Bellacosa A, Datta K, Bear S E, Patriotis C, Lazo P A, Copeland N G, Jenkins N A, Tsichlis P N
Fox Chase Cancer Center, Philadelphia, Pennsylvania 19111.
J Virol. 1994 Apr;68(4):2320-30. doi: 10.1128/JVI.68.4.2320-2330.1994.
The Tpl-1 locus was defined as a genomic DNA region which is targeted by provirus insertion during progression of Moloney murine leukemia virus-induced rat T-cell lymphomas. Using a panel of 156 (Mus musculus x Mus spretus) x Mus musculus interspecific backcross mice, we mapped Tpl-1 to mouse chromosome 9 at a distance of 1.2 +/- 0.9 centimorgans from the Ets-1 proto-oncogene (S.E. Bear, A. Bellacosa, P.A. Lazo, N.A. Jenkins, N.G. Copeland, C. Hanson, G. Levan, and P.N. Tsichlis, Proc. Natl. Acad. Sci. USA 86:7495-7499, 1989). In this report, we present evidence that all the known Tpl-1 provirus insertions occurred immediately 5' of the first exon of Ets-1 (exon A) and that the earlier detected distance between Tpl-1 and Ets-1 was due to the high frequency of meiotic recombination in the region between the site of provirus integration and exon III. Northern (RNA) blot analysis of polyadenylated RNA from normal adult rat tissues and Moloney murine leukemia virus-induced T-cell lymphomas and hybridization to a Tpl-1/Ets-1 probe derived from the 5' end of the gene revealed two lymphoid cell-specific RNA transcripts, of 5.5 and 2.2 kb. Sequence analysis of a near-full-length (4,991-bp) cDNA clone of the 5.5-kb RNA revealed a 441-amino-acid open reading frame encoding a protein identical to the human and mouse Ets-1 proteins with the exception of five and nine species-specific conservative amino acid differences, respectively. The steady-state level of the Tpl-1/Ets-1 RNA and of the Ets-1 protein was modestly elevated in tumors carrying a provirus in the Tpl-1 locus. The relative ratio of the two Ets-1 transcripts, which were shown to arise by differential polyadenylation, was not affected by provirus insertion. Moreover, the major site of transcriptional initiation, which was localized by primer extension 250 bp upstream of the 5' end of the Ets-1 cDNA clone, was shown to be identical in normal cells and tumors carrying a provirus in the Tpl-1 locus. Finally, the differential splicing of Ets-1 exon VII was shown by RNase protection to occur at a rate of 15 to 26% and to remain unaffected by provirus insertion. The subtlety of these effects, in contrast to the strong growth selection of cells with a provirus in the Tpl-1/Ets-1 locus, suggests that provirus insertion may affect the fine regulation of the gene, perhaps during cell cycle progression.
Tpl-1基因座被定义为在莫洛尼鼠白血病病毒诱导的大鼠T细胞淋巴瘤进展过程中被前病毒插入靶向的基因组DNA区域。利用一组156只(小家鼠×西班牙小家鼠)×小家鼠种间回交小鼠,我们将Tpl-1定位到小鼠9号染色体上,距离Ets-1原癌基因1.2±0.9厘摩(S.E.贝尔、A.贝拉科萨、P.A.拉佐、N.A.詹金斯、N.G.科普兰、C.汉森、G.莱万和P.N.齐赫利斯,《美国国家科学院院刊》86:7495 - 7499,1989年)。在本报告中,我们提供证据表明,所有已知的Tpl-1前病毒插入都发生在Ets-1第一个外显子(外显子A)的5'端紧邻处,并且先前检测到的Tpl-1与Ets-1之间的距离是由于前病毒整合位点与外显子III之间区域减数分裂重组的高频率。对来自正常成年大鼠组织和莫洛尼鼠白血病病毒诱导的T细胞淋巴瘤的多聚腺苷酸化RNA进行Northern(RNA)印迹分析,并与源自该基因5'端的Tpl-1/Ets-1探针杂交,发现了两种淋巴样细胞特异性RNA转录本,大小分别为5.5和2.2 kb。对5.5-kb RNA的一个近全长(4991-bp)cDNA克隆进行序列分析,揭示了一个441个氨基酸的开放阅读框,编码一种与人类和小鼠Ets-1蛋白相同的蛋白质,只是分别有5个和9个物种特异性保守氨基酸差异。在Tpl-1基因座携带前病毒的肿瘤中,Tpl-1/Ets-1 RNA和Ets-1蛋白的稳态水平适度升高。两种Ets-1转录本通过差异多聚腺苷酸化产生,其相对比例不受前病毒插入的影响。此外,通过引物延伸在Ets-1 cDNA克隆5'端上游250 bp处定位的转录起始主要位点,在正常细胞和Tpl-1基因座携带前病毒的肿瘤中显示是相同的。最后,通过RNA酶保护显示Ets-1外显子VII的差异剪接发生率为15%至26%,并且不受前病毒插入的影响。与Tpl-1/Ets-1基因座携带前病毒的细胞的强烈生长选择相比,这些效应的微妙性表明前病毒插入可能影响该基因的精细调控,也许是在细胞周期进程中。