Damke H, Baba T, Warnock D E, Schmid S L
Department of Cell Biology, Scripps Research Institute, La Jolla, California 92037.
J Cell Biol. 1994 Nov;127(4):915-34. doi: 10.1083/jcb.127.4.915.
Dynamin is the mammalian homologue to the Drosophila shibire gene product. Mutations in this 100-kD GTPase cause a pleiotropic defect in endocytosis. To further investigate its role, we generated stable HeLa cell lines expressing either wild-type dynamin or a mutant defective in GTP binding and hydrolysis driven by a tightly controlled, tetracycline-inducible promoter. Overexpression of wild-type dynamin had no effect. In contrast, coated pits failed to become constricted and coated vesicles failed to bud in cells overexpressing mutant dynamin so that endocytosis via both transferrin (Tfn) and EGF receptors was potently inhibited. Coated pit assembly, invagination, and the recruitment of receptors into coated pits were unaffected. Other vesicular transport pathways, including Tfn receptor recycling, Tfn receptor biosynthesis, and cathepsin D transport to lysosomes via Golgi-derived coated vesicles, were unaffected. Bulk fluid-phase uptake also continued at the same initial rates as wild type. EM immunolocalization showed that membrane-bound dynamin was specifically associated with clathrin-coated pits on the plasma membrane. Dynamin was also associated with isolated coated vesicles, suggesting that it plays a role in vesicle budding. Like the Drosophila shibire mutant, HeLa cells overexpressing mutant dynamin accumulated long tubules, many of which remained connected to the plasma membrane. We conclude that dynamin is specifically required for endocytic coated vesicle formation, and that its GTP binding and hydrolysis activities are required to form constricted coated pits and, subsequently, for coated vesicle budding.
发动蛋白是果蝇“麻木”基因产物在哺乳动物中的同源物。这种100-kD GTP酶的突变会导致内吞作用出现多效性缺陷。为了进一步研究其作用,我们构建了稳定的HeLa细胞系,这些细胞系表达野生型发动蛋白或由严格控制的四环素诱导型启动子驱动的GTP结合与水解缺陷型突变体。野生型发动蛋白的过表达没有影响。相比之下,在过表达突变体发动蛋白的细胞中,有被小窝无法收缩,有被小泡无法出芽,从而使通过转铁蛋白(Tfn)和表皮生长因子受体的内吞作用受到强烈抑制。有被小窝的组装、内陷以及受体募集到有被小窝的过程未受影响。其他囊泡运输途径,包括Tfn受体循环、Tfn受体生物合成以及组织蛋白酶D通过高尔基体衍生的有被小泡运输到溶酶体,均未受影响。大量液相摄取也以与野生型相同的初始速率持续进行。电子显微镜免疫定位显示,膜结合的发动蛋白与质膜上的网格蛋白包被小窝特异性相关。发动蛋白也与分离的有被小泡相关,这表明它在小泡出芽中起作用。与果蝇“麻木”突变体一样,过表达突变体发动蛋白的HeLa细胞积累了长管状结构,其中许多仍与质膜相连。我们得出结论,发动蛋白是内吞性有被小泡形成所特需的,并且其GTP结合和水解活性是形成收缩的有被小窝以及随后有被小泡出芽所必需的。