外源信号分子及肠道菌群对脆弱拟杆菌群体感应的影响
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摘要
研究背景：
细菌之间的交流是通过分泌和感应一种被称为自诱导剂(Autoinducer, AI)的信号分子来实现的。细菌根据这种特定信号分子浓度的变化来监测周围环境中其他细菌数量的变化,这种信号传递的过程称为群体感应(Quorum-sensing, QS)。现有报道的细菌QS环路有三种机制：革兰阳性菌以修饰多肽作为信号调节分子；革兰阴性菌主要以典型的LuxI/LuxR系统通过产生一类信号分子N-酰基-高丝氨酸内酯类(AHL)进行群体感应；另外一种QS分子目前认为是二类信号分子(呋喃硼酸二酯),为革兰阴性菌和阳性菌共有。最近几年,有关需氧菌的QS系统研究方兴未艾,研究范围越来越广。但是,厌氧菌的QS系统类似研究至今还未见报道。
脆弱拟杆菌占人体肠道菌群的比例约为1%-2%,是引起胃肠感染的最主要的厌氧菌,多种因素导致该菌可以在肠道内大量存在并形成生物膜。脆弱拟杆菌生物膜形成是否与QS有关,该菌是否产生一类QS信号分子AHL,以及AHL与肠道菌群对该菌生物学活性的影响,目前相关研究报道极少,因此,本研究重点研究了UPLC-MS(四极杆飞行时间质谱,Q-TOF-MS)检测AHL分子的影响因素,创建了基于UPLC-MS技术的细菌群体感应一类信号分子AHL的检测平台,为系统研究肠道微生态细菌的信号传导提供实验技术和方法。另外采用分子生物学技术探讨脆弱拟杆菌是否存在QS环路系统,同时分析了与脆弱拟杆菌密切相关的其他肠道细菌及外源性AHL对脆弱拟杆菌QS相关基因表达的影响,为进一步研究其群体感应发生机制奠定基础。
实验一N-酰基-高丝氨酸内酯类群体感应信号分子检测方法的建立
方法：采用UPLC/MS (Q-TOF-MS)代谢组学分析方法检测6种N-酰基-高丝氨酸内酯标准品,重点考察UPLC-MS检测AHL分子的各种参数,包括溶剂的选择、流动相的优化、滤膜孔径、质谱的电喷雾离子化模式、色谱柱的选择、质谱条件的优化、AHL提取分离的前处理条件、AHL分子在质谱中的离子化特点等。
结果：创建了基于UPLC-MS技术的细菌群体感应系统一类信号分子AHL的检测平台,其检测参数为：正离子电喷雾离子化模式、Aquity C181.7μm2.1mm×100mm色谱柱,乙腈作为AHL溶剂及流动相。AHL内酯环结构在质谱鉴定过程中不会被破坏,在电喷雾电离四极杆飞行时间质谱中总保持m/z为102.05和74.05的特征峰组合(N-已酰-DL-酰基高丝氨酸内酯只见102.05特征峰),三氯甲烷甲醇水混合液作为萃取剂提取细菌培养液中AHL的灵敏度为5ng/ml。
实验二基于UPLC-MS技术检测脆弱拟杆菌群体感应一类信号分子AHL
方法：采用PCR方法扩增实验菌株(包括脆弱拟杆菌、5种双歧杆菌、大肠埃希菌、屎肠球菌、金黄色葡萄球菌、副溶血弧菌、阴沟肠杆菌、鲍曼不动杆菌、伤寒沙门菌)的16SrRNA基因并测序,确保实验菌株鉴定的准确性；在确定UPLC-MS (Q-TOF-MS)检测AHL的各种检测参数后,采用该方法对脆弱拟杆菌和其他多种细菌的菌液提取物中是否存在AHL进行了检测,并以AHL标准品做对照实验。
结果：实验菌株16SrRNA基因序列与GenBank数据库序列同源性大于98%,采用UPLC-MS:法检测溶于不同细菌培养液中AHL标准品,其电喷雾电离四极杆飞行时间质谱均见m/z为102.05特征离子碎片或102.05和74.05特征碎片组合(N-已酰-DL-酰基高丝氨酸内酯只见102.05特征峰),而脆弱拟杆菌和其他多种细菌的菌液均未检测到AHL。
实验三脆弱拟杆菌生物学特性的研究
方法：采用甘油厌氧肉汤及甘油普通肉汤对脆弱拟杆菌进行保存,观察在不同时间及不同温度保存条件下的存活状况,同时观察反复冻融10次对其存活力的影响。另将脆弱拟杆菌菌液分别暴露在空气中5min、10min、30min、60min、90min、120min、150min、180min、24h,观察不同时间段脆弱拟杆菌的的生存力及生存率的变化。
结果：接种于甘油厌氧肉汤及普通肉汤中的脆弱拟杆菌在-20℃及-80℃两种温度下保存3个月后,反复冻融10次后均能培养成功,与空气接触180min时仍生存,24h后即死亡,与空气接触5min,10min,30min,60min后生存率分别为84.1%,68.1%,46.4%,34.8%。
实验四外源信号分子及肠道菌群对脆弱拟杆菌生物学活性的影响
方法：将5种肠道常见菌的培养上清液及2种AHL标准品加入厌氧血培养液中,用于脆弱拟杆菌培养,检测不同培养时间段的脆弱拟杆菌菌液OD值,分析其在不同培养环境的生长曲线变化。
结果：脆弱拟杆菌在厌氧血培养液、含十二烷基高丝氨酸内酯、大肠杆菌、金黄色葡萄球菌、屎肠球菌上清液、酮乙酰基高丝氨酸内酯的厌氧血培养中培养时,其进入对数期的时间分别为8h、6h、6h、6h、10h、10h,其进入稳定期时间分别为10h、8h、8h、8h、24h、24h,脆弱拟杆菌在含其他细菌上清液的厌氧血培养液中进入对数期及稳定期的时间与空白厌氧血培养液一致。
实验五荧光定量PCR法检测不同培养环境脆弱拟杆菌群体感应相关基因的表达
方法：采用PCR方法检测群体感应相关基因luxl及脆弱拟杆菌的luxR基因及其亚型,实时荧光定量PCR方法检测肠道菌群及两种AHL对脆弱拟杆菌不同亚型luxR基因的(?)RNA的相对表达量,并对其表达水平进行比较。
结果：在脆弱拟杆菌基因组中,检测到8种luxR同源的QS类基因,只有LuxR8检测阴性,表明这8种luxR同源基因参与该菌Ⅰ型QS环路,脆弱拟杆菌基因组中未检测到luxl和其他lux基因。不同luxR基因亚型表达量具有一定的时间趋势,即培养6小时或4小时的表达量最高,而培养1小时表达量相对较低,8小时后表达下降；luxRl由于表达量少,故其各时间段的表达量基本一致,而luxR8基因不存在,相应地也无mRNA表达。十二烷基高丝氨酸内酯、大肠杆菌、金黄色葡萄球菌对luxR基因表达上调,而酮乙酰基高丝氨酸内酯和屎肠球菌对luxR基因表达下调。
结论：
本课题在创建了基于UPLC-MS技术的细菌群体感应系统一类信号分子AHL的检测平台基础上,通过对参与脆弱拟杆菌群体感应的一类信号分子AHL和QS相关基因的筛查、不同外源性AHL及肠道菌群对脆弱拟杆菌生物学活性及luxR基因表达影响的实验研究,得出以下结论：
1.脆弱拟杆菌具有一类群体感应系统,该系统包括LuxR调节因子参与,且LuxR调节因子具有多种亚型,LuxR调节因子可能与外源性的信号分子结合而发挥群体感应。
2.脆弱拟杆菌不存在luxⅠ基因,不产生酰基高丝氨酸内酯类自诱导信号分子。
3.UPLC-MS (Q-TOF-MS)技术是检测QS群体感应系统一类信号分子—高丝氨酸内酯的一种有效工具,具有准确度高、灵敏度高、精密度高、重复性好、检测方便、线性范围广等特点,AHL在Q-TOF-MS中总保持m/z为102.05特征离子峰或102.05和74.05的特征峰组合(N-已酰-DL-酰基高丝氨酸内酯只见102.05特征峰),该特征峰可作为AHL的质谱鉴定依据。
4.化学结构不同的高丝氨酸内酯分子及肠道常见细菌的分泌物对脆弱拟杆菌的群体感应系统发挥不同的效应。
5.脆弱拟杆菌在液态培养液中具有一定的耐氧性,并非严格厌氧菌,且该菌在甘油保存液中对反复冻融具有一定的抵抗力。
6.氯仿、甲醇、水混合溶剂是分离提取细菌群体感应系统的一类信号分子—高丝氨酸内酯的有效萃取溶剂。
Background
Autoinducer conducts the communication among bacteria. Bacteria can survey the amount changes in other bacteria according to the concentration of this signal molecules; it is a course of Quorum-sensing (QS). The signaling pathway is involved in various of physicial activities, such as bioluminescence, virulence gene expression, biofilms formation, exocellular enzyme and polysaccharides synthesis, antibiotics production, etc. Different bacteria apply unique QS mechanism in regulating biological activities. There are three classes of QS mechanism, modified polypeptides acting as signal regulatory molecular in gram-positive bacteria, AHL produced by LuxI/LuxR system doing the same activity in gram-negative bacteria, and the third one in both gram-positive and gram-negative bacteria is normally supposed to the secondary signal molecule. There is a great interest in research of QS in recent years but details about the QS in anaerobic bacteria have not been mentioned yet.
Bacteroides fragilis accounts for1%-2%of human intestine micro flora and also is the main anaerobic bacteria causing gastrointestinal infection. Their large amount presentation and formation of biofilm are induced by a multiple factors, among which is their information communication mechanism so as to keep a coincidence of a whole biological activity in human.
There are few researches about Bacteroides fragilis in the following aspects, the relations between QS and biofilm formation, whether it produces the QS signal molecules AHL and the contribution of AHL and intestine microflora to its biological activity. This research focused on the deep study of AHL in the direction of their influence factors and their presentation in bacterial QS based on UPLC-MS. Another part of the research approached whether Bacteroides fragilis have QS circulation and the impaction of QS related gene expression in Bacteroides fragilis by AHL originated from both extrinsic and other closely related enterobacteria in the way of molecular biology. This research, as the base for further study in QS mechanism, established the technique and methods for systemic research of intestinal microecological signal transduction and a clue for intestinal microecology pharmaceutics according to the Bacteroides fragilis QS signal molecule.
Part I Establishment of detection for QS signal molecule AHL
Methods:A total of6AHL standard substance were detected and analyzed by UPLC-MS(Quadrupole Time-of-Flight, Q-TOF) metabonomics in the parameters of solvent selection, moblie phase optimization, filter membrane aperture, electrospray ionization mode in mass spectrometry, chromatographic column selection, conditional optimization in mass spectrometry, preparation for AHL extraction and separation, and the ionogenic feature of AHL in mass spectrometry.
Results:Technological platform for bacterial QS signaling molecule AHL detection was established in the following parameters:positive ion electrospray mode, chromatographic column of Aquity C181.7μm2.1mm×100mm, Methyl Cyanides as solvent. The lactonic ring of AHL was kept unspoiled featuring as apiece m/s of102.05or both102.05and74.05during the mass spectrometry assay (N-Tetradecanoyl-DL-homoserine lactone'm/s is102.05). The extraction solvent composed of trichloromethane, methanol and water showed the sensitivity at5ng/ml for bacterial culture extraction.
Part II Detection of QS signal molecule AHL in Bacteroides fragilis based on UPLC-MS
Methods:All of the experimental bacteria including Bacteroides fragilis, bacillus bifidus, Escherichia coli, Enterococcus faecium, Vibrio parahaemolyticus, Enterobacteria cloacae, Acinetobacter baumannii, Salmonella typhosa, were ensured by sequencing their16SrRNA. Extracted bacterial culture from Bacteroides fragilis and other experimental bacteria were analyzed by UPLC-MS (Q-TOF-MS) to determine the presentation of AHL following the established parameters in this study, AHL standard substance as a control.
Results:16SrRNA in all of the experimental bacteria shared the more than98%sequence homology with GenBank database. The AHL standard substance solving in different bacterial culture showed a featured piece of m/s of102.05or both102.05and74.05in electrospray ionization quadrupole time-of-flight (Q-TOF) mass spectrometry (N-Tetradecanoyl-DL-homoserine lactone'm/s is102.05), while AHL was detected in none of the Bacteroides fragilis and other bacteria.
Part III Study on Bacteroides fragilis culture feature
Methods:The survival of Bacteroides fragilis was recorded in the condition of different period, temperature and after10times freezing and thawing, when preserved in both anaerobic glycerol broth and common glycerol broth. Its viability and survival rate were compared when exposed in the air for5min,10min,30min,60min,90min,120min,150min,180min,24h.
Results:Bacteroides fragilis was cultured successfully when persevered at either-20℃or-80℃for as long as3months in both anaerobic broth and common broth, and survived even after freezing and thawing for10times. It was still live when exposed in the air for180min but dead for24h. Its survival rate after exposed in air for5min, lOmin,30min,60min was84.1%,68.1%,46.4%and34.8%respectively.
Part Ⅳ The impact of extrinsic N-Acyl-Homoserine Lactone signal molecules and intestine microflora on Bacteroides fragilis biological activity
Methods:Bacteroides fragilis was monitored by OD value during a rang of culture period with the culture supernatant from5common intestinal bacteria and2AHL standard substances in the anaerobic blood broth.
Results:Standard Bacteroides fragilis reached the log phase after culturing for8h,6h,6h,6h,10h, and10h, and stable phase after10h,8h,8h,8h,24h,and24h, in standard anaerobic hemoculture, anaerobic hemoculture added with N-Dodecanoyl-DL-homoserine lactone, culture supernatant from Enterococcus faecium, Staphylococcus aureus, Enterococcus faecium, N-(β-Ketocaproyl)-DL-homoserine lactone respectively. It took standard Bacteroides fragilis the same time in reaching the log phase and stable phase in anaerobic hemoculture added with other culture supernatant as in standard one.
Part Ⅴ Detection of Bacteroides fragilis QS related gene expression in different culture condition by fluorescent quantified PCR
Methods:Bacteroides fragilis QS related gene luxI, luxR and subtypes were detected by PCR. The mRNA of luxR subtypes in Bacteroides fragilis was quantitatived by real-time fluorescent quantitative PCR and further compared in the impact of intestine microflora and2AHLs.
Results:A total of8luxR-homologous genes, except luxR8, were detected in Bacteroides fragilis, while none of luxI and other lux genes were detected. This result indicated the8luxR-homologous genes were involved in class Ⅰ QS circulation. LuxR subtypes expression is different in period of culture, reaching the highest level after6h or4h culture while the lowest after1h or8h., with the except of luxR1that was expressed in low quality all the period and luxR8of non-presentation. LuxR expression was up-regulated by N-Dodecanoyl-DL-homoserine lactone, Enterococcus faecium and Staphylococcus aureus, on the contrary, down-regulated by N-(β-Ketocaproyl)-DL-homoserine lactone and Enterococcus faecium.
Conclusions
In this study, the bacterial QS Class Ⅰ signal molecule AHL detection platform based on UPLC-MS technique was established, class Ⅰ signal molecule AHL and QS related genes in Bacteroides fragilis was determined, and the impact of different extrinsic AHL and enterobacteria on the biological activity and luxR gene expression was further explored, so it arrived at conclusions as follows.
1. Bacteroides fragilis have class Ⅰ QS. LuxR regulatory factors and their subtypes were involved in this system and may play roles in QS when conjuncted with extrinsic signal molecule.
2. Neither LuxⅠ gene presented nor N-Acyl-Homoserine Lactone (AHL) was produced in Bacteroides fragilis.
3. UPLC-MS (Q-TOF-MS) technique is useful in detection of QS Class Ⅰ signal molecule AHL with the advantage of high accuracy, high sensitivity, good repeatability, easy operate, high efficiency. The lactonic ring of AHL was kept unspoiled featuring as apiece m/s of102.05or both102.05and74.05in Q-TOF-MS.
4. Different extrinsic AHLs and common enterobacteria secretions produced different effect on QS in Bacteroides fragilis.
5. Bacteroides fragilis was tolerant to oxygen to some extent.
6. The extraction solvent composed of trichloromethane, methanol and water was optimized in extraction QS Class I signal molecule AHL,
细菌之间的交流是通过分泌和感应一种被称为自诱导剂(Autoinducer, AI)的信号分子来实现的。细菌根据这种特定信号分子浓度的变化来监测周围环境中其他细菌数量的变化,这种信号传递的过程称为群体感应(Quorum-sensing, QS)。现有报道的细菌QS环路有三种机制：革兰阳性菌以修饰多肽作为信号调节分子；革兰阴性菌主要以典型的LuxI/LuxR系统通过产生一类信号分子N-酰基-高丝氨酸内酯类(AHL)进行群体感应；另外一种QS分子目前认为是二类信号分子(呋喃硼酸二酯),为革兰阴性菌和阳性菌共有。最近几年,有关需氧菌的QS系统研究方兴未艾,研究范围越来越广。但是,厌氧菌的QS系统类似研究至今还未见报道。
脆弱拟杆菌占人体肠道菌群的比例约为1%-2%,是引起胃肠感染的最主要的厌氧菌,多种因素导致该菌可以在肠道内大量存在并形成生物膜。脆弱拟杆菌生物膜形成是否与QS有关,该菌是否产生一类QS信号分子AHL,以及AHL与肠道菌群对该菌生物学活性的影响,目前相关研究报道极少,因此,本研究重点研究了UPLC-MS(四极杆飞行时间质谱,Q-TOF-MS)检测AHL分子的影响因素,创建了基于UPLC-MS技术的细菌群体感应一类信号分子AHL的检测平台,为系统研究肠道微生态细菌的信号传导提供实验技术和方法。另外采用分子生物学技术探讨脆弱拟杆菌是否存在QS环路系统,同时分析了与脆弱拟杆菌密切相关的其他肠道细菌及外源性AHL对脆弱拟杆菌QS相关基因表达的影响,为进一步研究其群体感应发生机制奠定基础。
实验一N-酰基-高丝氨酸内酯类群体感应信号分子检测方法的建立
方法：采用UPLC/MS (Q-TOF-MS)代谢组学分析方法检测6种N-酰基-高丝氨酸内酯标准品,重点考察UPLC-MS检测AHL分子的各种参数,包括溶剂的选择、流动相的优化、滤膜孔径、质谱的电喷雾离子化模式、色谱柱的选择、质谱条件的优化、AHL提取分离的前处理条件、AHL分子在质谱中的离子化特点等。
结果：创建了基于UPLC-MS技术的细菌群体感应系统一类信号分子AHL的检测平台,其检测参数为：正离子电喷雾离子化模式、Aquity C181.7μm2.1mm×100mm色谱柱,乙腈作为AHL溶剂及流动相。AHL内酯环结构在质谱鉴定过程中不会被破坏,在电喷雾电离四极杆飞行时间质谱中总保持m/z为102.05和74.05的特征峰组合(N-已酰-DL-酰基高丝氨酸内酯只见102.05特征峰),三氯甲烷甲醇水混合液作为萃取剂提取细菌培养液中AHL的灵敏度为5ng/ml。
实验二基于UPLC-MS技术检测脆弱拟杆菌群体感应一类信号分子AHL
方法：采用PCR方法扩增实验菌株(包括脆弱拟杆菌、5种双歧杆菌、大肠埃希菌、屎肠球菌、金黄色葡萄球菌、副溶血弧菌、阴沟肠杆菌、鲍曼不动杆菌、伤寒沙门菌)的16SrRNA基因并测序,确保实验菌株鉴定的准确性；在确定UPLC-MS (Q-TOF-MS)检测AHL的各种检测参数后,采用该方法对脆弱拟杆菌和其他多种细菌的菌液提取物中是否存在AHL进行了检测,并以AHL标准品做对照实验。
结果：实验菌株16SrRNA基因序列与GenBank数据库序列同源性大于98%,采用UPLC-MS:法检测溶于不同细菌培养液中AHL标准品,其电喷雾电离四极杆飞行时间质谱均见m/z为102.05特征离子碎片或102.05和74.05特征碎片组合(N-已酰-DL-酰基高丝氨酸内酯只见102.05特征峰),而脆弱拟杆菌和其他多种细菌的菌液均未检测到AHL。
实验三脆弱拟杆菌生物学特性的研究
方法：采用甘油厌氧肉汤及甘油普通肉汤对脆弱拟杆菌进行保存,观察在不同时间及不同温度保存条件下的存活状况,同时观察反复冻融10次对其存活力的影响。另将脆弱拟杆菌菌液分别暴露在空气中5min、10min、30min、60min、90min、120min、150min、180min、24h,观察不同时间段脆弱拟杆菌的的生存力及生存率的变化。
结果：接种于甘油厌氧肉汤及普通肉汤中的脆弱拟杆菌在-20℃及-80℃两种温度下保存3个月后,反复冻融10次后均能培养成功,与空气接触180min时仍生存,24h后即死亡,与空气接触5min,10min,30min,60min后生存率分别为84.1%,68.1%,46.4%,34.8%。
实验四外源信号分子及肠道菌群对脆弱拟杆菌生物学活性的影响
方法：将5种肠道常见菌的培养上清液及2种AHL标准品加入厌氧血培养液中,用于脆弱拟杆菌培养,检测不同培养时间段的脆弱拟杆菌菌液OD值,分析其在不同培养环境的生长曲线变化。
结果：脆弱拟杆菌在厌氧血培养液、含十二烷基高丝氨酸内酯、大肠杆菌、金黄色葡萄球菌、屎肠球菌上清液、酮乙酰基高丝氨酸内酯的厌氧血培养中培养时,其进入对数期的时间分别为8h、6h、6h、6h、10h、10h,其进入稳定期时间分别为10h、8h、8h、8h、24h、24h,脆弱拟杆菌在含其他细菌上清液的厌氧血培养液中进入对数期及稳定期的时间与空白厌氧血培养液一致。
实验五荧光定量PCR法检测不同培养环境脆弱拟杆菌群体感应相关基因的表达
方法：采用PCR方法检测群体感应相关基因luxl及脆弱拟杆菌的luxR基因及其亚型,实时荧光定量PCR方法检测肠道菌群及两种AHL对脆弱拟杆菌不同亚型luxR基因的(?)RNA的相对表达量,并对其表达水平进行比较。
结果：在脆弱拟杆菌基因组中,检测到8种luxR同源的QS类基因,只有LuxR8检测阴性,表明这8种luxR同源基因参与该菌Ⅰ型QS环路,脆弱拟杆菌基因组中未检测到luxl和其他lux基因。不同luxR基因亚型表达量具有一定的时间趋势,即培养6小时或4小时的表达量最高,而培养1小时表达量相对较低,8小时后表达下降；luxRl由于表达量少,故其各时间段的表达量基本一致,而luxR8基因不存在,相应地也无mRNA表达。十二烷基高丝氨酸内酯、大肠杆菌、金黄色葡萄球菌对luxR基因表达上调,而酮乙酰基高丝氨酸内酯和屎肠球菌对luxR基因表达下调。
结论：
本课题在创建了基于UPLC-MS技术的细菌群体感应系统一类信号分子AHL的检测平台基础上,通过对参与脆弱拟杆菌群体感应的一类信号分子AHL和QS相关基因的筛查、不同外源性AHL及肠道菌群对脆弱拟杆菌生物学活性及luxR基因表达影响的实验研究,得出以下结论：
1.脆弱拟杆菌具有一类群体感应系统,该系统包括LuxR调节因子参与,且LuxR调节因子具有多种亚型,LuxR调节因子可能与外源性的信号分子结合而发挥群体感应。
2.脆弱拟杆菌不存在luxⅠ基因,不产生酰基高丝氨酸内酯类自诱导信号分子。
3.UPLC-MS (Q-TOF-MS)技术是检测QS群体感应系统一类信号分子—高丝氨酸内酯的一种有效工具,具有准确度高、灵敏度高、精密度高、重复性好、检测方便、线性范围广等特点,AHL在Q-TOF-MS中总保持m/z为102.05特征离子峰或102.05和74.05的特征峰组合(N-已酰-DL-酰基高丝氨酸内酯只见102.05特征峰),该特征峰可作为AHL的质谱鉴定依据。
4.化学结构不同的高丝氨酸内酯分子及肠道常见细菌的分泌物对脆弱拟杆菌的群体感应系统发挥不同的效应。
5.脆弱拟杆菌在液态培养液中具有一定的耐氧性,并非严格厌氧菌,且该菌在甘油保存液中对反复冻融具有一定的抵抗力。
6.氯仿、甲醇、水混合溶剂是分离提取细菌群体感应系统的一类信号分子—高丝氨酸内酯的有效萃取溶剂。
Background
Autoinducer conducts the communication among bacteria. Bacteria can survey the amount changes in other bacteria according to the concentration of this signal molecules; it is a course of Quorum-sensing (QS). The signaling pathway is involved in various of physicial activities, such as bioluminescence, virulence gene expression, biofilms formation, exocellular enzyme and polysaccharides synthesis, antibiotics production, etc. Different bacteria apply unique QS mechanism in regulating biological activities. There are three classes of QS mechanism, modified polypeptides acting as signal regulatory molecular in gram-positive bacteria, AHL produced by LuxI/LuxR system doing the same activity in gram-negative bacteria, and the third one in both gram-positive and gram-negative bacteria is normally supposed to the secondary signal molecule. There is a great interest in research of QS in recent years but details about the QS in anaerobic bacteria have not been mentioned yet.
Bacteroides fragilis accounts for1%-2%of human intestine micro flora and also is the main anaerobic bacteria causing gastrointestinal infection. Their large amount presentation and formation of biofilm are induced by a multiple factors, among which is their information communication mechanism so as to keep a coincidence of a whole biological activity in human.
There are few researches about Bacteroides fragilis in the following aspects, the relations between QS and biofilm formation, whether it produces the QS signal molecules AHL and the contribution of AHL and intestine microflora to its biological activity. This research focused on the deep study of AHL in the direction of their influence factors and their presentation in bacterial QS based on UPLC-MS. Another part of the research approached whether Bacteroides fragilis have QS circulation and the impaction of QS related gene expression in Bacteroides fragilis by AHL originated from both extrinsic and other closely related enterobacteria in the way of molecular biology. This research, as the base for further study in QS mechanism, established the technique and methods for systemic research of intestinal microecological signal transduction and a clue for intestinal microecology pharmaceutics according to the Bacteroides fragilis QS signal molecule.
Part I Establishment of detection for QS signal molecule AHL
Methods:A total of6AHL standard substance were detected and analyzed by UPLC-MS(Quadrupole Time-of-Flight, Q-TOF) metabonomics in the parameters of solvent selection, moblie phase optimization, filter membrane aperture, electrospray ionization mode in mass spectrometry, chromatographic column selection, conditional optimization in mass spectrometry, preparation for AHL extraction and separation, and the ionogenic feature of AHL in mass spectrometry.
Results:Technological platform for bacterial QS signaling molecule AHL detection was established in the following parameters:positive ion electrospray mode, chromatographic column of Aquity C181.7μm2.1mm×100mm, Methyl Cyanides as solvent. The lactonic ring of AHL was kept unspoiled featuring as apiece m/s of102.05or both102.05and74.05during the mass spectrometry assay (N-Tetradecanoyl-DL-homoserine lactone'm/s is102.05). The extraction solvent composed of trichloromethane, methanol and water showed the sensitivity at5ng/ml for bacterial culture extraction.
Part II Detection of QS signal molecule AHL in Bacteroides fragilis based on UPLC-MS
Methods:All of the experimental bacteria including Bacteroides fragilis, bacillus bifidus, Escherichia coli, Enterococcus faecium, Vibrio parahaemolyticus, Enterobacteria cloacae, Acinetobacter baumannii, Salmonella typhosa, were ensured by sequencing their16SrRNA. Extracted bacterial culture from Bacteroides fragilis and other experimental bacteria were analyzed by UPLC-MS (Q-TOF-MS) to determine the presentation of AHL following the established parameters in this study, AHL standard substance as a control.
Results:16SrRNA in all of the experimental bacteria shared the more than98%sequence homology with GenBank database. The AHL standard substance solving in different bacterial culture showed a featured piece of m/s of102.05or both102.05and74.05in electrospray ionization quadrupole time-of-flight (Q-TOF) mass spectrometry (N-Tetradecanoyl-DL-homoserine lactone'm/s is102.05), while AHL was detected in none of the Bacteroides fragilis and other bacteria.
Part III Study on Bacteroides fragilis culture feature
Methods:The survival of Bacteroides fragilis was recorded in the condition of different period, temperature and after10times freezing and thawing, when preserved in both anaerobic glycerol broth and common glycerol broth. Its viability and survival rate were compared when exposed in the air for5min,10min,30min,60min,90min,120min,150min,180min,24h.
Results:Bacteroides fragilis was cultured successfully when persevered at either-20℃or-80℃for as long as3months in both anaerobic broth and common broth, and survived even after freezing and thawing for10times. It was still live when exposed in the air for180min but dead for24h. Its survival rate after exposed in air for5min, lOmin,30min,60min was84.1%,68.1%,46.4%and34.8%respectively.
Part Ⅳ The impact of extrinsic N-Acyl-Homoserine Lactone signal molecules and intestine microflora on Bacteroides fragilis biological activity
Methods:Bacteroides fragilis was monitored by OD value during a rang of culture period with the culture supernatant from5common intestinal bacteria and2AHL standard substances in the anaerobic blood broth.
Results:Standard Bacteroides fragilis reached the log phase after culturing for8h,6h,6h,6h,10h, and10h, and stable phase after10h,8h,8h,8h,24h,and24h, in standard anaerobic hemoculture, anaerobic hemoculture added with N-Dodecanoyl-DL-homoserine lactone, culture supernatant from Enterococcus faecium, Staphylococcus aureus, Enterococcus faecium, N-(β-Ketocaproyl)-DL-homoserine lactone respectively. It took standard Bacteroides fragilis the same time in reaching the log phase and stable phase in anaerobic hemoculture added with other culture supernatant as in standard one.
Part Ⅴ Detection of Bacteroides fragilis QS related gene expression in different culture condition by fluorescent quantified PCR
Methods:Bacteroides fragilis QS related gene luxI, luxR and subtypes were detected by PCR. The mRNA of luxR subtypes in Bacteroides fragilis was quantitatived by real-time fluorescent quantitative PCR and further compared in the impact of intestine microflora and2AHLs.
Results:A total of8luxR-homologous genes, except luxR8, were detected in Bacteroides fragilis, while none of luxI and other lux genes were detected. This result indicated the8luxR-homologous genes were involved in class Ⅰ QS circulation. LuxR subtypes expression is different in period of culture, reaching the highest level after6h or4h culture while the lowest after1h or8h., with the except of luxR1that was expressed in low quality all the period and luxR8of non-presentation. LuxR expression was up-regulated by N-Dodecanoyl-DL-homoserine lactone, Enterococcus faecium and Staphylococcus aureus, on the contrary, down-regulated by N-(β-Ketocaproyl)-DL-homoserine lactone and Enterococcus faecium.
Conclusions
In this study, the bacterial QS Class Ⅰ signal molecule AHL detection platform based on UPLC-MS technique was established, class Ⅰ signal molecule AHL and QS related genes in Bacteroides fragilis was determined, and the impact of different extrinsic AHL and enterobacteria on the biological activity and luxR gene expression was further explored, so it arrived at conclusions as follows.
1. Bacteroides fragilis have class Ⅰ QS. LuxR regulatory factors and their subtypes were involved in this system and may play roles in QS when conjuncted with extrinsic signal molecule.
2. Neither LuxⅠ gene presented nor N-Acyl-Homoserine Lactone (AHL) was produced in Bacteroides fragilis.
3. UPLC-MS (Q-TOF-MS) technique is useful in detection of QS Class Ⅰ signal molecule AHL with the advantage of high accuracy, high sensitivity, good repeatability, easy operate, high efficiency. The lactonic ring of AHL was kept unspoiled featuring as apiece m/s of102.05or both102.05and74.05in Q-TOF-MS.
4. Different extrinsic AHLs and common enterobacteria secretions produced different effect on QS in Bacteroides fragilis.
5. Bacteroides fragilis was tolerant to oxygen to some extent.
6. The extraction solvent composed of trichloromethane, methanol and water was optimized in extraction QS Class I signal molecule AHL,
引文
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