Affinage

SSU72

RNA polymerase II subunit A C-terminal domain phosphatase SSU72 · UniProt Q9NP77

Length
194 aa
Mass
22.6 kDa
Annotated
2026-06-10
44 papers in source corpus 34 papers cited in narrative 34 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

SSU72 is a cis-proline-dependent phosphatase structurally related to low-molecular-weight protein tyrosine phosphatases that coordinates the RNA polymerase II transcription cycle and pre-mRNA 3'-end processing, and which has been repurposed across diverse signaling contexts as a regulatory phosphatase (PMID:12606538, PMID:15125841, PMID:12704082). Its catalytic activity depends on a CX5R-motif cysteine, and it acts as the founding member of a distinct phosphatase subfamily that cleaves phosphotyrosine analogues in vitro (PMID:12606538). As a CTD phosphatase it preferentially dephosphorylates Ser5-P, and with ~4000-fold lower efficiency Ser7-P, recognizing the phosphoserine-proline bond only in its cis configuration—a conformation supplied by Pin1/Ess1 prolyl isomerase, making SSU72 the founding example of a cis-proline-specific enzyme (PMID:15125841, PMID:23070812, PMID:21159777, PMID:33410907). Crystal structures of SSU72 in ternary complex with the symplekin/Pta1 N-terminal domain and CTD phosphopeptides established the catalytic mechanism (a phosphoryl-enzyme intermediate captured by vanadate), the active-site residues that read the CTD, and the opposite-orientation binding of pSer5 versus pSer7 substrates (PMID:20861839, PMID:21204787, PMID:23070812). Through these activities SSU72 functions both at the initiation-to-elongation transition, where it removes Ser5-P and facilitates the transition, and at 3'-end processing as an integral subunit of the CPF complex—bridging Pta1/symplekin, TFIIB, and RNAP II via Rpb2—required for pre-mRNA cleavage and for both poly(A)-dependent and Nrd1-dependent (snoRNA/snRNA) termination, with its 3'-processing role being separable from its catalytic activity (PMID:17101794, PMID:12704082, PMID:12453421, PMID:19188448, PMID:15125841, PMID:25166011). Beyond transcription, SSU72 dephosphorylates Stn1 (Ser74) to control STN1 recruitment and terminate telomere replication (PMID:30796050); antagonizes Aurora B (which phosphorylates SSU72 at Ser19 to inactivate and degrade it) to maintain sister chromatid cohesion (PMID:24149858); and acts as a regulatory phosphatase in immune and metabolic signaling, targeting ZAP-70, the GM-CSF receptor β-chain, STAT3, PLCγ1-dependent Treg differentiation, and eIF2α in brown adipose tissue thermogenesis (PMID:34452999, PMID:32910932, PMID:28710354, PMID:33850312, PMID:36841836). In vivo, conditional loss of SSU72 in mouse liver produces hepatocyte polyploidization, dedifferentiation via HNF4α hypophosphorylation, and predisposition to NAFLD/NASH and HCC (PMID:26458163, PMID:34616001).

Mechanistic history

Synthesis pass · year-by-year structured walk · 19 steps
  1. 1996 Medium

    Before any biochemical role was known, genetics placed SSU72 at the transcription start-site selection step, linking it to the basal initiation machinery.

    Evidence Synthetic enhancement of a TFIIB (sua7-1) defect with start-site mapping and N-terminal/cysteine mutagenesis in yeast

    PMID:8657130

    Open questions at the time
    • No molecular activity defined
    • Direct physical contacts with the PIC not established
  2. 1999 Medium

    Allele-specific genetics formalized a functional relationship among Ssu72, TFIIB, and Sub1 in controlling start-site accuracy, sharpening SSU72's role at initiation.

    Evidence Allele-specific genetic interaction analysis and error-prone PCR mutagenesis in yeast

    PMID:10511545

    Open questions at the time
    • Mechanism of start-site control unresolved
    • No enzymatic activity yet assigned
  3. 2000 Medium

    A direct physical link to the polymerase was established, anchoring Ssu72 to the RNAP II core via Rpb2.

    Evidence Co-immunoprecipitation with purified RNAP II plus an rpb2 (R512C) suppressor of ssu72-2 in yeast

    PMID:11046131

    Open questions at the time
    • Single Co-IP without reciprocal mapping
    • Functional consequence of the interaction undefined
  4. 2002 Medium

    Ssu72 was shown to be a stable CPF subunit bridging Pta1, Ydh1/Cft2, TFIIB and RNAP II, connecting initiation factors to the 3'-end machinery and to elongation/termination control.

    Evidence Biochemical fractionation, interaction assays, and 6-azauracil genetic suppression of ssu72-2 in yeast

    PMID:12453421

    Open questions at the time
    • Catalytic substrate still unknown
    • How a single factor couples initiation and 3' processing unresolved
  5. 2003 High

    SSU72 was defined enzymatically as a phosphatase of the low-MW PTP class, and biochemically as a CPF subunit essential for pre-mRNA cleavage, with the surprising finding that its 3'-processing role is independent of catalysis.

    Evidence In vitro phosphatase assays with pNPP and catalytic-cysteine mutagenesis; biochemical depletion/reconstitution with recombinant protein in in vitro cleavage assays; genome-wide and microarray analyses of termination defects in yeast

    PMID:12606538 PMID:12660165 PMID:12704082 PMID:12944462

    Open questions at the time
    • Physiological phospho-substrate not yet identified
    • Catalysis-independent processing role mechanistically unexplained
  6. 2004 High

    The physiological substrate was identified as the RNAP II CTD with specificity for Ser5-P, defining SSU72 as a CTD phosphatase acting within CPF.

    Evidence In vitro CTD phosphatase assay showing Ser5-P (not Ser2-P) specificity, with recombinant-protein reconstitution of in vitro transcription in yeast

    PMID:15125841

    Open questions at the time
    • Structural basis of substrate selectivity unknown
    • Regulation of activity during the transcription cycle unclear
  7. 2006 High

    Catalytically impaired SSU72 was shown to accumulate Ser5-P and impair elongation, with genetic suppressors defining SSU72 as facilitating the initiation-to-elongation transition.

    Evidence In vitro CTD phosphatase and elongation assays, in vivo CTD phosphorylation, and suppressor screens (RPB1, RPB2, SPT4) in yeast

    PMID:17101794

    Open questions at the time
    • How dephosphorylation triggers the transition not resolved at structural level
  8. 2009 High

    Mapping showed the Pta1 N-terminus is required for and regulates Ssu72 function, neutralizing an inhibitory effect on 3'-end processing and linking Ssu72 to gene looping.

    Evidence Deletion mutagenesis, degron depletion, and in vitro cleavage/polyadenylation assays in yeast

    PMID:19188448

    Open questions at the time
    • Structural basis of Pta1-Ssu72 regulation not yet visualized
    • Mechanism of gene looping contribution unresolved
  9. 2010 High

    Crystal structures of the symplekin–Ssu72–CTD complex revealed the cis pSer5-Pro6 configuration in the active site and showed symplekin both stimulates Ssu72 catalysis and couples it to 3'-processing control, providing the structural logic for substrate recognition.

    Evidence X-ray crystallography (2.4 Å human symplekin–Ssu72–pSer5; Ssu72–pSer5 with NMR isomer analysis), mutagenesis, and in vitro phosphatase/polyadenylation assays

    PMID:20861839 PMID:21159777

    Open questions at the time
    • In vivo contribution of cis-proline preference not yet tested
    • Isomerase that supplies the cis substrate not yet identified in cells
  10. 2011 High

    A transition-state-analogue structure defined the phosphoryl-enzyme intermediate mechanism and the active-site groove residues required for CTD recognition.

    Evidence X-ray crystallography of apo and vanadate-bound Drosophila Ssu72 with mutagenesis and differential scanning fluorimetry

    PMID:21204787

    Open questions at the time
    • In vivo roles of the identified recognition residues not all validated
  11. 2012 High

    SSU72 was shown to act as a Ser7-P phosphatase as well, binding pSer7 in the opposite orientation to pSer5 but with ~4000-fold lower efficiency, and coupling CTD mark removal to termination.

    Evidence X-ray crystallography of the human symplekin–Ssu72–pSer7 ternary complex, kinetic comparison of pSer5/pSer7, and lethal phosphomimetic and genome-wide termination analyses

    PMID:22235117 PMID:23070812

    Open questions at the time
    • Physiological significance of the slow Ser7-P activity unclear
    • Coordination of multiple CTD marks at termination not fully resolved
  12. 2013 High

    SSU72 activity was shown to be tuned by neighboring CTD marks (Thr4-P reduces but does not abolish Ser5-P turnover) and counter-regulated in mitosis by Aurora B, which phosphorylates Ser19 to inactivate and degrade SSU72 and links it to sister chromatid cohesion.

    Evidence Crystallography of doubly phosphorylated CTD with kinetics; in vitro kinase assay, degradation and cohesin Co-IP, and phosphomimetic cohesion assays

    PMID:23844594 PMID:24149858

    Open questions at the time
    • The cohesin component dephosphorylated by SSU72 not identified
    • How a CTD phosphatase localizes to chromosome arms mechanistically unclear
  13. 2014 High

    Vertebrate and human studies confirmed conserved Ser5/Ser7 CTD phosphatase activity, defined entry of SSU72 at the PIC with an RNA-length-gated activity window, and revealed it as a host factor co-opted by HIV-1 Tat to stimulate proviral transcription.

    Evidence DT40 conditional knockout with 3'-end and ChIP analyses; in vitro complex fractionation with RNA-length cutoff; Tat interaction, ChIP-seq/GRO-seq and in vitro phosphatase stimulation

    PMID:25166011 PMID:25319827 PMID:25339178 PMID:30901332

    Open questions at the time
    • Trans-acting factors imposing the 28-nt activity cutoff unidentified
    • How Tat redirects SSU72 to promote rather than restrain transcription unresolved
  14. 2015 Medium

    In vivo loss-of-function in mouse liver showed SSU72 is required to restrain cell-cycle progression, preventing aberrant polyploidization and protecting against liver injury.

    Evidence Liver-specific conditional knockout with cell-cycle, flow cytometry, and histopathology analyses

    PMID:26458163

    Open questions at the time
    • Direct cell-cycle substrate of SSU72 in hepatocytes not defined
    • Link between transcriptional/processing role and ploidy control unclear
  15. 2019 High

    SSU72 was extended beyond transcription to telomere biology, dephosphorylating Stn1 Ser74 to control STN1 telomere recruitment and terminate telomere replication, a function conserved to human cells.

    Evidence Genetic deletion, telomere ChIP for Stn1, in vitro phosphatase assay, and overhang/telomerase analyses in fission yeast and human cells

    PMID:30796050

    Open questions at the time
    • How SSU72 is targeted to telomeres unresolved
    • Relationship to its transcriptional pool unclear
  16. 2020 High

    Pin1 prolyl isomerase was shown to directly recruit SSU72 and generate its cis-Pro substrate, integrating SSU72 into stress-responsive transcription and broad 3'-processing/termination programs.

    Evidence Protein interaction, ChIP, genetic epistasis (pin1Δ × ssu72) and transcriptome profiling in fission yeast and human cells; GM-CSFR β-chain Co-IP and dual conditional knockouts establishing an alveolar-macrophage role

    PMID:32282918 PMID:32910932 PMID:33410907

    Open questions at the time
    • Whether Pin1-SSU72 coupling operates at all SSU72 functions untested
    • Catalytic versus binding contribution to GM-CSFR regulation not fully separated
  17. 2021 High

    SSU72 was established as a direct immune-signaling phosphatase (dephosphorylating ZAP-70 tyrosine, complexing with PLCγ1 for Treg development) and as a hepatic differentiation regulator acting through HNF4α phosphorylation.

    Evidence AP-MS and in vitro phosphatase assay on ZAP-70 with conditional T-cell KO; PLCγ1 Co-IP and Treg differentiation assays; liver-specific KO with HNF4α phosphorylation, dedifferentiation and HCC models

    PMID:33850312 PMID:34452999 PMID:34616001

    Open questions at the time
    • How a CTD phosphatase achieves substrate specificity for tyrosine targets in cytoplasm unclear
    • Direct versus indirect basis of HNF4α dephosphorylation not fully resolved
  18. 2022 Medium

    SSU72 was shown to be targeted by viral antagonism: influenza NS1 binds and degrades SSU72, causing trans-strand transcriptional readthrough that disrupts STAT1/2 expression.

    Evidence NS1-SSU72 Co-IP, SSU72 overexpression rescue of readthrough and lung injury, patient PBMC analysis

    PMID:35332300

    Open questions at the time
    • Mechanism of trans-strand readthrough not biochemically dissected
    • Single-lab Co-IP for the NS1 interaction
  19. 2023 High

    SSU72 was placed in metabolic homeostasis as an eIF2α phosphatase in brown adipocytes that enables translation of thermogenic/OXPHOS effectors.

    Evidence In vitro eIF2α dephosphorylation, adipocyte-specific KO with polysome profiling, mitochondrial and cold-tolerance assays, and re-expression rescue

    PMID:36841836

    Open questions at the time
    • How SSU72 is partitioned to cytoplasmic translational control versus nuclear transcription unclear
    • Regulation of SSU72 by cold/thermogenic signaling not mechanistically defined

Open questions

Synthesis pass · forward-looking unresolved questions
  • A unifying account of how one cis-proline-dependent phosphatase is partitioned and substrate-targeted across the nuclear CTD/3'-processing machinery and the many cytoplasmic substrates (ZAP-70, GM-CSFR, STAT3, eIF2α, Stn1) remains unresolved.
  • No structural or regulatory model explaining tyrosine-substrate targeting
  • Mechanisms controlling nuclear versus cytoplasmic SSU72 pools undefined
  • Targeting determinants for non-CTD substrates unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0140096 catalytic activity, acting on a protein 10 GO:0016787 hydrolase activity 3 GO:0140098 catalytic activity, acting on RNA 2
Localization
GO:0005634 nucleus 3 GO:0005829 cytosol 2 GO:0005886 plasma membrane 1
Pathway
R-HSA-168256 Immune System 4 R-HSA-74160 Gene expression (Transcription) 3 R-HSA-8953854 Metabolism of RNA 3 R-HSA-1640170 Cell Cycle 2
Complex memberships
CPF (cleavage and polyadenylation factor)symplekin-Ssu72 complex

Evidence

Reading pass · 34 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
1996 SSU72 was identified as a synthetic enhancer of a TFIIB (sua7-1) defect, causing a downstream shift in transcription start site selection at ADH1; the N terminus of Ssu72 was shown to be essential for function, and cysteine residues in this region are critical, suggesting involvement in assembly of the transcription preinitiation complex. Genetic epistasis (synthetic enhancement screen), mutational analysis, in vivo transcription start site mapping Molecular and cellular biology Medium 8657130
1999 Functional interactions among TFIIB, Ssu72, and Sub1 in transcription start site selection were established; allele-specific interactions between ssu72-1 and sua7 alleles that affect start site accuracy were demonstrated, defining a functional relationship between Ssu72 and TFIIB. Allele-specific genetic interaction analysis, error-prone PCR mutagenesis screen, in vivo transcription assays Genetics Medium 10511545
2000 Ssu72 physically interacts directly with purified RNA polymerase II via its Rpb2 subunit, as shown by co-immunoprecipitation; an rpb2 suppressor mutation (R512C) was identified that genetically interacts with ssu72-2, linking Ssu72 to the RNAP II core machinery during transcription initiation. Co-immunoprecipitation with purified RNAP II, suppressor genetic screen, sequence analysis Molecular and cellular biology Medium 11046131
2002 Ssu72 is stably associated with yeast CPF and bridges CPF subunits Pta1 and Ydh1/Cft2p, TFIIB, and RNAP II via Rpb2; ssu72-2 mutants show defects in RNAP II transcription elongation and termination, and 6-azauracil (which slows elongation) suppresses the ssu72-2 growth defect, indicating Ssu72 exerts a negative influence on RNAP II elongation. Biochemical fractionation, protein interaction assays, genetic suppression with 6-AU, transcription analyses in ssu72-2 mutants Molecular cell Medium 12453421
2003 Ssu72 is a component of the yeast CPF complex required for 3' end cleavage of pre-mRNA but dispensable for poly(A) addition and RNAP II termination; the in vitro cleavage defect caused by Ssu72 depletion is rescued by recombinant Ssu72; Ssu72 physically interacts with Pta1 subunit of CPF; Sub1 interactions with Pta1 are mutually exclusive with Ssu72 interactions. Biochemical depletion and reconstitution with recombinant protein, in vitro cleavage assay, co-immunoprecipitation, genetic interaction analysis Genes & development High 12704082
2003 Ssu72 is a phosphatase resembling low-molecular-weight protein tyrosine phosphatases; recombinant Ssu72 cleaves the phosphotyrosine analogue p-nitrophenylphosphate; this activity is inhibited by PTPase-inhibiting agents; the CX5R signature motif catalytic cysteine is essential for activity in vitro and for viability in vivo; Ssu72 is proposed as the founding member of a new phosphatase subfamily. In vitro phosphatase assay with p-nitrophenylphosphate, site-directed mutagenesis of catalytic cysteine, inhibitor studies, secondary structure prediction The Journal of biological chemistry High 12606538
2003 Ssu72 mutations disrupt both Nrd1-dependent (poly(A)-independent, snoRNA) termination and poly(A)-dependent termination of RNAP II, demonstrating that Ssu72 mediates both termination pathways; Ssu72 was identified in a genome-wide selection for factors influencing the SNR13 snoRNA terminator. Genome-wide genetic selection, analysis of snoRNA read-through termination, in vivo transcription assays Molecular and cellular biology Medium 12944462
2003 Ssu72 is a phosphatase that physically interacts with CTD kinase Kin28 and functionally interacts with CTD phosphatase Fcp1; ssu72 mutants exhibit read-through transcription of snoRNAs and specific mRNAs, implicating Ssu72 in transcription termination of specific transcripts, possibly by promoting RNA polymerase pausing. Genome-wide expression analysis (DNA microarray), physical interaction assays, analysis of ssu72-ts69 mutant phenotype The EMBO journal Medium 12660165
2004 Ssu72, a component of yeast CPF, is a CTD phosphatase with specificity for Ser5-P (not Ser2-P); Ssu72 catalyzes CTD Ser5-P dephosphorylation in association with Pta1 component of CPF; depletion of Ssu72 impairs transcription in vitro and this defect is rescued by recombinant catalytically active Ssu72; the essential role of Ssu72 in 3' end processing is independent of its catalytic activity. In vitro CTD phosphatase assay, in vitro transcription assay, complementation with recombinant protein, biochemical fractionation with CPF Molecular cell High 15125841
2005 Human Ssu72 (hSsu72/HSPC182) was identified via yeast two-hybrid as a pRb-binding factor; interaction between hSsu72 and pRb was confirmed in transfected mammalian cells and involved multiple pRb domains; mammalian Ssu72 associates with TFIIB and yeast Pta1 and exhibits intrinsic phosphatase activity; endogenous and ectopically expressed mammalian Ssu72 resides primarily in the cytoplasm with only partial nuclear localization. Yeast two-hybrid, co-immunoprecipitation in transfected mammalian cells, in vitro phosphatase assay, subcellular fractionation and localization, siRNA knockdown Nucleic acids research Medium 15659578
2006 Ssu72-R129A (ssu72-2) is catalytically impaired in vitro and causes accumulation of Ser5-P form of RNAP II in vivo; ssu72-2 exhibits impaired elongation efficiency in vitro; suppressors in RPB1 (R1281A) and RPB2 (R983G), as well as slow RNAP II alleles rpb2-4 and rpb2-10 and deletion of SPT4 (Spt4-Spt5 complex subunit), suppress ssu72-2, defining Ssu72 as a transcription elongation factor that facilitates the transition from initiation to elongation. In vitro CTD phosphatase assay, in vivo CTD phosphorylation analysis, in vitro transcription elongation assay, suppressor genetic screen, multiple allele analysis Molecular and cellular biology High 17101794
2009 The N-terminal region (first 75 aa) of Pta1 is required for Ssu72 function: pta1-Δ75 mutant is defective for snoRNA termination, CTD Ser5-P dephosphorylation, and gene looping; the first 300 aa of Pta1 are sufficient for interaction with Ssu72; degron-mediated depletion of Pta1 leads to loss of Ssu72 protein; the Pta1 N-terminus has an inhibitory effect on 3'-end processing that is neutralized through interaction with Ssu72. Deletion mutagenesis, degron-mediated depletion, in vitro cleavage/polyadenylation assays, CTD phosphorylation analysis, protein interaction mapping Molecular and cellular biology High 19188448
2010 Crystal structure of the human symplekin N-terminal domain (ARM/HEAT fold) in ternary complex with Ssu72 and a CTD Ser5-phosphopeptide resolved at 2.4 Å; the pSer5-Pro6 peptide bond is in the cis configuration in the Ssu72 active site; Ssu72 has similarity to low-MW phosphotyrosine protein phosphatase with unique active-site features; engineered mutations in the symplekin-Ssu72 interface abolish their interaction; symplekin N-terminal domain stimulates Ssu72 CTD phosphatase activity in vitro; symplekin N-terminal domain inhibits polyadenylation in vitro only when coupled to transcription, and catalytically active Ssu72 overcomes this inhibition. X-ray crystallography at 2.4 Å, site-directed mutagenesis, in vitro phosphatase assay, in vitro polyadenylation coupled to transcription Nature High 20861839
2010 Crystal structure of Ssu72 in complex with a CTD Ser5-phosphopeptide reveals that the cis-Ser(P)5-Pro6 isomer is the preferred substrate; the cis-Ser(P)5-Pro6 isomer is the minor population in solution; Ess1 (Pin1 ortholog)-catalyzed cis-trans proline isomerization facilitates rapid Ser5-P dephosphorylation by Ssu72, providing the first structural evidence of a cis-proline-specific enzyme. X-ray crystallography, NMR analysis of cis/trans isomer populations, in vitro phosphatase kinetics with Ess1, mutagenesis The Journal of biological chemistry High 21159777
2011 Crystal structures of Drosophila Ssu72 in apo form and in complex with vanadate (transition state analogue) at 2.35 Å reveal a phosphoryl-enzyme intermediate mechanism; Ssu72 has a core fold similar to low-MW PTPs with a unique 'cap' domain sheltering the active site; mutagenesis identified five residues (Met17, Pro46, Asp51, Tyr77, Met85) in the active-site groove essential for CTD substrate recognition. X-ray crystallography (apo and vanadate-bound), site-directed mutagenesis, differential scanning fluorimetry The Biochemical journal High 21204787
2012 Ssu72 is identified as a Ser7-P phosphatase of the RNAP II CTD; phospho-Ser7 substitution with glutamate (phosphomimetic) is lethal; Ssu72 removal of Ser7-P is mechanistically coupled to removal of other CTD marks during transcription termination; inability to remove phospho-CTD marks prevents efficient Pol II termination. In vivo CTD phosphorylation analysis (mass spectrometry), lethal phosphomimetic mutant analysis, genome-wide termination assays, ChIP The Journal of biological chemistry High 22235117
2012 Crystal structure of a ternary complex of human symplekin N-terminal domain, human Ssu72, and a pSer7 CTD peptide shows the peptide is bound in the Ssu72 active site with its backbone running in the opposite direction compared with the pSer5 peptide; Ssu72 pSer7 phosphatase activity is ~4000-fold lower than pSer5 phosphatase activity toward peptide substrate. X-ray crystallography, in vitro phosphatase kinetic assay comparing pSer5 and pSer7 substrates Genes & development High 23070812
2013 Aurora B kinase directly interacts with and phosphorylates Ssu72 at Ser19 in vitro and in vivo; Aurora B-mediated phosphorylation of Ssu72 causes structural modification of Ssu72, downregulates its phosphatase activity, and triggers ubiquitin-dependent degradation of Ssu72; Ssu72 is identified as a new cohesin-binding phosphatase required for maintenance of chromosome arm cohesion; overexpression of the Aurora B phosphomimetic Ssu72 mutant prevents chromosome arm cohesion. In vitro kinase assay, in vivo phosphorylation analysis, ubiquitin-dependent degradation assay, cohesin co-immunoprecipitation, overexpression of phosphomimetic mutants, chromosome cohesion assays Nature communications High 24149858
2013 Thr4 phosphorylation of the CTD reduces Ssu72 Ser5-P phosphatase activity 4-fold but does not abolish it; Ssu72 does not dephosphorylate pThr4; the CTD adopts an almost identical cis conformation for Ssu72 recognition whether Ser5 alone or both Thr4/Ser5 are phosphorylated, despite loss of an intramolecular hydrogen bond; kinetic and structural data established that Thr4 phosphorylation fine-tunes but does not abolish Ssu72 activity. X-ray crystallography (Drosophila Ssu72-symplekin complex with doubly phosphorylated CTD peptide), mass spectrometry, in vitro phosphatase kinetics ACS chemical biology High 23844594
2014 Ssu72 dephosphorylates Ser5-P at the initiation-elongation transition in vivo; Ssu72 indirectly affects Ser2-P levels during elongation but does so independent of its catalytic activity; Ssu72 interacts with components of the initiation machinery yet is an integral component of CPF; Ssu72 is unique in having specificity for Ser5-P in one CTD orientation and Ser7-P in the opposite orientation. In vivo CTD phosphorylation analysis (ChIP with phospho-specific antibodies), catalytic mutant analysis, epistasis with initiation/elongation factors The Journal of biological chemistry Medium 25339178
2014 HIV-1 Tat interacts directly with Ssu72 and strongly stimulates its CTD phosphatase activity; Ssu72 is essential for Tat:P-TEFb-mediated phosphorylation of the Ser5-P CTD in vitro; Ssu72 is recruited by Tat to the integrated HIV-1 proviral promoter in T cells (ChIP); Ssu72 stimulates nascent HIV-1 transcription in a phosphatase-dependent manner; Ssu72 predominantly co-localizes with Ser5-P RNAP II at promoters genome-wide. Direct protein interaction assay, in vitro CTD phosphatase assay with Tat stimulation, ChIP, ChIP-seq, GRO-seq, in vivo transcription analysis Genes & development High 25319827
2014 Vertebrate (chicken) Ssu72 exhibits Ser5- and Ser7-specific CTD phosphatase activity in vitro; conditional inactivation of Ssu72 in DT40 cells causes defects in 3'-end formation of U2 and U4 snRNAs and GAPDH mRNA; Ssu72 inactivation increases efficiency of 3'-end formation of non-polyadenylated replication-dependent histone mRNA; Ssu72 depletion causes significant increases in both Ser5 and Ser7 phosphorylation of Pol II CTD at affected genes. Conditional knockout in DT40 cells, in vitro phosphatase assay, RT-PCR and Northern blot for 3'-end formation, ChIP with CTD phospho-specific antibodies PloS one High 25166011
2015 Liver-specific conditional knockout of Ssu72 in mice results in aberrant hepatocyte chromosome polyploidization, deregulation of cell cycle progression by overriding the G1 restriction point, and promotion of DNA endoreplication through G2/M arrest; Ssu72 depletion is associated with impaired liver damage response and markers of liver injury (fibrosis, steatosis, steatohepatitis). Conditional knockout mouse model (liver-specific), cell cycle analysis, flow cytometry, histopathology Hepatology (Baltimore, Md.) Medium 26458163
2017 Ssu72 overexpression suppresses STAT3 activation and Th17 cell responses in vitro; systemic infusion of Ssu72 attenuates experimental autoimmune arthritis by reducing STAT3 activity and Th17 cell differentiation, associated with reduced p-STAT3 levels. In vitro overexpression assays, systemic protein infusion in arthritis mouse model, p-STAT3 measurement, Th17 differentiation assays Scientific reports Medium 28710354
2019 Ssu72 phosphatase is responsible for terminating the cycle of telomere replication in fission yeast; Ssu72 controls recruitment of Stn1 to telomeres by regulating Stn1 phosphorylation at Ser74 within its conserved OB-fold domain; ssu72Δ mutants exhibit defective lagging-strand DNA synthesis with long 3'-ssDNA overhangs; human SSU72 regulates telomerase activation by controlling recruitment of hSTN1 to telomeres. Genetic deletion, telomere ChIP for Stn1 recruitment, in vitro phosphatase assay, telomere length and overhang analysis, human cell telomerase assays The EMBO journal High 30796050
2019 Human Ssu72 is physically associated with early RNAP II elongation complexes and enters the transcription cycle during PIC formation; Ssu72 phosphatase activity on early elongation complexes is strictly limited to complexes containing RNA shorter than 28 nt; when PICs are washed before initiation, this 28 nt cutoff is lost, suggesting trans-acting factors regulate Ssu72 activity during the initiation-to-pausing transition. Biochemical fractionation of RNAP II transcription complexes, in vitro transcription elongation assay, phosphatase activity assay on isolated complexes, salt wash experiments PloS one Medium 30901332
2020 In fission yeast, Pin1 prolyl isomerase directly recruits Ssu72 phosphatase to facilitate dephosphorylation of the RNAP II CTD for transcription elongation; in response to oxidative stress, Pin1 promotes dissociation of Sty1 MAPK from the CTD and recruits Ssu72 to dephosphorylate Ser5-P CTD, enabling productive transcription elongation. Protein interaction assays, genetic analysis (pin1Δ), in vivo ChIP, in vitro assays, analysis in fission yeast and human cancer cells Nucleic acids research Medium 33410907
2020 Transcriptome profiling in fission yeast pin1Δ overlaps 77–100% with the effects of Ssu72 inactivation, supporting Pin1 as a positive effector of 3' processing/termination that acts via its ability to generate cis-Pro6 substrate for Ssu72; CTD-S7A mutation is de-repressive for PHO genes and this effect is erased by pin1Δ in a Ssu72-dependent manner. Transcriptional profiling (RNA-seq), genetic epistasis analysis (pin1Δ × ssu72 inactivation), CTD mutant analysis Nucleic acids research Medium 32282918
2020 Following GM-CSF stimulation, Ssu72 directly binds to the GM-CSF receptor β-chain in alveolar macrophages, preventing its phosphorylation; Ssu72-deficient AMs show higher phosphorylation of GM-CSFR β-chain and downstream molecules; JAK2 inhibitor restores normal signaling in Ssu72-deficient AMs; LysM-Cre and CD11c-Cre conditional knockouts establish Ssu72 as required for AM development, maturation, and mitochondrial function. Co-immunoprecipitation (Ssu72-GM-CSFR β-chain binding), conditional knockout mice (LysM-Cre and CD11c-Cre), phosphorylation analysis, JAK2 inhibitor rescue, adoptive transfer experiments The Journal of allergy and clinical immunology High 32910932
2021 Ssu72 is activated by TCR and IL-2R signaling pathways and localizes at the cell membrane; Ssu72 forms a complex with PLCγ1; Ssu72 deficiency in T cells impairs PLCγ1 downstream signaling and results in failure of Foxp3 induction and Treg development; T-cell-specific Ssu72 deletion disrupts CD4+ T-cell differentiation into Tregs via overproduction of IL-2 and IFNγ. Conditional T-cell KO (Foxp3-Cre, CD4-Cre), co-immunoprecipitation (Ssu72-PLCγ1 complex), cytokine analysis, Foxp3 induction assays, mucosal tolerance patient samples Cellular & molecular immunology Medium 33850312
2021 Ssu72 phosphatase directly binds ZAP-70 in T cells (shown by affinity purification-mass spectrometry and in vitro assay); recombinant Ssu72 reduces tyrosine phosphorylation of ZAP-70 via phosphatase activity in vitro; Ssu72-deficient T cells show increased ZAP-70 phosphorylation and hyperresponsiveness; CD4-Cre Ssu72 fl/fl mice develop spontaneous inflammation with altered T cell subset distribution. Affinity purification-mass spectrometry, in vitro phosphatase assay (recombinant Ssu72 on ZAP-70), conditional T-cell KO, phosphorylation analysis Proceedings of the National Academy of Sciences of the United States of America High 34452999
2021 Liver-specific deletion of Ssu72 leads to Ssu72-mediated hypo-phosphorylation of HNF4α, a master hepatocyte regulator; loss of Ssu72 induces mature hepatocyte-to-progenitor cell conversion (dedifferentiation) orchestrated through HNF4α; Ssu72-deficient mice show high incidence of NAFLD and NASH, and increased probability of HCC in chemical/metabolic HCC models. Conditional liver-specific KO mice, phosphorylation analysis of HNF4α, hepatic progenitor cell assays, chemical and metabolic HCC induction models, histopathology Cell death and differentiation Medium 34616001
2022 Avian influenza NS1 protein directly binds to SSU72; NS1-mediated degradation of SSU72 induces transcriptional readthrough (TRT) at genes on the complementary strand; SSU72 overexpression reduces TRT and alleviates mouse lung injury post-AIV infection; SSU72 restoration suppresses TRT-mediated disruption of STAT1/2 expression, defining an NS1-SSU72-trans-TRT-STAT1/2 axis. Co-immunoprecipitation (NS1-SSU72 binding), SSU72 overexpression in cell lines and mouse lungs, TRT quantification, patient PBMC analysis, STAT1/2 expression analysis Cellular & molecular immunology Medium 35332300
2023 Ssu72 phosphatase dephosphorylates eIF2α in brown adipocytes; adipocyte-specific Ssu72 deletion causes hyperphosphorylation of eIF2α, alteration of cytosolic mRNA translation programs, reduced translation of mitochondrial OXPHOS subunits, mitochondrial dysfunction and defective thermogenesis; cold exposure increases Ssu72 expression in BAT; restoration of Ssu72 expression rescues metabolic dysfunction. Adipocyte-specific conditional KO, in vitro phosphatase assay (eIF2α dephosphorylation), polysome profiling/translation analysis, mitochondrial function assays, cold tolerance test, Ssu72 re-expression rescue Nature communications High 36841836

Source papers

Stage 0 corpus · 44 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
2004 Ssu72 Is an RNA polymerase II CTD phosphatase. Molecular cell 193 15125841
2010 Crystal structure of the human symplekin-Ssu72-CTD phosphopeptide complex. Nature 145 20861839
2002 A role for SSU72 in balancing RNA polymerase II transcription elongation and termination. Molecular cell 141 12453421
2003 Functional interactions between the transcription and mRNA 3' end processing machineries mediated by Ssu72 and Sub1. Genes & development 125 12704082
2010 cis-Proline-mediated Ser(P)5 dephosphorylation by the RNA polymerase II C-terminal domain phosphatase Ssu72. The Journal of biological chemistry 109 21159777
2003 Ssu72 is a phosphatase essential for transcription termination of snoRNAs and specific mRNAs in yeast. The EMBO journal 106 12660165
2003 Ssu72 protein mediates both poly(A)-coupled and poly(A)-independent termination of RNA polymerase II transcription. Molecular and cellular biology 102 12944462
1996 Synthetic enhancement of a TFIIB defect by a mutation in SSU72, an essential yeast gene encoding a novel protein that affects transcription start site selection in vivo. Molecular and cellular biology 86 8657130
2012 Ssu72 phosphatase-dependent erasure of phospho-Ser7 marks on the RNA polymerase II C-terminal domain is essential for viability and transcription termination. The Journal of biological chemistry 85 22235117
1999 Mutational analysis of yeast TFIIB. A functional relationship between Ssu72 and Sub1/Tsp1 defined by allele-specific interactions with TFIIB. Genetics 76 10511545
2000 Functional interaction between Ssu72 and the Rpb2 subunit of RNA polymerase II in Saccharomyces cerevisiae. Molecular and cellular biology 53 11046131
2009 The essential N terminus of the Pta1 scaffold protein is required for snoRNA transcription termination and Ssu72 function but is dispensable for pre-mRNA 3'-end processing. Molecular and cellular biology 50 19188448
2003 The mRNA transcription/processing factor Ssu72 is a potential tyrosine phosphatase. The Journal of biological chemistry 47 12606538
2006 Role for the Ssu72 C-terminal domain phosphatase in RNA polymerase II transcription elongation. Molecular and cellular biology 45 17101794
2012 An unexpected binding mode for a Pol II CTD peptide phosphorylated at Ser7 in the active site of the CTD phosphatase Ssu72. Genes & development 40 23070812
2011 Crystal structure of Ssu72, an essential eukaryotic phosphatase specific for the C-terminal domain of RNA polymerase II, in complex with a transition state analogue. The Biochemical journal 31 21204787
2013 novel modifications on C-terminal domain of RNA polymerase II can fine-tune the phosphatase activity of Ssu72. ACS chemical biology 28 23844594
2005 Conserved and specific functions of mammalian ssu72. Nucleic acids research 27 15659578
2015 Hepatocyte homeostasis for chromosome ploidization and liver function is regulated by Ssu72 protein phosphatase. Hepatology (Baltimore, Md.) 24 26458163
2014 The Ssu72 phosphatase mediates the RNA polymerase II initiation-elongation transition. The Journal of biological chemistry 24 25339178
2014 A gene-specific role for the Ssu72 RNAPII CTD phosphatase in HIV-1 Tat transactivation. Genes & development 20 25319827
2013 Functional interplay between Aurora B kinase and Ssu72 phosphatase regulates sister chromatid cohesion. Nature communications 20 24149858
2017 Ssu72 attenuates autoimmune arthritis via targeting of STAT3 signaling and Th17 activation. Scientific reports 18 28710354
2014 Vertebrate Ssu72 regulates and coordinates 3'-end formation of RNAs transcribed by RNA polymerase II. PloS one 18 25166011
2020 Genetic interactions and transcriptomics implicate fission yeast CTD prolyl isomerase Pin1 as an agent of RNA 3' processing and transcription termination that functions via its effects on CTD phosphatase Ssu72. Nucleic acids research 16 32282918
2020 Ssu72 regulates alveolar macrophage development and allergic airway inflammation by fine-tuning of GM-CSF receptor signaling. The Journal of allergy and clinical immunology 13 32910932
2019 Ssu72 phosphatase is a conserved telomere replication terminator. The EMBO journal 12 30796050
2023 Ssu72 phosphatase is essential for thermogenic adaptation by regulating cytosolic translation. Nature communications 9 36841836
2022 Avian influenza viruses suppress innate immunity by inducing trans-transcriptional readthrough via SSU72. Cellular & molecular immunology 9 35332300
2021 Ssu72 is a T-cell receptor-responsive modifier that is indispensable for regulatory T cells. Cellular & molecular immunology 9 33850312
2019 Functional interaction of human Ssu72 with RNA polymerase II complexes. PloS one 9 30901332
2005 Kinase Cak1 functionally interacts with the PAF1 complex and phosphatase Ssu72 via kinases Ctk1 and Bur1. Molecular genetics and genomics : MGG 9 16362371
2021 Ssu72 Dual-Specific Protein Phosphatase: From Gene to Diseases. International journal of molecular sciences 8 33917542
2021 Ssu72-HNF4α signaling axis classify the transition from steatohepatitis to hepatocellular carcinoma. Cell death and differentiation 8 34616001
2020 Ssu72 Regulates Fungal Development, Aflatoxin Biosynthesis and Pathogenicity in Aspergillus flavus. Toxins 7 33202955
2020 Diverse and conserved roles of the protein Ssu72 in eukaryotes: from yeast to higher organisms. Current genetics 7 33244642
2021 Ssu72 phosphatase directly binds to ZAP-70, thereby providing fine-tuning of TCR signaling and preventing spontaneous inflammation. Proceedings of the National Academy of Sciences of the United States of America 6 34452999
2013 Structurally conserved and functionally divergent yeast Ssu72 phosphatases. FEBS letters 6 23831060
2021 The fission yeast Pin1 peptidyl-prolyl isomerase promotes dissociation of Sty1 MAPK from RNA polymerase II and recruits Ssu72 phosphatase to facilitate oxidative stress induced transcription. Nucleic acids research 5 33410907
2023 Phosphatase Ssu72 Is Essential for Homeostatic Balance Between CD4+ T Cell Lineages. Immune network 2 37179750
2011 ¹H, ¹³C and ¹⁵N backbone and side-chain resonance assignments of Drosophila melanogaster Ssu72. Biomolecular NMR assignments 2 21732054
2024 Ssu72: a versatile protein with functions in transcription and beyond. Frontiers in molecular biosciences 1 38304578
2024 Ssu72 phosphatase deficiency leads to spindle crossing during the second meiotic division process. Yi chuan = Hereditas 0 38886153
2024 Deciphering significant interaction between Clp1 (CF IA) and Ssu72 (CPF) in pre-mRNA processing via in silico approaches. Journal of biomolecular structure & dynamics 0 39522172

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