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. 2017 Jul 25;8(41):70006–70034. doi: 10.18632/oncotarget.19549

Table 1. Nuclear localization summary of EBV-encoded proteins.

No. Protein name(s) Function/description # COS-7 subcellular localization Literature localization and references
1 BORF1 c Assembly/maturation, minor capsid protein-binding protein (mCP-BP), homology with triplex protein HSV-1 VP19C, VZV ORF20, MCMV M46 and KSHV ORF62 Subnuclear Pan-nuclear a [6] Speckles in the nucleus b [48]
2 BSLF2/BMLF1 t (EB2, Mta, SM) mRNA-export factor, mRNA splicing, interaction with human Spen proteins, homology with HSV-1 UL54 (ICP27), VZV ORF4, MCMV M69 and KSHV ORF57 Subnuclear, nucleolus like N/A
3 EBNA3A l (BLRF3/BERF1) Latency nuclear antigen 3A, negative effect on transactivator EBNA-2 and cell cycle, essential for immortalization Pan-nuclear, without nucleolus Pan-nuclear a [6, 49]
4 BZLF1 t (Zta, Z, Zebra, EB1) Trans-activator ZEBRA, origin binding protein (EB1, Zta), bZip similar to CCAAT/enhancer binding protein, homology with KSHV K08 Pan-nuclear, without nucleolus Pan-nuclear a [6] Nuclear b [50]
5 RPMS1 u Unknown function Subnuclear, nucleolus like Pan-nuclear a [34, 51]
6 BMLF1 t
(Mta)
Immediate-early transactivator, part of BSLF2/BMLF1 protein Subnuclear, nucleolus like Nuclear and speckles in the nucleus a [52, 53] Nuclear b [50]
7 BXRF1 u Nucleoprotein, homology with HSV-1 UL24, VZV ORF35, MCMV M76 and KSHV ORF20 Subnuclear, nucleolus like N/A
8 BALF2 r Major ssDNA binding protein, tegument protein, part of replication fork / machinery, homology with HSV-1 UL29, VZV ORF29, MCMV M57 and KSHV ORF6 Pan-nuclear, without nucleolus Pan-nuclear a [6]
9 BMRF1 r DNA polymerase processivity factor, transcriptional transactivators, sliding clamp, early antigen protein D (EA-D, polymerase accessory protein), homology with HSV-1 UL42, VZV ORF16, MCMV M44 and KSHV ORF59 Pan-nuclear Pan-nuclear a [6]
10 BLLF2 T Hypothetical protein, unknown function Subnuclear, nucleolus like Pan-nuclear a [6]
11 BFLF2 m Nuclear membrane phosphoprotein, part of intracellular virions, egress protein, herpesvirus UL31-like protein, complex with BFRF1, homology with HSV-1 UL31, VZV ORF27, MCMV M53 and KSHV ORF69 Pan-Nuclear Nuclear a [54]
12 BdRF1 c Capsid scaffolding protein, homology with HSV-1 UL26.5, VZV ORF33.5, MCMV M80.5 and KSHV ORF17.5 Pan-nuclear, without nucleolus N/A
13 EBNA1 l
(BKRF1)
Latency nuclear antigen 1, assures EBV episome maintenance replication, Gly-rich domain, essential for immortalization Pan-nuclear Nuclear a/b [55]
14 BLLF3 n dUTP pyrophosphatase, dUTPase, homology with HSV-1 UL50, VZV ORF8, MCMV M72 and KSHV ORF54 Nuclear obviously > Cytoplasmic, without nucleolus Pan-Nuclear a [6]
15 BGLF5 r Alkaline exonuclease, involved together with BALF2 in DNA recombination, homology with HSV-1 UL12, VZV ORF48, MCMV M98 and KSHV ORF37 Nuclear obviously > Cytoplasmic, without nucleolus Nucleus and cytoplasm a [56]
Nuclear a [57]
16 BRRF1 t Na, transcription factor, enhancement of the induction of the lytic cycle, homology with KSHV ORF49 Nuclear obviously > Cytoplasmic, without nucleolus Nuclear a [58]
17 BCRF2 u Unknown function Nuclear obviously > Cytoplasmic, without nucleolus N/A
18 BDLF4 u Putative metal binding protein (gp115), homology with MCMV M92 and KSHV ORF31 Nuclear > Cytoplasmic Pan-Cellular a [6]
19 BARF0 u Start at amino acid 298 of SWISS-PROT, homology with rhesus LCV Nuclear obviously > Cytoplasmic Nuclear a [59, 60]
20 BBLF1 T Myristoylated phosphoprotein in tegument (MyrP), homology with HSV-1 UL11, VZV ORF49, MCMV M99 and KSHV ORF38 Nuclear obviously > Cytoplasmic, without nucleolus Pan-Cellular or Cytoplasmic > Nuclear a [6]
trans-Golgi network (TGN) a/b [20]
21 BFRF3 c Smallest capsid protein (sCP) on outer capsid surface, homology with HSV-1 UL35 (VP26), VZV ORF23, MCMV M48.2 and KSHV ORF65 Nuclear obviously > Cytoplasmic, without nucleolus Pan-Nuclear a [6]
Nuclear b [61]
22 BGLF3 T gp118, not included in virions, homology with HSV-1 UL14, VZV ORF46, MCMV M95 and KSHV ORF34 Nuclear obviously > Cytoplasmic Nuclear > Cytoplasmic a [6]
23 BGLF4 n Ser/Thr kinase, phosphorylation of nucleoside analogues, homology with HSV-1 UL13, VZV ORF47, MCMV M97 and KSHV ORF36 Nuclear obviously > Cytoplasmic Nuclear a [62]
Nuclear > Cytoplasmic a [6]
Nuclear, a small portion is distributed in the cytoplasm at the late stage of virus replication b [63]
24 BKRF4 T Tegument phosphoprotein, homology with KSHV ORF45 Nuclear obviously > Cytoplasmic some with multiple small foci in the nucleus Pan-Nuclear a [6]
25 BRLF1 t
(Rta, R)
Transcriptional activator dimeric (TAF50), homology with KSHV ORF50 Nuclear obviously > Cytoplasmic, Cytoplasmic > Nuclear, Pan-cellular Pan-Nuclear, Pan-Cytoplasmic, Pan-Cellular a [6, 64]
Cytoplasmic and nuclear b [50]
26 BLRF2 T Tegument protein, homology with KSHV ORF52 Nuclear > Cytoplasmic, with speckles Nuclear > Cytoplasmic a [6]
Relocalized from the nucleus to the cytoplasm b [19]
27 BVRF2 c Proteinase/scaffold protein, capsid maturational protease, homology with HSV-1 UL26, VZV ORF33, MCMV M80 and KSHV ORF17 Nuclear slightly > Cytoplasmic Pan-Nuclear a [6]
28 BOLF1 T Tegument large tegument protein-binding protein (LTPBP) Nuclear obviously > Cytoplasmic, without nucleolus N/A

a: transfection

b: infection

N/A: Not Detected

r: replication related protein

n: nucleotide metabolism related protein

t: transcription related protein

p: packaging related protein

l: latency related protein

m: membrane (or glycosylated) protein

c: capsid related protein

T: tegument protein

u: unkown function protein

# Function/Description column compiled the following references: Tarbouriech et al. [5] and Salsman et al. [6].