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Aloysius J. Klingelhutz, PhD

Professor of Microbiology and Immunology
Professor of Radiation Oncology

Contact Information

Primary Office
3-612 Bowen Science Building
51 Newton Rd
Iowa City, IA 52242

3-615A Bowen Science Building
51 Newton Rd
Iowa City, IA 52242


BS, Biology, St John's University
PhD, Genetics, University of Wisconsin-Madison
Postdoctoral Fellow, Cancer Biology, Fred Hutchinson Cancer Research Center

Education/Training Program Affiliations

Biomedical Science Program, Department of Microbiology Graduate Program, Free Radical and Radiation Biology Graduate Program, Interdisciplinary Graduate Program in Genetics, Interdisciplinary Graduate Program in Human Toxicology, Interdisciplinary Graduate Program in Molecular Medicine, Medical Scientist Training Program

Center, Program and Institute Affiliations

Fraternal Order of Eagles Diabetes Research Center, Helen C. Levitt Center for Viral Pathogenesis, Holden Comprehensive Cancer Center

Research Summary

Our research has diversified into two main areas: 1) How viruses infect and disseminate in skin; 2) How microbial and environmental factors play a role in the development of metabolic syndrome. Our prior research addressed how human cells senesce, leading to aging, and how they become immortal, leading to cancer, with a particular interest in on how human papillomaviruses transform cells. Our expertise in cell immortalization and cell culture techniques has allowed us develop 3D cell culture models that recapitulate human tissue for our research.

1) How viruses infect and disseminate in skin. Collaborative studies were recently initiated with Wendy Maury’s lab to examine how Ebola virus (EBOV) infects and transmits through human skin. We found that EBOV can infect and replicate in different skin cell populations. We are currently working to understand the course of infection in skin, what specific receptors are being utilized by EBOV in skin cells, and what role skin infection plays in transmission and pathogenesis.

2) How microbial and environmental factors play a role in the development of metabolic syndrome. Our success with immortalizing human preadipocytes (pre-fat) cells has led to studies on how environmental and bacterial toxins cause or exacerbate type II diabetes through effects on fat tissue. We found that dioxin-like polychlorinated biphenyls (PCBs), which are persistent organic pollutants, can disrupt adipogenesis (i.e. the development of functional fat cells) through activation of the aryl hydrocarbon receptor (AhR). This causes a proinflammatory response and inhibits master regulatory genes involved in adipogenesis. Endogenous microbial-derived tryptophan metabolites are also able to activate AhR. Studies are underway to determine the mechanism by which AhR activation disrupts adipogenesis and to develop 3D cultures and in vivo genetic models to assess the role of AhR in the development of metabolic syndrome.


Schlievert, P. M., Gourronc, F. A., Leung, D. Y. & Klingelhutz, A. J. (2020). Human Keratinocyte Response to Superantigens. mSphere, 5(5), e00803-20. PMID: 33028686.

Gourronc, F. A., Markan, K. R., Kulhankova, K., Zhu, Z., Sheehy, R., Quelle, D. E., Zingman, L. V., Kurago, Z. B., Ankrum, J. A. & Klingelhutz, A. J. (2020). Pdgfra-Cre mediated knockout of the aryl hydrocarbon receptor protects mice from high-fat diet induced obesity and hepatic steatosis. PLoS One, 15(7), e0236741. PMID: 32730300.

El-Hattab, M. Y., Nagumo, Y., Gourronc, F. A., Klingelhutz, A. J., Ankrum, J. A. & Sander, E. A. (2020). Human Adipocyte Conditioned Medium Promotes In Vitro Fibroblast Conversion to Myofibroblasts. Sci Rep, 10(1), 10286. PMID: 32581231.

Gourronc, F. A., Perdew, G. H., Robertson, L. W. & Klingelhutz, A. J. (2020). PCB126 blocks the thermogenic beiging response of adipocytes. Environ Sci Pollut Res Int, 27(9), 8897-8904. PMID: 31721030.

Faris, R., Andersen, S. E., McCullough, A., Gourronc, F., Klingelhutz, A. J. & Weber, M. M. (2019). Chlamydia trachomatis Serovars Drive Differential Production of Proinflammatory Cytokines and Chemokines Depending on the Type of Cell Infected. Front Cell Infect Microbiol, 9, 399. PMID: 32039039.

Schlievert, P. M., Cahill, M. P., Hostager, B. S., Brosnahan, A. J., Klingelhutz, A. J., Gourronc, F. A., Bishop, G. A. & Leung, D. (2019). Staphylococcal Superantigens Stimulate Epithelial Cells through CD40 To Produce Chemokines. MBio, 10(2), e00214-19. PMID: 30890614.

Kolb, R., Kluz, P., Tan, Z. W., Borcherding, N., Bormann, N., Vishwakarma, A., Balcziak, L., Zhu, P., Davies, B. S., Gourronc, F., Liu, L. Z., Ge, X., Jiang, B. H., Gibson-Corley, K., Klingelhutz, A. J., Tan, N. S., Zhu, Y., Sutterwala, F. S., Shen, X. & Zhang, W. (2019). Obesity-associated inflammation promotes angiogenesis and breast cancer via angiopoietin-like 4. Oncogene, 38(13), 2351-2363. PMID: 30518876.

Gourronc, F. A., Robertson, L. W. & Klingelhutz, A. J. (2018). A delayed proinflammatory response of human preadipocytes to PCB126 is dependent on the aryl hydrocarbon receptor. Environ Sci Pollut Res Int, 25(17), 16481-16492. PMID: 28699004.

Kulhankova, K., Kinney, K. J., Stach, J. M., Gourronc, F. A., Grumbach, I. M., Klingelhutz, A. J. & Salgado-Pabón, W. (2018). The Superantigen Toxic Shock Syndrome Toxin 1 Alters Human Aortic Endothelial Cell Function. Infect Immun, 86(3), e00848-17. PMID: 29229737.

Klingelhutz, A. J., Gourronc, F. A., Chaly, A., Wadkins, D. A., Burand, A. J., Markan, K. R., Idiga, S. O., Wu, M., Potthoff, M. J. & Ankrum, J. A. (2018). Scaffold-free generation of uniform adipose spheroids for metabolism research and drug discovery. Sci Rep, 8(1), 523. PMID: 29323267.

Scorza, B. M., Wacker, M. A., Messingham, K., Kim, P., Klingelhutz, A., Fairley, J. & Wilson, M. E. (2017). Differential Activation of Human Keratinocytes by Leishmania spp. Causing Localized or Disseminated Disease. J Invest Dermatol, 137(10), 2149-2156. PMID: 28647347.

Zhang, Y., Xie, L., Gunasekar, S. K., Tong, D., Mishra, A., Gibson, W. J., Wang, C., Fidler, T., Marthaler, B., Klingelhutz, A. J., Abel, E. D., Samuel, I., Smith, J. K., Cao, L. & Sah, R. (2017). SWELL1 is a regulator of adipocyte size, insulin signalling and glucose homeostasis. Nat Cell Biol, 19(5), 504-517. PMID: 28436964.

Jang, J. H., Chand, H. S., Bruse, S., Doyle-Eisele, M., Royer, C., McDonald, J., Qualls, C., Klingelhutz, A. J., Lin, Y., Mallampalli, R., Tesfaigzi, Y. & Nyunoya, T. (2017). Connective Tissue Growth Factor Promotes Pulmonary Epithelial Cell Senescence and Is Associated with COPD Severity. COPD, 14(2), 228-237. PMID: 28026993.

Herrera, A., Kulhankova, K., Sonkar, V. K., Dayal, S., Klingelhutz, A. J., Salgado-Pabón, W. & Schlievert, P. M. (2017). Staphylococcal ß-Toxin Modulates Human Aortic Endothelial Cell and Platelet Function through Sphingomyelinase and Biofilm Ligase Activities. MBio, 8(2), e00273-17. PMID: 28325766.

Gadupudi, G. S., Klingelhutz, A. J. & Robertson, L. W. (2016). Diminished Phosphorylation of CREB Is a Key Event in the Dysregulation of Gluconeogenesis and Glycogenolysis in PCB126 Hepatotoxicity. Chem Res Toxicol, 29(9), 1504-9. PMID: 27509375.

Littlejohn, N. K., Keen, H. L., Weidemann, B. J., Claflin, K. E., Tobin, K. V., Markan, K. R., Park, S., Naber, M. C., Gourronc, F. A., Pearson, N. A., Liu, X., Morgan, D. A., Klingelhutz, A. J., Potthoff, M. J., Rahmouni, K., Sigmund, C. D. & Grobe, J. L. (2016). Suppression of Resting Metabolism by the Angiotensin AT2 Receptor. Cell Rep, 16(6), 1548-60. PMID: 27477281.

Gadupudi, G. S., Klaren, W. D., Olivier, A. K., Klingelhutz, A. J. & Robertson, L. W. (2016). PCB126-induced disruption in gluconeogenesis and fatty acid oxidation precedes fatty liver in male rats. Toxicol Sci, 149(1), 98-110. PMID: 26396156.

Merriman, J. A., Klingelhutz, A. J., Diekema, D. J., Leung, D. Y. & Schlievert, P. M. (2015). Novel Staphylococcus aureus Secreted Protein Alters Keratinocyte Proliferation and Elicits a Proinflammatory Response In Vitro and In Vivo. Biochemistry, 54(31), 4855-62. PMID: 26177220.

Vu, B. G., Stach, C. S., Kulhankova, K., Salgado-Pabón, W., Klingelhutz, A. J. & Schlievert, P. M. (2015). Chronic superantigen exposure induces systemic inflammation, elevated bloodstream endotoxin, and abnormal glucose tolerance in rabbits: possible role in diabetes. MBio, 6(2), e02554. PMID: 25714716.

Gadupudi, G., Gourronc, F. A., Ludewig, G., Robertson, L. W. & Klingelhutz, A. J. (2015). PCB126 inhibits adipogenesis of human preadipocytes. Toxicol In Vitro, 29(1), 132-41. PMID: 25304490.

Schlievert, P. M., Salgado-Pabón, W. & Klingelhutz, A. J. (2015). Does Staphylococcus aureus have a role in the development of Type 2 diabetes mellitus?. Future Microbiol, 10(10), 1549-52. PMID: 26439811.

Hagen, J., Muniz, V. P., Falls, K. C., Reed, S. M., Taghiyev, A. F., Quelle, F. W., Gourronc, F. A., Klingelhutz, A. J., Major, H. J., Askeland, R. W., Sherman, S. K., O'Dorisio, T. M., Bellizzi, A. M., Howe, J. R., Darbro, B. W. & Quelle, D. E. (2014). RABL6A Promotes G1-S Phase Progression and Pancreatic Neuroendocrine Tumor Cell Proliferation in an Rb1-Dependent Manner. Cancer Res, 74(22), 6661-70. PMID: 25273089.

Jang, J. H., Bruse, S., Huneidi, S., Schrader, R. M., Monick, M. M., Lin, Y., Carter, A. B., Klingelhutz, A. J. & Nyunoya, T. (2014). Acrolein-exposed normal human lung fibroblasts in vitro: cellular senescence, enhanced telomere erosion, and degradation of werner's syndrome protein. Environ Health Perspect, 122(9), 955-62. PMID: 24747221.

Gourronc, F. A., Rockey, W. M., Thiel, W. H., Giangrande, P. H. & Klingelhutz, A. J. (2013). Identification of RNA aptamers that internalize into HPV-16 E6/E7 transformed tonsillar epithelial cells. Virology, 446(1-2), 325-33. PMID: 24074596.

Vu, B. G., Gourronc, F. A., Bernlohr, D. A., Schlievert, P. M. & Klingelhutz, A. J. (2013). Staphylococcal superantigens stimulate immortalized human adipocytes to produce chemokines. PLoS One, 8(10), e77988. PMID: 24205055.

Pereboeva, L., Westin, E., Patel, T., Flaniken, I., Lamb, L., Klingelhutz, A. J. & Goldman, F. (2013). DNA damage responses and oxidative stress in dyskeratosis congenita. PLoS One, 8(10), e76473. PMID: 24124565.

Vande Pol, S. B., Klingelhutz, A. J. (2013). Papillomavirus E6 oncoproteins. Virology, 445(1-2), 115-37. PMID: 23711382.

Kimple, R. J., Smith, M. A., Blitzer, G. C., Torres, A. D., Martin, J. A., Yang, R. Z., Peet, C. R., Lorenz, L. D., Nickel, K. P., Klingelhutz, A. J., Lambert, P. F. & Harari, P. M. (2013). Enhanced radiation sensitivity in HPV-positive head and neck cancer. Cancer Res, 73(15), 4791-800. PMID: 23749640.

Hernandez, L. I., Flenker, K. S., Hernandez, F. J., Klingelhutz, A. J., McNamera, J. O. & Giangrande, P. H. (2013). Methods for Evaluating Cell-Specific, Cell-Internalizing RNA Aptamers. Pharmaceuticals (Basel), 6(3), 295-319. PMID: 23894227.

Buckingham, E. M., Goldman, F. D. & Klingelhutz, A. J. (2012). Dyskeratosis Congenita Dermal Fibroblasts are Defective in Supporting the Clonogenic Growth of Epidermal Keratinocytes. Aging Dis, 3(6), 427-37. PMID: 23251848.

Klingelhutz, A. J., Roman, A. (2012). Cellular transformation by human papillomaviruses: lessons learned by comparing high- and low-risk viruses. Virology, 424(2), 77-98. PMID: 22284986.

Bulysheva, A. A., Bowlin, G. L., Klingelhutz, A. J. & Yeudall, W. A. (2012). Low-temperature electrospun silk scaffold for in vitro mucosal modeling. J Biomed Mater Res A, 100(3), 757-67. PMID: 22238242.

Gourronc, F. A., Klingelhutz, A. J. (2012). Therapeutic opportunities: telomere maintenance in inducible pluripotent stem cells. Mutat Res, 730(1-2), 98-105. PMID: 21605571.

Vermeer, P. D., Bell, M., Lee, K., Vermeer, D. W., Wieking, B. G., Bilal, E., Bhanot, G., Drapkin, R. I., Ganesan, S., Klingelhutz, A. J., Hendriks, W. J. & Lee, J. H. (2012). ErbB2, EphrinB1, Src kinase and PTPN13 signaling complex regulates MAP kinase signaling in human cancers. PLoS One, 7(1), e30447. PMID: 22279592.

Senthikumar, P. K., Klingelhutz, A. J., Jacobus, J. A., Lehmler, H., Robertson, L. W. & Ludewig, G. (2011). Airborne polychlorinated biphenyls (PCBs) reduce telomerase activity and shorten telomere length in immortal human skin keratinocytes (HaCat). Toxicol Lett, 204(1), 64-70. PMID: 21530622.

Buckingham, E. M., Klingelhutz, A. J. (2011). The role of telomeres in the ageing of human skin. Exp Dermatol, 20(4), 297-302. PMID: 21371125.

Saccoccio, F. M., Sauer, A. L., Cui, X., Armstrong, A. E., Habib, e. l., Johnson, D. C., Ryckman, B. J., Klingelhutz, A. J., Adler, S. P. & McVoy, M. A. (2011). Peptides from cytomegalovirus UL130 and UL131 proteins induce high titer antibodies that block viral entry into mucosal epithelial cells. Vaccine, 29(15), 2705-11. PMID: 21310190.

Westin, E. R., Aykin-Burns, N., Buckingham, E. M., Spitz, D. R., Goldman, F. D. & Klingelhutz, A. J. (2011). The p53/p21(WAF/CIP) pathway mediates oxidative stress and senescence in dyskeratosis congenita cells with telomerase insufficiency. Antioxid Redox Signal, 14(6), 985-97. PMID: 21087144.

Agarwal, S., Loh, Y. H., McLoughlin, E. M., Huang, J., Park, I. H., Miller, J. D., Huo, H., Okuka, M., Dos Reis, R. M., Loewer, S., Ng, H. H., Keefe, D. L., Goldman, F. D., Klingelhutz, A. J., Liu, L. & Daley, G. Q. (2010). Telomere elongation in induced pluripotent stem cells from dyskeratosis congenita patients. Nature, 464(7286), 292-6. PMID: 20164838.

Gourronc, F. A., Robertson, M. M., Herrig, A. K., Lansdorp, P. M., Goldman, F. D. & Klingelhutz, A. J. (2010). Proliferative defects in dyskeratosis congenita skin keratinocytes are corrected by expression of the telomerase reverse transcriptase, TERT, or by activation of endogenous telomerase through expression of papillomavirus E6/E7 or the telomerase RNA component, TERC. Exp Dermatol, 19(3), 279-88. PMID: 19558498.

Hoover, A. C., Strand, G. L., Nowicki, P. N., Anderson, M. E., Vermeer, P. D., Klingelhutz, A. J., Bossler, A. D., Pottala, J. V., Hendriks, W. J. & Lee, J. H. (2009). Impaired PTPN13 phosphatase activity in spontaneous or HPV-induced squamous cell carcinomas potentiates oncogene signaling through the MAP kinase pathway. Oncogene, 28(45), 3960-70. PMID: 19734941.

Lace, M. J., Anson, J. R., Klingelhutz, A. J., Lee, J. H., Bossler, A. D., Haugen, T. H. & Turek, L. P. (2009). Human papillomavirus (HPV) type 18 induces extended growth in primary human cervical, tonsillar, or foreskin keratinocytes more effectively than other high-risk mucosal HPVs. J Virol, 83(22), 11784-94. PMID: 19740985.

Lace, M. J., Anson, J. R., Klingelhutz, A. J., Harada, H., Taniguchi, T., Bossler, A. D., Haugen, T. H. & Turek, L. P. (2009). Interferon-beta treatment increases human papillomavirus early gene transcription and viral plasmid genome replication by activating interferon regulatory factor (IRF)-1. Carcinogenesis, 30(8), 1336-44. PMID: 19541854.

Nyunoya, T., Monick, M. M., Klingelhutz, A. J., Glaser, H., Cagley, J. R., Brown, C. O., Matsumoto, E., Aykin-Burns, N., Spitz, D. R., Oshima, J. & Hunninghake, G. W. (2009). Cigarette smoke induces cellular senescence via Werner's syndrome protein down-regulation. Am J Respir Crit Care Med, 179(4), 279-87. PMID: 19011155.

Yu, L., Hitchler, M. J., Sun, W., Sarsour, E. H., Goswami, P. C., Klingelhutz, A. J. & Domann, F. E. (2009). AP-2alpha Inhibits c-MYC Induced Oxidative Stress and Apoptosis in HaCaT Human Keratinocytes. J Oncol, 2009, 780874. PMID: 20066163.

Goldman, F. D., Aubert, G., Klingelhutz, A. J., Hills, M., Cooper, S. R., Hamilton, W. S., Schlueter, A. J., Lambie, K., Eaves, C. J. & Lansdorp, P. M. (2008). Characterization of primitive hematopoietic cells from patients with dyskeratosis congenita. Blood, 111(9), 4523-31. PMID: 18310499.

Jacobus, J. A., Flor, S., Klingelhutz, A. J., Robertson, L. W. & Ludewig, G. (2008). 2-(4'-chlorophenyl)-1,4-benzoquinone increases the frequency of micronuclei and shortens telomeres. Environ Toxicol Pharmacol, 25(2), 267-72. PMID: 18438462.

Spanos, W. C., Hoover, A., Harris, G. F., Wu, S., Strand, G. L., Anderson, M. E., Klingelhutz, A. J., Hendriks, W., Bossler, A. D. & Lee, J. H. (2008). The PDZ binding motif of human papillomavirus type 16 E6 induces PTPN13 loss, which allows anchorage-independent growth and synergizes with ras for invasive growth. J Virol, 82(5), 2493-500. PMID: 18160445.

Spanos, W. C., Geiger, J., Anderson, M. E., Harris, G. F., Bossler, A. D., Smith, R. B., Klingelhutz, A. J. & Lee, J. H. (2008). Deletion of the PDZ motif of HPV16 E6 preventing immortalization and anchorage-independent growth in human tonsil epithelial cells. Head Neck, 30(2), 139-47. PMID: 17657785.

Westin, E. R., Chavez, E., Lee, K. M., Gourronc, F. A., Riley, S., Lansdorp, P. M., Goldman, F. D. & Klingelhutz, A. J. (2007). Telomere restoration and extension of proliferative lifespan in dyskeratosis congenita fibroblasts. Aging Cell, 6(3), 383-94. PMID: 17381549.

Hoover, A. C., Spanos, W. C., Harris, G. F., Anderson, M. E., Klingelhutz, A. J. & Lee, J. H. (2007). The role of human papillomavirus 16 E6 in anchorage-independent and invasive growth of mouse tonsil epithelium. Arch Otolaryngol Head Neck Surg, 133(5), 495-502. PMID: 17515506.

Berger, K. L., Barriga, F., Lace, M. J., Turek, L. P., Zamba, G. J., Domann, F. E., Lee, J. H. & Klingelhutz, A. J. (2006). Cervical keratinocytes containing stably replicating extrachromosomal HPV-16 are refractory to transformation by oncogenic H-Ras. Virology, 356(1-2), 68-78. PMID: 16945398.

Nyunoya, T., Monick, M. M., Klingelhutz, A. J., Yarovinsky, T. O., Cagley, J. R. & Hunninghake, G. W. (2006). Cigarette smoke induces cellular senescence. Am J Respir Cell Mol Biol, 35(6), 681-8. PMID: 16840774.

Darbro, B. W., Lee, K. M., Nguyen, N. K., Domann, F. E. & Klingelhutz, A. J. (2006). Methylation of the p16(INK4a) promoter region in telomerase immortalized human keratinocytes co-cultured with feeder cells. Oncogene, 25(56), 7421-33. PMID: 16767161.

James, M. A., Lee, J. H. & Klingelhutz, A. J. (2006). HPV16-E6 associated hTERT promoter acetylation is E6AP dependent, increased in later passage cells and enhanced by loss of p300. Int J Cancer, 119(8), 1878-85. PMID: 16708385.

James, M. A., Lee, J. H. & Klingelhutz, A. J. (2006). Human papillomavirus type 16 E6 activates NF-kappaB, induces cIAP-2 expression, and protects against apoptosis in a PDZ binding motif-dependent manner. J Virol, 80(11), 5301-7. PMID: 16699010.

Klingelhutz, A. J., Qian, Q., Phillips, S. L., Gourronc, F. A., Darbro, B. W. & Patil, S. R. (2005). Amplification of the chromosome 20q region is associated with expression of HPV-16 E7 in human airway and anogenital epithelial cells. Virology, 340(2), 237-44. PMID: 16051300.

Darbro, B. W., Schneider, G. B. & Klingelhutz, A. J. (2005). Co-regulation of p16INK4A and migratory genes in culture conditions that lead to premature senescence in human keratinocytes. J Invest Dermatol, 125(3), 499-509. PMID: 16117791.

Ault, K. A., Allen, H. K., Phillips, S. L., Zimmerman, M. B. & Klingelhutz, A. J. (2005). Telomerase activity as a potential diagnostic marker for triage of abnormal Pap smears. J Low Genit Tract Dis, 9(2), 93-9. PMID: 15870530.

Peterson, M. L., Ault, K., Kremer, M. J., Klingelhutz, A. J., Davis, C. C., Squier, C. A. & Schlievert, P. M. (2005). The innate immune system is activated by stimulation of vaginal epithelial cells with Staphylococcus aureus and toxic shock syndrome toxin 1. Infect Immun, 73(4), 2164-74. PMID: 15784559.

Landen, C. N., Klingelhutz, A. J., Coffin, J. E., Sorosky, J. I. & Sood, A. K. (2004). Genomic instability is associated with lack of telomerase activation in ovarian cancer. Cancer Biol Ther, 3(12), 1250-3. PMID: 15477760.

Martin, J. A., DeYoung, B. R., Gitelis, S., Weydert, J. A., Klingelhutz, A. J., Kurriger, G. & Buckwalter, J. A. (2004). Telomerase reverse transcriptase subunit expression is associated with chondrosarcoma malignancy. Clin Orthop Relat Res, 426, 117-24. PMID: 15346061.

Plug-DeMaggio, A. W., Sundsvold, T., Wurscher, M. A., Koop, J. I., Klingelhutz, A. J. & McDougall, J. K. (2004). Telomere erosion and chromosomal instability in cells expressing the HPV oncogene 16E6. Oncogene, 23(20), 3561-71. PMID: 15077181.

Martin, J. A., Klingelhutz, A. J., Moussavi-Harami, F. & Buckwalter, J. A. (2004). Effects of oxidative damage and telomerase activity on human articular cartilage chondrocyte senescence. J Gerontol A Biol Sci Med Sci, 59(4), 324-37. PMID: 15071075.

Luo, P., Tresini, M., Cristofalo, V., Chen, X., Saulewicz, A., Gray, M. D., Banker, D. E., Klingelhutz, A. J., Ohtsubo, M., Takihara, Y. & Norwood, T. H. (2004). Immortalization in a normal foreskin fibroblast culture following transduction of cyclin A2 or cdk1 genes in retroviral vectors. Exp Cell Res, 294(2), 406-19. PMID: 15023530.

Lee, J. H., Yi, S. M., Anderson, M. E., Berger, K. L., Welsh, M. J., Klingelhutz, A. J. & Ozbun, M. A. (2004). Propagation of infectious human papillomavirus type 16 by using an adenovirus and Cre/LoxP mechanism. Proc Natl Acad Sci U S A, 101(7), 2094-9. PMID: 14769917.

Morrison, T. E., Mauser, A., Klingelhutz, A. J. & Kenney, S. C. (2004). Epstein-Barr virus immediate-early protein BZLF1 inhibits tumor necrosis factor alpha-induced signaling and apoptosis by downregulating tumor necrosis factor receptor 1. J Virol, 78(1), 544-9. PMID: 14671137.

Morrison, J. A., Klingelhutz, A. J. & Raab-Traub, N. (2003). Epstein-Barr virus latent membrane protein 2A activates beta-catenin signaling in epithelial cells. J Virol, 77(22), 12276-84. PMID: 14581564.

Duffy, C. L., Phillips, S. L. & Klingelhutz, A. J. (2003). Microarray analysis identifies differentiation-associated genes regulated by human papillomavirus type 16 E6. Virology, 314(1), 196-205. PMID: 14517073.

Palanca-Wessels, M. C., Klingelhutz, A. J., Reid, B. J., Norwood, T. H., Opheim, K. E., Paulson, T. G., Feng, Z. & Rabinovitch, P. S. (2003). Extended lifespan of Barrett's esophagus epithelium transduced with the human telomerase catalytic subunit: a useful in vitro model. Carcinogenesis, 24(7), 1183-90. PMID: 12807723.

Allen, H., Klingelhutz, A. J. (2003). Papillomavirus infection and telomerase activation. Papillomavirus Report, 14(4), 155-162.

Zabner, J., Karp, P., Seiler, M., Phillips, S. L., Mitchell, C. J., Saavedra, M., Welsh, M. & Klingelhutz, A. J. (2003). Development of cystic fibrosis and noncystic fibrosis airway cell lines. Am J Physiol Lung Cell Mol Physiol, 284(5), L844-54. PMID: 12676769.

Mauser, A., Holley-Guthrie, E., Zanation, A., Yarborough, W., Kaufmann, W., Klingelhutz, A. J., Seaman, W. T. & Kenney, S. (2002). The Epstein-Barr virus immediate-early protein BZLF1 induces expression of E2F-1 and other proteins involved in cell cycle progression in primary keratinocytes and gastric carcinoma cells. J Virol, 76(24), 12543-52. PMID: 12438580.

Sprague, D. L., Phillips, S. L., Mitchell, C. J., Berger, K. L., Lace, M., Turek, L. P. & Klingelhutz, A. J. (2002). Telomerase activation in cervical keratinocytes containing stably replicating human papillomavirus type 16 episomes. Virology, 301(2), 247-54. PMID: 12359427.

Martin, J. A., Forest, E., Block, J. A., Klingelhutz, A. J., Whited, B., Gitelis, S., Wilkey, A. & Buckwalter, J. A. (2002). Malignant transformation in human chondrosarcoma cells supported by telomerase activation and tumor suppressor inactivation. Cell Growth Differ, 13(9), 397-407. PMID: 12354749.

Martin, J. A., Mitchell, C. J., Klingelhutz, A. J. & Buckwalter, J. A. (2002). Effects of telomerase and viral oncogene expression on the in vitro growth of human chondrocytes. J Gerontol A Biol Sci Med Sci, 57(2), B48-53. PMID: 11818423.

Sood, A. K., Coffin, J., Jabbari, S., Buller, R. E., Hendrix, M. J. & Klingelhutz, A. J. (2002). p53 null mutations are associated with a telomerase negative phenotype in ovarian carcinoma. Cancer Biol Ther, 1(5), 511-7. PMID: 12496479.

Farwell, D. G., Shera, K. A., Koop, J. I., Bonnet, G. A., Matthews, C. P., Reuther, G. W., Coltrera, M. D., McDougall, J. K. & Klingelhutz, A. J. (2000). Genetic and epigenetic changes in human epithelial cells immortalized by telomerase. Am J Pathol, 156(5), 1537-47. PMID: 10793065.

Klingelhutz, A. J. (1999). The roles of telomeres and telomerase in cellular immortalization and the development of cancer. Anticancer Res, 19(6A), 4823-30. PMID: 10697595.

Kiyono, T., Foster, S. A., Koop, J. I., McDougall, J. K., Galloway, D. A. & Klingelhutz, A. J. (1998). Both Rb/p16INK4a inactivation and telomerase activity are required to immortalize human epithelial cells. Nature, 396(6706), 84-8. PMID: 9817205.

Anderson, S., Shera, K., Ihle, J., Billman, L., Goff, B., Greer, B., Tamimi, H., McDougall, J. & Klingelhutz, A. J. (1997). Telomerase activation in cervical cancer. Am J Pathol, 151(1), 25-31. PMID: 9212727.

Klingelhutz, A. J. (1997). Telomerase activation and cancer. J Mol Med (Berl), 75(1), 45-9. PMID: 9020383.

Klingelhutz, A. J., Foster, S. A. & McDougall, J. K. (1996). Telomerase activation by the E6 gene product of human papillomavirus type 16. Nature, 380(6569), 79-82. PMID: 8598912.

Klingelhutz, A. J., Hedrick, L., Cho, K. R. & McDougall, J. K. (1995). The DCC gene suppresses the malignant phenotype of transformed human epithelial cells. Oncogene, 10(8), 1581-6. PMID: 7731713.

Klingelhutz, A. J., Barber, S. A., Smith, P. P., Dyer, K. & McDougall, J. K. (1994). Restoration of telomeres in human papillomavirus-immortalized human anogenital epithelial cells. Mol Cell Biol, 14(2), 961-9. PMID: 8289836.

Klingelhutz, A. J., Smith, P. P., Garrett, L. R. & McDougall, J. K. (1993). Alteration of the DCC tumor-suppressor gene in tumorigenic HPV-18 immortalized human keratinocytes transformed by nitrosomethylurea. Oncogene, 8(1), 95-9. PMID: 8380923.

Klingelhutz, A. J., Wu, S. Q., Huang, J. & Reznikoff, C. A. (1992). Loss of 3p13----p21.2 in tumorigenic reversion of a hybrid between isogeneic nontumorigenic and tumorigenic human uroepithelial cells. Cancer Res, 52(6), 1631-4. PMID: 1311637.

Klingelhutz, A. J., Wu, S. Q. & Reznikoff, C. A. (1991). Nonrandom chromosome losses in tumorigenic revertants of hybrids between isogeneic immortal and neoplastic human uroepithelial cells. Somat Cell Mol Genet, 17(6), 551-65. PMID: 1767334.

Klingelhutz, A. J., Wu, S. Q., Bookland, E. A. & Reznikoff, C. A. (1991). Allelic 3p deletions in high-grade carcinomas after transformation in vitro of human uroepithelial cells. Genes Chromosomes Cancer, 3(5), 346-57. PMID: 1686726.

Wu, S. Q., Storer, B. E., Bookland, E. A., Klingelhutz, A. J., Gilchrist, K. W., Meisner, L. F., Oyasu, R. & Reznikoff, C. A. (1991). Nonrandom chromosome losses in stepwise neoplastic transformation in vitro of human uroepithelial cells. Cancer Res, 51(12), 3323-6. PMID: 2040006.

Reznikoff, C. A., Kao, C., Bookland, E. A., Klingelhutz, A. J., Pratt, C. I., Wu, S. Q., Gilchrist, K. W. & Swaminathan, S. (1991). Transformation in vitro of human uroepithelial cells. New Jersey: Humana Press.

Reznikoff, C. A., Bookland, E. A., Klingelhutz, A. J., Pratt, C. I., Kao, C., Bhattacharya, M., Swaminathan, S. & Meisner, L. F. (1989). SV-HUC-1: A human urinary tract epithelial cell line for multistep in vitro transformation studies. England: Adam Hilger Press.