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Showing TIPRLTIPRL1 is a alias.

TIPRL

TIP41-like protein · UniProt O75663

Length
272 aa
Mass
31.4 kDa
Annotated
2026-06-10
17 papers in source corpus 11 papers cited in narrative 11 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

TIPRL is a conserved regulator of the type 2A serine/threonine phosphatases (PP2A, PP4, and PP6), binding their catalytic subunits and inhibiting their activity (PMID:17944932, PMID:27489114). Its crystal structure reveals a novel fold with a conserved cleft that engages the conserved C-terminal tail of PP2Ac (DYFL motif), preferentially recognizing the unmodified over the tyrosine-phosphorylated tail and blocking access to the phosphatase active site (PMID:27489114). TIPRL binds the catalytic subunits through a region distinct from the alpha4-binding site, allowing formation of a stable alpha4:PP2Ac:TIPRL ternary complex (PMID:17944932). The yeast ortholog TIP41 acts within a TOR-coupled feedback loop with TAP42 to amplify SIT4 phosphatase activity under TOR-inactivating conditions (PMID:11741537), whereas mammalian TIPRL positively regulates mTORC1 signaling by associating with PP2Ac and protecting mTORC1 substrate phosphorylation during nutrient stress (PMID:23892082). Through phosphatase inhibition TIPRL controls multiple stress and survival pathways: it suppresses PP4-mediated dephosphorylation of γ-H2AX to promote the DNA damage response (PMID:26717153), and ATM phosphorylates TIPRL at Ser265 upon irradiation, a modification required for its role in radioresistance alongside newly identified interactions with DNA-PKcs, RAD51, and nucleosomal histones (PMID:37971644). In cancer cells TIPRL additionally promotes survival by binding MKK7 to suppress JNK-mediated TRAIL-induced apoptosis (PMID:24969837, PMID:29348850), by interacting with eIF2α to activate the eIF2α-ATF4 autophagy axis (PMID:31862913), and by binding CaMKK2 to sustain a CaMK4-CREB stemness feedback loop that transcriptionally reinforces TIPRL expression (PMID:39076120).

Mechanistic history

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

    Established the ancestral function of the TIPRL family by showing the yeast ortholog couples TOR signaling to type 2A phosphatase regulation through a TAP42 feedback loop.

    Evidence Genetic epistasis, Co-IP, and phosphorylation/nuclear translocation assays in budding yeast

    PMID:11741537

    Open questions at the time
    • Did not address whether the mammalian ortholog conserves the inhibitory directionality on TOR
    • Direct phosphatase inhibition not biochemically demonstrated
  2. 2005 Medium

    Placed the Tip41 ortholog upstream of PP2A in nutrient-responsive cell cycle control, confirming phosphatase regulation as a conserved function.

    Evidence Genetic epistasis with ppa2 deletion, overexpression, and phosphatase activity assays in fission yeast

    PMID:16297994

    Open questions at the time
    • Direction of effect on phosphatase appeared activating, contrasting later mammalian inhibition data
    • No physical interaction mapping
  3. 2007 High

    Defined TIPRL as a direct inhibitor of multiple type 2A phosphatase catalytic subunits and mapped its binding to a site distinct from alpha4, enabling a ternary complex.

    Evidence Yeast two-hybrid, recombinant pull-downs, reverse two-hybrid mutagenesis, and in vitro phosphatase assays

    PMID:17944932

    Open questions at the time
    • Structural basis of inhibition unresolved
    • Functional consequence of the ternary complex in cells unknown
  4. 2013 Medium

    Showed that mammalian TIPRL positively regulates mTORC1 via PP2Ac, reversing the inhibitory directionality of the yeast ortholog.

    Evidence Overexpression/siRNA with mTORC1 substrate phosphorylation readouts and Co-IP

    PMID:23892082

    Open questions at the time
    • Mechanism of directionality switch from yeast not explained
    • Single-lab phosphorylation readouts
  5. 2014 Medium

    Extended TIPRL function to apoptosis control by identifying its interaction with MKK7 and showing it restrains the JNK death pathway.

    Evidence GST pull-down, ELISA interaction detection, and functional apoptosis assays with an interaction-disrupting extract

    PMID:24969837

    Open questions at the time
    • Whether MKK7 regulation depends on phosphatase activity unknown
    • Interaction interface not mapped
  6. 2015 Medium

    Connected TIPRL to the DNA damage response by showing it inhibits PP4 and thereby sustains γ-H2AX phosphorylation.

    Evidence siRNA/overexpression, phosphatase activity assays, PP4-C/PP4R2 Co-IP, and γ-H2AX foci imaging

    PMID:26717153

    Open questions at the time
    • Whether TIPRL competes with PP4R2 directly not resolved
    • Single-lab evidence
  7. 2016 High

    Provided the structural mechanism of phosphatase inhibition: a conserved cleft binds the PP2Ac C-terminal tail and occludes the active site.

    Evidence X-ray crystallography at 2.15 Å with mutagenesis, HDX-MS, peptide binding, and docking

    PMID:27489114

    Open questions at the time
    • No co-crystal structure with full-length phosphatase
    • Selectivity among PP2A/PP4/PP6 not structurally explained
  8. 2017 Medium

    Validated the MKK7-TIPRL interaction as a druggable node by sensitizing hepatocellular carcinoma to TRAIL-induced apoptosis upon disruption.

    Evidence ELISA-based screening, phosphorylation immunoblots, and xenograft tumor regression

    PMID:29348850

    Open questions at the time
    • Selectivity of small-molecule disruptors not established
    • Relationship to phosphatase function unclear
  9. 2019 Medium

    Identified TIPRL as a promoter of autophagy through interaction with eIF2α and activation of the eIF2α-ATF4 axis.

    Evidence Co-IP, eIF2α phosphorylation/ATF4 immunoblots, LC3-II/p62 autophagy assays, and xenografts in NSCLC

    PMID:31862913

    Open questions at the time
    • How TIPRL promotes eIF2α phosphorylation mechanistically unknown
    • Link to phosphatase inhibition not established
  10. 2023 Medium

    Revealed ATM-dependent phosphorylation of TIPRL at Ser265 as required for radioresistance and identified DDR-related interactors beyond phosphatases.

    Evidence MS phosphosite mapping and interactomics, CRISPR deletion, and Ser265Ala rescue in HNSCC

    PMID:37971644

    Open questions at the time
    • Functional roles of DNA-PKcs/RAD51/histone interactions not dissected
    • Whether Ser265 phosphorylation alters phosphatase inhibition unknown
  11. 2024 Medium

    Linked TIPRL to cancer stemness through a CaMKK2-CaMK4-CREB feedback loop that transcriptionally reinforces TIPRL expression.

    Evidence Co-IP, phosphorylation immunoblots, CREB reporter, knockdown, and xenograft metastasis assays in lung cancer stem cells

    PMID:39076120

    Open questions at the time
    • Direct vs indirect activation of CaMKK2 not resolved
    • Relationship of this loop to phosphatase inhibition unclear

Open questions

Synthesis pass · forward-looking unresolved questions
  • It remains unresolved how a single phosphatase-inhibitory scaffold mechanistically toggles between its PP2A/PP4/PP6 binding mode and its diverse kinase-pathway interactions (MKK7, eIF2α, CaMKK2) and DDR partners.
  • No structural model of TIPRL bound to non-phosphatase partners
  • Unknown whether phosphatase inhibition is required for the JNK, autophagy, and stemness phenotypes
  • Substrate selectivity among the three type 2A phosphatases not defined

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 3 GO:0140096 catalytic activity, acting on a protein 3 GO:0060090 molecular adaptor activity 2
Pathway
R-HSA-162582 Signal Transduction 3 R-HSA-5357801 Programmed Cell Death 2 R-HSA-73894 DNA Repair 2 R-HSA-9612973 Autophagy 1
Complex memberships
alpha4:PP2Ac:TIPRL ternary complex

Evidence

Reading pass · 11 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2001 Yeast TIP41 (ortholog of TIPRL) interacts with TAP42 (ortholog of alpha4) and negatively regulates the TOR signaling pathway; deletion of TIP41 confers rapamycin resistance and prevents dissociation of SIT4 phosphatase from TAP42, blocking SIT4-dependent dephosphorylation of NPR1 and nuclear translocation of GLN3. TIP41 binding to TAP42 is stimulated by rapamycin treatment via SIT4-dependent dephosphorylation of TIP41, establishing a feedback loop that amplifies SIT4 phosphatase activity under TOR-inactivating conditions. Genetic epistasis (deletion/suppressor analysis), co-immunoprecipitation, phosphorylation assays, nuclear translocation assays Molecular cell High 11741537
2007 TIPRL interacts with the C-terminal region (residues 210–309) of the catalytic subunits of PP2A, PP4, and PP6 via a region distinct from the alpha4-binding site; TIPRL and alpha4 can simultaneously bind PP2Ac, forming a stable ternary complex (alpha4:PP2Ac:TIPRL). TIPRL inhibits PP2Ac activity in vitro. Single amino acid substitutions D71L, I136T, M196V, and D198N on TIPRL disrupt its interaction with PP2Ac. The complex is rapamycin-insensitive in human cells. Yeast two-hybrid screen, pull-down assays with recombinant proteins, reverse two-hybrid mutagenesis, in vitro phosphatase activity assay, co-immunoprecipitation from human cells The FEBS journal High 17944932
2013 Mammalian TIPRL positively regulates mTORC1 signaling (in contrast to yeast TIP41 which is inhibitory); overexpression of TIPRL suppressed dephosphorylation of mTORC1 substrates under amino acid withdrawal, while knockdown of TIPRL attenuated phosphorylation of mTORC1 substrates after amino acid refeeding. This action requires TIPRL association with the catalytic subunit of PP2A (PP2Ac). Overexpression and siRNA knockdown with phosphorylation readouts (immunoblot), co-immunoprecipitation FEBS letters Medium 23892082
2015 TIPRL negatively regulates protein phosphatase 4 (PP4) activity; knockdown of TIPRL increases PP4 phosphatase activity and promotes formation of the active PP4-C/PP4R2 complex that dephosphorylates γ-H2AX. Overexpression of TIPRL promotes H2AX phosphorylation and increases γ-H2AX foci in response to DNA damage, while TIPRL knockdown inhibits γ-H2AX phosphorylation and protects cells from genotoxic stress. siRNA knockdown, overexpression, phosphatase activity assay, co-immunoprecipitation (PP4-C/PP4R2 complex), immunofluorescence (γ-H2AX foci), cell viability assays PloS one Medium 26717153
2016 Crystal structure of human TIPRL solved at 2.15 Å resolution reveals a novel fold organized around a central core of antiparallel beta-sheet with an N-terminal α/β region and a conserved cleft. Mutagenesis, pulldown, and hydrogen/deuterium exchange mass spectrometry demonstrate that this conserved cleft binds the conserved C-terminal tail of PP2Ac (mimicked by the peptide DYFL). TIPRL preferentially binds the unmodified PP2A C-terminal tail peptide over its tyrosine-phosphorylated version. Docking modeling suggests TIPRL blocks the phosphatase's active site. X-ray crystallography (2.15 Å), mutagenesis, pulldown assays, hydrogen/deuterium exchange mass spectrometry, docking modeling Scientific reports High 27489114
2014 TIPRL interacts with MKK7 (MAP kinase kinase 7) and this interaction contributes to resistance to TRAIL-induced apoptosis by inhibiting the MKK7-JNK pathway. Disruption of the MKK7-TIPRL interaction (by Tussilago farfara extract) restores MKK7/JNK phosphorylation and sensitizes cells to TRAIL-induced apoptosis. GST pull-down assay, ELISA-based interaction detection, co-treatment functional apoptosis assay (cell viability, caspase activation) Oncology reports Medium 24969837
2017 TIPRL directly interacts with MKK7, and inhibition of this interaction by small molecule compounds leads to increased MKK7 and JNK phosphorylation and sensitizes hepatocellular carcinoma cells to TRAIL-induced apoptosis in vitro and in vivo. ELISA-based high-throughput screening of MKK7-TIPRL interaction, cell viability assay, immunoblot for MKK7/JNK phosphorylation, xenograft in vivo tumor regression assay Oncotarget Medium 29348850
2019 TIPRL interacts with eIF2α and promotes eIF2α phosphorylation, activating the eIF2α-ATF4 pathway to induce autophagy in non-small cell lung cancer cells. TIPRL depletion reduces autophagic clearance and increases apoptosis. Co-immunoprecipitation (TIPRL-eIF2α interaction), immunoblot for eIF2α phosphorylation and ATF4 levels, autophagy assays (LC3-II/p62), siRNA knockdown, xenograft assay Cell death & disease Medium 31862913
2024 TIPRL binds directly to CaMKK2 (calcium/calmodulin-dependent protein kinase kinase 2) in lung cancer stem cells, causing sustained activation of the CaMKK2 signaling pathway, which phosphorylates CaMK4, leading to phosphorylation of CREB at Ser129 and Ser133. Activated CREB then drives expression of Bcl2 and HMG20A and transcriptionally activates TIPRL itself, forming a positive feedback loop. TIPRL depletion sensitizes lung cancer stem cells to afatinib and reduces distal metastasis in vivo. Co-immunoprecipitation (TIPRL-CaMKK2), immunoblot for CaMK4 and CREB phosphorylation, CREB transcriptional reporter, gene expression analysis, siRNA/CRISPR knockdown, xenograft metastasis assay Advanced science Medium 39076120
2023 ATM kinase phosphorylates TIPRL1 at Ser265 upon irradiation. A non-phosphorylatable Ser265Ala mutant cannot rescue the increased radiosensitivity of TIPRL1-depleted HNSCC cells, demonstrating that ATM-dependent phosphorylation of TIPRL1 at Ser265 is required for its role in radiotherapy resistance. TIPRL1 was also found to interact with DNA-PKcs, RAD51, and nucleosomal histones (novel interactors beyond PP2A-family phosphatases); histone binding is stimulated by RT but adversely affected by Ser265 phosphorylation. Mass spectrometry (phosphorylation site mapping and interactomics), CRISPR/Cas9 deletion, rescue with Ser265Ala mutant, immunoblot (DDR signaling), microscopy (micronuclei), flow cytometry (cell cycle) Cellular oncology Medium 37971644
2005 Fission yeast (S. pombe) Tip41 ortholog regulates type 2A phosphatase activity: overexpression of tip41+ increases type 2A phosphatase activity, and in a ppa2 deletion strain with reduced PP2A activity, overexpression of tip41+ no longer blocks the cell cycle G1 shift upon nitrogen starvation, placing Tip41 upstream of PP2A in cellular responses to nitrogen nutrient conditions. Genetic epistasis (tip41 deletion and ppa2 deletion double mutant), overexpression, phosphatase activity assay, cell cycle analysis Biochimica et biophysica acta Medium 16297994

Source papers

Stage 0 corpus · 17 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2001 TIP41 interacts with TAP42 and negatively regulates the TOR signaling pathway. Molecular cell 194 11741537
2000 Expression of the prolactin receptor (tiPRL-R) gene in tilapia Oreochromis niloticus: tissue distribution and cellular localization in osmoregulatory organs. Journal of molecular endocrinology 48 10750022
2013 A positive role of mammalian Tip41-like protein, TIPRL, in the amino-acid dependent mTORC1-signaling pathway through interaction with PP2A. FEBS letters 36 23892082
2021 Hsa_circ_0010235 functions as an oncogenic drive in non-small cell lung cancer by modulating miR-433-3p/TIPRL axis. Cancer cell international 31 33494763
2007 Interaction analysis of the heterotrimer formed by the phosphatase 2A catalytic subunit, alpha4 and the mammalian ortholog of yeast Tip41 (TIPRL). The FEBS journal 31 17944932
2019 TIPRL potentiates survival of lung cancer by inducing autophagy through the eIF2α-ATF4 pathway. Cell death & disease 29 31862913
2014 Tussilago farfara L. augments TRAIL-induced apoptosis through MKK7/JNK activation by inhibition of MKK7‑TIPRL in human hepatocellular carcinoma cells. Oncology reports 22 24969837
2015 TIPRL Inhibits Protein Phosphatase 4 Activity and Promotes H2AX Phosphorylation in the DNA Damage Response. PloS one 21 26717153
2020 TIPRL, a Novel Tumor Suppressor, Suppresses Cell Migration, and Invasion Through Regulating AMPK/mTOR Signaling Pathway in Gastric Cancer. Frontiers in oncology 17 32719745
2019 The positive correlation of TIPRL with LC3 and CD133 contributes to cancer aggressiveness: potential biomarkers for early liver cancer. Scientific reports 16 31727942
2016 Crystal structure of the human Tip41 orthologue, TIPRL, reveals a novel fold and a binding site for the PP2Ac C-terminus. Scientific reports 15 27489114
2022 TDP-43 Toxicity in Yeast Is Associated with a Reduction in Autophagy, and Deletions of TIP41 and PBP1 Counteract These Effects. Viruses 13 36298819
2024 TIPRL Regulates Stemness and Survival in Lung Cancer Stem Cells through CaMKK2-CaMK4-CREB Feedback Loop Activation. Advanced science (Weinheim, Baden-Wurttemberg, Germany) 12 39076120
2017 Novel indazole-based small compounds enhance TRAIL-induced apoptosis by inhibiting the MKK7-TIPRL interaction in hepatocellular carcinoma. Oncotarget 12 29348850
2023 TIPRL1 and its ATM-dependent phosphorylation promote radiotherapy resistance in head and neck cancer. Cellular oncology (Dordrecht, Netherlands) 8 37971644
2005 Fission yeast homologue of Tip41-like proteins regulates type 2A phosphatases and responses to nitrogen sources. Biochimica et biophysica acta 8 16297994
2024 TIPRL, a Potential Double-edge Molecule to be Targeted and Re-targeted Toward Cancer. Cell biochemistry and biophysics 1 38888871

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