CN117562189B - 维生素k在制备调控肉鸡肠道菌群饲料中的应用 - Google Patents
维生素k在制备调控肉鸡肠道菌群饲料中的应用 Download PDFInfo
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Abstract
本发明公开了维生素K在制备调控肉鸡肠道菌群饲料中的应用。本发明通过在肉鸡饲料中添加维生素K2或K3，证实可提高回肠肠道菌群的丰度，所述菌群包括Bacteroide菌，Bifidobacterium‑animalis菌和Ruegeria菌。通过在饲粮中添加维生素K2可以改善LPS诱导的肉鸡全身性炎症、肠道损伤和回肠微生物菌群失调，抑制有害菌在肠道定植，增加有益菌丰富度，保护肠道杯状细胞从而降低炎症，改善肠道屏障功能。其通过增加肉鸡回肠微生物多样性和缓解LPS注射诱导的菌肠失调，而在肠道疾病和骨代谢失调方面发挥治疗作用。
Description
技术领域
本发明涉及生物技术领域，具体涉及维生素K在制备调控肉鸡肠道菌群饲料中的应用。
背景技术
肠道是人和动物消化吸收的场所，也是最大的免疫系统。肠道健康保证了动物正常生长与生产，为抵御病原菌提供了免疫屏障。在当下禁用抗生素的实际生产背景下，饲料营养、疾病和饲养管理可能导致肠道问题更加突出。
肉鸡规模化集约化养殖模式解决了我国人民吃上肉的问题，但是密集饲养导致炎症发生频率加剧。感染、致病微生物和环境污染等多种因素可导致家禽的免疫应激和氧化应激。即使在正常饲养情况下，家禽也不可避免地面临着与细菌或其代谢产物(如脂多糖Lipopolysaccharide，LPS)相关的免疫或氧化应激等挑战，危害健康。因此，在家禽体内注射LPS已被用作研究家禽业系统性炎症和氧化应激的良好模型。已有研究表明，饲粮中添加维生素、合成抗氧化剂、天然植物提取物等饲料添加剂可以缓解LPS诱导的对肉鸡产生的负面影响。因此，寻找一种能够调节免疫和抗氧化系统且平衡菌群以保护肉鸡免受免疫应激影响的新型、有效的饲料添加剂具有重要的意义。
维生素K2(VK2)是一系列含有2-甲基-1,4-萘醌环及C3位带有数目不等异戊二烯结构单元的萜烯侧链化合物的统称。VK2又被称为甲萘醌(menaquinones)，MK-n为甲萘醌系列化合物的缩写，n代表异戊二烯侧链长度，范围为1 ~ 13。在所有的甲萘醌中，对MK-4和 MK-7的研究最充分。最近发表的一篇综述中提供了关于天然VK的详细信息。机体对VK的摄入主要包括VK1、MK-4和MK-7(分别大于60%、24%和7%)。近90%的MKs和10%的VK1通常分别储存在肝外组织和肝脏中，VK的生理需求量约为1.16 µg/kg 体重。
作为一种微量营养素，关于VK在肠道健康中免疫调节作用的新证据表明，VK在肠道疾病健康中具有超出其在止血中的经典功能以外的新作用。当病人出现肠道损伤、胆汁淤积性肝病、或遗传疾病、抗生素使用和抗凝血剂或任何脂肪吸收不良的症状会导致VK缺乏。VK缺乏时，由于缺乏γ -羧谷氨酸而出现异常凝血酶原，可导致严重机体出血和死亡。在肠炎患者中，VK缺乏是由于肠道损伤引起的吸收不良所致。VK缺乏在慢性胃肠疾病中也有报道，包括炎症性肠炎和短肠综合症。
在肉鸡实际生产中，肉鸡饲料中通过添加VK3为肉鸡提供维生素K。添加高剂量的VK3已被证明会导致潜在的毒性，如肝损伤和溶血性贫血，而且在一些国家，它只被允许作为宠物饲料的添加剂。20世纪90年代开始的研究表明，VK3是口服VK1在肠道中产生的分解代谢产物，可以输送到组织，随后转化为VK2。
最近研究发现，VK缺乏或添加对小鼠肠道菌群的影响已有少量报道。但是在肉鸡肠炎情况下，VK这种菌群调控作用是否存在尚不可知。
发明内容
因此，本发明探究了日粮添加维生素K2对肉鸡肠炎的缓解效果，揭示VK2对肠炎的免疫调控和屏障功能保护的调控作用及机制。
因此，本发明的目的在于针对现有技术中对于肉鸡肠炎的治疗技术不足的问题，提供一种能够调节肉鸡肠道菌群的方法，以应用于治疗和改善肉鸡肠道炎症。
为解决上述技术问题，本发明采用如下技术方案：
本发明公开了维生素K在制备调控肉鸡肠道菌群饲料中的应用，所述菌群包括：Bacteroide菌，Bifidobacterium-animalis菌和Ruegeria菌。
优选的，所述维生素K选自维生素K2和维生素K3中的一种或两种。
优选的，所述调控为提高菌群丰度。
优选的，所述维生素K2提高Bacteroide菌和Bifidobacterium-animalis菌的丰度。
优选的，所述维生素K2的含量为2mg/kg至4mg/kg，优选为3mg/kg饲料。
优选的，维生素K3提高Ruegeria菌的丰度。
优选的，所述维生素K3的含量为2mg/kg至4mg/kg，优选为3mg/kg饲料。
优选的，所述肠道菌群为回肠菌群。
优选的，所述饲料的配方为：玉米、面粉、豆粕、面粕、豆油、玉米面筋粉、碳酸钙、碳酸氢钙以及预混料。
优选的，所述肉鸡为患肠炎的肉鸡。
饲粮中添加VK2可以改善LPS诱导的肉鸡全身性炎症、肠道损伤和微回肠微生物菌群失调。VK2能缓解肉鸡全身性炎症反应和肠炎，抑制有害菌在肠道定植，增加有益菌丰富度，保护杯状细胞从而降低炎症，改善肠道屏障功能。
VK可能通过增加肉鸡回肠微生物多样性和缓解LPS注射诱导的菌肠失调，而在肠道疾病和骨代谢失调方面发挥治疗作用。由此可见，饲粮中添加VK通过调节LPS刺激下肉鸡炎症细胞因子的表达和肠道微生物菌群结构，对肉鸡炎症损伤和免疫应激产生保护作用。
附图说明
图1.日粮添加VK对LPS应激11d肉鸡生长性能的影响，其中A为肉鸡9-11d的生长性能；B为肉鸡1-11d的生长性能。
图2.11日龄肉鸡血清ucOC与ucMGP含量(n=6)。
图3.11日粮添加VK对LPS应激11d肉鸡免疫器官及炎症的影响(n=6)，其中，A为法氏囊和脾脏的相对重量；B为脾脏TNFα和IL-6的相对表达量。
图4.日龄肉鸡肝脏损伤，其中A为肝脏相对重量和肝脏COX-2的相对表达量；B为血清ALT和AST的酶活。
图5.11日龄肉鸡血清免疫蛋白和C反应蛋白水平，其中，A为血清TP、ALB和GLO的含量；B为血清免疫球蛋白IgY和C-反应蛋白（CRP）的含量。
图6.VK对LPS攻毒肉鸡的回肠菌群α-多样性的影响，其中，A为 Chao1指数；B为辛普森指数；C香农指数；D谱系多样性。
图7.门水平回肠菌群结构，其中，A为微生物门水平上相对丰度；B为厚壁菌门和拟杆菌门的相对丰度。
图8.属水平回肠菌群结构，其中，A为微生物门水平上相对丰度；B为拟杆菌属普拉梭菌属的相对丰度。
图9.LefSe分析的LDA分布状图和LefSe分析进化分支图，其中，A为LefSe分析的LDA分布状图；B为 LefSe分析进化分支图。
具体实施方式
以下结合附图对本发明的具体实施方式进行详细说明。应当理解的是，此处所描述的具体实施方式仅用于说明和解释本发明，并不用于限制本发明。
实施例1：试验动物的分组设计
本试验中的肉鸡饲养管理和样本采集程序遵守中国农业大学试验动物指南（伦理审查编号AW80302202-1-1）。
本试验在中国农业大学涿州基地进行。试验选取健康有活力的 1 日龄爱拔益加肉公鸡 264 只，随机分为4个处理组：生理盐水对照组、脂多糖(LPS)处理组、VK2添加组（3mg MK-7/kg）和VK3（3 mg /kg）添加组，每组 6 个重复，每个重复66只。试验期长11d。试验动物按照肉鸡饲养管理方法进行笼养，自由采食饮水，进行常规免疫接种。试验第9、11日龄，空腹进行腹腔注射1mg/kgBW的LPS 3 h后采样。
实施例2：试验基础日粮的配制以及营养水平的分析
试验基础日粮参照中国鸡饲养标准（2004）配置玉米-豆粕型日粮，试验基础日粮组成及其营养水平见表1。试验每个阶段的基础日粮一致，根据试验设计将VK2和VK3加入各组日粮中。
表1：肉鸡日粮成分和营养组成(风干基础)。
1每千克日粮微量元素含量： Cu, 8 mg (CuSO4·5H2O)； Zn 100 mg (ZnSO4·H2O)；Fe， 80 mg (FeSO4)；Mn， 100mg (MnSO4·H2O)； Se， 0.15 mg (Na2SeO3)； I， 0.35mg (KI)。
每千克日粮维生素含量：维生素 A 9500 IU；维生素D362.5ug； 维生素E 30 IU(α-生育酚)；维生素B12 mg；维生素B120.025 mg；维生素B26 mg；尼克酸50 mg；叶酸1.25mg；生物素0.0325 mg；泛酸12 mg。
2营养水平：根据NRC(1994)计算。
实施例3：日粮添加维生素K2对肉鸡生长性能的影响
参见图1，与生理盐水对照组比，腹腔注射LPS显著降低了LPS处理期间(9-11d)的体增重（P<0.001）和采食量（P<0.05），显著增加了FCR（P<0.05）；添加VK对BWG、FI和FCR影响不显著（P>0.05）。腹腔注射LPS有降低整个试验期间 (1-11d)的体增重的趋势（P=0.061），显著降低了采食量（P<0.05），对FCR没有显著影响（P>0.05）；添加VK对BWG（体增重（bodyweightgain））、FI（采食量（feedintake））、FCR（饲料转化率（feedconversionratio））影响不显著（P>0.05）。
实施例4：日粮添加维生素K2对肉鸡血清VK水平的影响
羧化不全基质γ-羧基谷氨酸蛋白(ucMGP)和骨钙素(ucOC)是间接反应体内VK水平的重要指标，添加VK2能提高肉鸡MGP和OC的羧化程度，反应体内VK水平升高。按照鸡ucOC(JK45112)和ucMGP(JK85113)酶联免疫吸附测定试剂盒(上海酶联生物技术有限公司)说明书测定血清ucOC和ucMGP浓度。在试验室中使用多光谱微孔板分光光度计(Spectra Maxi3x)在波长450 nm处读取吸光度。结果参见图2。
与LPS组相比，日粮添加VK2极显著降低了血清ucOC水平（P<0.01）。与CON组和LPS组相比，日粮添加VK2显著（P<0.05）和极显著（P<0.001）降低了血清ucMGP水平; 与LPS组相比，日粮添加VK3极显著降低了血清ucMGP水平（P<0.01）。
实施例5：日粮添加维生素K2对肉鸡免疫器官及炎症的影响
采样脾脏、肝脏、回肠和空肠样品液氮东西冻存后置于-80℃储存。按照说明书使用Trizol试剂(Genestar)从脾脏、肝脏和空肠(50mg)中提取总RNA。紫外分光光度计(NanoDrop-2000, Thermo Fisher Scientific)中260:280 nm的吸光度比用于估计总RNA的浓度和纯度(OD260/OD280: 1.8-2.0)。总RNA保存于-80℃，合成的cDNA保存于-20℃。引物序列见表2。采用ABI7500 RT-PCR 仪器进行测定，PCR反应参数设置为:95℃，5分钟一个周期，然后95℃，30秒40个周期，60℃ 30秒。每个样本3个重复。以 β-actin 为内参，进行相对定量计算，结果参见图3。
表2 目的基因序列
LPS导致了肉鸡法氏囊显著萎缩（P<0.05）和脾脏极显著肿大（P<0.001），表明腹腔注射LPS对肉鸡免疫器官产生了损伤。添加VK2和VK3对LPS诱导的胸腺萎缩没有显著影响（P>0.05）。同时，肉鸡脾脏的TNFα和IL-6基因表达结果显示，LPS极显著增加了脾脏TNFα（P<0.001）和IL-6的表达P<0.05）。日粮添加VK2极显著降低了TNFα（P<0.001）和IL-6（P<0.01）的表达上调，日粮添加VK3极显著降低了TNFα的表达上调（P<0.001），对IL-6的表达上调没有显著影响（P>0.05）。
实施例6：日粮添加维生素K2对肉鸡的免疫肝脏损伤和全身炎症影响
肝脏健康相关指标谷丙转氨酶(ALT)、谷草转氨酶(AST)及血清脂质水平如胆固醇(CHO)、甘油三酯(TG)、低密度脂蛋白(LDL)及高密度脂蛋白(HDL) 采用试剂盒(北京九强生物技术股份有限公司)于全自动生化分析仪检测。
根据鸡CRP (SEKCN-0100)、IgG (SEKCN-0126)和IgA (SEKCN-0018)酶联免疫吸附测定试剂盒(索莱宝生物科技有限公司)说明书，测定血清CRP、IgG和IgA含量。在试验室中使用多光谱微孔板分光光度计(Spectra Max i3x)在波长450 nm处读取吸光度。
与对照组比，肉鸡腹腔注射LPS导致肝脏肿大，肝脏器官指数极显著增加（P<0.01），日粮添加VK2显著缓解了肝脏肿大（P<0.05）;日粮添加VK3没有显著缓解肝脏肿大（P>0.05）。COX-2是触发后续炎症反应的关键环节，LPS导致肝脏COX2表达极显著上调，日粮添加VK2和VK3显著下调了COX2表达上调。临床上常用血清中天冬氨酸转氨酶(aspartateaminotransferase，AST)、丙氨酸转氨酶(alanine aminotransferase，ALT)活性作为反映肝损伤的具体指标。LPS导致血清AST（P<0.05）和ALB（P<0.01）水平显著增加，日粮添加VK2（P<0.001）和VK3（P<0.05）显著降低了血清AST水平。参见图4。
LPS显著增加了血清TP、GLO、IgY和CRP水平（P<0.05），VK2显著降低TP、IgY和CRP水平（P<0.05）；VK3对肉鸡血清免疫蛋白和C反应蛋白水平没有显著影响（P>0.05）。参见图5。
实施例7：日粮添加维生素K2对肉鸡的肠道微生物分析
肉鸡回肠食糜的16rDNA测序试验由北京百迈克生物科技有限公司对所有食糜样品进行微生物V3-V4区序列16srDNA扩增子测序分析。利用16rDNA V3-V4为靶标扩增的Illumina 标准 16S 元组测序文库，制备 16S rDNA 序列进行微生物区系分析, 使用Illumina Novaseq 6000 进行配对的末端测序(300 bp)，生成 200000 个序列/样本。随后经过 QIIME 软件根据 Barcode 序列拆分样本，分别使用Trimmomatic v0.3、Usearchv10、UCHIME v4.2软件分别对数据进行质量过滤、双端序列拼接、去除嵌合体，经vsearch软件去除长度小于 230 bp的序列，并根据已知数据库用uchime方法比对去除嵌合体序列。使用Usearch软件对Reads在97.%的相似度水平下进行聚类，获得OTU。使用 Silva128 16SrRNA 数据库(细菌和古菌)对每个 OTU 进行注释。Illumina Novaseq测序生成数据fastq文件上传至 NCBI。
通过α-多样性分析评估VK对注射LPS诱导炎症肉鸡回肠菌群物种丰富度和群落多样性影响。如图6所示，各组肉鸡之间回肠菌群的Chao1、Shannon和PD whole tree指数没有显著差异（P>0.05）。结果表明VK3和VK2具有增加回肠菌群物种丰富度和群落多样性的作用。
肉鸡回肠微生物在门水平变化如图7所示，4 个试验组共同的优势门水平细菌是厚壁菌门（Firmicutes）和拟杆菌门（Bacteroidetes），其次是变形菌门（Proteobacteria)。与 Con 组相比，LPS 组的厚壁菌门（Firmicutes）显著降低（P＜0.05），拟杆菌门（Bacteroidetes）显著增加（P＜0.05）。与 HF 组相比，VK3和V K2 组Firmicutes显著增加（P＜0.05），Bacteroidetes 显著降低（P＜0.05）。结果表明，LPS注射改变了回肠菌群的结构，而VK可以不同程度缓解LPS注射导致的肉鸡菌群失调。
在属水平上，与 Con 组相比，LPS 组Faecalibaculum粪棒状菌属和Anaerofilum厌氧细杆菌属相对比例显著降低（P＜0.05）；与LPS组比，VK3和VK2显著增加相对比例（P＜0.05）。与 Con 组相比，LPS 组Bacteroides 拟杆菌属降低显著增加（P＜0.05）；与LPS组比，VK3和VK2显著降低（P＜0.05）。参见图8。
图9A 显示了基于LEfSe分析的 LDA 分布柱状图，表明不同组中丰度有显著差异（LDA＞3，P＜0.05）的生物标志物，图中柱状图长度代表显著差异的生物标志物的影响程度。结果发现，Con 组、LPS组、VK3和VK2 组各有 2、11 、10和 3 个具有统计学差异的生物标记物（LDA＞3，P＜ 0.05）。
通过LEfSe分析进化分支图进一步确定与VK缓解作用密切相关的差异生物标记物信息。如图9B 可示，LEfSe进化分支图中由内至外辐射的圆圈代表由门至属的分类级别。每一个小圆圈代表该分类水平下的一个分类，其直径大小与相对丰度大小正相关。无显著差异的物种显示黄色，试验中各组的差异生物标记物分别显示为红色（Con组）、绿色（LPS组）、蓝色（VK2组）和紫色（VK3组）。结果显示，在 Con组中，Enterobacteriales目和Enterobacteriaceae科是最显著的生物标记物，在 LPS 和VK3组肉鸡肠道菌群含量显著升高，而在 VK2组回肠菌群含量显著降低。这表明早期注射LPS的肉鸡回肠菌群经过11d恢复后，肠道菌群丰富度提高，其中Bacteroidales目、Bacteroidaceae科、Bacteroide属、和Bacteroide菌种以及Bifidobacteriales目、Bifidobacteriaceae科、Bifidobacterium属和Bifidobacterium-animalis菌种；添加VK3增加了Bacterium-YE57菌种、Faecalibacterium科、Ruminococcaceae目、Bacterium属、Ruegeria菌种和Ruegeria属的丰度；而VK2添加使菌群更接近对照组，增加了Clostridiales梭菌目的丰度。
通过16S rRNA Illumina Miseq高通量测序对肉鸡回肠菌群结构进行α-多样性和β-多样性分析，结果表明LPS降低了肉鸡回肠的Simpson多样性（主要用于估算样本微生物多样性），而添加两种VK源恢复了LPS导致的α-多样性降低。同时通过PCA、PCoA在OUT水平对肉鸡回肠微生物群落信息进行可视化，解释四个组试验组菌群结构的相异性。结果显示，LPS组的肉鸡回肠菌群整体结构与其他三组之间表现出显著的差异性，反映LPS对肉鸡回肠菌群产生了长时间的整体失调的影响。PCA和PCoA结果显示添加两种VK源使肉鸡肠道菌群恢复与对照组接近。而PLS_DA结果表明，四组肉鸡回肠菌群结构发生了明显的差异，聚集在不同的区域。VK可能通过增加肉鸡回肠微生物多样性和缓解LPS注射诱导的菌肠失调，而在肠道疾病和骨代谢失调方面发挥治疗作用。由此可见，饲粮中添加VK通过调节LPS刺激下肉鸡炎症细胞因子的表达和肠道微生物菌群结构，对肉鸡炎症损伤和免疫应激产生保护作用。
本发明通过上述实施例来说明本发明的工艺方法，但本发明并不局限于上述工艺步骤，即不意味着本发明必须依赖上述工艺步骤才能实施。所属技术领域的技术人员应该明了，对本发明的任何改进，对本发明所选用原料的等效替换及辅助成分的添加、具体方式的选择等，均落在本发明的保护范围和公开范围之内。
Claims (6)
1.维生素K在制备提高肉鸡肠道菌群丰度的饲料中的应用，其特征在于，所述肠道菌群为回肠菌群；
且所述维生素K是维生素K3，其用于增加肉鸡肠道菌群的Ruegeria 属的丰度；或者所述维生素K是维生素K2，其用于增加肉鸡肠道菌群的Clostridiales 梭菌目的丰度。
2.根据权利要求1所述的应用，其特征在于，饲料中所述维生素K2的含量为2mg/kg至4mg/kg。
3.根据权利要求2所述的应用，其特征在于，饲料中所述维生素K2的含量为3mg/kg。
4.根据权利要求1所述的应用，其特征在于，饲料中所述维生素K3的含量为2mg/kg至4mg/kg。
5.根据权利要求4所述的应用，其特征在于，饲料中所述维生素K3的含量为3mg/kg。
6.根据权利要求1至5任一项所述的应用，其特征在于，所述饲料的配方为：玉米、面粉、豆粕、面粕、豆油、玉米面筋粉、碳酸钙、碳酸氢钙以及预混料。
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