海岛棉LRR-TM类抗病基因GbaVd1和GbaVd2的克隆与功能研究
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摘要
棉花黄萎病是土传性真菌病害,其致病菌为大丽轮枝菌,给棉花生产造成巨大的损失。对大丽轮枝菌的研究虽然开展了大量的工作,但尚无成果可应用于生产防治,主要原因是抗病基因资源匮乏和大丽轮枝菌致病机理复杂。一般认为,转基因抗病育种是解决棉花黄萎病的有效途径,因此,克隆对大丽轮枝菌有效的抗病基因并鉴定其功能就成为当前棉花抗黄萎病研究的主要内容之一。
本研究通过同源克隆、TAIL-PCR和RACE相结合的方法,从抗病棉种海7124中克隆了同番茄抗黄萎病Ve1和Ve2同源的LRR-TM类基因GbaVd1和GbaVd2。GbaVd1基因cDNA全长3262 bp,开放阅读框为3063bp,编码1020 aa,5'UTR为69bp,3'UTR为130bp,GbaVd2基因cDNA全长3466 bp,开放阅读框为3249 bp,编码1082 aa,5'UTR为68bp,3'UTR为149bp,两个基因均不含内含子。通过TAIL-PCR克隆了两个基因的侧翼序列分别为900 bp和1107 bp。尽管GbaVd1基因和GbaVd2基因与Ve1和Ve2的一致性不到60%,编码蛋白的一致性低于50%,但它们与LRR-TM类抗病基因类似,含有信号肽、N端成熟区、LRR重复单元、胞外结构域、跨膜区和胞内结构域。其中,GbaVd1含有32个LRR重复单元,GbaVd2含有33个LRR重复单元,分别含有22个和26个潜在的糖基化位点。Psort预测表明,GbaVd1和GbaVd2可能定位于细胞质膜上。侧翼序列分析表明,它们含有潜在的真菌激发子应答元件Box-W1、茉莉酸甲酯应答元件TGACG基序和CGTCA基序、脱落酸应答元件ABRE、应答防卫反应的顺式作用元件TC-rich repeat等。
根据GbaVd1和GbaVd2基因,从不同野生棉种中克隆了这两个基因(Vd1和Vd2)及其侧翼序列,并分析这些基因的差异性与棉种抗病性的关系。不同棉种Vd1和Vd2基因的差异主要是以单核苷酸变异为主,引起了编码蛋白多个有义突变,此外也存在着核酸片段的缺失,如纳尔逊棉GneVd1、戴维逊棉GdaVd2、斯托提棉GstVd2等,引起了编码蛋白信号肽缺失、N端成熟区改变、糖基化位点增加或删除、胞内短肽丧失等变化,但这些差异和棉种的抗感性并不存在明显的相关性。
转基因拟南芥实验表明,GbaVd1和GbaVd2过表达能够增强拟南芥对大丽轮枝菌毒素的耐受性,未发现叶片黄化和根系发育受抑制的病症。在大丽轮枝菌侵染的条件下,GbaVd1和GbaVd2基因明显提高了拟南芥对大丽轮枝菌的抗性,转基因株系分蘖能力恢复正常,生长延缓得到明显改善,株高受影响较小,种子结实正常,只有叶片表现出一定黄化症状。说明GbaVd1和GbaVd2是两个具有抗大丽轮枝菌功能的基因。
对抗性GbaVd1和GbaVd2转基因株系基因芯片表达分析表明,分别有44个和61个基因共同被上调和下调1.5倍以上,而且应激反应相关的基因被大量诱导表达,包括生物胁迫、非生物胁迫、免疫应答、应激调控系统等。对所有差异表达基因分析发现,GbaVd1和GbaVd2调控了多个抗病信号因子的表达,包括SERK2、SERK4、SGT1A、SGT1B、NDR1、EDS1和激酶类基因;影响了乙烯和茉莉酸信号转导途径相关基因表达,如EBF1、EBF2、EIN3、ABA1、AOS、AOC1、JMT、JAR1、JAZ7等;分别改变了黄酮化合物和萜类化合物的合成,DFR、TT5、LDOX、SQP2、GGPS1、SQE3等基因表达发生了变化;促使了细胞壁相关蛋白基因的表达发生了变化,包括Nodulin、凝聚素、富含脯氨酸蛋白、富含羟脯氨酸蛋白、富含甘氨酸蛋白、阿拉伯半乳糖蛋白等,转基因株系与细胞壁合成/降解相关的葡聚糖酶和木聚糖酶活性提高。
基于上述研究,提出了GbaVd1和GbaVd2抗病机理模式的假设:即两个基因编码类似于LRR-TM类抗病基因的跨膜受体蛋白,同大丽轮枝菌激发子直接或者间接结合,调控了抗病信号因子的表达,随后激活了乙烯和茉莉酸信号转导途径,进而改变植物抗毒素的合成和细胞壁相关蛋白的表达,形成具有抗真菌活性的植物抗毒素和增强细胞壁的组织障碍,提高寄主对大丽轮枝菌的抗性。
Cotton production was serious destroy with soil-borne fungal Verticillium dahliea and difficult to prevent. Although many research works about the Verticillium dahliea was carried out, there were no effective research achievements for disease control, due to lack of disease resistance gene resources in cotton and complexity pathogenicity of Verticillium dahliae. Therefore, cloning and identification the function of disease resistance genes to Verticillium dahliae is the urgent task to disease resistance transgenic breeding.
In this research, leucine-rich repeats transmembrane (LRR-TM) genes of GbaVd1 and GbaVd2 were cloned from the disease resistance cultivar Gossypium barbadense cv. Hai7124 by TAIL-CPR and RACE methods, which homologus to anti-Verticillium genes Ve1 and Ve2 in tomato. The full length cDNA of GbaVd1 is 3262 bp in which a 3063 bp ORF encoding a 1020 aa protein, a 3′UTR and 5′UTR of 130 bp and 69 bp, respectively. The full length cDNA of GbaVd2 comprised 3466 bp and included an ORF of 3249 bp, encoding a 1082 aa protein, a 68 bp and 149 bp long 5′UTR and 3′UTR, respectively. The flanking sequences of GbaVd1 and GbaVd2 were cloned by TAIL-PCR method and 900 bp and 1107 bp fragment were obtained, respectively, and there was no intron in GbaVd1 and GbaVd2. Although the identity of the nucleotides between GbaVd1/GbaVd2 and Ve1/Ve2 is less than 60% and code proteins is lower than 50%, their structures was similar to the LRR-TM disease resistance gene, which contains signal peptide, N terminus of the mature protein, leucine-rich repeats (LRRs), extracytoplasmic domain, transmembrane domain and cytoplasmic domain. In GbaVd1 and GbaVd2, there are 32 and 33 LRRs, also include 22 and 26 potential glycosylation sites, respectively. The subcellular localization of GbaVd1 and GbaVd2 was predicted in plasma membrane by Psort. Morever, the flanking sequences were possible contain many cis-elements, such as fungal elicitor responsive element of Box-W1, cis-acting regulatory element involved in the MeJA-responsiveness of TGACG motif and CGTCA motif, cis-acting element involved in the abscisic acid responsiveness of ABRE, cis-acting element involved in defense and stress responsiveness of TC-rich repeats and so on.
Vd1 and Vd2 genes and theirs flanking sequences were cloned from different wild cottons base on GbaVd1 and GbaVd2 sequences, and the relation between genes difference and cottons sensitivity to Verticillium dahliae was analysis. The difference of Vd1 and Vd2 genes were mainly shown single nucleotide polymorphisms which lead to many sense mutations. The lack of short nucleotide fragment was also exist in Vd1 and Vd2 genes, such as GneVd1 from Gossypium nelsonii, GdaVd2 from Gossypium davidsonii, GstVd2 from Gossypium sturtianum and so on, which result in the deletion of signal peptide, change the N terminus of mature protein, add or delete the glycosylation sites, lose of cytoplasmic domain and so on. However, there are no obvious correlation between the genes difference and the cottons sensitivity to Verticillium dahliae.
Overexpressin of GbaVd1 and GbaVd2 were shown to enhance the tolerance to Verticilliun dahliae toxin (VD-toxin) in transgenic Arabidopsis, and did not found the symptoms of leaf chlorosis and root depression. Morever, the resistance of GbaVd1 and GbaVd2 transgenic plants to V. dahliae was obvious improved. For example, compared to the infected of wild type Col-0, the tillers of transgenic plants were returned to normal, the growth retardants was improved, the influence of inflorescence height was reduced and the seeded was shown normal, but the leaves of transgenic plant were also shown chlorosis as the wild type. Therefore, the results were shown that GbaVd1 and GbaVd2 genes possess anti-Verticillium function in V. dahliae invade progress.
Analysis the gene expression profile were shown that 44 genes were co-upregulated and 61 genes co-downregulated more than 1.5 folds in both GbaVd1 and GbaVd2 transgenic plants, and many differential expression genes were related to response to stimulus which contained biotic stress, abiotic stress, immune response, regulation of response to stimulus and so on. The research was found that some signal factors of disease resistance were regulated in transgenic plants, such as SERK2, SERK4, SGT1A, SGT1B, NDR1, EDS1, kinase genes and so on. The ethylene and jasmonic acid signal transduction pathway were also effected in transgenic plants, the expression of EBF1, EBF2, EIN3, ABA1, AOS, AOC1, JMT, JAR1, JAZ7, etc were induced. The biosynthesis pathway of flavonoid and terpenoid were changed too, their related genes of DFR, TT5, LDOX, SQP2, GGPS1 and SQE3 were regulated. Overexpression of GbaVd1 and GbaVd2 were also promoted the expression of cell wall related protein genes, such as nodulin, lectin, proline-rich protein, hydroxyproline-rich glycoprotein, glycine-rich protein, arabinogalactan-protein and so on, and the activity ofβ-1,3-glucanase andβ-D-xylosidase were enhanced, which were related to cell wall synthesis or degradation.
Base on these results, the mechanism of GbaVd1 and GbaVd2 resistance model was supposed, two genes encode LRR-TM receptor like protein could directly or indirectly combine with the V. dahliae elicitors, this action would regulate the expression of signal factor of disease resistance, then active the ethylene and jasmonic acid signal transduction pathway which lead to change the synthesis of phytoalexin and expression of cell wall related proteins, form the antifungus activity compound and enhance the cell wall structure obstacle, finally increase the host resistance to V. dahliae.
本研究通过同源克隆、TAIL-PCR和RACE相结合的方法,从抗病棉种海7124中克隆了同番茄抗黄萎病Ve1和Ve2同源的LRR-TM类基因GbaVd1和GbaVd2。GbaVd1基因cDNA全长3262 bp,开放阅读框为3063bp,编码1020 aa,5'UTR为69bp,3'UTR为130bp,GbaVd2基因cDNA全长3466 bp,开放阅读框为3249 bp,编码1082 aa,5'UTR为68bp,3'UTR为149bp,两个基因均不含内含子。通过TAIL-PCR克隆了两个基因的侧翼序列分别为900 bp和1107 bp。尽管GbaVd1基因和GbaVd2基因与Ve1和Ve2的一致性不到60%,编码蛋白的一致性低于50%,但它们与LRR-TM类抗病基因类似,含有信号肽、N端成熟区、LRR重复单元、胞外结构域、跨膜区和胞内结构域。其中,GbaVd1含有32个LRR重复单元,GbaVd2含有33个LRR重复单元,分别含有22个和26个潜在的糖基化位点。Psort预测表明,GbaVd1和GbaVd2可能定位于细胞质膜上。侧翼序列分析表明,它们含有潜在的真菌激发子应答元件Box-W1、茉莉酸甲酯应答元件TGACG基序和CGTCA基序、脱落酸应答元件ABRE、应答防卫反应的顺式作用元件TC-rich repeat等。
根据GbaVd1和GbaVd2基因,从不同野生棉种中克隆了这两个基因(Vd1和Vd2)及其侧翼序列,并分析这些基因的差异性与棉种抗病性的关系。不同棉种Vd1和Vd2基因的差异主要是以单核苷酸变异为主,引起了编码蛋白多个有义突变,此外也存在着核酸片段的缺失,如纳尔逊棉GneVd1、戴维逊棉GdaVd2、斯托提棉GstVd2等,引起了编码蛋白信号肽缺失、N端成熟区改变、糖基化位点增加或删除、胞内短肽丧失等变化,但这些差异和棉种的抗感性并不存在明显的相关性。
转基因拟南芥实验表明,GbaVd1和GbaVd2过表达能够增强拟南芥对大丽轮枝菌毒素的耐受性,未发现叶片黄化和根系发育受抑制的病症。在大丽轮枝菌侵染的条件下,GbaVd1和GbaVd2基因明显提高了拟南芥对大丽轮枝菌的抗性,转基因株系分蘖能力恢复正常,生长延缓得到明显改善,株高受影响较小,种子结实正常,只有叶片表现出一定黄化症状。说明GbaVd1和GbaVd2是两个具有抗大丽轮枝菌功能的基因。
对抗性GbaVd1和GbaVd2转基因株系基因芯片表达分析表明,分别有44个和61个基因共同被上调和下调1.5倍以上,而且应激反应相关的基因被大量诱导表达,包括生物胁迫、非生物胁迫、免疫应答、应激调控系统等。对所有差异表达基因分析发现,GbaVd1和GbaVd2调控了多个抗病信号因子的表达,包括SERK2、SERK4、SGT1A、SGT1B、NDR1、EDS1和激酶类基因;影响了乙烯和茉莉酸信号转导途径相关基因表达,如EBF1、EBF2、EIN3、ABA1、AOS、AOC1、JMT、JAR1、JAZ7等;分别改变了黄酮化合物和萜类化合物的合成,DFR、TT5、LDOX、SQP2、GGPS1、SQE3等基因表达发生了变化;促使了细胞壁相关蛋白基因的表达发生了变化,包括Nodulin、凝聚素、富含脯氨酸蛋白、富含羟脯氨酸蛋白、富含甘氨酸蛋白、阿拉伯半乳糖蛋白等,转基因株系与细胞壁合成/降解相关的葡聚糖酶和木聚糖酶活性提高。
基于上述研究,提出了GbaVd1和GbaVd2抗病机理模式的假设:即两个基因编码类似于LRR-TM类抗病基因的跨膜受体蛋白,同大丽轮枝菌激发子直接或者间接结合,调控了抗病信号因子的表达,随后激活了乙烯和茉莉酸信号转导途径,进而改变植物抗毒素的合成和细胞壁相关蛋白的表达,形成具有抗真菌活性的植物抗毒素和增强细胞壁的组织障碍,提高寄主对大丽轮枝菌的抗性。
Cotton production was serious destroy with soil-borne fungal Verticillium dahliea and difficult to prevent. Although many research works about the Verticillium dahliea was carried out, there were no effective research achievements for disease control, due to lack of disease resistance gene resources in cotton and complexity pathogenicity of Verticillium dahliae. Therefore, cloning and identification the function of disease resistance genes to Verticillium dahliae is the urgent task to disease resistance transgenic breeding.
In this research, leucine-rich repeats transmembrane (LRR-TM) genes of GbaVd1 and GbaVd2 were cloned from the disease resistance cultivar Gossypium barbadense cv. Hai7124 by TAIL-CPR and RACE methods, which homologus to anti-Verticillium genes Ve1 and Ve2 in tomato. The full length cDNA of GbaVd1 is 3262 bp in which a 3063 bp ORF encoding a 1020 aa protein, a 3′UTR and 5′UTR of 130 bp and 69 bp, respectively. The full length cDNA of GbaVd2 comprised 3466 bp and included an ORF of 3249 bp, encoding a 1082 aa protein, a 68 bp and 149 bp long 5′UTR and 3′UTR, respectively. The flanking sequences of GbaVd1 and GbaVd2 were cloned by TAIL-PCR method and 900 bp and 1107 bp fragment were obtained, respectively, and there was no intron in GbaVd1 and GbaVd2. Although the identity of the nucleotides between GbaVd1/GbaVd2 and Ve1/Ve2 is less than 60% and code proteins is lower than 50%, their structures was similar to the LRR-TM disease resistance gene, which contains signal peptide, N terminus of the mature protein, leucine-rich repeats (LRRs), extracytoplasmic domain, transmembrane domain and cytoplasmic domain. In GbaVd1 and GbaVd2, there are 32 and 33 LRRs, also include 22 and 26 potential glycosylation sites, respectively. The subcellular localization of GbaVd1 and GbaVd2 was predicted in plasma membrane by Psort. Morever, the flanking sequences were possible contain many cis-elements, such as fungal elicitor responsive element of Box-W1, cis-acting regulatory element involved in the MeJA-responsiveness of TGACG motif and CGTCA motif, cis-acting element involved in the abscisic acid responsiveness of ABRE, cis-acting element involved in defense and stress responsiveness of TC-rich repeats and so on.
Vd1 and Vd2 genes and theirs flanking sequences were cloned from different wild cottons base on GbaVd1 and GbaVd2 sequences, and the relation between genes difference and cottons sensitivity to Verticillium dahliae was analysis. The difference of Vd1 and Vd2 genes were mainly shown single nucleotide polymorphisms which lead to many sense mutations. The lack of short nucleotide fragment was also exist in Vd1 and Vd2 genes, such as GneVd1 from Gossypium nelsonii, GdaVd2 from Gossypium davidsonii, GstVd2 from Gossypium sturtianum and so on, which result in the deletion of signal peptide, change the N terminus of mature protein, add or delete the glycosylation sites, lose of cytoplasmic domain and so on. However, there are no obvious correlation between the genes difference and the cottons sensitivity to Verticillium dahliae.
Overexpressin of GbaVd1 and GbaVd2 were shown to enhance the tolerance to Verticilliun dahliae toxin (VD-toxin) in transgenic Arabidopsis, and did not found the symptoms of leaf chlorosis and root depression. Morever, the resistance of GbaVd1 and GbaVd2 transgenic plants to V. dahliae was obvious improved. For example, compared to the infected of wild type Col-0, the tillers of transgenic plants were returned to normal, the growth retardants was improved, the influence of inflorescence height was reduced and the seeded was shown normal, but the leaves of transgenic plant were also shown chlorosis as the wild type. Therefore, the results were shown that GbaVd1 and GbaVd2 genes possess anti-Verticillium function in V. dahliae invade progress.
Analysis the gene expression profile were shown that 44 genes were co-upregulated and 61 genes co-downregulated more than 1.5 folds in both GbaVd1 and GbaVd2 transgenic plants, and many differential expression genes were related to response to stimulus which contained biotic stress, abiotic stress, immune response, regulation of response to stimulus and so on. The research was found that some signal factors of disease resistance were regulated in transgenic plants, such as SERK2, SERK4, SGT1A, SGT1B, NDR1, EDS1, kinase genes and so on. The ethylene and jasmonic acid signal transduction pathway were also effected in transgenic plants, the expression of EBF1, EBF2, EIN3, ABA1, AOS, AOC1, JMT, JAR1, JAZ7, etc were induced. The biosynthesis pathway of flavonoid and terpenoid were changed too, their related genes of DFR, TT5, LDOX, SQP2, GGPS1 and SQE3 were regulated. Overexpression of GbaVd1 and GbaVd2 were also promoted the expression of cell wall related protein genes, such as nodulin, lectin, proline-rich protein, hydroxyproline-rich glycoprotein, glycine-rich protein, arabinogalactan-protein and so on, and the activity ofβ-1,3-glucanase andβ-D-xylosidase were enhanced, which were related to cell wall synthesis or degradation.
Base on these results, the mechanism of GbaVd1 and GbaVd2 resistance model was supposed, two genes encode LRR-TM receptor like protein could directly or indirectly combine with the V. dahliae elicitors, this action would regulate the expression of signal factor of disease resistance, then active the ethylene and jasmonic acid signal transduction pathway which lead to change the synthesis of phytoalexin and expression of cell wall related proteins, form the antifungus activity compound and enhance the cell wall structure obstacle, finally increase the host resistance to V. dahliae.
引文
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