Affinage

EIF3D

Eukaryotic translation initiation factor 3 subunit D · UniProt O15371

Length
548 aa
Mass
64.0 kDa
Annotated
2026-06-09
38 papers in source corpus 19 papers cited in narrative 19 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 5/5 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

EIF3D is the subunit of the eIF3 translation initiation complex that carries an intrinsic mRNA 5' cap-binding domain, defining a non-canonical, eIF4E-independent but cap-dependent pathway of translation initiation (PMID:27462815, PMID:39107322). Its cap-binding domain is structurally homologous to RNA endonucleases and makes specific contacts with the m7G cap that are required to assemble initiation complexes on specialized mRNAs such as c-Jun, whose 5' inhibitory element blocks eIF4E recruitment (PMID:27462815); recruitment is quantitatively tuned by full cap methylation and operates on a defined subset of capped mRNAs that release eIF4E and engage eIF3d when eIF4E activity is suppressed (PMID:39107322, PMID:39971159). This pathway is switched on during cellular stress: glucose deprivation lowers CK2-mediated phosphorylation near the cap-binding pocket to activate eIF3d and drive selective translation of glucose-homeostasis and mTOR-pathway mRNAs essential for survival (PMID:33184215), and during persistent integrated stress response eIF3d activates GCN2 translation and upregulates the m6A demethylase ALKBH5 to enhance ATF4 translation via 5' UTR demethylation and uORF bypass (PMID:37683648). eIF3d additionally partners with DAP5/eIF4G2 to direct context-specific translational reprogramming of Treg-differentiation, EMT, migration and angiogenesis mRNAs (PMID:34848685, PMID:37314929, PMID:39971159), and the program is engaged downstream of mTOR inhibition, hypoxia (with eIF3e, controlling HIF1α), and viral infection (PMID:37494188, PMID:41364558, PMID:35508137). As a structural element of the complex, eIF3d is required to maintain eIF3 subunit integrity (PMID:11705997), and beyond translation it stabilizes specific client proteins (GRK2, GRP78) against proteasomal degradation (PMID:28594409, PMID:31669222) and is itself K27-polyubiquitinated at K153/K275 by the CUL3/KCTD10 ligase (PMID:31280863). Through these activities eIF3d governs selective translational responses controlling fibrogenesis (PMID:41197183) and pluripotency maintenance (PMID:40203091).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2001 High

    Established that eIF3d is a physical and functional core component of the eIF3 complex, answering whether it is integral to initiation machinery rather than peripheral.

    Evidence Co-IP, sucrose gradient sedimentation, and moe1 deletion in fission yeast

    PMID:11705997

    Open questions at the time
    • Did not reveal the cap-binding function discovered later
    • Ortholog-based, mammalian role not directly tested
  2. 2016 High

    Discovered that eIF3d harbors an intrinsic 5' cap-binding domain, defining an eIF4E-independent yet cap-dependent initiation pathway and explaining mRNA-selective initiation on transcripts like c-Jun.

    Evidence 1.4 Å crystal structure, cap analogue competition, mutagenesis, in vitro initiation complex assembly

    PMID:27462815

    Open questions at the time
    • Did not establish how the pathway is regulated in cells
    • Scope of the eIF3d-dependent mRNA program not defined
  3. 2018 Medium

    Demonstrated subunit-specific, mRNA-selective translational control by eIF3d via direct 5' UTR binding and cofactor (Hrp48) interactions, distinguishing its role from other eIF3 subunits.

    Evidence RNA chromatography, reporter assays, RNAi depletion and Co-IP in Drosophila

    PMID:29635389

    Open questions at the time
    • Drosophila ortholog, mammalian generality unclear
    • Direct cap-binding contribution to msl-2 control not dissected
  4. 2017 Medium

    Revealed a non-translational function: eIF3d stabilizes client proteins against ubiquitin-mediated degradation, here GRK2 to activate PI3K/Akt in cancer cells.

    Evidence Co-IP, ubiquitination assay, knockdown/overexpression, proliferation/migration assays

    PMID:28594409

    Open questions at the time
    • Mechanism of degradation blockade unclear
    • Direct vs indirect stabilization not resolved
  5. 2019 Medium

    Extended the protein-stabilizing role to GRP78 and identified eIF3d as a substrate of CUL3/KCTD10, defining how eIF3d is itself post-translationally regulated.

    Evidence Co-IP, ubiquitination assays, mass spectrometry of K153/K275 sites in cancer cells

    PMID:31280863 PMID:31669222

    Open questions at the time
    • Functional consequence of K27 ubiquitination on eIF3d activity not established
    • Reciprocal regulation between translational and stabilizing roles unknown
  6. 2020 High

    Identified the regulatory switch: CK2 phosphorylation near the cap-binding pocket inhibits eIF3d, and metabolic stress relieves this to activate selective translation, linking the pathway to nutrient sensing and survival.

    Evidence Phosphosite mapping, CK2 inhibition/knockdown, translation profiling, viability assays under glucose deprivation

    PMID:33184215

    Open questions at the time
    • Upstream signaling controlling CK2 in stress not defined
    • Full breadth of the regulated mRNA program incomplete
  7. 2021 Medium

    Showed eIF3d acts with DAP5/eIF4G2 in a distinct complex to support stress/context-specific translation, here Treg differentiation when eIF4E translation is inhibited.

    Evidence Translatome profiling, siRNA knockdown, T cell differentiation and polysome assays

    PMID:34848685

    Open questions at the time
    • Direct eIF3d cap-binding contribution within the DAP5 complex not isolated
    • How target mRNAs are selected unclear
  8. 2022 Medium

    Established eIF3d-dependent translation as a host pathway hijacked during HCMV infection as eIF4E activity wanes, identifying it as an antiviral target.

    Evidence eIF3d knockdown, polyribosome profiling, viral replication and gene expression assays

    PMID:35508137

    Open questions at the time
    • Viral mRNAs directly bound by eIF3d not mapped
    • Trigger of the eIF4E-to-eIF3d shift not defined
  9. 2023 High

    Defined eIF3d as a master coordinator of the integrated stress response by activating GCN2 and ALKBH5-driven m6A demethylation to enhance ATF4 translation, and showed DAP5/eIF3d drives EMT/metastasis programs.

    Evidence Ribosome profiling, m6A-seq, uORF reporters; translatome profiling and metastasis models

    PMID:37314929 PMID:37494188 PMID:37683648

    Open questions at the time
    • How eIF3d selects ISR and EMT target mRNAs not fully resolved
    • Interplay with RBP cofactors (hnRNPF/K/SSB) mechanistically incomplete
  10. 2024 High

    Genome-wide demonstration that eIF4E-independent translation of a defined subset of capped mRNAs is largely dependent on eIF3d cap-binding, mechanistically validating the cap-handoff model.

    Evidence Ribosome profiling under active 4E-BP, cap-binding pocket mutant, mRNA-eIF3d binding assays

    PMID:39107322

    Open questions at the time
    • Determinants of which mRNAs handoff to eIF3d not fully defined
    • In vivo physiological deployment of the handoff incomplete
  11. 2025 Medium

    Quantified cap recruitment of eIF3d/eIF4G2 dependent on full cap methylation and extended eIF3d roles to hypoxia (HIF1α), fibrogenesis (ATF4-S100P axis), pluripotency maintenance, and stress granule assembly via its RNA-binding domain.

    Evidence Fluorescence anisotropy and in vitro translation; ribosome profiling, CRISPRi, fibrosis mouse models, stress granule imaging and domain rescue (one preprint)

    PMID:39971159 PMID:40203091 PMID:41197183 PMID:41364558 PMID:bio_10.1101_2025.11.13.688230

    Open questions at the time
    • Stress granule role is from a preprint awaiting peer review
    • Structural basis of RNA-binding-domain-mediated granule assembly not defined

Open questions

Synthesis pass · forward-looking unresolved questions
  • How the eIF3d cap-binding switch is integrated across competing demands (translation initiation, protein stabilization, stress granule assembly) and how target mRNA selectivity is encoded remain unresolved.
  • No unified model linking phosphorylation, ubiquitination, and cofactor binding to target choice
  • Structural basis for mRNA selectivity beyond the cap unknown
  • Relative in vivo importance of translational vs non-translational functions unquantified

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0003723 RNA binding 5 GO:0045182 translation regulator activity 4 GO:0140313 molecular sequestering activity 2
Localization
GO:0005829 cytosol 2 GO:0005840 ribosome 1
Pathway
R-HSA-392499 Metabolism of proteins 4 R-HSA-8953854 Metabolism of RNA 3 R-HSA-8953897 Cellular responses to stimuli 2
Complex memberships
DAP5/eIF4G2 (eIF3d) complexeIF3 complex

Evidence

Reading pass · 19 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2016 eIF3d possesses a previously unknown cap-binding activity within the 800-kDa eIF3 complex. A 1.4 Å crystal structure of the eIF3d cap-binding domain reveals unexpected homology to endonucleases involved in RNA turnover. eIF3d makes specific contacts with the mRNA 5' cap (validated by cap analogue competition), and these interactions are essential for assembly of translation initiation complexes on eIF3-specialized mRNAs such as c-Jun. The c-Jun mRNA encodes an inhibitory RNA element that blocks eIF4E recruitment, enforcing alternative cap recognition by eIF3d, defining an eIF4E-independent, cap-dependent translation initiation pathway. X-ray crystallography (1.4 Å resolution), cap analogue competition assay, in vitro translation initiation complex assembly, mutagenesis Nature High 27462815
2020 eIF3d cap-binding activity is activated during metabolic stress (glucose deprivation) by reduced CK2-mediated phosphorylation near the eIF3d cap-binding pocket. This phosphorylation switch enables eIF3d to drive selective translation of a gene program enriched in glucose homeostasis factors including mTOR pathway members, and is essential for cell survival during chronic glucose deprivation. Phosphorylation site mapping, CK2 inhibitor and knockdown experiments, ribosome profiling/translation profiling, cell viability assays under glucose deprivation Science High 33184215
2023 During persistent integrated stress response (ISR), eIF3d activates translation of the kinase GCN2, which induces eIF2α phosphorylation and inhibits global protein synthesis. In parallel, eIF3d upregulates the m6A demethylase ALKBH5 to drive 5' UTR-specific demethylation of stress response genes including ATF4, increasing ribosome engagement and enhancing bypass of upstream open reading frames (uORFs) on ATF4 mRNA. Ribosome profiling, m6A sequencing, genetic knockdown/overexpression, uORF reporter assays, eIF2α phosphorylation assays Molecular cell High 37683648
2001 In fission yeast, Moe1 (homologue of mammalian eIF3d/p66) physically associates with eIF3 core subunits and 40S ribosomal particles as part of an eIF3 complex. Deletion of moe1 reduces translation rate by 30–40% and causes loss of stable association between eIF3 subunits upon ribosome dissociation, demonstrating that eIF3d is required for maintaining eIF3 subunit complex integrity. Co-immunoprecipitation, sucrose gradient sedimentation, deletion mutant analysis, translation rate measurement The Journal of biological chemistry High 11705997
2018 In Drosophila, eIF3d binds to the msl-2 5' UTR and is required for efficient translation of msl-2 mRNA. eIF3d also mediates translational repression of msl-2 by interacting with the co-factor Hrp48 (which binds the msl-2 3' UTR and is recruited by Sex-lethal). Depletion of eIF3d — but not of other eIF3 subunits — specifically de-represses msl-2 expression in female flies, indicating a subunit-specific role in mRNA-selective translation control. RNA chromatography, reporter assays, RNAi-mediated depletion in flies, co-immunoprecipitation Nucleic acids research Medium 29635389
2021 In human regulatory T cells (Tregs), a non-canonical cap-dependent translation mechanism utilizes DAP5 (eIF4G2) together with eIF3d, directed by 5' noncoding regions of Treg-specific mRNAs, to support translation of Treg differentiation and immune suppression mRNAs when mTORC1/eIF4E-dependent translation is inhibited. Silencing DAP5 impairs naive CD4+ T cell differentiation into Treg cells. Genome-wide transcriptomic and translatomic profiling, siRNA knockdown, T cell differentiation assays, polysome profiling Nature communications Medium 34848685
2017 EIF3D stabilizes GRK2 protein by blocking ubiquitin-mediated proteasomal degradation of GRK2, thereby activating PI3K/Akt signaling and promoting gallbladder cancer cell proliferation and migration. This represents a non-translational function of eIF3d. Co-immunoprecipitation, ubiquitination assay, knockdown/overexpression, in vitro and in vivo proliferation/migration assays Cell death & disease Medium 28594409
2019 EIF3D interacts with GRP78 and enhances GRP78 protein stability by blocking ubiquitin-mediated proteasomal degradation of GRP78, thereby promoting sunitinib resistance in renal cell carcinoma cells via unfolded protein response activation. Co-immunoprecipitation, Western blot, ubiquitination assay, knockdown/overexpression, in vitro and in vivo growth assays EBioMedicine Medium 31669222
2019 EIF3D is K27-polyubiquitinated at lysine residues K153 and K275 by the Cullin-3/KCTD10 ubiquitin ligase complex in human hepatocellular carcinoma HepG2 cells, as identified by mass spectrometry. Co-immunoprecipitation, mass spectrometry, site-directed mutagenesis of ubiquitination sites, ubiquitination assay Biochemical and biophysical research communications Medium 31280863
2022 During human cytomegalovirus (HCMV) infection, protein synthesis progressively shifts from eIF4E-dependent to eIF3d-dependent cap-dependent translation. Targeting eIF3d selectively inhibits HCMV replication, reduces polyribosome abundance, and interferes with expression of essential virus genes and a host chronic ER stress gene signature that supports HCMV reproduction. eIF3d knockdown/targeting, polyribosome profiling, viral replication assays, gene expression analysis Cell reports Medium 35508137
2023 The DAP5/eIF3d complex mediates selective cap-dependent, eIF4E-independent translation of mRNAs encoding EMT transcription factors, cell migration integrins, metalloproteinases, and angiogenesis factors in breast cancer cells. DAP5 is required for EMT, cell migration, invasion, metastasis, and angiogenesis in human and murine breast cancer models, but not for primary tumor growth. Genome-wide transcriptomic and translatomic profiling, siRNA knockdown, animal models of metastasis, cell migration/invasion assays Cell reports Medium 37314929
2023 mTOR inhibition activates eIF3d-mediated non-canonical translation, which cooperates with mRNA-binding proteins hnRNPF, hnRNPK, and SSB to support selective translation of mRNAs in INSR/IGF1R/IRS and IL-6ST/JAK/STAT signaling pathways, enabling cell phenotype switching from proliferative to migratory. Ribosome profiling, quantitative proteomics, eIF3d knockdown, mTOR inhibitor treatment, co-immunoprecipitation with hnRNPF/K/SSB Cell reports Medium 37494188
2024 eIF4E-independent translation of a subset of capped mRNAs is largely dependent on eIF3d cap-binding activity. Under eIF4E1 inactivation, these mRNAs preferentially release eIF4E1 and bind instead to eIF3d via its cap-binding pocket, enabling efficient translation. Ribosome profiling under constitutively active 4E-BP expression, eIF3d cap-binding pocket mutant, mRNA-eIF3d binding assays Nature communications High 39107322
2025 eIF3d and eIF3e mediate a selective translational response to acute hypoxia that controls HIF1α accumulation and cellular invasion. This translation program is dependent on the eIF3d/eIF3e module and can be inhibited by novel small molecules targeting eIF3e. Ribosome profiling in hypoxia, eIF3d/eIF3e knockdown, cellular invasion assays, HIF1α measurement, small molecule inhibitor characterization Cell reports Medium 41364558
2025 eIF3d quantitatively recruits itself and the eIF3d/eIF4G2 (DAP5) complex to specific capped mRNAs via its cap-binding activity, with binding affinity dependent on a fully methylated 5' mRNA cap. This eIF3d/eIF4G2 recruitment correlates with translation efficiency of these mRNAs in cap-dependent, eIF4E-independent manner as measured by in vitro translation assays. Fluorescence anisotropy equilibrium binding assays, in vitro luciferase reporter translation assays, cap analogue competition The Journal of biological chemistry Medium 39971159
2025 EIF3D-mediated translation of ATF4 drives ATF4-dependent S100P transcription in hepatic stellate cells (HSCs), activating JNK and NLRP3 signaling to promote HSC activation, survival, proliferation, and extracellular matrix production. Genetic and pharmacological inhibition of the EIF3D-ATF4-S100P axis suppresses metabolic reprogramming (mitochondrial activity and glycolysis) and fibrogenic markers in HSCs. Genetic knockdown/overexpression, HSC-specific ATF4 deletion mouse models, pharmacological ISR inhibitor (ERMT1), metabolic assays, fibrosis mouse models Redox biology Medium 41197183
2025 EIF3D is required for maintaining primed pluripotency by controlling translation of p53 regulators (keeping p53 activity low) and balancing pluripotency-associated signaling pathways. Loss of EIF3D disrupts this translational homeostasis, compromising the undifferentiated state. CRISPR interference screen, EIF3D knockdown in human PSCs, ribosome profiling, pluripotency marker analysis Science advances Medium 40203091
2025 The RNA-binding domain of eIF3d mediates its recruitment to cytoplasmic stress granules and is required for stress granule assembly in response to specific stresses. Deletion of this domain blocks granule formation, decreases cell viability, and the exogenous RNA-binding domain alone rescues stress granule assembly in eIF3d-depleted cells. This function is conserved in C. elegans. Live-cell imaging of stress granules, eIF3d domain deletion mutants, eIF3d depletion with rescue, C. elegans in vivo experiments bioRxivpreprint Medium bio_10.1101_2025.11.13.688230
2024 Src oncogene controls eIF3d-dependent non-canonical cap-dependent translation initiation pathway in addition to the canonical mTOR/eIF4E pathway. eIF3d (together with eIF3h and eIF3e) is essential for invadosome formation and extracellular matrix degradation downstream of Src. Both eIF4E and eIF3d pathways are required for invadosome function. eIF3d/eIF3h/eIF3e knockdown, invadosome formation assays, ECM degradation assays, Src inhibitor experiments, expression correlation analysis bioRxivpreprint Low bio_10.1101_2024.08.01.606119

Source papers

Stage 0 corpus · 38 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2016 eIF3d is an mRNA cap-binding protein that is required for specialized translation initiation. Nature 292 27462815
2020 A phosphorylation-regulated eIF3d translation switch mediates cellular adaptation to metabolic stress. Science (New York, N.Y.) 99 33184215
2017 EIF3D promotes gallbladder cancer development by stabilizing GRK2 kinase and activating PI3K-AKT signaling pathway. Cell death & disease 55 28594409
2023 eIF3d controls the persistent integrated stress response. Molecular cell 45 37683648
2021 A DAP5/eIF3d alternate mRNA translation mechanism promotes differentiation and immune suppression by human regulatory T cells. Nature communications 44 34848685
2019 EIF3D promotes sunitinib resistance of renal cell carcinoma by interacting with GRP78 and inhibiting its degradation. EBioMedicine 38 31669222
2016 EIF3D silencing suppresses renal cell carcinoma tumorigenesis via inducing G2/M arrest through downregulation of Cyclin B1/CDK1 signaling. International journal of oncology 36 27035563
2001 Moe1 and spInt6, the fission yeast homologues of mammalian translation initiation factor 3 subunits p66 (eIF3d) and p48 (eIF3e), respectively, are required for stable association of eIF3 subunits. The Journal of biological chemistry 36 11705997
2023 Breast cancer cell mesenchymal transition and metastasis directed by DAP5/eIF3d-mediated selective mRNA translation. Cell reports 35 37314929
2015 Knockdown of eIF3D inhibits breast cancer cell proliferation and invasion through suppressing the Wnt/β-catenin signaling pathway. International journal of clinical and experimental pathology 32 26617750
2017 Proteomics analysis of bladder cancer invasion: Targeting EIF3D for therapeutic intervention. Oncotarget 30 29050215
2015 The oncogenic role of EIF3D is associated with increased cell cycle progression and motility in prostate cancer. Medical oncology (Northwood, London, England) 24 26036682
2023 mTOR inhibition reprograms cellular proteostasis by regulating eIF3D-mediated selective mRNA translation and promotes cell phenotype switching. Cell reports 23 37494188
2015 Knockdown of EIF3D suppresses proliferation of human melanoma cells through G2/M phase arrest. Biotechnology and applied biochemistry 23 25322666
2024 eIF4E-independent translation is largely eIF3d-dependent. Nature communications 22 39107322
2015 RNAi-Mediated Silencing of EIF3D Alleviates Proliferation and Migration of Glioma U251 and U87MG Cells. Chemical biology & drug design 21 25682860
2015 Knockdown of eIF3d inhibits cell proliferation through G2/M phase arrest in non-small cell lung cancer. Medical oncology (Northwood, London, England) 21 26008152
2023 eIF3d: A driver of noncanonical cap-dependent translation of specific mRNAs and a trigger of biological/pathological processes. The Journal of biological chemistry 19 36997088
2018 Hrp48 and eIF3d contribute to msl-2 mRNA translational repression. Nucleic acids research 19 29635389
2022 An eIF3d-dependent switch regulates HCMV replication by remodeling the infected cell translation landscape to mimic chronic ER stress. Cell reports 17 35508137
2019 Cullin-3/KCTD10 complex is essential for K27-polyubiquitination of EIF3D in human hepatocellular carcinoma HepG2 cells. Biochemical and biophysical research communications 14 31280863
2019 Reduced eIF3d accelerates HIV disease progression by attenuating CD8+ T cell function. Journal of translational medicine 9 31118081
2023 EIF3D promotes resistance to 5-fluorouracil in colorectal cancer through upregulating RUVBL1. Journal of clinical laboratory analysis 8 36592991
2021 A Versatile eIF3d in Translational Control of Stress Adaptation. Molecular cell 7 33417853
2017 Knockdown of eukaryotic translation initiation factor 3 subunit D (eIF3D) inhibits proliferation of acute myeloid leukemia cells. Molecular and cellular biochemistry 7 28801778
2018 hmiR-34c-3p upregulation inhibits the proliferation of colon cancer cells by targeting EIF3D. Anti-cancer drugs 6 30096129
2025 eIF3d and eIF4G2 mediate an alternative mechanism of cap-dependent but eIF4E-independent translation initiation. The Journal of biological chemistry 5 39971159
2025 Integrated stress response-mediated metabolic reprogramming drives hepatic stellate cell activation and liver fibrosis via the noncanonical EIF3d-ATF4-S100P signaling pathway. Redox biology 4 41197183
2022 EIF3D promoted cervical carcinoma through Warburg effect by interacting with GRP78. Journal of obstetrics and gynaecology : the journal of the Institute of Obstetrics and Gynaecology 4 36264610
2021 EIF3D promotes the progression of preeclampsia by inhibiting of MAPK/ERK1/2 pathway. Reproductive toxicology (Elmsford, N.Y.) 4 34520790
2025 EIF3D safeguards the homeostasis of key signaling pathways in human primed pluripotency. Science advances 3 40203091
2024 RNA splicing regulator EIF3D regulates the tumor microenvironment through immunogene-related alternative splicing in head and neck squamous cell carcinoma. Aging 3 38535990
2024 Deciphering EIF3D's Role in Immune Regulation and Malignant Progression: A Pan-Cancer Analysis with a Focus on Colon Adenocarcinoma. Journal of inflammation research 3 39372593
2025 Surface Engineering of MXene and Functional Fullerenols for Cancer Biomarker 'eIF3d'. Langmuir : the ACS journal of surfaces and colloids 2 40114326
2025 eIF3d and eIF3e mediate selective translational control of hypoxia that can be inhibited by small molecules. Cell reports 2 41364558
2026 Quercetagitrin targets EIF3D to activate NCOA4-mediated ferritinophagy-dependent ferroptosis for the treatment of non-small cell lung cancer. Phytomedicine : international journal of phytotherapy and phytopharmacology 1 41653617
2019 Correction to: Expression of Concern: Knockdown of eIF3d inhibits cell proliferation through G2/M phase arrest in non-small cell lung cancer. Medical oncology (Northwood, London, England) 1 31053950
2025 eIF3d and eIF3e mediate selective translational control of hypoxia that can be inhibited by novel small molecules. bioRxiv : the preprint server for biology 0 40501575

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