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Michael R. Freeman, PhD


  • Wen H, Lee T, You S, Park SH, Song H, Eilber KS, Anger JT, Freeman MR, Park S, Kim J.
    Urinary metabolite profiling combined with computational analysis predicts interstitial cystitis-associated candidate biomarkers.
    J Proteome Res. 2015 Jan 2;14(1):541-8.
  • Minciacchi VR, Freeman MR, Di Vizio D.
    Extracellular Vesicles in Cancer: Exosomes, Microvesicles and the Emerging Role of Large Oncosomes.
    Semin Cell Dev Biol. 2015 Feb 23.
  • Kim J, Morley S, Le M, Bedoret D, Umetsu DT, Di Vizio D, Freeman MR.
    Enhanced shedding of extracellular vesicles from amoeboid prostate cancer cells: Potential effects on the tumor microenvironment.
    Cancer Biol Ther. 2014 Apr 1;15(4):409-18.
  • Morley S, Hager MH, Pollan SG, Knudsen B, Di Vizio D, Freeman MR.
    Trading in your spindles for blebs: the amoeboid tumor cell phenotype in prostate cancer.
    Asian J. Androl. 2014 Feb 21.
  • Morello M, Minciacchi VR, de Candia P, Yang J, Posadas E, Kim H, Griffiths D, Bhowmick N, Chung LW, Gandellini P, Freeman MR, Demichelis F, Di Vizio D.
    Large oncosomes mediate intercellular transfer of functional microRNA.
    Cell Cycle. 2013 Nov 15;12(22):3526-36.
  • Ayala G, Morello M, Frolov A, You S, Li R, Rosati F, Bartolucci G, Danza G, Adam RM, Thompson TC, Lisanti MP, Freeman MR, Di Vizio D.
    Loss of caveolin-1 in prostate cancer stroma correlates with reduced relapse-free survival and is functionally relevant to tumour progression.
    J Pathol. 2013 Sep;231(1):77-87.
  • Solomon KR, Allott EH, Freeman MR, Freedland SJ.
    Re: Dysregulation of Cholesterol Homeostasis in Human Prostate Cancer Through Loss of ABCA1.
    Eur Urol. 2013 Jun;63(6):1128-9.
  • Park JO, Choi DY, Choi DS, Kim HJ, Kang JW, Jung JH, Lee JH, Kim J, Freeman MR, Lee KY, Gho YS, Kim KP.
    Identification and characterization of proteins isolated from microvesicles derived from human lung cancer pleural effusions.
    Proteomics. 2013 Apr 14.
  • Freeman MR, Solomon KR.
    Statin Drugs and Prostate Cancer: Time to Consider Proactive Strategies in Patients.
    J Urol. 2013 Jan 18.
  • Kim J, Keay SK, You S, Loda M, Freeman MR.
    A synthetic form of frizzled 8-associated antiproliferative factor enhances p53 stability through USP2a and MDM2.
    PLoS ONE. 2012;7(12):e50392.
  • Di Vizio D, Morello M, Dudley AC, Schow PW, Adam RM, Morley S, Mulholland D, Rotinen M, Hager MH, Insabato L, Moses MA, Demichelis F, Lisanti MP, Wu H, Klagsbrun M, Bhowmick NA, Rubin MA, D'Souza-Schorey C, Freeman MR.
    Large oncosomes in human prostate cancer tissues and in the circulation of mice with metastatic disease.
    Am J Pathol. 2012 Nov;181(5):1573-84.
  • Hager MH, Morley S, Bielenberg DR, Gao S, Morello M, Holcomb IN, Liu W, Mouneimne G, Demichelis F, Kim J, Solomon KR, Adam RM, Isaacs WB, Higgs HN, Vessella RL, Di Vizio D, Freeman MR.
    DIAPH3 governs the cellular transition to the amoeboid tumour phenotype.
    EMBO Mol Med. 2012 Aug;4(8):743-60.
  • Pelton K, Freeman MR, Solomon KR.
    Cholesterol and prostate cancer.
    Curr Opin Pharmacol. 2012 Jul 21.
  • Kim J, Di Vizio D, Kim TK, Kim J, Kim M, Pelton K, Clinton SK, Hai T, Hwang D, Solomon KR, Freeman MR.
    The response of the prostate to circulating cholesterol: activating transcription factor 3 (ATF3) as a prominent node in a cholesterol-sensing network.
    PLoS ONE. 2012;7(7):e39448.
  • Yang W, Chung YG, Kim Y, Kim TK, Keay SK, Zhang CO, Ji M, Hwang D, Kim KP, Steen H, Freeman MR, Kim J.
    Quantitative proteomics identifies a {beta}-catenin network as an element of the signaling response to frizzled-8 protein-related antiproliferative factor.
    Mol Cell Proteomics. 2011 Mar 21.
  • Wu X, Gao H, Ke W, Hager M, Xiao S, Freeman MR, Zhu Z.
    VentX trans-activates p53 and p16ink4a to regulate cellular senescence.
    J Biol Chem. 2011 Feb 16.
  • Kim J, Ji M, DiDonato JA, Rackley RR, Kuang M, Sadhukhan PC, Mauney JR, Keay SK, Freeman MR, Liou LS, Adam RM.
    An hTERT-immortalized human urothelial cell line that responds to anti-proliferative factor.
    In Vitro Cell Dev Biol Anim. 2011 Jan;47(1):2-9.
  • Pelton K, Di Vizio D, Insabato L, Schaffner CP, Freeman MR, Solomon KR.
    Ezetimibe reduces enlarged prostate in an animal model of benign prostatic hyperplasia.
    J Urol. 2010 Oct;184(4):1555-9.
  • Peleg S, Sellin JH, Wang Y, Freeman MR, Umar S.
    Am J Physiol Gastrointest Liver Physiol. 2010 Sep;299(3):G593-601.
  • Yang W, Cai Q, Lui VW, Everley PA, Kim J, Bhola N, Quesnelle KM, Zetter BR, Steen H, Freeman MR, Grandis JR.
    Quantitative Proteomics Analysis Reveals Molecular Networks Regulated by Epidermal Growth Factor Receptor Level in Head and Neck Cancer.
    J Proteome Res. 2010 Jun 4;9(6):3073-82.
  • Yang W, Di Vizio D, Kirchner M, Steen H, Freeman MR.
    Proteome scale characterization of human S-acylated proteins in lipid raft-enriched and non-raft membranes.
    Mol Cell Proteomics. 2010 Jan;9(1):54-70.
  • Freeman MR.
    Growing evidence that radionuclide imaging identifies management strategies that improve outcome.
    J Nucl Cardiol.;16(6):844-5.
  • Vaezzadeh AR, Steen H, Freeman MR, Lee RS.
    Proteomics and opportunities for clinical translation in urological disease.
    J Urol. 2009 Sep;182(3):835-43. Review.
  • Di Vizio D, Morello M, Sotgia F, Pestell RG, Freeman MR, Lisanti MP.
    An absence of stromal caveolin-1 is associated with advanced prostate cancer, metastatic disease and epithelial Akt activation.
    Cell Cycle. 2009 Aug;8(15):2420-4.
  • Kim J,Yanagihara Y,Kikugawa T,Ji M,Tanji N,Masayoshi Y,Freeman MR.
    A Signaling Network in Phenylephrine-induced Benign Prostatic Hyperplasia.
    Endocrinology. 2009 Aug;150(8):3576-83.
  • Di Vizio D, Kim J, Hager MH, Morello M, Yang W, Lafargue CJ, True LD, Rubin MA, Adam RM, Beroukhim R, Demichelis F, Freeman MR.
    Oncosome formation in prostate cancer: association with a region of frequent chromosomal deletion in metastatic disease.
    Cancer Res. 2009 Jul 1;69(13):5601-9.
  • Mukhopadhyay NK,Kim J,Cinar B,Ramachandran A,Hager MH,Di Vizio D,Adam RM,Rubin MA,Raychaudhuri P,De Benedetti A,Freeman MR.
    Heterogeneous nuclear ribonucleoprotein K is a novel regulator of androgen receptor translation.
    Cancer Res. 2009 Mar 15;69(6):2210-8.
  • Solomon KR,Pelton K,Boucher K,Joo J,Tully C,Zurakowski D,Schaffner CP,Kim J,Freeman MR.
    Ezetimibe is an inhibitor of tumor angiogenesis.
    Am J Pathol. 2009 Mar;174(3):1017-26.
  • Di Vizio D,Solomon KR,Freeman MR.
    Cholesterol and cholesterol-rich membranes in prostate cancer: an update.
    Tumori. 2008 Sep-Oct;94(5):633-9. Review.
  • Kim J,Keay SK,Freeman MR.
    Heparin-binding epidermal growth factor-like growth factor functionally antagonizes interstitial cystitis antiproliferative factor via mitogen-activated protein kinase pathway activation.
    BJU Int. 2009 Feb;103(4):541-6.
  • Lee RS,Monigatti F,Briscoe AC,Waldon Z,Freeman MR,Steen H.
    Optimizing sample handling for urinary proteomics.
    J Proteome Res. 2008 Sep;7(9):4022-30.
  • Di Vizio D,Adam RM,Kim J,Kim R,Sotgia F,Williams T,Demichelis F,Solomon KR,Loda M,Rubin MA,Lisanti MP,Freeman MR.
    Caveolin-1 interacts with a lipid raft-associated population of fatty acid synthase.
    Cell Cycle. 2008 Jul 15;7(14):2257-67.
  • Saban MR,Backer JM,Backer MV,Maier J,Fowler B,Davis CA,Simpson C,Wu XR,Birder L,Freeman MR,Soker S,Hurst RE,Saban R.
    VEGF receptors and neuropilins are expressed in the urothelial and neuronal cells in normal mouse urinary bladder and are upregulated in inflammation.
    Am J Physiol Renal Physiol. 2008 Jul;295(1):F60-72.
  • Centeno CJ, Freeman M.
    Letters to the editor.
    Clin Rehabil. 2008 Apr;22(4):377-8.
  • Solomon KR, Freeman MR.
    Do the cholesterol-lowering properties of statins affect cancer risk?.
    Trends Endocrinol Metab. 2008 May-Jun;19(4):113-21. Review.
  • Lee RS, Monigatti F, Lutchman M, Patterson T, Budnik B, Steen JA, Freeman MR, Steen H.
    Temporal variations of the postnatal rat urinary proteome as a reflection of systemic maturation.
    Proteomics. 2008 Mar;8(5):1097-112.
  • Yang W, Steen H, Freeman MR.
    Proteomic approaches to the analysis of multiprotein signaling complexes.
    Proteomics. 2008 Feb;8(4):832-51. Review.
  • Cinar B, Fang PK, Lutchman M, Di Vizio D, Adam RM, Pavlova N, Rubin MA, Yelick PC, Freeman MR.
    The pro-apoptotic kinase Mst1 and its caspase cleavage products are direct inhibitors of Akt1.
    EMBO J. 2007 Oct 31;26(21):4523-34.
  • Dimitrakov J, Kroenke K, Steers WD, Berde C, Zurakowski D, Freeman MR, Jackson JL.
    Pharmacologic Management of Painful Bladder Syndrome/Interstitial Cystitis.
    Arch Intern Med. 2007 Oct 8;167(18):1922-9. Review.
  • Kim J, Jahng WJ, Di Vizio D, Lee JS, Jhaveri R, Rubin MA, Shisheva A, Freeman MR.
    The phosphoinositide kinase PIKfyve mediates epidermal growth factor receptor trafficking to the nucleus.
    Cancer Res. 2007 Oct 1;67(19):9229-37.
  • Di Vizio D, Sotgia F, Williams TM, Hassan GS, Capozza F, Frank PG, Pestell RG, Loda M, Freeman MR, Lisanti MP.
    Caveolin-1 is required for the upregulation of fatty acid synthase (FASN), a tumor promoter, during prostate cancer progression.
    Cancer Biol Ther. 2007 Aug;6(8):1263-8.
  • Cinar B, Mukhopadhyay NK, Meng G, Freeman MR.
    Phosphoinositide 3-kinase-independent non-genomic signals transit from the androgen receptor to Akt1 in membrane raft microdomains.
    J Biol Chem. 2007 Oct 5;282(40):29584-93.
  • Kim J, Keay SK, Dimitrakov JD, Freeman MR.
    p53 mediates interstitial cystitis antiproliferative factor (APF)-induced growth inhibition of human urothelial cells.
    FEBS Lett. 2007 Aug 7;581(20):3795-9.
  • Adam RM, Mukhopadhyay NK, Kim J, Di Vizio D, Cinar B, Boucher K, Solomon KR, Freeman MR.
    Cholesterol sensitivity of endogenous and myristoylated Akt.
    Cancer Res. 2007 Jul 1;67(13):6238-46.
  • Freeman MR.
    Beta2 microglobulin: a surprising therapeutic target for prostate cancer and renal cell carcinoma.
    J Urol. 2007 Jul;178(1):10-1.
  • Mukhopadhyay NK, Cinar B, Mukhopadhyay L, Lutchman M, Ferdinand AS, Kim J, Chung LW, Adam RM, Ray SK, Leiter AB, Richie JP, Liu BC, Freeman MR.
    The zinc finger protein ras-responsive element binding protein-1 is a coregulator of the androgen receptor: implications for the role of the Ras pathway in enhancing androgenic signaling in prostate cancer.
    Mol Endocrinol. 2007 Sep;21(9):2056-70.
  • Freeman MR, Cinar B, Kim J, Mukhopadhyay NK, Di Vizio D, Adam RM, Solomon KR.
    Transit of hormonal and EGF receptor-dependent signals through cholesterol-rich membranes.
    Steroids. 2007 Feb;72(2):210-7. Review.
  • Fang PK, Solomon KR, Zhuang L, Qi M, McKee M, Freeman MR, Yelick PC.
    Caveolin-1alpha and -1beta perform nonredundant roles in early vertebrate development.
    Am J Pathol. 2006 Dec;169(6):2209-22.
  • Duque JL, Loughlin KR, Adam RM, Kantoff P, Mazzucchi E, Freeman MR.
    Measurement of plasma levels of vascular endothelial growth factor in prostate cancer patients: relationship with clinical stage, Gleason score, prostate volume, and serum prostate-specific antigen.
    Clinics. 2006 Oct;61(5):401-8.
  • Estrada CR, Adam RM, Eaton SH, Bägli DJ, Freeman MR.
    Inhibition of EGFR signaling abrogates smooth muscle proliferation resulting from sustained distension of the urinary bladder.
    Lab Invest. 2006 Dec;86(12):1293-302.
  • Mukhopadhyay NK, Ferdinand AS, Mukhopadhyay L, Cinar B, Lutchman M, Richie JP, Freeman MR, Liu BC.
    Unraveling androgen receptor interactomes by an array-based method: discovery of proto-oncoprotein c-Rel as a negative regulator of androgen receptor.
    Exp Cell Res. 2006 Nov 15;312(19):3782-95.
  • Hager MH, Solomon KR, Freeman MR.
    The role of cholesterol in prostate cancer.
    Curr Opin Clin Nutr Metab Care. 2006 Jul;9(4):379-85. Review.
  • Dimitrakov JD, Kaplan SA, Kroenke K, Jackson JL, Freeman MR.
    Management of chronic prostatitis/chronic pelvic pain syndrome: an evidence-based approach.
    Urology. 2006 May;67(5):881-8. Review.
  • Maclellan DL, Steen H, Adam RM, Garlick M, Zurakowski D, Gygi SP, Freeman MR, Solomon KR.
    A quantitative proteomic analysis of growth factor-induced compositional changes in lipid rafts of human smooth muscle cells.
    Proteomics. 2005 Dec;5(18):4733-42.
  • Marcus K, Johnson M, Adam RM, O'Reilly MS, Donovan M, Atala A, Freeman MR, Soker S.
    Tumor cell-associated neuropilin-1 and vascular endothelial growth factor expression as determinants of tumor growth in neuroblastoma.
    Neuropathology. 2005 Sep;25(3):178-87.
  • Kim J, Adam RM, Freeman MR.
    Trafficking of nuclear heparin-binding epidermal growth factor-like growth factor into an epidermal growth factor receptor-dependent autocrine loop in response to oxidative stress.
    Cancer Res. 2005 Sep 15;65(18):8242-9.
  • Freeman MR, Cinar B, Lu ML.
    Membrane rafts as potential sites of nongenomic hormonal signaling in prostate cancer.
    Trends Endocrinol Metab. 2005 Aug;16(6):273-9.
  • Cinar B, De Benedetti A, Freeman MR.
    Post-transcriptional regulation of the androgen receptor by Mammalian target of rapamycin.
    Cancer Res. 2005 Apr 1;65(7):2547-53.
  • Zhuang L, Kim J, Adam RM, Solomon KR, Freeman MR.
    Cholesterol targeting alters lipid raft composition and cell survival in prostate cancer cells and xenografts.
    J Clin Invest. 2005 Apr;115(4):959-68.
  • Orsola A, Estrada CR, Nguyen HT, Retik AB, Freeman MR, Peters CA, Adam RM.
    Growth and stretch response of human exstrophy bladder smooth muscle cells: molecular evidence of normal intrinsic function.
    BJU Int. 2005 Jan;95(1):144-8.
  • Freeman MR.
    HER2/HER3 heterodimers in prostate cancer: Whither HER1/EGFR?.
    Cancer Cell. 2004 Nov;6(5):427-8.
  • Stehr M, Estrada CR, Khoury J, Danciu TE, Sullivan MP, Peters CA, Solomon KR, Freeman MR, Adam RM.
    J Urol. 2004 12;172(6, Part 1 of 2):2451-2455.
  • Kim J, Lin J, Adam RM, Lamb C, Shively SB, Freeman MR.
    An oxidative stress mechanism mediates chelerythrine-induced heparin-binding EGF-like growth factor ectodomain shedding.
    J Cell Biochem. 2005 Jan 1;94(1):39-49.
  • Danciu TE, Gagari E, Adam RM, Damoulis PD, Freeman MR.
    Mechanical strain delivers anti-apoptotic and proliferative signals to gingival fibroblasts.
    J Dent Res. 2004 Aug;83(8):596-601.
  • Cinar B, Yeung F, Konaka H, Mayo MW, Freeman MR, Zhau HE, Chung LW.
    Identification of a negative regulatory cis-element in the enhancer core region of the prostate-specific antigen promoter: implications for intersection of androgen receptor and nuclear factor-kappaB signalling in prostate cancer cells.
    Biochem J. 2004 Apr 15;379(Pt 2):421-31.
  • Freeman MR, Solomon KR.
    Cholesterol and prostate cancer.
    J Cell Biochem. 2004 Jan 1;91(1):54-69.
  • Kim J, Adam RM, Solomon KR, Freeman MR.
    Involvement of cholesterol-rich lipid rafts in interleukin-6-induced neuroendocrine differentiation of LNCaP prostate cancer cells.
    Endocrinology. 2004 Feb;145(2):613-9.
  • Kim J, Freeman MR.
    JNK/SAPK mediates doxorubicin-induced differentiation and apoptosis in MCF-7 breast cancer cells.
    Breast Cancer Res Treat. 2003 Jun;79(3):321-8.
  • Adam RM, Roth JA, Cheng HL, Rice DC, Khoury J, Bauer SB, Peters CA, Freeman MR.
    Signaling through PI3K/Akt mediates stretch and PDGF-BB-dependent DNA synthesis in bladder smooth muscle cells.
    J Urol. 2003 Jun;169(6):2388-93.
  • Adam RM, Danciu T, McLellan DL, Borer JG, Lin J, Zurakowski D, Weinstein MH, Rajjayabun PH, Mellon JK, Freeman MR.
    A nuclear form of the heparin-binding epidermal growth factor-like growth factor precursor is a feature of aggressive transitional cell carcinoma.
    Cancer Res. 2003 Jan 15;63(2):484-90.
  • Lin J, Freeman MR.
    Transactivation of ErbB1 and ErbB2 receptors by angiotensin II in normal human prostate stromal cells.
    Prostate. 2003 Jan 1;54(1):1-7.
  • Zhuang L, Lin J, Lu ML, Solomon KR, Freeman MR.
    Cholesterol-rich lipid rafts mediate akt-regulated survival in prostate cancer cells.
    Cancer Res. 2002 Apr 15;62(8):2227-31.
  • Kim J, Adam RM, Freeman MR.
    Activation of the Erk mitogen-activated protein kinase pathway stimulates neuroendocrine differentiation in LNCaP cells independently of cell cycle withdrawal and STAT3 phosphorylation.
    Cancer Res. 2002 Mar 1;62(5):1549-54.
  • Adam RM, Orsola A, Freeman MR.
    A novel method for implantation of LNCaP prostate tumor cells under the renal capsule.
    In Vitro Cell Dev Biol Anim. 2001 Jun;37(6):360-2.
  • Soker S, Kaefer M, Johnson M, Klagsbrun M, Atala A, Freeman MR.
    Vascular endothelial growth factor-mediated autocrine stimulation of prostate tumor cells coincides with progression to a malignant phenotype.
    Am J Pathol. 2001 Aug;159(2):651-9.
  • Peng JB, Zhuang L, Berger UV, Adam RM, Williams BJ, Brown EM, Hediger MA, Freeman MR.
    CaT1 expression correlates with tumor grade in prostate cancer.
    Biochem Biophys Res Commun. 2001 Apr 6;282(3):729-34.
  • Lin J, Hutchinson L, Gaston SM, Raab G, Freeman MR.
    BAG-1 is a novel cytoplasmic binding partner of the membrane form of heparin-binding EGF-like growth factor: a unique role for proHB-EGF in cell survival regulation.
    J Biol Chem. 2001 Aug 10;276(32):30127-32.
  • Duque JL, Adam RM, Mullen JS, Lin J, Richie JP, Freeman MR.
    Heparin-binding epidermal growth factor-like growth factor is an autocrine mediator of human prostate stromal cell growth in vitro.
    J Urol. 2001 Jan;165(1):284-8.
  • Nguyen HT, Adam RM, Bride SH, Park JM, Peters CA, Freeman MR.
    Cyclic stretch activates p38 SAPK2-, ErbB2-, and AT1-dependent signaling in bladder smooth muscle cells.
    Am J Physiol Cell Physiol. 2000 Oct;279(4):C1155-67.
  • Nguyen HT, Bride SH, Badawy AB, Adam RM, Lin J, Orsola A, Guthrie PD, Freeman MR, Peters CA.
    Heparin-binding EGF-like growth factor is up-regulated in the obstructed kidney in a cell- and region-specific manner and acts to inhibit apoptosis.
    Am J Pathol. 2000 Mar;156(3):889-98.
  • Borer JG, Park JM, Atala A, Nguyen HT, Adam RM, Retik AB, Freeman MR.
    Heparin-binding EGF-like growth factor expression increases selectively in bladder smooth muscle in response to lower urinary tract obstruction.
    Lab Invest. 1999 Nov;79(11):1335-45.
  • Duque JL, Loughlin KR, Adam RM, Kantoff PW, Zurakowski D, Freeman MR.
    Plasma levels of vascular endothelial growth factor are increased in patients with metastatic prostate cancer.
    Urology. 1999 Sep;54(3):523-7.
  • Park JM, Adam RM, Peters CA, Guthrie PD, Sun Z, Klagsbrun M, Freeman MR.
    AP-1 mediates stretch-induced expression of HB-EGF in bladder smooth muscle cells.
    Am J Physiol. 1999 Aug;277(2 Pt 1):C294-301.
  • Lin J, Adam RM, Santiestevan E, Freeman MR.
    The phosphatidylinositol 3'-kinase pathway is a dominant growth factor-activated cell survival pathway in LNCaP human prostate carcinoma cells.
    Cancer Res. 1999 Jun 15;59(12):2891-7.
  • Park JM, Yang T, Arend LJ, Schnermann JB, Peters CA, Freeman MR, Briggs JP.
    Obstruction stimulates COX-2 expression in bladder smooth muscle cells via increased mechanical stretch.
    Am J Physiol. 1999 Jan;276(1 Pt 2):F129-36.
  • Freeman MR, Yoo JJ, Raab G, Soker S, Adam RM, Schneck FX, Renshaw AA, Klagsbrun M, Atala A.
    Heparin-binding EGF-like growth factor is an autocrine growth factor for human urothelial cells and is synthesized by epithelial and smooth muscle cells in the human bladder.
    J Clin Invest. 1997 Mar 1;99(5):1028-36.