Clarke Christopher J, Shamseddine Achraf A, Jacob Joseph J, Khalife Gabrielle, Burns Tara A, Hannun Yusuf A
Department of Medicine and Stony Brook Cancer Center, Stony Brook University, Stony Brook, NY.
Department of Biochemistry and Molecular Biology, Medical University of South Carolina, Charleston, SC.
J Lipid Res. 2016 May;57(5):868-81. doi: 10.1194/jlr.M067447. Epub 2016 Mar 24.
Neutral sphingomyelinase-2 (nSMase2) is a key ceramide-producing enzyme in cellular stress responses. While many posttranslational regulators of nSMase2 are known, emerging evidence suggests a more protracted regulation of nSMase2 at the transcriptional level. Previously, we reported that nSMase2 is induced by all-trans retinoic acid (ATRA) in MCF7 cells and implicated nSMase2 in ATRA-induced growth arrest. Here, we further investigated how ATRA regulates nSMase2. We find that ATRA regulates nSMase2 transcriptionally through the retinoic acid receptor-α, but this is independent of previously identified transcriptional regulators of nSMase2 (Sp1, Sp3, Runx2) and is not through increased promoter activity. Epigenetically, the nSMase2 gene is not repressively methylated in MCF7 cells. However, inhibition of histone deacetylases (HDACs) with trichostatin A (TSA) induced nSMase2 comparably to ATRA; furthermore, combined ATRA and TSA treatment was not additive, suggesting ATRA regulates nSMase2 through direct modulation of histone acetylation. Confirming this, the histone acetyltransferases CREB-binding protein and p300 were required for ATRA induction of nSMase2. Finally, use of class-specific HDAC inhibitors suggested that HDAC4 and/or HDAC5 are negative regulators of nSMase2 expression. Collectively, these results identify a novel pathway of nSMase2 regulation and suggest that physiological or pharmacological modulation of histone acetylation can directly affect nSMase2 levels.
中性鞘磷脂酶2(nSMase2)是细胞应激反应中产生神经酰胺的关键酶。虽然已知许多nSMase2的翻译后调节因子,但新出现的证据表明nSMase2在转录水平上受到更持久的调节。此前,我们报道nSMase2在MCF7细胞中由全反式维甲酸(ATRA)诱导,并参与了ATRA诱导的生长停滞。在此,我们进一步研究了ATRA如何调节nSMase2。我们发现ATRA通过维甲酸受体-α在转录水平上调节nSMase2,但这独立于先前确定的nSMase2转录调节因子(Sp1、Sp3、Runx2),且不是通过增加启动子活性。在表观遗传学上,nSMase2基因在MCF7细胞中没有被抑制性甲基化。然而,用曲古抑菌素A(TSA)抑制组蛋白去乙酰化酶(HDACs)与ATRA诱导nSMase水平相当;此外,ATRA和TSA联合处理没有叠加效应,表明ATRA通过直接调节组蛋白乙酰化来调节nSMase2。证实这一点的是,组蛋白乙酰转移酶CREB结合蛋白和p300是ATRA诱导nSMase2所必需的。最后,使用类特异性HDAC抑制剂表明HDAC4和/或HDAC5是nSMase2表达的负调节因子。总的来说,这些结果确定了nSMase2调节的新途径,并表明组蛋白乙酰化的生理或药理调节可直接影响nSMase2水平。