香蕉枯萎病拮抗菌的筛选鉴定与生防作用

doi:10.13560/j.cnki.biotech.bull.1985.2025-0891

摘要/Abstract

摘要：

目的香蕉枯萎病严重影响香蕉的种植面积、产量和品质。为实现香蕉枯萎病绿色防控，筛选获得针对香蕉枯萎病菌（FOC）的生防菌。方法从香蕉根际土壤中筛选对FOC具有抑制作用的拮抗菌，并通过形态学、生理生化特性、16S rRNA基因序列和全基因组序列分析，明确菌株的分类地位并对其可能的生防相关功能基因进行注释分析。同时对拮抗菌开展防病促生特性研究、代谢产物抑菌能力检测、抑菌谱测定以及盆栽防效验证。结果筛选获得1株对FOC具有拮抗作用的细菌GX-13，鉴定为枯草芽胞杆菌（Bacillus subtilis）。GX-13具备溶磷、产氨、产吲哚乙酸，分泌蛋白酶、β-1，3-葡聚糖酶、纤维素酶及铁载体等抑菌促生特性。GX-13的非挥发性代谢物对FOC抑制率达76.85%，可破坏菌丝结构并削弱了菌丝侵袭力；挥发性代谢物和无菌发酵液抑菌作用较弱（20.01%和16.48%）。抑菌谱测定结果显示，GX-13对西瓜枯萎病菌、番茄枯萎病菌和稻瘟病菌等8种植物病原真菌均具有抑制作用。盆栽试验显示，GX-13对香蕉枯萎病的叶片黄化和球茎褐变防治率为68.42%和50.00%，并能促进植株生长，对根生长的促进作用达到极显著水平（P<0.01）。通过基因功能注释和次级代谢产物基因簇分析发现GX-13存在多个真菌细胞壁降解相关酶基因，以及抑菌物质、植物促生和病原菌拮抗等相关基因。结论 GX-13通过直接抑菌、促生和广谱拮抗等多重机制协同作用，实现对香蕉枯萎病的有效防控。

关键词: 香蕉枯萎病, 尖孢镰刀菌, 枯草芽胞杆菌, 拮抗作用, 促生作用

Abstract:

Objective Banana Fusarium wilt, caused by Fusarium oxysporum f. sp. cubense (FOC), is a devastating soil-borne disease that seriously impacts banana production in terms of cultivation area, yield, and fruit quality. This study is aimed to screen effective biocontrol strains against FOC to facilitate the green and sustainable management of this disease. Method Antagonistic bacteria were isolated from the rhizosphere soil of banana plants. A potent isolate, GX-13, was selected and identified based on its morphological, physiological and biochemical characteristics, 16S rRNA gene sequence, and whole genome sequence, with subsequent annotation for potential biocontrol-related genes. Additionally, we assessed its plant-growth-promoting traits (phosphate solubilization, ammonia production, indole-3-acetic acid (IAA) synthesis) and antifungal properties (production of siderophores and cell wall-degrading enzymes). The efficacy of its sterile fermentation supernatant, volatile and non-volatile metabolites against FOC were evaluated in vitro, along with its inhibitory spectrum against other plant pathogenic fungi. Finally, the biocontrol efficacy and plant-growth-promoting ability of GX-13 were validated in a pot experiment. Result A bacterial strain, GX-13, showing antagonistic activity against FOC, was identified as Bacillus subtilis. GX-13 presented plant growth-promoting traits, including phosphate solubilization, ammonia production, indole-3-acetic acid synthesis, and antifungal functional characteristics such as secretion of fungal cell wall-degrading enzymes (protease, β-1,3-glucanase, cellulase) and siderophore production. The non-volatile metabolites of GX-13 caused structural damage to FOC hyphae, resulting in an inhibition rate of 76.85% and significantly reducing hyphal invasiveness. In contrast, its volatile metabolites and sterile fermentation supernatant showed lower inhibition rates (20.01% and 16.48%, respectively). GX-13 also demonstrated a broad antifungal spectrum, effectively inhibiting eight other phytopathogenic fungi, including Fusarium oxysporum f. sp. niveum, Fusarium oxysporum f. sp. lycopersici, Magnaporthe oryzae, Colletotrichum acutatum, Aspergillus carbonarius, Colletotrichum musae, Phyllosticta graminicola and Pseudopestalotiopsis theae. In pot experiments, GX-13 achieved control efficacies of 68.42 % against leaf yellowing and 50.00% against corm browning. Furthermore, it significantly promoted plant growth, with a highly significant enhancement in root development (P<0.01). Gene function annotation and secondary metabolite biosynthetic gene cluster analysis revealed that GX-13 harbored numerous genes involved in the synthesis fungal cell wall-degrading enzymes, antimicrobial compounds, and pathways responsible for plant growth promotion and pathogen antagonism. Conclusion The Bacillus subtilis strain GX-13 effectively controls banana Fusarium wilt through a synergistic mechanisms, including direct antifungal activity, broad-spectrum antagonism, and plant growth promotion.

Key words: banana Fusarium wilt, Fusarium oxysporum f. sp. cubense, Bacillus subtilis, antagonism, growth promotion

林宝妹, 李珊珊, 李海明, 洪佳敏, 张帅, 吴妙鸿, 吴维坚, 吴水金. 香蕉枯萎病拮抗菌的筛选鉴定与生防作用[J]. 生物技术通报, 2026, 42(4): 272-286.

LIN Bao-mei, LI Shan-shan, LI Hai-ming, HONG Jia-min, ZHANG Shuai, WU Miao-hong, WU Wei-jian, WU Shui-jin. Screening and Identification of Antagonistic Bacterium against Banana Fusarium wilt and Its Biocontrol Effects[J]. Biotechnology Bulletin, 2026, 42(4): 272-286.

图/表 18

表1 香蕉枯萎病病害等级划分

Table 1 Disease severity classification of banana Fusarium wilt

评价指标 Evaluation indicator	病害等级 Pathogen severity grading	病害描述 Evaluation content description
叶片黄化度 Leaf yellowing degree	0	叶片无症状
	1	下部1-2片叶黄化，黄化面积≤25%
	3	3-4片叶黄化，25%<黄化面积≤50%
	5	外围叶片黄化，黄化面积>50%，叶片萎蔫
	7	大部分叶片黄化、萎蔫，仅叶心正常
	9	植物萎蔫、枯萎死亡
球茎褐变度 Bulb browning degree	0	球茎组织白色，无褐变
	1	球茎褐变面积≤25%
	3	25%<球茎褐变面积≤50%
	5	50%<球茎褐变面积≤75%
	7	球茎褐变面积>75%

图1 菌株GX-13对FOC的平板抑制作用A：对照；B：菌株GX-13与FOC的平板对峙

Fig. 1 Inhibition effect of strain GX-13 on FOC based on plate confrontation cultureA: Control. B: Plate confrontation between strain GX-13 and FOC

图2 菌株GX-13的形态学特征A：菌落形态；B：革兰氏染色；C：芽孢染色

Fig. 2 Morphological characteristics of strain GX-13A: Colony morphology. B: Gram staining. C: Spore staining

表2 菌株GX-13的生理生化特征

Table 2 Physiological and biochemical characteristics of strain GX-13

试验项目 Tested item		结果 Results	试验项目 Item tested		结果 Results
接触酶反应 Catalase reaction		+	V-P试验 Voges Proskauer test		+
明胶液化 Gelatine liquefication		+	甲基红试验 Methyl red test		-
吲哚试验 Indole test		-	苯丙氨酸脱氨酶 Amphetanine dehydrogenase reaction		-
淀粉水解 Starch hydrolysis		+	产氨试验 Ammonia production test		+
不同碳源 Different carbon sources	葡萄糖 Glucose	+	硝酸盐还原实验 Nitrate reduction reaction		+
	蔗糖 Sucrose	+	柠檬酸盐试验 Citrate test		+
	D-甘露醇 D-mannitol	+	不同氮源 Different nitrogen sources	硝酸钾 KNO₃	+
	D-麦芽糖 D-maltdust	+	不同氮源 Different nitrogen sources	硫酸铵（NH₄）₂SO₄	+

图3 菌株GX-13系统发育分析A：基于16S rRNA序列构建的系统发育树；B：基于全基因组序列构建的系统发育树；括号内数值为NCBI登录号；分枝处数值表示bootstrap值；标尺0.01和0.050代表序列的进化差异

Fig. 3 Phylogenetic analysis of strain GX-13A: The phylogenetic tree based on 16S rRNA sequencing analysis. B: The phylogenetic tree based on whole genome sequencing analysis. Numbers in parentheses are NCBI accession numbers. Numbers at branch nodes present bootstrap value. The scale bar 0.01 and 0.050 indicate sequence variance

表3 菌株GX-13与其相似菌株的ANI比值

Table 3 ANI values of strain GX-13 and its related strains

菌株 Strain	ANI （%）
B. subtilis ATCC6051	99.35
B. cabrialesii TE3	92.41
B. tequilensis ATCC BAA-819	91.71
B. halotolerans ZB201702	87.43
B. velezensis FZB42	77.24
B. amyloliquefaciens ATCC23350	77.02
B. paralicheniformis KJ-16	72.80
B. australimaris PWG	71.01
B. pumilus ATCC 7061	70.99

表4 菌株GX-13对8种植物病原真菌的抑制效果

Table 4 Antifungal activity of strain GX-13 against eight phytopathogenic fungal species

病原菌 Pathogenic fungi	菌落直径 Colony diameter （mm）		抑制率 Inhibition rate （%）
病原菌 Pathogenic fungi	对照 Control	处理 Treatment	抑制率 Inhibition rate （%）
西瓜枯萎病菌 F. oxysporum f. sp. niveum	76.00±0.84	44.04±0.24	42.04±0.89fF
番茄枯萎病菌 F. oxysporum f. sp. lycopersici	79.81±0.33	46.35±0.90	41.93±1.18fF
枇杷炭疽病菌 C. acutatum	77.03±6.04	25.80±0.32	66.38±2.32dD
香蕉炭疽病菌 C. musae	72.53±0.58	28.87±0.12	60.20±0.24eE
水稻稻瘟病菌 M. oryzae	76.33±0.09	7.87±0.11	89.70±0.14aA
葡萄黑霉病菌 A. carbonarius	77.82±5.53	29.83±0.39	61.55±2.41eE
茶叶芽枯病菌 P. graminicola	62.06±0.28	9.34±0.09	84.95±0.09bB
茶叶轮斑病菌 P. theae	70.70±0.56	13.13±1.10	81.43±1.55cC

图4 GX-13生长曲线

Fig. 4 Growth curve of GX-13 strain

表5 菌株GX-13次级代谢产物的抑菌效果

Table 5 Antimicrobial activity of secondary metabolites of strain GX-13

次级代谢物类型 Types of secondary metabolites		菌落直径 Colony diameter （mm）		抑菌率 Inhibition rate （%）
次级代谢物类型 Types of secondary metabolites		对照组 Control	处理组 Treatment	抑菌率 Inhibition rate （%）
无菌发酵滤液 Sterile fermentation filtrate		76.02±0.39	63.50±0.40	16.48±0.79gG
挥发性代谢产物 Volatile metabolites		82.09±0.17	65.66±0.29	20.01±0.30fF
非挥发性代谢产物 Non-volatile metabolites	累积3 d Accumulation for 3 d	82.65±0.3	49.89±1.63	39.64±2.19eE
	累积5 d Accumulation for 5 d	82.52±0.16	32.33±0.2	60.82±0.31dD
	累积7 d Accumulation for 7d	81.75±0.23	18.92±0.94	76.85±1.19aA
	累积9 d Accumulation for 9 d	82.06±0.09	27.36±0.41	66.66±0.53cC
	累积11 d Accumulation for 11 d	81.14±0.17	22.47±0.7	72.31±0.89bB

图5 GX-13的非挥发性代谢物处理后对FOC菌丝穿透能力的影响A：空白PDA平板+玻璃纸+FOC菌块；B：含代谢物的PDA平板+玻璃纸+FOC菌块；C：A去除玻璃纸和菌块后继续培养；D：B去除玻璃纸和菌块后继续培养

Fig. 5 Effects of non-volatile metabolites of GX-13 on mycelial penetration ability of FOCA: Blank PDA plate+cellophane+FOC. B: Metabolite-containing PDA plate+cellophane+FOC. C: Continued cultivation after removal of cellophane and FOC of A. D: Continued cultivation after removal of cellophane and FOC of B

图6 GX-13的非挥发性代谢物处理后FOC菌丝形态

Fig. 6 Effects of non-volatile metabolites of GX-13 on mycelial morphology of FOC

表6 菌株GX-13的防病促生特性

Table 6 Disease prevention and growth-promoting characteristics of strain GX-13

试验项目 Item tested	结果 Results	试验项目 Item tested	结果 Results
纤维素酶 Cellulase	+	解有机磷能力 Organophosphate solubility	-
β-1，3-葡聚糖酶 β-1，3-glucanase	+	解无机磷能力 Inorganic phosphorus solubility	+
蛋白酶 Protease	+	解钾能力 Potassium dissolving effect	-
几丁质酶 Chitinase	-	固氮能力 Nitrogen fixation capacity	-
产铁载体能力 Iron carrier capacity	+	产氨能力 NH₃ production	+

图7 菌株GX-13的产酶促生能力检测A：蛋白酶；B：纤维素酶；C：β-1，3-葡聚糖酶；D：解无机磷能力；E：产铁载体能力；F：产氨能力；G：产IAA能力

Fig. 7 Enzyme production and growth-promoting potential detection of GX-13 strainA: Protease; B: cellulase; C: β-1,3-glucanase; D: inorganic phosphorus solubility; E: iron carrier capacity; F: NH3 production; G: IAA production capacity

图8 不同处理下香蕉幼苗病情等级分布情况CK：空白对照；P：病原菌组；A+P：拮抗菌+病原菌组；C+P：多菌灵+病原菌组

Fig. 8 Distribution of disease severity grades in banana seedlings under different treatmentsCK: Blank control; P: pathogen group; A+P: antagonistic strain+pathogen group; C+P: carbendazim+pathogen group

表7 菌株GX-13对香蕉幼苗的促生效果

Table 7 Growth-promoting effects of strain GX-13 on banana seedlings

指标Index	空白对照组Blank control	拮抗菌组Antagonistic strain group	病原菌组Pathogen group	拮抗菌+病原菌组 Antagonistic strain + Pathogen group
株高 Plant height（cm）	17.45±1.46aA	19.20±2.74aA	12.20±1.57bB	12.95±2.03bB
茎粗 Stem size（mm）	17.06±0.98aA	15.97±1.47abA	10.91±1.67cB	14.76±1.90bA
粗根数量 Number of coarse roots	8.17±1.72bB	11.33±2.07aA	3.83±1.47cC	7.33±1.51bB
叶片数量 Leaf number	8.33±0.58aA	9.33±1.15aA	5.83±0.41cB	8.00±1.00bA
叶长 Leaf length（cm）	18.83±1.63aAB	19.25±2.46aA	15.75±1.51bB	18.33±1.69aAB
叶宽 Leaf width（cm）	8.60±0.78abAB	9.47±1.05aA	7.55±0.97bB	8.65±0.79abAB
叶绿素相对含量 Relative chlorophyll content	53.22±2.86aA	51.60±4.49aA	28.63±14.03bB	47.08±7.87aA

图9 基因组圈图C1，3-5：质粒基因组，C2：染色体基因组；最外面一圈为基因组大小的标示，每个刻度为5 kb；第2圈和第3圈分别为基因组正链和负链上的gene，不同颜色代表不同的COG功能分类；第4圈为重复序列；第5圈为tRNA和rRNA，蓝色为tRNA，紫色是rRNA；第6圈为GC含量，黄色部分表示该区域GC含量高于基因组的平均GC含量，峰值越高则与平均GC含量差异越大，蓝色部分则表示该区域GC含量低于基因组的平均GC含量；最内圈是GC-skew，深灰色代表G含量大于C的区域，红色代表C含量大于G的区域

Fig. 9 Genomic circle mapC1, 3-5: Plasmid genome, C2: Chromosome genome. The outermost circle indicates the size of the genome, with each scale being 5 kb; the second and third circles indicate genes on the positive and negative strands of the genome, with different colors representing different COG functional classifications. The fourth circle is a repeated sequence. The fifth circle indicates tRNA and rRNA, with blue indicating tRNA and purple indicating rRNA. The sixth circle indicates GC content. The yellow part indicates that the GC content in this region is higher than the average GC content of the genome. The higher the peak value, the greater the difference from the average GC content. The blue part indicates that the GC content in this region is lower than the average GC content of the genome. The innermost circle is GC-skew, where dark gray indicates the area where G content is greater than C, and red indicates the area where C content is greater than G

图10 基因功能注释结果A：KEGG注释分类统计图；B：注释CAZy家族分布

Fig. 10 Gene functional annotation resultsA: Statistics of pathways annotated in KEGG. B: CAZy family distribution of annotated enzymes

表 8 菌株GX-13次级代谢产物合成基因簇预测结果

Table 8 Predicted secondary metabolite biosynthetic gene clusters in strain GX-13

基因簇 ID Cluster ID	类型Type	起始Start	终止End	相似产物Similar production	相似度 Similar percentage （%）
1	核糖体肽类 Ranthipeptide， sactipeptide	203 800	225 871	产孢杀伤因子 Sporulation killing factor	100
2	非核糖体多肽类 NRPS	357 933	421 373	表面活性素 Surfactin	82
3	萜类 Terpene	1 132 993	1 153 506	-	-
4	反式AT型聚酮合酶、类聚酮合酶、Ⅲ型聚酮合酶和非核糖体合成肽transAT-PKS， PKS-like， T3PKS， NRPS	1 746 963	1 852 223	杆菌烯 Bacillaene	100
5	非核糖体多肽类、β-内酯类 NRPS， Betalactone	1 923 840	2 000 876	丰原素 Fengycin	100
6	萜类 Terpene	2 079 114	2 101 012	-	-
7	Ⅲ型聚酮合酶 T3PKS	2 149 489	2 190 586	-	-
8	非核糖体多肽类 NRPS	3 089 896	3 137 032	杆菌素 Bacillibactin	100
9	环二肽合成酶 CDPS	3 423 235	3 443 981	-	-
10	核糖体肽类 Sactipeptide	3 655 591	3 677 202	枯草菌素 A Subtilosin A	100
11	其他 Other	3 680 174	3 721 592	杆溶菌素 Bacilysin	100
12	表修饰肽 Epipeptide	3 940 581	3 962 279	泰兰他汀A Thailanstatin A	10

参考文献 40

[1]	de Jesus Rocha A, da Silva Soares JM, dos Santos Nascimento F, et al. Improvements in the resistance of the banana species to Fusarium wilt: a systematic review of methods and perspectives [J]. J Fungi, 2021, 7(4): 249.
[2]	Ruan YN, Nong CH, Jintrawet A, et al. A smooth vetch (Vicia villosa Var.) strain endogenous to the broad-spectrum antagonist Bacillus siamensis JSZ06 alleviates banana wilt disease [J]. Front Plant Sci, 2024, 15: 1410197.
[3]	Li CL, Xiang DD, Yang S, et al. Fusarium wilt of banana latency and onset detection based on visible/near infrared spectral technology [J]. Agronomy, 2024, 14(12): 2994.
[4]	赵阳, 刘爽, 王志彪, 等. 香蕉枯萎病菌候选效应蛋白FoSSP80能抑制植物免疫反应 [J]. 热带生物学报, 2025, 16(1): 87-97.
	Zhao Y, Liu S, Wang ZB, et al. Functional characterization of candidate effector FoSSP80 in Fusarium oxysporum f. sp. cubense [J]. J Trop Biol, 2025, 16(1): 87-97.
[5]	张妙宜, 周登博, 起登凤, 等. 香蕉枯萎病综合防控研究进展 [J]. 中国科学: 生命科学, 2024, 54(10): 1843-1852.
	Zhang MY, Zhou DB, Qi DF, et al. Research progress on the integrated control of Fusarium wilt disease in banana [J]. Sci Sin Vitae, 2024, 54(10): 1843-1852.
[6]	李华平, 李云锋, 聂燕芳. 香蕉枯萎病的发生及防控研究现状 [J]. 华南农业大学学报, 2019, 40(5): 128-136.
	Li HP, Li YF, Nie YF. Research status of occurrence and control of Fusarium wilt of banana [J]. J South China Agric Univ, 2019, 40(5): 128-136.
[7]	Shen NK, Li SY, Li SY, et al. The siderophore-producing bacterium, Bacillus siamensis Gxun-6, has an antifungal activity against Fusarium oxysporum and promotes the growth of banana [J]. Egypt J Biol Pest Control, 2022, 32(1): 34.
[8]	Hasan M, Hossain M, Jiang D. New endophytic strains of Trichoderma promote growth and reduce clubroot severity of rapeseed (Brassica napus) [J]. PLoS One, 2023, 18(10): e0287899.
[9]	De Simone N, Capozzi V, Amodio ML, et al. Microbial-based biocontrol solutions for fruits and vegetables: recent insight, patents, and innovative trends [J]. Recent Pat Food Nutr Agric, 2021, 12(1): 3-18.
[10]	Bubici G, Kaushal M, Prigigallo MI, et al. Corrigendum: biological control agents against Fusarium wilt of banana [J]. Front Microbiol, 2019, 10: 1290.
[11]	Yin C, Casa Vargas JM, Schlatter DC, et al. Rhizosphere community selection reveals bacteria associated with reduced root disease [J]. Microbiome, 2021, 9(1): 86.
[12]	Guo YQ, Tian L, Zhu XY, et al. Antagonism of Bacillus velezensis ZGE166 against the pathogenic fungi causing corm rot disease in saffron (Crocus sativus L.) [J]. Microb Ecol, 2025, 88(1): 40.
[13]	Vibha R, Granada DL, Skariyachan S, et al. In vitro and in silico investigation deciphering novel antifungal activity of endophyte Bacillus velezensis CBMB205 against Fusarium oxysporum [J]. Sci Rep, 2025, 15: 684.
[14]	Qi D, Liu Q, Zou L, et al. Taxonomic identification and antagonistic activity of Streptomyces luomodiensis sp. nov. against phytopathogenic fungi [J]. Front Microbiol, 2024, 15: 1402653.
[15]	Hernández-Melchor DJ, Guerrero-Chávez AC, Ferrera-Rodríguez MR, et al. Cellulase and chitinase activities and antagonism against Fusarium oxysporum f.sp. cubense race 1 of six Trichoderma strains isolated from Mexican maize cropping [J]. Biotechnol Lett, 2023, 45(3): 387-400.
[16]	Solórzano R, Ramírez Maguiña HA, Johnson L, et al. Current progress in microbial biocontrol of banana Fusarium wilt: a systematic review [J]. Agronomy, 2025, 15(3): 619.
[17]	Bagy HMMK, Hassan EA, Nafady NA, et al. Efficacy of arbuscular mycorrhizal fungi and endophytic strain Epicoccum nigrum ASU11 as biocontrol agents against blackleg disease of potato caused by bacterial strain Pectobacterium carotovora subsp. atrosepticum PHY7 [J]. Biol Control, 2019, 134: 103-113.
[18]	吴巳雨. 长枝木霉HK16-1抗香蕉枯萎病活性物质的挖掘及抑菌效果分析 [D]. 武汉: 华中农业大学, 2024.
	Wu SY. Extraction of active substance from Trichoderma longibranchus HK16-1 against banana Fusarium wilt and analysis of antifungal effect [D]. Wuhan: Huazhong Agricultural University, 2024.
[19]	Samaras A, Roumeliotis E, Ntasiou P, et al. Bacillus subtilis MBI600 promotes growth of tomato plants and induces systemic resistance contributing to the control of soilborne pathogens [J]. Plants, 2021, 10(6): 1113.
[20]	张德政, 聂太礼, 杨军, 等. 棉花黄萎病高效拮抗细菌的筛选及其防治效果 [J]. 湖北农业科学, 2024, 63(12): 67-72.
	Zhang DZ, Nie TL, Yang J, et al. Screening and control effect of highly effective antagonistic bacteria against cotton verticillium wilt [J]. Hubei Agric Sci, 2024, 63(12): 67-72.
[21]	东秀珠, 蔡妙英. 常见细菌系统鉴定手册 [M]. 北京: 科学出版社, 2001.
	Dong XZ, Cai MY. Manual for common bacterial system identification [M]. Beijing: Science Press, 2001.
[22]	赵斌, 何绍江. 微生物学实验 [M]. 北京: 科学出版社, 2002.
	Zhao B, He SJ. Microbiological experiment [M]. Beijing: Science Press, 2002.
[23]	毕钰, 张琳, 余红凤, 等. 小麦赤霉病拮抗菌JB7的拮抗活性及鉴定 [J]. 微生物学报, 2024, 64(5): 1580-1592.
	Bi Y, Zhang L, Yu HF, et al. Antagonistic activity and identification of antagonistic bacteria JB7 against Fusarium head blight [J]. Acta Microbiol Sin, 2024, 64(5): 1580-1592.
[24]	杨学宇, 谭琳, 沈程文, 等. 茶轮斑病拮抗细菌kc-16的抑菌性能及田间防效 [J]. 微生物学通报, 2024, 51(11): 4560-4573.
	Yang XY, Tan L, Shen CW, et al. Antimicrobial activity and field control effect of an antagonistic bacterial strain kc-16 against tea gray blight [J]. Microbiol China, 2024, 51(11): 4560-4573.
[25]	李文博, 张新祺, 赵亚婷, 等. 采前喷施壳寡糖对采后西梅黑斑病的控制 [J]. 食品科学, 2025, 46(7): 283-291.
	Li WB, Zhang XQ, Zhao YT, et al. Control of postharvest prune black spot disease by preharvest chitosan oligosaccharide spraying [J]. Food Sci, 2025, 46(7): 283-291.
[26]	黎晓丹,方献平,李水根,等.草莓果生炭疽菌效应子基因CfE4对产孢量及菌丝穿透能力的影响[J/OL].分子植物育种,1-11[2025-12-12].,
	Fang XP, Li SG, et al. The effect of effector gene CfE4 on conidia production and penetration of Colletotrichum fructicola in strawberry[J/OL]. [2025-5-27]. .
[27]	Hyder S, Gondal AS, Rizvi ZF, et al. Characterization of native plant growth promoting rhizobacteria and their anti-oomycete potential against Phytophthora capsici affecting chilli pepper (Capsicum annum L.) [J]. Sci Rep, 2020, 10: 13859.
[28]	甘林, 代玉立, 刘晓菲, 等. 香蕉枯萎病高效拮抗土著细菌的筛选及其防效 [J]. 西北农林科技大学学报: 自然科学版, 2024, 52(6): 95-105.
	Gan L, Dai YL, Liu XF, et al. Screening and control efficacies of highly effective antagonistic native bacteria against banana Fusarium wilt [J]. J Northwest A F Univ Nat Sci Ed, 2024, 52(6): 95-105.
[29]	Wang JR, Qin S, Fan RD, et al. Plant growth promotion and biocontrol of leaf blight caused by Nigrospora sphaerica on passion fruit by endophytic bacillus subtilis strain GUCC4 [J]. J Fungi, 2023, 9(2): 132.
[30]	Sun WL, Shahrajabian MH, Soleymani A. The roles of plant-growth-promoting rhizobacteria (PGPR)-based biostimulants for agricultural production systems [J]. Plants, 2024, 13(5): 613.
[31]	黄勋, 刘霞, 邓琳梅, 等. 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 X, Liu X, Deng LM, et al. Identification and study on its biocontrol and growth promoting properties of Bacillus velezensis 1X1Y against potato common scab [J]. Acta Hortic Sin, 2025, 52(1): 229-246.
[32]	Yang P, Yuan P, Liu WS, et al. Plant growth promotion and plant disease suppression induced by Bacillus amyloliquefaciens strain GD4a [J]. Plants, 2024, 13(5): 672.
[33]	邓玉莲, 谭琳, 吉进军, 等. 茶树根腐病拮抗真菌的筛选、鉴定及防病促生特性研究 [J]. 中国生物防治学报, 2024, 40(4): 914-926.
	Deng YL, Tan L, Ji JJ, et al. Screening and identification of antagonistic fungi against root rot of tea plants and their effect on disease control and plant growth promotion [J]. Chin J Biol Control, 2024, 40(4): 914-926.
[34]	Dutilloy E, Arias AA, Richet N, et al. Bacillus velezensis BE2 controls wheat and barley diseases by direct antagonism and induced systemic resistance [J]. Appl Microbiol Biotechnol, 2024, 108(1): 64.
[35]	Dai MM, Liu R, Jiang H, et al. Volatile organic compounds of Bacillus pumilus strain S1-10 exhibit fumigant activity against Meloidogyne incognita [J]. Plant Dis, 2023, 107(10): 3057-3063.
[36]	Guevara-Avendaño E, Bravo-Castillo KR, Monribot-Villanueva JL, et al. Diffusible and volatile organic compounds produced by avocado rhizobacteria exhibit antifungal effects against Fusarium kuroshium [J]. Braz J Microbiol, 2020, 51(3): 861-873.
[37]	刘高强, 李新鹏, 刘文钊, 等.一株高效降解血液蛋白的枯草芽孢杆菌NWMCC0137全基因组测序及分析 [J]. 江苏农业学报, 2023, 39(7): 1460-1471.
	Liu GQ, Li XP, Liu WZ, et al. Whole genome sequencing and analysis of Bacillus subtilis NWMCC0137, an efficient blood protein degrading strain [J]. Jiangsu J Agric Sci, 2023, 39(7): 1460-1471.
[38]	万青. 一株贝莱斯芽孢杆菌对香蕉枯萎病生防作用的研究 [D]. 海口: 海南大学, 2021.
	Wan Q. Study on the biocontrol effect of a Bacillus velezensis on banana Fusarium wilt [D]. Haikou: Hainan University, 2021.
[39]	李恒, 畅文军, 陈汉清, 等. 香蕉枯萎镰刀菌4号生理小种mon1基因敲除转化子的表型分析及致病力测定 [J]. 热带生物学报, 2019, 10(2): 127-134.
	Li H, Chang WJ, Chen HQ, et al. Functional analysis of the mon1 gene in Fusarium oxysporum f.sp. cubense Race 4 [J]. J Trop Biol, 2019, 10(2): 127-134.
[40]	王田, 陈代朋, 高雅, 等. 香蕉枯萎病菌效应蛋白研究进展 [J]. 农业生物技术学报, 2022, 30(8): 1614-1621.
	Wang T, Chen DP, Gao Y, et al. Research progress on the effector of Fusarium oxysporum f.sp. cubense [J]. J Agric Biotechnol, 2022, 30(8): 1614-1621.