果实采后病害拮抗菌的分离筛选及其抑菌效果的研究
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摘要
柑橘青霉病和梨黑斑病是柑橘、梨果实贮藏期的主要侵染性病害,是造成腐烂损失的重要原因。长期以来,使用化学杀菌剂一直被认为是防治果实采后病害最有效的方法,但是存在着农药残留、环境污染和病原菌抗药性等诸多弊端,迫切需要新的安全的保鲜方法来取代化学杀菌剂的使用。生物防治是选择对产品以及人体不造成危害的微生物,利用微生物之间的拮抗作用,抑制病原菌生长的一种有效防治采后病害的新途径。
本研究旨在分离、筛选能够抑制果实采后青霉、链格孢霉的拮抗菌株,鉴定其生物学特性,并测定所筛选出的菌株YF1,YF4和BF2及不同混配处理对柑橘青霉病和梨黑斑病的抑制效果,以及在果实伤口的定殖情况,并对其保鲜机理进行初步探讨。主要结果如下:
1.拮抗菌的分离和筛选:用稀释平板法,从果实伤口、果实表面、叶片以及果园根际土壤中分离得到102个菌株。应用平板对峙法,通过初筛和复筛,离体筛选出10个菌株对青霉菌(Penicillium italicum)有拮抗作用,9个菌株对链格孢霉(Alternaria kikuchiana)有拮抗作用。进一步通过果实接种法,活体筛选出分别能明显抑制青霉和链格孢霉的拮抗菌YF1,YF2和BF2,室温(约25℃)下抑菌率分别为79.12%,62.5%和66.67%。
2.拮抗菌的鉴定:根据酵母菌的形态特征、营养特征和生理特性,鉴定拮抗菌YF1为假丝酵母属(Candida sp.),YF4为柠檬形克勒克酵母(Kloeckera apiculata)。在对拮抗菌BF2的鉴定中,首先采用细菌传统的鉴定方法,对菌体的表型、生理生化特征进行了分类鉴定,结果发现,BF2符合芽孢杆菌属细菌的特征。进而采用16S rDNA分子鉴定方法,通过同源性比对及系统发育分析,鉴定拮抗菌BF2为芽孢杆菌属的蜡样芽孢杆菌(Bacillus cereus)。
3.拮抗菌的抑菌试验:当拮抗菌浓度为106-108 CFU/mL时,生物防治的效果与拮抗菌的浓度成正相关;分别在接种1×108 CFU/mL拮抗菌YF1、YF4和BF2的前48 h, 24 h, 2 h, 0 h、后2 h, 24 h, 48 h接种病菌,同时和推后病菌接种时间,都可以维持抑菌效果,但是提前接种病菌拮抗菌的抑菌力会有所降低。拮抗菌的接种时间越早防治病害的效果越明显;YF1与YF4等体积混合用于防治青霉病,果实发病率为12.50%,优于YF1和YF4单独使用的效果,差异显著; YF1, YF4和BF2两两混合使用抑制梨黑斑病,均可显著抑制梨黑斑病的发生,发病率分别为17.5%, 10.00%和15.83%,低于单独使用一种拮抗菌; CaCl2和低剂量化学杀菌剂能够明显增强拮抗菌的抑菌效果,水杨酸与拮抗菌结合使用与单独使用拮抗菌相比,对柑橘青霉病和梨黑斑病发病率的影响不大。
4.拮抗菌的生长动态:拮抗酵母菌YF1和YF4在有无青霉菌和链格孢霉菌作用的情况下都能够在果实伤口迅速生长定殖。菌株YF1接种48 h后,数量增加123.26倍;同时接种青霉病菌,96 h后YF1的数量可增加至144.17倍,与链格孢霉同时接种,96 h后YF1数量增加至178.29倍;拮抗菌YF4接种72 h后,数量增加了近200倍,同时接种青霉病菌,72 h后繁殖数量是起始值的344.54倍,同时接种链格孢霉,72 h后繁殖数量是起始值的294.12倍。
5.用1×108 CFU/mL的酵母菌YF1和YF4浸泡柑橘果实后,柑橘果皮的PAL(苯丙氨酸解氨酶)、POD(过氧化物酶)显著升高,PPO(多酚氧化酶)显著降低;在柑橘果实的伤口处添加青霉(Penicillium italicum)的孢子悬浮液可以显著诱导柑橘产生β-1, 3-葡聚糖酶和几丁质酶活性。说明两种酵母菌都能诱导柑橘果实产生抗病性。
Blue mold and black spot rot are important postharvest diseases of citrus and pear fruits. Pathogens attacks are main causes of postharvest losses. Generally, synthetic fungicide was considered to be an effective means for controlling postharvest diseases. However, its disadvantages such as chemical residues in foods, environmental pollution, and the occurrence of fungicide-tolerant strains of pathogens, have accelerated the development of alternative approaches. Among these, biological control using harmless antagonistic microorganisms was proved to be a promising strategy for managing fruit diseases.
The aim of this research is to isolate and screen effective antagonists against blue mold and pear black spot rot, and identify them. The inhibition effects and their population dynamics in wounds were evaluated after the antagonists were screened. Main conclusions were drawn as follows.
1. There were 102 strains isolated from the wounds and the surface of fruits, leaves, and rhizosphere soil using the method of serial dilution. After primary and further screening in vitro, 10 strains exhibited inhibitory activity to Penicillium italicum and 9 strains exhibited inhibitory activity to Alternaria kikuchiana. Among screened strains, it was found that the isolates YF1, YF4 and BF2 were effective antagonists against P. italicum and A. kikuchiana on fruits through Fruits-inoculation test. The inhibition rate of isolates YF1, YF4 and BF2 got to 79.12%, 62.5% and 66.67% at 25℃.
2. Identification of the antagonists: According to the morphological, nutritional and physiological character, we found YF1 belonged to Candida sp., and YF4 belonged to Kloeckera apiculata. In order to determine the taxonomic placement, the identification of BF2 was performed with traditional bacterial systemic identification method. And then the molecular identification of BF2 was performed. Through the comparative 16S rDNA sequences analysis and phylogenetic analysis, we found BF2 belonged to Bacillus cereus.
3. The effects of antagonists increased with the increase of antagonists cell population when the antagonists at the concentrations of 106-108 CFU/mL. Fruits were treated with antagonists suspension at 1×108 CFU/mL at various time intervals (48 h, 24 h, 2 h, 0 h) before and after challenge-inoculated with pathogenic spore suspension at 104 spores/mL. The efficacy of antagonists against A. kikuchiana and P. italicum was maintained when it was applied simultaneously with or before the pathogens. However, the efficacy decreased when applied after the spores. The result of inoculation-time test showed that the efficacy of biocontrol increased significantly with the time antagonists applied before pathogen inoculated. In mixed-antagonists inoculation test, the blue mold infection rate was reduced to 12.50% when the two yeasts, YF1 and YF4, suspensions were applied simultaneously. This meant that the efficacy of the mixture excelled they applied individually and indicated significant discrepancy between them. The infection rates of black spot rot were 17.5%, 10.00% and 15.83% when isolates YF1, YF4 and BF2 were mixed with each other. It suggested that the efficacy of the two mixtures was better than that of YF1, YF4 and BF2. In the antangonists-enhancement test, the result showed that CaCl2 and low synthetic fungicide elevated biocontrol efficacy of antagonists. But when antagonists combined with SA, there is no different from antagonist used alone.
4. Rapid colonization of isolates YF1 and YF4 in wounds was observed whether pathogenic spores inoculated. The population of YF1 increased 123.26 folds after 48 h. When P. italicum spores inoculated simultaneously, the population increased 144.17 folds after 96 h. The population of YF4 increased almost 200 folds after 72 h. While it was 344.54 folds compared with the start point at 72 h when P. italicum spores co-inoculated, and it was 294.12 folds compared with the start point at 72 h when A. kikuchiana spores co-inoculated.
5. When the citrus fruits were soaked in the two yeast cell suspensions of 1×108 CFU/mL, the activities of phenlalanine ammonialyase (PAL) and peroxidase (POD) in fruits of citrus fruits raised remarkably, while polyphenol oxidase (PPO) reduced prominently and activities ofβ-1,3-glucanase and exochitiase almost did not change. But when P. italicum and the two yeasts added into the wounds of the fruit simultaneously, the activities ofβ-1, 3-glucanase and exochitiase remarkably raised by being induced. These indicate that both of the yeasts can induce diseases resistance in citrus.
本研究旨在分离、筛选能够抑制果实采后青霉、链格孢霉的拮抗菌株,鉴定其生物学特性,并测定所筛选出的菌株YF1,YF4和BF2及不同混配处理对柑橘青霉病和梨黑斑病的抑制效果,以及在果实伤口的定殖情况,并对其保鲜机理进行初步探讨。主要结果如下:
1.拮抗菌的分离和筛选:用稀释平板法,从果实伤口、果实表面、叶片以及果园根际土壤中分离得到102个菌株。应用平板对峙法,通过初筛和复筛,离体筛选出10个菌株对青霉菌(Penicillium italicum)有拮抗作用,9个菌株对链格孢霉(Alternaria kikuchiana)有拮抗作用。进一步通过果实接种法,活体筛选出分别能明显抑制青霉和链格孢霉的拮抗菌YF1,YF2和BF2,室温(约25℃)下抑菌率分别为79.12%,62.5%和66.67%。
2.拮抗菌的鉴定:根据酵母菌的形态特征、营养特征和生理特性,鉴定拮抗菌YF1为假丝酵母属(Candida sp.),YF4为柠檬形克勒克酵母(Kloeckera apiculata)。在对拮抗菌BF2的鉴定中,首先采用细菌传统的鉴定方法,对菌体的表型、生理生化特征进行了分类鉴定,结果发现,BF2符合芽孢杆菌属细菌的特征。进而采用16S rDNA分子鉴定方法,通过同源性比对及系统发育分析,鉴定拮抗菌BF2为芽孢杆菌属的蜡样芽孢杆菌(Bacillus cereus)。
3.拮抗菌的抑菌试验:当拮抗菌浓度为106-108 CFU/mL时,生物防治的效果与拮抗菌的浓度成正相关;分别在接种1×108 CFU/mL拮抗菌YF1、YF4和BF2的前48 h, 24 h, 2 h, 0 h、后2 h, 24 h, 48 h接种病菌,同时和推后病菌接种时间,都可以维持抑菌效果,但是提前接种病菌拮抗菌的抑菌力会有所降低。拮抗菌的接种时间越早防治病害的效果越明显;YF1与YF4等体积混合用于防治青霉病,果实发病率为12.50%,优于YF1和YF4单独使用的效果,差异显著; YF1, YF4和BF2两两混合使用抑制梨黑斑病,均可显著抑制梨黑斑病的发生,发病率分别为17.5%, 10.00%和15.83%,低于单独使用一种拮抗菌; CaCl2和低剂量化学杀菌剂能够明显增强拮抗菌的抑菌效果,水杨酸与拮抗菌结合使用与单独使用拮抗菌相比,对柑橘青霉病和梨黑斑病发病率的影响不大。
4.拮抗菌的生长动态:拮抗酵母菌YF1和YF4在有无青霉菌和链格孢霉菌作用的情况下都能够在果实伤口迅速生长定殖。菌株YF1接种48 h后,数量增加123.26倍;同时接种青霉病菌,96 h后YF1的数量可增加至144.17倍,与链格孢霉同时接种,96 h后YF1数量增加至178.29倍;拮抗菌YF4接种72 h后,数量增加了近200倍,同时接种青霉病菌,72 h后繁殖数量是起始值的344.54倍,同时接种链格孢霉,72 h后繁殖数量是起始值的294.12倍。
5.用1×108 CFU/mL的酵母菌YF1和YF4浸泡柑橘果实后,柑橘果皮的PAL(苯丙氨酸解氨酶)、POD(过氧化物酶)显著升高,PPO(多酚氧化酶)显著降低;在柑橘果实的伤口处添加青霉(Penicillium italicum)的孢子悬浮液可以显著诱导柑橘产生β-1, 3-葡聚糖酶和几丁质酶活性。说明两种酵母菌都能诱导柑橘果实产生抗病性。
Blue mold and black spot rot are important postharvest diseases of citrus and pear fruits. Pathogens attacks are main causes of postharvest losses. Generally, synthetic fungicide was considered to be an effective means for controlling postharvest diseases. However, its disadvantages such as chemical residues in foods, environmental pollution, and the occurrence of fungicide-tolerant strains of pathogens, have accelerated the development of alternative approaches. Among these, biological control using harmless antagonistic microorganisms was proved to be a promising strategy for managing fruit diseases.
The aim of this research is to isolate and screen effective antagonists against blue mold and pear black spot rot, and identify them. The inhibition effects and their population dynamics in wounds were evaluated after the antagonists were screened. Main conclusions were drawn as follows.
1. There were 102 strains isolated from the wounds and the surface of fruits, leaves, and rhizosphere soil using the method of serial dilution. After primary and further screening in vitro, 10 strains exhibited inhibitory activity to Penicillium italicum and 9 strains exhibited inhibitory activity to Alternaria kikuchiana. Among screened strains, it was found that the isolates YF1, YF4 and BF2 were effective antagonists against P. italicum and A. kikuchiana on fruits through Fruits-inoculation test. The inhibition rate of isolates YF1, YF4 and BF2 got to 79.12%, 62.5% and 66.67% at 25℃.
2. Identification of the antagonists: According to the morphological, nutritional and physiological character, we found YF1 belonged to Candida sp., and YF4 belonged to Kloeckera apiculata. In order to determine the taxonomic placement, the identification of BF2 was performed with traditional bacterial systemic identification method. And then the molecular identification of BF2 was performed. Through the comparative 16S rDNA sequences analysis and phylogenetic analysis, we found BF2 belonged to Bacillus cereus.
3. The effects of antagonists increased with the increase of antagonists cell population when the antagonists at the concentrations of 106-108 CFU/mL. Fruits were treated with antagonists suspension at 1×108 CFU/mL at various time intervals (48 h, 24 h, 2 h, 0 h) before and after challenge-inoculated with pathogenic spore suspension at 104 spores/mL. The efficacy of antagonists against A. kikuchiana and P. italicum was maintained when it was applied simultaneously with or before the pathogens. However, the efficacy decreased when applied after the spores. The result of inoculation-time test showed that the efficacy of biocontrol increased significantly with the time antagonists applied before pathogen inoculated. In mixed-antagonists inoculation test, the blue mold infection rate was reduced to 12.50% when the two yeasts, YF1 and YF4, suspensions were applied simultaneously. This meant that the efficacy of the mixture excelled they applied individually and indicated significant discrepancy between them. The infection rates of black spot rot were 17.5%, 10.00% and 15.83% when isolates YF1, YF4 and BF2 were mixed with each other. It suggested that the efficacy of the two mixtures was better than that of YF1, YF4 and BF2. In the antangonists-enhancement test, the result showed that CaCl2 and low synthetic fungicide elevated biocontrol efficacy of antagonists. But when antagonists combined with SA, there is no different from antagonist used alone.
4. Rapid colonization of isolates YF1 and YF4 in wounds was observed whether pathogenic spores inoculated. The population of YF1 increased 123.26 folds after 48 h. When P. italicum spores inoculated simultaneously, the population increased 144.17 folds after 96 h. The population of YF4 increased almost 200 folds after 72 h. While it was 344.54 folds compared with the start point at 72 h when P. italicum spores co-inoculated, and it was 294.12 folds compared with the start point at 72 h when A. kikuchiana spores co-inoculated.
5. When the citrus fruits were soaked in the two yeast cell suspensions of 1×108 CFU/mL, the activities of phenlalanine ammonialyase (PAL) and peroxidase (POD) in fruits of citrus fruits raised remarkably, while polyphenol oxidase (PPO) reduced prominently and activities ofβ-1,3-glucanase and exochitiase almost did not change. But when P. italicum and the two yeasts added into the wounds of the fruit simultaneously, the activities ofβ-1, 3-glucanase and exochitiase remarkably raised by being induced. These indicate that both of the yeasts can induce diseases resistance in citrus.
引文
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