小麦赤霉病菌DON毒素及其降解基因Tri101的克隆和遗传转化研究
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摘要
赤霉病(Fusarium head blight, FHB)是由镰刀菌引起的谷类作物病害之一,其产生的单族毒素对人、畜毒性大,可引起多种中毒症状。禾谷镰刀菌(Fusarium graminearium Schwabe) Tri101基因编码的单端孢酶烯3-O-乙酰转移酶通过加乙酰基的形式使单族毒素降解为低毒物质。本文研究了禾谷镰刀菌的DON毒素及其降解基因Tri101的克隆、表达和在小麦中的遗传转化,为作物脱毒和有效防治赤霉病奠定理论基础,为经济、安全和高效净化食物和饲料中真菌毒素提供科学依据。主要包括:
1.野生型禾谷镰刀菌0623(F. graminearum 0623,简称Fg0623)在PDA平板上28℃分别培养3d、5d、7d,气生菌丝生长旺盛,中心菌丝在培养第7d开始老化;菌落基质由白色变为红色,部分基质呈深红色;在绿豆汤培养基中25℃、180rpm振动培养3d时,其镰刀状孢子清晰可见。在小麦培养基上25℃分别培养10d、20d、30d、40d、50d,高效液相色谱法测定DON的含量,结果显示DON产量与培养时间呈近似抛物线型关系,在30d时最高,30d后迅速下降。
2.利用RT-PCR克隆Fg0623的Tri101基因cDNA片段并测序,Tri101基因核苷酸序列阅读框架全长1 356bp(GenBank序列号:GQ907236),编码451个氨基酸的多肽,推测分子量为49.45 kDa,等电点为5.14。氨基酸序列同源性比对结果表明,它与Kimura报道的禾谷镰刀菌Tri101氨基酸序列同源性最高,为99.56%,与其它13种镰刀菌的Tri101氨基酸序列同源性分别为97.91%~75.68%。系统进化树分析结果表明,Fg0623与F. sporotrichioides属于同一进化枝且与F. asiaticum有较近的亲缘关系,而与F. oxysporum、F. moniliforme、F. nygamai、F. nisikadoi和F. decemcellulare的亲缘关系较远。
3.构建的原核表达载体pGEX-4T2/Tri101转化大肠杆菌BL21,用IPTG进行诱导表达。SDS-PAGE分析表明,Tri101基因在BL21中高效表达,融合蛋白GST-Tri101分子量为75.45 kDa。将该融合蛋白切胶纯化制备兔抗GST-Tri101多克隆抗体,经ELISA法测定抗体效价为1: 256 000,Western blot分析表明制备的抗体与原核细胞体外表达的Tri101蛋白可以特异性结合。应用该抗体验证了感赤霉病小麦中Tri101基因的表达。
4.采用农杆菌介导的小麦生长点转化法将pCAMBIA1301-Tri101-EHA105转入小麦品种豫麦18、郑麦366和郑麦9962中。100mg/L潮霉素筛选后的转化植株用PCR进一步验证,获得T0代小麦植株分别为32、15和29株,GUS染色结果显示有蓝色位点,证明Tri101基因已经整合进小麦基因组中并成功表达。
Fusarium head blight (FHB), caused by Fusarium spp., is one of major disease of grain species. Fusarium graminearum Tri101 encodes a trichothecene 3-O-acetyltransferase that converts DON to a less toxic acetylated form. The main purpose of this paper is to research deoxynivalenol toxin of Fusarium graminearum and cloning, expression and transformation of toxicity degradation gene Tri101 in wheat, lay the theoretical basis for the detoxification of food crops and the effective control of the F. graminearum,and provide the scientific bases for searching for economy, safety and high efficiency purification method of mycotoxin in food and feed. The main results were summarized as follows:
1. The basic morphology of Fusarium graminearum 0623 was observed by culturing it for 3d , 5d and 7d in PDA medium. Aerial mycelium grew more and more luxuriantly. The central old mycelium was sunk after 7d. With increasing day of culture, colonial stroma slowly changed from white to red, and a part of it was dark red. The spore morphology of it in greenbean soup for 3d was sickle shaped. Fusarium graminearum 0623 was cultured for 10d, 20d, 30d, 40d and 50d in wheat medium in order to detect DON content in different times with HPLC. Tests showed DON content was most highly when it was cultured for thirty days. However, DON content was fast setback after 30d.
2. The cDNA of Tri101 was amplied from the Fusarium graminearum 0623 total RNA by RT-PCR. The results showed that Tri101 gene was 1356 bp in size and encoded 451 amino acids (GenBank accession number:GQ907236).The molecular weight (Mw) and theoretical isoelectric point (pI) is 49.45 kD and 5.14 respectively. The deduced amino acid of Fusarium graminearum 0623 Tri101 gene shared 99.56% homology with the deduced amino acid published by Kimura in GenBank. Alignments with the deduced amino acid of mature proteins in other 13 species of Fusarium showed 97.91%~75.68%. Based on Tri101 amino acid sequence the phylogenic tree of trichothecene 3-O-acetyltransferase gene was established. The results showed that Fg0623 and F. sporotrichioides belong to the same group of organisms; it was closely related to F. asiaticum while distantly related to F. oxysporum, F. moniliforme, F. nygamai, F. nisikadoi and F. decemcellulare.
3. The Tri101 gene was ligated to the expression vector pGEX-4T2. The recombinant plasmid, pGEX-4T2/Tri101 was then transformed into E.coli BL21 strain, induced by IPTG. The results of SDS-PAGE and Western blot analysis showed that the Tri101 gene was highly expressed. The molecular weight of the recombinant fusion protein was 75.45 kDa, consistent with the predicted result. Polyclonal anti-GST-Tri101 rabbit antibody was successfully prepared by using purified Tri101 protein as immunogen. The ELISA titer of antiserum against GST-Tri101 was about 1∶2 56 000. Western blot analysis showed that the antiserum could specifically bind to the expressed Tri101 protein. Trichothecene 3-O-acetyltransferase encoded by Tri101 gene in Fusarium-damaged kernels of wheat was successfully detected with the antibody.
4. The plant expression vector pCAMBIA1301-Tri101, which was structured successfully, was transformed into Agrobacterium tumefaciens EHA105 by liquid nitrogen cryopreservation. Shoot apical meristem cells of YuMai 18, ZhengMai 366 and ZhengMai 9962 of were transformed by pCAMBIA1301-Tri101-EHA105. After selected with solution containing 100mg/L hygromyci, the Tri101 gene transgenic seedlings were then proved by PCR. The result indicated that 32, 15 and 29 T0 plants were obtained. GUS staining identification showed that blue dots.It was proved that the Tri101 genes were integrated into the wheat genomes and they could express in the transgenic wheat.
1.野生型禾谷镰刀菌0623(F. graminearum 0623,简称Fg0623)在PDA平板上28℃分别培养3d、5d、7d,气生菌丝生长旺盛,中心菌丝在培养第7d开始老化;菌落基质由白色变为红色,部分基质呈深红色;在绿豆汤培养基中25℃、180rpm振动培养3d时,其镰刀状孢子清晰可见。在小麦培养基上25℃分别培养10d、20d、30d、40d、50d,高效液相色谱法测定DON的含量,结果显示DON产量与培养时间呈近似抛物线型关系,在30d时最高,30d后迅速下降。
2.利用RT-PCR克隆Fg0623的Tri101基因cDNA片段并测序,Tri101基因核苷酸序列阅读框架全长1 356bp(GenBank序列号:GQ907236),编码451个氨基酸的多肽,推测分子量为49.45 kDa,等电点为5.14。氨基酸序列同源性比对结果表明,它与Kimura报道的禾谷镰刀菌Tri101氨基酸序列同源性最高,为99.56%,与其它13种镰刀菌的Tri101氨基酸序列同源性分别为97.91%~75.68%。系统进化树分析结果表明,Fg0623与F. sporotrichioides属于同一进化枝且与F. asiaticum有较近的亲缘关系,而与F. oxysporum、F. moniliforme、F. nygamai、F. nisikadoi和F. decemcellulare的亲缘关系较远。
3.构建的原核表达载体pGEX-4T2/Tri101转化大肠杆菌BL21,用IPTG进行诱导表达。SDS-PAGE分析表明,Tri101基因在BL21中高效表达,融合蛋白GST-Tri101分子量为75.45 kDa。将该融合蛋白切胶纯化制备兔抗GST-Tri101多克隆抗体,经ELISA法测定抗体效价为1: 256 000,Western blot分析表明制备的抗体与原核细胞体外表达的Tri101蛋白可以特异性结合。应用该抗体验证了感赤霉病小麦中Tri101基因的表达。
4.采用农杆菌介导的小麦生长点转化法将pCAMBIA1301-Tri101-EHA105转入小麦品种豫麦18、郑麦366和郑麦9962中。100mg/L潮霉素筛选后的转化植株用PCR进一步验证,获得T0代小麦植株分别为32、15和29株,GUS染色结果显示有蓝色位点,证明Tri101基因已经整合进小麦基因组中并成功表达。
Fusarium head blight (FHB), caused by Fusarium spp., is one of major disease of grain species. Fusarium graminearum Tri101 encodes a trichothecene 3-O-acetyltransferase that converts DON to a less toxic acetylated form. The main purpose of this paper is to research deoxynivalenol toxin of Fusarium graminearum and cloning, expression and transformation of toxicity degradation gene Tri101 in wheat, lay the theoretical basis for the detoxification of food crops and the effective control of the F. graminearum,and provide the scientific bases for searching for economy, safety and high efficiency purification method of mycotoxin in food and feed. The main results were summarized as follows:
1. The basic morphology of Fusarium graminearum 0623 was observed by culturing it for 3d , 5d and 7d in PDA medium. Aerial mycelium grew more and more luxuriantly. The central old mycelium was sunk after 7d. With increasing day of culture, colonial stroma slowly changed from white to red, and a part of it was dark red. The spore morphology of it in greenbean soup for 3d was sickle shaped. Fusarium graminearum 0623 was cultured for 10d, 20d, 30d, 40d and 50d in wheat medium in order to detect DON content in different times with HPLC. Tests showed DON content was most highly when it was cultured for thirty days. However, DON content was fast setback after 30d.
2. The cDNA of Tri101 was amplied from the Fusarium graminearum 0623 total RNA by RT-PCR. The results showed that Tri101 gene was 1356 bp in size and encoded 451 amino acids (GenBank accession number:GQ907236).The molecular weight (Mw) and theoretical isoelectric point (pI) is 49.45 kD and 5.14 respectively. The deduced amino acid of Fusarium graminearum 0623 Tri101 gene shared 99.56% homology with the deduced amino acid published by Kimura in GenBank. Alignments with the deduced amino acid of mature proteins in other 13 species of Fusarium showed 97.91%~75.68%. Based on Tri101 amino acid sequence the phylogenic tree of trichothecene 3-O-acetyltransferase gene was established. The results showed that Fg0623 and F. sporotrichioides belong to the same group of organisms; it was closely related to F. asiaticum while distantly related to F. oxysporum, F. moniliforme, F. nygamai, F. nisikadoi and F. decemcellulare.
3. The Tri101 gene was ligated to the expression vector pGEX-4T2. The recombinant plasmid, pGEX-4T2/Tri101 was then transformed into E.coli BL21 strain, induced by IPTG. The results of SDS-PAGE and Western blot analysis showed that the Tri101 gene was highly expressed. The molecular weight of the recombinant fusion protein was 75.45 kDa, consistent with the predicted result. Polyclonal anti-GST-Tri101 rabbit antibody was successfully prepared by using purified Tri101 protein as immunogen. The ELISA titer of antiserum against GST-Tri101 was about 1∶2 56 000. Western blot analysis showed that the antiserum could specifically bind to the expressed Tri101 protein. Trichothecene 3-O-acetyltransferase encoded by Tri101 gene in Fusarium-damaged kernels of wheat was successfully detected with the antibody.
4. The plant expression vector pCAMBIA1301-Tri101, which was structured successfully, was transformed into Agrobacterium tumefaciens EHA105 by liquid nitrogen cryopreservation. Shoot apical meristem cells of YuMai 18, ZhengMai 366 and ZhengMai 9962 of were transformed by pCAMBIA1301-Tri101-EHA105. After selected with solution containing 100mg/L hygromyci, the Tri101 gene transgenic seedlings were then proved by PCR. The result indicated that 32, 15 and 29 T0 plants were obtained. GUS staining identification showed that blue dots.It was proved that the Tri101 genes were integrated into the wheat genomes and they could express in the transgenic wheat.
引文
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