摘要
鸭病毒肠炎(duck viral enteritis,DVE),又名鸭瘟(duck plague,DP),是由鸭肠炎病毒(duckEnteritis virus,DEV)引起的鸭、鹅及多种雁形目禽类的一种急性、热性、败血性传染病。其特征是流行广泛,传播迅速,发病率和死亡率高,给养鸭业造成巨大的经济损失。
DEV具有疱疹病毒典型的形态和结构,因此划分为疱疹病毒科,其亚科、属、种分类地位尚未确定。大多数疱疹病毒有12种糖蛋白,gB是最保守的结构糖蛋白之一,在多数疱疹病毒中gB同源物由ul27基因编码。该糖蛋白是病毒复制所必须的,参与病毒吸附、融合穿入及病毒在细胞间的传播,能刺激机体产生中和抗体,介导细胞免疫,是重组疫苗和亚单位疫苗的候选抗原,也是哺乳动物疱疹病毒血清学诊断方法的候选抗原。目前还没有关于DEVul27基因及其编码产物的深入研究。
本试验以DEV ul27基因为研究对象,采用PCR技术克隆了DEV C-KCE株ul27基因,并从该基因与其他疱疹病毒的同源性、编码蛋白的结构及抗原性等方面进行了较为详尽的生物信息学分析,在此基础上利用原核表达载体pET-32a在Rosetta(DE3)PlysS宿主菌中分别表达了该基因9个抗原片段,以鸭肠炎病毒高免血清为第一抗体,通过Western blot确定了UL27蛋白的B细胞线性抗原表位优势区,以制备的鼠源抗原肽特异性抗血清进行蚀斑减数试验确定了中和抗原表位的优势区,利用体外淋巴细胞转化试验揭示了外周血抗原特异性T淋巴细胞对DEV的回忆应答规律,通过统计分析初步确定了T淋巴细胞的抗原表位。
结果表明:DEV C-KCE株ul27基因完整ORF为3003bp,编码1000个氨基酸,同源性分析表明该基因为α疱疹病毒gB同源物,其遗传进化分析提示DEV与α疱疹病毒马立克氏病毒属亲缘关系最近;ul27基因编码产物具有典型跨膜蛋白结构,推测其信号肽序列位于N端前105个氨基酸以内,胞外区从信号肽裂解位点至832位氨基酸,跨膜区为833～883位氨基酸,胞内区为884～1000位氨基酸;蛋白胞外区存在8个潜在的N糖基化位点,10个保守的半胱氨酸,具有潜在的翻译修饰后的酶解位点,HSPGs结合位点、PACS-1结合基序,符合α疱疹病毒gB的结构特征和功能特征;Western blot结果表明UL27-2、UL27-5、UL27-7、UL27-9抗原肽存在B细胞线性表位,其中UL27-5抗原性最强,UL27-7刺激机体产生的抗体持续时间最长;蚀斑减数试验确定UL27-5、UL27-6、UL27-7、UL7-8、UL27-9、UL27-10抗原肽的抗血清具有中和活性,其中UL27-7、UL27-8是中和表位优势区;体外淋巴细胞转化试验表明,9段抗原肽及其不同组合均具有诱导外周血淋巴细胞发生转化的能力,外周血抗原特异性淋巴细胞在回忆应答中对DEV及UL27抗原肽的反应能力呈现缓慢增强,达到高峰后又快速回落的趋势。经统计分析初步确定DEV UL27 T细胞表位优势区是UL27-6,候选优势区是UL27-4、UL27-10。
本研究为明确DEV的分类地位,阐明DEV gB的功能,了解其在机体免疫过程中发挥的作用及其在实践中应用的可能性提供了理论依据和物质基础。
Duck viral enteritis(DVE) named duck plague(DP) also,its pathogen is duck enteritis virus (DEV),which is acute,heat and haemorrhagic contagious viral disease in infected birds of the order Anseriformes(ducks,geese and swans).DEV is characterized by epidemic broadly,spread rapidly with high mortality and morbidity,it has been responsible for considerable economic loss to the commercial duck industry.
DEV is a member of the herpesviridae according to it has the classical morphology and structure of herpesivirus,but its definite taxonomic classification was unknown.Glycoprotein play the very important role during the course of pathopoiesia.There are 12 glycoprotein in herpesvirus, and gB homologue has been identified in every herpesvirus studied to date,which is the most highly conserved herpesvirus structural glycoprotein.It was encoded by ul27 gene usually. Glycoprotein B is required for herpesvirus infectivity and is involved in virus adsorption, membrane fusion,penetration and cell-cell spread.In addition,gB elicits neutralizing antibodies, cell-mediated immune responses,and has been shown to be a candidate antigen for recombinant, subunit vaccines,serology diagnostic assays in mammalian.There is very limited in study about ul27 genen and UL27 protein of DEV.
The DEV ul27gene is the interest of this study,and was cloned frome the C-KCE strain by PCR.The homology of ul27 genen and UL27 protein with other herpesvirus,the structure and the antigenicity of UL27 protein were analysised by bioinformatics.Nine truncated antigenic peptides were expressed in Escherichia coli Rosetta(DE3)PlysS by prokaryotic expression vector pET-32a.The immunodomiannt region of B cell antigen epitope were identified by western blot with the duck serum against DEV.And the immunodomiannt region of neutralization epitope were identified by plaque inhibition test with mice serum against the nine truncated antigenic peptides The rule of the peripheral blood lymphocytes response to UL27 after infection DEV was revealed by LTT.In addition the T cell epitopes were defined primarily by statistics analysis.
The results of study as followed:The ORF of ul27 gene from DEV C-KCE strain was consist of 3003 bsae pairs and it enconded a primary translation product of 1000 amino acids.The homology analysis with other herpesvirus indicated ul27 gene was the gB homologue,and Phylogenetic analysis to illustrate the evolutionary relationship of DEV was close to subfamily Alphaherpesvirinae and Mardivirus genus.The DEV UL27 possessed classical structure of transmembane glycoprotein,the putative signal sequence located in the 105 aa of N terminus,the extracellular domain of UL27 from the cleavage site of signal peptide to 832 amino acid long,and transmembane domain from 833 to 884 amino acid long,the cytoplasmic domain from 884 to 1000 amino acid.Bioinformatic analysis of UL27 indicated there were eight potential N-linked glycosylation sites,10 Cys conserved residues,HSPGs binding sites,PACS-1 binding motif in extraceilular domain,all this structure consisted with the structure and function characters of herpesvirus gB.The B cell epitopes were identified in ul27-2、ul27-5、ul27-7、ul27-9 by Western blot,and the ul27-5 was the strongest antigenicity epitope,the antibody against ul27-7 lasted longest.The antibody against ul27-5、ul27-6、ul27-7、ul27-8、ul27-9、ul27-10 derived from the mice have the ability of neutralized DEV by plaque inhibition test,and antibody against ul27-7、ul27-8 show off the greater neutralizing capacity,so ul27-7 and ul27-8 were thought to be region of the immunodomiannt neutralization epitope excist.The nine truncated antigenic peptides displayed the ability to induce the peripheral blood lymphocytes transformation in vitro by lymphocyte transformation test(LTT).The peripheral blood antigen specific lymphocytes presented some tendency in response to the DEV,nine truncated antigenic peptides and their different combination, it was the response ability of peripheral blood antigen specific lymphocytes strengthened slowly then declined rapidly after reaching the peak in recall response.The T cell immunodomiannt epitopes of UL27 located in ul27-6 by statistic analysis primarily,and candidate prevalent epitopes region are ul27-4、ul27-10.
The results of this study were very significance for illustration the classify of DEV, understanding the function of DEV gB,revealing the role in immune response of DEV gB,and the possibility to be candidate antigen in vaccine or diagnostic assays.
引文
贝怀国,冯连虎等.2005.雏鹅怎么暴发鸭病毒肠炎病毒病.中国家禽年27(13):59
蔡明法,周承永,胡仕福等.2000.鸭瘟流行的新特征和防治对策.浙江畜牧兽医,3:24
曹贺芳.2008.鸭瘟的症状及其防制.农村养殖技术,2:24
陈宏武,徐为燕,杜念兴.1986.鸭瘟病毒核酸限制性内切酶图谱.病毒学报.2(2):184-186
陈建君,张毓金,杨增岐等.2003.鸭肠炎病毒PCR检测及其产物分析.动物医学进展.24(5):71-73.
陈淑红.2006.鸭肠炎病毒部分基因组序列的克隆与分析.东北农业大学.博士学位论文.46-47、61-73
陈祥平,余兆昌.2001.浅谈鸭瘟免疫失败的几点原因.福建畜牧兽医.23(4):25
程安春,汪铭书,廖德惠等.1997.酶联免疫吸附试验检测鸭瘟抗体的研究和应用,四川农业大学学报.15(3):379-384
程安春,汪铭书,刘菲等.2004.PCR在鸭瘟临床诊断和免疫及致病机理研究中的初步应用.病毒学报.20(4):364-369
邓名义,E.C.Burgess,T.M.Yuill.1989.用反向被动血凝试验检测鸭瘟病毒.中国畜禽传染病.15(1):43-46
刁有祥,吕桂霞,郑福英,杨杰华,陈庆普,刘悦竹.2006.一种新型鸭瘟病原的分离鉴定及其特性.中国兽医学报,26(2):136-139
丁改法.2007.鸭传染性浆膜炎和鸭瘟混合感染的诊治.兽医导刊,3:39
丁巧玲,袁生,李建军,袁圣丹.2006.广东部分地区发生“新型鸭瘟”的初步诊断报告.广东畜牧兽医科技,31(4):28-32
傅涛.2004.噬菌体展示技术在病毒抗原表位筛选中的应用.国外医学病毒学分册.11(5):145-148
甘孟侯.中国禽病学.1999.中国农业出版社.107-199
郭霄峰,廖明,严英华.2002.利用PCR检测鸭肠炎病毒.中国兽医科技.32(4):13-16
郭宇飞,程安春,汪铭书.2006.鸭病毒性肠炎研究进展.中国兽医杂志.42(8):41-43
韩先杰,王君伟,布日额,李慧昕,高宏丽.2004.地高辛标记核酸探针检测鸭瘟病毒.中国兽医杂志,40(11):20-21
韩先杰,王君伟,布日额.2003.PCR检测鸭肠炎病毒的研究.中国兽医科技.33(8):10-14
何美珠.2000.大剂量鸭瘟疫苗紧急接种防治鸭瘟的体会.养禽与禽病防治.5:35
贺东生,赵善昌,苏丹萍.1998.鸭瘟病毒及其核酸的电镜观察.中国畜禽传染病.20(1):13-14
洪峰,黄引贤.1989.中国畜禽传染病.4.15-17
洪健,周雪平.2006.ICTV第八次报告的最新病毒分类系统.中国病毒学.21:84-96.
胡薛英,谷长勤,程国富等.2006.应用单克隆抗体的免疫组织化学法研究雏鸭体内鸭瘟病毒的分布.中国预防兽医学报.28(3):320-322
黄引贤,欧守杼,邝荣禄,林维庆.1980.鸭瘟病毒的研究.华南农学院学报.1(1):21-36
黄瑜,程龙飞,李文杨,苏敬良,庄向生,郭玉璞.2001.鸭疱疹病毒Ⅱ型(暂定名)的分离鉴定.中国预防兽医学报,23(2):95-98
黄瑜,苏敬良,王春风等.2003.鸭2型疱疹病毒的分子生物学依据.畜牧兽医学报,34(6):577-580
李海,毛旭虎.2007.蛋白质抗原表位研究进展.微生物学免疫学进展.35(1):54-58
李慧昕.2007.鸭肠炎病毒部分基因结构及相关功能的初步研究.中国农业科学院.博士学位论文.22-29、36-40
李玲,朱堃熹,林在尧.1988.淋巴细胞转化试验在鸭体上的应用Ⅱ鸭瘟病毒对鸭外周血淋巴细胞转化能力的影响.江苏农学院学报.9(2):5-8
林治华,吴玉章.2004.计算机辅助T细胞表位鉴定.生命的化学.24(5):397-399
刘亚刚,刘内生,岳华等.1996.安宁河流域鸭瘟、鸭霍乱综合防制研究Ⅰ鸭瘟母源抗体测定,中国兽医学报.16(2):121-124
陆承平.2005.最新动物病毒分类简介.中国病毒学.6:682-688
吕桂霞.2005.“新型鸭瘟”病理学研究.畜牧兽医杂志,4(2):41
马秀丽,宋敏训,李玉峰等.PCR用于鸭肠炎病毒诊断的研究[J].中国预防兽医学报.2005,27(5):408-411.
马秀丽,宋敏训,王莉莉等.2005.间接ELISA检测鸭瘟抗体的研究和应用.山东农业科学.1:62-64
毛秀丽.2006.N-连接蛋白质糖基化分析用微流控芯片平台和方法研究.中国科学院研究生院.博士学位论文.5-7
潘华奇.2007.鸭肠炎病毒gB基因的克隆与表达及其产物的初步应用,甘肃农业大学,硕士学位论文.15-21
齐雪峰,罗薇,杨晓燕,刘内生.2006.鸭瘟病理组织学动态观察.中国预防兽医学报,28(1):44-47
齐雪峰,程安春,汪铭书等.2007.鸭瘟病毒抗体间接ELISA检测试剂盒的研制.中国兽医科学.37(08):690-694
齐雪峰,杨晓燕,罗薇等.2003.利用PCR对鸭瘟和鸭“传染性肿头症”的鉴别诊断.四川畜牧兽医.10:21.
乔代蓉.1992.限制性内切酶分析鸭肠炎病毒DNA.四川农业大学学报.10(1):172-177
宋涌,程安春,汪铭书等.2005.聚合酶链反应(PCR)检测鸭肠炎病毒方法的建立和应用.中国兽医杂志.41(2):17-20.
孙健宏,曹殿军.2004.细胞的抗原表位研究方法.动物医学进展.25(5):18-21
索化夷,汤承,岳华等.2006.实时荧光定量TaqMan-PCR检测鸭瘟病毒方法的建立.中国预防兽医学报28(3):332-335
谭诗文,范存军,朱德荣.1990.鸭瘟弱毒疫苗早期免疫的研究Ⅰ.鸭瘟弱毒疫苗早期免疫途径和时间以及程序的研究.畜牧兽医学报.21(4):338-346
汤承,索化夷,岳华等.2006.SYBR Green Ⅰ实时定量PCR快速检测鸭瘟病毒的研究.中国动物检疫.23(9):25-27
汤承,岳华,索化夷等.2006.实时荧光定量PCR技术在鸭胚鸭瘟病毒检测中的应用.河南农业大学学报.40(3):284-287
王洪海,苏敬良,曹振等.2004.鸭肠炎病毒单克隆抗体的制备.中国兽医技.34(11):13-17.
王虎承.2005.计算机算法结合实验技术进行T细胞表位筛选.重庆工商大学学报(自然科学版).22(3):237-240
王老七,牛桂玲.2007.雏鹅感染鸭瘟的诊断及防治.安徽农业科学,35(8):2294-2296
王群.2007.禽类常见传染病的防治(1)鸭瘟.湖南农业,1:16
文明,程安春,汪铭书等.2006.鸭病毒性肠炎病毒的提纯及其结构蛋白SDS-PAGE分析.中国兽医学报.26(3)243-245
吴红专,朱道中,王星,贺东生,刘福安.1999.简介鹅的三种病毒性疾病—鹅的鸭瘟、鹅的副粘病毒病、禽流感.中国禽业导刊,16(4):46-47
谢芝勋,谢志勤,刘加波.2000.用聚合酶链反应检测鸭肠炎病毒的研究.中国兽药杂志,34(4):10-12
徐耀基,黄引贤.1992.Dot-ELISA检测鸭瘟病毒的研究.中国畜禽传染病.2:19-21
徐耀基,黄引贤.1992.应用琼脂凝胶沉淀试验检测鹅鸭病料中的鸭瘟病毒.中国畜禽传染病.65(4):30-31
徐耀基,黄引贤.1992.应用微量固相放射免疫测定法检测鸭肠炎病毒的初步研究.中国畜禽传染病.(3):35-38.
杨晓燕,罗薇,齐雪峰.2004.应用PCR检测成年鸭体内鸭瘟强毒的分布.中国兽医科技.34(1):66-69.
姚巧珍,董书青,杜红雅.2007.鸭瘟疫苗紧急接种防治鸭瘟的体会.河南畜牧兽医.28(6):41
叶联.1992.大批仔鸭病毒性肝炎和鸭瘟并发病例.广西畜牧兽医.3:34
叶润全,黄引贤.1991.中国畜禽传染病.2.35-36
殷震,刘景华.1997.动物病毒学.第二版.北京.科学出版社.988-998,1073-1081
余克伦,陆光进,贺东升.1993.鸭瘟病毒的某些分子生物学特性.病毒学报.9(3):284-286
余克伦,赵善昌,劳明波.1991.用生物素标记核酸探针检测鸭瘟病毒DNA的研究.广西农学院学报,10(3):7-14
余琳,马桂璋.2006.噬菌体展示肽库技术及其应用.国际检验医学杂志.27(8):696-671
袁桂萍,程安春,汪铭书等.2005.鸭病毒性肠炎病毒强毒在人工感染鸭体内形态结构和发生学的电镜观察.畜牧兽医学报.36(5):486-491
袁明龙,杨永纪,诸奎龙等.2001.鸭瘟抗体检测方法研究.中国预防兽医学报.23(2):140-143
岳华,杨发龙,景波,汤承,张焕容.2007.鸭瘟病毒在雏鸭体内的动态定量分布及其潜伏部位研究.中国预防兽医学报.29(9):671-675
翟中和,丁明孝,刘鹗书,宋福贞.1982.鸭瘟病毒的形态及其繁殖(复制)的电子显 微镜研究.病毒学集刊.1:63-69
张月昔,侯红霞,孙晓荣.2008.鸭瘟及鸭霍乱的鉴别与防治.水禽世界,1:13
章辉,朱荫昌.2006.B细胞表位预测的研究进展.中国血吸虫病防治杂志.18(3):236-240
郑福英,刁有祥,杨杰华等.2004.间接ELISA检测新型鸭瘟病毒抗体.中国预防兽医学报.26(3):226-230
郑福英,刁有祥.2003.应用间接免疫荧光法检测新型鸭肠炎病毒.中国兽医科技.33(10):47-50.
支轶,吴玉章.2005.万瑛.生物信息学方法在CTL表位预测中的应用.免疫学杂志.21(2):155-159
钟振东,向军俭.2005.噬菌体展示技术与肽疫苗的设计.中国生物制品学杂志.18(4):350-353
周洪.2002.T细胞表位的确定.细胞生物学杂志.24(4):196-199
Afonso CL,Tulman ER,Lu Z,Zsak L,Rock DL,Kutish GF.2001.The genome of turkey herpesvirus.J Virol.Jan;75(2):971-978
Arsenakis M,Hubenthal-Voss J,Campadelli-Fiume G,Pereira L,Roizman B,.1986.Construction and properties of a cell line constitutively expressing the herpes simplex virus glycoprotein B dependent on functional alpha 4 protein synthesis.J Virol.60(2):674-82
Bamdad T,Roostaee M H,Sadeghizadeh M,Mahboudi F,Kazemnejad A,Soleimanjahi H.2005.Immunogenicity and protective effect of a DNA construct encoding certain neutralizing epitopes of herpes simplex virus type-1 glycoprotein B.Folia Biol(Praha).51(4):109-1013
Banks T,Huo B,Kousoulas K,Spaete R,Pachl C,Pereira L.1989.A major neutralizing domain maps within the carboxyl-terminal half of the cleaved cytomegalovirus B glycoprotein.J Gen Virol.70(4):979-985
Baudet A.E.R.F.1923.Mortality in ducks in the Netherlands caused by a filterable virus:fowl plague.Tijdschr Diergeneeskd.50:455-459
Breese SS JR,Dardiri AH.1968.Electron microscopic characterization of duck plague virus.Virology.34:160-169
Breshnahan PA,Hayflick JS,Molloy SS,Thomas G.1993.Endoproteolysis of growth factors and other nonendocrine precursor proteins.In Mechanisms of Intracellular Trafficking and Processing of Proteins,225-250
Burgess EC,Ossa J,Yuiil T M.1979.Duck Plague:a carrier state in waterfowl.Avian Diseases.23(4):940-949.
Bzik DJ,Fox BA,Deluca NA,Person S.1984.Nucleotide sequence of a region of the herpes simplex virustype 1 gB glycoprotein gene:mutations affecting rate of virus entry and cell fusion.Virology.137:185-190.
Chen X,Velicer LF.1992.Expression of the Marek's disease virus homolog of herpes simplex virus glycoprotein B in Escherichia coli and its identification as B antigen.J Virol.66(7):4390-4398
Compton,T;Courtney,R J.1984.Evidence for post-translational glycosylation of a non -glycosylated precursor protein of herpes simplex virus type 1.J Virol.52(2):630-637
Cottral GE.1978. Manual of standardizl methods for veterinary microbiology ithaca and London. Cornell University press. 90-92, 148-150.
Crump,CM,Xiang Y, Thomas L, et al. 2001. PASC-1 binding to adaptors is requiredfor acidic cluster motif-mediated protein traffic. EMBO J. 20:2191-2201
Dardiri AH. 1975. Duck viral enteritis (duck plague) characteristics and immune response of the host. Am J Vet Res. 36(42):535-538.
Davison,A.J. and Scott,J.E. 1986. The complete DNA sequence of varicella-zoster virus. J. Gen. Virol. 67(9), 1759-1816
Eberle R, Black D. 1991. The simian herpesvirus SA8 homologue of the herpes simplex virus gB gene: mapping, sequencing, and comparison to the HSV gB. Arch Virol. 118(1):67-86
Edward T, Tomas B, Bo S, Stig J, Joseph C, Glorioso, Sigvard O. 1994. Localization of a functional site on herpes simplex virus type 1 glycoprotein C involved in binding to cell surface heparan sulphate. J Gen Virol. 75: 743-752.
Erik L, Maria J, Bo S ,Ulf L. 1991. Binding of herpes simplex virus to cellular heparan sulphate, an initial step in the adsorption process. J Gen Virol. 72:1131-1137.
Fitzpatrick D R, Redmond M J, Attah-Poku S K, van D, Littel-van den Hurk S, Babiuk L A, Zamb TJ. 1990. Mapping of 10 epitopes on bovine herpesvirus type 1 glycoproteins gI and gIII. Virology. 176:145-157.
Fu TM, Bonneau RH, Epler M, Tevethia MJ, Alam S, Verner K, Tevethia SS. 1996. Induction and persistence of a cytotoxic T lymphocyte (CTL) responseagainst a herpes simplex virus-specific CTL epitope expressed in a cellular protein. Virology. 222(1):269-74.
Gainer, H. 1993. Intracellular protein trafficking and proprotein processing : an overview. In Mechanisms of Intracellular Trafficking and Processing of Proteins. 1-17
Ghiasi H, Kaiwar R, Nesburn AB, Wechsler SL. 1992. Expression of herpes simplex virus type 1 glycoprotein B in insect cells. Initial analysis of its biochemical and immunological properties. Virus Res. 22(1)25-39
Griffin AM. 1991. The nucleotide sequence of the glycoprotein gB gene of infectious laryngotracheitis virus: analysis and evolutionary relationship to the homologous gene from other herpesviruses. J Gen Virol. 72 (2) 393-398
Guo PX. 1990.Characterization of the gene and an antigenic determinant of equine herpesvirus type-1 glycoprotein 14 with homology to gB-equivalent glycoproteins of other herpesviruses. Gene .87(2) 249-55
Han XJ, Wang JW, Ma B. 2006. Cloning and sequence of Glycoprotein H gene of duck plague virus. Agric.Sci.China. 5(5):397-402
Han XJ, Wang JW, 2005. Cloning of thymidine kinase gene of duck plague virus using degenerate PCR.Agric.Sci.China. 4(8):634-640
Hanke T, Graham FL, Rosenthal KL, Johnson DC. 1991. Identification of animmuno dominant cytotoxic T-lymphocyte recognition site in glycoprotein B of herpes simplex virus by using recombinant adenovirus vectors andsynthetic peptides. J Virol. 65(3): 1177-86.
Hansen WR, Brown SE, Nashold SW. 1999. Identification of duck plague virus by polymerase chainreaction. Avian Diseases.45:106-115.
Hansen WR, Nashold SW, Docherty DE, et al. 2000.Diagnosis of duck plague in waterfowl bypolymerase chain reaction[J]. Avian Diseases. 44(2): 266-274.
Hariharan K, Hariharan MJ, Zamb TJ, Krueger RJ, Srikumaran S. 1991. Bovine monoclonal anti-idiotypes induce antibodies specific for a synthetic peptide bearing a neutralizing epitope of bovine herpesvirus 1 glycoprotein gI (gB). J Immunol. 146(10):3489-95
Herman W F. 2006. The why's of Y-based motifs in alphaherpesvirus envelope proteins. Virus Research. 117:202-208
Huang Y, Babiuk LA, van Drunen Littel-van. 2005. Immunization with a bovine herpesvirus 1 glycoprotein B DNA vaccine induces cytotoxic T-lymphocyte responses in mice and cattle. J Gen Virol. 86(4):887-98
Isa G, Schelp C, Truyen U. 2003. Comparative studies with three different bovine blood sample BHV21 ELISA tests:indirect ELISA and gB-blocking-ELISA. Berl Munch Tierarztl Wochenschr. 116(526): 192-196.
Izumiya Y, Jang HK, Ono M, Mikami T. 2001. A complete genomic DNA sequence of Marek's disease virus type 2, strain HPRS24. Curr Top Microbiol Immunol. 255:191-221.
Johnson DC, Ghosh-Choudhury G, Smiley JR, Fallis L, Graham FL. 1988. Abundant expression of herpes simplex virus glycoprotein gB using an adenovirus vector. Virology. 164(1): 1-14
Kuhn JE, Kramer MD, Willenbacher W, et al. 1990. Identification of herpes simplex virus type 1 glycoproteins interacting with the cell surface. J Virol. 64(6): 2491-2497
Kulkarni DD, James PC, Sulochana S. 1998. Assessment of the immune response to duck plague vaccinations. Res Vet Sci. 64(3): 199-204.
Lam KM, Lin WQ. 1986. Antibody-mediated resistance against duck enteritis virus infection. Can J Vet Res. 50(3):380-383.
Lee WC, Fuller AO. 1993. Herpes simplex virus type 1 and pseudorabies virus bind to a common saturable receptor on Vero cells that is not heparan sulfate. J Virol. 67(9):5088-5097
Leung-Tack P, Audonnet JC, Riviere M. 1994.The complete DNA sequence and the genetic organi -zation of the short unique region (US) of the bovine herpesvirus type 1 (ST-strain). Virology. 199(2):409-21.
Li H, Liu S, Kong X. 2006. Characterization of the genes encoding UL24,TK,and gH proteins from duck enteritis virus(DEV):a proof for the classification of DEV. Virus Genes. Li, W; Minova-Foster, T J; Norton, D D; Muggeridge, M I. 2006. Identification of functional domains in herpes simplex virus 2 glycoprotein B. J Virol. 80(8):3792-3800
Li Y, van Drunen Littel-van, Babiuk LA, Liang. 1995. Characterization of cell-binding properties of bovine herpesvirus 1 glycoproteins B, C, and D: identification of a dual cell-binding function of gB. J Virol. 69 (8): 4758-4768
Loh L. 1991. Synthesis and processing of the major envelope glycoprotein of murine cytomegalovirus. Virology. 180:239-250.
Maeda K, Horimoto T, Norimine J, et al. 1992. Identification and nucleotide sequence of a gene in feline herpesvirus type 1 homologous to the herpes simplex virus gene encoding the glycoprotein B. Arch Virol. 127(1-4):387-397
McGeoch DJ, Dalrymple MA, Davison AJ, et al. 1988.The complete DNA sequence of the long unique region in the genome of herpes simplex virus type 1. J. Gen. Virol. 69 (7): 1531-1574
McLaughlin-Taylor E, Willey DE, Cantin EM, Eberle R, Moss B, Openshaw H. 1988. recombinant vaccinia virus expressing herpes simplex virus type 1 glycoprotein B induces cytotoxic T lymphocytes in mice. J Gen Virol. 69 (7): 1731-1734
Shieh MT, WuDunn D, Montgomery RI, Esko JD, Spear PG.. 1992. Cell Surface Receptors for Herpes Simplex Virus Are Heparan Sulfate Proteoglycans. The Journal of Cell Biology. 116(5):1273-1281
Miethke A, Keil GM, Weiland F, Mettenleiter TC. 1995. Unidirectional complementation between glycoprotein B homologues of pseudorabies virus and bovine herpesvirus 1 is determined by the carboxy-terminal part of the molecule. J Gen Virol. 76 (7): 1623-1635
Mikhail MZ, Oleg S. Morenkov NF, Attila B, Vadim VS, Istvan F. 1999.Distribution of B-cell epitopes on the pseudorabies virus glycoprotein B.Journal of General Virology. 80: 537-541.
Molloy SS, Anderson ED, Jean E, et al. 1999. Bi-cycling the furin pathway from TGN localization to parhogen activation and embryogenesis. Trends Cell Biol .9:28-35
Molloy SS, Thomas L, Kamibayashi C, et al. 1998. Regulation of endosome sorting by a specific PPZA isoform. J Cell Biol. 142:1399-1411
Montalvo EA, Grose C. 1987. Assembly and processing of the disulfide-linked varicella-zoster virus glycoprotein gpII(140). J.Virol. 61:2877-2884.
Mueller SN, Jones CM, Chen W, Kawaoka Y, Castrucci MR, Heath WR,Carbone FR. 2003. The early expression of glycoprotein B from herpes simplexvirus can be detected by antigen -specific CD8+ T cells. J Virol. 77(4):2445-2451.
Ross N, O'Sullivan G, Coudert F. 1996. Influence of chicken genotype on protection against Marek's disease by a herpesvirus of turkeys recombinant expressing the glycoprotein B(gB) of Marek's disease virus. Vaccine. 14(3): 187-189
Navarro D, Paz P, Pereira L.1992. Domains of herpes simplex virus 1 glycoprotein B that function in virus penetration, cell-to-cell spread, and cell fusion. Virology. 186(1):99-112
Navarro D, Qadri I, Pereira L. 1993. Transport and secretion of truncated derivatives of herpes simplex virus 1 glycoprotein B. Virology. 192(1):234-245
Neubauer A, Braun B, Brandmuller C, Kaaden OR, Osterrieder. 1997. Analysis of the contributions of the equine herpesvirus 1 glycoprotein gB homolog to virus entry and direct cell-to-cell spread. Virology. 227 (2) :281-294
Niikura M, Matsuura Y, Endoh D, Onuma M, Mikami T. 1992. Expression of the Marek's disease virus (MDV) homolog of glycoprotein B of herpes simplex virus by a recombinant baculovirus and its identification as the B antigen (gp100, gp60, gp49) of MDV. J Virol. 66(5): 2631-2638
Norais N, Tang D, Kaur S, Chamberlain SH, Masiarz FR, Burke RL, Marcus F. 1996. Disulfide bonds of herpes simplex virus type 2 glycoprotein gB. J Virol. 70(11):7379-7387
Nugent CT, Wolcott RM, Chervenak R, Jennings SR. 1994. Analysis of thecytolytic T-lymphocyte response to herpes simplex virus type 1 glycoproteinB during primary and secondary infection. J Virol. 68(11):7644-7648
Okazaki K, Fujii S, Takada A, Kida H. 2006. The amino-terminal residue of glycoprotein B is critical for neutralization of bovine herpesvirus 1. Virus Res. 115(2): 105- 111
Pereira L, Ali M, Kousoulas K, Huo B, Banks T. 1989. Domain structure of herpes simplex virus 1 glycoprotein B: neutralizing epitopes map in regions of continuous and discontinuous residues. Virology. 172(1): 11-24.
Pereira, L. 1994. Function of glycoprotein B homologues of the family herpesviridae. Infectious Agents and Disease 3:9-28
Pereira L, Ali M, Kousoulas K, Huo B, Banks T. 1989. Domain structure of herpes simplex virus 1 glycoprotein B: neutralizing epitopes map in regions of continuous and discontinuous residues. Virology. 172( 1): 11 -24
Person S, Warner SC, Bzik DJ, Debroy C, Fox BA. 1985. Expression in bacteria of gB-glycoprotein-coding sequences of Herpes simplex virus type 2. Gene. 35(3):279-287
Plummer PJ, Alefantis T, Kaplan S. 1998. Detection of duck enteritis virus by polymerase chainreaction. Avian Diseases.42: 554-564.
Poulsen D J, Keeler C. 1997. Characterization of the assembly and processing of infectious laryngotracheitis virus glycoprotein B. J Gen Virol. 78 (11):2945-2951
Proctor SJ. 1975. Pathogenesis of digestive tract lesions in duck plague.Vet Pathol. 12: 349-361.
Proctor SJ. Pathogenesis of duck plague in the bursa of Fabricius, thymus and spleen. American. Journal of Veterinary Research. 37:427-431
Robbins AK, Dorney DJ, Wathen MW, Whealy ME, Gold C, Watson RJ, Holland LE, Weed SD, Levine M, Glorioso JC, Enquist LW. 1987. The pseudorabies virus gll gene is closely related to the gB glycoprotein gene of herpessimplex virus. J. Virol. 61:2691-2701.
Robert N, James B, Keith Y, Robert M. 1995. pET TRX Fusion System for Increased Solubility of Proteins Expressed in E. coli. inNovation. 3:6-8
Ros C, Belak S. 2002. Characterization of the glycoprotein B gene from ruminant alphaherpesviruses. Virus Genes. 24(2):99-105
Ross LJ, Sanderson M, Scott SD, Binns, MM, Doel T, Milne B. 1989. Nucleotide sequence and characterization of the Marek's disease virus homologue of glycoprotein B of herpes simplex virus. J Gen Virol. 70 (7): 1789-1804
Shawky S, Sandhu T, Shivaprasad HL. 2000 .Pathogenicity of a low-virulence duck virus enteritisisolate with apparent immunosuppressive ability. Avian Diseases. 44(3): 590-599.
Shawky S, Schat KA. 2002. Latency sites and reactivation of duck enteritis virus[J]. Avian Diseases. 46(2): 308-313.
Shawky S. 2000.Target cells for duck enteritis virus in lymphoid organs. Avian Pathology. 29(6): 609-616
Smith IL, Hardwicke, MA, Sandri-Goldin, RM. 1992. Evidence that the herpes simplex virus immediate early protein ICP27 acts post-transcriptionally during infection to regulate gene expression. Virology. 186(1):74-86
Thomas CH, Nancy K, Susan H. 2000. Cytoplasmic Domain Signal Sequences That Mediate Transport of Varicella-Zoster Virus gB from the Endoplasmic Reticulum to the Golgi.journal of virology. 74(20):9421-9430
Thureen DR, Keeler CLJ. 2006. Psittacid herpesvirus 1 and infectious laryngotracheitis virus: Comparative genome sequence analysis of two avian alphaherpesviruses. J Virol. 80(16):7863-7872.
Toth TE. 1971. Active immunization of White Pekin ducks against Duck virus enteritis (duck plague) with modified-live-virus vaccine: serologic and immunologic response of breeder ducks. Am J Vet Res. 32(1):75-81
Vickery KGX. 1997. Antigen-specific blastogenesis assays for duck hepatitis B virus using duck peripheral blood and splenic mononuclear cells. Veterinary Immunology and Immunopathology. 59:349-358
Vickery K,Cossart Y, Dixon R. 1999. Comparison of the kinetics of the specific cellular immune response to duck hepatitis B virus in infected and immune ducks. Veterinary microbiology. 68:157-169
Vlcek C, Benes V, Lu Z, et al. 1995. Nucleotide sequence analysis of a 30-kb region of the bovine herpesvirus 1 genome which exhibits a colinear gene arrangement with the UL21 to UL4 genes of herpes simplex virus. J. Virol. 210 (1):100-108
Wan L, Molloy SS, Thomas L, et al. 1998. PACS-1 defines a novel gene family of cytosolic sorting proteins required for trans-Golgi network localization. Cell. 94:205-216
Weiqing L, Lam KM, Clark WE. 1984. Active and passive immunization of ducks against duck viral enteritis. Avian disease. 28(4):968-973
Welschinger R, Cossart Y, Pouliopoulos J, Dixon R, Vickery K, 2006. Identification of T-cell epitopes associated with immunity within the surface protein of duck hepatitis B virus Journal of Viral Hepatitis. 13:515-522
Whealy ME, Card JP, Meade RP, et al. 1991. Effect of brefeldin A on alphaherpesvirus membrane protein glycosylation and virus egress. J. Virol 65:1066-1081.
Whealy AE, Robbins AK, Enquist LW. 1990. The export pathway of the pseudorabies virus gB homolog gII involves oligomer formation in the endoplasmic reticulum and protease processing in the Golgi apparatus. J Virol. 64:1946-1955.
Whitbeck JC, Bello LJ, Lawrence WC. 1988. Comparison of the bovine herpesvirus 1 gI gene and the herpes simplex virus type 1 gB gene. J Virol. 62(9):3319-3327
Yang FL, Jia WX, Yue H, et al. 2005.Development of quantitative real-time polymerase virus chain reaction for duck enteritis. Avian Disease. 49: 397-400.