BAG6

UniProt ID: P46379
Organism: Homo sapiens
Review Status: COMPLETE
📝 Provide Detailed Feedback

Gene Description

BAG6 is a multifunctional nucleo-cytoplasmic proteostasis factor that sits at the boundary between the GET pathway and ubiquitin-proteasome-associated quality control. In the cytosol, the BAG6/UBL4A/GET4 (BAT3) complex captures hydrophobic tail-anchored or otherwise mislocalized secretory proteins, promotes their handoff to TRC40/ASNA1 for post-translational delivery to the endoplasmic reticulum, and routes failed clients toward ubiquitin-proteasome degradation. BAG6 also acts as a holdase for retrotranslocated ERAD substrates and contributes to ER stress-induced pre-emptive quality control. Additional nuclear and extracellular roles, including p53 acetylation after DNA damage and exosomal NKp30 ligand activity, are supported in specific contexts but are not the core conserved proteostasis functions.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0036503 ERAD pathway
IBA
GO_REF:0000033
ACCEPT
Summary: Human BAG6 literature supports BAG6 complex function in ER-associated degradation (ERAD), so the phylogenetic propagation is directionally correct and consistent with the core proteostasis role.
Reason: Bag6 keeps retrotranslocated hydrophobic clients soluble and engaged with ERAD machinery, matching ERAD pathway membership.
Supporting Evidence:
file:human/BAG6/BAG6-notes.md
GO:0031593 polyubiquitin modification-dependent protein binding
IBA
GO_REF:0000033
REMOVE
Summary: BAG6 operates in ubiquitin-linked quality control, but the reviewed core papers support ligase/DUB recruitment and client triage rather than a well-demonstrated direct polyubiquitin-binding activity.
Reason: The conserved BAG6 literature does not establish polyubiquitin-selective binding as a defensible core molecular function.
GO:0051787 misfolded protein binding
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic propagation agrees with the human holdase literature: BAG6 preferentially captures aggregation-prone hydrophobic or misfolded clients during quality control.
Reason: This matches the experimentally supported BAG6 holdase role in ERAD and mislocalized-protein triage.
Supporting Evidence:
file:human/BAG6/BAG6-notes.md
GO:0071818 BAT3 complex
IBA
GO_REF:0000033
ACCEPT
Summary: The phylogenetic BAT3 complex annotation agrees with multiple human studies placing BAG6 in a heterotrimeric BAG6/UBL4A/GET4(TRC35) complex.
Reason: Complex membership is well supported and central to both GET-pathway and UPS-adjacent BAG6 functions.
Supporting Evidence:
GO:0001822 kidney development
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: Developmental annotations are plausible downstream consequences from mammalian loss-of-function studies, but they are not BAG6's core conserved proteostasis role.
Reason: Keep as contextual biology rather than a core molecular/process function.
GO:0006511 ubiquitin-dependent protein catabolic process
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: BAG6 helps channel selected clients toward ubiquitin-dependent degradation, but this broad process term loses the more informative GET/ERAD/pre-emptive-QC context.
Reason: Retain more specific proteostasis terms instead of this umbrella catabolic label.
GO:0007130 synaptonemal complex assembly
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: This reproductive annotation reflects secondary mammalian biology rather than BAG6's central proteostasis function.
Reason: Treat as contextual/non-core.
GO:0007283 spermatogenesis
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: This reproductive-process annotation is best treated as contextual because BAG6's conserved role is proteostasis triage, not gametogenesis machinery.
Reason: Keep as non-core.
GO:0007420 brain development
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: Any developmental nervous-system consequence is downstream/contextual rather than part of BAG6's main evolved role.
Reason: Keep as non-core.
GO:0030324 lung development
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: This developmental annotation reflects contextual organismal phenotypes rather than BAG6's core GET/UPS-boundary biology.
Reason: Keep as non-core.
GO:0030544 Hsp70 protein binding
IEA
GO_REF:0000107
REMOVE
Summary: This looks like over-propagation from BAG family naming. BAG6 is explicitly not a canonical BAG-domain Hsp70 nucleotide-exchange factor.
Reason: Structural work shows the BAG6 C terminus is a mock/noncanonical BAG-similar domain, so Hsp70 binding should not be assumed from family membership.
Supporting Evidence:
file:human/BAG6/BAG6-notes.md
GO:0031593 polyubiquitin modification-dependent protein binding
IEA
GO_REF:0000107
REMOVE
Summary: This broad automated propagation is not well grounded in the direct BAG6 mechanistic literature.
Reason: The reviewed papers support client triage with E3s and DUBs, not a specific polyubiquitin-binding activity.
GO:0032435 negative regulation of proteasomal ubiquitin-dependent protein catabolic process
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: This regulatory proteolysis term reflects secondary spermatogenic/HSPA2 biology rather than BAG6's central proteostasis role.
Reason: Keep as contextual/non-core.
GO:0042981 regulation of apoptotic process
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: Apoptosis-related effects exist in some contexts, but this broad automated term is not the best summary of BAG6 function.
Reason: Retain only as non-core context.
GO:0043161 proteasome-mediated ubiquitin-dependent protein catabolic process
IEA
GO_REF:0000107
MARK AS OVER ANNOTATED
Summary: BAG6 contributes to selected proteasomal quality-control routes, but this automated term is broader and less informative than the specific BAG6-supported processes.
Reason: Prefer ERAD, tail-anchored insertion, or ER-stress pre-emptive quality control terms.
GO:0045861 negative regulation of proteolysis
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: This negative-regulation term reflects context-specific stabilization biology rather than the main BAG6 proteostasis program.
Reason: Keep as non-core.
GO:0045995 regulation of embryonic development
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: Embryonic-development phenotypes are contextual downstream consequences and not the core conserved BAG6 role.
Reason: Keep as non-core.
GO:0050821 protein stabilization
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: Protein stabilization is context-specific for selected partners and not the best core descriptor of BAG6.
Reason: Keep as non-core.
GO:0070059 intrinsic apoptotic signaling pathway in response to endoplasmic reticulum stress
IEA
GO_REF:0000107
KEEP AS NON CORE
Summary: ER-stress apoptosis is a context-dependent consequence, not BAG6's principal conserved function.
Reason: Keep as non-core.
GO:0070628 proteasome binding
IEA
GO_REF:0000107
REMOVE
Summary: The reviewed BAG6 papers place clients en route to the proteasome but do not establish a clean direct proteasome-binding activity.
Reason: This term is too specific for the available evidence.
GO:0005515 protein binding
IPI
PMID:14667819
Analysis of a high-throughput yeast two-hybrid system and it...
REMOVE
Summary: The source interaction is too generic to retain as a useful BAG6 annotation.
Reason: GO:0005515 is uninformative here and BAG6 already has more specific mechanistically grounded annotations.
GO:0005515 protein binding
IPI
PMID:16189514
Towards a proteome-scale map of the human protein-protein in...
REMOVE
Summary: The source interaction is too generic to retain as a useful BAG6 annotation.
Reason: GO:0005515 is uninformative here and BAG6 already has more specific mechanistically grounded annotations.
GO:0005515 protein binding
IPI
PMID:18852879
Dendritic cells release HLA-B-associated transcript-3 positi...
REMOVE
Summary: The exosome/NK paper supports a contextual receptor-ligand role, not a useful standalone generic protein-binding annotation.
Reason: Retain the more specific immune-context terms instead.
GO:0005515 protein binding
IPI
PMID:21903422
Mapping a dynamic innate immunity protein interaction networ...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:22046132
The SARS-coronavirus-host interactome: identification of cyc...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:22807449
The stalk domain and the glycosylation status of the activat...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:25036637
A quantitative chaperone interaction network reveals the arc...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:25416956
A proteome-scale map of the human interactome network.
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:26496610
A human interactome in three quantitative dimensions organiz...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:31515488
Extensive disruption of protein interactions by genetic vari...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:33961781
Dual proteome-scale networks reveal cell-specific remodeling...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:35271311
OpenCell: Endogenous tagging for the cartography of human ce...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005576 extracellular region
IEA
GO_REF:0000044
KEEP AS NON CORE
Summary: Extracellular release is real but limited to stress/tumor exosome contexts rather than BAG6's main intracellular proteostasis role.
Reason: Keep as contextual, non-core localization.
GO:0005634 nucleus
IEA
GO_REF:0000120
ACCEPT
Summary: BAG6 is a bona fide nucleo-cytoplasmic protein, and nuclear localization is repeatedly observed in the literature and UniProt curation.
Reason: Nuclear localization is real even though the principal PN role is cytosolic.
GO:0005829 cytosol
IEA
GO_REF:0000120
ACCEPT
Summary: Cytosolic localization is central to BAG6 GET/ERAD/mislocalized-protein triage functions.
Reason: The core proteostasis role depends on cytosolic BAG6 complex localization.
GO:0005654 nucleoplasm
IDA
GO_REF:0000052
ACCEPT
Summary: Nucleoplasmic signal is compatible with BAG6's documented nuclear pool and DNA-damage-associated functions.
Reason: This is a defensible subnuclear localization.
GO:0005829 cytosol
IDA
GO_REF:0000052
ACCEPT
Summary: Independent localization evidence also places BAG6 in the cytosol.
Reason: This is fully consistent with the core proteostasis model.
GO:0005515 protein binding
IPI
PMID:40105103
Definition of the human mitochondrial TOM interactome reveal...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0140597 protein carrier chaperone
IDA
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
ACCEPT
Summary: This is one of the strongest BAG6 annotations. The 2011 Mol Cell study directly supports a holdase/carrier role for retrotranslocated hydrophobic clients.
Reason: Bag6 maintains aggregation-prone clients in an unfolded yet soluble state and helps deliver them within quality-control pathways.
Supporting Evidence:
file:human/BAG6/BAG6-deep-research-falcon.md
BAG6 is widely characterized as a chaperone/holdase that binds exposed hydrophobic segments
GO:0005829 cytosol
NAS
PMID:25535373
Bag6 complex contains a minimal tail-anchor-targeting module...
ACCEPT
Summary: Cytosolic localization is consistent with the tail-anchor-targeting complex architecture and broader BAG6 literature, even if this individual assertion is author statement-level.
Reason: The BAG6 complex acts in the cytosol before ER insertion or proteasomal routing.
GO:0006511 ubiquitin-dependent protein catabolic process
IDA
PMID:20676083
A ribosome-associating factor chaperones tail-anchored membr...
MODIFY
Summary: PMID:20676083 is about ribosome-associated capture and handoff of tail-anchored proteins to TRC40, not a generic ubiquitin-dependent catabolic process.
Reason: Replace with the specific GET-pathway process terms directly supported by the paper; the separate ribosome-binding molecular function is already captured by its own annotation.
GO:0006620 post-translational protein targeting to endoplasmic reticulum membrane
IDA
PMID:25535373
Bag6 complex contains a minimal tail-anchor-targeting module...
ACCEPT
Summary: This matches the PN GET-pathway mapping and the structural/biochemical literature on tail-anchor targeting.
Reason: BAG6 participates in post-translational delivery of tail-anchored proteins toward the ER membrane.
Supporting Evidence:
file:human/BAG6/BAG6-deep-research-falcon.md
BAG6 is embedded in this pathway as both a substrate-holding factor and a quality-control adaptor
GO:0031647 regulation of protein stability
IDA
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
MODIFY
Summary: The paper supports holdase/chaperone behavior rather than a broad generic regulation-of-stability process.
Reason: A molecular chaperone/carrier term is a more faithful representation of the evidence.
Proposed replacements: protein carrier chaperone
GO:0048018 receptor ligand activity
IDA
PMID:18852879
Dendritic cells release HLA-B-associated transcript-3 positi...
KEEP AS NON CORE
Summary: Exosomal BAG6 can act as an NKp30 ligand, but this immune signaling role is contextual and not the core conserved BAG6 function.
Reason: Keep as non-core context.
Supporting Evidence:
GO:0051132 NK T cell activation
IDA
PMID:18852879
Dendritic cells release HLA-B-associated transcript-3 positi...
MODIFY
Summary: The paper concerns NK-cell activation, not NK T-cell activation.
Reason: Replace with natural killer cell activation to match the actual experiment.
Proposed replacements: natural killer cell activation
GO:0036503 ERAD pathway
NAS
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
ACCEPT
Summary: Although this is an author statement-level annotation, the underlying study directly demonstrates a BAG6 holdase role that improves ERAD efficiency.
Reason: The evidence supports ERAD pathway participation.
Supporting Evidence:
file:human/BAG6/BAG6-deep-research-falcon.md
BAG6 helps route failed membrane/secretory protein biogenesis products and certain ERAD substrates into ubiquitin-proteasome degradation
GO:0005515 protein binding
IPI
PMID:25713138
Structure of a BAG6 (Bcl-2-associated athanogene 6)-Ubl4a (u...
REMOVE
Summary: The structural paper supports complex architecture and adaptor function, not a useful generic protein-binding annotation.
Reason: Retain the specific adaptor/core-complex annotations instead.
GO:0060090 molecular adaptor activity
EXP
PMID:25713138
Structure of a BAG6 (Bcl-2-associated athanogene 6)-Ubl4a (u...
ACCEPT
Summary: BAG6 organizes UBL4A and TRC35/GET4 and supports substrate handoff, which is well captured by molecular adaptor activity.
Reason: This term fits BAG6's bridge/scaffold role at the GET-pathway and cytosolic quality-control boundary.
Supporting Evidence:
file:human/BAG6/BAG6-deep-research-falcon.md
BAG6’s C-terminal region is a structural part of the substrate-loading complex
GO:0140677 molecular function activator activity
IEP
PMID:25713138
Structure of a BAG6 (Bcl-2-associated athanogene 6)-Ubl4a (u...
REMOVE
Summary: The cited paper does not establish BAG6 as a direct molecular-function activator of another enzyme or receptor.
Reason: This annotation overstates the structural/adaptor evidence.
GO:0005515 protein binding
IPI
PMID:27113755
UBQLN4 recognizes mislocalized transmembrane domain proteins...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0045995 regulation of embryonic development
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Orthology-based developmental regulation is plausible but not part of BAG6's core proteostasis identity.
Reason: Keep as non-core context.
GO:0005515 protein binding
IPI
PMID:29042515
Structural basis for regulation of the nucleo-cytoplasmic di...
REMOVE
Summary: This paper is informative for localization regulation, but GO:0005515 remains too generic.
Reason: Remove generic protein-binding carryover.
GO:0005634 nucleus
IDA
PMID:29042515
Structural basis for regulation of the nucleo-cytoplasmic di...
ACCEPT
Summary: This study directly addresses BAG6 nucleo-cytoplasmic partitioning and supports nuclear localization.
Reason: Nuclear BAG6 is real and regulated by TRC35/GET4 binding.
GO:0005829 cytosol
IDA
PMID:29042515
Structural basis for regulation of the nucleo-cytoplasmic di...
ACCEPT
Summary: This study directly supports cytosolic retention of BAG6 by TRC35/GET4.
Reason: Cytosolic BAG6 is the state associated with core proteostasis functions.
GO:0071816 tail-anchored membrane protein insertion into ER membrane
IDA
PMID:25535373
Bag6 complex contains a minimal tail-anchor-targeting module...
ACCEPT
Summary: The minimal BAG6 complex directly supports transfer of tail-anchored substrates into the TRC40/GET pathway.
Reason: This is core PN-supported biology.
Supporting Evidence:
GO:0005515 protein binding
IPI
PMID:26876100
Selective Binding of AIRAPL Tandem UIMs to Lys48-Linked Tri-...
REMOVE
Summary: This generic interaction term adds no useful BAG6-specific functional information.
Reason: Remove generic protein-binding carryover.
GO:0005515 protein binding
IPI
PMID:14960581
Ricin triggers apoptotic morphological changes through caspa...
REMOVE
Summary: The ricin paper is informative for apoptosis context, but GO:0005515 remains too generic.
Reason: Remove generic protein-binding carryover.
GO:0061857 endoplasmic reticulum stress-induced pre-emptive quality control
IMP
PMID:26565908
Pre-emptive Quality Control Protects the ER from Protein Ove...
ACCEPT
Summary: This is a specific stress-responsive proteostasis process that the paper directly supports.
Reason: Bag6 contributes to degradation of rerouted ER pre-emptive quality-control substrates under ER stress.
Supporting Evidence:
GO:0010498 proteasomal protein catabolic process
IMP
PMID:26565908
Pre-emptive Quality Control Protects the ER from Protein Ove...
MARK AS OVER ANNOTATED
Summary: The specific process shown in the paper is ER stress-induced pre-emptive quality control, not a generic proteasomal catabolic pathway in isolation.
Reason: Keep the more specific GO:0061857 annotation instead.
GO:0036503 ERAD pathway
IMP
PMID:26565908
Pre-emptive Quality Control Protects the ER from Protein Ove...
MODIFY
Summary: PMID:26565908 is about ER stress-induced pre-emptive quality control, which is related to but distinct from canonical ERAD.
Reason: Replace with the specific ER pQC term.
GO:0016020 membrane
HDA
PMID:19946888
Defining the membrane proteome of NK cells.
MARK AS OVER ANNOTATED
Summary: A broad membrane term is too imprecise for BAG6, which is mainly cytosolic/nuclear with regulated ER-membrane association in specific QC contexts.
Reason: If membrane association is retained, it should be ER-membrane-specific rather than generic membrane.
GO:0002429 immune response-activating cell surface receptor signaling pathway
IDA
PMID:18852879
Dendritic cells release HLA-B-associated transcript-3 positi...
KEEP AS NON CORE
Summary: This immune signaling annotation is supported in exosome/NK-cell contexts but is clearly contextual rather than core BAG6 biology.
Reason: Keep as non-core.
Supporting Evidence:
GO:0005102 signaling receptor binding
IPI
PMID:18852879
Dendritic cells release HLA-B-associated transcript-3 positi...
MODIFY
Summary: The immune paper is better captured by receptor ligand activity than a generic signaling receptor binding term.
Reason: Replace with receptor ligand activity.
Proposed replacements: receptor ligand activity
GO:0005634 nucleus
IDA
PMID:14960581
Ricin triggers apoptotic morphological changes through caspa...
ACCEPT
Summary: The ricin study also observed nuclear BAG6, consistent with BAG6's established nucleo-cytoplasmic distribution.
Reason: Nuclear localization is defensible.
GO:0006915 apoptotic process
IDA
PMID:14960581
Ricin triggers apoptotic morphological changes through caspa...
KEEP AS NON CORE
Summary: Ricin-induced apoptosis is a real contextual role but not BAG6's core conserved proteostasis function.
Reason: Keep as non-core.
GO:0030101 natural killer cell activation
IDA
PMID:18852879
Dendritic cells release HLA-B-associated transcript-3 positi...
KEEP AS NON CORE
Summary: Natural killer cell activation is experimentally supported for exosomal BAG6 but remains a contextual immune role.
Reason: Keep as non-core.
Supporting Evidence:
GO:0070062 extracellular exosome
IDA
PMID:18852879
Dendritic cells release HLA-B-associated transcript-3 positi...
KEEP AS NON CORE
Summary: Exosomal BAG6 is experimentally documented, but this is a conditional extracellular context rather than the main cellular location.
Reason: Keep as non-core localization.
Supporting Evidence:
GO:0036503 ERAD pathway
IDA
PMID:23129660
SGTA antagonizes BAG6-mediated protein triage.
MODIFY
Summary: This paper studies SGTA antagonism of BAG6-mediated triage of mislocalized proteins in the cytosol, not canonical ERAD.
Reason: Replace with the narrower protein-quality-control term that better fits cytosolic degradation of mislocalized hydrophobic clients.
GO:0036503 ERAD pathway
IDA
PMID:24981174
Cytosolic quality control of mislocalized proteins requires ...
MODIFY
Summary: PMID:24981174 defines RNF126-dependent cytosolic quality control of mislocalized proteins rather than general ERAD.
Reason: Replace with the more specific protein-quality-control term rather than the broader ubiquitin-dependent catabolic umbrella term.
GO:1904294 positive regulation of ERAD pathway
IMP
PMID:24424410
USP13 antagonizes gp78 to maintain functionality of a chaper...
MODIFY
Summary: BAG6 is part of ERAD-associated machinery; this paper mostly shows that USP13 preserves BAG6 complex function rather than BAG6 acting as an upstream regulator of ERAD.
Reason: The direct BAG6 claim is better captured as ERAD pathway participation.
Proposed replacements: ERAD pathway
GO:0005737 cytoplasm
IDA
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
ACCEPT
Summary: Whole-cell cytoplasmic localization is fully consistent with the BAG6 complex's proteostasis role.
Reason: This is a defensible localization term.
GO:0005515 protein binding
IPI
PMID:25535373
Bag6 complex contains a minimal tail-anchor-targeting module...
REMOVE
Summary: The structural complex paper supports specific adaptor/complex roles, not generic protein binding.
Reason: Remove generic protein-binding carryover.
GO:0071818 BAT3 complex
IDA
PMID:25535373
Bag6 complex contains a minimal tail-anchor-targeting module...
ACCEPT
Summary: This paper directly analyzes the BAG6 heterotrimeric complex architecture.
Reason: BAT3 complex membership is well supported.
GO:0005515 protein binding
IPI
PMID:23246001
SGTA recognizes a noncanonical ubiquitin-like domain in the ...
REMOVE
Summary: This ERAD paper supports specific SGTA/BAG6 pathway functions rather than a useful generic binding term.
Reason: Remove generic protein-binding carryover.
GO:0005829 cytosol
IDA
PMID:23246001
SGTA recognizes a noncanonical ubiquitin-like domain in the ...
ACCEPT
Summary: The SGTA/BAG6 ERAD paper supports a cytosolic BAG6 pool.
Reason: This matches the core QC role.
GO:0016020 membrane
IDA
PMID:23246001
SGTA recognizes a noncanonical ubiquitin-like domain in the ...
MODIFY
Summary: The paper supports regulated ER-membrane association through ERAD machinery rather than a generic membrane localization.
Reason: Use endoplasmic reticulum membrane as the more faithful location.
Proposed replacements: endoplasmic reticulum membrane
GO:0005634 nucleus
IDA
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
ACCEPT
Summary: The 2011 ERAD study notes nuclear sequestration when Trc35 is absent, consistent with BAG6's known dynamic localization.
Reason: Nuclear localization is supported.
GO:0031625 ubiquitin protein ligase binding
IPI
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
ACCEPT
Summary: Bag6 physically associates with ERAD E3 ligases such as gp78/AMFR and SYVN1, which is central to its UPS-adjacent chaperone role.
Reason: This specific binding term is mechanistically informative and supported.
Supporting Evidence:
GO:0051787 misfolded protein binding
IDA
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
ACCEPT
Summary: This is the key direct evidence for BAG6 recognizing aggregation-prone hydrophobic/misfolded clients.
Reason: The term accurately captures BAG6 holdase specificity.
Supporting Evidence:
GO:0071818 BAT3 complex
IDA
PMID:21636303
A ubiquitin ligase-associated chaperone holdase maintains po...
ACCEPT
Summary: The paper directly studies the BAG6/Ubl4A/Trc35 complex.
Reason: Complex membership is well supported.
GO:1990381 ubiquitin-specific protease binding
IPI
PMID:24424410
USP13 antagonizes gp78 to maintain functionality of a chaper...
KEEP AS NON CORE
Summary: Direct interaction with USP13 is well supported, but this is a specific partner interaction rather than the best core BAG6 function summary.
Reason: Keep as non-core mechanistic context.
Supporting Evidence:
GO:0031625 ubiquitin protein ligase binding
IPI
PMID:24981174
Cytosolic quality control of mislocalized proteins requires ...
ACCEPT
Summary: RNF126 recruitment to the BAG6 UBL domain is a core mechanistic part of mislocalized-protein triage.
Reason: This specific ligase-binding annotation is supported and informative.
Supporting Evidence:
GO:0005515 protein binding
IPI
PMID:18765639
BAT3 and SET1A form a complex with CTCFL/BORIS to modulate H...
REMOVE
Summary: The chromatin-regulator interaction paper does not justify retaining a generic protein-binding term.
Reason: Remove generic protein-binding carryover.
GO:0001822 kidney development
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Mouse-based developmental evidence is plausible but not core to human BAG6 proteostasis biology.
Reason: Keep as non-core.
GO:0005515 protein binding
IPI
PMID:17403783
HLA-B-associated transcript 3 (Bat3)/Scythe is essential for...
REMOVE
Summary: The p53 acetylation paper supports a specific nuclear scaffold role, not a useful generic protein-binding annotation.
Reason: Remove generic protein-binding carryover.
GO:0005634 nucleus
IDA
PMID:17403783
HLA-B-associated transcript 3 (Bat3)/Scythe is essential for...
ACCEPT
Summary: This paper supports nuclear BAG6 during DNA-damage signaling.
Reason: Nuclear localization is well supported.
GO:0005829 cytosol
IDA
PMID:17403783
HLA-B-associated transcript 3 (Bat3)/Scythe is essential for...
ACCEPT
Summary: This paper also supports the broader nucleo-cytoplasmic distribution of BAG6.
Reason: Cytosolic BAG6 is consistent with its principal proteostasis role.
GO:0005829 cytosol
IDA
PMID:20676083
A ribosome-associating factor chaperones tail-anchored membr...
ACCEPT
Summary: The ribosome-associated TA-targeting study places BAG6 complex function in the cytosol.
Reason: Cytosolic localization is central to GET-pathway capture/handoff.
GO:0006511 ubiquitin-dependent protein catabolic process
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: The broad catabolic-process inference is directionally consistent with BAG6 QC biology but loses specificity.
Reason: Keep as non-core rather than using it as a core summary term.
GO:0007130 synaptonemal complex assembly
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Orthology-based reproductive biology is secondary/contextual.
Reason: Keep as non-core.
GO:0007283 spermatogenesis
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Orthology-based reproductive biology is secondary/contextual.
Reason: Keep as non-core.
GO:0007420 brain development
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Orthology-based developmental biology is secondary/contextual.
Reason: Keep as non-core.
GO:0018393 internal peptidyl-lysine acetylation
IDA
PMID:17403783
HLA-B-associated transcript 3 (Bat3)/Scythe is essential for...
KEEP AS NON CORE
Summary: BAG6 scaffolds p300-dependent p53 acetylation after DNA damage, but this is a specialized nuclear stress-response role rather than the core proteostasis function.
Reason: Keep as non-core context.
Supporting Evidence:
GO:0030324 lung development
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Orthology-based developmental biology is secondary/contextual.
Reason: Keep as non-core.
GO:0031593 polyubiquitin modification-dependent protein binding
ISS
GO_REF:0000024
REMOVE
Summary: The BAG6 literature reviewed here does not provide a solid basis for a direct polyubiquitin-binding activity.
Reason: Remove this inferred binding term.
GO:0032435 negative regulation of proteasomal ubiquitin-dependent protein catabolic process
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: This reflects secondary stabilization biology rather than BAG6's central PN role.
Reason: Keep as non-core.
GO:0042771 intrinsic apoptotic signaling pathway in response to DNA damage by p53 class mediator
IMP
PMID:17403783
HLA-B-associated transcript 3 (Bat3)/Scythe is essential for...
KEEP AS NON CORE
Summary: This DNA-damage apoptosis role is well supported but context-specific.
Reason: Keep as non-core nuclear stress biology.
Supporting Evidence:
GO:0043022 ribosome binding
IDA
PMID:20676083
A ribosome-associating factor chaperones tail-anchored membr...
KEEP AS NON CORE
Summary: Ribosome association is an important mechanistic feature of the GET-pathway capture step, but it is not the best standalone summary of BAG6 core function.
Reason: Keep as mechanistically informative but non-core.
GO:0045861 negative regulation of proteolysis
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Secondary stabilization/proteolysis control is contextual rather than core.
Reason: Keep as non-core.
GO:0050821 protein stabilization
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: Selected-client stabilization is context-specific and not the main BAG6 role.
Reason: Keep as non-core.
GO:0070059 intrinsic apoptotic signaling pathway in response to endoplasmic reticulum stress
ISS
GO_REF:0000024
KEEP AS NON CORE
Summary: ER-stress apoptosis is context-dependent and not BAG6's principal conserved function.
Reason: Keep as non-core.
GO:0070628 proteasome binding
ISS
GO_REF:0000024
REMOVE
Summary: Direct proteasome binding is not clearly established by the BAG6 papers reviewed here.
Reason: Remove the inferred binding term.
GO:0071816 tail-anchored membrane protein insertion into ER membrane
IDA
PMID:20676083
A ribosome-associating factor chaperones tail-anchored membr...
ACCEPT
Summary: This is the direct GET-pathway core BAG6 process supported by the landmark ribosome-associated TA-targeting paper.
Reason: The evidence specifically fits tail-anchored membrane protein insertion into the ER membrane.
Supporting Evidence:
GO:0071818 BAT3 complex
IDA
PMID:20676083
A ribosome-associating factor chaperones tail-anchored membr...
ACCEPT
Summary: The 2010 Nature study defines the Bat3/Trc35/Ubl4A complex as the relevant functional unit.
Reason: BAT3 complex membership is directly supported.

Core Functions

BAG6 acts as an ATP-independent holdase/carrier for hydrophobic, misfolded, retrotranslocated, or tail-anchored client proteins, keeping them soluble long enough for productive ER delivery or proteasome-directed quality control.

Supporting Evidence:

Within the BAG6/UBL4A/GET4 complex, BAG6 provides the scaffold/adaptor surface that links SGTA-bound hydrophobic clients to the TRC40/GET pathway and organizes the pretargeting complex architecture.

Supporting Evidence:
  • PMID:25535373
  • PMID:25713138
  • file:human/BAG6/BAG6-deep-research-falcon.md
    C-terminal region is a structural part of the substrate-loading complex that bridges UBL4A/SGTA to TRC35/TRC40

BAG6 preferentially recognizes aggregation-prone hydrophobic or misfolded clients generated during ERAD, cytosolic mislocalization, or ER stress rerouting, linking substrate capture to downstream quality-control decisions.

Supporting Evidence:
  • PMID:21636303
  • PMID:26565908
  • file:human/BAG6/BAG6-deep-research-falcon.md
    BAG6 captures hydrophobic or mislocalized clients, keeps them soluble, recruits ubiquitination machinery

References

file:human/BAG6/BAG6-notes.md
BAG6 review notes
file:human/BAG6/BAG6-deep-research-falcon.md
Falcon deep research report for BAG6
  • Falcon research supports BAG6 as a cytosolic BAG6/UBL4A/TRC35 holdase and adaptor linking tail-anchored protein targeting with ERAD/proteasome quality control.
    "BAG6 is embedded in this pathway as both a substrate-holding factor and a quality-control adaptor"
Manual transfer of experimentally-verified manual GO annotation data to orthologs by curator judgment of sequence similarity
Annotation inferences using phylogenetic trees
Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location vocabulary mapping, accompanied by conservative changes to GO terms applied by UniProt
Gene Ontology annotation based on curation of immunofluorescence data from the Human Protein Atlas
Automatic transfer of experimentally verified manual GO annotation data to orthologs using Ensembl Compara
Combined Automated Annotation using Multiple IEA Methods
A ribosome-associating factor chaperones tail-anchored membrane proteins.
A ubiquitin ligase-associated chaperone holdase maintains polypeptides in soluble states for proteasome degradation.
Protein targeting and degradation are coupled for elimination of mislocalized proteins.
SGTA antagonizes BAG6-mediated protein triage.
A ubiquitin-like domain recruits an oligomeric chaperone to a retrotranslocation complex in endoplasmic reticulum-associated degradation.
USP13 antagonizes gp78 to maintain functionality of a chaperone in ER-associated degradation.
Cytosolic quality control of mislocalized proteins requires RNF126 recruitment to Bag6.
Bag6 complex contains a minimal tail-anchor-targeting module and a mock BAG domain.
Structure of a BAG6 (Bcl-2-associated athanogene 6)-Ubl4a (ubiquitin-like protein 4a) complex reveals a novel binding interface that functions in tail-anchored protein biogenesis.
Structural basis for regulation of the nucleo-cytoplasmic distribution of Bag6 by TRC35.
HLA-B-associated transcript 3 (Bat3)/Scythe is essential for p300-mediated acetylation of p53.
Human leukocyte antigen-B-associated transcript 3 is released from tumor cells and engages the NKp30 receptor on natural killer cells.
Dendritic cells release HLA-B-associated transcript-3 positive exosomes to regulate natural killer function.
Analysis of a high-throughput yeast two-hybrid system and its use to predict the function of intracellular proteins encoded within the human MHC class III region.
Ricin triggers apoptotic morphological changes through caspase-3 cleavage of BAT3.
Towards a proteome-scale map of the human protein-protein interaction network.
BAT3 and SET1A form a complex with CTCFL/BORIS to modulate H3K4 histone dimethylation and gene expression.
Defining the membrane proteome of NK cells.
Mapping a dynamic innate immunity protein interaction network regulating type I interferon production.
The SARS-coronavirus-host interactome: identification of cyclophilins as target for pan-coronavirus inhibitors.
The stalk domain and the glycosylation status of the activating natural killer cell receptor NKp30 are important for ligand binding.
SGTA recognizes a noncanonical ubiquitin-like domain in the Bag6-Ubl4A-Trc35 complex to promote endoplasmic reticulum-associated degradation.
A quantitative chaperone interaction network reveals the architecture of cellular protein homeostasis pathways.
A proteome-scale map of the human interactome network.
A human interactome in three quantitative dimensions organized by stoichiometries and abundances.
Selective Binding of AIRAPL Tandem UIMs to Lys48-Linked Tri-Ubiquitin Chains.
UBQLN4 recognizes mislocalized transmembrane domain proteins and targets these to proteasomal degradation.
Extensive disruption of protein interactions by genetic variants across the allele frequency spectrum in human populations.
Dual proteome-scale networks reveal cell-specific remodeling of the human interactome.
OpenCell: Endogenous tagging for the cartography of human cellular organization.
Definition of the human mitochondrial TOM interactome reveals TRABD as a new interacting protein.
Pre-emptive Quality Control Protects the ER from Protein Overload via the Proximity of ERAD Components and SRP.

Deep Research

Falcon

(BAG6-deep-research-falcon.md)
Research Report: Human BAG6 (UniProt P46379; BAT3/Scythe) — Functional Annotation (2023–2024 prioritized) Falcon Edison Scientific Literature 29 citations 2026-05-10T19:00:13.355501

The research report should be a detailed narrative explaining the function, biological processes, and localization of the gene product. Citations should be given for all claims.

You should prioritize authoritative reviews and primary scientific literature when conducting research. You can supplement
this with annotations you find in gene/protein databases, but these can be outdated or inaccurate.

We are specifically interested in the primary function of the gene - for enzymes, what reaction is catalyzed, and what is the substrate specificity? For transporters, what is the substrate? For structural proteins or adapters, what is the broader structural role? For signaling molecules, what is the role in the pathway.

We are interested in where in or outside the cell the gene product carries out its function.

We are also interested in the signaling or biochemical pathways in which the gene functions. We are less interested in broad pleiotropic effects, except where these elucidate the precise role.

Include evidence where possible. We are interested in both experimental evidence as well as inference from structure, evolution, or bioinformatic analysis. Precise studies should be prioritized over high-throughput, where available.

Research Report: Human BAG6 (UniProt P46379; BAT3/Scythe) — Functional Annotation (2023–2024 prioritized)

0) Target verification (mandatory)

The literature synthesized here explicitly refers to human BAG6 using the aliases BAT3 and Scythe, and describes functions/complexes consistent with UniProt P46379 (N-terminal ubiquitin-like domain; BAG6/Bag6_BAGS regions; cytosolic PQC and TA-protein handling). For example, BAG6 is discussed as part of the mammalian TRC/GET tail-anchored pathway and as a cytosolic quality-control holdase and ubiquitin-ligase adaptor in multiple studies (hagiwara2023proteotoxicstressesstimulate pages 1-3, hagiwara2023proteotoxicstressesstimulate pages 3-5, miyauchi2023bag6supportsstress pages 1-2, zhou2024bag6inhibitsinfluenza pages 2-4, guna2018transmembranedomainrecognition pages 4-6).

1) Key concepts and definitions (current understanding)

1.1 Tail-anchored (TA) proteins and the TRC/GET pathway

Tail-anchored proteins are membrane proteins with a single hydrophobic transmembrane domain (TMD) near the C-terminus that typically insert post-translationally into the ER membrane. In mammals, TA protein targeting is carried out by the TRC40 pathway (homologous to the yeast GET pathway). BAG6 is embedded in this pathway as both a substrate-holding factor and a quality-control adaptor (guna2018transmembranedomainrecognition pages 4-6).

A key synthesis is that BAG6’s C-terminal region is a structural part of the substrate-loading complex that bridges UBL4A/SGTA to TRC35/TRC40, while the N-terminal ubiquitin-like (UBL) domain recruits ubiquitination machinery (notably RNF126) to route failed clients to proteasomal degradation (guna2018transmembranedomainrecognition pages 4-6).

1.2 Cytosolic protein quality control (PQC) for hydrophobic/mislocalized proteins

BAG6 is widely characterized as a chaperone/holdase that binds exposed hydrophobic segments of nascent or mislocalized proteins, preventing aggregation and enabling either productive targeting or degradative routing (abildgaard2020cochaperonesintargeting pages 9-11, abildgaard2020cochaperonesintargeting pages 21-23). Mechanistically, BAG6 links client capture to ubiquitin-mediated proteasomal degradation via recruitment of E3 ligases (e.g., RNF126) and via interactions enabling delivery to the proteasome (guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11).

1.3 “Triage” as a stress-sensitive decision point

Recent work emphasizes that BAG6 participates in stress-sensitive triage: allocating capacity between (i) biogenesis of TA proteins via TRC machinery and (ii) elimination of defective/aggregation-prone proteins. Under proteotoxic stress, BAG6 complex composition can shift, consistent with triage between these fates (hagiwara2023proteotoxicstressesstimulate pages 3-5, hagiwara2023proteotoxicstressesstimulate pages 10-12).

2) Molecular functions, domains, and interaction partners

2.1 Core complexes and partners

Canonical TA targeting / TRC pathway partners: BAG6 forms or participates in assemblies including UBL4A, TRC35, TRC40, and interacts functionally with SGTA and the ER insertase WRB–CAML in TA handover and insertion (roboti2022mitochondrialantiviralsignallingprotein pages 1-2, guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11).

Quality-control and degradation partners: BAG6’s UBL domain can recruit the E3 ligase RNF126 (review-level synthesis), and stress can increase BAG6 association with UBQLN4 in the context of proteasome inhibition (hagiwara2023proteotoxicstressesstimulate pages 3-5, guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11).

2.2 Biochemical and quantitative evidence for complex stability/remodeling (2023)

A notable quantitative contribution from 2023 is the demonstration that BAG6–UBL4A binding is extremely tight in vitro (Kd ~2.2 nM) and that estimated cellular concentrations in HEK293T are high (~720 nM BAG6; ~700 nM UBL4A), yet proteotoxic stress still induces dissociation in cells—supporting dynamic remodeling rather than a static assembly (hagiwara2023proteotoxicstressesstimulate pages 3-5).

3) Subcellular localization and where BAG6 acts

BAG6 is principally discussed as acting in the cytosol at the interface of newly synthesized hydrophobic proteins, TA substrate capture, and proteasomal delivery, with functional coupling to ER insertion machinery (TRC40; WRB–CAML) (roboti2022mitochondrialantiviralsignallingprotein pages 1-2, guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11). It can also appear in ubiquitin-positive inclusions/aggregates under proteotoxic stress conditions, consistent with a PQC role when aggregation risk is high (hagiwara2023proteotoxicstressesstimulate pages 3-5).

Some mechanistic evidence indicates BAG6 is a nucleocytoplasmic shuttling protein, with cytosolic retention promoted by GET4 masking a nuclear localization sequence; disease-linked perturbations of GET4 interactions can alter localization (wang2021analysisofthe pages 80-84).

4) Biological pathways and experimentally supported roles

4.1 TA protein biogenesis and hydrophobic-client triage

A high-impact conceptual model is that BAG6 preferentially binds long hydrophobic TMDs with slow off-rate, sequestering clients that fail timely membrane engagement and routing them toward ubiquitination and proteasomal degradation, while still supporting handover to the TRC40 targeting apparatus for productive TA insertion (guna2018transmembranedomainrecognition pages 4-6).

4.2 Stress-driven remodeling of TA-recognition vs degradative PQC (2023)

Hagiwara et al. (Biochemical Journal; Oct 2023; https://doi.org/10.1042/BCJ20230267) show that proteotoxic stressors (including polyQ aggregation, proteasome inhibition, mitochondrial depolarization) promote dissociation of UBL4A from BAG6, and propose this as a triage mechanism prioritizing degradation over TA synthesis under stress (hagiwara2023proteotoxicstressesstimulate pages 3-5, hagiwara2023proteotoxicstressesstimulate pages 10-12). The authors explicitly connect disruption of BAG6–UBL4A to impaired TA protein synthesis needed for vesicular trafficking and discuss relevance to aggregate-driven neurodegeneration mechanisms (hagiwara2023proteotoxicstressesstimulate pages 10-12).

4.3 Innate immunity: BAG6 regulation of MAVS TA client pool (2022; mechanistic bridge to immunity)

Roboti et al. (Journal of Cell Science; May 2022; https://doi.org/10.1242/jcs.259596) identify the mitochondrial TA protein MAVS as an endogenous client of both SGTA and the BAG6 complex, and propose that BAG6 binds a cytosolic pool of MAVS prior to misinsertion and retrieval, with the BAG6-associated fraction responding dynamically to innate immune activation (roboti2022mitochondrialantiviralsignallingprotein pages 1-2).

4.4 Novel role in actin cytoskeleton control via RhoA stabilization (2023)

Miyauchi et al. (Molecular Biology of the Cell; Apr 2023; https://doi.org/10.1091/mbc.e22-08-0355) identify BAG6 as a factor required to stabilize endogenous RhoA and thereby support stress fiber formation, focal adhesion assembly, and cell migration. Mechanistically, BAG6 depletion increases association of GDP-bound RhoA with CUL3-based E3 ligases, increasing RhoA polyubiquitination and degradation (miyauchi2023bag6supportsstress pages 1-2, miyauchi2023bag6supportsstress pages 11-13). The study includes quantitative ubiquitination comparisons with p < 0.05 (miyauchi2023bag6supportsstress pages 11-13).

4.5 Viral restriction: influenza A PB2 degradation and RdRp assembly interference (2024)

Zhou et al. (PLOS Pathogens; Mar 2024; https://doi.org/10.1371/journal.ppat.1012110) identify BAG6 as a host restriction factor for influenza A virus (IAV). Mechanistically, BAG6 binds the N-terminus of PB2, competes with PB1 for RdRp assembly, and promotes K48-linked ubiquitination and degradation of PB2, including mapping PB2 ubiquitination at K189 (zhou2024bag6inhibitsinfluenza pages 1-2).

Effect sizes and in vivo relevance (key statistics):
* In vitro, BAG6 overexpression reduced infectious titers by ~10–15-fold (TCID50), while BAG6 loss increased titers up to ~10-fold (zhou2024bag6inhibitsinfluenza pages 2-4).
* In mice, BAG6 knockdown increased disease severity with ~40% mortality vs 0% in controls (by 8 dpi) and ~10-fold higher lung viral loads at 3 and 5 dpi (zhou2024bag6inhibitsinfluenza pages 4-7).
* Domain requirements: antiviral activity required BAG6 N-terminus including UBL domain (aa 17–92) plus a PB2-binding region (aa 124–186) (zhou2024bag6inhibitsinfluenza pages 1-2). These quantitative/doman-mapping results are also supported by the extracted figure panels (zhou2024bag6inhibitsinfluenza media f7dcee05).

5) Recent developments and latest research highlights (2023–2024 focus)

  1. Stress-sensitive disassembly of BAG6–UBL4A despite nanomolar affinity suggests regulated triage and provides a mechanistic link between proteotoxic stress and impairment of TA biogenesis/vesicular trafficking (Hagiwara 2023) (hagiwara2023proteotoxicstressesstimulate pages 3-5, hagiwara2023proteotoxicstressesstimulate pages 10-12).
  2. Expansion of client scope to soluble signaling proteins: BAG6 acts as a GDP-RhoA holdase that protects against E3 access and degradation, connecting BAG6 PQC logic to cytoskeletal dynamics and migration (Miyauchi 2023) (miyauchi2023bag6supportsstress pages 1-2, miyauchi2023bag6supportsstress pages 11-13).
  3. Direct antiviral effector mechanism: BAG6 can function as an influenza restriction factor by driving degradation of a viral polymerase subunit and disrupting polymerase assembly, with large-magnitude in vitro and in vivo effects (Zhou 2024) (zhou2024bag6inhibitsinfluenza pages 2-4, zhou2024bag6inhibitsinfluenza pages 1-2, zhou2024bag6inhibitsinfluenza pages 4-7, zhou2024bag6inhibitsinfluenza media f7dcee05).
  4. Tumor biology via extracellular vesicles (EVs): BAG6 is implicated as a tumor-suppressive regulator in PDAC via suppression of IL33-presenting EV release and mast-cell activation (Alhamwe 2024) (alhamwe2024bag6restrictspancreatic pages 1-2, alhamwe2024bag6restrictspancreatic pages 8-10).
  5. Repair-associated gene expression signatures: In an emphysema/COPD repair model, Bag6 is among top differentially expressed genes in response to liver growth factor treatment, suggesting involvement in inflammation/immune-response regulation during repair (Carretero 2024) (carretero2024differentiallunggene pages 8-10).

6) Current applications and real-world implementations

6.1 Oncology: biomarker and therapy-stratification concepts (PDAC)

Alhamwe et al. (Cellular & Molecular Immunology; Jun 2024; https://doi.org/10.1038/s41423-024-01195-1) report that Bag6 deficiency accelerates PDAC tumor growth in an EV-dependent and mast-cell–dependent manner, and propose a translational rationale for considering mast-cell depletion (imatinib) in patients stratified by low BAG6 expression and high mast-cell infiltration (alhamwe2024bag6restrictspancreatic pages 1-2). They also discuss pharmacologic inhibition of EV release (GW4869) in vivo as a rescue approach (alhamwe2024bag6restrictspancreatic pages 11-12). The study notes that high BAG6 gene expression and high plasma BAG6 protein are associated with longer overall survival, supporting biomarker relevance (alhamwe2024bag6restrictspancreatic pages 1-2).

6.2 Host-directed antivirals / biomarkers (influenza A)

Zhou et al. (2024) show that BAG6 manipulation strongly affects viral replication (order-of-magnitude changes in titers) and disease outcomes in mice, implying BAG6-regulated pathways (ubiquitin-dependent PB2 turnover and polymerase assembly) could be explored for host-directed antiviral strategies or as biomarkers of susceptibility/severity (zhou2024bag6inhibitsinfluenza pages 2-4, zhou2024bag6inhibitsinfluenza pages 4-7).

6.3 Disease modeling and repair-response markers (COPD model)

In a cigarette-smoke emphysema model with liver growth factor treatment, Bag6 is upregulated with fold change 2.114 (FDR 3.69E-03) and qRT-PCR validation showing 1.390 ± 0.119 vs 1.061 ± 0.138; p = 0.0087, suggesting Bag6 as part of a repair-associated gene signature in this model (Carretero 2024; Aug 22, 2024; https://doi.org/10.1371/journal.pone.0309166) (carretero2024differentiallunggene pages 8-10).

7) Expert opinions and analysis (authoritative synthesis)

7.1 BAG6 as a metazoan “quality-control embellishment” of TA targeting

A widely cited synthesis (Guna & Hegde, Current Biology; Apr 23, 2018; https://doi.org/10.1016/j.cub.2018.02.004) frames BAG6 as a metazoan-specific adaptor that integrates TA targeting with a “fail-safe” degradative arm: BAG6 captures long hydrophobic TMDs with slow off-rate, recruits RNF126 via its UBL domain for ubiquitination, and uses its C-terminus to connect UBL4A/SGTA with TRC35/TRC40 to enable either insertion or disposal (guna2018transmembranedomainrecognition pages 4-6).

7.2 BAG6 as a proteasome-delivery co-chaperone/holdase

Abildgaard et al. (Biomolecules; Aug 2020; https://doi.org/10.3390/biom10081141) emphasize BAG6 as a holdase that can function largely independently of Hsp70, acting as part of a BAG6–UBL4A–TRC35 complex that can recruit ubiquitination machinery (RNF126) and interact with the proteasome to deliver substrates for degradation (abildgaard2020cochaperonesintargeting pages 9-11).

8) Relevant statistics and quantitative data (recent studies)

  • BAG6–UBL4A biochemical affinity and abundance: in vitro Kd 2.2 nM; estimated cellular concentrations ~720 nM BAG6 and ~700 nM UBL4A (Hagiwara 2023) (hagiwara2023proteotoxicstressesstimulate pages 3-5).
  • Influenza A restriction (in vitro): BAG6 overexpression yields ~10–15-fold lower titers; BAG6 loss yields up to ~10-fold higher titers (Zhou 2024) (zhou2024bag6inhibitsinfluenza pages 2-4).
  • Influenza A restriction (in vivo): BAG6 knockdown mice show ~40% mortality vs 0% and ~10-fold higher lung viral loads (Zhou 2024) (zhou2024bag6inhibitsinfluenza pages 4-7). These outcomes are visualized in extracted figure panels (zhou2024bag6inhibitsinfluenza media f7dcee05).
  • COPD model gene expression: Bag6 in top DEGs with FC 2.114 and FDR 3.69E-03; qRT-PCR validation p=0.0087 (Carretero 2024) (carretero2024differentiallunggene pages 8-10).
  • RhoA ubiquitination phenotype: BAG6 knockdown enhances RhoA polyubiquitination, with quantitative comparisons reported with p < 0.05 (Miyauchi 2023) (miyauchi2023bag6supportsstress pages 11-13).

9) Disease associations (database-level summary)

Open Targets reports BAG6 target–disease association evidence for neurodegenerative disease based on CRISPRi neuronal screen studies (PubMed 34031600) (OpenTargets Search: -BAG6). This is hypothesis-generating and should be interpreted as functional-genomics evidence rather than direct causal mechanistic proof.

10) Consolidated functional summary table

The table below summarizes BAG6’s best-supported functional modules (partners, locations, mechanisms, and recent quantitative evidence).

Function/module Key partners/complex Cellular location Mechanism (1 sentence) Key recent evidence (2023-2024) with effect sizes/statistics when available Key foundational evidence/reviews
Tail-anchored (TA) protein capture and handoff BAG6–UBL4A–TRC35 complex with SGTA, TRC40, WRB–CAML Cytosol; ER-targeting interface BAG6 acts as a hydrophobic-client holdase and scaffold that links SGTA-bound TA clients to TRC40-mediated ER insertion while retaining failed clients for triage. (roboti2022mitochondrialantiviralsignallingprotein pages 1-2, guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11) Proteotoxic stress and polyQ inclusions trigger UBL4A dissociation from BAG6; BAG6–UBL4A affinity is strong in vitro (Kd 2.2 nM), and estimated cellular concentrations are ~720 nM BAG6 and ~700 nM UBL4A, supporting a normally stable TA-recognition complex that is remodeled under stress. (hagiwara2023proteotoxicstressesstimulate pages 1-3, hagiwara2023proteotoxicstressesstimulate pages 3-5) Reviews place BAG6 as a metazoan quality-control layer in the TRC40 pathway, with its C-terminus bridging UBL4A/SGTA and TRC35/TRC40 during TA targeting. (guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11)
Cytosolic PQC and degradation of hydrophobic/mislocalized proteins RNF126, UBQLN4, SGTA, proteasome Cytosol; proteasome-associated; aggregate-prone compartments BAG6 captures hydrophobic or mislocalized clients, keeps them soluble, recruits ubiquitination machinery, and promotes proteasomal delivery/degradation. (guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11, abildgaard2020cochaperonesintargeting pages 21-23) Under proteasome inhibition, soluble BAG6 loses associated UBL4A but gains UBQLN4, indicating stress-dependent remodeling from TA biogenesis toward degradative PQC; BAG6 also colocalizes with ubiquitin-positive inclusions. (hagiwara2023proteotoxicstressesstimulate pages 3-5) BAG6’s N-terminal UBL recruits RNF126, and reviews describe BAG6 as a holdase for TA proteins, defective nascent chains, ERAD substrates, and mislocalized membrane proteins. (guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11, abildgaard2020cochaperonesintargeting pages 21-23)
ERAD-linked substrate processing SGTA, p97 pathway components, RNF126 Cytosol and ER-proximal quality-control interface BAG6 helps route failed membrane/secretory protein biogenesis products and certain ERAD substrates into ubiquitin-proteasome degradation rather than productive insertion. (abildgaard2020cochaperonesintargeting pages 9-11, roboti2022mitochondrialantiviralsignallingprotein pages 15-15, you2020paqr9modulatesbag6mediated pages 15-16) 2023 review context continues to position BAG6 in substrate processing and ER-proximal PQC, though BAG6-specific quantitative 2023–2024 ERAD effect sizes were not provided in the retrieved excerpts. (abildgaard2020cochaperonesintargeting pages 9-11, hagiwara2023proteotoxicstressesstimulate pages 14-15) Reviews summarize BAG6 as coupling client capture, ligase recruitment, and proteasome targeting for ERAD-like outcomes after failed membrane engagement. (guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11)
Stress-responsive remodeling of the BAG6 complex UBL4A, TRC35, TRC40, UBQLN4 Cytosol; insoluble aggregates under stress Proteotoxic stress redistributes BAG6 complexes by dissociating UBL4A and favoring aggregate-associated/degradative BAG6 states. (hagiwara2023proteotoxicstressesstimulate pages 1-3, hagiwara2023proteotoxicstressesstimulate pages 3-5) PolyQ inclusions, proteasome inhibition, and mitochondrial depolarization reduce BAG6–UBL4A association; overexpressed UBL4A suppresses BAG6 translocation into insoluble aggregates. (hagiwara2023proteotoxicstressesstimulate pages 3-5) Foundational models of BAG6 explain this as a triage switch between TA targeting and degradation of hydrophobic clients. (guna2018transmembranedomainrecognition pages 4-6, abildgaard2020cochaperonesintargeting pages 9-11)
Innate immunity-linked TA client control MAVS with SGTA and BAG6–UBL4A–TRC35 Cytosolic pool; ER and mitochondrial-signaling membrane interface BAG6 binds a cytosolic pool of the mitochondrial TA protein MAVS before misinsertion/retrieval, thereby modulating the fraction available for antiviral signaling. (roboti2022mitochondrialantiviralsignallingprotein pages 1-2) BioID2 proximity labeling identified MAVS as a high-confidence SGTA interactor (BFDR < 0.05), and BAG6-associated MAVS changed dynamically during innate immune activation. (roboti2022mitochondrialantiviralsignallingprotein pages 1-2) This extends the established BAG6 client-triage model from generic TA proteins to an endogenous immune signaling TA client. (roboti2022mitochondrialantiviralsignallingprotein pages 1-2, guna2018transmembranedomainrecognition pages 4-6)
Direct antiviral restriction of influenza A virus Viral PB2, PB1/RdRp complex Predominantly cytosolic/nuclear viral polymerase context BAG6 binds PB2, competes with PB1 for polymerase assembly, and promotes K48-linked ubiquitination-dependent PB2 degradation to suppress IAV replication. (zhou2024bag6inhibitsinfluenza pages 2-4, zhou2024bag6inhibitsinfluenza pages 1-2) Overexpression decreased IAV titers by ~10–15-fold across H1N1, H7N9, H9N2, and H5N1; BAG6 loss increased titers up to ~10-fold; in mice, BAG6 knockdown caused ~40% mortality by 8 dpi versus 0% in controls and ~10-fold higher lung viral loads. Figure review also mapped required BAG6 regions to the UBL domain (aa 17–92) plus a PB2-binding region (aa 124–186). (zhou2024bag6inhibitsinfluenza pages 2-4, zhou2024bag6inhibitsinfluenza pages 4-7, zhou2024bag6inhibitsinfluenza media f7dcee05) Recent primary work builds on broader BAG6 chaperone/PQC functions rather than a separate foundational antiviral literature in the retrieved corpus. (abildgaard2020cochaperonesintargeting pages 9-11, abildgaard2020cochaperonesintargeting pages 21-23)
Cytoskeletal regulation via small GTPase stabilization RhoA, CUL3-based ubiquitin ligases Cytosol; stress fibers/focal adhesions BAG6 acts as a holdase for GDP-bound RhoA, limiting its association with CUL3 ligases and thereby preserving RhoA-dependent actin assembly. (miyauchi2023bag6supportsstress pages 1-2) BAG6 depletion increased RhoA polyubiquitination and degradation, impairing stress fibers, focal adhesion assembly, and cell migration; RhoA re-expression rescued the phenotype. (miyauchi2023bag6supportsstress pages 1-2) This expands BAG6 client selectivity beyond membrane-hydrophobic clients to a soluble signaling protein stabilized through a related anti-degradation holdase mechanism. (miyauchi2023bag6supportsstress pages 1-2, abildgaard2020cochaperonesintargeting pages 21-23)
Tumor microenvironment and extracellular vesicle regulation p53, CBP/p300, ESCRT machinery, IL33-positive EVs, mast cells Cytosol/nucleus-linked vesicle biogenesis pathway; extracellular vesicles; tumor microenvironment BAG6 restrains pancreatic tumor progression by promoting EV biogenesis programs that prevent release of IL33-presenting EVs that activate mast cells and remodel the TME. (alhamwe2024bag6restrictspancreatic pages 1-2) In PDAC models, Bag6 deficiency accelerated subcutaneous and orthotopic tumor growth in an EV-dependent manner; high BAG6 gene expression and plasma BAG6 associated with longer overall survival, and imatinib-mediated mast-cell depletion reduced tumor growth in BAG6-low contexts. (alhamwe2024bag6restrictspancreatic pages 1-2) The study leverages prior evidence that BAG6 regulates membrane vesicle trafficking, EV cargo sorting, and immune-cell activation. (alhamwe2024bag6restrictspancreatic pages 1-2)
Nuclear-cytoplasmic shuttling and localization control GET4, UBL4A, FBXO7 Cytosol and nucleus BAG6 is a nucleocytoplasmic shuttling protein whose retention in the cytosol is promoted by GET4 masking its nuclear localization sequence. (wang2021analysisofthe pages 80-84) Dissertation-level mechanistic evidence reports that FBXO7-enhanced GET4 ubiquitination increases GET4–BAG6 binding and shifts BAG6 cytoplasmic localization; disease-associated FBXO7 variants weaken this control. (wang2021analysisofthe pages 80-84) This localization logic fits BAG6’s established need to remain cytosolic for TA targeting/PQC while retaining the capacity for alternative signaling roles. (wang2021analysisofthe pages 80-84, guna2018transmembranedomainrecognition pages 4-6)

Table: This table summarizes the best-supported functional modules of human BAG6 (UniProt P46379), integrating core mechanistic roles, partners, locations, and recent 2023-2024 evidence with quantitative findings where available. It is useful for quickly distinguishing BAG6’s foundational tail-anchored protein/PQC functions from newer roles in antiviral defense, cytoskeletal regulation, and tumor biology.

11) Key references (URLs and publication dates)

  • Zhou Y. et al. “BAG6 inhibits influenza A virus replication…” PLOS Pathogens. Mar 2024. https://doi.org/10.1371/journal.ppat.1012110 (zhou2024bag6inhibitsinfluenza pages 2-4, zhou2024bag6inhibitsinfluenza pages 4-7)
  • Alhamwe B.A. et al. “BAG6 restricts pancreatic cancer progression…” Cellular & Molecular Immunology. Jun 2024. https://doi.org/10.1038/s41423-024-01195-1 (alhamwe2024bag6restrictspancreatic pages 1-2)
  • Carretero L.S. et al. “Differential lung gene expression identified … Bag6 …” PLOS ONE. Aug 22, 2024. https://doi.org/10.1371/journal.pone.0309166 (carretero2024differentiallunggene pages 8-10)
  • Hagiwara T. et al. “Proteotoxic stresses stimulate dissociation of UBL4A …” Biochemical Journal. Oct 2023. https://doi.org/10.1042/BCJ20230267 (hagiwara2023proteotoxicstressesstimulate pages 3-5, hagiwara2023proteotoxicstressesstimulate pages 10-12)
  • Miyauchi M. et al. “BAG6 supports stress fiber formation …” Molecular Biology of the Cell. Apr 1, 2023. https://doi.org/10.1091/mbc.e22-08-0355 (miyauchi2023bag6supportsstress pages 11-13)
  • Roboti P. et al. “Mitochondrial antiviral-signalling protein is a client …” Journal of Cell Science. May 2022. https://doi.org/10.1242/jcs.259596 (roboti2022mitochondrialantiviralsignallingprotein pages 1-2)
  • Abildgaard A.B. et al. “Co-Chaperones in Targeting and Delivery …” Biomolecules. Aug 2020. https://doi.org/10.3390/biom10081141 (abildgaard2020cochaperonesintargeting pages 9-11)
  • Guna A., Hegde R.S. “Transmembrane Domain Recognition …” Current Biology. Apr 23, 2018. https://doi.org/10.1016/j.cub.2018.02.004 (guna2018transmembranedomainrecognition pages 4-6)

12) Limitations of this synthesis

Some frequently discussed BAG6 roles (e.g., antigen presentation, apoptosis regulation, mitophagy/autophagy) are mentioned in the retrieved PDAC/virology papers as contextual background but were not comprehensively supported with primary mechanistic excerpts within the currently retrieved text segments. Accordingly, the report emphasizes roles with direct mechanistic and/or quantitative support in the available evidence corpus.

References

  1. (hagiwara2023proteotoxicstressesstimulate pages 1-3): Takumi Hagiwara, Ryosuke Minami, Chizuru Ushio, Naoto Yokota, and Hiroyuki Kawahara. Proteotoxic stresses stimulate dissociation of ubl4a from the tail-anchored protein recognition complex. Biochemical Journal, 480:1583-1598, Oct 2023. URL: https://doi.org/10.1042/bcj20230267, doi:10.1042/bcj20230267. This article has 0 citations and is from a domain leading peer-reviewed journal.

  2. (hagiwara2023proteotoxicstressesstimulate pages 3-5): Takumi Hagiwara, Ryosuke Minami, Chizuru Ushio, Naoto Yokota, and Hiroyuki Kawahara. Proteotoxic stresses stimulate dissociation of ubl4a from the tail-anchored protein recognition complex. Biochemical Journal, 480:1583-1598, Oct 2023. URL: https://doi.org/10.1042/bcj20230267, doi:10.1042/bcj20230267. This article has 0 citations and is from a domain leading peer-reviewed journal.

  3. (miyauchi2023bag6supportsstress pages 1-2): Maho Miyauchi, Reina Matsumura, and Hiroyuki Kawahara. Bag6 supports stress fiber formation by preventing the ubiquitin-mediated degradation of rhoa. Molecular Biology of the Cell, Apr 2023. URL: https://doi.org/10.1091/mbc.e22-08-0355, doi:10.1091/mbc.e22-08-0355. This article has 7 citations and is from a domain leading peer-reviewed journal.

  4. (zhou2024bag6inhibitsinfluenza pages 2-4): Yong Zhou, Tian Li, Yunfan Zhang, Nianzhi Zhang, Yuxin Guo, Xiaoyi Gao, Wenjing Peng, Sicheng Shu, Chuankuo Zhao, Di Cui, Honglei Sun, Yipeng Sun, Jinhua Liu, Jun Tang, Rui Zhang, and Juan Pu. Bag6 inhibits influenza a virus replication by inducing viral polymerase subunit pb2 degradation and perturbing rdrp complex assembly. PLOS Pathogens, 20:e1012110, Mar 2024. URL: https://doi.org/10.1371/journal.ppat.1012110, doi:10.1371/journal.ppat.1012110. This article has 13 citations and is from a highest quality peer-reviewed journal.

  5. (guna2018transmembranedomainrecognition pages 4-6): Alina Guna and Ramanujan S. Hegde. Transmembrane domain recognition during membrane protein biogenesis and quality control. Current Biology, 28:R498-R511, Apr 2018. URL: https://doi.org/10.1016/j.cub.2018.02.004, doi:10.1016/j.cub.2018.02.004. This article has 151 citations and is from a highest quality peer-reviewed journal.

  6. (abildgaard2020cochaperonesintargeting pages 9-11): Amanda B. Abildgaard, Sarah K. Gersing, Sven Larsen-Ledet, Sofie V. Nielsen, Amelie Stein, Kresten Lindorff-Larsen, and Rasmus Hartmann-Petersen. Co-chaperones in targeting and delivery of misfolded proteins to the 26s proteasome. Biomolecules, 10:1141, Aug 2020. URL: https://doi.org/10.3390/biom10081141, doi:10.3390/biom10081141. This article has 54 citations.

  7. (abildgaard2020cochaperonesintargeting pages 21-23): Amanda B. Abildgaard, Sarah K. Gersing, Sven Larsen-Ledet, Sofie V. Nielsen, Amelie Stein, Kresten Lindorff-Larsen, and Rasmus Hartmann-Petersen. Co-chaperones in targeting and delivery of misfolded proteins to the 26s proteasome. Biomolecules, 10:1141, Aug 2020. URL: https://doi.org/10.3390/biom10081141, doi:10.3390/biom10081141. This article has 54 citations.

  8. (hagiwara2023proteotoxicstressesstimulate pages 10-12): Takumi Hagiwara, Ryosuke Minami, Chizuru Ushio, Naoto Yokota, and Hiroyuki Kawahara. Proteotoxic stresses stimulate dissociation of ubl4a from the tail-anchored protein recognition complex. Biochemical Journal, 480:1583-1598, Oct 2023. URL: https://doi.org/10.1042/bcj20230267, doi:10.1042/bcj20230267. This article has 0 citations and is from a domain leading peer-reviewed journal.

  9. (roboti2022mitochondrialantiviralsignallingprotein pages 1-2): Peristera Roboti, Craig Lawless, and Stephen High. Mitochondrial antiviral-signalling protein is a client of the bag6 protein quality control complex. Journal of Cell Science, May 2022. URL: https://doi.org/10.1242/jcs.259596, doi:10.1242/jcs.259596. This article has 1 citations and is from a domain leading peer-reviewed journal.

  10. (wang2021analysisofthe pages 80-84): Quan Wang. Analysis of the interaction of the parkinsonism-associated protein fbxo7 with the bag6 complex. Dissertation, Jan 2021. URL: https://doi.org/10.18154/rwth-2022-00532, doi:10.18154/rwth-2022-00532. This article has 0 citations.

  11. (miyauchi2023bag6supportsstress pages 11-13): Maho Miyauchi, Reina Matsumura, and Hiroyuki Kawahara. Bag6 supports stress fiber formation by preventing the ubiquitin-mediated degradation of rhoa. Molecular Biology of the Cell, Apr 2023. URL: https://doi.org/10.1091/mbc.e22-08-0355, doi:10.1091/mbc.e22-08-0355. This article has 7 citations and is from a domain leading peer-reviewed journal.

  12. (zhou2024bag6inhibitsinfluenza pages 1-2): Yong Zhou, Tian Li, Yunfan Zhang, Nianzhi Zhang, Yuxin Guo, Xiaoyi Gao, Wenjing Peng, Sicheng Shu, Chuankuo Zhao, Di Cui, Honglei Sun, Yipeng Sun, Jinhua Liu, Jun Tang, Rui Zhang, and Juan Pu. Bag6 inhibits influenza a virus replication by inducing viral polymerase subunit pb2 degradation and perturbing rdrp complex assembly. PLOS Pathogens, 20:e1012110, Mar 2024. URL: https://doi.org/10.1371/journal.ppat.1012110, doi:10.1371/journal.ppat.1012110. This article has 13 citations and is from a highest quality peer-reviewed journal.

  13. (zhou2024bag6inhibitsinfluenza pages 4-7): Yong Zhou, Tian Li, Yunfan Zhang, Nianzhi Zhang, Yuxin Guo, Xiaoyi Gao, Wenjing Peng, Sicheng Shu, Chuankuo Zhao, Di Cui, Honglei Sun, Yipeng Sun, Jinhua Liu, Jun Tang, Rui Zhang, and Juan Pu. Bag6 inhibits influenza a virus replication by inducing viral polymerase subunit pb2 degradation and perturbing rdrp complex assembly. PLOS Pathogens, 20:e1012110, Mar 2024. URL: https://doi.org/10.1371/journal.ppat.1012110, doi:10.1371/journal.ppat.1012110. This article has 13 citations and is from a highest quality peer-reviewed journal.

  14. (zhou2024bag6inhibitsinfluenza media f7dcee05): Yong Zhou, Tian Li, Yunfan Zhang, Nianzhi Zhang, Yuxin Guo, Xiaoyi Gao, Wenjing Peng, Sicheng Shu, Chuankuo Zhao, Di Cui, Honglei Sun, Yipeng Sun, Jinhua Liu, Jun Tang, Rui Zhang, and Juan Pu. Bag6 inhibits influenza a virus replication by inducing viral polymerase subunit pb2 degradation and perturbing rdrp complex assembly. PLOS Pathogens, 20:e1012110, Mar 2024. URL: https://doi.org/10.1371/journal.ppat.1012110, doi:10.1371/journal.ppat.1012110. This article has 13 citations and is from a highest quality peer-reviewed journal.

  15. (alhamwe2024bag6restrictspancreatic pages 1-2): Bilal Alashkar Alhamwe, Viviane Ponath, Fahd Alhamdan, Bastian Dörsam, Clara Landwehr, Manuel Linder, Kim Pauck, Sarah Miethe, Holger Garn, Florian Finkernagel, Anna Brichkina, Matthias Lauth, Dinesh Kumar Tiwari, Malte Buchholz, Daniel Bachurski, Sabrina Elmshäuser, Andrea Nist, Thorsten Stiewe, Lisa Pogge von Strandmann, Witold Szymański, Vanessa Beutgen, Johannes Graumann, Julia Teply-Szymanski, Corinna Keber, Carsten Denkert, Ralf Jacob, Christian Preußer, and Elke Pogge von Strandmann. Bag6 restricts pancreatic cancer progression by suppressing the release of il33-presenting extracellular vesicles and the activation of mast cells. Cellular and Molecular Immunology, 21:918-931, Jun 2024. URL: https://doi.org/10.1038/s41423-024-01195-1, doi:10.1038/s41423-024-01195-1. This article has 28 citations and is from a peer-reviewed journal.

  16. (alhamwe2024bag6restrictspancreatic pages 8-10): Bilal Alashkar Alhamwe, Viviane Ponath, Fahd Alhamdan, Bastian Dörsam, Clara Landwehr, Manuel Linder, Kim Pauck, Sarah Miethe, Holger Garn, Florian Finkernagel, Anna Brichkina, Matthias Lauth, Dinesh Kumar Tiwari, Malte Buchholz, Daniel Bachurski, Sabrina Elmshäuser, Andrea Nist, Thorsten Stiewe, Lisa Pogge von Strandmann, Witold Szymański, Vanessa Beutgen, Johannes Graumann, Julia Teply-Szymanski, Corinna Keber, Carsten Denkert, Ralf Jacob, Christian Preußer, and Elke Pogge von Strandmann. Bag6 restricts pancreatic cancer progression by suppressing the release of il33-presenting extracellular vesicles and the activation of mast cells. Cellular and Molecular Immunology, 21:918-931, Jun 2024. URL: https://doi.org/10.1038/s41423-024-01195-1, doi:10.1038/s41423-024-01195-1. This article has 28 citations and is from a peer-reviewed journal.

  17. (carretero2024differentiallunggene pages 8-10): Laura Sánchez Carretero, Adele Chloe Cardeñosa Pérez, Germán Peces-Barba, and Sandra Pérez-Rial. Differential lung gene expression identified zscan2 and bag6 as novel tissue repair players in an experimental copd model. PLOS ONE, 19:e0309166, Aug 2024. URL: https://doi.org/10.1371/journal.pone.0309166, doi:10.1371/journal.pone.0309166. This article has 2 citations and is from a peer-reviewed journal.

  18. (alhamwe2024bag6restrictspancreatic pages 11-12): Bilal Alashkar Alhamwe, Viviane Ponath, Fahd Alhamdan, Bastian Dörsam, Clara Landwehr, Manuel Linder, Kim Pauck, Sarah Miethe, Holger Garn, Florian Finkernagel, Anna Brichkina, Matthias Lauth, Dinesh Kumar Tiwari, Malte Buchholz, Daniel Bachurski, Sabrina Elmshäuser, Andrea Nist, Thorsten Stiewe, Lisa Pogge von Strandmann, Witold Szymański, Vanessa Beutgen, Johannes Graumann, Julia Teply-Szymanski, Corinna Keber, Carsten Denkert, Ralf Jacob, Christian Preußer, and Elke Pogge von Strandmann. Bag6 restricts pancreatic cancer progression by suppressing the release of il33-presenting extracellular vesicles and the activation of mast cells. Cellular and Molecular Immunology, 21:918-931, Jun 2024. URL: https://doi.org/10.1038/s41423-024-01195-1, doi:10.1038/s41423-024-01195-1. This article has 28 citations and is from a peer-reviewed journal.

  19. (OpenTargets Search: -BAG6): Open Targets Query (-BAG6, 5 results). Buniello, A. et al. (2025). Open Targets Platform: facilitating therapeutic hypotheses building in drug discovery. Nucleic Acids Research.

  20. (roboti2022mitochondrialantiviralsignallingprotein pages 15-15): Peristera Roboti, Craig Lawless, and Stephen High. Mitochondrial antiviral-signalling protein is a client of the bag6 protein quality control complex. Journal of Cell Science, May 2022. URL: https://doi.org/10.1242/jcs.259596, doi:10.1242/jcs.259596. This article has 1 citations and is from a domain leading peer-reviewed journal.

  21. (you2020paqr9modulatesbag6mediated pages 15-16): Xue You, Yijun Lin, Yongfan Hou, Lijiao Xu, Qianqian Cao, and Yan Chen. Paqr9 modulates bag6-mediated protein quality control of mislocalized membrane proteins. Biochemical Journal, 477:477-489, Jan 2020. URL: https://doi.org/10.1042/bcj20190620, doi:10.1042/bcj20190620. This article has 10 citations and is from a domain leading peer-reviewed journal.

  22. (hagiwara2023proteotoxicstressesstimulate pages 14-15): Takumi Hagiwara, Ryosuke Minami, Chizuru Ushio, Naoto Yokota, and Hiroyuki Kawahara. Proteotoxic stresses stimulate dissociation of ubl4a from the tail-anchored protein recognition complex. Biochemical Journal, 480:1583-1598, Oct 2023. URL: https://doi.org/10.1042/bcj20230267, doi:10.1042/bcj20230267. This article has 0 citations and is from a domain leading peer-reviewed journal.

Citations

  1. guna2018transmembranedomainrecognition pages 4-6
  2. hagiwara2023proteotoxicstressesstimulate pages 3-5
  3. wang2021analysisofthe pages 80-84
  4. hagiwara2023proteotoxicstressesstimulate pages 10-12
  5. roboti2022mitochondrialantiviralsignallingprotein pages 1-2
  6. carretero2024differentiallunggene pages 8-10
  7. abildgaard2020cochaperonesintargeting pages 9-11
  8. hagiwara2023proteotoxicstressesstimulate pages 1-3
  9. abildgaard2020cochaperonesintargeting pages 21-23
  10. roboti2022mitochondrialantiviralsignallingprotein pages 15-15
  11. hagiwara2023proteotoxicstressesstimulate pages 14-15
  12. https://doi.org/10.1042/BCJ20230267
  13. https://doi.org/10.1242/jcs.259596
  14. https://doi.org/10.1091/mbc.e22-08-0355
  15. https://doi.org/10.1371/journal.ppat.1012110
  16. https://doi.org/10.1038/s41423-024-01195-1
  17. https://doi.org/10.1371/journal.pone.0309166
  18. https://doi.org/10.1016/j.cub.2018.02.004
  19. https://doi.org/10.3390/biom10081141
  20. https://doi.org/10.1042/bcj20230267,
  21. https://doi.org/10.1091/mbc.e22-08-0355,
  22. https://doi.org/10.1371/journal.ppat.1012110,
  23. https://doi.org/10.1016/j.cub.2018.02.004,
  24. https://doi.org/10.3390/biom10081141,
  25. https://doi.org/10.1242/jcs.259596,
  26. https://doi.org/10.18154/rwth-2022-00532,
  27. https://doi.org/10.1038/s41423-024-01195-1,
  28. https://doi.org/10.1371/journal.pone.0309166,
  29. https://doi.org/10.1042/bcj20190620,

📚 Additional Documentation

Notes

(BAG6-notes.md)

BAG6 notes

PN framing

  • BAG6 is a proteostasis boundary-case gene rather than a canonical BAG-family Hsp70 nucleotide-exchange factor. Its C-terminal BAG-like region is a mock/noncanonical BAG domain and is better treated as a UBL4A-binding BAGS module than as a canonical Hsp70-binding BAG domain [PMID:25535373 Bag6 complex contains a minimal tail-anchor-targeting module and a mock BAG domain, "Bag6-BAG is not a canonical BAG domain"] [PMID:25713138 Structure of a BAG6-Ubl4a complex reveals a novel binding interface that functions in tail-anchored protein biogenesis, "previously classified as a BAG domain, were completely distinct from those of the canonical BAG domain"].
  • This matches the PN caution that BAG6 sits between ER proteostasis/GET-pathway targeting and UPS-associated chaperone roles, rather than fitting cleanly into a standard BAG-domain cochaperone bucket.

Core proteostasis biology

  • The strongest BAG6 function is cytosolic handling of hydrophobic client proteins. BAG6, UBL4A, and TRC35 form a ribosome-associated complex that captures tail-anchored membrane proteins and passes them to TRC40 for ER delivery [PMID:20676083 A ribosome-associating factor chaperones tail-anchored membrane proteins, "facilitates TA protein capture by TRC40"] [PMID:25535373 Bag6 complex contains a minimal tail-anchor-targeting module and a mock BAG domain, "facilitates tail-anchored substrate transfer from small glutamine-rich tetratricopeptide repeat-containing protein α to TRC40"].
  • BAG6 is also an ATP-independent holdase for hydrophobic quality-control clients. In ERAD, it keeps retrotranslocated substrates soluble and improves their proteasomal disposal [PMID:21636303 A ubiquitin ligase-associated chaperone holdase maintains polypeptides in soluble states for proteasome degradation, "maintains polypeptides in soluble states for proteasome degradation"] [PMID:23665563 A ubiquitin-like domain recruits an oligomeric chaperone to a retrotranslocation complex in endoplasmic reticulum-associated degradation, "anchor a disordered chaperone oligomer to the site of retrotranslocation to prevent protein aggregation in ERAD"].
  • BAG6 couples targeting and degradation for mislocalized membrane proteins. Clients that fail productive targeting can be ubiquitinated through BAG6-linked machinery rather than simply handed back to folding systems [PMID:21743475 Protein targeting and degradation are coupled for elimination of mislocalized proteins, "A subset of these Bag6 complex clients are transferred to TRC40 for insertion into the membrane, whereas the remainder are rapidly ubiquitinated"] [PMID:24981174 Cytosolic quality control of mislocalized proteins requires RNF126 recruitment to Bag6, "RNF126 is the primary Bag6-dependent ligase"] [PMID:23129660 SGTA antagonizes BAG6-mediated protein triage, "SGTA antagonizes BAG6-mediated protein triage"].
  • ER-stress-induced pre-emptive quality control is supported but should stay narrower than generic ERAD. Bag6 contributes to degradation of rerouted ER pQC substrates during stress [PMID:26565908 Pre-emptive Quality Control Protects the ER from Protein Overload via the Proximity of ERAD Components and SRP, "Bag6 and p97 contribute to the degradation of ER pQC substrates"].

Contextual, non-core roles

  • BAG6 has a documented nuclear stress-response role, promoting p300-mediated p53 acetylation and DNA-damage-associated apoptosis [PMID:17403783 HLA-B-associated transcript 3 (Bat3)/Scythe is essential for p300-mediated acetylation of p53, "Bat3 is required for the acetylation of p53 in response to DNA damage"]. This is real but not the core proteostasis identity.
  • Exosomal BAG6 can function as an NKp30 ligand and modulate NK-cell activation in tumor or dendritic-cell contexts [PMID:18055229 Human leukocyte antigen-B-associated transcript 3 is released from tumor cells and engages the NKp30 receptor on natural killer cells, "BAT3 ... engages the NKp30 receptor on natural killer cells"] [PMID:18852879 Dendritic cells release HLA-B-associated transcript-3 positive exosomes to regulate natural killer function, "BAT3-positive exosomes ... activation of NK cell-mediated cytokine release"]. This should be kept distinct from the core GET/UPS quality-control role.
  • BAG6 is genuinely nucleo-cytoplasmic, and TRC35/GET4 binding helps retain it in the cytosol where the main proteostasis functions occur [PMID:29042515 Structural basis for regulation of the nucleo-cytoplasmic distribution of Bag6 by TRC35, "TRC35 binding is critical ... to retain Bag6 in the cytosol"].

2026-05-29 - PROTEOSTASIS PN ER-protein-transport pass

  • PN places BAG6 under ER proteostasis > Protein transport > GET pathway component. The narrow GET-pathway mapping to GO:0006620 post-translational protein targeting to endoplasmic reticulum membrane is appropriate and is already represented in the local review/GOA.
  • The parent ER proteostasis > Protein transport mapping would additionally propagate broad GO:0015031 protein transport to BAG6. This is directionally related but less informative than the reviewed BAG6 terms, especially GO:0071816 tail-anchored membrane protein insertion into ER membrane and GO:0006620 post-translational protein targeting to endoplasmic reticulum membrane [PMID:20676083 "facilitates TA protein capture by TRC40"; PMID:25535373 "facilitates tail-anchored substrate transfer"].
  • Working curation conclusion: keep the GET-pathway-specific propagation for BAG6, but do not add the broad parent GO:0015031 protein transport candidate.

Pn Notes

(BAG6-pn-notes.md)

BAG6 PN Consistency Notes

  • Generated: 2026-06-18
  • Project: PROTEOSTASIS
  • Scope: priority-only PN rereview against local AIGR review and available deep-research artifacts
  • UniProt: P46379
  • AIGR review status: COMPLETE
  • Priority category: multibranch_boundary_case
  • Local AIGR project status: local_review_complete_not_phase1
  • Related project: ER_PHAGY.md

Source Files Checked

Deep Research Files

AIGR Review Snapshot

  • Description: BAG6 is a multifunctional nucleo-cytoplasmic proteostasis factor that sits at the boundary between the GET pathway and ubiquitin-proteasome-associated quality control. In the cytosol, the BAG6/UBL4A/GET4 (BAT3) complex captures hydrophobic tail-anchored or otherwise mislocalized secretory proteins, promotes their handoff to TRC40/ASNA1 for post-translational delivery to the endoplasmic reticulum, and routes failed clients toward ubiquitin-proteasome degradation. BAG6 also acts as a holdase for retrotranslocated ERAD substrates and contributes to ER stress-induced pre-emptive quality control. Additional nuclear and extracellular roles, including p53 acetylation after DNA damage and exosomal NKp30 ligand activity, are supported in specific contexts but are not the core conserved proteostasis functions.
  • Existing/core annotation action counts: ACCEPT: 30; KEEP_AS_NON_CORE: 32; MARK_AS_OVER_ANNOTATED: 4; MODIFY: 9; REMOVE: 29

PN Consistency Summary

  • Consistency: Priority-only record; no phase-1 dossier section exists yet. Local review status is local_review_complete_not_phase1. PN placement: ER proteostasis > GET pathway component; UPS > associated non-Ub enzyme chaperone; UPS > UBL domain > chaperones and related. Main issue: Review supports specific GET, ERAD, and holdase roles but not broad parent protein-transport propagation
  • PN story / NEW pressure: Current projection rows: GO:0006620 post-translational protein targeting to endoplasmic reticulum membrane (already_in_goa_exact).
  • Mapping strategy: Use narrow, evidence-aligned PN mappings, but preserve exceptions against broad parent propagation across mixed proteostasis branches.
  • Verdict: Add to boundary/phase1 tracking; keep broad protein-transport exception and positive GET/ERAD bridge outcomes

Priority Review Context

  • Category: multibranch_boundary_case
  • PN annotations: ER proteostasis > GET pathway component; UPS > associated non-Ub enzyme chaperone; UPS > UBL domain > chaperones and related
  • Why interesting: Review supports specific GET, ERAD, and holdase roles but not broad parent protein-transport propagation
  • Suggested next step: Add to boundary/phase1 tracking; keep broad protein-transport exception and positive GET/ERAD bridge outcomes
  • Related project: ER_PHAGY.md

PN Projection Rows

  • GO:0006620 post-translational protein targeting to endoplasmic reticulum membrane - already_in_goa_exact; scope=ok_for_propagation_to_go; mapping=er_proteostasis.yaml; PN=ER proteostasis|Protein transport|GET pathway component

PN Dossier Context

No phase-1 dossier exists for this priority-only gene. This note preserves the current PROTEOSTASIS boundary or exception decision and should be superseded by a dossier section if the gene is promoted into a full phase-1 batch.

Note

This file is generated from the current PROTEOSTASIS priority table, PN projection outputs, and local gene-review artifacts. Edit those source records rather than this generated note when correcting the underlying curation.

📄 View Raw YAML

id: P46379
gene_symbol: BAG6
product_type: PROTEIN
taxon:
  id: NCBITaxon:9606
  label: Homo sapiens
description: BAG6 is a multifunctional nucleo-cytoplasmic proteostasis factor that
  sits at the boundary between the GET pathway and ubiquitin-proteasome-associated
  quality control. In the cytosol, the BAG6/UBL4A/GET4 (BAT3) complex captures hydrophobic
  tail-anchored or otherwise mislocalized secretory proteins, promotes their handoff
  to TRC40/ASNA1 for post-translational delivery to the endoplasmic reticulum, and
  routes failed clients toward ubiquitin-proteasome degradation. BAG6 also acts as
  a holdase for retrotranslocated ERAD substrates and contributes to ER stress-induced
  pre-emptive quality control. Additional nuclear and extracellular roles, including
  p53 acetylation after DNA damage and exosomal NKp30 ligand activity, are supported
  in specific contexts but are not the core conserved proteostasis functions.
alternative_products:
- name: '1'
  id: P46379-1
- name: '2'
  id: P46379-2
  sequence_note: VSP_015695
- name: '3'
  id: P46379-3
  sequence_note: VSP_015695, VSP_030519
- name: '4'
  id: P46379-4
  sequence_note: VSP_015695, VSP_045910, VSP_045911,
- name: '5'
  id: P46379-5
  sequence_note: VSP_015695, VSP_045913
references:
- id: file:human/BAG6/BAG6-notes.md
  title: BAG6 review notes
- id: file:human/BAG6/BAG6-deep-research-falcon.md
  title: Falcon deep research report for BAG6
  findings:
  - statement: Falcon research supports BAG6 as a cytosolic BAG6/UBL4A/TRC35 holdase and adaptor linking tail-anchored protein targeting with ERAD/proteasome quality control.
    supporting_text: BAG6 is embedded in this pathway as both a substrate-holding factor and a quality-control adaptor
- id: GO_REF:0000024
  title: Manual transfer of experimentally-verified manual GO annotation data to orthologs
    by curator judgment of sequence similarity
- id: GO_REF:0000033
  title: Annotation inferences using phylogenetic trees
- id: GO_REF:0000044
  title: Gene Ontology annotation based on UniProtKB/Swiss-Prot Subcellular Location
    vocabulary mapping, accompanied by conservative changes to GO terms applied by
    UniProt
- id: GO_REF:0000052
  title: Gene Ontology annotation based on curation of immunofluorescence data from
    the Human Protein Atlas
- id: GO_REF:0000107
  title: Automatic transfer of experimentally verified manual GO annotation data to
    orthologs using Ensembl Compara
- id: GO_REF:0000120
  title: Combined Automated Annotation using Multiple IEA Methods
- id: PMID:20676083
  title: A ribosome-associating factor chaperones tail-anchored membrane proteins.
- id: PMID:21636303
  title: A ubiquitin ligase-associated chaperone holdase maintains polypeptides in
    soluble states for proteasome degradation.
- id: PMID:21743475
  title: Protein targeting and degradation are coupled for elimination of mislocalized
    proteins.
- id: PMID:23129660
  title: SGTA antagonizes BAG6-mediated protein triage.
- id: PMID:23665563
  title: A ubiquitin-like domain recruits an oligomeric chaperone to a retrotranslocation
    complex in endoplasmic reticulum-associated degradation.
- id: PMID:24424410
  title: USP13 antagonizes gp78 to maintain functionality of a chaperone in ER-associated
    degradation.
- id: PMID:24981174
  title: Cytosolic quality control of mislocalized proteins requires RNF126 recruitment
    to Bag6.
- id: PMID:25535373
  title: Bag6 complex contains a minimal tail-anchor-targeting module and a mock BAG
    domain.
- id: PMID:25713138
  title: 'Structure of a BAG6 (Bcl-2-associated athanogene 6)-Ubl4a (ubiquitin-like protein 4a) complex reveals a novel binding interface that functions in tail-anchored protein biogenesis.'
- id: PMID:29042515
  title: Structural basis for regulation of the nucleo-cytoplasmic distribution of
    Bag6 by TRC35.
- id: PMID:17403783
  title: HLA-B-associated transcript 3 (Bat3)/Scythe is essential for p300-mediated
    acetylation of p53.
- id: PMID:18055229
  title: Human leukocyte antigen-B-associated transcript 3 is released from tumor
    cells and engages the NKp30 receptor on natural killer cells.
- id: PMID:18852879
  title: Dendritic cells release HLA-B-associated transcript-3 positive exosomes to
    regulate natural killer function.
- id: PMID:14667819
  title: Analysis of a high-throughput yeast two-hybrid system and its use to predict
    the function of intracellular proteins encoded within the human MHC class III
    region.
- id: PMID:14960581
  title: Ricin triggers apoptotic morphological changes through caspase-3 cleavage
    of BAT3.
- id: PMID:16189514
  title: Towards a proteome-scale map of the human protein-protein interaction network.
- id: PMID:18765639
  title: BAT3 and SET1A form a complex with CTCFL/BORIS to modulate H3K4 histone dimethylation
    and gene expression.
- id: PMID:19946888
  title: Defining the membrane proteome of NK cells.
- id: PMID:21903422
  title: Mapping a dynamic innate immunity protein interaction network regulating
    type I interferon production.
- id: PMID:22046132
  title: 'The SARS-coronavirus-host interactome: identification of cyclophilins as
    target for pan-coronavirus inhibitors.'
- id: PMID:22807449
  title: The stalk domain and the glycosylation status of the activating natural killer
    cell receptor NKp30 are important for ligand binding.
- id: PMID:23246001
  title: SGTA recognizes a noncanonical ubiquitin-like domain in the Bag6-Ubl4A-Trc35
    complex to promote endoplasmic reticulum-associated degradation.
- id: PMID:25036637
  title: A quantitative chaperone interaction network reveals the architecture of
    cellular protein homeostasis pathways.
- id: PMID:25416956
  title: A proteome-scale map of the human interactome network.
- id: PMID:26496610
  title: A human interactome in three quantitative dimensions organized by stoichiometries
    and abundances.
- id: PMID:26876100
  title: Selective Binding of AIRAPL Tandem UIMs to Lys48-Linked Tri-Ubiquitin Chains.
- id: PMID:27113755
  title: UBQLN4 recognizes mislocalized transmembrane domain proteins and targets
    these to proteasomal degradation.
- id: PMID:31515488
  title: Extensive disruption of protein interactions by genetic variants across the
    allele frequency spectrum in human populations.
- id: PMID:33961781
  title: Dual proteome-scale networks reveal cell-specific remodeling of the human
    interactome.
- id: PMID:35271311
  title: 'OpenCell: Endogenous tagging for the cartography of human cellular organization.'
- id: PMID:40105103
  title: Definition of the human mitochondrial TOM interactome reveals TRABD as a
    new interacting protein.
- id: PMID:26565908
  title: Pre-emptive Quality Control Protects the ER from Protein Overload via the
    Proximity of ERAD Components and SRP.
existing_annotations:
- term:
    id: GO:0036503
    label: ERAD pathway
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: Human BAG6 literature supports BAG6 complex function in ER-associated
      degradation (ERAD), so the phylogenetic propagation is directionally correct
      and consistent with the core proteostasis role.
    action: ACCEPT
    reason: Bag6 keeps retrotranslocated hydrophobic clients soluble and engaged with
      ERAD machinery, matching ERAD pathway membership.
    supported_by:
    - reference_id: PMID:20676083
    - reference_id: PMID:21636303
    - reference_id: PMID:25535373
    - reference_id: file:human/BAG6/BAG6-notes.md
- term:
    id: GO:0031593
    label: polyubiquitin modification-dependent protein binding
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: BAG6 operates in ubiquitin-linked quality control, but the reviewed core
      papers support ligase/DUB recruitment and client triage rather than a well-demonstrated
      direct polyubiquitin-binding activity.
    action: REMOVE
    reason: The conserved BAG6 literature does not establish polyubiquitin-selective
      binding as a defensible core molecular function.
- term:
    id: GO:0051787
    label: misfolded protein binding
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: 'Phylogenetic propagation agrees with the human holdase literature: BAG6
      preferentially captures aggregation-prone hydrophobic or misfolded clients during
      quality control.'
    action: ACCEPT
    reason: This matches the experimentally supported BAG6 holdase role in ERAD and
      mislocalized-protein triage.
    supported_by:
    - reference_id: PMID:21636303
    - reference_id: file:human/BAG6/BAG6-notes.md
- term:
    id: GO:0071818
    label: BAT3 complex
  evidence_type: IBA
  original_reference_id: GO_REF:0000033
  review:
    summary: The phylogenetic BAT3 complex annotation agrees with multiple human studies
      placing BAG6 in a heterotrimeric BAG6/UBL4A/GET4(TRC35) complex.
    action: ACCEPT
    reason: Complex membership is well supported and central to both GET-pathway and
      UPS-adjacent BAG6 functions.
    supported_by:
    - reference_id: PMID:20676083
    - reference_id: PMID:25535373
- term:
    id: GO:0001822
    label: kidney development
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: Developmental annotations are plausible downstream consequences from
      mammalian loss-of-function studies, but they are not BAG6's core conserved proteostasis
      role.
    action: KEEP_AS_NON_CORE
    reason: Keep as contextual biology rather than a core molecular/process function.
- term:
    id: GO:0006511
    label: ubiquitin-dependent protein catabolic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: BAG6 helps channel selected clients toward ubiquitin-dependent degradation,
      but this broad process term loses the more informative GET/ERAD/pre-emptive-QC
      context.
    action: MARK_AS_OVER_ANNOTATED
    reason: Retain more specific proteostasis terms instead of this umbrella catabolic
      label.
- term:
    id: GO:0007130
    label: synaptonemal complex assembly
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: This reproductive annotation reflects secondary mammalian biology rather
      than BAG6's central proteostasis function.
    action: KEEP_AS_NON_CORE
    reason: Treat as contextual/non-core.
- term:
    id: GO:0007283
    label: spermatogenesis
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: This reproductive-process annotation is best treated as contextual because
      BAG6's conserved role is proteostasis triage, not gametogenesis machinery.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0007420
    label: brain development
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: Any developmental nervous-system consequence is downstream/contextual
      rather than part of BAG6's main evolved role.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0030324
    label: lung development
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: This developmental annotation reflects contextual organismal phenotypes
      rather than BAG6's core GET/UPS-boundary biology.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0030544
    label: Hsp70 protein binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: This looks like over-propagation from BAG family naming. BAG6 is explicitly
      not a canonical BAG-domain Hsp70 nucleotide-exchange factor.
    action: REMOVE
    reason: Structural work shows the BAG6 C terminus is a mock/noncanonical BAG-similar
      domain, so Hsp70 binding should not be assumed from family membership.
    supported_by:
    - reference_id: PMID:25535373
    - reference_id: PMID:25713138
    - reference_id: file:human/BAG6/BAG6-notes.md
- term:
    id: GO:0031593
    label: polyubiquitin modification-dependent protein binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: This broad automated propagation is not well grounded in the direct BAG6
      mechanistic literature.
    action: REMOVE
    reason: The reviewed papers support client triage with E3s and DUBs, not a specific
      polyubiquitin-binding activity.
- term:
    id: GO:0032435
    label: negative regulation of proteasomal ubiquitin-dependent protein catabolic
      process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: This regulatory proteolysis term reflects secondary spermatogenic/HSPA2
      biology rather than BAG6's central proteostasis role.
    action: KEEP_AS_NON_CORE
    reason: Keep as contextual/non-core.
- term:
    id: GO:0042981
    label: regulation of apoptotic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: Apoptosis-related effects exist in some contexts, but this broad automated
      term is not the best summary of BAG6 function.
    action: KEEP_AS_NON_CORE
    reason: Retain only as non-core context.
- term:
    id: GO:0043161
    label: proteasome-mediated ubiquitin-dependent protein catabolic process
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: BAG6 contributes to selected proteasomal quality-control routes, but
      this automated term is broader and less informative than the specific BAG6-supported
      processes.
    action: MARK_AS_OVER_ANNOTATED
    reason: Prefer ERAD, tail-anchored insertion, or ER-stress pre-emptive quality
      control terms.
- term:
    id: GO:0045861
    label: negative regulation of proteolysis
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: This negative-regulation term reflects context-specific stabilization
      biology rather than the main BAG6 proteostasis program.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0045995
    label: regulation of embryonic development
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: Embryonic-development phenotypes are contextual downstream consequences
      and not the core conserved BAG6 role.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0050821
    label: protein stabilization
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: Protein stabilization is context-specific for selected partners and not
      the best core descriptor of BAG6.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0070059
    label: intrinsic apoptotic signaling pathway in response to endoplasmic reticulum
      stress
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: ER-stress apoptosis is a context-dependent consequence, not BAG6's principal
      conserved function.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0070628
    label: proteasome binding
  evidence_type: IEA
  original_reference_id: GO_REF:0000107
  review:
    summary: The reviewed BAG6 papers place clients en route to the proteasome but
      do not establish a clean direct proteasome-binding activity.
    action: REMOVE
    reason: This term is too specific for the available evidence.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:14667819
  review:
    summary: The source interaction is too generic to retain as a useful BAG6 annotation.
    action: REMOVE
    reason: GO:0005515 is uninformative here and BAG6 already has more specific mechanistically
      grounded annotations.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:16189514
  review:
    summary: The source interaction is too generic to retain as a useful BAG6 annotation.
    action: REMOVE
    reason: GO:0005515 is uninformative here and BAG6 already has more specific mechanistically
      grounded annotations.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:18852879
  review:
    summary: The exosome/NK paper supports a contextual receptor-ligand role, not
      a useful standalone generic protein-binding annotation.
    action: REMOVE
    reason: Retain the more specific immune-context terms instead.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:21903422
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:22046132
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:22807449
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25036637
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25416956
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:26496610
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:31515488
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:33961781
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:35271311
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005576
    label: extracellular region
  evidence_type: IEA
  original_reference_id: GO_REF:0000044
  review:
    summary: Extracellular release is real but limited to stress/tumor exosome contexts
      rather than BAG6's main intracellular proteostasis role.
    action: KEEP_AS_NON_CORE
    reason: Keep as contextual, non-core localization.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: BAG6 is a bona fide nucleo-cytoplasmic protein, and nuclear localization
      is repeatedly observed in the literature and UniProt curation.
    action: ACCEPT
    reason: Nuclear localization is real even though the principal PN role is cytosolic.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IEA
  original_reference_id: GO_REF:0000120
  review:
    summary: Cytosolic localization is central to BAG6 GET/ERAD/mislocalized-protein
      triage functions.
    action: ACCEPT
    reason: The core proteostasis role depends on cytosolic BAG6 complex localization.
- term:
    id: GO:0005654
    label: nucleoplasm
  evidence_type: IDA
  original_reference_id: GO_REF:0000052
  review:
    summary: Nucleoplasmic signal is compatible with BAG6's documented nuclear pool
      and DNA-damage-associated functions.
    action: ACCEPT
    reason: This is a defensible subnuclear localization.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IDA
  original_reference_id: GO_REF:0000052
  review:
    summary: Independent localization evidence also places BAG6 in the cytosol.
    action: ACCEPT
    reason: This is fully consistent with the core proteostasis model.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:40105103
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0140597
    label: protein carrier chaperone
  evidence_type: IDA
  original_reference_id: PMID:21636303
  review:
    summary: This is one of the strongest BAG6 annotations. The 2011 Mol Cell study
      directly supports a holdase/carrier role for retrotranslocated hydrophobic clients.
    action: ACCEPT
    reason: Bag6 maintains aggregation-prone clients in an unfolded yet soluble state
      and helps deliver them within quality-control pathways.
    supported_by:
    - reference_id: PMID:21636303
    - reference_id: PMID:20676083
    - reference_id: file:human/BAG6/BAG6-deep-research-falcon.md
      supporting_text: BAG6 is widely characterized as a chaperone/holdase that binds exposed hydrophobic segments
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: NAS
  original_reference_id: PMID:25535373
  review:
    summary: Cytosolic localization is consistent with the tail-anchor-targeting complex
      architecture and broader BAG6 literature, even if this individual assertion
      is author statement-level.
    action: ACCEPT
    reason: The BAG6 complex acts in the cytosol before ER insertion or proteasomal
      routing.
- term:
    id: GO:0006511
    label: ubiquitin-dependent protein catabolic process
  evidence_type: IDA
  original_reference_id: PMID:20676083
  review:
    summary: PMID:20676083 is about ribosome-associated capture and handoff of tail-anchored
      proteins to TRC40, not a generic ubiquitin-dependent catabolic process.
    action: MODIFY
    reason: Replace with the specific GET-pathway process terms directly supported
      by the paper; the separate ribosome-binding molecular function is already captured
      by its own annotation.
    proposed_replacement_terms:
    - id: GO:0071816
      label: tail-anchored membrane protein insertion into ER membrane
- term:
    id: GO:0006620
    label: post-translational protein targeting to endoplasmic reticulum membrane
  evidence_type: IDA
  original_reference_id: PMID:25535373
  review:
    summary: This matches the PN GET-pathway mapping and the structural/biochemical
      literature on tail-anchor targeting.
    action: ACCEPT
    reason: BAG6 participates in post-translational delivery of tail-anchored proteins
      toward the ER membrane.
    supported_by:
    - reference_id: PMID:25535373
    - reference_id: PMID:20676083
    - reference_id: file:human/BAG6/BAG6-deep-research-falcon.md
      supporting_text: BAG6 is embedded in this pathway as both a substrate-holding factor and a quality-control adaptor
- term:
    id: GO:0031647
    label: regulation of protein stability
  evidence_type: IDA
  original_reference_id: PMID:21636303
  review:
    summary: The paper supports holdase/chaperone behavior rather than a broad generic
      regulation-of-stability process.
    action: MODIFY
    reason: A molecular chaperone/carrier term is a more faithful representation of
      the evidence.
    proposed_replacement_terms:
    - id: GO:0140597
      label: protein carrier chaperone
- term:
    id: GO:0048018
    label: receptor ligand activity
  evidence_type: IDA
  original_reference_id: PMID:18852879
  review:
    summary: Exosomal BAG6 can act as an NKp30 ligand, but this immune signaling role
      is contextual and not the core conserved BAG6 function.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core context.
    supported_by:
    - reference_id: PMID:18852879
    - reference_id: PMID:18055229
- term:
    id: GO:0051132
    label: NK T cell activation
  evidence_type: IDA
  original_reference_id: PMID:18852879
  review:
    summary: The paper concerns NK-cell activation, not NK T-cell activation.
    action: MODIFY
    reason: Replace with natural killer cell activation to match the actual experiment.
    proposed_replacement_terms:
    - id: GO:0030101
      label: natural killer cell activation
- term:
    id: GO:0036503
    label: ERAD pathway
  evidence_type: NAS
  original_reference_id: PMID:21636303
  review:
    summary: Although this is an author statement-level annotation, the underlying
      study directly demonstrates a BAG6 holdase role that improves ERAD efficiency.
    action: ACCEPT
    reason: The evidence supports ERAD pathway participation.
    supported_by:
    - reference_id: file:human/BAG6/BAG6-deep-research-falcon.md
      supporting_text: BAG6 helps route failed membrane/secretory protein biogenesis products and certain ERAD substrates into ubiquitin-proteasome degradation
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25713138
  review:
    summary: The structural paper supports complex architecture and adaptor function,
      not a useful generic protein-binding annotation.
    action: REMOVE
    reason: Retain the specific adaptor/core-complex annotations instead.
- term:
    id: GO:0060090
    label: molecular adaptor activity
  evidence_type: EXP
  original_reference_id: PMID:25713138
  review:
    summary: BAG6 organizes UBL4A and TRC35/GET4 and supports substrate handoff, which
      is well captured by molecular adaptor activity.
    action: ACCEPT
    reason: This term fits BAG6's bridge/scaffold role at the GET-pathway and cytosolic
      quality-control boundary.
    supported_by:
    - reference_id: PMID:25713138
    - reference_id: PMID:25535373
    - reference_id: file:human/BAG6/BAG6-deep-research-falcon.md
      supporting_text: BAG6’s C-terminal region is a structural part of the substrate-loading complex
- term:
    id: GO:0140677
    label: molecular function activator activity
  evidence_type: IEP
  original_reference_id: PMID:25713138
  review:
    summary: The cited paper does not establish BAG6 as a direct molecular-function
      activator of another enzyme or receptor.
    action: REMOVE
    reason: This annotation overstates the structural/adaptor evidence.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:27113755
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0045995
    label: regulation of embryonic development
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Orthology-based developmental regulation is plausible but not part of
      BAG6's core proteostasis identity.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core context.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:29042515
  review:
    summary: This paper is informative for localization regulation, but GO:0005515
      remains too generic.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:29042515
  review:
    summary: This study directly addresses BAG6 nucleo-cytoplasmic partitioning and
      supports nuclear localization.
    action: ACCEPT
    reason: Nuclear BAG6 is real and regulated by TRC35/GET4 binding.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IDA
  original_reference_id: PMID:29042515
  review:
    summary: This study directly supports cytosolic retention of BAG6 by TRC35/GET4.
    action: ACCEPT
    reason: Cytosolic BAG6 is the state associated with core proteostasis functions.
- term:
    id: GO:0071816
    label: tail-anchored membrane protein insertion into ER membrane
  evidence_type: IDA
  original_reference_id: PMID:25535373
  review:
    summary: The minimal BAG6 complex directly supports transfer of tail-anchored
      substrates into the TRC40/GET pathway.
    action: ACCEPT
    reason: This is core PN-supported biology.
    supported_by:
    - reference_id: PMID:25535373
    - reference_id: PMID:20676083
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:26876100
  review:
    summary: This generic interaction term adds no useful BAG6-specific functional
      information.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:14960581
  review:
    summary: The ricin paper is informative for apoptosis context, but GO:0005515
      remains too generic.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0061857
    label: endoplasmic reticulum stress-induced pre-emptive quality control
  evidence_type: IMP
  original_reference_id: PMID:26565908
  review:
    summary: This is a specific stress-responsive proteostasis process that the paper
      directly supports.
    action: ACCEPT
    reason: Bag6 contributes to degradation of rerouted ER pre-emptive quality-control
      substrates under ER stress.
    supported_by:
    - reference_id: PMID:26565908
- term:
    id: GO:0010498
    label: proteasomal protein catabolic process
  evidence_type: IMP
  original_reference_id: PMID:26565908
  review:
    summary: The specific process shown in the paper is ER stress-induced pre-emptive
      quality control, not a generic proteasomal catabolic pathway in isolation.
    action: MARK_AS_OVER_ANNOTATED
    reason: Keep the more specific GO:0061857 annotation instead.
- term:
    id: GO:0036503
    label: ERAD pathway
  evidence_type: IMP
  original_reference_id: PMID:26565908
  review:
    summary: PMID:26565908 is about ER stress-induced pre-emptive quality control,
      which is related to but distinct from canonical ERAD.
    action: MODIFY
    reason: Replace with the specific ER pQC term.
    proposed_replacement_terms:
    - id: GO:0061857
      label: endoplasmic reticulum stress-induced pre-emptive quality control
- term:
    id: GO:0016020
    label: membrane
  evidence_type: HDA
  original_reference_id: PMID:19946888
  review:
    summary: A broad membrane term is too imprecise for BAG6, which is mainly cytosolic/nuclear
      with regulated ER-membrane association in specific QC contexts.
    action: MARK_AS_OVER_ANNOTATED
    reason: If membrane association is retained, it should be ER-membrane-specific
      rather than generic membrane.
- term:
    id: GO:0002429
    label: immune response-activating cell surface receptor signaling pathway
  evidence_type: IDA
  original_reference_id: PMID:18852879
  review:
    summary: This immune signaling annotation is supported in exosome/NK-cell contexts
      but is clearly contextual rather than core BAG6 biology.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
    supported_by:
    - reference_id: PMID:18852879
    - reference_id: PMID:18055229
- term:
    id: GO:0005102
    label: signaling receptor binding
  evidence_type: IPI
  original_reference_id: PMID:18852879
  review:
    summary: The immune paper is better captured by receptor ligand activity than
      a generic signaling receptor binding term.
    action: MODIFY
    reason: Replace with receptor ligand activity.
    proposed_replacement_terms:
    - id: GO:0048018
      label: receptor ligand activity
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:14960581
  review:
    summary: The ricin study also observed nuclear BAG6, consistent with BAG6's established
      nucleo-cytoplasmic distribution.
    action: ACCEPT
    reason: Nuclear localization is defensible.
- term:
    id: GO:0006915
    label: apoptotic process
  evidence_type: IDA
  original_reference_id: PMID:14960581
  review:
    summary: Ricin-induced apoptosis is a real contextual role but not BAG6's core
      conserved proteostasis function.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0030101
    label: natural killer cell activation
  evidence_type: IDA
  original_reference_id: PMID:18852879
  review:
    summary: Natural killer cell activation is experimentally supported for exosomal
      BAG6 but remains a contextual immune role.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
    supported_by:
    - reference_id: PMID:18852879
    - reference_id: PMID:18055229
- term:
    id: GO:0070062
    label: extracellular exosome
  evidence_type: IDA
  original_reference_id: PMID:18852879
  review:
    summary: Exosomal BAG6 is experimentally documented, but this is a conditional
      extracellular context rather than the main cellular location.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core localization.
    supported_by:
    - reference_id: PMID:18852879
- term:
    id: GO:0036503
    label: ERAD pathway
  evidence_type: IDA
  original_reference_id: PMID:23129660
  review:
    summary: This paper studies SGTA antagonism of BAG6-mediated triage of mislocalized
      proteins in the cytosol, not canonical ERAD.
    action: MODIFY
    reason: Replace with the narrower protein-quality-control term that better fits
      cytosolic degradation of mislocalized hydrophobic clients.
    proposed_replacement_terms:
    - id: GO:0006515
      label: protein quality control for misfolded or incompletely synthesized proteins
- term:
    id: GO:0036503
    label: ERAD pathway
  evidence_type: IDA
  original_reference_id: PMID:24981174
  review:
    summary: PMID:24981174 defines RNF126-dependent cytosolic quality control of mislocalized
      proteins rather than general ERAD.
    action: MODIFY
    reason: Replace with the more specific protein-quality-control term rather than
      the broader ubiquitin-dependent catabolic umbrella term.
    proposed_replacement_terms:
    - id: GO:0006515
      label: protein quality control for misfolded or incompletely synthesized proteins
- term:
    id: GO:1904294
    label: positive regulation of ERAD pathway
  evidence_type: IMP
  original_reference_id: PMID:24424410
  review:
    summary: BAG6 is part of ERAD-associated machinery; this paper mostly shows that
      USP13 preserves BAG6 complex function rather than BAG6 acting as an upstream
      regulator of ERAD.
    action: MODIFY
    reason: The direct BAG6 claim is better captured as ERAD pathway participation.
    proposed_replacement_terms:
    - id: GO:0036503
      label: ERAD pathway
- term:
    id: GO:0005737
    label: cytoplasm
  evidence_type: IDA
  original_reference_id: PMID:21636303
  review:
    summary: Whole-cell cytoplasmic localization is fully consistent with the BAG6
      complex's proteostasis role.
    action: ACCEPT
    reason: This is a defensible localization term.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:25535373
  review:
    summary: The structural complex paper supports specific adaptor/complex roles,
      not generic protein binding.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0071818
    label: BAT3 complex
  evidence_type: IDA
  original_reference_id: PMID:25535373
  review:
    summary: This paper directly analyzes the BAG6 heterotrimeric complex architecture.
    action: ACCEPT
    reason: BAT3 complex membership is well supported.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:23246001
  review:
    summary: This ERAD paper supports specific SGTA/BAG6 pathway functions rather
      than a useful generic binding term.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IDA
  original_reference_id: PMID:23246001
  review:
    summary: The SGTA/BAG6 ERAD paper supports a cytosolic BAG6 pool.
    action: ACCEPT
    reason: This matches the core QC role.
- term:
    id: GO:0016020
    label: membrane
  evidence_type: IDA
  original_reference_id: PMID:23246001
  review:
    summary: The paper supports regulated ER-membrane association through ERAD machinery
      rather than a generic membrane localization.
    action: MODIFY
    reason: Use endoplasmic reticulum membrane as the more faithful location.
    proposed_replacement_terms:
    - id: GO:0005789
      label: endoplasmic reticulum membrane
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:21636303
  review:
    summary: The 2011 ERAD study notes nuclear sequestration when Trc35 is absent,
      consistent with BAG6's known dynamic localization.
    action: ACCEPT
    reason: Nuclear localization is supported.
- term:
    id: GO:0031625
    label: ubiquitin protein ligase binding
  evidence_type: IPI
  original_reference_id: PMID:21636303
  review:
    summary: Bag6 physically associates with ERAD E3 ligases such as gp78/AMFR and
      SYVN1, which is central to its UPS-adjacent chaperone role.
    action: ACCEPT
    reason: This specific binding term is mechanistically informative and supported.
    supported_by:
    - reference_id: PMID:21636303
- term:
    id: GO:0051787
    label: misfolded protein binding
  evidence_type: IDA
  original_reference_id: PMID:21636303
  review:
    summary: This is the key direct evidence for BAG6 recognizing aggregation-prone
      hydrophobic/misfolded clients.
    action: ACCEPT
    reason: The term accurately captures BAG6 holdase specificity.
    supported_by:
    - reference_id: PMID:21636303
- term:
    id: GO:0071818
    label: BAT3 complex
  evidence_type: IDA
  original_reference_id: PMID:21636303
  review:
    summary: The paper directly studies the BAG6/Ubl4A/Trc35 complex.
    action: ACCEPT
    reason: Complex membership is well supported.
- term:
    id: GO:1990381
    label: ubiquitin-specific protease binding
  evidence_type: IPI
  original_reference_id: PMID:24424410
  review:
    summary: Direct interaction with USP13 is well supported, but this is a specific
      partner interaction rather than the best core BAG6 function summary.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core mechanistic context.
    supported_by:
    - reference_id: PMID:24424410
- term:
    id: GO:0031625
    label: ubiquitin protein ligase binding
  evidence_type: IPI
  original_reference_id: PMID:24981174
  review:
    summary: RNF126 recruitment to the BAG6 UBL domain is a core mechanistic part
      of mislocalized-protein triage.
    action: ACCEPT
    reason: This specific ligase-binding annotation is supported and informative.
    supported_by:
    - reference_id: PMID:24981174
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:18765639
  review:
    summary: The chromatin-regulator interaction paper does not justify retaining
      a generic protein-binding term.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0001822
    label: kidney development
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Mouse-based developmental evidence is plausible but not core to human
      BAG6 proteostasis biology.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0005515
    label: protein binding
  evidence_type: IPI
  original_reference_id: PMID:17403783
  review:
    summary: The p53 acetylation paper supports a specific nuclear scaffold role,
      not a useful generic protein-binding annotation.
    action: REMOVE
    reason: Remove generic protein-binding carryover.
- term:
    id: GO:0005634
    label: nucleus
  evidence_type: IDA
  original_reference_id: PMID:17403783
  review:
    summary: This paper supports nuclear BAG6 during DNA-damage signaling.
    action: ACCEPT
    reason: Nuclear localization is well supported.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IDA
  original_reference_id: PMID:17403783
  review:
    summary: This paper also supports the broader nucleo-cytoplasmic distribution
      of BAG6.
    action: ACCEPT
    reason: Cytosolic BAG6 is consistent with its principal proteostasis role.
- term:
    id: GO:0005829
    label: cytosol
  evidence_type: IDA
  original_reference_id: PMID:20676083
  review:
    summary: The ribosome-associated TA-targeting study places BAG6 complex function
      in the cytosol.
    action: ACCEPT
    reason: Cytosolic localization is central to GET-pathway capture/handoff.
- term:
    id: GO:0006511
    label: ubiquitin-dependent protein catabolic process
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: The broad catabolic-process inference is directionally consistent with
      BAG6 QC biology but loses specificity.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core rather than using it as a core summary term.
- term:
    id: GO:0007130
    label: synaptonemal complex assembly
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Orthology-based reproductive biology is secondary/contextual.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0007283
    label: spermatogenesis
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Orthology-based reproductive biology is secondary/contextual.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0007420
    label: brain development
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Orthology-based developmental biology is secondary/contextual.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0018393
    label: internal peptidyl-lysine acetylation
  evidence_type: IDA
  original_reference_id: PMID:17403783
  review:
    summary: BAG6 scaffolds p300-dependent p53 acetylation after DNA damage, but this
      is a specialized nuclear stress-response role rather than the core proteostasis
      function.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core context.
    supported_by:
    - reference_id: PMID:17403783
- term:
    id: GO:0030324
    label: lung development
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Orthology-based developmental biology is secondary/contextual.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0031593
    label: polyubiquitin modification-dependent protein binding
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: The BAG6 literature reviewed here does not provide a solid basis for
      a direct polyubiquitin-binding activity.
    action: REMOVE
    reason: Remove this inferred binding term.
- term:
    id: GO:0032435
    label: negative regulation of proteasomal ubiquitin-dependent protein catabolic
      process
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: This reflects secondary stabilization biology rather than BAG6's central
      PN role.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0042771
    label: intrinsic apoptotic signaling pathway in response to DNA damage by p53
      class mediator
  evidence_type: IMP
  original_reference_id: PMID:17403783
  review:
    summary: This DNA-damage apoptosis role is well supported but context-specific.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core nuclear stress biology.
    supported_by:
    - reference_id: PMID:17403783
- term:
    id: GO:0043022
    label: ribosome binding
  evidence_type: IDA
  original_reference_id: PMID:20676083
  review:
    summary: Ribosome association is an important mechanistic feature of the GET-pathway
      capture step, but it is not the best standalone summary of BAG6 core function.
    action: KEEP_AS_NON_CORE
    reason: Keep as mechanistically informative but non-core.
- term:
    id: GO:0045861
    label: negative regulation of proteolysis
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Secondary stabilization/proteolysis control is contextual rather than
      core.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0050821
    label: protein stabilization
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Selected-client stabilization is context-specific and not the main BAG6
      role.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0070059
    label: intrinsic apoptotic signaling pathway in response to endoplasmic reticulum
      stress
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: ER-stress apoptosis is context-dependent and not BAG6's principal conserved
      function.
    action: KEEP_AS_NON_CORE
    reason: Keep as non-core.
- term:
    id: GO:0070628
    label: proteasome binding
  evidence_type: ISS
  original_reference_id: GO_REF:0000024
  review:
    summary: Direct proteasome binding is not clearly established by the BAG6 papers
      reviewed here.
    action: REMOVE
    reason: Remove the inferred binding term.
- term:
    id: GO:0071816
    label: tail-anchored membrane protein insertion into ER membrane
  evidence_type: IDA
  original_reference_id: PMID:20676083
  review:
    summary: This is the direct GET-pathway core BAG6 process supported by the landmark
      ribosome-associated TA-targeting paper.
    action: ACCEPT
    reason: The evidence specifically fits tail-anchored membrane protein insertion
      into the ER membrane.
    supported_by:
    - reference_id: PMID:20676083
- term:
    id: GO:0071818
    label: BAT3 complex
  evidence_type: IDA
  original_reference_id: PMID:20676083
  review:
    summary: The 2010 Nature study defines the Bat3/Trc35/Ubl4A complex as the relevant
      functional unit.
    action: ACCEPT
    reason: BAT3 complex membership is directly supported.
core_functions:
- molecular_function:
    id: GO:0140597
    label: protein carrier chaperone
  directly_involved_in:
  - id: GO:0036503
    label: ERAD pathway
  - id: GO:0071816
    label: tail-anchored membrane protein insertion into ER membrane
  locations:
  - id: GO:0005829
    label: cytosol
  in_complex:
    id: GO:0071818
    label: BAT3 complex
  description: BAG6 acts as an ATP-independent holdase/carrier for hydrophobic, misfolded,
    retrotranslocated, or tail-anchored client proteins, keeping them soluble long
    enough for productive ER delivery or proteasome-directed quality control.
  supported_by:
  - reference_id: PMID:21636303
  - reference_id: PMID:20676083
  - reference_id: PMID:25535373
  - reference_id: file:human/BAG6/BAG6-deep-research-falcon.md
    supporting_text: BAG6 captures long hydrophobic TMDs with slow off-rate, recruits RNF126 via its UBL domain for ubiquitination
- molecular_function:
    id: GO:0060090
    label: molecular adaptor activity
  directly_involved_in:
  - id: GO:0071816
    label: tail-anchored membrane protein insertion into ER membrane
  locations:
  - id: GO:0005829
    label: cytosol
  in_complex:
    id: GO:0071818
    label: BAT3 complex
  description: Within the BAG6/UBL4A/GET4 complex, BAG6 provides the scaffold/adaptor
    surface that links SGTA-bound hydrophobic clients to the TRC40/GET pathway and
    organizes the pretargeting complex architecture.
  supported_by:
  - reference_id: PMID:25535373
  - reference_id: PMID:25713138
  - reference_id: file:human/BAG6/BAG6-deep-research-falcon.md
    supporting_text: C-terminal region is a structural part of the substrate-loading complex that bridges UBL4A/SGTA to TRC35/TRC40
- molecular_function:
    id: GO:0051787
    label: misfolded protein binding
  directly_involved_in:
  - id: GO:0036503
    label: ERAD pathway
  - id: GO:0061857
    label: endoplasmic reticulum stress-induced pre-emptive quality control
  locations:
  - id: GO:0005829
    label: cytosol
  in_complex:
    id: GO:0071818
    label: BAT3 complex
  description: BAG6 preferentially recognizes aggregation-prone hydrophobic or misfolded
    clients generated during ERAD, cytosolic mislocalization, or ER stress rerouting,
    linking substrate capture to downstream quality-control decisions.
  supported_by:
  - reference_id: PMID:21636303
  - reference_id: PMID:26565908
  - reference_id: file:human/BAG6/BAG6-deep-research-falcon.md
    supporting_text: BAG6 captures hydrophobic or mislocalized clients, keeps them soluble, recruits ubiquitination machinery
status: COMPLETE