Li Hongbing, Sánchez-Torres Juan, Del Carpio Alan, Salas Valentina, Villalobo Antonio
Instituto de Investigaciones Biomédicas, Consejo Superior de Investigaciones Científicas and Universidad Autónoma de Madrid, Arturo Duperier 4, E-28029 Madrid, Spain.
Biochem J. 2004 Jul 1;381(Pt 1):257-66. doi: 10.1042/BJ20040515.
We have demonstrated previously that the EGFR (epidermal growth factor receptor) is a calmodulin (CaM)-binding protein. To establish whether or not the related receptor ErbB2/Neu/HER2 also binds CaM, we used human breast adenocarcinoma SK-BR-3 cells, because these cells overexpress this receptor thus facilitating the detection of this interaction. In the present paper, we show that ErbB2 could be pulled-down using CaM-agarose beads in a Ca2+-dependent manner, as detected by Western blot analysis using an anti-ErbB2 antibody. ErbB2 was also isolated by Ca2+-dependent CaM-affinity chromatography. We also demonstrate using an overlay technique with biotinylated CaM that CaM binds directly to the immunoprecipitated ErbB2. The binding of biotinylated CaM to ErbB2 depends strictly on the presence of Ca2+, since it was prevented by the presence of EGTA. Moreover, the addition of an excess of free CaM prevents the binding of its biotinylated form, demonstrating that this was a specific process. We excluded any interference with the EGFR, as SK-BR-3 cells express considerably lower levels of this receptor, and no detectable EGFR signal was observed by Western blot analysis in the immunoprecipitated ErbB2 preparations used to perform the overlay assays with biotinylated CaM. We also demonstrate that treating living cells with W7 [N-(6-aminohexyl)-5-chloro-1-naphthalenesulphonamide], a cell-permeant CaM antagonist, down-regulates ErbB2 phosphorylation, and show that W7 does not interfere non-specifically with the activity of ErbB tyrosine kinases. We also show that W7 inhibits the phosphorylation (activation) of both ERK1/2 (extracellular-signal-regulated kinases 1 and 2) and Akt/PKB (protein kinase B), in accordance with the inhibition observed in ErbB2 phosphorylation. In contrast, W7 treatment increased the phosphorylation (activation) of CREB (cAMP-response-element-binding protein) and ATF1 (activating transcription factor-1), two Ca2+-sensitive transcription factors that operate downstream of these ErbB2 signalling pathways, most likely because of the absence of calcineurin activity. We conclude that ErbB2 is a new CaM-binding protein, and that CaM plays a role in the regulation of this receptor and its downstream signalling pathways in vivo.
我们之前已经证明,表皮生长因子受体(EGFR)是一种钙调蛋白(CaM)结合蛋白。为了确定相关受体ErbB2/Neu/HER2是否也结合CaM,我们使用了人乳腺腺癌SK-BR-3细胞,因为这些细胞过度表达该受体,从而便于检测这种相互作用。在本文中,我们表明,使用抗ErbB2抗体进行蛋白质印迹分析检测到,ErbB2可以通过CaM琼脂糖珠以Ca2+依赖的方式被拉下。ErbB2也通过Ca2+依赖的CaM亲和色谱法分离出来。我们还使用生物素化CaM的覆盖技术证明,CaM直接结合到免疫沉淀的ErbB2上。生物素化CaM与ErbB2的结合严格依赖于Ca2+的存在,因为EGTA的存在会阻止这种结合。此外,添加过量的游离CaM会阻止其生物素化形式的结合,表明这是一个特异性过程。我们排除了对EGFR的任何干扰,因为SK-BR-3细胞中该受体的表达水平相当低,在用生物素化CaM进行覆盖分析的免疫沉淀ErbB2制剂中,通过蛋白质印迹分析未观察到可检测到的EGFR信号。我们还证明,用细胞可渗透的CaM拮抗剂W7 [N-(6-氨基己基)-5-氯-1-萘磺酰胺]处理活细胞会下调ErbB2的磷酸化,并表明W7不会非特异性地干扰ErbB酪氨酸激酶的活性。我们还表明,W7抑制细胞外信号调节激酶1和2(ERK1/2)以及蛋白激酶B(Akt/PKB)的磷酸化(激活),这与在ErbB2磷酸化中观察到的抑制一致。相反,W7处理增加了环磷酸腺苷反应元件结合蛋白(CREB)和激活转录因子-1(ATF1)的磷酸化(激活),这两种Ca2+敏感转录因子在这些ErbB2信号通路的下游发挥作用,最有可能是因为缺乏钙调神经磷酸酶活性。我们得出结论,ErbB2是一种新的CaM结合蛋白,并且CaM在体内对该受体及其下游信号通路的调节中发挥作用。