Affinage

Showing HERPUD1 — HERP is a alias.

HERPUD1

Homocysteine-responsive endoplasmic reticulum-resident ubiquitin-like domain member 1 protein · UniProt Q15011

Length
391 aa
Mass
43.7 kDa
Annotated
2026-06-10
100 papers in source corpus 30 papers cited in narrative 30 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

HERPUD1 (Herp/Mif1) is an ER-resident integral membrane protein with a cytoplasm-facing N-terminal ubiquitin-like domain that serves as a stress-inducible scaffolding hub of the ER-associated degradation (ERAD) machinery (PMID:10922362, PMID:16289116). It is among the most strongly induced targets of the unfolded protein response, regulated combinatorially through both the IRE1/XBP1 and ATF6 ER-stress branches and the shared PERK/eIF2α/ATF4 branch (PMID:14742429), and through additional transcription factors acting at its promoter elements, including Luman/CREB3 at ERSE-II and Nrf1 at antioxidant response elements (PMID:16940180, PMID:25637874). Mechanistically, HERPUD1 assembles a high-molecular-mass ERAD complex by binding directly to the E3 ubiquitin ligase HRD1 and recruiting the retrotranslocation factors p97, Derlin/DERL2 and VIMP, and by engaging ubiquilins to deliver substrates to the 26S proteasome (PMID:16289116, PMID:18042451, PMID:18307982, PMID:24366871). Its UBL domain is required for efficient HRD1-mediated ubiquitylation and degradation of lumenal substrates such as NHK α1-antitrypsin, and HERPUD1 compartmentalizes misfolded proteins and HRD1 within the ER quality control compartment, where it acts as a membrane-anchored shuttle analogous to hHR23 (PMID:21149444, PMID:24478453, PMID:24366871). Through ERAD-dependent turnover of ER Ca2+ release channels, principally IP3 receptors, HERPUD1 limits Ca2+ release to the cytosol and mitochondria, protecting cells against ER-stress- and oxidative-stress-induced apoptosis (PMID:15102845, PMID:22045699, PMID:26616647); loss of this function in vivo produces impaired glucose tolerance, ischemic vulnerability, and cardiac hypertrophy with elevated IP3R levels (PMID:22479592, PMID:29042597). HERPUD1 is itself a short-lived protein whose UBL domain drives rapid proteasomal turnover, including gp78/Ube2g2-mediated K48-linked ubiquitylation during ER-stress recovery and POSH-mediated K63-linked ubiquitylation that governs its trafficking between the trans-Golgi network and ER upon Ca2+ perturbation (PMID:14550564, PMID:17420289, PMID:24496447). Beyond core ERAD, HERPUD1 binds TBK1 to amplify MAVS-dependent IRF3/NF-κB signaling and type-I/III interferon production during RNA virus infection (PMID:28954889).

Mechanistic history

Synthesis pass · year-by-year structured walk · 12 steps
  1. 2000 High

    Established the basic identity of HERPUD1 as a stress-inducible ER membrane protein, defining the molecular object before any function was known.

    Evidence Antibody detection, membrane topology assays and stress induction in cultured cells; promoter reporter assays mapping a functional ERSE

    PMID:10708769 PMID:10922362

    Open questions at the time
    • Cytoplasmic function of the UBL domain not yet defined
    • No binding partners identified
  2. 2002 Medium

    First physical interactions and a functional context were assigned, linking Herp to presenilins and amyloid processing.

    Evidence Co-immunoprecipitation, overexpression and Aβ ELISA in presenilin-dependent cells; ULD deletion mapping

    PMID:11799129 PMID:14550564

    Open questions at the time
    • UBL/ULD role in Aβ generation dissociated from degradation but mechanism unclear
    • Single-lab Co-IP without reciprocal validation in independent systems
  3. 2004 High

    Resolved how HERPUD1 is induced and showed it is cytoprotective during ER stress, connecting transcriptional control to a survival phenotype.

    Evidence Genetic epistasis across UPR branch mutants; RNAi, overexpression, Ca2+ imaging, caspase assays and knockout F9 cells

    PMID:14742429 PMID:15102845 PMID:15147274

    Open questions at the time
    • Molecular basis of Ca2+ stabilization not yet defined
    • ERAD substrate spectrum not delineated
  4. 2005 High

    Defined HERPUD1 as a direct binding partner of HRD1 within a retrotranslocation complex, establishing its scaffolding role in ERAD.

    Evidence Reciprocal Co-IP, sucrose gradient sedimentation and pull-down assays

    PMID:16289116

    Open questions at the time
    • Substrate specificity of the complex not addressed
    • Stoichiometry of HERP within the complex unknown
  5. 2007 High

    Defined substrate selectivity and the proteasome-linking function, and identified POSH-mediated K63 ubiquitylation controlling HERP localization.

    Evidence siRNA knockdown, reciprocal Co-IP, proteasome association and pulse-chase assays for BiP-substrate ERAD; in vitro ubiquitination and confocal localization for POSH

    PMID:17420289 PMID:18042451

    Open questions at the time
    • Why HERP serves nonglycosylated/BiP but not calnexin substrates unresolved
    • Trigger linking Ca2+ perturbation to POSH ubiquitylation not defined
  6. 2006 High

    Identified Luman/CREB3 as a direct transcriptional activator at ERSE-II, expanding the regulatory inputs beyond the canonical UPR triad.

    Evidence ChIP, promoter mutagenesis, siRNA knockdown and luciferase reporter assays

    PMID:16940180

    Open questions at the time
    • Relative contribution of Luman versus ATF6/XBP1/ATF4 in vivo unclear
  7. 2008 Medium

    Showed ubiquilin recruitment by HERP is the link delivering specific substrates to the proteasome, mechanistically connecting scaffold to degradation.

    Evidence Co-IP, ubiquilin siRNA knockdown and transmembrane-deletion mutant rescue with pulse-chase degradation

    PMID:18307982

    Open questions at the time
    • Substrate subset requiring ubiquilins not fully mapped
    • Single-lab evidence
  8. 2010 High

    Established UBL-domain-dependent regulation of HRD1 ubiquitylation and HERP's role in degrading ER Ca2+ release channels, unifying ERAD scaffolding with Ca2+ homeostasis.

    Evidence UBL mutagenesis, ubiquitylation and pulse-chase assays for NHK; KD, UBL deletion, proteasome inhibition, Ca2+ imaging and A53T-αSyn transgenic mice for IP3R/RyR turnover

    PMID:21149444 PMID:22045699

    Open questions at the time
    • How HERP selects channel substrates not defined
    • Direct versus indirect role in channel ubiquitylation not separated
  9. 2013 High

    Defined HERPUD1's spatial role in the ERQC, its redundancy with HERP2, and a DERL2-recruitment mechanism, refining the architecture of the active retrotranslocation complex.

    Evidence Immunofluorescence/co-localization with KD; double siRNA knockdown, Co-IP, ubiquitination and retrotranslocation assays across multiple substrates

    PMID:24366871 PMID:24478453

    Open questions at the time
    • Structural model of the membrane-anchored hub is predictive only
    • DERL2-independent UBL function not mechanistically explained
  10. 2014 High

    Mapped the CUE-UBL interaction driving gp78/Ube2g2-mediated K48 ubiquitylation, explaining HERP's own rapid turnover during stress recovery.

    Evidence In vitro ubiquitylation reconstitution, domain-interaction mapping, siRNA and pulse-chase

    PMID:24496447

    Open questions at the time
    • Coordination between gp78-driven and POSH-driven ubiquitylation unresolved
  11. 2017 Medium

    Extended HERPUD1 function beyond ERAD into innate immunity and into in vivo cardiac physiology, broadening its biological scope.

    Evidence Co-IP, IFN reporter, phosphorylation and nuclear translocation assays with virus replication for TBK1/MAVS; knockout mice, echocardiography, KD, IP3R Western and Ca2+ imaging for cardiac hypertrophy

    PMID:28954889 PMID:29042597

    Open questions at the time
    • Direct versus scaffold-mediated role in TBK1 signaling unclear
    • Single-lab evidence for the immune axis
  12. 2022 Medium

    Implicated HERPUD1 in non-canonical contexts including ferroptosis, osteoblast differentiation, autophagy and metabolic regulation, indicating context-dependent roles.

    Evidence Knockdown/overexpression with ubiquitination, GSH and ferroptosis assays; differentiation, autophagy flux, and insulin-secretion readouts across separate studies

    PMID:23620059 PMID:24120520 PMID:29570393 PMID:36038536

    Open questions at the time
    • Whether these phenotypes depend on the core ERAD/Ca2+ functions not resolved
    • Each context characterized in a single study

Open questions

Synthesis pass · forward-looking unresolved questions
  • How HERPUD1's substrate selection, its self-turnover by competing ubiquitin ligases, and its diverse non-ERAD activities are mechanistically integrated remains unresolved.
  • No experimental structure of the membrane-anchored HERP-HRD1 complex
  • Rules governing substrate channeling to ubiquilins versus DERL2 undefined
  • Mechanistic link between Ca2+ regulation and the immune/metabolic roles unestablished

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0060090 molecular adaptor activity 4 GO:0140096 catalytic activity, acting on a protein 3 GO:0031386 protein tag activity 2
Localization
GO:0005783 endoplasmic reticulum 4 GO:0005794 Golgi apparatus 1
Pathway
R-HSA-392499 Metabolism of proteins 4 R-HSA-8953897 Cellular responses to stimuli 3 R-HSA-168256 Immune System 1
Partners
DERL2GP78HRD1P97/VCPPOSHTBK1UBQLNVIMP
Complex memberships
HRD1-SEL1L ERAD complex

Evidence

Reading pass · 30 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2000 HERPUD1 (Herp/Mif1) is an integral ER membrane protein with both N- and C-termini facing the cytoplasm, containing an N-terminal ubiquitin-like domain; it is strongly induced by ER stress agents (tunicamycin, thapsigargin, mercaptoethanol, A23187) and by homocysteine via the ER stress response pathway. Immunochemical analysis with Herp-specific antibodies, Western blot, mRNA expression analysis, membrane topology studies The Journal of biological chemistry High 10922362
2000 Mif1/HERPUD1 is a target of the unfolded protein response (UPR) pathway; its 5' flanking region contains a functional ER stress-responsive element (ERSE) sufficient for induction by tunicamycin. MMS activates Mif1 via a UPR-independent pathway. Overexpressed Mif1 protein localizes to the ER. Reporter gene assays with ERSE-containing promoter constructs, mRNA/protein induction analysis, subcellular localization by immunofluorescence FEBS letters Medium 10708769
2002 Herp physically interacts with both presenilin 1 (PS1) and presenilin 2 (PS2), and high Herp expression increases amyloid beta-protein (Aβ) generation in a presenilin-dependent manner; Herp was identified by a screen for cDNAs that increase gamma-cleavage. Co-immunoprecipitation, overexpression in cells with and without presenilin, Aβ ELISA The Journal of biological chemistry Medium 11799129
2003 The ubiquitin-like domain (ULD) of Herp is involved in its rapid proteasome-dependent degradation but is not required for enhancement of Aβ generation or interaction with presenilin. ULD deletion mutagenesis, pulse-chase degradation assay, co-immunoprecipitation FEBS letters Medium 14550564
2004 Herp is dually regulated by both the ER stress-specific branches (Ire1/XBP-1 and ATF6-dependent) and the shared cellular stress branch (PERK/eIF-2alpha/ATF4-dependent) of the UPR. ATF4 is required for Herp induction when only the shared branch is activated, but not when both branches are active. Genetic epistasis using knockout/knockdown of UPR branch components, Northern blot, reporter assays The Journal of biological chemistry High 14742429
2004 Herp stabilizes ER Ca2+ homeostasis and mitochondrial function in neurons during ER stress; overexpression promotes survival by preventing ER Ca2+ overload and suppressing caspase-3 and caspase-12 activation. Lethal ER stress causes caspase-mediated cleavage of Herp, generating a 30-kDa fragment; mutation of the caspase cleavage site enhances neuroprotection. RNA interference knockdown, Herp overexpression, Ca2+ imaging, mitochondrial function assays, caspase activity assays, mutagenesis of caspase cleavage site The Journal of biological chemistry High 15102845
2004 Herp knockout cells (F9) are more vulnerable to ER stress, show enhanced ER stress signaling and stabilization of an ERAD substrate in early stress, and are prone to apoptosis in late stress. The N-terminal ubiquitin-like domain of Herp is required for cell survival under ER stress. Herp itself is rapidly degraded by the proteasome. F9 Herp knockout cells via gene targeting, tunicamycin treatment, ERAD substrate stability assay, transfection with domain-deletion constructs Genes to cells High 15147274
2005 HERP forms a high-molecular-mass complex at the ER membrane containing the ubiquitin E3 ligase HRD1, retrotranslocation factors p97, Derlin-1, and VIMP. HERP binds directly to HRD1, while p97 interacts with Derlin-1 and HRD1. Co-immunoprecipitation, sucrose gradient sedimentation, pull-down assays Journal of molecular biology High 16289116
2006 Luman/CREB3, an ER membrane-bound transcription factor, directly activates Herp transcription through the ER stress response element II (ERSE-II; ATTGG-N-CCACG) in the Herp promoter. Luman physically associates with the ERSE-II second half-site. Luman knockdown attenuates Herp induction during ER stress. Chromatin immunoprecipitation (ChIP), promoter mutagenesis, siRNA knockdown, luciferase reporter assays Molecular and cellular biology High 16940180
2007 Herp is required for degradation of nonglycosylated BiP substrates but not calnexin substrates; Herp forms a complex with ubiquitinated proteins and the 26S proteasome, linking substrates to the proteasome. Herp associates with Derlin-1, and either p97/Hrd1 mutant expression or Herp knockdown inhibits BiP substrate degradation. Co-immunoprecipitation, siRNA knockdown, proteasome association assay, pulse-chase degradation assay Molecular cell High 18042451
2007 POSH, a ubiquitin E3 ligase, interacts with Herp and promotes K63-linked polyubiquitination of Herp via its UBL domain. This POSH-dependent ubiquitination controls Herp relocalization from the trans-Golgi network to the ER upon Ca2+ perturbation by thapsigargin, and regulates ER Ca2+ homeostasis. Co-immunoprecipitation, confocal microscopy, in vitro ubiquitination assay, dominant-negative overexpression, RNAi, Ca2+ measurement The Journal of cell biology High 17420289
2008 Herp interacts with members of the ubiquilin family; this interaction is required for efficient ERAD of a subset of substrates (e.g., CD3δ). Herp mutants lacking the transmembrane domain but retaining ubiquilin-binding capacity stabilize CD3δ, indicating that ubiquilin recruitment by Herp facilitates substrate delivery to the proteasome. Co-immunoprecipitation, siRNA knockdown of ubiquilins, overexpression of Herp transmembrane domain deletion mutants, pulse-chase degradation assay Biochemical and biophysical research communications Medium 18307982
2010 Herp regulates Hrd1-mediated ubiquitylation in a UBL domain-dependent manner; the Herp UBL domain is required for efficient ubiquitylation and degradation of the Hrd1-specific ERAD substrate NHK (null Hong Kong α1-antitrypsin). Upon ER stress, Herp is rapidly degraded and substituted by newly synthesized Herp at Hrd1 complexes, and multiple Hrd1 copies in a complex can bind variable numbers of Herp molecules. UBL domain mutagenesis, co-immunoprecipitation, pulse-chase assay, ubiquitylation assay The Journal of biological chemistry High 21149444
2010 Herp knockdown facilitates degradation of cytosolic proteins (α-synuclein, synphilin-1) and improves cell viability during proteasomal inhibition. Herp transiently binds α-synuclein, synphilin-1, and the E3 ligase SIAH1a during proteolytic stress (but not during ER stress), suggesting Herp delays ubiquitination of cytosolic substrates. siRNA knockdown, F9 Herp knockout cells, co-immunoprecipitation, protein stability assay, cell viability assay Genes to cells Medium 20604806
2011 Herp maintains ER Ca2+ homeostasis by facilitating proteasome-mediated degradation of ER-resident Ca2+ release channels (IP3 receptors and ryanodine receptors). UBL domain deletion or proteasome inhibition abolishes Herp-mediated ER Ca2+ stabilization. Aberrant accumulation of Ca2+ channels and chronic ER stress are observed in mutant A53T-αSyn transgenic mice. siRNA knockdown, UBL domain deletion, pharmacological proteasome inhibition, Ca2+ imaging, Western blot for IP3R/RyR levels, transgenic mouse model Human molecular genetics High 22045699
2012 Herp-deficient mice are viable but exhibit impaired glucose tolerance and increased vulnerability to brain ischemia. In Herp-deficient liver, degradation of ERAD substrates is reduced. Herp deficiency affects expression of ERAD-related proteins in a tissue-specific manner. Gene targeting knockout mice, glucose tolerance test, ischemia model, ERAD substrate stability assay, Western blot PloS one High 22479592
2013 Herp localizes to the ER quality control compartment (ERQC) and recruits HRD1 to this compartment; Herp is responsible for compartmentalization of misfolded proteins and the ubiquitin ligase HRD1 required for ERAD. Predicted structural similarity to ubiquitin-proteasome shuttle hHR23 (but with a transmembrane hairpin) suggests Herp functions as a membrane-anchored hub for ERAD machinery assembly. Immunofluorescence microscopy, siRNA knockdown, co-localization analysis, structural bioinformatics Molecular biology of the cell Medium 24478453
2013 HERP1 (HERPUD1) and HERP2 are both required for efficient HRD1-dependent ERAD of lumenal substrates (SHH, NHK). HERP2 is constitutively expressed while HERP1 is ER stress-induced. Both HERPs interact with HRD1 through a cytosol-facing region and help recruit DERL2 to the HRD1-SEL1L complex to form an active retrotranslocation complex. The UBL domain of HERP1 has an additional DERL2-independent function in ERAD. siRNA double knockdown, co-immunoprecipitation, ubiquitination assay, retrotranslocation assay, pulse-chase degradation The Journal of biological chemistry High 24366871
2013 Herp depletion leads to upregulation of autophagy regulators Atg5 and Beclin-1 (the latter via reduced Hrd1-dependent proteasomal degradation), resulting in increased autophagic flux that clears poly-ubiquitin protein aggregates and protects cells from glucose starvation-induced death. Stable shRNA knockdown, LC3-II Western blot, GFP-LC3 puncta assay, protein aggregation assay, cell viability assay Biochimica et biophysica acta Medium 24120520
2014 gp78 (an ER E3 ubiquitin ligase) together with Ube2g2 mediates polyubiquitylation of HERP, driven by a physical interaction between the CUE domain of gp78 and the UBL domain of HERP, leading to proteasomal degradation of HERP during ER stress recovery. In vitro ubiquitylation assay, co-immunoprecipitation, domain interaction mapping (CUE-UBL), siRNA knockdown, pulse-chase assay Journal of cell science High 24496447
2015 Nrf1 transcription factor directly activates Herpud1 expression during ER stress through antioxidant response elements in the Herpud1 promoter; loss of Nrf1 decreases Herpud1 expression and abolishes its ER stress induction. Nrf1 knockout cells, chromatin immunoprecipitation (ChIP), transactivation reporter assays, liver tissue analysis FEBS letters High 25637874
2015 HERPUD1 protects against oxidative stress-induced apoptosis by downregulating IP3 receptor (ITPR)-mediated Ca2+ release from the ER; HERPUD1 knockdown increases cytosolic and mitochondrial Ca2+ and cell death, which is abolished by ITPR antagonism or intracellular Ca2+ chelation. Stable shRNA knockdown, flow cytometry (cell death), Ca2+ imaging (cytosolic and mitochondrial), ITPR pharmacological inhibition, BAPTA-AM chelation Free radical biology & medicine Medium 26616647
2017 HERP interacts with TBK1 (TANK-binding kinase 1) and amplifies MAVS signaling, facilitating phosphorylation and nuclear translocation of IRF3 and NF-κB to enhance type-I and type-III IFN expression and inhibit RNA virus replication. HERP induction by EV71 is dependent on MAVS. Co-immunoprecipitation, reporter assays for IFN promoters, phosphorylation assays, nuclear translocation by immunofluorescence, siRNA knockdown, virus replication assay Journal of immunology Medium 28954889
2017 Herpud1 knockout mice develop cardiac hypertrophy and dysfunction; Herpud1 deficiency elevates IP3R protein levels and increases cytosolic and nuclear Ca2+ in cardiomyocytes. Herpud1 negatively regulates cardiac hypertrophy by promoting IP3R degradation via the ERAD pathway. Herpud1 knockout mouse model, echocardiography, siRNA knockdown in cardiomyocytes, IP3R Western blot, Ca2+ imaging, hypertrophic marker expression Scientific reports High 29042597
2018 HERPUD1 is required for osteoblast differentiation and mineralization; its expression increases as differentiation progresses with activation of ERAD and proteasomal degradation. HERPUD1 absence blocks mineralization and reduces alkaline phosphatase activity, while overexpression activates the osteoblast differentiation program. siRNA knockdown, overexpression in MC3T3-E1 and primary osteoblasts, alkaline phosphatase activity assay, calcium deposit measurement, Runx2/osterix expression FASEB journal Medium 29570393
2010 PRKCSH (glucosidase II beta subunit) interacts with HERP and inhibits HERP-mediated ubiquitination of TRPP2/polycystin-2, thereby protecting TRPP2 from ERAD; PRKCSH co-localizes with TRPP2 in the ER and binds TRPP2's C-terminal domain. Co-immunoprecipitation, ubiquitination assay, zebrafish overexpression/depletion, co-localization by immunofluorescence Human molecular genetics Medium 19801576
2006 Homocysteine-induced epigenetic hypermethylation of the HERP promoter correlates with decreased HERP mRNA expression, and promoter CpG island methylation is sufficient to suppress Herp transcription. Acute homocysteine causes transient promoter demethylation and Herp induction; the AARE and CREB binding sites in the Herp promoter mediate homocysteine-induced transcription. Luciferase reporter assays with AARE/CREB site mutations, methylation-specific PCR, mRNA quantitative PCR in patient samples and cell lines Experimental cell research Medium 17020760
2021 Hypoxia-induced p300 acetyltransferase interacts with and acetylates XBP1s, enhancing XBP1s stability and its transcriptional activation of Herpud1 through ERSE-II. The p300/XBP1s/Herpud1 axis promotes M2 macrophage polarization. Co-immunoprecipitation, acetylation assay, ChIP/promoter reporter assay, siRNA knockdown, macrophage polarization assay, mouse laser-induced CNV model Journal of cellular and molecular medicine Medium 34057287
2022 HERPUD1 promotes ferroptosis sensitivity in liver cancer cells by reducing MDM2 ubiquitination of glutathione synthetase (GSS), leading to GSS ubiquitination/degradation and decreased glutathione (GSH) levels, thus increasing susceptibility to ferroptosis. Overexpression/knockdown, ubiquitination assay for GSS/MDM2 interaction, GSH measurement, ferroptosis sensitivity assay, mouse xenograft model Cell death discovery Medium 36038536
2013 Herpud1 regulates insulin secretion in pancreatic beta cells; Herpud1 knockdown in DBA/2 islets reduces glucose-stimulated insulin secretion and decreases Nnt expression, while Herpud1 overexpression increases Nnt expression, placing Herpud1 upstream of Nnt in insulin secretion regulation. siRNA knockdown, Herpud1 overexpression in MIN6 cells and DBA/2 islets, glucose-stimulated insulin secretion assay, microarray profiling Diabetologia Medium 23620059

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2003 HES and HERP families: multiple effectors of the Notch signaling pathway. Journal of cellular physiology 1019 12548545
2000 Herp, a new ubiquitin-like membrane protein induced by endoplasmic reticulum stress. The Journal of biological chemistry 272 10922362
2017 Principles of the Kenzan Method for Robotic Cell Spheroid-Based Three-Dimensional Bioprinting<sup/>. Tissue engineering. Part B, Reviews 210 27917703
2001 HERP, a novel heterodimer partner of HES/E(spl) in Notch signaling. Molecular and cellular biology 187 11486045
2007 Characterization of an ERAD pathway for nonglycosylated BiP substrates, which require Herp. Molecular cell 180 18042451
2005 The ubiquitin-domain protein HERP forms a complex with components of the endoplasmic reticulum associated degradation pathway. Journal of molecular biology 177 16289116
2001 HERP, a new primary target of Notch regulated by ligand binding. Molecular and cellular biology 172 11486044
1994 Molecular cloning and functional characterization of a human VIP receptor from SUP-T1 lymphoblasts. Biochemical and biophysical research communications 145 7811244
1997 SUP-17, a Caenorhabditis elegans ADAM protein related to Drosophila KUZBANIAN, and its role in LIN-12/NOTCH signalling. Development (Cambridge, England) 139 9428412
2004 Herp is dually regulated by both the endoplasmic reticulum stress-specific branch of the unfolded protein response and a branch that is shared with other cellular stress pathways. The Journal of biological chemistry 135 14742429
2006 Epigenetic DNA hypermethylation of the HERP gene promoter induces down-regulation of its mRNA expression in patients with alcohol dependence. Alcoholism, clinical and experimental research 128 16573575
2017 Comparison of 3D-Printed Poly-ɛ-Caprolactone Scaffolds Functionalized with Tricalcium Phosphate, Hydroxyapatite, Bio-Oss, or Decellularized Bone Matrix<sup/>. Tissue engineering. Part A 125 28027692
2002 Endoplasmic reticulum stress-inducible protein, Herp, enhances presenilin-mediated generation of amyloid beta-protein. The Journal of biological chemistry 121 11799129
2018 Non-animal methods to predict skin sensitization (I): the Cosmetics Europe database<sup/>. Critical reviews in toxicology 117 29474128
2006 Luman/CREB3 induces transcription of the endoplasmic reticulum (ER) stress response protein Herp through an ER stress response element. Molecular and cellular biology 117 16940180
2004 Herp stabilizes neuronal Ca2+ homeostasis and mitochondrial function during endoplasmic reticulum stress. The Journal of biological chemistry 109 15102845
2004 Role of Herp in the endoplasmic reticulum stress response. Genes to cells : devoted to molecular & cellular mechanisms 104 15147274
1981 A second informational suppressor, SUP-7 X, in Caenorhabditis elegans. Genetics 91 7274656
1994 Mutations in the SUP-PF-1 locus of Chlamydomonas reinhardtii identify a regulatory domain in the beta-dynein heavy chain. The Journal of cell biology 88 8089181
2007 The Fox-1 family and SUP-12 coordinately regulate tissue-specific alternative splicing in vivo. Molecular and cellular biology 81 17923701
2008 Herp enhances ER-associated protein degradation by recruiting ubiquilins. Biochemical and biophysical research communications 78 18307982
1983 The genes sup-7 X and sup-5 III of C. elegans suppress amber nonsense mutations via altered transfer RNA. Cell 76 6571695
2016 The Scaffold Immune Microenvironment: Biomaterial-Mediated Immune Polarization in Traumatic and Nontraumatic Applications<sup/>. Tissue engineering. Part A 69 27736323
2017 Evaluation of Sofosbuvir (β-D-2'-deoxy-2'-α-fluoro-2'-β-C-methyluridine) as an inhibitor of Dengue virus replication<sup/>. Scientific reports 68 28740124
2009 Homocysteine-induced endoplasmic reticulum protein (herp) is up-regulated in parkinsonian substantia nigra and present in the core of Lewy bodies. Clinical neuropathology 68 19788048
2018 CapZyme-Seq Comprehensively Defines Promoter-Sequence Determinants for RNA 5' Capping with NAD<sup/>. Molecular cell 67 29681497
2014 Herp coordinates compartmentalization and recruitment of HRD1 and misfolded proteins for ERAD. Molecular biology of the cell 65 24478453
2003 sup-9, sup-10, and unc-93 may encode components of a two-pore K+ channel that coordinates muscle contraction in Caenorhabditis elegans. The Journal of neuroscience : the official journal of the Society for Neuroscience 64 14534247
1995 The unc-8 and sup-40 genes regulate ion channel function in Caenorhabditis elegans motorneurons. Genetics 62 7539392
2012 Derlin-1 deficiency is embryonic lethal, Derlin-3 deficiency appears normal, and Herp deficiency is intolerant to glucose load and ischemia in mice. PloS one 60 22479592
2010 Herp regulates Hrd1-mediated ubiquitylation in a ubiquitin-like domain-dependent manner. The Journal of biological chemistry 60 21149444
1996 The sup-pf-2 mutations of Chlamydomonas alter the activity of the outer dynein arms by modification of the gamma-dynein heavy chain. The Journal of cell biology 60 8991096
1995 Halotolerance in Methanosarcina spp.: Role of N(sup(epsilon))-Acetyl-(beta)-Lysine, (alpha)-Glutamate, Glycine Betaine, and K(sup+) as Compatible Solutes for Osmotic Adaptation. Applied and environmental microbiology 60 16535193
2011 The homocysteine-inducible endoplasmic reticulum (ER) stress protein Herp counteracts mutant α-synuclein-induced ER stress via the homeostatic regulation of ER-resident calcium release channel proteins. Human molecular genetics 57 22045699
1996 MoMuLV proviral integrations identified by Sup-F selection in tumors from infected myc/pim bitransgenic mice correlate with activation of the gfi-1 gene. Nucleic acids research 55 8692692
2006 Homocysteine-induced endoplasmic reticulum protein (Herp) is up-regulated in sporadic inclusion-body myositis and in endoplasmic reticulum stress-induced cultured human muscle fibers. Journal of neurochemistry 54 16441512
2019 Memantine protects thalamocortical hyper-glutamatergic transmission induced by NMDA receptor antagonism via activation of system xc<sup/>. Pharmacology research & perspectives 50 30784207
2004 The RNA-binding protein SUP-12 controls muscle-specific splicing of the ADF/cofilin pre-mRNA in C. elegans. The Journal of cell biology 50 15545320
1989 SUP-HD1: a new Hodgkin's disease-derived cell line with lymphoid features produces interferon-gamma. Blood 47 2554995
1987 Molecular analysis of TCRB and ABL in a t(7;9)-containing cell line (SUP-T3) from a human T-cell leukemia. Proceedings of the National Academy of Sciences of the United States of America 44 3025859
2013 Role of HERP and a HERP-related protein in HRD1-dependent protein degradation at the endoplasmic reticulum. The Journal of biological chemistry 43 24366871
2005 Evidence for functional redundancy between C. elegans ADAM proteins SUP-17/Kuzbanian and ADM-4/TACE. Developmental biology 43 16197940
2015 Induction of Herpud1 expression by ER stress is regulated by Nrf1. FEBS letters 42 25637874
2015 HERPUD1 protects against oxidative stress-induced apoptosis through downregulation of the inositol 1,4,5-trisphosphate receptor. Free radical biology & medicine 42 26616647
2018 HERP depletion inhibits zearalenone-induced apoptosis through autophagy activation in mouse ovarian granulosa cells. Toxicology letters 41 30394307
2010 PRKCSH/80K-H, the protein mutated in polycystic liver disease, protects polycystin-2/TRPP2 against HERP-mediated degradation. Human molecular genetics 41 19801576
2008 The psychology of psychiatric genetics: evidence that positive emotions in females moderate genetic sensitivity to social stress associated with the BDNF Val-sup-6-sup-6Met polymorphism. Journal of abnormal psychology 41 18729623
2007 The ubiquitin E3 ligase POSH regulates calcium homeostasis through spatial control of Herp. The Journal of cell biology 40 17420289
2000 The novel MMS-inducible gene Mif1/KIAA0025 is a target of the unfolded protein response pathway. FEBS letters 40 10708769
2018 Exercise Mitigates Alcohol Induced Endoplasmic Reticulum Stress Mediated Cognitive Impairment through ATF6-Herp Signaling. Scientific reports 39 29581524
2012 Muscle-specific splicing factors ASD-2 and SUP-12 cooperatively switch alternative pre-mRNA processing patterns of the ADF/cofilin gene in Caenorhabditis elegans. PLoS genetics 38 23071450
1989 Pharmacological characterization of the novel helodermin/VIP receptor present in human SUP-T1 lymphoma cell membranes. Regulatory peptides 38 2552509
2014 Ube2g2-gp78-mediated HERP polyubiquitylation is involved in ER stress recovery. Journal of cell science 37 24496447
2003 The ubiquitin-like domain of Herp is involved in Herp degradation, but not necessary for its enhancement of amyloid beta-protein generation. FEBS letters 37 14550564
2022 Corosolic acid sensitizes ferroptosis by upregulating HERPUD1 in liver cancer cells. Cell death discovery 35 36038536
2017 Electrospun Template Architecture and Composition Regulate Neutrophil NETosis In Vitro and In Vivo<sup/>. Tissue engineering. Part A 35 28068879
2017 Reactive Oxygen Species Shielding Hydrogel for the Delivery of Adherent and Nonadherent Therapeutic Cell Types<sup/>. Tissue engineering. Part A 35 28394196
2017 Stromal Cell-Derived Factor-1 Accelerates Cartilage Defect Repairing by Recruiting Bone Marrow Mesenchymal Stem Cells and Promoting Chondrogenic Differentiation<sup/>. Tissue engineering. Part A 35 28478702
2010 Water stress induces up-regulation of DOF1 and MIF1 transcription factors and down-regulation of proteins involved in secondary metabolism in amaranth roots (Amaranthus hypochondriacus L.). Plant biology (Stuttgart, Germany) 34 21489098
2018 The catalytic mechanism of electron-bifurcating electron transfer flavoproteins (ETFs) involves an intermediary complex with NAD<sup/>. The Journal of biological chemistry 33 30567738
2017 Construction of Three-Dimensional Dermo-Epidermal Skin Equivalents Using Cell Coating Technology and Their Utilization as Alternative Skin for Permeation Studies and Skin Irritation Tests<sup/>. Tissue engineering. Part A 33 28335694
1992 Mutations in the sup-38 gene of Caenorhabditis elegans suppress muscle-attachment defects in unc-52 mutants. Genetics 33 1427037
1984 The sup-7(st5) X gene of Caenorhabditis elegans encodes a tRNATrpUAG amber suppressor. Proceedings of the National Academy of Sciences of the United States of America 32 6093119
2018 Adjunctive S-adenosylmethionine (SAMe) in treating non-remittent major depressive disorder: An 8-week double-blind, randomized, controlled trial<sup/>. European neuropsychopharmacology : the journal of the European College of Neuropsychopharmacology 31 30115553
2017 MiR-9-3p augments apoptosis induced by H2O2 through down regulation of Herpud1 in glioma. PloS one 30 28430789
2013 Herp depletion protects from protein aggregation by up-regulating autophagy. Biochimica et biophysica acta 30 24120520
1981 Selective effects of alpha-MSH and MIF-1 on the blood-brain barrier. Peptides 30 6117842
2007 From MIF-1 to endomorphin: the Tyr-MIF-1 family of peptides. Peptides 29 17988762
1986 A visible allele of the muscle gene sup-10X of C. elegans. Genetics 29 3710144
2018 Application of combined porous tantalum scaffolds loaded with bone morphogenetic protein 7 to repair of osteochondral defect in rabbits<sup/>. International orthopaedics 28 29445961
2018 Knockdown of Herp alleviates hyperhomocysteinemia mediated atherosclerosis through the inhibition of vascular smooth muscle cell phenotype switching. International journal of cardiology 28 30017525
2017 In Vivo Immune Responses of Cross-Linked Electrospun Tilapia Collagen Membrane<sup/>. Tissue engineering. Part A 28 28375819
2006 Homocysteine regulates expression of Herp by DNA methylation involving the AARE and CREB binding sites. Experimental cell research 28 17020760
2021 The P300/XBP1s/Herpud1 axis promotes macrophage M2 polarization and the development of choroidal neovascularization. Journal of cellular and molecular medicine 27 34057287
2014 RBFOX and SUP-12 sandwich a G base to cooperatively regulate tissue-specific splicing. Nature structural & molecular biology 27 25132178
2013 A deficiency of Herp, an endoplasmic reticulum stress protein, suppresses atherosclerosis in ApoE knockout mice by attenuating inflammatory responses. PloS one 27 24204574
2001 Mif1: a missing link between the unfolded protein response pathway and ER-associated protein degradation? Current protein & peptide science 26 12370023
1989 Bacillus subtilis mutant allele sup-3 causes lysine insertion at ochre codons: use of sup-3 in studies of translational attenuation. Journal of bacteriology 26 2507520
2017 Bone Morphogenetic Protein 2 Alters Osteogenesis and Anti-Inflammatory Profiles of Mesenchymal Stem Cells Induced by Microtextured Titanium In Vitro<sup/>. Tissue engineering. Part A 24 28351289
2017 MicroRNA-384-mediated Herpud1 upregulation promotes angiotensin II-induced endothelial cell apoptosis. Biochemical and biophysical research communications 24 28483519
2010 Deletion of Herp facilitates degradation of cytosolic proteins. Genes to cells : devoted to molecular & cellular mechanisms 24 20604806
1998 Protection against MPTP treatment by an analog of Pro-Leu-Gly-NH2 (PLG, MIF-1). Peptides 24 9493876
2022 Role of MIF1/MIF2/CD74 interactions in bladder cancer. The Journal of pathology 23 36214539
2017 Biomaterial Encapsulation Is Enhanced in the Early Stages of the Foreign Body Reaction During Conditional Macrophage Depletion in Transgenic Macrophage Fas-Induced Apoptosis Mice<sup/>. Tissue engineering. Part A 23 28090808
2017 Activation of Macrophages by Lipopolysaccharide for Assessing the Immunomodulatory Property of Biomaterials<sup/>. Tissue engineering. Part A 23 28346799
2017 Herpud1 negatively regulates pathological cardiac hypertrophy by inducing IP3 receptor degradation. Scientific reports 23 29042597
2010 Somatic sex determination in Caenorhabditis elegans is modulated by SUP-26 repression of tra-2 translation. Proceedings of the National Academy of Sciences of the United States of America 23 20921392
2017 Extrinsic Calcitonin Gene-Related Peptide Inhibits Hyperoxia-Induced Alveolar Epithelial Type II Cells Apoptosis, Oxidative Stress, and Reactive Oxygen Species (ROS) Production by Enhancing Notch 1 and Homocysteine-Induced Endoplasmic Reticulum Protein (HERP) Expression. Medical science monitor : international medical journal of experimental and clinical research 22 29206808
2014 Targeting miR-21 sensitizes Ph+ ALL Sup-b15 cells to imatinib-induced apoptosis through upregulation of PTEN. Biochemical and biophysical research communications 22 25451263
1996 Characterization of the VIP receptor from SUP T1 lymphoblasts. Advances in neuroimmunology 22 8790781
2022 MIF1 and MIF2 Myostatin Peptide Inhibitors as Potent Muscle Mass Regulators. International journal of molecular sciences 21 35457038
2022 HERPUD1 promotes ovarian cancer cell survival by sustaining autophagy and inhibit apoptosis via PI3K/AKT/mTOR and p38 MAPK signaling pathways. BMC cancer 20 36544104
2014 Backbone-independent nucleic acid binding by splicing factor SUP-12 reveals key aspects of molecular recognition. Nature communications 20 25183497
2013 A novel mechanism regulating insulin secretion involving Herpud1 in mice. Diabetologia 20 23620059
1989 VIP and related peptides induce rapid homologous desensitization in the human lymphoma SUP T1 cell line. Peptides 20 2474155
2020 Circ_002664/miR-182-5p/Herpud1 pathway importantly contributes to OGD/R-induced neuronal cell apoptosis. Molecular and cellular probes 19 32376213
2018 A new role for HERPUD1 and ERAD activation in osteoblast differentiation and mineralization. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 19 29570393
2017 HERP Binds TBK1 To Activate Innate Immunity and Repress Virus Replication in Response to Endoplasmic Reticulum Stress. Journal of immunology (Baltimore, Md. : 1950) 19 28954889
2011 Coordinated conditional simulation with SLINK and SUP of many markers linked or associated to a trait in large pedigrees. Human heredity 19 21734403
1987 MIF-1 and Tyr-MIF-1 augment GABA-stimulated benzodiazepine receptor binding. Peptides 19 2893356

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