小麦孢囊线虫侵染特性、群体鉴定及品种抗病性研究
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摘要
2008年~2010年对山东省滨州、济南等6市进行了小麦孢囊线虫的调查,采集土壤样品269份,同时采用五点法调查分析了典型感病地块线虫群体的水平、行间和垂直分布;通过对卵、二龄幼虫(J2)及阴门锥的永久玻片的光学显微镜观察、测量和显微拍照,对山东省滨洲博兴县的孢囊线虫群体进行详细描述和鉴定;为明确山东省气候条件下小麦孢囊线虫病侵染发育特点,于2009年10月至2010年6月对品种济麦22和泰农18,采用网袋法收集根系,淘洗过筛法分离土壤中孢囊,酸性品红-次氯酸钠法对根内线虫染色方法,研究了济麦22和泰农18两品种根内及根围土壤中孢囊线虫的发育进程及动态;同时利用孢囊指数方法鉴定31个小麦品种(系)对Heterodera avenae群体的抗性。研究结果如下:
1.山东省小麦孢囊线虫病的调查及感病地块线虫群体的分布
在调查的6地级市的11个县/区检测到小麦孢囊线虫(CCN)孢囊,检出率为36. 4%,孢囊密度差异很大,枣庄的山亭区和台儿庄区没有检测出CCN孢囊,泰安岱岳区大汶口镇和肥城湖屯镇为发生最严重的地区。小麦孢囊线虫在田间的水平分布是不均匀的,行距20 cm种植方式在两种植行中间和种植行根部土壤中孢囊群体密度较大,距种植行5 cm群体密度较小。小麦孢囊线虫最高群体密度发生在土壤的耕作层,以0~30 cm土层中线虫群体密度较大,30 cm以下土层中孢囊线虫分布较少;孢囊数随着土层深度的增加呈现递减趋势。
2.山东省滨州小麦孢囊线虫群体的形态学鉴定
滨州小麦孢囊线虫群体的形态学特征为:J2蠕虫状,热杀死后虫体直或略弯,虫体长532.14±20.89μm(479.64~577.34),体宽360.25±45.05μm(277.73~444.1)。口针长21.77±1.04μm(18.35~24.02),基部球明显。DGO(背食道腺开口道口针末端的距离)为5.19±1.78μm(2.42~8.56)。尾长64.75±4.07μm(54.62~73.14),尾部稍钝,尾部透明区为38.87±2.66μm(33.44~45.69),占尾部总长的60%。按de Man公式计算二龄幼虫的a、b和c值,分别为26.28±1.59μm(22.65~30.08),8.2±0.45μm(7.54~9.64)和7.42±0.36μm(6.72~8.69)。孢囊柠檬形,褐色至深褐色,颈较长,阴门锥显著,阴门膜孔为双膜孔型,阴门下桥不明显,有大量明显的泡状突,阴门裂较短,为8.77±1.49μm(6.97~11.62)。鉴定该种线虫为燕麦孢囊线虫Heterodera avenae Wollenweber,1924。
3.小麦生长期间孢囊线虫的发育及动态
在山东省气候条件下,2个供试冬小麦品种根内均有小麦孢囊线虫能侵入并且能够顺利发育。冬前没有二龄幼虫侵入小麦根系,在越冬后期仅有少量二龄幼虫(J2)侵入。在返青期之前根内线虫形态均处于二龄,随着温度的升高,返青期J2侵染显著增加;之后J2依次发育成J3、J4。小麦孢囊线虫在山东泰安一年发生一代,在小麦的生长后期线虫完成生活史。2个供试品种单位根鲜重的线虫数无显著差异。另外,在研究过程中发现小麦根内有大量短体线虫侵入,与孢囊线虫共同危害小麦。
4.小麦品种(系)对Heterodera avenae群体的抗性鉴定
供试31个小麦品种中无高抗(HR)和中抗(MR)品种,大多数品种表现高度感病(HS)。淄麦12和山农8355两品种的单株孢囊数为11~20个,属中度感病(MS)品种,仅占供试品种的6.5%;其他29个品种单株孢囊数大于20个,均为高度感病(HS)品种,占供试品种的93.6%。在接种量相同的条件下,小麦根系的生长状况与最终形成的孢囊数没有明显相关性。
Cereal cyst nematode (CCN) was investigated in Shandong province in 2008, 2009 and 2010, 269 samples were collected from wheat roots and rhizosphere soils in 6 regional cities. CCN was also investigated using five-point method in typical susceptible wheat field. Cysts were extracted from soil samples from the wheat field in Binzhou. Cysts, eggs and second-stage juveniles (J2) were photographed and examined. Morphological observation of the cyst, vulval cone, eggs and J2 are described in this paper. In order to learn information about the developmental process and dynamics of CCN (Heterodera avena) juveniles in the root, nematodes in wheat root (variety Jimai 22 and Tainong 18) were recorded with interval times in 2009 and 2010. The root systems were collected with mesh bag method and nematodes in the root were stained by the method of acid fuchsin-NaCIO. The cereal cyst nematodes (including different stage juveniles) were counted under stereo microscope. Meanwhile, resistance against Heterodera avena was identified in 31 wheat lines. The results were showed as follows:
1. Occurrence of cereal cyst nematodes in Shandong province and its distribution in susceptible wheat field.
CCN were found in 11 counties/districts under the 6 regional cities of Binzhou, Jinan, Dezhou, Heze, Taian and Zaozhuang in Shandong. Occurrence of CCN is 36. 4% in this study. the cyst was not detected in Shanting and Taierzhuang districts of Zaozhuang city. CCN occurrence showed significant different in different regions. There were higher cyst density in Dawenkou and Hutun town in Taian city than other investigated area. Horizontal distribution of CCN in susceptible wheat field was not uniformity; more cysts were found in the middle of inter-row and planting row than other sampling point. Meanwhile, The abundances of CCN in four soil layers (0~10, 10~20, 20~30, and 30~40 cm) were analyzed. The greatest densities of CCN often occurred at of 0~30 cm depths, and then in 30~40 cm soil. The total number of CCN decreased with increasing of soil depth.
2. Observations and morphometrics of cereal cyst nematode from Binzhou, Shandong:
Morphological observation of the cyst, vulval cone, eggs and J2 are described in this paper. The species was determined to be Heterodera avenae characterized by J2 with body length 532±21μm(480~577), body width of 20.3±1.2μm (18.3~24.6), style length of 21.8±1.0μm (18.4~24.0) with well-developed style knobs, dorsal oesophageal gland orifice 5.2±1.8μm(2.4~8.6) from base of stylet, tail 64.8±4.1μm(54.6~73.1), hyaline tail terminal 38.9±2.7μm(33.4~45.7) and cysts were lemon-shaped, brown to dark brown, with protruding longer neck and vulva. A pair of semifenestra on vulval cone vulval underbridge absent, many bullae present obviously, vulval silt 8.8±1.5μm (7.0~11.6).
3. Development and dynamics of cereal cyst nematode during wheat growing season
CCN (Heterodera avenae) could invade the roots of 2 tested winter cultivars and complete its development under Shandong province climatic conditions. J2 was not detected in the root of 2 tested variety after seedling emergence (autumn season), J2 can invade root in January 15 (WP), but arrested development before February. With the increase of soil temperature, J2 density significantly increased in green stage of wheat growth (GS). J3 and J4 were detected in setting stage (SS) and heading stage (HS). There was only one generation per year in Shandong and the nematode completed its life cycle in mature stage of wheat. There was no significant difference in the number of nematodes per g fresh root between 2 tested variety Jimai 22 and Tainong 18 (p>0.05). Furthermore, Pratylenchus spp. was also detected in wheat root and rhizosphere. The coordinated damage of Pratylenchus spp. with Heterodera avena on the winter wheat should be attention.
4. Identification of wheat variety resistance to Heterodera avenae
There were no resistance variety to Heterodera avenae among tested wheat variety (line)., most cultivars were high susceptable (HS). Only Zimai 12 and Shannong 8355 were moderately susceptible (MS), 11~20 cysts/root, 6.5% of total tested variety (line),. Other 29 variety (line) were all high susceptable (HS), more than 20 cysts/root, 93.6% of total tested variety (line). In the present study, there was no relative between growth character of wheat root and numbers of new cysts under the same inoculation number of J2 .
1.山东省小麦孢囊线虫病的调查及感病地块线虫群体的分布
在调查的6地级市的11个县/区检测到小麦孢囊线虫(CCN)孢囊,检出率为36. 4%,孢囊密度差异很大,枣庄的山亭区和台儿庄区没有检测出CCN孢囊,泰安岱岳区大汶口镇和肥城湖屯镇为发生最严重的地区。小麦孢囊线虫在田间的水平分布是不均匀的,行距20 cm种植方式在两种植行中间和种植行根部土壤中孢囊群体密度较大,距种植行5 cm群体密度较小。小麦孢囊线虫最高群体密度发生在土壤的耕作层,以0~30 cm土层中线虫群体密度较大,30 cm以下土层中孢囊线虫分布较少;孢囊数随着土层深度的增加呈现递减趋势。
2.山东省滨州小麦孢囊线虫群体的形态学鉴定
滨州小麦孢囊线虫群体的形态学特征为:J2蠕虫状,热杀死后虫体直或略弯,虫体长532.14±20.89μm(479.64~577.34),体宽360.25±45.05μm(277.73~444.1)。口针长21.77±1.04μm(18.35~24.02),基部球明显。DGO(背食道腺开口道口针末端的距离)为5.19±1.78μm(2.42~8.56)。尾长64.75±4.07μm(54.62~73.14),尾部稍钝,尾部透明区为38.87±2.66μm(33.44~45.69),占尾部总长的60%。按de Man公式计算二龄幼虫的a、b和c值,分别为26.28±1.59μm(22.65~30.08),8.2±0.45μm(7.54~9.64)和7.42±0.36μm(6.72~8.69)。孢囊柠檬形,褐色至深褐色,颈较长,阴门锥显著,阴门膜孔为双膜孔型,阴门下桥不明显,有大量明显的泡状突,阴门裂较短,为8.77±1.49μm(6.97~11.62)。鉴定该种线虫为燕麦孢囊线虫Heterodera avenae Wollenweber,1924。
3.小麦生长期间孢囊线虫的发育及动态
在山东省气候条件下,2个供试冬小麦品种根内均有小麦孢囊线虫能侵入并且能够顺利发育。冬前没有二龄幼虫侵入小麦根系,在越冬后期仅有少量二龄幼虫(J2)侵入。在返青期之前根内线虫形态均处于二龄,随着温度的升高,返青期J2侵染显著增加;之后J2依次发育成J3、J4。小麦孢囊线虫在山东泰安一年发生一代,在小麦的生长后期线虫完成生活史。2个供试品种单位根鲜重的线虫数无显著差异。另外,在研究过程中发现小麦根内有大量短体线虫侵入,与孢囊线虫共同危害小麦。
4.小麦品种(系)对Heterodera avenae群体的抗性鉴定
供试31个小麦品种中无高抗(HR)和中抗(MR)品种,大多数品种表现高度感病(HS)。淄麦12和山农8355两品种的单株孢囊数为11~20个,属中度感病(MS)品种,仅占供试品种的6.5%;其他29个品种单株孢囊数大于20个,均为高度感病(HS)品种,占供试品种的93.6%。在接种量相同的条件下,小麦根系的生长状况与最终形成的孢囊数没有明显相关性。
Cereal cyst nematode (CCN) was investigated in Shandong province in 2008, 2009 and 2010, 269 samples were collected from wheat roots and rhizosphere soils in 6 regional cities. CCN was also investigated using five-point method in typical susceptible wheat field. Cysts were extracted from soil samples from the wheat field in Binzhou. Cysts, eggs and second-stage juveniles (J2) were photographed and examined. Morphological observation of the cyst, vulval cone, eggs and J2 are described in this paper. In order to learn information about the developmental process and dynamics of CCN (Heterodera avena) juveniles in the root, nematodes in wheat root (variety Jimai 22 and Tainong 18) were recorded with interval times in 2009 and 2010. The root systems were collected with mesh bag method and nematodes in the root were stained by the method of acid fuchsin-NaCIO. The cereal cyst nematodes (including different stage juveniles) were counted under stereo microscope. Meanwhile, resistance against Heterodera avena was identified in 31 wheat lines. The results were showed as follows:
1. Occurrence of cereal cyst nematodes in Shandong province and its distribution in susceptible wheat field.
CCN were found in 11 counties/districts under the 6 regional cities of Binzhou, Jinan, Dezhou, Heze, Taian and Zaozhuang in Shandong. Occurrence of CCN is 36. 4% in this study. the cyst was not detected in Shanting and Taierzhuang districts of Zaozhuang city. CCN occurrence showed significant different in different regions. There were higher cyst density in Dawenkou and Hutun town in Taian city than other investigated area. Horizontal distribution of CCN in susceptible wheat field was not uniformity; more cysts were found in the middle of inter-row and planting row than other sampling point. Meanwhile, The abundances of CCN in four soil layers (0~10, 10~20, 20~30, and 30~40 cm) were analyzed. The greatest densities of CCN often occurred at of 0~30 cm depths, and then in 30~40 cm soil. The total number of CCN decreased with increasing of soil depth.
2. Observations and morphometrics of cereal cyst nematode from Binzhou, Shandong:
Morphological observation of the cyst, vulval cone, eggs and J2 are described in this paper. The species was determined to be Heterodera avenae characterized by J2 with body length 532±21μm(480~577), body width of 20.3±1.2μm (18.3~24.6), style length of 21.8±1.0μm (18.4~24.0) with well-developed style knobs, dorsal oesophageal gland orifice 5.2±1.8μm(2.4~8.6) from base of stylet, tail 64.8±4.1μm(54.6~73.1), hyaline tail terminal 38.9±2.7μm(33.4~45.7) and cysts were lemon-shaped, brown to dark brown, with protruding longer neck and vulva. A pair of semifenestra on vulval cone vulval underbridge absent, many bullae present obviously, vulval silt 8.8±1.5μm (7.0~11.6).
3. Development and dynamics of cereal cyst nematode during wheat growing season
CCN (Heterodera avenae) could invade the roots of 2 tested winter cultivars and complete its development under Shandong province climatic conditions. J2 was not detected in the root of 2 tested variety after seedling emergence (autumn season), J2 can invade root in January 15 (WP), but arrested development before February. With the increase of soil temperature, J2 density significantly increased in green stage of wheat growth (GS). J3 and J4 were detected in setting stage (SS) and heading stage (HS). There was only one generation per year in Shandong and the nematode completed its life cycle in mature stage of wheat. There was no significant difference in the number of nematodes per g fresh root between 2 tested variety Jimai 22 and Tainong 18 (p>0.05). Furthermore, Pratylenchus spp. was also detected in wheat root and rhizosphere. The coordinated damage of Pratylenchus spp. with Heterodera avena on the winter wheat should be attention.
4. Identification of wheat variety resistance to Heterodera avenae
There were no resistance variety to Heterodera avenae among tested wheat variety (line)., most cultivars were high susceptable (HS). Only Zimai 12 and Shannong 8355 were moderately susceptible (MS), 11~20 cysts/root, 6.5% of total tested variety (line),. Other 29 variety (line) were all high susceptable (HS), more than 20 cysts/root, 93.6% of total tested variety (line). In the present study, there was no relative between growth character of wheat root and numbers of new cysts under the same inoculation number of J2 .
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