小麦—族毛麦6VS/6AL易位系叶片cDNA文库构建及6VS/6AL易位系中抗白粉病相关基因的克隆
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摘要
小麦白粉病是严重危害小麦生产的病害之一，南京农业大学细胞遗传研究
所培育的小麦-簇毛麦6VS/6AL易位系带有目前我国最有效的抗白粉病基因
Pm21，此外还对我国当前新优势条锈菌生理小种表现高抗，对叶锈也表现高抗。
对Pm21及抗白粉病相关基因进行克隆研究不仅有利于阐明其抗病机制，而且
对抗病育种有重大意义。
构建cDNA文库是基因克隆的基础，本研究用表达型质粒载体p~(spon-1)构建
了经白粉菌（Erysiphe graminis）诱导48小时和未经诱导的小麦-簇毛麦6VS/6AL
易位系（Triticum aestivum-Haynaldia villosa translocation line）叶片cDNA文库
二个。文库奇主菌为E.coli DH10B。非诱导文库包含约50万个重组cDNA克
隆，平均插入片段为1.25kb，主要分布在400bp-2.2kb之间。诱导文库包含约30
万个重组cDNA克隆，平均插入片段1.2kb。用本室克隆的小麦类甜蛋白基因
pWIR作探针，进行杂交筛库，杂交信号明显，重复性好。
用mRNA差异显示技术对经白粉菌诱导24小时，48小时和未经诱导的
6VS/6AL易位系叶片mRNA进行比较，共获得35个差异cDNA片段。Northern
杂交证明，其中8个cDNA克隆，在白粉菌诱导下其基因表达增强。
测序和同源性比较表明，其中4个分别与大豆蛋白激酶GmPK6、达提斯
卡（Datisca glomerata）硫代硫酸硫转移酶，推测的小麦In2-1人类似蛋白基因和
拟南芥第三染色体上的BAC克隆F15G16序列高度同源。它们都是小麦中的新
基因。另外三个推测为小麦叶绿体的三个基因：1.5-二磷酸核酮糖羧化酶
（Rubisco）大亚基基因，Rubisco活化酶基因和RNA聚合酶rpoA基因。这些
基因可能参与了小麦抗白粉病反应。
扬麦5号和6VS/6AL易位系基因组DNA经EcoRⅠ、EcoRⅤ和HindⅢ三
种酶切，用克隆的cDNA片段作探针，杂交结果表明，硫转移酶基因（NJS22）
在三种酶切中均有差异。小麦蛋白激酶基因（NJS4）、InZ八类似基因（NJS35）
和 beclin类似基因（NJSZ）在两者间均有不同程度的差异。
以 mRNA差异显示获得的 cDNA片段为基础，应用 RACE（Rpid
arnplification of c洲A ends）技术得至了 NJSZ，NJS4和 NJS22三个基因的全长
。DNA序列。在推测的开放读码框两侧设计引物，在cDNA中进行PCR扩增，
分别得到上述三个基因的全长。DNA克隆。
蛋白质一级结构分析表明，NJSZ与拟南芥的一个推导蛋白（CAB7101，l）
具有较高的同源性。NJS4推测为小麦的一个丝氨酸／苏氨酸蛋白激酶。NJS22
推测为小麦的一个硫代硫酸硫转移酶基因。
己克隆的植物抗病基因 Pto和 XaZI为丝氨酸／苏氨酸蛋白激酶，NJS4也属
于该基因家族。NJS4在6VS／6AL易位系抗白粉病反应中的作用有待于进一步
研究。白粉菌诱导后的基因表达分析表明，NJSZ、NJS22和 NJS35可能为小麦
中新的防卫反应基因。
以NJS22为探针，用中国春缺体／四体系进行Southern杂交定位，并用设计
的NJS22基因特异性引物在缺体／四体系和端二体系基因组DNA中扩增定位。
Southern杂交结果表明，N6B／T6A和 N6B／T6D缺一条杂交带。PCR扩增结果
表明，缺体四体系的N6B／T6A和N6B／T6D缺一条750hp扩增产物，端二体系
的6BL缺一条750hp扩增产物。从而将小麦硫代硫酸硫转移酶基因（NJS22）
定位在6B染色体短臂上。
Powdery mildew is one of the severest diseases in wheat production. A Triticum
aes(ivum-Haynaldia viiiosa 6VS/6AL translocation line containing powdery mildew
resistance gene Pm21 was developed in Cytogenetic Institute of Nanjing Agricultural
University,which is effective against all the current biotypes of Erysiphe
graminis .The line is also resistant to the new races of Puccinia strijformis (stripe
rust) and Puccinia recondita (leaf rust) in China. Cloning of Pm2 I and the related
genes for powdery mildew resistance is significance for understand its resistence
mechanism and disease resistance breeding.
To construct cDNA library is a prerequisite for gene cloning.In this study,two
cDNA libraries were constructed using mRNA from leaves of Triticum aestivum-
Haynaidia viiiosa 6VS/6AL translocation line, induced and non-induced with
Erysiphe graminis infection. The cloning vector was an expression plasmid vector
pSPORTI, introduced into E. coil DH I OB.The eDNA library of induced translocation
line contains about 300,000 recombinant eDNA clones, with average insert size of
1.2 Kb. the eDNA library of non-induced transloeation line contains about 500,000
recombinant cDNA clones, and the average insert was 1.25 Kb ,ranging basically
from 400bp to 2.2 Kb. The library was successfully screened with the probe of wheat
thaumatin-like protein gene pWIR cloned by ourselves.
mRNA differential-display reverse-transcription polymerase chain reaction
(DDRT-PCR) was used to compare mRNA from leaves of the translocation line
induced and non-induced with Erysiphe graminis ,total 35 different cDNAs were
obtained.Northern blot analysis confirmed that eight cDNAs showed increased signal
with mRNA from the induced leaves.Sequenee and database serehes revealed that
four of them are homologous to Giycine max protein kinase GmPK6,Dahsca
glomerata thiosulfate sulfurtransferase mRNA, Triticum aesfivum mRNA for putative
1n2. 1-like protein and Arabidopsis thaliana BAC clone Fl 5G1 6 DNA in chromosom
3 respectively,They are all new genes cloned in commen wheat.The other three genes
represent wheat chloroplast genes rbcl,psal for large subunit of ribulose 1,5-
bisphosphate carboxylase/oxygenase , apoprotein I,Rubisco activase gene and rpoA
gene respectively. These wheat genes may be involved in powdery mildew resistance
interaction.
EcoRI, EcoRV and HindIII-cut genomic DNA of Yanginai V and 6VS/6AL
translocation line was used for southern hybridization analysis with cloned cDNAs as
probes. The result showed that thiosulfate sulfurtransferase gene (NJS22) had
polymorphism bands between all three restriction enzyme-cut two genomic DNAs,
and the protein kinase gene (NJS4), 1n2-1-like protein gene (NJS35), beclinl like
gene (NJS2) all had some extent differences between two genome.
Based on cDNA fragments from DDRT-PCR, RACE technique was successfuly
used to obtain full-length eDNA sequences of NJS2, NJS4 and NJS22 genes. Then
gene specific primers were designed flanking putative open reading frames (OREs),
and three full-length eDNA clones of NJS2, NJS4 and NJS22 genes had been
obtained by PCR.
Protein database search revealed that NJS2 was significantly homologous to a
putative protein in Arabidopsls thaliana (accession number: CAB 7101.1). NJS4 was
a wheat serine/threonine protein kinase and NJS22 was a wheat thiosulfate
sulfurtransferase gene.
4
The cloned plant disease resistance genes of Pt
所培育的小麦-簇毛麦6VS/6AL易位系带有目前我国最有效的抗白粉病基因
Pm21，此外还对我国当前新优势条锈菌生理小种表现高抗，对叶锈也表现高抗。
对Pm21及抗白粉病相关基因进行克隆研究不仅有利于阐明其抗病机制，而且
对抗病育种有重大意义。
构建cDNA文库是基因克隆的基础，本研究用表达型质粒载体p~(spon-1)构建
了经白粉菌（Erysiphe graminis）诱导48小时和未经诱导的小麦-簇毛麦6VS/6AL
易位系（Triticum aestivum-Haynaldia villosa translocation line）叶片cDNA文库
二个。文库奇主菌为E.coli DH10B。非诱导文库包含约50万个重组cDNA克
隆，平均插入片段为1.25kb，主要分布在400bp-2.2kb之间。诱导文库包含约30
万个重组cDNA克隆，平均插入片段1.2kb。用本室克隆的小麦类甜蛋白基因
pWIR作探针，进行杂交筛库，杂交信号明显，重复性好。
用mRNA差异显示技术对经白粉菌诱导24小时，48小时和未经诱导的
6VS/6AL易位系叶片mRNA进行比较，共获得35个差异cDNA片段。Northern
杂交证明，其中8个cDNA克隆，在白粉菌诱导下其基因表达增强。
测序和同源性比较表明，其中4个分别与大豆蛋白激酶GmPK6、达提斯
卡（Datisca glomerata）硫代硫酸硫转移酶，推测的小麦In2-1人类似蛋白基因和
拟南芥第三染色体上的BAC克隆F15G16序列高度同源。它们都是小麦中的新
基因。另外三个推测为小麦叶绿体的三个基因：1.5-二磷酸核酮糖羧化酶
（Rubisco）大亚基基因，Rubisco活化酶基因和RNA聚合酶rpoA基因。这些
基因可能参与了小麦抗白粉病反应。
扬麦5号和6VS/6AL易位系基因组DNA经EcoRⅠ、EcoRⅤ和HindⅢ三
种酶切，用克隆的cDNA片段作探针，杂交结果表明，硫转移酶基因（NJS22）
在三种酶切中均有差异。小麦蛋白激酶基因（NJS4）、InZ八类似基因（NJS35）
和 beclin类似基因（NJSZ）在两者间均有不同程度的差异。
以 mRNA差异显示获得的 cDNA片段为基础，应用 RACE（Rpid
arnplification of c洲A ends）技术得至了 NJSZ，NJS4和 NJS22三个基因的全长
。DNA序列。在推测的开放读码框两侧设计引物，在cDNA中进行PCR扩增，
分别得到上述三个基因的全长。DNA克隆。
蛋白质一级结构分析表明，NJSZ与拟南芥的一个推导蛋白（CAB7101，l）
具有较高的同源性。NJS4推测为小麦的一个丝氨酸／苏氨酸蛋白激酶。NJS22
推测为小麦的一个硫代硫酸硫转移酶基因。
己克隆的植物抗病基因 Pto和 XaZI为丝氨酸／苏氨酸蛋白激酶，NJS4也属
于该基因家族。NJS4在6VS／6AL易位系抗白粉病反应中的作用有待于进一步
研究。白粉菌诱导后的基因表达分析表明，NJSZ、NJS22和 NJS35可能为小麦
中新的防卫反应基因。
以NJS22为探针，用中国春缺体／四体系进行Southern杂交定位，并用设计
的NJS22基因特异性引物在缺体／四体系和端二体系基因组DNA中扩增定位。
Southern杂交结果表明，N6B／T6A和 N6B／T6D缺一条杂交带。PCR扩增结果
表明，缺体四体系的N6B／T6A和N6B／T6D缺一条750hp扩增产物，端二体系
的6BL缺一条750hp扩增产物。从而将小麦硫代硫酸硫转移酶基因（NJS22）
定位在6B染色体短臂上。
Powdery mildew is one of the severest diseases in wheat production. A Triticum
aes(ivum-Haynaldia viiiosa 6VS/6AL translocation line containing powdery mildew
resistance gene Pm21 was developed in Cytogenetic Institute of Nanjing Agricultural
University,which is effective against all the current biotypes of Erysiphe
graminis .The line is also resistant to the new races of Puccinia strijformis (stripe
rust) and Puccinia recondita (leaf rust) in China. Cloning of Pm2 I and the related
genes for powdery mildew resistance is significance for understand its resistence
mechanism and disease resistance breeding.
To construct cDNA library is a prerequisite for gene cloning.In this study,two
cDNA libraries were constructed using mRNA from leaves of Triticum aestivum-
Haynaidia viiiosa 6VS/6AL translocation line, induced and non-induced with
Erysiphe graminis infection. The cloning vector was an expression plasmid vector
pSPORTI, introduced into E. coil DH I OB.The eDNA library of induced translocation
line contains about 300,000 recombinant eDNA clones, with average insert size of
1.2 Kb. the eDNA library of non-induced transloeation line contains about 500,000
recombinant cDNA clones, and the average insert was 1.25 Kb ,ranging basically
from 400bp to 2.2 Kb. The library was successfully screened with the probe of wheat
thaumatin-like protein gene pWIR cloned by ourselves.
mRNA differential-display reverse-transcription polymerase chain reaction
(DDRT-PCR) was used to compare mRNA from leaves of the translocation line
induced and non-induced with Erysiphe graminis ,total 35 different cDNAs were
obtained.Northern blot analysis confirmed that eight cDNAs showed increased signal
with mRNA from the induced leaves.Sequenee and database serehes revealed that
four of them are homologous to Giycine max protein kinase GmPK6,Dahsca
glomerata thiosulfate sulfurtransferase mRNA, Triticum aesfivum mRNA for putative
1n2. 1-like protein and Arabidopsis thaliana BAC clone Fl 5G1 6 DNA in chromosom
3 respectively,They are all new genes cloned in commen wheat.The other three genes
represent wheat chloroplast genes rbcl,psal for large subunit of ribulose 1,5-
bisphosphate carboxylase/oxygenase , apoprotein I,Rubisco activase gene and rpoA
gene respectively. These wheat genes may be involved in powdery mildew resistance
interaction.
EcoRI, EcoRV and HindIII-cut genomic DNA of Yanginai V and 6VS/6AL
translocation line was used for southern hybridization analysis with cloned cDNAs as
probes. The result showed that thiosulfate sulfurtransferase gene (NJS22) had
polymorphism bands between all three restriction enzyme-cut two genomic DNAs,
and the protein kinase gene (NJS4), 1n2-1-like protein gene (NJS35), beclinl like
gene (NJS2) all had some extent differences between two genome.
Based on cDNA fragments from DDRT-PCR, RACE technique was successfuly
used to obtain full-length eDNA sequences of NJS2, NJS4 and NJS22 genes. Then
gene specific primers were designed flanking putative open reading frames (OREs),
and three full-length eDNA clones of NJS2, NJS4 and NJS22 genes had been
obtained by PCR.
Protein database search revealed that NJS2 was significantly homologous to a
putative protein in Arabidopsls thaliana (accession number: CAB 7101.1). NJS4 was
a wheat serine/threonine protein kinase and NJS22 was a wheat thiosulfate
sulfurtransferase gene.
4
The cloned plant disease resistance genes of Pt
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