Perelman School of Medicine at the University of Pennsylvania



  1. Zhang D, Li A, Huang M, Yang F, Zhang L, Wellen KE, Xu X, Conn CS, Zou W, Kahn M, Rhoades SD, Weljie AM, Amankulor N, Yoshor D, Ye J, Koumenis C, Gong Y, & Fan Y.      PHGDH-mediated endothelial metabolism drives glioblastoma resistance to CAR T cell immunotherapy. Cell Metabolism 2023;35, 1-18.
  2. Yang F, Zhang D, Zhang L, Winkler J, Gong Y, & Fan Y.       Small-molecule reversal of macrophage-mediated immunosuppression overcomes glioblastoma resistance to immunotherapy. Science Translational Medicine 2023 February 15; 15: eabq3558.


  1. Zou W, Kim H, Diffenderfer ES, Carlson DJ, Koch CJ, Xiao Y, Teo BK, Kim MM, Metz JM, Fan Y, Maity A, Koumenis C, Busch TM, Wiersma R, Cengel KA, Dong L.     A phenomenological model of proton FLASH oxygen depletion effects depending on tissue vasculature and oxygen supply. Front Oncol. 2022 Nov 28;12:1004121. doi: 10.3389/fonc.2022.1004121. PMID: 36518319; PMCID: PMC9742361.
  2. Bayik D, Bartels CF, Lovrenert K, Watson DC, Zhang D, Kay K, Lee J, Lauko A, Johnson S, Lo A, Silver DJ, McGraw M, Grabowski M, Mohammadi AM, Veglia F, Fan Y, Vogelbaum MA, Scacheri P, Lathia JD.     Distinct Cell Adhesion Signature Defines Glioblastoma Myeloid-Derived Suppressor Cell Subsets. Cancer Res. 2022 Nov 15;82(22):4274-4287. doi: 10.1158/0008-5472.CAN-21-3840. PMID: 36126163; PMCID: PMC9664137.
  3. Verginadis II, Avgousti H, Monslow J, Skoufos G, Chinga F, Kim K, Leli NM, Karagounis IV, Bell BI, Velalopoulou A, Salinas CS, Wu VS, Li Y, Ye J, Scott DA, Osterman AL, Sengupta A, Weljie A, Huang M, Zhang D, Fan Y, Radaelli E, Tobias JW, Rambow F, Karras P, Marine JC, Xu X, Hatzigeorgiou AG, Ryeom S, Diehl JA, Fuchs SY, Puré E, Koumenis C.      A stromal Integrated Stress Response activates perivascular cancer-associated fibroblasts to drive angiogenesis and tumour progression. Nat Cell Biol. 2022 Jun;24(6):940-953. 2022 Jun 2. PMID: 35654839; PMCID: PMC9203279.
  4. Huang M., Yang F., Zhang D., Lin M., Duan H. EI-Mayta R., Zhang L., Qin L., Shewale S., Pei L., Mitchell M., Rader D., Fan Y*., Gong Y*.      Endothelial plasticity drives aberrant vascularization and impedes cardiac repair after myocardial infarction. Nature Cardiovascular Research. 2022 April 13; 1: 372–388 Notes: * Co-senior author.
  5. Paul R, Dorsey JF, Fan Y.      Cell plasticity, senescence, and quiescence in cancer stem cells: Biological and therapeutic implications. Pharmacol Ther. 2022 Mar;231:107985. PMID: 34480963.
  6. Jiang J, Yuan J, Hu Z, Zhang Y, Zhang T, Xu M, Long M, Fan Y, Tanyi JL, Montone KT, Tavana O, Vonderheide RH, Chan HM, Hu X, Zhang L.     Systematic illumination of druggable genes in cancer genomes. Cell Rep. 2022 Feb 22;38(8):110400. PMID: 35196490.
  7. Jiang J, Yuan J, Hu Z, Xu M, Zhang Y, Long M, Fan Y, Montone KT, Tanyi JL, Tavana O, Chan HM, Zhang L, Hu X.     Systematic Pan-Cancer Characterization of Nuclear Receptors Identifies Potential Cancer Biomarkers and Therapeutic Targets. Cancer Res. 2022 Jan 1;82(1):46-59. PMID: 34750098.


  1. Hu Z, Yuan J, Long M, Jiang J, Zhang Y, Zhang T, Xu M, Fan Y, Tanyi JL, Montone KT, Tavana O, Chan HM, Hu X, Vonderheide RH, Zhang L.    The Cancer Surfaceome Atlas integrates genomic, functional and drug response data to identify actionable targets. Nature Cancer. 2021 Dec;2(12):1406-1422. PMID: 35121907.
  2. Lamplugh Z, Fan Y.     Vascular Microenvironment, Tumor Immunity and Immunotherapy. Front Immunol. 2021 Dec 20;12:811485.811485. PMID: 34987525
  3. Wang L, Yao L, Duan H, Yang F, Lin M, Zhang R, He Z, Ahn J, Fan Y, Qin L, Gong Y.     Plasminogen Regulates Fracture Repair by Promoting the Functions of Periosteal Mesenchymal Progenitors. J Bone Miner Res. 2021 Nov;36(11):2229-2242. PMID: 34378815.
  4. Zou W, Diffenderfer ES, Cengel KA, Kim MM, Avery S, Konzer J, Cai Y, Boisseu P, Ota K, Yin L, Wiersma R, Carlson DJ, Fan Y, Busch TM, Koumenis C, Lin A, Metz JM, Teo BK, Dong L.     Current delivery limitations of proton PBS for FLASH. Radiother Oncol. 2021 Feb;155:212-218. PMID: 33186682.
  5. Yang F, He Z, Duan H, Zhang D, Li J, Yang H, Dorsey JF, Zou W, Nabavizadeh SA, Bagley SJ, Abdullah K, Brem S, Zhang L, Xu X, Byrne KT, Vonderheide RH*, Gong Y*, & Fan Y*.       Synergistic immunotherapy of glioblastoma by dual targeting of IL-6 and CD40. Nature Comm. 2021 Jun 8; 12(1): 3424. (PMID: 34103524). *Co-corresponding authors.
  6. Ou L, Wang H, Liu Q, Zhang J, Lu H, Luo L, Shi C, Lin S, Dong L, Guo Y, Huang L, Zhu J, Yin X, Huang AC, Karakousis G, Schuchter L, Amaravadi R, Zheng C, Fan Y, Guo W, Xu X.      Dichotomous and stable gamma delta T-cell number and function in healthy individuals. J Immunother Cancer. 2021 May; 9(5): e002274. (PMID: 34011536)
  7. Ma W, Wang Y, Zhang R, Yang F, Zhang D, Huang M, Zhang L, Dorsey JF, Binder ZA, O’Rourke DM, Fraietta JA, Gong Y*, Fan Y*.             Targeting PAK4 to reprogram vascular microenvironment and improve CAR T immunotherapy for glioblastoma. Nature Cancer. 2021; 2, 83–97. *Co-corresponding authors.


  1. Zou W, Diffenderfer ES, Cengel K, Kim M, Avery S, Konzer J, Cai Y, Boisseu P, Ota K, Yin L, Wiersma R, Carlson DJ, Fan Y, Busch TM, Koumenis C, Lin A, Metz JM, Teo BK and Dong L.      Current Delivery Limitations of Proton PBS for FLASH. Radiotherapy and Oncology 2020; 155:212-218.
  2. Duo H, He Z, Lin M, Wang Y, Yang F, Mitteer, R, Kim H, Yeo E, Han H, Qin L, Fan Y, and Gong Y.            Plasminogen regulates mesenchymal stem cellmediated tissue repair after ischemia through cyr61 activation. JCI insight 2020; 5(15), e131376 (PMID:  32759492).
  3. Shan W, Yuan J, Hu J, Jiang J, Wang Y, Loo N, Fan L, Tang Z, Zhang T, Xu M, Pan Y, Lu J, Long M, Tanyi LJ, Montone KT, Fan Y, Hu X, Zhang Y, Zhang L.       Systematic characterization of recurrent genomic alterations in cyclin-dependent kinases reveals potential therapeutic strategies for cancer treatment. Cell Reports 2020; 32 (2), 107884 (PMID:  32668240).
  4. Huang M, Zhang D, Wu J, Xing K, Yeo E, Li C, Zhang L, Holland E, Yao L, Qin L, Binder Z, O’Rourke D, Brem S, Koumenis C, Gong Y, and Fan Y.       Wnt-mediated endothelial transformation into mesenchymal stem cell-like cells induces chemoresistance in glioblastoma.       Science Translational Medicine 2020; 12(532) (PMID:  32102932).
  5. Yuan J, Kensler KH, Hu Z, Zhang Y, Zhang T, Jiang J, Xu M, Pan Y, Long M, Montone KT, Tanyi JL, Fan Y, Zhang R, Hu X, Rebbeck TR, Zhang L.       Integrative comparison of the genomic and transcriptomic landscape between prostate cancer patients of predominantly African or European genetic ancestry.         PLoS Genetics 16 (2), e1008641 (PMID:  32059012)


  1. Fan Y. Vascular detransformation for cancer therapy.      Trends in Cancer 2019; 5(8):460-463. (PMID:  31421902).
  2. Hu Z, Zhou J, Jiang J, Yuan J, Zhang Y, Wei X, Loo N, Wang Y, Pan Y, Zhang T, Zhong X, Long M, Montone K, Tanyi J, Fan Y, Wang TL, Shih IM, Hu X, Zhang L.      Genomic characterization of genes encoding histone acetylation modulator proteins identifies therapeutic targets for cancer treatment.       Nature Comm 2019; 10(1):733. (PMID: 30760718).


  1. Liu T, Xu H, Huang M, Ma W, Saxena D, Lustig RA, Alonso-Basanta M, Zhang Z, O’Rourke DM, Zhang L, Gong Y, Kao GD, Dorsey JF*, & Fan Y*.       Circulating glioma cells exhibit stem cell-like properties.       Cancer Res. 2018 Dec 1;78(23):6632-6642. *, corresponding authors. (PMID: 30322863).
  2. Yuan J, Hu Z, Mahal B, Zhao S, Kensler K, Pi J, Hu X, Wang Y, Jiang J, Zhang Y, Li C, Zhong X, Montone K, Guan G, Tanyi J, Fan Y, Xu X, Long M, Morgan M, Zhang Y, Zhang R, Sood A, Rebbeck T, Dang C, & Zhang L.       Integrated analysis of genetic ancestry and genomic alterations across cancers.        Cancer Cell 2018 Oct 8;34(4):549-560. (PMID: 30300578).
  3. Liu T, Ma W, Xu H, Huang M, Zhang D, He Z, Zhang L, Brem S, O'Rourke DM, Gong Y, Mou Y*, Zhang Z*, and Fan Y*.      PDGF-mediated mesenchymal transformation renders endothelial resistance to anti-VEGF treatment in glioblastoma.          Nature Comm. 2018; 9(1):3439. *corresponding authors (PMID: 30150753)
  4. Lin X, Shen J, Dan P, He X, Xu C, Chen X, Tanyi JL, Montone K, Fan Y, Huang Q, Zhang L, Zhong X.     RNA-binding protein LIN28B inhibits apoptosis through regulation of the AKT2/FOXO3A/BIM axis in ovarian cancer cells.          Signal Transduct Target Ther. 2018 Aug 31;3:23. (PMID: 30174831)
  5. Wang Q, He Z, Huang M, Liu T, Xu H, Ma P, Zhang L, Zamvil SS, Hidalgo J, Zhang Z, O’Rourke DM, Dahmane N, Brem S, Gong Y, & Fan Y.        Vascular niche IL-6 induces macrophage M2 polarization in gliomblastoma through HIF-2α.        Nature Comm. 2018; 9:559. (PMID: 29422647)
  6. Wang Y, Xu H, Liu T, Huang M, Butter P, Li C, Zhang L, Kao G, Gong Y, Maity A, Koumenis C, & Fan Y.      Temporal DNA-PK activation drives genomic instability and therapy resistance in glioma stem cells.       JCI Insight. 2018; 3(3): e98096. (PMID: 29415883)
  7. Guo Y, Hu J, Wang Y, Peng X, Min J, Wang J, Matthaiou E, Cheng Y, Sun K, Tong X, Fan Y, Zhang PJ, Kandalaft LE, Irving M, Coukos G, Li C.      Tumour endothelial marker 1/endosialin-mediated targeting of human sarcoma.   Eur J Cancer. 2018; 90: 111-121. (PMID: 29304474)


  1. Yang L, Zhang Y, Shan W, Hu Z, Yuan J, Pi J, Wang Y, Fan L, Tang Z, Li C, Hu X, Tanyi JL, Fan Y, Huang Q, Montone K, Dang CV, Zhang L.     Repression of BET activity sensitizes homologous recombination-proficient cancers to PARP inhibition.        Science Transl Med. 2017; 9(400): 1645. (PMID: 28747513)


  1. He Z, Mitteer RA, Mou Y, Fan Y.      Chapter 5: Multimodality targeting of glioma cells.      Glioblastoma, 1st edition (Ed. Brem S and Abdullah K), Elsevier, 2016.
  2. Brem S, Wong E, Desai A, Fan Y, Pan E.      Chapter 116: Angiogenesis and brain tumors.      Youmans and Winn Neurological Surgery, 7th edition (Ed. Winn HR), Elsevier, 2016.
  3. Huang M, T Liu, Ma P, Mitteer RA, Zhang Z, Kim HJ, Yeo E, Zhang D, Cai P, Li C, Zhang L, Zhao B, Roccograndi L, O’Rourke DM, Dahmane N, Gong Y, Koumenis C, Fan Y.      c-Met-mediated endothelial plasticity drives aberrant vascularization and chemoresistance in glioblastoma.      J Clin Invest. 2016; 126(5):1801-14. (PMID: 27043280)
  4. Zhang Y, He Q, Hu Z, Feng Y, Fan L, Tang Z, Yuan J, Shan W, Li C, Hu X, Tanyi JL, Fan Y, Huang Q, Montone K, Dang CV, Zhang L.      Long noncoding RNA LINP1 regulates repair of DNA double-strand breaks in triple-negative breast cancer.      Nat Struct Mol Biol. 2016; 23, 522–530. (PMID: 27111890)
  5. Zhang Y, Feng Y, Hu Z, Hu X, Yuan CX, Fan Y, Zhang L.      Characterization of Long Noncoding RNA-Associated Proteins by RNA-Immunoprecipitation.      Methods Mol Biol. 2016; 1402: 19-26. (PMID: 26721480)


  1. Yan X, Hu Z, Feng Y, Hu X, Yuan J, Zhao SD, Zhang Y, Yang L, Shan W, He Q, Fan L, Kandalaft LE, Tanyi JL, Li C, Yuan CX, Zhang D, Yuan H, Hua K, Lu Y, Katsaros D, Huang Q, Montone K, Fan Y, Coukos G, Boyd J, Sood AK, Rebbeck T, Mills GB, Dang CV, Zhang L.      Comprehensive Genomic Characterization of Long Non-coding RNAs across Human Cancers.        Cancer Cell. 2015; 28(4): 529-40. (PMID: 26461095)
  2. Mitteer RA, Wang YL, Shah J, Gordon S, Fager M, Guardiola-Salmeron C, Carabe-Fernandez A*, Fan Y*.      Proton beam radiation induces DNA damage and cell apoptosis in glioma stem cells through reactive oxygen species.        Sci Rep. 2015; 5: 13961. *, co-corresponding authors. (PMID: 26354413)
  3. Cai PQ, Lv XF, Tian L, Luo ZP, Mitteer RA Jr, Fan Y, Wu YP.      CT Characterization of Duodenal Gastrointestinal Stromal Tumors.       Am J Roentgenol. 2015; 204(5):988-93. (PMID: 25905932).
  4. Wu YP, Cai PQ, Tian L, Xu JH, Mitteer RA, Fan Y, Zhang ZF.       Hypertrophic adenoids in patients with nasopharyngeal carcinoma: appearance at magnetic resonance imaging before and after treatment.       Chin J Cancer. 2015; 34:4. (PMID: 25962737).


  1. Zhao B, Yang D, Jiang J, Li J, Fan C, Huang M, Fan Y, Jin Y, Jin Y.      Genome-wide mapping of miRNAs exxpresed in embryonic stem cells and pluripotent stem cells generated by different reprogramming strategies.     BMC Genomics. 2014; 15:488. (PMID: 24942538).
  2. Eswarappa SM, Potdar AA, Koch WJ, Fan Y, Vasu K, Lindner D, Willard B, Graham LM, DiCorleto PE, Fox PL.      Programmed Translational Readthrough Generates Antiangiogenic VEGF-Ax.      Cell. 2014; 157(7):1605-1618. (PMID: 24949972).
  3. Fan Y, Potdar AA, Gong Y, Eswarappa SM, Donnola S, Lathia J, Hambardzumyan D, Rich JN, and Fox PL.      Profilin-1 phosphorylation directs angiocrine expression and glioblastoma progression through induction of HIF-1α.      Nat Cell Biol. 2014; 16(5): 445-456. (PMID: 24747440).
  4. Gong Y, Zhao Y, Li Y, Fan Y, Hoover-Plow J.      Plasminogen regulates cardiac repair after myocardial infraction through its non-canonical function in stem cell homing to the infarcted heart.       J Am Coll Cardiol. 2014; 63(25): 2862-2872. (PMID: 24681141).


  1. Fan Y, Arif A, Gong Y, Jia J, Eswarappa SM, Willard B, Horowitz A, Graham LM, Penn MS, Fox PL.    Stimulus-dependent phosphorylation of profilin-1 in angiogenesis.       Nat Cell Biol. 2012; 14(10): 1046-1056. (PMID: 23000962).
  2. Fan Y, Eswarappa SM, Hitomi M, and Fox PL.    Myo1c facilitates G-actin transport to the leading edge ofmigrating endothelial cells.      J Cell Biol. 2012; 198(1): 47-55. (PMID: 22778278).
  3. Gong Y, Fan Y, Hoover-Plow J.    Plasminogen regulates SDF-1/CXCR4-mediated hematopoietic stem cell mobilization by activation of matrix metalloproteinase-9.       Arterioscler Thromb Vasc Biol. 2011; 31: 2035-2043. (PMID: 21719761).
  4. Fan Y, Gong Y, Ghosh P, Graham L, Fox PL.    Spatial coordination of actin polymerization and ILK/Akt2 activity during endothelial cell migration.        Dev Cell. 2009; 16: 661-674. (PMID: 19460343).