Countryman Jill K, Gradoville Lyndle, Miller George
Department of Molecular Biophysics and Biochemistry, Yale University School of Medicine, New Haven, CT 06520, USA.
J Virol. 2008 May;82(10):4706-19. doi: 10.1128/JVI.00116-08. Epub 2008 Mar 12.
Activation of the Epstein-Barr virus (EBV) lytic cycle is mediated through the combined actions of ZEBRA and Rta, the products of the viral BZLF1 and BRLF1 genes. During latency, these two genes are tightly repressed. Histone deacetylase inhibitors (HDACi) can activate viral lytic gene expression. Therefore, a widely held hypothesis is that Zp and Rp, the promoters for BZLF1 and BRLF1, are repressed by chromatin and that hyperacetylation of histone tails, by allowing the access of positively acting factors, leads to transcription of BZLF1 and BRLF1. To investigate this hypothesis, we used chromatin immunoprecipitation (ChIP) to examine the acetylation and phosphorylation states of histones H3 and H4 on Zp and Rp in three cell lines, Raji, B95-8, and HH514-16, which differ in their response to EBV lytic induction by HDACi. We studied the effects of three HDACi, sodium butyrate (NaB), trichostatin A (TSA), and valproic acid (VPA). We also examined the effects of tetradecanoyl phorbol acetate (TPA) and 5-aza-2'-deoxycytidine, a DNA methyltransferase inhibitor, on histone modification. In Raji cells, TPA and NaB act synergistically to activate the EBV lytic cycle and promote an increase in histone H3 and H4 acetylation and phosphorylation at Zp and Rp. Surprisingly, however, when Raji cells were treated with NaB or TSA, neither of which is sufficient to activate the lytic cycle, an increase of comparable magnitude of hyperacetylated and phosphorylated histone H3 at Zp and Rp was observed. In B95-8 cells, NaB inhibited lytic induction by TPA, yet NaB promoted hyperacetylation of H3 and H4. In HH514-16 cells, NaB and TSA strongly activated the EBV lytic cycle and caused hyperacetylation of histone H3 on Zp and Rp. However, when HH514-16 cells were treated with VPA, lytic cycle mRNAs or proteins were not induced, although histone H3 was hyperacetylated as measured by immunoblotting or by ChIP on Zp and Rp. Taken together, our data suggest that open chromatin at EBV BZLF1 and BRLF1 promoters is not sufficient to activate EBV lytic cycle gene expression.
爱泼斯坦-巴尔病毒(EBV)裂解周期的激活是通过ZEBRA和Rta的联合作用介导的,它们分别是病毒BZLF1和BRLF1基因的产物。在潜伏期间,这两个基因受到严格抑制。组蛋白脱乙酰酶抑制剂(HDACi)可以激活病毒裂解基因的表达。因此,一个广泛接受的假说是,Zp和Rp(BZLF1和BRLF1的启动子)被染色质抑制,并且组蛋白尾巴的超乙酰化通过允许正向作用因子的进入,导致BZLF1和BRLF1的转录。为了研究这个假说,我们使用染色质免疫沉淀(ChIP)来检测在三种细胞系Raji、B95-8和HH514-16中,Zp和Rp上组蛋白H3和H4的乙酰化和磷酸化状态,这三种细胞系对HDACi诱导的EBV裂解反应不同。我们研究了三种HDACi,丁酸钠(NaB)、曲古抑菌素A(TSA)和丙戊酸(VPA)的作用。我们还检测了十四酰佛波醇乙酸酯(TPA)和DNA甲基转移酶抑制剂5-氮杂-2'-脱氧胞苷对组蛋白修饰的影响。在Raji细胞中,TPA和NaB协同作用激活EBV裂解周期,并促进Zp和Rp处组蛋白H3和H4乙酰化和磷酸化的增加。然而,令人惊讶的是,当用NaB或TSA处理Raji细胞时,它们都不足以激活裂解周期,但在Zp和Rp处观察到组蛋白H3超乙酰化和磷酸化的增加幅度相当。在B95-8细胞中,NaB抑制TPA诱导的裂解,但NaB促进H3和H4的超乙酰化。在HH514-16细胞中,NaB和TSA强烈激活EBV裂解周期,并导致Zp和Rp上组蛋白H3的超乙酰化。然而,当用VPA处理HH514-16细胞时,虽然通过免疫印迹或ChIP检测到Zp和Rp上组蛋白H3超乙酰化,但未诱导出裂解周期的mRNA或蛋白质。综上所述,我们的数据表明,EBV BZLF1和BRLF1启动子处的开放染色质不足以激活EBV裂解周期基因的表达。