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Alan N. Engelman, PhD


  • Ballandras-Colas A, Naraharisetty H, Li X, Serrao E, Engelman A.Biochemical characterization of novel retroviral integrase proteins.PLoS ONE. 2013;8(10):e76638.
  • Ambrose Z, Lee K, Ndjomou J, Xu H, Oztop I, Matous J, Takemura T, Unutmaz D, Engelman A, Hughes SH, Kewalramani VN.Human Immunodeficiency Virus Type 1 Capsid Mutation N74D Alters Cyclophilin A Dependence and Impairs Macrophage Infection.J Virol. 2012 Apr;86(8):4708-14.
  • Kessl JJ, Jena N, Koh Y, Taskent-Sezgin H, Slaughter A, Feng L, de Silva S, Wu L, Le Grice SF, Engelman A, Fuchs JR, Kvaratskhelia M.A multimode, cooperative mechanism of action of allosteric HIV-1 integrase inhibitors.J Biol Chem. 2012 Mar 21.
  • Engelman A, Cherepanov P.The structural biology of HIV-1: mechanistic and therapeutic insights.Nat Rev Microbiol. 2012;10(4):279-90.
  • Li X, Koh Y, Engelman A.Correlation of Recombinant Integrase Activity and Functional Preintegration Complex Formation during Acute Infection by Replication-Defective Integrase Mutant Human Immunodeficiency Virus.J Virol. 2012 Apr;86(7):3861-79.
  • Matreyek KA, Engelman A.The requirement for nucleoporin NUP153 during human immunodeficiency virus type 1 infection is determined by the viral capsid.J Virol. 2011 Aug;85(15):7818-27.
  • Yan N, O'Day E, Wheeler LA, Engelman A, Lieberman J.HIV DNA is heavily uracilated, which protects it from autointegration.Proc Natl Acad Sci U S A. 2011 May 31;108(22):9244-9.
  • Zamborlini A, Coiffic A, Beauclair G, Delelis O, Paris J, Koh Y, Magne F, Giron ML, Tobaly-Tapiero J, Deprez E, Emiliani S, Engelman A, de The H, Saib A.Impairment of human immunodeficiency virus type-1 integrase SUMOylation correlates with an early replication defect.J Biol Chem. 2011 Apr 1.
  • Koh Y, Matreyek KA, Engelman A.Differential sensitivities of retroviruses to integrase strand transfer inhibitors.J Virol. 2011 Apr;85(7):3677-82.
  • Li X, Krishnan L, Cherepanov P, Engelman A.Structural biology of retroviral DNA integration.Virology. 2011 Mar 15;411(2):194-205.
  • Koh Y, Haim H, Engelman A.Identification and characterization of persistent intracellular human immunodeficiency virus type 1 integrase strand transfer inhibitor activity.Antimicrob Agents Chemother. 2011 Jan;55(1):42-9.
  • Krishnan L, Li X, Naraharisetty HL, Hare S, Cherepanov P, Engelman A.Structure-based modeling of the functional HIV-1 intasome and its inhibition.Proc Natl Acad Sci U S A. 2010 Sep 7;107(36):15910-5.
  • Lee K, Ambrose Z, Martin TD, Oztop I, Mulky A, Julias JG, Vandegraaff N, Baumann JG, Wang R, Yuen W, Takemura T, Shelton K, Taniuchi I, Li Y, Sodroski J, Littman DR, Coffin JM, Hughes SH, Unutmaz D, Engelman A, Kewalramani VN.Flexible Use of Nuclear Import Pathways by HIV-1.Cell Host Microbe. 2010 Mar 18;7(3):221-233.
  • Hare S, Gupta SS, Valkov E, Engelman A, Cherepanov P.Retroviral intasome assembly and inhibition of DNA strand transfer.Nature. 2010 Mar 11;464(7286):232-6.
  • Ferris AL, Wu X, Hughes CM, Stewart C, Smith SJ, Milne TA, Wang GG, Shun MC, Allis CD, Engelman A, Hughes SH.Lens epithelium-derived growth factor fusion proteins redirect HIV-1 DNA integration.Proc Natl Acad Sci U S A. 2010 Feb 16;107(7):3135-40.
  • Hughes S, Jenkins V, Dar MJ, Engelman A, Cherepanov P.Transcriptional co-activator LEDGF interacts with Cdc7-activator of S-phase kinase (ASK) and stimulates its enzymatic activity.J Biol Chem. 2010 Jan 1;285(1):541-54.
  • Krishnan L, Matreyek KA, Oztop I, Lee K, Tipper CH, Li X, Dar MJ, Kewalramani VN, Engelman A.The requirement for cellular transportin 3 (TNPO3 or TRN-SR2) during infection maps to human immunodeficiency virus type 1 capsid and not integrase.J Virol. 2010 Jan;84(1):397-406.
  • Dar MJ, Monel B, Krishnan L, Shun MC, Di Nunzio F, Helland DE, Engelman A.Biochemical and virological analysis of the 18-residue C-terminal tail of HIV-1 integrase.Retrovirology. 2009;6:94.
  • Hare S, Di Nunzio F, Labeja A, Wang J, Engelman A, Cherepanov P.Structural basis for functional tetramerization of lentiviral integrase.PLoS Pathog. 2009 Jul;5(7):e1000515.
  • Belshan M, Schweitzer CJ, Donnellan MR, Lu R, Engelman A.In vivo biotinylation and capture of HIV-1 matrix and integrase proteins.J Virol Methods. 2009 Aug;159(2):178-84.
  • Engelman A, Oztop I, Vandegraaff N, Raghavendra NK.Quantitative analysis of HIV-1 preintegration complexes.Methods. 2009 Apr;47(4):283-90.
  • Engelman A.Mechanistic and pharmacological analyses of HIV-1 integration.Methods. 2009 Apr;47(4):225-8.
  • Yan N,Cherepanov P,Daigle JE,Engelman A,Lieberman J.The SET complex acts as a barrier to autointegration of HIV-1.PLoS Pathog. 2009 Mar;5(3):e1000327.
  • Hare S, Shun MC, Gupta SS, Valkov E, Engelman A, Cherepanov P.A novel co-crystal structure affords the design of gain-of-function lentiviral integrase mutants in the presence of modified PSIP1/LEDGF/p75.PLoS Pathog. 2009 Jan;5(1):e1000259.
  • Engelman A.Isolation and analysis of HIV-1 preintegration complexes.Methods Mol Biol. 2009;485:135-49.
  • McKee CJ, Kessl JJ, Shkriabai N, Dar MJ, Engelman A, Kvaratskhelia M.Dynamic modulation of HIV-1 integrase structure and function by cellular lens epithelium-derived growth factor (LEDGF) protein.J Biol Chem. 2008 Nov 14;283(46):31802-12.
  • Shun MC,Botbol Y,Li X,Di Nunzio F,Daigle JE,Yan N,Lieberman J,Lavigne M,Engelman A.Identification and characterization of PWWP domain residues critical for LEDGF/p75 chromatin binding and human immunodeficiency virus type 1 infectivity.J Virol. 2008 Dec;82(23):11555-67.
  • Byers KB, Engelman A, Fontes B.General guidelines for experimenting with HIV.Curr Protoc Immunol. 2004 May;Chapter 12:Unit 12.1. Review.
  • Engelman A, Cherepanov P.The lentiviral integrase binding protein LEDGF/p75 and HIV-1 replication.PLoS Pathog. 2008 Mar 7;4(3):e1000046. Review.
  • Brass AL, Dykxhoorn DM, Benita Y, Yan N, Engelman A, Xavier RJ, Lieberman J, Elledge SJ.Identification of host proteins required for HIV infection through a functional genomic screen.Science. 2008 Feb 15;319(5865):921-6.
  • Botbol Y, Raghavendra NK, Rahman S, Engelman A, Lavigne M.Chromatinized templates reveal the requirement for the LEDGF/p75 PWWP domain during HIV-1 integration in vitro.Nucleic Acids Res. 2008 Mar;36(4):1237-46.
  • Zhao Z, McKee CJ, Kessl JJ, Santos WL, Daigle JE, Engelman A, Verdine G, Kvaratskhelia M.Subunit-specific protein footprinting reveals significant structural rearrangements and a role for N-terminal Lys-14 of HIV-1 Integrase during viral DNA binding.J Biol Chem. 2008 Feb 29;283(9):5632-41.
  • Shun MC, Raghavendra NK, Vandegraaff N, Daigle JE, Hughes S, Kellam P, Cherepanov P, Engelman A.LEDGF/p75 functions downstream from preintegration complex formation to effect gene-specific HIV-1 integration.Genes Dev. 2007 Jul 15;21(14):1767-78.
  • Engelman A.AIDS/HIV. A reversal of fortune in HIV-1 integration.Science. 2007 Jun 29;316(5833):1855-7.
  • Mbisa JL, Barr R, Thomas JA, Vandegraaff N, Dorweiler IJ, Svarovskaia ES, Brown WL, Mansky LM, Gorelick RJ, Harris RS, Engelman A, Pathak VK.Human immunodeficiency virus type 1 cDNAs produced in the presence of APOBEC3G exhibit defects in plus-strand DNA transfer and integration.J Virol. 2007 Jul;81(13):7099-110.
  • Vandegraaff N, Engelman A.Molecular mechanisms of HIV integration and therapeutic intervention.Expert Rev Mol Med. 2007;9(6):1-19. Review.
  • Daelemans D, Lu R, De Clercq E, Engelman A.Characterization of a replication-competent, integrase-defective human immunodeficiency virus (HIV)/simian virus 40 chimera as a powerful tool for the discovery and validation of HIV integrase inhibitors.J Virol. 2007 Apr;81(8):4381-5.
  • Raghavendra NK, Engelman A.LEDGF/p75 interferes with the formation of synaptic nucleoprotein complexes that catalyze full-site HIV-1 DNA integration in vitro: implications for the mechanism of viral cDNA integration.Virology. 2007 Mar 30;360(1):1-5.
  • Shun MC, Daigle JE, Vandegraaff N, Engelman A.Wild-type levels of human immunodeficiency virus type 1 infectivity in the absence of cellular emerin protein.J Virol. 2007 Jan;81(1):166-72.
  • Anderson JL, Campbell EM, Wu X, Vandegraaff N, Engelman A, Hope TJ.Proteasome inhibition reveals that a functional preintegration complex intermediate can be generated during restriction by diverse TRIM5 proteins.J Virol. 2006 Oct;80(19):9754-60.
  • Rahman S, Lu R, Vandegraaff N, Cherepanov P, Engelman A.Structure-based mutagenesis of the integrase-LEDGF/p75 interface uncouples a strict correlation between in vitro protein binding and HIV-1 fitness.Virology. 2007 Jan 5;357(1):79-90.
  • Diaz-Griffero F, Vandegraaff N, Li Y, McGee-Estrada K, Stremlau M, Welikala S, Si Z, Engelman A, Sodroski J.Requirements for capsid-binding and an effector function in TRIMCyp-mediated restriction of HIV-1.Virology. 2006 Aug 1;351(2):404-19.
  • Maertens GN, Cherepanov P, Engelman A.Transcriptional co-activator p75 binds and tethers the Myc-interacting protein JPO2 to chromatin.J Cell Sci. 2006 Jun 15;119(Pt 12):2563-71.
  • Si Z, Vandegraaff N, O'huigin C, Song B, Yuan W, Xu C, Perron M, Li X, Marasco WA, Engelman A, Dean M, Sodroski J.Evolution of a cytoplasmic tripartite motif (TRIM) protein in cows that restricts retroviral infection.Proc Natl Acad Sci U S A. 2006 May 9;103(19):7454-9.
  • Turlure F, Maertens G, Rahman S, Cherepanov P, Engelman A.A tripartite DNA-binding element, comprised of the nuclear localization signal and two AT-hook motifs, mediates the association of LEDGF/p75 with chromatin in vivo.Nucleic Acids Res. 2006;34(5):1653-75.
  • Vandegraaff N, Devroe E, Turlure F, Silver PA, Engelman A.Biochemical and genetic analyses of integrase-interacting proteins lens epithelium-derived growth factor (LEDGF)/p75 and hepatoma-derived growth factor related protein 2 (HRP2) in preintegration complex function and HIV-1 replication.Virology. 2006 Mar 15;346(2):415-26.
  • Cherepanov P, Ambrosio AL, Rahman S, Ellenberger T, Engelman A.Structural basis for the recognition between HIV-1 integrase and transcriptional coactivator p75.Proc Natl Acad Sci U S A. 2005 Nov 29;102(48):17308-13.
  • Lu R, Vandegraaff N, Cherepanov P, Engelman A.Lys-34, dispensable for integrase catalysis, is required for preintegration complex function and human immunodeficiency virus type 1 replication.J Virol. 2005 Oct;79(19):12584-91.
  • Lu R, Ghory HZ, Engelman A.Genetic analyses of conserved residues in the carboxyl-terminal domain of human immunodeficiency virus type 1 integrase.J Virol. 2005 Aug;79(16):10356-68.
  • Cherepanov P, Sun ZY, Rahman S, Maertens G, Wagner G, Engelman A.Solution structure of the HIV-1 integrase-binding domain in LEDGF/p75.Nat Struct Mol Biol. 2005 Jun;12(6):526-32.
  • Lu R, Limón A, Ghory HZ, Engelman A.Genetic analyses of DNA-binding mutants in the catalytic core domain of human immunodeficiency virus type 1 integrase.J Virol. 2005 Feb;79(4):2493-505.
  • Engelman A.The ups and downs of gene expression and retroviral DNA integration.Proc Natl Acad Sci U S A. 2005 Feb 1;102(5):1275-6.
  • Devroe E, Silver PA, Engelman A.HIV-1 incorporates and proteolytically processes human NDR1 and NDR2 serine-threonine kinases.Virology. 2005 Jan 5;331(1):181-9.
  • Lu R, Limón A, Devroe E, Silver PA, Cherepanov P, Engelman A.Class II integrase mutants with changes in putative nuclear localization signals are primarily blocked at a postnuclear entry step of human immunodeficiency virus type 1 replication.J Virol. 2004 Dec;78(23):12735-46.
  • Cherepanov P, Devroe E, Silver PA, Engelman A.Identification of an evolutionarily conserved domain in human lens epithelium-derived growth factor/transcriptional co-activator p75 (LEDGF/p75) that binds HIV-1 integrase.J Biol Chem. 2004 Nov 19;279(47):48883-92.
  • Turlure F, Devroe E, Silver PA, Engelman A.Human cell proteins and human immunodeficiency virus DNA integration.Front Biosci. 2004 Sep 1;9:3187-208.
  • Maertens G, Cherepanov P, Debyser Z, Engelborghs Y, Engelman A.Identification and characterization of a functional nuclear localization signal in the HIV-1 integrase interactor LEDGF/p75.J Biol Chem. 2004 Aug 6;279(32):33421-9.
  • Lu R, Nakajima N, Hofmann W, Benkirane M, Jeang KT, Sodroski J, Engelman A, Teh-Jeang K.Simian virus 40-based replication of catalytically inactive human immunodeficiency virus type 1 integrase mutants in nonpermissive T cells and monocyte-derived macrophages.J Virol. 2004 Jan;78(2):658-68.
  • Devroe E, Engelman A, Silver PA.Intracellular transport of human immunodeficiency virus type 1 integrase.J Cell Sci. 2003 Nov 1;116(Pt 21):4401-8.
  • Engelman A.The roles of cellular factors in retroviral integration.Curr Top Microbiol Immunol. 2003;281:209-38.
  • Lin CW, Engelman A.The barrier-to-autointegration factor is a component of functional human immunodeficiency virus type 1 preintegration complexes.J Virol. 2003 Apr;77(8):5030-6.
  • Limón A, Nakajima N, Lu R, Ghory HZ, Engelman A.Wild-type levels of nuclear localization and human immunodeficiency virus type 1 replication in the absence of the central DNA flap.J Virol. 2002 Dec;76(23):12078-86.
  • Limón A, Devroe E, Lu R, Ghory HZ, Silver PA, Engelman A.Nuclear localization of human immunodeficiency virus type 1 preintegration complexes (PICs): V165A and R166A are pleiotropic integrase mutants primarily defective for integration, not PIC nuclear import.J Virol. 2002 Nov;76(21):10598-607.
  • Chen H, Engelman A.Asymmetric processing of human immunodeficiency virus type 1 cDNA in vivo: implications for functional end coupling during the chemical steps of DNA transposition.Mol Cell Biol. 2001 Oct;21(20):6758-67.
  • Nakajima N, Lu R, Engelman A.Human immunodeficiency virus type 1 replication in the absence of integrase-mediated dna recombination: definition of permissive and nonpermissive T-cell lines.J Virol. 2001 Sep;75(17):7944-55.
  • Harris D, Engelman A.Both the structure and DNA binding function of the barrier-to-autointegration factor contribute to reconstitution of HIV type 1 integration in vitro.J Biol Chem. 2000 Dec 15;275(50):39671-7.
  • Chen H, Engelman A.Characterization of a replication-defective human immunodeficiency virus type 1 att site mutant that is blocked after the 3' processing step of retroviral integration.J Virol. 2000 Sep;74(17):8188-93.
  • Brown HE, Chen H, Engelman A.Structure-based mutagenesis of the human immunodeficiency virus type 1 DNA attachment site: effects on integration and cDNA synthesis.J Virol. 1999 Nov;73(11):9011-20.
  • Engelman A.In vivo analysis of retroviral integrase structure and function.Adv Virus Res. 1999;52:411-26.
  • Chen H, Wei SQ, Engelman A.Multiple integrase functions are required to form the native structure of the human immunodeficiency virus type I intasome.J Biol Chem. 1999 Jun 11;274(24):17358-64.
  • Chen H, Engelman A.The barrier-to-autointegration protein is a host factor for HIV type 1 integration.Proc Natl Acad Sci U S A. 1998 Dec 22;95(26):15270-4.
  • Jenkins TM, Esposito D, Engelman A, Craigie R.Critical contacts between HIV-1 integrase and viral DNA identified by structure-based analysis and photo-crosslinking.EMBO J. 1997 Nov 17;16(22):6849-59.
  • Du Z, Ilyinskii PO, Lally K, Desrosiers RC, Engelman A.A mutation in integrase can compensate for mutations in the simian immunodeficiency virus att site.J Virol. 1997 Nov;71(11):8124-32.
  • Engelman A, Liu Y, Chen H, Farzan M, Dyda F.Structure-based mutagenesis of the catalytic domain of human immunodeficiency virus type 1 integrase.J Virol. 1997 May;71(5):3507-14.
  • Taddeo B, Carlini F, Verani P, Engelman A.Reversion of a human immunodeficiency virus type 1 integrase mutant at a second site restores enzyme function and virus infectivity.J Virol. 1996 Dec;70(12):8277-84.
  • Engelman A.Biochemical characterization of recombinant equine infectious anemia virus integrase.Protein Expr Purif. 1996 Nov;8(3):299-304.
  • Jenkins TM, Engelman A, Ghirlando R, Craigie R.A soluble active mutant of HIV-1 integrase: involvement of both the core and carboxyl-terminal domains in multimerization.J Biol Chem. 1996 Mar 29;271(13):7712-8.
  • Engelman A, Craigie R.Efficient magnesium-dependent human immunodeficiency virus type 1 integrase activity.J Virol. 1995 Sep;69(9):5908-11.
  • Lodi PJ, Ernst JA, Kuszewski J, Hickman AB, Engelman A, Craigie R, Clore GM, Gronenborn AM.Solution structure of the DNA binding domain of HIV-1 integrase.Biochemistry. 1995 Aug 8;34(31):9826-33.
  • Englund G, Theodore TS, Freed EO, Engleman A, Martin MA.Integration is required for productive infection of monocyte-derived macrophages by human immunodeficiency virus type 1.J Virol. 1995 May;69(5):3216-9.
  • Engelman A, Englund G, Orenstein JM, Martin MA, Craigie R.Multiple effects of mutations in human immunodeficiency virus type 1 integrase on viral replication.J Virol. 1995 May;69(5):2729-36.
  • Dyda F, Hickman AB, Jenkins TM, Engelman A, Craigie R, Davies DR.Crystal structure of the catalytic domain of HIV-1 integrase: similarity to other polynucleotidyl transferases.Science. 1994 Dec 23;266(5193):1981-6.
  • Vicenzi E, Dimitrov DS, Engelman A, Migone TS, Purcell DF, Leonard J, Englund G, Martin MA.An integration-defective U5 deletion mutant of human immunodeficiency virus type 1 reverts by eliminating additional long terminal repeat sequences.J Virol. 1994 Dec;68(12):7879-90.
  • Hickman AB, Palmer I, Engelman A, Craigie R, Wingfield P.Biophysical and enzymatic properties of the catalytic domain of HIV-1 integrase.J Biol Chem. 1994 Nov 18;269(46):29279-87.
  • Engelman A.Most of the avian genome appears available for retroviral DNA integration.Bioessays. 1994 Nov;16(11):797-9.
  • Engelman A, Hickman AB, Craigie R.The core and carboxyl-terminal domains of the integrase protein of human immunodeficiency virus type 1 each contribute to nonspecific DNA binding.J Virol. 1994 Sep;68(9):5911-7.
  • Mazumder A, Engelman A, Craigie R, Fesen M, Pommier Y.Intermolecular disintegration and intramolecular strand transfer activities of wild-type and mutant HIV-1 integrase.Nucleic Acids Res. 1994 Mar 25;22(6):1037-43.
  • Engelman A, Bushman FD, Craigie R.Identification of discrete functional domains of HIV-1 integrase and their organization within an active multimeric complex.EMBO J. 1993 Aug;12(8):3269-75.
  • Bushman FD, Engelman A, Palmer I, Wingfield P, Craigie R.Domains of the integrase protein of human immunodeficiency virus type 1 responsible for polynucleotidyl transfer and zinc binding.Proc Natl Acad Sci U S A. 1993 Apr 15;90(8):3428-32.
  • Engelman A, Craigie R.Identification of conserved amino acid residues critical for human immunodeficiency virus type 1 integrase function in vitro.J Virol. 1992 Nov;66(11):6361-9.
  • Engelman A, Mizuuchi K, Craigie R.HIV-1 DNA integration: mechanism of viral DNA cleavage and DNA strand transfer.Cell. 1991 Dec 20;67(6):1211-21.
  • Craigie R, Mizuuchi K, Bushman FD, Engelman A.A rapid in vitro assay for HIV DNA integration.Nucleic Acids Res. 1991 May 25;19(10):2729-34.
  • Engelman A, Rosenberg N.Temperature-sensitive mutants of Abelson murine leukemia virus deficient in protein tyrosine kinase activity.J Virol. 1990 Sep;64(9):4242-51.
  • Engelman A, Rosenberg N.bcr/abl and src but not myc and ras replace v-abl in lymphoid transformation.Mol Cell Biol. 1990 Aug;10(8):4365-9.
  • Engelman A, Rosenberg N.The Abelson protein is required for initiation and maintenance for transformation in murine pre-B cells.Curr Top Microbiol Immunol. 1988;141:310-5.
  • Engelman A, Rosenberg N.Isolation of temperature-sensitive Abelson virus mutants by site-directed mutagenesis.Proc Natl Acad Sci U S A. 1987 Nov;84(22):8021-5.