CN116287335B - 评估阿拉伯木聚糖对肠道微生态调节作用的方法及其应用 - Google Patents
评估阿拉伯木聚糖对肠道微生态调节作用的方法及其应用 Download PDFInfo
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Abstract
本发明公开了一种评估阿拉伯木聚糖对肠道微生态调节作用的方法及其应用。1)体外进行模拟阿拉伯木聚糖胃‑小肠消化,获取未消化残渣作为发酵底物;2)收集猪新鲜粪菌;3)配制肠道微生物体外模型培养基;4)37℃体外孵育;5)测定发酵过程中样品中的短链脂肪酸含量变化、α‑L‑阿拉伯呋喃糖苷酶和β‑木聚糖酶的酶活性变化;并对发酵液中微生物进行16s高通量测序,检测样品中优势菌群相对丰度变化,判断阿拉伯木聚糖对肠道微生物的影响。本发明快速易行、简单高效,解决阿拉伯木聚糖功能活性研究中难以直接体内探究动态调控以及个性化评估差异大的问题,可评估不同品种肠道微生物利用阿拉伯木聚糖的代谢差异,实现个体化营养。
Description
技术领域
本发明属于生物技术和畜牧领域,具体涉及利用体外发酵和微生物测序技术快速评估一种纤维原料对肠道微生态促进作用的方法及将其用于靶向筛选代谢阿拉伯木聚糖的靶点肠道微生物。
背景技术
膳食纤维是一种自然来源的功能性物质,通过为微生物的生长提供底物来改变肠道的微生物环境,能利用膳食纤维的微生物会扩大他们自身的数量。肠道微生物拥有多种编码糖苷水解酶、多糖裂合酶,碳水化合物脂酶家族的基因,根据纤维物质的可利用性为微生物的动态变化提供不同的能量来源。膳食纤维能够维持肠道生态健康,调节营养元素和宿主的生理健康。通过为肠道微生物筛选新型纤维物质,理解其与微生物之间的相互作用,将有助于动物健康发展。
肠道微生物组的个性化特征受环境、生活方式和遗传的影响,导致宿主对饮食等干预措施的反应不同,如相同纤维干预不同的受试者,往往展现出不同的效果。由于肠道菌群本身的复杂性,如何通过有效手段准确识别差异生物标志物,预测干预措施引起的微生物反应有助于提高纤维干预的功效。
多数研究基于长期饮食对肠道微生物群落的影响,但是大量营养成分会在24小时内引起肠道微生物的显著变化,既肠道微生物群落会做出短期的应答反应。特别是,当饮食中纤维含量变化时,将导致具有降解纤维物质能力的微生物动态变化以及整个群落快速的重组。如果能够通过一套有效的方法在体外对肠道菌群代谢纤维的情况进行研究将非常有助于理解个体营养反应,从而解决动物营养问题。
阿拉伯木聚糖是由β-D-吡喃木糖残基经β-(1→4)-糖苷键连接而成的木聚糖主链和α-L-呋喃阿拉伯糖为侧链的多分支结构,是天然存在于粮谷类植物细胞壁的主要结构性多糖,是一种重要的膳食纤维和功能糖。作为一种不易消化的碳水化合物,具有促进肠道有益菌增殖,改善肠道菌群数量及结构的作用。阿拉伯木聚糖被肠道微生物降解生成SCFA可使肠道pH降低,促进营养物质吸收,改善肠道健康。同时其在调节肠道屏障、免疫功能、糖脂代谢等方面发挥着重要作用。因此研究肠道菌群与阿拉伯木聚糖代谢的关系可提供新的营养干预靶点。但是,目前有关阿拉伯木聚糖影响肠道微生物代谢的研究主要依赖于动物模型体内实验,具体体内与菌群的相互作用的活性机制研究较少。
发明内容
基于上述问题,本发明的目的是提供评估阿拉伯木聚糖对肠道微生态调节作用的方法及其应用。本发明提出了一种快速易行、简单高效的基于肠道菌群体外模型,解决阿拉伯木聚糖功能活性研究中难以直接体内探究动态调控以及个性化评估差异大的问题,建立评估阿拉伯木聚糖对肠道微生态调节作用的方法,可评估不同品种肠道微生物利用阿拉伯木聚糖的代谢差异,为个体化营养提供方法和指导。
一种评估阿拉伯木聚糖对肠道微生态调节作用的方法,步骤如下:
1)体外进行模拟阿拉伯木聚糖胃-小肠消化,获取未消化残渣作为发酵底物;
2)收集猪新鲜粪菌;
3)配制肠道微生物体外模型培养基;
4)在厌氧培养箱中,将步骤2)中的粪便微生物接种至步骤3)所述培养基,试验组加入步骤1的发酵底物进行体外共培养,不加发酵底物的发酵瓶作为空白对照组,试验组和对照组于37℃进行体外孵育;
5)测定发酵过程中样品中的短链脂肪酸(SCFA)含量变化、α-L-阿拉伯呋喃糖苷酶和β-木聚糖酶的酶活性变化;并对发酵液中微生物进行16s高通量测序,检测样品中优势菌群相对丰度变化,判断阿拉伯木聚糖对肠道微生物的影响。
所述的方法,步骤1)所述的发酵底物制备方法为:25g阿拉伯木聚糖溶于300 mL的PBS磷酸缓冲液中,加入2.25 ml α-淀粉酶,37℃,150 rpm,反应15 min。用1M HCl调整pH至2.5±0.1,加入10 mL 质量百分比10%的胃蛋白酶,37℃,150 rpm,反应30 min;随后加入50mL 0.1M马来酸钠缓冲液,加入1 M NaHCO3调整pH至6.9±0.1,再加入50 mL 质量百分比12.5% 胰蛋白酶,最后加入2 mL淀粉葡萄糖甘酶,37℃,150 rpm,反应2 h。采用95%乙醇进行沉淀处理,透析24 h后冻干得到消化后的阿拉伯木聚糖用于体外厌氧发酵。
步骤2)中粪便的采集为选择断奶后2个月健康的杜长大商品猪和金华猪各6头,所有猪只饲养管理一致;采食后同一时间段内从猪直肠中收集粪便样本,并将其转移到无菌冻存管中,所有操作均在厌氧条件下进行,液氮处理后在-80℃进行冷冻储存。
所述的方法,步骤3)中的培养基配方为:0.16 g/L 蛋白胨,0.1 g/L 酵母浸出物,0.16 g/L 吐温80,0.16 g/L NaHCO3,3.6 g/L NaCl,1.6 g/L K2HPO4,0.32 g/L L-半胱氨酸盐酸盐,0.36 g/L CaCl2·6H2O,0.5 g/L MgSO4·7H2O,0.01 g/L 血红素。
所述的方法,步骤4)包括:将1 g预消化处理的阿拉伯木聚糖溶于25 mL无菌发酵培养基中,100℃煮沸10 min;随后将其放入厌氧室中,冷却到室温,放置2 h,得到待发酵的阿拉伯木聚糖溶液。0.1 g粪便样本添加到5 mL的无菌培养基中,振荡涡旋混匀至均质状态得到待接种的粪菌悬液;将2.5 mL粪菌液与2.5 mL发酵底物溶液混合后得到最终发酵体系,37°C厌氧条件下以130 rpm振荡培养;在发酵的第1、3、6、9、12、15、18、21、24、48和72h进行采样;采取冰浴终止发酵,将发酵液离心,上清液和沉淀物80℃冻存。
所述的方法,测量各时间点pH值、气相色谱法定量分析短链脂肪酸含量、ELISA法测定α-L-阿拉伯呋喃糖苷酶和β-木聚糖酶活性;提取各个时间点发酵液离心后沉淀物中基因组DNA,所有样品均使用适用于细菌16S rDNA 的V3-V4区域进行PCR扩增,利用IlluminaMiSeq平台测序;得到的以上数据,作为评估阿拉伯木聚糖对肠道微生态影响的指标。
所述的方法的应用,用于评估不同品种肠道微生物利用阿拉伯木聚糖的代谢差异,筛选阿拉伯木聚糖靶向促进生长的微生物种类。
所述的方法的应用,用于研究个体内营养物质对肠道微生物的动态调节过程,实现个体化营养。
与现有技术相比,本发明具有如下优点:
由于动物体内评估的复杂性以及一些内源因素的干扰。为此,本发明建立了一种统一规范并简单易行的动物肠道菌群体外研究模型,在保证体外研究结果可靠的基础上,有效提高研究结果的可操作性。不仅有助于更快速的理解个体内营养物质对肠道微生物的动态调节过程,同时有利于精准筛选出阿拉伯木聚糖靶向促进生长的微生物种类。
附图说明
图1是肠道微生物体外培养模型示意图。
图2是发酵过程中pH和酶活性变化图。
图3是发酵过程中短链脂肪酸含量变化变化图。
图4是发酵过程中肠道微生物多样性变化图。
图5是发酵阶段图。
图6是肠道微生物在门水平的变化图。
图7是各阶段代表性时间点在属水平的差异微生物图。
具体实施方式
以下结合实施例和附图对本发明做进一步的阐述。
实施例1体外模拟消化阿拉伯木聚糖
如图1所示,25g阿拉伯木聚糖溶于300 mL的PBS磷酸缓冲液中,加入2.25 ml α-淀粉酶,37℃,150 rpm,反应15 min。用1M HCl调整pH至2.5±0.1,加入10 mL 质量百分比10%的胃蛋白酶,37℃,150 rpm,反应30 min。随后加入50 mL 0.1M马来酸钠缓冲液,加入1 MNaHCO3调整pH至6.9±0.1,再加入50 mL 质量百分比12.5% 胰蛋白酶,最后加入2 mL淀粉葡萄糖甘酶,37℃,150 rpm,反应2 h。采用95%乙醇进行沉淀处理,透析24 h后冻干得到消化后的阿拉伯木聚糖用于体外厌氧发酵。
实施例2阿拉伯木聚糖对肠道微生物的动态调控作用
1、粪便接种发酵
培养基配制按表1中的方法进行,选择断奶后2个月健康的杜长大商品猪(DLY)和金华猪(JH)各6头。具体操作过程按照图1所示,采集其粪菌并进行发酵底物(预消化处理阿拉伯木聚糖)体外肠道微生物共孵育。将1 g已进行体外消化的菊粉置于25 mL无菌发酵培养基中,煮沸10 min,煮沸后的纤维溶液转移至厌氧室,冷却至室温。将0.1 g粪便样本加入5 mL无菌发酵培养基中,均质。取2.5 mL均质粪菌悬液与2.5 mL纤维溶液混合,在厌氧条件下,37°C, 130 rpm振荡培养。所有发酵步骤都在厌氧室中进行。在体外发酵的第1、3、6、9、12、15、18、21、24、48和72 h进行采样。
表1. 培养基配制参数
2、测量pH、短链脂肪酸(SCFA)、α-L-阿拉伯呋喃糖苷酶和β-木聚糖酶活性
使用标准pH计测量每个采样时间点的pH值。取1 ml发酵液,20, 000× g离心15min,取上清液800 μL,与200 μL的25% (w/v)磷酸混合,随后取200 μL进行气相色谱上机,检测SCFA含量。样品8, 000 × g离心20 min后,取上清液,采用ELISA法测定α-L-阿拉伯呋喃糖苷酶和β-木聚糖酶活性。
3、微生物测序
采用E.Z.N.A.®粪便DNA试剂盒提取微生物基因组DNA。提取的DNA浓度和纯度用NanoDrop 2000 (Thermo Scientific, Wilmington, USA)测定,用1%琼脂糖凝胶检测。所有样本均采用V3-V4区进行PCR扩增,同一样品PCR产物混合后用2%琼脂糖凝胶电泳检测扩增效果,目的条带用DNA切胶回收试剂盒进行回收。将已构建好的PCR产物文库参照电泳初步定量结果,采用AxyPrep DNA凝胶提取试剂盒和Quantus™荧光计进行DNA荧光定量检测,样品等量混合,构建基因组测序上机文库,通过 Illumina MiSeq 测序平台进行双末端测序。
4、生物信息分析
使用Illumina-utils 拼接质量控制后的序列,使用UCHIME移除 PCR 扩增过程中的嵌合体;使用UPARSE 聚类操作分类单元(OTU)。细菌从门到属的分类由Greengenes数据库进行对比注释。采用QIIME1进行多样性分析。1)发酵阶段的分析:采用R语言中的vegdist包计算各个时间点的微生物群落相似性(Bray-Curtis距离),eclust进行层次聚类分析,根据K-means得到最佳分类簇。2)采用LEfSe(Linear discriminant analysis Effect Size, LEfSe)分析方法进行差异显著性分析,并根据线性判别分析(LDA)对数据进行分类和评估差异显著物种的影响力(即LDA score),LDA阈值默认为2-4,绘制LDA柱状图。
5、结果与分析
不同时间点表观指标变化
如图2所示,阿拉伯木聚糖体外发酵后,pH值呈现持续下降趋势,其中JH在21 h时的pH值显著高于DLY,而DLY在3h时和72 h显著高于JH(P<0.05)。发酵过程中两个猪种β-木聚糖酶活性呈现相似的变化,而α-L-阿拉伯呋喃糖苷酶活性是完全不同的变化趋势,JH在1h时的β-木聚糖酶活性高于DLY,而DLY在18 h和24 h时的α-L-阿拉伯呋喃糖苷酶活性显著高于JH(P<0.05)。此外,在发酵的前24 h金华猪的β-木聚糖酶与α-L-阿拉伯呋喃糖苷酶的变化为相反变化,而杜长大猪并无此变化现象,表明两者肠道菌群利用阿拉伯木聚糖的机制可能不同。如图3所示,金华猪和杜长大猪体外发酵阿拉伯木聚糖产生的SCFA水平有差异,与DLY相比,JH在48和72 h的乙酸含量较高(P<0.05),1 h时DLY丙酸产量较高(P<0.05),并且DLY中丁酸浓度在9、12、15、21、24和72 h时均高于JH(P<0.05)。总SCFA浓度JH在48和72h显著高于DLY,而在1 h和9 h时显著低于DLY(P<0.05)。以上结果说明了阿拉伯木聚糖调控两者肠道微生物的动态代谢差异。
微生物多样性的动态变化和发酵阶段分析
如图4所示,JH和DLY的微生物丰富度(OTUs数量)和多样性(Shannon指数)均呈下降趋势,并且JH的α多样性要高于DLY。两个品种的β多样性有显著差异,不同时间点微生物结构呈现出随时间分布的特征,在发酵前24 h时DLY的微生物结构变化幅度要高于JH。如图5所示,根据Bray-Curtis的群落结构相似性,JH可分为5个阶段:1 h、3-6 h、9-12 h、15-24h和48-72 h,DLY分为3个阶段:1-3 h、6-21 h和24-72 h。对两者的时间点共同聚类分析发现,JH和DLY在前24 h的微生物群落结构不同,48-72 h相似。这些结果表明,在短暂时间内阿拉伯木聚糖对两者肠道微生物的调控具有个体化差异,再次说明了体外发酵模型的构建对理解膳食纤维的动态调控作用的重要性。
微生物在门水平的动态变化和差异物种分析
如图6所示,两个猪种体外发酵阿拉伯木聚糖后,对应着发酵过程中微生物群落的动态演替。如图7所示,进一步地选取阿拉伯木聚糖调控肠道微生物不同阶段的代表性时间点:1 h、15 h和72 h进行LEfSe分析。结果显示,在1 h时,DLY中以巨球型菌、考拉杆菌和罕见小球菌为优势菌属,JH中以志贺氏菌、库特氏菌和毛螺菌为优势菌属;随着时间的变化,优势菌属也在发生改变,发酵中期15 h时,DLY中仍然以巨球型菌为主要代谢阿拉伯木聚糖的微生物,而JH主要为不动杆菌、芽孢杆菌、克里斯滕森菌和库特氏菌;在发酵后期,相比DLY,JH中显著富集的微生物为双歧杆菌和链球菌。以上结果再次表明了阿拉伯木聚糖调控两者肠道微生态过程中菌群的动态差异变化,该过程促进了纤维降解菌的增殖和有益菌的生长,说明了阿拉伯木聚糖对肠道菌群的益生作用。
以上所述实施例仅表达了本发明的几种实施方式,其描述较为具体和详细,但并不能因此而理解为对发明范围的限制。应当指出的是,对于本领域的普通技术人员来说,在不脱离本发明构思的前提下,还可以做出若干变形和改进,这些都属于本发明的保护范围。因此,本发明的保护范围应以所附权利要求为准。
Claims (1)
1.一种评估阿拉伯木聚糖对肠道微生态调节作用的方法,其特征在于,步骤如下:
1)体外进行模拟阿拉伯木聚糖胃-小肠消化,获取未消化残渣作为发酵底物;
2)收集金华猪和杜长大猪新鲜粪菌;
3)配制肠道微生物体外模型培养基;
4)在厌氧培养箱中,将步骤2)中的粪便微生物接种至步骤3)所述培养基,试验组加入步骤1)的发酵底物进行体外共培养,不加发酵底物的发酵瓶作为空白对照组,试验组和对照组于37℃进行体外孵育;
5)测定发酵过程中样品中的短链脂肪酸(SCFA)含量变化、α-L-阿拉伯呋喃糖苷酶和β-木聚糖酶的酶活性变化;并对发酵液中微生物进行16s高通量测序,检测样品中优势菌群相对丰度变化,判断阿拉伯木聚糖对肠道微生物的影响;
步骤1)所述的发酵底物制备方法为:25g阿拉伯木聚糖溶于300 mL的PBS磷酸缓冲液中,加入2.25 ml α-淀粉酶,37℃,150 rpm,反应15 min;用1M HCl调整pH至2.5±0.1,加入10 mL 质量百分比10%的胃蛋白酶,37℃,150 rpm,反应30 min;随后加入50 mL 0.1M马来酸钠缓冲液,加入1 M NaHCO3调整pH至6.9±0.1,再加入50 mL 质量百分比12.5% 胰蛋白酶,最后加入2 mL淀粉葡萄糖甘酶,37℃,150 rpm,反应2 h;采用95%乙醇进行沉淀处理,透析24 h后冻干得到消化后的阿拉伯木聚糖用于体外厌氧发酵;
步骤2)中粪便的采集为选择断奶后2个月健康的杜长大商品猪和金华猪各6头,所有猪只饲养管理一致;采食后同一时间段内从猪直肠中收集粪便样本,并将其转移到无菌冻存管中,所有操作均在厌氧条件下进行,液氮处理后在-80℃进行冷冻储存;
步骤3)中的培养基配方为:0.16 g/L 蛋白胨,0.1 g/L 酵母浸出物,0.16 g/L 吐温80,0.16 g/L NaHCO3,3.6 g/L NaCl,1.6 g/L K2HPO4,0.32 g/L L-半胱氨酸盐酸盐,0.36g/L CaCl2·6H2O,0.5 g/L MgSO4·7H2O,0.01 g/L 血红素;
步骤4)包括:将1 g预消化处理的阿拉伯木聚糖溶于25 mL无菌发酵培养基中,100℃煮沸10 min;随后将其放入厌氧室中,冷却到室温,放置2 h,得到待发酵的阿拉伯木聚糖溶液;0.1 g粪便样本添加到5 mL的无菌培养基中,振荡涡旋混匀至均质状态得到待接种的粪菌悬液;将2.5 mL粪菌液与2.5 mL发酵底物溶液混合后得到最终发酵体系,37°C厌氧条件下以130 rpm振荡培养;在发酵的第1、3、6、9、12、15、18、21、24、48和72 h进行采样;采取冰浴终止发酵,将发酵液离心,上清液和沉淀物于80℃冻存;
所述的方法,测量各时间点pH值、气相色谱法定量分析短链脂肪酸含量、ELISA法测定α-L-阿拉伯呋喃糖苷酶和β-木聚糖酶活性;提取各个时间点发酵液离心后沉淀物中基因组DNA,所有样品均使用适用于细菌16S rDNA 的V3-V4区域进行PCR扩增,利用IlluminaMiSeq平台测序;得到的以上数据,作为评估阿拉伯木聚糖对肠道微生态影响的指标;筛选得到不同时间点阿拉伯木聚糖靶向促进生长的微生物种类。
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