植物根际促生菌的筛选及应用
摘要
高产、优质和高效农业是我国农业发展的方向。这一进程中,化肥、农药、激素等化学农资产品的大量使用依然不可避免。化学农资产品的过度依赖必然造成环境污染和食品安全问题。因此,如何在相对减少农药化肥等用量的前提下,提高植物产量和品质,已成为农产品质量与安全领域的重要研究方向。对农学家和育种学家而言,改进栽培措施、培育优良品种,是解决这一问题的途径之一。除此之外,农用生物制剂如生物肥料和生物农药等因其效果显著、环境友好,也逐渐受到人们的关注。
农用生物制剂常用的微生物素材是植物根际促生菌(plant growth promoting rhizobacteria, PGPR),主要是定殖于植物根际系统,并能促进植物生长的一类细菌。PGPR能通过产生抵抗多种病原菌的抗生素类物质、毒素或诱导寄主植物产生病程相关蛋白等途径,帮助植物抵抗生物类侵害,包括病原细菌、真菌、病毒及线虫等；PGPR还能通过产生如ACC脱氨酶、IAA、气体信号分子等途径,帮助植物忍受多种非生物胁迫,包括重金属、干旱、盐分、肥力低下或过剩等。总之,PGPR作用机制多样,人们可以根据不同的需求,筛选合适的PGPR。然而,许多PGPR在实验室和温室条件下表现良好,但是在大田条件下的表现往往缺乏很好的重现性,这就限制了PGPR菌的大面积使用。因此,筛选高效广适的PGPR,并进行最适作用植物、土壤等条件的相关研究,成为研发生物制剂的关键。基于此,本研究致力于从不同植物根际筛选多株高效PGPR,并进行应用实践,以期为农用生物制剂的研制提供素材和依据。结果及结论如下：
(1)按国际通用方法,从盐城、扬州等地不同植物根际筛选到14株PGPR,体外促生指标显示,所筛菌株具有一定的应用前景,并从分子水平进行了菌株鉴定。分类鉴定结果显示：7株属于假单胞菌属(Pseudomonas)、3株属于类芽孢杆菌属(Paenibacillus)、2株为芽孢杆菌属(Bacillus)、1株为布克氏菌属(Burkholderia)、1株为欧文氏菌属(Erwinia)。
(2)通过体外促生试验明确：Pseudomonas sp. RA6和Bacillus sp. WP8可用于豇豆盆栽试验；Bacillus sp. RBB1、Bacillus sp. WP8和Pseudomonas sp. RBP1可用于水稻育秧试验；Erwinia sp. RA2和Bacillus sp.WP8可用于番茄青枯病的生物防治。
(3) PGPR对豇豆的促生作用因接种方式、接种量的不同而不同。WP8、RA6浸种处理的出苗率分别比对照(CK)提高14.29%和9.52%(p<0.05)；15d时的株高分别比CK提高14.39%和10.40%(p<0.05)；茎叶干物重分别比CK增加19.69%和17.71%(p<0.05)。WP8、 RA6低拌处理(104cfu/g soil)出苗率、15d时的株高以及茎叶干物重等指标与CK相比,均无显著差异(p>0.05); WP8. RA6中拌处理(106cfu/g soil)出苗率比CK提高4.76%,但未达显著差异(p>0.05)；15d时的株高分别比CK提高1.08%和5.65%(p>0.05)；茎叶干物重分别比CK增加12.71%和18.59%(p<0.05)。WP8、RA6高拌处理(108cfu/g soil)出苗率分别比CK提高9.52%和14.29%(p<0.05)；15d时的株高分别比CK提高6.37%和7.64%(p<0.05)；茎叶干物重分别比CK增加27.37%和20.43%(p<0.05)。说明供试菌株对豇豆均有一定程度的促生作用,表现在提高豇豆种子出苗率,增加茎叶干物重,提高植物高度,且浸种处理效果优于拌土处理。分析还表明,茎叶干物重和种子出苗率显著相关,说明用茎叶干物重作为PGPR促生的生物学指标更为敏感。
DGGE指纹图谱分析结果显示：各处理在15d和45d时,除WP8浸种处理外,其余土壤微生物群落多样性和CK均已发生明显变化,其中RA6菌株在土壤中可随时间推延,优势地位更趋明显,表现在45d时仍可明显检测到；WP8在土壤中存活时间不长,但拌十处理改变了土著细菌的群落结构。试验暗示WP8的促生作用很可能与土著微生物群落的变化有关。
(4)通过水稻塑盘育秧的促生试验,我们发现：①RBP1和WP8可不同程度地促进秧苗生长,主要表现在促使秧苗矮壮、增加干物质积累；②PGPR拌土普遍优于浸种方式；③PGPR的有效性受是否与壮秧剂混用的影响,其次是接种方式；④地上部干物重对PGPR不同处理方式较为敏感,是评价促生效果的理想指标；⑤PGPR对土壤细菌群落结构有一定的影响,但不十分明显。
(5)供试菌株WP8、RA2浸种处理12d时的出苗率分别达到90.4%和92.5%,显著高于青枯病菌处理(71.8%)和对照处理(60.2%)；健株率分别达62.8%和68.9%,远高于青枯病菌处理(22.4%)；生防效果分别为52.1%和59.9%；35d时的株高分别比病原菌对照处理高86.00%和81.56%p<0.05)；根长分别比病原菌对照处理长136.91%和118.12%p<0.05)；茎基粗分别比病原菌对照处理增粗109.65%和96.49%p<0.05)；茎叶干物重分别比病原菌对照处理增重110.82%和63.20%(p<0.05)；根干重分别比病原菌对照处理增重205.83%和159.13%(p<0.05)；35d时的土壤水稳性团聚体比例分别比病原菌对照处理增加156.88%和55.56%(p<0.05)。WP8、RA2拌土处理12d时的出苗率和对照相比未见增加,只有62.2%和72.4%；健株率分别比病原菌对照处理增加116.52%和133.48%(p<0.05)；生防效果为33.6%和38.5%,分别比各自浸种处理低35.51%和35.73%(p<0.05)；35d时的株高分别比病原菌对照处理高69.56%和92.00%(p<0.05)；根长分别比病原菌对照处理长120.40%和56.86%(p<0.05)；茎基粗分别比病原菌对照处理增粗131.58%和100.00%(p<0.05);茎叶干物重分别比病原菌对照处理增重53.68%和34.20%(p<0.05)；根干重分别比病原菌对照处理增重128.53%和164.52%(p<0.05)；35d时的土壤水稳性团聚体比例分别比病原菌对照处理增加42.87%(p<0.05)和降低31.88%(p<0.05)。
PGPR处理土壤中青枯病菌依然存在,说明生防途径并非通过减少该菌在土壤中的数量实现的；此外,论文还对病原菌处理的一些特征条带进行了分析。这些结果说明,2株PGPR都具有防治番茄青枯病的作用,并能不同程度地促进番茄幼苗生长,浸种处理的促进效应明显优于拌土处理；PGPR处理还能在一定程度上提高土壤水稳性团聚体(>0.25mm)比例,WP8浸种处理尤为明显。根际微生物群落受番茄种植的影响最大,其次是青枯病菌,受PGPR的影响最小；PGPR处理土壤中青枯病菌依然存在,说明生防途径并非通过减少该菌在土壤中的数量实现的。
通过本研究,明确了促生不同植物生长所需的最适PGPR,以及最佳的接种方式。PGPR对土壤土著微生物群落的影响等结论也为进一步应用提供理论基础；供试菌株中,WP8相对具有促生广适性,更具应用潜力。
High output, high quality and high efficiency are the objectives of modern agriculture. To achieve these goals, Chemical fertilizer, pesticide, hormones and other agricultural chemicals have been widely used. However such chemical-dependent agriculture inevitably results in environmental pollution and food safety issues. Therefore, scientific community is making great efforts to seek the ways to reduce chemical inputs but maintain high output and high quality of agricultural products. Improved cultural practices, breeding new varieties and application of plant growth promoting agents are the options.
Plant growth promoting rhizobacteria (PGPR) are the bacteria living in rhizospheric soils that promote plant growth. Growth promotion has been believed to be due to production of antibiotics and toxins (IAA and signal compounds) against pathogens. PGPRs also interact with plants by regulating plant metabolisms so as to help plant resist to pathogenic bacteria, fungus, virus and nematodes. Because of diversified acting modes of PGPRs, it is possible to obtain pathogen-, soil-, or plant-specific strains by screening. However, good results obtained in vitro cannot always be reproduced under field conditions, limiting the wide use of plant growth promoting agents. Therefore, screening strains with wide spectrum of acting mechanisms and finding their most effective plants are the keys to develop biocontrol agents. The purpose of present work is to screen high efficient PGPRs and test their effects in pots. Results are summarized as follows.
(1) Fourteen strains were isolated from rhizospheric soils sampled from Yangzhou and Yancheng, Jiangsu province, seven of which are Pseudomonas sps., three are Paenibacillus sps., two are Bacillus sps., one is Burkholderia sp., and one is Erwinia sp. according to their16S rDNA sequences. Assay in vitro showed these strains were all potential plant growth promoting agents.
(2) Assay in vitro showed that Bacillus sp. WP8, Pseudomonas sp. RA6promoted the growth of cowpea, whereas Bacillus sp. RBB1、Bacillus sp. WP8and Pseudomonas sp. RBP1promoted rice seedling growth. Erwinia sp. RA2and Bacillus sp. WPS showed antigenic effects on biotic Ralstonia solanacearum.
(3) PGPR effects of isolated strains on cowpea growth depended on inoculation methods and inoculation rates. By seed soaking with WP8and RA6, germination rate of cowpea seeds was increased by14.29%and9.52%respectively (p<0.05) as compared with control; plant height at15days after germination was increased by14.39%and10.40%(p<0.05) respectively; above-ground dry weight was increased by19.69%and17.71%(p<0.05) respectively. By soil drenching with WP8and RA6, plant height and above-ground dry weight were not significantly different from those of control when inoculation rate was low (104cfu/g soil). However when inoculation rate was increased to108cfu/g soil, seed germination rate was increased by9.52%and14.29%respectively (p<0.05); plant height at15days after germination was increased by6.37%and7.64%(p<0.05); above-ground dry weight was increased by27.37%and20.43%(p<0.05). These data suggested that the PGPR strains were potential bio-agents promoting cowpea growth. Seed soaking was better than soil drenching. Further analysis showed that above-ground dry weight was significantly correlated with germination rate, suggesting that above-ground dry weight was a sensitive indicator indicating growth promoting effects of strains.
Denaturing gradient gel electrophoresis (DGGE) analysis based on16S rRNA sequences indicated that microbial communities in strain-treated soils were obviously different from those in control soil. Strain RA6-specific band became intensified as the time passed. However, strain WP8-specific band became weak as time passed.
(4) Tested in rice seedling beds, it was found that strain RBP1and WP8promoted seedling growth to various extents. Seedlings treated with these strains showed short but strong growth and heavy dry weight. Soil drenching showed better results than seed soaking. Application of "ZYJ", a chemical that makes rice seedlings short and strong, overshadowed the effects of strain inoculation. Above-ground dry weight was a sensitive index to evaluate the effects of PGPR strains. Inoculation of PGPR strains insignificantly modified soil microbial community structure.
(5) Germination rate treated with WP8and RA2by seed soaking was90.4%and92.5%respectively, significantly higher than that treated with pathogen Ralstonia solanacearum (71.8%) and that of control (60.2%). The percentage of number of healthy plants to total number of plants in WP8-and RA2-treatments was62.8%and68.9%respectively, significantly higher than that in Ralstonia solanacearum-treatment (22.4%). Plant height measured at35days after germination was increased by86.00%and81.56%(p<0.05) as compared to that of pathogen-and control-treatment respectively. Root length, stem thickness, above-ground dry weight and root dry weight were all better in PGPR treatments than in pathogen-or control-treatments. Soil water stable aggregates was increased by156.88%and55.56%(p<0.05) as compared to that of pathogen-and control-treatment respectively. When the strains were inoculated by soil drenching, all these parameters were either better or insignificantly different in PGPR-treatments than those in pathogen-or control-treatments.
It was demonstrated that pathogen Ralstonia solanacearum was present in soils that were treated with PGPR strains, suggesting that bio-control effects were achieved not by reducing the abundance of pathogen, but by other yet-to-defined mechanisms.
Present work defined the suitable PGPR strains with respect to different plants and inoculation methods. The fact that inoculation of PGPR strains modified native microbial community to certain extent needs to be further studied. Strain WP8showed wide spectrum of acting modes as a bio-control and or plant growth promoting agent and was thus proposed as a potential applicable strain.
农用生物制剂常用的微生物素材是植物根际促生菌(plant growth promoting rhizobacteria, PGPR),主要是定殖于植物根际系统,并能促进植物生长的一类细菌。PGPR能通过产生抵抗多种病原菌的抗生素类物质、毒素或诱导寄主植物产生病程相关蛋白等途径,帮助植物抵抗生物类侵害,包括病原细菌、真菌、病毒及线虫等；PGPR还能通过产生如ACC脱氨酶、IAA、气体信号分子等途径,帮助植物忍受多种非生物胁迫,包括重金属、干旱、盐分、肥力低下或过剩等。总之,PGPR作用机制多样,人们可以根据不同的需求,筛选合适的PGPR。然而,许多PGPR在实验室和温室条件下表现良好,但是在大田条件下的表现往往缺乏很好的重现性,这就限制了PGPR菌的大面积使用。因此,筛选高效广适的PGPR,并进行最适作用植物、土壤等条件的相关研究,成为研发生物制剂的关键。基于此,本研究致力于从不同植物根际筛选多株高效PGPR,并进行应用实践,以期为农用生物制剂的研制提供素材和依据。结果及结论如下：
(1)按国际通用方法,从盐城、扬州等地不同植物根际筛选到14株PGPR,体外促生指标显示,所筛菌株具有一定的应用前景,并从分子水平进行了菌株鉴定。分类鉴定结果显示：7株属于假单胞菌属(Pseudomonas)、3株属于类芽孢杆菌属(Paenibacillus)、2株为芽孢杆菌属(Bacillus)、1株为布克氏菌属(Burkholderia)、1株为欧文氏菌属(Erwinia)。
(2)通过体外促生试验明确：Pseudomonas sp. RA6和Bacillus sp. WP8可用于豇豆盆栽试验；Bacillus sp. RBB1、Bacillus sp. WP8和Pseudomonas sp. RBP1可用于水稻育秧试验；Erwinia sp. RA2和Bacillus sp.WP8可用于番茄青枯病的生物防治。
(3) PGPR对豇豆的促生作用因接种方式、接种量的不同而不同。WP8、RA6浸种处理的出苗率分别比对照(CK)提高14.29%和9.52%(p<0.05)；15d时的株高分别比CK提高14.39%和10.40%(p<0.05)；茎叶干物重分别比CK增加19.69%和17.71%(p<0.05)。WP8、 RA6低拌处理(104cfu/g soil)出苗率、15d时的株高以及茎叶干物重等指标与CK相比,均无显著差异(p>0.05); WP8. RA6中拌处理(106cfu/g soil)出苗率比CK提高4.76%,但未达显著差异(p>0.05)；15d时的株高分别比CK提高1.08%和5.65%(p>0.05)；茎叶干物重分别比CK增加12.71%和18.59%(p<0.05)。WP8、RA6高拌处理(108cfu/g soil)出苗率分别比CK提高9.52%和14.29%(p<0.05)；15d时的株高分别比CK提高6.37%和7.64%(p<0.05)；茎叶干物重分别比CK增加27.37%和20.43%(p<0.05)。说明供试菌株对豇豆均有一定程度的促生作用,表现在提高豇豆种子出苗率,增加茎叶干物重,提高植物高度,且浸种处理效果优于拌土处理。分析还表明,茎叶干物重和种子出苗率显著相关,说明用茎叶干物重作为PGPR促生的生物学指标更为敏感。
DGGE指纹图谱分析结果显示：各处理在15d和45d时,除WP8浸种处理外,其余土壤微生物群落多样性和CK均已发生明显变化,其中RA6菌株在土壤中可随时间推延,优势地位更趋明显,表现在45d时仍可明显检测到；WP8在土壤中存活时间不长,但拌十处理改变了土著细菌的群落结构。试验暗示WP8的促生作用很可能与土著微生物群落的变化有关。
(4)通过水稻塑盘育秧的促生试验,我们发现：①RBP1和WP8可不同程度地促进秧苗生长,主要表现在促使秧苗矮壮、增加干物质积累；②PGPR拌土普遍优于浸种方式；③PGPR的有效性受是否与壮秧剂混用的影响,其次是接种方式；④地上部干物重对PGPR不同处理方式较为敏感,是评价促生效果的理想指标；⑤PGPR对土壤细菌群落结构有一定的影响,但不十分明显。
(5)供试菌株WP8、RA2浸种处理12d时的出苗率分别达到90.4%和92.5%,显著高于青枯病菌处理(71.8%)和对照处理(60.2%)；健株率分别达62.8%和68.9%,远高于青枯病菌处理(22.4%)；生防效果分别为52.1%和59.9%；35d时的株高分别比病原菌对照处理高86.00%和81.56%p<0.05)；根长分别比病原菌对照处理长136.91%和118.12%p<0.05)；茎基粗分别比病原菌对照处理增粗109.65%和96.49%p<0.05)；茎叶干物重分别比病原菌对照处理增重110.82%和63.20%(p<0.05)；根干重分别比病原菌对照处理增重205.83%和159.13%(p<0.05)；35d时的土壤水稳性团聚体比例分别比病原菌对照处理增加156.88%和55.56%(p<0.05)。WP8、RA2拌土处理12d时的出苗率和对照相比未见增加,只有62.2%和72.4%；健株率分别比病原菌对照处理增加116.52%和133.48%(p<0.05)；生防效果为33.6%和38.5%,分别比各自浸种处理低35.51%和35.73%(p<0.05)；35d时的株高分别比病原菌对照处理高69.56%和92.00%(p<0.05)；根长分别比病原菌对照处理长120.40%和56.86%(p<0.05)；茎基粗分别比病原菌对照处理增粗131.58%和100.00%(p<0.05);茎叶干物重分别比病原菌对照处理增重53.68%和34.20%(p<0.05)；根干重分别比病原菌对照处理增重128.53%和164.52%(p<0.05)；35d时的土壤水稳性团聚体比例分别比病原菌对照处理增加42.87%(p<0.05)和降低31.88%(p<0.05)。
PGPR处理土壤中青枯病菌依然存在,说明生防途径并非通过减少该菌在土壤中的数量实现的；此外,论文还对病原菌处理的一些特征条带进行了分析。这些结果说明,2株PGPR都具有防治番茄青枯病的作用,并能不同程度地促进番茄幼苗生长,浸种处理的促进效应明显优于拌土处理；PGPR处理还能在一定程度上提高土壤水稳性团聚体(>0.25mm)比例,WP8浸种处理尤为明显。根际微生物群落受番茄种植的影响最大,其次是青枯病菌,受PGPR的影响最小；PGPR处理土壤中青枯病菌依然存在,说明生防途径并非通过减少该菌在土壤中的数量实现的。
通过本研究,明确了促生不同植物生长所需的最适PGPR,以及最佳的接种方式。PGPR对土壤土著微生物群落的影响等结论也为进一步应用提供理论基础；供试菌株中,WP8相对具有促生广适性,更具应用潜力。
High output, high quality and high efficiency are the objectives of modern agriculture. To achieve these goals, Chemical fertilizer, pesticide, hormones and other agricultural chemicals have been widely used. However such chemical-dependent agriculture inevitably results in environmental pollution and food safety issues. Therefore, scientific community is making great efforts to seek the ways to reduce chemical inputs but maintain high output and high quality of agricultural products. Improved cultural practices, breeding new varieties and application of plant growth promoting agents are the options.
Plant growth promoting rhizobacteria (PGPR) are the bacteria living in rhizospheric soils that promote plant growth. Growth promotion has been believed to be due to production of antibiotics and toxins (IAA and signal compounds) against pathogens. PGPRs also interact with plants by regulating plant metabolisms so as to help plant resist to pathogenic bacteria, fungus, virus and nematodes. Because of diversified acting modes of PGPRs, it is possible to obtain pathogen-, soil-, or plant-specific strains by screening. However, good results obtained in vitro cannot always be reproduced under field conditions, limiting the wide use of plant growth promoting agents. Therefore, screening strains with wide spectrum of acting mechanisms and finding their most effective plants are the keys to develop biocontrol agents. The purpose of present work is to screen high efficient PGPRs and test their effects in pots. Results are summarized as follows.
(1) Fourteen strains were isolated from rhizospheric soils sampled from Yangzhou and Yancheng, Jiangsu province, seven of which are Pseudomonas sps., three are Paenibacillus sps., two are Bacillus sps., one is Burkholderia sp., and one is Erwinia sp. according to their16S rDNA sequences. Assay in vitro showed these strains were all potential plant growth promoting agents.
(2) Assay in vitro showed that Bacillus sp. WP8, Pseudomonas sp. RA6promoted the growth of cowpea, whereas Bacillus sp. RBB1、Bacillus sp. WP8and Pseudomonas sp. RBP1promoted rice seedling growth. Erwinia sp. RA2and Bacillus sp. WPS showed antigenic effects on biotic Ralstonia solanacearum.
(3) PGPR effects of isolated strains on cowpea growth depended on inoculation methods and inoculation rates. By seed soaking with WP8and RA6, germination rate of cowpea seeds was increased by14.29%and9.52%respectively (p<0.05) as compared with control; plant height at15days after germination was increased by14.39%and10.40%(p<0.05) respectively; above-ground dry weight was increased by19.69%and17.71%(p<0.05) respectively. By soil drenching with WP8and RA6, plant height and above-ground dry weight were not significantly different from those of control when inoculation rate was low (104cfu/g soil). However when inoculation rate was increased to108cfu/g soil, seed germination rate was increased by9.52%and14.29%respectively (p<0.05); plant height at15days after germination was increased by6.37%and7.64%(p<0.05); above-ground dry weight was increased by27.37%and20.43%(p<0.05). These data suggested that the PGPR strains were potential bio-agents promoting cowpea growth. Seed soaking was better than soil drenching. Further analysis showed that above-ground dry weight was significantly correlated with germination rate, suggesting that above-ground dry weight was a sensitive indicator indicating growth promoting effects of strains.
Denaturing gradient gel electrophoresis (DGGE) analysis based on16S rRNA sequences indicated that microbial communities in strain-treated soils were obviously different from those in control soil. Strain RA6-specific band became intensified as the time passed. However, strain WP8-specific band became weak as time passed.
(4) Tested in rice seedling beds, it was found that strain RBP1and WP8promoted seedling growth to various extents. Seedlings treated with these strains showed short but strong growth and heavy dry weight. Soil drenching showed better results than seed soaking. Application of "ZYJ", a chemical that makes rice seedlings short and strong, overshadowed the effects of strain inoculation. Above-ground dry weight was a sensitive index to evaluate the effects of PGPR strains. Inoculation of PGPR strains insignificantly modified soil microbial community structure.
(5) Germination rate treated with WP8and RA2by seed soaking was90.4%and92.5%respectively, significantly higher than that treated with pathogen Ralstonia solanacearum (71.8%) and that of control (60.2%). The percentage of number of healthy plants to total number of plants in WP8-and RA2-treatments was62.8%and68.9%respectively, significantly higher than that in Ralstonia solanacearum-treatment (22.4%). Plant height measured at35days after germination was increased by86.00%and81.56%(p<0.05) as compared to that of pathogen-and control-treatment respectively. Root length, stem thickness, above-ground dry weight and root dry weight were all better in PGPR treatments than in pathogen-or control-treatments. Soil water stable aggregates was increased by156.88%and55.56%(p<0.05) as compared to that of pathogen-and control-treatment respectively. When the strains were inoculated by soil drenching, all these parameters were either better or insignificantly different in PGPR-treatments than those in pathogen-or control-treatments.
It was demonstrated that pathogen Ralstonia solanacearum was present in soils that were treated with PGPR strains, suggesting that bio-control effects were achieved not by reducing the abundance of pathogen, but by other yet-to-defined mechanisms.
Present work defined the suitable PGPR strains with respect to different plants and inoculation methods. The fact that inoculation of PGPR strains modified native microbial community to certain extent needs to be further studied. Strain WP8showed wide spectrum of acting modes as a bio-control and or plant growth promoting agent and was thus proposed as a potential applicable strain.
引文
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