Affinage

PSMG4

Proteasome assembly chaperone 4 · UniProt Q5JS54

Length
123 aa
Mass
13.8 kDa
Annotated
2026-06-10
22 papers in source corpus 12 papers cited in narrative 12 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/6 claims corpus-supported (83%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

PSMG4 (PAC4) is a dedicated proteasome assembly chaperone that operates at the earliest stage of 20S core particle biogenesis as one half of the PAC3-PAC4 heterodimer, the mammalian counterpart of yeast Poc3/Poc4 acting upstream of the half-proteasome maturase UMP1 (PMID:17707236). The heterodimer functions as a molecular matchmaker that nucleates α-ring assembly: it is required to form the initial α4–α7 core intermediate and stabilizes the α4-α5-α6 subcomplex, without which the α-ring cannot assemble (PMID:30133132, PMID:31067643). Atomic-resolution crystallography of human PAC4 defined a hydrophobic surface ringed by charged residues that is complementary to PAC3 and charge-complementary to the proteasomal α4-α5 subunits, providing the structural basis for these interactions (PMID:28263418, PMID:31067643). Cryo-EM of chaperone-bound assembly intermediates places PAC3/PAC4 alongside PAC1-PAC2 on an early α-ring intermediate and shows that timed dissociation of PAC3/PAC4 is the trigger that licenses the transition from α-ring to β-ring assembly (PMID:39294158). Beyond its constitutive role, PAC3-PAC4 also directs selective incorporation of immunoproteasome-specific β subunits during immunoproteasome assembly (PMID:35563886). PSMG4 expression is controlled by SRSF5-dependent alternative splicing, where exon 2 skipping reduces full-length PSMG4 and contributes to pathological cardiac hypertrophy (PMID:41907183).

Mechanistic history

Synthesis pass · year-by-year structured walk · 9 steps
  1. 2007 High

    Established that mammals possess a dedicated early-acting proteasome assembly chaperone, defining PAC4 as a functional homolog working with PAC3 upstream of UMP1.

    Evidence Genetic phenotype screen and functional complementation across yeast and mammalian cells

    PMID:17707236

    Open questions at the time
    • Did not provide structural detail of PAC3-PAC4 interaction
    • Precise α-subunit binding partners not yet defined
  2. 2010 Medium

    Consolidated the model that PAC3-PAC4 acts in α-ring formation prior to β-subunit incorporation and UMP1 function, ordering the assembly pathway.

    Evidence Biochemical analysis of assembly intermediates synthesized in review form

    PMID:18786393 PMID:20074030

    Open questions at the time
    • Reviews rather than new primary data
    • Mechanism of which α-subunits PAC3-PAC4 directly engages unresolved
  3. 2017 High

    Provided the atomic structure of human PAC4, revealing the chemical basis for its complementarity with PAC3 and proteasomal α-subunits.

    Evidence X-ray crystallography at 1.90-Å resolution with surface characterization

    PMID:28263418

    Open questions at the time
    • Structure of the assembled PAC3-PAC4/α-subunit complex not determined
    • Dynamics of binding not addressed
  4. 2018 High

    Demonstrated that PAC3-PAC4 is strictly required to nucleate the α4–α7 core intermediate, establishing its non-redundant role at the start of α-ring assembly.

    Evidence Co-immunoprecipitation, knockdown, subcellular fractionation and overexpression of intermediates

    PMID:30133132

    Open questions at the time
    • Quantitative kinetics of intermediate formation not measured
    • Coordination with PAC1-PAC2 retention mechanism only partly defined
  5. 2019 High

    Defined PAC3-PAC4 as a molecular matchmaker stabilizing the α4-α5-α6 subcomplex and modeled the quintet complex at atomic resolution.

    Evidence 0.96-Å crystallography, NMR, structural modeling and biochemical binding assays

    PMID:31067643

    Open questions at the time
    • Functional consequence of loop 51-61 mobility not directly tested
    • Model not confirmed by a full experimental complex structure
  6. 2022 Medium

    Extended PAC3-PAC4 function to specialized assembly, showing it directs selective incorporation of immunoproteasome β subunits.

    Evidence Biochemical assembly intermediate analysis synthesized in review form

    PMID:35563886

    Open questions at the time
    • Review rather than primary data
    • Molecular basis for β1i/β2i selectivity not structurally defined
  7. 2024 High

    Captured the full chaperone-bound assembly trajectory, showing PAC3/PAC4 dissociation triggers the α-to-β ring transition and couples to proteolytic activation.

    Evidence Cryo-EM of endogenously CRISPR-tagged and recombinant chaperone-bound intermediates

    PMID:38328185 PMID:39294158

    Open questions at the time
    • Signal that times PAC3/PAC4 release not identified
    • Preprint companion study awaiting peer review
  8. 2025 Low

    Linked PSMG4 to cellular drug and damage responses through unbiased screens and expression correlation, though without direct mechanistic validation.

    Evidence Genome-wide CRISPR knockout screen and drug-treatment expression profiling in cancer cells

    PMID:39918307 PMID:40349826

    Open questions at the time
    • Colibactin screen validated PSMD4, not PSMG4
    • Caffeine data are correlative with no PSMG4 manipulation
  9. 2026 Medium

    Revealed regulatory control of PSMG4 abundance via SRSF5-dependent alternative splicing, connecting PSMG4 dosage to cardiac hypertrophy.

    Evidence RNA interaction assays, splicing reporters, AngII mouse and hiPSC-CM models with CHAtRF manipulation

    PMID:41907183

    Open questions at the time
    • Whether reduced PSMG4 impairs proteasome assembly in cardiomyocytes not shown
    • Single-lab finding awaiting independent replication

Open questions

Synthesis pass · forward-looking unresolved questions
  • The molecular trigger that times PAC3/PAC4 dissociation and how PSMG4 dosage perturbations propagate to proteasome capacity in disease remain unresolved.
  • No defined signal driving PAC3/PAC4 release
  • Causal link between PSMG4 splicing changes and proteasome dysfunction in hypertrophy not established

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 3 GO:0140096 catalytic activity, acting on a protein 3
Localization
GO:0005829 cytosol 1
Pathway
R-HSA-392499 Metabolism of proteins 3 R-HSA-1852241 Organelle biogenesis and maintenance 2
Partners
Complex memberships
20S proteasome assembly intermediatePAC3-PAC4 heterodimer

Evidence

Reading pass · 12 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2007 PAC4 (PSMG4) was identified as a mammalian proteasome assembly chaperone functioning as part of the PAC3-PAC4 heterodimer pair, acting at early stages of 20S proteasome assembly (alpha-ring formation), analogous to yeast Poc3/Poc4. It works upstream of the half-proteasome maturase UMP1. Genetic phenotype screen, yeast and mammalian cell-based functional complementation, identification of functional homologs Molecular cell High 17707236
2008 PAC3-PAC4 heterodimer promotes assembly of rings composed of seven alpha subunits of the 20S proteasome core particle, acting prior to beta-subunit incorporation and prior to UMP1-dependent half-proteasome dimerization. Biochemical fractionation, assembly intermediate analysis, review/synthesis of experimental data Structure (London, England : 1993) Medium 18786393
2010 PAC3-PAC4 heterodimer is implicated in alpha-ring formation during 20S proteasome core particle assembly; PAC3-PAC4 acts upstream of beta-subunit incorporation and UMP1 function. Biochemical analysis of assembly intermediates, review of experimental data Biochemical Society transactions Medium 20074030
2017 Crystal structure of human PAC4 (PSMG4) determined at 1.90-Å resolution, revealing a hydrophobic surface surrounded by charged residues that is complementary to its binding partner PAC3 and exhibits charge complementarity with proteasomal α4-5 subunits. X-ray crystallography at 1.90-Å resolution Protein science : a publication of the Protein Society High 28263418
2018 PAC3-PAC4 is required for formation of the core α4-α7 intermediate as the initial step of α-ring assembly. Without PAC3-PAC4, the α4-α7 core intermediate does not form. PAC1-PAC2 then prevents nonproductive dimerization of α-ring assembly intermediates and retains them in the cytoplasm by overriding nuclear localization signals of α-subunits. Co-immunoprecipitation, knockdown experiments, subcellular fractionation, overexpression of assembly intermediates Genes to cells : devoted to molecular & cellular mechanisms High 30133132
2019 The PAC3-PAC4 heterodimer functions as a molecular matchmaker, stabilizing the α4-α5-α6 subcomplex during α-ring assembly. A 0.96-Å crystal structure of PAC3 homodimer and NMR data revealed loop mobility (residues 51-61) and enabled modeling of the PAC3-4/α4/α5/α6 quintet complex. X-ray crystallography (0.96-Å resolution), NMR spectroscopy, 3D structural modeling, biochemical binding assays International journal of molecular sciences High 31067643
2022 PAC3-PAC4 is required for selective incorporation of immunoproteasome-specific beta subunits (β1i, β2i) before other beta subunits during immunoproteasome assembly, providing selectivity against constitutive subunits. Biochemical assembly intermediate analysis, review of experimental data Cells Medium 35563886
2024 Cryo-EM structures of chaperone-bound human 20S proteasome assembly intermediates show that PAC3/PAC4 stabilizes an early α-ring intermediate subcomplex together with PAC1-PAC2. Dissociation of PAC3/PAC4 (along with rearrangement of PAC1 N-terminal tail) triggers the transition to β-ring assembly. Completion of β-ring and half-proteasome dimerization repositions lysine K33 to trigger β pro-peptide cleavage, leading to concerted dissociation of POMP and PAC1/PAC2. Cryo-EM of endogenously CRISPR-tagged chaperone-bound complexes, structural analysis of assembly intermediates Nature communications High 39294158
2024 Cryo-EM reconstructions of seven recombinant human proteasome subcomplexes visualize PAC4 as part of the chaperone ensemble stabilizing early assembly intermediates; PAC3/PAC4 dissociation is required for progression from α-ring to β-ring assembly. Structural data explain the order of successive subunit additions and how assembly factors rearrange to coordinate proteolytic activation. Cryo-EM of recombinant human subcomplexes, structural comparison of intermediates and mature CP bioRxiv : the preprint server for biologypreprint Medium 38328185
2025 CRISPR/Cas9 genome-wide knockout screen identified PSMG4 as a gene whose loss confers resistance to colibactin-induced cytotoxicity, suggesting PSMG4 participates in the cellular response pathway to colibactin-mediated DNA damage. (Note: mechanistic validation was performed for PSMD4, not PSMG4, in this study.) Genome-wide CRISPR/Cas9 knockout screen with MAGeCK scoring mSphere Low 39918307
2025 Caffeine treatment of colorectal adenocarcinoma cells decreased expression of PAC4 (PSMG4) coincident with reduced immunoproteasome content and reduced oxidative stress, suggesting PAC4 expression is linked to immunoproteasome biogenesis in cancer cells. qPCR, Western blot, transcriptome analysis, flow cytometry of proteasome subunit expression Biochimie Low 40349826
2026 Alternative splicing of Psmg4 pre-mRNA regulated by the SRSF5 splicing factor promotes exon 2 skipping, reducing full-length Psmg4 isoform expression. A tRNA-derived fragment (CHAtRF) directly interacts with SRSF5, blocking its binding to Psmg4 pre-mRNA and thereby promoting exon 2 skipping, which inhibits full-length PSMG4 expression and contributes to pathological cardiac hypertrophy. RNA pulldown/interaction assays, alternative splicing reporter assays, cardiac hypertrophy models (AngII-treated mice, hiPSC-CMs), CHAtRF overexpression/knockdown Research (Washington, D.C.) Medium 41907183

Source papers

Stage 0 corpus · 22 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2007 20S proteasome assembly is orchestrated by two distinct pairs of chaperones in yeast and in mammals. Molecular cell 165 17707236
2011 Identification of genes that elicit disuse muscle atrophy via the transcription factors p50 and Bcl-3. PloS one 68 21249144
2008 PACemakers of proteasome core particle assembly. Structure (London, England : 1993) 51 18786393
1987 A complex gene superfamily encodes actin in petunia. The EMBO journal 47 3428258
1983 Platelet associated immunoglobulins and complement in idiopathic thrombocytopenic purpura. Clinical and experimental immunology 30 6683604
2018 PAC1-PAC2 proteasome assembly chaperone retains the core α4-α7 assembly intermediates in the cytoplasm. Genes to cells : devoted to molecular & cellular mechanisms 27 30133132
2022 The Molecular Mechanisms Governing the Assembly of the Immuno- and Thymoproteasomes in the Presence of Constitutive Proteasomes. Cells 25 35563886
2022 Sperm DNA methylation patterns at discrete CpGs and genes involved in embryonic development are related to bull fertility. BMC genomics 23 35585482
2019 Molecular and Structural Basis of the Proteasome α Subunit Assembly Mechanism Mediated by the Proteasome-Assembling Chaperone PAC3-PAC4 Heterodimer. International journal of molecular sciences 23 31067643
2010 Chaperone-assisted assembly of the proteasome core particle. Biochemical Society transactions 23 20074030
1990 Six actin gene subfamilies map to five chromosomes of Petunia hybrida. The Journal of heredity 16 1977797
2024 Structural basis of human 20S proteasome biogenesis. Nature communications 13 39294158
2017 Crystal structure of human proteasome assembly chaperone PAC4 involved in proteasome formation. Protein science : a publication of the Protein Society 11 28263418
2020 Design of potent ABA receptor antagonists based on a conformational restriction approach. Organic & biomolecular chemistry 7 32568332
2025 Genome-scale CRISPR/Cas9 screening reveals the role of PSMD4 in colibactin-mediated cell cycle arrest. mSphere 6 39918307
2025 A genome-wide association study identified candidate genes associated with egg quality traits in Muscovy duck. BMC genomics 6 40301754
2023 Antimicrobial Activity and Cytotoxicity of Prepolymer Allyl 2-cyanoacrylate and 2-Octyl Cyanoacrylate Mixture Adhesives for Topical Wound Closure. Materials (Basel, Switzerland) 3 37176306
2025 Caffeine modulates immunoproteasome activity and content in colorectal adenocarcinoma cells. Biochimie 1 40349826
2024 Visualizing chaperone-mediated multistep assembly of the human 20S proteasome. bioRxiv : the preprint server for biology 1 38328185
2024 Structural basis of human 20S proteasome biogenesis. bioRxiv : the preprint server for biology 1 39211201
2026 CHAtRF Modulates Cardiac Hypertrophy via SRSF5-Dependent Regulation of Psmg4 Alternative Splicing. Research (Washington, D.C.) 0 41907183
2025 Molecular and functional analysis of a putative pyocin S9, with endonuclease activity from P. chlororaphis subsp aurantiaca PB-St2. Archives of microbiology 0 40338343

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