网状内皮组织增生症病毒LAMP检测方法及感染性分子克隆的建立
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摘要
网状内皮组织增生症(Reticuloendotheliosis,RE)是禽类一种重要的免疫抑制性和致瘤性疾病。近年来,国内外鸡群中REV流行逐渐严重。REV与马立克氏病病毒(MDV)、鸡贫血病毒(CAV)和J-亚群禽白血病病毒(ALV-J)几种免疫抑制性病毒的混合感染在部分养鸡场中普遍存在,使得鸡群免疫力低下,导致禽流感、新城疫等重要疫苗免疫失败,并容易造成继发感染,使疫病的诊断和防控难度加大。另外,REV基因组易于整合到MDV和FPV基因组中,从而导致商品化疫苗受到污染。使用非SPF禽胚制备的疫苗也有污染REV的潜在危险。近来,RE的危害正逐渐被人们关注,但RE的诊断、流行病学以及基础研究仍不完善。为了更好地认识和防控RE,本文进行了如下方面的研究:
1 REV LAMP可视化快速检测方法的建立
借助新兴的环介导等温扩增技术(loop-mediated isothermal amplification, LAMP),设计了4条针对REV pol基因的6个位点的特异性引物,建立了一套简便、灵敏和特异的检测方法。该方法不与常见的感染鸡群的DNA病毒(MDV、CAV)及ALV-J发生交叉反应,比常规的PCR方法灵敏25倍;另外,该LAMP方法在临床样品方面的检测效率与传统的PCR方法相当;而且操作简单,不需要复杂的仪器,常规水浴锅即可进行,肉眼可直接观察检测结果。
2 REV gp90蛋白的表达及多克隆抗体的制备
利用PCR技术,从病料中扩增REV gp90基因,将其分别克隆至真核表达载体pFastBacHTA和原核表达载体pET-28a(+)中,从而构建了真核表达的供体质粒pF-gp90和原核表达质粒pET28-gp90。pF-gp90转化大肠杆菌DH10Bac感受态细胞,使gp90基因整合到Bacmid穿梭载体中,获得重组穿梭载体Bacmidgp90。通过脂质体介导将其转染Sf9昆虫细胞,获得重组杆状病毒rBacgp90。通过Western blot和IFA验证,rBacgp90可正确表达gp90蛋白,蛋白大小约为45kDa,变性和自然状态下均能够被识别,具有良好的生物活性。将pET28-gp90转化至大肠杆菌BL21 (DE3)感受态细胞,经IPTG诱导,进行gp90蛋白原核表达并通过Western blot检测,确定其以包涵体形式正确表达。纯化gp90蛋白并免疫6-8周龄Balb/c小鼠,制备抗gp90的多克隆抗体。免疫小鼠后获得的抗血清可与感染CEF细胞的REV发生特异性反应,ELISA效价达到1:12800。
3 RE流行病学研究
对我国9个省35个蛋鸡场进行了血清学调查,89%的鸡场REV抗体阳性,阳性率为1%-98%,平均阳性率为15%,说明REV在我国广泛流行,部分地区较为严重。将研磨后无菌处理的病料接种于CEF或DF1,置37℃CO_2培养箱培养,7d后收毒,连续盲传至少3代。做间接免疫荧光、PCR和电镜检测,确定是否分离到病毒。从东北地区的黑龙江、吉林、辽宁等省的一些鸡场分离到10株REV。以HLJR0901为例,测得其TCID_(50)为10~(5.2)/mL。通过绘制复制动力学曲线对其在CEF细胞上的增殖动态规律进行了研究,发现第六天和第七天REV增殖的滴度最高。将所分离的REV的主要保护性抗原基因-gp90基因,克隆至pMD-18T载体,测序并上传GenBank。以前病毒cDNA为模板,将HLJR0901基因组分六段进行了克隆和测序。最后确定:HLJR0901全长8284bp,GenBank登录号为GQ415646。序列分析结果显示:HLJR0901的LTR序列与FA株完全同源;与APC-566的同源性也高达99%。Pol是最稳定基因。LTR核苷酸、gag氨基酸、pol氨基酸、gp90氨基酸、env氨基酸以及全基因组核苷酸的遗传进化树显现了一个共同的特征:①所比较的毒株明显地分为三个分枝,分别代表REV的3个亚型;②东北分离株形成一个独立的群体,与我国早期南方毒株HA9901(亚型I)同源性较低,可能源于不同的祖先;③我国至少存在两种REV亚型(I型和III型),III型REV是目前东北地区的流行毒株。④东北分离株与目前美国和台湾的一些流行毒株亲缘关系较近。
4 REV感染性分子克隆的建立及应用
通过重叠延伸PCR (SOE PCR)和酶切拼接将HLJR0901的全基因组克隆于pBluescript II载体中,并在基因组中通过改变核苷酸(C2250G)消除了其中的MscI酶切位点作为分子标签,构建了REV感染性分子克隆pBlu-mHLJR0901。将纯化的pBlu-mHLJR0901转染鸡胚成纤维细胞(CEF)进行病毒拯救,经间接免疫荧光(IFA)、PCR、测序及电镜鉴定,成功拯救出REV。拯救病毒rHLJR0901在CEF细胞上具有与亲本毒相似的复制特性。基于感染性分子克隆,开展了表达EGFP的重组REV的构建:在HLJR0901全基因组934位点(gag基因启动子上游)和7713位点(env基因终止子下游)插入了EGFP基因,转染CEF细胞后,前者EGFP能够瞬时高效表达,后者不能表达。但最终均没有拯救出重组病毒,其具体机制有待进一步探讨。本研究还发现,REV LTR的启动子功能具有广谱性,不仅在鸡源细胞上能够被识别,而且在哺乳细胞上也能被识别。这些结果为REV的基础研究提供了参考。
5 REV动物感染实验
将拯救的REV毒株rHLJR0901通过腹腔接种途径感染1日龄SPF雏鸡,进行REV的致病性研究。结果显示:2周龄时REV能够明显影响鸡的生长性能;REV对肝脏、脾脏、胸腺、法氏囊等免疫相关器官均有一定程度的影响;感染后3w时血清开始阳转,但血清抗体阳性率一直是75%;整个实验周期内均可检测到病毒血症。动物感染实验为REV病毒感染模型的建立提供了必要的数据,并提示病毒血症可以作为REV病毒感染模型的稳定指标之一。
Reticuloendotheliosis is an important immunosuppressive and oncogenic disease in avian species. Recently, REV is prevailing in avian in the world. Co-infection of REV with subgroup J avian leukosis virus (ALV-J), Marek’s disease virus (MDV), and chicken anemia virus (CAV) has become endemic in layer and breeder flocks, which induce immunosuppressive and immuno-failure of some important disease (AIV, NDV, et al) so that secondary infection. So it is very difficult to diagnose and control REV. In addition, REV genome can integrate into MDV and fowl poxvirus (FPV) genome and contaminate these commercial vaccines. Besides, the vaccines developed with non SPF embryonated egg are also easy to be contaminated with REV. Now, the threat posed by REV has been got more and more attention. However, the diagnosis, epidemiology and basic research are not consummate. In order to further understand and control RE in a new efficient level, the following experiments were performed in this study.
1 Development of LAMP assay for REV
The convenient, specific and sensitive LAMP assay for detecting REV was developed, in which 4 primers aiming at six sites to amplify the pol gene. The REV LAMP assay did not cross react with common avian DNA viruses (MDV, CAV) and ALV-J. Additionally, the assay could detect different REV strains and has a detection limit of 5 copies, which has a 25 times sensitivity than traditional PCR methods. Furthermore, the efficiency of LAMP to detect REV in clinical samples was comparable to traditional PCR. The procedure of LAMP was straight-forward, not rely on any special equipment and performance was simple. The result of LAMP could be observed directly by the unaided eye.
2 Expression of gp90 protein and development of anti-gp90 monoclonal antibody
The gp90 gene was amplified by PCR and then cloned into pFastBacHTA, pET-28a (+) , respectively. The donator plamid pF-gp90 of eukaryotic expression and the prokaryotic expression plasmid pET28-gp90 were acquired. The recombinant donor plasmid pF-gp90 was transformed into E coli DH10Bac. The recombinant bacmid plasmid Bacmidgp90 was obtained by integrating gp90 gene into Bacmid vector in DH10Bac cells. The recombinant baculovirus rBacgp90 was obtained after transfecting Sf9 cells. Western blot and IFA showed that the gp90 protein was expressed successfully and its weight was about 45kDa. It has good reactionogenicity. The plasmid pET28-gp90 was transformed into E coli BL21 (DE3) and was induced by IPTG. Analysis by SDS-PAGE and Western blot showed that the gp90 protein was expressed successfully in the form of inclusion body. Then the gp90 protein was purified and immunized to 6-8 weeks Balb/c mice to prepare anti-gp90 serum. The protein has good immunogenicity and the mouse anti-gp90 serum with the ELISA titer of 1:12800 could recognize REV.
3 Research of RE epidemiology
Serological survey was performed in nine province. Serum samples were collected from thirty-five layer farms and REV antibody were detected by IDEXX ELISA kit. 89% farms were positive for REV antibody. The positive rate in each farms were 1%-98% and the average positive rate was 15%. These data suggested REV was popular in China and some farms were serious. Grinded and sterilized, the samples were inoculated CEF or DF1 and cultured in 37℃CO_2 incubator for 7 days. Virus was collected and blind passaged. Finally, REV was identificated by indirect immunofluorescence assay (IFA), PCR, electron micrograph. We isolated 10 REV strains from some chicken farm of Heilongjiang, jilin, liaoning, et al. The TCID_(50) of HLJ0901 strain was tittered as 10~(5.2)/mL. With the replication kinetics curve, the regular pattern of HLJR0901 replication in CEF cells was researched: the highest titer of HLJR0901 was observed at 6 and 7 days post-infection. Gp90 gene of fourteen strains, encoding the most important protective protein, was amplification and cloned into pMD-18T vector, then were sequenced and submitted to GenBnak. Six overlapping fragments covering the full-genome of HLJR0901 were cloned and sequenced. The full-genome sequence of HLJR0901 (8284bp) was ascertained. The GenBank accession number is GQ415646. Sequence analysis showed that the LTR of HLJR0901 was completely homologous with FA, and has 99% homology with APC-566. Pol was the most stable gene. The phylogenetic trees of LTR nucleotide, gag amino acids, pol amino acids, gp90 amino acids, env amino acids and the genome displayed a common character:①all the strains was diveided into three distinct branches, which represents three subtypes, respectively.②the northeast REV isolates formed a distinct cluster far from the early Chinese strain HA9901.(subtype I), which derive from different ancestors.③at least two subtypes of REV (subtype I and III) are popular in China, the northeast REV belong to subtype III.④the northeast REV isolates have high similarity with some popular strains in American and Taiwan.
4 Development and application of the infectious molecular clone of REV
The full-length genome of the HLJR0901 strain was jointed and cloned in the pBluescript II vector by SOE PCR and restriction enzyme, and the genetic tag C2250G (deleting the MscI site) was introduced. The infectious molecular clone pBlu-mHLJR0901 was constructed. pBlu-mHLJR0901 was transfected and then mind-blinded in CEF cells. With identification of IFA, PCR, sequencing, electron microscope assay, REV could be rescued efficiently. The rescued REV has similar characters of replication and infection. Based on the reverse genetics of REV,the construction of recombinant REV expressing EGFP was performed. EGFP was inserted into the 934 and 7713 nucleotide site of HLJR0901 genome, which was lied in the upper stream before gag gene promoter and the down stream after env gene terminator, respectively. After being transfected into CEF cells, EGFP was transient expressed transient with high performance in the former but not in the later. No recombinant virus was rescued. We also observed that the promoter activity of REV LTR was broad-spectrum not only in chicken cells but also in pig cell. The experiments provide some reference for the basic researches of REV.
5 Animal experiment of REV
The REV-infection-animal model with the resued rHLJR0901 was studied preliminarily. Infected 1d SPF chicken, REV could influence the growth and induced a little lesion in liver, spleen, thymus and bursa. Seroconversion was observed at 3w post-infection, but the positive rate was 75% till 8w post-infection. Viremia existed during all experiment period. The experiments provide necessary data for REV-infection-animal model. Viremia will be a stable index for REV-infection-animal model.
1 REV LAMP可视化快速检测方法的建立
借助新兴的环介导等温扩增技术(loop-mediated isothermal amplification, LAMP),设计了4条针对REV pol基因的6个位点的特异性引物,建立了一套简便、灵敏和特异的检测方法。该方法不与常见的感染鸡群的DNA病毒(MDV、CAV)及ALV-J发生交叉反应,比常规的PCR方法灵敏25倍;另外,该LAMP方法在临床样品方面的检测效率与传统的PCR方法相当;而且操作简单,不需要复杂的仪器,常规水浴锅即可进行,肉眼可直接观察检测结果。
2 REV gp90蛋白的表达及多克隆抗体的制备
利用PCR技术,从病料中扩增REV gp90基因,将其分别克隆至真核表达载体pFastBacHTA和原核表达载体pET-28a(+)中,从而构建了真核表达的供体质粒pF-gp90和原核表达质粒pET28-gp90。pF-gp90转化大肠杆菌DH10Bac感受态细胞,使gp90基因整合到Bacmid穿梭载体中,获得重组穿梭载体Bacmidgp90。通过脂质体介导将其转染Sf9昆虫细胞,获得重组杆状病毒rBacgp90。通过Western blot和IFA验证,rBacgp90可正确表达gp90蛋白,蛋白大小约为45kDa,变性和自然状态下均能够被识别,具有良好的生物活性。将pET28-gp90转化至大肠杆菌BL21 (DE3)感受态细胞,经IPTG诱导,进行gp90蛋白原核表达并通过Western blot检测,确定其以包涵体形式正确表达。纯化gp90蛋白并免疫6-8周龄Balb/c小鼠,制备抗gp90的多克隆抗体。免疫小鼠后获得的抗血清可与感染CEF细胞的REV发生特异性反应,ELISA效价达到1:12800。
3 RE流行病学研究
对我国9个省35个蛋鸡场进行了血清学调查,89%的鸡场REV抗体阳性,阳性率为1%-98%,平均阳性率为15%,说明REV在我国广泛流行,部分地区较为严重。将研磨后无菌处理的病料接种于CEF或DF1,置37℃CO_2培养箱培养,7d后收毒,连续盲传至少3代。做间接免疫荧光、PCR和电镜检测,确定是否分离到病毒。从东北地区的黑龙江、吉林、辽宁等省的一些鸡场分离到10株REV。以HLJR0901为例,测得其TCID_(50)为10~(5.2)/mL。通过绘制复制动力学曲线对其在CEF细胞上的增殖动态规律进行了研究,发现第六天和第七天REV增殖的滴度最高。将所分离的REV的主要保护性抗原基因-gp90基因,克隆至pMD-18T载体,测序并上传GenBank。以前病毒cDNA为模板,将HLJR0901基因组分六段进行了克隆和测序。最后确定:HLJR0901全长8284bp,GenBank登录号为GQ415646。序列分析结果显示:HLJR0901的LTR序列与FA株完全同源;与APC-566的同源性也高达99%。Pol是最稳定基因。LTR核苷酸、gag氨基酸、pol氨基酸、gp90氨基酸、env氨基酸以及全基因组核苷酸的遗传进化树显现了一个共同的特征:①所比较的毒株明显地分为三个分枝,分别代表REV的3个亚型;②东北分离株形成一个独立的群体,与我国早期南方毒株HA9901(亚型I)同源性较低,可能源于不同的祖先;③我国至少存在两种REV亚型(I型和III型),III型REV是目前东北地区的流行毒株。④东北分离株与目前美国和台湾的一些流行毒株亲缘关系较近。
4 REV感染性分子克隆的建立及应用
通过重叠延伸PCR (SOE PCR)和酶切拼接将HLJR0901的全基因组克隆于pBluescript II载体中,并在基因组中通过改变核苷酸(C2250G)消除了其中的MscI酶切位点作为分子标签,构建了REV感染性分子克隆pBlu-mHLJR0901。将纯化的pBlu-mHLJR0901转染鸡胚成纤维细胞(CEF)进行病毒拯救,经间接免疫荧光(IFA)、PCR、测序及电镜鉴定,成功拯救出REV。拯救病毒rHLJR0901在CEF细胞上具有与亲本毒相似的复制特性。基于感染性分子克隆,开展了表达EGFP的重组REV的构建:在HLJR0901全基因组934位点(gag基因启动子上游)和7713位点(env基因终止子下游)插入了EGFP基因,转染CEF细胞后,前者EGFP能够瞬时高效表达,后者不能表达。但最终均没有拯救出重组病毒,其具体机制有待进一步探讨。本研究还发现,REV LTR的启动子功能具有广谱性,不仅在鸡源细胞上能够被识别,而且在哺乳细胞上也能被识别。这些结果为REV的基础研究提供了参考。
5 REV动物感染实验
将拯救的REV毒株rHLJR0901通过腹腔接种途径感染1日龄SPF雏鸡,进行REV的致病性研究。结果显示:2周龄时REV能够明显影响鸡的生长性能;REV对肝脏、脾脏、胸腺、法氏囊等免疫相关器官均有一定程度的影响;感染后3w时血清开始阳转,但血清抗体阳性率一直是75%;整个实验周期内均可检测到病毒血症。动物感染实验为REV病毒感染模型的建立提供了必要的数据,并提示病毒血症可以作为REV病毒感染模型的稳定指标之一。
Reticuloendotheliosis is an important immunosuppressive and oncogenic disease in avian species. Recently, REV is prevailing in avian in the world. Co-infection of REV with subgroup J avian leukosis virus (ALV-J), Marek’s disease virus (MDV), and chicken anemia virus (CAV) has become endemic in layer and breeder flocks, which induce immunosuppressive and immuno-failure of some important disease (AIV, NDV, et al) so that secondary infection. So it is very difficult to diagnose and control REV. In addition, REV genome can integrate into MDV and fowl poxvirus (FPV) genome and contaminate these commercial vaccines. Besides, the vaccines developed with non SPF embryonated egg are also easy to be contaminated with REV. Now, the threat posed by REV has been got more and more attention. However, the diagnosis, epidemiology and basic research are not consummate. In order to further understand and control RE in a new efficient level, the following experiments were performed in this study.
1 Development of LAMP assay for REV
The convenient, specific and sensitive LAMP assay for detecting REV was developed, in which 4 primers aiming at six sites to amplify the pol gene. The REV LAMP assay did not cross react with common avian DNA viruses (MDV, CAV) and ALV-J. Additionally, the assay could detect different REV strains and has a detection limit of 5 copies, which has a 25 times sensitivity than traditional PCR methods. Furthermore, the efficiency of LAMP to detect REV in clinical samples was comparable to traditional PCR. The procedure of LAMP was straight-forward, not rely on any special equipment and performance was simple. The result of LAMP could be observed directly by the unaided eye.
2 Expression of gp90 protein and development of anti-gp90 monoclonal antibody
The gp90 gene was amplified by PCR and then cloned into pFastBacHTA, pET-28a (+) , respectively. The donator plamid pF-gp90 of eukaryotic expression and the prokaryotic expression plasmid pET28-gp90 were acquired. The recombinant donor plasmid pF-gp90 was transformed into E coli DH10Bac. The recombinant bacmid plasmid Bacmidgp90 was obtained by integrating gp90 gene into Bacmid vector in DH10Bac cells. The recombinant baculovirus rBacgp90 was obtained after transfecting Sf9 cells. Western blot and IFA showed that the gp90 protein was expressed successfully and its weight was about 45kDa. It has good reactionogenicity. The plasmid pET28-gp90 was transformed into E coli BL21 (DE3) and was induced by IPTG. Analysis by SDS-PAGE and Western blot showed that the gp90 protein was expressed successfully in the form of inclusion body. Then the gp90 protein was purified and immunized to 6-8 weeks Balb/c mice to prepare anti-gp90 serum. The protein has good immunogenicity and the mouse anti-gp90 serum with the ELISA titer of 1:12800 could recognize REV.
3 Research of RE epidemiology
Serological survey was performed in nine province. Serum samples were collected from thirty-five layer farms and REV antibody were detected by IDEXX ELISA kit. 89% farms were positive for REV antibody. The positive rate in each farms were 1%-98% and the average positive rate was 15%. These data suggested REV was popular in China and some farms were serious. Grinded and sterilized, the samples were inoculated CEF or DF1 and cultured in 37℃CO_2 incubator for 7 days. Virus was collected and blind passaged. Finally, REV was identificated by indirect immunofluorescence assay (IFA), PCR, electron micrograph. We isolated 10 REV strains from some chicken farm of Heilongjiang, jilin, liaoning, et al. The TCID_(50) of HLJ0901 strain was tittered as 10~(5.2)/mL. With the replication kinetics curve, the regular pattern of HLJR0901 replication in CEF cells was researched: the highest titer of HLJR0901 was observed at 6 and 7 days post-infection. Gp90 gene of fourteen strains, encoding the most important protective protein, was amplification and cloned into pMD-18T vector, then were sequenced and submitted to GenBnak. Six overlapping fragments covering the full-genome of HLJR0901 were cloned and sequenced. The full-genome sequence of HLJR0901 (8284bp) was ascertained. The GenBank accession number is GQ415646. Sequence analysis showed that the LTR of HLJR0901 was completely homologous with FA, and has 99% homology with APC-566. Pol was the most stable gene. The phylogenetic trees of LTR nucleotide, gag amino acids, pol amino acids, gp90 amino acids, env amino acids and the genome displayed a common character:①all the strains was diveided into three distinct branches, which represents three subtypes, respectively.②the northeast REV isolates formed a distinct cluster far from the early Chinese strain HA9901.(subtype I), which derive from different ancestors.③at least two subtypes of REV (subtype I and III) are popular in China, the northeast REV belong to subtype III.④the northeast REV isolates have high similarity with some popular strains in American and Taiwan.
4 Development and application of the infectious molecular clone of REV
The full-length genome of the HLJR0901 strain was jointed and cloned in the pBluescript II vector by SOE PCR and restriction enzyme, and the genetic tag C2250G (deleting the MscI site) was introduced. The infectious molecular clone pBlu-mHLJR0901 was constructed. pBlu-mHLJR0901 was transfected and then mind-blinded in CEF cells. With identification of IFA, PCR, sequencing, electron microscope assay, REV could be rescued efficiently. The rescued REV has similar characters of replication and infection. Based on the reverse genetics of REV,the construction of recombinant REV expressing EGFP was performed. EGFP was inserted into the 934 and 7713 nucleotide site of HLJR0901 genome, which was lied in the upper stream before gag gene promoter and the down stream after env gene terminator, respectively. After being transfected into CEF cells, EGFP was transient expressed transient with high performance in the former but not in the later. No recombinant virus was rescued. We also observed that the promoter activity of REV LTR was broad-spectrum not only in chicken cells but also in pig cell. The experiments provide some reference for the basic researches of REV.
5 Animal experiment of REV
The REV-infection-animal model with the resued rHLJR0901 was studied preliminarily. Infected 1d SPF chicken, REV could influence the growth and induced a little lesion in liver, spleen, thymus and bursa. Seroconversion was observed at 3w post-infection, but the positive rate was 75% till 8w post-infection. Viremia existed during all experiment period. The experiments provide necessary data for REV-infection-animal model. Viremia will be a stable index for REV-infection-animal model.
引文
阿合买提·买买提,杨丽萍,刘忠贵. 2000. SPF雏鸡感染REV后外周血液中免疫球蛋白含量和免疫器官与体重比值的动态变化[J].黑龙江畜牧兽医. 1:1~2.
崔治中. 2006.禽反转录病毒与DNA病毒间的基因重组及其流行病学意义[J].病毒学报. 22:150~154.
崔治中. 2008.鸭群中REV感染的流行病学调查[J].微生物学报. 48(4): 514~519.
崔治中,孟珊珊,姜世金等. 2006.我国白羽肉用型鸡群中CAV、REV和REOV感染状况的血清学调查[J].畜牧兽医学报. 37(2): 152~157.
崔治中,孙淑红,王辉等. 2009.鸡群中马立克氏病毒、J-亚群白血病毒和网状内皮增生病毒的多重感染[C].中英家禽肿瘤病理及肿瘤病毒研讨会.山东农业大学.
崔治中,孙怀昌,朱承如. 1987.禽白血病及网状内皮增生病感染情况的调查[J].中国畜禽传染病. l:37~38.
丁家波,崔治中,于立娟. 2004.含有禽网状内皮组织增生病病毒基因组片段的天然重组禽痘病毒的研究[J].微生物学报. 44:588~592.
杜岩,崔治中. 2001.禽网状内皮组织增生病病毒感染引起的鸡群免疫抑制[J].中国兽医学报. 21(2):122~124.
段宏安, Ibrahim Diallo , Peter Spradbrow ,等. 1999.应用聚合酶链式反应检测禽痘病毒田间分离株及疫苗株中网状内皮组织增殖病毒LTR片段, env及rel基因[J].中国兽医杂志. 25 (8): 9~10.
何宏虎,陈溥言,蔡宝祥. 1988.禽网状内皮组织增生病病毒的分离鉴定[J].中国畜禽传染病. 1~2.
何勇群,张中直,杨汉春.1998.北京地区鸡群网状内皮组织增殖病感染的血清学调查研究[J].畜牧兽医学报. 29:71~77.
胡北侠,黄艳艳,路希山等. 2009.肉种禽网状内皮增生症、鸡传染性贫血和禽白血病血清学调查[J].广东畜牧兽医科技. 34(1): 25~27.
吉荣,崔治中,丁家波. 2003.网状内皮组织增生病病毒囊膜糖蛋白gp90的原核表达[J]. 中国兽医学报. 23(1): 28~30.
吉荣,崔治中,王锡乐等. 2005.分子克隆化禽网状内皮组织增生症病毒传染性及其前病毒全基因组序列研究[J].病毒学报. 21:448~455.
吉荣,刘岳龙,秦爱建. 2001.免疫抑制鸡群鸡传染性贫血病毒和网状内皮增生症病毒共感染检测[J].中国兽药杂志. 35(4):1~3.
姜世金,孟珊珊,崔治中等. 2005.我国自然发病鸡群中MDV、REV和CAV共感染的检测[J].中国病毒学. 20:164~167.
金文杰,崔治中,刘岳龙等.2001.传染性法氏囊病病料中MDV、CAV、REV的共感染检测[J].中国兽医学报. 1:6~9.
匡宝晓,徐福南.1997.网状内皮组织增殖病病毒感染鸡红细胞免疫功能的动态变化[J].中国兽医科技. 27:7~9.
李广兴,刘忠贵,郎跃深. 2000. SPF雏鸡感染网状内皮组织增殖病病毒后白细胞介素2和干扰素诱生活性的变化[J].中国兽医科技. 30:3~5.
李凯,高宏雷,高立等. 2010.禽网状内皮组织增生病病毒TaqMan探针荧光定量PCR检测方法的建立及应用[J].中国兽医科学. 11:1137-1140.
李延鹏,崔治中,姜世金等. 2008.CAV与REV共感染SPF鸡对疫苗免疫反应的抑制作用[J].中国兽医学报. 28(11): 1243~1246.
孟珊珊,崔治中,孙淑红. 2006. REV和ALV-J共感染鸡病毒血症及抗体反应的相互影响[J].中国兽医学报. 26(4): 363-366.
秦立廷,高玉龙,潘伟等. 2010.我国部分地区蛋鸡群ALV~J及与REV、MDV、CAV混合感染检测[J].中国预防兽医学报. 32(2): 90~93.
荣骏弓,王笑梅,幸桂香等. 1997.禽网状内皮组织增殖病琼扩免疫扩散抗原的研制及应用[J].中国畜禽传染病. 5: 29~33.
孙淑红. 2007.博士论文:禽网状内皮组织增生症病毒和J-亚群白血病病毒的致病性及其疫苗研究.山东农业大学. 15-30.
王玉. 2005.博士论文:网状内皮组织增生症病毒分子流行病学调查及其在宿主免疫选择压下的分子变异.山东农业大学. 88-100.
王玉,崔志中,姜世金. 2006.网状内皮组织增生症病毒中国分离株HA9901全基因组核苷酸序列的测定与分析[J].中国科学C辑. 49(2): 149~157.
杨汉春,高云. 1997.应用改良ELISA检测禽网状内皮增殖病抗体血清[J].中国兽医杂志.23:3~5.
殷震,刘景华. 1997.动物病毒学(第二版).科学出版社. 329~331.
于立娟,崔治中. 2006.禽痘疫苗病毒中网状内皮组织增生病病毒5’LTR整合位点序列分析[J].微生物学报. 46:660~662.
郁晓岚,徐福南,蔡宝祥等. 1989.禽网状内皮增殖病组织病理学和超微结构的动态变化[J].南京农业大学学报. 12:91~96.
张志,王锡乐,庄国庆等. 2005.用单抗介导的免疫荧光试验检测组织切片中的禽网状内皮组织增生病病毒[J].中国兽医学报. 25(2) :122~124.
张志,崔治中,姜世金. 2004.从J亚群禽白血病肿瘤中检测出网状内皮组织增生病病毒[J].中国兽医学报. 1:10~23.
瞿晓兰,王宏俊,陈小玲. 2006.禽网状内皮组织增生病毒囊膜蛋白基因的克隆及其原核表达[J ].华北农学报. 21(增刊):154 ~157.
赵丽青,吴延功,王志亮. 2006.禽网状内皮组织增殖病的特异性荧光PCR检测方法的研究[C].湖南长沙:中国畜牧兽医学会禽病学分会第13次学术研讨会论文集. 10: 442~444.
Alien PT, Mullins JA, Harris CL, et al. 1979. Replication of reticuloendotheliosis virus in mammalian cells(abstract) [C]. Amer Soc Microbiol Annual Meeting. 100:256~256.
Aly MM, Fadly AM, Witter RL. 1992. Effects of serotype 2 Marek,s disease virus on development of viremia, antibody, and lymphoma induced by reticuloendotheliosis virus.4thInternational Symposium on Marek,s disease,19th world,s Poultry Congress, world,s Poultry Science Assn[C]. Amsterdam. l:272~276.
Aly MM, Hassan MK, Elzahr AA, et al. 1998. Serological survey on retieuloendotheliosis virus infection in comercial chicken and turkey flocks in EgyPt. Proceedings of the 5th Science Conference [C]. Egypt Vet Poultry Association. 51~68.
Aly MM, Smith EJ, Fadly AM. 1993. Detection of reticuloendotheliosis virus infection using the polymerase chain reaction[J]. Avian Pathology. 22:543~554.
Bachrach E, Duch M, Pelegrin M, et al. 2003. In vivo infection of mice by replication-competent MLV based retroviral vectors[J]. Methods Mol. Med. 76:343~352.
Bachrach E, Pelegrin M, Piechaczyk M, et al. 2002. Efficient gene transfer into spleen cells of newborn mice by a replication-competent retroviral vector. Virology. 293:328~334.
Bagust TJ, Grimes TM, Dennett DP. 1979. Infection studies on a reticuloendotheliosis virus contaminant of a commercial Marek’s disease vaccine [J]. Aust. Vet. J. 55: 153~157.
Bagust TJ, Grimes TM, Ratnamohan N. 1981. Experimental infection of chickens with an Australian strain of reticuloendotheliosis virus 3 Persistent infection and transmission by the adult hen[J]. Avian Patholoy. 10:375~385.
Barbosa T, Zavala G, Cheng S, et al. 2007. Full genome sequence and some biological properties of reticuloendotheliosis virus strain APC-566 isolated from endangered Attwater's prairie chickens[J]. Virus Res. 124, 68~77.
Barth CF, Humphries EH. 1988. Expression of v-relinduces mature B-cell lines that reflect the diversity of avian immunoglobulin heavy and light-chain rearrangements[J]. Molecular and Cellular Biology. 8:5358~5368.
Baxter-Gabbard KL, Campbell WF, Padgett F. 1971. Avian reticuloendotheliosis virus (strainT).Ⅱ.Biochemical and biophysical properties[J]. Avian Disease 15:850~862.
Berglund P, Pleeton MN, Smerdou C, et al. 1999. Immunization with recombinant Semliki Forest virus induces protection against influenza challenge in mice[J]. Vaccine.17 (5): 497~507.
Berglund P, Smerdou C, Fleeton MN, et al. 1998. Enhancing immune responses using suicidal DNA vaccines[J]. Nat Biotechnol.16 (6): 562~565.
Bohls RL, Linares JA, Gross SL, et al. 2006. Phylogenetic analyses indicate little variation among reticuloendotheliosis viruses infecting avian species, including the endangered Attwater’s prairie chicken[J]. Virus Res. 119:187~194.
Bose HR, Levine AS.1967. Replication of the reticuloendotheliosis virus (strainT) in chicken embryo cell culture[J]. Journal of Virology. l:1117~1121.
Breitman ML, Lai MMC, Vogt PK.1980. Attenuation of avian reticuloendotheliosis virus: Loss of the defective transforming component during serial passage of oncogenic virus in fibroblasts[J]. Virology. 101:304~306.
Celine G, Howard MT. 1986. Nondefective spleen necrosis virus-derived vectors define the upper size limit for packaging reticuloendotheliosis viruses[J]. Proc Nati Acad Sci. 83: 9211-9215.
Chang Z, Pan J, Logg C, et al. 2001. A replication-competent feline leukemia virus, subgroup A (FeLV-A), tagged with green fluorescent protein reporter exhibits in vitro biological properties similar to those of the parental FeLV-A[J]. J Virol. 75: 8837~ 8841.
Chen IS, Malk TW, O’Pear JJ, et al.1981. Characterization of retieuloendotheliosis virus strainT/DNA and isolation of a novel variant of reticuloendotheliosis virus strain T by molecular cloning[J]. J Virol. 40:800~811.
Chen PY, Cui ZZ, Lee LF, et al. 1987. Serologic difference among nondefective reticuloendotheliosis viruses[J]. Arch. Virol. 93:233~246.
Chen HT, Zhang J, Sun DH, et al. 2008. Rapid detection of porcine circovirus type 2 by loop-mediated isothermal amplification[J]. J Virol Methods. 149: 264~268.
Cheng WH, Huang YP, Wang CH. 2006. Serological and virological surveys of reticuloendotheliosis in chickens in Taiwan [J]. J Vet Med Sci. 68(12):1315~1320.
Coffin JM.1982. Structure of the retroviral genome. RNA tumor viruses. Molecular Biology of Tumor Viruses[J]. 2 Cold Spring Harbor Laboratory. Cold Spring Harbor. NY: 261~368.
Cohen M, Rein A, Stephens R.M, et al. 1981. Baboon endogenous virus genome.Ⅲ.non- Molecular cloning and structural characterization of defective viral genomes from the DNA of ababoon cell strain[J]. Proc.National Academy Science. 78:5207~5211.
Cook MK. 1969. Cultivation of filterable agent associated with Marek,s disease[J]. Joumal of the National Cancer Institute. 43:203~212.
Crittenden LB, Fadly AM, Smith EJ.1982. Effect of endogenous leukosis virus genes on response to infection with avian leucosis and reticuloendotheliosis virus[J]. Avian Disease. 26:279~294.
Cui ZZ, Sun SH, Zhang Z, et al. 2009a. Simultaneous endemic infections with subgroup J avian leukosis virus and reticuloendotheliosis virus in commercial and local breeds of chickens[J]. Avian Pathol. 38(6):443~448.
Cui ZZ, Zhuang G, Xu X, et al. 2009b. Molecular and biological characterization of a Marek’s disease virus field strain with reticuloendotheliosis virus LTR insert[J]. Virus gene. 437~445. Cui ZZ, Lee L Fetc. 1988. Monoclonal antibody mediated enzyme linked immunosorbent assay for detection of reticuloendotheliosis virus [J]. Avian Dis. 32: 32~40.
Davidson I, Borovskays A, Peri S, et al. 1995. Use of the polymerase chain reaction for the diagnosis of natural infection chickens and turkeys with Marek’s disease virus and reticuloendotheliosis virus[J]. Avian Pathology. 24: 69~94.
Davidson I, Borenshtain R. 2004. In vivo events of retroviral long terminal repeat integration into Marek's disease virus in commercial poultry: detection of chimeric molecules as a marker[J]. Avian Dis. 45: 102~121.
Davidson I, Yang H, Witter RL, et al. 1995. The immunodominant proteins of reticuloendotheliosis virus [J]. Vet Microbiol. 49: 273~284.
Diallo IS, Mackenzie MA, Spradbrow PB, et al. 1998. Field isolates of fowlpox viruscontaminated with reticuloendotheliosis virus[J]. Avian Pathol. 27: 60~66.
Dubensky T. W., Driver D. A., Polo J. M. 1996. Sindbis virus DNA-based expression vectors: utility for in vitro and in vivo gene transfer. J. Virol.70 (1): 508-519.
Fadly AM, Witter RL, Smith EJ, et al. 1996. An outbreak of lymphomas in commercial broiler breeder chickens vaccinated with a fowlpox vaccine contaminated with reticuloendotheliosis virus[J]. Avian Pathol. 25: 35~47.
Fadly AM, Witter RL. 1997.Comparative evaluation of in vitro and in vivo assays for the detection of reticuloendotheliosis virus as a contaminant in a live virus vaccine of poultry[J]. Avian Dis. 41: 695~701.
Fadly AM, Garcia MC. 2006. Detection of reticuloendotheliosis virus in live virus vaccines of poultry[J]. Dev Biol (Basel) 126: 301~305.
Fleeton M N, Liljestrom P, Sheahan B J, et al. Recombinant Semliki Forest virus particles expressing louping ill virus antigens induce a better protective response than plasmid-based DNA vaccines or an inactivated whole particle vaccine. J Gen Virol., 2000, 81 (3): 749-758.
Fritsch RB, Kang CY, Wan KMM, et al. 1977. Transformation of chick embryo fibroblasts by reticuloendotheliosis virus[J]. Virology. 83:313~321.
García MN, Narang N, Reed WM, et al. 2003. Molecular characterization of reticuloendotheliosis virus insertions in the genome of field and vaccine strains of fowl poxvirus[J]. Avian Dis. 47: 343~354.
Ge J Y, Deng G H, Wen Z Y, et al. 2007. Newcastle disease virus-based live attenuated vaccine completely protects chickens and mice from lethal challenge of homologous and heterologous H5N1 avian influenza viruses. J. Virol. 81 (1): 150~158.
Guillermo Zavala, Sunny Cheng, Taylor Barbosa, et al. 2006. Enzootic Reticuloendotheliosis in the Endangered Attwater’s and Greater Prairie Chickens[J]. Avian Dis. 50:520~525.
Hauck R, Prusas C, Hafez HM, et al. 2009. Serologic response against fowl poxvirus and reticuloendotheliosis virus after experimental and natural infections of chickens with fowl poxvirus[J]. Avian Dis. 53(2):205~210.
Hertig C, Coupar BE, Gould AR, et al. 1997. Field and vaccine strains of fowlpox virus carry integrated sequences from the avian retrovirus, reticuloendotheliosis virus[J]. Virology. 235: 367~376.
Hoelzer JD, Franklin RB, Bose JHR.1979. Transformation by reticuloendotheliosis virus: development of a focus assay and isolation of a non-transforming virus. Virology[J]. 93:20~30.
Hu Sylvia SF, Lai Michael MC, Wong TC. 1981. Avian reticuloendotheliosis virus : characterization of genome structure by heteroduplex mapping [J]. J Virol Methods. 37 (3): 899~907.
Humphries EH, Zhang G.1992. V-rel and C-rel modulate the expression of both bursal and non-bursal antigens on avian B-cell lymphomas. Current Topics in Microbiology and Immunology[J]. 182: 475~483.
Ianconescu M.1977. Reticuloendotheliosis antigen for the agar gel precipitation test[J]. Avian Pathology. 6:259~267.
Ianconeseu M, Aharonovici A.1978. Persistant viraemia in chickens subsequent to in ovo inoculation of reticuloendotheliosis virus[J]. Avian Pathology. 7:237~247.
Isfort R, Jones D, Kost R., et al. 1992. Retrovirus insertion into herpesvirus in vitro and in vivo[J]. Proc Natl Acad Sci. 89: 991~995.
Jackson CAW, Dunn SE, Smith DI, et al. 1977. Proventriculitis“Nakanuke”and reticuloendoheliosis in chickens following vaccination with herpesvirus of turkeys (HVT)[J]. Aust. Vet . J. 53: 457~458.
Jespersen T, Duch M, Carrasco ML, et al. 1999. Expression of heterologous genes from an IRES translational cassette in replication competent murine leukemia virus vectors[J]. Gene. 239: 227~235.
Jones D, Isfort R, Witter R, et al. 1993. Retroviral insertions into a herpesvirus are clustered at the junctions of the short repeat and short unique sequences[J]. Proc Natl Acad Sci. 90: 3855~3859.
Jones D, Brunovskis P, Witter R, et al. 1996. Retroviral insertional activation in a herpesvirus:Transcriptional activation of US genes by an integrated long terminal repeat in a Marek’s disease virus clone[J]. J Virol. 70: 2460~2467.
Kang CY, Howard MT. 1974. Reticuloendotheliosis virus nucleic acid sequences in cellular DNA [J]. J Virol. 14(5): 1179~1188.
Koyama H, Sasaki T, Ohwada Y, et al. 1980. The relationship between feathering abnormalities (“Nakanuke”) and tumour production in chickens inoculated with reticuloendotheliosis virus[J]. Avian Pathology. 9:331~340.
Larose RN, Sevoian M. 1965. Avian lymphomatosis. IX. Mortality and serological response of chickens of various ages to graded doses of T strain[J]. Avian Disease. 9: 604~610.
Li J, Calnek BW, Schat KA, et al. 1983. Pathogenesis of retieuloendotheliosis virus infection in ducks[J]. Avian Disease. 27:1090~1105.
Liu Q, Zhao J, Su J, et al. 2009. Full genome sequences of two reticuloendotheliosis viruses contaminating commercial Vaccines[J]. Avian Dis. 53: 341~346.
Logg CR, Logg A, Tai CK, et al. 2001. Genomic stability of murine leukemia viruses containing insertions at the Env-3V untranslated region boundary[J]. J Virol. 75: 6989~6998.
Masami T, Kiyoyasu I, Hideki N, et al. 1996. Detection of contamination of vaccines with the reticuloendotheliosis virus by reverse transcriptase polymerase chain reaction (RT-PCR) [J]. Virus Res. 40: 113~121.
Mays J, Silva R, Lee L, et al. 2010. Characterization of reticuloendotheliosis virus isolates obtained from broiler breeders, turkeys, and prairie chickens located in various geographical regions in the United States. Avian Pathol. 39: 383~389.
Mcdougall JS, Shilleto RFW, Biggs PM. 1980. Experimental infection and vertical transmission of reticuloendotheliosis virus in the turkey. Avian Pathologyg[J]. 9:445~454.
Moore KM, Davis JR, Sato T, et al. 2000. Reticuloendotheliosis virus (REV) long terminal repeats incorporated in the genomes of commercial fowl poxvirus vaccines and pigeon poxviruses without indication of the presence of infectious REV[J]. Avian Dis. 44: 827~841.
Mori Y, Nagamine K, Tomita N, et al. 2001. Detection of loop-mediated isothermal amplification reaction by turbidity derived from magnesium pyrophosphate formation. Biochem[J]. Biophys. Res. Commun. 289: 150~154.
Motha MXJ, Egerton JR, Sweeney AW. 1984. Some evidence of mechanical transmission of reticuloendotheliosis virus by mosquitos[J]. Avian Disease. 28:858~867.
Nicholas RAJ, Thornton DH. 1987. An enzymelinked immunosorbent assay for the detection of antibodies to avian reticuloendotheliosis virus using whole cell antigen[J]. Research in Veterinary Science. 43:403~404.
Notomi T, Okayama H, Masubuchi H, et al. 2000. Loop-mediated isothermal amplification of DNA[J]. Nucleic Acids Res. 28: 63.
Paul PS, Johnson KH, Pomeroy KA, et al. 1977. Experimental transmission of reticuloendothelios is in turkeys with the cell culture propagated reiculoendotheliosis viruses of turkey origin[J]. Joumal of the National Cancer Institute. 58:1819~1824.
Pedersen FS, Duch M. 2002. Replication competent retroviral vectors[J]. Bioforum Int. 6:100~104.
Peterson DA, Levine AS. 1971. Avian reticuloendotheliosis virus(strainT) IV Infectivity and transmissibility in day-old cockerels[J]. Avian Disease. 15: 874~883.
Racaniello V. R., Baltimore D. 1981. Cloned polivirus comlementary DNA is infectious in mammalian cells. Science. 214: 916–919.
Reimann I, Werner O. 1996. Use of the polymerase chain reaction for the detection of reticuloendotheliosis virus in Marek’s disease vaccines and chicken tissues[J]. J Vet Med. 43: 75~84.
Robinson FR, Twiehaus MJ.1974. Isolation of the avian reticuloendotheliosis virus (strainT) [J]. Avian Disease. 18:278~288.
Seong HW, Kim SJ, Kim JH, et al. 1996. Outbreaks of reticuloendotheliosis in Korea [J]. RDA Journal of Agrieultural Science Veterinary. 38:707~715.
Singh P, Kim TJ, Tripathy DN. 2000. Re-emerging fowPox: evaluation of isolates from vaccinated flocks[J]. Avian Pathology. 29: 449~455.
Singh P, Schnitzlein WM, Tripathy DN. 2003. Reticuloendotheliosis virus sequences within the genomes of field strains of fowlpox virus display variability[J]. J Virol. 77(10): 5855~5862.
Solly SK, Trajcevski S, Frisen C, et al. 2003. Replicative retroviral vectors for cancer gene therapy[J]. Cancer Gene Ther. 10: 30~39.
Sun AJ, Xu XY. 2009. Petherbridge L, et al. Functional evaluation of the role of reticuloendotheliosis virus long terminal repeat (LTR) integrated into the genome of a field strain of Marek's disease virus[J]. Virology. 11: 17~23.
Tadese T, Fitzgerald S, Reed W M. 2008. Detection and differentiation of reemerging fowlpox virus (FWPV) strains carrying integrated reticuloendotheliosis virus (FWPV-REV) by real-time PCR[J]. Vet Microbiol. 127: 39~49.
Taniguchi T., Palmieri M., Weissmann C. 1978. QB DNA-containing hybrid plasmids giving rise to QB phage formation in the bacterial host. Nature. 274 (5668): 223-228.
Temin HM, Kassner VK. 1974. Replication of reticuloendotheliosis viruses in cell culture: acute infection[J]. Journal of Virology. 13: 291~297.
Theilen GH, Zeigel RF, Twiehaus MJ.1966. Biological studies with REvirus (strain T) that induces reticuloendotheliosis in turkeys, chickens, and Japanese quail[J]. Journal of the National Cancer Institute. 37:.731~743.
Trager W. 1959. A new virus of ducks interfering with development of malaria parasite (Plasmodium lophurae) [J]. Proceedings of the Society for Experimental Biology and Medicine. 101: 578~582.
Witter RL, Fadly AM. 2003. Reticuloendotheliosis[C]. Diseases of poultry, 11th ed. State University Press. 517~535.
Witter RL, Johnson DC. 1985. Epidemiology of reticuloendotheliosis virus in broiler breeder flocks[J]. Avian Dis. 29:1140~1154.
Witter RL, Purchase HG, Burgoyne GH.1970. Peripheral nerve lesions similar to those of Marek,s disease in chickens inoculated with reticuloen-dotheliosis virus[J]. Journal of the National Cancer Institute. 45: 567~577.
Witter RL. 1997. Reticuloendothelisis. B W Calnek. Diseases of Poultry (10th) [M]. Iowa State University Press. 467~484.
Wong TC, Lai MMC.1981. Avian reticuloendotheliosis virus contains a new class of oneogene of turkey origin[J].Virology. 111:289~293.
Yuasa N, Yoshida I, Taniguchi T. 1976. Isolation of a reticuloendotheliosis virus from chickens inoculated with Marek’s disease vaccine[J]. Natl. Inst . Anim.Health. Q. 16: 141~151.
Zavala G, Cheng S, Barbosa T. 2006. Enzootic Reticuloendotheliosis in the Endangered Attwater’s and Greater Prairie Chickens Guillermo. Avian Dis. 50: 520-525.
Zhang Z,Cui ZZ. 2005. Isolation of reeombinant field strains of Marek’s disease virus integrated with retieuloendotheliosis virus genome fragments[J]. Science in China Sciences. 48: 81~88.
崔治中. 2006.禽反转录病毒与DNA病毒间的基因重组及其流行病学意义[J].病毒学报. 22:150~154.
崔治中. 2008.鸭群中REV感染的流行病学调查[J].微生物学报. 48(4): 514~519.
崔治中,孟珊珊,姜世金等. 2006.我国白羽肉用型鸡群中CAV、REV和REOV感染状况的血清学调查[J].畜牧兽医学报. 37(2): 152~157.
崔治中,孙淑红,王辉等. 2009.鸡群中马立克氏病毒、J-亚群白血病毒和网状内皮增生病毒的多重感染[C].中英家禽肿瘤病理及肿瘤病毒研讨会.山东农业大学.
崔治中,孙怀昌,朱承如. 1987.禽白血病及网状内皮增生病感染情况的调查[J].中国畜禽传染病. l:37~38.
丁家波,崔治中,于立娟. 2004.含有禽网状内皮组织增生病病毒基因组片段的天然重组禽痘病毒的研究[J].微生物学报. 44:588~592.
杜岩,崔治中. 2001.禽网状内皮组织增生病病毒感染引起的鸡群免疫抑制[J].中国兽医学报. 21(2):122~124.
段宏安, Ibrahim Diallo , Peter Spradbrow ,等. 1999.应用聚合酶链式反应检测禽痘病毒田间分离株及疫苗株中网状内皮组织增殖病毒LTR片段, env及rel基因[J].中国兽医杂志. 25 (8): 9~10.
何宏虎,陈溥言,蔡宝祥. 1988.禽网状内皮组织增生病病毒的分离鉴定[J].中国畜禽传染病. 1~2.
何勇群,张中直,杨汉春.1998.北京地区鸡群网状内皮组织增殖病感染的血清学调查研究[J].畜牧兽医学报. 29:71~77.
胡北侠,黄艳艳,路希山等. 2009.肉种禽网状内皮增生症、鸡传染性贫血和禽白血病血清学调查[J].广东畜牧兽医科技. 34(1): 25~27.
吉荣,崔治中,丁家波. 2003.网状内皮组织增生病病毒囊膜糖蛋白gp90的原核表达[J]. 中国兽医学报. 23(1): 28~30.
吉荣,崔治中,王锡乐等. 2005.分子克隆化禽网状内皮组织增生症病毒传染性及其前病毒全基因组序列研究[J].病毒学报. 21:448~455.
吉荣,刘岳龙,秦爱建. 2001.免疫抑制鸡群鸡传染性贫血病毒和网状内皮增生症病毒共感染检测[J].中国兽药杂志. 35(4):1~3.
姜世金,孟珊珊,崔治中等. 2005.我国自然发病鸡群中MDV、REV和CAV共感染的检测[J].中国病毒学. 20:164~167.
金文杰,崔治中,刘岳龙等.2001.传染性法氏囊病病料中MDV、CAV、REV的共感染检测[J].中国兽医学报. 1:6~9.
匡宝晓,徐福南.1997.网状内皮组织增殖病病毒感染鸡红细胞免疫功能的动态变化[J].中国兽医科技. 27:7~9.
李广兴,刘忠贵,郎跃深. 2000. SPF雏鸡感染网状内皮组织增殖病病毒后白细胞介素2和干扰素诱生活性的变化[J].中国兽医科技. 30:3~5.
李凯,高宏雷,高立等. 2010.禽网状内皮组织增生病病毒TaqMan探针荧光定量PCR检测方法的建立及应用[J].中国兽医科学. 11:1137-1140.
李延鹏,崔治中,姜世金等. 2008.CAV与REV共感染SPF鸡对疫苗免疫反应的抑制作用[J].中国兽医学报. 28(11): 1243~1246.
孟珊珊,崔治中,孙淑红. 2006. REV和ALV-J共感染鸡病毒血症及抗体反应的相互影响[J].中国兽医学报. 26(4): 363-366.
秦立廷,高玉龙,潘伟等. 2010.我国部分地区蛋鸡群ALV~J及与REV、MDV、CAV混合感染检测[J].中国预防兽医学报. 32(2): 90~93.
荣骏弓,王笑梅,幸桂香等. 1997.禽网状内皮组织增殖病琼扩免疫扩散抗原的研制及应用[J].中国畜禽传染病. 5: 29~33.
孙淑红. 2007.博士论文:禽网状内皮组织增生症病毒和J-亚群白血病病毒的致病性及其疫苗研究.山东农业大学. 15-30.
王玉. 2005.博士论文:网状内皮组织增生症病毒分子流行病学调查及其在宿主免疫选择压下的分子变异.山东农业大学. 88-100.
王玉,崔志中,姜世金. 2006.网状内皮组织增生症病毒中国分离株HA9901全基因组核苷酸序列的测定与分析[J].中国科学C辑. 49(2): 149~157.
杨汉春,高云. 1997.应用改良ELISA检测禽网状内皮增殖病抗体血清[J].中国兽医杂志.23:3~5.
殷震,刘景华. 1997.动物病毒学(第二版).科学出版社. 329~331.
于立娟,崔治中. 2006.禽痘疫苗病毒中网状内皮组织增生病病毒5’LTR整合位点序列分析[J].微生物学报. 46:660~662.
郁晓岚,徐福南,蔡宝祥等. 1989.禽网状内皮增殖病组织病理学和超微结构的动态变化[J].南京农业大学学报. 12:91~96.
张志,王锡乐,庄国庆等. 2005.用单抗介导的免疫荧光试验检测组织切片中的禽网状内皮组织增生病病毒[J].中国兽医学报. 25(2) :122~124.
张志,崔治中,姜世金. 2004.从J亚群禽白血病肿瘤中检测出网状内皮组织增生病病毒[J].中国兽医学报. 1:10~23.
瞿晓兰,王宏俊,陈小玲. 2006.禽网状内皮组织增生病毒囊膜蛋白基因的克隆及其原核表达[J ].华北农学报. 21(增刊):154 ~157.
赵丽青,吴延功,王志亮. 2006.禽网状内皮组织增殖病的特异性荧光PCR检测方法的研究[C].湖南长沙:中国畜牧兽医学会禽病学分会第13次学术研讨会论文集. 10: 442~444.
Alien PT, Mullins JA, Harris CL, et al. 1979. Replication of reticuloendotheliosis virus in mammalian cells(abstract) [C]. Amer Soc Microbiol Annual Meeting. 100:256~256.
Aly MM, Fadly AM, Witter RL. 1992. Effects of serotype 2 Marek,s disease virus on development of viremia, antibody, and lymphoma induced by reticuloendotheliosis virus.4thInternational Symposium on Marek,s disease,19th world,s Poultry Congress, world,s Poultry Science Assn[C]. Amsterdam. l:272~276.
Aly MM, Hassan MK, Elzahr AA, et al. 1998. Serological survey on retieuloendotheliosis virus infection in comercial chicken and turkey flocks in EgyPt. Proceedings of the 5th Science Conference [C]. Egypt Vet Poultry Association. 51~68.
Aly MM, Smith EJ, Fadly AM. 1993. Detection of reticuloendotheliosis virus infection using the polymerase chain reaction[J]. Avian Pathology. 22:543~554.
Bachrach E, Duch M, Pelegrin M, et al. 2003. In vivo infection of mice by replication-competent MLV based retroviral vectors[J]. Methods Mol. Med. 76:343~352.
Bachrach E, Pelegrin M, Piechaczyk M, et al. 2002. Efficient gene transfer into spleen cells of newborn mice by a replication-competent retroviral vector. Virology. 293:328~334.
Bagust TJ, Grimes TM, Dennett DP. 1979. Infection studies on a reticuloendotheliosis virus contaminant of a commercial Marek’s disease vaccine [J]. Aust. Vet. J. 55: 153~157.
Bagust TJ, Grimes TM, Ratnamohan N. 1981. Experimental infection of chickens with an Australian strain of reticuloendotheliosis virus 3 Persistent infection and transmission by the adult hen[J]. Avian Patholoy. 10:375~385.
Barbosa T, Zavala G, Cheng S, et al. 2007. Full genome sequence and some biological properties of reticuloendotheliosis virus strain APC-566 isolated from endangered Attwater's prairie chickens[J]. Virus Res. 124, 68~77.
Barth CF, Humphries EH. 1988. Expression of v-relinduces mature B-cell lines that reflect the diversity of avian immunoglobulin heavy and light-chain rearrangements[J]. Molecular and Cellular Biology. 8:5358~5368.
Baxter-Gabbard KL, Campbell WF, Padgett F. 1971. Avian reticuloendotheliosis virus (strainT).Ⅱ.Biochemical and biophysical properties[J]. Avian Disease 15:850~862.
Berglund P, Pleeton MN, Smerdou C, et al. 1999. Immunization with recombinant Semliki Forest virus induces protection against influenza challenge in mice[J]. Vaccine.17 (5): 497~507.
Berglund P, Smerdou C, Fleeton MN, et al. 1998. Enhancing immune responses using suicidal DNA vaccines[J]. Nat Biotechnol.16 (6): 562~565.
Bohls RL, Linares JA, Gross SL, et al. 2006. Phylogenetic analyses indicate little variation among reticuloendotheliosis viruses infecting avian species, including the endangered Attwater’s prairie chicken[J]. Virus Res. 119:187~194.
Bose HR, Levine AS.1967. Replication of the reticuloendotheliosis virus (strainT) in chicken embryo cell culture[J]. Journal of Virology. l:1117~1121.
Breitman ML, Lai MMC, Vogt PK.1980. Attenuation of avian reticuloendotheliosis virus: Loss of the defective transforming component during serial passage of oncogenic virus in fibroblasts[J]. Virology. 101:304~306.
Celine G, Howard MT. 1986. Nondefective spleen necrosis virus-derived vectors define the upper size limit for packaging reticuloendotheliosis viruses[J]. Proc Nati Acad Sci. 83: 9211-9215.
Chang Z, Pan J, Logg C, et al. 2001. A replication-competent feline leukemia virus, subgroup A (FeLV-A), tagged with green fluorescent protein reporter exhibits in vitro biological properties similar to those of the parental FeLV-A[J]. J Virol. 75: 8837~ 8841.
Chen IS, Malk TW, O’Pear JJ, et al.1981. Characterization of retieuloendotheliosis virus strainT/DNA and isolation of a novel variant of reticuloendotheliosis virus strain T by molecular cloning[J]. J Virol. 40:800~811.
Chen PY, Cui ZZ, Lee LF, et al. 1987. Serologic difference among nondefective reticuloendotheliosis viruses[J]. Arch. Virol. 93:233~246.
Chen HT, Zhang J, Sun DH, et al. 2008. Rapid detection of porcine circovirus type 2 by loop-mediated isothermal amplification[J]. J Virol Methods. 149: 264~268.
Cheng WH, Huang YP, Wang CH. 2006. Serological and virological surveys of reticuloendotheliosis in chickens in Taiwan [J]. J Vet Med Sci. 68(12):1315~1320.
Coffin JM.1982. Structure of the retroviral genome. RNA tumor viruses. Molecular Biology of Tumor Viruses[J]. 2 Cold Spring Harbor Laboratory. Cold Spring Harbor. NY: 261~368.
Cohen M, Rein A, Stephens R.M, et al. 1981. Baboon endogenous virus genome.Ⅲ.non- Molecular cloning and structural characterization of defective viral genomes from the DNA of ababoon cell strain[J]. Proc.National Academy Science. 78:5207~5211.
Cook MK. 1969. Cultivation of filterable agent associated with Marek,s disease[J]. Joumal of the National Cancer Institute. 43:203~212.
Crittenden LB, Fadly AM, Smith EJ.1982. Effect of endogenous leukosis virus genes on response to infection with avian leucosis and reticuloendotheliosis virus[J]. Avian Disease. 26:279~294.
Cui ZZ, Sun SH, Zhang Z, et al. 2009a. Simultaneous endemic infections with subgroup J avian leukosis virus and reticuloendotheliosis virus in commercial and local breeds of chickens[J]. Avian Pathol. 38(6):443~448.
Cui ZZ, Zhuang G, Xu X, et al. 2009b. Molecular and biological characterization of a Marek’s disease virus field strain with reticuloendotheliosis virus LTR insert[J]. Virus gene. 437~445. Cui ZZ, Lee L Fetc. 1988. Monoclonal antibody mediated enzyme linked immunosorbent assay for detection of reticuloendotheliosis virus [J]. Avian Dis. 32: 32~40.
Davidson I, Borovskays A, Peri S, et al. 1995. Use of the polymerase chain reaction for the diagnosis of natural infection chickens and turkeys with Marek’s disease virus and reticuloendotheliosis virus[J]. Avian Pathology. 24: 69~94.
Davidson I, Borenshtain R. 2004. In vivo events of retroviral long terminal repeat integration into Marek's disease virus in commercial poultry: detection of chimeric molecules as a marker[J]. Avian Dis. 45: 102~121.
Davidson I, Yang H, Witter RL, et al. 1995. The immunodominant proteins of reticuloendotheliosis virus [J]. Vet Microbiol. 49: 273~284.
Diallo IS, Mackenzie MA, Spradbrow PB, et al. 1998. Field isolates of fowlpox viruscontaminated with reticuloendotheliosis virus[J]. Avian Pathol. 27: 60~66.
Dubensky T. W., Driver D. A., Polo J. M. 1996. Sindbis virus DNA-based expression vectors: utility for in vitro and in vivo gene transfer. J. Virol.70 (1): 508-519.
Fadly AM, Witter RL, Smith EJ, et al. 1996. An outbreak of lymphomas in commercial broiler breeder chickens vaccinated with a fowlpox vaccine contaminated with reticuloendotheliosis virus[J]. Avian Pathol. 25: 35~47.
Fadly AM, Witter RL. 1997.Comparative evaluation of in vitro and in vivo assays for the detection of reticuloendotheliosis virus as a contaminant in a live virus vaccine of poultry[J]. Avian Dis. 41: 695~701.
Fadly AM, Garcia MC. 2006. Detection of reticuloendotheliosis virus in live virus vaccines of poultry[J]. Dev Biol (Basel) 126: 301~305.
Fleeton M N, Liljestrom P, Sheahan B J, et al. Recombinant Semliki Forest virus particles expressing louping ill virus antigens induce a better protective response than plasmid-based DNA vaccines or an inactivated whole particle vaccine. J Gen Virol., 2000, 81 (3): 749-758.
Fritsch RB, Kang CY, Wan KMM, et al. 1977. Transformation of chick embryo fibroblasts by reticuloendotheliosis virus[J]. Virology. 83:313~321.
García MN, Narang N, Reed WM, et al. 2003. Molecular characterization of reticuloendotheliosis virus insertions in the genome of field and vaccine strains of fowl poxvirus[J]. Avian Dis. 47: 343~354.
Ge J Y, Deng G H, Wen Z Y, et al. 2007. Newcastle disease virus-based live attenuated vaccine completely protects chickens and mice from lethal challenge of homologous and heterologous H5N1 avian influenza viruses. J. Virol. 81 (1): 150~158.
Guillermo Zavala, Sunny Cheng, Taylor Barbosa, et al. 2006. Enzootic Reticuloendotheliosis in the Endangered Attwater’s and Greater Prairie Chickens[J]. Avian Dis. 50:520~525.
Hauck R, Prusas C, Hafez HM, et al. 2009. Serologic response against fowl poxvirus and reticuloendotheliosis virus after experimental and natural infections of chickens with fowl poxvirus[J]. Avian Dis. 53(2):205~210.
Hertig C, Coupar BE, Gould AR, et al. 1997. Field and vaccine strains of fowlpox virus carry integrated sequences from the avian retrovirus, reticuloendotheliosis virus[J]. Virology. 235: 367~376.
Hoelzer JD, Franklin RB, Bose JHR.1979. Transformation by reticuloendotheliosis virus: development of a focus assay and isolation of a non-transforming virus. Virology[J]. 93:20~30.
Hu Sylvia SF, Lai Michael MC, Wong TC. 1981. Avian reticuloendotheliosis virus : characterization of genome structure by heteroduplex mapping [J]. J Virol Methods. 37 (3): 899~907.
Humphries EH, Zhang G.1992. V-rel and C-rel modulate the expression of both bursal and non-bursal antigens on avian B-cell lymphomas. Current Topics in Microbiology and Immunology[J]. 182: 475~483.
Ianconescu M.1977. Reticuloendotheliosis antigen for the agar gel precipitation test[J]. Avian Pathology. 6:259~267.
Ianconeseu M, Aharonovici A.1978. Persistant viraemia in chickens subsequent to in ovo inoculation of reticuloendotheliosis virus[J]. Avian Pathology. 7:237~247.
Isfort R, Jones D, Kost R., et al. 1992. Retrovirus insertion into herpesvirus in vitro and in vivo[J]. Proc Natl Acad Sci. 89: 991~995.
Jackson CAW, Dunn SE, Smith DI, et al. 1977. Proventriculitis“Nakanuke”and reticuloendoheliosis in chickens following vaccination with herpesvirus of turkeys (HVT)[J]. Aust. Vet . J. 53: 457~458.
Jespersen T, Duch M, Carrasco ML, et al. 1999. Expression of heterologous genes from an IRES translational cassette in replication competent murine leukemia virus vectors[J]. Gene. 239: 227~235.
Jones D, Isfort R, Witter R, et al. 1993. Retroviral insertions into a herpesvirus are clustered at the junctions of the short repeat and short unique sequences[J]. Proc Natl Acad Sci. 90: 3855~3859.
Jones D, Brunovskis P, Witter R, et al. 1996. Retroviral insertional activation in a herpesvirus:Transcriptional activation of US genes by an integrated long terminal repeat in a Marek’s disease virus clone[J]. J Virol. 70: 2460~2467.
Kang CY, Howard MT. 1974. Reticuloendotheliosis virus nucleic acid sequences in cellular DNA [J]. J Virol. 14(5): 1179~1188.
Koyama H, Sasaki T, Ohwada Y, et al. 1980. The relationship between feathering abnormalities (“Nakanuke”) and tumour production in chickens inoculated with reticuloendotheliosis virus[J]. Avian Pathology. 9:331~340.
Larose RN, Sevoian M. 1965. Avian lymphomatosis. IX. Mortality and serological response of chickens of various ages to graded doses of T strain[J]. Avian Disease. 9: 604~610.
Li J, Calnek BW, Schat KA, et al. 1983. Pathogenesis of retieuloendotheliosis virus infection in ducks[J]. Avian Disease. 27:1090~1105.
Liu Q, Zhao J, Su J, et al. 2009. Full genome sequences of two reticuloendotheliosis viruses contaminating commercial Vaccines[J]. Avian Dis. 53: 341~346.
Logg CR, Logg A, Tai CK, et al. 2001. Genomic stability of murine leukemia viruses containing insertions at the Env-3V untranslated region boundary[J]. J Virol. 75: 6989~6998.
Masami T, Kiyoyasu I, Hideki N, et al. 1996. Detection of contamination of vaccines with the reticuloendotheliosis virus by reverse transcriptase polymerase chain reaction (RT-PCR) [J]. Virus Res. 40: 113~121.
Mays J, Silva R, Lee L, et al. 2010. Characterization of reticuloendotheliosis virus isolates obtained from broiler breeders, turkeys, and prairie chickens located in various geographical regions in the United States. Avian Pathol. 39: 383~389.
Mcdougall JS, Shilleto RFW, Biggs PM. 1980. Experimental infection and vertical transmission of reticuloendotheliosis virus in the turkey. Avian Pathologyg[J]. 9:445~454.
Moore KM, Davis JR, Sato T, et al. 2000. Reticuloendotheliosis virus (REV) long terminal repeats incorporated in the genomes of commercial fowl poxvirus vaccines and pigeon poxviruses without indication of the presence of infectious REV[J]. Avian Dis. 44: 827~841.
Mori Y, Nagamine K, Tomita N, et al. 2001. Detection of loop-mediated isothermal amplification reaction by turbidity derived from magnesium pyrophosphate formation. Biochem[J]. Biophys. Res. Commun. 289: 150~154.
Motha MXJ, Egerton JR, Sweeney AW. 1984. Some evidence of mechanical transmission of reticuloendotheliosis virus by mosquitos[J]. Avian Disease. 28:858~867.
Nicholas RAJ, Thornton DH. 1987. An enzymelinked immunosorbent assay for the detection of antibodies to avian reticuloendotheliosis virus using whole cell antigen[J]. Research in Veterinary Science. 43:403~404.
Notomi T, Okayama H, Masubuchi H, et al. 2000. Loop-mediated isothermal amplification of DNA[J]. Nucleic Acids Res. 28: 63.
Paul PS, Johnson KH, Pomeroy KA, et al. 1977. Experimental transmission of reticuloendothelios is in turkeys with the cell culture propagated reiculoendotheliosis viruses of turkey origin[J]. Joumal of the National Cancer Institute. 58:1819~1824.
Pedersen FS, Duch M. 2002. Replication competent retroviral vectors[J]. Bioforum Int. 6:100~104.
Peterson DA, Levine AS. 1971. Avian reticuloendotheliosis virus(strainT) IV Infectivity and transmissibility in day-old cockerels[J]. Avian Disease. 15: 874~883.
Racaniello V. R., Baltimore D. 1981. Cloned polivirus comlementary DNA is infectious in mammalian cells. Science. 214: 916–919.
Reimann I, Werner O. 1996. Use of the polymerase chain reaction for the detection of reticuloendotheliosis virus in Marek’s disease vaccines and chicken tissues[J]. J Vet Med. 43: 75~84.
Robinson FR, Twiehaus MJ.1974. Isolation of the avian reticuloendotheliosis virus (strainT) [J]. Avian Disease. 18:278~288.
Seong HW, Kim SJ, Kim JH, et al. 1996. Outbreaks of reticuloendotheliosis in Korea [J]. RDA Journal of Agrieultural Science Veterinary. 38:707~715.
Singh P, Kim TJ, Tripathy DN. 2000. Re-emerging fowPox: evaluation of isolates from vaccinated flocks[J]. Avian Pathology. 29: 449~455.
Singh P, Schnitzlein WM, Tripathy DN. 2003. Reticuloendotheliosis virus sequences within the genomes of field strains of fowlpox virus display variability[J]. J Virol. 77(10): 5855~5862.
Solly SK, Trajcevski S, Frisen C, et al. 2003. Replicative retroviral vectors for cancer gene therapy[J]. Cancer Gene Ther. 10: 30~39.
Sun AJ, Xu XY. 2009. Petherbridge L, et al. Functional evaluation of the role of reticuloendotheliosis virus long terminal repeat (LTR) integrated into the genome of a field strain of Marek's disease virus[J]. Virology. 11: 17~23.
Tadese T, Fitzgerald S, Reed W M. 2008. Detection and differentiation of reemerging fowlpox virus (FWPV) strains carrying integrated reticuloendotheliosis virus (FWPV-REV) by real-time PCR[J]. Vet Microbiol. 127: 39~49.
Taniguchi T., Palmieri M., Weissmann C. 1978. QB DNA-containing hybrid plasmids giving rise to QB phage formation in the bacterial host. Nature. 274 (5668): 223-228.
Temin HM, Kassner VK. 1974. Replication of reticuloendotheliosis viruses in cell culture: acute infection[J]. Journal of Virology. 13: 291~297.
Theilen GH, Zeigel RF, Twiehaus MJ.1966. Biological studies with REvirus (strain T) that induces reticuloendotheliosis in turkeys, chickens, and Japanese quail[J]. Journal of the National Cancer Institute. 37:.731~743.
Trager W. 1959. A new virus of ducks interfering with development of malaria parasite (Plasmodium lophurae) [J]. Proceedings of the Society for Experimental Biology and Medicine. 101: 578~582.
Witter RL, Fadly AM. 2003. Reticuloendotheliosis[C]. Diseases of poultry, 11th ed. State University Press. 517~535.
Witter RL, Johnson DC. 1985. Epidemiology of reticuloendotheliosis virus in broiler breeder flocks[J]. Avian Dis. 29:1140~1154.
Witter RL, Purchase HG, Burgoyne GH.1970. Peripheral nerve lesions similar to those of Marek,s disease in chickens inoculated with reticuloen-dotheliosis virus[J]. Journal of the National Cancer Institute. 45: 567~577.
Witter RL. 1997. Reticuloendothelisis. B W Calnek. Diseases of Poultry (10th) [M]. Iowa State University Press. 467~484.
Wong TC, Lai MMC.1981. Avian reticuloendotheliosis virus contains a new class of oneogene of turkey origin[J].Virology. 111:289~293.
Yuasa N, Yoshida I, Taniguchi T. 1976. Isolation of a reticuloendotheliosis virus from chickens inoculated with Marek’s disease vaccine[J]. Natl. Inst . Anim.Health. Q. 16: 141~151.
Zavala G, Cheng S, Barbosa T. 2006. Enzootic Reticuloendotheliosis in the Endangered Attwater’s and Greater Prairie Chickens Guillermo. Avian Dis. 50: 520-525.
Zhang Z,Cui ZZ. 2005. Isolation of reeombinant field strains of Marek’s disease virus integrated with retieuloendotheliosis virus genome fragments[J]. Science in China Sciences. 48: 81~88.