Affinage

ATF4

Cyclic AMP-dependent transcription factor ATF-4 · UniProt P18848

Length
351 aa
Mass
38.6 kDa
Annotated
2026-04-28
100 papers in source corpus 46 papers cited in narrative 46 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

ATF4 is a bZIP transcription factor that serves as a central effector of the integrated stress response and a metabolic rheostat coupling nutrient and redox sensing to gene expression programs governing amino acid homeostasis, glutathione metabolism, autophagy, and cell fate. Translationally upregulated downstream of eIF2α kinases (PERK, GCN2, HRI) and also via an mTORC1-dependent, eIF2α-independent route, ATF4 directly activates target genes including SLC7A11/xCT (cystine uptake and ferroptosis suppression), REDD1 and Sestrin2 (mTORC1 feedback inhibition), LAMP3, HSPA5, HO-1, and NADPH-generating metabolic enzymes, with its transcriptional output shaped by heterodimerization partners (DISC1, FoxO1, Hop2, KDM4C, CTCF) and cooperative signaling with NRF2 and YAP/TAZ (PMID:16861899, PMID:33646118, PMID:36996941, PMID:31875479, PMID:26011642, PMID:37410595, PMID:34664408). ATF4 protein stability and activity are tuned by PRMT1-mediated Arg239 methylation, PARP-1-mediated PARylation, O-GlcNAcylation, PKA-dependent Ser219 phosphorylation linked to ubiquitin–proteasome degradation, and post-transcriptional control of its mRNA by DDX3, PRMT5-dependent splicing, and the lncRNA TINCR (PMID:31787756, PMID:28457938, PMID:33198401, PMID:29062139, PMID:35305370, PMID:33586907). Beyond transcription, ATF4 performs non-transcriptional roles: it is locally translated in axons as a retrograde injury signal and mediates TRPM3 membrane trafficking via KIF17 in sensory neurons to regulate heat nociception (PMID:25171414, PMID:33658516).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2006 High

    Establishing ATF4 as an effector of the PERK–eIF2α arm of the UPR resolved how ER and hypoxic stress converge on a single transcription factor to support tumor cell survival.

    Evidence PERK-deficient and eIF2α-mutant MEFs combined with xenograft tumor models

    PMID:16861899

    Open questions at the time
    • Identity of the full ATF4 transcriptional program under ER stress was not yet defined
    • Whether other eIF2α kinases converge on ATF4 in these contexts was not tested
  2. 2008 High

    Identification of REDD1 as a direct ATF4 transcriptional target explained how ER stress feeds back to suppress mTORC1, linking ATF4 to growth-factor signaling.

    Evidence ATF4-null and PERK-null MEFs with exogenous ATF4 rescue and mRNA analysis

    PMID:19114033

    Open questions at the time
    • Whether ATF4 binds the REDD1 promoter directly was not shown by ChIP at this stage
    • Functional consequence of REDD1 induction on mTORC1 was inferred, not directly measured
  3. 2012 Medium

    Discovery that FoxO1 physically interacts with ATF4 in osteoblasts and cooperates to regulate osteocalcin/glucose homeostasis revealed that ATF4 functions through heteromeric partnerships to control bone-derived endocrine signaling.

    Evidence Co-IP, colocalization, and compound FoxO1/ATF4 knockout mice with glucose tolerance tests

    PMID:22298775

    Open questions at the time
    • Structural basis of FoxO1–ATF4 interaction unknown
    • Whether this partnership operates outside osteoblasts was not addressed
  4. 2013 High

    Demonstrating that ATF4 stabilizes HIF-1α by blocking prolyl hydroxylase access and separately activates Sestrin2 to inhibit mTOR expanded ATF4's role from stress-responsive transcription factor to a coordinator of hypoxia adaptation and autophagy.

    Evidence ATF4 KO mice, Co-IP/ubiquitination assays (HIF-1α); ectopic ATF4 expression with mTOR activity and autophagy readouts (Sestrin2)

    PMID:23649506 PMID:23916134

    Open questions at the time
    • Whether the HIF-1α interaction is direct via the bZIP domain was not structurally resolved
    • Sestrin2 induction lacked ChIP evidence for direct promoter binding at this stage
  5. 2014 High

    Two parallel discoveries—that ATF4 is locally translated in axons as a retrograde injury signal and that it is sufficient to drive skeletal muscle atrophy—established that ATF4 has both neurodegenerative and muscle-wasting functions beyond classical ER stress.

    Evidence Axonal siRNA knockdown, retrograde transport inhibition, in vivo Aβ injection (axonal); muscle-specific KO/overexpression with immobilization model (atrophy)

    PMID:24895282 PMID:25171414

    Open questions at the time
    • Cargo mechanism for ATF4 retrograde transport not identified
    • Downstream atrophy genes beyond p21 were not fully catalogued
  6. 2015 High

    Showing that ATF4 co-activates HO-1 with NRF2 to confer anoikis resistance and enable lung colonization of tumor cells linked ATF4's antioxidant function to metastatic fitness.

    Evidence ATF4-deficient fibrosarcoma cells, HO-1 reconstitution, murine lung colonization assay

    PMID:26011642

    Open questions at the time
    • Whether ATF4 and NRF2 directly co-occupy the HO-1 promoter was not resolved by sequential ChIP
  7. 2016 Medium

    KDM4C was identified as a chromatin-modifying partner of ATF4 that removes H3K9me3 at serine biosynthesis genes, and ATF4 was shown to amplify IRE1α–XBP1 signaling, establishing cross-talk between UPR branches and epigenetic control of amino acid metabolism.

    Evidence Co-IP and ChIP for KDM4C–ATF4 at serine pathway genes; ATF4 knockdown/overexpression measuring IRE1α and XBP1 splicing

    PMID:26774480 PMID:27052593

    Open questions at the time
    • Whether KDM4C recruitment requires ATF4's bZIP DNA-binding activity is unknown
    • IRE1α crosstalk was demonstrated in one cell line
  8. 2017 Medium

    Identification of PARP-1-mediated PARylation of ATF4 (reducing its DNA binding) and DDX3-dependent translational promotion of ATF4 mRNA revealed two new regulatory layers—post-translational modification and translational control—that tune ATF4 output.

    Evidence In vitro PARylation assay plus ChIP-qPCR (PARylation); DDX3 siRNA with polyribosome profiling and luciferase reporters (translation)

    PMID:28457938 PMID:29062139

    Open questions at the time
    • PARylation sites on ATF4 were not mapped
    • DDX3 interaction with ATF4 mRNA structural elements not resolved
  9. 2018 High

    Demonstrating that ATF4 is required in CD4+ T cells for amino acid uptake, mTORC1 activation, redox homeostasis, and effector differentiation expanded ATF4's role to adaptive immunity.

    Evidence Atf4-deficient mice, T cell functional assays, metabolic measurements

    PMID:29742431

    Open questions at the time
    • Whether ATF4 operates through the same target genes in T cells as in fibroblasts was not fully resolved
    • Role in CD8+ T cells not addressed
  10. 2019 High

    Multiple discoveries converged: PRMT1 methylates ATF4-Arg239 to stabilize it; ATF4 directly binds the LAMP3 promoter; DISC1 restrains ATF4 nuclear activity (crystal structure solved); Hop2 enhances ATF4 transcription in bone; and ATF4 induces HSPA5/GPX4 to counteract ferroptosis—collectively defining how post-translational modifications, structural partnerships, and target gene specificity shape ATF4 output.

    Evidence R239K mutagenesis in cardiomyocytes (PRMT1); ChIP and luciferase on LAMP3 promoter; DISC1–ATF4 crystal structure with iPSC neurons; Hop2 KO mice and compound heterozygotes; siRNA epistaxis for PERK→ATF4→HSPA5→GPX4

    PMID:31433867 PMID:31444471 PMID:31519193 PMID:31787756 PMID:32312748

    Open questions at the time
    • Whether PRMT1 methylation affects ATF4 dimerization partner choice is unknown
    • DISC1 structural data has not been extended to a full-length complex
    • Hop2–ATF4 mechanism awaits reconstitution outside bone
  11. 2020 High

    ATF4 was shown to mediate reticulophagy via RETREG1/TEX264, drive TRAIL-R2-dependent apoptosis under glucose deprivation, undergo PKA-Ser219-dependent proteasomal degradation, be stabilized by FAM175B, regulate the mitochondrial UPR in alveolar cells, and suppress mTORC1 via dual REDD1/Sestrin2 induction during leucine deprivation—consolidating its role as a versatile stress integrator whose protein levels and transcriptional program are context-dependent.

    Evidence ATF4 KO with EM for reticulophagy; TRAIL-R2/FADD/caspase-8 ablation; phospho-Ser219 antibody and proteasome inhibitor; FAM175B Co-IP/ubiquitination assay; ATF4 KD with directional UPRmt analysis; triple KO MEFs with mTORC1 time-course

    PMID:28242652 PMID:30854784 PMID:31875479 PMID:32551949 PMID:33111629 PMID:33198401

    Open questions at the time
    • β-TrCP as the E3 ligase for ATF4 was suggested but not confirmed by reconstitution
    • FAM175B mechanism of ubiquitin inhibition unclear
    • UPRmt targets directly activated by ATF4 were not catalogued
  12. 2021 High

    A suite of studies revealed: mTORC1 activates ATF4 independently of eIF2α phosphorylation; asparagine signals mitochondrial status through ATF4; ATF4 mediates TRPM3 trafficking (non-transcriptional); Thbs1–PERK–ATF4 drives cardiac atrophy; ATF4 represses NRF1/TFAM for mitochondrial biogenesis; YAP/TAZ stabilize ATF4 for SLC7A11-dependent ferroptosis resistance; ATF4 promotes NADPH production in cardiomyocytes; lncRNA TINCR suppresses ATF4 translation; and O-GlcNAcylation tunes ATF4—collectively demonstrating that ATF4 integrates nutrient, mechanical, and oncogenic signals through transcription-dependent and -independent mechanisms.

    Evidence ATF4 KO MEFs with RNA-seq for mTORC1-specific program; asparagine rescue in mouse tumors; ATF4 KO DRG neurons with electrophysiology and Co-IP of TRPM3/KIF17; cardiac PERK KO and AAV9-ATF4; hepatocyte-specific ATF4 KO with ChIP on NRF1; shRNA screen with ferroptosis assays; cardiomyocyte-specific KO with isotope tracing; TINCR RIP and polyribosome profiling; OGT KD with ChIP-qPCR

    PMID:33177163 PMID:33586907 PMID:33609439 PMID:33646118 PMID:33658516 PMID:34168130 PMID:34664408 PMID:35574856 PMID:38192280

    Open questions at the time
    • Structural basis of mTORC1-dependent ATF4 translation unknown
    • How ATF4 switches between transcriptional and trafficking functions is unresolved
    • O-GlcNAcylation site(s) on ATF4 not mapped
  13. 2022 Medium

    New regulatory axes were defined: PRMT5 controls ATF4 mRNA splicing; GADD34 positively and CHOP negatively feed back on ATF4; ATF4 cooperates with CTCF in adipogenesis; and C. elegans ATF-4 extends lifespan via H2S/persulfidation—broadening ATF4 regulation to RNA processing, chromatin architecture in differentiation, and longevity.

    Evidence PRMT5 inhibitor with nuclear/cytoplasmic fractionation; siRNA of GADD34/CHOP with computational modeling; ChIP-seq and Co-IP in adipocytes; C. elegans atf-4 LOF with H2S measurement and lifespan assays

    PMID:33950334 PMID:35181679 PMID:35305370 PMID:36097827

    Open questions at the time
    • PRMT5 splicing regulation specificity for ATF4 vs. other transcripts not established
    • CTCF–ATF4 structural interaction not characterized
    • Conservation of H2S/persulfidation mechanism in mammals not tested
  14. 2023 High

    ATF4 was established as a bidirectional regulator of NRF2 (via CHAC1), an essential suppressor of hepatocyte ferroptosis through SLC7A11, a driver of age-related muscle decline, a mediator of Thbs1–TGFβ–Smad2/3 muscle atrophy, and a regulator of intestinal stem cell self-renewal via Sox9—completing a picture of ATF4 as a tissue-level determinant of redox fate, aging, and regeneration.

    Evidence ATF4 gain/loss with NRF2 reporter and CHAC1 KD; hepatocyte-specific ATF4 KO with SLC7A11 rescue in HCC model; muscle-specific ATF4 KO at 22 months; skeletal muscle Smad2/3 and ATF4 KO; circBtnl1 KO with RIP and organoid assays

    PMID:36715460 PMID:36996941 PMID:37014538 PMID:37410595 PMID:38678560

    Open questions at the time
    • Whether ATF4-driven aging phenotype is reversible by late-life deletion is untested
    • Whether circBtnl1–Ddx3y control of ATF4 operates in tissues beyond intestine is unknown
    • Genome-wide catalog of ATF4 direct targets across tissues remains incomplete

Open questions

Synthesis pass · forward-looking unresolved questions
  • Key unresolved questions include the structural basis for mTORC1-dependent (eIF2α-independent) ATF4 translational activation, the mechanism by which ATF4 switches between transcriptional and non-transcriptional (trafficking) functions, comprehensive mapping of ATF4 post-translational modification sites and their interplay, and the degree to which ATF4 target gene programs are tissue-specific versus universal.
  • No structural model for mTORC1–ATF4 translational mechanism
  • No systematic tissue-resolved ATF4 cistrome
  • Interplay among PRMT1 methylation, PARylation, O-GlcNAcylation, and phosphorylation at the protein level not addressed

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 12 GO:0003677 DNA binding 5
Localization
GO:0005634 nucleus 3 GO:0005829 cytosol 2
Pathway
R-HSA-74160 Gene expression (Transcription) 7 R-HSA-8953897 Cellular responses to stimuli 5 R-HSA-9612973 Autophagy 5 R-HSA-1430728 Metabolism 4 R-HSA-162582 Signal Transduction 4 R-HSA-392499 Metabolism of proteins 3 R-HSA-5357801 Programmed Cell Death 3
Partners
CTCFDISC1FAM175BFOXO1KDM4CNRF2PSME3IP1TRPM3

Evidence

Reading pass · 46 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2006 PERK phosphorylates eIF2α, which leads to translational upregulation of ATF4, placing ATF4 downstream of PERK-eIF2α in the UPR/integrated stress response. Cells with compromised PERK-eIF2α-ATF4 signaling are more sensitive to hypoxic stress in vitro and form slower-growing tumors in vivo. Genetic loss-of-function (PERK-deficient and eIF2α mutant MEFs), in vitro hypoxia assays, xenograft tumor models Cancer biology & therapy High 16861899
2008 ATF4 is necessary and sufficient for ER stress-induced transcriptional upregulation of REDD1. In MEFs lacking ATF4, ER stress fails to upregulate REDD1 mRNA, while exogenous ATF4 expression is sufficient to induce REDD1. ATF4-deficient MEFs, PERK-deficient MEFs, exogenous ATF4 overexpression, mRNA expression analysis Biochemical and biophysical research communications High 19114033
2013 ATF4 directly interacts with HIF-1α in hypoxic osteoblasts and promotes HIF-1α protein stability by reducing its binding to prolyl hydroxylases, thereby preventing pVHL-mediated proteasomal degradation. Loss of ATF4 increases HIF-1α ubiquitination and reduces VEGF expression, impairing bone angiogenesis. Co-immunoprecipitation, ubiquitination assay, ATF4 knockout mice, in vitro osteoblast studies, metatarsal sprouting assay Journal of bone and mineral research High 23649506
2013 ATF4 transcriptionally upregulates sestrin-2 (SESN2), which in turn inhibits mTOR complex activity and induces autophagy. Ectopic ATF4 expression (but not ATF3 or CHOP) is sufficient to drive SESN2 transcription. Ectopic expression of ATF4, ATF3, CHOP; mRNA and protein expression analysis; mTOR activity assay; autophagy readout Molecular oncology Medium 23916134
2014 ATF4 mRNA is locally translated in axons in response to Aβ1-42 and the resulting ATF4 protein is retrogradely transported to the nucleus where it drives transcriptional activity and cell loss. Inhibition of local translation, retrograde transport, or knockdown of axonal Atf4 mRNA abolishes these effects. Local Aβ application to axons, knockdown of axonal Atf4 mRNA with siRNA, retrograde transport inhibition, in vivo dentate gyrus injection in mice, analysis of AD patient brain tissue Cell High 25171414
2014 ATF4 and p53 mediate distinct and additive pathways to skeletal muscle atrophy during immobilization. Forced expression of ATF4 in muscle fibers is sufficient to induce atrophy, and ATF4-deficient fibers are partially resistant to immobilization-induced atrophy. ATF4 and p53 cooperate to induce p21 expression as a downstream effector of atrophy. Mouse muscle ATF4 and p53 KO/overexpression, limb immobilization model, genome-wide mRNA arrays, in vivo fiber size measurement American journal of physiology. Endocrinology and metabolism High 24895282
2015 ATF4 activates heme oxygenase 1 (HO-1) expression through simultaneous activation with NRF2, both converging on the HO1 promoter. HO-1 upregulation by ATF4 ameliorates oxidative stress and protects cells from anoikis following matrix detachment. Reconstitution of ATF4 or HO-1 in ATF4-deficient cells rescues anoikis resistance and tumor lung colonization. ATF4-deficient human fibrosarcoma cells, HO-1 reconstitution, murine lung colonization model, ATF4/NRF2 co-activation analysis The Journal of clinical investigation High 26011642
2016 KDM4C interacts with ATF4 and together they co-activate amino acid biosynthesis and transport genes. KDM4C activates ATF4 transcription and physically interacts with ATF4 to target serine pathway genes, removing repressive H3K9 trimethylation. Co-immunoprecipitation, ChIP, gene expression analysis, serine pathway reporter assays Cell reports Medium 26774480
2016 ATF4 enhances IRE1α expression, increasing the XBP1 mRNA splicing ratio under ER stress, establishing a cross-talk mechanism where the PERK-ATF4 branch amplifies IRE1α-XBP1 signaling. Knockdown and overexpression of ATF4, measurement of IRE1α expression and XBP1 splicing ratio under ER stress Scientific reports Medium 27052593
2017 ATF4 protein levels are regulated by PARylation: PARP-1 activation induces PARylation of ATF4 and reduces its binding to CRE sequences in vitro, while PARP inhibition increases ATF4 occupancy at the MKP-1 promoter (by ChIP-qPCR), leading to MKP-1 expression and JNK/p38 inactivation. PARylation assay in vitro, ChIP-qPCR, PARP inhibitor/siRNA knockdown, ATF4/MKP-1 functional pathway analysis Free radical biology & medicine Medium 28457938
2017 FLT3-ITD promotes autophagy in AML cells through ATF4. Cellular levels of ATF4 are highly dependent on FLT3-ITD activity; ATF4 downregulation inhibits autophagy-dependent AML cell proliferation. FLT3 inhibition (pharmacological and genetic), ATF4 knockdown, autophagy assays, xenograft mouse model Oncogene Medium 29059168
2018 ATF4 regulates a coordinated gene network in CD4+ T cells driving amino acid intake, mTORC1 activation, protein translation, and glutathione synthesis. ATF4-deficient CD4+ T cells have defects in redox homeostasis, proliferation, differentiation, and cytokine production. Atf4-deficient mice, CD4+ T cell functional assays, gene expression analysis, metabolic measurements Cell reports High 29742431
2018 EIF1AX-A113splice mutation stabilizes the preinitiation complex, induces ATF4, and ATF4 in turn suppresses eIF2α phosphorylation, enabling increased global protein synthesis. ATF4 cooperates with RAS-stabilized c-MYC to induce amino acid transporters and sensitize mTOR to amino acid supply. Isogenic cell lines with EIF1AX mutations, mouse tumorigenesis models, protein synthesis measurement, epistasis experiments Cancer discovery High 30305285
2019 PRMT1 methylates ATF4 at arginine 239, promoting ATF4 protein stability. The methylation-deficient ATF4 R239K mutant exacerbates ER stress and apoptosis. PRMT1 inhibition augments ER stress response via ATF4, while PRMT1 overexpression attenuates it in cardiomyocytes. ATF4 methylation-deficient mutant (R239K), PRMT1 overexpression/inhibition, ER stress assays in cardiomyocytes and PRMT1 null hearts Cell death & disease High 31787756
2019 DHA (dihydroartemisinin) causes ER stress in glioma cells, activating PERK, which upregulates ATF4, which in turn induces HSPA5 expression. HSPA5 increases GPX4 activity, attenuating ferroptosis in a negative feedback loop. siRNA knockdown of PERK, ATF4, HSPA5; small molecule inhibitors; lipid peroxidation and ferroptosis assays in vitro and in vivo Journal of experimental & clinical cancer research High 31519193
2019 ATF4 directly binds the LAMP3 gene promoter (verified by ChIP assay) and transcriptionally upregulates LAMP3 mRNA levels during the integrated stress response. LAMP3 regulation by ATF4 is confirmed by siRNA knockdown and exogenous ATF4 overexpression, plus dual-luciferase assay confirming the ATF4-binding site requirement. ChIP, siRNA knockdown of ATF4, exogenous ATF4 overexpression, dual-luciferase reporter assay The Journal of biological chemistry High 32312748
2019 DISC1 physically interacts with ATF4 (bZIP transcription factor). Mutation of DISC1 disrupts normal DISC1-ATF4 interaction, resulting in excessive ATF4 nuclear localization and DNA binding. The crystal structure of the DISC1-ATF4 complex was solved, revealing the atomic basis of their interaction. Co-immunoprecipitation, crystal structure determination, CRISPR-mediated ATF4 heterozygous knockout in iPSC-derived neurons, ChIP-seq Molecular psychiatry High 31444471
2020 ATF4 knockout significantly attenuates loperamide-induced autophagy and autophagic cell death in glioblastoma, and specifically mediates reticulophagy (selective ER degradation) through reticulophagy receptors RETREG1/FAM134B and TEX264. ATF4 knockout cells, electron microscopy, fluorescence microscopy for ER engulfment, reticulophagy receptor knockdown Autophagy Medium 33111629
2020 ATF4, but not ATF5, is the key regulator of the mitochondrial unfolded protein response (UPRmt) in alveolar epithelial cells. ER stress (UPRER) leads to UPRmt in an ATF4-dependent manner, while mitochondrial stress does not activate UPRER. ATF4 knockdown, ER stress induction, mitochondrial stress induction, UPRmt marker analysis, inducible ATF4 expression in mouse alveolar epithelial cells in vivo American journal of respiratory cell and molecular biology High 32551949
2020 ATF4 is induced by glucose deprivation and mediates CHOP-independent upregulation of TRAIL-R2 (DR5), leading to ligand-independent TRAIL receptor-mediated apoptosis. Ablation of TRAIL-R2, FADD, Bid, or caspase-8 attenuates this cell death. ATF4/CHOP knockdown, TRAIL receptor ablation, glucose deprivation model, apoptosis assays Molecular and cellular biology Medium 28242652
2020 ATF4 protein stability is regulated by ubiquitin-proteasome-mediated proteolysis. Phosphorylation of ATF4 at serine-219 by cAMP-dependent protein kinase increases during chemically induced LTP and decreases thereafter; proteasome inhibition prevents ATF4 degradation. β-TrCP may act as the ubiquitin ligase. Chemical LTP induction in hippocampal neurons, proteasome inhibitor treatment, phospho-specific antibody analysis, β-TrCP investigation International journal of molecular sciences Medium 33198401
2020 FAM175B interacts with ATF4 (co-localization by confocal microscopy and co-immunoprecipitation) and inhibits ubiquitin-dependent ATF4 degradation, elevating ATF4 protein levels and promoting CHOP-dependent apoptosis in esophageal squamous cell carcinoma. Co-IP, confocal colocalization, ubiquitination assay, siRNA knockdown, luciferase reporter Molecular oncology Medium 30854784
2021 mTORC1 activates ATF4 through mechanisms distinct from canonical ISR/eIF2α phosphorylation, and this mTORC1-ATF4 pathway stimulates amino acid uptake, synthesis, tRNA charging, and cellular cystine uptake for glutathione synthesis. Insulin-stimulated mTORC1 activation, ATF4 KO MEFs and cancer cell lines, RNA-seq comparison of mTORC1-ATF4 vs ISR-ATF4 transcriptional programs, metabolic assays eLife High 33646118
2021 Asparagine depletion (via ETC inhibition) increases ATF4 levels and impairs mTORC1. Exogenous asparagine restores ATF4 and mTORC1 activities, identifying asparagine as a signal communicating active mitochondrial respiration to ATF4 and mTORC1. ETC inhibitor (metformin), exogenous asparagine supplementation, asparaginase treatment, mouse tumor models, mTORC1 activity assays Cell metabolism High 33609439
2021 Thbs1 binds and activates PERK, inducing downstream ATF4 expression and causing lethal autophagy-mediated cardiac atrophy. Deletion of PERK in Thbs1 transgenic mice blunts ATF4 induction and autophagy and largely corrects lethal cardiac atrophy. AAV9-mediated overexpression of PERK or ATF4 alone is sufficient to promote cardiac atrophy. Transgenic mice, PERK cardiac-specific KO, AAV9 gene transfer, autophagy markers, cardiac phenotype analysis Nature communications High 34168130
2021 ATF4 selectively regulates heat nociception by interacting with TRPM3 and mediating its membrane trafficking via KIF17 in dorsal root ganglion neurons. Loss of ATF4 decreases TRPM3 current and KIF17-mediated trafficking, identifying a non-transcriptional function of ATF4. ATF4 KO in DRG neurons, co-immunoprecipitation of ATF4-TRPM3-KIF17, electrophysiology, membrane trafficking assays, heat sensitivity behavioral tests Nature communications High 33658516
2021 ATF4 represses NRF1 transcriptional activity by binding to the NRF1 promoter region, thereby suppressing TFAM expression and impairing mitochondrial biogenesis and respiratory function in hepatocytes during alcohol-induced steatohepatitis. Hepatocyte-specific ATF4 KO mice, TFAM siRNA, ChIP (ATF4 binding to NRF1 promoter), mitochondrial function assays, liver-specific TFAM overexpression mice Gut High 33177163
2021 YAP/TAZ sustain ATF4 protein stability, nuclear localization, and transcriptional activity in a TEAD-dependent manner. ATF4 cooperates with YAP/TAZ to induce SLC7A11 expression, maintaining glutathione homeostasis and enabling resistance to ferroptosis. shRNA screening, transcriptomic analysis, ATF4 localization studies, SLC7A11 reporter assays, ferroptosis assays EMBO molecular medicine Medium 34664408
2021 ATF4 activates expression of NADPH-producing enzymes in both the pentose phosphate pathway and mitochondrial serine/glycine/folate metabolic pathway in cardiomyocytes. Cardiomyocyte-specific ATF4 KO exacerbates cardiomyopathy under pressure overload due to inability to induce these NADPH-generating enzymes; stable isotope tracing confirms ATF4 augments metabolic flux in these pathways. Cardiomyocyte-specific ATF4 KO mice, RNA-seq, metabolomics, stable isotope tracer experiments, transverse aortic constriction model Circulation research High 35574856
2022 PRMT5 regulates ATF4 mRNA splicing; PRMT5 inhibition results in expression of intron-retaining, nuclear-detained ATF4 mRNA, reducing cytoplasmic spliced ATF4 transcript and protein levels, leading to decreased ATF4 target gene expression and increased oxidative stress. PRMT5 inhibitor treatment, RNA-seq, nuclear/cytoplasmic fractionation, ATF4 protein/mRNA measurement, oxidative stress assays Redox biology Medium 35305370
2022 ATF4 and CTCF cooperate in adipogenesis by co-localizing at promoters of key adipogenic genes (Cebpd, PPARγ). ATF4 directly regulates CTCF expression and physically interacts with CTCF in differentiated adipocytes. ATF4 genome-wide binding was mapped by ChIP-seq. ChIP-seq, RNA-seq, Co-IP, CTCF deletion and overexpression, in vivo adipose tissue analysis Cell biology and toxicology Medium 33950334
2022 In C. elegans, ATF-4 mediates longevity downstream of mTORC1 inhibition and translation inhibition by activating CTH-2 (transsulfuration enzyme) to increase hydrogen sulfide (H2S) production, which increases protein persulfidation as a protective modification. C. elegans genetic analysis, ATF-4 loss-of-function, H2S measurement, persulfidation analysis, lifespan assays Nature communications Medium 35181679
2022 GADD34 positively regulates ATF4 activity in a feedback loop within the PERK pathway, while CHOP inhibits ATF4 activity, providing fine-tuning of the ER stress response. These feedback loops were identified by combined molecular biology and computational modeling. siRNA knockdown of GADD34/CHOP, ATF4 activity measurement, mathematical modeling, HEK293T cells under ER stress FEBS open bio Medium 36097827
2023 ATF4 is an obligatory metabolic activator of NRF2: ATF4 increases NRF2 transcription and induces CHAC1 (glutathione-degrading enzyme), which is critically important for maintaining NRF2 activation. NRF2 in turn supports ATF4-induced stress response by increasing cystine uptake via xCT and upregulating thioredoxin pathways. ATF4 gain/loss-of-function, NRF2 reporter assays, CHAC1 knockdown, cystine uptake measurements, pathway analysis Cell reports Medium 37410595
2023 ATF4 suppresses ferroptosis in hepatocytes by transcriptionally activating SLC7A11 (xCT), maintaining cystine uptake and glutathione synthesis. Hepatocyte-specific ATF4 ablation increases susceptibility to ferroptosis and accelerates HCC development; ectopic SLC7A11 expression reverses these effects. Hepatocyte-specific ATF4 KO mice, SLC7A11 reconstitution, ferroptosis inhibitor experiments, RNA-seq, diethylnitrosamine HCC model Journal of hepatology High 36996941
2023 Thbs1 activates TGFβ-Smad2/3 signaling, which induces ATF4 expression in skeletal muscle. ATF4 in turn modulates the autophagy-lysosomal pathway and ubiquitin-proteasome system to facilitate muscle atrophy. Myofiber-specific deletion of Smad2/3 or ATF4 antagonizes Thbs1-induced muscle atrophy. Skeletal muscle-specific Thbs1 transgenic mice, Smad2/3 KO, ATF4 KO, denervation and caloric restriction atrophy models Cell reports High 38678560
2023 Circular RNA circBtnl1 competes with Atf4 mRNA for binding to the RNA helicase Ddx3y, thereby impairing ATF4 mRNA stability and reducing ATF4 expression in intestinal stem cells. ATF4 activates Sox9 transcription by binding to its promoter, enhancing ISC self-renewal. circBtnl1 KO mice, RIP (RNA immunoprecipitation) for Ddx3y binding, Atf4 mRNA stability assay, ATF4 ChIP on Sox9 promoter, organoid self-renewal assays The EMBO journal High 36715460
2020 ATF4 mediates mTORC1 re-suppression during prolonged leucine deprivation by transcriptionally upregulating both REDD1 and Sestrin2. In ATF4-deficient cells, neither REDD1 nor Sestrin2 is upregulated by leucine deprivation, and mTORC1 resuppression is absent. ATF4, REDD1, and/or Sestrin2 deficient MEFs, leucine deprivation time-course, mTORC1 activity measurement The Journal of nutrition High 31875479
2017 DDX3 (RNA-binding protein) promotes ATF4 mRNA translation under ER stress by interacting with the eIF4F complex. DDX3 depletion prevents phospho-eIF2α-mediated ATF4 expression; luciferase and polyribosome assays confirm DDX3 acts at the translational level on ATF4 mRNA. DDX3 siRNA knockdown, luciferase reporter assay, polyribosome profiling, protein interaction assay (DDX3-eIF4F binding) Scientific reports Medium 29062139
2019 LRPPRC deficiency in brown adipocytes causes mitochondrial ETC proteome imbalance and activates ATF4. ATF4 activation in brown adipocytes (by overexpression or low-protein diet) improves cold tolerance and systemic metabolism independently of UCP1 action. BA-specific Lrpprc KO, BA-specific ATF4 overexpression, low-protein diet model, Ucp1 KO mice, cold-tolerance tests, proteomics Cell reports Medium 34551310
2012 FoxO1 co-localizes with ATF4 in the osteoblast nucleus, physically interacts with ATF4, and promotes ATF4 transcriptional activity. FoxO1 and ATF4 cooperate to suppress osteocalcin activity by up-regulating expression of the phosphatase catalyzing osteocalcin inactivation, affecting glucose homeostasis. Co-immunoprecipitation, colocalization studies, genetic epistasis (FoxO1/ATF4 compound knockout), glucose tolerance tests The Journal of biological chemistry Medium 22298775
2019 Hop2 physically interacts with ATF4 via its Zip domain (identified by yeast two-hybrid, confirmed by deletional mapping) and enhances ATF4-dependent transcription. Hop2 deficiency in mice causes an osteopenic phenotype similar to ATF4-deficient mice; Atf4+/-:Hop2+/- compound heterozygotes display additive skeletal defects. Yeast two-hybrid, deletional domain mapping, Hop2 KO mice, compound heterozygous mice, osteocalcin/collagen expression Journal of bone and mineral research Medium 31433867
2021 O-GlcNAcylation regulates ATF4 protein levels during the mitochondrial integrated stress response. OGT knockdown increases ATF4 protein and mRNA expression, and ATF4 occupancy at the ATF5 promoter is increased in brains of TMG-treated mice (demonstrated by ChIP). OGT knockdown, O-GlcNAcase inhibition (Thiamet-G), ChIP-qPCR, iPSC-derived neurons, AD mouse model Frontiers in aging neuroscience Medium 38192280
2023 ATF4 is an essential mediator of skeletal muscle aging. Muscle-specific ATF4 KO mice are protected from age-related declines in strength, muscle quality, exercise capacity, and muscle mass at 22 months, with altered turnover of key structural/metabolic proteins. Muscle-specific ATF4 KO mice at 6 and 22 months, strength/exercise phenotyping, RNA-seq, proteomics GeroScience High 37014538
2021 ATF4 protein levels in AML cells are regulated downstream of oncogenic FLT3-ITD signaling. BRAF ensures ATF4 induction through mTOR and eIF4B as downstream regulators during GCN2 activation, providing a non-canonical route to ATF4 that remains transiently active during BRAF inhibitor treatment. BRAF inhibitors, GCN2 activation, mTOR/eIF4B epistasis experiments, ATF4 protein measurement, synergistic killing assays iScience Medium 32283529
2021 LNCR TINCR suppresses ATF4 translation by interacting with ATF4 mRNA and preventing its ribosome binding, establishing a non-canonical (eIF2α phosphorylation-independent) mechanism of ATF4 translational control in melanoma cells. TINCR knockdown/re-expression, polyribosome profiling, RNA immunoprecipitation of TINCR-ATF4 mRNA interaction, global protein synthesis measurement EMBO reports Medium 33586907

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2021 YAP/TAZ and ATF4 drive resistance to Sorafenib in hepatocellular carcinoma by preventing ferroptosis. EMBO molecular medicine 401 34664408
2023 ATF4 suppresses hepatocarcinogenesis by inducing SLC7A11 (xCT) to block stress-related ferroptosis. Journal of hepatology 290 36996941
2006 The PERK/eIF2alpha/ATF4 module of the UPR in hypoxia resistance and tumor growth. Cancer biology & therapy 288 16861899
2014 Axonally synthesized ATF4 transmits a neurodegenerative signal across brain regions. Cell 282 25171414
2019 Dihydroartemisinin-induced unfolded protein response feedback attenuates ferroptosis via PERK/ATF4/HSPA5 pathway in glioma cells. Journal of experimental & clinical cancer research : CR 260 31519193
2015 ATF4-dependent induction of heme oxygenase 1 prevents anoikis and promotes metastasis. The Journal of clinical investigation 234 26011642
2021 Asparagine couples mitochondrial respiration to ATF4 activity and tumor growth. Cell metabolism 219 33609439
2021 The mTORC1-mediated activation of ATF4 promotes protein and glutathione synthesis downstream of growth signals. eLife 184 33646118
2019 Artesunate activates the ATF4-CHOP-CHAC1 pathway and affects ferroptosis in Burkitt's Lymphoma. Biochemical and biophysical research communications 173 31537387
2009 Role of ATF4 in regulation of autophagy and resistance to drugs and hypoxia. Cell cycle (Georgetown, Tex.) 165 19887912
2008 ATF4 is necessary and sufficient for ER stress-induced upregulation of REDD1 expression. Biochemical and biophysical research communications 152 19114033
2009 JNK activity is essential for Atf4 expression and late-stage osteoblast differentiation. Journal of bone and mineral research : the official journal of the American Society for Bone and Mineral Research 138 19016586
2023 A stay of execution: ATF4 regulation and potential outcomes for the integrated stress response. Frontiers in molecular neuroscience 134 36825279
2021 Thbs1 induces lethal cardiac atrophy through PERK-ATF4 regulated autophagy. Nature communications 133 34168130
2010 ATF4 regulates lipid metabolism and thermogenesis. Cell research 132 20066008
2016 KDM4C and ATF4 Cooperate in Transcriptional Control of Amino Acid Metabolism. Cell reports 130 26774480
2000 Microphthalmia due to p53-mediated apoptosis of anterior lens epithelial cells in mice lacking the CREB-2 transcription factor. Developmental biology 121 10885750
2020 MicroRNA-214-3p enhances erastin-induced ferroptosis by targeting ATF4 in hepatoma cells. Journal of cellular physiology 120 31960438
2020 ATF4 activation promotes hepatic mitochondrial dysfunction by repressing NRF1-TFAM signalling in alcoholic steatohepatitis. Gut 120 33177163
2020 ATF4 links ER stress with reticulophagy in glioblastoma cells. Autophagy 117 33111629
2013 Nelfinavir and bortezomib inhibit mTOR activity via ATF4-mediated sestrin-2 regulation. Molecular oncology 116 23916134
2012 Targeting the ATF4 pathway in cancer therapy. Expert opinion on therapeutic targets 114 23009153
2016 Novel mechanism of enhancing IRE1α-XBP1 signalling via the PERK-ATF4 pathway. Scientific reports 110 27052593
2017 Glucose Deprivation Induces ATF4-Mediated Apoptosis through TRAIL Death Receptors. Molecular and cellular biology 109 28242652
2023 The integrated stress response effector ATF4 is an obligatory metabolic activator of NRF2. Cell reports 94 37410595
2022 ATF4 Protects the Heart From Failure by Antagonizing Oxidative Stress. Circulation research 93 35574856
2017 Oncogenic FLT3-ITD supports autophagy via ATF4 in acute myeloid leukemia. Oncogene 92 29059168
2014 p53 and ATF4 mediate distinct and additive pathways to skeletal muscle atrophy during limb immobilization. American journal of physiology. Endocrinology and metabolism 91 24895282
2017 PARP inhibition protects mitochondria and reduces ROS production via PARP-1-ATF4-MKP-1-MAPK retrograde pathway. Free radical biology & medicine 88 28457938
2022 ATF-4 and hydrogen sulfide signalling mediate longevity in response to inhibition of translation or mTORC1. Nature communications 86 35181679
2018 ATF4 Regulates CD4+ T Cell Immune Responses through Metabolic Reprogramming. Cell reports 86 29742431
2024 ATF4 in cellular stress, ferroptosis, and cancer. Archives of toxicology 80 38383612
2018 Role of the ISR-ATF4 pathway and its cross talk with Nrf2 in mitochondrial quality control. Journal of clinical biochemistry and nutrition 79 30705506
2018 EIF1AX and RAS Mutations Cooperate to Drive Thyroid Tumorigenesis through ATF4 and c-MYC. Cancer discovery 78 30305285
2017 Neurodegeneration: Keeping ATF4 on a Tight Leash. Frontiers in cellular neuroscience 68 29326555
2017 The oxido-metabolic driver ATF4 enhances temozolamide chemo-resistance in human gliomas. Oncotarget 62 28881638
2020 ATF4 Mediates Mitochondrial Unfolded Protein Response in Alveolar Epithelial Cells. American journal of respiratory cell and molecular biology 61 32551949
2023 PDIA4 confers resistance to ferroptosis via induction of ATF4/SLC7A11 in renal cell carcinoma. Cell death & disease 60 36906674
2012 FoxO1 protein cooperates with ATF4 protein in osteoblasts to control glucose homeostasis. The Journal of biological chemistry 60 22298775
2019 The PERK-EIF2α-ATF4 signaling branch regulates osteoblast differentiation and proliferation by PTH. American journal of physiology. Endocrinology and metabolism 58 30668150
2013 ATF4 promotes bone angiogenesis by increasing VEGF expression and release in the bone environment. Journal of bone and mineral research : the official journal of the American Society for Bone and Mineral Research 58 23649506
2017 ATF4 overexpression induces early onset of hyperlipidaemia and hepatic steatosis and enhances adipogenesis in zebrafish. Scientific reports 52 29180630
2019 Regulation of the unfolded protein response through ATF4 and FAM129A in prostate cancer. Oncogene 51 31312022
2020 ATF4-Mediated Upregulation of REDD1 and Sestrin2 Suppresses mTORC1 Activity during Prolonged Leucine Deprivation. The Journal of nutrition 48 31875479
2022 FOXO1 cooperates with C/EBPδ and ATF4 to regulate skeletal muscle atrophy transcriptional program during fasting. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 47 35061305
2013 An aryl hydrocarbon receptor induces VEGF expression through ATF4 under glucose deprivation in HepG2. BMC molecular biology 45 24330582
2022 Biology of Activating Transcription Factor 4 (ATF4) and Its Role in Skeletal Muscle Atrophy. The Journal of nutrition 44 34958390
2019 HER2 Upregulates ATF4 to Promote Cell Migration via Activation of ZEB1 and Downregulation of E-Cadherin. International journal of molecular sciences 43 31064130
2019 Activation of PERK-eIF2α-ATF4 pathway contributes to diabetic hepatotoxicity: Attenuation of ER stress by Morin. Cellular signalling 42 30877037
2022 Sevoflurane Induces Ferroptosis of Glioma Cells Through Activating the ATF4-CHAC1 Pathway. Frontiers in oncology 40 35372041
2017 ATF4 regulates CCL2 expression to promote endometrial cancer growth by controlling macrophage infiltration. Experimental cell research 39 28843961
2017 Role of ATF4 in skeletal muscle atrophy. Current opinion in clinical nutrition and metabolic care 38 28376050
2023 Advances in the roles of ATF4 in osteoporosis. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 37 37948991
2021 Activation of ATF4 triggers trabecular meshwork cell dysfunction and apoptosis in POAG. Aging 35 33714955
2013 ATF4 deficiency protects hepatocytes from oxidative stress via inhibiting CYP2E1 expression. Journal of cellular and molecular medicine 34 24373582
2021 Nox4 Promotes RANKL-Induced Autophagy and Osteoclastogenesis via Activating ROS/PERK/eIF-2α/ATF4 Pathway. Frontiers in pharmacology 33 34650437
2017 Up-Regulated ATF4 Expression Increases Cell Sensitivity to Apoptosis in Response to Radiation. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 33 28214891
2017 DDX3 regulates endoplasmic reticulum stress-induced ATF4 expression. Scientific reports 33 29062139
2017 Emerging Role for the PERK/eIF2α/ATF4 in Human Cutaneous Leishmaniasis. Scientific reports 31 29213084
2016 miRNA-1283 Regulates the PERK/ATF4 Pathway in Vascular Injury by Targeting ATF4. PloS one 31 27537404
2024 GPR68-ATF4 signaling is a novel prosurvival pathway in glioblastoma activated by acidic extracellular microenvironment. Experimental hematology & oncology 30 38291540
2022 PRMT5 regulates ATF4 transcript splicing and oxidative stress response. Redox biology 30 35305370
2019 Concomitant Nrf2- and ATF4-activation by Carnosic Acid Cooperatively Induces Expression of Cytoprotective Genes. International journal of molecular sciences 30 30959808
2023 The transcription regulator ATF4 is a mediator of skeletal muscle aging. GeroScience 29 37014538
2021 ATF4 selectively regulates heat nociception and contributes to kinesin-mediated TRPM3 trafficking. Nature communications 29 33658516
2019 Structural interaction between DISC1 and ATF4 underlying transcriptional and synaptic dysregulation in an iPSC model of mental disorders. Molecular psychiatry 29 31444471
2016 Crosstalk between ATF4 and MTA1/HDAC1 promotes osteosarcoma progression. Oncotarget 29 26797758
2024 Thbs1 regulates skeletal muscle mass in a TGFβ-Smad2/3-ATF4-dependent manner. Cell reports 28 38678560
2022 Gremlin-1 Promotes Colorectal Cancer Cell Metastasis by Activating ATF6 and Inhibiting ATF4 Pathways. Cells 28 35883579
2021 Long non-coding RNA TINCR suppresses metastatic melanoma dissemination by preventing ATF4 translation. EMBO reports 28 33586907
2019 PRMT1 suppresses ATF4-mediated endoplasmic reticulum response in cardiomyocytes. Cell death & disease 28 31787756
2013 Tetracyclines cause cell stress-dependent ATF4 activation and mTOR inhibition. Experimental cell research 28 24280420
2023 Tomatidine targets ATF4-dependent signaling and induces ferroptosis to limit pancreatic cancer progression. iScience 27 37554459
2018 GCN2 reduces inflammation by p-eIF2α/ATF4 pathway after intracerebral hemorrhage in mice. Experimental neurology 26 30529503
2021 Brown adipocyte ATF4 activation improves thermoregulation and systemic metabolism. Cell reports 25 34551310
2021 Cooperation of ATF4 and CTCF promotes adipogenesis through transcriptional regulation. Cell biology and toxicology 23 33950334
2020 Osteopromotive carbon dots promote bone regeneration through the PERK-eIF2α-ATF4 pathway. Biomaterials science 23 32307492
2020 The gene for the lysosomal protein LAMP3 is a direct target of the transcription factor ATF4. The Journal of biological chemistry 23 32312748
2022 A systems biological analysis of the ATF4-GADD34-CHOP regulatory triangle upon endoplasmic reticulum stress. FEBS open bio 22 36097827
2018 Investigation of Nrf2, AhR and ATF4 Activation in Toxicogenomic Databases. Frontiers in genetics 22 30333853
2018 P2X4R silence suppresses glioma cell growth through BDNF/TrkB/ATF4 signaling pathway. Journal of cellular biochemistry 22 30362154
2023 Noncoding RNA circBtnl1 suppresses self-renewal of intestinal stem cells via disruption of Atf4 mRNA stability. The EMBO journal 21 36715460
2021 Free Deoxycholic Acid Exacerbates Vascular Calcification in CKD through ER Stress-Mediated ATF4 Activation. Kidney360 21 34423309
2022 TRIB3 Mediates Fibroblast Activation and Fibrosis though Interaction with ATF4 in IPF. International journal of molecular sciences 20 36555349
2021 Inhibition of mTORC1 through ATF4-induced REDD1 and Sestrin2 expression by Metformin. BMC cancer 20 34253170
2024 Dihydroartemisinin induces ferroptosis of hepatocellular carcinoma via inhibiting ATF4-xCT pathway. Journal of cellular and molecular medicine 19 38652216
2023 Toosendanin induced hepatotoxicity via triggering PERK-eIF2α-ATF4 mediated ferroptosis. Toxicology letters 19 36801351
2023 Vitamin D receptor attenuate ischemia-reperfusion kidney injury via inhibiting ATF4. Cell death discovery 19 37173347
2020 Aspirin exerts anti-tumor effect through inhibiting Blimp1 and activating ATF4/CHOP pathway in multiple myeloma. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 19 32070879
2020 Disrupting ATF4 Expression Mechanisms Provides an Effective Strategy for BRAF-Targeted Melanoma Therapy. iScience 18 32283529
2017 Association of ATF4 Expression With Tissue Hypoxia and M2 Macrophage Infiltration in Infantile Hemangioma. The journal of histochemistry and cytochemistry : official journal of the Histochemistry Society 18 28438094
2020 Degradation of Transcriptional Repressor ATF4 during Long-Term Synaptic Plasticity. International journal of molecular sciences 17 33198401
2019 FAM175B promotes apoptosis by inhibiting ATF4 ubiquitination in esophageal squamous cell carcinoma. Molecular oncology 17 30854784
2023 O-GlcNAc regulates the mitochondrial integrated stress response by regulating ATF4. Frontiers in aging neuroscience 16 38192280
2020 Hydrogen peroxide induces nucleus pulposus cell apoptosis by ATF4/CHOP signaling pathway. Experimental and therapeutic medicine 16 32855694
2019 Hop2 Interacts with ATF4 to Promote Osteoblast Differentiation. Journal of bone and mineral research : the official journal of the American Society for Bone and Mineral Research 16 31433867
2016 Specificity of Stress-Responsive Transcription Factors Nrf2, ATF4, and AP-1. Journal of cellular biochemistry 16 27278863
2023 Shared Gene Targets of the ATF4 and p53 Transcriptional Networks. Molecular and cellular biology 15 37533313
2022 Hesperadin suppresses pancreatic cancer through ATF4/GADD45A axis at nanomolar concentrations. Oncogene 15 35551503
2022 ERH Interacts With EIF2α and Regulates the EIF2α/ATF4/CHOP Pathway in Bladder Cancer Cells. Frontiers in oncology 15 35774124