枯草芽胞杆菌JY-7-2L对乌头根腐病的生防能力及其潜在作用机制解析

doi:10.13560/j.cnki.biotech.bull.1985.2025-1200

摘要/Abstract

摘要：

目的探究枯草芽胞杆菌Bacillus subtilis JY-7-2L对乌头根腐病原菌尖孢镰刀菌（Fusarium oxysporum）的拮抗作用，解析其对大田根腐病的生防潜力，并从基因组层面剖析其抗菌机制。方法采用平板对峙试验检测JY-7-2L拮抗病原菌能力，大田试验检测菌株对根腐病的生防潜力和对乌头生长的影响，用Illumina NovaSeq和Oxford Nanopore ONT平台进行全基因组测序，通过COG、CAZy、GO和KEGG数据库对JY-7-2L基因组信息进行基因功能注释，并通过antiSMASH工具预测其次级代谢产物合成基因。结果 JY-7-2L显著抑制F. oxysporum的生长，并显著降低大田根腐病发病率，生防效率为89.26%-93.77%，同时增加了植株茎、主根和子根的干重。该菌的无细胞发酵滤液和挥发性物质均能显著抑制尖孢镰刀菌的生长。JY-7-2L基因组总长度为4.19 Mb，GC含量43.57%，共编码4 234个基因。JY-7-2L基因组中含有抗生素合成、生物膜形成、抑制病原菌、促进植物生长和诱导植物系统抗性相关基因。AntiSMASH分析表明，该菌株含有11个次级代谢产物合成基因簇，其中6个基因簇能够产生拮抗真菌或细菌的活性物质，3个基因簇具有合成新型活性物质的潜力。结论 JY-7-2L兼具生防和促生能力，具有开发作为生物肥料和生防材料的潜力。

关键词: 枯草芽胞杆菌, 生物防治, 乌头, 根腐病, 全基因组, 抑菌机制

Abstract:

Objective This study aimed to evaluate the antagonistic activity of Bacillus subtilis JY-7-2 against the Aconitum carmichaelii Debeaux root rot pathogen Fusarium oxysporum and its biocontrol efficacy against the root rot in the field, with genomic analysis to elucidate the underlying antimicrobial mechanisms. Method We evaluated the antagonistic activity of strain JY-7-2L with a plate confrontation assay. In field trials, we assessed its biocontrol efficacy against root rot and its effects on the growth of A. carmichaelii Debeaux. We performed whole-genome sequencing using the Illumina NovaSeq and Oxford Nanopore platforms, conducted functional annotation of the genome with the COG, CAZy, GO, and KEGG databases, and predicted secondary metabolite biosynthetic gene clusters with antiSMASH. Result Strain JY-7-2L significantly inhibited the growth of F. oxysporum and reduced the incidence of root rot in the field, achieving a biocontrol efficacy of 89.26%-93.77%. It also increased the dry weights of plant stems, main roots, and lateral roots. Both the cell-free fermentation filtrate and volatile organic compounds of JY-7-2L markedly suppressed the growth of F. oxysporum. The genome of JY-7-2L was 4.19 Mb in length with a GC content of 43.57%, encoding 4 234 genes. Genomic analysis revealed the presence of genes associated with antibiotic synthesis, biofilm formation, pathogen inhibition, plant growth promotion, and induction of plant systemic resistance. AntiSMASH analysis identified 11 secondary metabolite biosynthesis gene clusters, including 6 involved in the synthesis of antimicrobial compounds, and 3 potentially contribute to novel bioactive metabolites. Conclusion With its dual biocontrol and plant growth-promoting activities, Bacillus subtilis JY-7-2L shows strong potential for development as a biofertilizer and biocontrol agent.

Key words: Bacillus subtilis, biocontrol, Aconitum carmichaelii, root rot, whole genome, antifungal mechanism

邱宇洁, 王茜, 曾琳, 侯虹吉, 苟玉梅, 邹兰, 黄晶. 枯草芽胞杆菌JY-7-2L对乌头根腐病的生防能力及其潜在作用机制解析[J]. 生物技术通报, 2026, 42(8): 298-311.

QIU Yu-jie, WANG Qian, ZENG Lin, HOU Hong-ji, GOU Yu-mei, ZOU Lan, HUANG Jing. Biocontrol Efficacy of Bacillus subtilis JY-7-2L against Aconitum carmichaelii Debeaux Root Rot and Its Potential Mechanisms[J]. Biotechnology Bulletin, 2026, 42(8): 298-311.

图/表 12

参考文献 60

[1]	邹兰, 王茜, 李慕仪, 等. 乌头内生细菌JY-3-1R的鉴定及其生防和促生能力研究 [J]. 生物技术通报, 2023, 39(10): 246-255.
	Zou L, Wang Q, Li MY, et al. Identification, biocontrol and plant growth-promoting potential of endophytic bacterial strain JY-3-1R from Aconitum carmichaelii debx [J]. Biotechnol Bull, 2023, 39(10): 246-255.
[2]	胡芷涵, 张鼎, 胡跃强. 中药附子药理学及毒理学的研究进展 [J]. 湖南中医杂志, 2024, 40(12): 201-205.
	Hu ZH, Zhang D, Hu YQ. Research progress in pharmacology and toxicology of Radix aconiti lateralis [J]. Hunan J Tradit Chin Med, 2024, 40(12): 201-205.
[3]	崔颢, 曲媛, 黄虞枫, 等. 川陕滇乌头属药用植物资源调查 [J]. 中国现代中药, 2022, 24(7): 1264-1271.
	Cui H, Qu Y, Huang YF, et al. Investigation on medicinal plant resources of Aconitum in Sichuan, Shaanxi and Yunnan [J]. Mod Chin Med, 2022, 24(7): 1264-1271.
[4]	缪作清, 李世东, 刘杏忠, 等. 三七根腐病病原研究 [J]. 中国农业科学, 2006, 39(7): 1371-1378.
	Miao ZQ, Li SD, Liu XZ, et al. The causal microorganisms of Panax notoginseng root rot disease [J]. Sci Agric Sin, 2006, 39(7): 1371-1378.
[5]	王飞, 杨瑾, 李雪梦, 等. 一种优化的丹参根腐病离体抗性鉴定方法及应用 [J]. 植物病理学报, 2025: 1-11.
	Wang F, Yang J, Li XM, et al. An optimized in vitro method for evaluating resistance to root rot in Salvia miltiorrhiza and its application [J]. Acta Phytopathol Sin, 2025: 1-11.
[6]	赵林艳, 官会林, 向萍, 等. 白及根腐病植株根际土壤微生物群落组成特征分析 [J]. 生物技术通报, 2022, 38(2): 67-74.
	Zhao LY, Guan HL, Xiang P, et al. Composition features of microbial community in the rhizospheric soil of Bletilla striata with root rot [J]. Biotechnol Bull, 2022, 38(2): 67-74.
[7]	王瑞, 董林林, 徐江, 等. 农田栽参模式中人参根腐病原菌鉴定与防治 [J]. 中国中药杂志, 2016, 41(10): 1787-1791.
	Wang R, Dong LL, Xu J, et al. Identification and prevention of root rot pathogen in model of ginseng cultivated in farmlands [J]. China J Chin Mater Med, 2016, 41(10): 1787-1791.
[8]	陈茂婷, 胡琪琪, 书剑琴, 等. 乌头根腐病病原菌的分离与鉴定 [J]. 微生物学通报, 2020, 47(8): 2450-2457.
	Chen MT, Hu QQ, Shu JQ, et al. Isolation and identification of pathogens causing root rot on Aconitum carmichaelii Debx [J]. Microbiol China, 2020, 47(8): 2450-2457.
[9]	胡党振, 徐嫒媛, 于梦怡, 等. 芽胞杆菌处理对柑桔苗木黄龙病抑制效果 [J]. 中国农业大学学报, 2021, 26(8): 72-83.
	Hu DZ, Xu YY, Yu MY, et al. Inhibitory effect of Bacillus treatment on Citrus Huanglongbing disease in nursery growth plants [J]. J China Agric Univ, 2021, 26(8): 72-83.
[10]	Zhang B, Li XL, Fu J, et al. Production of acetoin through simultaneous utilization of glucose, xylose, and Arabinose by engineered Bacillus subtilis [J]. PLoS One, 2016, 11(7): e0159298.
[11]	姚凤琴, 林发壮, 陈昌铭, 等. 枯草芽胞杆菌Bv17对非洲菊根腐病抑菌作用及防治效果 [J]. 福建农业科技, 2019, 50(12): 55-59.
	Yao FQ, Lin FZ, Chen CM, et al. Antibacterial effect and control effect of Bacillus subtilis Bv17 on root rot of Gerbera jamesonii [J]. Fujian Agric Sci Technol, 2019, 50(12): 55-59.
[12]	Chandrasekaran M, Chun SC. Expression of PR-protein genes and induction of defense-related enzymes by Bacillus subtilis CBR05 in tomato (Solanum lycopersicum) plants challenged with Erwinia carotovora subsp. carotovora [J]. Biosci Biotechnol Biochem, 2016, 80(11): 2277-2283.
[13]	Yánez-Mendizábal V, Usall J, Viñas I, et al. Potential of a new strain of Bacillus subtilis CPA-8 to control the major postharvest diseases of fruit [J]. Biocontrol Sci Technol, 2011, 21(4): 409-426.
[14]	Gan KF, Chen M, Liang TT, et al. Biocontrol effects and underlying mechanism of Bacillus subtilis Pn1 on Panax notoginseng root rot caused by Fusarium solani [J]. Ind Crops Prod, 2025, 229: 120963.
[15]	Baard V, Bakare OO, Daniel AI, et al. Biocontrol potential of Bacillus subtilis and Bacillus tequilensis against four Fusarium species [J]. Pathogens, 2023, 12(2): 254.
[16]	Awan ZA, Shoaib A, Schenk PM, et al. Antifungal potential of volatiles produced by Bacillus subtilis BS-01 against Alternaria solani in Solanum lycopersicum [J]. Front Plant Sci, 2023, 13: 1089562.
[17]	Zou L, Wang Q, Wu RX, et al. Biocontrol and plant growth promotion potential of endophytic Bacillus subtilis JY-7-2L on Aconitum carmichaelii Debx [J]. Front Microbiol, 2023, 13: 1059549.
[18]	Walker BJ, Abeel T, Shea T, et al. Pilon: an integrated tool for comprehensive microbial variant detection and genome assembly improvement [J]. PLoS One, 2014, 9(11): e112963.
[19]	Lowe TM, Eddy SR. tRNAscan-SE a program for improved detection of transfer RNA genes in genomic sequence [J]. Nucleic Acids Res, 1997, 25(5): 955-964.
[20]	Kalvari I, Argasinska J, Quinones-Olvera N, et al. Rfam 13.0: shifting to a genome-centric resource for non-coding RNA families [J]. Nucleic Acids Res, 2018, 46(D1): D335-D342.
[21]	Galperin MY, Vera Alvarez R, Karamycheva S, et al. COG database update 2024 [J]. Nucleic Acids Res, 2025, 53(D1): D356-D363.
[22]	Lombard V, Henrissat B, Garron ML. CAZac: an activity descriptor for carbohydrate-active enzymes [J]. Nucleic Acids Res, 2025, 53(D1): D625-D633.
[23]	Aleksande SA, Balhoff JP, Carbon S. The Gene Ontology knowledgebase in 2026 [J]. Nucleic Acids Res, 2025, 54(D1): D1779-D1792.
[24]	Kanehisa M, Furumichi M, Sato Y, et al. KEGG for taxonomy-based analysis of pathways and genomes [J]. Nucleic Acids Res, 2023, 51(D1): D587-D592.
[25]	Blin K, Shaw S, Kloosterman AM, et al. antiSMASH 6.0: improving cluster detection and comparison capabilities [J]. Nucleic Acids Res, 2021, 49(W1): W29-W35.
[26]	Reddy BVB, Kallifidas D, Kim JH, et al. Natural product biosynthetic gene diversity in geographically distinct soil microbiomes [J]. Appl Environ Microbiol, 2012, 78(10): 3744-3752.
[27]	Gontang EA, Gaudêncio SP, Fenical W, et al. Sequence-based analysis of secondary-metabolite biosynthesis in marine Actinobacteria [J]. Appl Environ Microbiol, 2010, 76(8): 2487-2499.
[28]	Dahar GY, Wang HW, Rajer FU, et al. Comparative genomic analysis of Bacillus atrophaeus HAB-5 reveals genes associated with antimicrobial and plant growth-promoting activities [J]. Front Microbiol, 2024, 15: 1384691.
[29]	Chen L, Xu XX, Sun YX, et al. Surfactin inhibits Fusarium graminearum by accumulating intracellular ROS and inducing apoptosis mechanisms [J]. World J Microbiol Biotechnol, 2023, 39(12): 340.
[30]	Islam T, Rabbee MF, Choi J, et al. Biosynthesis, molecular regulation, and application of bacilysin produced by Bacillus species [J]. Metabolites, 2022, 12(5): 397.
[31]	Han XS, Shen DX, Xiong Q, et al. The plant-beneficial rhizobacterium Bacillus velezensis FZB42 controls the soybean pathogen Phytophthora sojae due to bacilysin production [J]. Appl Environ Microbiol, 2021, 87(23): e01601-e01621.
[32]	Chen ML, Wang H, Zhang C, et al. Recent advances in antimicrobial lipopeptide fengycin secreted by Bacillus: Structure, biosynthesis, antifungal mechanisms, and potential application in food preservation [J]. Food Chem, 2025, 489: 144937.
[33]	Algburi A, Zehm S, Netrebov V, et al. Subtilosin prevents biofilm formation by inhibiting bacterial quorum sensing [J]. Probiotics & Antimicro Prot, 2017, 9(1): 81-90.
[34]	Dinesen C, Vertot M, Jarmusch SA, et al. Subtilosin A production is influenced by surfactin levels in Bacillus subtilis [J]. microLife, 2025, 6: uqae029.
[35]	Kenig M, Abraham EP. Antimicrobial activities and antagonists of bacilysin and anticapsin [J]. J Gen Microbiol, 1976, 94(1): 37-45.
[36]	李腾杰, 梁孙妍, 郭健衡, 等. 特基拉芽胞杆菌XK29挥发物2-甲基丁酸对甘薯长喙壳菌的抑制作用研究 [J]. 微生物学报, 2022, 62(12): 5018-5028.
	Li TJ, Liang SY, Guo JH, et al. Inhibitory effects of volatile 2-methylbutyric acid produced by Bacillus tequilensis XK29 on Ceratocystis fimbriata [J]. Acta Microbiol Sin, 2022, 62(12): 5018-5028.
[37]	Shao JH, Li YC, Li ZF, et al. Participating mechanism of a major contributing gene ysnE for auxin biosynthesis in Bacillus amyloliquefaciens SQR9 [J]. J Basic Microbiol, 2021, 61(6): 569-575.
[38]	Jung J, Ahn S, Kim DH, et al. Triple interactions for induced systemic resistance in plants [J]. Front Plant Sci, 2024, 15: 1464710.
[39]	周江鸿, 夏菲, 仲丽, 等. 黄栌枯萎病拮抗细菌CCBC3-3-1的全基因组测序及比较基因组分析 [J]. 生物技术通报, 2024, 40(7): 235-246.
	Zhou JH, Xia F, Zhong L, et al. Whole genome sequencing and comparative genomic analysis of antagonistic bacterium CCBC3-3-1 against Verticillium dahlia [J]. Biotechnol Bull, 2024, 40(7): 235-246.
[40]	Amna, Xia Y, Farooq MA, et al. Multi-stress tolerant PGPR Bacillus xiamenensis PM₁4 activating sugarcane (Saccharum officinarum L.) red rot disease resistance [J]. Plant Physiol Biochem, 2020, 151: 640-649.
[41]	Wash P, Batool A, Mulk S, et al. Prevalence of antimicrobial resistance and respective genes among Bacillus spp., a versatile bio-fungicide [J]. Int J Environ Res Public Health, 2022, 19(22): 14997.
[42]	Wang J, Xie XY, Li B, et al. Complete genome analysis and antimicrobial mechanism of Bacillus velezensis GX0002980 reveals its biocontrol potential against mango anthracnose disease [J]. Microbiol Spectr, 2025, 13(6): e02685-e02624.
[43]	Ezrari S, Mhidra O, Radouane N, et al. Potential role of rhizobacteria isolated from Citrus rhizosphere for biological control of Citrus dry root rot [J]. Plants, 2021, 10(5): 872.
[44]	Mardanova AM, Fanisovna Hadieva G, Tafkilevich Lutfullin M, et al. Bacillus subtilis strains with antifungal activity against the phytopathogenic fungi [J]. Agric Sci, 2017, 8(1): 1-20.
[45]	Li Y, Zhang X, He K, et al. Isolation and identification of Bacillus subtilis LY-1 and its antifungal and growth-promoting effects [J]. Plants, 2023, 12(24): 4158.
[46]	Han SY, Chen JX, Zhao YJ, et al. Bacillus subtilis HSY21 can reduce soybean root rot and inhibit the expression of genes related to the pathogenicity of Fusarium oxysporum [J]. Pestic Biochem Physiol, 2021, 178: 104916.
[47]	Yang LY, Qu MB, Wang Z, et al. Biochemical properties of a novel cold-adapted GH19 chitinase with three chitin-binding domains from Chitinilyticum aquatile CSC-1 and its potential in biocontrol of plant pathogenic fungi [J]. J Agric Food Chem, 2024, 72(36): 19581-19593.
[48]	孙潇慧, 车程川, 季俊杰, 等. Bacillaene酮还原酶结构域的异源表达及底物特异性分析 [J]. 生物工程学报, 2015, 31(9): 1355-1362.
	Sun XH, Che CC, Ji JJ, et al. Heterologous expression and substrate specificity of ketoreductase domain in bacillaene polyketide synthase [J]. Chin J Biotechnol, 2015, 31(9): 1355-1362.
[49]	Miao S, Liang JH, Xu Y, et al. Bacillaene, sharp objects consist in the arsenal of antibiotics produced by Bacillus [J]. J Cell Physiol, 2023: jcp.30974.
[50]	Angelini LL, dos Santos RAC, Fox G, et al. Pulcherrimin protects Bacillus subtilis against oxidative stress during biofilm development [J]. NPJ Biofilms Microbiomes, 2023, 9: 50.
[51]	戚家明, 孙杉杉, 张东旭, 等. 芽胞杆菌BS-6基于全基因组数据的分类鉴定及拮抗能力分析 [J]. 生物技术通报, 2019, 35(10): 111-118.
	Qi JM, Sun SS, Zhang DX, et al. Identification and biocontrol activity analysis of Bacillus sp. BS-6 based on genome-wide data [J]. Biotechnol Bull, 2019, 35(10): 111-118.
[52]	Sood S, Steinmetz H, Beims H, et al. Paenilarvins: iturin family lipopeptides from the honey bee pathogen Paenibacillus larvae [J]. ChemBioChem, 2014, 15(13): 1947-1955.
[53]	Olishevska S, Nickzad A, Déziel E. Bacillus and Paenibacillus secreted polyketides and peptides involved in controlling human and plant pathogens [J]. Appl Microbiol Biotechnol, 2019, 103(3): 1189-1215.
[54]	McGowan SJ, Sebaihia M, O’Leary S, et al. Analysis of the carbapenem gene cluster of Erwinia carotovora: definition of the antibiotic biosynthetic genes and evidence for a novel β-lactam resistance mechanism [J]. Mol Microbiol, 1997, 26(3): 545-556.
[55]	Ma JY, Song XR, Li MC, et al. Global spread of carbapenem-resistant Enterobacteriaceae: Epidemiological features, resistance mechanisms, detection and therapy [J]. Microbiol Res, 2023, 266: 127249.
[56]	Wen F, Wu YJ, Jin J, et al. Integrating gene knockout, GC-O-MS and multivariate statistical analysis: revealing the impact of fadA and bkdB genes on the aroma characteristics of Bacillus subtilis BJ3-2 fermented soybeans [J]. RSC Adv, 2025, 15(30): 24437-24449.
[57]	Ali SAM, Sayyed RZ, Mir MI, et al. Induction of systemic resistance in maize and antibiofilm activity of surfactin from Bacillus velezensis MS20 [J]. Front Microbiol, 2022, 13: 879739.
[58]	Lam VB, Meyer T, Arias AA, et al. Bacillus cyclic lipopeptides iturin and fengycin control rice blast caused by Pyricularia oryzae in potting and acid sulfate soils by direct antagonism and induced systemic resistance [J]. Microorganisms, 2021, 9(7): 1441.
[59]	Peng G, Zhao XY, Li YZ, et al. Engineering Bacillus velezensis with high production of acetoin primes strong induced systemic resistance in Arabidopsis thaliana [J]. Microbiol Res, 2019, 227: 126297.
[60]	Afzal I, Shinwari ZK, Sikandar S, et al. Plant beneficial endophytic bacteria: Mechanisms, diversity, host range and genetic determinants [J]. Microbiol Res, 2019, 221: 36-49.

病原菌 Pathogen	处理 Treatment	菌落直径 Diameter of fungal colony （cm）
病原菌 Pathogen	处理 Treatment	48 h	72 h	96 h	120 h	144 h	168 h
F. oxysporum-1	CK	2.13±0.09a	3.83±0.07a	5.23±0.07a	6.90±0.06a	8.10±0.06a	8.50±0.00a
	JY-7-2L	1.13±0.03b	2.37±0.07b	3.37±0.17b	4.37±0.18b	6.07±0.32b	7.20±0.26b
F. oxysporum-2	CK	2.08±0.03a	3.50±0.04a	4.55±0.03a	5.85±0.09a	7.13±0.08a	8.08±0.06a
	JY-7-2L	0.93±0.03b	1.85±0.03b	2.48±0.05b	3.45±0.03b	4.48±0.09b	5.50±0.07b

病原菌 Pathogen	处理 Treatment	菌落直径 Diameter of fungal colony （cm）
病原菌 Pathogen	处理 Treatment	48 h	72 h	96 h	120 h	144 h	168 h
F. oxysporum-1	CK	2.13±0.09a	3.83±0.07a	5.23±0.07a	6.90±0.06a	8.10±0.06a	8.50±0.00a
	JY-7-2L	1.13±0.03b	2.37±0.07b	3.37±0.17b	4.37±0.18b	6.07±0.32b	7.20±0.26b
F. oxysporum-2	CK	2.08±0.03a	3.50±0.04a	4.55±0.03a	5.85±0.09a	7.13±0.08a	8.08±0.06a
	JY-7-2L	0.93±0.03b	1.85±0.03b	2.48±0.05b	3.45±0.03b	4.48±0.09b	5.50±0.07b

病原菌 Pathogen	处理 Treatment	菌落直径 Diameter of fungal colony （cm）
病原菌 Pathogen	处理 Treatment	48 h	72 h	96 h	120 h	144 h	168 h	192 h
F. oxysporum-1	CK	1.87±0.07a	3.63±0.24a	4.83±0.07a	5.70±0.09a	6.80±0.13a	7.90±0.23a	8.50±0.00a
	JY-7-2L	1.83±0.03a	2.53±0.24a	3.57±0.33b	4.43±0.07b	4.82±0.03b	5.56±0.09b	6.89±0.17b
F. oxysporum-2	CK	2.55±0.05a	4.13±0.05a	5.28±0.10a	6.38±0.09a	7.23±0.10a	8.08±0.25a	8.50±0.00a
	JY-7-2L	2.23±0.09b	3.53±0.09b	3.53±0.09b	4.83±0.12b	5.55±0.13b	6.18±0.06b	6.55±0.06b

病原菌 Pathogen	处理 Treatment	菌落直径 Diameter of fungal colony （cm）
病原菌 Pathogen	处理 Treatment	48 h	72 h	96 h	120 h	144 h	168 h	192 h
F. oxysporum-1	CK	1.87±0.07a	3.63±0.24a	4.83±0.07a	5.70±0.09a	6.80±0.13a	7.90±0.23a	8.50±0.00a
	JY-7-2L	1.83±0.03a	2.53±0.24a	3.57±0.33b	4.43±0.07b	4.82±0.03b	5.56±0.09b	6.89±0.17b
F. oxysporum-2	CK	2.55±0.05a	4.13±0.05a	5.28±0.10a	6.38±0.09a	7.23±0.10a	8.08±0.25a	8.50±0.00a
	JY-7-2L	2.23±0.09b	3.53±0.09b	3.53±0.09b	4.83±0.12b	5.55±0.13b	6.18±0.06b	6.55±0.06b

基因组富集通路 Genome enrichment pathway	富集基因数量 Number of enriched genes	通路编号 Pathway ID
Novobiocin biosynthesis	5	ko00401
Streptomycin biosynthesis	8	ko00521
Biosynthesis of ansamycins	1	ko01051
Penicillin and cephalosporin biosynthesis	2	ko00311
Vancomycin resistance	7	ko01502
Biosynthesis of vancomycin group antibiotics	1	ko01055
Biosynthesis of various antibiotics	10	ko00998
Acarbose and validamycin biosynthesis	2	ko00525
Cationic antimicrobial peptide resistance	12	ko01503
Carbapenem biosynthesis	3	ko00332
Prodigiosin biosynthesis	5	ko00333