徐　瑩，龍　淼,2*

(1.沈陽(yáng)工學(xué)院生命工程學(xué)院，遼寧撫順 113122；2.沈陽(yáng)農(nóng)業(yè)大學(xué)畜牧獸醫(yī)學(xué)院，遼寧沈陽(yáng) 110866)

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嘔吐毒素對(duì)動(dòng)物腸道結(jié)構(gòu)與功能影響研究進(jìn)展

徐瑩1，龍淼1,2*

(1.沈陽(yáng)工學(xué)院生命工程學(xué)院，遼寧撫順 113122；2.沈陽(yáng)農(nóng)業(yè)大學(xué)畜牧獸醫(yī)學(xué)院，遼寧沈陽(yáng) 110866)

脫氧雪腐鐮刀菌烯醇(DON)對(duì)食物和飼料的污染已成為全球化的問(wèn)題，人們也逐漸開(kāi)始重視飼料污染物DON對(duì)動(dòng)物機(jī)體造成的影響。隨著科學(xué)的不斷發(fā)展，DON神秘的面紗已被逐漸揭開(kāi)。近年來(lái)的研究證實(shí)了隨飼料經(jīng)消化道進(jìn)入動(dòng)物機(jī)體的DON，在通過(guò)改變腸道組織形態(tài)學(xué)，影響腸道功能的同時(shí)，也可通過(guò)改變腸道營(yíng)養(yǎng)物質(zhì)運(yùn)輸載體、緊密鏈接蛋白、mRNA轉(zhuǎn)錄以及某些炎性物質(zhì)等的表達(dá)，影響腸道營(yíng)養(yǎng)吸收作用、屏障功能、免疫功能以及滲透作用等，最終導(dǎo)致動(dòng)物機(jī)體產(chǎn)生營(yíng)養(yǎng)不良、腹瀉、嘔吐和腸道炎癥等不良反應(yīng)。研究還表明，不同情況下DON對(duì)不同的動(dòng)物腸道可以產(chǎn)生不同的影響。

嘔吐毒素；脫氧雪腐鐮刀菌烯醇；腸道屏障；免疫；家禽；豬

脫氧雪腐鐮刀菌烯醇 (deoxynivalenol,DON)又名嘔吐毒素(vomitoxin,VT)，是玉米、小麥、大麥等谷物及其產(chǎn)品中最為常見(jiàn)的一種真菌毒素，主要是由禾谷鐮刀菌和黃色鐮刀桿菌等真菌產(chǎn)生的一類(lèi)化學(xué)結(jié)構(gòu)和生物活性相似的次級(jí)代謝產(chǎn)物，歸類(lèi)于單端孢霉烯族毒素。嘔吐毒素會(huì)導(dǎo)致家畜和家禽生產(chǎn)性能降低，采食量下降，體增重、飼料轉(zhuǎn)化率差，采食減少，急性中毒可導(dǎo)致人和動(dòng)物嘔吐、腹瀉和神經(jīng)等癥狀[1]。由于DON主要是通過(guò)腸道吸收進(jìn)入動(dòng)物體內(nèi)，研究DON毒素對(duì)動(dòng)物正常胃腸道的影響具有重要意義。

1　DON對(duì)腸道形態(tài)結(jié)構(gòu)的影響

在動(dòng)物體內(nèi)，腸道上皮細(xì)胞能夠吸收和轉(zhuǎn)運(yùn)營(yíng)養(yǎng)物質(zhì)，進(jìn)而供動(dòng)物生長(zhǎng)發(fā)育需要。腸道穩(wěn)態(tài)的維持與腸道黏膜中一些細(xì)胞(上皮細(xì)胞和淋巴細(xì)胞)的結(jié)構(gòu)功能有關(guān)，如淋巴細(xì)胞可通過(guò)分泌免疫球蛋白中和毒素，有效地將毒素限制在腸道之內(nèi)[2]。但DON 能夠損傷腸道黏膜免疫，增加機(jī)體的易感性。一旦腸道結(jié)構(gòu)被DON破壞后，將直接影響腸道上皮細(xì)胞對(duì)營(yíng)養(yǎng)物質(zhì)的吸收。

DON 可造成動(dòng)物腸道形態(tài)結(jié)構(gòu)改變。試驗(yàn)證明，圈養(yǎng)雞連續(xù)攝入日糧中含DON(5 mg/kg或10 mg/kg)的日糧5周，可造成小腸(特別是十二指腸和空腸)絨毛變得更短更細(xì)[3-4]。同樣，飼喂雞含有DON 濃度為1 mg/kg或5 mg/kg日糧，小腸絨毛高度，表面積和肌層形態(tài)發(fā)生了明顯改變。飼喂肉雞DON濃度為7.54 mg/kg日糧3周后，小腸絨毛高度變短，皺襞深度變小[5]。可見(jiàn)，DON會(huì)影響雞的腸道形態(tài)，而且對(duì)腸道形態(tài)影響程度隨著DON濃度不同而程度不同。另外，大量試驗(yàn)也表明，DON可損傷豬腸道上皮細(xì)胞，而且具有劑量依賴(lài)性。Bracarense等試驗(yàn)證明，當(dāng)飼喂豬DON(3 mg/kg飼料)的污染料5周后，便可觀察到其空腸絨毛萎縮、溶解，絨毛高度降低和腸道細(xì)胞增殖受到影響等現(xiàn)象[6]。Pinton P等[7]研究表明，飼喂豬含有2.29 mg/kg DON 日糧4周后，空腸絨毛也縮短。表明DON可對(duì)雞腸道和豬腸道絨毛上皮細(xì)胞結(jié)構(gòu)形態(tài)造成損傷，并成劑量依賴(lài)性。但DON如何影響形態(tài)結(jié)構(gòu)還需要進(jìn)一步研究。

2　DON對(duì)腸道吸收營(yíng)養(yǎng)物質(zhì)的影響

試驗(yàn)證明，DON能夠損傷腸道運(yùn)輸并影響營(yíng)養(yǎng)物質(zhì)的吸收[8]。研究表明，DON是一種特殊的鈉離子-葡萄糖共轉(zhuǎn)運(yùn)載體(SGLT-1)抑制劑，可減少機(jī)體對(duì)葡萄糖的吸收[9]。Halawa A等[10]證實(shí)DON是葡萄糖抑制劑以及DON濃度為4 μg/mL或8 μg/mL時(shí)，能對(duì)豬空腸黏膜吸收丙氨酸的過(guò)程產(chǎn)生抑制作用 。Maresca等通過(guò)研究人類(lèi)腸道細(xì)胞系HT-29-D8對(duì)DON的反應(yīng)應(yīng)答方式，證明腸道細(xì)胞一旦暴露在DON的條件下，機(jī)體對(duì)葡萄糖和右旋絲氨酸的吸收將降低；這一結(jié)論也在雞體DON代謝試驗(yàn)中得到證實(shí)[4]。

對(duì)于DON改變腸道運(yùn)輸和吸收營(yíng)養(yǎng)物質(zhì)能力的機(jī)理，目前尚不明確。但一些試驗(yàn)表明， DON可造成多種運(yùn)輸?shù)鞍紫到y(tǒng)的改變，包括改變雞體內(nèi)空腸組織中SGLT-1和十二指腸組織中GLUT-2，下調(diào)多種運(yùn)輸?shù)鞍椎幕虮磉_(dá)水平，如下調(diào)分別負(fù)責(zé)糖(葡萄糖和果糖)、某些氨基酸(左旋絲氨酸和其他的中性的氨基酸)、軟脂酸(粗飼料中的長(zhǎng)鏈脂肪酸)和一元羥酸(短鏈脂肪酸)吸收運(yùn)輸?shù)腟LC2A5、SLC7A10、SLC27A4和SLC16A1基因的表達(dá)。

3　DON對(duì)腸道黏膜滲透性的影響

DON能夠?qū)δc道黏膜造成損傷，進(jìn)而導(dǎo)致腸道黏膜滲透性的改變。DON主要通過(guò)改變雞和豬腸道的TEER (跨膜上皮電阻)或Gt(經(jīng)上皮的離子導(dǎo)電性)，損傷雞和豬的腸道屏障功能、降低腸道細(xì)胞間常見(jiàn)離子傳遞和腸道細(xì)胞完整性[11-13]。體外研究證實(shí)，DON可增加豬腸道組織FITC-dextran的旁通路，破壞腸道屏障功能，從而嚴(yán)重威脅宿主健康[13-14]。而且DON使豬腸細(xì)胞系(即IPEC-J2)產(chǎn)生了高滲透性，使共生細(xì)菌和典型的革蘭陰性鼠傷寒沙門(mén)菌跨上皮通路的能力增強(qiáng)[15-16]，可能引發(fā)腸道免疫識(shí)別障礙，使腸道發(fā)生炎癥的可能性增加[17]。Pinton P等[14]試驗(yàn)表明，飼喂豬含有2.85 mg/kg DON飼料5周，DON降低了豬空腸上皮細(xì)胞緊密連接蛋白claudin-4的表達(dá)量。

4　DON對(duì)緊密連接蛋白表達(dá)和mRNA轉(zhuǎn)錄的影響

DON對(duì)腸道內(nèi)某些緊密連接蛋白的表達(dá)和相關(guān)的mRNA造成一定的影響。雖然對(duì)豬的體內(nèi)外試驗(yàn)結(jié)果表明,DON可改變緊密連接蛋白的表達(dá)已被證實(shí)[18-19]；但是，Diesing A K[18]團(tuán)隊(duì)在關(guān)于IPEC-J2細(xì)胞系的試驗(yàn)中，并未觀察到緊密連接蛋白claudin-3的顯著減少。近幾年研究表明， DON可導(dǎo)致豬腸道黏附連接蛋白E-cadherin和occludin的表達(dá)下降[6]，緊密連接蛋白-4生物蛋白合成的減少[10]。

但是，在雞腸道中DON對(duì)蛋白表達(dá)影響的研究結(jié)果卻與豬恰恰相反。試驗(yàn)表明， DON可選擇性地提高雞空腸組織中與claudin-5相關(guān)聯(lián)的mRNA表達(dá)，但claudin-1以及閉合小環(huán)蛋白1和2 mRNA表達(dá)并沒(méi)有任何顯著的變化[5]。報(bào)道還指出claudin-5 mRNA表達(dá)增加與TLR4 mRNA表達(dá)增加的同時(shí)出現(xiàn)，意味著GNB有可能會(huì)引起腸上皮細(xì)胞的炎性反應(yīng)。此外，DON還會(huì)改變腸壁的形態(tài)，使氧化應(yīng)激蛋白mRNA的表達(dá)增加。

諸多信息表明，緊密連接蛋白可通過(guò)激活MAPKs信號(hào)通路級(jí)聯(lián)中的傳導(dǎo)物質(zhì)來(lái)調(diào)節(jié)[19-21]。Pinton P[7]研究發(fā)現(xiàn)，IPEC-1在10 μmol/L 15- ADON(即DON的乙?；男问?條件下暴露48 h會(huì)導(dǎo)致claudin-3和claudin-4的表達(dá)減少，并揭示了15- ADON的毒性和它對(duì)MAPKs的活化、claudin表達(dá)的抑制之間呈正相關(guān)，從而導(dǎo)致腸屏障功能的高度損傷。此外，當(dāng)豬靜脈滴注DON 100 μg/kg體重后，其空腸中段閉合小環(huán)蛋白-1的表達(dá)峰值下調(diào)[13]。以上這些發(fā)現(xiàn)表明，DON暴露在腸道后，腸屏障功能受損是由MAPKs磷酸化表達(dá)的增加為起始，進(jìn)而導(dǎo)致緊密連接蛋白表達(dá)減少的協(xié)調(diào)過(guò)程。

5　DON對(duì)腸道先天免疫反應(yīng)的影響

DON誘導(dǎo)腸腔局部免疫細(xì)胞產(chǎn)生免疫刺激或免疫抑制的現(xiàn)象與其劑量、頻率和暴露時(shí)間有關(guān)[22-24]。DON免疫抑制效果可以通過(guò)其綁定的核糖體并抑制蛋白質(zhì)生物合成的能力來(lái)解釋[25]。試驗(yàn)表明，DON可提升人腸道上皮細(xì)胞中IL-8的水平[25]，可使Caco-2細(xì)胞中IL-8產(chǎn)生劑量依賴(lài)性增加現(xiàn)象[15,19,26]，可誘導(dǎo)IL-8、IL-1-α、 IL-1β和TNF-α表達(dá)顯著增加為主的促炎性反應(yīng)[27]。事實(shí)上，在對(duì)活體動(dòng)物長(zhǎng)時(shí)間DON持續(xù)暴露試驗(yàn)的觀察中發(fā)現(xiàn)，DON能在動(dòng)物體內(nèi)產(chǎn)生不同程度的毒性。例如，DON不能在短時(shí)間低劑量情況下使豬回腸中TGF-β、IFN-γ、IL-4和IL-6 mRNA的表達(dá)發(fā)生變化[28]，卻可以在長(zhǎng)時(shí)間低劑量情況下使豬回腸中IL-1β和IL-8表達(dá)下調(diào)[29]。這表明，DON暴露持續(xù)的時(shí)間對(duì)基因調(diào)控不僅有顯著的影響，而且還能刺激腸道產(chǎn)生局部免疫反應(yīng)。而且，2011年Becker團(tuán)隊(duì)關(guān)于DON短時(shí)間高劑量條件下的研究發(fā)現(xiàn)，豬回腸和空腸中TNF-α、IL-1β、IFN-γ、IL-6和IL-10水平顯著上調(diào)[6,29]。同樣，研究表明DON可引起肉雞腸道中IL-1β、TGF1和IFN-γ mRNA表達(dá)降低與TNF-α和IL-8血漿水平的降低，能對(duì)雞造成局部或是全身免疫抑制現(xiàn)象[30-31]。除此以外，經(jīng)消化道攝入的DON可降低禽類(lèi)體液免疫能力，抑制機(jī)體對(duì)IBV(雞傳染性支氣管炎病毒)疫苗的應(yīng)答，產(chǎn)生慢性中毒DON的免疫抑制[32]。

在禽類(lèi)一個(gè)新的研究報(bào)告指出，飼喂鳥(niǎo)類(lèi)DON 7.54 mg/kg飼料污染的飼料3周后，其十二指腸和空腸TLR-4 mRNA的表達(dá)顯著上調(diào)，從而表明革蘭陰性菌(GNB)參與了炎癥通路的活化[5]。雖然DON暴露與TLR-4炎癥通路的關(guān)系還不明確，但現(xiàn)有數(shù)據(jù)已經(jīng)確定，在腸道中DON和GNB之間存有一種相互作用[5]。此外，這些數(shù)據(jù)還可表明脂多糖(LPS)可以以類(lèi)似于DON的方式破壞胃腸屏障的功能[7]。

DON暴露在腸腔內(nèi)，增加了GNB感染腸上皮和遷移引起腸細(xì)胞發(fā)生炎癥的可能性。比如在豬腸道回腸中，DON不僅可以增加IL-12和TNF-α表達(dá)，還可以增加IL-1β、IL-8、MCP-1和IL-6表達(dá)[16]。Cano團(tuán)隊(duì)和Vandenbroucke團(tuán)隊(duì)研究都證明了MAPKs途徑能產(chǎn)生促炎性細(xì)胞因子，因此MAPKs信號(hào)轉(zhuǎn)導(dǎo)級(jí)聯(lián)可能是DON和GNB相互作用的一種促炎性途徑[16,27]。為了更好地表示DON和GNB在腸道內(nèi)促炎性途徑的特點(diǎn)，可進(jìn)一步針對(duì)GNB的LPS和純DON等免疫刺激劑進(jìn)行研究。

6　DON對(duì)腸細(xì)胞氧化應(yīng)激參數(shù)的影響

DON暴露能提高腸細(xì)胞自由基和活性氧的水平，并產(chǎn)生氧化應(yīng)激作用。此作用造成抗氧化劑和助氧化劑之間的不平衡可認(rèn)為是DON毒性的標(biāo)志。研究表明，肉雞攝取DON(7.54 mg/kg飼料)3周后，在肝臟和空腸氧化應(yīng)激標(biāo)記物(血紅素和黃嘌呤氧化還原酶)mRNA的表達(dá)發(fā)生改變[5]。此外，丙二醛(MDA)——一種人體腸道Caco-2細(xì)胞的脂質(zhì)過(guò)氧化生物標(biāo)志物，受DON影響后，自身分泌增加[33]。

有學(xué)者報(bào)道，肉雞攝取含DON 10 mg/kg 飼料，損傷了脾白細(xì)胞和血液淋巴細(xì)胞DNA，這表明了DON具有遺傳毒性[33-34]。氧化應(yīng)激途徑是DON誘導(dǎo)DNA損傷中必然存在的一種機(jī)制[35]。例如，DON能使HepG2細(xì)胞的細(xì)胞脂質(zhì)硫代巴比妥酸反應(yīng)物(TBARS)水平增高，造成DNA損傷[36]。而且，近來(lái)一項(xiàng)研究表明，飼喂給肉仔雞含DON 10 mg/kg飼料5周后，其空腸組織將產(chǎn)生以高劑量TBARS為標(biāo)志的氧化應(yīng)激反應(yīng)[37]。因此，氧化應(yīng)激是DON毒性的一個(gè)重要機(jī)制，可導(dǎo)致腸細(xì)胞DNA和細(xì)胞膜的改變，誘導(dǎo)腸細(xì)胞的細(xì)胞凋亡、萎縮和死亡。

7　小結(jié)

胃腸道是動(dòng)物機(jī)體吸收營(yíng)養(yǎng)物質(zhì)的主要場(chǎng)所。多數(shù)情況下，DON是經(jīng)過(guò)腸黏膜吸收。不同暴露時(shí)間、頻率和劑量的DON對(duì)動(dòng)物腸道細(xì)胞產(chǎn)生的毒性不盡相同。DON可使腸道絨毛和肌層組織發(fā)生改變，黏膜滲透性受到影響，導(dǎo)致?tīng)I(yíng)養(yǎng)吸收受阻；屏障功能紊亂，導(dǎo)致致病菌的遷移率和腸道傳染病的易感性增加等。但是，DON改變腸道運(yùn)輸和吸收營(yíng)養(yǎng)物質(zhì)能力的機(jī)理與DON和TLR-4炎癥通路之間的關(guān)系尚不明確。因此，仍需大量試驗(yàn)來(lái)驗(yàn)證DON在體內(nèi)的作用機(jī)制，以求從根本上了解DON對(duì)腸道產(chǎn)生的毒性作用，為將來(lái)降低DON毒性的研究提供依據(jù)。

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Progress on Effect of Vomitoxin on Intestinal Structure and Fuction of Animals

XU Ying1,LONG Miao1,2

(1.CollegeofLifeEngineeringInstitute,ShenyangCollegeofTechnology,Fushun,Liaoning,113122,China;2.CollegeofAnimalHusbandryandVeterinaryMedicine,ShenyangAgriculturalUinversity,Shenyang,Liaoning,110866,China)

Deoxynivalenol (DON),as a food and feed contamint,has became a global problem,and people are beginning to pay attention to the impact of DON.With the continuous development of science,DON is being discovered.In recent years,some studies have confirmed that DON is absorbed into the animal body by the way of feed intake,has an influence on intestinal morphology,barrier,immunity,penetration and etc.As consequences,DON has harmful effects on animal health,such as malnutrition,diarrhea,emesis and intestinal inflammation.What's more,DON could influence different animals by different ways.

vomitoxin；deoxynivalenol;intestinal barrier;immunity;poultry;pig

2016-04-11

國(guó)家自然科學(xué)基金項(xiàng)目(31201961，31302152，31640084)；中國(guó)博士后科學(xué)基金項(xiàng)目(2014M551125)；遼寧省教育廳科學(xué)研究一般項(xiàng)目(L2014561)

徐瑩(1993-)，女，四川樂(lè)山人，主要從事中毒病研究。*通訊作者

S859.82

A

1007-5038(2016)10-0103-05