马铃薯疮痂病生防菌1X1Y的鉴定及其生防促生特性研究

doi:10.16420/j.issn.0513-353x.2023-0647

摘要/Abstract

摘要：

从健康马铃薯根际土壤中分离获得1X1Y菌株，结合形态学特征、生理生化特性及多基因序列分析确定其为贝莱斯芽孢杆菌（Bacillus velezensis）。采用平板对峙培养法和抑菌圈法筛选拮抗菌株并验证其抑菌活性，其培养液均能显著抑制疮痂链霉菌Streptomyces acidiscabies和S. turgidiscabies的生长，抑菌圈直径为（34.92 ± 0.90）和（35.14 ± 1.56）mm，对疮痂病的相对防效为57.52%。该菌还可显著抑制马铃薯晚疫病菌、黑痣病菌、早疫病菌、炭疽病菌的生长，抑制率达56.44% ~ 71.47%。利用多对特异性引物检测菌株1X1Y抑菌活性物质合成关键基因，该菌含有srfAB、bmyB、fenD、bacA、ituC、bioA、beaS、dhbA、dfnA、mycB等10种抑菌活性物质合成关键基因。采用胞外酶和嗜铁素产生培养基、透明圈法、血平板、排油圈和紫外分光光度法对其胞外水解酶、抗性物质含量及防御酶活性等生防促生特性进行测定，该菌可分泌蛋白酶、β-1,3葡聚糖酶、纤维素酶、嗜铁素、脂肽类等抑菌相关物质，形成复杂生物膜结构，利于在根际定殖；同时可通过提高马铃薯可溶性蛋白含量及PAL、PPO防御酶活性，降低丙二醛含量来增强植株抗性。此外，1X1Y兼具固氮、ACC降解活性，可溶解磷酸钙、磷酸铁、磷酸铝、大豆卵磷脂、蛋黄卵磷脂、碳酸镁、氧化锌等多种难溶化合物，显著增加马铃薯株高、叶绿素相对含量、茎粗和单株产量。

关键词: 马铃薯, 疮痂病, 贝莱斯芽孢杆菌, 生物防治, 促生作用

Abstract:

The strain 1X1Y was isolated from the rhizosphere soil of healthy potato，and it was identified as Bacillus velezensis by morphological characteristics，physiological and biochemical characteristics and multi-gene sequence analysis. The antagonistic strain was screened and its antibacterial activity was verified by the plate confrontation culture method and the inhibition zone method. The culture solution could significantly inhibit the growth of Streptomyces acidiscabies and S. turgidiscabies，and the diameter of inhibition zone was（34.92 ± 0.90）and（35.14 ± 1.56）mm. The relative control effect on scab was 57.52%. It could also significantly inhibit the growth of Phytophthora infestans，Rhizoctonia solani，Alternaria alternata and Colletotrichum coccodes，with inhibition rate of 56.44%-71.47%. The key genes for the synthesis of antimicrobial substances in strain 1X1Y were detected by multiple pairs of specific primers. The strain 1X1Y contains 10 key genes for the synthesis of antimicrobial substances，such as srfAB，bmyB，fenD，bacA，ituC，bioA，beaS，dhbA，dfnA and mycB. The biocontrol and growth-promoting characteristics of extracellular hydrolase，resistance substance content and defense enzyme activity were determined by extracellular enzyme and siderophore production medium，transparent circle method，blood plate，oil discharge and ultraviolet spectrophotometry. The strain 1X1Y could secrete protease，β-1,3 glucanase，cellulase，siderophore，lipopeptide and other antibacterial related substances，and could form a complex biofilm structure，which was conducive to colonization in the rhizosphere. At the same time，the plant resistance was enhanced by increasing the soluble protein content of potato and the activities of PAL and PPO defense enzymes and reducing the content of malondialdehyde. In addition，strain 1X1Y has both nitrogen fixation and ACC degradation activity，which can dissolve calcium phosphate，iron phosphate，aluminium phosphate，soybean lecithin，egg yolk lecithin，magnesium carbonate，zinc oxide and other insoluble compounds，and significantly increase potato plant height，chlorophyll relative content，stem diameter and yield per plant.

Key words: potato, common scab, Bacillus velezensis, biological control, growth promoting effect

黄勋, 刘霞, 邓琳梅, 王兴国, 徐亚锦, 杨艳丽. 马铃薯疮痂病生防菌1X1Y的鉴定及其生防促生特性研究[J]. 园艺学报, 2025, 52(1): 229-246.

HUANG Xun, LIU Xia, DENG Linmei, WANG Xingguo, XU Yajin, YANG Yanli. Identification and Study on Its Biocontrol and Growth Promoting Properties of Bacillus velezensis 1X1Y Against Potato Common Scab[J]. Acta Horticulturae Sinica, 2025, 52(1): 229-246.

图/表 13

图1 菌株1X1Y在NA培养基上生长情况及菌落形态特征

Fig. 1 Growth and colony morphological characteristics of strain 1X1Y on NA medium

表1 拮抗菌株1X1Y的生理生化特征

Table1 Physiological and biochemical characteristics of antagonistic strain 1X1Y

项目名称 Object	试验现象 Experimental phenomenon	1X1Y	Bacillus velezensis
项目名称 Object	试验现象 Experimental phenomenon	1X1Y	R1B	XC1	QYQN-3
葡萄糖酵解 Glucose fermentation	培养基变为黄色，无气泡 The culture medium turns yellow and there are no bubbles	+	+	+	+
乳糖酵解 Lactose fermentation	培养基变为紫色，无气泡 The culture medium turns purple and there are no bubbles	-	/	/	+
淀粉水解 Starch hydrolysis	培养基颜色为紫色The culture medium turns purple	+	+	+	+
明胶液化 Gelatin liquefaction	培养基为液态 The culture medium is liquid	+	+	+	/
牛奶石蕊 Milk litmus	凝固胨化 Solidification and peptonization	+	/	/	+
脲酶产生 Urease	培养液颜色为红色 The color of the culture medium is red	+	/	/	/
H₂S产生 H₂S production	培养基中无黑色物质产生 No black substance was produced in the medium	-	-	-	/
甲基红试验 Methyl red test	培养液颜色为黄色 The color of the culture medium is yellow	-	-	-	-
柠檬酸盐利用 Citrate utilization	培养液颜色为蓝色 The color of the culture medium is blue	+	/	/	+
吲哚试验 Indole test	加入吲哚试剂后，培养液颜色为黄色 After adding indole reagent，the color of the culture medium is yellow	-	-	-	+
V-P试验 V-P test	培养液为红色 The culture medium is red	+	+	+	+

图2 菌株1X1Y基于16S rDNA、gyrA和gyrB序列构建的系统发育进化树

Fig. 2 Phylogenetic evolution tree of strain 1X1Y based on 16S rDNA，gyrA and gyrB sequences

图3 贝莱斯芽孢杆菌菌株1X1Y对马铃薯疮痂病原菌Streptomyces turgidiscabies和S. acidiscabies的平板对峙试验

Fig. 3 Plate face-off test of Bacillus velezensis strain 1X1Y against potato common scab pathogens Streptomyces turgidiscabies and S. acidiscabies

表2 菌株1X1Y培养液对马铃薯疮痂链霉菌的抑菌圈

Table 2 Inhibitory effect of strain 1X1Y on Streptomyces spp. mm

处理Treatment	S. turgidiscabies	S. acidiscabies
空白对照Control 1	0 b	0 b
纸片对照Control 2	0 b	0 b
1X1Y	35.14 ± 1.56 a	34.92 ± 0.90 a

表3 菌株1X1Y培养液对马铃薯疮痂病的防治效果

Table 3 1X1Y strain culture solution for the prevention and control of potato common scab disease

处理 Treatment	发病率/% Disease incidence	病情指数 Disease index	相对防效/% Relative control effect
对照 Control	100.00 ± 0.00 a	59.33 ± 5.84 a	—
JYC 314	88.33 ± 11.25 b	28.82 ± 7.47 b	51.42
1X1Y	77.42 ± 7.85 b	25.20 ± 1.77 b	57.52

图4 菌株1X1Y对4株马铃薯病原菌的抑制作用

Fig. 4 The inhibitory effect of strain 1X1Y on four potato pathogens

图5 菌株1X1Y抗菌物质合成相关基因PCR检测及溶血、排油试验结果

Fig. 5 Detection of genes related to synthesis of antimicrobial substances by PCR and results of hemolysis and oil discharge tests in strain 1X1Y

图6 菌株1X1Y胞外水解酶、嗜铁素产生及生物膜形成测定

Fig. 6 Determination of extracellular hydrolase，ferritin production and biofilm formation in strain 1X1Y

图7 菌株1X1Y对马铃薯植株可溶性蛋白和丙二醛含量及防御酶活性的影响图中同期小写字母表示经LSD法检验差异显著（P < 0.05）。下同

Fig. 7 Effect of 1X1Y on soluble protein，malondialdehyde content and defense enzyme activity of potato plants The lowercase letters of the same period in the figure indicate significant differences（P < 0.05）as tested by LSD method. The same below

图8 菌株1X1Y对难溶性无机磷酸盐和有机磷的溶解活性

Fig. 8 Solubility of strain 1X1Y to insoluble inorganic phosphate and organic phosphorus

图9 菌株1X1Y固氮、解钙、解镁、溶锌及产ACC脱氨酶活性检测

Fig. 9 Detection of the activities of strain 1X1Y for nitrogen fixation，calcium，magnesium，zinc dissolution and ACC deaminase production

图10 菌株1X1Y对马铃薯生长及产量的影响

Fig. 10 Effects of strain 1X1Y on growth and yield of potato

参考文献 59

[1]	Cao Weiping, Lu Qing, Lu Xiuyun, Zang Weiping, Song Jian. 2021. Identification of antagonistic bacterium Bacillus velezensis Hsg1949 and its biocontrol effect on peanut pod rot. Chinese Journal of Biological Control, 37 (4):761-770. (in Chinese) doi: 10.16409/j.cnki.2095-039x.2021.04.005
	曹伟平, 陆晴, 鹿秀云, 臧卫平, 宋健. 2021. 花生果腐病拮抗菌贝莱斯芽胞杆菌Hsg1949鉴定与防效. 中国生物防治学报, 37 (4):761-770. doi: 10.16409/j.cnki.2095-039x.2021.04.005
[2]	Chen Fangfang, Yu Xiaomei, Han Jingjing, Wu Chun, Pang Haiyue, Zhang Shudi, Chen Yupei. 2022. Antifungal activity and genomic analysis of Bacillus velezensis X49. Chinese Journal of Biotechnology, 38 (6):2292-2307. (in Chinese) doi: 10.13345/j.cjb.210804 pmid: 35786480
	陈芳芳, 余小妹, 韩晶晶, 吴春, 庞海月, 张书迪, 陈煜沛. 2022. 贝莱斯芽孢杆菌抗真菌能力及基因组分析. 生物工程学报, 38 (6):2292-2307. pmid: 35786480
[3]	Chen Shaoxian, Liu Yue, Yu Qian. 2022. Bacteriostatic effect of Bacillus velezensis zk 1 on rot-causing fungi of olecranon peach. China Food Additives, 33 (10):18-27. (in Chinese)
	陈少先, 刘悦, 余倩. 2022. 贝莱斯芽孢杆菌zk1对鹰嘴桃腐败真菌的抑菌作用. 中国食品添加剂, 33 (10):18-27.
[4]	Cheng Huanhuan, Yu Shui, Yao Weiwei, Li Zhong, Peng Lijuan, Ding Haixia. 2019. Screening of antagonistic Bacillus strains and control effect against pepper anthracnose in field trial. Journal of Henan Agricultural University, 53 (4):568-573. (in Chinese)
	程欢欢, 余水, 姚伟伟, 李忠, 彭丽娟, 丁海霞. 2019. 辣椒炭疽病生防芽孢杆菌的筛选及田间防效. 河南农业大学学报, 53 (4):568-573.
[5]	Deng Li, Sun Ran, Yao Jiangpang, Xie Yudan. 2021. Antibacterial mechanism of crude extract of Bacillus velezensis against Staphylococcus aureus. Food and Fermentation Sciences & Technology, 57 (3):1-9. (in Chinese)
	邓丽, 孙冉, 姚蒋庞, 谢毓丹. 2021 贝莱斯芽孢杆菌粗提物对金黄色葡萄球菌的抑菌机制. 食品与发酵科技, 57 (3):1-9.
[6]	Dong Xiuzhu, Cai Miaoying. 2001. Handbook of identification of common bacterial systems. Beijing: Science Press. (in Chinese)
	东秀珠, 蔡妙英. 2001. 常见细菌系统鉴定手册. 北京: 科学出版社.
[7]	Fang Yuan, Peng Yongzheng, Liao Changgui, Chen Lusheng, Zhou Qi, Huang Jian, Yan Yichao, Wang Muyuan, Zhang Yikun, Zou Lifang, Chen gongyou. 2022. Bacillus velezensis SF327,a potential biocontrol agent with the functions of preventing plant diseases and promoting plant growth. Acta Microbiologica Sinica, 62 (10):4071-4088. (in Chinese)
	方园, 彭勇政, 廖长贵, 陈路生, 周琦, 黄俭, 阎依超, 王慕媛, 张祎坤, 邹丽芳, 陈功友. 2022. 一株具有防病促生功能的贝莱斯芽孢杆菌SF327. 微生物学报, 62 (10):4071-4088.
[8]	Gao Zhenfeng, Zhao Jia. 2022. Effect of lipopeptide by Bacillus velezensis ZSY-1 on tomato postharvest softening and early blight. Acta Agriculturae Boreali-occidentalis Sinica, 31 (1):89-98. (in Chinese)
	振峰, 赵佳. 2022. 贝莱斯芽孢杆菌ZSY-1脂肽物质对番茄采后软化及早疫病发生的影响. 西北农业学报, 31 (1):89-98.
[9]	He Bipo, Hao Xuezheng, Liu Hongyan, Ni Yunxia, Liu Xintao, Zhao Xinbei, Zhao Hui. 2018. Analysis of growth-promotion,disease control effect and colonization capacity of Bacillus amyloliquefaciens B10-26 in sesame. Journal of Henan Agricultural Sciences, 47 (12):78-83. (in Chinese)
	何碧珀, 郝学政, 刘红彦, 倪云霞, 刘新涛, 赵新贝, 赵辉. 2018. 解淀粉芽孢杆菌B10-26对芝麻的促生防病效果及其定殖能力分析. 河南农业科学, 47 (12):78-83.
[10]	He Huyi, Tan Guanning, He Xinmin, Tang Zhouping, Li Lishu, Yang Xin. 2017. Identification of scab resistance in potato(Solanum tuberosum)Variety(line)resources. Journal of Plant Genetic Resources, 18 (4):786-793. (in Chinese) doi: 10.13430/j.cnki.jpgr.2017.04.023
	何虎翼, 谭冠宁, 何新民, 唐洲萍, 李丽淑, 杨鑫. 2017. 马铃薯品种(系)资源的疮痂病抗性鉴定. 植物遗传资源学报, 18 (4):786-793. doi: 10.13430/j.cnki.jpgr.2017.04.023
[11]	He Jianqing, Zhang Gejie, Zhao Weijin, Wang Xiaoxian. 2019. Screening and identification of superior associative nitrogen fixing bacteria in rhizosphere of Hordeum vulgare. Agricultural Research in the Arid Areas, 37 (5):182-186,192. (in Chinese)
	何建清, 张格杰, 赵伟进, 王孝先. 2019. 青稞根际优良联合固氮菌的筛选及鉴定. 干旱地区农业研究, 37 (5):182-186,192.
[12]	Huang Yonghui, Yang Mei, Quan Yongbing, Zhou Erxun. 2015. Induced resistance of benzothiadiazole to banana fusarium wilt. Journal of Huazhong Agricultural University, 34 (3):36-41. (in Chinese)
	黄永辉, 杨媚, 权永兵, 周而勋. 2015. 苯并噻二唑对香蕉枯萎病的诱导抗性. 华中农业大学学报, 34 (3):36-41.
[13]	Ji Wenxiu, Li Hulin, Leng Xue, Jin Zhengxun. 2019. Isolation and promoting growth properties of endophytic bacteria producing ACC deaminase of Panax ginseng. Journal of Jilin Agricultural University, 41 (2):168-174. (in Chinese)
	姬文秀, 李虎林, 冷雪, 金正勋. 2019. 产ACC脱氨酶人参内生细菌的分离和促生特性分析. 吉林农业大学学报, 41 (2):168-174.
[14]	Kim S Y, Lee S Y, Weon H Y, Sang M K, Song J. 2017. Complete genome sequence of Bacillus velezensis M75,a biocontrol agent against fungal plant pathogens,isolated from cotton waste. Journal of Biotechnology, 241:112-115. doi: S0168-1656(16)31623-6 pmid: 27899334
[15]	Li Dan, Li Rongmei, Qin Weiying, Zhou Yu, Shang Qiaoxia, Ren Zhengguang, Wei Yanmin, Zhao Xiaoyan. 2018. Control effects and related biocontrol factors detection of two Bacillus amyloliquefaciens strains on watermelon Fusarium Wilt. Chinese Journal of Biological Control, 34 (5):729-737. (in Chinese) doi: 10.16409/j.cnki.2095-039x.2018.05.012
	李丹, 李茸梅, 秦伟英, 周宇, 尚巧霞, 任争光, 魏艳敏, 赵晓燕. 2018. 两株解淀粉芽胞杆菌对西瓜枯萎病的防治作用及其相关生防因子检测. 中国生物防治学报, 34 (5):729-737. doi: 10.16409/j.cnki.2095-039x.2018.05.012
[16]	Li Miao, Li Yanming, Feng Ruizhang, Zhang Zixian, Zhou Wanhai, Yang Fuzhen, Wei Qin, Zuo Shan, Liu Yang. 2020. Identification and antagonistic activity of an endophytic strain Bacillus velezensis YZ 12 isolated from Cinnamomum longepaniculatum leaves. Biotic Resources, 42 (4):414-420. (in Chinese)
	李苗, 李彦铭, 冯瑞章, 张子娴, 周万海, 杨福珍, 魏琴, 左山, 刘洋. 2020. 特色香料作物油樟叶内生Bacillus velezensis YZ12的鉴定及拮抗病原菌活性. 生物资源, 42 (4):414-420.
[17]	Li Shengzhang, Chen Ying, Yang Ruihuan, Zhang Cuiping, Liu Zhao, Li Yilang, Chen Tao, Chen Gongyou, Zou Lifang. 2019. Isolation and identification of a Bacillus velezensis strain against plant pathogenic Xanthomonas spp. Acta Microbiologica Sinica, 59 (10):1969-1983. (in Chinese)
	李生樟, 陈颖, 杨瑞环, 张翠萍, 刘昭, 李逸朗, 陈涛, 陈功友, 邹丽芳. 2019. 一株拮抗黄单胞菌的贝莱斯芽孢杆菌的分离和鉴定. 微生物学报, 59 (10):1969-1983.
[18]	Li Yuyang, Xin Hanxiao, Fan Xueming, Liu Liying, Sun Zhongtao. 2017. Screening and identification of an antagonistic strain against Fusarium moniliforme in rice and its antifungal activity. Biotechnology Bulletin, 33 (5):190-196. (in Chinese) doi: 10.13560/j.cnki.biotech.bull.1985.2016-0826
	李玉洋, 辛寒晓, 范学明, 刘丽英, 孙中涛. 2017. 水稻恶苗病拮抗菌的筛选,鉴定及其抑菌活性. 生物技术通报, 33 (5):190-196. doi: 10.13560/j.cnki.biotech.bull.1985.2016-0826
[19]	Ling Juan, Dong Junde, Zhang Yanying, Yang Zhihao, Wang Youshao. 2010. Isolation and characterization of a N₂-fixing bacterium from the mangrove rhizosphere and study on its nitrogen-fixing ability. Journal of Tropical Oceanography, 29 (5):149-153. (in Chinese) doi: 10.11978/j.issn.1009-5470.2010.05.149
	凌娟, 董俊德, 张燕英, 杨志浩, 王友绍. 2010. 一株红树林根际固氮菌的分离,鉴定以及固氮活性测定. 热带海洋学报, 29 (5):149-153.
[20]	Luo Wenjian, Yang Fan, Shi Xuanjie, Zhao Xiuyun. 2018. Isolation and identification of antagonistic bacteria and biological control efficiency on cucumber Fusarium wilt. Journal of Huazhong Agricultural University, 37 (3):32-38. (in Chinese)
	罗文建, 杨凡, 史宣杰, 赵秀云. 2018. 黄瓜枯萎病拮抗菌的分离鉴定及其生物防效. 华中农业大学学报, 37 (3):32-38.
[21]	Meng Lulu, Yu Chunlei, Xue Desheng, Li Baohua, Liang Wenxing, Wang Caixia. 2019. Control efficiency of BHT against postharvest apple grey mold and its influence on the activity of defensive enzymes and content of malondialdehyde. Journal of Plant Protection, 46 (3):686-692. (in Chinese)
	孟璐璐, 于春蕾, 薛德胜, 李保华, 梁文星, 王彩霞. 2019. BHT对苹果采后灰霉病的防效及防御酶活性和丙二醛含量的影响. 植物保护学报, 46 (3):686-692.
[22]	Ou Ting, Jin Bikun, Gao Haiying, Wang Ruolin, Zhang Yuyang, Zuo Weidong, Xie Jie. 2022. Purification and research of inhibitory mechanism of antagonist substances from Bacillus velezensis SWUJ 1 strain. Journal of Southwest University(Natural Science Edition), 44 (1):75-87. (in Chinese)
	欧婷, 金必堃, 高海英, 王若琳, 张雨阳, 左伟东, 谢洁. 2022. Bacillus velezensis SWUJ1拮抗物质分离纯化及抑菌机理研究. 西南大学学报(自然科学版), 44 (1):75-87.
[23]	Ren Qingxin, Wang Jihua, Zeng Qi, Zhang Bixian, Liu Xiulin. 2022. Study on phosphorus solubilization characteristics of Talaromyces sp. WR1-4 in soybean(Glycine max)rhizosphere. Journal of Agricultural Biotechnology, 30 (1):114-124. (in Chinese)
	任氢欣, 王继华, 曾齐, 张必弦, 刘秀林. 2022. 大豆根际踝节真菌WR1-4的溶磷特性研究. 农业生物技术学报, 30 (1):114-124.
[24]	Sang Jianwei, Yang Yang, Chen Yipeng, Cai Jimiao, Lu Cuimei, Huang Guixiu. 2018. Antibacterial activity analysis of lipopeptide and polyketide compounds produced by endophytic bacteria Bacillus amyloliquefaciens BEB17. Acta Phytopathologica Sinica, 48 (3):402-412. (in Chinese)
	桑建伟, 杨扬, 陈奕鹏, 蔡吉苗, 陆翠梅, 黄贵修. 2018. 内生解淀粉芽孢杆菌BEB17脂肽类和聚酮类化合物的抑菌活性分析. 植物病理学报, 48 (3):402-412.
[25]	Shan Wenrong, Li Junxia, Liu Huafen. 2010. Study on the inhibitory effects of different active materials screened by the filter paper on cotton verticillium wilt. Chinese Agricultural Science Bulletin, 26 (19):285-289. (in Chinese) doi: 10.11924/j.issn.1000-6850.2010-1478
	单文荣, 李俊霞, 刘花粉. 2010. 滤纸片法筛选不同活性物对棉花黄萎病菌抑制效果研究. 中国农学通报, 26 (19):285-289.
[26]	Shi Yingying, Zhao Pan, Song Shuangwei, Xiong Minzi, Mo Chengbao, Zhong Naiqin. 2020. Isolation and characterization of the antagonistic bacterium YN-2- 2 against potato common scab. Microbiology China, 47 (8):2425-2435. (in Chinese)
	石莹莹, 赵盼, 宋双伟, 熊悯梓, 莫乘宝, 仲乃琴. 2020. 马铃薯疮痂病拮抗菌YN-2-2的分离与鉴定. 微生物学通报, 47 (8):2425-2435.
[27]	Sichuan Xinlanyue Biotechnology Co.,Ltd., Sichuan Lanyue Technology Co.,Ltd. 2021. A strain of Bacillus velezensis and its cultivation method and application:China,202011401120.4. 2021-04-30. (in Chinese)
	四川省新兰月生物科技有限公司, 四川省兰月科技有限公司. 2021. 一株贝莱斯芽孢杆菌及其培养方法和应用:中国:202011401120.4.2021-04-30.
[28]	Sun Yanchao, Yang Taixin, Yuan Mengjia. 2020. Study on the correlation between cold resistance and its correlation with physiological index of different Bupleurum chinense germplasms. Journal of Chinese Medicinal Materials, 43 (1):5-8. (in Chinese)
	孙延超, 杨太新, 袁梦佳. 2020. 不同柴胡种质抗寒性及其与生理指标的相关性研究. 中药材, 43 (1):5-8.
[29]	Tai Lianmei, Chen Zhiyao, Wang Peng, Wang Yuqi, Jin Guanghui, Jin Xuehui. 2021. Screening and identification of antagonistic bacteria of Streptomyces scabies. Journal of Henan Agricultural Sciences, 50 (11):108-116. (in Chinese)
	台莲梅, 陈志垚, 王鹏, 王钰琪, 金光辉, 靳学慧. 2021. 马铃薯疮痂病菌Streptomyces scabies拮抗细菌筛选及鉴定. 河南农业科学, 50 (11):108-116.
[30]	Tu Zhihua, Zhou Lingfeng, Huang Yanping, Chen Suyi, Chen Jinhui, Li Shengnan. 2021. Rhizosphere effects of soil enzyme activities in tropical cloud forests in Limushan national nature reserve on Hainan island. Bulletin of Soil and Water Conservation, 41 (3):1-7. (in Chinese)
	涂志华, 周凌峰, 黄艳萍, 陈夙怡, 陈金辉, 李胜男. 2021. 海南岛黎母山国家自然保护区热带云雾林土壤酶活性的根际效应. 水土保持通报, 41 (3):1-7.
[31]	Wang Di, Gao Yan, Wu Xiaoshuang, Zhang Wei, Shu Jinping, Zhang Yabo, Zhai Fengyan. 2022. Identification of two strains of Bacillus velezensis isloated from Carya illinoensis leaf and their antagonistic effects on pecan black spot pathogen. Chinese Journal of Biological Control, 38 (6):1572-1581. (in Chinese)
	王迪, 高岩, 吴小双, 张威, 舒金平, 张亚波, 翟凤艳. 2022. 两株贝莱斯芽胞杆菌鉴定及其对薄壳山核桃黑斑病病原菌的拮抗效果. 中国生物防治学报, 38 (6):1572-1581. doi: 10.16409/j.cnki.2095-039x.2022.08.003
[32]	Wang Mingyue, Liu Shaoxiong, Xiong Zhi, Ding Yadi, Wang Yanling, Li Keyan, Yang Tingdan, Tao Xi, Wang Jinhua. 2014. Study on leguminous plants rhizobia degradation of calcium carbonate and magnesium carbonate. Ecology and Environmental Sciences, 23 (10):1581-1585. (in Chinese)
	王明月, 刘绍雄, 熊智, 丁雅迪, 王艳玲, 李克艳, 阳廷丹, 陶茜, 王金华. 2014. 石漠化地区豆科植物根瘤菌降解碳酸钙、镁能力研究. 生态环境学报, 23 (10):1581-1585.
[33]	Wang Nan, Liao Yongqin, Shi Zhufeng, Shen Yunxin, Yang Tongyu, Feng Luyao, Yi Xiaopeng, Tang Jiacai, Chen Qibin, Yang Peiwen. 2024. Identification of three strains of Bacillus from the forest soil of Wuliang mountain and mining of their bioactivities. Biotechnology Bulletin, 40 (2):277-288. (in Chinese)
	王楠, 廖永琴, 施竹凤, 申云鑫, 杨童雨, 冯路遥, 矣小鹏, 唐加菜, 陈齐斌, 杨佩文. 2024. 三株无量山森林土壤芽孢杆菌鉴定及其生物活性挖掘. 生物技术通报, 40 (2):277-288. doi: 10.13560/j.cnki.biotech.bull.1985.2023-0918
[34]	Wang Ruolin, Xu Weifang, Wang Fei, Zhou Xiaolei, Zheng Yue, Jiang Hongsen, Xie Jie. 2019. Isolation and identification of an antagonistic endophytic bacterium from mulberry for biocontrol against Boeremia exigua. Acta Microbiologica Sinica, 59 (11):2130-2143. (in Chinese)
	王若琳, 徐伟芳, 王飞, 周小磊, 郑月, 江鸿森, 谢洁. 2019. 桑树内生拮抗菌的分离鉴定及其对桑断枝烂叶病的生防初探. 微生物学报, 59 (11):2130-2143.
[35]	Wang Weinan, Lan Zhiyong, Yu Wenli, Sun Qingpei, Li Peiqi, Fan Yonghong. 2021. Screening and identification of salt-tolerant and ACC deaminase producing strains in Halostachys caspica rhizosphere. Soil and Fertilizer Sciences in China,(2):270-275. (in Chinese)
	王伟楠, 兰智勇, 喻文丽, 孙庆培, 李佩琪, 樊永红. 2021. 盐穗木根际产ACC脱氨酶耐盐菌株的筛选及鉴定. 中国土壤与肥料,(2):270-275.
[36]	Wang Yuxia, Jiang Wei, Liu Yushuai, Meng Liqiang, Li Jing, Cao Xu, Hu Jihua, Chen Jingyu, Zhang Shumei. 2016. Study on isolation and characterization of a psychrotrophic antagonistic bacterium from cold area. Journal of Northeast Agricultural University, 47 (8):31-38. (in Chinese)
	王玉霞, 姜威, 刘宇帅, 孟利强, 李晶, 曹旭, 胡基华, 陈静宇, 张淑梅. 2016. 寒地耐冷生防菌株筛选鉴定及特性研究. 东北农业大学学报, 47 (8):31-38.
[37]	Wei Yanxia, Li Zhuoran, Zhang Bin, Yuan Yujin, Yu Weiwei, Chang Ruokui, Wang Yuanhong. 2021. Screening and function of plant immune proteins from Bacillus velezensis LJ02. Scientia Agricultura Sinica, 54 (16):3451-3460. (in Chinese)
	魏艳侠, 李卓然, 张斌, 苑瑜瑾, 于玮玮, 常若葵, 王远宏. 2021. 贝莱斯芽孢杆菌LJ02中植物免疫蛋白的筛选及其功能. 中国农业科学, 54 (16):3451-3460. doi: 10.3864/j.issn.0578-1752.2021.16.008
[38]	Wu Liqin, Shang Hongzhong, Gu Haike. 2019. Screening,identification of biocontrol bacterium R1B against Rhizopus stolonifer and analysis of its antagonistic characteristics. Biotechnology Bulletin, 35 (4):29-35. (in Chinese) doi: 10.13560/j.cnki.biotech.bull.1985.2018-0870
	武利勤, 尚宏忠, 顾海科. 2019. 拮抗匍枝根霉的生防菌R1B的筛选鉴定和抑菌活性分析. 生物技术通报, 35 (4):29-35. doi: 10.13560/j.cnki.biotech.bull.1985.2018-0870
[39]	Xu Xinyun, Wang Ning, Ding Jia, Chen Yan, Tian Guangming. 2021. Isolation and identification of antagonistic bacteria against tomato bacterial wilt and the mechanisms in disease prevention and plant growth promotion. Acta Microbiologica Sinica, 61 (10):3276-3290. (in Chinese)
	徐欣韵, 王宁, 丁佳, 陈研, 田光明. 2021 番茄青枯病拮抗菌的定向筛选及其抗病促生机制研究. 微生物学报, 61 (10):3276-3290.
[40]	Xu Yuanyuan, Hu Dangzhen, Nan jing, Zhan Ping, Jiang Ling. 2022. The effect of co treatment with two types of Bacillus on citrus seedlings infected with Huang long disease. South China Fruits, 51 (1):9-18. (in Chinese)
	徐媛媛, 胡党振, 南京, 詹平, 姜玲. 2022. 两种芽孢杆菌共处理对感染黄龙病的柑桔苗的作用. 中国南方果树, 51 (1):9-18.
[41]	Yan Zhubing, Wang Mei, Ming Changjun, Jiang Yuanmao, Shen Xiang, Chen Xuesen, Yin Chengmiao, Mao Zhiquan. 2021. Screening and identification of Bacillus velezensis XC1 and its influence on apple replant disease. Acta Horticulturae Sinica, 48 (3):409-420. (in Chinese) doi: 10.16420/j.issn.0513-353x.2020-0476 URL
	闫助冰, 王玫, 明常军, 姜远茂, 沈向, 陈学森, 尹承苗, 毛志泉. 2021. 贝莱斯芽孢杆菌XC1的筛选、鉴定及其对苹果连作障碍的影响. 园艺学报, 48 (3):409-420. doi: 10.16420/j.issn.0513-353x.2020-0476 URL
[42]	Yang Di, Du Chanjuan, Zhang Jin, Pan Lianfu, Ye Yunfeng, Huang Siliang, Fu Gang. 2021. Screening,identification and biological characteristics of Bacillus velezensis with antagonst activity against banana Fusarium wilt. Chinese Journal of Biological Control, 37 (1):165-171. (in Chinese)
	杨迪, 杜婵娟, 张晋, 潘连富, 叶云峰, 黄思良, 付岗. 2021. 香蕉枯萎病拮抗菌贝莱斯芽胞杆菌的筛选鉴定及其生物学特性. 中国生物防治学报, 37 (1):165-171. doi: 10.16409/j.cnki.2095-039x.2021.01.031
[43]	Yang Xiaolei, Qian Guoliang, Fan Jiaqin, Hu Baishi, Liu Fengquan. 2014. Selection and identification of antagonist Alternaria alternate and initial analysis of its antipathogenic activity. Journal of Nanjing Agricultural University, 37 (1):68-74. (in Chinese)
	杨晓蕾, 钱国良, 范加勤, 胡白石, 刘凤权. 2014. 梨黑斑病菌拮抗细菌的筛选鉴定及其拮抗活性的研究. 南京农业大学学报, 37 (1):68-74.
[44]	Yang Yanli, Liu Xia. 2019. Potato diseases. Beijing: Science Press:54-59. (in Chinese)
	杨艳丽, 刘霞. 2019. 马铃薯病害. 北京: 科学出版社:54-59.
[45]	Yunnan Agricultural University. 2022. Antagonistic strain JYC 314 and its application:China,202210041956.0. 2022-08-23. (in Chinese)
	云南农业大学. 2022. 拮抗菌株JYC 314及其应用:中国,202210041956.0. 2022-08-23.
[46]	Zang Chaoqun, Lin Ying, Xie Jinhui, Liu Xiaozhou, Pei Xue, Liang Chunhao. 2021. Preliminary study on antimicrobial mechanism of Bacillus velezensis against Peyronellaea arachidicola. Journal of Yangzhou University:Agricultural and Life Science Edition, 42 (5):130-134. (in Chinese)
	臧超群, 林英, 谢瑾卉, 刘晓舟, 裴雪, 梁春浩. 2021. 贝莱斯芽孢杆菌BP-1对花生网斑病菌的抑菌机制初探. 扬州大学学报:农业与生命科学版, 42 (5):130-134.
[47]	Zeng Xin, Zhang Yahui, Chi Huirong, Sun Zhongyu, Chen Weiliang, Mao Bizeng. 2019. Antimicrobial activity of endophytic bacterium strain B-11 isolated from Curcuma wenyujin. Microbiology China, 46 (5):1018-1029. (in Chinese)
	曾欣, 张亚惠, 迟惠荣, 孙钟毓, 陈卫良, 毛碧增. 2019. 温郁金内生拮抗细菌B-11的分离及其抑菌活性. 微生物学通报, 46 (5):1018-1029.
[48]	Zhang Caiwen, Cheng Kun, Zhang Xin, Liu Bo, Du Haibo, Yao Su. 2019. Progress in taxonomy and functional research of Bacillus velezensis. Food and Fermentation Industries, 45 (17):258-265. (in Chinese) doi: 10.13995/j.cnki.11-1802/ts.020737
	张彩文, 程坤, 张欣, 刘波, 都海渤, 姚粟. 2019. 贝莱斯芽胞杆菌(Bacillus velezensis)分类学及功能研究进展. 食品与发酵工业, 45 (17):258-265. doi: 10.13995/j.cnki.11-1802/ts.020737
[49]	Zhang Defeng, Gao Yanxia, Wang Yajun, Liu Chun, Shi Cunbin. 2020. Advances in taxonomy,antagonistic function and application of Bacillus velezensis. Microbiology China, 47 (11):3634-3649. (in Chinese)
	张德锋, 高艳侠, 王亚军, 刘春, 石存斌. 2020. 贝莱斯芽孢杆菌的分类,拮抗功能及其应用研究进展. 微生物学通报, 47 (11):3634-3649.
[50]	Zhang Dezhen, Chi Wenjuan, Li Tingting, Dai Huijie, Qiao Ning, Wang Cuicui. 2021. Effects of biocontrol bacteria against pepper phytophthora blight on disease control and growth promotion and its identification. Shandong Agricultural Sciences, 53 (10):91-96. (in Chinese)
	张德珍, 迟文娟, 李婷婷, 代惠洁, 乔宁, 王翠翠. 2021. 辣椒疫病生防菌的防病促生作用及其鉴定. 山东农业科学, 53 (10):91-96.
[51]	Zhang Lijuan, Huang Wei, Wang Ning, Song Bo, Zhu Jing, Abdukeyim Rayhangul, Abbas Bujannat, Wang Wei. 2021. Identification of radiation-resistant Bacillus velezensis JK23 and its biocontrol Activity. Chinese Journal of Biological Control, 37 (5):997-1006. (in Chinese) doi: 10.16409/j.cnki.2095-039x.2021.06.011
	张丽娟, 黄伟, 王宁, 宋博, 朱静, 热依罕姑丽 · 阿卜杜克依穆, 布坚乃提 · 阿巴斯, 王玮. 2021. 耐辐射芽胞杆菌JK23的鉴定及其生防活性研究. 中国生物防治学报, 37 (5):997-1006. doi: 10.16409/j.cnki.2095-039x.2021.06.011
[52]	Zhang Meng, Liu Bo, Yu Xiumei, Zhao Weiquan, Liu Daqun. 2010. Biological characteristics analysis of the pathogens caused potato common scab in China. Scientia Agricultura Sinica, 43 (12):2603-2610. (in Chinese)
	张萌, 刘伯, 于秀梅, 赵伟全, 刘大群. 2010. 中国马铃薯疮痂病菌生物学特性分析. 中国农业科学, 43 (12):2603-2610. doi: 10.3864/j.issn.0578-1752.2010.12.028
[53]	Zhang Qian, Chen Yushi, Xu Chunyan, Zou Man, Bao Wenting, Xi Liangqing, Xin Li, Chen Yilun. 2023. Efﬁcacy and action mechanism of Bacillus velezensis on controlling postharvest soft rot of sweet cherry fruits. Food Science, 44 (7):229-239. (in Chinese)
	张倩, 陈雨诗, 许春艳, 邹曼, 包汶婷, 郗良卿, 辛力, 陈义伦. 2023. 贝莱斯芽孢杆菌防治甜樱桃采后软腐病的效果和机理. 食品科学, 44 (7):229-239.
[54]	Zhang Qiong, Zabihullah Sherzad, Tang Canming. 2020. Biocontrol effect of Bacillus velezensis strain SZAD 1 on Verticillium dahliae. Cotton Science, 32 (4):329-338. (in Chinese)
	张琼, Zabihullah Sherzad, 唐灿明. 2020. 贝莱斯芽孢杆菌SZAD1对大丽轮枝菌的生物防治效果. 棉花学报, 32 (4):329-338. doi: 10.11963/1002-7807.zqtcm.20200609
[55]	Zhang Zhe, Liu Xia, Wang Xingguo, Yang Li, Chen Li, Huang Xun, Zhang Yamin, Yang Yanli. 2023. A preliminary study on the effects of potassium dihydrogen phosphate on the resistance of potato to late blight. Plant Protection, 49 (3):85-91,140. (in Chinese)
	张哲, 刘霞, 王兴国, 杨俐, 陈丽, 黄勋, 张雅敏, 杨艳丽. 2023. 磷酸二氢钾影响马铃薯对晚疫病抗性的初步研究. 植物保护, 49 (3):85-91,140.
[56]	Zhao Wenjuan, Fu Bo, Xu Shengyun, Ren Ping. 2017. Antagonism of biocontrol bacteria SF1103 and SF 1104 on wilt pathogen in cucumber. Acta Agriculturae Boreali-occidentalis Sinica, 26 (10):1537-1543. (in Chinese)
	赵文娟, 付博, 徐升运, 任平. 2017. 生防菌SF1103、SF1104对黄瓜枯萎病菌的拮抗作用. 西北农业学报, 26 (10):1537-1543.
[57]	Zhou Hongzi, Zhou Fangyuan, Duan Chengding, Wu Xiaoqing, Zhao Xiaoyan, Xie Xueying, Zhang Guangzhi, Fan Susu, Zhang Xinjian. 2021. Screening and identification of antagonistic bacteria of maize ear rot and its degradation effect on DON. Chinese Journal of Biological Control, 37 (5):1016-1023. (in Chinese) doi: 10.16409/j.cnki.2095-039x.2021.06.005
	周红姿, 周方园, 段成鼎, 吴晓青, 赵晓燕, 谢雪迎, 张广志, 范素素, 张新建. 2021. 玉米穗腐病菌的拮抗菌筛选,鉴定及降解DON毒素能力测定. 中国生物防治学报, 37 (5):1016-1023. doi: 10.16409/j.cnki.2095-039x.2021.06.005
[58]	Zhou Xiangping, Teng Kai, Xiao Qiming, Chen Wu, Wang Kaige, Zhang Hongbo, Liu Feng, Liu Tianbo. 2022. Tobacco growth promoting effect of Bacillus velezensis F10 and its control effect against tobacco bacterial wilt. Tobacco Science & Technology, 55 (7):9-16. (in Chinese)
	周向平, 滕凯, 肖启明, 陈武, 王凯歌, 张洪波, 刘峰, 刘天波. 2022. 贝莱斯芽胞杆菌F10促生作用及对烟草青枯病的防治效果. 烟草科技, 55 (7):9-16.
[59]	Zhou Y J, Li Q, Peng Z, Zhang J, Li J H. 2022. Biocontrol effect of Bacillus subtilis YPS-32 on potato common scab and its complete genome sequence analysis. Journal of Agricultural and Food Chemistry, 70 (17):5339-5348.