贝莱斯芽孢杆菌应用进展:从农牧业到食品加工

谢燿孺1,上官玲玲1,严彩柠1,刘子雄2*,代俊1*

1(湖北工业大学 生命科学与健康工程学院,湖北 武汉,430068)2(安琪酵母股份有限公司,湖北 宜昌,443008)

摘 要 贝莱斯芽孢杆菌(Bacillus velezensis)作为厚壁菌门(Firmicutes)芽孢杆菌属(Bacillus)的一种革兰氏阳性菌,具有显著的促植物生长、抑制病原菌及广谱抑菌特性,在农作物生产中应用广泛。研究表明,该菌株在生物防治领域潜力显著,可通过减少作物病虫害、提升饲料利用率、提高产量等方式助力农业生产,同时对畜牧及水产养殖业的病害防控具有积极作用。此外,贝莱斯芽孢杆菌可作为益生菌制剂,促进肠道蠕动、增强人体免疫力,并适用于多种食品发酵工艺。该文系统综述贝莱斯芽孢杆菌的生物学特性及其在食品相关领域的应用现状与发展前景,旨在为该菌株应用范围的拓展、功能开发及潜在益生价值的挖掘提供理论参考。

关键词 贝莱斯芽孢杆菌;农作物生产;水产养殖;益生菌;发酵剂

贝莱斯芽孢杆菌(Bacillus velezensis)是一种新界定的芽孢杆菌属物种,其作为典型的生防菌和微生态制剂,已广泛应用于农业、畜牧业和水产养殖(图1),该菌不仅能促进农作物的产量提升,还能有效地帮助植物抵御微生物的感染,使农作物免受病害[1-4]。作为益生菌,它还能促进肠道蠕动、增强免疫力,其益生功能正不断被开发[5]

图1 贝莱斯芽孢杆菌在农业与食品领域的广泛应用
Fig.1 Wide-ranging applications of B.velezensis in agriculture and food industries

在“后抗生素时代”,耐药菌增加、药物残留和继发感染等问题日益严峻,公众对肉、蛋、乳等食品的绿色安全关注不断提升[1]。禽类种源、饲料、饮水和环境等养殖环节,以及寄生虫、应激等不良因素,均对畜禽生产性能造成影响。在作物生产中,耐药性细菌、真菌和病毒同样导致农业减产和粮食短缺[2]。此外,多种致病性耐药菌可侵袭人体消化系统,影响营养吸收,成为追求健康生活的重要障碍。因此,亟需一种兼顾农业增产、畜禽食品安全与人体消化功能提升的综合策略。

B.velezensis不仅能应用于农业和养殖领域,其生物学基础(如分泌的脂肽类物质、抗菌蛋白和酶系统)也为其在食品加工和人类健康领域的应用提供了桥梁。通过从农场到餐桌的全链条机制,该菌株可减少化学农药和抗生素的使用,确保食品原料的安全性,同时在发酵过程中提升营养价值和风味,形成农业生产与食品安全的无缝衔接[6-7]B.velezensis有望为农业减产防控、食品安全保障及人体消化功能改善提供解决方案。

本文系统综述了其在农牧业和食品领域的研究现状及多领域应用前景,为B.velezensis应用范围的扩大和功能开发提供参考,从而挖掘贝莱斯芽孢杆菌更多的食品相关领域应用潜力。

1 贝莱斯芽孢杆菌的概述

B.velezensis属于芽孢菌属,经过多年的命名、更名和重新认证,最终在细菌分类学中确立了其身份,并获得国际学者的广泛认可。其菌落呈乳白色、圆形,表面光滑略凸,直径为1.0~1.5 mm,质地黏稠。细胞呈杆状或大卵圆形,通常为对生排列,是革兰氏阳性细菌。尽管不形成内生孢子,但能产生孢囊和荚膜黏液,通过周围鞭毛进行运动,属于好氧微生物。

B.velezensis广泛分布于自然界,具有抗逆性、快速生长和易分离等特点,同时易培养、安全无害、无污染,能发酵丰富的产物,具备优良的抗菌和抗应激能力[8]。研究表明,B.velezensis能够产生次级代谢产物,抑制霉菌并分解霉菌毒素。如酶类(包括几丁质酶、脂肪酶、淀粉酶等)和抗菌物质(如抗菌蛋白、脂肽类抗生素以及合成抗生素类聚酮化合物)[9-13]。因此,B.velezensis具有作为饲料添加剂的潜力,能改善畜禽肠道微生态、调控脂肪代谢、增强免疫功能并促进饲料的消化吸收。此外,研究表明,B.velezensis作为益生菌有助于促进肠道蠕动、提升免疫力[5],同时可作为发酵剂,参与食醋、豆豉等调味品的发酵,增强食品风味[13]

2 贝莱斯芽孢杆菌在农牧业相关领域的应用

2.1 贝莱斯芽孢杆菌防治农作物疫病

农作物作为提供能量和膳食纤维的粮食作物,包括蔬菜和水果,与人类健康密切相关。然而,农作物在生产过程中常受多种病害侵扰,而B.velezensis的生物防治特性可有效应对这些微生物感染问题。其发酵产物,如胰岛素、泛素和表面活性剂,均表现出显著的抑菌作用,对病原菌具有抑制和拮抗效应[14-16]。基于此,众多研究者深入研究B.velezensis的生防特性,发现该菌对植物病原菌(表1)的生长繁殖具有关键抑制作用。研究表明,B.velezensis对核桃炭疽病[17]、茄子软腐病[18]、黄瓜叶斑病[19]、柑橘溃疡病[20]和番茄灰霉病[21]等多种病害均具有显著防治效果。此外,该菌对稻瘟病菌(Magnaporthe oryzae)以及疫霉菌属(Phytophthora spp.)、丝核菌属(Rhizoctonia spp.)、核盘菌属(Sclerotinia spp.)、青霉菌属(Penicillium spp.)和交链孢霉属(Alternaria spp.)等多种重要病原菌也表现出强烈的抑制活性。

表1 贝莱斯芽孢杆菌的来源及防治目标
Table 1 Origins and pathogen targets of B.velezensis

菌株来源防治对象参考文献B.velezensis核桃树下的土壤胡桃炭疽病[17]B.Velezensis M2022571中国普通微生物培养收藏中心 (CCTCC)茄子果实软腐病[18]B.velezensis ZF2健康黄瓜的茎叶斑病[19]B.velezensis 25柑橘柑橘类细菌溃疡病[20]B.velezensis AMR25山葡萄(Vitis amurensis Rupr.)的叶子番茄灰霉病[21]B.velezensis KOF112日本本土酿酒葡萄(Vitis sp. Cv)葡萄霜霉病和黄瓜霜霉病[22]B.velezensis CB13猪沼液花生霉腐病[23]B.velezensis HC-8金银花的叶片组织尖孢镰刀菌、烟草疫霉菌、轮纹镰刀菌和金银花白粉病[24]B.velezensis HG-8-2健康辣椒的根际土壤辣椒采后炭疽病[25]B.velezensis GA1草莓(Fragaria sp.)花生茎腐病[26]B.velezensis P2-1苹果树枝苹果采后腐烂症[27]B.velezensis WZ-37土壤番茄猝倒病和根腐病[28]B.velezensis FZB42ABiTEP有限责任公司油菜中的长孢轮枝菌( Verticillium longisporum)[29]

HAMAOKA等[22]研究表明,B.velezensis KOF12发酵液对葡萄霜霉病和黄瓜霜霉病具有显著的保护作用,接种量为1×108 CFU/mL时防治效果最佳,疾病严重程度分别降低100%和80%,且具有一定的治疗作用,呈现较持久的防控效果。JIA等[23]研究发现,B.velezensis CB13发酵液通过增强防御酶活性,显著提升植物对花生霉病菌的抗性,同时提高花生根际有益菌的物种多样性和丰度,有效抑制花生腐霉病病原菌的增殖。黎燕珊等[24]研究指出,经过发酵工艺优化后的B.velezensis HC-8可增强对尖孢镰刀菌(Fusarium oxysporum)、烟草疫霉菌(P.nicotiana)和轮纹镰刀菌(F.rotalis)的拮抗能力,同时提高对白粉病菌分生孢子萌发的抑制效果。ZHONG等[25]研究表明,B.velezensis有利于辣椒生长,对辣椒炭疽病具有良好防治效果,其他研究也发现,其对花生茎腐病[26]、苹果轮纹病[27]、番茄茎基腐病[28]和油菜黄萎病[29]等植物病害也展现出显著的防控效能。该菌株在植物病害防治研究领域已获得广泛关注,其防控能力得到了多方实证。

然而,目前B.velezensis作为生物防治剂的应用仍存在若干瓶颈:多数分离菌株的防控机理尚不明确,田间防治效果缺乏系统性验证,且规模化生产工艺和应用模式尚不成熟,这些问题亟待在后续研究和产业化应用中突破。特别需要指出的是,生物制剂的类型选择、施用方式和施药周期显著影响其防控效能,因此在B.velezensis制剂的开发与应用中,需要重点关注其生产工艺优化和田间施用技术体系的标准化建设。

2.2 贝莱斯芽孢杆菌防治畜禽和水产疫病

畜牧业和水产业及其制品(如鱼类、蛋类、乳类等)为人体提供优质蛋白质和能量物质,成为人类重要的食物来源。B.velezensis对畜禽育肥、鱼类生长性能提升和肠道菌群结构优化具有积极作用(表2)。近年来,关于B.velezensis在畜禽养殖和水产品的品质提升等领域的应用研究不断深入。

表2 贝莱斯芽孢杆菌畜牧与水产养殖中的应用效果
Table 2 Application effect of B.velezensis in livestock and aquaculture

菌株来源应用效果参考文献B.velezensis JT3-1牦牛粪便具有良好的抑菌作用并能增强宿主免疫功能[30]B.velezensis CE100—改善垫料质量,降低FPD发生率[31]B.velezensis仔猪粪便改善鸡的产蛋性能和猪的肠道菌群结构[32]B.velezensis CL197麦田显著降解ZEN[33]B.velezensis CAU277环境样本(柑橘)抑制空肠弯曲菌的生长,并能优化鸡的肠道菌群结构[34]B.velezensis CML532盲肠样本对家禽具有增强肠道健康和促进生长的综合作用[35]B.velezensis KNF-209武汉科诺生物科技有限公司可通过提升生长性能、增强消化能力及改善肠道代谢环境,促进畜禽健康与生产效率[44]B.velezensis Y6 and F7当地农场对青枯雷尔氏菌和尖孢镰刀菌具有强烈的拮抗活性[36]B.velezensis CGS1.1鸡粪具备较强的胃肠道存活能力、良好的黏附与生物膜形成特性,能广谱抑制病原菌及分解多种底物[37]B.velezensis ZBG17—提升肉鸡生产性能与免疫力,并具备安全高效的抗菌作用,可作为抗生素替代品[38]B.velezensis FIO1408深海海水具有抑制多种病原菌和分解蛋白质的能力,且对养殖鱼类安全[39]B.velezensis TPS3 N淡水鱼尼罗罗非鱼的消化道在水产养殖中具有良好的环境适应性和广谱抗菌活性[40]B.velezensis V4海水循环水养殖系统可改善鱼类生长性能和饲料利用效率,同时增强免疫因子表达[41]B.velezensis D-18废水样本在水产动物中可增强先天免疫、维持肠道菌群稳态,并广谱拮抗多种病原菌[42]

注:“—”表示无相关数据。

2.2.1 对畜禽疫病的防治

在甘肃地区家畜牦牛粪便中分离得到的B.velezensis JT3-1菌株[30],不仅能抑制金黄色葡萄球菌(Staphylococcus aureus)和产气荚膜梭菌(Clostridium perfringens),还可显著提升牦牛血清中免疫球蛋白A、免疫球蛋白G、免疫球蛋白M和干扰素-γ水平,展现出卓越的抗菌特性。研究证实,在肉鸡饲料中联合添加金属蛋白酶和B.velezensis CE100,可改善垫料质量并降低足垫皮炎(footpad dermatitis,FPD)发病率,该复合添加剂能有效提升肉鸡健康水平,为人类提供更安全的禽肉产品[31]。YE等[32]通过实验证明,B.velezensis饲喂蛋鸡可显著提高孕酮和胃动素水平,增加采食量并使平均产蛋率提升7.6%。该菌株添加至猪饲料后,研究显示猪粪便中瘤胃球菌属(Ruminococcus spp.)及毛螺菌属(Lachnospiraceae spp.)菌群数量增加,而致病性链球菌属(Streptococcus spp.)菌群数量下降[33]。另有研究表明,饲料中添加B.velezensis CL197可近乎完全消除玉米赤霉烯酮(zearalenone,ZEN),其代谢产物毒性低于ZEN,且在猪体外消化模型中ZEN降解率高达64%,成为食品生产链中理想的霉菌毒素解毒剂[33]。CUI等[34]发现,鸡饲料中添加B.velezensis可有效抑制空肠弯曲菌(Campylobacter jejuni)引发的禽类健康问题。当该菌与空肠弯曲菌体外共培养时,病原菌活菌数降至初始值的14%,同时显著提高鸡回肠微生物物种丰度(如另枝菌属Alistipes spp.、克里斯滕森菌科Christensenellaceaes),降低梭状芽孢杆菌属(Lachnoclostridium spp.)的菌群数量。从鸡的盲肠中分离出的B.velezensis CML532具有最强的胆盐和酸耐受性,以及高产酶和抗菌活性[35]。将B.velezensis CML532添加至饲粮中,可促进鸡生长,改善肠道屏障和吸收功能,调节肠道菌群。B.velezensis CML532在存在产气荚膜梭菌的条件下,可减轻鸡肠道损伤,减少产气荚膜梭菌的回肠定植,从而提高鸡的生长性能。

在畜牧养殖中,该菌可降低养殖成本和感染风险,减少霉菌毒素危害,增强抗病力与生产性能,并减少废弃物中的抗生素残留,从而保护生态环境[36-37]。该菌株具有广谱抑菌作用,可完全抑制大肠杆菌和沙门氏菌的生长,并能水解淀粉、纤维素等多种底物。此外,通过饮食补充B.velezensis ZBG17可提高肉鸡饲料利用效率和体液免疫响应[38]。作为抗生素替代品,由于该菌株可在6~8 h内完全抑制沙门氏菌和大肠杆菌,且基因组无安全隐患,因此其用于提升生长性能和健康水平。

2.2.2 对水产疫病的防治

HUANG等[39]及其团队从印度洋400 m深水样本中分离出B.velezensis FIO1408,该菌可抑制鳗利斯特菌(Listeria ivanovii)、副溶血弧菌(Vibrio parahaemolyticus)和溶藻弧菌(V.alginolyticus)等水产病原菌,并具有显著的蛋白质水解能力。实验结果表明,该菌在比目鱼养殖体系中的存活率与对照组无显著差异,表明其安全性良好。此外,KUEBUTORNYE等[40]从罗非鱼肠道分离出B.velezensis,该菌具有高细胞表面疏水性、自聚集性和耐高温特性,在低pH值和低胆汁酸浓度下仍保持活性,并对多数抗生素敏感。该菌对无乳链球菌(S.agalactiae)、嗜水气单胞菌(Aeromonas hydrophila)和哈维氏弧菌(V.harveyi)等病原菌表现出较强的抗菌活性。研究表明,大西洋鲑对A.hydrophila高度敏感,感染后死亡率可达93%;在饲料中添加B.velezensis可提高其生长速率、改善饲料转化率,并降低死亡率[41]。此外,该菌可促进鲈鱼体内白细胞介素-1β、肿瘤坏死因子-α和环氧合酶-2等免疫相关因子的表达。B.velezensis可通过激活宿主细胞丝裂原活化蛋白激酶(mitogen-activated protein kinase,MAPK)和核因子激活的B细胞的κ-轻链增强信号通路,增强先天免疫功能,调控肠道菌群稳态,从而提高机体免疫应答能力[42]。该菌对海豚链球菌(S.iniae)、鲶鱼爱德华氏菌(Edwardsiella ictaluri)、迟缓爱德华氏菌(E.tarda)和鰤鱼诺卡氏菌(Nocardia seriolae)等20余种水产病原菌均具有拮抗作用。在罗非鱼养殖中,该菌可维持鱼体健康,具有广阔的水产益生菌应用前景。基因组分析表明,B.velezensis缺乏毒力因子和毒素编码基因[37],对水生动物无致病风险[8]。其应用可增强免疫球蛋白M、主要组织相容性复合体等免疫蛋白基因的表达,诱导宿主溶菌酶等免疫酶合成,从而提高鱼体抗病力并降低疾病发生率[8,43]

B.velezensis可通过黏附定植在动物肠道内形成生物膜,并维持一定存活率,从而持续拮抗病原菌、增强宿主免疫功能并促进饲料营养吸收[44]。值得注意的是,B.velezensis还具有抗病毒作用,可诱导干扰素刺激基因表达上调,从而抑制病毒复制周期[38]

B.velezensis在农业与畜牧业中展现出优异的促生长与抑菌特性,这些功能为其在食品加工中的拓展应用奠定了生物学基础[6-7]。该菌所分泌的脂肽类物质(如表面活性素、伊枯草菌素等)不仅能抑制植物及动物病原微生物[45],还具备较强的抗氧化与表面活性作用[46-47],可有效调控食品发酵体系中微生态平衡、促进风味形成和提升整体安全性[47-48]。此外,其产酶系统(蛋白酶、淀粉酶、脂肪酶等)在饲料中提高养分吸收率的同时[27],也可用于食品原料的预处理和营养活化,实现从“饲料酶”到“发酵酶”的功能转化[49]

从农场到餐桌的食品链过程中,B.velezensis通过生态安全机制实现了多环节联通:在农作物环节减少化学农药使用(作为生防菌与植物根际促生菌,提高植保可持续性)[6-7];在养殖环节替代抗生素、改善肠道健康与生产性能[44],并具有已获监管安全评估的产品化先例[50];在食品加工环节提升发酵效率和食品功能性[46]。因而,该菌株被视为支撑“绿色农业-安全养殖-健康食品”一体化发展的关键微生物资源[7]

3 贝莱斯芽孢杆菌在作为人类食品相关领域的应用

基于上述农业和养殖领域的应用基础,B.velezensis在人类食品相关领域的潜力进一步凸显。其作为安全发酵剂和益生菌,不仅可桥接从原料生产到终端消费的链条,还能通过代谢活性提升食品的营养与安全属性。

3.1 贝莱斯芽孢杆菌发酵健康食品

“民以食为天,食以安为先”,安全、健康的食品是人类生存的根本保障。B.velezensis是一种安全、健康的食品发酵剂,已广泛应用于酱油、豆豉、食醋等传统发酵食品,以及益生菌制剂的生产中(表3)。其发酵特性和代谢活性不仅有助于食品风味的形成,还能提升食品的安全性。

表3 贝莱斯芽孢杆菌与健康食品的关联
Table 3 Role of B.velezensis in functional food processing

研究对象 (菌株来源)核心功能与健康食品的关联参考文献B.velezensis KMU01 (泡菜分离)具备益生功能基因与耐盐分解酶系,支持发酵食品营养价值[51]B.velezensis与米曲霉协同发酵强化蛋白分解与代谢物生成,提升产品营养品质[52]B.velezensis DP-2 (大豆分离)双效蛋白酶系分解抗营养因子,并抑菌保障安全[13]B.velezensis高产纤溶酶支持功能活性,营养成分转化效率需优化[53]B.velezensis PMC (传统食品分离)跨阶段PAHs生物降解:从生产端到体内连续阻断致癌物暴露[54]

HEO等[51]从韩国泡菜中分离出B.velezensis KMU01菌株。基因组分析表明,该菌株含有γ-谷氨酰转肽酶、钠激酶及细菌素合成的关键基因,这些基因与人体健康密切相关。该菌株还携带淀粉酶、蛋白酶、脂肪酶等降解酶编码基因,以及耐盐相关基因簇,表明其适合作为高盐环境下食品发酵的优选菌种。GIL等[52]发现,B.velezensis与米曲霉(Aspergillus oryzae)协同发酵大豆酱时,氨基酸态氮含量比单一米曲霉发酵提高38.7%,并显著增加肌醇、葡萄糖、甘油和不饱和脂肪酸等有益代谢物。LIU等[13]从大豆中分离出B.velezensis DP-2菌株,该菌株可分泌5.85 U/mL的碱性蛋白酶和5.99 U/mL的中性蛋白酶,对霉菌、酵母菌及沙门氏菌的抑制率超过92%。在豆粕发酵中,该菌株使大豆球蛋白和β-伴球蛋白的分解率分别达到78.00%和43.07%,三氯乙酸可溶性蛋白含量提高至34.6 mg/g,粗蛋白利用率达到81.3%,证明其在豆类发酵中的优势。李杨等[53]发现,在35 ℃、接种量0.1%、发酵5 d的条件下,将B.velezensis接种于豆豉发酵体系,可使纤溶酶活性达到398.7 U/g。但氨基酸态氮和还原糖含量仍较低,提示需进一步优化接种时机、初始pH值及与纳豆芽孢杆菌(B.natto)的共培养周期。此外,从传统发酵食品中分离的B.velezensis PMC对苯并[a]芘等多环芳烃(polycyclic aromatic hydrocarbons,PAHs)具有显著降解能力。即使在菌体失活状态下,其胞外酶仍能降解83.6%的苯并[a]芘(50 mg/L),为食品污染物的生物消减提供了新思路。研究表明,B.velezensis PMC可通过双重解毒机制提升食品安全性:在食品发酵阶段有效降解PAHs污染物(降解率82.4%),并在人体肠道内持续发挥PAHs生物消减作用(体外模拟肠液环境下降解效率67.9%),从而形成覆盖食品加工到人体代谢的全链条健康保障体系[54]

3.2 贝莱斯芽孢杆菌作为人类肠胃益生菌

消化系统是人体重要且脆弱的系统之一,由口腔、咽喉、食管和胃肠等组成,负责分解食物并吸收营养。多种致病菌可经口腔进入并定植于胃肠道,破坏黏膜屏障,引发胃肠疾病,危害健康并削弱营养吸收。B.velezensis可从人体肠道分离,具促进蠕动和增强营养吸收等功能,已被列为重要候选益生菌[5],并被欧洲食品安全局(European Food Safety Authority,EFSA)纳入安全资格推定(Qualified Presumption of Safety,QPS)名录[5],显示其益生潜力获权威认可。

BRUTSCHER等[5]发现,B.velezensis的独特菌株BV379具有益生菌应用潜力,其基因组包含调控杆菌肽、梅加霉素和植物唑啉等抗菌物质合成的基因簇。该菌可响应食物相关刺激及胃肠道微环境中的应激信号,广泛利用多种碳水化合物,并表现出较强的蛋白酶和脂肪酶活性,从而促进日常膳食中蛋白质和膳食纤维的消化吸收。CAI等[55]研究表明,B.velezensis A2可通过调控猪肠上皮细胞的Wnt配体/卷曲相关蛋白/β-连环蛋白信号通路,影响细胞周期进程,缓解活性氧应激,减少ZEN诱导的细胞损伤,为益生菌维护肠道屏障功能提供分子机制参考。SAHAL等[56]从子宫颈分离的B.velezensis对低pH、高盐和0.3%胆酸盐均表现出良好耐受性,这为其作为肠道适应性益生菌提供了理论依据。

4 应用前景的展望

B.velezensis为兼性好氧芽孢菌,能在根际/肠道快速定殖并形成生物被膜;其基因组富含生物合成簇,可高产表面活性素、伊枯草菌素、丰原素等脂肽与多烯类活性物,抑菌促生并可干扰群体感应、破坏生物被膜。芽孢赋予其耐热、耐酸、耐贮运与制剂稳定性,生物安全性高且耐药选择压力低。基于这些特性,它成为替代抗生素与化学药剂的理想底座,进而支撑其在农业、渔业与食品工业中的多场景应用前景。

植物类经济作物产值高且与民生密切相关,但病虫害危害严重。现有植保主要依赖化学药剂,易致耐药性并通过食物链危害健康[57]。微生物拮抗菌剂具有工艺简便、毒性低、无残留等优势[58]。贝莱斯芽孢杆菌耐药突变概率低于1×10-9,可合成表面活性素、伊枯草菌素等代谢物,占据生态位、竞争营养抑制病原,并分泌植物生长素促进作物生长。在畜禽养殖中,可减少感染损失,降低霉菌毒素危害,使抗病力提升2.3倍、抗生素残留降92%,实现经济与生态双赢[59]

在生态压力叠加与农业结构调整背景下,水产养殖业已成为现代农业的重要支柱[60]。面对疫病、极端气候和地质异动等风险,亟需研发可替代抗生素的新型制剂,以在“禁抗令”与养殖需求之间建立技术缓冲,满足规模化养殖提质增效的要求[61]。当前防治无乳链球菌、海豚链球菌、鲫诺卡氏菌和嗜水气单胞菌等仍依赖抗生素和消毒剂,虽见效快但易致耐药并污染环境[61]。贝莱斯芽孢杆菌可高效抑制上述病原,对罗非鱼、斑马鱼等安全,其分泌的脂肽类物质可穿透生物被膜并干扰群体感应,阻断致病通路。

食品工业关系国民健康。2022年我国食源性疾病发病率为64.3/10万,凸显构建“最严谨标准、最严格监管”食品安全体系(GB 2760—2024《食品安全国家标准 食品添加剂使用标准》)的必要性[62]。贝莱斯芽孢杆菌在传统发酵食品和功能食品中应用广泛,可抑制霉菌、酵母菌和沙门氏菌,并分解大分子营养物质,使蛋白质消化率达91.3%[59]。作为潜力益生菌,其耐受胃酸和肠道高盐、高胆酸环境,促进肠蠕动和紧密连接蛋白表达,使钙、铁、锌吸收率分别达 89%、76%、81%,为功能食品开发提供解决方案[58]

贝莱斯芽孢杆菌兼具高抗逆性、广谱抑菌性与优良生物安全性,能够在农业、渔业及食品工业中实现多维度应用。其低耐药风险与环境友好特性,使其成为替代抗生素和化学药剂的理想微生物资源[61]。未来,结合基因组改良与产业化技术推广,该菌株有望在全球绿色可持续发展中发挥重要作用。

5 结论

B.velezensis作为新型益生菌,兼具防治动植物病害、提高饲料消化率、减少养殖损耗和抑制霉菌生长等多种功能。该菌可分泌纤维素酶、半纤维素酶和β-葡聚糖酶等多种酶,促进植物性原料和饲料的消化吸收。此外,在食品发酵领域,B.velezensis可激活谷物中γ-氨基丁酸合成途径,促进多酚类物质的生物转化,显著提升食品营养价值;其菌体成分还能促进肠道蠕动、增强消化功能、提高营养吸收效率,其代谢产物中的脂肽类物质(如表面活性素、伊枯草菌素)可通过多靶点机制发挥益生作用。在农业和食品工业中具有广阔的应用前景。

B.velezensis是一种多功能微生物资源,在农业、畜牧业、水产养殖、食品加工及人类健康领域展现出显著潜力。本文综述了其生物学特性,包括广泛的抗菌能力和促进作物产量提升的能力,并系统阐述了其在防治作物病害、畜禽水产疫病、发酵健康食品及作为肠道益生菌的应用进展。该菌株通过分泌脂肽、抗菌蛋白和酶类物质,不仅可替代化学农药和抗生素,减少环境污染和耐药风险,还能提升食品营养价值和人体消化功能。未来可通过多组学分析和基因工程优化,进一步拓展其应用范围,为绿色农业、食品安全及可持续发展的微生物解决方案提供重要支撑。

参考文献

[1] KONG W J, YAN Y C, LI X Y, et al.Draft genome sequence of Bacillus velezensis PEBA20, a strain with a plant growth-promoting effect and biocontrol potential[J].Genome Announcements, 2018, 6(21):e00286-e00218.

[2] RABBEE M F, ALI M S, CHOI J, et al.Bacillus velezensis:A valuable member of bioactive molecules within plant microbiomes[J].Molecules, 2019, 24(6):1046.

[3] KHALID F, KHALID A, FU Y C, et al.Potential of Bacillus velezensis as a probiotic in animal feed:A review[J].Journal of Microbiology, 2021, 59(7):627-633.

[4] SU T, SHEN B, HU X J, et al.Research advance of Bacillus velezensis:Bioinformatics, characteristics, and applications[J].Food Science and Human Wellness, 2024, 13(4):1756-1766.

[5] BRUTSCHER L M, GEBRECHRISTOS S, GARVEY S M, et al.Genetic and phenotypic characterization of Bacillus velezensis strain BV379 for human probiotic applications[J].Microorganisms, 2024, 12(3):436.

[6] KENFAOUI J, DUTILLOY E, BENCHLIH S, et al.Bacillus velezensis:A versatile ally in the battle against phytopathogens:Insights and prospects[J].Applied Microbiology and Biotechnology, 2024, 108:439.

[7] KESHMIRSHEKAN A, DE SOUZA MESQUITA L M, VENTURA S P M.Biocontrol manufacturing and agricultural applications of Bacillus velezensis[J].Trends in Biotechnology, 2024, 42(8):986-1001.

[8] LI J, WU Z B, ZHANG Z, et al.Effects of potential probiotic Bacillus velezensis K2 on growth, immunity and resistance to Vibrio harveyi infection of hybrid grouper (Epinephelus lanceolatus♂ × E.fuscoguttatus♀)[J].Fish &Shellfish Immunology, 2019, 93:1047-1055.

[9] LI C, LI S Z, DANG G Q, et al.Screening and characterization of Bacillus velezensis LB-Y-1 toward selection as a potential probiotic for poultry with multi-enzyme production property[J].Frontiers in Microbiology, 2023, 14:1143265.

[10] CHEN L, QU Z H, YU W, et al.Comparative genomic and transcriptome analysis of Bacillus velezensis CL-4 fermented corn germ meal[J].AMB Express, 2023, 13:10.

[11] JIN Q, JIANG Q Y, ZHAO L, et al.Complete genome sequence of Bacillus velezensis S3-1, a potential biological pesticide with plant pathogen inhibiting and plant promoting capabilities[J].Journal of Biotechnology, 2017, 259:199-203.

[12] XIONG Z R, COBO M, WHITTAL R M, et al.Purification and characterization of antifungal lipopeptide produced by Bacillus velezensis isolated from raw honey[J].PLoS One, 2022, 17(4):e0266470.

[13] LIU Z Y, GUAN X F, ZHONG X X, et al.Bacillus velezensis DP-2 isolated from Douchi and its application in soybean meal fermentation[J].Journal of the Science of Food and Agriculture, 2021, 101(5):1861-1868.

[14] LIU X Y, REN B, CHEN M, et al.Production and characterization of a group of bioemulsifiers from the marine Bacillus velezensis strain H3[J].Applied Microbiology and Biotechnology, 2010, 87(5):1881-1893.

[15] KIM S Y, LEE S Y, WEON H Y, et al.Complete genome sequence of Bacillus velezensis M75, a biocontrol agent against fungal plant pathogens, isolated from cotton waste[J].Journal of Biotechnology, 2017, 241:112-115.

[16] YU F T, SHEN Y Y, PANG Y Y, et al.Effects of branched-chain amino acids on surfactin structure and antibacterial activity in Bacillus velezensis YA215[J].World Journal of Microbiology and Biotechnology, 2024, 40(9):281.

[17] WANG L M, ZHU T H.Strong opponent of walnut anthracnose:Bacillus velezensis and its transcriptome analysis[J].Microorganisms, 2023, 11(8):1885.

[18] ZHANG X Y, XIN Y, WANG J Y, et al.Characterization of a Bacillus velezensis strain as a potential biocontrol agent against soft rot of eggplant fruits[J].International Journal of Food Microbiology, 2024, 410:110480.

[19] XU S, XIE X W, ZHAO Y R, et al.Whole-genome analysis of Bacillus velezensis ZF2, a biocontrol agent that protects Cucumis sativus against Corynespora leaf spot diseases[J].3 Biotech, 2020, 10(4):186.

[20] RABBEE M F, BAEK K H.Detection of antagonistic compounds synthesized by Bacillus velezensis against Xanthomonas citri subsp.citri by metabolome and RNA sequencing[J].Microorganisms, 2023, 11(6):1523.

[21] ANANEV A A, OGNEVA Z V, NITYAGOVSKY N N, et al.Whole genome sequencing of Bacillus velezensis AMR25, an effective antagonist strain against plant pathogens[J].Microorganisms, 2024, 12(8):1533.

[22] HAMAOKA K, AOKI Y, SUZUKI S.Isolation and characterization of endophyte Bacillus velezensis KOF112 from grapevine shoot xylem as biological control agent for fungal diseases[J].Plants, 2021, 10(9):1815.

[23] JIA S, SONG C, DONG H, et al.Evaluation of efficacy and mechanism of Bacillus velezensis CB13 for controlling peanut stem rot caused by Sclerotium rolfsii[J].Frontiers in Microbiology, 2023, 14:1111965.

[24] 黎燕珊, 崔文艳, 张陈芳, 等.抗金银花白粉病菌贝莱斯芽孢杆菌HC-8菌株培养基及发酵条件优化[J].南方农业学报, 2021, 52(8):2148-2157.
LI Y S, CUI W Y, ZHANG C F, et al.Optimization of culture medium and fermentation parameters of Bacillus velezensis HC-8 antagonistic to Erysiphe lonicerae[J].Journal of Southern Agriculture, 2021, 52(8):2148-2157.

[25] ZHONG J, WU X, GUO R, et al.Biocontrol potential of Bacillus velezensis HG-8-2 against postharvest anthracnose on chili pepper caused by Colletotrichum scovillei[J].Food Microbiology, 2024, 124:104613.

[26] KORANGI ALLELUYA V, ARGÜELLES ARIAS A, RIBEIRO B, et al.Bacillus lipopeptide-mediated biocontrol of peanut stem rot caused by Athelia rolfsii[J].Frontiers in Plant Science, 2023, 14:1069971.

[27] YUAN H B, SHI B K, WANG L, et al.Isolation and characterization of Bacillus velezensis strain P2-1 for biocontrol of apple postharvest decay caused by Botryosphaeria dothidea[J].Frontiers in Microbiology, 2022, 12:808938.

[28] ZHANG Y, LI Y Y, LIANG S B, et al.Study on the preparation and effect of tomato seedling disease biocontrol compound seed-coating agent[J].Life, 2022, 12(6):849.

[29] HAFIZ F B, MORADTALAB N, GOERTZ S, et al.Synergistic effects of a root-endophytic Trichoderma fungus and Bacillus on early root colonization and defense activation against Verticillium longisporum in rapeseed[J].Molecular Plant-Microbe Interactions, 2022, 35(5):380-392.

[30] LI Y Q, LI X, JIA D, et al.Complete genome sequence and antimicrobial activity of Bacillus velezensis JT3-1, a microbial germicide isolated from yak feces[J].3 Biotech, 2020, 10(5):231.

[31] PARK C J, SUN S S.Effect of dietary metallo-protease and Bacillus velezensis CE 100 supplementations on growth performance, footpad dermatitis and manure odor in broiler chickens[J].Animal Bioscience, 2022, 35(10):1628-1634.

[32] YE M, WEI C J, KHALID A, et al.Effect of Bacillus velezensis to substitute in-feed antibiotics on the production, blood biochemistry and egg quality indices of laying hens[J].BMC Veterinary Research, 2020, 16:400.

[33] ORSO P B, EVANGELISTA A G, DE MELO NAZARETH T, et al.Bacillus velezensis CL197:A zearalenone detoxifying strain isolated from wheat with potential to be used in animal production[J].Veterinary Research Communications, 2024, 48(6):3847-3857.

[34] CUI Y F, ZHU J J, LI P X, et al.Assessment of probiotic Bacillus velezensis supplementation to reduce Campylobacter jejuni colonization in chickens[J].Poultry Science, 2024, 103(8):103897.

[35] LA TENG ZHU LA A, WEN Q, XIAO Y X, et al.A new Bacillus velezensis strain CML532 improves chicken growth performance and reduces intestinal Clostridium perfringens colonization[J].Microorganisms, 2024, 12(4):771.

[36] CAO Y, PI H L, CHANDRANGSU P, et al.Antagonism of two plant-growth promoting Bacillus velezensis isolates against Ralstonia solanacearum and Fusarium oxysporum[J].Scientific Reports, 2018, 8:4360.

[37] SONI R, KEHARIA H, DUNLAP C, et al.Functional annotation unravels probiotic properties of a poultry isolate, Bacillus velezensis CGS1.1[J].LWT, 2022, 153:112471.

[38] SONI R, KEHARIA H, BOSE A, et al.Genome assisted probiotic characterization and application of Bacillus velezensis ZBG17 as an alternative to antibiotic growth promoters in broiler chickens[J].Genomics, 2021, 113(6):4061-4074.

[39] HUANG W H, QU L Y, GAO P, et al.Bioassay and whole-genome analysis of Bacillus velezensis FIO1408, a biocontrol agent against pathogenic bacteria in aquaculture[J].Current Microbiology, 2023, 80(11):354.

[40] KUEBUTORNYE F K A, LU Y S, ABARIKE E D, et al.In vitro assessment of the probiotic characteristics of three Bacillus species from the gut of Nile tilapia, Oreochromis niloticus[J].Probiotics and Antimicrobial Proteins, 2020, 12(2):412-424.

[41] WANG C, LIU Y, SUN G X, et al.Growth, immune response, antioxidant capability, and disease resistance of juvenile Atlantic salmon (Salmo salar L.) fed Bacillus velezensis V4 and Rhodotorula mucilaginosa compound[J].Aquaculture, 2019, 500:65-74.

[42] MONZN-ATIENZA L, BRAVO J, FERNNDEZ-MONTERO , et al.Dietary supplementation of Bacillus velezensis improves Vibrio anguillarum clearance in European sea bass by activating essential innate immune mechanisms[J].Fish &Shellfish Immunology, 2022, 124:244-253.

[43] REYES-ESTEBANEZ M, SANMARTN P, CAMACHO-CHAB J C, et al.Characterization of a native Bacillus velezensis-like strain for the potential biocontrol of tropical fruit pathogens[J].Biological Control, 2020, 141:104127.

[44] LIU Y, XIONG M Q, HU X, et al.Dietary Bacillus velezensis KNF-209 supplementation improves growth performance, enhances immunity, and promotes gut health in broilers[J].Poultry Science, 2024, 103(9):103946.

[45] DE FTIMA DIAS DINIZ G, FIGUEIREDO J E F, CANUTO K M, et al.Chemical and genetic characterization of lipopeptides from Bacillus velezensis and Paenibacillus ottowii with activity against Fusarium verticillioides[J].Frontiers in Microbiology, 2024, 15:1443327.

[46] ROY A, KHAN M R, MUKHERJEE A K.Recent advances in the application of microbial biosurfactants in food industries:Opportunities and challenges[J].Food Control, 2024, 163:110465.

[47] DUSSERT E, TOURRET M, DUPUIS C, et al.Evaluation of antiradical and antioxidant activities of lipopeptides produced by Bacillus subtilis strains[J].Frontiers in Microbiology, 2022, 13:914713.

[48] LUO Y Y, GUO Y, HU X Y, et al.Flavor improvement of fermented soybean foods by co-fermentation with Bacillus velezensis and Lactiplantibacillus plantarum[J].LWT, 2023, 186:115257.

[49] XUE J, WU J Y, JI Y L, et al.Effect of microbial fermentation on the quality of soybean meal[J].International Journal of Food Science &Technology, 2024, 59(1):72-83.

[50] EFSA Panel on Additives and Products or Substances used in Animal Feed (FEEDAP), BAMPIDIS V, AZIMONTI G, et al.Safety and efficacy of a feed additive consisting of Bacillus velezensis DSM 15544 (calsporin®) for dairy cows and other dairy ruminants (asahi biocycle Co.Ltd.)[J].EFSA Journal, 2022, 20(1):e06984.

[51] HEO S, KIM J H, KWAK M S, et al.Functional annotation genome unravels potential probiotic Bacillus velezensis strain KMU01 from traditional Korean fermented kimchi[J].Foods, 2021, 10(3):563.

[52] GIL N Y, JANG Y J, GWON H M, et al.Comparative evaluation of quality and metabolite profiles in meju using starter cultures of Bacillus velezensis and Aspergillus oryzae[J].Foods, 2022, 11(1):68.

[53] 李杨, 陈宇航, 刘雪薇, 等.响应面法优化贝莱斯芽孢杆菌发酵水豆豉的工艺条件[J].中国调味品, 2020, 45(2):49-53.
LI Y, CHEN Y H, LIU X W, et al.Optimization of technological conditions for fermented soya beans by Bacillus velezensis using response surface methodology[J].China Condiment, 2020, 45(2):49-53.

[54] SULTANA O F, LEE S, SEO H, et al.Biodegradation and removal of PAHs by Bacillus velezensis isolated from fermented food[J].Journal of Microbiology and Biotechnology, 2021, 31(7):999-1010.

[55] CAI J, YUAN X S, SUN Y H, et al.Bacillus velezensis A2 can protect against damage to IPEC-J2 cells induced by zearalenone via the Wnt/FRZB/β-catenin signaling pathway[J].Toxins, 2024, 16(1):44.

[56] SAHAL G, DONMEZ H G, BEKSAC M S.Cervicovaginal Bacillus velezensis isolate:A potential probiotic and an antagonist against Candida and Staphylococcus[J].Current Microbiology, 2023, 80(10):332.

[57] PRETTY J, BENTON T G, BHARUCHA Z P, et al.Global assessment of agricultural system redesign for sustainable intensification[J].Nature Sustainability, 2018, 1(8):441-446.

[58] OLANREWAJU O S, AYANGBENRO A S, GLICK B R, et al.Plant health:Feedback effect of root exudates-rhizobiome interactions[J].Applied Microbiology and Biotechnology, 2019, 103(3):1155-1166.

[59] ELSHAGHABEE F M F, ROKANA N, GULHANE R D, et al.Bacillus as potential probiotics:Status, concerns, and future perspectives[J].Frontiers in Microbiology, 2017, 8:1490.

[60] The state of world fisheries and aquaculture:2022:towards blue transformation[Z].Rome:Food and Agriculture Organization of the United Nations, 2022.

[61] Antimicrobial Resistance Collaborators.Global burden of bacterial antimicrobial resistance in 2019:A systematic analysis[J].Lancet (London, England), 2022, 399(10325):629-655.

[62] WANG T Y, WU Y B, LI W W, et al.Diarrheagenic Escherichia coli outbreak reporting to foodborne disease outbreaks surveillance system:China, 2011-2022[J].China CDC Weekly, 2024, 6(51):1370-1374.

Advances in the application of Bacillus velezensis:From agriculture and animal husbandry to food processing

XIE Yaoru1, SHANGGUAN Lingling1, YAN Caining1, LIU Zixiong2*, DAI Jun1*

1(Hubei Key Laboratory of Industrial Microbiology, School of Life and Health Sciences, Hubei University of Technology, Wuhan 430068, China)2(Angel Yeast Co.Ltd., Yichang 443008, China)

ABSTRACT Bacillus velezensis, a Gram-positive bacterium of the genus Bacillus belonging to the Firmicutes phylum, is notable for its significant plant growth-promoting, pathogen-inhibiting, and broad-spectrum antibacterial properties.This paper reviews recent studies on the biological characteristics of B.velezensis and synthesizes its application progress across multiple fields.We begin by discussing its role in disease prevention for crops, livestock, and aquaculture, then summarize its advancements in food processing and as an intestinal probiotic.The review also highlights the potential of B.velezensis as an alternative to chemical pesticides and antibiotics.By integrating existing research findings, this work aims to provide theoretical insights for expanding the application scope, developing novel functionalities, and exploring the untapped probiotic value of this promising strain.

Key words Bacillus velezensis;crop production;aquaculture;probiotics;starter culture

DOI:10.13995/j.cnki.11-1802/ts.044865

引用格式:谢燿孺,上官玲玲,严彩柠,等.贝莱斯芽孢杆菌应用进展:从农牧业到食品加工[J].食品与发酵工业,2026,52(15):401-408.XIE Yaoru, SHANGGUAN Lingling, YAN Caining, et al.Advances in the application of Bacillus velezensis:From agriculture and animal husbandry to food processing[J].Food and Fermentation Industries,2026,52(15):401-408.

第一作者:硕士研究生(代俊副教授和刘子雄助理工程师为共同通信作者,E-mail:jundai@hbut.edu.cn;liuzixiong@hbut.edu.cn)

基金项目:国家自然科学基金项目(31871789,41876114);湖北省自然科学基金项目(2024AFB803)

收稿日期:2025-10-18,改回日期:2025-12-16