应用根际促生细菌防治大豆胞囊线虫病的研究
摘要
本文针对大豆生产上危害严重的大豆胞囊线虫病筛选获得了可诱导大豆产生系统抗性的根际促生菌,并对其分类地位、摇瓶发酵条件、生防效果、定殖规律、作用机理等方面进行了系统的研究。研制了适宜生产的多功能PGPR制剂,并对其田间使用效果进行了反复论证,得出其可提高大豆产量,优化大豆品质,并可诱导大豆产生抗性。结果如下:
1.将大豆种子用1069株细菌菌悬液包衣后,施于辽宁省、黑龙江省大豆胞囊线虫重发田。35天后调查表明,有14株细菌包衣后的大豆对大豆胞囊线虫有明显的抑制作用。第二年重复试验验证了生防细菌的诱导抗性作用,并且我们结合田间试验和室内测定结果从中成功筛选出5株抗大豆胞囊线虫的大豆根际促生细菌,分别为Sneb207、Sneb353、Sneb364、Sneb382、Sneb482,其中以Sneb207、Sneb482效果最好。
2.在形态学和生理生化反应的基础上,通过分子生物学方法对上述根际促生菌进行分类鉴定,明确了筛选出的PGPR分类地位,共2属4种细菌。并首次报道了巨大芽孢杆菌对大豆胞囊线虫病的抑制作用。研究结果表明Sneb207、482均为巨大芽孢杆菌(Bacillus megaterium),Sneb353为枯草芽孢杆菌(Bacillus subtilis),Sneb364为阴沟肠杆菌(Enterobactet cloacae),Sneb382为成团肠杆菌(Enterobacter agglomerans)。
3.对根际促生菌Sneb207、482的剂型进行了初步研究。采用灭菌发酵液制成的种衣剂处理豆种,经辽宁省、黑龙江省三地多次田间使用效果证明对大豆有显著的促生防病作用。经PGPR制剂包衣后的大豆对大豆胞囊线虫胞囊的抑制率可达38.12%～82.13%,胞囊线虫发病越重地块其诱导抗性表现越强。同时PGPR制剂可使大豆生育期缩短,出现早熟现象。对大豆产量影响显著,单株荚数和粒数均比对照增加2～4倍,单株粒重增重200～380%,百粒重增重14～40%,植株鲜重增加。起到提高大豆产量并优化大豆品质的作用。
4.对Sneb207、482的培养条件进行了优化,获得了Sneb207、482的最佳配方。通过单因子试验和正交试验最终筛选出最佳碳氮源配方。菌株Sneb207的发酵配方为:蔗糖1.57%,牛肉膏0.897%,酵母浸膏0.842%。最佳摇瓶条件为:初始pH8.0、接种量为2%、装液量为40ml的100ml三角瓶在28℃、全光照振荡培养。菌株Sneb482的发酵配方为:蔗糖1.51%,牛肉膏0.836%,酵母浸膏0.68%。最佳摇瓶条件为:初始pH8.0、接种量为0.5%、装液量为30ml的100ml三角瓶在28℃、全光照振荡培养。
5.通过逐步提高药物浓度的方法,获得了抗红霉素300μg/ml的Sneb207标记菌株,利用标记菌株来研究菌株Sneb207在大豆根部的定殖和消长规律。结果表明菌株Sneb207可在大豆根际及根内定殖、繁殖。说明其不仅可在土壤中大量繁殖,还是一株内生细菌。菌株在根际及根内的消长动态测定表明,菌量在接种15d左右出现高峰后逐渐下降。大豆促生菌Sneb207对根际环境适应性的研究结果表明:Sneb207引入后能与寄主植物、环境、土壤微生物等因素建立其新的生态平衡,形成一个“综合的生防体系”,以提高生物防治的稳定性和防效。且Sneb207可与化学农药和肥料同时使用,因此生物与化学防治结合,更有效的防治大豆根部土传病害。
6.对Sneb207促进大豆生长的机理研究表明,Sneb207具有解钾、解磷、固氮等能力。Sneb207作为大豆根内生细菌,能够有效促进根瘤的形成,根瘤数明显增加且根瘤大,与大豆产生联合固氮作用;Sneb207可增加根系周围土壤中营养物质的可用性,分解土壤中难溶性的矿物,从而帮助植物吸收各种矿质元素:Sneb207对大豆根部生长和根部形态学有积极作用,另外Sneb207刺激质膜H~+-ATP酶活力的显著增强,提高其磷酸化水平。从而调节大豆许多重要的生理功能,促进其生长发育。
7.通过试验证明了Sneb207可诱导大豆产生抗线虫、真菌的广谱抗性。从Sneb207处理根部的冰冻切片发现了内部组织与对照相比有很大变化,抗病方面主要表现在Sneb207诱导后中柱鞘细胞加厚,结构更为紧密。大豆胞囊线虫的取食位点位于中柱鞘部位,其细胞越为紧密,线虫越不容易汲取养分,而无法维持其自身生长发育。因此即使线虫入侵到维管束组织内,也会有木质和酚类物质的大量沉积,细胞壁得到加厚,有效地阻止了线虫侵入,使其限制在木栓层和外皮层内。另外Sneb207处理大豆后还可以诱导大豆体内PPO、POD、PAL、SOD、CAT几种防御酶的活力提高。
PGPR induced soybean system resistance for biological control against soybean cyst nematode was screened and obtained.Taxonomy of PGPR,breeding of high-production strains and their fermentation conditions,effects of biological control,colonization of PGPR in soybean,action mechanism were studied systematically.The multifunctional preparation which increased soybean yield,optimized soybean qualty,induced soybean system resistance was manufactured.Results showed as follows:
1.Soybean seeds coated by 1069 bacteria strains were grown on farmland which Heterodera glycines occurred seriously in Liaoning and Heilongjiang province.After 35 days, 14 strains showed efficiency to control soybean cyst nematodes.System resistance induced by antagonistic rhizobacteria was validated by the retrial in the second year.Five PGPR induced soybean system resistance were screened from farm and greenhouse experiments.They were Sneb207,Sneb353,Sneb364,Sneb382,and Sneb482.Sneb207 and Sneb482 showed strong and stable resistance between them.
2.PGPR were classified and identified by the methods of classical morphological, biochemical and molecular biological taxonomy.The sequences of 16SrDNA gene were sequenced and analyzed.Results showed strains Sneb207,Sneb353,Sneb364,Sneb382,and Sneb482 belonged to Bacillus megaterium,Bacillus subtilis,Enterobacter cloacae, Enterobacter agglomerans and Bacillus megaterium,respectively.
3.The multifunctional preparation was manufactured.Soybean seeds coated by sterilizing fermentation filtrates 207 and 482 were grown on farmland which Heterodera glycines occurred seriously in Liaoning and Heilongjiang province.The results showed both prepatations promoted soybean seedlings growth significantly,and induced soybean system resistance efficiently.And the inhibition rate against soybean cyst of soybean which treated by preparation was between 38.12%~82.13%.In different district,soybean which treated by preparation showed stable resistance to SCN.As SCN occurred more seriously,the resistance became more efficient.Also the multifunctional preparation could make soybean prematurity, increase soybean yield,optimize soybean quality.Such as pods and seeds per plant increased 100～300%,weight of seeds per plant increased 200～380%,weight of 100-seeds increased 14～40%and plant flesh weight increased compering with blank from.
4.Breeding of high-production strains and their fermentation conditions were optimized. The fermentation conditions were investigated by single factor and designs of orthogonal experiments.And the results were as follows:the liquid medium of fermentation of strain Sneb207 constituded by sucrose 1.57%,beef extract 0.897%,yeast extract 0.842%.The fermentation conditions included the optimum initial pH8.0,2%of the inoculation amount, the volume of medium in shaking flasks about 40ml/100ml flask,rotate speed about 150r/min, all light cultivated at 28℃.And the liquid medium of fermentation of strain Sneb482 constituded by sucrose 1.51%,beef extract 0.836%,yeast extract 0.68%.The fermentation conditions included the optimum initial pH8.0,0.5%of the inoculation amount,the volume of medium in shaking flasks about 30ml/100ml flask,rotate speed about 150r/min,all light cultivated at 28℃.
5.Marked PGPR Sneb207,resistant to erythromycin at 300μg/ml by continuously screening on series concentration of erythromycin media was obtained.The marked strain would be re-isolated from rhizosphere soil and soybean root when it was inoculated in the root.The result showed that Sneb207 could colonize,survive and transfer in soil and soybean root.The population peaks of marked 207 strains in rhizosphere soil,corm and pseudostem of soybean were at 15d after inoculation.The strain population showed the trend“increased at the beginning and decreased afterward”after inoculation.The result of the environment adaptability of Sneb207 on soybean rhizosphere showed that a new ecological equilibrium was set up by Sneb207,soybean,environment and rhizosphere microorganism.They composed an integrated system to improve the biological effect and stability of soybean resistance.And PGPR 207 could be used together with mostly germicides and fertilizer.So it could eradicate soybean root diseases by biological and chemical control.
6.The mechanism how Sneb207 promoted soybean growth was studied.Sneb207 had the ability which fixed nitrogen and increased available P and K contents in rhizo-spheric soil. Sneb 207 as inner bacteria of soybean root could promote the root nodule forming,augment and enlargement.Sneb207 dissolved minerals,increased nutritious contents in rhizo-spheric soil.Sneb promoted soybean root growth,increased biomass,increased H~+-ATPase and decreased pH value in soybean root.So it adjusted soybean crucial physiological function and promoted soybean growth.
7.The root split system test proved soybean system resistance was induced by Sneb207. The mechanism of soybean system resistance was studied by section using a cryostat.The micro-biology of root inoculated by sneb207 was changed.It manifested pericycle cells thickening and the structure tightening.So it held nematodes back effictively,confining them in phellem layer or exodermis.And the activities of defence enzymes of soybean root were enhanced after Sneb207 induced.
1.将大豆种子用1069株细菌菌悬液包衣后,施于辽宁省、黑龙江省大豆胞囊线虫重发田。35天后调查表明,有14株细菌包衣后的大豆对大豆胞囊线虫有明显的抑制作用。第二年重复试验验证了生防细菌的诱导抗性作用,并且我们结合田间试验和室内测定结果从中成功筛选出5株抗大豆胞囊线虫的大豆根际促生细菌,分别为Sneb207、Sneb353、Sneb364、Sneb382、Sneb482,其中以Sneb207、Sneb482效果最好。
2.在形态学和生理生化反应的基础上,通过分子生物学方法对上述根际促生菌进行分类鉴定,明确了筛选出的PGPR分类地位,共2属4种细菌。并首次报道了巨大芽孢杆菌对大豆胞囊线虫病的抑制作用。研究结果表明Sneb207、482均为巨大芽孢杆菌(Bacillus megaterium),Sneb353为枯草芽孢杆菌(Bacillus subtilis),Sneb364为阴沟肠杆菌(Enterobactet cloacae),Sneb382为成团肠杆菌(Enterobacter agglomerans)。
3.对根际促生菌Sneb207、482的剂型进行了初步研究。采用灭菌发酵液制成的种衣剂处理豆种,经辽宁省、黑龙江省三地多次田间使用效果证明对大豆有显著的促生防病作用。经PGPR制剂包衣后的大豆对大豆胞囊线虫胞囊的抑制率可达38.12%～82.13%,胞囊线虫发病越重地块其诱导抗性表现越强。同时PGPR制剂可使大豆生育期缩短,出现早熟现象。对大豆产量影响显著,单株荚数和粒数均比对照增加2～4倍,单株粒重增重200～380%,百粒重增重14～40%,植株鲜重增加。起到提高大豆产量并优化大豆品质的作用。
4.对Sneb207、482的培养条件进行了优化,获得了Sneb207、482的最佳配方。通过单因子试验和正交试验最终筛选出最佳碳氮源配方。菌株Sneb207的发酵配方为:蔗糖1.57%,牛肉膏0.897%,酵母浸膏0.842%。最佳摇瓶条件为:初始pH8.0、接种量为2%、装液量为40ml的100ml三角瓶在28℃、全光照振荡培养。菌株Sneb482的发酵配方为:蔗糖1.51%,牛肉膏0.836%,酵母浸膏0.68%。最佳摇瓶条件为:初始pH8.0、接种量为0.5%、装液量为30ml的100ml三角瓶在28℃、全光照振荡培养。
5.通过逐步提高药物浓度的方法,获得了抗红霉素300μg/ml的Sneb207标记菌株,利用标记菌株来研究菌株Sneb207在大豆根部的定殖和消长规律。结果表明菌株Sneb207可在大豆根际及根内定殖、繁殖。说明其不仅可在土壤中大量繁殖,还是一株内生细菌。菌株在根际及根内的消长动态测定表明,菌量在接种15d左右出现高峰后逐渐下降。大豆促生菌Sneb207对根际环境适应性的研究结果表明:Sneb207引入后能与寄主植物、环境、土壤微生物等因素建立其新的生态平衡,形成一个“综合的生防体系”,以提高生物防治的稳定性和防效。且Sneb207可与化学农药和肥料同时使用,因此生物与化学防治结合,更有效的防治大豆根部土传病害。
6.对Sneb207促进大豆生长的机理研究表明,Sneb207具有解钾、解磷、固氮等能力。Sneb207作为大豆根内生细菌,能够有效促进根瘤的形成,根瘤数明显增加且根瘤大,与大豆产生联合固氮作用;Sneb207可增加根系周围土壤中营养物质的可用性,分解土壤中难溶性的矿物,从而帮助植物吸收各种矿质元素:Sneb207对大豆根部生长和根部形态学有积极作用,另外Sneb207刺激质膜H~+-ATP酶活力的显著增强,提高其磷酸化水平。从而调节大豆许多重要的生理功能,促进其生长发育。
7.通过试验证明了Sneb207可诱导大豆产生抗线虫、真菌的广谱抗性。从Sneb207处理根部的冰冻切片发现了内部组织与对照相比有很大变化,抗病方面主要表现在Sneb207诱导后中柱鞘细胞加厚,结构更为紧密。大豆胞囊线虫的取食位点位于中柱鞘部位,其细胞越为紧密,线虫越不容易汲取养分,而无法维持其自身生长发育。因此即使线虫入侵到维管束组织内,也会有木质和酚类物质的大量沉积,细胞壁得到加厚,有效地阻止了线虫侵入,使其限制在木栓层和外皮层内。另外Sneb207处理大豆后还可以诱导大豆体内PPO、POD、PAL、SOD、CAT几种防御酶的活力提高。
PGPR induced soybean system resistance for biological control against soybean cyst nematode was screened and obtained.Taxonomy of PGPR,breeding of high-production strains and their fermentation conditions,effects of biological control,colonization of PGPR in soybean,action mechanism were studied systematically.The multifunctional preparation which increased soybean yield,optimized soybean qualty,induced soybean system resistance was manufactured.Results showed as follows:
1.Soybean seeds coated by 1069 bacteria strains were grown on farmland which Heterodera glycines occurred seriously in Liaoning and Heilongjiang province.After 35 days, 14 strains showed efficiency to control soybean cyst nematodes.System resistance induced by antagonistic rhizobacteria was validated by the retrial in the second year.Five PGPR induced soybean system resistance were screened from farm and greenhouse experiments.They were Sneb207,Sneb353,Sneb364,Sneb382,and Sneb482.Sneb207 and Sneb482 showed strong and stable resistance between them.
2.PGPR were classified and identified by the methods of classical morphological, biochemical and molecular biological taxonomy.The sequences of 16SrDNA gene were sequenced and analyzed.Results showed strains Sneb207,Sneb353,Sneb364,Sneb382,and Sneb482 belonged to Bacillus megaterium,Bacillus subtilis,Enterobacter cloacae, Enterobacter agglomerans and Bacillus megaterium,respectively.
3.The multifunctional preparation was manufactured.Soybean seeds coated by sterilizing fermentation filtrates 207 and 482 were grown on farmland which Heterodera glycines occurred seriously in Liaoning and Heilongjiang province.The results showed both prepatations promoted soybean seedlings growth significantly,and induced soybean system resistance efficiently.And the inhibition rate against soybean cyst of soybean which treated by preparation was between 38.12%~82.13%.In different district,soybean which treated by preparation showed stable resistance to SCN.As SCN occurred more seriously,the resistance became more efficient.Also the multifunctional preparation could make soybean prematurity, increase soybean yield,optimize soybean quality.Such as pods and seeds per plant increased 100～300%,weight of seeds per plant increased 200～380%,weight of 100-seeds increased 14～40%and plant flesh weight increased compering with blank from.
4.Breeding of high-production strains and their fermentation conditions were optimized. The fermentation conditions were investigated by single factor and designs of orthogonal experiments.And the results were as follows:the liquid medium of fermentation of strain Sneb207 constituded by sucrose 1.57%,beef extract 0.897%,yeast extract 0.842%.The fermentation conditions included the optimum initial pH8.0,2%of the inoculation amount, the volume of medium in shaking flasks about 40ml/100ml flask,rotate speed about 150r/min, all light cultivated at 28℃.And the liquid medium of fermentation of strain Sneb482 constituded by sucrose 1.51%,beef extract 0.836%,yeast extract 0.68%.The fermentation conditions included the optimum initial pH8.0,0.5%of the inoculation amount,the volume of medium in shaking flasks about 30ml/100ml flask,rotate speed about 150r/min,all light cultivated at 28℃.
5.Marked PGPR Sneb207,resistant to erythromycin at 300μg/ml by continuously screening on series concentration of erythromycin media was obtained.The marked strain would be re-isolated from rhizosphere soil and soybean root when it was inoculated in the root.The result showed that Sneb207 could colonize,survive and transfer in soil and soybean root.The population peaks of marked 207 strains in rhizosphere soil,corm and pseudostem of soybean were at 15d after inoculation.The strain population showed the trend“increased at the beginning and decreased afterward”after inoculation.The result of the environment adaptability of Sneb207 on soybean rhizosphere showed that a new ecological equilibrium was set up by Sneb207,soybean,environment and rhizosphere microorganism.They composed an integrated system to improve the biological effect and stability of soybean resistance.And PGPR 207 could be used together with mostly germicides and fertilizer.So it could eradicate soybean root diseases by biological and chemical control.
6.The mechanism how Sneb207 promoted soybean growth was studied.Sneb207 had the ability which fixed nitrogen and increased available P and K contents in rhizo-spheric soil. Sneb 207 as inner bacteria of soybean root could promote the root nodule forming,augment and enlargement.Sneb207 dissolved minerals,increased nutritious contents in rhizo-spheric soil.Sneb promoted soybean root growth,increased biomass,increased H~+-ATPase and decreased pH value in soybean root.So it adjusted soybean crucial physiological function and promoted soybean growth.
7.The root split system test proved soybean system resistance was induced by Sneb207. The mechanism of soybean system resistance was studied by section using a cryostat.The micro-biology of root inoculated by sneb207 was changed.It manifested pericycle cells thickening and the structure tightening.So it held nematodes back effictively,confining them in phellem layer or exodermis.And the activities of defence enzymes of soybean root were enhanced after Sneb207 induced.
引文
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