微生物学报

Strain	Year	Biocontrol effect	Action mechanism	References
Bacillus subtilis BS-Z15	2019	BS-Z15 can effectively reduce the incidence of Verticillium wilt of cotton	Strain BS-Z15 secreted antagonistic active substances	[33]
Bacillus malacitensis Z-5	2019	Z-5 strain had remarkable control effect on Verticillium wilt of cotton	Z-5 strain can secrete lipopeptide antibiotics such as surfactant B and ictilicin A	[34]
Bacillus velezensis SZAD1	2020	The control efficiency of SZAD1 strain against Verticillium wilt was 60.10% and 56.00% in seed soaking and root irrigation, respectively	SZAD1 can produce cellulase and chitinase, which can reduce the ability of VD080 to settle cotton stems	[35]
Bacillus velezensis SZAD2	2020	The control efficiency of seed treatment was 60.31%, and that of root irrigation was 79.19%	The strain could systematically colonize the roots and induced systemic resistance of cotton roots by accumulating hydrogen peroxide in the roots and leaves	[36]
Bacillus circulans GN03	2021	Strain GN03 had good resistance to Verticillium wilt of cotton, with the highest control efficiency reaching 78%	GN03 inoculation altered the microflora in and around the plant roots, resulting in a significant accumulation of growth-related hormones	[37]
Bacillus amyloliquefaciens 489-2-2	2021	The control efficiency of seed treatment was 54.99%, and that of root irrigation was 60.31%	489-2-2 caused the mycelium of cotton verticillium wilt to lose pathogenicity, enhancing the systemic resistance of the plant by activating a large number of defense enzymes	[38]
Bacillus velezensis ND	2022	Application of ND fermentation liquid can increase the disease prevention effect from 36.00% to 92.99%	ND also has the activities of protease, cellulase and iron carrying, and has the ability to synthesize indole acetic acid, nitrogen fixation and phosphorus reduction	[39]
Bacillus velezensis EBV02	2022	The highest control effect of EBV02 on cotton Verticillium wilt was 68.33% and 37.25% in greenhouse and field tests, respectively	EBV02 inhibited the mycelia growth of Verticilliumdahliae, and induced active oxygen species outbreak and callus accumulation in cotton leaves	[40]
Bacillus T6	2023	The inhibition rate of T6 strain on Verticillium dahliae was 63.79%	T6 strain can produce volatile organic compound styrene, which can up-regulate the expression of some hydrolase genes in Charlottesia	[41]
Bacillus amyloliquefaciens YZU-SG 146	2023	The control effect of YZU-SG146 against Verticillium wilt of cotton was 84.21%, and it also promoted the growth of root length and seedling length of cotton seeds and seedlings	G146 can secrete ferric carrier, indoleacetic acid, cellulase, protease and amylase, and can trigger the outbreak of reactive oxygen species in cotton leaves	[42]
Bacillus velezensis BvZ45-1	2024	The indoor and field control efficiency of BvZ45-1 against Verticillium wilt of cotton was 46.53% and 47.27%, respectively	The bacterium can produce oxalate decarboxylase, inhibit the spore production of Verticilliumdahliae, and lead to mycelium rupture, cell membrane rupture and cell death	[6]
Bacillus altitudinis KRS010	2024	The effect of KRS010 strain on cotton Verticillium wilt was 93.59%	KRS010 induces plant immunity by inducing systemic resistance activated by salicylic acid and jasmonic acid signaling pathways	[32]

Strain	Year	Biocontrol effect	Action mechanism	References
Bacillus subtilis BS-Z15	2019	BS-Z15 can effectively reduce the incidence of Verticillium wilt of cotton	Strain BS-Z15 secreted antagonistic active substances	[33]
Bacillus malacitensis Z-5	2019	Z-5 strain had remarkable control effect on Verticillium wilt of cotton	Z-5 strain can secrete lipopeptide antibiotics such as surfactant B and ictilicin A	[34]
Bacillus velezensis SZAD1	2020	The control efficiency of SZAD1 strain against Verticillium wilt was 60.10% and 56.00% in seed soaking and root irrigation, respectively	SZAD1 can produce cellulase and chitinase, which can reduce the ability of VD080 to settle cotton stems	[35]
Bacillus velezensis SZAD2	2020	The control efficiency of seed treatment was 60.31%, and that of root irrigation was 79.19%	The strain could systematically colonize the roots and induced systemic resistance of cotton roots by accumulating hydrogen peroxide in the roots and leaves	[36]
Bacillus circulans GN03	2021	Strain GN03 had good resistance to Verticillium wilt of cotton, with the highest control efficiency reaching 78%	GN03 inoculation altered the microflora in and around the plant roots, resulting in a significant accumulation of growth-related hormones	[37]
Bacillus amyloliquefaciens 489-2-2	2021	The control efficiency of seed treatment was 54.99%, and that of root irrigation was 60.31%	489-2-2 caused the mycelium of cotton verticillium wilt to lose pathogenicity, enhancing the systemic resistance of the plant by activating a large number of defense enzymes	[38]
Bacillus velezensis ND	2022	Application of ND fermentation liquid can increase the disease prevention effect from 36.00% to 92.99%	ND also has the activities of protease, cellulase and iron carrying, and has the ability to synthesize indole acetic acid, nitrogen fixation and phosphorus reduction	[39]
Bacillus velezensis EBV02	2022	The highest control effect of EBV02 on cotton Verticillium wilt was 68.33% and 37.25% in greenhouse and field tests, respectively	EBV02 inhibited the mycelia growth of Verticilliumdahliae, and induced active oxygen species outbreak and callus accumulation in cotton leaves	[40]
Bacillus T6	2023	The inhibition rate of T6 strain on Verticillium dahliae was 63.79%	T6 strain can produce volatile organic compound styrene, which can up-regulate the expression of some hydrolase genes in Charlottesia	[41]
Bacillus amyloliquefaciens YZU-SG 146	2023	The control effect of YZU-SG146 against Verticillium wilt of cotton was 84.21%, and it also promoted the growth of root length and seedling length of cotton seeds and seedlings	G146 can secrete ferric carrier, indoleacetic acid, cellulase, protease and amylase, and can trigger the outbreak of reactive oxygen species in cotton leaves	[42]
Bacillus velezensis BvZ45-1	2024	The indoor and field control efficiency of BvZ45-1 against Verticillium wilt of cotton was 46.53% and 47.27%, respectively	The bacterium can produce oxalate decarboxylase, inhibit the spore production of Verticilliumdahliae, and lead to mycelium rupture, cell membrane rupture and cell death	[6]
Bacillus altitudinis KRS010	2024	The effect of KRS010 strain on cotton Verticillium wilt was 93.59%	KRS010 induces plant immunity by inducing systemic resistance activated by salicylic acid and jasmonic acid signaling pathways	[32]

生防微生物在棉花黄萎病防治中的研究进展

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刘延财 ¹^,² , 唐叶 ²^,³ , 吴家和 ² , 宋宪亮 ¹^,^* , 刘钢 ²^,³^,^*

微生物学报 | 综述 2025,65(3): 994-1006

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微生物学报 | 综述 2025, 65(3): 994-1006

生防微生物在棉花黄萎病防治中的研究进展

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刘延财¹^,², 唐叶²^,³, 吴家和², 宋宪亮¹^,^*, 刘钢²^,³^,^*

作者信息

1 山东农业大学农学院，山东泰安

2 中国科学院微生物研究所真菌学国家重点实验室，北京

3 中国科学院大学生命科学学院，北京

Research progress of biocontrol microbial strains in prevention of cotton wilt disease

Yancai LIU¹^,², Ye TANG²^,³, Jiahe WU², Xianliang SONG¹^,^*, Gang LIU²^,³^,^*

Affiliations

1 College of Agronomy, Shandong Agricultural University, Tai’an, Shandong, China

2 State Key Laboratory of Mycology, Institute of Microbiology, Chinese Academy of Sciences, Beijing, China

3 College of Life Sciences, University of Chinese Academy of Sciences, Beijing, China

出版时间: 2025-03-04 doi: 10.13343/j.cnki.wsxb.20240682

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摘要

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黄萎病是影响棉花种植业最重要的病害之一，可导致棉花减产甚至绝收。该病由丝状真菌大丽轮枝菌引发，属土传病害。传统的化学防治方法不仅影响人类健康，还带来环境污染问题，且连年使用易导致大丽轮枝菌产生抗药性。因此，研发针对棉花黄萎病的绿色环保、可持续发展的防治策略迫在眉睫，其中生物防治成为了一个优选方案。本文通过分析国内外最新研究进展，探讨了棉花黄萎病生防微生物菌株的筛选、作用机制及田间应用方式等，总结了生防微生物通过竞争、抗生作用、诱导植物防御反应等多种机制抑制病原菌生长的研究成果。尽管生防微生物的应用前景广阔，但仍面临环境适应性、稳定性和使用成本等挑战。未来研究应更加聚焦于生防微生物菌株的遗传改良、复配菌剂的研制和应用技术的优化，以进一步提升生防微生物菌株在农业生产中的实用性和有效性。

关键词

生防微生物 / 棉花黄萎病 / 大丽轮枝菌 / 作用机制 / 应用

Abstract

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Cotton Verticillium wilt is the most serious disease affecting cotton cultivation, which could cause a significant decrease in cotton yield or even complete crop failure. Cotton Verticillium wilt is caused by the filamentous fungus Verticillium dahliae. The traditional chemical control affects public health and brings about environmental pollution, and the continuous usage has induced the drug resistance of Verticillium dahliae. Therefore, it is urgent to develop environmental friendly and sustainable development control strategies against cotton Verticillium wilt. Biological control has become a good choice to prevent cotton Verticillium wilt. Based on the analysis of the recent research progress, this review discussed the screening, mechanism of action and field application of biocontrol microbial strains against cotton Verticillium wilt, and summarized the research progress of biocontrol microorganisms inhibiting the growth of pathogen through various mechanisms such as competition, antibiotic action, and inducing plant defense response. Although the application prospects of biocontrol microorganisms are expected, they still face challenges such as environmental adaptability, stability, and usage costs of these biocontrol microorganisms. To further improve the practicality of biocontrol microbial strains in agricultural production, future research should focus on genetic improvement of biocontrol microorganisms, development, and application of the microbial agents and so on.

Key words

biocontrol microbial strains / cotton wilt disease / Verticillium dahliae / action mechanism / application

引用本文

刘延财, 唐叶, 吴家和, 宋宪亮, 刘钢. 生防微生物在棉花黄萎病防治中的研究进展. 微生物学报, 2025 , 65 (3) : 994 -1006 . DOI: 10.13343/j.cnki.wsxb.20240682

Yancai LIU, Ye TANG, Jiahe WU, Xianliang SONG, Gang LIU. Research progress of biocontrol microbial strains in prevention of cotton wilt disease[J]. Acta Microbiologica Sinica, 2025 , 65 (3) : 994 -1006 . DOI: 10.13343/j.cnki.wsxb.20240682

正文

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棉花属于锦葵科棉属的一年生草本植物或亚灌木植物，是世界上重要的纤维和油料作物之一；我国自宋朝时期起便开始种植棉花，目前棉花种植主要集中在新疆、黄河流域、长江流域等地^[1]。其中，陆地棉(Gossypium hirsutum L.)是我国种植最广泛的品种，原产于美洲墨西哥，后被引入世界各地；棉花作为我国乃至全球范围内的重要经济作物，关乎国计民生，其作为大宗农产品影响着世界纺织品贸易格局；同其他农作物一样，棉花也容易遭受多种植物病虫害的侵袭；其中由丝状真菌大丽轮枝菌(Verticillium dahliae)侵染所导致的棉花黄萎病(cotton wilt disease)可造成棉花大面积减产甚至绝收，成为影响棉花产量及品质的主要病害之一，被誉为棉花的“癌症”，每年因黄萎病造成的经济损失高达数十亿元，严重阻碍了我国棉花种植业的发展^[2-3]。大丽轮枝菌是一种土传病原真菌，通过植株受损的根部侵入宿主体内，破坏棉花的维管束，进而导致茎内木质部以及叶片脉间组织变色，严重时还会导致叶片脱落，植株枯死。目前，针对棉花黄萎病的防治手段主要包括抗病品种的选育、农业栽培措施、化学农药防治和生物防治等；相较于其他防治手段，生物防治具有易实施、无污染、可持续等优点，已被农业生产广泛接受^[4]。

1 大丽轮枝菌及其致病机理

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大丽轮枝菌(Verticillium dahliae)是隶属于子囊菌门淡色菌科轮枝菌属的一种土传植物病原真菌，其致病力强，寄主范围广，能够危害包括棉花、番茄、茄子、辣椒、果树等在内的 660多种植物^[5]。在生长条件不适宜或无寄主植物存在的情况下，大丽轮枝菌会形成黑色的、形状不规则的微菌核，这些微菌核在土壤中能够存活长达10-15年。当生长条件适宜时，微菌核会重新萌发并长出菌丝，通过宿主植株的根毛或受损根部侵入，并延伸至维管束，进而侵染整个植株^[6]。

目前，关于大丽轮枝菌的致病机理，人们普遍接受的有“组织障碍假说”和“毒素假说”(图1)。“组织障碍假说”认为，大丽轮枝菌会吸附在棉花植株的根系上，受到根系分泌物的刺激后开始萌发，生长出的菌丝接触到棉花根系后，通过根毛或伤口侵入植株的表层、皮层、内皮层，然后进入木质部，在木质部中菌丝快速延伸生长并产生大量孢子，同时刺激邻近的薄壁细胞分泌大量胶状物质，导致导管中产生树胶和凝胶体聚集，从而阻塞了植物的木质部导管，造成植物体内水分和营养物质的运输障碍；同时由于植物的蒸腾作用，大丽轮枝菌的孢子会随着维管束系统扩散到植物的其他部位，扩大了侵染范围，最终使被侵染的植株出现萎蔫、黄化，并导致植株死亡^[7-9]。“毒素假说”则认为，大丽轮枝菌在侵染寄主后，会分泌毒素和其他物质，如蛋白质-脂多糖复合物、糖蛋白和细胞壁降解酶等；这些毒素物质能够破坏棉花植株叶片及根系组织的细胞膜，改变细胞膜的通透性，导致细胞内的钠、钾等离子大量渗漏，破坏电解质平衡，从而引起植物组织结构变化、维管束堵塞等，最终导致棉花植株萎蔫^[10-11]。

2 棉花黄萎病的防治方法

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目前，棉花黄萎病的防治手段主要包括抗病棉花品种的选育、农业栽培措施的改良、化学农药防治以及生物防治等。长期的生产实践证明，发掘和培育抗黄萎病的棉花新品种是世界范围内防治棉花黄萎病的根本举措；由于陆地棉中高抗黄萎病的种质资源匮乏，且大丽轮枝菌生理小种多样，使得抗病棉花品种的选育变得尤为困难^[12]。施用化学农药虽然可以在短时间内抑制植物病害，但同时会对环境和人体健康造成极大的危害；此外，化学农药在控制黄萎病方面并不容易，因为一旦大丽轮枝菌进入植物木质部，化学农药就会失去作用；长期使用化学农药会导致大丽轮枝菌抗药性的出现^[13]。相比之下，生物防治对环境友好，对人畜无害，且抑制病原微生物的范围广，被认为是未来最具发展潜力的一种植物病害防治方法。在生物防治中，尤其是生防微生物的应用，因其易于使用、绿色环保和可持续的特性，正逐渐成为该领域研究和应用的热点。

生防微生物是指从土壤、植物根际或植物体内筛选分离出的能够有效抑制病原菌的微生物，具有发展成为微生物菌剂的潜力。这些生防微生物的种类包括生防细菌、生防真菌和生防放线菌等，它们通过与病原菌竞争生存空间和营养、产生抑制病原菌生长和代谢的小分子化合物、诱导植物自身产生抗病性以及促进植物生长等方式起到预防和抵抗病原菌侵染的作用^[14]。

3 生防微生物在棉花黄萎病防治中的应用

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3.1 生防真菌

众多真菌对棉花黄萎病展现出良好的防治效果，例如木霉(Trichoderma spp.)、黄色篮状霉(Talaromyces flavus)、球毛壳菌(Chaetomium globosum)、尖孢镰孢菌(Fusarium oxysporum)^[15]、黏帚霉(Gliocladium spp.)^[16]、黑壳霉(Gibellulopsis nigrescens)^[17]等。其中，木霉、黄色篮状霉和球毛壳菌在文献中被频繁报道。

孟娜等^[18]对筛选出的4株木霉菌株进行抑菌试验，发现它们能有效抑制大丽轮枝菌菌丝的伸长。孙艳等^[19]通过田间试验发现，木霉菌剂能显著降低棉花黄萎病的发病率，防治效果超过30%，且随着使用年限的增加，防治效果更佳；木霉菌还能促进棉花出苗，提高产量，改善纤维品质。Fravel等^[20]和Kim等^[21-22]的研究表明，黄色篮状霉能产生多种具有抗菌活性的代谢产物，这些产物能抑制大丽轮枝菌菌丝生长、微菌核黑色素的形成以及微菌核萌发，并通过直接在微菌核内繁殖导致微菌核裂解。张芸等^[23]对黄萎病拮抗菌球毛壳菌的研究显示，该菌的非挥发性物质可以完全抑制大丽轮枝菌的生长，对分生孢子的抑制率高达70.5%。随着对木霉、黄色篮状霉等为代表的生防真菌的深入研究，它们在棉花黄萎病防治中的广应用前景将更加广阔。

3.2 生防放线菌

放线菌能够产生结构各异、活性广泛的代谢产物，成为创新药物的重要来源^[24]。近年来，放线菌也被作为生防微生物用于植物病虫害的防治^[25]。放线菌产生的代谢产物通过干扰病原菌细胞壁的形成、DNA和蛋白质合成以及能量代谢等过程，干扰病原真菌或病原细菌的正常生长和发育^[26]。这些活性次级代谢产物通常具有广泛的抗菌谱，能够同时对多种植物病原真菌或细菌产生抑制作用，阻止它们的生长，并可导致这些病原体的形态结构发生改变；在放线菌中研究最多的是链霉菌(Streptomyces spp.)。链霉菌不仅能够产生多种具有拮抗作用的次级代谢产物，还能够产生纤维素酶和几丁质酶，降解病原真菌细胞壁，分泌生长刺激素促进植物生长等^[27]。钱瑶等^[28]对一株分离自植物根际土壤的产二素链霉菌(Streptomyces ambofaciens)的研究表明，该菌株可以产生次级代谢产物borrelidin，对包括禾谷镰孢菌(Fusarium graminearum)在内的多种植物病原菌具有拮抗作用。Cao等^[29]从受感染的棉花根系中分离出链霉菌DHV3-2，该菌株在棉花根系中具有强大的定殖能力，并对大丽轮枝菌及其他病原真菌展现出显著的抑制效果。薛磊等^[30]通过对6株链霉菌进行微菌核萌发和菌丝生长速率分析，发现这些链霉菌能有效抑制大丽轮枝菌的菌丝生长和孢子萌发。宋东博等^[31]鉴定出一株白浅灰链霉菌(Streptomyces albogriseolus) KF-43-1，该菌株对大丽轮枝菌V991的抑制率达到82.05%，显示出其作为生物防治菌剂的巨大潜力。

3.3 生防细菌

在生防细菌中，研究最多的是芽孢杆菌(Bacillus spp.)和假单胞菌(Pseudomonas spp.)。芽孢杆菌适应能力强，其产生的内生芽孢具有极强的抗胁迫能力，不仅能在多种恶劣环境下存活而且方便运输。芽孢杆菌抑菌谱广泛，能够抑制多种植物病原真菌和细菌，增加了生物防治的灵活性。大多数芽孢杆菌对人类和动物是安全的，不会引起病害或污染^[32]。因此，芽孢杆菌成为生物防治中研究最多的一类细菌(表1)。在芽孢杆菌中，报道最多的有枯草芽孢杆菌(Bacillus subtilis)、解淀粉芽孢杆菌(Bacillus amyloliquefaciens)以及贝莱斯芽孢杆菌(Bacillus velezensis)等。

Sherzad等^[38]发现解淀粉芽孢杆菌489-2-2会导致大丽轮枝菌VD080失去致病力，并通过激活棉花植株产生大量防御相关酶来增强植物的系统抗性，且菌株489-2-2能够定殖于棉花根系内。白红燕等^[43]筛选出2株对大丽轮枝菌的生长具有拮抗作用的枯草芽孢杆菌EBS03和EBS10，发现它们能有效促进温室棉苗的生长发育，降低黄萎病发病率和病情指数。吴梦君等^[33]研究发现枯草芽孢杆菌BS-Z15产生的活性物质可有效抑制大丽轮枝菌侵染棉花植株，降低棉花黄萎病发病率。张琼等^[35]在进行抑菌实验时观察到贝莱斯芽孢杆菌SZAD1及其发酵液提取物对大丽轮枝菌VD080的菌丝生长和孢子萌发具有显著的抑制作用，显示出其对棉花黄萎病潜在的生物防治作用；此外，菌株SZAD1还能产生和分泌多种纤维素酶和几丁质酶，这些酶有助于减少VD080在棉花茎部的定殖。白红燕等^[40]研究表明，贝莱斯芽孢杆菌EBV02可以抑制大丽轮枝菌菌丝生长，诱导棉花叶片活性氧暴发和胼胝质积累，温室实验中EBV02对棉花黄萎病的最高防治效果达到68.33%。Song等^[44]发现枯草芽孢杆菌KRS015可以显著降低大丽轮枝菌在棉花幼苗根部的定殖，使发病率降低62%，同时促进棉花的生长。除了抑制大丽轮枝菌生长外，芽孢杆菌还会抑制多种植物病原菌的生长。贾慧慧等^[45]发现解淀粉芽胞杆菌BJ-6对15种植物病原菌均有不同程度的抑菌活性，盆栽试验发现该菌株发酵液对甜瓜细菌性果斑病有良好的防治效果。此外，蜡样芽孢杆菌(Bacillus cereus)^[46]、短小芽孢杆菌(Bacillus pumilus)^[47]、解木聚糖类芽孢杆菌(Paenibacillus xylanilyticus)^[48]、甲基营养芽孢杆菌(Bacillus methylotrophicus)^[49]、阿萨尔基亚芽孢杆菌(Bacillus axarquiensis)^[50]等也被发现具有防治黄萎病等植物病害的潜在能力。

除芽孢杆菌外，假单胞菌作为生防菌也被广泛研究。戚家明等^[51]用拮抗实验证明假单胞菌PF-1对棉花黄萎病菌的生长具有良好抑制效果，抑菌率为71.30%。Ni等^[52]研究了不同浓度桔黄假单胞菌(Pseudomonas aurantiaca) ST-TJ4细胞悬液产生的挥发性有机物(volatile organic compounds, VOCs)对大丽轮枝菌菌丝径向生长和生物量的影响，发现其VOCs对大丽轮枝菌菌丝生长的抑制率最高可达63.1%。Niu等^[53]从厨余垃圾发酵残留物中分离出的铜绿假单胞菌ZL6可以有效抑制孢子萌发和菌丝生长，其发酵液处理与对照组相比，防治效果提高了47.72%。

4 生防微生物在防治棉花黄萎病中的作用机制

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4.1 空间与营养竞争

空间和营养是微生物生长不可或缺的2个基本条件。研究表明，一些生防微生物能够优先在植株的根系及体内定殖，因此能优先获取大量的氧气、水分及营养物质等，从而使得病原微生物无法获得足够的营养物质进行生长和繁殖，进而降低其感染几率(图2)。铁载体作为一种铁螯合剂，能够促进生防微生物菌株对游离铁离子的吸收和利用^[42]。例如，木霉HZA14可以产生铁载体，螯合环境中的游离铁元素，并占据一定的生态位，使得黄萎病菌无法获得足够的铁营养，从而实现抑制病原菌生长的目的^[54]。镰孢菌10R-7作为一种内生菌，能够定殖在棉花根内部，可能通过改变根际微环境或争夺根际空间和营养来直接抑制大丽轮枝菌的生长^[55]。此外，一些生防细菌还能在植株易受侵染部位形成生物膜，阻止病原菌的感染，从而保护植物免受病原菌的侵袭。Montes-Osuna等^[56]从橄榄的根际土壤中成功分离出一株假单胞菌PICF7，研究发现该菌株能够通过影响群体感应机制，促进生物膜的形成，进而有效控制黄萎病的发生。

4.2 拮抗作用

拮抗作用是指一种微生物通过产生活性代谢产物，有效抑制或杀死另一种微生物。这些活性代谢产物包括抗菌肽、脂肽、水解酶、大环内酯以及多烯类抗生素等物质。目前，脂肽类抗菌物质的研究较为广泛，主要包括芽孢杆菌产生的表面活性素(surfactin)、伊枯草菌素(iturin)、丰原素(fengycin)，以及链霉菌产生的达托霉素(daptomycin)等^[57]。例如，解淀粉芽孢杆菌Oj-2.16通过产生表面活性素、伊枯草菌素和丰原素，对黄萎病菌具有显著的拮抗作用^[58]。芽孢杆菌ABLF-18和ABLF-50可分泌表面活性素、伊枯草菌素、纤维素酶、葡聚糖酶等拮抗物质，将棉花的病害指数和真菌生物量降低到对照的40%-70%^[57]。Bubici等^[59]分离出的4种链霉菌(StB-3、StB-6、StB-11和StB-12)能够分泌抗生素、纤维素酶、溶菌酶等次级代谢产物，显著降低黄萎病症状，其中StB-11分离株的防治效果最好，防治效果达64.9%。镰孢菌10R-7可以产生镰孢菌酸，当浓度达到20 μg/mL时，其对大丽轮枝菌菌丝生长的抑制作用达到了100%^[55]。Liu等^[42]发现解淀粉芽孢杆菌YZU-SG146可以分泌纤维素酶、蛋白酶和淀粉酶等，能促进大丽轮枝菌菌丝的溶解，从而有效地抑制棉花黄萎病的发生。Zhang等^[41]筛选到了一株芽孢杆菌T6菌株，该菌株可产生挥发性有机物苯乙烯，诱导大丽轮枝菌一些水解酶等基因表达上调，促进大丽轮枝菌的水解，从而抑制其危害。Sun等^[6]从贝莱斯芽孢杆菌BvZ45-1的粗蛋白提取物中发现了大量的抗真菌物质，其中水杨酸脱羧酶可以抑制大丽轮枝菌产孢，并且可以使大丽轮枝菌菌丝破裂、孢子形态改变、细胞膜破裂和细胞死亡，从而抑制棉花黄萎病的发生。Zhang等^[60]发现枯草芽孢杆菌J15的代谢产物C17霉菌素，可导致大丽轮枝菌孢子收缩、下沉，甚至损伤；菌丝扭曲粗糙，表面凹陷，内容物分布不均匀，甚至使大丽轮枝菌死亡。

4.3 诱导植物系统抗性

诱导系统抗性(induced systemic resistance, ISR)，又称植物免疫或系统获得抗性，是指植物在遭遇外界刺激后迅速增强自身的免疫防御机制。这种机制通过提高植株一系列免疫防御酶的活性，如过氧化氢酶、过氧化物酶、超氧化物歧化酶、苯丙氨酸解氨酶和多酚氧化酶等，以及产生抗菌性植保素、酚类化合物和丙二醛等物质，构建起一种天然的保护屏障，以帮助植物抵御病原菌的侵袭。Shan等^[32]发现高原芽孢杆菌KRS010通过诱导水杨酸(salicylic acid, SA)和茉莉酸(jasmonic acid, JA)信号通路来激活植物系统抗性，从而引发植物免疫反应；同时，高原芽孢杆菌KRS010产生的胞外代谢物和挥发性化合物还抑制了大丽轮枝菌黑色素的生物合成。Sherzad等^[36]通过对贝莱斯芽孢杆菌SZAD2的过氧化氢染色发现，该菌株能够诱导植株在根和叶中积累过氧化氢，从而增强植物的基础防御反应；此外，该菌株还能诱导棉花根系的系统抗性，并显著提高棉花苯丙氨酸解氨酶、多酚氧化酶、过氧化物酶等抗氧化酶的活性和酚的含量。黑壳霉CEF08111则可以通过提高植株木葡聚糖代谢、细胞壁多糖代谢和半纤维素代谢等过程来提高植株自身抗性，从而降低黄萎病的发生^[17]。

4.4 促进植物生长

Ahmed等^[61]研究发现，某些生防微生物具备固氮和合成吲哚乙酸(indole-3-acetic acid, IAA)的能力，同时可以增加土壤中钾和磷酸盐的可溶性，这些能力有助于促进植物根系和植株的生长，从而间接提升植物对病害的抵抗力。Verma等^[62]发现用棉花内生真菌CFR-1和CEL-48处理的棉花种子活力提高，植株总糖含量上升，生长旺盛。Mohamad等^[63]报道放线菌菌株XIEG63和XIEG55可以分泌生长素、铁载体等，与未接种放线菌的植株相比，接种了XIEG63和XIEG55的棉花地上部分、根的长度和生物量显著增加。Liu等^[42]用解淀粉芽孢杆菌YZU-SG146悬浮液处理棉花幼苗后，苗长增加了4.18 mm，根长增加了22.87 mm，苗鲜重增加了0.72 g，均显著高于对照组。

5 生防微生物的应用方式

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生防微生物的应用方式多种多样，其中报道较多的包括对植物种子进行处理和制备复合生防菌剂等。针对不同种类的生防微生物，采用各自适宜的处理方式可以使其发挥出最大的防治效果。

5.1 对棉花种子的处理

对棉花种子的处理主要包括浸种、拌种和种子包衣等方式。浸种和拌种是将棉花种子与生防微生物悬浮液或生防菌剂通过浸泡或搅拌的方式，使种子表面附着生防微生物。种子包衣则是一种先进的种子处理技术，它可以将种子均匀地包裹在含有特定生防微生物的保护膜中，在种植过程中，这层保护膜中的生防微生物可以有效促进棉花种子的萌发，并减少病虫害的发生^[64]。娄善伟等^[65]利用枯草芽孢杆菌对棉花种子进行包衣处理，结果发现包衣处理可以使种子提前出苗，增加出苗率，并且使棉花的真叶数和果枝数增加；利用枯草芽孢杆菌对棉花种子进行包衣处理对棉花黄萎病的防治效果达到34.1%，同时可使棉花增产36.92%。

5.2 复合生防菌剂

在生物防治领域，利用单一微生物防治植物病害的报道较多。然而，单一微生物菌剂往往受限于其活性成分，导致其功能相对有限，并且对应用环境的要求较高。同时，单一生防微生物的防治机制较为单一，可能需要大量的菌体才能达到理想的防治效果。因此，将具备不同功能的微生物进行组合，开发出相较于单一微生物菌剂具有更稳定防治效果和更强促生作用的复合微生物菌剂，已经成为近年来生防微生物研究的重要发展方向；“中棉菌乐土”是由中国农业科学院棉花研究所开发的以棉花内生真菌为主的复合微生物菌剂，在温室和病圃试验中，该菌剂对棉花和茄子等作物的枯萎病和黄萎病展现出了显著的防治效果；施用“中棉菌乐土”后30 d和80 d，其对重度、中度和轻度黄萎病田的相对防治效率分别达到 34.7%-74.6%；同时，对于同样程度的病田，该菌剂还能使籽棉产量提高10.8%-13.6%^[66]。

6 展望

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生防微生物在棉花黄萎病的防治中展现出巨大的应用潜力。然而，它们如何通过产生抗菌物质、诱导植物产生防御反应以及竞争营养等方式发挥作用，仍需系统深入的研究。利用生防微生物防治棉花黄萎病也面临一些限制因素：许多生防微生物在与植物竞争营养时，可能会使植物将能量从繁殖过程转移，导致植物产量并不高于对照植株；生防微生物可能产生次级代谢产物，这些物质积累到一定程度时会对植物细胞产生毒害作用；此外，使用生防微生物有时也可能增加病害流行的风险。目前，理想的生防微生物相对稀缺，生防机制的许多方面仍需进一步研究和阐明。一些生防微生物制剂的研发进程缓慢，效果也有待提升。未来的研究将更加注重从自然界中筛选对棉花黄萎病具有高效防治作用的生防菌株，并通过基因工程技术对这些菌株进行定向遗传改造，以增强其适应性和拮抗活性。随着生防微生物菌株的不断筛选和创制，相信将有更多高效的生防菌剂被开发出来，这些菌剂将具有更好的稳定性和更广的应用范围。生防微生物的应用将不仅限于单一的生物防治策略，而是与研究抗病新品种、化学防治、物理防治、农业管理措施等相结合，形成综合防治策略，以实现对棉花黄萎病的长期有效管理。考虑到不同地区环境条件的差异，未来的研究将更加关注生防微生物在不同环境条件下的适应性和稳定性，以确保其在实际应用中的有效性。随着生物防治产品的发展，相关的法规和市场准入标准也将不断完善，为生防微生物菌剂的商业化和市场推广提供支持。在全球范围内，加强国际合作与交流，共享生防微生物研究的最新成果，将促进生防微生物技术的创新和应用^[67]。

总之，生防微生物在棉花黄萎病防治中的应用前景广阔。未来的研究和应用将更加注重技术创新、机制研究、产品开发和综合防治策略的构建，以实现棉花生产的可持续发展。

基金

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中国科学院战略性先导科技专项(XDA28030000)
山东省现代农业产业技术研究体系(SDAIT-03-04/06)
山东省良种工程(2023LZGC007)
山东省自然科学基金(ZR2023QC049)

参考文献

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2025年第65卷第3期

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doi: 10.13343/j.cnki.wsxb.20240682

接收时间：2024-11-04
首发时间：2026-02-10
出版时间：2025-03-04

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收稿日期：2024-11-04
录用日期：2024-12-03

基金

Strategic Priority Research Program of Chinese Academy of Sciences(XDA28030000)

中国科学院战略性先导科技专项(XDA28030000)

Modern Agro-industry Technology Research System of Shandong Province(SDAIT-03-04/06)

山东省现代农业产业技术研究体系(SDAIT-03-04/06)

Agricultural Seed Projects of Shandong Province(2023LZGC007)

山东省良种工程(2023LZGC007)

Natural Science Foundation of Shandong Province(ZR2023QC049)

山东省自然科学基金(ZR2023QC049)

作者信息

1 山东农业大学农学院，山东泰安

2 中国科学院微生物研究所真菌学国家重点实验室，北京

3 中国科学院大学生命科学学院，北京

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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Strain	Year	Biocontrol effect	Action mechanism	References
Bacillus subtilis BS-Z15	2019	BS-Z15 can effectively reduce the incidence of Verticillium wilt of cotton	Strain BS-Z15 secreted antagonistic active substances	[33]
Bacillus malacitensis Z-5	2019	Z-5 strain had remarkable control effect on Verticillium wilt of cotton	Z-5 strain can secrete lipopeptide antibiotics such as surfactant B and ictilicin A	[34]
Bacillus velezensis SZAD1	2020	The control efficiency of SZAD1 strain against Verticillium wilt was 60.10% and 56.00% in seed soaking and root irrigation, respectively	SZAD1 can produce cellulase and chitinase, which can reduce the ability of VD080 to settle cotton stems	[35]
Bacillus velezensis SZAD2	2020	The control efficiency of seed treatment was 60.31%, and that of root irrigation was 79.19%	The strain could systematically colonize the roots and induced systemic resistance of cotton roots by accumulating hydrogen peroxide in the roots and leaves	[36]
Bacillus circulans GN03	2021	Strain GN03 had good resistance to Verticillium wilt of cotton, with the highest control efficiency reaching 78%	GN03 inoculation altered the microflora in and around the plant roots, resulting in a significant accumulation of growth-related hormones	[37]
Bacillus amyloliquefaciens 489-2-2	2021	The control efficiency of seed treatment was 54.99%, and that of root irrigation was 60.31%	489-2-2 caused the mycelium of cotton verticillium wilt to lose pathogenicity, enhancing the systemic resistance of the plant by activating a large number of defense enzymes	[38]
Bacillus velezensis ND	2022	Application of ND fermentation liquid can increase the disease prevention effect from 36.00% to 92.99%	ND also has the activities of protease, cellulase and iron carrying, and has the ability to synthesize indole acetic acid, nitrogen fixation and phosphorus reduction	[39]
Bacillus velezensis EBV02	2022	The highest control effect of EBV02 on cotton Verticillium wilt was 68.33% and 37.25% in greenhouse and field tests, respectively	EBV02 inhibited the mycelia growth of Verticilliumdahliae, and induced active oxygen species outbreak and callus accumulation in cotton leaves	[40]
Bacillus T6	2023	The inhibition rate of T6 strain on Verticillium dahliae was 63.79%	T6 strain can produce volatile organic compound styrene, which can up-regulate the expression of some hydrolase genes in Charlottesia	[41]
Bacillus amyloliquefaciens YZU-SG 146	2023	The control effect of YZU-SG146 against Verticillium wilt of cotton was 84.21%, and it also promoted the growth of root length and seedling length of cotton seeds and seedlings	G146 can secrete ferric carrier, indoleacetic acid, cellulase, protease and amylase, and can trigger the outbreak of reactive oxygen species in cotton leaves	[42]
Bacillus velezensis BvZ45-1	2024	The indoor and field control efficiency of BvZ45-1 against Verticillium wilt of cotton was 46.53% and 47.27%, respectively	The bacterium can produce oxalate decarboxylase, inhibit the spore production of Verticilliumdahliae, and lead to mycelium rupture, cell membrane rupture and cell death	[6]
Bacillus altitudinis KRS010	2024	The effect of KRS010 strain on cotton Verticillium wilt was 93.59%	KRS010 induces plant immunity by inducing systemic resistance activated by salicylic acid and jasmonic acid signaling pathways	[32]

Strain

Year

Biocontrol effect

Action mechanism

References

Bacillus subtilis BS-Z15

2019

BS-Z15 can effectively reduce the incidence of Verticillium wilt of cotton

Strain BS-Z15 secreted antagonistic active substances

[33]

Bacillus malacitensis Z-5

2019

Z-5 strain had remarkable control effect on Verticillium wilt of cotton

Z-5 strain can secrete lipopeptide antibiotics such as surfactant B and ictilicin A

[34]

Bacillus velezensis SZAD1

2020

The control efficiency of SZAD1 strain against Verticillium wilt was 60.10% and 56.00% in seed soaking and root irrigation, respectively

SZAD1 can produce cellulase and chitinase, which can reduce the ability of VD080 to settle cotton stems

[35]

Bacillus velezensis SZAD2

2020

The control efficiency of seed treatment was 60.31%, and that of root irrigation was 79.19%

The strain could systematically colonize the roots and induced systemic resistance of cotton roots by accumulating hydrogen peroxide in the roots and leaves

[36]

Bacillus circulans GN03

2021

Strain GN03 had good resistance to Verticillium wilt of cotton, with the highest control efficiency reaching 78%

GN03 inoculation altered the microflora in and around the plant roots, resulting in a significant accumulation of growth-related hormones

[37]

Bacillus amyloliquefaciens 489-2-2

2021

The control efficiency of seed treatment was 54.99%, and that of root irrigation was 60.31%

489-2-2 caused the mycelium of cotton verticillium wilt to lose pathogenicity, enhancing the systemic resistance of the plant by activating a large number of defense enzymes

[38]

Bacillus velezensis ND

2022

Application of ND fermentation liquid can increase the disease prevention effect from 36.00% to 92.99%

ND also has the activities of protease, cellulase and iron carrying, and has the ability to synthesize indole acetic acid, nitrogen fixation and phosphorus reduction

[39]

Bacillus velezensis EBV02

2022

The highest control effect of EBV02 on cotton Verticillium wilt was 68.33% and 37.25% in greenhouse and field tests, respectively

EBV02 inhibited the mycelia growth of Verticilliumdahliae, and induced active oxygen species outbreak and callus accumulation in cotton leaves

[40]

Bacillus T6

2023

The inhibition rate of T6 strain on Verticillium dahliae was 63.79%

T6 strain can produce volatile organic compound styrene, which can up-regulate the expression of some hydrolase genes in Charlottesia

[41]

Bacillus amyloliquefaciens YZU-SG 146

2023

The control effect of YZU-SG146 against Verticillium wilt of cotton was 84.21%, and it also promoted the growth of root length and seedling length of cotton seeds and seedlings

G146 can secrete ferric carrier, indoleacetic acid, cellulase, protease and amylase, and can trigger the outbreak of reactive oxygen species in cotton leaves

[42]

Bacillus velezensis BvZ45-1

2024

The indoor and field control efficiency of BvZ45-1 against Verticillium wilt of cotton was 46.53% and 47.27%, respectively

The bacterium can produce oxalate decarboxylase, inhibit the spore production of Verticilliumdahliae, and lead to mycelium rupture, cell membrane rupture and cell death

[6]

Bacillus altitudinis KRS010

2024

The effect of KRS010 strain on cotton Verticillium wilt was 93.59%

KRS010 induces plant immunity by inducing systemic resistance activated by salicylic acid and jasmonic acid signaling pathways

[32]