Gene Symbol: ATG9
Description: autophagy protein ATG9
Alias: APG9, AUT9, CVT6, CVT7, autophagy protein ATG9
Species: Saccharomyces cerevisiae S288c

Top Publications

  1. Reggiori F, Tucker K, Stromhaug P, Klionsky D. The Atg1-Atg13 complex regulates Atg9 and Atg23 retrieval transport from the pre-autophagosomal structure. Dev Cell. 2004;6:79-90 pubmed
    ..We analyzed the trafficking of Atg23 and the integral membrane protein Atg9 in the yeast Saccharomyces cerevisiae...
  2. Reggiori F, Shintani T, Nair U, Klionsky D. Atg9 cycles between mitochondria and the pre-autophagosomal structure in yeasts. Autophagy. 2005;1:101-9 pubmed
    ..In Saccharomyces cerevisiae, the integral membrane protein Atg9 that is required for autophagy cycles through the PAS...
  3. Sekito T, Kawamata T, Ichikawa R, Suzuki K, Ohsumi Y. Atg17 recruits Atg9 to organize the pre-autophagosomal structure. Genes Cells. 2009;14:525-38 pubmed publisher
    ..Under nutrient-rich condition, Atg9 is recruited to the PAS by binding to Atg11 for the Cvt pathway...
  4. Monastyrska I, He C, Geng J, Hoppe A, Li Z, Klionsky D. Arp2 links autophagic machinery with the actin cytoskeleton. Mol Biol Cell. 2008;19:1962-75 pubmed publisher
    ..The integral membrane protein Atg9 cycles between certain intracellular compartments and the vesicle nucleation site, presumably to supply membranes ..
  5. He C, Baba M, Cao Y, Klionsky D. Self-interaction is critical for Atg9 transport and function at the phagophore assembly site during autophagy. Mol Biol Cell. 2008;19:5506-16 pubmed publisher
    ..b>Atg9 is the sole integral membrane protein required in formation of the initial sequestering compartment, the phagophore,..
  6. Tucker K, Reggiori F, Dunn W, Klionsky D. Atg23 is essential for the cytoplasm to vacuole targeting pathway and efficient autophagy but not pexophagy. J Biol Chem. 2003;278:48445-52 pubmed
    ..We also demonstrate that Atg23 is a peripheral membrane protein that requires the presence of Atg9/Apg9 to be specifically targeted to lipid bilayers...
  7. Yen W, Legakis J, Nair U, Klionsky D. Atg27 is required for autophagy-dependent cycling of Atg9. Mol Biol Cell. 2007;18:581-93 pubmed
    ..The transmembrane protein Atg9 cycles between the mitochondria and the pre-autophagosomal structure, which is the site of autophagosome biogenesis...
  8. Reggiori F, Klionsky D. Atg9 sorting from mitochondria is impaired in early secretion and VFT-complex mutants in Saccharomyces cerevisiae. J Cell Sci. 2006;119:2903-11 pubmed
    ..The integral membrane protein Atg9 is essential for both autophagy and the cytoplasm-to-vacuole targeting pathway, a selective biosynthetic process in ..
  9. Yen W, Shintani T, Nair U, Cao Y, Richardson B, Li Z, et al. The conserved oligomeric Golgi complex is involved in double-membrane vesicle formation during autophagy. J Cell Biol. 2010;188:101-14 pubmed publisher
    ..In addition, mutations in the COG genes resulted in the mislocalization of Atg8 and Atg9, which are critical components involved in autophagosome formation.

More Information


  1. Ohashi Y, Munro S. Membrane delivery to the yeast autophagosome from the Golgi-endosomal system. Mol Biol Cell. 2010;21:3998-4008 pubmed publisher
    ..The integral membrane protein Atg9 is delivered to the autophagosome during starvation and in the related cytoplasm-to-vacuole (Cvt) pathway that ..
  2. He C, Song H, Yorimitsu T, Monastyrska I, Yen W, Legakis J, et al. Recruitment of Atg9 to the preautophagosomal structure by Atg11 is essential for selective autophagy in budding yeast. J Cell Biol. 2006;175:925-35 pubmed
    ..In the yeast Saccharomyces cerevisiae, the integral membrane protein Atg9 (autophagy-related protein 9) cycles between mitochondria and the preautophagosomal structure (PAS), the nucleating ..
  3. Legakis J, Yen W, Klionsky D. A cycling protein complex required for selective autophagy. Autophagy. 2007;3:422-32 pubmed
    ..Here, we examine three autophagy-related (Atg) proteins, Atg9, Atg23 and Atg27, which exhibit a unique localization pattern, residing both at the pre-autophagosomal structure (..
  4. Wang C, Kim J, Huang W, Abeliovich H, Stromhaug P, Dunn W, et al. Apg2 is a novel protein required for the cytoplasm to vacuole targeting, autophagy, and pexophagy pathways. J Biol Chem. 2001;276:30442-51 pubmed
    ..Apg2 localizes to the previously identified perivacuolar compartment that contains Apg9, the only characterized integral membrane protein that is required for autophagosome/Cvt vesicle formation.
  5. Lebesgue N, Megyeri M, Cristobal A, Scholten A, Chuartzman S, Voichek Y, et al. Combining Deep Sequencing, Proteomics, Phosphoproteomics, and Functional Screens To Discover Novel Regulators of Sphingolipid Homeostasis. J Proteome Res. 2017;16:571-582 pubmed publisher
    ..By following the rate of SL biosynthesis in those candidates that are both affecting growth and are phosphorylated in response to the drug, we uncovered Atg9, Stp4, and Gvp36 as putative new regulators of SL homeostasis.
  6. He C, Baba M, Klionsky D. Double duty of Atg9 self-association in autophagosome biogenesis. Autophagy. 2009;5:385-7 pubmed
    ..b>Atg9 is the only identified integral membrane protein required for autophagosome formation, and it is thought to cycle ..
  7. Lipatova Z, Segev N. A Role for Macro-ER-Phagy in ER Quality Control. PLoS Genet. 2015;11:e1005390 pubmed publisher
    ..Moreover, for the first time the role of Atg9, the only integral-membrane core Atg, is uncoupled from that of other core Atgs...
  8. Shirahama Noda K, Kira S, Yoshimori T, Noda T. TRAPPIII is responsible for vesicular transport from early endosomes to Golgi, facilitating Atg9 cycling in autophagy. J Cell Sci. 2013;126:4963-73 pubmed publisher
    ..Cargo proteins in this TRAPPIII-dependent pathway include Atg9, a transmembrane protein that is essential for autophagy, and Snc1, a SNARE unrelated to autophagy...
  9. Ma J, Bharucha N, Dobry C, Frisch R, Lawson S, Kumar A. Localization of autophagy-related proteins in yeast using a versatile plasmid-based resource of fluorescent protein fusions. Autophagy. 2008;4:792-800 pubmed
    ..Collectively, this plasmid-based resource of yeast gene-vYFP fusions provides an initial toolkit for a variety of systematic and large-scale localization studies exploring pathway biology in the budding yeast. ..
  10. Davis S, Wang J, Zhu M, Stahmer K, Lakshminarayan R, Ghassemian M, et al. Sec24 phosphorylation regulates autophagosome abundance during nutrient deprivation. elife. 2016;5: pubmed publisher
    ..During autophagy, Sec24 phosphorylation regulates autophagosome number and its interaction with the C-terminus of Atg9, a component of the autophagy machinery required for autophagosome initiation...
  11. Rao Y, Perna M, Hofmann B, Beier V, Wollert T. The Atg1-kinase complex tethers Atg9-vesicles to initiate autophagy. Nat Commun. 2016;7:10338 pubmed publisher
    ..Their biogenesis is initiated by recruitment of Atg9-vesicles to the phagophore assembly site...
  12. Backues S, Orban D, Bernard A, Singh K, Cao Y, Klionsky D. Atg23 and Atg27 act at the early stages of Atg9 trafficking in S. cerevisiae. Traffic. 2015;16:172-90 pubmed publisher
    b>Atg9 is a conserved multipass transmembrane protein with an essential role in autophagy. In Saccharomyces cerevisiae, it travels through the secretory pathway to a unique compartment, the Atg9 peripheral structures...
  13. Suzuki K, Akioka M, Kondo Kakuta C, Yamamoto H, Ohsumi Y. Fine mapping of autophagy-related proteins during autophagosome formation in Saccharomyces cerevisiae. J Cell Sci. 2013;126:2534-44 pubmed publisher
    ..We designate this localization the 'IM' pattern. The Atg2-Atg18 complex and Atg9 localized to the edge of the IM, appearing as two or three dots, in close proximity to the endoplasmic reticulum ..
  14. Wang K, Yang Z, Liu X, Mao K, Nair U, Klionsky D. Phosphatidylinositol 4-kinases are required for autophagic membrane trafficking. J Biol Chem. 2012;287:37964-72 pubmed publisher
    ..The PtdIns 4-kinase Pik1 is involved in Atg9 trafficking through the Golgi and is involved in both nonselective and selective types of autophagy, whereas the ..
  15. Yamamoto H, Kakuta S, Watanabe T, Kitamura A, Sekito T, Kondo Kakuta C, et al. Atg9 vesicles are an important membrane source during early steps of autophagosome formation. J Cell Biol. 2012;198:219-33 pubmed publisher
    ..In this study, we focus on the intracellular dynamics of Atg9, a multispanning membrane protein essential for autophagosome formation in yeast...
  16. Romanyuk D, Polak A, Maleszewska A, Sieńko M, Grynberg M, Zoładek T. Human hAtg2A protein expressed in yeast is recruited to preautophagosomal structure but does not complement autophagy defects of atg2? strain. Acta Biochim Pol. 2011;58:365-74 pubmed
    ..hAtg2A, in contrast to yeast Atg2, did not show interaction with the yeast autophagy protein Atg9 but both Atg2 proteins showed interaction with Atg18, a phospholipid-binding protein, in two-hybrid system...
  17. Webber J, Tooze S. New insights into the function of Atg9. FEBS Lett. 2010;584:1319-26 pubmed publisher
    ..Identification of more than 30 autophagy related proteins including a multi-spanning membrane protein, Atg9, has increased our understanding of the molecular mechanisms involved in autophagy...
  18. Cao Y, Nair U, Yasumura Yorimitsu K, Klionsky D. A multiple ATG gene knockout strain for yeast two-hybrid analysis. Autophagy. 2009;5:699-705 pubmed
    ..We also probed a new interaction using the MKO Y2H strain, and our results suggest that Atg29 and Atg31 interact independently of other known Atg proteins, and this interaction may mediate the interaction between Atg17 and Atg29. ..
  19. Kakuta S, Yamamoto H, Negishi L, Kondo Kakuta C, Hayashi N, Ohsumi Y. Atg9 vesicles recruit vesicle-tethering proteins Trs85 and Ypt1 to the autophagosome formation site. J Biol Chem. 2012;287:44261-9 pubmed publisher
    b>Atg9 is a transmembrane protein that is essential for autophagy. In the budding yeast Saccharomyces cerevisiae, it has recently been revealed that Atg9 exists on cytoplasmic small vesicles termed Atg9 vesicles...
  20. Araki Y, Ku W, Akioka M, May A, Hayashi Y, Arisaka F, et al. Atg38 is required for autophagy-specific phosphatidylinositol 3-kinase complex integrity. J Cell Biol. 2013;203:299-313 pubmed publisher
    ..These data suggest that the homodimer of Atg38 functions as a physical linkage between the Vps15-Vps34 and Atg14-Vps30 subcomplexes to facilitate complex I formation...
  21. Yao Z, Delorme Axford E, Backues S, Klionsky D. Atg41/Icy2 regulates autophagosome formation. Autophagy. 2015;11:2288-99 pubmed publisher
    ..Atg41 interacts with the transmembrane protein Atg9, a key component involved in autophagosome biogenesis, and both proteins display a similar localization profile...
  22. Lemus L, Ribas J, Sikorska N, Goder V. An ER-Localized SNARE Protein Is Exported in Specific COPII Vesicles for Autophagosome Biogenesis. Cell Rep. 2016;14:1710-1722 pubmed publisher
    ..increasingly exported from the ER and targeted to intracellular sites that contain the autophagy markers Atg8 and Atg9. In addition, Ufe1 interacts with non-ER SNARE proteins implicated in autophagosome formation...
  23. Mijaljica D, Prescott M, Devenish R. A late form of nucleophagy in Saccharomyces cerevisiae. PLoS ONE. 2012;7:e40013 pubmed publisher
    ..Moreover, the inhibition of LN in some mutants is accompanied by alterations in nuclear morphology. ..
  24. Jin M, He D, Backues S, Freeberg M, Liu X, Kim J, et al. Transcriptional regulation by Pho23 modulates the frequency of autophagosome formation. Curr Biol. 2014;24:1314-1322 pubmed publisher
    ..By RNA sequencing transcriptome profiling, we identified ATG9 as one of the key targets of Pho23, and our studies with strains expressing modulated levels of Atg9 show that the ..
  25. Kanki T, Wang K, Baba M, Bartholomew C, Lynch Day M, Du Z, et al. A genomic screen for yeast mutants defective in selective mitochondria autophagy. Mol Biol Cell. 2009;20:4730-8 pubmed publisher
    ..Accordingly, we have named this gene ATG33. The new mutants identified in this analysis will provide a useful foundation for researchers interested in the study of mitochondrial homeostasis and quality control. ..
  26. Mari M, Reggiori F. Atg9 trafficking in the yeast Saccharomyces cerevisiae. Autophagy. 2007;3:145-8 pubmed
    ..b>Atg9 is the only integral membrane component of the conserved Atg machinery and appears to be a crucial organizational ..
  27. He C, Klionsky D. Atg9 trafficking in autophagy-related pathways. Autophagy. 2007;3:271-4 pubmed
    ..Some important hints to these questions come from Atg9, which is the only integral membrane protein required for autophagosome formation and considered a membrane carrier ..
  28. Segarra V, Boettner D, Lemmon S. Atg27 tyrosine sorting motif is important for its trafficking and Atg9 localization. Traffic. 2015;16:365-78 pubmed publisher
    During autophagy, the transmembrane protein Atg27 facilitates transport of the major autophagy membrane protein Atg9 to the preautophagosomal structure (PAS)...
  29. Feng Y, Backues S, Baba M, Heo J, Harper J, Klionsky D. Phosphorylation of Atg9 regulates movement to the phagophore assembly site and the rate of autophagosome formation. Autophagy. 2016;12:648-58 pubmed publisher
    ..b>Atg9, conserved from yeast to mammals, is the only identified transmembrane protein in the yeast core macroautophagy ..