Affinage

DRAM1

DNA damage-regulated autophagy modulator protein 1 · UniProt Q8N682

Length
238 aa
Mass
26.3 kDa
Annotated
2026-06-09
98 papers in source corpus 31 papers cited in narrative 31 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 7/7 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

DRAM1 is a multi-pass lysosomal transmembrane protein and a direct transcriptional target of p53 that couples stress sensing to autophagy and regulated cell death (PMID:16839881, PMID:17397945). Its core lysosomal function is to drive autophagic flux: DRAM1 enhances lysosomal acidification by promoting V-ATPase activity and assembly, activates lysosomal proteases including cathepsin D, and accelerates autophagosome clearance (PMID:23696801, PMID:32943616). At the membrane-fusion step, DRAM1 binds VAMP8 on lysosomes and competitively blocks CHIP/STUB1-mediated ubiquitination of VAMP8 at Lys68/72/75, stabilizing VAMP8 and promoting assembly of the STX17-SNAP29-VAMP8 SNARE complex to license autophagosome-lysosome fusion (PMID:40595569, PMID:40884094). Beyond bulk autophagy, DRAM1 acts as a lysosomal scaffold that binds amino acid transporters SLC1A5 and LAT1 together with the carrier SCAMP3, directing them to lysosomes to support amino acid efflux and mTORC1 activation in an autophagy-independent manner with consequences for insulin signaling and adipocyte differentiation (PMID:31492633). DRAM1 also governs cell death decisions: it interacts directly with BAX, protects it from autophagic degradation, and recruits it to lysosomes to trigger cathepsin B release, BID cleavage, and mitochondrial apoptosis (PMID:25633293). In innate immunity, DRAM1 is induced downstream of TLR/IL1R-MYD88-NF-κB signaling and drives selective autophagy (xenophagy), LC3-associated phagocytosis, and acidification of pathogen-containing vesicles against intracellular bacteria, requiring STING and p62 (PMID:24922577, PMID:34731006, PMID:40950712). Additional lysosome-organelle activities include EPS15-dependent EGFR endocytic degradation (PMID:32943616) and STIM1-mediated ER-lysosome tethering that regulates calcium homeostasis and ER-phagy, with lysosomal targeting dependent on the intact cytosol-facing C-terminal domain (PMID:39292746).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 2006 High

    Established DRAM1 as the molecular link between p53 and autophagy, answering how the tumor suppressor engages the lysosomal degradation system to execute cell death.

    Evidence RNAi knockdown, overexpression, lysosomal fractionation and p53-epistasis assays in mammalian cells

    PMID:16839881

    Open questions at the time
    • Did not define the molecular activity of the lysosomal protein
    • Mechanism connecting DRAM1 induction to apoptosis unresolved
  2. 2007 High

    Defined the transcriptional control architecture, showing direct p53 binding at an upstream response element and a six-transmembrane topology, while extending regulation to the p53 family member p73.

    Evidence ChIP, luciferase reporter, isogenic p53 cell lines, GFP-fusion confocal microscopy; p73 isoform overexpression

    PMID:17304243 PMID:17397945

    Open questions at the time
    • p73-induced autophagy is DRAM-independent, leaving DRAM's role in p73 signaling unclear
    • No structural validation of the six-TM model
  3. 2009 Medium

    Distinguished DRAM1 from its paralogue DRAM2 and showed upstream JNK signaling feeds DRAM induction, refining the regulatory inputs and establishing functional specificity within the family.

    Evidence Paralogue localization/function comparison; siRNA of DRAM and JNK with autophagy/apoptosis readouts

    PMID:19556885 PMID:19706754

    Open questions at the time
    • Why DRAM2 localizes to lysosomes yet does not modulate autophagy unexplained
    • Direct biochemical activity still undefined
  4. 2012 High

    Resolved the lysosomal mechanism by which DRAM1 promotes flux—enhancing V-ATPase-driven acidification, cathepsin D activation, and autophagosome clearance—and linked DRAM1 to p62 trafficking and cancer cell invasion.

    Evidence siRNA knockdown, RFP-LC3 flux, lysosomal pH and cathepsin assays; p62 localization and invasion assays in glioblastoma stem cells

    PMID:22082963 PMID:22525272 PMID:23696801

    Open questions at the time
    • How DRAM1 mechanistically promotes V-ATPase activity not yet shown
    • Splice-variant-specific functions only partially mapped
  5. 2013 High

    Identified DRAM1 as a direct BAX partner and a node in viral-stress-induced death, establishing a transcription-independent apoptotic mechanism via lysosomal recruitment of BAX and cathepsin B release.

    Evidence Co-IP, BAX half-life and cathepsin B/cytochrome c assays; HIV infection model with p53/DRAM knockdown

    PMID:23658518 PMID:25633293

    Open questions at the time
    • Structural basis of DRAM1-BAX interaction unknown
    • How lysosomal BAX recruitment is gated relative to autophagy not defined
  6. 2014 High

    Connected DRAM1 to innate immune defense, showing a p53-independent MYD88-NF-κB axis induces DRAM1 to drive xenophagy against mycobacteria, and revealed AKT-dependent control of DRAM mitochondrial translocation.

    Evidence MYD88/NF-κB/STING/p62 epistasis in zebrafish and human macrophages; p-AKT/DRAM co-IP and localization in HCC cells

    PMID:24556693 PMID:24922577

    Open questions at the time
    • Molecular basis of STING/p62 requirement for DRAM1 xenophagy not defined
    • How p-AKT binding sequesters DRAM in cytoplasm structurally unclear
  7. 2018 Medium

    Expanded DRAM1's autophagy machinery contacts, showing it binds Atg7 to promote Atg12-Atg5 conjugation and is required for alternative Atg5-independent autophagy and isolation-membrane closure downstream of p53.

    Evidence Co-IP with Atg7/Atg5/Atg12, mRFP-GFP-LC3 flux, EM of isolation membranes, Atg5/p53 epistasis

    PMID:30144448 PMID:31225467

    Open questions at the time
    • Reconciliation of Atg7-dependent and Atg5-independent roles unresolved
    • Direct biochemical effect of DRAM1 on conjugation machinery not reconstituted
  8. 2019 High

    Revealed an autophagy-independent metabolic function: DRAM1 scaffolds amino acid transporters and SCAMP3 at lysosomes to drive amino acid efflux and mTORC1 activation, with whole-body consequences for glycemic and adipocyte biology, while also influencing Golgi/secretory trafficking and IGF-1R signaling.

    Evidence Co-IP, lysosomal fractionation, amino acid efflux and mTORC1 assays, DRAM1 knockout mouse; VSVG and CI-MPR trafficking assays

    PMID:30902093 PMID:31356858 PMID:31492633

    Open questions at the time
    • Mechanism by which DRAM1 selects transporter cargo for lysosomal delivery unknown
    • Apparent opposing effects on mTORC1/PI3K-Akt across contexts not reconciled
  9. 2020 High

    Defined DRAM1 as a recruiter of V-ATPase and EPS15 to drive EGFR endocytic degradation and confirmed in vivo that vesicle acidification underlies pathogen clearance, with loss leading to premature pyroptosis.

    Evidence BioID proteomics, EPS15 co-IP, lysosomal pH and EGFR degradation, xenografts; zebrafish dram1 CRISPR with caspa/gsdmeb epistasis

    PMID:32332700 PMID:32943616

    Open questions at the time
    • How DRAM1 physically enhances V-ATPase assembly remains mechanistically open
    • Link between failed acidification and pyroptosis induction not defined
  10. 2021 Medium

    Showed DRAM1 binds STOM and PKM2 to control lysosomal membrane permeabilization and extracellular vesicle/ectosome secretion from hepatocytes, extending its role to lipid- and ethanol-driven liver pathology and immune crosstalk.

    Evidence Co-IP, DRAM knockout and liver-specific overexpression mice, LMP and EV/ectosome assays, macrophage activation

    PMID:34731006 PMID:35036051

    Open questions at the time
    • Whether STOM and PKM2 interactions are direct or scaffold-mediated unclear
    • Mechanism coupling LMP to vesicle secretion not resolved
  11. 2024 High

    Established DRAM1 as an ER-lysosome contact factor: it tethers lysosomes to the ER via STIM1, controlling calcium homeostasis, ER stress, and ER-phagy, and mapped lysosomal targeting to its cytosol-facing C-terminal domain.

    Evidence Co-IP, calcium imaging, ER morphology and ER-phagy assays, AMPK-ULK1 readouts, domain-deletion mutants, STIM1 rescue

    PMID:39292746

    Open questions at the time
    • Physiological versus overexpression-driven STIM1 effects need separation
    • Structural determinants of the DRAM1-STIM1 contact undefined
  12. 2025 High

    Defined the molecular mechanism of DRAM1 at the fusion step—competitive inhibition of CHIP/STUB1 ubiquitination of VAMP8 to stabilize it and promote STX17-SNAP29-VAMP8 SNARE assembly—and tied this to HCC metastasis, while also implicating DRAM1 in AMPK-driven pro-senescent autophagy.

    Evidence Co-IP, VAMP8 lysine mutagenesis, ubiquitination and SNARE assembly assays, ATG5/ATG7 KO, mouse and zebrafish metastasis models; MSC/hepatocyte AMPK and senescence assays

    PMID:40595569 PMID:40884094 PMID:41037659

    Open questions at the time
    • How DRAM1 competes with CHIP at VAMP8 structurally unresolved
    • Relationship between SNARE stabilization and the distinct p62-sparing senescent autophagy unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • The biochemical activity intrinsic to the DRAM1 transmembrane protein—how it mechanistically promotes V-ATPase assembly and selects diverse cargo (transporters, BAX, VAMP8, STIM1) at the lysosomal membrane—remains undefined, as does a structural model unifying its many context-dependent partnerships.
  • No structure of DRAM1 or its complexes
  • No reconstituted assay defining a primary molecular activity
  • Determinants of cargo/partner selectivity unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 5 GO:0098772 molecular function regulator activity 4
Localization
GO:0005764 lysosome 7 GO:0005768 endosome 2 GO:0005783 endoplasmic reticulum 2 GO:0005886 plasma membrane 2 GO:0005794 Golgi apparatus 1
Pathway
R-HSA-9612973 Autophagy 5 R-HSA-162582 Signal Transduction 4 R-HSA-168256 Immune System 4 R-HSA-5357801 Programmed Cell Death 3 R-HSA-5653656 Vesicle-mediated transport 3
Partners
BAXEPS15LAT1SCAMP3SLC1A5STIM1STOMVAMP8

Evidence

Reading pass · 31 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2006 DRAM1 (DRAM) is a direct transcriptional target of p53 that encodes a lysosomal protein; p53 induces autophagy in a DRAM-dependent manner, and DRAM is essential for p53-mediated apoptosis. RNAi knockdown, overexpression, lysosomal localization by cell fractionation/imaging, p53-dependent induction assays Cell High 16839881
2007 TA-p73 transcriptionally regulates DRAM but, unlike p53, p73's induction of autophagy is DRAM-independent; deltaN-p73 negatively regulates p53-induced and p73-induced autophagy but not starvation-induced autophagy. RNAi knockdown of DRAM, overexpression of p73 isoforms, autophagy flux assays Cell death and differentiation Medium 17304243
2007 FLJ11259/DRAM1 is a direct p53 target gene with a functional p53 response element 22.3 kb upstream of the first coding exon; p53 binds this element in vivo (ChIP), and binding is enhanced after cisplatin treatment. The protein contains six transmembrane domains and localizes in a punctate cytoplasmic pattern. p53 siRNA knockdown, isogenic p53+/+ vs p53-suppressed cell lines, luciferase reporter assay, chromatin immunoprecipitation, GFP-fusion confocal microscopy Biochimica et biophysica acta High 17397945
2009 DRAM1 is a lysosomal protein; a closely related paralogue DRAM2 (45% identity) also localizes to lysosomes but does not modulate autophagy on overexpression and is not induced by p53 or p73. Drosophila DmDRAM, the single fly orthologue, retains the ability to modulate autophagy. Overexpression with subcellular localization (immunofluorescence), autophagy flux assays, comparison of p53/p73 induction across paralogues Cell cycle (Georgetown, Tex.) Medium 19556885
2009 JNK activation is required upstream of DRAM induction by 2-methoxyestradiol; JNK promotes DRAM expression in a p53-partially-regulated manner, and DRAM silencing reduces both autophagy and apoptosis triggered by 2-ME. siRNA knockdown of DRAM and JNK, pharmacological JNK inhibition, autophagy/apoptosis assays Cancer research Medium 19706754
2012 DRAM1 promotes autophagy flux by enhancing lysosomal acidification (V-ATPase activity), promoting autophagosome-lysosome fusion, and activating lysosomal cathepsin D; siRNA knockdown of DRAM1 reduces lysosomal V-ATPase activity and slows clearance of autophagosomes. siRNA knockdown, RFP-LC3 flux assay, lysosomal pH measurement, cathepsin D activity assay, rapamycin washout experiment PloS one High 23696801
2012 DRAM1 regulates p62 localization to autophagosomes and its autophagy-mediated degradation; DRAM1 knockdown decreases this process. DRAM1 and p62 cooperatively regulate cell motility and invasion in glioblastoma stem cells, associated with reduced ATP and lactate levels. Starvation- or mTOR/PI3K inhibition-induced autophagy is not affected by DRAM1 or p62 knockdown. siRNA knockdown, immunofluorescence localization of p62, invasion/migration assays, metabolic measurements Oncogene Medium 22525272
2012 DRAM-1 encodes multiple p53-inducible splice variants (SV1, SV4, SV5); SV1 localizes to lysosomes/endosomes, SV4 partially localizes to peroxisomes and ER, SV5 partially localizes to autophagosomes and ER. SV4 and SV5 modulate autophagy without inducing programmed cell death. Cloning of splice variants, immunofluorescence co-localization with organelle markers, autophagy assays Autophagy Medium 22082963
2013 DRAM1 interacts directly with BAX protein, inhibiting BAX degradation by autophagy, thereby increasing BAX protein levels in a transcription-independent manner. DRAM1 recruits BAX to lysosomes, triggering lysosomal cathepsin B release and BID cleavage, leading to mitochondrial cytochrome c release and caspase-3 activation. Co-immunoprecipitation, immunofluorescence, siRNA knockdown, BAX half-life measurement, cathepsin B/cytochrome c release assays Cell death & disease High 25633293
2013 HIV infection induces DRAM expression in a p53-dependent manner in CD4+ T cells; DRAM knockdown inhibits autophagy and lysosomal membrane permeabilization (LMP), cytochrome C release, MOMP, and cell death, but increases viral production. siRNA knockdown of DRAM and p53, LMP assay, cytochrome C release, MOMP measurement, viral titer PLoS pathogens Medium 23658518
2014 DRAM1 expression is induced downstream of the TLR/IL1R-MYD88-NF-κB innate immune sensing pathway in response to mycobacterial infection; DRAM1 activates selective autophagy against mycobacteria in a p53-independent manner requiring STING and p62/SQSTM1. MYD88/NF-κB inhibition, siRNA knockdown of DRAM1, overexpression, co-localization with Mtb, zebrafish and human macrophage infection models Cell host & microbe High 24922577
2014 Phosphorylated AKT (p-AKT) binds DRAM in the cytoplasm, blocking DRAM translocation to mitochondria; inactivation of PI3K/AKT pathway rescues DRAM translocation to mitochondria, where DRAM induces mitophagy and apoptosis in hepatocellular carcinoma cells. Co-immunoprecipitation of p-AKT and DRAM, immunofluorescence tracking of DRAM localization, PI3K inhibition, siRNA knockdown Cell death & disease Medium 24556693
2015 DRAM-3 (TMEM150B), a DRAM1-related protein, localizes to lysosomes/autolysosomes, endosomes, and plasma membrane; it modulates autophagy flux and promotes cell survival under glucose deprivation in an autophagy-independent manner. Immunofluorescence localization, CRISPR/Cas9 disruption, autophagy flux assays, clonogenic survival assay Cell death and differentiation Medium 25929859
2018 Dram1 is required for genotoxic stress-induced alternative (Atg5-independent) autophagy; Dram1 functions in the closure of isolation membranes downstream of p53 in this alternative autophagy pathway. Dram1 overexpression/knockdown in cells with or without Atg5, electron microscopy of isolation membranes, epistasis with p53 Cell stress Medium 31225467
2018 DRAM1 interacts with Atg7 (but not directly with Atg5 or Atg12), promoting formation of the Atg12-Atg5 conjugate; DRAM1 overexpression restores autophagic flux and autophagosome-to-autophagolysosome conversion in ischemic cardiomyocytes. Atg7 siRNA abolishes these effects. Co-immunoprecipitation (DRAM1 with Atg7, Atg5, Atg12), mRFP-GFP-LC3 flux assay, siRNA knockdown, adenoviral overexpression in rat AMI model Journal of molecular and cellular cardiology Medium 30144448
2019 DRAM1 binds the membrane carrier protein SCAMP3 and amino acid transporters SLC1A5 and LAT1, directing them to lysosomes to permit efficient efflux of amino acids from lysosomes into the cytoplasm, which is required for mTORC1 activation. Loss of DRAM1 impairs mTORC1 activation, insulin signaling, glycemic balance, and adipocyte differentiation. This effect is autophagy-independent. Co-immunoprecipitation, lysosomal fractionation, amino acid efflux assays, DRAM-1 knockout mouse, mTORC1 activity measurements, adipocyte differentiation assay Molecular cell High 31492633
2019 DRAM1 inhibits rpS6 phosphorylation in an mTORC1-dependent manner and inhibits activation of the PI3K-Akt pathway stimulated by growth factors; DRAM1 localizes at the plasma membrane and regulates phosphorylation of the IGF-1 receptor. FLAG-DRAM1 overexpression, siRNA knockdown, Western blot for phospho-rpS6/Akt/IGF-1R, immunostaining for DRAM1 localization, CCK-8 and colony formation assays Cell communication and signaling : CCS Medium 30902093
2019 DRAM1 deficiency causes fragmentation of the Golgi apparatus; DRAM1 is partially localized in the Golgi, and its knockdown delays ER-to-plasma membrane transport of VSVG-GFP and impedes retrograde trafficking of CI-MPR from plasma membrane to Golgi. siRNA knockdown, immunofluorescence with Golgi markers, ts045-VSVG-GFP transport assay, CI-MPR trafficking assay Cellular signalling Medium 31356858
2020 Dram1 deficiency in zebrafish reduces phagosome/vesicle acidification of Mycobacterium marinum-containing vesicles, impairs autophagic targeting of Mm, and leads to premature pyroptotic death of infected macrophages via caspase a/gasdermin Eb; knockdown of caspa and gsdmeb reverts the increased bacterial burden. Zebrafish dram1 CRISPR mutant, in vivo imaging of autophagic targeting, LysoTracker acidification assay, caspa/gsdmeb knockdown epistasis, RNA-seq Cell death & disease High 32332700
2020 DRAM1 interacts with EPS15 to promote EGFR endocytosis, and recruits V-ATPase (V-ATP6V1 subunit) to lysosomes, increasing lysosomal V-ATPase assembly, lowering lysosomal pH, and activating lysosomal proteases, resulting in accelerated EGFR lysosomal degradation and suppression of EGFR signaling. Proximity labeling (BioID) followed by proteomics, co-IP of DRAM1 with EPS15, lysosomal pH measurement, EGFR degradation assay, xenograft tumor model Cell death & disease High 32943616
2021 DRAM interacts with stomatin (STOM) and promotes its lysosomal localization; fatty acid-induced DRAM enhances lysosomal membrane permeabilization (LMP) and promotes exosome secretion from hepatocytes. DRAM knockout reverses high-fat diet-induced increase in exosome secretion. Co-immunoprecipitation of DRAM and STOM, DRAM knockout mouse model, siRNA knockdown, LMP assay, exosome quantification Science advances Medium 34731006
2021 DRAM1 interacts with PKM2 and increases PKM2 levels at the plasma membrane; ethanol-induced DRAM1 in hepatocytes increases secretion of PKM2-enriched extracellular vesicles/ectosomes that promote macrophage M1 activation. Co-immunoprecipitation of DRAM1 and PKM2, DRAM1 knockout and liver-specific overexpression mouse models, ectosome isolation, macrophage activation assay Molecular therapy. Nucleic acids Medium 35036051
2024 DRAM1 interacts with STIM1 to tether lysosomes to the ER, perturbing STIM1 function in maintaining intracellular calcium homeostasis; excess DRAM1 disrupts ER structure, triggers ER stress, and induces protective ER-phagy. Lysosomal localization of DRAM1 requires its intact cytosol-facing C-terminal domain. STIM1 overexpression restores calcium homeostasis, ER stress response, and AMPK-ULK1 signaling in cells with excess DRAM1. Co-immunoprecipitation of DRAM1 and STIM1, calcium imaging, ER morphology analysis, ER-phagy assay, AMPK-ULK1 signaling readouts, domain-deletion mutants for localization Proceedings of the National Academy of Sciences of the United States of America High 39292746
2023 In macrophages, DRAM1 localizes to mycobacteria-containing vesicles post-phagocytosis; DRAM1 knockdown reduces LC3 recruitment to mycobacteria and acidification of mycobacteria-containing vesicles, and reduces fusion with LAMP1-positive lysosomes, impairing intracellular killing. DRAM1 siRNA knockdown in RAW264.7 macrophages, immunofluorescence co-localization with LC3/LysoTracker/LAMP1, bacterial survival assay Cells Medium 36980169
2024 DRAM1 and xenophagy receptors Optn and p62 independently promote host defense against Mycobacterium marinum; Dram1 overexpression can compensate for loss of Optn or p62, and vice versa. Dram1 overexpression restores Lc3-Mm interaction in optn/p62 double mutants, indicating Dram1-mediated defense does not rely on specific xenophagy receptors. Single and double knockout zebrafish lines, overexpression rescue experiments, Lc3-Mm co-localization imaging, bacterial burden quantification Frontiers in cellular and infection microbiology Medium 38264729
2025 DRAM1 directly binds VAMP8 on lysosomes; this interaction is enhanced upon autophagy induction. DRAM1 competitively inhibits CHIP/STUB1-mediated ubiquitination of VAMP8 at Lys68, Lys72, and Lys75, stabilizing lysosomal VAMP8 and promoting assembly of the STX17-SNAP29-VAMP8 SNARE complex, thereby facilitating autophagosome-lysosome fusion and autophagic flux. DRAM1-mediated VAMP8 stabilization promotes HCC cell extravasation. Co-immunoprecipitation of DRAM1-VAMP8 and STUB1-VAMP8, ubiquitination assay, site-directed mutagenesis of VAMP8 lysines, SNARE complex assembly assay, ATG5/ATG7 knockout, mouse and zebrafish metastasis models Nature communications High 40595569
2025 DRAM1 activates AMPK and mediates pro-senescent autophagy (DMPA) in response to aging-associated metabolic cues (N-acetylhistamine, phosphatidylethanolamine); DRAM1-mediated autophagy in this context does not notably degrade SQSTM1/p62, distinguishing it from general macroautophagy. DRAM1 overexpression/knockdown in human MSCs and mouse hepatocytes, metabolomics, AMPK activity assay, autophagy flux assay, senescence markers, N-AcHA and ethanolamine supplementation Autophagy Medium 41037659
2023 Dram1/DRAM1 promotes LC3-associated phagocytosis (LAP) of Salmonella Typhimurium; Dram1 knockdown or mutation reduces GFP-Lc3 association with Salmonella and abolishes phagosomal NADPH oxidase-dependent ROS response to the bacteria. These results were confirmed in zebrafish and murine RAW264.7 macrophages. Morpholino knockdown, CRISPR mutation, mRNA overexpression in zebrafish, Salmonella ROS biosensor, GFP-Lc3 imaging, siRNA in RAW264.7 macrophages Autophagy reports Medium 40950712
2012 p73-regulated DRAM-1 is functionally required for neutrophil differentiation of APL cells; DRAM-1 expression is induced during ATRA-induced differentiation of NB4 APL cells, and its knockdown impairs this differentiation. p73 inhibition prevents both differentiation and DRAM-1 induction. siRNA knockdown of DRAM-1, ATRA differentiation assay, p73 inhibition, DRAM-1 expression measurement by qPCR/Western blot Leukemia research Medium 22981223
2012 Serum deprivation induces DRAM expression in liver cancer cells through epigenetic remodeling: active chromatin marks (diacetyl-H3, tetra-acetyl-H4, trimethyl-H3K4) increase and repressive mark (dimethyl-H3K9) decreases at the DRAM core promoter. The chromatin remodeling factor Brg-1 is enriched at the DRAM promoter and is required for serum deprivation-induced DRAM expression. ChIP for histone marks and Brg-1 at DRAM promoter, Brg-1 knockdown, luciferase reporter for DRAM promoter activity PloS one Medium 23251372
2025 DRAM1 interacts with VAMP8; this interaction is enhanced by autophagy stimulation. DRAM1 preferentially promotes autophagosome-lysosome fusion by enhancing assembly of the STX17-SNAP29-VAMP8 SNARE complex, and inhibits VAMP8 ubiquitination at Lys68, Lys72, and Lys75 by competing with CHIP/STUB1. Co-immunoprecipitation, ubiquitination assay, mutagenesis of VAMP8 lysine residues, SNARE complex co-IP, autophagy flux assay Autophagy High 40884094

Source papers

Stage 0 corpus · 98 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 DRAM, a p53-induced modulator of autophagy, is critical for apoptosis. Cell 1115 16839881
2012 The autophagy-associated factors DRAM1 and p62 regulate cell migration and invasion in glioblastoma stem cells. Oncogene 223 22525272
2007 DRAM links autophagy to p53 and programmed cell death. Autophagy 178 17102582
2012 Meta-analysis followed by replication identifies loci in or near CDKN1B, TET3, CD80, DRAM1, and ARID5B as associated with systemic lupus erythematosus in Asians. American journal of human genetics 175 23273568
2014 The DNA damage-regulated autophagy modulator DRAM1 links mycobacterial recognition via TLR-MYD88 to autophagic defense [corrected]. Cell host & microbe 133 24922577
2010 Evidence for the interplay between JNK and p53-DRAM signalling pathways in the regulation of autophagy. Autophagy 131 19949306
2015 DRAM1 regulates apoptosis through increasing protein levels and lysosomal localization of BAX. Cell death & disease 83 25633293
2007 p73 regulates DRAM-independent autophagy that does not contribute to programmed cell death. Cell death and differentiation 82 17304243
2013 DRAM1 regulates autophagy flux through lysosomes. PloS one 72 23696801
2009 c-Jun NH2-terminal kinase activation is essential for DRAM-dependent induction of autophagy and apoptosis in 2-methoxyestradiol-treated Ewing sarcoma cells. Cancer research 72 19706754
2009 Analysis of DRAM-related proteins reveals evolutionarily conserved and divergent roles in the control of autophagy. Cell cycle (Georgetown, Tex.) 62 19556885
2013 DRAM triggers lysosomal membrane permeabilization and cell death in CD4(+) T cells infected with HIV. PLoS pathogens 60 23658518
2012 Autophagy is induced through the ROS-TP53-DRAM1 pathway in response to mitochondrial protein synthesis inhibition. Autophagy 56 22576012
2012 DRAM-1 encodes multiple isoforms that regulate autophagy. Autophagy 53 22082963
2019 DRAM1 regulates autophagy and cell proliferation via inhibition of the phosphoinositide 3-kinase-Akt-mTOR-ribosomal protein S6 pathway. Cell communication and signaling : CCS 49 30902093
2019 mTORC1 Activation Requires DRAM-1 by Facilitating Lysosomal Amino Acid Efflux. Molecular cell 46 31492633
2019 TP53, TP53 Target Genes (DRAM, TIGAR), and Autophagy. Advances in experimental medicine and biology 45 31776983
2018 Dram1 regulates DNA damage-induced alternative autophagy. Cell stress 43 31225467
2016 Radiation induces autophagic cell death via the p53/DRAM signaling pathway in breast cancer cells. Oncology reports 41 27109777
2018 Overexpression of apoptosis-inducing factor mitochondrion-associated 1 (AIFM1) induces apoptosis by promoting the transcription of caspase3 and DRAM in hepatoma cells. Biochemical and biophysical research communications 38 29501488
2020 FTSJ1 regulates tRNA 2'-O-methyladenosine modification and suppresses the malignancy of NSCLC via inhibiting DRAM1 expression. Cell death & disease 37 32393790
2014 Phosphorylated AKT inhibits the apoptosis induced by DRAM-mediated mitophagy in hepatocellular carcinoma by preventing the translocation of DRAM to mitochondria. Cell death & disease 37 24556693
2021 Lipid-induced DRAM recruits STOM to lysosomes and induces LMP to promote exosome release from hepatocytes in NAFLD. Science advances 36 34731006
2020 Deficiency in the autophagy modulator Dram1 exacerbates pyroptotic cell death of Mycobacteria-infected macrophages. Cell death & disease 36 32332700
2013 Depending on the stage of hepatosteatosis, p53 causes apoptosis primarily through either DRAM-induced autophagy or BAX. Liver international : official journal of the International Association for the Study of the Liver 36 23875779
2018 Increased expression of DRAM1 confers myocardial protection against ischemia via restoring autophagy flux. Journal of molecular and cellular cardiology 33 30144448
2020 Tanshinone I induces cell apoptosis by reactive oxygen species-mediated endoplasmic reticulum stress and by suppressing p53/DRAM-mediated autophagy in human hepatocellular carcinoma. Artificial cells, nanomedicine, and biotechnology 32 32013613
2020 DRAM1 plays a tumor suppressor role in NSCLC cells by promoting lysosomal degradation of EGFR. Cell death & disease 32 32943616
2017 MicroRNA-26b suppresses autophagy in breast cancer cells by targeting DRAM1 mRNA, and is downregulated by irradiation. Oncology letters 32 29399189
2018 DRAM1 regulates the migration and invasion of hepatoblastoma cells via autophagy-EMT pathway. Oncology letters 29 30013633
2017 p53-Dependent PUMA to DRAM antagonistic interplay as a key molecular switch in cell-fate decision in normal/high glucose conditions. Journal of experimental & clinical cancer research : CR 29 28893313
2020 Role of DRAM1 in mitophagy contributes to preeclampsia regulation in mice. Molecular medicine reports 28 32582984
2014 DRAM1 protects neuroblastoma cells from oxygen-glucose deprivation/reperfusion-induced injury via autophagy. International journal of molecular sciences 28 25342320
2019 miR-199a-5p Represses Protective Autophagy and Overcomes Chemoresistance by Directly Targeting DRAM1 in Acute Myeloid Leukemia. Journal of oncology 26 31636666
2007 The direct p53 target gene, FLJ11259/DRAM, is a member of a novel family of transmembrane proteins. Biochimica et biophysica acta 26 17397945
2018 Clinical significance of serum DRAM1 mRNA, ARSA mRNA, hsa-miR-2053 and lncRNA-RP1-86D1.3 axis expression in malignant pleural mesothelioma. Journal of cellular biochemistry 25 30362153
2015 DRAM-3 modulates autophagy and promotes cell survival in the absence of glucose. Cell death and differentiation 25 25929859
2020 Long Non-Coding RNA LINC00511 Accelerates Proliferation and Invasion in Cervical Cancer Through Targeting miR-324-5p/DRAM1 Axis. OncoTargets and therapy 24 33116605
1988 Rapid detection of cytomegalovirus by fluorescent monoclonal antibody staining and in situ DNA hybridization in a dram vial cell culture system. Journal of clinical microbiology 24 2838514
2016 Another DRAM involved in autophagy and cell death. Autophagy 23 27046253
2013 Overexpression of DRAM enhances p53-dependent apoptosis. Cancer medicine 23 24133622
2020 Autophagy-mediated cytoplasmic accumulation of p53 leads to apoptosis through DRAM-BAX in cadmium-exposed human proximal tubular cells. Biochemical and biophysical research communications 22 33321290
2020 DNA Damage-Regulated Autophagy Modulator 1 (DRAM1) Mediates Autophagy and Apoptosis of Intestinal Epithelial Cells in Inflammatory Bowel Disease. Digestive diseases and sciences 21 33184797
2014 DRAM1 promotes the targeting of mycobacteria to selective autophagy. Autophagy 21 25484076
2018 Effect of autophagy gene DRAM on proliferation, cell cycle, apoptosis, and autophagy of osteoblast in osteoporosis rats. Journal of cellular physiology 19 30203495
2021 DRAM1 increases the secretion of PKM2-enriched EVs from hepatocytes to promote macrophage activation and disease progression in ALD. Molecular therapy. Nucleic acids 18 35036051
2012 Inhibition of damage-regulated autophagy modulator-1 (DRAM-1) impairs neutrophil differentiation of NB4 APL cells. Leukemia research 18 22981223
2015 XingNaoJing, prescription of traditional Chinese medicine, prevents autophagy in experimental stroke by repressing p53-DRAM pathway. BMC complementary and alternative medicine 16 26481508
2022 A review on morphotropic phase boundary in fluorite-structure hafnia towards DRAM technology. Nano convergence 15 36182997
2024 Investigating the Association between the Autophagy Markers LC3B, SQSTM1/p62, and DRAM and Autophagy-Related Genes in Glioma. International journal of molecular sciences 11 38203743
2017 DRAM Is Involved in Regulating Nucleoside Analog-Induced Neuronal Autophagy in a p53-Independent Manner. Molecular neurobiology 11 28265856
2024 Xenophagy receptors Optn and p62 and autophagy modulator Dram1 independently promote the zebrafish host defense against Mycobacterium marinum. Frontiers in cellular and infection microbiology 10 38264729
2024 The p53 target DRAM1 modulates calcium homeostasis and ER stress by promoting contact between lysosomes and the ER through STIM1. Proceedings of the National Academy of Sciences of the United States of America 10 39292746
2024 High performance Si-MoS2 heterogeneous embedded DRAM. Nature communications 10 39532875
2023 kb_DRAM: annotation and metabolic profiling of genomes with DRAM in KBase. Bioinformatics (Oxford, England) 10 36857575
2012 Serum starvation induces DRAM expression in liver cancer cells via histone modifications within its promoter locus. PloS one 10 23251372
2024 Oxide and 2D TMD semiconductors for 3D DRAM cell transistors. Nanoscale horizons 9 38563255
2019 DRAM is Involved in Hypoxia/Ischemia-Induced Autophagic Apoptosis in Hepatocytes. Aging and disease 9 30705770
2025 DRAM1 promotes the stability of lysosomal VAMP8 to enhance autolysosome formation and facilitates the extravasation. Nature communications 8 40595569
2023 DRAM1 Promotes Lysosomal Delivery of Mycobacterium marinum in Macrophages. Cells 8 36980169
2022 EGFR-AS1 Promotes Nonsmall Cell Lung Cancer (NSCLC) Progression via Downregulating the miR-524-5p/DRAM1 Axis and Inhibiting Autophagic Lysosomal Degradation. Journal of oncology 8 35222643
2019 DRAM1 deficiency affects the organization and function of the Golgi apparatus. Cellular signalling 8 31356858
2023 ARHGAP4 promotes leukemogenesis in acute myeloid leukemia by inhibiting DRAM1 signaling. Oncogene 7 37443303
2025 DRAM1 enhances the proliferation and metastasis of gastric cancer through the PI3K/AKT/mTOR signaling pathway and energy metabolism. Scientific reports 6 39875526
2022 DRAM-4 and DRAM-5 are compensatory regulators of autophagy and cell survival in nutrient-deprived conditions. The FEBS journal 6 35060334
2025 3D stacked IGZO 2T0C DRAM array with multibit capability for computing in memory applications. Science advances 5 40408494
2020 Analysis of the Sensing Margin of Silicon and Poly-Si 1T-DRAM. Micromachines 5 32102235
2017 DRAM-3 modulates autophagy and promotes cell survival in the absence of glucose. Cell death and differentiation 4 28665403
2024 Genome-wide association study identifies DRAM1 associated with Tourette syndrome in Taiwan. Biomedical journal 3 38608873
2023 Disturbance Characteristics of 1T DRAM Arrays Consisting of Feedback Field-Effect Transistors. Micromachines 3 37374723
2022 3-D stacked polycrystalline-silicon-MOSFET-based capacitorless DRAM with superior immunity to grain-boundary's influence. Scientific reports 3 36002621
2025 Flexible Organic Field-Effect Transistor (OFET) Based 2T0C DRAM Cells with 2-Bit Operation and Extended Retention. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 2 40051390
2024 ITZO-Based Self-Aligned Top Gate Thin-Film Transistor with Minimum Parasitic Capacitance for Long-Retention 2T0C DRAM. ACS omega 2 39829566
2022 DRAM1 plays a tumor suppressor role in clear cell renal cell carcinoma through modulating Akt signaling. Acta histochemica 2 35299128
2022 Simulation Study: The Impact of Structural Variations on the Characteristics of a Buried-Channel-Array Transistor (BCAT) in DRAM. Micromachines 2 36144099
2022 Design of a Capacitorless DRAM Based on a Polycrystalline-Silicon Dual-Gate MOSFET with a Fin-Shaped Structure. Nanomaterials (Basel, Switzerland) 2 36234653
2020 Correction: FTSJ1 regulates tRNA 2'-O-methyladenosine modification and suppresses the malignancy of NSCLC via inhibiting DRAM1 expression. Cell death & disease 2 32488007
2020 [MiR-199a-5p Affects Sensitivity of Acute Myeloid Leukemia to Adriamycin by Targeting DRAM1]. Zhongguo shi yan xue ye xue za zhi 2 32798383
2020 Analysis of a Lateral Grain Boundary for Reducing Performance Variations in Poly-Si 1T-DRAM. Micromachines 2 33105643
2025 Aging-rewired metabolic cues promote autophagy and senescence via DRAM1. Autophagy 1 41037659
2024 Enhanced Oxidation Resistance and Interface Stability of Atomic-Layer-Deposited MoN Electrodes via TiN Passivation for DRAM Cell Capacitor Applications. ACS applied materials & interfaces 1 39402816
2024 Doping- and capacitor-less 1T-DRAM cell using reconfigurable feedback mechanism. Nanotechnology 1 39514903
2023 Simulation of Capacitorless DRAM Based on the Polycrystalline Silicon Nanotube Structure with Multiple Grain Boundaries. Nanomaterials (Basel, Switzerland) 1 37446542
2022 Pseudo-Static Gain Cell of Embedded DRAM for Processing-in-Memory in Intelligent IoT Sensor Nodes. Sensors (Basel, Switzerland) 1 35684905
2017 The Programming Optimization of Capacitorless 1T DRAM Based on the Dual-Gate TFET. Nanoscale research letters 1 28875269
2026 Atomic-Layer-Deposition of Sn-Incorporated MoO2 Films as an Interface Control Layer for High-Performances TiO2-Based DRAM Capacitors. ACS applied materials & interfaces 0 41645779
2026 Androgen receptor-dependent DRAM1 activation drives autophagic resistance to BRAF inhibitors in BRAFV600-mutant melanoma. Cell death & disease 0 41839860
2026 Geometrical Design Schemes for the 2T0C DRAM Cell Using a Vertical Stack Transistor with a Distinctive Single IGZO Active Layer. ACS applied materials & interfaces 0 41927497
2026 Uniform TiN-Capped Co Buried Bit Lines for 4F2 DRAM Vertical Channel Transistors: Mitigating Agglomeration with Co Nitridation. ACS omega 0 42040442
2026 Unlocking the performance limits of 2T0C DRAM with Ω-shaped-gated single-crystal In2O3 FETs. Science advances 0 42213832
2025 Design of a gate-all-around arch-shaped tunnel-field-effect-transistor-based capacitorless DRAM. Discover nano 0 40172787
2025 VAMP8 stabilization by DRAM1 enables autophagosome-lysosome fusion and promotes metastatic extravasation. Autophagy 0 40884094
2025 DNA damage-regulated autophagy modulator 1 (DRAM1)-induced lipophagy facilitates Toxoplasma gondii nutrient acquisition and infection. Microbiological research 0 41435631
2023 DRAM1 confers resistance to Salmonella infection. Autophagy reports 0 40950712
2022 A Novel Capacitorless 1T DRAM with Embedded Oxide Layer. Micromachines 0 36296125
2021 Circuit Optimization Method to Reduce Disturbances in Poly-Si 1T-DRAM. Micromachines 0 34683260
2021 OBET: On-the-Fly Byte-Level Error Tracking for Correcting and Detecting Faults in Unreliable DRAM Systems. Sensors (Basel, Switzerland) 0 34960359
2016 DRAM Weak Cell Characterization for Retention Time. Journal of nanoscience and nanotechnology 0 27483878

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