大豆孢囊线虫的生物学特性及诱导抗性研究
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摘要
大豆孢囊线虫(Heterodera glycines Ichinohe)(SCN)是一类重要的植物内寄生线虫,目前已经扩散到全世界种植大豆的国家和地区。我国14个省市均有SCN发生的报道。SCN最主要的寄主是大豆,还可侵染赤豆,菜豆,绿豆,豌豆和条纹胡枝子等作物和一些野草。SCN寄生大豆根部,能影响根系的生长,减少大豆根瘤的数量,严重时造成大豆减产甚至绝产。本文对寄生大豆和烟草的SCN的生物学特性及诱导抗性进行了研究,主要研究进展如下：
1.对我国华东部分地区,包括安徽,江苏和上海等部分地区进行调查采样,从55份土壤样品中共分离并鉴定到14份SCN样品,SCN检出率为25.5%。在上海只获得了嘉定区一份样品,未检测到SCN。江苏省和安徽省的SCN检出率分别为26.7%和22.2%。其中,江苏省南通市的SCN检出率最高,为55.6%；而在昆山,大丰,盐城和泗洪等五个县市以及安徽的灵璧县均未检测到SCN；在所采集的6种作物样品中,只有大豆样品中检测到SCN。
2.对采自浙江杭州,山西太谷和黑龙江哈尔滨的SCN群体进行生理小种鉴定,太谷和哈尔滨SCN群体均为4号小种,而杭州SCN群体为3号小种,并明确了杭州SCN群体在杭州的春大豆生长季一年发生3代。
3.大豆孢囊线虫(SCN)和肾形肾状线虫(Rr)共同接种18天后,SCN的早期发育不受Rr的影响,并且对大豆的侵染能力比Rr强；接种线虫的处理均能降低大豆的根鲜重和株高：但是在接种30天后(dpi),单独接种Rr的大豆根鲜重和株鲜重均最低,而接种SCN的根鲜重比对照增加。南方根结线虫(Mi)和SCN均能抑制大豆的地上部生长,降低株鲜重和株高,但是对根系的生长影响不大,甚至能刺激根系生长；在30dpi,线虫数量随着接种量的增加而增多,但是接种量相同时,Mi的数量比SCN多。
4.对采自河南省许昌市襄县烟草根部的孢囊线虫(SCN-HN)进行形态学和分子学鉴定,确定其为大豆孢囊线虫；SCN-HN能侵染生理小种的鉴别寄主(Peking, Pickett, PI90763, PI88988和Lee71),甚至少数能发育到三龄幼虫(J3),但是不能完成发育。用SCN-HN接种烟草和大豆,在48dpi,SCN-HN在烟草根部形成孢囊,但只在大豆根部形成一个J3；而用寄生大豆的山西SCN群体(SCN-SX)接种烟草和大豆,12dpi在普通烟根部和30dpi在本氏烟根部均只形成一个J3。因此,大豆不是SCN-HN的寄主,烟草也不是SCN-SX的寄主,SCN-HN是SCN的一个新的致病型。
5.SCN-HN(?)(?)侵染供试植物辣椒,番茄和茄子,但是在辣椒,茄子和番茄品种茄门800上不能完成发育,而在番茄品种杭杂一号和浙粉701上能发育成雌虫和雄虫。番茄品种杭杂一号和浙粉701是SCN-HN的侵染力较差的寄主。
6.比较了寄生许昌烟草的SCN (SCN-HN)和南方根结线虫(Mi)对烟草的致病性差异。SCN-HN(?)中群密度低时,对烟草的生长无抑制作用,甚至能刺激烟草的生长,但种群密度高时,能降低烟草的株鲜重和根鲜重；而Mi对烟草生长的抑制作用比SCN-HN强,无论接种量高低,均降低株鲜重和根鲜重：混合接种与单独接种相同数量的线虫相比,对植物的危害减小,甚至有补偿因Mi造成危害的作用。随着接种量的增加,SCN-HN的雌虫数量虽然有所增加,但是当达到相当高的密度时,种群间的竞争变得更加激烈,反而抑制了种群的增长；然而Mi的种群一直处于增长状态：两者混合接种,接种量低时,Mi的种群增长比SCN快,但是接种量高时,SCN-HN会抑制Mi的种群增长。
7.水杨酸(SA),茉莉酸(JA),β-氨基丁酸(BABA),2,6-二氯异烟酸(1NA)和核黄素等五种化学诱导物处理大豆,均能一定程度上减少SCN的雌虫数量,但是只有BABA(?)INA的抑制率分别达到51%和53%。INA对大豆的生长具有毒害,能显著降低根鲜重,株鲜重和株高。BABA分别进行灌根、叶面喷雾和幼苗浸根处理,均能减少SCN的雌虫数量,其中灌根的效果最好。幼苗浸根导致植物矮小,灌根导致根鲜重下降,而叶面喷雾能降低株鲜重。
8.根据rDNA-ITS序列,用PAUP4.0软件对采自不同寄主的SCN群体进行系统发育分析,显示SCN与其它孢囊线虫种间存在一定的遗传差异,而在SCN种内有一定的稳定性。寄生大豆的SCN群体与寄生烟草的SCN群体存在一定的种内变异,但无特异性。通过rDNA-ITS序列不能鉴别SCN的致病型。
Soybean cyst nematodes (Heterodera glycines Ichinohe)(SCN) is an important endoparasitic nematodes, has spread to all the countries and regions that growing soybean nowadays. SCN have been reported in14provinces and municipalities in China. The most suitable host of SCN is soybean (Glycine max), some crops including adzuki bean (Vigna angularis), snap bean(Phaseolus vulgaris), mung bean (Phaseolus aureus), green pea(Pisum sativum) and common lespedeza(Lespedeza striata) to name a few, and some kinds of weeds are also hosts. Parasitizing on soybean roots, SCN can depress root growth, reduce the amount of soybean root nodules, and cause soybean yield reduction or even totally yield lost under favorable conditions. This study has focused on biological characteristics of SCN parasitizing on soybean and tobacco, and induced resistance to SCN, the main research achievements are as follows:
1. SCN were detected in14samples from55samples collected from Anhui, Jiangsu and Shanghai, the incidence was25.5%. SCN were not found in the only one sample from Jiading district in Shanghai. While the incidence of SCN in Jiangsu and Anhui Provience was26.7%and22.2%, respectively. The incidence was the highest in Nantong in Jiangsu, which was55.6%. However, SCN were not found in Kunshan, Dafeng, Yancheng and Sihong in Jiangsu, and Lingbi in Anhui. Among the samples collected from six crops, SCN only detected in soybean samples.
2. Races of three SCN populations from Hangzhou in Zhejiang, Taigu in Shanxi and Haerbin in Heilongjiang were determined. It showed that Shanxi and Heilongjiang populations were race4, which were the same with the reports before. Hangzhou population was race3, and it can propagate for three generations during the whole growing season of spring soybean in Hangzhou.
3. At18days post inoculation (dpi), competition between SCN and Rotylenchulus reniformis (Rr) showed that development of SCN in the early stage was not affected by Rr, and infectivity of SCN was stronger than Rr. Fresh root weight and shoot height were reduced by inoculation of nematodes. At30dpi, inoculation of Rr alone reduced freash root and shoot weight, while inoculation of SCN increased fresh root weight compared with control. Both Meloidogyne incognita (Mi) and SCN reduced fresh shoot weight and shoot height, stimulated root growth instead. Number of nematodes increased with the augment of inoculum amount, but at the same inoculum amount, the number of Mi was greater than SCN.
4. The cyst nematodes parasitizing on tobacco (SCN-HN) in Xiang county, Xuchang city in Henan province was identified as H. glycines by means of morphological and molecular indentification. SCN-HN can infect all of soybean differentals, Peking, Pickett, PI90763, PI88988and Lee71, used in race determination. Some even developed into third stage juveniles (J3), but full development never occured until40dpi. SCN-HN and SCN-SX revealed a significant virulence difference. SCN-HN successfully invaded soybean roots, minimal development (to J3only) occurred at48dpi, but adult female never formed. Inoculation of SCN-SX to tobacco got similar results. Nematode can infect both Nicotiana tobacum and N. benthamiana, but only one J3developed on N. tobacum at12dpi, and one J3developed on N. benthamiana at30dpi. These results indicated soybean is non-host of SCN-HN, and tobacco is non-host of SCN-SX. We suggest that SCN population parasitizing on tobacco is a new pathotype of SCN.
5. SCN-HN can infect all of Solanum melongena, Capsicum annuum and Lycopersicon esculentum studied at10dpi. However, adult female never formed on S. melongena (var. Hangjiaoyihao and Jizhuaxjilin), C. annuum (var. Hangzhouhongqie) and L. esculentum (var. Qiemen800) at33dpi, while female and male were developed on L. esculentum (var. Hangzayihao and Zhefen701). It seems that Hangzayihao and Zhefen701of L. esculentum are poor hosts of SCN-HN.
9. Pathogenicity was compared between SCN-HN and Mi that parasitizing on the same tobacco in Xuchang. Tobacco growth was not affected, even stimulated by SCN-HN if inoculation level was less than1000, but fresh shoot and root weight were reduced when the inoculation level was higher. The suppression by Mi was stronger than SCN-HN. Fresh shoot weight was reduced at any inoculation level of Mi. Compared with inoculated one species alone, inoculating two species together at the same inoculum level resulted in less damage to tobacco, even compensated the damage caused by Mi. Number of SCN was gradually increased as the inoculum augment, but decreased as a result of intence competetion at level10000. Mi increased gradually at all of the level. When inoculated together, Mi increased more quickly than SCN-HN at lower level, but was inhibited by SCN-HN at higher level.
7. Treatment of chemical elicitors salicylic acid (SA), jasmonic acid (JA) DL-β-amino butyric acid (BABA),2,6-dichloroisonicotinic acid (INA) and riboflavin to soybean, all reduced female number of SCN. BABA and INA were more effective, had suppression percentage of51%and53%, respectively. INA significantly reduced fresh root and shoot weight, and shoot height. BABA used as soil drench was more effective than used as foliar spray and root-drip. Root-drip decreased shoot height, soil drench caused reduction of fresh root weight, and foliar spray reduced fresh shoot weight.
8. Phylogenetic analysis was conducted based on rDNA-ITS sequence of SCN populations from soybean and tobacco using PAUP4.0. The phylogenetic tree showed some genetic variation among SCN and other cyst nematodes, while it has certain stability in species level of SCN. SCN populations parasitizing on soybean and tobacco showed some but not specific variation. However, the Maximum parsimony tree did not differentiate the two pathotypes distinctly.
1.对我国华东部分地区,包括安徽,江苏和上海等部分地区进行调查采样,从55份土壤样品中共分离并鉴定到14份SCN样品,SCN检出率为25.5%。在上海只获得了嘉定区一份样品,未检测到SCN。江苏省和安徽省的SCN检出率分别为26.7%和22.2%。其中,江苏省南通市的SCN检出率最高,为55.6%；而在昆山,大丰,盐城和泗洪等五个县市以及安徽的灵璧县均未检测到SCN；在所采集的6种作物样品中,只有大豆样品中检测到SCN。
2.对采自浙江杭州,山西太谷和黑龙江哈尔滨的SCN群体进行生理小种鉴定,太谷和哈尔滨SCN群体均为4号小种,而杭州SCN群体为3号小种,并明确了杭州SCN群体在杭州的春大豆生长季一年发生3代。
3.大豆孢囊线虫(SCN)和肾形肾状线虫(Rr)共同接种18天后,SCN的早期发育不受Rr的影响,并且对大豆的侵染能力比Rr强；接种线虫的处理均能降低大豆的根鲜重和株高：但是在接种30天后(dpi),单独接种Rr的大豆根鲜重和株鲜重均最低,而接种SCN的根鲜重比对照增加。南方根结线虫(Mi)和SCN均能抑制大豆的地上部生长,降低株鲜重和株高,但是对根系的生长影响不大,甚至能刺激根系生长；在30dpi,线虫数量随着接种量的增加而增多,但是接种量相同时,Mi的数量比SCN多。
4.对采自河南省许昌市襄县烟草根部的孢囊线虫(SCN-HN)进行形态学和分子学鉴定,确定其为大豆孢囊线虫；SCN-HN能侵染生理小种的鉴别寄主(Peking, Pickett, PI90763, PI88988和Lee71),甚至少数能发育到三龄幼虫(J3),但是不能完成发育。用SCN-HN接种烟草和大豆,在48dpi,SCN-HN在烟草根部形成孢囊,但只在大豆根部形成一个J3；而用寄生大豆的山西SCN群体(SCN-SX)接种烟草和大豆,12dpi在普通烟根部和30dpi在本氏烟根部均只形成一个J3。因此,大豆不是SCN-HN的寄主,烟草也不是SCN-SX的寄主,SCN-HN是SCN的一个新的致病型。
5.SCN-HN(?)(?)侵染供试植物辣椒,番茄和茄子,但是在辣椒,茄子和番茄品种茄门800上不能完成发育,而在番茄品种杭杂一号和浙粉701上能发育成雌虫和雄虫。番茄品种杭杂一号和浙粉701是SCN-HN的侵染力较差的寄主。
6.比较了寄生许昌烟草的SCN (SCN-HN)和南方根结线虫(Mi)对烟草的致病性差异。SCN-HN(?)中群密度低时,对烟草的生长无抑制作用,甚至能刺激烟草的生长,但种群密度高时,能降低烟草的株鲜重和根鲜重；而Mi对烟草生长的抑制作用比SCN-HN强,无论接种量高低,均降低株鲜重和根鲜重：混合接种与单独接种相同数量的线虫相比,对植物的危害减小,甚至有补偿因Mi造成危害的作用。随着接种量的增加,SCN-HN的雌虫数量虽然有所增加,但是当达到相当高的密度时,种群间的竞争变得更加激烈,反而抑制了种群的增长；然而Mi的种群一直处于增长状态：两者混合接种,接种量低时,Mi的种群增长比SCN快,但是接种量高时,SCN-HN会抑制Mi的种群增长。
7.水杨酸(SA),茉莉酸(JA),β-氨基丁酸(BABA),2,6-二氯异烟酸(1NA)和核黄素等五种化学诱导物处理大豆,均能一定程度上减少SCN的雌虫数量,但是只有BABA(?)INA的抑制率分别达到51%和53%。INA对大豆的生长具有毒害,能显著降低根鲜重,株鲜重和株高。BABA分别进行灌根、叶面喷雾和幼苗浸根处理,均能减少SCN的雌虫数量,其中灌根的效果最好。幼苗浸根导致植物矮小,灌根导致根鲜重下降,而叶面喷雾能降低株鲜重。
8.根据rDNA-ITS序列,用PAUP4.0软件对采自不同寄主的SCN群体进行系统发育分析,显示SCN与其它孢囊线虫种间存在一定的遗传差异,而在SCN种内有一定的稳定性。寄生大豆的SCN群体与寄生烟草的SCN群体存在一定的种内变异,但无特异性。通过rDNA-ITS序列不能鉴别SCN的致病型。
Soybean cyst nematodes (Heterodera glycines Ichinohe)(SCN) is an important endoparasitic nematodes, has spread to all the countries and regions that growing soybean nowadays. SCN have been reported in14provinces and municipalities in China. The most suitable host of SCN is soybean (Glycine max), some crops including adzuki bean (Vigna angularis), snap bean(Phaseolus vulgaris), mung bean (Phaseolus aureus), green pea(Pisum sativum) and common lespedeza(Lespedeza striata) to name a few, and some kinds of weeds are also hosts. Parasitizing on soybean roots, SCN can depress root growth, reduce the amount of soybean root nodules, and cause soybean yield reduction or even totally yield lost under favorable conditions. This study has focused on biological characteristics of SCN parasitizing on soybean and tobacco, and induced resistance to SCN, the main research achievements are as follows:
1. SCN were detected in14samples from55samples collected from Anhui, Jiangsu and Shanghai, the incidence was25.5%. SCN were not found in the only one sample from Jiading district in Shanghai. While the incidence of SCN in Jiangsu and Anhui Provience was26.7%and22.2%, respectively. The incidence was the highest in Nantong in Jiangsu, which was55.6%. However, SCN were not found in Kunshan, Dafeng, Yancheng and Sihong in Jiangsu, and Lingbi in Anhui. Among the samples collected from six crops, SCN only detected in soybean samples.
2. Races of three SCN populations from Hangzhou in Zhejiang, Taigu in Shanxi and Haerbin in Heilongjiang were determined. It showed that Shanxi and Heilongjiang populations were race4, which were the same with the reports before. Hangzhou population was race3, and it can propagate for three generations during the whole growing season of spring soybean in Hangzhou.
3. At18days post inoculation (dpi), competition between SCN and Rotylenchulus reniformis (Rr) showed that development of SCN in the early stage was not affected by Rr, and infectivity of SCN was stronger than Rr. Fresh root weight and shoot height were reduced by inoculation of nematodes. At30dpi, inoculation of Rr alone reduced freash root and shoot weight, while inoculation of SCN increased fresh root weight compared with control. Both Meloidogyne incognita (Mi) and SCN reduced fresh shoot weight and shoot height, stimulated root growth instead. Number of nematodes increased with the augment of inoculum amount, but at the same inoculum amount, the number of Mi was greater than SCN.
4. The cyst nematodes parasitizing on tobacco (SCN-HN) in Xiang county, Xuchang city in Henan province was identified as H. glycines by means of morphological and molecular indentification. SCN-HN can infect all of soybean differentals, Peking, Pickett, PI90763, PI88988and Lee71, used in race determination. Some even developed into third stage juveniles (J3), but full development never occured until40dpi. SCN-HN and SCN-SX revealed a significant virulence difference. SCN-HN successfully invaded soybean roots, minimal development (to J3only) occurred at48dpi, but adult female never formed. Inoculation of SCN-SX to tobacco got similar results. Nematode can infect both Nicotiana tobacum and N. benthamiana, but only one J3developed on N. tobacum at12dpi, and one J3developed on N. benthamiana at30dpi. These results indicated soybean is non-host of SCN-HN, and tobacco is non-host of SCN-SX. We suggest that SCN population parasitizing on tobacco is a new pathotype of SCN.
5. SCN-HN can infect all of Solanum melongena, Capsicum annuum and Lycopersicon esculentum studied at10dpi. However, adult female never formed on S. melongena (var. Hangjiaoyihao and Jizhuaxjilin), C. annuum (var. Hangzhouhongqie) and L. esculentum (var. Qiemen800) at33dpi, while female and male were developed on L. esculentum (var. Hangzayihao and Zhefen701). It seems that Hangzayihao and Zhefen701of L. esculentum are poor hosts of SCN-HN.
9. Pathogenicity was compared between SCN-HN and Mi that parasitizing on the same tobacco in Xuchang. Tobacco growth was not affected, even stimulated by SCN-HN if inoculation level was less than1000, but fresh shoot and root weight were reduced when the inoculation level was higher. The suppression by Mi was stronger than SCN-HN. Fresh shoot weight was reduced at any inoculation level of Mi. Compared with inoculated one species alone, inoculating two species together at the same inoculum level resulted in less damage to tobacco, even compensated the damage caused by Mi. Number of SCN was gradually increased as the inoculum augment, but decreased as a result of intence competetion at level10000. Mi increased gradually at all of the level. When inoculated together, Mi increased more quickly than SCN-HN at lower level, but was inhibited by SCN-HN at higher level.
7. Treatment of chemical elicitors salicylic acid (SA), jasmonic acid (JA) DL-β-amino butyric acid (BABA),2,6-dichloroisonicotinic acid (INA) and riboflavin to soybean, all reduced female number of SCN. BABA and INA were more effective, had suppression percentage of51%and53%, respectively. INA significantly reduced fresh root and shoot weight, and shoot height. BABA used as soil drench was more effective than used as foliar spray and root-drip. Root-drip decreased shoot height, soil drench caused reduction of fresh root weight, and foliar spray reduced fresh shoot weight.
8. Phylogenetic analysis was conducted based on rDNA-ITS sequence of SCN populations from soybean and tobacco using PAUP4.0. The phylogenetic tree showed some genetic variation among SCN and other cyst nematodes, while it has certain stability in species level of SCN. SCN populations parasitizing on soybean and tobacco showed some but not specific variation. However, the Maximum parsimony tree did not differentiate the two pathotypes distinctly.
引文
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