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First, ATG12 is activated by the E1 enzyme ATG7 and transferred to the E2 enzyme ATG10 after a sequential reaction to bind ATG5 and recruit ATG16L1 to form the ATG12-ATG5-ATG16L1 complex (Nakatogawa, 2013), which acts as an ubiquitin-like E3 enzyme and is recruited to the phagophore by WIPI2 (Dooley et al., 2014