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河南省小麦主推品种对2种禾谷孢囊线虫的抗性及其评价方法

本站小编 Free考研考试/2021-12-26

邢小萍, 袁虹霞, 孙君伟, 张洁, 孙炳剑, 李洪连*
河南农业大学植物保护学院, 河南郑州 450002
* 通讯作者(Corresponding author): 李洪连, E-mail:honglianli@sina.com, Tel: 0371-63558791
第一作者联系方式: E-mail:xxp168@163.com
收稿日期:2014-01-12 基金:?本研究由国家公益性行业(农业)科研专项(200903040), 国家高技术研究发展计划(863计划)项目(2012AA101105), 国家“十二五”粮食丰产科技工程(2012BAD04B07)和中国科学院重点部署项目(CXJQ120111)资助;

摘要通过室内人工接种和田间自然病圃鉴定, 以单株白雌虫数法、相对抗病指数法和繁殖系数法评价河南省47个主推的小麦品种对燕麦孢囊线虫Heterodera avenae荥阳群体和菲利普孢囊线虫H. filipjevi许昌群体的抗性。室内人工接种条件下, 依据单株白雌虫数评价, 所有小麦品种对两种孢囊线虫均表现感病或高度感病; 但依据相对抗病指数评价, 有4个品种(太空6号、新麦11、中育6号和新麦18)对H. avenae表现抗病, 其余品种表现中度以上感病。田间病圃鉴定条件下, 依据单株白雌虫数评价, 太空6号和新麦18对H. avenae表现高抗, 中育6号、新麦11等10个品种表现中抗, 其余品种均表现为感病或高感; 中育6号和太空6号对H. filipjevi表现高抗, 偃展4110、濮麦9号和豫农201表现中抗, 其余品种均表现感病。依据相对抗病指数评价, 太空6号对H. avenae表现高抗, 新麦18、中育6号和新麦11表现抗病; 中育6号、太空6号对H. filipjevi表现高抗, 偃展4110、濮麦9号、豫农201、豫农949表现抗病。依据繁殖系数Pf/Pi评价, 太空6号、新麦11、中育6号和新麦18对H. avenae表现抗病, 太空6号、中育6号、濮麦9号和濮优938对H. filipjevi表现抗病, 其余品种表现感病。3种方法的抗性评价结果不完全相同, 相对抗病指数法与单株白雌虫数法的评价结果在一定程度上有一致性, 可部分克服因感病程度悬殊而导致评价不一致的问题, 因而可作为小麦品种抗禾谷孢囊线虫新的鉴定方法。

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
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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( P f /P i比; 播前孢囊(卵)数, Initial population density, P i; 收获后孢囊(卵)数, Final population density, P f)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 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
品种 Cultivar Heterodera avenae Heterodera filipjevi
平均单株白雌虫数 Mean of female per plant抗性 Resis- tance相对抗病指数 Relative resistance index抗性 Resis- tance平均单株白雌虫数 Mean of female per plant抗性 Resis- tance相对抗病指数 Relative resistance index抗性 Resis- tance
太空6号 Taikong 619.71±3.85MS0.73R27.86±3.52HS0.62MS
新麦11 Xinmai 1120.20±4.62HS0.73R60.00±4.51HS0.18HS
中育6号 Zhongyu 620.86±3.60HS0.72R31.00±4.24HS0.58MS
新麦18 Xinmai 1822.17±1.63HS0.70R47.17±4.77HS0.36S
新麦19 Xinmai 1923.40±4.18HS0.68MS60.00±4.48HS0.18HS
周麦20 Zhoumai 2024.40±4.14HS0.67MS46.60±3.91HS0.37S
新麦9-998 Xinmai 9-99824.60±2.91HS0.67MS50.60±1.17HS0.31S
郑麦98 Zhengmai 9825.00±3.44HS0.66MS46.57±3.60HS0.37S
新麦208 Xinmai 20825.71±3.28HS0.65MS51.33±2.79HS0.30S
周麦17 Zhoumai 1725.71±1.38HS0.65MS39.40±2.34HS0.46S
濮麦9号 Pumai 926.60±6.01HS0.64MS33.17±4.16HS0.55MS
豫农201 Yunong 20128.14±5.01HS0.62MS37.67±2.09HS0.49S
温麦4号 Wenmai 428.17±5.88HS0.62MS37.80±1.96HS0.49S
豫农202 Yunong 20229.17±2.82HS0.60MS41.57±5.34HS0.43S
郑麦366 Zhengmai 36629.67±3.78HS0.60MS43.40±1.94HS0.41S
豫麦49 Yumai 4929.80±3.02HS0.60MS38.00±5.59HS0.48S
兰考906 Lankao 90630.17±5.00HS0.59MS59.71±4.18HS0.19HS
豫农949 Yunong 94931.40±3.33HS0.57MS38.00±3.70HS0.48S
周麦19 Zhoumai 1931.83±3.93HS0.57MS37.50±0.87HS0.49S
新麦13 Xinmai 1332.00±3.36HS0.57MS38.43±3.17HS0.48S
豫麦47 Yumai 4733.25±3.75HS0.55MS43.71±4.98HS0.41S
开麦18 Kaimai 1833.40±4.73HS0.55MS39.17±4.32HS0.47S
豫麦49-198 Yumai 49-19833.60±3.22HS0.54MS50.67±4.81HS0.31S
阜麦936 Fumai 93635.00±4.23HS0.53MS65.60±3.08HS0.11HS
周麦16 Zhoumai 1635.40±1.63HS0.52MS50.20±1.59HS0.32S
偃展4110 Yanzhan 411036.00±5.47HS0.51MS40.33±2.85HS0.45S
淮麦16 Huaimai 1638.00±5.47HS0.49S41.50±3.54HS0.44S
濮优938 Pumai 93838.00±2.84HS0.49S32.43±1.23HS0.56MS
郑麦9023 Zhengmai 902338.75±2.69HS0.48S41.83±0.70HS0.43S
新原958 Xinyuan 95838.83±4.12HS0.47S46.50±5.23HS0.37S
豫麦70 Yumai 7040.40±5.20HS0.45S60.00±6.87HS0.18HS
衡观35 Hengguan 3540.67±3.84HS0.45S53.67±4.01HS0.27HS
陕麦253 Shaanmai 25341.40±5.42HS0.44S52.17±2.43HS0.29HS
豫麦70-36 Yumai 70-3643.67±4.31HS0.41S51.43±2.72HS0.30S
郑麦9405 Zhengmai 940546.17±5.57HS0.37S50.50±5.42HS0.31S
郑麦004 Zhengmai 00447.67±2.80HS0.35S47.43±4.50HS0.36S
豫麦60 Yumai 6050.67±6.83HS0.31S59.80±4.55HS0.19HS
漯麦4号 Luomai 450.75±11.16HS0.31S55.80±2.85HS0.24HS
济麦1号 Jimai 150.75±5.06HS0.31S54.83±2.96HS0.25HS
豫麦34 Yumai 3452.00±3.07HS0.29HS57.43±2.98HS0.22HS
周麦18 Zhoumai 1853.33±4.31HS0.28HS44.50±2.74HS0.40S
豫麦18 Yumai 1855.40±2.71HS0.25HS51.20±3.88HS0.30S
温麦18 Wenmai 1855.40±5.12HS0.25HS47.80±2.08HS0.35S
豫麦58 Yumai 5856.67±6.03HS0.23HS39.40±3.20HS0.46S
温麦19 Wenmai 1969.67±6.76HS0.06HS73.57±2.05HSHS
泛麦5号 Fanmai 573.50±12.78HS0HS56.17±0.91HS0.24HS
矮抗58 Aikang 5873.83±4.81HSHS42.75±2.29HS0.42S
HR: 高抗; MR: 中抗; R: 抗; 中感: MS: 中感; S: 感; HS: 高感。
HR: highly resistant; MR: moderately resistant; R: resistant; MS: moderately susceptible; S: susceptible; HS: highly susceptible.

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

供试小麦禾谷孢囊线虫为 H. avenae荥阳群体和 H. filipjevi许昌群体。麦收后, 分别从河南省荥阳市乔楼镇秋社村和许昌县河街乡黄庄村小麦孢囊线虫病发生较重的田块采集土样, 晾干后, 采用Fenwick漂浮法[ 35]分离孢囊, 挑出成熟饱满的孢囊, 用无菌水反复冲洗干净后放入4℃冰箱备用。经形态和分子鉴定, 两地点的孢囊线虫种类分别为 H. av e nae[ 36] H. filipjevi[ 11, 12]
1.2 抗性鉴定方法1.2.1 室内人工接种鉴定 将孢囊用0.5% NaClO表面消毒10 min, 并用无菌水冲洗数次, 装入盛有无菌水的离心管, 置4℃冰箱低温处理。10周后, 在15℃左右人工气候箱内孵化。将孵出的二龄幼虫(J2)配成200条 mL-1的悬浮液待用。参照袁虹霞等[ 34]的方法, 接种后在室内(温度范围控制在15~20℃)自然光照下生长60 d后调查统计根表面及PVC管(直径 3 cm, 高13 cm)内土壤中的白雌虫数, 计算平均单株白雌虫数。
1.2.2 田间病圃鉴定 病圃选在河南省荥阳市乔楼镇秋社村和许昌县河街乡半坡铺村多年发生小麦禾谷孢囊线虫病的重病地块。前茬作物均为玉米, 2010年10月中旬播种, 品种随机排列, 每品种重复3次, 每小区9 m2 (3 m × 3 m), 管理同大田。次年5月上旬灌浆期取样调查, 每小区随机选取5点, 每点取20个单株, 统计根系及根围土中的白雌虫数, 计算平均单株白雌虫数。
1.2.3 繁殖系数( P f/ P i比)测定 在播种前和收获时用直径为25 mm的土钻分别取小麦根围0~20 cm深的耕作层土壤, 每小区取10个样点, 混匀装入样品袋中。室内晾干后采用过筛法调查土样中的孢囊数, 每样品3次重复, 每个重复取100 g土样。同时, 从每个重复随机取10个孢囊, 压破后定容至10 mL, 取1 mL查卵数。分别计算各品种种植前( P i)和收获后( P f)土壤孢囊密度或卵密度的比值。
1.3 抗病性评价方法1.3.1 依据单株白雌虫数评价 免疫(I)、高抗(HR)、中抗(MR)、中感(MS)和高感(HS)的评价标准分别为0、0.1~5.0、5.1~10.0、10.1~20.0和>20.0个白雌虫/株[ 34]
1.3.2 依据相对抗病指数(relative resistance index, RRI)评价 以当次试验发病最重的品种为感病对照, 计算各品种的RRI (RRI = 1 - 所测品种平均单株白雌虫数/发病最重品种平均单株白雌虫数)。免疫(I)为 RRI = 1, 高抗(HR)为0.90≤RRI<1, 抗病(R)为0.70≤RRI<0.90, 中感(MS)为0.50≤RRI<0.70, 感病(S)为 0.30≤RRI<0.50, 高感(HS)为RRI<0.30[ 34]
1.3.3 依据繁殖系数( P f /P i)评价 参照Soriano 等[ 29]的方法, 当 P f /P i≤1时为抗病反应(R), P f /P i>1时为感病反应(S)。
1.4 统计分析采用SPSS16.0统计软件对数据进行平均数及标准误分析。

2 结果与分析2.1 室内人工接种抗性鉴定小麦品种对 H. avenae荥阳群体的抗性, 不同方法的评价结果差异较大(表1)。依据平均单株白雌虫数评价, 供试小麦品种均超过10.0个, 表现为感病。其中, 太空6号平均单株白雌虫数为19.71, 表现中感, 其余46个品种平均单株白雌虫数均超过20.0个, 表现为高感。依据RRI评价, 供试小麦品种中无免疫和高抗品种; 太空6号、新麦11、新麦18和中育6号4个品种的RRI≥0.70, 表现为抗病; 新麦19、周麦20等22个品种的RRI在0.50~0.69之间, 为中感; 淮麦16、濮优938等13个品种的RRI在0.30~0.49之间, 表现为感病; 豫麦34、周麦18等8个品种的RRI<0.3, 为高感。
两种评价方法对 H. filipjevi的抗性评价结果也有一定差异(表1)。依据平均单株白雌虫数评价, 所有小麦品种平均超过20.0个, 表现高感。但依据RRI评价, 太空6号、中育6号等4个品种表现中感(MS), RRI在0.55~0.62之间; 周麦19、豫农201等30个品种表现感病, RRI在0.30~0.49之间; 陕麦253、衡观35等13个品种表现高感, RRI小于0.30。
2.2 田间自然病圃抗性鉴定小麦品种对 H. avenae的抗性依据平均单株白雌虫数评价, 太空6号和新麦18分别为2.62个和4.66个, 表现高抗, 中育6号、新麦11等10个品种在5.65~9.76个之间, 表现中抗; 郑麦9405、濮优938等25个品种在15.02~19.71个之间, 表现中感; 温麦19和泛麦5号等10个品达到20.0个以上, 表现高感。RRI评价结果表明, 太空6号的RRI为0.91, 表现高抗; 新麦18、中育6号和新麦11介于0.74~0.83之间, 表现抗病(R); 开麦18、新麦19等20个品种介于0.50~0.69之间, 表现中感; 周麦17、豫麦58等12个品种在0.31~0.49之间, 表现感病; 矮抗58、陕麦253等11个品种小于0.30, 表现高感(表2)。
表2
Table 2
表2(Table 2)
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
品种 Cultivar Heterodera av e nae He t erodera filipjevi
平均单株白雌虫数 Mean of female per plant抗性 Resis- tance相对抗病指数 Relative resistance index抗性 Resis- tance平均单株白雌虫数 Mean of female per plant抗性 Resis- tance相对抗病指数 Relative resistance index抗性 Resis- tance
太空6号 Taikong 62.62±0.76HR0.91HR3.65±0.83HR0.91HR
新麦18 Xinmai 184.66±1.17HR0.83R31.23±1.86HS0.24HS
中育6号 Zhongyu 65.65±0.76MR0.80R2.09±0.58HR0.95HR
新麦11 Xinmai 117.12±1.32MR0.74R17.67±1.05MS0.57MS
开麦18 Kaimai 188.56±0.26MR0.69MS12.69±0.95MS0.68MS
新麦19 Xinmai 198.62±0.15MR0.69MS17.09±2.51MS0.58MS
温麦4号 Wenmai 48.77±0.48MR0.68MS20.84±3.02HS0.49S
新麦208 Xinmai 2088.83±0.54MR0.68MS22.03±0.63HS0.47S
豫农201 Yunong 2019.25±0.08MR0.66MS9.27±1.54MR0.77R
豫农949 Yunong 9499.53±0.44MR0.65MS11.28±0.91MS0.72R
濮麦9号 Pumai 99.60±0.04MR0.65MS7.53±1.54MR0.82R
豫麦47 Yumai 479.76±0.17MR0.65MS33.25±2.92HS0.19HS
郑麦9405 Zhengmai 940510.32±0.80MS0.63MS18.43±2.32MS0.55MS
濮优938 Puyou 93811.03±0.77MS0.60MS19.03±1.70MS0.54MS
偃展4110 Yanzhan 411011.63±0.88MS0.58MS7.36±0.22MR0.82R
周麦16 Zhoumai 1611.73±0.55MS0.58MS23.93±2.13HS0.42S
豫农202 Yunong 20211.93±1.08MS0.57MS15.04±4.34MS0.63MS
济麦1号 Jimai 112.22±0.22MS0.56MS32.10±5.53HS0.22HS
周麦20 Zhoumai 2012.30±0.23MS0.55MS21.48±6.09HS0.48S
郑麦366 Zhengmai 36612.31±0.12MS0.55MS17.21±1.84MS0.58MS
豫麦70 Yumai 7012.70±1.40MS0.54MS22.10±3.68HS0.46S
豫麦49-198 Yumai 49-19812.90±0.26MS0.53MS21.72±0.95HS0.47S
豫麦18 Yumai 1813.46±0.51MS0.51MS24.10±0.34HS0.42S
周麦19 Zhoumai 1913.81±0.46MS0.50MS17.18±2.46MS0.58MS
周麦17 Zhoumai 1714.08±3.13MS0.49S20.44±4.77HS0.50MS
豫麦58 Yumai 5814.71±1.91MS0.47S23.90±2.73HS0.42S
新麦9-998 Xinmai 9-99814.84±1.38MS0.46S27.52±1.55HS0.33S
新原958 Xinyuan 95815.02±0.13MS0.46S12.43±3.50MS0.69MS
温麦6号 Wenmai 615.14±5.43MS0.45S21.16±3.36HS0.48S
豫麦60 Yumai 6015.96±2.44MS0.42S34.86±3.61HS0.16HS
兰考906 Lankao 90616.66±0.86MS0.40S41.00±1.38HSHS
郑麦98 Zhengmai 9817.21±1.10MS0.38S17.10±0.81MS0.58MS
新麦13 Xinmai 1317.49±1.03MS0.37S21.52±1.49HS0.48S
阜麦936 Fumai 93617.81±1.08MS0.35S37.77±0.82HS0.09HS
郑麦9023 Zhengmai 902318.38±0.69MS0.33S27.63±3.40HS0.33S
郑麦004 Zhengmai 00419.08±7.34MS0.31S13.00±0.87MS0.64MS
矮抗58 Aikang 5819.71±0.66MS0.29HS11.87±3.05MS0.69MS
陕麦253 Shaanmai 25320.53±0.06HS0.26HS12.51±2.06MS0.69MS
周麦18 Zhoumai 1821.03±1.16HS0.24HS19.94±1.22MS0.51MS
豫麦70-36 Yumai 70-3622.25±0.06HS0.19HS35.30±2.33HS0.15HS
温麦18 Wenmai 1823.06±4.97HS0.16HS21.95±2.42HS0.47S
淮麦16 Huaimai 1623.10±0.61HS0.16HS20.29±2.87HS0.51MS
衡观35 Hengguan 3523.15±2.36HS0.16HS32.27±3.66HS0.22HS
漯麦4号 Luomai 423.27±1.45HS0.16HS39.69±1.78HS0.04HS
豫麦34 Yumai 3424.40±0.86HS0.12HS35.36±0.86HS0.12HS
泛麦5号 Fanmai 525.78±0.24HS0.07HS28.92±3.92HS0.30S
温麦19 Wenmai 1927.60±4.63HSHS32.77±1.93HS0.21HS
HR: 高抗; MR: 中抗; R: 抗; 中感: MS: 中感; S: 感; HS: 高感。
HR: highly resistant; MR: moderately resistant; R: resistant; MS: moderately susceptible; S: susceptible; HS: highly susceptible.

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

H. filipjevi许昌群体进行田间抗性鉴定, 采用平均单株白雌虫数评价, 中育6号和太空6号表现高抗, 其平均单株白雌虫数分别为2.09和3.65; 偃展4110、濮麦9号和豫农201表现中抗, 平均单株白雌虫数分别为7.36、7.53和9.27; 豫农949、矮抗58等15个品种表现中感, 平均单株白雌虫数在11.28~19.94之间; 兰考906、漯麦4号等27个品种为高感, 平均单株白雌虫数均超过20.0。RRI评价结果表明, 中育6号和太空6号表现高抗, RRI分别为0.95和0.91; 豫农949、豫农201等4个品种表现抗病, RRI分别在0.72~0.82之间; 矮抗58、新原958等16个品种表现中感(MS), RRI分别在0.50~0.64之间; 温麦4号、温麦6号等14个品种表现感病, RRI在0.30~0.49之间; 新麦18、济麦1号等11个品种高感, RRI均小于0.25 (表2)。
2.3 繁殖系数法( P f /P i)抗性评价 对 H. avenae荥阳群体, 太空6号、新麦11等5个品种收获后根围土壤孢囊密度与种植前土壤孢囊密度的 P f/ P i值小于1.0, 表现抗病, 其余42个品种表现感病(表3)。太空6号、新麦11等4个品种表现抗病, 其余43个品种表现感病(表3)。
表3
Table 3
表3(Table 3)
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 P f /P i ratio
品种 Cultivar每100 g土孢囊数 Number of cysts per 100 g soil每克土卵数 Number of eggs per gram of soil
P i P f P f /P i抗性Resistance P i P f P f /P i抗性Resistance
太空6号 Taikong 69.25±0.858.00±1.000.86R16.10±0.7014.56±3.090.90R
新麦11 Xinmai 117.75±0.487.00±0.410.90R13.18±0.2211.83±1.220.90R
中育6号 Zhongyu 68.00±0.717.50±0.870.94R13.76±1.6112.60±0.770.92R
豫农202 Yunong 2029.50±0.659.00±1.350.95R14.25±1.5015.93±4.231.12S
新麦18 Xinmai 189.00±0.589.00±1.411.00R16.29±0.3115.93±2.970.98R
豫麦18 Yumai 188.50±0.299.00±1.081.06S13.69±0.2617.01±1.541.24S
新麦19 Xinmai 196.75±1.117.25±1.701.07S9.86±2.8913.05±0.311.32S
温麦6号 Wenmai 68.00±0.419.50±3.861.19S14.08±0.1617.10±3.661.21S
濮优938 Puyou 9389.50±0.6511.50±0.651.21S14.25±0.3822.08±3.921.55S
开麦18 Kaimai 189.25±0.9511.50±0.651.24S15.82±1.3819.90±1.901.26S
兰考906 Lankao 9068.25±0.8510.75±1.381.30S14.27±0.9921.18±1.801.48S
豫农201 Yunong 2016.50±0.298.50±0.651.31S10.86±0.2014.45±0.341.33S
温麦4号 Wenmai 48.00±0.0010.50±0.351.31S13.38±0.8520.16±2.361.51S
济麦1号 Jimai 18.50±0.9611.25±2.291.32S14.11±0.9318.90±4.111.34S
豫麦49-198 Yumai 49-1988.50±0.9611.25±2.291.32S14.20±1.8819.35±4.051.36S
郑麦366 Zhengmai 3668.25±0.6311.00±1.001.33S12.38±0.8319.58±1.061.58S
周麦17 Zhoumai 178.75±0.4812.00±1.291.37S13.21±1.0522.32±1.481.69S
衡观35 Hengguan 358.50±0.2912.25±0.631.44S13.01±0.6023.77±0.491.83S
温麦18 Wenmai 189.00±0.4113.00±1.471.44S14.49±0.2723.53±1.691.62S
周麦19 Zhoumai 198.25±0.2512.00±1.081.45S12.79±0.8120.40±0.661.60S
豫农949 Yunong 9498.50±0.2912.50±3.841.47S12.50±0.6520.50±3.711.64S
周麦16 Zhoumai 168.25±0.6312.50±2.401.52S13.61±0.0124.00±2.401.76S
新原958 Xinyuan 9588.00±0.4112.25±1.381.53S10.80±1.6121.68±4.712.01S
周麦20 Zhoumai 207.50±0.5012.25±1.651.63S11.25±0.1121.68±2.131.93S
濮麦9号 Pumai 97.75±0.2512.75±1.931.65S11.80±0.0522.57±0.381.91S
淮麦16 Huaimai 169.00±1.0815.50±1.191.72S13.41±1.2929.30±2.052.18S
豫麦70-36 Yumai 70-3610.00±1.8317.25±2.931.73S15.90±3.6032.26±3.412.03S
新麦208 Xinmai 2088.25±0.4814.25±4.961.73S15.51±3.5128.50±1.721.84S
周麦18 Zhoumai 187.25±0.4813.00±1.871.79S11.67±1.5923.53±3.772.02S
泛麦5号 Fanmai 57.75±0.6314.25±1.031.84S13.02±3.9223.80±1.621.83S
新麦9-998 Xinmai 9-99810.25±1.0319.00±2.381.85S17.22±1.6436.10±4.642.10S
偃展4110 Yanzhan 41109.25±0.9517.25±2.321.86S16.37±1.1731.91±5.471.95S
郑麦98 Zhengmai 988.50±0.5016.25±2.501.91S14.62±2.1230.71±6.342.10S
郑麦9023 Zhengmai 90237.50±0.5015.00±3.242.00S12.38±1.0228.05±2.632.27S
矮抗58 Aikang 588.50±0.8718.00±4.062.12S15.13±3.0632.22±2.572.13S
豫麦60 Yumai 608.00±0.4117.50±2.662.19S13.92±2.7430.63±4.772.20S
豫麦58 Yumai 589.50±0.6523.75±4.232.50S16.44±2.8847.26±5.842.88S
豫麦70 Yumai 707.75±0.8519.75±2.592.55S12.09±1.2337.53±4.213.10S
漯麦4号 Luomai 48.00±0.9120.75±2.212.59S12.74±0.8642.54±0.403.34S
温麦19 Wenmai 198.00±0.4121.00±1.412.63S13.20±0.2440.11±2.313.04S
新麦13 Xinmai 138.50±0.2922.75±6.762.68S13.35±0.7743.00±3.923.22S
豫麦47 Yumai 478.00±0.8222.25±1.652.78S12.96±0.1644.06±3.643.40S
郑麦9405 Zhengmai 94057.25±0.7520.25±0.952.79S11.46±1.1841.31±4.093.61S
陕麦253 Shaanmai 2538.50±0.5024.24±1.302.85S13.69±1.4548.02±3.423.51S
郑麦004 Zhengmai 0048.00±0.0026.50±6.743.31S13.44±1.7655.92±2.784.16S
豫麦34 Yumai 347.50±0.5025.00±4.883.33S12.75±2.1148.75±2.953.82S
阜麦936 Fumai 9368.50±0.5031.50±7.273.71S13.60±2.2457.96±6.064.26S
P i: 播前孢囊(卵)密度; P f: 收获后孢囊(卵)密度。HR: 高抗; MR: 中抗; R: 抗; 中感: MS: 中感; S: 感; HS: 高感。
P i: initial population density; P f: final population density. HR: highly resistant; MR: moderately resistant; R: resistant; MS: moderately susceptible; S: susceptible; HS: highly susceptible.

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 P f /P i ratio

H. filipjevi许昌群体, 太空6号、中育6号等4个品种收获后根围土壤孢囊密度与种植前孢囊密度的 P f /P i值小于1.0, 表现抗病, 其余43个品种感病(表4)。
表4
Table 4
表4(Table 4)
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 P f/ P i ratio
品种 Cultivar每100 g土孢囊数 Number of cysts in 100 g soil每克土卵数 Number of eggs per gram of soil
P i P f P f/ P i抗性Resistance P i P f P f/ P i抗性Resistance
太空6号 Taikong 632.67±1.8614.67±1.450.45R50.31±0.9931.00±1.700.62R
中育6号 Zhongyu 629.67±1.4518.00±2.080.61R51.43±1.3732.28±1.980.63R
濮麦9号 Pumai 953.67±2.3347.00±2.080.88R86.23±1.1379.27±1.770.92R
濮优938 Puyou 93840.00±1.5337.67±2.850.94R77.33±5.0368.06±4.030.88R
新麦208 Xinmai 20819.67±0.3320.00±2.311.02S38.36±1.4639.30±2.071.02S
周麦16 Zhoumai 1653.33±2.9158.33±2.401.09S100.26±0.26121.33±2.261.21S
郑麦9023 Zhengmai 902346.67±4.3753.33±4.101.14S84.01±1.29109.86±4.561.31S
周麦19 Zhoumai 1926.00±1.1531.00±1.151.19S35.19±1.7152.91±0.371.50S
温麦18 Wenmai 1860.67±3.2876.33±6.441.26S125.39±0.91145.54±2.561.16S
周麦20 Zhoumai 2044.67±1.4556.33±2.191.26S86.96±0.04106.65±0.251.23S
温麦6 Wenmai 628.33±0.3336.33±0.331.28S47.59±0.6181.86±2.341.72S
偃展4110 Yanzhang 411026.00±1.7333.67±2.191.30S44.03±1.2770.48±1.581.60S
周麦17 Zhoumai 1758.33±3.3376.33±7.171.31S111.22±1.18148.59±2.791.34S
新麦18 Xinmai 1840.67±2.6053.67±5.611.32S69.55±4.2586.59±2.391.25S
豫农201 Yunong 20131.67±3.2843.33±5.361.37S49.41±2.7194.46±1.361.91S
新麦13 Xinmai 1340.00±3.4656.67±2.331.42S68.53±1.23108.81±4.391.59S
豫麦47 Yumai 4757.33±4.3383.33±14.241.45S118.86±6.06134.44±5.041.13S
郑麦98 Zhengmai 9844.00±7.5565.33±1.761.48S69.81±1.39125.43±1.971.80S
开麦18 Kaimai 1818.33±0.8827.67±3.671.51S26.27±1.0348.51±0.081.85S
郑麦004 Zhengmai 00450.33±5.3679.00±7.211.57S94.96±0.26165.90±3.101.75S
新麦11 Xinmai 1118.67±4.6730.00±1.531.61S30.12±1.0256.60±3.901.88S
豫麦58 Yumai 5840.67±2.3366.67±7.671.64S74.02±1.02118.67±4.231.60S
淮麦16 Huaimai 1636.67±1.8660.33±1.451.65S57.69±0.79108.19±2.111.88S
漯麦4号 Luomai 430.33±5.2451.00±6.661.68S50.95±1.4596.22±3.021.89S
矮抗58 Aikang 5823.33±0.8841.67±1.451.79S45.26±0.1670.84±3.841.57S
新原958 Xinyuan 95834.67±1.7662.33±5.241.80S58.25±4.55127.98±4.782.20S
济麦1号 Jimai 151.33±2.3392.33±1.761.80S85.55±1.15171.12±5.722.00S
豫农202 Yunong 20234.00±0.5862.00±3.061.82S69.36±1.94112.42±8.371.62S
郑麦9405 Zhengmai 940557.00±5.57105.00±2.651.84S84.47±1.57185.50±1.002.20S
温麦4号 Wenmai 431.00±1.7358.00±1.151.87S47.33±2.4798.99±1.392.09S
泛麦5号 Fanmai 539.33±3.3874.33±5.611.89S75.25±1.05141.23±4.071.88S
衡观35 Hengguan 3532.67±1.4564.67±4.261.98S46.61±2.89143.13±3.113.07S
陕麦253 Shaanmai 25333.67±2.0366.67±4.261.98S59.71±1.41118.01±6.891.98S
周麦18 Zhoumai 1836.00±2.8972.00±5.512.00S75.60±0.30134.88±4.881.78S
新麦9-998 Xinmai 9-99835.67±1.4572.00±8.622.02S71.58±3.48149.76±1.462.09S
郑麦366 Zhengmai 36640.33±2.9185.00±3.512.11S65.06±3.24164.33±4.072.53S
豫农949 Yunong 94940.67±4.9187.67±9.532.16S65.07±5.47175.93±2.892.70S
豫麦49-198 Yumai 49-19840.67±3.3890.00±12.772.21S58.56±5.86170.40±3.002.91S
兰考906 Lankao 90636.67±1.2084.00±6.932.29S66.74±1.36173.60±3.002.60S
豫麦60 Yumai 6050.67±3.84120.00±8.662.37S93.23±3.83259.20±2.832.78S
豫麦34 Yumai 3423.33±2.0355.67±3.282.39S37.02±2.58103.92±1.382.81S
豫麦18 Yumai 1820.67±1.2051.00±9.002.47S25.49±1.81101.32±3.803.97S
豫麦70 Yumai 7021.67±3.3854.00±4.362.49S32.94±2.84114.12±2.223.46S
豫麦70-36 Yumai 70-3640.00±2.52107.33±10.042.68S69.33±3.53231.83±3.383.34S
新麦19 Xinmai 1930.67±0.8885.00±3.212.77S63.49±3.49171.70±2.672.70S
阜麦936 Fumai 93643.00±18.68119.67±19.272.78S86.09±2.69233.75±3.352.72S
温麦19 Wenmai 1930.67±1.8697.67±12.253.18S49.89±1.69210.97±2.574.23S
P i: 播前孢囊(卵)密度; P f: 收获后孢囊(卵)密度。HR: 高抗; MR: 中抗; R: 抗; 中感: MS: 中感; S: 感; HS: 高感。
P i: initial population density; P f: final population density. HR: highly resistant; MR: moderately resistant; R: resistant; MS: moderately susceptible; S: susceptible; HS: highly susceptible.

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 P f/ P i ratio

2.4 不同评价方法比较在人工接种条件下, 依据单株白雌虫数评价, 小麦品种对 H. avenae荥阳群体均表现为感病, 其中中感品种占2.1%, 其余品种均为高感; 依据RRI评价, 抗病品种占8.5%, 其余品种感病(其中中感占46.8%, 感占27.7%, 高感占17.0%)。田间病圃鉴定中, 依据单株白雌虫数评价, 高抗品种占4.3%, 中抗品种占21.3%, 其余品种表现感病; 依据RRI评价, 抗性品种占8.5% (高抗占2.1%, 抗占6.4%)。采用繁殖系数法鉴定, 依据孢囊密度评价, 抗病品种占10.6%; 依据卵密度评价, 抗病品种占8.5% (图1-a)。
室内人工接种条件下, 小麦品种对 H. filipjevi许昌群体均表现感病, 其中依据单株白雌虫数评价, 参试品种均为高感(HS); 依据RRI评价, 参试品种均为感病(中感占8.5%, 感病占63.8%, 高感占27.7%)。田间病圃鉴定中, 依据单株白雌虫数评价, 高抗品种占4.3%, 中抗品种占6.4%, 其余品种表现感病(中感占31.9%, 高感占57.4%); 依据RRI评价, 抗性品种占12.8% (高抗占4.3%, 抗占8.5%)。采用繁殖系数法鉴定, 依据孢囊密度和卵密度评价, 抗病品种比例均为4.7%, 其余品种感病(图1-b)。

3 讨论采用不同的抗性鉴定方法评价河南省推广的小麦品种对两种小麦禾谷孢囊线虫的抗性, 发现供试的47个小麦品种中, 没有对两种小麦禾谷孢囊线虫完全免疫的品种, 仅有少数品种表现一定抗性, 多数品种表现感病, 这与以前相关研究报道的结果基本一致[ 22, 23, 24, 25, 26], 说明目前我国小麦主产区河南省推广种植的小麦品种对两种小麦禾谷孢囊线虫的抗性普遍较差。鉴于小麦禾谷孢囊线虫病已经成为河南省及我国主要小麦生产区的重要病害, 应进一步加强抗病品种的选育和现有品种的抗病鉴定与筛选工作, 为该病害的防控提供可利用的品种材料。由于在普通栽培小麦品种中对小麦禾谷孢囊线虫可有效利用的抗性资源比较有限, 进一步发掘新的抗性资源是十分必要的。王振跃等[ 22]、袁虹霞等[ 34]、张佳佳等[ 37]、Li等[ 38]、Nicol等[ 39]发现一些国外种质材料和小麦近缘属种材料对小麦禾谷孢囊线虫表现较好抗性, 在抗病品种选育中应充分考虑利用这些种质资源。
图1
Fig. 1
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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 P f /P i ratio (density of cysts in soil before growing and after harvest); F: Based on P f /P i 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 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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