影响水稻抗白叶枯病主效基因Xa3/Xa26抗性发挥的遗传背景因素分析
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摘要
水稻白叶枯病是由黄单胞杆菌水稻致病变种(Xanthomonas oryzae pv. oryzae)引起的一种细菌性维管束病害,自二十世纪五十年代末以来,一直是影响我国水稻高产、稳产的主要病害之一。充分利用抗病基因资源,培育具有高效广谱抗性的优良品种是最经济有效地防治白叶枯病的技术措施。目前已经鉴定并克隆了大量主效抗病基因,部分基因在不同遗传背景中的抗病表型存在差异。研究清楚遗传背景如何影响抗病基因功能的发挥,为更好地利用主效抗病基因培育优良品种有着重要的指导意义。
从明恢63中鉴定并克隆的水稻抗白叶枯病主效基因Xa3/Xa26定位于第11染色体的长臂末端。该基因介导的抗性受遗传背景的影响,粳稻背景可以提高该基因的表达水平,随着表达量的增加植株的抗性也随之增强。为了证实Xa3/Xa26的剂量效应是否是影响其抗性发挥的唯一因素,本研究从基因表达谱和遗传学两个角度分析了遗传背景影响Xa3/Xa26抗性的原因。
首先,从牡丹江8/明恢63 F2分离群体中分别于苗期和成株期随机取材分析Xa3/Xa26的表达水平与植株抗性之间的关系,发现不管是在苗期还是成株期,两者之间均有一定的相关性,然而少数单株对PX061的抗性并不受抗病基因表达水平的影响。这说明遗传背景的确可以影响Xa3/Xa26的表达量,但该基因的剂量效应并非影响植株抗性的唯一因素。
其次,从牡丹江8／明恢63的F2群体和以明恢63为轮回亲本的多个回交群体中总共检测到5个QTLs,这些QTL位点的牡丹江8等位位点对增强植株抗性起到一定作用,即感病亲本中确实存在一些数量抗性位点,可以影响Xa3/Xa26对白叶枯病的抗性。还扫描到4个QTL位点,当它们为明恢63基因型时也可以增强植株的抗性。另外,对129个共显性标记进行两位点互作分析,发现这些QTLs之间几乎不存在互作关系,以非QTL之间的互作最多。这些结果都说明遗传背景中有很多位点可以影响Xa3/Xa26介导的抗性。
第三,用cDNA芯片对明恢63和Rb49接种白叶枯病病原后的基因表达谱进行分析,其中Rb49是一个携带有自身启动子驱动的Xa3/Xa26基因的单拷贝转基因株系。检测到的大多数差异表达基因在Rb49中能被病原快速激活,但它们在明恢63中却受到抑制表达或表达量基本不变。编码SNAC1蛋白的Os03g60080在明恢63和Rb49中均受病原的抑制表达,但从各时间点的表达趋势来看,该基因的表达模式与Xa3/Xa26, OsWRKY13和NH1相反,且它在OsWRKY13超量表达的转基因植株中的表达受抑制,这些都说明了这个基因不仅与非生物胁迫相关,而且很可能参与了Xa3/Xa26介导的抗病信号传导。结果表明某些基因在时空表达模式上的差异也是遗传背景影响主效抗病基因发挥功能的重要原因之一。
第四,通过生物信息学的方法将差异表达基因定位到遗传连锁图上,且部分基因位于QTLs置信区间内,那么这些基因在不同的遗传背景中可能通过不同的调节功能影响Xa3/Xa26介导的抗病反应。
Bacterial blight of rice is a vascular disease caused by Xanthomonas oryzae pv. oryzae. It became one of the most devastating diseases influencing the rice production in the late 1950s. The effective method to improve the rice disease resistance is taking the full advantage of host resistance genes to develop better varieties. Until now, a large number of resistance genes have been identified, while the resistant phenotypes of some resistance genes are diverse in different genetic background. Understanding how the genetic background affecting the R gene mediated resistance may facilitate molecular breeding to obtain higher resistance varieties.
Xa3/Xa26 was first identified in the Oryza sativa L. ssp. indica rice variety Minghui 63; it was mapped to the long arm of chromosome 11. The function of bacterial-blight resistance gene Xa3/Xa26 in rice is influenced by genetic background; Oryza sativa L. ssp. japonica background can increase Xa3/Xa26 expression, resulting in enhanced resistance. To identify whether Xa3/Xa26 transcript level is the only factor contributing to genetic background-controlled resistance, we combined the analyses of quantitative genetics and gene expression profile in this study.
First, some juvenile and adult plants were randomly chosen from a Mudanjiang 8/Minghui 63 F2 population to analyze the relationship between the transcription level of Xa3/Xa26 and resistance level to bacterial blight. Increasing Xa3/Xa26 expression was correlated with the enhanced resistance in theses F2 plants both at juvenile and adult stages. But a few F2 plants showed a disassociation of Xa3/Xa26 expression level and enhanced resistance. These results suggest that genetic background does influence Xa3/Xa26 expression and its expression level is not an exclusive factor which affects its function.
Second, five quantitative trait loci (QTLs) were identify from a Mudanjiang 8/Minghui 63 F2 population and some backcross populations using the Minghui 63 as a recurrent parent. The resistance alleles at these five QTLs were from the susceptible parent Mudanjiang 8. It means that some loci contributed by susceptible parent can influence Xa3/Xa26-mediated Xoo resistance. The other four QTLs also can enhance resistance when these alleles were homozygous for Minghui 63. Furthermore, One hundred and twenty-nine co-dominant markers were used for testing digenic interactions in this F2 segregating population. Few epistatic effects were existed between these QTLs. Most digenic interactions were detected between non-significant effect loci (non-QTL). These results suggest that Xa3/Xa26-mediated resistance was affected by multiple loci, including QTLs and epistatic interactions.
Third, analysis difference of the expression profile of Xa3/Xa26-mediated resistance between Minghui 63 and Rb49, using a cDNA microarray. Rb49 is a transgenic line carrying a single copy of Xa3/Xa26 regulated by its native promoter in the genetic background of susceptible Mudanjiang 8. Most differentially expressed genes were rapidly activated in the Rb49, but not in the indica background, during disease resistance. Os03g60080, encoding a SNAC1 protein, was suppressed by pathogen in both Minghui 63 and Rb49, while its expression pattern of four time point after inoculation was opposite to that of Xa3/Xa26,OsWRKY13 and NH1 This gene was also suppressed in OsWRKY13-overexpressing lines. It was indicated that the differentially expressed gene not only related with abiotic stress, but also participated in the signal transduction of Xa3/Xa26-mediated resistance. It means that the different spatiotemporal expression pattern of some genes was another important factor affecting the R gene function.
Fourth, the colocalization of some differentially expressed genes with QTLs in Xa3/Xa26-mediated resistance will help to characterize the genes underlying these QTLs and involved in regulation of genetic background-controlled resistance.
从明恢63中鉴定并克隆的水稻抗白叶枯病主效基因Xa3/Xa26定位于第11染色体的长臂末端。该基因介导的抗性受遗传背景的影响,粳稻背景可以提高该基因的表达水平,随着表达量的增加植株的抗性也随之增强。为了证实Xa3/Xa26的剂量效应是否是影响其抗性发挥的唯一因素,本研究从基因表达谱和遗传学两个角度分析了遗传背景影响Xa3/Xa26抗性的原因。
首先,从牡丹江8/明恢63 F2分离群体中分别于苗期和成株期随机取材分析Xa3/Xa26的表达水平与植株抗性之间的关系,发现不管是在苗期还是成株期,两者之间均有一定的相关性,然而少数单株对PX061的抗性并不受抗病基因表达水平的影响。这说明遗传背景的确可以影响Xa3/Xa26的表达量,但该基因的剂量效应并非影响植株抗性的唯一因素。
其次,从牡丹江8／明恢63的F2群体和以明恢63为轮回亲本的多个回交群体中总共检测到5个QTLs,这些QTL位点的牡丹江8等位位点对增强植株抗性起到一定作用,即感病亲本中确实存在一些数量抗性位点,可以影响Xa3/Xa26对白叶枯病的抗性。还扫描到4个QTL位点,当它们为明恢63基因型时也可以增强植株的抗性。另外,对129个共显性标记进行两位点互作分析,发现这些QTLs之间几乎不存在互作关系,以非QTL之间的互作最多。这些结果都说明遗传背景中有很多位点可以影响Xa3/Xa26介导的抗性。
第三,用cDNA芯片对明恢63和Rb49接种白叶枯病病原后的基因表达谱进行分析,其中Rb49是一个携带有自身启动子驱动的Xa3/Xa26基因的单拷贝转基因株系。检测到的大多数差异表达基因在Rb49中能被病原快速激活,但它们在明恢63中却受到抑制表达或表达量基本不变。编码SNAC1蛋白的Os03g60080在明恢63和Rb49中均受病原的抑制表达,但从各时间点的表达趋势来看,该基因的表达模式与Xa3/Xa26, OsWRKY13和NH1相反,且它在OsWRKY13超量表达的转基因植株中的表达受抑制,这些都说明了这个基因不仅与非生物胁迫相关,而且很可能参与了Xa3/Xa26介导的抗病信号传导。结果表明某些基因在时空表达模式上的差异也是遗传背景影响主效抗病基因发挥功能的重要原因之一。
第四,通过生物信息学的方法将差异表达基因定位到遗传连锁图上,且部分基因位于QTLs置信区间内,那么这些基因在不同的遗传背景中可能通过不同的调节功能影响Xa3/Xa26介导的抗病反应。
Bacterial blight of rice is a vascular disease caused by Xanthomonas oryzae pv. oryzae. It became one of the most devastating diseases influencing the rice production in the late 1950s. The effective method to improve the rice disease resistance is taking the full advantage of host resistance genes to develop better varieties. Until now, a large number of resistance genes have been identified, while the resistant phenotypes of some resistance genes are diverse in different genetic background. Understanding how the genetic background affecting the R gene mediated resistance may facilitate molecular breeding to obtain higher resistance varieties.
Xa3/Xa26 was first identified in the Oryza sativa L. ssp. indica rice variety Minghui 63; it was mapped to the long arm of chromosome 11. The function of bacterial-blight resistance gene Xa3/Xa26 in rice is influenced by genetic background; Oryza sativa L. ssp. japonica background can increase Xa3/Xa26 expression, resulting in enhanced resistance. To identify whether Xa3/Xa26 transcript level is the only factor contributing to genetic background-controlled resistance, we combined the analyses of quantitative genetics and gene expression profile in this study.
First, some juvenile and adult plants were randomly chosen from a Mudanjiang 8/Minghui 63 F2 population to analyze the relationship between the transcription level of Xa3/Xa26 and resistance level to bacterial blight. Increasing Xa3/Xa26 expression was correlated with the enhanced resistance in theses F2 plants both at juvenile and adult stages. But a few F2 plants showed a disassociation of Xa3/Xa26 expression level and enhanced resistance. These results suggest that genetic background does influence Xa3/Xa26 expression and its expression level is not an exclusive factor which affects its function.
Second, five quantitative trait loci (QTLs) were identify from a Mudanjiang 8/Minghui 63 F2 population and some backcross populations using the Minghui 63 as a recurrent parent. The resistance alleles at these five QTLs were from the susceptible parent Mudanjiang 8. It means that some loci contributed by susceptible parent can influence Xa3/Xa26-mediated Xoo resistance. The other four QTLs also can enhance resistance when these alleles were homozygous for Minghui 63. Furthermore, One hundred and twenty-nine co-dominant markers were used for testing digenic interactions in this F2 segregating population. Few epistatic effects were existed between these QTLs. Most digenic interactions were detected between non-significant effect loci (non-QTL). These results suggest that Xa3/Xa26-mediated resistance was affected by multiple loci, including QTLs and epistatic interactions.
Third, analysis difference of the expression profile of Xa3/Xa26-mediated resistance between Minghui 63 and Rb49, using a cDNA microarray. Rb49 is a transgenic line carrying a single copy of Xa3/Xa26 regulated by its native promoter in the genetic background of susceptible Mudanjiang 8. Most differentially expressed genes were rapidly activated in the Rb49, but not in the indica background, during disease resistance. Os03g60080, encoding a SNAC1 protein, was suppressed by pathogen in both Minghui 63 and Rb49, while its expression pattern of four time point after inoculation was opposite to that of Xa3/Xa26,OsWRKY13 and NH1 This gene was also suppressed in OsWRKY13-overexpressing lines. It was indicated that the differentially expressed gene not only related with abiotic stress, but also participated in the signal transduction of Xa3/Xa26-mediated resistance. It means that the different spatiotemporal expression pattern of some genes was another important factor affecting the R gene function.
Fourth, the colocalization of some differentially expressed genes with QTLs in Xa3/Xa26-mediated resistance will help to characterize the genes underlying these QTLs and involved in regulation of genetic background-controlled resistance.
引文
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