水稻白叶枯病抗性突变体的筛选与高分辨率溶解曲线在水稻中的应用
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摘要
水稻白叶枯病是水稻重要病害之一,对水稻生产造成了严重的危害。新的白叶枯病抗性种质资源的获得对于水稻抗白叶枯病育种具有重要意义。
本研究对5万个T-DNA突变体株系进行了系统的大田筛选。从5叶期开始,每个株系经过4轮的白叶枯病菌P10接种,研究初筛获得了一些白叶枯病抗性植株,对其中2个表型较好的植株进行了初步的生理研究。同时,在突变体库中观察获得了一些其他表型突变的株系。为加快水稻突变体基因定位,本研究尝试利用高分辨率溶解曲线技术(high resolution melting, HRM)进行水稻F2群体基因分型,为后期将HRM用于突变体中抗性基因定位探索条件。目前,已取得的主要研究结果有：
1.通过T-DNA突变体库大田种植观察及接种P10菌株,初筛获得了125个M0代白叶枯病抗性突变体。同时,获得了一些具有其他表型变化的突变体株系,如类病变突变体,叶色突变体,颖花退化,穗颈长度变化,有芒与无芒,颖壳变化,高矮秆突变,叶形突变等。
2.对初筛获得的125个M0代白叶枯病抗性突变体中的17个株系的M1代进行了对P10菌株的抗性鉴定,结果表明2个株系BBM14和BBM66的M1代对P10菌株抗性稳定,且BBM14和BBM66的M1代植株无明显抗性分离。同时,转基因PCR检测表明T-DNA插入已经纯合,因此,需要设计专门实验来判断BBM14和BBM66的抗性是否由T-DNA插入引起。对接种P10后叶片内部的菌株生长情况进行生长曲线分析表明,P10菌株在BBM14和BBM66中的生长相对于亲本中花11明显受到抑制,由此进一步证实了BBM14和BBM66对P10菌株的抗性。对BBM14和BBM66的抗谱分析发现,BBM14对白叶枯病菌株菲律宾生理小种P2 (PXO86), P3 (PXO79), P7 (PXO145), P8(PXO280), P10 (PXO124)表现抗性,对P4 (PXO71), P6(PXO99)表现感病。BBM66对P2,P7,P8,P10菌株表现出抗性,对P3,P4,P6表现感病。同时,将BBM14和BBM66抗谱与已知白叶枯病抗性基因抗谱比较后发现,两者的抗谱与这些白叶枯病抗性基因抗谱都不同,因此,初步判断BBM14和BBM66中均具有新的白叶枯病抗性基因。对BBM14和BBM66中的部分防卫相关基因进行表达分析,结果表明接种P10菌株后OsPRlb基因表达上调。
3.本研究尝试将HRM用于水稻F2群体基因分型,同时进行了HRM反应条件优化,为后续将HRM用于BBM14和BBM66中抗性基因定位打下了基础。利用本实验室创制的高抗白叶枯病材料Y73以及高感白叶枯病材料IR24杂交以构建F2群体,选取了22株F2植株。在5μl反应体系及普通PCR程序条件下,本研究对353对STS和SSR分子标记进行了Y73,IR24以及其F1之间多态HRM分子标记筛选,其中103对分子标记表现出多态的HRM曲线。将这103对多态HRM分子标记用于以上F2群体的HRM基因分型,结果显示在5μl反应体系及普通PCR程序条件下,12对STS和SSR分子标记成功地对该F2群体进行了基因型分析。传统的凝胶电泳结果显示,HRM基因分型结果与传统的PCR后电泳检测结果完全一致。为提高STS与SSR分子标记用于HRM基因分型的效率,本研究在另外两种HRM反应条件下,对这103对多态HRM分子标记用于该F2群体基因分型进行了重新的筛选。在5μl反应体系与降落PCR程序中,筛选效率没有明显提高,而在10μl反应体系与降落PCR程序中,除了这12对分子标记外,另外9对分子标记成功地对该F2群体进行了基因分型,筛选效率明显提高。因此,以上结果表明HRM用于水稻F2群体的基因分型具有可行性,且3种HRM体系中,10μl反应体系与降落PCR程序条件时的HRM体系更稳定,更适宜于水稻F2群体基因分型。
综合以上研究结果表明,突变体BBM14和BBM66对部分白叶枯病菌抗性稳定,可进行进一步的白叶枯病抗性基因研究。同时,HRM技术在水稻F2群体基因分型中具有可行性,因此,将HRM技术应用于BBM14和BBM66相关抗性基因的研究对于同时加快白叶枯病新抗性基因的定位以及推进HRM基因分型技术在水稻中的应用具有双重作用。
Bacterial blight is one of the most serious rice diseases, causing great damage to rice production. Novel bacterial blight resistance cultivars play an important role in bacterial blight resistance breeding in rice.
In this study, a large-scale screening was performed on a rice mutant population of 50,000 T-DNA insertion lines in the field. Plants at five-leaf stage were inoculated with Xanthomonas oryzae pv.oryzae (Xoo) P10 by cutting the leaf top with scissors and sprayed for at least four times. Some candidate bacterial blight resistance mutants were preliminarily screened from the population. Among them, two mutant lines were selected for further researches. At the same time, some other phenotype mutants were observed in the library. To make preparations for mapping the resistance genes in these mutants, high-resolution melting (HRM) was used to genotype the F2 population in rice. Major research results were summarized as follows:
1. A total of 125 bacterial blight resistance mutants' M0 lines were screened from the T-DNA mutant library. Simultaneously, some other phenotype mutants were observed, such as leaf colour mutants, lesion mimic mutant, degenerated spikelet, changes of ear stem length, beared and beardless rice, rice glume mutant, dwarf and tall-culm mutant, leaf shape mutant.
2. Resistance of 17 mutants' M1 progenies from the 125 lines was identified by the P10 strain. It shows that the M1 progenies of two mutants BBM14 and BBM66 were resistant to P10 strain stably and hereditably. However, no resistance separation was observed among the M1 progenies and transgene PCR detection shows that T-DNA insertion was homozygous, so further researches are needed to determine whether the resistance of BBM14 and BBM66 to the P10 strain was caused by T-DNA insertion was uncertain. The growth curves of the P10 strain in BBM14 and BBM66 leaves were analyzed, it shows that bacterial growth in the two mutants was suppressed compared to the parent zhonghuall, this confirmed the resistance of BBM14 and BBM66 to P10 strain further. Resistant spectrum showed that BBM14 was resistant to the strains P2 (PXO86), P3 (PXO79), P7 (PXO145), P8 (PXO280), and P10 (PXO124), and susceptible to the strains P4 (PXO71), P6 (PXO99). While BBM66 was resistant to the strains P2, P7, P8, P10 and susceptible to the strains P3, P4, P6. Comparing the resistance spectrum of BBM14 and BBM66 to the known bacterial blight resistance genes in this research, it was suggested that there were new bacterial blight resistance genes in the two mutants. The expression of some defense related genes were analyzed in the mutants after inoculated by P10, and it shows that OsPR1b was upregulated after P10 strain inoculation.
3. HRM technique was employed for F2 population genotyping in rice and the HRM reaction conditions were optimizated to make preparations for mapping the resistance genes in BBM14 and BBM66 For optimizing the reaction conditions, Twenty-two F2 lines were selected from the cross between the highly resistant cultivar Y73 and the susceptible cultivar IR24. With the 5μl reaction system and normal PCR procedure, a total of 353 STS and SSR markers were used to screen the markers showing polymorphic HRM curves between Y73, IR24 and F1. A total of 103 STS and SSR markers showed polymorphic melting curves among the three parents. These 103 markers were further used to genotype the F2 population. It showed that 12 STS and SSR markers genotyped the mapping population successfully. And the HRM genotyping results were consistent with traditional electrophoresis results completely. To raise the efficiency of STS and SSR markers for HRM genotyping, another two HRM reaction conditions were applied for screening the 103 polymorphic markers. In a 5μ1 reaction system and touchdown-PCR procedure, no screening efficiency improvement was observed. However, in a 10μl reaction system and touchdown-PCR procedure,9 new markers genotyped the mapping population successfully except for the 12 markers, HRM marker screening efficiency increased greatly. Therefore,it is practical for applying HRM in F2 genotyping in rice. And among the three HRM reaction conditions, the 10μl reaction system and touchdown-PCR procedure was more stable and suitable for genotyping in rice.
From the above results, it shows that BBM14 and BBM66 were resistant to part of the bacterial blight strains stably. Meanwhile, HRM was useful for F2 population genotyping in rice. So applying HRM for mapping the resistance gene in BBM14 and BBM66 can accelerate not only the gene identification but also the application of HRM genotyping in rice.
本研究对5万个T-DNA突变体株系进行了系统的大田筛选。从5叶期开始,每个株系经过4轮的白叶枯病菌P10接种,研究初筛获得了一些白叶枯病抗性植株,对其中2个表型较好的植株进行了初步的生理研究。同时,在突变体库中观察获得了一些其他表型突变的株系。为加快水稻突变体基因定位,本研究尝试利用高分辨率溶解曲线技术(high resolution melting, HRM)进行水稻F2群体基因分型,为后期将HRM用于突变体中抗性基因定位探索条件。目前,已取得的主要研究结果有：
1.通过T-DNA突变体库大田种植观察及接种P10菌株,初筛获得了125个M0代白叶枯病抗性突变体。同时,获得了一些具有其他表型变化的突变体株系,如类病变突变体,叶色突变体,颖花退化,穗颈长度变化,有芒与无芒,颖壳变化,高矮秆突变,叶形突变等。
2.对初筛获得的125个M0代白叶枯病抗性突变体中的17个株系的M1代进行了对P10菌株的抗性鉴定,结果表明2个株系BBM14和BBM66的M1代对P10菌株抗性稳定,且BBM14和BBM66的M1代植株无明显抗性分离。同时,转基因PCR检测表明T-DNA插入已经纯合,因此,需要设计专门实验来判断BBM14和BBM66的抗性是否由T-DNA插入引起。对接种P10后叶片内部的菌株生长情况进行生长曲线分析表明,P10菌株在BBM14和BBM66中的生长相对于亲本中花11明显受到抑制,由此进一步证实了BBM14和BBM66对P10菌株的抗性。对BBM14和BBM66的抗谱分析发现,BBM14对白叶枯病菌株菲律宾生理小种P2 (PXO86), P3 (PXO79), P7 (PXO145), P8(PXO280), P10 (PXO124)表现抗性,对P4 (PXO71), P6(PXO99)表现感病。BBM66对P2,P7,P8,P10菌株表现出抗性,对P3,P4,P6表现感病。同时,将BBM14和BBM66抗谱与已知白叶枯病抗性基因抗谱比较后发现,两者的抗谱与这些白叶枯病抗性基因抗谱都不同,因此,初步判断BBM14和BBM66中均具有新的白叶枯病抗性基因。对BBM14和BBM66中的部分防卫相关基因进行表达分析,结果表明接种P10菌株后OsPRlb基因表达上调。
3.本研究尝试将HRM用于水稻F2群体基因分型,同时进行了HRM反应条件优化,为后续将HRM用于BBM14和BBM66中抗性基因定位打下了基础。利用本实验室创制的高抗白叶枯病材料Y73以及高感白叶枯病材料IR24杂交以构建F2群体,选取了22株F2植株。在5μl反应体系及普通PCR程序条件下,本研究对353对STS和SSR分子标记进行了Y73,IR24以及其F1之间多态HRM分子标记筛选,其中103对分子标记表现出多态的HRM曲线。将这103对多态HRM分子标记用于以上F2群体的HRM基因分型,结果显示在5μl反应体系及普通PCR程序条件下,12对STS和SSR分子标记成功地对该F2群体进行了基因型分析。传统的凝胶电泳结果显示,HRM基因分型结果与传统的PCR后电泳检测结果完全一致。为提高STS与SSR分子标记用于HRM基因分型的效率,本研究在另外两种HRM反应条件下,对这103对多态HRM分子标记用于该F2群体基因分型进行了重新的筛选。在5μl反应体系与降落PCR程序中,筛选效率没有明显提高,而在10μl反应体系与降落PCR程序中,除了这12对分子标记外,另外9对分子标记成功地对该F2群体进行了基因分型,筛选效率明显提高。因此,以上结果表明HRM用于水稻F2群体的基因分型具有可行性,且3种HRM体系中,10μl反应体系与降落PCR程序条件时的HRM体系更稳定,更适宜于水稻F2群体基因分型。
综合以上研究结果表明,突变体BBM14和BBM66对部分白叶枯病菌抗性稳定,可进行进一步的白叶枯病抗性基因研究。同时,HRM技术在水稻F2群体基因分型中具有可行性,因此,将HRM技术应用于BBM14和BBM66相关抗性基因的研究对于同时加快白叶枯病新抗性基因的定位以及推进HRM基因分型技术在水稻中的应用具有双重作用。
Bacterial blight is one of the most serious rice diseases, causing great damage to rice production. Novel bacterial blight resistance cultivars play an important role in bacterial blight resistance breeding in rice.
In this study, a large-scale screening was performed on a rice mutant population of 50,000 T-DNA insertion lines in the field. Plants at five-leaf stage were inoculated with Xanthomonas oryzae pv.oryzae (Xoo) P10 by cutting the leaf top with scissors and sprayed for at least four times. Some candidate bacterial blight resistance mutants were preliminarily screened from the population. Among them, two mutant lines were selected for further researches. At the same time, some other phenotype mutants were observed in the library. To make preparations for mapping the resistance genes in these mutants, high-resolution melting (HRM) was used to genotype the F2 population in rice. Major research results were summarized as follows:
1. A total of 125 bacterial blight resistance mutants' M0 lines were screened from the T-DNA mutant library. Simultaneously, some other phenotype mutants were observed, such as leaf colour mutants, lesion mimic mutant, degenerated spikelet, changes of ear stem length, beared and beardless rice, rice glume mutant, dwarf and tall-culm mutant, leaf shape mutant.
2. Resistance of 17 mutants' M1 progenies from the 125 lines was identified by the P10 strain. It shows that the M1 progenies of two mutants BBM14 and BBM66 were resistant to P10 strain stably and hereditably. However, no resistance separation was observed among the M1 progenies and transgene PCR detection shows that T-DNA insertion was homozygous, so further researches are needed to determine whether the resistance of BBM14 and BBM66 to the P10 strain was caused by T-DNA insertion was uncertain. The growth curves of the P10 strain in BBM14 and BBM66 leaves were analyzed, it shows that bacterial growth in the two mutants was suppressed compared to the parent zhonghuall, this confirmed the resistance of BBM14 and BBM66 to P10 strain further. Resistant spectrum showed that BBM14 was resistant to the strains P2 (PXO86), P3 (PXO79), P7 (PXO145), P8 (PXO280), and P10 (PXO124), and susceptible to the strains P4 (PXO71), P6 (PXO99). While BBM66 was resistant to the strains P2, P7, P8, P10 and susceptible to the strains P3, P4, P6. Comparing the resistance spectrum of BBM14 and BBM66 to the known bacterial blight resistance genes in this research, it was suggested that there were new bacterial blight resistance genes in the two mutants. The expression of some defense related genes were analyzed in the mutants after inoculated by P10, and it shows that OsPR1b was upregulated after P10 strain inoculation.
3. HRM technique was employed for F2 population genotyping in rice and the HRM reaction conditions were optimizated to make preparations for mapping the resistance genes in BBM14 and BBM66 For optimizing the reaction conditions, Twenty-two F2 lines were selected from the cross between the highly resistant cultivar Y73 and the susceptible cultivar IR24. With the 5μl reaction system and normal PCR procedure, a total of 353 STS and SSR markers were used to screen the markers showing polymorphic HRM curves between Y73, IR24 and F1. A total of 103 STS and SSR markers showed polymorphic melting curves among the three parents. These 103 markers were further used to genotype the F2 population. It showed that 12 STS and SSR markers genotyped the mapping population successfully. And the HRM genotyping results were consistent with traditional electrophoresis results completely. To raise the efficiency of STS and SSR markers for HRM genotyping, another two HRM reaction conditions were applied for screening the 103 polymorphic markers. In a 5μ1 reaction system and touchdown-PCR procedure, no screening efficiency improvement was observed. However, in a 10μl reaction system and touchdown-PCR procedure,9 new markers genotyped the mapping population successfully except for the 12 markers, HRM marker screening efficiency increased greatly. Therefore,it is practical for applying HRM in F2 genotyping in rice. And among the three HRM reaction conditions, the 10μl reaction system and touchdown-PCR procedure was more stable and suitable for genotyping in rice.
From the above results, it shows that BBM14 and BBM66 were resistant to part of the bacterial blight strains stably. Meanwhile, HRM was useful for F2 population genotyping in rice. So applying HRM for mapping the resistance gene in BBM14 and BBM66 can accelerate not only the gene identification but also the application of HRM genotyping in rice.
引文
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