紫外线诱导小麦条锈菌致病性突变及其突变菌系的RAPD分析
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摘要
由条形柄锈菌小麦专化型（Puccinia striiformis f. sp. tritici）引起的小麦条锈病是我国小麦生产上最重要的病害，造成严重的产量和经济损失。种植抗病品种是当前防治小麦条锈病最经济有效的措施，但抗病品种常因小麦条锈菌新小种的不断产生和发展而“丧失”抗锈性。新小种产生主要是由于毒性变异所致，一般认为小麦条锈菌的毒性变异是通过基因突变、异核作用等途径发生的，其中基因突变是产生新的毒性基因的唯一途径。由紫外线照射引起的小麦条锈菌变异的生物学效应，前人已进行了研究，但尚未见对变异在分子水平上的研究报道，RAPD是揭示基因组DNA差异的有效方法，可以对整个基因组进行多态性检测，对单一引物来讲，DNA扩增片段的差异反映了基因组水平的差异，因此，本研究通过紫外线处理小麦条锈菌获得致病性变异的突变体，进而用RAPD技术对突变体进行分析，从而为研究小麦条锈菌的毒性基因变异机制奠定基础。并通过苗期接种试验，进一步明确致病性变异菌系预测小麦品种抗锈稳定性的可能性。
1. 用紫外线以不同时间处理小麦条锈菌水源致病类型1-4、条中29-3、条中23-2，当夏孢子致死率达90%左右时，确定为最适诱变时间分别为12min、8min、5min。以紫外线12min处理水源致病类型1时，经6批次筛选未获得突变体。以8min处理条中29-3，得到在Hybrid46上表现3型的菌系，在尤皮Ⅱ号上表现4型的菌系和在阿夫上表现2型的菌系，分别命名Hybrid46菌系、尤Ⅱ菌系和阿夫菌系，三菌系经4代转接该品种，最后获得2个稳定的突变菌系，包括毒性突变产生的尤Ⅱ菌系和非毒性突变产生的阿夫菌系。以5min处理条中23-2得到两个突变菌系：在尤皮Ⅱ号上表现3型的菌系和在水源11上表现2型的菌系，分别命名为尤Ⅱ-23菌系和水源11菌系，其中尤Ⅱ-23菌系经4次转接仍保持毒性。
以上获得的三个稳定突变菌系经鉴别寄主鉴定，毒性谱均发生了较大变化。
2. 对原始菌系和突变菌系进行RAPD分析后发现，条中29-3的突变菌系尤Ⅱ菌系、阿夫菌系与原始菌系DNA之间的多态性存在显著差异，多态率分别为10.58%和11.57%；条中23-2的突变菌系尤Ⅱ-23菌系与原始菌系DNA的多态性差异亦为10.73%，由此可知紫外线可使小麦条锈菌基因组DNA发生广泛的变异且突变位点比较复杂。
3. 通过用条中29号、条中31号和条中32号及条中29号的两个突变菌系、条中23号的突变菌系对2004年黄淮麦区区试品种进行苗期抗锈性鉴定，明确致病性变异菌系预测小麦品种抗锈稳定性的可能性。用混合菌系（条中29、条中31、条中32、水源致病类型14）进行成株期鉴定后与苗期比较发现，苗期仅有一个品种周麦17对三小种均表现抗病而成株期有10个品种表现抗病，大部分可能为慢锈型或高温抗锈性品种。
Wheat stripe rust, caused by Puccinia striiformis f.sp. tritici, is the most important disease in wheat production in China. Growing rust-resistant cultivars is the most economical and effective measure. However, the resistance usually was breakdown in wheat cultivars, and continuous occurrence of new virulent races was the main cause. Virulence variation caused occurrence of new virulent races. It had been proved that the approaches to virulence variation were mutation, heterokaryosis, and so on, in which gene mutation was the exclusive approach to producing new virulence genes. The biological effects on the stripe rust fungus induced by UV, had been reported; but the reports about variation on level of molecule in Puccinia striiformis f.sp. tritici were not found. RAPD analyses may detect the polymorphisms in genomic DNA. Most single nucleotide changes in a primer sequence caused a complete change in the pattern of amplified DNA segment. The stripe rust urediospores were treated by UV to obtain the mutant cultures in pathogenicity, and the cultures were analyzed with RAPD technique in order to lay the foundation for studying the mechanism of the virulent gene variation. The probability of forecasting the rust resistance stability of the wheat cultivars was verified by seeding inoculation tests.
1. Urediospores of the Shuiyuan 11 pathogenic type1-4, CY29-3 and CY23-2 were treat by UV. The proper time treated was12 minutes, 8 minutes and 5 minutes respectively, when the urediospores lethality rate reached to 90% or so. Mutants in pathogenicity were not obtained after urediospores of Su-1-4 were treated for 12 minutes through six bathes of selection. Urediospores of CY29-3 were treated for 8 minutes, and three mutant cultures in pathogenicity were obtained including Hybrid 46 isolate, Jubilejina2 isolate and Funo isolate. Two isolates were obtain after stabilization for 4 generations, including Jubilejina2 isolate and Funo isolate. Reaction type of Jubilejina2 isolate was type 4 on Jubilejina2, but that of the original race was type 0; and reaction type of Funo isolate was 2, while that of the original race was 4. Urediospores of CY23-2 were treated for 5 minutes, and two mutant cultures in pathogenicity were obtained, namely Jubilejina2-23 isolate and Shuiyuan 11 isolate, in which Jubilejina2-23 isolate kept its virulence still after stabilization for 4 generations.
The pathogenicity of three mutant cultures on the differentials has changed greatly.
2. RAPD analyses were conducted between the original race and the mutant cultures. DNA polymorphisms of Jubilejina2 isolate and Funo isolate were quite distinct from CY29-3. Their polymorphic rates were 10.58% and 11.57%. DNA polymorphisms of Jubilejina2-23 isolate were also quite distinct from CY23-2. So UV irradiation could cause great variation of the genome DNA of the wheat stripe rust of fungus and the mutation loci were complicated.
3. Wheat stripe rust races CY29, 31, 32, two mutant cultures of CY29-3 and the mutant culture of CY23-2 were used to identify the seeding resistance on the different cultivars grown in Huanghe and Huaihe river wheat area. The results testified that the resistance stability of the wheat varieties could been forecasted by inoculation of the mutant cultures. The results identifying the adult plant resistance with the mixed races (CY29,CY31,CY32,Su-14) indicated that 10 varieties were resistant to mixed races, while only one cultivars, Zhoumai 17, was resistant to three wild cultures. Most of them could be the slow rust type or high temperature resistance to strip rust on adult plant.
1. 用紫外线以不同时间处理小麦条锈菌水源致病类型1-4、条中29-3、条中23-2，当夏孢子致死率达90%左右时，确定为最适诱变时间分别为12min、8min、5min。以紫外线12min处理水源致病类型1时，经6批次筛选未获得突变体。以8min处理条中29-3，得到在Hybrid46上表现3型的菌系，在尤皮Ⅱ号上表现4型的菌系和在阿夫上表现2型的菌系，分别命名Hybrid46菌系、尤Ⅱ菌系和阿夫菌系，三菌系经4代转接该品种，最后获得2个稳定的突变菌系，包括毒性突变产生的尤Ⅱ菌系和非毒性突变产生的阿夫菌系。以5min处理条中23-2得到两个突变菌系：在尤皮Ⅱ号上表现3型的菌系和在水源11上表现2型的菌系，分别命名为尤Ⅱ-23菌系和水源11菌系，其中尤Ⅱ-23菌系经4次转接仍保持毒性。
以上获得的三个稳定突变菌系经鉴别寄主鉴定，毒性谱均发生了较大变化。
2. 对原始菌系和突变菌系进行RAPD分析后发现，条中29-3的突变菌系尤Ⅱ菌系、阿夫菌系与原始菌系DNA之间的多态性存在显著差异，多态率分别为10.58%和11.57%；条中23-2的突变菌系尤Ⅱ-23菌系与原始菌系DNA的多态性差异亦为10.73%，由此可知紫外线可使小麦条锈菌基因组DNA发生广泛的变异且突变位点比较复杂。
3. 通过用条中29号、条中31号和条中32号及条中29号的两个突变菌系、条中23号的突变菌系对2004年黄淮麦区区试品种进行苗期抗锈性鉴定，明确致病性变异菌系预测小麦品种抗锈稳定性的可能性。用混合菌系（条中29、条中31、条中32、水源致病类型14）进行成株期鉴定后与苗期比较发现，苗期仅有一个品种周麦17对三小种均表现抗病而成株期有10个品种表现抗病，大部分可能为慢锈型或高温抗锈性品种。
Wheat stripe rust, caused by Puccinia striiformis f.sp. tritici, is the most important disease in wheat production in China. Growing rust-resistant cultivars is the most economical and effective measure. However, the resistance usually was breakdown in wheat cultivars, and continuous occurrence of new virulent races was the main cause. Virulence variation caused occurrence of new virulent races. It had been proved that the approaches to virulence variation were mutation, heterokaryosis, and so on, in which gene mutation was the exclusive approach to producing new virulence genes. The biological effects on the stripe rust fungus induced by UV, had been reported; but the reports about variation on level of molecule in Puccinia striiformis f.sp. tritici were not found. RAPD analyses may detect the polymorphisms in genomic DNA. Most single nucleotide changes in a primer sequence caused a complete change in the pattern of amplified DNA segment. The stripe rust urediospores were treated by UV to obtain the mutant cultures in pathogenicity, and the cultures were analyzed with RAPD technique in order to lay the foundation for studying the mechanism of the virulent gene variation. The probability of forecasting the rust resistance stability of the wheat cultivars was verified by seeding inoculation tests.
1. Urediospores of the Shuiyuan 11 pathogenic type1-4, CY29-3 and CY23-2 were treat by UV. The proper time treated was12 minutes, 8 minutes and 5 minutes respectively, when the urediospores lethality rate reached to 90% or so. Mutants in pathogenicity were not obtained after urediospores of Su-1-4 were treated for 12 minutes through six bathes of selection. Urediospores of CY29-3 were treated for 8 minutes, and three mutant cultures in pathogenicity were obtained including Hybrid 46 isolate, Jubilejina2 isolate and Funo isolate. Two isolates were obtain after stabilization for 4 generations, including Jubilejina2 isolate and Funo isolate. Reaction type of Jubilejina2 isolate was type 4 on Jubilejina2, but that of the original race was type 0; and reaction type of Funo isolate was 2, while that of the original race was 4. Urediospores of CY23-2 were treated for 5 minutes, and two mutant cultures in pathogenicity were obtained, namely Jubilejina2-23 isolate and Shuiyuan 11 isolate, in which Jubilejina2-23 isolate kept its virulence still after stabilization for 4 generations.
The pathogenicity of three mutant cultures on the differentials has changed greatly.
2. RAPD analyses were conducted between the original race and the mutant cultures. DNA polymorphisms of Jubilejina2 isolate and Funo isolate were quite distinct from CY29-3. Their polymorphic rates were 10.58% and 11.57%. DNA polymorphisms of Jubilejina2-23 isolate were also quite distinct from CY23-2. So UV irradiation could cause great variation of the genome DNA of the wheat stripe rust of fungus and the mutation loci were complicated.
3. Wheat stripe rust races CY29, 31, 32, two mutant cultures of CY29-3 and the mutant culture of CY23-2 were used to identify the seeding resistance on the different cultivars grown in Huanghe and Huaihe river wheat area. The results testified that the resistance stability of the wheat varieties could been forecasted by inoculation of the mutant cultures. The results identifying the adult plant resistance with the mixed races (CY29,CY31,CY32,Su-14) indicated that 10 varieties were resistant to mixed races, while only one cultivars, Zhoumai 17, was resistant to three wild cultures. Most of them could be the slow rust type or high temperature resistance to strip rust on adult plant.
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