College of Life Science, Beijing Normal University, Beijing 100875, China.
National Institute of Biological Sciences, Beijing 102206, China.
Mol Biol Cell. 2018 Aug 15;29(17):2098-2112. doi: 10.1091/mbc.E17-11-0639. Epub 2018 Jun 21.
After endocytosis, transmembrane cargoes such as signaling receptors, channels, and transporters enter endosomes where they are sorted to different destinations. Retromer and ESCRT (endosomal sorting complex required for transport) are functionally distinct protein complexes on endosomes that direct cargo sorting into the recycling retrograde transport pathway and the degradative multivesicular endosome pathway (MVE), respectively. Cargoes destined for degradation in lysosomes are decorated with K63-linked ubiquitin chains, which serve as an efficient sorting signal for entry into the MVE pathway. Defects in K63-linked ubiquitination disrupt MVE sorting and degradation of membrane proteins. Here, we unexpectedly found that UBC-13, the E2 ubiquitin-conjugating enzyme that generates K63-linked ubiquitin chains, is essential for retrograde transport of multiple retromer-dependent cargoes including MIG-14/Wntless. Loss of ubc-13 disrupts MIG-14/Wntless trafficking from endosomes to the Golgi, causing missorting of MIG-14 to lysosomes and impairment of Wnt-dependent processes. We observed that retromer-associated SNX-1 and the ESCRT-0 subunit HGRS-1/Hrs localized to distinct regions on a common endosome in wild type but overlapped on ubc-13(lf) endosomes, indicating that UBC-13 is important for the separation of retromer and ESCRT microdomains on endosomes. Our data suggest that cargo ubiquitination mediated by UBC-13 plays an important role in maintaining the functionally distinct subdomains to ensure efficient cargo segregation on endosomes.
内吞作用后,跨膜货物(如信号受体、通道和转运体)进入内体,在内体中被分拣到不同的目的地。Retromer 和 ESCRT(内体分选复合物必需运输)是内体上功能不同的蛋白质复合物,分别将货物分拣到再循环逆行运输途径和降解多泡体内体途径(MVE)。在溶酶体中降解的货物被 K63 连接的泛素链修饰,K63 连接的泛素链作为进入 MVE 途径的有效分拣信号。K63 连接的泛素化缺陷破坏 MVE 分拣和膜蛋白的降解。在这里,我们出人意料地发现,UBC-13,即产生 K63 连接的泛素链的 E2 泛素连接酶,是多个依赖 retromer 的货物(包括 MIG-14/Wntless)逆行运输所必需的。ubc-13 的缺失破坏了 MIG-14/Wntless 从内体到高尔基体的运输,导致 MIG-14 错误分拣到溶酶体,并损害了 Wnt 依赖的过程。我们观察到,retromer 相关的 SNX-1 和 ESCRT-0 亚基 HGRS-1/Hrs 在野生型中定位于同一个内体上的不同区域,但在 ubc-13(lf)内体上重叠,表明 UBC-13 对内体上 retromer 和 ESCRT 微区室的分离很重要。我们的数据表明,UBC-13 介导的货物泛素化在维持功能不同的亚区室方面发挥着重要作用,以确保货物在内体上的有效分离。