Affinage

EMG1

Ribosomal RNA small subunit methyltransferase NEP1 · UniProt Q92979

Length
244 aa
Mass
26.7 kDa
Annotated
2026-06-09
77 papers in source corpus 18 papers cited in narrative 18 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

EMG1 (Nep1) is an essential nucleolar SPOUT-class methyltransferase that functions in biogenesis of the small (40S) ribosomal subunit, where its depletion selectively reduces 20S pre-rRNA and mature 18S rRNA (PMID:11694595, PMID:18063569). Enzymatically it is a pseudouridine-N1-specific methyltransferase that transfers a methyl group to the N1 of pseudouridine, generating the m1acp3-Ψ hypermodification at Ψ1191 of 18S rRNA in yeast (Ψ926 in archaeal 16S rRNA), with catalysis depending on a conserved active-site aspartate acting as a base (PMID:20047967, PMID:20972225, PMID:39918246, PMID:37649713). Structures show EMG1 adopts an alpha/beta knot fold with a preformed SAM-binding pocket, acts as a homodimer, and recognizes its target via base-specific contacts that remodel the bound rRNA stem-loop (PMID:18063569, PMID:18208838, PMID:21087996). EMG1 has a catalytically separable, essential role in ribosome assembly, since mutations that abolish SAM binding or block Ψ1191 methylation do not prevent ribosome biogenesis or growth (PMID:18063569, PMID:20972225). It is imported into the nucleus by Impα/β or Impβ/7, which also chaperone its basic regions against nonspecific RNA aggregation, and is recruited to the nucleolus within a subcomplex containing NOP14, NOC4L, and UTP14A (PMID:27798105). Beyond ribosome assembly, EMG1 is required for early embryonic development and nucleologenesis (PMID:20858271), cooperates with the transcription factor GRHL3 to drive nuclear GRHL3 localization and Wnt/β-catenin–mediated surface ectoderm differentiation (PMID:40761126), and its methyltransferase activity supports KSHV-induced specialized ribosome scanning of viral uORFs (PMID:40215162). The D86G missense mutation causes Bowen-Conradi syndrome; it does not abolish catalytic activity but enhances dimerization and RNA affinity, promotes aggregation and proteasomal degradation, and impairs nucleolar accumulation (PMID:19463982, PMID:20972225, PMID:27798105).

Mechanistic history

Synthesis pass · year-by-year structured walk · 11 steps
  1. 2001 High

    Established EMG1 as a 40S ribosomal subunit biogenesis factor whose nuclear function depends on a defined protein partner, framing it as a nucleolar assembly component rather than a free-standing enzyme.

    Evidence Genetic depletion, ts allele and rRNA Northern blotting in yeast with Nop14 interaction and localization assays

    PMID:11694595

    Open questions at the time
    • Molecular activity unknown at this stage
    • Did not define which pre-rRNA processing step requires EMG1
  2. 2007 High

    Defined EMG1 as a SPOUT-class methyltransferase by structure and, critically, showed that catalytic activity is dispensable for ribosome biogenesis, revealing a separable non-catalytic essential role.

    Evidence X-ray crystallography of yeast Emg1 with SAM, SAM- and RNA-binding mutagenesis, yeast complementation; archaeal Nep1 structures with cofactor analogs

    PMID:18063569 PMID:18208838

    Open questions at the time
    • RNA substrate and methylated position not yet identified
    • Nature of the catalysis-independent essential function unresolved
  3. 2010 High

    Identified the precise enzymatic activity and target: N1-methylation of pseudouridine at Ψ1191 of 18S rRNA, and confirmed a dual catalytic / assembly-factor function.

    Evidence In vitro methyltransferase assays with MS/HPLC/NMR, in vivo 14C-methionine labeling, ESI-MS, and genetic suppressor/SAM rescue in yeast; protein-RNA co-crystal structures

    PMID:20047967 PMID:20972225 PMID:21087996

    Open questions at the time
    • Mechanism by which assembly function is separable from methylation not defined
    • rRNA remodeling role inferred from static structures
  4. 2010 High

    Demonstrated organismal essentiality, showing EMG1 loss arrests early mouse development with nucleologenesis defects in a p53-independent manner.

    Evidence Mouse knockout, embryo phenotyping, genetic epistasis with p53 null

    PMID:20858271

    Open questions at the time
    • Did not resolve whether lethality reflects catalytic or assembly role
    • Cell-type-specific requirements not addressed
  5. 2009 Medium

    Linked EMG1 to human disease by identifying D86G as the cause of Bowen-Conradi syndrome and indicating a protein-stability/aggregation defect.

    Evidence Patient sequencing, Western blotting, yeast two-hybrid, mammalian overexpression, structural modeling

    PMID:19463982

    Open questions at the time
    • Aggregation inferred indirectly from solubility and two-hybrid data
    • Structural modeling is computational, not experimental
  6. 2012 Medium

    Placed EMG1 within a genetic network for early pre-ribosome assembly, linking it functionally to Utp30 and the surface for Rps19 loading onto the 90S pre-ribosome.

    Evidence Genome-wide synthetic genetic array with biochemical follow-up and complementation in yeast

    PMID:22588997

    Open questions at the time
    • Genetic interactions do not establish direct physical contacts
    • Single-lab screen without orthogonal validation of all hits
  7. 2016 High

    Resolved how EMG1 reaches and is protected within the nucleolus, defining importin-mediated import/chaperoning and a NOP14/NOC4L/UTP14A subcomplex required for nucleolar recruitment, and recast D86G pathology as a trafficking/degradation defect.

    Evidence Nuclear import and in vitro aggregation assays, Co-IP, proteasome inhibition, fluorescence microscopy

    PMID:27798105

    Open questions at the time
    • Stoichiometry and assembly order of the subcomplex not defined
    • Conflicts with patient-cell data on D86G nucleolar localization
  8. 2014 Medium

    Characterized cellular consequences of D86G in patient cells, showing delayed 18S processing and G2/M accumulation despite normal steady-state 40S and translation, indicating a partial/kinetic defect.

    Evidence Flow cytometry, proliferation and pulse-chase rRNA processing assays, immunofluorescence in BCS lymphoblasts

    PMID:26676230

    Open questions at the time
    • Reports normal D86G nucleolar localization, conflicting with overexpression studies
    • Mechanism connecting processing delay to G2/M arrest unresolved
  9. 2023 Medium

    Defined the chemical catalytic mechanism, identifying a conserved aspartate as the catalytic base for pseudouridine N1 deprotonation and a water-mediated proton-transfer pathway.

    Evidence Molecular dynamics, DFT calculations, site-directed mutagenesis and kinetic assays

    PMID:37649713

    Open questions at the time
    • Mechanism is largely computational and not independently replicated
    • Catalytic detail does not address the non-catalytic assembly role
  10. 2025 High

    Extended EMG1 function beyond ribosome assembly into development and viral biology, showing a GRHL3 partnership driving Wnt/β-catenin–dependent ectoderm/neural tube morphogenesis and a methyltransferase-dependent role in KSHV specialized ribosomes; also added a fresh archaeal structure with defined RNA target.

    Evidence Conditional knockout mice with protein complex identification and epistasis (GRHL3); EMG1 depletion with ribosome occupancy and viral output assays (KSHV); P. horikoshii crystal structures with in vitro methyltransferase assays

    PMID:39918246 PMID:40215162 PMID:40761126

    Open questions at the time
    • How a ribosome assembly factor controls GRHL3 nuclear localization mechanistically is unclear
    • Whether developmental roles require catalysis vs assembly function not separated
    • Generality of specialized-ribosome role beyond KSHV unknown
  11. 2025 Medium

    Tied EMG1 abundance to a cancer-relevant regulatory axis, showing FTO-mediated m6A demethylation of EMG1 mRNA reduces EMG1, ribosome biosynthesis, and bladder tumor cell aggressiveness.

    Evidence FTO gain/loss-of-function, m6A analysis, in vitro and in vivo bladder tumor models

    PMID:41962773

    Open questions at the time
    • Direct m6A site on EMG1 mRNA and reader involvement not fully defined
    • Causality between EMG1 level and tumor phenotype is correlative in part

Open questions

Synthesis pass · forward-looking unresolved questions
  • The molecular basis of EMG1's catalysis-independent essential ribosome assembly function, and how this is mechanistically distinct from its non-ribosomal roles in GRHL3-mediated development and specialized translation, remains unresolved.
  • No structure of EMG1 within an intact pre-40S particle
  • No separation-of-function alleles distinguishing assembly vs developmental roles
  • Mechanism linking nucleolar assembly factor to nuclear transcription-factor function unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140098 catalytic activity, acting on RNA 4 GO:0003723 RNA binding 3 GO:0016740 transferase activity 3
Localization
GO:0005730 nucleolus 3 GO:0005634 nucleus 2
Pathway
R-HSA-1266738 Developmental Biology 2 R-HSA-162582 Signal Transduction 2 R-HSA-8953854 Metabolism of RNA 2
Partners
GRHL3NOC4LNOP14RPS19UTP14AUTP30
Complex memberships
EMG1-GRHL3 complexEMG1/NOP14/NOC4L/UTP14A nucleolar subcomplex

Evidence

Reading pass · 18 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2001 EMG1 (Emg1) is required for biogenesis of the 40S ribosomal subunit in yeast; depletion causes selectively reduced levels of 20S pre-rRNA and mature 18S rRNA. Nuclear localization of Emg1 depends on physical interaction with the nucleolar protein Nop14. Genetic depletion, temperature-sensitive allele, rRNA Northern blotting, nuclear localization assay Molecular biology of the cell High 11694595
2007 Crystal structure of yeast Emg1 at 2 Å in complex with SAM identifies it as a novel member of the SPOUT-class (alpha/beta knot fold) methyltransferase superfamily. A point mutation in a basic patch abolished RNA binding in vitro; mutations disrupting SAM binding caused >100-fold reduction in SAM binding but did not prevent growth or ribosome biogenesis, indicating catalytic activity is not essential for ribosome biogenesis. X-ray crystallography, in vitro SAM binding assays, RNA binding assay, yeast complementation Nucleic acids research High 18063569
2008 Crystal structure of archaeal Nep1 (Methanocaldococcus jannaschii) in free, SAH-bound, and sinefungin-bound forms reveals an extended SPOUT-class methyltransferase fold with a preformed, pre-organized SAM-binding site topologically equivalent to other SPOUT methyltransferases. X-ray crystallography (2.15–2.25 Å resolution), ligand co-crystallization Nucleic acids research High 18208838
2010 Nep1 (Emg1) is a pseudouridine-N1-specific methyltransferase: it selectively methylates RNAs containing a pseudouridine at a position corresponding to the hypermodified m1acp3-Ψ in eukaryotic 18S rRNA, transferring the methyl group to the N1 of pseudouridine. This enzymatic activity is conserved in human NEP1. Fluorescence spectroscopy, NMR spectroscopy, MALDI-MS, HPLC, yeast three-hybrid screening for RNA-binding specificity, in vitro methyltransferase assay Nucleic acids research High 20047967
2010 In yeast, Nep1 catalyzes in vivo methylation of pseudouridine at position 1191 (Ψ1191) within loop 35 of 18S rRNA, the site of the hypermodification m1acp3Ψ. Nep1 is not required for the acp-modification but provides the Ψ1191 methylation. Nep1 has a dual function: as Ψ1191-methyltransferase and as an essential ribosome assembly factor, since restored growth of nep1-1(ts) by SAM addition was observed even when Ψ1191 methylation was prevented. The Bowen-Conradi D86G mutation does not affect methyltransferase activity but enhances dimerization and RNA-target affinity, and prevents nucleolar accumulation of Nep1. 14C-methionine labeling of 18S rRNA in yeast mutants, ESI-MS analysis of acp-modified Ψ-nucleosides in Δnep1 mutants, genetic suppressor analysis, SAM rescue experiments Nucleic acids research High 20972225
2009 A D86G mutation in human EMG1 causes Bowen-Conradi syndrome. In BCS patient fibroblasts, EMG1 protein is dramatically reduced without change in mRNA levels. Overexpression of EMG1-D86G in mammalian cells decreases soluble EMG1, and yeast two-hybrid analysis shows D86G increases interaction between EMG1 subunits, suggesting the mutation causes aggregation. Patient sequencing, Western blotting, yeast two-hybrid, mammalian overexpression, structural modeling American journal of human genetics Medium 19463982
2010 Crystal structures of S. cerevisiae Nep1 dimer and its RNA complexes show Nep1 recognizes its RNA site via base-specific interactions, stabilizes a stem-loop in bound RNA, and the bound RNA structure contradicts predicted rRNA secondary structures, suggesting Nep1 remodels rRNA upon binding. A uridine base is positioned in the active site for methyltransfer at C5, supporting N1-specific pseudouridine methyltransferase activity. Mutations reducing methyl-donor binding are not lethal. X-ray crystallography of protein-RNA complexes, mutagenesis Nucleic acids research High 21087996
2010 EMG1 knockout in mice arrests embryonic development prior to the blastocyst stage with defects in early cell lineage-specification and nucleologenesis. Loss of p53 failed to rescue Emg1-/- pre-implantation lethality, unlike some other ribosome biogenesis defects. Mouse gene targeting (knockout), embryo phenotyping, genetic epistasis with p53 null BMC developmental biology High 20858271
2012 Genetic interaction screen in yeast identified that NEP1 (EMG1) interacts genetically with ribosome biogenesis genes RPS18A/B, RRP8, EFG1, UTP30, ribosome quality control genes UBP3, BRE5, UBP6, and no-go decay factor DOM34. UTP30 deletion enforces all nep1-1(ts) phenotypes; UTP30 overexpression partially restores nep1-1(ts) growth; genetic and biochemical data suggest Utp30 and Nep1 act together during pre-ribosomal complex formation and provide the surface for Rps19 assembly to the 90S pre-ribosome. Genome-wide synthetic genetic array (SGA), biochemical co-analysis, yeast complementation Yeast (Chichester, England) Medium 22588997
2007 Tma23 and Nop6 mutations act as recessive suppressors of nep1(ts) and nep1 deletion in yeast. GFP fusions localized both proteins to the nucleolus, supporting their role in ribosome biogenesis. Both proteins have high lysine content; Nop6 contains an RNA-binding motif, suggesting RNA-binding functions. Suppressor screen, GFP localization, genetic complementation FEMS yeast research Medium 17425675
2016 EMG1 is imported into the nucleus by importins Impα/β or Impβ/7. Binding of the Impβ/7 heterodimer prevents unspecific aggregation of both EMG1 and EMG1-D86G on RNAs in vitro, indicating importins act as chaperones for EMG1 by binding to its basic regions. The BCS D86G mutation leads to reduced nucleolar localization, accumulation of EMG1-D86G in nuclear foci, and proteasome-dependent degradation. Pre-ribosomal factors NOP14, NOC4L, and UTP14A form a nucleolar subcomplex containing EMG1 required for its nucleolar recruitment. Nuclear import assays, Co-immunoprecipitation, in vitro aggregation assay, inhibitor treatments (proteasome inhibitor), fluorescence microscopy Human molecular genetics High 27798105
2014 In BCS patient lymphoblasts (D86G mutation), cells accumulate in G2/M resulting in reduced proliferation, and 18S rRNA processing is consistently delayed, though levels of 40S ribosomes and protein synthesis rates are not different from controls. The D86G substitution does not interfere with EMG1 nucleolar localization in mammalian cells. Cell cycle analysis (flow cytometry), proliferation assays, pulse-chase rRNA processing analysis, Western blotting, immunofluorescence BBA clinical Medium 26676230
2020 EMG1 interacts with NOP14 as shown by GST pulldown and co-immunoprecipitation. Together, EMG1 and NOP14 regulate cell growth, apoptosis, migration and invasion in melanoma cells, and simultaneous overexpression decreases WNT3a, β-catenin, phospho-GSK-3β, and c-Myc levels, implicating the Wnt/β-catenin pathway. GST pulldown, co-immunoprecipitation, overexpression, Western blotting Translational cancer research Medium 35117729
2025 EMG1 forms a protein complex with the transcription factor GRHL3 and is required for correct nuclear localization of GRHL3 and activation of the canonical Wnt/β-catenin signaling pathway. Conditional knockout of Emg1 in the GRHL3-positive surface ectoderm causes spina bifida (neural tube defects). Compound mutant phenotypes of Emg1 and Grhl3 indicate genetic interaction in neurulation and palate development. Conditional knockout mouse model, protein complex identification, immunofluorescence, genetic epistasis analysis Development (Cambridge, England) High 40761126
2025 EMG1 methyltransferase activity is required for KSHV-induced specialized ribosomes to scan through KSHV upstream open reading frames (uORFs) in late lytic genes. During KSHV lytic replication, EMG1 has enhanced association with precursor-40S ribosome complexes. Depletion of EMG1 reduces viral protein expression and infectious virus production. EMG1 depletion (knockdown), ribosome profiling/occupancy assays, viral replication assays, precursor-40S complex immunoprecipitation Cell reports High 40215162
2023 Molecular dynamics simulations and quantum-chemical calculations, combined with kinetic and mutational experiments, elucidate the catalytic mechanism of Nep1: a conserved aspartate (D101 in S. cerevisiae) is essential for catalysis, acting as a base for deprotonation of pseudouridine N1. A conserved hydroxyl-containing residue (S233 in S. cerevisiae) and active-site water molecules can mediate proton transfer. The water-mediated proton transfer pathway is the most energetically favorable. Mutation of D101 abolishes activity; mutation of S233 does not. Molecular dynamics simulation, DFT quantum-chemical calculations, site-directed mutagenesis, kinetic assays Computational and structural biotechnology journal Medium 37649713
2025 Crystal structures of Pyrococcus horikoshii Nep1 (PhNep1) in apo, adenosine-bound, and 5-methylthioadenosine-bound forms reveal an α/β SPOUT fold with a trefoil knot, two novel extensions (a globular loop and a β-α-β extension), a preformed cofactor-binding pocket, and a homodimer stabilized by inter-subunit hydrogen bonds and hydrophobic interactions. PhNep1 specifically methylates a pseudouridine at position 926 of helix 35 in 16S rRNA; conserved arginine residues at the dimeric interface assist RNA complex stability. X-ray crystallography (apo and holo forms), in vitro methyltransferase assay with consensus RNA substrates The FEBS journal High 39918246
2025 FTO-mediated m6A demethylation of EMG1 mRNA decreases EMG1 protein expression and reduces ribosome biosynthesis, suppressing bladder cancer cell proliferation, migration, and invasion in vitro and in vivo. FTO overexpression/knockdown, m6A modification analysis, in vitro and in vivo tumor models, Western blotting Biochimica et biophysica acta. Molecular cell research Medium 41962773

Source papers

Stage 0 corpus · 77 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Phytotoxicity and innate immune responses induced by Nep1-like proteins. The Plant cell 232 17194768
2006 Nep1-like proteins from plant pathogens: recruitment and diversification of the NPP1 domain across taxa. Phytochemistry 208 16430931
2013 Evidence for functional diversification within a fungal NEP1-like protein family. Molecular plant-microbe interactions : MPMI 125 23051172
2010 The Bowen-Conradi syndrome protein Nep1 (Emg1) has a dual role in eukaryotic ribosome biogenesis, as an essential assembly factor and in the methylation of Ψ1191 in yeast 18S rRNA. Nucleic acids research 109 20972225
2010 The ribosome assembly factor Nep1 responsible for Bowen-Conradi syndrome is a pseudouridine-N1-specific methyltransferase. Nucleic acids research 107 20047967
2009 Mutation of a gene essential for ribosome biogenesis, EMG1, causes Bowen-Conradi syndrome. American journal of human genetics 98 19463982
2001 Novel stress-responsive genes EMG1 and NOP14 encode conserved, interacting proteins required for 40S ribosome biogenesis. Molecular biology of the cell 89 11694595
2007 Phytotoxic Nep1-like proteins from the necrotrophic fungus Botrytis cinerea associate with membranes and the nucleus of plant cells. The New phytologist 86 18028294
2007 Nogo-66 receptor antagonist peptide (NEP1-40) administration promotes functional recovery and axonal growth after lateral funiculus injury in the adult rat. Neurorehabilitation and neural repair 80 18056009
2014 Comparative and functional analysis of the widely occurring family of Nep1-like proteins. Molecular plant-microbe interactions : MPMI 77 25025781
2019 Activity and Phylogenetics of the Broadly Occurring Family of Microbial Nep1-Like Proteins. Annual review of phytopathology 73 31283435
2012 Nontoxic Nep1-like proteins of the downy mildew pathogen Hyaloperonospora arabidopsidis: repression of necrosis-inducing activity by a surface-exposed region. Molecular plant-microbe interactions : MPMI 59 22235872
2010 Structural insight into the functional mechanism of Nep1/Emg1 N1-specific pseudouridine methyltransferase in ribosome biogenesis. Nucleic acids research 57 21087996
2007 The yeast ribosome synthesis factor Emg1 is a novel member of the superfamily of alpha/beta knot fold methyltransferases. Nucleic acids research 53 18063569
2008 Trans-activator of transcription-mediated delivery of NEP1-40 protein into brain has a neuroprotective effect against focal cerebral ischemic injury via inhibition of neuronal apoptosis. Anesthesiology 47 18497608
1996 DNA bending by the silencer protein NeP1 is modulated by TR and RXR. Nucleic acids research 42 8758989
2014 Characterization of the lipid binding properties of Otoferlin reveals specific interactions between PI(4,5)P2 and the C2C and C2F domains. Biochemistry 41 24999532
2016 Effects of the Bowen-Conradi syndrome mutation in EMG1 on its nuclear import, stability and nucleolar recruitment. Human molecular genetics 40 27798105
2008 The crystal structure of Nep1 reveals an extended SPOUT-class methyltransferase fold and a pre-organized SAM-binding site. Nucleic acids research 40 18208838
1993 NeP1. A ubiquitous transcription factor synergizes with v-ERBA in transcriptional silencing. Journal of molecular biology 38 8102652
2017 The Nep1-like protein family of Magnaporthe oryzae is dispensable for the infection of rice plants. Scientific reports 37 28663588
2012 The Nicotiana benthamiana mitogen-activated protein kinase cascade and WRKY transcription factor participate in Nep1(Mo)-triggered plant responses. Molecular plant-microbe interactions : MPMI 32 22835275
2005 Developmental expression of stress response genes in Theobroma cacao leaves and their response to Nep1 treatment and a compatible infection by Phytophthora megakarya. Plant physiology and biochemistry : PPB 29 15979314
2005 Caenorhabditis elegans neprilysin NEP-1: an effector of locomotion and pharyngeal pumping. Journal of molecular biology 27 16081104
2010 TAT-NEP1-40 as a novel therapeutic candidate for axonal regeneration and functional recovery after stroke. Journal of drug targeting 25 20367026
2021 Type 2 Nep1-Like Proteins from the Biocontrol Oomycete Pythium oligandrum Suppress Phytophthora capsici Infection in Solanaceous Plants. Journal of fungi (Basel, Switzerland) 24 34206578
2002 Expression of NEP1 by Fusarium oxysporum f. sp. erythroxyli After Gene Replacement and Overexpression Using Polyethylene Glycol-Mediated Transformation. Phytopathology 24 18942961
2019 Molecular basis for functional diversity among microbial Nep1-like proteins. PLoS pathogens 23 31479498
2009 Inducible expression of a Nep1-like protein serves as a model trigger system of camalexin biosynthesis. Phytochemistry 23 19155026
2005 NEP1 orthologs encoding necrosis and ethylene inducing proteins exist as a multigene family in Phytophthora megakarya, causal agent of black pod disease on cacao. Mycological research 23 16353637
2010 EMG1 is essential for mouse pre-implantation embryo development. BMC developmental biology 22 20858271
2011 Expression of an oxalate decarboxylase impairs the necrotic effect induced by Nep1-like protein (NLP) of Moniliophthora perniciosa in transgenic tobacco. Molecular plant-microbe interactions : MPMI 21 21405988
2005 Nep1, a Schizosaccharomyces pombe deneddylating enzyme. The Biochemical journal 19 15769255
2021 Nep1-like proteins as a target for plant pathogen control. PLoS pathogens 18 33857257
2012 Genetic interactions of yeast NEP1 (EMG1), encoding an essential factor in ribosome biogenesis. Yeast (Chichester, England) 18 22588997
2007 Mutations in the nucleolar proteins Tma23 and Nop6 suppress the malfunction of the Nep1 protein. FEMS yeast research 17 17425675
2022 Disruption of plant plasma membrane by Nep1-like proteins in pathogen-plant interactions. The New phytologist 16 36210522
2020 Identification and Characterization of Nep1-Like Proteins From the Grapevine Downy Mildew Pathogen Plasmopara viticola. Frontiers in plant science 16 32117400
2013 Nep1-like protein from Moniliophthora perniciosa induces a rapid proteome and metabolome reprogramming in cells of Nicotiana benthamiana. Physiologia plantarum 16 23586401
2014 Mutation of EMG1 causing Bowen-Conradi syndrome results in reduced cell proliferation rates concomitant with G2/M arrest and 18S rRNA processing delay. BBA clinical 15 26676230
2021 Functional Characterization of the Nep1-Like Protein Effectors of the Necrotrophic Pathogen - Alternaria brassicae. Frontiers in microbiology 14 34764943
2019 Necrotic and Cytolytic Activity on Grapevine Leaves Produced by Nep1-Like Proteins of Diplodia seriata. Frontiers in plant science 13 31749815
2022 Cytotoxic activity of Nep1-like proteins on monocots. The New phytologist 12 35383933
2021 NEP1‑40 promotes myelin regeneration via upregulation of GAP‑43 and MAP‑2 expression after focal cerebral ischemia in rats. Molecular medicine reports 12 34643252
2021 Nep1-like Proteins from Valsa mali Differentially Regulate Pathogen Virulence and Response to Abiotic Stresses. Journal of fungi (Basel, Switzerland) 12 34682251
2019 Otoferlin C2F Domain-Induced Changes in Membrane Structure Observed by Sum Frequency Generation. Biophysical journal 12 31587832
2016 Comparison of RNAi NgR and NEP1-40 in Acting on Axonal Regeneration After Spinal Cord Injury in Rat Models. Molecular neurobiology 12 27921243
2000 Nep1 Protein from Fusarium oxysporum Enhances Biological Control of Opium Poppy by Pleospora papaveracea. Phytopathology 12 18944501
2020 NEP1-40 alleviates behavioral phenotypes and promote oligodendrocyte progenitor cell differentiation in the hippocampus of cuprizone-induced demyelination mouse model. Neuroscience letters 11 32112820
2022 Functional analysis of the Nep1-like proteins from Plasmopara viticola. Plant signaling & behavior 9 35152834
2021 NEP1-40-overexpressing Neural Stem Cells Enhance Axon Regeneration by Inhibiting Nogo-A/NgR1 Signaling Pathway. Current neurovascular research 9 34544340
2022 Stemphylium lycopersici Nep1-like Protein (NLP) Is a Key Virulence Factor in Tomato Gray Leaf Spot Disease. Journal of fungi (Basel, Switzerland) 8 35628773
2009 Backbone resonance assignments of the 48 kDa dimeric putative 18S rRNA-methyltransferase Nep1 from Methanocaldococcus jannaschii. Biomolecular NMR assignments 8 19779849
2022 Nep1-Like Proteins From the Biocontrol Agent Pythium oligandrum Enhance Plant Disease Resistance Independent of Cell Death and Reactive Oxygen Species. Frontiers in plant science 7 35310640
2021 PLGA-PEG-PLGA hydrogel with NEP1-40 promotes the functional recovery of brachial plexus root avulsion in adult rats. PeerJ 7 34760354
2011 Neuroprotective effects of NEP1-40 and fasudil on Nogo-A expression in neonatal rats with hypoxic-ischemic brain damage. Genetics and molecular research : GMR 7 22180032
2008 Cupin: a candidate molecular structure for the Nep1-like protein family. BMC plant biology 7 18447914
2008 Purification, crystallization and preliminary X-ray diffraction analysis of an oomycete-derived Nep1-like protein. Acta crystallographica. Section F, Structural biology and crystallization communications 7 19052381
2020 EMG1 interacts with NOP14 to regulate the growth, migration, and invasion of melanoma cells via the Wnt/β-catenin pathway. Translational cancer research 6 35117729
2013 Molecular characterization and functional analysis of the Nep1-like protein-encoding gene from Phytophthora capsici. Genetics and molecular research : GMR 6 23661469
2008 Nogo-A inhibitory peptide (NEP1-40) increases pan-cadherin expression following mild cortical contusion injury in rats. Turkish neurosurgery 6 19107681
2018 l-Theanine and NEP1-40 promote nerve regeneration and functional recovery after brachial plexus root avulsion. Biochemical and biophysical research communications 5 30553451
2023 Nucleolar Essential Protein 1 (Nep1): Elucidation of enzymatic catalysis mechanism by molecular dynamics simulation and quantum mechanics study. Computational and structural biotechnology journal 4 37649713
2023 Probing the role of the C2F domain of otoferlin. Frontiers in molecular neuroscience 4 38152587
2009 Transforming a NEP1 toxin gene into two Fusarium spp. to enhance mycoherbicide activity on Orobanche--failure and success. Pest management science 4 19291699
2025 EMG1 methyltransferase activity affects ribosome occupancy at KSHV uORFs. Cell reports 3 40215162
2025 Improving neuronal recovery in spinal cord injury with NEP1-40-modified neural stem cells through RhoA/ROCK signaling pathway modulation. Biochimica et biophysica acta. Molecular basis of disease 2 40446939
2024 Interaction of an Oomycete Nep1-like Cytolysin with Natural and Plant Cell-Mimicking Membranes. The Journal of membrane biology 2 39692881
2022 Transplantation of NEP1-40 and NT-3 Gene-Co-Transduced Neural Stem Cells Improves Function and Neurogenesis after Spinal Cord Injury in a Rat Model. Neurology India 2 36412377
2025 Structural analysis of the ribosome assembly factor Nep1, an N1-specific pseudouridine methyltransferase, reveals mechanistic insights. The FEBS journal 1 39918246
2025 EMG1 cooperates with GRHL3 in β-catenin-mediated surface ectoderm differentiation to regulate neural tube closure. Development (Cambridge, England) 1 40761126
2025 Plasticity of the cytotoxic Nep1-like protein enables promiscuity in binding to its lipid receptor glycosylinositol phosphorylceramides. Science advances 1 41071873
2026 FTO-mediated m6A demethylation inhibits bladder cancer progression via decreasing EMG1 and reducing ribosome biosynthesis. Biochimica et biophysica acta. Molecular cell research 0 41962773
2025 NEP1-40 Regulates the Development of Hippocampal Neural Stem Cells in Schizophrenic Mice. Current medical science 0 40643877
2023 [Corrigendum] NEP1‑40 promotes myelin regeneration via upregulation of GAP‑43 and MAP‑2 expression after focal cerebral ischemia in rats. Molecular medicine reports 0 36695184
2010 [Expression of Nogo receptor in brain and neuroprotective effect of NEP1-40 on hypoxic ischemic brain damage in newborn rats]. Zhonghua er ke za zhi = Chinese journal of pediatrics 0 20426940
2006 [Cloning of NEP1-40 gene and expression of its protein]. Zhongguo xiu fu chong jian wai ke za zhi = Zhongguo xiufu chongjian waike zazhi = Chinese journal of reparative and reconstructive surgery 0 16457436

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