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CUKRICU UJM VITAE Name: Peter M. Howley, M.D. t J Date and Place of Birth: October 9, 1946; i:ev Brunswick, New Jerscy .,Citizenship: United States `I"farital Status: Harried, 1969; three Education: children June 1964 - Graduated from High School June 1968 - A.B. (Chemistry), Princeton University June 1970 - M.M.S., Rutgers-The State University ,lune 1972 - M.D., Harvard University Board Certification: Anatomic Pathology, May 1977 Brief Chronology of Employment: 1972-1973 - Intern, Pathology, Massachusetts General Hospital ~ 1973-1975 - Research Associate, National Institute of Allergy and 1975-1976 Infectious Diseases, National Institutes of Health Bethesda, 14aryland - Resident, Laboratory of Pathology, National Cancer 1976-1977 Institute, National Institutes of Health, Bethesda, t•1aryland - Junior Staff Pathologist, Laboratory of Pathology, 1977-1979 National Cancer Institute, National Institutes of Health, Bethesda, Maryland - Senior Investigator, Laboratory of Pathology, National Cancer Institute, National Institutes of Health, Bethesda, `laryland ~1979-date - Chief, Viral Oncology and Molecular Pathology Sectior., Laboratory of Pathology, National Cancer Institute, Na:ional Institutes of Health, Bethesda, tlaryland Military Service: July 1, 1973-date -.Senior Surgeon, USPAS Societies: Sigma Xi; American Association of Pathologists; American Society of tticrobiology; Association of University Pathologists (Pluto Clu`)); American Society of Virology Honors and Other Special Scientific Recognition: Sigma Xi; USPHS Commendation Award (1980) Editorial Board: Journal of Virology Research Interests: Hur.,an papovaviruses; DNA tumnr viruses; Fapillomaviruses; Patho;;enesis o! viral diseases; Recombinant DNA and the introduction of genetic infor-,ation into cul:aryotic cells Present Address: 5522 Oal: Place Bethesda, 1laryland 20317 65100 610=1 CTR HN 0 110 1 d2_

BI BLIOG Rf'1PHY 1. Kirkpatrick, J.B., Hyams, L., Tho©as, V.L., and Howley, P.`1.: Puri- fication of intact microtubules from brain. J. Cell Biol. 47: 384- 394, 1970. 2. Jones, M., Ando, W., Hendrick, M.E., Kulczycki, A., Howley, P.M., Hurnmel, K.F., and Malament, D.S.: Irradiation of methyl diazomalonate in solution. Reactions of singlet and triplet carbenes with carbon- carbon double bonds. J. Am. Chem. Soc. 94: 7469-7479, 1972. 3. Khoury, G., Howley, P.M., Brown, M., and Martin, M.: The detection and quantitation of SV40 nucleic acid sequences using single-stranded SV40 DNA probes. Cold Spring Harbor Symp. on Quant. Biol. 39: 147-152, 1974. 4. Howley, P.M., Mullarkey, M.F., Takemoto, K.K., and Martin, M.A.: Characterization of human papovavirus BK DNA. J. Virol. 15: 173-131, 1975. 5. Khoury, C., Howley, P.M., Nathans, D., and Martin, M.A.: Post-transcriptional selection of SV40-specific RNA. J. Virol. 15: 433-437, 1975. 6. Khoury, G., Howley, P.M., Caron, C., Mullarkey, '1.F., Takemoto, K.K., and Martin, 11.A.: Homology and relationship between the genomes of papovaviruses, BKV and S'J40.- Proc. Nat. Acad. Sci. USA 72: 2563-2567, 1975. 7. Howley, P.M., Khoury, G., Byrne, J.C., Takemoto, K.K., and Martin, ?:.A.: A physical map of the BKV genome. J. Virol. 16: 959-973, 1975. 8. Khoury, G., Carter, B.J., Ferdinand, F.-J., Howley, P.M., Brown, `1., and Martin, M.A.: Genore localization of simian virus 40 RNA species. J. Virol. 17: 832-840, 197b. 9. Martin, M.A., Howley, P.`:., Byrne, J.C., and Garon, C.F.: Characterization of supercoiled, oligometric SV40 DNA raolecules in productively infected cells. Virology 71: 26-40, 1976. 10. Howley, P.H., Khoury, G., Takemoto, K.K., and Martin, M.A.: Po1yn::cleo- tide sequences common to the genomes of simian virus 40 and the human papovaviruses JC and BK. Virology 73: 303-307, 1976. 11. Davoli, D., Canem, D., Nussbaum, A.L., Fareed, G.C., Hoaley, P.t:., Khoury, C., and Martin, M.A.: Genome structures of reiteration cutants of simian virus 40. Virology 77: 110-124, 1977. 12. Costa, J.C., ilowley, P.!:., Legallais, F.Y., Yee, C.L., Young, N.A., and Rabson, A.S.: Oncogenicity of a nude mouse cell line transformed by a human papovavirus. J. ?:atl. Cancer Inst. 58: 1147-1150, 1977. 13. Lucky, A.1+'., Howley, P.`:., Megysi, K., Spielberg, S.P., and Schulman, J.D.: Endocrine studies in cys:inosis: compensated primary hypothyroidisM. J. Pediatrics 91: 204-210, 1977. 6--100 6106 CTRHN 011013

14. Howley, P.M., and Martin, M.A.: Uniform representation of the human papovavirus BK genome in transformed hamster cells. J. Virol. 23: 205-208, 1977. 15. Holmberg, C.A., Gribble, D.H., Takemoto, K.K., Howley, P.M., Espana, C., and Osburn, B.I.: Isolation of simian virus 40 from rhesus monkeys (Hacaca mulatta) with spontaneous progressive multifocal leukoencephalopathy. J. Infect. Dis. 136: 593-596, 1977. 16. Khoury, G., Lai, C.-J., Solomon, D., Israel, M.A., and Howley, P.t;.: The human papovaviruses and their potential role in human diseases. Cold Spring Harbor Symposium - Origins of Human Cancer. Cold Spring llarbor, New York, 1977, pp. 971-988. 17. Takemoto, K.K., Solo3on, D., Israel, M.A., Howley, P.!1., Khoury, G., Aaronson, S.A., and Martin, M.A.: Search for evidence of papovavirus involvement in human cancer. In Bentvelzen, et al. (Eds.): Advances in Comparative Leukemia Research 1977. Elsevier/North-Holland Biomedical Press, 1978, pp. 260-263. 18. Bond, S.B., Howley, P.M., and Takemoto, K.K.: Cha.acterization of K virus and its comparison with polyoma virus. J. Virol. 28: 337- 343, 1978. 19. Khoury, C., Howley, P.M., Israel, M.A., Brown, D., and Lai, C.-J.: The papovaviruses: Their molecular biology and potential etiology in hum3n disease. In Allison, F., Jr. and Rothschild, F. (Eds.): liunan Diseases Caused by Cancer: Recent Developments. New York, Oxford Univ. ?ress, 1978, pp. 191-214. 20. Israel, M.A., Takemoto, K.K., Martin, 1;.A., Sclomon, D., Howley, P.?1., Aaronson, S.A., and K!:osrv, G.: Evaluation of normal and neoplastic human tissue for BK virus. Virology 90: 187-196, 1978. 21. Howley, P.M., Newell, N., Shah, K.V., Law, M.-F., Gruss, P., Sauer, G., and Kelly, T.J., Jr.: identification of the primate papovavirus HD as the stumptailed 1lacaque virus. J. Virol. 30: 400-403, 1979. 22. Law, M.-F., Takemoto, K.K., and Howley, P.ti.: Characterization of the genome of the murine papovavirus K. J. Virol. 30: 90-97, 1979. 23. Takemoto, K.K., Howley, P.M., and Miyamura, T.: JC human papovavirus replication in human amnion cells. J. Virol. 30: 334-389, 1979. 24. Howley, P.M., Israel, M.A., Law, M.-F., and Martin, M.A.: A rapid method for detecting and mapping homology between heterologuus Dh.4s. Evaluation of polyomavirus genomes. J. Biol. Checn. 254: 487b-4333, 1979. 25. Law, M.-F., Martin, J.D., Takemoto, K.K., and Howley, P.M.: The co-linear alignment of the genomes of papovaviruses JC, BK, and SV40. Virology 96: 576-587, 1979. 651()0 cle7 CTR Ht'°t 011014

-3- 26. Law, M.-F., Lancaster, h.D., and Howley, P.M.: Conserved polynucleo:ide sequences among the genomes of papillomaviruses. J. Virol. 32: 199-207, 1979. 27. Howley, P.M.: The molecular biology of SV40 and the human papovaviruses BK and JC. In Klein, C. (Ed.): Viral Oncology. New York, Raven Press, 1980, pp. 489-550. 28. Howley, P.M.: The DNA sequence of the human polyomavirus BK. (News and Views). Nature 284: 124-125, 1980. 29. Lowy, D.R., Dvoretzky, I., Shober, R., Law, M.-F., Engel, L.W., and Howley, P.M.: In vitro tumorigenic transformation by a defined subgeno-mic fragment of bovine papilioma virus DNA. Nature 287: 72-74, 1980. 30. Howley, P.M., Law, M.-F., Heilman, C., Engel, L., Alonso, M.D., Lancaster, W.D., Israel, M.A., and Lowy, D.R.: tiolecular characterizatiorn of papillomavirus genomes. In Essex, M., Todaro, C., and zur Hausen, H. (Eds.): Viruses in Naturally Occurring Cancers. Cold Spring Harbor, New York, Cold Spring Harbor Laboratory, 1980, pp. 233-247. 31. Stromberg, K., Donner, L., Yee, C.L., and Howley, P.M.: Growth potential of sheep and sea mammal cells transformed by SV40 early region DNA. Proc. Soc. Exp. Biol. Med. 164: 439-444, 1980. 32. Heilman, C.A., Law, *1.-F., Israel, M.A., and Howley, P.M.: Cloning of human papillomavirus genomic DNAs and analysis of homologous polynucleotide sequences. J. Virol. 36: 395-407, 1980. 33. Howley, P.M., Rentier-Deirue, F., Heilman, C.A., Law, M.-F, Chowdhury, K., Israel, P:.A., and Takemoto, K.K.: Cloned human polyomavirus JC D::n car, transform human amnion cells. J. Virol. 36: 878-882, 1980. 34. Costa, J. C. , Howley, Bowling, M. C. , Howard, R., and Bauer, 1+.C. : Presence of human papilloma viral antigens in juvenile multiple larynSeal papillomas. Am. J. C=i n. Pathol. 75: 194-197, 1981. 35. Goldman, N.D., Howley, and Khoury, G.: Functional interaction batwecn the early viral proteins of simian virus 40 and adenovirus. Virology 109: 303-313, 1981. 36. Law, M.-F., Lowy, D.R., Dvoretzky, I., and Howley, P.t1.: House cells ,rar.s- formed by bovine papillomavirus contain only episomal viral DNA sequences. Proc. Nat. Acad. Sci. USA 78: 2727-2731, 1981. 37. Sarver, N., Gruss, P., law, M.-F., Khourv, C, and lio::ley, P.M.: Bovine papilloma virus DtiA - A novel eukarvotic clonini, vl-~,tor . t:oie;.. E-3/. 1: 486-496, 1981. 65100 6108 CTR RtN 011015

-4 - 38. Rentier-Delrue, F., Lubiniecki, A., and Howley, P.M.: Analysis of JC virus DNA purified directly from human P"SL brains. J. Virol. 38: 761-769, 1981. 39. Patch, C.T., Howley, P.M., Hauser, J., and Levine, A.S.: Kinetics of inhibition. of papovavirus DNA synthesis by superinfection with adenovirus 2 and non-defective adenovirus 2 - simian virus 40 hybrid viruses. J. Gen. Virol. S5: 355-365, 1981. 40. Sarver, N., Gruss, P., Law, M.-F., Khoury, C., and Howley, P.M.: Rat insulin gene covalently linked to bovine papillomavirus DNA is expressed in transformed mouse cells. In Brown, D. and Fox, C.R. (Eds.): ICN- UCLA Symposia on Molecular and Cellular Biology. Vol. XXIII, Developrien:al Biology Using Purified Genes. New York, Academic Press, 1981, pp. 547-556. 41. Heilman, C.A., Engel, L.W., Lowy, D.R., and Howley, P.M.: Virus-specific transcription in bovine papillomavirus-transformed mouse cells. Virology 119: 22-34, 1982. 42. Law, M.-F., Howard, B., Sarver, N., and Howley, P.M.: Expression of selective traits in mouse cells transformed with a BPV DNA derived hybrid molecule containing E. coli _gpt. In Cluzman, Y. (Ed.): Eukaryotic Viral Vectors. Cold Spring Harbor, New York, Cold Spring Harbor Laboratory, 1982, pp. 79-86. 43. Howley, P.M.: The human papillomaviruses. (Special Article). Arch. Pathol. Lab. Med. 106: 429-432, 1982. 65100 6109 CTR HN 011016

ABSTRACTS AND PRESENTATIONS i , 1. Mullarkey, M.F., }lovley, P.M., Martin, M.A, Tal:erroto, K.K., Lee, T.h.H., and Nathans, D.: Genetic material of BK virus: A physical characteri- zation and comparison with SV40. American Society for Microbiology • Meeting, 1974. 2. Khoury, C., Howley, P.M., and Martin, H.A.: The detection and quanti:ation of SV40 genetic material. Cold Spring Harbor Symposium on Quantitative Biology (Tumor Viruses), 1974. 3. Howley, P.M., and Martin, M.A.: Human papovavirus BK: A physical map of the virus genome and quantitation of viral DNA sequences in transformed hamster cells. Sixth Tumor Virus Meeting. Cold Spring Harbor Laboratory, Cold Spring Harbor, New York, 1975. 4. Howley, P.M., Khoury, C., Takemoto, K.K., Mullarkey, M.F., and Martin, K.A.: Human papovavirus-BK DNA: Characterization and polynucleotide sequence homology to SV40 DNA. Fed. Proc. 34: 526, 1975. 5. Martin, M.A., and Howley, P.M.: Oligomeric forms of SV40 DNA in lytically infected cells. Sixth Tumor Virus Meeting. Cold Spring Harbor Laboratory, Cold Spring Harbor, New York, 1975. 6. Holmberg, C.A., Gribble, D.H.,.Takemoto, K.K., and Howley, P.M.: Papova- virus isolates from progressive multifocal leukoencehalopathy in Hacaques. Fed. Proc. 35: 570, 1976. 7. Lucky, A.W., Megyesi, K., Howley, P.M., and Schulman, J.D.: Endocrine function in cystinosis. The Society for Pediatric Research, 1976. 8. Howley, P.M., Khoury, G., Takemoto, K.K., and tlartin, M.A.: Polynucleotide sequences common to the genomes of SV40 and the human papovaviruses JC and BK. Seventh Tucor V_rus Meeting, Cold Spring Harbor Laboratory, Cold Spring Harbor, ":ew York, 1976. 9. Law, M.-F., Takemoto, K.K., and Howley, P./4.: Characterization of the D::A of K virus, a murine papovavirus. American Society for i•ticrobioloby, Las Vegas, Nevada, May 8-12, 1975. 10. Howley, P.M., Law, M.-F., and Israel, *1.A.: Detection of polynucleotide sequences shared among the genomes of papovaviruses. American Society for Microbiology, Las Vegas, Nevada, May 8-12, 1978. 11. Law, M.-F., Takemoto, K.K., and Howley, P.`1.: Characterization of the genome of Y, virus, a murine papovavirus. DNA Tumor Virus `teeting, Colci Spring Harbor Laboratory, Cold Spring Harbor, New York, July 26-3U, 1973. 12. Law, !1.-F., Lowy, D.R., Dvoretsky, I . , and Ilouley, P.M.: Transformat ion of mouse cells by cloned BPV-1 DNA segments and analysis of the state o: the viral genome. Tenth Tumor Virus Meeting, Cold Spring Harbor Laboratorv, Cold Spring Harbor, New York, August 13-17, 1980. 65100 611C CTR HN 0 110 Vt"

ABSTRACTS AND PRESENTATIONS (Cont'd) 13. Heilman, C.A., Law, M.-F., and Howley. P.M.: Clonin;, of huuan papilloma- virus genomic DNAs and analysis of homologous polynucleotide sequences. Tenth Tumor Virus Meeting, Cold Spring Harbor Laboratory, Cold Sprin.- •' Harbor, New York, August 13-17, 1980. 14. Rentier-Delrue, F., Israel, M.A., Takemoto, K.K., and Howley, P.M.. Molecular cloning of the human polyomavirus JC genome. Tenth Tunor Virus Meeting, Cold Spring Harbor Laboratory, Cold Spring Harbor, New York, August 13-17, 1980, p. 74. 15. Sarver, N., Rosenbaum, S., Law, M.-F., Byrne, J.C., and Howley, P.`1.: A bovine papillomavirus - p:4L2 hybrid vector which replicates as a plasmid in mouse cells and in bacteria. Workshop on Gene Transfer and Cancer, Frederick, Maryland, April 16-18, 1982. 16. Law, :1.-F. , Howard, B.:i. , Yang, Y.-C. , and Howley, P.t:. : The use of dominant selective markers in the BPV-1 vector system. Papilloma Viruses t(ecting, Cold Spring Harbor Labcratory, Cold Spring Harbor, New York, September 14-18, 1982. 17. Banks-Schlegel, S., and Howley, P.M.: Differentiation of human epidermal cells transformed by SV40. Papilloma Viruses Meeting, Cold Spring Harbor Laboratory, Cold Spring Harbor, New York, September 14-18, 1982. 18. Sarver, N., Byrne, J.C., and llowley, P.M.: Transformation and replication in mouse cells of a bov4ae papillonavirus/pifL2 plasmid vector that cau be rescued in bacteria. Papilloma Viruses Meeting, Cold SprinL .arbor Laboratory, Cold Spring Harbor, New York, September 14-18, 1962. 19. Graham, G.M., Babiss, l.•--., Byrne, J.C., Howley, P.M., and Fishcr, P.B.: Transformation of Fischer rat embryo (CREF) cells by bovine papi•11o•-~a virus (BYV) and plasmid cloned BPV DNA. Papilloma Viruses Meeting, Cold Spring Harbor Laboratory, Cold Spzing Harbor, New York, September 14-1S, 1982. 20. Alitrani-Rosenbaum, S., Maroteaux, L., Mory, Y., Revel, M., and liowley, ?.::.: Inducible expression of the human beta-interferon gene linked to a bovine papillomavirus DNA vector and maintained extrachromosomally in mouse cells. Papilloma Viruses Meeting, Cold Spring Harbor Laboratory, Cold Spring !larbor, New York, September 14-18, 1982. 21. Heilman, C.A., Engel, L.W., and Howley, P.M.: Analysis of the virus-speciiic transcripts in BPV-1 infected bovine fibropapillomas. Papillona Viruses efeetinb, Cold Spring Harbor Laboratory, Cold Spring Harbor, New York, September 14-18, 1982. 22. Turek, L.P., Byrne, ., .. , Low^y, D.R., Dvoretzky, ., FriedWan, and }Iowley, P.M.: in•er feron lnduces rorpholooic reversion witii of extr.+chromosomal viral genomes in buvine papi•1lomavirus-transiormc,I ce:ls. Papillona Viruses Mceting, Cold Spring Harbor Laboratory, Cold Sprin'4 New York, September 14-1b, 1982. 65100 6111 CTR MN 011016

ABSTRACTS A:~D PRESEFTATIONS (Cout'd) 23. Sarver, N., Yang, Y.-C., Byrne. J.C., and Hovley, P.M.: Analysis of genetic elements required for transformation and extrachromosocial nlasmid replication. Papilloma Viruses Neeting, Cold Spring Harbor Laboratory, Cold Spring }larbor, New York, September 14-18, 1982. e5:0G 6112 CTR HN 011019

PUBLICATIONS SUBMITTED OR IN PRESS 1. Howley, P.M.: Papovaviruses - search for evidence of possible association with human cancer. In Phillips, L.A. (Ed.): Viruses Associated with Human Cancer. Marcel Dekker, New York. Submitted: 3-81 In Press: X Journal: See Above 2. Chen, E.Y., Howley, P.M., Levinson, A., and Seeburg, P.H.: The primary structure and• genetic organization of the bovine papillomavirus (BPV) Type I genome. Submitted: 3-82 In Press: X Journal: Nature 3. Howley, P.M., Sarver, N., and Law, M.-F.: Eukaryotic cloning vectors derived from bovine papillomavirus DNA. In Wu, R., Grossman, L., and Moldave, K. (Eds.): Methods in Enzymology. New York, Academic Press, Inc. Submitted: 4-82 In Press: Journal: See Above 4. Turek, L.P., Byrne, J.C., Lowy, D.R., Dvoretzky, I., Friedman, R.M., and Howley, P.M.: Interferon inhibits bovine papillomavirus transformation of mouse cells and induces reversion of established transformants. ,n Merigan, T., Friedman, R.M., and Fox, C.F. (Eds.): UCLA Symposia on Molecular and Cellular Biology.. Vol. XXV, Chemistry and Biology of Interferons: Relationship to Therapeutics. New York, Academic Press. Submitted: 4-82 In Press: X Journal: See Above 5. Banks-Schlegel, S.P., and Howley, P.M.: Differentiation of human epider.mal cells transformed by SV40. Submitted: 5-82 In Press: X Journal: J. Cell Biol. 6. Sarver, N., Mitrani-Rosenbaum; S., Law, M.-F., Byrr.c, J.C., ..nd uo,:lcy, A bovine papillomavirus/p`IL2 hybrid vector -• a dual host replicon. In Sternberg, N.L., and Pearson, M.L. (Eds.): Gene Transfer and Cancer. New York, Raven Press. • Submitted: 6-82 In Press: X Journal: See Above 7. Turek, L.P., Byrne, .;.' , Lowy, D.R., Dvoretzky, I., Friedman, and Howley, P.M.: Interferon induces morphologic reversion with eliminatiorn of extrachromosomal viral genomes in bovine papillomavirus-transformed mouse cells. Sub-mitted: 7/82 In Press: X Journal: Proc. Nat. Acad. Sci. USA 8. Sarver, N., Byrne, J.C., and Howley, P.M.: Transformation and replication in mouse cells of a bovine papillomavirus/p`1L2 plasmid vector that can be rescued in bacteria. Submitted: 7/82 In Press: X Journa): Proc. Nat. Acad. Sci. USA 9. Maroteaux, L. Chen, L., Nitrani-Rosenbauro, S., Howlcy, P.1;., and "ncvel, Cycloheximide induces expression of the huruan interferon I genc in r_ouse cells transformed by bovine papiilomavirus-IFN= 1 recombinants. Submitted: 9-82 In Press: Journal: J. Virol. 651©© 6113 C T R H F"t 0 110~.'! %04