免费文献传递   相关文献

Genetic Variation of Natural Populations of Betula luminifera in Fujian and Its Relationship with the Habitat

福建光皮桦野生种群遗传变异及其与生境的关系


采用表型性状测定结合RAPD标记,研究福建境内8个光皮桦野生种群的遗传变异,探讨遗传变异与环境因子间的相关性。结果表明: 光皮桦野生种群在DNA和表型水平上的遗传变异基本一致,即约2/3的变异存在于种群内,1/3存在于种群间; 叶、果序和种子3类性状的变异系数(CV)分别为0.063 2,0.070 6和0.069 3,各类性状的形状指数(长宽比)的变异系数均比单个性状小; 种群平均多态位点百分比(PPL)、Neis基因多样性(h)和Shannon信息指数(I)分别为65.27%,0.246 1和0.362 8。相关分析表明:表型性状间及其与经纬度、年均温、年日照时数、海拔及有机质等环境因子间存在显著或极显著相关,与各环境因子相关的主要表型性状有叶长宽比、侧脉数、叶基角、种翅宽、种子千粒质量和果序长宽比等; PPL,h和I与年日照时数呈显著相关,与经纬度、年降水量呈负相关; 种群间的遗传距离与地理距离相关不显著,而与年均温分异显著相关。

Genetic variation was evaluated using phenotypic traits and RAPD markers in 8 natural populations of Betula luminifera from Fujian Province, China. The relationship between genetic variation and environmental factors was also examined. It was showed that the genetic variation of B. luminifera revealed by RAPD markers coincided well with those by phenotypic traits, and about 2/3 of the total genetic variation distributed within population while 1/3 among populations. The mean coefficient of variation (CV) of phenotypic traits of leaf, infructescence and seed was 0.063 2, 0.070 6 and 0.069 3, respectively. The CV of morphological index of leaf, infructescence and seed was much lower than those of other traits. The percentage of polymorphic loci (PPL), Neis gene diversity (h) and Shannon informative index (I) at mean population level were 65.27%, 0.246 1 and 0.362 8, respectively. The correlation analysis showed that there existed significant correlations between 15 phenotypic traits and environmental factors. The main phenotypic traits which were related with environmental factors, were leaf length/width, number of lateral vein, the angle of leaf base, seed wing width, weight of 1 000 seeds and infructescence length/width. The PPL, h and I were also significantly positively correlated with annual sunshine hours, negatively correlated with latitude/longitude and annual precipitation. The genetic distance between these populations (GD) was significantly correlated with their divergence of annual average temperature, however not with their geographic distance.


全 文 :! !"" ! ##
$ % % #$ # %
& ’ ( )
&’()*+(, &(-.,) &(*(’,)
./01!"**/1#
&234*
!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
$ % % #
XTH+"bH41$P=>&9LHK:T,
4\55Q$ m676 q185 m 95Q$ C:;5
A51 XT/&’()DEF X% 6"%%5$X$4 BC&’()DEFDEHF /W 5%%%#5X
64 XTH9[)H:())F X% 6"%%%7c
! "# KPoL0P$0mN 8,9:WX*DEXTK> ;hH+"bH41:$P=>*h[$P=>LJ
K5_<:ST0lmnopqH+"bH41d :*,xoL+t8:$P=>gbo‘*-u $<6:=>zd
B41>*5<6zdB41,-A’.c?Iw%1%=6 $*%1%7% =x%1%=# 6*Lr0P
:;PB-Ayn|c:=>,-u|rh0P{X41tu3I%S?@ g53i0A !cx
&ABCC/CZlB-A "c?Iw ="1$7D*%1$!= 5x%16=$ ;lST?@opqoL0P<&9L‘-*tu&Z2V2JK5_-*1gl*4jn*4_k=V9xn/yn|2X99-*! x " L$fb‘-{s2ST*L}STX41<:$P()LjI()ST!s2*ML$uF?>s2STl
$%&q H+"X$P=>XoL=>X8,9:XHK
’()*+#&75;1!=5&75;1" ,-./0#, ,12+#5%%5 E 7!;;3$%%#4%# E %%=% E %=
def#q$%%; E %7 E $6l
ghijqXT/&,4f(llTijAm&4$$%%=%kF "7!"#"$%& ’()%($%*# *+ ,($-)(. /*0-.($%*#1 *+ !"#$%& %$’()(*"+& %# 2-3%(# (#4
5$1 6".($%*#17%0 8%$7 $7" 9(:%$($
F>2 G>H>CI5*$ JKBCI 8KLAK6 -> MA>LA2C5 JKBCI G/CI5*$ GBCI M/CINK5
A5 4 #$%&’( )*’+,-. /0 #/1,231. #$4!/$ 6"%%5$X$ 4 51’+$’3, 6*!//7 /0 8!&(,2, )*’+,-. /0 #/1,231. 9,&%&(: 5%%%#5X
64 8/77,:, /0 ;&0, 6*&,(*,*#$%&’( ,12&3. #$4!/$ 6"%%%7c
;:1$)(&$q O2C2P>Q RBS>BP>/C NB@ 2RB0KBP2T K@>CI 3A2C/PU3>Q PSB>P@ BCT 8,9: VBSW2S@ >C ; CBPKSB0 3/3K0BP>/C@ /X 9,3$7’
7$-&(&0,1’ XS/V YKZ>BC 9S/R>CQ2*’A>CB 4 +A2 S20BP>/C@A>3 [2PN22C I2C2P>Q RBS>BP>/C BCT 2CR>S/CV2CPB0 XBQP/S@ NB@ B0@/ 2\BV>C2T4
(P NB@ @A/N2T PABP PA2 I2C2P>Q RBS>BP>/C /X 9 4 7$-&(&0,1’ S2R2B02T [U 8,9: VBSW2S@ Q/>CQ>T2T N200 N>PA PA/@2 [U 3A2C/PU3>Q
PSB>P@*BCT B[/KP $<6 /X PA2 P/PB0 I2C2P>Q RBS>BP>/C T>@PS>[KP2T N>PA>C 3/3K0BP>/C NA>02 5<6 BV/CI 3/3K0BP>/C@ 4 +A2 V2BC
Q/2XX>Q>2CP /X RBS>BP>/CA’.c/X 3A2C/PU3>Q PSB>P@ /X 02BX*>CXSKQP2@Q2CQ2 BCT @22T NB@ %1%=6 $*%1%7% = BCT %1%=# 6*
S2@32QP>R20U4 +A2 ’. /X V/S3A/0/I>QB0 >CT2\ /X 02BX*>CXSKQP2@Q2CQ2 BCT @22T NB@ VKQA 0/N2S PABC PA/@2 /X /PA2S PSB>P@ 4 +A2
32SQ2CPBI2 /X 3/0UV/S3A>Q 0/Q>A99-c**2>?@ I2C2 T>R2S@>PUA !cBCT &ABCC/C >CX/SVBP>R2 >CT2\A "cBP V2BC 3/3K0BP>/C 02R20
N2S2 ="1$7D*%1$!= 5 BCT %16=$ ;*S2@32QP>R20U 4 +A2 Q/SS20BP>/C BCB0U@>@ @A/N2T PABP PA2S2 2\>@P2T @>IC>X>QBCP Q/SS20BP>/C@
[2PN22C 5" 3A2C/PU3>Q PSB>P@ BCT 2CR>S/CV2CPB0 XBQP/S@ 4 +A2 VB>C 3A2C/PU3>Q PSB>P@ NA>QA N2S2 S20BP2T N>PA 2CR>S/CV2CPB0
XBQP/S@*N2S2 02BX 02CIPATPA*CKV[2S /X 0BP2SB0 R2>C*PA2 BCI02 /X 02BX [B@2*@22T N>CI N>TPA*N2>IAP /X 5 %%% @22T@ BCT
>CXSKQP2@Q2CQ2 02CIPATPA4 +A2 99-* ! BCT " N2S2 B0@/ @>IC>X>QBCP0U 3/@>P>R20U Q/SS20BP2T N>PA BCCKB0 @KC@A>C2 A/KS@*
C2IBP>R20U Q/SS20BP2T N>PA 0BP>PKT2<0/CI>PKT2 BCT BCCKB0 3S2Q>3>PBP>/C4 +A2 I2C2P>Q T>@PBCQ2 [2PN22C PA2@2 3/3K0BP>/C@AO:cNB@
@>IC>X>QBCP0U Q/SS20BP2T N>PA PA2>S T>R2SI2CQ2 /X BCCKB0 BR2SBI2 P2V32SBPKS2*A/N2R2S C/P N>PA PA2>S I2/ISB3A>Q T>@PBCQ2 4
<"= 8*)41q 9,3$7’ 7$-&(&0,1’XI2C2P>Q RBS>BP>/CX3A2C/PU3>Q RBS>BP>/CX8,9:XAB[>PBP
H+"A9,3$7’ 7$-&(&0,1’c*+TG1"*w&’
vB?",C:Wo4*x.?kBO-*npqr
YZL/[*9:?kBs-*B-[tu $"% ]
$ #%% V:>J=j*9Btu 7%% ] 5 $%% V$H+
":Fxxyz{&YpG0,1*O}HB-=&m
&&B;j*rHys*mf<*!*KVy>*PG
x*019[280*$o4%Z_x;v:ypo
H_EPK@4*o<$dBCZaj[xwV4
AUqT*$%%%cly#Y,*3)\H+"\]MN
R>!V*@VKIAP_‘9bH\]:N*gz
&?3w20&Y@0&*&++:ef*’BqI
] A^0Tv*!"""c*5lMN9bH4V\]-$
X"~Zlrs\H+":DE1%0*zfV&*1089xjNAP2a3*O}
A!""#cgB $%&’x ())$WX\XT/ **hH+
"^k1O ++hr‘:$P3i0mn?@X>o
K2A!"",cKP $%&’WX\ -h!wtuH+"
^k41$P=>mnDE*|\H+"bH41
oL=>Y&$P=>LHKIlbJKPoL0P$0mN $%&’WX*h[
XTK>H+"bH41:$P=>*Y&$P=
>LJK5_<:ST0*#ywH+"4V\]
:MN*$Pzp&NIAPJ\ZPZll
* KKLac
!"! 5QK>?k&G
Ki: ,hH+"bH41?I%BXTK>
:{t0uA&*c*0~{!A&!c*ogp3A&.c*p
g’vA&-c*FERkMN[A&+c*6t^b-RkMN[A&,clL
41z&:HK=9?-jI*z{x,15_Ao
*cl9B*+*fb‘-^L41Odjzzs\K_9[Z*
,1T)*T%*Z2VU9& 12v,">oK2
A!"",cac$0lH+"5UKK:K<&?zw
Xj7]:^k&*t%PRh41fL\ !"‘Y
8Ki*|^B[??’i41y{*hO-Z}*
Rh4112L’ */ h20"‘*"‘~/" 3Y8*V?IBL0i‘8Swa%12K<
5zgT:1qx20n/\LoL0P$0XK
69%:kEA0#A2*!""#cMzvjTUV*’4%J’l
9 ! ’ABChHfKhm
#$%& ! #’( ’$%)*$* +, - .)/0 1+12/$*)+34 +, ! & "#$%&%’()*
41;<
&;1<75=>;?
4;@
jS
A;B5=>;?
iV
A5=>=A4c
A;?E>=AFc
tu
F7DG5=>;?H
3
$}+
%??<57
1IDB>1>=5=>;?H
33
$uF
%??<57 3D5?
=D31DI5=J
fb‘-
)?D
L;T)
9;=57
1L;K1L5=DH
AEyMEN *c
T%
9;=57
?>=I;ED?H
AEyMEN *c
Z2V
OIE5?>B
35==DIH
AEyMEN *c
12
&* {t A>5?BLD?E !/P-"Q **#P/!Q #!/ * #-+ *RS" * ,!RSR "S.# *S+# .*S- +S#*
&! 0~ T;?E’5? !/P-/Q **+P",Q +"" * #-# *RS/ * ,+!S" "S.! *S/! .!S/ /S!/
&. og T; !#P"-Q **,P",Q +/" * #"! *,SR * +#-S" "S/- *S/* .RSR /S-*
&- pg V>?EU> !#P!+Q **+P"-Q #/" * +-- *,S" * +,,S" "S.. *S"# -+S" -SR/
&/ FE )L5U>5? !#P!,Q **+P-!Q +!/ * ##! *#S! * ,++S+ "S/" *S,- /!S. /S."
&# w6 )L5;W< !+P**Q **+P-.Q +!" * ,"! *+S+ * +"-S- "S., *S/. !!S/ /S"!
&+ 6y- XKL5? !+P/"Q **+P-/Q * *," * R"# *+SR * .""S" "S!! !S-/ ,.S/ -S/"
&, 6t X<1>?E !/P*.Q **#P*!Q ,/" * #!/ *RS/ * +/,S" "S/# !S"# #"S. +S!"
!"5 DEYl
*S!S* ^_‘2A*RR+cxZ,2
A!""/c:"7oL$0ac*P:a*9lr*?_
XW#a24r$91qy*1qn*es-*gn
(A1g+zn’()c*1ty*1gl2’(0P
xn/y*n/n*4_y*4_n*4jnx4_k
=V92He0PlRhr‘$0 !"qg?1q*
!"hmTn/x ."=4_l
*S!S! ’4%abc $%&’Ad H+"; ’4%J
’*67&L*$%&’:82t>oK2A!"",cac
mnl&L:PBbDE: *" Z1 1267A{R
8t )5?E;?yWc/Y:o !l
9 5 5!U=CKO6
#$%& 5 6(72(38(4 +, 5! (,,(8*)9( 1:);(:4
67;;
&I>3DI 4;@
/Y
)D[pv-
:5?C ?<3ZDI
67;;
&I>3DI 4;@
/Y
)D[pv-
:5?C ?<3ZDI
)!! 90660%0690 *! )*/! 99%9606666 ,
)!- %%96000690 ** )*/- 90600690%0 R
).* 6%%9606609 ** )*-"* 66096009%0 **
)-" 099060%966 ** )*-"! 00%%%66669 *!
)-/ 90%0600%6% ** )*-"/ 6660%%060% *-
)#, 900%660090 + )*-"# 090069900% **
)*"/ %096096666 *! )*-** 090606%%90 *"
)*"+ 6906%96090 ** )*-!" 69969600%6 **
)*!! 0%00%96669 *" )*-!+ 0900660%90 ,
)*-+ %0%906%066 ** )*-." %06%060%00 *.
)*/* 0%09696%00 ** ;_ 9;=57 !!/
*#! R# >oK2qXTH+"bH41$P=>&9LHK:T,
!"#"$ efDE !c;IBW-‘?@qP=>
,-A%&cx|"ar?@2acmnLoL0P
=>?@*PoL?},-A!’(c3441}UVAJ}2*!)**c*P ’+’ *"#=>mnCr?
@l#c,+-.-‘?@qS|~*x0Ug: ,+-.
pvmn !/0WX*ZvXw !*{vXw 0*TTj
!-‘e1lP -1-2343 !"$#=>_9L41:
3I%S’:;<<=-A$’(c2XP +>1&+=>A3?@=AA76B*!))CX>7DD6B*
!))Ec?@$P=>d41>*41<:?k*VP
(F-2+ !"$ =>A>;BG* !))EcmnC r ? @
AH-2>+cl$cST?@qP ’-’’ !$"0=>mno
L0P<*oL0PLJK5_&?_$P,-L
JK5_<:ST?@*9B$P()LjI()*
JK5_~<:ST?@,">oK2A#00*cl
# mnL?@
!"# ’ABChHf9bQRc;o $w *hH+"bH41LoL0P:=>
,-A%&clNoB^YZU*wo41>!woL
0PdW[r>l9B*=>UVz[:$4_k=V
9*9m$1ty*n/y*n/n*es-2*=>z
{:$n/*1x4_:;PB-Ayn|c*tp4
1>{J1*n/c$4_*9;PB-Wrh0P
x0X1*n/*4_ $ r0P:tu %& ?Iw
0"0I$ #*0"0E0 Ix0"0I) $*9B10P:tu=>,
-|n/x4_0P{*op10P|n/x4_0
P:x00AXN41+tZ*6t^b-RkMN
[41A-*c:=>z;z{Atu %&w0"0EC Ccl
9 $ ’ABChHfD9bSgKQR(%&’( $ )&*+&,+-. /-011+/+0.,233)4-1 450.-,64+/ ,*&+,2 -1 ! ( "#$%&%’()* +. 7 8+9: 4-4;9&,+-.2
0P (B;7J9 -! -# -$ -K -C -I -E -* ;O (=J;D
es- % DL 0"!K0 C 0"0)$ K 0"0*0 # 0"!#! 0 0"0$! # 0"0KC # 0"0#E 0 0"!CI ) 0"0EI K
1y & D 0"0E# # 0"0EI 0 0"0E! ! 0"0*0 E 0"0*I 0 0"0EK ! 0"0*K # 0"0I) ! 0"0$$ E
1n ’ D 0"!0# I 0"!0# $ 0"0*0 C 0"!0) ! 0"0)) * 0"0*E $ 0"0)$ # 0"!0! $ 0"0KK K
1yn| & D /’ D 0"0C$ ! 0"0I! # 0"0IK 0 0"0K) $ 0"0#C # 0"0I# ) 0"0#) ! 0"0$E E 0"0!E *
gn( ’M;? 0"#I$ # 0"#!! K 0"!0) ) 0"!KK ! 0"!0* * 0"!#$ 0 0"0)* C 0"0)* K 0"0#* C
1ty &N 0"$$! I 0"#0K ! 0"!)* E 0"#$! 0 0"!IC ! 0"!)* K 0"!IC $ 0"!IK C 0"!E) K
1gl ( DO 0"!)E 0 0"!$! C 0"!KK 0 0"!$# # 0"0$) # 0"!0) C 0"0C! K 0"!*E $ 0"0I# K
n/y & A 0"00) 0 0"0I) C 0"0#K ! 0"0EI * 0"0#) E 0"0!K $ 0"0#) 0 0"!)! C 0"!#0 E
n/n ’ A 0"0!# $ 0"0$I 0 0"0$C ) 0"0IC E 0"0KI C 0"0EI 0 0"0E* ) 0"!0! E 0"0EI K
n/yn| & A /’ A 0"00E E 0"0$! ! 0"00E * 0"0#K $ 0"00) I 0"00) K 0"0!! E 0"0CI C 0"0!K E
4_y &9 0"!## 0 0"!I$ 0 0"!EC # 0"!#! ) 0"0*K * 0"!)K # 0"!!E * 0"!#* I 0"0C$ K
4_n ’9 0"0I# I 0"!#0 ! 0"!0! $ 0"0I* C 0"0II ! 0"!0# * 0"!#! ! 0"0E! # 0"0KC $
4_yn| &9 /’9 0"0I* ! 0"0*) K 0"0)) C 0"0EK # 0"0*E 0 0"!0# E 0"!#C $ 0"0*) $ 0"0## I
4jn ’9P 0"!!* $ 0"!#E ) 0"!!* K 0"!!$ 0 0"0*) # 0"!K# 0 0"!!K ) 0"!!) C 0"0$C I
4_k=V9 )9 0"!I* E 0"$E) # 0"$EE E 0"0!E I 0"!IK ! 0"#EK ) 0"$0! * 0"$C) # 0"!*) K
tu >6;< 0"!!C $ 0"!#I K 0"!!# I 0"0)C $ 0"0EC C 0"!0E * 0"0)I I 0"!#* * 0"0I$ K
!% DLq4QMO6B =A D;J6B;D L67+D6|"ar?@mnAo Kcop*YQ: !Cho
L0Pd *hH+"bH41<*41>uzds
2Y%s2r>l1*n/*4_ !C hoL0P:
tuoL?},- !’( W 0"$I! EA=Ew0"$0I ! X
0"$)* Ic*-41<:tuoL=>*;=>:
$I"!EY*41>* I$"*$Y*tpH+"bH41>
=>$oL=>:1%,]l
NH+"bH41 !ChoL0P:ST?@
mnAo Cc^ YZU*’(0PB*1yL1n{s
2ST*L1gl{%s2STXes-L1gl*
Y&gn(L1ty{%s2STlHe0PB*
n/yLn/n{s2ST*L4_n{%s2S
TX4_yL4_n*4jnL4_k=V9u{
s2STldHe0PL’(0P<*1yn|L
4_yn|{%s2ST*es-Ln/y{s2
bST*gn(L4_y{s2}STl
*hH+"bH41: ) h1%oL0PB*
Z IhoL0PLHK5_ST*9Bes-*1g
lLiV{s2}STXes-LST*n/yn|L4_k=V9Ltu?I{s2}STx%s2b
STXes-*1glL$uFY&1yn|*4_
k=V9L$fb‘-u{s2STX1yn|L
Z2VU9{%s2}ST*es-*1glLNT
#I & ’ ( ) KC"
9 ! ’ABChHfD9bSgKFGY_)@\hk9b)x("#$% ! &’(()*)+,’#,’-+ .-)((’.’)+,/ #+0 1#*’#+.) #+#23/’/ -( 45)+-,34’. ,*#’,/ -( ! % "#$%&%’()* ’+ 6 7’20 4-482#,’-+/
0P
!"#$%&
uaAR^Vc’(A)*c !
41<
+,-./ 0-012#%$-.&
41>
3$%4$. 0-012#%$-.
2a
5#.)-, 6""-"&
41<
+,-./ 0-012#%$-.&
41>
3$%4$. 0-012#%$-.
oL?},-
7$**6"6.%$#%$-. 8-6**$8$6.%
-* 046.-%90$8 %"#$%&A"(!c
# 2: ;<=>;=? @A@c @>?A; @A;;Bc A>BA< C ;=>@=!! A>$ 2 ;?>;;A DA@c =>AB< BA;;Bc A>B=!! E>B;!! A>% 2 ;=>?EA EA@c ;>CDC DA;;Bc A>B;A @ ?>@C!! ?>@C!! A>$ 2 F% 2 A>??B =A@c A>AC< A ;;Bc A>ABA = ?>C;! <>B=!! A>%,#G =>DDB ?A@c B>;D< DA;;Bc A>;A; A <>DE!! C>E@!! A><@C =
$0 ;?>;@? DA@c ;>CB= ?A;;Bc A>A@E ? E>=D!! BB>?;!! A>& 2H ?EE>;DD @A@c ??< ;A;;Bc @>;DC @ B>@C!! A>$ * ;>D=D @A@c A>;D= =A;;Bc A>ADC ? E>@;!! <>?D!! A><@D E
% * B>CAC ;A@c A>;?? A ;;Bc A>A=A = ;<>?@!! D>EC!! A>$ * F% * A>?EA CA@c A>A?@ @A;;Bc A>A;B A ;A>@=!! E>@@! A>$& @>EC@ =A@c ;>;=B A ;;Bc A>A?E A <>CE! BB>@?!! A>%& <>EEB CA@c A>D;@ BA;;Bc A>ACC!! ;<>?E!! A>$& F%& A>DE@ DA@c A>A=? ;A;;Bc A>A;= = ;A>@?!! B*CD!! A><;? ?
%&I ;>CD< DA@c A>@;; A ;;Bc A>B?; C <>@?<== B
’& A>;B= @A@c A>ADD @A;;Bc A>A;; ; <>;;!! D><@!! A>!!*!!q?Io1d A>ADx A>A;+ts2l($/.$*$8#.% #% A>AD #.) A>A; 26:62&*"6&068%$:629J +wl!46 &#,6 H62-IJ
9 9 ’ABChHf9bSgPKU$S
"#$% 9 :-**)2#,’-+/ $),7))+ 45)+-,34’. ,*#’,/ -( ! % "#$%&%’()* ’+ 6 7’20 4-482#,’-+/
0P !"#$%& # 2: $ 2 % 2 $ 2 F% 2 %,#G $0 & 2H $ * % * $ * F% * $& %& $& F%& %&I
$ 2 K A>??@
% 2 A>DAA A>@C=!
$ 2 F% 2 A>BDB K A>?<@ K A>%,#G A>==D K A>B=C A>=B; A>=B=
$0 A>DA< K A><;@ A>BCE A>=B? A>C?@!!
& 2H A>E;E!! A>C@E!! A>;CA A>DBE A>DAC A>DED
$ * A>@<;! A>=A? K A>BB= A>B?= K A>

=A;
% * A><=C K A>AAB K A>AAD K A>D;@ A>;AC A>=B@ A><;? A>@D=!
$ * F% * A>AA? A>B;E A>;=B A>A;D A>BB? K A>;=C A>;?D A>AB; K A>B=A
$& K A>A=C A>AAA@ K A><@C K A>@<@! K A>?@E K A>;?? A>??B A><A;<
%& K A>?;E K A>=DA K A>;<< K A>;<@ K A>=BE A>ECA!! A>D?; K A>;;A A>@?E!
$& F%& K A>AB? K A>DE? K A>?A? A>C??!! A>AA;? A>BED K A>;B;= K A>AEA A>AA= K A>A?B
%&I K A>D;D A>A?@ K A>D;B A>BC; K A>;D< K A><B<; K A>;
AEE K A>;D?B
’& K A>?E= A>B=? K A>?BAB K A>?B@ K A>??E K A>D@E K A>;=@ A>D;A K A>;B< A>=?= K A>;DE A>B=@ A>@@D!
9 ; ’ABHf9bSgYrkH3d<"#$% ; :-**)2#,’-+/ $),7))+ )+1’*-+<)+,#2 (#.,-*/ #+0 45)+-,34’. ,*#’, <)#+/J=)+),’. 4#*#<),)*/ -( ! % "#$%&%’()*
’+ 6 7’20 4-482#,’-+/J*)/4).,’1)23
ij
L%6,
oL0Puv M46.-%90$8 %"#$% ,6#.& ?_$P,- N6.6%$8 0#"#,6%6"&
# 2: $ 2 F% 2 %,#G $0 & 2H $ * F% * $& F%& %&I ’& MMO ( ) *N
iV O#%$%1)6 K A>CB;! K A>B<= K A>=A; K A>@?C! K A>DDE A>AB@ A>=@@ A>?EC K A>B?; K A>D<< K A>D=? K A>;;A
CBD!! A>;B@ K A>B=A K A>?DA K A>@@=! A>B?; A>=@E A>DD@A K A>?;; K A>?A= A>A??
tu P26:#%$-. K A>=BE K A>D=? K A>D=? K A>D?= K A>=?= A>AED K A>;@< K A>@DE! A>CDB!! A>B=C K A>AB< K A>ADB K A>;==
jI()
N6-/"#04$8 )$&%#.86 K A>AC@ A>;== K A>B;@= K A>ADD K A>A;< A>==B A>AEE K A>;;C A>AEC A>;<@ K A>;DD K A>A?C
$}+ +..1#2
0"68$0$%#%$-. K A>D?? A>BCD K A>BBC K A>D@B A>BEC K A>ABB A>=
?AD; K A>AD?
$uF +..1#2
#:6"#/6 %6,06"#%1"6 A>@BA
! K A>DB; A>=AC A>C;=! A>=DA A>=
;E? K A>;?B A>;B= A>B@@ A>BC< A>Dfb‘-
(1.&4$.6 4-1"& A>=@D=
! A>=BE A><<; A>=;; A>;B@ A>A
DBC K A>C;E! A>D@A! A>DB?! A>DD=! A>AB@
T) !-%#2 04-&04#%6 A>B?C A>AAD K A><<@ A>B<= A>BCC A>ABB K A>=DD K A>B=@ K A>AB< A>AE; A>AE? K A>;A;
T% !-%#2 .$%"-/6. K A>B?@ A>?DD K A>BCD K A>BC= A>;;; K A>B;@ A>DCC A>D?? A>BC< A>B=? K A>;Z2V
Q"/#.$8 ,#%%6" K A>BAC K A>C;;
!!K A>DDA K A>DB< K A>=B@ A>DE= A>;A= A>D;@ A>B@D A>;AB A>A@? K A>;A=
0R A>@@=! A>A=?@ A>D@C?! A>;?C K A>?A@ K A>DE< A><;= A>
<=< K A>A?@
oK2qXTH+"bH41$P=>&9LHK:T,
vSTs2Ao !clopLHK5_ST:1%0
P$1yn|*es-*1gl*n/yn|*4jn
x4_k=V9l
!"! ’ABChHf #$%& QRc;$(
"#h #$ %& 1267\ ’ hH+"bH41
#"$hi‘ ()*mn +*,(?@*vr$Z ""- h
%S*tuRh67r$Z #$./h%SAo "cld
#$67 0#$-*0""x 0#1$"\41 ,’ ::8d8l
fP ,2,34)4 #.5"=>^9:41tu3I
%S0;<==>=ZlB-A "c?Iw !-."/?*$."1! #x
$.5!" ’*9B0~{!41A,"c:$P3i0zA
A,,6 @ !’.’A?*! @ $."/- $*" @ $.1$$ /c*ogp
341A,5czsA,,6 @ !$.11?*! @ $.""1 1*" @
$.55" -cl^ ,2,34)4 #.5" ^9: )789:g5?}
,-A#0B @ $.5$# 1cLAP *C2D* =>\ ’ hH
+"bH41:?@mngbo‘*-41<:$
P=>u*;=>: #E5Ao /cl
’hH+"bH41#"$hi‘gB+*,(W
d # 67 0#$-A*c*0""AFcx 0#1$"AGc
\41 ,’ ::8d8
H8IJ# +*,( &K>L8M7: >L &>&NM= )>J’ >L
$ J %&’()(*+,-A0#$-A*c*0""AFc<=P 0#1$"AGcc
X:mnZ 1h$P,-LHK5_ST*9B3
I%S=
ZlB-A "cL$fb‘-s2STAST,-?I
w $.-/$*$.--1x $.-"!c*$P()A.3cL$uF
?>s2STAST,-w $.-5-cAo !cl
9 ’ ’ABChHfPKH3)x
()*+ ’ ,-.-/01 2033-4-./0)/05. 53 ! + "#$%&%’()* 0. 6 7082 959:8)/05.;
BW
D,2,34)4 K7:NMO: *C2D* K7:NMO:
41>g53i0
37=7 P8Q7K:8OR S8O;8=
&>&NM=A!0c
;:g53i0
B>OA!Bc
g5?},-
G>7LL8T87=O >L I7=7
P8LL7K7=O8=A#0Bc
41<
*U>=I
&>&NM=:E?
41>
V8O;8=
&>&NM=E?
74+t 0&7T87: M7Q7M $."1! # $.5-# 5 $.5$# 1 5-.1"A/ W $.$$#c !1.-’A/ W $.$$#c
W\r 0( $."#1 5 $.#!’ /
!"< I*)@
gBoL0Px +*,(WX/T: ’hH+"
bH41Cr@:d " xd 5l^d^:*gB #-
hoL0P:4!()Cr@S ’ h41tw 5
rq41 ,5*,1*,-*,! x41 ,#*,"*,/ ?Iwor*
41 ,’ STworXgB "#h67 +*,(WX:
)789:{y$P()Cr@tS ’h41?w "r*
- ,#*,"*,5*,1*,’ - h41Cwor*,-*,! x ,/ 5
h41Cwborl^:*" rWX:Crmnz
dW[r>l
d " gB;IAoLc-‘:Cr@
H8IJ" (7=PK7IKL ’ S8MP &>&NM=: >L
$ J %&’()(*+,- %<:7P >= &;7=>OR&8T T;d 5 gB +*,(WX:Cr@
H8IJ5 (7=PK7IKL ’ S8MP &>&NM=: >L
$ J %&’()(*+,- %<:7P >= +*,( U1! & ’ ( ) 1-"
! mJL[J
$P3i0^NoL*hAO*2%k*?_W
XxEg/Y23h-3mnDEx:;lbDE
NoLx "#$?_+t8\XT % hH+"bH
41$P=>mn?@*mnop*H+"bH41
doLx "#$?_+t8uzdW;<:$P=
>l74+t8*&’hoL0Pd41>x41<
: ()uzdW[r>*9B4_k=V9=>z
[*n/*1x4_:;PB-Ayn|c=>z{*
10Px00Wn/x4_0PAX*$+"WX_
1 % hH+"bH41tu ++,*! x " ?Iw
-’.&/0*1.23- &x1.!-2 %*XLO}A211-c\XT
/ &&hH+"^k1O //hhO:?@mngb
o‘*LwC:YZ"A#$%&’( (’)*+,$-cA4567 $% (’ 8*
211!c$P3i0+tSX*ABo7N{1@4:
^_vAJ}*&9%%cloL& *$+"WXus1*H
+"bH41$P=>u 2:!zdB41>*&:!z
dB41<*XL@A@4-",A .+/+*,$),/*)
0!+)$)-$cA0 T v 2*2112c*h ]A 1(%!(2(
(/32/*4!2’’(cA#{T2*&99/c2Sgl
ar?@op*H+"LoL0P41<&4
1>uzds2Y%s2r>*Xtp!woL0
P\woHK5_:mf0&wooL0P.!w
HK5_:78!w*-oL=>$g5LxJK
5_"P:mnlH+"oL0PB*1yn|*e
s-x1glL2HK5_{s2Y%s2bY}ST*tpXh
JK5_\H+"1;I89:78Ws2X4j
nLtu{s2}ST*XLYZ":=>JhAZ
,2*211’co‘*_$^<$H+"4_\JK:
o4mf0og*!wtuJo!w*MJx4jn
4x784_}k:()X4_k=V9Ltu{
%s2bST*L$fb‘-{s2}ST*LVST!s2*tpQ?{HKsH+"NYmn
‘Lj:{z{=}*,1p>2^<\4_Hy
5zeHW[78*XZjsRmos]eDEl
gB *$+"WX:H+"bH41$P3i0
B- ++,7Y ! x " 4L$fb‘-{s2ST*
L‘-*$P3i0a37o0:F0"PlXLLBo2
A2112c\dY@A 5$3+0$/(- 0*/)+0&’(%(c41:DE*
Y&>oK2A211%c\!wtuH+"41:DE
mnSgll?*41<:$P()L$uF?>
^{s2ST*|L41:jI()ST!s2*o
pjI()"P!pslST?@mnop*HK
\XTH+"bH41$Pm/78W[*|^B
gzbH414|W{*5l{41=fxHKq
(}256^Z^<7841:$P?}l
gBoL0Px *$+"WX:CrmnzdW
[r>*1%^<$^BoL0Pf.JK78*?
_WX!.JK78~$lbDEopH+"$P
=>u 2:!]B41>*tp{HK$9=>:1
%,]l5l*d4V\]d<*:;&MNDE
‘*f[?"#HK5_:"P*29N3-m*9
%34DE+^*\KKmnjN:;*pj:;4V\]ll?*bDE~z&XTK>:
bHH+"41*9mnV8*C]-m:mosDEl
d e , -
O } 8 211- 8 H+"^k1O$P3i0DE 8 /W&’[))
**2%A-cq2% ; !38
J } 8 &9%% 8 w8kL&,1O$P=>DE 8 ZW&’[))
**&2A&cq-% ; //8
J }*ip%*OW6 8 &9%% 8 Pw4kDE-S21O:$Pm
/ 8 &’()*23A3cq!99 ; 3198
0Tv 8 2111 8 H+"^k&1089DE 8 &’()*!-A2cq
&22 ; &238
0Tv*]#5*QNH*2 8 2112 8 *$+"WXDE-",jI4
1:$P?} 8 &’()*!%A3cq-& ; --8
0#A*>oK*L0;*2 8 211- 8 !wMzac\H+"; "#$
J’=n:78 8 ?_"7z4*3A&cq&!& ; &!3 8
LBo*& E 8 2112 8 dY@41$P=>LJK=9:T, 8 H
I)**22A&&cq&9&2 ; &9&-8
UqT 8 2111 8 H+"\]:N5APL58o/ 8 \]N5Lt
n*&-A3cq221 ; 22&8
#{T*& "*x’(*2 8 &99/ 8 2(,b5z:?_H7)‘
‘ 8 "7?r)**!’A2cq9/ ; &118
>oK*0#A*L0;*2 8 211%8 6y-!wtuH+"41$P
3i0&9LHI5_:ST0 8 &’()*33A!cq’1 ; ’’8
Z ,*Nt+*!ap*2 8 211’ 8 xYYZ"^k01:oL=
> 8 &’()*3&A2cq’9 ; -’8
NIo 8 &9%’ 8 BC@,#q!j" 8 /WqBC&’U4w*2&&! ;
2&!&8
<=>?@@A5B ,8 &99’ 8 $CD)$ &.’’A$6EFGHAH ?@ C?F5>IFEB )EBAE6>5c8
J565KA>H E6L MA?N5KBG ,EO?BEK?BG*P6AQ5BHAKG ?@ J565QE*RIAKS5BFE6L8
,A +*M5EIFA5I T*M?IHUI5K T8 &99/ 8 J565KA> HKBI>KIB5 E6L VEKK5B6H ?@
7565KA> QEBAEKA?6 EN?67 V?VIFEKA?6H A6 5EHK5B6 WXAK5 RVBI>5A 6+0$(
3’(&0(c8 (E6ELAE6 T?IB6EF ?@ Y?B5HK *5H5EB>X*2/A2cq&%9 ; &9%8
CEBZ + C8 &99/8 [??FH @?B +?VIFEKA?6 J565KA> $6EFGHAHA[Y+J$c8 )5BHA?6
& 8! 8 XKKVq::\\\8 X5BO8 OA?8 6EI8 5LI:]NAFF5B 8
CAFF5B C +8 &99/ 8 $CD)$]+*<+*E VB?7BEN @?B KX5 VB5VEBEKA?6 ?@ $CD)$
A6VIK @AF5H @?B IH5 \AKX W^#$CD)$8 "5VEBKN56K ?@ MA?F?7A>EF
R>A56>5H*#?BKX5B6 $BAS?6E P6AQ5BHAKG*YFE7HKE@@*$48
4567 TA5*4?I _IVA67*MEA TAEGI*$% (’ 8 211! 8 *$+" E6EFGHAH ?@ 7565KA>
QEBAEKA?6 A6 \AFL V?VIFEKA?6H ?@ #$%&’( (’)*+,$- @B?N JIE67=A*(XA6E8
E*&!3A&cq!! ; 3&8
!"#$% - *,
’-! 9# >oK2qXTH+"bH41$P=>&9LHK:T,