根际施用生物有机肥防控西瓜土传枯萎病效果及机理研究
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摘要
西瓜土传枯萎病是西瓜连作生物障碍的主要病害之一,严重影响西瓜产业的发展,研究以生物防控为主导的综合防控具有重要意义。本论文以具有自主知识产权的三株能强烈抑制西瓜土传枯萎病病原真菌(尖孢镰刀菌西瓜专化型,Fusarium. oxysporum f.sp. niveum)的拮抗菌株(Paenibacillus polymyxa SQR21; Bacillus subtillus SQR-9; Trichoderma harzianum SQR-T037)为材料,经液体发酵后与合适的有机载体进行二次固体发酵,制成了西瓜专用生物有机肥。采用大田试验、盆栽试验和室内分析相结合的方法,研究了根际施用这些生物有机肥对西瓜连作土传枯萎病的防控效果、拮抗菌P. polymyxa SQR21(SQR21)与病原菌在西瓜根际和根表上的行为特征及与西瓜根系分泌物之间的相互影响机制。获得了如下主要结果：
1、营养钵施用西瓜专用生物有机肥(BIO)能显著增加西瓜苗的生物量,施用BIO的西瓜地上部鲜重分别为施用有机肥(OF)和施用化肥(CK)的1.2倍和2.1倍,西瓜地上部干重分别是施用OF和CK的1.2倍和1.7倍。施用BIO的西瓜植株根系生长最茂盛,CK次之,OF根系生长最差；各处理间西瓜总根长、根表面积、根体积及根尖差异显著,呈现的趋势顺序均为：BIO>CK>OF；各处理的根系活力也呈现BIO>OF>CK的趋势。施用BIO处理的西瓜根际土壤中的细菌和放线菌的数量均显著高于处理OF和CK,而真菌数量显著低于处理OF和CK。营养钵施用BIO能有效增加西瓜苗的净光合作用,可显著提高西瓜叶片中光合色素的含量,改善西瓜叶片的光合能力。
2、采用营养钵与移栽土壤双重施用BIO的模式对降低土传枯萎病的发生率效果显著,其防控效果可以达到73%以上。营养钵施用BIO的模式不仅能提高BIO的生防效果,还能促进西瓜植株生长,提高西瓜产量。该施肥模式有效促使BIO肥料中的生防菌在苗期就定殖于西瓜根际和根表,并随西瓜根系的不断生长而延伸,从而有效降低西瓜根际病原菌数量,使西瓜根系免受病原菌侵害。
3、拮抗菌株SQR21能在西瓜根部较好地定殖,随西瓜根系的生长在根际不断生长繁殖,在西瓜根尖表面形成一层生物膜,从而保护西瓜根部免受病原菌的侵害。揭示了生物有机肥中拮抗菌SQR21在土壤及西瓜根部的定殖及消长规律,为多粘芽孢杆菌SQR21的田间应用及其作用机制研究提供理论依据。
4、与对照相比,施用BIO肥料能够显著增加SQR21和降低FON在根际的数量；施用BIO处理的根际SQR21数量最大拷贝数为7.14lg拷贝/g土,离西瓜根系越远SQR21的拷贝数越低；对照中FON的数量比BIO处理高2个数量级,随着与根系间距的加大,根系接种SQR21处理的FON数量逐渐增加,而未接种SQR21处理的结果则相反。土体土壤由于被侵染土壤中FON背景值的存在,各处理间FON及SQR21的数量没有显著差异。以上结果表明,施用生物有机肥促进了SQR21在根系的定殖,大幅度地降低了FON的数量,从而有效地抑制了土传西瓜枯萎病的发生。
5、西瓜根系分泌物中的多种有机酸对多粘类芽孢杆菌SQR21具有趋化作用。在西瓜根系分泌物中检测到大量的三羧酸循环过程的中间产物,如：草酸、苹果酸、柠檬酸,其中苹果酸和柠檬酸对多粘类芽孢杆菌SQR21均有显著的趋化作用。三种有机酸中苹果酸的诱导效果最为显著,趋化性指数为3.9,群游菌圈是对照的1.5倍。另外,体外实验进一步证实了这些有机酸对多粘芽孢杆菌SQR21具有吸引作用。
6、与对照相比,接种SQR21的西瓜根系分泌物会减少FON分生孢子的萌发,而接种FON的西瓜根系分泌物则促进FON分生孢子的萌发。接种FON30天后,西瓜分泌物对分生孢子萌发率达到最大值,为对照的1.4倍；其根系分泌物中肉桂酸浓度为0-30μg/ml时,肉桂酸对FON分生孢子萌发有显著的刺激效应。不论是接种SQR21还是FON,西瓜根系分泌物的改变均具有局部性和系统性特性。
The bioorganic fertilizer (BIO) containing Paenibacillus polymyxa SQR21, Bacillus subtillus SQR-9and Trichoderma harzianum SQR-T037has been shown to suppress Fusarium wilt disease, a severe soil borne disease, and to promote growth of watermelon and cucumber plants in greenhouse and field experiments. The present study was carried out to develop a mode of application of bioorganic fertilizer for improving the biocontrolling efficacy to Fusarium wilt. The interactions that occur between plant roots and the antagonistic microbe (P. polymyxa SQR21) were in detail studied to provide guidelines of how to use biocontrol agents more effectively in actual field situations. The results were obtained as follows:
1. Nursery application of BIO could significantly increase the biomass of the watermelon plants compared with the treatments applied with organic fertilizer (OF) or with chemical fertilizer (CK). The fresh shoot weights of watermelon plant in BIO treatment were1.24and2.08times higher than both of OF and CK, and the dry root weights of watermelon plant in BIO treatment were1.22and1.67times higher than OF and CK, respectively; The length, surface area, volume, tips and average diam of roots were significantly increased by the application of BIO; The numbers of bacteria and actinomycetes were significantly increased by the application of BIO compared to the OF and CK, and the number of fungi was significantly decreased by application of BIO; Nursery application of BIO could effectively increase the photosynthetic capacity of watermelon leaves by increasing the content of photosynthetic pigments in watermelon leaves.
2. Pot and field experiments were carried out to evaluate the efficacy of the BIO application on control of watermelon Fusarium wilt.1) The new mode of nursery application of BIO could significantly decrease the Fusarium wilt incidence, and the best biocontrol efficacy could be achieved by applying BIO in the nursery stage plus application in the transplanted soil.2) The way of BIO application could not only suppress the Fusarium disease, but also increase the biomass of watermelon and thus increase the yields. This could be attributed to the fact that nursery application of BIO could ensure the efficient colonization of the antagonist at the seedling stage and decrease the number of the pathogen in the rhizosphere after transplanting.
3. The methods of color reaction in watermelon rhizosphere, GFP-labeling, petriplate cuture count were used to investigate the colonization-ability of SQR21in watermelon rhizosphere or on rhizoplane. SQR21could effectively colonize on the watermelon roots, propagate in rhizosphere as the growth of roots, and form biofilm on the surface of root tips of watermelon plants. These results indicated that SQR21could protect the watermelon roots from invasion by pathogenetic fungi, and could also support some basic theory for the research of mechanisms on biocontrol of Fusarium wilt by SQR21.
4. A rhizobox experiment was carried out to investigate the distribution of F. oxysporum f.sp. niveum (FON) and its antagonistic bacterium (SQR21), in the soil at different distances away from the watermelon rhizoplane by real-time PCR. The results confirmed that BIO application significantly suppressed Fusarium wilt and promoted the growth of the watermelon plants. Application of BIO significantly enhanced the abundance of SQR21and reduced the population of FON in the soil near the rhizoplane as compared to the control. A maximal quantity (7.14lgcopies/g soil) of P. polymyxa SQR21was obtained in the rhizosphere under the BIO treatment, and this number was decreased with increasing distance away from the rhizoplane. The population of FON in the rhizosphere of the control was2orders of magnitude higher than under the BIO treatment, and with increasing distance away from the rhizoplane, the FON abundance increased in the SQR21inoculated treatments and decreased in the treatments without SQR21. In the soil far away from the rhizoplane (bulk soil), both FON and SQR21reached a level that was not statistically different between the treatments, which might be attributed to the background level of the infested soil. These results revealed that the BIO application with the developed method suppressed Fusarium wilt by the promotion of SQR21colonization in the rhizosphere, which largely reduced the population of the pathogen.
5. Some organic acids were significantly detected in the root exudates of watermelon. Chemotaxis and swarming assays were performed to investigate the ability of these organic acids to induce the motility of P. polymyxa SQR21. The results showed that oxalic acid, malic acid and citric acid were present in the root exudates but only malic acid and citric acid could significantly induce motility in P. polymyxa SQR21. The maximal inducing ability was obtained from malic acid. Values for malic acid were3.9and1.5times higher than the control in the chemotaxis assay and the swarming assay, respectively. An in vitro experiment further confirmed that these organic acids could promote recruitment to P. polymyxa SQR21, thereby increasing the population in the rhizosphere.
6. Conidial germination of FON decreased in the presence of root exudates from SQR21inoculated plants, while it was enhanced in the present of root exudates from FON inoculated plants as compared with control. Maximum germination was found in the root exudates from FON inoculated plant after30days, which was1.35times more than control. A split-root system was designed to verify the fact that the alterations of the exudation pattern in SQR21or FON inoculated watermelon roots were not only local, but also systemic. Eleven kinds of phenolic acids were detected in the root exudates by HPLC. The changes of cinnamic acid concentrations were positively related to the conidial germination. Assay of cinnamic acid on conidial germination of FON revealed that relatively high concentration of this acid stimulated the conidial germination, and vice versa. All of these results will help in the better understanding of the plant-microbe communication and will guide to improve the biocontrol strategies against Fusarium wilt of watermelon plants.
1、营养钵施用西瓜专用生物有机肥(BIO)能显著增加西瓜苗的生物量,施用BIO的西瓜地上部鲜重分别为施用有机肥(OF)和施用化肥(CK)的1.2倍和2.1倍,西瓜地上部干重分别是施用OF和CK的1.2倍和1.7倍。施用BIO的西瓜植株根系生长最茂盛,CK次之,OF根系生长最差；各处理间西瓜总根长、根表面积、根体积及根尖差异显著,呈现的趋势顺序均为：BIO>CK>OF；各处理的根系活力也呈现BIO>OF>CK的趋势。施用BIO处理的西瓜根际土壤中的细菌和放线菌的数量均显著高于处理OF和CK,而真菌数量显著低于处理OF和CK。营养钵施用BIO能有效增加西瓜苗的净光合作用,可显著提高西瓜叶片中光合色素的含量,改善西瓜叶片的光合能力。
2、采用营养钵与移栽土壤双重施用BIO的模式对降低土传枯萎病的发生率效果显著,其防控效果可以达到73%以上。营养钵施用BIO的模式不仅能提高BIO的生防效果,还能促进西瓜植株生长,提高西瓜产量。该施肥模式有效促使BIO肥料中的生防菌在苗期就定殖于西瓜根际和根表,并随西瓜根系的不断生长而延伸,从而有效降低西瓜根际病原菌数量,使西瓜根系免受病原菌侵害。
3、拮抗菌株SQR21能在西瓜根部较好地定殖,随西瓜根系的生长在根际不断生长繁殖,在西瓜根尖表面形成一层生物膜,从而保护西瓜根部免受病原菌的侵害。揭示了生物有机肥中拮抗菌SQR21在土壤及西瓜根部的定殖及消长规律,为多粘芽孢杆菌SQR21的田间应用及其作用机制研究提供理论依据。
4、与对照相比,施用BIO肥料能够显著增加SQR21和降低FON在根际的数量；施用BIO处理的根际SQR21数量最大拷贝数为7.14lg拷贝/g土,离西瓜根系越远SQR21的拷贝数越低；对照中FON的数量比BIO处理高2个数量级,随着与根系间距的加大,根系接种SQR21处理的FON数量逐渐增加,而未接种SQR21处理的结果则相反。土体土壤由于被侵染土壤中FON背景值的存在,各处理间FON及SQR21的数量没有显著差异。以上结果表明,施用生物有机肥促进了SQR21在根系的定殖,大幅度地降低了FON的数量,从而有效地抑制了土传西瓜枯萎病的发生。
5、西瓜根系分泌物中的多种有机酸对多粘类芽孢杆菌SQR21具有趋化作用。在西瓜根系分泌物中检测到大量的三羧酸循环过程的中间产物,如：草酸、苹果酸、柠檬酸,其中苹果酸和柠檬酸对多粘类芽孢杆菌SQR21均有显著的趋化作用。三种有机酸中苹果酸的诱导效果最为显著,趋化性指数为3.9,群游菌圈是对照的1.5倍。另外,体外实验进一步证实了这些有机酸对多粘芽孢杆菌SQR21具有吸引作用。
6、与对照相比,接种SQR21的西瓜根系分泌物会减少FON分生孢子的萌发,而接种FON的西瓜根系分泌物则促进FON分生孢子的萌发。接种FON30天后,西瓜分泌物对分生孢子萌发率达到最大值,为对照的1.4倍；其根系分泌物中肉桂酸浓度为0-30μg/ml时,肉桂酸对FON分生孢子萌发有显著的刺激效应。不论是接种SQR21还是FON,西瓜根系分泌物的改变均具有局部性和系统性特性。
The bioorganic fertilizer (BIO) containing Paenibacillus polymyxa SQR21, Bacillus subtillus SQR-9and Trichoderma harzianum SQR-T037has been shown to suppress Fusarium wilt disease, a severe soil borne disease, and to promote growth of watermelon and cucumber plants in greenhouse and field experiments. The present study was carried out to develop a mode of application of bioorganic fertilizer for improving the biocontrolling efficacy to Fusarium wilt. The interactions that occur between plant roots and the antagonistic microbe (P. polymyxa SQR21) were in detail studied to provide guidelines of how to use biocontrol agents more effectively in actual field situations. The results were obtained as follows:
1. Nursery application of BIO could significantly increase the biomass of the watermelon plants compared with the treatments applied with organic fertilizer (OF) or with chemical fertilizer (CK). The fresh shoot weights of watermelon plant in BIO treatment were1.24and2.08times higher than both of OF and CK, and the dry root weights of watermelon plant in BIO treatment were1.22and1.67times higher than OF and CK, respectively; The length, surface area, volume, tips and average diam of roots were significantly increased by the application of BIO; The numbers of bacteria and actinomycetes were significantly increased by the application of BIO compared to the OF and CK, and the number of fungi was significantly decreased by application of BIO; Nursery application of BIO could effectively increase the photosynthetic capacity of watermelon leaves by increasing the content of photosynthetic pigments in watermelon leaves.
2. Pot and field experiments were carried out to evaluate the efficacy of the BIO application on control of watermelon Fusarium wilt.1) The new mode of nursery application of BIO could significantly decrease the Fusarium wilt incidence, and the best biocontrol efficacy could be achieved by applying BIO in the nursery stage plus application in the transplanted soil.2) The way of BIO application could not only suppress the Fusarium disease, but also increase the biomass of watermelon and thus increase the yields. This could be attributed to the fact that nursery application of BIO could ensure the efficient colonization of the antagonist at the seedling stage and decrease the number of the pathogen in the rhizosphere after transplanting.
3. The methods of color reaction in watermelon rhizosphere, GFP-labeling, petriplate cuture count were used to investigate the colonization-ability of SQR21in watermelon rhizosphere or on rhizoplane. SQR21could effectively colonize on the watermelon roots, propagate in rhizosphere as the growth of roots, and form biofilm on the surface of root tips of watermelon plants. These results indicated that SQR21could protect the watermelon roots from invasion by pathogenetic fungi, and could also support some basic theory for the research of mechanisms on biocontrol of Fusarium wilt by SQR21.
4. A rhizobox experiment was carried out to investigate the distribution of F. oxysporum f.sp. niveum (FON) and its antagonistic bacterium (SQR21), in the soil at different distances away from the watermelon rhizoplane by real-time PCR. The results confirmed that BIO application significantly suppressed Fusarium wilt and promoted the growth of the watermelon plants. Application of BIO significantly enhanced the abundance of SQR21and reduced the population of FON in the soil near the rhizoplane as compared to the control. A maximal quantity (7.14lgcopies/g soil) of P. polymyxa SQR21was obtained in the rhizosphere under the BIO treatment, and this number was decreased with increasing distance away from the rhizoplane. The population of FON in the rhizosphere of the control was2orders of magnitude higher than under the BIO treatment, and with increasing distance away from the rhizoplane, the FON abundance increased in the SQR21inoculated treatments and decreased in the treatments without SQR21. In the soil far away from the rhizoplane (bulk soil), both FON and SQR21reached a level that was not statistically different between the treatments, which might be attributed to the background level of the infested soil. These results revealed that the BIO application with the developed method suppressed Fusarium wilt by the promotion of SQR21colonization in the rhizosphere, which largely reduced the population of the pathogen.
5. Some organic acids were significantly detected in the root exudates of watermelon. Chemotaxis and swarming assays were performed to investigate the ability of these organic acids to induce the motility of P. polymyxa SQR21. The results showed that oxalic acid, malic acid and citric acid were present in the root exudates but only malic acid and citric acid could significantly induce motility in P. polymyxa SQR21. The maximal inducing ability was obtained from malic acid. Values for malic acid were3.9and1.5times higher than the control in the chemotaxis assay and the swarming assay, respectively. An in vitro experiment further confirmed that these organic acids could promote recruitment to P. polymyxa SQR21, thereby increasing the population in the rhizosphere.
6. Conidial germination of FON decreased in the presence of root exudates from SQR21inoculated plants, while it was enhanced in the present of root exudates from FON inoculated plants as compared with control. Maximum germination was found in the root exudates from FON inoculated plant after30days, which was1.35times more than control. A split-root system was designed to verify the fact that the alterations of the exudation pattern in SQR21or FON inoculated watermelon roots were not only local, but also systemic. Eleven kinds of phenolic acids were detected in the root exudates by HPLC. The changes of cinnamic acid concentrations were positively related to the conidial germination. Assay of cinnamic acid on conidial germination of FON revealed that relatively high concentration of this acid stimulated the conidial germination, and vice versa. All of these results will help in the better understanding of the plant-microbe communication and will guide to improve the biocontrol strategies against Fusarium wilt of watermelon plants.
引文
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