Affinage

CTDNEP1

CTD nuclear envelope phosphatase 1 · UniProt O95476

Length
244 aa
Mass
28.4 kDa
Annotated
2026-06-09
16 papers in source corpus 15 papers cited in narrative 15 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 4/4 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

CTDNEP1 (Dullard) is a non-canonical serine/threonine phosphatase that operates at the inner nuclear and ER membrane to terminate TGF-β/BMP superfamily signaling and to regulate membrane lipid homeostasis (PMID:38776370, PMID:21413788, PMID:17141153). Its catalytic activity depends on the regulatory subunit NEP1R1, which binds at a site distant from the active site to allosterically stabilize and activate the enzyme, while a conserved Arg residue in CTDNEP1 orients substrate peptides for dephosphorylation; co-depletion of CTDNEP1 or NEP1R1 produces identical ER expansion phenotypes, defining them as an obligate functional complex (PMID:38776370). Together with the inner nuclear membrane scaffold MAN1, the CTDNEP1-NEP1R1 phosphatase dephosphorylates nuclear receptor-regulated SMADs, and disruption of this complex causes accumulation of phosphorylated R-SMADs and aberrant signaling [PMID:bio_10.1101_2024.09.23.614427]. This signal-terminating role is conserved and context-dependent: CTDNEP1 dephosphorylates Mad/R-SMADs and promotes BMP receptor turnover in flies and Xenopus (PMID:17141153, PMID:27578171), and across mouse tissues its loss elevates pSmad1/5/8 or pSmad2/3 to drive defects in kidney maintenance, endochondral ossification, cardiac outflow tract development, and ovarian function, each rescuable by BMP or TGF-β receptor kinase inhibition (PMID:23360989, PMID:25155999, PMID:32105214, PMID:29521016). Independent of signaling termination, CTDNEP1 dephosphorylates lipin1 at its insulin-dependent Ser106 site without requiring an adaptor (PMID:21413788), stabilizes the genome by limiting MYC Ser62 phosphorylation and modulating mitotic regulators including TOP2A and CHEK1 (PMID:36765089), and interacts with the actin regulator Eps8L2 to organize dorsal actin cables for nuclear positioning during cell migration (PMID:33567288).

Mechanistic history

Synthesis pass · year-by-year structured walk · 14 steps
  1. 2006 High

    Established that the CTDNEP1 ortholog Dullard restrains BMP signaling, linking its phosphatase activity to receptor turnover and a developmental output.

    Evidence Co-IP, phosphatase-dead mutant, and morpholino knockdown with neural marker readout in Xenopus

    PMID:17141153

    Open questions at the time
    • Direct substrate of the phosphatase not defined
    • Mechanism coupling phosphatase activity to receptor ubiquitination unresolved
  2. 2011 High

    Defined CTDNEP1 as a quantitative phosphoserine phosphatase with defined substrate specificity, separating its lipid-regulatory function from signaling roles.

    Evidence Steady-state kinetics on phosphoserine nonapeptides corresponding to lipin1 sites

    PMID:21413788

    Open questions at the time
    • Activity measured on peptides, not full-length lipin1
    • Cellular consequences of lipin1 dephosphorylation not addressed here
  3. 2011 Medium

    Connected Dullard's BMP-modulating role to nuclear envelope and membrane lipid homeostasis, hinting at a membrane-based mechanism for signaling control.

    Evidence Genetic interaction with BMP components, pMad western, Importin-β/RanGAP localization and lipid staining in Drosophila

    PMID:21790556

    Open questions at the time
    • Causal link between lipid changes and BMP output not established
    • Ortholog data may not transfer directly to human CTDNEP1
  4. 2013 High

    Demonstrated cell-autonomous BMP restraint by Dullard in mammalian organogenesis using pharmacological epistasis.

    Evidence Conditional knockout mouse with pSmad1/5/8 staining and LDN-193189 rescue in kidney

    PMID:23360989

    Open questions at the time
    • Whether Dullard acts directly on R-SMADs or upstream receptors not distinguished
    • Substrate identity in vivo unconfirmed
  5. 2013 Medium

    Showed Dullard can act as a positive modulator of a distinct pathway (WNT/β-catenin), revealing context-dependent signaling roles beyond BMP.

    Evidence Conditional KO, Wnt3 compound heterozygote epistasis, β-catenin reporter, Dvl2 western in germ cell specification

    PMID:23469192

    Open questions at the time
    • Mechanism by which Dullard supports Dvl2 levels unknown
    • Direct substrate in WNT pathway not identified
  6. 2015 High

    Extended Dullard's signaling control to the TGF-β/Smad2/3 branch and implicated regulation of R-SMAD protein stability.

    Evidence Prx1-Cre conditional KO, pSmad2/3 and total Smad2/3 western, LY-364947 rescue in skeletal development

    PMID:25155999

    Open questions at the time
    • Mechanism linking phosphatase to Smad2/3 protein stability unresolved
    • Direct dephosphorylation of Smad2/3 not shown biochemically
  7. 2016 High

    Provided direct biochemical evidence that Dullard dephosphorylates R-SMADs (Mad) at both C-terminal and linker sites, defining an enzymatic mode of signal termination distinct from degradation.

    Evidence Reciprocal Co-IP, phosphatase-dead mutant separating binding from activity, loss-of-function with pMad readout in Drosophila

    PMID:27578171

    Open questions at the time
    • Human R-SMAD substrate not directly demonstrated here
    • Role of regulatory subunits in this context not examined
  8. 2018 Medium

    Confirmed BMP-suppressive role in additional tissue via inducible deletion and pharmacological rescue.

    Evidence Inducible/conditional KO mouse, Smad6/7 expression, LDN-193189 rescue in ovarian granulosa cells

    PMID:29521016

    Open questions at the time
    • Direct molecular substrate not identified
    • Single-lab observation
  9. 2020 High

    Showed Dullard tunes BMP signaling amplitude to control cell-fate decisions, connecting phosphatase activity to downstream transcriptional programs.

    Evidence NCC-specific conditional KO, BMP reporter, pSmad1/5/8 staining, Sema3c/Twist1 expression in cardiac outflow tract

    PMID:32105214

    Open questions at the time
    • Direct substrate in this context not biochemically defined
    • How signaling amplitude is quantitatively set unclear
  10. 2021 Medium

    Revealed a non-signaling role for CTDNEP1 in cytoskeletal organization and nuclear positioning through a direct partner.

    Evidence Direct interaction assay with Eps8L2, siRNA knockdown with nuclear positioning, migration, and actin cable imaging

    PMID:33567288

    Open questions at the time
    • Whether phosphatase activity is required for the Eps8L2 function not established
    • Substrate in actin regulation unknown
  11. 2023 High

    Implicated CTDNEP1 in genome stability and oncogenesis by limiting MYC phosphorylation and modulating mitotic regulators.

    Evidence Ctdnep1 KO mouse tumor model, phosphoproteomics, pSer62-MYC western, chromosomal instability assays, co-targeting MYC/CHEK1

    PMID:36765089

    Open questions at the time
    • Whether CTDNEP1 directly dephosphorylates MYC Ser62 not shown
    • Direct vs indirect modulation of TOP2A/CHEK1 unresolved
  12. 2024 High

    Resolved the structural and mechanistic basis of CTDNEP1 activation, defining NEP1R1 as an allosteric activating subunit and a conserved Arg as the substrate-orienting residue.

    Evidence X-ray crystallography of CTDNEP1-NEP1R1 with pseudosubstrate, in vitro phosphatase assays, mutagenesis, ER morphology readout

    PMID:38045299 PMID:38776370

    Open questions at the time
    • Physiological substrate repertoire engaged by the activated complex not fully enumerated
    • Regulation of complex assembly in vivo unclear
  13. 2024 High

    Integrated the phosphatase complex with the scaffold MAN1 to provide a molecular mechanism for nuclear R-SMAD dephosphorylation and signal termination.

    Evidence Domain mapping, structural prediction, mutagenesis, Co-IP, phosphoproteomics, TGF-β signaling assays (preprint)

    PMID:bio_10.1101_2024.09.23.614427

    Open questions at the time
    • Preprint; peer review pending
    • Quantitative contribution of MAN1 recruitment vs intrinsic activity not separated
  14. 2024 Medium

    Demonstrated CTDNEP1 negatively regulates osteoclast differentiation, broadening its tissue-level roles in signaling restraint.

    Evidence siRNA knockdown in RAW264.7 cells, TRAP staining, marker qPCR, pRANKL/Nfatc1 western, calcium resorption assay

    PMID:38749090

    Open questions at the time
    • Direct substrate in RANKL pathway not identified
    • Single cell-line model

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how a single phosphatase complex coordinates its distinct substrate sets — R-SMADs, lipin1, MYC, and mitotic regulators — and how substrate selection is controlled across membrane compartments and tissues.
  • No unified model of substrate selection
  • Spatial regulation between nuclear envelope and ER pools unclear
  • Direct in vivo human substrates largely inferred rather than demonstrated

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140096 catalytic activity, acting on a protein 5 GO:0016787 hydrolase activity 2
Localization
GO:0005635 nuclear envelope 2 GO:0005783 endoplasmic reticulum 1
Pathway
R-HSA-162582 Signal Transduction 6 R-HSA-1266738 Developmental Biology 4 R-HSA-1640170 Cell Cycle 1
Complex memberships
CTDNEP1-NEP1R1 phosphatase complexCTDNEP1-NEP1R1-MAN1 complex

Evidence

Reading pass · 15 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2024 Crystal structure of the human CTDNEP1-NEP1R1 complex reveals that NEP1R1 acts as an activating regulatory subunit that binds CTDNEP1 at a site distant from the active site to allosterically stabilize and activate CTDNEP1 phosphatase activity; substrate recognition is facilitated by a conserved Arg residue in CTDNEP1 that binds and orients substrate peptide in the active site; knockdown of NEP1R1 generates identical ER expansion phenotypes to loss of CTDNEP1, establishing them as a conserved complex that restricts ER membrane expansion. X-ray crystallography of CTDNEP1-NEP1R1 complex bound to pseudosubstrate peptide, in vitro phosphatase activity assays, mammalian cell knockdown with ER morphology readout Proceedings of the National Academy of Sciences of the United States of America High 38776370
2023 Crystal structure of the human CTDNEP1-NEP1R1 complex (preprint version confirming peer-reviewed findings): NEP1R1 allosterically activates CTDNEP1 by binding distant from the active site; conserved Arg in CTDNEP1 orients substrate in active site; CTDNEP1 and NEP1R1 co-knockdown generate identical ER expansion phenotypes. X-ray crystallography, in vitro phosphatase assay, cell knockdown with ER morphology readout bioRxivpreprint High 38045299
2024 CTDNEP1-NEP1R1, together with scaffold protein MAN1 at the inner nuclear membrane, forms a complex that dephosphorylates receptor-regulated SMADs (R-SMADs) in the nucleus to terminate TGF-β superfamily signaling; MAN1 binds independently to the CTDNEP1-NEP1R1 phosphatase and to R-SMADs to promote their inactivation; disruption of this complex causes nuclear accumulation of phosphorylated R-SMADs and aberrant signaling even without TGF-β ligand. Domain mapping, structural prediction, mutagenesis, co-immunoprecipitation, phosphoproteomics, cell-based TGF-β signaling assays with R-SMAD dephosphorylation readout bioRxivpreprint High bio_10.1101_2024.09.23.614427
2011 Human CTDNEP1/Dullard is a phosphoserine phosphatase that shows substrate specificity for the insulin-dependent phosphorylation site (Ser106) of lipin1, with a kcat/Km of 2.9×10^4 M−1 s−1, and does not require an adaptor protein for this activity. Steady-state kinetic analysis using phosphoserine-bearing nonapeptides corresponding to lipin1 phosphorylation sites Biochemistry High 21413788
2006 Xenopus Dullard (ortholog of CTDNEP1) promotes ubiquitin-mediated proteasomal degradation of BMP receptors; it preferentially complexes with BMPRII and partially colocalizes with caveolin-1-positive compartments, suggesting degradation via the lipid raft-caveolar pathway; it also associates with BMP type I receptors and represses BMP-dependent phosphorylation of type I receptors; phosphatase activity is required for BMPR degradation and neural induction. Co-immunoprecipitation, colocalization with caveolin-1, phosphatase-dead mutant analysis, Xenopus morpholino knockdown with neural marker gene readout Developmental cell High 17141153
2016 Drosophila Dullard (ortholog of CTDNEP1) physically interacts with phosphorylated Mad (R-Smad) via co-immunoprecipitation and dephosphorylates both C-terminal and linker phosphorylations of Mad to terminate BMP signaling; phosphatase-domain mutation abolishes dephosphorylation but not binding; Dullard acts as an alternative to proteasomal degradation for terminating BMP signaling. Co-immunoprecipitation, hypomorphic allele and RNAi knockdown with pMad western blot readout, Dullard overexpression, phosphatase-domain point mutation Scientific reports High 27578171
2021 CTDNEP1/Dullard, a nuclear envelope phosphatase, directly interacts with the actin regulator Eps8L2; this interaction regulates formation and thickness of dorsal actin cables required for TAN line engagement and nuclear movement during cell migration; loss of either protein impairs nuclear positioning and cell migration. Co-immunoprecipitation/direct interaction assay, siRNA knockdown with nuclear positioning and cell migration readouts, actin cable imaging Current biology : CB Medium 33567288
2023 CTDNEP1 deficiency stabilizes and activates MYC by elevating MYC Ser62 phosphorylation, and triggers chromosomal instability; phosphoproteomics reveals CTDNEP1 post-translationally modulates activities of chromosome segregation and mitotic checkpoint regulators including TOP2A and CHEK1. Ctdnep1 mouse knockout tumor model, phosphoproteomics, MYC phosphorylation (pSer62) western blot, chromosomal instability assays, co-targeting MYC and CHEK1 in tumor models Nature communications High 36765089
2013 Mouse Dullard/Ctdnep1 deletion in metanephric mesenchyme leads to massive nephron apoptosis associated with upregulation of phospho-Smad1/5/8, consistent with cell-autonomous inhibition of BMP signaling; BMP receptor kinase inhibitor LDN-193189 partially rescues the nephron defects, placing Dullard upstream of BMP receptor activity in kidney maintenance. Conditional knockout mouse, phospho-Smad1/5/8 immunostaining, pharmacological rescue with BMP receptor inhibitor LDN-193189 Nature communications High 23360989
2015 Dullard/Ctdnep1 suppresses TGF-β signaling during endochondral ossification; Dullard deficiency upregulates phospho-Smad2/3 and total Smad2/3 protein levels without increasing Smad2/3 mRNA, suggesting Dullard affects Smad2/3 protein stability; skeletal defects in Dullard-deficient mice are rescued by TGF-β type I receptor kinase blocker LY-364947. Conditional knockout mouse (Prx1-Cre), micromass culture, phospho-Smad2/3 and total Smad2/3 western blot, pharmacological rescue with LY-364947 in vivo Journal of bone and mineral research High 25155999
2013 Mouse Dullard/Ctdnep1 loss impairs primordial germ cell formation by reducing WNT/β-catenin signaling and Dishevelled 2 (Dvl2) protein levels; compound heterozygosity with Wnt3 has synergistic effects on PGC reduction, placing Dullard as a positive modulator of WNT signaling in germ cell specification. Conditional mouse knockout, compound heterozygote genetic epistasis with Wnt3, β-catenin reporter assay, Dvl2 western blot PloS one Medium 23469192
2020 Dullard, acting as a nuclear phosphatase, tunes BMP signaling amplitude during cardiac outflow tract development; it maintains neural crest cells in a mesenchymal state by attenuating Sema3c expression and promoting Twist1 expression; loss of Dullard leads to premature NCC condensation and tetralogy of Fallot-like defects, with elevated pSmad1/5/8. Conditional mouse knockout (NCC-specific), BMP signaling reporter, pSmad1/5/8 immunostaining, gene expression analysis of Sema3c and Twist1 eLife High 32105214
2011 Drosophila Dullard (ddd) genetically interacts with BMP signaling pathway components to regulate wing vein formation; manipulation of ddd expression alters p-Mad levels; Dullard overexpression mislocalizes nuclear import machinery components Importin-β and RanGAP and alters membrane lipid staining, suggesting Dullard regulates nuclear envelope/membrane lipid homeostasis to affect BMP signaling. Genetic interaction screen (double mutant wing vein phenotype), pMad western blot, Importin-β and RanGAP localization by immunofluorescence, membrane lipid staining Development, growth & differentiation Medium 21790556
2024 Ctdnep1 negatively regulates RANKL-induced osteoclast differentiation; Ctdnep1 knockdown increases phosphorylation of RANKL signaling components and elevates Nfatc1 protein even without RANKL stimulation, and enhances calcium-resorbing activity. siRNA knockdown in RAW264.7 cells, TRAP staining for osteoclast numbers, osteoclast marker gene qPCR, western blot for pRANKL signaling components and Nfatc1, calcium resorption assay Biochemical and biophysical research communications Medium 38749090
2018 Dullard/Ctdnep1 suppresses BMP signaling in ovarian granulosa cells; conditional deletion leads to hemorrhagic ovarian cysts and upregulation of BMP-inducible inhibitory Smads (Smad6/Smad7); pharmacological rescue with BMP receptor inhibitor LDN-193189 confirmed the mechanism is through BMP signaling. Conditional knockout mouse (Col1a1-Cre and Rosa26-CreER), Smad6/7 gene expression, BMP receptor inhibitor rescue Genes to cells Medium 29521016

Source papers

Stage 0 corpus · 16 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2006 Dullard promotes degradation and dephosphorylation of BMP receptors and is required for neural induction. Developmental cell 68 17141153
2002 Molecular cloning and characterization of dullard: a novel gene required for neural development. Biochemical and biophysical research communications 39 12083771
2023 Loss of phosphatase CTDNEP1 potentiates aggressive medulloblastoma by triggering MYC amplification and genomic instability. Nature communications 25 36765089
2013 The phosphatase Dullard negatively regulates BMP signalling and is essential for nephron maintenance after birth. Nature communications 25 23360989
2011 Homo sapiens dullard protein phosphatase shows a preference for the insulin-dependent phosphorylation site of lipin1. Biochemistry 24 21413788
2013 Dullard/Ctdnep1 modulates WNT signalling activity for the formation of primordial germ cells in the mouse embryo. PloS one 23 23469192
2011 Negative modulation of bone morphogenetic protein signaling by Dullard during wing vein formation in Drosophila. Development, growth & differentiation 19 21790556
2020 Dullard-mediated Smad1/5/8 inhibition controls mouse cardiac neural crest cells condensation and outflow tract septation. eLife 18 32105214
2015 Dullard/Ctdnep1 regulates endochondral ossification via suppression of TGF-β signaling. Journal of bone and mineral research : the official journal of the American Society for Bone and Mineral Research 18 25155999
2021 Ctdnep1 and Eps8L2 regulate dorsal actin cables for nuclear positioning during cell migration. Current biology : CB 17 33567288
2017 Two Triacylglycerol Pathway Genes, CTDNEP1 and LPIN1, are Down-Regulated by hsa-miR-122-5p in Hepatocytes. Archives of Iranian medicine 17 28287811
2016 Drosophila Dullard functions as a Mad phosphatase to terminate BMP signaling. Scientific reports 12 27578171
2018 Dullard deficiency causes hemorrhage in the adult ovarian follicles. Genes to cells : devoted to molecular & cellular mechanisms 10 29521016
2024 Structure and mechanism of the human CTDNEP1-NEP1R1 membrane protein phosphatase complex necessary to maintain ER membrane morphology. Proceedings of the National Academy of Sciences of the United States of America 7 38776370
2024 Ctdnep1 phosphatase is required for negative regulation of RANKL-induced osteoclast differentiation in RAW264.7 cells. Biochemical and biophysical research communications 4 38749090
2023 Structure and mechanism of the human CTDNEP1-NEP1R1 membrane protein phosphatase complex necessary to maintain ER membrane morphology. bioRxiv : the preprint server for biology 0 38045299

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