Article(id=1307007593197630431, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307007396681900041, articleNumber=null, orderNo=null, doi=10.11843/j.issn.0366-6964.2026.01.004, pmid=null, cstr=null, oa=null, hot=null, price=null, onlineType=0, articleFormat=0, articleType=null, articleTypeStr=research-article, receivedDate=1735488000000, receivedDateStr=2024-12-30, revisedDate=null, revisedDateStr=null, acceptedDate=null, acceptedDateStr=null, onlineDate=1789544493244, onlineDateStr=2026-09-16, pubDate=1769097600000, pubDateStr=2026-01-23, doiRegisterDate=null, doiRegisterDateStr=null, onlineIssueDate=1789544493244, onlineIssueDateStr=2026-09-16, onlineJustAcceptDate=null, onlineJustAcceptDateStr=null, onlineFirstDate=null, onlineFirstDateStr=null, sourceXml=null, magXml=null, createTime=1789544493244, creator=13701087609, updateTime=1789544493244, updator=13701087609, issue=Issue{id=1307007396681900041, tenantId=1146029695717560320, journalId=1306274286668972124, year='2026', volume='57', issue='1', pageStart='1', pageEnd='568', issueExtLink='null', onlineDate='null', pubDate='1769097600000', pubDateStr='2026-01-23', beforeIssueId=null, nextIssueId=null, price=null, status=1, issueComplete=1, articleOrder=1, issueType=-1, specialIssue=null, createTime=1789544446392, creator='13701087609', updateTime=1789609890784, updator='13701087609', preIssue=null, nextIssue=null, articleTotal=null, ext={EN=IssueExt(id=1307281890638057732, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307007396681900041, language=EN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=), CN=IssueExt(id=1307281890738721029, tenantId=1146029695717560320, journalId=1306274286668972124, issueId=1307007396681900041, language=CN, specialIssueTitle=, coverIllustrator=null, specialIssueEditor=, specialIssueAbout=)}, issueFiles=null, downloadFileDto=null}, startPage=30, endPage=45, ext={EN=ArticleExt(id=1307007593558340577, articleId=1307007593197630431, tenantId=1146029695717560320, journalId=1306274286668972124, language=EN, title=Biological Characteristics of Enterococcus faecalis and Its Application in Animal Husbandry, columnId=1307007397399126026, journalTitle=Acta Veterinaria et Zootechnica Sinica, columnName=REVIEW, runingTitle=null, highlight=null, articleAbstract=

Antibiotics once played an essential role in the process of China’s livestock industry. With the continuous increase in people’s attention to food safety and ecological environment issues, the adverse effects of antibiotic abuse have gradually become prominent. China’s feed industry has ushered in an era of a complete ban on antibiotics, and the livestock industry is facing significant challenges. Probiotics, as feed additive with great potential, can affect the composition of the intestinal flora after being ingested by animals, improve the intestinal barrier and regulate the immune response, showing unique advantages and value. It is expected to replace antibiotics and play a role in the livestock industry. Enterococcus faecalis, as a member of the lactic acid bacteria group, can be used as a feed additive and has attracted much attention. Enterococcus faecalis has relatively strong resistance to the environment, and its biological characteristics determine whether it can reach the host’s intestinal tract with a certain number of viable bacteria, and then determine whether this bacterium can be widely used in the livestock industry. As an active microorganism beneficial to the host, Enterococcus faecalis can regulate the balance of the body’s intestinal flora, enhance the function of the intestinal epithelial barrier, promote the metabolism of nutrients, and enhance the body's immunity. There is an increasing number of studies on the impact of Enterococcus faecalis on livestock and poultry. It has shown positive effects such as improving production performance, reducing the incidence of diarrhea and mortality, and enhancing the immune system in various livestock and poultry, including pigs, cattle, sheep, and chickens. This bacterium can optimize the digestive function of livestock and poultry, improve feed efficiency, and thus increase the economic benefits of farms. Its application range in the livestock industry is also constantly expanding. This article focuses on Enterococcus faecalis, summarizes its application in livestock and poultry in recent years, and reviews its biological characteristics, mechanisms of action, applications in the livestock and poultry industry, and the existing problems of Enterococcus faecalis. The aim is to provide a new insights from previous research about this bacterium and its impact in the livestock and poultry industry.

, authors=Ziqi TIAN1, 2, Yan TU1, Linshu JIANG2, Yanliang BI1, authorsList=Ziqi TIAN, Yan TU, Linshu JIANG, Yanliang BI, authorCompany=null, correspAuthors=Yanliang BI, authorNote=null, correspAuthorsNote=
BI Yanliang,E-mail:
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抗生素曾在我国畜牧业的养殖过程中扮演重要角色。随着人们对食品安全以及生态环境等问题关注度的持续提升,抗生素滥用的负面效应逐渐凸显,我国饲料业迎来全面禁抗时代,畜牧业面临重大挑战。益生菌作为一类极具潜力的饲料添加剂,被动物摄入后能影响肠道菌群的组成,改善肠道屏障以及调节免疫反应,展现出独特的优势和价值,有望代替抗生素在畜牧业中发挥作用。粪肠球菌作为一种乳酸菌,可用作饲料添加剂且备受关注。粪肠球菌对环境的抗性较强,其生物特性决定其能否以一定活菌数量抵达宿主肠道,进而决定该菌能否在畜牧业中广泛应用。粪肠球菌作为对宿主有益的活性微生物,能够调节机体肠道菌群平衡、增强肠道上皮屏障功能、促进营养物质代谢、增强机体免疫力。粪肠球菌对畜禽影响的研究日益增多,对猪、牛、羊、鸡等多种畜禽均表现出可提高生产性能、降低腹泻率和死亡率以及提高机体免疫力等积极作用,该菌能优化畜禽的消化功能,提高饲料利用率,进而提高养殖场的经济效益,在畜牧业中的应用范围也在不断拓展。本文围绕粪肠球菌,总结近些年粪肠球菌在畜禽中的应用,对粪肠球菌的生物特性、作用机制、在畜牧业上的应用以及目前存在的问题进行综述,旨在为该菌的后续研究提供参考,使其在畜牧业中发挥更大作用。

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田子琦,硕士生,主要从事反刍动物营养与饲料科学研究,E-mail:

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毕研亮,主要从事幼龄反刍动物营养与调控研究,E-mail:
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粪肠球菌生物特性及在畜牧业上的应用
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田子琦 1, 2 , 屠焰 1 , 蒋林树 2 , 毕研亮 1
畜牧兽医学报 | 综述 2026,57(1): 30-45
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粪肠球菌生物特性及在畜牧业上的应用
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田子琦1, 2 , 屠焰1, 蒋林树2, 毕研亮1
作者信息
  • 1.中国农业科学院饲料研究所 农业农村部饲料生物技术重点实验室 奶牛营养学北京市重点实验室,北京 100081
  • 2.北京农学院动物科学技术学院,北京 102206
通讯作者:
毕研亮,主要从事幼龄反刍动物营养与调控研究,E-mail:
作者简介:

田子琦,硕士生,主要从事反刍动物营养与饲料科学研究,E-mail:

Biological Characteristics of Enterococcus faecalis and Its Application in Animal Husbandry
Ziqi TIAN1, 2 , Yan TU1, Linshu JIANG2, Yanliang BI1
Affiliations
  • 1.Beijing Key Laboratory of Dairy Cow Nutrition,Key Laboratory of Feed Biotechnology,Ministry of Agriculture and Rural Affairs,Institute of Feed Research of Chinese Academy of Agricultural Sciences,Beijing 100081,China
  • 2.College of Animal Science and Technology,Beijing University of Agriculture,Beijing 102206,China
  • Corresponding Author:
    BI Yanliang,E-mail:
出版时间: 2026-01-23 doi: 10.11843/j.issn.0366-6964.2026.01.004
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抗生素曾在我国畜牧业的养殖过程中扮演重要角色。随着人们对食品安全以及生态环境等问题关注度的持续提升,抗生素滥用的负面效应逐渐凸显,我国饲料业迎来全面禁抗时代,畜牧业面临重大挑战。益生菌作为一类极具潜力的饲料添加剂,被动物摄入后能影响肠道菌群的组成,改善肠道屏障以及调节免疫反应,展现出独特的优势和价值,有望代替抗生素在畜牧业中发挥作用。粪肠球菌作为一种乳酸菌,可用作饲料添加剂且备受关注。粪肠球菌对环境的抗性较强,其生物特性决定其能否以一定活菌数量抵达宿主肠道,进而决定该菌能否在畜牧业中广泛应用。粪肠球菌作为对宿主有益的活性微生物,能够调节机体肠道菌群平衡、增强肠道上皮屏障功能、促进营养物质代谢、增强机体免疫力。粪肠球菌对畜禽影响的研究日益增多,对猪、牛、羊、鸡等多种畜禽均表现出可提高生产性能、降低腹泻率和死亡率以及提高机体免疫力等积极作用,该菌能优化畜禽的消化功能,提高饲料利用率,进而提高养殖场的经济效益,在畜牧业中的应用范围也在不断拓展。本文围绕粪肠球菌,总结近些年粪肠球菌在畜禽中的应用,对粪肠球菌的生物特性、作用机制、在畜牧业上的应用以及目前存在的问题进行综述,旨在为该菌的后续研究提供参考,使其在畜牧业中发挥更大作用。

粪肠球菌  /  生物特性  /  畜牧业  /  饲料添加剂

Antibiotics once played an essential role in the process of China’s livestock industry. With the continuous increase in people’s attention to food safety and ecological environment issues, the adverse effects of antibiotic abuse have gradually become prominent. China’s feed industry has ushered in an era of a complete ban on antibiotics, and the livestock industry is facing significant challenges. Probiotics, as feed additive with great potential, can affect the composition of the intestinal flora after being ingested by animals, improve the intestinal barrier and regulate the immune response, showing unique advantages and value. It is expected to replace antibiotics and play a role in the livestock industry. Enterococcus faecalis, as a member of the lactic acid bacteria group, can be used as a feed additive and has attracted much attention. Enterococcus faecalis has relatively strong resistance to the environment, and its biological characteristics determine whether it can reach the host’s intestinal tract with a certain number of viable bacteria, and then determine whether this bacterium can be widely used in the livestock industry. As an active microorganism beneficial to the host, Enterococcus faecalis can regulate the balance of the body’s intestinal flora, enhance the function of the intestinal epithelial barrier, promote the metabolism of nutrients, and enhance the body's immunity. There is an increasing number of studies on the impact of Enterococcus faecalis on livestock and poultry. It has shown positive effects such as improving production performance, reducing the incidence of diarrhea and mortality, and enhancing the immune system in various livestock and poultry, including pigs, cattle, sheep, and chickens. This bacterium can optimize the digestive function of livestock and poultry, improve feed efficiency, and thus increase the economic benefits of farms. Its application range in the livestock industry is also constantly expanding. This article focuses on Enterococcus faecalis, summarizes its application in livestock and poultry in recent years, and reviews its biological characteristics, mechanisms of action, applications in the livestock and poultry industry, and the existing problems of Enterococcus faecalis. The aim is to provide a new insights from previous research about this bacterium and its impact in the livestock and poultry industry.

Enterococcus faecalis  /  biological characteristics  /  animal husbandry  /  feed additives
田子琦, 屠焰, 蒋林树, 毕研亮. 粪肠球菌生物特性及在畜牧业上的应用. 畜牧兽医学报, 2026 , 57 (1) : 30 -45 . DOI: 10.11843/j.issn.0366-6964.2026.01.004
Ziqi TIAN, Yan TU, Linshu JIANG, Yanliang BI. Biological Characteristics of Enterococcus faecalis and Its Application in Animal Husbandry[J]. Acta Veterinaria et Zootechnica Sinica, 2026 , 57 (1) : 30 -45 . DOI: 10.11843/j.issn.0366-6964.2026.01.004
抗生素残留导致环境污染和细菌耐药的问题给食品安全及生物健康造成严重影响[1]。我国农业农村部第194号公告中指出,2020年7月1日起,饲料生产企业不能再生产含有促生长类药物饲料添加剂的商品饲料,进入全面禁抗时代,养殖业面临严峻挑战[2]。益生菌作为有益的活性微生物,可用作饲料添加剂,改善肠道微生态平衡以及畜禽的生长性能[3],是良好的抗生素替代品[4]。在益生菌中,粪肠球菌作为一种乳酸菌,能够产生有机酸,降低宿主肠道的pH,促进有益乳酸菌生长繁殖并抑制不耐酸病原菌生长[5]。此外,该菌的生物特性使其在饲料添加剂领域颇具应用潜力。1984年,美国饲料控制委员会(American Association of Feed Control Office,AAFCO)和美国食品药品监督管理局(Food and Drug Administration,FDA)已批准在饲料中使用屎肠球菌(Enterococcus faecium)和粪肠球菌(Enterococcus faecalis),我国农业农村部发布的《饲料添加剂品种目录(2008)》也将这两种菌纳入其中,说明它们可以被用作饲料添加剂[6]。粪肠球菌逐渐进入人们视野且受到广泛关注,诸多研究已经证实该菌具备优良的生物学性质,能以活菌状态抵达宿主胃肠道,进而发挥其益生菌功能,对宿主产生积极影响,在益生菌研究及饲料添加剂应用领域展现出特有的价值与潜力。本文综述了粪肠球菌的生物特性、作用机制、在畜牧业中的应用以及目前存在的问题与挑战,旨在推动粪肠球菌在畜牧生产中的应用并提供参考。
肠球菌属(Enterococcus)是乳酸菌的主要菌属之一,是普遍存在于人和动物肠道内的兼性厌氧菌,很容易在生产过程中的需氧条件下以及在瘤胃内的厌氧条件下培养和增殖[3]。粪肠球菌属于肠球菌属,为革兰阳性菌,菌体呈球形,直径为0.3~0.7 μm,没有芽孢和荚膜,分泌L型乳酸,是普遍存在于人和动物消化道内的微生物之一,有较强的肠黏膜定植和环境耐受能力[7⁃8]。该菌对其生长条件的要求相对宽松,具有繁殖速度快、环境适应性强等优点,适合作为益生菌用于饲料添加剂领域[9]。
温度是影响细菌生长的重要因素之一。粪肠球菌普遍存在于消化道内,其对温度的适应范围较为广泛。向盈盈等[10]发现,粪肠球菌(E.faecalis YN771)可在20 ℃~42 ℃的温度范围内生长繁殖,最适温度约为37 ℃。Wang等[11]在婴儿粪便中分离出7种粪肠球菌,所有菌株均可在25 ℃~45 ℃下分别进行有氧和无氧生长,且最适温度也为37 ℃。在50 ℃的发酵条件下粪肠球菌DUT1805仍能进行乳酸发酵,且乳酸的转化率可达90%左右[12]。30 ℃和40 ℃时,粪肠球菌的存活率较高且活性较高,50 ℃时粪肠球菌仍可存活[13]。由此表明,粪肠球菌对温度的耐受范围较广,能够很好地适应畜禽消化道内温度。
动物胃液中的主要成分是盐酸。单胃动物胃液pH一般在2.0~2.5[14],反刍动物的皱胃胃液pH约为2.0~3.0,胃液酸度会随动物摄食有所下降。在pH=3的环境下存活1.5~2 h是衡量益生菌低pH耐受性的标准之一。益生菌的耐酸性决定其能否以活菌状态在肠道定植并发挥作用[15]。因此,粪肠球菌是否具有一定的耐酸性至关重要。许喜林等[16]分离出的粪肠球菌EF-ZA1107-06在pH=2.0的酸溶液中耐受时间不足2.0 h,而在pH=2.5、3.0、4.0处理组中的菌株生长不受影响,表明该菌株具有较好的pH耐受性。粪肠球菌BN6株在pH=2.5和pH=3.0的环境3 h后,其存活率仍达到97.00%以上[15]。由此可知,粪肠球菌具有较好的耐酸能力,可以以活菌的形式通过宿主胃部抵达肠道进行定植。
胆盐是动物机体不可或缺的成分,在细胞外产生高渗透压环境,对菌体细胞的影响显著。益生菌需具备良好的耐胆盐性才能在动物胃肠道中保持一定的数量。因此,粪肠球菌要对胆盐具有一定耐受,才能在肠道内存活并定植,进而发挥益生功能[17]。动物消化道内胆汁的浓度为0.03%~0.3%,因此,通常以0.3%牛胆盐浓度作为菌株耐胆盐性的筛选标准,刘明珠[18]发现,浓度为0.2%~0.4%的胆盐对乳酸粪肠球菌存活率的影响不大。在0.3%浓度的胆盐环境下,粪肠球菌依然能够生长繁殖并且存活状态良好[19]。从中国健康婴儿粪便中分离的7种粪肠球菌,除了菌株MH385355外,其他六种菌株对胆盐表现出很强的耐受性[11],表明该菌对胆盐具有良好耐受,能够通过胃肠道并达到一定数量,进而在肠道中定植[20]。
细菌对宿主组织细胞黏附性的强弱是决定其入侵组织并定植的关键,是重要的起始步骤[21]。IPEC-J2细胞系是研究肠道细菌相互作用和益生菌对宿主作用的理想工具[22]。肠球菌的黏附蛋白可结合到细胞外基质蛋白质上(如胶原蛋白和黏蛋白)[11]。粪肠球菌能黏附到IPEC-J2表面以及细胞间的紧密连接处,呈不均匀分布状态,同时还会出现聚集现象[23]。与沙门菌或金色葡萄球菌共培养能够促进粪肠球菌N14对IPEC-J2细胞的黏附作用[24]。从健康婴儿粪便中分离的7种粪肠球菌表现出10.6%~54.9%的Caco-2细胞系黏附能力[11]。此外,粪肠球菌的胶原黏附素ace基因能促进菌株生物膜形成,增强黏附性[25],其菌毛也可加快黏附速度,对菌株的黏附十分重要[26]。由此可知,粪肠球菌到达肠道后能够黏附于肠道细胞表面,进而在肠道中定植并发挥益生作用。
肠球菌在生长过程中能产生有机酸、过氧化氢、肠球菌素(第II类细菌素)以及其他抑菌物质,其培养基上清液对某些病原体具有广谱的抑制作用[11]。粪肠球菌能产生细菌素等抑菌物质,不仅能够抑制沙门菌和大肠杆菌等病原菌的生长,进而改善宿主肠道微生态环境,还能够抑制肠道中代谢腐败菌和尿素酶的细菌生长,使肠道内尿素酶和内毒素含量降低,改善宿主的血液环境[27]。粪肠球菌L76可以通过黏附上皮细胞来有效保护细胞免受沙门菌侵袭[28]。车笑吟等[29]研究发现,粪肠球菌分泌的细菌素对奶牛乳房炎源S.aureus具有良好的抗菌活性,该菌的抗菌活性还可有效防治奶牛产后子宫内膜炎[30]。粪肠球菌与大肠杆菌O1和O78混合培养8 h后繁殖达到高峰,10 h后对大肠杆菌O1和O78表现出极显著的抑制作用[9]。从健康婴儿粪便中分离的7种粪肠球菌均展现出不同的抗菌活性[11],表明粪肠球菌在肠道定植后能抑制有害菌生长,调节肠道微生态环境。
粪肠球菌的生物特性表现出对环境的广泛适应性,菌株良好的抗逆性使其能在动物体内保持较高活性,其优良的耐酸耐胆盐能力,能够很好地适应不同动物的胃肠道坏境且能维持一定数量,该菌的黏附性为其在宿主肠道内完成定植并发挥益生菌的功能奠定基础。此外,粪肠球菌的抑菌性在一定程度上抑制病原菌生长繁殖,维持动物肠道健康,以上特性符合益生菌作为饲料添加剂在畜牧养殖领域的要求且表现出广阔的应用前景和潜力。
粪肠球菌代谢产生有机酸,可以在一定程度上使肠道环境的pH降低,抑制或阻止肠道内不耐酸的厌氧细菌生长[31],例如抑制致病性大肠杆菌和沙门菌的繁殖。王灿宇等[32]在锦江黄牛的饲料中添加粪肠球菌复合菌后,观察到牛后肠道中的大肠杆菌数量减少,而乳酸杆菌数量有显著上升的趋势。这一现象表明,粪肠球菌复合菌能够在一定程度上提升动物肠道内优势菌群的数量,有效改善锦江黄牛的肠道内环境,促进肠道健康。艾玉[33]研究发现,在饲料中添加粪肠球菌F7可增加凡纳滨对虾消化道内有益菌丰度,降低有害菌丰度,进而维持肠道健康。细菌在宿主肠道定植后才能具有生物活性。粪肠球菌较强的黏附性可以与有害病原菌竞争肠道细胞表面积,减少病原菌定植。同时与其他有益菌构成优势菌群,抑制病原菌的繁殖,进而优化宿主肠道环境,维持肠道微生态平衡[6],为宿主肠道健康提供保障。
肠道上皮作为机体关键的防御结构,能有效阻挡病原微生物的入侵,降低机体遭受外源微生物感染风险,更是重要的免疫屏障,能够阻止病原菌及内毒素穿越肠道黏膜,从而保障机体免受侵害,在维护机体肠道健康与整体稳态中发挥重要作用[34]。粪肠球菌可以通过竞争性黏附并聚集在肠道表面形成生物膜,阻隔病原菌在肠道表面的定植[35]。粪肠球菌等肠球菌可分泌亚油酸异构酶,将亚油酸转化为共轭亚油酸(CLA)。CLA作为信号刺激 CD4+CD8αα+上皮内淋巴细胞发育并驻留在小肠中,CLA通过与核受体HNF4γ结合,调控免疫细胞相关信号,影响肠道屏障功能[36]。肠道上皮细胞的紧密连接是肠道物理屏障的关键,其中紧密连接蛋白发挥着至关重要的作用。粪肠球菌能够上调紧密连接蛋白的mRNA相对表达量,通过增强紧密连接蛋白基因的表达对肠道屏障功能进行有效调节[37]。此外,Chichlowski等[38]发现,粪肠球菌等复合菌对雏鸡杯状细胞的生长和成熟有促进作用,杯状细胞所分泌的黏液素,能够促进机体黏液蛋白的分泌,进而增强菌株的黏附能力,在肠道内形成一道保护屏障,有效抵御毒素对肠道上皮细胞造成的侵害。
粪肠球菌不仅能够分泌乳酸和乙酸[39]供机体吸收利用,还具备分泌多种酶和维生素的能力,加快分解多种碳水化合物,促进营养物质吸收,提高动物生产水平[4]。在饲料中添加粪肠球菌F7有利于提高凡纳滨对虾肠道内脂肪酶和胰蛋白酶活性,增强对虾对营养物质的消化吸收[33]。荆祎等[40]研究发现,在饲料中添加粪肠球菌可以提高水貂对脂肪和蛋白质的消化率和吸收率,进而提高水貂的生产性能。此外,粪肠球菌还可以保护肠绒毛形态及其完整性,改善断奶仔猪肠道屏障功能。Wang等[4]研究发现,粪肠球菌能降低断奶仔猪隐窝深度,提高绒毛高度,有利于断奶仔猪的肠道健康发育和对养分的吸收。肠球菌分泌的挥发性脂肪酸可以激活mTOR信号通路,促进蛋白质合成,增加细胞的质量和体积,进而促进肠绒毛的生长[41⁃42]。杨珍珠等[43]在羊奶中分离的1株粪肠球菌DH9003能够使D-谷氨酰胺、L-亮氨酸和天冬氨酸等124种营养物质代谢上调,说明粪肠球菌对机体的代谢能力也具有一定的调节作用。
肠道是身体主要免疫系统之一,肠道上皮细胞形成关键屏障,被认为在免疫系统中起着重要作用[44]。粪肠球菌能够黏附于肠道上皮细胞刺激局部黏膜免疫[45],其分泌的生物素以及代谢物质能够在肠道表面形成屏障[33],阻止病原菌的入侵。Wang等[46]从新生儿、发酵乳和培养基中分离出的27株粪肠球菌中,发现有4株能通过降低白细胞介素-8(IL-8)的分泌和增加TGF-β(一种重要的炎症调节细胞因子)的表达来调节宿主免疫。鱼料中添加粪肠球菌B-8可诱导鲫鱼体液免疫,提升免疫球蛋白水平[47]。Tarabees等[48]成功地从健康鸡上分离到了1株粪肠球菌(E. faecalis-1),在肉鸡饲料中添加该粪肠球菌提高了动物机体免疫应答,降低了肉鸡死亡率。Pedicord等[49]发现粪肠球菌(E.faecalis OG1RF)限制小鼠沙门菌的发病机制,且其在定植时能够上调肠上皮细胞(IEC)相关基因的表达水平。IEC产生的干扰素-γ(IFN-γ)可促进IEC中主要组织相容性复合体(MHC)与程序性死亡受体-配体1(PD-L1)的表达,进而推动肠上皮内淋巴细胞(IEL)中CD4⁺T细胞分化。此外,IEC产生的缺氧诱导因子1α(HIF-1α)能促进IEL分化,以及CD4⁺CD8αα⁺T细胞亚群与T细胞受体γδ(TCRγδ)的表达[41],他们还发现,粪肠球菌的定植使上皮细胞和细菌分离,保护消化道不被细菌入侵,进而增强机体免疫力。此外,粪肠球菌F7能提高凡纳滨对虾对溶藻弧菌HN08155的抗感染能力[33],进而增强机体免疫力。
粪肠球菌通过相对较强的黏附性抢占定植位点,分泌有机酸形成酸性环境以及兼性厌氧的特性,均能使其在与有害病原菌竞争时体现较强的优势,很容易与其他有益菌形成优势菌群进一步抑制有害菌的生长繁殖,调节肠道菌群结构。此外,该菌的代谢产物不仅能够被动物机体吸收利用,还能积极促进其他营养物质的吸收,并且能够在肠道表面形成生物屏障,提高动物生长性能的同时保护肠道免受病原菌侵害,增强机体免疫力,为动物的健康生长和良好发育提供保障,在肠道健康和畜牧养殖领域具有重要意义。
在猪的不同生理阶段,日粮中添加粪肠球菌的目的不同。新生仔猪饲喂粪肠球菌可以帮助有益菌在其肠道内定植,降低仔猪腹泻率和死亡率,为仔猪的健康生长奠定良好基础。哺乳仔猪灌服粪肠球菌能使其日增重增加,提高断奶重[50],使仔猪以良好的机体状态进入下一个生长阶段。仔猪断奶后饲喂粪肠球菌的效果也很显著,不仅可以提高仔猪平均日增重、降低料重比和腹泻率,还能改善肠道消化吸收功能,提高断奶仔猪的生长性能[51-52]。此外,粪肠球菌能显著抑制大肠杆菌的增殖,促进肠道微生态平衡,显著增加血清中IgA、IgG浓度,提高仔猪免疫力[53-55]。粪肠球菌还能促进仔猪肠道发育,提高氨基酸的消化率。饲粮中同时添加枯草芽孢杆菌与粪肠球菌后,可提高仔猪空肠中短链脂肪酸的相对含量[56]。魏清甜等[57]在保育仔猪的饲料中添加不同剂量的粪肠球菌,对其生长性能与腹泻率均有显著改善效果,在保育前期表现尤为明显,还可提高保育仔猪的体液免疫,提升对粗蛋白质和粗脂肪的表观消化率,利于肠道发育,维持保育仔猪肠道微生态平衡。刘先高等[58]研究发现,粪肠球菌添加量会对仔猪后期生长性能产生一定程度的影响。薛洋[59]在其研究中发现,粪肠球菌促进了仔猪脾组织发育,显著降低回肠pH,提高仔猪空肠绒毛高度[4]。长期将粪肠球菌拌料饲喂或饮水饲喂,可以有效缩短育肥猪的出栏天数,提高猪的整齐度[60],便于管理和销售,能够从整体上提高养殖效益。
在猪的养殖生产中,粪肠球菌可以帮助新生仔猪肠道内有益菌的定植,降低仔猪腹泻率和死亡率,且有利于提高断奶重。仔猪断奶后饲喂粪肠球菌不仅能够增强机体免疫力,抑制大肠杆菌等有害菌的生长繁殖,减少相关疾病发生,还能有效促进营养物质吸收,提高饲料的利用率,降低料重比,减少饲料浪费,提高经济效益。整体来看,粪肠球菌在养猪业中表现出较高的应用价值。
在家禽生产中粪肠球菌的应用也较为广泛。雏鸡饲喂粪肠球菌后,回肠绒毛及绒毛长度增加,提高了雏鸡对营养的吸收[61]。王禹盟等[61]将母源粪菌移植给雏鸡后,发现其能够促进免疫器官发育和成熟。他们还发现粪肠球菌可以改善肠道菌群结构,丰富物种多样性。于会民等[62]研究发现,在给肉仔鸡饲喂葡萄糖氧化酶的基础上添加粪肠球菌能在不同程度上改善肉仔鸡的养分表观消化率,提高生长性能和抗氧化能力。此外,添加粪肠球菌可明显改善肉仔鸡屠宰性能,降低料肉比,改善肉仔鸡肉品质[63-64]。复合脂肪酸酯与粪肠球菌联用能促进肉鸡早期肠道形态发育,促进肉鸡早期肠道健康[65]。Huang等[66]研究发现,粪肠球菌能显著减轻肠黏膜病理损伤,改善肠黏膜屏障,减轻肠炎感染。在蛋鸡饲料中添加粪肠球菌,发现可以提高产蛋量、蛋黄颜色、蛋白高度和哈氏单位,但其作用机理有待进一步研究[67]。Li等[68]从鸡粪中分离出1株粪肠球菌(E.faecalis CML390),无论是否存在氧气都能降低尿酸,减少痛风的发生,保障家禽正常活动和生长发育。同时为家禽养殖过程中尿酸代谢异常问题提供了探索方向,推动家禽养殖业营养调控和管理领域发展。
粪肠球菌在家禽养殖中有诸多作用。雏禽阶段饲喂粪肠球菌可以优化肠道形态,促进营养物质吸收和免疫器官发育并丰富肠道菌群的多样性。在肉禽养殖中,粪肠球菌与其他有益菌或有益物质联合饲喂时,常表现出协同效应,缓解肠炎等疾病的感染,降低肠道损伤,保障肉禽肠道功能,提高肉禽的消化率和饲料的利用率,减少饲料浪费,降低饲养成本,提高养殖效益。
粪肠球菌在反刍动物上的应用研究较少。瘤胃是反刍动物的重要消化器官,幼龄反刍动物的瘤胃发育充分且功能健全是反刍动物发挥高生产性能的前提条件[69]。可见幼龄反刍动物瘤胃早期发育的重要性,而瘤胃微生物群的组成在瘤胃发育中发挥着重要作用[70]。Mamuad等[3]发现,添加肠球菌(E.faecium SROD KCCM11098P)可以调整瘤胃内微生物群落的组成和丰度,增加挥发性脂肪酸浓度。幼龄反刍动物饲喂益生菌,可调节瘤胃pH与胃肠道微生态区系,加速瘤胃发育,实现犊牛提前断奶[70],该发现为后续粪肠球菌同方向应用研究提供依据。在奶牛繁殖阶段,粪肠球菌能显著提高奶牛对饲料中粗蛋白质的消化率,使粪样中粗蛋白质含量降低,并可以抑制产奶量的下降趋势[71]。王灿宇等[32]发现,乳酸粪肠球菌复合菌能够增强锦江黄牛的免疫功能,改善消化道微生物环境。饲喂复合益生菌(酵母菌、乳酸粪肠球菌、芽孢杆菌)后,瘤胃内微生物繁殖旺盛,有机酸浓度和菌体蛋白升高,氨态氮浓度降低[72]。微生态制剂能够有效治疗奶牛瘤胃酸中毒[73-74],维持瘤胃pH的稳定,优化奶牛生产性能,降低经济损失。粪肠球菌和酿酒酵母菌复合物能明显缓解奶牛的亚急性瘤胃酸中毒[75]。肠球菌比牛链球菌的作用更加稳定,其代谢产生的细菌素可抑制瘤胃中产甲烷气体微生物并限制氨基酸的降解[76]。此外,粪肠球菌对奶牛子宫内膜上皮细胞有一定黏附能力,其抗菌活性可作为潜在益生菌,用于开发防治奶牛子宫内膜炎的微生态制剂[30,77]。
粪肠球菌在牛羊养殖领域的研究较少,屎肠球菌在该领域的研究较多,但粪肠球菌和屎肠球菌均属于肠球菌属,有所区别的同时也具有一定程度的相似性,可为粪肠球菌的研究提供指引与研究思路。
粪肠球菌是普遍存在于人和动物消化道内的活性菌群。由于不同的生存环境,菌株携带的基因也不完全相同,且菌株之间可通过重组或水平基因转移实现基因组多样性和基因组可塑性,有利于其适应不同的环境[78-79]。在Shehata等[80]的研究中粪肠球菌(E.faecalis-1)可促进肉鸡生长性能和免疫状态,对调节盲肠菌群有益,而Aalaei等[81]却得出其作用效果并不显著,但Shehata等[80]与Ateya等[81]一致的观点是,认为粪肠球菌能降低由致病性大肠杆菌导致的家禽死亡率,而Tarabees等[48]得出粪肠球菌对其死亡率影响不显著的结论。这些结果不一致可能是由于鸡的品种、饲料组成、饲养条件、添加剂的剂量和类型以及给药途径不同所致[80]。粪肠球菌FQ15能够抑制大肠杆菌O139和金黄色葡萄球菌生长繁殖,分别混合培养48 h、36 h后可将致病菌彻底消除,表现出很强的抑制和杀灭作用[82],而王锂韫等[83]从不同酸马奶酒样中分离的12株粪肠球菌对大肠杆菌ATCC10536和金黄色葡萄球菌B-250均没有抑制效果。不同粪肠球菌菌株对有害菌的抑制作用存在差异,可能是由于菌株来源、生理特性以及发酵液的浓度等不同而导致研究结果的抑菌作用强弱不同[9]。此外,粪肠球菌的菌株差异性还体现在其对药物的耐受性,不同地区的粪肠球菌往往表现出不同的耐药性。由于菌株差异性问题,不同地区、不同来源的粪肠球菌在分离鉴定后,应先对该菌的生物特性进行试验性研究,确认其特性后再进行其他方面的研究。
粪肠球菌普遍存在于消化道内,被认为是正常菌群。然而越来越多的研究发现,其在不断地进化中出现毒力基因[84]。某些粪肠球菌是条件性致病菌,能黏附并侵袭宿主肠上皮细胞,引起肠炎和中毒性疾病[85]。在特定情况下,某些粪肠球菌可穿过肠黏膜屏障,引起感染,例如菌血症、腹膜炎和心内膜炎[86-87],还可导致猪肠道炎症[23],牛乳腺炎[88],鸡败血症,实质器官肿大出血[89]。因此,在使用粪肠球菌作为益生菌添加到畜禽饲粮之前,应先对菌株进行筛选。药敏试验是筛选菌株的重要手段之一,通过药敏试验,可以清晰了解粪肠球菌对各类抗生素的敏感性[90],从而避免使用携带耐药基因的菌株,防止耐药性在畜禽群体以及养殖环境中的传播扩散,保障后续治疗畜禽疾病时抗生素的有效性。此外,利用分子标记技术,可以对不同来源的粪肠球菌菌株进行准确区分、鉴定和追踪,进一步确保其在畜禽体内的安全性和稳定性。
毒力岛又称为致病岛,在粪肠球菌感染中起重要作用,是一段主要与细菌毒力和代谢功能相关的DNA片段。粪肠球菌主要包含的毒力因子有溶血素(Cyl)、聚集物质(AS)、表面蛋白(Esp)、胶原结合蛋白(Ace)、心内膜炎抗原(EfaA)、明胶酶(Gel)和信息素等[87]。其中Cyl可以使革兰阳性菌、真核生物细胞和红细胞的胞浆膜出现小孔,使其破裂发生溶血;AS有利于菌株黏附于真核细胞表面的受体;Esp基因参与细胞壁的合成,促进细菌黏附;粪肠球菌在感染宿主细胞时,Ace辅助菌株黏附在宿主的细胞表面,完成定植;EfaA基因控制粪肠球菌表面黏附蛋白的表达;Gel基因参与粪肠球菌生物膜的形成,加速细菌的繁殖与扩散;信息素促进同属或不同属菌株之间质粒的接合转移[91-92]。由于粪肠球菌存在菌株差异,不同菌株携带的毒力因子不同,可通过基因检测技术精准检测出菌株中是否存在与致病性相关的基因,对菌株进行筛选。此外,在菌株投入应用之前,可将分离的菌株接种到动物模型中,观察动物的感染情况、疾病严重程度等,评估菌株的致病性和风险程度[93]。
耐抗生素肠球菌近年来有所上升,抗生素敏感性是评估肠球菌的重要因素[11]。近几年,粪肠球菌感染率的上升不仅是因为其包含的毒力因子存在致病性,其耐药性的增强也是重要因素之一,对头孢菌素等药物存在天然耐药性,后天也可获得新的耐药性[94]。粪肠球菌的耐药性除了因为在早期抗生素的使用中逐渐进化,该菌还可以通过转座子和质粒转移等方式在种内和种间进行抗生素耐药性传播[86, 92],且因为菌株差异性,不同地区致病菌携带的耐药因子不同,给当地的感染治疗带来一定困难。王东等[95]对在新疆阿克苏地区猪场分离的粪肠球菌进行耐药性测试,发现其中有322株菌对红霉素、泰乐菌素等五种抗生素耐药率达98%以上,部分菌株对庆大霉素已完全耐药。甘富斌等[96]在西藏林芝市某牧场的耗牛鼻拭子中分离到的粪肠球菌对多西环素、万古霉素等7种药物耐药。粪肠球菌多重耐药性的增加已对公共卫生安全造成威胁,尽管引入了几种新的抗生素和抗生素佐剂,但仍面临治疗挑战[97]。
粪肠球菌作为一种极具潜力的益生菌,在饲料添加剂领域有望发挥积极作用。该菌对环境的高抗性,使其能够在各种动物的消化道环境中良好生存,并且对宿主有诸多益处,在畜牧业中展现出广阔的应用前景。在畜牧业中,乳酸菌作为益生菌的组成部分,对畜禽的生长发育、健康维护等方面发挥着广泛且积极的作用,而粪肠球菌在其中占据着举足轻重的地位,为畜牧业的高效发展提供了有力支撑。然而,随着抗生素在畜牧业长期且大量的使用,粪肠球菌的致病性和耐药性日益严峻,这一现状促使科研人员和养殖从业者在使用该菌时必须秉持审慎态度。粪肠球菌的环境抗性较强,有多重耐药性且其耐药性可随着抗生素的持续使用而不断强化。一旦畜禽感染耐药性粪肠球菌后,常规治疗手段较难取得理想效果。因此,在将粪肠球菌应用到实际生产之前,需先对其进行全面的安全性评估,确保安全后方可投入使用。同时也要针对菌株开展药敏试验,以便在其引起畜禽疾病时,能迅速采取有效的治疗措施,保障生物安全,减少经济损失。
尽管面临上述挑战,粪肠球菌在畜牧业中的价值依然不可忽视。近年来,后生元被定义为对宿主健康有益、遗传背景明确的灭活微生物和/或菌体成分,多数研究认为它们须对宿主提供某种生理益处,并能提供类似于活性益生菌的作用[98]。鉴于此,以现有的后生元研究理论为依据,深入开展针对粪肠球菌后生元的研究,有望从根本上解决粪肠球菌在实际应用中的安全性难题,为其在畜牧业的广泛、安全、高效应用开辟新的路径,推动畜牧产业朝着更加绿色、健康、可持续的方向发展。
本文综述了粪肠球菌的生物特性、作用机制、在畜牧养殖中的应用以及目前存在的问题。粪肠球菌对各种环境的适应能力较强,有较广的温度、pH耐受范围以及良好的耐胆盐等特性,且具备优良的黏附性和抑菌性。粪肠球菌的多种作用机制能调节畜禽肠道微生态平衡,促进畜禽的健康生长,有效增加养殖场的经济效益。粪肠球菌在猪、家禽、牛羊等不同种类动物的养殖过程中均展现出提高生产性能、改善肠道环境、增强机体免疫力等诸多优势。然而,目前粪肠球菌也面临着一些问题与挑战,菌株差异使粪肠球菌的应用复杂化,某些菌株的潜在致病性、毒力因子以及日益严重的耐药性问题,威胁动物健康的同时,也对公共卫生安全构成了潜在风险,这在一定程度上限制了粪肠球菌在畜牧业更为广泛和安全有效的应用,未来还需进一步研究以解决这些问题。
  • 国家重点研发计划项目(2022YFA1304201)
  • 国家自然科学基金(32222081)
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2026年第57卷第1期
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doi: 10.11843/j.issn.0366-6964.2026.01.004
  • 接收时间:2024-12-30
  • 首发时间:2026-09-16
  • 出版时间:2026-01-23
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  • 收稿日期:2024-12-30
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国家重点研发计划项目(2022YFA1304201)
国家自然科学基金(32222081)
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    1.中国农业科学院饲料研究所 农业农村部饲料生物技术重点实验室 奶牛营养学北京市重点实验室,北京 100081
    2.北京农学院动物科学技术学院,北京 102206

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毕研亮,主要从事幼龄反刍动物营养与调控研究,E-mail:
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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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