Group Publications




Roulston, C., Luke, G.A., de Felipe, P., Ruan, L., Cope, J., Nicholson, J., Sukhodub, A., Tilsner, J. & Ryan, M.D. (2016). ‘2A-Like’ signal sequences mediating translational recoding: A novel form of dual protein targeting. Traffic (in the press).

Luke, G.A., Roulston, C., Tilsner, J. & Ryan, M.D. Growing Uses of 2A in Plant Biotechnology. in 'Biotechnology'. InTech (Publisher) Ekinci, D. & Mayis, O. (eds). ISBN 978-953-51-4157-0 (in the press)

Tulloch,F., Atkinson, N.J., Evans, D.J., Ryan, M.D. & Simmonds, P. (2014). The attenuated replication phenotype of codon pair de-optimised RNA viruses is an artefact of increasing CpG and UpA dinucleotide frequencies - implications for the design and safety evaluation of future live attenuated vaccines. eLIFE Dec 9;3. doi: 10.7554/eLife.04531.

Forrest, S., Lear, Z., Herod, M.R., Ryan, M.D., Rowlands, D.J & Stonehouse, N.J. (2014). Inhibition of the FMDV sub-genomic replicon by RNA aptamers. J. Gen. Virol. 95, 2649-2657.

Tulloch, F., Pathania, U., Luke, G.A., Nicholson, J., Stonehouse, N.J., Rowlands,, D.J., Jackson, T., Tuthill, T., Haas, J., Lamond, A.I. & Ryan, M.D. (2014). FMDV replicons encoding green fluorescent protein are replication competent. J. Virol. Methods 209, 35-40.

Luke, G.A., Pathania, U.S., Roulston, C., De Felipe, P. & Ryan, M.D. (2014). DxExNPGP - Motives for the motif. Recent Research Developments in Virology India: Research Signpost, 9, 25-42.

Luke, G.A. & Ryan, M.D. (2013). The protein co-expression problem in biotechnology and biomedicine: virus 2A and 2A-like sequences provide a solution. Future Virology 8, 983-981.

Minskaia, E., Nicholson, J. & Ryan, M.D. (2013). Optimisation of the foot-and-mouth disease virus 2A co-expression system for biomedical applications. BMC Biotech. 13, Article #67.

Luke, G.A., Roulston, C., Odon, V., de Felipe, P., Sukhodub, A. & Ryan, M.D. (2013). Lost in translation: The biogenesis of non-LTR retrotransposon proteins. Mobile Genetic Elements 3:6 e27525

Odon, V., Luke, G.A., de Felipe, P., Ruan, L., Roulston, C., Escuin-Ordinas, H., Brown, J.D., Ryan M.D. & Sukhodub, A. (2013). 2A translational recoding elements within APE-type non-LTR retrotransposons of multicellular organisms. Mol. Biol. Evol. 30, 1955-1965.

Minskaia, E. & Ryan, M.D. (2013). Protein co-expression using FMDV 2A: effect of ‘linker’ residues. Biomed. Res. Int. 2013:291730. doi: 10.1155/2013/291730.

Loughran, G., Libbey, J.E., Uddowla, S., Scallan, M.F., Ryan, M.D., Fujinami, R.S., Rieder, E. & Atkins, J.F. (2013). Theiler’s murine encephalomyelitis virus contrasts with Encephalomyocarditis and Foot-and-Mouth Disease viruses in its functional utilization of the StopGo non-standard translation mechanism. J. Gen. Virol. 94, 348-353.

Ryan, M.D. (2013). “MS.717 |Togavirin”. In ‘Handbook of Proteolytic Enzymes’. Rawlings, N.D. & Salvesen, G (eds). Elsevier Ltd.

Sharma, P., Yan, F., Doronina, V. Escuin-Ordinas, H., Ryan, M.D. & Brown, J. (2011). 2A peptides provide distinct solutions to driving stop-carry on translational recoding. Nuc. Acids Res. 40, 1-9.

Walter, C.T., Pringle, F.M., Nakayinga, R., de Felipe, P., Ryan, M.D., Ball, L.A. & Dorrington, R.A. (2010). Genome organization and translation products of Providence virus: insight into a unique tetravirus. J. Gen. Virol. 91, 2836-2835.

de Felipe, P., Luke, G.A., Brown, J.D. & Ryan, M.D. (2010). Inhibition of 2A-mediated 'cleavage' of certain artificial polyproteins bearing N-terminal signal sequences. Biotechnol. J. 5, 213-223.

Martínez-Salas, E. & Ryan, M.D. (2010). ‘Translation and Protein Processing’. In ‘Picornaviruses: Molecular Biology, Evolution and Pathogenesis’. Domingo, E., Ehrenfeld, E. & Roos, R.P. (eds). pp. 141-161, ASM Press.

Brown, J. & Ryan, M.D. (2010). ‘Ribosome “Skipping”: “Stop-Carry On” or “StopGo” Translation’. In ‘Recoding: Expansion of Decoding Rules Enriches Gene Expression’. Atkins, J.F. & Gestland, R.F. (eds.) pp. 101-122, Springer Science+Business Media.

Murray, L., Luke, G.A., Ryan, M.D., Wileman, T. & Knox, C.M. (2009). Amino acid substitutions within the 2C coding sequence of Theiler's Murine Encephalomyelitis virus alter virus growth and affect protein distribution. Virus Res. 144, 74-82

Luke, G.A., Escuin, H., de Felipe, P. & Ryan, M.D. (2009). 2A to the Fore - Research, Technology and Applications. Biotechnology & Genetic Engineering 26, 235-272.

Armer, H., Moffat, K., Wileman, T., Belsham, T., Jackson, T., Duprex, W.P., Ryan, M. & Monaghan, P. (2008). Foot-and-mouth disease virus, but not bovine enterovirus, targets the host cell cytoskeleton via the nonstructural protein 3Cpro. J. Virol. 82, 10556-10566.

Atkins, J.F. & Ryan, M.D. (2008). Foot and Mouth Achille’s heel? Nature Biotech. 26, 1335-1336.

Doronina, V.A., de Felipe, P., Wu, C., Sharma, P., Sachs, M.S., Ryan, M.D. & Brown, J.D. (2008). Dissection of a co-translational nascent chain separation event. Biochem. Soc. Trans. 36, 712-716.

Doronina, V.A., Wu, C., de Felipe, P., Sachs, M.S., Ryan, M.D. & Brown, J. (2008). Site-specific release of nascent chains from ribosomes at a sense codon. Mol. Cell. Biol. 28, 4227- 4239.

Hughes, L. E. & Ryan, M. D. (2008). 'Enteroviruses of Animals'. In Encyclopedia of Virology. (B.W.J. Mahy & M.H.V. Van Regenmortel, Editors Oxford: Elsevier Press.) pp. 123-129.

Luke, G.A., de Felipe, P., Lukashev, A., Kallioinen, S.E., Bruno, E.A & Ryan, M.D. (2008). The occurrence, function, and evolutionary origins of ‘2A-like’ sequences in virus genomes. J. Gen. Virol. 89, 1036-1042.

Funston, G.M., Kallioinen, S.E., de Felipe, P., Ryan, M.D. & Iggo, R.D. (2008). Expression of heterologous genes in oncolytic adenoviruses using picornaviral 2A sequences that trigger ribosome skipping. J. Gen. Virol. 89, 389-396.

Atkins, J.F., Wills, N.M., Loughran, G., Wu, C-Y., Parsawar, K. Ryan, M.D., Wang, C.H. & Nelson, C.C. (2007). A case for "StopGo": Reprogramming translation to augment codon meaning of GGN by promoting unconventional termination (Stop) after addition of glycine and then allowing continued translation (Go). RNA 13, 1-8.

Li, X. Halpin, C. & Ryan, M.D. (2007). A novel cleavage site within the P1 polyprotein of potato leafroll virus. J. Gen. Virol. 88, 1620-1623

Luke, G.A . (2007). Le peptide 2A du virus de la fievre aphteuse. Virologie 11, 371-380.

Moffat, K., Knox, C., Howell, G., Clark, S.J., Yang, Y.G., Belsham, G.J., Ryan, M.D & Wileman,T. (2007). Inhibition of the secretory pathway by the Foot and Mouth Disease Virus 2BC protein is reproduced by co-expression of 2B and 2C and the site of inhibition is determined by the subcellular location of 2C. J. Virol. 81, 1129-1139.

Heras, S.R., Thomas, M.C., García, M., de Felipe, P. García-Pérez, J.L., Ryan, M.D. & López, M.C. (2006). L1Tc non-LTR retrotransposons from Trypanosoma cruzi contain a functional viral-like self-cleaving 2A sequence in frame with the active proteins they encode. Cell. Mol. Life Sci. 63, 1449-1460.

de Felipe, P., Luke, G.A., Hughes, L.E., Gani, D., Halpin, C. & Ryan, M.D. (2006). E unum pluribus: multiple proteins from a self-processing polyprotein. Trends in Biotech. 24, 68-75.

Luke, G.A., de Felipe, P., Cowton, V.M., Hughes, L.E., Halpin, C. & Ryan, M.D. (2006). Self-processing polyproteins: A strategy for co-expression of multiple proteins in plants. Biotechnology & Genetic Engineering Reviews 23, 239-252.

Osborn, M.J., Panoskaltsis-Mortari, A., McElmurry, R.T., Bell, S.K., Vignali, D.A.A., Ryan, M.D., Wilber, A., McIvor, R.S., Tolar, J. & Blazar, B.R. (2005). A picornaviral ‘2A-like’ sequence based tricistronic vector allowing for high level therapeutic gene expression coupled to a dual reporter system. Mol. Ther. 12, 569-574.

Moffat,K., Howell, G., Knox, C., Belsham, G.J., Monaghan, P., Ryan, M.D. & Wileman, T. (2005). Effects of Foot-and-Mouth Disease virus non-structural proteins on the structure and function of the early secretory pathway: 2BC but not 3A blocks ER to Golgi transport. J. Virol. 79, 4382-4395.

Knox, C., Moffat, K., Ali, S., Ryan, M.D. & Wileman, T. (2005). Foot-and-mouth disease virus replication sites form next to the nucleus and close to the Golgi apparatus, but exclude marker proteins associated with host membrane compartments. J. Gen. Virol. 86, 687-696.

de Felipe, P. & Ryan, M.D. (2004). Targeting of proteins derived from self-processing polyproteins containing multiple signal sequences. Traffic 5, 616 - 626.

El Amrani, A., Barakate, A., Askari, B.M., Li, X., Roberts, A.G., Ryan, M.D. & Halpin, C. (2004) Coordinate expression and independent subcellular targeting of multiple proteins from a single transgene. Plant Physiology 135, 16-24.

Monaghan, P., Cook, H., Jackson, T., Ryan, M. & Wileman, T. (2004). The ultrastructure of the developing replication site in foot-and-mouth disease virus-infected BHK-38 cells. J. Gen. Virol. 85, 933-946.

Halpin, C. & Ryan, M.D. (2004). "Redirecting Metabolism by Co-ordinate Manipulation of Multiple Genes" in Metabolic Engineering in the Post-Genomic Era', Kholodenko, B. and Westerhoff, H. (eds.) Horizon Scientific Press, Norfolk, England.

Ryan, M.D., Donnelly, M.L.L., Flint, M., Cowton, V.M., Luke, G., Hughes, L.E., Knox, C. & de Felipe, P. (2004). "Foot-and-Mouth Disease Virus Proteinases". In Foot-And-Mouth Disease: current perspectives. E. Domingo & F. Sobrino (eds). Horizon Scientific Press, Norfolk, England. pp. 53-76.

de Felipe, P. Hughes, L.E. Ryan, M.D. & Brown, J.D. (2003). Co-translational, intraribosomal cleavage of polypeptides by the foot-and-mouth disease virus 2A peptide. J. Biol. Chem. 278, 11441-11448.

de Felipe, P. (2002). Polycistronic viral vectors. Current Gene Therapy 2, 355-378.

Ryan, M.D., Luke, G., Hughes, L.E., Cowton, V.M., ten Dam, E., Li, X., Donnelly, M.L.L., Mehrotra, A. & Gani, D. (2002). The aphtho- and cardiovirus ‘primary’ 2A/2B polyprotein ‘cleavage’.   In ‘Molecular Biology of Picornaviruses’ Semler, B. and Wimmer, E. (eds.) pp. 213-223. ASM Press, Washington, USA.

Halpin, C., Barakate, A., Askari, B., Abbott, J. and Ryan, M. D. (2001). Enabling technologies for manipulating multiple genes on complex pathways. Plant Mol. Biol. 47, 295-310.

Klump, H., Schiedlmeier, B., Vogt, B., Ryan, M.D., Ostertag, W. & Baum, C. (2001). Retroviral vector-mediated expression of HoxB4 in hematopoietic cells using a novel coexpression strategy. Gene Therapy  8, 811-817.

Luke, G. & Ryan M.D. (2001). Translating the message. Biologist  48, 79-82.

Donnelly, M.L.L., Luke, G., Mehrotra, A., Li, X., Hughes, L.E., Gani, D. & Ryan, M.D. (2001). Analysis of the aphthovirus 2A/2B polyprotein ‘cleavage’ mechanism indicates not a proteolytic reaction, but a novel translational effect: a putative ribosomal ‘skip’. J. Gen. Virol. 82, 1013-1025. (download this paper)

Donnelly, M.L.L., Hughes, L.E., Luke, G., Mendoza, H., ten Dam, E., Gani, D. & Ryan, M.D. (2001). The ‘cleavage’ activities of FMDV 2A site-directed mutants and naturally-occurring ‘2A-Like’ sequences. J. Gen. Virol. 82, 1027-1041.

Li, X,  Ryan, M.D. & Lamb, J. (2000). The P1 protein of potato leaf-roll luteovirus contains a serine proteinase domain. J. Gen. Virol. 81, 1857-1864.

Ryan, M.D., Flint, M., Donnelly, M.L.L., Byrne, E & Cowton, V. (2000). Analysis of RNA virus-encoded proteinases. In 'RNA Viruses - A Practical Approach' pp. 69-84 (ed. A. Cann), Oxford University Press.

Mehotra, A.P., Ryan, M.D. & Gani, D. (2000). Synthesis of 6-dimethyl amino-9- [3'- (O-methyl)S) -[UL-14C] -tyrosinylamino) -3'-deoxy-b-D -ribofuranosyl] Purine. J. Labelled Cpd. Radiopharm.43, 623-634.

ten Dam, E., Flint, M. & Ryan, M.D. (1999). Virus Encoded Proteinases of the Togaviridae. J. Gen. Virol. 80, 1879-1888. (download this paper)

Chaplin, P.J., Camon, E.B., Villarreal-Ramos, B., Flint, M., Ryan, M.D. & Collins, R.A. (1999). Production of interlukin-12 as a self-processing polypeptide. J. Interferon & Cytokine Res. 19, 235-241.

Ryan, M.D., Donnelly, M.L.L., Lewis, A., Mehrotra, A.P., Wilkie, J. & Gani, D. (1999). A model for non-stoichiometric, co-translational protein scission in eukaryotic ribosomes. Bioorganic Chem. 27, 55-79.

de Felipe, P., Martín, V., Cortés, M.L., Ryan, M.D. & Izquierdo, M. (1999). Use of the 2A sequence from foot-and-mouth disease virus in the generation of retroviral vectors for gene therapy. Gene Therapy 6, 198-208.

Halpin, C., Cooke, S.E., Barakate, A., El Amrani, A. & Ryan, M.D. (1999). Self-processing polyproteins - a system for co-ordinate expression of multiple proteins in transgenic plants. Plant J. 17, 453-459.

Ryan, M.D., Monoghan, S. & Flint, M. (1998).Virus-encoded proteinases of the Flaviviridae. J. Gen. Virol.  79,  947-959. (download this paper)

Gani, D., Lewis, A., Rutherford, T., Wilkie, J., Stirling, I., Thierry, J., Lightfoot, P. & Ryan, M.D. (1998). Design, synthesis, structure and properties of an a-helix cap template derived from N-[2(S)-2-choloropropionyl]- (2S)- Pro-(2R)-Ala-(2S,4S)- 4-thioPro-OMe which initiates a-helical structures.
Tetrahedron54, 15793-15819.

Lewis, A., Ryan, M.D.& Gani, D. (1998). Design, construction and properties of peptide N-terminal cap templates devised to initiate a-helices. Part 1. Caps derived from N- (-4-chlorobutyryl)-(2S)- Pro-(2S)-Pro-(2S)-Ala-Ome and -[(2S)- chloropropionyl] -(2S)- Pro-(2S) -Pro- (2S,4S) - 4-hydroxyPro- OMe. J. Chem. Soc.- Perkin Trans. 1, #22, 3767-3775.

Mackay, D.K.J., Forsyth, M.A., Davies, P.R., Berlinzani, A.,  Belsham, G.J.,  Flint, M. & Ryan, M.D.(1998). Differentiating infection from vaccination in foot-and-mouth disease using a panel of recombinant, non-structural proteins in ELISA. Vaccine 16, 446-459.

Collen, T., Baron, J., Childerstone, A., Corteyn, A.,  Doel, T.R., Flint, M., Garcia-Valcarcel, M., Parkhouse, R.M.E. & Ryan, M.D. (1998). Heterotypic recognition of recombinant FMDV proteins by bovine T-cells: The polymerase (P3Dpol) as an immunodominant immunogen. Virus Research  56, 125-133.

Ryan, M.D., Donnelly, M.L.L. & Gani, D. (1998). Aphtho-, cardiovirus 2A autolytic sequence. In ‘A Handbook of Proteolytic Enzymes’. Barrett, A. and Rawlings, N.J. eds. pp 1598-1600. Academic Press, London.

Halpin, C. & Ryan, M.D. (1998). Expression of self processing polyproteins in transgenic plants. U.S. Patent 5,846,767. Assignee - Zeneca Limited, London (PCT No. PCT/ GB94/ 02765).

Ryan, M.D. & Flint, M. (1997). Virus-encoded proteinases of the picornavirus super-group. J. Gen. Virol. 78, 699-723. (download this paper)

Donnelly, M., Gani, D., Flint, M., Monoghan, S. & Ryan, M.D. (1997). The cleavage activity of aphtho- and cardiovirus 2A proteins. J. Gen. Virol.78,  13-21.

Hartzoulakis, B., Rutherford, T.J., Ryan, M.D. & Gani, D. (1996). Synthesis and properties of a biocompatible analogue for b-turn protein structural motifs based on 5-amino-3-pentynoic acid. Tetrahedron Letters 37, 6911-66914.

Garcia-Valcarel, M., Doel,T., Collen,T., Ryan, M.D.& Parkhouse, R.M.E. (1996). Recognition of Foot-and-Mouth Disease Virus (FMDV) and its capsid protein VP1 by bovine peripheral T-lymphocytes. J. Gen. Virol. 77, 727-735.

Precious, B., Young, D.F., Bermingham, A., Fearns, R., Ryan, M.D. & Randall, R.E. (1995). Inducible expression of the P, V, and NP genes of the paramyxovirus simian virus 5 in cell lines and an examination of the NP-P and NP-V interactions. J. Virol.69, 8001-8010.

Newman, J.F.E., Piatti, P.G., Ryan, M.D. & Brown, F. (1994). Function of minor polypeptides in foot-and-mouth disease virus and poliovirus. Trends in Microbiology 2, 494-497.

Ryan, M.D. & Drew, J. (1994). Foot-and-mouth disease virus 2A oligopeptide mediated cleavage of an artificial polyprotein. EMBO J. 134, 928-933.

Newman, J.F.E., Piatti, P.G., Gorman, B.M., Burrage, T.G., Ryan, M.D., Flint, M. & Brown, F. (1994). Foot-and-mouth disease virus particles contain replicase protein 3D. Proc. Natl. Acad. Sci. U.S.A.  91, 733-737.

King, A.M.Q., Lomonossoff, G.P. & Ryan, M.D. (1991). Picornaviruses and their relatives in the plant kingdom. Seminars in Virology  2, 11-17.

Ryan, M.D., King, A.M.Q. & Thomas, G.P. (1991). Cleavage of foot-and-mouth disease virus polyprotein is mediated by residues located within a 19 amino acid sequence. J. Gen. Virol. 72, 2727-2732.

Roosien, J., Belsham, G.J., Ryan, M.D., King, A.M.Q. & Vlak, J.M. (1990). Synthesis of foot-and-mouth disease virus capsid proteins in insect cells using  baculovirus expression vectors. J. Gen. Virol. 71, 1703-1711.

Belsham, G.J., Brangwyn, J.K., Ryan, M.D., Abrams, C.C. & King, A.M.Q. (1990). Intracellular expression and processing of foot-and-mouth disease virus capsid precursors using vaccinia virus vectors; Influence of the L protease. Virology 176, 524-530.

Ryan, M.D., Jenkins, O., Hughes, P.J., Knowles, N.J., Brown, A., Booth, D., Minor, P. & Almond, J.W (1990). The complete nucleotide sequence of enterovirus type 70; Relationships with other members of the picornaviridae. J. Gen. Virol. 71, 2291-2299.

Ryan, M.D., Belsham, G.J. & King, A.M.Q. (1989). Specificity of enzyme-substrate interactions in foot-and-mouth disease virus polyprotein processing. Virology 173, 35-45.

Belsham, G.J., Ryan, M.D. & Brangwyn, J.K. (1989). ‘Expression of self-processing foot-and-mouth disease virus capsid precursors by vaccinia virus recombinants’. In ‘Vaccines 89’. Ginsberg, H., Brown, F., Lerner, R.A. & Chanock, R.M. eds pp 445-448 Cold Spring Harbor Laboratory Press, N.Y., USA.

Hyypia, T., Maaronen, M., Auvinen, P., Stalhandske, P., Petterson, U., Stanway,G., Hughes, P., Ryan, M.D., Almond, J.W., Stenvik, M. & Hovi, T. (1987). Nucleic acid sequence relationships between enterovirus serotypes. Mol. and Cellular  Probes 1, 169-176.


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