Department of Pathology and Immunology, Washington University School of Medicine, St Louis, Missouri 63110, USA.
J Biol Chem. 2010 May 21;285(21):16248-57. doi: 10.1074/jbc.M109.048785. Epub 2010 Mar 16.
NUP98 is a nucleoporin that plays complex roles in the nucleocytoplasmic trafficking of macromolecules. Rearrangements of the NUP98 gene in human leukemia result in the expression of numerous fusion oncoproteins whose effect on nucleocytoplasmic trafficking is poorly understood. The present study was undertaken to determine the effects of leukemogenic NUP98 fusion proteins on CRM1-mediated nuclear export. NUP98-HOXA9, a prototypic NUP98 fusion, inhibited the nuclear export of two known CRM1 substrates: mutated cytoplasmic nucleophosmin and HIV-1 Rev. In vitro binding assays revealed that NUP98-HOXA9 binds CRM1 through the FG repeat motif in a Ran-GTP-dependent manner similar to but stronger than the interaction between CRM1 and its export substrates. Two NUP98 fusions, NUP98-HOXA9 and NUP98-DDX10, whose fusion partners are structurally and functionally unrelated, interacted with endogenous CRM1 in myeloid cells as shown by co-immunoprecipitation. These leukemogenic NUP98 fusion proteins interacted with CRM1, Ran, and the nucleoporin NUP214 in a manner fundamentally different from that of wild-type NUP98. NUP98-HOXA9 and NUP98-DDX10 formed characteristic aggregates within the nuclei of a myeloid cell line and primary human CD34+ cells and caused aberrant localization of CRM1 to these aggregates. These NUP98 fusions caused nuclear accumulation of two transcription factors, NFAT and NFkappaB, that are regulated by CRM1-mediated export. The nuclear entrapment of NFAT and NFkappaB correlated with enhanced transcription from promoters responsive to these transcription factors. Taken together, the results suggest a new mechanism by which NUP98 fusions dysregulate transcription and cause leukemia, namely, inhibition of CRM1-mediated nuclear export with aberrant nuclear retention of transcriptional regulators.
NUP98 是一种核孔蛋白,在核质大分子的运输中发挥着复杂的作用。人类白血病中 NUP98 基因的重排导致表达许多融合癌蛋白,但其对核质运输的影响知之甚少。本研究旨在确定致白血病 NUP98 融合蛋白对 CRM1 介导的核输出的影响。NUP98-HOXA9 是一种典型的 NUP98 融合蛋白,它抑制了两种已知的 CRM1 底物的核输出:突变的细胞质核磷蛋白和 HIV-1 Rev。体外结合实验表明,NUP98-HOXA9 通过 FG 重复基序与 CRM1 结合,这种结合方式类似于 CRM1 与其出口底物之间的结合,但比这种结合更强,需要 Ran-GTP 的参与。两种 NUP98 融合蛋白,NUP98-HOXA9 和 NUP98-DDX10,其融合伙伴在结构和功能上不相关,通过共免疫沉淀在髓系细胞中与内源性 CRM1 相互作用。这些致白血病的 NUP98 融合蛋白与 CRM1、Ran 和核孔蛋白 NUP214 相互作用的方式与野生型 NUP98 有根本的不同。NUP98-HOXA9 和 NUP98-DDX10 在髓系细胞系和原代人 CD34+细胞的核内形成特征性的聚集体,并导致 CRM1 异常定位到这些聚集体。这些 NUP98 融合蛋白导致两种转录因子 NFAT 和 NFkappaB 的核内积累,NFAT 和 NFkappaB 受 CRM1 介导的出口调控。NFAT 和 NFkappaB 的核内捕获与这些转录因子的启动子的转录增强相关。总之,这些结果表明了 NUP98 融合蛋白通过异常的核内保留转录调节剂来调节转录和导致白血病的一种新机制,即抑制 CRM1 介导的核输出。