Affinage

TMUB1

Transmembrane and ubiquitin-like domain-containing protein 1 · UniProt Q9BVT8

Length
246 aa
Mass
26.3 kDa
Annotated
2026-04-28
100 papers in source corpus 5 papers cited in narrative 5 extracted findings

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

TMUB1 (also known as HOPS) is a ubiquitin-like domain-containing shuttling protein that regulates protein stability, localization, and turnover across multiple cellular contexts. It retains cytoplasmic p53 by binding it and competing with importin-α for nuclear import, thereby promoting p53 mitochondrial recruitment and intrinsic apoptosis; Hops−/− mice show reduced apoptosis upon chemotherapeutic challenge (PMID:31867855). TMUB1 stabilizes PD-L1 in the endoplasmic reticulum by competing with the E3 ligase HUWE1 to block polyubiquitination at K281 and by recruiting the oligosaccharyltransferase STT3A to enhance N-glycosylation, thereby promoting PD-L1 maturation and tumor immune evasion (PMID:36376293). Additional functions include facilitating synaptic surface recycling of GluR2-containing AMPA receptors through interaction with GluR2 and GRIP (PMID:18665261), and forming a trimeric complex with nucleophosmin (NPM) and p19(Arf) that stabilizes p19(Arf), controls its nucleolar localization, and prevents centrosome hyperamplification (PMID:22890319).

Mechanistic history

Synthesis pass · year-by-year structured walk · 5 steps
  1. 2005 Medium

    Identification of TMUB1 as a nucleocytoplasmic shuttling protein that binds eEF-1A and suppresses proliferation established it as a functionally active signaling intermediary rather than a passive structural component.

    Evidence Regenerating liver cDNA screen; co-IP with eEF-1A; CRM-1 export and cAMP regulation assays; overexpression growth suppression in H-35 and 3T3-NIH cells

    PMID:16014383

    Open questions at the time
    • Mechanism by which eEF-1A binding suppresses translation not delineated
    • Findings from a single lab without independent replication
    • In vivo relevance of the proliferation-suppressive effect not tested
  2. 2008 Medium

    Demonstrating that TMUB1 selectively promotes surface recycling of GluR2-containing AMPA receptors expanded its role to post-synaptic membrane trafficking, answering how GluR2 surface levels are maintained at synapses.

    Evidence RNAi knockdown in neurons with electrophysiology; surface biotinylation; co-IP of TMUB1 with GluR2 and GRIP from mouse brain; recycling assay

    PMID:18665261

    Open questions at the time
    • Single-lab study without independent replication
    • Structural basis of the TMUB1–GluR2–GRIP interaction unknown
    • Whether TMUB1 functions in long-term synaptic plasticity not tested
  3. 2012 Medium

    Discovery of a TMUB1–NPM–p19(Arf) trimeric complex revealed how TMUB1 integrates nucleolar surveillance with cell cycle control, explaining centrosome hyperamplification upon TMUB1 loss.

    Evidence Co-IP; protein stability assays; siRNA knockdown with centrosome and cell cycle phenotypes; overexpression inducing G0/G1 arrest

    PMID:22890319

    Open questions at the time
    • Single-lab study; trimeric complex not reconstituted in vitro
    • Whether TMUB1 loss causes centrosome defects through p19(Arf)-dependent or -independent pathways not resolved
    • Relevance of the complex to tumorigenesis in vivo not established
  4. 2019 High

    Using knockout mice, the p53-retention mechanism was resolved: TMUB1 competes with importin-α for p53 binding, blocking nuclear import and channeling p53 to mitochondria to activate intrinsic apoptosis—the first in vivo demonstration of TMUB1 function.

    Evidence Hops−/− mouse model; co-IP of TMUB1 with p53; importin-α competition assay; mitochondrial fractionation; in vivo apoptosis upon chemotherapy

    PMID:31867855

    Open questions at the time
    • Whether TMUB1–p53 interaction is direct or requires bridging factors not structurally resolved
    • Relative contributions of p53 stabilization versus nuclear import blockade not quantitatively separated
    • Tumor-suppressive consequences of TMUB1 loss not formally tested in cancer models
  5. 2022 High

    The PD-L1 stabilization mechanism answered how tumor cells upregulate immune checkpoint ligands post-translationally: TMUB1 outcompetes HUWE1 at K281 to prevent polyubiquitination and recruits STT3A for N-glycosylation, establishing TMUB1 as a targetable node in immune evasion.

    Evidence Co-IP of TMUB1 with PD-L1 and HUWE1; ubiquitination site mapping at K281; glycosylation assay with STT3A; TMUB1 KD/OE with PD-L1 stability readout; competing peptide in mouse tumor models

    PMID:36376293

    Open questions at the time
    • Whether TMUB1-mediated PD-L1 stabilization occurs in all tumor types or is context-dependent
    • Structural basis of TMUB1 competition with HUWE1 unknown
    • Whether TMUB1 regulates glycosylation of other ER clients besides PD-L1 not examined

Open questions

Synthesis pass · forward-looking unresolved questions
  • No structural model of TMUB1 exists, and how a single shuttling protein coordinates such diverse substrates (p53, PD-L1, GluR2, NPM/Arf) through its ubiquitin-like domain and transmembrane segments remains mechanistically unresolved.
  • No crystal or cryo-EM structure of TMUB1 or any of its complexes
  • How substrate selectivity is achieved across ER, cytoplasm, and synaptic compartments unknown
  • Functional significance of the ubiquitin-like domain not dissected by mutagenesis across all interaction partners

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 2 GO:0098772 molecular function regulator activity 2
Localization
GO:0005634 nucleus 2 GO:0005829 cytosol 2 GO:0005783 endoplasmic reticulum 1
Pathway
R-HSA-168256 Immune System 1 R-HSA-392499 Metabolism of proteins 1 R-HSA-5357801 Programmed Cell Death 1
Partners
CD274CDKN2AGRIA2GRIP1HUWE1NPM1STT3ATP53
Complex memberships
NPM–p19(Arf)–TMUB1 trimeric complex

Evidence

Reading pass · 5 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2005 TMUB1 (HOPS/Hepatocyte Odd Protein Shuttling) is a shuttling protein that binds elongation factor eEF-1A and interferes with protein synthesis. Its nuclear export is governed by cAMP signaling and mediated via the CRM-1 export receptor. Overexpression strongly reduces cell proliferation. Regenerating liver cDNA library screen; co-immunoprecipitation with eEF-1A; cAMP/CRM-1 export assays; overexpression in H-35 and 3T3-NIH cells with proliferation readout Journal of cell science Medium 16014383
2008 TMUB1 (Tmub1/HOPS) facilitates surface recycling of GluR2-containing AMPA receptors. It is enriched in synaptosomal membranes, co-immunoprecipitates with GluR2 and GRIP from mouse brain, and its knockdown reduces AMPA receptor currents and synaptic surface expression of GluR2 (but not GluR1), while overexpression increases GluR2 surface expression. RNAi knockdown in neurons with electrophysiological readout; overexpression with surface biotinylation assay; co-immunoprecipitation from mouse brain; recycling assay PloS one Medium 18665261
2012 TMUB1 (HOPS) directly interacts with nucleophosmin (NPM) and p19(Arf), forming a functionally active trimeric complex. NPM regulates HOPS half-life, while HOPS stabilizes p19(Arf) and controls its nucleolar localization. HOPS knockdown causes centrosome hyperamplification and multinucleated cells; overexpression causes G0/G1 arrest. Co-immunoprecipitation; protein stability assays; siRNA knockdown with cell cycle and centrosome phenotype readouts; overexpression with cell cycle analysis Oncogene Medium 22890319
2019 TMUB1 (HOPS) retains p53 in the cytoplasm by binding p53, inhibiting its proteasomal degradation, and interfering with importin-α to block nuclear import. This promotes p53 recruitment to mitochondria and induction of the intrinsic apoptosis pathway. Hops−/− mice show significantly reduced apoptosis upon chemotherapeutic challenge. Hops−/− mouse model; co-immunoprecipitation of HOPS with p53; importin-α interaction assay; mitochondrial fractionation; apoptosis assays in vivo and in cells EMBO reports High 31867855
2022 TMUB1 modulates PD-L1 post-translational modifications in tumor cells: it competes with the E3 ubiquitin ligase HUWE1 for interaction with PD-L1 and inhibits PD-L1 polyubiquitination at K281 in the endoplasmic reticulum. Additionally, TMUB1 recruits STT3A to enhance PD-L1 N-glycosylation and stability, promoting PD-L1 maturation and tumor immune evasion. Co-immunoprecipitation of TMUB1 with PD-L1 and HUWE1; ubiquitination assay identifying K281 site; glycosylation assays with STT3A recruitment; TMUB1 knockdown/overexpression with PD-L1 stability and immune evasion readouts; synthetic competing peptide in mouse tumor models Nature communications High 36376293

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2014 PLEKHM1 regulates autophagosome-lysosome fusion through HOPS complex and LC3/GABARAP proteins. Molecular cell 463 25498145
2013 CORVET and HOPS tethering complexes - coordinators of endosome and lysosome fusion. Journal of cell science 416 23645161
2004 Xanthohumol and related prenylflavonoids from hops and beer: to your good health! Phytochemistry 407 15231405
2020 ORF3a of the COVID-19 virus SARS-CoV-2 blocks HOPS complex-mediated assembly of the SNARE complex required for autolysosome formation. Developmental cell 301 33422265
1999 Antiproliferative and cytotoxic effects of prenylated flavonoids from hops (Humulus lupulus) in human cancer cell lines. Food and chemical toxicology : an international journal published for the British Industrial Biological Research Association 281 10418944
2012 Molecular architecture of the multisubunit homotypic fusion and vacuole protein sorting (HOPS) tethering complex. Proceedings of the National Academy of Sciences of the United States of America 231 22308417
2014 Interaction of the HOPS complex with Syntaxin 17 mediates autophagosome clearance in Drosophila. Molecular biology of the cell 211 24554766
2006 Purification of active HOPS complex reveals its affinities for phosphoinositides and the SNARE Vam7p. The EMBO journal 210 16601699
2013 Tethering complexes in the endocytic pathway: CORVET and HOPS. The FEBS journal 156 23351085
2006 Xanthohumol, a prenylflavonoid derived from hops induces apoptosis and inhibits NF-kappaB activation in prostate epithelial cells. Cancer letters 138 16563612
2008 Drosophila HOPS and AP-3 complex genes are required for a Deltex-regulated activation of notch in the endosomal trafficking pathway. Developmental cell 134 19000840
2015 Recruitment of VPS33A to HOPS by VPS16 Is Required for Lysosome Fusion with Endosomes and Autophagosomes. Traffic (Copenhagen, Denmark) 125 25783203
2005 The vacuolar kinase Yck3 maintains organelle fragmentation by regulating the HOPS tethering complex. The Journal of cell biology 125 15684030
2015 The small GTPase Arl8b regulates assembly of the mammalian HOPS complex on lysosomes. Journal of cell science 124 25908847
2010 Defined subunit arrangement and rab interactions are required for functionality of the HOPS tethering complex. Traffic (Copenhagen, Denmark) 122 20604902
2005 Sec17p and HOPS, in distinct SNARE complexes, mediate SNARE complex disruption or assembly for fusion. The EMBO journal 100 15889152
2012 The HOPS proteins hVps41 and hVps39 are required for homotypic and heterotypic late endosome fusion. Traffic (Copenhagen, Denmark) 98 23167963
2019 CORVET, CHEVI and HOPS - multisubunit tethers of the endo-lysosomal system in health and disease. Journal of cell science 96 31092635
2020 Loss-of-Function Variants in HOPS Complex Genes VPS16 and VPS41 Cause Early Onset Dystonia Associated with Lysosomal Abnormalities. Annals of neurology 94 32808683
2003 Inhibitors of nitric oxide production from hops (Humulus lupulus L.). Biological & pharmaceutical bulletin 91 12520174
1993 Site-specific genetic recombination: hops, flips, and flops. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 91 8392474
2005 Comparison of the in vitro estrogenic activities of compounds from hops (Humulus lupulus) and red clover (Trifolium pratense). Journal of agricultural and food chemistry 89 16076101
2015 Characterization of the Mammalian CORVET and HOPS Complexes and Their Modular Restructuring for Endosome Specificity. The Journal of biological chemistry 85 26463206
2017 Scc2/Nipbl hops between chromosomal cohesin rings after loading. eLife 83 28914604
2019 HOPS: automated detection and authentication of pathogen DNA in archaeological remains. Genome biology 82 31842945
2009 Vps41 phosphorylation and the Rab Ypt7 control the targeting of the HOPS complex to endosome-vacuole fusion sites. Molecular biology of the cell 82 19193765
2017 Pacer Mediates the Function of Class III PI3K and HOPS Complexes in Autophagosome Maturation by Engaging Stx17. Molecular cell 81 28306502
2013 Structural basis of Vps33A recruitment to the human HOPS complex by Vps16. Proceedings of the National Academy of Sciences of the United States of America 81 23901104
2011 HOPS drives vacuole fusion by binding the vacuolar SNARE complex and the Vam7 PX domain via two distinct sites. Molecular biology of the cell 76 21613544
2010 Xanthohumol, a chalcon derived from hops, inhibits hepatic inflammation and fibrosis. Molecular nutrition & food research 76 20087858
2018 Xanthohumol, a prenylated flavonoid from Hops, exerts anticancer effects against gastric cancer in vitro. Oncology reports 75 30272303
2022 Promoting anti-tumor immunity by targeting TMUB1 to modulate PD-L1 polyubiquitination and glycosylation. Nature communications 74 36376293
2009 HOPS interacts with Apl5 at the vacuole membrane and is required for consumption of AP-3 transport vesicles. Molecular biology of the cell 73 19741093
2022 Structure of the HOPS tethering complex, a lysosomal membrane fusion machinery. eLife 72 36098503
2000 Prenylflavonoids from hops inhibit the metabolic activation of the carcinogenic heterocyclic amine 2-amino-3-methylimidazo[4, 5-f]quinoline, mediated by cDNA-expressed human CYP1A2. Drug metabolism and disposition: the biological fate of chemicals 71 11038156
2016 CATCHR, HOPS and CORVET tethering complexes share a similar architecture. Nature structural & molecular biology 70 27428774
2010 The novel endosomal membrane protein Ema interacts with the class C Vps-HOPS complex to promote endosomal maturation. The Journal of cell biology 66 20194640
2014 Adjustment of host cells for accommodation of symbiotic bacteria: vacuole defunctionalization, HOPS suppression, and TIP1g retargeting in Medicago. The Plant cell 62 25217511
2014 Xanthohumol, a prenylated chalcone from beer hops, acts as an α-glucosidase inhibitor in vitro. Journal of agricultural and food chemistry 60 24897556
2010 Xanthohumol, a prenylated chalcone derived from hops, inhibits proliferation, migration and interleukin-8 expression of hepatocellular carcinoma cells. International journal of oncology 57 20043079
2019 The Multiple Biological Targets of Hops and Bioactive Compounds. Chemical research in toxicology 55 30608650
2008 SPE-39 family proteins interact with the HOPS complex and function in lysosomal delivery. Molecular biology of the cell 55 19109425
2013 The tethering complex HOPS catalyzes assembly of the soluble SNARE Vam7 into fusogenic trans-SNARE complexes. Molecular biology of the cell 54 24088569
2017 Adaptor Protein-3-Dependent Vacuolar Trafficking Involves a Subpopulation of COPII and HOPS Tethering Proteins. Plant physiology 53 28559361
2008 Structural basis of human pregnane X receptor activation by the hops constituent colupulone. Molecular pharmacology 53 18768384
2007 Inhibition of topoisomerase I activity and efflux drug transporters' expression by xanthohumol. from hops. Archives of pharmacal research 51 18087812
2013 The late endosomal HOPS complex anchors active G-protein signaling essential for pathogenesis in magnaporthe oryzae. PLoS pathogens 49 23935502
2017 Salmonella exploits the host endolysosomal tethering factor HOPS complex to promote its intravacuolar replication. PLoS pathogens 48 29084291
2004 In vitro binding experiments with a Valerian, hops and their fixed combination extract (Ze91019) to selected central nervous system receptors. Phytomedicine : international journal of phytotherapy and phytopharmacology 47 15636177
2024 De novo biosynthesis of the hops bioactive flavonoid xanthohumol in yeast. Nature communications 46 38177132
2018 Phosphoinositides control the localization of HOPS subunit VPS41, which together with VPS33 mediates vacuole fusion in plants. Proceedings of the National Academy of Sciences of the United States of America 44 30104351
2015 Toxoplasma gondii Vps11, a subunit of HOPS and CORVET tethering complexes, is essential for the biogenesis of secretory organelles. Cellular microbiology 44 25640905
2010 The ERM proteins interact with the HOPS complex to regulate the maturation of endosomes. Molecular biology of the cell 44 21148287
2018 Novel anti-obesity effects of beer hops compound xanthohumol: role of AMPK signaling pathway. Nutrition & metabolism 43 29946343
2019 HOPS-dependent endosomal fusion required for efficient cytosolic delivery of therapeutic peptides and small proteins. Proceedings of the National Academy of Sciences of the United States of America 42 30610181
2016 Xanthohumol, a Prenylated Chalcone from Hops, Inhibits the Viability and Stemness of Doxorubicin-Resistant MCF-7/ADR Cells. Molecules (Basel, Switzerland) 42 28036030
2021 Bi-allelic variants in HOPS complex subunit VPS41 cause cerebellar ataxia and abnormal membrane trafficking. Brain : a journal of neurology 41 33764426
2021 Neurodegenerative VPS41 variants inhibit HOPS function and mTORC1-dependent TFEB/TFE3 regulation. EMBO molecular medicine 41 33851776
2023 Consecutive functions of small GTPases guide HOPS-mediated tethering of late endosomes and lysosomes. Cell reports 40 36640308
2015 Inhibition of Osteoclastogenesis and Bone Resorption in vitro and in vivo by a prenylflavonoid xanthohumol from hops. Scientific reports 40 26620037
2013 Xanthohumol, a main prenylated chalcone from hops, reduces liver damage and modulates oxidative reaction and apoptosis in hepatitis C virus infected Tupaia belangeri. International immunopharmacology 39 23669332
2012 Impaired stimulation of p38α-MAPK/Vps41-HOPS by LPS from pathogenic Coxiella burnetii prevents trafficking to microbicidal phagolysosomes. Cell host & microbe 37 23245320
2006 In vitro studies of intestinal permeability and hepatic and intestinal metabolism of 8-prenylnaringenin, a potent phytoestrogen from hops (Humulus lupulus L.). Pharmaceutical research 37 16715376
2006 Molecular phylogeny of wild hops, Humulus lupulus L. Heredity 36 16685279
2011 Hops (Humulus lupulus) inhibits oxidative estrogen metabolism and estrogen-induced malignant transformation in human mammary epithelial cells (MCF-10A). Cancer prevention research (Philadelphia, Pa.) 35 21997247
2005 HOPS: a novel cAMP-dependent shuttling protein involved in protein synthesis regulation. Journal of cell science 34 16014383
2003 Comprehensive analysis of orthologous protein domains using the HOPS database. Genome research 33 14525933
2020 Xanthohumol, a Prenylated Flavonoid from Hops, Induces DNA Damages in Colorectal Cancer Cells and Sensitizes SW480 Cells to the SN38 Chemotherapeutic Agent. Cells 32 32290112
2020 AP-3 vesicle uncoating occurs after HOPS-dependent vacuole tethering. The EMBO journal 32 32840906
2015 The Habc domain of the SNARE Vam3 interacts with the HOPS tethering complex to facilitate vacuole fusion. The Journal of biological chemistry 32 25564619
2013 Differential regulation of detoxification enzymes in hepatic and mammary tissue by hops (Humulus lupulus) in vitro and in vivo. Molecular nutrition & food research 32 23512484
2016 Impact of xanthohumol (a prenylated flavonoid from hops) on DNA stability and other health-related biochemical parameters: Results of human intervention trials. Molecular nutrition & food research 30 26840505
2015 The inhibitory effects of xanthohumol, a prenylated chalcone derived from hops, on cell growth and tumorigenesis in human pancreatic cancer. Biomedicine & pharmacotherapy = Biomedecine & pharmacotherapie 30 26211581
2019 HOPS/TMUB1 retains p53 in the cytoplasm and sustains p53-dependent mitochondrial apoptosis. EMBO reports 29 31867855
2012 Phosphorylation of the effector complex HOPS by the vacuolar kinase Yck3p confers Rab nucleotide specificity for vacuole docking and fusion. Molecular biology of the cell 29 22787280
2008 Inhibitory effects of xanthohumol from hops (Humulus lupulus L.) on human hepatocellular carcinoma cell lines. Phytotherapy research : PTR 29 18814205
2019 Disruption of vacuolar protein sorting components of the HOPS complex leads to enhanced secretion of recombinant proteins in Pichia pastoris. Microbial cell factories 28 31269943
2015 Regulation of lipid droplet dynamics in Saccharomyces cerevisiae depends on the Rab7-like Ypt7p, HOPS complex and V1-ATPase. Biology open 28 25948753
2023 C9orf72-catalyzed GTP loading of Rab39A enables HOPS-mediated membrane tethering and fusion in mammalian autophagy. Nature communications 27 37821429
2021 Bi-allelic VPS16 variants limit HOPS/CORVET levels and cause a mucopolysaccharidosis-like disease. EMBO molecular medicine 27 33938619
2014 KDT501, a derivative from hops, normalizes glucose metabolism and body weight in rodent models of diabetes. PloS one 26 24498211
2018 6- and 8-Prenylnaringenin, Novel Natural Histone Deacetylase Inhibitors Found in Hops, Exert Antitumor Activity on Melanoma Cells. Cellular physiology and biochemistry : international journal of experimental cellular physiology, biochemistry, and pharmacology 25 30458450
2007 Isohumulones derived from hops ameliorate renal injury via an anti-oxidative effect in Dahl salt-sensitive rats. Hypertension research : official journal of the Japanese Society of Hypertension 25 17460388
2022 Arabidopsis HOPS subunit VPS41 carries out plant-specific roles in vacuolar transport and vegetative growth. Plant physiology 24 35417008
2019 Vps8 overexpression inhibits HOPS-dependent trafficking routes by outcompeting Vps41/Lt. eLife 24 31194677
2008 Transmembrane and ubiquitin-like domain-containing protein 1 (Tmub1/HOPS) facilitates surface expression of GluR2-containing AMPA receptors. PloS one 24 18665261
2007 Clinical safety and efficacy of NG440: a novel combination of rho iso-alpha acids from hops, rosemary, and oleanolic acid for inflammatory conditions. Canadian journal of physiology and pharmacology 24 18066133
2021 Xanthohumol, a Prenylated Chalcone Derived from Hops, Inhibits Growth and Metastasis of Melanoma Cells. Cancers 23 33572775
2020 Yeast phosphatidic acid phosphatase Pah1 hops and scoots along the membrane phospholipid bilayer. Journal of lipid research 23 32540926
2012 Hepatocyte odd protein shuttling (HOPS) is a bridging protein in the nucleophosmin-p19 Arf network. Oncogene 23 22890319
2011 Clathrin-dependent mechanisms modulate the subcellular distribution of class C Vps/HOPS tether subunits in polarized and nonpolarized cells. Molecular biology of the cell 23 21411634
2018 Synthesis and Antiproliferative Activity of Minor Hops Prenylflavonoids and New Insights on Prenyl Group Cyclization. Molecules (Basel, Switzerland) 22 29597299
2016 The HOPS/Class C Vps Complex Tethers High-Curvature Membranes via a Direct Protein-Membrane Interaction. Traffic (Copenhagen, Denmark) 22 27307091
2014 Caenorhabditis elegans HOPS and CCZ-1 mediate trafficking to lysosome-related organelles independently of RAB-7 and SAND-1. Molecular biology of the cell 22 24501423
2008 Genetic and epigenetic stability of cryopreserved and cold-stored hops (Humulus lupulus L.). Cryobiology 22 18822279
2017 VPS18 recruits VPS41 to the human HOPS complex via a RING-RING interaction. The Biochemical journal 21 28931724
2016 Hops for Menopausal Vasomotor Symptoms: Mechanisms of Action. Journal of menopausal medicine 21 27617238
2012 SNAREs, HOPS and regulatory lipids control the dynamics of vacuolar actin during homotypic fusion in S. cerevisiae. Journal of cell science 21 22357954
2009 The antimicrobial effects of hops (Humulus lupulus L.) on ruminal hyper ammonia-producing bacteria. Letters in applied microbiology 21 19413813
2022 Hops extract and xanthohumol ameliorate bone loss induced by iron overload via activating Akt/GSK3β/Nrf2 pathway. Journal of bone and mineral metabolism 20 35106609