Function
Locations
Serine/threonine kinase that starts phagophore formation: it assembles with Atg13 at the phagophore assembly site and phosphorylates downstream autophagy proteins.
Autophagosomes form from a cup-shaped phagophore that grows around cytoplasmic cargo and closes into a double-membrane vesicle. Formation needs a conserved set of autophagy-related (Atg) proteins: the Atg1/ULK kinase with Atg13 starts assembly; the class III phosphatidylinositol 3-kinase complex (Vps34, Vps15, Atg6/Beclin and Atg14) marks the membrane with phosphatidylinositol 3-phosphate; Atg9 vesicles seed the membrane and the Atg2-Atg18/WIPI bridge feeds it lipid from the ER; and two ubiquitin-like conjugation systems link Atg12 to Atg5 and Atg8 to phosphatidylethanolamine. This module is the eukaryote-wide protein core, written so that it can be evaluated in any genome.
Eukaryote-wide scope: PAINT places nodes for Atg1/ULK, Vps34, Vps15, Atg6/Beclin, Atg9, Atg2, Atg7, Atg5, Atg12 and Atg8 at Eukaryota (taxon:2759). The Atg18/WIPI node carries no taxon and has seeds only from animals and budding yeast, so for that step the eukaryote-wide reach rests on the family entries table (yeast, Arabidopsis, Dictyostelium and human members). Each node is shown to be an ancestor of the human protein by its IBA rows or its IBD seeds. The Atg13, Atg3 and Atg4 families have no PAINT table locally, and their selectors rest on family membership. Read as a satisfiability rule, every non-optional part is expected in any eukaryote. Optional: Atg14, whose family is opisthokont-only in the local tables, and Atg16, whose family lacks the fungal protein. Molecular functions come from the human gene reviews' core functions; Atg7 carries one annoton for each conjugation arm, and Atg4 is typed from the human ATG4C review. Vps15 is untyped because its review records only a contribution to the Vps34 complex's kinase activity. Locations follow the human reviews' core-function locations: the phagophore assembly site for Atg1/ULK, Atg13, Vps34 and Atg2, the phagophore for Atg9, Atg14, Atg18/WIPI and the Atg12-Atg5-Atg16 complex, the cytosol for Atg7 and Atg3, the cytoplasm for Atg4 and the autophagosome membrane for Atg8. Two are inferred instead: Vps15 and Beclin are given the phagophore assembly site of their complex partner Vps34, because their own reviews do not record that location. Selector caveats are on each family: the Atg1/ULK and PROPPIN families over-match, and no Dictyostelium member is listed for Atg13 or Atg2. Not modelled: FIP200/Atg17 and ATG101, which have no local PANTHER entry for the human proteins and are not conserved as such across eukaryotes; Atg10, the E2 for Atg12 (no local entry for human ATG10); the Atg11 and receptor scaffolds of selective autophagy; and autophagosome closure and fusion with the lysosome or vacuole. The modules metazoan_autophagosome_biogenesis and phagophore_assembly_site describe the metazoan pathway and the yeast assembly site in more detail.
All recommended fields populated.
✗ none found
No MODULE:autophagosome_core_biogenesis deep-research report alongside the module YAML.
✓ every leaf node grounds to a representative protein.
✓ every declared conforms_to bundle matches its template motif.
✓ every PRECEDES step chains, or its break is acknowledged via chaining_status.
17 gene-function check(s) reflected in core functions · 0 conflict(s) · 14 evidence or coverage gap(s) (34 comparison(s))
Retained noncore functions are reported separately; they do not require promotion to core functions. Support from representative genes does not establish conservation across the whole family. Missing reviews or identifiers leave a claim unverified.
| Annoton | Participant | Function | Assessment |
|---|---|---|---|
| atg1_ulk_kinase | ULK1 (human) UniProtKB:O75385 Family Representative |
protein serine/threonine kinase activity GO:0004674 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg13_regulator | ATG13 (human) UniProtKB:O75143 Family Representative |
protein kinase regulator activity GO:0019887 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| vps34_kinase | PIK3C3 (human) UniProtKB:Q8NEB9 Family Representative |
1-phosphatidylinositol-3-kinase activity GO:0016303 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg6_beclin | BECN1 (human) UniProtKB:Q14457 Family Representative |
protein-macromolecule adaptor activity GO:0030674 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg14_targeting | ATG14 (human) UniProtKB:Q6ZNE5 Family Representative |
protein-membrane adaptor activity GO:0043495 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg9_scramblase | ATG9A (human) UniProtKB:Q7Z3C6 Family Representative |
phospholipid scramblase activity GO:0017128 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg2_lipid_transfer | ATG2A (human) UniProtKB:Q2TAZ0 Family Representative |
lipid transfer activity GO:0120013 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg18_wipi | WIPI2 (human) UniProtKB:Q9Y4P8 Family Representative |
phosphatidylinositol-3-phosphate binding GO:0032266 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg7_e1 | ATG7 (human) UniProtKB:O95352 Family Representative |
Atg12 activating enzyme activity GO:0019778 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg7_e1_atg8 | ATG7 (human) UniProtKB:O95352 Family Representative |
Atg8 activating enzyme activity GO:0019779 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg12_ubl | ATG12 (human) UniProtKB:O94817 Family Representative |
protein tag activity GO:0031386 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg5_e3 | ATG5 (human) UniProtKB:Q9H1Y0 Family Representative |
Atg8-family ligase activity GO:0019776 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg3_e2 | ATG3 (human) UniProtKB:Q9NT62 Family Representative |
Atg8-family conjugating enzyme activity GO:0141046 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg4_protease | ATG4C (human) UniProtKB:Q96DT6 Family Representative |
protein-phosphatidylethanolamide deconjugating activity GO:0019786 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg4_protease | ATG4B (human) UniProtKB:Q9Y4P1 Family Representative |
protein-phosphatidylethanolamide deconjugating activity GO:0019786 |
Review Missing
No ai-review exists for this protein; its function and core-function coverage cannot yet be checked. |
| atg4_protease | ATG4D (human) UniProtKB:Q86TL0 Family Representative |
protein-phosphatidylethanolamide deconjugating activity GO:0019786 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg8_lc3 | MAP1LC3B (human) UniProtKB:Q9GZQ8 Family Representative |
phosphatidylethanolamine binding GO:0008429 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg16_complex | ATG16L1 (human) UniProtKB:Q676U5 Family Representative |
protein-membrane adaptor activity GO:0043495 |
Core Supported
The GO molecular function is reflected in core_functions (exact term or a reviewed subclass). |
| atg1_ulk_kinase | Atg1/ULK kinase family PANTHER:PTHR24348 Family |
protein serine/threonine kinase activity GO:0004674 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR24348. |
| atg13_regulator | Atg13 subfamily PANTHER:PTHR13430:SF4 Family |
protein kinase regulator activity GO:0019887 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR13430:SF4. |
| vps34_kinase | Vps34 subfamily PANTHER:PTHR10048:SF7 Family |
1-phosphatidylinositol-3-kinase activity GO:0016303 |
Family Supported
Family review supports GO:0016303 for PANTHER:PTHR10048:SF7. All three PI3K classes phosphorylate phosphatidylinositol at the 3-position: class III VPS34 (SF7, selected by the phosphoinositide_biosynthesis and phagosome_maturation modules, and its sole activity), the class II PI3Ks (SF28, SF29, SF30), and the class I catalytic subunits in vitro (SF107, SF33, SF34, SF35), whose PIK3CA and PIK3CD reviews keep the term. Not the type III PI4Ks. SF111 (C. elegans AGE-1, a class I catalytic subunit) is listed too; its review keeps the in vitro activity as non-core. For the class I subfamilies the scope records a measured in vitro activity, not the physiological one: in cells their main substrate is PI(4,5)P2. Read "safe" for those subfamilies with that caveat. |
| atg6_beclin | Atg6/Beclin family PANTHER:PTHR12768 Family |
protein-macromolecule adaptor activity GO:0030674 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR12768. |
| atg14_targeting | Atg14/Barkor family PANTHER:PTHR13664 Family |
protein-membrane adaptor activity GO:0043495 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR13664. |
| atg9_scramblase | Atg9 family PANTHER:PTHR13038 Family |
phospholipid scramblase activity GO:0017128 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR13038. |
| atg2_lipid_transfer | Atg2 family PANTHER:PTHR13190 Family |
lipid transfer activity GO:0120013 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR13190. |
| atg18_wipi | PROPPIN (Atg18/WIPI) family PANTHER:PTHR11227 Family |
phosphatidylinositol-3-phosphate binding GO:0032266 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR11227. |
| atg7_e1 | Atg7 subfamily PANTHER:PTHR10953:SF3 Family |
Atg12 activating enzyme activity GO:0019778 |
Family Supported
Family review supports GO:0019778 for PANTHER:PTHR10953:SF3. ATG12 activation is the specialization of the ATG7 subfamily SF3; other E1-like subfamilies activate different carriers. |
| atg7_e1_atg8 | Atg7 subfamily PANTHER:PTHR10953:SF3 Family |
Atg8 activating enzyme activity GO:0019779 |
Family Supported
Family review supports GO:0019779 for PANTHER:PTHR10953:SF3. ATG8-family activation is the second activity of the ATG7 subfamily SF3. |
| atg12_ubl | Atg12 family PANTHER:PTHR13385 Family |
protein tag activity GO:0031386 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR13385. |
| atg5_e3 | Atg5 family PANTHER:PTHR13040 Family |
Atg8-family ligase activity GO:0019776 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR13040. |
| atg3_e2 | Atg3 subfamily PANTHER:PTHR12866:SF2 Family |
Atg8-family conjugating enzyme activity GO:0141046 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR12866:SF2. |
| atg4_protease | Atg4 family PANTHER:PTHR22624 Family |
protein-phosphatidylethanolamide deconjugating activity GO:0019786 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR22624. |
| atg8_lc3 | Atg8 family PANTHER:PTHR10969 Family |
phosphatidylethanolamine binding GO:0008429 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR10969. |
| atg16_complex | Atg16-like family PANTHER:PTHR19878 Family |
protein-membrane adaptor activity GO:0043495 |
Family Review Missing
No supported structured family review is available for PANTHER:PTHR19878. |
8 complete review(s) · 7 with deep research · 1 missing review · 10 reviewed but lacking deep research
| Gene | Review | Complete | Deep research |
|---|---|---|---|
| ATG12 O94817 | ✓ | 54/55 | ✗ |
| ATG13 O75143 | ✓ | ✓ | ✗ |
| ATG14 Q6ZNE5 | ✓ | 93/95 | ✓ |
| ATG16L1 Q676U5 | ✓ | ✓ | ✗ |
| ATG2A Q2TAZ0 | ✓ | 30/31 | ✓ |
| ATG3 Q9NT62 | ✓ | ✓ | ✗ |
| ATG4C Q96DT6 | ✓ | ✓ | ✗ |
| ATG4D Q86TL0 | ✓ | ✓ | ✓ |
| ATG5 Q9H1Y0 | ✓ | 88/90 | ✓ |
| ATG7 O95352 | ✓ | ✓ | ✓ |
| ATG9A Q7Z3C6 | ✓ | ✓ | ✗ |
| BECN1 Q14457 | ✓ | 189/191 | ✓ |
| MAP1LC3B Q9GZQ8 | ✓ | ✓ | ✗ |
| PIK3C3 Q8NEB9 | ✓ | 108/119 | ✓ |
| PIK3R4 Q99570 | ✓ | 64/66 | ✗ |
| ATG4B (human) Q9Y4P1 | ✗ | — | — |
| ULK1 O75385 | ✓ | 137/139 | ✗ |
| WIPI2 Q9Y4P8 | ✓ | 43/45 | ✗ |
Serine/threonine kinase that starts phagophore formation: it assembles with Atg13 at the phagophore assembly site and phosphorylates downstream autophagy proteins.
HORMA-domain subunit that binds Atg1/ULK and recruits the kinase complex to the phagophore assembly site.
Lipid kinase that makes phosphatidylinositol 3-phosphate on the phagophore membrane.
Pseudokinase-like scaffold that anchors Vps34 to membranes and is required for its activity. It contributes to the complex's phosphatidylinositol 3-kinase activity rather than enabling it, and an annoton function would assert enabling, so the step is left untyped.
Coiled-coil scaffold that holds the Vps34-Vps15 core together with Atg14 or UVRAG.
Optional because the family entries table lists only fission yeast and animal members and the PAINT node is placed at Opisthokonta (taxon:33154); whether plants and other lineages have an Atg14 in another family has not been checked.
Subunit that directs the Vps34 complex to the phagophore assembly site.
Integral-membrane trimer, carried on small vesicles, that seeds the phagophore and equilibrates lipids between its leaflets.
Rod-shaped protein that tethers the phagophore rim to the ER and channels lipids into the growing membrane.
PROPPIN beta-propeller that binds phosphatidylinositol 3-phosphate on the phagophore and recruits Atg2 and the Atg16 complex.
E1-like enzyme that activates Atg12 for conjugation to Atg5.
The same enzyme activating Atg8 for transfer to Atg3.
Ubiquitin-like protein conjugated to Atg5.
Acceptor of Atg12; the Atg12-Atg5 conjugate acts as the E3-like enzyme that transfers Atg8 to phosphatidylethanolamine.
E2-like enzyme that carries Atg8 from Atg7 and transfers it to phosphatidylethanolamine.
Cysteine protease that exposes the C-terminal glycine of Atg8 for conjugation and later removes Atg8 from phosphatidylethanolamine.
Ubiquitin-like protein lipidated onto phosphatidylethanolamine of the phagophore membrane, where it drives expansion and binds cargo receptors.
Optional because the selector does not reach fungi: interpro/panther/PTHR19878/PTHR19878-entries.csv lists Arabidopsis, Dictyostelium and human Atg16 proteins but no yeast Atg16, which is a much smaller coiled-coil protein, so a fungal genome would fail the step for want of a selector rather than of the protein.
Binds Atg12-Atg5 and targets the conjugate to the phagophore, setting where Atg8 is lipidated.