副干酪乳杆菌
副干酪乳杆菌 | |
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科学分类 | |
域: | 细菌域 Bacteria |
门: | 芽孢杆菌门 Bacillota |
纲: | 芽孢杆菌纲 Bacilli |
目: | 乳杆菌目 Lactobacillales |
科: | 乳杆菌科 Lactobacillaceae |
属: | 乳杆菌属 Lactobacillus |
种: | 副干酪乳杆菌 L. paracasei
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二名法 | |
Lactobacillus paracasei Collins, Phillips & Zanoni, 1989
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副干酪乳杆菌(学名:Lactobacillus paracasei),常以其缩写称呼为LP菌,是一种革兰氏阳性菌,乳杆菌属中的一菌种。 为常见的益生菌。副干酪乳杆菌以偏利共生形式运作。常见于与人类相关的栖息地,包括人体内的肠道、口腔,以至人体外的粪渠、青贮饲料及前述的奶制品[1]。正如“乳杆菌”这名字所指,本菌种有著与芽孢杆菌属相似的杆状形态,宽约2.0到4.0μm,长约0.8到1.0μm。
1989年,透过DNA-DNA分子交杂法的结果,科学家将NCDO 151(=ATCC 25302)菌株从干酪乳杆菌分离出来[2]。 现时,科学家从不同的环境中分离出34种不同的菌株,当中有16种是从奶制品分离出来,10种从植物分离出来,8种从人类或其他动物的消化道分离出来[3]。副干酪乳杆菌在进入体外胃肠道运转(GIT)试验前,如经过长时间的冷藏,对于菌株的细胞耐受力并不会造成影响。[4]本物种无论是从基因型或从表型都难以从其他诸如干酪乳杆菌或鼠李糖乳杆菌(Lactobacillus rhamnosus)等其他物种分辨过来[5]。然而,科学家还是可以透过这些菌的发酵谱来将之分辨开来[6]。本菌种专门用于生物食品处理程序,以及用于制作营养补充剂,特别是帮助有消化道不适的病患[7]。
尽管益生菌被认为是安全的,但它们可能引起细菌与宿主的相互作用以及对健康的不利影响。在某些情况下,使用益生菌可能会导致菌血症[8][9]。现时尚未确定益生菌疗法的益生菌菌株、使用频率、剂量和持续时间之间的关系[8]。
生理学
副干酪乳杆菌在生理学属于革兰氏阳性菌,兼性异发酵,非芽孢形成的微生物[10]。副干酪乳杆菌的细胞通常为杆棒状,大小范围为宽度2.0μm至4.0μm,长度0.8至1.0μm[5]。这种生物体是不运动的。副干酪乳杆菌细胞通常具有方形末端,并且可以单一形式或链状存在[5]。
副干酪乳杆菌的最佳生长温度范围是 在10至37°C[11]。当温度超过40°C,乳杆菌就不会再生长。然而,即使在最高温度72°C下,乳杆菌仍然可存活约40秒[5]。由于副干酪乳杆菌是兼性异发酵的:大多数菌株会产生乳酸。
副干酪乳杆菌是正常人类肠道菌群的一部分[12][13]。副干酪乳杆菌无论是在人类胃肠道或一般的植物,均为其常见的栖所[10]。天然发酵的蔬菜、牛奶和肉也类可能含有副干酪乳杆菌[14]副干酪乳杆菌在冷藏(4 °C)下储存,明显比储存在非冷藏环境下(22°C)有较高的存活能力。[15]将益菌冷冻保存在摄氏零下 -20 度C和 -70 度C下,对益菌生存力的不利影响比储存在摄氏 7 度C下要少得多。[16]副干酪乳杆菌8700:2菌株已从健康的人的胃肠道黏膜和粪便中分离出来[12]。8700:2菌株可分解果寡糖和菊粉,而菌株同时可在两者上快速生长并产生乳酸作为最终产物[13]。
系统发生学
副干酪乳杆菌属于细菌界,它是厚壁菌门芽孢杆菌纲乳杆菌目乳杆菌科的物种。[5][17]关于副干酪乳杆菌(L. paracasei)与干酪乳杆菌(L. casei)的命名争论非常激烈,因为资料库中这两种菌株许多有用的序列资料被错误标记。[3]1989年,有人提议将副干酪乳杆菌指定为一个亚种(paracasei),以表明两个物种为DNA同源。[5]如此代表它们的名字在科学文献中可以互换使用。[3]16S 核糖体RNA序列同源性证实了这些物种之间的相关性,[5]但核心基因组系统发育证实了几个密切相关的物种Lc. casei、Lc. paracasei、Lc. rhamnosus(鼠李糖乳杆菌)与Lc. zeae(玉米乳杆菌)其实是不同的物种。[17][18]
从历史上看,副干酪乳杆菌与其他乳酸杆菌之间的差异是基于生化特性。干酪乳杆菌、副干酪乳杆菌和鼠李糖乳杆菌之间有大约90%的序列同一性。[3]但是,有一些区分标准通常用于区分它们。这些差异标准包括营养需求和生长环境。[3]已发现副干酪乳杆菌与其他乳杆菌有特定差异,因为它具有一定的耐热性,在成熟的奶酪中生长良好,并且具有高蛋白酶解活性。[12]
基因组学
副干酪乳杆菌的基因组包含环状DNA,并且在分离的不同菌株之间略有不同。平均而言,基因组有2.9至3.0百万个碱基对(通常缩写为Mb)。它的GC含量在46.2%到46.6%之间,预计编码大约2800到3100个蛋白质。[19]这些菌株的基因组差异在于不同的细胞外膜、分泌蛋白和多糖。许多常见的编码蛋白质是细胞表面相关的细胞壁水解酶,可保护细胞免于凋亡。这些酶已被证明可为人类上皮组织细胞提供细胞保护。[3]
使用多位点序列分型(MLST)和扩增片段长度多态性(AFLP)可评估不同副干酪乳杆菌基因组的遗传多样性。MLST是一种通过使用生物体必需基因的DNA片段对微生物进行分类的技术。[20]AFLP是一种聚合酶连锁反应(PCR)工具,用于DNA分析,使用限制酶和配体扩增所需的DNA片段。[21]
医学和临床研究
过敏性呼吸道疾病(过敏性鼻炎、结膜炎与气喘)
副干酪乳杆菌LP-33菌株治疗过敏性鼻炎提供了有益的临床和免疫学作用的重要证据。[22][23]
副干酪乳杆菌BRAP01菌株 是诱导台湾个体产生γ-干扰素 (IFN-γ)/介白素-10(IL-10) 的主要菌株。 [24]
副干酪乳杆菌 HB89菌株 可减轻“细悬浮微粒( PM 2.5)”刺激的呼吸道过敏。 [25]
对于过敏性鼻炎(AR),每日口服100亿以上菌数(1x1010CFU)的副干酪乳杆菌(食用前均保存于4°C中)并持续食用8周的时间,可改善鼻搔痒(AR的关键临床特征)并减少促发炎介质 IL-5 分泌。 [26]
异位性皮肤炎、荨麻疹
口服副干酪乳杆菌KBL382菌株可显著减少与 异位性皮肤炎(AD) 相关的皮肤损伤、表皮增厚、免疫球蛋白 E 的血清水平和免疫细胞浸润。 [27]
流行性感冒
口服热灭活的 副干酪乳杆菌MoLac-1菌株增加了脾脏中 NK 细胞的比例,并改善了小鼠 流行性感冒病毒(IFV) 感染的症状。 [28]
普通感冒
副干酪乳杆菌MCC1849菌株 有可能提高易感受试者对普通感冒感染的抵抗力,并保持理想的情绪状态,即使在精神压力条件下也是如此。 [29]
新型冠状病毒感染
副干酪乳杆菌DG 菌株显著诱导了参与保护性抗病毒免疫的基因表达,并阻止了由 SARS-CoV-2 感染引发的促炎基因的表达。 [30]
发炎性肠道疾病
包含副干酪乳杆菌的活菌制剂可与常规疗法结合使用,以治疗溃疡性结肠炎[31]。 副干酪乳杆菌的D3-5菌株经过“热杀”(heat killed)后,其细胞壁会析出脂磷壁酸,可改善实验鼠与衰老相关的肠道渗出、炎症,并改善其身体和认知功能[32]。
龋齿
除了各种和消化道相关的病变,2011年,副干酪乳杆菌首次用于预防龋齿。这是因为副干酪乳杆菌能够识别变形链球菌,使其附著并导致结块,而变形链球菌因此无法再附著在牙齿上,而会被唾液冲洗掉或在刷牙时被除去[33]。
腹泻
副干酪乳杆菌已被证明可抑制大肠杆菌的细菌活性,而大肠杆菌是引致腹泻的常见菌种,因此副干酪乳杆菌被应用于治疗腹泻[34]。包含副干酪乳杆菌的活菌制剂可与常规疗法结合使用,以治疗溃疡性结肠炎[31]。一项系统综述为副干酪乳杆菌LP-33菌株治疗变应性鼻炎提供了有益的临床和免疫学作用的重要证据。[22]
幽门螺旋杆菌感染
副干酪乳杆菌显示出对幽门螺旋杆菌的抑菌和杀菌活性。 [35]
大肠激躁症
副干酪乳杆菌可减轻胃肠道症状的严重程度,并改善某些大肠激躁症(IBS) 亚型个体的心理健康。 [36]
癌症
胃癌和结肠癌细胞的凋亡和生长易受已热灭活和有活力的IMPC2.1菌株的影响[1]。
健康问题
肠道微生物群的操纵很复杂,可能会导致细菌与宿主的相互作用。尽管益生菌被认为是安全的,但当它们通过口服用药时,存在活菌从胃肠道进入内脏(细菌易位)和随后菌血症的风险,这可能导致不利的健康后果。[8]有些人,如免疫功能受损、短肠综合症、中心静脉导管、心脏瓣膜疾病和早产儿,可能面临更高的不良事件风险。[9]
目前,益生菌治疗的益生菌菌株、频率、剂量和持续时间尚未确定。[8]活菌可能不是必需的,因为益生菌的有益作用似乎是由它们的DNA和分泌的可溶性因子介导的,其治疗效果可以通过全身给药而不是口服给药来获得。[8][37]
历史
乳酸菌(Lactic Acid Bacteria;一般缩写作LAB)是在1900年代初进行分类和分组的。它们受到注意,主要是由于科学家观察到这些细菌在不同食品(尤其是乳制品)中的相互作用。1991年,荷兰微生物学家马丁努斯·威廉·拜耶林克从之前已知的LAB组中分离出了革兰氏阳性细菌,就是乳酸杆菌[38]。 副干酪乳杆菌最近已被归类为干酪乳杆菌益生菌群组的成员[1]。但是,在生物分类学上,其具体位置并不明确。1996年,Dicks, Duplessis, Dellaglio & Lauer提出副干酪乳杆菌 并不是一个独立的物种[5]。
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