PATENT DOCUMENT

Abstract:
A determination of the nucleotide sequence for the genome of Equine rhinovirus 1 (ERhV1), a horse-respiratory pathogen of heretofor uncertain taxonomic status, reveals a structure and predicted amino acid sequence that are more similar to those of foot-and-mouth disease viruses, the only members of the aphthovirus genus, than of other picornaviruses. These insights inform an understanding of ERhV1 virion proteins and virus-like particles, as well as the design of probes, primers, antigens, vectors, diagnostics and tests of relevance to ERhV1.

Full Description:
This application is a divisional of application Ser. No. 09/091,219, filed Oct. 5, 1998, now U.S. Pat. No. 6,171,592, which is a National Stage of PCT/AU06/00815, filed Dec. 18, 1996, based on Australian Application No. PN 7201, filed Dec. 18, 1995. 
    
    
     INTRODUCTION TO INVENTION 
     This invention relates to the equine rhinovirus 1 (ERhV1) which has been sequenced and characterized. In particular, the invention relates to nucleotide and protein sequences of ERhV1 and a range of clinical and diagnostic products derived from ERhV1. 
     BACKGROUND OF INVENTION 
     Equine rhinovuris 1 (ERhV1) was first isolated from horses in the United Kingdom and subsequently from horses in mainland Europe, the USA and Australia. Most isolates were from the nasopharynx of horses with an acute, febrile respiratory disease. Virions had the characteristic size and morphology of picornaviruses and were acid-labile. Two other serologically distinct, acid-labile picornaviruses, ERhV2 and ERhV3, have also been isolated from horses. 
     Considerable uncertainity has surrounded the classification of ERhV1. Physicochemical studies have shown that the nucleic acid density and base composition of ERhV1 differ from those of rhinoviruses. In contrast to rhinoviruses, ERhV1 has a broad host-cell range in vitro and in vivo and there is no evidence of extensive antigenic variation. Infection of horses with ERhV1 causes a disease characterized by an acute febrile respiratory disease accompanied by anemia, fecal and urine shredding and viral persistance. The signs of systemic infection and persistance are not characteristic of rhinovirus infections in other species. The known host range of ERhV1 is broad and includes rabbits, guinea pigs, monkeys and humans, although in these species the virus does not appear to spread horizontally. There is both experimental and epidemiological evidence of ERhV1 infection of humans. A human volunteer inoculated intranasally with ERhV1 developed severe pharyngitis, lymphadenities, fever and viremia, and high ERhV1 antibody titers were found in the sera of 3 of 12 stable workers whereas no ERhV1 antibody was found in the sera of 159 non-stable workers. 
     In order to clarify the taxonomic status of ERhV1, a detailed study was undertaken to determine the nucleotide and amino acid sequence of ERhV1. The resultant studies provided the complete nucleotide sequence of the gene encoding the ERhV1 polyprotein and the 3′-nontranslated region (NTR) as well as part of the nucleotide sequence of the 5′NTR. The amino acid sequence of the various ERhV1 proteins was deduced from the nucleotide sequence. 
     The analysis of the nucleotide sequence of ERhV1 confirmed previous studies which indicated that many properties of ERhV1 are not consistent with those of other members of the genus Rhinovirus. Indeed many of the physiochemical and biological properties of ERhV1 have suggested ERhV1 is more closely related to foot-and-mouth disease virus (FMDV) the sole member of the Alpthovirus genus. In addition to the overall sequence similarity, several features of the ERhV1 genome are similar to those of FMDV. The ERhV1 L protein is most similar to its counterpart in aphthoviruses in both length, 207 amino acids in ERhV1 and 201 and FMDV, and in amino acid sequence identity. In aphthoviruses, the L protein catalyses its own cleavage from the polyprotein, and mediates cleavage of the p220 component of the cap-binding complex leading to inhibition of translation of capped mRNAs. Cardiovirus L proteins are only 67-76 amino acids long and are not auto catalytic. In contrast to the cardioviruses, aphthoviruses utilize two distinct initiation codons, which results in different forms of the L protein, Lab and Lb, differing from each other by 28 amino acids at their N-termini. 
     The second initiation codon occurs in a more favourable contexts, which is presumably the reason why Lb, the smaller of the two proteins, is the predominant species. Thus far, differences in the function of the two FMDV L proteins have not been detected. ERhV1 also possesses a second ATG, 63 bases downstream from the first optimal ATG, which is also present in a context optimal for initiation of translation. Translation from this ATG would result in an L protein with 21 fewer amino acids at its N-terminus. Therefore, it is probable that ERhV1 possesses a second species of L protein similar to the FMDV Lb protein. If so, the reason for the existence and conservation of two forms of the L protein in ERhV1 and FMDV is an intriguing question. Curiously, ERhV1 has tandemly repeated ATG codons at each of the possible initiation sites, where the first ATG in each case does not occur in a context optimal for translation. The role of these ATGs may be to ensure that translation is initiated from both possible initiation sites. 
     The 2A protease is only 16 amino acids in length in both FMDV and ERhV1, compared to 142-149 amino acids in other picornaviruses. In FMDV 2A protease cleaves at its C-terminus but, unlike the 2A protease of other picornaviruses, appears not to have a role in shut down of host cell macromolecular synthesis. The high degree of conservation of the FMDV and ERhV1 2A proteins is intriguing and suggests an important role for this protein in the diseases produced by these viruses. 
     It may be expected that the tree derived from the complete polyprotein coding sequence would provide the most representative view of the taxonomic status of ERhV1 by reducing any bias imparted by using restricted parts of the genome with highly variable evolutionary rates. However, such analysis is restricted because there are only a few complete polyprotein sequences available. The polymerase genes are the most conserved genes in positive strand RNA viruses and they have been used to construct a taxonomy, and to predict the ancient roots, of these viruses. In contrast to the polymerase gene, the VPI gene encodes the major antigenic determinants of the virus and evolves more rapidly than other regions in the genome. The diversity of VP1 regions make them useful for the study of closely related picornaviruses. Thus, trees based on the polymerase and VP1 genes presumably reflect the extremes of evolutionary rates from which the taxonomic status and evolutionary origin of ERhV1 could be identified. The ERhV1 VP1 amino acid sequence was more similar to FMDV that to any other sequence in the data base; this was true even when representative segments across the entire sequence were separately analysed. 
     Therefore, we consider that the difference in the topology of the VP1, compared to the other two trees, is most unlikely to be a consequence of genetic recombination. The topographic differences between the three ERhV1 trees compared to those of aphthoviruses, particularly the VP1 derived trees, as well as the presence of only one VPg gene in ERhV1 genome, leads us to conclude that ERhV1 is probably a member of a distinct genus proposed to be called Equirhinovirus. 
     The reassessment of the taxonomic status of ERhV1 focuses on a requirement to reassess the biology of the virus particularly with respect to the nature of clinical disease as well as means for control by vaccination and improved methods of diagnosis. For example, cardioviruses and aphthoviruses cause viremic infections accompanied by myocarditis. Clinical disease caused by ERhV1 is generally considered to be confirmed to the respiratory tract even though there is a viremia and the virus is shed in faces and urine. Whether ERhV1 infection produces systemic disease similar to that observed in apthovirus or cardiovirus infections, including the production of myocarditis, needs to be investigated. There is serological evidence that the incidence of ERhV1 infection is as high as 50% in some horse populations however, the number of reported isolations of ERhV1 is very small. We have clear evidence that primary isolation of the virus from clinical specimens is known to be difficult, suggesting that the true incidence of ERhV1 disease is much greater than reported. 
     The determination of the complete nucleotide sequence of ERhV1 polyprotein has important practical applications in developing novel methods for the diagnosis and control of ERhV1 disease in horses and other species. 
     OBJECT AND STATEMENT OF INVENTION 
     In one aspect, the invention provides a substantially pure nucleotide sequence for ERhV1 being: 
     
       
         
               
               
               
             
           
               
                 a substantially pure nucleotide 
                   
                   
               
               
                 sequence for ERhVI (SEQ ID NO: 1) being: 
               
               
                   
               
               
                 CCGTCAAGCC CGTTGCCTGT ATAGCCAGGT AACCGGACAG CGGCTTGCTG GATTTTCCCG 
                 −375 
               
               
                   
               
               
                 GTGCCATTGC TCTGGATGCT GTCACCAAGC TGACAAATGC GGAGTGAACC TCACAAAGCG 
                 −315 
               
               
                   
               
               
                 ACACGCCTGT GGTAGCGCTG CCCAAAAGGG AGCGGAACTC CCCGCCGAGG CGGTCCTCTC 
                 −255 
               
               
                   
               
               
                 TGGCCAAAAG CCCAGCGTTG ATAGCGCCTT TTGGGATGCA GGAACCCCAC CTGCCAGGTG 
                 −195 
               
               
                   
               
               
                 TGAAGTGGAG TGAGCGGATC TCCAATTTGG TCTGTTCTGA ACTACACCAT TTACTGCTGT 
                 −135 
               
               
                   
               
               
                 GAAGAATGCC CTGGAGGCAA GCTGGTTACA GCCCTGACCA GGCCCTGCCC GTGACTCTCG 
                 −75 
               
               
                   
               
               
                 ACCGGCGCAG GGTCAAAAAT TGTCTAAGCA GCAGCAGGAA CGCGGGAGCG  TTTCTTTTCC   
                 −15 
               
               
                   
               
               
                   TTTT GTACTG AC ATGATG GC GGCGTCTAAG GTGTATAGAG TTTGCGAGCA GACTCTGCTG 
                 45 
               
               
                   
               
               
                 GCAGGTGCCG TTCGC ATGAT   G GACAAATTC TTGCAAAAGA GAACTGTTTT TGTCCCCCAT 
                 105 
               
               
                   
               
               
                 CTTGACAAAA CAATTCGTTT GACTGGACTC CACAATTATG ACAATACTTG CTGGTTGAAT 
                 165 
               
               
                   
               
               
                 GCCTTGACAC AACTGACACA GATTCTTGGA ATTCGGCTTT TTGATGAACA CTTCGGCAAT 
                 225 
               
               
                   
               
               
                 AGAGGTCTGT TCACTCGGAA AACAATTGAT TGGGTGAGTG ACCAGACTGG TATAAAAGAT 
                 285 
               
               
                   
               
               
                 CTAAAATCAG GAGCACCGCC ACTCGTGGTG GTGTACAAAC TGTGGCAACA TGGACACTTG 
                 345 
               
               
                   
               
               
                 GATGTCGGTA CGATGGAGAA ACCCCGGTCG ATTACTCTAT GGTCTGGCCC CAAAGTGTGT 
                 405 
               
               
                   
               
               
                 CTTTCTGATT TCTGGGCCTG TGTTTCGGCA AAACCGGGAC ATGCAGTATT CTACCTTCTC 
                 465 
               
               
                   
               
               
                 ACAAGCGAGG GTTGGATCTG TGTTGATGAC AAGAAAATAT ACCCAGAAAC ACCCAAAACA 
                 525 
               
               
                   
               
               
                 GAGGATGTAC TTGTTTTTGC GCCCTATGAC TTTGAGTCAC TGGGCAAGGA CCCACCAAAG 
                 585 
               
               
                   
               
               
                 CTACACCAGA GATATGAAAA AGCATTTGAG CTCAGTGGCG GAGGTACATC CACTCCAACA 
                 645 
               
               
                   
               
               
                 ACTGGCAACC AAAACATGTC CGGAAACAGT GGTTCAATTG TTCAAAATTT TTACATGCAA 
                 705 
               
               
                   
               
               
                 CAGTACCAGA ATTCAATTGA CGCAGACCTC GGAGACAATG TGATTAGCCC TGAAGGCCAG 
                 765 
               
               
                   
               
               
                 GGCAGCAACA CTAGTAGTTC AACCTCATCA AGCCAATCCT CTGGCTTGGG CGGGTGGTTC 
                 825 
               
               
                   
               
               
                 TCTAGTTTGC TGAACCTTGG AACAAAACTA CTGGCTGACA AGAAGACAGA AGAGACTACA 
                 885 
               
               
                   
               
               
                 AACATTGAAG ACAGAATTGA AACAACAGTG GTTGGAGTCA CTATTATTAA TTCACAAGGA 
                 945 
               
               
                   
               
               
                 TCTGTTGGAA CAACCTACTG TTACTCCAAA CCGGATGGTA GACCACCATC CACAGTGTCA 
                 1005 
               
               
                   
               
               
                 GACCCAGTTA CCAGACTTTG ACCCACGCTT TCCAGGCACT ACACATTTAA GGTAGGTGAG 
                 1065 
               
               
                   
               
               
                 TGGCCCCATT CTCAATCACA TGGTCACGCA TGGATCTGTC CGTTGCCAGG TGACAAACTC 
                 1125 
               
               
                   
               
               
                 AAGAAGATGG GCAGTTTTCA TGAGGTTGTC AAAGCCCACC ACCTGGTCAA GAACGGCTGG 
                 1185 
               
               
                   
               
               
                 GATGTGGTTG TGCAGGTGAA TCCCTCATTT GCTCACTCCG GGCCGCTGTG TGTAGCAGCA 
                 1245 
               
               
                   
               
               
                 GTGCCGGAGT ACGAACACAC ACATGAGAAA GCACTCAAGT GGTCTGAGCT TGAGGAACCA 
                 1305 
               
               
                   
               
               
                 GCTTACACAT ACCAACAACT TTCAGTTTTT CCCCACCAGT TGCTAAATTT GAGGACAAAT 
                 1365 
               
               
                   
               
               
                 TCATCAGTGC ATTTGGTGAT GCCCTACATT GGGCCAGGCC AACCAACAAA TCTGACTTTG 
                 1435 
               
               
                   
               
               
                 CACAACCCGT GGACCATTGT TATTTTAATT TTGTCTGAAT TGACAGGACC TGGCCAAACT 
                 1485 
               
               
                   
               
               
                 GTGCCTGTGA CCATGTCGGT GGCTCCCATC GATGCAATGG TTAATGGGCC TCTTCCAAAT 
                 1545 
               
               
                   
               
               
                 CCAGAGGCAC CGATTAGAGT GGTGTCTGTG CCTGAATCAG ATTCTTTTAT GTCTTCAGTA 
                 1605 
               
               
                   
               
               
                 CCTGATAATT CGACTCCACT ATACCCCAAG GTTGTGGTCC CACCGCGCCA AGTTCCTGGC 
                 1665 
               
               
                   
               
               
                 CGGTTTACAA ATTTCATTGA TGTGGCAAAA CAGACATATT CATTTTGTTC CATTTCTGGA 
                 1725 
               
               
                   
               
               
                 AAACCTTATT TTGAGGTTAC CAACACCTCT GGGGACGAGC CACTGTTTCA GATGGATGTG 
                 1785 
               
               
                   
               
               
                 TCGCTCAGTG CGGCAGAGCT ACATGGCACT TACGTAGCTA GTTTGTCATC ATTTTTTGCA 
                 1845 
               
               
                   
               
               
                 CAGTACAGAG GCTCACTTAA TTTCAACTTT ATTTTCACTG GTGCAGCAGC CACTAAGGCA 
                 1905 
               
               
                   
               
               
                 AAGTTTCTGG TTGCTTTTGT GCCTCCCCAC AGTGCAGCGC CCAAAACGCG CGATGAAGCA 
                 1965 
               
               
                   
               
               
                 ATGGCGTGCA TCCATGCCGT GTGGGATGTT GGCTTGAACT CAGCTTTTTC TTTTAATGTA 
                 2025 
               
               
                   
               
               
                 CCTTATCCCT CCCCTGCTGA CTTCATGGCC GTTTATTCTG CGGAACGGAC GGTTGTGAAT 
                 2085 
               
               
                   
               
               
                 GTCTCTGGAT GGCTTCAAGT TTATGCACTA ACAGCTCTAA CTTCAACTGA CATTGCCGTG 
                 2145 
               
               
                   
               
               
                 AACAGTAAAG GCCGTGTGCT GGTTGCTGTT TCCGCCGGCC CAGACTTCTC CCTTCGTCAC 
                 2205 
               
               
                   
               
               
                 CCGGCGGACC TGCCCGACAA GCAGGTTACC AATGTGGGAG AGGATGGTGA ACCCGGTGAG 
                 2265 
               
               
                   
               
               
                 ACAGAGCCTC GTCATGCTTT GTCACCCGTG GACATGCACG TGCACACAGA TGTCAGTTTC 
                 2325 
               
               
                   
               
               
                 TTGCTTGACC GGTTCTTTGA TGTTGAGACA CTTGAGCTTT CAAATTTGAC AGGTTCTCCT 
                 2385 
               
               
                   
               
               
                 GCCACACATG TTCTGGATCC GTTTGGCTCG ACTGCCCAAC TGGCTTGGGC ACGTCTGCTA 
                 2445 
               
               
                   
               
               
                 AACACTTGCA CCTACTTCTT TTCTGATTTG GAATTGTCAA TCCAGTTTAA ATTTACCACC 
                 2505 
               
               
                   
               
               
                 ACTCCGTCCT CTGTTGGAGA GGGCTTTGTG TGGGTGAAGT GGCTCCCTGT TGGAGCACCA 
                 2565 
               
               
                   
               
               
                 ACCAAGACCA CAGATGCTTG GCAGTTAGAA GGAGGTGGAA ATTCAGTTAG AATTCAAAAA 
                 2625 
               
               
                   
               
               
                 TTGGCCGTTG CAGGGATGTG CCCCACTGTT GTGTTCAAGA TTGCAGGCTC CCGTTCACAA 
                 2685 
               
               
                   
               
               
                 GCCTGTGCTT CAGCGTTGCC ATATACATCA ATGTGGCGTG TTGTGCCAGT CTTTTACAAT 
                 2745 
               
               
                   
               
               
                 GGCTGGGGTG CACCTACCAA AGAAAAGGCA ACCTACAATT GGCTTCCTGG TGCACACTTT 
                 2805 
               
               
                   
               
               
                 GGTTCCATCT TGCTGACTTC TGATGCGCAT GATAAAGGAG GGTGCTACTT GCGGTATGCT 
                 2865 
               
               
                   
               
               
                 TTCCGCGCGC CAGCGATGTA TTGCCCTCGA CCCATTCCGC CGGCTTTTAC GCGTCCAGCG 
                 2925 
               
               
                   
               
               
                 GACAAAACCA GACATAAATT TCCCACTAAC ATCAAGAAAC AGTGTACTAA TTACTCTCTC 
                 2985 
               
               
                   
               
               
                 CTCAAATTGG CTGGAGATGT TGAGAGCAAC CCTGGCCCCA CTATTTTTTC CAAAGCATCA 
                 3045 
               
               
                   
               
               
                 GCAGACCTGA ATGCCTTGTC AACGTCGCTA GGTGAATTGA CTGGCATGCT AAAAGATCTT 
                 3105 
               
               
                   
               
               
                 AAAGCCAAGG CAGAAACTTA TTCCCCGTTT TACAAAATGG CCAAAATGCT TTTCAAACTT 
                 3165 
               
               
                   
               
               
                 GCAACACTAG CTGTGGCAGC TATGAGGACA AAGGACCCAG TAGTGGTGGT TATGTTGATT 
                 3225 
               
               
                   
               
               
                 GCTGATTTCG GATTGGAGGT CTTTGACACT GGGTTTTTCT TTTCCTACTT TCAAGAGAAG 
                 3285 
               
               
                   
               
               
                 TTGCAGCCTT ATATGAAAAC TATTCCTGGT AAGATTTCTG ATTTGGTCAC TGATGCGGCT 
                 3345 
               
               
                   
               
               
                 ACGGCTGCCG CCCAAATTCC AAAGGGAGTG TATTCTTTTG TGTCGTCATT TTTCGAAACG 
                 3405 
               
               
                   
               
               
                 CCTGAAGGAG TGGTTGAGAA GCAGGTGTCT CTTCGGACAG TGAATGACAT ATTTGCTTTG 
                 3465 
               
               
                   
               
               
                 CTTAAAAATT CTGATTGGTT CATAAAGACT CTTGTTGCCC TCAAGAAATG GCTGACATCC 
                 3525 
               
               
                   
               
               
                 TGGTTTGCTC AAGAACAACA GGCAGATGAT GCGCTCTATT CAGAATTGGA AAAATATCCC 
                 3565 
               
               
                   
               
               
                 TTGTACAAGT TAAAATTGAA GGAACCTGAT ACTCAAGAGG AAGCGCGCCA GTGGTTTAAA 
                 3645 
               
               
                   
               
               
                 GACATGCAGC AGCGTGCTCT CGCTGTGAAG GACAAAGGTC TCTTTTCCCT CCTGCAAATT 
                 3705 
               
               
                   
               
               
                 CCATTAGTTA ACTTGCCCCA GAGCCGTCCA GAGCCCGTTG TATGCGTCCT TCGGGGCGCA 
                 3765 
               
               
                   
               
               
                 TCAGGGCAAG GCAAATCTTA TTTGGCAAAT CTGATGGCTC AAGCAATTTC GCTTCTCTTG 
                 3825 
               
               
                   
               
               
                 GGTGGCAACC AGGACAGTGT GTGGAGTTGT CCTCCTGACC CCACATATTT TGATGGCTAT 
                 3885 
               
               
                   
               
               
                 AACGGACAGG CTGTGGTGAT TATGGATGCA TTGGGCCAGG ATCCGAATGG TGCTGACTTT 
                 3945 
               
               
                   
               
               
                 AAATATTTTT GCCAGATGGT CTCTACAACA GCTTTTGTAC CACCTATGGC CCATTTGGAT 
                 4005 
               
               
                   
               
               
                 GATAAAGGCA TTCCATTTAC TTCTCCTGTT GTTATTTGTA CTACAAATTT GCATTCATCT 
                 4065 
               
               
                   
               
               
                 TTTACCCCTA TTACTGTTTC TTGTCCTGAA GCTCTTAAGA GGAGGTTTCG GTTTGATGTG 
                 4125 
               
               
                   
               
               
                 ACGGTGTCCG CTAAACCGGG CTTTGTGCGC ACTGTTGGTT CAAACCAGCT TTTGAATCTC 
                 4185 
               
               
                   
               
               
                 CCACTTGCTC TTAAGCCAGC TGGTCTTCCC CCACACCCTA TCTTTGAAAA TGACATGCCC 
                 4245 
               
               
                   
               
               
                 ATTATAAATG GGCAGGCTGT TAAATTGGCT CTTTCTGGTG GAGAAGTGAC AGCTTTTGAG 
                 4305 
               
               
                   
               
               
                 CTTATTGAGA TGATACTGTC AGAAGTTCAA AACAGACAAG ACACACACAA AATGCCCATT 
                 4365 
               
               
                   
               
               
                 TTTAAACAAT CATGGTCTGA TTTGTTCAGA AAGTGTACAA CTGATGAGGA ACAGAAAATG 
                 4425 
               
               
                   
               
               
                 TTGCAGTTTT TAATTGACAA TAAAGATTCA GAAATTCTCA GGGCGTTTGT TTCAGAACGC 
                 4485 
               
               
                   
               
               
                 TCCATTTTAC TACATGAAGA GTATCTTAAA TGGGAGTCAT ATATGACCAG GAGAGCCAAG 
                 4545 
               
               
                   
               
               
                 TTTCACCGCC TGGCTGCTGA TTTTGCTATG TTTCTATCCA TTCTTACTTC ACTGATTGTT 
                 4605 
               
               
                   
               
               
                 ATTTTTTGTT TAGTTTATTC TATGTATCAA CTTTTTAAGA CCCCTGACGA GCAATCAGCT 
                 4665 
               
               
                   
               
               
                 TATGATCCTT CAACTAAGCC AAAACCAAAG ACCCAGGAAG TGAAAACACT GAAGATTAGG 
                 4725 
               
               
                   
               
               
                 ACTGAGACTG GTGTACCAGC AACTGACTTG CAACAATCCA TCATGAAAAA TGTTCAGCCA 
                 4785 
               
               
                   
               
               
                 ATTGAGCTTT ACCTTGACAA TGAATTGGTT ACTGACTGCT CTGCCTTGGG TGTTTATGAC 
                 4845 
               
               
                   
               
               
                 AATTCATATT TGGTGCCCCT TCATTTGTTT GAATTTGATT TTGATACCAT TGTGCTTGGT 
                 4905 
               
               
                   
               
               
                 GGACGTCATT ACAAGAAAGC TGAGTGTGAG AAGGTAGAGT TTGAGCTTGA AGTGAATGGA 
                 4965 
               
               
                   
               
               
                 GACGTGGTGT CATCAGATGC GTGTCTACTT CGAGTGTCAT CGGGGCCTAA AGTTAGAAAT 
                 5035 
               
               
                   
               
               
                 ATTGTTCATC TTTTTACAAA TGAAATTGAA TTGAAGAAAA TGACCCAAGT GACAGGAATC 
                 5085 
               
               
                   
               
               
                 ATGAATTCAC CACACCAGGC ACGCACTGTG TTTTTTGGCA GTTTTTTGAC AGTGAGGAAG 
                 5145 
               
               
                   
               
               
                 TCCATCTTAA CATCGGATGG GACTGTAATG CCCAATGTTT TGTCCTATGC CGCTCAGACC 
                 5205 
               
               
                   
               
               
                 TCGCGTGGGT ATTGTGGCGC TGCAATTGTT GCTGGCTCAC CTGCCCGCAT AATTGGTATC 
                 5265 
               
               
                   
               
               
                 CATTCAGCTG GCACTGGATC TGTTGCATTT TGCTCCCTGG TGTCCAGAGA CCAGCTGGAG 
                 5325 
               
               
                   
               
               
                 CAACTCTGGC CCCAGAAACA GGGCAACGTT AGTCGCCTTG ATGACGATGT GAGGGTGTCT 
                 5385 
               
               
                   
               
               
                 GTTCCGCGCC GCTCCAAATT GGTGAAATCA TTGGCTTACC CCATTTTCAA ACCTGACTAT 
                 5445 
               
               
                   
               
               
                 GGCCCAGCGC CACTCTCTCA ATTTGACAAG CGCCTGTCAG ACGGCGTGAA GCTGGATGAA 
                 5505 
               
               
                   
               
               
                 GTGGTTTTTG CTAAACATAC TGGAGACAAG GAGATTTCCG CACAGGACCA GAAATGGCTC 
                 5565 
               
               
                   
               
               
                 TTGCGTGCGG CGCATGTATA CGCCCAGAAG GTTTTCTCCC GGATTGGATT TGACAACCAG 
                 5625 
               
               
                   
               
               
                 GCTTTGACTG AAAAAGAGGC CATTTGTGGC ATTCCTGGCC TTGACAAGAT GGAGCAGGAC 
                 5685 
               
               
                   
               
               
                 ACCGCTCCGG GGCTGCCCTA TGCTCAGCAA AATAAGAGAA GGAAAGACAT CTGTGATTTT 
                 5745 
               
               
                   
               
               
                 GAAGAGGGCC GGCTGAAGGG CGCCGAACTC CAAAAGGACA GATTTATGGC TGGTGACTAC 
                 5805 
               
               
                   
               
               
                 TCTAATTTGG TCTATCAATC ATTTTTGAAA GATGAGATCC GCCCACTTGA GAAAGTTAGG 
                 5865 
               
               
                   
               
               
                 GCTGGAAAGA CCCGCCTGAT TGACGTGCCG CCGATGCCCC ATGTGGTGGT TGGTAGGCAG 
                 5925 
               
               
                   
               
               
                 CTCTTGGGCC GGTTTGTGGC AAAATTTCAT GAAGCAAATG GATTTGACAT TGGCTCAGCC 
                 5985 
               
               
                   
               
               
                 ATTGGATGTG ACCCAGATGT GGACTGGACT CGGTTTGGCC TCGAGTTGGA GCGTTTCAGG 
                 6045 
               
               
                   
               
               
                 TATGTATATG CCTGTGACTA CTCACGGTTC GATGCCAACC ATGCAGCTGA TGCAATGAGA 
                 6105 
               
               
                   
               
               
                 GTTGTGCTTA ACTACTTTTT CTCTGAGGAC CACGGTTTCG ACCCTGGTGT GCCTGCTTTT 
                 6165 
               
               
                   
               
               
                 ATTGAGTCAC TGGTTGATTC AGTGCATGCC TATGAAGAGA AAAGGTATAA CATCTACGGT 
                 6225 
               
               
                   
               
               
                 GGCTTGCCAT CCGGGTGTTC CTGCACATCA ATTTTGAATA CCATCTTGAA CAATGTTTAC 
                 6285 
               
               
                   
               
               
                 ATTCTTGCAG CTATGATGAA GGCTTATGAG AATTTTGAGC CAGATGACAT TCAGGTCATT 
                 6345 
               
               
                   
               
               
                 TGCTATGGGG ACGACTGCCT CATTGCTTCT GATTTTGAAA TTGATTTCCA ACAACTGGTG 
                 6405 
               
               
                   
               
               
                 CCTGTCTTTT CTAGTTTTGG ACAGGTAATA ACTACAGCTG ACAAGACTGA TTTTTTTAAA 
                 6465 
               
               
                   
               
               
                 CTGACAACCA TTTCGGAGGT GACCTTCCTT AAGCGCGCTT TTGTTCTGAC GGCCTTTTAC 
                 6525 
               
               
                   
               
               
                 AAGCCAGTGA TGGATGTGAA GACCCTTGAA GCAATCTTAA GCTTTGTTCG CCCAGGCACA 
                 6585 
               
               
                   
               
               
                 CAGGCTGAAA AGCTCCTGTC CGTGGCGCAG TTGGCAGGCC ACTGCGAACC GGAGCAGTAT 
                 6645 
               
               
                   
               
               
                 GAGCGCCTGT TTGAGCCCTT TGCTGGGATG TATTTCGTCC CTACTTGGCG ACTTGCGCCT 
                 6705 
               
               
                   
               
               
                 GCAGTGGTTG ATGAAGCTTG GTAGCTAAAT TCTTTTTGAC TTTGTTTTTC TTTGTTTTCT 
                 6765 
               
               
                   
               
               
                 TTTAGGCTTT TAAGGTGTTA AGTTTAAAGG TTAAGAGTTT TTAGAAGTTA AGATAGAGTT 
                 6825 
               
               
                   
               
               
                 TAGTTTTTAG TTTTGAGC-poly(A) 
               
             
          
         
       
     
     as disclosed in FIG.  2  and functional equivalents of said nucleotide sequence including naturally occurring derivatives, variants, degeneracy equivalents and deletion mutants thereof. 
     In another aspect, the invention provides a substantially pure amino acid sequence being: 
     
       
         
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
               
               
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
           
               
                   
               
               
                 a substantially pure amino acid sequence (SEQ ID NO:2) being: 
               
               
                   
               
             
             
               
                   
               
             
          
           
               
                 M 
                 A 
                 A 
                 S 
                 K 
                 V 
                 Y 
                 R 
                 V 
                 C 
                 E 
                 Q 
                 T 
                 L 
                 L 
                 A 
                 G 
                 A 
                 V 
                 R 
                 M 
                 M 
                 D 
                 K 
                 F 
               
               
                 L 
                 Q 
                 K 
                 R 
                 T 
                 V 
                 F 
                 V 
                 P 
                 H 
                 L 
                 D 
                 K 
                 T 
                 I 
                 R 
                 L 
                 T 
                 G 
                 L 
                 H 
                 N 
                 Y 
                 D 
                 N 
               
               
                 T 
                 C 
                 W 
                 L 
                 N 
                 A 
                 L 
                 T 
                 Q 
                 L 
                 T 
                 Q 
                 I 
                 L 
                 G 
                 I 
                 R 
                 L 
                 F 
                 D 
                 E 
                 H 
                 F 
                 G 
                 N 
               
               
                 R 
                 G 
                 L 
                 F 
                 T 
                 R 
                 K 
                 T 
                 I 
                 D 
                 W 
                 V 
                 S 
                 D 
                 Q 
                 T 
                 G 
                 I 
                 K 
                 D 
                 L 
                 K 
                 S 
                 G 
                 A 
               
               
                 P 
                 P 
                 L 
                 V 
                 V 
                 V 
                 Y 
                 K 
                 L 
                 W 
                 Q 
                 H 
                 G 
                 H 
                 L 
                 D 
                 V 
                 G 
                 T 
                 M 
                 E 
                 K 
                 P 
                 R 
                 S 
               
               
                 I 
                 T 
                 L 
                 W 
                 S 
                 G 
                 P 
                 K 
                 V 
                 C 
                 L 
                 S 
                 D 
                 F 
                 W 
                 A 
                 C 
                 V 
                 S 
                 A 
                 K 
                 P 
                 G 
                 H 
                 A 
               
               
                 V 
                 F 
                 Y 
                 L 
                 L 
                 T 
                 S 
                 E 
                 G 
                 W 
                 I 
                 C 
                 V 
                 D 
                 D 
                 K 
                 K 
                 I 
                 Y 
                 P 
                 E 
                 T 
                 P 
                 K 
                 T 
               
               
                 E 
                 D 
                 V 
                 L 
                 V 
                 F 
                 A 
                 P 
                 Y 
                 D 
                 F 
                 E 
                 S 
                 L 
                 G 
                 K 
                 D 
                 P 
                 P 
                 K 
                 L 
                 H 
                 Q 
                 R 
                 Y 
               
             
          
           
               
                   
                 L ↓ VP4 
                   
               
             
          
           
               
                 E 
                 K 
                 A 
                 F 
                 E 
                 L 
                 S 
                 G 
                 G 
                 G 
                 T 
                 S 
                 T 
                 P 
                 T 
                 T 
                 G 
                 N 
                 Q 
                 N 
                 M 
                 S 
                 G 
                 N 
                 S 
               
               
                 G 
                 S 
                 I 
                 V 
                 Q 
                 N 
                 F 
                 Y 
                 M 
                 Q 
                 Q 
                 Y 
                 Q 
                 N 
                 S 
                 I 
                 D 
                 A 
                 D 
                 L 
                 G 
                 D 
                 N 
                 V 
                 I 
               
               
                 S 
                 P 
                 E 
                 G 
                 Q 
                 G 
                 S 
                 N 
                 T 
                 S 
                 S 
                 S 
                 T 
                 S 
                 S 
                 S 
                 Q 
                 S 
                 S 
                 G 
                 L 
                 G 
                 G 
                 W 
                 F 
               
             
          
           
               
                   
                 VP4 ↓ VP2 
                   
               
             
          
           
               
                 S 
                 S 
                 L 
                 L 
                 N 
                 L 
                 G 
                 T 
                 K 
                 L 
                 L 
                 A 
                 D 
                 K 
                 K 
                 T 
                 E 
                 E 
                 T 
                 T 
                 N 
                 I 
                 E 
                 D 
                 R 
               
               
                 I 
                 E 
                 T 
                 T 
                 V 
                 V 
                 G 
                 V 
                 T 
                 I 
                 I 
                 N 
                 S 
                 Q 
                 G 
                 S 
                 V 
                 G 
                 T 
                 T 
                 Y 
                 C 
                 Y 
                 S 
                 K 
               
               
                 P 
                 D 
                 G 
                 R 
                 P 
                 P 
                 S 
                 T 
                 V 
                 S 
                 D 
                 P 
                 V 
                 T 
                 R 
                 L 
                 G 
                 P 
                 T 
                 L 
                 S 
                 R 
                 H 
                 Y 
                 T 
               
               
                 F 
                 K 
                 V 
                 G 
                 E 
                 W 
                 P 
                 H 
                 S 
                 Q 
                 S 
                 H 
                 G 
                 H 
                 A 
                 W 
                 I 
                 C 
                 P 
                 L 
                 P 
                 G 
                 D 
                 K 
                 L 
               
               
                 K 
                 K 
                 M 
                 G 
                 S 
                 F 
                 H 
                 E 
                 V 
                 V 
                 K 
                 A 
                 H 
                 M 
                 L 
                 V 
                 K 
                 N 
                 G 
                 W 
                 D 
                 V 
                 V 
                 V 
                 Q 
               
               
                 V 
                 N 
                 P 
                 S 
                 F 
                 A 
                 H 
                 S 
                 G 
                 P 
                 L 
                 C 
                 V 
                 A 
                 A 
                 V 
                 P 
                 E 
                 Y 
                 E 
                 H 
                 T 
                 H 
                 E 
                 K 
               
               
                 A 
                 L 
                 K 
                 W 
                 S 
                 E 
                 L 
                 E 
                 E 
                 P 
                 A 
                 Y 
                 T 
                 Y 
                 Q 
                 Q 
                 L 
                 S 
                 V 
                 F 
                 P 
                 H 
                 Q 
                 L 
                 L 
               
               
                 N 
                 L 
                 R 
                 T 
                 N 
                 S 
                 S 
                 V 
                 H 
                 L 
                 V 
                 M 
                 P 
                 Y 
                 I 
                 G 
                 P 
                 G 
                 Q 
                 P 
                 T 
                 N 
                 L 
                 T 
                 L 
               
               
                 H 
                 N 
                 P 
                 W 
                 T 
                 I 
                 V 
                 I 
                 L 
                 I 
                 L 
                 S 
                 E 
                 L 
                 T 
                 G 
                 P 
                 G 
                 Q 
                 T 
                 V 
                 P 
                 V 
                 T 
                 M 
               
             
          
           
               
                   
                 VP2 ↓ VP3 
                   
               
             
          
           
               
                 S 
                 V 
                 A 
                 P 
                 I 
                 D 
                 A 
                 M 
                 V 
                 N 
                 G 
                 P 
                 L 
                 P 
                 N 
                 P 
                 E 
                 A 
                 P 
                 I 
                 R 
                 V 
                 V 
                 S 
                 V 
               
               
                 P 
                 E 
                 S 
                 D 
                 S 
                 F 
                 M 
                 S 
                 S 
                 V 
                 P 
                 D 
                 N 
                 S 
                 T 
                 P 
                 L 
                 Y 
                 P 
                 K 
                 V 
                 V 
                 V 
                 P 
                 P 
               
               
                 R 
                 Q 
                 V 
                 P 
                 G 
                 R 
                 F 
                 T 
                 N 
                 F 
                 I 
                 D 
                 V 
                 A 
                 K 
                 Q 
                 T 
                 Y 
                 S 
                 F 
                 C 
                 S 
                 I 
                 S 
                 G 
               
               
                 K 
                 P 
                 Y 
                 F 
                 E 
                 V 
                 T 
                 N 
                 T 
                 S 
                 G 
                 D 
                 E 
                 P 
                 L 
                 F 
                 Q 
                 M 
                 D 
                 V 
                 S 
                 L 
                 S 
                 A 
                 A 
               
               
                 E 
                 L 
                 H 
                 G 
                 T 
                 Y 
                 V 
                 A 
                 S 
                 L 
                 S 
                 S 
                 F 
                 F 
                 A 
                 Q 
                 Y 
                 R 
                 G 
                 S 
                 L 
                 N 
                 F 
                 N 
                 F 
               
               
                 I 
                 F 
                 T 
                 G 
                 A 
                 A 
                 A 
                 T 
                 K 
                 A 
                 K 
                 F 
                 L 
                 V 
                 A 
                 F 
                 V 
                 P 
                 P 
                 H 
                 S 
                 A 
                 A 
                 P 
                 K 
               
               
                 T 
                 R 
                 D 
                 E 
                 A 
                 M 
                 A 
                 C 
                 I 
                 H 
                 A 
                 V 
                 N 
                 D 
                 V 
                 G 
                 L 
                 N 
                 S 
                 A 
                 F 
                 S 
                 F 
                 N 
                 V 
               
               
                 P 
                 Y 
                 P 
                 S 
                 P 
                 A 
                 D 
                 F 
                 M 
                 A 
                 V 
                 Y 
                 S 
                 A 
                 E 
                 R 
                 T 
                 V 
                 V 
                 N 
                 V 
                 S 
                 G 
                 W 
                 L 
               
               
                 Q 
                 V 
                 Y 
                 A 
                 L 
                 T 
                 A 
                 L 
                 T 
                 S 
                 T 
                 D 
                 I 
                 A 
                 V 
                 N 
                 S 
                 K 
                 G 
                 R 
                 V 
                 L 
                 V 
                 A 
                 V 
               
             
          
           
               
                   
                 VP3 ↓ VP2 
                   
               
             
          
           
               
                 S 
                 A 
                 G 
                 P 
                 D 
                 F 
                 S 
                 L 
                 R 
                 H 
                 P 
                 A 
                 D 
                 L 
                 P 
                 D 
                 K 
                 Q 
                 V 
                 T 
                 N 
                 V 
                 G 
                 E 
                 D 
               
               
                 G 
                 E 
                 P 
                 G 
                 E 
                 T 
                 E 
                 P 
                 R 
                 H 
                 A 
                 L 
                 S 
                 P 
                 V 
                 D 
                 M 
                 H 
                 V 
                 H 
                 T 
                 D 
                 V 
                 S 
                 F 
               
               
                 L 
                 L 
                 D 
                 R 
                 F 
                 F 
                 D 
                 V 
                 E 
                 T 
                 L 
                 E 
                 L 
                 S 
                 N 
                 L 
                 T 
                 G 
                 S 
                 P 
                 A 
                 T 
                 H 
                 V 
                 L 
               
               
                 D 
                 P 
                 F 
                 G 
                 S 
                 T 
                 A 
                 Q 
                 L 
                 A 
                 W 
                 A 
                 R 
                 L 
                 L 
                 N 
                 T 
                 C 
                 T 
                 Y 
                 F 
                 F 
                 S 
                 D 
                 L 
               
               
                 E 
                 L 
                 S 
                 I 
                 Q 
                 F 
                 K 
                 F 
                 T 
                 T 
                 T 
                 P 
                 S 
                 S 
                 V 
                 G 
                 E 
                 G 
                 F 
                 V 
                 W 
                 V 
                 K 
                 W 
                 L 
               
               
                 P 
                 V 
                 G 
                 A 
                 P 
                 T 
                 K 
                 T 
                 T 
                 D 
                 A 
                 W 
                 Q 
                 L 
                 E 
                 G 
                 G 
                 G 
                 N 
                 S 
                 V 
                 R 
                 I 
                 Q 
                 K 
               
               
                 L 
                 A 
                 V 
                 A 
                 G 
                 M 
                 C 
                 P 
                 T 
                 V 
                 V 
                 F 
                 K 
                 I 
                 A 
                 G 
                 S 
                 R 
                 S 
                 Q 
                 A 
                 C 
                 A 
                 S 
                 A 
               
               
                 L 
                 P 
                 Y 
                 T 
                 S 
                 M 
                 W 
                 R 
                 V 
                 V 
                 P 
                 V 
                 F 
                 Y 
                 N 
                 G 
                 W 
                 G 
                 A 
                 P 
                 T 
                 K 
                 E 
                 K 
                 A 
               
               
                 T 
                 Y 
                 N 
                 W 
                 L 
                 P 
                 G 
                 A 
                 H 
                 F 
                 G 
                 S 
                 I 
                 L 
                 L 
                 T 
                 S 
                 D 
                 A 
                 H 
                 D 
                 K 
                 G 
                 G 
                 C 
               
               
                 Y 
                 L 
                 R 
                 Y 
                 A 
                 F 
                 R 
                 A 
                 P 
                 A 
                 M 
                 Y 
                 C 
                 P 
                 R 
                 P 
                 I 
                 P 
                 P 
                 A 
                 F 
                 T 
                 R 
                 P 
                 A 
               
             
          
           
               
                   
                 VP1 ↓ 2A 
                   
               
             
          
           
               
                 D 
                 K 
                 T 
                 R 
                 H 
                 K 
                 F 
                 P 
                 T 
                 N 
                 I 
                 N 
                 K 
                 Q 
                 C 
                 T 
                 N 
                 Y 
                 S 
                 L 
                 L 
                 K 
                 L 
                 A 
                 G 
               
             
          
           
               
                   
                 2A ↓ 2B 
                   
               
             
          
           
               
                 D 
                 V 
                 E 
                 S 
                 N 
                 P 
                 G 
                 P 
                 T 
                 I 
                 F 
                 S 
                 K 
                 A 
                 S 
                 A 
                 D 
                 L 
                 N 
                 A 
                 L 
                 S 
                 T 
                 S 
                 L 
               
               
                 G 
                 E 
                 L 
                 T 
                 G 
                 M 
                 L 
                 K 
                 D 
                 L 
                 K 
                 A 
                 K 
                 A 
                 E 
                 T 
                 Y 
                 S 
                 P 
                 F 
                 Y 
                 K 
                 M 
                 A 
                 K 
               
               
                 M 
                 L 
                 F 
                 K 
                 L 
                 A 
                 T 
                 L 
                 A 
                 V 
                 A 
                 A 
                 M 
                 R 
                 T 
                 K 
                 D 
                 P 
                 V 
                 V 
                 V 
                 V 
                 M 
                 L 
                 I 
               
               
                 A 
                 D 
                 F 
                 G 
                 L 
                 E 
                 V 
                 F 
                 D 
                 T 
                 G 
                 F 
                 F 
                 F 
                 S 
                 Y 
                 F 
                 Q 
                 E 
                 K 
                 L 
                 Q 
                 P 
                 Y 
                 M 
               
               
                 K 
                 T 
                 I 
                 P 
                 G 
                 K 
                 I 
                 S 
                 D 
                 L 
                 V 
                 T 
                 D 
                 A 
                 A 
                 T 
                 A 
                 A 
                 A 
                 Q 
                 I 
                 P 
                 K 
                 G 
                 V 
               
             
          
           
               
                   
                 2B ↓ 2C 
                   
               
             
          
           
               
                 Y 
                 S 
                 F 
                 V 
                 S 
                 S 
                 F 
                 F 
                 E 
                 T 
                 P 
                 E 
                 G 
                 V 
                 V 
                 E 
                 K 
                 Q 
                 V 
                 S 
                 L 
                 R 
                 T 
                 V 
                 N 
               
               
                 D 
                 I 
                 F 
                 A 
                 L 
                 L 
                 K 
                 N 
                 S 
                 D 
                 W 
                 F 
                 I 
                 K 
                 T 
                 L 
                 V 
                 A 
                 L 
                 K 
                 K 
                 W 
                 L 
                 T 
                 S 
               
               
                 W 
                 F 
                 A 
                 Q 
                 E 
                 Q 
                 Q 
                 A 
                 D 
                 D 
                 A 
                 L 
                 Y 
                 S 
                 E 
                 L 
                 E 
                 K 
                 Y 
                 P 
                 L 
                 Y 
                 K 
                 L 
                 K 
               
               
                 L 
                 K 
                 E 
                 P 
                 D 
                 T 
                 Q 
                 E 
                 E 
                 A 
                 R 
                 Q 
                 W 
                 F 
                 K 
                 D 
                 M 
                 Q 
                 Q 
                 R 
                 A 
                 L 
                 A 
                 V 
                 K 
               
               
                 D 
                 K 
                 G 
                 L 
                 F 
                 S 
                 L 
                 L 
                 Q 
                 I 
                 P 
                 L 
                 V 
                 N 
                 L 
                 P 
                 Q 
                 S 
                 R 
                 P 
                 E 
                 P 
                 V 
                 V 
                 C 
               
               
                 V 
                 L 
                 R 
                 G 
                 A 
                 S 
                 G 
                 Q 
                 G 
                 K 
                 S 
                 Y 
                 L 
                 A 
                 N 
                 L 
                 M 
                 A 
                 Q 
                 A 
                 I 
                 S 
                 L 
                 L 
                 L 
               
               
                 V 
                 G 
                 K 
                 Q 
                 D 
                 S 
                 V 
                 W 
                 S 
                 C 
                 P 
                 P 
                 D 
                 P 
                 T 
                 Y 
                 F 
                 D 
                 G 
                 Y 
                 N 
                 G 
                 Q 
                 A 
                 V 
               
               
                 V 
                 I 
                 M 
                 D 
                 A 
                 L 
                 G 
                 Q 
                 D 
                 P 
                 N 
                 G 
                 A 
                 D 
                 F 
                 K 
                 Y 
                 F 
                 C 
                 Q 
                 M 
                 V 
                 S 
                 T 
                 T 
               
               
                 A 
                 F 
                 V 
                 P 
                 P 
                 M 
                 A 
                 H 
                 L 
                 D 
                 D 
                 K 
                 G 
                 I 
                 P 
                 F 
                 T 
                 S 
                 P 
                 V 
                 V 
                 I 
                 C 
                 T 
                 T 
               
               
                 N 
                 L 
                 H 
                 S 
                 S 
                 F 
                 T 
                 P 
                 I 
                 T 
                 V 
                 S 
                 C 
                 P 
                 E 
                 A 
                 L 
                 K 
                 R 
                 R 
                 F 
                 R 
                 F 
                 D 
                 V 
               
               
                 T 
                 V 
                 S 
                 A 
                 K 
                 P 
                 G 
                 F 
                 V 
                 R 
                 T 
                 V 
                 G 
                 S 
                 N 
                 Q 
                 L 
                 L 
                 N 
                 L 
                 P 
                 L 
                 A 
                 L 
                 K 
               
               
                 P 
                 A 
                 G 
                 L 
                 P 
                 P 
                 H 
                 P 
                 I 
                 F 
                 E 
                 N 
                 D 
                 M 
                 P 
                 I 
                 I 
                 N 
                 G 
                 Q 
                 A 
                 V 
                 K 
                 L 
                 A 
               
               
                 L 
                 S 
                 G 
                 G 
                 E 
                 V 
                 T 
                 A 
                 F 
                 E 
                 L 
                 I 
                 E 
                 M 
                 I 
                 L 
                 S 
                 E 
                 V 
                 Q 
                 N 
                 R 
                 Q 
                 D 
                 T 
               
             
          
           
               
                   
                 2C ↓ 3A 
                   
               
             
          
           
               
                 H 
                 K 
                 M 
                 P 
                 I 
                 F 
                 K 
                 Q 
                 S 
                 W 
                 S 
                 D 
                 L 
                 F 
                 R 
                 K 
                 C 
                 T 
                 T 
                 D 
                 E 
                 E 
                 Q 
                 K 
                 M 
               
               
                 L 
                 Q 
                 F 
                 L 
                 I 
                 D 
                 N 
                 K 
                 D 
                 S 
                 E 
                 I 
                 L 
                 R 
                 A 
                 F 
                 V 
                 S 
                 E 
                 R 
                 S 
                 I 
                 L 
                 L 
                 H 
               
               
                 E 
                 E 
                 Y 
                 L 
                 K 
                 W 
                 E 
                 S 
                 Y 
                 M 
                 T 
                 R 
                 R 
                 A 
                 K 
                 F 
                 H 
                 R 
                 L 
                 A 
                 A 
                 D 
                 F 
                 A 
                 M 
               
               
                 F 
                 L 
                 S 
                 I 
                 L 
                 T 
                 S 
                 L 
                 I 
                 V 
                 I 
                 F 
                 C 
                 L 
                 V 
                 Y 
                 S 
                 M 
                 Y 
                 Q 
                 L 
                 F 
                 K 
                 T 
                 P 
               
             
          
           
               
                   
                 3A ↓ 3B 
                   
               
             
          
           
               
                 D 
                 E 
                 Q 
                 S 
                 A 
                 Y 
                 D 
                 P 
                 S 
                 T 
                 K 
                 P 
                 K 
                 P 
                 K 
                 T 
                 Q 
                 E 
                 V 
                 K 
                 T 
                 L 
                 K 
                 T 
                 R 
               
             
          
           
               
                   
                 3B ↓ 3C 
                   
               
             
          
           
               
                 T 
                 E 
                 T 
                 G 
                 V 
                 P 
                 A 
                 T 
                 D 
                 L 
                 Q 
                 Q 
                 S 
                 I 
                 M 
                 K 
                 N 
                 V 
                 Q 
                 P 
                 I 
                 E 
                 L 
                 Y 
                 L 
               
               
                 D 
                 N 
                 E 
                 L 
                 V 
                 T 
                 D 
                 C 
                 S 
                 A 
                 L 
                 G 
                 V 
                 Y 
                 D 
                 N 
                 S 
                 Y 
                 L 
                 V 
                 P 
                 L 
                 H 
                 L 
                 F 
               
               
                 E 
                 F 
                 D 
                 F 
                 D 
                 T 
                 I 
                 V 
                 L 
                 G 
                 G 
                 R 
                 H 
                 Y 
                 K 
                 K 
                 A 
                 E 
                 C 
                 E 
                 K 
                 V 
                 E 
                 F 
                 E 
               
               
                 L 
                 E 
                 V 
                 N 
                 G 
                 D 
                 V 
                 V 
                 S 
                 S 
                 D 
                 A 
                 C 
                 L 
                 L 
                 R 
                 V 
                 S 
                 S 
                 G 
                 P 
                 K 
                 V 
                 R 
                 N 
               
               
                 I 
                 V 
                 K 
                 L 
                 F 
                 T 
                 N 
                 E 
                 I 
                 E 
                 L 
                 K 
                 K 
                 M 
                 T 
                 Q 
                 V 
                 T 
                 G 
                 I 
                 M 
                 N 
                 S 
                 P 
                 H 
               
               
                 Q 
                 A 
                 R 
                 T 
                 V 
                 F 
                 F 
                 G 
                 S 
                 F 
                 L 
                 T 
                 V 
                 R 
                 K 
                 S 
                 I 
                 L 
                 T 
                 S 
                 D 
                 G 
                 T 
                 V 
                 M 
               
               
                 P 
                 N 
                 V 
                 L 
                 S 
                 Y 
                 A 
                 A 
                 Q 
                 T 
                 S 
                 R 
                 G 
                 Y 
                 C 
                 G 
                 A 
                 A 
                 I 
                 V 
                 A 
                 G 
                 S 
                 P 
                 A 
               
               
                 R 
                 I 
                 I 
                 G 
                 I 
                 H 
                 S 
                 A 
                 G 
                 T 
                 G 
                 S 
                 V 
                 A 
                 F 
                 C 
                 S 
                 L 
                 V 
                 S 
                 R 
                 D 
                 A 
                 L 
                 S 
               
             
          
           
               
                   
                 3C ↓ 3D 
                   
               
             
          
           
               
                 Q 
                 L 
                 W 
                 P 
                 Q 
                 K 
                 Q 
                 G 
                 N 
                 V 
                 S 
                 R 
                 L 
                 D 
                 D 
                 D 
                 V 
                 R 
                 V 
                 S 
                 V 
                 P 
                 R 
                 R 
                 S 
               
               
                 K 
                 L 
                 V 
                 K 
                 S 
                 L 
                 A 
                 Y 
                 P 
                 I 
                 F 
                 K 
                 P 
                 D 
                 Y 
                 G 
                 P 
                 A 
                 P 
                 L 
                 S 
                 Q 
                 F 
                 D 
                 K 
               
               
                 R 
                 L 
                 S 
                 D 
                 G 
                 V 
                 K 
                 L 
                 D 
                 E 
                 V 
                 V 
                 F 
                 A 
                 K 
                 H 
                 T 
                 G 
                 D 
                 K 
                 E 
                 I 
                 S 
                 A 
                 Q 
               
               
                 D 
                 Q 
                 K 
                 W 
                 L 
                 L 
                 R 
                 A 
                 A 
                 H 
                 V 
                 Y 
                 A 
                 Q 
                 K 
                 V 
                 F 
                 S 
                 R 
                 I 
                 G 
                 F 
                 D 
                 N 
                 Q 
               
               
                 A 
                 L 
                 T 
                 E 
                 K 
                 E 
                 A 
                 I 
                 C 
                 G 
                 I 
                 P 
                 G 
                 L 
                 D 
                 K 
                 M 
                 E 
                 Q 
                 D 
                 T 
                 A 
                 P 
                 G 
                 L 
               
               
                 P 
                 Y 
                 A 
                 Q 
                 Q 
                 N 
                 K 
                 R 
                 R 
                 K 
                 D 
                 I 
                 C 
                 D 
                 F 
                 E 
                 E 
                 G 
                 R 
                 L 
                 K 
                 G 
                 A 
                 E 
                 L 
               
               
                 Q 
                 K 
                 D 
                 R 
                 F 
                 M 
                 A 
                 G 
                 D 
                 Y 
                 S 
                 N 
                 L 
                 V 
                 Y 
                 Q 
                 S 
                 F 
                 L 
                 K 
                 D 
                 E 
                 I 
                 R 
                 P 
               
               
                 L 
                 E 
                 K 
                 V 
                 R 
                 A 
                 G 
                 K 
                 T 
                 R 
                 L 
                 I 
                 D 
                 V 
                 P 
                 P 
                 M 
                 P 
                 H 
                 V 
                 V 
                 V 
                 G 
                 R 
                 Q 
               
               
                 L 
                 L 
                 G 
                 R 
                 F 
                 V 
                 A 
                 K 
                 F 
                 H 
                 E 
                 A 
                 N 
                 G 
                 F 
                 D 
                 I 
                 G 
                 S 
                 A 
                 I 
                 G 
                 C 
                 D 
                 P 
               
               
                 D 
                 V 
                 D 
                 W 
                 T 
                 R 
                 F 
                 G 
                 L 
                 E 
                 L 
                 E 
                 R 
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                 Y 
                 V 
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                 S 
                 R 
                 F 
               
               
                 D 
                 A 
                 N 
                 H 
                 A 
                 A 
                 D 
                 A 
                 M 
                 R 
                 V 
                 V 
                 L 
                 N 
                 Y 
                 F 
                 F 
                 S 
                 E 
                 D 
                 H 
                 G 
                 F 
                 D 
                 P 
               
               
                 G 
                 V 
                 P 
                 A 
                 F 
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                 E 
                 S 
                 L 
                 V 
                 D 
                 S 
                 V 
                 H 
                 A 
                 Y 
                 E 
                 E 
                 K 
                 R 
                 Y 
                 N 
                 I 
                 Y 
                 G 
               
               
                 G 
                 L 
                 P 
                 S 
                 G 
                 C 
                 S 
                 C 
                 T 
                 S 
                 I 
                 L 
                 N 
                 T 
                 I 
                 L 
                 N 
                 N 
                 V 
                 Y 
                 I 
                 L 
                 A 
                 A 
                 M 
               
               
                 M 
                 K 
                 A 
                 Y 
                 E 
                 N 
                 F 
                 E 
                 P 
                 D 
                 D 
                 I 
                 Q 
                 V 
                 I 
                 C 
                 Y 
                 G 
                 D 
                 D 
                 C 
                 L 
                 I 
                 A 
                 S 
               
               
                 D 
                 F 
                 E 
                 I 
                 D 
                 F 
                 Q 
                 Q 
                 L 
                 V 
                 P 
                 V 
                 F 
                 S 
                 S 
                 F 
                 G 
                 Q 
                 V 
                 I 
                 T 
                 T 
                 A 
                 D 
                 K 
               
               
                 T 
                 D 
                 F 
                 F 
                 K 
                 L 
                 T 
                 T 
                 L 
                 S 
                 E 
                 V 
                 T 
                 F 
                 L 
                 K 
                 R 
                 A 
                 F 
                 V 
                 L 
                 T 
                 A 
                 F 
                 Y 
               
               
                 K 
                 P 
                 V 
                 M 
                 D 
                 V 
                 K 
                 T 
                 L 
                 E 
                 A 
                 I 
                 L 
                 S 
                 F 
                 V 
                 R 
                 P 
                 G 
                 T 
                 Q 
                 A 
                 E 
                 K 
                 L 
               
               
                 L 
                 S 
                 V 
                 A 
                 Q 
                 L 
                 A 
                 G 
                 H 
                 C 
                 E 
                 P 
                 E 
                 Q 
                 Y 
                 E 
                 R 
                 L 
                 F 
                 E 
                 P 
                 F 
                 A 
                 G 
                 M 
               
             
          
           
               
                   
                 3D 
                   
               
             
          
           
               
                 Y 
                 F 
                 V 
                 P 
                 T 
                 W 
                 R 
                 L 
                 A 
                 P 
                 A 
                 V 
                 V 
                 D 
                 E 
                 A 
                 W 
                 M 
                 L 
                 N 
                 S 
                 F 
               
               
                   
               
             
          
         
       
     
     as disclosed in FIG.  2 . 
     In another aspect, the invention provides proteins derived from ERhV1 which exhibit virus like particles characteristics incorporating VP1 and having the following amino acid sequence: 
     
       
         
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
           
               
                   
               
               
                 a protein or virus like particle incorporating VP1, derived from ERhV1 and 
               
               
                 having the following amino acid sequence (SEQ ID NO:3): 
               
               
                   
               
             
             
               
                   
               
             
          
           
               
                 V 
                 T 
                 N 
                 V 
                 G 
                 E 
                 D 
                 G 
                 E 
                 P 
                 G 
                 E 
                 T 
                 E 
                 P 
                 R 
                 H 
                 A 
                 L 
                 S 
                 P 
                 V 
                 D 
                 M 
                 H 
               
               
                 V 
                 H 
                 T 
                 D 
                 V 
                 S 
                 F 
                 L 
                 L 
                 D 
                 R 
                 F 
                 F 
                 D 
                 V 
                 E 
                 T 
                 L 
                 E 
                 L 
                 S 
                 N 
                 L 
                 T 
                 G 
               
               
                 S 
                 P 
                 A 
                 T 
                 H 
                 V 
                 L 
                 D 
                 P 
                 F 
                 G 
                 S 
                 T 
                 A 
                 Q 
                 L 
                 A 
                 W 
                 A 
                 R 
                 L 
                 L 
                 N 
                 T 
                 C 
               
               
                 T 
                 Y 
                 F 
                 F 
                 S 
                 D 
                 L 
                 E 
                 L 
                 S 
                 I 
                 Q 
                 F 
                 K 
                 F 
                 T 
                 T 
                 T 
                 P 
                 S 
                 S 
                 V 
                 G 
                 E 
                 G 
               
               
                 F 
                 V 
                 W 
                 V 
                 K 
                 W 
                 L 
                 P 
                 V 
                 G 
                 A 
                 P 
                 T 
                 K 
                 T 
                 T 
                 D 
                 A 
                 W 
                 Q 
                 L 
                 E 
                 G 
                 G 
                 G 
               
               
                 N 
                 S 
                 V 
                 R 
                 I 
                 Q 
                 K 
                 L 
                 A 
                 V 
                 A 
                 G 
                 M 
                 C 
                 P 
                 T 
                 V 
                 V 
                 F 
                 K 
                 I 
                 A 
                 G 
                 S 
                 R 
               
               
                 S 
                 Q 
                 A 
                 C 
                 A 
                 S 
                 A 
                 L 
                 P 
                 Y 
                 T 
                 S 
                 M 
                 W 
                 R 
                 V 
                 V 
                 P 
                 V 
                 F 
                 Y 
                 N 
                 G 
                 W 
                 G 
               
               
                 A 
                 P 
                 T 
                 K 
                 E 
                 K 
                 A 
                 T 
                 Y 
                 N 
                 W 
                 L 
                 P 
                 G 
                 A 
                 H 
                 F 
                 G 
                 S 
                 I 
                 L 
                 L 
                 T 
                 S 
                 D 
               
               
                 A 
                 K 
                 D 
                 K 
                 G 
                 G 
                 C 
                 Y 
                 L 
                 R 
                 Y 
                 A 
                 F 
                 R 
                 A 
                 P 
                 A 
                 M 
                 Y 
                 C 
                 P 
                 R 
                 P 
                 I 
                 P 
               
               
                 P 
                 A 
                 F 
                 T 
                 R 
                 P 
                 A 
                 D 
                 K 
                 T 
                 R 
                 H 
                 K 
                 F 
                 P 
                 T 
                 N 
                 I 
                 N 
                 K 
                 Q 
                 C 
                 T 
               
               
                   
               
             
          
         
       
     
     In another aspect, the invention provides proteins derived from ERhV1 which exhibit virus like particle characteristics incorporating VP2 and having the following amino acid sequence: 
     
       
         
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
           
               
                   
               
               
                 a protein or virus like particle incorporating VP2 deived from ERhV1 and 
               
               
                 having the following amino acid sequence (SEQ ID NO:4): 
               
               
                   
               
             
             
               
                   
               
             
          
           
               
                 D 
                 K 
                 K 
                 T 
                 E 
                 E 
                 T 
                 T 
                 N 
                 I 
                 E 
                 D 
                 R 
                 I 
                 E 
                 T 
                 T 
                 V 
                 V 
                 G 
                 V 
                 T 
                 I 
                 I 
                 N 
               
               
                 S 
                 Q 
                 G 
                 S 
                 V 
                 G 
                 T 
                 T 
                 Y 
                 C 
                 Y 
                 S 
                 K 
                 P 
                 D 
                 G 
                 R 
                 P 
                 P 
                 S 
                 T 
                 V 
                 S 
                 D 
                 P 
               
               
                 V 
                 T 
                 R 
                 L 
                 G 
                 P 
                 T 
                 L 
                 S 
                 R 
                 H 
                 Y 
                 T 
                 F 
                 K 
                 V 
                 G 
                 E 
                 W 
                 P 
                 H 
                 S 
                 Q 
                 S 
                 H 
               
               
                 G 
                 H 
                 A 
                 W 
                 I 
                 C 
                 P 
                 L 
                 P 
                 G 
                 D 
                 K 
                 L 
                 K 
                 K 
                 M 
                 G 
                 S 
                 F 
                 H 
                 E 
                 V 
                 V 
                 K 
                 A 
               
               
                 H 
                 H 
                 L 
                 V 
                 K 
                 N 
                 G 
                 W 
                 D 
                 V 
                 V 
                 V 
                 Q 
                 V 
                 N 
                 P 
                 S 
                 F 
                 A 
                 H 
                 S 
                 G 
                 P 
                 L 
                 C 
               
               
                 V 
                 A 
                 A 
                 V 
                 P 
                 E 
                 Y 
                 E 
                 H 
                 T 
                 H 
                 E 
                 K 
                 A 
                 L 
                 K 
                 W 
                 S 
                 E 
                 L 
                 E 
                 E 
                 P 
                 A 
                 Y 
               
               
                 T 
                 Y 
                 Q 
                 Q 
                 L 
                 S 
                 V 
                 F 
                 P 
                 H 
                 Q 
                 L 
                 L 
                 N 
                 L 
                 R 
                 T 
                 N 
                 S 
                 S 
                 V 
                 H 
                 L 
                 V 
                 M 
               
               
                 P 
                 Y 
                 I 
                 G 
                 P 
                 G 
                 Q 
                 P 
                 T 
                 N 
                 L 
                 T 
                 L 
                 H 
                 N 
                 P 
                 W 
                 T 
                 I 
                 V 
                 I 
                 L 
                 I 
                 L 
                 S 
               
               
                 E 
                 L 
                 T 
                 G 
                 P 
                 G 
                 Q 
                 T 
                 V 
                 P 
                 V 
                 T 
                 M 
                 S 
                 V 
                 A 
                 P 
                 I 
                 D 
                 A 
                 M 
                 V 
                 N 
                 G 
                 P 
               
               
                 L 
                 P 
                 N 
                 P 
                 E 
               
               
                   
               
             
          
         
       
     
     In another aspect, the invention provides proteins derived from ERhV1 which exhibit virus like particle characteristics incorporating VP3 and having the following amino acid sequence: 
     
       
         
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
           
               
                   
               
               
                 a protein or virus like particle incorporating VP3; derived from ERhV1 and 
               
               
                 having the following amino acid sequence (SEQ ID NO:5): 
               
               
                   
               
             
             
               
                   
               
             
          
           
               
                 A 
                 P 
                 I 
                 R 
                 V 
                 V 
                 S 
                 V 
                 P 
                 E 
                 S 
                 D 
                 S 
                 F 
                 M 
                 S 
                 S 
                 V 
                 P 
                 D 
                 N 
                 S 
                 T 
                 P 
                 L 
               
               
                 Y 
                 P 
                 K 
                 V 
                 V 
                 V 
                 P 
                 P 
                 R 
                 Q 
                 V 
                 P 
                 G 
                 R 
                 F 
                 T 
                 N 
                 F 
                 I 
                 D 
                 V 
                 A 
                 K 
                 Q 
                 T 
               
               
                 Y 
                 S 
                 F 
                 C 
                 S 
                 I 
                 S 
                 G 
                 K 
                 P 
                 Y 
                 F 
                 E 
                 V 
                 T 
                 N 
                 T 
                 S 
                 G 
                 D 
                 E 
                 P 
                 L 
                 F 
                 Q 
               
               
                 M 
                 D 
                 V 
                 S 
                 L 
                 S 
                 A 
                 A 
                 E 
                 L 
                 H 
                 G 
                 T 
                 Y 
                 V 
                 A 
                 S 
                 L 
                 S 
                 S 
                 F 
                 F 
                 A 
                 Q 
                 Y 
               
               
                 R 
                 G 
                 S 
                 L 
                 N 
                 F 
                 N 
                 F 
                 I 
                 F 
                 T 
                 G 
                 A 
                 A 
                 A 
                 T 
                 K 
                 A 
                 K 
                 F 
                 L 
                 V 
                 A 
                 F 
                 V 
               
               
                 P 
                 P 
                 K 
                 S 
                 A 
                 A 
                 P 
                 K 
                 T 
                 R 
                 D 
                 E 
                 A 
                 M 
                 A 
                 C 
                 I 
                 H 
                 A 
                 V 
                 W 
                 D 
                 V 
                 G 
                 L 
               
               
                 N 
                 S 
                 A 
                 F 
                 S 
                 F 
                 N 
                 V 
                 P 
                 Y 
                 P 
                 S 
                 P 
                 A 
                 D 
                 F 
                 M 
                 A 
                 V 
                 Y 
                 S 
                 A 
                 E 
                 R 
                 T 
               
               
                 V 
                 V 
                 N 
                 V 
                 S 
                 G 
                 W 
                 L 
                 Q 
                 V 
                 Y 
                 A 
                 L 
                 T 
                 A 
                 L 
                 T 
                 S 
                 T 
                 D 
                 I 
                 A 
                 V 
                 N 
                 S 
               
               
                 K 
                 G 
                 R 
                 V 
                 L 
                 V 
                 A 
                 V 
                 S 
                 A 
                 G 
                 P 
                 D 
                 F 
                 S 
                 L 
                 R 
                 H 
                 P 
                 A 
                 D 
                 L 
                 P 
                 D 
                 K 
               
               
                 O 
               
               
                   
               
             
          
         
       
     
     In another aspect, the invention provides proteins derived from ERhV1 which exhibit virus like particle characteristics incorporating VP4 and having the following amino acid sequence: 
     
       
         
               
             
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
               
             
           
               
                   
               
               
                 a protein or virus like particle incorporating VP4, derived from ErhV1 and 
               
               
                 having the following amino acid sequence (SEQ ID NO:6): 
               
               
                   
               
             
             
               
                   
               
             
          
           
               
                 G 
                 G 
                 G 
                 T 
                 S 
                 T 
                 P 
                 T 
                 T 
                 G 
                 N 
                 Q 
                 N 
                 M 
                 S 
                 G 
                 N 
                 S 
                 G 
                 S 
                 I 
                 V 
                 Q 
                 N 
                 F 
               
               
                 Y 
                 M 
                 Q 
                 Q 
                 Y 
                 Q 
                 N 
                 S 
                 I 
                 D 
                 A 
                 D 
                 L 
                 G 
                 D 
                 N 
                 V 
                 I 
                 S 
                 P 
                 E 
                 G 
                 Q 
                 G 
                 S 
               
               
                 N 
                 T 
                 S 
                 S 
                 S 
                 T 
                 S 
                 S 
                 S 
                 Q 
                 S 
                 S 
                 G 
                 L 
                 G 
                 G 
                 W 
                 F 
                 S 
                 S 
                 L 
                 L 
                 N 
                 L 
                 G 
               
               
                 T 
                 K 
                 L 
                 L 
                 A 
               
               
                   
               
             
          
         
       
     
     The invention also provides a virus like particle comprising any one or a combination of VP1, VP2, VP3 and VP4. 
     
       
         
               
               
             
           
               
                 In another aspect, the invention provides a substantially 
                   
               
               
                 pure nucleotide sequence for VPI (SEQ ID NO:7) being: 
               
               
                   
               
               
                 GTTACCAATG TGGGAGAGGA TGGTGAACCC GGTGAGACAG AGCCTCGTCA TGCTTTGTCA 
               
               
                   
               
               
                 CCCGTGGACA TGCACGTGCA CACAGATGTC AGTTTCTTGC TTGACCGGTT CTTTGATGTT 
               
               
                   
               
               
                 GAGACACTTG AGCTTTCAAA TTTGACAGGT TCTCCTGCCA CACATGTTCT GGATCCGTTT 
               
               
                   
               
               
                 GGCTCGACTG CCCAACTGGC TTGGGCACGT CTGCTAAACA CTTGCACCTA CTTCTTTTCT 
               
               
                   
               
               
                 GATTTGGAAT TGTCAATCCA GTTTAAATTT ACCACCACTC CGTCCTCCGT TGGAGAGGGC 
               
               
                   
               
               
                 TTTGTGTGGG TGAAGTGGCT CCCTGTTGGA GCACCAACCA AGACCACAGA TGCTTGGCAG 
               
               
                   
               
               
                 TTAGAAGGAG GTGGAAATTC AGTTAGAATT CAAAAATTGG CCGTTGCAGG GATCTGCCCC 
               
               
                   
               
               
                 ACTGTTGTGT TCAAGATTGC AGGCTCCCGT TCACAAGCCT GTGCTTCAGC GTTGCCATAT 
               
               
                   
               
               
                 ACATCAATGT GGCGTGTTGT GCCAGTCTTT TACAATGGCT GGGGTGCACC TACCAAAGAA 
               
               
                   
               
               
                 AAGGCAACCT ACAATTGGCT TCCTGGTGCA CACTTTGGTT CCATCTTGCT GACTTCTGAT 
               
               
                   
               
               
                 GCGCATGATA AAGGAGGGTG CTACTTGCGG TATGCTTTCC GCGCGCCAGC GATGTATTGC 
               
               
                   
               
               
                 CCTCGACCCA TTCCGCCGGC TTTTACGCGT CCAGCGGACA AAACCAGACA TAAATTTCCC 
               
               
                   
               
               
                 ACTAACATCA ACAAACAGTG TACT 
               
             
          
         
       
     
     and functional equivalents of said nucleotide sequence including naturally occurring derivatives, variants and degeneracy equivalents. 
     
       
         
               
               
             
           
               
                 In another aspect, the invention provides a substantially 
                   
               
               
                 pure nucleotide sequence for VP2 (SEQ ID NO:8) being: 
               
               
                   
               
               
                 GACAAGAAGA CAGAAGAGAC TACAAACATT GAAGACACAA TTGAAACAAC AGTGGTTGGA 
               
               
                   
               
               
                 GTCACTATTA TTAATTCACA AGGATCTGTT GGAACAACCT ACTGTTACTC CAAACCGGAT 
               
               
                   
               
               
                 GGTAGACCAC CATCCACAGT GTCAGACCCA GTTACCAGAC TTGGACCCAC GCTTTCCAGG 
               
               
                   
               
               
                 CACTACACAT TTAAGGTAGG TGAGTGGCCC CATTCTCAAT CACATGGTCA CGCATGGATC 
               
               
                   
               
               
                 TGTCCGTTGC CAGGTGACAA ACTCAAGAAG ATGGGCAGTT TTCATGAGGT TGTCAAAGCC 
               
               
                   
               
               
                 CACCACCTGG TCAAGAACGG CTGGGATGTG GTTGTGCAGG TGAATCCCTC ATTTGCTCAC 
               
               
                   
               
               
                 TCCGGGCCGC TGTGTGTAGC AGCAGTGCCG GAGTACGAAC ACACACATGA GAAAGCACTC 
               
               
                   
               
               
                 AAGTGGTCTG AGCTTGAGGA ACCAGCTTAC ACATACCAAC AACTTTCAGT TTTTCCCCAC 
               
               
                   
               
               
                 CAGTTGCTAA ATTTGAGGAC AAATTCATCA GTGCATTTGG TGATGCCCTA CATTGGGCCA 
               
               
                   
               
               
                 GGCCAACCAA CAAATCTGAC TTTGCACAAC CCGTGGACCA TTGTTATTTT AATTTTGTCT 
               
               
                   
               
               
                 GAATTGACAG GACCTGGCCA AACTGTGCCT GTGACCATGT CGGTGGCTCC CATCGATGCA 
               
               
                   
               
               
                 ATGGTTAATG GGCCTCTTCC AAATCCAGAG 
               
             
          
         
       
     
     and functional equivalents of said nucleotide sequence including naturally occurring derivatives, variants and degeneracy equivalents. 
     
       
         
               
               
             
           
               
                 In another aspect, the invention provides a substantially 
                   
               
               
                 pure nucleotide sequence for VP3 (SEQ ID NO:9) being: 
               
               
                   
               
               
                 GCACCGATTA GAGTGGTGTC TGTGCCTGAA TCAGATTCTT TTATGTCTTC AGTACCTGAT 
               
               
                   
               
               
                 AATTCGACTC CACTATACCC CAAGGTTGTG GTCCCACCGC GCCAAGTTCC TGGCCGGTTT 
               
               
                   
               
               
                 ACAAATTTCA TTGATGTGGC AAAACAGACA TATTCATTTT GTTCCATTTC TGGAAAACCT 
               
               
                   
               
               
                 TATTTTGAGG TTACCAACAC CTCTGGGGAC GAGCCACTGT TTCAGATGGA TGTGTCGCTC 
               
               
                   
               
               
                 AGTGCGGCAG AGCTACATGG CACTTACGTA GCTAGTTTGT CATCATTTTT TGCACAGTAC 
               
               
                   
               
               
                 AGAGGCTCAC TTAATTTCAA CTTTATTTTC ACTGGTGCAG CAGCCACTAA GGCAAAGTTT 
               
               
                   
               
               
                 CTGCTTCCTT TTGTGCCTCC CCACAGTGCA GCGCCCAAAA CGCGCGATGA AGCAATGGCG 
               
               
                   
               
               
                 TGCATCCATG CCGTGTGGGA TGTTGGCTTG AACTCAGCTT TTTCTTTTAA TGTACCTTAT 
               
               
                   
               
               
                 CCCTCCCCTG CTGACTTCAT GGCCGTTTAT TCTGCGGAAC GGACGGTTGT GAATGTCTCT 
               
               
                   
               
               
                 GGATGGCTTC AAGTTTATGC ACTAACAGCT CTAACTTCAA CTGACATTGC CGTGAACAGT 
               
               
                   
               
               
                 AAAGGCCGTG TGCTGGTTGC TGTTTCCGCC GGCCCAGACT TCTCCCTTCG TCACCCGGCG 
               
               
                   
               
               
                 GACCTGCCCG ACAACCAG 
               
             
          
         
       
     
     and functional equivalents of said nucleotide sequence including naturally occurring derivatives, variants and degeneracy equivalents. 
     
       
         
               
               
             
           
               
                 In another aspect, the invention provides a substantially 
                   
               
               
                 pure nucleotide sequence for VP4 (SEQ ID NO:10) being: 
               
               
                   
               
               
                 GGCGGAGGTA CATCCACTCC AACAACTGGC AACCAAAACA TGTCCGGAAA CAGTGGTTCA 
               
               
                   
               
               
                 ATTGTTCAAA ATTTTTACAT GCAACAGTAC CAGAATTCAA TTGACGCAGA CCTGGGAGAC 
               
               
                   
               
               
                 AATGTGATTA GCCCTGAAGG CCAGGGCAGC AACACTAGTA GTTCAACCTC ATCAAGCCAA 
               
               
                   
               
               
                 TCCTCTGGCT TGGGCGGGTG GTTCTCTAGT TTGCTGAACC TTGGAACAAA ACTACTGGCT 
               
             
          
         
       
     
     and functional equivalents of said nucleotide sequence including naturally occurring derivatives, variants and degeneracy equivalents. 
     In another aspect, the invention provides oligonucleotide primers derived from the nucleotide sequence of FIG. 2 being highly specific for ERhV1 or cross reactive with other ERhV types. 
     The oligonucleotide primers may have any one of the following nucleotide sequences: 
     
       
         
               
               
               
               
             
           
               
                 VP1F 
                 (SEQ ID NO: 11) 
                 5′ GTTGTGTTCAAGATTGCAGGC 
                 3′ 
               
               
                   
               
               
                 VP1R1 
                 (SEQ ID NO: 12) 
                 5′ TTGCTCTCAACATCTCCAGC 
                 3′ 
               
               
                   
               
               
                 VP1R2 
                 (SEQ ID NO: 13) 
                 5′ TAGCACCCTCCTTTATCATGCG 
                 3′ 
               
             
          
         
       
     
     In another aspect, the invention provides an oligonucleotide probe derived from the sequence of FIG.  2 . 
     In another aspect, the invention provides diagnostic reagents, methods and kits characterised by the aforesaid oligonucleotide primers and probes. 
     In another aspect, the invention provides antigens comprising any one or a combination of the non-capsid proteins, being other than the individual VP1 to VP4 proteins, that are cleaved products of the polypeptide of FIG.  2 . 
     In another aspect, the invention provides vaccines and vectors incorporating any one or a combination of virion proteins VP1 to VP4. 
     In another aspect, the invention provides diagnostic tests for the detection of antibodies to ERhV1 in blood of horses or other animals characterised by the use of the aforesaid antigens. Such diagnostic tests may be ELISA based. 
     In a particularly preferred embodiment, the invention provides a test to distinguish horses infected with ERhV1 in which said virus had replicated from horses which have been vaccinated with the vaccine incorporating any one or a combination of virion proteins VP1 to VP4; comprising the steps of applying an antigen being any one of a combination of non-capsid proteins being other than VP1 to VP4, that are cleavage products of the polypeptide of FIG. 2 to a horse and testing for an immunoreaction thereto, wherein a positive immunoreaction would indicate that said horse had been infected with ERhV1 and a negative immunoreaction would indicate that said horse has not been infected with ERhV1. 
     In another aspect, the invention provides recombinant plasmids incorporating nucleotide sequences and subsequences derived from the nucleotide sequences of FIG.  2 . The recombinant plasmid may comprise the P1-2A-3C region of the ERhV1 genome. 
     In another aspect, the invention provides a host system characterised by incorporating the nucleoside sequence of FIG. 2 or part thereof. The host may be  E.coli,  vaccinia virus, baculovirus or yeast. 
     In another aspect, the invention provides a process for producing a protein product derived from ERhV1 comprising the shape of selecting out a gene of interest from the ERhV1 nucleotide sequence of FIG.  2  and expressing said protein product in a suitable host system. 
    
    
     SUMMARY OF THE DRAWINGS 
     The invention will now be described in detail with reference to FIGS. 1 to  6 : 
     FIGS.  1 (A) and (B) (A) Schematic representation of the ERhV1 genome and (B) comparison of the genomic structures of picornaviruses showing the predicted proteolytic cleavage pattern of the polyprotein,. The length of individual regions are drawn approximately to scale. The dashed line represents the unsequenced region of the ERhV1 5′-NTR. 
     FIGS.  2 A 1 - 2 A 7  Nucleotide and predicted amino acid sequence of the ERhV1 polyprotein (SEQ ID NOS 1 &amp; 2). The nucleotide sequences of the 3′-NTR and part of the 5′-NTR are also shown. Numbering is from the first ATG codon that occurs in a context optimal for translational initiation (Kozak, 1989). A polypyrimidine tract upstream of the putative insulating ATG and the two pairs of in-frame ATG codons are underlined. The predicted proteolytic cleavage sites are indicated by arrows. 
     FIG. 2B Nucleotide sequence of the ERhV1 5′-nontranslated region (SEQ ID NO:22). The polyC tract (dotted underline), polypyrimidine tract (underline) and potential initiation codons (double underline) are indicated. Predicted coding sequence is shown in bold type (SEQ ID NO:23). Numbering is from the ATG considered most likely to be used for translation initiation. 
     FIGS. 3A-3F Alignment of the predicted amino acid sequences of ERhV1.393/76 (SEQ ID NO:25) and FMDV.O1K (SEQ ID NO:24) polyprotein. Proteolytic cleavage sites, which are predicted in the case of ERhV1, are indicated by the arrows. Identical residues (*), highly conserved residues (:), and less conserved residues (.), are indicated. 
     FIGS. 4A-4C Unrooted phylogenic trees inferred using the picornavirus nucleotide sequences of (A) the complete polyprotein gene, (B) the polymerase gene and (C) VP1 gene of viruses representing the five recognised genera of the family Picornaviridae. The viruses used were: FMDV.A10, FMDV.O1K, FMDV.A12, FMDV.C3, FMDV.SAT3, EMCV, TMEV, Mengovirus, poliovirus 1 Mahoney (Polio 1), poliovirus 2.SAbin (Polio 2), poliovirus 3.Leon (Polio 2), coxsackievirus A9 (CV.A9), CV.B3, echovirus 22 (Echo 22), swine vesicular disease virus (SVDV), bovine enterovirus (BEV) heptatitis A virus (HAV) human rhinovirus 1B (HRV1B), HRV89 and HRV14. 
     Note: The branch lengths represent proportionate change only within each tree; they do not allow direct comparisons to be made between the three trees. 
     FIGS. 5A and 5B Diagram outlining the strategy for nested, reverse transcription-polymerase chain reaction (RT-PCR) for the detection of ERhV genome. The genome structure of ERhV1 is shown schematically (top), and the first round PCR product (352 bp), corresponding to VP1 and 2A regions, and the second round PCR product (210 bp), corresponding to part of VP1, are represented as black lines. 
     (B) the sequence of specific oligonucleotide primers used for RT-PCR are shown (SEQ ID NOS 11, 12 &amp; 13). VP1R1 was used for the RT reaction. 
     FIG. 6 Construction of ERhV1 expression plasmid for  E. coli  and baculovirus transfer vector for insect cells. The ERhV1 genome is shown (top) and oligonucleotide primers used to amplify P1.2A and 3C regions are depicted as arrows. The P1.2A fragment and subsequently the P1.2A.3C fragment, obtained through the ligation of P1.2A and 3C, were cloned separately into the multiple cloning sites of the pET15b and pBacbluII plasmid vectors to construct pET.P1.2A and pET.P1.2A.3C respectively for expression in  E. coli  an pBac.P1.2A and pBac.P1.2A.3C respectively for expression in insect cells. 
    
    
     DETAILED DESCRIPTION OF INVENTION 
     The sequence of specific oligonucleotide primers used for the construction of expression plasmids are: 
     
       
         
               
               
               
               
               
             
           
               
                 VP4F 
                 (SEQ ID NO:14) 
                 5′  GCTGGATCCATGAGTGGCGGAGGTACATCCACT 
                 3′ 
                   
               
               
                   
               
               
                 R2A 
                 (SEQ ID NO: 15) 
                 5′  GCTCTGCAGCAGGTCTGCTGATGCTTTGGA 
                 3′ 
               
               
                   
               
               
                 3CF 
                 (SEQ ID NO: 16) 
                 5′  GCTCTGCAGATGATTAGGACTGAGACTGGTGT 
                 3′ 
               
               
                   
               
               
                 3CR 
                 (SEQ ID NO: 17) 
                 5′  GCTGGATCCTTAGCCATAGTCAGGTTTGAA 
                 3′ 
               
             
          
         
       
     
     Virus growth and purification 
     ERhV1 strain 393/76 was isolated from a nasal swab taken from a thoroughbred horse in South Australia while it was being held in quarantine following importation from the United Kingdom. The mare had an acute, systemic febrile illness. The virus was passaged 14 times in quine fetal kidney (EFK) monolayer cell cultures and then once in Vero cells. ERhV1 virions were purified by a modification of the procedure described by Abraham and Colonno. Cells were harvested 48 hours after infection. The infected cells and supernatant fluid were frozen and thawed three times and clarified by centrifuging at 2,000× g for 20 min at 4 C. Polyethylene glycol 6000 and NaCl were added to the supernatant to final concentrations of 7% and 380 mM, respectively, and the mixture was stirred overnight at 4 C. The precipitated virions were recovered by centrifuging at 10,000× g for 15 min at 4 C and resuspended in 200-400 μl TNE buffer (10 mM Tris-HCl pH 8.0, 100 mM NaCl, 1 mM EDTA) containing 1% NP40. The suspension was clarified by centrifuging at 12,000× g for 3 min before layering onto 15% to 45% (wt/vol) linear sucrose gradients (35 ml) in TNE buffer and centrifuging at 100,000× g for 4 h at 4 C. Gradients were fractioned and the fractions analyzed by SDS-PAGE. Viral fractions were pooled, centrifuged at 200,000× g for 2 h at 4 C, and the viral pellet was resuspended in a small volume of TNE buffer, cDNA synthesis and cloning. Viral RNA was reverse transcribed using an oligo-dT primer (Amersham) or ERhV1 specific primers P1 (SEQ ID NO:18) (5′-ATCCAGCAAGCCGCTGTCCGGTTAC-3′) and P5 (SEQ ID NO:19) (5′-CGAAGAGACACCTGCTTC-3′). Viral RNA was prepared as described in (1987) Anal-Biochem. 162, 156-159. 
     Viral RNA and 100 pmol of primer were mixed, boiled for 2 min and cooled at room temperature. Fist strand cDNA was synthesized using 200 U of Maloney murine leukemia virus reverse transcriptase (Promega) in the presence of 0.8 mM dNTPs and 30 U of human placental RNAse inhibitor (Pharmacia) in a reaction volume of 25 μl. Second strand cDNA was synthesized using a cDNA synthesis kit (Amersham). The cDNA fragments were ligated into pUC18, either as blunt ended fragments or after ligating BamH I adaptors (Pharmacia), and the lighted products used to transform  E. coli  strain DH5α (Stratagene). Colonies were selected by hybridization, initially with an [32P]-dCTP-labelled cDNA probe derived from reverse transcribed viral RNA, and subsequently with [32P]-dCTP-labelled cloned viral cDNA (16). The sequence between two cDNA clones was obtained using the oligonucleotide primers P6 (SEQ ID NO: 20) (5′-TTCTGGTGGAGAAGTGACAGC-3′) and P7 (SEQ ID NO: 21) (5′-GTGAGCCAGCAACAATTGC-3′) in a polymerase chain reaction (PCR; 17) using the polymerase, Vent Exo+ (New England Biolabs). 
     DNA sequencing and analyses 
     Double-stranded DNA was prepared using the alkaline lysis method and sequenced by dideoxy chain termination using modified T7 DNA polymerase (Pharmacia) and [35S]-dATP (Amersham). Sequence was read and analyzed using the GeneWorks software package (IntelliGenetics, Mountain View, Calif.). The GenBank database was searched using the FASTA searching and comparison program. The protein alignment shown in FIG. 3 was performed using the Genetics Computer Group, Inc. (Madison, Wis., USA, 1994) GAP program with a gap creation penalty (GCP) of 3.0 and a gap extension penalty (GEP) of 0.1. The multiple alignments of nucleotide sequences were performed using ClustalW. For pairwise alignments the slow method was used with a GCP of 10 and a GEP of 0.1. For multiple alignments a GCP of 10 and a GEP of 0.05 was used with alignment of sequences which were more than 60% divergent delayed and using weighted transitions. Phylogenetic relationships were examined using the maximum likelihood method with the DNAML program of the Phylogeny Inference Package (Phylip) version 3.5c (1993, J. Felsenstein, Department of Genetics, University of Washington, Seattle). The model used allowed for unequal expected frequencies of the four nucleotides, with the frequencies determined empirically from those present in the sequences analysed, and unequal rates of transitions and transversions. A single rate of change was assumed for all sites. The program was allowed to perform global rearrangements to optimize the tree. Initial analyses were performed on polymerase sequences using a range of transition/transversion ratios to determine that which gave the maximal log likelihood. A ratio of 2.0 gave the maximal log likelihood and thus this ratio was used for all subsequent analyses of other sequences. 
     Cloning and Sequencing of the ERhV1 Genome 
     Sixty seven overlapping cDNA clones and one PCR product clone were obtained and sequenced from both ends. The nucleotide in each position was determine at least twice, and 95% of the sequence was obtained by sequencing in both directions. The predicted genomic structure of ERhV1 was characteristic of picornaviruses, possessing one long open reading frame (ORF) flanked by 5′- and 3′-NTR&#39;s (FIG.  1 ). 
     The nucleotide and predicted amino acid sequences of the ERhV1 polyprotein are shown in FIG. 2 a.  Partial sequence of the 5′-NTR (433 bases) was also obtained FIG. 2 b.  There was a tract of 9 Cs at position −550 to −542. PolyC tracts of various lengths have been observed in similar locations in FMDV and EMCV. The actual length of the ERhV1 polyC tract is uncertain as these sequences are known to be unstable when propagated in  E. coli.  A 14 nucleotide polypyrimidine tract, which possessed the TTTC motif common to all picornaviruses, was present near the potential translation initiation codons. A region of 450 nucleotides upstream of the most likely initiation codon is predicted to contain an internal ribosome entry site (IRES). This region showed most sequence identity (48-50%) with corresponding sequences in FMDV and EMCV. The 3′-NTR of ERhV1 was 102 nucleotides excluding the polyA tail (data not shown). 
     In picornaviruses, there are two factors that influence which ATG codon initiates translation, a requirement for the ATG to be located at the 3′-end of the IRES, and that this ATG occurs in a sequence optimal for initiating translation, that is, a purine at position −3 and a G in position +4. Two pairs of in-frame ATG codons were identified in the ERhV1 genome. The second ATG of the first pair is separated by 25 nucleotides from the beginning of the polypyrimidine tract (FIG. 2 b ), similar to the distance (25 to 27 nucleotides) found in the corresponding regions in FMDV and EMCV (24). The second ATG of each pair occurs in an optimal context. Therefore, the second ATG of the first pair is most likely to be the translation initiation codon but it is possible that translation is also initiated from the second optimal ATG, by a process of leaky scanning, or even from the other two, non-optimal ATG codons. The predicted ERhV1 coding sequence, beginning at the most likely initiation ATG, extended for 6,741 bases and would encode a polyprotein of 2,247 amino acids. 
     Alignment of the ERhV1 amino acid sequence with those of other picornaviruses showed that it was most similar to aphthoviruses and, to a lesser extent, to cardioviruses in all regions of the genome (data not shown). FIG. 3 shows a comparison of the predicted amino acid sequence of ERhV1 with that of FMDV.O1K. The two sequences were 40% identical. The more conserved regions include: the 3D/polymerase (50% identity), VP4 (49% identity) and some regions of the 2C protein. ERhV1 encodes a 2A protein of 16 amino acids, 14 of which were identical with those of FMDV 2A. ERhV1 possessed only one copy of the VPg sequence. This is in contrast to FMDV which has 3 tandemly repeated, non-identical VPg sequences (27-29). 
     Table 1 shows the proteolytic cleavage sites of ERhV1 predicted from the amino acid alignment (FIG.  3 ), and compares these with those of FMDV, EMCV and Theiler&#39;s murine encephalomyelitis virus (TMEV). Most of the ERhV1 cleavage sites could be assigned with reasonable confidence because of significant amino acid similarity with FMDV in the regions flanking the predicted cleavage site; an exception was the 3A/3B cleavage site where there was less sequence similarity. As is the case with FMDV, the predicted ERhV1 3C protease cleavage sites were more variable than those of the cardioviruses. EMCV and TMEV. 
     
       
         
               
             
               
               
               
             
               
               
               
               
               
             
           
               
                 TABLE 1 
               
             
             
               
                   
               
               
                 Comparison of the predicted proteolytic cleavage sites of the ERhVI 
               
               
                 polyprotein with those of FMDV, EMCV and TMEV. 
               
             
          
           
               
                   
                 Cleavage sites* 
                   
               
             
          
           
               
                 Proteins 
                 ERhV1 
                 FMDV 
                 EMCV 
                 TMEV 
               
               
                   
               
               
                 Leader/1A(VP4) 
                 S/G 
                 K/G 
                 Q/G 
                 Q/G 
               
               
                 1A(VP4)/1B(VP2) 
                 A/D 
                 A/D 
                 A/D 
                 L/D 
               
               
                 1B(VP2)/1C(VP3) 
                 E/A 
                 E/G 
                 Q/S 
                 Q/S 
               
               
                 1C(VP3)/1D(VP1) 
                 Q/V 
                 E/T 
                 Q/G 
                 Q/G 
               
               
                 1D(VP1)/2A 
                 T/N 
                 L/N 
                 E/S 
                 E/N 
               
               
                 2A/2B 
                 NPG/P 
                 NPG/P 
                 NPG/P 
                 NPG/P 
               
               
                 2B/2C 
                 Q/V 
                 Q/L 
                 Q/S 
                 Q/G 
               
               
                 2C/3A 
                 Q/S 
                 Q/I 
                 Q/G 
                 Q/S 
               
               
                 3A/3B 
                 Q/S 
                 E/G 
                 Q/G 
                 Q/G 
               
               
                 3B/3C 
                 E/T 
                 E/S 
                 Q/G 
                 Q/G 
               
               
                 3C/3D 
                 Q/G 
                 E/G 
                 Q/G 
                 Q/G 
               
               
                   
               
               
                 *Cleavage data from: FMDV.O1K (Forss et al. 1984), TMEV (Pevear et al. 1987) and EMCV (Palmenberg et al. 1984). The single amino acid code is used.  
               
             
          
         
       
     
     Phylogenetic analyses 
     A phylogenetic tree was derived from the nucleotide sequences of complete picornavirus polyproteins (FIG. 4 a ). Each branch of this tree was statistically, highly significant (P&lt;0.01), with the 95% confidence limits ranging from ±7% to ±15% of branch lengths. ERhV1 was found to be most closely related to the aphthoviruses, although it was clear that ERhV1 was considerably more distant from individual members of this genus than the aphthoviruses were from each other. A phylogenetic tree was also derived from the nucleotide sequences of picornavirus polymerase genes (FIG. 4 b ). Each branch of this tree was statistically, highly significant (P&lt;0.01) with 95% confidence limits ranging from ±14% to ±38% of the branch lengths. Again, ERhV1 grouped with the aphthoviruses and the topology of the tree was the same as that obtained using data of the entire polyprotein (FIG. 4 a ). The VP1 nucleotide sequences were also similarly analyzed (FIG. 4 c ). Most branches were statistically, highly significant (P&lt;0.01), although, that between the ERhV1 branch point and the branch point for the echovirus 22-hepatovirus cluster was less so (P&lt;0.05). The 95% confidence limits of the branch lengths of this tree were considerable greater than for the other two trees, ranging from ±18% to ±69%. This tree did not group ERhV1 with the aphthoviruses. With the exception of bovine enterovirus (BEV), the tree had the same topology as those derived from the complete polyprotein and the polymerase sequences. It was also apparent that picornaviruses formed three clusters: enteroviruses-rhinoviruses, echovirus 22-hepatovirus and cardioviruses-aphthoviruses-ERhV1. 
     (1) Diagnostic Reagents 
     Oligonucleotide primers: We have designed short oligonucleotide primers and used them in polymerase chain reactions (PCR) for the diagnosis of ERhV infected horses. Any of the ERhV nucleotide sequences may be used for the design primer sets for use as diagnostic reagents. They may be highly specific for ERhV1 or they may be designed to be more cross reactive so as to amplify single strand RNA template from other ERhV types e.g., ERhV 2, 3 and 4. As a specific example we have used the primer set shown in FIG. 5 to diagnose ERhV disease in several groups of seriously ill horses in circumstances in which, despite exhaustive efforts, we could not isolate the virus using conventional cell culture procedures. We no consider ERhV a very under reported disease simply because, most of the time, nasal samples collected from horses experiencing severe, systemic clinical disease because of ERhV infection do not yield the virus in cell culture. In one particular group of horses, we detected the presence of ERhV by PCR and confirmed that the horses were both actively infected and seriously ill with ERhV by use of paired serum samples which showed that there was a concomitant rise in ERhV1 serum neutralising antibody. Vigorous attempts to isolate the virus in cell cultures yielded negative results. 
     Oligonucleotide probes: Virus specific oligonucleotides are used as probes to detect the presence of the virus in infected samples from diseased horses and other animals. This may be especially important given the systemic nature of the illness, i.e., it is a foot-and-mouth-like, generalized disease with virus distributed throughout the body in many organs and tissues; it is not just a simple “common cold-like” illness as the name rhinovirus implies. The significance of the sequence in moving the virus out of the Rhinovirus genus and into a new genus proposed to be called “Equirhinovirus” in the Picornaviridae family does not represent merely a taxonomic change but represents a paradigm shift in how ERhV1 and related viruses must now be regarded as pathogens for the horse and other animal species. 
     Diagnostic antigens: Individual virion proteins, in particular VP1, VP2 and VP3, can be expressed in any one of a number of heterologous expression systems to provide antigens to detect specific antibody to ERhV1 present in blood. Such expression systems, which are well established for  E. coli,  yeast, vaccinia virus and baculovirus, allow for the production of large quantities of protein to a high degree of purity. The expressed virion proteins may be used in simple immunoassays, such as ELISA, to detect ERhV1 specific antibody. Virion proteins expressed in this way also serve as effective vaccines against ERhV1 disease. 
     (2) Vaccines 
     Production of virus like particles (VLPs): We have used the sequence information to construct recombinant plasmids containing the P1-2A-3C region of the genome (see FIG. 1 a  and FIG.  6 ). These plasmid constructions are of course critically dependent on the ERhV1 sequence that has been determined although the strategy that we are adopting, in general, is similar to that described in J. Virol 66, 4557-4564. Some early plasmid constructions have been inserted into  E. coli  and baculovirus expression systems based on prior art with similar viruses such as poliomyelitis of humans and foot-and-mouth disease virus of cattle and other cloven hoofed animals. The RT PCR double stranded DNA of the P1-2A-3C region of the ERhV1 genome is transcribed, within the transformed  E. coli  or insect cell for baculovirus, into messenger RNA as a single transcript which is then translated into a mini polyprotein. The 3C protease activity results in the cleavage of the mini polyprotein into its constituent parts namely 1A (VP4), 1B (VP2), 1C (VP3) and 1D(VP1), 2A and 3C (see FIG. 1 a  and FIG. 6) and that the VP component parts then self assemble into VLPs i.e., virus particles that lack nucleic acid and are therefore non infectious i.e., are unstable to cause disease. Two important applications of ERhV VLPs are as follows: 
     (a) The VLPs are very useful as highly effective, safe, high antigen-mass vaccines for the control ERhV1 disease. If ERhV1 disease is confirmed, as we believe to be the case, as significant and responsible for much hither to undiagnosed illness that results in many lost training days, many expensive treatments, much serious illness because of secondary infections following on the primary ERhV1 infection, and much poor performance, then the utility of the vaccine based on the VLPs that are the subject of this invention will be very great and likely to have world-wide application. 
     With improved methods for the diagnosis of ERhV1 infection such as by PCR and ELISA as described herein, it is likely that other members of the proposed new Equirhinovirus genus within the family Picornaviridae including for example ERhV2, ERhV3, may be similarly diagnosed. Indeed suitably selected PCR primer sets based on the ERhV1 sequence could be used to detect these other equine rhinoviruses. The sequencing of these genomes could provide a basis for their specific diagnosis. It is also evident that the construction of VLP&#39;s based on expression plasmids similar to those described herein for ERhV1, could be readily adapted to these other equine rhinoviruses leading for example to production of combined ERhV vaccines to cover all antigenic types as may be extant or as may emerge by antigenic variation, as is very much a part of the biology of FMDV, in the future. Polyvalent VLP vaccines incorporating a range of ERhV antigenic types are obvious extensions based on the work described herein. 
     (b) ERhV VLPs can be used as a delivery vector that will provide not only protection against ERhV disease but will be used to deliver other therapeutic and useful substances to the horses following administration by parenteral or other routes. Such delivery vectors can be produced by inserting into, for example the P1 region at some appropriate site, double stranded DNA coding for antigenic epitopes of other virus and infections agents of horse as well as epitopes derived from other non infectious sources for example reproductive hormones. 
     ERhV1 Diagnostic Tests 
     For the detection of ERhV1 antibodies in infected or vaccinated horses various stranded tests can be used. VLP&#39;s may be used in such test for example in an ELISA test for antibody. 
     Other diagnostic tests based on recombinant antigens derived from the ERhV1 sequence can be devised along similar lines to those reported for FMDV in which the absence of protein 2C from clarified inactivated whole virus FMD, FMDV or FMDV VLP vaccines may be used as the basis for distinguishing infected from vaccinated animals where the vaccine is a non-replicating form of ERhV1 or a deletion mutant of ERhV1 in which a particular non-structural protein gene has been deleted. Precedent for this comes from studies of FMDV as reported in for example Lubroth, Grubman, Burrage, Newman &amp; Brown, 1996, Absence of protein 2C from clarified foot-and-mouth disease virus vaccines provides the basis for distinguishing convalescent from vaccinated animals, Vaccine 14(5), 419-427. 
     Preparation and Use of Virus-Like Particles and Other Proteins Based on ERhV1 Sequence 
     From the sequence of ERhV1 it is possible to clone certain segments of the viral genome into a variety of vectors for expression in a variety of different expression systems. There is a straight forward and storing literature for FMDV that provides a very clear precedent for what can be done for ERhV1. Examples include the expression of FMDV P1-2A in a baculovirus (Abrams C C &amp; Belsham G J, 1994, The antigenicity of foot-and-mouth disease virus P1-2A polyprotein and empty capsids produced in vaccinia virus and baculovirus expression systems. In VIIth Meeting of the European Study Group on the Molecular Biology of Picornaviruses, Aug. 6-11, 1994, Korpilampi, Finland) or vaccina virus systems (Abrams C C, King A M Q &amp; Belsham G J, 1995, Assembly of foot-and-mouth disease virus empty capsids synthesized by a vaccinia virus expression system. Journal of General Virology 76:3089-3098) to obtain VLPs or viral proteins. We have prepared similar plasmids in which P1-2A, P1-2A-3C and these two sequences in a myristolated form have been inserted into p fastbac 1 baculovirus vector (Gibco/BRL) and into a PET vector (Novogene) for expression in insect cells and  E. coli  respectively. 
     These expressed products either as protein antigens or as VLPs, have utility as the basis for diagnostic tests or vaccines. 
     Accordingly, such references are herein incorporated in support of the full description and enablement of the invention where the disclosed methods of preparing diagnostics, vaccines, vectors, host systems and kits are fully described and applicable to the like aspects of the current invention. 
     (3) Applications in human medicine: 
     ERhV is also a human pathogen. We have unpublished data to confirm that humans have serum neutralising antibody to ERhV1 that is indicative of infection. One of the laboratory workers connected with the conduct of the sequencing and who handled infections virus has specific antibody in high amounts (serum neutralising antibody titre 1:640 to ERhV1). We are currently extending these studies and anticipate finding a significant incidence of infection in humans world wide particularly among those humans who work with horses. The improved diagnostic methods outlined above, perhaps also the vaccine, are expected to have application in human medicine. 
     
       
         
           
             25 
           
           
             1 
             7278 
             DNA 
             equine rhinovirus 1 
             
               CDS 
               (436)..(7176) 
             
           
            1
ccgtcaagcc cgttgcctgt atagccaggt aaccggacag cggcttgctg gattttcccg     60
gtgccattgc tctggatggt gtcaccaagc tgacaaatgc ggagtgaacc tcacaaagcg    120
acacgcctgt ggtagcgctg cccaaaaggg agcggaactc cccgccgagg cggtcctctc    180
tggccaaaag cccagcgttg atagcgcctt ttgggatgca ggaaccccac ctgccaggtg    240
tgaagtggag tgagcggatc tccaatttgg tctgttctga actacaccat ttactgctgt    300
gaagaatgcc ctggaggcaa gctggttaca gccctgacca ggccctgccc gtgactctcg    360
accggcgcag ggtcaaaaat tgtctaagca gcagcaggaa cgcgggagcg tttcttttcc    420
ttttgtactg acatg atg gcg gcg tct aag gtg tat aga gtt tgc gag cag     471
                 Met Ala Ala Ser Lys Val Tyr Arg Val Cys Glu Gln
                   1               5                  10
act ctg ctg gca ggt gcc gtt cgc atg atg gac aaa ttc ttg caa aag      519
Thr Leu Leu Ala Gly Ala Val Arg Met Met Asp Lys Phe Leu Gln Lys
         15                  20                  25
aga act gtt ttt gtc ccc cat ctt gac aaa aca att cgt ttg act gga      567
Arg Thr Val Phe Val Pro His Leu Asp Lys Thr Ile Arg Leu Thr Gly
     30                  35                  40
ctc cac aat tat gac aat act tgc tgg ttg aat gcc ttg aca caa ctg      615
Leu His Asn Tyr Asp Asn Thr Cys Trp Leu Asn Ala Leu Thr Gln Leu
 45                  50                  55                  60
aca cag att ctt gga att cgg ctt ttt gat gaa cac ttc ggc aat aga      663
Thr Gln Ile Leu Gly Ile Arg Leu Phe Asp Glu His Phe Gly Asn Arg
                 65                  70                  75
ggt ctg ttc act cgg aaa aca att gat tgg gtg agt gac cag act ggt      711
Gly Leu Phe Thr Arg Lys Thr Ile Asp Trp Val Ser Asp Gln Thr Gly
             80                  85                  90
ata aaa gat cta aaa tca gga gca ccg cca ctc gtg gtg gtg tac aaa      759
Ile Lys Asp Leu Lys Ser Gly Ala Pro Pro Leu Val Val Val Tyr Lys
         95                 100                 105
ctg tgg caa cat gga cac ttg gat gtc ggt acg atg gag aaa ccc cgg      807
Leu Trp Gln His Gly His Leu Asp Val Gly Thr Met Glu Lys Pro Arg
    110                 115                 120
tcg att act cta tgg tct ggc ccc aaa gtg tgt ctt tct gat ttc tgg      855
Ser Ile Thr Leu Trp Ser Gly Pro Lys Val Cys Leu Ser Asp Phe Trp
125                 130                 135                 140
gcc tgt gtt tcg gca aaa ccg gga cat gca gta ttc tac ctt ctc aca      903
Ala Cys Val Ser Ala Lys Pro Gly His Ala Val Phe Tyr Leu Leu Thr
                145                 150                 155
agc gag ggt tgg atc tgt gtt gat gac aag aaa ata tac cca gaa aca      951
Ser Glu Gly Trp Ile Cys Val Asp Asp Lys Lys Ile Tyr Pro Glu Thr
            160                 165                 170
ccc aaa aca gag gat gta ctt gtt ttt gcg ccc tat gac ttt gag tca      999
Pro Lys Thr Glu Asp Val Leu Val Phe Ala Pro Tyr Asp Phe Glu Ser
        175                 180                 185
ctg ggc aag gac cca cca aag cta cac cag aga tat gaa aaa gca ttt     1047
Leu Gly Lys Asp Pro Pro Lys Leu His Gln Arg Tyr Glu Lys Ala Phe
    190                 195                 200
gag ctc agt ggc gga ggt aca tcc act cca aca act ggc aac caa aac     1095
Glu Leu Ser Gly Gly Gly Thr Ser Thr Pro Thr Thr Gly Asn Gln Asn
205                 210                 215                 220
atg tcc gga aac agt ggt tca att gtt caa aat ttt tac atg caa cag     1143
Met Ser Gly Asn Ser Gly Ser Ile Val Gln Asn Phe Tyr Met Gln Gln
                225                 230                 235
tac cag aat tca att gac gca gac ctg gga gac aat gtg att agc cct     1191
Tyr Gln Asn Ser Ile Asp Ala Asp Leu Gly Asp Asn Val Ile Ser Pro
            240                 245                 250
gaa ggc cag ggc agc aac act agt agt tca acc tca tca agc caa tcc     1239
Glu Gly Gln Gly Ser Asn Thr Ser Ser Ser Thr Ser Ser Ser Gln Ser
        255                 260                 265
tct ggc ttg ggc ggg tgg ttc tct agt ttg ctg aac ctt gga aca aaa     1287
Ser Gly Leu Gly Gly Trp Phe Ser Ser Leu Leu Asn Leu Gly Thr Lys
    270                 275                 280
cta ctg gct gac aag aag aca gaa gag act aca aac att gaa gac aga     1335
Leu Leu Ala Asp Lys Lys Thr Glu Glu Thr Thr Asn Ile Glu Asp Arg
285                 290                 295                 300
att gaa aca aca gtg gtt gga gtc act att att aat tca caa gga tct     1383
Ile Glu Thr Thr Val Val Gly Val Thr Ile Ile Asn Ser Gln Gly Ser
                305                 310                 315
gtt gga aca acc tac tgt tac tcc aaa ccg gat ggt aga cca cca tcc     1431
Val Gly Thr Thr Tyr Cys Tyr Ser Lys Pro Asp Gly Arg Pro Pro Ser
            320                 325                 330
aca gtg tca gac cca gtt acc aga ctt gga ccc acg ctt tcc agg cac     1479
Thr Val Ser Asp Pro Val Thr Arg Leu Gly Pro Thr Leu Ser Arg His
        335                 340                 345
tac aca ttt aag gta ggt gag tgg ccc cat tct caa tca cat ggt cac     1527
Tyr Thr Phe Lys Val Gly Glu Trp Pro His Ser Gln Ser His Gly His
    350                 355                 360
gca tgg atc tgt ccg ttg cca ggt gac aaa ctc aag aag atg ggc agt     1575
Ala Trp Ile Cys Pro Leu Pro Gly Asp Lys Leu Lys Lys Met Gly Ser
365                 370                 375                 380
ttt cat gag gtt gtc aaa gcc cac cac ctg gtc aag aac ggc tgg gat     1623
Phe His Glu Val Val Lys Ala His His Leu Val Lys Asn Gly Trp Asp
                385                 390                 395
gtg gtt gtg cag gtg aat ccc tca ttt gct cac tcc ggg ccg ctg tgt     1671
Val Val Val Gln Val Asn Pro Ser Phe Ala His Ser Gly Pro Leu Cys
            400                 405                 410
gta gca gca gtg ccg gag tac gaa cac aca cat gag aaa gca ctc aag     1719
Val Ala Ala Val Pro Glu Tyr Glu His Thr His Glu Lys Ala Leu Lys
        415                 420                 425
tgg tct gag ctt gag gaa cca gct tac aca tac caa caa ctt tca gtt     1767
Trp Ser Glu Leu Glu Glu Pro Ala Tyr Thr Tyr Gln Gln Leu Ser Val
    430                 435                 440
ttt ccc cac cag ttg cta aat ttg agg aca aat tca tca gtg cat ttg     1815
Phe Pro His Gln Leu Leu Asn Leu Arg Thr Asn Ser Ser Val His Leu
445                 450                 455                 460
gtg atg ccc tac att ggg cca ggc caa cca aca aat ctg act ttg cac     1863
Val Met Pro Tyr Ile Gly Pro Gly Gln Pro Thr Asn Leu Thr Leu His
                465                 470                 475
aac ccg tgg acc att gtt att tta att ttg tct gaa ttg aca gga cct     1911
Asn Pro Trp Thr Ile Val Ile Leu Ile Leu Ser Glu Leu Thr Gly Pro
            480                 485                 490
ggc caa act gtg cct gtg acc atg tcg gtg gct ccc atc gat gca atg     1959
Gly Gln Thr Val Pro Val Thr Met Ser Val Ala Pro Ile Asp Ala Met
        495                 500                 505
gtt aat ggg cct ctt cca aat cca gag gca ccg att aga gtg gtg tct     2007
Val Asn Gly Pro Leu Pro Asn Pro Glu Ala Pro Ile Arg Val Val Ser
    510                 515                 520
gtg cct gaa tca gat tct ttt atg tct tca gta cct gat aat tcg act     2055
Val Pro Glu Ser Asp Ser Phe Met Ser Ser Val Pro Asp Asn Ser Thr
525                 530                 535                 540
cca cta tac ccc aag gtt gtg gtc cca ccg cgc caa gtt cct ggc cgg     2103
Pro Leu Tyr Pro Lys Val Val Val Pro Pro Arg Gln Val Pro Gly Arg
                545                 550                 555
ttt aca aat ttc att gat gtg gca aaa cag aca tat tca ttt tgt tcc     2151
Phe Thr Asn Phe Ile Asp Val Ala Lys Gln Thr Tyr Ser Phe Cys Ser
            560                 565                 570
att tct gga aaa cct tat ttt gag gtt acc aac acc tct ggg gac gag     2199
Ile Ser Gly Lys Pro Tyr Phe Glu Val Thr Asn Thr Ser Gly Asp Glu
        575                 580                 585
cca ctg ttt cag atg gat gtg tcg ctc agt gcg gca gag cta cat ggc     2247
Pro Leu Phe Gln Met Asp Val Ser Leu Ser Ala Ala Glu Leu His Gly
    590                 595                 600
act tac gta gct agt ttg tca tca ttt ttt gca cag tac aga ggc tca     2295
Thr Tyr Val Ala Ser Leu Ser Ser Phe Phe Ala Gln Tyr Arg Gly Ser
605                 610                 615                 620
ctt aat ttc aac ttt att ttc act ggt gca gca gcc act aag gca aag     2343
Leu Asn Phe Asn Phe Ile Phe Thr Gly Ala Ala Ala Thr Lys Ala Lys
                625                 630                 635
ttt ctg gtt gct ttt gtg cct ccc cac agt gca gcg ccc aaa acg cgc     2391
Phe Leu Val Ala Phe Val Pro Pro His Ser Ala Ala Pro Lys Thr Arg
            640                 645                 650
gat gaa gca atg gcg tgc atc cat gcc gtg tgg gat gtt ggc ttg aac     2439
Asp Glu Ala Met Ala Cys Ile His Ala Val Trp Asp Val Gly Leu Asn
        655                 660                 665
tca gct ttt tct ttt aat gta cct tat ccc tcc cct gct gac ttc atg     2487
Ser Ala Phe Ser Phe Asn Val Pro Tyr Pro Ser Pro Ala Asp Phe Met
    670                 675                 680
gcc gtt tat tct gcg gaa cgg acg gtt gtg aat gtc tct gga tgg ctt     2535
Ala Val Tyr Ser Ala Glu Arg Thr Val Val Asn Val Ser Gly Trp Leu
685                 690                 695                 700
caa gtt tat gca cta aca gct cta act tca act gac att gcc gtg aac     2583
Gln Val Tyr Ala Leu Thr Ala Leu Thr Ser Thr Asp Ile Ala Val Asn
                705                 710                 715
agt aaa ggc cgt gtg ctg gtt gct gtt tcc gcc ggc cca gac ttc tcc     2631
Ser Lys Gly Arg Val Leu Val Ala Val Ser Ala Gly Pro Asp Phe Ser
            720                 725                 730
ctt cgt cac ccg gcg gac ctg ccc gac aag cag gtt acc aat gtg gga     2679
Leu Arg His Pro Ala Asp Leu Pro Asp Lys Gln Val Thr Asn Val Gly
        735                 740                 745
gag gat ggt gaa ccc ggt gag aca gag cct cgt cat gct ttg tca ccc     2727
Glu Asp Gly Glu Pro Gly Glu Thr Glu Pro Arg His Ala Leu Ser Pro
    750                 755                 760
gtg gac atg cac gtg cac aca gat gtc agt ttc ttg ctt gac cgg ttc     2775
Val Asp Met His Val His Thr Asp Val Ser Phe Leu Leu Asp Arg Phe
765                 770                 775                 780
ttt gat gtt gag aca ctt gag ctt tca aat ttg aca ggt tct cct gcc     2823
Phe Asp Val Glu Thr Leu Glu Leu Ser Asn Leu Thr Gly Ser Pro Ala
                785                 790                 795
aca cat gtt ctg gat ccg ttt ggc tcg act gcc caa ctg gct tgg gca     2871
Thr His Val Leu Asp Pro Phe Gly Ser Thr Ala Gln Leu Ala Trp Ala
            800                 805                 810
cgt ctg cta aac act tgc acc tac ttc ttt tct gat ttg gaa ttg tca     2919
Arg Leu Leu Asn Thr Cys Thr Tyr Phe Phe Ser Asp Leu Glu Leu Ser
        815                 820                 825
atc cag ttt aaa ttt acc acc act ccg tcc tct gtt gga gag ggc ttt     2967
Ile Gln Phe Lys Phe Thr Thr Thr Pro Ser Ser Val Gly Glu Gly Phe
    830                 835                 840
gtg tgg gtg aag tgg ctc cct gtt gga gca cca acc aag acc aca gat     3015
Val Trp Val Lys Trp Leu Pro Val Gly Ala Pro Thr Lys Thr Thr Asp
845                 850                 855                 860
gct tgg cag tta gaa gga ggt gga aat tca gtt aga att caa aaa ttg     3063
Ala Trp Gln Leu Glu Gly Gly Gly Asn Ser Val Arg Ile Gln Lys Leu
                865                 870                 875
gcc gtt gca ggg atg tgc ccc act gtt gtg ttc aag att gca ggc tcc     3111
Ala Val Ala Gly Met Cys Pro Thr Val Val Phe Lys Ile Ala Gly Ser
            880                 885                 890
cgt tca caa gcc tgt gct tca gcg ttg cca tat aca tca atg tgg cgt     3159
Arg Ser Gln Ala Cys Ala Ser Ala Leu Pro Tyr Thr Ser Met Trp Arg
        895                 900                 905
gtt gtg cca gtc ttt tac aat ggc tgg ggt gca cct acc aaa gaa aag     3207
Val Val Pro Val Phe Tyr Asn Gly Trp Gly Ala Pro Thr Lys Glu Lys
    910                 915                 920
gca acc tac aat tgg ctt cct ggt gca cac ttt ggt tcc atc ttg ctg     3255
Ala Thr Tyr Asn Trp Leu Pro Gly Ala His Phe Gly Ser Ile Leu Leu
925                 930                 935                 940
act tct gat gcg cat gat aaa gga ggg tgc tac ttg cgg tat gct ttc     3303
Thr Ser Asp Ala His Asp Lys Gly Gly Cys Tyr Leu Arg Tyr Ala Phe
                945                 950                 955
cgc gcg cca gcg atg tat tgc cct cga ccc att ccg ccg gct ttt acg     3351
Arg Ala Pro Ala Met Tyr Cys Pro Arg Pro Ile Pro Pro Ala Phe Thr
            960                 965                 970
cgt cca gcg gac aaa acc aga cat aaa ttt ccc act aac atc aac aaa     3399
Arg Pro Ala Asp Lys Thr Arg His Lys Phe Pro Thr Asn Ile Asn Lys
        975                 980                 985
cag tgt act aat tac tct ctc ctc aaa ttg gct gga gat gtt gag agc     3447
Gln Cys Thr Asn Tyr Ser Leu Leu Lys Leu Ala Gly Asp Val Glu Ser
    990                 995                1000
aac cct ggc ccc act att ttt tcc aaa gca tca gca gac ctg aat gcc     3495
Asn Pro Gly Pro Thr Ile Phe Ser Lys Ala Ser Ala Asp Leu Asn Ala
1005               1010                1015                1020
ttg tca acg tcg cta ggt gaa ttg act ggc atg cta aaa gat ctt aaa     3543
Leu Ser Thr Ser Leu Gly Glu Leu Thr Gly Met Leu Lys Asp Leu Lys
               1025                1030                1035
gcc aag gca gaa act tat tcc ccg ttt tac aaa atg gcc aaa atg ctt     3591
Ala Lys Ala Glu Thr Tyr Ser Pro Phe Tyr Lys Met Ala Lys Met Leu
           1040                1045                1050
ttc aaa ctt gca aca cta gct gtg gca gct atg agg aca aag gac cca     3639
Phe Lys Leu Ala Thr Leu Ala Val Ala Ala Met Arg Thr Lys Asp Pro
       1055                1060                1065
gta gtg gtg gtt atg ttg att gct gat ttc gga ttg gag gtc ttt gac     3687
Val Val Val Val Met Leu Ile Ala Asp Phe Gly Leu Glu Val Phe Asp
   1070                1075                1080
act ggg ttt ttc ttt tcc tac ttt caa gag aag ttg cag cct tat atg     3735
Thr Gly Phe Phe Phe Ser Tyr Phe Gln Glu Lys Leu Gln Pro Tyr Met
1085               1090                1095                1100
aaa act att cct ggt aag att tct gat ttg gtc act gat gcg gct acg     3783
Lys Thr Ile Pro Gly Lys Ile Ser Asp Leu Val Thr Asp Ala Ala Thr
               1105                1110                1115
gct gcc gcc caa att cca aag gga gtg tat tct ttt gtg tcg tca ttt     3831
Ala Ala Ala Gln Ile Pro Lys Gly Val Tyr Ser Phe Val Ser Ser Phe
           1120                1125                1130
ttc gaa acg cct gaa gga gtg gtt gag aag cag gtg tct ctt cgg aca     3879
Phe Glu Thr Pro Glu Gly Val Val Glu Lys Gln Val Ser Leu Arg Thr
       1135                1140                1145
gtg aat gac ata ttt gct ttg ctt aaa aat tct gat tgg ttc ata aag     3927
Val Asn Asp Ile Phe Ala Leu Leu Lys Asn Ser Asp Trp Phe Ile Lys
   1150                1155                1160
act ctt gtt gcc ctc aag aaa tgg ctg aca tcc tgg ttt gct caa gaa     3975
Thr Leu Val Ala Leu Lys Lys Trp Leu Thr Ser Trp Phe Ala Gln Glu
1165               1170                1175                1180
caa cag gca gat gat gcg ctc tat tca gaa ttg gaa aaa tat ccc ttg     4023
Gln Gln Ala Asp Asp Ala Leu Tyr Ser Glu Leu Glu Lys Tyr Pro Leu
               1185                1190                1195
tac aag tta aaa ttg aag gaa cct gat act caa gag gaa gcg cgc cag     4071
Tyr Lys Leu Lys Leu Lys Glu Pro Asp Thr Gln Glu Glu Ala Arg Gln
           1200                1205                1210
tgg ttt aaa gac atg cag cag cgt gct ctc gct gtg aag gac aaa ggt     4119
Trp Phe Lys Asp Met Gln Gln Arg Ala Leu Ala Val Lys Asp Lys Gly
       1215                1220                1225
ctc ttt tcc ctc ctg caa att cca tta gtt aac ttg ccc cag agc cgt     4167
Leu Phe Ser Leu Leu Gln Ile Pro Leu Val Asn Leu Pro Gln Ser Arg
   1230                1235                1240
cca gag ccc gtt gta tgc gtc ctt cgg ggc gca tca ggg caa ggc aaa     4215
Pro Glu Pro Val Val Cys Val Leu Arg Gly Ala Ser Gly Gln Gly Lys
1245               1250                1255                1260
tct tat ttg gca aat ctg atg gct caa gca att tcg ctt ctc ttg gtt     4263
Ser Tyr Leu Ala Asn Leu Met Ala Gln Ala Ile Ser Leu Leu Leu Val
               1265                1270                1275
ggc aag cag gac agt gtg tgg agt tgt cct cct gac ccc aca tat ttt     4311
Gly Lys Gln Asp Ser Val Trp Ser Cys Pro Pro Asp Pro Thr Tyr Phe
           1280                1285                1290
gat ggc tat aac gga cag gct gtg gtg att atg gat gca ttg ggc cag     4359
Asp Gly Tyr Asn Gly Gln Ala Val Val Ile Met Asp Ala Leu Gly Gln
       1295                1300                1305
gat ccg aat ggt gct gac ttt aaa tat ttt tgc cag atg gtc tct aca     4407
Asp Pro Asn Gly Ala Asp Phe Lys Tyr Phe Cys Gln Met Val Ser Thr
   1310                1315                1320
aca gct ttt gta cca cct atg gcc cat ttg gat gat aaa ggc att cca     4455
Thr Ala Phe Val Pro Pro Met Ala His Leu Asp Asp Lys Gly Ile Pro
1325               1330                1335                1340
ttt act tct cct gtt gtt att tgt act aca aat ttg cat tca tct ttt     4503
Phe Thr Ser Pro Val Val Ile Cys Thr Thr Asn Leu His Ser Ser Phe
               1345                1350                1355
acc cct att act gtt tct tgt cct gaa gct ctt aag agg agg ttt cgg     4551
Thr Pro Ile Thr Val Ser Cys Pro Glu Ala Leu Lys Arg Arg Phe Arg
           1360                1365                1370
ttt gat gtg acg gtg tcc gct aaa ccg ggc ttt gtg cgc act gtt ggt     4599
Phe Asp Val Thr Val Ser Ala Lys Pro Gly Phe Val Arg Thr Val Gly
       1375                1380                1385
tca aac cag ctt ttg aat ctc cca ctt gct ctt aag cca gct ggt ctt     4647
Ser Asn Gln Leu Leu Asn Leu Pro Leu Ala Leu Lys Pro Ala Gly Leu
   1390                1395                1400
ccc cca cac cct atc ttt gaa aat gac atg ccc att ata aat ggg cag     4695
Pro Pro His Pro Ile Phe Glu Asn Asp Met Pro Ile Ile Asn Gly Gln
1405               1410                1415                1420
gct gtt aaa ttg gct ctt tct ggt gga gaa gtg aca gct ttt gag ctt     4743
Ala Val Lys Leu Ala Leu Ser Gly Gly Glu Val Thr Ala Phe Glu Leu
               1425                1430                1435
att gag atg ata ctg tca gaa gtt caa aac aga caa gac aca cac aaa     4791
Ile Glu Met Ile Leu Ser Glu Val Gln Asn Arg Gln Asp Thr His Lys
           1440                1445                1450
atg ccc att ttt aaa caa tca tgg tct gat ttg ttc aga aag tgt aca     4839
Met Pro Ile Phe Lys Gln Ser Trp Ser Asp Leu Phe Arg Lys Cys Thr
       1455                1460                1465
act gat gag gaa cag aaa atg ttg cag ttt tta att gac aat aaa gat     4887
Thr Asp Glu Glu Gln Lys Met Leu Gln Phe Leu Ile Asp Asn Lys Asp
   1470                1475                1480
tca gaa att ctc agg gcg ttt gtt tca gaa cgc tcc att tta cta cat     4935
Ser Glu Ile Leu Arg Ala Phe Val Ser Glu Arg Ser Ile Leu Leu His
1485               1490                1495                1500
gaa gag tat ctt aaa tgg gag tca tat atg acc agg aga gcc aag ttt     4983
Glu Glu Tyr Leu Lys Trp Glu Ser Tyr Met Thr Arg Arg Ala Lys Phe
               1505                1510                1515
cac cgc ctg gct gct gat ttt gct atg ttt cta tcc att ctt act tca     5031
His Arg Leu Ala Ala Asp Phe Ala Met Phe Leu Ser Ile Leu Thr Ser
           1520                1525                1530
ctg att gtt att ttt tgt tta gtt tat tct atg tat caa ctt ttt aag     5079
Leu Ile Val Ile Phe Cys Leu Val Tyr Ser Met Tyr Gln Leu Phe Lys
       1535                1540                1545
acc cct gac gag caa tca gct tat gat cct tca act aag cca aaa cca     5127
Thr Pro Asp Glu Gln Ser Ala Tyr Asp Pro Ser Thr Lys Pro Lys Pro
   1550                1555                1560
aag acc cag gaa gtg aaa aca ctg aag att agg act gag act ggt gta     5175
Lys Thr Gln Glu Val Lys Thr Leu Lys Ile Arg Thr Glu Thr Gly Val
1565               1570                1575                1580
cca gca act gac ttg caa caa tcc atc atg aaa aat gtt cag cca att     5223
Pro Ala Thr Asp Leu Gln Gln Ser Ile Met Lys Asn Val Gln Pro Ile
               1585                1590                1595
gag ctt tac ctt gac aat gaa ttg gtt act gac tgc tct gcc ttg ggt     5271
Glu Leu Tyr Leu Asp Asn Glu Leu Val Thr Asp Cys Ser Ala Leu Gly
           1600                1605                1610
gtt tat gac aat tca tat ttg gtg ccc ctt cat ttg ttt gaa ttt gat     5319
Val Tyr Asp Asn Ser Tyr Leu Val Pro Leu His Leu Phe Glu Phe Asp
       1615                1620                1625
ttt gat acc att gtg ctt ggt gga cgt cat tac aag aaa gct gag tgt     5367
Phe Asp Thr Ile Val Leu Gly Gly Arg His Tyr Lys Lys Ala Glu Cys
   1630                1635                1640
gag aag gta gag ttt gag ctt gaa gtg aat gga gac gtg gtg tca tca     5415
Glu Lys Val Glu Phe Glu Leu Glu Val Asn Gly Asp Val Val Ser Ser
1645               1650                1655                1660
gat gcg tgt cta ctt cga gtg tca tcg ggg cct aaa gtt aga aat att     5463
Asp Ala Cys Leu Leu Arg Val Ser Ser Gly Pro Lys Val Arg Asn Ile
               1665                1670                1675
gtt cat ctt ttt aca aat gaa att gaa ttg aag aaa atg acc caa gtg     5511
Val His Leu Phe Thr Asn Glu Ile Glu Leu Lys Lys Met Thr Gln Val
           1680                1685                1690
aca gga atc atg aat tca cca cac cag gca cgc act gtg ttt ttt ggc     5559
Thr Gly Ile Met Asn Ser Pro His Gln Ala Arg Thr Val Phe Phe Gly
       1695                1700                1705
agt ttt ttg aca gtg agg aag tcc atc tta aca tcg gat ggg act gta     5607
Ser Phe Leu Thr Val Arg Lys Ser Ile Leu Thr Ser Asp Gly Thr Val
   1710                1715                1720
atg ccc aat gtt ttg tcc tat gcc gct cag acc tcg cgt ggg tat tgt     5655
Met Pro Asn Val Leu Ser Tyr Ala Ala Gln Thr Ser Arg Gly Tyr Cys
1725               1730                1735                1740
ggc gct gca att gtt gct ggc tca cct gcc cgc ata att ggt atc cat     5703
Gly Ala Ala Ile Val Ala Gly Ser Pro Ala Arg Ile Ile Gly Ile His
               1745                1750                1755
tca gct ggc act gga tct gtt gca ttt tgc tcc ctg gtg tcc aga gac     5751
Ser Ala Gly Thr Gly Ser Val Ala Phe Cys Ser Leu Val Ser Arg Asp
           1760                1765                1770
gcg ctg gag caa ctc tgg ccc cag aaa cag ggc aac gtt agt cgc ctt     5799
Ala Leu Glu Gln Leu Trp Pro Gln Lys Gln Gly Asn Val Ser Arg Leu
       1775                1780                1785
gat gac gat gtg agg gtg tct gtt ccg cgc cgc tcc aaa ttg gtg aaa     5847
Asp Asp Asp Val Arg Val Ser Val Pro Arg Arg Ser Lys Leu Val Lys
   1790                1795                1800
tca ttg gct tac ccc att ttc aaa cct gac tat ggc cca gcg cca ctc     5895
Ser Leu Ala Tyr Pro Ile Phe Lys Pro Asp Tyr Gly Pro Ala Pro Leu
1805               1810                1815                1820
tct caa ttt gac aag cgc ctg tca gac ggc gtg aag ctg gat gaa gtg     5943
Ser Gln Phe Asp Lys Arg Leu Ser Asp Gly Val Lys Leu Asp Glu Val
               1825                1830                1835
gtt ttt gct aaa cat act gga gac aag gag att tcc gca cag gac cag     5991
Val Phe Ala Lys His Thr Gly Asp Lys Glu Ile Ser Ala Gln Asp Gln
           1840                1845                1850
aaa tgg ctc ttg cgt gcg gcg cat gta tac gcc cag aag gtt ttc tcc     6039
Lys Trp Leu Leu Arg Ala Ala His Val Tyr Ala Gln Lys Val Phe Ser
       1855                1860                1865
cgg att gga ttt gac aac cag gct ttg act gaa aaa gag gcc att tgt     6087
Arg Ile Gly Phe Asp Asn Gln Ala Leu Thr Glu Lys Glu Ala Ile Cys
   1870                1875                1880
ggc att cct ggc ctt gac aag atg gag cag gac acc gct ccc ggg ctg     6135
Gly Ile Pro Gly Leu Asp Lys Met Glu Gln Asp Thr Ala Pro Gly Leu
1885               1890                1895                1900
ccc tat gct cag caa aat aag aga agg aaa gac atc tgt gat ttt gaa     6183
Pro Tyr Ala Gln Gln Asn Lys Arg Arg Lys Asp Ile Cys Asp Phe Glu
               1905                1910                1915
gag ggc cgg ctg aag ggc gcc gaa ctc caa aag gac aga ttt atg gct     6231
Glu Gly Arg Leu Lys Gly Ala Glu Leu Gln Lys Asp Arg Phe Met Ala
           1920                1925                1930
ggt gac tac tct aat ttg gtc tat caa tca ttt ttg aaa gat gag atc     6279
Gly Asp Tyr Ser Asn Leu Val Tyr Gln Ser Phe Leu Lys Asp Glu Ile
       1935                1940                1945
cgc cca ctt gag aaa gtt agg gct gga aag acc cgc ctg att gac gtg     6327
Arg Pro Leu Glu Lys Val Arg Ala Gly Lys Thr Arg Leu Ile Asp Val
   1950                1955                1960
ccg ccg atg ccc cat gtg gtg gtt ggt agg cag ctc ttg ggc cgg ttt     6375
Pro Pro Met Pro His Val Val Val Gly Arg Gln Leu Leu Gly Arg Phe
1965               1970                1975                1980
gtg gca aaa ttt cat gaa gca aat gga ttt gac att ggc tca gcc att     6423
Val Ala Lys Phe His Glu Ala Asn Gly Phe Asp Ile Gly Ser Ala Ile
               1985                1990                1995
gga tgt gac cca gat gtg gac tgg act cgg ttt ggc ctc gag ttg gag     6471
Gly Cys Asp Pro Asp Val Asp Trp Thr Arg Phe Gly Leu Glu Leu Glu
           2000                2005                2010
cgt ttc agg tat gta tat gcc tgt gac tac tca cgg ttc gat gcc aac     6519
Arg Phe Arg Tyr Val Tyr Ala Cys Asp Tyr Ser Arg Phe Asp Ala Asn
       2015                2020                2025
cat gca gct gat gca atg aga gtt gtg ctt aac tac ttt ttc tct gag     6567
His Ala Ala Asp Ala Met Arg Val Val Leu Asn Tyr Phe Phe Ser Glu
   2030                2035                2040
gac cac ggt ttc gac cct ggt gtg cct gct ttt att gag tca ctg gtt     6615
Asp His Gly Phe Asp Pro Gly Val Pro Ala Phe Ile Glu Ser Leu Val
2045               2050                2055                2060
gat tca gtg cat gcc tat gaa gag aaa agg tat aac atc tac ggt ggc     6663
Asp Ser Val His Ala Tyr Glu Glu Lys Arg Tyr Asn Ile Tyr Gly Gly
               2065                2070                2075
ttg cca tcc ggg tgt tcc tgc aca tca att ttg aat acc atc ttg aac     6711
Leu Pro Ser Gly Cys Ser Cys Thr Ser Ile Leu Asn Thr Ile Leu Asn
           2080                2085                2090
aat gtt tac att ctt gca gct atg atg aag gct tat gag aat ttt gag     6759
Asn Val Tyr Ile Leu Ala Ala Met Met Lys Ala Tyr Glu Asn Phe Glu
       2095                2100                2105
cca gat gac att cag gtc att tgc tat ggg gac gac tgc ctc att gct     6807
Pro Asp Asp Ile Gln Val Ile Cys Tyr Gly Asp Asp Cys Leu Ile Ala
   2110                2115                2120
tct gat ttt gaa att gat ttc caa caa ctg gtg cct gtc ttt tct agt     6855
Ser Asp Phe Glu Ile Asp Phe Gln Gln Leu Val Pro Val Phe Ser Ser
2125               2130                2135                2140
ttt gga cag gta ata act aca gct gac aag act gat ttt ttt aaa ctg     6903
Phe Gly Gln Val Ile Thr Thr Ala Asp Lys Thr Asp Phe Phe Lys Leu
               2145                2150                2155
aca acg ctt tcg gag gtg acc ttc ctt aag cgc gct ttt gtt ctg acg     6951
Thr Thr Leu Ser Glu Val Thr Phe Leu Lys Arg Ala Phe Val Leu Thr
           2160                2165                2170
gcc ttt tac aag cca gtg atg gat gtg aag acc ctt gaa gca atc tta     6999
Ala Phe Tyr Lys Pro Val Met Asp Val Lys Thr Leu Glu Ala Ile Leu
       2175                2180                2185
agc ttt gtt cgc cca ggc aca cag gct gaa aag ctc ctg tcc gtg gcg     7047
Ser Phe Val Arg Pro Gly Thr Gln Ala Glu Lys Leu Leu Ser Val Ala
   2190                2195                2200
cag ttg gca ggc cac tgc gaa ccg gag cag tat gag cgc ctg ttt gag     7095
Gln Leu Ala Gly His Cys Glu Pro Glu Gln Tyr Glu Arg Leu Phe Glu
2205               2210                2215                2220
ccc ttt gct ggg atg tat ttc gtc cct act tgg cga ctt gcg cct gca     7143
Pro Phe Ala Gly Met Tyr Phe Val Pro Thr Trp Arg Leu Ala Pro Ala
               2225                2230                2235
gtg gtt gat gaa gct tgg atg cta aat tct ttt tgactttgtt tttctttgtt   7196
Val Val Asp Glu Ala Trp Met Leu Asn Ser Phe
           2240                2245
ttcttttagg cttttaaggt gttaagttta aaggttaaga gtttttagaa gttaagatag   7256
agtttagttt ttagttttga gc                                            7278
 
           
             2 
             2247 
             PRT 
             equine rhinovirus 1 
           
            2
Met Ala Ala Ser Lys Val Tyr Arg Val Cys Glu Gln Thr Leu Leu Ala
  1               5                  10                  15
Gly Ala Val Arg Met Met Asp Lys Phe Leu Gln Lys Arg Thr Val Phe
             20                  25                  30
Val Pro His Leu Asp Lys Thr Ile Arg Leu Thr Gly Leu His Asn Tyr
         35                  40                  45
Asp Asn Thr Cys Trp Leu Asn Ala Leu Thr Gln Leu Thr Gln Ile Leu
     50                  55                  60
Gly Ile Arg Leu Phe Asp Glu His Phe Gly Asn Arg Gly Leu Phe Thr
 65                  70                  75                  80
Arg Lys Thr Ile Asp Trp Val Ser Asp Gln Thr Gly Ile Lys Asp Leu
                 85                  90                  95
Lys Ser Gly Ala Pro Pro Leu Val Val Val Tyr Lys Leu Trp Gln His
            100                 105                 110
Gly His Leu Asp Val Gly Thr Met Glu Lys Pro Arg Ser Ile Thr Leu
        115                 120                 125
Trp Ser Gly Pro Lys Val Cys Leu Ser Asp Phe Trp Ala Cys Val Ser
    130                 135                 140
Ala Lys Pro Gly His Ala Val Phe Tyr Leu Leu Thr Ser Glu Gly Trp
145                 150                 155                 160
Ile Cys Val Asp Asp Lys Lys Ile Tyr Pro Glu Thr Pro Lys Thr Glu
                165                 170                 175
Asp Val Leu Val Phe Ala Pro Tyr Asp Phe Glu Ser Leu Gly Lys Asp
            180                 185                 190
Pro Pro Lys Leu His Gln Arg Tyr Glu Lys Ala Phe Glu Leu Ser Gly
        195                 200                 205
Gly Gly Thr Ser Thr Pro Thr Thr Gly Asn Gln Asn Met Ser Gly Asn
    210                 215                 220
Ser Gly Ser Ile Val Gln Asn Phe Tyr Met Gln Gln Tyr Gln Asn Ser
225                 230                 235                 240
Ile Asp Ala Asp Leu Gly Asp Asn Val Ile Ser Pro Glu Gly Gln Gly
                245                 250                 255
Ser Asn Thr Ser Ser Ser Thr Ser Ser Ser Gln Ser Ser Gly Leu Gly
            260                 265                 270
Gly Trp Phe Ser Ser Leu Leu Asn Leu Gly Thr Lys Leu Leu Ala Asp
        275                 280                 285
Lys Lys Thr Glu Glu Thr Thr Asn Ile Glu Asp Arg Ile Glu Thr Thr
    290                 295                 300
Val Val Gly Val Thr Ile Ile Asn Ser Gln Gly Ser Val Gly Thr Thr
305                 310                 315                 320
Tyr Cys Tyr Ser Lys Pro Asp Gly Arg Pro Pro Ser Thr Val Ser Asp
                325                 330                 335
Pro Val Thr Arg Leu Gly Pro Thr Leu Ser Arg His Tyr Thr Phe Lys
            340                 345                 350
Val Gly Glu Trp Pro His Ser Gln Ser His Gly His Ala Trp Ile Cys
        355                 360                 365
Pro Leu Pro Gly Asp Lys Leu Lys Lys Met Gly Ser Phe His Glu Val
    370                 375                 380
Val Lys Ala His His Leu Val Lys Asn Gly Trp Asp Val Val Val Gln
385                 390                 395                 400
Val Asn Pro Ser Phe Ala His Ser Gly Pro Leu Cys Val Ala Ala Val
                405                 410                 415
Pro Glu Tyr Glu His Thr His Glu Lys Ala Leu Lys Trp Ser Glu Leu
            420                 425                 430
Glu Glu Pro Ala Tyr Thr Tyr Gln Gln Leu Ser Val Phe Pro His Gln
        435                 440                 445
Leu Leu Asn Leu Arg Thr Asn Ser Ser Val His Leu Val Met Pro Tyr
    450                 455                 460
Ile Gly Pro Gly Gln Pro Thr Asn Leu Thr Leu His Asn Pro Trp Thr
465                 470                 475                 480
Ile Val Ile Leu Ile Leu Ser Glu Leu Thr Gly Pro Gly Gln Thr Val
                485                 490                 495
Pro Val Thr Met Ser Val Ala Pro Ile Asp Ala Met Val Asn Gly Pro
            500                 505                 510
Leu Pro Asn Pro Glu Ala Pro Ile Arg Val Val Ser Val Pro Glu Ser
        515                 520                 525
Asp Ser Phe Met Ser Ser Val Pro Asp Asn Ser Thr Pro Leu Tyr Pro
    530                 535                 540
Lys Val Val Val Pro Pro Arg Gln Val Pro Gly Arg Phe Thr Asn Phe
545                 550                 555                 560
Ile Asp Val Ala Lys Gln Thr Tyr Ser Phe Cys Ser Ile Ser Gly Lys
                565                 570                 575
Pro Tyr Phe Glu Val Thr Asn Thr Ser Gly Asp Glu Pro Leu Phe Gln
            580                 585                 590
Met Asp Val Ser Leu Ser Ala Ala Glu Leu His Gly Thr Tyr Val Ala
        595                 600                 605
Ser Leu Ser Ser Phe Phe Ala Gln Tyr Arg Gly Ser Leu Asn Phe Asn
    610                 615                 620
Phe Ile Phe Thr Gly Ala Ala Ala Thr Lys Ala Lys Phe Leu Val Ala
625                 630                 635                 640
Phe Val Pro Pro His Ser Ala Ala Pro Lys Thr Arg Asp Glu Ala Met
                645                 650                 655
Ala Cys Ile His Ala Val Trp Asp Val Gly Leu Asn Ser Ala Phe Ser
            660                 665                 670
Phe Asn Val Pro Tyr Pro Ser Pro Ala Asp Phe Met Ala Val Tyr Ser
        675                 680                 685
Ala Glu Arg Thr Val Val Asn Val Ser Gly Trp Leu Gln Val Tyr Ala
    690                 695                 700
Leu Thr Ala Leu Thr Ser Thr Asp Ile Ala Val Asn Ser Lys Gly Arg
705                 710                 715                 720
Val Leu Val Ala Val Ser Ala Gly Pro Asp Phe Ser Leu Arg His Pro
                725                 730                 735
Ala Asp Leu Pro Asp Lys Gln Val Thr Asn Val Gly Glu Asp Gly Glu
            740                 745                 750
Pro Gly Glu Thr Glu Pro Arg His Ala Leu Ser Pro Val Asp Met His
        755                 760                 765
Val His Thr Asp Val Ser Phe Leu Leu Asp Arg Phe Phe Asp Val Glu
    770                 775                 780
Thr Leu Glu Leu Ser Asn Leu Thr Gly Ser Pro Ala Thr His Val Leu
785                 790                 795                 800
Asp Pro Phe Gly Ser Thr Ala Gln Leu Ala Trp Ala Arg Leu Leu Asn
                805                 810                 815
Thr Cys Thr Tyr Phe Phe Ser Asp Leu Glu Leu Ser Ile Gln Phe Lys
            820                 825                 830
Phe Thr Thr Thr Pro Ser Ser Val Gly Glu Gly Phe Val Trp Val Lys
        835                 840                 845
Trp Leu Pro Val Gly Ala Pro Thr Lys Thr Thr Asp Ala Trp Gln Leu
    850                 855                 860
Glu Gly Gly Gly Asn Ser Val Arg Ile Gln Lys Leu Ala Val Ala Gly
865                 870                 875                 880
Met Cys Pro Thr Val Val Phe Lys Ile Ala Gly Ser Arg Ser Gln Ala
                885                 890                 895
Cys Ala Ser Ala Leu Pro Tyr Thr Ser Met Trp Arg Val Val Pro Val
            900                 905                 910
Phe Tyr Asn Gly Trp Gly Ala Pro Thr Lys Glu Lys Ala Thr Tyr Asn
        915                 920                 925
Trp Leu Pro Gly Ala His Phe Gly Ser Ile Leu Leu Thr Ser Asp Ala
    930                 935                 940
His Asp Lys Gly Gly Cys Tyr Leu Arg Tyr Ala Phe Arg Ala Pro Ala
945                 950                 955                 960
Met Tyr Cys Pro Arg Pro Ile Pro Pro Ala Phe Thr Arg Pro Ala Asp
                965                 970                 975
Lys Thr Arg His Lys Phe Pro Thr Asn Ile Asn Lys Gln Cys Thr Asn
            980                 985                 990
Tyr Ser Leu Leu Lys Leu Ala Gly Asp Val Glu Ser Asn Pro Gly Pro
        995                1000                 1005
Thr Ile Phe Ser Lys Ala Ser Ala Asp Leu Asn Ala Leu Ser Thr Ser
   1010                1015                1020
Leu Gly Glu Leu Thr Gly Met Leu Lys Asp Leu Lys Ala Lys Ala Glu
1025                1030                1035               1040
Thr Tyr Ser Pro Phe Tyr Lys Met Ala Lys Met Leu Phe Lys Leu Ala
               1045                1050                1055
Thr Leu Ala Val Ala Ala Met Arg Thr Lys Asp Pro Val Val Val Val
           1060                1065                1070
Met Leu Ile Ala Asp Phe Gly Leu Glu Val Phe Asp Thr Gly Phe Phe
       1075                1080                 1085
Phe Ser Tyr Phe Gln Glu Lys Leu Gln Pro Tyr Met Lys Thr Ile Pro
   1090                1095                1100
Gly Lys Ile Ser Asp Leu Val Thr Asp Ala Ala Thr Ala Ala Ala Gln
1105                1110                1115               1120
Ile Pro Lys Gly Val Tyr Ser Phe Val Ser Ser Phe Phe Glu Thr Pro
               1125                1130                1135
Glu Gly Val Val Glu Lys Gln Val Ser Leu Arg Thr Val Asn Asp Ile
           1140                1145                1150
Phe Ala Leu Leu Lys Asn Ser Asp Trp Phe Ile Lys Thr Leu Val Ala
       1155                1160                 1165
Leu Lys Lys Trp Leu Thr Ser Trp Phe Ala Gln Glu Gln Gln Ala Asp
   1170                1175                1180
Asp Ala Leu Tyr Ser Glu Leu Glu Lys Tyr Pro Leu Tyr Lys Leu Lys
1185                1190                1195               1200
Leu Lys Glu Pro Asp Thr Gln Glu Glu Ala Arg Gln Trp Phe Lys Asp
               1205                1210                1215
Met Gln Gln Arg Ala Leu Ala Val Lys Asp Lys Gly Leu Phe Ser Leu
           1220                1225                1230
Leu Gln Ile Pro Leu Val Asn Leu Pro Gln Ser Arg Pro Glu Pro Val
       1235                1240                 1245
Val Cys Val Leu Arg Gly Ala Ser Gly Gln Gly Lys Ser Tyr Leu Ala
   1250                1255                1260
Asn Leu Met Ala Gln Ala Ile Ser Leu Leu Leu Val Gly Lys Gln Asp
1265                1270                1275               1280
Ser Val Trp Ser Cys Pro Pro Asp Pro Thr Tyr Phe Asp Gly Tyr Asn
               1285                1290                1295
Gly Gln Ala Val Val Ile Met Asp Ala Leu Gly Gln Asp Pro Asn Gly
           1300                1305                1310
Ala Asp Phe Lys Tyr Phe Cys Gln Met Val Ser Thr Thr Ala Phe Val
       1315                1320                 1325
Pro Pro Met Ala His Leu Asp Asp Lys Gly Ile Pro Phe Thr Ser Pro
   1330                1335                1340
Val Val Ile Cys Thr Thr Asn Leu His Ser Ser Phe Thr Pro Ile Thr
1345                1350                1355               1360
Val Ser Cys Pro Glu Ala Leu Lys Arg Arg Phe Arg Phe Asp Val Thr
               1365                1370                1375
Val Ser Ala Lys Pro Gly Phe Val Arg Thr Val Gly Ser Asn Gln Leu
           1380                1385                1390
Leu Asn Leu Pro Leu Ala Leu Lys Pro Ala Gly Leu Pro Pro His Pro
       1395                1400                 1405
Ile Phe Glu Asn Asp Met Pro Ile Ile Asn Gly Gln Ala Val Lys Leu
   1410                1415                1420
Ala Leu Ser Gly Gly Glu Val Thr Ala Phe Glu Leu Ile Glu Met Ile
1425                1430                1435               1440
Leu Ser Glu Val Gln Asn Arg Gln Asp Thr His Lys Met Pro Ile Phe
               1445                1450                1455
Lys Gln Ser Trp Ser Asp Leu Phe Arg Lys Cys Thr Thr Asp Glu Glu
           1460                1465                1470
Gln Lys Met Leu Gln Phe Leu Ile Asp Asn Lys Asp Ser Glu Ile Leu
       1475                1480                 1485
Arg Ala Phe Val Ser Glu Arg Ser Ile Leu Leu His Glu Glu Tyr Leu
   1490                1495                1500
Lys Trp Glu Ser Tyr Met Thr Arg Arg Ala Lys Phe His Arg Leu Ala
1505                1510                1515               1520
Ala Asp Phe Ala Met Phe Leu Ser Ile Leu Thr Ser Leu Ile Val Ile
               1525                1530                1535
Phe Cys Leu Val Tyr Ser Met Tyr Gln Leu Phe Lys Thr Pro Asp Glu
           1540                1545                1550
Gln Ser Ala Tyr Asp Pro Ser Thr Lys Pro Lys Pro Lys Thr Gln Glu
       1555                1560                 1565
Val Lys Thr Leu Lys Ile Arg Thr Glu Thr Gly Val Pro Ala Thr Asp
   1570                1575                1580
Leu Gln Gln Ser Ile Met Lys Asn Val Gln Pro Ile Glu Leu Tyr Leu
1585                1590                1595               1600
Asp Asn Glu Leu Val Thr Asp Cys Ser Ala Leu Gly Val Tyr Asp Asn
               1605                1610                1615
Ser Tyr Leu Val Pro Leu His Leu Phe Glu Phe Asp Phe Asp Thr Ile
           1620                1625                1630
Val Leu Gly Gly Arg His Tyr Lys Lys Ala Glu Cys Glu Lys Val Glu
       1635                1640                 1645
Phe Glu Leu Glu Val Asn Gly Asp Val Val Ser Ser Asp Ala Cys Leu
   1650                1655                1660
Leu Arg Val Ser Ser Gly Pro Lys Val Arg Asn Ile Val His Leu Phe
1665                1670                1675               1680
Thr Asn Glu Ile Glu Leu Lys Lys Met Thr Gln Val Thr Gly Ile Met
               1685                1690                1695
Asn Ser Pro His Gln Ala Arg Thr Val Phe Phe Gly Ser Phe Leu Thr
           1700                1705                1710
Val Arg Lys Ser Ile Leu Thr Ser Asp Gly Thr Val Met Pro Asn Val
       1715                1720                 1725
Leu Ser Tyr Ala Ala Gln Thr Ser Arg Gly Tyr Cys Gly Ala Ala Ile
   1730                1735                1740
Val Ala Gly Ser Pro Ala Arg Ile Ile Gly Ile His Ser Ala Gly Thr
1745                1750                1755               1760
Gly Ser Val Ala Phe Cys Ser Leu Val Ser Arg Asp Ala Leu Glu Gln
               1765                1770                1775
Leu Trp Pro Gln Lys Gln Gly Asn Val Ser Arg Leu Asp Asp Asp Val
           1780                1785                1790
Arg Val Ser Val Pro Arg Arg Ser Lys Leu Val Lys Ser Leu Ala Tyr
       1795                1800                 1805
Pro Ile Phe Lys Pro Asp Tyr Gly Pro Ala Pro Leu Ser Gln Phe Asp
   1810                1815                1820
Lys Arg Leu Ser Asp Gly Val Lys Leu Asp Glu Val Val Phe Ala Lys
1825                1830                1835               1840
His Thr Gly Asp Lys Glu Ile Ser Ala Gln Asp Gln Lys Trp Leu Leu
               1845                1850                1855
Arg Ala Ala His Val Tyr Ala Gln Lys Val Phe Ser Arg Ile Gly Phe
           1860                1865                1870
Asp Asn Gln Ala Leu Thr Glu Lys Glu Ala Ile Cys Gly Ile Pro Gly
       1875                1880                 1885
Leu Asp Lys Met Glu Gln Asp Thr Ala Pro Gly Leu Pro Tyr Ala Gln
   1890                1895                1900
Gln Asn Lys Arg Arg Lys Asp Ile Cys Asp Phe Glu Glu Gly Arg Leu
1905                1910                1915               1920
Lys Gly Ala Glu Leu Gln Lys Asp Arg Phe Met Ala Gly Asp Tyr Ser
               1925                1930                1935
Asn Leu Val Tyr Gln Ser Phe Leu Lys Asp Glu Ile Arg Pro Leu Glu
           1940                1945                1950
Lys Val Arg Ala Gly Lys Thr Arg Leu Ile Asp Val Pro Pro Met Pro
       1955                1960                 1965
His Val Val Val Gly Arg Gln Leu Leu Gly Arg Phe Val Ala Lys Phe
   1970                1975                1980
His Glu Ala Asn Gly Phe Asp Ile Gly Ser Ala Ile Gly Cys Asp Pro
1985                1990                1995               2000
Asp Val Asp Trp Thr Arg Phe Gly Leu Glu Leu Glu Arg Phe Arg Tyr
               2005                2010                2015
Val Tyr Ala Cys Asp Tyr Ser Arg Phe Asp Ala Asn His Ala Ala Asp
           2020                2025                2030
Ala Met Arg Val Val Leu Asn Tyr Phe Phe Ser Glu Asp His Gly Phe
       2035                2040                 2045
Asp Pro Gly Val Pro Ala Phe Ile Glu Ser Leu Val Asp Ser Val His
   2050                2055                2060
Ala Tyr Glu Glu Lys Arg Tyr Asn Ile Tyr Gly Gly Leu Pro Ser Gly
2065                2070                2075               2080
Cys Ser Cys Thr Ser Ile Leu Asn Thr Ile Leu Asn Asn Val Tyr Ile
               2085                2090                2095
Leu Ala Ala Met Met Lys Ala Tyr Glu Asn Phe Glu Pro Asp Asp Ile
           2100                2105                2110
Gln Val Ile Cys Tyr Gly Asp Asp Cys Leu Ile Ala Ser Asp Phe Glu
       2115                2120                 2125
Ile Asp Phe Gln Gln Leu Val Pro Val Phe Ser Ser Phe Gly Gln Val
   2130                2135                2140
Ile Thr Thr Ala Asp Lys Thr Asp Phe Phe Lys Leu Thr Thr Leu Ser
2145                2150                2155               2160
Glu Val Thr Phe Leu Lys Arg Ala Phe Val Leu Thr Ala Phe Tyr Lys
               2165                2170                2175
Pro Val Met Asp Val Lys Thr Leu Glu Ala Ile Leu Ser Phe Val Arg
           2180                2185                2190
Pro Gly Thr Gln Ala Glu Lys Leu Leu Ser Val Ala Gln Leu Ala Gly
       2195                2200                 2205
His Cys Glu Pro Glu Gln Tyr Glu Arg Leu Phe Glu Pro Phe Ala Gly
   2210                2215                2220
Met Tyr Phe Val Pro Thr Trp Arg Leu Ala Pro Ala Val Val Asp Glu
2225                2230                2235               2240
Ala Trp Met Leu Asn Ser Phe
               2245
 
           
             3 
             248 
             PRT 
             equine rhinovirus 1 
           
            3
Val Thr Asn Val Gly Glu Asp Gly Glu Pro Gly Glu Thr Glu Pro Arg
  1               5                  10                  15
His Ala Leu Ser Pro Val Asp Met His Val His Thr Asp Val Ser Phe
             20                  25                  30
Leu Leu Asp Arg Phe Phe Asp Val Glu Thr Leu Glu Leu Ser Asn Leu
         35                  40                  45
Thr Gly Ser Pro Ala Thr His Val Leu Asp Pro Phe Gly Ser Thr Ala
     50                  55                  60
Gln Leu Ala Trp Ala Arg Leu Leu Asn Thr Cys Thr Tyr Phe Phe Ser
 65                  70                  75                  80
Asp Leu Glu Leu Ser Ile Gln Phe Lys Phe Thr Thr Thr Pro Ser Ser
                 85                  90                  95
Val Gly Glu Gly Phe Val Trp Val Lys Trp Leu Pro Val Gly Ala Pro
            100                 105                 110
Thr Lys Thr Thr Asp Ala Trp Gln Leu Glu Gly Gly Gly Asn Ser Val
        115                 120                 125
Arg Ile Gln Lys Leu Ala Val Ala Gly Met Cys Pro Thr Val Val Phe
    130                 135                 140
Lys Ile Ala Gly Ser Arg Ser Gln Ala Cys Ala Ser Ala Leu Pro Tyr
145                 150                 155                 160
Thr Ser Met Trp Arg Val Val Pro Val Phe Tyr Asn Gly Trp Gly Ala
                165                 170                 175
Pro Thr Lys Glu Lys Ala Thr Tyr Asn Trp Leu Pro Gly Ala His Phe
            180                 185                 190
Gly Ser Ile Leu Leu Thr Ser Asp Ala His Asp Lys Gly Gly Cys Tyr
        195                 200                 205
Leu Arg Tyr Ala Phe Arg Ala Pro Ala Met Tyr Cys Pro Arg Pro Ile
    210                 215                 220
Pro Pro Ala Phe Thr Arg Pro Ala Asp Lys Thr Arg His Lys Phe Pro
225                 230                 235                 240
Thr Asn Ile Asn Lys Gln Cys Thr
                245
 
           
             4 
             230 
             PRT 
             equine rhinovirus 1 
           
            4
Asp Lys Lys Thr Glu Glu Thr Thr Asn Ile Glu Asp Arg Ile Glu Thr
  1               5                  10                  15
Thr Val Val Gly Val Thr Ile Ile Asn Ser Gln Gly Ser Val Gly Thr
             20                  25                  30
Thr Tyr Cys Tyr Ser Lys Pro Asp Gly Arg Pro Pro Ser Thr Val Ser
         35                  40                  45
Asp Pro Val Thr Arg Leu Gly Pro Thr Leu Ser Arg His Tyr Thr Phe
     50                  55                  60
Lys Val Gly Glu Trp Pro His Ser Gln Ser His Gly His Ala Trp Ile
 65                  70                  75                  80
Cys Pro Leu Pro Gly Asp Lys Leu Lys Lys Met Gly Ser Phe His Glu
                 85                  90                  95
Val Val Lys Ala His His Leu Val Lys Asn Gly Trp Asp Val Val Val
            100                 105                 110
Gln Val Asn Pro Ser Phe Ala His Ser Gly Pro Leu Cys Val Ala Ala
        115                 120                 125
Val Pro Glu Tyr Glu His Thr His Glu Lys Ala Leu Lys Trp Ser Glu
    130                 135                 140
Leu Glu Glu Pro Ala Tyr Thr Tyr Gln Gln Leu Ser Val Phe Pro His
145                 150                 155                 160
Gln Leu Leu Asn Leu Arg Thr Asn Ser Ser Val His Leu Val Met Pro
                165                 170                 175
Tyr Ile Gly Pro Gly Gln Pro Thr Asn Leu Thr Leu His Asn Pro Trp
            180                 185                 190
Thr Ile Val Ile Leu Ile Leu Ser Glu Leu Thr Gly Pro Gly Gln Thr
        195                 200                 205
Val Pro Val Thr Met Ser Val Ala Pro Ile Asp Ala Met Val Asn Gly
    210                 215                 220
Pro Leu Pro Asn Pro Glu
225                 230
 
           
             5 
             226 
             PRT 
             equine rhinovirus 1 
           
            5
Ala Pro Ile Arg Val Val Ser Val Pro Glu Ser Asp Ser Phe Met Ser
  1               5                  10                  15
Ser Val Pro Asp Asn Ser Thr Pro Leu Tyr Pro Lys Val Val Val Pro
             20                  25                  30
Pro Arg Gln Val Pro Gly Arg Phe Thr Asn Phe Ile Asp Val Ala Lys
         35                  40                  45
Gln Thr Tyr Ser Phe Cys Ser Ile Ser Gly Lys Pro Tyr Phe Glu Val
     50                  55                  60
Thr Asn Thr Ser Gly Asp Glu Pro Leu Phe Gln Met Asp Val Ser Leu
 65                  70                  75                  80
Ser Ala Ala Glu Leu His Gly Thr Tyr Val Ala Ser Leu Ser Ser Phe
                 85                  90                  95
Phe Ala Gln Tyr Arg Gly Ser Leu Asn Phe Asn Phe Ile Phe Thr Gly
            100                 105                 110
Ala Ala Ala Thr Lys Ala Lys Phe Leu Val Ala Phe Val Pro Pro His
        115                 120                 125
Ser Ala Ala Pro Lys Thr Arg Asp Glu Ala Met Ala Cys Ile His Ala
    130                 135                 140
Val Trp Asp Val Gly Leu Asn Ser Ala Phe Ser Phe Asn Val Pro Tyr
145                 150                 155                 160
Pro Ser Pro Ala Asp Phe Met Ala Val Tyr Ser Ala Glu Arg Thr Val
                165                 170                 175
Val Asn Val Ser Gly Trp Leu Gln Val Tyr Ala Leu Thr Ala Leu Thr
            180                 185                 190
Ser Thr Asp Ile Ala Val Asn Ser Lys Gly Arg Val Leu Val Ala Val
        195                 200                 205
Ser Ala Gly Pro Asp Phe Ser Leu Arg His Pro Ala Asp Leu Pro Asp
    210                 215                 220
Lys Gln
225
 
           
             6 
             80 
             PRT 
             equine rhinovirus 1 
           
            6
Gly Gly Gly Thr Ser Thr Pro Thr Thr Gly Asn Gln Asn Met Ser Gly
  1               5                  10                  15
Asn Ser Gly Ser Ile Val Gln Asn Phe Tyr Met Gln Gln Tyr Gln Asn
             20                  25                  30
Ser Ile Asp Ala Asp Leu Gly Asp Asn Val Ile Ser Pro Glu Gly Gln
         35                  40                  45
Gly Ser Asn Thr Ser Ser Ser Thr Ser Ser Ser Gln Ser Ser Gly Leu
     50                  55                  60
Gly Gly Trp Phe Ser Ser Leu Leu Asn Leu Gly Thr Lys Leu Leu Ala
 65                  70                  75                  80
 
           
             7 
             744 
             DNA 
             equine rhinovirus 1 
           
            7
gttaccaatg tgggagagga tggtgaaccc ggtgagacag agcctcgtca tgctttgtca     60
cccgtggaca tgcacgtgca cacagatgtc agtttcttgc ttgaccggtt ctttgatgtt    120
gagacacttg agctttcaaa tttgacaggt tctcctgcca cacatgttct ggatccgttt    180
ggctcgactg cccaactggc ttgggcacgt ctgctaaaca cttgcaccta cttcttttct    240
gatttggaat tgtcaatcca gtttaaattt accaccactc cgtcctctgt tggagagggc    300
tttgtgtggg tgaagtggct ccctgttgga gcaccaacca agaccacaga tgcttggcag    360
ttagaaggag gtggaaattc agttagaatt caaaaattgg ccgttgcagg gatgtgcccc    420
actgttgtgt tcaagattgc aggctcccgt tcacaagcct gtgcttcagc gttgccatat    480
acatcaatgt ggcgtgttgt gccagtcttt tacaatggct ggggtgcacc taccaaagaa    540
aaggcaacct acaattggct tcctggtgca cactttggtt ccatcttgct gacttctgat    600
gcgcatgata aaggagggtg ctacttgcgg tatgctttcc gcgcgccagc gatgtattgc    660
cctcgaccca ttccgccggc ttttacgcgt ccagcggaca aaaccagaca taaatttccc    720
actaacatca acaaacagtg tact                                           744
 
           
             8 
             690 
             DNA 
             equine rhinovirus 1 
           
            8
gacaagaaga cagaagagac tacaaacatt gaagacagaa ttgaaacaac agtggttgga     60
gtcactatta ttaattcaca aggatctgtt ggaacaacct actgttactc caaaccggat    120
ggtagaccac catccacagt gtcagaccca gttaccagac ttggacccac gctttccagg    180
cactacacat ttaaggtagg tgagtggccc cattctcaat cacatggtca cgcatggatc    240
tgtccgttgc caggtgacaa actcaagaag atgggcagtt ttcatgaggt tgtcaaagcc    300
caccacctgg tcaagaacgg ctgggatgtg gttgtgcagg tgaatccctc atttgctcac    360
tccgggccgc tgtgtgtagc agcagtgccg gagtacgaac acacacatga gaaagcactc    420
aagtggtctg agcttgagga accagcttac acataccaac aactttcagt ttttccccac    480
cagttgctaa atttgaggac aaattcatca gtgcatttgg tgatgcccta cattgggcca    540
ggccaaccaa caaatctgac tttgcacaac ccgtggacca ttgttatttt aattttgtct    600
gaattgacag gacctggcca aactgtgcct gtgaccatgt cggtggctcc catcgatgca    660
atggttaatg ggcctcttcc aaatccagag                                     690
 
           
             9 
             678 
             DNA 
             equine rhinovirus 1 
           
            9
gcaccgatta gagtggtgtc tgtgcctgaa tcagattctt ttatgtcttc agtacctgat     60
aattcgactc cactataccc caaggttgtg gtcccaccgc gccaagttcc tggccggttt    120
acaaatttca ttgatgtggc aaaacagaca tattcatttt gttccatttc tggaaaacct    180
tattttgagg ttaccaacac ctctggggac gagccactgt ttcagatgga tgtgtcgctc    240
agtgcggcag agctacatgg cacttacgta gctagtttgt catcattttt tgcacagtac    300
agaggctcac ttaatttcaa ctttattttc actggtgcag cagccactaa ggcaaagttt    360
ctggttgctt ttgtgcctcc ccacagtgca gcgcccaaaa cgcgcgatga agcaatggcg    420
tgcatccatg ccgtgtggga tgttggcttg aactcagctt tttcttttaa tgtaccttat    480
ccctcccctg ctgacttcat ggccgtttat tctgcggaac ggacggttgt gaatgtctct    540
ggatggcttc aagtttatgc actaacagct ctaacttcaa ctgacattgc cgtgaacagt    600
aaaggccgtg tgctggttgc tgtttccgcc ggcccagact tctcccttcg tcacccggcg    660
gacctgcccg acaagcag                                                  678
 
           
             10 
             240 
             DNA 
             equine rhinovirus 1 
           
            10
ggcggaggta catccactcc aacaactggc aaccaaaaca tgtccggaaa cagtggttca     60
attgttcaaa atttttacat gcaacagtac cagaattcaa ttgacgcaga cctgggagac    120
aatgtgatta gccctgaagg ccagggcagc aacactagta gttcaacctc atcaagccaa    180
tcctctggct tgggcgggtg gttctctagt ttgctgaacc ttggaacaaa actactggct    240
 
           
             11 
             21 
             DNA 
             equine rhinovirus 1 
           
            11
gttgtgttca agattgcagg c                                               21
 
           
             12 
             20 
             DNA 
             equine rhinovirus 1 
           
            12
ttgctctcaa catctccagc                                                 20
 
           
             13 
             22 
             DNA 
             equine rhinovirus 1 
           
            13
tagcaccctc ctttatcatg cg                                              22
 
           
             14 
             33 
             DNA 
             equine rhinovirus 1 
           
            14
gctggatcca tgagtggcgg aggtacatcc act                                  33
 
           
             15 
             30 
             DNA 
             equine rhinovirus 1 
           
            15
gctctgcagc aggtctgctg atgctttgga                                      30
 
           
             16 
             32 
             DNA 
             equine rhinovirus 1 
           
            16
gctctgcaga tgattaggac tgagactggt gt                                   32
 
           
             17 
             30 
             DNA 
             equine rhinovirus 1 
           
            17
gctggatcct tagccatagt caggtttgaa                                      30
 
           
             18 
             25 
             DNA 
             equine rhinovirus 1 
           
            18
atccagcaag ccgctgtccg gttac                                           25
 
           
             19 
             18 
             DNA 
             equine rhinovirus 1 
           
            19
cgaagagaca cctgcttc                                                   18
 
           
             20 
             21 
             DNA 
             equine rhinovirus 1 
           
            20
ttctggtgga gaagtgacag c                                               21
 
           
             21 
             19 
             DNA 
             equine rhinovirus 1 
           
            21
gtgagccagc aacaattgc                                                  19
 
           
             22 
             865 
             DNA 
             equine rhinovirus 1 
             
               CDS 
               (794)..(865) 
             
           
            22
taagtaaaac gctgtaactg catgatttgc gcctgtagcg ccagtaaaac gcagaaacca     60
caagcaaaaa cctgtagcgt cagtaaaacg cgcacattca catacagagc ttcccggctt    120
taagggttac tgctcgtaat gagagcacat gacaacttgt cgagattacg gcaactgtca    180
cgggagagag gagcccgttt tcgggcactt gtctcctaaa caatgttggc gcgcatttgc    240
gcgccccccc cctttttcag ccccctgtca ttgactggtc gaagcgttcg caataagact    300
ggtcgtcact tggctgttct atcgtttcag gctttagcgc gcccttgcgc ggcgggccgt    360
caagcccgtg cgctgtatag cgccaggtaa ccggacagcg gcgtgctgga ttttcccggt    420
gccattgctc tggatggtgt caccaagctg acaaatgcgg agtgaacctc acaaagcgac    480
acgcctgtgg tagcgctgcc caaaagggag cggaactccc cgccgaggcg gtcctctctg    540
gccaaaagcc cagcgttgat agcgcctttt gggatgcagg aaccccacct gccaggtgtg    600
aagtggagtg agcggatctc caatttggtc tgttctgaac tacaccattt actgctgtga    660
agaatgccct ggaggcaagc tggttacagc cctgaccagg ccctgcccgt gactctcgac    720
cggcgcaggg tcaaaaattg tctaagcagc agcaggaacg cgggagcgtt tcttttcctt    780
ttgtactgac atg atg gcg gcg tct aag gtg tat aga gtt tgc gag cag       829
               Met Ala Ala Ser Lys Val Tyr Arg Val Cys Glu Gln
                 1               5                  10
act ctg ctg gca ggt gcc gtt cgc atg atg gac aaa                      865
Thr Leu Leu Ala Gly Ala Val Arg Met Met Asp Lys
         15                  20
 
           
             23 
             24 
             PRT 
             equine rhinovirus 1 
           
            23
Met Ala Ala Ser Lys Val Tyr Arg Val Cys Glu Gln Thr Leu Leu Ala
  1               5                  10                  15
Gly Ala Val Arg Met Met Asp Lys
             20
 
           
             24 
             2318 
             PRT 
             Foot-and-mouth disease virus 
           
            24
Met Asn Thr Thr Asp Cys Phe Ile Ala Leu Val Gln Ala Ile Arg Glu
  1               5                  10                  15
Ile Lys Ala Leu Phe Leu Ser Arg Thr Thr Gly Lys Met Glu Leu Thr
             20                  25                  30
Leu Tyr Asn Gly Glu Lys Lys Thr Phe Tyr Ser Arg Pro Asn Asn His
         35                  40                  45
Asp Asn Cys Trp Leu Asn Ala Ile Leu Gln Leu Phe Arg Tyr Val Glu
     50                  55                  60
Glu Pro Phe Phe Asp Trp Val Tyr Ser Ser Pro Glu Asn Leu Thr Leu
 65                  70                  75                  80
Glu Ala Ile Lys Gln Leu Glu Asp Leu Thr Gly Leu Glu Leu His Glu
                 85                  90                  95
Gly Gly Pro Pro Ala Leu Val Ile Trp Asn Ile Lys His Leu Leu His
            100                 105                 110
Thr Gly Ile Gly Thr Ala Ser Arg Pro Ser Glu Val Cys Met Val Asp
        115                 120                 125
Gly Thr Asp Met Cys Leu Ala Asp Phe His Ala Gly Ile Phe Leu Lys
    130                 135                 140
Gly Gln Glu His Ala Val Phe Ala Cys Val Thr Ser Asn Gln Trp Tyr
145                 150                 155                 160
Ala Ile Asp Asp Glu Asp Phe Tyr Pro Trp Thr Pro Asp Pro Ser Asp
                165                 170                 175
Val Leu Val Phe Val Pro Tyr Asp Gln Glu Pro Leu Asn Gly Glu Trp
            180                 185                 190
Lys Ala Lys Val Gln Arg Lys Leu Lys Gly Ala Gly Gln Ser Ser Pro
        195                 200                 205
Ala Thr Gly Ser Gln Asn Gln Ser Gly Asn Thr Gly Ser Ile Ile Asn
    210                 215                 220
Asn Tyr Tyr Met Gln Gln Tyr Gln Asn Ser Met Asp Thr Gln Leu Gly
225                 230                 235                 240
Asp Asn Ala Ile Ser Gly Gly Ser Asn Glu Gly Ser Thr Asp Thr Thr
                245                 250                 255
Ser Thr His Thr Thr Asn Thr Gln Asn Asn Asp Trp Phe Ser Lys Leu
            260                 265                 270
Ala Ser Ser Ala Phe Ser Gly Leu Phe Gly Ala Leu Leu Ala Asp Lys
        275                 280                 285
Lys Thr Glu Glu Thr Thr Leu Leu Glu Asp Arg Ile Leu Thr Thr Arg
    290                 295                 300
Asn Gly His Thr Thr Ser Thr Thr Gln Ser Ser Val Gly Val Thr Tyr
305                 310                 315                 320
Gly Tyr Ala Thr Ala Glu Asp Phe Val Ser Gly Pro Asn Thr Ser Gly
                325                 330                 335
Leu Glu Thr Arg Val Val Gln Ala Glu Arg Phe Phe Lys Thr His Leu
            340                 345                 350
Phe Asp Trp Val Thr Ser Asp Ser Phe Gly Arg Cys His Leu Leu Glu
        355                 360                 365
Leu Pro Thr Asp His Lys Gly Val Tyr Gly Ser Leu Thr Asp Ser Tyr
    370                 375                 380
Ala Tyr Met Arg Asn Gly Trp Asp Val Glu Val Thr Ala Val Gly Asn
385                 390                 395                 400
Gln Phe Asn Gly Gly Cys Leu Leu Val Ala Met Val Pro Glu Leu Tyr
                405                 410                 415
Ser Ile Gln Lys Arg Glu Leu Tyr Gln Leu Thr Leu Phe Pro His Gln
            420                 425                 430
Phe Ile Asn Pro Arg Thr Asn Met Thr Ala His Ile Thr Val Pro Phe
        435                 440                 445
Val Gly Val Asn Arg Tyr Asp Gln Tyr Lys Val His Lys Pro Trp Thr
    450                 455                 460
Leu Val Val Met Val Val Ala Pro Leu Thr Val Asn Thr Glu Gly Ala
465                 470                 475                 480
Pro Gln Ile Lys Val Tyr Ala Asn Ile Ala Pro Thr Asn Val His Val
                485                 490                 495
Ala Gly Glu Phe Pro Ser Lys Glu Gly Ile Phe Pro Val Ala Cys Ser
            500                 505                 510
Asp Gly Tyr Gly Gly Leu Val Thr Thr Asp Pro Lys Thr Ala Asp Pro
        515                 520                 525
Val Tyr Gly Lys Val Phe Asn Pro Pro Arg Asn Gln Leu Pro Gly Arg
    530                 535                 540
Phe Thr Asn Leu Leu Asp Val Ala Glu Ala Cys Pro Thr Phe Leu Arg
545                 550                 555                 560
Phe Glu Gly Gly Val Pro Tyr Val Thr Thr Lys Thr Asp Ser Asp Arg
                565                 570                 575
Val Leu Ala Gln Phe Asp Met Ser Leu Ala Ala Lys Gln Met Ser Asn
            580                 585                 590
Thr Phe Leu Ala Gly Leu Ala Gln Tyr Tyr Thr Gln Tyr Ser Gly Thr
        595                 600                 605
Ile Asn Leu His Phe Met Phe Thr Gly Pro Thr Asp Ala Lys Ala Arg
    610                 615                 620
Tyr Met Val Ala Tyr Ala Pro Pro Gly Met Glu Pro Pro Lys Thr Pro
625                 630                 635                 640
Glu Ala Ala Ala His Cys Ile His Ala Glu Trp Asp Thr Gly Leu Asn
                645                 650                 655
Ser Lys Phe Thr Phe Ser Ile Pro Tyr Leu Ser Ala Ala Asp Tyr Ala
            660                 665                 670
Tyr Thr Ala Ser Gly Val Ala Glu Thr Thr Asn Val Gln Gly Trp Val
        675                 680                 685
Cys Leu Phe Gln Ile Thr His Gly Lys Ala Asp Gly Asp Ala Leu Val
    690                 695                 700
Val Leu Ala Ser Ala Gly Lys Asp Phe Glu Leu Arg Leu Pro Val Asp
705                 710                 715                 720
Ala Arg Ala Glu Thr Thr Ser Ala Gly Glu Ser Ala Asp Pro Val Thr
                725                 730                 735
Thr Thr Val Glu Asn Tyr Gly Gly Glu Thr Gln Ile Gln Arg Arg Gln
            740                 745                 750
His Thr Asp Val Ser Phe Ile Met Asp Arg Phe Val Lys Val Thr Pro
        755                 760                 765
Gln Asn Gln Ile Asn Ile Leu Asp Leu Met Gln Ile Pro Ser His Thr
    770                 775                 780
Leu Val Gly Ala Leu Leu Arg Ala Ser Thr Tyr Tyr Phe Ser Asp Leu
785                 790                 795                 800
Glu Ile Ala Val Lys His Glu Gly Asp Leu Thr Trp Val Pro Asn Gly
                805                 810                 815
Ala Pro Glu Lys Ala Leu Asp Asn Thr Thr Asn Pro Thr Ala Tyr His
            820                 825                 830
Lys Ala Pro Leu Thr Arg Leu Ala Leu Pro Tyr Thr Ala Pro His Arg
        835                 840                 845
Val Leu Ala Thr Val Tyr Asn Gly Glu Cys Arg Tyr Asn Arg Asn Ala
    850                 855                 860
Val Pro Asn Leu Arg Gly Asp Leu Gln Val Leu Ala Gln Lys Val Ala
865                 870                 875                 880
Arg Thr Leu Pro Thr Ser Phe Asn Tyr Gly Ala Ile Lys Ala Thr Arg
                885                 890                 895
Val Thr Glu Leu Leu Tyr Arg Met Lys Arg Ala Glu Thr Tyr Cys Pro
            900                 905                 910
Arg Pro Leu Leu Ala Ile His Pro Thr Glu Ala Arg His Lys Gln Lys
        915                 920                 925
Ile Val Ala Pro Val Lys Gln Thr Leu Asn Phe Asp Leu Leu Lys Leu
    930                 935                 940
Ala Gly Asp Val Glu Ser Asn Pro Gly Pro Phe Phe Phe Ser Asp Val
945                 950                 955                 960
Arg Ser Asn Phe Ser Lys Leu Val Glu Thr Ile Asn Gln Met Gln Glu
                965                 970                 975
Asp Met Ser Thr Lys His Gly Pro Asp Phe Asn Arg Leu Val Ser Ala
            980                 985                 990
Phe Glu Glu Leu Ala Ile Gly Val Lys Ala Ile Arg Thr Gly Leu Asp
        995                1000                1005
Glu Ala Lys Pro Trp Tyr Lys Leu Ile Lys Leu Leu Ser Arg Leu Ser
   1010                1015                1020
Cys Met Ala Ala Val Ala Ala Arg Ser Lys Asp Pro Val Leu Val Ala
1025               1030                1035                1040
Ile Met Leu Ala Asp Thr Gly Leu Glu Ile Leu Asp Ser Thr Phe Val
               1045                1050                1055
Val Lys Lys Ile Ser Asp Ser Leu Ser Ser Leu Phe His Val Pro Ala
           1060                1065                1070
Pro Val Phe Ser Phe Gly Ala Pro Val Leu Leu Ala Gly Leu Val Lys
       1075                1080                1085
Val Ala Ser Ser Phe Phe Arg Ser Thr Pro Glu Asp Leu Glu Arg Ala
   1090                1095                1100
Glu Lys Gln Leu Lys Ala Arg Asp Ile Asn Asp Ile Phe Ala Ile Leu
1105               1110                1115                1120
Lys Asn Gly Glu Trp Leu Val Lys Leu Ile Leu Ala Ile Arg Asp Trp
               1125                1130                1135
Ile Lys Ala Trp Ile Ala Ser Glu Glu Lys Phe Val Thr Met Thr Asp
           1140                1145                1150
Leu Val Pro Gly Ile Leu Glu Lys Gln Arg Asp Leu Asn Asp Pro Ser
       1155                1160                1165
Lys Tyr Lys Glu Ala Lys Glu Trp Leu Asp Asn Ala Arg Gln Ala Cys
   1170                1175                1180
Leu Lys Ser Gly Asn Val His Ile Ala Asn Leu Cys Lys Val Val Ala
1185               1190                1195                1200
Pro Ala Pro Ser Lys Ser Arg Pro Glu Pro Val Val Val Cys Leu Arg
               1205                1210                1215
Gly Lys Ser Gly Gln Gly Lys Ser Phe Leu Ala Asn Val Leu Ala Gln
           1220                1225                1230
Ala Ile Ser Thr His Phe Thr Gly Arg Ile Asp Ser Val Trp Tyr Cys
       1235                1240                1245
Pro Pro Asp Pro Asp His Phe Asp Gly Tyr Asn Gln Gln Thr Val Val
   1250                1255                1260
Val Met Asp Asp Leu Gly Gln Asn Pro Asp Gly Lys Asp Phe Lys Tyr
1265               1270                1275                1280
Phe Ala Gln Met Val Ser Thr Thr Gly Phe Ile Pro Pro Met Ala Ser
               1285                1290                1295
Leu Glu Asp Lys Gly Lys Pro Phe Asn Ser Lys Val Ile Ile Ala Thr
           1300                1305                1310
Thr Asn Leu Tyr Ser Gly Phe Thr Pro Arg Thr Met Val Cys Pro Asp
       1315                1320                1325
Ala Leu Asn Arg Arg Phe His Phe Asp Ile Asp Val Ser Ala Lys Asp
   1330                1335                1340
Gly Tyr Lys Ile Asn Ser Lys Leu Asp Ile Ile Lys Ala Leu Glu Asp
1345               1350                1355                1360
Thr His Ala Asn Pro Val Ala Met Phe Gln Tyr Asp Cys Ala Leu Leu
               1365                1370                1375
Asn Gly Met Ala Val Glu Met Lys Arg Met Gln Gln Asp Met Phe Lys
           1380                1385                1390
Pro Gln Pro Pro Leu Gln Asn Val Tyr Gln Leu Val Gln Glu Val Ile
       1395                1400                1405
Asp Arg Val Glu Leu His Glu Lys Val Ser Ser His Pro Ile Phe Lys
   1410                1415                1420
Gln Ile Ser Ile Pro Ser Gln Lys Ser Val Leu Tyr Phe Leu Ile Glu
1425               1430                1435                1440
Lys Gly Gln His Glu Ala Ala Ile Glu Phe Phe Glu Gly Met Val His
               1445                1450                1455
Asp Ser Ile Lys Glu Glu Leu Arg Pro Leu Ile Gln Gln Thr Ser Phe
           1460                1465                1470
Val Lys Arg Ala Phe Lys Arg Leu Lys Glu Asn Phe Glu Ile Val Ala
       1475                1480                1485
Leu Cys Leu Thr Leu Leu Ala Asn Ile Val Ile Met Ile Arg Glu Thr
   1490                1495                1500
Arg Lys Arg Gln Lys Met Val Asp Asp Ala Val Asn Glu Tyr Ile Glu
1505               1510                1515                1520
Lys Ala Asn Ile Thr Thr Asp Asp Lys Thr Leu Asp Glu Ala Glu Lys
               1525                1530                1535
Ser Pro Leu Glu Thr Ser Gly Ala Ser Thr Val Gly Phe Arg Glu Arg
           1540                1545                1550
Thr Leu Pro Gly Gln Lys Ala Cys Asp Asp Val Asn Ser Glu Pro Ala
       1555                1560                1565
Gln Pro Val Glu Glu Gln Pro Gln Ala Glu Gly Pro Tyr Ala Gly Pro
   1570                1575                1580
Leu Glu Arg Gln Lys Pro Leu Lys Val Arg Ala Lys Leu Pro Gln Gln
1585               1590                1595                1600
Glu Gly Pro Tyr Ala Gly Pro Met Glu Arg Gln Lys Pro Leu Lys Val
               1605                1610                1615
Lys Ala Lys Ala Pro Val Val Lys Glu Gly Pro Tyr Glu Gly Pro Val
           1620                1625                1630
Lys Lys Pro Val Ala Leu Lys Val Lys Ala Lys Asn Leu Ile Val Thr
       1635                1640                1645
Glu Ser Gly Ala Pro Pro Thr Asp Leu Gln Lys Met Val Met Gly Asn
   1650                1655                1660
Thr Lys Pro Val Glu Leu Ile Leu Asp Gly Lys Thr Val Ala Ile Cys
1665               1670                1675                1680
Cys Ala Thr Gly Val Phe Gly Thr Ala Tyr Leu Val Pro Arg His Leu
               1685                1690                1695
Phe Ala Glu Lys Tyr Asp Lys Ile Met Val Asp Gly Arg Ala Met Thr
           1700                1705                1710
Asp Ser Asp Tyr Arg Val Phe Glu Phe Glu Ile Lys Val Lys Gly Gln
       1715                1720                1725
Asp Met Leu Ser Asp Ala Ala Leu Met Val Leu His Arg Gly Asn Arg
   1730                1735                1740
Val Arg Asp Ile Thr Lys His Phe Arg Asp Thr Ala Arg Met Lys Lys
1745               1750                1755                1760
Gly Thr Pro Val Val Gly Val Ile Asn Asn Ala Asp Val Gly Arg Leu
               1765                1770                1775
Ile Phe Ser Gly Glu Ala Leu Thr Tyr Lys Asp Ile Val Val Cys Met
           1780                1785                1790
Asp Gly Asp Thr Met Pro Gly Leu Phe Ala Tyr Arg Ala Ala Thr Lys
       1795                1800                1805
Ala Gly Tyr Cys Gly Gly Ala Val Leu Ala Lys Asp Gly Ala Asp Thr
   1810                1815                1820
Phe Ile Val Gly Thr His Ser Ala Gly Gly Asn Gly Val Gly Tyr Cys
1825               1830                1835                1840
Ser Cys Val Ser Arg Ser Met Leu Leu Lys Met Lys Ala His Ile Asp
               1845                1850                1855
Pro Glu Pro His His Glu Gly Leu Ile Val Asp Thr Arg Asp Val Glu
           1860                1865                1870
Glu Arg Val His Val Met Arg Lys Thr Lys Leu Ala Pro Thr Val Ala
       1875                1880                1885
His Gly Val Phe Asn Pro Glu Phe Gly Pro Ala Ala Leu Ser Asn Lys
   1890                1895                1900
Asp Pro Arg Leu Asn Glu Gly Val Val Leu Asp Glu Val Ile Phe Ser
1905               1910                1915                1920
Lys His Lys Gly Asp Thr Lys Met Ser Glu Glu Asp Lys Ala Leu Phe
               1925                1930                1935
Arg Arg Cys Ala Ala Asp Tyr Ala Ser Arg Leu His Ser Val Leu Gly
           1940                1945                1950
Thr Ala Asn Ala Pro Leu Ser Ile Tyr Glu Ala Ile Lys Gly Val Asp
       1955                1960                1965
Gly Leu Asp Ala Met Glu Pro Asp Thr Ala Pro Gly Leu Pro Trp Ala
   1970                1975                1980
Leu Gln Gly Lys Arg Arg Gly Ala Leu Ile Asp Phe Glu Asn Gly Thr
1985               1990                1995                2000
Val Gly Pro Glu Val Glu Ala Ala Leu Lys Leu Met Glu Lys Arg Glu
               2005                2010                2015
Tyr Lys Phe Val Cys Gln Thr Phe Leu Lys Asp Glu Ile Arg Pro Leu
           2020                2025                2030
Glu Lys Val Arg Ala Gly Lys Thr Arg Ile Val Asp Val Leu Pro Val
       2035                2040                2045
Glu His Ile Leu Tyr Thr Arg Met Met Ile Gly Arg Phe Cys Ala Gln
   2050                2055                2060
Met His Ser Asn Asn Gly Pro Gln Ile Gly Ser Ala Val Gly Cys Asn
2065               2070                2075                2080
Pro Asp Val Asp Trp Gln Arg Phe Gly Thr His Phe Ala Gln Tyr Arg
               2085                2090                2095
Asn Val Trp Asp Val Asp Tyr Ser Ala Phe Asp Ala Asn His Cys Ser
           2100                2105                2110
Asp Ala Met Asn Ile Met Phe Glu Glu Val Phe Arg Thr Glu Phe Gly
       2115                2120                2125
Phe His Pro Asn Ala Glu Trp Ile Leu Lys Thr Leu Val Asn Thr Glu
   2130                2135                2140
His Ala Tyr Glu Asn Lys Arg Ile Thr Val Gly Gly Gly Met Pro Ser
2145               2150                2155                2160
Gly Cys Ser Ala Thr Ser Ile Ile Asn Thr Ile Leu Asn Asn Ile Tyr
               2165                2170                2175
Val Leu Tyr Ala Leu Arg Arg His Tyr Glu Gly Val Glu Leu Asp Thr
           2180                2185                2190
Tyr Thr Met Ile Ser Tyr Gly Asp Asp Ile Val Val Ala Ser Asp Tyr
       2195                2200                2205
Asp Leu Asp Phe Glu Ala Leu Lys Pro His Phe Lys Ser Leu Gly Gln
   2210                2215                2220
Thr Ile Thr Pro Ala Asp Lys Ser Asp Lys Gly Phe Val Leu Gly His
2225               2230                2235                2240
Ser Ile Thr Asp Val Thr Phe Leu Lys Arg His Phe His Met Asp Tyr
               2245                2250                2255
Gly Thr Gly Phe Tyr Lys Pro Val Met Ala Ser Lys Thr Leu Glu Ala
           2260                2265                2270
Ile Leu Ser Phe Ala Arg Arg Gly Thr Ile Gln Glu Lys Leu Ile Ser
       2275                2280                2285
Val Ala Gly Leu Ala Val His Ser Gly Pro Asp Glu Tyr Arg Arg Leu
   2290                2295                2300
Phe Glu Pro Phe Gln Gly Leu Phe Glu Ile Pro Ser Tyr Arg
2305               2310                2315
 
           
             25 
             2232 
             PRT 
             equine rhinovirus 1 
           
            25
Met Ala Ala Ser Lys Val Tyr Arg Val Cys Glu Gln Thr Leu Leu Ala
  1               5                  10                  15
Gly Ala Val Arg Met Met Asp Lys Phe Leu Gln Lys Arg Thr Val Phe
             20                  25                  30
Val Pro His Leu Asp Lys Thr Ile Arg Leu Thr Gly Leu His Asn Tyr
         35                  40                  45
Asp Asn Thr Cys Trp Leu Asn Ala Leu Thr Gln Leu Thr Gln Ile Leu
     50                  55                  60
Gly Ile Arg Leu Phe Asp Glu His Phe Gly Asn Arg Gly Leu Phe Thr
 65                  70                  75                  80
Arg Lys Thr Ile Asp Trp Val Ser Asp Gln Thr Gly Ile Lys Asp Leu
                 85                  90                  95
Lys Ser Gly Ala Pro Pro Leu Val Val Val Tyr Lys Leu Trp Gln His
            100                 105                 110
Gly His Leu Asp Val Gly Thr Met Glu Lys Pro Arg Ser Ile Thr Leu
        115                 120                 125
Trp Ser Gly Pro Lys Val Cys Leu Ser Asp Phe Trp Ala Cys Val Ser
    130                 135                 140
Ala Lys Pro Gly His Ala Val Phe Tyr Leu Leu Thr Ser Glu Gly Trp
145                 150                 155                 160
Ile Cys Val Asp Asp Lys Lys Ile Tyr Pro Glu Thr Pro Lys Thr Glu
                165                 170                 175
Asp Val Leu Val Phe Ala Pro Tyr Asp Phe Glu Ser Leu Gly Lys Asp
            180                 185                 190
Pro Pro Lys Leu His Gln Arg Tyr Glu Lys Ala Phe Glu Leu Ser Gly
        195                 200                 205
Gly Gly Thr Ser Thr Pro Thr Thr Gly Asn Gln Asn Met Ser Gly Asn
    210                 215                 220
Ser Gly Ser Ile Val Gln Asn Phe Tyr Met Gln Gln Tyr Gln Asn Ser
225                 230                 235                 240
Ile Asp Ala Asp Leu Gly Asp Asn Val Ile Ser Pro Glu Gly Gln Gly
                245                 250                 255
Ser Asn Thr Ser Ser Ser Thr Ser Ser Ser Gln Ser Ser Gly Leu Gly
            260                 265                 270
Gly Trp Phe Ser Ser Leu Leu Asn Leu Gly Thr Lys Leu Leu Ala Asp
        275                 280                 285
Lys Lys Thr Glu Glu Thr Thr Asn Ile Glu Asp Arg Ile Glu Thr Thr
    290                 295                 300
Val Val Gly Val Thr Ile Ile Asn Ser Gln Gly Ser Val Gly Thr Thr
305                 310                 315                 320
Tyr Cys Tyr Ser Lys Pro Asp Gly Arg Pro Pro Ser Thr Val Ser Asp
                325                 330                 335
Pro Val Thr Arg Leu Gly Pro Thr Leu Ser Arg His Tyr Thr Phe Lys
            340                 345                 350
Val Gly Glu Trp Pro His Ser Gln Ser His Gly His Ala Trp Ile Cys
        355                 360                 365
Pro Leu Pro Gly Asp Lys Leu Lys Lys Met Gly Ser Phe His Glu Val
    370                 375                 380
Val Lys Ala His His Leu Val Lys Asn Gly Trp Asp Val Val Val Gln
385                 390                 395                 400
Val Asn Pro Ser Phe Ala His Ser Gly Pro Leu Cys Val Ala Ala Val
                405                 410                 415
Pro Glu Tyr Glu His Thr His Glu Lys Ala Leu Lys Trp Ser Glu Leu
            420                 425                 430
Glu Glu Pro Ala Tyr Thr Tyr Gln Gln Leu Ser Val Phe Pro His Gln
        435                 440                 445
Leu Leu Asn Leu Arg Thr Asn Ser Ser Val His Leu Val Met Pro Tyr
    450                 455                 460
Ile Gly Pro Gly Gln Pro Thr Asn Leu Thr Leu His Asn Pro Trp Thr
465                 470                 475                 480
Ile Val Ile Leu Ile Leu Ser Glu Leu Thr Gly Pro Gly Gln Thr Val
                485                 490                 495
Pro Val Thr Met Ser Val Ala Pro Ile Asp Ala Met Val Asn Gly Pro
            500                 505                 510
Leu Pro Asn Pro Glu Ala Pro Ile Arg Val Val Ser Val Pro Glu Ser
        515                 520                 525
Asp Ser Phe Met Ser Ser Val Pro Asp Asn Ser Thr Pro Leu Tyr Pro
    530                 535                 540
Lys Val Val Val Pro Pro Arg Gln Val Pro Gly Arg Phe Thr Asn Phe
545                 550                 555                 560
Ile Asp Val Ala Lys Gln Thr Tyr Ser Phe Cys Ser Ile Ser Gly Lys
                565                 570                 575
Pro Tyr Phe Glu Val Thr Asn Thr Ser Gly Asp Glu Pro Leu Phe Gln
            580                 585                 590
Met Asp Val Ser Leu Ser Ala Ala Glu Leu His Gly Thr Tyr Val Ala
        595                 600                 605
Ser Leu Ser Ser Phe Phe Ala Gln Tyr Arg Gly Ser Leu Asn Phe Asn
    610                 615                 620
Phe Ile Phe Thr Gly Ala Ala Ala Thr Lys Ala Lys Phe Leu Val Ala
625                 630                 635                 640
Phe Val Pro Pro His Ser Ala Ala Pro Lys Thr Arg Asp Glu Ala Met
                645                 650                 655
Ala Cys Ile His Ala Val Trp Asp Val Gly Leu Asn Ser Ala Phe Ser
            660                 665                 670
Phe Asn Val Pro Tyr Pro Ser Pro Ala Asp Phe Met Ala Val Tyr Ser
        675                 680                 685
Ala Glu Arg Thr Val Val Asn Val Ser Gly Trp Leu Gln Val Tyr Ala
    690                 695                 700
Leu Thr Ala Leu Thr Ser Thr Asp Ile Ala Val Asn Ser Lys Gly Arg
705                 710                 715                 720
Val Leu Val Ala Val Ser Ala Gly Pro Asp Phe Ser Leu Arg His Pro
                725                 730                 735
Ala Asp Leu Pro Asp Lys Gln Val Thr Asn Val Gly Glu Asp Gly Glu
            740                 745                 750
Pro Gly Glu Thr Glu Pro Arg His Ala Leu Ser Pro Val Asp Met His
        755                 760                 765
Val His Thr Asp Val Ser Phe Leu Leu Asp Arg Phe Phe Asp Val Glu
    770                 775                 780
Thr Leu Glu Leu Ser Asn Leu Thr Gly Ser Pro Ala Thr His Val Leu
785                 790                 795                 800
Asp Pro Phe Gly Ser Thr Ala Gln Leu Ala Trp Ala Arg Leu Leu Asn
                805                 810                 815
Thr Cys Thr Tyr Phe Phe Ser Asp Leu Glu Leu Ser Ile Gln Phe Lys
            820                 825                 830
Phe Thr Thr Thr Pro Ser Ser Val Gly Glu Gly Phe Val Trp Val Lys
        835                 840                 845
Trp Leu Pro Val Gly Ala Pro Thr Lys Thr Thr Asp Ala Trp Gln Leu
    850                 855                 860
Glu Gly Gly Gly Asn Ser Val Arg Ile Gln Lys Leu Ala Val Ala Gly
865                 870                 875                 880
Met Cys Pro Thr Val Val Phe Lys Ile Ala Gly Ser Arg Ser Gln Ala
                885                 890                 895
Cys Ala Ser Ala Leu Pro Tyr Thr Ser Met Trp Arg Val Val Pro Val
            900                 905                 910
Phe Tyr Asn Gly Trp Gly Ala Pro Thr Lys Glu Lys Ala Thr Tyr Asn
        915                 920                 925
Trp Leu Pro Gly Ala His Phe Gly Ser Ile Leu Leu Thr Ser Asp Ala
    930                 935                 940
His Asp Lys Gly Gly Cys Tyr Leu Arg Tyr Ala Phe Arg Ala Pro Ala
945                 950                 955                 960
Met Tyr Cys Pro Arg Pro Ile Pro Pro Ala Phe Thr Arg Pro Ala Asp
                965                 970                 975
Lys Thr Arg His Lys Phe Pro Thr Asn Ile Asn Lys Gln Cys Thr Asn
            980                 985                 990
Tyr Ser Leu Leu Lys Leu Ala Gly Asp Val Glu Ser Asn Pro Gly Pro
        995                1000                1005
Thr Ile Phe Ser Lys Ala Ser Ala Asp Leu Asn Ala Leu Ser Thr Ser
   1010                1015                1020
Leu Gly Glu Leu Thr Gly Met Leu Lys Asp Leu Lys Ala Lys Ala Glu
1025               1030                1035                1040
Thr Tyr Ser Pro Phe Tyr Lys Met Ala Lys Met Leu Phe Lys Leu Ala
               1045                1050                1055
Thr Leu Ala Val Ala Ala Met Arg Thr Lys Asp Pro Val Val Val Val
           1060                1065                1070
Met Leu Ile Ala Asp Phe Gly Leu Glu Val Phe Asp Thr Gly Phe Phe
       1075                1080                1085
Phe Ser Tyr Phe Gln Glu Lys Leu Gln Pro Tyr Met Lys Thr Ile Pro
   1090                1095                1100
Gly Lys Ile Ser Asp Leu Val Thr Asp Ala Ala Thr Ala Ala Ala Gln
1105               1110                1115                1120
Ile Pro Lys Gly Val Tyr Ser Phe Val Ser Ser Phe Phe Glu Thr Pro
               1125                1130                1135
Glu Gly Val Val Glu Lys Gln Val Ser Leu Arg Thr Val Asn Asp Ile
           1140                1145                1150
Phe Ala Leu Leu Lys Asn Ser Asp Trp Phe Ile Lys Thr Leu Val Ala
       1155                1160                1165
Leu Lys Lys Trp Leu Thr Ser Trp Phe Ala Gln Glu Gln Gln Ala Asp
   1170                1175                1180
Asp Ala Leu Tyr Ser Glu Leu Glu Lys Tyr Pro Leu Tyr Lys Leu Lys
1185               1190                1195                1200
Leu Lys Glu Pro Asp Thr Gln Glu Glu Ala Arg Gln Trp Phe Lys Asp
               1205                1210                1215
Met Gln Gln Arg Ala Leu Ala Val Lys Asp Lys Gly Leu Phe Ser Leu
           1220                1225                1230
Leu Gln Ile Pro Leu Val Asn Leu Pro Gln Ser Arg Pro Glu Pro Val
       1235                1240                1245
Val Cys Val Leu Arg Gly Ala Ser Gly Gln Gly Lys Ser Tyr Leu Ala
   1250                1255                1260
Asn Leu Met Ala Gln Ala Ile Ser Leu Leu Leu Val Gly Lys Gln Asp
1265               1270                1275                1280
Ser Val Trp Ser Cys Pro Pro Asp Pro Thr Tyr Phe Asp Gly Tyr Asn
               1285                1290                1295
Gly Gln Ala Val Val Ile Met Asp Ala Leu Gly Gln Asp Pro Asn Gly
           1300                1305                1310
Ala Asp Phe Lys Tyr Phe Cys Gln Met Val Ser Thr Thr Ala Phe Val
       1315                1320                1325
Pro Pro Met Ala His Leu Asp Asp Lys Gly Ile Pro Phe Thr Ser Pro
   1330                1335                1340
Val Val Ile Cys Thr Thr Asn Leu His Ser Ser Phe Thr Pro Ile Thr
1345               1350                1355                1360
Val Ser Cys Pro Glu Ala Leu Lys Arg Arg Phe Arg Phe Asp Val Thr
               1365                1370                1375
Val Ser Ala Lys Pro Gly Phe Val Arg Thr Val Gly Ser Asn Gln Leu
           1380                1385                1390
Leu Asn Leu Pro Leu Ala Leu Lys Pro Ala Gly Leu Pro Pro His Pro
       1395                1400                1405
Ile Phe Glu Asn Asp Met Pro Ile Ile Asn Gly Gln Ala Val Lys Leu
   1410                1415                1420
Ala Leu Ser Gly Gly Glu Val Thr Ala Phe Glu Leu Ile Glu Met Ile
1425               1430                1435                1440
Leu Ser Glu Val Gln Asn Arg Gln Asp Thr His Lys Met Pro Ile Phe
               1445                1450                1455
Lys Gln Ser Trp Ser Asp Leu Phe Arg Lys Cys Thr Thr Asp Glu Glu
           1460                1465                1470
Gln Lys Met Leu Gln Phe Leu Ile Asp Asn Lys Asp Ser Glu Ile Leu
       1475                1480                1485
Arg Ala Phe Val Ser Glu Arg Ser Ile Leu Leu His Glu Glu Tyr Leu
   1490                1495                1500
Lys Trp Glu Ser Tyr Met Thr Arg Arg Ala Lys Phe His Arg Leu Ala
1505               1510                1515                1520
Ala Asp Phe Ala Met Phe Leu Ser Ile Leu Thr Ser Leu Ile Val Ile
               1525                1530                1535
Phe Cys Leu Val Tyr Ser Met Tyr Gln Leu Phe Lys Thr Pro Asp Glu
           1540                1545                1550
Gln Ser Ala Tyr Asp Pro Ser Thr Lys Pro Lys Pro Lys Thr Gln Glu
       1555                1560                1565
Val Lys Thr Leu Lys Ile Arg Thr Glu Thr Gly Val Pro Ala Thr Asp
   1570                1575                1580
Leu Gln Gln Ser Ile Met Lys Asn Val Gln Pro Ile Glu Leu Tyr Leu
1585               1590                1595                1600
Asp Asn Glu Leu Val Thr Asp Cys Ser Ala Leu Gly Val Tyr Asp Asn
               1605                1610                1615
Ser Tyr Leu Val Pro Leu His Leu Phe Glu Phe Asp Phe Asp Thr Ile
           1620                1625                1630
Val Leu Gly Gly Arg His Tyr Lys Lys Ala Glu Cys Glu Lys Val Glu
       1635                1640                1645
Phe Glu Leu Glu Val Asn Gly Asp Val Val Ser Ser Asp Ala Cys Leu
   1650                1655                1660
Leu Arg Val Ser Ser Gly Pro Lys Val Arg Asn Ile Val His Leu Phe
1665               1670                1675                1680
Thr Asn Glu Ile Glu Leu Lys Lys Met Thr Gln Val Thr Gly Ile Met
               1685                1690                1695
Asn Ser Pro His Gln Ala Arg Thr Val Phe Phe Gly Ser Phe Leu Thr
           1700                1705                1710
Val Arg Lys Ser Ile Leu Thr Ser Asp Gly Thr Val Met Pro Asn Val
       1715                1720                1725
Leu Ser Tyr Ala Ala Gln Thr Ser Arg Gly Tyr Cys Gly Ala Ala Ile
   1730                1735                1740
Val Ala Gly Ser Pro Ala Arg Ile Ile Gly Ile His Ser Ala Gly Thr
1745               1750                1755                1760
Gly Ser Val Ala Phe Cys Ser Leu Val Ser Arg Asp Ala Leu Glu Gln
               1765                1770                1775
Leu Trp Pro Gln Lys Gln Gly Asn Val Ser Arg Leu Asp Asp Asp Val
           1780                1785                1790
Arg Val Ser Val Pro Arg Arg Ser Lys Leu Val Lys Ser Leu Ala Tyr
       1795                1800                1805
Pro Ile Phe Lys Pro Asp Tyr Gly Pro Ala Pro Leu Ser Gln Phe Asp
   1810                1815                1820
Lys Arg Leu Ser Asp Gly Val Lys Leu Asp Glu Val Val Phe Ala Lys
1825               1830                1835                1840
His Thr Gly Asp Lys Glu Ile Ser Ala Gln Asp Gln Lys Trp Leu Leu
               1845                1850                1855
Arg Ala Ala His Val Tyr Ala Gln Lys Val Phe Ser Arg Ile Gly Phe
           1860                1865                1870
Asp Asn Gln Ala Leu Thr Glu Lys Glu Ala Ile Cys Gly Ile Pro Gly
       1875                1880                1885
Leu Asp Lys Met Glu Gln Asp Thr Ala Pro Gly Leu Pro Tyr Ala Gln
   1890                1895                1900
Gln Asn Lys Arg Arg Lys Asp Ile Cys Asp Phe Glu Glu Gly Arg Leu
1905               1910                1915                1920
Lys Gly Ala Glu Leu Gln Lys Asp Arg Phe Met Ala Gly Asp Tyr Ser
               1925                1930                1935
Asn Leu Val Tyr Gln Ser Phe Leu Lys Asp Glu Ile Arg Pro Leu Glu
           1940                1945                1950
Lys Val Arg Ala Gly Lys Thr Arg Leu Ile Asp Val Pro Pro Met Pro
       1955                1960                1965
His Val Val Val Gly Arg Gln Leu Leu Gly Arg Phe Val Ala Lys Phe
   1970                1975                1980
His Glu Ala Asn Gly Phe Asp Ile Gly Ser Ala Ile Gly Cys Asp Pro
1985               1990                1995                2000
Asp Val Asp Trp Thr Arg Phe Gly Leu Glu Leu Glu Arg Phe Arg Tyr
               2005                2010                2015
Val Tyr Ala Cys Asp Tyr Ser Arg Phe Asp Ala Asn His Ala Ala Asp
           2020                2025                2030
Ala Met Arg Val Val Leu Asn Tyr Phe Phe Ser Glu Asp His Gly Phe
       2035                2040                2045
Asp Pro Gly Val Pro Ala Phe Ile Glu Ser Leu Val Asp Ser Val His
   2050                2055                2060
Ala Tyr Glu Glu Lys Arg Tyr Asn Ile Tyr Gly Gly Leu Pro Ser Gly
2065               2070                2075                2080
Cys Ser Cys Thr Ser Ile Leu Asn Thr Ile Leu Asn Asn Val Tyr Ile
               2085                2090                2095
Leu Ala Ala Met Met Lys Ala Tyr Glu Asn Phe Glu Pro Asp Asp Ile
           2100                2105                2110
Gln Val Ile Cys Tyr Gly Asp Asp Cys Leu Ile Ala Ser Asp Phe Glu
       2115                2120                2125
Ile Asp Phe Gln Gln Leu Val Pro Val Phe Ser Ser Phe Gly Gln Val
   2130                2135                2140
Ile Thr Thr Ala Asp Lys Thr Asp Phe Phe Lys Leu Thr Thr Leu Ser
2145               2150                2155                2160
Glu Val Thr Phe Leu Lys Arg Ala Phe Val Leu Thr Ala Phe Tyr Lys
               2165                2170                2175
Pro Val Met Asp Val Lys Thr Leu Glu Ala Ile Leu Ser Phe Val Arg
           2180                2185                2190
Pro Gly Thr Gln Ala Glu Lys Leu Leu Ser Val Ala Gln Leu Ala Gly
       2195                2200                2205
His Cys Glu Pro Glu Gln Tyr Glu Arg Leu Phe Glu Pro Phe Ala Gly
   2210                2215                2220
Met Tyr Phe Val Pro Thr Trp Arg
2225               2230