atg16

UniProt ID: O94656
Organism: Schizosaccharomyces pombe (strain 972 / ATCC 24843)
Review Status: DRAFT
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Gene Description

Atg16 is a core autophagy protein that functions as an essential structural component of the Atg12-Atg5-Atg16 complex (E3-like ligase), which promotes the conjugation of Atg8 to phosphatidylethanolamine (PE) during autophagosome biogenesis. The protein contains an N-terminal Atg5-binding domain and a C-terminal coiled-coil domain required for homodimerization. In S. pombe, Atg16 localizes to the phagophore assembly site (PAS) in a manner dependent on Atg18a and is required for efficient macroautophagy.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005737 cytoplasm
IEA
GO_REF:0000044
ACCEPT
Summary: IEA annotation derived from UniProtKB-SubCell mapping. Atg16 is predominantly cytosolic in vegetative cells, with a fraction localizing to the PAS upon autophagy induction. High-throughput localization data from PMID:16823372 confirms cytoplasmic localization.
Reason: Cytoplasmic localization is consistent with the known biology of Atg16. The protein exists in the cytoplasm and is recruited to the PAS upon autophagy induction. This annotation correctly captures the baseline localization.
Supporting Evidence:
PMID:16823372
we determined the localization of 4,431 proteins, corresponding to approximately 90% of the fission yeast proteome
GO:0006914 autophagy
IEA
GO_REF:0000043
MODIFY
Summary: IEA annotation based on UniProtKB keyword mapping from the Autophagy keyword. Atg16 is a core autophagy protein essential for autophagosome biogenesis. Direct experimental evidence supports its role in macroautophagy [PMID:23950735].
Reason: While not incorrect, this annotation is too general. More specific experimental evidence (IMP from PMID:23950735) supports a role in macroautophagy (GO:0016236), which is the correct specific term for this protein. The IEA should be retained but the more specific macroautophagy term (already annotated with IMP) better captures the function.
Proposed replacements: macroautophagy
Supporting Evidence:
PMID:23950735
we obtained a comprehensive catalog of autophagy genes in this highly tractable organism, including genes encoding three heretofore unidentified core Atg proteins, Atg10, Atg14, and Atg16
file:SCHPO/atg16/atg16-deep-research-falcon.md
**atg16ฮ”** cells are defective in CFP-Atg8 processing under nitrogen starvation
GO:0015031 protein transport
IEA
GO_REF:0000043
MARK AS OVER ANNOTATED
Summary: IEA annotation based on UniProtKB keyword mapping from the Protein Transport keyword. This appears to be an indirect annotation reflecting that autophagy involves protein transport to the vacuole/lysosome.
Reason: Atg16 does not directly function as a protein transporter. Its role is as a scaffold/E3-like ligase component in the Atg12-Atg5-Atg16 complex that promotes Atg8 lipidation. The protein transport annotation is too generic and does not capture the specific molecular function. Autophagy does involve protein transport to the vacuole, but this is not Atg16's specific function.
Supporting Evidence:
PMID:23950735
Atg18a uniquely required for the targeting of the Atg12-Atg5ยทAtg16 complex
file:SCHPO/atg16/atg16-deep-research-falcon.md
Atg16 is the scaffold subunit of the **Atg12โ€“Atg5โ€“Atg16** complex, which functions as the **E3-like factor for Atg8 conjugation to phosphatidylethanolamine (PE)** and helps determine where Atg8 lipidation occurs on autophagic membranes.
GO:0034045 phagophore assembly site membrane
IEA
GO_REF:0000044
MODIFY
Summary: IEA annotation based on UniProtKB subcellular location mapping. UniProt notes that Atg16 localizes to the PAS membrane as a peripheral membrane protein. Direct experimental evidence from PMID:23950735 and PMID:31941401 confirms PAS localization.
Reason: This annotation is accurate and consistent with experimental evidence showing Atg16 localizes to the PAS upon autophagy induction. The protein is known to function at the PAS membrane during autophagosome biogenesis. [2026-08 re-review against GO issue #29437] The supporting evidence quoted here describes PAS puncta / PAS accumulation, not a membrane. GO:0034045 asserts, via bounding_layer_of, that a membrane bounds the phagophore assembly site; the PAS is a liquid-like protein condensate with no bounding bilayer (PMID:32025038), and the term additionally carries 'phagophore' and 'isolation membrane' as synonyms of the distinct term GO:0061908. This annotation derives solely from the UniProt SL-0221 subcellular location via GO_REF:0000044, which is the propagation source rather than independent evidence. Generalize to the structure the cited experiments actually show.
Proposed replacements: phagophore assembly site
Supporting Evidence:
PMID:23950735
atg18aฮ” abolished the starvation-induced puncta formation by Atg5 and Atg16
PMID:31941401
PAS accumulation of Atg2, Atg18b, Atg24b, Atg5, Atg16, and Atg8 reduced by the Atg38 AIM mutation was recovered by expressing 3ร—EEEWEEL-Atg14 or Atg14-3ร—EEEWEEL
file:SCHPO/atg16/atg16-deep-research-falcon.md
Atg16 localizes to the **PAS (phagophore assembly site / pre-autophagosomal structure)** during starvation and colocalizes with **CFP-Atg8**
GO:0051321 meiotic cell cycle
IEA
GO_REF:0000043
REMOVE
Summary: IEA annotation based on UniProtKB keyword mapping from the Meiosis keyword. The Meiosis keyword was assigned to atg16 based on PMID:16169489, a high-throughput knockout screen that identified genes affecting meiotic chromosome segregation. However, this is a classic case of over-annotation: autophagy is upregulated during meiosis/sporulation in yeast, and loss of autophagy genes affects meiotic processes indirectly rather than through direct involvement in the meiotic cell cycle machinery.
Reason: This annotation represents an over-annotation resulting from the indirect effects of autophagy defects on meiosis. Atg16 is a core autophagy protein, not a meiotic cell cycle regulator. The screen in PMID:16169489 identified atg16 because autophagy is required during nutrient starvation that triggers meiosis/sporulation in fission yeast. The meiotic defects observed in atg16 mutants are secondary consequences of impaired autophagy, not evidence that Atg16 directly participates in meiotic cell cycle regulation. ATG genes are autophagy machinery components, and this keyword assignment conflates upstream/downstream relationships. The protein's core function is in autophagosome biogenesis, not meiotic regulation.
Supporting Evidence:
file:SCHPO/atg16/atg16-deep-research-perplexity.md
The atg16 gene in Schizosaccharomyces pombe encodes autophagy protein 16, a conserved component of the autophagy machinery
GO:0000407 phagophore assembly site
IDA
PMID:31941401
Atg38-Atg8 interaction in fission yeast establishes a positi...
ACCEPT
Summary: IDA annotation showing Atg16 localizes to the phagophore assembly site (PAS). This study by Yu et al. (2020) demonstrated that the Atg38-Atg8 interaction affects PAS accumulation of multiple Atg proteins including Atg16. The study showed that PAS accumulation of Atg16 was reduced by the Atg38 AIM mutation and could be recovered by artificial Atg14-Atg8 interaction constructs.
Reason: Direct experimental evidence from fluorescence microscopy demonstrates Atg16 localizes to the PAS. This is a core localization for Atg16 function in autophagosome biogenesis.
Supporting Evidence:
PMID:31941401
PAS accumulation of Atg2, Atg18b, Atg24b, Atg5, Atg16, and Atg8 reduced by the Atg38 AIM mutation was recovered by expressing 3ร—EEEWEEL-Atg14 or Atg14-3ร—EEEWEEL
GO:0000407 phagophore assembly site
IDA
PMID:23950735
Global analysis of fission yeast mating genes reveals new au...
ACCEPT
Summary: IDA annotation showing Atg16 localizes to the PAS. The study by Sun et al. 2013 systematically examined subcellular localization of fission yeast autophagy factors and showed that Atg16 forms cytoplasmic puncta that colocalize with Atg8 at the PAS upon starvation induction. The study also showed that Atg18a is required for Atg16 recruitment to the PAS.
Reason: This is the primary experimental study demonstrating PAS localization of Atg16 in S. pombe. The evidence includes colocalization with Atg8 puncta and demonstration that recruitment requires the PI3P-binding protein Atg18a.
Supporting Evidence:
PMID:23950735
Fifteen Atg proteins colocalized with CFP-Atg8 at cytoplasmic puncta induced by starvation
PMID:23950735
atg18aฮ” abolished the starvation-induced puncta formation by Atg5 and Atg16
PMID:23950735
Atg18a may serve as a binding platform for the recruitment of the Atg12โ€“Atg5ยทAtg16 complex to PAS
file:SCHPO/atg16/atg16-deep-research-falcon.md
Atg18a is uniquely required to target the Atg12โ€“Atg5ยทAtg16 complex to the PAS
GO:0016236 macroautophagy
IMP
PMID:23950735
Global analysis of fission yeast mating genes reveals new au...
ACCEPT
Summary: IMP annotation showing Atg16 is involved in macroautophagy. The study identified atg16 as a core autophagy gene through genome-wide screening. Deletion of atg16 causes defects in CFP-Atg8 processing, a standard assay for autophagy function. The study demonstrated that Atg16 is part of the Atg12-Atg5-Atg16 complex required for autophagosome biogenesis.
Reason: This is the core biological process annotation for Atg16. Direct genetic evidence shows that atg16 deletion impairs autophagy, and the protein functions in the conserved Atg12-Atg5-Atg16 complex that promotes Atg8 lipidation. This is the primary function of Atg16.
Supporting Evidence:
PMID:23950735
we obtained a comprehensive catalog of autophagy genes in this highly tractable organism, including genes encoding three heretofore unidentified core Atg proteins, Atg10, Atg14, and Atg16
PMID:23950735
SPBC405.05 is currently annotated as a sequence orphan. We found that it shares homology with S. cerevisiae Atg16 in both the N-terminal Atg5-binding domain and the C-terminal coiled-coil domain
PMID:23950735
S. pombe SPBC405.05/Atg16 protein interacts with Atg5 both in the presence and in the absence of Atg12
file:SCHPO/atg16/atg16-deep-research-falcon.md
**atg16ฮ”** cells are defective in CFP-Atg8 processing under nitrogen starvation
GO:0019776 Atg8-family ligase activity
ISO
GO_REF:0000024
ACCEPT
Summary: ISO annotation transferred from S. cerevisiae Atg16 (SGD:S000004769). The Atg12-Atg5-Atg16 complex functions as an E3-like ubiquitin ligase that promotes the conjugation of Atg8 to phosphatidylethanolamine. This activity is well characterized in budding yeast and mammals and is conserved in S. pombe.
Reason: This molecular function annotation is appropriate for Atg16. The protein is part of the Atg12-Atg5-Atg16 complex that acts as an E3-like enzyme for Atg8 lipidation. The deep research document confirms this is a well-conserved function. The "contributes_to" qualifier is appropriate since Atg16 is one component of a multi-subunit complex.
Supporting Evidence:
file:SCHPO/atg16/atg16-deep-research-perplexity.md
The ATG12โ€“ATG5โ€“ATG16 complex represents one of the most critical assemblies in the autophagy pathway, functioning as a specialized E3-like ubiquitin ligase
file:SCHPO/atg16/atg16-deep-research-falcon.md
Atg16 is the scaffold subunit of the **Atg12โ€“Atg5โ€“Atg16** complex, which functions as the **E3-like factor for Atg8 conjugation to phosphatidylethanolamine (PE)** and helps determine where Atg8 lipidation occurs on autophagic membranes.
GO:0005737 cytoplasm
HDA
PMID:16823372
ORFeome cloning and global analysis of protein localization ...
ACCEPT
Summary: HDA (High-throughput Direct Assay) annotation from a large-scale protein localization study. Matsuyama et al. performed ORFeome cloning and global analysis of protein localization in S. pombe using fluorescent protein tagging.
Reason: This annotation is supported by systematic localization data. Atg16 is predominantly cytoplasmic in unstressed cells, consistent with its known biology as an autophagy protein that is recruited to the PAS upon autophagy induction.
Supporting Evidence:
PMID:16823372
we determined the localization of 4,431 proteins, corresponding to approximately 90% of the fission yeast proteome
GO:0005829 cytosol
HDA
PMID:16823372
ORFeome cloning and global analysis of protein localization ...
ACCEPT
Summary: HDA annotation from the same large-scale localization study (PMID:16823372). The cytosol annotation is more specific than cytoplasm and indicates the protein is soluble rather than membrane-associated under basal conditions.
Reason: Cytosolic localization is consistent with Atg16 being a soluble protein that is recruited to the PAS membrane upon autophagy induction. The protein does not have transmembrane domains and functions as a peripheral membrane protein when localized to the PAS.
Supporting Evidence:
PMID:16823372
we determined the localization of 4,431 proteins, corresponding to approximately 90% of the fission yeast proteome

Core Functions

Atg16 is an essential component of the Atg12-Atg5-Atg16 complex that functions as an E3-like ubiquitin ligase to promote the conjugation of Atg8 to phosphatidylethanolamine during autophagosome biogenesis. The protein contains an N-terminal Atg5-binding domain and a C-terminal coiled-coil domain required for homodimerization. Upon autophagy induction, Atg16 is recruited to the phagophore assembly site (PAS) in a manner dependent on the PI3P-binding protein Atg18a. At the PAS, the Atg12-Atg5-Atg16 complex promotes Atg8 lipidation, which is essential for autophagosome formation. Deletion of atg16 causes defects in Atg8 processing and impairs macroautophagy.

Molecular Function:
Atg8-family ligase activity
Directly Involved In:
Cellular Locations:
Supporting Evidence:
  • PMID:23950735
    Atg18a uniquely required for the targeting of the Atg12-Atg5ยทAtg16 complex
  • PMID:31941401
    PAS accumulation of Atg2, Atg18b, Atg24b, Atg5, Atg16, and Atg8 reduced by the Atg38 AIM mutation
  • file:SCHPO/atg16/atg16-deep-research-falcon.md
    Atg16 is the scaffold subunit of the **Atg12โ€“Atg5โ€“Atg16** complex, which functions as the **E3-like factor for Atg8 conjugation to phosphatidylethanolamine (PE)** and helps determine where Atg8 lipidation occurs on autophagic membranes.

References

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Suggested Questions for Experts

Q: What is the precise stoichiometry of the Atg12-Atg5-Atg16 complex in S. pombe? While the budding yeast and mammalian complexes have been well characterized structurally, the S. pombe complex stoichiometry has not been directly determined.

Q: Does S. pombe Atg16 have membrane-binding activity via an amphipathic helix like S. cerevisiae Atg16? The membrane-binding capability of Atg16 via an amphipathic helix has been characterized in budding yeast but not specifically confirmed in fission yeast.

Suggested Experiments

Experiment: Structure determination of S. pombe Atg12-Atg5-Atg16 complex to confirm conserved structural features and enable comparison with budding yeast and mammalian complexes.

Hypothesis: The S. pombe Atg12-Atg5-Atg16 complex shares the same structural organization as the budding yeast and mammalian complexes.

Experiment: Direct measurement of Atg8-PE conjugation activity with purified S. pombe Atg12-Atg5-Atg16 complex to provide direct biochemical evidence for the E3-like ligase activity in the fission yeast system.

Hypothesis: The S. pombe Atg12-Atg5-Atg16 complex has E3-like ligase activity that promotes Atg8 lipidation in vitro.

Deep Research

Falcon

(atg16-deep-research-falcon.md)

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Perplexity

(atg16-deep-research-perplexity.md)

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๐Ÿ“š Additional Documentation

Bioreason Rl Predictions

(atg16-bioreason-rl-predictions.md)

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Bioreason Rl Review

(atg16-bioreason-rl-review.md)

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๐Ÿ“„ View Raw YAML

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