稻瘟病菌（Magnaporthe grisea）致病相关基因的分子标记及克隆
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摘要
利用近年来发展起来的真菌限制性内切酶介导整合(REMI)插入诱变技术，用含有HygB抗性标记的质粒pUCATPH在稻瘟病菌菌株M131中建立一个转化体系，结果发现用不同的限制酶介导转化，原生质体的转化效率有不同程度的提高，经过多次转化得到639个转化子，对其中的200个转化子进行致病性测定，发现其中6个转化子对一些品种的致病性和原始菌株M131对比发生了变化，即为致病性突变体，同时还观察到部分表型发生变化的表型突变体。
用DIG辛标记质粒pUCATPH作为探针对6个致病性突变菌株和两个表型突变菌株进行点杂交检测，结果都呈杂交阳性，说明质粒的插入使原始菌株M131的某个致病相关基因或黑色素产生相关基因发生了突变。对这些突变菌株进行rep-PCR分析，发现变菌株的rep-PCR图谱不仅和原始菌株M131相比有一定差异而且相互间也有区别，可以认为这些菌株在分子水平上发生了突变，并且在REMI诱变中质粒插入基因组的位点不同。将其中两个表型突变菌株及rep-PCR图谱差异性较大的两个致病性突变菌株和M131的DNA分别用Hind Ⅲ和EcoR Ⅴ酶切电泳后，以DIG辛标记的质粒为探针进行RFLP分析，结果发现突变菌株中不仅在质粒插入位点有杂交信号，而且在分子量较小处也有强弱不一的信号出现。这些现象可能是质粒多位点串联整合并且整合后发生重组的结果。
突变菌株01-36对已知含有抗病基因Pita的K1品种致病性发现了变化，根据基因-基因学说，可以推测菌株中无毒基因avr-Pita可能发生了突变。依据已经发表的无毒基因avr-Pita序列设计出引物P1、P2，分别对M131和突变菌株01-36基因组DNA进行PCR扩增，结果都出现一条1100bp左右的扩增带。将M131的扩增片段测序后得到长度为1057bp的序列，并且将所测序列和已知的avr-Pita一段序列比较发现达98％的同源，即该序列为无毒基因的某一段序列，但插入的位点并不在所扩增片段内。另外选取一些来自不同地区的致病性差异较大的稻瘟病菌菌株进行PCR扩增，以M131扩增产物为探针，对各扩增产物进行点杂交分析，结果都为阳性反应，即所有扩增片段都可能和无毒基因avr-Pita有部分同源性。本文通过无毒基因的克隆和分析，对基因-基因学说中病原菌无毒基因和抗病基因互作方式进行了初步探讨。
To perform insertional mutation to tag pathogenicity related gene in Magnaporthe grisea, the pathogen strain Ml31 was used to establish a high-efficient Restriction Enzyme Mediated Integration(REMI) transformation system with the plasmid pUCATPH. The plasmid pUCATPH was linearized by restriction endonucleases Hind III, Sac I, and Kpn I, respectively. Resulted fragments were introduced individually into Ml31 in the presence of corresponding enzymes. Transformation ration was found to be increased by the incorporation of restriction enzymes, while levels of transformation enhancement varied with enzymes.
Based on pathogenicity and pigment production of 200 transfomants, two transformants with altered phenotypes and six transformants with modified pathogenicity were obtained. DNA dot hybridization analysis revealed that the plasmid DNA was integrated into the fungal genomes in these transformants. Moreover, genes involved in pathogenicity or pigment production experimentally were mutated in these mutants. Rep-PCR fingerprint patterns of the trasformants and M131 revealed that mutants were different from M131, and from one another, suggesting that they have mutated at molecular levels. To determine patterns of the plasmid integration, DNA from four transformants were digested with Hind III and EcoR V ,respectively, and tested by dot hybridization using pUCATPH as the probe. It is found that M131 was not homologous to pUCATPH, and the four transformants all showed many bands of varied sizes. These patterns could result from REMI events, tandem integrations, or a combination of both.
The transformant 01-36 was nonpathogenic to the rice varity K1 with resistance gene Pita, which was different from M131. It could be deduced that avirulence gene avr-Pita might had mutated in the transformant 01-36. The genome DNA of the strains 01-36 and M131 was PCR amplified using a couple of primers PI and P2 designed according to the sequence of avirulence gene avr-Pita, and a 1057bp fragment was obtained. The amplified
band from M131 was sequenced and then aligned with the sequence of avr-Pila gene. The result was that the sequence was 98% homologous to one fragment of avr-Pita sequence, but the integration site was not within this fragment. The other amplified fragments of the strains those are incompatible to Kl and from different regions, was subjected to dot hybridization analysis with the probe of tagged avr-Pita. Positive responses showed that they all could included homologous fragments to avr-Pita gene. The interaction fashions between avirulence genes and resistant genes were pilot studied.
用DIG辛标记质粒pUCATPH作为探针对6个致病性突变菌株和两个表型突变菌株进行点杂交检测，结果都呈杂交阳性，说明质粒的插入使原始菌株M131的某个致病相关基因或黑色素产生相关基因发生了突变。对这些突变菌株进行rep-PCR分析，发现变菌株的rep-PCR图谱不仅和原始菌株M131相比有一定差异而且相互间也有区别，可以认为这些菌株在分子水平上发生了突变，并且在REMI诱变中质粒插入基因组的位点不同。将其中两个表型突变菌株及rep-PCR图谱差异性较大的两个致病性突变菌株和M131的DNA分别用Hind Ⅲ和EcoR Ⅴ酶切电泳后，以DIG辛标记的质粒为探针进行RFLP分析，结果发现突变菌株中不仅在质粒插入位点有杂交信号，而且在分子量较小处也有强弱不一的信号出现。这些现象可能是质粒多位点串联整合并且整合后发生重组的结果。
突变菌株01-36对已知含有抗病基因Pita的K1品种致病性发现了变化，根据基因-基因学说，可以推测菌株中无毒基因avr-Pita可能发生了突变。依据已经发表的无毒基因avr-Pita序列设计出引物P1、P2，分别对M131和突变菌株01-36基因组DNA进行PCR扩增，结果都出现一条1100bp左右的扩增带。将M131的扩增片段测序后得到长度为1057bp的序列，并且将所测序列和已知的avr-Pita一段序列比较发现达98％的同源，即该序列为无毒基因的某一段序列，但插入的位点并不在所扩增片段内。另外选取一些来自不同地区的致病性差异较大的稻瘟病菌菌株进行PCR扩增，以M131扩增产物为探针，对各扩增产物进行点杂交分析，结果都为阳性反应，即所有扩增片段都可能和无毒基因avr-Pita有部分同源性。本文通过无毒基因的克隆和分析，对基因-基因学说中病原菌无毒基因和抗病基因互作方式进行了初步探讨。
To perform insertional mutation to tag pathogenicity related gene in Magnaporthe grisea, the pathogen strain Ml31 was used to establish a high-efficient Restriction Enzyme Mediated Integration(REMI) transformation system with the plasmid pUCATPH. The plasmid pUCATPH was linearized by restriction endonucleases Hind III, Sac I, and Kpn I, respectively. Resulted fragments were introduced individually into Ml31 in the presence of corresponding enzymes. Transformation ration was found to be increased by the incorporation of restriction enzymes, while levels of transformation enhancement varied with enzymes.
Based on pathogenicity and pigment production of 200 transfomants, two transformants with altered phenotypes and six transformants with modified pathogenicity were obtained. DNA dot hybridization analysis revealed that the plasmid DNA was integrated into the fungal genomes in these transformants. Moreover, genes involved in pathogenicity or pigment production experimentally were mutated in these mutants. Rep-PCR fingerprint patterns of the trasformants and M131 revealed that mutants were different from M131, and from one another, suggesting that they have mutated at molecular levels. To determine patterns of the plasmid integration, DNA from four transformants were digested with Hind III and EcoR V ,respectively, and tested by dot hybridization using pUCATPH as the probe. It is found that M131 was not homologous to pUCATPH, and the four transformants all showed many bands of varied sizes. These patterns could result from REMI events, tandem integrations, or a combination of both.
The transformant 01-36 was nonpathogenic to the rice varity K1 with resistance gene Pita, which was different from M131. It could be deduced that avirulence gene avr-Pita might had mutated in the transformant 01-36. The genome DNA of the strains 01-36 and M131 was PCR amplified using a couple of primers PI and P2 designed according to the sequence of avirulence gene avr-Pita, and a 1057bp fragment was obtained. The amplified
band from M131 was sequenced and then aligned with the sequence of avr-Pila gene. The result was that the sequence was 98% homologous to one fragment of avr-Pita sequence, but the integration site was not within this fragment. The other amplified fragments of the strains those are incompatible to Kl and from different regions, was subjected to dot hybridization analysis with the probe of tagged avr-Pita. Positive responses showed that they all could included homologous fragments to avr-Pita gene. The interaction fashions between avirulence genes and resistant genes were pilot studied.
引文
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