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Susan Hafenstein

Professor of Biochemistry and Molecular Biology, and The College of Medicine
Image of Susan Hafenstein

Department or University Committees

  • Promotion and Tenure Committee, Biochemistry and Molecular Biology Department
  • Peer Teaching Review Committee, Biochemistry and Molecular Biology Department


Program or Departmental Affiliations

The BMMB Graduate Program Molecular, Cellular, and Integrative Biosciences Program 


Editorial Boards

NSF Review EMSL Review



Center of Structural Oncology Penn State Cancer Institute Center for Infectious Disease Dynamics


Research Summary

We use a structure/function approach to study virus and host interactions.


Selected Publications

  • Subramanian S, Maurer AC, Bator CM, Makhov AM, Conway JF, Marden, JH, Vandenberghe L, Hafenstein S. Filling AAV capsids: estimating success by cryo-EM. Human Gene Therapy (accepted, in press).
  • Lee H, Callaway HM, Bator CM, Parrish CR, Hafenstein SL. Structure of canine parvovirus in complex with transferrin receptor type-1 explains the controls of viral infection and host range. PNAS (accepted, in press)
  • Goetschius DJ, Parrish CR, Hafenstein S. Asymmetry in icosahedral viruses. Curr Opin Virol. 2019 Jun;36:67-73. doi: 10.1016/j.coviro.2019.05.006. Epub 2019 Jun 28. Review. PMID: 31255982
  • Heather Callaway, Kathrin Welsch, Wendy Weichert, Andrew Allison, Susan Hafenstein, Kai Huang, Sho Iketani, and Colin Parrish. Complex and dynamic interactions between parvovirus capsids, transferrin receptors and antibodies control cell infection and host range J Virol. 2018 Apr 25. pii: JVI.00460-18. doi: 10.1128/JVI.00460-18 (JVI00460-18R1)
  • Minns AM, Hart KJ, Subramanian S, Hafenstein S, Lindner SE. Nuclear, Cytosolic, and Surface-Localized Poly(A)-Binding Proteins of Plasmodium yoelii. mSphere. 2018 Jan 10;3(1). pii: e00435-17. doi: 10.1128/mSphere.00435-17. eCollection 2018 Jan-Feb. PMID: 29359180
  • Guan J, Bywaters SM, Brendle SA, Ashley RE, Makhov AM, Conway JF, Christensen ND, Hafenstein S. High-Resolution Structure Analysis of Antibody V5 and U4 Conformational Epitopes on Human Papillomavirus 16. Viruses. 2017 Dec 6;9(12). pii: E374. doi: 10.3390/v9120374 PMID: 29211035
  • Subramanian S, Organtini LJ, Grossman A, Domeier PP, Cifuente JO, Makhov AM, Conway JF, D'Abramo A Jr, Cotmore SF, Tattersall P, Hafenstein S. Cryo-EM maps reveal five-fold channel structures and their modification by gatekeeper mutations in the parvovirus minute virus of mice (MVM) capsid. Virology. 2017 Oct;510:216-223. doi: 10.1016/j.virol.2017.07.015. Epub 2017 Jul 24. PMID: 28750325 
  • Guan J, Bywaters SM, Brendle SA, Lee H, Ashley RE, Christensen ND, Hafenstein S. (2016) Human Cryoelectron Microscopy Maps of Human Papillomavirus 16 Reveal L2 Densities and Heparin Binding Site. Structure. 2017 Feb 7;25(2):253-263. doi: 10.1016/j.str.2016.12.001. Epub 2017 Jan 5.PMID: 28065506
  • Carson SD, Hafenstein S, Lee H. (2017) MOPS and coxsackievirus B3 stability. Virology. Jan 15;501:183-187. doi: 10.1016/j.virol.2016.12.002. PMID: 27940223
  • Organtini LJ, Shingler KL, Durrani K, Capaldi E, Ashley RE, Dryden K, Makhov AM, Conway JF, Pizzorno M, Hafenstein S. (2016) Honey bee deformed wing virus structures reveal changes required for genome release. J Virol. 2017 Jan 3;91(2). pii: e01795-16. doi: 10.1128/JVI.01795-16. PMID: 27852845 Selected as Spotlight by Editor
  • Hafenstein S. (2016) Structural biology: A picornavirus unlike the others. Nature Microbiology. 26;1(11):16217. doi: 10.1038/nmicrobiol.2016.217 
  • Organtini LJ, Iketani S, Huang K, Ashley RE, Makhov AM, Conway JF, Parrish CR, Hafenstein S. (2016) Near-atomic resolution structure of a highly neutralizing Fab complexed with a virus capsid. Journal of Virology. 90(21):9733-9742. PMID: 27535057
  • Lee H*, Shingler KL*, Organtini LJ, Ashley RE, Subramanian S, Makhov AM, Conway JF, Hafenstein S. (2016)The novel asymmetric entry intermediate of a picornavirus captured with nanodiscs. Science Advances 2(8):e1501929. doi: 10.1126/sciadv.1501929. *first two authors contributed equally.
  • Guan J, Bywaters SM, Brendle SA, Lee H, Ashley RE, Christensen ND, Hafenstein S. (2015) The U4 antibody epitope on human papillomavirus 16 identified by cryo-EM. Journal of Virology. 89(23):12108-17. doi: 10.1128/JVI.02020-15. PMID: 264010
  • Guan J, Bywaters SM, Brendle SA, Lee H, Ashley RE, Makhov AM, Conway JF, Christensen ND, Hafenstein S. (2015) Structural comparison of four different neutralizing antibodies interacting with human papillomavirus 16 and mechanisms of neutralization. Virology. 483:253-63. doi: 10.1016/j.virol.2015.04.016. PMID: 25996608
  • 11(5):e1004838. doi: 10.1371/journal.ppat.1004838. PMID: 25941809
  • Shingler KL, Cifuente JO, Ashley RE, Makhov AM, Conway JF, Hafenstein S. (2015) The Enterovirus 71 procapsid binds neutralizing antibodies and rescues virus infection in vitro. Journal of Virology. 89(3):1900-8. doi: 10.1128/JVI.03098-14. PMID: 25428877 
  • Organtini LJ, Allison AB, Lukk T, Parrish CR, Hafenstein S. (2015) Global displacement of canine parvovirus by a host-adapted variant: A structural comparison between pandemic viruses. Journal of Virology. 89(3):1909-12. doi: 10.1128/JVI.02611-14. PMID: 25410876
  • Lee H, Brendle SA, Bywaters SM, Guan J, Ashley RE, Yoder JD, Makhov AM, Conway JF, Christensen ND, Hafenstein S. (2015) A Cryo-electron microscopy study reveals the mechanism of HPV capsid stabilization used by neutralizing antibody H16.V5 and identifies the complete epitope. Journal of Virology. 15;89(2):1428-38. doi: 10.1128/JVI.02898-14. PMID: 25392224
  • Shingler KL, Organtini LJ, Hafenstein S. (2014) Enterovirus 71 Propagation and Purification Biological Protocols. 5;4(9). pii: e1117. PMID: 27547789
  • Organtini LJ, Makhov AM, Conway JF, Hafenstein S*, Carson SD*. (2014) Kinetic and structural analysis of coxsackievirus B3 receptor interactions and formation of the A-particle. Journal of Virology. 88(10):5755-65. doi: 10.1128/JVI.00299-14. PMID: 24623425 *last two authors share corresponding authorship 
  • Lee H, Cifuente JO, Ashley RE, Conway JF, Makhov AM, Tano Y, Shimizu H, Nishimura Y, Hafenstein S. (2013) A strain-specific epitope of Enterovirus 71 identified by cryo-EM of the complex with Fab from neutralizing antibody. Journal of Virology. 87(21):11363-70. doi: 10.1128/JVI.01926-13. PMID: 23946455 
  • Cifuente JO, Lee H, Yoder JD, Shingler KL, Carnegie MS, Yoder JL, Ashley RE, Makhov AM, Conway JF, Hafenstein S. (2013) The EV71 strain 1095 structures of procapsid and mature virion. Journal of Virology. 87(13):7637-45. doi: 10.1128/JVI.03519-12. PMID: 23637404 
  • Shingler KL, Yoder JL, Carnegie MS, Ashley RE, Makhov AM, Conway JF, Hafenstein SL. (2013) The Enterovirus 71 A-Particle Forms a Gateway to Allow Genome Release: A Cryo-EM Study of Picornavirus Uncoating. PLoS Pathogens. 2013 Mar;9(3):e1003240. doi: 10.1371/journal.ppat.1003240. Erratum in: PLoS Pathog. 9(9). doi: 10.1371/annotation/e92d19e0-996a-4bfa-afdd-20dce770ed75. PMID: 23555253 
  • Yoder JD, Cifuente JO, Pan J, Bergelson JM, Hafenstein S. (2012) The crystal structure of a coxsackievirus B3-RD variant and a refined 9Ǻ cryo-EM reconstruction of the virus complexed with decay accelerating factor (DAF), provide a new footprint of DAF onto the virus surface. Journal of Virology. 86(23):12571-81. doi: 10.1128/JVI.01592-12. PMID: 22973031 
  • Cifuente JO, Ferrer MF, de Giusti CJ, Song WC, Romanowski V, Hafenstein S, Gomez RM. (2011) Molecular determinants of disease in coxsackievirus B1 murine infection. Journal of Medical Virology. 83(9):1571-81. doi: 10.1002/jmv.22133. PMID: 21739448 
  • Pan J, Narayanan B, Shah S, Yoder JD, Cifuente JO, Hafenstein S, Bergelson JM. (2011) Single amino acid changes in the virus capsid permit coxsackievirus B3 to bind decay accelerating factor. Journal of Virology. 85(14):7436-43. doi: 10.1128/JVI.00503-11. PMID: 2156196 
  • Plevka P*, Hafenstein S*, Harris K, Cifuente JO, Zhang Y, Bowman VD, Chipman PR, Bator CM, Lin F, Medof ME, Rossmann MG. (2010) The interaction of Decay-accelerating factor with Echovirus 7. Journal of Virology. 84(24):12665-74. doi: 10.1128/JVI.00837-10. PMID: 20881044 *First two authors contributed equally
  • Hafenstein S, Bowman VD, Sun T, Nelson CDS, Palermo LM, Chipman PR, Battisti AJ, Parrish CR, Rossmann MG. (2009) Structural comparison of different antibodies interacting with parvovirus capsids. Journal of Virology. 83(11):5556-66. doi: 10.1128/JVI.02532-08 PMID: 19321620 
  • Hafenstein S, Chipman PR, Bowman VD, Kelly CB, Lin F, Medof ME, Rossmann MG. (2007) The interaction of CVB3 with decay accelerating factor. Journal of Virology. 81(23):12927-35 PMID: 17804498 
  • Hafenstein S, Palermo LM, Xiao C, Kostyuchenko VA, Morais M, Nelson CDS, Chipman PR, Bowman VD, Battisti AJ, Parrish CR, Rossmann MG. (2007) Asymmetric binding of transferrin receptor to parvovirus capsids. Proceedings of the National Academy of Sciences USA. 104(16):6585-9. PMID: 17420467 
  • Rossmann MG, Arisaka F, Battisti AJ, Bowman VD, Chipman PR, Fokine A, Hafenstein S, Kanamaru S, Kostyuchenko VA, Mesyanzhinov VV, Shneider MM, Morais MC, Leiman PG, Palermo LM, Parrish CR, Xiao C. (2007) From structure of the complex to understanding of the biology. Acta Crystallographica D Biological Crystallography. 63(Pt 1):9-16. PMID: 17164521 
  • Bernal RA, Hafenstein S, Esmeralda R, Fane BA, Rossmann MG. (2004) The phiX174 protein J mediates DNA packaging and viral attachment to host cells. Journal of Molecular Biology. 337(5):1109-22.PMID: 15046981 
  • Hafenstein SL, Chen M, Fane BA. (2004) Genetic and functional analyses of the phiX174 DNA binding protein: the effects of substitutions for amino acid residues that spatially organize the two DNA binding domains. Virology. 318(1):204-13. PMID: 14972548 
  • Hafenstein S, Fane BA. (2002) phiX174 genome-capsid interactions influence the biophysical properties of the virion: evidence for a scaffolding-like function of the genome during the final stages of morphogenesis. Journal of Virology. 76(11):5350-6. PMID: 11991963