Faculty

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Zea Borok, MD
Professor of Medicine
Ralph Edgington Chair in Medicine
Chief, Division of Pulmonary and Critical Care Medicine
Medicine
IRD 720 2020 Zonal Avenue Off Campus Los Angeles
+1 323 226 7923

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.

Awards

American Thoracic Society : Elizabeth Rich Award, 2017

Western Society for Clinical Investigation: Mayo Soley Award , 2017

Association for American Physicians: Elected to Association for American Physicians, 2014

American Thoracic Society: Recognition Award for Scientific Accomplishment, 2014

NHLBI: Method to Extend Research in Time (MERIT) Award, 2010

Western Society for Clinical Investigation: Outstanding Investigator Award, 2007

NIH: Clinical Investigator Development Award, 1993

Publications

Pleiotropic Analysis of Lung Cancer and Blood Triglycerides. J Natl Cancer Inst. 2016 Dec; 108(12). View in: PubMed

IL-1R1-MyD88 axis elicits papain-induced lung inflammation. Eur J Immunol. 2016 Nov; 46(11):2531-2541. View in: PubMed

Cross-species Transcriptome Profiling Identifies New Alveolar Epithelial Type I Cell-specific Genes. Am J Respir Cell Mol Biol. 2016 Oct 17. View in: PubMed

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. View in: PubMed

The importance of detailed epigenomic profiling of different cell types within organs. Epigenomics. 2016 Jun; 8(6):817-29. View in: PubMed

An American Thoracic Society Official Research Statement: Future Directions in Lung Fibrosis Research. Am J Respir Crit Care Med. 2016 Apr 1; 193(7):792-800. View in: PubMed

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. View in: PubMed

Mesodermal ALK5 controls lung myofibroblast versus lipofibroblast cell fate. BMC Biol. 2016; 14(1):19. View in: PubMed

Non-canonical WNT signalling in the lung. J Biochem. 2015 Nov; 158(5):355-65. View in: PubMed

Progenitors of secondary crest myofibroblasts are developmentally committed in early lung mesoderm. Stem Cells. 2015 Mar; 33(3):999-1012. View in: PubMed

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. View in: PubMed

Transforming Growth Factor-ß1 Induced Epithelial Mesenchymal Transition is blocked by a chemical antagonist of translation factor eIF4E. Sci Rep. 2015; 5:18233. View in: PubMed

Transcriptomic Profiling of Primary Alveolar Epithelial Cell Differentiation in Human and Rat. Genom Data. 2014 Dec 1; 2:105-109. View in: PubMed

Claudin 4 knockout mice: normal physiological phenotype with increased susceptibility to lung injury. Am J Physiol Lung Cell Mol Physiol. 2014 Oct 1; 307(7):L524-36. View in: PubMed

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. View in: PubMed

Alveolar epithelium: beyond the barrier. Am J Respir Cell Mol Biol. 2014 May; 50(5):853-6. View in: PubMed

Future Directions in Idiopathic Pulmonary Fibrosis Research. An NHLBI Workshop Report. Am J Respir Crit Care Med. 2014 Jan 15; 189(2):214-22. View in: PubMed

Dynamic involvement of ATG5 in cellular stress responses. Cell Death Dis. 2014; 5:e1478. View in: PubMed

Characterizing the genetic basis of methylome diversity in histologically normal human lung tissue. Nat Commun. 2014; 5:3365. View in: PubMed

Deletion of ATG5 shows a role of autophagy in salivary homeostatic control. J Dent Res. 2013 Oct; 92(10):911-7. View in: PubMed

Nanoparticle translocation across mouse alveolar epithelial cell monolayers: species-specific mechanisms. Nanomedicine. 2013 Aug; 9(6):786-94. View in: PubMed

Apc deficiency alters pulmonary epithelial cell fate and inhibits Nkx2. 1 via triggering TGF-beta signaling. Dev Biol. 2013 Jun 1; 378(1):13-24. View in: PubMed

Integrated transcriptomic and epigenomic analysis of primary human lung epithelial cell differentiation. PLoS Genet. 2013 Jun; 9(6):e1003513. View in: PubMed

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. View in: PubMed

Aqp5 is a new transcriptional target of Dot1a and a regulator of Aqp2. PLoS One. 2013; 8(1):e53342. View in: PubMed

Mesodermal Pten inactivation leads to alveolar capillary dysplasia- like phenotype. J Clin Invest. 2012 Nov 1; 122(11):3862-72. View in: PubMed

EMT and interstitial lung disease: a mysterious relationship. Curr Opin Pulm Med. 2012 Sep; 18(5):517-23. View in: PubMed

NOTCH1 Is Required for Regeneration of Clara Cells During Repair of Airway Injury. Stem Cells. 2012 May; 30(5):946-55. View in: PubMed

Cigarette smoke extract stimulates epithelial-mesenchymal transition through Src activation. Free Radic Biol Med. 2012 Apr 15; 52(8):1437-42. View in: PubMed

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 2; 287(10):7026-38. View in: PubMed

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. View in: PubMed

In search of the fibrotic epithelial cell: opportunities for a collaborative network. Thorax. 2012 Feb; 67(2):179-82. View in: PubMed

Troglitazone Attenuates TGF-ß1-Induced EMT in Alveolar Epithelial Cells via a PPAR?-Independent Mechanism. Troglitazone Attenuates TGF-ß1-Induced EMT in Alveolar Epithelial Cells via a PPAR? -Independent Mechanism. PLoS One. 2012; 7(6):e38827. View in: PubMed

Regulation of breast cancer metastasis by Runx2 and estrogen signaling: the role of SNAI2. Breast Cancer Res. 2011 Dec; 13(6):R127. View in: PubMed

Polystyrene nanoparticle trafficking across MDCK-II. Nanomedicine. 2011 Oct; 7(5):588-94. View in: PubMed

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. View in: PubMed

Stem cells and cell therapies in lung biology and lung diseases. Proc Am Thorac Soc. 2011 Jun; 8(3):223-72. View in: PubMed

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. View in: PubMed

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 1; 4(1):65-78. View in: PubMed

Epithelium-specific deletion of TGF-ß receptor type II protects mice from bleomycin-induced pulmonary fibrosis. J Clin Invest. 2011 Jan 4; 121(1):277-87. View in: PubMed

Translocation of PEGylated quantum dots across rat alveolar epithelial cell monolayers. Int J Nanomedicine. 2011; 6:2849-57. View in: PubMed

Alveolar epithelial cell injury due to zinc oxide nanoparticle exposure. Am J Respir Crit Care Med. 2010 Dec 1; 182(11):1398-409. View in: PubMed

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. View in: PubMed

Directed expression of Cre in alveolar epithelial type 1 cells. Am J Respir Cell Mol Biol. 2010 Aug; 43(2):173-8. View in: PubMed

Cell type-specific expression of adenomatous polyposis coli in lung development, injury, and repair. Dev Dyn. 2010 Aug; 239(8):2288-97. View in: PubMed

Mechanisms of alveolar epithelial translocation of a defined population of nanoparticles. Am J Respir Cell Mol Biol. 2010 May; 42(5):604-14. View in: PubMed

Signaling via Alk5 controls the ontogeny of lung Clara cells. Development. 2010 Mar; 137(5):825-33. View in: PubMed

More life for a "terminal" cell. Am J Physiol Lung Cell Mol Physiol. 2009 Dec; 297(6):L1042-4. View in: PubMed

N-acetylcysteine inhibits alveolar epithelial-mesenchymal transition. Am J Physiol Lung Cell Mol Physiol. 2009 Nov; 297(5):L805-12. View in: PubMed

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. View in: PubMed

Epithelial-mesenchymal transition: potential role in obliterative bronchiolitis? Thorax. Epithelial-mesenchymal transition: potential role in obliterative bronchiolitis? Thorax. 2009 Sep; 64(9):742-3. View in: PubMed

Characterization of mouse alveolar epithelial cell monolayers. Am J Physiol Lung Cell Mol Physiol. 2009 Jun; 296(6):L1051-8. View in: PubMed

Conjugation with cationic cell-penetrating peptide increases pulmonary absorption of insulin. Mol Pharm. 2009 Mar-Apr; 6(2):492-503. View in: PubMed

Role for alpha3 integrin in EMT and pulmonary fibrosis. J Clin Invest. 2009 Jan; 119(1):7-10. View in: PubMed

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. View in: PubMed

GATA-6 mediates transcriptional activation of aquaporin-5 through interactions with Sp1. Am J Physiol Cell Physiol. 2008 Nov; 295(5):C1141-50. View in: PubMed

Progressive recruitment of Runx2 to genomic targets despite decreasing expression during osteoblast differentiation. J Cell Biochem. 2008 Nov 1; 105(4):965-70. View in: PubMed

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. View in: PubMed

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. View in: PubMed

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. View in: PubMed

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. View in: PubMed

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. View in: PubMed

Polystyrene nanoparticle trafficking across alveolar epithelium. Nanomedicine. 2008 Jun; 4(2):139-45. View in: PubMed

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. View in: PubMed

Expression and biological activity of ABCA1 in alveolar epithelial cells. Am J Respir Cell Mol Biol. 2008 Mar; 38(3):283-92. View in: PubMed

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. View in: PubMed

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. View in: PubMed

Nanoparticle effects on rat alveolar epithelial cell monolayer barrier properties. Toxicol In Vitro. 2007 Dec; 21(8):1373-81. View in: PubMed

TGF-beta-induced EMT: mechanisms and implications for fibrotic lung disease. Am J Physiol Lung Cell Mol Physiol. 2007 Sep; 293(3):L525-34. View in: PubMed

Enhancement of insulin transport across primary rat alveolar epithelial cell monolayers by endogenous cellular factor(s). Pharm Res. 2007 Sep; 24(9):1713-9. View in: PubMed

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. View in: PubMed

Conserved elements within first intron of aquaporin-5 (Aqp5) function as transcriptional enhancers. Biochem Biophys Res Commun. 2007 Apr 27; 356(1):26-31. View in: PubMed

Hypertonic induction of aquaporin-5: novel role of hypoxia-inducible factor-1alpha. Am J Physiol Cell Physiol. 2007 Apr; 292(4):C1280-90. View in: PubMed

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. View in: PubMed

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. View in: PubMed

Epithelial origin of myofibroblasts during fibrosis in the lung. Proc Am Thorac Soc. 2006 Jun; 3(4):377-82. View in: PubMed

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. View in: PubMed

Adult stem cells, lung biology, and lung disease. NHLBI/Cystic Fibrosis Foundation Workshop. Proc Am Thorac Soc. 2006 May; 3(3):193-207. View in: PubMed

Developmental pathways and specification of intrapulmonary stem cells. Pediatr Res. 2006 Apr; 59(4 Pt 2):84R-93R. View in: PubMed

Induction of epithelial-mesenchymal transition in alveolar epithelial cells by transforming growth factor-beta1: potential role in idiopathic pulmonary fibrosis. Am J Pathol. 2005 May; 166(5):1321-32. View in: PubMed

Estimation of paracellular conductance of primary rat alveolar epithelial cell monolayers. J Appl Physiol. 2005 Jan; 98(1):138-43. View in: PubMed

Development of a lung slice preparation for recording ion channel activity in alveolar epithelial type I cells. Respir Res. 2005; 6:40. View in: PubMed

Effects of transdifferentiation and EGF on claudin isoform expression in alveolar epithelial cells. J Appl Physiol. 2005 Jan; 98(1):322-8. View in: PubMed

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. View in: PubMed

Alveolar epithelial type I cells express beta2-adrenergic receptors and G-protein receptor kinase 2. J Histochem Cytochem. 2004 Jun; 52(6):759-67. View in: PubMed

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. View in: PubMed

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. View in: PubMed

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. View in: PubMed

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. View in: PubMed

Timeless in lung morphogenesis. Dev Dyn. 2003 Sep; 228(1):82-94. View in: PubMed

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. View in: PubMed

Lung edema clearance: 20 years of progress: invited review: role of aquaporin water channels in fluid transport in lung and airways. J Appl Physiol. 2002 Dec; 93(6):2199-206. View in: PubMed

Wnt5a participates in distal lung morphogenesis. Dev Biol. 2002 Aug 1; 248(1):68-81. View in: PubMed

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. View in: PubMed

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. View in: PubMed

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. View in: PubMed

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 1; 536(Pt 3):693-701. View in: PubMed

Extracellular glutathione inhibits oxygen-induced permeability changes in alveolar epithelial monolayers. J Appl Physiol. 2001 Aug; 91(2):748-54. View in: PubMed

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. View in: PubMed

A useful in vitro model for transport studies of alveolar epithelial barrier. Pharm Res. 2001 Mar; 18(3):253-5. View in: PubMed

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. View in: PubMed

Integrin alpha(3)-subunit expression modulates alveolar epithelial cell monolayer formation. Am J Physiol Lung Cell Mol Physiol. 2000 Jul; 279(1):L183-93. View in: PubMed

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. View in: PubMed

Glutathione aerosol suppresses lung epithelial surface inflammatory cell-derived oxidants in cystic fibrosis. J Appl Physiol. 1999 Jul; 87(1):438-43. View in: PubMed

KGF prevents hyperoxia-induced reduction of active ion transport in alveolar epithelial cells. Am J Physiol. 1999 Jun; 276(6 Pt 1):C1352-60. View in: PubMed

Mechanisms of EGF-induced stimulation of sodium reabsorption by alveolar epithelial cells. Am J Physiol. 1998 Jul; 275(1 Pt 1):C82-92. View in: PubMed

Modulation of t1alpha expression with alveolar epithelial cell phenotype in vitro. Am J Physiol. 1998 Jul; 275(1 Pt 1):L155-64. View in: PubMed

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. View in: PubMed

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. View in: PubMed

Identification of Na(+)-K(+)-ATPase beta-subunit in alveolar epithelial cells. Am J Physiol. 1997 Jan; 272(1 Pt 1):L85-94. View in: PubMed

Effects of EGF on alveolar epithelial junctional permeability and active sodium transport. Am J Physiol. 1996 Apr; 270(4 Pt 1):L559-65. View in: PubMed

Reversible transdifferentiation of alveolar epithelial cells. Am J Respir Cell Mol Biol. 1995 May; 12(5):497-502. View in: PubMed

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. View in: PubMed

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. View in: PubMed

Systemic deficiency of glutathione in cystic fibrosis. J Appl Physiol. 1993 Dec; 75(6):2419-24. View in: PubMed

Pulmonary Langerhans cell granulomatosis (histiocytosis X). A clinicopathologic study of 48 cases. Am J Surg Pathol. 1993 Oct; 17(10):971-86. View in: PubMed

Correction of glutathione deficiency in the lower respiratory tract of HIV seropositive individuals by glutathione aerosol treatment. Thorax. 1993 Oct; 48(10):985-9. View in: PubMed

Activation of alveolar macrophages in asymptomatic HIV-infected individuals. J Immunol. 1993 Feb 1; 150(3):1019-28. View in: PubMed

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. View in: PubMed

Effect of glutathione aerosol on oxidant-antioxidant imbalance in idiopathic pulmonary fibrosis. Lancet. 1991 Jul 27; 338(8761):215-6. View in: PubMed

Region-specific role for Pten in maintenance of epithelial phenotype and integrity. Am J Physiol Lung Cell Mol Physiol. 2016 Nov 18; ajplung. 00005. 2015. View in: PubMed

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