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Christophe Fraser

Senior Group Leader in Pathogen Dynamics

Education

Affiliations

Projects

Papers

Ratmann O, Grabowski MK, Hall M, Golubchik T, Wymant C, Abeler-Dörner L, Bonsall D, Hoppe A, Brown AL, de Oliveira T, Gall A, Kellam P, Pillay D, Kagaayi J, Kigozi G, Quinn TC, Wawer MJ, Laeyendecker O, Serwadda D, Gray RH, Fraser C, . (2019). Inferring HIV-1 transmission networks and sources of epidemic spread in Africa with deep-sequence phylogenetic analysis. Nature communications, 10(1)

Hall MD, Holden MT, Srisomang P, Mahavanakul W, Wuthiekanun V, Limmathurotsakul D, Fountain K, Parkhill J, Nickerson EK, Peacock SJ, Fraser C. (2019). Improved characterisation of MRSA transmission using within-host bacterial sequence diversity. eLife, (8)

Abeler-Dörner L, Grabowski MK, Rambaut A, Pillay D, Fraser C, . (2019). PANGEA-HIV 2: Phylogenetics And Networks for Generalised Epidemics in Africa. Current opinion in HIV and AIDS, 14(3)

Lehtinen S, Blanquart F, Lipsitch M, Fraser C, . (2019). On the evolutionary ecology of multidrug resistance in bacteria. PLoS pathogens, 15(5)

Volz EM, Le Vu S, Ratmann O, Tostevin A, Dunn D, Orkin C, O'Shea S, Delpech V, Brown A, Gill N, Fraser C, . (2018). Molecular Epidemiology of HIV-1 Subtype B Reveals Heterogeneous Transmission Risk: Implications for Intervention and Control. The Journal of infectious diseases, 217(10)

Blanquart F, Lehtinen S, Lipsitch M, Fraser C. (2018). The evolution of antibiotic resistance in a structured host population. Journal of the Royal Society, Interface, 15(143)

Wymant C, Hall M, Ratmann O, Bonsall D, Golubchik T, de Cesare M, Gall A, Cornelissen M, Fraser C, . (2018). PHYLOSCANNER: Inferring Transmission from Within- and Between-Host Pathogen Genetic Diversity. Molecular biology and evolution, 35(3)

Wymant C, Blanquart F, Golubchik T, Gall A, Bakker M, Bezemer D, Croucher NJ, Hall M, Hillebregt M, Ong SH, Ratmann O, Albert J, Bannert N, Fellay J, Fransen K, Gourlay A, Grabowski MK, Gunsenheimer-Bartmeyer B, Günthard HF, Kivelä P, Kouyos R, Laeyendecker O, Liitsola K, Meyer L, Porter K, Ristola M, van Sighem A, Berkhout B, Cornelissen M, Kellam P, Reiss P, Fraser C, . (2018). Easy and accurate reconstruction of whole HIV genomes from short-read sequence data with shiver. Virus evolution, 4(1)

Nouvellet P, Cori A, Garske T, Blake IM, Dorigatti I, Hinsley W, Jombart T, Mills HL, Nedjati-Gilani G, Van Kerkhove MD, Fraser C, Donnelly CA, Ferguson NM, Riley S. (2018). A simple approach to measure transmissibility and forecast incidence. Epidemics, (22)

Lehtinen S, Blanquart F, Croucher NJ, Turner P, Lipsitch M, Fraser C. (2017). Evolution of antibiotic resistance is linked to any genetic mechanism affecting bacterial duration of carriage. Proceedings of the National Academy of Sciences of the United States of America, 114(5)

Blanquart F, Lehtinen S, Fraser C. (2017). An evolutionary model to predict the frequency of antibiotic resistance under seasonal antibiotic use, and an application to Streptococcus pneumoniae. Proceedings. Biological sciences, 284(1855)

Blanquart F, Wymant C, Cornelissen M, Gall A, Bakker M, Bezemer D, Hall M, Hillebregt M, Ong SH, Albert J, Bannert N, Fellay J, Fransen K, Gourlay AJ, Grabowski MK, Gunsenheimer-Bartmeyer B, Günthard HF, Kivelä P, Kouyos R, Laeyendecker O, Liitsola K, Meyer L, Porter K, Ristola M, van Sighem A, Vanham G, Berkhout B, Kellam P, Reiss P, Fraser C, . (2017). Viral genetic variation accounts for a third of variability in HIV-1 set-point viral load in Europe. PLoS biology, 15(6)

Li LM, Grassly NC, Fraser C. (2017). Quantifying Transmission Heterogeneity Using Both Pathogen Phylogenies and Incidence Time Series. Molecular biology and evolution, 34(11)

Mostowy RJ, Croucher NJ, De Maio N, Chewapreecha C, Salter SJ, Turner P, Aanensen DM, Bentley SD, Didelot X, Fraser C. (2017). Pneumococcal Capsule Synthesis Locus cps as Evolutionary Hotspot with Potential to Generate Novel Serotypes by Recombination. Molecular biology and evolution, 34(10)

Ratmann O, Hodcroft EB, Pickles M, Cori A, Hall M, Lycett S, Colijn C, Dearlove B, Didelot X, Frost S, Hossain AS, Joy JB, Kendall M, Kühnert D, Leventhal GE, Liang R, Plazzotta G, Poon AF, Rasmussen DA, Stadler T, Volz E, Weis C, Leigh Brown AJ, Fraser C, . (2017). Phylogenetic Tools for Generalized HIV-1 Epidemics: Findings from the PANGEA-HIV Methods Comparison. Molecular biology and evolution, 34(1)

Cobey S, Baskerville EB, Colijn C, Hanage W, Fraser C, Lipsitch M. (2017). Host population structure and treatment frequency maintain balancing selection on drug resistance. Journal of the Royal Society, Interface, 14(133)

Ratmann O, Wymant C, Colijn C, Danaviah S, Essex M, Frost SDW, Gall A, Gaiseitsiwe S, Grabowski M, Gray R, Guindon S, von Haeseler A, Kaleebu P, Kendall M, Kozlov A, Manasa J, Minh BQ, Moyo S, Novitsky V, Nsubuga R, Pillay S, Quinn TC, Serwadda D, Ssemwanga D, Stamatakis A, Trifinopoulos J, Wawer M, Leigh Brown A, de Oliveira T, Kellam P, Pillay D, Fraser C. (2017). HIV-1 full-genome phylogenetics of generalized epidemics in sub-Saharan Africa: impact of missing nucleotide characters in next-generation sequences. AIDS research and human retroviruses, 33(11)

Didelot X, Fraser C, Gardy J, Colijn C. (2017). Genomic Infectious Disease Epidemiology in Partially Sampled and Ongoing Outbreaks. Molecular biology and evolution, 34(4)

Blanquart F, Wymant C, Cornelissen M, Gall A, Bakker M, Bezemer D, Hall M, Hillebregt M, Ong SH, Albert J, Bannert N, Fellay J, Fransen K, Gourlay AJ, Grabowski MK, Gunsenheimer-Bartmeyer B, Günthard HF, Kivelä P, Kouyos R, Laeyendecker O, Liitsola K, Meyer L, Porter K, Ristola M, van Sighem A, Vanham G, Berkhout B, Kellam P, Reiss P, Fraser C, . (2017). Correction: Viral genetic variation accounts for a third of variability in HIV-1 set-point viral load in Europe. PLoS biology, 15(7)

Cauchemez S, Nouvellet P, Cori A, Jombart T, Garske T, Clapham H, Moore S, Mills HL, Salje H, Collins C, Rodriquez-Barraquer I, Riley S, Truelove S, Algarni H, Alhakeem R, AlHarbi K, Turkistani A, Aguas RJ, Cummings DA, Van Kerkhove MD, Donnelly CA, Lessler J, Fraser C, Al-Barrak A, Ferguson NM. (2016). Unraveling the drivers of MERS-CoV transmission. Proceedings of the National Academy of Sciences of the United States of America, 113(32)

Ratmann O, van Sighem A, Bezemer D, Gavryushkina A, Jurriaans S, Wensing A, de Wolf F, Reiss P, Fraser C, . (2016). Sources of HIV infection among men having sex with men and implications for prevention. Science translational medicine, 8(320)

Blanquart F, Grabowski MK, Herbeck J, Nalugoda F, Serwadda D, Eller MA, Robb ML, Gray R, Kigozi G, Laeyendecker O, Lythgoe KA, Nakigozi G, Quinn TC, Reynolds SJ, Wawer MJ, Fraser C. (2016). A transmission-virulence evolutionary trade-off explains attenuation of HIV-1 in Uganda. Elife, (5)

Pinsent A, Fraser C, Ferguson NM, Riley S. (2016). A systematic review of reported reassortant viral lineages of influenza A. BMC infectious diseases, (16)

Lythgoe KA, Blanquart F, Pellis L, Fraser C. (2016). Large Variations in HIV-1 Viral Load Explained by Shifting-Mosaic Metapopulation Dynamics. PLoS biology, 14(10)

Croucher NJ, Mostowy R, Wymant C, Turner P, Bentley SD, Fraser C. (2016). Horizontal DNA Transfer Mechanisms of Bacteria as Weapons of Intragenomic Conflict. PLoS biology, 14(3)

Herbeck JT, Mittler JE, Gottlieb GS, Goodreau SM, Murphy JT, Cori A, Pickles M, Fraser C. (2016). Evolution of HIV virulence in response to widespread scale up of antiretroviral therapy: a modeling study. Virus evolution, 2(2)

Cori A, Pickles M, van Sighem A, Gras L, Bezemer D, Reiss P, Fraser C. (2015). CD4+ cell dynamics in untreated HIV-1 infection: overall rates, and effects of age, viral load, sex and calendar time. AIDS (London, England), 29(18)

Lipsitch M, Donnelly CA, Fraser C, Blake IM, Cori A, Dorigatti I, Ferguson NM, Garske T, Mills HL, Riley S, Van Kerkhove MD, Hernán MA. (2015). Potential Biases in Estimating Absolute and Relative Case-Fatality Risks during Outbreaks. PLoS neglected tropical diseases, 9(7)

Nouvellet P, Garske T, Mills HL, Nedjati-Gilani G, Hinsley W, Blake IM, Van Kerkhove MD, Cori A, Dorigatti I, Jombart T, Riley S, Fraser C, Donnelly CA, Ferguson NM. (2015). The role of rapid diagnostics in managing Ebola epidemics. Nature, 528(7580)

Hollingsworth TD, Pilcher CD, Hecht FM, Deeks SG, Fraser C. (2015). High Transmissibility During Early HIV Infection Among Men Who Have Sex With Men-San Francisco, California. The Journal of infectious diseases, 211(11)

Bezemer D, Cori A, Ratmann O, van Sighem A, Hermanides HS, Dutilh BE, Gras L, Rodrigues Faria N, van den Hengel R, Duits AJ, Reiss P, de Wolf F, Fraser C, . (2015). Dispersion of the HIV-1 Epidemic in Men Who Have Sex with Men in the Netherlands: A Combined Mathematical Model and Phylogenetic Analysis. PLoS medicine, 12(11)

Li LM, Grassly NC, Fraser C. (2014). Genomic analysis of emerging pathogens: methods, application and future trends. Genome biology, 15(11)

Fraser C, Lythgoe K, Leventhal GE, Shirreff G, Hollingsworth TD, Alizon S, Bonhoeffer S. (2014). Virulence and pathogenesis of HIV-1 infection: an evolutionary perspective. Science (New York, N.Y.), 343(6177)

Pelat C, Ferguson NM, White PJ, Reed C, Finelli L, Cauchemez S, Fraser C. (2014). Optimizing the precision of case fatality ratio estimates under the surveillance pyramid approach. American journal of epidemiology, 180(10)

Cori A, Ayles H, Beyers N, Schaap A, Floyd S, Sabapathy K, Eaton JW, Hauck K, Smith P, Griffith S, Moore A, Donnell D, Vermund SH, Fidler S, Hayes R, Fraser C, . (2014). HPTN 071 (PopART): a cluster-randomized trial of the population impact of an HIV combination prevention intervention including universal testing and treatment: mathematical model. PLoS One, 9(1)

Kucharski A, Mills H, Pinsent A, Fraser C, Van Kerkhove M, Donnelly CA, Riley S. (2014). Distinguishing Between Reservoir Exposure and Human-to-Human Transmission for Emerging Pathogens Using Case Onset Data. PLoS currents, (6)

Jombart T, Aanensen DM, Baguelin M, Birrell P, Cauchemez S, Camacho A, Colijn C, Collins C, Cori A, Didelot X, Fraser C, Frost S, Hens N, Hugues J, Höhle M, Opatowski L, Rambaut A, Ratmann O, Soubeyrand S, Suchard MA, Wallinga J, Ypma R, Ferguson N. (2014). OutbreakTools: a new platform for disease outbreak analysis using the R software. Epidemics, (7)

Cauchemez S, Fraser C, Van Kerkhove MD, Donnelly CA, Riley S, Rambaut A, Enouf V, van der Werf S, Ferguson NM. (2014). Middle East respiratory syndrome coronavirus: quantification of the extent of the epidemic, surveillance biases, and transmissibility. The Lancet. Infectious diseases, 14(1)

Mostowy R, Croucher NJ, Hanage WP, Harris SR, Bentley S, Fraser C. (2014). Heterogeneity in the frequency and characteristics of homologous recombination in pneumococcal evolution. PLoS genetics, 10(5)

Jombart T, Cori A, Didelot X, Cauchemez S, Fraser C, Ferguson N. (2014). Bayesian reconstruction of disease outbreaks by combining epidemiologic and genomic data. PLoS computational biology, 10(1)

Shepheard MA, Fleming VM, Connor TR, Corander J, Feil EJ, Fraser C, Hanage WP. (2013). Historical zoonoses and other changes in host tropism of Staphylococcus aureus, identified by phylogenetic analysis of a population dataset. PLoS One, 8(5)

Alizon S, Fraser C. (2013). Within-host and between-host evolutionary rates across the HIV-1 genome. Retrovirology, (10)

Lythgoe KA, Pellis L, Fraser C. (2013). Is HIV short-sighted? Insights from a multistrain nested model. Evolution; international journal of organic evolution, 67(10)

Shirreff G, Alizon S, Cori A, Günthard HF, Laeyendecker O, van Sighem A, Bezemer D, Fraser C. (2013). How effectively can HIV phylogenies be used to measure heritability?. Evolution, medicine, and public health, 2013(1)

Cori A, Ferguson NM, Fraser C, Cauchemez S. (2013). A new framework and software to estimate time-varying reproduction numbers during epidemics. American journal of epidemiology, 178(9)

Ratmann O, Donker G, Meijer A, Fraser C, Koelle K. (2012). Phylodynamic inference and model assessment with approximate bayesian computation: influenza as a case study. PLoS computational biology, 8(12)

Lythgoe KA, Fraser C. (2012). New insights into the evolutionary rate of HIV-1 at the within-host and epidemiological levels. Proceedings. Biological sciences, 279(1741)

van Sighem A, Jansen I, Bezemer D, De Wolf F, Prins M, Stolte I, Fraser C. (2012). Increasing sexual risk behaviour among Dutch men who have sex with men: mathematical models versus prospective cohort data. AIDS (London, England), 26(14)

de Silva E, Ferguson NM, Fraser C. (2012). Inferring pandemic growth rates from sequence data. Journal of the Royal Society, Interface, 9(73)

Truscott J, Fraser C, Cauchemez S, Meeyai A, Hinsley W, Donnelly CA, Ghani A, Ferguson N. (2012). Essential epidemiological mechanisms underpinning the transmission dynamics of seasonal influenza. Journal of the Royal Society, Interface, 9(67)

Shirreff G, Pellis L, Laeyendecker O, Fraser C. (2011). Transmission selects for HIV-1 strains of intermediate virulence: a modelling approach. PLoS computational biology, 7(10)

Opatowski L, Fraser C, Griffin J, de Silva E, Van Kerkhove MD, Lyons EJ, Cauchemez S, Ferguson NM. (2011). Transmission characteristics of the 2009 H1N1 influenza pandemic: comparison of 8 Southern hemisphere countries. PLoS pathogens, 7(9)

Fraser C, Cummings DA, Klinkenberg D, Burke DS, Ferguson NM. (2011). Influenza transmission in households during the 1918 pandemic. American journal of epidemiology, 174(5)

Pellis L, Ferguson NM, Fraser C. (2011). Epidemic growth rate and household reproduction number in communities of households, schools and workplaces. Journal of mathematical biology, 63(4)

Müller V, Fraser C, Herbeck JT. (2011). A strong case for viral genetic factors in HIV virulence. Viruses, 3(3)

Fraser C, Hollingsworth TD. (2010). Interpretation of correlations in setpoint viral load in transmitting couples. AIDS (London, England), 24(16)

Colijn C, Cohen T, Fraser C, Hanage W, Goldstein E, Givon-Lavi N, Dagan R, Lipsitch M. (2010). What is the mechanism for persistent coexistence of drug-susceptible and drug-resistant strains of Streptococcus pneumoniae?. Journal of the Royal Society, Interface, 7(47)

Hollingsworth TD, Laeyendecker O, Shirreff G, Donnelly CA, Serwadda D, Wawer MJ, Kiwanuka N, Nalugoda F, Collinson-Streng A, Ssempijja V, Hanage WP, Quinn TC, Gray RH, Fraser C. (2010). HIV-1 transmitting couples have similar viral load set-points in Rakai, Uganda. PLoS pathogens, 6(5)

Van Kerkhove MD, Asikainen T, Becker NG, Bjorge S, Desenclos JC, dos Santos T, Fraser C, Leung GM, Lipsitch M, Longini IM Jr., McBryde ES, Roth CE, Shay DK, Smith DJ, Wallinga J, White PJ, Ferguson NM, Riley S, . (2010). Studies needed to address public health challenges of the 2009 H1N1 influenza pandemic: insights from modeling. PLoS medicine, 7(6)

Baggaley RF, Fraser C. (2010). Modelling sexual transmission of HIV: testing the assumptions, validating the predictions. Current opinion in HIV and AIDS, 5(4)

Hanage WP, Finkelstein JA, Huang SS, Pelton SI, Stevenson AE, Kleinman K, Hinrichsen VL, Fraser C. (2010). Evidence that pneumococcal serotype replacement in Massachusetts following conjugate vaccination is now complete. Epidemics, 2(2)

Bezemer D, de Wolf F, Boerlijst MC, van Sighem A, Hollingsworth TD, Fraser C. (2010). 27 years of the HIV epidemic amongst men having sex with men in the Netherlands: an in depth mathematical model-based analysis. Epidemics, 2(2)

Cauchemez S, Donnelly CA, Reed C, Ghani AC, Fraser C, Kent CK, Finelli L, Ferguson NM. (2009). Household transmission of 2009 pandemic influenza A (H1N1) virus in the United States. The New England journal of medicine, 361(27)

Fraser C, Donnelly CA, Cauchemez S, Hanage WP, Van Kerkhove MD, Hollingsworth TD, Griffin J, Baggaley RF, Jenkins HE, Lyons EJ, Jombart T, Hinsley WR, Grassly NC, Balloux F, Ghani AC, Ferguson NM, Rambaut A, Pybus OG, Lopez-Gatell H, Alpuche-Aranda CM, Chapela IB, Zavala EP, Guevara DM, Checchi F, Garcia E, Hugonnet S, Roth C, . (2009). Pandemic potential of a strain of influenza A (H1N1): early findings. Science (New York, N.Y.), 324(5934)

Hanage WP, Fraser C, Tang J, Connor TR, Corander J. (2009). Hyper-recombination, diversity, and antibiotic resistance in pneumococcus. Science (New York, N.Y.), 324(5933)

Fraser C, Alm EJ, Polz MF, Spratt BG, Hanage WP. (2009). The bacterial species challenge: making sense of genetic and ecological diversity. Science (New York, N.Y.), 323(5915)

Lipsitch M, Colijn C, Cohen T, Hanage WP, Fraser C. (2009). No coexistence for free: neutral null models for multistrain pathogens. Epidemics, 1(1)

Bezemer D, de Wolf F, Boerlijst MC, van Sighem A, Hollingsworth TD, Prins M, Geskus RB, Gras L, Coutinho RA, Fraser C. (2008). A resurgent HIV-1 epidemic among men who have sex with men in the era of potent antiretroviral therapy. AIDS (London, England), 22(9)

Halloran ME, Ferguson NM, Eubank S, Longini IM Jr., Cummings DA, Lewis B, Xu S, Fraser C, Vullikanti A, Germann TC, Wagener D, Beckman R, Kadau K, Barrett C, Macken CA, Burke DS, Cooley P. (2008). Modeling targeted layered containment of an influenza pandemic in the United States. Proceedings of the National Academy of Sciences of the United States of America, 105(12)

Pellis L, Ferguson NM, Fraser C. (2008). The relationship between real-time and discrete-generation models of epidemic spread. Mathematical biosciences, 216(1)

Grassly NC, Fraser C. (2008). Mathematical models of infectious disease transmission. Nature reviews. Microbiology, 6(6)

Hollingsworth TD, Anderson RM, Fraser C. (2008). HIV-1 transmission, by stage of infection. The Journal of infectious diseases, 198(5)

Fraser C, Hollingsworth TD, Chapman R, de Wolf F, Hanage WP. (2007). Variation in HIV-1 set-point viral load: epidemiological analysis and an evolutionary hypothesis. Proceedings of the National Academy of Sciences of the United States of America, 104(44)

Fraser C, Hanage WP, Spratt BG. (2007). Recombination and the nature of bacterial speciation. Science (New York, N.Y.), 315(5811)

Fraser C, Tomassini JE, Xi L, Golm G, Watson M, Giuliano AR, Barr E, Ault KA. (2007). Modeling the long-term antibody response of a human papillomavirus (HPV) virus-like particle (VLP) type 16 prophylactic vaccine. Vaccine, 25(21)

Fraser C. (2007). Estimating individual and household reproduction numbers in an emerging epidemic. PLoS One, 2(8)

Fraser C. (2007). Influenza pandemic vaccines: spread them thin?. PLoS medicine, 4(6)

Hanage WP, Fraser C, Spratt BG. (2006). Sequences, sequence clusters and bacterial species. Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 361(1475)

Hanage WP, Spratt BG, Turner KM, Fraser C. (2006). Modelling bacterial speciation. Philosophical transactions of the Royal Society of London. Series B, Biological sciences, 361(1475)

Ferguson NM, Cummings DA, Fraser C, Cajka JC, Cooley PC, Burke DS. (2006). Strategies for mitigating an influenza pandemic. Nature, 442(7101)

Hanage WP, Fraser C, Spratt BG. (2006). The impact of homologous recombination on the generation of diversity in bacteria. Journal of theoretical biology, 239(2)

Grassly NC, Fraser C. (2006). Seasonal infectious disease epidemiology. Proceedings. Biological sciences, 273(1600)

Wu JT, Riley S, Fraser C, Leung GM. (2006). Reducing the impact of the next influenza pandemic using household-based public health interventions. PLoS medicine, 3(9)

Fraser C. (2005). HIV recombination: what is the impact on antiretroviral therapy?. Journal of the Royal Society, Interface, 2(5)

Ferguson NM, Cummings DA, Cauchemez S, Fraser C, Riley S, Meeyai A, Iamsirithaworn S, Burke DS. (2005). Strategies for containing an emerging influenza pandemic in Southeast Asia. Nature, 437(7056)

Fraser C, Hanage WP, Spratt BG. (2005). Neutral microepidemic evolution of bacterial pathogens. Proceedings of the National Academy of Sciences of the United States of America, 102(6)

Hanage WP, Fraser C, Spratt BG. (2005). Fuzzy species among recombinogenic bacteria. BMC biology, (3)

Fraser C, Riley S, Anderson RM, Ferguson NM. (2004). Factors that make an infectious disease outbreak controllable. Proceedings of the National Academy of Sciences of the United States of America, 101(16)

Ferguson NM, Fraser C, Donnelly CA, Ghani AC, Anderson RM. (2004). Public health. Public health risk from the avian H5N1 influenza epidemic. Science (New York, N.Y.), 304(5673)

Riley S, Fraser C, Donnelly CA, Ghani AC, Abu-Raddad LJ, Hedley AJ, Leung GM, Ho LM, Lam TH, Thach TQ, Chau P, Chan KP, Lo SV, Leung PY, Tsang T, Ho W, Lee KH, Lau EM, Ferguson NM, Anderson RM. (2003). Transmission dynamics of the etiological agent of SARS in Hong Kong: impact of public health interventions. Science (New York, N.Y.), 300(5627)

Garnett GP, Fraser C. (2003). Let it be sexual--selection, aggregation and distortion used to construct a case against sexual transmission. International journal of STD & AIDS, 14(11)

Fraser C, Ferguson NM, De Wolf F, Ghani AC, Garnett GP, Anderson RM. (2002). Antigen-driven T-cell turnover. Journal of theoretical biology, 219(2)