关键词:禾谷孢囊线虫; 相对抗病指数; 繁殖系数(Pf /Pi比); 抗性评价方法 Resistance to Two Species of Cereal Cyst Nematode and Evaluation Methods in Major Wheat Cultivars from Henan Province, China XING Xiao-Ping, YUAN Hong-Xia, SUN Jun-Wei, ZHANG Jie, SUN Bing-Jian, LI Hong-Lian* College of Plant Protection, Henan Agricultural University, Zhengzhou 450002, China Fund: AbstractResistance to cereal cyst nematode (CCN) is an important breeding target in wheat. In this study, we evaluated the resistance to CCN Xingyang population (Heterodera avenae) and Xuchang population (H. filipjevi) in 47 wheat cultivars from Henan Province, China through greenhouse and field tests. The CCN resistance was evaluated by the methods of average number of white female per plant, relative resistance index (RRI) andPf/Pi ratio. All cultivars were susceptible toH. avenae andH. filipjevi evaluated by average number of white female per plant after artificial inoculation in greenhouse. However, Taikong 6, Xinmai 11, Zhongyu 6, and Xinmai 18 showed resistance toH. avenae based on RRI evaluation. In field test, when evaluated with average number of white female per plant, there were a few resistant cultivars, including Taikong 6 and Xinmai 18 with high resistance and 10 cultivars (such as Zhongyu 6 and Xinmai 11) with moderate resistance toH. avenae, and Zhongyu 6 and Taikong 6 with high resistance and three cultivars (Yanzhan 4110, Pumai 9, and Yunong 201) with moderate resistance toH. filipjevi. The result of RRI evaluation showed that Taikong 6 was highly resistant toH. avenae, and Xinmai 18, Zhongyu 6 and Xinmai 11 were resistant toH. avenae; Zhongyu 6 and Taikong 6 were highly resistant toH. filipjevi and 4 cultivars (Yanzhan 4110, Pumai 9, Yunong 201, and Yunong 949) were resistant toH. filipjevi. UsingPf/Piratio as the evaluation index, four cultivars (Taikong 6, Xinmai 11, Zhongyu 6, and Xinmai 18) were resistant toH. avenae, and four cultivars (Taikong 6, Zhongyu 6, Pumai 9, and Puyou 938) were resistant toH. filipjevi. The resistance evaluation based on RRI method was partially identical to that based on white female number per plant method, and RRI method has the advantage to reduce the great evaluation error among susceptible cultivars. This method can be considered in CCN resistance evaluation in wheat.
Keyword:Cereal cyst nematode; Relative resistance index; Pf /Pi ratio; Methods for resistance evaluation Show Figures Show Figures
禾谷孢囊线虫(cereal cyst nematode, CCN)是麦类作物上重要的病原线虫, 以燕麦孢囊线虫( Heterodera avenae)、菲利普孢囊线虫( H. filipjevi)和麦类孢囊线虫( H. latipons)分布普遍, 危害严重[ 1]。1874年在德国首先报道 H. avenae, 现已50多个国家被发现[ 2, 3]。CCN是一个复合种群, 由已确定的12个种和几个未确定的种组成[ 4], 其中我国于1989年在湖北省首次发现 H. avenae[ 5], 目前已在河南、河北、山东等16个省(市)发现有该种分布[ 6, 7, 8, 9, 10], H. filipjevi仅在河南省被报道[ 11, 12]。河南省是我国小麦主产区, 其种植面积和总产量分别占全国的23%和27%, 小麦CCN发生面积达117万公顷, 是受该病为害最严重的省份, 而且近年呈不断加重趋势[ 13, 14]。CCN侵染小麦后, 破坏根系的正常代谢和机能, 一般造成小麦减产15%~40%, 严重时高达50%以上[ 2, 6, 15, 16, 17]。 小麦禾谷孢囊线虫病主要靠化学药剂和农业措施防治[ 18]。虽然利用轮作倒茬、土地休闲等农艺措施可以有效降低土壤中的孢囊基数[ 15, 18, 19], 但我国小麦主产区难以实施长期轮作, 而化学防治又面临土壤污染和成本较高等问题[ 17]。因此, 抗病品种作为高效、持续和经济的防病措施, 被寄予厚望。在澳大利亚等国家, 利用抗病品种已经基本控制了该病的危害[ 20, 21]。在我国, 生产上推广种植的小麦品种普遍抗性较差。王振跃等[ 22]采用田间自然病圃法对一些小麦品种和种质材料的 H. avenae抗性进行了鉴定, 没有发现抗病或免疫品种, 仅有少数国外引进的种质材料发病较轻。郑经武等[ 23]用人工接种法对国内22个栽培小麦品种和澳大利亚的25份抗病材料对 H. avenae的抗性进行鉴定, 发现国内品种大部分感病, 而澳大利亚的材料多数表现抗病。刘炳良等[ 24]采用室内人工接种和田间病圃法鉴定了40个小麦品种对 H. avenae江苏沛县群体的抗性, 发现仅有少数几个品种表现抗病。赵洪海等[ 25]和刘静等[ 26]采用室内人工接种、田间自然病圃等方法鉴定了山东省主要小麦栽培品种对 H. avenae的抗性, 发现供试小麦品种多数感病。 小麦品种对CCN抗性的评价方法和评价标准报道不尽相同[ 22, 23, 27, 28]。多数研究采用室内人工接种条件下, 依据根系上的白雌虫数进行评价。Soriano等[ 29]采用繁殖系数法( Pf /Pi比; 播前孢囊(卵)数, Initial population density, Pi; 收获后孢囊(卵)数, Final population density, Pf)评价水稻对禾谷根结线虫的抗性, 但鉴定结果只有抗、感两个级别。利用试验中发病最重或某一高度感病的品种作为对照, 将其他被鉴定品种的发病程度与其比较, 计算出相对抗病指数, 并建立一套评价体系, 是一种植物品种抗病性鉴定的有效方法, 可以避免因为鉴定条件的变化造成的误差, 目前已在多种病害上成功应用[ 30, 31, 32, 33]。本课题组曾采用该方法评价了国外小麦品种资源对菲利普孢囊线虫的抗性, 取得了较好的结果[ 34]。 鉴于我国小麦主产区河南省当前主推小麦品种对 H. ave nae和 H. filipjevi的抗性还不十分明确, 本研究用不同抗性鉴定方法评价了河南省主推的47个小麦品种对2种小麦禾谷孢囊线虫的抗性, 旨在为生产上利用抗病品种防控该病提供依据。 1 材料与方法1.1 小麦品种和线虫群体河南省生产上大面积推广的47个冬小麦品种(表1)分别由国家小麦工程技术中心、河南农业大学小麦育种研究室、河南省农业科学院小麦研究中心等育种单位提供。 表1 Table 1 表1(Table 1)
表1 室内人工接种条件下47个小麦品种对 Heterodera avenae荥阳群体和 Heterodera filipjevi许昌群体的抗性鉴定 Table 1 Resistance of 47 wheat cultivars to Heterodera avenae Xingyang population and Heterodera filipjeviXuchang population by inoculation with the second-stage juveniles in greenhouse
表1 室内人工接种条件下47个小麦品种对 Heterodera avenae荥阳群体和 Heterodera filipjevi许昌群体的抗性鉴定 Table 1 Resistance of 47 wheat cultivars to Heterodera avenae Xingyang population and Heterodera filipjeviXuchang population by inoculation with the second-stage juveniles in greenhouse
表2 田间自然病圃条件下47个小麦品种对 Heterodera avenae荥阳群体和 Heterodera filipjevi许昌群体的抗性鉴定 Table 2 Field resistance of 47 wheat cultivars to Heterodera avenae Xingyang population and Heterodera filipjeviXuchang population
表2 田间自然病圃条件下47个小麦品种对 Heterodera avenae荥阳群体和 Heterodera filipjevi许昌群体的抗性鉴定 Table 2 Field resistance of 47 wheat cultivars to Heterodera avenae Xingyang population and Heterodera filipjeviXuchang population
表3 Heterodera avena 荥阳群体在47个小麦品种根围的繁殖系数及抗性评价 Table 3 Evaluation of 47 wheat cultivars resistant to Xingyang population of Heterodera avenae in wheat plant rhizosphere based on Pf /Pi ratio
表3 Heterodera avena 荥阳群体在47个小麦品种根围的繁殖系数及抗性评价 Table 3 Evaluation of 47 wheat cultivars resistant to Xingyang population of Heterodera avenae in wheat plant rhizosphere based on Pf /Pi ratio
表4 Heterodera filipjevi许昌群体在47个小麦品种根围的繁殖系数及抗性评价 Table 4 Evaluation of 47 wheat cultivars resistant to Xuchang population of Heterodera filipjevi in wheat plant rhizosphere based on Pf/ Pi ratio
表4 Heterodera filipjevi许昌群体在47个小麦品种根围的繁殖系数及抗性评价 Table 4 Evaluation of 47 wheat cultivars resistant to Xuchang population of Heterodera filipjevi in wheat plant rhizosphere based on Pf/ Pi ratio
图1 不同评价方法鉴定小麦品种对 Heterodera avenae荥阳群体(a)和 Heterodera fili p jevi许昌群体(b)各类抗性水平比例A: 室内人工接种单株孢囊数评价法; B: 室内人工接种RRI评价法; C: 田间病圃单株孢囊数评价法; D: 田间病圃相对病情指数法; E: 繁殖系数法(孢囊密度); F: 繁殖系数法(卵密度)。HR: 高抗; MR: 中抗; R: 抗; 中感: MS: 中感; S: 感; HS: 高感。Fig. 1 Percentages of various resistance degrees of wheat cultivars to Heterodera avenae Xingyang population (a) and He t e - rodera filipjevi Xuchang population (b) evaluated by different methodsA: Based on number of cyst per plant inoculated in greenhouse; B: Based on relative resistance index (RRI) in greenhouse; C: Based on cyst number per plant in field trail; D: Based on RRI in field trial; E: Based on Pf /Pi ratio (density of cysts in soil before growing and after harvest); F: Based on Pf /Pi ratio (density of eggs in soil before growing and after harvest). HR: highly resistant; MR: moderately resistant; R: resistant; MS: moderately susceptible; S: susceptible; HS: highly susceptible.
本研究发现, 尽管多数品种对两种小麦禾谷孢囊线虫表现感病, 但不同小麦品种之间对两种禾谷孢囊线虫的抗性差异较大, 单株平均孢囊数差别达到几倍甚至十多倍。在田间自然病圃条件下, 采用不同方法评价的结果表明, 太空6号和中育6号对 H. avenae和 H. filipjevi均表现较好的抗性, 新麦11、新麦18仅对 H. avenae表现抗病, 濮麦9号对 H. filipjevi表现抗病。在重病区可以考虑推广使用太空6号等抗性较好的品种。在以 H. avenae危害为主的地区, 还可以考虑推广应用新麦11、新麦18等品种; 以 H. filipjevi危害为主的地区则可以考虑推广应用濮麦9号等品种。温麦19、豫麦34等品种对两种禾谷孢囊线虫均表现高度感病, 在重病区不宜推广应用。鉴于各地的小麦禾谷孢囊线虫种类和致病型存在差异, 在筛选和利用抗病品种时还应根据当地的病原线虫种类和致病型进行抗性鉴定和评价。对抗性表现较好的品种, 应深入研究和开发利用其抗性基因。 本研究发现, 利用单株白雌虫数进行评价, 在室内人工接种和自然病圃条件下, 参试品种的抗性反应级别存在较大差异, 如对 H. avenae荥阳群体抗病和感病品种的比例分别为0∶47和12∶35; 但采用RRI进行评价, 室内人工接种和自然病圃的鉴定结果一致, 对 H. avenae荥阳群体抗感病品种的比例均为4∶43; 而且相对抗病指数法评价结果与繁殖系数法结果也比较吻合, 对 H. avenae荥阳群体抗感品种比例分别为4∶43和5∶42。室内人工接种条件下, 依据单株白雌虫数评价品种抗性时, 所有参试品种对两种小麦禾谷孢囊线虫均表现感病。而采用RRI评价时, 以当次实验中发病最重的品种为感病对照品种, 计算相对抗病指数, 这样就可以有效避免由于不同实验条件而造成的评价结果的差异。因此, 综合考虑各种抗性评价方法的利弊, 作者认为RRI能较准确地评价参试材料的抗性水平, 而且比较简便, 可以作为小麦品种对CCN抗性评价的一种有效方法。建议无论在室内接种鉴定或田间病圃鉴定中以单株孢囊数作为主要鉴定依据的同时, 可以考虑以其中最感病的品种作为对照品种, 采用相对抗病指数法评价, 以避免年度和地区间由于试验条件不同而造成较大的误差。 4 结论河南省主推的47个小麦品种对两种小麦禾谷孢囊线虫群体的抗性整体表现较差, 多数品种表现感病, 但品种之间抗性差异明显, 少数品种表现较好抗性, 在重病区有一定推广利用价值。利用不同的抗性鉴定和评价方法, 结果存在一定的差异。采用田间病圃鉴定和繁殖系数法评价, 太空6号、新麦18、中育6号、新麦11对 H. avenae抗病性较好, 太空6号、中育6号对 H. filipjevi抗性较好。相对抗病指数法(RRI)简便易行, 可以作为小麦品种对小麦禾谷孢囊线虫抗性鉴定的新方法。 The authors have declared that no competing interests exist. 作者已声明无竞争性利益关系。The authors have declared that no competing interests exist.
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