趙廣偉 馮廣鵬 李婉濤 張景鋒
摘 要:杜仲葉提取物作為中草藥飼料添加劑可以調(diào)節(jié)動(dòng)物的營(yíng)養(yǎng)代謝、免疫反應(yīng)和腸道健康,顯示出作為膳食抗生素替代品的良好潛力。研究發(fā)現(xiàn),飼糧中添加杜仲葉提取物能促進(jìn)動(dòng)物生長(zhǎng)、增強(qiáng)免疫力、防治疾病。主要綜述了近年來(lái)杜仲葉提取物促進(jìn)斷奶仔豬生長(zhǎng)和提高動(dòng)物免疫功能的作用及其作用機(jī)制的研究進(jìn)展,特別是對(duì)畜禽健康的營(yíng)養(yǎng)調(diào)控策略進(jìn)行了前景展望,以期為杜仲葉提取物在動(dòng)物營(yíng)養(yǎng)研究及科學(xué)應(yīng)用上提供理論參考。
關(guān)鍵詞:杜仲葉提取物;斷奶仔豬;生產(chǎn)性能;免疫反應(yīng)
中圖分類(lèi)號(hào):S 828.5 ? 文獻(xiàn)標(biāo)志碼:A ? 文章編號(hào):0253-2301(2021)06-0058-06
DOI: 10.13651/j.cnki.fjnykj.2021.06.010
Abstract: The extract of Eucommia folium, as a feed additive of Chinese herbal medicine, could regulate the nutrition metabolism, immune response and intestinal health of animals, showing a good potential as a substitute for dietary antibiotics. The research showed that the dietary added with the extract of Eucommia folium could promote the growth of animals, enhance the immunity, and prevent and cure the diseases. In this paper, the effects of the extract of Eucommia folium on the growth and immune function of weaned piglets and its mechanism were reviewed, especially the nutritional regulation strategies for the health of livestock and poultry were prospected, so as to provide theoretical references for the animal nutrition research and the scientific application of the extract of Eucommia folium.
Key words: Extract of Eucommia folium; Weaned piglets; Production performance; Immune response; Intestinal health
斷奶被認(rèn)為是集中飼養(yǎng)的豬的應(yīng)激期[1],豬在應(yīng)激期間會(huì)降低對(duì)營(yíng)養(yǎng)物質(zhì)的消化吸收,引起氧化應(yīng)激,導(dǎo)致斷奶前后出現(xiàn)生長(zhǎng)遲緩和腹瀉,進(jìn)而對(duì)牲畜生產(chǎn)造成巨大的經(jīng)濟(jì)損失[2]。杜仲葉(Eucomia Ulmoides,EL)作為傳統(tǒng)中草藥杜仲的副產(chǎn)物,在我國(guó)相當(dāng)普遍[3]。杜仲葉提取物含有豐富的生物活性化合物(如黃酮類(lèi)化合物、綠原酸、桃葉珊瑚素、京尼平苷酸等),具有抗炎、抗氧化、抗病毒和保肝功能[4]。研究發(fā)現(xiàn),飼糧中添加杜仲葉提取物可以提高斷奶仔豬的生長(zhǎng)性能[5]、抗氧化能力及消化吸收能力[6]。本文就杜仲葉提取物在仔豬營(yíng)養(yǎng)功能、腸道功能和免疫功能的機(jī)制進(jìn)行綜述,以期為杜仲葉提取物在豬生產(chǎn)上的進(jìn)一步應(yīng)用提供參考。
1 杜仲葉提取物的主要功能介紹
1.1 杜仲葉提取物的成分及代謝
京尼平苷酸和綠原酸(Chlorogenic acid,CGA)是杜仲葉中重要的活性成分,前者可促進(jìn)膠原合成[7],提高角質(zhì)層的轉(zhuǎn)化率[8],后者具有抗菌、消炎、致突變、抗氧化等生物活性[9]。綠原酸由反式肉桂酸和奎寧酸酯化而形成的酚類(lèi)化合物,是一類(lèi)酯的總稱(chēng),最常見(jiàn)的綠原酸是5咖啡??鼘幩幔?Ocaffeoylquinic acid,5CQA)[10]。CGA的結(jié)構(gòu)不穩(wěn)定,在碳1和碳3上有軸向羥基,在碳4和碳5上有平方羥基,其容易水解產(chǎn)生奎尼酸和各種肉桂酸,釋放的肉桂酸可以脫羧,降解成一系列簡(jiǎn)單的酚類(lèi)物質(zhì)并進(jìn)一步轉(zhuǎn)化,從而進(jìn)一步發(fā)揮作用[11]。
1.2 杜仲葉提取物的吸收途徑及代謝產(chǎn)物
前人研究結(jié)果發(fā)現(xiàn),尿液中可以檢測(cè)出少量綠原酸[12-14]。Lafay等[15]在大鼠上的研究發(fā)現(xiàn),部分綠原酸以咖啡酸的形式從腸道流出物中回收,說(shuō)明在腸道黏膜中存在微量酯酶活性,同時(shí)在血漿和膽汁中均未檢出綠原酸,以上研究結(jié)果表明,綠原酸在小腸中被吸收和水解。同時(shí)在大鼠和人上的研究發(fā)現(xiàn),咖啡酸及其O甲基化代謝物普遍參在于血液和尿液[16-18],結(jié)果表明,綠原酸可以在體內(nèi)水解。以上研究結(jié)果表明,綠原酸可能在腸道內(nèi)水解為具有生物活性的代謝產(chǎn)物,其經(jīng)消化道吸收后通過(guò)血液循環(huán)運(yùn)送到全身各處,從而發(fā)揮相應(yīng)的生理功能。
2 杜仲葉提取物對(duì)仔豬生產(chǎn)性能的影響及可能作用機(jī)制
2.1 杜仲葉提取物促進(jìn)仔豬生長(zhǎng)
研究發(fā)現(xiàn),杜仲葉提取物提高了斷奶仔豬[19]、凡納濱對(duì)蝦(Litopenaeus vannamei)[20]、肉雞[21]、異育銀鯽(Carassius gibelio)[22]、三黃雞[23]的體重,增加仔豬[24]、肉兔[25]的采食量,以及羔羊[26]的飼料效率。同時(shí),前人研究結(jié)果發(fā)現(xiàn),飼糧中添加杜仲葉中提取物對(duì)羔羊的平均日增重、干物質(zhì)采食量和飼料效率均無(wú)顯著影響[26],這可能與杜仲葉提取物的添加量、動(dòng)物種類(lèi)不同有關(guān)。
2.2 杜仲葉提取物促進(jìn)仔豬生長(zhǎng)的可能作用機(jī)制
斷奶后,飼料組成和飼養(yǎng)條件的突然變化又會(huì)引起消化功能的急劇變化,從而導(dǎo)致腸道對(duì)營(yíng)養(yǎng)物質(zhì)的吸收不良[27-28]。動(dòng)物對(duì)飼料中營(yíng)養(yǎng)物質(zhì)的利用是體現(xiàn)飼料營(yíng)養(yǎng)價(jià)值的一個(gè)重要指標(biāo)。動(dòng)物生長(zhǎng)與消化功能密切相關(guān)。小腸消化酶(脂肪酶、胰蛋白酶)活性的提高有利于斷奶仔豬的生長(zhǎng)[29]。Ding等[29]研究結(jié)果表明,飼糧中添加杜仲葉提取物提高斷奶仔豬十二指腸脂肪酶活性和空腸胰蛋白酶活性。腸道吸收能力也是體內(nèi)充分利用營(yíng)養(yǎng)物質(zhì)的基礎(chǔ)[30]。堿性磷酸酶(Alkaline phosphatase,AKP)活性提高可以反映腸道上皮細(xì)胞的吸收能力增強(qiáng)[31]。石英等[22]研究結(jié)果表明,飼糧中補(bǔ)充杜仲葉提取物可以提高異育銀鯽血清中AKP的活性。同時(shí),冷向軍等[32]研究結(jié)果表明,飼糧中補(bǔ)充杜仲葉提取物對(duì)草魚(yú)(Ctenopharyngodon idella)血清AKP的活性無(wú)顯著影響。以上研究結(jié)果表明,杜仲葉提取物可能通過(guò)增強(qiáng)仔豬腸道消化吸收能力促進(jìn)仔豬生長(zhǎng),但是可能與動(dòng)物的種類(lèi)有關(guān)。
腸道健康是保證動(dòng)物健康生長(zhǎng)的前提條件[33]。腸道黏膜的改變,如絨毛高度降低、隱窩深度增加、腸絨毛高度與隱窩深度之比(VH/CD)增加等,被認(rèn)為是由入侵的病原體引起的組織損傷[34]。之前的研究結(jié)果表明,飼糧中添加杜仲葉提取物提高了斷奶仔豬十二指腸和空腸絨毛高度[29]。同時(shí),Peng等[3]在斷奶仔豬上的研究結(jié)果表明,飼糧中補(bǔ)充杜仲葉提取物降低了隱窩深度,增加了絨毛高度與隱窩深度的比值。腸道健康與腸道抗氧化能力[35]、腸道結(jié)構(gòu)完整性[35]、腸炎抵抗能力[36]密切相關(guān)。血清中丙二醛(Malondialdehyde,MDA)水平的降低,過(guò)氧化氫酶(Catalase,CAT)、超氧化物歧化酶(Superoxide dismutase,SOD)和谷胱甘肽過(guò)氧化物酶(Glutathione peroxidase,GSH-Px)水平的增加以及腸道緊密連接相關(guān)蛋白基因表達(dá)水平的增加可以反映動(dòng)物腸道健康水平增加[37]。研究表明,飼糧中添加杜仲葉提取物可以降低仔豬血清和肝臟MDA水平[29]、增加凡納濱對(duì)蝦血清CAT、SOD和GSH-Px水平。Peng等[3]研究結(jié)果表明,飼糧中添加杜仲提取物可以增加仔豬空腸黏膜緊密連接相關(guān)蛋白claudin3的基因表達(dá)水平。以上研究結(jié)果表明,杜仲葉提取物可能通過(guò)提高仔豬消化吸收能力、改善腸道形態(tài)及降低腸道通透性維持腸道功能健康,進(jìn)而促進(jìn)仔豬的生長(zhǎng)。
揮發(fā)性脂肪酸(Volatile Fatty Acid,VFA)水平是可以反映腸道健康和微生物活性的指標(biāo)[38]。Zhao等[39]研究結(jié)果表明,杜仲葉提取物可以提高SPF小鼠血清中揮發(fā)性脂肪酸乙酸和異丁酸的濃度。Freire等[40]研究表明,含有甜菜堿的飼糧飼喂仔豬時(shí)其盲腸中揮發(fā)性脂肪酸含量較高。同時(shí),Che等[41]研究表明,飼糧中添加了黃芪莖葉纖維的仔豬具有較高的乙酸、丙酸、丁酸和總揮發(fā)性脂肪酸含量。腸道微生物通過(guò)與宿主的營(yíng)養(yǎng)利用和腸道系統(tǒng)的發(fā)育相互作用,在宿主的營(yíng)養(yǎng)和健康、采食量和效率以及增重方面發(fā)揮重要作用[42-44]。Peng等[3]研究結(jié)果表明,飼糧中添加杜仲葉提取物可以提高仔豬結(jié)腸微生物群落豐富度和多樣性,與呂武興等[23]在三黃雞上的研究結(jié)果一致。杜仲葉提取物可能通過(guò)增加腸道中揮發(fā)性脂肪酸的含量及腸道微生物多樣性來(lái)維持腸道健康。
以上研究結(jié)果表明,杜仲葉提取物可能通過(guò)增強(qiáng)營(yíng)養(yǎng)物質(zhì)的消化吸收、維持腸道功能健康來(lái)促進(jìn)動(dòng)物生長(zhǎng)。
3 杜仲葉提取物對(duì)仔豬免疫功能的影響及可能作用機(jī)制
3.1 杜仲葉提取物對(duì)仔豬免疫功能的影響
病原菌入侵可以刺激機(jī)體激活免疫系統(tǒng)清除病原菌,若不能及時(shí)清除,則可能導(dǎo)致機(jī)體被病原菌感染而發(fā)病,嚴(yán)重時(shí)可能導(dǎo)致死亡。Chen等[45]研究結(jié)果表明,魚(yú)類(lèi)的疾病抵抗能力與其攻毒后的存活率呈正相關(guān)。氣單胞菌引起魚(yú)類(lèi)運(yùn)動(dòng)性氣單胞菌敗血癥,導(dǎo)致水產(chǎn)養(yǎng)殖業(yè)的高死亡率和嚴(yán)重的經(jīng)濟(jì)損失,嗜水氣單胞菌被認(rèn)為是氣單胞菌感染的主要病原[46]。羅慶華[47]研究結(jié)果表明,飼糧中補(bǔ)充杜仲葉提取物可以降低嗜水氣單胞菌攻毒后鯉魚(yú)(Cyprinus carpio)的存活率。
腸道中變形菌門(mén)的豐富度增加表明腸道菌群失調(diào)或者不平衡[48]。Peng等[3]研究結(jié)果表明,杜仲葉提取物可以提高斷奶仔豬結(jié)腸微生物群落豐富度,降低變形菌門(mén)的豐富度。斷奶仔豬腹瀉事件是由大腸菌群過(guò)剩引起的,控制其傳播是降低腹瀉發(fā)病率的有效手段[49]。在仔豬上的研究發(fā)現(xiàn),飼糧中杜仲葉提取物的添加降低了腹瀉率[3],與Ding等[29]在斷奶仔豬上的研究結(jié)果一致。Baratta等
[50]研究表明,中草藥添加劑具有很強(qiáng)的抗菌活性,可以抑制大腸桿菌的生長(zhǎng)[51]。飼糧中添加一定量的中草藥黃芪莖葉纖維可以降低仔豬腹瀉發(fā)病率(P<0.05)[41],與Li等[49]研究結(jié)果相似。同時(shí),Chen等[19]研究結(jié)果表明,飼糧中綠原酸的添加降低了仔豬腹瀉率。
3.2 杜仲葉提取物對(duì)仔豬免疫功能影響的可能作用機(jī)制
血清抗體水平可以反映體液免疫的水平[52]。王鑫等[53]研究結(jié)果表明,飼糧中補(bǔ)充杜仲葉提取物提高仔豬血清中IgA的含量,與李曉梅等[54]在仔豬上的研究結(jié)果一致。Tan等[55]研究表明,飼糧中加入黃芪多糖可以提高母豬初乳中IgM和IgG水平,與Kajimura等[56]在小鼠上的研究一致。豬瘟(Classical swine fever,CSF)是一種傳染性病毒疾病,可以造成嚴(yán)重的經(jīng)濟(jì)損失的疾病[57]。前人研究表明,母源性CSF抗體的存在可以保護(hù)仔豬免受這種疾病的侵害[58]。Tan等[55]研究表明,分娩前在飼糧中補(bǔ)充黃芪多糖可增加初乳中母源性抗CSF特異性抗體。黃芪多糖能增加抗體產(chǎn)生可能與其能激活B細(xì)胞和巨噬細(xì)胞有關(guān)[59-60]。淋巴細(xì)胞對(duì)刀豆蛋白(Con A)的增殖反應(yīng)是反映T淋巴細(xì)胞功能和細(xì)胞免疫的重要標(biāo)準(zhǔn)。在小鼠上的研究表明,杜仲葉提取物可增強(qiáng)脾細(xì)胞對(duì)刀豆蛋白(Con)A的增殖反應(yīng)和腹腔巨噬細(xì)胞吞噬功能[61]。同時(shí),Yuan等[52]研究表明,飼糧中補(bǔ)充黃芪多糖可以改善斷奶仔豬淋巴細(xì)胞對(duì)Con A的增殖反應(yīng),與Kong等[62]在雞上的研究結(jié)果一致。
細(xì)胞因子介導(dǎo)的炎癥反應(yīng)是血腦屏障細(xì)胞免疫反應(yīng)的重要組成部分[45]。腫瘤壞死因子(Tumor necrosis factor,TNF)α是重要的促炎細(xì)胞因子,參與宿主對(duì)細(xì)菌定殖或入侵的防御機(jī)制[63-64]。在大鼠中的研究發(fā)現(xiàn),下調(diào)促炎細(xì)胞因子可減弱炎癥反應(yīng)[65]。Jiang等[35]研究結(jié)果表明,飼糧中補(bǔ)充杜仲葉提取物可以提高仔豬肝臟中TNFα的基因表達(dá)水平。
以上研究結(jié)果表明,杜仲葉提取物可能通過(guò)提高動(dòng)物的體液免疫和細(xì)胞免疫,減弱炎癥反應(yīng)來(lái)增強(qiáng)仔豬的免疫功能,但還需要更加深入系統(tǒng)的研究。
4 結(jié)論及展望
綜上所述,杜仲葉提取物能夠通過(guò)提高營(yíng)養(yǎng)物質(zhì)的消化率、維持腸道功能健康及免疫功能進(jìn)而促進(jìn)仔豬的生長(zhǎng)。杜仲葉提取物通過(guò)提高仔豬腸道脂肪酶和胰蛋白酶的活性,增加絨毛高度來(lái)促進(jìn)仔豬對(duì)營(yíng)養(yǎng)物質(zhì)的消化吸收,通過(guò)提高腸道抗氧化能力、增強(qiáng)腸道緊密連接及增加腸道微生物的多樣性及揮發(fā)性脂肪酸的含量提高腸道功能,進(jìn)而提高仔豬的生長(zhǎng)性能;通過(guò)提高仔豬體液免疫、細(xì)胞免疫和減弱炎癥反應(yīng)來(lái)提高仔豬的抗病力。
近年來(lái)關(guān)于植物性提取物替代抗生素的研究逐漸成為動(dòng)物營(yíng)養(yǎng)學(xué)中的一大熱點(diǎn)。杜仲葉提取物中含有的多種活性成分使得其在畜牧業(yè)中的應(yīng)用可能為改善畜牧業(yè)生產(chǎn)和控制疾病提供一個(gè)全新的途徑。然而,有關(guān)斷奶仔豬抗氧化能力、腸道功能和消化酶活性的影響的信息很少。本文綜述了杜仲葉提取物對(duì)仔豬生產(chǎn)性能、腸道形態(tài)、抗氧化活性、免疫相關(guān)蛋白和血液指標(biāo)的影響,清晰闡明杜仲葉提取物對(duì)仔豬機(jī)體健康影響的機(jī)制,這可能為預(yù)防和治療疾病提供新思路,具有廣闊的應(yīng)用前景。
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