根际有益芽孢杆菌N11及SQR9与植物根系的互作研究
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摘要
土传枯萎病的防治是世界性难题,传统的农业防控措施只能起到局部的控制作用,抗病品种选育和化学防治效果有限,且后者会带来严重的环境问题。近年来,生物防控以其环保、有效等优点日益受到人们关注。
本文研究了由实验室筛选到的香蕉枯萎病拮抗菌N6(Bacillus amyloliquefaciens)或/和N11(Bacillus subtilis)制得的专用型生物有机肥及广谱型有机肥在盆栽实验中对土传香蕉枯萎病的防控效果,拮抗菌N11和广谱型拮抗菌SQR9(Bacillus amyloliquefaciens)在香蕉根际的定殖情况,拮抗菌生物膜形成的表型特征、定量方法和动态变化,三株拮抗菌对香蕉和黄瓜根系分泌物及其中各种有机酸成份的响应,及不同作物根系分泌物对SQR9基因表达影响的转录组分析。最后研究了cheA基因突变对SQR9生物膜形成能力的影响。获得了以下结果：
1.通过盆栽实验研究了由实验室筛选到的土传香蕉枯萎病拮抗菌N6(Bacillus amyloliquefaciens)或/和N11(Bacillus subtilis)制得的专用型生物有机肥(BIO2-4)在盆栽实验中对土传香蕉枯萎病的防控效果,并与实验室之前研发的由拮抗菌SQR21(Paenibacillus subtilis)和SQR-T37(Trichoderma harzianum)制得的生物有机肥(BIO1)的效果进行比较,发现生物有机肥可将香蕉植株发病率由93.3%降到33.3%以下,专用有机肥处理的植株发病率更低于16.7%。利用实时荧光定量PCR法对香蕉植株根际土及土体土中的尖孢镰刀菌进行了检测,结果表明处理的根际土总DNA的尖孢镰刀菌ITS基因拷贝数为1.59×105个g-1土,而施用BIO1可将其拷贝数下降到6×104个g-1土,香蕉专用有机肥BIO2-4(T3-5)可进一步将拷贝数抑制到1.5×104个g-1土以下。利用PCR-DGGE分析了各处理香蕉根际土壤的微生物群落结构,发现根际土壤的真菌群落变化不大,但专用型香蕉生物有机肥(BIO2-4)处理可以显著增加细菌种群的多样性。
2.试图通过电转化将持续表达绿色荧光蛋白(Green Fluorescent Protein, GFP)的大肠-枯草杆菌穿梭质粒pHAPII引入拮抗菌N11,成功得到了GFP标记的N11菌株(N11-gfp),其菌落和菌体在荧光显微镜下可发出明亮的绿色荧光。N11-gfp的生长速度和对香蕉枯萎病病原菌FOC的拮抗能力与原始菌株N11基本相同。随后通过水培、砂培和土培系统,利用激光共聚焦荧光显微镜(CLSM)和扫描电子显微镜(SEM)研究了N11-gfp菌株和实验室之前构建的SQR9-gfp在香蕉根系的定殖情况。发现三种系统中拮抗菌在香蕉根表定殖的情况相似,在接种后2-4天,拮抗菌即可在香蕉植株根表形成微菌落和生物膜,定殖主要发生在主根的伸长区和分生区,次生根,主根与侧根的交界处以及根毛区,而附着于根尖的拮抗菌数量很少。拮抗菌定量检测的结果表明接种4天后N11-gfp与SQR9-gfp在香蕉根表的密度保持在每克根105-106CFU,以及每克根际土106-107CFU。
3.通过体外实验研究了芽孢杆菌在细胞培养板中的生物膜形成情况,发现枯草芽孢杆菌168在LB与MSgg中形成的生物膜都较弱,而野生型的FZB42、N6、N11和SQR9都能形成较厚的生物膜,且在MSgg中形成的生物膜更厚、结构更复杂。荧光显微镜观察发现N11-gfp与SQR9-gfp在盖玻片和培养板中形成的生物膜具有一定的厚度和立体结构。CLSM观察也表明SQR9-gfp形成的生物膜具有一定的立体结构。传统的结晶紫染色法与震荡打碎法定量测定生物膜生物量的相关性较高,R2达到0.9313。实验中发现芽孢杆菌在细胞培养板中的成膜具有一定边缘效应。最后研究了4株野生型芽孢杆菌在LB与MSgg培养基中生物膜与悬浮细胞生物量的动态变化规律,发现各菌生物膜的生物量从24h开始增加,到60h时保持稳定,同时悬浮细胞开始增多,96h后生物膜逐渐消解。各菌在MSgg中形成的生物膜生物量都高于LB,FZB42与SQR9在两种培养基中形成的生物膜生物量都高于N6与N11。
4.从香蕉根际分离的拮抗菌N11和从黄瓜根际分离的拮抗菌SQR9在其原始宿主根表的定殖能力都显著强于另一种宿主根部。N6和N11对浓缩的香蕉和黄瓜根系分泌物均表现出强烈的趋化性,SQR9对两种分泌物的趋化反应则一般。香蕉分泌物可以显著诱导N6和N11生物膜的形成。利用HPLC分析从香蕉根系分泌物中鉴定出草酸、苹果酸和延胡索酸这3种主要的有机酸成分,在黄瓜根系分泌物中则鉴定出草酸、苹果酸和柠檬酸。N6和N11对苹果酸、柠檬酸和延胡索酸都表现出趋化性,SQR9只对苹果酸和柠檬酸有较强的趋化反应。延胡索酸可以显著增加N6、N11和SQR9形成生物膜的生物量,柠檬酸则都能显著诱导N6和SQR9生物膜的形成。体外定殖实验发现延胡索酸可以诱导N11-gfp在两种作物上的定殖,而苹果酸、柠檬酸和延胡索酸可促进SQR9-gfp在香蕉或／和黄瓜根际的定殖。
5.通过接合的方法将携带有在木糖诱导型启动子(PxylA)控制下表达的感受态因子ComK的穿梭质粒引入野生型芽孢杆菌SQR9,发现通过木糖诱导可实现外源质粒的转化,转化效率在103-104CFU·μg-1质粒。通过响应曲面法设计Box-Behnken实验对诱导转化的条件进行了优化,发现最佳条件为菌液OD600达到0.5时,用0.2%浓度的木糖诱导1h,该条件下转化质粒的效率可以接近104CFU·μg-1。将cheA基因的双同源臂突变片段,以pMD19-T为母体的自杀载体,及该质粒的多聚体通过诱导转化的方式引入9kz,经过PCR验证发现得到了正确的双交换的cheA突变体。利用传代培养删除了质粒pXKZ.突变体对趋化物柠檬酸的反应消失,生长速度与野生型相似,生物膜形成有滞后现象。透射电镜的观察发现突变体的鞭毛基本缺失,而野生型具周生鞭毛。推测对cheA的删除影响了下游sigD的转录和表达,从而影响了鞭毛的形成进而造成趋化性的消失和成膜的滞后,具体情况需要cheA和sigD的互补实验验证。
6.浓缩的黄瓜、玉米及香蕉的根系分泌物可显著促进SQR9在1/2浓度的MSgg中的生长(接种后6h)。玉米根系分泌物可诱导SQR9在1/2MSgg中的生物膜形成,利用Solexa高通量转录组测序研究了在振荡培养条件下三种作物分泌物,以及静态生物膜形成条件下玉米根系分泌物对SQR9基因表达的影响。结果表明主要调控的基因类群包括营养代谢(主要包括碳水化合物代谢、氨基酸代谢、脂类代谢和核酸代谢等)、蛋白质翻译与调控、跨膜运输(主要是ABC转运蛋白)、信号转导(主要为双组份调控系统)以及趋化及运动相关。不同根系分泌物调控的基因类群有一定差异。SQR9在静态条件下对玉米根系分泌物的转录组在接种后24h和48h有较大差异。
The control of Fusarium wilt is a world wide problem. The traditionally farming control methods have rarely provided long-term control in any production area, while selection of resistant cultivars and chemical application are not effective enough for control this disease, and also the latter causes environmental problems. Recently, biological control of soil-born disease is an efficient and environmentally friendly way and is receiving increased attention.
In this study, the capability of the specific bio-organic fertilizers made of organic fertilizer (pig manure compost and amino acid fertilizer) fermented with antagonisms Bacillus amyloliquefaciens N6or/and Bacillus subtilis N11, as well as the normal BIO developed before in this laboratory, to control the Fusarium wilt of banana was evaluated in pot experiments. The colonization pattern of GFP tagged N11and SQR9, another biocontrol agent isolated from the cucumber rhizosphere and being shown a very broad antagonistic spectrum to Fusarium oxysporum, on banana roots was determined in hydroponic, sand and natural soil system, respectively. The appearance of the biofilm formation of the two Bacillus strains in vitro, the method for quantitative measurement of the biofilm biomass, and the dynamics of the biofilm biomass were studied. The response of N6, N11and SQR9towards the root exudates of banana and cucumber, as well as the organic acids composition, was investigated. Subsequently, the transcriptional profiling of SQR9towards different root exudates were analysed by Solexa high-throughout transcriptome sequence. At last, the effect of cheA deletion on biofilm formation of SQR9was studied. The main results obtained are listed as follows.
1. The capability of the specific bio-organic fertilizers (BIO2-4) for banana plants made of Bacillus amyloliquefaciens N6or/and Bacillus subtilis N11isolated from the banana rhizosphere to control the Fusarium wilt of banana was evaluated in pot experiments, and the normal bio-organic fertilizer (BIO1) made of Paenibacillus polymyxa SQR21and Trichoderma harzianum SQR-T37was also used for comparison. Results indicated that the application of BIO could decrease the disease incidence (DI) of banana from93.3%to below33.3%, and the DI in treatments applied with specific BIO (BIO2-4) was even lower than16.7%. The F. oxysporum was recovered from the bulk and rhizosphere soil by Quantitative Real-time PCR. Results revealed that the copy numbers of ITS gene of FOC in bulk soil were similar among all treatments. However, application of BIO could decrease the copy numbers of ITS gene of FOC inrhizophere soil from1.59×105g-1soil (control) to less than6×104g-1soil, and the treatments with specific BIOs (BIO2-4, T3-5) even suppressed the copy numbers to below1.5×104g"1soil. The microbial communities in the banana rhizosphere in all the treatments were analyzed by PCR-DGGE. No significant dissimilarities in species composition of the fungal communities were found. However, application of specific BIOs (T3-5) significantly increased the bacterial species.
2. An E. coli-Bacillus shuttle vector pHAPII, which contained a continually expressed Green Fluorescent Protein (GFP), was attempted introduce into N11by electroporation to obtain the GFP-tagged N11(N11-gfp). The cells and colonies of N11-gfp could emit bright green fluorescent under the microcopy. The growth speed and the antagonistic ability against FOC of N11-gfp were identical with that of the original strain. Later, the distribution of N11-gfp and SQR9-gfp, constructed by this laboratory previously, in banana rhizosphere was determined in hydroponic, sand and natural soil system by confocal laser scanning microscopy (CLSM) and scanning electron microscopy (SEM), respectively. Results indicated the colonization patterns of the biocontrol agents were similar among the three systems. Micrographs showed that2to4days after inoculation, the antagonists formed microcolonies and biofilm on the roots surface of banana plants, and the colonization mainly occurred on the elongation and differentiation zones of the main roots, the lateral roots, the junctions between roots, as well as the root hairs, while very few cells attached along the root tip. The quantitative analysis revealed that4days after inoculation, the population of N11-gfp and SQR9-gfp maintained at about105-106CFU·g-1roots, and106-107CFU·g-1soil.
3. The biofilm formation of several Bacillus strains in microtitre was studied in vitro. The biofilm formed by B. subtilis168in both LB and MSgg medium was thick, while the wide type strains FZB42, N6, N11and SQR9could form thin biofilm in both medium. It was found the biofilm formed in MSgg was thinner, also with more complex structure. The florescent microscopy pictures revealed that the biofilm of N11-gfp and SQR9-gfp formed on cover slides and microtitre presented complex three-dimensional architecture. The CLSM graphs of biofilm formed by SQR9-gfp confirmed this phenomenon. The traditionary violent staining method and a vortex method for quantitative measurement of the biofilm biomass were found to be identical, with a correlation index R2of0.9313. The biofilm formed by Bacillus strains displayed some edge-effects in the microtitre. At last, the dynamics of the biomass of biofilm and suspended cells of the four wide type strains were monitored. The biofilm biomass of each strain continuously increased from24h after inoculation, and maintained at about60h post inoculation. At this time the biomass of suspend cells began to increase. The disassembly of the biofilm occurred at96h after inoculation. The biofilm biomass of all the four strains in MSgg were higher than those in LB, and the biomass of the biofilm formed by FZB42and SQR9in both of the medium were larger than N6and N11.
4. The colonization of N11-gfp (isolated from banana rhizosphere) and SQK9-gfp (isolated from cucumber rhizosphere) on their original host plants was more effective than on the other plant. N6and N11displayed strong chemotactic reaction towards both the root exudates of banana and cucumber. SQR9showed ordinary reaction to the two kinds of root exudates. The root exudates of banana could significantly stimulated the biofilm formation of N6and N11, while both the two kinds of root exudates could significantly induced the biofilm formation of SQR9. High performance liquid chromatography (HPLC) analysis of the root exudates of banana revealed the existence of oxalic acid, malic acid, and fumaric acid, while oxalic acid, malic acid, and citric acid in those of cucumber. N6and N11showed intense chemotaxis reaction to malic acid, citric acid, and fumaric acid, while only the first two organic acids could significantly attract SQR9. Fumaric could significantly stimulate the biofilm formation of all the three strains, while citric acid could induce the biofilm formation of N6and SQR9. Colonization assay indicated that fumaric acid could improve the colonization of N11-gfp on banana and cucumber; while malic acid, citric acid, and fumaric could induce the survival of SQR9-gfp on banana or/and cucumber.
5. The shuttle vector pXKZ, which contained a competence factor ComK under the control of a xylose induced promoter (PxylA), was introduced into wide type Bacillus strains SQR9by conjunction. It was found that transformation of9kz by different plasmids was available with induction of xylose, and the transformation efficiency ranged from103to10CFU·μg-1. The transformation condition was optimized by Box-Behnken experiment with Response Surface Methodology (RSM), and the best situation was found to be:induction with0.2%of xylose for1h when the OD600of the suspension reached0.5, the transformation efficiency was approximately10CFU·μg-1DNA. The fragments contained the upstream and downstream arm for recombination with cheA, the suicide vector constructed by this fragment and pMD19-T, and the multimerzation of this suicide plasmid, were introduced into SQR9kz by induced transformation, respectively. Several corrected mutants were obtained by confirmation with PCR, and the highest efficiency were obtained for multimer transformation, which was still lower than10CFU·μg-1DNA. Then the pXKZ were eliminated by culture the mutant for several generations. The chemotactic response of the cheA null mutant towards citric acid was abolished. Its growth speed was similar with the wide type strain, while the biofilm formation delayed. The transmission electron microscopy (TEM) graphs revealed that the mutant failed to form flagellum, while the flagellums were found around the wide type cells. We speculated that the knockout of cheA caused some polar effects on the sigD in the downstream, thus influence the formation of flagellum, and disturb the chemotactic reaction, as well as delay of the biofilm formation, which needs complementation of cheA and sigD for confirmation.
6. The concentrated root exudates of cucumber, maize and banana could all significantly stimulate the growth of SQR9in1/2MSgg medium. Root exudates of maize could induce the biofilm formation of SQR9in1/2MSgg medium. The transcriptional profiling of SQR9towards the three kinds of root exudates under shaken condition, as well as its respond to maize exudates under static condition, were analysed by Solexa high-throughout transcriptome sequence. Results indicated the main regulated genes belonged to metabolism (mainly regarding carbohydrate, amino acids, lipid and nucleotide), protein translation and folding, membrane transport (ABC transporters), signal transduction (two-component system) and cell chemotaxis and motility. Some differences exist within the genes mediated by different root exudates. In static condition, the transcriptome of SQR9respond to maize roots after incubation for24h and48h, were quite different.
本文研究了由实验室筛选到的香蕉枯萎病拮抗菌N6(Bacillus amyloliquefaciens)或/和N11(Bacillus subtilis)制得的专用型生物有机肥及广谱型有机肥在盆栽实验中对土传香蕉枯萎病的防控效果,拮抗菌N11和广谱型拮抗菌SQR9(Bacillus amyloliquefaciens)在香蕉根际的定殖情况,拮抗菌生物膜形成的表型特征、定量方法和动态变化,三株拮抗菌对香蕉和黄瓜根系分泌物及其中各种有机酸成份的响应,及不同作物根系分泌物对SQR9基因表达影响的转录组分析。最后研究了cheA基因突变对SQR9生物膜形成能力的影响。获得了以下结果：
1.通过盆栽实验研究了由实验室筛选到的土传香蕉枯萎病拮抗菌N6(Bacillus amyloliquefaciens)或/和N11(Bacillus subtilis)制得的专用型生物有机肥(BIO2-4)在盆栽实验中对土传香蕉枯萎病的防控效果,并与实验室之前研发的由拮抗菌SQR21(Paenibacillus subtilis)和SQR-T37(Trichoderma harzianum)制得的生物有机肥(BIO1)的效果进行比较,发现生物有机肥可将香蕉植株发病率由93.3%降到33.3%以下,专用有机肥处理的植株发病率更低于16.7%。利用实时荧光定量PCR法对香蕉植株根际土及土体土中的尖孢镰刀菌进行了检测,结果表明处理的根际土总DNA的尖孢镰刀菌ITS基因拷贝数为1.59×105个g-1土,而施用BIO1可将其拷贝数下降到6×104个g-1土,香蕉专用有机肥BIO2-4(T3-5)可进一步将拷贝数抑制到1.5×104个g-1土以下。利用PCR-DGGE分析了各处理香蕉根际土壤的微生物群落结构,发现根际土壤的真菌群落变化不大,但专用型香蕉生物有机肥(BIO2-4)处理可以显著增加细菌种群的多样性。
2.试图通过电转化将持续表达绿色荧光蛋白(Green Fluorescent Protein, GFP)的大肠-枯草杆菌穿梭质粒pHAPII引入拮抗菌N11,成功得到了GFP标记的N11菌株(N11-gfp),其菌落和菌体在荧光显微镜下可发出明亮的绿色荧光。N11-gfp的生长速度和对香蕉枯萎病病原菌FOC的拮抗能力与原始菌株N11基本相同。随后通过水培、砂培和土培系统,利用激光共聚焦荧光显微镜(CLSM)和扫描电子显微镜(SEM)研究了N11-gfp菌株和实验室之前构建的SQR9-gfp在香蕉根系的定殖情况。发现三种系统中拮抗菌在香蕉根表定殖的情况相似,在接种后2-4天,拮抗菌即可在香蕉植株根表形成微菌落和生物膜,定殖主要发生在主根的伸长区和分生区,次生根,主根与侧根的交界处以及根毛区,而附着于根尖的拮抗菌数量很少。拮抗菌定量检测的结果表明接种4天后N11-gfp与SQR9-gfp在香蕉根表的密度保持在每克根105-106CFU,以及每克根际土106-107CFU。
3.通过体外实验研究了芽孢杆菌在细胞培养板中的生物膜形成情况,发现枯草芽孢杆菌168在LB与MSgg中形成的生物膜都较弱,而野生型的FZB42、N6、N11和SQR9都能形成较厚的生物膜,且在MSgg中形成的生物膜更厚、结构更复杂。荧光显微镜观察发现N11-gfp与SQR9-gfp在盖玻片和培养板中形成的生物膜具有一定的厚度和立体结构。CLSM观察也表明SQR9-gfp形成的生物膜具有一定的立体结构。传统的结晶紫染色法与震荡打碎法定量测定生物膜生物量的相关性较高,R2达到0.9313。实验中发现芽孢杆菌在细胞培养板中的成膜具有一定边缘效应。最后研究了4株野生型芽孢杆菌在LB与MSgg培养基中生物膜与悬浮细胞生物量的动态变化规律,发现各菌生物膜的生物量从24h开始增加,到60h时保持稳定,同时悬浮细胞开始增多,96h后生物膜逐渐消解。各菌在MSgg中形成的生物膜生物量都高于LB,FZB42与SQR9在两种培养基中形成的生物膜生物量都高于N6与N11。
4.从香蕉根际分离的拮抗菌N11和从黄瓜根际分离的拮抗菌SQR9在其原始宿主根表的定殖能力都显著强于另一种宿主根部。N6和N11对浓缩的香蕉和黄瓜根系分泌物均表现出强烈的趋化性,SQR9对两种分泌物的趋化反应则一般。香蕉分泌物可以显著诱导N6和N11生物膜的形成。利用HPLC分析从香蕉根系分泌物中鉴定出草酸、苹果酸和延胡索酸这3种主要的有机酸成分,在黄瓜根系分泌物中则鉴定出草酸、苹果酸和柠檬酸。N6和N11对苹果酸、柠檬酸和延胡索酸都表现出趋化性,SQR9只对苹果酸和柠檬酸有较强的趋化反应。延胡索酸可以显著增加N6、N11和SQR9形成生物膜的生物量,柠檬酸则都能显著诱导N6和SQR9生物膜的形成。体外定殖实验发现延胡索酸可以诱导N11-gfp在两种作物上的定殖,而苹果酸、柠檬酸和延胡索酸可促进SQR9-gfp在香蕉或／和黄瓜根际的定殖。
5.通过接合的方法将携带有在木糖诱导型启动子(PxylA)控制下表达的感受态因子ComK的穿梭质粒引入野生型芽孢杆菌SQR9,发现通过木糖诱导可实现外源质粒的转化,转化效率在103-104CFU·μg-1质粒。通过响应曲面法设计Box-Behnken实验对诱导转化的条件进行了优化,发现最佳条件为菌液OD600达到0.5时,用0.2%浓度的木糖诱导1h,该条件下转化质粒的效率可以接近104CFU·μg-1。将cheA基因的双同源臂突变片段,以pMD19-T为母体的自杀载体,及该质粒的多聚体通过诱导转化的方式引入9kz,经过PCR验证发现得到了正确的双交换的cheA突变体。利用传代培养删除了质粒pXKZ.突变体对趋化物柠檬酸的反应消失,生长速度与野生型相似,生物膜形成有滞后现象。透射电镜的观察发现突变体的鞭毛基本缺失,而野生型具周生鞭毛。推测对cheA的删除影响了下游sigD的转录和表达,从而影响了鞭毛的形成进而造成趋化性的消失和成膜的滞后,具体情况需要cheA和sigD的互补实验验证。
6.浓缩的黄瓜、玉米及香蕉的根系分泌物可显著促进SQR9在1/2浓度的MSgg中的生长(接种后6h)。玉米根系分泌物可诱导SQR9在1/2MSgg中的生物膜形成,利用Solexa高通量转录组测序研究了在振荡培养条件下三种作物分泌物,以及静态生物膜形成条件下玉米根系分泌物对SQR9基因表达的影响。结果表明主要调控的基因类群包括营养代谢(主要包括碳水化合物代谢、氨基酸代谢、脂类代谢和核酸代谢等)、蛋白质翻译与调控、跨膜运输(主要是ABC转运蛋白)、信号转导(主要为双组份调控系统)以及趋化及运动相关。不同根系分泌物调控的基因类群有一定差异。SQR9在静态条件下对玉米根系分泌物的转录组在接种后24h和48h有较大差异。
The control of Fusarium wilt is a world wide problem. The traditionally farming control methods have rarely provided long-term control in any production area, while selection of resistant cultivars and chemical application are not effective enough for control this disease, and also the latter causes environmental problems. Recently, biological control of soil-born disease is an efficient and environmentally friendly way and is receiving increased attention.
In this study, the capability of the specific bio-organic fertilizers made of organic fertilizer (pig manure compost and amino acid fertilizer) fermented with antagonisms Bacillus amyloliquefaciens N6or/and Bacillus subtilis N11, as well as the normal BIO developed before in this laboratory, to control the Fusarium wilt of banana was evaluated in pot experiments. The colonization pattern of GFP tagged N11and SQR9, another biocontrol agent isolated from the cucumber rhizosphere and being shown a very broad antagonistic spectrum to Fusarium oxysporum, on banana roots was determined in hydroponic, sand and natural soil system, respectively. The appearance of the biofilm formation of the two Bacillus strains in vitro, the method for quantitative measurement of the biofilm biomass, and the dynamics of the biofilm biomass were studied. The response of N6, N11and SQR9towards the root exudates of banana and cucumber, as well as the organic acids composition, was investigated. Subsequently, the transcriptional profiling of SQR9towards different root exudates were analysed by Solexa high-throughout transcriptome sequence. At last, the effect of cheA deletion on biofilm formation of SQR9was studied. The main results obtained are listed as follows.
1. The capability of the specific bio-organic fertilizers (BIO2-4) for banana plants made of Bacillus amyloliquefaciens N6or/and Bacillus subtilis N11isolated from the banana rhizosphere to control the Fusarium wilt of banana was evaluated in pot experiments, and the normal bio-organic fertilizer (BIO1) made of Paenibacillus polymyxa SQR21and Trichoderma harzianum SQR-T37was also used for comparison. Results indicated that the application of BIO could decrease the disease incidence (DI) of banana from93.3%to below33.3%, and the DI in treatments applied with specific BIO (BIO2-4) was even lower than16.7%. The F. oxysporum was recovered from the bulk and rhizosphere soil by Quantitative Real-time PCR. Results revealed that the copy numbers of ITS gene of FOC in bulk soil were similar among all treatments. However, application of BIO could decrease the copy numbers of ITS gene of FOC inrhizophere soil from1.59×105g-1soil (control) to less than6×104g-1soil, and the treatments with specific BIOs (BIO2-4, T3-5) even suppressed the copy numbers to below1.5×104g"1soil. The microbial communities in the banana rhizosphere in all the treatments were analyzed by PCR-DGGE. No significant dissimilarities in species composition of the fungal communities were found. However, application of specific BIOs (T3-5) significantly increased the bacterial species.
2. An E. coli-Bacillus shuttle vector pHAPII, which contained a continually expressed Green Fluorescent Protein (GFP), was attempted introduce into N11by electroporation to obtain the GFP-tagged N11(N11-gfp). The cells and colonies of N11-gfp could emit bright green fluorescent under the microcopy. The growth speed and the antagonistic ability against FOC of N11-gfp were identical with that of the original strain. Later, the distribution of N11-gfp and SQR9-gfp, constructed by this laboratory previously, in banana rhizosphere was determined in hydroponic, sand and natural soil system by confocal laser scanning microscopy (CLSM) and scanning electron microscopy (SEM), respectively. Results indicated the colonization patterns of the biocontrol agents were similar among the three systems. Micrographs showed that2to4days after inoculation, the antagonists formed microcolonies and biofilm on the roots surface of banana plants, and the colonization mainly occurred on the elongation and differentiation zones of the main roots, the lateral roots, the junctions between roots, as well as the root hairs, while very few cells attached along the root tip. The quantitative analysis revealed that4days after inoculation, the population of N11-gfp and SQR9-gfp maintained at about105-106CFU·g-1roots, and106-107CFU·g-1soil.
3. The biofilm formation of several Bacillus strains in microtitre was studied in vitro. The biofilm formed by B. subtilis168in both LB and MSgg medium was thick, while the wide type strains FZB42, N6, N11and SQR9could form thin biofilm in both medium. It was found the biofilm formed in MSgg was thinner, also with more complex structure. The florescent microscopy pictures revealed that the biofilm of N11-gfp and SQR9-gfp formed on cover slides and microtitre presented complex three-dimensional architecture. The CLSM graphs of biofilm formed by SQR9-gfp confirmed this phenomenon. The traditionary violent staining method and a vortex method for quantitative measurement of the biofilm biomass were found to be identical, with a correlation index R2of0.9313. The biofilm formed by Bacillus strains displayed some edge-effects in the microtitre. At last, the dynamics of the biomass of biofilm and suspended cells of the four wide type strains were monitored. The biofilm biomass of each strain continuously increased from24h after inoculation, and maintained at about60h post inoculation. At this time the biomass of suspend cells began to increase. The disassembly of the biofilm occurred at96h after inoculation. The biofilm biomass of all the four strains in MSgg were higher than those in LB, and the biomass of the biofilm formed by FZB42and SQR9in both of the medium were larger than N6and N11.
4. The colonization of N11-gfp (isolated from banana rhizosphere) and SQK9-gfp (isolated from cucumber rhizosphere) on their original host plants was more effective than on the other plant. N6and N11displayed strong chemotactic reaction towards both the root exudates of banana and cucumber. SQR9showed ordinary reaction to the two kinds of root exudates. The root exudates of banana could significantly stimulated the biofilm formation of N6and N11, while both the two kinds of root exudates could significantly induced the biofilm formation of SQR9. High performance liquid chromatography (HPLC) analysis of the root exudates of banana revealed the existence of oxalic acid, malic acid, and fumaric acid, while oxalic acid, malic acid, and citric acid in those of cucumber. N6and N11showed intense chemotaxis reaction to malic acid, citric acid, and fumaric acid, while only the first two organic acids could significantly attract SQR9. Fumaric could significantly stimulate the biofilm formation of all the three strains, while citric acid could induce the biofilm formation of N6and SQR9. Colonization assay indicated that fumaric acid could improve the colonization of N11-gfp on banana and cucumber; while malic acid, citric acid, and fumaric could induce the survival of SQR9-gfp on banana or/and cucumber.
5. The shuttle vector pXKZ, which contained a competence factor ComK under the control of a xylose induced promoter (PxylA), was introduced into wide type Bacillus strains SQR9by conjunction. It was found that transformation of9kz by different plasmids was available with induction of xylose, and the transformation efficiency ranged from103to10CFU·μg-1. The transformation condition was optimized by Box-Behnken experiment with Response Surface Methodology (RSM), and the best situation was found to be:induction with0.2%of xylose for1h when the OD600of the suspension reached0.5, the transformation efficiency was approximately10CFU·μg-1DNA. The fragments contained the upstream and downstream arm for recombination with cheA, the suicide vector constructed by this fragment and pMD19-T, and the multimerzation of this suicide plasmid, were introduced into SQR9kz by induced transformation, respectively. Several corrected mutants were obtained by confirmation with PCR, and the highest efficiency were obtained for multimer transformation, which was still lower than10CFU·μg-1DNA. Then the pXKZ were eliminated by culture the mutant for several generations. The chemotactic response of the cheA null mutant towards citric acid was abolished. Its growth speed was similar with the wide type strain, while the biofilm formation delayed. The transmission electron microscopy (TEM) graphs revealed that the mutant failed to form flagellum, while the flagellums were found around the wide type cells. We speculated that the knockout of cheA caused some polar effects on the sigD in the downstream, thus influence the formation of flagellum, and disturb the chemotactic reaction, as well as delay of the biofilm formation, which needs complementation of cheA and sigD for confirmation.
6. The concentrated root exudates of cucumber, maize and banana could all significantly stimulate the growth of SQR9in1/2MSgg medium. Root exudates of maize could induce the biofilm formation of SQR9in1/2MSgg medium. The transcriptional profiling of SQR9towards the three kinds of root exudates under shaken condition, as well as its respond to maize exudates under static condition, were analysed by Solexa high-throughout transcriptome sequence. Results indicated the main regulated genes belonged to metabolism (mainly regarding carbohydrate, amino acids, lipid and nucleotide), protein translation and folding, membrane transport (ABC transporters), signal transduction (two-component system) and cell chemotaxis and motility. Some differences exist within the genes mediated by different root exudates. In static condition, the transcriptome of SQR9respond to maize roots after incubation for24h and48h, were quite different.
引文
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