USP26 Data Analysis

HGNC Gene Name
ubiquitin specific peptidase 26
HGNC Gene Symbol
USP26
Identifiers
hgnc:13485 NCBIGene:83844 uniprot:Q9BXU7
Orthologs
mgi:1933247 rgd:1563443
INDRA Statements
deubiquitinations all statements
Pathway Commons
Search for USP26
Number of Papers
58 retrieved on 2022-05-22

DepMap Analysis

The Dependency Map (DepMap) is a genome-wide pooled CRISPR-Cas9 knockout proliferation screen conducted in more than 700 cancer cell lines spanning many different tumor lineages. Each cell line in the DepMap contains a unique barcode, and each gene knockout is assigned a “dependency score” on a per cell-line basis which quantifies the rate of CRISPR-Cas9 guide drop. It has been found that proteins with similar DepMap scores across cell lines, a phenomenon known as co-dependent genes, have closely related biological functions. This can include activity in the same or parallel pathways or membership in the same protein complex or the same pathway.

We identified the strongest seven co-dependent genes (“Symbol”) for DUBs and ran GO enrichment analysis. We used Biogrid, IntAct, and Pathway Commons PPIDs, and the NURSA protein-protein interaction databases (PPIDs) to determine whether co-dependent genes interact with one another. The “Evidence” column contains the PPIDs in which the interaction appears as well as whether there is support for the association by an INDRA statement. As another approach to identify potential interactors, we looked at proteomics data from the Broad Institute's Cancer Cell Line Encyclopedia (CCLE) for proteins whose expression across ~375 cell lines strongly correlated with the abundance of each DUB; it has previously been observed that proteins in the same complex are frequently significantly co-expressed. The correlations and associated p-values in the CCLE proteomics dataset are provided. And, we determined whether co-dependent genes yield similar transcriptomic signatures in the Broad Institute's Connectivity Map (CMap). A CMap score greater than 90 is considered significantly similar.

DepMap Correlations

Symbol Name DepMap Correlation Evidence CCLE Correlation CCLE Z-score CCLE p-value (adj) CCLE Significant CMAP Score CMAP Type

Dependency GO Term Enrichment

Gene set enrichment analysis was done on the genes correlated with USP26using the terms from Gene Ontology and gene sets derived from the Gene Ontology Annotations database via MSigDB.

Using the biological processes and other Gene Ontology terms from well characterized DUBs as a positive control, several gene set enrichment analyses were considered. Threshold-less methods like GSEA had relatively poor results. Over-representation analysis with a threshold of of the top 7 highest absolute value Dependency Map correlations yielded the best results and is reported below.

GO Identifier GO Name GO Type p-value p-value (adj.) q-value

Transcriptomics

The following table shows the significantly differentially expressed genes after knocking out USP26 using CRISPR-Cas9.

Knockout Differential Expression

Symbol Name log2-fold-change p-value p-value (adj.)
RPL17-C18orf32 RPL17-C18orf32 readthrough 1.08e+00 6.24e-09 1.38e-04
RPL17 ribosomal protein L17 1.07e+00 1.64e-07 1.81e-03

Gene Set Enrichment Analysis

There were too few differentially expressed genes to run a meaningful GSEA.

Literature Mining

INDRA was used to automatically assemble known mechanisms related to USP26 from literature and knowledge bases. The first section shows only DUB activity and the second shows all other results.

Deubiquitinase Activity

psp cbn pc bel_lc signor biogrid lincs_drug tas hprd trrust ctd vhn pe drugbank omnipath conib crog dgi | rlimsp isi tees geneways eidos trips medscan sparser reach
USP26 deubiquitinates AR. 6 / 6
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Taken together, the effects of AR deubiquitinated by USP26 could modulate sperm maturation and spermatogenesis through the androgen receptor signaling pathway.

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Elevated cellular ROS levels might then inhibit USP26 activity to increase the ubiquitination of androgen receptor (AR) and AR splice variant 7 (ARv7) and their ubiquitin and proteasome dependent degradation, which contributed to the increase of Enz sensitivity.

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USP26 deubiquitinates androgen receptor (AR) in the maintenance of sperm maturation and spermatogenesis through the androgen receptor signaling pathway.

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Our data suggest that USP26 assembles with AR and other cofactors in subnuclear foci, and serves to counteract hormone-induced AR ubiquitination, thereby contributing to the regulation of AR transcriptional activity.

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USP26 can deubiquitylate AR in the presence of androgen, while USP10 and 2A-DUB both regulate AR transcriptional activity but do not alter protein stability.

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USP26 deubiquitinates androgen receptor (AR) in the maintenance of sperm maturation and spermatogenesis through the androgen receptor signaling pathway.
USP26 deubiquitinates MDM2. 4 / 4
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USP26 also deubiquitinates and stabilizes Mdm2 [70] thus underlining the complex regulation of AR transcriptional activity.

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USP26 deubiquitinates Mdm2 and stabilizes it.

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USP26 deubiquitinates Mdm2 and stabilizes it.
USP26 deubiquitinates SMAD7. 3 / 3
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USP26 regulates TGF-b signaling by deubiquitinating and stabilizing SMAD7.

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USP26 deubiquitinates SMAD7.

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Examples include ubiquitin specific protease 10 (USP10), which can act on SMAD4 to make it deubiquitinated and stable, further promoting TGF-beta signaling [XREF_BIBR], and USP26, which promotes SMAD7 deubiquitination, thereby amplifying the inhibitory effect of SMAD7 and strengthening the inhibition of the TGF-beta signaling pathway [XREF_BIBR].
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USP26 deubiquitinates PRC1. 2 / 2
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In some cancer types, PRC1 can be deubiquitinated by USP7, USP11 and USP26 [XREF_BIBR, XREF_BIBR].
USP26 deubiquitinates H2AC17. 1 / 1
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Other Statements

psp cbn pc bel_lc signor biogrid lincs_drug tas hprd trrust ctd vhn pe drugbank omnipath conib crog dgi | rlimsp isi tees geneways eidos trips medscan sparser reach
USP26 affects SMAD7
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USP26 activates SMAD7.
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USP26 activates SMAD7. 3 / 3
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Conversely, knockdown of USP26 rapidly degrades SMAD7 resulting in TGF-beta receptor stabilization and enhanced levels of p-SMAD2.

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In contrast, loss of USP26 rapidly degrades SMAD7 stabilizing the TGF-beta receptors leading to enhanced TGF-beta activity as observed by increased levels of phosphorylated SMAD2.

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As loss of USP26 degrades SMAD7 leading to the stabilization of TbetaR, we evaluated the role of TbetaR and SMAD7 on clinical outcome.
USP26 inhibits SMAD7.
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USP26 inhibits SMAD7. 2 / 2
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As observed with USP26 knockdown, ectopic expression of the DUB dead mutant recapitulated the augmentation of SMAD7 induction by TGF-beta whereas wild-type USP26 significantly diminished the induction of SMAD7 in the presence of TGF-beta (Fig XREF_FIG F).

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Therefore, we analyzed whether loss of USP26 increases the levels of SMAD7 Lys48 or Lys63 incorporated ubiquitin chains.
USP26 increases the amount of SMAD7.
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Modified mutated USP26 increases the amount of SMAD7. 1 / 1
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Overexpression of USP26, but not the catalytically inactive mutant USP26 C/S, markedly increased the expression of SMAD7 (Fig XREF_FIG A).
Modified USP26 increases the amount of SMAD7. 1 / 1
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Overexpression of USP26, but not the catalytically inactive mutant USP26 C/S, markedly increased the expression of SMAD7 (Fig XREF_FIG A).
USP26 decreases the amount of SMAD7.
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USP26 decreases the amount of SMAD7. 1 / 1
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In addition, depletion of USP26 significantly reduced the expression of SMAD7 in both the MDA-MB-231 and U373 cell lines (Fig XREF_FIG F and G).

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USP26 expression promotes ESC differentiation.

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Conversely, lesser extent of CBX7 protein binding to the promoters of Cbx4 and Cbx6 is observed during Usp26 mediated ESC differentiation and RA induced ESC differentiation.

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Correlative with the Oct4 gene expression data, occupancy at the Oct4 promoter by USP26, CBX4, CBX6, and H2A-ubi1 was low even during USP26 mediated ESC differentiation.

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Furthermore, infection of ESCs with GFP tagged mouse (m) USP26 led to ESC differentiation in an RA independent manner.

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Using screening of shRNA specific for the DUB subfamily USPs, we identified a novel cellular reprogramming repressor gene Usp26, which blocked somatic cell reprogramming into iPSCs and promoted differentiation by stabilizing PRC1 complexes.

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We also found significant decrease in the gene expression of Cbx7, a Polycomb protein, upon Usp26 induced ESC differentiation.

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These results suggest that USP26 expression promotes ESC differentiation.

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Given that USP26 and USP37 are able to reverse RNF8/168 mediated ubiquitylation and promote HR, these enzymes would be ideal candidates to facilitate the repositioning of ubiquitylated substrates away from the DSB.

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A striking conclusion from our study is that both over-expression as well as knockdown of USP26 or USP37 impairs HR.

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Loss of USP26 or USP37 impairs HR by counteracting RAP80 dependent sequestration of BRCA1.

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Loss of USP26 and USP37 function markedly impairs the assembly of PALB2, RAD51 and efficient HR.

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Depletion of USP26 or USP37 disrupts the execution of HR and this effect is alleviated by the simultaneous depletion of RAP80.

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We reasoned that USP26 and USP37 may antagonize the RAP80 dependent sequestration of BRCA1 by removing RNF8 and RNF168 mediated ubiquitylation and thereby promote HR.
USP26 affects TGFB
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USP26 inhibits TGFB.
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USP26 inhibits TGFB. 5 / 5
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These results suggest that USP26 may potentially act in conjunction with SMAD7 and SMURF2 to downregulate canonical TGF-beta signaling.

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Taken together, these findings confirmed our hypothesis that loss of USP26 enhances TGF-beta signaling and confers poor prognosis in glioblastoma patients.

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Taken together, these data indicate that USP26 inhibits TGF-beta pathway activity by limiting Lys48 ubiquitin mediated degradation of SMAD7 consequently resulting in enhanced ubiquitylation and degradation of the TbetaR complex.

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As knockdown of USP26 potently activated the TGF-beta pathway in breast cancer, we investigated whether depletion of USP26 in the TGF-beta-responsive metastatic cell line MDA-MB-231 enhanced cellular motility and invasion.

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In line with our other models, knockdown of USP26 in this patient derived cell line enhanced TGF-beta activity as evidenced by increased levels of phospho-SMAD2 and increased expression of TGF-beta target genes CTGF, LIF, and SMAD7 (Fig XREF_FIG D and E).
USP26 activates TGFB.
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USP26 activates TGFB. 1 / 1
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USP26 depletion enhances TGF-beta activity and TGF-beta biological responses in breast cancer and glioma.
USP26 affects BRCA1
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USP26 inhibits BRCA1.
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USP26 inhibits BRCA1. 3 / 3
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Together, these results suggest that USP26 and USP37 promote the BRCA1 dependent loading of PALB2 and RAD51 by counteracting the repressive impact of RAP80 dependent BRCA1 sequestration and RAP80 dependent inhibition of end-resection during HR.

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We reasoned that USP26 and USP37 may antagonize the RAP80 dependent sequestration of BRCA1 by removing RNF8 and RNF168 mediated ubiquitylation and thereby promote HR.

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In agreement with this hypothesis, we found that USP26 or USP37 depletion inhibits the efficient association of BRCA1 with PALB2, a unique subunit of the BRCC complex.
USP26 activates BRCA1.
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USP26 activates BRCA1. 3 / 3
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Loss of USP26 or USP37 impairs HR by counteracting RAP80 dependent sequestration of BRCA1.

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On the other hand, we also found that USP26 and USP37 promote the efficient association of BRCA1 with PALB2.

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Together, these results suggest that USP26 and USP37 promote the BRCA1 dependent loading of PALB2 and RAD51 by counteracting the repressive impact of RAP80 dependent BRCA1 sequestration and RAP80 dependent inhibition of end-resection during HR.
TGFB affects USP26
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TGFB increases the amount of USP26.
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TGFB increases the amount of USP26. 5 / 5
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Further experimentation will be required to elucidate the precise mechanism of how TGF-beta enhances USP26 expression.

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In all cell lines tested, TGF-beta enhanced the mRNA expression levels of both USP26 and SMAD7.

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We demonstrate that TGF-beta rapidly enhances the expression of USP26 and reinforces SMAD7 stability by limiting the ubiquitin mediated turnover of SMAD7.

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TGFbeta enhances the expression of USP26 and reinforces SMAD7 stability by limiting the ubiquitin mediated turnover of SMAD7.

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Our data suggest that as part of a negative feedback loop, TGF-beta enhances the expression of not only SMAD7 but also USP26, which acts to deubiquitinate and stabilize SMAD7.
TGFB activates USP26.
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TGFB activates USP26. 1 / 1
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To verify whether USP26 induction by TGF-beta is prevalent in other cell types, we analyzed the induction of USP26 and SMAD7 at early time points in the TGF-beta-responsive breast cancer cell lines MCF7, MDA-MB-231, T47D, and CAL51 and glioma cell lines U373, PCTC, and A172 (Fig XREF_FIG A-G).
USP26 affects CBX4
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USP26 activates CBX4. 2 / 4
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This result suggests that the increased Usp26 expression during ESC differentiation may lead to increased CBX4 and CBX6 protein expression not only by CBX4 and CBX6 protein stabilization but also indirectly by reversing the repression of their gene expression.

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Taken together, these results provide molecular mechanisms of Usp26 mediated increased CBX4 and CBX6 protein stability for suppressing pluripotent gene expression.
USP26 affects UIMC1
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USP26 inhibits UIMC1.
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USP26 inhibits UIMC1. 2 / 2
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We reasoned that USP26 and USP37 may antagonize the RAP80 dependent sequestration of BRCA1 by removing RNF8 and RNF168 mediated ubiquitylation and thereby promote HR.

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Together, these results suggest that USP26 and USP37 promote the BRCA1 dependent loading of PALB2 and RAD51 by counteracting the repressive impact of RAP80 dependent BRCA1 sequestration and RAP80 dependent inhibition of end-resection during HR.
USP26 activates UIMC1.
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USP26 activates UIMC1. 2 / 2
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Loss of USP26 or USP37 impairs HR by counteracting RAP80 dependent sequestration of BRCA1.

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Together, this model may explain how USP26 and USP37 limit the repressive impact of RAP80 on HR.
USP26 affects SOX2
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USP26 inhibits SOX2.
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USP26 inhibits SOX2. 1 / 2
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By contrast, knockdown of Usp26 enhances the efficiency of reprogramming by reactivating Sox2 and Nanog through degradation of CBX4 and CBX6.
USP26 decreases the amount of SOX2.
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Modified USP26 decreases the amount of SOX2. 1 / 1
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Overexpression of Usp26 in ESCs led to decreased mRNA levels of Sox2, Nanog, and Oct4 compared with control cells.
USP26 decreases the amount of SOX2. 1 / 1
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Our results showed that after Usp26 knockdown during Dox induced OSKM mediated MEF reprogramming led to increased mRNA levels of Sox2 and Nanog, but not Oct4, compared with control.
USP26 affects NANOG
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USP26 inhibits NANOG.
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USP26 inhibits NANOG. 1 / 2
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By contrast, knockdown of Usp26 enhances the efficiency of reprogramming by reactivating Sox2 and Nanog through degradation of CBX4 and CBX6.
USP26 decreases the amount of NANOG.
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USP26 decreases the amount of NANOG. 1 / 1
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Our results showed that after Usp26 knockdown during Dox induced OSKM mediated MEF reprogramming led to increased mRNA levels of Sox2 and Nanog, but not Oct4, compared with control.
USP26 activates NANOG.
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Modified USP26 activates NANOG. 1 / 1
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Overexpression of Usp26 in ESCs led to decreased mRNA levels of Sox2, Nanog, and Oct4 compared with control cells.
USP26 affects CBX6
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USP26 activates CBX6. 1 / 3
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Taken together, these results provide molecular mechanisms of Usp26 mediated increased CBX4 and CBX6 protein stability for suppressing pluripotent gene expression.

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The increased ROS levels generated by ABT263 can inhibit USP26 activity to ubiquitinate AR/ARv7, and further sensitize PCa cells to Enz treatment.

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In our current study, we screened several AR related USPs, including USP7, USP12, USP14, USP22, and USP26 and found that the USP26 activity was repressed by the induced cellular ROS, which is responsible for AR/ARv7 protein degradation and Enz sensitivity increase by ABT263.

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An in vitro DUB assay was applied to further detect USP26 activity, and results revealed that ABT263 could significantly decrease USP26 activity, which could be reversed by inhibition of cellular ROS (XREF_FIG H-J).

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Silencing of USP26 significantly enhanced TGF-beta-induced migration and invasion (Fig XREF_FIG A-D).

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In comparison, stable expression of USP26, but not the catalytically inactive mutant USP26 C/S, attenuated invasion of MDA-MB-231 cells following exposure to TGF-beta (Fig XREF_FIG E-G).
| 1

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As knockdown of USP26 potently activated the TGF-beta pathway in breast cancer, we investigated whether depletion of USP26 in the TGF-beta-responsive metastatic cell line MDA-MB-231 enhanced cellular motility and invasion.
USP26 affects POU5F1
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USP26 activates POU5F1.
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Modified USP26 activates POU5F1. 1 / 1
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Overexpression of Usp26 in ESCs led to decreased mRNA levels of Sox2, Nanog, and Oct4 compared with control cells.
USP26 activates POU5F1. 1 / 1
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Correlative with the Oct4 gene expression data, occupancy at the Oct4 promoter by USP26, CBX4, CBX6, and H2A-ubi1 was low even during USP26 mediated ESC differentiation.
USP26 decreases the amount of POU5F1.
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USP26 decreases the amount of POU5F1. 1 / 1
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Our results showed that after Usp26 knockdown during Dox induced OSKM mediated MEF reprogramming led to increased mRNA levels of Sox2 and Nanog, but not Oct4, compared with control.
USP26 affects Ubiquitin
| 2
| 2

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Therefore, we analyzed whether loss of USP26 increases the levels of SMAD7 Lys48 or Lys63 incorporated ubiquitin chains.

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USP26 or USP37 antagonize ubiquitin dependent spreading of RAP80-BRCA1.
USP26 affects RAD51
| 2
USP26 activates RAD51. 2 / 2
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Loss of USP26 and USP37 function markedly impairs the assembly of PALB2, RAD51 and efficient HR.

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Together, these results suggest that USP26 and USP37 promote the BRCA1 dependent loading of PALB2 and RAD51 by counteracting the repressive impact of RAP80 dependent BRCA1 sequestration and RAP80 dependent inhibition of end-resection during HR.
USP26 affects PALB2
| 2
USP26 activates PALB2. 2 / 2
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Together, these results suggest that USP26 and USP37 promote the BRCA1 dependent loading of PALB2 and RAD51 by counteracting the repressive impact of RAP80 dependent BRCA1 sequestration and RAP80 dependent inhibition of end-resection during HR.

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Loss of USP26 and USP37 function markedly impairs the assembly of PALB2, RAD51 and efficient HR.
USP26 affects Lys-Ser
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Mutated USP26 activates Lys-Ser. 2 / 2
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Thus, some KS should originate from XY spermatozoa, and paternal USP26 mutations increase the risk of producing KS offspring.

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Consistent with our mouse results, a clinical study shows that some USP26 variations increase the proportion of XY aneuploid spermatozoa in fertile men, and we identified two families with KS offspring wherein the father of the KS patient harbored a USP26 mutated haplotype, further supporting that paternal USP26 mutation can cause KS offspring production.
USP26 affects SMAD2
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Modified USP26 increases the amount of SMAD2. 1 / 1
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Overexpression of the catalytically inactive USP26 C/S mutant also enhanced the levels of p-SMAD2 whereas ectopic expression of USP26 slightly diminished the levels of phosphorylated SMAD2 (Fig XREF_FIG B).
USP26 increases the amount of SMAD2. 1 / 1
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Conversely, knockdown of USP26 rapidly degrades SMAD7 resulting in TGFbeta receptor stabilization and enhanced levels of p-SMAD2, ultimately enhancing cellular proliferation, invasion, and metastasis [XREF_BIBR].
USP26 affects AR
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USP26 decreases the amount of AR.
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USP26 decreases the amount of AR. 1 / 1
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To further dissect the molecular mechanism (s) how miR-367-3p increases AR expression, according to the screening strategy and bioinformatics analysis, we focused on the 2 miR-367-3p predicted targets, MDM2 and USP26, that were reported to modulate AR protein expression.
USP26 activates AR.
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USP26 activates AR. 1 / 1
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In LNCaP cells, Usp26 depletion by siRNA enhances AR dependent transcription, suggesting it acts on a factor that is negatively regulated by ubiquitination (Dirac and Bernards, 2010).
Hairpins affects USP26
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About 2x10 5 MDA-MB-231 cells expressing either control vector, or hairpins targeting USP26 were seeded per well in a six-well plate.
Cadmium dichloride increases the amount of USP26. 1 / 1
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ctd
No evidence text available
ZHX2 affects USP26
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ZHX2 decreases the amount of USP26. 1 / 1
1 |

biopax:msigdb
No evidence text available

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In LNCaP cells, Usp26 depletion by siRNA enhances AR dependent transcription, suggesting it acts on a factor that is negatively regulated by ubiquitination (Dirac and Bernards, 2010).
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Conversely, lesser extent of CBX7 protein binding to the promoters of Cbx4 and Cbx6 is observed during Usp26 mediated ESC differentiation and RA induced ESC differentiation.
USP26 affects maleate(2-)
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Novel Mutations in X Linked, USP26 Induced Asthenoteratozoospermia and Male Infertility.
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These data suggest that USP26 and USP37 limit the magnitude of BRCA1-A complex assembly in DSB neighboring chromatin by counteracting RNF168 mediated H2A ubiquitylation.

eidos
Moreover , the results from co-immunoprecipitation ( CO-IP ) , immunofluorescence and western blot assays showed that the physiological process due to USP26 interacted with AR and influenced AR deubiquitination , thus upregulating the proteins CCND1 and SPATA46 - which are associated with cell cycle progression and spermatogenesis - as well as decreasing the expression of TP73 .
USP26 affects USP26
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USP26 decreases the amount of USP26. 1 / 1
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We isolated the two hairpin vectors from the USP26 DUB pool, which had previously been demonstrated to most effectively inhibit USP26 expression and tested their ability to knockdown USP26 24.
USP26 affects RNF168
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USP26 activates RNF168. 1 / 1
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Interestingly, the DUBs USP26 and USP27 were found to modulate RNF168 mediated protein ubiquitylation at DSB sites, preventing excessive spreading of RAP80-BRCA1, promoting association of BRCA1 with P[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]

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USP26 promotes esophageal squamous cell carcinoma metastasis through stabilizing Snail.
USP26 affects CBX7
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USP26 decreases the amount of CBX7. 1 / 1
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Our results demonstrate that USP26 can diminish the expression of the Cbx7 gene and inhibit cellular reprogramming.
USP26 affects Ala-Pro
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In contrast, the overexpression of Usp26 decreased the number of AP + colonies, while overexpression of Usp20 increased the number of AP + colonies.
SOX2 affects USP26
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SOX2 activates USP26. 1 / 1
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Sox2 and Nanog mRNA levels, but not Oct4, dramatically increased in Cbx4 or Cbx6 knockdown cells, similar to the increased Sox2 and Nanog mRNA levels in Usp26 knockdown cells.
NANOG affects USP26
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NANOG activates USP26. 1 / 1
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Sox2 and Nanog mRNA levels, but not Oct4, dramatically increased in Cbx4 or Cbx6 knockdown cells, similar to the increased Sox2 and Nanog mRNA levels in Usp26 knockdown cells.
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To this end, we co-transfected HEK293T cells with expression plasmids encoding a constitutively active TbetaR (TRICA) and HA tagged ubiquitin in the presence of shRNA vectors targeting USP26.
GATA6 affects USP26
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GATA6 decreases the amount of USP26. 1 / 1
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biopax:msigdb
No evidence text available
CBX7 affects USP26
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CBX7 inhibits USP26. 1 / 1
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To understand how Usp26 expression is regulated, we assessed the promoter binding activity of Usp26 by PRC1 subunits using the ChIP-qPCR assay during RA induced ESC differentiation, and observed that PRC1 components PCGF2 and Cbx7 occupation as well as H2A-ubi1 modifications decreased significantly in Usp26 promoter, thus indicating that CBX7 containing PRC1 complex is a potential repressor of Usp26.
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ctd
No evidence text available