微生物学报

Function	Gene	Comment
Bacteriocin leader domain-containing protein	breB	Bacteriocin synthesis gene
breC
Immune factors encoding bacteriocins	breE
Catalytic decarboxylation of L-glutamic acid to produce gamma-aminobutyric acid	gadA	Glutamic acid decarboxylase (GAD) encoding gene
gadB
Glutamate/gamma-aminobutyrate antiporter	gadC	Accessory gene
Surface layer protein SlpB	slpB	S-layer protein
Surface layer protein SlpC	slpC
Surface layer protein SlpD	slpD
Encodes a family of membrane-bound glycosyltransferases	gtf27	Membrane bound-glycosyltransferase family
gtf28
Responsible for the transport of extracellular polysaccharides	orf29	Transporter gene
Rhodanese-related sulfurtransferase	TstT	Participation in cyanide detoxification processes
Transcriptional regulator	TstR
Encodes an organophosphorus hydrolase	OpdB	The member of the GDSVG family of esterolytic enzymes

Function	Gene	Comment
Bacteriocin leader domain-containing protein	breB	Bacteriocin synthesis gene
breC
Immune factors encoding bacteriocins	breE
Catalytic decarboxylation of L-glutamic acid to produce gamma-aminobutyric acid	gadA	Glutamic acid decarboxylase (GAD) encoding gene
gadB
Glutamate/gamma-aminobutyrate antiporter	gadC	Accessory gene
Surface layer protein SlpB	slpB	S-layer protein
Surface layer protein SlpC	slpC
Surface layer protein SlpD	slpD
Encodes a family of membrane-bound glycosyltransferases	gtf27	Membrane bound-glycosyltransferase family
gtf28
Responsible for the transport of extracellular polysaccharides	orf29	Transporter gene
Rhodanese-related sulfurtransferase	TstT	Participation in cyanide detoxification processes
Transcriptional regulator	TstR
Encodes an organophosphorus hydrolase	OpdB	The member of the GDSVG family of esterolytic enzymes

短促生乳杆菌基因组学研究进展

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吕慧敏 ²^,³^,⁴ , 李伟程 ¹^,²^,³^,⁴ , 张和平 ¹^,²^,³^,⁴^,^*

微生物学报 | 综述 2024,64(9): 3157-3167

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微生物学报 | 综述 2024, 64(9): 3157-3167

短促生乳杆菌基因组学研究进展

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吕慧敏²^,³^,⁴, 李伟程¹^,²^,³^,⁴, 张和平¹^,²^,³^,⁴^,^*

作者信息

1 乳酸菌与发酵乳制品省部共建协同创新中心, 内蒙古呼和浩特 010018

2 内蒙古农业大学, 乳品生物技术与工程教育部重点实验室, 内蒙古呼和浩特 010018

3 农业农村部奶制品加工重点实验室, 内蒙古呼和浩特 010018

4 内蒙古自治区乳品生物技术与工程重点实验室, 内蒙古呼和浩特 010018

Research progress in genomics of Levilactobacillus brevis

Huimin LYU²^,³^,⁴, Weicheng LI¹^,²^,³^,⁴, Heping ZHANG¹^,²^,³^,⁴^,^*

Affiliations

1 Collaborative Innovative Center of Ministry of Education for Lactic Acid Bacteria and Fermented Dairy Products, Hohhot 010018, Inner Mongolia, China

2 Key Laboratory of Dairy Biotechnology and Engineering, Ministry of Education, Inner Mongolia Agricultural University, Hohhot 010018, Inner Mongolia, China

3 Key Laboratory of Dairy Products Processing, Ministry of Agriculture and Rural Affairs, Hohhot 010018, Inner Mongolia, China

4 Inner Mongolia Key Laboratory of Dairy Biotechnology and Engineering, Hohhot 010018, Inner Mongolia, China

出版时间: 2024-05-10 doi: 10.13343/j.cnki.wsxb.20240118

文章导航

摘要

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短促生乳杆菌(Levilactobacillus brevis)是常见乳酸菌物种，主要存在于植物茎叶表面、泡菜、乳制品以及肠道等生态位。L. brevis具有优良的生理功能，是潜在的益生菌物种。随着基因组学研究兴起，从基因水平揭示L. brevis的遗传学特征及功能基因特性对该菌的应用具有重要意义。本文主要围绕L. brevis的遗传背景及重要功能基因进行综述，以期为L. brevis的应用研究奠定理论基础。

关键词

短促生乳杆菌 / 基因组学 / 功能基因

Abstract

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Levilactobacillus brevis is a common species of lactic acid bacteria mainly detected on the surface of plant stems and leaves and in pickles, dairy products, and intestines. With excellent physiological functions, L. brevis is a potential probiotic species. With the rise of genomics, it is of great significance to reveal the genetic characteristics and functional gene properties of L. brevis at the gene level for application of this bacterium. This paper reviews the genetic background and major functional genes of L. brevis, aiming to lay a theoretical foundation for the application of L. brevis.

Key words

Levilactobacillus brevis / genomics / functional gene

引用本文

吕慧敏, 李伟程, 张和平. 短促生乳杆菌基因组学研究进展. 微生物学报, 2024 , 64 (9) : 3157 -3167 . DOI: 10.13343/j.cnki.wsxb.20240118

Huimin LYU, Weicheng LI, Heping ZHANG. Research progress in genomics of Levilactobacillus brevis[J]. Acta Microbiologica Sinica, 2024 , 64 (9) : 3157 -3167 . DOI: 10.13343/j.cnki.wsxb.20240118

正文

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短促生乳杆菌(Levilactobacillus brevis)为乳杆菌科革兰氏染色阳性菌，具有专性异型发酵的生物学特性^[1]，最适生长温度29.5−30.0 ℃^[2]。其可以在较宽的pH范围内存活(pH 4.0−8.0)，尤其能够耐受pH为2.0的外界环境^[3]。L. brevis的分离源十分广泛，在植物茎叶表面、泡菜、乳制品以及肠道等多种生态位存在，其可以用作发酵剂改善产品的风味和质地、提升产品的营养价值，并抑制腐败菌及病原体的生长，延长产品货架期^{[2, 4]}。益生菌作为对宿主有益的微生物，能在肠道中定殖并发挥其益生功能是至关重要的先决条件。研究表明L. brevis可在胆盐环境下存活，并抑制大肠杆菌、葡萄球菌和芽孢杆菌等常见肠道病原菌，并且无溶血活性和耐药性，具有作为益生菌的潜力^[5-7]。L. brevis具有高产γ-氨基丁酸(γ-aminobutyric acid, GABA)^[8]、合成乳蛋白活性肽^[9-10]等生物学特性，使其在医药行业具有良好的发展前景^[11]。2008年，本研究团队刘文俊等^[12]完成了国内首株乳酸菌Lacticaseibacillus casei Zhang的全基因组测序工作，开启了国内益生乳酸菌基因组学研究的序幕；近年来，还建成了全球最大的乳酸菌基因组数据库(https://www.imhpc.com/iLABdb)，为乳酸菌研究领域提供了宝贵的资源和平台，而对于L. brevis的基因组学研究较为薄弱。本文主要综述了L. brevis的基因组学研究进展，以期揭示L. brevis的遗传背景和重要功能基因，为优良特性L. brevis的挖掘与利用提供理论基础。

1 短促生乳杆菌的遗传背景

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二代高通量测序技术的不断发展，为基因组学的研究提供了大量数据。1999年，第一株乳酸菌Lactococcus lactis subsp. lactis IL1403基因组草图测序与组装工作的完成标志着乳酸菌研究进入基因组时代^[13]。起初基因组学主要应用在乳酸菌分类地位^[14]、发酵能力以及益生特性^[15]等方面。如今，研究者们通过全基因组测序技术，获取菌株的全部遗传物质信息以及编码基因的功能分类，通过比较种内或种间不同菌株的基因组，分析其遗传进化关系，并挖掘重要功能基因。2006年首株L. brevis ATCC367^T全基因组测序完成^[16]，标志着L. brevis的研究开始深入到分子水平。截至2024年1月，共有159株L. brevis完成基因组测序，其中完成图有29株(https://www.ncbi.nlm.nih.gov/datasets/genome/?taxon=1580)，基因组大小2.34−2.91 Mb，G+C含量45.2%−46.4%。L. brevis具有基因组较小、G+C含量较低的特点^[17]，这为其在基因组水平上分析提供了便利条件。

Feyereisen等^[18]筛选出6株来源于啤酒、啤酒酿造环境以及青贮饲料的L. brevis，与NCBI公共数据库中筛选的13株不同分离源L. brevis进行比较基因组学分析发现，啤酒源的L. brevis菌株基因组中的基因数目多于其他分离源菌株，说明啤酒源菌株适应啤酒环境获得了新的功能基因；对这些啤酒特异性的“获得性”基因序列进行同源性分析表明，大约25%的基因编码氧化还原反应相关蛋白、22%编码转录相关蛋白、21%编码膜和细胞表面蛋白、14%编码膜转运相关蛋白，说明L. brevis在啤酒酿造过程中发生了适应性进化。Fraunhofer等^[19]通过比较基因组学研究24株不同分离源的L. brevis发现，啤酒及酿造环境分离株与昆虫分离株在质粒组内存在共享基因库，而且共享基因库中的horC基因是经过早期研究证实在啤酒生产中具有啤酒花抗性的基因，该研究首次在非啤酒酿造环境分离株中发现horC基因，推测啤酒酿造环境中的horC基因是通过水平基因转移而获得。Panahi等^[20]对83株L. brevis的CRISPR-Cas多样性进行分析发现，127个已确认的CRISPR以及31个Cas基因分布在69株L. brevis中，这些基因被归类为Ⅱ-A、Ⅱ-C和Ⅰ-E亚型，其中Ⅱ-A亚型是L. brevis中对抗外源DNA及噬菌体最活跃的系统。

综上所述，L. brevis在基因组水平上具有很高的相似性，但因菌株的生活环境不同，导致不同菌株发生了环境适应性进化，并拥有了各自独特的功能基因。通过了解这种基因型和表型之间的相互联系，能够进一步探究不同菌株的水平基因转移和生理功能特性。目前，针对除啤酒酿造环境外的其他分离源L. brevis遗传多样性及适应性进化机制研究较少，因此从更多、更广的角度分析不同分离源L. brevis的基因组特征、遗传进化规律及功能基因差异具有重要意义。

2 短促生乳杆菌的重要功能基因

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L. brevis已被美国食品药物监督管理局(Food and Drug Administration, FDA)和美国饲料公定协会(Association of American Feed Control Officials, AAFCO)评价为可直接饲喂且通常认为是安全的微生物(generally recognized as safe, GRAS)，被欧洲食品安全局(European Food Safety Authority, EFSA)授予合格安全资格认定(qualified presumption of safety, QPS)地位^[21-22]。L. brevis作为潜在的益生菌在乳制品、医药等行业中具有良好的发展前景。利用L. brevis的基因组信息及相关生物信息学分析该菌基因型与表型的相互联系，对L. brevis的工业化应用至关重要，有关L. brevis功能基因及应用研究备受研究者们青睐。研究发现，L. brevis具有高产γ-氨基丁酸、抑菌以及降低有害物质含量的能力^{[15, 23-24]}。表1详细列出了本文中提及的与L. brevis相关的功能基因，这些基因对于确保L. brevis菌株的生长、代谢以及环境适应性起到不可或缺的作用。通过筛选优良L. brevis菌株，有效挖掘和调控关键功能基因，进而促进其产生更多有益代谢产物。这不仅有助于提高L. brevis菌株在生物体内的应用潜力，也成为未来L. brevis在食品、饲料以及医药等领域得到广泛应用的重要途径。

2.1 细菌素合成基因簇

细菌素是由细菌核糖体合成的一类具有抗菌特性的活性多肽，能够抑制除自身以外的多种微生物。由于抗生素耐药性及一些化学合成类食品添加剂的潜在不良作用，乳酸菌细菌素所具有的独特属性使其在作为天然食品防腐剂用以替代化学合成防腐剂和抗生素药物方面拥有巨大潜力^[25-26]。L. brevis产生的细菌素属于Ⅱ类细菌素，具有分布广泛和热稳定的特点，目前仅发现这类细菌素具有抗李斯特菌(Listeria)的作用^[27]。

2009年，Wada等^[28]从韩国传统泡菜中分离出L. brevis 925A，并分离得到了一种细菌素brevicin 925A，基因组研究发现该细菌素合成基因(breB和breC)和免疫基因(breE)位于该菌中碱基数最多的pLB925A04质粒上。2015年，Noda等^[29]从柑橘中分离出L. brevis 174A，并发现了Ⅱ b型细菌素brevicin 174A合成基因位于质粒上的由8个开放阅读框(open reading frame, ORF)组成的基因簇中，该基因簇主要包括生物合成基因、自身细菌素抗性基因和转录调控蛋白基因。2018年，Noda等^[30]进行后续研究发现L. brevis 174A细菌素合成基因簇上存在2个假定调控基因(breD和breG)，它们编码转录调节蛋白并正向调控brevicin 174A的合成，继而研究表明brevicin 174A不仅抑制与L. brevis近缘的乳酸菌生长，还能抑制金黄色葡萄球菌(Staphylococcus aureus)、单核增生李斯特菌(Listeria monocytogenes)和沙门氏菌属(Salmonella)等致病菌的生长^[31]。

L. brevis中细菌素基因簇的发现，有助于在分子水平上进一步了解L. brevis产生细菌素的功能特性，同时也为L. brevis细菌素在食品防腐剂领域的应用以及作为替代抗生素药物的研究提供了相关理论依据。然而，目前关于L. brevis细菌素的研究相对较少，需要更多的深入研究来探索其潜在的应用价值。

2.2 γ-氨基丁酸合成基因

γ-氨基丁酸(gamma-aminobutyric acid, GABA)是哺乳动物中枢神经系统中主要的抑制性神经递质^[32]。在生产GABA的微生物中，乳酸菌被认为是一类较为安全的微生物。目前已经报道了几种产生GABA的乳杆菌，包括发酵黏液乳杆菌(Limosilactobacillus fermentum)、植物乳植杆菌(Lactiplantibacillus plantarum)、乳酸乳球菌(Lactococcus lactis)、L. brevis等，其中L. brevis在胃肠道中产生的GABA浓度最高^[33]。研究表明食用富含GABA的食物可以降血压^[34-35]、抑制癌细胞增殖^[36]、改善记忆和学习能力^[37]。GABA已被列为食品和药品中的生物活性成分^[38]，并在医药以及食品领域得到广泛应用。

Renes等^[39]从手工奶酪中成功分离出85株乳酸菌，其中10株具有生产GABA的能力(包括6株L. brevis和4株乳酸乳球菌)；该研究检测了产生和不产生GABA的菌株是否存在编码谷氨酸脱羧酶系统的相关基因，研究结果显示，所有能够产GABA的6株L. brevis均检测到含有编码该系统的3个基因；此外，该研究提出谷氨酸脱羧酶系统编码基因的检测或可以作为筛选生产GABA菌株的一种方法，但这一方法仍需在更多实验中进一步验证。Shi等^[40]从泡菜中分离得到一株高产GABA的L. brevis Lb85，并鉴定到2个谷氨酸脱羧酶(glutamic acid decarboxylase, GAD)基因，分别为gadB1和gadB2，及其辅助基因gadC和gadR；gadB1和gadB2基因均具有谷氨酸脱羧酶活性，而且gadB2基因的表达活性优于gadB1基因，而不含gad基因的菌株则不产GABA；此外，当GABA的增长高于40 mg/(L·h)时，表达gadC基因有助于GABA的转运，而当初始葡萄糖浓度达到160 g/L时，表达gadR基因可提高GABA的产量。彭春龙等^[41]从鲜奶中分离出一株具有高产GABA特性的L. brevis CGMCC NO. 1306，并克隆得到gadA和gadB及其辅助基因gadC，他们还探究了酸胁迫对基因表达的影响，结果表明该菌株中gadB与gadC基因位于同一操纵子中，在酸性环境下会促进gadCB表达；此外，当菌株达到对数生长期后，gadCB基因的表达量也会大幅度增加，而培养条件的改变对gadA基因的表达影响较小。Pakdeeto等^[42]在泰国泡菜中分离出16株产GABA的乳酸菌，通过基因组分析发现菌株GPB7-4与L. brevis ATCC367^T亲缘关系较近，基因组平均遗传相似度(average nucleotide identity, ANI)值为99.94%。该菌株含有产GABA的基因gadA和gadB；在16株乳酸菌中L. brevis GPB7-4产GABA的能力最高，因此在发酵食品领域具有良好的应用前景。Gong等^[43]研究证实，L. brevis中GAD系统的gadR基因是控制该菌株GABA转化及耐酸性的正转录调控因子，因此，高表达gadR的L. brevis菌株是工业规模生产GABA的优良候选菌株。本研究团队对本实验室分离得到的145株L. brevis结合NCBI RefSeq数据库已公布的157株L. brevis对gad基因进行分析，结果发现gadB基因存在于所有302株菌中，而gadC基因存在于301株菌中，而且gadB和gadC基因的拷贝数在各菌株中有所差异。

谷氨酸脱羧酶是可以催化L-谷氨酸脱羧产生γ-氨基丁酸唯一的关键限速酶。然而，由于L. brevis菌体本身具有厌氧、生长速度慢、发酵周期长等特性，导致GABA的生产效率较低^[44]。因此，探究L. brevis中谷氨酸脱羧酶基因特征及其高效表达条件，对GABA的高效且大规模生产，以及在食品医药等行业的应用具有重要意义。

2.3 S-层蛋白合成基因

S-层蛋白(S-layer protein, Slps)是一种生物活性大分子，广泛存在于古生菌、革兰氏阳性菌和革兰氏阴性菌细胞壁表面，大多数Slps都由单一蛋白质或糖蛋白组成^[45-46]。Slps在瑞士乳杆菌(Lactobacillus helveticus)、嗜酸乳杆菌(Lactobacillus acidophilus)、L. brevis等13种乳酸杆菌中较为丰富，而不同乳酸菌中Slps基因的总体序列相似性较低，但通常在近缘菌种中较高^[47-48]。

研究表明，乳酸杆菌表达系统外源蛋白表达量与启动子活性有关，L. brevis的slp基因启动子为强启动子，因此Slps蛋白表达量较高，占菌体总蛋白含量的10%−15%；研究发现L. brevis的Slps蛋白信号肽本身由slp基因编码，能够一直表达其目的基因不受其他外源因素影响，有利于构建乳酸菌外源蛋白表达载体^[49]。早期有学者^[50-51]对L. brevis ATCC8287的Slps进行分离、纯化及基因测序，结果表明，该蛋白具有一个ORF，此ORF编码一个蛋白质以及一个典型革兰氏阳性菌信号肽。Jakava-Viljanen等^[52]在L. brevis ATCC14869^T中分离得到2个Slps (SlpB和SlpD蛋白)，测序了3个slp基因(slpB、slpC和slpD)并探究其在菌株中的表达情况，结果表明，在不同生长条件下，ATCC14869^T菌株形成光滑(S)和粗糙(R) 2种不同的菌落类型，在有氧条件下合成SlpB和SlpD蛋白，产生R型菌落，而厌氧条件下只合成SlpB蛋白，产生S型菌落；此外，不同于其他乳杆菌，ATCC14869^T的slpB基因下游还具有基因slpC，该基因在测试的生长条件下不表达，而slpD基因与slpB-slpC基因位于基因组上不同位置，并且通气条件会诱导slpD基因的表达。Banić等^[53]从酸菜中分离出一株L. brevis SF9B，通过全基因组序列发现了3个编码slpB基因，但仅表达了一个与L. brevis ATCC14869^T相似的slpB基因；研究结果发现，在胃肠道(gastrointestinal, GI)条件和冷冻干燥过程中，slpB基因的表达使菌株具有更高的存活率，而且Slps对人肠道细胞株Caco-2细胞的黏附能力更强，去除Slps则完全消除了SF9B细胞的黏附作用。

Slps具有对胆盐及消化酶的耐受能力，参与乳杆菌对宿主细胞的黏附，并抑制病原菌对宿主细胞的黏附及侵入作用。因此，它在疫苗、生物传感器、微载体、口服药物载体的纳米涂层等方面有着巨大的发展潜力^[54-55]。关于L. brevis中Slps的研究，当前主要聚焦于探索其在调节肠道炎症过程中的作用^[56]。然而，对于L. brevis独特的slp基因表达机制还有待于继续研究。

2.4 胞外多糖合成基因

胞外多糖(exopoly saccharides, EPS)是长链、高分子量聚合物，其分为与细胞表面紧密结合的荚膜多糖(capsular polysaccharide, CPS)以及能够释放到菌体周围环境介质中的黏液多糖(slime polysaccharide, SPS)。乳酸菌的胞外多糖具有保护菌体免受环境pH值、渗透压、噬菌体、抗生素等应激条件的影响，促进细胞表面黏附及营养吸收的作用^[57]，其用在食品工业中可以改善产品的风味、质地以及营养价值^[58]。

Fukao等^[59-60]对L. brevis KB290进行测序及基因组分析发现，KB290是EPS的天然生产者，而且KB290的质粒pKB290-1中含有3个EPS基因(gtf27、gtf28和orf29)能够合成EPS，使其具有细胞聚集和胆汁抵抗能力，并推测gtf27、gtf28可能属于膜结合糖基转移酶家族负责生物合成，而orf29则负责EPS的输出。2019年，Fukao等^[61]继续以pKB290-1的EPS基因为研究对象，成功克隆了该质粒上的EPS基因位点(gtf27、gtf28和orf29)并表达在L. brevis以及植物乳植杆菌上，研究发现这3个基因与菌落形成的褶皱形态有关。在酒类或其他饮料中，细菌产生的EPS会使其黏度增加，影响产品的品质。Fraunhofer等^[62]对3株啤酒及其酿造环境分离出的L. brevis进行了全基因组测序，通过分析发现在所有的3株菌中均含有编码EPS合成关键酶——糖基转移酶(Gtf-2)的基因gtf-2；通过对从黏稠变质啤酒中分离出的L. brevis TMW 1.2112分析发现，Gtf-2是L. brevis TMW 1.2112合成β-葡聚糖的必需酶，gtf-2基因位于TMW 1.2112的一个质粒上，其可以作为使啤酒黏稠变质菌检测的标记基因。L. brevis的EPS生产基因是其益生特性的体现之一^[63]，但是对于潜在益生菌L. brevis的EPS基因表达机制及其代谢途径还有待于深入挖掘。

2.5 降解氰化物、有机磷等有害物质基因

L. brevis通常可以在植物和发酵蔬菜中被分离出来^[64]。在豆科、蔷薇科、禾本科、亚麻科和菊科等植物中可能含有氰苷^[65]，氰苷在遇酸或生物酶作用下水解产生有毒的氢氰酸。Pagliai等^[66]在L. brevis ATCC367^T中发现2个基因TstT和TstR，它们各自编码参与氰化物解毒的硫代硫酸盐和转录调节器；该研究还确定了TstT与TstR基因均参与L. brevis的氰化物解毒过程。Obilie等^[67]研究表明，植物乳植杆菌、L. brevis以及肠膜状明串珠菌乳脂亚种(Leuconostoc mesenteroides subsp. cremoris)组合成的微生物群能够使木薯根在加工过程中总氰原含量降低约98%。Cho等^[68]发现，在泡菜发酵过程中，部分乳酸菌如肠膜明串珠菌WCP907、L. brevis WCP902、植物乳植杆菌WCP931和清酒乳杆菌(Lactobacillus sakei) WCP904对有机磷(organophosphorus, OP)杀虫剂毒死蜱(chlorpyrifos, CP)有降解作用。Islam等^[69]从泡菜中成功分离到一株L. brevis WCP902，通过研究发现了其有机磷水解酶编码基因(OpdB)，同时证明了L. brevis WCP902具有降解CP的能力。L. brevis中降低有害物质含量相关基因的发现及表达，对泡菜发酵过程中降低氰原含量及农药残留和提高产品品质及安全性有着重要意义。

随着工业化的快速发展，水、食物及环境污染问题日益严峻，这些不利影响加剧了重金属进入并累积于人类食物链中的风险。研究发现，L. brevis在小肠内对重金属汞有很强的结合能力，能够有效减少肠道炎症并改善肠道损伤^[70-71]，但对于L. brevis中有关汞的抗性基因还有待继续挖掘。此外，L. brevis还具有抑制黄曲霉菌产毒素^[72]和降解青贮饲料中伏马菌素^[73]的作用，其对亚硝酸盐的降解率可达82.84%^[74]。然而，目前针对L. brevis中降解有害物质的基因及其脱毒机理的研究尚处于初级阶段。

3 展望

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L. brevis是一种在工业发酵中具有重要应用价值的菌种，已逐渐被应用于食品、医药等领域。该菌种具有良好的耐酸、耐胆盐及黏附性等特性，具有促进健康及作为益生菌的潜力。然而，也有报道称L. brevis是导致啤酒腐败的病原菌。为实现高效筛选L. brevis优良菌株及菌株安全性评价，利用基因组信息挖掘不同菌株的耐酸、耐胆盐、细菌素操纵子等功能基因特征，探究其遗传进化机制，对L. brevis作为潜在益生菌的发展具有重要意义，为其高效利用提供理论依据。

基金

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国家自然科学基金(U22A20540)

参考文献

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文献

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参考文献引证文献

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2024年第64卷第9期

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doi: 10.13343/j.cnki.wsxb.20240118

接收时间：2024-02-26
首发时间：2026-03-20
出版时间：2024-05-10

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收稿日期：2024-02-26
录用日期：2024-05-06

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National Natural Science Foundation of China(U22A20540)

国家自然科学基金(U22A20540)

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1 乳酸菌与发酵乳制品省部共建协同创新中心, 内蒙古呼和浩特 010018

2 内蒙古农业大学, 乳品生物技术与工程教育部重点实验室, 内蒙古呼和浩特 010018

3 农业农村部奶制品加工重点实验室, 内蒙古呼和浩特 010018

4 内蒙古自治区乳品生物技术与工程重点实验室, 内蒙古呼和浩特 010018

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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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