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Cloning and Sequence Analysis of β Tubulin Gene Families in Willows

柳树β微管蛋白基因家族的克隆和序列分析



全 文 :书!"#$%&
,2016,36(5):0902-0909
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Ž‘
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|}ŒŸ 74.0%l86.6%…¡,’›a¢qrž,yz{|} 85.8%…¡,k“£v¤#$βopqr›
‹ž,yz{|} 81.5%…¡。~€AŒ‘,k“βopqr(”¥Œ¦4—§¨,Ž©ª“βop
qr,s‚ƒ>„

«¬k“βopqr,s(”­®iªk+¯,s¨R°:±l²³R°:±,´ k“
犜犝犅11l犜犝犅12µ¶·¢¸¹ºR°»G¼½。,s‡ˆ‰ŠŒ€^,e(”˜J‹‡ˆ¾–¿>‹¨ÀÁ
Â}

tڌR°,su YĬ¨ÀW‡ˆÅÂÆ。
k“βopqr,s(”˜Jyz‹ÇÈ{|}、˜
JÉʋwËÌH

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ØuÍ/KQÙ#$‹K
ԀAÚ}ÛÜR。
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JKCL$
:Q785;Q786 !MNOP:A
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,βtubulingenesof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.Theresultshowedthateachwilowcontained20βtubulin
genes,andeachgenesharedmorethan74.0%and86.6%nucleotideandaminoacidsequencesimilarity
withoneanotherineachwilowspecies.Therewasover85.8%aminoacidsequencesimilarityoftheβtu
bulinbetweenthetwowilows.Whiletheaminoacidsequencesimilarityamong犛犪犾犻狓andotherplants
werelargerthan81.5%.Inthephylogeneticanalysis,βtubulinproteinsformedfourclasses,anditwas
presumedthat犛犪犾犻狓βtubulingenefamilieshadundergonesalicoidduplicationandtandemduplicatione
ventscombinedwiththestudyofchromosomallocationofpoplarβtubulin.However,犜犝犅11and犜犝犅12
werepossiblyderivedfromsegmentalduplicationorreplicativetransposition.Theexpressionpatternanal
ysisshowedatissuespecificityofβtubulingenesinwilows,andasectionofduplicatedgenepairsshowed
differentexpressionpatternsintestedtissues.Highdegreeofsequencesimilarity,evolutionaryexpansion
andofmembersandexpressionpatterndiversityof犛犪犾犻狓βtubulingenefamiliesmightconferflexibilityin
celdivisionandgrowthwhichisofimportantsignificancetothedevelopmentandgrowthhabitofperenni
alwoodyplants.
犓犲狔狑狅狉犱狊:wilow;βtubulin;phylogenetic;sequencesimilarity;expressionpattern
  op(MT)ÝÞαopqr(TUA)lβopq
r
(TUB)ߘ‹Âàáâƒ,¤ÝãœK$ÑÒä
å‹R8¨˜šŒ

 ÑҖæŒÓ
[1]
lçɌ
Ó
[2]
W¾–R8Fè

#$opäåé£êEë‹
ìL
[3],
é£ÑҋíîI˜
[4],
t #$ÑÒïí
˜ðñWòLóôõoóô‹ö÷
[5]。


BC
opqrµ¶ù–ú¸û‘#$‹íîI˜»#$
üšíî혋Œýþÿ

!"€^‹#$

#$

$KK$lã%W‹
TUAlTUBqrž,yz{|}Lj88%…
¡
[612]。
&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

(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.)‹6—µ
45犜犝犃,s[16]l8—–6¶‹犜犝犅 ,s[17],
78
(犗狉狔狕犪狊犪狋犻狏犪L.)W–4—犜犝犃,s[18]l8
—犜犝犅,s[19]9。
ÙBC…WÁ–‹jk
(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]¦;<,fgi2’k“‹
犜犝犅,s,tufg‹,swxi¨ÀÁÂ}‡ˆ
Œ

wxiœlž,yz{|}l~€A
Œ…†‚ƒ>„

¦QÙ#$opqr‹BC
=>i¿>‹?@,A

1 ;1.1 QR
,sfg…jk£mnk7@CDEF¦RNA
=F;<

TG>ÊPCRTUY苿/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
rž,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$lm<¾(ht
tp://www.ncbi.nlm.nih.gov/tools/primerblast/
index.cgi?LINK_LOC=BlastHome)lmò$,f
gjk£mnk犜犝犅,s。jèEASYspin#$
RNAno=Fpqr("ZstuK$+v–wx
y

lSuperRTcDNAz¿{©˜pqr("Z|¦
}~K$+v–wxy

wxjk£mnkRNA
=FlcDNAz¿{©˜,PCRfgJè2×EsTaq
MasterMix("Z|¦}~K$+v–wxy)wx。
Yèò$+‡1Y‘。fgYPQ‹CDNAEºÞ
"ZW+€=K$+v–w‚xy¬y

1.2.2 犜犝犅,:;±„fgPQ‹jklmnk犜犝犅 ,s45¹
(CDS)…†˜ž,yz,£cdk、MNk、.
ª

,[-

78

23

ÇV

áWXYZl[\Z
9犜犝犅,swx$’›l$’™šœlž,
yz‡u

Œ犜犝犅 ,s(” a¿$’ˆa˜
J›‹yz{|}

…† ˆa$’W‹ef}

1.2.3 犜犝犅89=WXYZCD jèClustalW
„k“£v¤#$‹TUBž,yzwxyz‡
u

‰è Mega5.0ƒ±ßI~€A“,b芋
B
(NeighborJoining,NJ),ŒŒµEÈĬ)Ž
(bootstrap)¦1000R°,ž,‘‰’bè
JTT(JonesTaylorThornton)‰’[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:k“βopqr,s(”‹fglyzŒ
`1 45β6789,:ab
Table1 Primersforβtubulingenesinwilow
,sGene
ò$Primer(5′→3′)
¡™Forward S™Reverse
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)Œ犜犝犅 ,s v‚
ƒ¡‹>„

1.2.5 45犜犝犅,:;<=cd^e犘犆犚 „j
k
(Sm)£mnk(Smt)¿/KHIŒ,š(S1)、W
º
(S2)、¡ššÔ¹(S3)l›œ(S4)4—š„ŒŸ
=F RNA,jè SYBRFastqPCR Mixpqr
(Takara)wxTG>ÊPCR,Œ犜犝犅 ,s 2
’k“¿/KHIˆaš„‹‡ˆ‰Š

2 Ž£Œ
2.1 犜犝犅,:;fgPQjklmnk犜犝犅,s•20—,(
”˜JžŸ5 
(ORF)ÔÈ¡¢¦1335~1353
bp,œyz{|}ŒŸ¦74.1%~99.5%l
74.0%~99.9%。£¬‹qràž,yzÔȦ
445~451。jkW犛犿犜犝犅2l犛犿犜犝犅11,s4
5‹ž,yz{a
,犛犿犜犝犅7l犛犿犜犝犅1245
‹ž,yz{a

v¤˜J›‹ž,yz{|
}¦86.6%~98.8%。mnkW 犛犿狋犜犝犅7l
犛犿狋犜犝犅12,s45‹ž,yz{a,v¤˜J
›‹ž,yz{|}¦87.0%~99.5%。¤*
k“TUBqref})Ç,¥vC¦§¨©v¤
#$¿ÎÝÍ:‹

jk SmTUB4、SmTUB6、
SmTUB10、SmTUB1415、SmTUB17、SmTUB19
20£mnkª~a¢qr‹ž,yz–¯¿«,
v¤ª~a¢qr‹ž,yz{|}¦98.0%~
99.7%;¬~a¢qr{|}¦85.8%~99.7%。
„k“TUB£v¤¥ý#$βopqrwx
ž,yz‡u

€^ TUB ¥ý#$W­®e
f

vW

jkSmTUB4、SmTUB14lSmTUB19
£cdkª~a¢qr‹ž,yz–¯¿«

SmTUB14£.ªª~a¢qr‹ž,yz–
¯¿«
;´
MNk‹ TUB(”˜J ]^ðñW
¯PQi12I–°yz,±–€^£jk–¯¿«
‹TUBa¢qr。jk£¡²3’ªk+#$‹
v¤TUBª~a¢qr›‹yz{|}³´¦
95.9%~99.7%、91.5%~99.7%l90.5%~
99.7%,¬~a¢qr‹yz{|}´¦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‹ž
,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¢qr›‹yz{|}£jkW½¾,
Ù{a

¤£cdk›–¯¿«‹ª~a¢qra
ΦTUB4、TUB14lTUB19;£.ª˜TUB14
ü¿–TUB18¨–¯¿«;£MNk›aα–
€^¿«‹TUBª~a¢qr。mnk£…¡3
’#$‹v¤TUBª~a¢qr›‹yz{|}
´¦95.9%~99.7%、90.9%~99.5%l90.5%
~99.7%,¬~a¢qr›‹yz{|}´¦
86.0%~99.3%、85.8%~99.3%l81.8%~
98.8%。ajk¿Î,mnk£,[-、78、23、
ÇV

áWXYZl[\ZW TUBa¢qr‹ž
,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®iÀ‹Ì
H

vWÁÂÃýD#$¯,s¨°ÿ:±

ªk+
#$¯,s¨°ÿ:±…†²³R°

ÄʼnQª
k+‹4—$’jk、mnk、cdkl.ª
犜犝犅,sw˦^ ‹•20—。
~€AŒŽ‘
,TUBqr(”µŒ¦
Æ\Ç

È1),Ø£Oakley9[15]‹BC¿«。z¿
ÇÁÂClassI£ClassIlike,v˜JÄÍ,ɯš
Œyz‹41.13%(51/124)。Ø¿ŒŒWÁÊ
SmTUB、SmtTUB£CaTUB•8—,Ëɕ)˜
Jɋ¿Ì
,´ PtTUB³ÍΖ¿ÌâÇ ø
ŒŒW

Þ¸ÏЎ߈ÑÒ
,3’k“•–2—
TUBqr(SmTUB9~10、SmtTUB9~10£ Ca
TUB9~10)¥ÓŒ ClassIlikeW,.ªW–4
—TUBqr(PtTUB9~12)Ô¸øÇ。øü,Class
IWÁʖ2—78ÞÕýD#$TUBqr¨˜
‹ÖŒŒ

vWÁÂ12—ÕýDTUBl1—,[
-TUB(AtTUB6)。
zጌClassⅡ˜JÉ׏¸ClassI,YÉ
‡Ø¦33.87%(42/124)。vW\šŒ¦ÃýD#
$TUBqr,ÁÂ6—jk(SmTUB1、SmTUB2/
11、SmTUB3~6)、7—mnk(SmtTUB1~6l
SmtTUB11)、7 — c d k (CaTUB1 ~ 6 l
CaTUB11)、6—ª“(PtTUB1~6)、5—MNk
(SsTUB1lSsTUB3~6)£3—,[-TUB(At
TUB2、AtTUB7lAtTUB8);ÕýD#$TUB¯
– 2 3
(ZmTUB1)、Ç V (SbTUB2)、7 8
(OsTUB2)láWXYZ(BdTUB2)•1—,4—Õ
ýD#$TUB٘¿Œ。
zÚlzƌŒ˜JÆÛ

ŒŸ¦16l19—,
ÏЎߨ‡ÆÜÕ

vWÕ

ÃýD#$TUBÝ
݌âÇ

ÃClassⅢ£ClassⅣW•–2—,[
-TUB)˜¿Œ,ŒŸ¦ AtTUB1、AtTUB5l
AtTUB4、AtTUB9。ClassⅢWÁÂ10—ªk+
TUB,jk、mnk、cdk、MNkl.ªqr
•2—,ŒŸÝSmTUB19~20、SmtTUB19~20、
CaTUB19~20、SsTUB19~20lPtTUB19~20,Ø
Þqr‹¿—ßaÁSÝÝ vyzz39„¡‡
v¤TUBqrÍi3—ž,à,,´ v¤qr
 39„†vἨ݇ÆÍ:‹。ٌŒ‹4—
ÕýD#$ TUBŒŸ¦ OsTUB8、SbTUB6、Zm
TUB3lZmTUB4。ClassⅣWÁÂ2—jkTUB
(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),ß9—ªk+TUBqr。âü¿ÁÂ
3—78、3—ÇV、1—23l1—áWXYZß8
—ÕýD#$TUBqr。
2.3 犜犝犅,:;<[\]^_CD
犜犝犅,s‚ƒ>„‘,20—犘狋犜犝犅,s
Œã äðÌɋ‚ƒ¡
(10/19)。vW1ë
‚ƒ¡Œã‹犜犝犅,sÄÍ(6—);vÝ9ë
‚ƒ

Œã–3—;3、6l16끂ƒ•Œã–2
—
,´
v偂ƒ
(2、5、11、12l15ë)¡•–1
—

.ª¯,s¨¬y
[22]
‡æª“çÛ­®i
ڏ¯,s¨R°:±

…†è·‹Í¹ºR°

²
³R°l¼½

u‡Ä¼‹¿ªk+¯,s¨°
ÿ:±èª“‚ƒéêÈë

€^20—ª“
犜犝犅,sÝ­®iُ°ÿ:±,tÃ>„ R°
¹º‹2—,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
°

*´¤í‹wË®îtˆ{a



~€A
Œ‘2u²³R°,sW犜犝犅3l犜犝犅4
5095"              2-3,9:k“βopqr,s(”‹fglyzŒ
ïð‡‘JTT‰’Sñò‹Õ„óìôõ;SmTUB1~SmTUB20,ÙBCfg‹20—jkTUBqr;SmtTUB1~SmtTUB20,
ÙBCfg‹20—mnkTUBqr;SsTUB1~SsTUB6、SsTUB8~SsTUB10、SsTUB14lSsTUB8~SsTUB10,ÙOblastPQ
‹14—MNkTUBqr;CaTUB1~CaTUB20,20—cdkTUBqr;PtTUB1~PtTUB20,20—.ªTUBqr;AtTUB1~
AtTUB9,9—,[-TUBqr;OsTUB1~ OsTUB8,8—78TUBqr;ZmTUB1~ZmTUB7,7—23TUBqr;SbTUB1~
SbTUB6,6—ÇVTUBqr;BdTUB1~BdTUB5,5—áWXYZTUBqr;SvTUB1~SvTUB2,2—[\ZTUBqr
È1 #$TUBqr~€AŒ
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¿—Œ
Œ¡

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¿ŒŒ,søµ…«¬¤í
öÞ¿—÷ö,s­ð,s¨R°:±w˘
609 ! " # $ % &                   36µ
犜犝犅13/15l犜犝犅14/1692—,s,ûèg€K
²³R°ü˜,søù

í˜4—,s。
  ßk}Œ‡æk“£ª“‚ƒŽßÇÈ{
|
[23],
Rø«¬k“犜犝犅,s(”aέ®iÚ
¯,s¨R°:±l²³R°:±

L«v(”
˜Jýþ&20—。*´,k“犜犝犅11l犜犝犅12
,s

È2)‹wˮ“ˆa,ª“PtTUB11
ÿk鋋Ý犜犝犅狊°ÿ,s,Tk鋋Ýk“犜犝犅11
l犜犝犅12…†¤íµ¶‹R°,s。
È2 犜犝犅,s‚ƒ>„£,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  ClassIlikeW,´ k“
TUB11!£TUB1lTUB2ßaâÇ ClassⅡ
W
,TUB12£TUB7ž,yz{a,t£ TUB8
âÇ   Class Ⅳ W。s ø,£ ª “ ˆ a,k “
犜犝犅11l犜犝犅12ˆÝ·¢¸a¿—÷ö,s,¤
íµ¶ÝÞ犜犝犅1/犜犝犅2l犜犝犅7ŒŸDð¹º
R°»¼½"K

2.4 45犜犝犅,:`ijk
¼ÌÉ
(9—)k“犜犝犅,s YĬ¨ÀW
ˆ‡ˆ

vW\šŒÝR°,s
。ClassIW¯Ä¬
&犜犝犅13l 犜犝犅18 jklmnkHI,š
(S1)、Wš(S2)、šÔ¹(S3)l#§(S4)4—¨ÀW
‡ˆ

È3),Ã{u‡ˆ7$Ý)¸(ÄÇ{u‡ˆ
ʦ0.007121)。 jkW,犜犝犅13l犜犝犅18 
HI‹#§‡ˆ7$ÄÇ

,š¶

WšlšÔ¹
Ƹ
;´
 mnkW
,犛犿狋犜犝犅13 HIšÔ¹‡
ˆ7$ÄÇ

vÝ#§ŒK¹

,šlWš
,犛犿狋
犜犝犅18HI#§lšÔ¹‡ˆ7$݇ÆÇ,v
Ý,šlWš

F¦犜犝犅13‹R°,s,犜犝犅14、
犜犝犅15l犜犝犅16%ˆ ¡²¨ÀW‡ˆ;犜犝犅17
F¦犜犝犅18‹R°,s¨ˆ‡ˆ。ClassIlikeW
‹2—,s犜犝犅9l犜犝犅10¨ˆ ¡²¨ÀW‡
ˆ
,´
¤í‹R°,s犜犝犅3l犜犝犅4(ClassⅡ) 
¡²4—¨ÀWݖ‡ˆ。
ClassⅡW‹犜犝犅11Ý¡²4—k“¨ÀW
‹78‡ˆ,s

È3),{u‡ˆ7$% 0.25…
Sm.jk;Smt.mnk;S1.HI,š;S2.HIWš;S3.HIšÔ¹;S4.HI#§
È3 k“犜犝犅,s‡ˆ‰Š
Sm.犛.犿犪狋狊狌犱犪狀犪;Smt.犛.犿犪狋狊狌犱犪狀犪var.狋狅狉狋狌狊狅犪;S1.Basalregionofbranch;
S2.Middleregionofbranch;S3.Elongationregionofbranch;S4.Apicalregionofbranch
Fig.3 Theexpressionprofileof犜犝犅inwilows
7095"              2-3,9:k“βopqr,s(”‹fglyzŒ
¡,
 jkHIšÔ¹‹‡ˆ7$ÄÇ

{u‡ˆÊ
¦9.05),´  mnk{&¨ÀW‹‡ˆ7$æ
¸¸jk

{u‡ˆÊ¦1.80),e,s 2’k“
‹#§W‡ˆ7$¨‡ÆÇLj7${
,´
 
HIWšl,š‡ˆ7$Ƹ
。犜犝犅1l犜犝犅2
F¦犜犝犅11‹R°,sݘ犜犝犅11ü‡ˆÊÆ
Nj,s
,犜犝犅3l犜犝犅4¶。´jk犜犝犅1~4
 ‹HIWšl#§‡ˆ7$%Ǹmnk

šÔ
¹‡ˆ7$£mnk{
;犜犝犅2 mnkHI,
š‡ˆ7$Ǹjk
。犜犝犅5l犜犝犅6 (’k
“‹¡²4—¨ÀW%)‡ˆ。
ClassⅢW¯–犜犝犅19‡ˆ(È3),e,s 
2’k“‹HI#§‡ˆ7$ÄÇ, mnkHI
šÔ¹‡ˆ7$¨‡ÆÇ

 v¤¨ÀW‡ˆ7$
Ƹ

)Ĭ&vR°,s犜犝犅20 ¡²¨ÀW
‡ˆ
。Class Ⅳ W‹ 3 —,s (Þ¸ 犜犝犅7 l
犜犝犅12œyz{|}­®Ç,L«*©+ò$µ
…„2—,s‹‡ˆ7$Œ¬>) ¡²¨ÀW
%–‡ˆ

vW犜犝犅8 jkHIšÔ¹‹‡ˆ7
$æÇ¸jklmnkv¤HI¨À
。犜犝犅7/12
 jkHI,š



šÔ¹l#§ŒK¹‹‡ˆ
7$%Ǹmnk‹{&¨À

vW… jkHIš
Ô¹‹‡ˆ7$ÄÇ



#§ŒK¹l,š´
¶

e,s mnkHI•¨ÀW‡ˆ7$Ǹ
y´¦šÔ¹



,šl#§ŒK¹

3 , @
DðfgPQjklmnk犜犝犅 ,s•20
—

£.ªl/01ª‹犜犝犅,s(”[15]˜J
É{9

Ãv(”™š˜J›œlž,yz{
|}¨£ª“{

-çǸª“

 Œý7$¿>
ñÈ¡.æjklmnk¹º»~)¼

/0iW
#$1WmnkÝjk‹:’¿.
[20]。
jkl
mnk TUBqr(”˜J‹ž,yzÇÈe
f

vC¦§£v¤#$¿Î¨ÝÍ:‹,–BC‡
æTUBqrC¦§23op‹¨4[24],5>opÍ
⃋â©67
[25],
µ¶ß˜8ˆqr‹Ž©‡
9
[26],
tÿÝ)ÍopŽ©qr‹:F¹;
[2733]。


µ…«¬k“TUBqr‹C¦§¨ vop
ܬ4

†£opŽ©qr‹:FW
~€Al‚ƒ>„Œ‘

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
¡‹¹Ÿ=‘

 ªk+#$,s¨R°:±€K
èªhlkh#$•)¾–ˆa‹wË®î

â
ü

Þ1,s(”˜J‹ÉÊ>?

ªk+#$
犜犝犅,s(”‹wËÌH‡ZÙ#$Õæ。‡
+

,[-W¯–9—犜犝犅,s[14]、23W1¯
h>@8—犜犝犅,s[17]、78W¨¯–8—犜犝犅
,s
[19]
9
,´
ª“lk“Ý–20—犜犝犅,s。Ø
’,sÌH‹ˆu9µ¶uQÙ#$Í/K‹KÔ
Ú}ÛÜR。
k“£ª“犜犝犅 ,s‹‡ˆ‰ŠÅÂÆ\,
ª“ ClassIl ClassIlike § ¨ W ‹ 犜犝犅9、
犜犝犅13、犜犝犅15l犜犝犅16ÝQàšW‹78‡ˆ
,s
[15],´
k“¿/KHI4—¨ÀW)Ĭ&
犜犝犅9、犜犝犅15l犜犝犅16‹‡ˆ,Ã犜犝犅13‹‡
ˆ7$¨)¸

{‡¶S
,ClassⅡW‹犜犝犅11†
vR°,s犜犝犅1l犜犝犅2 k“W‹‡ˆÊÄ
Ç

søk“犜犝犅11、犜犝犅1l犜犝犅2A–µ¶_
iª“犜犝犅9、犜犝犅13、犜犝犅15l犜犝犅16‹6¶,2
3k“Qàš‹KԀA

¥Ý

k“犜犝犅12t±
–©犜犝犅11¿Î k“¿/KHIWLJˆ,=‘
R°:±"K‹c,swËBC݈a‹

âü

犜犝犅13 ‹ R ° , s 犜犝犅14、犜犝犅15 l 犜犝犅16,
犜犝犅18‹R°,s犜犝犅17,犜犝犅3l犜犝犅4‹R°
,s 犜犝犅9l 犜犝犅10,…† 犜犝犅19‹R°,s
犜犝犅209,s YĬ¨ÀWˆ‡ˆ,.æR°,
s wËðñW€Ki6¶“”»6¶ŒD

lm!M

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