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. 2004 Aug 13;36(12):2503–2518. doi: 10.1016/j.biocel.2004.05.009

Table 1.

Mammalian autophagy-related genes

Gene Designation
Yeast Human Mouse Comment Reference

ATG1 ULK1 Unc-51-like kinase interacts with GATE-16 and GABARAP Larsen and Sulzer, 2002, Lemasters et al., 2002
ATG3 hATG3/hAPG3 mATG3/mAPG3 an E2-like enzyme for LC3, GABARAP, and GATE-16 Liang et al., 1999, Lockshin and Zakeri, 2002, Mann and Hammarback, 1994
ATG4 hATG4A/HsATG4A/HsAPG4A/autophagin-2 Cysteine protease for GATE-16 Liang et al., 1999, Marino et al., 2003, Matsuura et al., 1997, Meier and Silke, 2003
hATG4B/HsATG4B/hAPG4B/autophagin-1 Cysteine protease for LC3, Matsuura et al., 1997, Meijer and Dubbelhuis, 2004
GABARAP, and GATE-16 unpublished results
Delipidating enzyme for LC3-II and GABARAP unpublished results
hAtg4C/HsAUTL1/autophagin-3 Cysteine protease Matsuura et al., 1997
hATG4D/autophagin-4 Matsuura et al., 1997
ATG5 hATG5/hAPG5 target protein of Atg12 Mizushima et al., 1998, Mizushima et al., 1998, Mizushima et al., 2001, Mizushima et al., 2003, Mizushima et al., 2003
ATG6 beclin 1 beclin 1 related to tumorigenesis Mizushima et al., 1999, Mizushima et al., 1998, Mizushima et al., 1998
ATG7 hATG7/HsGSA7/hAPG7 mATG7/mAPG7 an E1-like enzyme for Atg12 and Atg 8 homologues Mizushima et al., 2001, Mizushima et al., 2004, Mizushima et al., 2002, Mizushima et al., 2003, Mizushima et al., 2003, Nakanishi, 2003, Nemoto et al., 2003
ATG8 LC3 modifier for autophagosomes Marino et al., 2003, Nishino et al., 2000, Nishino, 2003
GABARAP modifier Noguchi et al., 2004
GATE-16 modifier Ogier-Denis and Codogno, 2003
ATG8L/APG8L unknown Meijer and Dubbelhuis, 2004
ATG10 mATG10/mAPG10 an E2-like enzyme for Atg12 Ohsumi, 2001, Okazaki et al., 2000
ATG12 hATG12/hAPG12 mATG12/mAPG12 modifier for autophagosomal Mizushima et al., 2001, Mizushima et al., 2003, Mizushima et al., 2003, Nemoto et al., 2003
ATG16 ATG16L/APG16L interacts with Atg5 Otto et al., 2004, Otto et al., 2003