蒲公英(Taraxacum officinale),俗称婆婆丁,又名华花郎、黄花三七、白鼔丁等,属于菊科蒲公英属,是一种多年生草本植物,主要分布在南北半球温带地区,原产于欧洲,在我国东北、华北、西北、华中、西南地区均有分布[1-3]。蒲公英的主要化学成分有多糖类、黄酮类、三萜类、倍半萜内酯类、有机酸类、香豆素、挥发油类等物质,具有抗氧化、抗菌消炎、保肝利胆、降血糖、抗肿瘤、增强免疫力等生物学功能[4-5]。蒲公英全草可入药,但其根、叶和花所含的有效成分有所不同,蒲公英根中多糖含量较高,可达83.31%,远高于叶和花[6],不同研究中所测得的多糖含量有所差别,总体上来说多糖含量根>花>叶[7];叶中酚酸和类黄酮含量丰富,其中,对香豆酸、咖啡酸、木犀草苷含量均达到根中的100倍以上;叶中木犀草素含量达200 μg/g以上,而根中木犀草素含量则为0[8]。研究表明,同一成分在不同产地、不同药用部位中的含量具有明显的差异[9],因而其药理效果也有所不同。此外,提取方法不同,提取物的药理作用也会有所不同[10]。随着我国禁抗时代的到来,开发各种可应用的中草药作为饲料添加剂以替代抗生素的作用成为必然发展趋势。我国蒲公英分布广泛、资源丰富、廉价易得,在畜、禽、鱼类饲养中有明显效果,具有良好的开发应用前景。本文对蒲公英根、叶、花提取物的生物学功能及其在畜、禽、鱼类生产中的应用进行了综述,为蒲公英的开发利用和在动物生产上的应用提供参考。
1 蒲公英提取物的生物学功能 1.1 抗氧化功能活性氧(reactive oxygen species,ROS)和活性氮(reactive nitrogen species,RNS)是代谢途径中产生的2类主要的自由基,生物体内的自由基不断产生和消除,始终维持动态平衡。自由基如果产生过多或者消除过慢而在细胞、组织、器官积累时,就会造成氧化失衡和机体损伤,进而导致一些疾病如癌症、肝脏疾病、类风湿性关节炎、动脉粥样硬化等的发生,并参与衰老等退行性过程[11-14]。蒲公英多糖能上调锰超氧化物歧化酶(Mn-superoxide dismutase,Mn-SOD)、铜锌超氧化物歧化酶(Cu-Zn superoxide dismutase,Cu-Zn-SOD)、谷胱甘肽过氧化物酶-1(glutathione peroxidase-1,GPX-1)、谷胱甘肽过氧化物酶-4(glutathione peroxidase-4,GPX-4)等氧化应激相关基因的相对表达量,提高抗氧化酶的活性,有效清除自由基,避免其对机体的损伤[15]。Guo等[16]采用超声辅助提取法,使用高效液相分离纯化得到蒲公英中的水溶性多糖(polysaccharides from dandelion,PD),具有良好的1, 1-二苯基-2-三硝基苯肼(1, 1-diphenyl-2-trinitrobenzene hydrazine,DPPH)自由基清除活性,能有效地清除生物体内的羟基自由基,具有较强的抗氧化活性。除蒲公英多糖外,Huang等[11]采用醇沉超声提取法提取蒲公英黄酮,蒲公英黄酮也具有抗DPPH活性,能够抑制氨酸酶的活性,快速清除自由基,在氧化反应中,黄酮能够在链转移阶段和阻断链式反应中直接捕获和清除过氧自由基,起到抗氧化的作用[17]。Liu等[18]采用响应面法对蒲公英发酵条件进行优化,使发酵蒲公英粗提物(extract from fermented dandelion,FDF)中黄酮类化合物含量达到(183.72±2.24) mg/g,使FDF在体外DPPH自由基清除能力和还原能力方面表现出更好的活性。酚类物质也能起到抗氧化作用,Hu等[19]研究表明,蒲公英花提取物(dandelion flower extract,DFE)与α-生育酚的协同作用及自由基的清除能力与酚类物质的含量有关。DFE的抗氧化活性能够防止活细胞受到过氧自由基诱导的氧化。蒲公英中的倍半萜内酯(sesquiterpene lactones,SLs)对血浆脂质和蛋白质具有保护作用,抵抗过氧化氢(H2O2)/铁(Fe)的氧化作用[20]。Choi等[21]研究表明,2 ∶ 1(质量比)混合的柠檬香脂和蒲公英提取物(lemon balm and dandelion extracts,LD)能够清除DPPH自由基,保护HepG2细胞免受叔丁基过氧化氢(tert-butylhydroperoxide,tBHP)介导的氧化应激。LD在增强生物体抗氧化能力的同时,还能降低核因子E2相关因子2(nuclear factor E2-related factor 2,Nrf2)的表达水平,从而显著降低乙醇诱导的肝脏氧化应激反应,降低肿瘤坏死因子-α(tumor necrosis factor-α,TNF-α)含量,保护肝脏免受乙醇的影响。多酚可以通过阻止自由基的形成或解毒来保护DNA、蛋白质和脂类免受氧化应激,从而防止各种病理生理过程[22]。蒲公英水提物(dandelion aqueous extract,DAE)可以抑制脂多糖(lipopolysaccharide,LPS)诱导的氧化应激,并且是Nrf2信号通路的潜在诱导剂,是一种有效的抗氧化剂[23]。
1.2 抗菌抗炎功能蒲公英具有良好的抗炎作用,其活性成分是蒲公英寡肽(dandelion oligopeptides,DOPs),可有效抑制LPS诱导的炎性细胞因子过表达,并抑制细胞外调节蛋白激酶(extracellular regulated protein kinases,ERK)1/2、磷脂酰肌醇3-激酶(phos-phatidylinositol 3-kinase,PI3K)、c-Jun氨基末端激酶(c-jun N-terminal kinase,JNK)和蛋白激酶B(protein kinase B,Akt)的磷酸化以及核因子-κB(nuclear factor-kappa B,NF-κB)的活化[24]。Xue等[3]研究表明,在体外培养的人结肠上皮细胞中,蒲公英叶提取物(dandelion leaf extract,DLE)中的菊苣酸能有效抑制肠道氧化应激和炎症小体HT-29细胞的形成。Choi等[21]研究表明,LD抑制LPS刺激的RAW264.7细胞中促炎介质的产生可能是由于Nrf2的激活和LPS依赖的NF-κB为中心的信号通路的抑制。蒲公英多糖和寡糖(dandelion-derived oligosaccharides,DOs)具有广谱抗菌作用,对金黄色葡萄球菌、大肠杆菌和枯草芽孢杆菌等细菌均具有较强的抑制活性[25-26]。Kenny等[27]研究证明,蒲公英根提取物(dandelion root extract,DRE)对金黄色葡萄球菌和蜡样芽胞杆菌有很好的抑菌作用。Yang等[28]研究表明,重金属可诱导细菌产生抗生素耐药性,DRE和蒲公英根中主要天然成分蒲公英甾醇(taraxasterol,TS)通过抑制β-内酰胺酶或乙酰转移酶活性来帮助逆转镍、镉、砷引起的细菌抗生素耐药性。Yang等[29]研究发现,蒲公英及其成分蒲公英醇具有对乙型肝炎B病毒(hepatitis B virus,HBV)的抑制作用,其重要的结合位点为宿主因子多聚嘧啶区结合蛋白1(polypyrimidine tract binding protein 1,PTBP1)和组蛋白去乙酰化酶1(sirtuin 1,SIRT1)。
1.3 抗肿瘤功能蒲公英具有抗癌作用,研究表明DRE能诱导侵袭性和耐药的人慢性粒细胞白血病细胞(chronic myelomonocytic leukemia,CMML)的凋亡[30]。DLE对乳腺癌细胞系MCF-7/AZ细胞的增殖有抑制作用,而DRE和DLE分别阻断了人乳腺癌细胞系MCF-7/AZ和人前列腺癌LNCaP细胞的侵袭[31]。Yarnell等[32]研究表明,蒲公英具有选择性诱导人类黑色素瘤和白血病细胞凋亡的能力,对非癌性外周血单核细胞无毒性。Ovadje等[33]证明,DRE有诱导人胰腺癌细胞凋亡和自噬的潜力,而对非癌细胞没有显著影响。满瑾等[34]试验结果表明,蒲公英提取物可以影响能量及多种氨基酸代谢,从而扰乱A549细胞的生长和增殖,达到抑制A549细胞活力的作用。Ovadje等[35]研究表明,DRE水溶液依赖剂量和时间诱导人白血病细胞凋亡,并且DRE中含有可选择性诱导白血病细胞凋亡的成分,为白血病的治疗提供了一种新的可能。Chatterjee等[36]发现,DRE能特异且有效地诱导人类黑色素瘤细胞(A375)凋亡,而不会对非癌细胞产生毒性。Menke等[37]研究表明,所有癌细胞株对蒲公英的敏感性均高于正常人成纤维细胞(normal human fibroblast cell line,NHDF-C),在神经母细胞瘤细胞系中,蒲公英能够引起细胞凋亡,抑制癌细胞的迁移,导致线粒体完整性的丧失,当与槲寄生提取物同时使用时,表现出协同效应。Duan等[38]研究表明,DRE对食管鳞状细胞癌(esophageal squamous cell carcinoma,ESCC)具有抑制作用,其原理是通过调控PI3K/Akt、Ras/Raf/ERK信号通路以及内源性CBS/H2S系统来影响ESCC的进展,诱导细胞凋亡,选择性地抑制ESCC细胞的生长、增殖、迁移和侵袭。Zhang等[39]研究表明,从蒲公英根中分离得到的菊粉果聚糖,体外评价其具有对肿瘤细胞增殖中等或弱的抑制作用。化疗是目前治愈癌症的主要方法之一,但是其副作用及肿瘤细胞耐药性的产生极大地限制了化疗在肿瘤治疗领域的应用,DRE、DLE具有抗肿瘤活性,并且无副作用,可作为一种抗肿瘤的备选药材进行合理的开发与利用。
1.4 其他功能除上述生物学功能外,蒲公英提取物还具有保护肝脏、增强免疫力、降血糖等生物学功能。Cai等[40]研究表明,从蒲公英根多糖(dandelion root polysaccharide,DRP)中分离得到2种α型多糖DRP1和DRP2,这2种多糖可以阻止对乙酰氨基酚(n-acetyl-p-aminophenol,n-APAP)诱导的肝脏氧化损伤。DRP通过降低抑制蛋白Kelch样环氧氯丙烷相关蛋白(Kelch-like epichlorohydrin-associated protein 1,Keap1)的表达而激活Nrf2,从而增强下游抗氧化蛋白血红素氧合酶-1(heme oxygenase-1,HO-1)和依赖还原型辅酶Ⅰ(Ⅱ)醌氧化还原酶1[NAD(P)H: quinine oxidoreductase 1,NQO1]的表达,起到抗氧化作用。蒲公英中提取的黄酮可以对胰腺α-淀粉酶具有抑制作用,起到降低餐后血糖的作用[41]。小鼠在疲劳状态下,蒲公英可以降低小鼠乳糖脱氢酶活性,提高葡萄糖含量,说明蒲公英可以减少小鼠肌肉损伤,缓解疲劳,为小鼠提供更多的能量来源,并且在小鼠腹腔巨噬细胞中,蒲公英可以增加TNF-α、白细胞介素-12p70(interleukin-12p70,IL-12p70)、白细胞介素-10(interleukin-10,IL-10)以及一氧化氮(NO)含量,具有增强免疫力的作用[42]。蒲公英根中的次级代谢产物氨基酸及其衍生物和4-羟基苯乙酸肌醇酯(hydroxyphenylacetate inositol esters,PIEs)对血浆中H2O2/Fe氧化的整体保护效果最好,对超氧阴离子(O2-)抑制效果最好,抗血小板潜力最佳,而羟基肉桂酸(hydroxycinnamic acids,HCAs)的抗凝效果则更强[20, 43]。
2 蒲公英及其提取物在畜、禽、鱼类生产中的应用 2.1 在猪生产中的应用饲粮中添加蒲公英干粉或提取物能提高猪的免疫力,增强抗病性。李玉荣[44]研究发现,饲粮中添加500 g/t蒲公英干粉可以提高肥育猪平均日增重和平均日采食量,同时可以显著降低体内甘油三酯和总胆固醇含量及腹泻率。王留等[45]试验证明,饮用水中添加6 g/L的DAE可以降低断奶仔猪腹泻率,并且也能够降低血清中促炎性细胞因子白细胞介素-22(interleukin-22,IL-22)和TNF-α含量,增加血清中抗炎因子IL-10含量。饲粮中添加蒲公英中草药合剂对猪链球菌病具有一定的防治作用,能够降低肥育猪的得病率[46]。体外研究表明,蒲公英对猪链球菌具有抑制作用[47]。蒲公英提取物对猪胸膜肺炎放线杆菌的抑菌浓度为14.6 g/L,对猪链球菌的抑菌浓度为9.3 g/L,对多杀性巴氏杆菌的抑菌浓度为11.7 g/L[48]。于文会等[49]通过水提法提取得到蒲公英中的绿原酸,发现该提取物对猪链球菌生物被膜的形成具有抑制作用。杜学林[50]研究表明,仔猪饲粮中添加1%蒲公英干粉对提高猪群平均日采食量、平均日增重有明显效果,同时还会降低疾病发生率,主要是由于蒲公英所含的酮类、酸类和糖类起到提高猪群免疫力的作用。Zhao等[51]试验结果表明,饲粮中添加DRE可以提高断奶仔猪粪便中丁酸含量,并且能够改善免疫功能,同时粪便菌群的相对丰度和组成也发生了相应的变化。
2.2 在家禽生产中的应用饲粮中添加蒲公英能改善肉鸡的生长性能,提高其免疫力。饲粮中添加蒲公英叶和葫芦巴籽对肉鸡肠道形态有一定的保护作用,同时在肉鸡生长期间添加蒲公英叶片可以改善肉鸡的生长性能[52-53]。饲粮中同时添加冻干大蒜(5 g/kg)和蒲公英根粉(50 g/kg)可提高肉鸡生长性能,改善胴体性状和营养脂肪品质[54]。蒲公英能促进乌骨鸡新城疫病毒(newcastle disease virus,NDV)抗体的产生和胸腺发育,抑制脾脏发育[55]。饲粮中添加2.0%的蒲公英可以提高动物产品质量和品质以及免疫抗体效价和免疫器官发育,从而增强肉鸡新城疫免疫效果[56]。邓海涛[57]研究发现,肉鸡饲粮中添加1%的蒲公英粉剂对改善肉鸡的生长性能、提高抗病能力起到很好的效果。王树艳等[58]研究结果显示,饮用水中添加5%、10%的蒲公英提取物可以提高三黄鸡胸腺指数、脾脏指数、免疫后抗体水平和免疫功能,提高抗禽流感水平。王留等[59]研究表明,饮水中添加DAE(3~5 g/L)可以显著提高肉仔鸡外周血淋巴细胞转化率和血清中免疫球蛋白(IgG)含量,同时血清中的总抗氧化能力(T-AOC)以及谷胱甘肽过氧化物酶(GSH-Px)和超氧化物歧化酶(SOD)活性显著提高,丙二醛(MDA)含量显著降低,肉仔鸡的整体免疫功能和抗氧化能力均有所提高。
2.3 在反刍动物生产中的应用开发中草药作为新型添加剂对瘤胃微生物群落、瘤胃发酵、动物生产和健康状况均具有深远影响。Li等[60]试验结果表明,饲粮中添加蒲公英可以通过改善奶牛瘤胃厚壁菌门的相对丰度和瘤胃代谢途径来促进瘤胃发酵。蒲公英提取物对金黄色葡萄球菌和大肠杆菌都有抑制作用,能够有效地减少奶牛隐性乳房炎的发生[61]。在奶牛饲粮中添加蒲公英提取物(0.5%)可以缓解其氧化应激并改善泌乳性能[62]。在精料中添加6.0%的蒲公英可以促进放牧肉牛增重、血清球蛋白含量升高,改善肉牛血液代谢指标和免疫力[63]。
以上研究表明,蒲公英提取物能够通过改善瘤胃微生物,抑制奶牛乳房炎的发生,提高奶牛抗氧化能力,改善肉牛生长性能。反刍动物饲粮中添加蒲公英的研究还比较少,其提取物在精料中的应用还有待于进一步研究。
2.4 在鱼类生产中的应用在鱼类生产过程中,将蒲公英添加到饲料中,对其生长性能的改善、抗病力的提高均具有积极作用。Sun等[64]研究结果表明,将柴胡提取物、蒲公英提取物和银杏提取物按照1 ∶ 4 ∶ 1(质量比)进行混合后,按照0.75 g/kg添加到啤酒糟中喂食杂交石斑鱼,可以提高石斑鱼的增重率和生长率,降低血清天冬氨酸转氨酶和乳酸脱氢酶活性,同时还可以保护肝脏和肠道形态结构的完整性,降低肝脏脂肪的积累,增强免疫力。在饲料中添加枯草芽孢杆菌和蒲公英提取物可以显著改善鲤鱼的生长性能,提高血清生化指标,促进肝脏抗氧化能力,增强免疫力[65]。饲料中添加2%的DAE能提高消化酶活性,促进鲫鱼生长,改善鲫鱼肉质[66]。饲料中添加2.83~3.13 g/kg的DFE能够提高虹鳟鱼的生长性能,添加2.59~2.75 g/kg的DFE能增强其抗氧化能力[67]。虹鳟鱼饲料中添加2.49~2.74 g/kg的DFE可以增强免疫系统功能,还可以提高对海豚链球菌的抗性[68]。在金鲳鱼的饲料中添加一定量的蒲公英提取物(1.0 g/kg)可以提高其生长性能,提高肝脏和血浆抗氧化酶活性,增强对哈维弧菌感染的抵抗力,提高免疫力、抗氧化能力以及饲料的利用率等。饲粮中添加蒲公英提取物可以改善金鲳幼鱼的肠道形态,增强抗氧化能力和肠道屏障功能,从而促进肠道健康[69];并可以调控抗氧化基因的表达,改善金鲳幼鱼的肠道内抗氧化酶活性,增加肠道绒毛的长度、宽度、数量和肌肉厚度,增加杯状细胞数量和调节免疫相关基因表达,且可通过上调闭锁小带蛋白-1(zonula occludens-1,ZO-1)和闭合蛋白(occludin)表达水平和下调封闭蛋白(claudin)-12和claudin-15表达水平,改善金鲳幼鱼的肠道形态、免疫功能、抗氧化能力和肠道屏障功能[70]。
3 小结蒲公英作为药用植物有着悠久的应用历史,蒲公英资源丰富,富含多种药理活性成分,其提取物具有抗氧化、抗肿瘤、抗炎杀菌、降低血糖、保护肝脏、增强免疫力等多种生物学功能,并且安全有效,毒副作用小,在治疗相关疾病方面极具优势。在畜、禽、鱼类等动物饲料全面禁抗的背景下,蒲公英由于其原料廉价易得、抗菌消炎效果优良等优势,成为一种较好的替抗饲料添加剂。蒲公英提取物应用于畜、禽和鱼类生产中具有提高动物免疫力、促进生长性能、抗病抗菌等,应用前景广阔。虽然蒲公英的根、叶和花都已经用于膳食补充剂和药物制剂,但具体的有效成分和治病机理还有待进一步研究。蒲公英提取物作为饲料添加剂,针对不同的添加目的和动物生长阶段,其添加量和添加形式还有待于进一步确定。
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