Mark Steinberg


Main Affiliation

Chemistry and Biochemistry

Additional Departments/Affiliated Programs

MARC Program

Areas of Expertise/Research

  • Chemistry


Marshak Science Building



Mark Steinberg

Mark Steinberg


After finishing his doctoral work at the University of Pennsylvania Dr. Steinberg did postdoctoral research at the NYU school of medicine where he began his research on human epithelial cell model systems. His group was the first to document transformation and immortalization of human epidermal cells by the oncogenic virus, SV40 (PNAS 76:801, 1979). After joining the Department of Chemistry at City College in 1986 he has received a number of NIH grants to extend this work to include other carcinogenic stimuli including organic carcinogens, arsenic and ultraviolet radiation.
Dr. Steinberg is director of the NIH-funded RISE program at City College. The purpose and goal of the RISE program is to help students from underrepresented minority groups enter doctoral programs, and ultimately professional careers, in the biomedical and behavioral sciences.  Towards this end the RISE program provides funding to help defray the financial burden of paying for college as a full time student.  The program also provides money for extracurricular training including attendance at scientific conferences.


University of Michigan           B.A.     1966   Psychology
University of Pennsylvania    Ph.D.   1976   Biology

Courses Taught

Chem 48005/A8005: Advanced Biochemistry

Chem 45902: Biochemistry 1

Chem 10301: General Chemistry 1

Research Interests

Our laboratory utilizes an in vitro model system of human epidermal keratinocytes to study oncogenic gene expression in response to various oncogenic stimuli. There are three main areas of research:

1. Transcriptional activation of the cyclin D1 promoter in response to UVB irradiation and exposure to arsenic.

2. To study oncogenic gene expression in SV40-transformed human keratinocytes using RNA interference techniques to silence the SV40 early genes.

3. The role of reactive oxygen species (ROS) related to UVB induced deletions of mitochondrial DNA in the process of altered gene expression in cellular senescence and aging.


1.  Steinberg, M.L. and Whittaker, J.R.: Stimulation of melanotic expression in a melanoma cell line by theophyllline. J. Cell. Physiol., 87:265-275, 1976.

2.  Steinberg ,M.L. and Whittaker, J.R.: Theophylline incorporation into the nucleic acids of theophylline-stimulated melanoma cells.  J. Invest. Derm., 71:250-256, 1978.

3.  Steinberg, M.L. and Defendi, V.: Altered pattern of growth and differentiation in human keratinocytes infected by SV40. Proc. Natl. Acad. Sci., 76:801-805, 1979.

4.  Wolman, S.R., Steinberg, M.L. and Defendi, V.: Simian virus 40-induced chromosome changes in human epidermal cultures. Cancer Genet. and Cytogenet., 2:39-46, 1980.

5.  Steinberg, M.L. and Defendi V.: Patterns of cell communication and differentiation in SV40 transformed human keratinocytes. J. Cell. Physiol., 109:153-159, 1981.

6.  Steinberg, M.L. and Defendi, V.: Fusion induced differentiation of SV40 transformed human keratinocytes. Exp. Cell Res., 139:369-375, 1982.

7.  Mufson, R.A., Steinberg, M.L. and Defendi, V.: The effects of 12-O-tetradecanoyl phorbol-13-acetate on the differentiation of SV40-infected human keratinocytes. Cancer Res., 42:4600-4605, 1982.

8.  Friedman, E.A. and Steinberg, M.L.: Disrupted communication between late-stage premalignant human colon epithelial cells by 12-O-tetradecanoyl-13-acetate. Cancer Res., 42:5096-5102, 1982.

9.  Defendi, V., Naimski, P., and Steinberg, M.L.: Human cells transformed by SV40 revisited: The Epithelial Cells. J. Cell. Physiol., Suppl. 2:131-140, 1982.

10. Steinberg, M.L., Cassai, N. and Defendi, V.: A scanning electron microscope study of normal and SV40-infected human keratinocytes. Scanning Electron Microscopy, Vol. I pt.1:343-349, 1983.
11. Steinberg, M.L. and Defendi, V.: Transformation and immortalization of human keratinocytes by SV40. J. Invest. Derm., 81(1):131s-136s, 1983.

12. Okada, N., Steinberg, M.L. and Defendi, V.: Re-expression of differentiated properties in SV40-infected human epidermal keratinocytes induced by 5-azacytidine.  Exp. Cell Res., 153:198-207, 1984.

13. Hronis, T.S., Steinberg, M.L., Defendi, V. and Sun T.-T.: The simple epithelial nature of some simian virus- 40 transformed human epidermal keratinocytes. Cancer Res., 44:5797-5804, 1984.

14. Edelman, B., Steinberg, M.L. and Defendi, V.: Changes in fibronectin synthesis and binding distribution in SV40-transformed human keratinocytes.  Int. J. Cancer., 35(2):219-225, 1985.

15. Steinberg, M.L. and Defendi, V.: Altered patterns of keratin synthesis in human epidermal keratinocytes transformed by SV40. J. Cell. Physiol., 123:117-125, 1985.

16. Naimski, P. and Steinberg, M.L.: Analysis of stable and unstable viral forms in SV40 infected human keratinocytes., J. Cell. Bioch., 29:95-103, 1985.
17. Morris, A., Steinberg, M.L. and Defendi, V.: Keratin gene expression in SV40 transformed human keratinocytes. Proc. Natl. Acad. Sci., 82:8498-8502, 1985.

18. Steinberg, M.L., Morris, A. and Goodman, S.: Oncogenic properties of human epidermal keratinocytes transformed by SV40. In: Processes in Cutaneous Epidermal Differentiation, I.A. Bernstein and T. Hirone, eds. Praeger Scientific, New York, N.Y. 1986. pp. 287-299.

19. Okada, N. Kitano, Y. Miyagawa, S., Sakimoto, K. and Steinberg, M.L.: Expression of pemphigoid antigen by SV40 transformed human keratinocytes. J. Invest. Derm., 86:399-401, 1986.

20. Baden, H.P., Kubilus, J., Kvedar, J.C., Steinberg. M.L. and Wolman, S.R.: Isolation and characterization of a spontaneously arising long-lived line of human keratinocytes (NM 1). In Vitro, 23(3):205-213, 1987.

21. Rossman, T.G., Rubin, L.M., Gilmer, T, and Steinberg, M.L.: Heterogeneity in the response of inserted viral sequences to carcinogen treatment: a clonal approach. in: Accomplishments in Oncology Vol. II no.1: The Role of DNA amplification in Carcinogenesis. H. Zur Hausen and J. Schlehoffer eds. Lippincott, Philadelphia, 1987. pp.83-94.

22. Baden, H.P., Kubilus, J., Wolman, S.R., Steinberg, M.L. and Philips, S.B.: The NM1 keratinocyte line is cytogenetically and biologically stable and exhibits a unique structural protein.  J. Invest. Derm., 89:574-579, 1987.

23. Miyagawa, S., Okada, N., Kitano, Y., Sakamoto, K. and Steinberg, M.L.: SSA/Ro antigen expression in SV40-transformed human keratinocytes.  J. Invest. Derm. 90(3):342-345, 1988.
23. Okada, N., Miyagawa, S., Hiriguchi, Y. Kitano, Y. Yoshikawa, K., Sakamoto, K. and Steinberg, M.L.: Synthesis of epidermolysis bullosa acquisita antigen by simian virus 40-transformed human keratinocytes. Arch. Derm. Res. 281:1-4, 1989.

24. Morris, A.E., Steinberg, M.L., Li, J.-Y. and Defendi, V.: Isolation of novel cDNA transformation markers from SV40 transformed human keratinocytes. Exp. Cell Res. 182:461-472, 1989.

25. Miyagawa, S., Okada, N., Takasaki, Y., Takashi, I., Kitano, Y. Yoshikawa, K., Sakamoto, K. and Steinberg, M.L.:Expression of proliferating cell nuclear antigen/cyclin in human keratinocytes. J. Invest. Derm. 93(5):678-681, 1989.

26. Steinberg, M.L., Rossman, T.G., Morris, A. and Chen, G.: Specific high frequency rearrangements induced by MNNG in SV40-infected human keratinocytes. Carcinogenesis, 10(10):1801-1807, 1989.                 

27. Staiano-Coico, L., Steinberg, M., and Higgins, P.J.: Butyrate-induced terminal differentiation in normal and SV40-transformed human keratinocytes: regulation of the actin-based cytoskeleton and production of cornified envelopes.  Int. J. of Cancer, 46:733-738, 1990.

28. Okada, N., Miyagawa, S., Steinberg, M.L. and  Yoshikawa, K.: Proliferating cell nuclear antigen/cyclin in cultured human keratinocytes. J. of Derm., 17(9):521-525, 1990.

29. W. Sha Glassman, Steinberg, M. and Alfano, R.R.:Steady state and time resolved fluorescence spectroscopy from normal and malignant human breast cell lines.  Lasers in the Life Sciences, 6(2):91-98, 1994.

30. Chen,W., Ruml, T. and Steinberg, M.L.: Characterization of a 1.8 kb DNA segment located upstream from the human Ha-ras protooncogene and possibly regulating its function.  Gene, 155(2):253-256, 1995.

31. Guangtian Chen, G.T, Baez, J. and Steinberg, M.: integration of the SV40 promoter/enhancer sequences in an anchorage independent clonal subline of SV40-infected human keratinocytes. Gene, 183:41-45, 1996.

32. Papadopoulos, A.J., Zhadin, N.N., Steinberg, M.L. and Alfano, R.R.: Fluorescence spectroscopy of normal, SV40-transformed human keratinocytes and carcinoma cells.  Cancer Biochemistry Biophysics, 17:13-23, 1999.

33. Forman, S., Kas, J., Fini, F., Steinberg, M. and Ruml, T.: The effect of different solvents on the ATP/ADP content and growth properties of HeLa cells. Journal of Biochemical and Molecular Toxicology, 13(1):11-15, 1999.

34. Vazquez, R. and Steinberg, M.L.: Enhancement of Genomic DNA Templates for Polymerase Chain Reactions by Partial Restriction Digestion. Biotechniques 26:91-95, 1999.

35. M. Steinberg, G. T. Chen, R. Vazquez, T.Ruml, J. Kas: SV40-immortalized human keratinocytes as an in vitro model system for studying environmental carcinogens.  International Biodeterioration and Bioremediation, 44(1):7-16, 1999.

36. M. Steinberg and R. Vazquez: SV40-immortalized human keratinocytes as an in vitro model system to analyze nucleotide alterations caused by alkylating agents. Proceedings of the 2000 Medical Defense Bioscience Review. vol. II, pp. 907-908.

37. R. Vazquez, R., Nelson, M., Guzman, J., Corun, C., Graham, J., Hwang, BJ., Otoo, H., and Steinberg, M: Analysis of the anti-inflammatory effects motrin (ibuprofen) has on SV40-immortalized keratinocytes (IHEK) exposed to the chemical warfare agent sulfur mustard (HD).   Proceeding of the 2002 Medical Defense Bioscience Review. p.216, 2002.

38. Vazquez R. and M. Steinberg: SV40-immortalized Human Keratinocytes as an In Vitro Model   System to Analyze Sequence Specific Nucleotide Alterations Caused by Alkylating Agents. J.Burns 2002:II;No.3.:17-24

39. R. Vazquez, R., Nelson, M., Guzman, J., Corun, C., and  Steinberg, M: Immortalized Human Keratinocytes: A Model System to Study the Efficacy of Therapeutic Drugs in Response to the Chemical Warfare Agent Sulfur Mustard (HD). Electronic Journal of Biotechnology, 7(2):124-129, 2004.

40. Qabar, A. Nelson, M. Guzman, J., Charlene, C., Hwang, B.-J.,, and Steinberg, M.: Modulation of Sulfur Mustard Induced Cell Death in Human Epidermal Keratinocytes Using IL-10 and TNF-α. J.Biochem Molecular Toxicology, 19(4):213-225  2005

41. Hwang, B.-J., Utti, C., and Steinberg, M.: Induction of cyclin D1 by submicromolar concentrations of arsenite in human epidermal keratinocytes. Toxicol. Appl. Pharm. 217: 161-167, 2006.

42. Fang, J., Pierre, Z., Liu, S., Hwang, B.-J., Hill, H.Z., Hubbard, K., and Steinberg, M. Novel mitochondrial deletions in human epithelial cells irradiated with an FS20 ultraviolet light source in vitro. J. Photochem. Photobiol. 184:340–346, 2006.

43. Hubbard,K, Steinberg, Mark L, Hill, HZ, Orlow, I. Mitochondrial DNA deletions in skin from melanoma patients.  Ethnicity and Disease, 18(S2):38-43, 2008.  PMID: 19161404

44. Steinberg, ML, Hwang, B-J, Tang, L, Shah, MA. E cadherin gene alterations in gastric cancers in different ethnic populations.  Ethnicity and Disease, 18(S2):70-74, 2008.    PMID: 18646324

45. Steinberg, M. L., Hubbard, K., Utti, C., Clas, B., Hwang, B-J., Hill, H.Z., Orlow, I. Patterns of persistent DNA damage associated with sun exposure and the glutathione S-transferase M1 genotype in melanoma patients.  Photochem. Photobiol. 85: 379-386, 2009.  PMID: 19161404, PMCID: PMC2743971

46. Hwang, B-J. Kuttamperoor, F., Wu, J. and Steinberg, M.L. Spectrum of mitochondrial DNA deletions within the Common Deletion region induced by low levels of UVB irradiation of human keratinocytes in vitro. Gene 440: 23-27, 2009.
PMID: 19345726, PMCID: PMC2706008

47.  Suqing Liu, Julian Gonzalez, Bor-Jang Hwang and Mark L Steinberg. Induction of Cyclin D1 by Arsenite and UVB-irradiation in Human Keratinocytes. Journal of Health Care for the Poor and Underserved Volume 22, Number 4, November 2011, Supplement pp. 110-121   E-ISSN: 1548-6869 Print ISSN: 1049-2089

Book Chapters

Steinberg, M. Immortalization of Human Epidermal Keratinocytes by SV40. in: Culture of Immortalized Cells. Chapter 5, pp. 95-120,  Ian Freshney ed., Wiley-Liss, New York, 1996

Steinberg, Mark L. Mitochondrial DNA Biomarkers in Melanoma. In: Diagnostic and Prognostic Biomarkers and Therapeutic Targets in Melanoma. Chapter 11, pp. 145-157, Michael J. Murphy ed., Humana Press, New York, NY, 2012 ISBN: 9781607614326 

Reference Textbooks

Steinberg, M., Guyden, J., Calhoun, D., Coico, R. and Staiaino-Coico, L. "Recombinant DNA Technology: Concepts and Biomedical Applications", Prentice Hall, Englewood Cliffs, New Jersey Inc., 1993

Steinberg, M. and Cosloy, S. "The Facts on File Dictionary of Biotechnology and Genetic Engineering", 3nd edition  Facts on File, Inc., New York, NY, 2006

Professional Experience

1992-Present   Professor, Dept. of Chemistry & Biochemistry, The City College of New York, New York, N.Y.

1986-2000   Adjunct Professor of Pathology, N.Y.U Medical Center, New York, N.Y.

1988-1991   Associate Professor, Dept. of Chemistry, The City College of New York, New York, N.Y.

1986-1988   Assistant Professor, Dept. of Chemistry, The City College of New York, New York, N.Y.

1982-1986   Assistant Professor, Dept. of Pathology, New York University Medical Center, New York, N.Y

1979-1982   Research Assistant Professor, Dept. of Pathology, N.Y.U. Medical Center, New York, NY

1976-1979  NCI,NRSA Postdoctoral fellowship, Dept. of Pathology, N.Y.U. Medical Center, New York, NY