我国O型泛亚1系口蹄疫病毒表型差异的分子基础
|
摘要
本论文对3株不同宿主来源的O型泛亚1系口蹄疫病毒（foot-and-mouth disease virus, FMDV）进行了分子变异和表型差异研究。经全基因组序列测定、核苷酸/氨基酸同源性比较分析，结果表明：1999年2株牛源分离毒（O/NX/CHA/99和O/HK/HB/CHA/99）和2008年1株猪源分离毒（O/ZheJ/CHA/8/2008）划分为PanAsia-1a和PanAsia-1b两个亚系。VP2的136（E-F环）、175位（G-H环）和214位（C-端），VP3的174位（G-H环），VP1的142、152、153位（G-H环）和199位（C-端）为这两个亚系的特征性氨基酸残基位点。BHK-21细胞、乳鼠毒力测定以及蚀斑形成和/或抑制试验结果显示：O/NX/CHA/99的毒力（TCID_50=6.625, LD_50=7.33）明显低于O/HK/HB/CHA/99（TCID_50=7.75, LD_50=8.33）和O/ZheJ/CHA/8/2008（TCID_50=7.875, LD50=8.5）；O/NX/CHA/99呈小而密致的蚀斑，O/HK/HB/CHA/99呈大而浑浊的蚀斑，O/ZheJ/CHA/8/2008呈大而清晰的蚀斑；肝素钠可抑制O/NX/CHA/99的成斑能力，但完全不能抑制O/HK/HB/CHA/99和O/ZheJ/CHA/8/2008的呈斑水平。有趣的是，在CHO-K1细胞上，O/NX/CHA/99和O/HK/HB/CHA/99长斑，肝素钠可以完全抑制它们的成斑能力，而O/ZheJ/CHA/8/2008不长斑。提示O/NX/CHA/99具有与硫酸乙酰肝素（heparan sulfate, HS）受体的高亲和力，O/HK/HB/CHA/99可以利用HS受体，O/ZheJ/CHA/8/2008不能利用HS受体。
为了寻找决定O型泛亚1系FMDV蚀斑大小、利用HS受体的关键性氨基酸位点，以改造的不能利用HS受体的FMDV OZK93全长cDNA质粒作为基础骨架，利用定点突变和盒式替换策略，分别构建了12种VP3基因不同的嵌合全长cDNA质粒、3种不同VP1基因的嵌合全长cDNA质粒和22种VP0基因不同的嵌合全长cDNA质粒。经体内转录，分别拯救了9、3、13种VP3、VP1、VP0基因不同的嵌合病毒。BHK-21、CHO-K1和HS缺陷型细胞（pgsA745和pgsB618）的蚀斑形成和/或抑制试验结果显示，1）O/NX/CHA/99VP1基因的嵌合病毒不仅在BHK-21细胞上呈大小迥异的混合斑，肝素钠只能部分抑制其呈斑水平，而且在CHO-K1细胞上长斑，可被肝素钠完全抑制，同时在pgsA745和pgsB618上的成斑能力明显被抑制，说明它能够同时利用整联蛋白和HS受体；VP1基因区段是决定O/NX/CHA/99利用HS受体的功能区域，从序列比对和氨基酸性质推测K83可能是关键性氨基酸。VP2的Q80是决定O/HK/HB/CHA/99利用HS受体的关键性氨基酸。2）VP2的79位为H（碱性）的嵌合基因工程毒与该位点为Y（中性）的嵌合基因工程毒相比，蚀斑趋于变小；136位为E（酸性）的嵌合基因工程毒与该位点为G（中性）的嵌合基因工程毒相比，蚀斑趋于变大。VP4上第8位氨基酸残基S（亲水、极性）→A（疏水、非极性）对病毒与肝素的亲和力具有一定的辅助作用。
综合以上研究结果得出：1）猪源O型泛亚1系FMDV不能利用HS受体，而牛源O型泛亚1系FMDV能够利用HS受体；病毒与HS受体的结合效力与其在BHK-21细胞上的蚀斑大小和毒力有关。2）FMDV蚀斑形态差异与其衣壳蛋白上关键性作用位点的氨基酸残基性质改变有关。
Foot-and-mouth disease (FMD), one of serious animal diseases of pigs, cattle, sheep and goats, iscaused by foot-and-mouth disease virus (FMDV). Serotype O is the most prevalent serotype comparedto the other serotypes (A, Asia1) of FMDV in China. To understand genetic variations and phenotypicdifferences at the molecular level, in this study, we selected three virus isolates of the PanAsia-1strainfrom bovine and porcine based on sequence determination and comparison of nucleotides and/ordeduced amino acids homology. The analyses of the complete genome sequences showed: Two bovineisolates in1999and one swine isolate in2008were subdivided into PanAsia-1a and PanAsia-1bsubstrains of the PanAsia-1strain of Middle-East South-Asia (ME-SA) topotype, respectively. Instructural protein, the distinct amino acid residues located at positions136(E-F loop),175(G-H loop)and214(C-terminal) of VP2,174(G-H loop) of VP3,142,152and153(G-H loop) and199(C-terminal) of VP1. The virulence to BHK-21cells and suckling mice of FMDV O/NX/CHA/99(TCID_50=6.625, LD_50=7.33) was lower than O/HK/HB/CHA/99(TCID_50=7.75, LD_50=8.33) andO/ZheJ/CHA/8/2008(TCID_50=7.875, LD_50=8.5). These three viruses differed from their plaquephenotypes on BHK-21cells. O/NX/CHA/99formed small and compact plaques, O/HK/HB/CHA/99formed large and turbid plaques, and O/ZheJ/CHA/8/2008formed large and clear plaques.Concentrations of heparin sodium from0to1mg/ml greatly decreased the plaque formation ofO/NX/CHA/99, but had no effect of O/HK/HB/CHA/99and O/ZheJ/CHA/8/2008on BHK-21cells.Interestingly, O/NX/CHA/99and O/HK/HB/CHA/99produced plaques on CHO-K1cells, and plaqueformation could be inhibited by addition of heparin sodium. However, O/ZheJ/CHA/8/2008can notproduce plaques on CHO-K1cells. These results indicate that O/NX/CHA/99has a high affinity toheparan sulfate (HS) receptor, O/HK/HB/CHA/99can utilize HS as cellular receptors, whereasO/ZheJ/CHA/8/2008propagated in cultured cells independent of HS binding.
To identify the key amino acid residues determing the plaque size and the binding ability to HSreceptor, we constructed chimeric full-length cDNA clones and generated chimeric FMDV mutants asdescribed. By using site-directed mutagenesis and exchange-cassette strategy,9,3, and12cDNAsencoding the VP3, VP1, and VP0capsid proteins of an infectious genome-length cDNA copy of HSutilizing-deficient OZK93(rOZK, hs~-) were replaced with appropriate variants and cDNAscorresponding to PanAsia-1strains, producing chimeric FMDV mutants for providing additional insightinto the close relationship among virulence, plaque phenotypes, alternative mechanisms to receptorsusage and amino acid substitutions in capsid proteins of FMDV.25different DNA fragment patternswere designed and stably maintained the sequence of the original rescued viruses, based on T7RNApolymerase in vivo transcription system, respectively. The plaque forming and/or inhibiting assays onBHK-21, CHO-K1, pgsA745(xylosyltransferase I-deficient) and pgsB618(galactosyltransferaseI-deficient) confirmed:1) Chimeric virus containing capsid protein VP1of O/NX/CHA/99formed smalland large blend plaques on BHK-21cells, produced plaques on CHO-K1cells, and grew poorly onpgsA745and pgsB618. Plaque formation was inhibited on CHO-K1cells but had moderate effect on BHK-21cells when heparin sodium was added to diluted virus suspension. The results demonstratedthat the adaptation to HS receptor by this virus appears not to compromise their ability to make use ofintegrins. The capsid protein VP1of O/NX/CHA/99was identified as the critical determinant relevantto FMDV-HS receptor interaction. Residue K83in VP1potentially functions as one of critical aminoacids for HS receptor binding in cultured cells. Q80L change within B-C loop of VP2can lead to loss ofHS receptor binding ability of O/HK/HB/CHA/99.2) Residues79and136were measured in VP2of thePanAsia-1strain as the critical amino acid determinants of plaque phenotypes, which have an influenceon the average size of plaque production.2079H (basic) and2136G (neutral) facilitated the formation ofsmall plaques of rescued FMDV variants.2079Y (neutral) and2136E (acidic) enlarged the plaquesformed by other FMDV mutants. In addition, a single amino acid replacement at residue8(Ser→Ala,hydrophilic, polar→hydrophobic, nonpolar) of VP4can play a helper role in heparin recognition.
In conclusion, these studies confirmed the ability of FMDV to bind different classes of receptors isrelated to host tropism. The bovine isolates can infect cells via HS-mediated adsorption, while FMDVisolated from porcine targets susceptible cells via integrins interaction. The efficiency of FMDV-HSreceptor interaction is associated with phenotypic differences on BHK-21cells and virulence. Mutationsat critical amino acid sites (basic-neutral-acidic) in capsid proteins might generate change of plaquephenotypes, of the PanAsia-1strain of FMDV serotype O.
为了寻找决定O型泛亚1系FMDV蚀斑大小、利用HS受体的关键性氨基酸位点，以改造的不能利用HS受体的FMDV OZK93全长cDNA质粒作为基础骨架，利用定点突变和盒式替换策略，分别构建了12种VP3基因不同的嵌合全长cDNA质粒、3种不同VP1基因的嵌合全长cDNA质粒和22种VP0基因不同的嵌合全长cDNA质粒。经体内转录，分别拯救了9、3、13种VP3、VP1、VP0基因不同的嵌合病毒。BHK-21、CHO-K1和HS缺陷型细胞（pgsA745和pgsB618）的蚀斑形成和/或抑制试验结果显示，1）O/NX/CHA/99VP1基因的嵌合病毒不仅在BHK-21细胞上呈大小迥异的混合斑，肝素钠只能部分抑制其呈斑水平，而且在CHO-K1细胞上长斑，可被肝素钠完全抑制，同时在pgsA745和pgsB618上的成斑能力明显被抑制，说明它能够同时利用整联蛋白和HS受体；VP1基因区段是决定O/NX/CHA/99利用HS受体的功能区域，从序列比对和氨基酸性质推测K83可能是关键性氨基酸。VP2的Q80是决定O/HK/HB/CHA/99利用HS受体的关键性氨基酸。2）VP2的79位为H（碱性）的嵌合基因工程毒与该位点为Y（中性）的嵌合基因工程毒相比，蚀斑趋于变小；136位为E（酸性）的嵌合基因工程毒与该位点为G（中性）的嵌合基因工程毒相比，蚀斑趋于变大。VP4上第8位氨基酸残基S（亲水、极性）→A（疏水、非极性）对病毒与肝素的亲和力具有一定的辅助作用。
综合以上研究结果得出：1）猪源O型泛亚1系FMDV不能利用HS受体，而牛源O型泛亚1系FMDV能够利用HS受体；病毒与HS受体的结合效力与其在BHK-21细胞上的蚀斑大小和毒力有关。2）FMDV蚀斑形态差异与其衣壳蛋白上关键性作用位点的氨基酸残基性质改变有关。
Foot-and-mouth disease (FMD), one of serious animal diseases of pigs, cattle, sheep and goats, iscaused by foot-and-mouth disease virus (FMDV). Serotype O is the most prevalent serotype comparedto the other serotypes (A, Asia1) of FMDV in China. To understand genetic variations and phenotypicdifferences at the molecular level, in this study, we selected three virus isolates of the PanAsia-1strainfrom bovine and porcine based on sequence determination and comparison of nucleotides and/ordeduced amino acids homology. The analyses of the complete genome sequences showed: Two bovineisolates in1999and one swine isolate in2008were subdivided into PanAsia-1a and PanAsia-1bsubstrains of the PanAsia-1strain of Middle-East South-Asia (ME-SA) topotype, respectively. Instructural protein, the distinct amino acid residues located at positions136(E-F loop),175(G-H loop)and214(C-terminal) of VP2,174(G-H loop) of VP3,142,152and153(G-H loop) and199(C-terminal) of VP1. The virulence to BHK-21cells and suckling mice of FMDV O/NX/CHA/99(TCID_50=6.625, LD_50=7.33) was lower than O/HK/HB/CHA/99(TCID_50=7.75, LD_50=8.33) andO/ZheJ/CHA/8/2008(TCID_50=7.875, LD_50=8.5). These three viruses differed from their plaquephenotypes on BHK-21cells. O/NX/CHA/99formed small and compact plaques, O/HK/HB/CHA/99formed large and turbid plaques, and O/ZheJ/CHA/8/2008formed large and clear plaques.Concentrations of heparin sodium from0to1mg/ml greatly decreased the plaque formation ofO/NX/CHA/99, but had no effect of O/HK/HB/CHA/99and O/ZheJ/CHA/8/2008on BHK-21cells.Interestingly, O/NX/CHA/99and O/HK/HB/CHA/99produced plaques on CHO-K1cells, and plaqueformation could be inhibited by addition of heparin sodium. However, O/ZheJ/CHA/8/2008can notproduce plaques on CHO-K1cells. These results indicate that O/NX/CHA/99has a high affinity toheparan sulfate (HS) receptor, O/HK/HB/CHA/99can utilize HS as cellular receptors, whereasO/ZheJ/CHA/8/2008propagated in cultured cells independent of HS binding.
To identify the key amino acid residues determing the plaque size and the binding ability to HSreceptor, we constructed chimeric full-length cDNA clones and generated chimeric FMDV mutants asdescribed. By using site-directed mutagenesis and exchange-cassette strategy,9,3, and12cDNAsencoding the VP3, VP1, and VP0capsid proteins of an infectious genome-length cDNA copy of HSutilizing-deficient OZK93(rOZK, hs~-) were replaced with appropriate variants and cDNAscorresponding to PanAsia-1strains, producing chimeric FMDV mutants for providing additional insightinto the close relationship among virulence, plaque phenotypes, alternative mechanisms to receptorsusage and amino acid substitutions in capsid proteins of FMDV.25different DNA fragment patternswere designed and stably maintained the sequence of the original rescued viruses, based on T7RNApolymerase in vivo transcription system, respectively. The plaque forming and/or inhibiting assays onBHK-21, CHO-K1, pgsA745(xylosyltransferase I-deficient) and pgsB618(galactosyltransferaseI-deficient) confirmed:1) Chimeric virus containing capsid protein VP1of O/NX/CHA/99formed smalland large blend plaques on BHK-21cells, produced plaques on CHO-K1cells, and grew poorly onpgsA745and pgsB618. Plaque formation was inhibited on CHO-K1cells but had moderate effect on BHK-21cells when heparin sodium was added to diluted virus suspension. The results demonstratedthat the adaptation to HS receptor by this virus appears not to compromise their ability to make use ofintegrins. The capsid protein VP1of O/NX/CHA/99was identified as the critical determinant relevantto FMDV-HS receptor interaction. Residue K83in VP1potentially functions as one of critical aminoacids for HS receptor binding in cultured cells. Q80L change within B-C loop of VP2can lead to loss ofHS receptor binding ability of O/HK/HB/CHA/99.2) Residues79and136were measured in VP2of thePanAsia-1strain as the critical amino acid determinants of plaque phenotypes, which have an influenceon the average size of plaque production.2079H (basic) and2136G (neutral) facilitated the formation ofsmall plaques of rescued FMDV variants.2079Y (neutral) and2136E (acidic) enlarged the plaquesformed by other FMDV mutants. In addition, a single amino acid replacement at residue8(Ser→Ala,hydrophilic, polar→hydrophobic, nonpolar) of VP4can play a helper role in heparin recognition.
In conclusion, these studies confirmed the ability of FMDV to bind different classes of receptors isrelated to host tropism. The bovine isolates can infect cells via HS-mediated adsorption, while FMDVisolated from porcine targets susceptible cells via integrins interaction. The efficiency of FMDV-HSreceptor interaction is associated with phenotypic differences on BHK-21cells and virulence. Mutationsat critical amino acid sites (basic-neutral-acidic) in capsid proteins might generate change of plaquephenotypes, of the PanAsia-1strain of FMDV serotype O.
引文
1.洪健,周雪平. ICTV第八次报告的最新病毒分类系统.中国病毒学,2006,21(1):8496.
2.刘在新.口蹄疫病毒基因组及其编码蛋白一级结构研究.[博士学位论文].北京:中国农业科学院,2002.
3.刘增胜,谷继承,陈伟生.中国畜牧业年鉴.北京:中国农业出版社,2007.
4.浦华.动物疾病防控的经济学研究综述.农业经济问题,2006,6:6164.
5.谢庆阁.主编.口蹄疫.北京:中国农业出版社,2004.
6.熊忠良,徐涤平.我国重大动物疫病流行现状及其防控策略.湖北畜牧兽医,2007,12:2224.
7.徐耀先,周晓峰,刘立德.主编.分子病毒学.武汉:湖北科学技术出版社,2000.
8.殷震,刘景华.主编.动物病毒学.第二版.北京:科学出版社,1997.
9. Abdul-Hamid N. F., Hussein N. M., Wadsworth J., Radford A. D., Knowles N. J., King D. P.Phylogeography of foot-and-mouth disease virus types O and A in Malaysia and surroundingcountries. Infect Genet Evol.,2011,11(2):320328.
10. Acharya R., Fry E., Stuart D., Fox G., Rowlands D., Brown F. The three-dimensional structure offoot-and-mouth disease virus at2.9resolution. Nature,1989,337(6209):709–716.
11. Aggarwal N., Barnett P. V. Antigenic sites of foot-and-mouth disease virus (FMDV): an analysis ofthe specificities of anti-FMDV antibodies after vaccination of naturally susceptible host species. JGen Virol.,2002,83(Pt4):775–782.
12. Ahn I., Bae S. E., Son H. S. Comparative study of codon substitution patterns in foot-and-mouthdisease virus (serotype O). Exp Mol Med.,2011,43(10):587595.
13. Ansell D. M., Samuel A. R., Carpenter W. C., Knowles N. J. Genetic relationships betweenfoot-and-mouth disease type Asia1viruses. Epidemiol Infect.,1994,112(1):213224.
14. Arias A., Agudo R., Ferrer-Orta C., Pérez-Luque R., Airaksinen A., Brocchi E., Domingo E.,Verdaguer N., Escarmís C. Mutant viral polymerase in the transition of virus to error catastropheidentifies a critical site for RNA binding. J Mol Biol.,2005,353(5):1021–1032.
15. Arias A., Perales C., Escarmís C., Domingo E. Deletion mutants of VPg reveal new cytopathologydeterminants in a picornavirus. PLoS One,2010,5(5): e10735.
16. Arzt J., Pacheco J. M., Rodriguez L. L. The early pathogenesis of foot-and-mouth disease in cattleafter aerosol inoculation. Identification of the nasopharynx as the primary site of infection. VetPathol.,2010,47(6):1048–1063.
17. Asokan A., Hamra J. B., Govindasamy L., Agbandje-McKenna M., Samulski R. J.Adeno-associated virus type2contains an integrin alpha5beta1binding domain essential for viralcell entry. J Virol.,2006,80(18):8961–8969.
18. Ayelet G., Mahapatra M., Gelaye E., Egziabher B. G., Rufeal T., Sahle M., Ferris N. P., WadsworthJ., Hutchings G. H., Knowles N. J. Genetic characterization of foot-and-mouth disease viruses,Ethiopia,1981-2007. Emerg Infect Dis.,2009,15(9):14091417.
19. Bao H. F., Li D., Sun P., Zhou Q., Hu J., Bai X. W., Fu Y. F., Lu Z. J., Liu Z. X. The infectivity andpathogenicity of a foot-and-mouth disease virus persistent infection strain fromoesophageal-pharyngeal fluid of a Chinese cattle in2010. Virol J.,2011,8:536.
20. Baranowski E., Molina N., Nú ez J. I., Sobrino F., Sáiz M. Recovery of infectious foot-and-mouthdisease virus from suckling mice after direct inoculation with in vitro-transcribed RNA. J Virol.,2003,77(20):1129011295.
21. Baranowski E., Ruiz-Jarabo C. M., Sevilla N., Andreu D., Beck E., Domingo E. Cell recognitionby foot-and-mouth disease virus that lacks the RGD integrin-binding motif: flexibility inaphthovirus receptor usage. J Virol.,2000,74(4):1641–1647.
22. Baranowski E., Sevilla N., Verdaguer N., Ruiz-Jarabo C. M., Beck E., Domingo E. Multiplevirulence determinants of foot-and-mouth disease virus in cell culture. J Virol.,1998,72(8):6362–6372.
23. Barnett P. V., Ouldridge E. J., Rowlands D. J., Brown F., Parry N. R. Neutralizing epitopes of typeO foot-and-mouth disease virus. I. Identification and characterization of three functionallyindependent, conformational sites. J Gen Virol.,1989,70(Pt6):1483–1491.
24. Baxt B., Mason P. W. Foot-and-mouth disease virus undergoes restricted replication in macrophagecell cultures following Fc receptor-mediated adsorption. Virology,1995,207(2):503–509.
25. Bayry J., Tough D. F. Interaction of foot-and-mouth disease virus with dendritic cells. TrendsMicrobiol.,2006,14(8):346–347.
26. Beard C. W., Mason P. W. Genetic determinants of altered virulence of Taiwanese foot-and-mouthdisease virus. J Virol.,2000,74(2):987991.
27. Belnap D. M., Filman D. J., Trus B. L., Cheng N., Booy E. P., Conway J. E., Curry S., Hiremath C.N., Tsang S. K., Steven A. C., Hogle J. M. Molecular tectonic model of virus structure transitions:the putative cell entry states of poliovirus. J Virol.,2000,74:13421354.
28. Belsham G. J., Jamal S. M., Tj rneh j K., B tner A. Rescue of foot-and-mouth disease viruses thatare pathogenic for cattle from preserved viral RNA samples. PLoS One,2011,6(1): e14621.
29. Berinstein A., Roivainen M., Hovi T., Mason P. W., Baxt B. Antibodies to the vitronectin receptor(integrin alpha V beta3) inhibit binding and infection of foot-and-mouth disease virus to culturedcells. J Virol.,1995,69(4):2664–2666.
30. Berryman S., Clark S., Monaghan P., Jackson T. Early events in integrin alphavbeta6-mediated cellentry of foot-and-mouth disease virus. J Virol.,2005,79(13):8519–8534.
31. Bittle J. L., Houghen R. A., Alexander H., Shinnick T. M., Sutcliffe J. G., Lerner R. A., RowlandsD. J., Brown F. Protection against foot-and-mouth disease by immunization with a chemicallysynthesized peptide predicted from the viral nucleotide sequence. Nature,1982,298(5869):3033.
32. Bolwell C., Brown A. L., Barnett P. V., Campbell R. O., Clarke B. E., Parry N. R., Ouldridge E. J.,Brown F., Rowlands D. J. Host cell selection of antigenic variants of foot-and-mouth disease virus.J Gen Virol.,1989,70(Pt1):4557.
33. Borca M. V., Pacheco J. M., Holinka L. G., Carrillo C., Hartwig E., Garriga D., Kramer E.,Rodriguez L., Piccone M. E. Role of arginine-56within the structural protein VP3offoot-and-mouth disease virus (FMDV) O1Campos in virus virulence. Virology,2012,422(1):37–45.
34. B tner A., Kakker N. K., Barbezange C., Berryman S., Jackson T., Belsham G. J. Capsid proteinsfrom field strains of foot-and-mouth disease virus confer a pathogenic phenotype in cattle on anattenuated, cell-culture-adapted virus. J Gen Virol.,2011,92(Pt5):1141–1151.
35. Brown F. The history of research in foot-and-mouth disease. Virus Res.,2003,91:37.
36. Brown J. K., McAleese S. M., Thornton E. M., Pate J. A., Schock A., Macrae A. I., Scott P. R.,Miller H. R., Collie D. D. Integrin-alphavbeta6, a putative receptor for foot-and-mouth diseasevirus, is constitutively expressed in ruminant airways. J Histochem Cytochem.,2006,54(7):807–816.
37. Burman A., Clark S., Abrescia N. G., Fry E. E., Stuart D. I., Jackson T. Specificity of the VP1GHloop of foot-and-mouth disease virus for alphav integrins. J Virol.,2006,80(19):9798–9810.
38. Camper N., Byrne T., Burden R. E., Lowry J., Gray B., Johnston J. A., Migaud M. E., Olwill S. A.,Buick R. J., Scott C. J. Stable expression and purification of a functional processed Fab' fragmentfrom a single nascent polypeptide in CHO cells expressing the mCAT-1retroviral receptor. JImmunol Methods,2011,372(1-2):30–41.
39. Cao Y., Lu Z., Sun P., Fu Y., Tian F., Hao X., Bao H., Liu X., Liu Z. A pseudotype baculovirusexpressing the capsid protein of foot-and-mouth disease virus and a T-cell immunogen showsenhanced immunogenicity in mice. Virol J.,2011,8:77.
40. Carrillo C., Lu Z., Borca M. V., Vagnozzi A., Kutish G. F., Rock D. L. Genetic and phenotypicvariation of foot-and-mouth disease virus during serial passages in a natural host. J Virol.,2007,81(20):11341–11351.
41. Carrillo C., Tulman E. R., Delhon G., Lu Z., Carreno A., Vagnozzi A., Kutish G. F., Rock D. L.Comparative genomics of foot-and-mouth disease virus. J Virol.,2005,79(10):64876504.
42. Chang Y., Zheng H., Shang Y., Jin Y., Wang G., Shen X., Liu X. Recovery of infectiousfoot-and-mouth disease virus from full-length genomic cDNA clones using an RNA polymerase Isystem. Acta Biochim Biophys Sin.,2009,41(12):9981007.
43. Charpentier N., Dávila M., Domingo E., Escarmís C. Long-term, large-population passage ofaphthovirus can generate and amplify defective noninterfering particles deleted in the leaderprotease gene. Virology,1996,223(1):1018.
44. Chen J. D., Zhao M. Q., Hui K. H., Leung F. C. Molecular characterization of foot-and-mouthdisease virus in Hong Kong during2001-2002. Virus Genes,2006,32(2):139143.
45. Cottam E. M., Haydon D. T., Paton D. J., Gloster J., Wilesmith J. W., Ferris N. P., Hutchings G. H.,King D. P. Molecular epidemiology of the foot-and-mouth disease virus outbreak in the UnitedKingdom in2001. J Virol.,2006,80(22):1127411282.
46. Cottam E. M., Wadsworth J., Shaw A. E., Rowlands R. J., Goatley L., Maan S., Maan N. S.,Mertens P. P., Ebert K., Li Y., Ryan E. D., Juleff N., Ferris N. P., Wilesmith J. W., Haydon D. T.,King D. P., Paton D. J., Knowles N. J. Transmission pathways of foot-and-mouth disease virus inthe United Kingdom in2007. PLoS Pathog.,2008,4(4): e1000050.
47. Cowan K. M. Immunochemical studies of foot-and-mouth disease. V. Antigenic variants of virusdemonstrated by immunodiffusion analyses with19S but not7S antibodies. J Exp Med.,1969,129(2):333350.
48. Cowan K. M., Erol N., Whiteland A. P. Heterogeneity of type Asia1foot-and-mouth disease virusand BHK-21cells and the relationship to vaccine preparation. O. I. E Bull.,1974,81:12711298.
49. Crowther J. R., Farias S., Carpenter W. C., Samuel A. R. Identification of a fifth neutralizable siteon type O foot-and-mouth disease virus following characterization of single and quintuplemonoclonal antibody escape mutants. J Gen Virol.,1993,74:17471953.
50. Curry S., Abrams C. C., Fry E., Crowther J. C., Belsham G. J., Stuart D. I., King A. M. Viral RNAmodulates the acid sensitivity of foot-and-mouth disease virus capsids. J Virol.,1995,69(1):430–438.
51. Delputte P. L., Costers S., Nauwynck H. J. Analysis of porcine reproductive and respiratorysyndrome virus attachment and internalization: distinctive roles for heparan sulphate andsialoadhesin. J Gen Virol.,2005,86(Pt5):1441–1445.
52. Dicara D., Burman A., Clark S., Berryman S., Howard M. J., Hart I. R., Marshall J. F., Jackson T.Foot-and-mouth disease virus forms a highly stable, EDTA-resistant complex with its principalreceptor, integrin alphavbeta6: implications for infectiousness. J Virol.,2008,82(3):1537–1546.
53. Díez J., Dávila M., Escarmís C., Mateu M. G., Dominguez J., Pérez J. J., Giralt E., Melero J. A.,Domingo E. Unique amino acid substitutions in the capsid proteins of foot-and-mouth disease virusfrom a persistent infection in cell culture. J Virol.,1990,64(11):5519–5528.
54. Domingo E., Escarmís C., Baranowski E., Ruiz-Jarabo C. M., Carrillo E., Nú ez J. I., Sobrino F.Evolution of foot-and-mouth disease virus. Virus Res.,2003,91(1):47–63.
55. Domingo E., Escarmis C., Martinez M. A., Martinez-Salas E., Mateu M. G. Foot-and-mouthdisease virus populations are quasispecies. Curr Top Microbiol Immunol.,1992,176:3347.
56. Domingo E., Mateu M. C., Martinez M. A., Dopazo J., Moya A., Sobrino F. Genetic variability andantigenic diversity of foot-and-mouth disease virus. In: Kurstak E., Marusyk R. G., Murphy F. A.,van Regenmortel M. H. V. eds. Applied virology research, Vol2, Virus variation and epidemiology.New York: Plenum Publishing Corp.,1990.233266.
57. Domingo E., Ruiz-Jarabo C. M., Sierra S., Arias A., Pariente N., Baranowski E., Escarmis C.Emergence and selection of RNA virus variants: memory and extinction. Virus Res.,2002,82(1-2):3944.
58. Drake J. W. Rates of spontaneous mutation among RNA viruses. Proc Natl Acad Sci U S A,1993,90(9):41714175.
59. Drake J. W., Holland J. J. Mutation rates among RNA viruses. Proc Natl Acad Sci U S A,1999,96(24):1391013913.
60. Du J., Chang H., Cong G., Shao J., Lin T., Shang Y., Liu Z., Liu X., Cai X., Xie Q. Completenucleotide sequence of a Chinese serotype Asia1vaccine strain of foot-and-mouth disease virus.Virus Genes,2007,35(3):635642.
61. Du J., Chang H., Gao S., Xue S., Cong G., Shao J., Lin T., Liu Z., Liu X., Cai X. Molecularcharacterization and expression analysis of porcine integrins alphavbeta3, alphavbeta6andalphavbeta8that are potentially involved in FMDV infection. Mol Cell Probes,2010,24(5):256–265.
62. Dunn C. S., Donaldson A. I. Natural adaption to pigs of a Taiwanese isolate of foot-and-mouthdisease virus. Vet Rec.,1997,141(7):174175.
63. Duque H., Baxt B. Foot-and-mouth disease virus receptors: comparison of bovine alpha(V)integrin utilization by type A and O viruses. J Virol.,2003,77(4):2500–2511.
64. Duque H., LaRocco M., Golde W. T., Baxt B. Interactions of foot-and-mouth disease virus withsoluble bovine alphaVbeta3and alphaVbeta6integrins. J Virol.,2004,78(18):9773–9781.
65. Durand S., Murphy C., Zhang Z., Alexandersen S. Epithelial distribution and replication offoot-and-mouth disease virus RNA in infected pigs. J Comp Pathol.,2008,139(2-3):86–96.
66. Edwards J., Morrissy C. China-EU training to improve prevention and control of foot-and-mouthdisease.812August2011, P. R. China: Lanzhou.2011.
67. Ellard F. M., Drew J., Blakemore W. E., Stuart D. I., King A. M. Evidence for the role of His-142of protein1C in the acid-induced disassembly of foot-and-mouth disease virus capsids. J GenVirol.,1999,80(Pt8):1911–1918.
68. Escarmís C., Carrillo E. C., Ferrer M., Arriaza J. F., Lopez N., Tami C., Verdaguer N., Domingo E.,Franze-Fernández M. T. Rapid selection in modified BHK-21cells of a foot-and-mouth diseasevirus variant showing alterations in cell tropism. J Virol.,1998,72(12):10171–10179.
69. Escarmís C., Gómez-Mariano G., Dávila M., Lázaro E., Domingo E. Resistance to extinction oflow fitness virus subjected to plaque-to-plaque transfers: diversification by mutation clustering. JMol Biol.,2002,315(4):647–661.
70. Escarmís C., Lázaro E., Manrubia S. C. Population bottlenecks in quasispecies dynamics. Curr TopMicrobiol Immunol.,2006,299:141–170.
71. Escarmís C., Perales C., Domingo E. Biological effect of Muller's Ratchet: distant capsid site canaffect picornavirus protein processing. J Virol.,2009,83(13):6748–6756.
72. Esko J. D., Weinke J. L., Taylor W. H., Ekborg G., Rodén L., Anantharamaiah G., Gawish A.Inhibition of chondroitin and heparan sulfate biosynthesis in Chinese hamster ovary cell mutantsdefective in galactosyltransferase I. J Biol Chem.,1987,262(25):12189–12195.
73. Fauquet C. M., Mayo M. A., Maniloff J., Desselberger U., Ball L. A. Virus taxonomy. VIIIth reportof the international committee on taxonomy of viruses. San Diego, California: Elservier AcademicPress,2005.
74. Feliu J. X., Benito A., Oliva B., Avilés F. X., Villaverde A. Conformational flexibility in a highlymobile protein loop of foot-and-mouth disease virus: distinct structural requirements for integrinand antibody binding. J Mol Biol.,1998,283(2):331–338.
75. Fox G., Parry N. R., Barnett P. V., McGinn B., Rowlands D. J., Brown F. The cell attachment siteon foot-and-mouth disease virus includes the amino acid sequence RGD (arginine-glycine-asparticacid). J Gen Virol.,1989,70(Pt3):625–637.
76. Fry E. E., Lea S. M., Jackson T., Newman J. W., Ellard F. M., Blakemore W. E., Abu-Ghazaleh R.,Samuel A., King A. M., Stuart D. I. The structure and function of a foot-and-mouth diseasevirus-oligosaccharide receptor complex. EMBO J.,1999,18(3):543554.
77. Fry E. E., Newman J. W., Curry S., Najjam S., Jackson T., Blakemore W., Lea S. M., Miller L.,Burman A., King A. M., Stuart D. I. Structure of Foot-and-mouth disease virus serotype A1061alone and complexed with oligosaccharide receptor: receptor conservation in the face of antigenicvariation. J Gen Virol.,2005,86(Pt7):19091920.
78. Gloster J., Jones A., Redington A., Burgin L., S rensen J. H., Turner R., Dillon M., Hullinger P.,Simpson M., Astrup P., Garner G., Stewart P., D'Amours R., Sellers R., Paton D. Airborne spread offoot-and-mouth disease—model intercomparison. Vet J.,2010,183(3):278286.
79. Gloster J., Williams P., Doel C., Esteves I., Coe H., Valarcher J. F. Foot-and-mouthdisease-quantification and size distribution of airborne particles emitted by healthy and infectedpigs. Vet J.,2007,174(1):4253.
80. Grubman M. J. Development of novel strategies to control foot-and-mouth disease: markervaccines and antivirals. Biologicals,2005,33(4):227234.
81. Grubman M. J., Baxt B. Foot-and-mouth disease. Clin Microbiol Rev.,2004,17(2):465493.
82. Guo H., Liu X., Liu Z., Yin H., Ma J., Wang Y., Shang Y., Zhang Q., Li D., Guo J., Lu Z., Xie Q.Recent outbreaks of foot-and-mouth disease type Asia1in China. J Vet Med B Infect Dis VetPublic Health,2006,53(s1):2933.
83. Gutiérrez-Rivas M., Pulido M. R., Baranowski E., Sobrino F., Sáiz M. Tolerance to mutations inthe foot-and-mouth disease virus integrin-binding RGD region is different in cultured cells and invivo and depends on the capsid sequence context. J Gen Virol.,2008,89(Pt10):2531–2539.
84. Harwood L. J., Gerber H., Sobrino F., Summerfield A., McCullough K. C. Dendritic cellinternalization of foot-and-mouth disease virus: influence of heparan sulfate binding on virusuptake and induction of the immune response. J Virol.,2008,82(13):6379–6394.
85. Haydon D. T., Bastos A. D., Awadalla P. Low linkage disequilibrium indicative of recombination infoot-and-mouth disease virus gene sequence alignments. J Gen Virol.,2004,85(Pt5):10951100.
86. He D. S., Li K. N., Lin X. M., Lin S. R., Su D. P., Liao M. Genomic comparison of foot-and-mouthdisease virus R strain and its chick-passaged attenuated strain. Vet Microbiol.,2011,150(1-2):185–190.
87. Hemadri D., Tosh C., Sanyal A., Venkataramanan R. Emergence of a new strain of type Ofoot-and-mouth disease virus: its phylogenetic and evolutionary relationship with the PanAsiapandemic strain. Virus Genes,2002,25(1):2334.
88. Hernández J., Valero M. L., Andreu D., Domingo E., Mateu M. G. Antibody and host cellrecognition of foot-and-mouth disease virus (serotype C) cleaved at the Arg-Gly-Asp (RGD) motif:a structural interpretation. J Gen Virol.,1996,77(Pt2):257–264.
89. Hewat E. A., Verdaguer N., Fita I., Blakemore W., Brookes S., King A., Newman J., Domingo E.,Mateu M. G., Stuart D. I. Structure of the complex of an Fab fragment of a neutralizing antibodywith foot-and-mouth disease virus: positioning of a highly mobile antigenic loop. EMBO J.,1997,16(7):1492–1500.
90. Horsington J., Zhang Z. Consistent change in the B-C loop of VP2observed in foot-and-mouthdisease virus from persistently infected cattle: implications for association with persistence. VirusRes.,2007,125(1):114118.
91. Huang C. C., Lin Y. L., Huang T. S., Tu W. J., Lee S. H., Jong M. H., Lin S. Y. Molecularcharacterization of foot-and-mouth disease virus isolated from ruminants in Taiwan in1999-2000.Vet Microbiol.,2001,81(3):193205.
92. Huang C., Jong M., Lin S. Characteristics of foot and mouth disease virus in Taiwan. J Vet MedSci.,2000,62(7):677679.
93. Hui E. K. Reasons for the increase in emerging and re-emerging viral infectious diseases. MicrobesInfect.,2006,8(3):905916.
94. Jackson A. L., O'Neill H., Maree F., Blignaut B., Carrillo C., Rodriguez L., Haydon D. T. Mosaicstructure of foot-and-mouth disease virus genomes. J Gen Virol.,2007,88(Pt2):487492.
95. Jackson T., Blakemore W., Newman J. W., Knowles N. J., Mould A. P., Humphries M. J., King A.M. Foot-and-mouth disease virus is a ligand for the high-affinity binding conformation of integrinalpha5beta1: influence of the leucine residue within the RGDL motif on selectivity of integrinbinding. J Gen Virol.,2000a,81(Pt5):1383–1391.
96. Jackson T., Clark S., Berryman S., Burman A., Cambier S., Mu D., Nishimura S., King A. M.Integrin alphavbeta8functions as a receptor for foot-and-mouth disease virus: role of thebeta-chain cytodomain in integrin-mediated infection. J Virol.,2004,78(9):4533–4540.
97. Jackson T., Ellard F. M., Ghazaleh R. A., Brookes S. M., Blakemore W. E., Corteyn A. H., Stuart D.I., Newman J. W., King A. M. Efficient infection of cells in culture by type O foot-and-mouthdisease virus requires binding to cell surface heparan sulfate. J Virol.,1996,70(8):5282–5287.
98. Jackson T., King A. M., Stuart D. I., Fry E. Structure and receptor binding. Virus Res.,2003,91(1):3346.
99. Jackson T., Mould A. P., Sheppard D., King A. M. Integrin alphavbeta1is a receptor forfoot-and-mouth disease virus. J Virol.,2002,76(3):935–941.
100.Jackson T., Sharma A., Ghazaleh R. A., Blakemore W. E., Ellard F. M., Simmons D. L., Newman J.W., Stuart D. I., King A. M. Arginine-glycine-aspartic acid-specific binding by foot-and-mouthdisease viruses to the purified integrin alpha(v)beta3in vitro. J Virol.,1997,71(11):8357–8361.
101.Jackson T., Sheppard D., Denyer M., Blakemore W., King A. M. The epithelial integrinalphavbeta6is a receptor for foot-and-mouth disease virus. J Virol.,2000b,74(11):4949–4956.
102.Jamal S. M., Ferrari G., Ahmed S., Normann P., Belsham G. J. Genetic diversity of foot-and-mouthdisease virus serotype O in Pakistan and Afghanistan,1997-2009. Infect Genet Evol.,2011,11(6):12291238.
103.Jin H., Xiao C., Zhao G., Du X., Yu Y., Kang Y., Wang B. Induction of immature dendritic cellapoptosis by foot and mouth disease virus is an integrin receptor mediated event before viralinfection. J Cell Biochem.,2007,102(4):980–991.
104.Jin N. Y., Zhang H. Y., Yin G. F., Zheng M., Liu T., Jiang W. Z., Li Z. J. Immunogenicity ofrecombinant fowl-pox virus co-expressing structural protein precursor P1-2A and proteinase3C ofFMDV. Chinese Sci Bull.,2004,47(8):823827.
105.Jnaoui K., Minet M., Michiels T. Mutations that affect the tropism of DA and GDVII strains ofTheiler's virus in vitro influence sialic acid binding and pathogenicity. J Virol.,2002,76(16):8138–8147.
106.Johns H. L., Berryman S., Monaghan P., Belsham G. J., Jackson T. A dominant-negative mutant ofrab5inhibits infection of cells by foot-and-mouth disease virus: implications for virus entry. JVirol.,2009,83(12):6247–6256.
107.Kitching R. P. Global epidemiology and prospects for control of foot-and-mouth disease. Curr TopMicrobiol Immunol.,2005,288:133148.
108.Kitson J. D., McCahon D., Belsham G. J. Sequence analysis of monoclonal antibody resistantmutants of type O foot and mouth disease virus: evidence for the involvement of the three surfaceexposed capsid proteins in four antigenic sites. Virology,1990,179(1):2634.
109.Klein J. Understanding the molecular epidemiology of foot-and-mouth-disease virus. Infect GenetEvol.,2009,9:153161.
110.Klein J., Hussain M., Ahmad M., Normann P., Afzal M., Alexandersen S. Genetic characterisationof the recent foot-and-mouth disease virus subtype A/IRN/2005. Virol J.,2007,4:122.
111.Knipe T., Rieder E., Baxt B., Ward G., Mason P. W. Characterization of synthetic foot-and-mouthdisease virus provirions separates acid-mediated disassembly from infectivity. J Virol.,1997,71(4):2851–2856.
112.Knowles N. J., Davies P. R., Henry T., O'Donnell V., Pacheco J. M., Mason P. W. Emergence inAsia of foot-and-mouth disease viruses with altered host range: characterization of alterations inthe3A protein. J Virol.,2001,75(3):15511556.
113.Knowles N. J., Samuel A. R. Molecular epidemiology of foot-and-mouth disease virus. Virus Res.,2003,91:6580.
114.Knowles N. J., Samuel A. R., Davies P. R., Midgley R. J., Valarcher J. F. Pandemic strain offoot-and-mouth disease virus serotype O. Emerg Infect Dis.,2005,11(12):18871893.
115.Krebs O., Marquardt O. Identification and characterization of foot-and-mouth disease virus O1Burgwedel/1987as an intertypic recombinant. J Gen Virol.,1992,73(Pt3):613619.
116.Le V. P., Nguyen T., Lee K. N., Ko Y. J., Lee H. S., Nguyen V. C., Mai T. D., Do T. H., Kim S. M.,Cho I. S., Park J. H. Molecular characterization of serotype A foot-and-mouth disease virusescirculating in Vietnam in2009. Vet Microbiol.,2010,144(1-2):5866.
117.Lee K. N., Oem J. K., Park J. H., Kim S. M., Lee S. Y., Tserendorj Sh., Sodnomdarjaa R., Joo Y. S.,Kim H. Evidence of recombination in a new isolate of foot-and-mouth disease virus serotype Asia
1. Virus Res.,2009,139(1):117121.
118.Leippert M., Beck E., Weiland F., Pfaff E. Point mutations within the betaG-betaH loop offoot-and-mouth disease virus O1K affect virus attachment to target cells. J Virol.,1997,71(2):1046–1051.
119.Leippert M., Pfaff E. Foot-and-mouth disease virus can utilize the C-terminal extension ofcoxsackievirus A9VP1for cell infection. J Gen Virol.,2001,82(Pt7):1703–1711.
120.Leister D., Adam K. H., Marquardt O. Co-replication of several isotypes of foot-and-mouth diseasevirus. J Gen Virol.,1993,74(Pt12):2753–2757.
121.Lewis-Rogers N., McClellan D. A., Crandall K. A. The evolution of foot-and-mouth disease virus:impacts of recombination and selection. Infect Genet Evol.,2008,8(6):786798.
122.Li D., Liu Z. X., Bao H. F., Lu Z. J., Guo J. H., Cao Y. M., Li P. H., Bai X. W., Chen Y. L., Xie B.X., Cai X. P., Xie Q. G. The complete genome sequence of foot-and-mouth disease virusO/Akesu/58strain and its some molecular characteristics. Arch Virol.,2007a,152(11):20792085.
123.Li D., Shang Y. J., Liu Z. X., Liu X. T., Cai X. P. Comparisons of the complete genomes of twoChinese isolates of a recent foot-and-mouth disease type Asia1virus. Arch Virol.,2007b,152(9):16991708.
124.Li D., Shang Y. J., Liu Z. X., Liu X. T., Cai X. P. Molecular relationships between type Asia1newstrain from China and type O Panasia strains of foot-and-mouth-disease virus. Virus Genes,2007c,35(2):273279.
125.Li F., Browning G. F., Studdert M. J., Crabb B. S. Equine rhinovirus1is more closely related tofoot-and-mouth disease virus than to other picornaviruses. Proc Natl Acad Sci U S A,1996,93:990995.
126.Li J. Situation and control of FMD in mainland, China.16th OIE Sub-Commission Meeting forFMD Control in South East Asia,1519March2010, Lao PDR: Vientiane.2010.
127.Li P., Lu Z., Bao H., Li D., King D. P., Sun P., Bai X., Cao W., Gubbins S., Chen Y., Xie B., Guo J.,Yin H., Liu Z. In-vitro and in-vivo phenotype of type Asia1foot-and-mouth disease virusesutilizing two non-RGD receptor recognition sites. BMC Microbiol.,2011,11:154.
128.Loeffler F., Frosch P. Summarischer bericht über die ergebnisse der untersuchungen dercommission zur erforschung der maul-und klauenseuche be idem institute fürinfektionskrankheiten in Berlin. Centralbl. Bakteriol.,1897,22:257259.
129.Luna E., Rodríguez-Huete A., Rincón V., Mateo R., Mateu M. G. Systematic study of the geneticresponse of a variable virus to the introduction of deleterious mutations in a functional capsidregion. J Virol.,2009,83(19):10140–10151.
130.Luo M., Vriend G., Kamer G., Minor I., Arnold E., Rossmann M. G., Boege U., Scraba D. G., DukeG. M., Palmenberg A. C. The atomic structure of Mengo virus at3.0resolution. Science,1987,235(4785):182191.
131.Mahy B. W. J. Foot-and-mouth disease virus. Verlag Berlin Heidelberg: Springer,2005.
132.Malirat V., Bergmann I. E., Campos R. M., Salgado G., Sánchez C., Conde F., Quiroga J. L., OrtizS. Phylogenetic analysis of Foot-and-Mouth Disease Virus type O circulating in the Andean regionof South America during2002-2008. Vet Microbiol.,2011,152(1-2):7487.
133.Malirat V., Bergmann I. E., de Mendon a Campos R., Conde F., Quiroga J. L., Villamil M.,Salgado G., Ortiz S. Molecular epidemiology of foot-and-mouth disease virus type A in SouthAmerica. Vet Microbiol.,2012,[Epub ahead of print].
134.Maree F. F., Blignaut B., Aschenbrenner L., Burrage T., Rieder E. Analysis of SAT1typefoot-and-mouth disease virus capsid proteins: influence of receptor usage on the properties of virusparticles. Virus Res.,2011,155(2):462–472.
135.Maree F. F., Blignaut B., de Beer T. A., Visser N., Rieder E. A. Mapping of amino acid residuesresponsible for adhesion of cell culture-adapted foot-and-mouth disease SAT type viruses. VirusRes.,2010,153(1):82–91.
136.Martín M. J., Nú ez J. I., Sobrino F., Dopazo J. A procedure for detecting selection in highlyvariable viral genomes: evidence of positive selection in antigenic regions of capsid protein VP1offoot-and-mouth disease virus. J Virol Methods,1998,74(2):215221.
137.Martín-Acebes M. A., González-Magaldi M., Sandvig K., Sobrino F., Armas-Portela R. Productiveentry of type C foot-and-mouth disease virus into susceptible cultured cells requires clathrin and isdependent on the presence of plasma membrane cholesterol. Virology,2007,369(1):105–118.
138.Martín-Acebes M. A., González-Magaldi M., Vázquez-Calvo A., Armas-Portela R., Sobrino F.Internalization of swine vesicular disease virus into cultured cells: a comparative study withfoot-and-mouth disease virus. J Virol.,2009,83(9):4216–4226.
139.Martín-Acebes M. A., Rincón V., Armas-Portela R., Mateu M. G., Sobrino F. A single amino acidsubstitution in the capsid of foot-and-mouth disease virus can increase acid lability and conferresistance to acid-dependent uncoating inhibition. J Virol.,2010,84(6):2902–2912.
140.Martín-Acebes M. A., Vázquez-Calvo A., González-Magaldi M., Sobrino F. Foot-and-mouthdisease virus particles inactivated with binary ethylenimine are efficiently internalized into culturedcells. Vaccine,2011a,29(52):9655–9662.
141.Martín-Acebes M. A., Vázquez-Calvo A., Rincón V., Mateu M. G., Sobrino F. A single amino acidsubstitution in the capsid of foot-and-mouth disease virus can increase acid resistance. J Virol.,2011b,85(6):2733–2740.
142.Martínez M. A., Verdaguer N., Mateu M. G., Domingo E. Evolution subverting essentiality:dispensability of the cell attachment Arg-Gly-Asp motif in multiply passaged foot-and-mouthdisease virus. Proc Natl Acad Sci U S A,1997,94(13):6798–6802.
143.Martinsen J. S. Heterogeneity of foot-and-mouth disease virus: further studies on plaque formationby two plaque-size variants. Can J Comp Med.,1972,36(1):26–33.
144.Mason P. W., Grubman M. J., Baxt B. Molecular basis of pathogenesis of FMDV. Virus Res.,2003,91(1):932.
145.Mason P. W., Rieder E., Baxt B. RGD sequence of foot-and-mouth disease virus is essential forinfecting cells via the natural receptor but can be bypassed by an antibody-dependent enhancementpathway. Proc Natl Acad Sci U S A,1994,91(5):1932–1936.
146.Mateo R., Luna E., Mateu M. G. Thermostable variants are not generally represented infoot-and-mouth disease virus quasispecies. J Gen Virol.,2007,88(Pt3):859–864.
147.Mateo R., Mateu M. G. Deterministic, compensatory mutational events in the capsid offoot-and-mouth disease virus in response to the introduction of mutations found in viruses frompersistent infections. J Virol.,2007,81(4):1879–1887.
148.Mateu M. G. Antibody recognition of picornaviruses and escape from neutralization: a structureview. Virus Res.,1995,38:124.
149.Mateu M. G., Martínez M. A., Capucci L., Andreu D., Giralt E., Sobrino F., Brocchi E., Domingo E.A single amino acid substitution affects multiple overlapping epitopes in the major antigenic site offoot-and-mouth disease virus of serotype C. J Gen Virol.,1990,71(Pt3):629637.
150.Mateu M. G., Valero M. L., Andreu D., Domingo E. Systematic replacement of amino acid residueswithin an Arg-Gly-Asp-containing loop of foot-and-mouth disease virus and effect on cellrecognition. J Biol Chem.,1996,271(22):12814–12819.
151.McCahon D., Slade W. R., King A. M., Saunders K., Pullen L., Lake J. R., Priston R. A. Effect ofmutation on the virulence in mice of a strain of foot-and-mouth disease virus. J Gen Virol.,1981,54(Pt2):263–272.
152.McKenna T. S., Lubroth J., Rieder E., Baxt B., Mason P. W. Receptor binding site-deletedfoot-and-mouth disease (FMD) virus protects cattle from FMD. J Virol.,1995,69(9):57875790.
153.McVicar J. W., Sutmoller P. Three variants of foot-and-mouth disease virus type O: agar geldiffusion reactions. Am J Vet Res.,1972,33(8):16351639.
154.Miller L. C., Blakemore W., Sheppard D., Atakilit A., King A. M., Jackson T. Role of thecytoplasmic domain of the beta-subunit of integrin alpha(v)beta6in infection by foot-and-mouthdisease virus. J Virol.,2001,75(9):4158–4164.
155.Monaghan P., Gold S., Simpson J., Zhang Z., Weinreb P. H., Violette S. M., Alexandersen S.,Jackson T. The alpha(v)beta6integrin receptor for foot-and-mouth disease virus is expressedconstitutively on the epithelial cells targeted in cattle. J Gen Virol.,2005,86(Pt10):2769–2780.
156.Moore D. M., Cowan K. M. Effect of trypsin and chymotrypsin on the polypeptides of large andsmall plaque variants of foot-and-mouth disease virus: relationship to specific antigenicity andinfectivity. J Gen Virol.,1978,41(3):549–562.
157.Morioka K., Fukai K., Ohashi S., Sakamoto K., Tsuda T., Yoshida K. Comparison of the charactersof the plaque-purified viruses from foot-and-mouth disease virus O/JPN/2000. J Vet Med Sci.,2008,70(7):653–658.
158.Murphy F. A., Gibbs E., Horzinek M. C., Studdert M. J. Veterinary virology.3rd. San Diego,California: Academic Press,1999.
159.Neff S., Baxt B. The ability of integrin alpha(v)beta(3) to function as a receptor for foot-and-mouthdisease virus is not dependent on the presence of complete subunit cytoplasmic domains. J Virol.,2001,5(1):527–532.
160.Neff S., Mason P. W., Baxt B. High-efficiency utilization of the bovine integrin alpha(v)beta(3) asa receptor for foot-and-mouth disease virus is dependent on the bovine beta(3) subunit. J Virol.,2000,74(16):7298–7306.
161.Neff S., Sá-Carvalho D., Rieder E., Mason P. W., Blystone S. D., Brown E. J., Baxt B.Foot-and-mouth disease virus virulent for cattle utilizes the integrin alpha(v)beta3as its receptor. JVirol.,1998,72(5):3587–3594.
162.Nilsson E. C., Jamshidi F., Johansson S. M., Oberste M. S., Arnberg N. Sialic acid is a cellularreceptor for coxsackievirus A24variant, an emerging virus with pandemic potential. J Virol.,2008,82(6):3061–3068.
163.Nobiron I., Rémond M., Kaiser C., Lebreton F., Zientara S., Delmas B. The nucleotide sequence offoot-and-mouth disease virus O/FRA/1/2001and comparison with its British parental strainO/UKG/35/2001. Virus Res.,2005,108(1-2):225229.
164.Norkin L. C. Virus receptors: implications for pathogenesis and the design of antiviral agents. ClinMicrobiol Rev.,1995,8(2):293315.
165.Novella I. S., Borrego B., Mateu M. G., Domingo E., Giralt E., Andreu D. Use of substituted andtandem-repeated peptides to probe the relevance of the highly conserved RGD tripeptide in theimmune response against foot-and-mouth disease virus. FEBS Lett.,1993,330(3):253259.
166.Nú ez J. I., Martín M. J., Piccone M. E., Carrillo E., Palma E. L., Dopazo J., Sobrino F.Identification of optimal regions for phylogenetic studies on VP1gene of foot-and-mouth diseasevirus: analysis of types A and O Argentinean viruses. Vet Res.,2001,32(1):31–45.
167.Nú ez J. I., Molina N., Baranowski E., Domingo E., Clark S., Burman A., Berryman S., Jackson T.,Sobrino F. Guinea pig-adapted foot-and-mouth disease virus with altered receptor recognition canproductively infect a natural host. J Virol.,2007,81(16):8497–8506.
168.O'Donnell V., Larocco M., Baxt B. Heparan sulfate-binding foot-and-mouth disease virus enterscells via caveola-mediated endocytosis. J Virol.,2008,82(18):9075–9085.
169.O'Donnell V., LaRocco M., Duque H., Baxt B. Analysis of foot-and-mouth disease virusinternalization events in cultured cells. J Virol.,2005,79(13):8506–8518.
170.O'Donnell V., Pacheco J. M., Gregg D., Baxt B. Analysis of foot-and-mouth disease virus integrinreceptor expression in tissues from na ve and infected cattle. J Comp Pathol.,2009,141(2-3):98–112.
171.O'Donnell V., Pacheco J. M., LaRocco M., Burrage T., Jackson W., Rodriguez L. L., Borca M. V.,Baxt B. Foot-and-mouth disease virus utilizes an autophagic pathway during viral replication.Virology,2011,410(1):142–150.
172.Oem J. K., Yeh M. T., McKenna T. S., Hayes J. R., Rieder E., Giuffre A. C., Robida J. M., Lee K.N., Cho I. S., Fang X., Joo Y. S., Park J. H. Pathogenic characteristics of the Korean2002isolate offoot-and-mouth disease virus serotype O in pigs and cattle. J Comp Pathol.,2008,138(4):204–214.
173.Office International des Epizooties. Handistatus II. In: International Office of Epizootics,2006.
174.Office International des Epizooties. O.I.E Bull.,1999, N°3:202.
175.Pacheco J. M., Arzt J., Rodriguez L. L. Early events in the pathogenesis of foot-and-mouth diseasein cattle after controlled aerosol exposure. Vet J.,2010,183(1):4653.
176.Pacheco J. M., Mason P. W. Evaluation of infectivity and transmission of different Asianfoot-and-mouth disease viruses in swine. J Vet Sci.,2010,11(2):133–142.
177.Parry N., Fox G., Rowlands D., Brown F., Fry E., Acharya R., Logan D., Stuart D. Structural andserological evidence for a novel mechanism of antigenic variation in foot-and-mouth disease virus.Nature,1990,347(6293):569572.
178.Paton D. J., King D. P., Knowles N. J., Hammond J. Recent spread of foot-and-mouth disease inthe Far East. Vet Rec.,2010,166(18):569570.
179.Pfaff M. Recognition sites of RGD-dependent integrins. In: Eble J. A., Kühn K., eds.Intergrin-ligand interaction. Heidelberg: Springer-Verlag,1997,101121.
180.Piccone M. E., Pacheco J. M., Pauszek S. J., Kramer E., Rieder E., Borca M. V., Rodriguez L. L.The region between the two polyprotein initiation codons of foot-and-mouth disease virus iscritical for virulence in cattle. Virology,2010,396(1):152–159.
181.Rieder E., Baxt B., Mason P. W. Animal-derived antigenic variants of foot-and-mouth disease virustype A12have low affinity for cells in culture. J Virol.,1994,68(8):5296–5299.
182.Rieder E., Berinstein A., Baxt B., Kang A., Mason P. W. Propagation of an attenuated virus bydesign: engineering a novel receptor for a noninfectious foot-and-mouth disease virus. Proc NatlAcad Sci U S A,1996,93(19):10428–10433.
183.Rieder E., Henry T., Duque H., Baxt B. Analysis of a foot-and-mouth disease virus type A24isolate containing an SGD receptor recognition site in vitro and its pathogenesis in cattle. J Virol.,2005,79(20):12989–12998.
184.Robinson L., Windsor M., McLaughlin K., Hope J., Jackson T., Charleston B. Foot-and-mouthdisease virus exhibits an altered tropism in the presence of specific immunoglobulins, enablingproductive infection and killing of dendritic cells. J Virol.,2011,85(5):2212–2223.
185.Rojas E. R., Carrillo E., Schiappacassi M., Campos R. Modification of foot-and-mouth diseasevirus O1Caseros after serial passages in the presence of antiviral polyclonal sera. J Virol.,1992,66(6):33683372.
186.Ruiz-Jarabo C. M., Arias A., Baranowski E., Escarmís C., Domingo E. Memory in viralquasispecies. J Virol.,2000,74(8):35433547.
187.Ruíz-Jarabo C. M., Arias A., Molina-París C., Briones C., Baranowski E., Escarmís C., Domingo E.Duration and fitness dependence of quasispecies memory. J Mol Biol.,2002,315(3):285–296.
188.Ruiz-Jarabo C. M., Sevilla N., Dávila M., Gómez-Mariano G., Baranowski E., Domingo E.Antigenic properties and population stability of a foot-and-mouth disease virus with an alteredArg-Gly-Asp receptor-recognition motif. J Gen Virol.,1999,80(Pt8):1899–1909.
189.Ruiz-Sáenz J., Goez Y., Tabares W., López-Herrera A. Cellular receptors for foot and mouthdisease virus. Intervirology,2009,52(4):201–212.
190.Sa-Carvalho D., Rieder E., Baxt B., Rodarte R., Tanuri A., Mason P. W. Tissue culture adaptationof foot-and-mouth disease virus selects viruses that bind to heparin and are attenuated in cattle. JVirol.,1997,71(7):5115–5123.
191.Saitou N., Nei M. The neighbor-joining method: a new method for reconstructing phylogenetictrees. Mol Biol Evol.,1987,4(4):406425.
192.Sáiz M., Gómez S., Martínez-Salas E., Sobrino F. Deletion or substitution of the aphthovirus3'NCR abrogates infectivity and virus replication. J Gen Virol.,2001,82(Pt1):93101.
193.Sakamoto K., Yoshida K. Recent outbreaks of foot and mouth disease in countries of East Asia.Rev Sci Tech.,2002,21(3):459463.
194.Sanz-Ramos M., Díaz-San Segundo F., Escarmís C., Domingo E., Sevilla N. Hidden virulencedeterminants in a viral quasispecies in vivo. J Virol.2008,82(21):10465–10476.
195.Schneider-Schaulies J. Cellular receptors for viruses: links to tropism and pathogenesis. J GenVirol.,2000,81(Pt6):14131429.
196.Schols D. HIV co-receptor inhibitors as novel class of anti-HIV drugs. Antiviral Res.,2006,71(2-3):216226.
197.Schumann K. R., Knowles N. J., Davies P. R., Midgley R. J., Valarcher J. F., Raoufi A. Q.,McKenna T. S., Hurtle W., Burans J. P., Martin B. M., Rodriguez L. L., Beckham T. R. Geneticcharacterization and molecular epidemiology of foot-and-mouth disease viruses isolated formAfghanistan in2003-2005. Virus Genes,2008,36(2):401413.
198.Semler B. L., Wimmer E. Molecular Biology of Picornaviruses. Washington, DC: AmericanSociety for Microbiology Press,2002.
199.Sevilla N., Domingo E. Evolution of a persistent aphthovirus in cytolytic infections: partialreversion of phenotypic traits accompanied by genetic diversification. J Virol.,1996,70(10):6617–6624.
200.Sevilla N., Ruiz-Jarabo C. M., Gómez-Mariano G., Baranowski E., Domingo E. An RNA virus canadapt to the multiplicity of infection. J Gen Virol.,1998,79(Pt12):2971–2980.
201.Sharma A., Rao Z., Fry E., Booth T., Jones E. Y., Rowlands D. J., Simmons D. L., Stuart D. I.Specific interactions between human integrin alpha v beta3and chimeric hepatitis B virus coreparticles bearing the receptor-binding epitope of foot-and-mouth disease virus. Virology,1997,239(1):150–157.
202.Sieczkarski S. B., Whittaker G. R. Viral entry. Curr Top Microbiol Immunol.,2005,285:123.
203.Sobrino F., Domingo E. Foot and mouth disease: current perspectives. Norfolk: HorizonBioscience,2004.
204.Sobrino F., Sáiz M., Jiménez-Clavero M. A., Nú ez J. I., Rosas M. F., Baranowski E., Ley V.Foot-and-mouth disease virus: a long known virus, but a current threat. Vet Res.,2001,32(1):130.
205.Storey P., Theron J., Maree F. F., O'Neill H. G. A second RGD motif in the1D capsid protein of aSAT1type foot-and-mouth disease virus field isolate is not essential for attachment to target cells.Virus Res.,2007,124(1-2):184–192.
206.Tami C., Taboga O., Berinstein A., Nú ez J. I., Palma E. L., Domingo E., Sobrino F., Carrillo E.Evidence of the coevolution of antigenicity and host cell tropism of foot-and-mouth disease virusin vivo. J Virol.,2003,77(2):1219–1226.
207.Thompson D., Muriel P., Russell D., Osborne P., Bromley A., Rowland M., Creigh-Tyte S., BrownC. Economic costs of the foot and mouth disease outbreak in the United Kingdom in2001. Rev SciTech.,2002,21(3):675687.
208.Tosh C., Hemadri D., Sanyal A. Evidence of recombination in the capsid-coding region of type Afoot-and-mouth disease virus. J Gen Virol.,2002,83(Pt10):24552460.
209.Tuthill T. J., Harlos K., Walter T. S., Knowles N. J., Groppelli E., Rowlands D. J., Stuart D. I., FryE. E. Equine rhinitis A virus and its low pH empty particle: clues towards an aphthovirus entrymechanism? PLoS Pathog.,2009,5(10): e1000620.
210.Twomey T., France L. L., Hassard S., Burrage T. G., Newman J. F., Brown F. Characterization ofan acid-resistant mutant of foot-and-mouth disease virus. Virology,1995,206(1):69–75.
211.Valarcher J. F., Knowles N. J., Ferris N. P., Paton D. J., Zakharov V., Sherbakov A., Shang Y. J.,Liu Z. X., Liu X. T., Sanyal A., Hemadri D., Tosh C., Rasool T. J. Recent spread of FMD virusserotype Asia1. Vet Rec.,2005,157(1):30.
212.Valarcher J. F., Knowles N. J., Fernandez R., Davies P. R., Midgley R. J., Hutchings G., Newman B.J., Ferris N. P., Paton D. J. FMD global update2003-2004. In: Open sessions of the EUFMDresearch group. Greece: Chaina,2004.
213.van Rensburg H. G., Henry T. M., Mason P. W. Studies of genetically defined chimeras of aEuropean type A virus and a South African Territories type2virus reveal growth determinants forfoot-and-mouth disease virus. J Gen Virol.,2004,85(Pt1):6168.
214.van Vlijmen H. W., Curry S., Schaefer M., Karplus M. Titration calculations of foot-and-mouthdisease virus capsids and their stabilities as a function of pH. J Mol Biol.,1998,275(2):295–308.
215.Verdaguer N., Mateu M. G., Andreu D., Giralt E., Domingo E., Fita I. Structure of the majorantigenic loop of foot-and-mouth disease virus complexed with a neutralizing antibody: directinvolvement of the Arg-Gly-Asp motif in the interaction. EMBO J.,1995,14(8):16901696.
216.Verdaguer N., Sevilla N., Valero M. L., Stuart D., Brocchi E., Andreu D., Giralt E., Domingo E.,Mateu M. G., Fita I. A similar pattern of interaction for different antibodies with a major antigenicsite of foot-and-mouth disease virus: implications for intratypic antigenic variation. J Virol.,1998,72(1):739748.
217.Vlodavsky I., Ilan N., Naggi A., Casu B. Heparanase: structure, biological functions, and inhibitionby heparin-derived mimetics of heparan sulfate. Curr Pharm Des.,2007,13(20):20572073.
218.Wang H., Zhao L., Li W., Zhou G., Yu L. Identification of a conformational epitope on the VP1G-H Loop of type Asia1foot-and-mouth disease virus defined by a protective monoclonal antibody.Vet Microbiol.,2011,148(2-4):189199.
219.Wilke C. O., Novella I. S. Phenotypic mixing and hiding may contribute to memory in viralquasispecies. BMC Microbiol.,2003,3:11.
220.Woodbury E. L., Samuel A. R., Knowles N. J., Hafez S. M., Kitching R. P. Analysis of mixedfoot-and-mouth disease virus infections in Saudi Arabia: prolonged circulation of an exoticserotype. Epidemiol Infect.,1994,112(1):201211.
221.Yang D., Zhang C., Zhao L., Zhou G., Wang H., Yu L. Identification of a conserved linear epitopeon the VP1protein of serotype O foot-and-mouth disease virus by neutralising monoclonalantibody8E8. Virus Res.,2011,155(1):291299.
222.Yang X., Zhou Y. S., Wang H. N., Zhang Y., Wei K., Wang T. Isolation, identification and completegenome sequence analysis of a strain of foot-and-mouth disease virus serotype Asia1from pigs insouthwest of China. Virol J.,2011,8:175.
223.Yi J., Liu Ch. Integrity of foot-and-mouth disease virions bound to the cells. Acta Virol.,2010,54(4):307–310.
224.Yoon S. H., Park W., King D. P., Kim H. Phylogenomics and molecular evolution offoot-and-mouth disease virus. Mol Cells,2011,31(5):413421.
225.Zhang L., David G., Esko J. D. Repetitive Ser-Gly sequences enhance heparan sulfate assembly inproteoglycans. J Biol Chem.,1995,270(45):27127–27135.
226.Zhang X. S., Liu Z. X., Zhao Q. Z., Chang H. Y., Xie Q. G. Sequencing and analysis for thefull-length genome RNA of foot-and-mouth disease virus China/99. Chinese Sci Bull.,2004,47(1):7481.
227.Zhao M. Q., Suo Q. L., Chen J. D., Chen L. J., Xu Y. F. Sequence analysis of the protein-codingregions of foot-and-mouth disease virus O/HK/2001. Vet Microbiol.,2008,130(3-4):238246.
228.Zhao Q., Pacheco J. M., Mason P. W. Evaluation of genetically engineered derivatives of a Chinesestrain of foot-and-mouth disease virus reveals a novel cell-binding site which functions in cellculture and in animals. J Virol.,2003,77(5):3269–3280.
229.Zheng H. X., Guo J. H., Jin Y., Shang Y. J., Tian H., Yang Y. M., Liu X. T., Cai X. P. Infectivity ofAsia1type foot-and-mouth disease virus was increased via an alternative RDD motif. Chinese SciBull.,2010,55:13701375.
230.Zheng H. X., He J. J., Guo J. H., Jin Y., Yang F., Lv L., Liu X. T. Genetic characterization of a newpandemic Southeast Asia topotype strain of serotype O foot-and-mouth disease virus isolated inChina during2010. Virus Genes,2012,44(1):8088.
231.Zhong J., Li Y., Zhao S., Liu S., Zhang Z. Mutation pressure shapes codon usage in the GC-Richgenome of foot-and-mouth disease virus. Virus Genes,2007,35(3):767776.
232.Zhou J. H., Zhang J., Chen H. T., Ma L. N., Ding Y. Z., Pejsak Z., Liu Y. S. The codon usage modelof the context flanking each cleavage site in the polyprotein of foot-and-mouth disease virus. InfectGenet Evol.,2011,11(7):18151819.
233.Zhou J. H., Zhang J., Chen H. T., Ma L. N., Liu Y. S. Analysis of synonymous codon usage infoot-and-mouth disease virus. Vet Res Commun.,2010a,34:393404.
234.Zhou J. H., Zhang J., Ding Y. Z., Chen H. T., Ma L. N., Liu Y. S. Characteristics of codon usagebias in two regions downstream of the initiation codons of foot-and-mouth disease virus.BioSystems,2010b,101:2028.
2.刘在新.口蹄疫病毒基因组及其编码蛋白一级结构研究.[博士学位论文].北京:中国农业科学院,2002.
3.刘增胜,谷继承,陈伟生.中国畜牧业年鉴.北京:中国农业出版社,2007.
4.浦华.动物疾病防控的经济学研究综述.农业经济问题,2006,6:6164.
5.谢庆阁.主编.口蹄疫.北京:中国农业出版社,2004.
6.熊忠良,徐涤平.我国重大动物疫病流行现状及其防控策略.湖北畜牧兽医,2007,12:2224.
7.徐耀先,周晓峰,刘立德.主编.分子病毒学.武汉:湖北科学技术出版社,2000.
8.殷震,刘景华.主编.动物病毒学.第二版.北京:科学出版社,1997.
9. Abdul-Hamid N. F., Hussein N. M., Wadsworth J., Radford A. D., Knowles N. J., King D. P.Phylogeography of foot-and-mouth disease virus types O and A in Malaysia and surroundingcountries. Infect Genet Evol.,2011,11(2):320328.
10. Acharya R., Fry E., Stuart D., Fox G., Rowlands D., Brown F. The three-dimensional structure offoot-and-mouth disease virus at2.9resolution. Nature,1989,337(6209):709–716.
11. Aggarwal N., Barnett P. V. Antigenic sites of foot-and-mouth disease virus (FMDV): an analysis ofthe specificities of anti-FMDV antibodies after vaccination of naturally susceptible host species. JGen Virol.,2002,83(Pt4):775–782.
12. Ahn I., Bae S. E., Son H. S. Comparative study of codon substitution patterns in foot-and-mouthdisease virus (serotype O). Exp Mol Med.,2011,43(10):587595.
13. Ansell D. M., Samuel A. R., Carpenter W. C., Knowles N. J. Genetic relationships betweenfoot-and-mouth disease type Asia1viruses. Epidemiol Infect.,1994,112(1):213224.
14. Arias A., Agudo R., Ferrer-Orta C., Pérez-Luque R., Airaksinen A., Brocchi E., Domingo E.,Verdaguer N., Escarmís C. Mutant viral polymerase in the transition of virus to error catastropheidentifies a critical site for RNA binding. J Mol Biol.,2005,353(5):1021–1032.
15. Arias A., Perales C., Escarmís C., Domingo E. Deletion mutants of VPg reveal new cytopathologydeterminants in a picornavirus. PLoS One,2010,5(5): e10735.
16. Arzt J., Pacheco J. M., Rodriguez L. L. The early pathogenesis of foot-and-mouth disease in cattleafter aerosol inoculation. Identification of the nasopharynx as the primary site of infection. VetPathol.,2010,47(6):1048–1063.
17. Asokan A., Hamra J. B., Govindasamy L., Agbandje-McKenna M., Samulski R. J.Adeno-associated virus type2contains an integrin alpha5beta1binding domain essential for viralcell entry. J Virol.,2006,80(18):8961–8969.
18. Ayelet G., Mahapatra M., Gelaye E., Egziabher B. G., Rufeal T., Sahle M., Ferris N. P., WadsworthJ., Hutchings G. H., Knowles N. J. Genetic characterization of foot-and-mouth disease viruses,Ethiopia,1981-2007. Emerg Infect Dis.,2009,15(9):14091417.
19. Bao H. F., Li D., Sun P., Zhou Q., Hu J., Bai X. W., Fu Y. F., Lu Z. J., Liu Z. X. The infectivity andpathogenicity of a foot-and-mouth disease virus persistent infection strain fromoesophageal-pharyngeal fluid of a Chinese cattle in2010. Virol J.,2011,8:536.
20. Baranowski E., Molina N., Nú ez J. I., Sobrino F., Sáiz M. Recovery of infectious foot-and-mouthdisease virus from suckling mice after direct inoculation with in vitro-transcribed RNA. J Virol.,2003,77(20):1129011295.
21. Baranowski E., Ruiz-Jarabo C. M., Sevilla N., Andreu D., Beck E., Domingo E. Cell recognitionby foot-and-mouth disease virus that lacks the RGD integrin-binding motif: flexibility inaphthovirus receptor usage. J Virol.,2000,74(4):1641–1647.
22. Baranowski E., Sevilla N., Verdaguer N., Ruiz-Jarabo C. M., Beck E., Domingo E. Multiplevirulence determinants of foot-and-mouth disease virus in cell culture. J Virol.,1998,72(8):6362–6372.
23. Barnett P. V., Ouldridge E. J., Rowlands D. J., Brown F., Parry N. R. Neutralizing epitopes of typeO foot-and-mouth disease virus. I. Identification and characterization of three functionallyindependent, conformational sites. J Gen Virol.,1989,70(Pt6):1483–1491.
24. Baxt B., Mason P. W. Foot-and-mouth disease virus undergoes restricted replication in macrophagecell cultures following Fc receptor-mediated adsorption. Virology,1995,207(2):503–509.
25. Bayry J., Tough D. F. Interaction of foot-and-mouth disease virus with dendritic cells. TrendsMicrobiol.,2006,14(8):346–347.
26. Beard C. W., Mason P. W. Genetic determinants of altered virulence of Taiwanese foot-and-mouthdisease virus. J Virol.,2000,74(2):987991.
27. Belnap D. M., Filman D. J., Trus B. L., Cheng N., Booy E. P., Conway J. E., Curry S., Hiremath C.N., Tsang S. K., Steven A. C., Hogle J. M. Molecular tectonic model of virus structure transitions:the putative cell entry states of poliovirus. J Virol.,2000,74:13421354.
28. Belsham G. J., Jamal S. M., Tj rneh j K., B tner A. Rescue of foot-and-mouth disease viruses thatare pathogenic for cattle from preserved viral RNA samples. PLoS One,2011,6(1): e14621.
29. Berinstein A., Roivainen M., Hovi T., Mason P. W., Baxt B. Antibodies to the vitronectin receptor(integrin alpha V beta3) inhibit binding and infection of foot-and-mouth disease virus to culturedcells. J Virol.,1995,69(4):2664–2666.
30. Berryman S., Clark S., Monaghan P., Jackson T. Early events in integrin alphavbeta6-mediated cellentry of foot-and-mouth disease virus. J Virol.,2005,79(13):8519–8534.
31. Bittle J. L., Houghen R. A., Alexander H., Shinnick T. M., Sutcliffe J. G., Lerner R. A., RowlandsD. J., Brown F. Protection against foot-and-mouth disease by immunization with a chemicallysynthesized peptide predicted from the viral nucleotide sequence. Nature,1982,298(5869):3033.
32. Bolwell C., Brown A. L., Barnett P. V., Campbell R. O., Clarke B. E., Parry N. R., Ouldridge E. J.,Brown F., Rowlands D. J. Host cell selection of antigenic variants of foot-and-mouth disease virus.J Gen Virol.,1989,70(Pt1):4557.
33. Borca M. V., Pacheco J. M., Holinka L. G., Carrillo C., Hartwig E., Garriga D., Kramer E.,Rodriguez L., Piccone M. E. Role of arginine-56within the structural protein VP3offoot-and-mouth disease virus (FMDV) O1Campos in virus virulence. Virology,2012,422(1):37–45.
34. B tner A., Kakker N. K., Barbezange C., Berryman S., Jackson T., Belsham G. J. Capsid proteinsfrom field strains of foot-and-mouth disease virus confer a pathogenic phenotype in cattle on anattenuated, cell-culture-adapted virus. J Gen Virol.,2011,92(Pt5):1141–1151.
35. Brown F. The history of research in foot-and-mouth disease. Virus Res.,2003,91:37.
36. Brown J. K., McAleese S. M., Thornton E. M., Pate J. A., Schock A., Macrae A. I., Scott P. R.,Miller H. R., Collie D. D. Integrin-alphavbeta6, a putative receptor for foot-and-mouth diseasevirus, is constitutively expressed in ruminant airways. J Histochem Cytochem.,2006,54(7):807–816.
37. Burman A., Clark S., Abrescia N. G., Fry E. E., Stuart D. I., Jackson T. Specificity of the VP1GHloop of foot-and-mouth disease virus for alphav integrins. J Virol.,2006,80(19):9798–9810.
38. Camper N., Byrne T., Burden R. E., Lowry J., Gray B., Johnston J. A., Migaud M. E., Olwill S. A.,Buick R. J., Scott C. J. Stable expression and purification of a functional processed Fab' fragmentfrom a single nascent polypeptide in CHO cells expressing the mCAT-1retroviral receptor. JImmunol Methods,2011,372(1-2):30–41.
39. Cao Y., Lu Z., Sun P., Fu Y., Tian F., Hao X., Bao H., Liu X., Liu Z. A pseudotype baculovirusexpressing the capsid protein of foot-and-mouth disease virus and a T-cell immunogen showsenhanced immunogenicity in mice. Virol J.,2011,8:77.
40. Carrillo C., Lu Z., Borca M. V., Vagnozzi A., Kutish G. F., Rock D. L. Genetic and phenotypicvariation of foot-and-mouth disease virus during serial passages in a natural host. J Virol.,2007,81(20):11341–11351.
41. Carrillo C., Tulman E. R., Delhon G., Lu Z., Carreno A., Vagnozzi A., Kutish G. F., Rock D. L.Comparative genomics of foot-and-mouth disease virus. J Virol.,2005,79(10):64876504.
42. Chang Y., Zheng H., Shang Y., Jin Y., Wang G., Shen X., Liu X. Recovery of infectiousfoot-and-mouth disease virus from full-length genomic cDNA clones using an RNA polymerase Isystem. Acta Biochim Biophys Sin.,2009,41(12):9981007.
43. Charpentier N., Dávila M., Domingo E., Escarmís C. Long-term, large-population passage ofaphthovirus can generate and amplify defective noninterfering particles deleted in the leaderprotease gene. Virology,1996,223(1):1018.
44. Chen J. D., Zhao M. Q., Hui K. H., Leung F. C. Molecular characterization of foot-and-mouthdisease virus in Hong Kong during2001-2002. Virus Genes,2006,32(2):139143.
45. Cottam E. M., Haydon D. T., Paton D. J., Gloster J., Wilesmith J. W., Ferris N. P., Hutchings G. H.,King D. P. Molecular epidemiology of the foot-and-mouth disease virus outbreak in the UnitedKingdom in2001. J Virol.,2006,80(22):1127411282.
46. Cottam E. M., Wadsworth J., Shaw A. E., Rowlands R. J., Goatley L., Maan S., Maan N. S.,Mertens P. P., Ebert K., Li Y., Ryan E. D., Juleff N., Ferris N. P., Wilesmith J. W., Haydon D. T.,King D. P., Paton D. J., Knowles N. J. Transmission pathways of foot-and-mouth disease virus inthe United Kingdom in2007. PLoS Pathog.,2008,4(4): e1000050.
47. Cowan K. M. Immunochemical studies of foot-and-mouth disease. V. Antigenic variants of virusdemonstrated by immunodiffusion analyses with19S but not7S antibodies. J Exp Med.,1969,129(2):333350.
48. Cowan K. M., Erol N., Whiteland A. P. Heterogeneity of type Asia1foot-and-mouth disease virusand BHK-21cells and the relationship to vaccine preparation. O. I. E Bull.,1974,81:12711298.
49. Crowther J. R., Farias S., Carpenter W. C., Samuel A. R. Identification of a fifth neutralizable siteon type O foot-and-mouth disease virus following characterization of single and quintuplemonoclonal antibody escape mutants. J Gen Virol.,1993,74:17471953.
50. Curry S., Abrams C. C., Fry E., Crowther J. C., Belsham G. J., Stuart D. I., King A. M. Viral RNAmodulates the acid sensitivity of foot-and-mouth disease virus capsids. J Virol.,1995,69(1):430–438.
51. Delputte P. L., Costers S., Nauwynck H. J. Analysis of porcine reproductive and respiratorysyndrome virus attachment and internalization: distinctive roles for heparan sulphate andsialoadhesin. J Gen Virol.,2005,86(Pt5):1441–1445.
52. Dicara D., Burman A., Clark S., Berryman S., Howard M. J., Hart I. R., Marshall J. F., Jackson T.Foot-and-mouth disease virus forms a highly stable, EDTA-resistant complex with its principalreceptor, integrin alphavbeta6: implications for infectiousness. J Virol.,2008,82(3):1537–1546.
53. Díez J., Dávila M., Escarmís C., Mateu M. G., Dominguez J., Pérez J. J., Giralt E., Melero J. A.,Domingo E. Unique amino acid substitutions in the capsid proteins of foot-and-mouth disease virusfrom a persistent infection in cell culture. J Virol.,1990,64(11):5519–5528.
54. Domingo E., Escarmís C., Baranowski E., Ruiz-Jarabo C. M., Carrillo E., Nú ez J. I., Sobrino F.Evolution of foot-and-mouth disease virus. Virus Res.,2003,91(1):47–63.
55. Domingo E., Escarmis C., Martinez M. A., Martinez-Salas E., Mateu M. G. Foot-and-mouthdisease virus populations are quasispecies. Curr Top Microbiol Immunol.,1992,176:3347.
56. Domingo E., Mateu M. C., Martinez M. A., Dopazo J., Moya A., Sobrino F. Genetic variability andantigenic diversity of foot-and-mouth disease virus. In: Kurstak E., Marusyk R. G., Murphy F. A.,van Regenmortel M. H. V. eds. Applied virology research, Vol2, Virus variation and epidemiology.New York: Plenum Publishing Corp.,1990.233266.
57. Domingo E., Ruiz-Jarabo C. M., Sierra S., Arias A., Pariente N., Baranowski E., Escarmis C.Emergence and selection of RNA virus variants: memory and extinction. Virus Res.,2002,82(1-2):3944.
58. Drake J. W. Rates of spontaneous mutation among RNA viruses. Proc Natl Acad Sci U S A,1993,90(9):41714175.
59. Drake J. W., Holland J. J. Mutation rates among RNA viruses. Proc Natl Acad Sci U S A,1999,96(24):1391013913.
60. Du J., Chang H., Cong G., Shao J., Lin T., Shang Y., Liu Z., Liu X., Cai X., Xie Q. Completenucleotide sequence of a Chinese serotype Asia1vaccine strain of foot-and-mouth disease virus.Virus Genes,2007,35(3):635642.
61. Du J., Chang H., Gao S., Xue S., Cong G., Shao J., Lin T., Liu Z., Liu X., Cai X. Molecularcharacterization and expression analysis of porcine integrins alphavbeta3, alphavbeta6andalphavbeta8that are potentially involved in FMDV infection. Mol Cell Probes,2010,24(5):256–265.
62. Dunn C. S., Donaldson A. I. Natural adaption to pigs of a Taiwanese isolate of foot-and-mouthdisease virus. Vet Rec.,1997,141(7):174175.
63. Duque H., Baxt B. Foot-and-mouth disease virus receptors: comparison of bovine alpha(V)integrin utilization by type A and O viruses. J Virol.,2003,77(4):2500–2511.
64. Duque H., LaRocco M., Golde W. T., Baxt B. Interactions of foot-and-mouth disease virus withsoluble bovine alphaVbeta3and alphaVbeta6integrins. J Virol.,2004,78(18):9773–9781.
65. Durand S., Murphy C., Zhang Z., Alexandersen S. Epithelial distribution and replication offoot-and-mouth disease virus RNA in infected pigs. J Comp Pathol.,2008,139(2-3):86–96.
66. Edwards J., Morrissy C. China-EU training to improve prevention and control of foot-and-mouthdisease.812August2011, P. R. China: Lanzhou.2011.
67. Ellard F. M., Drew J., Blakemore W. E., Stuart D. I., King A. M. Evidence for the role of His-142of protein1C in the acid-induced disassembly of foot-and-mouth disease virus capsids. J GenVirol.,1999,80(Pt8):1911–1918.
68. Escarmís C., Carrillo E. C., Ferrer M., Arriaza J. F., Lopez N., Tami C., Verdaguer N., Domingo E.,Franze-Fernández M. T. Rapid selection in modified BHK-21cells of a foot-and-mouth diseasevirus variant showing alterations in cell tropism. J Virol.,1998,72(12):10171–10179.
69. Escarmís C., Gómez-Mariano G., Dávila M., Lázaro E., Domingo E. Resistance to extinction oflow fitness virus subjected to plaque-to-plaque transfers: diversification by mutation clustering. JMol Biol.,2002,315(4):647–661.
70. Escarmís C., Lázaro E., Manrubia S. C. Population bottlenecks in quasispecies dynamics. Curr TopMicrobiol Immunol.,2006,299:141–170.
71. Escarmís C., Perales C., Domingo E. Biological effect of Muller's Ratchet: distant capsid site canaffect picornavirus protein processing. J Virol.,2009,83(13):6748–6756.
72. Esko J. D., Weinke J. L., Taylor W. H., Ekborg G., Rodén L., Anantharamaiah G., Gawish A.Inhibition of chondroitin and heparan sulfate biosynthesis in Chinese hamster ovary cell mutantsdefective in galactosyltransferase I. J Biol Chem.,1987,262(25):12189–12195.
73. Fauquet C. M., Mayo M. A., Maniloff J., Desselberger U., Ball L. A. Virus taxonomy. VIIIth reportof the international committee on taxonomy of viruses. San Diego, California: Elservier AcademicPress,2005.
74. Feliu J. X., Benito A., Oliva B., Avilés F. X., Villaverde A. Conformational flexibility in a highlymobile protein loop of foot-and-mouth disease virus: distinct structural requirements for integrinand antibody binding. J Mol Biol.,1998,283(2):331–338.
75. Fox G., Parry N. R., Barnett P. V., McGinn B., Rowlands D. J., Brown F. The cell attachment siteon foot-and-mouth disease virus includes the amino acid sequence RGD (arginine-glycine-asparticacid). J Gen Virol.,1989,70(Pt3):625–637.
76. Fry E. E., Lea S. M., Jackson T., Newman J. W., Ellard F. M., Blakemore W. E., Abu-Ghazaleh R.,Samuel A., King A. M., Stuart D. I. The structure and function of a foot-and-mouth diseasevirus-oligosaccharide receptor complex. EMBO J.,1999,18(3):543554.
77. Fry E. E., Newman J. W., Curry S., Najjam S., Jackson T., Blakemore W., Lea S. M., Miller L.,Burman A., King A. M., Stuart D. I. Structure of Foot-and-mouth disease virus serotype A1061alone and complexed with oligosaccharide receptor: receptor conservation in the face of antigenicvariation. J Gen Virol.,2005,86(Pt7):19091920.
78. Gloster J., Jones A., Redington A., Burgin L., S rensen J. H., Turner R., Dillon M., Hullinger P.,Simpson M., Astrup P., Garner G., Stewart P., D'Amours R., Sellers R., Paton D. Airborne spread offoot-and-mouth disease—model intercomparison. Vet J.,2010,183(3):278286.
79. Gloster J., Williams P., Doel C., Esteves I., Coe H., Valarcher J. F. Foot-and-mouthdisease-quantification and size distribution of airborne particles emitted by healthy and infectedpigs. Vet J.,2007,174(1):4253.
80. Grubman M. J. Development of novel strategies to control foot-and-mouth disease: markervaccines and antivirals. Biologicals,2005,33(4):227234.
81. Grubman M. J., Baxt B. Foot-and-mouth disease. Clin Microbiol Rev.,2004,17(2):465493.
82. Guo H., Liu X., Liu Z., Yin H., Ma J., Wang Y., Shang Y., Zhang Q., Li D., Guo J., Lu Z., Xie Q.Recent outbreaks of foot-and-mouth disease type Asia1in China. J Vet Med B Infect Dis VetPublic Health,2006,53(s1):2933.
83. Gutiérrez-Rivas M., Pulido M. R., Baranowski E., Sobrino F., Sáiz M. Tolerance to mutations inthe foot-and-mouth disease virus integrin-binding RGD region is different in cultured cells and invivo and depends on the capsid sequence context. J Gen Virol.,2008,89(Pt10):2531–2539.
84. Harwood L. J., Gerber H., Sobrino F., Summerfield A., McCullough K. C. Dendritic cellinternalization of foot-and-mouth disease virus: influence of heparan sulfate binding on virusuptake and induction of the immune response. J Virol.,2008,82(13):6379–6394.
85. Haydon D. T., Bastos A. D., Awadalla P. Low linkage disequilibrium indicative of recombination infoot-and-mouth disease virus gene sequence alignments. J Gen Virol.,2004,85(Pt5):10951100.
86. He D. S., Li K. N., Lin X. M., Lin S. R., Su D. P., Liao M. Genomic comparison of foot-and-mouthdisease virus R strain and its chick-passaged attenuated strain. Vet Microbiol.,2011,150(1-2):185–190.
87. Hemadri D., Tosh C., Sanyal A., Venkataramanan R. Emergence of a new strain of type Ofoot-and-mouth disease virus: its phylogenetic and evolutionary relationship with the PanAsiapandemic strain. Virus Genes,2002,25(1):2334.
88. Hernández J., Valero M. L., Andreu D., Domingo E., Mateu M. G. Antibody and host cellrecognition of foot-and-mouth disease virus (serotype C) cleaved at the Arg-Gly-Asp (RGD) motif:a structural interpretation. J Gen Virol.,1996,77(Pt2):257–264.
89. Hewat E. A., Verdaguer N., Fita I., Blakemore W., Brookes S., King A., Newman J., Domingo E.,Mateu M. G., Stuart D. I. Structure of the complex of an Fab fragment of a neutralizing antibodywith foot-and-mouth disease virus: positioning of a highly mobile antigenic loop. EMBO J.,1997,16(7):1492–1500.
90. Horsington J., Zhang Z. Consistent change in the B-C loop of VP2observed in foot-and-mouthdisease virus from persistently infected cattle: implications for association with persistence. VirusRes.,2007,125(1):114118.
91. Huang C. C., Lin Y. L., Huang T. S., Tu W. J., Lee S. H., Jong M. H., Lin S. Y. Molecularcharacterization of foot-and-mouth disease virus isolated from ruminants in Taiwan in1999-2000.Vet Microbiol.,2001,81(3):193205.
92. Huang C., Jong M., Lin S. Characteristics of foot and mouth disease virus in Taiwan. J Vet MedSci.,2000,62(7):677679.
93. Hui E. K. Reasons for the increase in emerging and re-emerging viral infectious diseases. MicrobesInfect.,2006,8(3):905916.
94. Jackson A. L., O'Neill H., Maree F., Blignaut B., Carrillo C., Rodriguez L., Haydon D. T. Mosaicstructure of foot-and-mouth disease virus genomes. J Gen Virol.,2007,88(Pt2):487492.
95. Jackson T., Blakemore W., Newman J. W., Knowles N. J., Mould A. P., Humphries M. J., King A.M. Foot-and-mouth disease virus is a ligand for the high-affinity binding conformation of integrinalpha5beta1: influence of the leucine residue within the RGDL motif on selectivity of integrinbinding. J Gen Virol.,2000a,81(Pt5):1383–1391.
96. Jackson T., Clark S., Berryman S., Burman A., Cambier S., Mu D., Nishimura S., King A. M.Integrin alphavbeta8functions as a receptor for foot-and-mouth disease virus: role of thebeta-chain cytodomain in integrin-mediated infection. J Virol.,2004,78(9):4533–4540.
97. Jackson T., Ellard F. M., Ghazaleh R. A., Brookes S. M., Blakemore W. E., Corteyn A. H., Stuart D.I., Newman J. W., King A. M. Efficient infection of cells in culture by type O foot-and-mouthdisease virus requires binding to cell surface heparan sulfate. J Virol.,1996,70(8):5282–5287.
98. Jackson T., King A. M., Stuart D. I., Fry E. Structure and receptor binding. Virus Res.,2003,91(1):3346.
99. Jackson T., Mould A. P., Sheppard D., King A. M. Integrin alphavbeta1is a receptor forfoot-and-mouth disease virus. J Virol.,2002,76(3):935–941.
100.Jackson T., Sharma A., Ghazaleh R. A., Blakemore W. E., Ellard F. M., Simmons D. L., Newman J.W., Stuart D. I., King A. M. Arginine-glycine-aspartic acid-specific binding by foot-and-mouthdisease viruses to the purified integrin alpha(v)beta3in vitro. J Virol.,1997,71(11):8357–8361.
101.Jackson T., Sheppard D., Denyer M., Blakemore W., King A. M. The epithelial integrinalphavbeta6is a receptor for foot-and-mouth disease virus. J Virol.,2000b,74(11):4949–4956.
102.Jamal S. M., Ferrari G., Ahmed S., Normann P., Belsham G. J. Genetic diversity of foot-and-mouthdisease virus serotype O in Pakistan and Afghanistan,1997-2009. Infect Genet Evol.,2011,11(6):12291238.
103.Jin H., Xiao C., Zhao G., Du X., Yu Y., Kang Y., Wang B. Induction of immature dendritic cellapoptosis by foot and mouth disease virus is an integrin receptor mediated event before viralinfection. J Cell Biochem.,2007,102(4):980–991.
104.Jin N. Y., Zhang H. Y., Yin G. F., Zheng M., Liu T., Jiang W. Z., Li Z. J. Immunogenicity ofrecombinant fowl-pox virus co-expressing structural protein precursor P1-2A and proteinase3C ofFMDV. Chinese Sci Bull.,2004,47(8):823827.
105.Jnaoui K., Minet M., Michiels T. Mutations that affect the tropism of DA and GDVII strains ofTheiler's virus in vitro influence sialic acid binding and pathogenicity. J Virol.,2002,76(16):8138–8147.
106.Johns H. L., Berryman S., Monaghan P., Belsham G. J., Jackson T. A dominant-negative mutant ofrab5inhibits infection of cells by foot-and-mouth disease virus: implications for virus entry. JVirol.,2009,83(12):6247–6256.
107.Kitching R. P. Global epidemiology and prospects for control of foot-and-mouth disease. Curr TopMicrobiol Immunol.,2005,288:133148.
108.Kitson J. D., McCahon D., Belsham G. J. Sequence analysis of monoclonal antibody resistantmutants of type O foot and mouth disease virus: evidence for the involvement of the three surfaceexposed capsid proteins in four antigenic sites. Virology,1990,179(1):2634.
109.Klein J. Understanding the molecular epidemiology of foot-and-mouth-disease virus. Infect GenetEvol.,2009,9:153161.
110.Klein J., Hussain M., Ahmad M., Normann P., Afzal M., Alexandersen S. Genetic characterisationof the recent foot-and-mouth disease virus subtype A/IRN/2005. Virol J.,2007,4:122.
111.Knipe T., Rieder E., Baxt B., Ward G., Mason P. W. Characterization of synthetic foot-and-mouthdisease virus provirions separates acid-mediated disassembly from infectivity. J Virol.,1997,71(4):2851–2856.
112.Knowles N. J., Davies P. R., Henry T., O'Donnell V., Pacheco J. M., Mason P. W. Emergence inAsia of foot-and-mouth disease viruses with altered host range: characterization of alterations inthe3A protein. J Virol.,2001,75(3):15511556.
113.Knowles N. J., Samuel A. R. Molecular epidemiology of foot-and-mouth disease virus. Virus Res.,2003,91:6580.
114.Knowles N. J., Samuel A. R., Davies P. R., Midgley R. J., Valarcher J. F. Pandemic strain offoot-and-mouth disease virus serotype O. Emerg Infect Dis.,2005,11(12):18871893.
115.Krebs O., Marquardt O. Identification and characterization of foot-and-mouth disease virus O1Burgwedel/1987as an intertypic recombinant. J Gen Virol.,1992,73(Pt3):613619.
116.Le V. P., Nguyen T., Lee K. N., Ko Y. J., Lee H. S., Nguyen V. C., Mai T. D., Do T. H., Kim S. M.,Cho I. S., Park J. H. Molecular characterization of serotype A foot-and-mouth disease virusescirculating in Vietnam in2009. Vet Microbiol.,2010,144(1-2):5866.
117.Lee K. N., Oem J. K., Park J. H., Kim S. M., Lee S. Y., Tserendorj Sh., Sodnomdarjaa R., Joo Y. S.,Kim H. Evidence of recombination in a new isolate of foot-and-mouth disease virus serotype Asia
1. Virus Res.,2009,139(1):117121.
118.Leippert M., Beck E., Weiland F., Pfaff E. Point mutations within the betaG-betaH loop offoot-and-mouth disease virus O1K affect virus attachment to target cells. J Virol.,1997,71(2):1046–1051.
119.Leippert M., Pfaff E. Foot-and-mouth disease virus can utilize the C-terminal extension ofcoxsackievirus A9VP1for cell infection. J Gen Virol.,2001,82(Pt7):1703–1711.
120.Leister D., Adam K. H., Marquardt O. Co-replication of several isotypes of foot-and-mouth diseasevirus. J Gen Virol.,1993,74(Pt12):2753–2757.
121.Lewis-Rogers N., McClellan D. A., Crandall K. A. The evolution of foot-and-mouth disease virus:impacts of recombination and selection. Infect Genet Evol.,2008,8(6):786798.
122.Li D., Liu Z. X., Bao H. F., Lu Z. J., Guo J. H., Cao Y. M., Li P. H., Bai X. W., Chen Y. L., Xie B.X., Cai X. P., Xie Q. G. The complete genome sequence of foot-and-mouth disease virusO/Akesu/58strain and its some molecular characteristics. Arch Virol.,2007a,152(11):20792085.
123.Li D., Shang Y. J., Liu Z. X., Liu X. T., Cai X. P. Comparisons of the complete genomes of twoChinese isolates of a recent foot-and-mouth disease type Asia1virus. Arch Virol.,2007b,152(9):16991708.
124.Li D., Shang Y. J., Liu Z. X., Liu X. T., Cai X. P. Molecular relationships between type Asia1newstrain from China and type O Panasia strains of foot-and-mouth-disease virus. Virus Genes,2007c,35(2):273279.
125.Li F., Browning G. F., Studdert M. J., Crabb B. S. Equine rhinovirus1is more closely related tofoot-and-mouth disease virus than to other picornaviruses. Proc Natl Acad Sci U S A,1996,93:990995.
126.Li J. Situation and control of FMD in mainland, China.16th OIE Sub-Commission Meeting forFMD Control in South East Asia,1519March2010, Lao PDR: Vientiane.2010.
127.Li P., Lu Z., Bao H., Li D., King D. P., Sun P., Bai X., Cao W., Gubbins S., Chen Y., Xie B., Guo J.,Yin H., Liu Z. In-vitro and in-vivo phenotype of type Asia1foot-and-mouth disease virusesutilizing two non-RGD receptor recognition sites. BMC Microbiol.,2011,11:154.
128.Loeffler F., Frosch P. Summarischer bericht über die ergebnisse der untersuchungen dercommission zur erforschung der maul-und klauenseuche be idem institute fürinfektionskrankheiten in Berlin. Centralbl. Bakteriol.,1897,22:257259.
129.Luna E., Rodríguez-Huete A., Rincón V., Mateo R., Mateu M. G. Systematic study of the geneticresponse of a variable virus to the introduction of deleterious mutations in a functional capsidregion. J Virol.,2009,83(19):10140–10151.
130.Luo M., Vriend G., Kamer G., Minor I., Arnold E., Rossmann M. G., Boege U., Scraba D. G., DukeG. M., Palmenberg A. C. The atomic structure of Mengo virus at3.0resolution. Science,1987,235(4785):182191.
131.Mahy B. W. J. Foot-and-mouth disease virus. Verlag Berlin Heidelberg: Springer,2005.
132.Malirat V., Bergmann I. E., Campos R. M., Salgado G., Sánchez C., Conde F., Quiroga J. L., OrtizS. Phylogenetic analysis of Foot-and-Mouth Disease Virus type O circulating in the Andean regionof South America during2002-2008. Vet Microbiol.,2011,152(1-2):7487.
133.Malirat V., Bergmann I. E., de Mendon a Campos R., Conde F., Quiroga J. L., Villamil M.,Salgado G., Ortiz S. Molecular epidemiology of foot-and-mouth disease virus type A in SouthAmerica. Vet Microbiol.,2012,[Epub ahead of print].
134.Maree F. F., Blignaut B., Aschenbrenner L., Burrage T., Rieder E. Analysis of SAT1typefoot-and-mouth disease virus capsid proteins: influence of receptor usage on the properties of virusparticles. Virus Res.,2011,155(2):462–472.
135.Maree F. F., Blignaut B., de Beer T. A., Visser N., Rieder E. A. Mapping of amino acid residuesresponsible for adhesion of cell culture-adapted foot-and-mouth disease SAT type viruses. VirusRes.,2010,153(1):82–91.
136.Martín M. J., Nú ez J. I., Sobrino F., Dopazo J. A procedure for detecting selection in highlyvariable viral genomes: evidence of positive selection in antigenic regions of capsid protein VP1offoot-and-mouth disease virus. J Virol Methods,1998,74(2):215221.
137.Martín-Acebes M. A., González-Magaldi M., Sandvig K., Sobrino F., Armas-Portela R. Productiveentry of type C foot-and-mouth disease virus into susceptible cultured cells requires clathrin and isdependent on the presence of plasma membrane cholesterol. Virology,2007,369(1):105–118.
138.Martín-Acebes M. A., González-Magaldi M., Vázquez-Calvo A., Armas-Portela R., Sobrino F.Internalization of swine vesicular disease virus into cultured cells: a comparative study withfoot-and-mouth disease virus. J Virol.,2009,83(9):4216–4226.
139.Martín-Acebes M. A., Rincón V., Armas-Portela R., Mateu M. G., Sobrino F. A single amino acidsubstitution in the capsid of foot-and-mouth disease virus can increase acid lability and conferresistance to acid-dependent uncoating inhibition. J Virol.,2010,84(6):2902–2912.
140.Martín-Acebes M. A., Vázquez-Calvo A., González-Magaldi M., Sobrino F. Foot-and-mouthdisease virus particles inactivated with binary ethylenimine are efficiently internalized into culturedcells. Vaccine,2011a,29(52):9655–9662.
141.Martín-Acebes M. A., Vázquez-Calvo A., Rincón V., Mateu M. G., Sobrino F. A single amino acidsubstitution in the capsid of foot-and-mouth disease virus can increase acid resistance. J Virol.,2011b,85(6):2733–2740.
142.Martínez M. A., Verdaguer N., Mateu M. G., Domingo E. Evolution subverting essentiality:dispensability of the cell attachment Arg-Gly-Asp motif in multiply passaged foot-and-mouthdisease virus. Proc Natl Acad Sci U S A,1997,94(13):6798–6802.
143.Martinsen J. S. Heterogeneity of foot-and-mouth disease virus: further studies on plaque formationby two plaque-size variants. Can J Comp Med.,1972,36(1):26–33.
144.Mason P. W., Grubman M. J., Baxt B. Molecular basis of pathogenesis of FMDV. Virus Res.,2003,91(1):932.
145.Mason P. W., Rieder E., Baxt B. RGD sequence of foot-and-mouth disease virus is essential forinfecting cells via the natural receptor but can be bypassed by an antibody-dependent enhancementpathway. Proc Natl Acad Sci U S A,1994,91(5):1932–1936.
146.Mateo R., Luna E., Mateu M. G. Thermostable variants are not generally represented infoot-and-mouth disease virus quasispecies. J Gen Virol.,2007,88(Pt3):859–864.
147.Mateo R., Mateu M. G. Deterministic, compensatory mutational events in the capsid offoot-and-mouth disease virus in response to the introduction of mutations found in viruses frompersistent infections. J Virol.,2007,81(4):1879–1887.
148.Mateu M. G. Antibody recognition of picornaviruses and escape from neutralization: a structureview. Virus Res.,1995,38:124.
149.Mateu M. G., Martínez M. A., Capucci L., Andreu D., Giralt E., Sobrino F., Brocchi E., Domingo E.A single amino acid substitution affects multiple overlapping epitopes in the major antigenic site offoot-and-mouth disease virus of serotype C. J Gen Virol.,1990,71(Pt3):629637.
150.Mateu M. G., Valero M. L., Andreu D., Domingo E. Systematic replacement of amino acid residueswithin an Arg-Gly-Asp-containing loop of foot-and-mouth disease virus and effect on cellrecognition. J Biol Chem.,1996,271(22):12814–12819.
151.McCahon D., Slade W. R., King A. M., Saunders K., Pullen L., Lake J. R., Priston R. A. Effect ofmutation on the virulence in mice of a strain of foot-and-mouth disease virus. J Gen Virol.,1981,54(Pt2):263–272.
152.McKenna T. S., Lubroth J., Rieder E., Baxt B., Mason P. W. Receptor binding site-deletedfoot-and-mouth disease (FMD) virus protects cattle from FMD. J Virol.,1995,69(9):57875790.
153.McVicar J. W., Sutmoller P. Three variants of foot-and-mouth disease virus type O: agar geldiffusion reactions. Am J Vet Res.,1972,33(8):16351639.
154.Miller L. C., Blakemore W., Sheppard D., Atakilit A., King A. M., Jackson T. Role of thecytoplasmic domain of the beta-subunit of integrin alpha(v)beta6in infection by foot-and-mouthdisease virus. J Virol.,2001,75(9):4158–4164.
155.Monaghan P., Gold S., Simpson J., Zhang Z., Weinreb P. H., Violette S. M., Alexandersen S.,Jackson T. The alpha(v)beta6integrin receptor for foot-and-mouth disease virus is expressedconstitutively on the epithelial cells targeted in cattle. J Gen Virol.,2005,86(Pt10):2769–2780.
156.Moore D. M., Cowan K. M. Effect of trypsin and chymotrypsin on the polypeptides of large andsmall plaque variants of foot-and-mouth disease virus: relationship to specific antigenicity andinfectivity. J Gen Virol.,1978,41(3):549–562.
157.Morioka K., Fukai K., Ohashi S., Sakamoto K., Tsuda T., Yoshida K. Comparison of the charactersof the plaque-purified viruses from foot-and-mouth disease virus O/JPN/2000. J Vet Med Sci.,2008,70(7):653–658.
158.Murphy F. A., Gibbs E., Horzinek M. C., Studdert M. J. Veterinary virology.3rd. San Diego,California: Academic Press,1999.
159.Neff S., Baxt B. The ability of integrin alpha(v)beta(3) to function as a receptor for foot-and-mouthdisease virus is not dependent on the presence of complete subunit cytoplasmic domains. J Virol.,2001,5(1):527–532.
160.Neff S., Mason P. W., Baxt B. High-efficiency utilization of the bovine integrin alpha(v)beta(3) asa receptor for foot-and-mouth disease virus is dependent on the bovine beta(3) subunit. J Virol.,2000,74(16):7298–7306.
161.Neff S., Sá-Carvalho D., Rieder E., Mason P. W., Blystone S. D., Brown E. J., Baxt B.Foot-and-mouth disease virus virulent for cattle utilizes the integrin alpha(v)beta3as its receptor. JVirol.,1998,72(5):3587–3594.
162.Nilsson E. C., Jamshidi F., Johansson S. M., Oberste M. S., Arnberg N. Sialic acid is a cellularreceptor for coxsackievirus A24variant, an emerging virus with pandemic potential. J Virol.,2008,82(6):3061–3068.
163.Nobiron I., Rémond M., Kaiser C., Lebreton F., Zientara S., Delmas B. The nucleotide sequence offoot-and-mouth disease virus O/FRA/1/2001and comparison with its British parental strainO/UKG/35/2001. Virus Res.,2005,108(1-2):225229.
164.Norkin L. C. Virus receptors: implications for pathogenesis and the design of antiviral agents. ClinMicrobiol Rev.,1995,8(2):293315.
165.Novella I. S., Borrego B., Mateu M. G., Domingo E., Giralt E., Andreu D. Use of substituted andtandem-repeated peptides to probe the relevance of the highly conserved RGD tripeptide in theimmune response against foot-and-mouth disease virus. FEBS Lett.,1993,330(3):253259.
166.Nú ez J. I., Martín M. J., Piccone M. E., Carrillo E., Palma E. L., Dopazo J., Sobrino F.Identification of optimal regions for phylogenetic studies on VP1gene of foot-and-mouth diseasevirus: analysis of types A and O Argentinean viruses. Vet Res.,2001,32(1):31–45.
167.Nú ez J. I., Molina N., Baranowski E., Domingo E., Clark S., Burman A., Berryman S., Jackson T.,Sobrino F. Guinea pig-adapted foot-and-mouth disease virus with altered receptor recognition canproductively infect a natural host. J Virol.,2007,81(16):8497–8506.
168.O'Donnell V., Larocco M., Baxt B. Heparan sulfate-binding foot-and-mouth disease virus enterscells via caveola-mediated endocytosis. J Virol.,2008,82(18):9075–9085.
169.O'Donnell V., LaRocco M., Duque H., Baxt B. Analysis of foot-and-mouth disease virusinternalization events in cultured cells. J Virol.,2005,79(13):8506–8518.
170.O'Donnell V., Pacheco J. M., Gregg D., Baxt B. Analysis of foot-and-mouth disease virus integrinreceptor expression in tissues from na ve and infected cattle. J Comp Pathol.,2009,141(2-3):98–112.
171.O'Donnell V., Pacheco J. M., LaRocco M., Burrage T., Jackson W., Rodriguez L. L., Borca M. V.,Baxt B. Foot-and-mouth disease virus utilizes an autophagic pathway during viral replication.Virology,2011,410(1):142–150.
172.Oem J. K., Yeh M. T., McKenna T. S., Hayes J. R., Rieder E., Giuffre A. C., Robida J. M., Lee K.N., Cho I. S., Fang X., Joo Y. S., Park J. H. Pathogenic characteristics of the Korean2002isolate offoot-and-mouth disease virus serotype O in pigs and cattle. J Comp Pathol.,2008,138(4):204–214.
173.Office International des Epizooties. Handistatus II. In: International Office of Epizootics,2006.
174.Office International des Epizooties. O.I.E Bull.,1999, N°3:202.
175.Pacheco J. M., Arzt J., Rodriguez L. L. Early events in the pathogenesis of foot-and-mouth diseasein cattle after controlled aerosol exposure. Vet J.,2010,183(1):4653.
176.Pacheco J. M., Mason P. W. Evaluation of infectivity and transmission of different Asianfoot-and-mouth disease viruses in swine. J Vet Sci.,2010,11(2):133–142.
177.Parry N., Fox G., Rowlands D., Brown F., Fry E., Acharya R., Logan D., Stuart D. Structural andserological evidence for a novel mechanism of antigenic variation in foot-and-mouth disease virus.Nature,1990,347(6293):569572.
178.Paton D. J., King D. P., Knowles N. J., Hammond J. Recent spread of foot-and-mouth disease inthe Far East. Vet Rec.,2010,166(18):569570.
179.Pfaff M. Recognition sites of RGD-dependent integrins. In: Eble J. A., Kühn K., eds.Intergrin-ligand interaction. Heidelberg: Springer-Verlag,1997,101121.
180.Piccone M. E., Pacheco J. M., Pauszek S. J., Kramer E., Rieder E., Borca M. V., Rodriguez L. L.The region between the two polyprotein initiation codons of foot-and-mouth disease virus iscritical for virulence in cattle. Virology,2010,396(1):152–159.
181.Rieder E., Baxt B., Mason P. W. Animal-derived antigenic variants of foot-and-mouth disease virustype A12have low affinity for cells in culture. J Virol.,1994,68(8):5296–5299.
182.Rieder E., Berinstein A., Baxt B., Kang A., Mason P. W. Propagation of an attenuated virus bydesign: engineering a novel receptor for a noninfectious foot-and-mouth disease virus. Proc NatlAcad Sci U S A,1996,93(19):10428–10433.
183.Rieder E., Henry T., Duque H., Baxt B. Analysis of a foot-and-mouth disease virus type A24isolate containing an SGD receptor recognition site in vitro and its pathogenesis in cattle. J Virol.,2005,79(20):12989–12998.
184.Robinson L., Windsor M., McLaughlin K., Hope J., Jackson T., Charleston B. Foot-and-mouthdisease virus exhibits an altered tropism in the presence of specific immunoglobulins, enablingproductive infection and killing of dendritic cells. J Virol.,2011,85(5):2212–2223.
185.Rojas E. R., Carrillo E., Schiappacassi M., Campos R. Modification of foot-and-mouth diseasevirus O1Caseros after serial passages in the presence of antiviral polyclonal sera. J Virol.,1992,66(6):33683372.
186.Ruiz-Jarabo C. M., Arias A., Baranowski E., Escarmís C., Domingo E. Memory in viralquasispecies. J Virol.,2000,74(8):35433547.
187.Ruíz-Jarabo C. M., Arias A., Molina-París C., Briones C., Baranowski E., Escarmís C., Domingo E.Duration and fitness dependence of quasispecies memory. J Mol Biol.,2002,315(3):285–296.
188.Ruiz-Jarabo C. M., Sevilla N., Dávila M., Gómez-Mariano G., Baranowski E., Domingo E.Antigenic properties and population stability of a foot-and-mouth disease virus with an alteredArg-Gly-Asp receptor-recognition motif. J Gen Virol.,1999,80(Pt8):1899–1909.
189.Ruiz-Sáenz J., Goez Y., Tabares W., López-Herrera A. Cellular receptors for foot and mouthdisease virus. Intervirology,2009,52(4):201–212.
190.Sa-Carvalho D., Rieder E., Baxt B., Rodarte R., Tanuri A., Mason P. W. Tissue culture adaptationof foot-and-mouth disease virus selects viruses that bind to heparin and are attenuated in cattle. JVirol.,1997,71(7):5115–5123.
191.Saitou N., Nei M. The neighbor-joining method: a new method for reconstructing phylogenetictrees. Mol Biol Evol.,1987,4(4):406425.
192.Sáiz M., Gómez S., Martínez-Salas E., Sobrino F. Deletion or substitution of the aphthovirus3'NCR abrogates infectivity and virus replication. J Gen Virol.,2001,82(Pt1):93101.
193.Sakamoto K., Yoshida K. Recent outbreaks of foot and mouth disease in countries of East Asia.Rev Sci Tech.,2002,21(3):459463.
194.Sanz-Ramos M., Díaz-San Segundo F., Escarmís C., Domingo E., Sevilla N. Hidden virulencedeterminants in a viral quasispecies in vivo. J Virol.2008,82(21):10465–10476.
195.Schneider-Schaulies J. Cellular receptors for viruses: links to tropism and pathogenesis. J GenVirol.,2000,81(Pt6):14131429.
196.Schols D. HIV co-receptor inhibitors as novel class of anti-HIV drugs. Antiviral Res.,2006,71(2-3):216226.
197.Schumann K. R., Knowles N. J., Davies P. R., Midgley R. J., Valarcher J. F., Raoufi A. Q.,McKenna T. S., Hurtle W., Burans J. P., Martin B. M., Rodriguez L. L., Beckham T. R. Geneticcharacterization and molecular epidemiology of foot-and-mouth disease viruses isolated formAfghanistan in2003-2005. Virus Genes,2008,36(2):401413.
198.Semler B. L., Wimmer E. Molecular Biology of Picornaviruses. Washington, DC: AmericanSociety for Microbiology Press,2002.
199.Sevilla N., Domingo E. Evolution of a persistent aphthovirus in cytolytic infections: partialreversion of phenotypic traits accompanied by genetic diversification. J Virol.,1996,70(10):6617–6624.
200.Sevilla N., Ruiz-Jarabo C. M., Gómez-Mariano G., Baranowski E., Domingo E. An RNA virus canadapt to the multiplicity of infection. J Gen Virol.,1998,79(Pt12):2971–2980.
201.Sharma A., Rao Z., Fry E., Booth T., Jones E. Y., Rowlands D. J., Simmons D. L., Stuart D. I.Specific interactions between human integrin alpha v beta3and chimeric hepatitis B virus coreparticles bearing the receptor-binding epitope of foot-and-mouth disease virus. Virology,1997,239(1):150–157.
202.Sieczkarski S. B., Whittaker G. R. Viral entry. Curr Top Microbiol Immunol.,2005,285:123.
203.Sobrino F., Domingo E. Foot and mouth disease: current perspectives. Norfolk: HorizonBioscience,2004.
204.Sobrino F., Sáiz M., Jiménez-Clavero M. A., Nú ez J. I., Rosas M. F., Baranowski E., Ley V.Foot-and-mouth disease virus: a long known virus, but a current threat. Vet Res.,2001,32(1):130.
205.Storey P., Theron J., Maree F. F., O'Neill H. G. A second RGD motif in the1D capsid protein of aSAT1type foot-and-mouth disease virus field isolate is not essential for attachment to target cells.Virus Res.,2007,124(1-2):184–192.
206.Tami C., Taboga O., Berinstein A., Nú ez J. I., Palma E. L., Domingo E., Sobrino F., Carrillo E.Evidence of the coevolution of antigenicity and host cell tropism of foot-and-mouth disease virusin vivo. J Virol.,2003,77(2):1219–1226.
207.Thompson D., Muriel P., Russell D., Osborne P., Bromley A., Rowland M., Creigh-Tyte S., BrownC. Economic costs of the foot and mouth disease outbreak in the United Kingdom in2001. Rev SciTech.,2002,21(3):675687.
208.Tosh C., Hemadri D., Sanyal A. Evidence of recombination in the capsid-coding region of type Afoot-and-mouth disease virus. J Gen Virol.,2002,83(Pt10):24552460.
209.Tuthill T. J., Harlos K., Walter T. S., Knowles N. J., Groppelli E., Rowlands D. J., Stuart D. I., FryE. E. Equine rhinitis A virus and its low pH empty particle: clues towards an aphthovirus entrymechanism? PLoS Pathog.,2009,5(10): e1000620.
210.Twomey T., France L. L., Hassard S., Burrage T. G., Newman J. F., Brown F. Characterization ofan acid-resistant mutant of foot-and-mouth disease virus. Virology,1995,206(1):69–75.
211.Valarcher J. F., Knowles N. J., Ferris N. P., Paton D. J., Zakharov V., Sherbakov A., Shang Y. J.,Liu Z. X., Liu X. T., Sanyal A., Hemadri D., Tosh C., Rasool T. J. Recent spread of FMD virusserotype Asia1. Vet Rec.,2005,157(1):30.
212.Valarcher J. F., Knowles N. J., Fernandez R., Davies P. R., Midgley R. J., Hutchings G., Newman B.J., Ferris N. P., Paton D. J. FMD global update2003-2004. In: Open sessions of the EUFMDresearch group. Greece: Chaina,2004.
213.van Rensburg H. G., Henry T. M., Mason P. W. Studies of genetically defined chimeras of aEuropean type A virus and a South African Territories type2virus reveal growth determinants forfoot-and-mouth disease virus. J Gen Virol.,2004,85(Pt1):6168.
214.van Vlijmen H. W., Curry S., Schaefer M., Karplus M. Titration calculations of foot-and-mouthdisease virus capsids and their stabilities as a function of pH. J Mol Biol.,1998,275(2):295–308.
215.Verdaguer N., Mateu M. G., Andreu D., Giralt E., Domingo E., Fita I. Structure of the majorantigenic loop of foot-and-mouth disease virus complexed with a neutralizing antibody: directinvolvement of the Arg-Gly-Asp motif in the interaction. EMBO J.,1995,14(8):16901696.
216.Verdaguer N., Sevilla N., Valero M. L., Stuart D., Brocchi E., Andreu D., Giralt E., Domingo E.,Mateu M. G., Fita I. A similar pattern of interaction for different antibodies with a major antigenicsite of foot-and-mouth disease virus: implications for intratypic antigenic variation. J Virol.,1998,72(1):739748.
217.Vlodavsky I., Ilan N., Naggi A., Casu B. Heparanase: structure, biological functions, and inhibitionby heparin-derived mimetics of heparan sulfate. Curr Pharm Des.,2007,13(20):20572073.
218.Wang H., Zhao L., Li W., Zhou G., Yu L. Identification of a conformational epitope on the VP1G-H Loop of type Asia1foot-and-mouth disease virus defined by a protective monoclonal antibody.Vet Microbiol.,2011,148(2-4):189199.
219.Wilke C. O., Novella I. S. Phenotypic mixing and hiding may contribute to memory in viralquasispecies. BMC Microbiol.,2003,3:11.
220.Woodbury E. L., Samuel A. R., Knowles N. J., Hafez S. M., Kitching R. P. Analysis of mixedfoot-and-mouth disease virus infections in Saudi Arabia: prolonged circulation of an exoticserotype. Epidemiol Infect.,1994,112(1):201211.
221.Yang D., Zhang C., Zhao L., Zhou G., Wang H., Yu L. Identification of a conserved linear epitopeon the VP1protein of serotype O foot-and-mouth disease virus by neutralising monoclonalantibody8E8. Virus Res.,2011,155(1):291299.
222.Yang X., Zhou Y. S., Wang H. N., Zhang Y., Wei K., Wang T. Isolation, identification and completegenome sequence analysis of a strain of foot-and-mouth disease virus serotype Asia1from pigs insouthwest of China. Virol J.,2011,8:175.
223.Yi J., Liu Ch. Integrity of foot-and-mouth disease virions bound to the cells. Acta Virol.,2010,54(4):307–310.
224.Yoon S. H., Park W., King D. P., Kim H. Phylogenomics and molecular evolution offoot-and-mouth disease virus. Mol Cells,2011,31(5):413421.
225.Zhang L., David G., Esko J. D. Repetitive Ser-Gly sequences enhance heparan sulfate assembly inproteoglycans. J Biol Chem.,1995,270(45):27127–27135.
226.Zhang X. S., Liu Z. X., Zhao Q. Z., Chang H. Y., Xie Q. G. Sequencing and analysis for thefull-length genome RNA of foot-and-mouth disease virus China/99. Chinese Sci Bull.,2004,47(1):7481.
227.Zhao M. Q., Suo Q. L., Chen J. D., Chen L. J., Xu Y. F. Sequence analysis of the protein-codingregions of foot-and-mouth disease virus O/HK/2001. Vet Microbiol.,2008,130(3-4):238246.
228.Zhao Q., Pacheco J. M., Mason P. W. Evaluation of genetically engineered derivatives of a Chinesestrain of foot-and-mouth disease virus reveals a novel cell-binding site which functions in cellculture and in animals. J Virol.,2003,77(5):3269–3280.
229.Zheng H. X., Guo J. H., Jin Y., Shang Y. J., Tian H., Yang Y. M., Liu X. T., Cai X. P. Infectivity ofAsia1type foot-and-mouth disease virus was increased via an alternative RDD motif. Chinese SciBull.,2010,55:13701375.
230.Zheng H. X., He J. J., Guo J. H., Jin Y., Yang F., Lv L., Liu X. T. Genetic characterization of a newpandemic Southeast Asia topotype strain of serotype O foot-and-mouth disease virus isolated inChina during2010. Virus Genes,2012,44(1):8088.
231.Zhong J., Li Y., Zhao S., Liu S., Zhang Z. Mutation pressure shapes codon usage in the GC-Richgenome of foot-and-mouth disease virus. Virus Genes,2007,35(3):767776.
232.Zhou J. H., Zhang J., Chen H. T., Ma L. N., Ding Y. Z., Pejsak Z., Liu Y. S. The codon usage modelof the context flanking each cleavage site in the polyprotein of foot-and-mouth disease virus. InfectGenet Evol.,2011,11(7):18151819.
233.Zhou J. H., Zhang J., Chen H. T., Ma L. N., Liu Y. S. Analysis of synonymous codon usage infoot-and-mouth disease virus. Vet Res Commun.,2010a,34:393404.
234.Zhou J. H., Zhang J., Ding Y. Z., Chen H. T., Ma L. N., Liu Y. S. Characteristics of codon usagebias in two regions downstream of the initiation codons of foot-and-mouth disease virus.BioSystems,2010b,101:2028.