德?tīng)柋吧抽T菌的研究進(jìn)展

鄭慧娟，潘志明，焦新安

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德?tīng)柋吧抽T菌的研究進(jìn)展

鄭慧娟，潘志明，焦新安

德?tīng)柋吧抽T菌是重要的人獸共患病原菌，可引起人食物中毒和敗血癥等癥狀，而且對(duì)牲畜的繁殖及健康帶來(lái)嚴(yán)重威脅。本文就德?tīng)柋吧抽T菌的流行現(xiàn)狀、基因組學(xué)及致病機(jī)制等方面的研究進(jìn)行總結(jié)，以期為該菌的防治提供指導(dǎo)。

德?tīng)柋吧抽T菌；流行現(xiàn)狀；基因組學(xué)；致病機(jī)制

Funded by the National Key Research and Development Program Special Project (No.2016YFD0501607), Agroscientific Research in the Public Interest (No.201403054), the National Natural Science Foundation of China (Nos. 31320103907 and 31230070), the Research and Development Program of Jiangsu (BE2015343), the "six Talent Peaks Program" of Jiangsu Province (No.NY-028), the Yangzhou University Science and Technology Innovation Team, the Priority Academic Program Development of Jiangsu Higher Education Institutions (PAPD)

沙門菌是引起世界范圍內(nèi)人類食源性疾病的首要因素[1]。絕大多數(shù)沙門菌對(duì)人和動(dòng)物有致病性，具有重要的公共衛(wèi)生意義[2]。近年來(lái)在豬肉生產(chǎn)鏈中德?tīng)柋吧抽T菌的分離率逐漸趨于首位，其在健康人群的攜帶率也位于前列。本文綜述了德?tīng)柋吧抽T菌在流行現(xiàn)狀、基因組學(xué)和致病機(jī)制等方面的研究進(jìn)展，以期為德?tīng)柋吧抽T菌的防治提供理論依據(jù)。

1 德?tīng)柋吧抽T菌流行現(xiàn)狀

1.1 流行病學(xué)調(diào)查 生豬屠宰場(chǎng)及豬肉的流行病學(xué)研究發(fā)現(xiàn)，德?tīng)柋吧抽T菌已成為優(yōu)勢(shì)血清型之一[3-5]。德?tīng)柋吧抽T菌是1923年P(guān)eckham在引起食物中毒的豬肉餅中首次分離到的[6]。它在1923-1944年間引起英格蘭和威爾士11起暴發(fā)性的食物中毒，1946年澳大利亞的一家醫(yī)院也暴發(fā)了由德?tīng)柋吧抽T菌引起的嬰兒腹瀉，其中68個(gè)嬰兒發(fā)病，10例死亡[7]。美國(guó)每年約有上百萬(wàn)人因食用沙門菌污染的食品而引起腸熱癥、胃腸炎和敗血癥等疾病[8]。德?tīng)柋吧抽T菌主要在豬肉中分離到[9]，沙門菌污染的豬肉是導(dǎo)致人感染沙門菌的第二大來(lái)源，歐洲26個(gè)國(guó)家豬源沙門菌的主要血清型分別是鼠傷寒沙門菌、德?tīng)柋吧抽T菌和腸炎沙門菌，而且德?tīng)柋吧抽T菌分離率有超越鼠傷寒沙門菌的趨勢(shì)[10]。在2006-2007年間，歐盟對(duì)其25個(gè)成員國(guó)及挪威生豬屠宰場(chǎng)淋巴結(jié)樣品及13個(gè)成員國(guó)生豬屠宰場(chǎng)豬胴體擦拭樣品沙門菌的污染狀況進(jìn)行流行病學(xué)分析，發(fā)現(xiàn)沙門菌污染率分別為10.3%和8.3%；淋巴結(jié)樣品沙門菌分離株血清型中，德?tīng)柋吧抽T菌分離率為24.3%，位居第二；而胴體來(lái)源沙門菌2個(gè)主要血清型分別為鼠傷寒沙門菌占49.4%，德?tīng)柋吧抽T菌占24.3%[11]。近年來(lái)，德?tīng)柋吧抽T菌引起的沙門菌病逐漸上升[12]，這不僅對(duì)人的健康造成了嚴(yán)重的威脅，同時(shí)影響了經(jīng)濟(jì)的發(fā)展，所以應(yīng)該引起人們對(duì)德?tīng)柋吧抽T菌的重視。

在我國(guó)德?tīng)柋吧抽T菌也逐漸成為最優(yōu)勢(shì)血清型。李昱辰等[13]和蔡銀強(qiáng)等[14]對(duì)江蘇省屠宰場(chǎng)及農(nóng)貿(mào)市場(chǎng)豬肉樣品沙門菌的污染情況進(jìn)行調(diào)查發(fā)現(xiàn)德?tīng)柋吧抽T菌的分離率均占首位。1990年山東一家醫(yī)院內(nèi)暴發(fā)了由德?tīng)柋吧抽T菌引起的嬰幼兒腹瀉，其中有3名患者并發(fā)敗血癥[15]。2006年通過(guò)對(duì)武漢同濟(jì)醫(yī)院0～3歲孩子的3 746份門診腹瀉樣本沙門菌的分離鑒定發(fā)現(xiàn)，德?tīng)柋吧抽T菌的分離率占首位[16]。從這些數(shù)據(jù)可以看出德?tīng)柋吧抽T菌在國(guó)內(nèi)食品中流行比較嚴(yán)重，尤其是豬肉樣品，而中國(guó)作為豬肉生產(chǎn)和消費(fèi)大國(guó)，應(yīng)加強(qiáng)對(duì)德?tīng)柋吧抽T菌的監(jiān)測(cè)。目前，發(fā)達(dá)國(guó)家正逐步建立食源性致病菌監(jiān)測(cè)網(wǎng)和同源性分析網(wǎng)等來(lái)應(yīng)對(duì)食源性疾病的暴發(fā)和新病原菌的出現(xiàn)，世界衛(wèi)生組織(World Health Organization, WHO)于2000年建立了全球沙門菌監(jiān)測(cè)網(wǎng)(World Health Organization Global Salmonella Survey, WHO GSS)[17]。

1.2 分子流行病學(xué)研究 多位點(diǎn)序列分型(Multilocus sequence typing, MLST)通過(guò)對(duì)細(xì)菌多個(gè)管家基因核酸序列的測(cè)定從而對(duì)細(xì)菌進(jìn)行分型，此分型方法操作簡(jiǎn)單，重復(fù)性好[18]，而且Sukhnanand等[19]研究發(fā)現(xiàn)沙門菌ST型與其血清型之間有很好的對(duì)應(yīng)關(guān)系，可以對(duì)沙門菌血清型進(jìn)行預(yù)測(cè)。通過(guò)查閱沙門菌MLST數(shù)據(jù)庫(kù) (http://mlst.warwick.ac.uk/mlst/dbs/Senterica/Get

TableInfo_html)發(fā)現(xiàn)德?tīng)柋吧抽T菌一共有13種ST型，分別為ST15、ST39、ST40、ST71、ST72、ST678、ST682、ST683、ST695、ST774、ST813、ST1326、ST1585，而ST40沙門菌大多分離于豬及豬肉中，與此同時(shí)引起了大量的人沙門菌病。而目前我國(guó)主要有ST40和ST71兩種ST型，其中以ST40為主，而ST71在近幾年才開始出現(xiàn)[13-14]。在國(guó)外德?tīng)柋吧抽T菌ST型種類比較豐富，比如丹麥德?tīng)柋吧抽T菌ST型有6種，分別為ST39、ST40、ST71、ST678、ST682、ST683[20]，在德國(guó)德?tīng)柋吧抽T菌ST型有5種，ST39、ST40、ST71、ST682、ST774，其中ST39是分離率最高的一種ST型，而ST774是新發(fā)現(xiàn)的一種ST型[21]。從以上數(shù)據(jù)可以看出德?tīng)柋吧抽T菌ST型比較豐富，但是由于管家基因相對(duì)比較保守，其在區(qū)分高度關(guān)聯(lián)的菌株時(shí)仍存在一定的局限性，無(wú)法區(qū)分各ST型之間的親緣關(guān)系，因此需要分辨力更高的分型方法。

2 德?tīng)柋吧抽T菌基因組學(xué)研究

根據(jù)GenBank公布的序列，目前已有12株德?tīng)柋吧抽T菌完成了基因組測(cè)序，這推動(dòng)了德?tīng)柋吧抽T菌基因組的研究，加深了對(duì)其致病機(jī)理的認(rèn)識(shí)，為德?tīng)柋吧抽T菌的防控研究奠定了基礎(chǔ)。德?tīng)柋吧抽T菌主要在豬和豬肉上分離到，這是否說(shuō)明德?tīng)柋吧抽T菌對(duì)豬有宿主適應(yīng)性還有待研究。Hayward等[22]對(duì)兩株豬源德?tīng)柋吧抽T菌D1、D2進(jìn)行全基因組測(cè)序,通過(guò)序列比對(duì)發(fā)現(xiàn)德?tīng)柋吧抽T菌基因組中存在一個(gè)新的毒力島(SPI-23)，長(zhǎng)37 Kb，42個(gè)開放閱讀框，除此之外德?tīng)柋吧抽T菌的SPI-2, SPI-4與豬霍亂沙門菌SC-B67在基因組序列上完全一致，而SPI-18與傷寒沙門菌CT-18在基因組序列上完全一致。但是德?tīng)柋吧抽T菌基因組中缺失10個(gè)毒力島(SPI-7, SPI-8, SPI-10, SPI-15, SPI-16, SPI-17, SPI-19, SPI-20, SPI-21, SPI-22)。

細(xì)菌質(zhì)粒是獨(dú)立存在于細(xì)菌染色體DNA之外的遺傳物質(zhì)，其具有可獨(dú)立復(fù)制和轉(zhuǎn)錄的能力，而且質(zhì)粒攜帶的基因與宿主菌的表型密切相關(guān)[23]。耐藥質(zhì)粒在選擇性壓力下可以發(fā)生水平轉(zhuǎn)移[24]。在德?tīng)柋吧抽T菌基因組上存在IncL和ColE1兩個(gè)耐β-內(nèi)酰胺類抗生素質(zhì)粒[25-26]，Anne Bleicher等[27]研究發(fā)現(xiàn)了德?tīng)柋吧抽T菌4個(gè)新質(zhì)粒(pSD4.0, pSD4.6, pSD5.6, pSD107)，而且其基因組上存在的大質(zhì)粒與小質(zhì)粒間可以發(fā)生穩(wěn)定的轉(zhuǎn)移。但是目前有關(guān)德?tīng)柋吧抽T菌質(zhì)粒的數(shù)據(jù)相對(duì)較少，還有許多未知問(wèn)題有待研究。

3 德?tīng)柋吧抽T菌致病機(jī)制研究

沙門菌的致病性與入侵非吞噬細(xì)胞的能力、在胞內(nèi)存活的能力、以及在吞噬細(xì)胞內(nèi)復(fù)制和增殖的能力有關(guān)[28]。通過(guò)德?tīng)柋吧抽T菌感染小鼠模型發(fā)現(xiàn)其是低毒力菌株，口服低于2×108CFU(Colony Forming Units)對(duì)小鼠是不致死的，腹腔注射量為5×106CFU(Colony Forming Units)時(shí)死亡率為40%。德?tīng)柋吧抽T菌不僅可在小鼠糞便和尿液中存在5個(gè)月之久，而且其在多個(gè)臟器內(nèi)都可分離到，如心臟、脾臟，小腸、膀胱等[7]。德?tīng)柋吧抽T菌也可引起豬的長(zhǎng)程感染，并且在多個(gè)臟器內(nèi)長(zhǎng)期定植，如扁桃體、回盲淋巴結(jié)、脾臟、空腸、結(jié)腸、盲腸等，這可導(dǎo)致生豬屠宰時(shí)對(duì)豬肉的污染，所以其在豬肉中的分離率居高不下。細(xì)菌引起的免疫應(yīng)答反應(yīng)較弱，這也許是機(jī)體無(wú)法將其快速清除的一個(gè)原因[29]。德?tīng)柋吧抽T菌SPI-23上的potR基因在德?tīng)柋吧抽T菌對(duì)IPEC-J2細(xì)胞的黏附和侵襲有著重要的作用，缺失potR基因后，細(xì)菌對(duì)IPEC-J2細(xì)胞的黏附率和侵襲率顯著下降。當(dāng)細(xì)菌侵襲豬的空腸上皮細(xì)胞時(shí)SPI-23上有9個(gè)基因的表達(dá)量會(huì)上調(diào)(potR,chlR,docB,genE,shaU,dumE,sadZ,tinY,talN)，由此可看出這些基因與德?tīng)柋吧抽T菌的致病性密切相關(guān)[9]。德?tīng)柋吧抽T菌感染豬時(shí)往往不會(huì)引起明顯的臨床癥狀，豬只是長(zhǎng)程帶菌成為傳染源，危害整個(gè)群體，而且如果豬群感染其它高致病性病原菌時(shí)，其可引起繼發(fā)感染，導(dǎo)致豬的死亡率增加。由于這些低毒力菌株不會(huì)引起明顯的臨床癥狀，容易被忽視，所以對(duì)食品安全的威脅會(huì)更嚴(yán)重。

4 展 望

目前對(duì)德?tīng)柋吧抽T菌的研究數(shù)據(jù)還較少，而且細(xì)菌在哺乳動(dòng)物模型中的致病機(jī)制的相關(guān)信息有限，所以這給防治帶來(lái)一定的困難。慶幸的是，現(xiàn)在國(guó)內(nèi)外已經(jīng)對(duì)德?tīng)柋吧抽T菌重視起來(lái)，全基因組數(shù)據(jù)也逐漸增多，相信在不久的將來(lái)，有關(guān)德?tīng)柋吧抽T菌的致病機(jī)理、免疫應(yīng)答以及分子流行病學(xué)等方面的研究會(huì)取得重大突破。

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Research progress onSalmonellaDerby

ZHENG Hui-juan, PAN Zhi-ming, JIAO Xin-an

(JiangsuKeyLaboratoryofZoonosis,JiangsuCo-innovationCenterforPreventionandControlofImportantAnimalInfectiousDiseasesandZoonoses,JiangsuYangzhou,China,JointInternationalResearchLaboratoryofAgricultureandAgri-productSafetyoftheMinistryofEducation,Yangzhou,Jiangsu,China,KeyLaboratoryofPreventionandControlofBiologicalHazardFactors(AnimalOrigin)forAgrifoodSafetyandQuality,MinistryofAgricultureofChina,YangzhouUniversity,Yangzhou, 225009China)

SalmonellaDerby is recognized as a major human food-borne pathogen causing food poisoning, septicaemia and other symptoms. Meanwhile, it can represent a severe threat to livestock breeding and health. The objective of this review is to summarize novel research progress on epidemic, genomics, pathogenic mechanism ofSalmonellaDerby for provide reference to related research.

SalmonellaDerby; epidemic; genomics; pathogenic mechanism

國(guó)家重點(diǎn)研發(fā)計(jì)劃課題(No.2016YFD0501607)；公益性行業(yè)(農(nóng)業(yè))科研專項(xiàng)(No.201403054)；國(guó)家自然科學(xué)基金(No.31320103907,No.31230070)；江蘇省重點(diǎn)研發(fā)項(xiàng)目(No.BE2015343)；江蘇省第九批"六大人才高峰"高層次人才項(xiàng)目(No.NY-028)；揚(yáng)州大學(xué)科技創(chuàng)新團(tuán)隊(duì)；江蘇省高校優(yōu)勢(shì)學(xué)科建設(shè)工程項(xiàng)目聯(lián)合資助

潘志明，Email:zmpan@yzu.edu.cn

江蘇省人獸共患病學(xué)重點(diǎn)實(shí)驗(yàn)室/江蘇省動(dòng)物重要疫病與人獸共患病防控協(xié)同創(chuàng)新中心/農(nóng)業(yè)與農(nóng)產(chǎn)品安全國(guó)際合作聯(lián)合實(shí)驗(yàn)室/農(nóng)業(yè)部農(nóng)產(chǎn)品質(zhì)量安全生物性危害因子(動(dòng)物源)控制重點(diǎn)實(shí)驗(yàn)室

10.3969/j.issn.1002-2694.2017.07.013

R378.2

A

1002-2694(2017)07-0642-04

2016-04-14 編輯：張智芳