内生芽孢杆菌及抗菌物质的鉴定和对作物黄萎病的防治效果
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摘要
大丽轮枝菌(Verticillium dahliae Kleb.)在世界上很多地区造成多种作物,其中包括棉花(Gossypium hirsutum L.)和茄子(Solanum melongena L.),发生微管束萎蔫病害。由该病菌引起的黄萎病已成为严重危害我国棉花和茄子生产的制约因素之一。利用内生细菌作为生防菌来防治土传微管束病害具有独特的优势,是很有应用前景的防治措施。
我们从茄子茎组织中分离获得一株内生细菌Jaas edl,其菌体和无细胞滤液对大丽轮枝菌(Verticillium dahliae)都有很强的拮抗活性。根据形态特性、生理生化特性和16s rDNA序列系统发育分析,鉴定该菌株为枯草芽孢杆菌(Bacillius subtilis),其16s rDNA序列在GenBank上的登录号为EF178293。通过平板法测定,内生枯草芽孢杆菌Jaas ed1可产生挥发性抗菌物质、嗜铁素和蛋白酶,但不产生几丁质酶。对内生枯草芽孢杆菌Jaas ed1的胞外抗菌粗提物进行理化性质测定,结果表明其胞外抗菌粗提物的抗菌活性对热稳定,耐碱性不耐酸性,对蛋白酶K和胰蛋白酶都不敏感。抑菌谱测定表明该胞外抗菌粗提物对10种病原真菌和1种病原细菌都具有拮抗作用。在光学显微镜下观察,经该胞外抗菌粗提物处理的大丽轮枝菌部分菌丝细胞异常膨大,细胞膜破裂,细胞质外泄。进一步将内生枯草芽孢杆菌Jaas ed1的培养液离心后的上清液,通过硫酸铵沉淀、Sephedex G-25分子筛层析、Cellulose DEAE-52阴离子交换层析、FPLC300SB-C18反相层析,获得在Tricine-SDS-PAGE上显示为分子量远小于6.5KD的一条带的抗菌物质。再通过电喷雾四极杆飞行时间串联质谱(Q-TOF2)分析发现,该抗菌物质包含有两个分子量相差14Da脂肽类化合物系列,推测为3个Fengycin家族同系物和4个Surfactin-like compound同系物。进一步用Q-TOF2测序鉴定其中m/z718.38(双电荷离子)的物质为C-14fengycinA。
通过灌根接种内生枯草芽孢杆菌Jaas ed1的Rif突变菌株Jaas ed1R,发现菌株Jaased1R能够从根部进入植株,并能迅速向上转移到茎和叶中。我们推测该菌株能够定殖在植物的微管束组织中,从而快速扩散到整个植株。温室盆栽试验表明,内生枯草芽孢杆菌Jaas ed1的菌体细胞悬浮液和无细胞滤液都对茄子黄萎病有一定的防治效果,但菌体细胞悬浮液的防治效果显著高于无细胞滤液。因此,不但内生枯草芽孢杆菌Jaasedl产生的抗菌物质具有生防作用,而且细菌本身在植株中的定殖可能是其能够有效防治茄子黄萎病的一个重要因素。此外,无论接种内生枯草芽孢杆菌Jaas edl菌体细胞悬浮液,还是其无细胞滤液都未在茄子植株上观察到任何病害症状。将内生枯草芽孢杆菌Jaas ed1作为可用于田间防治茄子黄萎病的生防菌剂还需要进一步进行田间大面积应用试验。
另外,我们对原有的一株棉花黄萎病生防内生细菌Jaas cd(原名73a)进行了菌种鉴定和田间应用方式的改进。通过形态观察、生理生化鉴定、Biolog鉴定以及16SrDNA序列比对,表明菌株Jaas cd为多粘类芽孢杆菌(Paenibacillus polymyxa),其16S rDNA序列在GenBank上的登录号为AY942618。抑菌谱测定表明,内生多粘芽孢杆菌Jaas cd对12种病原真菌均有较强的抑制作用。比较内生多粘芽孢杆菌Jaas cd和土壤拮抗细菌JB52以及化学药剂黄腐酸绿源宝单独使用、混合使用对棉苗生长、棉花产量的影响以及对棉花黄萎病的防治效果,结果表明无论将内生多粘芽孢杆菌Jaas cd与拮抗细菌JB52混用还是与黄腐酸绿源宝混用都不如单独使用效果好。内生多粘芽孢杆菌Jaas cd的促生长作用最强,棉花苗期株高可增长29.9%,鲜重可增加45.2%,田间试验可增产11.5%,并且用内生多粘芽孢杆菌Jaas cd的菌液灌根对棉花黄萎病的防治效果也最好,在7月底和9月初的防效分别达到51.17%和49.90%。田间大区示范试验表明用内生多粘芽孢杆菌Jaas cd的发酵液在棉苗移栽前喷施苗床和移栽时灌根两种方式处理都能有效防治棉花黄萎病和提高棉花产量,尤其喷施苗床的田间操作更加简便易行。
Verticillium dahliae Kleb. can cause vascular wilt, a serious disease of many plant species, including eggplant(Solanum melongena L.) and cotton(Gossypium hirsutum L.)in many areas of the world. Verticillium wilt is one of the major constraints on cotton and eggplant production in China. Because of the special contributions of endophytic bacteria to biological control, it is attractive to use this type of bacteria as an alternative tool to control soilborne vascular wilt disease.
An endophytic bacterial strain, Jaas edl,was isolated from the interior of eggplant (Solanum melongena L.)stems. The strain Jaas ed1and its cell-free filtrate had strong antifungal activity against Verticillium dahliae. According to the morphological and physiological characteristics and phylogenetic analysis of the16s rDNA sequence, it was identified as Bacillius subtilis. The sequence data for the16S rDNA gene of the strain Jaas edl has been deposited in GenBank with the accession number EF178293. The assay results using plates showed that endophytic B. subtilis Jaas edl produced some volatiles which strongly inhibited the mycelial growth of V. dahliae. In addition, the strain also had the capacity of producing siderophores and protease, but it didn't produce chitinase. The characterization of crude extracellular antifungal metabolites indicated that the antifungal activity was thermostable, tolerant to alkalescence pH, untolerant to acid pH, unsensible to proteinase K and trypsin. The crude extracellular antifungal metabolites showed strong antifungal activity against10fungal pathogens and1bacterial pathogen. Observed through optics microscopy, the crude extracellular antifungal metabolites caused cell membrane of Verticillium dahlia swelled and couldn't retain integrity, and then the intracellular substances leaked out. By approaches of ammonium sulphate precipitation, sephedex G-25molecular sieve chromatography, Cellulose DEAE-52ion-exchange chromatography, FPLC 300SB-C18reversion phase chromatography, antifungal substances were able to be isolated and purified from the culture of endophytic B. subtilis Jaas edl. The purified antifungal substances only showed one band with far lower than6.5KDa molecular weight compared with standard protein on the Tricine-SDS-PAGE. The Q-TOF2mass spectrum of purified antifungal substances revealed that two clusters each containing several molecules with the difference of14Da. According to the molecular weight, we speculated1cluster compounds with the doubly charged ion at [M+H] m/z=718.38,725.40,732.40were homologous compound of Fengycin family, and the other with the doubly charged ion at [M+H] m/z=722.37,729.37,736.39,743.39were homologous compound of surfactin-like compound family. After sequenced the amino acid by Q-TOF2, the compound with the doubly charged ion at [M+H] m/z=718.38was identified as C-14fengycinA.
We used the Rif mutant strain of Jaas ed1R instead of strain Jaas edl to study endophytic colonization ability. We inoculated endophytic B. subtilis Jaas ed1R to the eggplant roots, the bacteria could enter the eggplants from the roots and traveled upward to the stems and leaves quickly. The results suggest that the strain prefers to colonize inside the vascular tissues, so it can move to the whole plant quickly. In greenhouse experiments, the strain cell suspension effectively controlled Verticillium wilt of eggplant, and its control efficiency was far more significant than that of the cell-free filtrate after inoculation of V. dahliae. Therefore, not only the antifungal substance produce by the endophytic B. subtilis Jaas edl had the biocontrol potential, but also that the colonization of the bacterial inside plants might be another major factor in their effectiveness in controlling Verticillium wilt. No disease appeared in any eggplant inoculated with endophytic B. subtilis Jaas edl cell suspension or its cell-free filtrate. Further studies are currently underway to test the practical of usage of strain Jaas edl working as a biocontrol agent against Verticillium wilt in a large scale field experiment.
In addition, the endophytic bacterial strain Jaas cd (original name73a) against cotton verticillium wilt was identified and its application method for using in field was improved. According to the morphologlcal, physiological characteristics, Biolog identification and phylogenetic analysis, strain Jaas cd was identified as Paenibacillus polymyxa. The sequence data for the16S rDNA gene of the endophytic P. polymyxa Jaas cd has been deposited in GenBank with the accession number AY942618. The strain showed the strong antifungal activity against12fungal pathogens including Verticillium dahliae Kleb, Fusarium oxysporum f. sp. vasinfectum, Fusarium moniliforme and Rhizoctonia solani. Effects of endophytic bacterial strain Jaas cd, antagonistic bacterial strain JB52and fulivic acid on growth,yield and Verticillium wilt of cotton were investigated through single and combine applications, respectively. The results showed that the endophytic P. polymyxa Jaas cd was the best among all treatments. The efficiency of combine application with the strain Jaas cd and strain JB52and combine application with the strain Jaas cd and fulvic acid were weaker than that single application with the strain Jaas cd. Height and fresh weight of individual cotton seedlings was increased up to29.9%,45.2%respectively by the application with the endophytic P. polymyxa Jaas cd in pots. Yield of cotton was increased up to11.5%and the control efficiency against Verticillium wilt of cotton was up to51.17%in the end of July and49.90%at the beginning of September by the application with the endophytic P. polymyxa Jaas cd in field. The results in the field trial experiments showed that the disease indexes of cotton Verticillium wilt were decreased and the yields of cotton were increased by spraying the culture liquid of the endophytic P. polymyxa Jaas cd to cotton seeding beds before transplantation or watering seedling roots after transplantation. Spraying the culture liquid of the endophytic P. polymyxa Jaas cd to the cotton seeding bed before transplantation was more convenient to operate than watering the seedling roots after transplantation.
我们从茄子茎组织中分离获得一株内生细菌Jaas edl,其菌体和无细胞滤液对大丽轮枝菌(Verticillium dahliae)都有很强的拮抗活性。根据形态特性、生理生化特性和16s rDNA序列系统发育分析,鉴定该菌株为枯草芽孢杆菌(Bacillius subtilis),其16s rDNA序列在GenBank上的登录号为EF178293。通过平板法测定,内生枯草芽孢杆菌Jaas ed1可产生挥发性抗菌物质、嗜铁素和蛋白酶,但不产生几丁质酶。对内生枯草芽孢杆菌Jaas ed1的胞外抗菌粗提物进行理化性质测定,结果表明其胞外抗菌粗提物的抗菌活性对热稳定,耐碱性不耐酸性,对蛋白酶K和胰蛋白酶都不敏感。抑菌谱测定表明该胞外抗菌粗提物对10种病原真菌和1种病原细菌都具有拮抗作用。在光学显微镜下观察,经该胞外抗菌粗提物处理的大丽轮枝菌部分菌丝细胞异常膨大,细胞膜破裂,细胞质外泄。进一步将内生枯草芽孢杆菌Jaas ed1的培养液离心后的上清液,通过硫酸铵沉淀、Sephedex G-25分子筛层析、Cellulose DEAE-52阴离子交换层析、FPLC300SB-C18反相层析,获得在Tricine-SDS-PAGE上显示为分子量远小于6.5KD的一条带的抗菌物质。再通过电喷雾四极杆飞行时间串联质谱(Q-TOF2)分析发现,该抗菌物质包含有两个分子量相差14Da脂肽类化合物系列,推测为3个Fengycin家族同系物和4个Surfactin-like compound同系物。进一步用Q-TOF2测序鉴定其中m/z718.38(双电荷离子)的物质为C-14fengycinA。
通过灌根接种内生枯草芽孢杆菌Jaas ed1的Rif突变菌株Jaas ed1R,发现菌株Jaased1R能够从根部进入植株,并能迅速向上转移到茎和叶中。我们推测该菌株能够定殖在植物的微管束组织中,从而快速扩散到整个植株。温室盆栽试验表明,内生枯草芽孢杆菌Jaas ed1的菌体细胞悬浮液和无细胞滤液都对茄子黄萎病有一定的防治效果,但菌体细胞悬浮液的防治效果显著高于无细胞滤液。因此,不但内生枯草芽孢杆菌Jaasedl产生的抗菌物质具有生防作用,而且细菌本身在植株中的定殖可能是其能够有效防治茄子黄萎病的一个重要因素。此外,无论接种内生枯草芽孢杆菌Jaas edl菌体细胞悬浮液,还是其无细胞滤液都未在茄子植株上观察到任何病害症状。将内生枯草芽孢杆菌Jaas ed1作为可用于田间防治茄子黄萎病的生防菌剂还需要进一步进行田间大面积应用试验。
另外,我们对原有的一株棉花黄萎病生防内生细菌Jaas cd(原名73a)进行了菌种鉴定和田间应用方式的改进。通过形态观察、生理生化鉴定、Biolog鉴定以及16SrDNA序列比对,表明菌株Jaas cd为多粘类芽孢杆菌(Paenibacillus polymyxa),其16S rDNA序列在GenBank上的登录号为AY942618。抑菌谱测定表明,内生多粘芽孢杆菌Jaas cd对12种病原真菌均有较强的抑制作用。比较内生多粘芽孢杆菌Jaas cd和土壤拮抗细菌JB52以及化学药剂黄腐酸绿源宝单独使用、混合使用对棉苗生长、棉花产量的影响以及对棉花黄萎病的防治效果,结果表明无论将内生多粘芽孢杆菌Jaas cd与拮抗细菌JB52混用还是与黄腐酸绿源宝混用都不如单独使用效果好。内生多粘芽孢杆菌Jaas cd的促生长作用最强,棉花苗期株高可增长29.9%,鲜重可增加45.2%,田间试验可增产11.5%,并且用内生多粘芽孢杆菌Jaas cd的菌液灌根对棉花黄萎病的防治效果也最好,在7月底和9月初的防效分别达到51.17%和49.90%。田间大区示范试验表明用内生多粘芽孢杆菌Jaas cd的发酵液在棉苗移栽前喷施苗床和移栽时灌根两种方式处理都能有效防治棉花黄萎病和提高棉花产量,尤其喷施苗床的田间操作更加简便易行。
Verticillium dahliae Kleb. can cause vascular wilt, a serious disease of many plant species, including eggplant(Solanum melongena L.) and cotton(Gossypium hirsutum L.)in many areas of the world. Verticillium wilt is one of the major constraints on cotton and eggplant production in China. Because of the special contributions of endophytic bacteria to biological control, it is attractive to use this type of bacteria as an alternative tool to control soilborne vascular wilt disease.
An endophytic bacterial strain, Jaas edl,was isolated from the interior of eggplant (Solanum melongena L.)stems. The strain Jaas ed1and its cell-free filtrate had strong antifungal activity against Verticillium dahliae. According to the morphological and physiological characteristics and phylogenetic analysis of the16s rDNA sequence, it was identified as Bacillius subtilis. The sequence data for the16S rDNA gene of the strain Jaas edl has been deposited in GenBank with the accession number EF178293. The assay results using plates showed that endophytic B. subtilis Jaas edl produced some volatiles which strongly inhibited the mycelial growth of V. dahliae. In addition, the strain also had the capacity of producing siderophores and protease, but it didn't produce chitinase. The characterization of crude extracellular antifungal metabolites indicated that the antifungal activity was thermostable, tolerant to alkalescence pH, untolerant to acid pH, unsensible to proteinase K and trypsin. The crude extracellular antifungal metabolites showed strong antifungal activity against10fungal pathogens and1bacterial pathogen. Observed through optics microscopy, the crude extracellular antifungal metabolites caused cell membrane of Verticillium dahlia swelled and couldn't retain integrity, and then the intracellular substances leaked out. By approaches of ammonium sulphate precipitation, sephedex G-25molecular sieve chromatography, Cellulose DEAE-52ion-exchange chromatography, FPLC 300SB-C18reversion phase chromatography, antifungal substances were able to be isolated and purified from the culture of endophytic B. subtilis Jaas edl. The purified antifungal substances only showed one band with far lower than6.5KDa molecular weight compared with standard protein on the Tricine-SDS-PAGE. The Q-TOF2mass spectrum of purified antifungal substances revealed that two clusters each containing several molecules with the difference of14Da. According to the molecular weight, we speculated1cluster compounds with the doubly charged ion at [M+H] m/z=718.38,725.40,732.40were homologous compound of Fengycin family, and the other with the doubly charged ion at [M+H] m/z=722.37,729.37,736.39,743.39were homologous compound of surfactin-like compound family. After sequenced the amino acid by Q-TOF2, the compound with the doubly charged ion at [M+H] m/z=718.38was identified as C-14fengycinA.
We used the Rif mutant strain of Jaas ed1R instead of strain Jaas edl to study endophytic colonization ability. We inoculated endophytic B. subtilis Jaas ed1R to the eggplant roots, the bacteria could enter the eggplants from the roots and traveled upward to the stems and leaves quickly. The results suggest that the strain prefers to colonize inside the vascular tissues, so it can move to the whole plant quickly. In greenhouse experiments, the strain cell suspension effectively controlled Verticillium wilt of eggplant, and its control efficiency was far more significant than that of the cell-free filtrate after inoculation of V. dahliae. Therefore, not only the antifungal substance produce by the endophytic B. subtilis Jaas edl had the biocontrol potential, but also that the colonization of the bacterial inside plants might be another major factor in their effectiveness in controlling Verticillium wilt. No disease appeared in any eggplant inoculated with endophytic B. subtilis Jaas edl cell suspension or its cell-free filtrate. Further studies are currently underway to test the practical of usage of strain Jaas edl working as a biocontrol agent against Verticillium wilt in a large scale field experiment.
In addition, the endophytic bacterial strain Jaas cd (original name73a) against cotton verticillium wilt was identified and its application method for using in field was improved. According to the morphologlcal, physiological characteristics, Biolog identification and phylogenetic analysis, strain Jaas cd was identified as Paenibacillus polymyxa. The sequence data for the16S rDNA gene of the endophytic P. polymyxa Jaas cd has been deposited in GenBank with the accession number AY942618. The strain showed the strong antifungal activity against12fungal pathogens including Verticillium dahliae Kleb, Fusarium oxysporum f. sp. vasinfectum, Fusarium moniliforme and Rhizoctonia solani. Effects of endophytic bacterial strain Jaas cd, antagonistic bacterial strain JB52and fulivic acid on growth,yield and Verticillium wilt of cotton were investigated through single and combine applications, respectively. The results showed that the endophytic P. polymyxa Jaas cd was the best among all treatments. The efficiency of combine application with the strain Jaas cd and strain JB52and combine application with the strain Jaas cd and fulvic acid were weaker than that single application with the strain Jaas cd. Height and fresh weight of individual cotton seedlings was increased up to29.9%,45.2%respectively by the application with the endophytic P. polymyxa Jaas cd in pots. Yield of cotton was increased up to11.5%and the control efficiency against Verticillium wilt of cotton was up to51.17%in the end of July and49.90%at the beginning of September by the application with the endophytic P. polymyxa Jaas cd in field. The results in the field trial experiments showed that the disease indexes of cotton Verticillium wilt were decreased and the yields of cotton were increased by spraying the culture liquid of the endophytic P. polymyxa Jaas cd to cotton seeding beds before transplantation or watering seedling roots after transplantation. Spraying the culture liquid of the endophytic P. polymyxa Jaas cd to the cotton seeding bed before transplantation was more convenient to operate than watering the seedling roots after transplantation.
引文
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