河南省三个小麦孢囊线虫群体种类和致病型鉴定及品种抗性研究
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摘要
小麦禾谷孢囊线虫病(cereal cyst nematode)在我国北方小麦主要产区发生分布十分广泛,对小麦生产构成了潜在的威胁。河南是我国小麦第一生产大省,种植面积和总产量都居国内首位。近年来小麦孢囊线虫病在河南郑州和许昌两地区危害严重,造成了严重的产量损失。为了给该病防控工作提供理论依据,我们对郑州和许昌地区3个禾谷孢囊线虫群体的种类和致病型进行了鉴定,同时对国内外小麦品种及相关资源材料对孢囊线虫郑州和许昌群体的抗性进行了筛选,初步探索了小麦品种抗禾谷孢囊线虫的分子机理。主要研究结果如下:
通过对郑州和许昌小麦孢囊线虫群体的形态鉴定和分子鉴定,明确了郑州和许昌地区小麦孢囊线虫的种类:郑州须水和荥阳两个群体都属于禾谷孢囊线虫组中的燕麦孢囊线虫(Heterodera avenae Wollenweber, 1924),许昌群体属于禾谷孢囊线虫组中的菲利普孢囊线虫〔H.filipjevi(Madzhidov,1981)Stelter,1984〕。小麦孢囊线虫郑州两个群体成熟孢囊柠檬形,褐色至黑褐色。颈较长,阴门锥显著。阴门膜孔为双膜孔,卵圆形,有阴门桥,但没有下桥。尾锥内不规则分布着大量褐色泡状突。二龄幼虫蠕虫形,口针粗壮,口针基部球锚形。中食道球卵圆形。尾圆锥形,有较长的透明尾。许昌群体成熟孢囊褐色,柠檬形,头部与尾部明显。头部有颈,尾部有尾锥。阴门膜孔为双膜孔,马蹄形。有阴门桥和下桥结构。尾锥内不规则分布着褐色泡状突,泡状突较郑州的两个群体少。二龄幼虫蠕虫形,口针粗壮,口针基部球锚形。中食道球卵圆形。尾圆锥形,有较长的透明尾。
扩增郑州和许昌3个小麦孢囊线虫群体核糖体DNA的ITS区,PCR产物都产生1条大小约为980bp的片段。用TaqⅠ、AluⅠ、HaeⅢ、HinfⅠ、RsaⅠ、PstⅠ、CfoⅠ、Tru9Ⅰ等8种限制性内切酶酶切3个群体的rDNA–ITS–PCR产物,结果表明,郑州须水和荥阳两个群体的酶切图谱相同,它们的ITS属于“B”型;许昌群体的酶切图谱与已报道H. filipjevi的酶切图谱一致;TaqⅠ酶与PstⅠ酶均可将许昌与郑州两地的小麦孢囊线虫区分开来。rDNA-ITS序列聚类分析结果表明,郑州须水和荥阳两个群体的序列与国内过去报道的H.avenae序列亲缘关系较近,与澳大利亚的H. australis序列也有较近的亲缘关系。许昌群体的序列与国外已报道的H. filipjevi序列聚在同一簇。
利用燕麦孢囊线虫致病型国际鉴别寄主对郑州荥阳、须水和许昌3个群体的致病型进行鉴定,结果表明,根据国际鉴别寄主A组的鉴定结果,小麦孢囊线虫荥阳群体与须水群体为同一致病型,它们不同于国际上已命名的13个致病型,与国内已报道的5个燕麦孢囊线虫群体的致病型有较大差异,该致病型与澳大利亚致病型Ha13较为相近。根据致病型命名的原则,郑州地区小麦孢囊线虫应属于第三致病型组,命名为Ha43。
小麦孢囊线虫许昌群体致病型与国际上命名的13个致病型也不相同。小麦孢囊线虫许昌群体致病型与郑州的两个群体存在极大的差异,有5个大麦品种、2个燕麦品种和1个小麦品种对两个群体的反应是不同的。与国内已报道的致病型比较,许昌群体与北京房山和河南新乡群体对国际鉴别寄主的反应是一致的,而与河北定州群体、山西太谷和安徽固镇群体有一定差异。许昌群体致病型与瑞典报道的G(West)有些相近。小麦孢囊线虫郑州两个群体对A组鉴别寄主材料的致病谱大于许昌群体。
采用室内接种和田间病圃鉴定方法测定了黄淮麦区主推小麦品种对小麦孢囊线虫郑州荥阳群体和许昌群体的抗性。结果表明,46个主推小麦品种对荥阳群体的抗性总体较差,多数品种表现为感病(中感、感病或高感),缺乏免疫和高抗品种,只有太空6号、新麦18、中育6号和新麦11等4个品种对燕麦孢囊线虫表现出一定的抗性。对许昌群体的抗性总体表现也较差,室内接种鉴定没有发现抗病品种,全部表现为中感、感病或高感;田间鉴定表现抗病的有中育6号、太空6号、偃展4110、濮麦9号、豫农201、豫农949等6个品种。对来源于CIMMYT的66份小麦品种资源材料和41份小麦近缘材料对小麦孢囊线虫许昌群体或荥阳群体的抗性进行了测定,66份国外小麦品种多数表现感病,6R(6D)、MACKELLER、CP1133842等12个品种表现较好的抗性,具有很好的利用价值。同时,鉴定结果发现,小麦近缘材料中存在着丰富的抗禾谷孢囊线虫资源,三羊草属(Aegilops)、冰草属(Agropyron)、偃麦草属(Elytrigria)、簇毛麦属(Haynaldia)等中的一些材料对H. avenae和H. filipjevi均表现出较好的抗性,在抗禾谷孢囊线虫育种中可加以利用。
以小麦稳定表达的肌动蛋白基因(Actin)为对照,利用半定量RT-PCR技术对接种H.avenae后小麦不同抗性品种、不同时期、不同部位的几丁质酶基因表达进行了分析,结果表明:小麦受到禾谷孢囊线虫侵染后,能够诱导植株体内几丁质酶基因表达的上调,小麦抗、感品种之间几丁质酶基因表达量没有显著差异,说明几丁质酶活性变化不是小麦品种抗禾谷孢囊线虫的主要作用机制。
The cereal cyst nematode (CCN) occurred and distributed widely in the major wheat growing regions which brought a potential menace on the wheat production in China. Henan province was the main wheat production region and its cultivated area and total output of wheat ranked first in China. In recent years, CCN has caused serious yield losses in Zhengzhou and Xuchang, Henan province. In order to provide basis for the disease control, species and pathotypes of CCN from both areas and the resistance of wheat and its relatives to CCN were identified, the disease-resistant mechanism of wheat to Heterodera avenae were preliminarily studied .The main research results presented as follows.
The species of cereal cyst nematode from Zhengzhou (Xingyang population and Xushui population) and Xuchang (Xuchang population) were definited based on morphological and molecular methods. Xingyang population and Xushui population belonged to Heterodera avenae Wollenweber, 1924and Xuchang population belonged to H. filipjevi( ( Madzhidov,1981 )Stelter,1984. The mature cysts of both populations from Zhengzhou were lemon shaped, with colors from brown to dark brown, long neck, vulval cone, bifenestrate, vulval bridge. Cysts of Zhengzhou population didn′t have underbridge and their vulval cone had many bullae, yet the cysts of Xuchang population had underbridge, their vulval cone had less bullae. Second-stage juveniles of three populations were worm-shaped, with stylet of anchor-shaped basal knobs, ovoid oesophageal bulb, conical tail, hyaline tail.
Amplification of the rDNA-ITS region yielded a single PCR fragment of approximately 1000bp for each of the 3 populations. Eight kinds of restriction enzymes, TaqⅠ、AluⅠ、HaeⅢ、HinfⅠ、RsaⅠ、PstⅠ、CfoⅠ、Tru9Ⅰwere used to digest production of rDNA–ITS–PCR. The result showed restriction patterns of Xingyang population and Xushui population were the same, their ITS type were type B.The restriction pattern of Xuchang population was the same as the roports before. Zhengzhou cereal cyst nematode could be separated from Xuchang cereal cyst nematode using restriction enzymes Taq I and Pst I.
Cluster analysis for rDNA-ITS sequence, the populations of Xingyang and Xushui closely related to the herrings and native to the H. avenae populations from China and H.australis from Australia. The populations of Xuchang was clustered with high bootstrap supported with all H. filipjevi reported from foreign countries.
The pathotypes of three populations of cereal cyst nematodes from Zhengzhou and Xuchang were identified using the International Host Differentials (IHD). According to the reactions of IHD (group A) to the nematodes, the pathotypes of both populations from Zhengzhou were the same, but they were different from ones which had been nominated and reported. The pathotype of Zhengzhou populations were similar to Hal3, which was the dominant pathotype in Australia. Accoding to nomenclature, the two populations of Zhengzhou should belong to pathotype group 3, so the pathotype of Zhengzhou populations were named as Ha43.
The pathotype of Xuchang population was different from ones which had been nominated. It was different from the two populations of Zhengzhou. The resistant reaction of five barley, two oats and a wheat cultivars were different. Compared with other pathotypes in China, the pathotype of Xuchang population was the same as Fangshan population in Beijing and Xinxiang population in Henan, with little difference from Dingzhou population in Hebei, Taigu population in Shanxi and Guzhen population in Anhui. The pathogenicity of Zhengzhou populations was stronger than Xuchang population to IHD (group A).
Resistance of main released wheat cultivars in Huanghuai wheat area was tested to H.avenae and H. filipjevi by inoculation in greenhouse or in diseased field. The results indicated that the reactions of 46 cultivars were not good in general for Xingyang population, most of cultivars were susceptible (moderate susceptible, susceptible or high susceptible), immune and high resistant cultivars were lacking, Taikong 6, Xinmai18, Zhongyu6 and Xinmai11 were resistant to H.avenae. Resistance of 46 cultivars was not good in general for Xuchang population, also. There was no resistant cultivars in greenhouse identification and all cultivars were moderate susceptible, susceptible or high susceptible. Six cultivars were resistant to Xuchang population in the field testing, they were Zhongyu6, Zhongyu6, Yanzhan4110, Pumai9, Yunong201 and Yunong949.
The resistance of 66 wheat variety resources providing by CIMMYT and 41 relatives of wheat were tested to H. filipjevi of Xuchang population or H.avenae of Xingyang population. The results showed that 12 wheat cultivars were resistant to H. filipjevi of Xuchang population, for instance, 6R(6D), MACKELLER, CPI 133842, ect. There were abundant resistant resources in the relatives of wheat to cereal cyst nematode. Some materials in Aegilops, Agropyron, Elytrigria, Haynaldia were resistant to H.avenae and H. filipjevi, and they could be utilized in breeding for cereal cyst nematode disease resistance.
Taking wheat Actin gene as a control, expression of wheat chitinase gene 4(Chi4) were analyzed in the different growing periods, in different positions of different resistant wheat cultivars after inoculating H. avenae by semi-quantitative RT-PCR. The results showed that the expression of Chi4 was up-regulated when wheat plants were infected by H. avenae, no matter the cultivar was resistant or susceptible. There was no significant difference for Upregulation of Chi4 between resistant and susceptible cultivars. The change of Chi4 activities was not the principal mechanism for the resistance of wheat to cereal cyst nematode.
通过对郑州和许昌小麦孢囊线虫群体的形态鉴定和分子鉴定,明确了郑州和许昌地区小麦孢囊线虫的种类:郑州须水和荥阳两个群体都属于禾谷孢囊线虫组中的燕麦孢囊线虫(Heterodera avenae Wollenweber, 1924),许昌群体属于禾谷孢囊线虫组中的菲利普孢囊线虫〔H.filipjevi(Madzhidov,1981)Stelter,1984〕。小麦孢囊线虫郑州两个群体成熟孢囊柠檬形,褐色至黑褐色。颈较长,阴门锥显著。阴门膜孔为双膜孔,卵圆形,有阴门桥,但没有下桥。尾锥内不规则分布着大量褐色泡状突。二龄幼虫蠕虫形,口针粗壮,口针基部球锚形。中食道球卵圆形。尾圆锥形,有较长的透明尾。许昌群体成熟孢囊褐色,柠檬形,头部与尾部明显。头部有颈,尾部有尾锥。阴门膜孔为双膜孔,马蹄形。有阴门桥和下桥结构。尾锥内不规则分布着褐色泡状突,泡状突较郑州的两个群体少。二龄幼虫蠕虫形,口针粗壮,口针基部球锚形。中食道球卵圆形。尾圆锥形,有较长的透明尾。
扩增郑州和许昌3个小麦孢囊线虫群体核糖体DNA的ITS区,PCR产物都产生1条大小约为980bp的片段。用TaqⅠ、AluⅠ、HaeⅢ、HinfⅠ、RsaⅠ、PstⅠ、CfoⅠ、Tru9Ⅰ等8种限制性内切酶酶切3个群体的rDNA–ITS–PCR产物,结果表明,郑州须水和荥阳两个群体的酶切图谱相同,它们的ITS属于“B”型;许昌群体的酶切图谱与已报道H. filipjevi的酶切图谱一致;TaqⅠ酶与PstⅠ酶均可将许昌与郑州两地的小麦孢囊线虫区分开来。rDNA-ITS序列聚类分析结果表明,郑州须水和荥阳两个群体的序列与国内过去报道的H.avenae序列亲缘关系较近,与澳大利亚的H. australis序列也有较近的亲缘关系。许昌群体的序列与国外已报道的H. filipjevi序列聚在同一簇。
利用燕麦孢囊线虫致病型国际鉴别寄主对郑州荥阳、须水和许昌3个群体的致病型进行鉴定,结果表明,根据国际鉴别寄主A组的鉴定结果,小麦孢囊线虫荥阳群体与须水群体为同一致病型,它们不同于国际上已命名的13个致病型,与国内已报道的5个燕麦孢囊线虫群体的致病型有较大差异,该致病型与澳大利亚致病型Ha13较为相近。根据致病型命名的原则,郑州地区小麦孢囊线虫应属于第三致病型组,命名为Ha43。
小麦孢囊线虫许昌群体致病型与国际上命名的13个致病型也不相同。小麦孢囊线虫许昌群体致病型与郑州的两个群体存在极大的差异,有5个大麦品种、2个燕麦品种和1个小麦品种对两个群体的反应是不同的。与国内已报道的致病型比较,许昌群体与北京房山和河南新乡群体对国际鉴别寄主的反应是一致的,而与河北定州群体、山西太谷和安徽固镇群体有一定差异。许昌群体致病型与瑞典报道的G(West)有些相近。小麦孢囊线虫郑州两个群体对A组鉴别寄主材料的致病谱大于许昌群体。
采用室内接种和田间病圃鉴定方法测定了黄淮麦区主推小麦品种对小麦孢囊线虫郑州荥阳群体和许昌群体的抗性。结果表明,46个主推小麦品种对荥阳群体的抗性总体较差,多数品种表现为感病(中感、感病或高感),缺乏免疫和高抗品种,只有太空6号、新麦18、中育6号和新麦11等4个品种对燕麦孢囊线虫表现出一定的抗性。对许昌群体的抗性总体表现也较差,室内接种鉴定没有发现抗病品种,全部表现为中感、感病或高感;田间鉴定表现抗病的有中育6号、太空6号、偃展4110、濮麦9号、豫农201、豫农949等6个品种。对来源于CIMMYT的66份小麦品种资源材料和41份小麦近缘材料对小麦孢囊线虫许昌群体或荥阳群体的抗性进行了测定,66份国外小麦品种多数表现感病,6R(6D)、MACKELLER、CP1133842等12个品种表现较好的抗性,具有很好的利用价值。同时,鉴定结果发现,小麦近缘材料中存在着丰富的抗禾谷孢囊线虫资源,三羊草属(Aegilops)、冰草属(Agropyron)、偃麦草属(Elytrigria)、簇毛麦属(Haynaldia)等中的一些材料对H. avenae和H. filipjevi均表现出较好的抗性,在抗禾谷孢囊线虫育种中可加以利用。
以小麦稳定表达的肌动蛋白基因(Actin)为对照,利用半定量RT-PCR技术对接种H.avenae后小麦不同抗性品种、不同时期、不同部位的几丁质酶基因表达进行了分析,结果表明:小麦受到禾谷孢囊线虫侵染后,能够诱导植株体内几丁质酶基因表达的上调,小麦抗、感品种之间几丁质酶基因表达量没有显著差异,说明几丁质酶活性变化不是小麦品种抗禾谷孢囊线虫的主要作用机制。
The cereal cyst nematode (CCN) occurred and distributed widely in the major wheat growing regions which brought a potential menace on the wheat production in China. Henan province was the main wheat production region and its cultivated area and total output of wheat ranked first in China. In recent years, CCN has caused serious yield losses in Zhengzhou and Xuchang, Henan province. In order to provide basis for the disease control, species and pathotypes of CCN from both areas and the resistance of wheat and its relatives to CCN were identified, the disease-resistant mechanism of wheat to Heterodera avenae were preliminarily studied .The main research results presented as follows.
The species of cereal cyst nematode from Zhengzhou (Xingyang population and Xushui population) and Xuchang (Xuchang population) were definited based on morphological and molecular methods. Xingyang population and Xushui population belonged to Heterodera avenae Wollenweber, 1924and Xuchang population belonged to H. filipjevi( ( Madzhidov,1981 )Stelter,1984. The mature cysts of both populations from Zhengzhou were lemon shaped, with colors from brown to dark brown, long neck, vulval cone, bifenestrate, vulval bridge. Cysts of Zhengzhou population didn′t have underbridge and their vulval cone had many bullae, yet the cysts of Xuchang population had underbridge, their vulval cone had less bullae. Second-stage juveniles of three populations were worm-shaped, with stylet of anchor-shaped basal knobs, ovoid oesophageal bulb, conical tail, hyaline tail.
Amplification of the rDNA-ITS region yielded a single PCR fragment of approximately 1000bp for each of the 3 populations. Eight kinds of restriction enzymes, TaqⅠ、AluⅠ、HaeⅢ、HinfⅠ、RsaⅠ、PstⅠ、CfoⅠ、Tru9Ⅰwere used to digest production of rDNA–ITS–PCR. The result showed restriction patterns of Xingyang population and Xushui population were the same, their ITS type were type B.The restriction pattern of Xuchang population was the same as the roports before. Zhengzhou cereal cyst nematode could be separated from Xuchang cereal cyst nematode using restriction enzymes Taq I and Pst I.
Cluster analysis for rDNA-ITS sequence, the populations of Xingyang and Xushui closely related to the herrings and native to the H. avenae populations from China and H.australis from Australia. The populations of Xuchang was clustered with high bootstrap supported with all H. filipjevi reported from foreign countries.
The pathotypes of three populations of cereal cyst nematodes from Zhengzhou and Xuchang were identified using the International Host Differentials (IHD). According to the reactions of IHD (group A) to the nematodes, the pathotypes of both populations from Zhengzhou were the same, but they were different from ones which had been nominated and reported. The pathotype of Zhengzhou populations were similar to Hal3, which was the dominant pathotype in Australia. Accoding to nomenclature, the two populations of Zhengzhou should belong to pathotype group 3, so the pathotype of Zhengzhou populations were named as Ha43.
The pathotype of Xuchang population was different from ones which had been nominated. It was different from the two populations of Zhengzhou. The resistant reaction of five barley, two oats and a wheat cultivars were different. Compared with other pathotypes in China, the pathotype of Xuchang population was the same as Fangshan population in Beijing and Xinxiang population in Henan, with little difference from Dingzhou population in Hebei, Taigu population in Shanxi and Guzhen population in Anhui. The pathogenicity of Zhengzhou populations was stronger than Xuchang population to IHD (group A).
Resistance of main released wheat cultivars in Huanghuai wheat area was tested to H.avenae and H. filipjevi by inoculation in greenhouse or in diseased field. The results indicated that the reactions of 46 cultivars were not good in general for Xingyang population, most of cultivars were susceptible (moderate susceptible, susceptible or high susceptible), immune and high resistant cultivars were lacking, Taikong 6, Xinmai18, Zhongyu6 and Xinmai11 were resistant to H.avenae. Resistance of 46 cultivars was not good in general for Xuchang population, also. There was no resistant cultivars in greenhouse identification and all cultivars were moderate susceptible, susceptible or high susceptible. Six cultivars were resistant to Xuchang population in the field testing, they were Zhongyu6, Zhongyu6, Yanzhan4110, Pumai9, Yunong201 and Yunong949.
The resistance of 66 wheat variety resources providing by CIMMYT and 41 relatives of wheat were tested to H. filipjevi of Xuchang population or H.avenae of Xingyang population. The results showed that 12 wheat cultivars were resistant to H. filipjevi of Xuchang population, for instance, 6R(6D), MACKELLER, CPI 133842, ect. There were abundant resistant resources in the relatives of wheat to cereal cyst nematode. Some materials in Aegilops, Agropyron, Elytrigria, Haynaldia were resistant to H.avenae and H. filipjevi, and they could be utilized in breeding for cereal cyst nematode disease resistance.
Taking wheat Actin gene as a control, expression of wheat chitinase gene 4(Chi4) were analyzed in the different growing periods, in different positions of different resistant wheat cultivars after inoculating H. avenae by semi-quantitative RT-PCR. The results showed that the expression of Chi4 was up-regulated when wheat plants were infected by H. avenae, no matter the cultivar was resistant or susceptible. There was no significant difference for Upregulation of Chi4 between resistant and susceptible cultivars. The change of Chi4 activities was not the principal mechanism for the resistance of wheat to cereal cyst nematode.
引文
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