Table 1 Published list of ULK1 substrates
ProteinResidue(s)FunctionReferences
ULK1T180, S1042, T1046T180 important for ULK1 kinase activity. S1042 and T1046 regulate ULK1 ubiquitylation and degradation during prolonged starvation[21,35]
ATG13S318 (S355 isoform 1)S318 required for clearance of depolarized mitochondria[74]
FIP200S943, S986, S1323Unknown[10]
ATG101S11, S203Unknown[10]
ATG9S14Promotes ATG9 trafficking in response to starvation[50]
BECLIN-1S14 (mouse)Required for VPS34 activation[44]
AMBRA1S465, S635Regulates dissociation of AMPRA1-VPS34-BECLIN-1 from the dynein complex, to promote interaction with ULK1 and autophagy[10,46]
ATG4BS316Inhibits ATG4B activity and LC3 processing[51]
FUNDC1S17Promotes mitophagy by enhancing FUNDC1 binding to LC3[54]
VPS34S249Unknown[10]
ATG14LS29Promotes autophagy by increasing VPS34 complex activity[45]
DENND3S554, S572Activates Rab12 to facilitate autophagosome trafficking[75]
HK1S124During amino acid and growth factor starvation in order to maintain homoeostasis of cellular energy and redox levels[76]
ENO1S282During amino acid and growth factor starvation in order to maintain homoeostasis of cellular energy and redox levels[76]
PFK1S762During amino acid and growth factor starvation in order to maintain homoeostasis of cellular energy and redox levels[76]
FBP1S63During amino acid and growth factor starvation in order to maintain homoeostasis of cellular energy and redox levels[76]
RaptorS855, S859, S792Inhibition of mTORC1 during starvation[77]
Sec23AS207, S312Inhibition of ER to Golgi trafficking during starvation[78]
Cdc37S339Disruption of Cdc37 client proteins stability[79]
Smrc8S400, S492, S562, T666Unknown function in C9orf72 regulation of the ULK1 complex[42]
p62/SQSTM1S409To promote aggregate clearance during proteotoxic stress[43]
StingS366To inhibit an excessive transcription of innate immune genes during their activation by cyclic dinucleotides[80]
  • Abbreviations: DENND3, DENN domain containing 3; HK1, hexokinase 1; ENO1, enolase 1; PFK1, phosphofructokinase 1; FBP1, fructose-1,6-bisphosphatase 1; FUNDC1, FUN14 domain containing 1; p62/SQSTM1, sequestosome-1.