全 文 :书!"#$%&
,2016,36(5):0902-0909
犃犮狋犪犅狅狋.犅狅狉犲犪犾.犗犮犮犻犱犲狀狋.犛犻狀.
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E F:eBCfgh>ijklmnkβopqr,s,tuvwxiyz{|}、~A、>
。
,2kβopqr,s(20J,(Jl,yz{
|} 74.0%l86.6%
¡,a¢qr,yz{|} 85.8%
¡,k£v¤#$βopqr
,yz{|} 81.5%
¡。~A,kβopqr(¥¦4§¨,©ªβop
qr,s>
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«¬kβopqr,s(®iªk+¯,s¨R°:±l²³R°:±,´ k
犜犝犅11l犜犝犅12µ¶·¢¸¹ºR°»G¼½。,s^,e(J¾¿>¨ÀÁ
Â}
,
tÃR°,su YĬ¨ÀWÅÂÆ。
kβopqr,s(JyzÇÈ{|}、
JÉÊwËÌH
、
ÍÎ}µ¶ÏÐiÑÒÓ£KÔÕÇÖ×}
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ØuÍ/KQÙ#$K
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犆犾狅狀犻狀犵犪狀犱犛犲狇狌犲狀犮犲犃狀犪犾狔狊犻狊狅犳β犜狌犫狌犾犻狀犌犲狀犲犉犪犿犻犾犻犲狊犻狀犠犻犾狅狑狊
SUIJinkai1,RAOGuodong1,2,ZHANGJianguo1,2
(1ResearchInstituteofForestry,ChineseAcademyofForestry,KeyLaboratoryofTreeBreedingandCultivation,StateForest
ryAdministration,Beijing100091,China;2ColaborativeInnovationCenterofSustainableForestryinSouthernChina,Nanjing
ForestryUniversity,Nanjing210037,China)
犃犫狊狋狉犪犮狋:Inthisstudy,βtubulingenesof犛犪犾犻狓犿犪狋狊狌犱犪狀犪 Koidz.and犛.犿犪狋狊狌犱犪狀犪var.狋狅狉狋狌狊狅犪
(Vilm.)Rehd.wereclonedandidentified.Thesequencesimilarity,phylogeny,chromosomallocalization
andexpressionpatternswerefurtheranalyzed.Theresultshowedthateachwilowcontained20βtubulin
genes,andeachgenesharedmorethan74.0%and86.6%nucleotideandaminoacidsequencesimilarity
withoneanotherineachwilowspecies.Therewasover85.8%aminoacidsequencesimilarityoftheβtu
bulinbetweenthetwowilows.Whiletheaminoacidsequencesimilarityamong犛犪犾犻狓andotherplants
werelargerthan81.5%.Inthephylogeneticanalysis,βtubulinproteinsformedfourclasses,anditwas
presumedthat犛犪犾犻狓βtubulingenefamilieshadundergonesalicoidduplicationandtandemduplicatione
ventscombinedwiththestudyofchromosomallocationofpoplarβtubulin.However,犜犝犅11and犜犝犅12
werepossiblyderivedfromsegmentalduplicationorreplicativetransposition.Theexpressionpatternanal
ysisshowedatissuespecificityofβtubulingenesinwilows,andasectionofduplicatedgenepairsshowed
differentexpressionpatternsintestedtissues.Highdegreeofsequencesimilarity,evolutionaryexpansion
andofmembersandexpressionpatterndiversityof犛犪犾犻狓βtubulingenefamiliesmightconferflexibilityin
celdivisionandgrowthwhichisofimportantsignificancetothedevelopmentandgrowthhabitofperenni
alwoodyplants.
犓犲狔狑狅狉犱狊:wilow;βtubulin;phylogenetic;sequencesimilarity;expressionpattern
op(MT)ÝÞαopqr(TUA)lβopq
r
(TUB)ßÂàáâ,¤ÝãK$ÑÒä
åR8¨
,
ÑÒæÓ
[1]
lçÉ
Ó
[2]
W¾R8Fè
,
#$opäåé£êEë
ìL
[3],
é£ÑÒíîI
[4],
t #$ÑÒïí
ðñWòLóôõoóôö÷
[5]。
sø
,
BC
opqrµ¶ùú¸û#$íîI»#$
üíîíýþÿ
。
!"^#$
、
#$
、
$KK$lã%W
TUAlTUBqr,yz{|}Ç88%
¡
[612]。
&1¦(
,
)Í#$opqr,s*
¥fg»h>
,
+,[-6犜犝犃[13]l9
犜犝犅,s[14],.ª(犘狅狆狌犾狌狊狋狉犻犮犺狅犮犪狉狆犪Torr.
&Gray)8犜犝犃 l20犜犝犅,s[15],/0
1ª
(犘狅狆狌犾狌狊狋狉犲犿狌犾狅犻犱犲狊Michx.)8犜犝犃l
20犜犝犅,s[15],23(犣犲犪犿犪狔狊L.)6µ
45犜犝犃,s[16]l86¶犜犝犅 ,s[17],
78
(犗狉狔狕犪狊犪狋犻狏犪L.)W4犜犝犃,s[18]l8
犜犝犅,s[19]9。
ÙBC
WÁjk
(犛犪犾犻狓犿犪狋狊狌犱犪狀犪
Koidz.)v:mnk[犛.犿犪狋狊狌犱犪狀犪var.狋狅狉狋狌
狊狅犪(Vilm.)Rehd.][20]¦;<,fgi2k
犜犝犅,s,tufg,swxi¨ÀÁÂ}
,
wxil,yz{|}l~A
>
,
¦QÙ#$opqrBC
=>i¿>?@,A
。
1 ;
,sfg
jk£mnk7@CDEF¦RNA
=F;<
;
TG>ÊPCRTUYè¿/KHI;
<¸/\¡JK
,
KÔ¸"Z#$L
。
MNk
(犛.狊狌犮犺狅狑犲狀狊犻狊W.C.ChengexG.
Zhu)犜犝犅,sDðÙOBlastPQ,v,s¨ÉR
ST¸http://115.29.234.170/wilow/;.ª
犜犝犅,sST¸.ª¯,s¨ÉRU(https://
phytozome.jgi.doe.gov/pz/portal.html);,[-
犜犝犅,sST¸,[-¯,s¨ÉRU(http://
www.arabidopsis.org/);78、23、ÇV(犛狅狉
犵犺狌犿犫犻犮狅犾狅狉L.)、áWXYZ(犅狉犪犮犺狔狆狅犱犻狌犿犱犻狊
狋犪犮犺狔狌犿 (L.)Beauv.)l[\Z(犛犲狋犪狉犻犪狏犻狉犻犱犻狊
(L.)Beauv.)9#$ 犜犝犅 ,sDð Blast]^
GenbankÉRUPQ。
1.2 ST
1.2.1 45犜犝犅,:=>?
.ªβopq
r,yz¦_`Blast]^GenbankÉRU,
ab P Q 20 c d k (犆犺狅狊犲狀犻犪犪狉犫狌狋犻犳狅犾犻犪
(Pal.)A.Skv.)犜犝犅,s。Þ¸犜犝犅,s
$ef})Ç
,
jk
、
mnk£cdkgah
¸ªk+kh#$
,
søÙBC
cdk犜犝犅 ,
s¦i
,
jè NCBI kò$lm<¾(ht
tp://www.ncbi.nlm.nih.gov/tools/primerblast/
index.cgi?LINK_LOC=BlastHome)lmò$,f
gjk£mnk犜犝犅,s。jèEASYspin#$
RNAno=Fpqr("ZstuK$+vwx
y
)
lSuperRTcDNAz¿{©pqr("Z|¦
}~K$+vwxy
)
wxjk£mnkRNA
=FlcDNAz¿{©,PCRfgJè2×EsTaq
MasterMix("Z|¦}~K$+vwxy)wx。
Yèò$+1Y。fgYPQCDNAEºÞ
"ZW+=K$+vwxy¬y
。
1.2.2 犜犝犅,:;
(CDS)
,yz,£cdk、MNk、.
ª
、
,[-
、
78
、
23
、
ÇV
、
áWXYZl[\Z
9犜犝犅,swx$l$l,
yzu
,
犜犝犅 ,s( a¿$a
Jyz{|}
,
a$Wef}
。
1.2.3 犜犝犅89=WXYZCD jèClustalW
k£v¤#$TUB,yzwxyz
u
,
è Mega5.0±ßI~A,bè
B
(NeighborJoining,NJ),µEÈĬ)
(bootstrap)¦1000R°,,bè
JTT(JonesTaylorThornton)[21],S
?]B¦¯
。
1.2.4 犜犝犅,:=[\]^_ jè,s(¯
,s¨><¾ WGMapping(http://bioinformat
ics.psb.ugent.be/plaza/versions/plaza_v3_dicots/
3095" 2-3,9:kβopqr,s(fglyz
`1 45β6789,:ab
Table1 Primersforβtubulingenesinwilow
,sGene
ò$Primer(5′→3′)
¡Forward SReverse
犜犝犅1 TTCCGCTCATCCTCTCTGTCT CCAGCAACTGGAAGTTGAGGT
犜犝犅2 CTTCCATCTGCCCGACACTCTC CCAGCAACTGGAAGTTGAGGTT
犜犝犅3 ATGAGGGAAATCCTTCACATCC CAGAACCAATTTGAAAGGATGC
犜犝犅4 CACTGCATCCTCTCCTCGAC GAAACAAGCAAAGCAACATTGC
犜犝犅5 TCTCTCTCCATTTCCTCTGC TGTCTACCTCGATCACAACC
犜犝犅6 ATCCCACCCCGTCCCTGCAT GGCCTACTTCTGCCTGGAAT
犜犝犅7 CACGTTCCTTCCTCTTCGAT GAACCCACAACCCATTCACAT
犜犝犅8 TATACGCTCTCATCCGTTCCTT GGAGTCACAAAATCCCATCTTC
犜犝犅9 CCATTCCATTCTTCCTCACTCTC CCTCATCGACGGCGGCATCTTG
犜犝犅10 CGCAATCTTCTACTCTCTCTAC GCCAAGCCATAACCACAGCCTG
犜犝犅11 ATGCGAGAGATCCTTCACATCC CTACTGATAGTCTTGCTGGTAT
犜犝犅12 CTTAAGATCCACCAAGAACACC CAGAAACATAATCACTCACTCGC
犜犝犅13 ATCGGATCTGATGTCTCTGTGC GTAGCAGGACCGCTCATACTTC
犜犝犅14 ATGAGAGAAATCCTTCACATTC GTAGCAGGATCGTTTACACCTCC
犜犝犅15 CTCTTACCATTAATTCGATCAT CCAACCCTACTTGCAATCCAGTG
犜犝犅16 ATGAGAGAAATCCTTCACGTTC ACTTCTCGCATCAGCCACACGG
犜犝犅17 ATGAGAGAGATCCTTCACGTGC ATACCTGTTGACATTACCGTGC
犜犝犅18 GTGTGGTGCTGATTGATAGT CTTTATCTCCTGGTGAAGTTC
犜犝犅19 ACGAGATTCGTTCCTCTCTG CTAACGAAATGACCCTGTGG
犜犝犅20 TCGCTCGTTACCCATTTCCTTG GTGACAGCCACGTAGTATTAGAT
genome_mapping/index)犜犝犅 ,s v
¡>
。
1.2.5 45犜犝犅,:;<=cd^e犘犆犚 j
k
(Sm)£mnk(Smt)¿/KHI,(S1)、W
º
(S2)、¡Ô¹(S3)l(S4)4
=F RNA,jè SYBRFastqPCR Mixpqr
(Takara)wxTG>ÊPCR,犜犝犅 ,s 2
k¿/KHIa
。
2 £
2.1 犜犝犅,:;
J5
(ORF)ÔÈ¡¢¦1335~1353
bp,yz{|}¦74.1%~99.5%l
74.0%~99.9%。£¬qrà,yzÔȦ
445~451。jkW犛犿犜犝犅2l犛犿犜犝犅11,s4
5,yz{a
,犛犿犜犝犅7l犛犿犜犝犅1245
,yz{a
,
v¤J,yz{|
}¦86.6%~98.8%。mnkW 犛犿狋犜犝犅7l
犛犿狋犜犝犅12,s45,yz{a,v¤J
,yz{|}¦87.0%~99.5%。¤*
kTUBqref})Ç,¥vC¦§¨©v¤
#$¿ÎÝÍ:
。
jk SmTUB4、SmTUB6、
SmTUB10、SmTUB1415、SmTUB17、SmTUB19
20£mnkª~a¢qr,yz¯¿«,
v¤ª~a¢qr,yz{|}¦98.0%~
99.7%;¬~a¢qr{|}¦85.8%~99.7%。
kTUB£v¤¥ý#$βopqrwx
,yzu
,
^ TUB ¥ý#$W®e
f
。
vW
,
jkSmTUB4、SmTUB14lSmTUB19
£cdkª~a¢qr,yz¯¿«
;
SmTUB14£.ªª~a¢qr,yz
¯¿«
;´
MNk TUB(J ]^ðñW
¯PQi12I°yz,±^£jk¯¿«
TUBa¢qr。jk£¡²3ªk+#$
v¤TUBª~a¢qryz{|}³´¦
95.9%~99.7%、91.5%~99.7%l90.5%~
99.7%,¬~a¢qryz{|}´¦85.8%
409 ! " # $ % & 36µ
~99.3%、85.6%~99.1%l81.5%~98.8%。{
¶S
,
jk£v¤¥ý#$+,[-
、
78
、
23
、
ÇV
、
áWXYZl[\ZW TUBa¢qr
,yz{|}·¸
,´
¦86.0%~97.1%、
86.8%~97.7%、86.6% ~97.3%、87.0% ~
97.3%、88.8%~96.8%l88.8%~96.6%。
Þ¸ljk¹º»~®¼
,
mnk£v¤#
$TUBa¢qryz{|}£jkW½¾,
Ù{a
。
¤£cdk¯¿«ª~a¢qra
ΦTUB4、TUB14lTUB19;£.ªTUB14
ü¿TUB18¨¯¿«;£MNkaα
^¿«TUBª~a¢qr。mnk£
¡3
#$v¤TUBª~a¢qryz{|}
´¦95.9%~99.7%、90.9%~99.5%l90.5%
~99.7%,¬~a¢qryz{|}´¦
86.0%~99.3%、85.8%~99.3%l81.8%~
98.8%。ajk¿Î,mnk£,[-、78、23、
ÇV
、
áWXYZl[\ZW TUBa¢qr
,yz{|}¨¸¸ªk+#$
,
yz{|}É
R´¦ 86.2% ~97.3%、87.0% ~97.7%、
86.8%~97.3%、87.2%~97.5%、89.0%~96.8%
l89.0%~96.6%。
2.2 犜犝犅89WXYZCD
TUBqr( wËðñW®iÀÌ
H
,
vWÁÂÃýD#$¯,s¨°ÿ:±
、
ªk+
#$¯,s¨°ÿ:±
²³R°
,
ÄÅQª
k+4$jk、mnk、cdkl.ª
犜犝犅,sw˦^ 20。
~A
,TUBqr(µ¦
Æ\Ç
(
È1),Ø£Oakley9[15]BC¿«。z¿
ÇÁÂClassI£ClassIlike,vJÄÍ,ɯ
yz41.13%(51/124)。Ø¿WÁÊ
SmTUB、SmtTUB£CaTUB8,ËÉ)
JÉ¿Ì
,´ PtTUB³ÍοÌâÇ ø
W
。
Þ¸ÏÐßÑÒ
,3k2
TUBqr(SmTUB9~10、SmtTUB9~10£ Ca
TUB9~10)¥Ó ClassIlikeW,.ªW4
TUBqr(PtTUB9~12)Ô¸øÇ。øü,Class
IWÁÊ278ÞÕýD#$TUBqr¨
Ö
,
vWÁÂ12ÕýDTUBl1,[
-TUB(AtTUB6)。
záClassⅡJÉ׸ClassI,YÉ
ئ33.87%(42/124)。vW\¦ÃýD#
$TUBqr,ÁÂ6jk(SmTUB1、SmTUB2/
11、SmTUB3~6)、7mnk(SmtTUB1~6l
SmtTUB11)、7 c d k (CaTUB1 ~ 6 l
CaTUB11)、6ª(PtTUB1~6)、5MNk
(SsTUB1lSsTUB3~6)£3,[-TUB(At
TUB2、AtTUB7lAtTUB8);ÕýD#$TUB¯
2 3
(ZmTUB1)、Ç V (SbTUB2)、7 8
(OsTUB2)láWXYZ(BdTUB2)1,4Õ
ýD#$TUBÙ¿。
zÚlzÆJÆÛ
,
¦16l19,
ÏÐߨÆÜÕ
,
vWÕ
、
ÃýD#$TUBÝ
ÝâÇ
,
ÃClassⅢ£ClassⅣW2,[
-TUB)¿,¦ AtTUB1、AtTUB5l
AtTUB4、AtTUB9。ClassⅢWÁÂ10ªk+
TUB,jk、mnk、cdk、MNkl.ªqr
2,ÝSmTUB19~20、SmtTUB19~20、
CaTUB19~20、SsTUB19~20lPtTUB19~20,Ø
Þqr¿ßaÁSÝÝ vyzz39¡
v¤TUBqrÍi3,à,,´ v¤qr
39vἨÝÆÍ:。Ù4
ÕýD#$ TUB¦ OsTUB8、SbTUB6、Zm
TUB3lZmTUB4。ClassⅣWÁÂ2jkTUB
(SmTUB7/12 l SmTUB8)、2 m n k TUB
(SmtTUB7/12l SmtTUB8)、2 cdk TUB
(CaTUB7/12l CaSmTUB8)、2.ª TUB
(PtTUB7 l PtTUB8)l 1 M N k TUB
(SsTUB8),ß9ªk+TUBqr。âü¿ÁÂ
378、3ÇV、123l1áWXYZß8
ÕýD#$TUBqr。
2.3 犜犝犅,:;<[\]^_CD
犜犝犅,s>,20犘狋犜犝犅,s
ã äðÌÉ¡
(10/19)。vW1ë
¡ã犜犝犅,sÄÍ(6);vÝ9ë
,
ã3;3、6l16ëã2
,´
vå
(2、5、11、12l15ë)¡1
。
.ª¯,s¨¬y
[22]
æªçÛ®i
Ú¯,s¨R°:±
,
è·Í¹ºR°
、
²
³R°l¼½
。
uļ¿ªk+¯,s¨°
ÿ:±èªéêÈë
,
^20ª
犜犝犅,sÝ®iÙ°ÿ:±,tÃ> R°
¹º2,sÝ¥eìS·(È2)。vW,犜犝犅3
l犜犝犅9、犜犝犅4l犜犝犅10、犜犝犅13l犜犝犅15
犜犝犅14l犜犝犅1694u,s¿Ki²³R
°
,
*´¤íwË®ît{a
。
s¦
,
~A
2u²³R°,sW犜犝犅3l犜犝犅4
5095" 2-3,9:kβopqr,s(fglyz
ïðJTTSñòÕóìôõ;SmTUB1~SmTUB20,ÙBCfg20jkTUBqr;SmtTUB1~SmtTUB20,
ÙBCfg20mnkTUBqr;SsTUB1~SsTUB6、SsTUB8~SsTUB10、SsTUB14lSsTUB8~SsTUB10,ÙOblastPQ
14MNkTUBqr;CaTUB1~CaTUB20,20cdkTUBqr;PtTUB1~PtTUB20,20.ªTUBqr;AtTUB1~
AtTUB9,9,[-TUBqr;OsTUB1~ OsTUB8,878TUBqr;ZmTUB1~ZmTUB7,723TUBqr;SbTUB1~
SbTUB6,6ÇVTUBqr;BdTUB1~BdTUB5,5áWXYZTUBqr;SvTUB1~SvTUB2,2[\ZTUBqr
È1 #$TUBqr~A
ThebarmeansunitgeneticdistanceestimatedbyJTTmodel;SmTUB1~SmTUB20,20犛.犿犪狋狊狌犱犪狀犪TUBproteinsclonedin
thisstudy;SmtTUB1~SmtTUB20,20犛.犿犪狋狊狌犱犪狀犪var.狋狅狉狋狌狊狅犪TUBproteinsclonedinthisstudy;SsTUB1~SsTUB6,
SsTUB8~SsTUB10,SsTUB14andSsTUB8~SsTUB10,14犛.狊狌犮犺狅狑犲狀狊犻狊TUBproteinsobtainedbylocalblast;CaTUB1~
CaTUB20,20犆.犪狉犫狌狋犻犳狅犾犻犪TUBproteins;PtTUB1~PtTUB20,20犘.狋狉犻犮犺狅犮犪狉狆犪TUBproteins;AtTUB1~AtTUB9,nine
犃.狋犺犪犾犻犪狀犪TUBproteins;OsTUB1~ OsTUB8,eight犗.狊犪狋犻狏犪TUBproteins;ZmTUB1~ZmTUB7,seven犣.犿犪狔狊
TUBproteins;SbTUB1~SbTUB6,six犛.犫犻犮狅犾狅狉TUBproteins;BdTUB1~BdTUB5,five犅.犱犻狊狋犪犮犺狔狅狀TUBproteins;
SvTUB1~SvTUB2,two犛.狏犻狉犻犱犻狊TUBproteins
Fig.1 PhylogeneticanalysisofplantTUBproteins
»~Ƽ
,
aÎ犜犝犅9l犜犝犅10¨ a¿
¡
,
Rø«¬¤íÝö®i²³R°:±
,
Þ¿
÷ö,sw˦犜犝犅3/4l犜犝犅9/10,*è
Kiªk+,s¨R°:±Q2,søùç4
:犜犝犅3、犜犝犅4、犜犝犅9l犜犝犅10。{ú,è2u
,s³Ý 犜犝犅13l 犜犝犅15wË»~Ƽ,´
犜犝犅14l犜犝犅16 a¿,søµ
«¬¤í
öÞ¿÷ö,sð,s¨R°:±wË
609 ! " # $ % & 36µ
犜犝犅13/15l犜犝犅14/1692,s,ûègK
²³R°ü,søù
,
í4,s。
ßk}æk£ªßÇÈ{
|
[23],
Rø«¬k犜犝犅,s(aήiÚ
¯,s¨R°:±l²³R°:±
,
L«v(
Jýþ&20。*´,k犜犝犅11l犜犝犅12
,s
(
È2)wˮa,ªPtTUB11
ÿkéÝ犜犝犅狊°ÿ,s,TkéÝk犜犝犅11
l犜犝犅12
¤íµ¶R°,s。
È2 犜犝犅,s>£,sR°:±
Theduplicatedparalogouspairsof犜犝犅狊areconnected
wi thdottedlines.犜犝犅11and犜犝犅12from犛犪犾犻狓areconnected
withtheirpotentialduplicatedparalogouswithsolidlines
Fig.2 Chromosomallocationandgeneduplication
eventsof犜犝犅genes
lPtTUB12ßaâÇ ClassIlikeW,´ k
TUB11!£TUB1lTUB2ßaâÇ ClassⅡ
W
,TUB12£TUB7,yz{a,t£ TUB8
âÇ Class Ⅳ W。s ø,£ ª a,k
犜犝犅11l犜犝犅12Ý·¢¸a¿÷ö,s,¤
íµ¶ÝÞ犜犝犅1/犜犝犅2l犜犝犅7Dð¹º
R°»¼½"K
。
2.4 45犜犝犅,:`ijk
¼ÌÉ
(9)k犜犝犅,s YĬ¨ÀW
,
vW\ÝR°,s
。ClassIW¯Ä¬
&犜犝犅13l 犜犝犅18 jklmnkHI,
(S1)、W(S2)、Ô¹(S3)l#§(S4)4¨ÀW
(
È3),Ã{u7$Ý)¸(ÄÇ{u
ʦ0.007121)。 jkW,犜犝犅13l犜犝犅18
HI#§7$ÄÇ
,
,¶
,
WlÔ¹
Ƹ
;´
mnkW
,犛犿狋犜犝犅13 HIÔ¹
7$ÄÇ
,
vÝ#§K¹
、
,lW
,犛犿狋
犜犝犅18HI#§lÔ¹7$ÝÆÇ,v
Ý,lW
。
F¦犜犝犅13R°,s,犜犝犅14、
犜犝犅15l犜犝犅16% ¡²¨ÀW;犜犝犅17
F¦犜犝犅18R°,s¨。ClassIlikeW
2,s犜犝犅9l犜犝犅10¨ ¡²¨ÀW
,´
¤íR°,s犜犝犅3l犜犝犅4(ClassⅡ)
¡²4¨ÀWÝ。
ClassⅡW犜犝犅11Ý¡²4k¨ÀW
78,s
(
È3),{u7$% 0.25
Sm.jk;Smt.mnk;S1.HI,;S2.HIW;S3.HIÔ¹;S4.HI#§
È3 k犜犝犅,s
Sm.犛.犿犪狋狊狌犱犪狀犪;Smt.犛.犿犪狋狊狌犱犪狀犪var.狋狅狉狋狌狊狅犪;S1.Basalregionofbranch;
S2.Middleregionofbranch;S3.Elongationregionofbranch;S4.Apicalregionofbranch
Fig.3 Theexpressionprofileof犜犝犅inwilows
7095" 2-3,9:kβopqr,s(fglyz
¡,
jkHIÔ¹7$ÄÇ
(
{uÊ
¦9.05),´ mnk{&¨ÀW7$æ
¸¸jk
(
{uʦ1.80),e,s 2k
#§W7$¨ÆÇÃ7${
,´
HIWl,7$Ƹ
。犜犝犅1l犜犝犅2
F¦犜犝犅11R°,sÝ犜犝犅11üÊÆ
Ç,s
,犜犝犅3l犜犝犅4¶。´jk犜犝犅1~4
HIWl#§7$%Ǹmnk
,
Ô
¹7$£mnk{
;犜犝犅2 mnkHI,
7$Ǹjk
。犜犝犅5l犜犝犅6 (k
¡²4¨ÀW%)。
ClassⅢW¯犜犝犅19(È3),e,s
2kHI#§7$ÄÇ, mnkHI
Ô¹7$¨ÆÇ
,
v¤¨ÀW7$
Ƹ
。
)Ĭ&vR°,s犜犝犅20 ¡²¨ÀW
。Class Ⅳ W 3 ,s (Þ¸ 犜犝犅7 l
犜犝犅12yz{|}®Ç,L«*©+ò$µ
2,s7$¬>) ¡²¨ÀW
%
,
vW犜犝犅8 jkHIÔ¹7
$æǸjklmnkv¤HI¨À
。犜犝犅7/12
jkHI,
、
W
、
Ô¹l#§K¹
7$%Ǹmnk{&¨À
,
vW
jkHI
Ô¹7$ÄÇ
,
W
、
#§K¹l,´
¶
;
e,s mnkHI¨ÀW7$Ǹ
y´¦Ô¹
、
W
、
,l#§K¹
。
3 , @
DðfgPQjklmnk犜犝犅 ,s20
,
£.ªl/01ª犜犝犅,s([15]J
É{9
,
Ãv(Jl,yz{
|}¨£ª{
,
-çǸª
,
ý7$¿>
ñÈ¡.æjklmnk¹º»~)¼
,
/0iW
#$1WmnkÝjk:¿.
[20]。
jkl
mnk TUBqr(J,yzÇÈe
f
,
vC¦§£v¤#$¿Î¨ÝÍ:,BC
æTUBqrC¦§23op¨4[24],5>opÍ
ââ©67
[25],
µ¶ß8qr©
9
[26],
tÿÝ)Íop©qr:F¹;
[2733]。
sø
,
µ
«¬kTUBqrC¦§¨ vop
¨4
,
£op©qr:FW
~Al>
,
k犜犝犅
,s(aª
[22]
¿Î®i3¯,s¨R°
:±
,
è·¹ºR°
、
²³R°l¼½9,s
R°:±
。
k£ª犜犝犅11l犜犝犅12 wË
¡¹=
,
ªk+#$,s¨R°:±K
èªhlkh#$)¾awË®î
。
â
ü
,
Þ1,s(JÉÊ>?
,
ªk+#$
犜犝犅,s(wËÌHZÙ#$Õæ。
+
,
,[-W¯9犜犝犅,s[14]、23W1¯
h>@8犜犝犅,s[17]、78W¨¯8犜犝犅
,s
[19]
9
,´
ªlkÝ20犜犝犅,s。Ø
,sÌHu9µ¶uQÙ#$Í/KKÔ
Ú}ÛÜR。
k£ª犜犝犅 ,sÅÂÆ\,
ª ClassIl ClassIlike § ¨ W 犜犝犅9、
犜犝犅13、犜犝犅15l犜犝犅16ÝQàW78
,s
[15],´
k¿/KHI4¨ÀW)Ĭ&
犜犝犅9、犜犝犅15l犜犝犅16,Ã犜犝犅13
7$¨)¸
。
{¶S
,ClassⅡW犜犝犅11
vR°,s犜犝犅1l犜犝犅2 kWÊÄ
Ç
,
søk犜犝犅11、犜犝犅1l犜犝犅2Aµ¶_
iª犜犝犅9、犜犝犅13、犜犝犅15l犜犝犅166¶,2
3kQàKÔA
。
¥Ý
,
k犜犝犅12t±
©犜犝犅11¿Î k¿/KHIWÇ,=
R°:±"Kc,swËBCÝa
。
âü
,
犜犝犅13 R ° , s 犜犝犅14、犜犝犅15 l 犜犝犅16,
犜犝犅18R°,s犜犝犅17,犜犝犅3l犜犝犅4R°
,s 犜犝犅9l 犜犝犅10,
犜犝犅19R°,s
犜犝犅209,s YĬ¨ÀW,.æR°,
s wËðñWKi6¶»6¶D
。
lm!M
:
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