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Philipp Altrock, PhD

Program: Integrated Mathematical Oncology

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Darwinian selection is a universal feature of life. We unravel how Darwinian evolution works in cancer, which helps to understand and improve cancer therapy and cancer patients’ lives.


    • Integrated Mathematical Oncology
    • Malignant Hematology
    • Blood and Marrow Transplant and Cellular Immunotherapy
    • Cancer Biology and Evolution Program
    • Evolutionary Therapy
    • Cancer Biology and Evolution Program

Education & Training


  • University of Kiel, PhD - Natural Sciences
  • University of Leipzig, MSc - Physics

We apply theoretical evolutionary biology models, together with computational, bioinformatics and statistical approaches to cancer cell biology. Of particular interest to us is how the tumor microenvironment shapes selective pressures that play a fundamental role in the evolution of tumor progression and resistance to therapy.  


  • Johnson B, Altrock PM, Kimmel GJ. Two-dimensional adaptive dynamics of evolutionary public goods games: finite-size effects on fixation probability and branching time. R Soc Open Sci. 2021 May.8(5):210182. Pubmedid: 34084549. Pmcid: PMC8150049.
  • Miroshnychenko D, Baratchart E, Ferrall-Fairbanks MC, Velde RV, Laurie MA, Bui MM, Tan AC, Altrock PM, Basanta D, Marusyk A. Spontaneous cell fusions as a mechanism of parasexual recombination in tumour cell populations. Nat Ecol Evol. 2021 Mar.5(3):379-391. Pubmedid: 33462489.
  • Kimmel GJ, Locke FL, Altrock PM. The roles of T cell competition and stochastic extinction events in chimeric antigen receptor T cell therapy. Proc Biol Sci. 2021 Mar.288(1947):20210229. Pubmedid: 33757357. Pmcid: PMC8059581.
  • Damaghi M, West J, Robertson-Tessi M, Xu L, Ferrall-Fairbanks MC, Stewart PA, Persi E, Fridley BL, Altrock PM, Gatenby RA, Sims PA, Anderson ARA, Gillies RJ. The harsh microenvironment in early breast cancer selects for a Warburg phenotype. Proc Natl Acad Sci U S A. 2021 Jan.118(3). Pubmedid: 33452133. Pmcid: PMC7826394.
  • Chakiryan NH, Kimmel GJ, Kim Y, Hajiran A, Aydin AM, Zemp L, Katende E, Nguyen J, Lopez-Blanco N, Chahoud J, Spiess PE, Fournier M, Dhillon J, Wang L, Moran-Segura C, El-Kenawi A, Mulé J, Altrock PM, Manley BJ. Spatial clustering of CD68+ tumor associated macrophages with tumor cells is associated with worse overall survival in metastatic clear cell renal cell carcinoma. PLoS One. 2021 Apr.16(4):e0245415. Pubmedid: 33882057. Pmcid: PMC8059840.
  • Ferrall-Fairbanks MC, Altrock PM. Investigating Inter- and Intrasample Diversity of Single-Cell RNA Sequencing Datasets. Methods Mol Biol. 2020 Sep.2194:177-186. Pubmedid: 32926367.
  • Kimmel GJ, Dane M, Heiser LM, Altrock PM, Andor N. Integrating Mathematical Modeling with High-Throughput Imaging Explains How Polyploid Populations Behave in Nutrient-Sparse Environments. Cancer Res. 2020 Nov.80(22):5109-5120. Pubmedid: 32938640. Pmcid: PMC7671819.
  • Enderling H, Altrock PM, Andor N, Basanta D, Brown JS, Gatenby RA, Marusyk A, Rejniak KA, Silva A, Anderson ARA. High School Internship Program in Integrated Mathematical Oncology (HIP IMO): Five-Year Experience at Moffitt Cancer Center. Bull Math Biol. 2020 Jul.82(7):91. Pubmedid: 32648152.
  • Kimmel GJ, Gerlee P, Altrock PM. Time scales and wave formation in non-linear spatial public goods games. PLoS Comput Biol. 2019 Sep.15(9):e1007361. Pubmedid: 31545788. Pmcid: PMC6776369.
  • Gerlee P, Altrock PM. Persistence of cooperation in diffusive public goods games. Phys Rev E. 2019 Jun.99(6-1):062412. Pubmedid: 31330651.
  • Ferrall-Fairbanks MC, Glazar DJ, Brady RJ, Kimmel GJ, Zahid MU, Altrock PM, Enderling H. Re: Simulation analysis for tumor radiotherapy based on three-component mathematical models. J Appl Clin Med Phys. 2019 Jul.20(7):204-205. Pubmedid: 31145529. Pmcid: PMC6612696.
  • Kimmel GJ, Gerlee P, Brown JS, Altrock PM. Neighborhood size-effects shape growing population dynamics in evolutionary public goods games. Commun Biol. 2019 Feb.2:53. Pubmedid: 30729189. Pmcid: PMC6363775.
  • Kimmel GJ, Gerlee P, Brown JS, Altrock PM. Neighborhood size-effects shape growing population dynamics in evolutionary public goods games. Commun Biol. 2019 Feb.2(1):53. Pubmedid: 31924922.
  • Ferrall-Fairbanks MC, Ball M, Padron E, Altrock PM. Leveraging Single-Cell RNA Sequencing Experiments to Model Intratumor Heterogeneity. JCO Clin Cancer Inform. 2019 Apr.3:1-10. Pubmedid: 30995123. Pmcid: PMC6873939.
  • Altrock PM, Ferlic J, Galla T, Tomasson MH, Michor F. Computational Model of Progression to Multiple Myeloma Identifies Optimum Screening Strategies. JCO Clin Cancer Inform. 2018 Dec.2:1-12. Pubmedid: 30652561. Pmcid: PMC6873949.
  • Gerlee P, Altrock PM. Extinction rates in tumour public goods games. J R Soc Interface. 2017 Sep.14(134). Pubmedid: 28954847. Pmcid: PMC5636271.
  • Altrock PM, Traulsen A, Nowak MA. Evolutionary games on cycles with strong selection. Phys Rev E. 2017 Feb.95(2-1):022407. Pubmedid: 28297871.
  • Altrock PM, Brendel C, Renella R, Orkin SH, Williams DA, Michor F. Mathematical modeling of erythrocyte chimerism informs genetic intervention strategies for sickle cell disease. Am J Hematol. 2016 Sep.91(9):931-937. Pubmedid: 27299299. Pmcid: PMC5093908.
  • Werner B, Scott JG, Sottoriva A, Anderson AR, Traulsen A, Altrock PM. The Cancer Stem Cell Fraction in Hierarchically Organized Tumors Can Be Estimated Using Mathematical Modeling and Patient-Specific Treatment Trajectories. Cancer Res. 2016 Apr.76(7):1705-1713. Pubmedid: 26833122. Pmcid: PMC4900896.
  • Gerlee P, Altrock PM. Complexity and stability in growing cancer cell populations. Proc Natl Acad Sci U S A. 2015 May.112(21):E2742-E2743. Pubmedid: 25941414. Pmcid: PMC4450388.
  • Li XY, Pietschke C, Fraune S, Altrock PM, Bosch TC, Traulsen A. Which games are growing bacterial populations playing?. J R Soc Interface. 2015 Jul.12(108):20150121. Pubmedid: 26236827. Pmcid: PMC4528578.
  • Altrock PM, Liu LL, Michor F. The mathematics of cancer: integrating quantitative models. Nat Rev Cancer. 2015 Dec.15(12):730-745. Pubmedid: 26597528.
  • Marusyk A, Tabassum DP, Altrock PM, Almendro V, Michor F, Polyak K. Non-cell-autonomous driving of tumour growth supports sub-clonal heterogeneity. Nature. 2014 Oct.514(7520):54-58. Pubmedid: 25079331. Pmcid: PMC4184961.
  • Ashcroft P, Altrock PM, Galla T. Fixation in finite populations evolving in fluctuating environments. J R Soc Interface. 2014 Nov.11(100):20140663. Pubmedid: 25165604. Pmcid: PMC4191092.
  • Reeves RG, Bryk J, Altrock PM, Denton JA, Reed FA. First steps towards underdominant genetic transformation of insect populations. PLoS One. 2014 May.9(5):e97557. Pubmedid: 24844466. Pmcid: PMC4028297.
  • Du J, Wu B, Altrock PM, Wang L. Aspiration dynamics of multi-player games in finite populations. J R Soc Interface. 2014 May.11(94):20140077. Pubmedid: 24598208. Pmcid: PMC3973373.
  • Hilbe C, Abou Chakra M, Altrock PM, Traulsen A. The evolution of strategic timing in collective-risk dilemmas. PLoS One. 2014 Jan.8(6):e66490. Pubmedid: 23799109. Pmcid: PMC3682992.
  • Franzenburg S, Fraune S, Altrock PM, Künzel S, Baines JF, Traulsen A, Bosch TC. Bacterial colonization of Hydra hatchlings follows a robust temporal pattern. ISME J. 2013 Apr.7(4):781-790. Pubmedid: 23344242. Pmcid: PMC3603398.
  • Altrock PM, Traulsen A, Galla T. The mechanics of stochastic slowdown in evolutionary games. J Theor Biol. 2012 Oct.311:94-106. Pubmedid: 22814474.
  • Altrock PM, Traulsen A, Reed FA. Stability properties of underdominance in finite subdivided populations. PLoS Comput Biol. 2011 Nov.7(11):e1002260. Pubmedid: 22072956. Pmcid: PMC3207953.
  • Wu B, Altrock PM, Wang L, Traulsen A. Universality of weak selection. Phys Rev E Stat Nonlin Soft Matter Phys. 2010 Oct.82(4 Pt 2):046106. Pubmedid: 21230344.
  • Altrock PM, Traulsen A, Reeves RG, Reed FA. Using underdominance to bi-stably transform local populations. J Theor Biol. 2010 Nov.267(1):62-75. Pubmedid: 20691703.
  • Altrock PM, Gokhale CS, Traulsen A. Stochastic slowdown in evolutionary processes. Phys Rev E Stat Nonlin Soft Matter Phys. 2010 Jul.82(1 Pt 1):011925. Pubmedid: 20866666.
  • Altrock PM, Traulsen A. Deterministic evolutionary game dynamics in finite populations. Phys Rev E Stat Nonlin Soft Matter Phys. 2009 Jul.80(1 Pt 1):011909. Pubmedid: 19658731.


Title: Impact of Stromal Architecture on the Response of Lung Cancers to Targeted Therapies
Sponsor: Florida Biomedical Research Program (FBRP)
PI: Marusyk, A., CO-PI: Altrock, P., CO-PI: Basanta Gutierrez, D.