水稻纹枯病接种鉴定体系新探索及抗病新种质YSBR1抗性分析
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摘要
纹枯病是世界性的水稻3大病害之一,严重发生时可造成产量的巨大损失和品质的严重降低。水稻对纹枯病的抗性属于典型的数量性状,不存在主基因抗性,现有种质资源中也未发现表现免疫或高抗的品种。
水稻纹枯病接种鉴定体系涉及到接种方法、接种时间、调查时间、调查标准等多种因素,其合适与否关系到水稻抗纹枯病遗传育种研究的进展。合适的接种鉴定体系不仅可减轻相关研究工作的工作强度,而且能提高试验的可靠性、重复性及结果的精确性。鉴此,本试验以抗感反应不同的5个水稻品种[Lemont、武育粳3号(Wuyujing3)、YSBR1、特青(Teqing)和Jasmine85(J-85)]为试验材料,在人工气候箱、控温室中进行水稻苗期抗纹枯病接种试验,并与田间相应的成株期抗性试验进行比较,初步建立了水稻苗期接种鉴定体系。在此基础上,采用牙签嵌入法在控温室、田间对各试验水稻材料进行成株期不同阶段(分蘖末期、小孕穗期)抗纹枯病接种试验,并于接种后3-11d、抽穗后30d测量病斑长度和采用3种不同调查标准调查病级,对水稻成株期纹枯病的接种鉴定方法进行了发展,并对田间水稻抽穗后30d适宜的纹枯病病级调查标准进行了新探索。
YSBR1是本研究组选育并经过多年鉴定为较抗病的新种质,目前对其抗性研究报道较少。本试验以YSBR1及易感品种Lemont为试验材料,通过构建这2个品种的F2无性系群体,对其开展了多年份的纹枯病抗性鉴定及农艺性状调查,同时对YSBR1中相关纹枯病抗性QTL、农艺性状QTL进行初步定位,研究这2类QTL之间的相互关系。在此基础上,又以这2个品种为试验材料,对其进行纹枯病病原菌接种,分别于接种后10、20h取样用于基因芯片表达谱分析,从中筛选差异表达基因,尤其是YSBR1特异差异表达基因。最终对YSBR1特异差异表达基因进行pathway分析和GO (gene ontology)分析,并将其与已检测到YSBR1的QTL区间进行整合。
主要研究结果如下：
1.85%的相对湿度为纹枯病菌侵染危害水稻苗期植株的适宜湿度；苗期5个水稻品种对纹枯病抗性差异极显著,可将其分为相对感病(Lemont、武育粳3号)和相对抗病(YSBR1、Jasmine85、特青)2大类；接种叶龄对发病程度有显著的影响,5个品种在4叶期接种时的平均病级显著高于5叶期接种的平均病级；苗期水稻品种间抗感差异小于田间鉴定试验结果,但两者间品种抗感趋势基本一致。表明,苗期快速鉴定技术可用于大规模水稻品种(组合)的抗性筛选或初步鉴定。
2.以病斑长度为调查指标,控温室中分蘖末期接种的5个水稻品种间抗感差异极显著,Lemont和YSBR1表现为稳定的感病、抗病,且和另外3个品种(武育粳3号、特青、Jasmine85)差异极显著,各品种间的抗感趋势与田间抽穗后30d的验证试验结果基本一致；而田间分蘖末期接种的5个水稻品种间虽然抗感差异极显著,但品种间抗感趋势与同期温室及田间抽穗后30d的验证试验结果差异不一致。田间小孕穗期接种的5个水稻品种间抗感差异极显著,品种间抗感趋势与分蘖末期接种并于抽穗后30d调查病级的田间验证试验结果完全一致。表明,温室条件下分蘖末期或田间条件下小孕穗期以病斑长度作为衡量纹枯病抗性的调查指标具有一定的可行性。
3.在分蘖末期接种的各参试品种,采用叶鞘位调查标准的结果与常用的Rush调查标准的结果相比较,两者在不同年份间、地点间差异均不显著,相关分析结果显示,2种调查标准间相关性在不同年份、地点间分别为0.9455、0.9846,均达到极显著水平,表明叶鞘位标准作为1个新的调查标准在田间纹枯病病级调查时完全可替代Rush调查标准；对于小孕穗期接种的各品种,采用相对病级调查标准,各品种间不仅抗性差异极显著,而且品种间抗感差异大于分蘖末期接种采用叶鞘位调查标准的相应对照,并且这2种调查标准间各品种的抗、感趋势完全一致。在水稻抗纹枯病遗传研究中,小孕穗期接种以相对病级作为调查标准,其适用范围是有限制的。
4.两年间(2009-2010年)共初步定位到12个抗性QTL,除qSB-1Le、qSB-5Le和qSB-8Le来自感病亲本Lemont外,其余均来自抗病亲本YSBR1,其中qSB-2Y、qSB-12Y被认为是主效QTL,两年的平均贡献率分别为28%和17%。此外,两年间13个主要农艺性状中有4个农艺性状(株高、生育期、穗长、有效穗数)和纹枯病发病程度存在显著或极显著相关,且多为负相关。表明水稻纹枯病的发病情况受到株高、生育期、穗长、有效穗数等性状的影响,尤其是株高的影响最大。
5.接种后短期内(5d),YSBR1与Lemont间抗感差异明显,接种后10h均产生侵染垫,接种后20h均出现病斑；不同生物学重复芯片质量可靠,按自定筛选标准,在Lemont全基因组共筛选到2826个差异表达基因,在YSBR1全基因组筛选到748个差异表达基因,其中在YSBR1中特异表达的差异基因有247个；247个YSBR1中特异表达基因分布水稻12条染色体上,落入已检测到的QTL区间的为31个,占总数比例为12.55%;落入2个主效QTLqSB-2Y、qSB-12Y区间的YSBRl特异表达基因分别有9、3个,其中qSB-2Y区间9个特异差异表达基因中有4个上调表达,5个下调表达；qSB-12Y区间3个特异差异表达基因中有2个上调表达,1个下调表达。
Sheath blight (SB) disease is one of the three most serious diseases in rice worldwide, and causes severe loss in yield and quality under favorable environmental condition. The resistance of rice to SB is a typical quantitative trait, and resources with strong resistance against this disease in the existing rice germplasm are not found.
Rice SB inoculation identification system involves the inoculation method, inoculation time, investigation time, investigation standard and other factors, and the appropriateness of the system can affect genetic and breeding research progress of the resistance to SB in rice. Appropriate inoculation identification system can not only reduce the strength of the research work, but also improve the reliability, repeatability, and accuracy of the test results. In view of this point, five rice cultivars with different levels of resistance to SB were adopted to identify their resistance to SB at seedling stage in artificial climate incubator or temperature-controlled greenhouse, and the test results were verified through a field experiment during rice adult stage, then a SB inoculation identification system in rice seedling stage was established preliminarily. Based on the above study, five rice cultivars were inoculated with toothpick-inserting inoculation method during different times in rice adult stage (the late tillering stage, the early booting stage) in field and temperature-controlled greenhouse. The lesion length was measured during3-11days after inoculation, and the SB disease rating at the30th day after heading was investigated with the reference of three different investigation standards. Then, some new inoculation identification methods in rice adult stage were developed and some new investigation standards of SB disease rating at the30th day after heading stage in field were also explored in this study.
YSBR1developed by our own research group is a new germplasm with stable and high resistance to sheath blight, which has been identified by using many methods in different experimental sites for many years. However, information of its genetic background relevant to SB resistance has been in scarce yet. A F2clonal populations (F2CP) derived from the cross between YSBR1and Lemont (a susceptible cultivar) was constructed, and the identification of resistance to SB and the investigation of agronomic traits for F2CP in a multi-year experiment were carried out in this study. At the same time, the QTLs mapping of resistance to SB in YSBR1and agronomic traits were conducted, and the interrelation between resistance to SB and agronomic traits were also analyzed. On the basis of this study, the two cultivars (YSBRl, Lemont) were used as materials to inoculate SB pathogens, and their leaf sheaths were sampled at10and20hours after inoculation respectively for gene chip expression profiling analysis and screening of differentially expressed genes, especially YSBR1specific differentially expressed gene. Furthermore, the pathway and GO (gene ontology) of YSBR1specific differentially expressed gene and its integration with QTL detected in YSBRl were analyzed.
The main results were as follows:
1. The optimal relative humidity for SB mycelium to develop infection structures and to infect rice seedling was approximately85%. The SB resistance levels in5cultivar seedlings differed significantly (P<0.01) and could be divided into two groups, relatively susceptible (Lemont, WuYujing3) and relatively resistant (YSBR1, Jasmine85, Teqing). The symptom severeness degrees of the seedlings at different leaf-age (4-leaf stage and5-leaf stage) were significantly different and the average disease rating at4-leaf stage was significantly higher than that at5-leaf stage. The differences of resistance among different cultivars in greenhouse were smaller than that in the field, but the resistant-to-susceptible order among the cultivars were consistent with that in the field. Rapid identification technique of rice SB resistance in seedling stage proposed in this study could be used for screening resistance of rice germplasm on a large-scale scope or for primary resistance identification.
2. The SB resistance levels evaluated with the lesion length among5cultivars inoculated at the late tillering stage in temperature-controlled greenhouse differed significantly(.P<0.01) and could be divided into three groups, relatively susceptible (Lemont), moderate susceptible to moderate resistant(Wuyujing3, Jasmine85, Teqing) and relatively resistant (YSBRl), and the resistant-to-susceptible order among five cultivars were consistent with the results tested in field verifying experiment at30th day after heading. Although the SB resistance levels among5cultivars inoculated at the same time in field differed significantly at1%levels, the resistant-to-susceptible order among five cultivars was inconsistent with that of the30th day after heading. Furthermore, the SB resistance levels among5cultivars inoculated at the early booting stage in field differed significantly at1%levels, and the resistant-to-susceptible order among five cultivars were consistent with that of the30th day after heading. These results indicated that the lesion length used as investigation index to evaluate rice SB resistance at the late tillering stage in greenhouse or at the early booting stage in field was feasible.
3. There were no differences between the SB disease rating data measured with the common Rush's investigation standard and with the "order of leaf sheath" investigation standard in five rice cultivars inoculated at the late tillering stage in different years and locations, and the correlation index of SB disease rating data between two investigation standards in different years and locations were0.9455,0.9846(P<0.01), respectively, which indicated that the "order of leaf sheath" investigation standard as a new standard could replace Rush's investigation standard completely in evaluating rice SB disease rating. The difference of the SB disease rating data measured with the relative disease rating investigation standard among five rice cultivars inoculated at the early booting stage was not only significant (P<0.01), but also greater than that among the cultivars inoculated at the late tillering stage by means of the "order of leaf sheath" investigation standard. The resistant-to-susceptible order among the five cultivars between two investigation standards were consistent completely, which showed that the relative disease rating investigation standard as a new standard could also be used for evaluating rice SB disease rating in some feasible conditions. The application scopes of relative disease rating investigation standard in genetic research of resistance to rice SB was limited.
4. In two years experiment(2009-2010), twelve resistant QTLs were located, of which qSB-1Le, qSB-5Le and qSB-8Le were from susceptible parent Lemont, and remaining QTLs were all from YSBR1. Among QTLs detected, qSB-2Y and qSB-12Y could be considered as the major effective QTLs, with the average contribution rate of28%and17%in2009and2010, respectively. Furthermore, the QTLs of13major agronomic traits were located and their relationship with the SB QTLs were analyzed. There existed significant correlation (mostly negative, P<0.01or P<0.05) between four agronomic traits (plant height, growth duration, panicle length, effective panicles) and SB disease rating. This result showed that the incidence of rice SB was affected by the traits of plant height, growth duration, panicle length, effective panicles, of which plant height had the most powerful effect on the SB disease rating.
5. The difference of SB resistance levels within a short-term (5days after inoculation) between Lemont and YSBR1was significant (P<0.05), and the infection cushion and lesion appaered in10and20h after inoculation, respectively. The quality of gene chips of different biological replicates were reliable,2826differentially expressed genes in Lemont whole genome and748differentially expressed genes in YSBR1whole genome were screened out by using our screening standard respectively. Furthermore,247specific expressed differentially genes in YSBR1were also screened out. The specific expressed differentially genes in YSBR1were distributed among12rice chromosomes, and31genes fell into the detected QTL intervals, with the proportion to the total of12.55%. There were9and3YSBR1specific expressed differentially genes fell into the intervals of two major QTLs (qSB-2Y, qSB-12Y). Four and2genes were upregulated, and5and1genes were downregulated in qSB-2Y and qS.B-12Y intervals, respectively.
水稻纹枯病接种鉴定体系涉及到接种方法、接种时间、调查时间、调查标准等多种因素,其合适与否关系到水稻抗纹枯病遗传育种研究的进展。合适的接种鉴定体系不仅可减轻相关研究工作的工作强度,而且能提高试验的可靠性、重复性及结果的精确性。鉴此,本试验以抗感反应不同的5个水稻品种[Lemont、武育粳3号(Wuyujing3)、YSBR1、特青(Teqing)和Jasmine85(J-85)]为试验材料,在人工气候箱、控温室中进行水稻苗期抗纹枯病接种试验,并与田间相应的成株期抗性试验进行比较,初步建立了水稻苗期接种鉴定体系。在此基础上,采用牙签嵌入法在控温室、田间对各试验水稻材料进行成株期不同阶段(分蘖末期、小孕穗期)抗纹枯病接种试验,并于接种后3-11d、抽穗后30d测量病斑长度和采用3种不同调查标准调查病级,对水稻成株期纹枯病的接种鉴定方法进行了发展,并对田间水稻抽穗后30d适宜的纹枯病病级调查标准进行了新探索。
YSBR1是本研究组选育并经过多年鉴定为较抗病的新种质,目前对其抗性研究报道较少。本试验以YSBR1及易感品种Lemont为试验材料,通过构建这2个品种的F2无性系群体,对其开展了多年份的纹枯病抗性鉴定及农艺性状调查,同时对YSBR1中相关纹枯病抗性QTL、农艺性状QTL进行初步定位,研究这2类QTL之间的相互关系。在此基础上,又以这2个品种为试验材料,对其进行纹枯病病原菌接种,分别于接种后10、20h取样用于基因芯片表达谱分析,从中筛选差异表达基因,尤其是YSBR1特异差异表达基因。最终对YSBR1特异差异表达基因进行pathway分析和GO (gene ontology)分析,并将其与已检测到YSBR1的QTL区间进行整合。
主要研究结果如下：
1.85%的相对湿度为纹枯病菌侵染危害水稻苗期植株的适宜湿度；苗期5个水稻品种对纹枯病抗性差异极显著,可将其分为相对感病(Lemont、武育粳3号)和相对抗病(YSBR1、Jasmine85、特青)2大类；接种叶龄对发病程度有显著的影响,5个品种在4叶期接种时的平均病级显著高于5叶期接种的平均病级；苗期水稻品种间抗感差异小于田间鉴定试验结果,但两者间品种抗感趋势基本一致。表明,苗期快速鉴定技术可用于大规模水稻品种(组合)的抗性筛选或初步鉴定。
2.以病斑长度为调查指标,控温室中分蘖末期接种的5个水稻品种间抗感差异极显著,Lemont和YSBR1表现为稳定的感病、抗病,且和另外3个品种(武育粳3号、特青、Jasmine85)差异极显著,各品种间的抗感趋势与田间抽穗后30d的验证试验结果基本一致；而田间分蘖末期接种的5个水稻品种间虽然抗感差异极显著,但品种间抗感趋势与同期温室及田间抽穗后30d的验证试验结果差异不一致。田间小孕穗期接种的5个水稻品种间抗感差异极显著,品种间抗感趋势与分蘖末期接种并于抽穗后30d调查病级的田间验证试验结果完全一致。表明,温室条件下分蘖末期或田间条件下小孕穗期以病斑长度作为衡量纹枯病抗性的调查指标具有一定的可行性。
3.在分蘖末期接种的各参试品种,采用叶鞘位调查标准的结果与常用的Rush调查标准的结果相比较,两者在不同年份间、地点间差异均不显著,相关分析结果显示,2种调查标准间相关性在不同年份、地点间分别为0.9455、0.9846,均达到极显著水平,表明叶鞘位标准作为1个新的调查标准在田间纹枯病病级调查时完全可替代Rush调查标准；对于小孕穗期接种的各品种,采用相对病级调查标准,各品种间不仅抗性差异极显著,而且品种间抗感差异大于分蘖末期接种采用叶鞘位调查标准的相应对照,并且这2种调查标准间各品种的抗、感趋势完全一致。在水稻抗纹枯病遗传研究中,小孕穗期接种以相对病级作为调查标准,其适用范围是有限制的。
4.两年间(2009-2010年)共初步定位到12个抗性QTL,除qSB-1Le、qSB-5Le和qSB-8Le来自感病亲本Lemont外,其余均来自抗病亲本YSBR1,其中qSB-2Y、qSB-12Y被认为是主效QTL,两年的平均贡献率分别为28%和17%。此外,两年间13个主要农艺性状中有4个农艺性状(株高、生育期、穗长、有效穗数)和纹枯病发病程度存在显著或极显著相关,且多为负相关。表明水稻纹枯病的发病情况受到株高、生育期、穗长、有效穗数等性状的影响,尤其是株高的影响最大。
5.接种后短期内(5d),YSBR1与Lemont间抗感差异明显,接种后10h均产生侵染垫,接种后20h均出现病斑；不同生物学重复芯片质量可靠,按自定筛选标准,在Lemont全基因组共筛选到2826个差异表达基因,在YSBR1全基因组筛选到748个差异表达基因,其中在YSBR1中特异表达的差异基因有247个；247个YSBR1中特异表达基因分布水稻12条染色体上,落入已检测到的QTL区间的为31个,占总数比例为12.55%;落入2个主效QTLqSB-2Y、qSB-12Y区间的YSBRl特异表达基因分别有9、3个,其中qSB-2Y区间9个特异差异表达基因中有4个上调表达,5个下调表达；qSB-12Y区间3个特异差异表达基因中有2个上调表达,1个下调表达。
Sheath blight (SB) disease is one of the three most serious diseases in rice worldwide, and causes severe loss in yield and quality under favorable environmental condition. The resistance of rice to SB is a typical quantitative trait, and resources with strong resistance against this disease in the existing rice germplasm are not found.
Rice SB inoculation identification system involves the inoculation method, inoculation time, investigation time, investigation standard and other factors, and the appropriateness of the system can affect genetic and breeding research progress of the resistance to SB in rice. Appropriate inoculation identification system can not only reduce the strength of the research work, but also improve the reliability, repeatability, and accuracy of the test results. In view of this point, five rice cultivars with different levels of resistance to SB were adopted to identify their resistance to SB at seedling stage in artificial climate incubator or temperature-controlled greenhouse, and the test results were verified through a field experiment during rice adult stage, then a SB inoculation identification system in rice seedling stage was established preliminarily. Based on the above study, five rice cultivars were inoculated with toothpick-inserting inoculation method during different times in rice adult stage (the late tillering stage, the early booting stage) in field and temperature-controlled greenhouse. The lesion length was measured during3-11days after inoculation, and the SB disease rating at the30th day after heading was investigated with the reference of three different investigation standards. Then, some new inoculation identification methods in rice adult stage were developed and some new investigation standards of SB disease rating at the30th day after heading stage in field were also explored in this study.
YSBR1developed by our own research group is a new germplasm with stable and high resistance to sheath blight, which has been identified by using many methods in different experimental sites for many years. However, information of its genetic background relevant to SB resistance has been in scarce yet. A F2clonal populations (F2CP) derived from the cross between YSBR1and Lemont (a susceptible cultivar) was constructed, and the identification of resistance to SB and the investigation of agronomic traits for F2CP in a multi-year experiment were carried out in this study. At the same time, the QTLs mapping of resistance to SB in YSBR1and agronomic traits were conducted, and the interrelation between resistance to SB and agronomic traits were also analyzed. On the basis of this study, the two cultivars (YSBRl, Lemont) were used as materials to inoculate SB pathogens, and their leaf sheaths were sampled at10and20hours after inoculation respectively for gene chip expression profiling analysis and screening of differentially expressed genes, especially YSBR1specific differentially expressed gene. Furthermore, the pathway and GO (gene ontology) of YSBR1specific differentially expressed gene and its integration with QTL detected in YSBRl were analyzed.
The main results were as follows:
1. The optimal relative humidity for SB mycelium to develop infection structures and to infect rice seedling was approximately85%. The SB resistance levels in5cultivar seedlings differed significantly (P<0.01) and could be divided into two groups, relatively susceptible (Lemont, WuYujing3) and relatively resistant (YSBR1, Jasmine85, Teqing). The symptom severeness degrees of the seedlings at different leaf-age (4-leaf stage and5-leaf stage) were significantly different and the average disease rating at4-leaf stage was significantly higher than that at5-leaf stage. The differences of resistance among different cultivars in greenhouse were smaller than that in the field, but the resistant-to-susceptible order among the cultivars were consistent with that in the field. Rapid identification technique of rice SB resistance in seedling stage proposed in this study could be used for screening resistance of rice germplasm on a large-scale scope or for primary resistance identification.
2. The SB resistance levels evaluated with the lesion length among5cultivars inoculated at the late tillering stage in temperature-controlled greenhouse differed significantly(.P<0.01) and could be divided into three groups, relatively susceptible (Lemont), moderate susceptible to moderate resistant(Wuyujing3, Jasmine85, Teqing) and relatively resistant (YSBRl), and the resistant-to-susceptible order among five cultivars were consistent with the results tested in field verifying experiment at30th day after heading. Although the SB resistance levels among5cultivars inoculated at the same time in field differed significantly at1%levels, the resistant-to-susceptible order among five cultivars was inconsistent with that of the30th day after heading. Furthermore, the SB resistance levels among5cultivars inoculated at the early booting stage in field differed significantly at1%levels, and the resistant-to-susceptible order among five cultivars were consistent with that of the30th day after heading. These results indicated that the lesion length used as investigation index to evaluate rice SB resistance at the late tillering stage in greenhouse or at the early booting stage in field was feasible.
3. There were no differences between the SB disease rating data measured with the common Rush's investigation standard and with the "order of leaf sheath" investigation standard in five rice cultivars inoculated at the late tillering stage in different years and locations, and the correlation index of SB disease rating data between two investigation standards in different years and locations were0.9455,0.9846(P<0.01), respectively, which indicated that the "order of leaf sheath" investigation standard as a new standard could replace Rush's investigation standard completely in evaluating rice SB disease rating. The difference of the SB disease rating data measured with the relative disease rating investigation standard among five rice cultivars inoculated at the early booting stage was not only significant (P<0.01), but also greater than that among the cultivars inoculated at the late tillering stage by means of the "order of leaf sheath" investigation standard. The resistant-to-susceptible order among the five cultivars between two investigation standards were consistent completely, which showed that the relative disease rating investigation standard as a new standard could also be used for evaluating rice SB disease rating in some feasible conditions. The application scopes of relative disease rating investigation standard in genetic research of resistance to rice SB was limited.
4. In two years experiment(2009-2010), twelve resistant QTLs were located, of which qSB-1Le, qSB-5Le and qSB-8Le were from susceptible parent Lemont, and remaining QTLs were all from YSBR1. Among QTLs detected, qSB-2Y and qSB-12Y could be considered as the major effective QTLs, with the average contribution rate of28%and17%in2009and2010, respectively. Furthermore, the QTLs of13major agronomic traits were located and their relationship with the SB QTLs were analyzed. There existed significant correlation (mostly negative, P<0.01or P<0.05) between four agronomic traits (plant height, growth duration, panicle length, effective panicles) and SB disease rating. This result showed that the incidence of rice SB was affected by the traits of plant height, growth duration, panicle length, effective panicles, of which plant height had the most powerful effect on the SB disease rating.
5. The difference of SB resistance levels within a short-term (5days after inoculation) between Lemont and YSBR1was significant (P<0.05), and the infection cushion and lesion appaered in10and20h after inoculation, respectively. The quality of gene chips of different biological replicates were reliable,2826differentially expressed genes in Lemont whole genome and748differentially expressed genes in YSBR1whole genome were screened out by using our screening standard respectively. Furthermore,247specific expressed differentially genes in YSBR1were also screened out. The specific expressed differentially genes in YSBR1were distributed among12rice chromosomes, and31genes fell into the detected QTL intervals, with the proportion to the total of12.55%. There were9and3YSBR1specific expressed differentially genes fell into the intervals of two major QTLs (qSB-2Y, qSB-12Y). Four and2genes were upregulated, and5and1genes were downregulated in qSB-2Y and qS.B-12Y intervals, respectively.
引文
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