稻鸭种养生态系统抑制水稻纹枯病的发生流行规律及拮抗菌SU8生防潜力研究
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摘要
本文拟从植物病害流行学的角度阐述稻鸭种养生态系统抑制水稻纹枯病在田间流行与发生的机理。设置稻鸭种养生态田与常规稻田两个处理,调查两者田间漂浮菌核量与稻株叶鞘菌核附着量变化、观察稻丛基部距水面10cm部位的田间小气候因子动态、测定稻株体内保护性酶的活性动态。研究结果表明,稻鸭种养生态系统中田间漂浮菌核较常规稻田少86%-91%,稻株叶鞘附着菌核少67%～78%。稻鸭种养生态系统稻丛中的相对湿度要显著低于常规稻田,光照强度显著高于常规稻田。从秧苗移栽到水稻分蘖末期,稻鸭种养生态系统中稻丛中的温度较常规稻田低,而从水稻拔节期到灌浆期显著高于常规稻田。稻鸭种养生态系统中稻株内的多酚氧化酶活性较常规稻田低,但丙氨酸解氨酶解氨酶、过氧化物酶、几丁质酶活性要显著高于常规稻田。水稻成熟期病情指数在稻鸭种养生态系统中要比常规稻田低76%,稻谷产量在稻鸭种养生态系统中要比常规稻田高40%。
在上述研究过程中,作者在稻鸭种养生态系统的田间水稻根际土壤中还分离获得了一株对水稻纹枯病菌具有拮抗作用的细菌SU8。因此,进一步对SU8进行了系统研究,评价其作为生防菌的潜力。研究结果如下：①通过形态特征、生理生化和16S rDNA序列测定以及系统进化树构建对拮抗细菌SU8进行鉴定,该菌体呈长杆状,菌体长短均匀,一般长度在1-1.5μmm范围内,革兰氏阴性,其培养性状与生理化特征符合铜绿假单胞菌(Pseudomonas aeruginosa)的相关鉴定描述。其16S rDNA序列提交GenBank数据库,收录号为HQ283487,该序列在系统发育树中与其它已知的铜绿假单胞菌序列的同源性达99%。因此,SU8被鉴定为铜绿假单胞菌(Pseudomonas aeruginosa)。②铜绿假单胞菌株SU8的抑菌谱测定结果显示,该菌株对禾谷镰刀菌(Fusarium graminearum)、烟草黑胫病菌(Phytophthora parasitica)、辣椒炭疽病菌(Colletotrichum capsici)、水稻稻瘟病菌(Magnapor the oryzae)、水稻稻曲病菌(Ustilaginoidea virens)、水稻纹枯病菌(Rhizoctonia solani)、烟草赤星病菌(?)(Alternaria alternata)均具有强拮抗作用。菌株SU8菌悬液及其过滤液均能有效降低水稻纹枯病菌致病力。浓度108cfu/mL的SU8菌悬液对水稻纹枯病的盆栽防效最高可达70%。③对铜绿假单胞菌株SU8过滤液中活性物质的粗提取方法进行了探索,确立了按乙酸乙酯：过滤液1：1,反复萃取3次,合并有机相(即粗提活性物)的提取方法。将合并有机相进行两次硅胶柱层析,得到活性样品Sample7-4。样品为淡黄色针状结晶,经HPLC色谱检测,样品纯度达97%,④对样品Sample7-4进行GC-MS(?)分析,并结合谱库检索得到其化学分子式为：C13H9N3O。根据1H-NMR(300MHz)和13C-NMR(300MHz)检测结果,确定铜绿假单胞菌株SU8抑菌活性物质为吩嗪～(-1)-甲酰胺(Phenazine-1-carboxamide, PCN),并得到其结构式：
根据上述研究结果,表明：①稻鸭种养生态系统抑制水稻纹枯病在田间流行与发生的机理为稻鸭种养生态系统中,田间菌核密度的降低有效控制了水稻纹枯病初侵染源的在田间的扩散,为减少病株率与病情指数提供了条件；田间稻丛中的相对湿度的降低、光照强度的增加、温度动态变化特征形成了不利于水稻纹枯病菌生长的环境,抑制了水稻纹枯病的发生；在稻鸭种养生态系统中水稻植株的抗性蛋白丙氨酸解氨酶、过氧化物酶、几丁质酶的活性在水稻的各个生育期均有提高,对水稻纹枯病的发病与传播起到抑制作用。②铜绿假单胞菌株SU8的抑菌活性物质为吩嗪～(-1)-甲酰胺,是一种广泛存在于假单胞菌代谢产物中的抑制真菌的抗生素,与众多报道的假单胞生防菌的抑菌活性物质相同,说明菌株SU8具有作为生防菌的潜力。同时,抑菌活性物质吩嗪～(-1)-甲酰胺结构式的确立可为开发新型高效、安全、稳定的仿生农药奠定基础。
Researches in this paper were conducted aiming to investigate the epidemical mechanism of Rice-Duck Integrated System (RD) suppressing Rice Sheath Blight. Experiment was carried out between two treatments, rice combined with ducks (RD) and conventional rice field without ducks rearing (CK) in early season rice paddy, to investigate the variations of sclerotia in floodwater and on rice plant, microclimate10cm above the waterline in rice paddy and activity of protective enzymes in rice plants. The results showed that the floating sclerotia in floodwater in RD was86%～91%lower than that in CK, and adhering sclerotia in rice plant in RD was67%～78%lower than that in CK. The relative humidity tested significantly lower and light intensity tested significantly higher in RD. The temperature in the early rice growth stages in RD was slightly lower than that in CK, but it was significantly higher (32.3～36.5℃) in the middle stage rice growth stages. The polyphenoloxidase (PPO) activity in RD were lower than that in CK, but the enhanced activity of phenylalanine ammonia-lyase (PAL), peroxidase (POD) and Chitinase was observed in different stages of rice growth in RD, especially the Chitinase which showed higher activity in all investigating days. At dough stage of growing rice, the disease index in RD was76%lower than that in CK, and the rice yield in RD was40%higher than that in CK.
During the above researches, author isolated anagonistic bacteria strain SU8from rhizoshperic soil in RD, which exhibited superior antifungal activity against Rhizoctonia solani. In the fallowing research, bacteria stain SU8was studied based on morphological, physiological and biochemical characteristics, sequence of16S rDNA genes and phylogenetic analysis. And, in order to ascertain its functioning mechanism in Rhizoctonia solani control, the active compound from the sterilized fermentation liquid was isolated and purified. The molecular structure of the compound was detected through GC-MS,'H-NMR(300MHz) and13C-NMR(300MHz). The results show that:
1. According to the result of morphological research, strain SU8was shaped as long rod, the length ranged1～1.5μm, G negative. The feature in petri dish and its physiobiochemcial characters matched the description of standard Pseudomonas aeruginosa. The family tree based on the16S rDNA sequence showed SU8had99%similarity with other known strains of Pseudomonas aeruginosa, and the sequence was recorded in GeneBank, obtained sequence number HQ283487.
2. According to the test of antibacterial spectrum, strain SU8had inhibitory effect on Fusarium graminearum, Phytophthora parasitica, Colletotrichum capsici, Magnaporthe oryzae, Ustilaginoidea virens, Rhizoctonia solani, Alternaria alternata. Its sterile zymotic liquid was used to evaluate suppressive effect on pathogenicity of two diseases, and pot culture experiments were also conducted to evaluate its control efficiency. And the control efficiencies both reached up to70%on infected rice plants in pot culture. Strain SU8showed superior potential as a bio-control agent in suppressing ShB and RB.
3. Identification of the activity compound in the metabolites of Pseudomonas aeruginosa SU8is the main target of this research. Through the selection of crude extraction method, solvent extraction was selected to be the most effective method by ethyl acetate to sterile zymotic liquid with ratio of1:1,3repetitive operation. Two times of Silica column chromatography were established to obtain the purified Sample-4, with the first elution agent system of petroleum ether:chloroform as ratios of10:1、10:2、10:3、10:4、10:5、10:8、10:10、8:10、5:10,and second elution agent system of petroleum ether:chloroform as ratios of petroleum ether:ethyl acetate as ratios of10:2、10:4、10:6、10:8、10:10、0:1.Through bio-test, Sample7-4was selected out to be active compound, which weighed21.5mg and was light-yellow crystal. HPLC test showed that the purity reached up to97%.
4. Sample7-4was analyzed by GC-MS. According to the database, the molecular formula was given as C13H9N3O. Through the'H-NMR(300MHz) and C-NMR(300MHz) detection, the active compound of Pseudomonas aeruginosa SU8was identified to be Phenazine-1-carboxamide, PCN and the molecular structure:, and acquired spectrum parameters showed highly resemblance with the report by R. Sunish Kumar.
According to the given results,①the density of the sclerotia was considerably decreased, contributing the inhibited proliferation of primary source of infection, which led to the reduction of diseased plant rate and diseased hill rate. In the rice canopy, reduced relative humid, illumination intensity and dynamics of temperature created the condition adverse to the growth of Rhizoctonia solani. And, under the condition of duck intervention, the rice plant cells underwent the physiobiochemical changes which enabled the cells to generate the induced resistance to plant disease, like the enhanced activity of PPO, PLA, POD and Chitinase. Those changes made positive contribution to the suppression of rich sheath blight.②Pseudomonas aeruginosa SU8has broad-spectrum antifungal activity. The control efficiency on rice sheath blight in culture pot indicates strain SU8has the valuable property as an biological control agent, which provides the quality to be used for the development of live bacteria agent. The isolation and purification of the active compound from sterilized fermentation liquid help to give referable information for active compound extraction. And, the identified molecular structure of active compound contributes to ascertaining the antifungal mechanism of strain SU8, which also provides the important information for develop stable, safe and efficient bionic fungicides for rice sheath blight control.
在上述研究过程中,作者在稻鸭种养生态系统的田间水稻根际土壤中还分离获得了一株对水稻纹枯病菌具有拮抗作用的细菌SU8。因此,进一步对SU8进行了系统研究,评价其作为生防菌的潜力。研究结果如下：①通过形态特征、生理生化和16S rDNA序列测定以及系统进化树构建对拮抗细菌SU8进行鉴定,该菌体呈长杆状,菌体长短均匀,一般长度在1-1.5μmm范围内,革兰氏阴性,其培养性状与生理化特征符合铜绿假单胞菌(Pseudomonas aeruginosa)的相关鉴定描述。其16S rDNA序列提交GenBank数据库,收录号为HQ283487,该序列在系统发育树中与其它已知的铜绿假单胞菌序列的同源性达99%。因此,SU8被鉴定为铜绿假单胞菌(Pseudomonas aeruginosa)。②铜绿假单胞菌株SU8的抑菌谱测定结果显示,该菌株对禾谷镰刀菌(Fusarium graminearum)、烟草黑胫病菌(Phytophthora parasitica)、辣椒炭疽病菌(Colletotrichum capsici)、水稻稻瘟病菌(Magnapor the oryzae)、水稻稻曲病菌(Ustilaginoidea virens)、水稻纹枯病菌(Rhizoctonia solani)、烟草赤星病菌(?)(Alternaria alternata)均具有强拮抗作用。菌株SU8菌悬液及其过滤液均能有效降低水稻纹枯病菌致病力。浓度108cfu/mL的SU8菌悬液对水稻纹枯病的盆栽防效最高可达70%。③对铜绿假单胞菌株SU8过滤液中活性物质的粗提取方法进行了探索,确立了按乙酸乙酯：过滤液1：1,反复萃取3次,合并有机相(即粗提活性物)的提取方法。将合并有机相进行两次硅胶柱层析,得到活性样品Sample7-4。样品为淡黄色针状结晶,经HPLC色谱检测,样品纯度达97%,④对样品Sample7-4进行GC-MS(?)分析,并结合谱库检索得到其化学分子式为：C13H9N3O。根据1H-NMR(300MHz)和13C-NMR(300MHz)检测结果,确定铜绿假单胞菌株SU8抑菌活性物质为吩嗪～(-1)-甲酰胺(Phenazine-1-carboxamide, PCN),并得到其结构式：
根据上述研究结果,表明：①稻鸭种养生态系统抑制水稻纹枯病在田间流行与发生的机理为稻鸭种养生态系统中,田间菌核密度的降低有效控制了水稻纹枯病初侵染源的在田间的扩散,为减少病株率与病情指数提供了条件；田间稻丛中的相对湿度的降低、光照强度的增加、温度动态变化特征形成了不利于水稻纹枯病菌生长的环境,抑制了水稻纹枯病的发生；在稻鸭种养生态系统中水稻植株的抗性蛋白丙氨酸解氨酶、过氧化物酶、几丁质酶的活性在水稻的各个生育期均有提高,对水稻纹枯病的发病与传播起到抑制作用。②铜绿假单胞菌株SU8的抑菌活性物质为吩嗪～(-1)-甲酰胺,是一种广泛存在于假单胞菌代谢产物中的抑制真菌的抗生素,与众多报道的假单胞生防菌的抑菌活性物质相同,说明菌株SU8具有作为生防菌的潜力。同时,抑菌活性物质吩嗪～(-1)-甲酰胺结构式的确立可为开发新型高效、安全、稳定的仿生农药奠定基础。
Researches in this paper were conducted aiming to investigate the epidemical mechanism of Rice-Duck Integrated System (RD) suppressing Rice Sheath Blight. Experiment was carried out between two treatments, rice combined with ducks (RD) and conventional rice field without ducks rearing (CK) in early season rice paddy, to investigate the variations of sclerotia in floodwater and on rice plant, microclimate10cm above the waterline in rice paddy and activity of protective enzymes in rice plants. The results showed that the floating sclerotia in floodwater in RD was86%～91%lower than that in CK, and adhering sclerotia in rice plant in RD was67%～78%lower than that in CK. The relative humidity tested significantly lower and light intensity tested significantly higher in RD. The temperature in the early rice growth stages in RD was slightly lower than that in CK, but it was significantly higher (32.3～36.5℃) in the middle stage rice growth stages. The polyphenoloxidase (PPO) activity in RD were lower than that in CK, but the enhanced activity of phenylalanine ammonia-lyase (PAL), peroxidase (POD) and Chitinase was observed in different stages of rice growth in RD, especially the Chitinase which showed higher activity in all investigating days. At dough stage of growing rice, the disease index in RD was76%lower than that in CK, and the rice yield in RD was40%higher than that in CK.
During the above researches, author isolated anagonistic bacteria strain SU8from rhizoshperic soil in RD, which exhibited superior antifungal activity against Rhizoctonia solani. In the fallowing research, bacteria stain SU8was studied based on morphological, physiological and biochemical characteristics, sequence of16S rDNA genes and phylogenetic analysis. And, in order to ascertain its functioning mechanism in Rhizoctonia solani control, the active compound from the sterilized fermentation liquid was isolated and purified. The molecular structure of the compound was detected through GC-MS,'H-NMR(300MHz) and13C-NMR(300MHz). The results show that:
1. According to the result of morphological research, strain SU8was shaped as long rod, the length ranged1～1.5μm, G negative. The feature in petri dish and its physiobiochemcial characters matched the description of standard Pseudomonas aeruginosa. The family tree based on the16S rDNA sequence showed SU8had99%similarity with other known strains of Pseudomonas aeruginosa, and the sequence was recorded in GeneBank, obtained sequence number HQ283487.
2. According to the test of antibacterial spectrum, strain SU8had inhibitory effect on Fusarium graminearum, Phytophthora parasitica, Colletotrichum capsici, Magnaporthe oryzae, Ustilaginoidea virens, Rhizoctonia solani, Alternaria alternata. Its sterile zymotic liquid was used to evaluate suppressive effect on pathogenicity of two diseases, and pot culture experiments were also conducted to evaluate its control efficiency. And the control efficiencies both reached up to70%on infected rice plants in pot culture. Strain SU8showed superior potential as a bio-control agent in suppressing ShB and RB.
3. Identification of the activity compound in the metabolites of Pseudomonas aeruginosa SU8is the main target of this research. Through the selection of crude extraction method, solvent extraction was selected to be the most effective method by ethyl acetate to sterile zymotic liquid with ratio of1:1,3repetitive operation. Two times of Silica column chromatography were established to obtain the purified Sample-4, with the first elution agent system of petroleum ether:chloroform as ratios of10:1、10:2、10:3、10:4、10:5、10:8、10:10、8:10、5:10,and second elution agent system of petroleum ether:chloroform as ratios of petroleum ether:ethyl acetate as ratios of10:2、10:4、10:6、10:8、10:10、0:1.Through bio-test, Sample7-4was selected out to be active compound, which weighed21.5mg and was light-yellow crystal. HPLC test showed that the purity reached up to97%.
4. Sample7-4was analyzed by GC-MS. According to the database, the molecular formula was given as C13H9N3O. Through the'H-NMR(300MHz) and C-NMR(300MHz) detection, the active compound of Pseudomonas aeruginosa SU8was identified to be Phenazine-1-carboxamide, PCN and the molecular structure:, and acquired spectrum parameters showed highly resemblance with the report by R. Sunish Kumar.
According to the given results,①the density of the sclerotia was considerably decreased, contributing the inhibited proliferation of primary source of infection, which led to the reduction of diseased plant rate and diseased hill rate. In the rice canopy, reduced relative humid, illumination intensity and dynamics of temperature created the condition adverse to the growth of Rhizoctonia solani. And, under the condition of duck intervention, the rice plant cells underwent the physiobiochemical changes which enabled the cells to generate the induced resistance to plant disease, like the enhanced activity of PPO, PLA, POD and Chitinase. Those changes made positive contribution to the suppression of rich sheath blight.②Pseudomonas aeruginosa SU8has broad-spectrum antifungal activity. The control efficiency on rice sheath blight in culture pot indicates strain SU8has the valuable property as an biological control agent, which provides the quality to be used for the development of live bacteria agent. The isolation and purification of the active compound from sterilized fermentation liquid help to give referable information for active compound extraction. And, the identified molecular structure of active compound contributes to ascertaining the antifungal mechanism of strain SU8, which also provides the important information for develop stable, safe and efficient bionic fungicides for rice sheath blight control.
引文
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