董 輝，陸瑤瑤，楊玉榮

DOI：10.3969/j.issn.1002-2694.2017.09.014

綿羊和山羊住肉孢子蟲流行病學(xué)及分型研究進(jìn)展

董 輝，陸瑤瑤，楊玉榮1

動(dòng)物住肉孢子蟲病可引起畜牧養(yǎng)殖業(yè)的經(jīng)濟(jì)損失，影響食品質(zhì)量安全，威脅人類健康。國(guó)際上已確定的綿羊(Ovisaries)住肉孢子蟲有4種：S.tenella、S.arieticanis、S.gigantea、S.medusiformis，山羊(Caprahircus)住肉孢子蟲有3種：S.capracanis、S.hircicanis、S.moulei。本文綜述綿羊和山羊住肉孢子蟲的生活史、國(guó)內(nèi)和國(guó)際的流行情況、致病性、形態(tài)學(xué)特點(diǎn)和診斷方法，以期為中國(guó)綿羊和山羊住肉孢子蟲的生物學(xué)特征研究及其診斷分型提供有力基礎(chǔ)依據(jù)。

住肉孢子蟲；綿羊；山羊；流行病學(xué)；形態(tài)學(xué)；分型; 分子生物學(xué)

住肉孢子蟲(Sarcocystis)為頂復(fù)亞門，孢子蟲綱，真球蟲目，肉孢子蟲科，孢子蟲屬的一種原生性細(xì)胞內(nèi)寄生蟲[1]。住肉孢子蟲在世界分布廣泛，人畜均會(huì)感染[2]。1843年Miescher第1次在鼠橫紋肌中發(fā)現(xiàn)了白色線狀的住肉孢子蟲包囊[3-4]，到目前為止在動(dòng)物中一共發(fā)現(xiàn)有196種住肉孢子蟲[5]。住肉孢子蟲具有嚴(yán)格的宿主選擇性，其生活史需要在中間宿主和終末宿主中完成，中間宿主多為草食動(dòng)物或者雜食動(dòng)物，終末宿主多為肉食動(dòng)物[6-7]。住肉孢子蟲可以在豬、牛和羊等中間宿主的肌肉形成包囊[8]，住肉孢子蟲蟲型和感染數(shù)量的不同，其致病性也具有明顯的差異，可引起家畜的厭食、流產(chǎn)、甚至死亡，人食用含住肉孢子蟲的肉后，會(huì)引起胃腸道疾病。羊住肉孢子蟲病可引起畜牧業(yè)經(jīng)濟(jì)損失，降低肉品質(zhì)量，危害公共健康[9]。本文綜述我國(guó)和國(guó)際上已確定的綿羊和山羊住肉孢子蟲的流行情況、生活史、形態(tài)學(xué)特點(diǎn)和分型、致病性以及分子學(xué)診斷方法，為深入研究國(guó)內(nèi)綿羊和山羊住肉孢子蟲生物學(xué)特征提供有力依據(jù)。

1 生活史

住肉孢子蟲與哈蒙球蟲、弓形蟲和新孢子蟲的[10-12]生活史十分相似，分為有性生殖和無(wú)性生殖。以S.tenalla的生活史為例解釋綿羊和山羊住肉孢子蟲的生活史[4]。當(dāng)犬科動(dòng)物食入含有S.tenella包囊的組織后，包囊在胃和小腸內(nèi)消化釋放出緩殖子，緩殖子在侵入小腸粘膜的過(guò)程中非常有活力，攝入包囊6 HPI(hours post inoculation)，可以在杯狀細(xì)胞內(nèi)發(fā)現(xiàn)緩殖子。然后，緩殖子進(jìn)入配子生殖階段，發(fā)育為雌、雄配子，其數(shù)量比約為95∶5，雌配子和雄配子受精后產(chǎn)生合子，合子外分泌囊壁形成卵囊。卵囊在腸壁內(nèi)進(jìn)行孢子化，每個(gè)孢子化的卵囊內(nèi)含有兩個(gè)孢子囊，每個(gè)孢子囊內(nèi)形成4個(gè)子孢子。孢子化的卵囊壁薄，易于破裂，游離的孢子囊會(huì)進(jìn)入腸腔，隨終末宿主的糞便排出到外界的環(huán)境中[1]。當(dāng)中間宿主綿羊誤食飼料和水中攜帶的S.tenella的卵囊，卵囊進(jìn)入小腸后釋放出的子孢子經(jīng)腸系膜和腸系膜淋巴結(jié)動(dòng)脈，進(jìn)入全身動(dòng)脈血管內(nèi)皮細(xì)胞完成第一代裂殖生殖(9～21 DPI)，然后進(jìn)入全身毛細(xì)血管完成第二代裂殖生殖(16～40 DPI)，蟲血癥時(shí)間為14～32 DPI，然后裂殖子進(jìn)入中樞神經(jīng)組織、心臟形成包囊。

2 流行病學(xué)

2.1 世界各國(guó)的感染情況 綿羊和山羊住肉孢子蟲病在世界分布廣泛，文獻(xiàn)統(tǒng)計(jì)綿羊住肉孢子蟲病陽(yáng)性率為31.42%～93.31%，山羊住肉孢子蟲病陽(yáng)性率為28.30%～83.69%。國(guó)際上廣泛承認(rèn)和確定的綿羊住肉孢子蟲有4種，S.tenella、S.arieticanis、S.gigantea、S.medusiformis。目前S.tenella、S.gigantea、S.arieticanis世界廣泛分布，S.medusiformis只在澳大利亞、新西蘭、伊朗和意大利有報(bào)道[7]。國(guó)際上廣泛承認(rèn)和確定的山羊住肉孢子蟲有S.capracanis、S.hircicanis、S.moulei。山羊住肉孢子蟲在歐洲和北美洲文獻(xiàn)報(bào)道尚少。世界綿羊和山羊住肉孢子蟲感染和分布情況詳見(jiàn)表1。

表1 世界綿羊及山羊住肉孢子蟲病流行情況
Tab.1 Prevalence of Sarcocystis in sheep or goats from the world

宿主Host位置Location陽(yáng)性率Positive%蟲型Species參考文獻(xiàn)References綿羊大洋洲93.31(990/1061)S.tenella、S.arieticanis、S.gigantea、S.medusiformis[1]歐洲31.42(3158/10051)S.tenella、S.arieticanis、S.gigantea、S.medusiformis[1,7][13]非洲54.67(621/1136)S.tenella、S.arieticanis[1]南美36.22(343/947)S.tenella、S.arieticanis、S.gigantea[1,14][12,15]北美67.40(1141/1693)S.tenella、S.arieticanis、S.gigantea[1]亞洲48.02(26061/54270)S.tenella、S.arieticanis、S.gigantea、S.medusiformis[1,4,8,10,16-87]山羊大洋洲28.33(17/60)—[1]非洲76.80(907/1181)S.capracanis[1]南美83.69(585/699)S.capracanis[1,14]亞洲53.89(5522/10246)S.capracanis、S.hircicanis、S.moulei[1,5,6,8,20,26,33,50,51,61,73,80,84,88-91]

2.2 在我國(guó)的流行情況 我國(guó)已確定的綿羊住肉孢子蟲有3種：S.tenella、S.arieticanis、S.gigantea，已確定的山羊住肉孢子蟲有3種：S.hircianis、S.capracanis、S.moulei。綿羊住肉孢子蟲病的陽(yáng)性率在7.74%～100%，山羊住肉孢子蟲病的陽(yáng)性率在9.73%～77.94%。根據(jù)已報(bào)道的綿羊和山羊住肉孢子蟲病的流行病學(xué)調(diào)查，統(tǒng)計(jì)中國(guó)綿羊和山羊的住肉孢子蟲病的分布情況，發(fā)現(xiàn)中國(guó)多地區(qū)羊感染住肉孢子蟲，且感染率較高，見(jiàn)表2。

表2 中國(guó)綿羊和山羊住肉孢子蟲流行病學(xué)調(diào)查
Tab.2 Epidemiological investigation of Sarcocystis in sheep and goats of China

宿主Host地區(qū)Region年份Year陽(yáng)性率Positive%蟲型Species被檢組織Tissue檢測(cè)方法Method參考文獻(xiàn)Reference綿羊北京、河北199954.50(12/22)——MS,EM[26]甘肅19807.74(1152/14891)—MMS[80]黑龍江198591.82(359/391)—D,HGR,MS[81]河南1988-201611.60(105/905)S.arieticanis,S.tenellaD,E,H,MGR,MS,PS[25,48]遼寧1993-200027.83(59/212)—D,E,H,MGR,MS[43-45,82]內(nèi)蒙199957.14(44/77)——MS,EM[26]青海1983-201561.34(7490/12210)S.tenella,S.gigantea,S.arieticanisD,E,H,MGR,PS,PS[27-31,36,37,39-42,46,47,49-74,76-79,83-87]新疆1985-199082.77(5418/6546)S.arieticanisD,H,S,M,GR,MS,PS[32-35,38,75]山羊北京、河北199957.60(19/33)——MS,EM[26]甘肅19809.73(114/1172)—MMS[80]內(nèi)蒙199933.30(12/36)——MS,EM[26]青海1983-201373.52(261/355)S.mouleiD,E,H,MGR,MS[50,51,61,73,84,91]新疆1987-198842.00(5/12)—HMS[33]云南1999,201677.94(212/272)S.hircianis,S.capracanis—MS,PS,EM[5,26]

D: diaphragm 膈肌; E: esophagus 食道肌; H: heart 心肌; M: skeletal muscle 骨骼肌; —:未報(bào)道;

EM: electron microscope 電子顯微鏡; GR: Gross 肉眼觀察; MS: muscle squash 肌肉壓片; PS: paraffin section 石蠟切片

3 致病性

綿羊住肉孢子蟲S.tenella和S.arieticanis的終末宿主為犬科動(dòng)物[7](表3)。S.tenella強(qiáng)致病性，綿羊感染后會(huì)出現(xiàn)厭食、體重減輕、貧血、產(chǎn)奶量降低、早產(chǎn)、流產(chǎn)、神經(jīng)癥狀和死亡等[15,21]。S.arieticanis的致病性稍弱于S.tenalla[21]，感染S.arieticanis可以引起綿羊腹瀉、食欲下降、失去活動(dòng)精神、毛發(fā)減少、虛弱等[4]。S.gigantea和S.medusiformis的終末宿主是貓科動(dòng)物，對(duì)綿羊無(wú)致病性[7]，當(dāng)組織間寄生的住肉孢子蟲包囊破裂時(shí)，引起肌肉的炎癥反應(yīng)將影響肉的品質(zhì)[15]。山羊住肉孢子蟲中，S.capracanis和S.hircicanis的終末宿主是犬科動(dòng)物，S.capracanis致病性強(qiáng)于S.hircicanis，可以引起山羊體溫升高、體重減輕、流產(chǎn)和死亡[6]。S.moulei的終末宿主是貓科動(dòng)物，無(wú)致病性[6]。

綿羊和山羊住肉孢子蟲對(duì)中間宿主的致病性高于終末宿主，通過(guò)犬科動(dòng)物傳播的住肉孢子蟲的致病性要強(qiáng)于通過(guò)貓科動(dòng)物傳播住的肉孢子蟲(對(duì)比見(jiàn)表3)。對(duì)于強(qiáng)致病性的S.tenella和S.arieticanis引起的羊住肉孢子蟲病，其感染嚴(yán)重程度與最初卵囊的攝入量、宿主的免疫狀態(tài)及營(yíng)養(yǎng)狀態(tài)有關(guān)[4,7]，但強(qiáng)致病性的綿羊和山羊住肉孢子蟲對(duì)人沒(méi)有致病性。綿羊可以同時(shí)感染兩種類型的住肉孢子蟲(有致病性和無(wú)致病性的蟲型)[7]。

4 形態(tài)學(xué)特征

4.1 光學(xué)顯微鏡下的形態(tài)結(jié)構(gòu) 很難根據(jù)臨床癥狀診斷住肉孢子蟲病[21]。2016年，Damboriarena就報(bào)道了綿羊食道肌大量隨機(jī)分布的白色橢圓形的S.gigantea包囊[15]，因?yàn)榫d羊和山羊住肉孢子蟲的包囊通常寄生在肌肉組織。住肉孢子蟲與肌纖維平行，形態(tài)呈紡錘形、圓柱形、卵圓型等，顏色呈灰白色、乳白色。S.gigantea和S.medusiformis包囊可以通過(guò)肉眼觀察，S.tenella和S.arieticanis的包囊需要通過(guò)顯微鏡觀察。山羊住肉孢子蟲包囊均可以通過(guò)肉眼觀察。

蛋白酶可以將肌組織和包囊壁消化破解，釋放緩殖子，緩殖子呈月牙或香蕉型，通過(guò)光學(xué)顯微鏡可以檢查住肉孢子蟲的緩殖子[19,21]；組織學(xué)壓片技術(shù)可以觀察到宿主肌肉組織中非肉眼可見(jiàn)的住肉孢子蟲包囊[21]，當(dāng)住肉孢子蟲的感染密度高時(shí)，組織學(xué)壓片方法優(yōu)于其它方法[14]，住肉孢子蟲對(duì)宿主選擇和在組織器官內(nèi)分布均有特異性，用光學(xué)顯微鏡觀察H&E染色的切片可以判定住肉孢子蟲包囊壁的薄厚并觀察包囊壁內(nèi)的緩殖子[21]，所以結(jié)合組織學(xué)方法可以對(duì)住肉孢子蟲進(jìn)鑒別和分型[25]。比如綿羊住肉孢子蟲中S.tenella是厚壁的住肉孢子蟲，寄生于綿羊的心肌和舌肌，S.arieticanis是薄壁的住肉孢子蟲，寄生于心肌，而S.gigantea和S.medusiformis為薄壁的住肉孢子蟲未寄生于心肌。

4.2 超微結(jié)構(gòu) 住肉孢子蟲包囊壁由兩層構(gòu)成，包囊外有囊壁和突起(Villar protrusions, VP)，內(nèi)壁向內(nèi)延伸形成隔，將囊腔分成小室，小室內(nèi)有大量香蕉狀或腎形的緩殖子和母細(xì)胞。包囊壁的超微結(jié)構(gòu)在不同類型的住肉孢子蟲中具有明顯的區(qū)別，目前為止住肉孢子蟲包囊壁的種類至少已有82種[1]。不同住肉孢子蟲包囊壁VP的類型有較大差別[1]，所以通過(guò)電子顯微鏡對(duì)比超微結(jié)構(gòu)的差異是對(duì)住肉孢子蟲分型的一種有效方法[4]，也是住肉孢子分類學(xué)中最為認(rèn)可的手段[7]。綿羊和山羊住肉孢子蟲形態(tài)特征見(jiàn)表3[1,5]。

表3 綿羊及山羊住肉孢子蟲形態(tài)特征
Tab.3 Morphology of the Sarcocystis in the sheep or goats

蟲型Species宿主Host肉眼Gross包囊大小和壁厚度Cystssize(mm)andWal(μm)突起類型Villarprotrusions分布Distribution致病性Pathogenicity膈肌D心肌H中樞神經(jīng)系統(tǒng)CNS綿羊或山羊SheeporGoat人Human終末宿主DefinitiveHostsS.gigantea綿羊可見(jiàn)7.5×5薄壁<2Type21:菜花狀有無(wú)無(wú)無(wú)無(wú)家貓S.medusiformis可見(jiàn)8×0.2薄壁<2Type20:四邊形有無(wú)無(wú)無(wú)無(wú)家貓S.tenella不可見(jiàn)最長(zhǎng)0.7厚壁1～3Type14:柵欄狀無(wú)有有強(qiáng)無(wú)家犬、狼、狐S.arieticanis不可見(jiàn)最長(zhǎng)0.9薄壁<1Type7b:細(xì)絲狀有有無(wú)較強(qiáng)無(wú)家犬S.capracanis山羊可見(jiàn)1×0.1厚壁2-3Type14:柵欄狀有有有有無(wú)家犬、狼、狐S.hircicanis可見(jiàn)最長(zhǎng)2.5薄壁<1Type7a:細(xì)絲狀有有無(wú)無(wú)無(wú)家犬S.moulei可見(jiàn)17×7厚壁3～4Type21:菜花狀有無(wú)無(wú)—無(wú)家貓

D: diaphragm 膈肌；H: heart 心??；CNS: central nervous system 中樞神經(jīng)系統(tǒng)；S: sheep 綿羊；G: goat 山羊；Hu: human 人；—: 未報(bào)道

5 分子生物學(xué)分型

近年來(lái)隨著分子生物學(xué)技術(shù)的發(fā)展，關(guān)于住肉孢子蟲基因序列的研究主要集中在分子流行病學(xué)調(diào)查、分型研究和種系發(fā)育。目前，通過(guò)核糖體小亞基(18S rRNA)、核糖體大亞基(28S rRNA)鑒別住肉孢子蟲較為普遍[7]。內(nèi)轉(zhuǎn)錄間隔區(qū)-ITS(Internal transcribed spacer)、線粒體細(xì)胞色素氧化酶Ⅰ-cox1(Cytochrome C oxidase subunit I)[5,93]也是良好的基因序列研究的選擇。已報(bào)道的關(guān)于綿羊和山羊住肉孢子蟲PCR引物序列，見(jiàn)表4。

6 討 論

國(guó)內(nèi)對(duì)于住肉孢子蟲病的研究多集中在黃牛[94]、水牛，關(guān)于綿羊和山羊住肉孢子蟲的研究多集中于流行病學(xué)方面的研究，關(guān)于其生活史和其它生物學(xué)特征尚未完全證實(shí)。研究住肉孢子蟲，一方面是探究住肉孢子蟲的整體感染情況、季節(jié)動(dòng)態(tài)變化、地理區(qū)域分布，另一方面是明確我國(guó)存在的住肉孢子蟲，對(duì)其分型，從而提高流行病學(xué)調(diào)查的信息多樣性和準(zhǔn)確性，確定是否會(huì)對(duì)畜牧業(yè)及人類的食品安全造成威脅并有效控制該病。

表4 綿羊和山羊住肉孢子蟲PCR引物序列
Tab.4 Sequences of PCR primers of Sarcocystis in sheep or goats

序號(hào)NO.檢測(cè)方法Methods引物Primer片段大小Size(bp)蟲型Species參考文獻(xiàn)References1PCRSAD2F(5'-GGAAGCGATTGGAACC-3')300-350S.capracanis[6]18SrRNASAD2R(5'-CCTTGGTCCGTGTTTCA-3')2PCR1L(5'-CCATGCATGTCTAAGTATAAGC)-3')800S.tenella[14,88]3H(5'-GGCAAATGCTTTCGCAGTAG-3')3PCR-RFLPForward(5'-TTCTATGGCTAATACATGCG-3')722and242S.gigantea[19]Reverse(5'-CCCTAATCCTTCGAAACAGGA-3')901and63S.medusiformis4PCRForward(5'-CGCAAATTACCCAATCCTGA-3')700S.tenellaorS.arieticanis[12]Reverse(5'-ATTTCTCATAAGGTGCAGGAG-3')5Nest-PCRAP2(5'CCCGGGATCCAAGCTTGATCCTTCTGCAGGTTCACCTAC-3')530S.tenella[92]18SrRNAnested(5'-TTTGACTCAACACGGG-3')ST3(5'-CGTTGCCGCGCGTTAA-3')externalST1(5'-GGATCGCATTATGGTCAT-3')6Nest-PCRexternalSTA(5'-TTTCGCAAGGAAGAGGA-3')andSA2(5'-TGAAACGGCGCGTAGA-3')375S.arieticanis[92]18SrRNAinternal(5'-AGGGTTCGATTCCGGAG-3')SA1(5'-GCGGGAAGAGGAGAAT-3')7RT-PCRSar-F1(5'-GCACTTGATGAATTCTGGCA-3')600S.tenella[17]Sar-R1(5'-CACCACCCATAGAATCAAG-3')Sar-antisense:(5'-CCCTAATCCTTCGAAACAGGA-3')

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EpidemiologyandclassificationofSarcocystisinsheepandgoat

DONG Hui, LU Yao-yao, YANG Yu-rong

(CollegeofAnimalScienceandVeterinaryMedicine,HenanAgriculturalUniversity,Zhengzhou450002,China)

Sarcocystiscould cause significant economic losses of animal husbandry, and threaten to public health and the quality of food safety.S.tenella,S.arieticanis,S.giganteaandS.medusiformisare four speciesSarcocystisin sheep (Ovisaries). WhileS.capracanis,S.hircicanisandS.mouleiare three species ofSarcocystisin goat (Caprahircus). The life cycle, epidemiological, pathogenicity, morphological characteristics and diagnostic methods of sheep and goats were reviewed in this paper. The research of biological characteristics ofSarcocystisprovides a powerful basis for diagnosis in sheep and goat in China.

Sarcocystis; sheep; goat; epidemiology; morphology; classification; molecular biology

Yang Yu-rong, Email: yangyu7712@sina.com

國(guó)家重點(diǎn)研發(fā)計(jì)劃(2016YFD0500707)，河南省高校科技創(chuàng)新人才支持計(jì)劃(17HASTIT038)資助項(xiàng)目

楊玉榮, yangyu7712@sina.com

河南農(nóng)業(yè)大學(xué)牧醫(yī)工程學(xué)院, 鄭州 450002

R382.3

：A

：1002-2694(2017)09-0828-09

2017-03-23編輯：劉岱偉

Supported by the National Key Research and Development Program of China (No. 2016YFD0500707) and the Program for Science and Technology Innovation Talents in Universities of Henan Province (No. 17HASTIT038)