综述

芽孢杆菌在饲料霉菌毒素脱毒中的应用及机制研究进展

  • 段滔 ,
  • 王若菲 ,
  • 丁励 ,
  • 周盼 , *
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  • 西南科技大学生命科学与农林学院, 绵阳 621010
*周 盼,讲师,硕士生导师,E-mail:

段 滔(2001—),男,四川遂宁人,硕士研究生,研究方向为动物营养与饲料加工。E-mail:

Office editor: 靳爽

收稿日期: 2025-09-16

  网络出版日期: 2026-04-14

基金资助

国家重点研发计划项目(2023YFD1300802)

四川省科技厅项目(2022NSFSC1778)

Research Progress on Application and Mechanism of Bacillus in Detoxification of Mycotoxins in Feed

  • DUAN Tao ,
  • WANG Ruofei ,
  • DING Li ,
  • ZHOU Pan , *
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  • College of Life Science and Agri-Forestry, Southwest University of Science and Technology, Mianyang 621010, China
*lecturer, E-mail:

Received date: 2025-09-16

  Online published: 2026-04-14

摘要

霉菌毒素是由镰刀菌属、曲霉菌属等真菌产生的有毒次级代谢产物,广泛污染饲料原料,严重威胁动物健康与畜产品安全。传统物理和化学脱毒方法(如添加吸附剂、物理加工、化学反应等)虽可在一定程度上破坏毒素分子结构,但普遍存在脱毒效率低、营养成分损失及潜在残留风险等问题。近年来,生物脱毒技术因其高效性、安全性与环境友好性等优势成为研究热点。其中,芽孢杆菌凭借其分布广泛、抗逆性强及代谢多样等特性,在霉菌毒素降解领域展现出广泛应用潜力。尽管已有大量研究总结了霉菌毒素的生物降解方法,但针对芽孢杆菌在饲料霉菌毒素脱毒中的具体应用及其作用机制的系统梳理仍相对有限。本文系统综述了饲料中常见霉菌毒素(如黄曲霉毒素B1、玉米赤霉烯酮、脱氧雪腐镰刀菌烯醇、赭曲霉毒素A和伏马毒素)的毒性特征及现有脱毒方法的优缺点,重点总结了芽孢杆菌对多种霉菌毒素的降解效率与作用机制,以期为高效脱霉菌株的选育、芽孢杆菌类脱霉剂的开发及其在饲料中的安全应用提供参考。

本文引用格式

段滔 , 王若菲 , 丁励 , 周盼 . 芽孢杆菌在饲料霉菌毒素脱毒中的应用及机制研究进展[J]. 动物营养学报, 2026 , 38(4) : 2520 -2533 . DOI: 10.12418/CJAN2026.203

Abstract

Mycotoxins are toxic secondary metabolites produced by fungi such as Fusarium spp. and Aspergillus spp. They widely contaminate feed materials, posing serious threats to animal health and livestock product safety. Traditional physical and chemical detoxification methods(e.g., adsorbents, physical processing, chemical reactions) can partially disrupt toxin molecular structures but are limited by low efficiency, nutrient loss and potential residue risks. In recent years, biological detoxification has emerged as a research hotspot due to its advantages of high efficiency, safety and environmental friendliness. Among microbial agents, Bacillus show broad potential in mycotoxin degradation owing to their wide distribution, high stress tolerance and diverse metabolic capabilities. Although many studies have summarized the biodegradation methods for mycotoxins, systematic reviews focusing specifically on the application and mechanisms of Bacillus in the detoxification of mycotoxins in feed remain relatively limited. This paper systematically reviews the toxicological characteristics of common mycotoxins found in feed (e.g., aflatoxin B1, zearalenone, deoxynivalenol, ochratoxin A and fumonisin) and the advantages and disadvantages of existing detoxification methods. It focuses on summarizing the degradation efficiency and mechanisms of action of Bacillus against various mycotoxins, aiming to provide a reference for the screening of highly efficient detoxifying strains, the development of Bacillus-based detoxifying agents, and their safe application in feed.

霉菌毒素是镰刀菌属、曲霉菌属等真菌在生长繁殖过程中产生的一类有毒次级代谢产物[1-2]。霉菌毒素污染在饲料原料及配合饲料中普遍存在,呈现出高检出率和复合污染的特征。据统计,全球60%~80%的农作物均受到霉菌毒素污染[3]。动物摄入被污染的饲料后,不仅会损害肝脏、肾脏等器官功能,还可通过畜产品残留对人类健康构成潜在威胁[4-5]。饲料霉菌毒素污染已成为制约畜牧业可持续发展的重要因素,因此开发高效、安全的脱毒技术成为饲料与养殖行业的迫切需求。随着微生物制剂在养殖业中的广泛应用,其在饲料质量与安全防控方面的研究也逐渐深入。其中,芽孢杆菌因分布广泛、抗逆性强、稳定性高等特点,被广泛应用于饲料霉菌毒素的防治研究[6]。本文系统介绍了饲料中常见霉菌毒素的种类及其降解方法,重点综述了芽孢杆菌对霉菌毒素的降解效率与作用机制,以期为芽孢杆菌类脱霉剂的开发、高效降解菌株的筛选及其在饲料中的安全应用提供参考。

1 饲料中常见霉菌毒素种类

霉菌毒素种类繁多,目前已知的有300余种。在饲料及饲料原料中常见的霉菌毒素包括黄曲霉毒素B1(aflatoxin B1,AFB1)、玉米赤霉烯酮(zearalenone,ZEN)、赭曲霉毒素A(ochratoxin A,OTA)、脱氧雪腐镰刀菌烯醇(deoxynivalenol,DON)和伏马毒素(fumonisin,FB)等[7]

1.1 AFB1

黄曲霉毒素(AF)是由黄曲霉和寄生曲霉产生的次生代谢产物,常见类型包括AFB1、AFB2、AFM1、AFM2、AFG1和AFG2等,其中AFB1存在最为广泛且毒性最强[8]。AFB1的分子式为C17H13O6,化学结构式如图1所示。双呋喃环结构是其主要的毒性基团,香豆素环结构则介导其强致癌性[9]。多项研究表明,谷物、玉米及各类饼粕产品极易受到AFB1污染,且污染率因地区和原料类型不同而存在较大差异。其中玉米的污染率普遍在20%~50%,部分高温高湿地区可超过70%[10-12]。鉴于AFB1对动物及人体肝脏的强危害性,世界卫生组织(World Health Organization,WHO)已将其列为ⅠA级危险物,国际癌症研究机构(International Agency for Research on Cancer,IARC)将其列为Ⅰ类致癌物[13]
图1 AFB1化学结构式

红色为双呋喃环,蓝色为香豆素环。

Fig.1 Chemical structure formula of AFB1

Red indicates the difuran ring, and blue indicates the coumarin ring.

1.2 ZEN

ZEN是由镰刀菌属代谢产生的具有雌激素作用的毒素,其分子式为C18H22O5,化学结构式如图2所示。ZEN的主要毒性基团包括内酯环和苯环上的C-4游离酚羟基。由于ZEN的结构与雌二醇相似,其C-4羟基可直接竞争性结合雌激素受体,从而对动物生殖系统造成损害[14]。除生殖毒性外,ZEN还会对动物产生细胞毒性、免疫毒性及遗传毒性等多种毒性作用[15]。在饲料原料中,玉米及其副产物最易受到ZEN污染,同时该毒素也广泛分布于小麦、高粱等谷物中。调查显示,我国多个省份超20 000份饲料样本中,ZEN检出率高达85.0%~100.0%[16-17]
图2 ZEN化学结构式

红色为C-4游离酚羟基,蓝色为内酯环。

Fig.2 Chemical structure formula of ZEN

Red indicates the free phenolic hydroxyl group at C-4, and blue indicates the lactone ring.

1.3 OTA

OTA是赭曲霉、黑曲霉及青霉菌属等真菌产生的次生代谢产物,其分子式为C20H18CINO6,化学结构式如图3所示。OTA的主要毒性结构为异香豆素部分与苯丙氨酸部分的羧基,其分子上的氯原子还可增强毒性[18]。在机体中,OTA的主要毒性作用靶器官是肝脏和肾脏,其不仅能够抑制葡萄糖代谢,还可以与血浆蛋白高度结合形成难降解的物质,从而对机体产生持续毒性作用[19-20]。OTA广泛污染谷物及其副产品、肉类、乳制品、干果和配合饲料等。据报道,湿热地区谷物类饲料OTA污染率可达40%以上[21-22]。鉴于其致癌性,IARC已将其列为Ⅰ类致癌物[23]
图3 OTA化学结构式

红色为L-苯丙氨酸部分,蓝色为异香豆素部分。

Fig.3 Chemical structure formula of OTA

Red indicates the L-phenylalanine moiety, and blue indicates the isocoumarin moiety.

1.4 DON

DON由孢子丝核镰刀菌和梨孢镰刀菌等真菌产生,因可引发猪呕吐反应,又名呕吐毒素,其分子式为C15H20O6,化学结构式如图4所示。DON的主要毒性结构为12,13-环氧环与C-3羟基,且12,13-环氧环是其毒性的必需基团[24]。在机体内,DON通过抑制蛋白质合成和干扰细胞膜完整性等途径,损害胃肠道细胞和淋巴细胞的正常功能,从而破坏肠道内稳态并引发免疫功能障碍[25-27]。DON是小麦及其副产品中检出率最高且危害最严重的真菌毒素之一。多项调查结果表明,在小麦、大麦和玉米等多种饲料原料中均可检测到DON,检出率高达96.4%~100.0%[28-30]
图4 DON化学结构式

红色为C-3羟基,蓝色为12,13-环氧环。

Fig.4 Chemical structure formula of DON

Red indicates the hydroxyl group at C-3, and blue indicates the 12,13-epoxy ring.

1.5 FB

FB是由串珠镰刀菌和轮枝镰刀菌等镰刀菌属产生的次生代谢产物,有伏马毒素B1(FB1)、伏马毒素B2(FB2)和伏马毒素B3(FB3)等形式,其中FB1被认为是毒性最强的镰刀菌毒素之一[31]。FB的分子式为C34H59NO15,化学结构式如图5所示。其主要毒性结构为C-2位氨基与三羧酸侧链[32]。FB可侵害动物肝脏、肾脏和肠道等器官,通过诱发氧化应激反应阻碍其生长发育,并具有强致癌性[33-34]。其主要污染玉米及其副产物,以及燕麦、大麦和小麦等谷物[35]。多地调查结果显示,不同饲料样品均存在一定程度的FB污染,其中FB1和FB2的最高检出率分别可达96.7%和98.3%[36-37]
图5 FB化学结构式

红色为三羧酸侧链酯键,蓝色为C-2位游离氨基。

Fig.5 Chemical structure formula of FB

Red indicates the tricarboxylic acid side chain ester linkages, and blue indicates the free amino group at position C-2.

2 饲料霉菌毒素的脱毒方法

饲料中的霉菌毒素具有较高的热稳定性,传统处理方法难以有效降解毒素,反而可能破坏饲料中的营养成分,降低其饲用价值[38]。因此,开发高效、安全的霉菌毒素脱毒技术具有重要意义。目前常用的霉菌毒素脱毒方法分为物理脱毒法、化学脱毒法和生物脱毒法3类。

2.1 物理脱毒法

由于霉菌毒素具有较高的热稳定性,常规物理脱毒方法(如水洗、加热以及吸附剂吸附等)难以实现有效脱毒,因此当前研究多集中于采用蒙脱石、活性炭和沸石等吸附剂对饲料中的霉菌毒素进行物理去除。以蒙脱石为例,其拥有高比表面积和强吸附能力,被广泛用于吸附重金属及霉菌毒素等污染物[39]。天然蒙脱石对AFB1、FB、ZEN和DON等有一定的脱毒作用,并可降低其在动物组织中的残留[40-41]。然而,天然蒙脱石对ZEN和DON等极性较弱毒素的吸附能力有限[42]。因此,常通过有机改性、无机改性和复合改性等手段提升蒙脱石的吸附性能[43]。例如,Olopade等[44]报道,经钠离子改性后,蒙脱石对T-2毒素的吸附率从48%提高至66%。张立阳等[45]研究发现,载金属离子和季铵盐改性均能显著增强蒙脱石的脱毒能力,其中季铵盐改性蒙脱石对AFB1的吸附率可达98%以上。改性蒙脱石除具备优良的抗菌活性和吸附性能外,还对动物生长具有一定促进作用[46]。蔡孟君[47]研究发现,与天然蒙脱石相比,载铜蒙脱石对AFB1和DON的吸附率进一步提高;且添加至仔猪饲粮后可降低仔猪腹泻率,改善其生长性能、免疫功能和抗氧化能力。综上所述,改性蒙脱石等物理吸附剂在霉菌毒素脱毒中表现出良好的应用潜力,兼具减毒、促生长和增强免疫等多重功能。然而,其大规模应用仍面临一系列挑战,未来需进一步探索复合毒素的协同吸附机制、长期饲喂安全性以及低成本、规模化的改性工艺。

2.2 化学脱毒法

化学脱毒法主要利用碱化、酸化或臭氧处理等手段,将霉菌毒素降解或转化为无毒或弱毒的产物。传统化学脱毒方法(如利用化学试剂)虽可部分降解霉菌毒素,但受限于操作安全性、处理效果及潜在残留等问题,在实际生产中应用较少[48-49]。当前研究更多聚焦于新型化学脱毒剂,其中腐殖酸钠、臭氧和氨水等显示出较好的应用潜力[50]。腐殖酸钠被证实可以改善动物生长性能,并对重金属和真菌毒素等有害物质具有一定的吸附能力[51]。叶盛群[52]研究显示,腐殖酸钠对自然霉变玉米中常见的AFB1、ZEN和DON均具脱毒效果,在人工胃液中的去除率分别为25.00%、23.69%和10.27%,在人工肠液中则分别达到52.32%、38.66%和14.41%。此外,氨气熏蒸法不仅对饲料营养价值破坏小、成本较低,也适用于大规模处理。陈志娟等[53]报道,在温度37 ℃、氨气体积分数7.05%、氨熏96 h的条件下,霉变玉米中AFB1的降解率可达92%。尽管新型化学法在安全性与脱毒效能上优于传统方法,但与物理或生物法相比,仍面临化学物质残留风险较高、适用性较窄以及环境可持续性不足等瓶颈问题,其广泛应用仍需进一步探索与优化。

2.3 生物脱毒法

与传统物理和化学方法相比,生物脱毒法凭借其高效、安全及环保等优势,已成为当前霉菌毒素脱毒研究的热点。该方法主要通过微生物或其代谢产物作用于霉菌毒素分子的关键基团,将其转化为无毒或低毒的代谢产物,从而显著降低其对动物和人类的危害。
大量研究表明,多种微生物及其代谢产物(包括细菌、真菌及酶制剂)能够有效吸附或降解霉菌毒素[54-56]。例如,Sangmanee等[57]采用微量稀释法评估了源自发酵米粉的植物乳杆菌(Lactobacillus plantarum)K35的抗真菌活性,发现其可显著抑制黄曲霉的生长。Wang等[55]从鸡大肠消化物中分离出一株赖氨酸芽孢杆菌(Lysinibacillus lysinolyticus),其对ZEN的降解率超过90%。此外,部分真菌如酵母和曲霉也显示出抑制霉菌毒素生成并降解其毒性的能力。李梦珂等[58]报道,黑曲霉培养物对AFB1和ZEN的同步降解率分别达到71.54%和98.28%。邵春山等[59]以DON为唯一碳源分离得到一株毕赤酵母菌(Komagataella phaffii)MC-1,对DON的降解率为54.76%。在酶制剂方面,漆酶和锰过氧化物酶等因其高特异性、高效力及环境友好特性,展现出良好的应用潜力。谷晓丹[60]研究发现,漆酶Lac-W可广谱降解包括AFB1在内的6种霉菌毒素,降解率为12%~93%,尤以对AFB1的降解率最高。基于此,利用微生物降解霉菌毒素是一种具有广阔应用前景的防控策略。在众多微生物中,芽孢杆菌属因其高效降解特性成为当前研究的重点对象。

3 芽孢杆菌降解饲料中霉菌毒素的作用效果及机制

3.1 芽孢杆菌对不同种类霉菌毒素的降解作用

芽孢杆菌属是一类好氧或兼性厌氧的革兰氏阳性细菌,广泛分布于土壤、水、空气及动物肠道等环境中,具有较强的抗逆性[61]。该菌属常见菌种包括枯草芽孢杆菌(Bacillus subtilis)、地衣芽孢杆菌(Bacillus licheniformis)和蜡样芽孢杆菌(Bacillus cereus)等。芽孢杆菌属不仅可通过营养物质与空间竞争、诱导植物系统抗性等方式抑制病原真菌,还能分泌多种生物活性物质(如抗菌肽、细胞壁降解酶和环状脂肽类抗生素),直接抑制或杀伤产毒真菌,因此在食品、医疗和畜牧领域具有广泛的应用价值[62-63]
研究表明,常见的霉菌毒素如AFB1、ZEN、DON和OTA等均可被至少1种芽孢杆菌降解,其中AFB1与ZEN是主要的靶向毒素[64]。例如,地衣芽孢杆菌QT338和暹罗芽孢杆菌(Bacillus siamensis)DY3123对AFB1的降解率均超过90%[65-66];而ZEN可被贝莱斯芽孢杆菌(Bacillus velezensis)2X-7、斯氏芽孢杆菌(Bacillus spizizenii)B73和斯氏芽孢杆菌YQ-1高效降解,降解率分别达到98.30%、99.30%和99.07%[67-69]。在众多芽孢杆菌中,枯草芽孢杆菌作为脱霉剂的研究最为广泛,目前已有80余个菌株被鉴定具有霉菌毒素降解能力;此外,贝莱斯芽孢杆菌、蜡样芽孢杆菌和地衣芽孢杆菌等也是当前研究的重点[64]
饲料中的霉菌毒素常以复合形式存在,其协同毒性远高于单一毒素,因此近年来芽孢杆菌对复合毒素的降解能力备受关注。曹艳子等[70]利用自主分离的贝莱斯芽孢杆菌BSL-1进行降解试验,发现其发酵液原液对FB、DON、AFB1、ZEN和OTA均具降解能力,其中对AFB1和ZEN的降解率分别为50.60%和62.04%。Riahi等[71]研制了一种由枯草芽孢杆菌和地衣芽孢杆菌等组成的复合脱霉剂,可缓解OTA和T-2毒素对肉鸡生长性能的负面影响。Kumar等[72]则利用白芽孢杆菌(Bacillu albus)YUN5的无细胞上清液同步降解AFB1和AFG1,最高降解率分别为76.28%和98.98%。这些结果表明,芽孢杆菌在降解多种霉菌毒素方面表现出良好的广谱潜力。然而,由于多数芽孢杆菌缺乏对FB或T-2毒素等特定霉菌毒素的高效专一酶降解途径,单一菌株的降解能力仍有限。因此,在实际应用中多采用复合脱毒策略,例如将芽孢杆菌与蒙脱石等吸附剂、特定酶制剂复配,或整合物理、化学及生物脱毒方法,以全面提升脱毒效率。表1列举了其他研究中常用作脱霉剂的芽孢杆菌及其对多种霉菌毒素的降解效果。
表1 常见芽孢杆菌菌种对不同种类霉菌毒素的降解效果

Table 1 Degradation efficiency of common Bacillus species against different types of mycotoxins

芽孢杆菌菌种
Bacillus species
菌株编号
Strain numbers
降解霉菌毒素种类
Degraded mycotoxin species
降解率
Degradation rate/%
参考文献
References
枯草芽孢杆菌
Bacillus subtilis
JSW-1 AFB1 67.2 [73]
ZENL09 ZEN 87.0 [74]
KU-153 OTA 90.0 [75]
GMb2 DON 75.7 [76]
BIOUFLA2 FB1 40.0 [77]
贝莱斯芽孢杆菌
Bacillus velezensis
IS-6 AFB1 58.0 [78]
IS-6 ZEN 68.0 [78]
IS-6 OTA 89.0 [78]
E2 DON 31.0 [79]

蜡样芽孢杆菌
Bacillus cereus
XSWW9 AFB1 86.7 [80]
BC7 ZEN 100.0 [81]
地衣芽孢杆菌
Bacillus licheniformis
S51 AFB1 61.3 [82]
ANSB821 ZEN 90.0 [83]
DTM06 OTA 40.3 [84]
YB9 DON 82.7 [85]

AFB1:黄曲霉毒素B1 aflatoxin B1;ZEN:玉米赤霉烯酮 zearalenone;OTA:赭曲霉毒素A ochratoxin A;DON:脱氧雪腐镰刀菌烯醇 deoxynivalenol;FB1:伏马毒素B1 fumonisin B1

3.2 芽孢杆菌的脱毒作用机制

芽孢杆菌对霉菌毒素的脱毒作用机制主要包括吸附与降解2种方式(图6),其吸附机制依赖于细胞壁与霉菌毒素之间的非共价键结合。芽孢杆菌的细胞壁是由肽聚糖构成的多层网状结构,具有高比表面积和丰富的微孔通道,为物理吸附提供了结构基础[86]。为探究该机制,当前研究通常将菌液分离为不同组分(如灭活菌体、上清液等),并分别与霉菌毒素进行共培养。研究表明,部分芽孢杆菌以吸附作用为主要脱毒方式。例如,张家林[87]比较了贝莱斯芽孢杆菌CBMB205的灭活菌体(以吸附为主)、细胞壁组分(以吸附为主)及上清液(以酶降解为主)对AFB1和ZEN的脱除效果,发现细胞壁组分的脱毒效率显著高于其他组分,表明该菌株对这2种毒素的脱除以细胞壁吸附为主导机制。姜富贵等[88]也报道,纳豆枯草芽孢杆菌(Bacillus subtilis natto)对AFB1具有一定的物理吸附作用,最大吸附率可达17.76%。然而,多数研究显示,芽孢杆菌的上清液组分往往表现出更显著的脱毒效果[6,89-90],提示在芽孢杆菌属微生物的脱毒过程中,生物降解(尤其是胞外酶介导的降解)是其发挥高效脱毒功效的主要机制。
图6 芽孢杆菌对部分霉菌毒素的脱毒作用机制

AFB1:黄曲霉毒素B1 aflatoxin B1;ZEN:玉米赤霉烯酮 zearalenone;OTA:赭曲霉毒素A ochratoxin A;FB:伏马毒素 fumonisin。

Fig.6 Detoxification mechanisms of Bacillus against partial mycotoxins

降解是指益生菌通过其分泌的代谢产物(如酶)将毒素分解或转化为低毒或无毒产物的过程。在芽孢杆菌中,介导这一过程的关键酶主要包括漆酶、酯酶、磷酸转移酶、羧肽酶和过氧化物酶等[91-95]。这些酶可通过多种分子机制高效降解霉菌毒素,例如裂解酯键、打开环氧结构或在毒素核心骨架上引入官能团等,从而显著降低其毒性。Xiong等[96]发现,源自解淀粉芽孢杆菌(Bacillus amyloliquefaciens)B10的漆酶对AFB1的降解率可达79.3%,其机制在于漆酶活性位点的氨基酸残基可氧化AFB1的毒性关键部分——呋喃环末端,并添加羟基官能团,从而降低其毒性。González等[97]报道,源于芽孢杆菌的AHL内酯酶能够水解AFB1内酯环中的酯键,破坏其毒性基团。Xu等[98]研究表明,枯草芽孢杆菌CW14主要通过其D-丙氨酸羧肽酶A(DacA)裂解OTA的酰胺键,移除毒性基团L-苯丙氨酸,生成低毒产物赭曲霉毒素α(OTα)。Shi等[99]发现,枯草芽孢杆菌YT-4分泌的内酯水解酶可裂解ZEN的内酯环酯键,生成无雌激素活性的代谢物。Duviick等[100]则证实,源自解淀粉芽孢杆菌的羧酸酯酶能水解FB1的三羧酸侧链酯键,使其完全丧失神经毒性。
需指出的是,芽孢杆菌对霉菌毒素的脱除通常是酶降解与物理吸附协同作用的结果。例如,张家林[87]在研究贝莱斯芽孢杆菌CBMB205和暹罗芽孢杆菌KCTC 13613时发现,两者对AFB1、ZEN和DON兼具吸附与酶降解能力。李根[101]也报道,饲料类芽孢杆菌(Paenibacillus pabuli)E1除可分泌N-酰基高丝氨酸内酯酶破坏AFB1的呋喃环结构实现降解外,其细胞壁同时对AFB1具有一定的吸附作用。

3.3 芽孢杆菌相关脱霉剂在动物生产上的应用

鉴于微生物对霉菌毒素的良好降解效果,以及全面禁抗背景下对抗生素替代品的迫切需求,以芽孢杆菌制剂为代表的微生物制剂展现出巨大应用潜力。研究表明,芽孢杆菌具有调节动物肠道微生态平衡、促进营养物质消化与吸收、增强动物免疫功能,以及提高动物生产性能等益生效应[102]。在实际应用中,多项研究证实了芽孢杆菌制剂的有效性。例如,肖丹等[103]在艾维茵肉鸡试验中发现,枯草芽孢杆菌制剂不仅可降低黄曲霉毒素在机体内的残留,还能缓解AFB1引起的生长抑制、免疫器官损伤和抗氧化能力下降。林洋等[104]报道,饲喂贝莱斯芽孢杆菌制剂可减少肉鸡粪便中AFB1、FB、DON和ZEN的排泄量,同时显著提高肉鸡的试验末体重,降低死淘率和料重比。Zhou等[105]研究发现,枯草芽孢杆菌ANSB01G能够缓解ZEN对妊娠母猪造成的生殖激素紊乱、阴户肿胀、器官损伤及细胞凋亡等负面影响。但需要注意的是,并非所有芽孢杆菌均为益生菌,部分菌株如某些蜡样芽孢杆菌可能引起恶心、呕吐等轻微中毒症状,甚至导致严重的心内膜炎或脓毒症等并发症[106]。因此,系统评估芽孢杆菌制剂的安全性并深入研究其对动物生产性能的影响,是推动其在养殖业中广泛应用的重要前提。
除体内试验外,细胞毒性试验和饲粮营养成分分析也常用于评估芽孢杆菌的脱毒效果及安全性。Guo等[107]发现,在肉鸡饲粮中添加含枯草芽孢杆菌CGMCC1.0504的复合益生菌,可改善肉鸡肠道菌群,缓解霉菌毒素对其生产性能的不利影响,并减轻AFB1与ZEN对肝细胞的毒性作用。王睿等[108]研究表明,地衣芽孢杆菌不仅能有效降低玉米青贮饲料中DON、FB和AFB1的含量,还可提高饲料粗脂肪含量,降低中性洗涤纤维和酸性洗涤纤维含量。石楠等[109]利用蜡样芽孢杆菌BT-1对霉变玉米进行体外发酵,结果显示,AFB1、DON和ZEN的脱毒率分别达到67.59%、73.03%和65.71%,且对青贮饲料和玉米籽粒的营养成分含量无显著负面影响。这些结果突显了芽孢杆菌在脱除霉菌毒素的同时,兼具保持甚至改善饲料营养价值的双重优势。

3.4 芽孢杆菌脱毒应用的挑战

尽管芽孢杆菌在饲料霉菌毒素脱毒方面展现出广阔的应用前景,其实际推广仍面临一系列挑战。首先,不同菌株的降解谱相对狭窄,对部分毒素的降解效率仍不理想;此外,在多种毒素共存的复杂基质中,其降解效率往往进一步下降。如Hassan等[110]研究发现,巨大芽孢杆菌(Priestia megaterium)BM344-1对ZEN的降解率接近100%,但对FB1和FB2的降解率仅为14%和12%;尤其在ZEN、T-2毒素、FB1和FB2混合污染的条件下,其对ZEN和T-2毒素的降解效率更是被严重抑制。其次,关键降解酶及其分子机制尚未完全阐明,制约了针对性的菌株改造与功能强化[111]。同时,现有研究中仍相对缺乏对芽孢杆菌降解霉菌毒素后代谢产物的系统毒理学评估。如AFB1的代谢产物AFB2a、ZEN的代谢产物α-玉米赤霉烯醇虽较原始毒素毒性降低,但仍可能引发氧化应激、DNA损伤或雌激素样反应等潜在危害[112-113]。此外,部分菌株的生物安全性、工业化制剂的稳定性与生产成本等问题也亟待解决。

4 小结与展望

饲料中常见的霉菌毒素(如AFB1、ZEN、DON、OTA和FB等)不仅严重威胁动物和人类健康,也制约着畜牧业的可持续发展。虽然物理和化学脱毒方法可在一定程度上降低霉菌毒素的毒性,但存在营养物质损失、成本较高和易造成二次污染等问题,难以满足当前饲料工业对绿色、安全与高效生产的需要。因此,生物脱毒技术逐渐成为霉菌毒素治理的研究和应用热点。近年来,大量研究证实多种芽孢杆菌菌株对上述霉菌毒素具有显著的降解能力,其作用机制主要包括酶促降解与细胞吸附。其中,漆酶、酯酶、磷酸转移酶等代谢酶类能够定向破坏毒素的关键毒性结构,并将其转化为低毒或无毒的产物,表现出良好的应用潜力。在动物生产上,部分芽孢杆菌不仅能有效降解毒素,还可改善动物肠道菌群、增强机体免疫力以及缓解毒素引发的器官损伤,展现出其作为安全脱霉剂的极大潜力。然而,该技术目前仍面临一些挑战,包括菌株降解谱窄、对某些毒素降解效率不高、作用机制尚未完全明晰等。今后的研究应致力于筛选广谱高效菌株、解析关键代谢途径及功能酶基因、评估复合毒素条件下的降解稳定性与安全性,并推动其与物理、化学方法的协同使用,以实现多种霉菌毒素的高效协同脱除,为保障饲料安全与畜禽健康提供更加可靠的技术支撑和解决方案。
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