Department of Cell Biology and Physiology, Washington University School of Medicine, St Louis, Missouri 63110, USA.
J Biol Chem. 2010 Nov 12;285(46):35428-38. doi: 10.1074/jbc.M110.126318. Epub 2010 Aug 30.
VPS4 proteins are AAA(+) ATPases required to form multivesicular bodies, release viral particles, and complete cytokinesis. They act by disassembling ESCRT-III heteropolymers during or after their proposed function in membrane scission. Here we show that purified human VPS4A is essentially inactive but can be stimulated to hydrolyze ATP by ESCRT-III proteins in a reaction that requires both their previously defined MIT interacting motifs and ∼50 amino acids of the adjacent sequence. Importantly, C-terminal fragments of all ESCRT-III proteins tested, including CHMP2A, CHMP1B, CHMP3, CHMP4A, CHMP6, and CHMP5, activated VPS4A suggesting that it disassembles ESCRT-III heteropolymers by affecting each component protein. VPS4A is thought to act as a ring-shaped cylindrical oligomer like other AAA(+) ATPases, but this has been difficult to directly demonstrate. We found that concentrating His(6)-VPS4A on liposomes containing Ni(2+)-nitrilotriacetic acid-tagged lipid increased ATP hydrolysis, confirming the importance of inter-subunit interactions for activity. We also found that mutating pore loops expected to line the center of a cylindrical oligomer changed the response of VPS4A to ESCRT-III proteins. Based on these data, we propose that ESCRT-III proteins facilitate assembly of functional but transient VPS4A oligomers and interact with sequences inside the pore of the assembled enzyme. Deleting the N-terminal MIT domain and adjacent linker from VPS4A increased both basal and liposome-enhanced ATPase activity, indicating that these elements play a role in autoinhibiting VPS4A until it encounters ESCRT-III proteins. These findings reveal new ways in which VPS4 activity is regulated and specifically directed to ESCRT-III polymers.
VPS4 蛋白是 AAA(+)ATP 酶,需要形成多泡体、释放病毒颗粒并完成胞质分裂。它们通过在膜分裂过程中或之后分解 ESCRT-III 杂多体来发挥作用。在这里,我们表明纯化的人 VPS4A 基本上没有活性,但可以通过 ESCRT-III 蛋白刺激水解 ATP,该反应需要它们先前定义的 MIT 相互作用基序和相邻序列中的约 50 个氨基酸。重要的是,测试的所有 ESCRT-III 蛋白的 C 末端片段,包括 CHMP2A、CHMP1B、CHMP3、CHMP4A、CHMP6 和 CHMP5,都激活了 VPS4A,表明它通过影响每个组成蛋白来分解 ESCRT-III 杂多体。VPS4A 被认为像其他 AAA(+)ATP 酶一样作为环形圆柱形寡聚体发挥作用,但这很难直接证明。我们发现,将 His(6)-VPS4A 浓缩在含有 Ni(2+)- 亚氨基二乙酸标记脂质的脂质体上会增加 ATP 水解,从而证实了亚基间相互作用对活性的重要性。我们还发现,突变预期排列在圆柱形寡聚体中心的孔环改变了 VPS4A 对 ESCRT-III 蛋白的反应。基于这些数据,我们提出 ESCRT-III 蛋白促进功能性但瞬时 VPS4A 寡聚体的组装,并与组装酶孔内的序列相互作用。从 VPS4A 中删除 N 端 MIT 结构域和相邻连接子会增加基础和脂质体增强的 ATP 酶活性,这表明这些元件在 VPS4A 遇到 ESCRT-III 蛋白之前发挥作用,自动抑制 VPS4A。这些发现揭示了 VPS4 活性被调节并特别靶向 ESCRT-III 聚合物的新方式。