0H>E>M?G 0[ (7B%%*(7B%7*(7#%%*(7#%7*(7#%$ 6HG ($%%% 5 -1M?J6ME0H> EH J?16MEF? D0HM?HM 0[ M0H0N16>M [6MMR
6DEG D0@N0>EME0H 3HG?J +6(1 >MJ?>> P?J? 0Q>?JF?G5 )H J00M> MK? ?]M?HM 0[ 3H>6M3J6ME0H 0[ M0H0N16>M [6MMR 6DEG> G?DJ?6>?G*6HG MK?H
EM EHDJ?6>?G 3HG?J 7"% @@01A.A 7 +6(1 5 )H 1?6F?>*MK? ?]M?HM 0[ 3H>6M3J6ME0H 0[ M0H0N16>M [6MMR 6DEG> G?DJ?6>?G 3HG?J
"% @@01A.A 7 +6(1 MJ?6M@?HM*K0P?F?J MK? ?]M?HM EHDJ?6>?61 PEMK MK? EHDJ?6>? 0[ >61EHEMR*1?6F?> 6HG P6> KE4K?J MK6H MK6M EH
D0HMJ01 5 ,K? DK6H4?> EH M0H0N16>M [13EGEMR D0EHDEG?G PEMK MK0>? EH 3H>6M3J6ME0H 0[ [6MMR 6DEG> 3HG?J +6(1 >MJ?>> 5 ,0H0N16>M [13EGEMR
G?DJ?6>?G PEMK MK? G?D1EH? 0[ 3H>6M3J6ME0H 0[ [6MMR 6DEG> 5 ,K? 0QO?DMEF?> 0[ MKE> >M3GR P?J? M0 46EH Q?MM?J 3HG?J>M6HG 0[ MK?
6G6NMEF? @?DK6HE>@ 0[ :; <-,=6>?*[6MMR 6DEG D0@N0>EME0H 6HG @?@QJ6H? [13EGEMR 0[ M0H0N16>M 0[ P00GR N16HM> 6HG MK?EJ
J?16ME0H>KEN 3HG?J >61M >MJ?>> 5
=.# >&7,$% >("-))"0$*4/ ?/?+(4&$(/S>61M >MJ?>>SM0H0N16>MS[6MMR 6DEGS@?@QJ6H? [13EGEMRS:; <-,=6>?
’61EHEMR E> 0H? 0[ MK? @6O0J 6QE0MED >MJ?>>?>*PKEDK
K6> 6 @6O0J E@N6DM 0H N16HM NJ0G3DME0H 6HG NJ0G3DMEFEMR 5
*]D?>> 6@03HM 0[ >61M EH MK? >0E1 6GF?J>?1R 6[[?DM> N16HM
4J0PMK 6HG G?F?10N@?HMA’6EJ6@ $* /# 5*$%%!Y5 )M E> H0P
4?H?J611R 6DD?NM?G MK6M MK? D?11 @?@QJ6H? E> MK? NJE@6JR
>EM? 0[ EHO3JR PK?H N16HM> 6J? >3QO?DM?G M0 >61M >MJ?>>
A’3JO3> $* /# 5*7&&BY5 I?@QJ6H?> N16R 6H E@N0JM6HM J01?
EH @6HR D?11316J 6DMEFEME?> EHD13GEH4 E0H MJ6H>N0JM*NJ0M0H<
!"#!$%& ’()*+,*+$&%*- ./*%+0"1.$2%*,.1A3"*%&*4556Y7
8. $+ %2. "%.$- /,1,%.-9 .:*. .:, +$&%$;$1*%1, 2; #,#-$!$0+ $% !-*%. +*-$%$.9 .2-,/*%1, :*+ <,,% *!!/,1$*.,07 8%
.:, !/,=$2"+ +."0$,+ +2#, /,+,*/1:,/+ :*=, $%=,+.$&*.,0
.:, 1:*%&,+ $% ;*..9 *1$0+*!:2+!:2-$!$0+ *%0 +.,/2-+
A>"/?"+ !" #$ 7*4556S@" !" #$ 7*ABBCSD$*%& !" #$ 7*
ABBCY$% 2/0,/ .2 ,+.*<-$+: * /,-*.$2%+:$! <,.E,,% -$!$0
*-.,/*.$2% *%0 +*-. +./,++ $% !-*%.+ 7 32E,=,/*.:, /,+"-.+
*/, 2;.,% 12%./2=,/+$*-A>"/?"+ !" #$ 7*4556Y7
(:, *<$-$.9 2; !-*%.+ .2 &/2E $% :$&: F*G-
12%1,%./*.$2%+ $+ *++21$*.,0 E$.: .:, *<$-$.9 2; .:, !-*%.+
.2 ./*%+!2/.*12#!*/.#,%.*-$H, 2/ ,I./"0, F*JA’!+, !"
#$ 7*ABBKY7 (:, !"#!$%& 2; F*J $%.2 .:, =*1"2-, $+
1*.*-9H,0 <9 =*+1"-*/ F*J L3J *%.$!2/.,/A(,+.,/ !" #$ 7*
ABBMY7 3J N,-,1./21:,#$1*- !2.,%.$*- &/*0$,%.*E:$1:
!/2=$0,+ .:, 0/$=$%& ;2/1,*E*+ $%$.$*--9 ,+.*<-$+:,0 <9 .:,
3J N’()*+, *%0 3J N!9/2!:2+!:*.*+, 2% .:, .2%2!-*+.
A(,+.,/ !" #$ 7*ABBMY7 O*1"2-*/ 3J N’()*+,+AON 3J N
’()*+,Y*/, #"-.$N+"<"%$. ,%H9#,+ .:*. !"#! !/2.2%+
$%.2 $%./*1,--"-*/ 12#!*/.#,%.+AG2P,/ !" #$ 7*ABBMY7 ON
3J N’()*+, !-*9+ $#!2/.*%. /2-,+ $% !3 :2#,2+.*+$+ *%0
=*/$2"+ +./,++ /,+!2%+,+AG2P,/ !" #$ 7*ABBMS>H, !" #$ 7*
ABBAY7 (:, ON3J N’()*+, :*+ <,,% /,!2/.,0 .2 <,
$%=2-=,0 $% +*-. .2-,/*%1, $% .:, !/,=$2"+ +."0$,+AD$*%& !"
#$ 7*ABBCSQ:*%& !" #$ 7*ABBASR" !" #$ 7*ABBCY7
S,#1/"1$*- $% !-*%. *0*!.*.$2% .2 *<$2.$1 +./,++,+AD$*%& !"
#$ 7*ABBCY7 (:, #*$%.*$%+ 2; #,#!/,/,T"$+$., ;2/ .:, ;"%1.$2%*=$*<$-$.9*&/2E.: *%0
/,!/20"1.$2% 2; 1,--+AD" !" #$ 7*ABB6Y7 G:*%&,+ $%
#,#,%=$/2%#,%. +"//2"%0$%& !/2.,$%+*E:$1: $% ."/% $%;-",%1,+
#,#*%0 .:, *1.$=$.9 2; #,#’()*+, *1.$=$.9A>"/?"+ !" #$ 7*4556Y7
U%.$- %2E*.:,/, $+ ;,E +."0$,+ :*=, <,,% 1*//$,0 2".
.2 *%*-9H, .:, 1:*%&,+ 2; -$!$0 12#!2+$.$2% *%0 #,#;-"$0$.9 2; E2209 !-*%.+ "%0,/ +*-. +./,++ 7 8% .:$+ +."09*
.:, .2%2!-*+. =,+$1-,+ ;/2# .:, -,*=,+ *%0 /22.+ 2;
%&’())’*!"+# ,#,-&+.!&# &/,E "%0,/ F*G- +./,++ E,/,
$+2-*.,0 *%0 .:, ,;;,1. 2; F*G- 2% .2%2!-*+. 3J N’()*+,
*1.$=$.9 *%0 #,#$%=,+.$&*.,07 (:, #,#E*+ *-+2 +."0$,07 (:, *$# 2; 2"/ +."09 $+ .2 &*$% <,..,/
"%0,/+.*%0$%& 2; .:, #,1:*%$+#+ 2; +*-. .2-,/*%1, $%
E2209 !-*%. 7
! "#$%&’#() #*+ ,%$-.+)
!/! 0(#*$ ,#$%&’#(
% 7 ,#,-&+.!&# "+,0 $% .:$+ +."09 E*+ $%./20"1,0
;/2# V*!*%,+ *%0 E*+ 1"-.$=*.,0 $% .:, %"/+,9 2; V$*%&+"
’1*0,#9 2; W2/,+./9 7 U%$;2/#N+$H,0 % 7 ,#,-&+.!&# +*
/+"&’ /,&,%,/*.,0 /22.$%& !-*%.-,.+ E,/, !-*%.,0 $%.2 !-*+.$1
!2.+ ;$--,0 E$.: CBB #D 2; :*-;N+./,%&.: S> +2-".$2%+ $% *
&/2E.: 1:*#<,/ 7 (:, %"./$,%. +2-".$2% E*+ /,%,E,0 ,=,/9
2.:,/ 0*97 (:, ,I!,/$#,%.*- 0,+$&% 12%+$+.,0 2; * 12%./2-
A %2 F*G-Y *%0 .:/,, ./,*.#,%.+ A CB*4BB *%0
4CB ##2-ADX 4 F*G-Y *%0 E*+ ,+.*<-$+:,0 $% *
/*%02#$H,0 0,+$&% E$.: .:/,, /,!-$1*.,+ 7 ’;.,/ ;$=, 0*9+
./,*.#,%. .:, /22.+ *%0 -,*=,+ E,/, :*/=,+.,0 ;2/ $+2-*.$2%
2; .2%2!-*+. =,+$1-,+ 7
!/1 2).(#$’.* .3 $.*.4(#)$5%*&’6-%+ 7%)’6(%)
(2%2!-*+. =,+$1-,+ E,/, $+2-*.,0 *112/0$%& .2 .:,
!/,=$2"+-9 /,!2/.,0 #,.:20AD$*%& !" #$ 7*ABBCYE$.:
#$%2/ #20$;$1*.$2%+7 Y22.+ *%0 -,*=,+ E,/, :2#2&,%$H,0
E$.: *% $1,N12-0 #2/.*/ *%0 !,+.-, $% .:, :2#2&,%$H*.$2%
<";;,/ACB ##2-ADX4 3Z)Z>N(/$+*!3 K[6*ACB ##2-ADX4
+2/<$.2-* 4AC ##2-ADX 4 \G-* C ##2-ADX 4 Z](’*
A[C ##2-ADX 4 \A>A C^*A ##2-ADX 4 )S>W*4[C_ )O)*
B[4_ ‘>’*A ##2-ADX 4 a((Y7 (:, :2#2&,%*.,+ E,/,
;$-.,/,0 .:/2"&: ;2"/ -*9,/+ 2; 1:,,+,1-2.: *%0 +"0$;;,/,%.$*- 1,%./$;"&*.$2% *. 4B BBB b & ;2/ 4C #$% *%0
+"<+,T",%.-9 *. CB BBB b & ;2/ MB #$% *. c d 7 (:,
4B BBB b & .2 CB BBB b & !,--,.+ E,/, +"+!,%0,0 $% 6 #D
2; <";;,/ ’AMBB ##2-ADX 4 +"1/2+,*4B ##2-ADX 4 \G-*
4 ##2-ADX 4 Z](’**%0 A ##2-ADX 4 a((*!3 K[eY7
(:,% c #D 2; <";;,/ ‘ A ACB ##2-ADX 4 +2/<$.2-*
4 ##2-ADX 4 Z](’*C ##2-ADX 4 3Z)Z>*!3 K[MYE,/,
-*9,/,0 2=,/ .:, <";;,/ ’ &/*0$,%. 7 (:, &/*0$,%.+ E,/,
.:,% +"c dY E$.: * ($5B /2.2/ $% *% ^!.$#* DNeBf)
"-./*1,%./$;"&,A‘,1P#*% G2"-.,/ 8%1 7*W"--,/.2%*G’*
U>’Y7 O,+$1-,+ <*%0$%& *. .:, <";;,/ ’L‘ $%.,/;*1, E,/,
12--,1.,0 *%0 +.2/,0 *. X eB d "%.$- "+, 7
!/8 9%$%&,’*#$’.* .3 :; 5<=0#)% #6$’7’$> ’*
$.*.4(#)$
’1.$=$.9 2; ON3J N’()*+, E*+ #,*+"/,0 *112/0$%& .2
>:$ !" #$ 7AABBKYE$.: #$%2/ #20$;$1*.$2% 7 (:, /,*1.$2%
E*+ 1*//$,0 2". $% * B[C #D #$I."/,AMB ##2-ADX 4
3Z)Z>N(/$+*!3K[C*M ##2-ADX 4 S&> c^*4BB ##2-ADX 4
4C! e# ^ ?=%F*G-[\8’mz{A%y6~6Br=
!"#*$%& ’’(#A)* + ,-,.S$%+ ’’(#A)* + ,-./01*
$%+ ’’(#A)* +A,21Y13(01*$%$45 6789(: ;<+$$*
. ’’(#A)* + =6>,-4Y? @A-B98(: C-D D9-79AE FG -EE8:H
&$!) 9(:(I#-D9 JAD8B#AD ? 6KA 7A-B98(: C-D 9A7’8:-9AE FG
9KA -EE898(: (L $%+ ’) (L +$5 MNM -L9A7 8:BOF-98(: -9
.P Q L(7 .$ ’8:? 6KA 7A#A-DA (L >8 L7(’ =6> C-D
EA9A7’8:AE FG 9KA R8DSA+T4&Y*OD8:H ,-24>01 -D 9KA D9-:E-7E?
!"# $%&%’()*+ ,)++- )./0 )&)(-*/*
R-99G -B8ED (L 9(:(I#-D9 JAD8B#AD CA7A -:-#GUAE OD8:H
9KA ’A9K(E EADB78FAE FG )8-:H !" #$ ?A4$$&YC89K D(’A
’(E8L8B-98(:D ? R-99G -B8ED CA7A ’A9KG#-9AE C89K F(7(:
978L#O(78EA+$!H (L KAI9-EAB-:(8B -B8E DA7JAE -D 8:9A7:-# D9-:E-7E?
6KA L-99G -B8E ’A9KG# AD9A7D CA7A 9KA: -:-#GUAE FG 6KA7’(
67-BA X" Y#97- H-D BK7(’-9(H7-IK AZO8IAE C89K - L#-’A
8(:8U-98(: EA9AB9(7AR[NY-:E -: MI<4&\$ ZO-79U B-I8##-7G
B(#O’:A+$$ ’ ] $%4& ’’ ] $%4!’Y? 28HK#G IO7A ,4
C-D ODAE -D 9KA B-778A7 H-D ? 6KA 8:^AB98(: -:E EA9AB98(:
9A’IA7-9O7A CA7A F(9K DA9 -9 44$ Q ? 6KA L(##(C8:H
9A’IA7-9O7A I7(H7-’ C-D ODAE%6KA 8:898-98(: 9A’IA7-9O7A
C-D DA9 -9 +1$ Q -:E ’-8:9-8:AE L(7 & ’8:*L(##(CAE FG
1 QA’8:* + 9( 44$ Q L(7 4& ’8:? _O-:989-98(: C-D
-BK8AJAE FG :(7’-#8U-98(: C89K -: 8:9A7:-# D9-:E-7E (L
KAI9-EAB-:(8B ’A9KG# AD9A7 ? 6KA L-99G -B8E B(’I(D898(:
C-D A‘I7ADDAE -D ’(# 5 ?
!"1 23+345/&)+/%& %, +%&%’()*+ ,(6/0/+-
N>2A+*\- L#O(7ADBA:9 I7(FA 9( EA9A7’8:A 9KA L#O8E89G (L 9(:(I#-D9
JAD8B#AD 8: 9K8D D9OEG? = D9(BS D(#O98(: (L N>2
A4 ’’(#A)* +Y C-D (F9-8:AE FG E8DD(#J8:H N>2 8:
9A97-KGE7(LO7-:? R(7 #AFA#8:H (L 9(:(I#-D9 JAD8B#AD*N>2
C-D ODAE -9 - B(:BA:97-98(: (L 4!’(#A)
* + ? 6KA ’8‘9O7A
C-D 8:BOF-9AE -9 .P Q L(7 + K 8: 9KA E-7S? 6KA
L#O(7ADBA:BA I(#-78U-98(: (L N>2 C-D EA9A7’8:AE (: -
289-BK8 a&$ L#O(7ADBA:BA DIAB97(’A9A7 L899AE C89K -
I(#-78U-98(: -99-BK’A:9 -D EADB78FAE FG >-:H !" #$ ?
A4$$&Y? 6KA D-’I#AD CA7A A‘B89AE -9 .\$ :’*-:E 9KA
A’8DD8(:D -9 14T :’ CA7A 7AB(7EAE? b(9K A‘B89-98(: -:E
A’8DD8(: D#89D CA7A DA9 -9 & :’? 6KA EAH7AA (L
L#O(7ADBA:BA I(#-78U-98(:A&YC-D B-#BO#-9AE -BB(7E8:H 9(
9KA L(##(C8:H L(7’O#-%& cA ’ VV * ’!YVA ’ VV d X[!Y*8:
CK8BK ’ VV -:E ’! -7A 9KA L#O(7ADBA:BA 8:9A:D898AD
’A-DO7AE C89K I-7-##A# -:E IA7IA:E8BO#-7 (78A:9AE
I(#-78UA7D*7ADIAB98JA#G*-:E ( 8D 9KA B-#8F7-98(: L-B9(7 ?
2A7A*( c ’! V ’ VV ?
!"7 8+)+/*+/.)( )&)(-*/*
M9-98D98B-# -:-#GD8D C-D B-778AE (O9 FG (:A=,0/= OD8:H M>MM +$%$ D(L9C-7A 9( EA9A7’8:A 9KA
E8LLA7A:9 D8H:8L8B-:BA? eKA: 9KA =,0/= C-D D8H:8L8B-:9
-9 & f $%$&*9KA NO:B-: ’O#98I#A 7-:HA 9AD9 C-D ODAE L(7
’A-: B(’I-78D(:? N-9- I7ADA:9AE CA7A ’A-: g MN (L
9K7AA A‘IA78’A:9D ?
9 :3*6(+*
9"! ;,,3.+ %, <)=( *+43** %& >?@A ?B$C)*3 ).+/D/+-
,4%5 4%%+* )&0 (3)D3* %, ! E "#"$%&’(%#
6( -DDADD 9KA 8:L#OA:BA (L ,-"# (: 9KA -B98J89G /<
2d <=6>-DA*) ? *#*+,-.!,# CA7A 97A-9AE C89K E8LLA7A:9
B(:BA:97-98(:D (L ,-"# -:E 9(:(I#-D9 JAD8B#AD CA7A 8D(#-9AE
L7(’ 9KA 7((9D -:E #A-JAD ? 6KA 7ADO#9D DK(CAE 9K-9 9KA
-B98J89G (L /<2d <=6>-DA 8: 9KA 7((9D 8:B7A-DAE C89K 9KA
8:B7A-DA (L ,-"# B(:BA:97-98(: ? =:E 9KA: /<2d <=6>-DA
-B98J89G EAB7A-DAE CKA: ,-"# B(:BA:97-98(: 7A-BKAE
+&$ ’’(#A)* +*FO9 D8’8#-7 C89K 9KA B(:97(#AR8H ? +Y?
2(CAJA7*9KA BK-:HAD (L /<2d <=6>-DA -B98J89G 8: 9KA
#A-JAD CA7A :(9 B(:D8D9A:9 C89K 9KA 7((9D ? [: #A-JAD*/<
2d <=6>-DA K-E :( D8H:8L8B-:9 E8LLA7A:BA 9( B(:97(# CKA:
97A-9AE C89K #(C ,-"# B(:BA:97-98(: ? =9 D-#8:89G (L
+$$ ’’(#A)* + 9KA -B98J89G EAB7A-DAE D8H:8L8B-:9#G -:E
7A-BKAE 9KA #(CAD9 #AJA# ? =L9A7 9KA:*9KA -B98J89G 7AB(JA7AE
-9 K8HKAD9 B(:BA:97-98(: (L ,-"#A+&$ ’’(#A)* +Y-:E 89
-#’(D9 B#(DA 9( 9KA #AJA# -D B(:97(# ?
R8H? + "K-:HAD 8: /<2d <=6>-DA -B98J89G L7(’ 7((9D
-:E #A-JAD (L ),/011/2!"-# *#*+,-.!,#
h-BK ’A-DO7A’A:9 C-D B(:EOB9AE C89K 9K7AA 7AI#8B-9AD*-:E E-9- CA7A
’A-: g MN? N8LLA7A:9 #A99A7D 8:E8B-9A D8H:8L8B-:9 E8LLA7A:BA FG NO:B-:iD
’O#98I#A 7-:HA 9AD9 -9 $%$& #AJA# ?
9"9 =F)&G3* /& ,)++- )./0 .%5’%*/+/%& %, +%&%’()*+
6KA -:-#GD8D (L L-99G -B8E B(’I(D898(: (L 9KA 9(:(I#-D9
4& & ’ ( ) 1&"
!"#$%&"# &$’$( ")*+,%* +"!",&"( *-,* *-" .,**/ ,%$(# $0
*101’&,#* 1. ! 2 "#"$%&’(%# +11*# ,0( &",!"# 3,$0&/
%10#$#*"( 1. 456%7*456%5*458%7*458%5*458%9 ,0(
497%7*:-"+",# 458%; :,# .1<0( $0 *-" +11*# +,*-"+ *-,0
$0 *-" &",!"#A=,>2 5 ,0( 9Y2 =-" +,*$1 1. <0#,*<+,*"(
.,**/ ,%$(# *1 #,*<+,*"( .,**/ ,%$(#A?@AY("%+",#"( $0 *-"
*101’&,#*# 1. +11*# ,0( &",!"# ,* &1:"+ &"!"&# 1. B,4& 2
C-$&" *-" ?@A 1. +11*# $0%+",#"( ,0( :,# #&$D-*&/ -$D-"+
*-,0 %10*+1& <0("+ 5E7 331&AFG 5 B,4& 2 H0 &",!"#**-"
?@A $0%+",#"( D+,(<,&&/ ,* 577 ,0( 5E7 331&AFG 5 B,4&
*+",*3"0*A=,>2 5Y2 I1+"1!"+**-" $0(") 1. <0#,*<+,*"(
.,**/ ,%$(#A H?JKY("%+",#"( $0 *-" +11*# ,.*"+ B,4&
*+",*3"02 H0 %10*+,#* *-" H?JK ("%+",#"( $0 *-" &",!"#
<0("+ E7 331&AFG 5 B,4&*,0( *-"0 $0%+",#"( <0("+ 577
,0( 5E7 331&AFG 5 B,4&A=,>2 9Y2
!"#$% &’’()* +’ ,"-. +/ ’"**0 ")12 )+34+51*1+/ +’ *+/+4."5* 15+."*(2 ’6+3 6++*5 +’ ! $ "#"$%&’(%#!
J,**/ ,%$( %13’1#$*$10A31& LY B,4&@
A331&AFG 5Y
7 E7 577 5E7
M,&3$*$% ,%$(A456%7Y
M,&3$*1&"$% ,%$(A456%5Y
A*",+$% ,%$(A458%7Y
N&,$($% ,%$(A458%50O*Y
P&"$% ,%$(A458%50O%Y
F$01&"&,$($% ,%$(A458%906*Y
F$01&"$% ,%$(A458%906%Y
K+,%-$($% ,%$(A497%7Y
F$01&"0$% ,%$(A458%;0;Y
?@A
H?JK
5OQ69 R 52;6
9;Q67 R 72ST
OQ77 R 72;;
TQ7E R 7267
7QE5 R 727T
;TQT7 R ;2;T
SQ59 R 725;
7QT7 R 725S
5Q9E R 7279
9QTT R 72;E
55TQOE R 7276
59Q9T R 7267
5TQTT R 72;9
T;Q77 R 72O;
EQ59 R 7289
;Q8O R 72TS
58QSO R 92TE
5QO6 R 72S9
7Q56 R 7278
7QT7 R 727T
7Q85 R 727S
66Q5E R 72T;
5TQ;7 R 727O
97Q77 R 728S
9;Q5S R 7289
TQE7 R 72S8
9Q69 R 7287
;;Q58 R T2T7
9Q9; R 72;T
3
3
5Q6S R 72TS
OSQOT R 72O7
57QO; R 7296
;TQ98 R 52O6
5EQE7 R 52T8
SQSS R 52E7
7QTE R 727E
9OQ85 R 5285
5Q9T R 727E
3
3
9QS8 R 72O5
57TQ65 R 52E;
! ?@A +,*$1 1. <0#,*<+,*"( .,**/ ,%$(# *1 #,*<+,*"( .,**/ ,%$(#YUAL *1*,& <0#,*<+,*"( .,**/ ,%$(#@ *1*,& #,*<+,*"( .,**/ ,%$(#YSH?JKA$0(") 1. <0#,*<+,*"(
.,**/ ,%$(#YUA5 V L 310"0"# W 9 V L ($"0"# W ; V L *+$"0"#YSQ3R*%,0 01* >" ("*"+3$0"( :$*- D,# %-+13,*1D+,’-2 =-" #,3" >"&1:2
!"#$7 &’’()* +’ ,"-. +/ ’"**0 ")12 )+34+51*1+/ +’ *+/+4."5* 15+."*(2 ’6+3 .("8(5 +’ ! $ "#"$%&’(%#
J,**/ ,%$( %13’1#$*$10A31& LY
B,4&@A331&AFG 5Y
7 E7 577 5E7
M,&3$*$% ,%$(A456%7Y
M,&3$*1&"$% ,%$(A456%5Y
A*",+$% ,%$(A458%7Y
N&,$($% ,%$(A458%50O*Y
P&"$% ,%$(A458%50O%Y
F$01&"&,$($% ,%$(A458%906*Y
F$01&"$% ,%$(A458%906%Y
K+,%-$($% ,%$(A497%7Y
?@A
H?JK
95Q;E R 52;7
5SQ58 R 726O
;7QTE R 52;9
56Q6E R 528T
;Q59 R 72E;
8QET R 729S
9Q;E R 727;
7Q;E R 7257
7QO9 R 727T
E8QS; R 92;9
59QES R 727S
OQT; R 72E;
E5Q;T R 92TS
58QEO R 927;
;QT5 R 727E
7Q96 R 727T
;Q65 R 72T5
7Q87 R 7276
7QEE R 727;
;OQ5S R 52ST
58Q6S R 527O
99QO5 R 52;7
56QO7 R 52E;
96QO6 R 9297
8QS7 R 72OT
;Q57 R 72;T
5Q7O R 7279
5Q66 R 729T
5Q6O R 727T
66QOE R 5286
5OQT9 R 7295
;EQ65 R 92ST
97Q8E R T275
TQS9 R 7276
5Q9E R 725E
5SQ5T R 72;9
7QE; R 7276
7QTS R 7259
5QTE R 7278
S6QO9 R 52;T
79: ,"-. *6("*3(/* );"/<(2 *;( ’.=121*0 +’ *+/+4."5*
8(51).(5
=-" *101’&,#* !"#$%&"# :"+" &,>"&"( :$*- *-"
.&<1+"#%"0* +",D"0* XMY *1 "!,&<,*" *-" %-,0D"# $0 *-"
.&<$($*/ 1. *101’&,#* !"#$%&"# <0("+ B,4& #*+"## 2 =-"
.&<1+"#%"0%" &,>"&"( *101’&,#* !"#$%&"# :"+" #%,00"( :$*-
, .&<1+"#%"0%" #’"%*+13"*"+*,0( *-" ’",Z !,&<" 1.
")%$*,*$10 ,0( "3$##$10 :,# .1<0( ,* ;67 ,0( T9O 03*
+"#’"%*$!"&/AJ$D 2 9 K ,0( [Y2 =-" 3"3>+,0" .&<$($*/
:,# ("*"%*"( >/ *-" #*",(/\#*,*" .&<1+"#%"0%" ’1&,+$],*$10
3",#<+"3"0*# 2 =-" +"#<&*# $0 =,>2; ("310#*+,*"( *-,* $0
*-" +11*# *-" ) !,&<" $0%+",#"( <0("+ E7 ,0(
!"#$: -;"/<(5 1/ *+/+4."5* ’.=121*0 +’ 6++*5 "/2 .("8(5
’6+3 ! $ "#"$%&’(%#
B,4&@
A331&AFG 5Y
I"3>+,0" .&<$($*/AMY
^11* F",.
7 7 2978 R 72776 7 297T R 72757
E7 729T7 R 72755 7 29;5 R 7275;
577 7299; R 7277S 7Q567 R 7275E
5E7 7297; R 7277S 7Q58; R 72755
577 331&AFG 5 B,4&*:-$%- $0($%,*"( *-" ("%+",#" $0
*101’&,#* .&<$($*/ 2 K0( *101’&,#* .&<$($*/ $0%+",#"( :-"0
B,4& %10%"0*+,*$10 +",%-"( 5E7 331&AFG 5 2 C-"+",#*$0
*-" &",!"# *101’&,#* .&<$($*/ ("%+",#"( <0("+
E7 331&AFG 5 B,4&*,0( *-"0 $* $0%+",#"( :$*- *-"
;E! 8# ^ ?=%B,4&[\8’mz{A%y6~6Br=
!"#$ % &’(")*)"+,A-Y*,. /0"11"+,A2Y13/()450 +6
675+4/1(/,(/ 7*8/7/. )+,+37*1) 9/1"(7/
",(4/*1/ +6 :*;7 (+,(/,)4*)"+, $
! "#$%&$$#’(
<=/ 65,.*0/,)*7 8*1"1 +6 )=/ *.*3)*)"+, +6 37*,)1 )+
1*7",")> 1)4/11 "1 )=/ (+,)4+7 +6 )4*,13+4) +6 1*7) *(4+11
0/084*,/1 $ 37*10*7/00* *,. )=/ )+,+37*1)**4/ 3*4)"(57*47> ",9+79/.
", )=/ 34+(/11 +6 (+03*4)0/,)*)"+, +6 )=/ :*;7 )*A/, 53
ABC))#/*DEEFY$ <=/ )+,+37*1)1 +6 37*,) (/771 (+,)*",
34+)/",1 (*)*7>G",# 34"0*4>H*,. 1/(+,.*4>H*()"9/ 34+(/11/1
+6 "+, )4*,13+4)*?="(= *4/ /11/,)"*7 ", )=/ 1*7)
(+03*4)0/,)*)"+, ",9+79/. ", *.*3)*)"+, )+ 1*7",")>
ABC))#/*DEEFY$ IHJK H-1 * 65,.*0/,)*7 4+7/
", /,/4#"G",# :*K MJK *,)"3+4) *()"9")> ", (/771
*((5057*)",# 1"#,"6"(*,) N5*,)")"/1 +6 :*;7AO=*,# !"
#$ $*%PP%Y$ Q) =*1 8//, 1=+?, )=*) )=/ *()"9")> +6 )=/ IH
JK H-64+0 8*47> 4++)1 5,./4 )4/*)0/,) ?")= :*;7AO=*,# !"
#$ $*%PP%Y$ Q, )="1 1)5.>*?/ 6+5,. )=*) *) 7+?
(+,(/,)4*)"+,1 +6 :*;7 )4/*)0/,) )=/ *()"9")> +6 IHJK H
-1)"057*)/.$ -,. )=/ *()"9")> ./(4/*1/. 17"#=)7> 5,./4
DRP 00+7ABS D :*;7*85) ,+) 1"#,"6"(*,) (+03*4/. ?")=
)=/ (+,)4+7 $ J+?/9/4*)=/ (=*,#/1 ", )=/ *()"9")> +6 IH
JK H-)=/ 4++)1 $ <=/ IHJK H- ", )=/ 7/*9/1
./(4/*1/. ", 4/13+,1/ )+ :*;7 1)4/11 $ <=/1/ ."66/4/,(/1
+81/49/. 0*> 8/ 8/(*51/ )=*) ."66/4/,)"*)/. (/771 ",
9*4"+51 37*,) )"115/1 *,. +4#*,1 0*> 65,()"+, ."66/4/,)7> ",
(++4.",*)"+, ?")= ,/"#=8+4",# (/771 )+ *(="/9/ 1*7",")>
)+7/4*,(/AT5 !" #$ $*%PPRY$
U/084*,/1 =*9/ 8//, 1=+?, )+ 37*> *, "03+4)*,)
4+7/ ", )=/ *8"7")> +6 37*,)1 )+ (+3/ ?")= 1*7",")>AU*,1+54
!" #$ $*%PPRY$ I*4"+51 A",.1 +6 /,9"4+,0/,)*7 1)4/11*
15(= *1 )/03/4*)54/ 1)4/11 *,. +10+)"( 1)4/11*(*51/
*7)/4*)"+,1 ", )=/ 3=>1"(*7 34+3/4)"/1 +6 )=/ 0/084*,/
7"3".1 ", 7"9",# (/771AB+1 !" #$ $*%PPVY$ <=/ 3=>1"(*7
1)*)/ +6 0/084*,/ 7"3".1 *71+ *()1 ."4/()7> )+ 4/#57*)/ )=/
*()"9")> +6 0/084*,/H8+5,. 34+)/",1*15(= *1 )=/ "+,
(=*,,/71AW5A=*4/9*DEEEY*1/,1+4 34+)/",1AW5#"54* !"
#$ $*DEEVY**,. 34+)+,H3503",# -DEEXY$ Y/)/40",",# =+? 1*7",")> *66/()1 )=/ 0/084*,/
(+03+1")"+, +6 )=/ ?++.> 37*,) % $ &#&’()*!(# ?"77
",(4/*1/ +54 5,./41)*,.",# +6 37*,) 1*7) )+7/4*,(/$ T/
+81/49/. )=*) ", )=/ 4++)1*:*;7 (+,(/,)4*)"+, *) ?="(=
JK H- ",(4/*1/. ?*1 "./,)"(*7 ?")= )=*) *)
?="(= )+,+37*1) 675".")> ./(4/*1/.$ <="1 3=/,+0/,+, ?*1
*71+ 4/3+4)/. ", )=/ )+,+37*1) "1+7*)/. 64+0 8*47/> 4++)1
5,./4 1*7",")>AO=*,# !" #$ $*%PP%Y$ <=/ 34/9"+51 1)5.>
=*1 15##/1)/. )=*) 34+3/4 675".")> +6 0/084*,/ ?*1
"03+4)*,) 6+4 +3)"0*7 1)45()54/ *,. ="#= *()"9")> +6 JK H
-(=*,#/1 ", )=/ *()"9")> +6 IHJK H-,+) (+,1"1)/,) ?")= )=/ 4++)1 $ IHJK H-
./(7",/. ?")= )=/ ",(4/*1/ ", :*;7 (+,(/,)4*)"+, $
<4*.")"+,*77>**7)/4*)"+,1 ", 6*))> *(". 5,1*)54*)"+,
./#4// *4/ 4/7*)/. )+ (=*,#/1 ", 0/084*,/ 675".")>**,. ")
"1 A,+?, )=*) * ./(4/*1/ ", 6*))> *(". 5,1*)54*)"+, 4/157)1
", * ./(4/*1/ ", 0/084*,/ 675".")>A:*9*4"HQGG+ !" #$ $*
%PPPSZ5*4)*((" !" #$ $*%PP%Y$ Q, +54 1)5.>*)=/
+81/49/. ./(7",/ ", )+,+37*1) 675".")> 0"#=) .5/ )+ )=/
./(4/*1/ ", )=/ 4*)"+ +6 5,1*)54*)/. 6*))> *(".1 )+ 1*)54*)/.
6*))> *(".1A[MWY$
Q, * 34/9"+51 4/9"/?*U5,,1ADEEFY=*1 (+,(75./.
)=*)Q-.9*,(/1 ", 1*7) )+7/4*,(/ *) )=/ 0+7/(57*4 7/9/7 ?"77
8/ ", 0*,"357*)",# )=/ /’34/11"+, *,. 1)45()54/ +6 34+)/",1
VR & ’ ( ) VR"
!"#! $%&!’%( !") !’#&*+%’! %, *#(! #$’%** -)-.’#&)*R/
0")’),%’)*1! 1* %, 2’)#! 1-+%’!#&! !% 1&3)*!12#!) !")
$"#&2)* %, !") 4567 5809#*) :&;)’ *#(1&1!<*)*+)$1#((< 1&
=%%;< +(#&!* /
>& $%&$(:*1%&*!") ;)!#1(); #&#(<*1* %, !") 1*%(#!);
!%&%+(#*! 3)*1$() $%-+%&)&! -#< +’%31;) 3#(:#.()
1&,%’-#!1%& =1!" ’)*+)$! !% !") +"<*1%(%21$#( ’)*+%&*)* %,
+(#&!* !% *#(! *!’)** /
!"#"$"%&"’
8+*) ? 9*@(:-=#’; A/ BCCD / E#7 !’#&*+%’! 1& +(#&!*/ FA@G H)!!)’*IJK%
BBLD M BBIL /
N%O)’ P G*P%&)* Q*Q#31)* A/ BCCR / >;)&!1,1$#!1%&*$%&*)’3#!1%&*#&; ’)(#!13)
)S+’)**1%& %, 45809#*) $QE8* 1& !%-#!% +(#&!*/ 9(#&! ?%()$:(#’
@1%(%2< T)+%’!)’*BK%KLI M KIJ /
F1*O) N 6*G:..#’%= U/ KVBI / 0") $%(%’1-)!’1$ ;)!)’-1&#!1%& %,
+"%*+"%’:*/ P%:’&#( %, @1%(%21$#( N")-1*!’<*WW%RDI M LCC/
6:#&2 N U/ KVVW / G#(!5*!’)** 1&;:$)* (1+1; ;)2’#;#!1%& #&; (1+1; +"#*)
!’#&*1!1%& 1& +(#*-# -)-.’#&) %, *%<.)#& +(#&!*/ 0#1=#&1#*LK%VW M
KCL /
H1#&2 U N*X"#&2 Y 6*N")& Z*!" #$ / BCCI / A,,)$!* %, *1(1$%& %& 67 5
809#*) #&; 67 589#*) #$!131!<*,#!!< #$1; $%-+%*1!1%& #&; ,(:1;1!< %,
!%&%+(#*! 3)*1$()* ,’%- ’%%!* %, *#(!5*!’)**); .#’()H/Y/ A&31’%&-)&!#( #&; AS+)’1-)&!#( @%!#&<*IR%BV M RD/
H%* Q 8*?:’#!# E/ BCCL / ?)-.’#&) ,(:1;1!< #&; 1!* ’%()* 1& !") +)’$)+!1%&
%, )&31’%&-)&!#( *12&#(*/ @1%$"1-1$# )! @1%+"<*1$# 8$!#*KWWW%KLB M
KID /
H: U E*Z1: Z U*Y#&2 U*!" #$ / BCCW / A,,)$!* %, +")((%;)&;’%& #&; 1!*
-#1& $%-+%&)&!* %& !") $)(( -)-.’#&) ,(:1;1!+#!"%+"1*1%(%2<*BB%KIW M KIV /
H[!!2) \/ KVVR / 9(#&! $)(( -)-.’#&)* #&; *#(1&1!<%*!’:$!:’#(*.1%$")-1$#(
#&; .1%+"<*1$#( $"#&2)*/ T @’#* F1*1%( 4)2*I%BKD M BBL/
?#&*%:’ ? ? F*G#(#-# ] 6 8*8(5?:!#=# ? ?*!" #$ / BCCB / A,,)$! %,
E#N( #&; +%(<#-1&)* %& +(#*-# -)-.’#&) (1+1;* %, =")#! ’%%!*/
@1%(%21# 9(#&!#’:-*LI%BRI M BRV/
?:&&* T/ KVVR / 9"<*1%(%21$#( +’%$)**)* (1-1!1&2 +(#&! 2’%=!" 1& *#(1&) *%1(*%
*%-) ;%2-#* #&; "<+%!")*)*/ 9(#&!*N)(( ^ A&31’%&-)&!*KW%KI M BL/
E#3#’15>__% F*Z:#’!#$$1 ? F*91&_1&% N*!" #$ / BCCC / 9’%!)1& ;<&#-1$* 1&
!"<(#O%1;* %, !") ;)*1$$#!1%&5!%()’#&! +(#&! @%)# "<2’%*$%+1$# ;:’1&2
;)"<;’#!1%& #&; ’)"<;’#!1%&/ 9(#&! 9"<*1%(%2<*KBL%KLBD M KLRW/
9#&2 U 6*X": 6*Y: 9*!" #$ / BCCI / 0") $"#’#$!)’1_#!1%& %, +(#*-#
-)-.’#&) N#B 7 5809#*) 1& ’1$" *+"1&2%-<)(1&5$"%()*!)’%( ;%-#1&*/
FA@G H)!!)’**IDV%BRVD M BLCR/
Z:#’!#$$1 ? F*‘(1a1b$ c*G!)3#&%31b$ @*!" #$ / BCCB / 9(#*-# -)-.’#&) (1+1;*
1& !") ’)*:’’)$!1%& +(#&! -#*&.(# /!’012# ,%((%=1&2 ;)"<;’#!1%& #&;
’)"<;’#!1%&/ P%:’&#( %, AS+)’1-)&!#( @%!#&<*IR%BKIV M BKWW/
G#1’#- T ]*0<#21 8/ BCCL / 9"<*1%(%2< #&; -%()$:(#’ .1%(%2< %, *#(1&1!<
*!’)** !%()’#&$) 1& +(#&!*/ N:’’)&! G$1)&$)*JW%LCD M LBK/
G"1 Z 6*Q1&2 F*Y#&2 d F*!" #$ / BCCD / AS%2)&%:* &1!’1$ %S1;) +’%!)$!
$:$:-.)’ ’%%!* #2#1&*! %S1;#!13) *!’)** 1&;:$); .< *#(! *!’)**/ 9(#&!
9"<*1%(%2< #&; @1%$")-1*!’<*LI%ILB M IIC /
G:21:’# 8*61’%O#=# ]*E#O#*"1-# ]*!" #$ / KVVL / G12&#(5*)&*1&2
-)$"#&1*-* %, !") +:!#!13) %*-%*)&*%’ ];+Q 1& 3/24!’1241# 2&$1 /
?%()$:(#’ ?1$’%.1%(%2<*KL%VBV M VRJ/
G:O"#’)3 G/ KVVV / ?)$"#&%*)&*1!13) $"#&&)(* 1& .#$!)’1# #* -)-.’#&)
!)&*1%& ’)+%’!)’*/ F8GA@ P%:’&#(*KR%GII M GWKAG:++( /Y/
G:’e:* 8*Q:’#&; ?/ KVVW / H1+1; $"#&2)* 1& *%<.)#& ’%%! -)-.’#&)* 1&
’)*+%&*) !% *#(! !’)#!-)&! / P%:’&#( %, AS+)’1-)&!#( @%!#&<*LD%KD M BR/
G_) 6*G$":-#$")’ ]*?[(()’ ? H*!" #$ / BCCB / 8 *1-+() &%-)&$(#!:’) ,%’ #
$%-+()S +’%!%& +:-+%5%6 2)&)* )&$%;) !") 3#$:%(#’ 67 5809#*)/
0’)&;* 1& 9(#&! G$1)&$)*D%KID M KWK /
0)*!)’ ?*Q#3)&+%’! T/ BCCR / E#7 !%()’#&$) #&; E#7 !’#&*+%’! 1& "12")’
+(#&!*/ 8&&#(* %, @%!#&<*VK%ICR M IBD/
Y: P H*G)(1*O#’ Q ?*‘#((#2")’ P H/ BCCI / 0") ’)*+%&*) %, +(#*-#
-)-.’#&) (1+1; $%-+%*1!1%& 1& $#((:* %, !") "#(%+"A9%#$)#)Y!% *#(1&1!< *!’)**/ 8-)’1$#& P%:’&#( %, @%!#&<*VB%JIB M
JIJ/
U: @ P*H#- 6 ?*G"#% ‘ 6*!" #$ / BCCI / A,,)$!* %, *#(1&1!< %& #$!131!1)* %,
67 5809#*)*67 599#*) #&; -)-.’#&) (1+1; $%-+%*1!1%& 1& +(#*-#
-)-.’#&) #&; !%&%+(#*! 3)*1$()* 1*%(#!); ,’%- *%<.)#&A 9$:21.! *#;
H/Y*));(1&2*/ P%:’&#( %, A&31’%&-)&!#( G$1)&$)**KD%BIV M BWB/
X"#&2 Y 6*H1: U H/ BCCB / T)(#!1%&*"1+ .)!=))& !%&%+(#*! 67 5809#*)
#$!131!<*1%& :+!#O) #&; N#($1:- 1& .#’()< ’%%!* :&;)’ E#N( *!’)**/ 8$!#
@%!#&1$# G1&1$#*LL%WWD M WDB/
!"#$% 789)
II! J# ^ ?=%E#N([\8’mz{A%y6~6Br=

Physiological and Biochemical Response of Tonoplast Vesicles Isolated from Broussonetia papyrifera to NaCl Stress

NaCl胁迫下构树幼苗液泡膜生理生化响应

相关文献