Table 1.

Post‐translational modifications of and by the UNC51‐like kinase 1 (ULK1) complex in autophagy

Substrate	Modification	Site	Modifying enzyme	Effect in autophagy	References
ULK1	P	Ser317	AMPK	Direct activating ULK1 in a mTOR‐independent manner	81
		Thr575
		Ser555
		Ser777
	P	Ser757	mTOR	Prevent ULK1 action & disrupting the interaction between ULK1 and AMPK	37, 81
		Ser638
		Ser758
	P	Ser774	Akt		37
	P	Thr180	ULK1	Autophosphorylation to form the kinase‐activation loop	37
	P	C‐terminal regions residues 829–1051	mAtg13	Binding with ULK1	41, 54
	P	C‐terminal regions residues 651–1051	FIP200
	Ac	Lys162	Tip60	Active ULK1 in serum starvation
		Lys606			53
	Ub	Unknown	TRAF6	AMBRA1, interacting with the E3‐ligase TRAF6, supports ULK1 ubiquitylation by LYS‐63‐linked chains	53, 82
AMPK	P	Thr172	ULK1	Decreased the starvation induced activation of AMPK (Feedback mechanisms)	37, 44
AMBRA1	P	Ser52	mTOR	Inhibit AMBRA1 by phosphorylation, whereas on autophagy induction	52
AMBRA1	P	Unknown	ULK1	The cross‐talk between ULK1 and Beclin‐1 complexes	52
FIP200	P	Unknown	ULK1	Binding with ULK1	41, 54
Raptor	P	Ser855		Inactivating mTORC1	83
		Ser859	ULK1
		Ser863
		Ser792
mTOR	P	Ser2481	ULK1	Induced by ULK1	83
mAtg13	P	C‐terminal residues 384–517	ULK1	Binding with ULK1	41, 54
Sqa (ZIPK homology)	P	Thr194	ULK1	Phosphorylation of Sqa at Thr‐279 is required for Atg1‐mediated myosin II activation	59
		Thr239	ULK1
		Thr279	ULK1
ZIPK	P	Unknown	ULK1	May process the same function of Atg1 acting with Sqa.	59
Beclin‐1	P	Ser14	ULK1	Enhance the activity of the Atg14L‐containing VPS34 complexes	58