高抗菌核病野生甘蓝C01抗性机理及其应用
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摘要
油菜是人类主要的植物油来源之一,也是不可再生能源物质的重要替代品,广泛栽培于欧洲、北美洲及亚洲。菌核病是油菜最严重的病害之一,其流行可导致油菜减产10—70%,含油量下降1%—5%。随着机械化栽培倡导高密度栽培和气候条件的变迁,该病害可能日趋严重。油菜菌核病的病源菌为核盘菌,该真菌对植物的侵染具有广谱性,可对400多种植物产生侵染。目前人们采用多种方法来防治油菜菌核病。轮作法因为核盘菌的寄主广谱性而收效甚微,化学防控的效果也受多种因素的限制,且会增加油菜生产成本、污染环境。防治菌核病最有效、最经济的方法就是选育抗性品种。然而,现在油菜中匮乏抗病性资源,阻碍了油菜菌核病抗性改良育种和相关的基础研究。
近缘物种抗性资源的发掘与利用是改良油菜菌核病抗性的有效途径。甘蓝与甘蓝型油菜同属十字花科芸薹属物种。与甘蓝型油菜相比,甘蓝有多种栽培型和野生型,遗传多样性丰富,可用于拓宽甘蓝型油菜的遗传基础。重庆市油菜工程技术研究中心从国内外收集了大量的野生甘蓝和栽培甘蓝资源,经菌核病抗性鉴定,发现一份来自Brassica incana野生甘蓝(C01)的菌核病抗性突出,为甘蓝型油菜菌核病抗性改良带来了新的希望。
本研究以野生甘蓝C01及另一份感病甘蓝C41为主要研究材料,观察了该野生甘蓝对核盘菌菌丝的生长、入侵的影响,测定了核盘菌胁迫下叶片中草酸含量的时空变化,分析了多聚半乳糖醛酸酶抑制蛋白编码基因、类萌发素编码基因及SA/JA信号转导途径中相关基因相对表达量动态变化,并以C01为亲本创造了人工合成甘蓝型油菜,检测了人工合成甘蓝型油菜的菌核病抗性,分析了抗性突出的人工合成甘蓝型油菜RB165在核盘菌胁迫下几个病程蛋白编码基因相对表达的动态变化。现将主要结果阐述如下：
(1)野生甘蓝C01影响核盘菌菌丝的生长、侵染垫的发育
通过光学显微镜和扫描电镜观察了C01、C41对核盘菌菌丝形态及侵染垫发育的影响,结果表明,较C41而言,菌丝在C01叶片生长缓慢,侵染垫的发育延缓,接种后期形成的侵染垫疏松,且在叶面的分布密度小。
(2)野生甘蓝C01草酸本底含量高且受核盘菌胁迫影响较小
用HPLC测定了核盘菌胁迫下,C01、C41叶片中草酸含量的动态变化,发现C01中的草酸本底值高于C41,其草酸含量随接种时间的延长有所增高,但其变化幅度小于C41的变化幅度。在侵染后期,病斑处的草酸含量较高,但低于病健区草酸含量,在病健区外围,草酸含量更低。
(3)C01中PGlP基因受核盘菌胁迫诱导表达
检测核盘菌胁迫下,C01、C41叶片多聚半乳糖醛酸酶抑制蛋白编码基因(PGIP)5个成员相对表达量的动态变化,结果表明PGIP1、PGIP5受核盘菌诱导表达,而PGIP9.PGIPl2表达受核盘菌胁迫抑制,PGIP2在甘蓝(C01、C41)中表达量低,在甘蓝型油菜中油821(ZY821)中受核盘菌胁迫诱导,C01和C41中PGIP基因的表达模式相似,但在C01中的相对表达量较高。
(4)C01中GLP基因的表达量受核盘菌胁迫诱导表达
CLP基因受核盘菌的诱导表达,但GLP3在接种早期受到诱导表达,GLP12在接种后期被诱导表达,且两基因在C01中的相对表达量高于C41。
(5)C01的防御反应受SA、JA途径协作调控
核盘菌胁迫下,SA、JA信号途径部分标志基因相对表达都不同程度受到诱导表达,SA、JA信号转导途径都被激活,可能两条途径交叉作用,利于甘蓝、甘蓝型油菜抗性产生,但是SA途径较JA途径被较早激活。
(6)以C01为亲本获得了抗性改良的人工合成甘蓝型油菜
以C01为亲本,获得了人工合成甘蓝型油菜。通过离体叶片鉴定、离体茎杆鉴定检测了人工合成甘蓝型油菜的菌核病抗性,结果表明,人工合成甘蓝型油菜的抗性强于对照甘蓝型油菜中油821,且甘蓝材料抗性与人工合成甘蓝型油菜抗性中度相关。
(7)人工合成甘蓝型油菜的抗性与几个病程蛋白编码基因表达水平并无直接关系
人工合成甘蓝型油菜RB165的菌核病抗性低于其抗病亲本C01,但高于目前广泛栽培的耐菌核病油菜品种ZY821(中油821)和ZS9(中双9号)；试材对菌核病的抗性水平与OXO、Cu/Zn SOD、PR2、PR3的表达动态变化没有直接关系。
Oilseed rape, not only one of the main sources of plant edible oil for humankind but also the alternative of the non-renewable energy resource, is widely grown in Europe, North America and Asia. Sclerotinia stem rot, infected by Sclerotinia sclerotiorum, is a devastating threat to oilseed rape production with substantial yield loss and quality decline. The burst of the pathogen decrease the yield by10%-70%and deprive the oil content by1%-5%. S. slcerotiorum is a non-specific plant pathogen, which can infect more than400plant species.As the advocation of cultivation with high plant density and the exacerbation of the climate, the damage of the disease may become more serious. Many approaches have been adopted to control the disease including soil amendment, crop rotation, chemical application and variety breeding. However, the effect of cultural practice is limited because of the persistent nature of sclerotia and the wide host range of this pathogen. And chemical control effect dependes on the match of ascospores release with fungicide application during the grow season. In addition, chemical fungicide possiblly pollutes the environment and increases the cost. Among the measures, variety breeding focusing on high resistance is the most effective and economical choice. Whereas, there is no resistance in current Brassica napus gene pools, which hinders the resistance breeding and the relative basic research.
Compared with B. napus, B. oleracea possesses various cultivated and wild genotypes, which can be used to widen the genetic basis of B. napus. A line of wild B. oleracea, B. incana (C01) was identified with high resistance against sclerotinia rot at Chongqing Rapeseed Engineering and Technology Research Center, which is tremendously potential to to improve the resistance againt S. sclerotiorum in B. napus.
Using C01and C41(another B.oleracea, susceptible to S. sclerotiorum) as the main experimental materials, in the present study we observed the different influence of them on the growth and invasion of the hyphae; measured the dynamics change of oxalic acid content; detected the relative expression of the genes encoding PGIPs and GLP, as well as the marker genes involved in SA and JA signal transduction pathway; resynthesized RS line using C01as parent and identified the resistance; analysized relationship between the relative expression of PR genes and the resistance in RB165, a newly resynthsized B. napus with high resistance. The results are shown as following,
1) C01affected the growth and infection cushions of S. sclerotiorum
The morphological character of the hyphae and the formation of the infection cushions on the infected leaves of C01, C41was observed with light microscope and scanning electronic microscope. Compared with C41, C01inhibited the growth and delayed the formation of infection cushions. In addition, the infection cushions on C01leaves were fluffy and scattered on the infected leaves.
2) Immanent content of oxalic acid in C01is higher and the increment is lessly influenced by S. sclerotiorum infection
The content of oxalic acid in the leaves of C01and C41infected by S. slcerotiorum was measured with HPLC. The result showed that C01possessed higher oxalaic acid inherently. After the inoculation with S. sclerotiorum, the content of oxalic acid increased but the increase was lower than that in C41. The content of oxalic acid in the lesion is lower than that in the interface part between the lesion and the healthy tissues, but higher than that of the outside of the interface part.
3) Expression of several members of PGIPs were induced by S. slcerotiorum
The relative expression of five members of PGIPs was detected in the leaves of C01and C41challenged by S. sclerotiorum. It indicated that PGIP1and PGIP5were induced by the pathogen. On the contrast, PGIP9and PGIP12were inhibited. The expression of PGIP2was induced in B. napus but inhibited in B. oleracea. The expression pattern of PGIP in C01and C41was similar, but the expression level in C01was higher than that in C41.
4) Expression of GLPs was induced by S. sclerotiorum
Two members of GLPs, namely GLP3and GLP12, were induced by S. sclerotiorum, but the pattern was different among the members. GLP3was induced in the early inoculation stage and the induced effect in GLP12emerged in the later stage. As far as the expression level was concerned, it was higher in CO1than that in C41.
5) Defense response in C01was regulated by the coolaboration of SA and JA signal transduction
Partial marker genes involved in SA and JA signal transduction were induced by S. slcerotiorum to some extent. It is implied that the crosstalk between the ways regulated the response, but the induction related to SA was earlier than JA.
6) Improvement of resistance against S. sclerotiorum in resynthesized B. napus using C01as resistance donor
Resynthesized B. napus were obtained using C01as parent hybred with other Brassicas. The resistance in the resynthesized B. napus was identified through detched leaves and stems. The results showed that the resistance in resynthesized B napus was better than that of ZY821and the correlation of the resistance between resynthesized B. napus and the corresponding donor was moderate. RB165is one of the resynthesized B. napus with high resistance. It was more resistant to S. sclerotiorum than natural rapeseed, such ZY821and ZS9(Zhongshuang9). However, the resistance of RB165is not consistent to the expression of some PR genes, i.e., OXO, Cu/Zn SOD, PR2and PR3.
近缘物种抗性资源的发掘与利用是改良油菜菌核病抗性的有效途径。甘蓝与甘蓝型油菜同属十字花科芸薹属物种。与甘蓝型油菜相比,甘蓝有多种栽培型和野生型,遗传多样性丰富,可用于拓宽甘蓝型油菜的遗传基础。重庆市油菜工程技术研究中心从国内外收集了大量的野生甘蓝和栽培甘蓝资源,经菌核病抗性鉴定,发现一份来自Brassica incana野生甘蓝(C01)的菌核病抗性突出,为甘蓝型油菜菌核病抗性改良带来了新的希望。
本研究以野生甘蓝C01及另一份感病甘蓝C41为主要研究材料,观察了该野生甘蓝对核盘菌菌丝的生长、入侵的影响,测定了核盘菌胁迫下叶片中草酸含量的时空变化,分析了多聚半乳糖醛酸酶抑制蛋白编码基因、类萌发素编码基因及SA/JA信号转导途径中相关基因相对表达量动态变化,并以C01为亲本创造了人工合成甘蓝型油菜,检测了人工合成甘蓝型油菜的菌核病抗性,分析了抗性突出的人工合成甘蓝型油菜RB165在核盘菌胁迫下几个病程蛋白编码基因相对表达的动态变化。现将主要结果阐述如下：
(1)野生甘蓝C01影响核盘菌菌丝的生长、侵染垫的发育
通过光学显微镜和扫描电镜观察了C01、C41对核盘菌菌丝形态及侵染垫发育的影响,结果表明,较C41而言,菌丝在C01叶片生长缓慢,侵染垫的发育延缓,接种后期形成的侵染垫疏松,且在叶面的分布密度小。
(2)野生甘蓝C01草酸本底含量高且受核盘菌胁迫影响较小
用HPLC测定了核盘菌胁迫下,C01、C41叶片中草酸含量的动态变化,发现C01中的草酸本底值高于C41,其草酸含量随接种时间的延长有所增高,但其变化幅度小于C41的变化幅度。在侵染后期,病斑处的草酸含量较高,但低于病健区草酸含量,在病健区外围,草酸含量更低。
(3)C01中PGlP基因受核盘菌胁迫诱导表达
检测核盘菌胁迫下,C01、C41叶片多聚半乳糖醛酸酶抑制蛋白编码基因(PGIP)5个成员相对表达量的动态变化,结果表明PGIP1、PGIP5受核盘菌诱导表达,而PGIP9.PGIPl2表达受核盘菌胁迫抑制,PGIP2在甘蓝(C01、C41)中表达量低,在甘蓝型油菜中油821(ZY821)中受核盘菌胁迫诱导,C01和C41中PGIP基因的表达模式相似,但在C01中的相对表达量较高。
(4)C01中GLP基因的表达量受核盘菌胁迫诱导表达
CLP基因受核盘菌的诱导表达,但GLP3在接种早期受到诱导表达,GLP12在接种后期被诱导表达,且两基因在C01中的相对表达量高于C41。
(5)C01的防御反应受SA、JA途径协作调控
核盘菌胁迫下,SA、JA信号途径部分标志基因相对表达都不同程度受到诱导表达,SA、JA信号转导途径都被激活,可能两条途径交叉作用,利于甘蓝、甘蓝型油菜抗性产生,但是SA途径较JA途径被较早激活。
(6)以C01为亲本获得了抗性改良的人工合成甘蓝型油菜
以C01为亲本,获得了人工合成甘蓝型油菜。通过离体叶片鉴定、离体茎杆鉴定检测了人工合成甘蓝型油菜的菌核病抗性,结果表明,人工合成甘蓝型油菜的抗性强于对照甘蓝型油菜中油821,且甘蓝材料抗性与人工合成甘蓝型油菜抗性中度相关。
(7)人工合成甘蓝型油菜的抗性与几个病程蛋白编码基因表达水平并无直接关系
人工合成甘蓝型油菜RB165的菌核病抗性低于其抗病亲本C01,但高于目前广泛栽培的耐菌核病油菜品种ZY821(中油821)和ZS9(中双9号)；试材对菌核病的抗性水平与OXO、Cu/Zn SOD、PR2、PR3的表达动态变化没有直接关系。
Oilseed rape, not only one of the main sources of plant edible oil for humankind but also the alternative of the non-renewable energy resource, is widely grown in Europe, North America and Asia. Sclerotinia stem rot, infected by Sclerotinia sclerotiorum, is a devastating threat to oilseed rape production with substantial yield loss and quality decline. The burst of the pathogen decrease the yield by10%-70%and deprive the oil content by1%-5%. S. slcerotiorum is a non-specific plant pathogen, which can infect more than400plant species.As the advocation of cultivation with high plant density and the exacerbation of the climate, the damage of the disease may become more serious. Many approaches have been adopted to control the disease including soil amendment, crop rotation, chemical application and variety breeding. However, the effect of cultural practice is limited because of the persistent nature of sclerotia and the wide host range of this pathogen. And chemical control effect dependes on the match of ascospores release with fungicide application during the grow season. In addition, chemical fungicide possiblly pollutes the environment and increases the cost. Among the measures, variety breeding focusing on high resistance is the most effective and economical choice. Whereas, there is no resistance in current Brassica napus gene pools, which hinders the resistance breeding and the relative basic research.
Compared with B. napus, B. oleracea possesses various cultivated and wild genotypes, which can be used to widen the genetic basis of B. napus. A line of wild B. oleracea, B. incana (C01) was identified with high resistance against sclerotinia rot at Chongqing Rapeseed Engineering and Technology Research Center, which is tremendously potential to to improve the resistance againt S. sclerotiorum in B. napus.
Using C01and C41(another B.oleracea, susceptible to S. sclerotiorum) as the main experimental materials, in the present study we observed the different influence of them on the growth and invasion of the hyphae; measured the dynamics change of oxalic acid content; detected the relative expression of the genes encoding PGIPs and GLP, as well as the marker genes involved in SA and JA signal transduction pathway; resynthesized RS line using C01as parent and identified the resistance; analysized relationship between the relative expression of PR genes and the resistance in RB165, a newly resynthsized B. napus with high resistance. The results are shown as following,
1) C01affected the growth and infection cushions of S. sclerotiorum
The morphological character of the hyphae and the formation of the infection cushions on the infected leaves of C01, C41was observed with light microscope and scanning electronic microscope. Compared with C41, C01inhibited the growth and delayed the formation of infection cushions. In addition, the infection cushions on C01leaves were fluffy and scattered on the infected leaves.
2) Immanent content of oxalic acid in C01is higher and the increment is lessly influenced by S. sclerotiorum infection
The content of oxalic acid in the leaves of C01and C41infected by S. slcerotiorum was measured with HPLC. The result showed that C01possessed higher oxalaic acid inherently. After the inoculation with S. sclerotiorum, the content of oxalic acid increased but the increase was lower than that in C41. The content of oxalic acid in the lesion is lower than that in the interface part between the lesion and the healthy tissues, but higher than that of the outside of the interface part.
3) Expression of several members of PGIPs were induced by S. slcerotiorum
The relative expression of five members of PGIPs was detected in the leaves of C01and C41challenged by S. sclerotiorum. It indicated that PGIP1and PGIP5were induced by the pathogen. On the contrast, PGIP9and PGIP12were inhibited. The expression of PGIP2was induced in B. napus but inhibited in B. oleracea. The expression pattern of PGIP in C01and C41was similar, but the expression level in C01was higher than that in C41.
4) Expression of GLPs was induced by S. sclerotiorum
Two members of GLPs, namely GLP3and GLP12, were induced by S. sclerotiorum, but the pattern was different among the members. GLP3was induced in the early inoculation stage and the induced effect in GLP12emerged in the later stage. As far as the expression level was concerned, it was higher in CO1than that in C41.
5) Defense response in C01was regulated by the coolaboration of SA and JA signal transduction
Partial marker genes involved in SA and JA signal transduction were induced by S. slcerotiorum to some extent. It is implied that the crosstalk between the ways regulated the response, but the induction related to SA was earlier than JA.
6) Improvement of resistance against S. sclerotiorum in resynthesized B. napus using C01as resistance donor
Resynthesized B. napus were obtained using C01as parent hybred with other Brassicas. The resistance in the resynthesized B. napus was identified through detched leaves and stems. The results showed that the resistance in resynthesized B napus was better than that of ZY821and the correlation of the resistance between resynthesized B. napus and the corresponding donor was moderate. RB165is one of the resynthesized B. napus with high resistance. It was more resistant to S. sclerotiorum than natural rapeseed, such ZY821and ZS9(Zhongshuang9). However, the resistance of RB165is not consistent to the expression of some PR genes, i.e., OXO, Cu/Zn SOD, PR2and PR3.
引文
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