Kowluru A, Seavey S E, Li G, Sorenson R L, Weinhaus A J, Nesher R, Rabaglia M E, Vadakekalam J, Metz S A
Department of Medicine and Division of Endocrinology, University of Wisconsin School of Medicine, Madison, Wisconsin 53792, USA.
J Clin Invest. 1996 Jul 15;98(2):540-55. doi: 10.1172/JCI118822.
Several GTP-binding proteins (G-proteins) undergo post-translational modifications (isoprenylation and carboxyl methylation) in pancreatic beta cells. Herein, two of these were identified as CDC42 and rap 1, using Western blotting and immunoprecipitation. Confocal microscopic data indicated that CDC42 is localized only in islet endocrine cells but not in acinar cells of the pancreas. CDC42 undergoes a guanine nucleotide-specific membrane association and carboxyl methylation in normal rat islets, human islets, and pure beta (HIT or INS-1) cells. GTPgammaS-dependent carboxyl methylation of a 23-kD protein was also demonstrable in secretory granule fractions from normal islets or beta cells. AFC (a specific inhibitor of prenyl-cysteine carboxyl methyl transferases) blocked the carboxyl methylation of CDC42 in five types of insulin-secreting cells, without blocking GTPgammaS-induced translocation, implying that methylation is a consequence (not a cause) of transfer to membrane sites. High glucose (but not a depolarizing concentration of K+) induced the carboxyl methylation of CDC42 in intact cells, as assessed after specific immunoprecipitation. This effect was abrogated by GTP depletion using mycophenolic acid and was restored upon GTP repletion by coprovision of guanosine. In contrast, although rap 1 was also carboxyl methylated, it was not translocated to the particulate fraction by GTPgammaS; furthermore, its methylation was also stimulated by 40 mM K+ (suggesting a role which is not specific to nutrient stimulation). AFC also impeded nutrient-induced (but not K+-induced) insulin secretion from islets and beta cells under static or perifusion conditions, whereas an inactive structural analogue of AFC failed to inhibit insulin release. These effects were reproduced not only by S-adenosylhomocysteine (another methylation inhibitor), but also by GTP depletion. Thus, the glucose- and GTP-dependent carboxyl methylation of G-proteins such as CDC42 is an obligate step in the stimulus-secretion coupling of nutrient-induced insulin secretion, but not in the exocytotic event itself. Furthermore, AFC blocked glucose-activated phosphoinositide turnover, which may provide a partial biochemical explanation for its effect on secretion, and implies that certain G-proteins must be carboxyl methylated for their interaction with signaling effector molecules, a step which can be regulated by intracellular availability of GTP.
几种GTP结合蛋白(G蛋白)在胰腺β细胞中会经历翻译后修饰(异戊二烯化和羧基甲基化)。在此,通过蛋白质印迹法和免疫沉淀法确定其中两种为CDC42和rap 1。共聚焦显微镜数据表明,CDC42仅定位于胰岛内分泌细胞,而不在胰腺腺泡细胞中。在正常大鼠胰岛、人胰岛和纯β(HIT或INS-1)细胞中,CDC42会发生鸟嘌呤核苷酸特异性膜结合和羧基甲基化。在来自正常胰岛或β细胞的分泌颗粒组分中也可证实23-kD蛋白的GTPγS依赖性羧基甲基化。AFC(异戊烯基半胱氨酸羧基甲基转移酶的特异性抑制剂)可阻断五种胰岛素分泌细胞中CDC42的羧基甲基化,但不阻断GTPγS诱导的易位,这意味着甲基化是转移至膜位点的结果(而非原因)。如特异性免疫沉淀后所评估,高糖(而非去极化浓度的K+)可诱导完整细胞中CDC42的羧基甲基化。使用霉酚酸消耗GTP可消除此效应,通过同时提供鸟苷补充GTP后效应得以恢复。相比之下,尽管rap 1也会发生羧基甲基化,但它不会被GTPγS转运至颗粒组分;此外,其甲基化也受到40 mM K+的刺激(表明其作用并非特定于营养刺激)。在静态或灌流条件下,AFC也会阻碍胰岛和β细胞中营养物质诱导的(而非K+诱导的)胰岛素分泌,而AFC的无活性结构类似物无法抑制胰岛素释放。这些效应不仅可被S-腺苷同型半胱氨酸(另一种甲基化抑制剂)重现,也可被GTP消耗重现。因此,诸如CDC42等G蛋白的葡萄糖和GTP依赖性羧基甲基化是营养物质诱导的胰岛素分泌刺激-分泌偶联中的一个必要步骤,但不是胞吐事件本身中的必要步骤。此外,AFC可阻断葡萄糖激活的磷酸肌醇周转,这可能为其对分泌的影响提供部分生化解释,并意味着某些G蛋白必须进行羧基甲基化才能与信号效应分子相互作用,这一步骤可由细胞内GTP的可用性调节。