蛋鸡J亚群禽白血病病毒的分离和部分序列测定
|
摘要
J亚群禽白血病(Avian leucosis virus subgroup J,ALV-J)是一种病毒引起的禽类肿瘤性疾病，又称作骨髓细胞瘤病(Myelocytomatosis)。J亚群禽白血病自发现以来，仅见于肉鸡发病的报道，该病给世界上许多国家的养鸡业带来了巨大的经济损失。2002年中国农业大学徐镔蕊发现了蛋鸡J亚群禽白血病。自然发病的蛋鸡表现为贫血、消瘦、产蛋下降、死亡率达10％，尸体剖检可见卵巢、输卵管发育不良，给蛋鸡业造成很大的经济损失，经过病理学、免疫组化学以及PCR方法检测ALV-Jgp85抗原，并对该病毒亚群特异性的序列进行测定，在国内外首次发现并报道了蛋鸡J亚群禽白血病的自然病例。
本研究采用临床发病蛋鸡的病料进行了病毒分离。病料(肝、脾、肾)过滤除菌后接种于鸡胚成纤维细胞(CEF)。培养7天后，用ALV-Jgp85特异性单抗进行间接荧光抗体法(IFA)检测接种的CEF，结果发现CEF的胞浆内有明显的特异性荧光，细胞核未被染色，用免疫组化的方法检测到ALV-J囊膜特异性蛋白gp85的存在，为了进一步对病毒核酸序列进行分析，我们提取接种病料的CEF的总RNA，做反转录聚合酶链式反应(RT-PCR)，在pol基因的3′端和gp85编码基冈的内部设计了一对引物，扩增得到长度为545bp的ALV-J特异性片段，与预期片段大小相符，经TaKaRa公司序列测定后，与ALV-J原型株HPRS-103的序列进行了比较，发现其核苷酸同源性为97.4％，所编码氨基酸的同源性为96.1％。
本研究结果显示我们从蛋鸡中首次成功分离到J亚群禽白血病病毒，说明本实验所研究蛋鸡的骨髓细胞瘤病是由ALV-J引起。对蛋鸡ALV-J的分离，一方面为之前人们所认识的引起蛋鸡产蛋下降的疾病的病原增添了新的成员，同时也将为更好的防治和根除ALV-J、减少养鸡业的经济损失具有重要的理论和实际意义。
Avian leucosis virus subgroup J in chickens ( ALV-J) is a tumor disease induced by virus, namely Myelocytomatosis. Since ALV-J was discovered, it has just been reported in meat-type chickens, resulting in considerable losses of chickens industry all over the world. In 2002, Xu Binrui, professor of China Agricultural University, discovered ALV-J in layer chickens. Naturally infected layer flocks showed the symptom of tabes and serious anemia, decrease of egg production, and the morality was as high as 10%. Maldevelopment was present in testes, ovaries, and oviducts. It caused considerable losses of layer chickens industry. Specific protein gp85 of ALV-J was detected through pathology, immunohistochemistry and PCR method. The layer chickens natural infected with ALV-J were first dicovered and reported in the world.
In the study isolation of virus was performed using the samples of layer chickens naturally infected with ALV-J. The tissue samples (hearts, livers, spleens etc) of illed chickens were filtrated and inoculated into chicken embryo fibroblast (CEF). The CEFs were detected using the indirect fluorescence assay (I FA) with specific monoclonal antibody against ALV-J gp85 at 7 day after inoculation. Specific fluorescence was detected in the cytoplasm of CEFs and the caryon was unstained. Specific envelope protein gp85 was detected by using immunohistochemistry. For further detection RT-PCR was performed on the CEF inoculated with the samples. A 545bp specific product was amplified using primer set H5 and H7 that was designed in 3' region of pol gene and 5' region of the gp85 encoding gene respectively. The product was as same size as expected and was sequenced by TaKaRa company. The sequence data showed that the identity of the nucleotide sequence was 97.4% and that of the amino acid sequence was 96.1% compared with that of the prototypical strain of ALV-J HPRS-103. The results of RT-PCR illuminated that the gp85 gene of ALV-J isolated from egg-type chickens have some variation compared with that of prototype HPRS-103 strain.
The results of this study demonstrated that ALV-J was successfully isolated from naturally infected egg-type chickens for the first time. The isolation of ALV-J in layer chickens will develope a new member for previously known diseases that affected the productivity of chickens, meanwhile, it will also provide theory and practice basis for better preventing, controlling the occurrence of ALV-J and decreasing the economic losses.
本研究采用临床发病蛋鸡的病料进行了病毒分离。病料(肝、脾、肾)过滤除菌后接种于鸡胚成纤维细胞(CEF)。培养7天后，用ALV-Jgp85特异性单抗进行间接荧光抗体法(IFA)检测接种的CEF，结果发现CEF的胞浆内有明显的特异性荧光，细胞核未被染色，用免疫组化的方法检测到ALV-J囊膜特异性蛋白gp85的存在，为了进一步对病毒核酸序列进行分析，我们提取接种病料的CEF的总RNA，做反转录聚合酶链式反应(RT-PCR)，在pol基因的3′端和gp85编码基冈的内部设计了一对引物，扩增得到长度为545bp的ALV-J特异性片段，与预期片段大小相符，经TaKaRa公司序列测定后，与ALV-J原型株HPRS-103的序列进行了比较，发现其核苷酸同源性为97.4％，所编码氨基酸的同源性为96.1％。
本研究结果显示我们从蛋鸡中首次成功分离到J亚群禽白血病病毒，说明本实验所研究蛋鸡的骨髓细胞瘤病是由ALV-J引起。对蛋鸡ALV-J的分离，一方面为之前人们所认识的引起蛋鸡产蛋下降的疾病的病原增添了新的成员，同时也将为更好的防治和根除ALV-J、减少养鸡业的经济损失具有重要的理论和实际意义。
Avian leucosis virus subgroup J in chickens ( ALV-J) is a tumor disease induced by virus, namely Myelocytomatosis. Since ALV-J was discovered, it has just been reported in meat-type chickens, resulting in considerable losses of chickens industry all over the world. In 2002, Xu Binrui, professor of China Agricultural University, discovered ALV-J in layer chickens. Naturally infected layer flocks showed the symptom of tabes and serious anemia, decrease of egg production, and the morality was as high as 10%. Maldevelopment was present in testes, ovaries, and oviducts. It caused considerable losses of layer chickens industry. Specific protein gp85 of ALV-J was detected through pathology, immunohistochemistry and PCR method. The layer chickens natural infected with ALV-J were first dicovered and reported in the world.
In the study isolation of virus was performed using the samples of layer chickens naturally infected with ALV-J. The tissue samples (hearts, livers, spleens etc) of illed chickens were filtrated and inoculated into chicken embryo fibroblast (CEF). The CEFs were detected using the indirect fluorescence assay (I FA) with specific monoclonal antibody against ALV-J gp85 at 7 day after inoculation. Specific fluorescence was detected in the cytoplasm of CEFs and the caryon was unstained. Specific envelope protein gp85 was detected by using immunohistochemistry. For further detection RT-PCR was performed on the CEF inoculated with the samples. A 545bp specific product was amplified using primer set H5 and H7 that was designed in 3' region of pol gene and 5' region of the gp85 encoding gene respectively. The product was as same size as expected and was sequenced by TaKaRa company. The sequence data showed that the identity of the nucleotide sequence was 97.4% and that of the amino acid sequence was 96.1% compared with that of the prototypical strain of ALV-J HPRS-103. The results of RT-PCR illuminated that the gp85 gene of ALV-J isolated from egg-type chickens have some variation compared with that of prototype HPRS-103 strain.
The results of this study demonstrated that ALV-J was successfully isolated from naturally infected egg-type chickens for the first time. The isolation of ALV-J in layer chickens will develope a new member for previously known diseases that affected the productivity of chickens, meanwhile, it will also provide theory and practice basis for better preventing, controlling the occurrence of ALV-J and decreasing the economic losses.
引文
[1] Payne, L. N., A. M. Gillespie, and K. Howes. 1992. Myeloid leukaemogenicity and transmission of the HPRS-103 strain of avian leukosis virus. Leukemia. 6:1167-1176
[2] Bai, J., L.N. Payne, and M.A. Skinner. 1995. HPRS-103 (exogenous avian leukosis virus, subgroup J) has an env gene related to those of endogenous elements EAV-0 and E51 and an E element found previously only in sarcoma viruses. J. Virol. 69:779-784
[3] Kim, Y., S. M. Gharaibeh, N. L. Stedman, et al. Comparison and verification of quantitive competitive reverse transcription polymerase chain reaction (QC-RT-PCR) and real time RT-PCR for avian leucosis virus subgroup J. J. Virol. Methods. 2002, 102:1-8
[4] Venugopal, K., L. M. Smith, K. Howes, et al. Antigentic variants of subgroup J avian leukosisvirus:Sequence analysis reveals mutiple changes in the env gene. J. Gen. Virol. 1998.79: 575-766
[5] 徐镔蕊，吕艳丽，董卫星等．蛋鸡骨髓细胞瘤病的病理学诊断．畜牧兽医学报，2002，33(6)：562-564
[6] Xu, B. R., W. X. Dong, Ch. M. Yu, et al. 2004. Occurrence of avian leukosis virus subgroup J in commercial layer flocks in China. Avian Pathol. 33:13-17
[7] 徐镔蕊，董卫星，何召庆，等．间接荧光抗体法快速诊断海兰褐蛋鸡J亚群禽白血病的研究．中国兽医杂志，2002，9(38)：7-9
[8] 徐镔蕊，董卫星，冯小宇等．采用SP法检测商品蛋鸡感染J亚群禽白血病病毒对产蛋性能的影响．中国家禽，2003，26(6)：9-12．
[9] Pentimalli, F. 1915. Ueber die Geschwuelste bei huehnern. I. Mittrilung. Ailgemeine morphologie der spontanen und der translantablen Huehnergeschuelste. Z Krebsforsh 15:111-153.
[10] Furth, J. 1931 .Erythroleukosis and the anemias of the fowl. Arch pathol 12: 1-30.
[11] Beard, J. W. Biology of avian oncornavirus. In G. Klein(ed.). Viral Oncology. Raven Press, New York, 1980. pp. 55-87.
[12] Mladenov, Z., U. Heine, D. Beard, et al. Strain MC29 avian leucosis virus. Myelocytoma, endothelima, and renal growths: Pathomorphological and ultrastructural aspects. J Natl Cancer Inst 1967. 16:877-888
[13] Payne L. N. The emergence of myeloid leucosis in meat type chickens [A]. 99 international conference and exibition on veterinary poultry proceeding [C]. Beijng: National Animal Husbandry and Veterinary Servive, China International Conference for Science and Technology, 1999.1-10
[14] Payne, L.N., S. R. Brown, N. Bumstead,et al Thouless. A novel subgroup of exogenous avian leukosis virus in chicken. J. Gen. Virol. 1991, 72:801-807
[15] Weiss, R.A., N. Teich, H. Varmus, et al. RNA Tumor Viruses, 2nd ed. Cold Spring Harbor Laboratory, Cold Spring Harbor, New York. 1982
[16] Wiebe Van der sluis著，宁长申，张龙现译．1998年世界禽病最新动态．预防兽医学进展，1999，1(2)：33-35
[17] Fadly A. M. and Smith E. J. Isolation and some characteristics of a subgroup J-like avian leukosisi virus associated with myeloid leucosis in meat-type chickens in the United States. Avian Dis. 1999. 43: 291-400
[18] Benson S. J., Ruis B. L., Garbes A. L., et al. Independent isolates of the emerging subgroup J avian leucosis virus derive from a common ancestor. J Virol. 1998.12:10301-10304 Veterinary Servive, China International Conference for Science and Technology, 1999.1-10
[19] 卡尔尼克BW.禽病学[M]．第9版．高福，刘文军译．北京：北京农业大学出版社，1991．334～381
[20] 殷震，刘景华．动物病毒学[M]．第2版．北京：科学出版社，1997．870～885
[21] Calnek, B. W. 1968. Lymphiod leukosis virus: A survey of cornmercial breeding flocks for genetic resistance and incidence of embryo infection. Avian Dis12:104-111
[22] Morgan, H. R. 1973. Avian leucosis-sarcoma virus antibodies in wildfowl, domestic chickens, and man in Kenya. Pro Soc Exp Biol Med 144:1-4
[23] Sandelin, K., T. Estola.1974. occurrence of different subgroup of avian leucosis virus in Finnish poultry. Avian Pathol 3:159-168
[24] Smith, E. J. 1987. Endogenous avian leukemia viruses. In G. F. DeBoer(ed.). Avian Leukosis. Martinus Nijhoff. Boston, MA, pp. 101-120
[25] Guillermo Zavala, et al. An Overview of Myeloid leucosis in Meat-type chickens. Technical News. Special Technical Bulletin, 1998,1:1-4
[26] Beard, J. W. 1963. Avian virus growths and their etiological agents. Adv Cancer Res 7:1-127
[27] Bauer, H. 1974. Virion and tumor cell antigens of C type RNA tumors viruses. Adv Cancer Res 20: 275-341
[28] Matthews, R. E. F. 1982. Fourth report of the international committee on taxonomy of viruses. Classification and nomenclature of viruses. Intervirology 17, Nos. 1-3
[29] Coffin, J. M. 1992. Structure and classification of retroviruses. In J. Levy(ed.). The Retroviridae, vol 1. Plenum Press, New York, p19-49
[30] Nowinski, R. C., E. Fleissner, and N. H. Sarkar. 1973. Structural and serological aspects of the oncornaviruses in vol. 8: Persisitent virus infections. Perspect Virol 8: 31-60
[31] Temin, H. M. 1974. The cellular and molecular biology of RNA tumor viruses, especially avian leucosis-sarcoma viruses, and their relatives. Adv Cancer Res19:47-104
[32] Beard, J. W. 1973. Oncornaviruses. I. The avian tumor viruses. In A. J. Dalton and F. Haguenau(eds.). Ultrastructure in Biological Systems, vol. 5. Ultrastructure of Animal Viruses and Bacteriophages. Academic Press, New York, pp. 261-181
[33] Robinson, W. S., AND P. H. Duesberg. The chemistry, of RNA tumor viruses. In H. Fraenkel-Conrat(ed.). Molecular Basis of Virology. Reinhold Book, New York, 1968. 306-331
[34] Friesen, B., and H. Rubin. 1961. Some physicochemical and immunologcal properties of an avian
leucosis virus (RIF). Virology 15:387-396
[35] Vogt, P. K. 1965. Avian tumor viruses. Adv Virus Res 11:293-385
[36] Eckert, E. A., I. Grenn, D. G. Sharp, D. Beard. and J. W. Virus of avian erythromyeloblastic leucosis. Ⅶ. Thermal stability of virus infectivity; of the virus particle; and of the enzyme dephosphorylating adenosinetriphosphate. J Natl Cancer Inst 1955.16:153-161
[37] Bryan, W. R. , J. B. Moloney, and D. Calnan. 1954. Stable standard preparations of the Rous sarcoma virus preserved by freezing and storage at low temperatures. J Natl Cancer Inst15:315-329
[38] Hayward, W. S., and B. G. Neel. Retroviral gene expression. Cure Top Microbiol Immunol. 1981. 217-276
[39] Young J. A. T. Avian leucosis virus-receptor interaction[J].AvianPathol, 1998,27:21～25
[40] 杜岩，崔治中，秦爱建，等．鸡的J亚群白血病病毒的分离及部分序列比较．病毒学报．2000，16(4)：341-346
[41] Bai, J., K. Howes, L.N. Payne, et al. 1995. Sequence of host range determinants in the env gene of a full length, infectious proviral clone of exogenous avian leucosis virus HPRS-103 confirms that it represents a new subgroup (designed J). J. Gen. Virol. 76:181-187
[42] Smith, L. M., A. A. Toye, K. Howes, et al. 1999. Novel endogenous retroviral sequence in the chicken genome closely related to HPRS-103 (subgroup J) avian leucosis virus. J. Gen. Virol. 80:261-268
[43] Benson, S. J., B. L. Ruis, A. M. Fadly, et al. 1998. The unique envelope gene of the subgroup J avian leukosis virus derives from ev/J proviruses, a novel family of avian endogenous viruses. J. Virol. 72:10157-10164
[44] Bolognesi, D. P. 1974. Structure components of RNAtumor viruses. Adv Virus Res 19:315-359
[45] Weiss, R.A., N. Teich, H. Varmus, et al. 1985. RNA Tumor Viruses, 2nd ed. Supplements and Appendices. Cold Spring Harbor Laboratory, Cold Spring Harbor, New York
[46] Guillermo Zavala.A new Challenge for the Poultry Industry Myeloid Leukosis.Vineland Update. 1998, 61(4): 665-668
[47] Young J. A. T. Avian leucosis virus-receptor interaction[J].AvianPathol, 1998,27:21～25
[48] Temin, H. M. 1974. The Bertner Foundation Memorial Award Lecture—from proviruses to protoviruses: RNA-directed DNA sybthesis by RNA tumor viruses and cells. Molecular Study in Viral Neoplasia. Williams& Wilkins, Baltinore, MD, pp. 7-38
[49] 杜岩，崔治中．J亚群禽白血病病毒中国分离株的人工致病性试验．中国农业科学，2002，35(4)：430-433
[50] H. W. Sung, S. M. Reddy, and A. M. Fadly. High virus titer in feather pulp of chickens infected with subgroup J avian leucosis virus
[51] Luciw, P. A. and N. J. Leung. Mechanisms of retroviral replication. In J. Levy(ed.). The Retrovirdae, vol 1, Plenum Press, New York. 1992. pp. 159-298
[52] Piraino, F. The mechaniam of genetic resistance of chick embryo cells to infection by Rous sarcoma virus-Bryan strain(BS-RSV). Virology. 1967.32:700-707
[53] Dales, S., and H. Hanafusa. Penetration and intracellular release of the genomes of avian RNA tumor viruses. Virology. 1972. 163: 452-461
[54] Cooper, G. M. 1982. Cellular transforming genes. Science 217: 801-806
[55] Enrietto, P. J., and M. J. Hayman. 1987. Structure and virus-associated oncogenes of avian sarcoma and leukemia viruses. In G. F. de Boer(ed.). Avian Leukosis. Martinus Nijhoff, Boston, MA, pp. 29-46
[56] Kung, H .J., and N. J. Maihle. 1987. Molecular basis of oncogenesis by non-avian retroviruses. In G. F. de Boer(ed.). Avian Leukosis. Martinus Nijhoff, Boston, MA, pp. 77-99
[57] Payne, L. N. 1992. Biology of avian retroviruses. In J. Levy(ed.). The Retriviridae, vol 1. Plenum Press, New York, pp. 299-404
[58] Rasheed, S. Retroviruses and Oncogenes [M]. New York: Plenum Press, 1995. 293～408
[59] Moscovici, C., and L. Gazzolo. 1987.Virus-cell interactions of avian sarcoma and defective leukosisi. Martinus Nijhoff, Boston, MA, pp. 151-169
[60] Cottral, G. E., B. R. Burmester, and N. F. Waters. 1954. Egg transmission of avian lymphomatosis. Poult Sci 33: 1174-1184
[61] Rubin, H., A. Cornelius, and L. Fishier. 1961. The pattern of congenital transmission of an avian leucosis virus. Prov Natl Acad Sci USA47: 1058-1060
[62] Rubin, H., L. Fishier, A. Cornelius, et al. 1962. Tolerance and immunity in chickens after congenital and contact infection with an avian leucosis virus. Virology 17:143-156
[63] 余兵译，朱国强，王水坤校．一种正在威胁养禽业的新病—J亚型白细胞增生病．中国禽业导刊．1999，16(3)：11
[64] Purchase, H. G., W. Okazaki, and B. R. Burmester. Long-term field trials with the herpesvirus of turkeys vaccine against Marek's disease. Avian Dis. 1972.16: 57-71
[65] Mathey, W. J. 1977. Personal communication
[66] Payne. L. N., et al. Recovery of acutely transforming virus from myeloid leucosis induced by the HPRS-103 strain of avian leucosis virus. Avian Dis. 1993, 37: 428-450
[67] Payne. L. N. HPRS-103: retrovirus strikes back. The emergence of subgroup J avian leucosis virus. Avian Pathol. 1998, 27: 36-45
[68] 张志．我国J亚群禽白血病病毒的流行病学调查和囊膜蛋白env基因的克隆测序及序列分析．[硕士学位论文]．山东农业大学，2001
[69] Hwang, C., W. Ching-Ho. Serologic profiles of chickens infected with subgroup J avian leucosis virus. Avian Dis. 2002, 46:598-604
[70] 杜岩，崔治中，秦爱建，等．从市场商品肉鸡中检出J亚群禽白血病病毒．中国家禽学报．1999， 3(1)：1-4
[71] 杜岩．J亚群禽白血病病毒中国株的分离、分子鉴定和致病性研究及其囊膜糖蛋白gp85基因的表达[D]．江苏：扬州大学，2000
[72] 张志等，商品代肉鸡J亚群禽白血病的病理及病毒分离鉴定．中国兽医杂志．2002，38(6)：6-8．
[73] Smith, E. J., Williams, S. M., and Fadly, A. M. Detection of avian leucosis virus subgroup J using the polymerase chain reaction. Avian Dis.1998.42: 375-380
[74] 关云涛，李昌文，张立成等．应用PCR和琼脂扩散检测禽白血病病毒的比较．中国试验动物杂志．2002，12(5)：306-308
[75] Arshad, S. S., Smith, L. M., Howes, K., et al. Tropism of subgroup J avian leucosis virus as detected by in situ hybridization. Avian Pathol. 1999, 28: 163-169.
[76] 秦爱建，Lucy,L.禽白血病病毒J亚群重组囊膜蛋白基因的表达效果．中国兽医学报．2001，21(5)：425-428
[77] Venugopal, K., K. Howes, G. S. Barron, et al. Recominat env-gp85 of HPRS-103(subgroup J) avian ieucosis virus: antigenic characteristic and usefulness as a diagnostic reagent. Avian Dis. 1997, 41: 283-288
[78] Kreager K S.Chicken industry strategies for control of tumor virus infections [J]. Poult Sci, 1998, 77:1213-1216
[79] Williams, S. M., W. M. Reed, and A. M. Fadly. Influence of age of exposure on the response of line 0 and line 63 chickens to infection with subgroup J avian leucosis virus. In: Proc. International Symposium on ALV-J and other avian retroviruses, Rauischholzhausen, Germany. 2000. pp. 57-76
[80] Gingrich, E., Porter, R. E., Lupiani, B., et al. Diagnosis of myeloid leucosis induced by a recombinant avian leucosis virus in commercial white leghorn egg laying flocks. Avian Dis. 2002. 46: 745-748
[81] 徐镔蕊，冯小宇，董卫星等．端粒、端粒酶与肿瘤的研究进展．动物医学进展．2003．24(3)：1-4
[82] 徐镔蕊，董卫星，何召庆，等．突变型抑癌基因p53在蛋鸡J亚群禽白血病中的表达．农业生物技术学报．2003，11(3)：276-279
[83] Bishop J. M. Cellular oncogenes and retroviruses. Ann. Rev. Biochem. 1983.52:301-354
[84] Boyce-Jacino M. T., Donoghue K. O., and Faras A. J. Multipe complex families of endogenous retrovirus are highly conserved in the genus Gallus. J Virol. 1992.66:4919-4929
[85] Bowers W. J. and Ruddell A. a1/EBP: a leucine zipper protein that binds CCAAT/enhancer elements in the avian leukosis virus long terminal repeat enhancer. J. Virol. 1992 66:6578-6586
[86] Sambrook J．分子克隆实验指南[M]．第2版．北京：科学出版社，1999
[87] 卢圣栋．现在分子生物学实验技术．北京：高等教育出版社．1993
[88] 苏慧慈，刘彦仿．原位PCR．北京：科学出版社．1995
[89] 杜岩．J.亚群禽白血病病毒中国株的分离、分子鉴定和致病性研究及其囊膜糖蛋白gp85基因的表达．江苏扬州：扬州大学，2000
[90] Lupiani B., Hunt H., Silva R., et al. Identification and characterization of recombinant subgroup J avian leucosis viruses (ALV) expressing subgroup A ALV envelope. Virology. 2000. 276(1): 37-43
[91] Smith, L. M., S. R. Brown, K. Howes, et al. 1998. Development and application of polymerase chain reaction (PCR) tests for the detection of subgroup J avian leucosis virus. Virus. Res. 54: 87-98
[92] 赵振华，顾玉芳，郝永清等，禽骨髓细胞瘤病的病理学研究(初报)．动物医学进展，1999，20(3)：85～86
[93] 杜岩，崔治中，秦爱建等．用PCR从市场商品肉鸡中检出J亚群白血病病毒感染．99
北京国际禽病会议暨展览会论文集．北京：全国畜牧兽医总站，中国国际科技会议中心，1999，284．
[94] 杜岩，崔治中，秦爱建等．鸡的J亚群白血病病毒的分离及部分序列比较。病毒学报，2000，16(4)：341—346．
[95] 成子强，哈斯阿古拉，赵振华．禽骨髓细胞瘤病病毒NM0138株gp85基因的分子克隆和序列分析．中国畜牧兽医学会兽医病理学分会第十一次兽医病理学、第十次动物病理生理学学术研讨会沦文集，杭州：中国畜牧兽医学会兽医病理学分会，2001，192．197
[96] 成了强，赵振华，郝永清等．禽骨髓细胞瘤病的血清学调查及PCR诊断．中国病毒学报，2002，17(4)：371-373
[97] 成子强，赵心力，郝永清等．应用ELISA方法调查禽骨髓细胞瘤病，中国家禽，2002，24(6)：9-10．
[98] Crittenden, L. B., et al. A comparision of test materials for differentiating avian leucosis virus group-specific antigens of exogenous and endogenous origin. Avian Dis, 1984, 28: 1057-1070
[99] Fadly, A. M., et al. Relative efficiencyof test proceducers to detect lymphoid leucosis virus infection. Poult. Sci, 1981,60:2037-2044
[100] Korec, E., et al. A rapid detection of avian oncovirus group-specific antigens in feather pulp by the enzyme-linked immunosorbent assay. Folia boil. 1984,30:15-23
[101] Mizuno, Y., et al. Enzyme-linked immunosorbent assay to detect subgroup-specific antibodies to avian leucosis viruses. Am. J. Vet. Res. 1986.47:551-556
[102] Nicholas, R. A. J.,et al. Detection of acian leucosis virus: comparision of five techniques. Res. Vet. Sci. 1984,37:371-373
[103] Stehelin D., Varmus H. E., Bishop J. M., et al, DNA related to the transforming gene(S) of avian sarcoma virus is present in normal avian DNA. Nature. 1976. 260:170-173
[104] 秦爱建．禽白血病病毒J亚群囊膜糖蛋白基因的生物学和生物化学特性．博士学位论文[D]，扬州大学，1999．
[105] 王海艳，张二芹，王金玲，等．用ELISA对种蛋中禽骨髓细胞瘤病毒的母源抗体调查．动物医学进展．2003，24(5)：110-111
[106] Hauptli D., Bruckner L., Ottiger H. P. Use of reverse transcriptase polymerase chain reaction for detection of vaccine contamination by avian leucosis virus. J Virol MethodS. 1997.66(1): 71-81
[107] Pham T. D., Spencer J. L., Vicki L. T. Detection of exogenous and endogenous avian leucosis virus in commercial chicken eggs using reverse transcription and polymerase chain reaction assay. Avian Pathol. 1999.28:385-392
[2] Bai, J., L.N. Payne, and M.A. Skinner. 1995. HPRS-103 (exogenous avian leukosis virus, subgroup J) has an env gene related to those of endogenous elements EAV-0 and E51 and an E element found previously only in sarcoma viruses. J. Virol. 69:779-784
[3] Kim, Y., S. M. Gharaibeh, N. L. Stedman, et al. Comparison and verification of quantitive competitive reverse transcription polymerase chain reaction (QC-RT-PCR) and real time RT-PCR for avian leucosis virus subgroup J. J. Virol. Methods. 2002, 102:1-8
[4] Venugopal, K., L. M. Smith, K. Howes, et al. Antigentic variants of subgroup J avian leukosisvirus:Sequence analysis reveals mutiple changes in the env gene. J. Gen. Virol. 1998.79: 575-766
[5] 徐镔蕊，吕艳丽，董卫星等．蛋鸡骨髓细胞瘤病的病理学诊断．畜牧兽医学报，2002，33(6)：562-564
[6] Xu, B. R., W. X. Dong, Ch. M. Yu, et al. 2004. Occurrence of avian leukosis virus subgroup J in commercial layer flocks in China. Avian Pathol. 33:13-17
[7] 徐镔蕊，董卫星，何召庆，等．间接荧光抗体法快速诊断海兰褐蛋鸡J亚群禽白血病的研究．中国兽医杂志，2002，9(38)：7-9
[8] 徐镔蕊，董卫星，冯小宇等．采用SP法检测商品蛋鸡感染J亚群禽白血病病毒对产蛋性能的影响．中国家禽，2003，26(6)：9-12．
[9] Pentimalli, F. 1915. Ueber die Geschwuelste bei huehnern. I. Mittrilung. Ailgemeine morphologie der spontanen und der translantablen Huehnergeschuelste. Z Krebsforsh 15:111-153.
[10] Furth, J. 1931 .Erythroleukosis and the anemias of the fowl. Arch pathol 12: 1-30.
[11] Beard, J. W. Biology of avian oncornavirus. In G. Klein(ed.). Viral Oncology. Raven Press, New York, 1980. pp. 55-87.
[12] Mladenov, Z., U. Heine, D. Beard, et al. Strain MC29 avian leucosis virus. Myelocytoma, endothelima, and renal growths: Pathomorphological and ultrastructural aspects. J Natl Cancer Inst 1967. 16:877-888
[13] Payne L. N. The emergence of myeloid leucosis in meat type chickens [A]. 99 international conference and exibition on veterinary poultry proceeding [C]. Beijng: National Animal Husbandry and Veterinary Servive, China International Conference for Science and Technology, 1999.1-10
[14] Payne, L.N., S. R. Brown, N. Bumstead,et al Thouless. A novel subgroup of exogenous avian leukosis virus in chicken. J. Gen. Virol. 1991, 72:801-807
[15] Weiss, R.A., N. Teich, H. Varmus, et al. RNA Tumor Viruses, 2nd ed. Cold Spring Harbor Laboratory, Cold Spring Harbor, New York. 1982
[16] Wiebe Van der sluis著，宁长申，张龙现译．1998年世界禽病最新动态．预防兽医学进展，1999，1(2)：33-35
[17] Fadly A. M. and Smith E. J. Isolation and some characteristics of a subgroup J-like avian leukosisi virus associated with myeloid leucosis in meat-type chickens in the United States. Avian Dis. 1999. 43: 291-400
[18] Benson S. J., Ruis B. L., Garbes A. L., et al. Independent isolates of the emerging subgroup J avian leucosis virus derive from a common ancestor. J Virol. 1998.12:10301-10304 Veterinary Servive, China International Conference for Science and Technology, 1999.1-10
[19] 卡尔尼克BW.禽病学[M]．第9版．高福，刘文军译．北京：北京农业大学出版社，1991．334～381
[20] 殷震，刘景华．动物病毒学[M]．第2版．北京：科学出版社，1997．870～885
[21] Calnek, B. W. 1968. Lymphiod leukosis virus: A survey of cornmercial breeding flocks for genetic resistance and incidence of embryo infection. Avian Dis12:104-111
[22] Morgan, H. R. 1973. Avian leucosis-sarcoma virus antibodies in wildfowl, domestic chickens, and man in Kenya. Pro Soc Exp Biol Med 144:1-4
[23] Sandelin, K., T. Estola.1974. occurrence of different subgroup of avian leucosis virus in Finnish poultry. Avian Pathol 3:159-168
[24] Smith, E. J. 1987. Endogenous avian leukemia viruses. In G. F. DeBoer(ed.). Avian Leukosis. Martinus Nijhoff. Boston, MA, pp. 101-120
[25] Guillermo Zavala, et al. An Overview of Myeloid leucosis in Meat-type chickens. Technical News. Special Technical Bulletin, 1998,1:1-4
[26] Beard, J. W. 1963. Avian virus growths and their etiological agents. Adv Cancer Res 7:1-127
[27] Bauer, H. 1974. Virion and tumor cell antigens of C type RNA tumors viruses. Adv Cancer Res 20: 275-341
[28] Matthews, R. E. F. 1982. Fourth report of the international committee on taxonomy of viruses. Classification and nomenclature of viruses. Intervirology 17, Nos. 1-3
[29] Coffin, J. M. 1992. Structure and classification of retroviruses. In J. Levy(ed.). The Retroviridae, vol 1. Plenum Press, New York, p19-49
[30] Nowinski, R. C., E. Fleissner, and N. H. Sarkar. 1973. Structural and serological aspects of the oncornaviruses in vol. 8: Persisitent virus infections. Perspect Virol 8: 31-60
[31] Temin, H. M. 1974. The cellular and molecular biology of RNA tumor viruses, especially avian leucosis-sarcoma viruses, and their relatives. Adv Cancer Res19:47-104
[32] Beard, J. W. 1973. Oncornaviruses. I. The avian tumor viruses. In A. J. Dalton and F. Haguenau(eds.). Ultrastructure in Biological Systems, vol. 5. Ultrastructure of Animal Viruses and Bacteriophages. Academic Press, New York, pp. 261-181
[33] Robinson, W. S., AND P. H. Duesberg. The chemistry, of RNA tumor viruses. In H. Fraenkel-Conrat(ed.). Molecular Basis of Virology. Reinhold Book, New York, 1968. 306-331
[34] Friesen, B., and H. Rubin. 1961. Some physicochemical and immunologcal properties of an avian
leucosis virus (RIF). Virology 15:387-396
[35] Vogt, P. K. 1965. Avian tumor viruses. Adv Virus Res 11:293-385
[36] Eckert, E. A., I. Grenn, D. G. Sharp, D. Beard. and J. W. Virus of avian erythromyeloblastic leucosis. Ⅶ. Thermal stability of virus infectivity; of the virus particle; and of the enzyme dephosphorylating adenosinetriphosphate. J Natl Cancer Inst 1955.16:153-161
[37] Bryan, W. R. , J. B. Moloney, and D. Calnan. 1954. Stable standard preparations of the Rous sarcoma virus preserved by freezing and storage at low temperatures. J Natl Cancer Inst15:315-329
[38] Hayward, W. S., and B. G. Neel. Retroviral gene expression. Cure Top Microbiol Immunol. 1981. 217-276
[39] Young J. A. T. Avian leucosis virus-receptor interaction[J].AvianPathol, 1998,27:21～25
[40] 杜岩，崔治中，秦爱建，等．鸡的J亚群白血病病毒的分离及部分序列比较．病毒学报．2000，16(4)：341-346
[41] Bai, J., K. Howes, L.N. Payne, et al. 1995. Sequence of host range determinants in the env gene of a full length, infectious proviral clone of exogenous avian leucosis virus HPRS-103 confirms that it represents a new subgroup (designed J). J. Gen. Virol. 76:181-187
[42] Smith, L. M., A. A. Toye, K. Howes, et al. 1999. Novel endogenous retroviral sequence in the chicken genome closely related to HPRS-103 (subgroup J) avian leucosis virus. J. Gen. Virol. 80:261-268
[43] Benson, S. J., B. L. Ruis, A. M. Fadly, et al. 1998. The unique envelope gene of the subgroup J avian leukosis virus derives from ev/J proviruses, a novel family of avian endogenous viruses. J. Virol. 72:10157-10164
[44] Bolognesi, D. P. 1974. Structure components of RNAtumor viruses. Adv Virus Res 19:315-359
[45] Weiss, R.A., N. Teich, H. Varmus, et al. 1985. RNA Tumor Viruses, 2nd ed. Supplements and Appendices. Cold Spring Harbor Laboratory, Cold Spring Harbor, New York
[46] Guillermo Zavala.A new Challenge for the Poultry Industry Myeloid Leukosis.Vineland Update. 1998, 61(4): 665-668
[47] Young J. A. T. Avian leucosis virus-receptor interaction[J].AvianPathol, 1998,27:21～25
[48] Temin, H. M. 1974. The Bertner Foundation Memorial Award Lecture—from proviruses to protoviruses: RNA-directed DNA sybthesis by RNA tumor viruses and cells. Molecular Study in Viral Neoplasia. Williams& Wilkins, Baltinore, MD, pp. 7-38
[49] 杜岩，崔治中．J亚群禽白血病病毒中国分离株的人工致病性试验．中国农业科学，2002，35(4)：430-433
[50] H. W. Sung, S. M. Reddy, and A. M. Fadly. High virus titer in feather pulp of chickens infected with subgroup J avian leucosis virus
[51] Luciw, P. A. and N. J. Leung. Mechanisms of retroviral replication. In J. Levy(ed.). The Retrovirdae, vol 1, Plenum Press, New York. 1992. pp. 159-298
[52] Piraino, F. The mechaniam of genetic resistance of chick embryo cells to infection by Rous sarcoma virus-Bryan strain(BS-RSV). Virology. 1967.32:700-707
[53] Dales, S., and H. Hanafusa. Penetration and intracellular release of the genomes of avian RNA tumor viruses. Virology. 1972. 163: 452-461
[54] Cooper, G. M. 1982. Cellular transforming genes. Science 217: 801-806
[55] Enrietto, P. J., and M. J. Hayman. 1987. Structure and virus-associated oncogenes of avian sarcoma and leukemia viruses. In G. F. de Boer(ed.). Avian Leukosis. Martinus Nijhoff, Boston, MA, pp. 29-46
[56] Kung, H .J., and N. J. Maihle. 1987. Molecular basis of oncogenesis by non-avian retroviruses. In G. F. de Boer(ed.). Avian Leukosis. Martinus Nijhoff, Boston, MA, pp. 77-99
[57] Payne, L. N. 1992. Biology of avian retroviruses. In J. Levy(ed.). The Retriviridae, vol 1. Plenum Press, New York, pp. 299-404
[58] Rasheed, S. Retroviruses and Oncogenes [M]. New York: Plenum Press, 1995. 293～408
[59] Moscovici, C., and L. Gazzolo. 1987.Virus-cell interactions of avian sarcoma and defective leukosisi. Martinus Nijhoff, Boston, MA, pp. 151-169
[60] Cottral, G. E., B. R. Burmester, and N. F. Waters. 1954. Egg transmission of avian lymphomatosis. Poult Sci 33: 1174-1184
[61] Rubin, H., A. Cornelius, and L. Fishier. 1961. The pattern of congenital transmission of an avian leucosis virus. Prov Natl Acad Sci USA47: 1058-1060
[62] Rubin, H., L. Fishier, A. Cornelius, et al. 1962. Tolerance and immunity in chickens after congenital and contact infection with an avian leucosis virus. Virology 17:143-156
[63] 余兵译，朱国强，王水坤校．一种正在威胁养禽业的新病—J亚型白细胞增生病．中国禽业导刊．1999，16(3)：11
[64] Purchase, H. G., W. Okazaki, and B. R. Burmester. Long-term field trials with the herpesvirus of turkeys vaccine against Marek's disease. Avian Dis. 1972.16: 57-71
[65] Mathey, W. J. 1977. Personal communication
[66] Payne. L. N., et al. Recovery of acutely transforming virus from myeloid leucosis induced by the HPRS-103 strain of avian leucosis virus. Avian Dis. 1993, 37: 428-450
[67] Payne. L. N. HPRS-103: retrovirus strikes back. The emergence of subgroup J avian leucosis virus. Avian Pathol. 1998, 27: 36-45
[68] 张志．我国J亚群禽白血病病毒的流行病学调查和囊膜蛋白env基因的克隆测序及序列分析．[硕士学位论文]．山东农业大学，2001
[69] Hwang, C., W. Ching-Ho. Serologic profiles of chickens infected with subgroup J avian leucosis virus. Avian Dis. 2002, 46:598-604
[70] 杜岩，崔治中，秦爱建，等．从市场商品肉鸡中检出J亚群禽白血病病毒．中国家禽学报．1999， 3(1)：1-4
[71] 杜岩．J亚群禽白血病病毒中国株的分离、分子鉴定和致病性研究及其囊膜糖蛋白gp85基因的表达[D]．江苏：扬州大学，2000
[72] 张志等，商品代肉鸡J亚群禽白血病的病理及病毒分离鉴定．中国兽医杂志．2002，38(6)：6-8．
[73] Smith, E. J., Williams, S. M., and Fadly, A. M. Detection of avian leucosis virus subgroup J using the polymerase chain reaction. Avian Dis.1998.42: 375-380
[74] 关云涛，李昌文，张立成等．应用PCR和琼脂扩散检测禽白血病病毒的比较．中国试验动物杂志．2002，12(5)：306-308
[75] Arshad, S. S., Smith, L. M., Howes, K., et al. Tropism of subgroup J avian leucosis virus as detected by in situ hybridization. Avian Pathol. 1999, 28: 163-169.
[76] 秦爱建，Lucy,L.禽白血病病毒J亚群重组囊膜蛋白基因的表达效果．中国兽医学报．2001，21(5)：425-428
[77] Venugopal, K., K. Howes, G. S. Barron, et al. Recominat env-gp85 of HPRS-103(subgroup J) avian ieucosis virus: antigenic characteristic and usefulness as a diagnostic reagent. Avian Dis. 1997, 41: 283-288
[78] Kreager K S.Chicken industry strategies for control of tumor virus infections [J]. Poult Sci, 1998, 77:1213-1216
[79] Williams, S. M., W. M. Reed, and A. M. Fadly. Influence of age of exposure on the response of line 0 and line 63 chickens to infection with subgroup J avian leucosis virus. In: Proc. International Symposium on ALV-J and other avian retroviruses, Rauischholzhausen, Germany. 2000. pp. 57-76
[80] Gingrich, E., Porter, R. E., Lupiani, B., et al. Diagnosis of myeloid leucosis induced by a recombinant avian leucosis virus in commercial white leghorn egg laying flocks. Avian Dis. 2002. 46: 745-748
[81] 徐镔蕊，冯小宇，董卫星等．端粒、端粒酶与肿瘤的研究进展．动物医学进展．2003．24(3)：1-4
[82] 徐镔蕊，董卫星，何召庆，等．突变型抑癌基因p53在蛋鸡J亚群禽白血病中的表达．农业生物技术学报．2003，11(3)：276-279
[83] Bishop J. M. Cellular oncogenes and retroviruses. Ann. Rev. Biochem. 1983.52:301-354
[84] Boyce-Jacino M. T., Donoghue K. O., and Faras A. J. Multipe complex families of endogenous retrovirus are highly conserved in the genus Gallus. J Virol. 1992.66:4919-4929
[85] Bowers W. J. and Ruddell A. a1/EBP: a leucine zipper protein that binds CCAAT/enhancer elements in the avian leukosis virus long terminal repeat enhancer. J. Virol. 1992 66:6578-6586
[86] Sambrook J．分子克隆实验指南[M]．第2版．北京：科学出版社，1999
[87] 卢圣栋．现在分子生物学实验技术．北京：高等教育出版社．1993
[88] 苏慧慈，刘彦仿．原位PCR．北京：科学出版社．1995
[89] 杜岩．J.亚群禽白血病病毒中国株的分离、分子鉴定和致病性研究及其囊膜糖蛋白gp85基因的表达．江苏扬州：扬州大学，2000
[90] Lupiani B., Hunt H., Silva R., et al. Identification and characterization of recombinant subgroup J avian leucosis viruses (ALV) expressing subgroup A ALV envelope. Virology. 2000. 276(1): 37-43
[91] Smith, L. M., S. R. Brown, K. Howes, et al. 1998. Development and application of polymerase chain reaction (PCR) tests for the detection of subgroup J avian leucosis virus. Virus. Res. 54: 87-98
[92] 赵振华，顾玉芳，郝永清等，禽骨髓细胞瘤病的病理学研究(初报)．动物医学进展，1999，20(3)：85～86
[93] 杜岩，崔治中，秦爱建等．用PCR从市场商品肉鸡中检出J亚群白血病病毒感染．99
北京国际禽病会议暨展览会论文集．北京：全国畜牧兽医总站，中国国际科技会议中心，1999，284．
[94] 杜岩，崔治中，秦爱建等．鸡的J亚群白血病病毒的分离及部分序列比较。病毒学报，2000，16(4)：341—346．
[95] 成子强，哈斯阿古拉，赵振华．禽骨髓细胞瘤病病毒NM0138株gp85基因的分子克隆和序列分析．中国畜牧兽医学会兽医病理学分会第十一次兽医病理学、第十次动物病理生理学学术研讨会沦文集，杭州：中国畜牧兽医学会兽医病理学分会，2001，192．197
[96] 成了强，赵振华，郝永清等．禽骨髓细胞瘤病的血清学调查及PCR诊断．中国病毒学报，2002，17(4)：371-373
[97] 成子强，赵心力，郝永清等．应用ELISA方法调查禽骨髓细胞瘤病，中国家禽，2002，24(6)：9-10．
[98] Crittenden, L. B., et al. A comparision of test materials for differentiating avian leucosis virus group-specific antigens of exogenous and endogenous origin. Avian Dis, 1984, 28: 1057-1070
[99] Fadly, A. M., et al. Relative efficiencyof test proceducers to detect lymphoid leucosis virus infection. Poult. Sci, 1981,60:2037-2044
[100] Korec, E., et al. A rapid detection of avian oncovirus group-specific antigens in feather pulp by the enzyme-linked immunosorbent assay. Folia boil. 1984,30:15-23
[101] Mizuno, Y., et al. Enzyme-linked immunosorbent assay to detect subgroup-specific antibodies to avian leucosis viruses. Am. J. Vet. Res. 1986.47:551-556
[102] Nicholas, R. A. J.,et al. Detection of acian leucosis virus: comparision of five techniques. Res. Vet. Sci. 1984,37:371-373
[103] Stehelin D., Varmus H. E., Bishop J. M., et al, DNA related to the transforming gene(S) of avian sarcoma virus is present in normal avian DNA. Nature. 1976. 260:170-173
[104] 秦爱建．禽白血病病毒J亚群囊膜糖蛋白基因的生物学和生物化学特性．博士学位论文[D]，扬州大学，1999．
[105] 王海艳，张二芹，王金玲，等．用ELISA对种蛋中禽骨髓细胞瘤病毒的母源抗体调查．动物医学进展．2003，24(5)：110-111
[106] Hauptli D., Bruckner L., Ottiger H. P. Use of reverse transcriptase polymerase chain reaction for detection of vaccine contamination by avian leucosis virus. J Virol MethodS. 1997.66(1): 71-81
[107] Pham T. D., Spencer J. L., Vicki L. T. Detection of exogenous and endogenous avian leucosis virus in commercial chicken eggs using reverse transcription and polymerase chain reaction assay. Avian Pathol. 1999.28:385-392