Kessels Michael M, Qualmann Britta
Research Group Membrane Trafficking and Cytoskeleton, Department of Neurochemistry and Molecular Biology, D-39118 Magdeburg, Germany.
Research Group Cell Biology, Leibniz Institute for Neurobiology, D-39118 Magdeburg, Germany.
J Biol Chem. 2006 May 12;281(19):13285-13299. doi: 10.1074/jbc.M510226200. Epub 2006 Mar 15.
Syndapins were proposed to interconnect the machineries for vesicle formation and actin polymerization, as they interact with dynamin and the Arp2/3 complex activator N-WASP (neural Wiskott-Aldrich syndrome protein). Syndapins, however, have only one Src homology 3 domain mediating both interactions. Here we show that syndapins self-associate via direct syndapin/syndapin interactions, providing a molecular mechanism for the coordinating role of syndapin. Cross-link studies with overexpressed and endogenous syndapins suggest that predominantly dimers form in vivo. Our analyses show that the N-terminal Fes/Cip4 homology domain but not the central coiled-coil domain is sufficient for oligomerization. Additionally, a second interface located further C-terminally mediated interactions with the N terminus. The Src homology 3 domain and the NPF region are not involved and thus available for further interactions interconnecting different syndapin binding partners. Our analyses showed that self-association is crucial for syndapin function. Both syndapin-mediated cytoskeletal rearrangements and endocytosis were disrupted by a self-association-deficient mutant. Consistent with a role of syndapins in linking actin polymerization bursts with endocytic vesicle formation, syndapin-containing complexes had a size of 300-500 kDa in gel filtration analysis and contained both dynamin and N-WASP. The existence of an interconnection of the GTPase dynamin with N-WASP via syndapin oligomers was demonstrated both by coimmunoprecipitations and by reconstitution at membranes in intact cells. The interconnection was disrupted by coexpression of syndapin mutants incapable of self-association. Syndapin oligomers may thus act as multivalent organizers spatially and temporally coordinating vesicle fission with local actin polymerization.
Syndapins被认为可将囊泡形成机制与肌动蛋白聚合作用相互连接起来,因为它们能与发动蛋白以及Arp2/3复合物激活剂N-WASP(神经威斯科特-奥尔德里奇综合征蛋白)相互作用。然而,Syndapins只有一个Src同源3结构域介导这两种相互作用。在此我们表明,Syndapins通过直接的Syndapin/Syndapin相互作用进行自我缔合,这为Syndapin的协调作用提供了一种分子机制。对过表达和内源性Syndapins进行的交联研究表明,在体内主要形成二聚体。我们的分析表明,N端的Fes/Cip4同源结构域而非中央的卷曲螺旋结构域就足以进行寡聚化。此外,位于更靠C端的第二个界面介导了与N端的相互作用。Src同源3结构域和NPF区域不参与其中,因此可用于与不同的Syndapin结合伴侣进行进一步的相互作用。我们的分析表明,自我缔合对Syndapin功能至关重要。自我缔合缺陷型突变体破坏了Syndapin介导的细胞骨架重排和内吞作用。与Syndapins在将肌动蛋白聚合爆发与内吞囊泡形成相联系中所起的作用一致,在凝胶过滤分析中,含有Syndapins的复合物大小为300 - 500 kDa,并且同时含有发动蛋白和N-WASP。通过共免疫沉淀以及在完整细胞的膜上进行重组,均证明了GTP酶发动蛋白通过Syndapin寡聚体与N-WASP之间存在相互连接。这种相互连接被不能进行自我缔合的Syndapin突变体的共表达所破坏。因此,Syndapin寡聚体可能作为多价组织者在空间和时间上协调囊泡裂变与局部肌动蛋白聚合。