Affinage

BMAL1

Basic helix-loop-helix ARNT-like protein 1 · UniProt O00327

Length
626 aa
Mass
68.8 kDa
Annotated
2026-06-09
100 papers in source corpus 49 papers cited in narrative 49 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

BMAL1 is the obligate positive-arm transcription factor of the mammalian circadian clock, functioning as a pioneer-like factor that binds nucleosomes, promotes rhythmic chromatin opening with H2A.Z incorporation, and enables time-of-day binding of other transcription factors to generate genome-wide rhythmic output (PMID:24395244). Its rhythmic transcription is itself sustained by a REV-ERB/ROR (RRE) element feedback loop that confers resistance to perturbation of the core E-box oscillation (PMID:35999195), while promoter CpG methylation can silence BMAL1 expression entirely (PMID:24103761). BMAL1 activity and abundance are tightly tuned by post-translational control: GSK3β phosphorylates Ser17/Thr21 to prime ubiquitin-proteasomal degradation downstream of dopamine D2 receptor signaling (PMID:20049328), S6K1 phosphorylation of Ser42 drives promoter dissociation and an EZH2-dependent repressive chromatin switch (PMID:35338256), synaptic Ser42 phosphorylation gates CaMKIIα-dependent hippocampal LTP (PMID:37878694), and BMAL1 protein is stabilized by a necdin–SGT1–HSP90 chaperone module (PMID:32667666). As a transcription factor BMAL1 directly drives diverse tissue programs, activating metabolic and drug-handling targets including Dgat2, Cyp3a11, Sult1a1, and an intestinal Dbp/Rev-erbα→Mrp2 cascade (PMID:34493722, PMID:31633069, PMID:30064996, PMID:31244920), and controlling cell-cycle, survival, and differentiation genes such as CDK1/CCNA1 in endothelial cells, MITF in melanocytes, and hedgehog effectors Ptch1/Ihh in chondrocytes (PMID:37052172, PMID:34160901, PMID:31747713). Beyond canonical CLOCK partnership, BMAL1 forms a non-canonical, diurnally regulated heterodimer with HIF2A—whose structure was resolved by cryo-EM—to stabilize HIF2A and drive the rhythmic hypoxic target AREG (PMID:40269168), and it acts upstream of HIF1α in hepatic and skeletal-muscle hypoxia and glucose-metabolism programs (PMID:39106859, PMID:40127275). BMAL1 also performs transcription-independent functions: it stabilizes heterochromatin at LINE1 loci to suppress the cGAS-STING pathway during aging (PMID:35286396) and sequesters MYH9 to modulate MRTF-SRF cytoskeletal signaling and melanoma cell state (PMID:38245503). These activities underlie cell-type-specific roles spanning macrophage mitochondrial metabolism and RhoA-dependent phagocytosis (PMID:32396064, PMID:31900362), astrocyte BAG3-mediated protein clearance and endolysosomal function (PMID:37315555, PMID:37155839), skeletal-muscle control of sleep amount (PMID:28726633), oligodendrocyte precursor proliferation and myelination (PMID:37657440), and dopaminergic neuron survival (PMID:38032732).

Mechanistic history

Synthesis pass · year-by-year structured walk · 15 steps
  1. 2010 High

    Established that BMAL1 abundance is controlled by signal-driven phosphodegradation, linking neurotransmitter signaling to clock amplitude.

    Evidence In vitro kinase and ubiquitylation assays with Ser17/Thr21 mutagenesis and D2 receptor pharmacology in striatal neurons

    PMID:20049328

    Open questions at the time
    • E3 ligase mediating the ubiquitylation not defined here
    • in vivo phosphorylation stoichiometry over the circadian cycle not measured
  2. 2014 High

    Defined how a single heterodimer generates heterogeneous genome-wide rhythms: CLOCK:BMAL1 acts as a pioneer-like factor remodeling chromatin to license other factors.

    Evidence Genome-wide ChIP-seq, ATAC-seq, and histone variant (H2A.Z) profiling

    PMID:24395244

    Open questions at the time
    • chromatin remodelers recruited by BMAL1 not identified
    • which secondary factors are licensed at each locus left open
  3. 2016 High

    Connected hepatic BMAL1 to systemic lipid and cholesterol handling through defined intermediary transcription factors.

    Evidence Global and liver-specific Bmal1 KO on ApoE-/- and Ldlr-/- backgrounds with SHP/GATA4 gain-of-function rescue

    PMID:27721414

    Open questions at the time
    • direct vs indirect regulation of Shp/Gata4 not fully resolved
    • human relevance not tested
  4. 2018 High

    Showed BMAL1 transcriptional output is gated by tissue-specific repressors and reciprocal feedback in liver and colon oscillators.

    Evidence Reciprocal luciferase assays, genome-wide HNF4A/CLOCK:BMAL1 co-occupancy ChIP-seq, and HNF4A knockdown

    PMID:30530698

    Open questions at the time
    • mechanism of HNF4A transrepression at chromatin not defined
    • generalizability beyond liver/colon untested
  5. 2018 Medium

    Demonstrated that direct BMAL1 transcriptional targets extend to xenobiotic-handling enzymes, establishing a clock basis for drug pharmacokinetics.

    Evidence EMSA, ChIP, and luciferase reporters at the Sult1a1 E-box in Bmal1-deficient mice and serum-shocked cells

    PMID:30064996

    Open questions at the time
    • single-site analysis; broader enzyme network not mapped
    • human SULT1A1 regulation not addressed
  6. 2019 High

    Mapped a multi-tier BMAL1 transcriptional cascade controlling intestinal drug transport and CYP enzyme rhythms.

    Evidence Intestine-specific Bmal1 KO with EMSA/ChIP/reporter and in vivo pharmacokinetics for Mrp2 and Cyp3a11

    PMID:31244920 PMID:31633069

    Open questions at the time
    • relative contribution of each intermediary (Dbp, Rev-erba, Hnf4a) in vivo not quantified
  7. 2020 High

    Identified a chaperone machinery that maintains BMAL1 protein stability, defining a post-translational input to clock pace independent of transcription.

    Evidence Yeast two-hybrid, Co-IP with domain mapping, HSP90/proteasome inhibitor rescue, and necdin-KO mouse phenotyping

    PMID:32667666

    Open questions at the time
    • how chaperone availability is itself regulated rhythmically not addressed
  8. 2020 High

    Revealed BMAL1 as a metabolic and cytoskeletal regulator in macrophages, linking the clock to mitochondrial function, HIF-1α reprogramming, and RhoA-dependent phagocytosis.

    Evidence Myeloid-specific Bmal1 KO with metabolic readouts, phosphoproteomics, RhoA activity assays, ChIP-seq, and pharmacological rescue

    PMID:31900362 PMID:32396064

    Open questions at the time
    • whether cytoskeletal control is transcriptional or post-translational in macrophages not fully separated
  9. 2022 High

    Established a transcription-independent genome-protective role for BMAL1 in heterochromatin maintenance at LINE1 elements during aging.

    Evidence BMAL1 loss-of-function in primate MPCs with chromatin fractionation, LINE1 ChIP, and cGAS-STING pathway assays

    PMID:35286396

    Open questions at the time
    • how BMAL1 is recruited to LINE1 heterochromatin not defined
    • relationship to its E-box DNA binding unresolved
  10. 2022 High

    Showed that rhythmic Bmal1 transcription via RRE elements buffers the clock against perturbation, defining the functional purpose of the auxiliary feedback loop.

    Evidence CRISPR deletion of RRE elements in cells and mice with mathematical modeling and CRY1 manipulation

    PMID:35999195

    Open questions at the time
    • molecular nature of the perturbation buffering not mechanistically dissected
  11. 2022 High

    Defined an S6K1-Ser42 phosphorylation switch that couples nutrient/fasting signaling to a BMAL1→EZH2 chromatin state change at metabolic gene promoters.

    Evidence S6K1 kinase assay, phospho-Ser42 detection, ChIP for BMAL1/EZH2/H3K27me3 at Adipoq, and S6K1 KO mice

    PMID:35338256

    Open questions at the time
    • whether the EZH2 switch operates at other BMAL1 targets genome-wide untested
  12. 2023 High

    Connected BMAL1 to neuronal physiology through synaptic localization and cell-autonomous roles in protein clearance, axon regeneration, and synaptic maturation.

    Evidence Astrocyte-, neuron-, and Purkinje-cell-specific Bmal1 KO with synaptic fractionation, electrophysiology, BAG3/lysosomal assays, and 5hmC ChIP

    PMID:35301425 PMID:37155839 PMID:37315555 PMID:37620297 PMID:37878694

    Open questions at the time
    • extent to which these roles are circadian vs constitutive not always separated
    • direct BMAL1 target genes in each neural context only partially mapped
  13. 2024 High

    Positioned BMAL1 upstream of HIF1α in tissue hypoxia and glucose-metabolism programs through epistatic rescue.

    Evidence Liver and skeletal-muscle conditional Bmal1 KO with genetic HIF1α stabilization rescue, metabolomics, and time-dependent physiology

    PMID:39106859 PMID:40127275

    Open questions at the time
    • molecular mechanism by which BMAL1 restrains HIF1α accumulation not defined
  14. 2025 High

    Resolved a non-canonical BMAL1-HIF2A heterodimer structurally and functionally, demonstrating direct circadian-hypoxia transcriptional cross-talk.

    Evidence Cryo-EM of the BMAL1-HIF2A-DNA complex with Co-IP, HIF2A stability assays, ChIP, and cardiac ischemia models

    PMID:40269168

    Open questions at the time
    • genome-wide scope of BMAL1-HIF2A targets beyond AREG not mapped
    • competition with CLOCK for BMAL1 not quantified
  15. 2025 High

    Demonstrated that a small molecule binding the BMAL1 PASB cavity can allosterically reshape its transcription factor activity, validating BMAL1 as a directly druggable target.

    Evidence Small-molecule design with structural and biochemical binding studies, PER2-Luc oscillation readouts, and macrophage assays

    PMID:40133642

    Open questions at the time
    • selectivity over related PAS-domain proteins in vivo not fully established
    • therapeutic window unaddressed

Open questions

Synthesis pass · forward-looking unresolved questions
  • How BMAL1 partitions between its canonical CLOCK-dependent transcriptional role and its transcription-independent functions (heterochromatin stabilization, MYH9 sequestration, HIF partnerships) within a single cell remains unresolved.
  • no quantitative model of how BMAL1 pools are allocated among functions
  • recruitment mechanisms to non-E-box sites undefined
  • in vivo consequences of decoupling transcriptional from non-transcriptional roles untested

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140110 transcription regulator activity 10 GO:0003677 DNA binding 3 GO:0140313 molecular sequestering activity 1
Localization
GO:0005634 nucleus 3 GO:0000228 nuclear chromosome 2 GO:0005886 plasma membrane 1
Pathway
R-HSA-74160 Gene expression (Transcription) 5 R-HSA-1430728 Metabolism 3 R-HSA-4839726 Chromatin organization 3 R-HSA-8953897 Cellular responses to stimuli 3 R-HSA-9909396 Circadian clock 3 R-HSA-168256 Immune System 2
Partners
CLOCKHIF2AHNF4AMYH9NDNRELASGT1
Complex memberships
BMAL1-HIF2A heterodimerCLOCK:BMAL1 heterodimer

Evidence

Reading pass · 49 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2010 GSK3β phosphorylates BMAL1 specifically on Ser17 and Thr21, priming it for ubiquitylation and proteasomal degradation. In the absence of GSK3β-mediated phosphorylation, BMAL1 is stabilized. Dopamine D2 receptor signaling via the Akt-GSK3β pathway regulates BMAL1 stability and circadian gene expression amplitude in striatal neurons. In vitro phosphorylation assays, site-directed mutagenesis (Ser17/Thr21), ubiquitylation assays, D2 receptor pharmacology in striatal neurons PloS one High 20049328
2014 CLOCK:BMAL1 heterodimer functions as a pioneer-like transcription factor: it binds to nucleosomes, promotes rhythmic chromatin opening (including incorporation of histone variant H2A.Z), and enables rhythmic binding of other transcription factors at adjacent sites, thereby driving genome-wide heterogeneous transcriptional output. ChIP-seq, ATAC-seq (chromatin accessibility), histone variant profiling, genome-wide binding analysis Genes & development High 24395244
2020 BMAL1 functions as a metabolic checkpoint in macrophages: upon M1 inflammatory stimulation, myeloid-specific Bmal1 KO renders macrophages unable to sustain mitochondrial function, enhancing succinate dehydrogenase (SDH)-mediated ROS production and HIF-1α-dependent metabolic reprogramming. This Bmal1-HIF-1α regulatory loop controls macrophage effector functions and tumor microenvironment. Myeloid-specific Bmal1 knockout (M-BKO), mitochondrial function assays, ROS measurement, HIF-1α activation assays, SDH inhibitor rescue experiments, tumor burden measurement eLife High 32396064
2025 BMAL1 forms a transcriptionally active heterodimer with HIF2A (a non-canonical partner) in a diurnal manner to regulate myocardial hypoxic response. Cryo-EM structure of the BMAL1-HIF2A-DNA complex reveals structural rearrangements within BMAL1 that enable circadian-hypoxia cross-talk. BMAL1 enhances HIF2A transcriptional activity and stabilizes the HIF2A protein. Amphiregulin (AREG) is identified as a rhythmic target of this complex, critical for daytime variations in myocardial injury. Cryo-EM structure determination, Co-IP, transcriptional activity assays, HIF2A protein stability assays, ChIP, cardiac ischemia models with time-of-day pharmacological targeting Nature High 40269168
2018 The nuclear receptor HNF4A strongly transrepresses the transcriptional activity of the CLOCK:BMAL1 heterodimer and maintains cell-autonomous circadian oscillations in a tissue-specific manner in liver and colon cells. Genome-wide ChIP-seq reveals co-occupancy of HNF4A and CLOCK:BMAL1 at metabolic genes, defining a feedback loop in tissue-specific oscillators. Luciferase transcriptional activity assays, ChIP-seq (genome-wide co-occupancy), siRNA knockdown of HNF4A in liver and colon cells, circadian period measurement Proceedings of the National Academy of Sciences of the United States of America High 30530698
2022 BMAL1 plays a transcription-independent role in stabilizing heterochromatin to prevent activation of the LINE1-cGAS-STING pathway. In senescent primate mesenchymal progenitor cells (MPCs), BMAL1 binding to LINE1 loci is decreased, leading to heterochromatin destabilization and aberrant LINE1 transcription, accelerating aging phenotype. BMAL1 loss-of-function in human and cynomolgus monkey MPCs, chromatin fractionation, ChIP for BMAL1 at LINE1, cGAS-STING pathway activation assays, heterochromatin marker analysis in BMAL1-deficient monkey tissues Nucleic acids research High 35286396
2015 BMAL1 in chondrocytes directs circadian expression of catabolic, anabolic, and apoptotic genes critical for cartilage homeostasis. Loss of chondrocyte BMAL1 reduces phosphorylated SMAD2/3 and NFATC2, decreases expression of Sox9, Acan, and Col2a1, and increases p-SMAD1/5 levels, causing progressive cartilage degeneration. Chondrocyte-specific Bmal1 knockout mice, RNA-seq, western blot for SMAD2/3 and SMAD1/5 phosphorylation, histological cartilage assessment The Journal of clinical investigation High 26657859
2020 BMAL1 regulates macrophage actin cytoskeletal organization, motility, and phagocytosis via a RhoA-dependent mechanism. BMAL1-/- macrophages show reduced phosphocofilin, increased active RhoA, altered cell morphology and increased motility. BMAL1 controls a network of 148 cell movement genes within 100 kb of high-confidence BMAL1 binding sites, including 29 genes impacting RhoA expression or activation. RhoA inhibition restores phagocytic phenotype. Bmal1-/- macrophage characterization, phosphoproteomics, RhoA activity assay, phagocytosis assay in vivo and in vitro, RhoA inhibitor rescue, BMAL1 ChIP-seq for binding sites near cytoskeletal genes Proceedings of the National Academy of Sciences of the United States of America High 31900362
2023 BMAL1 rhythmically localizes to hippocampal synapses in a manner dependent on its phosphorylation at Ser42 (pBMAL1-S42). Synaptic pBMAL1(S42) regulates autophosphorylation of CaMKIIα and circadian rhythms of CaMKIIα-dependent molecular interactions and LTP, but not global rest/activity behavior. Immunofluorescence/subcellular fractionation of synaptic BMAL1, phospho-specific antibodies, electrophysiology (LTP), CaMKIIα autophosphorylation assays, Ser42 phosphorylation-state analysis Science advances High 37878694
2018 P2-HNF4α (an alternative isoform induced in hepatocellular carcinoma) represses BMAL1 (ARNTL) expression and causes nuclear-to-cytoplasmic re-localization of P1-HNF4α. Forced BMAL1 expression in HNF4α-positive HCC prevents tumor growth in vivo. Promoter reporter assays, immunofluorescence for BMAL1 localization, forced BMAL1 expression in HCC xenografts, isoform-specific expression analysis Nature communications Medium 30341289
2020 Necdin stabilizes BMAL1 protein through SGT1-HSP90 chaperone machinery. Necdin interacts with BMAL1 (N-terminal fragment) and SGT1 (C-terminal fragment) as identified by yeast two-hybrid screening. Depletion of necdin or SGT1/HSP90 leads to BMAL1 degradation via the ubiquitin-proteasome system, altering clock gene expression and circadian rhythms. Yeast two-hybrid screening, Co-IP, domain mapping (N- and C-terminal fragments), HSP90 inhibitor treatment, proteasome inhibitor rescue, necdin-KO mouse behavioral and gene expression analysis Nucleic acids research High 32667666
2014 BMAL1 deficiency leads to increased mTORC1 activity both in vivo and in cell culture. mTORC1 inhibition with rapamycin increases lifespan of Bmal1-/- mice by 50%, placing BMAL1 as a negative regulator of mTORC1 signaling upstream of aging control. Bmal1-/- mice, mTORC1 activity assays (S6K phosphorylation), rapamycin treatment, lifespan measurement Aging Medium 24481314
2022 The RRE-mediated feedback regulation of Bmal1 (rhythmic transcription driven by REV-ERB/ROR elements) underpins the E-box-mediated core oscillation. Deletion of RRE elements in mutant mice produced apparently normal rhythms but made circadian period and amplitude more susceptible to disturbance of CRY1 protein rhythm, demonstrating that rhythmic Bmal1 transcription confers perturbation resistance to the clock. CRISPR deletion of RRE elements in cells and mice, circadian period/amplitude measurements, mathematical modeling, CRY1 protein manipulation Nature communications High 35999195
2019 BMAL1 coordinates circadian MRP2 expression in the intestine by directly activating transcription of Dbp and Rev-erbα through E-box elements in their promoters, which in turn regulate Mrp2 via a D-box element. BMAL1 also negatively regulates E4BP4 (a Mrp2 repressor) via Rev-erbα, generating diurnal Mrp2 expression and time-dependent drug pharmacokinetics. Intestine-specific Bmal1 KO mice, qPCR, western blot, luciferase reporter assays, EMSA, ChIP assays for E-box binding, intestinal perfusion and pharmacokinetic experiments Theranostics High 31244920
2019 BMAL1 regulates circadian Cyp3a11 expression by directly activating Hnf4α and Dbp transcription via E-box elements. Cellular regulation of Cyp3a11 by BMAL1 is Dbp- and HNF4α-dependent, as Dbp activates and Hnf4α activates Cyp3a11 transcription via a D-box and DR1 element respectively in the Cyp3a11 promoter. Bmal1-deficient mice, luciferase reporter, ChIP, promoter mutation analysis, Dbp/Hnf4α overexpression/knockdown, drug toxicity assays Communications biology High 31633069
2018 BMAL1 activates Sult1a1 transcription by directly binding an E-box element (-571 to -554 bp) in the Sult1a1 promoter, controlling circadian expression and activity of the sulfotransferase enzyme. Bmal1-deficient mice, luciferase reporter assay, EMSA, ChIP assay, serum-shocked cell circadian model Drug metabolism and disposition Medium 30064996
2018 CLOCK and BMAL1 cooperate to upregulate RHOA expression by inhibiting CUL3-mediated ubiquitination, and activate RHOA by reducing its interaction with RhoGDI. Consequently, CLOCK:BMAL1 controls the RHOA-ROCK-cofilin pathway, altering F-actin/G-actin dynamics and promoting cancer cell proliferation, migration, and invasion. Co-IP for RHOA-RhoGDI and CUL3 interactions, ubiquitination assay, F-actin/G-actin ratio measurement, CLOCK/BMAL1 overexpression and knockdown in cancer cells, cell migration and invasion assays Experimental & molecular medicine Medium 30287810
2016 Hepatic BMAL1 controls lipoprotein production and biliary cholesterol excretion by regulating expression of Mtp and Abcg5/Abcg8 via Shp and Gata4. BMAL1 deficiency reduces SHP (increasing MTP and lipoprotein secretion) and reduces GATA4 (diminishing ABCG5/8 expression and biliary cholesterol excretion). Forced SHP or GATA4 expression rescues respective phenotypes. Global and liver-specific Bmal1 KO mice on ApoE-/- and Ldlr-/- backgrounds, forced expression of SHP and GATA4, lipoprotein secretion assays, biliary cholesterol measurement, western blot/qPCR Nature communications High 27721414
2021 BMAL1 directly transactivates the Dgat2 gene (encoding triacylglycerol synthesis enzyme DGAT2) via direct binding to an E-box in the promoter, promoting dietary fat absorption. Intestinal Bmal1 deficiency impairs lipid resynthesis and reduces fat secretion; Rev-erbα deficiency (enhancing BMAL1 activity) exacerbates this. Intestine-specific Bmal1 KO mice, ChIP assay for E-box binding in Dgat2 promoter, luciferase reporter, lipid absorption assays, Rev-erbα intestinal KO Nature communications High 34493722
2022 S6K1 phosphorylates BMAL1 at serine 42, causing its dissociation from the Adipoq promoter. This leads to EZH2 recruitment and H3K27me3 modification of the Adipoq promoter, suppressing adiponectin expression. Fasting inactivates S6K1, triggering an EZH2-to-BMAL1 transcriptional switch that promotes adiponectin expression. S6K1 kinase assay, ChIP for BMAL1 and EZH2/H3K27me3 at Adipoq promoter, S6K1 KO mice, phospho-Ser42 BMAL1 detection, promoter reporter assays Experimental & molecular medicine High 35338256
2023 Astrocyte-specific BMAL1 deletion induces Bag3 expression, a chaperone critical for macroautophagy, enhancing astrocyte phagocytosis of alpha-synuclein and tau in a BAG3-dependent manner. Astrocyte Bag3 overexpression is sufficient to mitigate alpha-synuclein spreading in vivo, placing BMAL1 upstream of BAG3-mediated protein clearance. Astrocyte-specific Bmal1 KO, Bag3 KO rescue experiments, Bag3 overexpression in vivo, phagocytosis assays for alpha-synuclein and tau, tauopathy and alpha-synucleinopathy mouse models Neuron High 37315555
2023 BMAL1 deletion in astrocytes influences endolysosomal function, autophagy, and protein degradation dynamics in a cell-autonomous manner. Bmal1-deficient astrocytes exhibit increased endocytosis, lysosome-dependent protein cleavage, and accumulation of LAMP1- and RAB7-positive organelles in vitro; astrocyte-specific Bmal1 KO brains show accumulation of autophagosome-like structures in vivo by electron microscopy. Transcriptional dysregulation of lysosome function pathways occurs independently of TFEB activation. Astrocyte-specific Bmal1 KO, electron microscopy, LAMP1/RAB7 immunostaining, endocytosis assays, lysosomal protease activity, RNA-seq of isolated astrocytes Proceedings of the National Academy of Sciences of the United States of America High 37155839
2018 BMAL1 cell-autonomously regulates astrocyte activation (astrogliosis) and inflammatory gene expression. Astrocyte-specific Bmal1 deletion induces astrocyte activation in vitro and in vivo, mediated in part by suppression of glutathione-S-transferase signaling. Loss of astrocyte BMAL1 promotes neuronal death in vitro. Astrocyte-specific Bmal1 KO (in vitro and in vivo), inflammatory gene expression profiling, glutathione-S-transferase pathway analysis, neuronal co-culture death assay Cell reports High 30282019
2017 BMAL1 expression in skeletal muscle is both necessary and sufficient to regulate total sleep amount. Restoring Bmal1 only in skeletal muscle of Bmal1-KO mice rescued sleep-amount phenotypes; muscle-specific Bmal1 KO reproduced them. Overexpression of skeletal-muscle Bmal1 reduced the recovery response to sleep loss. Brain-specific and muscle-specific Bmal1 rescue in global KO mice, muscle-specific Bmal1 KO, polysomnographic sleep recording eLife High 28726633
2023 Bmal1 inhibits axon regeneration of dorsal root ganglia neurons by acting as a gatekeeper of neuroepigenetic responses to axonal injury, limiting Tet3 expression and restricting 5-hydroxymethylcytosine (5hmC) modifications at regeneration-associated genes. Conditional neuronal Bmal1 deletion enhances axon regeneration in vitro and in vivo. Diurnal oscillation of Tet3 and 5hmC in DRG neurons corresponds to time-of-day effect on axon growth potential. Neuron-specific conditional Bmal1 KO, in vitro and in vivo neurite outgrowth assays, ChIP for 5hmC, RNA-seq, Tet3 expression analysis Nature communications High 37620297
2019 BMAL1 directly binds the MITF promoter and transcriptionally regulates its expression, which positively influences melanin synthesis. BMAL1 overexpression increases MITF and melanin levels in human melanocytes/melanoma cells and protects against UVB-induced DNA damage. ChIP for BMAL1 at MITF promoter, luciferase reporter assay, BMAL1 overexpression/knockdown, melanin quantification, UVB treatment Pigment cell & melanoma research Medium 34160901
2024 BMAL1 directly regulates neuronal survival in dopaminergic (TH+) neurons of the substantia nigra pars compacta in a cell-autonomous manner. Pan-neuronal or TH neuron-specific Bmal1 deletion caused spontaneous TH+ neuron loss. Transcriptomics revealed dysregulation of oxidative phosphorylation and Parkinson disease pathways. Cell-type-specific Cre-lox Bmal1 KO (pan-neuronal, TH-specific, astrocyte, microglia), stereological TH+ neuron counting, RNA-seq JCI insight High 38032732
2021 BMAL1 transcriptionally regulates Rab27a (a key exosome secretion gene) via luciferase assay, promoting exosome release from colorectal cancer cells and facilitating cancer cell migration. Luciferase reporter assay for Rab27a promoter, BMAL1 overexpression/knockdown, exosome isolation and quantification, migration assays Molecular biology reports Medium 34727291
2022 Polyamines stimulate BMAL1 and REV-ERBα protein synthesis at the translational level through enhancement of ribosomal shunting within the 5'-UTR of their mRNAs. In polyamine-reduced cells, BMAL1 synthesis is reduced and circadian period is lengthened. EGFP fusion reporter with wild-type or mutated 5'-UTR of Bmal1 mRNA, polyamine depletion experiments, circadian period measurement in NIH3T3 cells International journal of molecular sciences Medium 33525630
2019 BMAL1 (ARNTL) associates with telomere DNA in a rhythmic/diurnal fashion in zebrafish and mice, and controls a circadian rhythm in TERRA (Telomeric Repeat-containing RNA). BMAL1-dependent H3K9me3 rhythm at telomeres is lost in aged animals, linking BMAL1 to telomere heterochromatin maintenance. ChIP for BMAL1 at telomere DNA, TERRA RT-qPCR in Bmal1-/- mouse liver, H3K9me3 ChIP at telomeres in zebrafish and mice, aging comparison PloS one Medium 31634367
2021 BMAL1 dephosphorylation by protein phosphatase 4 (PPP4) participates in modulating circadian timing and determines the pace of the circadian clock (as reviewed based on findings by Klemz et al.). Biochemical/genetic analysis of PPP4-mediated BMAL1 dephosphorylation (referenced in review/perspective based on original research) Genes & development Low 34341001
2025 A small molecule (CCM) targeting the PASB domain cavity of BMAL1 causes expansion of the cavity, conformational changes in the PASB domain, and alters BMAL1 function as a transcription factor. CCM induces dose-dependent alterations in PER2-Luc oscillations and downregulates inflammatory and phagocytic pathways in macrophages. Crystal/structural and biochemical studies validate selectivity. Small molecule design, structural studies (BMAL1 PASB domain), biochemical binding assays, PER2-Luc reporter assay, macrophage inflammatory assays Nature chemical biology High 40133642
2020 BMAL1 directly interacts with NF-κB p65 (RelA) subunit in theca cells and negatively regulates PI3K/NF-κB signaling, thereby controlling steroidogenesis-associated gene expression including Lhcgr, Hsd3β2, and Cyp17a1. Co-IP for BMAL1-RelA interaction, PI3K inhibitor rescue, siRNA-mediated Bmal1 knockdown in granulosa and theca cells, Bmal1 KO mice, hormone measurement Reproduction Medium 33112769
2019 BMAL1 (ARNTL) acts as a transcription factor that suppresses NPC cell proliferation and enhances cisplatin sensitivity by directly binding the CDK5 promoter and activating CDK5 transcription, as validated by dual luciferase reporter and ChIP assays. CDK5 overexpression reverses the suppressive effects of BMAL1. Dual luciferase reporter assay for CDK5 promoter, ChIP assay, ARNTL overexpression and CDK5 overexpression epistasis experiment, proliferation and cisplatin sensitivity assays Journal of experimental & clinical cancer research Medium 30621723
2022 BMAL1 promotes OPC (oligodendrocyte precursor cell) proliferation in a time-of-day-dependent manner. OPC-specific Bmal1 KO disrupts circadian gene expression, proliferation, density, morphology, and migration, leading to thinner myelin and sleep fragmentation. In adults, OPC-specific Bmal1 loss impairs remyelination via changes in OPC morphology and migration. OPC-specific Bmal1 KO during development and adulthood, RNA-seq, immunostaining, electron microscopy for myelin thickness, demyelination lesion model, sleep recording Neuron High 37657440
2023 BMAL1 binds the promoters of CCNA1 and CDK1 genes in endothelial cells (by ChIP-seq) and controls their expression, thereby regulating EC cell cycle progression. EC-specific BMAL1 impairment causes angiogenesis defects in neonatal vascular tissues and adult tumor angiogenesis settings. EC-specific Bmal1 KO in vivo, BMAL1 ChIP-seq in ECs, cell cycle analysis, angiogenesis assays (neonatal retina and tumor), RNA-seq Cardiovascular research High 37052172
2024 Hepatic BMAL1 and HIF1α cooperate in the circadian response to hypoxia through shared and distinct regulatory roles. HIF1α accumulation upon hypoxia is temporally regulated and Bmal1-dependent. Loss of both hepatic Bmal1 and Hif1α causes hypoxemia, pulmonary vasodilation (via ERK-eNOS-NO), and mortality in a daytime-dependent manner, resembling hepatopulmonary syndrome. Liver-specific Bmal1 KO, liver-specific Hif1α KO, double KO, hypoxia exposure at different times of day, transcriptomics, eNOS/NO measurement, ERK activation assays Cell metabolism High 39106859
2018 LIN28A directly binds the BMAL1 promoter under H. pylori infection and activates BMAL1 transcription. In turn, BMAL1 functions as a transcription factor to enhance expression of proinflammatory TNF-α, promoting gastric inflammation. ChIP assay for LIN28A at BMAL1 promoter, dual luciferase reporter assay, H. pylori infection models in vitro and in vivo EBioMedicine Medium 30502053
2022 BMAL1 non-canonically sequesters myosin heavy chain 9 (MYH9) to increase MRTF-SRF activity and AP-1 transcriptional signature, shifting melanoma cells from a Sox10-high to Sox9-high mesenchymal/immune-resistant cell state. This occurs independently of BMAL1's transcriptional activity, as a transcriptionally inactive BMAL1 mutant retains this function. Co-IP for BMAL1-MYH9 interaction, BMAL1 transcriptionally inactive mutant expression, MRTF-SRF reporter assays, scRNA-seq cell state analysis, tumor growth assays Nature communications Medium 38245503
2022 BMAL1 in cerebellar Purkinje cells (PCs) is required cell-autonomously for normal synaptic transmission and dendritic spine maturation. Bmal1 KO in PCs causes enhanced excitatory and inhibitory synaptic transmission, reduced PC firing, aberrant spine density/morphology, hyperactivated mTORC1 signaling, and autistic-like behaviors. Metformin reversal of mTORC1 hyperactivation rescues behavioral and cellular deficits. Global and PC-specific Bmal1 KO, electrophysiology, dendritic spine morphology, mTORC1 activity assays, metformin rescue, behavioral tests Molecular psychiatry High 35301425
2025 In skeletal muscle during diet-induced obesity, BMAL1 regulates HIF-driven glycolysis and metabolic flexibility. Muscle-specific BMAL1 deficiency impairs early glycolytic steps and worsens glucose tolerance. Genetic HIF1α stabilization in muscle Bmal1-deficient mice restores glucose tolerance and rescues 217/736 dysregulated genes, establishing BMAL1 upstream of HIF1α in muscle glucose metabolism. Muscle-specific Bmal1 KO, metabolite profiling, glucose tolerance tests, HIF1α genetic stabilization rescue, RNA-seq in high-fat diet conditions Proceedings of the National Academy of Sciences of the United States of America High 40127275
2019 BMAL1 directly activates Ptch1 and Ihh promoters (binding to their promoter regions), regulating hedgehog signaling in mandibular condylar chondrocytes. BMAL1 deficiency impairs sequential chondrocyte differentiation and endochondral ossification; hedgehog signaling activator rescues the short stature phenotype. ChIP for BMAL1 at Ptch1 and Ihh promoters, RNA-seq in Bmal1-/- mandibular condyle, hedgehog activator rescue of BMAL1-deficient phenotype, micro-CT and histology Cell proliferation Medium 31747713
2022 PPAR-γ trans-activates Slc1a5 via a response element in its promoter, driving glutamine/methionine uptake in adipocytes. Impaired PPAR-γ in obesity reduces SLC1A5 and glutamine/methionine, decreasing H3K27ac and H3K4me3 at the Bmal1 promoter and disrupting Bmal1 transcription. ChIP assay for H3K27ac and H3K4me3 at Bmal1 promoter, luciferase reporter for PPAR-γ/SLC1A5 axis, in vitro and in vivo glutamine/methionine supplementation, qPCR/western blot Theranostics Medium 35198059
2013 DNA methylation of the BMAL1 promoter CpG islands silences circadian BMAL1 expression. Treatment with the demethylating agent 5-aza-2'-deoxycytidine recovers BMAL1 expression and circadian oscillation. Reporter gene assays confirm that transcriptional machinery for BMAL1 is active, and methylation (not promoter dysfunction) is the cause of silencing. Bisulfite sequencing of BMAL1 promoter CpG islands, 5-aza-dC treatment, luciferase reporter assay, qPCR for circadian oscillation recovery Biochemical and biophysical research communications Medium 24103761
2020 BMAL1 promotes mitophagy via SIRT1 to alleviate myocardial injury in sepsis. BMAL1 knockdown inhibits SIRT1 and mitophagy-associated proteins; SIRT1 overexpression partially reverses effects of BMAL1 knockdown, placing BMAL1 upstream of SIRT1 in a pro-mitophagy pathway in cardiomyocytes. siRNA/shRNA Bmal1 knockdown and overexpression in H9C2 cells, SIRT1 overexpression rescue, mitophagy protein expression (western blot), apoptosis/ROS by flow cytometry, cecal ligation-puncture in vivo model International immunopharmacology Medium 38678672
2024 BMAL1 overexpression reduces YAP transcriptional activity and inhibits the Hippo signaling pathway in renal tubular epithelial cells; ChIP-qPCR identifies YAP as a direct BMAL1 target. YAP positively regulates ACSL4, promoting ferroptosis. Inhibiting YAP (by Verteporfin) reverses BMAL1-downregulation-induced ferroptosis. BMAL1 overexpression, ChIP-qPCR for BMAL1 at YAP promoter, Verteporfin rescue, ferroptosis markers, transcriptome sequencing, in vivo sepsis-AKI model International immunopharmacology Medium 39303541
2021 BMAL1 (ARNTL) in EPIYA L-cells is required for rhythmic GLP-1 secretion; L-cell-specific Arntl KO reduces GLP-1 secretion at peak time points and impairs time-dependent GLP-1 release. L-cell Arntl disruption also alters intestinal CD4+ intraepithelial lymphocytes, proinflammatory cytokine expression, and colonic microbiome composition. Inducible L-cell-specific Arntl KO mice, oral glucose tolerance tests at ZT2 and ZT14, GLP-1 measurement, siRNA knockdown in mGLUTag cells, transcriptomic analysis, microbiome 16S rRNA sequencing Molecular metabolism Medium 34520858
2022 Circadian rhythm disruption (by melatonin receptor 1/AMPKβ1 pathway perturbation) impairs endochondral bone formation. Mechanistically, melatonin receptor 1 (MTR1) periodically activates AMPKβ1 phosphorylation, which destabilizes CRY1 and triggers BMAL1 expression, coordinating circadian rhythms of chondrocyte proliferation and matrix synthesis. Circadian disruption mouse models, AMPKβ1 agonist rescue, AMPKβ1 phosphorylation assays, CRY1 stability assays, BMAL1 expression analysis, bone formation histology Cell death and differentiation Medium 35094018
2024 Circadian disruption causes BMAL1 to suppress MUC4 expression in corneal epithelial cells, contributing to dry eye pathology. BMAL1 silencing reduces MUC4 expression and BMAL1 overexpression increases it in cultured human corneal epithelial cells in vitro. Genetic BMAL1 ablation in mice recapitulates MUC4 deficiency and dry eye disease. BMAL1 siRNA and overexpression in human corneal epithelial cells, Bmal1 KO mice, MUC4 protein/mRNA measurement, corneal surface assessment Experimental & molecular medicine Medium 38956298

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2010 Regulation of BMAL1 protein stability and circadian function by GSK3beta-mediated phosphorylation. PloS one 242 20049328
2014 CLOCK:BMAL1 is a pioneer-like transcription factor. Genes & development 206 24395244
2020 BMAL1-Downregulation Aggravates Porphyromonas Gingivalis-Induced Atherosclerosis by Encouraging Oxidative Stress. Circulation research 196 32078488
2014 BMAL1-dependent regulation of the mTOR signaling pathway delays aging. Aging 176 24481314
2015 The chondrocyte clock gene Bmal1 controls cartilage homeostasis and integrity. The Journal of clinical investigation 173 26657859
2020 Bmal1 integrates mitochondrial metabolism and macrophage activation. eLife 137 32396064
2018 Cell-Autonomous Regulation of Astrocyte Activation by the Circadian Clock Protein BMAL1. Cell reports 137 30282019
2017 Bmal1 function in skeletal muscle regulates sleep. eLife 123 28726633
2020 The clock gene Bmal1 inhibits macrophage motility, phagocytosis, and impairs defense against pneumonia. Proceedings of the National Academy of Sciences of the United States of America 116 31900362
2016 Global and hepatocyte-specific ablation of Bmal1 induces hyperlipidaemia and enhances atherosclerosis. Nature communications 104 27721414
2018 Incompatibility of the circadian protein BMAL1 and HNF4α in hepatocellular carcinoma. Nature communications 95 30341289
2022 PPAR-γ integrates obesity and adipocyte clock through epigenetic regulation of Bmal1. Theranostics 93 35198059
2010 ARNTL (BMAL1) and NPAS2 gene variants contribute to fertility and seasonality. PloS one 85 20368993
2023 An astrocyte BMAL1-BAG3 axis protects against alpha-synuclein and tau pathology. Neuron 82 37315555
2022 BMAL1 moonlighting as a gatekeeper for LINE1 repression and cellular senescence in primates. Nucleic acids research 80 35286396
2013 Impaired glucocorticoid production and response to stress in Arntl-deficient male mice. Endocrinology 77 24189141
2016 Dysregulated circadian rhythm pathway in human osteoarthritis: NR1D1 and BMAL1 suppression alters TGF-β signaling in chondrocytes. Osteoarthritis and cartilage 73 27884645
2021 Deficiency of intestinal Bmal1 prevents obesity induced by high-fat feeding. Nature communications 72 34493722
2016 Myeloid Bmal1 deletion increases monocyte recruitment and worsens atherosclerosis. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 71 27927724
2018 Nuclear receptor HNF4A transrepresses CLOCK:BMAL1 and modulates tissue-specific circadian networks. Proceedings of the National Academy of Sciences of the United States of America 70 30530698
2022 Rhythmic transcription of Bmal1 stabilizes the circadian timekeeping system in mammals. Nature communications 69 35999195
2019 The Circadian Clock Gene Bmal1 Controls Intestinal Exporter MRP2 and Drug Disposition. Theranostics 66 31244920
2020 Deficiency of the Circadian Clock Gene Bmal1 Reduces Microglial Immunometabolism. Frontiers in immunology 64 33363534
2019 Bmal1 regulates circadian expression of cytochrome P450 3a11 and drug metabolism in mice. Communications biology 62 31633069
2017 Cell-intrinsic, Bmal1-dependent Circadian Regulation of Temozolomide Sensitivity in Glioblastoma. Journal of biological rhythms 62 28470120
2018 Potential Roles of Dec and Bmal1 Genes in Interconnecting Circadian Clock and Energy Metabolism. International journal of molecular sciences 60 29518061
2014 Epigenetic silencing of ARNTL, a circadian gene and potential tumor suppressor in ovarian cancer. International journal of oncology 55 25175925
2018 Opposite Carcinogenic Effects of Circadian Clock Gene BMAL1. Scientific reports 54 30375470
2023 Circadian clock protein BMAL1 broadly influences autophagy and endolysosomal function in astrocytes. Proceedings of the National Academy of Sciences of the United States of America 52 37155839
2019 Aryl hydrocarbon receptor nuclear translocator-like (ARNTL/BMAL1) is associated with bevacizumab resistance in colorectal cancer via regulation of vascular endothelial growth factor A. EBioMedicine 52 31300350
2022 Emerging Insight Into the Role of Circadian Clock Gene BMAL1 in Cellular Senescence. Frontiers in endocrinology 49 35733785
2020 The biological function of BMAL1 in skeleton development and disorders. Life sciences 47 32251631
2022 Autistic-like behavior and cerebellar dysfunction in Bmal1 mutant mice ameliorated by mTORC1 inhibition. Molecular psychiatry 46 35301425
2019 Effects of BMAL1 Manipulation on the Brain's Master Circadian Clock and Behavior. The Yale journal of biology and medicine 45 31249486
2015 Association of CLOCK, ARNTL, and NPAS2 gene polymorphisms and seasonal variations in mood and behavior. Chronobiology international 43 26134245
2017 Conditional Deletion of Bmal1 Accentuates Microvascular and Macrovascular Injury. The American journal of pathology 42 28432873
2022 Circadian rhythm modulates endochondral bone formation via MTR1/AMPKβ1/BMAL1 signaling axis. Cell death and differentiation 40 35094018
2025 BMAL1-HIF2A heterodimer modulates circadian variations of myocardial injury. Nature 39 40269168
2019 ARNTL hypermethylation promotes tumorigenesis and inhibits cisplatin sensitivity by activating CDK5 transcription in nasopharyngeal carcinoma. Journal of experimental & clinical cancer research : CR 39 30621723
2023 BMAL1 loss in oligodendroglia contributes to abnormal myelination and sleep. Neuron 38 37657440
2023 The circadian protein BMAL1 supports endothelial cell cycle during angiogenesis. Cardiovascular research 36 37052172
2013 Common genetic variants in ARNTL and NPAS2 and at chromosome 12p13 are associated with objectively measured sleep traits in the elderly. Sleep 36 23449886
2018 CLOCK and BMAL1 stabilize and activate RHOA to promote F-actin formation in cancer cells. Experimental & molecular medicine 35 30287810
2020 Necdin regulates BMAL1 stability and circadian clock through SGT1-HSP90 chaperone machinery. Nucleic acids research 33 32667666
2019 Long noncoding RNA FLRL2 alleviated nonalcoholic fatty liver disease through Arntl-Sirt1 pathway. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 33 31311301
2024 Circadian misalignment impairs oligodendrocyte myelination via Bmal1 overexpression leading to anxiety and depression-like behaviors. Journal of pineal research 31 38241675
2019 Circadian BMAL1 regulates mandibular condyle development by hedgehog pathway. Cell proliferation 31 31747713
2018 The Circadian Oscillator of the Cerebral Cortex: Molecular, Biochemical and Behavioral Effects of Deleting the Arntl Clock Gene in Cortical Neurons. Cerebral cortex (New York, N.Y. : 1991) 31 28052921
2020 The circadian clock gene Bmal1 facilitates cisplatin-induced renal injury and hepatization. Cell death & disease 30 32522976
2017 Smad3 and Bmal1 regulate p21 and S100A4 expression in myocardial stromal fibroblasts via TNF-α. Histochemistry and cell biology 30 28721450
2023 Synaptic BMAL1 phosphorylation controls circadian hippocampal plasticity. Science advances 29 37878694
2023 Neural function of Bmal1: an overview. Cell & bioscience 28 36593479
2024 Hepatic BMAL1 and HIF1α regulate a time-dependent hypoxic response and prevent hepatopulmonary-like syndrome. Cell metabolism 27 39106859
2018 BMAL1 and CLOCK proteins in regulating UVB-induced apoptosis and DNA damage responses in human keratinocytes. Journal of cellular physiology 27 29943823
2018 The Clock Protein Bmal1 Regulates Circadian Expression and Activity of Sulfotransferase 1a1 in Mice. Drug metabolism and disposition: the biological fate of chemicals 27 30064996
2018 H. pylori infection induced BMAL1 expression and rhythm disorder aggravate gastric inflammation. EBioMedicine 27 30502053
2021 BMAL1 induces colorectal cancer metastasis by stimulating exosome secretion. Molecular biology reports 26 34727291
2025 Pharmacological targeting of BMAL1 modulates circadian and immune pathways. Nature chemical biology 25 40133642
2024 Neuronal deletion of the circadian clock gene Bmal1 induces cell-autonomous dopaminergic neurodegeneration. JCI insight 24 38032732
2020 The role of circadian clock gene BMAL1 in vascular proliferation. European journal of pharmacology 24 31958455
2022 Critical Roles of the Circadian Transcription Factor BMAL1 in Reproductive Endocrinology and Fertility. Frontiers in endocrinology 23 35311235
2022 Haploinsufficiency of a Circadian Clock Gene Bmal1 (Arntl or Mop3) Causes Brain-Wide mTOR Hyperactivation and Autism-like Behavioral Phenotypes in Mice. International journal of molecular sciences 22 35682995
2024 BMAL1 alleviates myocardial damage in sepsis by activating SIRT1 signaling and promoting mitochondrial autophagy. International immunopharmacology 21 38678672
2023 Behavioral circatidal rhythms require Bmal1 in Parhyale hawaiensis. Current biology : CB 21 36977416
2022 BMAL1 regulates Propionibacterium acnes-induced skin inflammation via REV-ERBα in mice. International journal of biological sciences 21 35414779
2021 The circadian clock gene Bmal1: Role in COVID-19 and periodontitis. Chronobiology international 21 33792447
2021 Orthodontic Force-Induced BMAL1 in PDLCs Is a Vital Osteoclastic Activator. Journal of dental research 21 34157911
2021 Circadian clock protein BMAL1 regulates melanogenesis through MITF in melanoma cells. Pigment cell & melanoma research 21 34160901
2018 Deficiency of the clock gene Bmal1 affects neural progenitor cell migration. Brain structure & function 21 30341743
2023 Ceramide-1-phosphate alleviates high-altitude pulmonary edema by stabilizing circadian ARNTL-mediated mitochondrial dynamics. Journal of advanced research 20 37479181
2019 BMAL1 associates with chromosome ends to control rhythms in TERRA and telomeric heterochromatin. PloS one 20 31634367
2024 Cell state dependent effects of Bmal1 on melanoma immunity and tumorigenicity. Nature communications 19 38245503
2022 Bmal1-knockout mice exhibit reduced cocaine-seeking behaviour and cognitive impairments. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 19 35779420
2022 Neuronal Bmal1 regulates retinal angiogenesis and neovascularization in mice. Communications biology 18 35933488
2019 Deficiency of Bmal1 disrupts the diurnal rhythm of haemostasis. Experimental gerontology 18 30610898
2013 DNA methylation of the BMAL1 promoter. Biochemical and biophysical research communications 18 24103761
2024 Circadian disruption reduces MUC4 expression via the clock molecule BMAL1 during dry eye development. Experimental & molecular medicine 17 38956298
2020 BMAL1 coordinates energy metabolism and differentiation of pluripotent stem cells. Life science alliance 17 32284354
2023 Circadian clock regulator Bmal1 gates axon regeneration via Tet3 epigenetics in mouse sensory neurons. Nature communications 16 37620297
2021 Translational Regulation of Clock Genes BMAL1 and REV-ERBα by Polyamines. International journal of molecular sciences 16 33525630
2021 L-cell Arntl is required for rhythmic glucagon-like peptide-1 secretion and maintenance of intestinal homeostasis. Molecular metabolism 16 34520858
2020 Circadian expression of Fabp7 mRNA is disrupted in Bmal1 KO mice. Molecular brain 16 32093736
2015 Depression-associated ARNTL and PER2 genetic variants in psychotic disorders. Chronobiology international 16 25799324
2025 Control of circadian muscle glucose metabolism through the BMAL1-HIF axis in obesity. Proceedings of the National Academy of Sciences of the United States of America 15 40127275
2023 BMAL1 regulates osteoblast differentiation through mTOR/GSK3β/β-catenin pathway. Journal of molecular endocrinology 15 36942818
2022 S6K1 controls adiponectin expression by inducing a transcriptional switch: BMAL1-to-EZH2. Experimental & molecular medicine 15 35338256
2020 The clock regulator Bmal1 protects against muscular dystrophy. Experimental cell research 15 33130178
2016 Circadian adaptation to cell injury stresses: a crucial interplay of BMAL1 and HSF1. The journal of physiological sciences : JPS 15 26910317
2025 Atheroma transcriptomics identifies ARNTL as a smooth muscle cell regulator and with clinical and genetic data improves risk stratification. European heart journal 14 39552248
2024 Melatonin ameliorates hepatic fibrosis via the melatonin receptor 2-mediated upregulation of BMAL1 and anti-oxidative enzymes. European journal of pharmacology 14 38246330
2024 BMAL1/PGC1α4-FNDC5/irisin axis impacts distinct outcomes of time-of-day resistance exercise. Journal of sport and health science 14 39187065
2023 Ziwuliuzhu Acupuncture Modulates Clock mRNA, Bmal1 mRNA and Melatonin in Insomnia Rats. Journal of acupuncture and meridian studies 14 37381033
2017 Circadian regulation of kidney function: finding a role for Bmal1. American journal of physiology. Renal physiology 14 29357439
2024 BMAL1 alleviates sepsis-induced AKI by inhibiting ferroptosis. International immunopharmacology 13 39303541
2023 Denervation aggravates renal ischemia reperfusion injury via BMAL1-mediated Nrf2/ARE pathway. Archives of biochemistry and biophysics 13 37657745
2022 Melatonin upregulates BMAL1 to attenuate chronic sleep deprivation-related cognitive impairment by alleviating oxidative stress. Brain and behavior 13 36563187
2021 BMAL1 dephosphorylation determines the pace of the circadian clock. Genes & development 13 34341001
2020 Core clock gene Bmal1 deprivation impairs steroidogenesis in mice luteinized follicle cells. Reproduction (Cambridge, England) 13 33112769
2022 The circadian clock gene ARNTL overexpression suppresses oral cancer progression by inducing apoptosis via activating autophagy. Medical oncology (Northwood, London, England) 12 36180647
2023 The circadian rhythm gene Bmal1 ameliorates acute deoxynivalenol-induced liver damage. Archives of toxicology 11 36602574

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