P. Howley

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NCoR1 Mediates Papillomavirus E8^E2C Transcriptional Repression

Jennifer A. Smith, M. Sowa, J. Harper, 2010, Journal of Virology.

The E6 oncoprotein encoded by human papillomavirus types 16 and 18 promotes the degradation of p53

Arnold J. Levine, Martin Scheffner, A. Levine, 1990, Cell.

Human papillomavirus immortalization and transformation functions.

Karl Münger, K. Münger, P. Howley, 2002, Virus research.

Functions of human papillomavirus proteins.

K. Münger, M. Scheffner, P. Howley, 1994, Current topics in microbiology and immunology.

Functional domains of wild-type and mutant p53 proteins involved in transcriptional regulation, transdominant inhibition, and transformation suppression

M. Scheffner, P. Howley, T. Unger, 1993 .

Cloning and expression of the cDNA for E6-AP, a protein that mediates the interaction of the human papillomavirus E6 oncoprotein with p53

M. Scheffner, P. Howley, J. Huibregtse, 1993 .

Localization of the E6-AP regions that direct human papillomavirus E6 binding, association with p53, and ubiquitination of associated proteins

M. Scheffner, P. Howley, J. Huibregtse, 1993 .

p300/MDM2 complexes participate in MDM2-mediated p53 degradation.

D. Livingston, S. Grossman, P. Howley, 1998, Molecular cell.

Live cell, image-based high-throughput screen to quantitate p53 stabilization and viability in human papillomavirus positive cancer cells.

Jennifer A. Smith, V. C. Vieira, P. Howley, 2021, Virology.

The human E6-AP gene (UBE3A) encodes three potential protein isoforms generated by differential splicing.

P. Howley, J. Huibregtse, Y. Yamamoto, 1997, Genomics.

Structure of an E6AP-UbcH7 complex: insights into ubiquitination by the E2-E3 enzyme cascade.

P. Howley, N. Pavletich, J. Huibregtse, 1999, Science.

NOTES TheE6andE7GenesoftheHumanPapillomavirus Type16 Together AreNecessary andSufficient forTransformation ofPrimary HumanKeratinocytes

K. Münger, P. Howley, R. Schlegel, 1989 .

Together Are Necessary and Sufficient for Transformation

P. Howley, K. Munger, R. Schlegel, 2022 .

Proteomic Analysis and Identification of Cellular Interactors of the Giant Ubiquitin Ligase HERC2

J. Harper, P. Howley, T. Chittenden, 2014, Journal of proteome research.

DNA-repair protein hHR23a alters its protein structure upon binding proteasomal subunit S5a

P. Howley, A. Goh, Qinghua Wang, 2003, Proceedings of the National Academy of Sciences of the United States of America.

Virus infection induces the assembly of coordinately activated transcription factors on the IFN-beta enhancer in vivo.

T. Maniatis, C. H. Lin, P. Howley, 1998, Molecular cell.

Human papillomavirus 16 E6 oncoprotein binds to interferon regulatory factor-3 and inhibits its transcriptional activity.

M. Vidal, L. Ronco, A. Karpova, 1998, Genes & development.

A rapid method for detecting and mapping homology between heterologous DNAs. Evaluation of polyomavirus genomes.

M. Israel, P. Howley, M. Martin, 1979, The Journal of biological chemistry.

Structure-function analysis of the human papillomavirus type 16 E7 oncoprotein

K. Münger, P. Howley, C. Yee, 1992, Journal of virology.

Characterization of K virus and its comparison with polyoma virus

P. Howley, S. Bond, K. Takemoto, 1978, Journal of virology.

Transcriptional trans-activation by the human papillomavirus type 16 E2 gene product

P. Howley, W. Phelps, 1987, Journal of virology.

Ubiquitination and degradation of the substrate recognition subunits of SCF ubiquitin-protein ligases.

P. Howley, P. Zhou, Pengbo Zhou, 1998, Molecular cell.

A family of proteins structurally and functionally related to the E6-AP ubiquitin-protein ligase.

M. Scheffner, P. Howley, J. Huibregtse, 1995, Proceedings of the National Academy of Sciences of the United States of America.

Trans-activation of the human immunodeficiency virus long terminal repeat sequence by DNA viruses.

H. Gendelman, P. Howley, M. Martin, 1986, Proceedings of the National Academy of Sciences of the United States of America.

Complex formation of human papillomavirus E7 proteins with the retinoblastoma tumor suppressor gene product.

K. Münger, P. Howley, N. Dyson, 1989, The EMBO journal.

The human papilloma virus-16 E7 oncoprotein is able to bind to the retinoblastoma gene product.

K. Münger, P. Howley, N. Dyson, 1989, Science.

The transcriptional transactivation function of wild‐type p53 is inhibited by SV40 large T‐antigen and by HPV‐16 E6 oncoprotein.

P. Howley, T. Unger, J. Huibregtse, 1992, The EMBO journal.

Differentiation SV 40 of Human Epidermal Cells Transformed by

P. Howley, S. Banks-Schlegel, 2003 .

Differentiation of human epidermal cells transformed by SV40

P. Howley, J. Bonner, S. Banks-Schlegel, 1983, The Journal of cell biology.

Harnessing the ubiquitination machinery to target the degradation of specific cellular proteins.

P. Howley, P. Zhou, L. McReynolds, 2000, Molecular cell.

The Human Papillomavirus E6 Oncoprotein Targets USP15 and TRIM25 To Suppress RIG-I-Mediated Innate Immune Signaling

K. Münger, P. Howley, C. Meyers, 2017, Journal of Virology.

Repression of the Integrated Papillomavirus E6/E7 Promoter Is Required for Growth Suppression of Cervical Cancer Cells

P. Howley, D. A. Francis, S. Schmid, 2000, Journal of Virology.

Inhibition of cyclin D-CDK4/CDK6 activity is associated with an E2F-mediated induction of cyclin kinase inhibitor activity.

J. Nevins, F. Kaye, P. Howley, 1996, Proceedings of the National Academy of Sciences of the United States of America.

Biochemical Analysis of Angelman Syndrome-associated Mutations in the E3 Ubiquitin Ligase E6-associated Protein*

E. M. Cooper, P. Howley, J. Wagstaff, 2004, Journal of Biological Chemistry.

Characterization of papillomavirus E 1 helicase mutants defective for interaction with the SUMO-conjugating enzyme Ubc 9

P. Howley, S. Titolo, J. Archambault, 2009 .

E 2 polypeptides encoded by bovine papillomavirus type 1 form dimers through the common carboxyl-terminal domain : Transactivation is mediated by the conserved amino-terminal domain ( transcriptional regulation / DNA-protein interaction )

P. Howley, J. Byrne, A. McBride, 2022 .

Negative regulation of the bovine papillomavirus E5, E6, and E7 oncogenes by the viral E1 and E2 genes

P. Howley, S. V. Vande Pol, 1995, Journal of virology.

Transient replication of human papillomavirus DNAs

P. Howley, H. Romanczuk, C. Baker, 1992, Journal of virology.

The viral transcriptional regulatory region upstream of the E6 and E7 genes is a major determinant of the differential immortalization activities of human papillomavirus types 16 and 18

R. Schlegel, P. Howley, H. Romanczuk, 1991, Journal of virology.

A bovine papillomavirus constitutive enhancer is negatively regulated by the E2 repressor through competitive binding for a cellular factor

P. Howley, S. V. Vande Pol, Peter M. Howley, 1990, Journal of virology.

Functional and sequence similarities between HPV16 E7 and adenovirus E1A.

K. Münger, P. Howley, C. Yee, 1989, Current topics in microbiology and immunology.

The Biology of the Papillomaviruses

P. Howley, 1987 .

Human papillomaviruses: present knowledge, future approaches.

P. Howley, Peter M. Howley, 1985, Clinics in dermatology.

Biochemical and biological differences between E7 oncoproteins of the high- and low-risk human papillomavirus types are determined by amino-terminal sequences

K. Münger, P. Howley, H. Moses, 1991, Journal of virology.

Interaction of tSNARE Syntaxin 18 with the Papillomavirus Minor Capsid Protein Mediates Infection

I. Bossis, P. Howley, R. Roden, 2005, Journal of Virology.

Transcriptional transActivation by the Human Papillomavirus Type 16 E 2 Gene Product

P. Howley, W. Phelps, 2022 .

Molecular mechanisms of transformation by the human papillomaviruses.

R. Schlegel, K. Münger, P. Howley, 1989, Princess Takamatsu symposia.

The concurrence of saethre‐chotzen syndrome and malignancy in a family with in vitro immune dysfunction

P. Levine, P. Howley, J. Mulvihill, 1984, Cancer.

Factor-binding element in the human c-myc promoter involved in transcriptional regulation by transforming growth factor beta 1 and by the retinoblastoma gene product.

K. Münger, R. Stein, P. Howley, 1991, Proceedings of the National Academy of Sciences of the United States of America.

Ubiquitin recognition by the DNA repair protein hHR23a.

P. Howley, A. Goh, Qinghua Wang, 2003, Biochemistry.

Identification of HHR23A as a Substrate for E6-associated Protein-mediated Ubiquitination*

P. Howley, Sushant Kumar, A. Talis, 1999, The Journal of Biological Chemistry.

Oncoproteins encoded by the cancer-associated human papillomaviruses target the products of the retinoblastoma and p53 tumor suppressor genes.

K. Münger, M. Scheffner, P. Howley, 1991, Cold Spring Harbor symposia on quantitative biology.

Warts, cancer and ubiquitylation: lessons from the papillomaviruses.

P. Howley, 2006, Transactions of the American Clinical and Climatological Association.

Suppression of Cervical Cancer Cells E 6 / E 7 Promoter Is Required for Growth Repression of the Integrated Papillomavirus

P. Howley, D. A. Francis, S. Schmid, 2000 .

Inducible expression of the human interferon beta 1 gene linked to a bovine papilloma virus DNA vector and maintained extrachromosomally in mouse cells

P. Howley, L. Maroteaux, S. Mitrani‐Rosenbaum, 1983, Molecular and cellular biology.

Transcriptional organization of bovine papillomavirus type 1

P. Howley, C. Heilman, L. Engel, 1983, Journal of virology.

Virus-specific transcription in bovine papillomavirus-transformed mouse cells.

D. Lowy, P. Howley, C. Heilman, 1982, Virology.

Identification of a human chromosome 11 gene which is differentially regulated in tumorigenic and nontumorigenic somatic cell hybrids of HeLa cells.

P. Howley, H. Seuánez, W. Modi, 1992, Cell growth & differentiation : the molecular biology journal of the American Association for Cancer Research.

Cycloheximide induces expression of the human interferon beta 1 gene in mouse cells transformed by bovine papillomavirus-interferon beta 1 recombinants

P. Howley, L. Maroteaux, S. Mitrani‐Rosenbaum, 1983, Journal of virology.

Early regions of JC virus and BK virus induce distinct and tissue-specific tumors in transgenic mice.

P. Howley, G. Khoury, G. Jay, 1986, Proceedings of the National Academy of Sciences of the United States of America.

Analysis of trans activation by human papillomavirus type 16 E7 and adenovirus 12S E1A suggests a common mechanism

J. Nevins, K. Münger, P. Howley, 1991, Journal of virology.

Regulation of Aurora B Expression by the Bromodomain Protein Brd4

P. Howley, Matthias Ottinger, Jianxin You, 2009, Molecular and Cellular Biology.

Bovine Papillomavirus DNA Vectors

P. Howley, N. Sarver, 1986 .

Enhancer-dependent expression of the rat preproinsulin gene in bovine papillomavirus type 1 vectors

P. Howley, R. Muschel, G. Khoury, 1985, Molecular and cellular biology.

Transformation and replication in mouse cells of a bovine papillomavirus--pML2 plasmid vector that can be rescued in bacteria.

P. Howley, N. Sarver, J. Byrne, 1982, Proceedings of the National Academy of Sciences of the United States of America.

Bovine papilloma virus deoxyribonucleic acid: a novel eucaryotic cloning vector

P. Gruss, P. Howley, G. Khoury, 1981, Molecular and cellular biology.

In vitro tumorigenic transformation by a defined sub-genomic fragment of bovine papilloma virus DNA

D. Lowy, P. Howley, M. Law, 1980, Nature.

Chapter 6 – Infectious Agents and Cancer

P. Howley, J. DeMasi, 2008 .

Functional analysis of the papilloma virus E2 trans-activator in Saccharomyces cerevisiae.

P. Howley, M. Yaniv, N. Dostatni, 1989, Genes & development.

Kaposi's Sarcoma-Associated Herpesvirus Latency-Associated Nuclear Antigen Interacts with Bromodomain Protein Brd4 on Host Mitotic Chromosomes

P. Howley, M. Ballestas, W. Harrington, 2006, Journal of Virology.

The molecular biology of papillomavirus transformation. Warner-Lambert Parke-Davis Award Lecture.

P. Howley, 1983, The American journal of pathology.

The human papillomaviruses.

P. Howley, 1982, Archives of pathology & laboratory medicine.

Cloning of human papilloma virus genomic DNAs and analysis of homologous polynucleotide sequences

M. Israel, P. Howley, C. Heilman, 1980, Journal of virology.

The human papillomaviruses. An overview.

R. Schlegel, P. Howley, 1988, The American journal of medicine.

c-myc and pRB: Role in TGF-β1 Inhibition of Keratinocyte Proliferation

K. Münger, P. Howley, H. Moses, 1991 .

Differentiation of human epidermal cells transformed by SV40

P. Howley, S. Banks-Schlegel, 1983, The Journal of cell biology.

A stable bovine papillomavirus hybrid plasmid that expresses a dominant selective trait

P. Howley, J. Byrne, M. Law, 1983, Molecular and cellular biology.

Type 1 human T-cell leukemia virus small envelope protein expressed in mouse cells by using a bovine papilloma virus-derived shuttle vector

M. Reitz, P. Howley, A. Fisher, 1985, Molecular and cellular biology.

Human Papillomavirus Type 16 E6 Induces Self-Ubiquitination of the E6AP Ubiquitin-Protein Ligase

P. Howley, J. Huibregtse, S. Beaudenon, 2000, Journal of Virology.

The HPV-16 E6 and E6-AP complex functions as a ubiquitin-protein ligase in the ubiquitination of p53

M. Scheffner, P. Howley, R. Vierstra, 1993, Cell.

The Ubiquitin-Specific Peptidase USP15 Regulates Human Papillomavirus Type 16 E6 Protein Stability

P. Howley, E. White, Jennifer Altreuter, 2009, Journal of Virology.

The bovine papillomavirus E6 oncoprotein interacts with paxillin and disrupts the actin cytoskeleton.

P. Howley, X. Tong, Xiao Tong, 1997, Proceedings of the National Academy of Sciences of the United States of America.

Targeted degradation of the retinoblastoma protein by human papillomavirus E7‐E6 fusion proteins.

K. Münger, M. Scheffner, P. Howley, 1992, The EMBO journal.

NCoR1 Mediates Papillomavirus E8 (cid:1) E2C Transcriptional Repression (cid:1) †

Jennifer A. Smith, M. Sowa, J. Harper, 2010 .

NCoR 1 Mediates Papillomavirus E 8 E 2 C Transcriptional Repression †

Jennifer A. Smith, M. Sowa, J. Harper, 2010 .

Efficiency of binding the retinoblastoma protein correlates with the transforming capacity of the E7 oncoproteins of the human papillomaviruses.

K. Münger, P. Howley, C. Yee, 1992, Proceedings of the National Academy of Sciences of the United States of America.

The bovine papillomavirus constitutive enhancer is essential for viral transformation, DNA replication, and the maintenance of latency

P. Howley, S. V. Vande Pol, 1992, Journal of virology.

Bovine papillomavirus with a mutation in the E2 serine 301 phosphorylation site replicates at a high copy number

P. Howley, A. McBride, 1991, Journal of virology.

Phenotypic analysis of bovine papillomavirus type 1 E2 repressor mutants

P. Howley, P. Lambert, B. Monk, 1990, Journal of virology.

Genetic assignment of multiple E2 gene products in bovine papillomavirus-transformed cells

P. Howley, J. Schiller, P. Lambert, 1989, Journal of virology.

Bovine papillomavirus type 1 E1 replication-defective mutants are altered in their transcriptional regulation

P. Howley, P. Lambert, 1988, Journal of virology.

Mutational analysis of the 3' open reading frames and the splice junction at nucleotide 3225 of bovine papillomavirus type 1

P. Howley, P. Hermonat, 1987, Journal of virology.

Bovine papillomavirus type 1 3' early region transformation and plasmid maintenance functions

P. Howley, C. Yee, Yuh Cheng Yang, 1986, Journal of virology.

The bovine papillomavirus distal "enhancer" is not cis essential for transformation or for plasmid maintenance

P. Howley, E. Schenborn, E. Lund, 1985, Molecular and cellular biology.

Transactivation of a bovine papilloma virus transcriptional regulatory element by the E2 gene product

P. Howley, Yu-Chung Yang, B. Spalholz, 1985, Cell.

Bovine Papillomavirus Shuttle Vectors

P. Howley, W. Mcallister, N. Sarver, 1983 .

Expression oftheRatPreproinsulin Genein BovinePapillomavirus Type1 Vectors

P. Howley, R. Muschel, G. Khoury, 1985 .

Eukaryotic cloning vectors derived from bovine papillomavirus DNA.

P. Howley, N. Sarver, M. Law, 1983, Methods in enzymology.

Analysis of JC virus DNA purified directly from human progressive multifocal leukoencephalopathy brains

P. Howley, A. Lubiniecki, F. Rentier-Delrue, 1981, Journal of virology.

Cloned human polyomavirus JC DNA can transform human amnion cells

M. Israel, K. Chowdhury, P. Howley, 1980, Journal of virology.

RAT INSULIN GENE COVALENTLY LINKED TO BOVINE PAPILLOMAVIRUS DNA IS EXPRESSED IN TRANSFORMED MOUSE CELLS

P. Gruss, P. Howley, G. Khoury, 1981 .

Bovine Papillomavirus E7 Oncoprotein Inhibits Anoikis

P. Howley, M. C. Chao, Michael C. Chao, 2007, Journal of Virology.

Specific chromosomal abnormalities characterize fibrosarcomas of bovine papillomavirus type 1 transgenic mice.

D. Hanahan, P. Howley, J. Skowroński, 1989, Proceedings of the National Academy of Sciences of the United States of America.

Endocrine studies in cystinosis: compensated primary hypothyroidism.

P. Howley, J. Schulman, S. Spielberg, 1977, The Journal of pediatrics.

PURIFICATION OF INTACT MICROTUBULES FROM BRAIN

P. Howley, J. Kirkpatrick, V. Thomas, 1970, The Journal of cell biology.

The role of papillomaviruses in human cancer.

P. Howley, 1987, Important advances in oncology.

Evidence for cooperativity between E2 binding sites in E2 trans-regulation of bovine papillomavirus type 1

P. Howley, J. Byrne, B. Spalholz, 1988, Journal of virology.

Bovine papillomavirus transcriptional regulation: localization of the E2-responsive elements of the long control region

P. Howley, C. Yee, P. Lambert, 1987, Journal of Virology.

Dissociation of transforming and trans-activation functions for bovine papillomavirus type 1

P. Howley, Yu-Chung Yang, B. Spalholz, 1985, Nature.

The genetics of bovine papillomavirus type 1.

P. Howley, P. Lambert, C. Baker, 1988, Annual review of genetics.

The ER‐Lumenal Domain of the HHV‐7 Immunoevasin U21 Directs Class I MHC Molecules to Lysosomes

H. Ploegh, P. Howley, D. Blom, 2003, Traffic.

Genome localization of simian virus 40 RNA species

P. Howley, M. Martin, G. Khoury, 1976, Journal of virology.

Papillomavirus E2 induces senescence in HPV‐positive cells via pRB‐ and p21CIP‐dependent pathways

J. D. Benson, A. Karpova, P. Howley, 2000, The EMBO journal.

Transcriptional trans-Activation bytheHumanPapillomavirus Type16E2GeneProduct

P. Howley, 1987 .

Identification of a human ubiquitin-conjugating enzyme that mediates the E6-AP-dependent ubiquitination of p53.

M. Scheffner, P. Howley, J. Huibregtse, 1994, Proceedings of the National Academy of Sciences of the United States of America.

Brd 4-Independent Transcriptional Repression Function of the Papillomavirus E 2 Proteins

P. Howley, Matthias Ottinger, M. Schweiger, 2007 .

Erratum: A family of proteins structurally and functionally related to the E6-AP ubiquitin-protein ligase (Proceedings of the National Academy of Sciences of the United States of America (March 28, 1996) 92:7 (2563-2564))

M. Scheffner, P. Howley, J. Huibregtse, 1995 .

Phosphorylation sites of the E2 transcriptional regulatory proteins of bovine papillomavirus type 1

P. Howley, J. Bolen, A. McBride, 1989, Journal of virology.

The Bovine Papillomavirus E6 Protein Binds to the LD Motif Repeats of Paxillin and Blocks Its Interaction with Vinculin and the Focal Adhesion Kinase*

R. Salgia, P. Howley, Jian-Liang Li, 1997, The Journal of Biological Chemistry.

Posttranscriptional Selection of Simian Virus 40-Specific RNA

P. Howley, G. Khoury, D. Nathans, 1975, Journal of virology.

Identification of TRAIL as an Interferon Regulatory Factor 3 Transcriptional Target

A. Karpova, P. Howley, J. Kirshner, 2005, Journal of Virology.

Maren Kops and Peter Regulatory Factor 3 Transcriptional Target Identification of TRAIL as an Interferon

A. Karpova, P. Howley, J. Kirshner, 2005 .

Mouse cells transformed by bovine papillomavirus contain only extrachromosomal viral DNA sequences.

D. Lowy, P. Howley, M. Law, 1981, Proceedings of the National Academy of Sciences of the United States of America.

INTERFERON INHIBITS BOVINE PAPILLOMAVIRUS TRANSFORMATION OF MOUSE CELLS AND INDUCES REVERSION OF ESTABLISHED TRANSFORMANTS

D. Lowy, R. Friedman, P. Howley, 1982 .

Tumor progression in transgenic mice containing the bovine papillomavirus genome.

P. Howley, 1991, Basic life sciences.

The functional BPV-1 E2 trans-activating protein can act as a repressor by preventing formation of the initiation complex.

P. Howley, M. Yaniv, N. Dostatni, 1991, Genes & development.

Oncogenicity of a nude mouse cell line transformed by a human papovavirus.

P. Howley, C. Yee, N. Young, 1977, Journal of the National Cancer Institute.

NOTES Genetic Assignment ofMultiple E2GeneProducts inBovine Papillomavirus-Transformed Cells

P. Howley, J. Schiller, P. Lambert, 1989 .

Kinetics of inhibition of papovavirus DNA synthesis by superinfection with adenovirus 2 and non-defective adenovirus 2-simian virus 40 hybrid viruses.

P. Howley, A. Levine, J. Hauser, 1981, The Journal of general virology.

Characterization of the genome of the murine papovavirus K

P. Howley, M. Law, K. Takemoto, 1979, Journal of virology.

JC human papovavirus replication in human amnion cells

P. Howley, T. Miyamura, K. Takemoto, 1979, Journal of virology.

JCHumanPapovavirus Replication inHumanAmnionCells

P. Howley, T. Miyamura, 1979 .

Isolation of simian virus 40 from rhesus monkeys (Macaca mulatta) with spontaneous progressive multifocal leukoencephalopathy.

P. Howley, K. Takemoto, C. Holmberg, 1977, The Journal of infectious diseases.

Uniform representation of the human papovavirus BK genome in transformed hamster cells

P. Howley, M. Martin, M. Martin, 1977, Journal of virology.

Polynucleotide sequences common to the genomes of simian virus 40 and the human papovaviruses JC and BK.

P. Howley, M. Martin, G. Khoury, 1976, Virology.

Characterization of supercoiled oligomeric SV40 DNA molecules in productively infected cells.

P. Howley, M. Martin, J. Byrne, 1976, Virology.

Physical map of the BK virus genome

P. Howley, M. Martin, G. Khoury, 1975, Journal of virology.

Characterization of human papovavirus BK DNA

P. Howley, M. Martin, K. Takemoto, 1975, Journal of virology.

Mapping and characterization of the interaction domains of human papillomavirus type 16 E1 and E2 proteins

J. D. Benson, M. Vidal, P. Howley, 1997, Journal of virology.

A Fifteen-Amino-Acid Peptide Inhibits Human Papillomavirus E1-E2 Interaction and Human Papillomavirus DNA Replication In Vitro

J. D. Benson, P. Howley, H. Kasukawa, 1998, Journal of Virology.

The carboxy‐terminal domain shared by the bovine papillomavirus E2 transactivator and repressor proteins contains a specific DNA binding activity.

R. Schlegel, P. Howley, A. McBride, 1988, The EMBO journal.

Two classes of human papillomavirus type 16 E1 mutants suggest pleiotropic conformational constraints affecting E1 multimerization, E2 interaction, and interaction with cellular proteins

J. D. Benson, M. Vidal, P. Howley, 1997, Journal of virology.

p35, the Neuronal-specific Activator of Cyclin-dependent Kinase 5 (Cdk5) Is Degraded by the Ubiquitin-Proteasome Pathway*

L. Tsai, Young T. Kwon, P. Howley, 1998, The Journal of Biological Chemistry.

Irradiation of methyl diazomalonate in solution. Reactions of singlet and triplet carbenes with carbon-carbon double bonds

P. Howley, Maitland Jones, W. Andō, 1972 .

Structural and transcriptional analysis of human papillomavirus type 16 sequences in cervical carcinoma cell lines

M. Braun, M. Gonda, P. Howley, 1987, Journal of virology.

Proteomic approaches to the study of papillomavirus-host interactions.

P. Howley, E. White, 2013, Virology.

Interaction of the Bovine Papillomavirus E2 Protein with Brd4 Tethers the Viral DNA to Host Mitotic Chromosomes

P. Howley, K. Ozato, A. Nishimura, 2004, Cell.

The primary structure and genetic organization of the bovine papillomavirus type 1 genome

Peter H. Seeburg, Ellson Y. Chen, E. Chen, 1982, Nature.

Bovine papillomavirus E7 transformation function correlates with cellular p600 protein binding.

K. Münger, P. Howley, Y. Nakatani, 2005, Proceedings of the National Academy of Sciences of the United States of America.

Differential promoter utilization by the bovine papillomavirus in transformed cells and productively infected wart tissues.

P. Howley, C. Baker, 1987, The EMBO journal.

Progressive squamous epithelial neoplasia in K14-human papillomavirus type 16 transgenic mice

D. Hanahan, K. Münger, J. Arbeit, 1994, Journal of virology.

Brd4 Regulation of Papillomavirus Protein E2 Stability

Jennifer A. Smith, J. Harper, P. Howley, 2009, Journal of Virology.

Inhibition of E2 Binding to Brd4 Enhances Viral Genome Loss and Phenotypic Reversion of Bovine Papillomavirus-Transformed Cells

P. Howley, M. Schweiger, Jianxin You, 2005, Journal of Virology.

Functional analysis of E2-mediated repression of the HPV18 P105 promoter.

P. Howley, F. Thierry, 1991, The New biologist.

E2 polypeptides encoded by bovine papillomavirus type 1 form dimers through the common carboxyl-terminal domain: transactivation is mediated by the conserved amino-terminal domain.

P. Howley, J. Byrne, A. McBride, 1989, Proceedings of the National Academy of Sciences of the United States of America.

A transcriptional repressor encoded by BPV-1 shares a common carboxy-terminal domain with the E2 transactivator

P. Howley, P. Lambert, B. Spalholz, 1987, Cell.

Localization and analysis of bovine papillomavirus type 1 transforming functions

P. Howley, Y. Yang, N. Sarver, 1984, Journal of virology.

THE BIOLOGY OF THE PAPILLOMA VIRUSES

P. Howley, 1987 .

Human papillomavirus-11 DNA in a patient with chronic laryngotracheobronchial papillomatosis and metastatic squamous-cell carcinoma of the lung.

M. Tsao, P. Howley, J. Byrne, 1987, The New England journal of medicine.

The Papovaviridae

P. Howley, N. Salzman, H. Ahola, 1987, The Viruses.

Interferon induces morphologic reversion with elimination of extrachromosomal viral genomes in bovine papillomavirus-transformed mouse cells.

D. Lowy, R. Friedman, P. Howley, 1982, Proceedings of the National Academy of Sciences of the United States of America.

Presence of human papilloma viral antigens in juvenile multiple laryngeal papilloma.

P. Howley, J. Costa, W. Bauer, 1981, American journal of clinical pathology.

Conserved polynucleotide sequences among the genomes of papillomaviruses

P. Howley, W. Lancaster, M. Law, 1979, Journal of virology.

The state of the p53 and retinoblastoma genes in human cervical carcinoma cell lines.

K. Münger, M. Scheffner, P. Howley, 1991, Proceedings of the National Academy of Sciences of the United States of America.

Adenovirus E1A, simian virus 40 tumor antigen, and human papillomavirus E7 protein share the capacity to disrupt the interaction between transcription factor E2F and the retinoblastoma gene product.

J. Nevins, P. Howley, K. Munger, 1992, Proceedings of the National Academy of Sciences of the United States of America.

The SMC5/6 Complex Interacts with the Papillomavirus E2 Protein and Influences Maintenance of Viral Episomal DNA

P. Howley, A. McBride, E. White, 2018, Journal of Virology.

Two AP1 sites binding JunB are essential for human papillomavirus type 18 transcription in keratinocytes

P. Howley, M. Yaniv, G. Spyrou, 1992, Journal of virology.

Interaction of the Bovine Papillomavirus E6 Protein with the Clathrin Adaptor Complex AP-1

T. Kirchhausen, P. Howley, W. Boll, 1998, Journal of Virology.

The papillomavirus E2 regulatory proteins.

P. Howley, H. Romanczuk, A. McBride, 1991, The Journal of biological chemistry.

The E6 and E7 genes of the human papillomavirus type 16 together are necessary and sufficient for transformation of primary human keratinocytes

R. Schlegel, K. Münger, P. Howley, 1989, Journal of virology.

Bovine papillomavirus contains multiple transforming genes.

P. Howley, H. Okayama, Yuh Cheng Yang, 1985, Proceedings of the National Academy of Sciences of the United States of America.

Association of the human papillomavirus type 16 E7 oncoprotein with the 600-kDa retinoblastoma protein-associated factor, p600.

K. Münger, P. Howley, Y. Nakatani, 2005, Proceedings of the National Academy of Sciences of the United States of America.

Human Papillomavirus E6 and E7 Oncogenes

K. Münger, P. Howley, D. DiMaio, 2007 .

Role of the human papillomaviruses in human cancer.

P. Howley, Peter M. Howley, 1991, Cancer research.

Presence and expression of human papillomavirus sequences in human cervical carcinoma cell lines.

R. Schlegel, P. Howley, C. Yee, 1985, The American journal of pathology.

A cellular protein mediates association of p53 with the E6 oncoprotein of human papillomavirus types 16 or 18.

M. Scheffner, P. Howley, J. Huibregtse, 1991, The EMBO journal.

Association of human papillomavirus types 16 and 18 E6 proteins with p53.

A. Levine, P. Howley, B. Werness, 1990, Science.

Targeted mutagenesis of the human papillomavirus type 16 E2 transactivation domain reveals separable transcriptional activation and DNA replication functions

J. D. Benson, P. Howley, T. Yasugi, 1996, Journal of virology.

Characterization of the cis elements involved in basal and E2-transactivated expression of the bovine papillomavirus P2443 promoter

P. Howley, S. V. Vande Pol, B. Spalholz, 1991, Journal of virology.

Mutational analysis of cis elements involved in E2 modulation of human papillomavirus type 16 P97 and type 18 P105 promoters

P. Howley, H. Romanczuk, F. Thierry, 1990, Journal of virology.

Cell-heritable stages of tumor progression in transgenic mice harboring the bovine papillomavirus type 1 genome

D. Hanahan, P. Howley, M. Sippola-Thiele, 1989, Molecular and cellular biology.

Transcriptional Regulation by Papillomavirus E2 Gene Products

P. Howley, P. Lambert, A. McBride, 1989 .

Chronic estrogen-induced cervical and vaginal squamous carcinogenesis in human papillomavirus type 16 transgenic mice.

D. Hanahan, J. Arbeit, P. Howley, 1996, Proceedings of the National Academy of Sciences of the United States of America.

SMCX and components of the TIP60 complex contribute to E2 regulation of the HPV E6/E7 promoter.

Jennifer A. Smith, D. Livingston, J. Decaprio, 2014, Virology.

Genome-wide siRNA screen identifies SMCX, EP400, and Brd4 as E2-dependent regulators of human papillomavirus oncogene expression

Jennifer A. Smith, M. Sowa, J. Harper, 2010, Proceedings of the National Academy of Sciences.

Cell-type specific transcriptional activities among different papillomavirus long control regions and their regulation by E2.

Jennifer A. Smith, M. L. Powell, P. Howley, 2009, Virology.

Brd4-Independent Transcriptional Repression Function of the Papillomavirus E2 Proteins

P. Howley, Matthias Ottinger, M. Schweiger, 2007, Journal of Virology.

Transcriptome signature of irreversible senescence in human papillomavirus-positive cervical cancer cells

B. Aronow, P. Howley, S. Wells, 2003, Proceedings of the National Academy of Sciences of the United States of America.

Mechanisms of Human Papillomavirus E2-Mediated Repression of Viral Oncogene Expression and Cervical Cancer Cell Growth Inhibition

P. Howley, T. Ono, A. Ishimoto, 2000, Journal of Virology.

Suppression of cellular proliferation by the papillomavirus E2 protein

P. Howley, A. McBride, J. Dowhanick, 1995, Journal of virology.

Conserved interaction of the papillomavirus E2 transcriptional activator proteins with human and yeast TFIIB proteins

J. D. Benson, P. Howley, R. Lawande, 1997, Journal of virology.

Phosphorylation Sites oftheE2Transcriptional Regulatory Proteins ofBovinePapillomavirus Type1

P. Howley, 1989 .

Disruption of either the E1 or the E2 regulatory gene of human papillomavirus type 16 increases viral immortalization capacity.

P. Howley, H. Romanczuk, 1992, Proceedings of the National Academy of Sciences of the United States of America.

Amino-terminal domains of the bovine papillomavirus type 1 E1 and E2 proteins participate in complex formation

J. D. Benson, P. Howley, John D. Benson, 1995, Journal of virology.

Gordon Wilson Lecture: Infectious Disease Causes of Cancer: Opportunities for Prevention and Treatment.

P. Howley, 2015, Transactions of the American Clinical and Climatological Association.

The human papillomavirus type 16 E7 gene encodes transactivation and transformation functions similar to those of adenovirus E1A

K. Münger, P. Howley, C. Yee, 1988, Cell.

Localization of the E6-AP regions that direct human papillomavirus E6 binding, association with p53, and ubiquitination of associated proteins

M. Scheffner, P. Howley, J. Huibregtse, 1993, Molecular and cellular biology.

Cloning and expression of the cDNA for E6-AP, a protein that mediates the interaction of the human papillomavirus E6 oncoprotein with p53

M. Scheffner, P. Howley, J. Huibregtse, 1993, Molecular and cellular biology.

Beta genus papillomaviruses and skin cancer.

P. Howley, H. Pfister, 2015, Virology.

Systematic identification of interactions between host cell proteins and E7 oncoproteins from diverse human papillomaviruses

M. Sowa, K. Münger, J. Harper, 2012, Proceedings of the National Academy of Sciences.

Papillomaviruses: Molecular and clinical aspects

P. Howley, T. Broker, 1985 .

Interactions of HPV E6 and E7 oncoproteins with tumour suppressor gene products.

K. Münger, M. Scheffner, P. Howley, 1992, Cancer surveys.

Cutaneous β-human papillomavirus E6 proteins bind Mastermind-like coactivators and repress Notch signaling

J. Aster, M. Sowa, J. Harper, 2012, Proceedings of the National Academy of Sciences.

Comprehensive Analysis of Host Cellular Interactions with Human Papillomavirus E6 Proteins Identifies New E6 Binding Partners and Reflects Viral Diversity

J. Harper, P. Howley, Rebecca E. Kramer, 2012, Journal of Virology.

Ubiquitination of p53 and p21 is differentially affected by ionizing and UV radiation

P. Howley, C. Maki, Peter M. Howley, 1997, Molecular and cellular biology.

In vivo ubiquitination and proteasome-mediated degradation of p53(1).

P. Howley, J. Huibregtse, C. Maki, 1996, Cancer research.

Ubiquitin-mediated degradation of active Src tyrosine kinase.

Santhosh K. P. Kumar, K. F. Harris, E. M. Cooper, 1999, Proceedings of the National Academy of Sciences of the United States of America.

Mutations in serines 15 and 20 of human p53 impair its apoptotic activity

P. Howley, T. Unger, M. Oren, 1999, Oncogene.

Small DNA tumor viruses: large contributors to biomedical sciences.

D. Livingston, P. Howley, 2009, Virology.

Human Papillomavirus E6 Triggers Upregulation of the Antiviral and Cancer Genomic DNA Deaminase APOBEC3B

V. C. Vieira, M. Soares, P. Howley, 2014, mBio.

Merkel cell polyomavirus recruits MYCL to the EP400 complex to promote oncogenesis

Benjamin J. Strober, Alexander J. Federation, J. Bradner, 2017, PLoS pathogens.

Comparative analysis of the human type 1a and bovine type 1 papillomavirus genomes

E. Chen, P. Howley, M. Yaniv, 1983, Journal of virology.

The colinear alignment of the genomes of papovaviruses JC, BK, and SV40.

P. Howley, M. Law, K. Takemoto, 1979, Virology.

Ubiquitin family proteins and their relationship to the proteasome: a structural perspective.

Gerhard Wagner, G. Wagner, P. Howley, 2004, Biochimica et biophysica acta.

Cellular targets of the oncoproteins encoded by the cancer associated human papillomaviruses.

K. Münger, M. Scheffner, P. Howley, 1991, Princess Takamatsu symposia.

Comparison of Southern blot hybridization and polymerase chain reaction methods for the detection of human papillomavirus DNA

M. Schiffman, A. Glass, P. Howley, 1991, Journal of clinical microbiology.

Evaluation of normal and neoplastic human tissue for BK virus.

M. Israel, P. Howley, M. Martin, 1978, Virology.

NSD3-NUT fusion oncoprotein in NUT midline carcinoma: implications for a novel oncogenic mechanism.

C. Antonescu, P. D. Dal Cin, P. Howley, 2014, Cancer discovery.

TGF-β1 inhibition of c-myc transcription and growth in keratinocytes is abrogated by viral transforming proteins with pRB binding domains

P. Yaciuk, E. Moran, K. Münger, 1990, Cell.

The hPLIC proteins may provide a link between the ubiquitination machinery and the proteasome.

P. Howley, P. Zhou, S. Kumar, 2000, Molecular cell.

Functional domains of wild-type and mutant p53 proteins involved in transcriptional regulation, transdominant inhibition, and transformation suppression

M. Scheffner, P. Howley, T. Unger, 1993, Molecular and cellular biology.

Homology and relationship between the genomes of papovaviruses, BK virus and simian virus 40.

P. Howley, M. Martin, G. Khoury, 1975, Proceedings of the National Academy of Sciences of the United States of America.

Regulation of the Src family tyrosine kinase Blk through E6AP-mediated ubiquitination.

Santhosh K. P. Kumar, P. Howley, H. Oda, 1999, Proceedings of the National Academy of Sciences of the United States of America.

The Role of E6AP in the Regulation of p53 Protein Levels in Human Papillomavirus (HPV)-positive and HPV-negative Cells*

P. Howley, J. Huibregtse, A. Talis, 1998, The Journal of Biological Chemistry.

Identification of MicroRNAs That Stabilize p53 in Human Papillomavirus-Positive Cancer Cells

Jennifer A. Smith, P. Howley, Gustavo Martínez-Noël, 2020, bioRxiv.

1 1 Stabilization of p 53 by microRNAs in HPV-positive 2 cervical cancer cells 3 4

Jennifer A. Smith, P. Howley, Gustavo Martínez-Noël, 2022 .

Functional interaction between the early viral proteins of simian virus 40 and adenovirus.

P. Howley, G. Khoury, N. Goldman, 1981, Virology.

Requirement of E7 oncoprotein for viability of HeLa cells.

P. Howley, S. Kyo, T. Ueno, 2006, Microbes and infection.

The ubiquitin-associated domain of hPLIC-2 interacts with the proteasome.

P. Howley, M. Kleijnen, R. Alarcon, 2003, Molecular biology of the cell.

The Brd4 Extraterminal Domain Confers Transcription Activation Independent of pTEFb by Recruiting Multiple Proteins, Including NSD3

Jennifer A. Smith, M. Sowa, J. Harper, 2011, Molecular and Cellular Biology.

Dual utilization of an acceptor/donor splice site governs the alternative splicing of the IRF-3 gene.

A. Karpova, P. Howley, L. Ronco, 2000, Genes & development.

The Molecular Basis of Cancer

Joe W. Gray, Mark A. Israel, John Mendelsohn, 1985 .

High-Risk Human Papillomavirus E7 Proteins Target PTPN14 for Degradation

K. Münger, P. Howley, E. White, 2016, mBio.

Inhibition of cyclin D-CDK 4 / CDK 6 activity is associated with an E 2 F-mediated induction of cyclin kinase inhibitor activity ( human papillomaviruses / cervical cancer / E 7 oncoprotein / retinoblastoma protein )

F. Kaye, P. Howley, G. Otterson, 2022 .

The assembly of the measles virus nucleoprotein into nucleocapsid-like particles is modulated by the phosphoprotein.

P. Howley, D. Spehner, R. Drillien, 1997, Virology.

Physical Interaction between Specific E2 and Hect E3 Enzymes Determines Functional Cooperativity*

P. Howley, Sushant Kumar, Wynn H. Kao, 1997, The Journal of Biological Chemistry.

NSD3-NUT Fusion Oncoprotein in NUT Midline Carcinoma: Implications for a Novel 1

C. Antonescu, P. Cin, P. Howley, 2014 .

Functional Characterization of Interferon Regulatory Factor 3a (IRF-3a), an Alternative Splice Isoform of IRF-3

L. Ronco, A. Karpova, P. Howley, 2001, Molecular and Cellular Biology.

Interferon regulatory factor-3 is an in vivo target of DNA-PK

L. Cantley, A. Karpova, P. Howley, 2002, Proceedings of the National Academy of Sciences of the United States of America.

Genome Structures of reiteration mutants of simian virus 40.

P. Howley, M. Martin, G. Khoury, 1977, Virology.

Ubiquitination of the p53 Tumor Suppressor

P. Howley, J. Huibregtse, C. Maki, 1998 .

CDYL bridges REST and histone methyltransferases for gene repression and suppression of cellular transformation.

S. Elledge, J. Barretina, T. Westbrook, 2008, Molecular cell.

Comprehensive Genomic Profiling of Advanced Penile Carcinoma Suggests a High Frequency of Clinically Relevant Genomic Alterations.

P. Stephens, R. Yelensky, D. Lipson, 2016, The oncologist.

The Molecular Basis of Cancer: Fourth Edition

J. Gray, M. Israel, P. Howley, 2014 .

Interaction of the human papillomavirus type 16 E6 oncoprotein with wild-type and mutant human p53 proteins

J. Minna, M. Scheffner, P. Howley, 1992, Journal of virology.

Components of the ubiquitin-proteasome pathway compete for surfaces on Rad23 family proteins

R. Deshaies, D. Finley, P. Howley, 2008, BMC Biochemistry.

Bromodomain Protein 4 Mediates the Papillomavirus E2 Transcriptional Activation Function

P. Howley, M. Schweiger, Jianxin You, 2006, Journal of Virology.

The bovine papillomavirus P2443 promoter is E2 trans‐responsive: evidence for E2 autoregulation.

P. Howley, P. Hermonat, B. Spalholz, 1988, The EMBO journal.

Acquired immunodeficiency syndrome with progressive multifocal leukoencephalopathy and monoclonal B-cell proliferation.

D. Snydman, P. Howley, D. McEniry, 1984, Annals of internal medicine.

A Human Herpesvirus 7 Glycoprotein, U21, Diverts Major Histocompatibility Complex Class I Molecules to Lysosomes

H. Ploegh, P. Howley, A. Hudson, 2001, Journal of Virology.

Papillomavirus E7 Oncoproteins Share Functions with Polyomavirus Small T Antigens

W. Hahn, D. Livingston, T. Roberts, 2014, Journal of Virology.

Neuroepithelial carcinomas in mice transgenic with human papillomavirus type 16 E6/E7 ORFs.

D. Hanahan, K. Münger, J. Arbeit, 1993, The American journal of pathology.

A vaccinia virus transfer vector using a GUS reporter gene inserted into the I4L locus.

P. Howley, D. Spehner, R. Drillien, 1996, Gene.

A functional measles virus replication and transcription machinery encoded by the vaccinia virus genome.

M. Billeter, P. Howley, D. Spehner, 1999, Journal of virological methods.

Network Analysis of UBE3A/E6AP-Associated Proteins Provides Connections to Several Distinct Cellular Processes.

M. Vidal, Q. Zhong, Katja Luck, 2018, Journal of molecular biology.

Identification and Proteomic Analysis of Distinct UBE3A/E6AP Protein Complexes

M. Sowa, J. Harper, P. Howley, 2012, Molecular and Cellular Biology.

E6AP Ubiquitin Ligase Mediates Ubiquitylation and Degradation of Hepatitis C Virus Core Protein

P. Howley, Y. Matsuura, T. Wakita, 2006, Journal of Virology.

Conserved cysteine residue in the DNA-binding domain of the bovine papillomavirus type 1 E2 protein confers redox regulation of the DNA-binding activity in vitro.

R D Klausner, R. Klausner, P. Howley, 1992, Proceedings of the National Academy of Sciences of the United States of America.

Association of distinct yeast Not2 functional domains with components of Gcn5 histone acetylase and Ccr4 transcriptional regulatory complexes

J. D. Benson, K. Struhl, P. Howley, 1998, The EMBO journal.

BovinePapillomavirus witha Mutation intheE2Serine 301 Phosphorylation SiteReplicates ata HighCopyNumber

P. Howley, 1991 .

Characterization of papillomavirus E1 helicase mutants defective for interaction with the SUMO-conjugating enzyme Ubc9.

P. Howley, S. Titolo, J. Archambault, 2009, Virology.

E6AP and Calmodulin Reciprocally Regulate Estrogen Receptor Stability*

P. Howley, D. Sacks, Lu Li, 2006, Journal of Biological Chemistry.

E6-AP directs the HPV E6-dependent inactivation of p53 and is representative of a family of structurally and functionally related proteins.

M. Scheffner, P. Howley, J. Huibregtse, 1994, Cold Spring Harbor symposia on quantitative biology.

Angelman syndrome–associated point mutations in the Zn2+-binding N-terminal (AZUL) domain of UBE3A ubiquitin ligase inhibit binding to the proteasome

J. Harper, P. Howley, Gustavo Martínez-Noël, 2018, The Journal of Biological Chemistry.

Genus Beta Human Papillomavirus E6 Proteins Vary in Their Effects on the Transactivation of p53 Target Genes

P. Howley, H. Javanbakht, E. White, 2014, Journal of Virology.

Identification of the structural and functional human homolog of the yeast ubiquitin conjugating enzyme UBC9.

P. Howley, T. Yasugi, 1996, Nucleic acids research.

A family of proteins structurally and functionally related to the E6-AP ubiquitin-protein ligase.

M. Scheffner, P. Howley, J. Huibregtse, 1995, Proceedings of the National Academy of Sciences of the United States of America.

Structural studies of the interaction between ubiquitin family proteins and proteasome subunit S5a.

Gerhard Wagner, G. Wagner, P. Howley, 2002, Biochemistry.

Identification of the primate papovavirus HD as the stump-tailed macaque virus

K. Shah, P. Gruss, P. Howley, 1979, Journal of virology.

Characterization ofHumanPapovavirus BK DNA

P. Howley, 1975 .

Role of the human papillomavirus oncoproteins in transformation and carcinogenic progression.

K. Münger, P. Howley, B. Werness, 1991, Important advances in oncology.

The detection and quantitation of SV40 nucleic acid sequences using single-stranded SV40 DNA probes.

P. Howley, G. Khoury, M. Brown, 1975, Cold Spring Harbor symposia on quantitative biology.

Identification of MicroRNAs That Stabilize p53 in Human Papillomavirus-Positive Cancer Cells

Jennifer A. Smith, P. Howley, Gustavo Martínez-Noël, 2021, Journal of virology.

A family of proteins structurally and functionally related to the E 6-AP ubiquitin-protein ligase ( papillomavirus / thioester )

P. Howley, J. Huibregtse, S. Beaudenon, 2022 .

Bovine papillomavirus E 7 transformation function correlates with cellular p 600 protein binding

K. Münger, P. Howley, Y. Nakatani, 2006 .

CDYL bridges REST and histone methyltransferases for gene repression and suppression of cellular transformation.

S. Elledge, J. Barretina, T. Westbrook, 2008, Molecular cell.

Structure-function analysis of the human papillomavirus type 16 E7 oncoprotein

K. Münger, P. Howley, W. Phelps, 1992, Journal of virology.

Beta genus papillomaviruses and skin cancer.

P. Howley, H. Pfister, 2015, Virology.

Evidence forCooperativity between E2Binding Sites inE2 trans-Regulation ofBovine Papillomavirus Type1

P. Howley, 1988 .