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    Zea Borok, MD

    TitleProfessor of Medicine
    SchoolKeck School of Medicine of USC
    DepartmentMedicine
    AddressIRD 720 2020 Zonal Avenue
    Off Campus
    Los Angeles California 90033
    Phone+1 323 226 7923
    vCardDownload vCard
      Other Positions
      TitleRalph Edgington Chair in Medicine

      TitleChief, Division of Pulmonary and Critical Care Medicine


      Collapse Overview 
      Collapse Overview
      Dr. Borok's clinical interests include acute lung injury and intensive care. Her research interests include transcriptional and epigenetic regulation of alveolar epithelial cell differentiation/plasticity, modulation of alveolar epithelial cell phenotype and recovery following lung injury, role of alveolar epithelium in pulmonary fibrosis, biology of lung stem cells and alveolar epithelial tight junction regulation.

      After completing medical school in her native South Africa, Dr. Borok completed her residency in Internal Medicine at the University of Pittsburgh. Following a fellowship in Critical Care Medicine at the University of Pittsburgh, she completed fellowship training in Pulmonary Medicine at the National Institutes of Health. Dr. Borok joined the faculty of USC in 1990 and quickly distinguished herself as a prominent medical researcher, educator and practitioner. She holds several administrative and academic positions and directs an active research program in mechanisms of lung injury repair.


      Collapse Biography 
      Collapse Awards and Honors
      American Thoracic Society 2017Elizabeth Rich Award
      Western Society for Clinical Investigation2017Mayo Soley Award
      American Thoracic Society2014Recognition Award for Scientific Accomplishment
      Association for American Physicians2014Elected to Association for American Physicians
      NHLBI2010Method to Extend Research in Time (MERIT) Award
      Western Society for Clinical Investigation2007Outstanding Investigator Award
      NIH1993Clinical Investigator Development Award

      Collapse Bibliographic 
      Collapse Publications
      Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Researchers can login to make corrections and additions, or contact us for help.
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      1. Lax S, Rayes J, Wichaiyo S, Haining EJ, Lowe K, Grygielska B, Laloo R, Flodby P, Borok Z, Crandall ED, Thickett D, Watson SP. Platelet CLEC-2 protects against lung injury via effects of its ligand podoplanin on inflammatory alveolar macrophages in the mouse. Am J Physiol Lung Cell Mol Physiol. 2017 Aug 24; ajplung.00023.2017. PMID: 28839100.
        View in: PubMed
      2. Gukasyan HJ, Uchiyama T, Kim KJ, Ehrhardt C, Wu SK, Borok Z, Crandall ED, Lee VHL. Oligopeptide Transport in Rat Lung Alveolar Epithelial Cells is Mediated by Pept2. Pharm Res. 2017 Aug 22. PMID: 28831683.
        View in: PubMed
      3. Stueve TR, Li WQ, Shi J, Marconett CN, Zhang T, Yang C, Mullen D, Yan C, Wheeler W, Hua X, Zhou B, Borok Z, Caporaso NE, Pesatori AC, Duan J, Laird-Offringa IA, Landi MT. Epigenome-wide analysis of DNA methylation in lung tissue shows concordance with blood studies and identifies tobacco smoke-inducible enhancers. Hum Mol Genet. 2017 Aug 01; 26(15):3014-3027. PMID: 28854564.
        View in: PubMed
      4. Horng S, Therattil A, Moyon S, Gordon A, Kim K, Argaw AT, Hara Y, Mariani JN, Sawai S, Flodby P, Crandall ED, Borok Z, Sofroniew MV, Chapouly C, John GR. Astrocytic tight junctions control inflammatory CNS lesion pathogenesis. J Clin Invest. 2017 Aug 01; 127(8):3136-3151. PMID: 28737509.
        View in: PubMed
      5. Debebe A, Medina V, Chen CY, Mahajan IM, Jia C, Fu D, He L, Zeng N, Stiles BW, Chen CL, Wang M, Aggarwal KR, Peng Z, Huang J, Chen J, Li M, Dong T, Atkins S, Borok Z, Yuan W, Machida K, Ju C, Kahn M, Johnson D, Stiles BL. Wnt/ß-catenin activation and macrophage induction during liver cancer development following steatosis. Oncogene. 2017 Jul 03. PMID: 28671671.
        View in: PubMed
      6. Flodby P, Li C, Liu Y, Wang H, Rieger ME, Minoo P, Crandall ED, Ann DK, Borok Z, Zhou B. Cell-specific expression of aquaporin-5 (Aqp5) in alveolar epithelium is directed by GATA6/Sp1 via histone acetylation. Sci Rep. 2017 Jun 14; 7(1):3473. PMID: 28615712.
        View in: PubMed
      7. Marconett CN, Zhou B, Sunohara M, Pouldar TM, Wang H, Liu Y, Rieger ME, Tran E, Flodby P, Siegmund KD, Crandall ED, Laird-Offringa IA, Borok Z. Cross-Species Transcriptome Profiling Identifies New Alveolar Epithelial Type I Cell-Specific Genes. Am J Respir Cell Mol Biol. 2017 Mar; 56(3):310-321. PMID: 27749084.
        View in: PubMed
      8. Flodby P, Liebler JM, Sunohara M, Castillo DR, McConnell AM, Krishnaveni MS, Banfalvi A, Li M, Stripp B, Zhou B, Crandall ED, Minoo P, Borok Z. Region-specific role for Pten in maintenance of epithelial phenotype and integrity. Am J Physiol Lung Cell Mol Physiol. 2017 Jan 01; 312(1):L131-L142. PMID: 27864284.
        View in: PubMed
      9. Agoro R, Piotet-Morin J, Palomo J, Michaudel C, Vigne S, Maillet I, Chenuet P, Guillou N, Le Bérichel J, Kisielow M, Flodby P, Borok Z, Crandall ED, Le Bert M, Quesniaux V, Muller M, Di Padova F, Ryffel B, Gabay C, Couturier-Maillard A. IL-1R1-MyD88 axis elicits papain-induced lung inflammation. Eur J Immunol. 2016 Nov; 46(11):2531-2541. PMID: 27569535.
        View in: PubMed
      10. Flodby P, Kim YH, Beard LL, Gao D, Ji Y, Kage H, Liebler JM, Minoo P, Kim KJ, Borok Z, Crandall ED. Knockout Mice Reveal a Major Role for Alveolar Epithelial Type I Cells in Alveolar Fluid Clearance. Am J Respir Cell Mol Biol. 2016 Sep; 55(3):395-406. PMID: 27064541; PMCID: PMC5023030 [Available on 09/01/17].
      11. Zuber V, Marconett CN, Shi J, Hua X, Wheeler W, Yang C, Song L, Dale AM, Laplana M, Risch A, Witoelar A, Thompson WK, Schork AJ, Bettella F, Wang Y, Djurovic S, Zhou B, Borok Z, van der Heijden HF, de Graaf J, Swinkels D, Aben KK, McKay J, Hung RJ, Bikeböller H, Stevens VL, Albanes D, Caporaso NE, Han Y, Wei Y, Panadero MA, Mayordomo JI, Christiani DC, Kiemeney L, Andreassen OA, Houlston R, Amos CI, Chatterjee N, Laird-Offringa IA, Mills IG, Landi MT. Pleiotropic Analysis of Lung Cancer and Blood Triglycerides. J Natl Cancer Inst. 2016 Dec; 108(12). PMID: 27565901.
        View in: PubMed
      12. Stueve TR, Marconett CN, Zhou B, Borok Z, Laird-Offringa IA. The importance of detailed epigenomic profiling of different cell types within organs. Epigenomics. 2016 Jun; 8(6):817-29. PMID: 27305639.
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      13. White ES, Borok Z, Brown KK, Eickelberg O, Guenther A, Jenkins RG, Kolb M, Martinez FJ, Roman J, Sime P. An American Thoracic Society Official Research Statement: Future Directions in Lung Fibrosis Research. Am J Respir Crit Care Med. 2016 Apr 01; 193(7):792-800. PMID: 27035782.
        View in: PubMed
      14. Li A, Ma S, Smith SM, Lee MK, Fischer A, Borok Z, Bellusci S, Li C, Minoo P. Mesodermal ALK5 controls lung myofibroblast versus lipofibroblast cell fate. BMC Biol. 2016 Mar 16; 14:19. PMID: 26984772; PMCID: PMC4793501.
      15. Rieger ME, Zhou B, Solomon N, Sunohara M, Li C, Nguyen C, Liu Y, Pan JH, Minoo P, Crandall ED, Brody SL, Kahn M, Borok Z. p300/ß-Catenin Interactions Regulate Adult Progenitor Cell Differentiation Downstream of WNT5a/Protein Kinase C (PKC). J Biol Chem. 2016 Mar 18; 291(12):6569-82. PMID: 26833564; PMCID: PMC4813575 [Available on 03/18/17].
      16. Smith KA, Zhou B, Avdulov S, Benyumov A, Peterson M, Liu Y, Okon A, Hergert P, Braziunas J, Wagner CR, Borok Z, Bitterman PB. Transforming Growth Factor-ß1 Induced Epithelial Mesenchymal Transition is blocked by a chemical antagonist of translation factor eIF4E. Sci Rep. 2015 Dec 18; 5:18233. PMID: 26678431; PMCID: PMC4683370.
      17. Li C, Bellusci S, Borok Z, Minoo P. Non-canonical WNT signalling in the lung. J Biochem. 2015 Nov; 158(5):355-65. PMID: 26261051; PMCID: PMC4751232 [Available on 11/01/16].
      18. Li C, Li M, Li S, Xing Y, Yang CY, Li A, Borok Z, De Langhe S, Minoo P. Progenitors of secondary crest myofibroblasts are developmentally committed in early lung mesoderm. Stem Cells. 2015 Mar; 33(3):999-1012. PMID: 25448080; PMCID: PMC4650896.
      19. Catalán MA, Kondo Y, Peña-Munzenmayer G, Jaramillo Y, Liu F, Choi S, Crandall E, Borok Z, Flodby P, Shull GE, Melvin JE. A fluid secretion pathway unmasked by acinar-specific Tmem16A gene ablation in the adult mouse salivary gland. Proc Natl Acad Sci U S A. 2015 Feb 17; 112(7):2263-8. PMID: 25646474; PMCID: PMC4343136.
      20. Marconett CN, Zhou B, Siegmund KD, Borok Z, Laird-Offringa IA. Transcriptomic Profiling of Primary Alveolar Epithelial Cell Differentiation in Human and Rat. Genom Data. 2014 Dec 01; 2:105-109. PMID: 25343132.
        View in: PubMed
      21. Lin HH, Lin SM, Chung Y, Vonderfecht S, Camden JM, Flodby P, Borok Z, Limesand KH, Mizushima N, Ann DK. Dynamic involvement of ATG5 in cellular stress responses. Cell Death Dis. 2014 Oct 23; 5:e1478. PMID: 25341032; PMCID: PMC4649523.
      22. Kage H, Flodby P, Gao D, Kim YH, Marconett CN, DeMaio L, Kim KJ, Crandall ED, Borok Z. Claudin 4 knockout mice: normal physiological phenotype with increased susceptibility to lung injury. Am J Physiol Lung Cell Mol Physiol. 2014 Oct 01; 307(7):L524-36. PMID: 25106430; PMCID: PMC4187039.
      23. Li G, Flodby P, Luo J, Kage H, Sipos A, Gao D, Ji Y, Beard LL, Marconett CN, DeMaio L, Kim YH, Kim KJ, Laird-Offringa IA, Minoo P, Liebler JM, Zhou B, Crandall ED, Borok Z. Knockout mice reveal key roles for claudin 18 in alveolar barrier properties and fluid homeostasis. Am J Respir Cell Mol Biol. 2014 Aug; 51(2):210-22. PMID: 24588076; PMCID: PMC4148039.
      24. Borok Z. Alveolar epithelium: beyond the barrier. Am J Respir Cell Mol Biol. 2014 May; 50(5):853-6. PMID: 24783957; PMCID: PMC4068954.
      25. Shi J, Marconett CN, Duan J, Hyland PL, Li P, Wang Z, Wheeler W, Zhou B, Campan M, Lee DS, Huang J, Zhou W, Triche T, Amundadottir L, Warner A, Hutchinson A, Chen PH, Chung BS, Pesatori AC, Consonni D, Bertazzi PA, Bergen AW, Freedman M, Siegmund KD, Berman BP, Borok Z, Chatterjee N, Tucker MA, Caporaso NE, Chanock SJ, Laird-Offringa IA, Landi MT. Characterizing the genetic basis of methylome diversity in histologically normal human lung tissue. Nat Commun. 2014 Feb 27; 5:3365. PMID: 24572595; PMCID: PMC3982882.
      26. Blackwell TS, Tager AM, Borok Z, Moore BB, Schwartz DA, Anstrom KJ, Bar-Joseph Z, Bitterman P, Blackburn MR, Bradford W, Brown KK, Chapman HA, Collard HR, Cosgrove GP, Deterding R, Doyle R, Flaherty KR, Garcia CK, Hagood JS, Henke CA, Herzog E, Hogaboam CM, Horowitz JC, King TE, Loyd JE, Lawson WE, Marsh CB, Noble PW, Noth I, Sheppard D, Olsson J, Ortiz LA, O'Riordan TG, Oury TD, Raghu G, Roman J, Sime PJ, Sisson TH, Tschumperlin D, Violette SM, Weaver TE, Wells RG, White ES, Kaminski N, Martinez FJ, Wynn TA, Thannickal VJ, Eu JP. Future directions in idiopathic pulmonary fibrosis research. An NHLBI workshop report. Am J Respir Crit Care Med. 2014 Jan 15; 189(2):214-22. PMID: 24160862; PMCID: PMC3983890.
      27. Morgan-Bathke M, Lin HH, Chibly AM, Zhang W, Sun X, Chen CH, Flodby P, Borok Z, Wu R, Arnett D, Klein RR, Ann DK, Limesand KH. Deletion of ATG5 shows a role of autophagy in salivary homeostatic control. J Dent Res. 2013 Oct; 92(10):911-7. PMID: 23884556; PMCID: PMC3775371.
      28. Marconett CN, Zhou B, Rieger ME, Selamat SA, Dubourd M, Fang X, Lynch SK, Stueve TR, Siegmund KD, Berman BP, Borok Z, Laird-Offringa IA. Integrated transcriptomic and epigenomic analysis of primary human lung epithelial cell differentiation. PLoS Genet. 2013 Jun; 9(6):e1003513. PMID: 23818859; PMCID: PMC3688557.
      29. Li A, Chan B, Felix JC, Xing Y, Li M, Brody SL, Borok Z, Li C, Minoo P. Tissue-dependent consequences of Apc inactivation on proliferation and differentiation of ciliated cell progenitors via Wnt and notch signaling. PLoS One. 2013; 8(4):e62215. PMID: 23646120; PMCID: PMC3639955.
      30. Li C, Li A, Xing Y, Li M, Chan B, Ouyang R, Taketo MM, Kucherlapati R, Borok Z, Minoo P. Apc deficiency alters pulmonary epithelial cell fate and inhibits Nkx2.1 via triggering TGF-beta signaling. Dev Biol. 2013 Jun 01; 378(1):13-24. PMID: 23562608; PMCID: PMC3838671.
      31. Fazlollahi F, Kim YH, Sipos A, Hamm-Alvarez SF, Borok Z, Kim KJ, Crandall ED. Nanoparticle translocation across mouse alveolar epithelial cell monolayers: species-specific mechanisms. Nanomedicine. 2013 Aug; 9(6):786-94. PMID: 23454523; PMCID: PMC3702651.
      32. Wu H, Chen L, Zhang X, Zhou Q, Li JM, Berger S, Borok Z, Zhou B, Xiao Z, Yin H, Liu M, Wang Y, Jin J, Blackburn MR, Xia Y, Zhang W. Aqp5 is a new transcriptional target of Dot1a and a regulator of Aqp2. PLoS One. 2013; 8(1):e53342. PMID: 23326416; PMCID: PMC3542343.
      33. Tiozzo C, Carraro G, Al Alam D, Baptista S, Danopoulos S, Li A, Lavarreda-Pearce M, Li C, De Langhe S, Chan B, Borok Z, Bellusci S, Minoo P. Mesodermal Pten inactivation leads to alveolar capillary dysplasia- like phenotype. J Clin Invest. 2012 Nov; 122(11):3862-72. PMID: 23023706; PMCID: PMC3484434.
      34. Kage H, Borok Z. EMT and interstitial lung disease: a mysterious relationship. Curr Opin Pulm Med. 2012 Sep; 18(5):517-23. PMID: 22854509; PMCID: PMC3914631.
      35. Zhou B, Buckley ST, Patel V, Liu Y, Luo J, Krishnaveni MS, Ivan M, DeMaio L, Kim KJ, Ehrhardt C, Crandall ED, Borok Z. Troglitazone attenuates TGF-ß1-induced EMT in alveolar epithelial cells via a PPAR?-independent mechanism. PLoS One. 2012; 7(6):e38827. PMID: 22745681; PMCID: PMC3380041.
      36. Xing Y, Li A, Borok Z, Li C, Minoo P. NOTCH1 is required for regeneration of Clara cells during repair of airway injury. Stem Cells. 2012 May; 30(5):946-55. PMID: 22331706; PMCID: PMC4005608.
      37. Zhang H, Liu H, Borok Z, Davies KJ, Ursini F, Forman HJ. Cigarette smoke extract stimulates epithelial-mesenchymal transition through Src activation. Free Radic Biol Med. 2012 Apr 15; 52(8):1437-42. PMID: 22342303; PMCID: PMC3312989.
      38. Zhou B, Liu Y, Kahn M, Ann DK, Han A, Wang H, Nguyen C, Flodby P, Zhong Q, Krishnaveni MS, Liebler JM, Minoo P, Crandall ED, Borok Z. Interactions between ß-catenin and transforming growth factor-ß signaling pathways mediate epithelial-mesenchymal transition and are dependent on the transcriptional co-activator cAMP-response element-binding protein (CREB)-binding protein (CBP). J Biol Chem. 2012 Mar 02; 287(10):7026-38. PMID: 22241478; PMCID: PMC3293544.
      39. Chimge NO, Baniwal SK, Little GH, Chen YB, Kahn M, Tripathy D, Borok Z, Frenkel B. Regulation of breast cancer metastasis by Runx2 and estrogen signaling: the role of SNAI2. Breast Cancer Res. 2011; 13(6):R127. PMID: 22151997; PMCID: PMC3326569.
      40. DeMaio L, Buckley ST, Krishnaveni MS, Flodby P, Dubourd M, Banfalvi A, Xing Y, Ehrhardt C, Minoo P, Zhou B, Crandall ED, Borok Z. Ligand-independent transforming growth factor-ß type I receptor signalling mediates type I collagen-induced epithelial-mesenchymal transition. J Pathol. 2012 Mar; 226(4):633-44. PMID: 21984393; PMCID: PMC4012689.
      41. Fazlollahi F, Sipos A, Kim YH, Hamm-Alvarez SF, Borok Z, Kim KJ, Crandall ED. Translocation of PEGylated quantum dots across rat alveolar epithelial cell monolayers. Int J Nanomedicine. 2011; 6:2849-57. PMID: 22131830; PMCID: PMC3224711.
      42. Jenkins G, Blanchard A, Borok Z, Bradding P, Ehrhardt C, Fisher A, Hirani N, Johnson S, Königshoff M, Maher TM, Millar A, Parfrey H, Scotton C, Tetley T, Thickett D, Wolters P. In search of the fibrotic epithelial cell: opportunities for a collaborative network. Thorax. 2012 Feb; 67(2):179-82. PMID: 21680569.
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      43. Borok Z, Whitsett JA, Bitterman PB, Thannickal VJ, Kotton DN, Reynolds SD, Krasnow MA, Bianchi DW, Morrisey EE, Hogan BL, Kurie JM, Walker DC, Radisky DC, Nishimura SL, Violette SM, Noble PW, Shapiro SD, Blaisdell CJ, Chapman HA, Kiley J, Gail D, Hoshizaki D. Cell plasticity in lung injury and repair: report from an NHLBI workshop, April 19-20, 2010. Proc Am Thorac Soc. 2011 Jun; 8(3):215-22. PMID: 21653526; PMCID: PMC3132783.
      44. Weiss DJ, Bertoncello I, Borok Z, Kim C, Panoskaltsis-Mortari A, Reynolds S, Rojas M, Stripp B, Warburton D, Prockop DJ. Stem cells and cell therapies in lung biology and lung diseases. Proc Am Thorac Soc. 2011 Jun; 8(3):223-72. PMID: 21653527; PMCID: PMC3132784.
      45. Yacobi NR, Fazllolahi F, Kim YH, Sipos A, Borok Z, Kim KJ, Crandall ED. Nanomaterial interactions with and trafficking across the lung alveolar epithelial barrier: implications for health effects of air-pollution particles. Air Qual Atmos Health. 2011 Mar 01; 4(1):65-78. PMID: 25568662.
        View in: PubMed
      46. Fazlollahi F, Angelow S, Yacobi NR, Marchelletta R, Yu AS, Hamm-Alvarez SF, Borok Z, Kim KJ, Crandall ED. Polystyrene nanoparticle trafficking across MDCK-II. Nanomedicine. 2011 Oct; 7(5):588-94. PMID: 21310266; PMCID: PMC3130091.
      47. Zhong Q, Zhou B, Ann DK, Minoo P, Liu Y, Banfalvi A, Krishnaveni MS, Dubourd M, Demaio L, Willis BC, Kim KJ, duBois RM, Crandall ED, Beers MF, Borok Z. Role of endoplasmic reticulum stress in epithelial-mesenchymal transition of alveolar epithelial cells: effects of misfolded surfactant protein. Am J Respir Cell Mol Biol. 2011 Sep; 45(3):498-509. PMID: 21169555; PMCID: PMC3175581.
      48. Li M, Krishnaveni MS, Li C, Zhou B, Xing Y, Banfalvi A, Li A, Lombardi V, Akbari O, Borok Z, Minoo P. Epithelium-specific deletion of TGF-ß receptor type II protects mice from bleomycin-induced pulmonary fibrosis. J Clin Invest. 2011 Jan; 121(1):277-87. PMID: 21135509; PMCID: PMC3007138.
      49. Li A, Xing Y, Chan B, Heisterkamp N, Groffen J, Borok Z, Minoo P, Li C. Cell type-specific expression of adenomatous polyposis coli in lung development, injury, and repair. Dev Dyn. 2010 Aug; 239(8):2288-97. PMID: 20658693; PMCID: PMC2911649.
      50. Henderson WR, Chi EY, Ye X, Nguyen C, Tien YT, Zhou B, Borok Z, Knight DA, Kahn M. Inhibition of Wnt/beta-catenin/CREB binding protein (CBP) signaling reverses pulmonary fibrosis. Proc Natl Acad Sci U S A. 2010 Aug 10; 107(32):14309-14. PMID: 20660310; PMCID: PMC2922550.
      51. Kim YH, Fazlollahi F, Kennedy IM, Yacobi NR, Hamm-Alvarez SF, Borok Z, Kim KJ, Crandall ED. Alveolar epithelial cell injury due to zinc oxide nanoparticle exposure. Am J Respir Crit Care Med. 2010 Dec 01; 182(11):1398-409. PMID: 20639441; PMCID: PMC3029930.
      52. Xing Y, Li C, Li A, Sridurongrit S, Tiozzo C, Bellusci S, Borok Z, Kaartinen V, Minoo P. Signaling via Alk5 controls the ontogeny of lung Clara cells. Development. 2010 Mar; 137(5):825-33. PMID: 20147383; PMCID: PMC2827691.
      53. Borok Z, Crandall ED. More life for a "terminal" cell. Am J Physiol Lung Cell Mol Physiol. 2009 Dec; 297(6):L1042-4. PMID: 19837848; PMCID: PMC2793181.
      54. Flodby P, Borok Z, Banfalvi A, Zhou B, Gao D, Minoo P, Ann DK, Morrisey EE, Crandall ED. Directed expression of Cre in alveolar epithelial type 1 cells. Am J Respir Cell Mol Biol. 2010 Aug; 43(2):173-8. PMID: 19767448; PMCID: PMC2937230.
      55. Willis BC, Borok Z. Epithelial-mesenchymal transition: potential role in obliterative bronchiolitis? Thorax. 2009 Sep; 64(9):742-3. PMID: 19717708; PMCID: PMC4172368.
      56. Felton VM, Borok Z, Willis BC. N-acetylcysteine inhibits alveolar epithelial-mesenchymal transition. Am J Physiol Lung Cell Mol Physiol. 2009 Nov; 297(5):L805-12. PMID: 19648289; PMCID: PMC2777496.
      57. Tiozzo C, De Langhe S, Yu M, Londhe VA, Carraro G, Li M, Li C, Xing Y, Anderson S, Borok Z, Bellusci S, Minoo P. Deletion of Pten expands lung epithelial progenitor pools and confers resistance to airway injury. Am J Respir Crit Care Med. 2009 Oct 15; 180(8):701-12. PMID: 19574443; PMCID: PMC2778148.
      58. Yacobi NR, Malmstadt N, Fazlollahi F, DeMaio L, Marchelletta R, Hamm-Alvarez SF, Borok Z, Kim KJ, Crandall ED. Mechanisms of alveolar epithelial translocation of a defined population of nanoparticles. Am J Respir Cell Mol Biol. 2010 May; 42(5):604-14. PMID: 19574531; PMCID: PMC2874446.
      59. Demaio L, Tseng W, Balverde Z, Alvarez JR, Kim KJ, Kelley DG, Senior RM, Crandall ED, Borok Z. Characterization of mouse alveolar epithelial cell monolayers. Am J Physiol Lung Cell Mol Physiol. 2009 Jun; 296(6):L1051-8. PMID: 19329539; PMCID: PMC2692796.
      60. Patel LN, Wang J, Kim KJ, Borok Z, Crandall ED, Shen WC. Conjugation with cationic cell-penetrating peptide increases pulmonary absorption of insulin. Mol Pharm. 2009 Mar-Apr; 6(2):492-503. PMID: 19228019; PMCID: PMC2798810.
      61. Borok Z. Role for alpha3 integrin in EMT and pulmonary fibrosis. J Clin Invest. 2009 Jan; 119(1):7-10. PMID: 19104143; PMCID: PMC2613449.
      62. Li M, Li C, Liu YH, Xing Y, Hu L, Borok Z, Kwong KY, Minoo P. Mesodermal deletion of transforming growth factor-beta receptor II disrupts lung epithelial morphogenesis: cross-talk between TGF-beta and Sonic hedgehog pathways. J Biol Chem. 2008 Dec 26; 283(52):36257-64. PMID: 18990706; PMCID: PMC2605990.
      63. Pregizer S, Baniwal SK, Yan X, Borok Z, Frenkel B. Progressive recruitment of Runx2 to genomic targets despite decreasing expression during osteoblast differentiation. J Cell Biochem. 2008 Nov 01; 105(4):965-70. PMID: 18821584; PMCID: PMC2591066.
      64. Zhou B, Francis TA, Yang H, Tseng W, Zhong Q, Frenkel B, Morrisey EE, Ann DK, Minoo P, Crandall ED, Borok Z. GATA-6 mediates transcriptional activation of aquaporin-5 through interactions with Sp1. Am J Physiol Cell Physiol. 2008 Nov; 295(5):C1141-50. PMID: 18768929; PMCID: PMC2584994.
      65. Patel LN, Uchiyama T, Kim KJ, Borok Z, Crandall ED, Shen WC, Lee VH. Molecular and functional expression of multidrug resistance-associated protein-1 in primary cultured rat alveolar epithelial cells. J Pharm Sci. 2008 Jun; 97(6):2340-9. PMID: 17854063.
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      66. Liebler JM, Lutzko C, Banfalvi A, Senadheera D, Aghamohammadi N, Crandall ED, Borok Z. Retention of human bone marrow-derived cells in murine lungs following bleomycin-induced lung injury. Am J Physiol Lung Cell Mol Physiol. 2008 Aug; 295(2):L285-92. PMID: 18515407; PMCID: PMC2519849.
      67. Zhou B, Ann DK, Flodby P, Minoo P, Liebler JM, Crandall ED, Borok Z. Rat aquaporin-5 4.3-kb 5'-flanking region differentially regulates expression in salivary gland and lung in vivo. Am J Physiol Cell Physiol. 2008 Jul; 295(1):C111-20. PMID: 18448628; PMCID: PMC2493550.
      68. Yacobi NR, Demaio L, Xie J, Hamm-Alvarez SF, Borok Z, Kim KJ, Crandall ED. Polystyrene nanoparticle trafficking across alveolar epithelium. Nanomedicine. 2008 Jun; 4(2):139-45. PMID: 18375191.
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      69. Uchiyama T, Fujita T, Gukasyan HJ, Kim KJ, Borok Z, Crandall ED, Lee VH. Functional characterization and cloning of amino acid transporter B(0,+) (ATB(0,+)) in primary cultured rat pneumocytes. J Cell Physiol. 2008 Mar; 214(3):645-54. PMID: 17960566.
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      70. Bahhady R, Kim KJ, Borok Z, Crandall ED, Shen WC. Characterization of protein factor(s) in rat bronchoalveolar lavage fluid that enhance insulin transport via transcytosis across primary rat alveolar epithelial cell monolayers. Eur J Pharm Biopharm. 2008 Aug; 69(3):808-16. PMID: 18406118; PMCID: PMC3638983.
      71. Zhou B, Zhong Q, Minoo P, Li C, Ann DK, Frenkel B, Morrisey EE, Crandall ED, Borok Z. Foxp2 inhibits Nkx2.1-mediated transcription of SP-C via interactions with the Nkx2.1 homeodomain. Am J Respir Cell Mol Biol. 2008 Jun; 38(6):750-8. PMID: 18239190; PMCID: PMC2396252.
      72. Minoo P, Hu L, Zhu N, Borok Z, Bellusci S, Groffen J, Kardassis D, Li C. SMAD3 prevents binding of NKX2.1 and FOXA1 to the SpB promoter through its MH1 and MH2 domains. Nucleic Acids Res. 2008 Jan; 36(1):179-88. PMID: 18003659; PMCID: PMC2248754.
      73. Bates SR, Tao JQ, Yu KJ, Borok Z, Crandall ED, Collins HL, Rothblat GH. Expression and biological activity of ABCA1 in alveolar epithelial cells. Am J Respir Cell Mol Biol. 2008 Mar; 38(3):283-92. PMID: 17884990; PMCID: PMC2258448.
      74. Qiao R, Yan W, Clavijo C, Mehrian-Shai R, Zhong Q, Kim KJ, Ann D, Crandall ED, Borok Z. Effects of KGF on alveolar epithelial cell transdifferentiation are mediated by JNK signaling. Am J Respir Cell Mol Biol. 2008 Feb; 38(2):239-46. PMID: 17872496; PMCID: PMC2214671.
      75. Willis BC, Borok Z. TGF-beta-induced EMT: mechanisms and implications for fibrotic lung disease. Am J Physiol Lung Cell Mol Physiol. 2007 Sep; 293(3):L525-34. PMID: 17631612.
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      76. Yacobi NR, Phuleria HC, Demaio L, Liang CH, Peng CA, Sioutas C, Borok Z, Kim KJ, Crandall ED. Nanoparticle effects on rat alveolar epithelial cell monolayer barrier properties. Toxicol In Vitro. 2007 Dec; 21(8):1373-81. PMID: 17555923; PMCID: PMC3855017.
      77. Bahhady R, Kim KJ, Borok Z, Crandall ED, Shen WC. Enhancement of insulin transport across primary rat alveolar epithelial cell monolayers by endogenous cellular factor(s). Pharm Res. 2007 Sep; 24(9):1713-9. PMID: 17443400.
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      78. Jain R, Shaul PW, Borok Z, Willis BC. Endothelin-1 induces alveolar epithelial-mesenchymal transition through endothelin type A receptor-mediated production of TGF-beta1. Am J Respir Cell Mol Biol. 2007 Jul; 37(1):38-47. PMID: 17379848; PMCID: PMC1899351.
      79. Flodby P, Zhou B, Ann DK, Kim KJ, Minoo P, Crandall ED, Borok Z. Conserved elements within first intron of aquaporin-5 (Aqp5) function as transcriptional enhancers. Biochem Biophys Res Commun. 2007 Apr 27; 356(1):26-31. PMID: 17339032; PMCID: PMC2366028.
      80. Lin HH, Xiong Y, Ho YS, Zhou B, Nguyen HV, Deng H, Lee R, Yen Y, Borok Z, Ann DK. Transcriptional regulation by targeted expression of architectural transcription factor high mobility group A2 in salivary glands of transgenic mice. Eur J Oral Sci. 2007 Feb; 115(1):30-9. PMID: 17305714.
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      81. Minoo P, Hu L, Xing Y, Zhu NL, Chen H, Li M, Borok Z, Li C. Physical and functional interactions between homeodomain NKX2.1 and winged helix/forkhead FOXA1 in lung epithelial cells. Mol Cell Biol. 2007 Mar; 27(6):2155-65. PMID: 17220277; PMCID: PMC1820505.
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      83. Willis BC, duBois RM, Borok Z. Epithelial origin of myofibroblasts during fibrosis in the lung. Proc Am Thorac Soc. 2006 Jun; 3(4):377-82. PMID: 16738204; PMCID: PMC2658689.
      84. Weiss DJ, Berberich MA, Borok Z, Gail DB, Kolls JK, Penland C, Prockop DJ. Adult stem cells, lung biology, and lung disease. NHLBI/Cystic Fibrosis Foundation Workshop. Proc Am Thorac Soc. 2006 May; 3(3):193-207. PMID: 16636086.
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      85. Nguyen HV, Chen JL, Zhong J, Kim KJ, Crandall ED, Borok Z, Chen Y, Ann DK. SUMOylation attenuates sensitivity toward hypoxia- or desferroxamine-induced injury by modulating adaptive responses in salivary epithelial cells. Am J Pathol. 2006 May; 168(5):1452-63. PMID: 16651613; PMCID: PMC1606580.
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      89. Chen SP, Zhou B, Willis BC, Sandoval AJ, Liebler JM, Kim KJ, Ann DK, Crandall ED, Borok Z. Effects of transdifferentiation and EGF on claudin isoform expression in alveolar epithelial cells. J Appl Physiol (1985). 2005 Jan; 98(1):322-8. PMID: 15361518.
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      90. Kim KJ, Borok Z, Ehrhardt C, Willis BC, Lehr CM, Crandall ED. Estimation of paracellular conductance of primary rat alveolar epithelial cell monolayers. J Appl Physiol (1985). 2005 Jan; 98(1):138-43. PMID: 15273240.
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      91. Liebler JM, Borok Z, Li X, Zhou B, Sandoval AJ, Kim KJ, Crandall ED. Alveolar epithelial type I cells express beta2-adrenergic receptors and G-protein receptor kinase 2. J Histochem Cytochem. 2004 Jun; 52(6):759-67. PMID: 15150284.
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      92. Kim KJ, Fandy TE, Lee VH, Ann DK, Borok Z, Crandall ED. Net absorption of IgG via FcRn-mediated transcytosis across rat alveolar epithelial cell monolayers. Am J Physiol Lung Cell Mol Physiol. 2004 Sep; 287(3):L616-22. PMID: 15169676.
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      93. Qiao R, Zhou B, Harboe-Schmidt E, Kasahara N, Kim KJ, Liebler JM, Crandall ED, Borok Z. Subunit-specific coordinate upregulation of sodium pump activity in alveolar epithelial cells by lentivirus-mediated gene transfer. Hum Gene Ther. 2004 May; 15(5):457-68. PMID: 15144576.
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      94. Blumenthal M, Kardosh A, Dubeau L, Borok Z, Schönthal AH. Suppression of the transformed phenotype and induction of differentiation-like characteristics in cultured ovarian tumor cells by chronic treatment with progesterone. Mol Carcinog. 2003 Dec; 38(4):160-9. PMID: 14639655.
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      95. Xiao J, Li C, Zhu NL, Borok Z, Minoo P. Timeless in lung morphogenesis. Dev Dyn. 2003 Sep; 228(1):82-94. PMID: 12950082.
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      96. You Y, Huang T, Richer EJ, Schmidt JE, Zabner J, Borok Z, Brody SL. Role of f-box factor foxj1 in differentiation of ciliated airway epithelial cells. Am J Physiol Lung Cell Mol Physiol. 2004 Apr; 286(4):L650-7. PMID: 12818891.
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      97. Willis BC, Kim KJ, Li X, Liebler J, Crandall ED, Borok Z. Modulation of ion conductance and active transport by TGF-beta 1 in alveolar epithelial cell monolayers. Am J Physiol Lung Cell Mol Physiol. 2003 Dec; 285(6):L1192-200. PMID: 12730080.
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      98. Qiao R, Zhou B, Liebler JM, Li X, Crandall ED, Borok Z. Identification of three genes of known function expressed by alveolar epithelial type I cells. Am J Respir Cell Mol Biol. 2003 Jul; 29(1):98-105. PMID: 12600825.
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      99. Borok Z, Verkman AS. Lung edema clearance: 20 years of progress: invited review: role of aquaporin water channels in fluid transport in lung and airways. J Appl Physiol (1985). 2002 Dec; 93(6):2199-206. PMID: 12433939.
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      100. Li C, Xiao J, Hormi K, Borok Z, Minoo P. Wnt5a participates in distal lung morphogenesis. Dev Biol. 2002 Aug 01; 248(1):68-81. PMID: 12142021.
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      101. Mairbäurl H, Mayer K, Kim KJ, Borok Z, Bärtsch P, Crandall ED. Hypoxia decreases active Na transport across primary rat alveolar epithelial cell monolayers. Am J Physiol Lung Cell Mol Physiol. 2002 Apr; 282(4):L659-65. PMID: 11880290.
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      102. Borok Z, Liebler JM, Lubman RL, Foster MJ, Zhou B, Li X, Zabski SM, Kim KJ, Crandall ED. Na transport proteins are expressed by rat alveolar epithelial type I cells. Am J Physiol Lung Cell Mol Physiol. 2002 Apr; 282(4):L599-608. PMID: 11880284.
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      103. Borok Z, Harboe-Schmidt JE, Brody SL, You Y, Zhou B, Li X, Cannon PM, Kim KJ, Crandall ED, Kasahara N. Vesicular stomatitis virus G-pseudotyped lentivirus vectors mediate efficient apical transduction of polarized quiescent primary alveolar epithelial cells. J Virol. 2001 Dec; 75(23):11747-54. PMID: 11689655; PMCID: PMC114760.
      104. Kemp PJ, Kim KJ, Borok Z, Crandall ED. Re-evaluating the Na(+) conductance of adult rat alveolar type II pneumocytes: evidence for the involvement of cGMP-activated cation channels. J Physiol. 2001 Nov 01; 536(Pt 3):693-701. PMID: 11691865; PMCID: PMC2278905.
      105. Roum JH, Aledia AS, Carungcong LA, Kim KJ, Borok Z. Extracellular glutathione inhibits oxygen-induced permeability changes in alveolar epithelial monolayers. J Appl Physiol (1985). 2001 Aug; 91(2):748-54. PMID: 11457790.
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      106. Liu RM, Borok Z, Forman HJ. 4-Hydroxy-2-nonenal increases gamma-glutamylcysteine synthetase gene expression in alveolar epithelial cells. Am J Respir Cell Mol Biol. 2001 Apr; 24(4):499-505. PMID: 11306445.
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      107. Kim KJ, Borok Z, Crandall ED. A useful in vitro model for transport studies of alveolar epithelial barrier. Pharm Res. 2001 Mar; 18(3):253-5. PMID: 11442260.
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      108. Borok Z, Li X, Fernandes VF, Zhou B, Ann DK, Crandall ED. Differential regulation of rat aquaporin-5 promoter/enhancer activities in lung and salivary epithelial cells. J Biol Chem. 2000 Aug 25; 275(34):26507-14. PMID: 10849430.
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      109. Lubman RL, Zhang XL, Zheng J, Ocampo L, Lopez MZ, Veeraraghavan S, Zabski SM, Danto SI, Borok Z. Integrin alpha(3)-subunit expression modulates alveolar epithelial cell monolayer formation. Am J Physiol Lung Cell Mol Physiol. 2000 Jul; 279(1):L183-93. PMID: 10893217.
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      110. Kemp PJ, Borok Z, Kim KJ, Lubman RL, Danto SI, Crandall ED. Epidermal growth factor regulation in adult rat alveolar type II cells of amiloride-sensitive cation channels. Am J Physiol. 1999 Dec; 277(6 Pt 1):C1058-65. PMID: 10600757.
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      111. Roum JH, Borok Z, McElvaney NG, Grimes GJ, Bokser AD, Buhl R, Crystal RG. Glutathione aerosol suppresses lung epithelial surface inflammatory cell-derived oxidants in cystic fibrosis. J Appl Physiol (1985). 1999 Jul; 87(1):438-43. PMID: 10409605.
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      112. Borok Z, Mihyu S, Fernandes VF, Zhang XL, Kim KJ, Lubman RL. KGF prevents hyperoxia-induced reduction of active ion transport in alveolar epithelial cells. Am J Physiol. 1999 Jun; 276(6 Pt 1):C1352-60. PMID: 10362598.
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      113. Borok Z, Danto SI, Lubman RL, Cao Y, Williams MC, Crandall ED. Modulation of t1alpha expression with alveolar epithelial cell phenotype in vitro. Am J Physiol. 1998 Jul; 275(1 Pt 1):L155-64. PMID: 9688947.
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      114. Danto SI, Borok Z, Zhang XL, Lopez MZ, Patel P, Crandall ED, Lubman RL. Mechanisms of EGF-induced stimulation of sodium reabsorption by alveolar epithelial cells. Am J Physiol. 1998 Jul; 275(1 Pt 1):C82-92. PMID: 9688838.
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      115. Borok Z, Lubman RL, Danto SI, Zhang XL, Zabski SM, King LS, Lee DM, Agre P, Crandall ED. Keratinocyte growth factor modulates alveolar epithelial cell phenotype in vitro: expression of aquaporin 5. Am J Respir Cell Mol Biol. 1998 Apr; 18(4):554-61. PMID: 9533944.
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      116. Borok Z, Danto SI, Dimen LL, Zhang XL, Lubman RL. Na(+)-K(+)-ATPase expression in alveolar epithelial cells: upregulation of active ion transport by KGF. Am J Physiol. 1998 Jan; 274(1 Pt 1):L149-58. PMID: 9458813.
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      117. Zhang XL, Danto SI, Borok Z, Eber JT, Martín-Vasallo P, Lubman RL. Identification of Na(+)-K(+)-ATPase beta-subunit in alveolar epithelial cells. Am J Physiol. 1997 Jan; 272(1 Pt 1):L85-94. PMID: 9038907.
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      118. Borok Z, Hami A, Danto SI, Lubman RL, Kim KJ, Crandall ED. Effects of EGF on alveolar epithelial junctional permeability and active sodium transport. Am J Physiol. 1996 Apr; 270(4 Pt 1):L559-65. PMID: 8928815.
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      119. Danto SI, Shannon JM, Borok Z, Zabski SM, Crandall ED. Reversible transdifferentiation of alveolar epithelial cells. Am J Respir Cell Mol Biol. 1995 May; 12(5):497-502. PMID: 7742013.
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      120. Borok Z, Hami A, Danto SI, Zabski SM, Crandall ED. Rat serum inhibits progression of alveolar epithelial cells toward the type I cell phenotype in vitro. Am J Respir Cell Mol Biol. 1995 Jan; 12(1):50-5. PMID: 7811470.
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      121. Borok Z, Danto SI, Zabski SM, Crandall ED. Defined medium for primary culture de novo of adult rat alveolar epithelial cells. In Vitro Cell Dev Biol Anim. 1994 Feb; 30A(2):99-104. PMID: 8012662.
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      122. Roum JH, Buhl R, McElvaney NG, Borok Z, Crystal RG. Systemic deficiency of glutathione in cystic fibrosis. J Appl Physiol (1985). 1993 Dec; 75(6):2419-24. PMID: 8125859.
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      123. Travis WD, Borok Z, Roum JH, Zhang J, Feuerstein I, Ferrans VJ, Crystal RG. Pulmonary Langerhans cell granulomatosis (histiocytosis X). A clinicopathologic study of 48 cases. Am J Surg Pathol. 1993 Oct; 17(10):971-86. PMID: 8372949.
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      124. Holroyd KJ, Buhl R, Borok Z, Roum JH, Bokser AD, Grimes GJ, Czerski D, Cantin AM, Crystal RG. Correction of glutathione deficiency in the lower respiratory tract of HIV seropositive individuals by glutathione aerosol treatment. Thorax. 1993 Oct; 48(10):985-9. PMID: 8256245; PMCID: PMC464806.
      125. Buhl R, Jaffe HA, Holroyd KJ, Borok Z, Roum JH, Mastrangeli A, Wells FB, Kirby M, Saltini C, Crystal RG. Activation of alveolar macrophages in asymptomatic HIV-infected individuals. J Immunol. 1993 Feb 01; 150(3):1019-28. PMID: 8380824.
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      126. Borok Z, Gillissen A, Buhl R, Hoyt RF, Hubbard RC, Ozaki T, Rennard SI, Crystal RG. Augmentation of functional prostaglandin E levels on the respiratory epithelial surface by aerosol administration of prostaglandin E. Am Rev Respir Dis. 1991 Nov; 144(5):1080-4. PMID: 1952435.
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      127. Borok Z, Buhl R, Grimes GJ, Bokser AD, Hubbard RC, Holroyd KJ, Roum JH, Czerski DB, Cantin AM, Crystal RG. Effect of glutathione aerosol on oxidant-antioxidant imbalance in idiopathic pulmonary fibrosis. Lancet. 1991 Jul 27; 338(8761):215-6. PMID: 1676780.
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