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The E2 either transfers ubiquitin directly to the E3 in the case of HECT E3s or binds the E3 and transfers the ubiquitin to the substrate. 泛素结合酶结合泛素蛋白连接酶并把泛素转移到底物蛋白上使底物泛素化，或把泛素转移到泛素蛋白连接酶再使底物泛素化。