Affinage

GABARAPL2

Gamma-aminobutyric acid receptor-associated protein-like 2 · UniProt P60520

Length
117 aa
Mass
13.7 kDa
Annotated
2026-06-09
100 papers in source corpus 25 papers cited in narrative 25 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

GABARAPL2 (GATE-16) is a ubiquitin-like ATG8-family protein that functions as a membrane-conjugated adaptor coordinating autophagosome biogenesis, membrane fusion, and selective cargo capture (PMID:10856287, PMID:21497758). Following C-terminal processing, it is lipidated to a phosphatidylethanolamine-conjugated form (form II) that associates with autophagosomal membranes (PMID:15169837), and during non-canonical autophagy (CASM) it is alternatively conjugated to phosphatidylserine, which serves as a molecular signature distinguishing this pathway and is differentially delipidated by ATG4 enzymes (PMID:33909989). Its two N-terminal α-helices drive membrane tethering and fusion through hydrophobic interactions—a 10-residue N-terminal peptide is sufficient to fuse membranes in vitro and these residues are essential for autophagosome biogenesis—and tethering occurs by reversible trans-assembly that is curvature-sensitive, favoring larger/flatter vesicles in contrast to the small-curvature preference of LC3B (PMID:21497758, PMID:31960529). GABARAPL2 recruits autophagy receptors and adaptors, including p62/SQSTM1, through LIR–LDS interactions whose selectivity is dictated by subfamily-specific residues in the LIR docking site (PMID:17580304, PMID:31053714). Together with ESCRT machinery it maintains autophagosomal membranes in a sealed state required for maturation into autolysosomes (PMID:37272163), and via SNARE interactions it contributes to autophagosome–lysosome fusion and lysosome biogenesis (PMID:31625181). Beyond autophagy, GABARAPL2 acts in innate immunity: it cooperates with IRGM2 to dampen caspase-11 non-canonical inflammasome activation by limiting GBP2-dependent targeting of intracellular bacteria, and Gate-16/Gabarap double-deficient mice die after low-dose LPS in a GBP2-dependent manner (PMID:33124769, PMID:33042141); it is also required for IFN-γ–mediated restriction of Toxoplasma gondii, localizing to parasitophorous vacuole membranes (PMID:32094251). Its endogenous interactome links it to ER lipid-droplet biology and ufmylation via ACSL3, which anchors the UFM1-activating enzyme UBA5 at the ER (PMID:32843575), to transcriptional control of autophagy through LMX1B in dopaminergic neurons (PMID:37014324), and to regulation of the TNF receptor Fn14 and TWEAK/NF-κB signaling (PMID:30218067).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 2000 High

    Established the structural basis for GATE-16's adaptor activity and its first functional role in membrane trafficking, framing it as a ubiquitin-fold protein with extra N-terminal helices that engages NSF and Golgi SNAREs.

    Evidence X-ray crystallography at 1.8 Å with biochemical interaction studies

    PMID:10856287

    Open questions at the time
    • Did not define how the N-terminal helices engage membranes
    • Golgi SNARE-priming role left mechanistically informal until later reviews
  2. 2004 High

    Showed GATE-16 is post-translationally lipidated like LC3 and GABARAP, placing it in the ATG8 conjugation system and on autophagosomal membranes.

    Evidence Subcellular fractionation, [14C]-ethanolamine metabolic labeling, and Atg4B deconjugation in mammalian cells

    PMID:15169837

    Open questions at the time
    • Did not resolve the functional role of lipidated GATE-16 in autophagosome formation
    • PE identity inferred from labeling/deconjugation rather than direct lipid structure
  3. 2007 High

    Identified the direct receptor-binding function by mapping p62/SQSTM1 binding to a LIR-containing motif, explaining how GABARAPL2 links ubiquitinated cargo to autophagy.

    Evidence Direct binding, reciprocal co-IP, and pH-sensitive tandem-tag imaging of aggregate degradation

    PMID:17580304

    Open questions at the time
    • Did not establish what distinguishes GABARAPL2 from other ATG8s in receptor selection
  4. 2011 High

    Defined the membrane fusion mechanism, showing the N-terminal α-helices are sufficient and necessary for tethering/fusion and autophagosome biogenesis.

    Evidence Cell-free fusion assays with synthetic N-terminal peptides plus cellular autophagosome biogenesis assays

    PMID:21497758

    Open questions at the time
    • Did not address membrane-curvature preference or how fusion is regulated in vivo
  5. 2011 Medium

    Linked GATE-16 to ORP7 and SNARE (GS28) stability control, extending its trafficking role beyond direct SNARE priming.

    Evidence Yeast two-hybrid, BiFC, truncation mapping, and protein stability assays

    PMID:21669198

    Open questions at the time
    • Relied on overexpression of ORP7
    • Physiological relevance of GS28 regulation not established
  6. 2013 Medium

    Identified GIMAP6 as a specific GABARAPL2 partner degraded during starvation, suggesting GABARAPL2-mediated selective autophagy of an immune GTPase.

    Evidence Biotin tag-affinity purification, chemical cross-linking, co-IP, and starvation degradation assays in Jurkat T cells

    PMID:24204963

    Open questions at the time
    • Single lab
    • Functional consequence of GIMAP6 degradation not defined
  7. 2013 Medium

    Demonstrated a cellular requirement for GABARAPL2 in autophagic flux and ATRA-induced myeloid differentiation.

    Evidence siRNA knockdown with differentiation and autophagic flux readouts in APL cells

    PMID:23891751

    Open questions at the time
    • Did not separate autophagy-dependent from autophagy-independent contributions to differentiation
  8. 2015 High

    Revealed a C-terminus-centered conformational polymorphism crucial for GABARAPL2 activity, connecting structural dynamics to function.

    Evidence X-ray crystallography, NMR, and molecular dynamics simulations

    PMID:26284781

    Open questions at the time
    • Did not link the conformational transition to a specific binding or lipidation step
  9. 2016 Medium

    Distinguished GABARAPL2 from LC3B functionally by showing it does not bind cardiolipin or translocate to mitochondria, implying it is not a primary mitophagy effector.

    Evidence Quantitative in vitro biophysical binding assays and mitochondrial translocation imaging

    PMID:27764541

    Open questions at the time
    • Negative result for a single condition (rotenone); other mitophagy contexts untested
  10. 2018 Medium

    Showed GABARAPL2 and GABARAP have non-redundant roles in handling the TNF receptor Fn14, with GABARAPL2 controlling endosomal Fn14 and TWEAK/NF-κB signaling.

    Evidence Knockout cell lines, microscopy, NF-κB reporter assays, and immunoprecipitation

    PMID:30218067

    Open questions at the time
    • Mechanism of Fn14 endosomal retention not resolved
    • Single lab
  11. 2019 High

    Defined the molecular basis of GABARAP-subfamily receptor selectivity, showing LIR-motif and LDS residues drive selective binding and that altering them impairs receptor degradation.

    Evidence In vitro binding, mutagenesis, cellular degradation assays, and live-cell PCM1 imaging

    PMID:31053714

    Open questions at the time
    • Did not enumerate the full GABARAPL2-specific receptor repertoire
  12. 2019 Medium

    Connected GABARAP-subfamily proteins to SNARE-dependent autophagosome-lysosome fusion and lysosome biogenesis via LIR-mediated Stx16 binding.

    Evidence LIR mapping, co-IP, knockout cell lines, and autophagic flux/lysosome biogenesis assays

    PMID:31625181

    Open questions at the time
    • GABARAPL2-specific contribution not isolated from other subfamily members
  13. 2020 High

    Identified ACSL3 as an endogenous stabilizing partner recruiting GABARAPL2 to the ER and anchoring UBA5, linking GABARAPL2 to ufmylation and lipid-droplet/ER-phagy biology.

    Evidence CRISPR endogenous tagging, IP-MS, LIR-site mutagenesis, fractionation, and siRNA depletion

    PMID:32843575

    Open questions at the time
    • Direct enzymatic role of GABARAPL2 in ufmylation not established
  14. 2020 High

    Established GABARAPL2 as a brake on the caspase-11 non-canonical inflammasome, acting with IRGM2 to limit GBP-dependent bacterial targeting, pyroptosis, and lethal endotoxemia.

    Evidence Knockout and double-knockout macrophages and mice, inflammasome/pyroptosis assays, and GBP2-deficiency rescue, replicated across two labs

    PMID:33042141 PMID:33124769

    Open questions at the time
    • Whether membrane lipidation of GABARAPL2 is required for inflammasome dampening not fully resolved
  15. 2020 High

    Defined GABARAPL2's curvature-sensitive tethering mechanism, distinguishing it from LC3B by its preference for larger/flatter vesicles via reversible trans-assembly.

    Evidence Cell-free reconstitution with purified lipidated proteins and curvature-defined synthetic liposomes

    PMID:31960529

    Open questions at the time
    • In vivo relevance of the curvature preference to specific fusion events untested
  16. 2020 Medium

    Showed GABARAPL2 is required for IFN-γ-mediated restriction of Toxoplasma gondii and localizes to parasitophorous vacuole membranes.

    Evidence Knockdown/knockout, microscopy, and parasite growth assays under IFN-γ stimulation

    PMID:32094251

    Open questions at the time
    • Molecular mechanism of vacuole targeting and parasite killing not defined
    • Single lab
  17. 2020 High

    Implicated GABARAPL2-subfamily proteins, alongside ESCRT-I, in maintaining sealed autophagosomal membranes required for autolysosome maturation.

    Evidence mATG8 knockout cell lines, a novel in vitro membrane sealing assay, ESCRT interaction studies, and flux assays

    PMID:37272163

    Open questions at the time
    • Direct ESCRT binding shown for GABARAP/LC3A; GABARAPL2-specific ESCRT contacts not separately mapped
  18. 2021 High

    Established phosphatidylserine conjugation of GABARAPL2 as a hallmark of non-canonical autophagy (CASM), distinct from canonical PE lipidation in dynamics and ATG4 processing.

    Evidence Lipidomics, multiple CASM inducers, ATG4 delipidation assays, and ATG16L1 WD40 mutants

    PMID:33909989

    Open questions at the time
    • Functional consequences specific to the PS-conjugated species not fully dissected
  19. 2022 Medium

    Linked GABARAPL2 to transcriptional control of autophagy by showing ATG8 binding stimulates LMX1B-mediated transcription and stress protection in dopaminergic neurons.

    Evidence Co-IP, live-cell imaging, iPSC-derived neuron differentiation, and transcription/toxicity assays

    PMID:37014324

    Open questions at the time
    • GABARAPL2-specific (vs other ATG8) contribution to LMX1B regulation not isolated

Open questions

Synthesis pass · forward-looking unresolved questions
  • How GABARAPL2's distinct biochemical properties (curvature preference, PS-lipidation, LDS selectivity) are deployed to assign it non-redundant roles versus other ATG8 family members across autophagy, immunity, and trafficking remains unresolved.
  • No unified model partitioning GABARAPL2 vs GABARAP/LC3 functions in vivo
  • Many findings inferred at the subfamily level rather than GABARAPL2 alone

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0008289 lipid binding 4 GO:0005198 structural molecule activity 3 GO:0060090 molecular adaptor activity 3 GO:0031386 protein tag activity 2
Localization
GO:0005764 lysosome 1 GO:0005768 endosome 1 GO:0005783 endoplasmic reticulum 1 GO:0005794 Golgi apparatus 1 GO:0005829 cytosol 1
Pathway
R-HSA-168256 Immune System 3 R-HSA-5653656 Vesicle-mediated transport 3 R-HSA-9612973 Autophagy 3 R-HSA-5357801 Programmed Cell Death 2
Partners
ACSL3GIMAP6IRGM2LMX1BNSFSQSTM1STX16TNFRSF12A

Evidence

Reading pass · 25 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2000 Crystal structure of GATE-16 (GABARAPL2) was solved at 1.8 Å resolution, revealing a ubiquitin fold decorated by two additional N-terminal helices. The structure suggests GATE-16 binds targets via pseudo-continuous beta-sheets similar to Ras effectors, and a second potential protein-protein interaction site may explain its adapter activity. GATE-16 was shown to associate with N-ethylmaleimide-sensitive fusion protein (NSF) and Golgi SNAREs, and participates in intra-Golgi transport. X-ray crystallography (1.8 Å); biochemical interaction studies The Journal of biological chemistry High 10856287
2004 GATE-16 (GABARAPL2), like LC3 and GABARAP, undergoes post-translational lipidation (form II generation) and the lipidated form associates with autophagosomal membranes. [14C]-ethanolamine incorporation and sensitivity to mammalian Atg4B support that form II is a phosphatidylethanolamine (PE)-conjugated species. Subcellular fractionation, [14C]-ethanolamine metabolic labeling, Atg4B deconjugation assay, fluorescence microscopy Journal of cell science High 15169837
2007 p62/SQSTM1 directly interacts with GABARAPL2 (GATE-16) and other mammalian ATG8 homologs (LC3A, LC3B, GABARAP) via a conserved 22-residue sequence containing the LIR motif, facilitating autophagic degradation of ubiquitinated protein aggregates. Direct binding assay, co-immunoprecipitation, fluorescence microscopy with pH-sensitive tandem tag The Journal of biological chemistry High 17580304
2011 GATE-16 (GABARAPL2) promotes membrane tethering and fusion via its N-terminal α-helices, mediated by hydrophobic interactions. A 10-amino acid peptide from the GATE-16 N-terminus is sufficient to promote membrane fusion in a cell-free system. These N-terminal residues are essential for autophagosome biogenesis in cells. Cell-free membrane fusion assay, synthetic N-terminal peptides, autophagosome biogenesis assay in cells Developmental cell High 21497758
2013 GIMAP6, a cytosolic GTPase expressed in immune cells, specifically interacts with GABARAPL2 (identified by biotin tag-affinity purification and chemical cross-linking in Jurkat T cells). The interaction requires the last 10 amino acids of GIMAP6 (not its AIM motif). Upon starvation, GIMAP6 co-localizes with GABARAPL2 and MAP1LC3B at autophagosomes and is degraded. GIMAP6 overexpression increases endogenous GABARAPL2 levels. Biotin tag-affinity purification, chemical cross-linking, co-immunoprecipitation, fluorescence microscopy, starvation-induced degradation assay PloS one Medium 24204963
2013 Knockdown of GABARAPL2/GATE-16 in acute promyelocytic leukemia (APL) cells attenuates ATRA-induced neutrophil differentiation and decreases autophagic flux, demonstrating a functional requirement for GATE-16 in myeloid differentiation and autophagosome formation. siRNA knockdown, differentiation assays, autophagic flux measurement Biochemical and biophysical research communications Medium 23891751
2015 Conformational polymorphism of GATE-16 (GABARAPL2) was characterized: X-ray crystallography, NMR spectroscopy, and molecular dynamics simulations revealed a structural transition centered on the C-terminus that is crucial for biological activity of the protein. X-ray crystallography, NMR spectroscopy, molecular dynamics simulations Biochemistry High 26284781
2016 Among human ATG8 orthologs (LC3B, GABARAPL2, GABARAP), only LC3B and GABARAP interact with cardiolipin (CL) in a biophysically characterized manner in vitro; GABARAPL2 does not interact with cardiolipin. Correspondingly, neither GABARAPL2 nor GABARAP translocated to mitochondria in rotenone-treated cells, while LC3B did, suggesting distinct roles for ATG8 orthologs in mitophagy. Quantitative biophysical binding assays (in vitro), fluorescence microscopy of mitochondrial translocation in human glioblastoma cells Autophagy Medium 27764541
2018 GATE-16 (GABARAPL2) plays a distinct role from GABARAP in regulating the TNF receptor Fn14: GATE-16 absence causes Fn14 accumulation within endosomes in the vicinity of autophagic membranes and regulates TWEAK signaling by Fn14 and NF-κB activity, whereas GABARAP (not GATE-16) regulates overall cellular levels of Fn14 and controls its accumulation at the ERGIC. Knockout cell lines, fluorescence microscopy, NF-κB activity assay, immunoprecipitation Nature communications Medium 30218067
2019 Residues within the core LIR motif and adjacent C-terminal region, plus ATG8 subfamily-specific residues in the LIR docking site (LDS), are critical for selective binding of autophagy receptors/adaptors to GABARAP subfamily proteins including GABARAPL2. Rendering GABARAP more LC3B-like impairs autophagy receptor degradation. The centriolar satellite protein PCM1's binding specificity for GABARAPL2 was shown to alter its cellular dynamics. In vitro binding assays, mutagenesis, cellular autophagy receptor degradation assays, live-cell imaging of PCM1 dynamics Nature communications High 31053714
2019 GABARAP subfamily proteins including GABARAPL2 interact with mammalian Syntaxin 16 (Stx16) and other SNAREs via LIR motifs, and mAtg8s regulate lysosome biogenesis. Stx16 knockout caused defects in lysosome biogenesis, while Stx16/Stx17 double knockout completely blocked autophagic flux. LIR motif identification, co-immunoprecipitation, knockout cell lines, autophagic flux assay, lysosome biogenesis assay The EMBO journal Medium 31625181
2020 GABARAP (and by extension GABARAPL2 as a member of the GABARAP subfamily) interacts with TFEB and IRGM, and GABARAP deletion affects global transcriptional responses to starvation and downregulates TFEB targets. IRGM and GABARAPs counter mTOR's negative regulation of TFEB, and this pathway is activated during M. tuberculosis and HIV infections. Co-immunoprecipitation, knockout cell lines, transcriptome analysis, mTOR/TFEB reporter assays, infection models Nature cell biology Medium 32753672
2020 ACSL3, an ER-associated lipid droplet biogenesis factor, is a stabilizing binding partner of endogenous GABARAPL2, identified using CRISPR/Cas9-tagged endogenous GABARAPL2 and interaction proteomics. GABARAPL2 binds ACSL3 via its LC3-interacting region binding site; this interaction recruits GABARAPL2 to the ER and anchors the UFM1-activating enzyme UBA5 at the ER. ACSL3 depletion and LD induction affect ufmylation components and ER-phagy. CRISPR/Cas9 endogenous tagging, interaction proteomics (IP-MS), mutagenesis of LIR-binding site, subcellular fractionation, siRNA depletion Journal of cell science High 32843575
2020 Irgm2 and Gate-16 (GABARAPL2) cooperatively dampen Gram-negative bacteria-induced caspase-11 non-canonical inflammasome activation in macrophages and in vivo. Gate-16 deficiency leads to increased GBP-dependent and GBP-independent caspase-11 targeting to intracellular bacteria, enhanced pyroptosis, and cytokine release. Knockout macrophages and mice, inflammasome activation assays, pyroptosis measurement, cytokine measurement, bacterial infection models EMBO reports High 33042141 33124769
2020 Gate-16/GABARAPL2 and Gabarap deficiency results in over-activation of caspase-11 inflammasomes (but not canonical inflammasomes) due to formation of GBP2-containing aggregates that promote IL-1β production. Gate-16/Gabarap double knockout mice show high mortality after low-dose LPS challenge, rescued by compound GBP2 deficiency. Double knockout mice and macrophages, inflammasome assays, in vivo LPS/poly(I:C) challenge, GBP2 knockout rescue Frontiers in immunology High 33042141
2020 GATE-16 (GABARAPL2) mediates curvature-sensitive membrane tethering via trans-assembly: in a reconstituted system with synthetic liposomes, GATE-16-PE more efficiently tethers flat large vesicles (200–400 nm diameter) compared to LC3B, while LC3B is more potent for highly curved small vesicles (50 nm). Membrane tethering requires trans-assembly and is reversible. Cell-free reconstitution with purified proteins and synthetic liposomes of defined curvature, quantitative tethering assays Protein science High 31960529
2020 GABARAP (as a member of GABARAP subfamily, with GABARAPL2 being a subfamily member) interact with SNAREs (Stx17 and Stx16) via LIR motifs to regulate autophagosome-lysosome fusion and lysosome biogenesis. GABARAPL2 was specifically identified as secreted inside small extracellular vesicles (sEVs) in a lipidation-dependent manner upon chloroquine treatment. Proteomics of extracellular vesicles, ATG16L1 mutant distinguishing single vs. double membrane lipidation, nanoparticle tracking Autophagy Medium 35220892
2020 GABARAP and LC3A bind key ESCRT-I components, contributing—along with other ESCRTs—to maintaining autophagosomal membranes in a sealed state. In cells lacking principal mATG8 proteins (including GABARAPL2), autophagosomal membranes are permeable and fail to mature into autolysosomes; autophagic organelles are arrested as amphisomes. mATG8 knockout cell lines, novel in vitro membrane sealing assay, ESCRT interaction studies, autophagic flux assays The EMBO journal High 37272163
2020 GABARAPL2 is critical for IFN-γ-induced growth restriction of Toxoplasma gondii in HeLa cells. GABARAPL2 is recruited to membrane structures surrounding parasitophorous vacuoles (PV). Autophagy adaptors are required for proper GABARAPL2 localization and function in this IFN-γ-induced immune response. GABARAPL2 knockdown/knockout, fluorescence microscopy, parasite growth assays, IFN-γ stimulation Infection and immunity Medium 32094251
2011 ORP7 interacts with GATE-16 (GABARAPL2) via residues 1-142 of ORP7 and residues 30-117 of GATE-16 (mapped by yeast two-hybrid and bimolecular fluorescence complementation). ORP7 overexpression negatively regulates GS28 stability via proteasomal degradation in a GATE-16-binding-dependent manner. Excess ORP7 also leads to formation of autophagic vacuoles containing GATE-16. Yeast two-hybrid screening, bimolecular fluorescence complementation (BiFC), ORP7 truncation mutants, siRNA knockdown, protein stability assays Experimental cell research Medium 21669198
2021 Septin-3 binds GABARAPL2 (and LC3B) directly; co-localization of septin-3 with LC3B increases upon chemical autophagy induction in primary neuronal cells. Septin-3 localizes to LC3B-positive membranes by electron microscopy. Co-immunoprecipitation/binding assays, fluorescence co-localization, electron microscopy, chemical autophagy induction Cellular and molecular life sciences Low 35932293
2021 Non-canonical autophagy (CASM) induces alternative lipidation of all ATG8 proteins including GABARAPL2 to phosphatidylserine (PS), in addition to the canonical PE conjugation. ATG8-PS and ATG8-PE adducts are differentially delipidated by ATG4 family members and have different cellular dynamics. ATG8-PS serves as a molecular signature for the non-canonical autophagy pathway. Lipidomics, pharmacological CASM induction (monensin, LC3-associated phagocytosis, influenza A), ATG4 delipidation assays, ATG16L1 WD40 mutants, mass spectrometry Molecular cell High 33909989
2022 The LMX1B transcription factor binds multiple ATG8 proteins including GABARAPL2. Binding is dependent on subcellular localization and nutrient status. ATG8 binding stimulates LMX1B-mediated transcription for efficient autophagy and stress protection in human iPSC-derived midbrain dopaminergic neurons. Co-immunoprecipitation, live-cell imaging, iPSC-derived neuronal differentiation, transcriptional reporter assays, rotenone toxicity assay The Journal of cell biology Medium 37014324
2023 The crystal structure of GABARAP (a close GABARAPL2 subfamily member) in complex with the non-canonical LIR motif of TAX1BP1 was solved, revealing a unique binding mode. TAX1BP1 selectively interacts with ATG8 family members via this non-canonical LIR. Crystal structure determination, isothermal titration calorimetry, mutagenesis, pull-down assays Proceedings of the National Academy of Sciences of the United States of America Medium 38437556
2003 GATE-16 (GABARAPL2) is proposed as an adapter in intra-Golgi SNARE priming: it assists NSF/SNAP-mediated SNARE dissociation (priming) in the Golgi, keeping dissociated cis-SNAREs apart to allow multiple fusion rounds. This review consolidates earlier biochemical evidence for GATE-16's role in membrane trafficking. Review consolidating earlier biochemical and cell-biology data; original biochemical evidence cited includes NSF/SNARE interaction studies Biochimica et biophysica acta Low 12914955

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 p62/SQSTM1 binds directly to Atg8/LC3 to facilitate degradation of ubiquitinated protein aggregates by autophagy. The Journal of biological chemistry 3849 17580304
2004 LC3, GABARAP and GATE16 localize to autophagosomal membrane depending on form-II formation. Journal of cell science 1205 15169837
2008 Atg8 controls phagophore expansion during autophagosome formation. Molecular biology of the cell 610 18508918
2008 The Atg8 and Atg12 ubiquitin-like conjugation systems in macroautophagy. 'Protein modifications: beyond the usual suspects' review series. EMBO reports 610 18704115
2019 Selective Autophagy: ATG8 Family Proteins, LIR Motifs and Cargo Receptors. Journal of molecular biology 548 31310766
2011 Atg8: an autophagy-related ubiquitin-like protein family. Genome biology 443 21867568
2010 Atg8-family interacting motif crucial for selective autophagy. FEBS letters 438 20083108
2016 Role of the mammalian ATG8/LC3 family in autophagy: differential and compensatory roles in the spatiotemporal regulation of autophagy. BMB reports 299 27418283
2011 LC3 and GATE-16 N termini mediate membrane fusion processes required for autophagosome biogenesis. Developmental cell 261 21497758
2013 The Atg8 family: multifunctional ubiquitin-like key regulators of autophagy. Essays in biochemistry 211 24070471
2021 Non-canonical autophagy drives alternative ATG8 conjugation to phosphatidylserine. Molecular cell 168 33909989
2020 Activation and targeting of ATG8 protein lipidation. Cell discovery 166 32377373
2009 Junctional adhesion molecule A interacts with Afadin and PDZ-GEF2 to activate Rap1A, regulate beta1 integrin levels, and enhance cell migration. Molecular biology of the cell 146 19176753
2016 ATG8 Expansion: A Driver of Selective Autophagy Diversification? Trends in plant science 137 28038982
2000 Structure of GATE-16, membrane transport modulator and mammalian ortholog of autophagocytosis factor Aut7p. The Journal of biological chemistry 136 10856287
2019 Molecular determinants regulating selective binding of autophagy adapters and receptors to ATG8 proteins. Nature communications 132 31053714
2013 Phosphorylation of mitophagy and pexophagy receptors coordinates their interaction with Atg8 and Atg11. EMBO reports 129 23559066
2009 Characterization of unusual families of ATG8-like proteins and ATG12 in the protozoan parasite Leishmania major. Autophagy 85 19066473
2020 Multiple Functions of ATG8 Family Proteins in Plant Autophagy. Frontiers in cell and developmental biology 84 32596242
2015 PI3P binding by Atg21 organises Atg8 lipidation. The EMBO journal 84 25691244
2020 Mammalian Atg8 proteins and the autophagy factor IRGM control mTOR and TFEB at a regulatory node critical for responses to pathogens. Nature cell biology 78 32753672
2015 Autophagy-Related Protein ATG8 Has a Noncanonical Function for Apicoplast Inheritance in Toxoplasma gondii. mBio 78 26507233
2023 Lysosome damage triggers direct ATG8 conjugation and ATG2 engagement via non-canonical autophagy. The Journal of cell biology 77 37796195
2023 Atg8 family proteins, LIR/AIM motifs and other interaction modes. Autophagy reports 74 38214012
2018 Coordination of Autophagosome-Lysosome Fusion by Atg8 Family Members. Current biology : CB 74 29689234
2017 Conserved Atg8 recognition sites mediate Atg4 association with autophagosomal membranes and Atg8 deconjugation. EMBO reports 71 28330855
2016 Human Atg8-cardiolipin interactions in mitophagy: Specific properties of LC3B, GABARAPL2 and GABARAP. Autophagy 71 27764541
2020 Atg8-Family Proteins-Structural Features and Molecular Interactions in Autophagy and Beyond. Cells 70 32882854
2019 Mechanisms and Pathophysiological Roles of the ATG8 Conjugation Machinery. Cells 70 31450711
2019 Noncanonical ATG8-ABS3 interaction controls senescence in plants. Nature plants 68 30664732
2022 Chloroquine treatment induces secretion of autophagy-related proteins and inclusion of Atg8-family proteins in distinct extracellular vesicle populations. Autophagy 62 35220892
2018 Potent and specific Atg8-targeting autophagy inhibitory peptides from giant ankyrins. Nature chemical biology 62 29867141
2020 The functions of Atg8-family proteins in autophagy and cancer: linked or unrelated? Autophagy 59 32255730
2016 Mechanism of cargo-directed Atg8 conjugation during selective autophagy. eLife 56 27879200
2023 AlphaFold2-multimer guided high-accuracy prediction of typical and atypical ATG8-binding motifs. PLoS biology 52 36753519
2016 Comparative analyses of ubiquitin-like ATG8 and cysteine protease ATG4 autophagy genes in the plant lineage and cross-kingdom processing of ATG8 by ATG4. Autophagy 52 27540766
2021 ATI1 (ATG8-interacting protein 1) and ATI2 define a plant starvation-induced reticulophagy pathway and serve as MSBP1/MAPR5 cargo receptors. Autophagy 50 33487099
2019 Mammalian Atg8 proteins regulate lysosome and autolysosome biogenesis through SNAREs. The EMBO journal 48 31625181
2020 Irgm2 and Gate-16 cooperatively dampen Gram-negative bacteria-induced caspase-11 response. EMBO reports 47 33124769
2012 Roles of putative Rho-GEF Gef2 in division-site positioning and contractile-ring function in fission yeast cytokinesis. Molecular biology of the cell 45 22298427
2019 N-terminal β-strand underpins biochemical specialization of an ATG8 isoform. PLoS biology 44 31329577
2012 Structural characterization and inhibition of the Plasmodium Atg8-Atg3 interaction. Journal of structural biology 44 22982544
2023 Shuffled ATG8 interacting motifs form an ancestral bridge between UFMylation and autophagy. The EMBO journal 43 36762703
2006 Haplotypes spanning SPEC2, PDZ-GEF2 and ACSL6 genes are associated with schizophrenia. Human molecular genetics 43 17030554
2016 Fluorescence-based ATG8 sensors monitor localization and function of LC3/GABARAP proteins. The EMBO journal 42 28028054
2015 Phospholipid methylation controls Atg32-mediated mitophagy and Atg8 recycling. The EMBO journal 42 26438722
2008 The RapGEF PDZ-GEF2 is required for maturation of cell-cell junctions. Cellular signalling 40 18585005
2022 Phosphorylation by casein kinase 2 enhances the interaction between ER-phagy receptor TEX264 and ATG8 proteins. EMBO reports 38 35417087
2013 Cooperation between Rho-GEF Gef2 and its binding partner Nod1 in the regulation of fission yeast cytokinesis. Molecular biology of the cell 38 23966468
2020 Analysis of Drosophila Atg8 proteins reveals multiple lipidation-independent roles. Autophagy 36 33249988
2017 Beyond Atg8 binding: The role of AIM/LIR motifs in autophagy. Autophagy 36 28121222
2019 Leishmania donovani parasite requires Atg8 protein for infectivity and survival under stress. Cell death & disease 35 31649242
2023 Mammalian ATG8 proteins maintain autophagosomal membrane integrity through ESCRTs. The EMBO journal 33 37272163
2017 Atg7 Activates an Autophagy-Essential Ubiquitin-like Protein Atg8 through Multi-Step Recognition. Journal of molecular biology 31 29237558
2012 ATI1, a newly identified atg8-interacting protein, binds two different Atg8 homologs. Plant signaling & behavior 29 22580699
2013 Inhibition of GATE-16 attenuates ATRA-induced neutrophil differentiation of APL cells and interferes with autophagosome formation. Biochemical and biophysical research communications 28 23891751
2022 Atg8-PE protein-based in vitro biochemical approaches to autophagy studies. Autophagy 26 35072587
2020 The trehalose-6-phosphate phosphatase Tps2 regulates ATG8 transcription and autophagy in Saccharomyces cerevisiae. Autophagy 26 32240040
2013 The immune system GTPase GIMAP6 interacts with the Atg8 homologue GABARAPL2 and is recruited to autophagosomes. PloS one 25 24204963
2022 Apicoplast biogenesis mediated by ATG8 requires the ATG12-ATG5-ATG16L and SNAP29 complexes in Toxoplasma gondii. Autophagy 24 36095096
2021 Phosphorylation of the LIR Domain of SCOC Modulates ATG8 Binding Affinity and Specificity. Journal of molecular biology 24 33845085
2020 Structure and Dynamics in the ATG8 Family From Experimental to Computational Techniques. Frontiers in cell and developmental biology 24 32587856
2020 ACSL3 is a novel GABARAPL2 interactor that links ufmylation and lipid droplet biogenesis. Journal of cell science 24 32843575
2017 Methods for Studying Interactions Between Atg8/LC3/GABARAP and LIR-Containing Proteins. Methods in enzymology 23 28253953
2003 Involvement of LMA1 and GATE-16 family members in intracellular membrane dynamics. Biochimica et biophysica acta 23 12914955
2022 Canonical and non-canonical roles for ATG8 proteins in autophagy and beyond. Frontiers in molecular biosciences 22 36601581
2021 ATG8-Interacting Motif: Evolution and Function in Selective Autophagy of Targeting Biological Processes. Frontiers in plant science 21 34912364
2020 Atg38-Atg8 interaction in fission yeast establishes a positive feedback loop to promote autophagy. Autophagy 21 31941401
2020 Role of Gate-16 and Gabarap in Prevention of Caspase-11-Dependent Excess Inflammation and Lethal Endotoxic Shock. Frontiers in immunology 21 33042141
2013 Characterization of Atg8 in lepidopteran insect cells. Archives of insect biochemistry and physiology 21 23959953
2009 Mutation at the cargo-receptor binding site of Atg8 also affects its general autophagy regulation function. Autophagy 21 19398890
2008 Association of haplotypes spanning PDZ-GEF2, LOC728637 and ACSL6 with schizophrenia in Han Chinese. Journal of medical genetics 21 18718982
2023 Post-translational modifications of ATG8 proteins - an emerging mechanism of autophagy control. Journal of cell science 20 37589340
2022 The evolutionary and functional divergence of the Atg8 autophagy protein superfamily. Biologia futura 20 35731422
2015 Atg8 is involved in endosomal and phagosomal acidification in the parasitic protist Entamoeba histolytica. Cellular microbiology 18 25923949
2011 OSBP-related protein 7 interacts with GATE-16 and negatively regulates GS28 protein stability. Experimental cell research 18 21669198
2019 CpATG8, a Homolog of Yeast Autophagy Protein ATG8, Is Required for Pathogenesis and Hypovirus Accumulation in the Chest Blight Fungus. Frontiers in cellular and infection microbiology 17 31355148
2019 Role of Atg8 in the regulation of vacuolar membrane invagination. Scientific reports 17 31616012
2014 In vitro systems for Atg8 lipidation. Methods (San Diego, Calif.) 17 25461810
2024 DDHD2, whose mutations cause spastic paraplegia type 54, enhances lipophagy via engaging ATG8 family proteins. Cell death and differentiation 16 38332048
2024 Phosphorylation of the selective autophagy receptor TAX1BP1 by TBK1 and IKBKE/IKKi promotes ATG8-family protein-dependent clearance of MAVS aggregates. Autophagy 16 39193925
2023 ATG8-dependent LMX1B-autophagy crosstalk shapes human midbrain dopaminergic neuronal resilience. The Journal of cell biology 16 37014324
2021 Long Non-coding RNAs Gabarapl2 and Chrnb2 Positively Regulate Inflammatory Signaling in a Mouse Model of Dry Eye. Frontiers in medicine 16 34957168
2013 Identification of atg8 isoform in encysting Acanthamoeba. The Korean journal of parasitology 16 24327773
2021 Overexpression of ATG8/LC3 enhances wound-induced somatic reprogramming in Physcomitrium patens. Autophagy 15 34612155
2020 Curvature-sensitive trans-assembly of human Atg8-family proteins in autophagy-related membrane tethering. Protein science : a publication of the Protein Society 15 31960529
2018 Autophagy differentially regulates TNF receptor Fn14 by distinct mammalian Atg8 proteins. Nature communications 15 30218067
2015 Conformational Polymorphism in Autophagy-Related Protein GATE-16. Biochemistry 15 26284781
2024 Mechanistic insights into the interactions of TAX1BP1 with RB1CC1 and mammalian ATG8 family proteins. Proceedings of the National Academy of Sciences of the United States of America 14 38437556
2020 GABARAPL2 Is Critical for Growth Restriction of Toxoplasma gondii in HeLa Cells Treated with Gamma Interferon. Infection and immunity 14 32094251
2024 Prohibitins, Phb1 and Phb2, function as Atg8 receptors to support yeast mitophagy and also play a negative regulatory role in Atg32 processing. Autophagy 13 38964378
2022 Unlocking the gate to GABARAPL2. Biologia futura 13 35486231
2022 Neuronal-specific septin-3 binds Atg8/LC3B, accumulates and localizes to autophagosomes during induced autophagy. Cellular and molecular life sciences : CMLS 13 35932293
2022 Development of new tools to study membrane-anchored mammalian Atg8 proteins. Autophagy 13 36250672
2020 Laodelphax striatellus Atg8 facilitates Rice stripe virus infection in an autophagy-independent manner. Insect science 13 32108430
2025 ATG8 delipidation is not universally critical for autophagy in plants. Nature communications 12 39757240
2025 Mycobacterium tuberculosis phagosome Ca2+ leakage triggers multimembrane ATG8/LC3 lipidation to restrict damage in human macrophages. Science advances 12 40138395
2020 Irgm2 and Gate-16 put a break on caspase-11 activation. EMBO reports 12 33135287
2025 TM9SF3 is a Golgi-resident ATG8-binding protein essential for Golgi-selective autophagy. Developmental cell 11 40609542
2022 Engineered ATG8-binding motif-based selective autophagy to degrade proteins and organelles in planta. The New phytologist 11 36263708

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