大豆胞囊线虫不同生理小种的致病性及其分子基础研究
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摘要
本论文从大豆胞囊线虫不同生理小种的致病性差异入手,系统地研究了大豆胞囊线虫不同生理小种对大豆膜脂过氧化作用、光合能力、物质代谢能力、防御酶系统等多个方面与致病性差异的关系；本文首次利用蛋白质组和SSH的方法从SCN不同生理小种本身寻找差异蛋白和差异序列,从分子水平揭示了SCN不同生理小种的致病性差异,主要研究结果如下：
1.将来源不同的4份胞囊土样重新进行生理分化表型的研究。生理小种的鉴定结果为：采自辽宁省康平县的S1为1号生理小种；辽宁省沈阳市的S2为3号生理小种；山西省汾阳市的S3为4号生理小种；黑龙江省安达市S4为14号生理小种；均与土样来源地的原生理小种相同。因此可以确定这4份土样中大豆胞囊线虫的生理小种。HG类型的鉴定结果为：S1的HG类型为2,5,7、S2的HG类型为5,7、S3的HG类型为1,2,3,5,7、S4的HG类型为1,3,7。
2.大豆胞囊线虫不同生理小种对大豆生理指标的影响与其致病性密切相关。试验采用辽豆10、PI88788、PI90763,3个品种均同时接种大豆胞囊线虫的4个生理小种：1号、3号、4号和14号。测定了这3个品种组织的丙二醛含量、电解质渗透率以及叶绿素含量3项生理指标,分析不同品种的大豆接种大豆胞囊线虫不同生理小种后生理变化和致病性关系。结果表明：PI88788接种大豆胞囊线虫1号、4号生理小种和PI90763接种4号、14号生理小种后3种生理指标均显著高于本品种的对照水平；而PI88788接种大豆胞囊线虫3号和14号生理小种后3种生理指标略高于对照水平,PI90763接种大豆胞囊线虫1号和3号生理小种后3种生理指标略高于对照水平。研究证实大豆胞囊线虫不同生理小种对大豆不同品种的致病性与接种SCN不同生理小种后大豆植株内丙二醛含量、电解质渗透率、叶绿素含量的变化密切相关。
3.物质代谢与大豆胞囊线虫致病性有着密切关系。接种大豆胞囊线虫后,供试大豆品种根内可溶性糖含量有不同程度的提高,接种同一个SCN生理小种相对感病的品种根内可溶性糖含量低于抗病品种,认为寄主植物可能需要适当的糖类物质作为抗病反应的激发子来抵御线虫的侵染,根系中低糖有利于大豆胞囊线虫的侵入；PI88788接种大豆胞囊线虫3号、14号生理小种和PI90763接种1号、3号生理小种后根内可溶性蛋白含量变化较大,升高明显,这一现象与寄主植物大豆受侵染后诱导产生与抗性相关的蛋白有一定关系。
4.受大豆胞囊线虫不同生理小种侵染后大豆根内防御相关酶系,包括苯丙氨酸解氨酶(PAL)、多酚氧化酶(PPO)、超氧化物歧化酶(SOD)、过氧化物酶(POD)和几丁质酶活性出现了明显的变化,表现出酶活的动态变化与大豆胞囊线虫不同生理小种的致病性密切相关。
5.大豆胞囊线虫的4个生理小种经双向电泳各检测到近500个蛋白点(其中3号和4号生理小种为一个对照组；1号和14号生理小种为一个对照组),从2-DE图谱上找到30个明显差异表达的蛋白点,通过基质辅助激光解析电离飞行时间质谱技术(MALDI-TOF/TOF)获得了这30个蛋白点的肽质量指纹图谱(PMF), Mascot数据库搜索比对后,鉴定了17个蛋白质点,它们分别是nebulin-related-anchoring protein isoform 1(80); myosin heavy chain(240); CRE-UGT-56 protein(119); cornified envelope precursor protein(136); similar to Rho-associated kinase(352); GG18879(206a); GF17954(206b); golgi antigen gcp372(253,389); Rp49(405); predicted protein(274); hypothetical protein(114, 127,128,209,296)。
6.利用抑制消减杂交技术构建了大豆胞囊线虫不同生理小种J2 cDNA的正反向抑制消减文库(其中3号和4号生理小种为一个对照组；1号和14号生理小种为一个对照组)。结果表明：应用T/A克隆法,共获得2000多个正、反向消减片段的重组克隆,得到的cDNA片段的长度分布在200～900bp左右,平均长度约为500bp左右。以上结果表明：通过SSH技术所构建的cDNA文库能够满足后续试验的要求。
7.对所得到的2对正反向消减文库进行斑点杂交,分别从分库中挑选插入片段在500bp左右的质粒768个,与DIG标记的SSH-探针分别进行杂交,筛选到972个候选阳性克隆。为进一步确定差异表达基因,我们从斑点杂交的阳性克隆中挑选了20个差异基因进行RT-PCR验证,结果表明：在挑选的这20个基因中得到10个真正差异表达的cDNA片段。其中LFSSH-875、LFSSH-950、LFSSH-963、LFSSH-1000、LFSSH-1103只出现在4号生理小种,不出现在3号生理小种中,即在4号生理小种中高表达；而LRSSH-1209只出现在3号生理小种中,不出现在4号生理小种中,即在3号生理小种中高表达；XFSSH-938只出现在14号生理小种中,不出现在1号生理小种中,即在14号生理小种中高表达；而XRSSH-1018、XRSSH-1250、XRSSH-1258只出现在1号生理小种中,不出现在14号生理小种中,即在1号生理小种中高表达。其余的基因只是在表达量上有区别。这说明,致病性较弱的生理小种中也会有特异性表达的基因。
The researches on the pathogenic differences of different races of Heterodera glycines were systematically conducted to the relationship between pathogenic differences and the soybean's membrane lipid peroxidation, photosynthetic capacity, material metabolism, oxidases system. Proteomics and SSH were used in research on different races of SCN to study different proteomics and different genes. It was important for fundamentally revelation pathogenic differences of different races of Heterodera glycines. The main results of this study were as follows:
1. Four Heterodera glycines samples from different areas were collected, race tests were conducted on this population. The sample collected from Kangping is race 1; the sample S2 collected from Shenyang is race 3; the sample collected from Hefei is race 4; the sample collected from Anda is race 14. Employ by HG type appraisal results as follows:The sample collected from Kangping is 2,5,7; the sample S2 collected from Shenyang is 5,7; the sample collected from Hefei is 1,2,3,5,7; the sample collected from Anda is 1,3,7.
2. Three soybean varieties'Liaodou 10','PI88788'and'PI90763'were inoculated with different races of Heterodera glycines (race 1, race 3, race 4 and race 14) in the same time. The three physiological indices of MDA contents, Electronic efflux percentage and Chlorophyll contents were analyzed. Results showed that the three physiological indices of soybean cultivars'PI88788'colonized by H.glycines race 1、race 4and 'PI90763' colonized by H.glycines race 4、race 14 were both higher than those in control soybean cultivars. But this three physiological indices of soybean cultivars'PI88788'colonized by H.glycines race 3 and race 14 were a little higher than those in control roots; three physiological indices of soybean cultivars'PI90763'colonized by H.glycines race 1 and race 3 were a little higher than those in control Soybean cultivars. It is concluded that the activities of this three physiological indices had closely related to pathogenicity of different races of H.glycines.
3. The study on the relationships between the metabolism and resistance to H.glycines indicated that the soluble sugar content of resistant varieties root was increased in different degree after inoculating by the same race of SCN, the soluble sugar content of susceptible varieties root was relatively lower than CK. Host plants need carbohydrate to resist SCN. Low sugar content in soybean roots was beneficial to nematode invasion. Moreover, the soluble protein content in the roots of soybean cultivars'PI88788'colonized by H.glycines race 3 and race 14 and soybean cultivars'PI90763'colonized by H.glycines race 1 and race 3 were obviously increased, maybe soybean infected by SCN were induced to express resistant protein
4. Defensive enzymes were stimulated after infected by different races of SCN. The dynamic changes of defensive enzymes, include phenylalanine ammonialyase (PAL), polyphenoloxidase (PPO), superoxide dismutase (SOD) and peroxidase (POD) were studied. The results showed that the change of enzymes had closely related to pathogenicity of different races of H.glycines.
5. Differential proteomics were studied of different races of SCN(race 3 and race 4 as a control group; race 1 and race 14 as a control group). Two-dimensional gel electrophoresis (2-DE) were adopted to identify proteins from different races of SCN. Nearly 500 protein spots were detected. Furthermore, analysis with MALDI-TOF-MASS identification Peptide Mass Fingerprinting of 30 protein spots. The 17 protein spots were obtained by Mascot database searching and intercomparison preliminary identification Mascot, The proteins were respectively nebulin-related-anchoring protein isoform 1(80,114); myosin heavy chain(240); CRE- UGT- 56 protein(119); cornified envelope precursor protein(136); similar to Rho-associated kinase (352); GG18879(206a); GF17954(206b); golgi antigen gcp372(253, 389); Rp49(405); predicted protein(274); hypothetical protein(127,128,209,296).
6. Forward and reverse suppressive subtractive hybridization (SSH) libraries were constructed by using RNA prepared from different races of H.glycines(race 3 and race 4 as a control group; race 1 and race 14 as a control group). Had more than 2000 clones. The both subtraction libraries with the inserted fragments between 200bp and 900bp. All these results indicated that both SSH libraries were qualified for further studies.
7. Plasmids with 500bp inserted fragment were selected from each of the two forward and reverse suppressive subtractive hybridization libraries for dot blotting hybridization in which DIG was used to label probes from SSH PCR products, respectively. A total of 972 genes were found. To confirm further the differential expressions of the genes,20 genes were chosen for RT-PCR analysis. The results indicated that:LFSSH-875, LFSSH-950, LFSSH-963, LFSSH-1000, LFSSH-1103 were expressed mainly in race 4; LRSSH-1209 was expressed mainly in race 3; XFSSH-938 was expressed mainly in race 14; XRSSH-1018、 XRSSH-1250、XRSSH-1258 were expressed mainly in race 1.
1.将来源不同的4份胞囊土样重新进行生理分化表型的研究。生理小种的鉴定结果为：采自辽宁省康平县的S1为1号生理小种；辽宁省沈阳市的S2为3号生理小种；山西省汾阳市的S3为4号生理小种；黑龙江省安达市S4为14号生理小种；均与土样来源地的原生理小种相同。因此可以确定这4份土样中大豆胞囊线虫的生理小种。HG类型的鉴定结果为：S1的HG类型为2,5,7、S2的HG类型为5,7、S3的HG类型为1,2,3,5,7、S4的HG类型为1,3,7。
2.大豆胞囊线虫不同生理小种对大豆生理指标的影响与其致病性密切相关。试验采用辽豆10、PI88788、PI90763,3个品种均同时接种大豆胞囊线虫的4个生理小种：1号、3号、4号和14号。测定了这3个品种组织的丙二醛含量、电解质渗透率以及叶绿素含量3项生理指标,分析不同品种的大豆接种大豆胞囊线虫不同生理小种后生理变化和致病性关系。结果表明：PI88788接种大豆胞囊线虫1号、4号生理小种和PI90763接种4号、14号生理小种后3种生理指标均显著高于本品种的对照水平；而PI88788接种大豆胞囊线虫3号和14号生理小种后3种生理指标略高于对照水平,PI90763接种大豆胞囊线虫1号和3号生理小种后3种生理指标略高于对照水平。研究证实大豆胞囊线虫不同生理小种对大豆不同品种的致病性与接种SCN不同生理小种后大豆植株内丙二醛含量、电解质渗透率、叶绿素含量的变化密切相关。
3.物质代谢与大豆胞囊线虫致病性有着密切关系。接种大豆胞囊线虫后,供试大豆品种根内可溶性糖含量有不同程度的提高,接种同一个SCN生理小种相对感病的品种根内可溶性糖含量低于抗病品种,认为寄主植物可能需要适当的糖类物质作为抗病反应的激发子来抵御线虫的侵染,根系中低糖有利于大豆胞囊线虫的侵入；PI88788接种大豆胞囊线虫3号、14号生理小种和PI90763接种1号、3号生理小种后根内可溶性蛋白含量变化较大,升高明显,这一现象与寄主植物大豆受侵染后诱导产生与抗性相关的蛋白有一定关系。
4.受大豆胞囊线虫不同生理小种侵染后大豆根内防御相关酶系,包括苯丙氨酸解氨酶(PAL)、多酚氧化酶(PPO)、超氧化物歧化酶(SOD)、过氧化物酶(POD)和几丁质酶活性出现了明显的变化,表现出酶活的动态变化与大豆胞囊线虫不同生理小种的致病性密切相关。
5.大豆胞囊线虫的4个生理小种经双向电泳各检测到近500个蛋白点(其中3号和4号生理小种为一个对照组；1号和14号生理小种为一个对照组),从2-DE图谱上找到30个明显差异表达的蛋白点,通过基质辅助激光解析电离飞行时间质谱技术(MALDI-TOF/TOF)获得了这30个蛋白点的肽质量指纹图谱(PMF), Mascot数据库搜索比对后,鉴定了17个蛋白质点,它们分别是nebulin-related-anchoring protein isoform 1(80); myosin heavy chain(240); CRE-UGT-56 protein(119); cornified envelope precursor protein(136); similar to Rho-associated kinase(352); GG18879(206a); GF17954(206b); golgi antigen gcp372(253,389); Rp49(405); predicted protein(274); hypothetical protein(114, 127,128,209,296)。
6.利用抑制消减杂交技术构建了大豆胞囊线虫不同生理小种J2 cDNA的正反向抑制消减文库(其中3号和4号生理小种为一个对照组；1号和14号生理小种为一个对照组)。结果表明：应用T/A克隆法,共获得2000多个正、反向消减片段的重组克隆,得到的cDNA片段的长度分布在200～900bp左右,平均长度约为500bp左右。以上结果表明：通过SSH技术所构建的cDNA文库能够满足后续试验的要求。
7.对所得到的2对正反向消减文库进行斑点杂交,分别从分库中挑选插入片段在500bp左右的质粒768个,与DIG标记的SSH-探针分别进行杂交,筛选到972个候选阳性克隆。为进一步确定差异表达基因,我们从斑点杂交的阳性克隆中挑选了20个差异基因进行RT-PCR验证,结果表明：在挑选的这20个基因中得到10个真正差异表达的cDNA片段。其中LFSSH-875、LFSSH-950、LFSSH-963、LFSSH-1000、LFSSH-1103只出现在4号生理小种,不出现在3号生理小种中,即在4号生理小种中高表达；而LRSSH-1209只出现在3号生理小种中,不出现在4号生理小种中,即在3号生理小种中高表达；XFSSH-938只出现在14号生理小种中,不出现在1号生理小种中,即在14号生理小种中高表达；而XRSSH-1018、XRSSH-1250、XRSSH-1258只出现在1号生理小种中,不出现在14号生理小种中,即在1号生理小种中高表达。其余的基因只是在表达量上有区别。这说明,致病性较弱的生理小种中也会有特异性表达的基因。
The researches on the pathogenic differences of different races of Heterodera glycines were systematically conducted to the relationship between pathogenic differences and the soybean's membrane lipid peroxidation, photosynthetic capacity, material metabolism, oxidases system. Proteomics and SSH were used in research on different races of SCN to study different proteomics and different genes. It was important for fundamentally revelation pathogenic differences of different races of Heterodera glycines. The main results of this study were as follows:
1. Four Heterodera glycines samples from different areas were collected, race tests were conducted on this population. The sample collected from Kangping is race 1; the sample S2 collected from Shenyang is race 3; the sample collected from Hefei is race 4; the sample collected from Anda is race 14. Employ by HG type appraisal results as follows:The sample collected from Kangping is 2,5,7; the sample S2 collected from Shenyang is 5,7; the sample collected from Hefei is 1,2,3,5,7; the sample collected from Anda is 1,3,7.
2. Three soybean varieties'Liaodou 10','PI88788'and'PI90763'were inoculated with different races of Heterodera glycines (race 1, race 3, race 4 and race 14) in the same time. The three physiological indices of MDA contents, Electronic efflux percentage and Chlorophyll contents were analyzed. Results showed that the three physiological indices of soybean cultivars'PI88788'colonized by H.glycines race 1、race 4and 'PI90763' colonized by H.glycines race 4、race 14 were both higher than those in control soybean cultivars. But this three physiological indices of soybean cultivars'PI88788'colonized by H.glycines race 3 and race 14 were a little higher than those in control roots; three physiological indices of soybean cultivars'PI90763'colonized by H.glycines race 1 and race 3 were a little higher than those in control Soybean cultivars. It is concluded that the activities of this three physiological indices had closely related to pathogenicity of different races of H.glycines.
3. The study on the relationships between the metabolism and resistance to H.glycines indicated that the soluble sugar content of resistant varieties root was increased in different degree after inoculating by the same race of SCN, the soluble sugar content of susceptible varieties root was relatively lower than CK. Host plants need carbohydrate to resist SCN. Low sugar content in soybean roots was beneficial to nematode invasion. Moreover, the soluble protein content in the roots of soybean cultivars'PI88788'colonized by H.glycines race 3 and race 14 and soybean cultivars'PI90763'colonized by H.glycines race 1 and race 3 were obviously increased, maybe soybean infected by SCN were induced to express resistant protein
4. Defensive enzymes were stimulated after infected by different races of SCN. The dynamic changes of defensive enzymes, include phenylalanine ammonialyase (PAL), polyphenoloxidase (PPO), superoxide dismutase (SOD) and peroxidase (POD) were studied. The results showed that the change of enzymes had closely related to pathogenicity of different races of H.glycines.
5. Differential proteomics were studied of different races of SCN(race 3 and race 4 as a control group; race 1 and race 14 as a control group). Two-dimensional gel electrophoresis (2-DE) were adopted to identify proteins from different races of SCN. Nearly 500 protein spots were detected. Furthermore, analysis with MALDI-TOF-MASS identification Peptide Mass Fingerprinting of 30 protein spots. The 17 protein spots were obtained by Mascot database searching and intercomparison preliminary identification Mascot, The proteins were respectively nebulin-related-anchoring protein isoform 1(80,114); myosin heavy chain(240); CRE- UGT- 56 protein(119); cornified envelope precursor protein(136); similar to Rho-associated kinase (352); GG18879(206a); GF17954(206b); golgi antigen gcp372(253, 389); Rp49(405); predicted protein(274); hypothetical protein(127,128,209,296).
6. Forward and reverse suppressive subtractive hybridization (SSH) libraries were constructed by using RNA prepared from different races of H.glycines(race 3 and race 4 as a control group; race 1 and race 14 as a control group). Had more than 2000 clones. The both subtraction libraries with the inserted fragments between 200bp and 900bp. All these results indicated that both SSH libraries were qualified for further studies.
7. Plasmids with 500bp inserted fragment were selected from each of the two forward and reverse suppressive subtractive hybridization libraries for dot blotting hybridization in which DIG was used to label probes from SSH PCR products, respectively. A total of 972 genes were found. To confirm further the differential expressions of the genes,20 genes were chosen for RT-PCR analysis. The results indicated that:LFSSH-875, LFSSH-950, LFSSH-963, LFSSH-1000, LFSSH-1103 were expressed mainly in race 4; LRSSH-1209 was expressed mainly in race 3; XFSSH-938 was expressed mainly in race 14; XRSSH-1018、 XRSSH-1250、XRSSH-1258 were expressed mainly in race 1.
引文
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