甘蓝型油菜黄化和光叶突变体的基因定位及克隆
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摘要
作物中叶片的突变体多种多样,包括有叶色,叶形态等突变体类型。他们为研究作物的光合作用、呼吸作用、光能利用、生长发育,基因功能及某些组分(蜡质、色素等)的合成代谢调控等提供良好的实验材料。一些突变体的表型明显稳定(如棉花牙黄,水稻紫叶,油菜黄叶等)被应用于杂种优势的利用中。本研究对甘蓝型油菜黄化突变体BnaC.ygl和光叶突变体B. napus Nilla glossy进行了初步的表型解析,遗传分析和基因定位,并成功克隆到了叶色基因BnaC.YGL,这为探究表型机理及其在育种中的应用奠定了基础。主要研究结果如下：
1甘蓝型油菜隐性叶色突变体BnaC.ygl基因的精细定位及克隆
1)与野生型相比,突变体BnaC.ygl叶片颜色在苗期叶片黄化,现蕾期和抽薹期叶色接近于野生型。对其叶绿素含量的测定结果表明,苗期突变体和野生型的叶绿素a,叶绿素b,总叶绿素含量和叶绿素a/b值均差异显著,开花期突变体叶绿素含量与野生型差异不显著。
2)经透射电镜分析结果表明苗期突变体叶绿体类囊体膜和基粒片层垛叠数有所减少,基粒片层间稀疏,无规则,叶绿体发育不良。在开花期时,突变体叶绿体类囊体膜和基粒片层垛叠数量明显增加,与野生型无明显差异。
3)利用了两个群体对BnaC.YGL基因精细定位,将BnaC.YGL基因定在甘蓝型油菜N17的0.35cM的区间,同时获得18对与BnaC.YGL共分离的标记。并推测标记SC25附近的与甘蓝中Bo1027145(HO1)同源的基因为BnaC.YGL的候选基因。比较测序分析结果表明该基因在突变体中缺失了C基因组上的同源拷贝(BnaC.HO1)。遗传转化的结果证明BnaC.HO1是BnaC.ygl叶色突变体的突变基因。
4)甘蓝型油菜BnaC.HO1与甘蓝HO1的相似度最高,与BnaA.HO1的同源性也在90%以上,与拟南芥和白菜达80%以上,表明BnaC.HO1在进化进程中比较保守。5) RT-PCR结果表明BnaC.YGL基因在苗期展开叶和角果中表达较强,而发芽的种子和根中表达较弱。亚细胞定位的结果表明,BnaC.YGL基因定位于原生质的叶绿体中。
6)与BnaC.YGL基因共分离的16个标记在突变体和野生型中均检测到多态性,这些标记覆盖了甘蓝C07染色体至少1.2Mb的区段。另外检测到在突变体中有3个基因均丢失了N17染色体定位区段的同源拷贝,因此我们认为经EMS诱变获得的突变体BnaC.ygl可能在N17染色体上有基因组的大片段的缺失。
2甘蓝型油菜隐性光叶突变体的遗传分析和BnaCNgll基因定位
1)光叶突变体Nilla glossy的茎杆,叶片,花蕾及角果皮均不被蜡粉,表面油亮,整个生育期此表型都明显可见。扫描电镜扫描也显示光叶突变体的茎干表面被少量的片状和颗粒状蜡质晶体覆盖而叶表面被少量颗粒状晶体覆盖。
2)遗传分析结果表明该光叶表型受到两对隐性基因控制。
3)利用F2群体筛选到5个可能的多态性标记,选取G1组合中BC2代1：1分离的16个株系取样验证。结果表明只有SSR标记CB10443能在7个株系中表型出多态性,其他标记均无多态性,说明CB10443与控制光叶性状的基因中一对基因连锁,并将其命名为"BnaC.NGL1"。
4)通过AFLP技术和比较基因组开发标记,并利用包含1191个单株G1BC2群体将BnaC.NGL1锁定在甘蓝型油菜N11连锁群SSR标记gls41和gls79之间,它们之前的遗传距离为0.16cM。两个标记在甘蓝C01上的距离大约为230kb,包含约51个注释的基因。
Leaf mutants of crops are abundant and diverse such as the mutation of leaf coloration, leaf morphology and so on. Leaf mutants may be good materials for the researchs about photosynthesis, respiration, energy use, plant development, gene function, pigment metabolism, wax metabolism and so on.The mutation of Leaf is very obvious trait mutation and the mutation frequency is high. They may also be used as a morphological marker for crossbreeding. In our study, we obtained a chlorophyll-deficient mutant and a gossly mutant of Brassica napus. The researchs about preliminary phenotypic analysis, genetic analysis and mapping of the chlorophyll-deficient mutant (BnaC.ygl) and the gossly mutant (Nilla glossy) had been done. And the BnaC.ygl gene was isolated by a map-based cloning approach with the candidate gene approach. The major points were described as follows:
1The mapping and coloning of Bna C.YGL gene in a chlorophyll-deficient mutant of Brassica napus
1) There was no changes in the pigment composition of BnaC.ygl mutant.The Chl a and Ch1b levels in BnaC.ygl mutant and the wild type were significant differences at seeding stage, but not at flowering stage.
2) In BnaC.ygl mutant, the chloroplasts displayed less dense granal stacks and fewer granal membranes compared with those of the wild type. However, in twenty-week-old plants, Chloroplasts in the BnaC.ygl mutant and wild-type had well-ordered granal stacks and normal granal membranes. There was no apparent difference in the structures of chloroplasts in the BnaC.ygl mutant and wild-type.
3) Mapping of the gene was subsequently conducted in two populations with yellow-green leaves (population Ⅰ BCs and Ⅰ BC4, which comprised3472and5288individuals respectively).And the IP, SSR makers were developed from the sequences of B. oleracea. Then the BnaC. YGL gene was narrowed into a0.35cM region in the linkage group N17of Brassica napus. And in the two populations,18makers cosegregated with BnaC.YGL. The results suggested the recombination between cosegregated markers and BnaC.YGL was severely suppressed. BLAST analysis revealed that the sequences of the makers displayed highly conserved homo logy with C07of B. oleracea. The analysis of the information342genes from
B. oleracea homologous region indicated that an ORF(Bol027145) in the vicinity of
SC25maker had homologous to HOI which can catalytic decomposition of heme in the heme/bilin branch. Multiple mutant of HOI in Arabidopsis exhibited yellow-green leaves. Thus we hypothesized that this gene in B.napus, the ortholog of atHO1may be a candidate gene of BnaC. YGL. Comparative sequencing results showed that homologous copy (BnaC.HO1) of HO1on the C genome was losted in the mutant. Two resulting complementation constructs (gDNA和cDNA) were introduced into the BnaC.ygl mutant by Agrobacterium tumefaciens-mediated transformation. The genetic complementation suggested BnaC.HO1corresponds to the mutant gene in the BnaC.ygl mutant.
4) Comparison of the deduced amino acid sequences between BnaC.HO1and other Brassica HO1s further revealed several highly conserved regions as signature sequence and conserved amino acid residues. These results suggested that BnaC.HO1is highly homologous to BoHO1. Moreover, BnaC.HO1shared more than90%,80%,80%amino acid identities with BnaA.HO1, AtHO1, BrHO1.
5) Semi-quantitative RT-PCR analysis indicated that BnaC.YGL displayed higher expression level in young leaves and pods and lower in germinating seeds and roots. The subcelluar localization assay showed the BnaC. YGL protein was localized in the chloroplast of the protoplast.
6) The analysis of16cosegregated makers covering C07at least1.2Mb in B. oleracea showed that16cosegregated makers were detected polymorphism in the BnaC.ygl mutant and wild-type. And comparative sequencing results indicated that the C7homologous copy of three genes were losted. These suggested that there may be a large fragment deletion on N17in the BnaC.ygl mutant.
2Genetic characterization and fine mapping of a gossly mutant (Nilla glossy)in Brassica napus
1) The wax of stems, leaves, buds and pods in the glossy mutant was defective in the entire growth period. Scanning electron microscopy indicated that the wax on the leaf and stem surface in Nilla glossy mutan was significantly reduced.
2) The phenotype of the filial generations obtained by the crosses of the BnaC.ygl mutant and the normal parents (No2127-17and Bing409) was thoroughly investigated. The reciprocal F1plants exhibited glossy leaves as No2127-17and Bing409, which indicated that the phenotype of the glossy mutant was controlled by nuclear genes. The F2populations from both crosses showed an expected Mendelian inheritance ratio of15:1. And the ratio of normal plants to glossy plants in the BC1progenies was approximately3:1. These data indicated that the phenotype of the Nilla glossy mutant was controlled by two recessive genes.
3) Five possible polymorphic markers from F2population were used to detect the polymorphisms in16lines with1:1separation in BC2generation of G1combination.The results showed that CB10443had polymorphisms in7lines, which indicated that CB10443was linked to one of the two genes and the gene linked with CB10443was named as "BnaC.NGLl".
4) The rough mapping mapped BnaC.Ngll to linkage group N11. Eight AFLP makers were developed by AFLP technique combined with BAS. The sequences of the CB10443and AFLP markers were submitted to the public database for identification of putative orthologues, which showed sequence homology to the CO I of B. oleracea. The SSR makers were designed from the sequence of C01in B. oleracea and analyzed in population G1BC2with1191plants. Finally, BnaC.YGL locus was mapped to the interval between the markers gls41and gls79, corresponding to a genetic distance of0.16cM. Compared with C01in B. oleracea, the interval between the markers gls41and gls79was about230kb, including51annotated genes.
1甘蓝型油菜隐性叶色突变体BnaC.ygl基因的精细定位及克隆
1)与野生型相比,突变体BnaC.ygl叶片颜色在苗期叶片黄化,现蕾期和抽薹期叶色接近于野生型。对其叶绿素含量的测定结果表明,苗期突变体和野生型的叶绿素a,叶绿素b,总叶绿素含量和叶绿素a/b值均差异显著,开花期突变体叶绿素含量与野生型差异不显著。
2)经透射电镜分析结果表明苗期突变体叶绿体类囊体膜和基粒片层垛叠数有所减少,基粒片层间稀疏,无规则,叶绿体发育不良。在开花期时,突变体叶绿体类囊体膜和基粒片层垛叠数量明显增加,与野生型无明显差异。
3)利用了两个群体对BnaC.YGL基因精细定位,将BnaC.YGL基因定在甘蓝型油菜N17的0.35cM的区间,同时获得18对与BnaC.YGL共分离的标记。并推测标记SC25附近的与甘蓝中Bo1027145(HO1)同源的基因为BnaC.YGL的候选基因。比较测序分析结果表明该基因在突变体中缺失了C基因组上的同源拷贝(BnaC.HO1)。遗传转化的结果证明BnaC.HO1是BnaC.ygl叶色突变体的突变基因。
4)甘蓝型油菜BnaC.HO1与甘蓝HO1的相似度最高,与BnaA.HO1的同源性也在90%以上,与拟南芥和白菜达80%以上,表明BnaC.HO1在进化进程中比较保守。5) RT-PCR结果表明BnaC.YGL基因在苗期展开叶和角果中表达较强,而发芽的种子和根中表达较弱。亚细胞定位的结果表明,BnaC.YGL基因定位于原生质的叶绿体中。
6)与BnaC.YGL基因共分离的16个标记在突变体和野生型中均检测到多态性,这些标记覆盖了甘蓝C07染色体至少1.2Mb的区段。另外检测到在突变体中有3个基因均丢失了N17染色体定位区段的同源拷贝,因此我们认为经EMS诱变获得的突变体BnaC.ygl可能在N17染色体上有基因组的大片段的缺失。
2甘蓝型油菜隐性光叶突变体的遗传分析和BnaCNgll基因定位
1)光叶突变体Nilla glossy的茎杆,叶片,花蕾及角果皮均不被蜡粉,表面油亮,整个生育期此表型都明显可见。扫描电镜扫描也显示光叶突变体的茎干表面被少量的片状和颗粒状蜡质晶体覆盖而叶表面被少量颗粒状晶体覆盖。
2)遗传分析结果表明该光叶表型受到两对隐性基因控制。
3)利用F2群体筛选到5个可能的多态性标记,选取G1组合中BC2代1：1分离的16个株系取样验证。结果表明只有SSR标记CB10443能在7个株系中表型出多态性,其他标记均无多态性,说明CB10443与控制光叶性状的基因中一对基因连锁,并将其命名为"BnaC.NGL1"。
4)通过AFLP技术和比较基因组开发标记,并利用包含1191个单株G1BC2群体将BnaC.NGL1锁定在甘蓝型油菜N11连锁群SSR标记gls41和gls79之间,它们之前的遗传距离为0.16cM。两个标记在甘蓝C01上的距离大约为230kb,包含约51个注释的基因。
Leaf mutants of crops are abundant and diverse such as the mutation of leaf coloration, leaf morphology and so on. Leaf mutants may be good materials for the researchs about photosynthesis, respiration, energy use, plant development, gene function, pigment metabolism, wax metabolism and so on.The mutation of Leaf is very obvious trait mutation and the mutation frequency is high. They may also be used as a morphological marker for crossbreeding. In our study, we obtained a chlorophyll-deficient mutant and a gossly mutant of Brassica napus. The researchs about preliminary phenotypic analysis, genetic analysis and mapping of the chlorophyll-deficient mutant (BnaC.ygl) and the gossly mutant (Nilla glossy) had been done. And the BnaC.ygl gene was isolated by a map-based cloning approach with the candidate gene approach. The major points were described as follows:
1The mapping and coloning of Bna C.YGL gene in a chlorophyll-deficient mutant of Brassica napus
1) There was no changes in the pigment composition of BnaC.ygl mutant.The Chl a and Ch1b levels in BnaC.ygl mutant and the wild type were significant differences at seeding stage, but not at flowering stage.
2) In BnaC.ygl mutant, the chloroplasts displayed less dense granal stacks and fewer granal membranes compared with those of the wild type. However, in twenty-week-old plants, Chloroplasts in the BnaC.ygl mutant and wild-type had well-ordered granal stacks and normal granal membranes. There was no apparent difference in the structures of chloroplasts in the BnaC.ygl mutant and wild-type.
3) Mapping of the gene was subsequently conducted in two populations with yellow-green leaves (population Ⅰ BCs and Ⅰ BC4, which comprised3472and5288individuals respectively).And the IP, SSR makers were developed from the sequences of B. oleracea. Then the BnaC. YGL gene was narrowed into a0.35cM region in the linkage group N17of Brassica napus. And in the two populations,18makers cosegregated with BnaC.YGL. The results suggested the recombination between cosegregated markers and BnaC.YGL was severely suppressed. BLAST analysis revealed that the sequences of the makers displayed highly conserved homo logy with C07of B. oleracea. The analysis of the information342genes from
B. oleracea homologous region indicated that an ORF(Bol027145) in the vicinity of
SC25maker had homologous to HOI which can catalytic decomposition of heme in the heme/bilin branch. Multiple mutant of HOI in Arabidopsis exhibited yellow-green leaves. Thus we hypothesized that this gene in B.napus, the ortholog of atHO1may be a candidate gene of BnaC. YGL. Comparative sequencing results showed that homologous copy (BnaC.HO1) of HO1on the C genome was losted in the mutant. Two resulting complementation constructs (gDNA和cDNA) were introduced into the BnaC.ygl mutant by Agrobacterium tumefaciens-mediated transformation. The genetic complementation suggested BnaC.HO1corresponds to the mutant gene in the BnaC.ygl mutant.
4) Comparison of the deduced amino acid sequences between BnaC.HO1and other Brassica HO1s further revealed several highly conserved regions as signature sequence and conserved amino acid residues. These results suggested that BnaC.HO1is highly homologous to BoHO1. Moreover, BnaC.HO1shared more than90%,80%,80%amino acid identities with BnaA.HO1, AtHO1, BrHO1.
5) Semi-quantitative RT-PCR analysis indicated that BnaC.YGL displayed higher expression level in young leaves and pods and lower in germinating seeds and roots. The subcelluar localization assay showed the BnaC. YGL protein was localized in the chloroplast of the protoplast.
6) The analysis of16cosegregated makers covering C07at least1.2Mb in B. oleracea showed that16cosegregated makers were detected polymorphism in the BnaC.ygl mutant and wild-type. And comparative sequencing results indicated that the C7homologous copy of three genes were losted. These suggested that there may be a large fragment deletion on N17in the BnaC.ygl mutant.
2Genetic characterization and fine mapping of a gossly mutant (Nilla glossy)in Brassica napus
1) The wax of stems, leaves, buds and pods in the glossy mutant was defective in the entire growth period. Scanning electron microscopy indicated that the wax on the leaf and stem surface in Nilla glossy mutan was significantly reduced.
2) The phenotype of the filial generations obtained by the crosses of the BnaC.ygl mutant and the normal parents (No2127-17and Bing409) was thoroughly investigated. The reciprocal F1plants exhibited glossy leaves as No2127-17and Bing409, which indicated that the phenotype of the glossy mutant was controlled by nuclear genes. The F2populations from both crosses showed an expected Mendelian inheritance ratio of15:1. And the ratio of normal plants to glossy plants in the BC1progenies was approximately3:1. These data indicated that the phenotype of the Nilla glossy mutant was controlled by two recessive genes.
3) Five possible polymorphic markers from F2population were used to detect the polymorphisms in16lines with1:1separation in BC2generation of G1combination.The results showed that CB10443had polymorphisms in7lines, which indicated that CB10443was linked to one of the two genes and the gene linked with CB10443was named as "BnaC.NGLl".
4) The rough mapping mapped BnaC.Ngll to linkage group N11. Eight AFLP makers were developed by AFLP technique combined with BAS. The sequences of the CB10443and AFLP markers were submitted to the public database for identification of putative orthologues, which showed sequence homology to the CO I of B. oleracea. The SSR makers were designed from the sequence of C01in B. oleracea and analyzed in population G1BC2with1191plants. Finally, BnaC.YGL locus was mapped to the interval between the markers gls41and gls79, corresponding to a genetic distance of0.16cM. Compared with C01in B. oleracea, the interval between the markers gls41and gls79was about230kb, including51annotated genes.
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