Q12142 (ATG9_YEAST) Reviewed, UniProtKB/Swiss-Prot

Names and origin

Protein names

Recommended name:
Autophagy-related protein 9

Alternative name(s):
Cytoplasm to vacuole targeting protein 7

Gene names

Name:	ATG9
Synonyms:	APG9, AUT9, CVT7
Ordered Locus Names:	YDL149W
ORF Names:	D1560

Organism

Saccharomyces cerevisiae (strain ATCC 204508 / S288c) (Baker's yeast)

Taxonomic identifier

559292 [NCBI]

Taxonomic lineage

Eukaryota › Fungi › Dikarya › Ascomycota › Saccharomycotina › Saccharomycetes › Saccharomycetales › Saccharomycetaceae › Saccharomyces

Protein attributes

Sequence length	997 AA.
Sequence status	Complete.
Protein existence	Evidence at protein level

General annotation (Comments)

Function	Involved in autophagy and cytoplasm to vacuole transport (Cvt) vesicle formation. Plays a key role in the organization of the preautophagosomal structure/phagophore assembly site (PAS), the nucleating site for formation of the sequestering vesicle. Cycles between the PAS and the cytoplasmic vesicle pool and may participate in supplying membrane for the growing autophagosome. Different machineries are required for anterograde trafficking to the PAS during either the Cvt pathway or bulk autophagy and for retrograde trafficking. Anterograde trafficking requires an intact actin cytoskeleton and proper retrograde trafficking of from the PAS to the cytoplasmic vesicles requires the ATG1-ATG13 complex and the phosphatidylinositol 3-kinase complex I. Recruits ATG23 and ATG8 to the PAS. Ref.4 Ref.5 Ref.6 Ref.7 Ref.8 Ref.9 Ref.12 Ref.17 Ref.18
Subunit structure	Self-interacts (via C-terminus); required for anterograde transport to the PAS. Interacts with ATG23 and ATG27 to form a cycling complex for trafficking to the PAS. Interacts (via N-terminus) with ATG11, required for recruitment of ATG9 to the PAS for the Cvt pathway during nutrient-rich conditions. Interacts (via N-terminus) with ATG17; required for recruitment to the PAS during autophagy and starved conditions. Interacts with ATG2 and ATG18; required for the retrieval of ATG9 from the PAS to the cytoplasmic pool. Ref.10 Ref.12 Ref.13 Ref.15 Ref.18 Ref.20
Subcellular location	Preautophagosomal structure membrane; Multi-pass membrane protein. Cytoplasmic vesicle membrane; Multi-pass membrane protein. Note: The peripheral pool of ATG9 partially colocalizes with mitochondria and it has been speculated that ATG9 resides either in the mitochondria or within vesicles in very close appostition to the mitochondria. Ref.7 Ref.8 Ref.13 Ref.14 Ref.20
Sequence similarities	Belongs to the ATG9 family.

Ontologies

Keywords
Biological process	Autophagy Protein transport Transport
Cellular component	Cytoplasmic vesicle Membrane
Domain	Transmembrane Transmembrane helix
PTM	Phosphoprotein
Technical term	Complete proteome Reference proteome
Gene Ontology (GO)
Biological process	CVT pathway Inferred from mutant phenotype Ref.8. Source: SGD autophagic vacuole assembly Inferred from mutant phenotype Ref.8. Source: SGD mitochondrion degradation Inferred from mutant phenotype. Source: SGD piecemeal microautophagy of nucleus Inferred from mutant phenotype. Source: SGD protein homooligomerization Inferred from mutant phenotype. Source: SGD
Cellular component	cytoplasmic vesicle membrane Inferred from electronic annotation. Source: UniProtKB-SubCell integral to membrane Inferred from electronic annotation. Source: UniProtKB-KW membrane fraction Inferred from direct assay Ref.8. Source: SGD mitochondrion Inferred from direct assay Ref.14. Source: SGD pre-autophagosomal structure membrane Inferred from electronic annotation. Source: UniProtKB-SubCell
Complete GO annotation...

Sequence annotation (Features)

	Feature key	Position(s)	Length	Description	Graphical view	Feature identifier
Molecule processing
	Chain	1 – 997	997	Autophagy-related protein 9		PRO_0000119839
Regions
	Topological domain	1 – 318	318	Cytoplasmic Probable
	Transmembrane	319 – 339	21	Helical; Potential
	Topological domain	340 – 376	37	Lumenal Probable
	Transmembrane	377 – 397	21	Helical; Potential
	Topological domain	398 – 538	141	Cytoplasmic Probable
	Transmembrane	539 – 559	21	Helical; Potential
	Topological domain	560 – 620	61	Lumenal Probable
	Transmembrane	621 – 641	21	Helical; Potential
	Topological domain	642 – 656	15	Cytoplasmic Probable
	Transmembrane	657 – 677	21	Helical; Potential
	Topological domain	678 – 723	46	Lumenal Probable
	Transmembrane	724 – 744	21	Helical; Potential
	Topological domain	745 – 997	253	Cytoplasmic Probable
	Compositional bias	907 – 978	72	Asn-rich
Amino acid modifications
	Modified residue	111	1	Phosphothreonine Ref.19
	Modified residue	112	1	Phosphoserine Ref.19
	Modified residue	787	1	Phosphoserine Ref.19
Experimental info
	Mutagenesis	192	1	H → L: Aboloshes interaction with ATG11 and disrupts Cvt, but not bulk autophagy. Ref.15

Sequences

	Sequence		Length	Mass (Da)
	Q12142 [UniParc]. Last modified November 1, 1996. Version 1. Checksum: E900AD5E3AC5D111	FASTA	997	115,403
	10 20 30 40 50 60 MERDEYQLPN SHGKNTFLSR IFGLQSDEVN PSLNSQEMSN FPLPDIERGS SLLHSTNDSR 70 80 90 100 110 120 EDVDENDLRV PESDQGTSTE EEDEVDEEQV QAYAPQISDG LDGDHQLNSV TSKENVLETE 130 140 150 160 170 180 KSNLERLVEG STDDSVPKVG QLSSEEEEDN EFINNDGFDD DTPLFQKSKI HEFSSKKSNT 190 200 210 220 230 240 IEDGKRPLFF RHILQNNRPQ RDTQKLFTSS NAIHHDKDKS ANNGPRNING NQKHGTKYFG 250 260 270 280 290 300 SATQPRFTGS PLNNTNRFTK LFPLRKPNLL SNISVLNNTP EDRINTLSVK ERALWKWANV 310 320 330 340 350 360 ENLDIFLQDV YNYYLGNGFY CIILEKILNI CTLLFVVFVS TYMGHCVDYS KLPTSHRVSD 370 380 390 400 410 420 IIIDKCYSNS ITGFTKFFLW MFYFFVILKI VQLYFDVQKL SELQNFYKYL LNISDDELQT 430 440 450 460 470 480 LPWQNVIQQL MYLKDQNAMT ANVVEVKAKN RIDAHDVANR IMRRENYLIA LYNSDILNLS 490 500 510 520 530 540 LPIPLFRTNV LTKTLEWNIN LCVMGFVFNE SGFIKQSILK PSQREFTREE LQKRFMLAGF 550 560 570 580 590 600 LNIILAPFLV TYFVLLYFFR YFNEYKTSPG SIGARQYTPI AEWKFREYNE LYHIFKKRIS 610 620 630 640 650 660 LSTTLANKYV DQFPKEKTNL FLKFVSFICG SFVAILAFLT VFDPENFLNF EITSDRSVIF 670 680 690 700 710 720 YITILGAIWS VSRNTITQEY HVFDPEETLK ELYEYTHYLP KEWEGRYHKE EIKLEFCKLY 730 740 750 760 770 780 NLRIVILLRE LTSLMITPFV LWFSLPSSAG RIVDFFRENS EYVDGLGYVC KYAMFNMKNI 790 800 810 820 830 840 DGEDTHSMDE DSLTKKIAVN GSHTLNSKRR SKFTAEDHSD KDLANNKMLQ SYVYFMDDYS 850 860 870 880 890 900 NSENLTGKYQ LPAKKGYPNN EGDSFLNNKY SWRKQFQPGQ KPELFRIGKH ALGPGHNISP 910 920 930 940 950 960 AIYSTRNPGK NWDNNNNGDD IKNGTNNATA KNDDNNGNND HEYVLTESFL DSGAFPNHDV 970 980 990 IDHNKMLNSN YNGNGILNKG GVLGLVKEYY KKSDVGR

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"Analysis of a 23 kb region on the left arm of yeast chromosome IV." Delaveau T.T.D., Blugeon C., Jacq C., Perea J. Yeast 12:1587-1592(1996) [PubMed: 8972581] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [GENOMIC DNA].
[2]	"The nucleotide sequence of Saccharomyces cerevisiae chromosome IV." Jacq C., Alt-Moerbe J., Andre B., Arnold W., Bahr A., Ballesta J.P.G., Bargues M., Baron L., Becker A., Biteau N., Bloecker H., Blugeon C., Boskovic J., Brandt P., Brueckner M., Buitrago M.J., Coster F., Delaveau T. , del Rey F., Dujon B., Eide L.G., Garcia-Cantalejo J.M., Goffeau A., Gomez-Peris A., Granotier C., Hanemann V., Hankeln T., Hoheisel J.D., Jaeger W., Jimenez A., Jonniaux J.-L., Kraemer C., Kuester H., Laamanen P., Legros Y., Louis E.J., Moeller-Rieker S., Monnet A., Moro M., Mueller-Auer S., Nussbaumer B., Paricio N., Paulin L., Perea J., Perez-Alonso M., Perez-Ortin J.E., Pohl T.M., Prydz H., Purnelle B., Rasmussen S.W., Remacha M.A., Revuelta J.L., Rieger M., Salom D., Saluz H.P., Saiz J.E., Saren A.-M., Schaefer M., Scharfe M., Schmidt E.R., Schneider C., Scholler P., Schwarz S., Soler-Mira A., Urrestarazu L.A., Verhasselt P., Vissers S., Voet M., Volckaert G., Wagner G., Wambutt R., Wedler E., Wedler H., Woelfl S., Harris D.E., Bowman S., Brown D., Churcher C.M., Connor R., Dedman K., Gentles S., Hamlin N., Hunt S., Jones L., McDonald S., Murphy L.D., Niblett D., Odell C., Oliver K., Rajandream M.A., Richards C., Shore L., Walsh S.V., Barrell B.G., Dietrich F.S., Mulligan J.T., Allen E., Araujo R., Aviles E., Berno A., Carpenter J., Chen E., Cherry J.M., Chung E., Duncan M., Hunicke-Smith S., Hyman R.W., Komp C., Lashkari D., Lew H., Lin D., Mosedale D., Nakahara K., Namath A., Oefner P., Oh C., Petel F.X., Roberts D., Schramm S., Schroeder M., Shogren T., Shroff N., Winant A., Yelton M.A., Botstein D., Davis R.W., Johnston M., Andrews S., Brinkman R., Cooper J., Ding H., Du Z., Favello A., Fulton L., Gattung S., Greco T., Hallsworth K., Hawkins J., Hillier L.W., Jier M., Johnson D., Johnston L., Kirsten J., Kucaba T., Langston Y., Latreille P., Le T., Mardis E., Menezes S., Miller N., Nhan M., Pauley A., Peluso D., Rifkin L., Riles L., Taich A., Trevaskis E., Vignati D., Wilcox L., Wohldman P., Vaudin M., Wilson R., Waterston R., Albermann K., Hani J., Heumann K., Kleine K., Mewes H.-W., Zollner A., Zaccaria P. Nature 387:75-78(1997) [PubMed: 9169867] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA]. Strain: ATCC 204508 / S288c.
[3]	Saccharomyces Genome Database Submitted (DEC-2009) to the EMBL/GenBank/DDBJ databases Cited for: GENOME REANNOTATION. Strain: ATCC 204508 / S288c.
[4]	"Isolation and characterization of autophagy-defective mutants of Saccharomyces cerevisiae." Tsukada M., Ohsumi Y. FEBS Lett. 333:169-174(1993) [PubMed: 8224160] [Abstract] Cited for: FUNCTION.
[5]	"Isolation and characterization of yeast mutants in the cytoplasm to vacuole protein targeting pathway." Harding T.M., Morano K.A., Scott S.V., Klionsky D.J. J. Cell Biol. 131:591-602(1995) [PubMed: 7593182] [Abstract] Cited for: FUNCTION.
[6]	"Genetic and phenotypic overlap between autophagy and the cytoplasm to vacuole protein targeting pathway." Harding T.M., Hefner-Gravink A., Thumm M., Klionsky D.J. J. Biol. Chem. 271:17621-17624(1996) [PubMed: 8663607] [Abstract] Cited for: FUNCTION.
[7]	"Autophagy and the cvt pathway both depend on AUT9." Lang T., Reiche S., Straub M., Bredschneider M., Thumm M. J. Bacteriol. 182:2125-2133(2000) [PubMed: 10735854] [Abstract] Cited for: FUNCTION, SUBCELLULAR LOCATION.
[8]	"Apg9p/Cvt7p is an integral membrane protein required for transport vesicle formation in the Cvt and autophagy pathways." Noda T., Kim J., Huang W.-P., Baba M., Tokunaga C., Ohsumi Y., Klionsky D.J. J. Cell Biol. 148:465-480(2000) [PubMed: 10662773] [Abstract] Cited for: FUNCTION, SUBCELLULAR LOCATION.
[9]	"The pre-autophagosomal structure organized by concerted functions of APG genes is essential for autophagosome formation." Suzuki K., Kirisako T., Kamada Y., Mizushima N., Noda T., Ohsumi Y. EMBO J. 20:5971-5981(2001) [PubMed: 11689437] [Abstract] Cited for: FUNCTION.
[10]	"Apg2 is a novel protein required for the cytoplasm to vacuole targeting, autophagy, and pexophagy pathways." Wang C.-W., Kim J., Huang W.-P., Abeliovich H., Stromhaug P.E., Dunn W.A. Jr., Klionsky D.J. J. Biol. Chem. 276:30442-30451(2001) [PubMed: 11382760] [Abstract] Cited for: INTERACTION WITH ATG2.
[11]	"A unified nomenclature for yeast autophagy-related genes." Klionsky D.J., Cregg J.M., Dunn W.A. Jr., Emr S.D., Sakai Y., Sandoval I.V., Sibirny A., Subramani S., Thumm M., Veenhuis M., Ohsumi Y. Dev. Cell 5:539-545(2003) [PubMed: 14536056] [Abstract] Cited for: NOMENCLATURE.
[12]	"Atg23 is essential for the cytoplasm to vacuole targeting pathway and efficient autophagy but not pexophagy." Tucker K.A., Reggiori F., Dunn W.A. Jr., Klionsky D.J. J. Biol. Chem. 278:48445-48452(2003) [PubMed: 14504273] [Abstract] Cited for: FUNCTION, INTERACTION WITH ATG23.
[13]	"The Atg1-Atg13 complex regulates Atg9 and Atg23 retrieval transport from the pre-autophagosomal structure." Reggiori F., Tucker K.A., Stromhaug P.E., Klionsky D.J. Dev. Cell 6:79-90(2004) [PubMed: 14723849] [Abstract] Cited for: SUBCELLULAR LOCATION, INTERACTION WITH ATG18, TRAFFICKING.
[14]	"Atg9 cycles between mitochondria and the pre-autophagosomal structure in yeasts." Reggiori F., Shintani T., Nair U., Klionsky D.J. Autophagy 1:101-109(2005) [PubMed: 16874040] [Abstract] Cited for: SUBCELLULAR LOCATION.
[15]	"Recruitment of Atg9 to the preautophagosomal structure by Atg11 is essential for selective autophagy in budding yeast." He C., Song H., Yorimitsu T., Monastyrska I., Yen W.-L., Legakis J.E., Klionsky D.J. J. Cell Biol. 175:925-935(2006) [PubMed: 17178909] [Abstract] Cited for: INTERACTION WITH ATG11; ATG23 AND ATG27, MUTAGENESIS OF HIS-192.
[16]	"A global topology map of the Saccharomyces cerevisiae membrane proteome." Kim H., Melen K., Oesterberg M., von Heijne G. Proc. Natl. Acad. Sci. U.S.A. 103:11142-11147(2006) [PubMed: 16847258] [Abstract] Cited for: TOPOLOGY [LARGE SCALE ANALYSIS]. Strain: ATCC 208353 / W303-1A.
[17]	"Atg9 trafficking in autophagy-related pathways." He C., Klionsky D.J. Autophagy 3:271-274(2007) [PubMed: 17329962] [Abstract] Cited for: FUNCTION.
[18]	"Self-interaction is critical for Atg9 transport and function at the phagophore assembly site during autophagy." He C., Baba M., Cao Y., Klionsky D.J. Mol. Biol. Cell 19:5506-5516(2008) [PubMed: 18829864] [Abstract] Cited for: FUNCTION, SUBUNIT.
[19]	"A multidimensional chromatography technology for in-depth phosphoproteome analysis." Albuquerque C.P., Smolka M.B., Payne S.H., Bafna V., Eng J., Zhou H. Mol. Cell. Proteomics 7:1389-1396(2008) [PubMed: 18407956] [Abstract] Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT THR-111; SER-112 AND SER-787, MASS SPECTROMETRY.
[20]	"Atg17 recruits Atg9 to organize the pre-autophagosomal structure." Sekito T., Kawamata T., Ichikawa R., Suzuki K., Ohsumi Y. Genes Cells 14:525-538(2009) [PubMed: 19371383] [Abstract] Cited for: INTERACTION WITH ATG11 AND ATG17, SUBCELLULAR LOCATION.
+	Additional computationally mapped references.

Cross-references

Sequence databases
EMBL GenBank DDBJ	X97751 Genomic DNA. Translation: CAA66342.1. Z74197 Genomic DNA. Translation: CAA98723.1. BK006938 Genomic DNA. Translation: DAA11709.1.
PIR	S67697.
RefSeq	NP_010132.1. NM_001180209.1.
3D structure databases
ProteinModelPortal	Q12142.
ModBase	Search...
Protein-protein interaction databases
DIP	DIP-1938N.
IntAct	Q12142. 10 interactions.
MINT	MINT-404580.
STRING	Q12142.
Protocols and materials databases
StructuralBiologyKnowledgebase	Search...
Genome annotation databases
EnsemblFungi	YDL149W; YDL149W; YDL149W.
GeneID	851406.
KEGG	sce:YDL149W.
NMPDR	fig|4932.3.peg.869.
Organism-specific databases
CYGD	YDL149w.
SGD	S000002308. ATG9.
Phylogenomic databases
eggNOG	fuNOG04500.
GeneTree	EFGT00050000004380.
HOGENOM	HBG447720.
OMA	EYTHYLP.
OrthoDB	EOG4Q2HPN.
Gene expression databases
ArrayExpress	Q12142.
Genevestigator	Q12142.
GermOnline	YDL149W. Saccharomyces cerevisiae.
Family and domain databases
InterPro	IPR007241. Autophagy-rel_prot_9. [Graphical view]
PANTHER	PTHR13038. APG9. 1 hit.
Pfam	PF04109. APG9. 1 hit. [Graphical view]
ProtoNet	Search...
Other
NextBio	968579.

Entry information

Entry name

ATG9_YEAST

Accession

Primary (citable) accession number: Q12142
Secondary accession number(s): D6VRJ9

Entry history

Integrated into UniProtKB/Swiss-Prot:	March 15, 2005
Last sequence update:	November 1, 1996
Last modified:	December 14, 2011
This is version 86 of the entry and version 1 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation program

Fungal Protein Annotation Program