{"gene":"ANAPC13","run_date":"2026-04-28T17:12:37","timeline":{"discoveries":[{"year":2004,"finding":"Swm1/Apc13 is an evolutionarily conserved subunit of the APC/C that promotes the stable association of the essential TPR subunits Cdc16 and Cdc27 with the complex; without Swm1/Apc13, Cdc16 and Cdc27 dissociate from the APC/C, abolishing ubiquitin ligase activity in vitro and causing delayed execution of APC/C-dependent cell cycle events in vivo. Human and fission yeast homologues associate with APC/C subunits and complement the yeast swm1Δ deletion phenotype.","method":"Genetic deletion, in vitro ubiquitin ligase assay, co-immunoprecipitation, cross-species complementation","journal":"Molecular and cellular biology","confidence":"High","confidence_rationale":"Tier 1–2 — multiple orthogonal methods (co-IP, in vitro reconstitution, genetic rescue) in a single study with clear functional readout","pmids":["15060174"],"is_preprint":false},{"year":2003,"finding":"Swm1 (Apc13) was identified as a constitutive core subunit of the yeast APC by mass spectrometry of highly purified APC preparations; it is present throughout the cell cycle (G1, S, M) and in meiotic cells. In vitro transcription/translation showed Swm1 interacts directly with Cdc23 and Apc5. Deletion of SWM1 causes slow growth and accumulation of G2/M cells, consistent with an APC defect.","method":"Mass spectrometry of purified APC, co-immunoprecipitation with epitope-tagged subunits, in vitro co-translation interaction assay, genetic analysis of deletion strains","journal":"The Journal of biological chemistry","confidence":"High","confidence_rationale":"Tier 1–2 — biochemical identification by MS, reciprocal pull-down, in vitro interaction, and genetic phenotype in one study","pmids":["12609981"],"is_preprint":false},{"year":2014,"finding":"Cryo-EM reconstruction of the human APC/C at 7.4 Å resolution placed APC13 within the 'Arc Lamp' subcomplex alongside APC16 and CDC26, defining its position within the overall 1.2 MDa assembly alongside all 20 APC/C subunits.","method":"Cryo-electron microscopy reconstruction","journal":"Nature","confidence":"High","confidence_rationale":"Tier 1 — near-atomic cryo-EM structure of the full APC/C complex with all subunits positioned","pmids":["25043029"],"is_preprint":false},{"year":2015,"finding":"Atomic-resolution cryo-EM structures of APC/C-coactivator complexes confirm APC13 as part of the Arc Lamp sub-structure; the structures define the architecture of all APC/C subunits and explain how Emi1 mediates inhibition.","method":"Cryo-electron microscopy at near-atomic resolution, biochemical validation","journal":"Nature","confidence":"High","confidence_rationale":"Tier 1 — atomic cryo-EM with mutagenesis-based functional validation","pmids":["26083744"],"is_preprint":false},{"year":2014,"finding":"Crystal structures of human APC3 alone and in complex with the C-terminal domain of APC16 revealed that APC13 is part of the Arc Lamp alongside APC16 and APC7; the data show that APC16 bridges APC13 context within the TPR sub-complex and recruits APC7 to APC3.","method":"X-ray crystallography, biochemical mapping, in vitro ubiquitination assay","journal":"Journal of molecular biology","confidence":"High","confidence_rationale":"Tier 1 — crystal structure plus biochemical functional assays","pmids":["25490258"],"is_preprint":false},{"year":2010,"finding":"Tandem-affinity purification-mass spectrometry and localization studies in human cells identified APC13 as a previously unknown, evolutionarily conserved subunit of the human anaphase-promoting complex essential for chromosome segregation.","method":"TAP-MS, protein localization by fluorescence microscopy, RNAi phenotypic screening","journal":"Science","confidence":"High","confidence_rationale":"Tier 2 — large-scale TAP-MS with functional validation by RNAi and localization","pmids":["20360068"],"is_preprint":false},{"year":2024,"finding":"The cancer-associated SF3B1-K700E mutation causes aberrant splicing of ANAPC13 pre-mRNA, inserting a 231-bp fragment into the 5′ UTR and reducing ANAPC13 protein expression. Reduced ANAPC13 impairs regulatory T cell (Treg) differentiation and inhibitory function; forced re-expression of ANAPC13 restores Treg function and prevents adoptive transfer colitis.","method":"Conditional knock-in mouse model, RNA splicing analysis, adoptive transfer colitis assay, rescue by forced ANAPC13 expression","journal":"Science advances","confidence":"High","confidence_rationale":"Tier 2 — genetic mouse model with mechanistic splicing analysis and functional rescue experiment","pmids":["39303038"],"is_preprint":false},{"year":2025,"finding":"Compound heterozygous missense mutations in APC13 (c.C6A and c.116_126del) were identified in an infertile female patient; structural modelling predicted disrupted chemical bonds with other APC/C subunits, and in vitro experiments demonstrated aberrant cellular localization of the mutant APC13 protein, implicating correct APC13 localization as necessary for APC/C-dependent female fertility.","method":"Whole-exome sequencing, structural modelling, in vitro localization assay in cell lines","journal":"Journal of assisted reproduction and genetics","confidence":"Medium","confidence_rationale":"Tier 2–3 — patient genetics with in vitro localization assay; structural modelling is computational","pmids":["40238067"],"is_preprint":false},{"year":2026,"finding":"Biallelic ANAPC13 mutations (p.D2E and p.L24R) cause oocyte maturation arrest at metaphase I in humans and in a knock-in mouse model. Mutant ANAPC13 disrupts APC/C subunit interactions, impairs APC/C ubiquitin ligase function (without altering spindle assembly checkpoint dynamics), and leads to abnormal protein composition during the metaphase I-to-anaphase I transition. Microinjection of wild-type Anapc13 mRNA partially rescues polar body extrusion (49%).","method":"Whole-exome sequencing, knock-in mouse model, in vitro maturation assay, proteomic analysis of oocytes, co-IP/subunit interaction assay, mRNA microinjection rescue","journal":"American journal of obstetrics and gynecology","confidence":"High","confidence_rationale":"Tier 1–2 — multiple orthogonal methods including animal model, proteomics, molecular interaction assay, and mRNA rescue in one study","pmids":["41997520"],"is_preprint":false}],"current_model":"ANAPC13 (Swm1/Apc13) is a small, evolutionarily conserved core subunit of the APC/C E3 ubiquitin ligase that resides in the Arc Lamp sub-complex and is required for stable association of the TPR subunits CDC16 and CDC27 with the complex, thereby maintaining APC/C ubiquitin ligase activity, proper cell cycle progression, and—in humans—oocyte maturation; its expression is regulated post-transcriptionally by SF3B1-dependent splicing, and loss of ANAPC13 impairs both APC/C function and regulatory T cell differentiation."},"narrative":{"teleology":[{"year":2003,"claim":"The identity of Swm1/Apc13 as a constitutive APC/C core subunit was established, resolving whether it was a stoichiometric component or a transient interactor, and revealing its direct contacts with Cdc23 and Apc5.","evidence":"Mass spectrometry of purified yeast APC, in vitro co-translation interaction assay, and genetic deletion analysis in S. cerevisiae","pmids":["12609981"],"confidence":"High","gaps":["No structural information on how Swm1 integrates into the APC/C architecture","No direct evidence for the human orthologue at this stage"]},{"year":2004,"claim":"The functional requirement for Swm1/Apc13 was defined: it stabilizes TPR subunits Cdc16 and Cdc27 within the APC/C, and its absence abolishes ubiquitin ligase activity, explaining the cell cycle delays seen in swm1Δ cells.","evidence":"Genetic deletion, in vitro ubiquitin ligase assay, co-immunoprecipitation, and cross-species complementation using human and fission yeast homologues in budding yeast","pmids":["15060174"],"confidence":"High","gaps":["Molecular contacts mediating TPR subunit stabilization were undefined","Whether the human protein was an endogenous APC/C subunit in human cells remained unproven"]},{"year":2010,"claim":"APC13 was confirmed as a bona fide subunit of the human APC/C, closing the gap between yeast genetics and human cell biology and demonstrating its requirement for proper chromosome segregation.","evidence":"Tandem-affinity purification–mass spectrometry, fluorescence microscopy localization, and RNAi phenotypic screening in human cells","pmids":["20360068"],"confidence":"High","gaps":["Precise position of APC13 within the ~1.2 MDa human APC/C structure was unknown","Mechanism by which APC13 supports chromosome segregation was not molecularly resolved"]},{"year":2014,"claim":"Structural studies placed APC13 within the Arc Lamp sub-complex of the human APC/C alongside APC16 and CDC26, defining its three-dimensional context and how it interfaces with the TPR lobe.","evidence":"Cryo-EM at 7.4 Å resolution and X-ray crystallography of APC3–APC16 complexes with biochemical mapping","pmids":["25043029","25490258"],"confidence":"High","gaps":["Atomic-resolution contacts of APC13 with neighbouring subunits were not fully resolved","No disease-associated mutations had been linked to APC13"]},{"year":2015,"claim":"Atomic-resolution cryo-EM structures of APC/C–coactivator complexes confirmed and refined the Arc Lamp placement of APC13, completing the architectural picture of how the full APC/C is assembled and regulated.","evidence":"Near-atomic cryo-EM with mutagenesis-based functional validation","pmids":["26083744"],"confidence":"High","gaps":["Physiological consequences of APC13 loss in mammalian tissues remained untested","Whether APC13 has tissue-specific roles was unknown"]},{"year":2024,"claim":"A new regulatory axis was revealed: the cancer-associated SF3B1-K700E splicing-factor mutation reduces ANAPC13 protein by aberrant 5′ UTR splicing, linking APC/C function to splicing fidelity and demonstrating that ANAPC13 deficiency impairs Treg differentiation and immune homeostasis.","evidence":"Conditional SF3B1-K700E knock-in mouse model, RNA splicing analysis, adoptive transfer colitis assay, and forced ANAPC13 re-expression rescue","pmids":["39303038"],"confidence":"High","gaps":["Whether ANAPC13 reduction affects APC/C substrate repertoire in T cells is unresolved","Other SF3B1-dependent APC/C subunit transcripts have not been examined"]},{"year":2025,"claim":"Compound heterozygous ANAPC13 mutations were linked to female infertility, with in vitro evidence of aberrant protein localization, providing the first patient-level genetic evidence that APC13 integrity is required for human oocyte function.","evidence":"Whole-exome sequencing, structural modelling, and in vitro localization assay in cell lines","pmids":["40238067"],"confidence":"Medium","gaps":["Structural modelling is computational and awaits experimental confirmation of disrupted contacts","Single patient study without independent replication cohort","No direct APC/C activity measurement in patient-derived cells"]},{"year":2026,"claim":"Biallelic ANAPC13 mutations were causally linked to oocyte metaphase I arrest through a knock-in mouse model and proteomic analysis, establishing that APC13 is required for the metaphase I–anaphase I transition via APC/C-dependent substrate degradation rather than spindle assembly checkpoint regulation.","evidence":"Knock-in mouse model, in vitro maturation, oocyte proteomics, co-IP subunit interaction assay, and mRNA microinjection rescue (49% polar body extrusion recovery)","pmids":["41997520"],"confidence":"High","gaps":["Incomplete rescue (~49%) suggests additional factors or irreversible damage from early APC/C disruption","Identity of the specific APC/C substrates whose failed degradation causes metaphase I arrest was not determined","Whether ANAPC13 mutations affect male meiosis is untested"]},{"year":null,"claim":"The specific APC/C substrates whose degradation is impaired by ANAPC13 loss in oocytes and T cells remain unidentified, and whether ANAPC13 has APC/C-independent functions has not been addressed.","evidence":"","pmids":[],"confidence":"Low","gaps":["No substrate-level resolution of APC/C targets affected by ANAPC13 deficiency","No study has tested APC/C-independent roles for ANAPC13","Tissue-specific essentiality beyond oocytes and T cells is unexplored"]}],"mechanism_profile":{"molecular_activity":[{"term_id":"GO:0140096","term_label":"catalytic activity, acting on a protein","supporting_discovery_ids":[0,1,8]},{"term_id":"GO:0005198","term_label":"structural molecule activity","supporting_discovery_ids":[0,2,3,4]}],"localization":[{"term_id":"GO:0005634","term_label":"nucleus","supporting_discovery_ids":[5]},{"term_id":"GO:0005829","term_label":"cytosol","supporting_discovery_ids":[5]}],"pathway":[{"term_id":"R-HSA-1640170","term_label":"Cell Cycle","supporting_discovery_ids":[0,1,5,8]},{"term_id":"R-HSA-392499","term_label":"Metabolism of proteins","supporting_discovery_ids":[0,8]},{"term_id":"R-HSA-168256","term_label":"Immune System","supporting_discovery_ids":[6]},{"term_id":"R-HSA-1474165","term_label":"Reproduction","supporting_discovery_ids":[7,8]}],"complexes":["APC/C"],"partners":["CDC16","CDC27","CDC23","APC5","APC16","CDC26","APC7"],"other_free_text":[]},"mechanistic_narrative":"ANAPC13 (Swm1/Apc13) is a small, evolutionarily conserved core subunit of the anaphase-promoting complex/cyclosome (APC/C) E3 ubiquitin ligase that is essential for maintaining complex integrity and cell cycle progression. It resides in the Arc Lamp sub-structure of the APC/C alongside APC16 and CDC26, and promotes the stable association of the TPR-repeat subunits CDC16 (APC6) and CDC27 (APC3) with the complex; loss of ANAPC13 causes their dissociation and abolishes APC/C ubiquitin ligase activity [PMID:15060174, PMID:25043029, PMID:26083744]. ANAPC13 expression is regulated post-transcriptionally by SF3B1-dependent splicing, and its reduction impairs regulatory T cell differentiation, while biallelic loss-of-function mutations cause human oocyte maturation arrest at metaphase I [PMID:39303038, PMID:41997520]. Biallelic ANAPC13 mutations have been identified as a cause of female infertility due to oocyte maturation failure, with partial rescue by wild-type mRNA microinjection confirming a causal role [PMID:41997520, PMID:40238067]."},"prefetch_data":{"uniprot":{"accession":"Q9BS18","full_name":"Anaphase-promoting complex subunit 13","aliases":["Cyclosome subunit 13"],"length_aa":74,"mass_kda":8.5,"function":"Component of the anaphase promoting complex/cyclosome (APC/C), a cell cycle-regulated E3 ubiquitin ligase that controls progression through mitosis and the G1 phase of the cell cycle (PubMed:15060174, PubMed:18485873). The APC/C complex acts by mediating ubiquitination and subsequent degradation of target proteins: it mainly mediates the formation of 'Lys-11'-linked polyubiquitin chains and, to a lower extent, the formation of 'Lys-48'- and 'Lys-63'-linked polyubiquitin chains (PubMed:15060174, PubMed:18485873). The APC/C complex catalyzes assembly of branched 'Lys-11'-/'Lys-48'-linked branched ubiquitin chains on target proteins (PubMed:29033132)","subcellular_location":"Nucleus","url":"https://www.uniprot.org/uniprotkb/Q9BS18/entry"},"depmap":{"release":"DepMap","has_data":true,"is_common_essential":true,"resolved_as":"","url":"https://depmap.org/portal/gene/ANAPC13","classification":"Common Essential","n_dependent_lines":851,"n_total_lines":1208,"dependency_fraction":0.7044701986754967},"opencell":{"profiled":false,"resolved_as":"","ensg_id":"","cell_line_id":"","localizations":[],"interactors":[{"gene":"ANAPC16","stoichiometry":10.0},{"gene":"ANAPC2","stoichiometry":10.0},{"gene":"CDC16","stoichiometry":10.0},{"gene":"CDC23","stoichiometry":10.0},{"gene":"CDC26","stoichiometry":10.0},{"gene":"ANAPC4","stoichiometry":4.0}],"url":"https://opencell.sf.czbiohub.org/search/ANAPC13","total_profiled":1310},"omim":[{"mim_id":"614484","title":"ANAPHASE-PROMOTING COMPLEX, SUBUNIT 13; ANAPC13","url":"https://www.omim.org/entry/614484"}],"hpa":{"profiled":true,"resolved_as":"","reliability":"Approved","locations":[{"location":"Mitochondria","reliability":"Approved"},{"location":"Cytosol","reliability":"Approved"}],"tissue_specificity":"Low tissue specificity","tissue_distribution":"Detected in all","driving_tissues":[],"url":"https://www.proteinatlas.org/search/ANAPC13"},"hgnc":{"alias_symbol":["SWM1","APC13","DKFZP566D193"],"prev_symbol":[]},"alphafold":{"accession":"Q9BS18","domains":[{"cath_id":"-","chopping":"32-74","consensus_level":"medium","plddt":67.6502,"start":32,"end":74}],"viewer_url":"https://alphafold.ebi.ac.uk/entry/Q9BS18","model_url":"https://alphafold.ebi.ac.uk/files/AF-Q9BS18-F1-model_v6.cif","pae_url":"https://alphafold.ebi.ac.uk/files/AF-Q9BS18-F1-predicted_aligned_error_v6.png","plddt_mean":73.56},"mouse_models":{"mgi_url":"https://www.informatics.jax.org/marker/summary?nomen=ANAPC13","jax_strain_url":"https://www.jax.org/strain/search?query=ANAPC13"},"sequence":{"accession":"Q9BS18","fasta_url":"https://rest.uniprot.org/uniprotkb/Q9BS18.fasta","uniprot_url":"https://www.uniprot.org/uniprotkb/Q9BS18/entry","alphafold_viewer_url":"https://alphafold.ebi.ac.uk/entry/Q9BS18"}},"corpus_meta":[{"pmid":"21441434","id":"PMC_21441434","title":"The 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Human and fission yeast homologues complement the budding yeast swm1Δ phenotype and associate with APC/C subunits.\",\n      \"method\": \"Genetic deletion, in vitro ubiquitin ligase assay, co-immunoprecipitation, complementation assays, mass spectrometry\",\n      \"journal\": \"Molecular and cellular biology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1-2 — in vitro ubiquitin ligase assay + reciprocal co-IP + genetic complementation across species; moderate-strong evidence\",\n      \"pmids\": [\"15060174\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2003,\n      \"finding\": \"Swm1 (Apc13) is a core, constitutive subunit of the yeast APC/C present throughout the cell cycle (G1, S, M, and meiosis), identified by mass spectrometry of highly purified APC preparations; Swm1 interacts with Cdc23 and Apc5 in an in vitro transcription/translation system. Deletion of SWM1 causes slow growth and G2/M accumulation consistent with an APC defect.\",\n      \"method\": \"Mass spectrometry, co-purification, epitope-tag co-immunoprecipitation, in vitro transcription/translation interaction assay, cell cycle analysis\",\n      \"journal\": \"The Journal of biological chemistry\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1-2 — MS identification + reciprocal co-IP + in vitro interaction + cell cycle phenotype; replicated by independent lab (PMID:15060174)\",\n      \"pmids\": [\"12609981\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 1999,\n      \"finding\": \"Swm1p is a nuclear protein required for the late sporulation program in S. cerevisiae, specifically for spore wall assembly; swm1Δ diploids complete meiotic divisions but fail to produce mature asci and show altered expression of mid-late and late sporulation genes. Epistasis analysis indicates Swm1p acts in a pathway distinct from the Sps1p-Smk1p MAP kinase cascade.\",\n      \"method\": \"Genetic deletion, epistasis analysis, gene expression analysis, electron microscopy, sensitivity assays\",\n      \"journal\": \"Molecular and cellular biology\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 — genetic epistasis + localization + specific phenotypic readout; single lab\",\n      \"pmids\": [\"10022899\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2004,\n      \"finding\": \"Swm1p (yeast APC13 ortholog), as an APC/C subunit, is required for cell wall integrity during growth at high temperature; swm1 mutants show reduced glucan synthase activity (7-fold), delocalized chitin deposition, and cell lysis, with the chitin excess dependent on chitin synthase III (Chs3p) as shown by swm1 chs3 double mutant analysis.\",\n      \"method\": \"Electron microscopy, glucan synthase activity assay, chitin content measurement, double-mutant analysis\",\n      \"journal\": \"FEMS microbiology letters\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2 — enzymatic activity assay + double-mutant epistasis + EM; single lab, single study\",\n      \"pmids\": [\"15135545\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2014,\n      \"finding\": \"APC13 is a component of the 'Arc Lamp' subcomplex of the human APC/C; crystal structures of APC3 alone and in complex with the C-terminal domain of APC16 show how APC16 (which recruits APC7 to APC3) interacts asymmetrically with the symmetric APC3 dimer, providing structural context for APC13's location within the Arc Lamp alongside APC16 and CDC26.\",\n      \"method\": \"Crystal structure determination, biochemical mapping, mutagenesis, in vitro ubiquitination assay\",\n      \"journal\": \"Journal of molecular biology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1 — crystal structure with functional validation by mutagenesis and ubiquitination assays\",\n      \"pmids\": [\"25490258\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2024,\n      \"finding\": \"The cancer-associated SF3B1-K700E mutation causes aberrant splicing of ANAPC13, inserting a 231-bp fragment into the 5' UTR and reducing ANAPC13 protein expression in regulatory T cells (Tregs); reduced ANAPC13 impairs Treg differentiation and inhibitory function, and forced re-expression of ANAPC13 restores Treg differentiation and ability to prevent adoptive transfer colitis.\",\n      \"method\": \"Conditional knock-in mouse model, RNA splicing analysis, adoptive transfer colitis assay, forced expression rescue experiment, flow cytometry\",\n      \"journal\": \"Science advances\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 — mechanistic splicing analysis + loss-of-function + rescue in vivo with defined functional readout; single lab but multiple orthogonal methods\",\n      \"pmids\": [\"39303038\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2025,\n      \"finding\": \"Compound heterozygous missense variants in APC13 (c.C6A and c.116_126del) cause female infertility with oocyte/early embryo defects; structural modeling indicates these mutations alter chemical bonds with other APC/C subunits, and in vitro experiments show the missense mutation in APC13 results in aberrant cellular localization patterns.\",\n      \"method\": \"Whole-exome sequencing, structural modeling, in vitro cell-based localization assay\",\n      \"journal\": \"Journal of assisted reproduction and genetics\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2-3 — human genetics + structural modeling + in vitro localization; single study, limited functional follow-up\",\n      \"pmids\": [\"40238067\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2026,\n      \"finding\": \"Biallelic ANAPC13 mutations (p.D2E and p.L24R) cause oocyte maturation arrest at metaphase I in humans and in a knock-in mouse model; mutant ANAPC13 disrupts APC/C function by causing abnormal subunit interactions, impairing the protein composition remodeling during the metaphase I-to-anaphase I transition without altering spindle assembly checkpoint dynamics. Microinjection of Anapc13 mRNA partially rescues first polar body extrusion (~49%).\",\n      \"method\": \"Whole-exome sequencing, knock-in mouse model, in vitro maturation assay, proteomic analysis of human oocytes, molecular interaction studies with cell lines and plasmids, mRNA microinjection rescue\",\n      \"journal\": \"American journal of obstetrics and gynecology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1-2 — human mutations + mouse knock-in recapitulation + proteomic mechanistic analysis + mRNA rescue; multiple orthogonal methods in single study\",\n      \"pmids\": [\"41997520\"],\n      \"is_preprint\": false\n    }\n  ],\n  \"current_model\": \"ANAPC13 (Swm1/Apc13) is a small, evolutionarily conserved core subunit of the APC/C E3 ubiquitin ligase that stabilizes the association of essential TPR subunits (Cdc16/APC6 and Cdc27/APC8) within the Arc Lamp subcomplex, is required for APC/C ubiquitin ligase activity and timely cell cycle progression, and plays a critical role in oocyte meiotic maturation by maintaining proper APC/C subunit interactions and protein composition remodeling at the metaphase I-to-anaphase I transition; its expression is also regulated post-transcriptionally by SF3B1-dependent splicing in immune cells, where it supports regulatory T cell differentiation and function.\"\n}\n```\n\n--- ROUND 2 (R2/R3, augmented corpus) ---\n\n```json\n{\n  \"discoveries\": [\n    {\n      \"year\": 2004,\n      \"finding\": \"Swm1/Apc13 is an evolutionarily conserved subunit of the APC/C that promotes the stable association of the essential TPR subunits Cdc16 and Cdc27 with the complex; without Swm1/Apc13, Cdc16 and Cdc27 dissociate from the APC/C, abolishing ubiquitin ligase activity in vitro and causing delayed execution of APC/C-dependent cell cycle events in vivo. Human and fission yeast homologues associate with APC/C subunits and complement the yeast swm1Δ deletion phenotype.\",\n      \"method\": \"Genetic deletion, in vitro ubiquitin ligase assay, co-immunoprecipitation, cross-species complementation\",\n      \"journal\": \"Molecular and cellular biology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1–2 — multiple orthogonal methods (co-IP, in vitro reconstitution, genetic rescue) in a single study with clear functional readout\",\n      \"pmids\": [\"15060174\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2003,\n      \"finding\": \"Swm1 (Apc13) was identified as a constitutive core subunit of the yeast APC by mass spectrometry of highly purified APC preparations; it is present throughout the cell cycle (G1, S, M) and in meiotic cells. In vitro transcription/translation showed Swm1 interacts directly with Cdc23 and Apc5. Deletion of SWM1 causes slow growth and accumulation of G2/M cells, consistent with an APC defect.\",\n      \"method\": \"Mass spectrometry of purified APC, co-immunoprecipitation with epitope-tagged subunits, in vitro co-translation interaction assay, genetic analysis of deletion strains\",\n      \"journal\": \"The Journal of biological chemistry\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1–2 — biochemical identification by MS, reciprocal pull-down, in vitro interaction, and genetic phenotype in one study\",\n      \"pmids\": [\"12609981\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2014,\n      \"finding\": \"Cryo-EM reconstruction of the human APC/C at 7.4 Å resolution placed APC13 within the 'Arc Lamp' subcomplex alongside APC16 and CDC26, defining its position within the overall 1.2 MDa assembly alongside all 20 APC/C subunits.\",\n      \"method\": \"Cryo-electron microscopy reconstruction\",\n      \"journal\": \"Nature\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1 — near-atomic cryo-EM structure of the full APC/C complex with all subunits positioned\",\n      \"pmids\": [\"25043029\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2015,\n      \"finding\": \"Atomic-resolution cryo-EM structures of APC/C-coactivator complexes confirm APC13 as part of the Arc Lamp sub-structure; the structures define the architecture of all APC/C subunits and explain how Emi1 mediates inhibition.\",\n      \"method\": \"Cryo-electron microscopy at near-atomic resolution, biochemical validation\",\n      \"journal\": \"Nature\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1 — atomic cryo-EM with mutagenesis-based functional validation\",\n      \"pmids\": [\"26083744\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2014,\n      \"finding\": \"Crystal structures of human APC3 alone and in complex with the C-terminal domain of APC16 revealed that APC13 is part of the Arc Lamp alongside APC16 and APC7; the data show that APC16 bridges APC13 context within the TPR sub-complex and recruits APC7 to APC3.\",\n      \"method\": \"X-ray crystallography, biochemical mapping, in vitro ubiquitination assay\",\n      \"journal\": \"Journal of molecular biology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1 — crystal structure plus biochemical functional assays\",\n      \"pmids\": [\"25490258\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2010,\n      \"finding\": \"Tandem-affinity purification-mass spectrometry and localization studies in human cells identified APC13 as a previously unknown, evolutionarily conserved subunit of the human anaphase-promoting complex essential for chromosome segregation.\",\n      \"method\": \"TAP-MS, protein localization by fluorescence microscopy, RNAi phenotypic screening\",\n      \"journal\": \"Science\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 — large-scale TAP-MS with functional validation by RNAi and localization\",\n      \"pmids\": [\"20360068\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2024,\n      \"finding\": \"The cancer-associated SF3B1-K700E mutation causes aberrant splicing of ANAPC13 pre-mRNA, inserting a 231-bp fragment into the 5′ UTR and reducing ANAPC13 protein expression. Reduced ANAPC13 impairs regulatory T cell (Treg) differentiation and inhibitory function; forced re-expression of ANAPC13 restores Treg function and prevents adoptive transfer colitis.\",\n      \"method\": \"Conditional knock-in mouse model, RNA splicing analysis, adoptive transfer colitis assay, rescue by forced ANAPC13 expression\",\n      \"journal\": \"Science advances\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 2 — genetic mouse model with mechanistic splicing analysis and functional rescue experiment\",\n      \"pmids\": [\"39303038\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2025,\n      \"finding\": \"Compound heterozygous missense mutations in APC13 (c.C6A and c.116_126del) were identified in an infertile female patient; structural modelling predicted disrupted chemical bonds with other APC/C subunits, and in vitro experiments demonstrated aberrant cellular localization of the mutant APC13 protein, implicating correct APC13 localization as necessary for APC/C-dependent female fertility.\",\n      \"method\": \"Whole-exome sequencing, structural modelling, in vitro localization assay in cell lines\",\n      \"journal\": \"Journal of assisted reproduction and genetics\",\n      \"confidence\": \"Medium\",\n      \"confidence_rationale\": \"Tier 2–3 — patient genetics with in vitro localization assay; structural modelling is computational\",\n      \"pmids\": [\"40238067\"],\n      \"is_preprint\": false\n    },\n    {\n      \"year\": 2026,\n      \"finding\": \"Biallelic ANAPC13 mutations (p.D2E and p.L24R) cause oocyte maturation arrest at metaphase I in humans and in a knock-in mouse model. Mutant ANAPC13 disrupts APC/C subunit interactions, impairs APC/C ubiquitin ligase function (without altering spindle assembly checkpoint dynamics), and leads to abnormal protein composition during the metaphase I-to-anaphase I transition. Microinjection of wild-type Anapc13 mRNA partially rescues polar body extrusion (49%).\",\n      \"method\": \"Whole-exome sequencing, knock-in mouse model, in vitro maturation assay, proteomic analysis of oocytes, co-IP/subunit interaction assay, mRNA microinjection rescue\",\n      \"journal\": \"American journal of obstetrics and gynecology\",\n      \"confidence\": \"High\",\n      \"confidence_rationale\": \"Tier 1–2 — multiple orthogonal methods including animal model, proteomics, molecular interaction assay, and mRNA rescue in one study\",\n      \"pmids\": [\"41997520\"],\n      \"is_preprint\": false\n    }\n  ],\n  \"current_model\": \"ANAPC13 (Swm1/Apc13) is a small, evolutionarily conserved core subunit of the APC/C E3 ubiquitin ligase that resides in the Arc Lamp sub-complex and is required for stable association of the TPR subunits CDC16 and CDC27 with the complex, thereby maintaining APC/C ubiquitin ligase activity, proper cell cycle progression, and—in humans—oocyte maturation; its expression is regulated post-transcriptionally by SF3B1-dependent splicing, and loss of ANAPC13 impairs both APC/C function and regulatory T cell differentiation.\"\n}\n```","stage2_raw":"```json\n{\n  \"mechanistic_narrative\": \"ANAPC13 (Swm1/Apc13) is a small, evolutionarily conserved core subunit of the anaphase-promoting complex/cyclosome (APC/C) E3 ubiquitin ligase, where it stabilizes the association of essential TPR-repeat subunits (Cdc16/APC6 and Cdc27/APC3) within the Arc Lamp subcomplex and is required for APC/C ubiquitin ligase activity and timely cell cycle progression [PMID:12609981, PMID:15060174, PMID:25490258]. ANAPC13 is present constitutively throughout the cell cycle and during meiosis; biallelic loss-of-function mutations cause oocyte maturation arrest at metaphase I by disrupting APC/C subunit interactions and impairing protein composition remodeling at the metaphase I-to-anaphase I transition, establishing ANAPC13 as a cause of female infertility [PMID:41997520, PMID:40238067]. In immune cells, SF3B1-dependent aberrant splicing reduces ANAPC13 protein expression and impairs regulatory T cell differentiation and suppressive function [PMID:39303038]. In yeast, loss of Swm1 additionally compromises sporulation, cell wall integrity, and glucan synthase activity, reflecting broader APC/C-dependent roles beyond mitotic progression [PMID:10022899, PMID:15135545].\",\n  \"teleology\": [\n    {\n      \"year\": 1999,\n      \"claim\": \"Before its connection to the APC/C was known, SWM1 was identified as a nuclear gene required for late sporulation and spore wall assembly in yeast, establishing its first biological role in meiotic development.\",\n      \"evidence\": \"Genetic deletion, epistasis analysis, and electron microscopy in S. cerevisiae diploids\",\n      \"pmids\": [\"10022899\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"Molecular mechanism linking Swm1 to spore wall gene expression was unresolved\",\n        \"No physical interaction partners identified\",\n        \"Single lab study\"\n      ]\n    },\n    {\n      \"year\": 2003,\n      \"claim\": \"Swm1/Apc13 was identified as a constitutive core subunit of the APC/C present throughout the cell cycle, reframing its sporulation role as part of a ubiquitin ligase complex; in vitro interaction mapping placed it in contact with Cdc23 and Apc5.\",\n      \"evidence\": \"Mass spectrometry of purified APC/C, co-immunoprecipitation, in vitro transcription/translation interaction assay, cell cycle analysis in yeast\",\n      \"pmids\": [\"12609981\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Direct contribution to APC/C catalytic activity was not yet measured\",\n        \"Human ortholog function was not demonstrated\"\n      ]\n    },\n    {\n      \"year\": 2004,\n      \"claim\": \"Swm1/Apc13 was shown to stabilize the association of essential TPR subunits Cdc16 and Cdc27 with the APC/C and to be required for ubiquitin ligase activity in vitro; cross-species complementation by human and fission yeast orthologs demonstrated evolutionary conservation of this function.\",\n      \"evidence\": \"In vitro ubiquitin ligase assay, co-immunoprecipitation, genetic complementation in budding yeast with human and S. pombe homologs\",\n      \"pmids\": [\"15060174\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Structural basis for how Apc13 stabilizes TPR subunit association was unknown\",\n        \"Role in mammalian cells not directly tested in vivo\"\n      ]\n    },\n    {\n      \"year\": 2004,\n      \"claim\": \"Loss of Swm1 was found to compromise cell wall integrity via reduced glucan synthase activity and delocalized chitin deposition, extending APC/C-dependent phenotypes beyond canonical cell cycle control.\",\n      \"evidence\": \"Glucan synthase activity assay, chitin content measurement, swm1 chs3 double-mutant epistasis, electron microscopy in yeast\",\n      \"pmids\": [\"15135545\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"Whether cell wall defects reflect direct APC/C substrate targeting or indirect effects was unclear\",\n        \"Single lab, single study\"\n      ]\n    },\n    {\n      \"year\": 2014,\n      \"claim\": \"Structural analysis placed APC13 within the Arc Lamp subcomplex of the human APC/C alongside APC16 and CDC26, providing the first atomic-resolution context for its architectural role in bridging TPR subunits.\",\n      \"evidence\": \"Crystal structure of APC3 and APC3–APC16 complex, mutagenesis, in vitro ubiquitination assay\",\n      \"pmids\": [\"25490258\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"No high-resolution structure of APC13 itself was resolved\",\n        \"Mechanism by which APC13 stabilizes Cdc16/Cdc27 at atomic level remained unresolved\"\n      ]\n    },\n    {\n      \"year\": 2024,\n      \"claim\": \"SF3B1-K700E mutation was shown to cause aberrant ANAPC13 splicing, reducing its protein levels in regulatory T cells, which impaired Treg differentiation and suppressive function — revealing a post-transcriptional regulatory axis and an immune-cell-specific consequence of ANAPC13 deficiency.\",\n      \"evidence\": \"SF3B1-K700E conditional knock-in mouse model, RNA splicing analysis, adoptive transfer colitis assay, forced expression rescue, flow cytometry\",\n      \"pmids\": [\"39303038\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Which APC/C substrates are dysregulated in Tregs upon ANAPC13 reduction was not identified\",\n        \"Whether other SF3B1-dependent mis-spliced targets contribute to the Treg phenotype was not fully resolved\"\n      ]\n    },\n    {\n      \"year\": 2025,\n      \"claim\": \"Biallelic ANAPC13 mutations were identified as a cause of human female infertility through oocyte maturation arrest at metaphase I; a knock-in mouse model recapitulated the phenotype and proteomic analysis revealed that mutant ANAPC13 disrupts APC/C subunit interactions and impairs protein composition remodeling during the MI-to-AI transition.\",\n      \"evidence\": \"Whole-exome sequencing, knock-in mouse model, in vitro maturation, oocyte proteomics, mRNA microinjection rescue (~49% polar body extrusion)\",\n      \"pmids\": [\"40238067\", \"41997520\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Specific APC/C substrates whose degradation is blocked at MI were not identified\",\n        \"Partial rescue (~49%) suggests additional factors contributing to the phenotype\",\n        \"Whether ANAPC13 mutations affect male fertility is unknown\"\n      ]\n    },\n    {\n      \"year\": null,\n      \"claim\": \"The precise structural mechanism by which ANAPC13 stabilizes TPR subunit interactions within the APC/C remains unresolved, and the specific APC/C substrates whose degradation depends on ANAPC13 integrity during meiosis and Treg differentiation have not been identified.\",\n      \"evidence\": \"\",\n      \"pmids\": [],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"No high-resolution structure of ANAPC13 within the complete APC/C\",\n        \"Meiosis-specific and Treg-specific APC/C substrates affected by ANAPC13 deficiency are unknown\",\n        \"Genotype–phenotype relationships for different ANAPC13 mutations not systematically characterized\"\n      ]\n    }\n  ],\n  \"mechanism_profile\": {\n    \"molecular_activity\": [\n      {\n        \"term_id\": \"GO:0140096\",\n        \"supporting_discovery_ids\": [0, 1, 7]\n      }\n    ],\n    \"localization\": [\n      {\n        \"term_id\": \"GO:0005634\",\n        \"supporting_discovery_ids\": [2, 6]\n      }\n    ],\n    \"pathway\": [\n      {\n        \"term_id\": \"R-HSA-1640170\",\n        \"supporting_discovery_ids\": [0, 1, 7]\n      },\n      {\n        \"term_id\": \"R-HSA-392499\",\n        \"supporting_discovery_ids\": [0, 4]\n      },\n      {\n        \"term_id\": \"R-HSA-168256\",\n        \"supporting_discovery_ids\": [5]\n      },\n      {\n        \"term_id\": \"R-HSA-1474165\",\n        \"supporting_discovery_ids\": [6, 7]\n      }\n    ],\n    \"complexes\": [\n      \"APC/C\"\n    ],\n    \"partners\": [\n      \"CDC16\",\n      \"CDC27\",\n      \"CDC23\",\n      \"APC5\",\n      \"APC16\",\n      \"CDC26\"\n    ],\n    \"other_free_text\": []\n  }\n}\n```\n\n--- ROUND 2 (R2/R3, augmented corpus) ---\n\n```json\n{\n  \"mechanistic_narrative\": \"ANAPC13 (Swm1/Apc13) is a small, evolutionarily conserved core subunit of the anaphase-promoting complex/cyclosome (APC/C) E3 ubiquitin ligase that is essential for maintaining complex integrity and cell cycle progression. It resides in the Arc Lamp sub-structure of the APC/C alongside APC16 and CDC26, and promotes the stable association of the TPR-repeat subunits CDC16 (APC6) and CDC27 (APC3) with the complex; loss of ANAPC13 causes their dissociation and abolishes APC/C ubiquitin ligase activity [PMID:15060174, PMID:25043029, PMID:26083744]. ANAPC13 expression is regulated post-transcriptionally by SF3B1-dependent splicing, and its reduction impairs regulatory T cell differentiation, while biallelic loss-of-function mutations cause human oocyte maturation arrest at metaphase I [PMID:39303038, PMID:41997520]. Biallelic ANAPC13 mutations have been identified as a cause of female infertility due to oocyte maturation failure, with partial rescue by wild-type mRNA microinjection confirming a causal role [PMID:41997520, PMID:40238067].\",\n  \"teleology\": [\n    {\n      \"year\": 2003,\n      \"claim\": \"The identity of Swm1/Apc13 as a constitutive APC/C core subunit was established, resolving whether it was a stoichiometric component or a transient interactor, and revealing its direct contacts with Cdc23 and Apc5.\",\n      \"evidence\": \"Mass spectrometry of purified yeast APC, in vitro co-translation interaction assay, and genetic deletion analysis in S. cerevisiae\",\n      \"pmids\": [\"12609981\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"No structural information on how Swm1 integrates into the APC/C architecture\",\n        \"No direct evidence for the human orthologue at this stage\"\n      ]\n    },\n    {\n      \"year\": 2004,\n      \"claim\": \"The functional requirement for Swm1/Apc13 was defined: it stabilizes TPR subunits Cdc16 and Cdc27 within the APC/C, and its absence abolishes ubiquitin ligase activity, explaining the cell cycle delays seen in swm1Δ cells.\",\n      \"evidence\": \"Genetic deletion, in vitro ubiquitin ligase assay, co-immunoprecipitation, and cross-species complementation using human and fission yeast homologues in budding yeast\",\n      \"pmids\": [\"15060174\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Molecular contacts mediating TPR subunit stabilization were undefined\",\n        \"Whether the human protein was an endogenous APC/C subunit in human cells remained unproven\"\n      ]\n    },\n    {\n      \"year\": 2010,\n      \"claim\": \"APC13 was confirmed as a bona fide subunit of the human APC/C, closing the gap between yeast genetics and human cell biology and demonstrating its requirement for proper chromosome segregation.\",\n      \"evidence\": \"Tandem-affinity purification–mass spectrometry, fluorescence microscopy localization, and RNAi phenotypic screening in human cells\",\n      \"pmids\": [\"20360068\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Precise position of APC13 within the ~1.2 MDa human APC/C structure was unknown\",\n        \"Mechanism by which APC13 supports chromosome segregation was not molecularly resolved\"\n      ]\n    },\n    {\n      \"year\": 2014,\n      \"claim\": \"Structural studies placed APC13 within the Arc Lamp sub-complex of the human APC/C alongside APC16 and CDC26, defining its three-dimensional context and how it interfaces with the TPR lobe.\",\n      \"evidence\": \"Cryo-EM at 7.4 Å resolution and X-ray crystallography of APC3–APC16 complexes with biochemical mapping\",\n      \"pmids\": [\"25043029\", \"25490258\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Atomic-resolution contacts of APC13 with neighbouring subunits were not fully resolved\",\n        \"No disease-associated mutations had been linked to APC13\"\n      ]\n    },\n    {\n      \"year\": 2015,\n      \"claim\": \"Atomic-resolution cryo-EM structures of APC/C–coactivator complexes confirmed and refined the Arc Lamp placement of APC13, completing the architectural picture of how the full APC/C is assembled and regulated.\",\n      \"evidence\": \"Near-atomic cryo-EM with mutagenesis-based functional validation\",\n      \"pmids\": [\"26083744\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Physiological consequences of APC13 loss in mammalian tissues remained untested\",\n        \"Whether APC13 has tissue-specific roles was unknown\"\n      ]\n    },\n    {\n      \"year\": 2024,\n      \"claim\": \"A new regulatory axis was revealed: the cancer-associated SF3B1-K700E splicing-factor mutation reduces ANAPC13 protein by aberrant 5′ UTR splicing, linking APC/C function to splicing fidelity and demonstrating that ANAPC13 deficiency impairs Treg differentiation and immune homeostasis.\",\n      \"evidence\": \"Conditional SF3B1-K700E knock-in mouse model, RNA splicing analysis, adoptive transfer colitis assay, and forced ANAPC13 re-expression rescue\",\n      \"pmids\": [\"39303038\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Whether ANAPC13 reduction affects APC/C substrate repertoire in T cells is unresolved\",\n        \"Other SF3B1-dependent APC/C subunit transcripts have not been examined\"\n      ]\n    },\n    {\n      \"year\": 2025,\n      \"claim\": \"Compound heterozygous ANAPC13 mutations were linked to female infertility, with in vitro evidence of aberrant protein localization, providing the first patient-level genetic evidence that APC13 integrity is required for human oocyte function.\",\n      \"evidence\": \"Whole-exome sequencing, structural modelling, and in vitro localization assay in cell lines\",\n      \"pmids\": [\"40238067\"],\n      \"confidence\": \"Medium\",\n      \"gaps\": [\n        \"Structural modelling is computational and awaits experimental confirmation of disrupted contacts\",\n        \"Single patient study without independent replication cohort\",\n        \"No direct APC/C activity measurement in patient-derived cells\"\n      ]\n    },\n    {\n      \"year\": 2026,\n      \"claim\": \"Biallelic ANAPC13 mutations were causally linked to oocyte metaphase I arrest through a knock-in mouse model and proteomic analysis, establishing that APC13 is required for the metaphase I–anaphase I transition via APC/C-dependent substrate degradation rather than spindle assembly checkpoint regulation.\",\n      \"evidence\": \"Knock-in mouse model, in vitro maturation, oocyte proteomics, co-IP subunit interaction assay, and mRNA microinjection rescue (49% polar body extrusion recovery)\",\n      \"pmids\": [\"41997520\"],\n      \"confidence\": \"High\",\n      \"gaps\": [\n        \"Incomplete rescue (~49%) suggests additional factors or irreversible damage from early APC/C disruption\",\n        \"Identity of the specific APC/C substrates whose failed degradation causes metaphase I arrest was not determined\",\n        \"Whether ANAPC13 mutations affect male meiosis is untested\"\n      ]\n    },\n    {\n      \"year\": null,\n      \"claim\": \"The specific APC/C substrates whose degradation is impaired by ANAPC13 loss in oocytes and T cells remain unidentified, and whether ANAPC13 has APC/C-independent functions has not been addressed.\",\n      \"evidence\": \"\",\n      \"pmids\": [],\n      \"confidence\": \"Low\",\n      \"gaps\": [\n        \"No substrate-level resolution of APC/C targets affected by ANAPC13 deficiency\",\n        \"No study has tested APC/C-independent roles for ANAPC13\",\n        \"Tissue-specific essentiality beyond oocytes and T cells is unexplored\"\n      ]\n    }\n  ],\n  \"mechanism_profile\": {\n    \"molecular_activity\": [\n      {\"term_id\": \"GO:0140096\", \"supporting_discovery_ids\": [0, 1, 8]},\n      {\"term_id\": \"GO:0005198\", \"supporting_discovery_ids\": [0, 2, 3, 4]}\n    ],\n    \"localization\": [\n      {\"term_id\": \"GO:0005634\", \"supporting_discovery_ids\": [5]},\n      {\"term_id\": \"GO:0005829\", \"supporting_discovery_ids\": [5]}\n    ],\n    \"pathway\": [\n      {\"term_id\": \"R-HSA-1640170\", \"supporting_discovery_ids\": [0, 1, 5, 8]},\n      {\"term_id\": \"R-HSA-392499\", \"supporting_discovery_ids\": [0, 8]},\n      {\"term_id\": \"R-HSA-168256\", \"supporting_discovery_ids\": [6]},\n      {\"term_id\": \"R-HSA-1474165\", \"supporting_discovery_ids\": [7, 8]}\n    ],\n    \"complexes\": [\n      \"APC/C\"\n    ],\n    \"partners\": [\n      \"CDC16\",\n      \"CDC27\",\n      \"CDC23\",\n      \"APC5\",\n      \"APC16\",\n      \"CDC26\",\n      \"APC7\"\n    ],\n    \"other_free_text\": []\n  }\n}\n```"}