菌根真菌和生物有机肥结合对棉花土传黄萎病防控作用及其机制
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摘要
棉花土传黄萎病是棉花连作障碍的主要病害之一,是一种典型的土传病害,其病原菌为大丽轮枝菌(Verticillium dahliae Kleb).传统的化学防控通常会带来环境问题而田间防控措施如轮作和采用抗病品种等也存在许多限制性因素,生物防控日渐成为克服土传植物病害的重要途径。已有的研究表明菌根真菌或死谷芽孢杆菌菌株(Bacillus vallismortis,HJ-5)单独施用对防控棉花土传黄萎病有效果,但二者共同接种对防控棉花土传黄萎病的防控效果和协同作用机制尚未见报道。本文首先通过盆栽试验筛选了适于双接种试验的菌根菌菌株和菌株HJ-5的菌剂接种量。通过三次独立的温室盆栽试验研究了结合施用地表球囊霉(Glomus versiforme, Gv)和死谷芽孢杆菌强化的生物有机肥对棉花土传黄萎病的防病效果。采用绿色荧光蛋白(Green fluorescent protein, GFP)标记技术构建了菌株HJ-5的GFP标记菌株(GHJ-5),研究了菌株GHJ-5在植株根表和根际的定殖行为。在温室条件下利用分根试验研究了菌株HJ-5和Gv单独接种和混合接种对棉花根系分泌物及抑病效果的影响。另外,还通过一系列的实验室和温室试验了研究病原菌和拮抗菌之间的互作关系。主要结果如下。
1.筛选菌根菌和菌株HJ-5接种量的试验结果表明,双接种处理的最适菌根菌菌剂为菌株Gv,菌株HJ-5的最适用量为107CFU·g-1.菌株Gv和HJ-5双接种处理的土传黄萎病发病率最低,发病指数下降了26.71%,植株生物量显著增加。
2.三次独立的盆栽试验一致表明,育苗时先接种菌根真菌菌株Gv,移栽时再在连作土壤中施用死谷芽孢杆菌发酵的生物有机肥,防控棉花土传黄萎病的效果最好,发病指数平均降低65%,植株地上部高度,叶面积,地上,下部干重量三次平均分别增加75%,44%,216%,和71%。植株地上部和地下部N和P含量显著增加。对照相比,植株地上部和地下部N含量分别增加83%和65%；地上部和地下部P含量分别增加49%和45%。根际土壤Verticillium dahliae数量显著降低,下降幅度达83%。值得注意的是,菌株Gv在植物根的定殖没有受施用HJ-5生物有机肥的影响。
3.实验室分析表明,菌株HJ-5能产生脂肽类物质、吲哚乙酸(IAA)、铁载体、氰化氢(HCN)、氨气,但不产脱氨酶(ACC),不具有解磷能力。GFP标记菌株GHJ-5能够定殖在土传黄萎病病原菌的菌丝上并使其变形和裂解,表明菌株HJ-5为土传黄萎病的拮抗菌。
4.双接种菌株Gv和GHJ-5的盆栽试验结果表明,与单接种菌株GHJ-5或Gv相比,棉花植株的生物量、根系密度、植株磷含量、棉花植株根内和根际的菌株GHJ-5的数量都有显著增加,说明菌株Gv与GHJ-5双接种能够促进棉花的生长。
5.分根试验表明,双接种菌株Gv和HJ-5处理棉花土传黄萎病发病指数比对照下降了63.3%。棉花根系分泌物中发现4种酚酸,所有酚酸在低浓度时促进大丽轮枝菌的生长和孢子的萌发,浓度升高到一定程度,抑制病原菌孢子的生长。接种菌株Gv和HJ-5显著地减低了根系分泌物中酚酸类物质的浓度。接种V. dahliae、菌株Gv或HJ-5均系统性地改变了根系分泌物的组成。
6.对盆栽试验植株样品的分析表明,双接种菌株Gv和HJ-5显著提高了植株防卫酶如内过氧化物酶(POD)、苯丙氨酸解氨酶(PAL)、几丁质酶等酶的活性,增加了叶片水杨酸和叶绿素含量、降低了植株茎基部胼胝质含量,减轻棉花土传黄萎病发病症状。
综上所述,本文证实了菌株Gv和HJ-5双接种处理具有协同促进植物生长和抑制植物病原菌生长的作用。菌株Gv能够帮助植物吸收土壤中的养分元素,菌株HJ-5具有产生拮抗物质的能力,二者在土壤和植株上的成功定殖是防控棉花土传黄萎病的关键。接种菌株HJ-5和Gv对棉花根系分泌物的改变作用与植物系统抗性有关,这与接种菌株HJ-5和Gv之后植物防御性酶活性等特征的变化是一致的。本文的研究结果有助于研发新型棉花土传黄萎病防控技术。今后应进一步研究生物有机肥中功能菌在修复土壤障碍因子中的分子生物学机制。
Verticillium wilt of cotton caused by fungal pathogen Verticillium dahliae Kleb is a severe disease that limits the development of the cotton industry in the world. While chemical control is challenged by environmental problems and field control practices such as crop rotation and disease-resistant cultivar adoption exists many limting factores, biological control becomes increasingly an important way in control of soilborne plant diseases. Previous reports have shown that arbarscular mycorrhizal fungi (AMF) and Bacillus vallismortis HJ-5have biological control effects against Verticillium wilt of cotton. However, little has been known the effect of dual inoculation of soil with AMF and the bacterium and their synergetic mechanisms. Greenhouse pot experiments were firstly conducted to study several AMF and the inoculation doses of HJ-5for dual inoculation. Three independent greenhouse pot experiments were conducted to study the effects of the combined application of Glomus versiforme (Gv) propagules and HJ-5fortified bio-organic fertilizer on control of Verticillium wilt of cotton. Green fluorescent protein (GFP) labeled strain (GHJ-5) of strain HJ-5was constructed by using green GFP labeling techniques and the ability of GHJ-5to colonize roots and rhizosphere soil of cotton was also studied. To identify the effects of HJ-5and Gv alone or both on root exudates during the incidence of the wilt disease, a split-root experiment was conducted in greenhouse. Additionally, a series of laboratory experiments were performed on observation of the interaction between the plant pathogen and the biocontrol agents. Results were listed as follows.
(1) The primary screening experiments for selection of the best AMF strain and dosage of HJ-5showed that Gv was the suitable AMF strain and the proper innoclation dose of HJ-5was107CFU·g-1. The dual inoculation treatment with Gv and HJ-5had the lowest disease incidence of Verticillium wilt, which was reduced by26.71%as compared with other treatments, and the highest increase of plant biomass.
(2) Inoculation of nursery soil with propagula of Gv and application of the HJ-5enhanced bio-organic fertilizer in the diseased pot soil achieved agreeable high effecenies in suppressing Verticillium wilt by three independent pot experiments. Combined treatment decreased the disease index of Verticillium wilt by65%and increased plant height, leaf area, shoot and root dry weigh by75%,44%,216%, and71%respectively, on the average of three trials as compared with the non-inoculation control. Dual inoculation also significantly increased N in shoot and root and P in shoot and root by83%and65%,49%and45%on the average, respectively, while decreased the number of phytopathogenic Verticillium dahliae in rhizosphere soil of cotton by83%on the average, as compared with the non-inoculation control. Interestingly, the colonization of cotton roots by Gv was not infected by inoculation with HJ-5.
(3) Measurement from laboratory tests showed that the HJ-5was able to produce lipopeptide substances, growth-promoting substances indole acetic acid (IAA), siderophore, hydrogen cyanide (HCN) and ammonia, but not produce deaminase (ACC) and was unable to dissolve P in medium. Colonization, deformation and dissociation of hypha of pathogen V. dahliae Kleb by the green fluorescent protein labelled HJ-5(GHJ-5) was also obsverved, suggesting that HJ-5was an effective antagonist against the pathogen of Verticillium wilt of cotton.
(4) Dual inoculation of soil with GHJ-5and Gv in the greenhouse experiment showed that the cotton plant biomass, root density, plant phosphorus content significantly as well as the number of colonization of the rhizosphere of GHJ-5increased as compared with single inoculation with GHJ-5or Gv, suggesting that synergistic effect existed in promotion of the growth of cotton and suppression of Verticillium wilt of cotton.
(5) In root-splicing experiment, the co-inoculation with HJ-5and Gv reduced disease index by63.3%as compared with the control. Four phenolic acids were found in the cotton root exudates, and all the phenolic acids at low concentrations stimulated germination of V. dahliae spores, while higher concentrations were inhibitory. The phenolic acid concentrations in the root exudates decreased significantly with the application of HJ-5and Gv. The the exudation pattern of cotton roots by HJ-5, Gv and V. dahliae was systemically altered.
(6) In the pot experiment, inoculation of Gv and HJ-5significantly increased the activities of plant defence enzymes such as polyphenoloxidase (PPO), phenylalanine ammonialyase (PAL) and chitinase, increased salicylic acid chlorophyll in plant leaves, and decreased callose in crown of cotton, and consequently relieved the symptoms of Verticillium wilt of cotton.
In conclusion, the present study confirmed that co-inoculation of soil with AMF and rhizosphere bacteria had synergetic promotion of plant growth and resistance to phytopathogens. Abilities of Gv to facilitate plant uptake of nutrients and abilities of HJ-5to produce beneficial secondary metabolics against V. dahliae Kleb as well as successful colonization of soil and plant roots were essential to control Verticillium wilt of cotton. The alteration of the exudation pattern of cotton roots by HJ-5and Gv was related to plant systemic resistance to the phatopathogen. This was also supported by the enhanced activities of plant defence enzymes after inoculation of Gv and HJ-5. Results in this study would be useful in the developping new methods to better control cotton wilt disease. Further researches should be given to the molecular microbilogical aspects of the functions performed by bio-organic fertilizers in remediation of continuously cropping obstacles and in the promotion of plant growth and health exerted by soil functional microbes.
1.筛选菌根菌和菌株HJ-5接种量的试验结果表明,双接种处理的最适菌根菌菌剂为菌株Gv,菌株HJ-5的最适用量为107CFU·g-1.菌株Gv和HJ-5双接种处理的土传黄萎病发病率最低,发病指数下降了26.71%,植株生物量显著增加。
2.三次独立的盆栽试验一致表明,育苗时先接种菌根真菌菌株Gv,移栽时再在连作土壤中施用死谷芽孢杆菌发酵的生物有机肥,防控棉花土传黄萎病的效果最好,发病指数平均降低65%,植株地上部高度,叶面积,地上,下部干重量三次平均分别增加75%,44%,216%,和71%。植株地上部和地下部N和P含量显著增加。对照相比,植株地上部和地下部N含量分别增加83%和65%；地上部和地下部P含量分别增加49%和45%。根际土壤Verticillium dahliae数量显著降低,下降幅度达83%。值得注意的是,菌株Gv在植物根的定殖没有受施用HJ-5生物有机肥的影响。
3.实验室分析表明,菌株HJ-5能产生脂肽类物质、吲哚乙酸(IAA)、铁载体、氰化氢(HCN)、氨气,但不产脱氨酶(ACC),不具有解磷能力。GFP标记菌株GHJ-5能够定殖在土传黄萎病病原菌的菌丝上并使其变形和裂解,表明菌株HJ-5为土传黄萎病的拮抗菌。
4.双接种菌株Gv和GHJ-5的盆栽试验结果表明,与单接种菌株GHJ-5或Gv相比,棉花植株的生物量、根系密度、植株磷含量、棉花植株根内和根际的菌株GHJ-5的数量都有显著增加,说明菌株Gv与GHJ-5双接种能够促进棉花的生长。
5.分根试验表明,双接种菌株Gv和HJ-5处理棉花土传黄萎病发病指数比对照下降了63.3%。棉花根系分泌物中发现4种酚酸,所有酚酸在低浓度时促进大丽轮枝菌的生长和孢子的萌发,浓度升高到一定程度,抑制病原菌孢子的生长。接种菌株Gv和HJ-5显著地减低了根系分泌物中酚酸类物质的浓度。接种V. dahliae、菌株Gv或HJ-5均系统性地改变了根系分泌物的组成。
6.对盆栽试验植株样品的分析表明,双接种菌株Gv和HJ-5显著提高了植株防卫酶如内过氧化物酶(POD)、苯丙氨酸解氨酶(PAL)、几丁质酶等酶的活性,增加了叶片水杨酸和叶绿素含量、降低了植株茎基部胼胝质含量,减轻棉花土传黄萎病发病症状。
综上所述,本文证实了菌株Gv和HJ-5双接种处理具有协同促进植物生长和抑制植物病原菌生长的作用。菌株Gv能够帮助植物吸收土壤中的养分元素,菌株HJ-5具有产生拮抗物质的能力,二者在土壤和植株上的成功定殖是防控棉花土传黄萎病的关键。接种菌株HJ-5和Gv对棉花根系分泌物的改变作用与植物系统抗性有关,这与接种菌株HJ-5和Gv之后植物防御性酶活性等特征的变化是一致的。本文的研究结果有助于研发新型棉花土传黄萎病防控技术。今后应进一步研究生物有机肥中功能菌在修复土壤障碍因子中的分子生物学机制。
Verticillium wilt of cotton caused by fungal pathogen Verticillium dahliae Kleb is a severe disease that limits the development of the cotton industry in the world. While chemical control is challenged by environmental problems and field control practices such as crop rotation and disease-resistant cultivar adoption exists many limting factores, biological control becomes increasingly an important way in control of soilborne plant diseases. Previous reports have shown that arbarscular mycorrhizal fungi (AMF) and Bacillus vallismortis HJ-5have biological control effects against Verticillium wilt of cotton. However, little has been known the effect of dual inoculation of soil with AMF and the bacterium and their synergetic mechanisms. Greenhouse pot experiments were firstly conducted to study several AMF and the inoculation doses of HJ-5for dual inoculation. Three independent greenhouse pot experiments were conducted to study the effects of the combined application of Glomus versiforme (Gv) propagules and HJ-5fortified bio-organic fertilizer on control of Verticillium wilt of cotton. Green fluorescent protein (GFP) labeled strain (GHJ-5) of strain HJ-5was constructed by using green GFP labeling techniques and the ability of GHJ-5to colonize roots and rhizosphere soil of cotton was also studied. To identify the effects of HJ-5and Gv alone or both on root exudates during the incidence of the wilt disease, a split-root experiment was conducted in greenhouse. Additionally, a series of laboratory experiments were performed on observation of the interaction between the plant pathogen and the biocontrol agents. Results were listed as follows.
(1) The primary screening experiments for selection of the best AMF strain and dosage of HJ-5showed that Gv was the suitable AMF strain and the proper innoclation dose of HJ-5was107CFU·g-1. The dual inoculation treatment with Gv and HJ-5had the lowest disease incidence of Verticillium wilt, which was reduced by26.71%as compared with other treatments, and the highest increase of plant biomass.
(2) Inoculation of nursery soil with propagula of Gv and application of the HJ-5enhanced bio-organic fertilizer in the diseased pot soil achieved agreeable high effecenies in suppressing Verticillium wilt by three independent pot experiments. Combined treatment decreased the disease index of Verticillium wilt by65%and increased plant height, leaf area, shoot and root dry weigh by75%,44%,216%, and71%respectively, on the average of three trials as compared with the non-inoculation control. Dual inoculation also significantly increased N in shoot and root and P in shoot and root by83%and65%,49%and45%on the average, respectively, while decreased the number of phytopathogenic Verticillium dahliae in rhizosphere soil of cotton by83%on the average, as compared with the non-inoculation control. Interestingly, the colonization of cotton roots by Gv was not infected by inoculation with HJ-5.
(3) Measurement from laboratory tests showed that the HJ-5was able to produce lipopeptide substances, growth-promoting substances indole acetic acid (IAA), siderophore, hydrogen cyanide (HCN) and ammonia, but not produce deaminase (ACC) and was unable to dissolve P in medium. Colonization, deformation and dissociation of hypha of pathogen V. dahliae Kleb by the green fluorescent protein labelled HJ-5(GHJ-5) was also obsverved, suggesting that HJ-5was an effective antagonist against the pathogen of Verticillium wilt of cotton.
(4) Dual inoculation of soil with GHJ-5and Gv in the greenhouse experiment showed that the cotton plant biomass, root density, plant phosphorus content significantly as well as the number of colonization of the rhizosphere of GHJ-5increased as compared with single inoculation with GHJ-5or Gv, suggesting that synergistic effect existed in promotion of the growth of cotton and suppression of Verticillium wilt of cotton.
(5) In root-splicing experiment, the co-inoculation with HJ-5and Gv reduced disease index by63.3%as compared with the control. Four phenolic acids were found in the cotton root exudates, and all the phenolic acids at low concentrations stimulated germination of V. dahliae spores, while higher concentrations were inhibitory. The phenolic acid concentrations in the root exudates decreased significantly with the application of HJ-5and Gv. The the exudation pattern of cotton roots by HJ-5, Gv and V. dahliae was systemically altered.
(6) In the pot experiment, inoculation of Gv and HJ-5significantly increased the activities of plant defence enzymes such as polyphenoloxidase (PPO), phenylalanine ammonialyase (PAL) and chitinase, increased salicylic acid chlorophyll in plant leaves, and decreased callose in crown of cotton, and consequently relieved the symptoms of Verticillium wilt of cotton.
In conclusion, the present study confirmed that co-inoculation of soil with AMF and rhizosphere bacteria had synergetic promotion of plant growth and resistance to phytopathogens. Abilities of Gv to facilitate plant uptake of nutrients and abilities of HJ-5to produce beneficial secondary metabolics against V. dahliae Kleb as well as successful colonization of soil and plant roots were essential to control Verticillium wilt of cotton. The alteration of the exudation pattern of cotton roots by HJ-5and Gv was related to plant systemic resistance to the phatopathogen. This was also supported by the enhanced activities of plant defence enzymes after inoculation of Gv and HJ-5. Results in this study would be useful in the developping new methods to better control cotton wilt disease. Further researches should be given to the molecular microbilogical aspects of the functions performed by bio-organic fertilizers in remediation of continuously cropping obstacles and in the promotion of plant growth and health exerted by soil functional microbes.
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