免费文献传递   相关文献

Phenotypic Plasticity of the Seedlings of Camptotheca acuminata and Gordonia acuminata under Different Light Regimes

不同光环境下喜树与四川大头茶幼苗的表型可塑性*


研究不同光环境对常绿阔叶林中喜树和四川大头茶幼苗生长的影响。结果表明: 1) 落叶阔叶树种喜树幼苗通过提高自遮荫程度、增大叶悬挂角、增加根生物量比和减小比叶面积适应旷地环境,在旷地中具有最大的相对生长速率,而常绿阔叶树种四川大头茶幼苗没有表现出对旷地光环境的良好适应; 2) 四川大头茶幼苗在林窗中通过增加叶片数、提高单叶面积、比叶面积和叶生物量比接受更多光能,相对于旷地和林下具有较快的高度生长,喜树幼苗比四川大头茶幼苗在林窗中具有更大的单叶面积、叶柄长度、总叶面积和总叶面积增长率; 3) 在3—8月,喜树在林窗中的株高增加约35.3 cm,约为四川大头茶(约6.6 cm)的5.3倍,喜树在林窗中具有明显的光资源竞争优势,表现出比常绿阔叶树种四川大头茶更高的相对生长速率,这就是喜树存在于常绿阔叶林的一个重要原因; 4) 郁闭林下的光资源有限,2个树种幼苗都不能在林下良好生长; 5) 喜树幼苗叶性状在不同光环境的可塑性指数远大于四川大头茶,这是喜树能够在常绿阔叶林中存在的另一重要原因。

The effects of different light regimes (such as open, gap and understory) on the phonotypic plasticity of the seedlings of Camptotheca acuminata and Gordonia acuminata were studied, based on the leaf traits. The results showed that: 1) Seedings of C. acuminata, as a deciduous broadleaf tree, mainly improved the self-shading, increased the midrib angle and ratio of root biomass to total biomass, and decreased the specific leaf area to adapt the full light in the open sites. Therefore those seedlings exhibited the highest relative growth rate in the open sites in compared with the other light regimes. However, the seedling of G. acuminata did not show the similar adaptability in the open light environment. 2) In order to receive more light energy in gap, the seedlings of G. acuminata developed more leaves, and larger single leaf area and specific leaf area, and increased the ratio of leaf biomass to total biomass. Therefore, the seedlings of G. acuminata could grow relatively higher in gap than that in open and understory. Compared with G. acuminata, the seedlings of C. acuminatain gap had larger single leaf area and total leaf area, longer petiole, and higher growth rates in height and total leaf area. 3) From March to August, the height of the seedlings of C. acuminata in gap increased by 35.3 cm,which was about 5.3 times of the height growth (6.6 cm) of the seedlings of G. acuminata. Therefore, in the gap, C. acuminata seedlings with a distinct competitive advantage of light resources showed a higher relative growth rate than G. acuminata, which was one of the major reasons why the deciduous broad-leaved tree of C. acuminata in evergreen broad-leaved forests. 4) Because of the lack of sufficient solar energy, the two trees’ seedlings could not grow well in the understory. 5) The phenotypic plasticity index based on leaf traits in different light regimes of C. acuminatawas much higher than the those of G. acuminata, which was another important reason for the existence of C. acuminata in evergreen broad-leaved forests.


全 文 :! !"" ! ##
$ % % #$ # %
& ’ ( )
&’()*+(, &(-.,) &(*(’,)
./01!"**/1#
&234*
!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!
$ % % #
!wHJK+‘@Lab[cdef:oL^g0"
012 # 3 456
ABC&’()DEFG&HIJKLMNDEO /W 5%%%#5c
! "# DE!wHJK\O({1&B‘@xab[cdefHy:78lmnopq5c01{1@4‘@
ef~@JARhiUV*8[1jkl*8e!H79|xN{|13EmfnjJK*dnjBrZz[:S
\Hyo(*MO({1@4ab[cdefYZogU\njHJK:p1mfX$cab[cdefd&kB
~@8e1q-*JAr13E*|13Ex1H79|D.C3H<*S\Bnjx&+rZWs:AVHy*
‘@ef|ab[cdefd&kBrZC[:r13E*1tyV*;13Ex;13E8y(X6cd 6s7
%*‘@d&kB:‘A8eu 6"16 89*uwab[cdAu :1: 89c: "16v*‘@d&kBrZps:H\]w
xyF*ogU|O({1@4ab[cdCA:S\Hyo(*XL$‘@zdBO({1&:ohR%W5X
!cFG&+:H\]Z}*$h@4ef4!G[Bab[cd*X$‘@$%&q HJKX10PXoL^g0X‘@Xab[cd
’()*+#&;571!6 ,-./0#, ,12+#5%%5 < ;!773$%%#4%# < %%%: < %;
def#q$%%; < 5$ < 56Xzjf#q$%%# < %: < $"l
ghijqCPRk()gh38ijA6%6;55!5cX(l?yz0ijm{m|[}jHI,bjN:;L~!qA$%%6=(>!?5!cXC"F
{m#LCP&’Q$N\%ijA&@$%%5 < %5$cXCP&’QRSTUVghl
"&J5w~!"#l
!"#$%&’()* !+,-&)*)&’ %. &"# /##0+)$1- %. !"#$%&%’()" ")*#+,"%" ,$0
-&./&,+" ")*#+,"%" 2$0#3 4)..#3#$& 5)1"& 6#1)7#-
AB/ CDEDBF GFHI J/HI @EF/ G2HKF
A !"#$%$&$’ () *(+’#$ ,-(.(/0*,"1%+("2’"$ 3"4 5+($’-$%("*67* 8’%9%"/ 5%%%#5c
89-&3,*&q +D2 2KK28LM /K NEKK2O2HL 0EIDL O2IE92MAMB8D FM /32H*IF3 FHN BHN2OML/OPc/H LD2 3D/H/LP3E8 30FMLE8ELP /K LD2
M22N0EHIM /K 632:$($;’-3 3-&2%"3$3 FHN <(+4("%3 3-&2%"3$3 Q2O2 MLBNE2N*RFM2N /H LD2 02FK LOFELM1 +D2 O2MB0LM MD/Q2N LDFLq5c
&22NEHIM /K 6 4 3-&2%"3$3*FM F N28ENB/BM RO/FN02FK LO22*9FEH0P E93O/S2N LD2 M20KTMDFNEHI*EH8O2FM2N LD2 9ENOER FHI02 FHN
OFLE/ /K O//L RE/9FMM L/ L/LF0 RE/9FMM*FHN N28O2FM2N LD2 M328EKE8 02FK FO2F L/ FNF3L LD2 KB00 0EIDL EH LD2 /32H MEL2M 4 +D2O2K/O2 LD/M2
M22N0EHIM 2UDEREL2N LD2 DEID2ML O20FLES2 IO/QLD OFL2 EH LD2 /32H MEL2M EH 8/93FO2N QELD LD2 /LD2O 0EIDL O2IE92M4 V/Q2S2O*LD2
M22N0EHI /K < 4 3-&2%"3$3 NEN H/L MD/Q LD2 ME9E0FO FNF3LFRE0ELP EH LD2 /32H 0EIDL 2HSEO/H92HL 4 $c(H /ON2O L/ O282ES2 9/O2 0EIDL
2H2OIP EH IF3*LD2 M22N0EHIM /K < 4 3-&2%"3$3 N2S20/32N 9/O2 02FS2M*FHN 0FOI2O MEHI02 02FK FO2F FHN M328EKE8 02FK FO2F*FHN
EH8O2FM2N LD2 OFLE/ /K 02FK RE/9FMM L/ L/LF0 RE/9FMM 4 +D2O2K/O2*LD2 M22N0EHIM /K < 4 3-&2%"3$3 8/B0N IO/Q O20FLES20P DEID2O EH IF3
LDFH LDFL EH /32H FHN BHN2OML/OP 4 ’/93FO2N QELD < 4 3-&2%"3$3*LD2 M22N0EHIM /K 6 4 3-&2%"3$3 EH IF3 DFN 0FOI2O MEHI02 02FK
FO2F FHN L/LF0 02FK FO2F*0/HI2O 32LE/02*FHN DEID2O IO/QLD OFL2M EH D2EIDL FHN L/LF0 02FK FO2F 4 6cWO/9 XFO8D L/ ,BIBML*LD2
D2EIDL /K LD2 M22N0EHIM /K 6 4 3-&2%"3$3 EH IF3 EH8O2FM2N RP 6"16 89*QDE8D QFM FR/BL "16 LE92M /K LD2 D2EIDL IO/QLDA:1: 89c
/K LD2 M22N0EHIM /K < 4 3-&2%"3$3 4 +D2O2K/O2*EH LD2 IF3*6 4 3-&2%"3$3 M22N0EHIM QELD F NEMLEH8L 8/932LELES2 FNSFHLFI2 /K 0EIDL
O2M/BO82M MD/Q2N F DEID2O O20FLES2 IO/QLD OFL2 LDFH < 4 3-&2%"3$3*QDE8D QFM /H2 /K LD2 9FY/O O2FM/HM QDP LD2 N28ENB/BM RO/FNT
02FS2N LO22 /K 6 4 3-&2%"3$3 EH 2S2OIO22H RO/FNT02FS2N K/O2MLM 4 !c>28FBM2 /K LD2 0F8Z /K MBKKE8E2HL M/0FO 2H2OIP*LD2 LQ/ LO22M’
M22N0EHIM 8/B0N H/L IO/Q Q200 EH LD2 BHN2OML/OP 4 "c+D2 3D2H/LP3E8 30FMLE8ELP EHN2U RFM2N /H 02FK LOFELM EH NEKK2O2HL 0EIDL O2IE92M
/K 6 4 3-&2%"3$3 QFM 9B8D DEID2O LDFH LD2 LD/M2 /K < 4 3-&2%"3$3*QDE8D QFM FH/LD2O E93/OLFHL O2FM/H K/O LD2 2UEML2H82 /K 6 4
3-&2%"3$3 EH 2S2OIO22H RO/FNT02FS2N K/O2MLM 4
:#’ ;%30-q 0EIDL O2IE92MX02FK LOFELMX3D2H/LP3E8 30FMLE8ELPX632:$($;’-3 3-&2%"3$3X<(+4("%3 3-&2%"3$3
G&:HJK()3=A’DF[N/H ’$ 3. 4*5##:cl
"7mf>VJK:<*\9HzxHe*;2+
TR%*oL^g0:[{$,9"7\>VJK
mf<*:R%BW*-2wo,HI)DE:o
hR%prA!"# !" #$ $*%&&’X()*+,-"". !" #$ $*
%&&/ X0+#1234+-+ !" #$ $*%&&/cl-ZDEop*"
71%$~@;I)=}Ymf!w:HJK*.
$Z:HI=}^1%/5B|1R:=}
A5##,+6+72 !" #$ $*899:clq;4|0i4rZCA
:oL^g0A;".."-"<+2 !" #$ $*%&&&cqL0i0@
4S|*q;4:1Hy(x&2]u(A <"7+ )=
7#22>+ 7><,)?+@4|13EA2@+3#=#3 .+"= "<+"*A!Bc}s*14l
xO51-8e*678es*5lq;4OrZW
A:9:Hy(A?+<7#3". C<)D74 <"7+c*;23-@D
YJARhiAE..#2 !" #$ $*%&&&XF"."?"2# !" #$ $*
%&&8X G2"-" !" #$ $*%&&HX I+"<3J !" #$ $* %&&’X
()*+,-"". !" #$ $*%&&/cl;".."-"<+22A%&&&cBU@
;\"7oL^g0:DE3=}:HJK*M\>B"7?O^g0a3:D
EAs"‘HyxH79?@:^g0c*Mb$X
4?O:^g0F0B"7BxHyl
‘@A %#&’"(")!*# #*+&,-#"#cC D n @ (
A5J22"3+"+c"7*A) H& 6*xkBECyQYZL
/&yQY/??j[*$&’vO({1&/Z
:o4A[01F,lab[cdA .(/0(-,#
#*+&,-#"#cC-d(AK4+"3+"+c*O({1[F,*R
k?kBb_*bZtu %&& L 8 M’& 6:j[*w&
’vO({1&yF4~ol
bDEYR^GH-RkMN[:ab[cd
x‘@wDE\z*fgHIJj*&kx&+2!
wHJK*|W %h@4d!wHJKB:10P*
"‘Hy*H79&9?@:=}*h[O({1@
4x01{1@4\!wHJK:oL^g0r>
Y&A[01{1@4"wO({1&/Z2?:
W5l
8 KKLac
ijLdYZ[)HIM>*()GH-Rk
MN[u % N6*9$z{P*L-<*p@:GH
-z{P*SoANO2*89::cldM>fgnj
AHIJjc*&kx&+ H4HJK*R4HJK5
Uiju 8&& 6%lnj>{F+,X&k^Ao
%& 6:‘@*oPA%,--#&(&+& *#&’)(/#c2F,&
ohA[+,/2X&+HJKfgd^A[F,
xQ%+,/2:RSJK*9FGVu &O:lD
E#<0#GN*M-HJK:gbx0l
%&&’$ H%9*NGH-fTjL’[{xH
yP*o‘: %$H‘@xab[cdefL %&&
I‘*9B‘@eftu‘AA%HO/ P 8O%c36*gC
A%O: P &OHc66*{1qXab[cdeftu‘A
A8’O’ P &O/c36*gCA%O8 P &O%c66*tu1q-
9OM P 8O’lefUVd] H& 36*:Cw %’ 36:Y
WB*VX,’RGH-efHyj*eeo09Z
2YZ[lef2BR*UgBfg1:nj*&k
x&+*R4HJKRh@44" ’& ‘Y8ef*
5U#<29\f+Yl
%&&’$ :%*vL\ ’h]^*cPzF_*T
xOFV_A^_zz‘oac*!#Q8M&& bV_x
!#QR)< /M&&kc"HN$0,bAdC !#QR)?I$0 HhHJK:ftuzF*S\xV*HN
Z=efxgx+zhr2JK5_l
%&&’$ :%+i*d HhHJK+12L’‘
@xab[cdefL 9‘*AP !#QH&&&kc"1
3E‘AdC !#QR)0R‘:r13E*r1yV*r1nV*Poa$
0Rq1:1tyV*zR_9r‘;1-l
|13EAA!BcKPpqrc$0ldRj$
‘8’no1Bs:{tq H& h*STRUTV
9lA!B S 1 T2*"B 1 Wo{tq3E*2 Wo
{tqTV9l’ 9‘$0:tuv"w>4d>
HJK+: A!Bl
Y :%| ’h]^>$0:1jklx1tl
f=}:tuv"w>4d’oHJK+:1jk
lx1tlv*ac,"O~u2A%&&8cl
d$10P:w‘*$ 9‘ef:‘AxgC*
r‘;13E:tuv"w>4d>HJK+:H
yBWl!‘ H%9_:SfBW*_9‘A*gC
x;13E:%tu8y(l
dZLj$"‘*?!*6*1d :M UST+v
V9*kR?IU9TV9*V_9;H79*!H
79|*6H79|x1H79|l"7:He*
PS\Hyo(A6CyCV 8 "V 8c,o1A5)-" !" #$ O*
%&&Mc*_9y"s+q
(W( SA.,&% V .,&8cTA "% V "8cl
"Bq(W(A<+."7#?+ C<)D74 <"7+cwS\Hy(*&8 x
&% ?Iw H %x : %$0:efH79*"8 x "%
w %m$0:‘^g0B-:_9q^g0B-2B>^g0
BWd HhHJKB:z[vN;z{vdwYz
[v*ac," B247),2A899:cl
B5G;B?@x !AX 3R|Wd ABA :O& +
mnl
Y! 9# 4#.2q!wHJK+‘@Lab[cdef:oL^g0
! mnL?@
!"# fSg
!wHJK+*‘@ef:10Pr>s2
Ao "cl9B*nj‘@ef:tu1q-zAA##$% &
"!$!c*%s2[B&kx&+A! ’ ($("cX‘@ef:
r13Ed&kz[A%%$) & "!$(c*+!*uw&+:
!$,vXnjx&kB‘@ef:1yx1nu%s
2[B&+A! ’ ($("cXl?*&kB‘@ef:1t
yV%s2[Bnjx&+A! ’ ($("cl
w1tyV?*ab[cdef:9I10P
BWd!wHJK+urZs2r>Ao "c*9d
&kB:1q-*r13E*1yx1nu%s2Y
s2[Bnjx&+l
‘@ef1jkl:ftuvd!wHJKB
rZ%s2r>A! ’ ($("cAd "cl1Hb!V8
e*‘@ef:1jkls28[*nj:1jkl
zAA#) & )$#c-*w&+: )vY8XMab[c
def1jkld!wHJK<{s2r>
A! . ($(#c*uw "/ 0 !)-Ad "cl
9 # fSg3! $ %4!
&’()# *+’, -.’/-03! $ %4
74
123*435
10P
6378 9:7495
&+
;<=3:59>:?
&k
@72
nj
A23<
‘@
"#$%&’&()*# #*+$,-#&#
1q- 6378 r13E 141y 6378 J31n 6378 N4=9MK*+ !$) & ( E)F )$D & ( E%H )$" & ( E#H
1ty O394>J3 J3ab[cd
.’/0’-,# #*+$,-#&#
1q- 6378 r13E 141y 6378 J31n 6378 N4=9MK*+ "$# & ( E!C !$" & ( E!7 "$, & ( E)C
1ty O394>J3 J3! won>:!w[xyzo1uvd!P ($("+t8r>s2*won>:!w{xyzo1uvd!P ($(#+t8r>s2l+
wlQ49M4< 37*M :>N*9M3 R7JB35 5M7:4N*9M3 R7JB35 5M7:4N3:*753
J3993:5 +37< =4883:3<*35 79!P ($(#$ SM3 57+3 C3J>N$
d " jklx|13E
T4IE " U4=:4C 7!wHJK+*‘@ef:|13Er>%s
2A! ’ ($("cAd "cl&+‘@ef:|13Ew
AD"D$% & ,D$)c*+!yIV "*$nj: !$/v|{l1
7Hb!V:8e*‘@ef:|13E[[}s*
1q%s28NA! ’ ($("clab[cdef|1
3Ed!wHJK+r>s2A! ’ ($(#c*nja
b[cdef:|13EA"(/$! & !"$#c*+!yIV "W
&kx&+u}s )(Wl
!"! hij.
/%|*!wHJK+‘@ef:‘A*gCx
;13Eurr>s2A! ’ ($(#cAd !cl&kB
‘@ef:‘Az[A#/$G & !$Dc*+*&+z{
A!G$% & %$Dc*+X‘@ef:gC17HKBHb
!V:8eM8[X‘@:;13Ednjx&k
{s2r>*d " !#( 0 " %D( *+! ~<*$&+‘@
;13E: DvY8l
/ & ’ ( ) %#"
ab[cdef:‘AA!!"# $ %"!c&’xgC
A("! $ )"%c&’ ud&kBz[*d&+z{ll
?*&kBab[cdef:;13EA#*#"+ $
(+"+c&’s2ABnjx&+*?Iuwnjx&
+: !",x ("+vl
R (%9+ -%|*‘@xab[cdef‘
A*gCx;13E:%tu8y(d!wHJK
+r>s2A! . )"),cAd !cl‘@efd&kB
:%9:Hy(z[*)A,"/ $ #"!c&’*njm~*
M&+z{X9gC%8y(1H!:8[M8
[*njB‘@efgC:%8y(uwA#") $
)"#c&’*w&+: ,,v|{Xl?*njx&kB
‘@ef:;13E%8y(us2AB&+la
b[cdef:‘A*gCx;13E%8y(u
d&kBz[*d&+z{XVQ9;13E8y
(d&kBwbvAu !) &’!c*Mdnjx&+t
Ug}8y*?Iw 0 #"*x 0 %"# &’!*Xopab
[cd!d ! !wHJKBef:HyBW
1234 ! 56789: 2;<=> 7? 9:= @==!"# hCEkhlm
-%|*‘@xab[cdef:H79d!
wHJK+:r>%s2Ao !cl‘@ef:!V
9*1V9x;H79ogU1HyH!:8[M
8[:EF*njB:vu%s2[B&kx&+
A! . )")#cX|dnjx&kB*‘@ef:6V9
{s2r>A! C )"),c*u|&+‘@:6V9A
, D *vlab[cdef:1V9x;H79d&
kBzA*%s2[Bnjx&+:SfvA! .
)")#cX|ab[cdef:!V9x6V9dn
jx&kB{s2r>A! C )"),c*u|&+:S
fvAu !vYY8l
9 ! nohCE
$%&’! ()*+%,, *- ./0 ,0012)34, 3
74 EF=&2=@ &? G7’F7;=;9@ &+ H;<=6@976I &k 5JF nj KF=;
‘@
"#$%&’&()*# #*+$,-#&#
! L779 )"#, $ )"),G !"(, $ )"(-M +")( $ #"%/N
6 E9=’ )",+ $ )"#,M ("!- $ )"*!N ("/, $ #")!N
1 O=J? )"(( $ )"#)G ("-- $ )"+)M +"%/ $ #"!%N
?‘ P79JA #")% $ )"!+G /",# $ )"/!M #+"%* $ ("#+N
ab[cd
.’/0’-,# #*+$,-#&#
! L779 )"!! $ )")*M )"+, $ )"!,J )",- $ )"!*N
6 E9=’ )"(# $ )")/M )"+, $ )"!)J )",/ $ )"#/N
1 O=J? )"(! $ )"#%M #"#* $ )"%(N )"+* $ )"(*M
?‘ P79JA )"-, $ )"!*G !"%* $ )"*%N #"-( $ )"*!M
N (%9+ -%|*‘@xab[cdef:
S\Hyo(d!wHJK+r>%s2A ! .
)")#cAd (cl‘@ef:S\Hy(1HyH!
:8[M8[*njB‘@ef:S\Hy(Au
#,"* ’3y30 #c?I$&kx&+: #"! x %*", vX
Mab[cdefd&kB:S\Hy(z[Au
*"% ’3y30 #c*&+z{Au #", ’3y30 #c*&+ab[
cdef:S\Hy(~$&+‘@efS\Hy
/! /# 4#.2q!wHJK+‘@Lab[cdef:oL^g0
d ! !wHJK+:S\Hyo(
"#$% ! &’()*#+’ $,-.*/ ,)*’ 012’, 2#33’,’1* (#$/* ,’$#4’
(: 5v|{l
‘@efd!wHJKB:H79?@rZs
2r>A! 6 7875cAd 9cl17HKBHb!V:
8[*‘@ef:!H79|s2JA*|6H79
|!s2+}Xl?*njx&kB‘@ef:1
H79|us2AB&+lL‘@ef!w*ab
[cdef:!H79|x6H79|d!wHJ
K+{s2r>A! : 7875c*~1H79|d&k
Bs2ABnjx&+l
!"# pqSj.
dgB10P:^g0BWB*‘@ef:L
^g0vu[Bab[cdefXdgBHy: !
hBWB*‘@ef:‘A*gCx;13E:^g
0v?I$ab[cdef: ;8<*;8=x <8!vX
dgBH79&9?@:L^g0BWB*w1H
79|xS\Hyo( ;hBW:^g0vSr!
[?*‘@:9ILBW:^g0vu[Bab[
cdljN|WL^g0BWop*‘@ef\!
wHJK:oL^g0[Bab[cdef*9t
u^g0v$ab[cdef: <8=vAo !cl
d 9 !wHJK+:H79?@
"#$% 9 >#-4)?? )((-@)*#-1 012’, 2#33’,’1* (#$/* ,’$#4’?
9 $ 9bpqSr
%&’($ )&*+,- ./ 01, 21,3.04256 2*&-056504
BW
A12’B
‘@
"#$%&’&()*#
#*+$,-#&#
ab[cd
.’/0’-,#
#*+$,-#&#
1q- C’)3 104D’, 78E= 789E
r13E F#1$(’ (’)3 ),’) 78=! 789=
1y C’)3 (’1$*/ 78!! 78!;
1n C’)3 .#2*/ 78!E 78;;
1tyV G’*#-(’ (’1$*/ 78;H 787=
|13E FI’@#3#@ (’)3 ),’) 78== 78!J
1jkl K#2,#D )1$(’ 78=J 78;<
‘A G()1* /’#$/* 7857 78;9
gC F*’4 2#)4’*’, 78== 78;5
;13E L-*)( (’)3 ),’) 78J= 78=E
!V9 &--* .’#$/* 78HJ 78==
6V9 F*’4 .’#$/* 78J= 785;
1V9 C’)3 .’#$/* 78H5 78E!
;H79 L-*)( .’#$/* 78H9 78==
!H79| &--* .’#$/*M*-*)( .’#$/* 78=; 78<=
6H79| F*’4 .’#$/*M*-*)( .’#$/* 785= 78;E
1H79| C’)3 .’#$/*M*-*)( .’#$/* 78;; 78;=
! mJL[J
dnj*‘@:H79xS\Hy(zA*?I
w&k: <8E x <8; v|{*ogU9\!HJK
:p1mf0qe1q-x;13EAX ;iBWudnjzAc,
JA\H<:AP*YTgzA:S\Hy(X;c
L&kS|*nj‘@:‘AW{Q6H79|s
2}s*NM8[B1qS[R":UV*JABR
hiA?’(3N?/)2#1$c+t*Z=N{BAH\"7:#
$AK)()+)?# )& #1 8* ;77&-P’12))( )& #1 8*;77=cX!cnjB‘@ef~@8
e1tyV*8[1jkl,N{!H\ef1q
:%$*Tg\!H:p1mfA4#.2*;77=cX
9c‘@ef~@JA!H79|x}s|13E*
cefdnjB~@!?[?Z’+?x&V’(
V[9:8zdNVA1q=Nc:1qB*N{B
njB!Hx+?’(\ef:#$l
S3*ab[cd)k$q;@4*9ef~k
7< & ’ ( ) 95"
!b[cdef:‘AL&kr>!s2A ! "
#$#%c*|9dnj:H79~w&k: &’(*1q
-*;13ExS\Hy(us2sB&kA! )
#$#%clR‘AWA*1q-WI*t!@JA"‘:Rhi+t,*mnj!H%$lw
‘*ab[cdef:1ti+*~ #$* +,*!AB
1qL)Xb?*nj:ab[cdefVMSC
3:HNe7?@X!*9!H79|L&k{s
2r>A! " #$#%cl5l*ab[cdef!<~
@8e!H79|,JA"7\+?:,d<*Y
mf!H+:A+?-.ldnj*‘@ef:H
79*S\Hy(2uG[Bab[cd*‘@\T
HbJK:oL^g0xmf<*uG[Bab[
cdl
ab[cdx‘@ef4\&kXoB2H!
:HJKogUp1:mf<*lLb? *hHJ
KS|*&kB:ab[cdef:;H79xS
\Hy(zA*1q-*r13Ex;13Ez[*
1H79|s2JA*|13ES\Bnjps8
[*Q%tu9:Hy(x;13E8y(uz[l
Xhmf089ZABab[cdefd&kB/
@xD.C3:H<*NMZ[Ws:Hy*ogU
AS\Hy(x;H79l&kB*)k‘@ef
:1H79LnjZ%s2r>A! ) #$#-c*|9
;13ELnj{s2r>A! " #$#%cl‘@ef
d!NI;13E:w‘ALnj|*! " #$#%c*o
a3~@NI1q-*8er13EAo -c,NI
Rhi*boa3~@8e|13E,N{1qN
V*8e;13E*z[}Vj8e\H<:0Z<
*lw‘*d&k*‘@cSC3HNe7?@X
6*96H79|Lnj{s2r>Ao *c*gC!
s2{Bnj*5l*d&k*‘@efrZz[:
‘Ax9:Hy(*ogUWA:\]z[AP(l
Lab[cdS|*&kB:‘@efrZC
[:r13E*1tyVx;13E*VrZCs:
9:Hy(x;13E8y(*Xh89c‘@e
fdLO({1@4wxH\]‘*ZpsyF*
NMogU|ab[cdC[:S\Hy(ld&
k*.%9*‘@ef:‘AA*.$/ +,cuwab[c
defA-%$% +,c: -$%vX|d !%|*‘@:‘
AuwA%!$0 1 *$/c+,*$ab[cdA**$- 1
’$*c+,: *$& v*GABab[cdld .s! %
<*‘@ef:9:8yAVwA.%$. 1 .$-c+,*$
ab[cdA/$/ 1 .$’c+,: %$. vXw‘*&kB
‘@ef:S\Hy($ab[cd: -$!vl{
1j*Lab[cdS|*&kJKCmN‘@:H
y*X$01{1@4zdBO({1&:R%W
5~ol
23‘@xab[cd:ef4\&+~HJ
KogUo0:mf0*s8[|13Ec1q=
3*kBH:,dx/@*‘@efc~@N{1j
klY8[.H3E*|^B&+:H\]Z}**
h@4u!\Hy(psAB‘@*^ 4‘Bab[cdef
:0i0C!A&562*-000cl
Lab[cdS|*‘@efgB10P*H
y*H79&9?@:^g0BWvX7[Bab
[cdef*‘@ef:oL^g0v:tuv$
ab[cdef: -$/ v*5l*^ Y8w*‘@:
oL^g0ABab[cd*9\JK:mf<*
W!l@3DEopq19:+:"7|19:y
:"7rZCA:oL^g0A2344353678 "# $% $*
*###X9343:38; "# $% $**##-X<=>7?5334 "# $% $**##/cl
‘@ef:oL^g0A^<^$5w(:19:
|ab[cd+l
1l:=}$"7mH0x0H0:QLo
gl‘@efd!wHJK+:1jkl*1tl
rZs2r>ldnj*‘@efdd;]^:1
qL)z[*‘@:1jklx1tlz[*-1q
y)+t3UVz[X-/q##|{*‘@:1jkl
|38A %#@Y8*1tl|38A -#@Y8*1q<
={+9PIA4#.2**##/c*X4mfmn!H:f\1q:%$VNI+?}>lS
3*ab[cdefd!wHJK+:1jkl=
}!s2A! " #$#%c*?+!<.$Z1jkl:
=}l5l*ab[cdef:1qdnj%$*1q@A^{gUB|A*VQ@A17Hb
!V:}sMN~*d&+1q{g](AA&56
2*-000cl‘@xab[cdef1qL):s
2r>*^ ) . +,Y8*$ab[cd:1tyVAu #$* +,c
: -%v|{l
d e , -
4#.*CDE*i 8*2 $ *##/$ ‘@ef:1jklx1tl!wHJK:8fLmf $ &’()DE*-0A%cq/’& A /%*$
&56*IE2 $ -000$ !wHJK:ab[cdef:HImf $
HI)**-0A.cq’** A ’*/$
O~u*A8F $ *##-$ HG_+)45{0,1lx1(O:L
) B HI)***-A-cq/! A &’$
--! 0# 4#.2q!wHJK+‘@Lab[cdef:oL^g0
NO2 ! "#$$! O({1&HI)DE % R^qYZH9[)U4
w*&’( ) &*+ %
,-./01 2 3 4*567189: ; 2*<9:6=1 > 2*!" #$ % "##$ % ?.71@9- A1 697B
-/:CD/C:9 A1 :967/A01 /0 D:0E1 F0-A/A01 718 /:99 -AG9 0B %!"&$# ’#’()*+!)#
EA/.A1 BA:9H0:A@A1 -/718- 0B A1/9:A0: D987:H.9I60DJ% ?7178A71 K0C:176
<0/71=**LA*cq""$+ ) ""$* %
?.7G801 M ;*297:D= M*;99 N% "##L % 2.0/0-=1/.9/AD :9-F01-9- 0B /:0FAD76
B0:9-/ F671/- /0 D01/:7-/A1@ 6A@./ 91OA:01I91/- PP 3C6J9= 4 4% Q:0FAD76
B0:9-/ F671/ 9D0F.=-A060@=% R9E S0:Jq?.7FI71 718 T766*( ) ((%
U66A- , M*TCVV966 4 2*20/OA1 ?% W+++ % X1 -A/C BA968 I97-C:9I91/- 0B
F.0/0-=1/.9/AD :7/9- 0B /:0FAD76 /:99 -F9DA9-q7 /9-/ 0B /.9 BC1D/A0176
@:0CF .=F0/.9-A-% ?7178A71 K0C:176 <0/71=**$A"+cq"&&L ) "&’*%
;7A X ;*4D.7:: T*Y:7C-9: ?*!" #$ % W++( % ;97B @:0E/. 8=17IAD- 0B /E0
D01@919: @=I10-F9:I /:99 -F9DA9- :9B69D/ /.9 .9/9:0@919A/= 0B 6A@./
A1/91-A/A9- @AO91 A1 /.9A: 17/C:76 9D060@AD76 1AD.9% 2671/*?966 718
U1OA:01I91/*W$A"Wcq"’#L ) "(+( %
3767O7-A Z ?*3767O7-A 3 3% W++" % ;97B D.7:7D/9:A-/AD- 718 D.60:0F.=66
D01D91/:7/A01 0B ,-.(/0$01*&2 ’#)#.(1&2 718 3(2!4#!# 5"*10-#)’#
-9986A1@- @:0E1 A1 8ABB9:91/ 6A@./ :9@AI9-% Q:99 2.=-A060@=*W"A"+cq
*+" ) *+& %
RAA19I9/- Z*YC66 5*Q91.C191 K N% "##$ % ,1 7176=-A- 0B 6A@./ 9BB9D/- 01
B06A7: I0:F.060@=*718 6A@./ A1/9:D9F/A01 A1 /9IF9:7/9 89DA8C0C- E008=
-F9DA9- 0B D01/:7-/A1@ -.789 /069:71D9% Q:99 2.=-A060@=*"$A"+cqL$" )
L#L%
R087 T*3C:70J7 T*[7-.A/71A ;% W++’ % 30:F.060@AD76 718 F.=-A060@AD76
7DD6AI7/A01 :9-F01-9- /0 D01/:7-/A1@ 6A@./ 718 E7/9: :9@AI9- A1 6)*2&$#
5*!10$7** % UD060@AD76 M9-97:D.*"#A&cq&&" ) &’+%
5-787 R*Q7J987 T*YA/7\AI7 Y% W++& % ]C1D/A0176 D0::967/9- 0B 697B
89I0@:7F.AD :9-F01-9 /0 @7F :9697-9 A1 -7F6A1@- 0B -.789H/069:71/ /:99*
U67/9:A0-F9:ICI /7F0-% 59D060@A7*"&*AWcq"$" ) "$*%
297:D= M [*3C:70J7 T*^766787:9- ]% W++( % ?:0E1 7:D.A/9D/C:9 A1 -C1
718 -.789 91OA:01I91/-q7--9--A1@ BC1D/A01 718 /:789H0BB- EA/. 7 /.:99H
8AI91-A0176 -AIC67/A01 I0896 % R9E 2.=/060@A-/*"LLA&cq*#" ) $++%
M0G918776 N 3 ,*TC:/780 ^ T*200:/9: ;% W++L % 267-/ADA/= A1 697B /:7A/- 0B
&$ /:0FAD76 /:99 -F9DA9- A1 :9-F01-9 /0 6A@./X:967/A01-.AF- EA/. 6A@./
89I718 718 78C6/ -/7/C:9% ]C1D/A0176 UD060@=*W+AWcqW+* ) W"L%
^766787:9- ]*[:A@./ 4 K*;7--0 U*!" #$ % W+++ % 267-/AD F.910/=FAD :9-F01-9
/0 6A@./ 0B "L D01@919:AD -.:CV- B:0I 7 F717I71A71 :7A1B0:9-/ % UD060@=*
$"A*cq"#W( ) "#&L %
[9A\-D.989 K*37:/A1J0O7 K*Y:001 T*!" #$ % W++L % 4.789 7O0A871D9 A1
8)*+0$*&2 )!’!45qD0-/- 718 V919BA/- 0B F67-/ADA/= A1 F9/A069 691@/. 718
697B -AG9% R9E 2.=/060@A-/*"*WA’cqL(( ) LLL %
!"#$% &’( )*+,
stuv !"#"wxyz{|}~!
I&’YQJtK"#*L$LM*$^HZ/&’YQJtFxYZj[&’ZlB*vw1#:C>?
yN5n:&’(lM7l#TC3P#M-‘|dOlwBC(lP*#M*BC&’P*#M*pQ
I?,KH?NJ9KRny7#MlbMNSBG&RkMN[Tf*G&MN*&UDE*’V&lB*HIWX*&e’N5*{PJY2ZjlbMYlB
0*TP0*Lq0wWt*rZW!:B_0*V[0x^\0*$x[NQ&’He*(D*]):(l4
"#xp_!^Y.:,"\Kl
bMw^%M*,’ Nb*_r%boM< ?R (& ) ""*WP4l
W+"+$R# "&+‘*R#0; "+!*T$ L+!X8M Wa*T$ ’+!Xv_ "++!l^;b?R#5n*-f
7*cdwxLSef*efr%Yh3^~@gQYhnijlcdx[&’(l3klmne*81R
op.*^ cdwxLSMqxr*81;zl
"#$%6&#HZ/spt,u[3v_w W$#<
HZ/&’YQJtFI&’YQJtK;b?
’(c)%*#x y " 2#L(++("
+ , - .#sptzn^urn / +#W"#W+"+’+++"##&
- . , 0#HZ/&’YQJtF
1 2#A+$*"c&&"$&’*X&&&W(&$ 3 4#A+$*"c&&"$&’*
$H%&’(q=1BAF_OAF%"L& % D0I )**+qPP6=8D@.% F9:A08AD76- % 19/ % D1P
W" & ’ ( ) ’("