建立牛梨形虫病血清学诊断方法及新疆流行病学调查
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摘要
牛梨形虫病(piroplasmosis)主要是由巴贝斯虫、双芽巴贝斯虫和环形泰勒虫引起的一种蜱传性血液原虫病,多种动物可感染,尤其是反刍动物,有时也可感染人。本病共同的临诊特征是发热、贫血、血红蛋白尿,严重者可死亡。按照病程,可将牛的梨形虫病分为急性型和慢性型；按感染类型,可将其分为显性、亚临诊性以及持续性感染。传统的检测和鉴定主要依靠显微镜检查,主要用于急性临诊感染的诊断,但难以进行梨形虫虫种鉴定。在持续性感染的动物中,由于末梢血液中的原虫数量少,容易漏诊。血清学检测可诊断亚临诊性感染和持续性带虫感染,所用的方法主要包括补体结合试验(Complement Fixation Test,CFT)、间接血凝试验(Indirect Hemagglutination Test, IHA)、乳胶凝集试验(Latex Agglutination Test, LAT)、间接荧光抗体试验(Indirect Fluorescent Antibody test, IFAT)、酶联免疫吸附试验(Enzyme-Linked Immunosorbent Assays, ELISA)等,但所使用的诊断抗原大多是粗制的自然抗原,难以克服抗原种间的交叉反应。分子生物学诊断方法例如DNA探针、rRNA探针和PCR方法因辐射、需要专业人员和特殊的实验室材料与仪器等原因仅限于实验室内使用。因此,提高血清学方法的特异性是诊断和鉴定牛梨形虫病的关键所在。目前发现牛巴贝斯虫的裂殖子表面抗原2c (MSA-2c).牛双芽巴贝斯虫的热休克蛋白20(HSP20),牛环形泰勒虫裂殖子表面抗原1(Tams1)的共同特点是保守性好、免疫原性和反应原性强、特异性高,不仅可以作为检测牛梨形虫病的诊断抗原,也可作为预防这些疾病的亚单位疫苗或核酸疫苗的候选者。我国尚未见相关的研究报道。本研究的目的是选择MSA-2C基因、HSP20基因以及Tamsl基因进行克隆与表达,通过优化基因表达获得相应的重组蛋白,初步建立牛梨形虫病的新型血清学诊断方法,并对新疆牛梨形虫病的流行病学进行调查,为制定新疆牛梨形虫病的防治计划提供了第一手资料。通过实验研究得到如下结果：
1.通过分别设计MSA-2c基因、HSP20基因以及Tams1基因序列的特异性引物,经PCR扩增,从血液原虫病患牛的全血样品的基因组DNA中克隆到MSA-2c基因、HSP20基因和Tams1基因片段。又通过设计特异性引物,经RT-PCR扩增,从血液原虫病患牛的全血样品的总RNA中,获得HSP20外显子基因片段。经序列分析发现：(1)克隆出的MSA-2c基因全长798bp,有完整的开放阅读框,与国际标准虫株(德克萨斯株AY052538)的同源性高达99.90%,与其他14个不同国家和地区分离株的MSA-2c基因序列有97.7%的同源性；(2)克隆到HSP20基因和HSP20基因外显子,全长分别为699bp和534bp,与国际标准虫株(墨西哥株)的同源性分别为99.43%和99.63%。(3)克隆的Tams1基因全长846bp,有一个完整的开放阅读框,编码281个氨基酸,与国外虫株的同源性为99.36%～99.80%。系统进化树分析显示,新疆牛环形泰勒虫Tams1基因与印度株、毛利塔里亚株、土耳其、北非共和国株进化途径一致。抗原决定簇分析显示Tams1基因编码的氨基酸具有13个交叉的抗原决定簇。上述结果表明牛巴贝斯虫的MSA-2c基因、牛双芽巴贝斯虫的HSP20基因和HSP20外显子基因以及牛环形泰勒虫Tams1基因都具有高度的保守性。
2.将克隆的MSA-2c基因、HSP20外显子基因以及Tams1基因片段分别再克隆至原核表达载体pGEX-4T-2中,将合成的构建物转入大肠杆菌中,进行IPTG诱导表达。通过优化这些基因表达获得了相应的重组蛋白。SDS-PAGE和Western blot分析发现,MSA-2c基因、HSP20外显子基因以及Tams1基因所表达的融合蛋白分别为55kDa、46kDa、56kDa,它们分别能被抗牛巴贝斯虫、抗双芽巴贝斯虫和抗环形泰勒虫的抗血清所识别。这一结果表明表达出的融合蛋白具有很强的免疫原性和反应原性,能够作为诊断抗原建立牛梨形虫病的新型血清学检测方法。
3.分别以纯化的GST-MSA-2c融合蛋白、GST-HSP20(exon)融合蛋白和GST-Tams1融合蛋白作为诊断抗原,通过优化ELISA反应条件,分别建立了可以检测牛巴贝斯虫、牛双芽巴贝斯虫和牛环形泰勒虫血清特异性抗体的新型间接ELISA方法。方阵试验确定(1)GST-MSA-2c抗原的最适包被浓度为2.5μg/ml,血清最佳稀释倍数为40倍,ELISA阳性反应的临界值为OD450>0.347,批内和批间重复试验的变异系数均小于10%。(2)GST-HSP20抗原的最适包被浓度为5μg/ml,血清最佳稀释倍数为40倍,ELISA阳性反应的临界值为OD450>0.292,批内和批间重复试验的变异系数均小于10%。(3)GST-Tams1抗原的最适包被浓度为10μg/ml,血清最佳稀释倍数为80倍,ELISA阳性反应的临界值为OD450>0.282,批内和批间重复试验的变异系数均小于15%。特异性试验结果为(1) MSA-2c间接ELISA可以检出已知的牛巴贝斯虫阳性血清,对已知的牛双芽巴贝斯虫、环形泰勒虫、东方巴贝斯虫的阳性血清和牛巴贝斯虫阴性血清为阴性。(2)HSP20间接ELISA可以检出已知的双芽巴贝斯虫阳性血清,对已知的牛巴贝斯虫、环形泰勒虫、东方巴贝斯虫的阳性血清和双芽巴贝斯虫阴性血清为阴性。(3)Tams1司接ELISA可以检出已知的牛环形泰勒虫的阳性血清,对已知的牛双芽巴贝斯虫、牛巴贝斯虫虫、东方巴贝斯虫的阳性血清和牛环形泰勒虫的阴性血清为阴性。这些结果表明检测牛梨形虫病的这三种新型间接ELISA方法特异性好,能进行梨形虫虫种的诊断与鉴定。与对应的三种巢式PCR检测结果相比,MSA-2c间接ELISA、HSP20间接ELISA以及Tams1间接ELISA的阳性符合率为96%。上述结果表明所建立的新型重组蛋白间接ELSIA检测方法法重复性好、特异性强、灵敏度高,为大规模地进行新疆牛梨形虫病的流行病学调查和血清学诊断提供有效的技术手段。
4.使用所建立的牛梨形虫病新型血清学诊断方法对2006年～2008年新疆14个地州的牛梨形虫病流行病学进行了调查。检测结果显示(1)新疆存在着牛巴贝斯虫病。在2006年采集的278份牛血清样品中,阳性血清7份,感染率为2.52%。在2007年的532份牛血清样品中检出阳性血清32份,感染率为3.13%。在2008年的530份牛血清中检出阳性血清43份,感染率为5.28%。2008年,在地方流行性疫病区牛巴贝斯虫感染率高达28%。牛巴贝斯虫病所在的地州由2006年的2个扩大到2008年的9个。(2)新疆还存在着牛双芽巴贝斯虫病,而且比牛巴贝斯虫病严重。在2006年采集的278份牛血清样品中,阳性血清11份,感染率为5.40%。在2007年的532份牛血清样品中检出阳性血清25份,感染率为4.70%。在2008年的530份牛血清中检出阳性血清53份,感染率为7.17%。2008年,在地方流行性疫病区牛巴贝斯虫感染率高达30%。牛双芽巴贝斯虫病所在的地州由2006年的8个扩大到2008年的13个。(3)牛环形泰勒虫病仍是新疆牛梨形病的主要病原。在2006年采集的278份牛血清样品中,阳性血清31份,感染率为11.15%。在2007年的532份牛血清样品中检出阳性血清61份,感染率为11.47%。在2008年的530份牛血清中检出阳性血清61份,感染率为11.51%。2008年,在地方流行性疫病区牛环形泰勒虫感染率高达24%。(4)在三年采集的1340份血清样品中共检出牛梨形虫阳性血清280例,感染率为20.90%,其中牛巴贝斯虫、双芽巴贝斯虫、泰勒梨形虫单一感染率分别为8.21%(23/280),15.71%(44/280)和43.93%(123/280)。新疆牛梨形虫混合感染为15.35%,甚至出现三重感染现象。2008年在流行区内的牛梨形虫感染率高达74%(37/50),单一感染与混合感染率分别为64.85%(24/37)和35.15%(13/37),流行区的混合感染率明显增高。这些结果表明新疆牛梨形虫病由原来的一种牛环形泰勒虫病增加到三种牛梨形虫病,其流行病学特征为感染率逐年增加,疫区不断扩大,混合感染使疫情趋于复杂化,对动物和人类构成一定的威胁,不容忽视。
本研究是新疆首次利用新型血清学方法对全疆牛梨形虫病进行大规模的流行病学调查,初步掌握了新疆梨形虫病流行的第一手数据和流行特点,为今后制定包括牛巴贝斯虫病、牛双芽巴贝斯虫病和牛环形泰勒虫在内的新疆牛梨形虫防控计划和综合防治措施的制定提供了依据。
The bovine piroplasmosis is a tick-born hemoparasitic disease which mainly caused by Babesia bovis, Babesia bigemina and Theilaria annulata. Many kinds of animals can be infected, especially in ruminants, sometime in man. The common clinical symptoms are fever, anaemia, haemoglobinuria, even death in severe cases. The bovine piroplasmosis can be divided into acute and chronic infections, according to the course of disease, and can also be grouped into clinical, sub-clinic and persistent infections by their infection types. The traditional detection and identification of piroplasms depends mainly on microscope, especially for the acute and clinical infection, but is hard for the sub-clinical and persistent infected cases, because the number of hemoparasites in the peripheral blood is too low to detect and easily to make misdiagnosis. The sub-clinical, chronic and persistent infected cases, even carriers can be detected by serological methods, such as complement fixation test (CFT), indirect hemagglutination test (IHA), latex agglutination test (LAT), indirect fluorescent antibody test (IFAT), and enzyme-linked immunosorbent assay (ELISA), etc. The crude, natural proteins as diagnostic antigens are used in the most of existing serological tests for bovine piroplasmosis which are hard to eliminate the cross reactions among the species of piroplasms. The tests of molecular biology, such as DNA probes, rRNA probe and PCR have highly sensitive and specific tools for diagnosis of bovine piroplasmosis, but still hard to use as routine method, because vitriol conditions, such as irradiation, professional personnel and special laboratory materials and equipment, are unsuitable for use in the field. Therefore, it is an important turning point to improve the specificity of serological tests for detection of specific antibodies against bovine piroplasmas. Recently years, some specific antigens have been found, such as merozoite surface antigen-2c (MSA-2c) in Babesia bovis, heat shock protein20(HSP20) in Babesia bigemina, and merozoite/piroplasm surface antigen1(Tams1) in Theileria annulata, etc. The common features for the antigens are highly conserved and immunogenic. Some scientists postulates that they may be not only the useful diagnostic candidates for detection of bovine piroplasmosis, but also the useful subunit or NDA vaccine candidates for improved control of bovine piroplasmosis. At present, the reports of the same or relative studies have not been found in China.
The purpose of this study are:(1) to clone and express the MSA-2c, HSP20and Tams1and to obtain three fusion proteins by the optimal conditions for expression of these genes;(2) to develop the improved indirect enzyme-linked immunosorbent assays (iELISAs) using recombinant proteins as diagnostic antigens;(3) to perform the epidemiologic surveys of bovine piroplasmosis in Xinjiang;(4) to obtain the firsthand information for development of prevent and control program against bovine piroplasmosis in the future. The results from this study are as following:
1. Three genes MSA-2c, HSP20and Tams1were cloned from extracted genome DNA in bovine blood samples infected with hemoparasite in Xinjiang by PCR with specific designed primers, respectively, and HSP20exon gene was loned from extracted total RNA in bovine blood samples infected with hemoparasites by RT-PCR with specific designed primers. The sequence analysis was shown:(1) the total length of MSA-2c gene was798kb, and had an intact open reading frame encoding281amino acids. It shared99.90%nucleotide homology with that of type strain, Texas AY052538, and also shared97.7%nucleotide homology with that of stains isolated from other different countries and regions.(2) The total lengths of HSP20gene and HSP20exon gene were699kb and534kb, respectively, and they shared99.43%and99.63%nucleotide homology, respectively, with those of the Mexico type strain.(3) The total length of Tamsl gene was846kb, and had an intact open reading frame encoding281amino acids. It shared99.36%-99.80%nucleotide homology with that published in GenBank. Analysis of phylogenetic tree was shown that it was closely related to others isolated from India, Mauritania, Turkey and Tunisia. Analysis of antigen determinants was shown that it had13putative cross-antigen determinants.
The results above mentioned suggested these genes MSA-2c for B. bovis, HSP20/HSP20(exon) for B. bigemina and Tamsl for T. annulata were highly conserved.
2. The cloned MSA-2c, HSP20(exon) and Tamsl genes were sub-cloned into expression vector pGEX-4T-2and the resultant constructs were respectively transformed into E. coli cells and induced by IPTG. SDS-PAGE and Western blot analysis were shown that fussion proteins of55kDa for MSA-2c gene,46kDa for HSP20(exon) and56kDa for Tamsl were expressed which could be recognized by bovine antisera against B. bovis, B. bigemina and T. annulata, respectively. These results were indicated the expressed fusion proteins by these genes possessed strong immunogenicity and reactionogenicity, which could be used as recombinant antigens to develop the improved serological tests for detection of specific antibodies against bovine piroplasmosis.
3.Three improved indirect enzyme-linked immunosorbent assays (indirect ELISAs) for the detection of specific antibody against bovine piroplasmosis were developed using MSA-2c, HSP20(exon) and Tams1fusion proteins after optimizing its reacting conditions.(1) For indirect MSA-2c ELISA, the optimal concentration of coating recombinant antigen was2.5μg/ml and the optimal dilution of serum sample was1:40in the cross assay. The cutoff was chosen as an OD450>0.347for positive response. The variation coefficient of intra-batch and the inter-batch in the repeating tests was less than10%.(2) For indirect HSP20(exon) ELISA, The optimal concentration of coating recombinant antigen was5μg/ml and the optimal dilution of serum sample was1:40in the cross assay. The cutoff was chosen as an OD450>0.292for positive response. The variation coefficient of intra-batch and the inter-batch in the repeating tests was less than10%.(3) For indirect Tams1ELISA, the optimal concentration of coating antigen was10μg/ml and the optimal dilution of serum sample was1:80in the cross assay. The cutoff was chosen as an OD450≥0.282for positive response. The variation coefficient of intra-batch and the inter-batch in the repeating tests was less than15%.
The results of specificity for the three indirect ELISAs were shown that (1) positive serum samples for B. bovis identified by Npcr were still detected positive in the indirect MSA2c ELISA, but not from the sera of B. bigemina, T. annulata and B. orientalis, and negative control sera.(2) Identified positive serum samples for B. bigemina were detected positive in the indirect HSP20(exon) ELISA, but not from the sera of B. bovis, T. annulata and B. orientalis, and negative control sera.(3) Identified positive serum samples for T. annulata were detected positive in the indirect Tamsl ELISA, not for sera of B. bigemina, B. bovis and B. orientalis, and negative control sera. The concordances of identified positive serum samples for bovine piroplasmosis in indirect MSA-2c, HSP20(exon) and Tamsl ELISAs were96%(14/16, n=50),96%(9/11, n=50) and96%(21/23, n=50), respectively, comparing with those by nested-PCR. The results above mentioned were shown three improved indirect ELISAs were highly sensitive and specific for the diagnosis and identification of bovine piroplsmas, which could be useful tools of epidemiological survey for detection of bovine piroplasmosis in large scale.
4. The epidemiologic survey for detection of bovine piroplasmosis in Xinjiang by the three improved indirect ELISAs was performed in this study. The results were shown that (1) Bovine babesiosis caused by B. bovis was presented in Xinjiang which was first time to be confirmed by the serological test. The prevalence for B. bovis from the tested cattle in2006,2007and2008were2.25%(7/278),3.31%(32/532) and5.28%(43/530), respectively. The rate of B. bovis infection was increased to28%(14/50) in some endemic regions where the disease occurred in2008. The number of infected prefectures in Xinjiang was also increased from2in2006to9in2008.(2) Bovine babesiosis caused by B. bigemina was also presented in Xinjiang, which was first time to be confirmed by the serological test. The infection rates for B. bigemina in2006,2007and2008were5.40%(11/278),4.70%(25/532) and7.17%(53/530), respectively. The infection rate was increased to30%(15/50) in some endemic regions where the disease occurred in2008. The number of infected prefectures in Xinjiang was also increased from8in2006to13in2008.(3) Bovine theileriosis caused by T. annulata was still the main hemoparasitic disease in cattle of Xinjiang. The infection rates in2006,2007and2008were11.15%(31/278),11.47%(61/532) and11.51%(61/530), respectively. The infection rate was increased to24%(15/50) in some regions where the disease occurred in2008.(4)280positive serum examples for bovine piroplasmosis were detected from1340sera collected from14prefectures in Xinjiang during2006to2008. The total infection rate reached to20.90%(280/1340). of those, the infection rates of bovine piroplasmosis caused by B. bovis, B. bigemina and T. annulata alone were8.92%(23/280),15.71%(44/280) and43.92%(123/280), respectively, during that time. The rate of mix infections among them was15.35%(43/280) which included some triple infections. The infection rate was unexpectedly increased to74%(37/50) in some endemic regions where the disease occurred in2008. The mixed infections reached to35.15%(13/50) in that areas. These results were indicated the category of bovine piroplasmosis was increased from1kind of hemoparasitic disease caused by T. annulata in past years to3kinds of ones caused by T. annulata, B. bigemina and B. bovis, respectively. The features of bovine piroplasmosis in Xinjiang were that rate of single infection and mixed infections were increased year by year, which made epidemic situations more complicated. It was more dangerous for animal and human health and a question which could be disregarded.
This is a first time to perform the epidemiological survey in large-scale for detection the specific antibodies against bovine piroplasmosis in Xinjiang by improved indirect ELISAs using recombinant proteins as diagnostic antigens and the first time that B. bovis and B. bigemina have been detected in Xinjiang. We basically obtain the firsthand informa-tion for development of prevent and control program against bovine piroplasmosis in the future in Xinjiang.
1.通过分别设计MSA-2c基因、HSP20基因以及Tams1基因序列的特异性引物,经PCR扩增,从血液原虫病患牛的全血样品的基因组DNA中克隆到MSA-2c基因、HSP20基因和Tams1基因片段。又通过设计特异性引物,经RT-PCR扩增,从血液原虫病患牛的全血样品的总RNA中,获得HSP20外显子基因片段。经序列分析发现：(1)克隆出的MSA-2c基因全长798bp,有完整的开放阅读框,与国际标准虫株(德克萨斯株AY052538)的同源性高达99.90%,与其他14个不同国家和地区分离株的MSA-2c基因序列有97.7%的同源性；(2)克隆到HSP20基因和HSP20基因外显子,全长分别为699bp和534bp,与国际标准虫株(墨西哥株)的同源性分别为99.43%和99.63%。(3)克隆的Tams1基因全长846bp,有一个完整的开放阅读框,编码281个氨基酸,与国外虫株的同源性为99.36%～99.80%。系统进化树分析显示,新疆牛环形泰勒虫Tams1基因与印度株、毛利塔里亚株、土耳其、北非共和国株进化途径一致。抗原决定簇分析显示Tams1基因编码的氨基酸具有13个交叉的抗原决定簇。上述结果表明牛巴贝斯虫的MSA-2c基因、牛双芽巴贝斯虫的HSP20基因和HSP20外显子基因以及牛环形泰勒虫Tams1基因都具有高度的保守性。
2.将克隆的MSA-2c基因、HSP20外显子基因以及Tams1基因片段分别再克隆至原核表达载体pGEX-4T-2中,将合成的构建物转入大肠杆菌中,进行IPTG诱导表达。通过优化这些基因表达获得了相应的重组蛋白。SDS-PAGE和Western blot分析发现,MSA-2c基因、HSP20外显子基因以及Tams1基因所表达的融合蛋白分别为55kDa、46kDa、56kDa,它们分别能被抗牛巴贝斯虫、抗双芽巴贝斯虫和抗环形泰勒虫的抗血清所识别。这一结果表明表达出的融合蛋白具有很强的免疫原性和反应原性,能够作为诊断抗原建立牛梨形虫病的新型血清学检测方法。
3.分别以纯化的GST-MSA-2c融合蛋白、GST-HSP20(exon)融合蛋白和GST-Tams1融合蛋白作为诊断抗原,通过优化ELISA反应条件,分别建立了可以检测牛巴贝斯虫、牛双芽巴贝斯虫和牛环形泰勒虫血清特异性抗体的新型间接ELISA方法。方阵试验确定(1)GST-MSA-2c抗原的最适包被浓度为2.5μg/ml,血清最佳稀释倍数为40倍,ELISA阳性反应的临界值为OD450>0.347,批内和批间重复试验的变异系数均小于10%。(2)GST-HSP20抗原的最适包被浓度为5μg/ml,血清最佳稀释倍数为40倍,ELISA阳性反应的临界值为OD450>0.292,批内和批间重复试验的变异系数均小于10%。(3)GST-Tams1抗原的最适包被浓度为10μg/ml,血清最佳稀释倍数为80倍,ELISA阳性反应的临界值为OD450>0.282,批内和批间重复试验的变异系数均小于15%。特异性试验结果为(1) MSA-2c间接ELISA可以检出已知的牛巴贝斯虫阳性血清,对已知的牛双芽巴贝斯虫、环形泰勒虫、东方巴贝斯虫的阳性血清和牛巴贝斯虫阴性血清为阴性。(2)HSP20间接ELISA可以检出已知的双芽巴贝斯虫阳性血清,对已知的牛巴贝斯虫、环形泰勒虫、东方巴贝斯虫的阳性血清和双芽巴贝斯虫阴性血清为阴性。(3)Tams1司接ELISA可以检出已知的牛环形泰勒虫的阳性血清,对已知的牛双芽巴贝斯虫、牛巴贝斯虫虫、东方巴贝斯虫的阳性血清和牛环形泰勒虫的阴性血清为阴性。这些结果表明检测牛梨形虫病的这三种新型间接ELISA方法特异性好,能进行梨形虫虫种的诊断与鉴定。与对应的三种巢式PCR检测结果相比,MSA-2c间接ELISA、HSP20间接ELISA以及Tams1间接ELISA的阳性符合率为96%。上述结果表明所建立的新型重组蛋白间接ELSIA检测方法法重复性好、特异性强、灵敏度高,为大规模地进行新疆牛梨形虫病的流行病学调查和血清学诊断提供有效的技术手段。
4.使用所建立的牛梨形虫病新型血清学诊断方法对2006年～2008年新疆14个地州的牛梨形虫病流行病学进行了调查。检测结果显示(1)新疆存在着牛巴贝斯虫病。在2006年采集的278份牛血清样品中,阳性血清7份,感染率为2.52%。在2007年的532份牛血清样品中检出阳性血清32份,感染率为3.13%。在2008年的530份牛血清中检出阳性血清43份,感染率为5.28%。2008年,在地方流行性疫病区牛巴贝斯虫感染率高达28%。牛巴贝斯虫病所在的地州由2006年的2个扩大到2008年的9个。(2)新疆还存在着牛双芽巴贝斯虫病,而且比牛巴贝斯虫病严重。在2006年采集的278份牛血清样品中,阳性血清11份,感染率为5.40%。在2007年的532份牛血清样品中检出阳性血清25份,感染率为4.70%。在2008年的530份牛血清中检出阳性血清53份,感染率为7.17%。2008年,在地方流行性疫病区牛巴贝斯虫感染率高达30%。牛双芽巴贝斯虫病所在的地州由2006年的8个扩大到2008年的13个。(3)牛环形泰勒虫病仍是新疆牛梨形病的主要病原。在2006年采集的278份牛血清样品中,阳性血清31份,感染率为11.15%。在2007年的532份牛血清样品中检出阳性血清61份,感染率为11.47%。在2008年的530份牛血清中检出阳性血清61份,感染率为11.51%。2008年,在地方流行性疫病区牛环形泰勒虫感染率高达24%。(4)在三年采集的1340份血清样品中共检出牛梨形虫阳性血清280例,感染率为20.90%,其中牛巴贝斯虫、双芽巴贝斯虫、泰勒梨形虫单一感染率分别为8.21%(23/280),15.71%(44/280)和43.93%(123/280)。新疆牛梨形虫混合感染为15.35%,甚至出现三重感染现象。2008年在流行区内的牛梨形虫感染率高达74%(37/50),单一感染与混合感染率分别为64.85%(24/37)和35.15%(13/37),流行区的混合感染率明显增高。这些结果表明新疆牛梨形虫病由原来的一种牛环形泰勒虫病增加到三种牛梨形虫病,其流行病学特征为感染率逐年增加,疫区不断扩大,混合感染使疫情趋于复杂化,对动物和人类构成一定的威胁,不容忽视。
本研究是新疆首次利用新型血清学方法对全疆牛梨形虫病进行大规模的流行病学调查,初步掌握了新疆梨形虫病流行的第一手数据和流行特点,为今后制定包括牛巴贝斯虫病、牛双芽巴贝斯虫病和牛环形泰勒虫在内的新疆牛梨形虫防控计划和综合防治措施的制定提供了依据。
The bovine piroplasmosis is a tick-born hemoparasitic disease which mainly caused by Babesia bovis, Babesia bigemina and Theilaria annulata. Many kinds of animals can be infected, especially in ruminants, sometime in man. The common clinical symptoms are fever, anaemia, haemoglobinuria, even death in severe cases. The bovine piroplasmosis can be divided into acute and chronic infections, according to the course of disease, and can also be grouped into clinical, sub-clinic and persistent infections by their infection types. The traditional detection and identification of piroplasms depends mainly on microscope, especially for the acute and clinical infection, but is hard for the sub-clinical and persistent infected cases, because the number of hemoparasites in the peripheral blood is too low to detect and easily to make misdiagnosis. The sub-clinical, chronic and persistent infected cases, even carriers can be detected by serological methods, such as complement fixation test (CFT), indirect hemagglutination test (IHA), latex agglutination test (LAT), indirect fluorescent antibody test (IFAT), and enzyme-linked immunosorbent assay (ELISA), etc. The crude, natural proteins as diagnostic antigens are used in the most of existing serological tests for bovine piroplasmosis which are hard to eliminate the cross reactions among the species of piroplasms. The tests of molecular biology, such as DNA probes, rRNA probe and PCR have highly sensitive and specific tools for diagnosis of bovine piroplasmosis, but still hard to use as routine method, because vitriol conditions, such as irradiation, professional personnel and special laboratory materials and equipment, are unsuitable for use in the field. Therefore, it is an important turning point to improve the specificity of serological tests for detection of specific antibodies against bovine piroplasmas. Recently years, some specific antigens have been found, such as merozoite surface antigen-2c (MSA-2c) in Babesia bovis, heat shock protein20(HSP20) in Babesia bigemina, and merozoite/piroplasm surface antigen1(Tams1) in Theileria annulata, etc. The common features for the antigens are highly conserved and immunogenic. Some scientists postulates that they may be not only the useful diagnostic candidates for detection of bovine piroplasmosis, but also the useful subunit or NDA vaccine candidates for improved control of bovine piroplasmosis. At present, the reports of the same or relative studies have not been found in China.
The purpose of this study are:(1) to clone and express the MSA-2c, HSP20and Tams1and to obtain three fusion proteins by the optimal conditions for expression of these genes;(2) to develop the improved indirect enzyme-linked immunosorbent assays (iELISAs) using recombinant proteins as diagnostic antigens;(3) to perform the epidemiologic surveys of bovine piroplasmosis in Xinjiang;(4) to obtain the firsthand information for development of prevent and control program against bovine piroplasmosis in the future. The results from this study are as following:
1. Three genes MSA-2c, HSP20and Tams1were cloned from extracted genome DNA in bovine blood samples infected with hemoparasite in Xinjiang by PCR with specific designed primers, respectively, and HSP20exon gene was loned from extracted total RNA in bovine blood samples infected with hemoparasites by RT-PCR with specific designed primers. The sequence analysis was shown:(1) the total length of MSA-2c gene was798kb, and had an intact open reading frame encoding281amino acids. It shared99.90%nucleotide homology with that of type strain, Texas AY052538, and also shared97.7%nucleotide homology with that of stains isolated from other different countries and regions.(2) The total lengths of HSP20gene and HSP20exon gene were699kb and534kb, respectively, and they shared99.43%and99.63%nucleotide homology, respectively, with those of the Mexico type strain.(3) The total length of Tamsl gene was846kb, and had an intact open reading frame encoding281amino acids. It shared99.36%-99.80%nucleotide homology with that published in GenBank. Analysis of phylogenetic tree was shown that it was closely related to others isolated from India, Mauritania, Turkey and Tunisia. Analysis of antigen determinants was shown that it had13putative cross-antigen determinants.
The results above mentioned suggested these genes MSA-2c for B. bovis, HSP20/HSP20(exon) for B. bigemina and Tamsl for T. annulata were highly conserved.
2. The cloned MSA-2c, HSP20(exon) and Tamsl genes were sub-cloned into expression vector pGEX-4T-2and the resultant constructs were respectively transformed into E. coli cells and induced by IPTG. SDS-PAGE and Western blot analysis were shown that fussion proteins of55kDa for MSA-2c gene,46kDa for HSP20(exon) and56kDa for Tamsl were expressed which could be recognized by bovine antisera against B. bovis, B. bigemina and T. annulata, respectively. These results were indicated the expressed fusion proteins by these genes possessed strong immunogenicity and reactionogenicity, which could be used as recombinant antigens to develop the improved serological tests for detection of specific antibodies against bovine piroplasmosis.
3.Three improved indirect enzyme-linked immunosorbent assays (indirect ELISAs) for the detection of specific antibody against bovine piroplasmosis were developed using MSA-2c, HSP20(exon) and Tams1fusion proteins after optimizing its reacting conditions.(1) For indirect MSA-2c ELISA, the optimal concentration of coating recombinant antigen was2.5μg/ml and the optimal dilution of serum sample was1:40in the cross assay. The cutoff was chosen as an OD450>0.347for positive response. The variation coefficient of intra-batch and the inter-batch in the repeating tests was less than10%.(2) For indirect HSP20(exon) ELISA, The optimal concentration of coating recombinant antigen was5μg/ml and the optimal dilution of serum sample was1:40in the cross assay. The cutoff was chosen as an OD450>0.292for positive response. The variation coefficient of intra-batch and the inter-batch in the repeating tests was less than10%.(3) For indirect Tams1ELISA, the optimal concentration of coating antigen was10μg/ml and the optimal dilution of serum sample was1:80in the cross assay. The cutoff was chosen as an OD450≥0.282for positive response. The variation coefficient of intra-batch and the inter-batch in the repeating tests was less than15%.
The results of specificity for the three indirect ELISAs were shown that (1) positive serum samples for B. bovis identified by Npcr were still detected positive in the indirect MSA2c ELISA, but not from the sera of B. bigemina, T. annulata and B. orientalis, and negative control sera.(2) Identified positive serum samples for B. bigemina were detected positive in the indirect HSP20(exon) ELISA, but not from the sera of B. bovis, T. annulata and B. orientalis, and negative control sera.(3) Identified positive serum samples for T. annulata were detected positive in the indirect Tamsl ELISA, not for sera of B. bigemina, B. bovis and B. orientalis, and negative control sera. The concordances of identified positive serum samples for bovine piroplasmosis in indirect MSA-2c, HSP20(exon) and Tamsl ELISAs were96%(14/16, n=50),96%(9/11, n=50) and96%(21/23, n=50), respectively, comparing with those by nested-PCR. The results above mentioned were shown three improved indirect ELISAs were highly sensitive and specific for the diagnosis and identification of bovine piroplsmas, which could be useful tools of epidemiological survey for detection of bovine piroplasmosis in large scale.
4. The epidemiologic survey for detection of bovine piroplasmosis in Xinjiang by the three improved indirect ELISAs was performed in this study. The results were shown that (1) Bovine babesiosis caused by B. bovis was presented in Xinjiang which was first time to be confirmed by the serological test. The prevalence for B. bovis from the tested cattle in2006,2007and2008were2.25%(7/278),3.31%(32/532) and5.28%(43/530), respectively. The rate of B. bovis infection was increased to28%(14/50) in some endemic regions where the disease occurred in2008. The number of infected prefectures in Xinjiang was also increased from2in2006to9in2008.(2) Bovine babesiosis caused by B. bigemina was also presented in Xinjiang, which was first time to be confirmed by the serological test. The infection rates for B. bigemina in2006,2007and2008were5.40%(11/278),4.70%(25/532) and7.17%(53/530), respectively. The infection rate was increased to30%(15/50) in some endemic regions where the disease occurred in2008. The number of infected prefectures in Xinjiang was also increased from8in2006to13in2008.(3) Bovine theileriosis caused by T. annulata was still the main hemoparasitic disease in cattle of Xinjiang. The infection rates in2006,2007and2008were11.15%(31/278),11.47%(61/532) and11.51%(61/530), respectively. The infection rate was increased to24%(15/50) in some regions where the disease occurred in2008.(4)280positive serum examples for bovine piroplasmosis were detected from1340sera collected from14prefectures in Xinjiang during2006to2008. The total infection rate reached to20.90%(280/1340). of those, the infection rates of bovine piroplasmosis caused by B. bovis, B. bigemina and T. annulata alone were8.92%(23/280),15.71%(44/280) and43.92%(123/280), respectively, during that time. The rate of mix infections among them was15.35%(43/280) which included some triple infections. The infection rate was unexpectedly increased to74%(37/50) in some endemic regions where the disease occurred in2008. The mixed infections reached to35.15%(13/50) in that areas. These results were indicated the category of bovine piroplasmosis was increased from1kind of hemoparasitic disease caused by T. annulata in past years to3kinds of ones caused by T. annulata, B. bigemina and B. bovis, respectively. The features of bovine piroplasmosis in Xinjiang were that rate of single infection and mixed infections were increased year by year, which made epidemic situations more complicated. It was more dangerous for animal and human health and a question which could be disregarded.
This is a first time to perform the epidemiological survey in large-scale for detection the specific antibodies against bovine piroplasmosis in Xinjiang by improved indirect ELISAs using recombinant proteins as diagnostic antigens and the first time that B. bovis and B. bigemina have been detected in Xinjiang. We basically obtain the firsthand informa-tion for development of prevent and control program against bovine piroplasmosis in the future in Xinjiang.
引文
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