The EBNA-2 N-Terminal Transactivation Domain Folds into a Dimeric Structure Required for Target Gene Activation

PLoS Pathog. 2015 May 29;11(5):e1004910. doi: 10.1371/journal.ppat.1004910. eCollection 2015 May.

Abstract

Epstein-Barr virus (EBV) is a γ-herpesvirus that may cause infectious mononucleosis in young adults. In addition, epidemiological and molecular evidence links EBV to the pathogenesis of lymphoid and epithelial malignancies. EBV has the unique ability to transform resting B cells into permanently proliferating, latently infected lymphoblastoid cell lines. Epstein-Barr virus nuclear antigen 2 (EBNA-2) is a key regulator of viral and cellular gene expression for this transformation process. The N-terminal region of EBNA-2 comprising residues 1-58 appears to mediate multiple molecular functions including self-association and transactivation. However, it remains to be determined if the N-terminus of EBNA-2 directly provides these functions or if these activities merely depend on the dimerization involving the N-terminal domain. To address this issue, we determined the three-dimensional structure of the EBNA-2 N-terminal dimerization (END) domain by heteronuclear NMR-spectroscopy. The END domain monomer comprises a small fold of four β-strands and an α-helix which form a parallel dimer by interaction of two β-strands from each protomer. A structure-guided mutational analysis showed that hydrophobic residues in the dimer interface are required for self-association in vitro. Importantly, these interface mutants also displayed severely impaired self-association and transactivation in vivo. Moreover, mutations of solvent-exposed residues or deletion of the α-helix do not impair dimerization but strongly affect the functional activity, suggesting that the EBNA-2 dimer presents a surface that mediates functionally important intra- and/or intermolecular interactions. Our study shows that the END domain is a novel dimerization fold that is essential for functional activity. Since this specific fold is a unique feature of EBNA-2 it might provide a novel target for anti-viral therapeutics.

Publication types

  • Research Support, Non-U.S. Gov't

MeSH terms

  • Adult
  • Amino Acid Sequence
  • Blotting, Western
  • Crystallography, X-Ray
  • Epstein-Barr Virus Nuclear Antigens / chemistry*
  • Epstein-Barr Virus Nuclear Antigens / genetics
  • Fluorescent Antibody Technique
  • HeLa Cells
  • Humans
  • Immunoenzyme Techniques
  • Immunoprecipitation
  • Molecular Sequence Data
  • Mutant Proteins / chemistry*
  • Mutant Proteins / genetics
  • Mutation / genetics
  • Nuclear Magnetic Resonance, Biomolecular
  • Protein Conformation
  • Protein Multimerization
  • Protein Structure, Tertiary
  • RNA, Messenger / genetics
  • Real-Time Polymerase Chain Reaction
  • Reverse Transcriptase Polymerase Chain Reaction
  • Sequence Homology, Amino Acid
  • Trans-Activators / genetics*
  • Transcriptional Activation*
  • Viral Proteins / chemistry*
  • Viral Proteins / genetics

Substances

  • EBNA-2 protein, Human herpesvirus 4
  • Epstein-Barr Virus Nuclear Antigens
  • Mutant Proteins
  • RNA, Messenger
  • Trans-Activators
  • Viral Proteins

Associated data

  • PDB/2N2J

Grants and funding

AF was supported by a PhD fellowship from Helmholtz Zentrum München (HMGU), and by the International PhD program in Protein Dynamics from Elitenetzwerk Bayern. JH gratefully acknowledges the Swedish Research Council (Vetenskapsrådet) and the European Molecular Biology Organization (EMBO, ALTF-276-2010) for postdoctoral fellowships. ST was supported by the HELENA graduate school. The authors acknowledge NMR measurement time at the Bavarian NMR Centre, Garching, Germany. This work was supported by the Deutsche Forschungsgemeinschaft, GRK1721 (MS) and SFB1035 (MS), by the Deutsche Krebshilfe 109258 (BK) and Deutsche Carreras Leukämie Stiftung DJCLS R07/11 (BK). The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.