Affinage

ATG9A

Autophagy-related protein 9A · UniProt Q7Z3C6

Length
839 aa
Mass
94.4 kDa
Annotated
2026-04-28
100 papers in source corpus 50 papers cited in narrative 50 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

ATG9A is the sole multi-spanning transmembrane protein of the core autophagy machinery, functioning as a homotrimeric lipid scramblase that equilibrates phospholipids across membrane bilayers to drive autophagosome membrane expansion in concert with the lipid transfer protein ATG2A (PMID:33106659, PMID:36347259, PMID:39174844). ATG9A cycles between the trans-Golgi network, recycling endosomes, and the plasma membrane via AP-4/RUSC2/kinesin-1-dependent trafficking, and its proper itinerary—regulated by AP-2, SNX4, retromer, RHOD, and post-translational modifications including ubiquitination and S-palmitoylation—is essential for delivering lipid-metabolizing enzymes (PI4KIIIβ, PI4K2A) and nucleating autophagosome biogenesis through its C-terminal interaction with the ATG13–ATG101 HORMA dimer of the ULK1 complex (PMID:29180427, PMID:30917996, PMID:36791199, PMID:34369648, PMID:40394978). Beyond canonical autophagy, ATG9A performs autophagy-independent functions: it organizes IQGAP1–ESCRT-III at the plasma membrane to mediate membrane repair, promotes LC3-independent lysosomal degradation of TNF complex IIa to prevent apoptosis, negatively regulates STING–TBK1 innate immune signaling, controls lipid mobilization from lipid droplets, facilitates unconventional galectin-9 secretion, and supports neurite outgrowth and axonal integrity (PMID:34257406, PMID:36520901, PMID:19926846, PMID:34799570, PMID:40335523, PMID:28513333). ATG9A also partners with IQGAP1 and CHMP2A to mediate autophagosome closure, establishing it as a central hub operating at every stage of autophagosome biogenesis (PMID:39745851).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 2009 High

    Establishing that ATG9A has autophagy-independent roles: ATG9A was shown to negatively regulate STING–TBK1 assembly during innate immune sensing of dsDNA, a function not shared by the downstream autophagy factor ATG7, revealing that ATG9A acts at the interface of membrane trafficking and innate immunity.

    Evidence Atg9a KO macrophages with STING/TBK1 co-IP and IFN-β induction assays, compared with Atg7 KO

    PMID:19926846

    Open questions at the time
    • Mechanism by which ATG9A suppresses STING–TBK1 assembly not defined at molecular level
    • Whether ATG9A's scramblase activity contributes to STING regulation unknown
  2. 2016 Medium

    Defining ATG9A's intracellular trafficking itinerary: a series of studies identified sorting motifs controlling ER-to-Golgi transport (G516), Golgi retention (L711YM713), recycling endosome exit (AP-2 binding motifs), and showed that ATG9A self-interaction promotes biosynthetic transport, establishing the multi-compartment cycling model essential for autophagosome nucleation.

    Evidence Sequential mutagenesis of ATG9A sorting motifs, AP-2 co-IP, TRAPPC8 knockdown, subcellular localization in HeLa cells

    PMID:27316455 PMID:27587839 PMID:27663665

    Open questions at the time
    • Structural basis of ATG9A self-interaction not resolved
    • Relative contributions of each sorting motif to autophagy flux not quantified
  3. 2016 High

    Revealing ATG9A-specific neuronal functions: brain-specific Atg9a conditional knockout caused neurite outgrowth defects and corpus callosum dysgenesis not phenocopied by Atg7 or Atg16l1 KO, and Atg9a-dependent necrosis was identified during bone morphogenesis, demonstrating developmental roles beyond canonical autophagy.

    Evidence Conditional Atg9a KO mice, diffusion tensor MRI, primary neuron culture, Atg5 KO comparison for bone phenotype

    PMID:27811852 PMID:28513333

    Open questions at the time
    • Molecular mechanism of autophagy-independent neurite outgrowth promotion unknown
    • Targets of Atg9a-dependent developmental necrosis not identified
  4. 2017 High

    Identifying AP-4 as the adaptor sorting ATG9A from the TGN: AP-4 loss caused ATG9A retention at the TGN, impaired LC3B lipidation, and blocked preautophagosomal structure maturation, establishing the AP-4–ATG9A axis as a critical supply route for autophagosome membranes.

    Evidence AP-4 subunit KO/KD in multiple cell types, LC3B lipidation assay, AP-4 ε KO mice with axonal phenotypes and patient fibroblasts

    PMID:29180427 PMID:29698489 PMID:30262884

    Open questions at the time
    • Direct recognition motif on ATG9A for AP-4 binding not mapped at structural resolution
    • Whether AP-4 sorts other autophagy-relevant cargo alongside ATG9A remains open
  5. 2019 High

    Demonstrating that ATG9A vesicles deliver lipid-metabolizing enzymes to autophagosome initiation sites: immunoisolation and mass spectrometry revealed ATG9A vesicles carry PI4KIIIβ and BAR-domain proteins; PI4KIIIβ interacts with both ATG9A and ATG13 to generate PI4P at initiation membranes.

    Evidence Immunoisolation of ATG9A vesicles with quantitative MS, reciprocal co-IP of PI4KIIIβ–ATG9A–ATG13, siRNA with PI4P readouts

    PMID:30917996

    Open questions at the time
    • Whether PI4P production is coupled to ATG9A scramblase activity not tested
    • Full lipid composition of ATG9A vesicles not determined
  6. 2020 High

    Solving the structure and enzymatic activity of ATG9A: two independent cryo-EM studies revealed a domain-swapped homotrimer with a central pore and lateral cavities, establishing ATG9A as a lipid scramblase whose pore mutations reduce phospholipid flipping and autophagosome size, and whose architecture induces membrane curvature.

    Evidence Cryo-EM at 2.9 Å, in vitro scramblase reconstitution assay, MD simulation, mutagenesis with autophagosome size readout

    PMID:32610138 PMID:33106659

    Open questions at the time
    • Lipid substrate specificity of scramblase not fully defined
    • How scrambling rate is regulated in vivo (e.g., by post-translational modifications) unknown
  7. 2020 Medium

    Expanding ATG9A trafficking regulators: SNX4–SNX7 heterodimer, FHF/dynein, and RUSC2/kinesin-1 were shown to coordinate bidirectional microtubule-based transport of ATG9A vesicles, with OPTN recruiting ATG9A vesicles to ubiquitinated mitochondria for selective mitophagy.

    Evidence SNX4/SNX7 CRISPR KO, FHF KD with ATG9A localization, RUSC2–kinesin-1 co-IP, OPTN–ATG9A co-IP with mitophagy rescue

    PMID:32073997 PMID:32513819 PMID:32556086

    Open questions at the time
    • How cargo receptor binding switches ATG9A from recycling to autophagosome nucleation not resolved
    • Relative contributions of anterograde vs retrograde transport to ATG9A steady-state distribution not quantified
  8. 2021 High

    Discovering autophagy-independent membrane repair and lipid mobilization functions: ATG9A organizes IQGAP1 and ESCRT at the plasma membrane to protect against permeabilization by diverse agents, and separately controls fatty acid transfer from lipid droplets to mitochondria, broadening ATG9A's functional repertoire well beyond autophagy.

    Evidence ATG9A KO with plasma membrane permeabilization by gasdermin/MLKL/ORF3a, IQGAP1/ESCRT co-IP; ATG9A depletion in human cells and C. elegans with lipid droplet and mitochondrial respiration assays

    PMID:34257406 PMID:34799570

    Open questions at the time
    • Whether ATG9A scramblase activity is required for membrane repair not tested
    • Molecular mechanism linking ATG9A to lipid droplet–mitochondria fatty acid transfer undefined
  9. 2021 Medium

    Mapping the ATG9A–ULK1 complex interface: BioID and split-fluorescence approaches showed ATG9A directly contacts the ATG13–ATG101 HORMA dimer independently of ULK1, and loss of ATG13/ATG101 mislocalizes ATG9A to stalled p62–ubiquitin clusters.

    Evidence BioID proximity proteomics, split-mVenus ATG13–ATG101 capture, ATG13/ATG101 KO with ATG9A localization

    PMID:34369648 PMID:35442099

    Open questions at the time
    • Stoichiometry of ATG9A trimer to ATG13–ATG101 dimer on membranes not established
    • Whether ATG13–ATG101 activates ATG9A scramblase activity not tested
  10. 2022 High

    Reconstituting the ATG2A–ATG9A lipid transfer–scramblase couple and revealing non-autophagic TNF checkpoint: structural and biochemical studies defined how ATG2A directly delivers lipids into ATG9A's perpendicular branch for scrambling; separately, ATG9A was shown to promote LC3-independent lysosomal degradation of TNF complex IIa, constituting a cell-death checkpoint preventing embryonic lethality.

    Evidence HDX-MS/crosslinking MS/cryo-EM of ATG2A–ATG9A interface with mutagenesis; ATG9A KO mice with TNF complex IIa isolation and lysosomal targeting assays

    PMID:36347259 PMID:36520901

    Open questions at the time
    • Whether other lipid transfer proteins besides ATG2A can couple with ATG9A scramblase not tested
    • Mechanism of LC3-independent lysosomal targeting by ATG9A not molecularly defined
  11. 2022 Medium

    Identifying post-translational ubiquitination as a regulatory switch: TRAF6-mediated K48/K63 ubiquitination enhances ATG9A–Beclin 1 association and VPS34 activation under ROS, while MARCH9-mediated K63 ubiquitination disperses ATG9A from the Golgi to cause GRASP55-dependent Golgi fragmentation under heat stress.

    Evidence TRAF6/A20 co-IP with ubiquitin chain analysis and VPS34 activity assay; MARCH9 KO with GRASP55 oligomerization and Golgi morphology

    PMID:35196483 PMID:35977480

    Open questions at the time
    • Whether the two ubiquitination events compete for the same sites not determined
    • In vivo relevance of MARCH9-dependent Golgi fragmentation beyond heat stress unclear
  12. 2023 High

    Atomic-resolution structure of ATG9A C-tail bound to ATG13–ATG101 HORMA dimer: the 2.4 Å crystal structure revealed β-sheet complementation in the HORMA cleft, and disruption impaired PINK1/Parkin mitophagy, providing the structural basis for autophagosome nucleation.

    Evidence X-ray crystallography at 2.4 Å, structure-guided mutations with mitophagy assay

    PMID:36791199

    Open questions at the time
    • Whether HORMA binding allosterically activates ATG9A scramblase not tested
    • Full-length ATG9A trimer–ATG13–ATG101 complex structure not yet available
  13. 2024 High

    Visualizing the complete ATG2A–WIPI4–ATG9A lipid transfer complex: cryo-EM at 7 Å showed 3:1 stoichiometry (ATG9A trimer:ATG2A) with ATG9A lateral pore aligned to ATG2A lipid cavity; cryo-ET demonstrated ATG2A tethers donor and acceptor vesicles.

    Evidence Cryo-EM, cryo-electron tomography, MD simulation

    PMID:39174844

    Open questions at the time
    • How WIPI4 PI3P binding coordinates complex assembly on phagophore membranes not resolved in situ
    • Dynamic regulation of lipid transfer rate not measured
  14. 2025 Medium

    Extending ATG9A to autophagosome closure and unconventional secretion: ATG9A partners with IQGAP1–CHMP2A/ESCRT-III for autophagosome closure, making it active at all biogenesis stages; separately, ATG9A vesicles mediate galectin-9 unconventional secretion via TMED10-dependent cargo loading and STX13–SNAP23–VAMP3 SNARE-mediated plasma membrane fusion.

    Evidence ATG9A KO reconstitution with in vitro closure assay (SolVit) and IQGAP1/CHMP2A co-IP; ATG9A KO with galectin-9 secretion assay, N-terminus mutagenesis, SNARE identification

    PMID:39745851 PMID:40335523

    Open questions at the time
    • How ATG9A transitions from expansion to closure function mechanistically undefined
    • Spectrum of unconventionally secreted cargoes dependent on ATG9A vesicles not catalogued
  15. 2025 Medium

    Discovering ATG9A vesicle roles in lysosomal repair and lipid modification-dependent trafficking: ATG9A vesicles deliver PI4K2A to damaged lysosomes cooperating with ARFIP2 for PI4P control, and S-palmitoylation of ATG9A coordinates its trafficking for autophagy initiation.

    Evidence ATG9A KO with lysosomal damage assays and PI4K2A co-IP; acyl-biotin exchange and palmitoylation-site mutagenesis with trafficking assays

    PMID:40394978 PMID:40460835

    Open questions at the time
    • Palmitoyl-transferase responsible for ATG9A palmitoylation not identified
    • Whether PI4K2A delivery requires ATG9A scramblase activity not tested
    • Interplay between palmitoylation and ubiquitination in ATG9A trafficking regulation not examined

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions: how ATG9A's scramblase activity is dynamically regulated in vivo (by post-translational modifications, lipid environment, or binding partners); how ATG9A transitions between its distinct functions at autophagosome initiation, expansion, closure, plasma membrane repair, and unconventional secretion; and the full spectrum of non-autophagic cargoes and contexts dependent on ATG9A vesicle trafficking.
  • No in vivo measurement of ATG9A scramblase rate or regulation exists
  • Complete lipidome of ATG9A vesicles undetermined
  • Structural basis for ATG9A's multi-functionality (IQGAP1/ESCRT binding vs ATG2A coupling) not resolved

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0005215 transporter activity 3 GO:0008289 lipid binding 3 GO:0060090 molecular adaptor activity 3
Localization
GO:0031410 cytoplasmic vesicle 8 GO:0005794 Golgi apparatus 6 GO:0005768 endosome 4 GO:0005886 plasma membrane 3 GO:0005783 endoplasmic reticulum 2
Pathway
R-HSA-9612973 Autophagy 9 R-HSA-5653656 Vesicle-mediated transport 6 R-HSA-9609507 Protein localization 5 R-HSA-168256 Immune System 2 R-HSA-5357801 Programmed Cell Death 2 R-HSA-1430728 Metabolism 1
Partners
AP4E1ATG101ATG13ATG2ACHMP2AIQGAP1PI4KBRUSC2
Complex memberships
ATG2A-WIPI4-ATG9A lipid transfer complexATG9A homotrimerATG9A-IQGAP1-CHMP2A closure complex

Evidence

Reading pass · 50 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2020 ATG9A is a lipid scramblase that equilibrates phospholipids across membrane bilayers. Cryo-EM structures reveal a homotrimeric architecture with a solvated central pore connected laterally to the cytosol through cavities within each protomer, analogous to ABC exporters. Molecular dynamics simulations suggest the central pore opens laterally to accommodate lipid headgroups enabling lipid flipping. Mutations in the pore reduce scrambling activity and yield markedly smaller autophagosomes, establishing that lipid scrambling by ATG9A is essential for autophagosome membrane expansion. Cryo-EM structure determination, in vitro scramblase assay, molecular dynamics simulation, mutagenesis with functional readout (autophagosome size) Nature structural & molecular biology High 33106659
2020 Human ATG9A adopts a domain-swapped homotrimeric fold with multiple membrane spans and a network of branched internal cavities consistent with a membrane transporter. Structure-guided molecular simulations predict ATG9A causes membrane bending, explaining its localization to small vesicles and highly curved edges of growing autophagosomes. Mutational analyses of cavity-lining residues support their role in ATG9A function. Cryo-EM structure at 2.9-Å resolution, molecular dynamics simulation, in cellulo mutagenesis Cell reports High 32610138
2022 ATG9A and ATG2A form a heteromeric complex in which ATG2A (a lipid transfer protein) directly transfers lipids into the lipid-binding perpendicular branch of ATG9A (the scramblase), enabling coupled lipid transport and re-equilibration during autophagosome membrane expansion. The complex interface was defined by peptide arrays, crosslinking, hydrogen-deuterium exchange MS, and cryo-EM, and interface mutations impair autophagy. Peptide arrays, crosslinking MS, HDX-MS, cryo-EM, integrative structure modeling, mutagenesis + functional autophagy assays Molecular cell High 36347259
2024 Cryo-EM structures of the ATG2A–WIPI4–ATG9A complex at 7 Å reveal 3:1 stoichiometry of ATG9A trimer to ATG2A, directly aligning the ATG9A lateral pore with the ATG2A lipid transfer cavity, and show ATG9A interacts with both the N-terminal and C-terminal tip of rod-shaped ATG2A. Cryo-ET showed ATG2A tethers lipid vesicles at different orientations, and MD simulations propose a mechanism for lipid extraction from donor membranes. Cryo-EM, cryo-electron tomography, molecular dynamics simulation Nature structural & molecular biology High 39174844
2023 The C-terminal tail of ATG9A contains a 'HORMA dimer-interacting region' (HDIR) that binds the ATG13:ATG101 HORMA dimer of the ULK1 complex via β-sheet complementation, nestled in a deep cleft at the ATG13:ATG101 interface. X-ray crystallographic structure at 2.4 Å was determined; disruption of this complex in cells impairs PINK1/Parkin mitophagy mediated by the cargo receptor NDP52. X-ray crystallography (2.4 Å), cell-based mitophagy assays with structure-guided mutations Science advances High 36791199
2009 ATG9A co-localizes with STING after dsDNA stimulation and negatively regulates STING–TBK1 assembly. Loss of Atg9a (but not Atg7) greatly enhances STING–TBK1 assembly and causes aberrant activation of innate immune responses to dsDNA, establishing ATG9A as an autophagy-independent regulator of innate immunity. Atg9a knockout macrophages, co-immunoprecipitation of STING/TBK1, immunofluorescence co-localization, IFN-β induction assays Proceedings of the National Academy of Sciences of the United States of America High 19926846
2017 AP-4 promotes signal-mediated export of ATG9A from the trans-Golgi network to the peripheral cytoplasm. Loss of AP-4 causes ATG9A retention in the TGN, impairs LC3B lipidation, and blocks maturation of preautophagosomal structures, identifying ATG9A as a specific AP-4 cargo. AP-4 subunit knockdown/knockout, immunofluorescence and subcellular fractionation of ATG9A, LC3B lipidation assay Proceedings of the National Academy of Sciences of the United States of America High 29180427
2018 AP-4 sorts ATG9A into vesicles that are transported to the cell periphery by the accessory protein RUSC2. AP-4 deficiency causes ATG9A missorting in multiple cell types including patient-derived fibroblasts, leading to dysregulated autophagy. ATG9A-positive peripheral vesicles cluster near autophagosomes, suggesting they form the 'ATG9A reservoir' required for autophagosome biogenesis. Dynamic Organellar Maps proteomics, AP-4 KO cell lines, patient fibroblasts, immunofluorescence, co-localization with autophagosomes Nature communications High 30262884
2020 The autophagy adaptor OPTN forms a complex with ATG9A vesicles during mitophagy. Disruption of OPTN–ATG9A interactions prevents mitophagy initiation, demonstrating that autophagy adaptors directly recruit ATG9A vesicles (core autophagy units) to ubiquitinated mitochondria to nucleate autophagosomal membrane de novo. Chemical-induced mitochondrial ubiquitination system, phase-separated fluorescent foci assay, OPTN–ATG9A co-immunoprecipitation, knockout/rescue experiments The Journal of cell biology High 32556086
2019 ATG9A vesicles are enriched in BAR-domain proteins (Arfaptins) and phosphoinositide-metabolizing enzymes. Arfaptin2 regulates starvation-dependent distribution of ATG9A vesicles. These vesicles deliver PI4-kinase PI4KIIIβ to the autophagosome initiation site. PI4KIIIβ interacts with both ATG9A and ATG13 to control PI4P production at the initiation membrane site. Immunoisolation of ATG9A vesicles + quantitative mass spectrometry, co-immunoprecipitation of PI4KIIIβ–ATG9A–ATG13, siRNA knockdown with PI4P and autophagy readouts The Journal of cell biology High 30917996
2021 ATG9A depletion increases the size and/or number of lipid droplets and blocks transfer of fatty acids from lipid droplets to mitochondria, impairing mitochondrial respiration. ATG9A localizes to vesicular-tubular clusters (VTCs) closely associated with an ER subdomain enriched in TMEM41B and in proximity to lipid droplets and mitochondria, indicating a non-autophagic role in lipid mobilization. ATG9A depletion in human cells and C. elegans, lipid droplet imaging, fatty acid transfer assay (isotope tracing), mitochondrial respiration assay, subcellular localization by immunofluorescence and electron microscopy Nature communications High 34799570
2021 ATG9A organizes IQGAP1 and ESCRT system components at the plasma membrane to protect against plasma membrane damage. ATG9A knockout sensitizes cells to permeabilization by diverse microbial and endogenous agents including gasdermin, MLKL, and coronavirus ORF3a, revealing an autophagy-independent role in plasma membrane integrity maintenance. ATG9A KO cells, plasma membrane permeabilization assays with multiple agents, co-immunoprecipitation of ATG9A–IQGAP1–ESCRT, live-cell imaging Nature cell biology High 34257406
2022 ATG9A and FIP200 promote degradation of the cytotoxic complex IIa formed upon TNF sensing through an LC3-independent lysosomal targeting pathway. This mechanism constitutes a cell-death checkpoint that prevents TNF-induced apoptosis and TNFR1-mediated embryonic lethality and inflammatory skin disease in mouse models. ATG9A KO mice and cells, TNF complex IIa isolation, lysosomal targeting assays, mouse model of dermatitis and embryonic lethality Science (New York, N.Y.) High 36520901
2025 ATG9A facilitates autophagosome closure (the final stage of autophagosome biogenesis) by partnering with IQGAP1 and the ESCRT-III component CHMP2A. This function is distinct from ATG9A's role in phagophore initiation and expansion, making ATG9A a central hub for all stages of autophagosome membrane biogenesis. Co-immunoprecipitation of ATG9A–IQGAP1–CHMP2A, in vitro closure assay (SolVit), ATG9A KO reconstitution, high-content microscopy The Journal of cell biology High 39745851
2016 ATG9A trafficking through recycling endosomes is an essential step for autophagosome formation. Sorting motifs in the N-terminal cytosolic stretch of ATG9A interact with the adaptor protein AP-2; mutations in these motifs block autophagy and cause abnormal ATG9A accumulation at recycling endosomes. TRAPPC8 knockdown similarly causes ATG9A accumulation at recycling endosomes. AP-2 co-immunoprecipitation, site-directed mutagenesis of sorting motifs, TRAPPC8 siRNA knockdown, immunofluorescence localization, autophagy flux assays Journal of cell science High 27587839
2018 SNX18 regulates ATG9A trafficking from recycling endosomes by recruiting Dynamin-2. ATG9A is recruited to SNX18-induced tubules from recycling endosomes. SNX18 binding to Dynamin-2 is required for ATG9A trafficking from recycling endosomes and for formation of ATG16L1- and WIPI2-positive autophagosome precursor membranes. SNX18 KO cells, immunofluorescence of ATG9A-positive tubules, SNX18–Dynamin-2 Co-IP, autophagosome precursor membrane assays EMBO reports Medium 29437695
2020 The FHF (FTS-Hook-FHIP) complex interacts with AP-4 through direct binding of AP-4 μ4 subunit to coiled-coil domains in Hook1 and Hook2. FHF knockdown disperses AP-4 and ATG9A from the perinuclear region, indicating FHF couples AP-4/ATG9A vesicles to the microtubule retrograde motor dynein-dynactin for perinuclear distribution. Affinity purification-mass spectrometry, direct binding assay (μ4–Hook interaction), FHF subunit knockdown + ATG9A localization Molecular biology of the cell Medium 32073997
2021 RUSC2 couples ATG9A-containing vesicles to the plus-end-directed microtubule motor kinesin-1 via an interaction between a disordered region of RUSC2 and kinesin-1 light chain, driving peripheral distribution. The microtubule-associated protein WDR47 counteracts this interaction and negatively regulates peripheral ATG9A distribution. Kinesin-1 Co-IP with RUSC2, RUSC2/WDR47 knockout + rescue with truncation mutants, ATG9A localization by immunofluorescence Molecular biology of the cell Medium 34432492
2022 TRAF6 E3 ubiquitin ligase mediates K48/K63-linked non-proteolytic ubiquitination of ATG9A in response to ROS. This ubiquitination enhances ATG9A's association with Beclin 1 and promotes assembly of the VPS34-UVRAG complex, stimulating autophagy. The deubiquitinase A20 reverses this modification. ATG9A ubiquitination mutants impair ROS-induced VPS34 activation and autophagy. Co-immunoprecipitation of TRAF6–ATG9A, ubiquitin chain linkage analysis, ATG9A ubiquitination site mutagenesis, VPS34 activity assay Cell reports Medium 35196483
2022 Upon heat stress, the E3 ubiquitin ligase MARCH9 ubiquitinates ATG9A via K63 linkage. The non-degradable ubiquitinated ATG9A disperses from the Golgi to the cytoplasm and inhibits GRASP55 oligomerization, causing Golgi fragmentation. Knockout of ATG9A or MARCH9 prevents Golgi fragmentation under heat stress, revealing a noncanonical role for ATG9A in Golgi dynamics. MARCH9 KO and ATG9A KO cells, K63 ubiquitin chain analysis, GRASP55 oligomerization assay, Golgi morphology imaging Cell reports Medium 35977480
2021 BioID proximity proteomics identified ATG9A interactions with the ULK1 complex (particularly ATG13 and ATG101), as well as TRAPP, EARP, GARP, exocyst, AP-1, and AP-4 complexes. ATG9A interacts with the ATG13–ATG101 dimer independently of ULK1. Deletion of ATG13 or ATG101 shifts ATG9A distribution, causing accumulation at stalled p62/SQSTM1–ubiquitin clusters. Quantitative BioID proteomics, split-mVenus ATG13–ATG101 dimer capture, ATG13/ATG101 knockout + ATG9A localization EMBO reports Medium 34369648
2017 Rab1B, a small GTPase essential for ER-to-Golgi vesicle trafficking, is associated with ATG9A vesicles. Knockdown of Rab1B suppresses autophagy and causes ATG9A to accumulate in intermediate membrane structures at autophagosome formation sites, indicating Rab1B regulates proper development of autophagosomes through ATG9A vesicle dynamics. Immunoisolation of ATG9A vesicles + proteomics, Rab1B siRNA knockdown, ATG9A localization by immunofluorescence FASEB journal Medium 28522593
2022 ATG9A plays a critical role in chemotactic cell migration. ATG9A-positive vesicles are targeted toward the migration front where their exocytosis correlates with protrusive activity (TIRF live-cell imaging). ATG9A controls delivery of β1 integrin to the leading edge and regulates adhesion dynamics through interaction with clathrin adaptor complexes. ATG9A depletion + directional migration assays, ATG9A-pHluorin TIRF live imaging, β1 integrin delivery assay, Co-IP with clathrin adaptors The Journal of cell biology Medium 35180289
2021 BECN2 interacts with inflammasome sensors and mediates their degradation through a ULK1- and ATG9A-dependent but BECN1-WIPI2-ATG16L1-LC3-independent non-canonical autophagic pathway. BECN2 recruits inflammasome sensors on ATG9A+ vesicles to form a BECN2–ATG9A–sensor complex upon ULK1 activation, and SNARE proteins SEC22A, STX5, and STX6 mediate subsequent degradation. BECN2 Co-IP with ATG9A and inflammasome sensors, ATG9A KO cells, SNARE knockdown, NLRP3/AIM2/NLRP1/NLRC4 activity assays Autophagy Medium 34152938
2025 ATG9A-containing vesicles deliver PI4K2A to damaged lysosomes, cooperating with ARFIP2 to control PI4P levels for lysosomal repair. ARFIP2, a component of ATG9A vesicles, binds and sequesters PI4P on lysosomes, balancing OSBPL-dependent lipid transfer, and promotes retrieval of ATG9A vesicles by recruiting AP-3. ATG9A KO cells, lysosomal damage assays (chemical + bacterial), Co-IP of PI4K2A with ATG9A, ARFIP2–PI4P binding assay, AP-3 recruitment assay Developmental cell Medium 40460835
2024 ATG9A directly interacts with VPS13A, forming a complex distinct from the ATG9A–ATG2A complex, revealed by interactome mass spectrometry analysis. The ATG9A interactome is also enriched for proteins involved in lipid synthesis and trafficking including ACSL3 and VPS13C. Immunoprecipitation-mass spectrometry interactome, validation by Co-IP, biochemical distinction from ATG9A–ATG2A complex Journal of cell science Medium 38294121
2020 A heterodimeric SNX4–SNX7 complex coordinates ATG9A trafficking within the endocytic network. SNX4 partially co-localizes with juxtanuclear ATG9A-positive membranes, and SNX4 disruption causes mis-trafficking/retention of ATG9A in the Golgi region, impairing early autophagosome assembly. siRNA and CRISPR-Cas9 KO of SNX4/SNX7, ATG9A localization by immunofluorescence, LC3 lipidation assay, image-based early autophagosome assembly analysis Journal of cell science Medium 32513819
2021 SNX4 mediates recycling of ATG9A (the lipid scramblase) from endolysosomes to early endosomes, from where ATG9A is recycled to the TGN in a retromer-dependent manner. SNX4 or retromer (VPS35) depletion causes ATG9A accumulation on endolysosomes or early endosomes respectively, and impairs starvation-induced autophagosome biogenesis. siRNA knockdown of SNX4 and VPS35, ATG9A localization by immunofluorescence, autophagic flux assays Journal of cell science Medium 33468622
2018 VAMP7 localizes in ATG9A-resident vesicles of recycling endosomes and interacts with Hrb, Syntaxin16, and SNAP-47. Hrb recruits VAMP7 and ATG9A from the plasma membrane to recycling endosomes. VAMP7 forms a SNARE complex with Syntaxin16 and SNAP-47 that mediates fusion of ATG9A-resident vesicles during autophagosome formation. Co-immunoprecipitation of VAMP7–ATG9A–Hrb–Syntaxin16–SNAP-47, VAMP7-KO β-cells, immunofluorescence co-localization, autophagosome formation assays Endocrinology Medium 30215699
2020 ATG9A has a non-autophagic role in Golgi integrity and works in concert with GRASP55. ATG9A is not merely a passenger in the TGN but actively participates in Golgi dynamics (elaborated in follow-up MARCH9/ATG9A axis paper). ATG9A KO, Golgi morphology assays Autophagy Low 36198086
2016 A conserved glycine residue (G516) in the C-terminal region of human ATG9A (within the V515–C519 motif) is required for efficient transport of ATG9A from the ER to the Golgi apparatus. Human ATG9A self-interacts, and this self-interaction promotes transport through biosynthetic compartments, but via a mechanism distinct from yeast Atg9. Sequential mutagenesis of V515–C519, immunofluorescence localization, co-immunoprecipitation of ATG9A self-interaction Biochemical and biophysical research communications Medium 27663665
2016 Specific intramolecular sequences in ATG9A control its subcellular trafficking: deletion of L340–L354 causes ER retention; mutation of the L711YM713 sequence in the C-terminal region impairs Golgi transport. ATG9A oligomerization with co-expressed wild-type ATG9A can correct this Golgi transport defect, and some ATG9A can bypass the Golgi to reach the plasma membrane directly. Sequential deletion and point mutation analysis, immunofluorescence localization in HeLa cells Biochimica et biophysica acta Medium 27316455
2018 In AP-4 ε knockout mice, ATG9A is retained in the TGN and depleted from axons. Axonal ATG9A depletion leads to defective autophagosome generation and axonal swellings with accumulated ER. This mislocalization is also observed in human patient fibroblasts with AP-4 μ4 subunit mutations, linking AP-4-mediated ATG9A sorting to axonal integrity. AP-4 ε KO mice, immunohistochemistry and immunofluorescence in neurons and fibroblasts, electron microscopy of axonal swellings, mutant huntingtin aggregate assay PLoS genetics High 29698489
2020 ATG9A acetylation within the ER lumen regulates induction of reticulophagy. Although acetylation occurs in the ER lumen, reticulophagy induction requires ATG9A to engage reticulophagy receptors FAM134B and SEC62 on the cytosolic side of the ER. Interactome analysis in AT-1 dysregulation models identified CALR and HSPB1 as ATG9A partners that regulate reticulophagy as a function of ATG9A acetylation. AT-1 transgenic mouse models (overexpression + haploinsufficiency), ATG9A interactome by AP-MS, reticulophagy assays, Co-IP of ATG9A–FAM134B/SEC62 iScience Medium 33870132
2020 ATG9A regulates epithelial barrier function and tight junction biogenesis. HIF-1 binds the ATG9A promoter and induces its expression under hypoxia. ATG9A knockdown prevents epithelial barrier formation by >95%, mislocalizes multiple tight junction proteins, and causes intrinsic abnormalities in actin cytoskeleton including mislocalization of vasodilator-stimulated phosphoprotein. ChIP-chip identifying HIF-1 binding to ATG9A promoter, luciferase promoter assay, lentiviral ATG9A KD, transepithelial resistance measurement, tight junction protein immunofluorescence, actin cytoskeleton imaging Molecular biology of the cell Medium 32726170
2016 Excess sphingomyelin (due to SMPD1 deficiency) traps ATG9A in juxtanuclear recycling endosomes, preventing its normal trafficking to autophagic membranes. This ATG9A mistrafficking causes accumulation of elongated unclosed autophagic membranes. Ectopic ATG9A expression reverts the autophagy defect, and exogenous C12-sphingomyelin induces the same ATG9A mislocalization in healthy cells. SMPD1 KD cells and Niemann-Pick patient fibroblasts, ATG9A localization by immunofluorescence, ectopic ATG9A rescue, C12-sphingomyelin treatment, smpd1 KO mouse tissue Autophagy Medium 27070082
2016 Atg9a-deficient conditional knockout neurons show accumulation of SQSTM1/p62, NBR1, and ubiquitin, confirming autophagy block. Notably, Atg9a deficiency also impairs neurite outgrowth independently of autophagy (not phenocopied by Atg7 or Atg16l1 KO), and causes dysgenesis of corpus callosum and anterior commissure detected by diffusion tensor MRI. Brain-specific conditional Atg9a KO mice, immunohistochemistry, electron microscopy, diffusion tensor MRI, primary neuron culture neurite outgrowth assay, Atg7-KO/Atg16l1-KO comparison Autophagy High 28513333
2016 Atg9a-dependent necrosis occurs at the bone surface during developmental bone formation in mice. This form of necrosis is not observed in Atg5 knockout embryos, establishing Atg9a-specific (autophagy-independent) function in developmental morphogenesis through programmed necrosis. Atg9a knockout mice display abnormalities of the bone surface. Propidium iodide vital staining in mouse embryos, Atg9a KO and Atg5 KO comparison, bone surface morphology analysis Nature communications Medium 27811852
2024 RHOD (an atypical Rho GTPase) interacts with ATG9A upon starvation and accompanies ATG9A trafficking from the Golgi toward phagophores. RHOD promotes Golgi fragmentation to enhance ATG9A vesicle export from the TGN. WHAMM forms a complex with RHOD and participates in this process in a RHOD-dependent manner. RHOD mutants lacking ATG9A-binding or membrane-targeting domains fail to stimulate ATG9A trafficking. Co-immunoprecipitation of RHOD–ATG9A, bimolecular fluorescence complementation, PUP-IT interaction tagging, RHOD KO cells + ATG9A localization, RHOD mutant rescue experiments Autophagy Medium 40143438
2024 IRP2, but not IRP1, is indispensable for ferritinophagy. IRP2 ablation results in defective ATG9A endosomal trafficking (dependent on compromised AMPK activation), leading to decreased engulfment of the NCOA4-ferritin complex by endosomes and dysregulated endosomal microferritinophagy. IRP2 KO cells, ATG9A endosomal localization by immunofluorescence, ferritinophagy flux assay, AMPK activation analysis The Journal of biological chemistry Medium 39276939
2025 ATG9A vesicles serve as specialized carriers for galectin-9 secretion via unconventional secretion independent of classical autophagy, secretory autophagy, or LC3-dependent extracellular vesicle secretion. The N-terminus of ATG9A and both carbohydrate recognition domains of galectin-9 are critical for the process. TMED10 mediates galectin-9 incorporation into ATG9A vesicles, which fuse with the plasma membrane via the STX13-SNAP23-VAMP3 SNARE complex. ATG9A KO cells, galectin-9 secretion assay, ATG9A N-terminus mutagenesis, Co-IP of galectin-9 with ATG9A, TMED10 KD, SNARE complex identification Nature communications Medium 40335523
2025 ATG9A undergoes S-palmitoylation, a lipid modification that coordinates its trafficking to mediate autophagy initiation. Palmitoylation-deficient ATG9A (C155S/C156S double mutant) shows altered subcellular distribution and impaired autophagy initiation. Acyl-biotin exchange assay, APEX2 proximity labeling, mutagenesis of palmitoylation sites, ATG9A trafficking and autophagy assays Autophagy Medium 40394978
2024 SEC31a (outer coat protein of COPII vesicles) interacts with ATG9A on autophagosomal seed vesicles, mediating recruitment of COPII vesicles as a membrane source for autophagosome formation during osteogenic differentiation of mesenchymal stem cells. Co-immunoprecipitation of SEC31a–ATG9A, SEC31a siRNA disruption + autophagosome number/size analysis, in vivo bone tissue analysis Advanced science Low 39361436
2035 ATG9A vesicles are a subtype of intracellular nanovesicles (INVs). Virtually all ATG9A vesicles are INVs, but only ~20% of INVs are ATG9A vesicles. ATG9A vesicles carry other ATG9A vesicle cargoes as confirmed by in-cell vesicle capture assays. Perturbing ATG9A-flavor INVs impairs starvation-induced autophagy. In-cell vesicle capture assay, quantitative imaging of ATG9A/synaptophysin co-localization, INV proteomics Journal of cell science Low 40067248
2023 ATG9A supports Chlamydia trachomatis inclusion growth via autophagy-independent mechanisms; rescue experiments with ATG9A mutants revealed this function is mediated not by scramblase/autophagic activity but by ATG9A's binding ability to clathrin adaptor proteins. ATG9A KO HeLa cells, C. trachomatis growth assay, rescue with ATG9A mutants (scramblase-dead vs clathrin adaptor-binding mutants) Microbiology spectrum Medium 37707289
2024 ATG9A vesicles and synaptophysin-positive synaptic vesicles segregate into distinct phases within synapsin condensates at synapses. In fibroblasts and nerve terminals, ATG9A localizes on a distinct class of vesicles from synaptophysin-positive vesicles, indicating differential sorting mechanisms. ATG9A undergoes activity-dependent exo-endocytosis at synapses. Ectopic expression in fibroblasts, live-cell imaging of synapsin condensates, nerve terminal immunofluorescence, fluorescence correlation with synapsin/synaptophysin Nature communications Medium 36709207
2022 miR-34a directly binds the 3'-UTR of ATG9A mRNA and inhibits ATG9A protein expression, thereby suppressing autophagic activity. In angiotensin II-induced cardiomyocyte hypertrophy, downregulation of miR-34a increases ATG9A expression and autophagic activity, which promotes cardiomyocyte hypertrophy. This places ATG9A downstream of miR-34a in the hypertrophy pathway. miR-34a 3'-UTR luciferase reporter assay, ATG9A overexpression/knockdown in cardiomyocytes, LC3 II/I and p62 western blot, TEM of autophagosomes PloS one Medium 24728149
2024 ULK1 phosphorylates ATG9A at S14, and ATG9A interacts with ULK1 at S467 of ULK1. ATG9A functions as a superior signaling node upstream of Rab9 in the ULK1/Atg9a/Rab9 signaling pathway regulating inflammasome activation, Golgi fragmentation, and mitochondrial oxidative stress in asthma. ULK1 KO cells and mice, lentiviral ULK1 WT and S467A reconstitution, Co-IP of ULK1–ATG9A, phosphorylation site mapping Redox biology Low 38373380
2023 ATG9A directly interacts with the HORMA dimer of ATG13 and ATG101 independently of ULK1. An ULK1-independent ATG13 complex (ATG13–ATG101 dimer) promotes autophagy in fed cells, and this is regulated through ATG9A interaction. BioID, split-mVenus ATG13–ATG101 capture, Co-IP, ATG13/ATG101 KO + ATG9A localization, autophagy flux in fed conditions Autophagy Medium 35442099
2023 ATG9A vesicles are present at the transition from a seed vesicle to a mature autophagosome; live-cell imaging tracking shows ATG9A vesicles at initiation sites prior to LC3B/GABARAPL1 recruitment, and the ATG9A-positive structure persists through early autophagosome assembly. Live-cell multi-color fluorescence microscopy in U2OS cells tracking ATG9A, ATG13, WIPI2, LC3B, GABARAPL1, SQSTM1 dynamics Autophagy Low 37405380

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2009 Atg9a controls dsDNA-driven dynamic translocation of STING and the innate immune response. Proceedings of the National Academy of Sciences of the United States of America 698 19926846
2020 Structure, lipid scrambling activity and role in autophagosome formation of ATG9A. Nature structural & molecular biology 244 33106659
2020 Critical role of mitochondrial ubiquitination and the OPTN-ATG9A axis in mitophagy. The Journal of cell biology 161 32556086
2019 ATG9A shapes the forming autophagosome through Arfaptin 2 and phosphatidylinositol 4-kinase IIIβ. The Journal of cell biology 154 30917996
2017 AP-4 mediates export of ATG9A from the trans-Golgi network to promote autophagosome formation. Proceedings of the National Academy of Sciences of the United States of America 139 29180427
2020 Structure of Human ATG9A, the Only Transmembrane Protein of the Core Autophagy Machinery. Cell reports 137 32610138
2016 Atg9A trafficking through the recycling endosomes is required for autophagosome formation. Journal of cell science 132 27587839
2018 AP-4 vesicles contribute to spatial control of autophagy via RUSC-dependent peripheral delivery of ATG9A. Nature communications 127 30262884
2022 ATG9A and ATG2A form a heteromeric complex essential for autophagosome formation. Molecular cell 108 36347259
2021 The autophagy protein ATG9A enables lipid mobilization from lipid droplets. Nature communications 101 34799570
2018 Altered distribution of ATG9A and accumulation of axonal aggregates in neurons from a mouse model of AP-4 deficiency syndrome. PLoS genetics 95 29698489
2018 Atg9a deficiency causes axon-specific lesions including neuronal circuit dysgenesis. Autophagy 87 28513333
2018 SNX18 regulates ATG9A trafficking from recycling endosomes by recruiting Dynamin-2. EMBO reports 77 29437695
2018 ATG7 and ATG9A loss-of-function variants trigger autophagy impairment and ovarian failure. Genetics in medicine : official journal of the American College of Medical Genetics 70 30224786
2019 Axonal autophagosome maturation defect through failure of ATG9A sorting underpins pathology in AP-4 deficiency syndrome. Autophagy 69 31142229
2021 ATG9A protects the plasma membrane from programmed and incidental permeabilization. Nature cell biology 68 34257406
2014 miR-34a modulates angiotensin II-induced myocardial hypertrophy by direct inhibition of ATG9A expression and autophagic activity. PloS one 66 24728149
2017 Activation of miR-34a impairs autophagic flux and promotes cochlear cell death via repressing ATG9A: implications for age-related hearing loss. Cell death & disease 65 28981097
2022 K48/K63-linked polyubiquitination of ATG9A by TRAF6 E3 ligase regulates oxidative stress-induced autophagy. Cell reports 64 35196483
2019 The lncRNA NEAT1/miR-29b/Atg9a axis regulates IGFBPrP1-induced autophagy and activation of mouse hepatic stellate cells. Life sciences 60 31610195
2017 Small GTPase Rab1B is associated with ATG9A vesicles and regulates autophagosome formation. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 56 28522593
2016 Excess sphingomyelin disturbs ATG9A trafficking and autophagosome closure. Autophagy 52 27070082
2023 Structural basis for ATG9A recruitment to the ULK1 complex in mitophagy initiation. Science advances 47 36791199
2021 BioID reveals an ATG9A interaction with ATG13-ATG101 in the degradation of p62/SQSTM1-ubiquitin clusters. EMBO reports 42 34369648
2023 Synaptic vesicle proteins and ATG9A self-organize in distinct vesicle phases within synapsin condensates. Nature communications 40 36709207
2010 Atg9A protein, an autophagy-related membrane protein, is localized in the neurons of mouse brains. The journal of histochemistry and cytochemistry : official journal of the Histochemistry Society 40 20124090
2020 Two different axes CALCOCO2-RB1CC1 and OPTN-ATG9A initiate PRKN-mediated mitophagy. Autophagy 38 32892694
2020 A heterodimeric SNX4--SNX7 SNX-BAR autophagy complex coordinates ATG9A trafficking for efficient autophagosome assembly. Journal of cell science 37 32513819
2015 Role of the Atg9a gene in intrauterine growth and survival of fetal mice. Reproductive biology 36 26370455
2014 Age-enhanced endoplasmic reticulum stress contributes to increased Atg9A inhibition of STING-mediated IFN-β production during Streptococcus pneumoniae infection. Journal of immunology (Baltimore, Md. : 1950) 36 24670807
2016 Aberrant methylation of ATG2B, ATG4D, ATG9A and ATG9B CpG island promoter is associated with decreased mRNA expression in sporadic breast carcinoma. Gene 35 27265029
2022 ATG9A prevents TNF cytotoxicity by an unconventional lysosomal targeting pathway. Science (New York, N.Y.) 34 36520901
2021 The phosphatidylinositol 3-phosphate-binding protein SNX4 controls ATG9A recycling and autophagy. Journal of cell science 33 33468622
2020 HANR Enhances Autophagy-Associated Sorafenib Resistance Through miR-29b/ATG9A Axis in Hepatocellular Carcinoma. OncoTargets and therapy 30 32210579
2020 NMMHC IIA triggers neuronal autophagic cell death by promoting F-actin-dependent ATG9A trafficking in cerebral ischemia/reperfusion. Cell death & disease 29 32513915
2020 The FTS-Hook-FHIP (FHF) complex interacts with AP-4 to mediate perinuclear distribution of AP-4 and its cargo ATG9A. Molecular biology of the cell 27 32073997
2018 VAMP7 Regulates Autophagosome Formation by Supporting Atg9a Functions in Pancreatic β-Cells From Male Mice. Endocrinology 27 30215699
2024 Structural basis for lipid transfer by the ATG2A-ATG9A complex. Nature structural & molecular biology 26 39174844
2021 BECN2 (beclin 2) Negatively Regulates Inflammasome Sensors Through ATG9A-Dependent but ATG16L1- and LC3-Independent Non-Canonical Autophagy. Autophagy 26 34152938
2022 Nondegradable ubiquitinated ATG9A organizes Golgi integrity and dynamics upon stresses. Cell reports 25 35977480
2018 ATG9A Is Overexpressed in Triple Negative Breast Cancer and Its In Vitro Extinction Leads to the Inhibition of Pro-Cancer Phenotypes. Cells 25 30563263
2017 TMEM74 promotes tumor cell survival by inducing autophagy via interactions with ATG16L1 and ATG9A. Cell death & disease 23 29048433
2016 ATG9A loss confers resistance to trastuzumab via c-Cbl mediated Her2 degradation. Oncotarget 23 27050377
2020 The HIF target ATG9A is essential for epithelial barrier function and tight junction biogenesis. Molecular biology of the cell 22 32726170
2016 Vital staining for cell death identifies Atg9a-dependent necrosis in developmental bone formation in mouse. Nature communications 22 27811852
2014 Lentivirus-mediated Bos taurus bta-miR-29b overexpression interferes with bovine viral diarrhoea virus replication and viral infection-related autophagy by directly targeting ATG14 and ATG9A in Madin-Darby bovine kidney cells. The Journal of general virology 22 25234643
2022 The core autophagy protein ATG9A controls dynamics of cell protrusions and directed migration. The Journal of cell biology 21 35180289
2024 NDRG1 overcomes resistance to immunotherapy of pancreatic ductal adenocarcinoma through inhibiting ATG9A-dependent degradation of MHC-1. Drug resistance updates : reviews and commentaries in antimicrobial and anticancer chemotherapy 20 38228036
2024 IL-4 activates ULK1/Atg9a/Rab9 in asthma, NLRP3 inflammasomes, and Golgi fragmentation by increasing autophagy flux and mitochondrial oxidative stress. Redox biology 19 38373380
2024 Emerging roles of ATG9/ATG9A in autophagy: implications for cell and neurobiology. Autophagy 19 39099167
2021 RUSC2 and WDR47 oppositely regulate kinesin-1-dependent distribution of ATG9A to the cell periphery. Molecular biology of the cell 18 34432492
2025 ATG9A and ARFIP2 cooperate to control PI4P levels for lysosomal repair. Developmental cell 17 40460835
2024 Exploring the ATG9A interactome uncovers interaction with VPS13A. Journal of cell science 17 38294121
2018 Deregulation of ATG9A by impaired AR signaling induces autophagy in prostate stromal fibroblasts and promotes BPH progression. Cell death & disease 17 29568063
2022 miR-34a/ATG9A/TFEB Signaling Modulates Autophagy in Cochlear Hair Cells and Correlates with Age-related Hearing Loss. Neuroscience 16 35367291
2019 The autophagy protein ATG9A promotes HIV-1 infectivity. Retrovirology 16 31269971
2021 High-throughput imaging of ATG9A distribution as a diagnostic functional assay for adaptor protein complex 4-associated hereditary spastic paraplegia. Brain communications 15 34729478
2025 ATG9A facilitates the closure of mammalian autophagosomes. The Journal of cell biology 14 39745851
2021 ATG9A regulates proteostasis through reticulophagy receptors FAM134B and SEC62 and folding chaperones CALR and HSPB1. iScience 14 33870132
2018 MiR-29a inhibited intestinal epithelial cells autophagy partly by decreasing ATG9A in ulcerative colitis. Anti-cancer drugs 14 29916896
2022 Trehalose Protects Keratinocytes against Ultraviolet B Radiation by Activating Autophagy via Regulating TIMP3 and ATG9A. Oxidative medicine and cellular longevity 13 35450405
2023 Ap4b1-knockout mouse model of hereditary spastic paraplegia type 47 displays motor dysfunction, aberrant brain morphology and ATG9A mislocalization. Brain communications 12 36632189
2023 circAP1M2 activates ATG9A-associated autophagy by inhibiting miR-1249-3p to promote cisplatin resistance in oral squamous cell carcinoma. Journal of cellular physiology 12 37661341
2023 ATG9A modulated by miR-195-5p can boost the malignant progression of cervical cancer cells. Epigenetics 10 37782756
2023 ATG9B is a tissue-specific homotrimeric lipid scramblase that can compensate for ATG9A. Autophagy 10 37938170
2025 YBX1 promotes 5-Fluorouracil resistance in gastric cancer via m5C-dependent ATG9A mRNA stabilization through autophagy. Oncogene 9 40251390
2022 Autophagy-associated immune dysregulation and hyperplasia in a patient with compound heterozygous mutations in ATG9A. Autophagy 9 35838483
2024 SEC31a-ATG9a Interaction Mediates the Recruitment of COPII Vesicles for Autophagosome Formation. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 8 39361436
2023 S100a9 inhibits Atg9a transcription and participates in suppression of autophagy in cardiomyocytes induced by β1-adrenoceptor autoantibodies. Cellular & molecular biology letters 8 37723445
2019 ATG9A supplies PtdIns4P to the autophagosome initiation site. Autophagy 8 31204568
2024 Iron regulatory protein two facilitates ferritinophagy and DNA damage/repair through guiding ATG9A trafficking. The Journal of biological chemistry 7 39276939
2022 Atg9A-mediated mitophagy is required for decidual differentiation of endometrial stromal cells. Reproductive biology 7 36343573
2020 The structure of human ATG9A and its interplay with the lipid bilayer. Autophagy 7 33016201
2023 HSF4 Transcriptionally Activates Autophagy by Regulating ATG9a During Lens Terminal Differentiation. Investigative ophthalmology & visual science 6 37266953
2023 Tracking the transition from an ATG9A vesicle to an autophagosome. Autophagy 6 37405380
2022 A noncanonical autophagy function of ATG9A for Golgi integrity and dynamics. Autophagy 6 36198086
2024 miRNA family miR-29 inhibits PINK1-PRKN dependent mitophagy via ATG9A. bioRxiv : the preprint server for biology 5 38293184
2024 Tepsin binds LC3B to promote ATG9A trafficking and delivery. Molecular biology of the cell 5 38381558
2016 A conserved glycine residue in the C-terminal region of human ATG9A is required for its transport from the endoplasmic reticulum to the Golgi apparatus. Biochemical and biophysical research communications 5 27663665
2025 Autophagy-independent role of ATG9A vesicles as carriers for galectin-9 secretion. Nature communications 4 40335523
2025 S-palmitoylation coordinates the trafficking of ATG9A to mediate autophagy initiation. Autophagy 4 40394978
2022 Mapping the proximity interactome of ATG9A reveals unexpected dynamics of ULK1 complex proteins. Autophagy 4 35442099
2022 Chemotactic cell migration: the core autophagy protein ATG9A is at the leading edge. Autophagy 4 35468023
2016 Molecular determinants that mediate the sorting of human ATG9A from the endoplasmic reticulum. Biochimica et biophysica acta 4 27316455
2025 ATG9A vesicles are a subtype of intracellular nanovesicle. Journal of cell science 3 40067248
2025 Anoctamin-5 deficiency enhances ATG9A-dependent autophagy, inducing osteogenesis and gnathodiaphyseal dysplasia-like bone formation. JCI insight 3 40067389
2025 Progress on multifunctional transmembrane protein ATG9A. Cell communication and signaling : CCS 3 40598533
2024 HCMV miR-UL70-3p downregulates the rapamycin-induced autophagy by targeting the autophagy-related protein 9A (ATG9A). International reviews of immunology 3 38164951
2023 A novel role of ATG9A and RB1CC1/FIP200 in mediating cell-death checkpoints to repress TNF cytotoxicity. Autophagy 3 36892222
2023 ATG9A supports Chlamydia trachomatis infection via autophagy-independent mechanisms. Microbiology spectrum 3 37707289
2021 Downregulation of microRNA-96-5p protects TM3 cells against zearalenone toxicity via targeting ATG9A. Experimental and therapeutic medicine 3 34584554
2025 ATG2A-WDR45/WIPI4-ATG9A complex-mediated lipid transfer and equilibration during autophagosome formation. Autophagy 2 40116844
2025 ATG9A controls all stages of autophagosome biogenesis. Autophagy 2 40241347
2025 ATG9A-mediated autophagy prevents inflammatory skin disease by limiting TNFR1-driven STING activation and ZBP1-dependent cell death. Immunity 2 41118755
2024 The differential expression patterns of Atg9a and Atg9b in cells of the reproductive organs. Clinical and experimental reproductive medicine 2 38757275
2024 Ubiquitin-mediated recruitment of the ATG9A-ATG2 lipid transfer complex drives clearance of phosphorylated p62 aggregates. Molecular biology of the cell 2 39718773
2023 Imaging ATG9A, a Multi-Spanning Membrane Protein. Journal of visualized experiments : JoVE 2 37395569
2023 Tepsin binds LC3B to promote ATG9A export and delivery at the cell periphery. bioRxiv : the preprint server for biology 2 37502979
2025 RHOD mediates ATG9A trafficking to promote autophagosome formation during autophagy in cancer. Autophagy 1 40143438
2024 Absence of ATG9A and synaptophysin demixing on Rab5 mutation-induced giant endosomes. Molecular brain 1 39223639