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

doi:10.16420/j.issn.0513-353x.2020-0476

摘要/Abstract

摘要：

从连作苹果园健康苹果树根际土壤中筛选出1株对苹果连作障碍镰孢属病原菌具有较强拮抗作用的细菌XC1,该菌对尖孢镰孢菌的抑菌率最高,达到79.83%。根据其形态、生理生化特征和基于16S rDNA、gyrA基因的系统发育分析,鉴定该菌为贝莱斯芽孢杆菌（Bacillus velezensis）。目前菌株XC1已保藏于中国微生物菌种保藏管理委员会普通微生物中心,保藏号为CGMCC No.20057。苹果连作土盆栽试验表明,XC1菌肥处理能显著促进平邑甜茶（Malus hupehensis Rehd.）幼苗的生长,其株高、地径、鲜质量和干质量分别比对照提高了54.8%、34.8%、121.1%和96.0%;显著提高连作土壤中的蔗糖酶、脲酶、过氧化氢酶和中性磷酸酶的活性,分别为对照的1.57倍、2.71倍、1.21倍和1.79倍;显著减少连作土壤中真菌的数量,提高细菌和放线菌的数量,且对尖孢镰孢菌（Fusarium oxysporum）有很好的抑制作用,尖孢镰孢菌的基因拷贝数比对照降低了76.4%。这说明贝莱斯芽孢杆菌抑制连作障碍病原菌的效果明显,可改善土壤环境,促进平邑甜茶幼苗的生长,能在一定程度上减轻苹果连作障碍。

关键词: 苹果, 连作障碍, 平邑甜茶, 贝莱斯芽孢杆菌, 镰孢菌

Abstract:

In this study,a strain XC1 was screened from the rhizosphere soil of healthy apple trees in the replanted apple orchard,which could antagonize the main pathogenic fungi（Fusarium solani,F. proliferatum,F. oxysporum,F. moniliforme）of apple replant disease. Among them,the inhibitory rate of F. oxysporumwas the highest,reaching 79.83%. Through morphological,physiological and biochemical characteristics,16S rDNA and gyrA sequence analysis,the bacterium was identified as Bacillus velezensis,which is a new biocontrol bacterium with development potential. At present,strain XC1 has been collected by the China General Microbiological Culture Collection Center,with the preservation number CGMCC No.20057. The pot experiment showed that compared with the control（replanted soil）,XC1 treatment could significantly promote the seedlings growth of Malus hupehensisRehd. and its plant height,ground diameter,fresh and dry mass were 54.8%,34.8%,121.1% and 96.0% higher than those of the control respectively;the activities of sucrase,urease,catalase and neutral phosphatase in the replanted soil treated with XC1 fertilizer were significantly increased,which were 1.57,2.71,1.21 and 1.79 times higher than those of the control,respectively. The number of fungi in the soil were significantly reduced;and the number of bacteria and actinomycetes was increased;and the gene copy number of F. oxysporum treated with XC1 fertilizer was 76.4% lower than that of the control. This indicated that B. velezensis has obvious bacteriostatic effect,which can improve the soil environment,promote the growth of M. hupehensisRehd. seedlings,and reduce the apple replant disease to a certain extent.

Key words: apple, replant disease, Malus hupehensis, Bacillus velezensis, Fusariumspp.

中图分类号:

S661.1

闫助冰, 王玫, 明常军, 姜远茂, 沈向, 陈学森, 尹承苗, 毛志泉. 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[J]. 园艺学报, 2021, 48(3): 409-420.

YAN Zhubing, WANG Mei, MING Changjun, JIANG Yuanmao, SHEN Xiang, CHEN Xuesen, YIN Chengmiao, MAO Zhiquan. Screening and Identification of Bacillus velezensis XC1 and Its Influence on Apple Replant Disease[J]. Acta Horticulturae Sinica, 2021, 48(3): 409-420.

图/表 9

参考文献 43

[1]	Alamer I S A, Tomah A A, Li B, Zhang J Z. 2020. Isolation,identification and characterization of rhizobacteria strains for biological control of bacterial wilt( Ralstonia solanacearum)of eggplant in China. Agriculture, 10 (2):1-16. doi: 10.3390/agriculture10010001 URL
[2]	Allison V J, Condron L M, Peltzer D A, Richardson S J, Turner B L. 2007. Changes in enzyme activities and soil microbial community composition along carbon and nutrient gradients at the Franz Josef chronosequence. New Zealand. Soil Biology and Biochemistry, 39 (7):1770-1781.
[3]	cai Chang-ping, Huang Jun, Zeng Yan, Bi Shi-yu, Huang Bin-bin, Guo Zhao-hui, Liu Qing-shu. 2018. Isolation and primary identification of an endophytic antagonistic bacteria from pepper. Hunan Agricultural Sciences, 394 (7):9-12. (in Chinese)
	蔡长平, 黄军, 曾艳, 毕世宇, 黄彬彬, 郭照辉, 刘清术. 2018. 一株辣椒内生拮抗细菌的筛选及初步鉴定. 湖南农业科学, 394 (7):9-12.
[4]	Cai Miao-ying, Dong Xiu-zhu. 2001. Identification of common bacterial system manual. Beijing: Science Press. (in Chinese)
	蔡妙英, 东秀珠. 2001. 常见细菌系统鉴定手册. 北京: 科学出版社.
[5]	Caputo F, Nicoletti F, Picione F D L, Manici L M. 2015. Rhizospheric changes of fungal and bacterial communities in relation to soil health of multi-generation apple orchards. Biological Control, 88 (9):8-17. doi: 10.1016/j.biocontrol.2015.04.019 URL
[6]	Chen Long, Wu Xing-li, Yan Xiao-gang, Wei Bing-dong, Zhang Fang-yu. 2020. The classification,secondary metabolites and application of Bacillus velezensis. Journal of Domestic Animal Ecology, 41 (1):1-8. (in Chinese)
	陈龙, 吴兴利, 闫晓刚, 魏炳栋, 张芳毓. 2020. 贝莱斯芽孢杆菌的分类、次级代谢产物及应用. 家畜生态学报, 41 (1):1-8.
[7]	Chen Xue-sen, Mao Zhi-quan, Shen Xiang, Jiang Yuan-mao, Wang Zhi-gang, Zhang Zong-ying, Wang Nan, Xu Hai-feng, Wang Yi-cheng. 2017. The breeding of new variety of easy colouring functional apple and non-bag cultivation technology. Chin Fruit,(1):1-5. (in Chinese)
	陈学森, 毛志泉, 沈向, 姜远茂, 王志刚, 张宗营, 王楠, 许海峰, 王意程. 2017. 易着色功能型苹果新品种培育及无袋化栽培技术. 中国果树,(1):1-5.
[8]	Du Shu-tao, Li Shu-na, Zhu Bao-cheng. 2010. Screening and identification of antagonistic strain DL- 59 of Bacillus velezensis against Alternaria brassicae and biocontrol efficiency. Journal of Hebei Agricultural University, 33 (6):51-56. (in Chinese)
	杜淑涛, 李术娜, 朱宝成. 2010. 白菜黑斑病拮抗细菌 Bacillus velezensis DL-59 的筛选鉴定及田间防效实验. 河北农业大学学报, 33 (6):51-56.
[9]	Guan Song-yin. 1986. Soil enzyme and its research methods. Beijing: Agriculture Press:274-340. (in Chinese)
	关松荫. 1986. 土壤酶及其研究法. 北京: 农业出版社:274-340.
[10]	He Bi-po, Zhao Hui, Liu Hong-yan, Ni Yun-xia, Wen Yi, Liu Xin-tao. 2019. Inhibitory effects,growth-promotion evaluation and identification of two biocontrol bacteria against fungal diseases of sesame. Journal of Henan Agricultural Sciences, 48 (11):92-98. (in Chinese)
	何碧珀, 赵辉, 刘红彦, 倪云霞, 文艺, 刘新涛. 2019. 2株芝麻真菌病害生防菌的抑菌特征、促生评价及鉴定. 河南农业科学, 48 (11):92-98.
[11]	Ju R, Zhao Y, Li J, Jiang H, Liu P, Yang T, Bao Z, Zhou B, Zhou X, Liu X. 2014. Identification and evaluation of a potential biocontrol agent, Bacillus subtilis,against Fusarium sp. in apple seedlings. Annals of Microbiology, 64 (6):377-383. doi: 10.1007/s13213-013-0672-3 URL
[12]	Kelderer M, Manici L M, Caputo F, Thalheimer M. 2012. Planting in the‘inter-row’to overcome replant disease in apple orchards:a study on the effectiveness of the practice based on microbial indicators. Plant and Soil, 357:381-393. doi: 10.1007/s11104-012-1172-0 URL
[13]	Li Jia-jia, Xiang Li, Pan Feng-bing, Chen Xue-sen, Shen Xiang, Yin Cheng-miao, Mao Zhi-quan. 2016. Effects of Malus hupehensis seedlings and Allium fistulosum mixed cropping on replanted soil environment. Acta Horticulturae Sinica, 43 (10):1853-1862. (in Chinese)
	李家家, 相立, 潘凤兵, 陈学森, 沈向, 尹承苗, 毛志泉. 2016. 平邑甜茶幼苗与葱混作对苹果连作土壤环境的影响. 园艺学报, 43 (10):1853-1862.
[14]	Li Xin-yue, Wang Chang-quan, Li Bing, He Jie, Li Yu-hao, Zhang Jing-sheng, Liang Jing-yue, Chen Lan, Yin Bin. 2019. Effect of controlled release blend bulk urea on soil microorganism and enzyme activity in rice and wheat rotation. Soil and Fertilizer Sciences in China, 279 (1):41-49. (in Chinese)
	李新悦, 王昌全, 李冰, 何杰, 李玉浩, 张敬昇, 梁靖越, 陈兰, 尹斌. 2019. 控释掺混氮肥对稻麦土壤微生物及酶活性的影响. 中国土壤与肥料, 279 (1):41-49.
[15]	Liu En-tai, Wang gong-shuai, Li yuan-yuan, Shen xiang, Chen xue-sen, Song fu-hai, Wu shu-jing, Chen qiang, Mao zhi-quan. 2014. Replanting affects the tree growth and fruit quality of Gala apple. J Integ Agri, 13 (8):1699-1706.
[16]	Liu Guo-qiang, Kong Ying-ying, Fan Ya-jing, Geng Ce, Peng Dong-hai, Sun Ming. 2017. Whole-genome sequencing of Bacillus velezensis LS69,a strain with a broad inhibitory spectrum against pathogenic bacteria. Journal of Biotechnology, 249:20-24. doi: 10.1016/j.jbiotec.2017.03.018 URL
[17]	Liu Li-ying, Xu Chao, Liu Ke-xin, Chi Xiao-li, Zhu Hao, Zhang Xiao, Jin Xiao, Liu Wei-wei, Sun Zhong-tao, Mao Zhi-quan. 2019. Effects of two endophytic antagonistic bacteria on growth of pot experiment Malus hupehensis seedlings and the soil enzymes activity under continuous cropping conditions. Acta Horticulturae Sinica, 46 (7):1238-1248. (in Chinese)
	刘丽英, 许超, 刘珂欣, 迟晓丽, 朱浩, 张潇, 金晓, 刘维维, 孙中涛, 毛志泉. 2019. 两株植物内生拮抗菌对连作土盆栽平邑甜茶幼苗生长及土壤酶活性的影响. 园艺学报, 46 (7):1238-1248.
[18]	Liu Zhi. 2013. The isolation and identification of fungi pathogen from apple replant disease in Bohai Bay region and the screening of the antagonistic Trichoderma strains[M. D. Dissertation]. Tai'an:Shandong Agricultural University. (in Chinese)
	刘志. 2013. 环渤海湾地区苹果再植病病原真菌分离鉴定及生防木霉菌株的筛选[硕士论文]. 泰安:山东农业大学.
[19]	Ma Hui, Yang Rui-xiu, Gai Xiao-tong, Liu Chang, Zhang Zhao-ran, Tang Shuang-shuang, Gao Zeng-gui. 2018. Alleviating of muskmelon soil of continuous cropping obstacles by different soil treatment. Agent Agrochemicals, 57 (10):764-767. (in Chinese)
	马慧, 杨瑞秀, 盖晓彤, 刘畅, 张照然, 唐爽爽, 高增贵. 2018. 不同土壤处理剂对甜瓜连作障碍的修复作用. 农药, 57 (10):764-767.
[20]	Ma Zhong-ming, Du Shao-ping, Wang Ping, Bao Xing-guo. 2012. Effects of long-term located fertilization on soil invertase activity in irrigated desert soils. Acta Agriculturae Boreali-Occidentalis Sinica, 21 (1):151-155. (in Chinese)
	马忠明, 杜少平, 王平, 包兴国. 2012. 长期定位施肥对灌漠土蔗糖酶活性的影响. 西北农业学报, 21 (1):151-155.
[21]	Masum M M I, Liu L, Yang M, Hossain M M, Siddiqa M M, Supty M E, Ogunyemi S O, Hossain A, An Q, Li B. 2018. Halotolerant bacteria belonging to operational group Bacillus amyloliquefaciens in biocontrol of the rice brown stripe pathogen Acidovorax oryzae. Journal of Applied Microbiology, 125:1852-1867. doi: 10.1111/jam.2018.125.issue-6 URL
[22]	Mazzola M, Manici L M. 2012. Apple replant disease:role of microbial ecology in cause and control. Annual Review of Phytopathology, 50:45-65. doi: 10.1146/annurev-phyto-081211-173005 pmid: 22559069
[23]	Mei Xin-lan, Zhao Qing-yun, Tan Shi-yong, Xu Yang-chun, Shen Biao, Shen Qi-rong. 2010. Screening,identification,and biocontrol effect of antagonistic bacteria against Phytophthora capsici. Chinese Journal of Applied Ecology, 21 (10):2652-2658. (in Chinese) pmid: 21328956
	梅新兰, 赵青云, 谭石勇, 徐阳春, 沈标, 沈其荣. 2010. 辣椒疫病拮抗菌株筛选、鉴定及其防效. 应用生态学报, 21 (10):2652-2658. pmid: 21328956
[24]	Schoor L V, Denman S, Cook N C. 2009. Characterisation of apple replant disease under South African conditions and potential biological management strategies. Scientia Horticulturae, 119 (2):153-162. doi: 10.1016/j.scienta.2008.07.032 URL
[25]	Shen Ping, Chen Xiang-dong. 2007. Experiment of Microbiology. Beijing: Higher Education Press. (in Chinese)
	沈萍, 陈向东. 2007. 微生物学实验. 北京教育出版社.
[26]	Shen Zong-zhuan, Sun Li, Wang Dong-sheng, Lü Na-na, Xue Chao, Li Rong, Shen Qi-rong. 2017. Effects of lime-ammonium bicarbonate fumigation and biofertilizer application on Fusarium wilt and biomass of continuous cropping cucumber and watermelon. Chinese Journal of Applied Ecology, 28 (10):3351-3359. (in Chinese)
	沈宗专, 孙莉, 王东升, 吕娜娜, 薛超, 李荣, 沈其荣. 2017. 石灰碳铵熏蒸与施用生物有机肥对连作黄瓜和西瓜枯萎病及生物量的影响. 应用生态学报, 28 (10):3351-3359.
[27]	Sheng Yue-fan, Wang Hai-yan, Wang Mei, Li Han-hao, Li Xiang, Pan Feng-bing, Chen Xue-sen, Shen Xiang, Yin Cheng-miao, Mao Zhi-quan. 2020. Effects of soil texture on the growth of young apple trees and soil microbial community structure under replanted conditions. Horticultural Plant Journal, 6 (3):123-131. doi: 10.1016/j.hpj.2020.04.003 URL
[28]	Song Yi-ling, Yu Jian, Chen Shi-geng, Xiao Cheng-ze, Li Yu-huan, Su Xiu-rong, Ding Fang-jun. 2018. Effects of reduced chemical fertilizer with application of bio-organic fertilizer on rape growth,microorganism and enzymes activities in soil. Journal of Soil and Water Conservation, 32 (1):352-360. (in Chinese)
	宋以玲, 于建, 陈士更, 肖承泽, 李玉环, 苏秀荣, 丁方军. 2018. 化肥减量配施生物有机肥对油菜生长及土壤微生物和酶活性影响. 水土保持学报, 32 (1):352-360.
[29]	Tao Yong-mei, Pan Hong-ji, Huang Jian, Xi Xin, Li Peng, Lu Zhi-jun. 2019. Research and application of a novel bio-control microbial factor Bacillus velezensis. China Plant Protection, 39 (9):26-33. (in Chinese)
	陶永梅, 潘洪吉, 黄健, 席昕, 李鹏, 卢志军. 2019. 新型生防微生物因子贝莱斯芽孢杆菌(Bacillus velezensis)的研究与应用. 中国植保导刊, 39 (9):26-33.
[30]	Tewoldemedhin Y T, Mazzola M, Labuschagne I, Mcleod A. 2011. A multi-phasic approach reveals that apple replant disease is caused by multiple biological agents,with some agents acting synergistically. Soil Biology and Biochemistry, 43 (9):1917-1927. doi: 10.1016/j.soilbio.2011.05.014 URL
[31]	Wang Gong-shuai. 2018. Studies on fungal community in replanted soil around Bohai Gulf and alleviation apple replanted disease by mixed cropping with Allium fistulosum L.[M. D. Dissertation]. Tai'an:Shandong Agricultural University. (in Chinese)
	王功帅. 2018. 环渤海连作土壤真菌群落结构分析及混作葱减轻苹果连作障碍的研究[硕士论文]. 泰安:山东农业大学.
[32]	Wang Xiao-fang, Xiang Kun, Wang Yan-fang, Li Qian-jin, Jiang Wei-tao, Sheng Yue-fan, Wang Hai-yan, Chen Xue-sen, Yin Cheng-miao, Mao Zhi-quan. 2019. Effects of Tagetes erecta biofumigation on apple replanted soil environment and physiological characteristics of Malus hupehensis seedlings. Acta Horticulturae Sinica, 46 (12):2383-2396. (in Chinese)
	王晓芳, 相昆, 王艳芳, 李前进, 姜伟涛, 盛月凡, 王海燕, 陈学森, 尹承苗, 毛志泉. 2019. 万寿菊植株粉末熏蒸对苹果连作土壤环境及平邑甜茶生理特性的影响. 园艺学报, 46 (12):2383-2396.
[33]	Wang Xiao-qi, Jiang Wei-tao, Yao Yuan-yuan, Yin Cheng-miao, Chen Xue-sen, Mao Zhi-quan. 2020. Research advance of apple replant disease based on soil microorganism. Acta Horticulturae Sinica, 47 (11):2223-2237. (in Chinese)
	王晓琪, 姜伟涛, 姚媛媛, 尹承苗, 陈学森, 毛志泉. 2020. 苹果连作障碍土壤微生物的研究进展. 园艺学报, 47 (11):2223-2237.
[34]	Wang Yan-fang, Xiang Li, Xu Shao-zhuo, Wang Sen, Wang Xiao-wei, Chen Xue-sen, Mao Zhi-quan, Zhang Min. 2017. Effects of biochar and chitin combined application on Malus hupehensis Rehd. seedlings and soil environment under replanting conditions. Scientia Agricultura Sinica, 50 (4):711-719. (In Chinese).
	王艳芳, 相立, 徐少卓, 王森, 王晓伟, 陈学森, 毛志泉, 张民. 2017. 生物炭与甲壳素配施对连作平邑甜茶幼苗及土壤环境的影响. 中国农业科学, 50 (4):711-719.
[35]	Wei Xiao-ya, Lin Yu, Chen Ting, Tao Zi-xi, Zhao Han-yu, Lin Sheng, Lin Wen-xiong. 2018. Effects of plant growth-promoting rhizobacteria on alleviating consecutive monoculture problem of Pseudostellaria heterophylla under field conditions. Chinese Journal of Ecology, 37 (2):399-408. (in Chinese)
	位小丫, 林煜, 陈婷, 陶子曦, 赵涵予, 林生, 林文雄. 2018. 田间条件下植物促生细菌缓解太子参连作障碍的效果评价. 生态学杂志, 37 (2):399-408.
[36]	Yang Sheng-qing, Zhang Fan, Ma Gui-long. 2017. Purification and inhibitive mechanism of antagonist substances from Bacillus velezensis S6 strain. Agrochemicals, 56 (9):645-648,660. (in Chinese)
	杨胜清, 张帆, 马贵龙. 2017. 贝莱斯芽孢杆菌S6拮抗物质分离纯化及抑菌机理. 农药, 56 (9):645-648,660.
[37]	Zhang Dan-yu, Qu Tian-tian, Liu Ying-ying, Guo Ying, Bu Ning. 2019. Antibacterial function and promoting effect of two plant growth promoting rhizobiain from Glycine soja. Soybean Science, 38 (4):563-569. (in Chinese)
	张丹雨, 曲甜甜, 刘莹莹, 郭英, 卜宁. 2019. 2株野生大豆根际促生菌抑菌促生作用研究. 大豆科学, 38 (4):563-569.
[38]	Zhang Hong, Lü Jia-long, Cao Ying-fei, Xu Wen-xin. 2014. Decomposition characteristics of different plant straws and soil microbial functional diversity. Acta Pedologica Sinica, 51 (4):743-752. (in Chinese)
	张红, 吕家珑, 曹莹菲, 徐温新. 2014. 不同植物秸秆腐解特性与土壤微生物功能多样性研究. 土壤学报, 51 (4):743-752.
[39]	Zhang Xian-fu, Xiang Li, Wang Yan-fang, Wang Gong-shuai, Liu Hui-xiang, Sun Geng-wu, Shen Xiang, Chen Xue-sen, Zhou Hui, Yin Cheng-miao, Mao Zhi-quan. 2016. Identification of Penicillium oxalicum A 1 strain and antagonistic effects on four species of Fusarium pathogen of apple. Acta Horticulturae Sinica, 43 (5):841-852. (in Chinese)
	张先富, 相立, 王艳芳, 王功帅, 刘会香, 孙庚午, 沈向, 陈学森, 周慧, 尹承苗, 毛志泉. 2016. 草酸青霉A1菌株的鉴定及对苹果4种镰孢病菌的拮抗作用. 园艺学报, 43 (5):841-852.
[40]	Zhang Zhi-xiao, Ze Sang-zi, Hu Lian-rong, Liu Ling, Ji Mei. 2018. Regulatory factors of soil urease activity and phylogenetic analysis of urease bacteria. Journal of West China Forestry Scienc, 47 (1):65-73. (in Chinese)
	张知晓, 泽桑梓, 户连荣, 刘凌, 季梅. 2018. 土壤脲酶活性调控因素和脲酶活性细菌系统发育研究. 西部林业科学, 47 (1):65-73. (in Chinese)
[41]	Zheng Xue-hao, Sun Li-na, Liu Ke-bin, Sun Jia-jun, Wang Xiao-xu, Zheng Jia-yu, Ma Xin-yu, Zhang Hong-ling, Wang Hui. 2018. Regulation of plant growth promoting rhizobacteria and lignin on remediation of POPs contaminated soil and their impacts on soil enzyme activity. Chinese Journal of Ecology, 37 (6):1807-1814. (in Chinese)
	郑学昊, 孙丽娜, 刘克斌, 孙家君, 王晓旭, 郑佳玉, 马欣雨, 张鸿龄, 王辉. 2018. 根际促生菌及木质素对持久性有机污染土壤修复的调控和酶活性的影响. 生态学杂志, 37 (6):1807-1814.
[42]	Zhou Wen-jie, Lü De-guo, Qin Si-jun, 2016. Research progress in interaction between plant and rhizosphere microorganism. Journal of Jilin Agricultural University, 38 (3):253-260. (in Chinese)
	周文杰, 吕德国, 秦嗣军. 2016. 植物与根际微生物相互作用关系研究进展. 吉林农业大学学报, 38 (3):253-260.
[43]	Zhu Zhan-ling, Bai Yan, Lü Ming-lu, Tian Ge, Zhang Xin, Li Li, Jiang Yuan-mao, Ge Shun-feng. 2020. Soil fertility,microbial biomass,and microbial functional diversity responses to four years fertilization in an apple orchard in north China. Horticultural Plant Journal, 6 (3):123-131. doi: 10.1016/j.hpj.2020.04.003 URL

特征 Characteristic	结果 Result	特征 Characteristic	结果 Result
V-P试验 V-P test	+	硝酸盐还原 Nitrate reduction	+
甲基红试验 Methyl red test	-	H₂S试验 H₂S test	-
淀粉水解试验 Starch hydrolysis test	+	动力学试验 Dynamic test	+
接触酶试验 Catalase test	+	明胶液化试验 Gelatin liquefaction test	+
吲哚测定 Indole test	-	D-葡萄糖 D-Glucose	+

特征 Characteristic	结果 Result	特征 Characteristic	结果 Result
V-P试验 V-P test	+	硝酸盐还原 Nitrate reduction	+
甲基红试验 Methyl red test	-	H₂S试验 H₂S test	-
淀粉水解试验 Starch hydrolysis test	+	动力学试验 Dynamic test	+
接触酶试验 Catalase test	+	明胶液化试验 Gelatin liquefaction test	+
吲哚测定 Indole test	-	D-葡萄糖 D-Glucose	+

处理 Treatment	株高/cm Plant height	地径/mm Ground diameter	鲜质量/g Fresh mass	干质量/g Dry mass
对照Control	49.56 ± 2.21 d	6.63 ± 0.09 c	25.13 ± 1.74 c	12.37 ± 0.26 c
溴甲烷Methyl bromide	87.77 ± 1.80 a	11.44 ± 0.61 a	98.07 ± 3.71 a	40.50 ± 1.71 a
菌肥载体Biofertilizer carrier	59.87 ± 3.12 c	6.84 ± 0.42 c	32.35 ± 3.69 c	15.38 ± 0.84 c
XC1菌肥XC1 biofertilize	76.70 ± 1.61 b	8.94 ± 0.33 b	55.56 ± 2.00 b	24.25 ± 1.03 b

处理 Treatment	株高/cm Plant height	地径/mm Ground diameter	鲜质量/g Fresh mass	干质量/g Dry mass
对照Control	49.56 ± 2.21 d	6.63 ± 0.09 c	25.13 ± 1.74 c	12.37 ± 0.26 c
溴甲烷Methyl bromide	87.77 ± 1.80 a	11.44 ± 0.61 a	98.07 ± 3.71 a	40.50 ± 1.71 a
菌肥载体Biofertilizer carrier	59.87 ± 3.12 c	6.84 ± 0.42 c	32.35 ± 3.69 c	15.38 ± 0.84 c
XC1菌肥XC1 biofertilize	76.70 ± 1.61 b	8.94 ± 0.33 b	55.56 ± 2.00 b	24.25 ± 1.03 b

处理 Treatment	细菌（× 10⁵ CFU·g^-1soil） Bacteria	真菌（×10³ CFU·g^-1soil） Fungi	放线菌（×10⁴ CFU·g^-1soil） Actinomyces
对照Control	20.33 ± 1.45 c	36.33 ± 3.48 a	42.33 ± 2.03 c
溴甲烷Methyl bromide	4.67 ± 0.33 d	5.00 ± 0.58 b	15.67 ± 1.76 d
菌肥载体Biofertilizer carrier	31.33 ± 2.03 b	31.33 ± 4.06 a	55.67 ± 0.67 b
XC1菌肥XC1 biofertilize	54.67 ± 2.33 a	7.67 ± 0.67 b	73.00 ± 1.53 a