張永寧,梅 琳,張 舟,唐麗杰,吳紹強(qiáng)
(1.中國(guó)檢驗(yàn)檢疫科學(xué)研究院動(dòng)物檢疫研究所,北京 100176;2.東北農(nóng)業(yè)大學(xué)動(dòng)物醫(yī)學(xué)學(xué)院,哈爾濱 150030)
豬圓環(huán)病毒3型研究進(jìn)展
張永寧1,梅 琳1,張 舟1,唐麗杰2,吳紹強(qiáng)1
(1.中國(guó)檢驗(yàn)檢疫科學(xué)研究院動(dòng)物檢疫研究所,北京 100176;2.東北農(nóng)業(yè)大學(xué)動(dòng)物醫(yī)學(xué)學(xué)院,哈爾濱 150030)
2016年,美國(guó)堪薩斯州立大學(xué)科研人員利用宏基因組測(cè)序技術(shù),從患有皮炎腎病綜合征(PDNS)和繁殖障礙母豬及其流產(chǎn)胎兒體內(nèi)首次鑒定出一種新型豬圓環(huán)病毒,命名為豬圓環(huán)病毒3型(Porcine circovirus 3,PCV3)。流行病學(xué)調(diào)查發(fā)現(xiàn)PCV3感染多見于具有PDNS和繁殖障礙癥狀豬群,目前已于美國(guó)多個(gè)州豬群內(nèi)普遍流行。同時(shí),美國(guó)舊金山血液系統(tǒng)研究所科研人員從3頭不明病因?qū)е碌男呐K和多器官炎癥仔豬體內(nèi)也檢測(cè)出PCV3。隨后,我國(guó)湖北、廣東等多個(gè)省市豬群中也檢測(cè)出PCV3。近來,波蘭和韓國(guó)相繼報(bào)道存在PCV3感染豬群。作為一種新發(fā)現(xiàn)病原體,PCV3對(duì)豬致病性及在豬圓環(huán)病毒相關(guān)疾病(PCVAD)中作用以及現(xiàn)有商品化PCV2疫苗對(duì)其免疫效力等有待進(jìn)一步研究。文章綜述PCV3最新進(jìn)展,以期為后續(xù)研究提供參考。
宏基因組測(cè)序;豬圓環(huán)病毒3型;豬皮炎腎病綜合征;繁殖障礙;豬圓環(huán)病毒相關(guān)疾病
豬圓環(huán)病毒(Porcine circovirus,PCV)為一種共價(jià)閉合、單股環(huán)狀DNA病毒,是迄今發(fā)現(xiàn)最小動(dòng)物病毒[1]。2016年前,全球范圍內(nèi)僅發(fā)現(xiàn)PCV1和PCV2兩個(gè)基因型[2-3]。其中,PCV1被認(rèn)為是豬腎細(xì)胞PK-15污染物,但不產(chǎn)生細(xì)胞病變效應(yīng)[4],對(duì)豬無(wú)致病性[5]。PCV2是引起斷奶仔豬多系統(tǒng)衰竭綜合征(PMWS)主要病原[6],與豬皮炎和腎病綜合征(PDNS)[7]、豬呼吸道疾病綜合征(PRDC)[8]、繁殖障礙[9]、增生性壞死性肺炎(PNP)[10]和腸炎[11-12]等疾病密切相關(guān),均為豬圓環(huán)病毒相關(guān)疾?。≒CVAD)[13-14]。臨床上,PCV2常與豬繁殖與呼吸綜合征病毒(PRRSV)[15]、豬細(xì)小病毒(PPV)[16]和豬肺炎支原體(M.hyo)[17]等病原體混合/繼發(fā)感染,造成養(yǎng)豬業(yè)巨大經(jīng)濟(jì)損失。
2003年全球范圍內(nèi)PCV2發(fā)生由PCV2a向PCV2b基因型轉(zhuǎn)變,病毒致病力增強(qiáng),對(duì)養(yǎng)豬業(yè)危害加重[18]。2012年P(guān)CV2在世界范圍內(nèi)再次發(fā)生基因型轉(zhuǎn)變,由PCV2b向致病力更強(qiáng)PCV2d轉(zhuǎn)變[19-20],現(xiàn)有疫苗免疫效果降低。2015年6月,美國(guó)北卡羅來納州某商品化豬場(chǎng)暴發(fā)PDNS疫情,與歷史平均值相比,該豬場(chǎng)母豬死亡率升高10.2%、受孕率下降0.6%?;疾∧肛i厭食、皮膚呈現(xiàn)多灶性丘疹、斑點(diǎn)和淺表性皮炎,所產(chǎn)胎兒(包括弱胎、死胎、木乃伊胎)具有相似臨床癥狀。Palinski等借助新一代測(cè)序(NGS)技術(shù),從患病母豬及流產(chǎn)胎兒體內(nèi)首次鑒定出一種新病毒[21],該病毒與圓環(huán)病毒科病毒具有類似基因組結(jié)構(gòu)和遺傳相似性,但與其他圓環(huán)病毒衣殼蛋白氨基酸序列同源性低于70%,根據(jù)國(guó)際病毒分類委員會(huì)(ICTV)圓環(huán)病毒分類標(biāo)準(zhǔn)[22],將其歸類為一種新型圓環(huán)病毒,命名為PCV3[21]。作為新動(dòng)物病原體,PCV3對(duì)豬致病性和在PCVAD中作用以及現(xiàn)有商品化PCV2疫苗對(duì)其免疫效果等有待進(jìn)一步研究。
文章綜述PCV3在病原學(xué)、流行病學(xué)、臨床癥狀與病理變化、診斷與防控對(duì)策等方面國(guó)內(nèi)外最新進(jìn)展,以期為后續(xù)科學(xué)研究提供參考。
1.1 分類
PCV3在分類上屬于圓環(huán)病毒科(Circoviridae)圓環(huán)病毒屬(Circovirus)成員[21]。
1.2 形態(tài)與結(jié)構(gòu)
與PCV1和PCV2一樣,PCV3病毒粒子呈二十面體結(jié)構(gòu),無(wú)囊膜,直徑為17~20 nm。基因組為單股DNA,全長(zhǎng)2 000 bp,包含3個(gè)主要開放閱讀框(ORF)。其中,ORF1編碼由297個(gè)氨基酸(aa)組成復(fù)制酶蛋白R(shí)ep,ORF2編碼由214個(gè)aa組成衣殼蛋白Cap,兩者復(fù)制方向相反(見圖1)。在rep和cap基因間隔區(qū)5'端存在莖-環(huán)結(jié)構(gòu)(見圖1),具有與PCV1和PCV2完全一致、保守九核苷酸基序(TAGTATTAC)[23-24]。
圖1 PCV3基因組結(jié)構(gòu)Fig.1 PCV3 genomic structure
按照慣例,將莖-環(huán)結(jié)構(gòu)中位于九核苷酸基序內(nèi)第8位核苷酸殘基“A”定義為基因組起始位置“1”[25]。但Rep蛋白編碼基因序列中無(wú)標(biāo)準(zhǔn)起始密碼子ATG,rep基因5'端存在密碼子GTC,該替代性起始密碼子也見于豬圓環(huán)病毒PorkNW2/USA/2009(GenBank登錄號(hào)ADU77001.1)及禽類圓環(huán)病毒(如鵝圓環(huán)病毒、鴿圓環(huán)病毒和喙羽病病毒)[26-27]。與其他圓環(huán)病毒Cap蛋白類似[28],PCV3 Cap蛋白N端富含精氨酸,為病毒核定位信號(hào)區(qū)域,與病毒在細(xì)胞核內(nèi)定位有關(guān)。ORF3編碼是一個(gè)由231個(gè)aa組成的功能未知蛋白質(zhì),該蛋白與PorkNW2/USA/2009編碼蛋白質(zhì)具有94%同源性,與鼠皰疹病毒M169具有39%同源性,ORF3編碼方向與rep基因相同[29]。與rep基因類似,ORF3起始密碼子尚不清楚,其5'端密碼子為TCG。位于ORF3第55位aa處甲硫氨酸可能為另一個(gè)翻譯起始位點(diǎn),產(chǎn)生一個(gè)由177個(gè)aa組成蛋白質(zhì)[21]。
1.3 抗原性
Cap蛋白為PCV主要結(jié)構(gòu)蛋白,誘導(dǎo)動(dòng)物機(jī)體產(chǎn)生特異性免疫應(yīng)答[30-32]。PCV2與PCV3 Cap蛋白aa同源性僅為30%,兩者間存在交叉免疫保護(hù)可能性較低[21,33]。湛洋等對(duì)PCV3 Cap蛋白三維結(jié)構(gòu)分析發(fā)現(xiàn),PCV3和PCV2 Cap蛋白具有類似由兩個(gè)β-片層結(jié)構(gòu)組成jelly roll折疊,但PCV3 Cap蛋白其中一個(gè)片層僅由3條反平行β-折疊組成,而PCV2 Cap蛋白兩個(gè)片層均由4條反平行β-折疊組成[33]。借助B細(xì)胞抗原表位在線預(yù)測(cè)軟件分析發(fā)現(xiàn),PCV3 Cap蛋白抗原表位主要分布于8個(gè)區(qū)域(見表1)。
表1 PCV3 Cap蛋白B細(xì)胞抗原表位預(yù)測(cè)Table 1 B-cell epitope prediction for the PCV3 Cap protein
1.4 培養(yǎng)特性
目前,豬腎細(xì)胞(PK-15、SK)和豬睪丸細(xì)胞(ST)常用于PCV2分離與培養(yǎng)[34],推測(cè)PCV3可在此類細(xì)胞中增殖[21]。
1.5 理化特性
PCV對(duì)不良環(huán)境抵抗力較強(qiáng),可于酸性環(huán)境中(pH=3)存活較長(zhǎng)時(shí)間,無(wú)血凝性[35]。Allan和O'Dea等對(duì)PCV熱穩(wěn)定性分析發(fā)現(xiàn),PCV1于56和70℃加熱15 min后感染性無(wú)明顯變化[35-36]。PCV2于75℃處理15 min仍有感染性,但80℃處理15 min后完全失去感染性[36],耐熱性良好。Welch等研究發(fā)現(xiàn),常規(guī)巴氏殺菌、干熱處理和濕熱處理均難以徹底滅活血液制品中PCV2[37]。由于PCV無(wú)囊膜,因此對(duì)氯仿、碘酒、酒精等有機(jī)溶劑不敏感,但對(duì)苯酚、季胺類化合物、氫氧化鈉和氧化劑等較敏感[35,38]。鑒于PCV3與PCV2屬于同一病毒科,推測(cè)兩者理化特性相似,可采用相同消毒劑和消毒方法處理PCV3污染圈舍、環(huán)境、運(yùn)輸工具等。
2.1 傳染源
處于病毒血癥期感染豬為主要傳染源,包括發(fā)病豬和隱性感染豬。病毒隨患病豬鼻、口腔分泌物及糞便排出[39-40]。
2.2 傳播途徑
PCV2可通過呼吸道、消化道和泌尿生殖道分泌物,以直接或間接接觸方式傳播[40-42]。懷孕母豬感染PCV2后,可經(jīng)胎盤垂直傳染仔豬[43-44]。Palinski等從患有PDNS癥狀母豬及其流產(chǎn)胎兒體內(nèi)均發(fā)現(xiàn)PCV3全基因組序列[21],間接證實(shí)PCV3垂直傳播方式。PCV2感染公豬精液中存在間歇性排毒現(xiàn)象[45],將摻入PCV2的健康豬精液人工授精接種健康母豬,可導(dǎo)致母豬發(fā)生繁殖障礙,產(chǎn)弱胎、死胎和木乃伊胎[46]。Ku等通過PCR技術(shù)從豬精液中檢出PCV3核酸,揭示PCV3也可以通過精液傳播[47]。
2.3 易感動(dòng)物
不同年齡、性別和品種均可感染PCV3,但以妊娠母豬和仔豬更易感[48-49]。目前,尚不清楚野豬是否對(duì)PCV3易感,也未發(fā)現(xiàn)其他物種感染PCV3報(bào)道。
2.4 發(fā)生與分布
圖2 PCV3在美國(guó)和中國(guó)的流行與分布Fig.2 Prevalenceand distribution of PCV3 in the USA and China
2015年6月,美國(guó)北卡羅來納州某豬場(chǎng)暴發(fā)PDNS疫情,與歷史平均值相比,母豬死亡率升高10.2%、受孕率下降0.6%?;疾∧肛i出現(xiàn)厭食癥狀,皮膚呈現(xiàn)多灶性丘疹、斑點(diǎn)和淺表性皮炎,所產(chǎn)胎兒(包括弱胎、死胎、木乃伊胎)具有類似臨床癥狀。2016年,Palinski等借助NGS技術(shù)從患病母豬及其流產(chǎn)胎兒體內(nèi)首次鑒定出PCV3,美國(guó)多個(gè)州豬群證實(shí)存在PCV3感染[21,48],如北卡羅來納州和愛荷華州等(見圖2A)。我國(guó)湖北、廣東某些發(fā)生繁殖障礙母豬以及急性死亡仔豬體內(nèi)也檢測(cè)出PCV3[50]。截至目前,我國(guó)已有13個(gè)省豬場(chǎng)存在PCV3感染與流行[47,51],且主要分布于中東部地區(qū)(見圖2B),其他省市是否存在PCV3感染目前尚不清楚。Stadejek等對(duì)波蘭境內(nèi)14個(gè)商品化豬場(chǎng)母豬、仔豬和育成豬血清作PCV3檢測(cè),首次證實(shí)歐洲地區(qū)存在PCV3感染[49]。Kwon等對(duì)采自韓國(guó)6個(gè)省73個(gè)豬場(chǎng)的豬唾液樣品作PCV3核酸檢測(cè),首次發(fā)現(xiàn)韓國(guó)豬群也存在PCV3感染與流行[52]。
PCV3感染可導(dǎo)致母豬出現(xiàn)厭食癥狀,皮膚出現(xiàn)多灶性丘疹、斑點(diǎn)和淺表性皮炎,生產(chǎn)性能下降;妊娠母豬發(fā)生繁殖障礙,產(chǎn)弱胎、死胎、木乃伊胎和弱仔豬,病情嚴(yán)重甚至造成急性死亡[21,47];不同胎齡胎兒均可發(fā)生流產(chǎn)。組織學(xué)檢查可見,真皮和皮下組織呈以纖維蛋白樣變、透壁性嗜中性粒細(xì)胞浸潤(rùn)、出血和纖維蛋白滲出為特征的壞死性血管炎;炎性細(xì)胞浸潤(rùn)通常蔓延至周圍真皮和皮下組織,以“袖套”形式圍繞于血管和皮膚附件周圍;偶見輕度角化過度表皮增生癥。腎臟呈現(xiàn)彌散性膜增生性腎小球腎炎,表現(xiàn)為腎小球硬化和波曼氏囊(Bowman's capsule)增厚、皮質(zhì)小管衰退和不同程度的間質(zhì)纖維化;腎小管發(fā)生擴(kuò)張,衰退上皮細(xì)胞堆積于管腔,皮質(zhì)間質(zhì)和腎小球出現(xiàn)大簇淋巴細(xì)胞和巨噬細(xì)胞擴(kuò)散性浸潤(rùn)。肺臟出現(xiàn)不同程度支氣管間質(zhì)性肺炎,偶爾繼發(fā)化膿性支氣管肺炎;中小型氣道和小血管周圍出現(xiàn)由淋巴細(xì)胞和漿細(xì)胞組成聚集物,圍繞于細(xì)支氣管和血管周圍形成“袖套”樣結(jié)構(gòu);相鄰肺泡隔膜發(fā)生淋巴細(xì)胞和漿細(xì)胞浸潤(rùn);肺泡腔內(nèi)充滿中等數(shù)量泡沫巨噬細(xì)胞、少量多核巨細(xì)胞和小簇嗜中性粒細(xì)胞,形成腔內(nèi)水腫。淋巴結(jié)呈現(xiàn)彌漫性肉芽腫淋巴結(jié)炎,表現(xiàn)為中度皮質(zhì)淋巴細(xì)胞消耗,甚至出現(xiàn)組織細(xì)胞和多核巨細(xì)胞替代[21]。
患有體重減輕、呼吸、神經(jīng)系統(tǒng)疾病以及心臟和全身多發(fā)性炎癥仔豬可能與PCV3感染有關(guān)[48]。但患病仔豬組織同時(shí)檢出豬星狀病毒4型、輪狀病毒A、豬巨細(xì)胞病毒和豬血凝性腦脊髓炎病毒,目前尚未證實(shí)上述臨床癥狀與病理變化是否與PCV3感染直接相關(guān)。
4.1 臨床診斷
除PCV2感染可以導(dǎo)致PDNS癥狀外[53],無(wú)PCV2存在情況下,將PRRSV以及含有細(xì)環(huán)病毒(Torque teno virus,TTV)組織勻漿液接種健康豬,也可產(chǎn)生PDNS癥狀[54]。鑒于PCV3是從患有PDNS癥狀母豬及其流產(chǎn)胎兒體內(nèi)檢出唯一病原體,PCV3感染豬臨床癥狀與病理變化與PCV2高度相似[21],因此,須結(jié)合臨床與實(shí)驗(yàn)室診斷確診是否為PCV3感染。
4.2 實(shí)驗(yàn)室診斷
4.2.1 病毒分離與鑒定
利用無(wú)PCV1和PCV2污染PK-15、SK或ST細(xì)胞從患病豬血清、皮膚、腎臟、肺和淋巴結(jié)等組織樣品中分離PCV3[21],借助DNA測(cè)序、中和試驗(yàn)、免疫組化和熒光定量PCR等技術(shù)鑒定病毒[21]。
4.2.2 熒光定量PCR
表2 PCV3熒光定量PCR引物和探針Table 2 Primers and probesused for real-time PCR detection of PCV3
美國(guó)堪薩斯州立大學(xué)、河北出入境檢驗(yàn)檢疫局和中國(guó)檢驗(yàn)檢疫科學(xué)研究院針對(duì)PCV3cap基因設(shè)計(jì)引物和探針,建立PCV3TaqMan探針熒光定量PCR檢測(cè)方法(見表2)。
4.2.3 DNA測(cè)序
Palinski等、Ku等針對(duì)PCV3基因組不同區(qū)域設(shè)計(jì)擴(kuò)增片段相互重疊引物,借助PCR與測(cè)序技術(shù)獲得PCV3全基因組序列[21,47]。
4.2.4 免疫組化
利用PCV3特異性單克隆抗體染色患病豬組織切片[21],觀察PCV3組織嗜性及其組織內(nèi)分布情況。
4.2.5 ELISA
Palinski等利用原核表達(dá)PCV3 Cap蛋白包被ELISA板,建立檢測(cè)PCV3抗體間接ELISA[21]。
5.1 疫病風(fēng)險(xiǎn)評(píng)估
目前,PCV3感染尚未列入世界動(dòng)物衛(wèi)生組織(OIE)通報(bào)性疫病名錄及《中華人民共和國(guó)進(jìn)境動(dòng)物檢疫疫病名錄》,該病毒隨生豬及其產(chǎn)品國(guó)際貿(mào)易傳播風(fēng)險(xiǎn)尚待評(píng)估。相關(guān)部門有必要根據(jù)該病毒最新研究進(jìn)展開展風(fēng)險(xiǎn)評(píng)估工作,并基于風(fēng)險(xiǎn)評(píng)估結(jié)果作貿(mào)易決定。
5.2 檢測(cè)技術(shù)儲(chǔ)備
鑒于PCV3具有傳播途徑多、傳播速度快、流行范圍廣等特點(diǎn),應(yīng)盡快研發(fā)適用于出入境口岸PCV3快速篩查與準(zhǔn)確鑒定需求的檢疫技術(shù)。
5.3 疫情處置
我國(guó)已出現(xiàn)PCV3感染病例[47,51],主要集中于中東部地區(qū)。全國(guó)范圍內(nèi)急需開展PCV3流行病學(xué)調(diào)查,“早發(fā)現(xiàn)、早報(bào)告、早隔離、早診斷、早撲滅”,及時(shí)控制和撲滅疫情。
5.4 疫苗研制
鑒于PCV2與PCV3存在交叉免疫保護(hù)可能性較低[21,33],應(yīng)加快PCV3毒株分離步伐,同時(shí)利用感染性克隆技術(shù)構(gòu)建含有PCV3 cap基因PCV1或PCV2嵌合病毒,為疫苗生產(chǎn)奠定基礎(chǔ)。
目前針對(duì)PCV3研究尚處于起步階段,需盡快開展相關(guān)檢測(cè)技術(shù)研究,建立PCV3快速、靈敏、特異病原學(xué)和血清學(xué)檢測(cè)方法,分離與鑒定PCV3毒株,為開展疫苗研制及動(dòng)物回歸試驗(yàn)工作奠定基礎(chǔ),保護(hù)我國(guó)生豬產(chǎn)業(yè)健康發(fā)展。
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Research progress on porcine circovirus 3/
ZHANG Yongning1,Mei Lin1,ZHANG
Zhou1,TANG Lijie2,WU Shaoqiang2
(1.Institute of Animal Quarantine,Chinese Academy of Inspection and Quarantine,Beijing 100176,China;2.School of Veterinary Medicine,Northeast Agricultural University,Harbin 150030,China)
In 2016,a novel porcine circovirus,designated porcine circovirus 3(PCV3),was first identified in diseased sows with porcine dermatitis and nephropathy syndrome(PDNS)-like clinical signs as well as in the aborted fetuses by researchers from the Kansas State University using metagenomic sequencing technology.Epidemiological survey further demonstrated that PCV3 infection was mainly distributed in the pig herds exhibiting PDNS and reproductive failure symptoms and was widely prevalent in the swine population in several states of the USA.The Blood Systems Research Institute(San Francisco,USA)also discovered PCV3 in three sick piglets with cardiac and multi-systemic inflammation of unknown etiology.PCV3 was also detected in the Chinese pig herds in Hubei and Guangdong Provinces,etc.The presence of PCV3 infection was successively reported in the Polish and Korean pig populations.As a newly emerging pathogen,the pathogenicity of PCV3 for swine,the role of PCV3 played in porcine circovirusassociated diseases(PCVAD),and its potential influence on the immune efficacy of the existing commercial porcine circovirus 2(PCV2)vaccines need to be further studied.In order to provide a reference for follow-up studies,the latest research progress of PCV3 was summarized in this review.
metagenomic sequencing;porcine circovirus 3(PCV3);porcine dermatitis and nephropathy syndrome(PDNS);reproductive failure;porcine circovirus associated diseases(PCVAD)
S852.659
A
1005-9369(2017)09-0089-08
時(shí)間 2017-10-19 17:14:28 [URL]http://kns.cnki.net/kcms/detail/23.1391.S.20171019.1714.006.
張永寧,梅琳,張舟,等.豬圓環(huán)病毒3型研究進(jìn)展[J].東北農(nóng)業(yè)大學(xué)學(xué)報(bào),2017,48(9):89-96.
Zhang Yongning,Mei Lin,Zhang Zhou,et al.Research progress on Porcine Circovirus 3[J].Journal of Northeast Agricultural University,2017,48(9):89-96.(in Chinese with English abstract)
2017-07-20
“十三五”國(guó)家重點(diǎn)研發(fā)計(jì)劃項(xiàng)目(2016YFD0501102,2017YFD0501105);中國(guó)檢驗(yàn)檢疫科學(xué)研究院“青年英才”計(jì)劃(CAIQ-YC-20140205)
張永寧(1980-),男,副研究員,博士,研究方向?yàn)橥鈦韯?dòng)物疫病檢疫。 E-mail:zhangyn@caiq.gov.cn