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Publications

Papers

  1. SE Strawbridge, AK Schrattel, P Humphreys, KA Jones, J Artus, A-K Hadjantonakis, AG Fletcher, J Nichols (2025). Donor embryonic stem cells displace host cells of 8-cell stage chimeras to the extraembryonic lineages by spatial crowding and FGF4 signalling. Development. In press. DOI

  2. L Trinidad, AG Fletcher, D Strutt (2025). The Fat-Dachsous planar polarity pathway competes with hinge contraction to orient polarised cell behaviours during Drosophila wing morphogenesis. Curr Biol. 35:422-430.e3. DOI

  3. CM Cammarota, NS Dawney, PM Bellomio, M Jüng, AG Fletcher, TM Finegan, DT Berglstralh (2024). The mechanical influence of densification on epithelial architecture. PLOS Comput Biol. 20:e1012001. DOI

  4. I Groves, J Holmshaw, D Furley, E Manning, K Chinnaiya, M Towers, BD Evans, M Placzek, AG Fletcher (2023). Accurate staging of chick embryonic tissues via deep learning of salient features. Development. dev.202068. DOI

  5. S Montes-Olivas, D Legge, A Lund, AG Fletcher, AC Williams, L Marucci, M Homer (2023). In-silico and in-vitro morphometric analysis of intestinal organoids. PLOS Comput Biol. 19:e1011386. DOI

  6. SE Strawbridge, E Corujo-Simon, AN Fletcher, J Nichols, AG Fletcher (2023). insideOutside: an accessible algorithm for classifying interior and exterior points, with applications in embryology. Biol Open 12:bio060055 DOI

  7. A Schmidt, T Finegan, M Häring, D Kong, AG Fletcher, Z Alam, J Grosshans, F Wolf, M Peifer (2023). Polychaetoid/ZO-1 strengthens cell junctions under tension while localizing differently than core adherens junction proteins. Mol Biol Cell 4:ar81 DOI

  8. CGM Johnson, AG Fletcher, OS Soyer (2022). ChemChaste: Simulating spatially inhomogenous biochemical reaction-diffusion systems for modelling cell-environment feedbacks. GigaScience. 11:giac051 DOI

  9. A Nestor-Bergmann, G Blanchard, N Hervieux, AG Fletcher, J Etienne, B Sanson (2022). Adhesion-regulated junction slippage controls cell intercalation dynamics in an Apposed-Cortex Adhesion Model. PLOS Comput Biol. 18:e1009812. DOI

  10. R Niehus, NM Oliveira, A Li, AG Fletcher, K Foster (2021). The evolution of strategy in bacterial warfare: quorum sensing, stress responses, and the regulation of bacteriocins and antibiotics. eLife. 10:e69756 DOI

  11. L Gonay, C Spourquet, M Baudoin, L Lepers, P Lemoine, AG Fletcher, E Hanert, CE Pierreux (2021). Modelling of epithelial growth, fission and lumen formation during embryonic thyroid development: a combination of computational and experimental approaches. Front Endocrinol. 12:516 DOI

  12. AG Fletcher, JM Osborne (2021). Seven challenges in the multiscale modelling of multicellular tissues. WIREs Mech Dis. e1527. DOI

  13. K Campbell, E Noel, AG Fletcher, N Bulgakova (2020). Contemporary morphogenesis. Philos Trans R Soc B 375:20190549. DOI

  14. I Groves, M Placzek, AG Fletcher (2020). Of mitogens and morphogens: modelling Sonic Hedgehog mechanisms in vertebrate development. Philos Trans R Soc B 375:20190660. DOI

  15. L Ruske, J Kursawe, A Tsakiridis, V Wilson, AG Fletcher, RA Blythe, LJ Schumacher (2020). Coupled differentiation and division of embryonic stem cells inferred from clonal snapshots. Phys Biol. 17:065009 DOI

  16. TE Gorochowski, S Hauert, J Kreft, L Marucci, NR Stillman, TD Tang, L Bandiera, V Bartoli, DOR Dixon, AJH Fedorec, H Fellermann, AG Fletcher, T Foster, L Giuggioli, A Matyjaszkiewicz, S McCormick, S Montes Olivas, J Naylor, A Rubio Denniss, D Ward (2020). Towards engineering biosystems with emergent collective functions. Front Bioeng Biotechnol. 8:705 DOI

  17. FR Cooper, RE Baker, MO Bernabeu, R Bordas, L Bowler, A Bueno-Orovio, HM Byrne, V Carapella, L Cardone-Noott, J Cooper, S Dutta, BD Evans, AG Fletcher, JA Grogan, W Guo, DG Harvey, M Hendrix, D Kay, J Kursawe, PK Maini, B McMillan, GR Mirams, JM Osborne, P Pathmanathan, JM Pitt-Francis, M Robinson, B Rodriguez, RJ Spiteri, and David J Gavaghan (2020). Chaste: Cancer, Heart and Soft Tissue Environment. J Open Source Softw. 5:1848 DOI

  18. D Ward, S Montes Olivas, AG Fletcher, M Homer, L Marucci (2020). Cross-talk between Hippo and Wnt signalling pathways in intestinal crypts: insights from an agent-based model. Comput Struct Biotechnol J. 18:230-240 DOI

  19. DR Grimes, AG Fletcher (2020). Close encounters of the cell kind: The impact of contact inhibition on tumor growth and cancer models. Bull Math Biol. 82: 20 DOI

  20. TM Finegan, N Hervieux, A Nestor-Bergmann, AG Fletcher, GB Blanchard, B Sanson (2019). The tricellular vertex-specific adhesion molecule Sidekick facilitates polarised cell intercalation during Drosophila axis extension. PLOS Biol. 7:e3000522 DOI

  21. JG Scott, PK Maini, ARA Anderson, AG Fletcher (2019). Inferring tumour proliferative organisation from phylogenetic tree measures in a computational model. Syst Biol. syz070 DOI

  22. RL Cooper, VJ Lloyd, N Di-Poï, AG Fletcher, PM Barrett, G Fraser (2019). Conserved gene signalling and a derived patterning mechanism underlie the development of avian footpad scales. EvoDevo. 10:19 DOI

  23. KH Fisher, D Strutt, AG Fletcher (2019). Experimental and theoretical evidence for bidirectional signalling via core planar polarity protein complexes in Drosophila. iScience. 17:49-66 DOI

  24. JG Scott, A Dhawan, A Hjelmeland, J Lathia, A Chumakova, M Hitomi, AG Fletcher, PK Maini, ARA Anderson (2019). Recasting the cancer stem cell hypothesis: unification using a continuum model of microenvironmental forces. Curr Stem Cell Rep. 5:22-30 DOI

  25. C Pin, A Parker, L Vaux, A Patterson, A Modasia, D Muraro, AG Fletcher, HM Byrne, PK Maini, AJM Watson (2018). Elevated apoptosis impairs epithelial cell turnover and shortens villi in TNF-driven intestinal inflammation. Cell Death Dis. 10:108 DOI

  26. TM Finegan, D Na, AV Skeeters, C Cammarota, TJ Nadasi, NS Dawney, AG Fletcher, PW Oakes, DT Bergstralh (2018). Tissue tension and not interphase cell shape determines cell division orientation in the Drosophila follicular epithelium. EMBO J. 38:e100072 DOI / Cover image

  27. R Cooper, A Thiery, AG Fletcher, D Delbarre, L Rasch, G Fraser (2018). An ancient Turing reaction-diffusion mechanism regulates skin denticle patterning in sharks. Sci Adv. 4:eaau5484 DOI

  28. D Muraro, A Parker, L Vaux, S Filippi, AA Almet, AG Fletcher, AJM Watson, C Pin, PK Maini, HM Byrne (2018). TNFα-driven injury delays cell migration to villi in the intestinal epithelium. J R Soc Interface. 15:20180037 DOI

  29. GB Blanchard, AG Fletcher, LJ Schumacher (2018). The devil is in the mesoscale: mechanical and behavioural heterogeneity in collective cell movement. Semin Cell Dev Biol. S1084-9521(17)30557-8 DOI

  30. J Kursawe, RE Baker, AG Fletcher (2018). Approximate Bayesian computation reveals the importance of repeated measurements for parameterising cell-based models of growing tissues. J Theor Biol. 443:66-81 DOI

  31. B Lambert, AL MacLean, AG Fletcher, AN Combes, MH Little, HM Byrne (2018). Bayesian inference of agent-based models: a tool for studying kidney branching morphogenesis. J Math Biol. 76:1673-1697 DOI

  32. F Cooper, RE Baker, AG Fletcher (2017). Numerical analysis of the immersed boundary method for cell-based simulation. SIAM J Sci Comput. 39:B943-B967. DOI

  33. KH Fisher, D Strutt, AG Fletcher (2017). Integrating planar polarity and tissue mechanics in computational models of epithelial morphogenesis. Curr Opin Syst Biol. 5:41-49. DOI

  34. OJ Maclaren, A Parker, C Pin, SR Carding, AJM Watson, AG Fletcher, HM Byrne, PK Maini (2017). A hierarchical Bayesian framework for understanding the spatiotemporal dynamics of the intestinal epithelium. PLOS Comput Biol. 13:e1005688. DOI

  35. J Kursawe, RE Baker, AG Fletcher (2017). Impact of implementation choices on quantitative predictions of cell-based computational models. J Comput Phys. 345:752-767. DOI

  36. S Godwin, D Ward, E Pedone, M Homer, AG Fletcher, L Marucci (2017). An extended model for culture-dependent heterogeneous gene expression and proliferation dynamics in mouse embryonic stem cells. npj Syst Biol Appl. 3:19. DOI

  37. A Bucksch, A Atta-Boateng, AF Azihou, D Battogtokh, A Baumgartner, BM Binder, SA Braybrook, C Chang, V Coneva, TJ DeWitt, AG Fletcher, MA Gehan, DH Diaz-Martinez, L Hong, AS Iyer-Pascuzzi, LL Klein, S Leiboff, M Li, JP Lynch, A Maizel, JN Maloof, RJC Markelz, CC Martinez, LA Miller, W Mio, W Palubicki, H Poorter, C Pradal, CA Price, E Puttonen, JB Reese, R Rellán-Álvarez, EP Spalding, EE Sparks, CN Topp, JH Williams, DH Chitwood (2017). Morphological plant modeling: Unleashing geometric and topologic potential within the plant sciences. Front Plant Sci. 8:900. DOI

  38. JM Osborne, AG Fletcher, JM Pitt-Francis, PK Maini, DJ Gavaghan (2017). Comparing individual-based approaches to modelling the self-organization of multicellular tissues. PLOS Comput Biol. 13:e1005387. DOI

  39. AG Fletcher, F Cooper, RE Baker (2017). Mechanocellular models of epithelial morphogenesis. Philos Trans R Soc B. 372. DOI

  40. A Parker, OJ Maclaren, AG Fletcher, D Muraro, P Kreuzaler, HM Byrne, PK Maini, AJM Watson, C Pin (2017). Proliferation within small intestinal crypts is the principal driving force for cell migration on villi. FASEB J. 31:636-649. DOI

  41. J Kursawe, R Bardenet, JJ Zartman, RE Baker, AG Fletcher (2016). Robust cell tracking in epithelial tissues through identification of maximum common subgraphs. J R Soc Interface. 13. DOI

  42. RJ Tetley, GB Blanchard, AG Fletcher, RJ Adams, B Sanson (2016). Unipolar distributions of junctional Myosin II identify cell stripe boundaries that drive cell intercalation throughout Drosophila axis extension. eLife 5:e12094. DOI

  43. A Dhawan, TA Graham, AG Fletcher (2016). A computational modelling approach for deriving biomarkers to predict cancer risk in premalignant disease. Cancer Prev Res. 9:283-295. DOI / Cover image

  44. JG Scott, AG Fletcher, ARA Anderson, PK Maini (2016). Spatial metrics of tumour vascular organisation predict radiation efficacy in a computational model. PLOS Comput Biol. 12:e1004712. DOI

  45. J Kursawe, PA Brodskiy, JJ Zartman, RE Baker, AG Fletcher (2015). Capabilities and limitations of tissue size control through passive mechanical forces. PLOS Comput Biol. 11:e1004679. DOI

  46. R Niehus, S Mitri, AG Fletcher, KR Foster (2015). Migration and horizontal gene transfer divide microbial genomes into multiple niches. Nat Commun. 6. DOI

  47. D Nichol, P Jeavons, AG Fletcher, RA Bonomo, PK Maini, JL Paul, RA Gatenby, ARA Anderson, JG Scott (2015). Steering evolution with sequential therapy to prevent the emergence of bacterial antibiotic resistance. PLOS Comput Biol. 11:e1004493. DOI

  48. C Narciso, Q Wu, P Brodskiy, G Garston, RE Baker, AG Fletcher, JJ Zartman (2015). Patterning of wound-induced intercellular Ca2+ flashes in a developing epithelium. Phys Biol. 12:056005. DOI

  49. AG Fletcher, PJ Murray, PK Maini (2015). Multiscale modelling of intestinal crypt organization and carcinogenesis. Math Mod Meth Appl Sci. 25:2563-2585. DOI

  50. OJ Maclaren, HM Byrne, AG Fletcher, PK Maini (2015). Models, measurement and inference in epithelial tissue dynamics. Math Biosci Eng. 12:1321-1340. DOI

  51. DG Harvey, AG Fletcher, JM Osborne, JM Pitt-Francis (2014). A parallel implementation of an off-lattice individual-based model of multicellular populations. Comput Phys Commun. 192:130-137. DOI

  52. B Knapp, R Bardenet, MO Bernabeu, R Bordas, M Bruna, B Calderhead, J Cooper, AG Fletcher, D Groen, B Kuijper, J Lewis, G McInerny, T Minssen, JM Osborne, V Paulitschke, J Pitt-Francis, J Todoric, CA Yates, DJ Gavaghan, CM Deane (2014). Ten simple rules for a successful cross-disciplinary collaboration. PLoS Comput Biol. 11:e1004214. DOI

  53. JG Scott, AG Fletcher, PK Maini, ARA Anderson, P. Gerlee (2014). A filter-flow perspective of hematogenous metastasis offers a non-genetic paradigm for personalized cancer therapy. Eur J Cancer. 50:3068-3075. DOI

  54. DR Grimes, AG Fletcher, M Partridge (2014). Oxygen consumption dynamics in steady-state tumour models. R Soc Open Sci. 1:140080. DOI / Cover image

  55. AM Baker, B Cereser, S Melton, AG Fletcher, M Rodriguez-Justo, PJ Tadrous, A Humphries, G Elia, SAC McDonald, NA Wright, BD Simons, M Jansen, TA Graham (2014). Quantification of crypt and stem cell evolution in the normal and neoplastic human colon. Cell Rep. 8:940-947. DOI

  56. G Rosser, RE Baker, JP Armitage, AG Fletcher (2014). Modelling and analysis of bacterial tracks suggest an active reorientation mechanism in Rhodobacter sphaeroides. J R Soc Interface. 11:20140320. DOI

  57. AG Fletcher, M Osterfield, RE Baker, SY Shvartsman (2014). Vertex models of epithelial morphogenesis. Biophys J. 106:2291-2304. DOI

  58. JM Osborne, MO Bernabeu, M Bruna, B Calderhead, J Cooper, N Dalchau, SJ Dunn, AG Fletcher, R Freeman, D Groen, B Knapp, GJ McInerny, GR Mirams, JM Pitt-Francis, B Sengupta, DW Wright, CA Yates, DJ avaghan, S Emmott, C Deane (2014). Ten simple rules for effective computational research. PLOS Comput Biol. 10:e1003506. DOI

  59. AG Fletcher, JM Osborne, PK Maini, DJ Gavaghan (2013). Implementing vertex dynamics models of cell populations in biology within a consistent computational framework. Prog Biophys Mol Biol. 113:299-326. DOI

  60. K Bloch, H Smith, V van Hamel Parsons, DJ Gavaghan, C Kelly, AG Fletcher, PK Maini, R Callaghan (2013). Metabolic alterations during the growth of tumour spheroids. Cell Biochem Biophys. 68:615-628. DOI

  61. G Rosser, AG Fletcher, DA Wilkinson, JA de Beyer, CA Yates, JP Armitage, PK Maini, RE Baker (2013). Novel methods for analysing bacterial tracks reveal persistence in Rhodobacter sphaeroides. PLoS Comput Biol. 9:e1003276. DOI

  62. JP Vincent, AG Fletcher, LA Baena-Lopez (2013). Mechanisms and mechanics of cell competition in epithelia. Nat Rev Mol Cell Biol. 14:1-11. DOI

  63. G Rosser, AG Fletcher, PK Maini, RE Baker (2013). The effect of sampling rate on observed statistics in a correlated random walk. J R Soc Interface. 10:20130273. DOI

  64. B Lloyd-Lewis, AG Fletcher, TC Dale, HM Byrne (2013). Towards a quantitative understanding of the Wnt/beta-catenin pathway through simulation and experiment. WIREs Syst Biol Med. 5:391-407. DOI

  65. GR Mirams, CJ Arthurs, MO Bernabeu, R Bordas, J Cooper, A Corrias, Y Davit, SJ Dunn, AG Fletcher, DG Harvey, ME Marsh, JM Osborne, P Pathmanathan, JM Pitt-Francis, J Southern, N Zemzemi, DJ Gavaghan (2013). Chaste: an open source C++ library for computational physiology and biology. PLoS Comput Biol. 9:e1002970. DOI

  66. GR Mirams, AG Fletcher, PK Maini, HM Byrne (2012). A theoretical investigation of the effect of dysregulated proliferation and adhesion on monoclonal conversion in the colonic crypt. J Theor Biol. 312:143-156. DOI

  67. R Dello Ioio, C Galinha, AG Fletcher, SP Grigg, A Molnar, V Willemsen, B Scheres, S Sabatini, D Baulcombe, PK Maini, M Tsiantis (2012). A PHABULOSA/cytokinin feedback loop controls root growth in Arabidopsis. Curr Biol. 22:1699-1704. DOI

  68. AG Fletcher, CJW Breward, SJ Chapman (2012). Mathematical modelling of monoclonal conversion in the colonic crypt. J Theor Biol. 300:118-133. DOI

  69. SJ Dunn, AG Fletcher, SJ Chapman, DJ Gavaghan, JM Osborne (2011). Modelling the role of the basement membrane beneath a growing epithelial monolayer. J Theor Biol. 298:82-91. DOI

  70. PJ Murray, A Walter, AG Fletcher, CM Edwards, MJ Tindall, PK Maini (2011). Comparing a discrete and continuum model of the intestinal crypt. Phys Biol. 8:026011. DOI

  71. JM Osborne, A Walter, SK Kershaw, GR Mirams, AG Fletcher, P Pathmanathan, DJ Gavaghan, OE Jensen, PK Maini, HM Byrne (2010). A hybrid approach to multiscale modelling of cancer. Phil Trans R Soc A. 368:5013-5028. DOI

  72. IMM van Leeuwen, GR Mirams, A Walter, AG Fletcher, PJ Murray, JM Osborne, S Varma, SJ Young, J Cooper, B Doyle, JM Pitt-Francis, L Momtahan, P Pathmanathan, JP Whiteley, SJ Chapman, DJ Gavaghan, OE Jensen, JR King, PK Maini, SL Waters, HM Byrne (2009). An integrative computational model for intestinal tissue renewal. Cell Prolif. 42:617-636. DOI

  73. P Pathmanathan, J. Cooper, AG Fletcher, GR Mirams, L. Momtahan, PJ Murray, JM Osborne, JM Pitt-Francis, SJ Chapman (2009). A computational study of discrete mechanical tissue models. Phys Biol. 6:036001. DOI

  74. JM Pitt-Francis, P Pathmanathan, MO Bernabeu, R Bordas, J Cooper, AG Fletcher, GR Mirams, P Murray, JM Osborne, A Walter, SJ Chapman, A Garny, IMM van Leeuwen, PK Maini, B Rodriguez, SL Waters, JP Whiteley, HM Byrne, DJ Gavaghan (2009). Chaste: a test-driven approach to software development for biological modelling. Comput Phys Commun. 180:2452-2471. DOI

Book chapters

  1. JG Scott, P Gerlee, D Basanta, AG Fletcher, PK Maini, ARA Anderson (2013). Mathematical modeling of the metastatic process. In Experimental Metastasis: Modelling and Analysis, 189-208, Springer Netherlands. DOI

  2. AG Fletcher, GR Mirams, PJ Murray, A Walter, JW Kang, KH Cho, PK Maini, HM Byrne (2010). Multiscale modeling of colonic crypts and early colorectal cancer. In Multiscale Cancer Modeling, 111-134, CRC Press. DOI

Preprints

  1. AN Fletcher, AG Fletcher (2024). Variance of the distance to the boundary of convex domains in R2 and R3. arXiv

  2. SH Friedman, ARA Anderson, DM Bortz, AG Fletcher, HB Frieboes, A Ghaffarizadeh, DR Grimes, A Hawkins-Daarud, S Hoehme, EF Juarez, C Kesselman, RMH Merks, SM Mumenthaler, PK Newton, K-A Norton, R Rawat, RC Rockne, D Ruderman, J Scott, SS Sindi, JL Sparks, K Swanson, DB Agus, P Macklin (2016). MultiCellDS: a standard and a community for sharing multicellular data. bioRxiv

  3. A Buttenschoen, A Elkenawi, F El Moustaid, AG Fletcher, C Grassberger, E Kim, A Marusyk, H-L McClelland, D Miroshnychenko, D Nichol, J Mullinax, C O’Farrely (2016). Harnessing the lymphocyte meta-phenotype to optimize adoptive cell therapy. bioRxiv

  4. PA Brodskiy, PM Eberts, C Narciso, J Kursawe, AG Fletcher, JJ Zartman (2016). QuickStitch for seamless stitching of confocal mosaics through high-pass filtering and recursive normalization. bioRxiv