IndraLab

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"Fig. 3 shows that USP22 promoted cell proliferation via the c-Myc/cyclin D2 pathway, BMI-1 pathway and p53, as evident by the results of knocking down USP22 in HeLa cells."

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"Studies have shown that overexpression of USP22 can enhance the inhibitory effect of cell cycle inhibitors such as p21 and enhance the proliferation of tumor cells, thus promoting the occurrence and development of tumors [XREF_BIBR]."

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"18 In lung cancer, USP22 promotes cell survival and proliferation by upregulating the expression levels of stemness-related markers."

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"XREF_BIBR - XREF_BIBR Zhang and colleagues have demonstrated that ectopic overexpression of USP22 promotes cell proliferation and that suppression of USP22 expression by small hairpin RNA induces cell cycle arrest in human lung cancer cells."

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"Our results indicate that USP22 promotes cell proliferation, migration, invasion, and cell cycle transition, while inhibiting apoptosis in gastric cancer cells."

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"Accordingly, we observed similar patterns in the colony forming efficiency of LvUSP22 transfected PC9 cells and USP22-shRNA transfected H1975 cells, suggesting that USP22 positively regulates cell pro[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"Consistent with these findings, our data demonstrate that USP22 promotes gastric cancer cell proliferation, migration, and invasion in vitro and enhances tumor growth in vivo."

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"USP22 promotes cell proliferation, migration, and invasion in multiple cancers, including glioma, lung adenocarcinoma, thyroid carcinoma, colorectal cancer, and gastric cancer [27–31]."

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"USP22 is also upregulated in many cancer cells and activates the proliferation, migration, and invasion of gastric cancer, colorectal cancer, and breast cancer XREF_BIBR - XREF_BIBR."

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"Previous studies have demonstrated that the USP22 gene promotes the proliferation of esophageal carcinoma and lung breast, colorectal and bladder cancer cells."

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"Downregulation of USP22 inhibited OS cell proliferation, invasion, and EMT in vitro."

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"The overexpression or knockdown of USP22 promoted the proliferation of NPC."

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"As shown in Figure XREF_FIG to XREF_FIG, silencing USP22 significantly inhibited proliferation and generated a smaller number of colonies in Bel/Fu cells."

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"In this study, we found that silencing of USP22 inhibited proliferation of gastric cancer cells and suppressed the cancer stem cell spheroid formation in serum-free culture."

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"In summary, USP22 downregulation reduces cancer cell proliferation, migration, and invasion and reduces tumor growth and metastasis in vivo."

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"In further functional analysis, ADAM9 deletion abolished USP22-triggered suppression in proliferation ( Fig. 5 B–D), promotion in apoptosis ( Fig. 5 E), and weakness in invasion and migration abilitie[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"USP22 silencing inhibits GC cells proliferation."

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"Depletion of USP22 suppressed cell proliferation in vitro and tumor growth in vivo."

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"Together, these data showed that USP22 silencing inhibited the proliferation of ATC cells in vitro ."

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"Consistently, we demonstrated that USP22 silencing inhibited the proliferation of human ATC cells (8505C and CAL-62)."
USP22 affects SIRT1
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USP22 binds SIRT1.
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sparser
"SIRT1 physically interacts with USP22 as a mediator of USP22."

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"USP22 can also form a complex with the histone deacetylase SIRT1, which serves a similar repressive action on Sox2."

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"Moreover, USP22 directly interacts with sirtuin 1 (SIRT1) and positively regulates SIRT1 protein expression, leading to activation of the SIRT1/AKT/ABCC1 pathway and MDR of hepatocellular carcinoma [ xref ]."

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"Sgf73 can also physically interact with Sir2 and Ubp8, a finding replicated in mammals by interaction of USP22 with human SIRT1 ( xref ; xref ), suggesting that altered chromatin-modifying activities in the central nervous system may contribute to SCA7 neurodegeneration."

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"These results indicate that USP22 directly interacts with SIRT1 and contributes to stabilization of SIRT1 protein in FLT3-ITD AML cells."

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"We show that USP22 directly interacts with SIRT1 in FLT3-ITD AML cells in a FLT3-kinase-independent manner."

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"By using Co-immunoprecipitation, the interaction between USP22 and Sirt1 was validated in cells transfected with KLF3-AS1 overexpression plasmid or not (Fig.  xref E)."

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"USP22 Interacts with SIRT1 and Maintains Its Protein Expression."

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"These results indicate that USP22 directly interacts with SIRT1 and contributes to stabilization of SIRT1 protein in FLT3-ITD AML cells."

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"It has been demonstrated that USP22 interacts with Sirt1 and removes Sirt1 polyubiquitination to enhance its protein stability [14] ."
USP22 deubiquitinates SIRT1.
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"Ubiquitin specific protease 22 (USP22) reduced Sirt1 ubiquitination and degradation and decreased FN and TGF-beta1 expression in GMCs under both basal and AGEs treated conditions."

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"Our previous study has already found that USP22 could deubiquitinate and stabilize Sirt1 in GMCs ( Lin et al., 2012 )."

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"As shown in the data in Fig. 3 B, USP22-FLAG rather than USP22–C185S-FLAG could reduce the Sirt1 ubiquitination level, indicating that the USP22 plasmids functioned normally in the EMT model."

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"USP22 can deubiquitinate and stabilize the expression of Sirt1."

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"Meanwhile, USP22 may also antagonize p53 transcriptional activation by deubiquitinating and stabilizing Sirt1 [14] ."

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"Our previous research has shown that USP22 deubiquitinates and stabilizes Sirt1 to alleviate fibrotic changes in GMCs under HG conditions ( Huang et al., 2015 )."

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"In addition, USP22 knockdown prevented c-MYC-mediated reduction of SIRT1 ubiquitination (XREF_FIG) and increase in SIRT1 expression (XREF_SUPPLEMENTARY)."

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"Third, since USP7 has key roles in the p53 tumor suppressor pathway through stabilization of p53 via increasing MDM2 [6, 52], while USP22 is reported to antagonize p53 transcriptional activation by deubiquitinating Sirt1 to suppress [34], we further investigated p53p53 the effect of USP22 and USP7 inhibition on the p53 pathway."

trips
"Ubiquitin-specific peptidase USP22 negatively regulates the STAT signaling pathway by deubiquitinating SIRT1."

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"Knockdown of USP22 reversed the deubiquitination of Sirt1 mediated by KLF3-AS1 (Fig. 4D)."
USP22 activates SIRT1.
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"Collectively, these results revealed that CTRP9 promoted USP22 expression, which removed the conjugated poly-ubiquitin chains from Sirt1 and enhanced the stabilization of Sirt1 protein.As shown in Fig[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"The lncRNA highly upregulated in liver cancer ( HULC ) enhances expression of ubiquitin-specific peptidase (USP22) which increases stability of Sirt1 protein."

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"We found that USP22 promotes multidrug resistance in HCC cells by activating SIRT1/protein kinase B (Akt)/multidrug resistance-associated protein 1 (MRP1) pathway, while inhibition of USP22 and SIRT1 [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"Simply put, USP22 may activate the SIRT1–AKT–MRP1 pathway and consequently promote MDR in human HCC cells (226)."

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"USP22 positively regulates the histone deacetylase Sirt1, resulting in suppression of p53 function by reducing p53 acetylation [58] ."

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"USP22 mediates stabilization of Sirt1 by interacting and removing poly-ubiquitin chains previously conjugated to Sirt1."

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"In our study, after treatment with FO, the expression of USP22 was markedly increased, and the increase in USP22 increased the activity of Sirt1 and decreased myocardial cell death."

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"Our data showed that USP22 overexpression increased the stability of SIRT1, thereby prolonging the half-life of SIRT1 protein (Fig. 4C)."

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"Furthermore, c-Myc-induced upregulation of USP22 deubiquitinase promotes SIRT1 stability and subsequently confers protection against FLT3 inhibitors in FLT3-ITD AML LSCs [136]."

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"In fact, expression of USP22 significantly prolonged the half-life of Sirt1 ( Figures 3 A and 3B )."
USP22 inhibits SIRT1.
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"The USP22 inhibition‐induced decrease in SIRT1 expression was partially reversed by treatment with the proteasome inhibitor MG132, but not the autophagy inhibitor chloroquine (Figure 5G), suggesting that USP22 may inhibit the degradation of SIRT1 through a proteasome‐dependent pathway."

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"USP22 depletion enhanced Sirt1 degradation and displayed combined effects with AGEs to further promote FN and TGF-beta1 expression."

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"USP22 inhibits activation of apoptotic pathways by stabilizing SIRT1."

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"USP22 inhibits p53 activity and transcriptional activation of the p53 target gene by stabilizing SIRT1 through deubiquitination to suppress cell apoptosis [ 29 ]."

sparser
"Moreover, we found that USP22 inhibited the SIRT1 gene to regulate ferroptosis-induced cardiomyocyte death."

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"USP22 Inhibits SIRT1 to Regulate Ferroptosis-Induced Cardiomyocyte Death A previous study has shown that USP22 possesses the ability to stabilize SIRT1 by the process of deubiquitination ( Lin et al ., 2012 ) ."

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"Furthermore, USP22 reduces hepatic steatosis and obesity by stabilizing Sirt1 protein and regulating Sirt1-dependent mitochondrial respiration (145).3 Discussion."

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"This study demonstrated that teneligliptin acts as a negative regulator of PRDX3 acetylation through USP22-mediated SIRT1 activation [12]."

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"Additionally, SIRT1 instability caused by loss of USP22, a deubiquitinating enzyme that stabilizes SIRT1, is associated with the defective embryogenesis in USP22 null mice [XREF_BIBR]."

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"USP22 Inhibits SIRT1 to Regulate Ferroptosis-Induced Cardiomyocyte Death."
USP22 increases the amount of SIRT1.
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"In addition, SIRT1 is a functional mediator of USP22, and USP22 promotes the expression of SIRT1 by directly combining with SIRT1."

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"In addition, USP22 knockdown prevented c-MYC-mediated reduction of SIRT1 ubiquitination (XREF_FIG) and increase in SIRT1 expression (XREF_SUPPLEMENTARY)."

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"USP22, for instance, can directly regulate and upregulate SIRT1 protein levels, which enhances hepatoma cell resistance to sorafenib [20]."

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"USP22 knockdown reduced SIRT1 expression in FLT3-ITD AML cells, whereas USP22 overexpression increased SIRT1 levels by increasing protein stability, indicating that USP22 is an important positive regulator of SIRT1 in FLT3-ITD cells."

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"Depletion of USP22 exhibited no effect on Sirt1 mRNA level, but significantly inhibited Sirt1 protein expression ( Fig. 3 F and G)."

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"USP22 overexpression can increase the SIRT1 protein level and decrease the p53 acetylation level, promoting SLC7A11 expression."

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"Finally, genetic deletion of usp22 gene in mice leads to early embryonic lethality, and loss of USP22 functions causes decreased Sirt1 protein expression.USP22 is critically required for embryogenesis[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"Given that SIRT1 is deubiquitinated by USP22 and stabilized at the protein level (Armour etal., 2013; Lin etal., 2012) and previous studies have reported that SIRT1 could negatively influence the chemosensitivity of HCC cells (Chen etal., 2012), our results supported the notion that USP22 increases SIRT1 protein levels in HCC cells."

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"Moreover, depletion of USP22 or usage of EX527 reduced the Sirt1 protein level and LC3II/LC3I ratio, and enhanced SQSTM1 protein level compared with CTRP9 treatment ( Fig. 4 C)."

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"USP22, a member of the deubiquitinase family, USP22-mediated deubiquitination can stabilize the expression of SIRT1."
USP22 ubiquitinates SIRT1.
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"USP22 negatively regulates STAT signaling by deubiquitinating SIRT1 in colon cancer cells [15] ."

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"USP22 negatively regulates STAT signaling by deubiquitinating SIRT1 [15] ."

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"As shown in the data in Fig. 3 B, USP22-FLAG rather than USP22–C185S-FLAG could reduce the Sirt1 ubiquitination level, indicating that the USP22 plasmids functioned normally in the EMT model."

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"Sirt1 ubiquitination can be modulated by USP22, MDM2 and JNK proteins (Gao et al. 2011; Peng et al. 2015; Xiong et al. 2017)."

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"For example, Lin et al. (2012) proposed that USP22 can suppress TP53 transcriptional activation by deubiquitinating SIRT1 ."

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"Third, since USP7 has key roles in the p53 tumor suppressor pathway through stabilization of p53 via increasing MDM2 [6, 52], while USP22 is reported to antagonize p53 transcriptional activation by deubiquitinating Sirt1 to suppress [34], we further investigated p53p53 the effect of USP22 and USP7 inhibition on the p53 pathway."

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"Additionally, by deubiquitinating Sirt1, the increased expression of USP22 decreased the frequency of p53-dependent apoptosis and oxidative response (Figures 5, 6)."

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"Recent studies have found that USP22 negatively regulates STAT signaling and p53 activation [9] by deubiquitinating Sirt1 [15,16] ."

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"Meanwhile, USP22 may also antagonize p53 transcriptional activation by deubiquitinating and stabilizing Sirt1 [14] ."

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"Furthermore, previous studies have revealed that Sirt1 ubiquitination can be modulated by ubiquitin specific peptidase 22 (USP22), mouse double minute 2 homolog (MDM2) and c-Jun N-terminal kinase (JNK) proteins (Gao et al. 2011; Peng et al. 2015; Xiong et al. 2017)."
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"As shown in the data in Fig. 3 B, USP22-FLAG rather than USP22–C185S-FLAG could reduce the Sirt1 ubiquitination level, indicating that the USP22 plasmids functioned normally in the EMT model."

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"These results indicate that Rottlerin or Morusin is a potent USP22-specific small molecule inhibitor that downregulates Sirt1 and PD-L1 expression in HCT116 cells, which might be important for further[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"In addition, USP22 could also decrease the acetylation of Ku70 by stabilizing the expression of Sirt1, thus inhibiting Bax mediated apoptosis and contributing to cisplatin resistance."

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"USP22 can reduce Ku70 acetylation by stabilizing SIRT1 expression, thereby inhibiting Bax-mediated apoptosis and promoting cisplatin resistance.ALDH1A3, a major ALDH isoenzyme, is important for the stem cell signature of lung cancer and is associated with enhanced cisplatin resistance in lung adenocarcinoma."

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"USP22 suppressed HG-induced ferroptosis in pancreatic beta cells by stabilizing SIRT1 expression."

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"Knockdown of endogenous USP22 in MV4-11 cells decreased SIRT1 protein levels (XREF_FIG) and significantly reduced cell growth (XREF_SUPPLEMENTARY)."
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eidos
"Downregulation of USP22 raised the sensitivity of PC cells to cisplatin , reduced the levels of stem cell markers , reduced the tumor sphere formation and migration , and promoted apoptosis ."

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"As a consequence, gain of USP22 functions promotes cell cycle progression and inhibits cell apoptosis, leading to cancer cell hyper-proliferation and tumorigenesis."

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"Also, USP22 silencing promotes apoptosis and cell cycle arrest in human brain gliomas."

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"Silencing USP22 promoted Bel/Fu cell apoptosis, but had no effect on cell cycle progression."

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"USP22 Silencing Inhibits Proliferation and Induces Apoptosis and Cell Cycle Arrest in NSCLC Cells."

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"As expected, overexpression of USP22 in PC9 cells significantly reduced apoptosis (0.7667 +/- 0.06667% vs. 2.267 +/- 0.5044%, p < 0.05, Fig. 2 D)."

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"Therefore, these data suggest that USP22 decreases cell arrest in G1 phase and reduces apoptosis of EGFR-mutant lung ADC cells in vitro."

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"Overexpression of USP22 in macrophages inhibits foam cell formation, inflammation, and macrophage apoptosis."

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"Finally, we observed that USP22 inhibited macrophage early apoptosis under basal and inflammatory conditions (Fig. 3H–J)."

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"We next explored the pathways by which USP22 silencing promoted HepG2 cell apoptosis."
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"In further functional analysis, ADAM9 deletion abolished USP22-triggered suppression in proliferation ( Fig. 5 B–D), promotion in apoptosis ( Fig. 5 E), and weakness in invasion and migration abilitie[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"Both elevated exosomal miR-let-7a or silenced USP22 reduced the apoptosis of renal cells and improved kidney function."

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"Down-regulation of USP22 expression by siRNA induces the mitochondrial apoptosis of HCC cells."

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"These results suggested that the lack of USP22 could induce apoptosis in Ang II-treated cells and that FO treatment could protect against this reaction."

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"We found that downregulation of USP22 could inhibit Sao-2 cell proliferation, migration, invasion, and induce apoptosis."

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"Therefore, like USP7, accumulating evidence indicates that USP22 also represents a promising target for cancer therapy, and USP22 knockdown markedly decreases cancer growth, induces apoptosis, and sensitizes cancer cells to chemo-radio and immune therapies [17, 34–36]."

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"Downregulation of USP22 induces apoptosis in Sao-2 cells."

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"Additionally, investigation into the underlying mechanism, using small interfering RNA, revealed that the downregulation of USP22 inhibited proliferation and promoted apoptosis though the phosphoinositide 3-kinase/protein kinase B signaling pathway."

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"Under the treatment of Lenvatinib, USP22 knockdown inhibited the cell viability of drug-resistant HCC cells and promoted the apoptosis of drug-resistant cells."

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"Moreover, Usp22 silencing also suppressed renal cell apoptosis, decreased B-cell/CLL lymphoma 2 (Bcl-2)-associated X protein (Bax) mRNA, and concomitantly increased Bcl-2 [137]."
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"In addition, we present evidence that USP22 promotes Bel/Fu cell growth, migration, invasion, EMT and chemoresistance."

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"USP22 plays as an interactor with LINC01426 and activates the hedgehog pathway to promote the invasion and metastasis in lung adenocarcinoma via stabilizing SHH protein [ 48 ]."

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"In further functional analysis, ADAM9 deletion abolished USP22-triggered suppression in proliferation ( Fig. 5 B–D), promotion in apoptosis ( Fig. 5 E), and weakness in invasion and migration abilitie[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"Our results indicate that USP22 promotes cell proliferation, migration, invasion, and cell cycle transition, while inhibiting apoptosis in gastric cancer cells."

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"USP22 knockdown significantly decreased in vitro survival, proliferation, migration, and invasiveness of GC cells compared with the controls."

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"For example, USP22 mentioned above promotes LUAD cell invasion and migration62."

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"In this study, we show that USP22 knockdown significantly decreases migration and invasiveness of GC cells."

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"USP22 silencing inhibits the proliferation, invasion, and EMT of OS cells in vitro ."

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"These results suggest that while USP22 promotes cell migration and invasion, loss of USP22 sensitizes EGFR mutant NSCLC cells to erlotinib in vitro."

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"USP22 promotes cell proliferation, migration, and invasion in multiple cancers, including glioma, lung adenocarcinoma, thyroid carcinoma, colorectal cancer, and gastric cancer [27–31]."

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"USP22 knockdown reduced the overall infiltration of late‐stage exhausted TIM‐3+ CD8 T‐cells and attenuated the upregulatory effect induced by Taz or combination therapy (Figure 6H)."

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"Furthermore, the knockdown of USP22 reduced the infiltration of M2 macrophages, but no significant differences in M2 infiltration were observed with Taz treatment or immunotherapy."

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"Similar results have been reported in liver tumors (19), where ablation of USP22 in liver tumor cells has been shown to increase tumor immunogenicity and promote T-cell infiltration into the resulting liver tumors."

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"Moreover, given that USP22 is positively correlated with M2 macrophage infiltration in public datasets, our study indeed found that knockdown of USP22 can reduce M2‐type macrophage infiltration."

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"Studies have found that USP22 deletion may lead to a significant reduction of MDSCs in the TME and promote the infiltration of T cells and NK cells while the expression of USP22 confers tumor resistance to immunotherapy (64)."

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"USP22 can regulate PD-L1 by deubiquitination and modulate the infiltration of T cells into malignancies 10."

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"99 The downregulation of USP22, according to a prior research, restricts the penetration of myeloid cells and enhances the infiltration of natural killer cells and T cells, hence improving tumor eradication in cancer immunotherapy."

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"We report that deletion of USP22 in pancreatic tumor cells reduced the infiltration of myeloid cells and promoted the infiltration of T cells and NK cells, leading to an improved response to combination immunotherapy."

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"Ablation of USP22 in liver tumor cells has been shown to increase tumor immunogenicity and promote infiltration of T cells into the resulting liver tumors."
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"Downregulation of USP22 reduces in vitro cancer cell proliferation, survival, migration, and invasion, and decreases in vivo tumor growth and metastasis [6, 10–13]."

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"The assay results showed that USP22 downregulation significantly inhibited the proliferation (XREF_FIG) and invasion (XREF_FIG) of U2OS and MG-63 cells."

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"Ablation of USP22 expression remarkably attenuates melanoma migration, invasion, and epithelial-mesenchymal transition in vitro and suppresses melanoma metastasis in vivo."

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"We found that downregulation of USP22 could inhibit Sao-2 cell proliferation, migration, invasion, and induce apoptosis."

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"However, Ao et al (31) reported that USP22 promoted cell proliferation but inhibited cell invasion in SW480 colon cancer through the STAT3/MMP9 pathway (31)."

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"In summary, USP22 downregulation reduces cancer cell proliferation, migration, and invasion and reduces tumor growth and metastasis in vivo."

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"USP22 depletion could significantly inhibit the proliferation and invasion, and promote the apoptosis of ATC cells in vitro."

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"We also suggested that downregulation of USP22 inhibited OS cell proliferation and invasion in vitro."

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"Downregulation of USP22 inhibits migration and invasion of Sao-2 cells."

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"In conclusion, USP22 attenuated the invasion capacity of colon cancer cells by inhibiting the STAT3 and MMP9 signaling pathway."

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"The knockout of USP22 in pancreatic ductal adenocarcinoma cells results in reduced myeloid cells infiltration and increased tumor infiltration of NK cells and T cells, leading to a synergistic response with combined immunotherapy (59)."

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"Increased infiltration of NK cells within the TME is a favorable prognostic factor in several solid tumors [35]; however, USP22 has been reported to suppress NK cell infiltration by altering the transcriptome of pancreatic cancer (PC) cells [36]."

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"Inhibiting USP22 would lead to less activation of immune cells to immunosuppressive microenvironment or more infiltration of immune cells to immune-sensitive status, this issue of deeply associated mechanisms needs to consistently be solved."

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"USP22 in PC cells interacts with SAGA/STAGA to affect the immune microenvironment, and USP22 deficiency promotes T-cell and NK cell infiltration and inhibits tumor metastasis (116)."

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"We report that deletion of USP22 in pancreatic tumor cells reduced the infiltration of myeloid cells and promoted the infiltration of T cells and NK cells, leading to an improved response to combination immunotherapy."

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"The decreased expression of the deubiquitinase USP22 in pancreatic cancer cells promoted the infiltration of natural killer and T cells, thereby enhancing the anti-tumor immune response of the TME.91 Ubiquitination modification also regulates T cells to promote the differentiation of other cells."

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"In mouse models of hepatocellular carcinoma, knockout of Usp22 increases the infiltration of tumor-infiltrating lymphocytes, augments anti-tumor immunity, and synergizes with anti-PD-L1 treatments and chemotherapy (29)."

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"Tumor cell-derived CXCL1 and ubiquitin-specific protease 22 (USP22) were shown to decrease T cell infiltration and response to immunotherapy in PDAC."
USP22 affects CD274
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USP22 binds CD274.
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"USP22 directly interacts with the C-terminus of PD-L1, thereby inducing its deubiquitination and stabilization."

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"USP22 physically interacted with PD-L1."

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"Now that USP22 could regulate PD-L1 protein level, we asked whether USP22 interacted with PD-L1."

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"We performed co-immunoprecipitation assays in HEK293FT and H157 cells respectively, and we found that USP22 interacted with PD-L1 (Fig.  xref a, Additional file xref : Fig. S2A)."

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"Thus, we validated the interaction between USP22 and PD-L1."

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"The loss of USP22 causes PD-L1 to be degraded at the post-translational level, which has been shown to reduce carcinogenesis and increase T cell-mediated cell death. xref PD-L1 targeted immunotherapy and CDDP-based chemotherapy were both shown to benefit from USP22 reduction in another research, highlighting the complex and important functions of the USP22-PD-L1 axis in cancer treatment. xref According to a recent research, the tumor-promoting long noncoding RNA (lncRNA) KCNQ1OT1 controls the ubiquitination of PD-L1 and decreases the response of CD8 + T cells via the miR-30a-5p/USP22 pathway."

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"USP22 may be a particularly attractive target for cancer immunotherapy, because USP22 can limit the effect of anti-tumor immune response in Treg cells, and there is currently evidence that USP22 can i[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"The deubiquitinase USP22 can bind with PD-L1 and enhance its stability, leading to reduced T cell cytotoxicity in tumor cells (76)."

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"In addition, USP22 could facilitated the interaction between CSN5 and PD-L1, and CSN5 promoted the interaction between USP22 and PD-L1."

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"Another study also showed that USP22 interacted with PD-L1, enhancing its stability by removing ubiquitin and preventing its breakdown by the proteasome."
USP22 deubiquitinates CD274.
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"However, our previous study demonstrated that USP22 induces the deubiquitination of PD-L1 and prevents PD-L1 degradation."

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"Deubiquitinase USP22 deubiquitinates PD-L1, preventing its proteasomal degradation (151)."

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"USP22 deubiquitinated PD-L1 and inhibited its proteasome degradation."

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"Among them, USP9X, CSN5, USP22 and USP7 are responsible for the deubiquitination and stabilization of PD-L1 in tumor cells [98–101]."

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"Huang et al. reported that USP22 deubiquitinated PD-L1 and maintained its stabilization, leading to inhibition of anticancer immunity."

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"In HCC, USP22 impaired anticancer immunity by deubiquitinating CD274 44 and enhanced hypoxia-induced stemness via HIF-1α deubiquitination."

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"In addition, USP22 could inhibit the ubiquitination of PD-L1 protein."

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"In addition, USP22 deubiquitinated CD274 leading to the stabilization of its expression, knockdown of USP22 enhanced the anticancer immunity of CD274 in liver cancer [ 31 ]."

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"Contrary, it is reported that PD-L1 can be deubiquitinated by CSN5, USP22, USP7, USP9X, and OTUB1 (Feng et al.)."

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"USP22 deubiquitinates CD274 to suppress anti-cancer immunity."
USP22 activates CD274.
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"Stability, USP22 can promote PD-L1 to remove the ubiquitin chain and prevent it from being degraded by the proteasome [13–16] ."

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"Either CSN5 or USP22 enhanced the binding of PD-L1 with the other one."

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"Additionally, ubiquitin-specific peptidase 22 (USP22) in HCC [141], ubiquitin-specific peptidase 9, X-linked (USP9X) in OSCC [142], and ubiquitin C-terminal hydrolase L1 (UCHL1) in NSCLC [143] deubiquitinate and upregulate the PD-L1 expression.As a ubiquitously expressed protein, CMTM6 binds PD-L1 either at the plasma membrane or in recycling endosomes, repressing lysosome-mediated degradation of PD-L1 and upregulating protein half-time of PD-L1 without affecting the transcription level of PD-L1 [144, 145]."

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"On the one hand, USP22 could directly regulate PD-L1 stability through deubiquitination."

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"Knockdown of USP22 or OTUB1 only significantly reduces PD-L1 abundance in NCI-H358 cells but not in SK-MES-1 cells."

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"By modulating the stability of the CD274 protein by its deubiquitinase activity, USP22 prevents the proteasomal degradation of CD274 [74]."

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"On the one hand, USP22 can directly regulate the stability of PD-L1 through deubiquitination, leading to tumor immune resistance."

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"On the one hand, USP22 can directly regulate PD-L1 stability through deubiquitination [ 81 ]."

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"Our hypothesis is that the poor clinical efficacy of the treatments targeting PD-L1 may be due to the stabilization of PD-L1 mediated by USP22, abolishing the effect of anti-PD-L1 drugs."

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"USP22 knockdown reduces T cell-dependent tumor metastasis, increases the immunogenicity of tumors, increases the lymphatic invasion of tumors and natural killer cell activity [ 82 ], and enhances anti[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 inhibits CD274.
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"USP22 also deubiquitinated and stabilized CDC274 (PD-L1) to decrease the efficacy of CD274 targeted immunotherapy in mice."

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"In addition, USP22 deubiquitinated CD274 leading to the stabilization of its expression, knockdown of USP22 enhanced the anticancer immunity of CD274 in liver cancer [ 31 ]."

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"The loss of USP22 causes PD-L1 to be degraded at the post-translational level, which has been shown to reduce carcinogenesis and increase T cell-mediated cell death.101 PD-L1 targeted immunotherapy and CDDP-based chemotherapy were both shown to benefit from USP22 reduction in another research, highlighting the complex and important functions of the USP22-PD-L1 axis in cancer treatment."

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"By modulating the stability of the CD274 protein by its deubiquitinase activity, USP22 prevents the proteasomal degradation of CD274 [74]."

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"Replacing lysine with arginine in the intercellular domain of PD-L1 prevents USP22 depletion-induced downregulation of PD-L1, suggesting that USP22 stabilizes PD-L1 through deubiquitination of lysine (23)."

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"On the one hand, USP22 can directly regulate PD-L1 degradation through deubiquitination."

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"EZH2 inhibition transcriptionally upregulates USP22 expression, and upregulated USP22 further stabilizes PD-L1."

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"In a liver cancer mouse model, USP22 knockdown significantly enhanced the efficacy of combined PD-L1 targeted immunotherapy and cisplatin by boosting tumor immunogenicity [210]."

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"In LIHC and NSCLC, USP22 acts as a deubiquitinating enzyme for PD-L1, inhibiting the ubiquitin-proteasome degradation pathway of PD-L1 and thus leading to tumor immune escape (Huang et al., 2019; Wang et al., 2020)."
USP22 ubiquitinates CD274.
| 5 2
USP22 ubiquitinates CD274. 7 / 7
| 5 2

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"The tumor-promoting effect of lncRNA KCNQ1OT1 was through the autocrine effect of tumor cell-derived exosomes, which mediates the miR-30a-5p/USP22 pathway to regulate the ubiquitination of PD-L1 and inhibits CD8+ T-cell response, thereby promoting colorectal cancer development.Conclusion: Tumor cell-derived exosomes' KCNQ1OT1 could regulate PD-L1 ubiquitination through miR-30a-5p/USP22 to promote colorectal cancer immune escape."

sparser
"Conclusion Exosomal miR-335-5p promotes ubiquitination of PD-L1 by USP22 through down-regulating USP22, and inhibits TNBC immune escape mediated by PD-L1."

reach
"Among them, the tumor-promoting effect of lncRNA KCNQ1OT1 is through the autocrine effect of tumor cell-derived exosomes, which mediates the miR-30a-5p/USP22 pathway to regulate the ubiquitination of PD-L1 and inhibits CD8+ T-cell response, thereby promoting CRC development (Figure 10)."

reach
"Conclusion Exosomal miR-335-5p promotes ubiquitination of PD-L1 by USP22 through down-regulating USP22, and inhibits TNBC immune escape mediated by PD-L1."

sparser
"Exosomal miR-335-5p suppresses TNBC immune escape mediated by PD-L1 by promoting the ubiquitination of PD-L1 by USP22 via downregulation of USP22 and PD-L2 [ xref ]."

reach
"Exosomal miR-335-5p suppresses TNBC immune escape mediated by PD-L1 by promoting the ubiquitination of PD-L1 by USP22 via downregulation of USP22 and PD-L2 [92]."

reach
"Although PD-L1 has been reported to be ubiquitinated or deubiquitinated by CSN5, USP22, OTUB1, FBXO38, STUB1, SPOP or HRD1 [28], only OTUB1 silencing blocked the regulatory effect of PKP3 on PD-L1 expression (Fig. S6D-I, Fig. 6F-G)."
USP22 increases the amount of CD274.
| 3
USP22 increases the amount of CD274. 3 / 3
| 3

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"Increased USP22 transcription under the influence of PRDM1 upregulates PD-L1 transcription, mediated through SPI1 deubiquitination and degradation, which subsequently causes CD8 + T cell exhaustion [5[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, it was demonstrated that targeting USP22 via the miR-30-5p family inhibited the induction of PD-L1 expression in hypoxic conditions, thus preventing activated Tcells from killing LUAD cells."

reach
"Herein, we found that USP22-induced deubiquitination of SPI1 dominantly enhanced PD-L1 transcription."
| 3 73
| 3 60

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"These findings suggested that USP22 promotes melanoma metastasis both in vitro and in vivo.2.3 Elevated USP22 induces EMT activation in melanoma."

reach
"USP22 knockdown reduces the invasiveness and metastasis of multiple cancers by downregulating pathways driven by the oncoprotein, BMI-1 [27, 29, 33]."

reach
"Oncogenic USP22 supports gastric cancer growth and metastasis by activating c-Myc/NAMPT/SIRT1-dependent FOXO1 and YAP signaling."

reach
"The USP22 increases growth and metastasis of HCC cells via inducing Wnt/β-catenin signaling."

reach
"These results demonstrated that USP22 might promotes melanoma metastasis by inducing EMT activation.2.4 USP22 potentiates melanoma metastasis and EMT through activating PI3K/Akt/mTOR pathway."

reach
"In addition, USP22 silencing blocks OS tumour growth and metastasis in vivo ."

reach
"Meanwhile, USP22 is reported to promote GC distant metastasis (9, 10, 23)."

reach
"Downregulation of USP22 reduces in vitro cancer cell proliferation, survival, migration, and invasion, and decreases in vivo tumor growth and metastasis [6, 10–13]."

reach
"USP22 knockdown reduces T cell-dependent tumor metastasis, increases the immunogenicity of tumors, increases the lymphatic invasion of tumors and natural killer cell activity [ 82 ], and enhances anti[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Both genetic and pharmacological inhibition of USP22 largely impaired breast CSCs self-renewal and prevented their metastasis."
| 13

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"As expected, downregulation of USP22 suppressed OS tumor growth and metastasis in vivo."

reach
"Collectively, these results suggested that USP22 depletion attenuates tumor growth and metastasis of ATC."

reach
"Moreover, USP22 overexpression can promote EMT and TGF-beta expression, whereas depletion of USP22 can reverse EMT and reduce metastasis of lung adenocarcinomas."

reach
"Meanwhile, downregulation of USP22 in anaplastic thyroid carcinoma cells may impede lung metastasis in vivo."

reach
"Similarly, Zhao et al. reported that USP22 depletion suppressed cell survival and proliferation as well as tumor growth and lung metastasis of anaplastic thyroid carcinoma cells XREF_BIBR."

reach
"USP22 inhibition abolished EPI-induced breast cancer metastasis."

reach
"In addition, downregulation of USP22 suppressed OS tumor growth and metastasis in vivo."

reach
"XREF_BIBR Downregulation of USP22 has been shown to suppress osteosarcoma growth and metastasis through PI3K and Akt pathway."

reach
"Moreover, USP22 ablation completely prevented lung metastasis, even in mice with tumors treated with EPI, because the lung luminol fluorescence activity and tumor nodules were both markedly decreased by USP22 deletion (Fig. 2, S to U)."

reach
"Consistent with in vitro findings, downregulation of USP22 in ATC cells impeded tumor growth and lung metastasis in vivo."
USP22 affects MYC
10 1 | 46 13
USP22 activates MYC.
| 29
USP22 activates MYC. 10 / 32
| 29

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"In addition, USP22 expression was shown to increase the abundance of c-Myc and the androgen receptor itself [XREF_BIBR, XREF_BIBR, XREF_BIBR]."

reach
"Furthermore, it has been reported that USP22 positively regulates c-Myc and promotes tumorigenic activity in human breast cancer (22)."

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"48 Consistent with the results of the present study, USP22 has been shown to increase c-Myc stability via deubiquitination to enhance growth and colony formation of BC cells (T47D and MCF7 cells)."

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"21 USP22 overexpression accelerated c‐Myc/NRasGV12‐induced HCC in this study."

reach
"USP22 depletion in the lung cancer cells decreases the transcriptional ability of Myc [4]."

reach
"Decreased levels of USP22 reduce the ability of c-MYC, which can stimulate activation of AP4 to directly or indirectly induce EMT [XREF_BIBR], activating the transcription of its targets [XREF_BIBR, XREF_BIBR]."

reach
"61 In contrast, c-Myc promotes apoptosis and accelerates cell turnover during cellular carcinogenesis, thereby promoting the progression of cells toward increasingly malignant phenotypes.73 Therefore, it is hypothesized that USP22 promotes cancer progression in BC cells by stabilizing c-Myc to promote apoptosis."

reach
"In this study, we found that USP22 overexpression induces c-myc upregulation in both SGC7901 cells and tumor tissue, consistent with previous reports."

reach
"Taken together, USP22 accelerates c‐Myc/NrasGV12‐induced tumorigenesis and promotes tumor progression through an FKBP12/mTORC1/autophagy positive feedback loop in HCC cells."

reach
"Interestingly, a pioneer study from Kim et al. showed for the first time that USP22 promotes breast cancer by stabilizing the proto-oncogene c-Myc [37]."
USP22 binds MYC.
10 | 2 13
10 | 2 13

sparser
"Reciprocal co-IP assays in 293T cells demonstrated that ectopically expressed FLAG-BRD4 interacted with ectopically expressed Myc-USP22 ( Fig. 5 E and F)."

sparser
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected Myc-USP22 and Flag-FoxM1, but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone ( xref )."

sparser
"As shown in Fig. 5 G, the ubiquitination level of BRD4 was decreased when Myc-USP22 was co-transfected into 293T cells."

reach
"The data presented here suggest that the recruitment of USP22 to Myc targets as part of the hSAGA complex may provide this critical function.An outstanding question raised by these studies is the iden[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"It has been reported that SIRT1 protein level is increased in LSCs in both chronic myeloid leukemia (CML) and acute myeloid leukemia (AML), by either transcriptional induction in CML or by c-Myc-USP22 mediated stabilization of SIRT1 protein in AML [ xref ]."

sparser
"These unique cellular effects are mediated through USP22's interaction with c-Myc which enhances c-Myc deubiquitination and reduces intracellular glycolysis [ xref ]."

sparser
"Western blotting analysis detected the Flag-tagged FOXO1 in anti-Myc immunoprecipitants of human embryonic kidney (HEK) 293T cells when myc-USP22 but not empty vector was cotransfected ( xref )."

sparser
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected with Myc-USP22 and Flag-FoxM1 but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone ( xref C)."

sparser
"USP22 interacts with MYC through its N-terminal region containing zinc finger motif and abrogates Fbw7-mediated polyubiquitination and degradation of MYC."

sparser
"These results support a role for USP22 in MYC-mediated increase in SIRT1 protein stabilization, and indicate that FLT3-ITD, c-MYC and USP22 form an oncogenic network that enhances SIRT1 expression and activity in leukemic cells."
USP22 deubiquitinates MYC.
1 | 7
USP22 deubiquitinates MYC. 7 / 8
1 | 7

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"Skp2 is a ubiquitin ligase of Myc, which promotes the polyubiquitination and degradation of Myc during normal lymphocytes G1 to S phase transition [145], and USP22 mediates the deubiquitination of c-Myc to promote breast cancer progression [146]."

reach
"Similar results were obtained in the exogenous Co-IP assays that were carried out on HEK293T cells transfected with Flag-USP22 or/and HA-c-Myc ( Figures 5 C and D), which corroborated that c-Myc is a [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"For example, overexpression of USP22 upregulates the level of oncogenic c-MYC in breast cancer by promoting the deubiquitination of c-MYC, which is closely related with the progression of breast cance[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"It could be through a direct mechanism where USP22 deubiquitylates c-MYC inducing its stabilization and activation, or indirectly through ubiquitin removal from histones at c-MYC target genes, recruitment of other transcriptional machinery or deubiquitylation of proteins important for c-MYC activity."

reach
"It has been established that USP22 is required for c-MYC transcriptional activity, through a direct mechanism where USP22 deubiquitylates c-MYC, causing its stabilization or activation, or indirectly through the recruitment of other transcriptional machinery, or by the removal of ubiquitin from histones at c-MYC target genes."

reach
"USP22 promotes the deubiquitination of c-MYC in breast cancer cells, resulting in an increase in c-MYC level (65)."

reach
"We found that USP22 promotes deubiquitination of c-Myc in several breast cancer cell lines, resulting in increased levels of c-Myc."
USP22 increases the amount of MYC.
| 5
USP22 increases the amount of MYC. 5 / 5
| 5

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"Levels of MYC, a critical transcription factor driver in many cancers, are enhanced by USP22 (REF."

reach
"For example, overexpression of USP22 upregulates the level of oncogenic c-MYC in breast cancer by promoting the deubiquitination of c-MYC, which is closely related with the progression of breast cance[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"These data suggest that USP22 promotes GC progression.USP22 and c-Myc are co-expressed in GC tissues [7]."

reach
"It was found that USP22 positively regulates MYC protein levels and that USP22 promotes cancer progression by targeting k48-linked polyubiquitin chains to deubiquitinate MYC (137)."

reach
"The loss of USP22 can reduce the transcription of Myc and p53 by removing Ub from H2B.27 USP3 and USP21 also have functions in chromatin modification, leading to aberrant gene transcription.28 Loss of USP3 results in defects of the cell cycle, while USP21 can promote regenerative functions in hepatocytes."
USP22 inhibits MYC.
| 3
USP22 inhibits MYC. 3 / 3
| 3

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"Noticeably, overexpression of Usp22 reduced the survival time of the c‐Myc/NRasGV12 mouse model of HCC (Figure 1B)."

reach
"Consistently, overexpression of USP22 stimulates while downregulation of USP22 suppresses breast cancer cell growth, migration and tumorigenesis in a MYC dependent manner.113 USP13."

reach
"Because the Myc protein controls the expression of thousands of other genes, the depletion of cellular USP22 can inhibit Myc function, inhibiting the invasive growth of cancer cells."
USP22 affects TP53
| 1 62 7
USP22 inhibits TP53.
| 1 28 2
USP22 inhibits TP53. 10 / 33
| 1 28 2

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"Usp22 promotes oncogenic c-Myc activation as well as indirectly antagonizes the tumor-suppressive function of p53, and clearly diminishes tumor growth in in vitro and in vivo LLC1 models (55)."

reach
"USP22 Silencing Down-Regulates MDMX and Up-Regulates the p53 Pathway in NSCLC Cells."

reach
"In retinoblastoma, the depletion of USP22 has been shown to induce cancer cell apoptosis by suppressing the TERT/P53 signal pathway ."

reach
"High USP22 expression levels potently suppress the transcriptional activity of p53 that contributes to cancer cell proliferation and inhibition of apoptosis or senescence [6]."

reach
"Additionally, by deubiquitinating Sirt1, the increased expression of USP22 decreased the frequency of p53-dependent apoptosis and oxidative response (Figures 5, 6)."

reach
"USP22 can be inhibited by shRNA, activates the p53 pathway in tumours and downregulates MDMX protein, thereby inducing apoptosis in NSCLC cells (9)."

sparser
"Collectively, our data indicates that USP22 inhibits p53 functions through Sirt1 deacetylase."

reach
"It has been reported that USP22 antagonizes p53 in bladder cancer cells through up-regulating MDM2 [XREF_BIBR]."

reach
"On the bases of these results, we speculated that USP22 silencing activates the p53 pathway in NSCLC cells by post-transcriptional down-regulation of MDMX."

reach
"As for Myc, USP22 depletion blocked the ability of p53 to activate the transcription of its targets."
USP22 decreases the amount of TP53.
| 11
USP22 decreases the amount of TP53. 10 / 11
| 11

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"However, at the protein level, USP22 overexpression significantly increased SIRT1 and inhibited p53 and pSTAT3 expression."

reach
"In addition, USP22 knockdown might downregulate MDM2 and MDMX and upregulate p53 expression, leading to cell cycle arrest [ 64 ]."

reach
"USP22 overexpression can increase the SIRT1 protein level and decrease the p53 acetylation level, promoting SLC7A11 expression."

reach
"In fact, USP22 dose-dependently inhibited the levels of acetylation but not protein expression of p53 ( Figures 4 A and 4B )."

reach
"Furthermore, a positive feedback loop exists between c-MYC and SIRT1, where USP22 increases SIRT1 stability through MYC mediation, concurrently decreasing p53 levels [125]."

reach
"Downregulation of USP22 by siRNA in the HeLa cell line results in the decreased expression of BMI-1, c-MYC and cyclin D2, but increased expression of p53."

reach
"The data indicate that the USP22 gene is involved in the regulation of HeLa cell a cell cycle, and USP22 was demonstrated to play an important role in the regulation of human HeLa cell proliferation.I[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP22 reduces P53 levels by stabilizing SIRT1, thereby inhibiting apoptosis during DNA damage and embryonic development (44)."

reach
"In HeLa cells, knock-down of USP22 could up-regulate the expression of p53, and overexpression of USP22 could down-regulate the expression of p53."

reach
"We found that USP22 silencing in A549 and NCI-H460 cells increased the protein expression of p53, p21 and Bax, the key p53 signal molecules (XREF_FIG A), suggesting that p53 activation plays a role in USP22 silencing induced growth inhibition."
USP22 activates TP53.
| 7 1
USP22 activates TP53. 8 / 8
| 7 1

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"Silencing of USP22 promotes human retinoblastoma cell apoptosis by inhibiting TERT and P53 pathway 36."

reach
"Since Sirt1 is a suppressor for p53 functions, it is possible that USP22 promotes cell proliferation and suppresses apoptosis through regulation of Sirt1 to antagonize p53 function."

reach
"Usp22 promotes oncogenic c-Myc activation as well as indirectly antagonizes the tumor-suppressive function of p53, and clearly diminishes tumor growth in in vitro and in vivo LLC1 models (55)."

reach
"USP22 can also suppress apoptosis and promote cell proliferation by antagonizing p53 function through the regulation of SIRT1[29,30]."

reach
"USP22 promotes hypoxia induced hepatocellular carcinoma stemness by a HIF1alpha and USP22 positive feedback loop upon TP53 inactivation."

reach
"USP22 promotes hypoxia induced HCC stemness by a HIF1alpha and USP22 positive feedback loop on TP53 inactivation."

sparser
"Several studies demonstrated that USP22 silencing could activate p53. xref , xref Because USP22, Ube2d4, and Ube3b were important downstream genes of YWHAZ, we infer that YWHAZ downregulates p53 by activating USP22, Ube2d4, and Ube3b, which make P53 ubiquitination and lead to its proteasomal degradation."

reach
"In TP53-mutated hepatocellular carcinoma, USP22 and HIF-1α promote the stabilization of each other, forming a positive feedback loop (29)."
USP22 deacetylates TP53.
| 7
USP22 leads to the deacetylation of TP53. 7 / 7
| 7

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"The deacetylase activity of Sirt1 is required for its synergy with USP22 in suppressing p53 acetylation, as expression of the HDAC inactive mutant of Sirt1, Sirt1/HY, failed to inhibit p53 acetylation[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Notably, expression of USP22 failed to suppress p53 acetylation in sirt1 knockdown HCT116 cells ( Figure S3 A) or in sirt1 null MEF cells ( Figure 5 C)."

reach
"Therefore, our studies collectively demonstrated that USP22-mediated deubiquitination of Sirt1 inhibits p53 acetylation and transcriptional activity and p53-mediated apoptosis in response to DNA damag[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP22 overexpression can increase the SIRT1 protein level and decrease the p53 acetylation level, promoting SLC7A11 expression."

reach
"Ubiquitin-specific peptidase 22 stabilizes sirtuin 1 (SIRT1) through deubiquitination, and increased SIRT1 expression results in decreased p53 acetylation and protein expression."

reach
"USP22 mediated stabilization of Sirt1 results in decreased levels of p53 acetylation and consequently in suppression of p53 mediated functions."

reach
"As shown in Figure 5 A , USP22 and Sirt1 synergistically inhibited p53 acetylation."
USP22 increases the amount of TP53.
| 6
USP22 increases the amount of TP53. 6 / 6
| 6

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"USP22 also play roles in cell cycle regulation, where depletion of USP22 increases the expression of p53 and p21, inhibits proliferation, and induces cell cycle arrest at G1 phase."

reach
"USP22 siRNA transfection increased the expression of p53 and p21, decreased cyclin E expression in bladder cancer cells."

reach
"In HCT116 cells, downregulation of USP22 could up-regulate the expression of p53 ( Lv et al., 2011 )."

reach
"In HeLa cells, knock-down of USP22 could up-regulate the expression of p53, and overexpression of USP22 could down-regulate the expression of p53."

reach
"In contrast, the expression of p53 was increased by Ang II infusion, and si-USP22 treatment further increased the expression of p53 (Figure 6A)."

reach
"The loss of USP22 can reduce the transcription of Myc and p53 by removing Ub from H2B.27 USP3 and USP21 also have functions in chromatin modification, leading to aberrant gene transcription.28 Loss of USP3 results in defects of the cell cycle, while USP21 can promote regenerative functions in hepatocytes."
USP22 binds TP53.
| 4
| 4

sparser
"In addition, the identified USP22-p53 interaction is of potential biological significance."

sparser
"USP22 appears to form a complex with Sirt1 and p53 to suppress p53 acetylation, since we detected the interaction of USP22 with p53 in transiently transfected HEK293 cells ( Figure 5 E)."

sparser
"However, the interaction of USP22 with p53 was only detected in wild-type but not in sirt1 null MEF cells ( Figure 5 F)."

sparser
"Therefore, the interaction of USP22 with p53 is probably mediated by Sirt1."
USP22 acetylates TP53.
| 3
USP22 leads to the acetylation of TP53. 3 / 3
| 3

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"The USP22-mediated accumulation of SIRT1 inhibits p53 acetylation and its transcriptional activity."

reach
"Collectively, our data indicates that USP22 inhibits p53 functions through Sirt1 deacetylase.To further prove that USP22 acts on p53 by stabilizing Sirt1, we tested whether inactivation of USP22 incre[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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"Therefore, USP22 suppresses p53 transcriptional activity and proapoptotic functions.Our finding that USP22 inhibits the p53 transcriptional activity and stabilizes Sirt1 suggests that USP22 inhibits p[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects Neoplasms
| 7 58
USP22 activates Neoplasms.
| 6 43
| 6 43

reach
"Moreover, the depletion of USP22 was shown to decrease in vivo tumor growth ."

reach
"We performed rescue experiments to test whether ALKBH5’s tumor-promoting function may be mediated by USP22 and RNF40."

reach
"In summary, USP22 downregulation reduces cancer cell proliferation, migration, and invasion and reduces tumor growth and metastasis in vivo."

reach
"Other studies also suggested that USP22 promotes tumor migration by promoting procancer pathways such as Wnt/β-Catenin and signal transducer and activator of transcription 3 (STAT3)/MMP9 [ 73 , 74 ]."

reach
"Knockdown of USP22 in an in vivo model was shown to decrease tumor angiogenesis, impair non-homologous DNA damage repair pathways and significantly improve the therapeutic efficacy of cisplatin."

reach
"Similar results have been reported in liver tumors (19), where ablation of USP22 in liver tumor cells has been shown to increase tumor immunogenicity and promote T-cell infiltration into the resulting liver tumors."

reach
"Downregulation of USP22 reduces in vitro cancer cell proliferation, survival, migration, and invasion, and decreases in vivo tumor growth and metastasis [6, 10–13]."

reach
"USP22 has been reported to promote tumor progression by participating in the mediation of DNA repair processes [38]."

reach
"For example, in hepatocellular carcinoma USP22 upregulates PD‐L1 and induces the exhaustion of tumor infiltrating CD8+ T cells, 32 while in the colorectal carcinoma it targets BMI‐1, c‐MYC, CYCLIN D2, p16INK4a, and p14ARF, so exerting a crucial role in tumor proliferation."

eidos
"Apart from activating oncogenes such as BMI-1 and c-MYC , USP22 can inhibit the expression of tumor suppressors such as TP53 through ubiquitination , thus promoting proliferation of tumors [ 87 ] ."
USP22 inhibits Neoplasms.
| 1 15
| 1 15

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"The knockout of USP22 in pancreatic ductal adenocarcinoma cells results in reduced myeloid cells infiltration and increased tumor infiltration of NK cells and T cells, leading to a synergistic response with combined immunotherapy (59)."

reach
"Moreover, genetic depletion of USP22 increased tumor immunogenicity and tumor-infiltrating lymphocytes."

reach
"Notably, although knocking down USP22 only mildly reduced tumor growth, it significantly increased the sensitivity of tumors to Taz."

reach
"In addition to its cancer cell-intrinsic roles, USP22 has been recently discovered to suppress tumor immunosurveillance through potentiating Foxp3 + regulatory functions as well as upregulating the expression of checkpoint receptors PD-L1 and CD73 ."

reach
"Usp22 promotes oncogenic c-Myc activation as well as indirectly antagonizes the tumor-suppressive function of p53, and clearly diminishes tumor growth in in vitro and in vivo LLC1 models (55)."

reach
"USP22 also inhibits tumor proliferation via contributing to the deubiquitination of PTEN (Ren et al., 2022)."

reach
"USP22 was found to positively regulate stable FOXP3 expression and reduce tumor volume when ablated in Tregs."

reach
"Moreover, in NSCIC (86), USP22 deletion can promote the therapeutic effect of PD-L1-targeted tumor immunotherapy."

reach
"17 About the possible relationship among BMI1, EZH2, and USP22, it has been reported that USP22 increases BMI1: in gastric cancer, USP22 inactivation decreases the neoplasm growth, while high levels of BMI1 accelerate the cell cycle via INK4a/ARF and AKT."

reach
"Interestingly, it was demonstrated that silencing USP22 inhibited proliferation in bladder cancer cells (Lv et al., 2011) leading to cell cycle arrest, inhibition of autophagy and decreased tumor progression representing a potential target for cancer therapy."
SIRT1 affects USP22
18 | 1 17 29
SIRT1 binds USP22.
18 | 12 29
18 | 12 29

sparser
"SIRT1 physically interacts with USP22 as a mediator of USP22."

reach
"USP22 can also form a complex with the histone deacetylase SIRT1, which serves a similar repressive action on Sox2."

sparser
"Moreover, USP22 directly interacts with sirtuin 1 (SIRT1) and positively regulates SIRT1 protein expression, leading to activation of the SIRT1/AKT/ABCC1 pathway and MDR of hepatocellular carcinoma [ xref ]."

sparser
"Sgf73 can also physically interact with Sir2 and Ubp8, a finding replicated in mammals by interaction of USP22 with human SIRT1 ( xref ; xref ), suggesting that altered chromatin-modifying activities in the central nervous system may contribute to SCA7 neurodegeneration."

reach
"These results indicate that USP22 directly interacts with SIRT1 and contributes to stabilization of SIRT1 protein in FLT3-ITD AML cells."

reach
"We show that USP22 directly interacts with SIRT1 in FLT3-ITD AML cells in a FLT3-kinase-independent manner."

sparser
"By using Co-immunoprecipitation, the interaction between USP22 and Sirt1 was validated in cells transfected with KLF3-AS1 overexpression plasmid or not (Fig.  xref E)."

sparser
"USP22 Interacts with SIRT1 and Maintains Its Protein Expression."

sparser
"These results indicate that USP22 directly interacts with SIRT1 and contributes to stabilization of SIRT1 protein in FLT3-ITD AML cells."

sparser
"It has been demonstrated that USP22 interacts with Sirt1 and removes Sirt1 polyubiquitination to enhance its protein stability [14] ."
SIRT1 activates USP22.
| 1 3
SIRT1 activates USP22. 4 / 4
| 1 3

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"This suggested SIRT1 might be a mediator of miR-144-3p/USP22 in regulating the ferroptosis of INS-1 cells."

reach
"In addition, SIRT1 is a functional mediator of USP22, and USP22 promotes the expression of SIRT1 by directly combining with SIRT1."

eidos
"Interestingly , Sirt-1 also activates USP22 expression through activation of c-Myc [ 34 ] , suggesting that a positively regulatory loop exists between Sirt-1 and USP22 ."

reach
"On the other hand, by downregulating USP22, the RAS system can reduce Sirt 1 protein expression (Wang et al., 2021)."
SIRT1 inhibits USP22.
| 2
SIRT1 inhibits USP22. 2 / 2
| 2

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"Mechanistically, KLF3-AS1 inhibited the ubiquitination of Sirt1 protein through inducing USP22."

reach
"In particular, SIRT1 stabilization is due to the overexpression, in HCC, of lncRNA HULC (highly upregulated in liver cancer) through a mechanism in which HULC inhibits SIRT1 ubiquitination and degrada[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects BMI1
3 1 | 27 30
USP22 binds BMI1.
3 | 5 30
3 | 5 30

sparser
"Meanwhile, USP22 and BMI1 co-activation may be associated with GC progression and poor prognosis [ xref ]."

sparser
"USP22 and BMI1 were positively associated with TNM staging (Figure xref )."

sparser
"Overexpression of USP22 and BMI1 was previously associated with GC progression and therapy failure through clinical specimen analysis, and our study is consistent with these results."

sparser
"In this study, we hypothesized RNF220 promoted the stemness and progression of colon cancer cells via the USP22-BMI1 axis, with the aim to investigate the specific role and related mechanism of RNF220 in colon cancer."

sparser
"USP22 and BMI1 together form a multiprotein complex acting on their target homologous gene clusters."

sparser
"USP22-BMI1 silences the Hox gene and increases tumor resistance."

sparser
"In this study, we found that RNF220 promoted the stemness and progression of colon cancer cells via the USP22-BMI1 axis."

sparser
"These results indicate that USP22 interacts with BMI1 and protects BMI1 from degradation through a deubiquitination mechanism in glioma cells."

sparser
"Furthermore, our in vivo model suggested that RNF220 promoted tumor growth by regulating the USP22-BMI1 axis."

reach
"USP22 interacts with and deubiquitinates BMI1 for post-translational stabilization."
USP22 activates BMI1.
| 10
USP22 activates BMI1. 10 / 10
| 10

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"USP22 also upregulates BMI-1 and may upregulate Survivin via BMI-1 overexpression (20)."

reach
"Moreover, WT-USP22 prolonged GFP-BMI1 half-life compared with that in cells transfected with control vector or CI-USP22."

reach
"17 About the possible relationship among BMI1, EZH2, and USP22, it has been reported that USP22 increases BMI1: in gastric cancer, USP22 inactivation decreases the neoplasm growth, while high levels of BMI1 accelerate the cell cycle via INK4a/ARF and AKT."

reach
"USP22 maintains gastric cancer stem cell stemness and promotes gastric cancer progression by stabilizing BMI1 protein."

reach
"Ma et al (19) demonstrated that USP22 maintained gastric cancer cell stemness and promoted gastric cancer progression by stabilizing the Bmi-1 protein (19)."

reach
"USP22 promotes stem cell-like features of NSCLC cells by regulating BMI1 signaling (78)."

reach
"An in vitro study showed that the upregulation of USP22 mediated the enhanced expression of BMI1 and Cyclin D2, and was responsible for increased cell proliferation and the metastatic behavior of colon cancer cells ."

reach
"Previous studies have confirmed that USP22 can promote the biological process of NSCLC cells by regulating BMI-1 and AKT signaling pathway [XREF_BIBR]."

reach
"Additionally, USP22 silencing led to down-regulated BMI1."

reach
"In this study, we demonstrated that USP22 mediated protein stabilization of BMI1 promotes gastric CSC stemness maintenance and GC progression, thereby providing a rationale for USP22 targeting as a potential therapeutic approach against GC."
USP22 increases the amount of BMI1.
| 8
USP22 increases the amount of BMI1. 8 / 8
| 8

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"We have reported that USP22 mediates cell survival and proliferation by promoting the expression of BMI-1 and upregulation of activated AKT pathway in colon cancer cells."

reach
"It is posited that increased USP22 expression contributes to 5-Fu resistance in HCC cells by elevating BMI1 expression."

reach
"We found USP22 silencing led to decreased expression of BMI1, as well as decreased P21 levels."

reach
"Our results showed that USP22 silencing significantly down-regulated BMI1 protein expression and further affected gastric CSC self-renewal."

reach
"Above, we demonstrated that USP22 silencing predominantly decreases the BMI1 protein levels rather than mRNA expression, and further alters GC cell proliferation, gastric CSC formation and maintenance of stem cell stemness, indicating post-transcriptional regulation of BMI1."

reach
"USP22 knockdown inhibits glioma stemness partially by downregulating the protein level but not the transcriptional level of BMI1."

reach
"USP22 knockdown remarkably decreased BMI1 protein level, but it barely affected BMI1 mRNA level in glioma cell lines."

reach
"USP22 silencing significantly downregulated BMI1 protein expression and further affected gastric CSC self-renewal."
USP22 deubiquitinates BMI1.
1 | 4
USP22 deubiquitinates BMI1. 4 / 5
1 | 4

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"To test whether the endogenous BMI1 is also deubiquitinated by USP22 in glioma cells, we knocked down endogenous USP22 in U251 cells pretreated with CHX and found that endogenous BMI1 also became unstable and degraded rapidly."

reach
"Our work also discovered PRC1 component Bmi1 deubiquitinated and stabilized by USP22 in glioblastoma cells (data not published)."

reach
"USP22 interacts with and deubiquitinates BMI1 for post-translational stabilization."

reach
"3.3 USP22 deubiquitinates BMI1 for protein stabilization."
USP22 affects AR
2 | 1 40 5
USP22 activates AR.
| 1 21
USP22 activates AR. 10 / 28
| 1 21

reach
"USP22 promotes lethal tumor phenotypes by modulating androgen receptor accumulation and oncogenic signaling in prostate cancer [ 25 ]."

reach
"Overexpression of USP22 enhanced AR protein accumulation, which in turn activated downstream target genes regulated by AR and MYC."

reach
"USP22 Enhances AR Activity and Promotes Bypass of AR Antagonists."

reach
"By contrast, USP22 deregulation induced marked enhancement of ligand independent AR activity, determined by analyses of multiple, clinically relevant AR target genes (XREF_FIG)."

reach
"Moreover, USP22 and DHT acted cooperatively to further enhance AR activity (XREF_FIG)."

reach
"These data demonstrate that USP22 potentiates both ligand dependent and ligand independent AR function."

reach
"These data put forth the provocative hypothesis that USP22 increases the activity of AR through altering AR protein levels, thus identifying a novel role for USP22 in modulating steroid receptor function."

reach
"Strikingly, USP22 deregulation significantly increased AR occupancy at known ARORs (1.15-2.6% input; XREF_FIG), but not in control regions of the KLK3 and PSA (' EF ' region, XREF_SUPPLEMENTARY)."

reach
"Together, these data demonstrate that USP22 regulates ligand independent AR residence at target gene loci and promotes AR driven CRPC gene profiles, which may have specificity for USP22 perturbation, further implicating USP22 as an independent effector of aggressive tumor phenotypes."

reach
"In addition to the role of USP22 in the cell cycle, USP22 promotes the activity of the androgen receptor (AR)."
USP22 binds AR.
2 | 6 5
2 | 6 3

reach
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Cotransfection in HEK293T cells and coimmunoprecipitation experiments revealed that AR interacted with ATXN7L3 in a ligand-dependent manner, similarly to GCN5, while interaction between AR and USP22 w[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Subsequent immunoprecipitation also confirmed the association between AR and USP22 in vitro, which is consistent with the result of previous studies (Fig. 3I)."

sparser
"As AR and USP22 interacted in vivo, we next tested whether AR could be a substrate of the TFTC/STAGA deubiquitination activity."

sparser
"No significant association of AR and USP22 with chromatin at an intergenic region could be detected, and the amount of histone H3 at KLK2 promoter remained constant, further demonstrating specificit[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, our immunofluorescence analysis showed that Importin-7 binds to AR or USP22 in both the nucleus and cytoplasm (Fig. 4C–E), suggesting that Importin-7 may regulate the nuclear translocation of AR and USP22.Next, we conducted a more in-depth study of the Importin-7–ARUSP22 complex by the treatment of AR inhibitor (enzalutamide) or AR ligands, including DHT and synthetic androgen R1881."

reach
"Here, to further investigate whether the arrest of the nuclear translocation of ARUSP22 complex triggered by the knockdown of Importin-7 could enhance the sensitivity of BC to enzalutamide.We first confirmed that the knockdown of Importin-7 could enhance the growth inhibition of cancer cells by enzalutamide using CCK8 assay (Fig. 6A, B) and EdU staining assay (Fig. 6C, D)."

reach
"Interestingly, we found that USP22, one of the maintainers of AR [22–24], whose nuclear translocation is also regulated by Importin-7, although knockdown of Importin-7 may not affect the formation of the ARUSP22 complex.In addition, we also observed that AR expression levels were upregulated after DHT and R1881 stimulation, and Importin-7-bound AR was also up-regulated to an almost equal extent, indicating that the level of AR carried by Importin-7 was not saturated."

reach
"Cotransfection in HEK293T cells and coimmunoprecipitation experiments revealed that AR interacted with ATXN7L3 in a ligand dependent manner, similarly to GCN5, while interaction between AR and USP22 w[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
AR binds USP22 and ATXN7L3. 2 / 2
| 2

sparser
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR-dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"To further investigate this hypothesis, we analyzed putative interactions of USP22 or ATXN7L3 with AR in mammalian cells."
USP22 inhibits AR.
| 5
USP22 inhibits AR. 5 / 7
| 5

reach
"First, depletion of USP22 significantly reduced AR protein (~ 73%) compared to control (XREF_FIG, compare lanes 3,4 and 7,8)."

reach
"As shown, in CRPC cells, USP22 depletion suppressed AR protein accumulation in both the androgen stimulated and deprived conditions (XREF_FIG, compare lanes 1,2 and 3,4)."

reach
"In androgen deprived conditions, USP22 depletion modestly decreased basal AR activity."

reach
"Additionally, in both culture conditions, USP22 depletion diminished AR activity (XREF_FIG)."

reach
"It is worth noting that the knockdown of USP22, although leading to an insignificant down-regulation of AR, allows Importin-7 to bind more AR, suggesting that USP22 is possibly one of the cargoes of Importin-7 (Fig. 4H)."
USP22 increases the amount of AR.
| 4
USP22 increases the amount of AR. 4 / 5
| 4

reach
"The observations that USP22 upregulation is sufficient to promote ligand independent AR expression and activity, and induce Casodex resistance are clinically relevant, as these attributes reflect key biochemical characteristics of CRPC."

reach
"Further, USP22 mediates AR expression through a proteasome dependent mechanism, since modeling clinically-relevant USP22 upregulation results in an increased AR protein half-life and proteasome inhibition rescued the decreased AR expression following USP22 depletion."

reach
"Mechanistically, USP22 promotes the progression of PCa by stabilizing the AR protein levels and regulating the AR-Myc signaling pathway (Schrecengost et al., 2014)."

reach
"And correspondingly, overexpression of USP22 stabilized AR-V7 expression in a manner similar to USP14 (Fig. 5A–D), demonstrating that USP22 may also be involved in stabilizing the protein level of AR-V7, similar to USP14.In fact, the regulation of AR expression and function by USP22 in prostate cancer has been reported [37], but whether AR-V7 could be regulated by USP22 is elusive."
USP22 decreases the amount of AR.
| 4
USP22 decreases the amount of AR. 4 / 5
| 4

reach
"USP22 Depletion Suppresses Ligand Dependent and Castration Resistant AR Expression and Activity."

reach
"Conversely, depletion of USP22 dramatically down-regulates AR protein levels and abrogates basal and DHT stimulated AR activity in both ADT sensitive and CRPC cells."

reach
"In contrast, USP22 depletion reduced AR protein levels in the absence of androgen, and inhibited DHT induced AR expression, within in models of therapy sensitive disease (XREF_FIG, compare lanes 2, 4)."

reach
"Schrecengost et al XREF_BIBR previously showed that USP22 depletion dramatically downregulated androgen receptor protein levels and abolished androgen receptor activity in both androgen deprivation therapy and castration resistant prostate adenocarcinoma cells."
USP22 is modified
1 | 43 11
USP22 is phosphorylated.
| 23 11
USP22 is phosphorylated. 10 / 34
| 23 11

sparser
"Our data suggest that ATK-mediated phosphorylation of USP22 is critical for its stabilization because mutation of the phosphorylation residues increased USP22 ubiquitination and facilitated its proteasomal degradation."

sparser
"Moreover, cyclin B1-Cdk1 leads to USP22 phosphorylation on residues Thr147 and Ser237, which in turns promotes its deubiquitinase activity."

sparser
"Notably, USP22 phosphorylation levels are positively associated with EPI and with downstream pathways involving both FOXO1 and ATGL in breast cancers."

sparser
"Only weak USP22 phosphorylation was detected in HCT116 cells, presumably catalyzed by endogenous CDK1, and CDK1 overexpression significantly increased USP22 phosphorylation."

sparser
"In addition, expression of the constitutively active form of CDK1 (CDK1/AF) [ xref , xref ] further enhanced USP22 phosphorylation in HCT116 cells ( xref )."

sparser
"Moreover, mutation of the aspartic acid at position 146 of CDK1 (CDK1/D146N), which is a kinase-inactive mutant [ xref ], completely abolished its ability to promote USP22 phosphorylation ( xref )."

sparser
"In an in vitro kinase assay, co-incubation of the purified GST-USP22 fusion proteins and the constitutive forms of CDK1 (CDK1/AF) immunoprecipitated from transiently transfected HCT116 cells confirmed USP22 phosphorylation by CDK1 ( xref )."

sparser
"Endogenous USP22 was phosphorylated in G2/M phase, but not in G1 phase, as the phosphorylated forms of USP22 were detected in cells treated with thymidine-nocodazole but not double thymidine ( xref )."

sparser
"These findings show that CDK1 is a kinase that catalyzes USP22 phosphorylation in a cell cycle-specific manner."

sparser
"These results indicate that T147 and S237 are the potential CDK1 phosphorylation sites and imply that these residues have important functions in CDK1-mediated USP22 phosphorylation."
USP22 is ubiquitinated.
| 11
USP22 is ubiquitinated. 10 / 11
| 11

sparser
"Fourth, USP22 is ubiquitinated and degraded by CDC20-containing APC/C complex during cell exit from M phase, presumably to release the brake on CCNB1 degradation."

sparser
"Besides, USP22 could remove poly-ubiquitin chain of CSN5, while CSN5 seemed no effect on USP22 ubiquitination (Fig. xref c, Fig. xref d)."

sparser
"Stimulation with EPI markedly reduced USP22 ubiquitination levels ( xref ), supporting our speculation that EPI protects USP22 from ubiquitination-mediated degradation."

sparser
"Lastly, ubiquitylation of USP22 mediated by the anaphase promoter complex/cyclosome (APC/C) induces USP22 protein degradation during the cell cycle [ xref ]."

sparser
"Our data suggest that ATK-mediated phosphorylation of USP22 is critical for its stabilization because mutation of the phosphorylation residues increased USP22 ubiquitination and facilitated its proteasomal degradation."

sparser
"Among these differentially ubiquitylated USP22 targets numerous DNA repair proteins, including XPC and RAD23B, were identified."

sparser
"Conversely, treatment with the AKT-specific inhibitor facilitated USP22 protein degradation ( xref ), increased the USP22 ubiquitination ( xref ), and inhibited the AKT and USP22 interaction in breast cancer cells ( xref )."

sparser
"Interestingly, we also observed Morusin caused conformational changes in the active pocket residues 255–262 after MD-50 ns, indicating that Morusin may inhibit USP22 ubiquitination activity through al[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Loss of AKT-mediated phosphorylation markedly increased USP22 ubiquitination and abolished EPI-induced inhibitory efficacy ( xref )."

sparser
"However, of those four DUBs, only USP22 substantially decreased KDM1A ubiquitination ( xref )."
USP22 is acetylated.
1 | 9
USP22 is acetylated. 7 / 7
| 7

sparser
"These results suggest that USP22 acetylation, incorporation into SAGA, or its deubiquitinase activity is unlikely contribute to its role as a barrier to reprogramming."

sparser
"USP22 is acetylated [ xref ], and USP7 is phosphorylated [ xref ]."

sparser
"In an attempt to separate the HAT activity from the DUB interaction domain to assess the contribution of both acetylation and deubiquitylation of PCAF and USP22, we generated an enzymatically dead PCAF by installing two mutations in the core HAT domain (PCAF-YFAA [Y616A/F617A] mutant) (; xref )."

sparser
"Furthermore, our data suggest that ATAC and SAGA control of gene transcription may be more specific than control of promoter H3K9 acetylation, indicating that specificity of transcriptional regulation by either complex may rely on unique effects of their additional enzymatic activities, namely H2B deubiquitination by USP22 and H4 acetylation by KAT14. xref Accordingly, our inspection of USP22 requirements in erythropoiesis suggest that they align with the stage specificity of SAGA but may exceed the effects of loss of SUPT20H and more directly resemble postcommitment contributions of KAT2A . Future studies investigating single and combined requirements of ATAC and SAGA enzymatic subunits in locus and cellular regulation will enhance our understanding of the transcriptional control of cell state and fate decisions, including in leukemia and the normal blood system."

sparser
"Interestingly, SIRT1 was identified as a mediator of acetylation of USP22 and the SAGA coactivator complex (Armour et al ., xref )."

sparser
"Interestingly, SIRT1 was also identified as a mediator of acetylation of USP22 and the SAGA coactivator complex [ xref ]."

sparser
"SIRT1 is a mediator of acetylation of the USP22 and SAGA coactivator complex."
USP22 is acetylated on K129. 2 / 3
1 | 2

sparser
"Acetylation of USP22 on lysine 129 (K129) regulates USP22 deubiquitylase activity and its association with the SAGA complex."

sparser
"Acetylation of K129 on USP22 can regulate its enzymatic activity as well as its binding to GCN5 xref ."
USP22 affects cell cycle
| 3 43
USP22 activates cell cycle.
| 3 28
| 3 28

reach
"These findings together with previous reports that PI3K and Akt signaling upregulates cyclin D2 via repression of the transcriptional factor FOXO1 [XREF_BIBR] suggest that USP22 promotes cell cycle progression possibly via PI3K/Akt/cyclin D2 pathway in ATC cells."

reach
"In addition, the mechanism of USP22 on promoting cell cycle progression in colorectal cancer has been reported recently, which further confirmed our results[37]."

reach
"Mechanistically, there is evidence that USP22 promotes cell cycle progression by increasing β-catenin nuclear localization, which is required for Wnt pathway activation."

reach
"USP22 has been shown to induce cell cycle progression via BMI-1 and p-Akt pathways in colorectal carcinoma [37] and promote the epithelial-mesenchymal transition (EMT) by regulating EMT marker express[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Studies have shown that overexpression of USP22 can enhance the inhibitory effect of cell cycle inhibitors such as p21 and enhance the proliferation of tumor cells, thus promoting the occurrence and development of tumors [XREF_BIBR]."

reach
"Our previous studies have confirmed that defects in USP22 can not only cause cell cycle arrest in G0/G1 phase, but also inhibit apoptosis and promote tumor cell proliferation [XREF_BIBR]."

reach
"However, the expression and functions of USP22 in pancreatic ductal adenocarcinoma (PDA) and whether FoxM1 is involved in USP22 mediated cell cycle regulation have not been studied."

reach
"While it is clear that USP22 is overexpressed in various cancer types and may promote oncogenesis by altering gene expression, cell death and cell cycle progression, emerging evidence suggests that USP22 also harbors tumor suppressor like properties."

reach
"As a consequence, gain of USP22 functions promotes cell cycle progression and inhibits cell apoptosis, leading to cancer cell hyper-proliferation and tumorigenesis."

reach
"USP22 depletion causes cells to accumulate in the G1 phase of the cell cycle and results in concomitant decreases in the percentage of cells in the synthesis and second gap and mitosis phases."
USP22 inhibits cell cycle.
| 15
| 15

reach
"XREF_BIBR - XREF_BIBR Zhang and colleagues have demonstrated that ectopic overexpression of USP22 promotes cell proliferation and that suppression of USP22 expression by small hairpin RNA induces cell cycle arrest in human lung cancer cells."

reach
"The USP22 silencing both in CNE-1 and CNE-2 cells caused them to accumulate in the G0/G1 phase of the cell cycle."

reach
"USP22 Silencing Inhibits Proliferation and Induces Apoptosis and Cell Cycle Arrest in NSCLC Cells."

reach
"Also, USP22 silencing promotes apoptosis and cell cycle arrest in human brain gliomas."

reach
"On the other hand, USP22 suppresses the pro apoptotic and cell cycle arrest activity of p53 through SIRT1 [XREF_BIBR]."

reach
"USP22, Aurora-B and Survivin are highly expressed in OSCC, and serve an important role in the tumorigenesis of oral cancer most likely by disrupting cell cycle progression (23,26,27)."

reach
"Knockdown of USP22 was found to suppress cell proliferation in vitro and tumour growth in vivo by inducing G1 phase cell cycle arrest through synergy with TGF-beta1 (Ji et al., 2015)."

reach
"However, Ling et al reported that knockdown of USP22 by siRNA induced cells G0/G1 cell cycle arrest via the c-Myc and cyclin D2 pathway in HepG2 cells."

reach
"Our group first ensured that knockdown of USP22 can induce cell cycle arrest and inhibit cell growth in the HCC cell line HepG2 (Ling etal., 2012a)."

reach
"RNA interference mediated USP22 gene silencing promotes human brain glioma apoptosis and induces cell cycle arrest."

reach
"These results indicate that USP22 increases CRC cell migration and invasion abilities by promoting EMT."

reach
"Subsequent experiments showed that USP22 knockdown resulting from up-regulation of miR-30e-5p could inhibit proliferation, invasion, migration, and EMT in 5-8F cells."

reach
"As expected, silencing of USP22 has been proved to reverse the EMT phenotype of DDP-resistant TNBC cells in the present study."

reach
"Moreover, USP22 promotes epithelial–mesenchymal transition (EMT) in a variety of cancers by upregulating TGF-β1 expression [ 70 ], activating protein-activating enhancer binding protein 4 (AP4) [ 71 ][MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In the meantime, USP22 overexpression aggravated the EMT phenotype of TNBC cells, which revealed that USP22 contributes to the EMT process in TNBC.The causative role of glycolysis in stemness and EMT [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"These findings suggested that USP22 promotes melanoma metastasis both in vitro and in vivo.2.3 Elevated USP22 induces EMT activation in melanoma."

reach
"These results demonstrated that USP22 might promotes melanoma metastasis by inducing EMT activation.2.4 USP22 potentiates melanoma metastasis and EMT through activating PI3K/Akt/mTOR pathway."

reach
"USP22 silencing inhibits the proliferation, invasion, and EMT of OS cells in vitro ."

reach
"In the context of EMT induction, miR-30e is an important EMT inhibitor that works by targeting ITGB1, TUSC3, USP22, and SOX9 mRNAs, which reportedly induce EMT by activating transcription factors including SNAIL, SLUG, TWIST, and ZEB1/2 [XREF_BIBR, XREF_BIBR, XREF_BIBR, XREF_BIBR]."

reach
"The present study has revealled an additional novel mechanism by which USP22 induces EMT."

reach
"Moreover, USP22 overexpression can promote EMT and TGF-beta expression, whereas depletion of USP22 can reverse EMT and reduce metastasis of lung adenocarcinomas."

reach
"Moreover, USP22 downregulation repressed cell proliferation, invasion and epithelial-mesenchymal transition (EMT) via inactivation of PI3K/Akt signaling pathway in osteosarcoma cells (28)."

reach
"In this study, we found that quercetin significantly inhibited the expression of USP22 and Snail1 in high glucose (HG)-induced renal tubular epithelial cells (TECs), and reversed the expression of EMT-related proteins and inhibited the overproduction of fibronectin (FN) and Collage Type IV (Collagen IV) induced by high glucose."

reach
"Downregulation of USP22 inhibited OS cell proliferation, invasion, and EMT in vitro."

reach
"Ablation of USP22 expression remarkably attenuates melanoma migration, invasion, and epithelial-mesenchymal transition in vitro and suppresses melanoma metastasis in vivo."

reach
"The results suggested that USP22 downregulation obviously suppressed the EMT process in OS cells."
CDK1 affects USP22
2 1 | 12 14 11
CDK1 phosphorylates USP22.
1 | 10 13 11
CDK1 phosphorylates USP22. 10 / 27
1 | 8 7 10

sparser
"Phosphorylation of USP22 by CDK1 enhances its activity in deubiquitinating CCNB1."

sparser
"CDK1 phosphorylates USP22 to optimize its activity in CCNB1 stabilization during the G2/M phase."

sparser
"From our proteomic analysis, we noticed that CCNB1 appears to form a complex with both USP22 and CDK1; this prompted us to ask whether CDK1 phosphorylates USP22."

sparser
"We then analyzed whether USP22 is phosphorylated by CDK1."

sparser
"In an in vitro kinase assay, co-incubation of the purified GST-USP22 fusion proteins and the constitutive forms of CDK1 (CDK1/AF) immunoprecipitated from transiently transfected HCT116 cells confirmed USP22 phosphorylation by CDK1 ( xref )."

reach
"Phosphorylation of USP22 by CDK1 enhances its activity in deubiquitinating CCNB1."

reach
"CDK1 phosphorylates USP22 to optimize its activity in CCNB1 stabilization during the G2/M phase."

reach
"From our proteomic analysis, we noticed that CCNB1 appears to form a complex with both USP22 and CDK1; this prompted us to ask whether CDK1 phosphorylates USP22."

reach
"We then analyzed whether USP22 is phosphorylated by CDK1."

reach
"In addition, expression of the constitutively active form of CDK1 (CDK1/AF) [XREF_BIBR, XREF_BIBR] further enhanced USP22 phosphorylation in HCT116 cells (XREF_FIG)."
CDK1 phosphorylates USP22 on S237. 4 / 5
| 1 3 1

sparser
"Phosphorylation of USP22 at T147 and S237 by CDK1 increases the deubiquitination status of cyclin B1 in a cell cycle dependent manner."

reach
"In addition, phosphorylation of Thr147 and Ser237 of USP22 by CDK1 enhances the deubiquitination activity of USP22 on cyclin B1."

sparser
"In addition, phosphorylation of Thr147 and Ser237 of USP22 by CDK1 enhances the deubiquitination activity of USP22 on cyclin B1."

sparser
"Phosphorylation of USP22 at T147 and S237 by cyclin-dependent kinase 1 (CDK1) was shown to activate USP22 to deubiquitylate cyclin B1."
CDK1 phosphorylates USP22 on T147. 4 / 4
| 1 3

sparser
"Phosphorylation of USP22 at T147 and S237 by CDK1 increases the deubiquitination status of cyclin B1 in a cell cycle dependent manner."

sparser
"Phosphorylation of USP22 at T147 and S237 by cyclin-dependent kinase 1 (CDK1) was shown to activate USP22 to deubiquitylate cyclin B1."

sparser
"In addition, phosphorylation of Thr147 and Ser237 of USP22 by CDK1 enhances the deubiquitination activity of USP22 on cyclin B1."

reach
"In addition, phosphorylation of Thr147 and Ser237 of USP22 by CDK1 enhances the deubiquitination activity of USP22 on cyclin B1."
CDK1 activates USP22.
1 | 2
CDK1 activates USP22. 2 / 4
1 | 2

reach
"On the other hand, CDK1 enhances USP22 activity to stabilize CCNB1 during the G2/M phase."

reach
"USP22 is activated by CDK1 phosphorylation and deubiquitinates and stabilizes Cyclin B1 to promote cell cycle progression [XREF_BIBR]."
CDK1 binds USP22.
1 | 1
1 | 1

sparser
"From our proteomic analysis, we noticed that CCNB1 appears to form a complex with both USP22 and CDK1; this prompted us to ask whether CDK1 phosphorylates USP22."
BMI1 affects USP22
3 | 8 30
BMI1 binds USP22.
3 | 5 30
3 | 5 30

sparser
"Meanwhile, USP22 and BMI1 co-activation may be associated with GC progression and poor prognosis [ xref ]."

sparser
"USP22 and BMI1 were positively associated with TNM staging (Figure xref )."

sparser
"Overexpression of USP22 and BMI1 was previously associated with GC progression and therapy failure through clinical specimen analysis, and our study is consistent with these results."

sparser
"In this study, we hypothesized RNF220 promoted the stemness and progression of colon cancer cells via the USP22-BMI1 axis, with the aim to investigate the specific role and related mechanism of RNF220 in colon cancer."

sparser
"USP22 and BMI1 together form a multiprotein complex acting on their target homologous gene clusters."

sparser
"USP22-BMI1 silences the Hox gene and increases tumor resistance."

sparser
"In this study, we found that RNF220 promoted the stemness and progression of colon cancer cells via the USP22-BMI1 axis."

sparser
"These results indicate that USP22 interacts with BMI1 and protects BMI1 from degradation through a deubiquitination mechanism in glioma cells."

sparser
"Furthermore, our in vivo model suggested that RNF220 promoted tumor growth by regulating the USP22-BMI1 axis."

reach
"USP22 interacts with and deubiquitinates BMI1 for post-translational stabilization."
BMI1 activates USP22.
| 3
BMI1 activates USP22. 3 / 3
| 3

reach
"These results showed that BMI1 mediates the effect of USP22 on glioma stemness."

reach
"Therefore, Bmi-1 and Cyclin D2 are important oncogenes in cancer and may mediate the promoting role of USP22 in human colon cancer.In conclusion, the present study systematically examined the oncogenic role of USP22 in human colon cancer."

reach
"Considering BMI1 is a potential target of USP22 in other cancers,21, 22 we attempted to examine whether BMI1 may mediate the effect of USP22 on glioma stemness."
USP22 affects KDM1A
5 1 | 24 5
USP22 binds KDM1A.
5 | 5 5
5 | 5 5

sparser
"Moreover, GSK3β knockdown in GSC11 cells attenuated the interaction between endogenous USP22 and KDM1A ( xref ), which was confirmed in 293T cells by transfection of GSK3β-KD as compared with GSK3β-CA ( xref )."

sparser
"At the same time, the loss of OTUD7B promotes the binding of LSD1 to P62, a polyubiquitin chain binding protein, leading to LSD1 proteasomal degradation. xref USP28 (ubiquitin-specific protease 28) catalyzes deubiquitination of LSD1 through the N-terminal region of USP28 and the AOL domain of LSD1 to stabilize LSD1 protein, which enables direct regulation of the expression of differentiation genes, indirectly regulating the expression of the pluripotency activators. xref In glioma, a lack of methionine inhibits the growth of cancer cells and enhances the binding of LSD1 to E3 ubiquitin ligase CBL (casitas B-lineage lymphoma), and increased ubiquitination of LSD1 enhances expression of CXCL8 (CXC motif chemokine ligand 8) and methylated histone H3, which in turn affects the metabolism of the glycerophospholipid. xref USP7 (ubiquitin-specific protease 7) is able to interact with LSD1, exercising deubiquitination functions and enhancing the inhibitory effect of LSD1 on p53 (tumor protein p53). xref , xref , xref Phosphorylation of LSD1 catalyzed by GSK3β promotes the binding of LSD1 to USP22 (ubiquitin specific protease 22), and deubiquitination of LSD1 promotes tumor formation. xref , xref USP38 (ubiquitin-specific protease 38) interacts with LSD1 at amino acid 454 to stabilize protein level."

reach
"Using a series of KDM1A deletion mutants, we found that KDM1A 's AOL domain was required for KDM1A 's binding with USP22 (XREF_FIG)."

reach
"Moreover, GSK3beta knockdown in GSC11 cells attenuated the interaction between endogenous USP22 and KDM1A (XREF_FIG), which was confirmed in 293T cells by transfection of GSK3beta-KD as compared with GSK3beta-CA (XREF_SUPPLEMENTARY)."

reach
"Furthermore, KDM1A S683A significantly decreased the interaction between USP22 and KDM1A in the presence of GSK3beta-CA (XREF_FIG)."

sparser
"The target relationship of USP22 and miR-362-3p as well as the interaction of USP22 and LSD1 in RB was verified."

reach
"The target relationship of USP22 and miR-362-3p as well as the interaction of USP22 and LSD1 in RB was verified."

sparser
"Likewise, glycogen synthase kinase-3β (GSK-3β) phosphorylates LSD1 at S683 and stabilizes LSD1 via increasing the binding of ubiquitin specific peptidase 22 with LSD1 [ 36 ]."

reach
"40 In glioma, a lack of methionine inhibits the growth of cancer cells and enhances the binding of LSD1 to E3 ubiquitin ligase CBL (casitas B-lineage lymphoma), and increased ubiquitination of LSD1 enhances expression of CXCL8 (CXC motif chemokine ligand 8) and methylated histone H3, which in turn affects the metabolism of the glycerophospholipid.41 USP7 (ubiquitin-specific protease 7) is able to interact with LSD1, exercising deubiquitination functions and enhancing the inhibitory effect of LSD1 on p53 (tumor protein p53).42, 43, 44 Phosphorylation of LSD1 catalyzed by GSK3β promotes the binding of LSD1 to USP22 (ubiquitin specific protease 22), and deubiquitination of LSD1 promotes tumor formation.29 45 USP38 (ubiquitin-specific protease 38) interacts with LSD1 at amino acid 454 to stabilize protein level."

sparser
"Using a series of KDM1A deletion mutants, we found that KDM1A’s AOL domain was required for KDM1A’s binding with USP22 ( xref )."
USP22 activates KDM1A.
| 10
USP22 activates KDM1A. 10 / 12
| 10

reach
"USP22 could also mediate the stability of LSD1 via deubiquitination to promote cancer progression [ 46 , 47 ]."

reach
"These findings demonstrate that nuclear GSK3β- and USP22-mediated LSD1 stabilization promotes GSC stemness and glioblastoma tumorigenesis.More recently, DUB-mediated LSD1 stabilization has been suggested to contribute to breast cancer metastasis."

reach
"Our findings demonstrate that nuclear GSK3beta and USP22 mediated KDM1A stabilization is essential for glioblastoma tumorigenesis."

reach
"miR-140 inhibits osteosarcoma progression by impairing USP22 mediated LSD1 stabilization and promoting p21 expression."

reach
"In contrast, USP22 overexpression in 293T cells increased KDM1A stability (XREF_SUPPLEMENTARY)."

reach
"MiR-140 suppresses osteosarcoma progression by inhibiting USP22-mediated LSD1 stability, resulting in promoting ubiquitination of LSD1 and increasing p21 expression (105)."

reach
"Likewise, knockdown of USP22 inhibited GSC stemness (XREF_FIG and XREF_SUPPLEMENTARY), and exogenous KDM1A rescued the effect of USP22 depletion on GSC stemness (XREF_FIG and XREF_SUPPLEMENTARY)."

reach
"USP22 and GSK3β together in the nucleus mediate the stability of lysine (K)-specific demethylase 1A (KDM1A), a histone demethylase, and changes in histone H3K4 methylation level inhibit the transcript[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Furthermore, microRNA 140 inhibits the USP22 mediated LSD1 stabilization, which halts the cell cycle (78)."

reach
"For instance, deubiquitinating enzyme USP10 promotes OS metastasis by stabilizing YAP1 [33], miR-140 inhibits OS progression by impairing USP22-mediated LSD1 stabilization [34]."
USP22 deubiquitinates KDM1A.
1 | 7
USP22 deubiquitinates KDM1A. 7 / 8
1 | 7

reach
"A screen for DUBs that could stabilize KDM1 demonstrates USP15, USP21, USP22 and USP28 as potential hits, among which ectopic expression of USP22 reduced KDM1A ubiquitination levels strikingly."

reach
"In addition, deubiquitination of KDM1A by USP22 was attenuated after GSK3beta knockdown (XREF_FIG)."

reach
"USP22 deubiquitinated LSD1 in RB."

reach
"However, of those four DUBs, only USP22 substantially decreased KDM1A ubiquitination (XREF_SUPPLEMENTARY)."

reach
"Moreover, using in vitro deubiquitination assay, we found that KDM1A ubiquitination was decreased by incubating with recombinant USP22, suggesting that USP22 deubiquitinates KDM1A directly (XREF_FIG)."

reach
"Furthermore, knockdown of USP22 in GSC11 cells increased KDM1A ubiquitination (XREF_FIG)."

reach
"These results indicate that USP22 deubiquitinates and stabilizes KDM1A."
USP22 increases the amount of KDM1A.
| 2
USP22 increases the amount of KDM1A. 2 / 2
| 2

reach
"In GSC11 cells, USP22 knockdown decreased KDM1A protein level (XREF_FIG), but not KDM1A mRNA (XREF_SUPPLEMENTARY)."

reach
"We therefore screened a panel of DUBs in which 23 DUBs ' cDNA plasmids were transfected into 293T cells, and found that USP15, USP21, USP22, and USP28 upregulated KDM1A levels (XREF_SUPPLEMENTARY)."
USP22 affects cell growth
| 1 30 1
USP22 activates cell growth.
| 1 24
| 1 24

reach
"Indeed, compared with control, we found that the USP22 knockdown cells grew significantly more slowly in H1650 (p < 0.05, Fig. 2 B) and the USP22 overexpression in A549 accelerated cell growth (p < 0.[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"However, targeted deletion of USP22 completely abolished EPI-induced 4T1 cell growth and migration (fig."

reach
"Additionally, it was observed that cell growth was suppressed by USP22 siRNA in OSCC cells."

reach
"In addition, we present evidence that USP22 promotes Bel/Fu cell growth, migration, invasion, EMT and chemoresistance."

reach
"Knockdown of USP22 significantly retarded cell growth in H1975 cells compared to control (p < 0.05, Fig. 2 A), whereas overexpression of USP22 in PC9 cells accelerated cell growth (p < 0.01, Fig. 2 A)[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Collectively, our results show that USP22 silencing in HepG2 cells suppressed cell growth through mitochondrial apoptosis and that this suppression was dependent on caspase activation."

reach
"These observations indicate that USP22 gene silencing by RNA interference inhibits HCC cell growth."

reach
"Our data showed that USP22 silencing led to significantly slower cell growth compared with the control (p < 0.01 after 120 h) (XREF_FIG B)."

reach
"Moreover, it has been shown that USP22 promotes cell growth by regulating the far upstream element (FUSE)-binding protein 1 (FBP1), a transcriptional regulator of p21 [XREF_BIBR]."

reach
"Our results demonstrated that USP22 silencing suppressed cell growth and induced cell apoptosis."
USP22 inhibits cell growth.
| 6 1
| 6 1

reach
"USP22 siRNA suppressed cell growth."

sparser
"These observations indicate that USP22 gene silencing by RNA interference inhibits HCC cell growth (Fig. xref )."

reach
"Furthermore, ALKBH5-mediated m6A deficiency increases the expression of USP22 and RNF40 in osteosarcomas, promoting osteosarcoma cell growth and proliferation [40]."

reach
"These results suggested that USP22 siRNA effectively inhibited OSCC cell growth in vitro."

reach
"It was identified that USP22 siRNA decreased the expression of USP22 protein (Fig. 2A) and also inhibited cell growth in OSCC cells (Fig. 2B)."

reach
"Consistently, overexpression of USP22 stimulates while downregulation of USP22 suppresses breast cancer cell growth, migration and tumorigenesis in a MYC dependent manner.113 USP13."

reach
"Hence, we determined whether downregulation of USP22 suppresses colon cancer cell growth and cancer progression."
MYC affects USP22
10 | 13 13
MYC binds USP22.
10 | 2 13
10 | 2 13

sparser
"Reciprocal co-IP assays in 293T cells demonstrated that ectopically expressed FLAG-BRD4 interacted with ectopically expressed Myc-USP22 ( Fig. 5 E and F)."

sparser
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected Myc-USP22 and Flag-FoxM1, but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone ( xref )."

sparser
"As shown in Fig. 5 G, the ubiquitination level of BRD4 was decreased when Myc-USP22 was co-transfected into 293T cells."

reach
"The data presented here suggest that the recruitment of USP22 to Myc targets as part of the hSAGA complex may provide this critical function.An outstanding question raised by these studies is the iden[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"It has been reported that SIRT1 protein level is increased in LSCs in both chronic myeloid leukemia (CML) and acute myeloid leukemia (AML), by either transcriptional induction in CML or by c-Myc-USP22 mediated stabilization of SIRT1 protein in AML [ xref ]."

sparser
"These unique cellular effects are mediated through USP22's interaction with c-Myc which enhances c-Myc deubiquitination and reduces intracellular glycolysis [ xref ]."

sparser
"Western blotting analysis detected the Flag-tagged FOXO1 in anti-Myc immunoprecipitants of human embryonic kidney (HEK) 293T cells when myc-USP22 but not empty vector was cotransfected ( xref )."

sparser
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected with Myc-USP22 and Flag-FoxM1 but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone ( xref C)."

sparser
"USP22 interacts with MYC through its N-terminal region containing zinc finger motif and abrogates Fbw7-mediated polyubiquitination and degradation of MYC."

sparser
"These results support a role for USP22 in MYC-mediated increase in SIRT1 protein stabilization, and indicate that FLT3-ITD, c-MYC and USP22 form an oncogenic network that enhances SIRT1 expression and activity in leukemic cells."
MYC activates USP22.
| 7
MYC activates USP22. 7 / 8
| 7

reach
"At the posttranscriptional level, c-MYC has been shown to increase USP22 protein but not mRNA levels."

reach
"Instead, the increased SIRT1 expression observed was linked to a myc-dependent induction of USP22 deuibiquitinase, leading to reduced SIRT1 ubiquitination and increase SIRT1 stability in leukemic cells."

reach
"For example, as part of the SAGA complex USP22 promotes transcriptional activation by the Myc oncogene (Zhang et al., 2008b)."

reach
"C-Myc induces the post-transcriptional upregulation of USP22, which stabilizes SIRT1 in LSCs [165]."

reach
"This SIRT1 overexpression is related to enhanced expression of the USP22 deubiquitinase induced by c-MYC, leading to reduced SIRT1 ubiquitination and enhanced stability."

reach
"SIRT1 overexpression is related to enhanced expression of the USP22 deubiquitinase induced by c-MYC, leading to reduced SIRT1 ubiquitination and enhanced stability, but inhibition of SIRT1 expression or activity reduced the growth of FLT3-ITD AML LSCs and significantly enhanced TKI-mediated killing of the cells."

reach
"Furthermore, USP22 acts as an enzymatic component of the SAGA transcriptional cofactor complex and is activated by Myc as an oncogene [38]."
| PMC
MYC increases the amount of USP22.
| 4
MYC increases the amount of USP22. 4 / 4
| 4

reach
"This is significant, as AR upregulation drives the CRPC phenotype, MYC is a known PCa oncogene, and USP22 regulates MYC transcription."

reach
"The proto-oncogene c-Myc further enhances this expression by inducing elevated levels of USP22, a deubiquitinase."

reach
"These results suggest that USP22 positively regulates MYC dependent transcription, which may at least partially explain its oncogenic properties."

reach
"Importantly, c-MYC has been reported to induce the expression of the deubiquitinase USP22, which in turn reduced ubiquitination and enhanced the stability of SIRT1 in CD34 + Flt3-ITD cells."
CD274 affects USP22
9 | 8 20
9 | 8 20

sparser
"USP22 directly interacts with the C-terminus of PD-L1, thereby inducing its deubiquitination and stabilization."

sparser
"USP22 physically interacted with PD-L1."

sparser
"Now that USP22 could regulate PD-L1 protein level, we asked whether USP22 interacted with PD-L1."

sparser
"We performed co-immunoprecipitation assays in HEK293FT and H157 cells respectively, and we found that USP22 interacted with PD-L1 (Fig.  xref a, Additional file xref : Fig. S2A)."

sparser
"Thus, we validated the interaction between USP22 and PD-L1."

sparser
"The loss of USP22 causes PD-L1 to be degraded at the post-translational level, which has been shown to reduce carcinogenesis and increase T cell-mediated cell death. xref PD-L1 targeted immunotherapy and CDDP-based chemotherapy were both shown to benefit from USP22 reduction in another research, highlighting the complex and important functions of the USP22-PD-L1 axis in cancer treatment. xref According to a recent research, the tumor-promoting long noncoding RNA (lncRNA) KCNQ1OT1 controls the ubiquitination of PD-L1 and decreases the response of CD8 + T cells via the miR-30a-5p/USP22 pathway."

sparser
"USP22 may be a particularly attractive target for cancer immunotherapy, because USP22 can limit the effect of anti-tumor immune response in Treg cells, and there is currently evidence that USP22 can i[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"The deubiquitinase USP22 can bind with PD-L1 and enhance its stability, leading to reduced T cell cytotoxicity in tumor cells (76)."

sparser
"In addition, USP22 could facilitated the interaction between CSN5 and PD-L1, and CSN5 promoted the interaction between USP22 and PD-L1."

sparser
"Another study also showed that USP22 interacted with PD-L1, enhancing its stability by removing ubiquitin and preventing its breakdown by the proteasome."
USP22 affects CCNB1
4 1 | 20 7
USP22 binds CCNB1.
4 | 5 7
4 | 5 7

reach
"Our studies suggest that SIRT1 and CCNB1 interact with USP22 through different regions."

sparser
"However, this report clearly demonstrates that USP22 binds and deubiquitinates cyclin B1, leading to its stabilization."

sparser
"USP22 binding to Cyclin B1, promotes cyclin B1 accumulation in the nucleus and inhibits its degradation [ xref ]."

sparser
"USP22 interacts with CCNB1."

sparser
"The interaction between endogenous USP22 and CCNB1 in human colon cancer cells was also detected, as anti-CCNB1-specific antibody but not normal mouse immunoglobulin G immunoprecipitated USP22 protein ( xref )."

sparser
"To further refine our understanding of the molecular interaction between USP22 and CCNB1, we generated truncated USP22 mutants ( xref )."

reach
"USP22 binding to Cyclin B1, promotes cyclin B1 accumulation in the nucleus and inhibits its degradation [XREF_BIBR]."

sparser
"In addition, as S237 is also within the portion of USP22 that interacts with CCNB1, it is possible that CDK1-mediated S237 phosphorylation enhances USP22 interaction with CCNB1."

reach
"USP22 interacts with CCNB1."

reach
"The interaction between endogenous USP22 and CCNB1 in human colon cancer cells was also detected, as anti-CCNB1-specific antibody but not normal mouse immunoglobulin G immunoprecipitated USP22 protein (XREF_FIG)."
USP22 activates CCNB1.
| 8
USP22 activates CCNB1. 8 / 10
| 8

reach
"Phosphorylation of USP22 at T147 and S237 by CDK1 increases the deubiquitination status of cyclin B1 in a cell cycle dependent manner."

reach
"Our discovery here, that both USP22 and CCNB1 proteins are elevated and positively associated in human colon cancers, indicates that USP22 mediated CCNB1 stabilization is one possible molecular mechanism underlying its proto-oncogenicity."

reach
"In addition, phosphorylation of Thr147 and Ser237 of USP22 by CDK1 enhances the deubiquitination activity of USP22 on cyclin B1."

reach
"USP22 mediated CCNB1 stabilization is regulated by both CDK1 and the APC/C E3 ubiquitin ligase complex."

reach
"In addition, Usp22 promotes CRC by stabilizing cyclin B1."

reach
"Moreover, the proteasome inhibitor MG132 protected CCNB1 from degradation in usp22 knockdown cells (XREF_FIG), implying that USP22 mediates CCNB1 stabilization through regulating the proteasomal pathway."

reach
"Given that degradation of CCNB1 is required for cells to exit M phase and enter anaphase [XREF_BIBR, XREF_BIBR], we proposed that USP22 degradation, which presumably allows CCNB1 degradation during late M phase, is necessary for cell cycle regulation."

reach
"Therefore, APC and CDC20 E3 ligase complex promotes USP22 protein degradation, presumably allowing CCNB1 degradation for cells to exit M phase."
USP22 deubiquitinates CCNB1.
1 | 7
USP22 deubiquitinates CCNB1. 7 / 8
1 | 7

reach
"In addition, USP22 promotes cell cycle progression and proliferation of cancer cells by regulating and inducing the deubiquitination of the cyclin B1 protein [ 67 ]."

reach
"USP22 can inhibit the ubiquitination of CCNB1 and induce the G2/M transition of CRC cells."

reach
"Indeed, USP22 inhibited CCNB1 ubiquitination both in vivo and in vitro (XREF_FIG and XREF_SUPPLEMENTARY)."

reach
"The deubiquitinase catalytic activity of USP22 is required for CCNB1 deubiquitination because the catalytically inactive USP22 (USP22 and C185A) mutant failed to suppress CCNB1 ubiquitination without affecting its interaction with CCNB1 (XREF_SUPPLEMENTARY)."

reach
"USP22 deubiquitinates cyclin B1, stabilizes cyclin B1 by antagonizing proteasome mediated degradation, and promotes itaccumulation in the nucleus (Lin etal., 2015; Melo-Cardenas etal., 2016)."

reach
"In addition to histones, USP22 deubiquitinates TRF1, CCNB1, CCND1, and SIRT1, thereby regulating involvement in metabolism, cycling, and apoptosis."

reach
"[Somewhat paradoxically, a role for USP22 in M phase has been proposed based on evidence that CCNB1 is deubiquitylated by USP22 (43)]."
ATXN7L3 affects USP22
16 | 5 13
ATXN7L3 binds USP22.
16 | 3 13
16 | 3 3

sparser
"As shown in xref , we found that association of USP22 and ATXN7L3 was greatly impaired upon addition of increasing amounts of USP27X (compare lane 7 to 8, 9 and10)."

reach
"Further, Pfam analysis of protein domain structures confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity (XREF_FIG and XREF_SUPPLEMENTARY)."

reach
"Thus, the formation of this module would be driven by interactions between ATXN7L3 and USP22 through their respective ZnF-Sgf11 and ZnF-UBP domains, and between ATXN7L3 ZnF-Sgf11 and ENY2."

reach
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Thus, the formation of this module would be driven by interactions between ATXN7L3 and USP22 through their respective ZnF-Sgf11 and ZnF-UBP domains, and between ATXN7L3 ZnF-Sgf11 and ENY2 ( Figure 2 D[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Immunoblot analysis of purified complexes revealed an interaction between ATXN7L3 and USP22 ( Figure 2 A, lane 1) and between ATXN7L3 and ENY2 ( Figure 2 A, lanes 3 and 7), but not between ENY2 and US[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 8

sparser
"The SAGA DUB module is highly conserved both in subunit composition and structural organization ( xref ; xref ; xref ; xref ; xref ; xref ), The deubiquitinating enzyme USP22 (Ubp8 in yeast) closely associates with two adapter proteins, ATXN7L3 and ENY2 (Sgf11 and Sus1 in yeast, respectively) ( xref ; xref ; xref ), and a fourth protein, ATXN7 (Sgf73), anchors the DUB module to the larger SAGA complex."

sparser
"It has been reported that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator Spt-Ada-Gcn5-acetyltransferase (SAGA) complex to regulate global levels of H2B monoubiquitination, thereby promoting antibody class switch recombination by facilitating nonhomologous end joining ( xref ; xref ; xref )."

sparser
"Similarly, USP22, the human homolog of Ubp8, is bound to the STAGA complex through interactions with ATXN7L3 and ENY2, which are homologs of Sgf11 and Sus1, respectively ( xref )."

sparser
"Further, Pfam analysis of protein domain structures ( xref ) confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity ( xref and xref ) ( xref )."

sparser
"We show that the ubiquitin protease USP22 forms a subcomplex with ATXN7L3 (ySgf11 homolog) and ENY2 (ySus1 homolog)."

sparser
"These results together suggest that ATXN7L3, USP22, and ENY2 form a stable subcomplex and that USP22 and ENY2 are recruited into TFTC/STAGA by ATXN7L3 ( Figure 2 D)."

sparser
"Cotransfection and coimmunoprecipitation experiments revealed that the ATXN7L3 ZnF-Sgf11 domain alone is able to interact with both ENY2 and USP22 as efficiently as the full-length ATXN7L3 (see Figur[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Previous study has shown that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator SAGA complex to regulate global levels of H2B monoubiquitination ( xref )."
AR binds USP22 and ATXN7L3. 2 / 2
| 2

sparser
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR-dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"To further investigate this hypothesis, we analyzed putative interactions of USP22 or ATXN7L3 with AR in mammalian cells."
ATXN7L3 activates USP22.
| 2
ATXN7L3 activates USP22. 2 / 2
| 2

reach
"However, depletion of the DUBm adaptors ENY2 and/or ATXN7L3 would eliminate the activity of all the USP22-related DUBm-s (independently of the cellular expression levels of the three related USPs)."

reach
"Together with SAGA subunit ENY2, ATXN7L3 functions to activate the SAGA deubiquitinase USP22 XREF_BIBR."
USP22 affects FOXM1
| 1 21 9
USP22 binds FOXM1.
| 5 9
| 5 9

sparser
"These results indicate that FoxM1 physically interacts with USP22 in breast cancer cells."

sparser
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected with Myc-USP22 and Flag-FoxM1 but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone ( xref C)."

sparser
"Our data indicate physical interaction between USP22 and FoxM1 is required for USP22-mediated suppression of FoxM1 ubiquitination, because mutation of cystines 61 and 63, which disrupts the zinc finger structure and its interaction with FoxM1 ( xref ), totally abolished USP22 activity in suppressing FoxM1 ubiquitination ( xref )."

sparser
"The endogenous interaction between USP22 and FoxM1 was further validated in patient-derived breast cancer L2G + TN1 and 4T1 cells ( xref D and xref B)."

reach
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected Myc-USP22 and Flag-FoxM1, but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone (Fig. 3C)."

reach
"The endogenous interaction between USP22 and FoxM1 in patient-derived breast cancer L2G TN1 cells was further validated (Fig. 3D and s3B)."

sparser
"Truncated mutation analysis revealed that the zinc finger-containing N-terminus is sufficient for USP22 interaction with FoxM1, while the C-terminus ubiquitin-specific peptidase domain is not involved in mediating its FoxM1 interaction ( xref F)."

reach
"Our data indicate physical interaction between USP22 and FoxM1 is required for USP22-mediated suppression of FoxM1 ubiquitination, because mutation of cystines 61 and 63, which disrupts the zinc finger structure and its interaction with FoxM1 (Fig. 3I), totally abolished USP22 activity in suppressing FoxM1 ubiquitination (Fig. 3G)."

sparser
"These results indicate that FoxM1 physically interacts with USP22 in breast cancer cells."

reach
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected with Myc-USP22 and Flag-FoxM1 but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone (Figure 3C)."
USP22 increases the amount of FOXM1.
| 8
USP22 increases the amount of FOXM1. 8 / 9
| 8

reach
"USP22 promotes the G1/S phase transition by upregulating FoxM1 expression via beta-catenin nuclear localization and is associated with poor prognosis in stage II pancreatic ductal adenocarcinoma."

reach
"USP22 promotes the G1/S phase transition by upregulating FoxM1 expression via promoting beta-catenin nuclear localization."

reach
"Similarly, in pancreatic ductal adenocarcinoma cells, USP22 promotes the G1/S transition and proliferation by upregulating the expression of FoxM1 [8], a key regulator of the G1/S and G2/M cell cycle transitions [32]."

reach
"USP22 promotes the G1/S phase transition by upregulating FoxM1 expression via beta-catenin nuclear localization and is associated with poor prognosis in stage II pancreatic ductal adenocarcinoma."

reach
"By contrast, treatment with NH Cl, an inhibitor of endosome-lysosome degradation pathway, fails to protect FoxM1 from degradation (Figure s3A), suggesting that USP22 promotes FoxM1 level through inhibiting its proteasomal degradation.As a deubiquitinase, USP22 exerts its biological function largely through protecting its downstream substrates from ubiquitination-mediated degradation ."

reach
"By contrast, treatment with NH Cl, an inhibitor of endosome-lysosome degradation pathway, fails to protect FoxM1 from degradation (Figure S3A), suggesting that USP22 promotes FoxM1 expression by inhibiting its proteasomal degradation.As a deubiquitinase, USP22 exerts its biological function largely through protecting its downstream substrates from ubiquitination-mediated degradation.27 Accordingly, we speculated that USP22 could be a deubiquitinase of FoxM1."

reach
"USP22 can increase the expression of foxhead box M1(FoxM1) by inducing β-catenin nuclear location which promotes the expression of FoxM1.Besides, in a study about USP5 in PC, it promoted tumorigenesis[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Mechanistically, it has been reported that USP22 induces β-catenin nuclear localisation and upregulates FoxM1 expression to promote G1/S cell cycle transition and cell proliferation (Ning et al., 2014)."
USP22 deubiquitinates FOXM1.
| 5
USP22 deubiquitinates FOXM1. 5 / 5
| 5

reach
"Importantly, transient USP22 expression largely diminished FoxM1 ubiquitination (Fig. 3G)."

reach
"Our data indicate physical interaction between USP22 and FoxM1 is required for USP22-mediated suppression of FoxM1 ubiquitination, because mutation of cystines 61 and 63, which disrupts the zinc finger structure and its interaction with FoxM1 (Fig. 3I), totally abolished USP22 activity in suppressing FoxM1 ubiquitination (Fig. 3G)."

reach
"As expected, expression of the catalytically inactive USP22, through C185A mutation, failed to inhibit FoxM1 ubiquitination despite not altering its interaction with FoxM1 (Figures 3G and 3M)."

reach
"USP22 is required for human endometrial stromal cell proliferation and decidualization by deubiquitinating FoxM1."

reach
"Higher molecular weight bands were detected in FoxM1 immunoprecipitants, indicating FoxM1 is ubiquitinated possibly by its endogenous E3 ubiquitin ligases such as FBWX7.29 Importantly, transient USP22 expression largely diminished FoxM1 ubiquitination (Figure 3M)."
USP22 activates FOXM1.
| 1 3
USP22 activates FOXM1. 4 / 4
| 1 3

reach
"The result of GSIS assay further indicated that overexpressing USP22 could rescue the insulin secretion ability of INS-1 cells that was impaired by HG; but miR-144-3p mimic abated this impact (Fig. 4H)."

eidos
"Mechanistically , it has been reported that USP22 induces beta-catenin nuclear localisation and upregulates FoxM1 expression to promote G1 / S cell cycle transition and cell proliferation ( Ning et al ., 2014 ) ."

reach
"In concordance with this conclusion, USP22 overexpression dramatically prolonged FoxM1 half-life as measured by pulse-chase analysis (Fig. 3J & 3K)."

reach
"In pancreatic cancer cell lines a different mechanism in USP22 was found to modulate the lacatenin/Wnt signaling and therefore increase the abundance of FoxM1, a transcription factor that normally represses the expression of p21 and p27."
USP22 affects ATXN7L3
16 | 3 13
16 | 3 3

sparser
"As shown in xref , we found that association of USP22 and ATXN7L3 was greatly impaired upon addition of increasing amounts of USP27X (compare lane 7 to 8, 9 and10)."

reach
"Further, Pfam analysis of protein domain structures confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity (XREF_FIG and XREF_SUPPLEMENTARY)."

reach
"Thus, the formation of this module would be driven by interactions between ATXN7L3 and USP22 through their respective ZnF-Sgf11 and ZnF-UBP domains, and between ATXN7L3 ZnF-Sgf11 and ENY2."

reach
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Thus, the formation of this module would be driven by interactions between ATXN7L3 and USP22 through their respective ZnF-Sgf11 and ZnF-UBP domains, and between ATXN7L3 ZnF-Sgf11 and ENY2 ( Figure 2 D[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Immunoblot analysis of purified complexes revealed an interaction between ATXN7L3 and USP22 ( Figure 2 A, lane 1) and between ATXN7L3 and ENY2 ( Figure 2 A, lanes 3 and 7), but not between ENY2 and US[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 8

sparser
"The SAGA DUB module is highly conserved both in subunit composition and structural organization ( xref ; xref ; xref ; xref ; xref ; xref ), The deubiquitinating enzyme USP22 (Ubp8 in yeast) closely associates with two adapter proteins, ATXN7L3 and ENY2 (Sgf11 and Sus1 in yeast, respectively) ( xref ; xref ; xref ), and a fourth protein, ATXN7 (Sgf73), anchors the DUB module to the larger SAGA complex."

sparser
"It has been reported that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator Spt-Ada-Gcn5-acetyltransferase (SAGA) complex to regulate global levels of H2B monoubiquitination, thereby promoting antibody class switch recombination by facilitating nonhomologous end joining ( xref ; xref ; xref )."

sparser
"Similarly, USP22, the human homolog of Ubp8, is bound to the STAGA complex through interactions with ATXN7L3 and ENY2, which are homologs of Sgf11 and Sus1, respectively ( xref )."

sparser
"Further, Pfam analysis of protein domain structures ( xref ) confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity ( xref and xref ) ( xref )."

sparser
"We show that the ubiquitin protease USP22 forms a subcomplex with ATXN7L3 (ySgf11 homolog) and ENY2 (ySus1 homolog)."

sparser
"These results together suggest that ATXN7L3, USP22, and ENY2 form a stable subcomplex and that USP22 and ENY2 are recruited into TFTC/STAGA by ATXN7L3 ( Figure 2 D)."

sparser
"Cotransfection and coimmunoprecipitation experiments revealed that the ATXN7L3 ZnF-Sgf11 domain alone is able to interact with both ENY2 and USP22 as efficiently as the full-length ATXN7L3 (see Figur[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Previous study has shown that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator SAGA complex to regulate global levels of H2B monoubiquitination ( xref )."
AR binds USP22 and ATXN7L3. 2 / 2
| 2

sparser
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR-dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"To further investigate this hypothesis, we analyzed putative interactions of USP22 or ATXN7L3 with AR in mammalian cells."
KAT2A affects USP22
14 | 8 8
KAT2A binds USP22.
14 | 5 8
14 | 5 8

sparser
"Gcn5 is closely associated with USP22 or its fly orthologue, Nonstop ( xref ; xref ; xref ; xref )."

reach
"This raises the hypothesis that once USP22 associates with SAGA and GCN5, it may oppose telomere recombination in APBs by promoting the binding of POT1 to telomeres via TRF1, thereby inhibiting ATR activation."

reach
"GCN5 binds to ubiquitin-specific peptidase 22 (USP22) which deubiquitinates and thus salvages the telomeric repeat-binding factor 1 (TRF1) shelterin protein."

sparser
"There were 408 overlapping putative target genes bound by both USP22 and GCN5."

sparser
"Motif analysis showed that the sequences bound by USP22 and GCN5 shared two common motifs."

sparser
"Of these genes, 408 genes were bound potentially by both USP22 and GCN5, which accounted of 16.8% of genes recognized by USP22 and 18.1% recognized by GCN5 ( xref )."

reach
"The polypeptide composition of these purified complexes was assessed by silver staining, which revealed a pattern of 10-12 comigrating proteins shared by the USP22 and hGCN5 complexes."

reach
"Normalized ChIP‐seq signals for the individual SAGA subunits demonstrated inducible binding of GCN5, USP22, and ATXN7L3 after treatment (Fig 1C)."

sparser
"We found either USP22 antibody or GCN5 antibody precipitated a large number of DNA fragments, indicating that either USP22 or GCN5 widely interacted with different genes in the genome of HeLa cells."

sparser
"MACS analysis indicated that there were many genes potentially bound by USP22 or GCN5."
KAT2A inhibits USP22.
| 1
KAT2A inhibits USP22. 1 / 2
| 1

reach
"Loss of Gcn5 leads to depletion of Usp22 and the DUB module from SAGA, compromising Usp22 activity, leading to telomere fusions [XREF_BIBR]."
KAT2A activates USP22.
| 2
KAT2A activates USP22. 2 / 2
| 2

reach
"GCN5 has been shown to support the association of USP22 with the SAGA complex."

reach
"Loss of Gcn5 impairs the deubiquitinating activity of Usp22 [XREF_BIBR], which partners with Atxn7 in the DUB module."
USP22 affects Histone_H2B
| 29 1
USP22 deubiquitinates Histone_H2B.
| 27
USP22 deubiquitinates Histone_H2B. 10 / 27
| 27

reach
"In addition, reduced expression of USP22 is associated with chromosomal instability (CIN), since efficient chromosome compaction during mitosis requires USP22 deubiquitination of H2B in metaphase (Jeusset et al., 2021)."

reach
"Interestingly, USP22 alone deubiquitinates very inefficiently histone H2B, and its activation requires association with TFTC/STAGA."

reach
"USP22 might de-ubiquitinate H2A and H2B, subunits of the human SAGA complex that are intimately linked to the transcriptional activation of the MYC gene and increased cell proliferation in HCC [XREF_BIBR]."

reach
"Apart from deubiquitinating histones H2A and H2B, USP22 also modulates transcription through stabilization of the p53-antagonizing deacetylase sirtuin-1 (SIRT1) to suppress p53-controlled transcription and to abrogate pre-mature senescence in pluripotent progenitor cells during embryonic development [5]."

reach
"As we discussed previously, USP22 is a component of a transcriptional activator complex SAGA and can deubiquitinate histones H2A and H2B, as well as several other substrates (Zhang et al., 2008a, b)."

reach
"Our observations regarding differentiation could not be correlated to H2B and H2Bub1 levels suggesting that H2B is only partially deubiquitinated by Usp22."

reach
"USP22 deubiquitylates both, H2A and H2B."

reach
"In yeast, the USP22 ortholog deubiquitylates H2B, resulting in Pol II Ser2 phosphorylation and subsequent transcriptional elongation."

reach
"USP22 is able to deubiquitinate histone H2A and H2B in vitro and is required for androgen receptor transcription activation."

reach
"However, while USP22 is capable of deubiquitinating H2B in vitro [36,53,60] and USP22 knockdown increases H2Bub1 levels on the IRF1 gene [18] , none of the published reports demonstrated a significant[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 ubiquitinates Histone_H2B.
| 2 1
USP22 ubiquitinates Histone_H2B. 3 / 3
| 2 1

reach
"The modulation of histone H2B ubiquitination by USP22 was shown to underlie the processing of JAK-STAT-inducible genes, suggesting a potential mechanism of USP22 in cancer [28]."

sparser
"The modulation of histone H2B ubiquitination by USP22 was shown to underlie the processing of JAK-STAT-inducible genes, suggesting a potential mechanism of USP22 in cancer ."

reach
"Reduced USP22 Expression Impairs Mitotic Removal of H2B Monoubiquitination, Alters Chromatin Compaction and Induces Chromosome Instability That May Promote Oncogenesis."
USP22 affects AKT
| 25 4
USP22 activates AKT.
| 17
USP22 activates AKT. 10 / 17
| 17

reach
"USP22 overexpression in gastric cancer cells induces the upregulation of SOS1 and activation of the RAS/ERK and PI3K/AKT pathways."

reach
"3.5 USP22 activates the AKT and MRP1 pathway depending on SIRT1 in HCC cells."

reach
"Western blot analysis showed that USP22 overexpression also induced activation of the RAS/ERK and PI3K/AKT pathways in SGC7901 cells and xenograft tumor tissues."

reach
"Studies have demonstrated that USP22 directly interacts with SIRT1, activating the AKT/GSK-3β/multidrug resistance-associated protein 1 (MRP1) pathway, thereby enhancing 5-Fu efflux and reducing 5-Fu-induced apoptosis in HCC cells [124]."

reach
"XREF_FIG A, knockdown of USP22 by shRNA inhibited the AKT and MRP1 pathway compared with control shRNA cells and wild-type cells."

reach
"We found that USP22 promotes multidrug resistance in HCC cells by activating SIRT1/protein kinase B (Akt)/multidrug resistance-associated protein 1 (MRP1) pathway, while inhibition of USP22 and SIRT1 [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Similarly, overexpression of USP22 in BEL/7402 cells activated the AKT and MRP1 pathway."

reach
"Previous study also showed that USP22 promotes cell cycle progression by positively regulating the PI3K and Akt pathway [XREF_BIBR]."

reach
"Moreover, SIRT1 deficiency attenuated USP22 induced activation of the AKT and MRP1 pathway in BEL7402 cells, suggesting that USP22 regulated the AKT and MRP1 pathway in a SIRT1 dependent manner."

reach
"For example, USP22 can stabilize the E2F6 stability and activate Akt pathway in hepatocellular carcinoma (HCC), leading to aggressive progression of HCC (25)."
USP22 inhibits AKT.
| 5
USP22 inhibits AKT. 5 / 6
| 5

reach
"Smad silencing can block the downregulation of Akt induced by USP22 knockdown and reverse 5-FU resistance [ 77 ]."

reach
"Furthermore, our results showed that USP22 deletion also caused down-regulation of Akt and GSK3beta activity, which can also contribute to the reduction of cyclin D2."

reach
"Silencing Smad4 blocked USP22 knockdown induced Akt inhibition in Bel/Fu cells."

reach
"USP22 prevents ubiquitination-mediated EGFR degradation, thereby inducing persistent activation of EGFR-mediated oncogenic signaling pathways, such as STAT3, AKT/mTOR, and MEK/ERK pathways."

reach
"Downregulation of USP22 Inhibited the Activation of PI3K and Akt Signaling Pathway."
USP22 binds AKT.
| 1 4
| 1 4

sparser
"The interaction of AKT with USP22 was detected in transiently transfected HEK293T cells as well as in human MDA-MB-231 breast cancer cells ( xref )."

sparser
"Stimulation of MDA-MB-231 breast cancer cells with EPI markedly enhanced both AKT and USP22 interaction ( xref ) as well as their phosphorylation (fig."

sparser
"Conversely, treatment with the AKT-specific inhibitor facilitated USP22 protein degradation ( xref ), increased the USP22 ubiquitination ( xref ), and inhibited the AKT and USP22 interaction in breast cancer cells ( xref )."

reach
"Conversely, treatment with the AKT-specific inhibitor facilitated USP22 protein degradation (Fig. 5, K and L), increased the USP22 ubiquitination (Fig. 5M), and inhibited the AKT and USP22 interaction in breast cancer cells (Fig. 5N)."

sparser
"Of note, mass spectrometry (MS) analysis also detected 10 AKT phosphorylated peptides in the anti-USP22 immunoprecipitants ( xref ), confirming our initial discovery of AKT-USP22 interaction and suggesting that USP22 interacts with the phosphorylated AKT upon EPI stimulation in breast cancer cells."
USP22 increases the amount of AKT.
| 2
USP22 increases the amount of AKT. 2 / 2
| 2

reach
"We have reported that USP22 mediates cell survival and proliferation by promoting the expression of BMI-1 and upregulation of activated AKT pathway in colon cancer cells."

reach
"USP22 knockdown decreased MDR related genes expression through up-regulation of Smad4 and suppression of Akt."
USP22 affects ZEB1
2 | 16 11
USP22 binds ZEB1.
2 | 3 11
2 | 3 11

sparser
"In addition, HEK-293 cells were transfected with USP22 and ZEB1 expression plasmids as indicated and the interaction between USP22 and ZEB1 was conformed in Co-IP experiment (Supplementary Fig. S xref C)."

sparser
"The results showed that only the N terminal of USP22 directly binds to ZEB1 (Fig. xref ), while the C terminal of USP22 may indirectly bind to ZEB1."

sparser
"Through a co-IP assay, we found that USP22 can bind to ZEB1 mRNA and that knockdown of USP22 markedly facilitates the ubiquitination of ZEB1 and thus enhances its protein degradation."

sparser
"Taken together, these data demonstrated that USP22 physically associates with ZEB1 in HCC cells."

sparser
"Here, our results have demonstrated that USP22 interacts with ZEB1, and USP22/ZEB1 is recruited to the ZEB1-binding elements of VEGFA promoter region and USP22 decrease the accumulation of histone H2Bub."

sparser
"Co-IP was used to validate the interaction between USP22 and ZEB1."

sparser
"Having established that USP22 associates with ZEB1, and USP22/ZEB1 is recruited to the promoter of VEGFA in HCC cells, we thus turn to examine whether USP22 participates in maintenance of ZEB1 stability."

sparser
"Next, we employed a Co-IP assay and found that USP22 interacts with ZEB1 ( xref )."

reach
"In addition, HEK-293 cells were transfected with USP22 and ZEB1 expression plasmids as indicated and the interaction between USP22 and ZEB1 was conformed in Co-IP experiment (Supplementary Fig. S3C)."

sparser
"USP22 interacts with ZEB1, and USP22/ZEB1 is recruited to the ZEB1-binding elements of VEGFA promoter region in HCC cell lines."
USP22 increases the amount of ZEB1.
| 5
USP22 increases the amount of ZEB1. 5 / 5
| 5

reach
"USP22 depletion or USP22 deubiquitinase inactive mutant USP22 (m) enhanced the ubiquitination level of ZEB1 (Fig. 4H, I)."

reach
"Thus, analysis of novel co-regulator for modulation of ZEB1 action may provide the effective strategy for anti-angiogenic therapy resistance in HCC.In this study, our results have demonstrated that USP22 up-regulates ZEB1-mediated VEGFA transcription."

reach
"Upregulation of USP22 can increase the expression of the ZEB1 and Snail transcription factors, significantly reduce the expression of E-cadherin at cell–cell junctions, and upregulate the expression of mesenchymal markers [143]."

reach
"Our results showed that ectopic expression of USP22 increased ZEB1 protein expression and USP22 deletion reduced ZEB1 protein expression, while USP22 had no obvious effect on mRNA expression of ZEB1 in HCC-derived cell lines (Fig. 4A–C)."

reach
"Ectopic expression of USP22 did not increase the protein expression of ZEB1 in the presence of ubiquitin proteasome inhibitor MG132."
USP22 decreases the amount of ZEB1.
| 4
USP22 decreases the amount of ZEB1. 4 / 4
| 4

reach
"Our results showed that ectopic expression of USP22 increased ZEB1 protein expression and USP22 deletion reduced ZEB1 protein expression, while USP22 had no obvious effect on mRNA expression of ZEB1 in HCC-derived cell lines (Fig. 4A–C)."

reach
"USP22 depletion or USP22 deubiquitinase inactive mutant USP22 (m) enhanced the ubiquitination level of ZEB1 (Fig. 4H, I)."

reach
"The results showed that overexpression of USP22 also reduced the ubiquitination level of ZEB1 in HCCLM3 cells (Fig. 4J)."

reach
"The results demonstrated that the ubiquitination level of ZEB1 was significantly decreased by ectopic expression of USP22 in HCCLM3 and Huh7 cells (Fig. 4G and Supplementary Fig. S5B)."
USP22 activates ZEB1.
| 4
USP22 activates ZEB1. 4 / 4
| 4

reach
"Reduction of ZEB1 caused by knockdown of USP22 was also prevented by MG132, on the other hand, USP22 depletion accelerated ZEB1 degradation with the treatment of cycloheximide (CHX), which is protein synthesis inhibitor, in HCC-derived cells (Fig. 4D–F and Supplementary Fig. S5A)."

reach
"While USP22 depletion decreased ZEB1 protein, not mRNA level (Fig. 6E, F and Supplementary Fig. S7A)."

reach
"According to the results of previous studies, USP22 can induce EMT by regulating TGF-beta1 in lung cancer cells [XREF_BIBR], and elevated USP22 expression can promote EMT by up-regulating ZEB1 and Snail in pancreatic cancer cells though a process that involves focal adhesion kinase (FAK) signaling [XREF_BIBR]."

reach
"ZEB1 has illustrated to be modulated by USP7, USP22 and USP51 in cancers."
USP22 affects FASN
4 | 11 14
USP22 binds FASN.
4 | 14
4 | 14

sparser
"These results suggest that H 2 O 2 induces cells apoptosis by repressing the USP22-FASN axis in p53 +/+ colorectal cancer cells."

sparser
"Depletion of the USP22-FASN axis inhibits cell growth and colorectal tumorigenesis."

sparser
"Our study demonstrates that overproduced ROS in colorectal cancer controls lipid synthesis by critically regulating the USP22-FASN axis through a p53-dependent manner."

sparser
"To determine the potential clinical relevance of the USP22-FASN axis, we next assessed the expression of USP22 and FASN proteins in serial sections of 108 human CRC specimens, and found that FASN expression levels were significantly correlated with USP22 levels (Fig. xref , r  = 0.6626, P  < 0.0001)."

sparser
"Our findings establish that the USP22-FASN axis plays an important role in regulating lipid accumulation and colorectal tumorigenesis, and targeting this axis may represent a promising therapeutic strategy for colorectal cancer."

sparser
"Because p53 mutation usually loss the transcriptional activity and responses to upstream signals differently [ xref , xref ], p53-mediated repression of the USP22-FASN axis will be abolished in p53-mutated cancers, resulting in FASN stabilization, lipid accumulation, and tumor growth."

sparser
"Thus, the USP22-FASN axis revealed in this study may be a critical mechanism for the maintenance of CSC properties and therapy resistance of human cancer, which deserves further investigation."

sparser
"Due to a high-frequency of p53 mutation and dysfunction in human cancer, activation of the USP22-FASN axis may represent a prevailing mechanism that drives tumorigenesis, indicating a promising strategy for the treatment of colorectal cancer."

sparser
"Our study demonstrates that ROS critically regulates lipid synthesis and tumorigenesis through the USP22-FASN axis in a p53-dependent manner, and targeting the USP22-FASN axis may represent a potential strategy for the treatment of colorectal cancer."

sparser
"We have demonstrated that the USP22-FASN axis is highly activated in CRC and is critical for lipid accumulation and tumorigenesis."
USP22 deubiquitinates FASN.
| 5
USP22 deubiquitinates FASN. 5 / 5
| 5

reach
"Whereas, depletion of USP22 in RKO E6 cells promoted FASN ubiquitination (Fig. 2E)."

reach
"Together, these results support that USP22 interacts with FASN and inhibits FASN ubiquitination in CRC cells."

reach
"We have found H O -induced p53 expression inhibits the transcription of USP22, which otherwise deubiquitinates and stabilizes FASN."

reach
"In p53 wild-type colorectal cancer (CRC) cells, hydrogen peroxide (H 2 O 2 )-induced p53 expression represses the transcription of deubiquitinase USP22, which otherwise deubiquitinates and stabilizes Fatty Acid Synthase (FASN), and thus inhibits fatty acid synthesis."

reach
"Overexpression of USP22 decreased FASN ubiquitination in the presence of MG132 (Fig. 2D and Fig. S2D)."
USP22 increases the amount of FASN.
| 3
USP22 increases the amount of FASN. 3 / 3
| 3

reach
"Moreover, the depletion of USP22 decreased the levels of FASN and Ki-67 in mouse xenograft tumors, and exogenous FASN rescued those effects (Fig. 6H)."

reach
"We, therefore, screened a panel of DUBs by transfecting the cDNA plasmids of 36 DUBs into 293T cells, and found USP2, USP14, USP22, and USP30 upregulated FASN levels (Fig. S2A)."

reach
"As we expected, depletion of USP22 decreased the levels of FASN in CRC cells (Fig. 3A)."
USP22 activates FASN.
| 3
USP22 activates FASN. 3 / 3
| 3

reach
"Together, these results support that USP22 promotes CRC cell proliferation and tumorigenesis by stabilizing FASN."

reach
"We next investigated the role of USP22-mediated stabilization of FASN in CRC cell proliferation and tumor growth."

reach
"While overexpression of USP22 stabilized FASN in 293T cells (Fig. 3B), USP22 depletion promoted FASN degradation in CRC cells under CHX treatment (Fig. 3C)."
USP22 affects H2Bub1
| 28
USP22 activates H2Bub1.
| 9
USP22 activates H2Bub1. 9 / 9
| 9

reach
"The deubiquitinating enzyme USP22 catalyzes H2Bub1 removal in interphase and may also be required for H2Bub1 removal in early mitosis to maintain chromosome stability."

reach
"Mechanistically, reduced USP22 expression was determined to increase local H2Bub1 abundance at the inflammation regulator SPARC locus resulting in an increase in expression [ 161 ]."

reach
"Interestingly however, this compensatory mechanism does not function in all cellular processes in which USP22 mediated H2Bub1 regulation is involved, including transcriptional activation and DNA damage repair [XREF_BIBR, XREF_BIBR, XREF_BIBR] (see Section 6 and Section 9)."

reach
"To test this possibility, we sought to determine whether reduced USP22 expression impairs H2Bub1 removal specifically within prophase chromosomes."

reach
"Recently, USP22 depletion was also determined to impair H2Bub1 removal from mitotic chromosomes, inducing mitotic chromatin compaction defects and segregation errors that promote CIN [ 151 ]."

reach
"Using complementary genetic and QuantIM approaches, we show that USP22 silencing impairs H2Bub1 removal from chromosomes during prophase and correlates with increases in multiple CIN phenotypes."

reach
"Importantly, USP22 depletion decreases H2Bub1 abundance throughout the FOXP3 locus which corresponds with FOXP3 repression [ 207 ]."

reach
"Consistent with that finding is another study that implicates the ubiquitin protease, USP22, which targets H2Bub1 for turnover in malignant colon carcinoma."

reach
"Mechanistically, USP22 depletion in human CRC cells induced a profound upregulation of secreted protein acidic and rich in cysteine (SPARC) by affecting H3K27ac and H2Bub1 occupancy on the SPARC gene."
USP22 deubiquitinates H2Bub1.
| 7
USP22 deubiquitinates H2Bub1. 7 / 7
| 7

reach
"Interestingly, neither free recombinant USP22 nor Ubp8p can efficiently deubiquitylate H2Bub1 or H2Aub1 (in the case of USP22) in vitro , indicating that an interaction(s) between the ZnF-UBP and othe[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, USP22 can also enhance DNA damage repair by deubiquitinating histones H2A and H2Bub1."

reach
"Thus, ATXN7L3 is required for the full activity of the three related DUB modules to regulate global H2Bub1 levels, whereas USP22-containing DUB module is less involved in genome-wide deubiquitylation of H2Bub1."

reach
"Along these lines it is interesting to note that neither free USP22 nor a stable recombinant subcomplex, composed of TAF5L, ATXN7L3, ENY2, and USP22, can deubiquitinate H2Aub1 or H2Bub1 in vitro, sugg[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Finally, USP22 and the 2A-DUB (KIAA1915), both target H2Aub1 (although USP22 can also deubiquitylate H2Bub1), and both were described as co-activators of androgen receptor (AR)-mediated transcription."

reach
"Unlike the ATXN7L3 DUB complex, a USP22, ATXN7L3B, and ENY2 complex can not deubiquitinate H2Bub1 efficiently in vitro Moreover, ATXN7L3B knockdown inhibits migration of breast cancer cells in vitro and limits expression of ER target genes."

reach
"USP22, USP27X, or USP51 require ATXN7L3 and ENY2 for their full activity and are unable to deubiquitinate H2Bub1 alone."
USP22 inhibits H2Bub1.
| 6
USP22 inhibits H2Bub1. 6 / 6
| 6

reach
"Western blot analysis of H2Bub1 abundance within asynchronous USP22 silenced and control cells indicates that USP22 silencing induces a moderate increase (1.1- to 1.5-fold) in the global abundance of H2Bub1."

reach
"In agreement with previous observations [49, 71], loss of USP22 expression in hIECs did indeed increase H2Bub1, a hallmark of transcriptionally active chromatin [72–74]."

reach
"We show that USP22 deficiency impairs H2Bub1 removal and induces chromatin compaction defects."

reach
"In addition, we reveal that USP22 deficiency impairs H2Bub1 removal in early mitosis and induces CIN."

reach
"Moreover, Usp22 knockout in mice corresponds with an overall increase in H2Bub1 abundance and enhances H2Bub1 accumulation in response to irradiation, which is associated with reduced γH2AX foci forma[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Compared with Baohuoside I, treatment with Rottlerin or Morusin did not enhance USP22 depletion–induced H2Bub1 up regulation, PD-L1 and Sirt1 down regulation."
USP22 decreases the amount of H2Bub1.
| 4
USP22 decreases the amount of H2Bub1. 4 / 4
| 4

reach
"To investigate whether the observed decrease in H2Bub1 levels upon GANP depletion are the result of an increased DUB activity, we tested whether the concomitant depletion of the three USP22-related DUBm-s through ENY2 KD or ATXN7L3 KD would restore normal H2Bub1 levels."

reach
"As expected, USP22 depletion led to increased H2Bub1 levels, as well as reduced Sirt1 and PD-L1 levels ( Fig. 3 C – E)."

reach
"Functionally, USP22 depletion led to increased protein level of H2Bub1, reduced PD-L1 and Sirt1in HCT116 cells."

reach
"However, while USP22 is capable of deubiquitinating H2B in vitro [36,53,60] and USP22 knockdown increases H2Bub1 levels on the IRF1 gene [18] , none of the published reports demonstrated a significant[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 increases the amount of H2Bub1.
| 2
USP22 increases the amount of H2Bub1. 2 / 2
| 2

reach
"In this study, we demonstrate that siRNA mediated USP22 depletion increases H2Bub1 levels in early mitosis and induces CIN phenotypes associated with mitotic chromatin compaction defects revealed by super-resolution microscopy."

reach
"Similarly, Atanassov et al recently reported that depletion of USP22, a major H2B DUB, not only failed to increase H2Bub1 levels but even caused a mild decrease; this was ascribed to the fact that two newly identified H2B DUBs, USP27X and USP51, compete with USP22 for binding to the SAGA complex adaptor proteins ATXN7L3 and ENY2 XREF_BIBR."
| 3 23 1
USP22 activates Carcinogenesis.
| 3 21
| 3 21

reach
"Together, these results support that USP22 promotes CRC cell proliferation and tumorigenesis by stabilizing FASN."

reach
"Together, these results suggest that FASN is stabilized by USP22 in colorectal cancer, and the dysregulated USP22/FASN axis is a important driver for tumorigenesis."

reach
"In the current study, we demonstrated that the ubiquitin hydrolase USP22 contributed to HCC tumorigenesis and promoted tumorigenic potential of HCC, suggesting an oncogenic role of USP22 consistent with the majority of published studies."

eidos
"Further , USP22 is shown to facilitate cell-cycle progression and colorectal tumorigenesis by targeting CCNB1 while in glioblastoma , USP22 promotes tumorigenesis via stabilizing KDM1A [ 89,90 ] ."

reach
"Genetic and pharmacological inhibition of PPARγ abolished the regulatory effect on ACC and ACLY transcription, and significantly decreased lipogenesis and tumorigenesis caused by USP22 expression both in HCC cells and xenograft tissues."

reach
"Silencing of USP22 enhanced T cell cytotoxicity and blocked lung tumorigenesis."

eidos
"In addition , we find that USP22 promotes de novo fatty acid synthesis and contributes to HCC tumorigenesis , however , this tumorigenicity is suppressed by inhibiting the expression of PPARgamma , ACLY , or ACC in in vivo tumorigenesis experiments ."

reach
"In our study, we found that knockdown of DUSP1 enhanced the AKT signaling pathway, whereas repression of DUSP1 by E2F6 was required for USP22-mediated carcinogenesis."

reach
"We wanted to elucidate the mechanisms by which miR-30e-5p inhibits USP22-mediated tumorigenesis in NSCLC.Previous studies have demonstrated a role for miR-30e-5p in downregulating oncogenic pathways, [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"The deubiquitylase USP22 is frequently overexpressed in cancer and contributes to tumorigenesis by driving cell cycle progression."
USP22 inhibits Carcinogenesis.
| 2 1
| 2 1

reach
"Interestingly, USP22 deletion, rather than overexpression, has been shown to drive tumorigenesis in colorectal cancer and myeloid leukemia."

sparser
"Therefore, our results from the mouse xenograft model demonstrate that USP22 silencing inhibits NSCLC tumorigenesis in vivo through regulating the MDMX–p53 pathway."

reach
"Consistently, overexpression of USP22 stimulates while downregulation of USP22 suppresses breast cancer cell growth, migration and tumorigenesis in a MYC dependent manner.113 USP13."
USP22 affects KAT2A
14 | 5 8
14 | 5 8

sparser
"Gcn5 is closely associated with USP22 or its fly orthologue, Nonstop ( xref ; xref ; xref ; xref )."

reach
"This raises the hypothesis that once USP22 associates with SAGA and GCN5, it may oppose telomere recombination in APBs by promoting the binding of POT1 to telomeres via TRF1, thereby inhibiting ATR activation."

reach
"GCN5 binds to ubiquitin-specific peptidase 22 (USP22) which deubiquitinates and thus salvages the telomeric repeat-binding factor 1 (TRF1) shelterin protein."

sparser
"There were 408 overlapping putative target genes bound by both USP22 and GCN5."

sparser
"Motif analysis showed that the sequences bound by USP22 and GCN5 shared two common motifs."

sparser
"Of these genes, 408 genes were bound potentially by both USP22 and GCN5, which accounted of 16.8% of genes recognized by USP22 and 18.1% recognized by GCN5 ( xref )."

reach
"The polypeptide composition of these purified complexes was assessed by silver staining, which revealed a pattern of 10-12 comigrating proteins shared by the USP22 and hGCN5 complexes."

reach
"Normalized ChIP‐seq signals for the individual SAGA subunits demonstrated inducible binding of GCN5, USP22, and ATXN7L3 after treatment (Fig 1C)."

sparser
"We found either USP22 antibody or GCN5 antibody precipitated a large number of DNA fragments, indicating that either USP22 or GCN5 widely interacted with different genes in the genome of HeLa cells."

sparser
"MACS analysis indicated that there were many genes potentially bound by USP22 or GCN5."
USP22 affects PALB2
2 | 12 12
USP22 binds PALB2.
2 | 7 12
2 | 7 12

sparser
"To elucidate if USP22 was directly binding PALB2 WD40 we performed in-vitro pulldown experiments using MBP-PALB2 WD40 as the bait along with His-USP22 ( xref )."

sparser
"To evaluate USP22-PALB2 binding in cells an IP was performed with FLAG-PALB2 in H1299 lung adenocarcinoma cells in which endogenous USP22 was successfully pulled down ( xref )."

sparser
"USP22 directly interacts with PALB2 to promote DNA homologous recombination repair and inhibit the killing effect of cisplatin on cancer cells."

sparser
"USP22 DUB Domain Interacts with PALB2 WD40 Domain."

sparser
"Our in-silico analysis showed an interaction between the C-terminal DUB domain of USP22 and the C-terminal WD40 domain of PALB2 that has been previously crystallized ( xref , PDB: 2W18) xref ."

sparser
"PALB2 WD40 Domain Directly Binds USP22 and stimulates its Catalytic Activity."

sparser
"Using our in-silico model of PALB2-USP22 binding ( xref ) we predicted several residues on the PALB2 WD40 domain that looked critical for this interaction; S951, N953 (upper panel), R942 (middle panel), and H913 (lower panel)."

sparser
"We identified a direct interaction between the C-terminal WD40 domain of PALB2 and the DUB domain of USP22 and that this interaction was necessary and sufficient to activate USP22 catalytic DUB activity in-vitro ."

reach
"To assay for this we used the U2OS cell line and overexpressed either FLAG-H2B or FLAG-H2B and analyzed recruitment of GFP-USP22, Rad51, PALB2, and BRCA2 after induction of mCherry-LacI-FOKI (Figure 2d,e)."

sparser
"Understanding the interaction of USP22-PALB2 and indeed the interaction of other DUBs with WD40 domain containing proteins could present a new way to target DUBs for cancer therapies."
USP22 increases the amount of PALB2.
| 5
USP22 increases the amount of PALB2. 5 / 5
| 5

reach
"USP22 modulates PALB2 and BRCA2 levels to promote chemoresistance in lung adenocarcinoma."

reach
"To determine whether USP22 is modulating PALB2 and BRCA2 protein levels at the transcriptional level we performed qPCR of BRCA2 and PALB2 with and without USP22 knockdown after 48 hours (Figure S1b)."

reach
"To further validate USP22 was modulating and stabilizing BRCA2 and PALB2 levels at the translational level, cells were depleted of USP22 again with and without treatment of the proteasome inhibitor MG132 to see if this could rescue PALB2 and BRCA2 levels in a USP22 knockdown background (Figure S1c)."

reach
"Lastly, we show USP22 positively regulates BRCA2 and PALB2 levels at the protein level."

reach
"Our study reveals USP22 positively regulates PALB2 and BRCA2 levels at the translational level."
| 1 24
| 1 21

reach
"USP22 silencing suppresses in vitro GC cell migration and invasiveness."

eidos
"Liu et al. reported that USP22 promoted GC progression through the activation of c-Myc / NAMPT / SIRT1-dependent FOXO1 and YAP signaling pathways [ 32 ] ."

reach
"USP22 knockdown significantly decreased in vitro survival, proliferation, migration, and invasiveness of GC cells compared with the controls."

reach
"In this study, we show that USP22 knockdown significantly decreases migration and invasiveness of GC cells."

reach
"Meanwhile, USP22 is reported to promote GC distant metastasis (9, 10, 23)."

reach
"USP22 silencing inhibits GC cells proliferation."

reach
"These results suggest that USP22 promotes in vitro GC cell migration and invasion."

reach
"USP22 promotes in vitro GC progression by modulating c-Myc/NAMPT/SIRT1- dependent FOXO1 and YAP signaling pathways."

reach
"Meanwhile, the colony formation assays of MGC-803 cells also revealed that USP22 knockdown suppressed the proliferative ability of GC cells."

reach
"Liu et al. reported that USP22 promoted GC progression through the activation of c-Myc/NAMPT/SIRT1-dependent FOXO1 and YAP signaling pathways [32]."
| 3

reach
"USP22 depletion promotes in vitro GC cell apoptosis."

reach
"Moreover, USP22 knockdown induces apoptosis in GC cells."

reach
"Knockdown of USP22 suppresses GC xenografts growth."
USP22 affects cisplatin
| 24
USP22 activates cisplatin.
| 14
| 14

reach
"Wang et al. [10] reported that USP22 induced cisplatin resistance in LUAD by modulating DNA damage repair mediated by γH2AX and apoptosis mediated by Ku70/Bax."

reach
"To further confirm that USP22 induces cisplatin resistance via Sirt1, we added flow cytometric analysis results."

reach
"The study reveal the dual mechanism of USP22 involvement in cisplatin resistance : (1) USP22 enhances DNA damage repair and induce cisplatin resistance by promoting the phosphorylation of histone H2AX via deubiquitinating histone H2A."

reach
"USP22 regulates ALDH activity by transcriptionally regulating ALDH1A3 levels.Enhanced DNA repair processes, increased cell stemness characteristics, and inhibition of apoptotic pathways may be major factors in USP22-induced cisplatin resistance."

reach
"These results confirm that USP22 is involved in the cisplatin resistance of A549 and CDDP cells and H2AX, gammaH2AX, and Sirt1 may be responsible for USP22 mediated cisplatin resistance."

reach
"USP22 Induces Cisplatin Resistance in Lung Adenocarcinoma by Regulating gammaH2AX Mediated DNA Damage Repair and Ku70 and Bax Mediated Apoptosis."

reach
"Overall, downregulation of USP22 is expected to enhance the therapeutic effect of cisplatin in tumors."

reach
"To verify whether inhibition of USP22 expression could reverse the cisplatin resistance of A549 and CDDP cells, CCK8 assays showed that, after the inhibition of USP22 expression, the 48h IC50 of A549 and CDDP decreased from 0.925 +/- 0.04 muM to 0.337 +/- 0.03 muM."

reach
"According to that model, USP22 enhances DNA damage repair and cisplatin resistance by deubiquitinating histone H2A, which in turn facilitates the phosphorylation of histone H2AX."

reach
"To verify whether inhibition of USP22 expression could reverse the cisplatin resistance of A549 and CDDP cells, CCK8 assays showed that, after the inhibition of USP22 expression, the 48h IC50 of A549 and CDDP decreased from 0.925 +/- 0.04 muM to 0.337 +/- 0.03 muM."
USP22 inhibits cisplatin.
| 10
| 10

reach
"USP22 inhibition restores cisplatin sensitivity in cisplatin-resistant lung cancer cells.Cisplatin can induce cell death by activating mitochondrial apoptosis."

reach
"The Cisplatin Sensitivity in Cisplatin Resistant A549 and CDDP Cells Was Restored by USP22 Inhibition."

reach
"These results indicate that the cisplatin sensitivity in cisplatin resistant A549 and CDDP cells was restored by USP22 inhibition."

reach
"In addition, the cisplatin sensitivity in cisplatin resistant A549 and CDDP cells was restored by USP22 inhibition in vivo and vitro."

reach
"USP22 directly interacts with PALB2 to promote DNA homologous recombination repair and inhibit the killing effect of cisplatin on cancer cells."

reach
"The cisplatin sensitivity in cisplatin resistant A549 and CDDP cells was restored by USP22 inhibition in vivo and vitro."

reach
"Silencing USP22 can inhibit the Wnt/β-catenin pathway to reduce cell stemness and enhance the sensitivity of PC cells to cisplatin."

reach
"Inhibiting USP22 Expression Enhanced Cisplatin Sensitivity in Cisplatin Resistant A549 and CDDP Cell Nude Mouse Xenografts."

reach
"These results suggest that inhibiting USP22 expression enhanced cisplatin sensitivity in lung adenocarcinoma by downregulation of Sirt1 and gammaH2AX in vivo."

reach
"Furthermore, USP22 knockout significantly impaired non homologous DNA damage repair capacity, enhanced cisplatin and irradiation induced apoptosis in these cells."
USP22 affects SNAI1
| 18 6
USP22 binds SNAI1.
| 5 6
| 5 6

sparser
"WB results showed that the mutation of lysine residues in C-terminal or N-terminal had no effect on the binding of USP22 and Snail1 ( Fig. 8 C)."

sparser
"Despite the mutation of lysine residues in C-terminal or N-terminal having no effect on the binding of USP22 and Snail1, the WT Snail1 and the Snail1(C-8KR) mutant plasmid could decrease the ubiquityl[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Therefore, the USP22-Snail1 signal axis may be a new target for delaying diabetes TIF. (i) EMT induced by HG can promote TIF, which is one of the main pathological changes induced by DN."

reach
"Co-IP experiment results also showed that USP22 could bind to Snail1 in NRK-52E cells ( Fig. 7 C), and HG stimulation obviously increased their interactions ( Fig. 7 C)."

reach
"We further performed Co-IP experiment to determine whether the mutation of lysine residues in C-terminal and N-terminal affected the binding between USP22 and Snail1."

sparser
"Further study found that quercetin inhibited the interaction between USP22 and Snail1, thereby reducing the stability of Snail1."

reach
"Further study found that quercetin inhibited the interaction between USP22 and Snail1, thereby reducing the stability of Snail1."

reach
"Despite the mutation of lysine residues in C-terminal or N-terminal having no effect on the binding of USP22 and Snail1, the WT Snail1 and the Snail1(C-8KR) mutant plasmid could decrease the ubiquityl[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Co-IP experiment results also showed that USP22 could bind to Snail1 in NRK-52E cells ( Fig. 7 C), and HG stimulation obviously increased their interactions ( Fig. 7 C)."

sparser
"Co-IP assay was used to verify whether USP22 and Snail1 can interact with each other."
USP22 increases the amount of SNAI1.
| 7
USP22 increases the amount of SNAI1. 7 / 7
| 7

reach
"Results revealed that USP22 overexpression promoted the EMT process and increased the expression of the EMT transcriptional factor Snail1."

reach
"WB showed that USP22 overexpression increased the protein level of Snail1 compared with those in NG and HG, whereas Snail1 knockdown reversed the increase in Snail1 caused by USP22 overexpression ( Fi[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"However, USP22 deficiency reversed HG-induced renal EMT and reduced Snail1 levels, thereby reducing the accumulation of TGF-β and ECM components, such as FN, Collagen I, and Collagen Ⅳ."

reach
"These foundations demonstrated that USP22 could deubiquitinate Snail1 to increase Snail1 protein expression in model cells, which may aggravate renal EMT process."

reach
"Upregulation of USP22 can increase the expression of the ZEB1 and Snail transcription factors, significantly reduce the expression of E-cadherin at cell–cell junctions, and upregulate the expression of mesenchymal markers [143]."

reach
"These results indicated that USP22 silencing ameliorated renal damage in db/db mice.IHC staining and WB results showed that AAV9-USP22-RNAi adeno-associated virus downregulated Snail1 expression in ki[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP22 promotes EMT by stabilizing Snail expression, further affecting tubulointerstitial fibrosis98."
USP22 activates SNAI1.
| 4
USP22 activates SNAI1. 4 / 4
| 4

reach
"Moreover, the deficiency of USP22 partly reversed the upregulation of Snail1 induced by HG ( Fig. 5 E)."

reach
"WB showed that USP22 overexpression increased the protein level of Snail1 compared with those in NG and HG, whereas Snail1 knockdown reversed the increase in Snail1 caused by USP22 overexpression ( Fi[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"These results indicated that USP22 deficiency reversed renal EMT and TIF in diabetic kidneys by blocking the Snail1 signaling pathway (see Fig. 11 )."

reach
"According to the results of previous studies, USP22 can induce EMT by regulating TGF-beta1 in lung cancer cells [XREF_BIBR], and elevated USP22 expression can promote EMT by up-regulating ZEB1 and Snail in pancreatic cancer cells though a process that involves focal adhesion kinase (FAK) signaling [XREF_BIBR]."
USP22 deubiquitinates SNAI1.
| 2
USP22 deubiquitinates SNAI1 on lysine. 2 / 2
| 2

reach
"Therefore, USP22 deubiquitinated Snail1 through the N-terminal lysine residues of Snail1."

reach
"Thus, USP22 deubiquitinated Snail1 through the N-terminal lysine residues of Snail1."
USP22 affects STING1
1 1 | 5 16
USP22 binds STING1.
1 | 16
1 | 16

sparser
"In line with this, USP22-induced ISG expression could be reversed as well in USP22-STING dKO HT-29 cells (Fig. xref )."

sparser
"Additionally, USP22-mediated increases in IFN-λ expression could also largely be reduced upon USP22-STING dKO, whereas expression of IFN-α and IFN-β remained largely unaffected (Fig. xref )."

sparser
"The differential response to ISD, but not poly(I:C), and the reversal of the IFN signature in USP22-STING dKO hIECs suggests an important role of USP22 in the control of STING-induced type III IFN signaling."

sparser
"E. Western blot 831 analysis of STING and USP22 expression levels in control-NHT, control-USP22 KO 832 #1 and #6, STING-NHT and STING-USP22 KO #1 and #6 double KO (dKO) HT-29 833 cells."
| DOI

sparser
"To investigate the significance of USP22 and the 326 resulting STING-mediated upregulation of type III IFN and ISG signaling for viral327 defense, the role of the USP22-STING axis was tested during SARSexpression of STING, compared to wild-type (WT) and NHT 332 CRISPR/Cas9 control Caco-2 cells (Figure 6A)."
| DOI

sparser
"Intriguingly, 351 USP22-STING dKO hIECs exhibit higher SARS-CoV-2 replication rates as well as the formation of more de novo infectious viral particles compared to USP22 inflammatory diseases and cancer 1,2 ."
| DOI

sparser
"To confirm the role of STING in USP22-induced type III IFN signaling, USP22-STING dKO HT-29 cells were generated (Fig. xref )."

sparser
"USP22-STING dKO cells exhibited strikingly reduced levels of basal and phosphorylated STAT1 protein compared to USP22 KO HT-29 cells (Fig. xref ), suggesting a STING-dependent rescue of the USP22-dependent IFN signature."

sparser
"F. Quantification of relative SARS-CoV-2 835 genome expression of SARS-CoV-2-infected control-NHT, control-USP22 KO #1 and 836 #6, STING-NHT and STING-USP22 KO #1 and #6 dKO HT-29 cells at 24 hpi."
| DOI

sparser
"In line with this, USP22-283 induced ISG expression could be reversed as well in USP22-STING dKO HT-29 cells 284 (Figure 4H)."
| DOI
USP22 deubiquitinates STING1.
1 | 5
USP22 deubiquitinates STING1. 5 / 6
1 | 5

reach
"We furthermore demonstrate for the first time that in the absence of viral 421 infections or exogenous IFN, loss of USP22 expression resulted in basal and 2'3'-422 cGAMP-induced STING ubiquitination in hIECs."
| DOI

reach
"USP22 negatively regulates STING activation and ubiquitination."
| DOI

reach
"USP22 negatively regulates STING activation and ubiquitination."

reach
"288 289 USP22 negatively regulates STING activation and ubiquitination 290 The differential response to ISD, but not poly(I:C), and the reversal of the IFN signature 291 in USP22-STING dKO hIECs suggests an important role of USP22 in the control of 292 STING-induced type III IFN signaling."
| DOI

reach
"However, USP22 does not seem to have a clear specificity in terms of its selectivity for PPARγ deubiquitination, and studies have reported that USP22 can deubiquitinate ATG5 and STING ."
USP22 affects SPI1
3 | 15 5
USP22 binds SPI1.
3 | 5 5
3 | 5 5

sparser
"In line with the truncated mutant of USP22, the C-terminal ubiquitin-specific peptidase domain of USP22 containing a point mutation could bind SPI1 weakly, whereas the N-terminal zinc finger domain of USP22 containing a point mutation could bind SPI1 more strongly (Fig.  xref )."

sparser
"These results indicate that USP22 interacts with SPI1 and enhances SPI1 stability through deubiquitination."

reach
"Co-IP assays were then performed to verify the interaction between USP22 and SPI1."

reach
"The C-terminal ubiquitin-specific peptidase domain of USP22 binds SPI1 strongly, whereas the N-terminal zinc finger domain binds SPI1 weakly (Fig. 5d)."

reach
"In line with the truncated mutant of USP22, the C-terminal ubiquitin-specific peptidase domain of USP22 containing a point mutation could bind SPI1 weakly, whereas the N-terminal zinc finger domain of USP22 containing a point mutation could bind SPI1 more strongly (Fig. 5g)."

reach
"Thus, the C-terminal ubiquitin-specific peptidase domain of USP22 is the dominant regulator of binding between USP22 and SPI1."

reach
"Co-IP assays were performed to determine the interaction between USP22 and SPI1."

sparser
"USP22 interacts with SPI1."

sparser
"Co-IP assays were then performed to verify the interaction between USP22 and SPI1."

sparser
"To identify the region within USP22 interacting with SPI1, we generated two truncated mutants of USP22 in HEK-293T cells."
USP22 activates SPI1.
| 6
USP22 activates SPI1. 6 / 6
| 6

reach
"On the other hand, USP22 deficiency decreases PU.1 stability to promote myeloid leukemia [10]."

reach
"Research has shown that USP22 can enhance myeloid differentiation in Kras mice by stabilizing the PU.1 protein and decreasing the risk of AML [14]."

reach
"Further, following CHX treatment, USP22 overexpression significantly attenuated SPI1 degradation, which in contrast, was strikingly facilitated in USP22 knockdown cells (Fig. 4i, j and Supplementary Fig. 4h, i)."

reach
"These results indicate that USP22 interacts with SPI1 and enhances SPI1 stability through deubiquitination."

reach
"Mechanistically, PRDM1 enhances USP22 transcription, thus reducing SPI1 protein degradation through deubiquitination, which enhances PD-L1 transcription."

reach
"Here, by utilizing the DUB siRNA library, we found that USP22 could reduce SPI1 protein degradation through deubiquitination."
USP22 deubiquitinates SPI1.
| 4
USP22 deubiquitinates SPI1. 4 / 4
| 4

reach
"USP22 can deubiquitinate SPI1 to transcriptionally upregulate PD-L1."

reach
"USP22 deubiquitinates SPI1 and enhances its stability, thereby regulating PD-L1 expression."

reach
"Mechanistically, SPI1 serves as a transcription factor for PD-L1 and is deubiquitinated by USP22 (Fig. 3)."

reach
"33 , 34 In addition, USP22 can also deubiquitinate and stabilize the SPI1 protein, thereby enhancing PD‐L1 transcription and expression."
USP22 affects CDKN1A
| 22 1
USP22 decreases the amount of CDKN1A.
| 15
USP22 decreases the amount of CDKN1A. 10 / 15
| 15

reach
"In MGC-803 cells and SGC-7901 cells, knockdown of USP22 and BMI1 both increased P21 expression and reduced the expression of CSC stemness genes of CD133 and SOX2."

reach
"Moreover, USP22 knockdown upregulates the transcription of p21 by upregulating the ubiquitylation of FBP1."

reach
"Recent studies have demonstrated that USP22 can inhibit the transcription of the p21 gene by de-ubiquitinating the transcriptional regulator FBP1, leading to cell proliferation and tumorigenesis [XREF_BIBR]."

reach
"Furthermore, our results showed that USP22 deletion caused down -- regulation of cyclin D2 expression and up -- regulation of p15 and p21 expression."

reach
"We found that USP22 depletion inhibited the proliferation of the human GC cell lines MGC-803 and SGC-7901 cells and increased expression of P21, indicating cell cycle arrest [XREF_BIBR]."

reach
"In pharyngeal carcinoma, knockdown of USP22 not only upregulated p21 expression but also upregulated p27 expression and suppressed retinoblastoma protein (RB) phosphorylation."

reach
"USP22 has also been demonstrated to inhibit transcription of the p21 gene by deubiquitinating the transcriptional regulator, FBP1, leading to cell proliferation and tumorigenesis."

reach
"USP22-mediated deubiquitination of PTEN inhibits pancreatic cancer progression by inducing p21 expression."

reach
"A previous study demonstrated that USP22 could inhibit the transcription of the cyclin-dependent kinase inhibitor 1A (p21) gene by deubiquitinating the transcriptional regulator fructose-1,6-bisphosphatase (FBP1) (5)."

reach
"Furthermore, USP22 small interfering (si)RNA decreased the expression of Cyclin B and Survivin, and increased the expression of p21 in HCC (7)."
USP22 increases the amount of CDKN1A.
| 4 1
USP22 increases the amount of CDKN1A. 5 / 5
| 4 1

reach
"We found USP22 silencing led to decreased expression of BMI1, as well as decreased P21 levels."

reach
"USP22 also play roles in cell cycle regulation, where depletion of USP22 increases the expression of p53 and p21, inhibits proliferation, and induces cell cycle arrest at G1 phase."

reach
"Furthermore, USP22 small interfering (si)RNA decreased the expression of Cyclin B and Survivin, and increased the expression of p21 in HCC (7)."

reach
"USP22 siRNA transfection increased the expression of p53 and p21, decreased cyclin E expression in bladder cancer cells."

sparser
"USP22 induced p21 expression via PTEN in pancreatic cancer."
USP22 inhibits CDKN1A.
| 3
USP22 inhibits CDKN1A. 3 / 3
| 3

reach
"USP22 knockdown, using lentivirus delivered siRNA, increased the expression levels of cell cycle proteins P21 and P27, but reduced the levels of phosphorylated retinoblastoma protein, resulting in the inhibition of FaDu cell growth and proliferation."

reach
"Hence USP22 silencing reduces the capacity of FBP1 to repress p21 (independently of TP53 status), which in turn inhibits CDKs to prevent the G1/S transition and resulting in G1 accumulation [XREF_BIBR]."

reach
"In addition, USP22 was shown to induce cell cycle protein-dependent kinase inhibitor 1A (CDKN1A) in pancreatic cancer, and MDM2 inhibitors enhanced the anti-pancreatic cancer effect of USP22 overexpression [166]."
USP22 affects COPS5
1 | 10 10
USP22 binds COPS5.
| 6 10
| 6 10

sparser
"Moreover, USP22 also interacted with CSN5 and stabilized CSN5 through deubiquitination."

sparser
"USP22 also interacted with CSN5 and kept its stability via deubiquitination."

reach
"Moreover, USP22 also interacted with CSN5 and stabilized CSN5 through deubiquitination."

reach
"USP22, a ubiquitin carboxyl-terminal hydrolase, interacted with JAB1 and stabilized JAB1 through deubiquitination (Wang et al. 2020b)."

sparser
"On the other hand, USP22 interacts with CSN5 and deubiquitinates it, thereby facilitating the interaction between PD-L1 and CSN5 ( xref )."

sparser
"USP22 interacted with CSN5 and deubiquitinated CSN5."

sparser
"Additionally, USP22 forms a complex with CSN5, maintaining its stability via deubiquitination."

sparser
"Since CSN5 bound to PD-L1 [ xref ], we wondered if USP22 interacted with CSN5 physically."

reach
"Additionally, USP22 forms a complex with CSN5, maintaining its stability via deubiquitination."

sparser
"As expected, USP22 interacted with CSN5 exogenously and endogenously (Fig.  xref a, Fig. xref b)."
USP22 deubiquitinates COPS5.
1 | 4
USP22 deubiquitinates COPS5. 4 / 5
1 | 4

reach
"On the other hand, USP22 deubiquitinates CSN5 and regulates PD-L1 protein levels through the USP22-CSN5-PD-L1 axis."

reach
"Furthermore, when the PD-L1 protein is depleted, USP22 deubiquitinates CSN5 and controls the level of PD-L1 protein through the USP22-CSN5-PD-L1 axis, thereby strengthening antitumour immune reactions [75]."

reach
"USP22 can deubiquitinate PD-L1 itself or CSN5 for their stabilization [233] (Figure 3)."
| PMC

reach
"USP22 deubiquitinates CSN5 polyubiquitin chains and stabilizes CSN5 protein, resulting in enhanced PD-L1 expression [102]."
PALB2 affects USP22
2 | 7 12
2 | 7 12

sparser
"To elucidate if USP22 was directly binding PALB2 WD40 we performed in-vitro pulldown experiments using MBP-PALB2 WD40 as the bait along with His-USP22 ( xref )."

sparser
"To evaluate USP22-PALB2 binding in cells an IP was performed with FLAG-PALB2 in H1299 lung adenocarcinoma cells in which endogenous USP22 was successfully pulled down ( xref )."

sparser
"USP22 directly interacts with PALB2 to promote DNA homologous recombination repair and inhibit the killing effect of cisplatin on cancer cells."

sparser
"USP22 DUB Domain Interacts with PALB2 WD40 Domain."

sparser
"Our in-silico analysis showed an interaction between the C-terminal DUB domain of USP22 and the C-terminal WD40 domain of PALB2 that has been previously crystallized ( xref , PDB: 2W18) xref ."

sparser
"PALB2 WD40 Domain Directly Binds USP22 and stimulates its Catalytic Activity."

sparser
"Using our in-silico model of PALB2-USP22 binding ( xref ) we predicted several residues on the PALB2 WD40 domain that looked critical for this interaction; S951, N953 (upper panel), R942 (middle panel), and H913 (lower panel)."

sparser
"We identified a direct interaction between the C-terminal WD40 domain of PALB2 and the DUB domain of USP22 and that this interaction was necessary and sufficient to activate USP22 catalytic DUB activity in-vitro ."

reach
"To assay for this we used the U2OS cell line and overexpressed either FLAG-H2B or FLAG-H2B and analyzed recruitment of GFP-USP22, Rad51, PALB2, and BRCA2 after induction of mCherry-LacI-FOKI (Figure 2d,e)."

sparser
"Understanding the interaction of USP22-PALB2 and indeed the interaction of other DUBs with WD40 domain containing proteins could present a new way to target DUBs for cancer therapies."
USP22 affects ferroptosis
| 20
USP22 inhibits ferroptosis.
| 17
| 17

reach
"Overexpression of USP22 reversed LPS-induced cellular inflammation, attenuated decidualization, and inhibited ferroptosis."

reach
"In particular, USP22 raised GSH content and reduced the occurrence of ferroptosis in cardiomyocytes by activating the Sirt1-p53/SLC7A11 axis [252]."

reach
"A recent study uncovered that USP22 can repress the transcriptional activity of p53 to upregulate SLC7A11 expression, thereby suppressing ferroptosis in cardiomyocytes [28]."

reach
"USP22 inhibits Sorafenib‐induced ferroptosis through transcriptional suppression of TFRC mediated by deubiquitinating H2BK120ub."

reach
"Additionally, A549 and H1975 LUAD cells with USP22 silencing exhibited a higher expression of ACSL4 and lower levels of SLC7A11 and GPX4 than sh‐Ctrl controls; however, these changes could be markedly reversed by COL17A1 reexpression (Figure 6J,K), indicating that USP22 silencing induces cell ferroptosis via COL17A1 downregulation."

reach
"USP22 promotes gefitinib resistance and inhibits ferroptosis in non-small cell lung cancer by deubiquitination of MDM2."

reach
"Additionally, USP22 acts as a novel inducer of Sorafenib resistance and suppresses Sorafenib-triggered ferroptosis in hepatocellular carcinoma cells."

reach
"Our results showed that USP22 knockout enhanced ferroptosis induced by different drugs, as evidenced by decreased cell viability (Figure S5A) and elevated cellular lipid ROS levels (Figure S5B) compared to control cells."

reach
"Similarly, through the SIRT1/p53/SLC7A11 connection, USP22 overexpression may prevent ferroptosis-induced cardiomyocyte death and shield against MI/RI [40]."

reach
"Finally, in vivo study proved that miR-144-3p antagomir could restore the impaired function of pancreatic β-cells by suppressing ferroptosis to exert anti-hyperglycemic effect on T2DM mice.In short, the abnormal up-regulation of miR-144-3p clearly suppressed the expression of USP22 mRNA and inhibited its downstream effects on the ferroptosis pathway to cause the dysfunction of pancreatic β cells (Fig. 6), thereby promoting the progression of T2DM."
USP22 activates ferroptosis.
| 3
| 3

reach
"Overexpression of USP22 ameliorates LPS-induced endometrial stromal cells inflammation and modulates cells decidualization by inhibiting ferroptosis."

reach
"USP22 promotes the proliferation of hepatocellular carcinoma cells by stabilising CDK11B and enhances resistance to Sorafenib by inhibiting ferroptosis through the USP22/H2BK120ub/TFRC axis.So far, more than 40 USPs have been directly or indirectly associated with relevant cancer processes."

reach
"Besides, USP22 overexpression enhanced gefitinib resistance and ferroptosis protection in NSCLC cells."
USP22 affects USP22
| 18
USP22 inhibits USP22.
| 5
USP22 inhibits USP22. 5 / 5
| 5

reach
"However, the increase in SIRT1 levels enhances its deacetylation activity, which in turn, deacetylates USP22 and other SAGA components, thereby decreasing the enzymatic activity of USP22 ."

reach
"Lastly, ubiquitylation of USP22 mediated by the anaphase promoter complex and cyclosome (APC/C) induces USP22 protein degradation during the cell cycle [XREF_BIBR]."

reach
"Supporting the regulation of USP22 by m A modification, knockdown of the methyltransferase METTL3 increased USP22 expression in ALKBH5-silenced 143B cells (Fig. 4H; Supplementary Fig. S9A)."

reach
"Loss of USP22 leads to upregulation of many ISGs, some with important functions in viral defense, like OAS1, -2 and -3, MX1 and IFI27, suggesting a potential role of USP22 in the regulation of IFN signaling and viral responses (Fig. 2B)."

reach
"Stimulation with EPI markedly reduced USP22 ubiquitination levels (Fig. 5A), supporting our speculation that EPI protects USP22 from ubiquitination-mediated degradation."
USP22 decreases the amount of USP22.
| 4
USP22 decreases the amount of USP22. 4 / 5
| 4

reach
"The results of western blot analysis showed that USP22R restored the levels of USP22 and COX-2 downregulated by USP22 siRNA."

reach
"XREF_FIG, USP22 specific siRNA effectively inhibited USP22 gene transcription and protein expression."

reach
"USP22 siRNA transfection (58 nM) for 24 h was observed to significantly reduce expression of USP22 mRNA and protein in U87 and U251 cells (XREF_FIG)."

reach
"However, ablation of RALY resulted in a considerable decrease in USP22 protein levels; whereas RALY overexpression increased USP22 protein levels (Figure 6F)."
USP22 activates USP22.
| 5
USP22 activates USP22. 5 / 5
| 5

reach
"In addition, phosphorylation of Thr147 and Ser237 of USP22 by CDK1 enhances the deubiquitination activity of USP22 on cyclin B1."

reach
"Phosphorylation of USP22 at T147 and S237 by cyclin dependent kinase 1 (CDK1) was shown to activate USP22 to deubiquitylate cyclin B1."

reach
"Unsurprisingly, the expression level of USP22 protein in HG-induced INS-1 cells was dramatically increased after transfecting USP22 OE."

reach
"Interestingly, USP22 activity is regulated by CDK1, which catalyzes USP22 phosphorylation to elevate USP22 ability in CCNB1 deubiquitination and stabilization."

reach
"These findings suggested that activation of the mTORC1 pathway after USP22 overexpression induces therapeutic susceptibility to rapamycin.2.4 USP22 regulates FKBP12 to activate mTORC1 through deubiquitylation."
USP22 increases the amount of USP22.
| 4
USP22 increases the amount of USP22. 4 / 4
| 4

reach
"In HG-treated NRK-52E cells, USP22-FLAG overexpression further increased USP22 expression ( Fig. 3 C), downregulated E-cadherin and ZO-1expression, and upregulated Vimentin expression ( Fig. 3 D)."

reach
"The LSL Usp22 allele inserted at the Rosa26 locus allows tissue-specific overexpression (OE) of USP22 upon exposure to Cre recombinase [17]."

reach
"USP22 knockdown (Figure 3A) by using siRNA (si-USP22) dramatically decreased USP22 level and, in parallel, the levels of cyclin D1 and of cell vitality (Figure 3B) and viability (Figure 3C) compared with the negative control."

reach
"It was identified that USP22 siRNA decreased the expression of USP22 protein (Fig. 2A) and also inhibited cell growth in OSCC cells (Fig. 2B)."
USP22 affects MDM4
2 | 11 6
USP22 binds MDM4.
2 | 8 6
2 | 8 6

sparser
"Our data suggest that MDMX directly binds to USP22 in NSCLC cells."

reach
"Our co-immunoprecipitation analysis shows that USP22 interacts with MDMX in NSCLC cells."

reach
"MDMX Directly Binds to USP22 in NSCLC Cells."

sparser
"These data indicate that MDMX directly binds to Flag-tagged USP22 in A549-USP22 cells."

sparser
"Furthermore, we confirmed that MDMX directly interacts with endogenous USP22 in NCl-H460 cells, as MDMX was detected in USP22 immunoprecipitates and vice versa ( xref C,D)."

reach
"To investigate the interaction between MDMX and USP22, co-immunoprecipitation analysis was performed in NSCLC cell lines."

reach
"Furthermore, we confirmed that MDMX directly interacts with endogenous USP22 in NCl-H460 cells, as MDMX was detected in USP22 immunoprecipitates and vice versa (XREF_FIG C, D)."

reach
"Our data suggest that MDMX directly binds to USP22 in NSCLC cells."

sparser
"Our co-immunoprecipitation analysis shows that USP22 interacts with MDMX in NSCLC cells."

reach
"USP22 interacts with MDMX and leading to the inhibition of p53 dependent apoptosis [29]."
USP22 increases the amount of MDM4.
| 3
USP22 increases the amount of MDM4. 3 / 3
| 3

reach
"In addition, similar to our in vitro findings, USP22 silencing led to a decreased level of MDMX and increased levels of p53 pathway proteins in xenograft tumor tissues (XREF_FIG C)."

reach
"Interestingly, we found that in A549 and NCI-H460 cells, USP22 silencing, while activating the p53 pathway, decreased MDMX protein expression (XREF_FIG C), which was confirmed with immunofluorescence (IF) analysis (XREF_FIG D)."

reach
"Additionally, USP22 silencing downregulates MDMX protein expression and activates the p53 pathway."
USP22 affects E2F6
3 | 6 10
USP22 binds E2F6.
3 | 3 10
3 | 3 10

reach
"Therefore, we validated the interaction of E2F6 and USP22; the results revealed that E2F6, a pocket protein-independent transcription repressor, was capable of interacting with USP22 ( Fig. 1 C–E), in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Importantly, we found that the function of the USP22-E2F6 axis in promoting tumor cell growth was fulfilled by a decrease in DUSP1 activities, wherein E2F6 could directly bind to the promoter region o[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Therefore, the USP22-E2F6 axis has a tumor-promoting capacity in HCC, with the function to downregulate DUSP1 activities."

sparser
"Therefore, we validated the interaction of E2F6 and USP22; the results revealed that E2F6, a pocket protein-independent transcription repressor, was capable of interacting with USP22 ( Fig. 1 C–E), in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Thus, herein we provide the evidence that DUSP1 inhibition by the USP22-E2F6 axis is the upstream event that activates AKT signaling and drives HCC growth."

reach
"These results indicate a functional significance of the interaction between USP22 and E2F6 in HCC."

sparser
"To explore the functional importance of USP22-E2F6 regulation in HCC, we examined the transcriptional profiles of Huh7 cells with or without USP22 or E2F6 silencing by RNA-seq."

sparser
"These results indicate a functional significance of the interaction between USP22 and E2F6 in HCC."

sparser
"Overall, these results suggest that USP22-E2F6 regulation is a clinically relevant event in HCC."

reach
"Moreover, it has been demonstrated that the deubiquitinating enzyme USP22 can directly interact with and stabilize E2F6 in HCC cells, which indicates the presence of corresponding E3 ubiquitin ligases of E2F6 forming a dynamic bidirectional regulatory system for ubiquitin modifications 47."
USP22 activates E2F6.
| 3
USP22 activates E2F6. 3 / 3
| 3

reach
"Therefore, we validated the interaction of E2F6 and USP22; the results revealed that E2F6, a pocket protein-independent transcription repressor, was capable of interacting with USP22 ( Fig. 1 C–E), in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Notably, the reduction in the E2F6 protein level caused by USP22 inhibition could be reversed by the proteasome inhibitor MG132 ( Fig. 1 H), indicating that USP22 increased E2F6 stability by inhibitin[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"We then generated four signatures based on the transcriptional alterations caused by E2F6 or USP22 deletion, namely E2F6 ACTIVITY UP or USP22 ACTIVITY UP (the downregulated genes upon E2F6 or USP22 de[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
MDM4 affects USP22
2 | 11 6
MDM4 binds USP22.
2 | 8 6
2 | 8 6

sparser
"Our data suggest that MDMX directly binds to USP22 in NSCLC cells."

reach
"Our co-immunoprecipitation analysis shows that USP22 interacts with MDMX in NSCLC cells."

reach
"MDMX Directly Binds to USP22 in NSCLC Cells."

sparser
"These data indicate that MDMX directly binds to Flag-tagged USP22 in A549-USP22 cells."

sparser
"Furthermore, we confirmed that MDMX directly interacts with endogenous USP22 in NCl-H460 cells, as MDMX was detected in USP22 immunoprecipitates and vice versa ( xref C,D)."

reach
"To investigate the interaction between MDMX and USP22, co-immunoprecipitation analysis was performed in NSCLC cell lines."

reach
"Furthermore, we confirmed that MDMX directly interacts with endogenous USP22 in NCl-H460 cells, as MDMX was detected in USP22 immunoprecipitates and vice versa (XREF_FIG C, D)."

reach
"Our data suggest that MDMX directly binds to USP22 in NSCLC cells."

sparser
"Our co-immunoprecipitation analysis shows that USP22 interacts with MDMX in NSCLC cells."

reach
"USP22 interacts with MDMX and leading to the inhibition of p53 dependent apoptosis [29]."
MDM4 activates USP22.
| 3
MDM4 activates USP22. 3 / 3
| 3

reach
"These results demonstrate that MDMX mediates USP22 's regulation effect in the NSCLC cell line."

reach
"MDMX Mediates USP22 's Regulation Effect in NSCLC Cells."

reach
"Importantly, overexpression of MDMX reversed USP22 silencing, induced p53 activation, growth inhibition, cell cycle arrest and apoptosis in A549 cells (XREF_FIG B-E)."
KDM1A affects USP22
5 | 7 5
KDM1A binds USP22.
5 | 5 5
5 | 5 5

sparser
"Moreover, GSK3β knockdown in GSC11 cells attenuated the interaction between endogenous USP22 and KDM1A ( xref ), which was confirmed in 293T cells by transfection of GSK3β-KD as compared with GSK3β-CA ( xref )."

sparser
"At the same time, the loss of OTUD7B promotes the binding of LSD1 to P62, a polyubiquitin chain binding protein, leading to LSD1 proteasomal degradation. xref USP28 (ubiquitin-specific protease 28) catalyzes deubiquitination of LSD1 through the N-terminal region of USP28 and the AOL domain of LSD1 to stabilize LSD1 protein, which enables direct regulation of the expression of differentiation genes, indirectly regulating the expression of the pluripotency activators. xref In glioma, a lack of methionine inhibits the growth of cancer cells and enhances the binding of LSD1 to E3 ubiquitin ligase CBL (casitas B-lineage lymphoma), and increased ubiquitination of LSD1 enhances expression of CXCL8 (CXC motif chemokine ligand 8) and methylated histone H3, which in turn affects the metabolism of the glycerophospholipid. xref USP7 (ubiquitin-specific protease 7) is able to interact with LSD1, exercising deubiquitination functions and enhancing the inhibitory effect of LSD1 on p53 (tumor protein p53). xref , xref , xref Phosphorylation of LSD1 catalyzed by GSK3β promotes the binding of LSD1 to USP22 (ubiquitin specific protease 22), and deubiquitination of LSD1 promotes tumor formation. xref , xref USP38 (ubiquitin-specific protease 38) interacts with LSD1 at amino acid 454 to stabilize protein level."

reach
"Using a series of KDM1A deletion mutants, we found that KDM1A 's AOL domain was required for KDM1A 's binding with USP22 (XREF_FIG)."

reach
"Moreover, GSK3beta knockdown in GSC11 cells attenuated the interaction between endogenous USP22 and KDM1A (XREF_FIG), which was confirmed in 293T cells by transfection of GSK3beta-KD as compared with GSK3beta-CA (XREF_SUPPLEMENTARY)."

reach
"Furthermore, KDM1A S683A significantly decreased the interaction between USP22 and KDM1A in the presence of GSK3beta-CA (XREF_FIG)."

sparser
"The target relationship of USP22 and miR-362-3p as well as the interaction of USP22 and LSD1 in RB was verified."

reach
"The target relationship of USP22 and miR-362-3p as well as the interaction of USP22 and LSD1 in RB was verified."

sparser
"Likewise, glycogen synthase kinase-3β (GSK-3β) phosphorylates LSD1 at S683 and stabilizes LSD1 via increasing the binding of ubiquitin specific peptidase 22 with LSD1 [ 36 ]."

reach
"40 In glioma, a lack of methionine inhibits the growth of cancer cells and enhances the binding of LSD1 to E3 ubiquitin ligase CBL (casitas B-lineage lymphoma), and increased ubiquitination of LSD1 enhances expression of CXCL8 (CXC motif chemokine ligand 8) and methylated histone H3, which in turn affects the metabolism of the glycerophospholipid.41 USP7 (ubiquitin-specific protease 7) is able to interact with LSD1, exercising deubiquitination functions and enhancing the inhibitory effect of LSD1 on p53 (tumor protein p53).42, 43, 44 Phosphorylation of LSD1 catalyzed by GSK3β promotes the binding of LSD1 to USP22 (ubiquitin specific protease 22), and deubiquitination of LSD1 promotes tumor formation.29 45 USP38 (ubiquitin-specific protease 38) interacts with LSD1 at amino acid 454 to stabilize protein level."

sparser
"Using a series of KDM1A deletion mutants, we found that KDM1A’s AOL domain was required for KDM1A’s binding with USP22 ( xref )."
KDM1A activates USP22.
| 2
KDM1A activates USP22. 2 / 2
| 2

reach
"14 However, the Lnc-ZFAS1-mediated mechanism in osteosarcoma still remains insufficient.Recently, the dysregulation of miRNAs has been verified closely correlated with tumors growth and metastasis,15 and accumulating studies have proved the crucial role of miRNAs in osteosarcoma, like miR-411-5p,2 miR-53916 and miR-4660.17 Liu W et al. suggested miR-140 could weak the protein stabilization of LSD1 mediated by targeting USP22 and facilitate the expression of p21 to block osteosarcoma progression.18 Interestingly, Wang J et al. have reported the suppressive influence of miR-520b on growth and metastasis in osteosarcoma."

reach
"As a result, the GSK3β-dependent phosphorylation of LSD1 could induce the combination of USP22 and LSD1, thus leading to the deubiquitylation of LSD1 and enhancing its stability."
USP22 affects Histone
| 18
USP22 deubiquitinates Histone-H2A. 9 / 9
| 9

reach
"Several DUBs have been implicated in histone deubiquitination, including USP3, USP12, USP22, and USP46, which deubiquitinate both histones H2A and histones H2B [XREF_BIBR]."

reach
"USP22 was initially reported to promote deubiquitylation of histones H2A and H2B, leading to transcription activation."

reach
"USP22 induces changes in gene promoter regions by deubiquitinating histones H2A and H2B, thereby controlling transcription."

reach
"Previous studies have demonstrated that USP22 deubiquitinates histones H2A and H2B and regulates cell growth, cell-cycle regulation and signal transduction (4,5,17)."

reach
"Apart from deubiquitinating histones H2A and H2B, USP22 also modulates transcription through stabilization of the p53-antagonizing deacetylase sirtuin-1 (SIRT1) to suppress p53-controlled transcription and to abrogate pre-mature senescence in pluripotent progenitor cells during embryonic development [5]."

reach
"In addition, USP22 can also enhance DNA damage repair by deubiquitinating histones H2A and H2Bub1."

reach
"USP22 is one subunit of SAGA (Spt-Ada-Gcn5-acetyltransferase), and is involved in the deubiquitination of histones H2A and H2B and the acetylation of histone H4 in order to regulate the transcription and expression of numerous genes (15,16)."

reach
"As we discussed previously, USP22 is a component of a transcriptional activator complex SAGA and can deubiquitinate histones H2A and H2B, as well as several other substrates (Zhang et al., 2008a, b)."

reach
"As a subunit of hSAGA, USP22 participates in the deubiquitination of histones H2A and H2B and the acetylation of histone H4 to regulate gene transcription and expression."
USP22 deubiquitinates Histone. 9 / 9
| 9

reach
"USP22 deubiquitylates histone and non histone substrates and has been associated with cancer progression and spinocerebellar ataxia."

reach
"USP22 deubiquitinates histone H2Bub and H2Aub."

reach
"USP22 can deubiquitylate both histone and non‐histone substrates."

reach
"USP22 reverses the polycomb-catalyzed ubiquitination of histones, including H2A and H2B, as well as nonhistone protein TRF1 (TBP [TATA box-binding protein]-related factor 1) to regulate the transcript[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition to histones, USP22 deubiquitinates TRF1, CCNB1, CCND1, and SIRT1, thereby regulating involvement in metabolism, cycling, and apoptosis."

reach
"Our studies indicate that USP22 acting as a histone deubiquitinase participates in deubiquitination of histone H2Bub on the promoter of VEGFA, co-activating ZEB1-mediated VEGFA transcription.It has been reported that ubiquitination modification on ZEB1 involved in maintenance of ZEB1 stability plays important roles in tumor progression."

reach
"However, the mechanisms by which miR-30e-5p regulates NSCLC pathogenesis remain unclear.Ubiquitin-specific peptidase 22 (USP22) is part of a multiprotein complex and regulates gene transcription withi[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP22 also deubiquitinates non histone proteins, including telomeric repeat binding factor 1 (TRF1), sirtuin 1 (SIRT1), cyclin B1 and others, leading to protein stabilization by preventing proteasome mediated degradation."

reach
"In addition, certain non histone proteins such as sirtuin 1 (Sirt1) and fructose-bisphosphatase 1 (FBP1) could be deubiquitinated by USP22."
USP22 affects BIRC5
1 | 15 1
USP22 decreases the amount of BIRC5.
| 6
USP22 decreases the amount of BIRC5. 6 / 6
| 6

reach
"Moreover, USP22 siRNA decreased survivin expression together with upregulation of CDK inhibitor, p21 and downregulation of cyclinB."

reach
"Furthermore, USP22 small interfering (si)RNA decreased the expression of Cyclin B and Survivin, and increased the expression of p21 in HCC (7)."

reach
"It was reported that USP22 silence by small interfering RNA reduced the expression of survivin in oral squamous cell carcinoma ."

reach
"Furthermore, USP22 small interfering RNA inhibited cell growth and reduced the expression levels of Aurora-B, Survivin and Cyclin B, together with the upregulation of cyclin-dependent kinase inhibitor 1A (p21)."

reach
"Indeed, the present study identified that USP22 siRNA decreased Aurora-B and Survivin expression in OSCC cells."

reach
"Notably, USP22 siRNA reduced the protein levels of Aurora-B and Survivin in OSCC cells (Fig. 2B)."
USP22 activates BIRC5.
| 5
USP22 activates BIRC5. 5 / 5
| 5

reach
"USP22 promotes proliferation in renal cell carcinoma by stabilizing survivin."

reach
"USP22 also upregulates BMI-1 and may upregulate Survivin via BMI-1 overexpression (20)."

reach
"Consistently, our results uncovered that USP22 overexpression significantly increased nuclear survivin in CA1 after tGCI."

reach
"These data indicate that USP22 promotes the development of OSCC together with Aurora-B and Survivin."

reach
"These data indicate that USP22 promotes the development of OSCC together with Survivin and Aurora-B."
USP22 binds BIRC5.
1 | 2 1
1 | 2 1

reach
"In order to understand the role of USP22, the present study examined the association between USP22, Aurora-B, Survivin and Ki-67 in OSCC."

sparser
"Finally, it was confirmed that USP22 interacted with survivin and stabilized it by downregulating its ubiquitination."

reach
"The functional associations between USP22, Aurora-B and Survivin in OSCC were also investigated."
USP22 increases the amount of BIRC5.
| 2
USP22 increases the amount of BIRC5. 2 / 3
| 2

reach
"Subsequently, it was demonstrated that USP22 knockdown inhibited the growth of an RCC cell line ACHN and downregulated the protein level of survivin, accompanied by an increased level of cleaved-caspase-3."

reach
"Our previous research demonstrated that USP22 can positively regulate the expression of Survivin in HCC (7); however, it is unclear whether USP22 regulates the expression of Aurora-B or Survivin in OSCC."
FASN affects USP22
4 | 14
4 | 14

sparser
"These results suggest that H 2 O 2 induces cells apoptosis by repressing the USP22-FASN axis in p53 +/+ colorectal cancer cells."

sparser
"Depletion of the USP22-FASN axis inhibits cell growth and colorectal tumorigenesis."

sparser
"Our study demonstrates that overproduced ROS in colorectal cancer controls lipid synthesis by critically regulating the USP22-FASN axis through a p53-dependent manner."

sparser
"To determine the potential clinical relevance of the USP22-FASN axis, we next assessed the expression of USP22 and FASN proteins in serial sections of 108 human CRC specimens, and found that FASN expression levels were significantly correlated with USP22 levels (Fig. xref , r  = 0.6626, P  < 0.0001)."

sparser
"Our findings establish that the USP22-FASN axis plays an important role in regulating lipid accumulation and colorectal tumorigenesis, and targeting this axis may represent a promising therapeutic strategy for colorectal cancer."

sparser
"Because p53 mutation usually loss the transcriptional activity and responses to upstream signals differently [ xref , xref ], p53-mediated repression of the USP22-FASN axis will be abolished in p53-mutated cancers, resulting in FASN stabilization, lipid accumulation, and tumor growth."

sparser
"Thus, the USP22-FASN axis revealed in this study may be a critical mechanism for the maintenance of CSC properties and therapy resistance of human cancer, which deserves further investigation."

sparser
"Due to a high-frequency of p53 mutation and dysfunction in human cancer, activation of the USP22-FASN axis may represent a prevailing mechanism that drives tumorigenesis, indicating a promising strategy for the treatment of colorectal cancer."

sparser
"Our study demonstrates that ROS critically regulates lipid synthesis and tumorigenesis through the USP22-FASN axis in a p53-dependent manner, and targeting the USP22-FASN axis may represent a potential strategy for the treatment of colorectal cancer."

sparser
"We have demonstrated that the USP22-FASN axis is highly activated in CRC and is critical for lipid accumulation and tumorigenesis."
E2F6 affects USP22
3 | 5 10
E2F6 binds USP22.
3 | 3 10
3 | 3 10

reach
"Therefore, we validated the interaction of E2F6 and USP22; the results revealed that E2F6, a pocket protein-independent transcription repressor, was capable of interacting with USP22 ( Fig. 1 C–E), in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Importantly, we found that the function of the USP22-E2F6 axis in promoting tumor cell growth was fulfilled by a decrease in DUSP1 activities, wherein E2F6 could directly bind to the promoter region o[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Therefore, the USP22-E2F6 axis has a tumor-promoting capacity in HCC, with the function to downregulate DUSP1 activities."

sparser
"Therefore, we validated the interaction of E2F6 and USP22; the results revealed that E2F6, a pocket protein-independent transcription repressor, was capable of interacting with USP22 ( Fig. 1 C–E), in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Thus, herein we provide the evidence that DUSP1 inhibition by the USP22-E2F6 axis is the upstream event that activates AKT signaling and drives HCC growth."

reach
"These results indicate a functional significance of the interaction between USP22 and E2F6 in HCC."

sparser
"To explore the functional importance of USP22-E2F6 regulation in HCC, we examined the transcriptional profiles of Huh7 cells with or without USP22 or E2F6 silencing by RNA-seq."

sparser
"These results indicate a functional significance of the interaction between USP22 and E2F6 in HCC."

sparser
"Overall, these results suggest that USP22-E2F6 regulation is a clinically relevant event in HCC."

reach
"Moreover, it has been demonstrated that the deubiquitinating enzyme USP22 can directly interact with and stabilize E2F6 in HCC cells, which indicates the presence of corresponding E3 ubiquitin ligases of E2F6 forming a dynamic bidirectional regulatory system for ubiquitin modifications 47."
E2F6 activates USP22.
| 2
E2F6 activates USP22. 2 / 2
| 2

reach
"Indeed, silencing E2F6 in liver tumor cells profoundly reversed the effects of USP22 on cell growth ( Fig. 3 D and E)."

reach
"We then generated four signatures based on the transcriptional alterations caused by E2F6 or USP22 deletion, namely E2F6 ACTIVITY UP or USP22 ACTIVITY UP (the downregulated genes upon E2F6 or USP22 de[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 1 16

reach
"Accordingly, USP22 downregulation also attenuated cerebral I/R-induced oxidative stress, inflammation, and cell apoptosis in mice, thereby reducing nerve injury and neurological dysfunction [20]."

reach
"Thus, we infer that the possible mechanism is that in the early stage of inflammation, USP22 reduces the level of cellular inflammation through compensatory cytoplasmic translocation, and when the inflammatory stimulus is prolonged, USP22 is in a state of loss of compensation, its expression decreases markedly, and the cell undergoes a strong inflammation."

reach
"Overexpression of USP22 reversed LPS-induced cellular inflammation, attenuated decidualization, and inhibited ferroptosis."

reach
"USP22 in macrophages promotes LRR and PYD structural domain protein 3 (NLRP3) degradation and attenuates inflammation through autophagy-associated 5 homolog (ATG5)-mediated autophagy ."

reach
"In addition, USP22/SIRT1-regulated mitochondrial respiration can affect liver steatosis, and the silencing of USP22 in diabetic rats confers a similar protective effect against high glucose-induced ap[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Addition of USP22 reduces Ang II-induced cardiac inflammation and fibrosis."

reach
"A previous study showed that USP22 deficiency in macrophages significantly promotes alum-induced peritoneal inflammation ."

reach
"Overexpression of USP22 in macrophages inhibits foam cell formation, inflammation, and macrophage apoptosis."

reach
"The knockout of Usp22 increased inflammation associated symptoms after DSS treatment locally and systemically."
| 1 7

reach
"Silencing of USP22 suppressed ROS production and inflammation while inhibition of USP14 reduced the accumulation of oxidized proteins."

reach
"In addition, USP22 expression increased the inflammatory response."

eidos
"Accordingly , USP22 downregulation also attenuated cerebral I / R-induced oxidative stress , inflammation , and cell apoptosis in mice , thereby reducing nerve injury and neurological dysfunction [ 20 ] ."

reach
"Overexpression of USP22 ameliorates LPS-induced endometrial stromal cells inflammation and modulates cells decidualization by inhibiting ferroptosis."

reach
"USP22 contributes to the inflammatory response and apoptosis in podocytes through oxidative stress and secretion of inflammatory mediators, up-regulation of caspase-3, and an increase in the BAX/Bcl-2 ratio96."

reach
"Therefore, USP22/SIRT1 signaling pathway may contribute to liver inflammation in ALD."

reach
"Taken together, the findings of the present study have demonstrated for the first time that USP22 inhibition attenuates high glucose induced podocyte injuries and inflammation."

reach
"In the current study, we show that the overexpression of USP22 induced by treatment with FO greatly improved Ang II-induced cardiac dysfunction, heart hypertrophy, inflammation, and fibrosis (Figures 1–4)."
| 17
| 13

reach
"Since these pathways all converge in the same target gene, USP22 can be used as a new target in cancer treatment.USP22 may be a particularly attractive target for cancer immunotherapy, because USP22 c[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Usp22 and Usp21 deletion in T reg cells synergizes to enhance antitumor immunity."

reach
"USP22 also restricts antitumor immunity through potentiating regulatory T cell (T ) immune suppressive activity (16) and T adaptation in the tumor microenvironment (17)."

reach
"What more excited is that USP22 can inhibit the antitumor immunity, efficacy of PDL1 targeted immunotherapy by inhibiting the deubiquitinase of PDL1 (CD274) [XREF_BIBR], which is consistent with our results."

reach
"In addition, USP22 deubiquitinated CD274 leading to the stabilization of its expression, knockdown of USP22 enhanced the anticancer immunity of CD274 in liver cancer [ 31 ]."

reach
"The expression of USP22 in tumor cells suppresses anti-tumor immunity and confers resistance to immunotherapy (18)."

reach
"Based on these observations, the authors hypothesized that USP22 inhibition could simultaneously promote anti-tumor immunity and target tumors driven by USP22 upregulation and pathogenic H2Bub1 deplet[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Tumor cell-intrinsic USP22 suppresses antitumor immunity in pancreatic cancer."

reach
"Also, Gregory et al. confirmed an important role for USP22 in suppressing the neoantigen-specific immune response via stabilizing the ecto-5’-nucleotidase (NT5E) in BC [39]."

reach
"These findings are consistent with the idea that USP22 promotes the non T cell inflamed TME and suppresses antitumor immunity in PDA."
USP22 activates immune response.
| 4

reach
"Therefore, our results indicate that Rottlerin and Morusin exerts potent anti-tumor effect and enhances antitumor immunity mediated by USP22 and merits further development.In order to gain a deeper in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Inactivation of Usp22, Brd9, and Ccdc101 enhanced antitumor immunity but also led to spontaneous inflammation in some cases (96–98)."

reach
"Based on these observations, the authors hypothesized that USP22 inhibition could simultaneously promote anti-tumor immunity and target tumors driven by USP22 upregulation and pathogenic H2Bub1 deplet[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"KO of Usp22 destabilized Foxp3 and subsequently increased antitumor immunity in vivo."
USP22 affects TERF1
1 | 12 3
USP22 deubiquitinates TERF1.
1 | 7
USP22 deubiquitinates TERF1. 7 / 8
1 | 7

reach
"In addition to histones, USP22 deubiquitinates TRF1, CCNB1, CCND1, and SIRT1, thereby regulating involvement in metabolism, cycling, and apoptosis."

reach
"This dissociation leads to lowered USP22 activity, which in turn leads to increased ubiquitination and turnover of TRF1 (XREF_FIG)."

reach
"Given the known regulation of TRF1 by ubiquitination, we hypothesized that the SAGA complex might facilitate Usp22 dependent deubiquitination of TRF1 and that loss of Gcn5 might alter this activity by compromising complex integrity."

reach
"Empirical studies have demonstrated that USP22 can deubiquitylate nonhistone substrates as well (e.g., TRF1 and FBP) (14, 15)."

reach
"Additionally, Usp22 directly deubiquitinates TRF1 (TBP (TATA box binding protein)-related factor 1) to regulate the transcription of cell cycle and apoptosis genes [XREF_BIBR] and inhibits the transcriptional activity of p53 by deubiquitinating SIRT1 histone deacetylase [XREF_BIBR] and by regulating MDMX stability [XREF_BIBR]."

reach
"GCN5 and USP22 also protect telomeres from DNA damage response through the stabilization of a shelterin component called TRF1, and interestingly, this regulation is not transcriptional but involves USP22 mediated deubiquitination of TRF1 [XREF_BIBR]."

reach
"For example, USP22 promotes TRF1 deubiquitylation to enhance TRF1 protein stability and maintain telomere integrity."
USP22 binds TERF1.
| 3 3
| 3 3

reach
"USP22 interacts with TRF1 and is required for TRF1 stability."

sparser
"USP22 interacts with TRF1 and is required for TRF1 stability."

reach
"For example, USP22 binding to TRF1, a protein that regulates telomere length, induces TRF1 protein stability."

sparser
"USP22 could bind to TRF1 and regulate telomere length (Atanassov et al ., xref )."

reach
"USP22 could bind to TRF1 and regulate telomere length (Atanassov etal., 2009)."

sparser
"For example, USP22 binding to TRF1, a protein that regulates telomere length, induces TRF1 protein stability."
USP22 increases the amount of TERF1.
| 2
USP22 increases the amount of TERF1. 2 / 3
| 2

reach
"Depletion of USP22 decreases TRF1 protein levels whereas overexpression of wild-type USP22 increases TRF1 levels in a manner dependent upon its enzymatic activity ( Atanassov et al., 2009 )."

reach
"Depleting USP22 decreases TRF1 levels and enhances cell death by genotoxic insults ."
USP22 affects ENY2
6 | 2 9
6 | 2 1

reach
"Further, Pfam analysis of protein domain structures confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity (XREF_FIG and XREF_SUPPLEMENTARY)."

reach
"Streptavidin pull-down of SFB-tagged PCAF validated our AP-MS, as interactions between PCAF and the SAGA DUB components USP22, ENY2, and ATXN7L were observed (Fig. 4B)."

sparser
"The association of USP22 and ENY2 in the same TFTC/STAGA module would therefore suggest that two activities, histone deubiquitination and mRNA export, may also be coregulated within this complex."
| 8

sparser
"The SAGA DUB module is highly conserved both in subunit composition and structural organization ( xref ; xref ; xref ; xref ; xref ; xref ), The deubiquitinating enzyme USP22 (Ubp8 in yeast) closely associates with two adapter proteins, ATXN7L3 and ENY2 (Sgf11 and Sus1 in yeast, respectively) ( xref ; xref ; xref ), and a fourth protein, ATXN7 (Sgf73), anchors the DUB module to the larger SAGA complex."

sparser
"It has been reported that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator Spt-Ada-Gcn5-acetyltransferase (SAGA) complex to regulate global levels of H2B monoubiquitination, thereby promoting antibody class switch recombination by facilitating nonhomologous end joining ( xref ; xref ; xref )."

sparser
"Similarly, USP22, the human homolog of Ubp8, is bound to the STAGA complex through interactions with ATXN7L3 and ENY2, which are homologs of Sgf11 and Sus1, respectively ( xref )."

sparser
"Further, Pfam analysis of protein domain structures ( xref ) confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity ( xref and xref ) ( xref )."

sparser
"We show that the ubiquitin protease USP22 forms a subcomplex with ATXN7L3 (ySgf11 homolog) and ENY2 (ySus1 homolog)."

sparser
"These results together suggest that ATXN7L3, USP22, and ENY2 form a stable subcomplex and that USP22 and ENY2 are recruited into TFTC/STAGA by ATXN7L3 ( Figure 2 D)."

sparser
"Cotransfection and coimmunoprecipitation experiments revealed that the ATXN7L3 ZnF-Sgf11 domain alone is able to interact with both ENY2 and USP22 as efficiently as the full-length ATXN7L3 (see Figur[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Previous study has shown that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator SAGA complex to regulate global levels of H2B monoubiquitination ( xref )."
USP22 affects EGFR
1 | 16
USP22 activates EGFR.
| 8
USP22 activates EGFR. 8 / 8
| 8

reach
"We show here that overexpression of USP22 prevents EGFR down-regulation, while shRNA mediated silencing of USP22 enhances EGFR degradation."

reach
"Thus, stabilization and activation of EGFR by USP22 are likely to be important factors in the mechanism underlying the oncogenicity and drug-resistance in EGFR-mutant lung ADCs."

reach
"Our study suggests that USP22 antagonizes EGFR degradation and amplifies EGFR signaling activity to promote EGFR-TKIs resistance in EGFR mutated lung ADCs."

reach
"USP22 can protect epidermal growth factor (EGR) receptor (EGFR) on late endosomes from degradation by deubiquitination and enhance EGFR recirculation after EGF stimulation, thereby activating multiple[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Cancer cells express proteins such as sortilin, tribbles homolog 3 protein (TRIB3), and ubiquitin specific peptidase 22 (USP22), which suppress EGFR degradation via EGFR stabilization, and their knockdown inhibits cancer cells [39–41]."

reach
"Of note, more EGFR colocalized with Rab11 in the PC9 cells with overexpression of USP22 than the control cells, which confirms that USP22 enhances EGFR recycling."

reach
"However, our finding on prevention of endocytosis mediated EGFR degradation by USP22 reveals one vital aspect of USP22 's diverse functions."

reach
"Ubiquitin-specific protease 22 (USP22) has been shown to prevent ubiquitination-mediated EGFR degradation and activate multiple EGFR downstream signaling pathways, thus conferring resistance to EGFR-TKIs in mutant lung adenocarcinoma cells [169]."
USP22 deubiquitinates EGFR.
1 | 3
USP22 deubiquitinates EGFR. 3 / 4
1 | 3

reach
"Similarly, shRNA knockdown of USP22 resulted in accumulated Ubn-EGFR, which further confirmed that USP22 antagonizes EGFR ubiquitination."

reach
"In this model, late endosome localized USP22 deubiquitinates EGFR and impedes sorting of EGFR to the lysosome, thus sustaining the trafficking of EGFR to the plasma membrane (Figs. 6 and 9)."

reach
"Concurrently, USP22 deubiquitinates EGFR on late endosomes, enhancing its recycling and the sustained activation of various downstream signaling pathways upon EGF stimulation [191]."
USP22 inhibits EGFR.
| 3
USP22 inhibits EGFR. 3 / 3
| 3

reach
"We show here that overexpression of USP22 prevents EGFR down-regulation, while shRNA mediated silencing of USP22 enhances EGFR degradation."

reach
"Our current study demonstrates that USP22 promotes EGFR-TKIs resistance by preventing EGFR degradation in EGFR-mutant lung ADC."

reach
"Cancer cells express proteins such as sortilin, tribbles homolog 3 protein (TRIB3), and ubiquitin specific peptidase 22 (USP22), which suppress EGFR degradation via EGFR stabilization, and their knockdown inhibits cancer cells [39–41]."
USP22 increases the amount of EGFR.
| 2
USP22 increases the amount of EGFR. 2 / 2
| 2

reach
"These results indicate that USP22 upregulates EGFR protein levels via enhancing EGFR stability."

reach
"In lung adenocarcinoma, USP22 can induce EGFR-TKI resistance by regulating the endocytosis and transport of EGFR through deubiquitination modification, stabilising the intracellular EGFR protein levels and promoting the continuous activation of EGFR-dependent signalling pathways (112)."
STING1 affects USP22
1 | 16
1 | 16

sparser
"In line with this, USP22-induced ISG expression could be reversed as well in USP22-STING dKO HT-29 cells (Fig. xref )."

sparser
"Additionally, USP22-mediated increases in IFN-λ expression could also largely be reduced upon USP22-STING dKO, whereas expression of IFN-α and IFN-β remained largely unaffected (Fig. xref )."

sparser
"The differential response to ISD, but not poly(I:C), and the reversal of the IFN signature in USP22-STING dKO hIECs suggests an important role of USP22 in the control of STING-induced type III IFN signaling."

sparser
"E. Western blot 831 analysis of STING and USP22 expression levels in control-NHT, control-USP22 KO 832 #1 and #6, STING-NHT and STING-USP22 KO #1 and #6 double KO (dKO) HT-29 833 cells."
| DOI

sparser
"To investigate the significance of USP22 and the 326 resulting STING-mediated upregulation of type III IFN and ISG signaling for viral327 defense, the role of the USP22-STING axis was tested during SARSexpression of STING, compared to wild-type (WT) and NHT 332 CRISPR/Cas9 control Caco-2 cells (Figure 6A)."
| DOI

sparser
"Intriguingly, 351 USP22-STING dKO hIECs exhibit higher SARS-CoV-2 replication rates as well as the formation of more de novo infectious viral particles compared to USP22 inflammatory diseases and cancer 1,2 ."
| DOI

sparser
"To confirm the role of STING in USP22-induced type III IFN signaling, USP22-STING dKO HT-29 cells were generated (Fig. xref )."

sparser
"USP22-STING dKO cells exhibited strikingly reduced levels of basal and phosphorylated STAT1 protein compared to USP22 KO HT-29 cells (Fig. xref ), suggesting a STING-dependent rescue of the USP22-dependent IFN signature."

sparser
"F. Quantification of relative SARS-CoV-2 835 genome expression of SARS-CoV-2-infected control-NHT, control-USP22 KO #1 and 836 #6, STING-NHT and STING-USP22 KO #1 and #6 dKO HT-29 cells at 24 hpi."
| DOI

sparser
"In line with this, USP22-283 induced ISG expression could be reversed as well in USP22-STING dKO HT-29 cells 284 (Figure 4H)."
| DOI
ENY2 affects USP22
6 | 2 9
6 | 2 1

reach
"Further, Pfam analysis of protein domain structures confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity (XREF_FIG and XREF_SUPPLEMENTARY)."

reach
"Streptavidin pull-down of SFB-tagged PCAF validated our AP-MS, as interactions between PCAF and the SAGA DUB components USP22, ENY2, and ATXN7L were observed (Fig. 4B)."

sparser
"The association of USP22 and ENY2 in the same TFTC/STAGA module would therefore suggest that two activities, histone deubiquitination and mRNA export, may also be coregulated within this complex."
| 8

sparser
"The SAGA DUB module is highly conserved both in subunit composition and structural organization ( xref ; xref ; xref ; xref ; xref ; xref ), The deubiquitinating enzyme USP22 (Ubp8 in yeast) closely associates with two adapter proteins, ATXN7L3 and ENY2 (Sgf11 and Sus1 in yeast, respectively) ( xref ; xref ; xref ), and a fourth protein, ATXN7 (Sgf73), anchors the DUB module to the larger SAGA complex."

sparser
"It has been reported that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator Spt-Ada-Gcn5-acetyltransferase (SAGA) complex to regulate global levels of H2B monoubiquitination, thereby promoting antibody class switch recombination by facilitating nonhomologous end joining ( xref ; xref ; xref )."

sparser
"Similarly, USP22, the human homolog of Ubp8, is bound to the STAGA complex through interactions with ATXN7L3 and ENY2, which are homologs of Sgf11 and Sus1, respectively ( xref )."

sparser
"Further, Pfam analysis of protein domain structures ( xref ) confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity ( xref and xref ) ( xref )."

sparser
"We show that the ubiquitin protease USP22 forms a subcomplex with ATXN7L3 (ySgf11 homolog) and ENY2 (ySus1 homolog)."

sparser
"These results together suggest that ATXN7L3, USP22, and ENY2 form a stable subcomplex and that USP22 and ENY2 are recruited into TFTC/STAGA by ATXN7L3 ( Figure 2 D)."

sparser
"Cotransfection and coimmunoprecipitation experiments revealed that the ATXN7L3 ZnF-Sgf11 domain alone is able to interact with both ENY2 and USP22 as efficiently as the full-length ATXN7L3 (see Figur[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Previous study has shown that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator SAGA complex to regulate global levels of H2B monoubiquitination ( xref )."
CCNB1 affects USP22
4 | 5 7
4 | 5 7

reach
"Our studies suggest that SIRT1 and CCNB1 interact with USP22 through different regions."

sparser
"However, this report clearly demonstrates that USP22 binds and deubiquitinates cyclin B1, leading to its stabilization."

sparser
"USP22 binding to Cyclin B1, promotes cyclin B1 accumulation in the nucleus and inhibits its degradation [ xref ]."

sparser
"USP22 interacts with CCNB1."

sparser
"The interaction between endogenous USP22 and CCNB1 in human colon cancer cells was also detected, as anti-CCNB1-specific antibody but not normal mouse immunoglobulin G immunoprecipitated USP22 protein ( xref )."

sparser
"To further refine our understanding of the molecular interaction between USP22 and CCNB1, we generated truncated USP22 mutants ( xref )."

reach
"USP22 binding to Cyclin B1, promotes cyclin B1 accumulation in the nucleus and inhibits its degradation [XREF_BIBR]."

sparser
"In addition, as S237 is also within the portion of USP22 that interacts with CCNB1, it is possible that CDK1-mediated S237 phosphorylation enhances USP22 interaction with CCNB1."

reach
"USP22 interacts with CCNB1."

reach
"The interaction between endogenous USP22 and CCNB1 in human colon cancer cells was also detected, as anti-CCNB1-specific antibody but not normal mouse immunoglobulin G immunoprecipitated USP22 protein (XREF_FIG)."
End3 affects USP22
| 4 12
| 4 12

sparser
"SIRT1, a class III histone deacetylase, serves as an acetylation mediator within the USP22 and SAGA coactivator complex [ xref , xref ]."

sparser
"USP22 forms part of the SAGA complex and requires the participation of cofactors like ATXN7L3 and ENY2 to deubiquitinate H2B, while USP27X functions independently of SAGA but also needs ATXN7L3 and ENY2 to act on H2B [ xref ]."

sparser
"GCN5 has been shown to support the association of USP22 with the SAGA complex."

reach
"Although this shows USP22 can bind both PALB2 and SAGA it does not delineate whether these are two separate populations or whether USP22 can interact with all of these components including PALB2 at the same time."

sparser
"Like in Drosophila , USP22 associates with human SAGA, and is recruited to specific genes by activators such as the MYC oncoprotein for transcription [ xref ]."

sparser
"On the contrary, as a part of the SAGA complex, USP22, by deubiqitinating TRF1, restores its protein level and thereby maintains telomeric length."

sparser
"We successfully identified many protein complexes known to associate with glucose starvation and revealed connections with many other complexes, of which identified modulated Emerin complex 1 and USP22-SAGA complexes are experimentally validated by co-IP."

sparser
"SAGA and USP22 inducibly bind to the promoter region of ER stress response genes."

sparser
"GCN5 was found to be required for USP22 to properly associate with the SAGA complex and to be able to deubiquitinate TRF1 and prevent its turnover ( xref )."

sparser
"One is Emerin complex 1 which function to organize the nuclear membrane during cytokinesis xref , and the other is USP22-SAGA complex which function is a regulatory center for signaling, chromatin modification, DNA damage repair, and gene control xref ."
ZEB1 affects USP22
2 | 3 11
2 | 3 11

sparser
"In addition, HEK-293 cells were transfected with USP22 and ZEB1 expression plasmids as indicated and the interaction between USP22 and ZEB1 was conformed in Co-IP experiment (Supplementary Fig. S xref C)."

sparser
"The results showed that only the N terminal of USP22 directly binds to ZEB1 (Fig. xref ), while the C terminal of USP22 may indirectly bind to ZEB1."

sparser
"Through a co-IP assay, we found that USP22 can bind to ZEB1 mRNA and that knockdown of USP22 markedly facilitates the ubiquitination of ZEB1 and thus enhances its protein degradation."

sparser
"Taken together, these data demonstrated that USP22 physically associates with ZEB1 in HCC cells."

sparser
"Here, our results have demonstrated that USP22 interacts with ZEB1, and USP22/ZEB1 is recruited to the ZEB1-binding elements of VEGFA promoter region and USP22 decrease the accumulation of histone H2Bub."

sparser
"Co-IP was used to validate the interaction between USP22 and ZEB1."

sparser
"Having established that USP22 associates with ZEB1, and USP22/ZEB1 is recruited to the promoter of VEGFA in HCC cells, we thus turn to examine whether USP22 participates in maintenance of ZEB1 stability."

sparser
"Next, we employed a Co-IP assay and found that USP22 interacts with ZEB1 ( xref )."

reach
"In addition, HEK-293 cells were transfected with USP22 and ZEB1 expression plasmids as indicated and the interaction between USP22 and ZEB1 was conformed in Co-IP experiment (Supplementary Fig. S3C)."

sparser
"USP22 interacts with ZEB1, and USP22/ZEB1 is recruited to the ZEB1-binding elements of VEGFA promoter region in HCC cell lines."
USP22 affects end3
| 4 12
| 4 12

sparser
"SIRT1, a class III histone deacetylase, serves as an acetylation mediator within the USP22 and SAGA coactivator complex [ xref , xref ]."

sparser
"USP22 forms part of the SAGA complex and requires the participation of cofactors like ATXN7L3 and ENY2 to deubiquitinate H2B, while USP27X functions independently of SAGA but also needs ATXN7L3 and ENY2 to act on H2B [ xref ]."

sparser
"GCN5 has been shown to support the association of USP22 with the SAGA complex."

reach
"Although this shows USP22 can bind both PALB2 and SAGA it does not delineate whether these are two separate populations or whether USP22 can interact with all of these components including PALB2 at the same time."

sparser
"Like in Drosophila , USP22 associates with human SAGA, and is recruited to specific genes by activators such as the MYC oncoprotein for transcription [ xref ]."

sparser
"On the contrary, as a part of the SAGA complex, USP22, by deubiqitinating TRF1, restores its protein level and thereby maintains telomeric length."

sparser
"We successfully identified many protein complexes known to associate with glucose starvation and revealed connections with many other complexes, of which identified modulated Emerin complex 1 and USP22-SAGA complexes are experimentally validated by co-IP."

sparser
"SAGA and USP22 inducibly bind to the promoter region of ER stress response genes."

sparser
"GCN5 was found to be required for USP22 to properly associate with the SAGA complex and to be able to deubiquitinate TRF1 and prevent its turnover ( xref )."

sparser
"One is Emerin complex 1 which function to organize the nuclear membrane during cytokinesis xref , and the other is USP22-SAGA complex which function is a regulatory center for signaling, chromatin modification, DNA damage repair, and gene control xref ."
USP22 affects SOX2
| 14
USP22 decreases the amount of SOX2.
| 11
USP22 decreases the amount of SOX2. 10 / 11
| 11

reach
"USP22 is located directly on the Sox2 promoter and negatively regulates Sox2 transcription in ESCs [XREF_BIBR]."

reach
"DUBs that have been reported to control developmental processes through deubiquitylating H2B include USP44, which represses genes involved in lineage commitment during mESC maintenance [XREF_BIBR], and USP22, which specifically inhibits expression of the pluripotency factor SOX2 during hESC differentiation [XREF_BIBR]."

reach
"Specifically, USP22 occupies the Sox2 promoter and represses Sox2 transcription [105]."

reach
"USP22 is known to bind to the promoter region of Sox2 and negatively regulates Sox2 transcription in embryonic stem cells (ESCs) 47."

reach
"In the recent study by Sussman et al., USP22 was shown to repress the expression of the Sox2 locus that encodes one of the core transcriptional regulators of ES-cell pluripotency (Sussman et al., 2013[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In mouse embryonic stem cells (mESC), USP22 knockdown upregulates SOX2 expression and attenuates the induction of differentiation marker genes in embryoid body formation assays ."

reach
"Therefore, we attempt to induce the degradation of SOX2 by targeting its DUBs.However, despite that USP22 has been shown to repress the transcription of SOX2, the deubiquitinating enzymes for controll[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP22 has been found to be located directly on the Sox2 promoter and catalyzes deubiquitination of H2B and attenuates Sox2 transcription."

reach
"In mouse ES cells, Usp22 represses the transcription of the pluripotency factor Sox2, thereby promoting differentiation [XREF_BIBR]."

reach
"In addition, USP22 occupies the Sox2 promoter and hydrolyzes mono-ubiquitin from ubiquitinated H2B (uH2B), and blocks transcription of the Sox2 locus."
USP22 activates SOX2.
| 2
USP22 activates SOX2. 2 / 3
| 2

reach
"Our data revealed that knockdown of USP22 decreased the expression of stemness related genes Sox2 and CD133, redox related genes Nrf2 and Sirt1, and quiescence related genes PP2A and c-Myc in GSCs und[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"As presented in Fig. 2 A–B , knockdown of USP22 inhibited the expression of stemness related genes Sox2 and CD133, redox related genes Nrf2 and Sirt1, and quiescence related genes PP2A and c-Myc."
USP22 inhibits SOX2.
| 1
USP22 inhibits SOX2. 1 / 2
| 1

reach
"Moreover they found USP22 represses the SOX2 promoter in order to control the embryonic stem cell transition from self-renewal to differentiation [XREF_BIBR] Therefore, not only is SOX2 an essential stem cell marker but its suppression is mandatory for cellular differentiation."
USP22 affects PTGS2
4 | 7 3
USP22 binds PTGS2.
4 | 2 3
4 | 2 3

sparser
"To examine the potential interaction between USP22 and COX-2, A549 cells were transfected with vectors expressing USP22 and Myc-tagged COX-2 and cell lysates were immunoprecipitated against anti-Myc [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"To verify the direct interaction between USP22 and COX-2, recombinant USP22 was incubated with His tagged COX-2 or COX-1 and proteins were isolated by affinity chromatography."

reach
"As shown in Fig. 2 C, USP22 was pulled down together with COX-2 but not with COX-1, confirming the specificity of the interaction between USP22 and COX-2."

sparser
"To verify the direct interaction between USP22 and COX-2, recombinant USP22 was incubated with His-tagged COX-2 or COX-1 and proteins were isolated by affinity chromatography."

sparser
"As shown in C, USP22 was pulled down together with COX-2 but not with COX-1, confirming the specificity of the interaction between USP22 and COX-2."
USP22 activates PTGS2.
| 3
USP22 activates PTGS2. 3 / 4
| 3

reach
"Here, we identified COX-2 as a substrate of USP22 and investigated the role of USP22 in tumorigenesis via the modulation of COX-2 stability and activity.In the present study, we showed that silencing [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Silencing of USP22 downregulates COX-2 and thus inhibits cancer cell proliferation in OSCC by direct interaction leading to modulation of stability and activity of COX-2 through controlling of its ubiquitination status."

reach
"Our results showed that USP22 silencing downregulated COX-2 and FBP1 in A549 and NCI-H460 cells compared to control siRNA."
USP22 decreases the amount of PTGS2.
| 2
USP22 decreases the amount of PTGS2. 2 / 3
| 2

reach
"The results of western blot analysis showed that USP22R restored the levels of USP22 and COX-2 downregulated by USP22 siRNA."

reach
"Silencing of USP22 or COX-2 significantly decreased the production of PGE2, whereas overexpression of COX-2 restored PGE2 levels downregulated by USP22, confirming that USP22 modulates COX-2 expressio[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects NFATC2
2 1 | 10 3
USP22 deubiquitinates NFATC2.
1 | 7
USP22 deubiquitinates NFATC2. 7 / 8
1 | 7

reach
"In this study, we found that an ubiquitin-specific protease, USP22, interacts with and deubiquitinates NFATc2."

reach
"Consistent with previous results, overexpression of USP22 but not the USP22C185A mutant could reduce ubiquitination of NFATc2."

reach
"USP22 can promote the expression of interleukin-2 in T cells by deubiquitinating and stabilizing NFATc2 (an important regulator of T-cell activation)—a novel role for USP22 as a positive regulator of NFATc2 in the control of T-cell immune responses (25)."

reach
"NFATc2 is a regulatory molecule in T-cell activation, and USP22 promotes interleukin-2 expression in T cells by deubiquitinating and maintaining the stability of NFATc2."

reach
"Gao et al found that USP22 interacts with and deubiquitinates NFATc2, and also stabilizes NFATc2 protein and promotes NFATc2 function to facilitate IL-2 expression in T cells."

reach
"USP22 can deubiquitinate and stabilize NFATC2 to activate T cells and upregulate IL-2 release."

reach
"Thus, our data indicated that USP22 deubiquitinates NFATc2, which requires its enzymatic activity.Since we found that USP22 deubiquitinates NFATc2, we reasoned that USP22 might control NFATc2 stabilit[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 binds NFATC2.
2 | 3 3
2 | 3 3

sparser
"We then examined the interaction between USP22 and NFATc2."

reach
"We found that USP22 could physically interact with and stabilize NFATc2 by deubiquitination."

reach
"Gao et al found that USP22 interacts with and deubiquitinates NFATc2, and also stabilizes NFATc2 protein and promotes NFATc2 function to facilitate IL-2 expression in T cells."

reach
"We then examined the interaction between USP22 and NFATc2."

sparser
"Collectively these results indicated that USP22 can interact with NFATc2."

sparser
"Because the deubiquitinase USP22 interacts with NFATc2, we hypothesized that USP22 might facilitate NFATc2 deubiquitination."
USP22 affects Melanoma
| 1 15
USP22 activates Melanoma.
| 1 12
| 1 12

reach
"Moreover, we found that USP22 expression is relatively elevated in melanoma when compared with normal skin or tumor‐adjacent normal tissues in Xiangya dataset and two different GSE datasets (GSE15605 and GSE46517) (Figures 1F and S1B)."

eidos
"USP22 promotes melanoma mainly through YAP As a transcriptional activator , YAP exerts its oncogenic functions by promoting the transcription of downstream target genes , such as CTGF and Cyr61 ( 41 ) ."

reach
"Together, our results demonstrated the upregulation of USP22 in melanoma and its correlation with poor clinical outcomes.2.2 USP22 promotes melanoma metastasis both in vitro and in vivo."

reach
"These findings suggested that USP22 promotes melanoma metastasis both in vitro and in vivo.2.3 Elevated USP22 induces EMT activation in melanoma."

reach
"Based on our results, the ferroptosis inhibitor liproxstatin‐1 partially rescued the inhibitory effect of topotecan on metastasis, indicating that pharmacological inhibition of USP22 could partially suppress melanoma metastasis by enhancing ferroptosis sensitivities.Previously, Yi et al. 53 revealed that activating mutations in PI3K or loss of PTEN mediate the ferroptosis resistance of cancer cells."

reach
"Besides, USP22 was found to augment melanoma and resistance to BRAF inhibitors by stabilizing YAP [34] ."

reach
"These results demonstrated that USP22 might promotes melanoma metastasis by inducing EMT activation.2.4 USP22 potentiates melanoma metastasis and EMT through activating PI3K/Akt/mTOR pathway."

reach
"These results suggested that USP22 potentiates melanoma metastasis and EMT through activating the PI3K/Akt/mTOR pathway.2.5 USP22 activates PI3K/Akt/mTOR pathway via SIRT1/PTEN axis."

reach
"In mechanism, USP22 enhances the stability of STAT1 via deubiquitination and promotes the activation of IFNγ stimulation in melanoma [ 43 ]."

reach
"In this study, we demonstrate that the ubiquitin-specific peptidase 22 (USP22) is essential for HGF-induced melanoma metastasis."
USP22 inhibits Melanoma.
| 3
| 3

reach
"Ablation of USP22 expression remarkably attenuates melanoma migration, invasion, and epithelial-mesenchymal transition in vitro and suppresses melanoma metastasis in vivo."

reach
"We found that USP22 deficiency only enhances the vulnerability of melanoma to ferroptosis inducer RSL3 and has no sensitizing effect on other antimelanoma drugs or cell death inducers (Figures 7A and S7A), suggesting that USP22 has a specific sensitization effect on melanoma ferroptosis."

reach
"Pharmacological inhibition of USP22 by topotecan exerts a similar effect to USP22 knockout in inhibiting melanoma metastasis both in vivo and in vitro."
USP22 affects HYCC1
| 16
USP22 activates HYCC1. 10 / 16
| 16

reach
"These results suggest that the expression of USP22 may promote VM formation in HCC-derived cell lines.Next, we turn to determine the effect of USP22 on migratory and invasive behaviors of HCC cells, transwell assays were performed in shUSP22 Huh7 or shUSP22 PLC/PRF/5 cells."

reach
"Taken together, our results indicate that USP22 promotes HCC cell growth and VM formation in mice.Having revealed that USP22 maintains ZEB1 stability, we next examined the expression of USP22 and ZEB1 in 24 pairs of human HCC pathological sections by immunohistochemistry."

reach
"Our results revealed that USP22 promoted tumorigenesis and progression via an FKBP12/mTORC1/autophagy positive feedback loop in HCC."

reach
"Therefore, USP22 activates the mTORC1 signaling pathway both in vitro and in vivo.2.3 USP22 promotes tumorigenic potential and sensitizes HCC toward rapamycin in vitro and in vivo."

reach
"2.1 Overexpression of Usp22 accelerates c-Myc/NRasGV12-induced HCC in mice."

reach
"Under the treatment of Lenvatinib, USP22 knockdown inhibited the cell viability of drug-resistant HCC cells and promoted the apoptosis of drug-resistant cells."

reach
"Simply put, USP22 may activate the SIRT1–AKT–MRP1 pathway and consequently promote MDR in human HCC cells (226)."

reach
"Our data demonstrate that USP22 deletion inhibits the migration and invasion of HCC cells (Fig. 5F, G and Supplementary Fig. S6D)."

reach
"Ablation of USP22 in immunosuppressive regulatory T cells leads to reduced tumor burden in several cancers.56 Importantly, USP22 promotes tumorigenesis and progression in HCC, by promoting stemness.53,57As hepatic expression of CXCL13 and SCF genes strongly correlated with tumor burden, we tested their potential as circulating HCC biomarkers."

reach
"USP22 promotes HCC-derived cell growth/invasion/Vascular Mimicry (VM) formation and angiogenesis."
USP22 affects FlaG
| 16
| 16

sparser
"Consistent with our LacO array recruitment experiment, ( xref ) mCherry-PALB2 WD40 4X could not pull-down FLAG-USP22 WT while mCherry-PALB2 WD40 WT robustly bound FLAG-USP22 WT ."

sparser
"The fragment mCherry-PALB2 Nterm that does not contain the WD40 domain was not able to pull-down FLAG-USP22 WT validating our previous LacO array data ( xref )."

sparser
"Subsequently, after transfecting the cells with the USP22-FLAG and USP22–C185S-FLAG plasmids under the NG and HG conditions, we detected the ubiquitination level of Snail1."

sparser
"To determine whether USP22 could affect the development of DKD, we used a USP22 overexpression plasmid (USP22-FLAG) and a USP22 deubiquitinase activity mutant plasmid (USP22–C185S-FLAG) to transfect t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Moreover, the results of immunofluorescence experiments showed that transfected FLAG-USP22 (FITC, green) and endogenous ZEB1(Cy3, red) were distributed in the nucleus in HCC-derived cells or HEK293 cells (Fig. xref and Supplementary Fig. S xref D)."

sparser
"WB results showed that USP22 expression significantly increased in NRK-52E cells transfected with USP22-FLAG and USP22–C185S-FLAG ( Fig. 3 A), which verified the success of USP22 overexpression."

sparser
"Therefore, we detected the ubiquitination level of Sirt1 to verify the enzyme activity of USP22 plasmids in NRK-52E cells transfected with USP22-FLAG and USP22–C185S-FLAG."

sparser
"As shown in the data in Fig. 3 B, USP22-FLAG rather than USP22–C185S-FLAG could reduce the Sirt1 ubiquitination level, indicating that the USP22 plasmids functioned normally in the EMT model."

sparser
"Furthermore, overexpression of siRNA resistant FLAG-USP22 in U2OS FOKI cells rescues the localization of BRCA2, PALB2, and Rad51 to the array after DNA damage."

sparser
"In HG-treated NRK-52E cells, USP22-FLAG overexpression further increased USP22 expression ( Fig. 3 C), downregulated E-cadherin and ZO-1expression, and upregulated Vimentin expression ( Fig. 3 D)."
USP22 affects FOXP3
| 16
USP22 activates FOXP3.
| 5
USP22 activates FOXP3. 5 / 5
| 5

reach
"Recent studies demonstrate that USP22 is a promising target for cancer immunotherapy, since USP22 positively regulates FOXP3 activity in mouse Treg cells that suppress antitumor immune responses, and Treg-specific USP22-knockout (USP22-Ko) suppressed in vivo Treg function to improve antitumor immunity [31]."

reach
"USP22 positively regulated transcription factor FOXP3 activity in mouse regulatory T (T reg) cells."

reach
"Our findings that hypoxia induced Usp22 in a HIF-dependent manner, and loss of Usp22 resulted in diminished Foxp3 stability, imply that hypoxia can stabilize T cells through Usp22-mediated Foxp3 stabilization."

reach
"USP22 positively regulated transcription factor FOXP3 activity in mouse regulatory T (T reg) cells."

reach
"T cells from the dKO mice displayed a GSEA and bulk gene expression signature that merged the changes found in each of the single KO mice, suggesting that the loss of both Usp22 and Usp21 synergizes to diminish T cell function (Fig. 4G) in both Foxp3-depedent and Foxp3-independent manner.As we demonstrated that both Usp22 and Usp21 are regulated by metabolic alterations in the TME, it was particularly interesting to identify disruption of multiple metabolic pathways in each of the KO groups."
USP22 increases the amount of FOXP3.
| 4
USP22 increases the amount of FOXP3. 4 / 4
| 4

reach
"Loss of Usp22 in Treg reduces Foxp3 transcript levels, increases FOXP3 ubiquitination and degradation, and reduces suppressive activity in vivo in mice."

reach
"Together with our recent discovery that Usp22 functions as a Foxp3-specific deubiquitinase, our data suggest that Usp22 promotes Foxp3 expression in itT cells (14)."

reach
"Targeted knockdown of USP22 in Treg cells using clustered regularly interspaced short palindromic repeats (CRISPR) technology reduces Foxp3 protein expression and inhibits tumor growth in multiple tum[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Hypoxia selectively induces T reg Usp22, which supports Foxp3 expression."
USP22 decreases the amount of FOXP3.
| 4
USP22 decreases the amount of FOXP3. 4 / 4
| 4

reach
"T reg -specific ablation of Usp22 in mice reduced Foxp3 protein levels and caused defects in their suppressive function that led to spontaneous autoimmunity but protected against tumour growth in multiple cancer models."

reach
"Screen hits were confirmed using individual ribonucleoprotein sgRNA-Cas9 complexes (RNPs), which showed that targeting Usp22 in both mouse and human Tregs (USP22) decreased FOXP3 expression, again suggesting a role in positively regulating FOXP3 expression."

reach
"Therefore, these results indicate that Usp22i-S02 is a potent Usp22-specific small-molecule inhibitor that down-regulates Foxp3 expression in T cells in a Usp22- but not Usp21-dependent manner."

reach
"Deletion of Usp22 drives reduction in Foxp3 transcript levels and through increased ubiquitin at the chromatin level ultimately leading to Foxp3 degradation."
USP22 deubiquitinates FOXP3.
| 3
USP22 deubiquitinates FOXP3. 3 / 3
| 3

reach
"The results revealed that USP22 can promote FoxP3 deubiquitination and stabilization, which is counteracted by the E3 ligase ring finger protein 20 (Rnf20)."

reach
"Similarly, USP21, USP22, and USP44 also deubiquitinate Foxp3, preventing its degradation (195–199)."

reach
"Loss of Usp22 in Treg reduces Foxp3 transcript levels, increases FOXP3 ubiquitination and degradation, and reduces suppressive activity in vivo in mice."
FlaG affects USP22
| 16
| 16

sparser
"Consistent with our LacO array recruitment experiment, ( xref ) mCherry-PALB2 WD40 4X could not pull-down FLAG-USP22 WT while mCherry-PALB2 WD40 WT robustly bound FLAG-USP22 WT ."

sparser
"The fragment mCherry-PALB2 Nterm that does not contain the WD40 domain was not able to pull-down FLAG-USP22 WT validating our previous LacO array data ( xref )."

sparser
"Subsequently, after transfecting the cells with the USP22-FLAG and USP22–C185S-FLAG plasmids under the NG and HG conditions, we detected the ubiquitination level of Snail1."

sparser
"To determine whether USP22 could affect the development of DKD, we used a USP22 overexpression plasmid (USP22-FLAG) and a USP22 deubiquitinase activity mutant plasmid (USP22–C185S-FLAG) to transfect t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Moreover, the results of immunofluorescence experiments showed that transfected FLAG-USP22 (FITC, green) and endogenous ZEB1(Cy3, red) were distributed in the nucleus in HCC-derived cells or HEK293 cells (Fig. xref and Supplementary Fig. S xref D)."

sparser
"WB results showed that USP22 expression significantly increased in NRK-52E cells transfected with USP22-FLAG and USP22–C185S-FLAG ( Fig. 3 A), which verified the success of USP22 overexpression."

sparser
"Therefore, we detected the ubiquitination level of Sirt1 to verify the enzyme activity of USP22 plasmids in NRK-52E cells transfected with USP22-FLAG and USP22–C185S-FLAG."

sparser
"As shown in the data in Fig. 3 B, USP22-FLAG rather than USP22–C185S-FLAG could reduce the Sirt1 ubiquitination level, indicating that the USP22 plasmids functioned normally in the EMT model."

sparser
"Furthermore, overexpression of siRNA resistant FLAG-USP22 in U2OS FOKI cells rescues the localization of BRCA2, PALB2, and Rad51 to the array after DNA damage."

sparser
"In HG-treated NRK-52E cells, USP22-FLAG overexpression further increased USP22 expression ( Fig. 3 C), downregulated E-cadherin and ZO-1expression, and upregulated Vimentin expression ( Fig. 3 D)."
COPS5 affects USP22
| 6 10
| 6 10

sparser
"Moreover, USP22 also interacted with CSN5 and stabilized CSN5 through deubiquitination."

sparser
"USP22 also interacted with CSN5 and kept its stability via deubiquitination."

reach
"Moreover, USP22 also interacted with CSN5 and stabilized CSN5 through deubiquitination."

reach
"USP22, a ubiquitin carboxyl-terminal hydrolase, interacted with JAB1 and stabilized JAB1 through deubiquitination (Wang et al. 2020b)."

sparser
"On the other hand, USP22 interacts with CSN5 and deubiquitinates it, thereby facilitating the interaction between PD-L1 and CSN5 ( xref )."

sparser
"USP22 interacted with CSN5 and deubiquitinated CSN5."

sparser
"Additionally, USP22 forms a complex with CSN5, maintaining its stability via deubiquitination."

sparser
"Since CSN5 bound to PD-L1 [ xref ], we wondered if USP22 interacted with CSN5 physically."

reach
"Additionally, USP22 forms a complex with CSN5, maintaining its stability via deubiquitination."

sparser
"As expected, USP22 interacted with CSN5 exogenously and endogenously (Fig.  xref a, Fig. xref b)."
ATXN7 affects USP22
10 | 4
ATXN7 binds USP22.
10 | 1
10 | 1

reach
"Western blot analysis of complexes obtained after an ATXN7 immunopurification showed that USP22 and ATXN7L3 associate with ATXN7 together with other components of TFTC and STAGA."
ATXN7 inhibits USP22.
| 3
ATXN7 inhibits USP22. 3 / 4
| 3

reach
"This indicated that poly (Q) Ataxin-7 likely decreased USP22 activity."

reach
"Recently, Lan et al. reported that poly (Q) Ataxin-7 does not directly decrease USP22 enzymatic activity, but instead, it forms aggregates that impair USP22 binding to its substrates [XREF_BIBR]."

reach
"For example, polyQ ataxin-7 sequesters USP22 into aggregates and inhibits its DUB activity."
AKT affects USP22
| 10 6
AKT binds USP22.
| 1 4
| 1 4

sparser
"The interaction of AKT with USP22 was detected in transiently transfected HEK293T cells as well as in human MDA-MB-231 breast cancer cells ( xref )."

sparser
"Stimulation of MDA-MB-231 breast cancer cells with EPI markedly enhanced both AKT and USP22 interaction ( xref ) as well as their phosphorylation (fig."

sparser
"Conversely, treatment with the AKT-specific inhibitor facilitated USP22 protein degradation ( xref ), increased the USP22 ubiquitination ( xref ), and inhibited the AKT and USP22 interaction in breast cancer cells ( xref )."

reach
"Conversely, treatment with the AKT-specific inhibitor facilitated USP22 protein degradation (Fig. 5, K and L), increased the USP22 ubiquitination (Fig. 5M), and inhibited the AKT and USP22 interaction in breast cancer cells (Fig. 5N)."

sparser
"Of note, mass spectrometry (MS) analysis also detected 10 AKT phosphorylated peptides in the anti-USP22 immunoprecipitants ( xref ), confirming our initial discovery of AKT-USP22 interaction and suggesting that USP22 interacts with the phosphorylated AKT upon EPI stimulation in breast cancer cells."
AKT inhibits USP22.
| 3 1
AKT inhibits USP22. 4 / 4
| 3 1

reach
"These results suggest that USP22 knockdown induced chemosensitivity of HCC cells by down-regualting PI3K and Akt, and Smad4 mediated Akt suppression as well."

reach
"Inhibition of the PI3K/Akt pathway could rescue the effects of USP22 on cell migration."

reach
"USP22 knockdown enhanced chemosensitivity of hepatocellular carcinoma cells to 5-Fu by up-regulation of Smad4 and suppression of Akt."

sparser
"Silencing Smad4 blocked USP22 knockdown-induced Akt inhibition in Bel/Fu cells."
AKT activates USP22.
| 3
AKT activates USP22. 3 / 3
| 3

reach
"In addition to USP22, many other downstream factors, such as breast cancer resistance protein of AKT, are involved in drug resistance of breast cancers (56); therefore, future works are needed to dissect each substrate of phosphatidylinositol 3-kinase–AKT pathways in breast cancer tumorigenesis.One unexpected observation is the EPI-induced posttranscriptional stabilization of USP22, the cancer stem cell gene with increased expression in breast cancer stem cells (18)."

reach
"In addition, the deubiquitinase-inactive mutation completely abolished AKT-mediated USP22 stabilization."

reach
"These findings suggest that SOS1/RAS and downstream ERK and PI3K/AKT pathways mediate the oncogenic role of USP22 in gastric cancer."
AKT phosphorylates USP22.
| 1 1
AKT phosphorylates USP22. 2 / 2
| 1 1

sparser
"As shown in xref , USP22 peptides carrying each of the five phosphorylated amino acid residues including T50, T147, T168, T173, and S355 were markedly enriched in cells when AKT was overexpressed ( xref ), indicating that AKT phosphorylates USP22 through multiple sites."

reach
"As shown in Fig. 6 (A and B), USP22 peptides carrying each of the five phosphorylated amino acid residues including T50, T147, T168, T173, and S355 were markedly enriched in cells when AKT was overexpressed (Fig. 6, A and B), indicating that AKT phosphorylates USP22 through multiple sites."
AKT deubiquitinates USP22.
| 2
AKT leads to the deubiquitination of USP22. 2 / 2
| 2

reach
"Loss of AKT-mediated phosphorylation markedly increased USP22 ubiquitination and abolished EPI-induced inhibitory efficacy (Fig. 6D)."

reach
"As a consequence, overexpression of AKT markedly inhibited USP22 ubiquitination and prolonged its half-life (Fig. 5, H to J)."
| 15

reach
"USP22 silencing suppresses in vitro GC cell migration and invasiveness."

reach
"These results suggest that while USP22 promotes cell migration and invasion, loss of USP22 sensitizes EGFR mutant NSCLC cells to erlotinib in vitro."

reach
"Our results also demonstrated that USP22 can increase cell migration and invasion abilities via EMT induction."

reach
"These results suggest that USP22 promotes in vitro GC cell migration and invasion."

reach
"Indeed, compared with control, we found that the USP22 knockdown cells grew significantly more slowly in H1650 (p < 0.05, Fig. 2 B) and the USP22 overexpression in A549 accelerated cell growth (p < 0.[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"The results showed cancer cell migration was significantly inhibited by knockdown of USP22 in H1650 (p < 0.05, Fig. 2 C)."

reach
"Meanwhile, overexpression of USP22 in A549 markedly promoted the cell migration when compared with untreated A549 controls (p < 0.05, Fig. 2 C)."

reach
"Cell-scratch test results showed that in HG-treated NRK-52E cells, USP22-FLAG overexpression significantly increased cell migration, whereas USP22–C185S-FLAG overexpression had no such function ( Fig.[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP22 increases CRC cell migration and invasion by inducing EMT."

reach
"Taken together, these results indicate that USP22 increases cell migration and invasion by inducing EMT by binding to the promoter of AP4 to activate its transcription."
USP22 affects YAP1
2 1 | 11 1
USP22 binds YAP1.
2 | 4 1
2 | 4 1

sparser
"Using IP assays, USP22 interaction with YAP at the endogenous level in A375 melanoma cells was observed ( xref )."

reach
"In melanoma, USP22 interacts with and deubiquitinates YAP, thereby favoring melanoma cell proliferation."

reach
"The interaction between USP22 and KDM3A and between KDM3A and YAP1 was further validated."

reach
"USP22 interacts with and deubiquitinates YAP to prevent YAP turnover [26]."

reach
"A recent study demonstrated that ubiquitin-specific peptidase 22 (USP22) interacted with and stabilized YAP, which in turn conferred resistance to the BRAF inhibitor vemurafenib and provided new therapeutic avenues to target USP22/YAP as an option for melanoma treatment [224]."
USP22 activates YAP1.
| 5
USP22 activates YAP1. 5 / 5
| 5

reach
"USP22 promotes GC progression by modulating FOXO1 or YAP signaling pathways via c-Myc/NAMPT/SIRT1."

reach
"Besides, USP22 was found to augment melanoma and resistance to BRAF inhibitors by stabilizing YAP [34] ."

reach
"Western blot analysis of control and USP22-silenced GC cells showed that USP22 modulates the c-Myc/NAMPT/SIRT1-dependent FOXO1 and YAP signaling pathways."

reach
"USP22 maintained GC cell stemness by stabilizing BMI1 and promoted proliferation and metastasis by activating the FOXO1 and YAP signaling pathway [47,50]."

reach
"Oncogenic USP22 supports gastric cancer growth and metastasis by activating c-Myc/NAMPT/SIRT1-dependent FOXO1 and YAP signaling."
USP22 deubiquitinates YAP1.
1 | 2
USP22 deubiquitinates YAP1. 2 / 3
1 | 2

reach
"USP22 interacts with and deubiquitinates YAP to prevent YAP turnover [26]."

reach
"In melanoma, USP22 interacts with and deubiquitinates YAP, thereby favoring melanoma cell proliferation."
USP22 affects Ubiquitin
| 15
USP22 inhibits Ubiquitin.
| 8
| 8

reach
"HULC is overexpressed in HCC as it promotes growth in cancer cells through enhancing ubiquitin specific peptidase 22 (USP22) which reduces ubiquitin mediated degradation of COX-2 protein hence stabilizing and upregulating COX-2 protein [XREF_BIBR]."

reach
"Moreover, the study of the corresponding mechanism by which HULC upregulated COX-2 showed that HULC enhanced the level of ubiquitin specific peptidase 22 (USP22), which decreased ubiquitin mediated degradation of COX-2 protein by removing the conjugated polyubiquitin chains from COX-2 and finally stabilized COX2 protein."

reach
"Coexpression of USP22 significantly inhibited ubiquitin-conjugation to Sirt1 ( Figure 2 A)."

reach
"It was also indicated that lncRNA HULC could up-regulate expression of ubiquitin-specific peptidase 22 (USP22) and reduce the ubiquitin-mediated degradation of Sirt1 protein by removing the conjugated polyubiquitin chain of Sirt1 [135]."

reach
"In liver cancer, upregulation of lncRNA HULC activates autophagy by increasing the expression of ubiquitin specific peptidase 22 (USP22) which in turn prevents the ubiquitin mediated degradation of silent information regulator 1 (SIRT1) by removing the conjugated polyubiquitin chains from SIRT1."

reach
"On the contrary, the deubiqutinase USP22 antagonizes the ubiquitin ligase complex RNF20/40 on H2B monoubiquitination."

reach
"It belongs to the DUB subset of Machado–Joseph disease (MJD) proteic domain-containing peptidases with Atxn3 (a.k.a. MJD1 and SCA3), encoded by the gene mutated in MJD, also termed type-3 spinocerebellar ataxia (SCA3), and Josephin domain-containing DUbs JosD1 and JosD2 (Table 1.7).27Aggregates formed by polyglutamine-expanded ataxin-7 sequester ubiquitin-specific peptidase USP22 that cannot then fulfill its deubiquitinating function in the SAGA complex, causing cytotoxicity and neurodegeneration [109]."
| PMC

reach
"As expected, USP22 overexpression significantly inhibited ubiquitin conjugation to c-Myc ( Figure 5 E)."
USP22 activates Ubiquitin.
| 7
| 7

reach
"Stability, USP22 can promote PD-L1 to remove the ubiquitin chain and prevent it from being degraded by the proteasome [13–16] ."

reach
"As suspected from its domain structure, USP22 is able to hydrolyze a ubiquitin linkage from histone H2B in vitro and endogenous USP22 contributes ubiquitin hydrolase activity to the hSAGA complex."

reach
"USP22 inhibits ubiquitin-mediated proteasomal degradation of SPI1."

reach
"Sevoflurane postconditioning upregulated USP22, which increased the protein content of lysine-specific demethylase 3A (KDM3a) in the promoter region of YAP1 by cutting the ubiquitin chains of the protein."

reach
"Interestingly, the human and fly SAGA complexes possess a module that houses ubiquitin hydrolase activity mediated by the USP22 (human) and Nonstop (fly) proteins."

reach
"37 It can be upregulated along with the poor prognosis by USP22 (ubiquitin‐specific peptidase 22) expression through MAPK1 pathway."

reach
"Another study also showed that USP22 interacted with PD-L1, enhancing its stability by removing ubiquitin and preventing its breakdown by the proteasome."

reach
"USP22 enhances atherosclerotic plaque stability and macrophage efferocytosis by stabilizing PPARgamma."

reach
"We unexpectedly found that USP22 in macrophages enhanced efferocytosis both in vitro and in vivo."

reach
"In this study, we likewise found that silencing USP22 promotes the uptake of oxidized lipoproteins, which may inhibit macrophage efferocytosis.Interestingly, in this study, we found that USP22 promoted the expression of efferocytosis-associated receptors as well as bridging molecules and that this regulation is dependent on the activation of PPARγ."

reach
"In addition, to determine the mechanism by which silencing USP22 inhibits efferocytosis, we used CytD to inhibit actin polymerization in THP-1-derived macrophages, which prevented the internalization of ACs."

reach
"We subsequently used adeno-associated virus overexpressing USP22 in macrophages to investigate its biological properties and found that USP22 could promote efferocytosis and stabilize plaques by regulating PPARγ."

reach
"We next investigated whether USP22 enhances efferocytosis by regulating PPARγ."

reach
"Finally, we tested whether the decrease in PPARγ might be one of the molecular mechanisms by which USP22 silencing inhibits efferocytosis."

reach
"Our results showed that the PPARγ agonist rosiglitazone reversed the decrease in macrophage efferocytosis caused by USP22 silencing (Fig. 5H–J, Supplementary Fig. S5A, B. Furthermore, for THP-1-derived macrophages under inflammatory conditions, USP22 overexpression also promoted efferocytosis."

reach
"In addition, we cannot completely exclude the possible role of USP22 in other cell types in atherosclerosis in this study.In conclusion, we showed that USP22 in macrophages promotes efferocytosis and alleviates the progression of atherosclerosis in ApoE mice by regulating PPARγ."

reach
"We subsequently determined whether GW9662 influenced the promotion of efferocytosis by USP22 overexpression in macrophages in vivo."
USP22 affects PPARG
4 | 3 7
USP22 binds PPARG.
4 | 1 7
4 | 1 7

sparser
"For instance, the USP22-PPARγ axis plays a significant role in modulating both adipogenesis and inflammation, whereas disorders in bile acid metabolism and alterations in gut microbiota exhibit bidirectional regulation. xref In light of these mechanisms, preclinical studies have led to the development of small-molecule inhibitors that target USP22 and FXR."

sparser
"In vitro pulldown assays with purified recombinant proteins demonstrated that USP22 directly bound to PPARγ (Fig.  xref )."

sparser
"We subsequently performed coimmunoprecipitation experiments to confirm the interaction between USP22 and PPARγ (Fig.  xref ), and predicted their possible binding sites via HDOCK software (Fig.  xref )."

sparser
"Then, the specific interaction between USP22 and PPARγ was confirmed by co-immunoprecipitation assay with exogenously transduced USP22 and PPARγ in HEK293T cells (Fig.  xref )."

sparser
"Overall, our results confirm that USP22 specifically interacts with PPARγ, which is a key transcription factor related to lipid metabolism."

sparser
"Interestingly, we found that USP22 strongly bound to the PPARγ DBD domain (Fig.  xref ) as well as pVHL and CUL4B proteins in the HCC cells (Supplementary Fig.  xref ), and the interaction between PPARγ and USP22 significantly decreased the pVHL and CRL4B AhR involved ubiquitination (Fig.  xref ), indicating that USP22 regulates deubiquitination of PPARγ through other lysine sites."

sparser
"Taken together, these results indicate that USP22 directly interacts with PPARγ and functions as a bona fide PPARγ deubiquitinase in cells."

reach
"Mechanistically, USP22 bound to peroxisome proliferator-activated receptor gamma (PPARgamma) and inhibited its ubiquitination, thereby stabilizing PPARgamma and promoting efferocytosis."
USP22 activates PPARG.
| 2
USP22 activates PPARG. 2 / 2
| 2

reach
"USP22 enhances atherosclerotic plaque stability and macrophage efferocytosis by stabilizing PPARgamma."

reach
"USP22 promotes lipogenesis in Tregs through stabilization of peroxisome proliferator-activated receptor gamma (PPARγ) [51] ."
| 1 12
USP22 activates Osteosarcoma.
| 1 8
| 1 8

reach
"In addition, deficiency of ALKBH5‐mediated m6A modification in osteosarcoma causes an increase expression of histone deubiquitinase USP22 and ubiquitin ligase RNF40, which inhibits histone H2AK199 monoubiquitination, induces the expression of genes related to DNA repair, and promotes the progression of osteosarcoma."

reach
"In addition, the expression of USP22 promotes the proliferation of osteosarcoma cells in a glycolytic dependent manner both in vitro and in vivo, while the knockout of USP22 is the opposite."

reach
"We also confirmed that USP22 enhanced viability and motility of osteosarcoma cells."

reach
"The detailed mechanism of USP22-mediated osteosarcoma is still elusive.In the present study, we investigated the expression and biological functions of linc00265 in osteosarcoma cells."

reach
"The molecular mechanisms by which USP22 promotes osteosarcoma involve multiple signaling pathways, including glycolysis, oxidative phosphorylation, Spliceosome, Thermogenesis, and Cell cycle."

eidos
"Significance : RNA demethylase ALKBH5 upregulates USP22 and RNF40 to inhibit histone H2A ubiquitination and induces expression of key replication and DNA repair-associated genes , driving osteosarcoma progression ."

reach
"Silencing of either USP22 or RNF40 reduced short-term cell viability and clonogenic growth of osteosarcoma cells (Fig. 6A and B)."

reach
"USP22 Promotes Osteosarcoma Progression by Stabilising β-Catenin and Upregulating HK2 and Glycolysis."

reach
"CCK-8 data showed that USP22 overexpression accelerated viability of osteosarcoma cells, which was rescued by miR-485-5p upregulation ( Figure 6A )."
USP22 inhibits Osteosarcoma.
| 4
| 4

reach
"Downregulation of USP22 can inhibit the glycolysis and growth of osteosarcoma cells."

reach
"Furthermore, ALKBH5-mediated m6A deficiency increases the expression of USP22 and RNF40 in osteosarcomas, promoting osteosarcoma cell growth and proliferation [40]."

reach
"Western blotting data further validated that miR-485-5p overexpression reduced the expression of USP22 protein, whereas miR-485-5p downregulation increased the USP22 protein levels in osteosarcoma cells ( Figure 5D )."

reach
"In addition, deficiency of ALKBH5‐mediated m6A modification in osteosarcoma causes an increase expression of histone deubiquitinase USP22 and ubiquitin ligase RNF40, which inhibits histone H2AK199 monoubiquitination, induces the expression of genes related to DNA repair, and promotes the progression of osteosarcoma."
USP22 affects FOXO1
| 9 5
USP22 binds FOXO1.
| 2 5
| 2 5

sparser
"The endogenous USP22 interaction with FOXO1 was further confirmed in MDA-MB-231 cells ( xref )."

sparser
"Further analysis of FOXO1 interactions with USP22 truncated mutants revealed that the zinc finger–containing N terminus of USP22 mediates its interaction with FOXO1 ( xref )."

sparser
"Stimulation with EPI markedly increased USP22 interaction with FOXO1 in breast cancer cells ( xref )."

sparser
"EPI stimulation, which induced USP22 protein expression and promoted USP22-FOXO1 interaction, markedly inhibited FOXO1 polyubiquitination in USP22 WT cancer cells, and targeted USP22 deletion resulted in a significant increase in FOXO1 ubiquitination and reduced FOXO1 protein expression even with EPI stimulation ( xref )."

reach
"The endogenous USP22 interaction with FOXO1 was further confirmed in MDA-MB-231 cells (Fig. 4L)."

reach
"Further analysis of FOXO1 interactions with USP22 truncated mutants revealed that the zinc finger–containing N terminus of USP22 mediates its interaction with FOXO1 (Fig. 4K)."

sparser
"To define USP22 functions as a possible FOXO1-specific deubiquitinating enzyme, we first determined if USP22 interacts with FOXO1."
USP22 activates FOXO1.
| 7
USP22 activates FOXO1. 7 / 7
| 7

reach
"Oncogenic USP22 supports gastric cancer growth and metastasis by activating c-Myc/NAMPT/SIRT1-dependent FOXO1 and YAP signaling."

reach
"USP22 promotes GC progression by modulating FOXO1 or YAP signaling pathways via c-Myc/NAMPT/SIRT1."

reach
"Further IHC analysis confirmed that EPI treatment markedly increased the protein expression of USP22, FOXO1, and ATGL in xenograft MDA-MB-231 breast cancer tissues, while USP22 inhibition largely suppressed EPI-induced up-regulation of both FOXO1 and ATGL (Fig. 4, F to I)."

reach
"Western blot analysis of control and USP22-silenced GC cells showed that USP22 modulates the c-Myc/NAMPT/SIRT1-dependent FOXO1 and YAP signaling pathways."

reach
"Therefore, our data suggest that EPI promotes USP22-mediated FOXO1 protein stabilization for ATGL up-regulation in breast cancer cells."

reach
"USP22 maintained GC cell stemness by stabilizing BMI1 and promoted proliferation and metastasis by activating the FOXO1 and YAP signaling pathway [47,50]."

reach
"Collectively, our study revealed a previously unappreciated molecular mechanism underlying how USP22 controls EPI-induced cancer progression and metastasis through potentiating FOXO1-medited ATGL expression and lipolysis."

reach
"USP22 promotes the proliferation of hepatocellular carcinoma cells by stabilising CDK11B and enhances resistance to Sorafenib by inhibiting ferroptosis through the USP22/H2BK120ub/TFRC axis.So far, more than 40 USPs have been directly or indirectly associated with relevant cancer processes."

reach
"In contrast, other studies have shown that USP22 promotes fatty acid synthesis in hepatocellular carcinoma cells and, thus, hepatocellular carcinoma progression ."

reach
"After we had confirmed that USP22 promotes the growth of hepatocellular carcinoma, we further focused on exploring whether USP22 regulates Sorafenib resistance in hepatocellular carcinoma."

reach
"To further validate that USP22 increases hepatocellular carcinoma cell proliferation by stabilising CDK11B protein, we overexpressed His‐CDK11B in USP22 knockout cells (Figure 4F) and evaluated cell proliferation."

reach
"Our data further indicates that USP22 promotes the proliferation of hepatocellular carcinoma cells via deubiquitinating and stabilizing cyclin-dependent kinase 11B (CDK11B)."

reach
"Why USP22 has a preference for H2BK120ub as its substrate under Sorafenib treatment is worth further research.In summary, our study demonstrated that USP22 promotes the growth of hepatocellular carcinoma and resistance to Sorafenib‐induced ferroptosis by removing ubiquitin moieties from non‐histone protein CDK11B and histone H2B."

reach
"Altogether, these results provided evidence that USP22 promotes the proliferation of hepatocellular carcinoma cells by stabilising CKD11B and increases Sorafenib resistance through reducing H2BK120ub level and TFRC transcription in vivo.3 DISCUSSION."

reach
"For example, USP22 can stabilize the E2F6 stability and activate Akt pathway in hepatocellular carcinoma (HCC), leading to aggressive progression of HCC (25)."

reach
"19 However, there has been a lack of thorough investigation into the role of USP22 in hepatocellular carcinoma development and Sorafenib resistance.In this study, we demonstrate that USP22 promotes the proliferation of hepatocellular carcinoma cells by stabilising cyclin‐dependent kinase 11B (CDK11B)."

reach
"USP22 promotes MDR in hepatocellular carcinoma cells partly by activating the SIRT1/Akt/MRP1 pathway [ 17 ]."
TFPI affects USP22
| 9 5
TFPI phosphorylates USP22.
| 3 4
TFPI leads to the phosphorylation of USP22. 7 / 7
| 3 4

sparser
"Treatment of mice with the AKT inhibitor completely abolished EPI-induced USP22 phosphorylation in 4T1 tumors (fig."

reach
"These results indicate that EPI stabilizes USP22 through AKT-mediated USP22 phosphorylation."

reach
"These results indicate that EPI induces USP22 phosphorylation both in vitro and in tumor tissues in an AKT-dependent manner.We then analyzed the phospho-USP22 (p-USP22) levels by IHC staining using specific antibodies to one of the USP22 phosphorylation sites, T147, that is catalyzed by AKT (Fig. 6, A and B)."

sparser
"These results indicate that EPI induces USP22 phosphorylation both in vitro and in tumor tissues in an AKT-dependent manner."

sparser
"Further treatment with the AKT-specific inhibitor largely diminished EPI-induced USP22 phosphorylation ( xref ), indicating that AKT is a kinase for USP22 phosphorylation in breast cancer cells upon EPI stimulation."

reach
"Consistent with our conclusion that EPI induces USP22 phosphorylation through AKT activation, we observed a statistically substantial increase in p-USP22 levels in breast cancers from EPI versus EPI groups (Fig. 6, J and K)."

sparser
"Consistent with our conclusion that EPI induces USP22 phosphorylation through AKT activation, we observed a statistically substantial increase in p-USP22 levels in breast cancers from EPI high versus EPI low groups ( xref )."
TFPI increases the amount of USP22.
| 6 1
TFPI increases the amount of USP22. 7 / 7
| 6 1

reach
"These observations suggest the possibility that EPI induces USP22 protein expression posttranslationally in human breast cancers."

reach
"Further IHC analysis confirmed that EPI treatment markedly increased the protein expression of USP22, FOXO1, and ATGL in xenograft MDA-MB-231 breast cancer tissues, while USP22 inhibition largely suppressed EPI-induced up-regulation of both FOXO1 and ATGL (Fig. 4, F to I)."

reach
"EPI stimulation, which induced USP22 protein expression and promoted USP22-FOXO1 interaction, markedly inhibited FOXO1 polyubiquitination in USP22 WT cancer cells, and targeted USP22 deletion resulted in a significant increase in FOXO1 ubiquitination and reduced FOXO1 protein expression even with EPI stimulation (Fig. 4P)."

reach
"Because EPI induces USP22 protein expression without affecting its mRNA levels, we tested if EPI-mediated signaling controls USP22 protein expression at posttranslational levels."

reach
"These findings suggest a potential mechanism whereby EPI may contribute to tumorigenesis, at least in part, through the up-regulation of USP22.We sought to investigate if EPI stimulates USP22 expression in breast cancer cells."

sparser
"Similarly, EPI-induced USP22, FOXO1, and ATGL protein expression was also observed in 4T1 syngeneic breast cancer tissues by IHC staining."

reach
"Intriguingly, the induction of USP22 protein expression by EPI reached a plateau as early as 2 hours after treatment (Fig. 1G)."
FOXM1 affects USP22
| 5 9
| 5 9

sparser
"These results indicate that FoxM1 physically interacts with USP22 in breast cancer cells."

sparser
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected with Myc-USP22 and Flag-FoxM1 but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone ( xref C)."

sparser
"Our data indicate physical interaction between USP22 and FoxM1 is required for USP22-mediated suppression of FoxM1 ubiquitination, because mutation of cystines 61 and 63, which disrupts the zinc finger structure and its interaction with FoxM1 ( xref ), totally abolished USP22 activity in suppressing FoxM1 ubiquitination ( xref )."

sparser
"The endogenous interaction between USP22 and FoxM1 was further validated in patient-derived breast cancer L2G + TN1 and 4T1 cells ( xref D and xref B)."

reach
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected Myc-USP22 and Flag-FoxM1, but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone (Fig. 3C)."

reach
"The endogenous interaction between USP22 and FoxM1 in patient-derived breast cancer L2G TN1 cells was further validated (Fig. 3D and s3B)."

sparser
"Truncated mutation analysis revealed that the zinc finger-containing N-terminus is sufficient for USP22 interaction with FoxM1, while the C-terminus ubiquitin-specific peptidase domain is not involved in mediating its FoxM1 interaction ( xref F)."

reach
"Our data indicate physical interaction between USP22 and FoxM1 is required for USP22-mediated suppression of FoxM1 ubiquitination, because mutation of cystines 61 and 63, which disrupts the zinc finger structure and its interaction with FoxM1 (Fig. 3I), totally abolished USP22 activity in suppressing FoxM1 ubiquitination (Fig. 3G)."

sparser
"These results indicate that FoxM1 physically interacts with USP22 in breast cancer cells."

reach
"Indeed, USP22 interaction with FoxM1 was detected in HEK-293T cells transiently transfected with Myc-USP22 and Flag-FoxM1 but not in control cells transfected with Flag-FoxM1 or Myc-USP22 alone (Figure 3C)."
| 7

reach
"Research has shown that USP22 can enhance myeloid differentiation in Kras mice by stabilizing the PU.1 protein and decreasing the risk of AML [14]."

reach
"On the contrary, Usp22 overexpression leads to increased differentiation of ESCs into EBs even in the absence of stimuli that drives differentiation [XREF_BIBR]."

reach
"USP22 deficiency in Ras-driven myeloproliferative neoplasm blocks myeloid differentiation promoting acute myeloid leukemia (37)."

reach
"Knockdown of Usp22 shortened the half-life of Hes1, delayed its oscillation, and enhanced neuronal differentiation in mouse developing brain, whereas mis expression of Usp27x reduced neuronal differentiation."

reach
"In addition, Sussman RT, et al. have reported that USP22 promotes embryonic stem cell differentiation through transcriptional repression of Sex determining region Y-box 2 (Sox2) XREF_BIBR."

reach
"USP22 modulated embryonic stem cell differentiation through acting as a transcription repressor of SOX2 , which contributes to the transition from self-renewal to differentiation in embryonic stem cel[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Additionally, the oncogenic PML-RARα fusion protein is also stabilized in an USP22- and K394-mediated manner, through which USP22 defines sensitivity towards ATRA-mediated APL differentiation."
| 6

reach
"Another study in mice showed that Usp22 negatively regulates neuronal differentiation in the mouse developing brain."

reach
"On the contrary, depletion of USP22 delays Hes1 oscillation and thereby, induces neuronal differentiation from neuronal progenitor stem cells ."

reach
"For instance, two histone directed DUBs, USP44 and USP22, are antagonistically regulated in their mRNA expression to ensure faithful stem cell differentiation [XREF_BIBR - XREF_BIBR]."

reach
"Moreover, USP22 depletion promotes the differentiation of NSCs, both in vitro and in vivo."

reach
"In contrast, USP22 overexpression inhibits NSC differentiation into neurons."

reach
"Interestingly, our data showed that USP22 promotes the proliferation but inhibits the differentiation of NSCs in the dentate gyrus (DG)of the hippocampus soon after TBI."
USP22 affects FBP1
1 | 12
USP22 deubiquitinates FBP1.
1 | 5
USP22 deubiquitinates FBP1. 5 / 6
1 | 5

reach
"A previous study demonstrated that USP22 could inhibit the transcription of the cyclin-dependent kinase inhibitor 1A (p21) gene by deubiquitinating the transcriptional regulator fructose-1,6-bisphosphatase (FBP1) (5)."

reach
"Furthermore, USP22 regulates cell proliferation by deubiquitinating the transcriptional regulator, FBP1 (5)."

reach
"18 USP22 also induced deubiquitination of FBP1, thereby promoting the progression of non-small cell lung cancer."

reach
"Empirical studies have demonstrated that USP22 can deubiquitylate nonhistone substrates as well (e.g., TRF1 and FBP) (14, 15)."

reach
"Moreover, USP22 knockdown upregulates the transcription of p21 by upregulating the ubiquitylation of FBP1."
USP22 ubiquitinates FBP1.
| 4
USP22 ubiquitinates FBP1. 4 / 4
| 4

reach
"Ablation of USP22 does not affect the protein stability of FBP1 but increases ubiquitination of FBP1 and decreases FBP1 occupancy at the p21 gene [13] ."

reach
"Recent studies have demonstrated that USP22 can inhibit the transcription of the p21 gene by de-ubiquitinating the transcriptional regulator FBP1, leading to cell proliferation and tumorigenesis [XREF_BIBR]."

reach
"A previous study demonstrated that USP22 could inhibit the transcription of the cyclin-dependent kinase inhibitor 1A (p21) gene by deubiquitinating the transcriptional regulator fructose-1,6-bisphosphatase (FBP1) (5)."

reach
"USP22 also suppresses CDKN1A and p21 -locus expression by deubiquitinating its transcriptional repressor FBP1 (Atanassov and Dent, 2011), and acts as a negative regulator of the tumor suppressor p53 b[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 activates FBP1.
| 3
USP22 activates FBP1. 3 / 3
| 3

reach
"Moreover, it has been shown that USP22 promotes cell growth by regulating the far upstream element (FUSE)-binding protein 1 (FBP1), a transcriptional regulator of p21 [XREF_BIBR]."

reach
"Hence USP22 silencing reduces the capacity of FBP1 to repress p21 (independently of TP53 status), which in turn inhibits CDKs to prevent the G1/S transition and resulting in G1 accumulation [XREF_BIBR]."

reach
"Our results showed that USP22 silencing downregulated COX-2 and FBP1 in A549 and NCI-H460 cells compared to control siRNA."
USP22 affects CTNNB1
| 11 2
USP22 binds CTNNB1.
| 2 2
| 2 2

sparser
"Next, we examined the interaction between USP22 and β-catenin within the nucleus after tGCI with or without hypoxia by co-immunoprecipitation assay."

sparser
"The results showed that the interaction between USP22 and β-catenin in the nucleus was significantly weakened at 50 h after tGCI."

reach
"Next, we examined the interaction between USP22 and β-catenin within the nucleus after tGCI with or without hypoxia by co-immunoprecipitation assay."

reach
"The results showed that the interaction between USP22 and β-catenin in the nucleus was significantly weakened at 50 h after tGCI."
USP22 methylates CTNNB1.
| 3
USP22 leads to the methylation of CTNNB1. 3 / 3
| 3

reach
"In addition, considering that the ubiquitination of β-catenin could be reduced by the inhibition of demethylase activity of KDM2A and the methylation of β-catenin increased by USP22, we speculate that there may be a competition between the methylation and ubiquitination modification of β-catenin."

reach
"Further, the upregulation of nuclear USP22 induced by HPC promotes the methylation of nuclear β-catenin after USP22-mediated deubiquitination, which enhances the stability of nuclear β-catenin in CA1 and protects ischemic neuron against cerebral ischemia."

reach
"Our data revealed that nuclear β-catenin deubiquitination induced by USP22 overexpression significantly increased the methylation of nuclear β-catenin after tGCI."
USP22 deubiquitinates CTNNB1.
| 3
USP22 leads to the deubiquitination of CTNNB1. 3 / 3
| 3

reach
"In contrast, the overexpression of USP22 promoted nuclear beta-catenin deubiquitination and enhanced the neuroprotective effects offered by HPC."

reach
"Further, the upregulation of nuclear USP22 induced by HPC promotes the methylation of nuclear β-catenin after USP22-mediated deubiquitination, which enhances the stability of nuclear β-catenin in CA1 and protects ischemic neuron against cerebral ischemia."

reach
"Our data revealed that nuclear β-catenin deubiquitination induced by USP22 overexpression significantly increased the methylation of nuclear β-catenin after tGCI."
USP22 activates CTNNB1.
| 3
USP22 activates CTNNB1. 3 / 3
| 3

reach
"By up- and downregulation of USP22 expression, we also proved that USP22 can activate the Wnt and beta-Catenin pathway, which in turn affected the proliferation and migration of HepG2 cells."

reach
"USP22 was found to promote β-catenin nuclear localization, thus increasing the activity of the Wnt/β-catenin pathway in cancer cells [41] ."

reach
"In addition, USP22 can also activate Wnt/beta-catenin signaling axis to promote stemness and chemoresistance [ 67 ] (see Fig. 2 )."

reach
"Owing to its relationship with the chemotherapeutic resistance of several types of human cancers (Glinsky, 2005), cyclin B1 may also mediate the USP22 induced MDR in HCC cells, which is valuable for future exploration."

reach
"Together, these results indicate that USP22 could promote the MDR in HCC cells by activating the SIRT1/AKT/MRP1 pathway."

reach
"First, we defined SIRT1 as a significant mediator in USP22 driven MDR in HCC."

reach
"Simply put, USP22 may activate the SIRT1–AKT–MRP1 pathway and consequently promote MDR in human HCC cells (226)."

reach
"These results confirmed the role of SIRT1 in USP22 induced MDR in HCC cells."

reach
"USP22 promotes MDR in hepatocellular carcinoma cells partly by activating the SIRT1/Akt/MRP1 pathway [ 17 ]."

reach
"XREF_BIBR Ubiquitin specific protease 22 (USP22) mediates the MDR of HCC via the SIRT1/AKT/MRP1 signaling pathway."

reach
"Interestingly, we found that modulation of USP22 or SIRT1 could influence the intracellular ADR concentration, which might partly bridge the induction of MDR by USP22 and SIRT1 in HCC cells."

eidos
"] Our previous work revealed that USP22 was able to promote HCC stemness by a HIF1alpha / USP22 positive feedback loop and mediate multidrug resistance ( MDR ) by activating the SIRT1 / AKT / MRP1 pathway ."

reach
"Taken together, the present study found that USP22 can promote the MDR in HCC cells via activating the SIRT1/AKT/MRP1 pathway."
USP22 increases the amount of diphenylmethane-4,4'-diisocyanate.
| 3
| 3

reach
"USP22 is also found to indirectly promote the expression of MDR-related genes through upregulation of AKT and subsequently activation of the PI3K pathway in HCC [69]."

reach
"Moreover, USP22 knockdown also decreases MDR related genes expression by inhibiting Akt phosphorylation, and USP22 knockdown mediated Smad4 up-regulation is crucial for Akt suppression."

reach
"USP22 knockdown decreased MDR related genes expression through up-regulation of Smad4 and suppression of Akt."
SPI1 affects USP22
3 | 5 5
3 | 5 5

sparser
"In line with the truncated mutant of USP22, the C-terminal ubiquitin-specific peptidase domain of USP22 containing a point mutation could bind SPI1 weakly, whereas the N-terminal zinc finger domain of USP22 containing a point mutation could bind SPI1 more strongly (Fig.  xref )."

sparser
"These results indicate that USP22 interacts with SPI1 and enhances SPI1 stability through deubiquitination."

reach
"Co-IP assays were then performed to verify the interaction between USP22 and SPI1."

reach
"The C-terminal ubiquitin-specific peptidase domain of USP22 binds SPI1 strongly, whereas the N-terminal zinc finger domain binds SPI1 weakly (Fig. 5d)."

reach
"In line with the truncated mutant of USP22, the C-terminal ubiquitin-specific peptidase domain of USP22 containing a point mutation could bind SPI1 weakly, whereas the N-terminal zinc finger domain of USP22 containing a point mutation could bind SPI1 more strongly (Fig. 5g)."

reach
"Thus, the C-terminal ubiquitin-specific peptidase domain of USP22 is the dominant regulator of binding between USP22 and SPI1."

reach
"Co-IP assays were performed to determine the interaction between USP22 and SPI1."

sparser
"USP22 interacts with SPI1."

sparser
"Co-IP assays were then performed to verify the interaction between USP22 and SPI1."

sparser
"To identify the region within USP22 interacting with SPI1, we generated two truncated mutants of USP22 in HEK-293T cells."
AR affects USP22
2 | 6 5
2 | 6 3

reach
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Cotransfection in HEK293T cells and coimmunoprecipitation experiments revealed that AR interacted with ATXN7L3 in a ligand-dependent manner, similarly to GCN5, while interaction between AR and USP22 w[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Subsequent immunoprecipitation also confirmed the association between AR and USP22 in vitro, which is consistent with the result of previous studies (Fig. 3I)."

sparser
"As AR and USP22 interacted in vivo, we next tested whether AR could be a substrate of the TFTC/STAGA deubiquitination activity."

sparser
"No significant association of AR and USP22 with chromatin at an intergenic region could be detected, and the amount of histone H3 at KLK2 promoter remained constant, further demonstrating specificit[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, our immunofluorescence analysis showed that Importin-7 binds to AR or USP22 in both the nucleus and cytoplasm (Fig. 4C–E), suggesting that Importin-7 may regulate the nuclear translocation of AR and USP22.Next, we conducted a more in-depth study of the Importin-7–ARUSP22 complex by the treatment of AR inhibitor (enzalutamide) or AR ligands, including DHT and synthetic androgen R1881."

reach
"Here, to further investigate whether the arrest of the nuclear translocation of ARUSP22 complex triggered by the knockdown of Importin-7 could enhance the sensitivity of BC to enzalutamide.We first confirmed that the knockdown of Importin-7 could enhance the growth inhibition of cancer cells by enzalutamide using CCK8 assay (Fig. 6A, B) and EdU staining assay (Fig. 6C, D)."

reach
"Interestingly, we found that USP22, one of the maintainers of AR [22–24], whose nuclear translocation is also regulated by Importin-7, although knockdown of Importin-7 may not affect the formation of the ARUSP22 complex.In addition, we also observed that AR expression levels were upregulated after DHT and R1881 stimulation, and Importin-7-bound AR was also up-regulated to an almost equal extent, indicating that the level of AR carried by Importin-7 was not saturated."

reach
"Cotransfection in HEK293T cells and coimmunoprecipitation experiments revealed that AR interacted with ATXN7L3 in a ligand dependent manner, similarly to GCN5, while interaction between AR and USP22 w[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
AR binds USP22 and ATXN7L3. 2 / 2
| 2

sparser
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR-dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"To further investigate this hypothesis, we analyzed putative interactions of USP22 or ATXN7L3 with AR in mammalian cells."
ALKBH5 affects USP22
| 1 12
ALKBH5 increases the amount of USP22.
| 7
ALKBH5 increases the amount of USP22. 7 / 7
| 7

reach
"In addition, ALKBH5-mediated m6A methylation upregulates the expression of USP22 and RNF40, subsequently inhibiting the ubiquitination of histone H2A, promoting osteosarcoma growth and metastasis (16)."

reach
"Methyl RNA immunoprecipitation sequencing analysis and functional studies showed that ALKBH5 mediates its protumorigenic function by regulating m 6 A levels of histone deubiquitinase USP22 and the ubiquitin ligase RNF40."

reach
"Furthermore, ALKBH5-mediated m6A deficiency increases the expression of USP22 and RNF40 in osteosarcomas, promoting osteosarcoma cell growth and proliferation [40]."

reach
"Because ALKBH5 KD increased m A levels and significantly decreased USP22 mRNA steady-state levels, we performed mRNA stability analyses in scrambled siRNA and ALKBH5-silenced osteosarcoma cells."

reach
"ALKBH5 silencing led to significantly reduced expression of USP22 in 143B, U2OS, and OS-17 osteosarcoma cell lines (Fig. 4F; Supplementary Fig. S8D–S8G)."

reach
"Conversely, ALKBH5 overexpression increased USP22 expression in osteosarcoma cells (Fig. 4G; Supplementary Fig. S8H)."

reach
"Taken together, our data suggests that USP22 is highly sensitive to changes in m A levels and that ALKBH5 induces USP22 expression by maintaining an undermethylated state as increased methylation promotes accelerated decay of USP22 transcript in ALKBH5-depleted cells."
ALKBH5 decreases the amount of USP22.
| 3
ALKBH5 decreases the amount of USP22. 3 / 3
| 3

reach
"Because ALKBH5 KD reduced USP22 expression, we wondered whether changes in ALKBH5 levels or activity affects H2A and H2B monoubiquitylation levels."

reach
"Because ALKBH5 KD increased m A levels and significantly decreased USP22 mRNA steady-state levels, we performed mRNA stability analyses in scrambled siRNA and ALKBH5-silenced osteosarcoma cells."

reach
"In addition, deficiency of ALKBH5‐mediated m6A modification in osteosarcoma causes an increase expression of histone deubiquitinase USP22 and ubiquitin ligase RNF40, which inhibits histone H2AK199 monoubiquitination, induces the expression of genes related to DNA repair, and promotes the progression of osteosarcoma."
ALKBH5 activates USP22.
| 1 2
ALKBH5 activates USP22. 3 / 3
| 1 2

reach
"ALKBH5 mediates its protumorigenic function by inducing the stability of histone deubiquitinase ubiquitin specific peptidase 22 (USP22) and ubiquitin ligase RING finger protein 40 (RNF40) in a RNA methylation-dependent manner."

eidos
"Significance : RNA demethylase ALKBH5 upregulates USP22 and RNF40 to inhibit histone H2A ubiquitination and induces expression of key replication and DNA repair-associated genes , driving osteosarcoma progression ."

reach
"In one study, the protumorigenic function of ALKBH5 was mediated by regulating m6A of histone deubiquitination ubiquitin-specific peptidase 22 (USP22) and ubiquitin ligase RING finger protein 40 (RNF40)."

reach
"Taken together, these results suggest that USP22 silencing inhibits the inflammatory response and lipid accumulation in ALD.BRD4 can bind to acetylated histones and transcription factors through its b[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, USP22 binds to the promoter region of AP4 to activate its transcription."

reach
"USP22 positively regulated transcription factor FOXP3 activity in mouse regulatory T (T reg) cells."

reach
"Consistently two of its complex partners, ATXN7L3 and ENY2, are required for optimal USP22-mediated transcription activation by nuclear receptors in vivo [67] ."

reach
"Moreover, SOX9 was also reported to bind the promoter of ubiquitin-specific peptidase 22 (USP22), a deubiquitinating enzyme, promote its transcription, and activate the Wnt/β-catenin pathway (Miao et al., 2022)."

sparser
"At the biochemical level, these Polycomb proteins function as global transcriptional repressors by catalyzing the ubiquitylation of histone H2A. In yeast, the USP22 homolog functions as a transcriptional coactivator by removing ubiquitin from a distinct core histones, H2B. Given that USP22 is expressed in cancer as part of an 11 gene signature that includes transcriptional repressors which ubiquitylate H2A, it seemed possible that USP22 might activate transcription in part via the deubiquitylation of this same substrate."

reach
"USP22 overexpression enhances colon cancer migration and invasiveness by inducing EMT via activating AP4 (activating enhancer binding protein-4) transcription by binding to its promoter[54]."

reach
"Transcriptionally, USP22 leads to H2BK120Ub on chromatins among the FOXP3 locus to enhance its transcription."

reach
"USP22 positively regulated transcription factor FOXP3 activity in mouse regulatory T (T reg) cells."

reach
"Additionally, USP22 downregulates transferrin receptor (TFRC) transcription by removing H2B lysine 120 ubiquitination (H2BK120ub) from TFRC transcription start site (TSS) downstream region, thereby inhibiting Sorafenib‐induced ferroptosis in hepatocellular carcinoma."

sparser
"Recent studies have demonstrated that USP22 can inhibit the transcription of the p21 gene by de-ubiquitinating the transcriptional regulator FBP1, leading to cell proliferation and tumorigenesis [ xref ]."

reach
"By evaluating the USP22-mediated downregulated genes in tumor cells by Landscape In Silico deletion Analysis (Lisa) [ 22 ], we found that the three transcriptional repressors present in USP22-immunopr[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects Cyclin
| 12
USP22 decreases the amount of Cyclin.
| 5
USP22 decreases the amount of Cyclin. 5 / 5
| 5

reach
"Mechanistically, we found that USP22 depletion dramatically decreased Akt phosphorylation and cyclin D2 expression."

reach
"Furthermore, USP22 small interfering (si)RNA decreased the expression of Cyclin B and Survivin, and increased the expression of p21 in HCC (7)."

reach
"USP22 siRNA also increased the expression of p21 and reduced the expression of Cyclin B in OSCC cells."

reach
"Furthermore, USP22 small interfering RNA inhibited cell growth and reduced the expression levels of Aurora-B, Survivin and Cyclin B, together with the upregulation of cyclin-dependent kinase inhibitor 1A (p21)."

reach
"Moreover, USP22 siRNA increased the p21 protein levels and decreased the Cyclin B protein levels in OSCC cells (Fig. 2A)."
USP22 activates Cyclin.
| 5
USP22 activates Cyclin. 5 / 5
| 5

reach
"USP22 can stabilize cyclin D1 and promote the nuclear accumulation of cyclin D1 (16)."

reach
"An in vitro study showed that the upregulation of USP22 mediated the enhanced expression of BMI1 and Cyclin D2, and was responsible for increased cell proliferation and the metastatic behavior of colon cancer cells ."

reach
"19 However, there has been a lack of thorough investigation into the role of USP22 in hepatocellular carcinoma development and Sorafenib resistance.In this study, we demonstrate that USP22 promotes the proliferation of hepatocellular carcinoma cells by stabilising cyclin‐dependent kinase 11B (CDK11B)."

reach
"Hence, USP22 overexpression was associated with reduced p21 and p27 levels and increased abundance of Cyclin D1, CDK4 and CDK6, which collectively form a complex that promotes G1 progression [XREF_BIBR]."

reach
"In addition, Usp22 promotes CRC by stabilizing cyclin B1."
USP22 phosphorylates Cyclin.
| 2
USP22 leads to the phosphorylation of Cyclin. 2 / 2
| 2

reach
"In vitro assays showed that USP22 depletion suppressed ATC cell survival and proliferation by decreasing Rb phosphorylation and cyclin D2, inactivating Akt, and simultaneously upregulating Rb; USP22 silencing restrained cell migration and invasion by inhibiting epithelial-mesenchymal transition; USP22 knockdown promoted mitochondrion- mediated and caspase dependent apoptosis by upregulating Bax and Bid and promoting caspase-3 activation."

reach
"Molecular analysis of the tumor tissues showed that USP22 knockdown reduced the levels of cyclin D2, Akt phosphorylation, vimentin, and Bcl-2, whereas upregulated the expressions of E-cadherin, Bax, and cleaved (cl)-caspase-3, which confirmed in vitro findings."
USP22 affects ATXN7
10 | 1
10 | 1

reach
"Western blot analysis of complexes obtained after an ATXN7 immunopurification showed that USP22 and ATXN7L3 associate with ATXN7 together with other components of TFTC and STAGA."
PPARG affects USP22
4 | 1 7
4 | 1 7

sparser
"For instance, the USP22-PPARγ axis plays a significant role in modulating both adipogenesis and inflammation, whereas disorders in bile acid metabolism and alterations in gut microbiota exhibit bidirectional regulation. xref In light of these mechanisms, preclinical studies have led to the development of small-molecule inhibitors that target USP22 and FXR."

sparser
"In vitro pulldown assays with purified recombinant proteins demonstrated that USP22 directly bound to PPARγ (Fig.  xref )."

sparser
"We subsequently performed coimmunoprecipitation experiments to confirm the interaction between USP22 and PPARγ (Fig.  xref ), and predicted their possible binding sites via HDOCK software (Fig.  xref )."

sparser
"Then, the specific interaction between USP22 and PPARγ was confirmed by co-immunoprecipitation assay with exogenously transduced USP22 and PPARγ in HEK293T cells (Fig.  xref )."

sparser
"Overall, our results confirm that USP22 specifically interacts with PPARγ, which is a key transcription factor related to lipid metabolism."

sparser
"Interestingly, we found that USP22 strongly bound to the PPARγ DBD domain (Fig.  xref ) as well as pVHL and CUL4B proteins in the HCC cells (Supplementary Fig.  xref ), and the interaction between PPARγ and USP22 significantly decreased the pVHL and CRL4B AhR involved ubiquitination (Fig.  xref ), indicating that USP22 regulates deubiquitination of PPARγ through other lysine sites."

sparser
"Taken together, these results indicate that USP22 directly interacts with PPARγ and functions as a bona fide PPARγ deubiquitinase in cells."

reach
"Mechanistically, USP22 bound to peroxisome proliferator-activated receptor gamma (PPARgamma) and inhibited its ubiquitination, thereby stabilizing PPARgamma and promoting efferocytosis."
| 11
| 7

reach
"USP22 up-regulates the SIRT1/AKT/MRP1 signaling pathway and promotes the efflux of 5-Fluorouracil (5-FU), leading to the treatment resistance of HCC [17]."

reach
"This study further discovers that USP22 activated the SRC signaling pathway by upregulating the molecule KDEL endoplasmic reticulum protein retention receptor 1 (KDELR1), thereby contributing to LUAD cell progression."

reach
"Collectively, our findings indicate that miR-200b-5p-mediated inhibition of USP22 attenuates cell proliferation by targeting the NF-κB signaling pathway in GC, suggesting that miR-200b-5p and USP22 could serve as potential diagnostic or therapeutic targets for gastric cancer and other related human diseases."

reach
"These results indicated that USP22 deficiency reversed renal EMT and TIF in diabetic kidneys by blocking the Snail1 signaling pathway (see Fig. 11 )."

reach
"These results further validated that the tumors with Usp22 overexpression were significantly larger and more aggressive than those in the control group, whereas Usp22 ablation diminished this effect.2.2 USP22 activates the mTORC1 signaling pathway."

reach
"Inhibition of USP22 by miR-200b-5p represses gastric cancer cell proliferation and migration by targeting the NF-κB signaling pathway."

reach
"USP22 maintained GC cell stemness by stabilizing BMI1 and promoted proliferation and metastasis by activating the FOXO1 and YAP signaling pathway [47,50]."
| 4

reach
"Knockdown of USP22 can activate STAT1 signaling pathway, inhibit T-cell depletion, and promote the proliferation and activation of NK cells (75)."

reach
"These results further validated that the tumors with Usp22 overexpression were significantly larger and more aggressive than those in the control group, whereas Usp22 ablation diminished this effect.2.2 USP22 activates the mTORC1 signaling pathway."

reach
"Conversely, ectopic expression of USP22 reversed this effect by modulating the NF-κB signaling pathway."

reach
"USP22 can also enhance the telomerase reverse transcriptase (TERT)/p53 signaling pathway [ 65 ] and promote cell proliferation and DNA repair (see Fig. 1 )."

reach
"The siRNA-mediated knock-down of USP22 could effectively induce cell cycle arrest by regulating target molecules, such as cyclin D2, p21, p15 and p53, and could also inhibit cell growth."

reach
"In this study, we used siRNA to specifically suppress expression of USP22, and we observed that the knock-down of USP22 could effectively inhibit HeLa cell proliferation and induce cell cycle arrest."

reach
"USP22 knockdown inhibits cell proliferation and induces cell cycle arrest in IEC-6 cells after hypoxia/reoxygenation injury."

reach
"Mechanistic analysis has further demonstrated that knockdown of ALKBH5 induces cell cycle arrest by destabilizing ubiquitin-specific peptidase (USP22) and ubiquitin ligase ring finger protein 40 (RNF40), resulting in the upregulation of p27Kip1 and Wee1, which are cyclin-dependent kinase inhibitory proteins, and downregulation of cell cycle and DNA damage and repair-related genes in osteosarcoma cells."

reach
"Our study also shows that USP22 silencing in GC cells decreases cell proliferation and induces cell cycle arrest and apoptosis in vitro, and suppresses tumor growth and metastasis in vivo."

reach
"Therefore, up-regulation of USP22 expression will lead to abnormal activation of multiple pathways to promote cell survival while down-regulation of USP22 expression can induce cell cycle arrest at G0/G1 phase in different types of cancer cells (Zhang et al., 2008)."

reach
"In bladder cancer, silencing USP22 by siRNAs induced cell cycle arrest and attenuated cell proliferation (50)."

reach
"Silencing USP22 by interfering with RNA inhibits proliferation and induces cell cycle arrest in BCa cells (133)."

reach
"Besides, Lv et al. discovered that knockdown of USP22 suppressed cell growth and induced cell cycle arrest in bladder cancer cells [ 16 ]."

reach
"USP22 silencing promotes G1-phase cell cycle arrest, thereby suggesting that USP22 promotes G1-to-S cell cycle transition [13, 28, 29]."

reach
"[Retracted] RNA interference-mediated USP22 gene silencing promotes human brain glioma apoptosis and induces cell cycle arrest."
| 2 9

eidos
"In particular , Gal-SLPs can induce a trio synergetic effect : i ) sorafenib elevated intracellular levels of ROS , which oxidize B-PDEAEA to trigger rapid shUSP22 release for efficient gene downregulation ; ii ) the downregulation of USP22 led to downregulation of multidrug resistance-associated protein 1 ( MRP1 ) and inhibition of glycolysis , dramatically impairing MDR and achieving higher intracellular sorafenib accumulation , thus generating an ROS-responsive positive feedback loop ; iii ) the downregulation of USP22 suppressed the cell metabolism of cancer cells and further influenced cancer stemness ."

reach
"In the meantime, USP22 overexpression aggravated the EMT phenotype of TNBC cells, which revealed that USP22 contributes to the EMT process in TNBC.The causative role of glycolysis in stemness and EMT [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

eidos
"Thus , we speculated that the downregulation of USP22 by Gal-SLPs could block the glycolysis and further suppress stemness features in HCC cells ."

reach
"USP22 can stabilize HIF-1α by deubiquitination and promote glycolysis induced by the hypoxia and stemness of hepatocellular carcinoma (HCC) cells."

reach
"Overexpression of USP22 in HCC cell lines, under hypoxic conditions, significantly enhanced glycolysis, as it upregulated the mRNA expression of key glycolytic enzymes (HK2, PDK1, and ENO1)."

reach
"HIF-1α knock-down resulted in the abrogation of the USP22-enhanced cancer stemness and glycolysis under hypoxic conditions [92]."

reach
"USP22 promotes hypoxia induced HCC stemness and glycolysis by deubiquitinating and stabilising HIF1alpha."

reach
"Studies have identified that USP22 promotes glycolysis in HCC via the deubiquitination of HIF-1α and activates the de novo synthesis of fatty acids in HCC by deubiquitinating PPARγ [ 22 , 36 ]."

reach
"An illustrative example is the positive feedback loop between USP22 and HIF1α, which promotes glycolysis and stemness in HCC following TP53 mutation [94]."

reach
"USP22 Promotes Osteosarcoma Progression by Stabilising β-Catenin and Upregulating HK2 and Glycolysis."
USP22 affects autophagy
| 10
| 10

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"USP22 may promote drug resistance by inducing autophagy."

reach
"Beyond regulating apoptosis, the USP22 and SIRT1 pathway may also modulate autophagy."

reach
"In addition, in a pancreatic cancer cell line (Panc-1), USP22 overexpression stimulates autophagy through the ERK1/2 pathway and hereby promotes resistance to gemcitabine treatment [XREF_BIBR]."

reach
"Through SIRT1 deubiquitination, USP22 potentially triggers autophagy, diminishing HCC cell sensitivity to chemotherapeutic agents, including 5-Fu [74] (Fig. 3B)."

reach
"USP22 might induce autophagy through deubiquitination of SIRT1, thereby reducing the sensitivity of HCC cells to chemotherapeutic drugs [ 31 ]."

reach
"USP22 upregulation may thus inhibit apoptosis and stimulate autophagy in response to treatment with DNA damaging agents or targeted inhibitors to promote resistance to chemotherapy in cancer patients."

reach
"Taken together, these findings reveal a potential mechanism underlying the chemoresistance of PC cells mediated by the regulation of USP22 mediated autophagy by miR-29c, suggesting potential targets and therapeutic strategies in PC."

reach
"USP22 induced autophagy was also found to enhance cell proliferation and resistance to starvation and chemotherapeutic drugs in Panc-1 cells, therefore expressing an overall effect that promotes cell survival."

reach
"USP22 suppresses the NLRP3 inflammasome by degrading NLRP3 via ATG5-dependent autophagy."

reach
"In addition, USP22 may induce autophagy through deubiquitination of SIRT1, thereby reducing the sensitivity of hepatoma cells to chemotherapeutic drugs (45)."
USP22 affects VIM
| 11
USP22 increases the amount of VIM.
| 8
USP22 increases the amount of VIM. 8 / 8
| 8

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"Western blot analysis showed that USP22 knockdown in H1650 suppressed expression of the mesenchymal markers N-cadherin and vimentin and increased expression of the epithelial markers E-cadherin and al[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"AP4 down-regulation partially reversed the increases in N-cadherin and vimentin expression levels induced by USP22 up-regulation; however, no changes in USP22 expression levels were observed."

reach
"Molecular analysis of the tumor tissues showed that USP22 knockdown reduced the levels of cyclin D2, Akt phosphorylation, vimentin, and Bcl-2, whereas upregulated the expressions of E-cadherin, Bax, and cleaved (cl)-caspase-3, which confirmed in vitro findings."

reach
"The results showed that USP22 knockdown significantly decreased the mRNA levels of MSRB3 and VIM, but had no obvious effect on that of MMP14 (Fig. 2J, K)."

reach
"WB results further showed that USP22 overexpression downregulated E-cadherin and ZO-1 expression, upregulated Vimentin expression ( Fig. 6 C), and upregulated the expression of TGF-β1 and ECM componen[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Moreover, knockdown of USP22 effectively decreased the expression levels of Vimentin and Twist proteins, and increased those of E-cadherin protein in DDP-resistant TNBC cells ( Figure 2 G and Suppleme[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"IHC staining and WB results also showed that USP22 deficiency increased the protein expressions of E-cadherin and ZO-1, decreased the protein expression of α-SMA and Vimentin ( Fig. 10 C and D), and r[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In HG-treated NRK-52E cells, USP22-FLAG overexpression further increased USP22 expression ( Fig. 3 C), downregulated E-cadherin and ZO-1expression, and upregulated Vimentin expression ( Fig. 3 D)."
USP22 decreases the amount of VIM.
| 3
USP22 decreases the amount of VIM. 3 / 3
| 3

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"USP22 downregulation significantly increased the expression of E-cadherin (epithelial marker) but decreased the expression of N-cadherin and vimentin (mesenchymal markers) (XREF_FIG)."

reach
"In this study, we showed that USP22 depletion significantly decreased the expressions of BMI-1, vimentin, and snail and increased E-cadherin expression in ATC cells."

reach
"USP22 transfection significantly reversed these changes by suppressing E-cadherin and promoting vimentin expression."
USP22 affects ITGB1
| 11
USP22 increases the amount of ITGB1.
| 5
USP22 increases the amount of ITGB1. 5 / 5
| 5

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"Therefore, USP22 controls breast cancer stem cell self-renewal through protecting FoxM1 from ubiquitination-mediated proteasomal degradation to enhance ITGB1 transcription."

reach
"The fact that USP22 deletion reduced ITGB1 mRNA expression suggest that USP22 regulates integrin b1 expression at transcriptional level."

reach
"Therefore, USP22 controls BCSCs self-renewal through protecting FoxM1 from ubiquitination-mediated proteasomal degradation to enhance ITGB1 transcription."

reach
"The fact that USP22 deletion reduced ITGB1 mRNA expression suggests that USP22 regulates integrin β1 expression at the transcriptional level."

reach
"This further corroborates our discovery that upregulation of USP22 in BCSCs promotes breast cancer lung metastasis through FoxM1-mediated ITGB1 gene transcription."
USP22 inhibits ITGB1.
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USP22 inhibits ITGB1. 3 / 3
| 3

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"In addition to integrin β1, USP22 deletion led to a slight cell surface reduction of several additional integrin family members including integrin α1-6 and integrin β2-3 and β5-7 but not β4, β8, and α7-8 expression determined by flow cytometry (Figure S2B)."

reach
"In addition, the loss of USP22 resulted in a significant reduction in FoxM1 binding to ITGB1 promoter (Figures 4A and 4B)."

reach
"In addition to integrin b1, USP22 deletion led to a modest but statistically significant reduction in several additional integrin family members including integrin a1–6 and integrin b2–3, b5–7, but not b4, b8 and a7–8 expression (Figure s2B), In contrast, integrin b6 expression is slightly increased in USP22-null breast cancer cells (Figure s2B)."
USP22 activates ITGB1.
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USP22 activates ITGB1. 3 / 3
| 3

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"Therefore, USP22 appears to regulate the expression of multiple integrin family members with b1 as the dominant one.We then focused on studying the functional consequences of USP22-mediated integrin b1 upregulation and assessed whether USP22 maintains breast CSC self-renewal and promotes breast cancer metastasis through integrin b1 upregulation by ectopic reconstitution of ITGB1 in USP22 knockout cells (Figure s2C-E)."

reach
"Collectively, these results demonstrate that USP22 enhances BCSCs tumorigenic potential, in part, through integrin b1 upregulation."

reach
"We then focused on studying the functional consequences of USP22-mediated integrin β1 upregulation and assessed whether USP22 maintains BCSCs self-renewal and promotes breast cancer metastasis through integrin β1 upregulation by ectopic reconstitution of ITGB1 in USP22 knockout cells (Figures S2C–S2E)."
USP22 affects IRF3
| 11
| 11

sparser
"USP22 interacted with IRF3 after viral infection in a KPNA2-dependent manner."

sparser
"USP22 is associated with IRF3 after viral infection."

sparser
"Knockdown of KPNA2 impaired virus-induced USP22-IRF3 association in MEFs ( xref ), indicating that USP22 interacts with IRF3 through KPNA2 after viral infection."

sparser
"Viral infection induced USP22-IRF3 association in the cytoplasm in a KPNA2-depedent manner, and knockdown or knockout of USP22 or KPNA2 impaired IRF3 nuclear translocation and expression of downstream genes after viral infection."

sparser
"In this context, we observed that IRF3(IL139/140AA), which is predominantly located in the nucleus, did not interact with USP22, and USP22(RR164/165AA) interacted with IRF3, as did wild-type USP22."

sparser
"We observed that USP22 interacted with IRF3 and phosphorylated IRF3 (pIRF3) in the cytoplasm but not in the nucleus after vesicular stomatitis virus (VSV) or HSV-1 infection in BMDCs ( xref )."

sparser
"Interestingly, we found that the USP22(RR164/165AA) was still associated with IRF3 ( xref )."

sparser
"These data suggest that USP22 interacts with IRF3 in the cytoplasm after viral infection."

sparser
"Because USP22-mediated virus-triggered signaling requires its enzyme activity and USP22 interacts with IRF3, we hypothesized that USP22 might catalyze deubiquitination of IRF3 and regulate the nuclear translocation of IRF3."

sparser
"Together with the observations that knockdown of KPNA2 impaired IRF3-USP22 associations after viral infection and that reconstitution of KPNA2 into USP22 knockout cells restored virus-induced nuclear translocation of IRF3 and expression of downstream genes, we concluded that USP22 promotes nuclear translocation of IRF3 primarily through deubiquitinating and stabilizing KPNA2."
USP22 affects HIF1
| 11
USP22 activates HIF1.
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USP22 activates HIF1. 6 / 6
| 6

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"USP22 boosts the stability and transcriptional activity of HIF-1α under hypoxia through deubiquitination and induces upregulation of HIF-1α downstream genes."

reach
"USP22 positively regulates c-Myc, androgen receptor, and HIF-1α actions to promote breast cancer, prostate cancer, and HCC progression [16, 53, 54]."

reach
"USP22 enhances the stability and transcriptional activity of HIF-1α under hypoxia through deubiquitination, and it induces upregulation of HIF-1α downstream genes ( Figure 3 ) (86)."

reach
"Under hypertrophic stress conditions, USP22 enhances the stability of HIF-1α through its deubiquitination activity, which further activates the TAK1-(JNK1/2)/P38 signaling pathway to lead to cardiac hypertrophy."

reach
"G protein α12 elevates USP22 expression in hepatocytes by activating hypoxia-inducible factor 1α (HIF1α) [195]."

reach
"USP22 enhances the stability and transcriptional activity of HIF-1α under hypoxia through deubiquitination and induces upregulation of HIF-1α downstream genes."
USP22 deubiquitinates HIF1.
| 3
USP22 deubiquitinates HIF1. 3 / 3
| 3

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"Mechanically, H-ADSCs carrying USP22 accounted for deubiquitinating and stabilizing HIF-1α."

reach
"Our previous study revealed that phosphorylated non-muscle myosin heavy chain 9 (p-MYH9) at Ser1943, could deubiquitinate and stabilize HIF-1α by recruiting (ubiquitin-specific protease 22) USP22 in HCC, which led to the development of LR 15."

reach
"HIF-1α is deubiquitinated and stabilised by USP22, promoting the Warburg effect in HCC cells under hypoxic conditions [96] ."
USP22 increases the amount of HIF1.
| 2
USP22 increases the amount of HIF1. 2 / 2
| 2

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"Thus, we further analyzed the effect of USP22 on HIF-1α and confirmed that ubiquitin binding to HIF-1α was significantly reduced after USP22 overexpression in HCC cells, suggesting that USP22 enhances[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Subsequently, USP22 promotes HIF-1α protein levels by deubiquitinating and facilitating the transcription of glycolytic enzymes, thereby promoting tumor progression and glycolysis in HCC."
USP22 affects H2AX
| 8 3
USP22 phosphorylates H2AX.
| 5 3
USP22 leads to the phosphorylation of H2AX. 8 / 8
| 5 3

reach
"The results confirm that USP22 could deubiquitinate H2A and promote the phosphorylation of histone H2AX, thus contributing to DNA damage repair and inducing cisplatin resistance in A549 cells."

reach
"USP22 enhances DNA damage repair and induce cisplatin resistance by promoting the phosphorylation of histone H2AX via deubiquitinating histone H2A."

reach
"Our findings in lung adenocarcinoma cell line and xenografts support that USP22 could promote the phosphorylation of histone H2AX via deubiquitinating histone H2A, thus contributing to the DNA damage repair induced by cisplatin and leading to cisplatin resistance."

reach
"USP22 Enhances DNA Damage Repair and Induces Cisplatin Resistance by Promoting the Phosphorylation of Histone H2AX via Deubiquitinating Histone H2A."

reach
"USP22 promotes phosphorylation of histone H2AX by deubiquitinating histone H2A, enhancing DNA damage repair, and inducing cisplatin resistance (56)."

sparser
"Our findings in lung adenocarcinoma cell line (Figure xref ) and xenografts (Figure xref ) support that USP22 could promote the phosphorylation of histone H2AX via deubiquitinating histone H2A, thus contributing to the DNA damage repair induced by cisplatin and leading to cisplatin resistance."

sparser
"In cisplatin-resistant lung adenocarcinoma cells, USP22 not only promotes H2A histone family member X (H2AX) phosphorylation through deubiquitination of H2A, thereby promoting DNA damage repair, but a[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"USP22 promotes phosphorylation of histone H2AX by deubiquitinating histone H2A, enhancing DNA damage repair, and inducing cisplatin resistance ( xref )."
USP22 increases the amount of H2AX.
| 3
USP22 increases the amount of H2AX. 3 / 3
| 3

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"The expression of H2AFX (H2A histone family, member X) histone was promoted by USP22 (Ubiquitin-specific protease 22), which was involved in the occurrence and progression of LUAD [48]."

reach
"USP22 induces the occurrence and development of LUAD by promoting the expression of H2AFX histone."

reach
"However, formation of γH2AX may require dynamic regulation of H2B monoubiquitination, as loss of the H2B-specific deubiquitinase USP22 led to reduced levels of γH2AX (11)."
TP53 affects USP22
| 7 4
TP53 inhibits USP22.
| 4
TP53 inhibits USP22. 4 / 4
| 4

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"The upregulation of TP53 inhibited the upregulation of USP22."

reach
"To further validate the effect of p53 on USP22 expression, we depleted p53 in RKO and HCT116 cells, and found USP22 was increased by p53 depletion at both protein and mRNA levels (Fig. 4C, D)."

reach
"Our study demonstrates that p53 represses USP22-mediated stabilization of FASN, thus provides an alternative pathway connecting p53 and fatty acid synthesis."

reach
"In HCC cells with wild-type TP53 , p53 blocks the HIF-1α-induced upregulation of USP22."
TP53 binds USP22.
| 4
| 4

sparser
"In addition, the identified USP22-p53 interaction is of potential biological significance."

sparser
"USP22 appears to form a complex with Sirt1 and p53 to suppress p53 acetylation, since we detected the interaction of USP22 with p53 in transiently transfected HEK293 cells ( Figure 5 E)."

sparser
"However, the interaction of USP22 with p53 was only detected in wild-type but not in sirt1 null MEF cells ( Figure 5 F)."

sparser
"Therefore, the interaction of USP22 with p53 is probably mediated by Sirt1."
TP53 decreases the amount of USP22.
| 3
TP53 decreases the amount of USP22. 3 / 3
| 3

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"We have found H O -induced p53 expression inhibits the transcription of USP22, which otherwise deubiquitinates and stabilizes FASN."

reach
"In p53 wild-type colorectal cancer (CRC) cells, hydrogen peroxide (H 2 O 2 )-induced p53 expression represses the transcription of deubiquitinase USP22, which otherwise deubiquitinates and stabilizes Fatty Acid Synthase (FASN), and thus inhibits fatty acid synthesis."

reach
"P53 transcriptionally represses USP22 expression under H 2 O 2 treatment."
SP1 affects USP22
| 1 3 4
SP1 inhibits USP22.
| 1 2
SP1 inhibits USP22. 3 / 4
| 1 2

eidos
"In summary , our results demonstrate that USP22 expression is promoted by AP2 and c-Myc and is suppressed by SP1 and ATF3 at the transcriptional level ."

reach
"To demonstrate the upregulation of USP22 by USP7 knockdown is associated with desuppression of SP1 transcriptional activity, USP7 was knocked down, 24 h later, control plasmid and SP1 plasmids were individually introduced to overexpress SP1 in H1299 cells, additional 48 h later, proteins were extracted, Western blot analysis showed that reintroduction of SP1 significantly attenuated the upregulation of USP22 by USP7 knockdown (Fig. 2D)."

reach
"In contrast, knockdown of Sp1 enhanced USP22 promoter activity and mRNA levels."
SP1 binds USP22.
| 4
| 4

sparser
"For example, SP1 and protein kinase A/cAMP response element-binding protein could bind to the USP22 promoter to suppress or promote USP22 transcription, respectively ( xref , xref )."

sparser
"Interaction between Sp1 and the USP22 Promoter."

sparser
"Immunoprecipitation of cross-linked chromatin from HFL1 cells with an anti-Sp1 antibody followed by PCR amplification of the region (the sequence between-210 and +52) confirmed that the endogenous Sp1 protein does bind to this region of the USP22 promoter in HFL1 ( xref )."

sparser
"Results showed that Sp1 binds to the USP22 promoter in fibroblasts, suggesting that Sp1-DNA interaction may be required for USP22 repression."
SP1 decreases the amount of USP22.
| 1
SP1 decreases the amount of USP22. 1 / 3
| 1

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"For example, SP1 and protein kinase A/cAMP response element binding protein could bind to the USP22 promoter to suppress or promote USP22 transcription, respectively."
SNAI1 affects USP22
| 5 6
| 5 6

sparser
"WB results showed that the mutation of lysine residues in C-terminal or N-terminal had no effect on the binding of USP22 and Snail1 ( Fig. 8 C)."

sparser
"Despite the mutation of lysine residues in C-terminal or N-terminal having no effect on the binding of USP22 and Snail1, the WT Snail1 and the Snail1(C-8KR) mutant plasmid could decrease the ubiquityl[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Therefore, the USP22-Snail1 signal axis may be a new target for delaying diabetes TIF. (i) EMT induced by HG can promote TIF, which is one of the main pathological changes induced by DN."

reach
"Co-IP experiment results also showed that USP22 could bind to Snail1 in NRK-52E cells ( Fig. 7 C), and HG stimulation obviously increased their interactions ( Fig. 7 C)."

reach
"We further performed Co-IP experiment to determine whether the mutation of lysine residues in C-terminal and N-terminal affected the binding between USP22 and Snail1."

sparser
"Further study found that quercetin inhibited the interaction between USP22 and Snail1, thereby reducing the stability of Snail1."

reach
"Further study found that quercetin inhibited the interaction between USP22 and Snail1, thereby reducing the stability of Snail1."

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"Despite the mutation of lysine residues in C-terminal or N-terminal having no effect on the binding of USP22 and Snail1, the WT Snail1 and the Snail1(C-8KR) mutant plasmid could decrease the ubiquityl[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Co-IP experiment results also showed that USP22 could bind to Snail1 in NRK-52E cells ( Fig. 7 C), and HG stimulation obviously increased their interactions ( Fig. 7 C)."

sparser
"Co-IP assay was used to verify whether USP22 and Snail1 can interact with each other."
IRF3 affects USP22
| 11
| 11

sparser
"USP22 interacted with IRF3 after viral infection in a KPNA2-dependent manner."

sparser
"USP22 is associated with IRF3 after viral infection."

sparser
"Knockdown of KPNA2 impaired virus-induced USP22-IRF3 association in MEFs ( xref ), indicating that USP22 interacts with IRF3 through KPNA2 after viral infection."

sparser
"Viral infection induced USP22-IRF3 association in the cytoplasm in a KPNA2-depedent manner, and knockdown or knockout of USP22 or KPNA2 impaired IRF3 nuclear translocation and expression of downstream genes after viral infection."

sparser
"In this context, we observed that IRF3(IL139/140AA), which is predominantly located in the nucleus, did not interact with USP22, and USP22(RR164/165AA) interacted with IRF3, as did wild-type USP22."

sparser
"We observed that USP22 interacted with IRF3 and phosphorylated IRF3 (pIRF3) in the cytoplasm but not in the nucleus after vesicular stomatitis virus (VSV) or HSV-1 infection in BMDCs ( xref )."

sparser
"Interestingly, we found that the USP22(RR164/165AA) was still associated with IRF3 ( xref )."

sparser
"These data suggest that USP22 interacts with IRF3 in the cytoplasm after viral infection."

sparser
"Because USP22-mediated virus-triggered signaling requires its enzyme activity and USP22 interacts with IRF3, we hypothesized that USP22 might catalyze deubiquitination of IRF3 and regulate the nuclear translocation of IRF3."

sparser
"Together with the observations that knockdown of KPNA2 impaired IRF3-USP22 associations after viral infection and that reconstitution of KPNA2 into USP22 knockout cells restored virus-induced nuclear translocation of IRF3 and expression of downstream genes, we concluded that USP22 promotes nuclear translocation of IRF3 primarily through deubiquitinating and stabilizing KPNA2."
EZH2 affects USP22
2 | 8 1
EZH2 decreases the amount of USP22.
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EZH2 decreases the amount of USP22. 7 / 7
| 7

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"Consequently, EZH2 suppresses the expression of ubiquitin-specific peptidase 22 (USP22) by modifying H3K27me3, leading to enhanced OS, inflammation, and apoptosis in rat cardiomyocytes [52]."

reach
"The results indicate that EZH2 inhibition can upregulate USP22 and PD‐L1 expression in four of five primary cells (Figure 7D,E)."

reach
"Inhibition of ubiquitin-specific peptidase 22 (USP22) expression by EZH2 through H3K27me3 modification aggravates cardiac injury in SIMD rats 30."

reach
"EZH2 repressed ubiquitin-specific peptidase 22 (USP22) expression through H3K27me3 modification."

reach
"EZH2 inhibition transcriptionally upregulates USP22 expression, and upregulated USP22 further stabilizes PD-L1."

reach
"By contrast, the deubiquitinating enzyme USP22 promotes PD-L1 stabilization in colon cancer (COAD), while EZH2 inhibits USP22 transcription via H3K27me3, leading to PD-L1 degradation [165]."

reach
"Overexpression of EZH2 could modestly inhibit the transcription of USP22 and compensate for the increased USP22 expression resulting from EZH2 knockdown (Figure 3F,G)."
EZH2 binds USP22.
2 | 1 1
2 | 1 1

sparser
"About the possible relationship among BMI1 , EZH2 , and USP22 , it has been reported that USP22 increases BMI1 : in gastric cancer, USP22 inactivation decreases the neoplasm growth, while high levels of BMI1 accelerate the cell cycle via INK4a/ARF and AKT. xref Moreover, a reciprocal interaction between EZH2 and USP22 has been described in the sepsis‐induced myocardial dysfunction model: in this case, EZH2 represses USP22 through the H3K27me3 modification. xref In acute myeloid leukemia, the activation of BAX and the inhibition of BCL2 lead to a reduced expression of BMI1 and EZH2 . xref Finally, both BMI1 and EZH2 seem to be targets for the same miRNA, the miR200 that is a key regulator of the epithelial‐mesenchymal transition; indeed, a reduced expression of these genes reduces the migration capacity of colon cancer cells. xref "

reach
"18 Moreover, a reciprocal interaction between EZH2 and USP22 has been described in the sepsis‐induced myocardial dysfunction model: in this case, EZH2 represses USP22 through the H3K27me3 modification."
USP7 affects USP22
| 10
USP7 inhibits USP22.
| 5
USP7 inhibits USP22. 5 / 5
| 5

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"Taken together, our findings demonstrate that USP7 inhibition can dramatically upregulate USP22 in cancer cells; and targeting USP7 and USP22 may represent a more effective approach for targeted cancer therapy, which warrants further study."

reach
"In this study, through screening known USPs inhibitors against USP22 DUB complex, we, for the first time, surprisedly found that pharmaceutical inhibition or knockdown of USP7 can dramatically upregulate USP22 protein in cancer cells independent of these cells’ p53 status."

reach
"To further validate the specificity of the upregulation, we made siRNA-mediated USP7-knockdown (USP7-Kd) in A549 and H1299 lung cancer cells and found that USP7-Kd also markedly upregulates USP22 (Fig. 1D) in both cancer cell lines."

reach
"Upregulation of USP22 by USP7 inhibition through SP1 degradation."

reach
"Inhibition of USP7 upregulates USP22 and activates its downstream cancer-related signaling pathways in human cancer cells."
USP7 activates USP22.
| 5
USP7 activates USP22. 5 / 5
| 5

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"In the study, we found that USP7 inhibition by three inhibitors: HBX4418 [45], P5091 [41], and FT671 [12] dramatically upregulates USP22 in cancer cells through transcriptional mechanisms, and given the critical role of USP22 in carcinogenesis and anticancer response, we propose that the upregulated USP22 may represent a critical side-effects of USP7 inhibition.We further examined the expression and activation of downstream signaling of USP22 pathway by USP7 inhibition and found that this feedback upregulation has a significant impact on USP22 pathway and USP22-related oncogenes."

reach
"In the study, we found that targeting USP7 via either siRNA-mediated knockdown or pharmaceutical inhibitors dramatically upregulates USP22 in cancer cells."

reach
"We hypothesized that USP7-induced USP22 may be responsible for the insensitivity."

reach
"Targeting USP7 significantly upregulated USP22 in cancer cells."

reach
"We found that the USP7-specific inhibitor FT671 [12] dramatically upregulates USP22 protein in a dose-dependent manner, even at a very low concentration that barely affects cellular proliferation in lung cancer cell line A549 (Fig. 1A)."
USP22 affects RCAN1
3 | 3 2
USP22 binds RCAN1.
3 | 2 2
3 | 2 2

reach
"In the present study, we have identified a novel interaction between USP22 and RCAN1 (RCAN1-1S) in the mammalian cells."

reach
"Moreover, we found that RCAN1 was bound to USP22 in basal conditions, and interferon-alpha (IFN-alpha) treatment caused the dissociation of RCAN1 from USP22, which subsequently triggered RCAN1 ubiquitination and proteasome degradation."

sparser
"Moreover, we found that RCAN1 was bound to USP22 in basal conditions, and interferon-α (IFN-α) treatment caused the dissociation of RCAN1 from USP22, which subsequently triggered RCAN1 ubiquitination and proteasome degradation."

sparser
"In the present study, we have identified a novel interaction between USP22 and RCAN1 (RCAN1-1S) in the mammalian cells."
USP22 activates RCAN1.
| 1
USP22 activates RCAN1. 1 / 3
| 1

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"In addition, the overexpression of USP22 caused the increase of RCAN1 protein stability."
USP22 affects IL2
| 8
USP22 increases the amount of IL2. 8 / 10
| 8

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"Depletion of USP22 in T cells inhibits IL2 transcription by regulating NFATc2 stability."

reach
"USP22 can deubiquitinate and stabilize NFATC2 to activate T cells and upregulate IL-2 release."

reach
"Knockdown of USP22 decreases IL2 expression in Jurkat cells upon activation."

reach
"Taken together, these data suggest that USP22 positively regulates TCR-induced T cell activation by restricting IL2 production.To investigate how USP22 upregulates IL2 expression, we hypothesized that[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Furthermore, we showed that USP22 stabilizes NFATc2 protein and promotes NFATc2 function to facilitate IL2 expression in T cells.Previous studies reported that USP22 regulates gene transcription in a [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP22 can promote the expression of interleukin-2 in T cells by deubiquitinating and stabilizing NFATc2 (an important regulator of T-cell activation)—a novel role for USP22 as a positive regulator of NFATc2 in the control of T-cell immune responses (25)."

reach
"Interestingly, USP22 has recently been identified to positively regulate NFATc2 and promote IL-2 expression, which might function redundantly with USP15 in regulating Th cell differentiation [113] ."

reach
"NFATc2 is a regulatory molecule in T-cell activation, and USP22 promotes interleukin-2 expression in T cells by deubiquitinating and maintaining the stability of NFATc2."
USP22 affects DPP4
1 | 3 6
1 | 3 6

reach
"These results suggest that HuCD26mAb mediates the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus.We further examined a functional analysis on HuCD26mAb-mediated removal of[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"On the other hand, HuCD26mAb-mediated internalization of cell surface CD26 leads to the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus to counteract heterochromatin silenc[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"These data suggest that USP22 expression regulates CD26 expression through its physical interaction of a deubiquitinating enzyme activity in USP22 + CD26 + MPM cells."

sparser
"In view of their cellular localization, the mechanisms involved in CD26-USP22 interaction in MPM cells would need to be elucidated."

sparser
"We therefore hypothesize that nuclear localization of CD26 molecule by HuCD26mAb potentiates an association of USP22 with CD26, leading to the abrogation of USP22 protein and p21 upregulation in MPM c[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"As shown in Fig. 3 C, treatment with HuCD26mAb induced the formation of a CD26-USP22 complex in CD26 + MPM cells in a dose dependent manner of exogenous HuCD26mAb."

sparser
"These results suggest that HuCD26mAb mediates the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus."

reach
"As shown in Fig. 3 C, treatment with HuCD26mAb induced the formation of a CD26-USP22 complex in CD26 + MPM cells in a dose dependent manner of exogenous HuCD26mAb."

reach
"On the other hand, HuCD26mAb-mediated internalization of cell surface CD26 leads to the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus to counteract heterochromatin silenc[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 1 9
USP22 activates Cell Survival.
| 1 6
| 1 6

reach
"USP22 was highly expressed in NPC cells and promoted cell viability and proliferation."

reach
"Moreover, the knockdown of USP22 significantly reduced the viability of CRC cells overexpressing ZRANB1, while the elevation of USP22 in ZRANB1-deficient cells successfully enhanced cell viability to [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Under the treatment of Lenvatinib, USP22 knockdown inhibited the cell viability of drug-resistant HCC cells and promoted the apoptosis of drug-resistant cells."

reach
"Therefore, up-regulation of USP22 expression will lead to abnormal activation of multiple pathways to promote cell survival while down-regulation of USP22 expression can induce cell cycle arrest at G0/G1 phase in different types of cancer cells (Zhang et al., 2008)."

reach
"Silencing of either USP22 or RNF40 reduced short-term cell viability and clonogenic growth of osteosarcoma cells (Fig. 6A and B)."

eidos
"Similarly , Zhao et al. reported that USP22 depletion suppressed cell survival and proliferation as well as tumor growth and lung metastasis of anaplastic thyroid carcinoma cells25 ."

reach
"18 In lung cancer, USP22 promotes cell survival and proliferation by upregulating the expression levels of stemness-related markers."
USP22 inhibits Cell Survival.
| 3

reach
"Cell viability assessed with the MTT assay showed that knock-down of USP22 and COX-2 significantly reduced cell viability to approximately 63% and 47%, respectively, of control cells, whereas overexpr[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Similarly, Zhao et al. reported that USP22 depletion suppressed cell survival and proliferation as well as tumor growth and lung metastasis of anaplastic thyroid carcinoma cells XREF_BIBR."

reach
"CCK-8 assay showed that overexpressing USP22 in INS-1 cells could rescue the cell viability suppression induced by HG (Fig. 3G)."
HULC affects USP22
| 10
HULC activates USP22.
| 4
HULC activates USP22. 4 / 4
| 4

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"HULC upregulated ubiquitin specific peptidase 22 (USP22), leading to the decrease of ubiquitin mediated degradation of Sirt1 protein by removing the conjugated polyubiquitin chains from Sirt1 [XREF_BIBR]."

reach
"XREF_BIBR Similarly, HULC is able to stabilize silent information regulator 1 (Sirt1) protein in hepatocellular carcinoma cells because HULC can upregulate ubiquitin specific peptidase 22 (USP22), and suppress ubiquitin mediated degradation of Sirt1 protein by removing the conjugated polyubiquitin chains from Sirt1, XREF_BIBR leading to autophagy and chemoresistance."

reach
"HULC downregulates the abovementioned miRs and strongly upregulates USP22."

reach
"The investigation for the corresponding mechanism by which HULC stabilized Sirt1 revealed that HULC upregulated ubiquitin specific peptidase 22 (USP22), leading to the decrease of ubiquitin mediated degradation of Sirt1 protein by removing the conjugated polyubiquitin chains from Sirt1."
HULC inhibits USP22.
| 3
HULC inhibits USP22. 3 / 3
| 3

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"MiR-6825-5p, miR-6845-5p, and miR-6886-3p were down-regulated by HULC, resulting in the elevation of Sirt1, USP22, and protective autophagy, thus attenuating the sensitivity of HCC cells to chemotherapeutic agents [82]."

reach
"In HepG2 and Hep3B cells, HULC downregulates miR-6825-5p, miR-6845-5p, and miR-6886-3p, resulting in the upregulation of their target ubiquitin-specific peptidase 22 (USP22) and the inhibition of ubiquitin-mediated degradation of SIRT1 [81]."

reach
"By enhancing ubiquitin-specific peptidase 22, HULC suppresses ubiquitin-mediated degradation of cyclooxygenase-2 and silent information regulator 1 in HCC[118]."
HULC increases the amount of USP22.
| 3
HULC increases the amount of USP22. 3 / 3
| 3

reach
"In turn, HULC strongly increased USP22 protein levels and promoted SIRT1 deubiquitylation, consequently decreasing the UPP-mediated degradation of SIRT1 and increased its protein stability."

reach
"HULC can increase the expression of USP22 (Ubiquitin Specific Peptidase 22) via inhibiting miR-6886−3p, miR-6825−5p, and miR-6845−5p."

reach
"The overexpression of HULC in HCC cells increased the expression level of USP22, leading to a reduction in the ubiquitin-mediated degradation of COX-2 proteins, resulting in the upregulation of COX-2 expression and half-life prolongation."
DPP4 affects USP22
1 | 3 6
1 | 3 6

reach
"These results suggest that HuCD26mAb mediates the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus.We further examined a functional analysis on HuCD26mAb-mediated removal of[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"On the other hand, HuCD26mAb-mediated internalization of cell surface CD26 leads to the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus to counteract heterochromatin silenc[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"These data suggest that USP22 expression regulates CD26 expression through its physical interaction of a deubiquitinating enzyme activity in USP22 + CD26 + MPM cells."

sparser
"In view of their cellular localization, the mechanisms involved in CD26-USP22 interaction in MPM cells would need to be elucidated."

sparser
"We therefore hypothesize that nuclear localization of CD26 molecule by HuCD26mAb potentiates an association of USP22 with CD26, leading to the abrogation of USP22 protein and p21 upregulation in MPM c[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"As shown in Fig. 3 C, treatment with HuCD26mAb induced the formation of a CD26-USP22 complex in CD26 + MPM cells in a dose dependent manner of exogenous HuCD26mAb."

sparser
"These results suggest that HuCD26mAb mediates the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus."

reach
"As shown in Fig. 3 C, treatment with HuCD26mAb induced the formation of a CD26-USP22 complex in CD26 + MPM cells in a dose dependent manner of exogenous HuCD26mAb."

reach
"On the other hand, HuCD26mAb-mediated internalization of cell surface CD26 leads to the formation of a CD26-USP22 complex and the removal of USP22 from the nucleus to counteract heterochromatin silenc[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects VEGFA
| 8 1
USP22 increases the amount of VEGFA. 9 / 9
| 8 1

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"The results demonstrated that the transcription of VEGFA was downregulated by USP22 deletion, in contrast, ectopic expression of USP22 increased VEGFA transcription (Fig. 2D–F)."

reach
"Dual luciferase assay results showed that USP22 significantly upregulated VEGFA transcription (Fig. 2G)."

reach
"Western blotting experiments showed that the upregulation of VEGFA by USP22 was inhibited in the absence of ZEB1 (Fig. 2I), this suggested that the up-regulation of VEGFA transcription by USP22 was partly dependent on ZEB1 expression."

sparser
"Western blotting experiments showed that the upregulation of VEGFA by USP22 was inhibited in the absence of ZEB1 (Fig. xref ), this suggested that the up-regulation of VEGFA transcription by USP22 was partly dependent on ZEB1 expression."

reach
"The results demonstrated USP22 knockdown reduced mRNA or protein expression of VEGFA."

reach
"The results showed that, overexpression of USP22 significantly promoted the growth of HCC, and knockdown of ZEB1 inhibited the effect of USP22 on tumor growth (Fig. 6G–J), suggesting that ZEB1 was involved in the development of HCC, and the promotion of HCC growth by USP22 was partially related to ZEB1.In addition, the results from immunohistochemistry showed that USP22 depletion decreased USP22, ZEB1, or VEGFA expression in xenograft tumors."

reach
"The results showed that USP22 deletion inhibited VEGFA secretion (Fig. 5H)."

reach
"USP22 increases ZEB1-induced VEGFA transcription in HCC cells."

reach
"Western blotting results showed that the protein expression of VEGFA was decreased by knockdown of USP22, while the protein expression of VEGFA was upregulated by overexpression of USP22 (Fig. 2A–C)."
USP22 affects TGFB1
| 6 1
USP22 increases the amount of TGFB1.
| 5
USP22 increases the amount of TGFB1. 3 / 4
| 3

reach
"And overexpression USP22 in A549 induced an increase of TGF-beta1 expression."

reach
"Ubiquitin specific protease 22 (USP22) reduced Sirt1 ubiquitination and degradation and decreased FN and TGF-beta1 expression in GMCs under both basal and AGEs treated conditions."

reach
"Ubiquitin specific protease 22 (USP22) reduces the degradation of sirtuin-1 and the expression of FN and TGF-beta1 in AGE treated GMCs, whereas depletion of USP22 promotes sirtuin-1 degradation and the expression of FN and TGF-beta1 in this cell model."
Modified USP22 increases the amount of TGFB1. 2 / 2
| 2

reach
"Furthermore, we have found overexpression of USP22 correlated with high TGF-beta1 expression in the lung ADC tissues.To determine whether USP22 expression in lung ADC cells induced EMT by changing the[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In diabetic nephropathy, the increased expression of USP22 reduced the Sirt1 ubiquitination and degradation, and decreased fibronectin and TGF-beta1 expression in glomerular mesangial cells under both[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 activates TGFB1.
| 1 1
USP22 activates TGFB1. 2 / 3
| 1 1

sparser
"AP4 transcription is increased by TGF-β1 [ xref ], which, in turn, can be activated by USP22."

reach
"Differential regulation of gene expression, as well as loss of USP22 expression, was also demonstrated by independent qRT-PCR of the USP22-dependent upregulated genes TGFB1, SLFN5, TGM2, and DDX60, as well as downregulation of USP22, CXCR4, and AK4 (Fig. 1C), confirming the quality of the microarray."
USP22 affects Flag
| 9
| 9

sparser
"To investigate the effect of USP22 overexpression on hepatocellular carcinoma cell proliferation, we infected HepG2 and Hep3B cells with lentiviruses and established cell lines stably overexpressing FlagUSP22 or not (Figure xref )."

sparser
"We found that transduction efficiencies were more than 95% and comparable between Jurkat (Flag) and Jurkat (Flag-USP22) cells ( Fig. 4 C)."

sparser
"The lysates of HEK‐293FT cells expressing FlagUSP22 were prepared and subjected to Flag affinity purification."

sparser
"Total proteins from HEK‐293FT cells expressing FlagUSP22 or control vectors were first immunoprecipitated with anti‐Flag, followed by immunoblotting with antibodies against CDK11B. The results showed that FlagUSP22 indeed interacted with CDK11B (Figure  xref )."

sparser
"USP22 protein contains an N‐terminal zinc‐finger (ZnF) and a C19 ubiquitin‐specific peptidase (C19 peptidase) domain. xref , xref To illustrate the molecular detail involved, co‐IP assays were performed in HEK‐293FT cells expressing Flag (vector), FlagUSP22, FlagUSP22 (1‒160 aa) or FlagUSP22 (161‒525 aa) using anti‐Flag."

sparser
"Similar results were obtained in the exogenous Co-IP assays that were carried out on HEK293T cells transfected with Flag-USP22 or/and HA-c-Myc ( Figures 5 C and D), which corroborated that c-Myc is a [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Cells were transfected with HA-c-Myc combined with or without Flag-USP22 for 48 hours."

sparser
"The plasmids encoding FlagUSP22 wild‐type or catalytically inactive FlagUSP22 HH/AA (Figure  xref ) were co‐transfected with HA‐ubiquitin into HEK‐293FT cells."

sparser
"The endogenous NFATc2 protein level was higher in Jurkat (Flag-USP22) cells and Jurkat (Flag) cells treated with MG132 than Jurkat (Flag) cells ( Fig. 4 H)."
USP22 affects BRD4
3 | 4 2
USP22 binds BRD4.
3 | 1 2
3 | 1 2

sparser
"We further showed that USP22 specifically bound to BRD4 and promoted its deubiquitination in HEK239T and AML-12 cells."

reach
"We further showed that USP22 specifically bound to BRD4 and promoted its deubiquitination in HEK239T and AML-12 cells."

sparser
"These data suggest that BRD4 interacts with USP22 and is deubiquitinated by USP22 in ALD."
USP22 deubiquitinates BRD4.
| 3
USP22 deubiquitinates BRD4. 3 / 3
| 3

reach
"These data suggest that BRD4 interacts with USP22 and is deubiquitinated by USP22 in ALD.ALD is a common chronic liver disease; clinically, the disease first involves alcoholic fatty liver development[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"For example, USP22 ameliorates chronic ALD by deubiquitinating BRD4 ."

reach
"Third, USP22 facilitated the inflammatory response by deubiquitinating BRD4 in ALD."
PTGS2 affects USP22
4 | 2 3
4 | 2 3

sparser
"To examine the potential interaction between USP22 and COX-2, A549 cells were transfected with vectors expressing USP22 and Myc-tagged COX-2 and cell lysates were immunoprecipitated against anti-Myc [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"To verify the direct interaction between USP22 and COX-2, recombinant USP22 was incubated with His tagged COX-2 or COX-1 and proteins were isolated by affinity chromatography."

reach
"As shown in Fig. 2 C, USP22 was pulled down together with COX-2 but not with COX-1, confirming the specificity of the interaction between USP22 and COX-2."

sparser
"To verify the direct interaction between USP22 and COX-2, recombinant USP22 was incubated with His-tagged COX-2 or COX-1 and proteins were isolated by affinity chromatography."

sparser
"As shown in C, USP22 was pulled down together with COX-2 but not with COX-1, confirming the specificity of the interaction between USP22 and COX-2."
HPC affects USP22
| 9
HPC increases the amount of USP22.
| 4
HPC increases the amount of USP22. 4 / 4
| 4

reach
"Despite that USP22 was detectable in both nucleus and cytoplasm of CA1 cells, cytoplasmic USP22 exhibited no significant change in our ischemic model, whereas HPC restored the tGCI-induced downexpression of nuclear USP22."

reach
"Hence, it is worth investigating whether HPC induces USP22 expression in CA1 to catalyze nuclear β-catenin deubiquitination, thereby promoting nuclear β-catenin stabilization and mediating neuroprotection against tGCI.Herein we aim to explore the mechanisms by which HPC regulates β-catenin ubiquitination to stabilize nuclear β-catenin in CA1 after tGCI."

reach
"On the other hand, HPC increases USP22 expression to induce the deubiquitination of nuclear β-catenin, thereby inhibiting the degradation of nuclear β-catenin to enhance nuclear β-catenin stability and finally promoting neuronal survival after tGCI."

reach
"On the other hand, HPC increased the expression of USP22 within the neuronal nucleus in CA1 after ischemia to promote nuclear β-catenin deubiquitination."
HPC activates USP22.
| 3
HPC activates USP22. 3 / 3
| 3

reach
"HPC reduced SCF KDM2A complex and the K48-Ub of beta-catenin, and increased ubiquitin-specific peptidase 22 (USP22) in nucleus after tGCI."

reach
"HPC enhanced the neuroprotective effects of USP22."

reach
"However, HPC remarkably inhibited the reduction of nuclear USP22 induced by tGCI, but had no effect on cytoplasmic USP22 (Fig. 6E, F)."
HPC inhibits USP22.
| 2
HPC inhibits USP22. 2 / 2
| 2

reach
"In line with those findings, HPC reduces the K48-Ub of nuclear β-catenin and maintains the stability of nuclear β-catenin by increasing nuclear USP22 in CA1 after tGCI, thereby promoting the survival of neurons."

reach
"Conversely, HPC reversed tGCI-induced reductions of USP22-positive cells to a certain extent (Fig. 6B)."
Flag affects USP22
| 9
| 9

sparser
"To investigate the effect of USP22 overexpression on hepatocellular carcinoma cell proliferation, we infected HepG2 and Hep3B cells with lentiviruses and established cell lines stably overexpressing FlagUSP22 or not (Figure xref )."

sparser
"We found that transduction efficiencies were more than 95% and comparable between Jurkat (Flag) and Jurkat (Flag-USP22) cells ( Fig. 4 C)."

sparser
"The lysates of HEK‐293FT cells expressing FlagUSP22 were prepared and subjected to Flag affinity purification."

sparser
"Total proteins from HEK‐293FT cells expressing FlagUSP22 or control vectors were first immunoprecipitated with anti‐Flag, followed by immunoblotting with antibodies against CDK11B. The results showed that FlagUSP22 indeed interacted with CDK11B (Figure  xref )."

sparser
"USP22 protein contains an N‐terminal zinc‐finger (ZnF) and a C19 ubiquitin‐specific peptidase (C19 peptidase) domain. xref , xref To illustrate the molecular detail involved, co‐IP assays were performed in HEK‐293FT cells expressing Flag (vector), FlagUSP22, FlagUSP22 (1‒160 aa) or FlagUSP22 (161‒525 aa) using anti‐Flag."

sparser
"Similar results were obtained in the exogenous Co-IP assays that were carried out on HEK293T cells transfected with Flag-USP22 or/and HA-c-Myc ( Figures 5 C and D), which corroborated that c-Myc is a [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Cells were transfected with HA-c-Myc combined with or without Flag-USP22 for 48 hours."

sparser
"The plasmids encoding FlagUSP22 wild‐type or catalytically inactive FlagUSP22 HH/AA (Figure  xref ) were co‐transfected with HA‐ubiquitin into HEK‐293FT cells."

sparser
"The endogenous NFATc2 protein level was higher in Jurkat (Flag-USP22) cells and Jurkat (Flag) cells treated with MG132 than Jurkat (Flag) cells ( Fig. 4 H)."
USP22 affects sorafenib
| 8
USP22 inhibits sorafenib.
| 5
| 5

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"USP22 inhibits Sorafenib‐induced ferroptosis through transcriptional suppression of TFRC mediated by deubiquitinating H2BK120ub."

reach
"It has been reported that depletion of USP22 suppresses angiogenesis in mouse xenograft model of Non-small cell lung cancer [21], and promotes the effect of sorafenib in HCC [22]."

reach
"39 , 40 It was speculated that TFRC was essential for USP22‐mediated resistance to Sorafenib‐induced ferroptosis."

reach
"Taken together, our findings demonstrated that USP22 inhibits Sorafenib‐induced ferroptosis through the transcriptional inhibition of TFRC.2.8 USP22 depletion inhibits hepatocellular carcinoma growth and sensitises hepatocellular carcinoma to Sorafenib treatment in mice."

reach
"We have demonstrated that USP22 depletion inhibits hepatocellular carcinoma cell proliferation and promotes the sensitivity of hepatocellular carcinoma cells to Sorafenib‐induced ferroptosis by in vitro assays."
USP22 activates sorafenib.
| 3
| 3

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"Why USP22 has a preference for H2BK120ub as its substrate under Sorafenib treatment is worth further research.In summary, our study demonstrated that USP22 promotes the growth of hepatocellular carcinoma and resistance to Sorafenib‐induced ferroptosis by removing ubiquitin moieties from non‐histone protein CDK11B and histone H2B."

reach
"USP22 promotes the proliferation of hepatocellular carcinoma cells by stabilising CDK11B and enhances resistance to Sorafenib by inhibiting ferroptosis through the USP22/H2BK120ub/TFRC axis.So far, more than 40 USPs have been directly or indirectly associated with relevant cancer processes."

reach
"The USP22 targeting lipopolyplexes caused high tumour inhibition and high sorafenib sensitivity in mice bearing HCC."
USP22 affects cell death
| 1 6
USP22 activates cell death.
| 1 3
| 1 3

reach
"While it is clear that USP22 is overexpressed in various cancer types and may promote oncogenesis by altering gene expression, cell death and cell cycle progression, emerging evidence suggests that USP22 also harbors tumor suppressor like properties."

reach
"USP22 accelerates necroptotic cell death in several cancer cells via regulating RIPK3 ubiquitination [55]."

eidos
"In addition to USP2-1 , USP2-2 also caused caspase-8 activation , cleavage of poly ADP-ribose polymerase ( PARP ) , and promotion of cell death [ 78 ] ."

reach
"For instance, loss of ubiquitin-specific peptidase 22 expressions significantly decreases the TNF-α-induced necroptosis cell death ( Roedig et al., 2021 ), whereas, administration of pterostilbene inh[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 inhibits cell death.
| 3
| 3

reach
"Moreover, USP22, SIRT1, or SLC7A11 elevation contributed to enhanced cardiomyocyte viability and attenuated ferroptosis induced cell death in vitro, accompanied by increased GSH levels, as well as decreased reactive oxygen species production, lipid peroxidation, and iron accumulation."

reach
"USP22 inhibition restores cisplatin sensitivity in cisplatin-resistant lung cancer cells.Cisplatin can induce cell death by activating mitochondrial apoptosis."

reach
"In our study, after treatment with FO, the expression of USP22 was markedly increased, and the increase in USP22 increased the activity of Sirt1 and decreased myocardial cell death."
| 1 7
| 1 7

reach
"In addition, hypoxic preconditioning has been used as a tool for the regulation of angiogenesis, and hypoxic adipose stem cell-derived sEVs carrying a high abundance of USP22 promote angiogenesis and cutaneous wound healing through upregulation of lncRNA H19 [83]."

reach
"It has been reported that depletion of USP22 suppresses angiogenesis in mouse xenograft model of Non-small cell lung cancer [21], and promotes the effect of sorafenib in HCC [22]."

reach
"For instance, USP22, a ubiquitin hydrolase, enhances the proliferation, angiogenesis, and recurrence of NSCLC."

reach
"Knockdown of USP22 in an in vivo model was shown to decrease tumor angiogenesis, impair non-homologous DNA damage repair pathways and significantly improve the therapeutic efficacy of cisplatin."

reach
"Notably, USP22 knockout dramatically suppressed in vitro proliferation, colony formation; and angiogenesis, growth, metastasis of A549 and H1299 in mouse xenograft model, and significantly prolonged survival of metastatic cancer bearing mice."

reach
"Moreover, USP22 affected the function and interaction of different cell types in placental labyrinth vessels, while other studies have shown that USP22 knockout could significantly reduce tumour angio[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"These data suggest that USP22 promotes angiogenesis in HCC partially related to VEGFA expression."

eidos
"USP22 depletion decreased cell proliferation , migration , Vascular Mimicry ( VM ) formation , and angiogenesis ."
USP22 affects Wnt/β-catenin
| 8
USP22 activates Wnt/β-catenin.
| 5
USP22 activates Wnt/β-catenin. 5 / 5
| 5

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"Silencing USP22 can inhibit the Wnt/β-catenin pathway to reduce cell stemness and enhance the sensitivity of PC cells to cisplatin."

reach
"USP22 was found to promote β-catenin nuclear localization, thus increasing the activity of the Wnt/β-catenin pathway in cancer cells [41] ."

reach
"Besides that, our findings also verified that USP22 interacted with ADAM9 to mediate the activation of Wnt/β-catenin pathway in trophoblast cells.DUBs consists of five families: USPs, UCHs, OTUs, JAMM[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Moreover, SOX9 was also reported to bind the promoter of ubiquitin-specific peptidase 22 (USP22), a deubiquitinating enzyme, promote its transcription, and activate the Wnt/β-catenin pathway (Miao et al., 2022)."

reach
"Silencing USP22 inhibited the Wnt/β-catenin pathway."
USP22 inhibits Wnt/β-catenin.
| 3
USP22 inhibits Wnt/β-catenin. 3 / 3
| 3

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"Down-regulation of USP22 reduces cell stemness and enhances the sensitivity of pancreatic cancer cells to cisplatin by inactivating the Wnt/β-catenin pathway."

reach
"Inhibition of USP22 reduced the cell stemness and augmented the sensitivity of PC cells to cisplatin by inhibiting the Wnt/β-catenin pathway."

reach
"It has also been shown that micro RNA-30-5p attenuated the stemness and chemoresistance of CRC stem cells via attenuating the expression of USP22 and suppressing the Wnt/β-catenin signaling [42] ."
USP22 affects TGFB
| 8
USP22 increases the amount of TGFB.
| 5
USP22 increases the amount of TGFB. 5 / 5
| 5

reach
"In addition to promoting EMT, USP22 overexpression upregulated the expression of TGF-β1 and ECM components including FN, Collagen I, and Collagen Ⅳ under HG conditions ( Fig. 3 F)."

reach
"WB results further showed that USP22 overexpression downregulated E-cadherin and ZO-1 expression, upregulated Vimentin expression ( Fig. 6 C), and upregulated the expression of TGF-β1 and ECM componen[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"A recent study suggested that USP22 could increase TGF-beta expression and promote the epithelial-mesenchymal transition (EMT)."

reach
"IHC staining and WB results also showed that USP22 deficiency increased the protein expressions of E-cadherin and ZO-1, decreased the protein expression of α-SMA and Vimentin ( Fig. 10 C and D), and r[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"The overexpression of USP22 can promote EMT occurrence and increase TGF-β1 expression in tumor cells ( Hu et al., 2015 )."
USP22 activates TGFB.
| 3
USP22 activates TGFB. 3 / 3
| 3

reach
"Together, these data indicate that Usp22 and Usp21 are mediated through distinct TGF-β signaling pathways, and the TGF-β–induced Usp22 as positive feedback to stabilize SMAD2 and SMAD4 (fig."

reach
"TGF-beta signaling up-regulates Usp22 and Usp21 through distinctive pathways."

reach
"Therefore, our data suggest that Usp22 functions to reciprocally enhance TGF-β signaling through SMAD2 and SMAD4 protein stabilization."
USP22 affects STAT1
| 5 3
USP22 phosphorylates STAT1.
| 1 3
USP22 leads to the phosphorylation of STAT1. 4 / 4
| 1 3

sparser
"Indeed, this finding could be confirmed by blocking type I/II IFN signaling with a neutralizing antibody against IFNAR2 that did not affect USP22-induced STAT1 phosphorylation (Fig. xref )."

sparser
"Interestingly, despite efficient KO of the individual PRRs in both NHT and USP22 KO HT-29 cells, additional deletion of RIG-I, MDA5, or TLR3 did not decrease USP22-dependent STAT1 phosphorylation or ISG56 expression (Fig. xref )."

sparser
"Interestingly, despite efficient KO of the 245 individual PRRs in both NHT and USP22 KO HT-29 cells, additional deletion of RIG-I, 246 MDA5 or TLR3 did not decrease USP22-dependent STAT1 phosphorylation or ISG56 247 expression (Figure 4B -D)."
| DOI

reach
"Loss of USP22 expression in Caco-2 cells triggered phosphorylation of STAT1 and increased expression of STING, compared to wild-type (WT) and NHT CRISPR/Cas9 control Caco-2 cells (Fig. 6A)."
USP22 inhibits STAT1.
| 4
USP22 inhibits STAT1. 4 / 4
| 4

reach
"Knockdown of USP22 can activate STAT1 signaling pathway, inhibit T-cell depletion, and promote the proliferation and activation of NK cells (75)."

reach
"USP22-deficient hIECs strongly 46 upregulate genes involved in IFN signaling and viral defense, including numerous IFN-47 stimulated genes (ISGs), with increased secretion of IFN-λ and enhanced STAT1 48 signaling, even in the absence of exogenous IFNs or viral infection."
| DOI

reach
"USP22-deficient hIECs strongly upregulate genes involved in IFN signaling and viral defense, including numerous IFN-stimulated genes (ISGs), with increased secretion of IFN-lambda and enhanced STAT1 signaling, even in the absence of exogenous IFNs or viral infection."

reach
"211 212 USP22 negatively regulates STAT1 signaling and IFN-λ1 expression 213 The expression of ISGs is typically induced upon activation of IFN signaling pathways 214 during pathogen invasion or autoinflammatory disease and serves to control 215 inflammation and other defensive mechanisms 9 ."
| DOI
USP22 affects PPAR
| 8
USP22 activates PPAR.
| 6
USP22 activates PPAR. 6 / 6
| 6

reach
"USP22 promotes lipogenesis in Tregs through stabilization of peroxisome proliferator-activated receptor gamma (PPARγ) [51] ."

reach
"Unexpectedly, the antiatherosclerotic effect of USP22 was associated with increased efferocytosis through its ability to bind to peroxisome proliferator-activated receptor γ (PPARγ) and modulate PPARγ stability."

reach
"We next investigated whether USP22 enhances efferocytosis by regulating PPARγ."

reach
"Thus, these results suggest that USP22 increases the stability of the PPARγ protein by binding to PPARγ in the nucleus of macrophages."

reach
"To determine the mechanism of PPARγ reduction, we used the protease inhibitor MG132 to intervene in THP-1-derived macrophages, and the protein reduction in PPARγ caused by Si-USP22 was blocked by MG132 (Fig. 5G)."

reach
"In addition, we cannot completely exclude the possible role of USP22 in other cell types in atherosclerosis in this study.In conclusion, we showed that USP22 in macrophages promotes efferocytosis and alleviates the progression of atherosclerosis in ApoE mice by regulating PPARγ."
USP22 deubiquitinates PPAR.
| 2
USP22 deubiquitinates PPAR. 2 / 2
| 2

reach
"Taken together, in this study, Ning et al. demonstrated that USP22 promotes de novo synthesis of fatty acids and tumorigenesis by deubiquitinating PPARγ in HCC (Fig. 1) [18]."

reach
"Furthermore, USP22 significantly promoted PPARγ deubiquitination (Supplementary Fig. S4)."
USP22 affects NSCLC
| 8
USP22 activates NSCLC. 8 / 8
| 8

reach
"Our study has deepened the understanding of how USP22 promotes NSCLC tumorigenesis."

reach
"Taken together, our results suggest that USP22 promotes NSCLC tumorigenesis in vitro and in vivo through MDMX upregulation and subsequent p53 inhibition."

reach
"For instance, USP22, a ubiquitin hydrolase, enhances the proliferation, angiogenesis, and recurrence of NSCLC."

reach
"However, whether USP22 promotes tumorigenesis in NSCLC remains unclear."

reach
"In summary, our results suggest that USP22 promotes NSCLC tumorigenesis in vitro and in vivo through MDMX upregulation and subsequent p53 inhibition."

reach
"The mechanism by which USP22 promotes NSCLC tumorigenesis is that USP22 can directly bind and upregulate MDMX (E3 ubiquitin ligase) in NSCLC cells and subsequently inhibit the P53 pathway to promote NSCLC tumorigenesis (76) ( Table 1 )."

reach
"USP22 promotes the invasion of NSCLC in a TFAP2A-dependent manner [11]."

reach
"Therefore, our results from the mouse xenograft model demonstrate that USP22 silencing inhibits NSCLC tumorigenesis in vivo through regulating the MDMX-p53 pathway."
USP22 affects Mice
| 8
USP22 activates Mice.
| 5
USP22 activates Mice. 5 / 5
| 5

reach
"In vivo experiments reveal that USP22 knockdown in GC cells significantly reduces xenograft tumor growth and lung metastasis in SCID mice."

reach
"Moreover, inhibition of USP22 reduced osteosarcoma tumor growth and metastasis in mice (28)."

reach
"2.1 Overexpression of Usp22 accelerates c-Myc/NRasGV12-induced HCC in mice."

reach
"IHC staining and WB results in the kidney of db/db mice showed that USP22 expression significantly increased in db/db mice compared with those in control group."

reach
"PAS staining results showed that USP22 deficiency attenuated the renal tubular hypertrophy and mesangial expansion of diabetic mice ( Fig. 9 C)."
USP22 inhibits Mice.
| 3
USP22 inhibits Mice. 3 / 3
| 3

reach
"Indeed, USP22 was recently shown to be involved in the early development of mouse embryos, with USP22 deletion causing embryonic lethality and mice carrying a hypomorphic USP22 allele displaying viabi[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Accordingly, USP22 downregulation also attenuated cerebral I/R-induced oxidative stress, inflammation, and cell apoptosis in mice, thereby reducing nerve injury and neurological dysfunction [20]."

reach
"However, deletion of Usp22 significantly decreased tumor burden and increased survival of MMTV-NIC mice."
USP22 affects MSH3
| 8
USP22 activates MSH3.
| 5
USP22 activates MSH3. 5 / 5
| 5

reach
"3.5 USP22 activates the AKT and MRP1 pathway depending on SIRT1 in HCC cells."

reach
"XREF_FIG A, knockdown of USP22 by shRNA inhibited the AKT and MRP1 pathway compared with control shRNA cells and wild-type cells."

reach
"Similarly, overexpression of USP22 in BEL/7402 cells activated the AKT and MRP1 pathway."

reach
"Simply put, USP22 may activate the SIRT1–AKT–MRP1 pathway and consequently promote MDR in human HCC cells (226)."

reach
"Both of these findings indicated that USP22 upregulated MRP1 depending on the AKT pathway."
USP22 increases the amount of MSH3.
| 3
USP22 increases the amount of MSH3. 3 / 3
| 3

reach
"Subsequently, using qPCR array analysis, we found that knockdown of USP22 could drastically inhibit the expression of MRP1, but not P-gp, in BEL/FU cells."

reach
"Collectively, USP22 might deubiquitinate SIRT1 and subsequently activate the AKT pathway, increasing the expression of MRP1 to induce MDR in HCC cells."

reach
"LY294002 (10mum) also suppressed USP22 induced high expression of MRP1 via inhibition of AKT pathway in BEL7402 cells."
USP22 affects KAT2B
3 | 3 2
3 | 3 2

sparser
"We observed similar results using the PCAF HAT inhibitor garcinol ( xref ), which blocked H2BK120 acetylation in vitro () while not affecting PCAF-USP22 interactions or recruitment of USP22 to DNA damage sites in treated cells ()."

sparser
"Regardless, our data have identified and mapped interactions between PCAF and USP22, as well as further revealing the importance of the HAT and BRD domains of PCAF in mediating DNA damage signaling and repair of DSBs by HR."

reach
"Streptavidin pull-down of SFB-tagged PCAF validated our AP-MS, as interactions between PCAF and the SAGA DUB components USP22, ENY2, and ATXN7L were observed (Fig. 4B)."

reach
"In addition to PARP inhibitor sensitivity, GSK4027 treatment also reduced recruitment of PCAF and USP22 to DNA damage sites, again consistent with the BRD of PCAF being an essential domain linking PCAF to its DNA damage functions (Fig. 4I)."

reach
"Regardless, our data have identified and mapped interactions between PCAF and USP22, as well as further revealing the importance of the HAT and BRD domains of PCAF in mediating DNA damage signaling and repair of DSBs by HR.The observation of a H2BK120 ubiquitin to acetyl switch at DSBs (Clouaire et al. 2018) and the potential involvement of PCAF in mediating these events prompted us to test directly the involvement of PCAF in regulating H2BK120 acetylation."
USP22 affects HK2
| 6 2
USP22 activates HK2.
| 4 2
USP22 activates HK2. 6 / 6
| 4 2

reach
"In addition of USP22, USP29 also deubiquitinates HIF1α, contributing to sorafenib resistance in HCC cells by promoting the transcriptional activation of target genes, especially hexokinase 2 (HK2), a key enzyme in glycolytic pathway [168]."

reach
"In addition of USP22, USP29 also deubiquitinates HIF1α, contributing to sorafenib resistance in HCC cells by promoting the transcriptional activation of target genes, especially hexokinase 2 (HK2), a key enzyme in glycolytic pathway [168]."

reach
"Overexpression of USP22 in HCC cell lines, under hypoxic conditions, significantly enhanced glycolysis, as it upregulated the mRNA expression of key glycolytic enzymes (HK2, PDK1, and ENO1)."

sparser
"Therefore, to investigate the molecular mechanism by which USP22 activates HK2 in osteosarcoma cells, we verified whether endogenous USP22 and HK2 directly bind to 143B cells."

sparser
"Therefore, to investigate the molecular mechanism by which USP22 activates HK2 in osteosarcoma cells, we verified whether endogenous USP22 and HK2 directly bind to 143B cells."

reach
"USP22 Promotes Osteosarcoma Progression by Stabilising β-Catenin and Upregulating HK2 and Glycolysis."
USP22 increases the amount of HK2.
| 2
USP22 increases the amount of HK2. 2 / 2
| 2

reach
"In addition, USP22 knockout reduced the protein expression of β-catenin and hexokinase 2 (HK2) in osteosarcoma cells."

reach
"In addition, the regulation of HK2 expression induced by USP22 depends on β-catenin."
USP22 affects AR-V7
| 8
USP22 binds AR-V7.
| 4
AR-V7 binds USP22. 4 / 4
| 4

reach
"Unlike rutaecarpine, nobiletin failed to alter the GRP78-mediated degradation of AR-V7.In conclusion, this research not only demonstrates the reason why nobiletin suppressed the growth process of CRPC through the selective degradation of AR-V7, but also enriches our understanding of the degradation mechanism of AR-V7 and provides an efficient treatment target to overcome CRPC via targeting the interaction between AR-V7 and USP14/USP22 (Fig. 7G)."

reach
"Immunofluorescence and western blot experiments showed that these molecular events mainly occurred in the nucleus.We also confirmed that nobiletin can significantly decrease the interactions between AR-V7 and USP14/USP22 in a time and concentration-dependent manner, but did not affect the interactions between AR-FL and USP14/USP22, which may explain why nobiletin exhibits a certain selectivity in inducing the degradation of AR-V7."

reach
"Binding of USP22 to AR-V7 prevents AR-V7 protein degradation."

reach
"We confirmed the endogenous interaction between AR-V7 and USP22."
USP22 activates AR-V7.
| 4
USP22 activates AR-V7. 4 / 4
| 4

reach
"Protein analysis showed that USP22 depletion significantly reduced the half-life of AR-V7."

reach
"Furthermore, CHX-tracking assay showed that USP22 depletion significantly decreased the half-life of AR-V7."

reach
"In reverse, overexpression of USP22 slowed down the attenuation of AR-V7 (Fig. 5E–H)."

reach
"Protein analysis showed that USP22 reduction significantly reduced the half-life of AR-V7."
USP22 affects ADAM9
| 5 3
USP22 binds ADAM9.
| 2 3
| 2 3

sparser
"To clarify the interaction between USP22 and ADAM9, Co-IP assay was performed."

sparser
"Besides that, our findings also verified that USP22 interacted with ADAM9 to mediate the activation of Wnt/β-catenin pathway in trophoblast cells."

sparser
"In addition, we also found USP22 bound to ADAM9 to suppress the activation of Wnt/β-catenin pathway."

reach
"In addition, we also found USP22 bound to ADAM9 to suppress the activation of Wnt/β-catenin pathway."

reach
"To clarify the interaction between USP22 and ADAM9, Co-IP assay was performed."
USP22 deubiquitinates ADAM9.
| 3
USP22 deubiquitinates ADAM9. 3 / 3
| 3

reach
"Here, we found a higher expression of USP22 in PE samples, and USP22 induced the deubiquitination of ADAM9 in trophoblast cells to promote cell dysfunction during PE process."

reach
"Mechanistically, the deubiquitinase USP22 deubiquitinated ADAM9 and stabilized its expression in trophoblast cells."

reach
"ADAM9 is highly expressed in PE, and functionally, ADAM9 is deubiquitinated by USP22 to suppress the proliferation, migration, and invasion of trophoblasts [33]."
| 6

reach
"As shown in Figure XREF_FIG, USP22 activates AP4 expression during EMT induction."

reach
"In summary, USP22 induces EMT by activating AP4 transcription to enhance CRC cell migration and invasion."

reach
"USP22 up-regulation enhances CRC cell migration and invasion and EMT related marker and AP4 expression, but these effects are partly blocked by AP4 knockdown."

reach
"The key findings of the study are as follows : (i) USP22 enhances CRC cell migration and invasion by inducing EMT, (ii) USP22 directly increases AP4 transcription to induce EMT and promote CRC cell metastasis to the lungs in vivo, and (iii) USP22 and AP4 overexpression is related to CRC progression and liver metastasis and poor outcomes in CRC patients."

reach
"Moreover, USP22 up-regulation induces EMT by directly increasing AP4 transcription, resulting in CRC cell metastasis to the lungs in vivo."

reach
"USP22 induces EMT by activating AP4 transcription."

reach
"The association between USP22 and AP4 and liver, but not lymph node, metastasis may be due to these proteins driving blood stream, but not lymphatic, metastasis of CRC cells."

reach
"Consistent with the above results, ChIP analysis revealed that USP22 binds the promoter region of AP4."
Neoplasms affects USP22
| 6
Neoplasms activates USP22.
| 5
| 5

reach
"In tumors overexpressing USP22, specific inhibitors such as those already developed for USP7 and USP8 are imaginable [88] ."

reach
"Furthermore, our study indicated that SOS1 was upregulated in USP22-overexpressing gastric cancer cells and xenograft tumor tissue, accompanied by activation of the RAS/ERK and PI3K/AKT pathways, suggesting that SOS1/RAS signaling mediates the oncogenic role of USP22 in gastric cancer."

reach
"Tumor-derived TGF-beta selectively induces Usp22 and Usp21 in T reg cells."

reach
"IHC staining showed that the tumors developed from USP22-overexpressing SGC7901 cells had significantly increased USP22 and Ki-67 staining and decreased TUNEL staining, compared with the control group (Fig. 4d)."

reach
"Two weeks later, when the tumor size was approximately 100 mm , mice carrying USP22-overexpressing tumors were randomly divided into two groups (n = 4): siRNA of ZEB1 (7.5 μg) was injected into the tumor, and the other group was injected with the same amount of negative control every 72 h for another 2 weeks."
Neoplasms increases the amount of USP22.
| 1
Neoplasms increases the amount of USP22. 1 / 3
| 1

reach
"Supporting that, a more quantitative me-RIP-qPCR analysis on tumor tissues from ALKBH5 KD group and ALKBH5-silenced osteosarcoma cells showed significantly increased m A levels and decreased expression of USP22 compared with scrambled group (Supplementary Fig. S8B and S8C)."
NFATC2 affects USP22
2 | 3 3
2 | 3 3

sparser
"We then examined the interaction between USP22 and NFATc2."

reach
"We found that USP22 could physically interact with and stabilize NFATc2 by deubiquitination."

reach
"Gao et al found that USP22 interacts with and deubiquitinates NFATc2, and also stabilizes NFATc2 protein and promotes NFATc2 function to facilitate IL-2 expression in T cells."

reach
"We then examined the interaction between USP22 and NFATc2."

sparser
"Collectively these results indicated that USP22 can interact with NFATc2."

sparser
"Because the deubiquitinase USP22 interacts with NFATc2, we hypothesized that USP22 might facilitate NFATc2 deubiquitination."
KAT2B affects USP22
3 | 3 2
3 | 3 2

sparser
"We observed similar results using the PCAF HAT inhibitor garcinol ( xref ), which blocked H2BK120 acetylation in vitro () while not affecting PCAF-USP22 interactions or recruitment of USP22 to DNA damage sites in treated cells ()."

sparser
"Regardless, our data have identified and mapped interactions between PCAF and USP22, as well as further revealing the importance of the HAT and BRD domains of PCAF in mediating DNA damage signaling and repair of DSBs by HR."

reach
"Streptavidin pull-down of SFB-tagged PCAF validated our AP-MS, as interactions between PCAF and the SAGA DUB components USP22, ENY2, and ATXN7L were observed (Fig. 4B)."

reach
"In addition to PARP inhibitor sensitivity, GSK4027 treatment also reduced recruitment of PCAF and USP22 to DNA damage sites, again consistent with the BRD of PCAF being an essential domain linking PCAF to its DNA damage functions (Fig. 4I)."

reach
"Regardless, our data have identified and mapped interactions between PCAF and USP22, as well as further revealing the importance of the HAT and BRD domains of PCAF in mediating DNA damage signaling and repair of DSBs by HR.The observation of a H2BK120 ubiquitin to acetyl switch at DSBs (Clouaire et al. 2018) and the potential involvement of PCAF in mediating these events prompted us to test directly the involvement of PCAF in regulating H2BK120 acetylation."
ENY2 affects ATXN7L3
| 8
| 8

sparser
"The SAGA DUB module is highly conserved both in subunit composition and structural organization ( xref ; xref ; xref ; xref ; xref ; xref ), The deubiquitinating enzyme USP22 (Ubp8 in yeast) closely associates with two adapter proteins, ATXN7L3 and ENY2 (Sgf11 and Sus1 in yeast, respectively) ( xref ; xref ; xref ), and a fourth protein, ATXN7 (Sgf73), anchors the DUB module to the larger SAGA complex."

sparser
"It has been reported that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator Spt-Ada-Gcn5-acetyltransferase (SAGA) complex to regulate global levels of H2B monoubiquitination, thereby promoting antibody class switch recombination by facilitating nonhomologous end joining ( xref ; xref ; xref )."

sparser
"Similarly, USP22, the human homolog of Ubp8, is bound to the STAGA complex through interactions with ATXN7L3 and ENY2, which are homologs of Sgf11 and Sus1, respectively ( xref )."

sparser
"Further, Pfam analysis of protein domain structures ( xref ) confirmed that in addition to the catalytic UCH domain, both USP27X and USP51 have an N-terminal Znf-UBP domain highly similar to that found in USP22, which is critical for USP22 interaction with ATXN7L3 and ENY2 and deubiquitination activity ( xref and xref ) ( xref )."

sparser
"We show that the ubiquitin protease USP22 forms a subcomplex with ATXN7L3 (ySgf11 homolog) and ENY2 (ySus1 homolog)."

sparser
"These results together suggest that ATXN7L3, USP22, and ENY2 form a stable subcomplex and that USP22 and ENY2 are recruited into TFTC/STAGA by ATXN7L3 ( Figure 2 D)."

sparser
"Cotransfection and coimmunoprecipitation experiments revealed that the ATXN7L3 ZnF-Sgf11 domain alone is able to interact with both ENY2 and USP22 as efficiently as the full-length ATXN7L3 (see Figur[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Previous study has shown that USP22 interacts with ENY2 and ATXN7L3 and forms the transcriptional coactivator SAGA complex to regulate global levels of H2B monoubiquitination ( xref )."
YAP1 affects USP22
2 | 4 1
2 | 4 1

sparser
"Using IP assays, USP22 interaction with YAP at the endogenous level in A375 melanoma cells was observed ( xref )."

reach
"In melanoma, USP22 interacts with and deubiquitinates YAP, thereby favoring melanoma cell proliferation."

reach
"The interaction between USP22 and KDM3A and between KDM3A and YAP1 was further validated."

reach
"USP22 interacts with and deubiquitinates YAP to prevent YAP turnover [26]."

reach
"A recent study demonstrated that ubiquitin-specific peptidase 22 (USP22) interacted with and stabilized YAP, which in turn conferred resistance to the BRAF inhibitor vemurafenib and provided new therapeutic avenues to target USP22/YAP as an option for melanoma treatment [224]."
USP22 affects pathway
| 7
USP22 activates pathway. 7 / 7
| 7

sparser
"USP22 activates PI3K/Akt/mTOR pathway via SIRT1/PTEN axis."

sparser
"To verify whether USP22 activates the SRC pathway by stabilizing KDELR1 expression, we overexpressed KDELR1 in H1299 and H1975 cells with USP22 knockdown and depleted KDELR1 in PC9 cells with USP22 overexpression."

sparser
"By up- and downregulation of USP22 expression, we also proved that USP22 can activate the Wnt/β-Catenin pathway, which in turn affected the proliferation and migration of HepG2 cells."

sparser
"SIRT1 has been reported to deacetylate PTEN and activate the PI3K/Akt/mTOR pathway. xref Deacetylated PTEN is well known to activate PI3K/AKT/mTOR pathway through promoting its phosphatase activity. xref , xref , xref To determine whether USP22 activates the PI3K/Akt/mTOR pathway via SIRT1‐mediated PTEN deacetylation, we used siRNA to silence the expression of SIRT1 in USP22‐overexpressing melanoma cells."

sparser
"These findings suggested that USP22 activates the PI3K/Akt/mTOR pathway and induces melanoma EMT via the SIRT1/PTEN axis."

sparser
"In CRC, because of the low expression of miR-30-5p, USP22 activates the Wnt/β-catenin pathway by increasing the nuclear concentration of β-catenin, and enhancing cancer stemness and tumorigenesis [ xref ]."

sparser
"Gene Set Enrichment Analysis (GSEA) further illustrated that USP22 markedly activates PI3K/Akt/mTOR pathway (Figure  xref )."
USP22 affects mTORC1
| 5 2
USP22 activates mTORC1. 7 / 7
| 5 2

reach
"In summary, USP22 activates mTORC1 by deubiquitinating FKBP12.2.5 Activated mTORC1 further inhibits the autophagic degradation of USP22."

sparser
"In addition, USP22 activated mTORC1 by deubiquitinating FK506‐binding protein 12 (FKBP12) and activated mTORC1, in turn, further stabilizing USP22 by inhibiting autophagic degradation."

reach
"Therefore, high USP22 expression in tumor tissues could predict the benefit of sirolimus in patients with HCC after LT, which supports the significant role of USP22 in activating mTORC1 in HCC.In summary, in HCC, USP22 overexpression increased mTORC1 activity and sensitized HCC toward rapamycin."

reach
"Mechanistically, USP22 interacted with FKBP12 to activate the mTORC1 pathway through deubiquitylation."

reach
"These results further validated that the tumors with Usp22 overexpression were significantly larger and more aggressive than those in the control group, whereas Usp22 ablation diminished this effect.2.2 USP22 activates the mTORC1 signaling pathway."

sparser
"In summary, USP22 activates mTORC1 by deubiquitinating FKBP12."

reach
"Because USP22 was shown to activate mTORC1 activity, we hypothesized that USP22 interacts with certain proteins associated with mTORC1, including regulatory‐associated protein of mTOR (Raptor), mTOR‐associated protein, LST8 homolog (MLST8), mTOR, and FK506‐binding protein 12 (FKBP12)."
USP22 affects XRCC6
| 7
USP22 deacetylates XRCC6. 7 / 7
| 7

reach
"USP22 can reduce Ku70 acetylation by stabilizing SIRT1 expression, thereby inhibiting Bax-mediated apoptosis and promoting cisplatin resistance.ALDH1A3, a major ALDH isoenzyme, is important for the stem cell signature of lung cancer and is associated with enhanced cisplatin resistance in lung adenocarcinoma."

reach
"USP22 Decreases the Acetylation of Ku70 by Stabilizing Sirt1, thus Inhibiting Bax Mediated Apoptosis and Inducing Cisplatin Resistance."

reach
"In addition, USP22 could also decrease the acetylation of Ku70 by stabilizing the expression of Sirt1, thus inhibiting Bax mediated apoptosis and contributing to cisplatin resistance."

reach
"These results suggest that USP22 decreases the acetylation of Ku70 by stabilizing Sirt1, thus inhibiting Bax mediated apoptosis and inducing cisplatin resistance."

reach
"(2) USP22 decreases the acetylation of Ku70 by stabilizing Sirt1, thus inhibiting Bax mediated apoptosis and inducing cisplatin resistance."

reach
"In addition, USP22 decreases the acetylation of Ku70 by stabilizing Sirt1, thus inhibiting Bax mediated apoptosis and inducing cisplatin resistance."

reach
"In addition, USP22 was shown to decrease the acetylation of Ku70 by stabilizing SIRT11 via deubiquitination."
USP22 affects STAT3
| 5
USP22 activates STAT3.
| 4
USP22 activates STAT3. 4 / 4
| 4

reach
"In colon cancer, USP22 was reported to attenuate the invasion capacity of colon cancer cells by inhibiting the STAT3 and MMP9 signaling pathway."

reach
"USP22 or MUC1 sustained activation of STAT3 signaling pathways contributes to EGFR-TKI resistance [ 51 , 60 ]."

reach
"Additionally, a recent publication showed that USP22 modulated the activity of STAT3 indirectly by stabilizing EGFR, which indicated that USP22 and USP28 may have redundant effects in NSCLC."

reach
"In conclusion, USP22 attenuated the invasion capacity of colon cancer cells by inhibiting the STAT3 and MMP9 signaling pathway."
USP22 inhibits STAT3.
| 1
USP22 inhibits STAT3. 1 / 3
| 1

reach
"USP22 prevents ubiquitination-mediated EGFR degradation, thereby inducing persistent activation of EGFR-mediated oncogenic signaling pathways, such as STAT3, AKT/mTOR, and MEK/ERK pathways."
USP22 affects LINC01426
| 7
LINC01426 binds USP22. 7 / 7
| 7

reach
"Pull-down sliver staining and RIP assay revealed that LINC01426 could interact with USP22."

reach
"LINC01426 binds with USP22 to promote SHH deubiquitination."

reach
"Pull-down Silver staining followed by western blot analysis further demonstrated the interaction between LINC01426 and USP22 in PC-9 and Calu3 cells."

reach
"Here, we uncovered that LINC01426 could interact with USP22 in LUAD cells."

reach
"Here, we determined that LINC01426 interacted with USP22 to stabilize SHH protein."

reach
"Researchers have found that LINC01426 enhances the stability of SHH protein by binding to USP22 in tumor tissues, thereby promoting lung adenocarcinoma growth and stemness."

reach
"In lung adenocarcinoma (LUAD), LINC01426 recruits and binds the USP22 protein to promote Shh protein stabilization, which activates the Hh pathway and promotes LUAD cells proliferation, migration, and EMT [149]."
USP22 affects ISG
| 7
USP22 increases the amount of ISG.
| 4
USP22 increases the amount of ISG. 4 / 4
| 4

reach
"Interestingly, 49 USP22 controls basal and 2'3'-cGAMP-induced STING activation and loss of STING 50 reversed STAT activation and ISG and IFN-λ expression."
| DOI

reach
"USP22 controls basal and 2′3′-cGAMP-induced STING ubiquitination, phosphorylation, and activation, and combined loss of USP22 and STING rescues ISG expression, STAT signaling, and IFN-λ production."

reach
"Since type III IFN-induced target genes largely overlap with genes regulated by type I IFNs [64, 65], type III IFN is likely the main IFN contributing to USP22-dependent induction of ISG expression and STAT1 activation."

reach
"Interestingly, USP22 controls basal and 2'3'-cGAMP-induced STING activation and loss of STING reversed STAT activation and ISG and IFN-lambda expression."
USP22 decreases the amount of ISG.
| 3
USP22 decreases the amount of ISG. 3 / 3
| 3

reach
"USP22 controls basal and 2′3′-cGAMP-induced STING ubiquitination, phosphorylation, and activation, and combined loss of USP22 and STING rescues ISG expression, STAT signaling, and IFN-λ production."

reach
"loss of USP22 and STING rescues ISG expression, STAT signaling and 142 IFN-λ production."
| DOI

reach
"At present, the functional role of 3CLpro-mediated inactivation of RNF20 for H2Bub1 still remains to be addressed.Since loss of USP22 mostly affects type III IFN expression and secretion, USP22 likely mediates ISG expression both via epigenetic regulatory mechanisms as well as through long-term IFN-mediated priming effects in hIECs."
USP22 affects HIF1A
2 1 | 2 2
USP22 binds HIF1A.
2 | 2
2 | 2

sparser
"In TP53-mutant HCC cells, USP22 and HIF1α formed a positive feedback loop and promote the stemness of HCC."

sparser
"Our laboratory found that HIF-1α can bind to the USP22 promoter."
USP22 deubiquitinates HIF1A.
1 | 2
USP22 deubiquitinates HIF1A. 2 / 3
1 | 2

reach
"A new study reveals that USP22 promotes the pathological process of myocardial hypertrophy by deubiquitinating HIF-1a and further activate the TAK1-(JNK1/2)/P38 signaling pathway [75]."

reach
"USP22 promotes the pathological process of myocardial hypertrophy by deubiquitinating HIF-1a which further activate the TAK1-(JNK1/2)/p38 signaling pathway [75]."
USP22 affects FKBP1A
| 2 5
| 2 5

reach
"These results indicated that the upregulation of USP22 activated the mTROC1 pathway via FKBP12.Moreover, we identified an interaction between USP22 and FKBP12."

reach
"The interaction between endogenous FKBP12 and USP22 in SNU‐449 cells is shown in Figure 5E, while the co‐localization of FKBP12 and USP22 was observed by confocal microscopy (Figure 5F)."

sparser
"Mechanistically, USP22 interacted with FKBP12 to activate mTORC1 pathway through deubiquitylation."

sparser
"Moreover, we identified an interaction between USP22 and FKBP12."

sparser
"The interaction between endogenous FKBP12 and USP22 in SNU‐449 cells is shown in Figure  xref , while the co‐localization of FKBP12 and USP22 was observed by confocal microscopy (Figure  xref )."

sparser
"Mechanistically, USP22 interacted with FKBP12 to activate the mTORC1 pathway through deubiquitylation."

sparser
"Mechanistically, USP22 interacted with FKBP12 and stabilized it via deubiquitylation."
RCAN1 affects USP22
3 | 2 2
3 | 2 2

reach
"In the present study, we have identified a novel interaction between USP22 and RCAN1 (RCAN1-1S) in the mammalian cells."

reach
"Moreover, we found that RCAN1 was bound to USP22 in basal conditions, and interferon-alpha (IFN-alpha) treatment caused the dissociation of RCAN1 from USP22, which subsequently triggered RCAN1 ubiquitination and proteasome degradation."

sparser
"Moreover, we found that RCAN1 was bound to USP22 in basal conditions, and interferon-α (IFN-α) treatment caused the dissociation of RCAN1 from USP22, which subsequently triggered RCAN1 ubiquitination and proteasome degradation."

sparser
"In the present study, we have identified a novel interaction between USP22 and RCAN1 (RCAN1-1S) in the mammalian cells."
LINC01426 affects USP22
| 7
LINC01426 binds USP22. 7 / 7
| 7

reach
"Pull-down sliver staining and RIP assay revealed that LINC01426 could interact with USP22."

reach
"LINC01426 binds with USP22 to promote SHH deubiquitination."

reach
"Pull-down Silver staining followed by western blot analysis further demonstrated the interaction between LINC01426 and USP22 in PC-9 and Calu3 cells."

reach
"Here, we uncovered that LINC01426 could interact with USP22 in LUAD cells."

reach
"Here, we determined that LINC01426 interacted with USP22 to stabilize SHH protein."

reach
"Researchers have found that LINC01426 enhances the stability of SHH protein by binding to USP22 in tumor tissues, thereby promoting lung adenocarcinoma growth and stemness."

reach
"In lung adenocarcinoma (LUAD), LINC01426 recruits and binds the USP22 protein to promote Shh protein stabilization, which activates the Hh pathway and promotes LUAD cells proliferation, migration, and EMT [149]."
IPO7 affects USP22
| 6 1
IPO7 binds USP22.
| 4 1
| 4 1

sparser
"Together, the above results indicate that Importin–7–AR–USP22 might form a relatively stable complex, even if AR and USP22 may competitively bind to Importin-7."

reach
"Mechanistically, through co-immunoprecipitation, immunofluorescence, and nuclear-cytoplasmic protein separation experiments, we discovered that AR and USP22 can bind to Importin-7 as cargoes to promote BC progression."

reach
"Importin-7 binds to AR and USP22."

reach
"In addition, our immunofluorescence analysis showed that Importin-7 binds to AR or USP22 in both the nucleus and cytoplasm (Fig. 4C–E), suggesting that Importin-7 may regulate the nuclear translocation of AR and USP22.Next, we conducted a more in-depth study of the Importin-7–AR–USP22 complex by the treatment of AR inhibitor (enzalutamide) or AR ligands, including DHT and synthetic androgen R1881."

reach
"Together, the above results indicate that Importin–7–AR–USP22 might form a relatively stable complex, even if AR and USP22 may competitively bind to Importin-7."
IPO7 activates USP22.
| 2
IPO7 activates USP22. 2 / 2
| 2

reach
"Here, our study demonstrated that Importin-7 promotes the progression of BC by controlling the nuclear transport of AR and its maintainer USP22."

reach
"We showed that the downregulation of Importin-7 causes AR and USP22 proteins to be retained in the cytoplasm and reduced in the nucleus (Fig. 5A, B)."
FOXO1 affects USP22
| 2 5
| 2 5

sparser
"The endogenous USP22 interaction with FOXO1 was further confirmed in MDA-MB-231 cells ( xref )."

sparser
"Further analysis of FOXO1 interactions with USP22 truncated mutants revealed that the zinc finger–containing N terminus of USP22 mediates its interaction with FOXO1 ( xref )."

sparser
"Stimulation with EPI markedly increased USP22 interaction with FOXO1 in breast cancer cells ( xref )."

sparser
"EPI stimulation, which induced USP22 protein expression and promoted USP22-FOXO1 interaction, markedly inhibited FOXO1 polyubiquitination in USP22 WT cancer cells, and targeted USP22 deletion resulted in a significant increase in FOXO1 ubiquitination and reduced FOXO1 protein expression even with EPI stimulation ( xref )."

reach
"The endogenous USP22 interaction with FOXO1 was further confirmed in MDA-MB-231 cells (Fig. 4L)."

reach
"Further analysis of FOXO1 interactions with USP22 truncated mutants revealed that the zinc finger–containing N terminus of USP22 mediates its interaction with FOXO1 (Fig. 4K)."

sparser
"To define USP22 functions as a possible FOXO1-specific deubiquitinating enzyme, we first determined if USP22 interacts with FOXO1."
FKBP1A affects USP22
| 2 5
| 2 5

reach
"These results indicated that the upregulation of USP22 activated the mTROC1 pathway via FKBP12.Moreover, we identified an interaction between USP22 and FKBP12."

reach
"The interaction between endogenous FKBP12 and USP22 in SNU‐449 cells is shown in Figure 5E, while the co‐localization of FKBP12 and USP22 was observed by confocal microscopy (Figure 5F)."

sparser
"Mechanistically, USP22 interacted with FKBP12 to activate mTORC1 pathway through deubiquitylation."

sparser
"Moreover, we identified an interaction between USP22 and FKBP12."

sparser
"The interaction between endogenous FKBP12 and USP22 in SNU‐449 cells is shown in Figure  xref , while the co‐localization of FKBP12 and USP22 was observed by confocal microscopy (Figure  xref )."

sparser
"Mechanistically, USP22 interacted with FKBP12 to activate the mTORC1 pathway through deubiquitylation."

sparser
"Mechanistically, USP22 interacted with FKBP12 and stabilized it via deubiquitylation."
APC_C affects USP22
| 7
APC_C inhibits USP22.
| 5
APC_C inhibits USP22. 5 / 5
| 5

reach
"In addition, as described above, APC/C degrades USP22 through CDC20 during withdrawal from the M phase of cell cycle [ 25 ] (see Fig. 4 )."

reach
"In contrast, the APC and CDC20 E3 ligase complex negatively regulates USP22 activity by targeting it for degradation, presumably allowing CCNB1 downregulation so that cells can exit mitosis and enter anaphase."

reach
"USP22 is degraded by the APC/C E3 ubiquitin ligase complex, which also targets CCNB1 for destruction [XREF_BIBR, XREF_BIBR], thereby allowing cells to exit from mitosis, presumably by facilitating CCNB1 degradation."

reach
"Our study demonstrates that USP22 is also a substrate of the APC/C E3 ligase complex, which degrades USP22 during cell exit from M phase."

reach
"Therefore, APC and CDC20 E3 ligase complex promotes USP22 protein degradation, presumably allowing CCNB1 degradation for cells to exit M phase."
APC_C ubiquitinates USP22.
| 2
APC_C ubiquitinates USP22. 2 / 2
| 2

reach
"Fourth, USP22 is ubiquitinated and degraded by CDC20 containing APC/C complex during cell exit from M phase, presumably to release the brake on CCNB1 degradation."

reach
"Lastly, ubiquitylation of USP22 mediated by the anaphase promoter complex and cyclosome (APC/C) induces USP22 protein degradation during the cell cycle [XREF_BIBR]."

reach
"Moreover, USP22 promotes both HR (23) and NHEJ (48, 59)."

reach
"In this study we show that USP22 is necessary for HR, localizes at sites of DNA damage, and is necessary for recruitment of the PALB2-BRCA2-Rad51 complex through stabilizing PALB2 and BRCA2 protein levels at the translational level."

reach
"USP22 Is Necessary for Efficient HR."

reach
"USP22 directly interacts with PALB2 via the C-terminal WD40 domain to promote DNA homologous recombination repair (55)."

reach
"USP22 directly interacts with PALB2 through the latter's C-terminal WD40 domain to promote DNA homologous recombination repair [ 78 ]."

reach
"USP22 directly interacts with PALB2 to promote DNA homologous recombination repair and inhibit the killing effect of cisplatin on cancer cells."
USP22 affects XPC
1 | 3 2
USP22 binds XPC.
1 | 2
1 | 2

sparser
"An interaction between USP22 and XPC also occurred in the GEMM-derived MAFs, GFP and hUSP22 cells ( xref )."

sparser
"This interaction occurred in control cells as well, indicating that XPC and USP22 interact regardless of USP22 overexpression ( xref )."
USP22 activates XPC.
| 3
USP22 activates XPC. 3 / 3
| 3

eidos
"After treatment with irradiation , XPC foci were significantly increased in LN-USP22hi and C42-USP22hi cells as well as decreased in LN-shUSP22 and C42-shUSP22 compared to corresponding controls ( Figure 5G ) , suggesting that USP22 modulates XPC activity basally and in response to genotoxic insult ."

eidos
"Moreover , as Rad23B partners with XPC to sense and initiate NER , Rad23B foci formation was also increased basally and after irradiation in LN-USP22hi and C42-USP22hi cells ( Supplemental Figure 5F ) , further suggesting USP22 upregulation augments XPC activity ."

eidos
"USP22 induced deubiquitylation of XPC without an increase of the protein half-life ( Supplemental Figure 5A ) , suggesting the alternative hypothesis that USP22 may modulate XPC activity through de-polyubiquitylation , as XPC is known to be polyubiquitylated to promote efficient DNA repair [ 32,33,39 ] ."
USP22 affects TADA2B
4 | 2
4 | 2

sparser
"The association of USP22 with ADA2B was also compromised by the SANT domain triple mutations, especially the alanine substitutions."

sparser
"Association of USP22 with ADA2B was also compromised by the SANT domain triple mutations, especially with FLAG-3xHA-ADA2B W70A W90A Y110A protein."
USP22 affects SRSF9
| 6
| 6

sparser
"Furthermore, we analyzed potential candidate mRNAs that can be bound by SRSF9 using the ENCORI dataset () and found that SRSF9 could bind to USP22 mRNA."

sparser
"SRSF9 binds to USP22 mRNA to increase its expression."

sparser
"Through bioinformatic analysis via the ENCORI website, we predicted that SRSF9 could bind to USP22 mRNA."

sparser
"These results suggest that SRSF9 binds to USP22 mRNA to improve its stability."

sparser
"SRSF9 can bind to USP22 mRNA, increasing its stability."

sparser
"Furthermore, we demonstrated that SRSF9 can bind to USP22 mRNA and improve its stability."
USP22 affects SOS1
| 6
USP22 increases the amount of SOS1.
| 3
USP22 increases the amount of SOS1. 3 / 3
| 3

reach
"Indeed, USP22 overexpression induces significant elevations in protein levels of SOS1, RAS, p-ERK, p-PI3K, and p-AKT in both SGC7901 cells and tumors."

reach
"Lim et al. proposed that USP22 upregulates SOS1 expression in GC, thus leading to the activation of RAS/ERK and RAS/PI3K/AKT pathways [52]."

reach
"The results showed that overexpression of USP22 induced considerable upregulation of SOS1 expression in SGC7901 cells, accompanied by significant upregulation of RAS, p-ERK, c-myc, p-PI3K, and p-AKT (Fig. 5b, left panel)."
USP22 activates SOS1.
| 3
USP22 activates SOS1. 3 / 3
| 3

reach
"In this study, using a ChIP assay, we demonstrated, for the first time, that the overexpression of USP22 induced the upregulation of RAS activator SOS1 in SGC7901 cells."

reach
"USP22 overexpression in gastric cancer cells induces the upregulation of SOS1 and activation of the RAS/ERK and PI3K/AKT pathways."

reach
"Chromatin immunoprecipitation revealed that the overexpression of USP22 induced the upregulation of RAS activator son of sevenless 1 (SOS1) in SGC7901 cells."
USP22 affects RCOR1
| 6
| 4

reach
"Ultimately, we demonstrate that GSE1 is required for the recruitment of the deubiquitinase USP22 to the CoREST complex, potentially serving as a scaffold, and that loss of GSE1 inhibits deubiquitination of histone H2B at lysine 120, which was previously shown to facilitate γH2AX formation (11,35)."

reach
"Binding of USP22 to CoREST was independent of HDAC1 (Figure 6D) and also not affected by treatment with the HDAC inhibitor trichostatin A (TSA) (Supplementary Figure S6D)."

reach
"One of key requirements in this process seems to be the GSE1-dependent binding of USP22 to CoREST."

reach
"Of note, we observed that GSE1 is essential for the binding of USP22 to CoREST, suggesting a role as a scaffolder.Why has this multi-eraser complex gone unnoticed in previous research?"
GSE1 binds USP22 and RCOR1. 2 / 2
| 2

reach
"What is the biological function of the GSE1-USP22-CoREST complex?"

reach
"The GSE1-USP22-CoREST complex seems to be stable, as we do not see changes in its composition following DNA damage induction."
USP22 affects MYO1B
1 | 5
1 | 5

sparser
"MS analysis showed that USP22 interacted with MYO1B in H1299 cells."

sparser
"A previous MS study on HEK293T cells transfected with USP22 detected MYO1B as an interacting protein. [ xref ] MYO1B is a type of myosin and a motor protein involved in various biological processes, such as cell migration, cytoskeleton organization, and vesicle transport. [ xref ] Molecular docking simulations were performed to verify the interaction between USP22 and MYO1B (Figure xref )."

sparser
"Co‐IP experiments showed that USP22 interacted with MYO1B (Figure xref )."

sparser
"After identifying the interaction between USP22 and MYO1B, further verification was performed."

sparser
"The N‐terminal zinc finger domain of USP22 binds strongly to MYO1B, whereas the C‐terminal ubiquitin‐specific peptidase domain binds weakly to MYO1B (Figure  xref )."
USP22 affects EZH2
2 | 1 2 1
USP22 binds EZH2.
2 | 1 1
2 | 1 1

sparser
"About the possible relationship among BMI1 , EZH2 , and USP22 , it has been reported that USP22 increases BMI1 : in gastric cancer, USP22 inactivation decreases the neoplasm growth, while high levels of BMI1 accelerate the cell cycle via INK4a/ARF and AKT. xref Moreover, a reciprocal interaction between EZH2 and USP22 has been described in the sepsis‐induced myocardial dysfunction model: in this case, EZH2 represses USP22 through the H3K27me3 modification. xref In acute myeloid leukemia, the activation of BAX and the inhibition of BCL2 lead to a reduced expression of BMI1 and EZH2 . xref Finally, both BMI1 and EZH2 seem to be targets for the same miRNA, the miR200 that is a key regulator of the epithelial‐mesenchymal transition; indeed, a reduced expression of these genes reduces the migration capacity of colon cancer cells. xref "

reach
"18 Moreover, a reciprocal interaction between EZH2 and USP22 has been described in the sepsis‐induced myocardial dysfunction model: in this case, EZH2 represses USP22 through the H3K27me3 modification."
USP22 inhibits EZH2.
| 1 1
USP22 inhibits EZH2. 2 / 2
| 1 1

eidos
"EZH2 elevation aggravated the cardiac injury in SIMD rats , while USP22 upregulation inhibited the effect of EZH2 , which reduced the cardiac injury in SIMD rats ."

reach
"EZH2 elevation aggravated the cardiac injury in SIMD rats, while USP22 upregulation inhibited the effect of EZH2, which reduced the cardiac injury in SIMD rats."

reach
"USP22 Silencing Induces CIN Associated Phenotypes."

reach
"Furthermore, two-sample Kolmogorov-Smirnov (KS) tests revealed statistically significant increases (siUSP22-2, siUSP22-3) and a decrease (siUSP22-Pool) in cumulative nuclear area frequency distributions relative to siControl (XREF_FIG D; Table S5) that are consistent with reduced USP22 expression inducing CIN."

reach
"Having established that reduced USP22 expression induces CIN in short-term (< 1 week) siRNA based experiments, we now sought to determine the impact long-term USP22 depletion has on CIN."

reach
"In agreement with this possibility, we show that diminished USP22 expression induces CIN, highlighting a novel role for USP22 as a tumor suppressor that is essential to maintain mitotic fidelity and chromosome stability."

reach
"In this regard, while USP22 has been proposed as a novel therapeutic target based on its oncogenic functions [XREF_BIBR, XREF_BIBR, XREF_BIBR], our work suggests that USP22 inhibition will induce CIN that may promote cancer progression and/or the development of secondary malignancies."

reach
"Further, we identify USP22 as a novel CIN gene, indicating that USP22 deletions in tumors may drive CIN and contribute to oncogenesis."
USP22 affects CDH1
| 6
USP22 increases the amount of CDH1.
| 3
USP22 increases the amount of CDH1. 3 / 3
| 3

reach
"USP22 downregulation significantly increased the expression of E-cadherin (epithelial marker) but decreased the expression of N-cadherin and vimentin (mesenchymal markers) (XREF_FIG)."

reach
"Western blot analysis showed that USP22 knockdown in H1650 suppressed expression of the mesenchymal markers N-cadherin and vimentin and increased expression of the epithelial markers E-cadherin and al[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In this study, we showed that USP22 depletion significantly decreased the expressions of BMI-1, vimentin, and snail and increased E-cadherin expression in ATC cells."
USP22 decreases the amount of CDH1.
| 3
USP22 decreases the amount of CDH1. 3 / 3
| 3

reach
"IHC staining and WB results also showed that USP22 deficiency increased the protein expressions of E-cadherin and ZO-1, decreased the protein expression of α-SMA and Vimentin ( Fig. 10 C and D), and r[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In this study, we showed that USP22 depletion significantly decreased the expressions of BMI-1, vimentin, and snail and increased E-cadherin expression in ATC cells."

reach
"WB results further showed that USP22 overexpression downregulated E-cadherin and ZO-1 expression, upregulated Vimentin expression ( Fig. 6 C), and upregulated the expression of TGF-β1 and ECM componen[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects CCND1
| 6
USP22 ubiquitinates CCND1.
| 3
USP22 ubiquitinates CCND1. 3 / 3
| 3

reach
"While the screen confirmed known substrates of USP22, including histones H2A and H2B, the sole component of the core cell cycle machinery identified as a USP22 substrate in this screen was the G1 phase cyclin CCND1.Further analysis of the USP22CCND1 relationship revealed that USP22 directly reverses polyubiquitylation of CCND1 via a pathway distinct from protein-destabilizing GSK3β phosphorylation and ultimately protects CCND1 from proteasome-mediated degradation (16, 17)."

reach
"Control of CCND1 ubiquitylation by the catalytic SAGA subunit USP22 is essential for cell cycle progression through G1 in cancer cells."

reach
"Control of CCND1 ubiquitylation by the catalytic SAGA subunit USP22 is essential for cell cycle progression through G1 in cancer cells."
USP22 deubiquitinates CCND1.
| 3
USP22 deubiquitinates CCND1. 3 / 3
| 3

reach
"In addition to histones, USP22 deubiquitinates TRF1, CCNB1, CCND1, and SIRT1, thereby regulating involvement in metabolism, cycling, and apoptosis."

reach
"The elevated ubiquitylation and decreased steady-state levels of CCND1 seen after USP22 depletion suggest that deubiquitylation of CCND1 by USP22 may protect it from proteasomal degradation.Levels of both USP22 and CCND1 are elevated in human cancer, raising the possibility that elevated USP22 might protect CCND1 from degradation."

reach
"Collectively these findings support a model in which elevated expression of USP22 contributes to the aggressive growth of cancer cells in part via its ability to deubiquitylate and stabilize the rate-limiting cyclin CCND1, thereby promoting the G1–S transition.The identification of the USP22-CCND1 enzyme–substrate relationship may explain the previously reported phosphorylation-independent ubiquitylation and degradation of CCND1 (50)."
USP22 affects BCL2
| 6
USP22 increases the amount of BCL2.
| 4
USP22 increases the amount of BCL2. 4 / 4
| 4

reach
"We also noticed that overexpression of USP22 induced significant alterations in apoptosis-related BAD and bcl-2 expression."

reach
"Molecular analysis of the tumor tissues showed that USP22 knockdown reduced the levels of cyclin D2, Akt phosphorylation, vimentin, and Bcl-2, whereas upregulated the expressions of E-cadherin, Bax, and cleaved (cl)-caspase-3, which confirmed in vitro findings."

reach
"This may explain changes in BAD and bcl-2 expression in response to USP22 overexpression.To investigate whether SOS1 plays an essential role in mediating the tumorigenic roles of USP22 in gastric cancer, we silenced its expression in USP22-overexpressing SGC7901 cells."

reach
"USP22 knockdown also decreased the expression of Bcl-XL and Bcl-2 and increased the expression of cleaved-caspase 3 and cleaved-caspase 9."
USP22 activates BCL2.
| 2
USP22 activates BCL2. 2 / 2
| 2

reach
"The downregulation of USP1, USP8 and USP22 through their respective siRNA decreased Bcl-2 protein, as shown by western blot analysis."

reach
"Further analyses showed that USP22 silencing in HepG2 cells decreased the Bcl-2 and Bax ratio and enhanced the release of cytochrome c into the cytoplasm, suggesting the initiation of mitochondrial mediated apoptosis."
USP22 affects ABCC1
| 6
USP22 activates ABCC1. 6 / 6
| 6

eidos
"The downregulation of USP22 suppresses multidrug resistance-associated protein 1 ( MRP1 ) to induce intracellular sorafenib accumulation and hampers glycolysis of HCC cells ."

eidos
"] Downregulation of USP22 could inhibit the AKT / GSK-3beta pathway and further suppress the expression of MRP1 ( Figure 4a [ ."

eidos
"] Our study provided strong evidence that the downregulation of USP22 by Gal-SLPs suppressed the expression of MRP1 and caused high intracellular sorafenib accumulation ."

eidos
"In our previous study , USP22 bound to SIRT1 and subsequently activated the AKT pathway , increasing the expression of MRP1 to induce 5-FU resistance in HCC cells [ 15 ] ."

eidos
"iv ) The downregulation of USP22 not only suppresses multidrug resistance-associated protein 1 ( MRP1 ) , enhances the intracellular accumulation of sorafenib , and finally inhibits cell proliferation and cancer angiogenesis , but also inhibits glycolysis in hepatocellular carcinoma ( HCC ) cells , enhancing sorafenib chemosensitivity and impairing cell metabolism ."

eidos
"During sorafenib treatment , upregulation of USP22 increases ABCC1 expression and subsequently contributes to sorafenib resistance in HCC cells ."
TERF1 affects USP22
| 3 3
| 3 3

reach
"USP22 interacts with TRF1 and is required for TRF1 stability."

sparser
"USP22 interacts with TRF1 and is required for TRF1 stability."

reach
"For example, USP22 binding to TRF1, a protein that regulates telomere length, induces TRF1 protein stability."

sparser
"USP22 could bind to TRF1 and regulate telomere length (Atanassov et al ., xref )."

reach
"USP22 could bind to TRF1 and regulate telomere length (Atanassov etal., 2009)."

sparser
"For example, USP22 binding to TRF1, a protein that regulates telomere length, induces TRF1 protein stability."
TADA2B affects USP22
4 | 2
4 | 2

sparser
"The association of USP22 with ADA2B was also compromised by the SANT domain triple mutations, especially the alanine substitutions."

sparser
"Association of USP22 with ADA2B was also compromised by the SANT domain triple mutations, especially with FLAG-3xHA-ADA2B W70A W90A Y110A protein."
SRSF9 affects USP22
| 6
| 6

sparser
"Furthermore, we analyzed potential candidate mRNAs that can be bound by SRSF9 using the ENCORI dataset () and found that SRSF9 could bind to USP22 mRNA."

sparser
"SRSF9 binds to USP22 mRNA to increase its expression."

sparser
"Through bioinformatic analysis via the ENCORI website, we predicted that SRSF9 could bind to USP22 mRNA."

sparser
"These results suggest that SRSF9 binds to USP22 mRNA to improve its stability."

sparser
"SRSF9 can bind to USP22 mRNA, increasing its stability."

sparser
"Furthermore, we demonstrated that SRSF9 can bind to USP22 mRNA and improve its stability."
SOS1 affects USP22
| 6
SOS1 activates USP22. 6 / 6
| 6

reach
"These findings suggest that SOS1/RAS and downstream ERK and PI3K/AKT pathways mediate the oncogenic role of USP22 in gastric cancer."

reach
"Knockdown of SOS1 reverses the oncogenic effects of USP22 in gastric cancer cells."

reach
"Furthermore, SOS1 silencing could reverse the effects of USP22 on gastric cancer cell behavior and RAS signaling both in vitro and in vivo."

reach
"Furthermore, SOS1 upregulation in USP22-overexrpessing gastric cancer cells and xenograft tumor tissue is accompanied by activation of the RAS/ERK and RAS/PI3K/AKT pathways, which is attenuated by SOS1 silencing."

reach
"These results suggest that SOS1/RAS signaling mediates the oncogenic role of USP22 in gastric cancer."

reach
"Furthermore, our study indicated that SOS1 was upregulated in USP22-overexpressing gastric cancer cells and xenograft tumor tissue, accompanied by activation of the RAS/ERK and PI3K/AKT pathways, suggesting that SOS1/RAS signaling mediates the oncogenic role of USP22 in gastric cancer."
RCOR1 affects USP22
| 6
| 4

reach
"Ultimately, we demonstrate that GSE1 is required for the recruitment of the deubiquitinase USP22 to the CoREST complex, potentially serving as a scaffold, and that loss of GSE1 inhibits deubiquitination of histone H2B at lysine 120, which was previously shown to facilitate γH2AX formation (11,35)."

reach
"Binding of USP22 to CoREST was independent of HDAC1 (Figure 6D) and also not affected by treatment with the HDAC inhibitor trichostatin A (TSA) (Supplementary Figure S6D)."

reach
"One of key requirements in this process seems to be the GSE1-dependent binding of USP22 to CoREST."

reach
"Of note, we observed that GSE1 is essential for the binding of USP22 to CoREST, suggesting a role as a scaffolder.Why has this multi-eraser complex gone unnoticed in previous research?"
GSE1 binds USP22 and RCOR1. 2 / 2
| 2

reach
"What is the biological function of the GSE1-USP22-CoREST complex?"

reach
"The GSE1-USP22-CoREST complex seems to be stable, as we do not see changes in its composition following DNA damage induction."
PTEN affects USP22
| 3 3
PTEN binds USP22.
| 1 3
| 1 3

reach
"USP22 bound to PTEN and stabilized PTEN expression by decreasing its ubiquitination."

sparser
"USP22 bound to PTEN and stabilized PTEN expression by decreasing its ubiquitination."

sparser
"We found that PTEN might interact with USP22 (Fig.  xref , and xref )."

sparser
"Subsequently, the PTENUSP22 interaction was confirmed by endogenous immunoprecipitation analysis in both SW1990 and HPAC cells (Fig.  xref )."
PTEN activates USP22.
| 2
PTEN activates USP22. 2 / 2
| 2

reach
"PTEN overexpression reversed the promoting effect of USP22 knockdown on cell viability and the inhibitory effects of USP22 knockdown on apoptosis, oxidative stress, and LDH release rate in PC12 cells subjected to OGD/R."

reach
"Moreover, PTEN inhibition could also reverse the USP22‐mediated activation of the PI3K/Akt/mTOR pathway (Figure 5K)."
MYO1B affects USP22
1 | 5
1 | 5

sparser
"MS analysis showed that USP22 interacted with MYO1B in H1299 cells."

sparser
"A previous MS study on HEK293T cells transfected with USP22 detected MYO1B as an interacting protein. [ xref ] MYO1B is a type of myosin and a motor protein involved in various biological processes, such as cell migration, cytoskeleton organization, and vesicle transport. [ xref ] Molecular docking simulations were performed to verify the interaction between USP22 and MYO1B (Figure xref )."

sparser
"Co‐IP experiments showed that USP22 interacted with MYO1B (Figure xref )."

sparser
"After identifying the interaction between USP22 and MYO1B, further verification was performed."

sparser
"The N‐terminal zinc finger domain of USP22 binds strongly to MYO1B, whereas the C‐terminal ubiquitin‐specific peptidase domain binds weakly to MYO1B (Figure  xref )."
| 1 5
| 1 5

eidos
"Moreover , anaphase-promoting complex cell division cycle protein 20 ( APCCDC20 ) , an E3 ubiquitin ligase , promotes USP22 degradation in a cell cycle-dependent manner ( 61 ) ."

reach
"In contrast, the APC and CDC20 E3 ligase complex negatively regulates USP22 activity by targeting it for degradation, presumably allowing CCNB1 downregulation so that cells can exit mitosis and enter anaphase."

reach
"USP22 is degraded by the APC/C E3 ubiquitin ligase complex, which also targets CCNB1 for destruction [XREF_BIBR, XREF_BIBR], thereby allowing cells to exit from mitosis, presumably by facilitating CCNB1 degradation."

reach
"Our study demonstrates that USP22 is also a substrate of the APC/C E3 ligase complex, which degrades USP22 during cell exit from M phase."

reach
"To identify the E3 ligase that degrades USP22, we tested the interaction between USP22 and FBW7, CDC20, and CDH1, all of which are active during exit from mitosis [XREF_BIBR, XREF_BIBR]."

reach
"Therefore, APC and CDC20 E3 ligase complex promotes USP22 protein degradation, presumably allowing CCNB1 degradation for cells to exit M phase."
BRD4 affects USP22
3 | 1 2
3 | 1 2

sparser
"We further showed that USP22 specifically bound to BRD4 and promoted its deubiquitination in HEK239T and AML-12 cells."

reach
"We further showed that USP22 specifically bound to BRD4 and promoted its deubiquitination in HEK239T and AML-12 cells."

sparser
"These data suggest that BRD4 interacts with USP22 and is deubiquitinated by USP22 in ALD."

reach
"In light of these findings, it could be suggested that YAP1 silencing could reverse the protective effects of SPC on H/R-treated H9c2 cells.Altogether, our findings indicated that SPC up-regulated USP22, which stabilized KDM3A protein level via deubiquitination, and KDM3A inhibited H3K9me2 levels in the YAP1 promoter region to encourage YAP1 transcription, leading to reduction of MI/RI."

reach
"However, we failed to elucidate whether SPC could directly up-regulate USP22 to target KDM3A."

reach
"USP 22, KDM3A, and YAP1 were found to be down-regulated in MI/RI and SPC protected MI/RI rats, as evidenced by up-regulated expressions of USP22, KDM3A, and YAP1, reduced pathological injury in the myocardium, myocardial infarction areas, apoptosis, and levels of CK-MB, cTnI, and LDH, and enhanced H9c2 cell viability; while the protective effects of Sevo were counteracted by silencing of USP22, KDM3A, and SPC upregulated USP22, which stabilized KDM3A protein levels via deubiquitination, and KDM3A inhibited histone 3 lysine 9 di-methylation (H3K9me2) levels in the YAP1 promoter to encourage YAP1 transcription, to reduce MI/RI."
Sphingosine-1-phosphocholine increases the amount of USP22.
| 2
Sphingosine-1-phosphocholine increases the amount of USP22. 2 / 2
| 2

reach
"Lastly, rescue experimentation was carried out by inhibiting KDM3A or YAP1.3.1 SPC protects MI/RI rats and upregulates USP22 expression in the myocardium."

reach
"Collectively, our findings indicated that SPC ameliorated rat MI/RI and up-regulated USP22 expression levels in the myocardium."
Rcor1 affects USP22
| 5
| 5

sparser
"One of key requirements in this process seems to be the GSE1-dependent binding of USP22 to CoREST."

sparser
"While loss of GSE1 does not affect the histone deacetylation activity of CoREST, GSE1 appears to be essential for binding of the deubiquitinase USP22 to CoREST and for the deubiquitination of H2B K120 in response to DNA damage."

sparser
"Binding of USP22 to CoREST was independent of HDAC1 (Figure xref ) and also not affected by treatment with the HDAC inhibitor trichostatin A (TSA) ( xref )."

sparser
"However, none of these interactions were observed in GSE1 KO cells, showing that GSE1 is indeed required for the binding of USP22 to the CoREST complex (Figure xref )."

sparser
"Of note, we observed that GSE1 is essential for the binding of USP22 to CoREST, suggesting a role as a scaffolder."

reach
"In terms of underlying mechanisms, RNA sequencing and gene ontology enrichment analysis demonstrated that USP22 knockout significantly suppressed angiogenesis, proliferation, EMT, RAS, c-Myc pathways, concurrently enhanced oxidative phosphorylation and tight junction pathways in A549 and H1299 NSCLC cells."

reach
"Interestingly, we identified that USP22 promotes CSC properties and drug tolerance by supporting the oxidative phosphorylation program, known to be largely responsible for the poor response to conventional therapies in this particularly aggressive BC subtype."

reach
"Moreover, our findings demonstrate that USP22 epigenetically mediates the OXPHOS-dependent CSC- and drug-tolerant features, thereby, unveiling the potential therapeutic value of this DUB to combat the metabolic vulnerabilities of these particularly aggressive BC subtypes."

reach
"The USP22-specific DUBm of the SAGA complex mediates the OXPHOS gene expression program and respiratory capacity in TNBC."

reach
"Collectively, USP22-specific DUBm of the SAGA complex mediates the OXPHOS gene expression program and respiratory capacity in BC cells."

reach
"USP22 facilitated EPI-induced breast cancer progression and metastasis by enhancing adipose triglyceride lipase (ATGL)-mediated lipolysis."

reach
"This study has defined a USP22-mediated lipolysis circuit that links the chronic stress–induced EPI production to breast cancer pathogenesis through the AKT-USP22-FOXO1-ATGL pathway."

reach
"In addition to lactate and glucose metabolism, we show here that EPI promotes breast cancer metastasis through a USP22-mediated lipolysis circuit."

reach
"Future studies integrated with a psychologist team are needed to address this critical question.Our study revealed that USP22-mediated lipolysis circuit links the chronic stress–induced EPI production to breast cancer pathogenesis through the AKT-USP22-FOXO1-ATGL pathway across a wide range of human breast cancers, irrespective of their HER expression status."

reach
"Epinephrine promotes breast cancer metastasis through a ubiquitin-specific peptidase 22-mediated lipolysis circuit."
USP22 affects glucose
| 5
USP22 activates glucose. 5 / 5
| 5

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"The results showed that compared to the control group, USP22 expression knockdown significantly decreased glucose consumption, lactate production, and ATP generation in Sao-2 cells (p < 0.05)."

reach
"Overexpressing USP22 attenuated deleterious roles of HG in INS-1 cells; but its roles were reversed by up-regulating miR-144-3p."

reach
"In HG-treated NRK-52E cells, USP22-FLAG overexpression further increased USP22 expression ( Fig. 3 C), downregulated E-cadherin and ZO-1expression, and upregulated Vimentin expression ( Fig. 3 D)."

reach
"Cell-scratch test results showed that in HG-treated NRK-52E cells, USP22-FLAG overexpression significantly increased cell migration, whereas USP22–C185S-FLAG overexpression had no such function ( Fig.[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Moreover, Usp22 knockdown abolished high glucose stimulation–induced cell cycle arrest, apoptosis, and proinflammatory cytokine production in podocytes [138]."
| 5

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": not significant)and requests for resources and reagents should be directed to and 662 will be fulfilled by the corresponding author, Sjoerd J. L. van Wijk (vanWijk@med.uni-663 frankfurt.de; s.wijk@kinderkrebsstiftung-frankfurt.de)Profiling USP22-mediated gene expression in HT-29 hIECs."
| DOI

reach
"Profiling USP22-mediated gene expression in HT-29 hIECs."

reach
"In addition, USP22 was previously shown to modulate apoptosis in various cellular pathways to influence gene expression and protein transduction [42]."

reach
"The USP22-specific DUBm of the SAGA complex mediates the OXPHOS gene expression program and respiratory capacity in TNBC."

reach
"Collectively, USP22-specific DUBm of the SAGA complex mediates the OXPHOS gene expression program and respiratory capacity in BC cells."
USP22 affects Sirt1/JAK/STAT3
| 5
USP22 activates Sirt1/JAK/STAT3. 5 / 5
| 5

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"In NSCLC, miR-30e-5p suppressed tumorigenesis and metastasis by inhibiting USP22-mediated Sirt1/JAK/STAT3 signaling."

reach
"26 Xu et al suggested that miR-30e-5p suppresses NSCLC tumorigenesis by targeting ubiquitin carboxyl-terminal hydrolase 22 mediated Sirt1/JAK/STAT3 signaling."

reach
"Taken together, our findings suggest that miR-30e-5p reduces NSCLC tumorigenesis by targeting USP22-mediated Sirt1/JAK/STAT3 signaling."

reach
"miR-30e-5p, a member of the miR-30 family, regulates the Sirt1/JAK/STAT3 signaling mediated by USP22, serving as an inhibitor in NSCLC [18]."

reach
"It remains to be determined whether miR-30e-5p can reduce NSCLC tumorigenesis by inhibiting USP22-mediated regulation of MDMX and cyclooxygenase 2.Taken together, our findings indicate that miR-30e-5p[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects RAS
| 5
USP22 activates RAS. 5 / 5
| 5

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"SOS1 silencing also attenuated the USP22-induced activation of RAS/ERK and PI3K/AKT signaling both in vitro and in vivo."

reach
"USP22 overexpression in gastric cancer cells induces the upregulation of SOS1 and activation of the RAS/ERK and PI3K/AKT pathways."

reach
"Chromatin immunoprecipitation revealed that the overexpression of USP22 induced the upregulation of RAS activator son of sevenless 1 (SOS1) in SGC7901 cells."

reach
"Western blot analysis showed that USP22 overexpression also induced activation of the RAS/ERK and PI3K/AKT pathways in SGC7901 cells and xenograft tumor tissues."

reach
"In this study, using a ChIP assay, we demonstrated, for the first time, that the overexpression of USP22 induced the upregulation of RAS activator SOS1 in SGC7901 cells."
| 5
| 3

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"Conversely, ectopic expression of USP22 caused an enhancement in PD‐L1 protein expression, and further attenuating the upregulatory effect of Taz on PD‐L1 expression (Figure 3D,E)."

reach
"Conversely, ectopic USP22 expression enhanced the stability of PD‐L1 and prolonged its half‐life, as well as enhanced resistance to T‐cell cytotoxic effects and resulted in an increase in cell viability (Figure 4D–F; Figure S4B, Supporting Information)."

reach
"For example, in hepatocellular carcinoma USP22 upregulates PD‐L1 and induces the exhaustion of tumor infiltrating CD8+ T cells, 32 while in the colorectal carcinoma it targets BMI‐1, c‐MYC, CYCLIN D2, p16INK4a, and p14ARF, so exerting a crucial role in tumor proliferation."
USP22 deubiquitinates Parkinson Disease.
| 2
USP22 deubiquitinates Parkinson Disease on L1. 2 / 2
| 2

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"56 USP22 physically interacts with the C‐terminus of the PD‐L1 protein and CSN5, a known DUB of PD‐L1, therefore deubiquitinates and stabilizes PD‐L1."

reach
"For instance, USP22 can enhance the deubiquitination and stabilization of PD‐L1 in cancer cells, suggesting its protumorigenic role in cancer [31]."
USP22 affects PI3K/Akt/mTOR
| 5
USP22 activates PI3K/Akt/mTOR. 5 / 5
| 5

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"32 , 33 , 34 To determine whether USP22 activates the PI3K/Akt/mTOR pathway via SIRT1‐mediated PTEN deacetylation, we used siRNA to silence the expression of SIRT1 in USP22‐overexpressing melanoma cells."

reach
"Moreover, PTEN inhibition could also reverse the USP22‐mediated activation of the PI3K/Akt/mTOR pathway (Figure 5K)."

reach
"These findings suggested that USP22 activates the PI3K/Akt/mTOR pathway and induces melanoma EMT via the SIRT1/PTEN axis.2.6 Drug screening identifies topotecan as a USP22-targeting molecule to suppress melanoma metastasis."

reach
"These results demonstrated that USP22 might promotes melanoma metastasis by inducing EMT activation.2.4 USP22 potentiates melanoma metastasis and EMT through activating PI3K/Akt/mTOR pathway."

reach
"These results suggested that USP22 potentiates melanoma metastasis and EMT through activating the PI3K/Akt/mTOR pathway.2.5 USP22 activates PI3K/Akt/mTOR pathway via SIRT1/PTEN axis."
USP22 affects PI3K
| 5
USP22 activates PI3K. 5 / 5
| 5

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"USP22 overexpression in gastric cancer cells induces the upregulation of SOS1 and activation of the RAS/ERK and PI3K/AKT pathways."

reach
"Western blot analysis showed that USP22 overexpression also induced activation of the RAS/ERK and PI3K/AKT pathways in SGC7901 cells and xenograft tumor tissues."

reach
"Previous study also showed that USP22 promotes cell cycle progression by positively regulating the PI3K and Akt pathway [XREF_BIBR]."

reach
"These results suggest that USP22 downregulation inhibits OS cells by suppressing the PI3K and Akt pathway."

reach
"To further dissect the precise mechanism by which USP22 activates the PI3K/Akt signaling pathway, coimmunoprecipitation (Co‐IP) and mass spectrometry (MS) were utilized to identify the potential direct binding partners of USP22."
USP22 affects NT5E
4 | 1
4 | 1

sparser
"In breast cancer, the deubiquitinating enzyme USP22 interacts with CD73, inhibiting CD73 ubiquitination and proteasomal degradation, thereby stabilizing its protein levels."
USP22 affects NLRP3
| 1 4
USP22 inhibits NLRP3. 5 / 5
| 1 4

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"In NLRP3 inflammasome-related diseases, USP22 inhibits NLRP3 inflammatory bodies through Atg5-dependent autophagy to degrade NLRP3 [23]."

reach
"USP22 in macrophages promotes LRR and PYD structural domain protein 3 (NLRP3) degradation and attenuates inflammation through autophagy-associated 5 homolog (ATG5)-mediated autophagy ."

reach
"Here, we demonstrated that USP22 (ubiquitin specific peptidase 22) promotes NLRP3 degradation and inhibits NLRP3 inflammasome activation."

eidos
"Here , we demonstrated that USP22 ( ubiquitin specific peptidase 22 ) promotes NLRP3 degradation and inhibits NLRP3 inflammasome activation ."

reach
"USP22 suppresses the NLRP3 inflammasome by degrading NLRP3 via ATG5-dependent autophagy."
USP22 affects KDELR1
| 5
| 5

sparser
"USP22 Interacts with KDELR1 and Stabilizes KDELR1 Expression via Deubiquitination."

sparser
"We confirmed the interaction between USP22 and KDELR1 using molecular docking simulations and co‐IP assays (Figure  xref )."

sparser
"These results confirmed that the C19 ubiquitin‐specific peptidase domain of USP22 was the dominant mediator of the interaction between USP22 and KDELR1."

sparser
"The interaction between USP22 and KDELR1 was validated using a PLA assay (Figure  xref )."

sparser
"Therefore, we hypothesized that the USP22KDELR1 axis regulates the invadopodia process to drive LUAD metastasis."
USP22 affects IPO7
| 4 1
| 4 1

sparser
"Together, the above results indicate that Importin–7–AR–USP22 might form a relatively stable complex, even if AR and USP22 may competitively bind to Importin-7."

reach
"Mechanistically, through co-immunoprecipitation, immunofluorescence, and nuclear-cytoplasmic protein separation experiments, we discovered that AR and USP22 can bind to Importin-7 as cargoes to promote BC progression."

reach
"Importin-7 binds to AR and USP22."

reach
"In addition, our immunofluorescence analysis showed that Importin-7 binds to AR or USP22 in both the nucleus and cytoplasm (Fig. 4C–E), suggesting that Importin-7 may regulate the nuclear translocation of AR and USP22.Next, we conducted a more in-depth study of the Importin-7–AR–USP22 complex by the treatment of AR inhibitor (enzalutamide) or AR ligands, including DHT and synthetic androgen R1881."

reach
"Together, the above results indicate that Importin–7–AR–USP22 might form a relatively stable complex, even if AR and USP22 may competitively bind to Importin-7."
USP22 affects Death
| 5
USP22 inhibits Death. 5 / 5
| 5

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"A recent study shows that the expression of USP22 protein, a kind of deubiquitinase, can inhibit ferroptotic cardiomyocyte death through the SIRT1/p53/SLC7A11 axis and further protect against myocardial ischemia/reperfusion injury [64]."

reach
"Overexpression of USP22 could inhibit ferroptotic cardiomyocyte death to protect against IRI (102)."

reach
"Ma et al. found that ubiquitin-specific peptidase 22 (USP22) inhibits cardiomyocyte death induced by ferroptosis in myocardial I/R injury via the SIRT1/p53/SLC7A11 axis, providing a novel therapeutic target for the treatment [83]."

reach
"Previous studies have shown that USP22 is widely expressed in various tissues and that USP22 gene deletion can lead to the early death of mouse embryos, whereas heterozygous embryos display stunted gr[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"According to a recent study, USP22 may prevent cardiac ischemia‒reperfusion damage by preventing cardiomyocyte death (Ma S et al., 2020)."
USP22 affects DNA repair
| 5
| 5

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"It is likely that changes in USP22 and RNF40 status due to altered levels of m A may direct the DNA repair machinery to employ or not employ HR or NHEJ to repair damaged DNA."

reach
"USP22 can also enhance the telomerase reverse transcriptase (TERT)/p53 signaling pathway [ 65 ] and promote cell proliferation and DNA repair (see Fig. 1 )."

reach
"Furthermore, USP22 can regulate DNA repair events by targeting the deubiquitination of proteins other than H2A (22, 23, 37, 45), it is possible that in addition to H2Aub, other USP22 target proteins may contribute to ALKBH5’s DNA repair and oncogenic functions."

reach
"Unlike the mechanism of 5-FU drug resistance, USP22 enhances a tumor's DNA repair ability and decreases its sensitivity to cisplatin."

reach
"Unlike 5-FU resistance mechanisms, USP22 increases tumor resistance to cisplatin by enhancing DNA repair capacity."

reach
"For example, USP22 was reported to promote vascularization in the mouse placenta and aberrant angiogenesis in hepatocellular carcinoma and non-small cell lung cancer [43–45]."

reach
"USP22 promotes non-small cell lung cancer and retinoblastoma tumorigenesis via p53 inhibition [74,75] ."

eidos
"As reported previously , USP22 promotes the proliferation of non-small cell lung cancer by regulating the ubiquitination of COX-221 ."
USP22 affects CDK11B
| 3 2
USP22 activates CDK11B.
| 3
USP22 activates CDK11B. 3 / 3
| 3

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"To further validate that USP22 increases hepatocellular carcinoma cell proliferation by stabilising CDK11B protein, we overexpressed His‐CDK11B in USP22 knockout cells (Figure 4F) and evaluated cell proliferation."

reach
"19 However, there has been a lack of thorough investigation into the role of USP22 in hepatocellular carcinoma development and Sorafenib resistance.In this study, we demonstrate that USP22 promotes the proliferation of hepatocellular carcinoma cells by stabilising cyclin‐dependent kinase 11B (CDK11B)."

reach
"31 To establish the relationship between the promotion of cell proliferation and the stabilisation of CDK11B protein caused by USP22 overexpression, we intended to first examine the effects of CDK11B knockdown on hepatocellular carcinoma cell proliferation."
USP22 binds CDK11B.
| 2
| 2

sparser
"USP22 interacts with CDK11B."

sparser
"Taken together, these results indicated that USP22 interacts with CDK11B in hepatocellular carcinoma cells."
USP22 affects BRCA2
| 5
USP22 increases the amount of BRCA2. 5 / 5
| 5

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"Lastly, we show USP22 positively regulates BRCA2 and PALB2 levels at the protein level."

reach
"Our study reveals USP22 positively regulates PALB2 and BRCA2 levels at the translational level."

reach
"USP22 modulates PALB2 and BRCA2 levels to promote chemoresistance in lung adenocarcinoma."

reach
"To determine whether USP22 is modulating PALB2 and BRCA2 protein levels at the transcriptional level we performed qPCR of BRCA2 and PALB2 with and without USP22 knockdown after 48 hours (Figure S1b)."

reach
"To further validate USP22 was modulating and stabilizing BRCA2 and PALB2 levels at the translational level, cells were depleted of USP22 again with and without treatment of the proteasome inhibitor MG132 to see if this could rescue PALB2 and BRCA2 levels in a USP22 knockdown background (Figure S1c)."
SOX9 affects USP22
| 5
SOX9 increases the amount of USP22.
| 3
SOX9 increases the amount of USP22. 3 / 3
| 3

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"In this study, we reported that ZRANB1 enhanced the features of CSCs and accelerated tumor progression by regulating the stability of Sox9 protein, which then mediated USP22 transcription and the acti[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"The upregulation of Sox9 induced mRNA expression of USP22 in CRC cells, whereas Sox9 knockdown led to significantly decreased USP22 expression compared to the control cells ( Fig. 5 A )."

reach
"In HCT116 and SW620 cells, the downregulation of ZRANB1 significantly suppressed the expression of USP22, while the overexpression of Sox9 restored the mRNA and protein levels of USP22 in CRC cells ( [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
SOX9 decreases the amount of USP22.
| 2
SOX9 decreases the amount of USP22. 2 / 2
| 2

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"The upregulation of Sox9 induced mRNA expression of USP22 in CRC cells, whereas Sox9 knockdown led to significantly decreased USP22 expression compared to the control cells ( Fig. 5 A )."

reach
"Consistently, the protein level of USP22 in cells overexpressing Sox9 was markedly elevated compared to the control group, while the downregulation of Sox9 effectively inhibited the expression of USP2[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
PRDM1 affects USP22
| 5
PRDM1 increases the amount of USP22. 5 / 5
| 5

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"Mechanistically, PRDM1 enhances USP22 transcription, thus reducing SPI1 protein degradation through deubiquitination, which enhances PD-L1 transcription."

reach
"It was recently shown that BLIMP1 enhances transcription of USP22 deubiquitinating enzyme leading to decreased degradation of SPI1 transcription factor and subsequent enhanced expression of programmed death ligand 1 (PD-L1), which leads to infiltrated CD8+ T cell exhaustion and memory responses [3]."

reach
"PRDM1/BLIMP1 regulates USP22 transcription."

reach
"PRDM1 overexpression only increased the mRNA and protein levels of USP22, whereas PRDM1 knockout decreased the mRNA and protein levels of USP22 (Fig. 4g, h and Supplementary Fig. 4g)."

reach
"This upregulation may be related to the enhancement of USP22 transcription level by PRDM1, which decreases SPI1 protein degradation through ubiquitination and consequently increases PD-L1 expression [49]."
NT5E affects USP22
4 | 1
4 | 1

sparser
"In breast cancer, the deubiquitinating enzyme USP22 interacts with CD73, inhibiting CD73 ubiquitination and proteasomal degradation, thereby stabilizing its protein levels."
KDELR1 affects USP22
| 5
| 5

sparser
"USP22 Interacts with KDELR1 and Stabilizes KDELR1 Expression via Deubiquitination."

sparser
"We confirmed the interaction between USP22 and KDELR1 using molecular docking simulations and co‐IP assays (Figure  xref )."

sparser
"These results confirmed that the C19 ubiquitin‐specific peptidase domain of USP22 was the dominant mediator of the interaction between USP22 and KDELR1."

sparser
"The interaction between USP22 and KDELR1 was validated using a PLA assay (Figure  xref )."

sparser
"Therefore, we hypothesized that the USP22KDELR1 axis regulates the invadopodia process to drive LUAD metastasis."
ADAM9 affects USP22
| 2 3
| 2 3

sparser
"To clarify the interaction between USP22 and ADAM9, Co-IP assay was performed."

sparser
"Besides that, our findings also verified that USP22 interacted with ADAM9 to mediate the activation of Wnt/β-catenin pathway in trophoblast cells."

sparser
"In addition, we also found USP22 bound to ADAM9 to suppress the activation of Wnt/β-catenin pathway."

reach
"In addition, we also found USP22 bound to ADAM9 to suppress the activation of Wnt/β-catenin pathway."

reach
"To clarify the interaction between USP22 and ADAM9, Co-IP assay was performed."
MiR-4490 affects USP22
| 4
MiR-4490 activates USP22. 4 / 4
| 4

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"Together, these results indicate that miR-4490 may inhibit GC metastasis and EMT by targeting USP22."

reach
"USP22 can also be modulated by the POU2F1/miR-4490 axis to further increase GC proliferation and metastasis [51]."

reach
"Although werecently developed an algorithm to predict targeting of USP22 by miR-4490, it remains to be established whether this miRNA participates in the regulation of USP22 expression in GC."

reach
"These findings indicate that miR-4490 targets a complementary sequence in the USP22 3 '-UTR."
| 2

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"A previous study demonstrated that USP22 could inhibit the transcription of the cyclin-dependent kinase inhibitor 1A (p21) gene by deubiquitinating the transcriptional regulator fructose-1,6-bisphosphatase (FBP1) (5)."

reach
"Recent studies have demonstrated that USP22 can inhibit the transcription of the p21 gene by de-ubiquitinating the transcriptional regulator FBP1, leading to cell proliferation and tumorigenesis [XREF_BIBR]."
USP22 deubiquitinates transcriptional regulator.
| 2

reach
"Furthermore, USP22 regulates cell proliferation by deubiquitinating the transcriptional regulator, FBP1 (5)."

reach
"A previous study demonstrated that USP22 could inhibit the transcription of the cyclin-dependent kinase inhibitor 1A (p21) gene by deubiquitinating the transcriptional regulator fructose-1,6-bisphosphatase (FBP1) (5)."
USP22 affects miR-34b
| 4
USP22 activates miR-34b. 4 / 4
| 4

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"To confirm that USP22 is a direct target of miR-34b, we cloned the 3 '-UTR of USP22 into a reporter plasmid downstream of the luciferase gene."

reach
"Overexpression of USP22 attenuates the inhibitory effect of miR-34b on NPCcell proliferation."

reach
"Overexpression of USP22 attenuates the inhibitory effect of miR-34b on NPC cell viability and proliferation, thus confirming our hypothesis that miR-34b inhibits NPC proliferation by downregulating USP22."

reach
"MTT (XREF_FIG), cell cycle (XREF_FIG), and colony formation analyses (XREF_FIG) show that overexpression of USP22 can significantly attenuate the suppressive effect of miR-34b."
USP22 affects miR-132-3p
| 4
USP22 activates miR-132-3p. 4 / 4
| 4

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"USP22 Was a Target of miR-132-3p in CRC."

reach
"In terms of mechanism, our results revealed that USP22 was a target of miR-132-3p."

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"Overexpressed USP22 Restored the Inhibition Effects of miR-132-3p on CRC Cell Progression."

reach
"Besides, USP22 was a target of miR-132-3p, and its overexpression restored the inhibition effect of miR-132-3p mimic on CRC cell progression."
USP22 affects integrin b1
| 4
USP22 increases the amount of integrin b1. 4 / 4
| 4

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"Consistently, re-expression of WT USP22, but not its mutants restored integrin b1 expression in USP22-null breast cancer cells (Fig. 3N & 3O)."

reach
"USP22 promotes integrin b1 expression through FoxM1 stabilization.."

reach
"These results support our hypothesis that USP22 specifically promote integrin b1 expression through FoxM1 stabilization."

reach
"USP22 appears to positively regulate integrin b1 expression at the transcriptional level because its targeted deletion resulted in a more than 70% reduction in ITGB1 mRNA levels (Fig. 2E)."
USP22 affects TFRC
| 4
USP22 increases the amount of TFRC.
| 2
USP22 increases the amount of TFRC. 2 / 2
| 2

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"These findings collectively demonstrated that USP22 negatively regulates ferroptosis in hepatocellular carcinoma cells.2.6 USP22 modulates cellular H2BK120ub level and the transcription of TFRC ."

reach
"In summary, these findings suggested that USP22 knockout enhances the occupation of H2BK120ub on the TSS downstream region of TFRC, thereby promoting TFRC transcription.2.7 USP22 inhibits Sorafenib-induced ferroptosis through downregulating TFRC transcription."
USP22 decreases the amount of TFRC.
| 2
USP22 decreases the amount of TFRC. 2 / 2
| 2

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"In addition, IHC staining of Ki‐67, H2BK120ub and TFRC in frozen sections from all four groups indicated that USP22 knockout promoted H2BK120ub and TFRC levels in transplanted tumours, and inhibited hepatocellular carcinoma malignancy as indicated by Ki‐67 staining, especially after Sorafenib treatment (Figure 8G‒J)."

reach
"Likewise, the increase of GPX4 protein level, as well as the decrease of TFR1 protein level that caused by USP22 OE in HG-induced INS-1 cells were also partly abrogated by miR-144-3p mimic (Fig. 3L)."
USP22 affects SP1
| 4
| 4

sparser
"For example, SP1 and protein kinase A/cAMP response element-binding protein could bind to the USP22 promoter to suppress or promote USP22 transcription, respectively ( xref , xref )."

sparser
"Interaction between Sp1 and the USP22 Promoter."

sparser
"Immunoprecipitation of cross-linked chromatin from HFL1 cells with an anti-Sp1 antibody followed by PCR amplification of the region (the sequence between-210 and +52) confirmed that the endogenous Sp1 protein does bind to this region of the USP22 promoter in HFL1 ( xref )."

sparser
"Results showed that Sp1 binds to the USP22 promoter in fibroblasts, suggesting that Sp1-DNA interaction may be required for USP22 repression."
USP22 affects PTEN
| 1 3
| 1 3

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"USP22 bound to PTEN and stabilized PTEN expression by decreasing its ubiquitination."

sparser
"USP22 bound to PTEN and stabilized PTEN expression by decreasing its ubiquitination."

sparser
"We found that PTEN might interact with USP22 (Fig.  xref , and xref )."

sparser
"Subsequently, the PTENUSP22 interaction was confirmed by endogenous immunoprecipitation analysis in both SW1990 and HPAC cells (Fig.  xref )."

reach
"Additionally, USP22 promotes CSC maintenance through the Wnt/β-catenin pathway (81)."

reach
"Interestingly, we identified that USP22 promotes CSC properties and drug tolerance by supporting the oxidative phosphorylation program, known to be largely responsible for the poor response to conventional therapies in this particularly aggressive BC subtype."

reach
"In addition, USP22 promotes CSC maintenance through the Wnt/β-catenin pathway (37)."

reach
"In this study, we demonstrated that USP22 mediated protein stabilization of BMI1 promotes gastric CSC stemness maintenance and GC progression, thereby providing a rationale for USP22 targeting as a potential therapeutic approach against GC."
USP22 affects MMP9
| 4
USP22 activates MMP9. 4 / 4
| 4

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"USP22 and STAT3 co-depletion partly rescued the MMP9 proteolytic activity and invasion of SW480 cells, compared with that of STAT3 depletion alone."

reach
"In colon cancer, USP22 was reported to attenuate the invasion capacity of colon cancer cells by inhibiting the STAT3 and MMP9 signaling pathway."

reach
"Other inflammatory mediators secreted by PDA cells include granulocyte colony-stimulating factor (G-CSF) (41), IL-6 (42), IL-1α (43), IL-1β (44, 45), ubiquitin specific peptidase 22 (USP22) (46), C-X-C motif chemokine ligand 8 (CXCL8) (47), matrix metallopeptidase 9 (MMP-9) and indoleamine-2,3-dioxygenase (IDO) (48), which all contribute to the establishment of immunosuppressive TME in pancreatic cancer ( Figure 1 )."

reach
"In conclusion, USP22 attenuated the invasion capacity of colon cancer cells by inhibiting the STAT3 and MMP9 signaling pathway."
USP22 affects MED1
| 4
USP22 binds MED1. 4 / 4
| 4

sparser
"The interaction between the endogenous USP22 and MED1 was further validated in mouse primary iNKT cells because MED1 was detected in anti-USP22 immunoprecipitates but not the normal rabbit IgG controls ( xref ), indicating a possibility that USP22 regulates iNKT cell development through, at least partially, MED1 interaction."

sparser
"Since USP22 interacts with MED1, both of which are transcription coactivators involved in promoting T-bet and IL-2R gene transcription in iNKT cells, we then asked whether MED1 binds to the promoters of T-bet and IL-2Rβ in a USP22-dependent manner, or vice versa."

sparser
"However, assessment of ER stress‐induced changes in MED1 and USP22 binding shows poor correlation (Fig  xref )."

sparser
"USP22 interacts with MED1 in iNKT cells."
USP22 affects KDM3A
| 2 2
| 2 2

sparser
"Accordingly, we speculated whether USP22 could influence KDM3A expression and detected the interaction between USP22 and KDM3A using a ChIP assay, and found that USP22 could indeed interact with KDM3A ( xref )."

reach
"Accordingly, we speculated whether USP22 could influence KDM3A expression and detected the interaction between USP22 and KDM3A using a ChIP assay, and found that USP22 could indeed interact with KDM3A (Figure 5a)."

reach
"The interaction between USP22 and KDM3A and between KDM3A and YAP1 was further validated."

sparser
"The interaction between USP22 and KDM3A and between KDM3A and YAP1 was further validated."
USP22 affects Interferon
| 4
| 4

reach
"However, USP13, USP19, and USP22 inhibit virus-induced IFN production by removing K27-linked polyubiquitin chains from STING (40, 81) or TRIF (90)."

reach
"In mechanism, USP22 enhances the stability of STAT1 via deubiquitination and promotes the activation of IFNγ stimulation in melanoma [ 43 ]."

reach
"18 Besides, USP22 in melanoma cells stabilizes STAT1 and promotes the interferon pathway, which improves the cytotoxic effects of CD8 T cells."

reach
"Increased USP22-dependent upregulation of IFN signaling in Caco-2 cells was also reflected in the elevated expression of the antiviral ISGs IRF9 and OAS3 (Fig. 6B)."
USP22 affects Integrins
| 4
USP22 increases the amount of Integrins. 4 / 4
| 4

reach
"In addition to integrin β1, USP22 appears to promote the transcription of several additional integrin family members."

reach
"Notably, CRISPR-mediated deletion of USP22 completely eliminates HGF-induced integrin expression in melanoma cells."

reach
"In addition to integrin b1, USP22 appears to promote the transcription of several additional integrin family members."

reach
"USP22 promotes integrin beta1 expression through FoxM1 stabilization."
USP22 affects IL1B
| 4
USP22 increases the amount of IL1B. 4 / 4
| 4

reach
"In addition, the rescue experiment revealed that USP22 siRNA administration decreased IL-6 and activated IL-1β levels, which were unchanged by BRD4 siRNA treatment."

reach
"To verify this inference, we further showed that USP22 overexpression increased the expression of IL-6 and activated IL-1β, which was attenuated by BRD4 siRNA ( Fig. 4 E)."

reach
"Conversely, the protein levels of IL-6 and activated IL-1β were obviously increased by USP22 overexpression in AML-12 cells ( Fig. 2 K)."

reach
"In addition, USP22 overexpression increased the levels of the inflammatory factors IL-6 and activated IL-1β."
USP22 affects Hypoxia
| 4
USP22 activates Hypoxia. 4 / 4
| 4

reach
"19 Moreover, current evidence suggests that USP22 could promote hypoxia‐induced HCC glycolysis and stemness by deubiquitinating and stabilizing HIF‐1α."

reach
"In addition, USP22 can promote hypoxia-induced generation of liver cancer stem cells through the HIF1α/USP22 positive feedback loop after p53 inactivation, and lipoprotein complexes targeting USP22 can inhibit the growth of liver cancer cells and enhance sorafenib sensitivity (77)."

reach
"27 Ubiquitin‐specific protease 22 (USP22) can promote hypoxia‐induced hepatocellular carcinoma (HCC) stemness and glycolysis by deubiquitinating and stabilizing HIF1α."

reach
"23 In the absence of TP53, we found that USP22 enhanced hypoxia‐induced stemness, 13 which facilitated us to further investigate the relationship between USP22 and hepatotumorigenesis."
USP22 affects Fibrosis
| 4
USP22 inhibits Fibrosis.
| 2
| 2

reach
"USP22 in podocytes promotes diabetic nephropathy by enhancing oxidative stress [138], while USP22 in mesangial cells suppresses fibrosis through the stabilization of Sirt-1 [135]."

reach
"Addition of USP22 reduces Ang II-induced cardiac inflammation and fibrosis."
USP22 activates Fibrosis.
| 2
| 2

reach
"In the current study, we show that the overexpression of USP22 induced by treatment with FO greatly improved Ang II-induced cardiac dysfunction, heart hypertrophy, inflammation, and fibrosis (Figures 1–4)."

reach
"OTUD1 and USP22 have been shown to promote fibrosis and injury in renal tubular epithelial cells ."
USP22 affects FN1
| 4
USP22 increases the amount of FN1.
| 2
USP22 increases the amount of FN1. 2 / 2
| 2

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"Ubiquitin specific protease 22 (USP22) reduces the degradation of sirtuin-1 and the expression of FN and TGF-beta1 in AGE treated GMCs, whereas depletion of USP22 promotes sirtuin-1 degradation and the expression of FN and TGF-beta1 in this cell model."

reach
"Ubiquitin specific protease 22 (USP22) reduced Sirt1 ubiquitination and degradation and decreased FN and TGF-beta1 expression in GMCs under both basal and AGEs treated conditions."
USP22 decreases the amount of FN1.
| 2
USP22 decreases the amount of FN1. 2 / 2
| 2

reach
"Ubiquitin specific protease 22 (USP22) reduced Sirt1 ubiquitination and degradation and decreased FN and TGF-beta1 expression in GMCs under both basal and AGEs treated conditions."

reach
"Ubiquitin specific protease 22 (USP22) reduces the degradation of sirtuin-1 and the expression of FN and TGF-beta1 in AGE treated GMCs, whereas depletion of USP22 promotes sirtuin-1 degradation and the expression of FN and TGF-beta1 in this cell model."
USP22 affects ERK
| 3 1
USP22 activates ERK. 4 / 4
| 3 1

reach
"Western blot analysis showed that USP22 overexpression also induced activation of the RAS/ERK and PI3K/AKT pathways in SGC7901 cells and xenograft tumor tissues."

sparser
"USP22 can activate extracellular signal–regulated kinase 1/2 (ERK1/2) to promote the conversion of microtubule-associated protein 1A/1B-light chain 3 (LC3) to the lipid-modified form of LC3 (LC3-II) i[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Activation of ERK1/2 kinase rather than AKT1 by USP22 was found to be one of the mechanisms promoting LC3 processing."

reach
"USP22 overexpression in gastric cancer cells induces the upregulation of SOS1 and activation of the RAS/ERK and PI3K/AKT pathways."
USP22 affects ERBB2
| 4
USP22 activates ERBB2. 4 / 4
| 4

reach
"USP22 promotes HER2 driven mammary carcinoma aggressiveness by suppressing the unfolded protein response."

reach
"The current work provides evidence that the USP22 deubiquitinase, part of the DUBm of the SAGA complex, supports the mitochondrial biogenesis transcriptomic program to foster the CSC traits and promote the drug resistant features of HER2 -BC and TNBC cells (Fig. 7)."

reach
"To determine whether Usp22 overexpression might enhance ERBB2-driven formation of mammary tumors, we introduced our previously described Lox-Stop-Lox (LSL) Usp22 allele [17] into mice carrying the MMTV-NIC transgene (Fig 1A)."

reach
"In conclusion, USP22 strongly supports the drug-tolerant behavior of HER2 -BC and TNBC and is a promising therapeutic target to improve standard as well as OXPHOS-based therapies (Fig. 7E)."
USP22 affects CRC
| 4
USP22 activates CRC. 4 / 4
| 4

reach
"However, USP22 knockdown diminished the tumorigenic effects of ZRANB1 in CRC as evidenced by reduced Wnt/β-catenin activity, downregulated stem cell markers, weakened viability and sphere-forming capa[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"He et al (178) concluded that SNHG16 could activate USP22 expression to promote CRC progression via absorbing miR-132-3p."

reach
"However, the mechanism by which USP22 promotes CRC metastasis remains largely unknown."

reach
"In addition, Usp22 promotes CRC by stabilizing cyclin B1."
USP22 affects COL17A1
| 4
USP22 deubiquitinates COL17A1. 4 / 4
| 4

reach
"Mechanistically, USP22 stabilized and upregulated COL17A1 by enhancing the deubiquitination of COL17A1."

reach
"In addition, the reduced expression of the ferroptosis suppressor GPX4 was observed in sh‐COL17A1 xenograft tumors (Figure 4D).3.5 USP22 Promotes the Deubiquitination of COL17A1 to Stabilize COL17A1."

reach
"In this paper, we establish the fact, for the first time, that USP22 stabilizes COL17A1 by deubiquitinating COL17A1 in LUAD cells, highlighting a novel deubiquitination mechanism underlying COL17A1 upregulation in LUAD."

reach
"More intriguingly, in A549 LUAD cells, immunoblot analysis with an antibody against ubiquitin (UB) after IP experiments using the anti‐COL17A1 antibody revealed that USP22 silencing resulted in a distinct increase in ubiquitinated COL17A1 protein level and COL17A1 degradation (Figure 5M), suggesting that USP22 induces the deubiquitination of the COL17A1 protein in LUAD cells."
USP22 affects AP4
| 4
USP22 binds AP4. 4 / 4
| 4

sparser
"In addition, USP22 binds to the promoter region of AP4 to activate its transcription."

sparser
"Consistent with the above results, ChIP analysis revealed that USP22 binds the promoter region of AP4 (Figure xref )."

sparser
"We also found that USP22 binds to the promoter region of AP4 to mediate its transcription and thus induce CRC cell EMT."

sparser
"Additionally, assessment of CRC clinicopathological characteristics demonstrated that USP22 and AP4 expression levels were significantly associated with pT classification, pM classification, AJCC stage and tumor markers used as prognostic indicators of postoperative recurrence and metastasis in CRC, including CEA and CA199."
MED1 affects USP22
| 4
USP22 binds MED1. 4 / 4
| 4

sparser
"The interaction between the endogenous USP22 and MED1 was further validated in mouse primary iNKT cells because MED1 was detected in anti-USP22 immunoprecipitates but not the normal rabbit IgG controls ( xref ), indicating a possibility that USP22 regulates iNKT cell development through, at least partially, MED1 interaction."

sparser
"Since USP22 interacts with MED1, both of which are transcription coactivators involved in promoting T-bet and IL-2R gene transcription in iNKT cells, we then asked whether MED1 binds to the promoters of T-bet and IL-2Rβ in a USP22-dependent manner, or vice versa."

sparser
"However, assessment of ER stress‐induced changes in MED1 and USP22 binding shows poor correlation (Fig  xref )."

sparser
"USP22 interacts with MED1 in iNKT cells."
KDM3A affects USP22
| 2 2
| 2 2

sparser
"Accordingly, we speculated whether USP22 could influence KDM3A expression and detected the interaction between USP22 and KDM3A using a ChIP assay, and found that USP22 could indeed interact with KDM3A ( xref )."

reach
"Accordingly, we speculated whether USP22 could influence KDM3A expression and detected the interaction between USP22 and KDM3A using a ChIP assay, and found that USP22 could indeed interact with KDM3A (Figure 5a)."

reach
"The interaction between USP22 and KDM3A and between KDM3A and YAP1 was further validated."

sparser
"The interaction between USP22 and KDM3A and between KDM3A and YAP1 was further validated."
HIF1A affects USP22
2 | 2
2 | 2

sparser
"In TP53-mutant HCC cells, USP22 and HIF1α formed a positive feedback loop and promote the stemness of HCC."

sparser
"Our laboratory found that HIF-1α can bind to the USP22 promoter."
HIF1 affects USP22
| 4
HIF1 activates USP22. 4 / 4
| 4

reach
"As a key regulator of hypoxia, HIF-1α contributes to HCC development by regulating USP22 and SENP1 to control HCC tumor growth and upregulating angiogenesis related genes."

reach
"In HCC cells with wild-type TP53 , p53 blocks the HIF-1α-induced upregulation of USP22."

reach
"As the direct target genes of HIF-1α, USP22 and TP53 can be upregulated by HIF-1α transcription in hypoxia."

reach
"Specifically, the hypoxia-mediated activation of HIF1α induced the upregulation of USP22, which in turn acted to deubiquitinate HIF1α, leading to the expression of HIF1α target genes, including glycolysis-related genes HK2, PDK1 and enolase 1 (ENO1) [123]."
Gal-SLPs affects USP22
| 4
Gal-SLPs inhibits USP22. 4 / 4
| 4

eidos
"Besides , Gal-SLPs downregulated in vivo USP22 expression to a much greater extent than Gal-LPs ( Figure 7g ) , which was well correlated with the in vitro Western blot analysis , convincingly proving the self-activated cascade-responsive process and synergy of sorafenib and shUSP22 ."

eidos
"Thus , we speculated that the downregulation of USP22 by Gal-SLPs could block the glycolysis and further suppress stemness features in HCC cells ."

eidos
"Thus , Gal-SLPs dramatically suppressed the expression of USP22 ."

eidos
"] Our study provided strong evidence that the downregulation of USP22 by Gal-SLPs suppressed the expression of MRP1 and caused high intracellular sorafenib accumulation ."
CTNNB1 affects USP22
| 2 2
| 2 2

sparser
"Next, we examined the interaction between USP22 and β-catenin within the nucleus after tGCI with or without hypoxia by co-immunoprecipitation assay."

sparser
"The results showed that the interaction between USP22 and β-catenin in the nucleus was significantly weakened at 50 h after tGCI."

reach
"Next, we examined the interaction between USP22 and β-catenin within the nucleus after tGCI with or without hypoxia by co-immunoprecipitation assay."

reach
"The results showed that the interaction between USP22 and β-catenin in the nucleus was significantly weakened at 50 h after tGCI."
C1QTNF9 affects USP22
| 4
C1QTNF9 increases the amount of USP22. 4 / 4
| 4

reach
"We found that CTRP9 increased the expression of USP22, which triggered the de-ubiquitination of Sirt1 and promoted cell autophagy.In conclusion, the present study demonstrated that CTRP9 alleviated li[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"As illustrated, both mRNA and protein expression of USP22 were upregulated by CTRP9 treatment in a dose-dependent manner ( Fig. 3 A&3B)."

reach
"However, the relationship between CTRP9 and Sirt1 has not been fully explored in AS progression.In the present study, we demonstrated that CTRP9 upregulated the expression of USP22, a kind of deubiqui[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Collectively, these results revealed that CTRP9 promoted USP22 expression, which removed the conjugated poly-ubiquitin chains from Sirt1 and enhanced the stabilization of Sirt1 protein.As shown in Fig[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
BIRC5 affects USP22
1 | 2 1
1 | 2 1

reach
"In order to understand the role of USP22, the present study examined the association between USP22, Aurora-B, Survivin and Ki-67 in OSCC."

sparser
"Finally, it was confirmed that USP22 interacted with survivin and stabilized it by downregulating its ubiquitination."

reach
"The functional associations between USP22, Aurora-B and Survivin in OSCC were also investigated."
AR-V7 affects USP22
| 4
AR-V7 binds USP22. 4 / 4
| 4

reach
"Unlike rutaecarpine, nobiletin failed to alter the GRP78-mediated degradation of AR-V7.In conclusion, this research not only demonstrates the reason why nobiletin suppressed the growth process of CRPC through the selective degradation of AR-V7, but also enriches our understanding of the degradation mechanism of AR-V7 and provides an efficient treatment target to overcome CRPC via targeting the interaction between AR-V7 and USP14/USP22 (Fig. 7G)."

reach
"Immunofluorescence and western blot experiments showed that these molecular events mainly occurred in the nucleus.We also confirmed that nobiletin can significantly decrease the interactions between AR-V7 and USP14/USP22 in a time and concentration-dependent manner, but did not affect the interactions between AR-FL and USP14/USP22, which may explain why nobiletin exhibits a certain selectivity in inducing the degradation of AR-V7."

reach
"Binding of USP22 to AR-V7 prevents AR-V7 protein degradation."

reach
"We confirmed the endogenous interaction between AR-V7 and USP22."
AP4 affects USP22
| 4
USP22 binds AP4. 4 / 4
| 4

sparser
"In addition, USP22 binds to the promoter region of AP4 to activate its transcription."

sparser
"Consistent with the above results, ChIP analysis revealed that USP22 binds the promoter region of AP4 (Figure xref )."

sparser
"We also found that USP22 binds to the promoter region of AP4 to mediate its transcription and thus induce CRC cell EMT."

sparser
"Additionally, assessment of CRC clinicopathological characteristics demonstrated that USP22 and AP4 expression levels were significantly associated with pT classification, pM classification, AJCC stage and tumor markers used as prognostic indicators of postoperative recurrence and metastasis in CRC, including CEA and CA199."
Topotecan affects USP22
| 2 1
| 2 1

reach
"Bortezomib, a proteasome inhibitor, but not chloroquine, an autophagy inhibitor, prevented topotecan‐induced USP22 protein degradation (Figure 6H)."

reach
"Pharmacological inhibition of USP22 by topotecan exerts a similar effect to USP22 knockout in inhibiting melanoma metastasis both in vivo and in vitro."

sparser
"Pharmacological inhibition of USP22 by topotecan exerts a similar effect to USP22 knockout in inhibiting melanoma metastasis both in vivo and in vitro."
MiR-144-3p affects USP22
| 3
MiR-144-3p binds USP22. 3 / 3
| 3

reach
"The interaction between miR-144-3p and USP22 was validated by dual-luciferase reporter assay."

reach
"In order to explore the functional relevance of USP22/miR-144-3p interaction on ferroptosis of pancreatic β cells, we transfected USP22 OE alone or together with miR-144-3p mimic into INS-1 cells."

reach
"Hence, miR-144-3p/USP22 interaction might be a crucial event that regulated ferroptosis during T2DM development, which served as a novel target for T2DM treatment."
MiR-140 affects USP22
| 3
MiR-140 inhibits USP22. 3 / 3
| 3

reach
"In addition, overexpression of miR-140 inhibits the proliferation, migration, and invasion of OS cells by directly targeting USP22, suggesting that the miR-140/USP22 axis may represent a novel therape[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Overexpression of miR-140 inhibited cell proliferation, migration, and invasion and promoted cell apoptosis by directly targeting USP22, resulting in its decreased expression."

reach
"Liu et al. also found that miR-140 attenuated osteosarcoma progression via interference of USP22-involved lysine-specific demethylase 1 (LSD1) stabilization and elevating p21 expression (40)."
Macrocycle affects USP22
| 3
| 3

reach
"Michael Morgan et al. firstly reported macrocycle inhibitors of USP22/SAGA DUB module, demonstrating the ability of the cyclic peptides to enter human cells and inhibit H2B deubiquitination [17] ."

reach
"Recent research has identified potent macrocycle inhibitors of USP22, including compound S02, which binds tightly to the catalytic domain pocket of USP22."

reach
"Michael Morgan et al. reported the first macrocycle inhibitors of USP22/SAGA DUB module, demonstrating the ability of the cyclic peptides to enter human cells and inhibit H2B deubiquitination [17] ."
XPC affects USP22
1 | 2
1 | 2

sparser
"An interaction between USP22 and XPC also occurred in the GEMM-derived MAFs, GFP and hUSP22 cells ( xref )."

sparser
"This interaction occurred in control cells as well, indicating that XPC and USP22 interact regardless of USP22 overexpression ( xref )."

reach
"Immune Evasion and Drug Resistance Mediated by USP22 in Cancer: Novel Targets and Mechanisms."

reach
"Overall, lactate contributes to the immune suppressive microenvironment in the context of glioblastoma.In the review article, Guo et al. focus on the immune evasion and drug resistance mediated by USP22 in cancer."

reach
"Co-expression of USP22 and BMI1 can accelerate tumor proliferation, stemness, and drug resistance (35)."

reach
"Collectively, these results suggest that USP22 activates AKT signaling via E2F6-mediated repression of DUSP1.DUSP1 has been identified as a tumor suppressor that negatively regulates tumor proliferati[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Moreover, USP22 promotes cell cycle progression by regulating BMI-mediated INK4a/ARF and Akt signaling pathways [12, 32–34]."

reach
"To further dissect the precise mechanism by which USP22 activates the PI3K/Akt signaling pathway, coimmunoprecipitation (Co‐IP) and mass spectrometry (MS) were utilized to identify the potential direct binding partners of USP22."
USP22 affects miR-144-3p
| 3
MiR-144-3p binds USP22. 3 / 3
| 3

reach
"The interaction between miR-144-3p and USP22 was validated by dual-luciferase reporter assay."

reach
"In order to explore the functional relevance of USP22/miR-144-3p interaction on ferroptosis of pancreatic β cells, we transfected USP22 OE alone or together with miR-144-3p mimic into INS-1 cells."

reach
"Hence, miR-144-3p/USP22 interaction might be a crucial event that regulated ferroptosis during T2DM development, which served as a novel target for T2DM treatment."

reach
"USP22 promotes lipogenesis in Tregs through stabilization of peroxisome proliferator-activated receptor gamma (PPARγ) [51] ."

reach
"In support of this, gain-of-function or loss-of-function studies revealed that USP22 stimulates lipogenesis from glucose in cultured hepatocytes and subsequently contributes to tumorigenesis [193]."

reach
"Genetic and pharmacological inhibition of PPARγ abolished the regulatory effect on ACC and ACLY transcription, and significantly decreased lipogenesis and tumorigenesis caused by USP22 expression both in HCC cells and xenograft tissues."

reach
"In contrast, other studies have shown that USP22 promotes fatty acid synthesis in hepatocellular carcinoma cells and, thus, hepatocellular carcinoma progression ."

reach
"In addition, we find that USP22 promotes de novo fatty acid synthesis and contributes to HCC tumorigenesis, however, this tumorigenicity is suppressed by inhibiting the expression of PPARγ, ACLY, or ACC in in vivo tumorigenesis experiments."

reach
"In p53 wild-type colorectal cancer (CRC) cells, hydrogen peroxide (H 2 O 2 )-induced p53 expression represses the transcription of deubiquitinase USP22, which otherwise deubiquitinates and stabilizes Fatty Acid Synthase (FASN), and thus inhibits fatty acid synthesis."
USP22 increases the amount of extracellular matrix. 3 / 3
| 3

reach
"Under NG conditions, the half-life of Snail1 was about 0.8 h, whereas HG stimulation significantly increased the of Snail1 level and extended its half-life to 1.4 h ( Fig. 7 A), indicating that HG inc[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In vitro experiments confirmed that USP22 promoted the EMT process to increase the expression of ECM components."

reach
"In addition to promoting EMT, USP22 overexpression upregulated the expression of TGF-β1 and ECM components including FN, Collagen I, and Collagen Ⅳ under HG conditions ( Fig. 3 F)."
USP22 affects cellular survival
| 3
USP22 activates cellular survival. 3 / 3
| 3

eidos
"Moreover , USP22 depletion decreased cellular survival in response to cisplatin at 2.5 muM ( 1.9-fold ; p =0 .004 ) and 5 muM ( 3.5-fold ; p =0 .0002 ) in CRPC cells ( Figure 2E , right ) as well as HT-sensitive cells ( p =0 .001 ; Supplemental Figure 2E , right ) , indicating that USP22 is required for cellular survival after DNA damage ."

eidos
"Accordingly , USP22 promotes cellular survival after double strand break-inducing irradiation ( Figure 2 ; Supplemental Figure 2 ; Figure 3 ) ."

eidos
"The USP22-sensitive ubiquitylome reveals altered modification of DNA repair-related proteins The USP22-sensitive transcriptome implicated USP22 in affecting DDR-related pathways ( Figure 2 ) , and in vitro and in vivo studies demonstrated that USP22 modulates proliferative phenotypes as well as cellular survival ( Figures 2-3 ) ."
USP22 affects cell stemness
| 3
USP22 activates cell stemness. 3 / 3
| 3

eidos
"Down-regulation of USP22 reduces cell stemness and enhances the sensitivity of pancreatic cancer cells to cisplatin by inactivating the Wnt / beta-catenin pathway ."

eidos
"Inhibition of USP22 reduced the cell stemness and augmented the sensitivity of PC cells to cisplatin by inhibiting the Wnt / beta-catenin pathway ."

eidos
"Jiang et al.18 pointed out that USP22 contributes to CRC cell stemness and decreases the sensitivity of CRC cells to chemoresistance via the Wnt / beta-Catenin pathway ."
USP22 affects c-NHEJ
| 3
USP22 activates c-NHEJ. 3 / 3
| 3

reach
"We demonstrate that Usp22 promotes c-NHEJ and CSR in vivo."

reach
"These results further suggest that Usp22 promotes the c-NHEJ but not the A-EJ pathway."

reach
"Early pro B ablation of Usp22 results in the blockade of B-cell development and defects in V to DJ recombination, supporting other findings that Usp22 promotes c-NHEJ."
USP22 affects Wnt
| 3
USP22 activates Wnt. 3 / 3
| 3

reach
"In pancreatic cancer cell lines a different mechanism in USP22 was found to modulate the lacatenin/Wnt signaling and therefore increase the abundance of FoxM1, a transcription factor that normally represses the expression of p21 and p27."

reach
"By up- and downregulation of USP22 expression, we also proved that USP22 can activate the Wnt and beta-Catenin pathway, which in turn affected the proliferation and migration of HepG2 cells."

reach
"In addition, USP22 can also activate Wnt/beta-catenin signaling axis to promote stemness and chemoresistance [ 67 ] (see Fig. 2 )."
USP22 affects USP14
| 3
| 3

reach
"Immunofluorescence and western blot experiments showed that these molecular events mainly occurred in the nucleus.We also confirmed that nobiletin can significantly decrease the interactions between AR-V7 and USP14/USP22 in a time and concentration-dependent manner, but did not affect the interactions between AR-FL and USP14/USP22, which may explain why nobiletin exhibits a certain selectivity in inducing the degradation of AR-V7."

reach
"Unlike rutaecarpine, nobiletin failed to alter the GRP78-mediated degradation of AR-V7.In conclusion, this research not only demonstrates the reason why nobiletin suppressed the growth process of CRPC through the selective degradation of AR-V7, but also enriches our understanding of the degradation mechanism of AR-V7 and provides an efficient treatment target to overcome CRPC via targeting the interaction between AR-V7 and USP14/USP22 (Fig. 7G)."

reach
"However, it is worth noting that nobiletin had no effect on the interactions between AR-FL and USP14/USP22, which may explain why nobiletin has a certain selectivity in inducing the degradation of AR-V7.Our previous study has been demonstrated that rutaecarpinecan selectively trigger the GRP78-dependent AR-V7 degradation [23]."
USP22 affects TNFRSF10B
| 3

sparser
"Results: CCK-8 assay showed that the IC 50 values of SW480-USP22 (SW480 cells overexpressing USP22) treated with oxaliplatin for 24 h and 48 h was (4.62±0.05)μmol/L and (2.32±0.04)μmol/L respectively; which was 2.7 times and 3.0 times higher than that in control cells, respectively."

sparser
"The fluorescence intensity of calcein-AM and rhodamine123 in the SW480-USP22 group were significantly increased when compared with that in the control cells (both P <0.01)."

sparser
"The protein expression levels of MRP1 and P-gp in SW480-USP22 cells were significantly increased when compared with that in the control cells(both P <0.01)."
USP22 affects TIMM8A
| 3
USP22 activates TIMM8A. 3 / 3
| 3

reach
"USP22 overexpression significantly enhanced resistance of TNBC cells to DDP ( Figure 3 C and Supplemental Figure S2C)."

reach
"As expected, silencing of USP22 has been proved to reverse the EMT phenotype of DDP-resistant TNBC cells in the present study."

reach
"28 We therefore investigated the regulatory role of USP22 in the chemoresistance, stemness, and EMT phenotype of TNBC cells in vitro and in vivo in the following experiments.The results revealed that [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects TERT
| 3
USP22 inhibits TERT. 3 / 3
| 3

reach
"In retinoblastoma, the depletion of USP22 has been shown to induce cancer cell apoptosis by suppressing the TERT/P53 signal pathway ."

reach
"In retinoblastoma, USP22 depletion induces cancer cell apoptosis by inhibiting the TERT/P53 signaling pathway (89)."

reach
"USP22 can also enhance the telomerase reverse transcriptase (TERT)/p53 signaling pathway [ 65 ] and promote cell proliferation and DNA repair (see Fig. 1 )."
USP22 affects OS tumor growth
| 3
USP22 activates OS tumor growth. 3 / 3
| 3

eidos
"These results suggest that USP22 downregulation inhibited OS tumor growth and metastasis in vivo ."

eidos
"As expected , downregulation of USP22 suppressed OS tumor growth and metastasis in vivo ."

eidos
"In addition , downregulation of USP22 suppressed OS tumor growth and metastasis in vivo ."
USP22 affects MTOR
| 3
USP22 inhibits MTOR. 3 / 3
| 3

reach
"USP22 reduces the mTOR activity to retard the progression of CRC[8]."

reach
"USP22 prevents ubiquitination-mediated EGFR degradation, thereby inducing persistent activation of EGFR-mediated oncogenic signaling pathways, such as STAT3, AKT/mTOR, and MEK/ERK pathways."

reach
"USP22 knockdown protects against cerebral ischemia/reperfusion injury via destabilizing PTEN protein and activating the mTOR/TFEB pathway."
USP22 affects LUAD
| 3
USP22 activates LUAD. 3 / 3
| 3

reach
"In this study, we speculated that USP22 might be involved in a variety of ways to promote the development of LUAD, including ubiquitination and immunosuppression."

reach
"According to the USP22 comprehensive regulation network, we propose that USP22 may promote the development of LUAD through ubiquitination and immunosuppression."

reach
"In conclusion, we constructed a global regulation network to show that USP22 may promote the development of LUAD through ubiquitination and immunosuppression."
USP22 affects KPNA2
| 3
| 3

sparser
"Results from endogenous immunoprecipitation assays suggest that KPNA2 interacted with USP22 constitutively but interacted with IRF3 or pIRF3 in THP-1 cells in a manner dependent on viral infection ( xref )."

sparser
"In our study, we found that knockout of USP22 impaired virus-triggered nuclear translocation of p65 and induction of p65-target genes such as Tnf and Il6 , which was restored by reconstitution of KPNA2, suggesting a role of the USP22KPNA2 axis in NF-κB activation after viral infection (data not shown)."

sparser
"In contrast, mutation of USP22 NLS (USP22-RR/AA) did not affect its association with KPNA2 ( xref ), suggesting that USP22 interacts with KPNA2 independently of its NLS."
USP22 affects IFN-λ1
| 3
USP22 decreases the amount of IFN-λ1. 3 / 3
| 3

reach
"These findings suggest that USP22 negatively regulates IFN-λ1 expression and ISG induction."

reach
"Mean and SD of three independent experiments in triplicate USP22 negatively regulates STAT1 signaling and IFN-λ1 expression."
| DOI

reach
"Nat Rev Immunol 21, 852 548-569, doi:10.1038/s41577-021-00524-z (2021These findings suggest that USP22 negatively regulates IFN-λ1 expression and ISG 232 induction."
| DOI
USP22 affects Hypertrophy
| 3
| 3

reach
"In the current study, we show that the overexpression of USP22 induced by treatment with FO greatly improved Ang II-induced cardiac dysfunction, heart hypertrophy, inflammation, and fibrosis (Figures 1–4)."

reach
"Recent research has shown that USP22 protects against myocardial ischemia‒reperfusion injury (Ma et al., 2020), but it is still unknown how USP22 affects Ang II-induced cardiac remodeling and hypertrophy."

reach
"PAS staining results showed that USP22 deficiency attenuated the renal tubular hypertrophy and mesangial expansion of diabetic mice ( Fig. 9 C)."
USP22 affects HES1
1 | 2
USP22 deubiquitinates HES1. 2 / 3
1 | 2

reach
"Another study demonstrated that USP27x, Usp22, and Usp51 deubiquitinate and stabilize Hes1 protein, a transcriptional repressor necessary for the maintenance of neural stem/progenitor cells."

reach
"Hes1, which undergoes a fast turnover rate due to its degradation by the proteasome, is deubiquitinated and stabilized by USP22 [102]."
USP22 affects H2BK120ub
| 3
USP22 deubiquitinates H2BK120ub. 3 / 3
| 3

reach
"USP22 is involved in the deubiquitylation of H2BK120ub, a mark that is found at DSB sites, USP22 could be a “reader” for this mark and potentiate the recruitment of other DNA damage factors through this ubiquitin mark."

reach
"14 , 38 To detect whether USP22 participates in the deubiquitination of H2AK119ub and H2BK120ub during Sorafenib‐induced ferroptosis, we initially conducted Western blotting to assess the levels of H2AK119ub and H2BK120ub in USP22 knockout or control cells treated with DMSO or Sorafenib."

reach
"Other major DUBs with specificity for histone H2AK119ub over H2BK120ub are BAP1 and USP16, while USP3, USP12, USP22, and USP44 deubiquitinate both H2AK119ub and H2BK120ub, as well as different non histone substrates [XREF_BIBR]."
USP22 affects ESR1
| 3
| 3

sparser
"METTL14-mediated m6A modification enhances USP22-ERα axis to drive breast cancer malignancy."

sparser
"However, the post-transcriptional modifications on the USP22-ERα axis remain elusive."

sparser
"Moreover, USP22 associates with ERα to be involved in maintenance of ERα stability."
USP22 affects DNA Damage
| 3
| 3

reach
"According to that model, USP22 enhances DNA damage repair and cisplatin resistance by deubiquitinating histone H2A, which in turn facilitates the phosphorylation of histone H2AX."

reach
"The study reveal the dual mechanism of USP22 involvement in cisplatin resistance : (1) USP22 enhances DNA damage repair and induce cisplatin resistance by promoting the phosphorylation of histone H2AX via deubiquitinating histone H2A."

reach
"Knockdown of USP22 in an in vivo model was shown to decrease tumor angiogenesis, impair non-homologous DNA damage repair pathways and significantly improve the therapeutic efficacy of cisplatin."
USP22 affects CRPC
| 3
USP22 activates CRPC. 3 / 3
| 3

reach
"USP22 promotes CRPC through enhanced ligand independent and androgen stimulated AR transcriptional activity and sustained AR activity in the presence of antagonists, concomitant with AR protein accumulation."

reach
"In prostate cancer, USP22 predicts disease outcome and promotes the CRPC phenotype by controlling AR and MYC dual regulation (63)."

reach
"Given the ability of USP22 to enhance AR accumulation, AR activity, and CRPC, the biological impact of a model of tetracycline inducible shUSP22 was developed in therapy sensitive PCa cells."
USP22 affects CD101
| 3
| 3

sparser
"This Study demonstrated that targeting PCBP1-AS1 increases the sensitivity of prostate cancer cells to Enzalutamide by reducing the binding of USP22 to AR-V7 or AR and decreasing the stability of the complex, providing new ideas for the clinical treatment of drug-resistant patients with CRPC."

sparser
"Collectively, nobiletin selectively induces AR-V7 degradation by inhibiting the interaction of AR-V7 with USP22."

sparser
"Binding of USP22 to AR-V7 prevents AR-V7 protein degradation."
USP22 affects Aurora-B
| 3
USP22 decreases the amount of Aurora-B. 3 / 3
| 3

reach
"Indeed, the present study identified that USP22 siRNA decreased Aurora-B and Survivin expression in OSCC cells."

reach
"Furthermore, USP22 small interfering RNA inhibited cell growth and reduced the expression levels of Aurora-B, Survivin and Cyclin B, together with the upregulation of cyclin-dependent kinase inhibitor 1A (p21)."

reach
"Notably, USP22 siRNA reduced the protein levels of Aurora-B and Survivin in OSCC cells (Fig. 2B)."

reach
"USP22 promotes cell proliferation, migration, and invasion in multiple cancers, including glioma, lung adenocarcinoma, thyroid carcinoma, colorectal cancer, and gastric cancer [27–31]."

reach
"USP22 plays as an interactor with LINC01426 and activates the hedgehog pathway to promote the invasion and metastasis in lung adenocarcinoma via stabilizing SHH protein [ 48 ]."

reach
"The Deubiquitinase USP22-Stabilized COL17A1 Promotes Lung Adenocarcinoma Progression."
USP22 affects AR-FL
| 3
AR-FL binds USP22. 3 / 3
| 3

reach
"However, it is worth noting that nobiletin had no effect on the interactions between AR-FL and USP14/USP22, which may explain why nobiletin has a certain selectivity in inducing the degradation of AR-V7.Our previous study has been demonstrated that rutaecarpinecan selectively trigger the GRP78-dependent AR-V7 degradation [23]."

reach
"Immunofluorescence and western blot experiments showed that these molecular events mainly occurred in the nucleus.We also confirmed that nobiletin can significantly decrease the interactions between AR-V7 and USP14/USP22 in a time and concentration-dependent manner, but did not affect the interactions between AR-FL and USP14/USP22, which may explain why nobiletin exhibits a certain selectivity in inducing the degradation of AR-V7."

reach
"In addition, the interaction of USP22 and AR (including AR-FL and AR-Vs) was also confirmed."
USP22 affects AKT1
| 2
USP22 activates AKT1. 2 / 3
| 2

reach
"We found that USP22 promotes multidrug resistance in HCC cells by activating SIRT1/protein kinase B (Akt)/multidrug resistance-associated protein 1 (MRP1) pathway, while inhibition of USP22 and SIRT1 [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Activation of ERK1/2 kinase rather than AKT1 by USP22 was found to be one of the mechanisms promoting LC3 processing."
USP22 affects ABCD1
| 3
USP22 activates ABCD1. 3 / 3
| 3

reach
"Second, USP22 silencing reduced ALD in vivo."

reach
"These results indicate that USP22 contributes to ALD in vitro."

reach
"And this will provide more sufficient theoretical basis for USP22 as a therapeutic target for ALD.In summary, our findings provide the first confirmation of the essential role of USP22 in Lieber-DeCar[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 3

reach
"In conclusion, USP22 is upregulated in a variety of malignancies, and studies have shown that USP22 induces tumor resistance to 5-FU by acting on CSCs, SIRT1, and C-MYC."

reach
"Studies have demonstrated that USP22 directly interacts with SIRT1, activating the AKT/GSK-3β/multidrug resistance-associated protein 1 (MRP1) pathway, thereby enhancing 5-Fu efflux and reducing 5-Fu-induced apoptosis in HCC cells [124]."

reach
"USP22 up-regulates the SIRT1/AKT/MRP1 signaling pathway and promotes the efflux of 5-Fluorouracil (5-FU), leading to the treatment resistance of HCC [17]."
USP14 affects USP22
| 3
| 3

reach
"Immunofluorescence and western blot experiments showed that these molecular events mainly occurred in the nucleus.We also confirmed that nobiletin can significantly decrease the interactions between AR-V7 and USP14/USP22 in a time and concentration-dependent manner, but did not affect the interactions between AR-FL and USP14/USP22, which may explain why nobiletin exhibits a certain selectivity in inducing the degradation of AR-V7."

reach
"Unlike rutaecarpine, nobiletin failed to alter the GRP78-mediated degradation of AR-V7.In conclusion, this research not only demonstrates the reason why nobiletin suppressed the growth process of CRPC through the selective degradation of AR-V7, but also enriches our understanding of the degradation mechanism of AR-V7 and provides an efficient treatment target to overcome CRPC via targeting the interaction between AR-V7 and USP14/USP22 (Fig. 7G)."

reach
"However, it is worth noting that nobiletin had no effect on the interactions between AR-FL and USP14/USP22, which may explain why nobiletin has a certain selectivity in inducing the degradation of AR-V7.Our previous study has been demonstrated that rutaecarpinecan selectively trigger the GRP78-dependent AR-V7 degradation [23]."
TNFRSF10B affects USP22
| 3

sparser
"Results: CCK-8 assay showed that the IC 50 values of SW480-USP22 (SW480 cells overexpressing USP22) treated with oxaliplatin for 24 h and 48 h was (4.62±0.05)μmol/L and (2.32±0.04)μmol/L respectively; which was 2.7 times and 3.0 times higher than that in control cells, respectively."

sparser
"The fluorescence intensity of calcein-AM and rhodamine123 in the SW480-USP22 group were significantly increased when compared with that in the control cells (both P <0.01)."

sparser
"The protein expression levels of MRP1 and P-gp in SW480-USP22 cells were significantly increased when compared with that in the control cells(both P <0.01)."
SOX2 affects USP22
| 3
SOX2 increases the amount of USP22. 3 / 3
| 3

reach
"USP22 is located directly on the Sox2 promoter and negatively regulates Sox2 transcription in ESCs [XREF_BIBR]."

reach
"USP22 has been found to be located directly on the Sox2 promoter and catalyzes deubiquitination of H2B and attenuates Sox2 transcription."

reach
"USP22 is known to bind to the promoter region of Sox2 and negatively regulates Sox2 transcription in embryonic stem cells (ESCs) 47."
RAS affects USP22
| 3
RAS activates USP22. 3 / 3
| 3

reach
"These findings suggest that SOS1/RAS and downstream ERK and PI3K/AKT pathways mediate the oncogenic role of USP22 in gastric cancer."

reach
"These results suggest that SOS1/RAS signaling mediates the oncogenic role of USP22 in gastric cancer."

reach
"Furthermore, our study indicated that SOS1 was upregulated in USP22-overexpressing gastric cancer cells and xenograft tumor tissue, accompanied by activation of the RAS/ERK and PI3K/AKT pathways, suggesting that SOS1/RAS signaling mediates the oncogenic role of USP22 in gastric cancer."
KPNA2 affects USP22
| 3
| 3

sparser
"Results from endogenous immunoprecipitation assays suggest that KPNA2 interacted with USP22 constitutively but interacted with IRF3 or pIRF3 in THP-1 cells in a manner dependent on viral infection ( xref )."

sparser
"In our study, we found that knockout of USP22 impaired virus-triggered nuclear translocation of p65 and induction of p65-target genes such as Tnf and Il6 , which was restored by reconstitution of KPNA2, suggesting a role of the USP22KPNA2 axis in NF-κB activation after viral infection (data not shown)."

sparser
"In contrast, mutation of USP22 NLS (USP22-RR/AA) did not affect its association with KPNA2 ( xref ), suggesting that USP22 interacts with KPNA2 independently of its NLS."
KLF3-AS1 affects USP22
| 3
KLF3-AS1 activates USP22. 3 / 3
| 3

reach
"Here, we validated that KLF3-AS1 sponged miR-206 to downregulate its level, thereby upregulating USP22."

reach
"KLF3-AS1 induced USP22 to promote the deubiquitination of Sirt1 protein."

reach
"Furthermore, overexpression of KLF3-AS1 upregulated Sirt1 and USP22, but silencing of USP22 reversed these effects (Fig. 4C)."
HDAC1 affects USP22
| 1 1 1
HDAC1 inhibits USP22. 3 / 3
| 1 1 1

sparser
"As shown in xref , linear hD1 inhibits USP22 at a ~4-fold higher concentration, indicating the constraining the peptide does not play a major role in the effectiveness of this inhibitor."

reach
"As shown in figure 3B, linear hD1 inhibits USP22 at a ~4-fold higher concentration, indicating the constraining the peptide does not play a major role in the effectiveness of this inhibitor."

eidos
"The most potent inhibitor , hD1 , inhibits USP22 in vitro with a Ki of 180 nM ( Figure 3A ) ."
ESR1 affects USP22
| 3
| 3

sparser
"METTL14-mediated m6A modification enhances USP22-ERα axis to drive breast cancer malignancy."

sparser
"However, the post-transcriptional modifications on the USP22-ERα axis remain elusive."

sparser
"Moreover, USP22 associates with ERα to be involved in maintenance of ERα stability."
CD101 affects USP22
| 3
| 3

sparser
"This Study demonstrated that targeting PCBP1-AS1 increases the sensitivity of prostate cancer cells to Enzalutamide by reducing the binding of USP22 to AR-V7 or AR and decreasing the stability of the complex, providing new ideas for the clinical treatment of drug-resistant patients with CRPC."

sparser
"Collectively, nobiletin selectively induces AR-V7 degradation by inhibiting the interaction of AR-V7 with USP22."

sparser
"Binding of USP22 to AR-V7 prevents AR-V7 protein degradation."
AR-FL affects USP22
| 3
AR-FL binds USP22. 3 / 3
| 3

reach
"However, it is worth noting that nobiletin had no effect on the interactions between AR-FL and USP14/USP22, which may explain why nobiletin has a certain selectivity in inducing the degradation of AR-V7.Our previous study has been demonstrated that rutaecarpinecan selectively trigger the GRP78-dependent AR-V7 degradation [23]."

reach
"Immunofluorescence and western blot experiments showed that these molecular events mainly occurred in the nucleus.We also confirmed that nobiletin can significantly decrease the interactions between AR-V7 and USP14/USP22 in a time and concentration-dependent manner, but did not affect the interactions between AR-FL and USP14/USP22, which may explain why nobiletin exhibits a certain selectivity in inducing the degradation of AR-V7."

reach
"In addition, the interaction of USP22 and AR (including AR-FL and AR-Vs) was also confirmed."
| 3

reach
"Remarkably, silencing USP22 reduced the tumor volume of BEL/FU cells treated with 5-FU (BEL/FU control shRNA cells treated with 5-FU vs BEL/FU USP22 shRNA treated with 5-FU, 945 +/-545mm 3 vs 372 +/-228mm 3, P < 0.05)."

reach
"After the Bel/Fu cells were injected into nude mice, the mean diameter of the subcutaneous xenografts that developed was 1.5 cm, whereas for mice injected with Bel/Fu cells with stable expression of USP22 siRNA, the mean diameter was smaller (1.4 cm, n = 6, P < 0.05, XREF_FIG); after the mice injected with Bel/Fu cells were treated with 5-FU, the mean diameter of xenografts was smaller (1.2 cm, n = 6, P < 0.05, XREF_FIG); after the mice injected with Bel/Fu cells with stable expression of USP22 siRNA were treated with 5-FU, the mean diameter of xenografts was 0.9 cm, significantly smaller than that observed in mice injected with Bel/Fu cells alone (n = 6, P < 0.05, XREF_FIG)."

reach
"After the mice injected with Bel/Fu cells with siRNA interference of USP22 expression were treated with 5-FU, the size of the subcutaneous xenografts significantly decreased, suggesting that downregulation of USP22 expression mitigated 5-FU resistance and increased the sensitivity of HCC to chemotherapy drugs, which may be related to the physiological function of USP22 to form a complex with BMI1."
Peptide affects USP22
| 1 1
| 1 1

eidos
"Cells treated with the cyclic peptide inhibitors had increased H2B-Ub levels , indicating that the peptides can also inhibit USP22 in vivo ."

reach
"Macrocyclic peptide inhibitors of USP22 have been developed by Morgan et al. (53) with one representative named hD1 ( Figure 8D , compound 19)."
P38 affects USP22
| 2
P38 decreases the amount of USP22. 2 / 2
| 2

reach
"It has been also discovered that p38 MAPK reduces USP22 expression in HeLa cells [37]."

reach
"Cisplatin, the activator of p38 MAPK, also suppressed USP22 expression."
Ethanol affects USP22
| 1 1
Ethanol increases the amount of USP22. 2 / 2
| 1 1

sparser
"Together with the fact that ALD often progress toward cirrhosis and ultimately to liver cancer [46] and that USP22 has been known as an oncogene [48] , our study here implies a potential role of alcoh[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Hence, USP22 expression and BRD4 stability was enhanced by chronic alcohol consumption.To determine how alcohol stimulation induces USP22 expression in liver cells, we analyzed the effect of alcohol o[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
Chloroform affects USP22
| 2
Chloroform increases the amount of USP22. 2 / 2
| 2

reach
"TCM from B16 and LLC1, but not EG7 cells, enhanced Usp22 and Usp21 mRNA levels (Fig. 2A)."

reach
"To support this notion, we further demonstrated that the TGF-β inhibitor completely diminished the TCM-induced mRNA enhancement of Usp22 (Fig. 2C), signifying that TGF-β is the primary factor in the B16 and LLC1 TCM that enhances Usp22 expression."
ZRANB1 affects USP22
| 2
ZRANB1 increases the amount of USP22. 2 / 2
| 2

reach
"These results implied that ZRANB1 induces the expression of USP22 via regulating Sox9.To better understand the correlation of ZRANB1, Sox9, and USP22 during the progression of CRC, we co-transfected S[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"These results implied that ZRANB1 induces the expression of USP22 via regulating Sox9.Finally, to ascertain that ZRANB1 regulated CRC progression through USP22, we co-transfected SW480 cells with vect[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects ubH2B
| 2
USP22 deubiquitinates ubH2B. 2 / 2
| 2

reach
"While other events, such as histone exchange or direct or indirect RNF20 inactivation, are also possible, studies in the lab are ongoing to determine if USP22 (the mammalian homolog to Ubp8) also deubiquitinates ubH2B."

reach
"Deubiquitination of ubH2B is catalyzed by the ubiquitin protease subunit of the Spt-Ada-Gcn5 Acetyltransferase (SAGA) transcriptional coactivator complex : Nonstop (FBgn0013717) in Drosophila melanogaster, Ubp8 in Saccharomyces cerevisiae, and USP22 in humans."
USP22 affects tumor growth
| 1
USP22 activates tumor growth. 1 / 2
| 1

eidos
"Moreover , the DUB USP7 , USP13 , USP22 , USP28 , USP36 and USP37 stabilize c-Myc , thereby stimulating tumor growth [ 155 , 157 , 171-174 ] ."
| PMC
USP22 affects proteolysis
| 2
| 2

reach
"Mechanistically, PRDM1 enhances USP22 transcription, thus reducing SPI1 protein degradation through deubiquitination, which enhances PD-L1 transcription."

reach
"Here, by utilizing the DUB siRNA library, we found that USP22 could reduce SPI1 protein degradation through deubiquitination."
USP22 affects hydrolase
| 2
| 2

reach
"Interestingly, the human and fly SAGA complexes possess a module that houses ubiquitin hydrolase activity mediated by the USP22 (human) and Nonstop (fly) proteins."

reach
"As suspected from its domain structure, USP22 is able to hydrolyze a ubiquitin linkage from histone H2B in vitro and endogenous USP22 contributes ubiquitin hydrolase activity to the hSAGA complex."

reach
"Immune Evasion and Drug Resistance Mediated by USP22 in Cancer: Novel Targets and Mechanisms."

reach
"Overall, lactate contributes to the immune suppressive microenvironment in the context of glioblastoma.In the review article, Guo et al. focus on the immune evasion and drug resistance mediated by USP22 in cancer."

reach
"Moreover, USP22 promotes both HR (23) and NHEJ (48, 59)."

eidos
"Moreover , USP22 promotes both HR ( 23 ) and NHEJ ( 48 , 59 ) ."
USP22 affects conjugation
| 2
| 2

reach
"Coexpression of USP22 significantly inhibited ubiquitin-conjugation to Sirt1 ( Figure 2 A)."

reach
"As expected, USP22 overexpression significantly inhibited ubiquitin conjugation to c-Myc ( Figure 5 E)."
| 2

reach
"Our data revealed that knockdown of USP22 decreased the expression of stemness related genes Sox2 and CD133, redox related genes Nrf2 and Sirt1, and quiescence related genes PP2A and c-Myc in GSCs und[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"As presented in Fig. 2 A–B , knockdown of USP22 inhibited the expression of stemness related genes Sox2 and CD133, redox related genes Nrf2 and Sirt1, and quiescence related genes PP2A and c-Myc."

reach
"Our results indicate that USP22 promotes cell proliferation, migration, invasion, and cell cycle transition, while inhibiting apoptosis in gastric cancer cells."

reach
"Gain- and loss-of-function assays showed that USP22 promoted gastric cancer cell growth and cell cycle transition while suppressing apoptosis in vitro."
USP22 affects cell apoptosis
| 1
USP22 activates cell apoptosis. 1 / 2
| 1

eidos
"Accordingly , USP22 downregulation also attenuated cerebral I / R-induced oxidative stress , inflammation , and cell apoptosis in mice , thereby reducing nerve injury and neurological dysfunction [ 20 ] ."
USP22 affects SIRT1/AKT/MRP1
| 2
USP22 activates SIRT1/AKT/MRP1. 2 / 2
| 2

reach
"Together, these results indicate that USP22 could promote the MDR in HCC cells by activating the SIRT1/AKT/MRP1 pathway."

reach
"USP22 up-regulates the SIRT1/AKT/MRP1 signaling pathway and promotes the efflux of 5-Fluorouracil (5-FU), leading to the treatment resistance of HCC [17]."
USP22 affects RNF220
| 1 1
| 1 1

reach
"Mechanistically, RNF220 bound to USP22, and interference of RNF220 downregulated the Wnt/β-catenin axis via USP22, which was confirmed by the overexpression of USP22 in both cell lines."

sparser
"Mechanistically, RNF220 bound to USP22, and interference of RNF220 downregulated the Wnt/β-catenin axis via USP22, which was confirmed by the overexpression of USP22 in both cell lines."
USP22 affects RNF20
| 2
| 2

sparser
"Future studies should also examine the mechanism by which the transcriptional regulations of RNF20 and USP22 interact."

sparser
"To determine the association of USP22, H2Bub1 and RNF20 protein expression with CICs, we measured the expression level of these proteins in both CD133+ and CD133− cells."
USP22 affects RB1
| 2
USP22 leads to the phosphorylation of RB1. 2 / 2
| 2

reach
"In pharyngeal carcinoma, knockdown of USP22 not only upregulated p21 expression but also upregulated p27 expression and suppressed retinoblastoma protein (RB) phosphorylation."

reach
"In vitro assays showed that USP22 depletion suppressed ATC cell survival and proliferation by decreasing Rb phosphorylation and cyclin D2, inactivating Akt, and simultaneously upregulating Rb; USP22 silencing restrained cell migration and invasion by inhibiting epithelial-mesenchymal transition; USP22 knockdown promoted mitochondrion- mediated and caspase dependent apoptosis by upregulating Bax and Bid and promoting caspase-3 activation."
USP22 affects Proteasome
| 1
| 1

reach
"CCND1 is protected from proteasome-mediated degradation by USP22, and aggressive growth phenotypes in CRC cells are inhibited [87]."
USP22 affects PRAM1
| 2
USP22 activates PRAM1. 2 / 2
| 2

reach
"To further analyze the dynamics of USP22-mediated PML-RARα stabilization, the long isoform of WT PML-RARα was expressed in control and USP22 KO HEK293T cells and PML-RARα stability was analyzed upon exposure to CHX."

reach
"Next, we aimed to elucidate the biological relevance of USP22-mediated PML-RARα stabilization."
USP22 affects NP
| 2
NP binds USP22. 2 / 2
| 2

sparser
"We demonstrated that SARS-CoV-2 NP proteins undergo ubiquitination-dependent degradation in host cells, while USP22 interacts with SARS-CoV-2 NP and downregulates K63-linked polyubiquitination of SARS-CoV-2 NP, thereby protecting SARS-CoV-2 NP from degradation."

sparser
"In this study, we provide evidence that SARS-CoV-2 NP interacts with the deubiquitinase USP22 in host cells, which downregulates SARS-CoV-2 NP ubiquitination."
USP22 affects NFE2L2
| 2
USP22 activates NFE2L2. 2 / 2
| 2

reach
"Our data revealed that knockdown of USP22 decreased the expression of stemness related genes Sox2 and CD133, redox related genes Nrf2 and Sirt1, and quiescence related genes PP2A and c-Myc in GSCs und[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"As presented in Fig. 2 A–B , knockdown of USP22 inhibited the expression of stemness related genes Sox2 and CD133, redox related genes Nrf2 and Sirt1, and quiescence related genes PP2A and c-Myc."
USP22 affects NFATc1
| 2
USP22 binds NFATc1. 2 / 2
| 2

sparser
"As NFATc1 has high homology with NFATc2, we also tested the interaction between USP22 and NFATc1."

sparser
"The reciprocal co-immunoprecipitation assay showed that USP22 could not interact with NFATc1 ( Fig. 2 C and D)."
USP22 affects MYH9
| 1 1
| 1 1

sparser
"Furthermore, we performed co-IP assays in HEK-293T cells and found that MYH9 could interact with USP22, especially MYH9-S1943E, but not MYH9-S1943A (Fig. xref ; Supplementary Fig. xref )."

reach
"This study confirmed that the p‐MYH9/USP22 complex stabilizes HIF‐1α under normoxic conditions, providing a promising therapeutic target for HCC cells that have metastasized into the bloodstream following drug resistance."
USP22 affects MKI67
| 1 1
| 1 1

sparser
"Ki67 expression was associated with USP22 overexpression in cervical cancer, prostate cancer and oral squamous cell carcinoma [ xref , xref , xref ]."

reach
"In order to understand the role of USP22, the present study examined the association between USP22, Aurora-B, Survivin and Ki-67 in OSCC."
USP22 affects HSPA5
| 2
USP22 deubiquitinates HSPA5. 2 / 2
| 2

reach
"It should be noted that while upregulation of the best characterized H2Bub1 DUB, USP22, has been associated with poorer breast cancer patient outcomes [ 167 ], to date, mechanistic investigations indi[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"These authors also reported down regulation of the Heat shock protein family A (Hsp70) member 5 (HSPA5), an HSP70 family member involved in the unfolded protein response, in tumors from Usp22 null mice, consistent with reports that USP22 mediates deubiquitination of HSPA5 in human prostate cancer cells [15]."
USP22 affects H2Aub1
| 2
USP22 deubiquitinates H2Aub1. 2 / 2
| 2

reach
"Along these lines it is interesting to note that neither free USP22 nor a stable recombinant subcomplex, composed of TAF5L, ATXN7L3, ENY2, and USP22, can deubiquitinate H2Aub1 or H2Bub1 in vitro, sugg[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Interestingly, neither free recombinant USP22 nor Ubp8p can efficiently deubiquitylate H2Bub1 or H2Aub1 (in the case of USP22) in vitro , indicating that an interaction(s) between the ZnF-UBP and othe[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP22 affects H2AC20
1 | 1
1 | 1

sparser
"Subsequent immunoprecipitation (IP) experiments confirmed that USP22 bind to endogenous H2A in A549 cell line (Figures xref )."
USP22 affects GSE1
| 2
GSE1 binds USP22 and RCOR1. 2 / 2
| 2

reach
"What is the biological function of the GSE1-USP22-CoREST complex?"

reach
"The GSE1-USP22-CoREST complex seems to be stable, as we do not see changes in its composition following DNA damage induction."
USP22 affects CXCL8
| 2
USP22 decreases the amount of CXCL8. 2 / 2
| 2

reach
"In addition, loss of USP22 expression also induced elevated basal secretion of the pro-inflammatory cytokines CXCL10 and IL-8 and minor changes in the secretion of IFN-α2 and GM-CSF, compared to controls (Fig. 3C)."

reach
"In addition, loss of USP22 expression also induced 229 elevated basal secretion of the pro-inflammatory cytokines CXCL10 and IL-8 and minor 230 changes in the secretion of IFN-α2 and GM-CSF, compared to controls."
| DOI
USP22 affects CDKN
| 2
USP22 decreases the amount of CDKN. 2 / 2
| 2

reach
"USP22 increases the expression of cyclin-dependent kinase inhibitor 1A (CDKI) including p21 and p27, and it decreases the expression of Rb protein in HNSCC cells [74,75,76]."

reach
"A previous study demonstrated that USP22 could inhibit the transcription of the cyclin-dependent kinase inhibitor 1A (p21) gene by deubiquitinating the transcriptional regulator fructose-1,6-bisphosphatase (FBP1) (5)."
USP22 affects CDK1
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1 | 1

sparser
"From our proteomic analysis, we noticed that CCNB1 appears to form a complex with both USP22 and CDK1; this prompted us to ask whether CDK1 phosphorylates USP22."

reach
"USP22 promotes the proliferation of NPC."

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"By contrast, USP22 was overexpressed in NPC cells and promoted the proliferation of NPC."
USP22 affects ATP
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USP22 activates ATP. 2 / 2
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"We found that USP22 knockdown could inhibit the glycolytic pathway in osteosarcoma cells, suppress cell proliferation, migration and invasion, and promote cancer cell apoptosis.Glycolysis is one of the major pathways for cells to generate energy by converting glucose into lactate and producing ATP (Lunt & Vander Heiden, 2011)."

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"The results showed that compared to the control group, USP22 expression knockdown significantly decreased glucose consumption, lactate production, and ATP generation in Sao-2 cells (p < 0.05)."
USP22 affects AR-V7 protein
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USP22 activates AR-V7 protein. 2 / 2
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"Our current research focuses on the regulation of AR-V7 protein stability mediated by USP22."

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"Our CHX-tracking assay confirmed that USP22 promoted the stability of AR-V7 protein."
USP22 affects ADR
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USP22 inhibits ADR. 2 / 2
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"In addition, overexpression of USP22 also reduced the intracellular ADR concentration in BEL7402 cells."

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"As we previously described, USP22 silencing increased the concentration of intracellular ADR."
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"USP22 knockdown enhanced chemosensitivity of hepatocellular carcinoma cells to 5-Fu by up-regulation of Smad4 and suppression of Akt."

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"We also found that USP22 knockdown enhanced the anti-growth and pro apoptotic effect of 5-Fu in Bel/Fu cells."
UMOD affects USP22
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UMOD decreases the amount of USP22. 2 / 2
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"In terms of inhibitors, 4′-O-tetrahydropyranyl-adriamycin (THP; pirarubicin) partially reduces USP22 expression and promotes HeLa cell apoptosis [ 92 ]."

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"The present study demonstrated that THP decreased USP22 expression and promoted HeLa cell apoptosis partially by inhibiting the phosphorylation of CREB-1."
TAFAZZIN affects USP22
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TAFAZZIN increases the amount of USP22. 2 / 2
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"Taz enhanced USP22 expression at both the mRNA and protein levels in various colon cancer cell lines, including HCT116, SW480, LS180, and MC38 cells (Figure 3A; Figure S3C,D, Supporting Information)."

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"This suggests that EZH2 regulates USP22 expression through its SET domain‐mediated histone methylation activity.The immunofluorescence results similarly indicated that Taz treatment upregulated the cellular expression of both USP22 and PD‐L1 (Figure 3H)."
SNHG16 affects USP22
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SNHG16 increases the amount of USP22. 2 / 2
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"It has also been found that SNHG16 promotes the progression of CRC by activating the expression of USP22 via sponging miR-132-3p[65,66]."

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"He et al (178) concluded that SNHG16 could activate USP22 expression to promote CRC progression via absorbing miR-132-3p."

reach
"Whereas, in p53-deficient CRC cells, relief of ROS-mediated USP22 repression promotes FASN stabilization and lipid accumulation, and thus promotes tumorigenesis (Fig. 7D)."

reach
"Whereas, in p53-deficient CRC cells, ROS-mediated inhibition of USP22 is relieved, leading to FASN stabilization, which thus promotes lipid synthesis and tumor growth."
RNF220 affects USP22
| 1 1
| 1 1

reach
"Mechanistically, RNF220 bound to USP22, and interference of RNF220 downregulated the Wnt/β-catenin axis via USP22, which was confirmed by the overexpression of USP22 in both cell lines."

sparser
"Mechanistically, RNF220 bound to USP22, and interference of RNF220 downregulated the Wnt/β-catenin axis via USP22, which was confirmed by the overexpression of USP22 in both cell lines."
RNF20 affects USP22
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sparser
"Future studies should also examine the mechanism by which the transcriptional regulations of RNF20 and USP22 interact."

sparser
"To determine the association of USP22, H2Bub1 and RNF20 protein expression with CICs, we measured the expression level of these proteins in both CD133+ and CD133− cells."
RCOR1 affects GSE1
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GSE1 binds USP22 and RCOR1. 2 / 2
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"What is the biological function of the GSE1-USP22-CoREST complex?"

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"The GSE1-USP22-CoREST complex seems to be stable, as we do not see changes in its composition following DNA damage induction."
RALY affects USP22
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RALY activates USP22. 2 / 2
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"Here, we show that RALY interacts directly with ALYREF and promotes USP22 mRNA nuclear export by transferring it to ALYREF.Due to the fact that RALY serves as a key oncogene, the biologically targeted degradation of RALY could become an effective therapeutic strategy."

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"Stable RALY promotes tumor growth by regulating the ALYREF-dependent nuclear export of USP22 mRNA."
PI3K affects USP22
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PI3K inhibits USP22. 2 / 2
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"Inhibition of the PI3K/Akt pathway could rescue the effects of USP22 on cell migration."

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"These results suggest that USP22 knockdown induced chemosensitivity of HCC cells by down-regualting PI3K and Akt, and Smad4 mediated Akt suppression as well."
NP affects USP22
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NP binds USP22. 2 / 2
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sparser
"We demonstrated that SARS-CoV-2 NP proteins undergo ubiquitination-dependent degradation in host cells, while USP22 interacts with SARS-CoV-2 NP and downregulates K63-linked polyubiquitination of SARS-CoV-2 NP, thereby protecting SARS-CoV-2 NP from degradation."

sparser
"In this study, we provide evidence that SARS-CoV-2 NP interacts with the deubiquitinase USP22 in host cells, which downregulates SARS-CoV-2 NP ubiquitination."
NFATc1 affects USP22
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USP22 binds NFATc1. 2 / 2
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sparser
"As NFATc1 has high homology with NFATc2, we also tested the interaction between USP22 and NFATc1."

sparser
"The reciprocal co-immunoprecipitation assay showed that USP22 could not interact with NFATc1 ( Fig. 2 C and D)."
Mice affects USP22
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Mice activates USP22. 2 / 2
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"Consistent with these results, no increase in ERBB2, phospho-ERBB2, or ERα protein levels were observed in endpoint tumor lysates isolated from USP22 overexpressing mice, relative to tumor lysates isolated from MMTV-NIC mice (Fig 2A top)."

reach
"We observed over branching of the mammary glands in USP22 overexpressing mice, along with aberrant up regulation of many of the same signaling pathways that were down regulated in Usp22 null embryos, including estrogen receptor, extracellular signal-regulated kinase /Mitogen-activated protein kinase (ERK/MAPK), and TGFβ signaling [16, 17]."
MYH9 affects USP22
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sparser
"Furthermore, we performed co-IP assays in HEK-293T cells and found that MYH9 could interact with USP22, especially MYH9-S1943E, but not MYH9-S1943A (Fig. xref ; Supplementary Fig. xref )."

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"This study confirmed that the p‐MYH9/USP22 complex stabilizes HIF‐1α under normoxic conditions, providing a promising therapeutic target for HCC cells that have metastasized into the bloodstream following drug resistance."
MKI67 affects USP22
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sparser
"Ki67 expression was associated with USP22 overexpression in cervical cancer, prostate cancer and oral squamous cell carcinoma [ xref , xref , xref ]."

reach
"In order to understand the role of USP22, the present study examined the association between USP22, Aurora-B, Survivin and Ki-67 in OSCC."

reach
"To further investigate the effect of USP22 on HCC progression, we first generated Huh7 cells and PLC/PRF/5 cells with stable knockdown of USP22 (shUSP22) by lentivirus infection, a scramble shRNA as a control (shCtrl) as indicated."

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"To examine whether USP22 affected the histone H2Bub or ZEB1 recruitment to the VEGFA-promoter I region, we first constructed HCCLM3 cell lines with stable knockdown of USP22 (shUSP22) by lentivirus infection, a scramble shRNA as a control (shCtrl)."
H2AC20 affects USP22
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1 | 1

sparser
"Subsequent immunoprecipitation (IP) experiments confirmed that USP22 bind to endogenous H2A in A549 cell line (Figures xref )."
GSE1 affects USP22
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GSE1 binds USP22 and RCOR1. 2 / 2
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"What is the biological function of the GSE1-USP22-CoREST complex?"

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"The GSE1-USP22-CoREST complex seems to be stable, as we do not see changes in its composition following DNA damage induction."
FO affects USP22
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FO increases the amount of USP22. 2 / 2
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"Furthermore, another study showed that FO could increase the expression and activity of the Sirt 1 protein by upregulating the expression of USP22 (Yu et al., 2020)."

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"These findings suggested that FO therapy could increase USP22 expression, which might reduce SBP and cardiac dysfunction."
Cyclin affects USP22
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Cyclin activates USP22. 2 / 2
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"USP22 function reverses cyclin D1 ubiquitylation; overexpression of cyclin D1 can partially rescue cell cycle arrest in USP22 knockdown cells."

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"Therefore, Bmi-1 and Cyclin D2 are important oncogenes in cancer and may mediate the promoting role of USP22 in human colon cancer.In conclusion, the present study systematically examined the oncogenic role of USP22 in human colon cancer."
COL17A1 affects USP22
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COL17A1 activates USP22. 2 / 2
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"Together, these results suggest that USP22 stabilizes COL17A1 by deubiquitinating COL17A1.3.6 Reexpression of COL17A1 Reverses USP22 Silencing-Induced In Vitro Phenotype Changes of LUAD Cells."

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"Since our data have demonstrated the regulation of USP22 in COL17A1 stability in LUAD cells, we next hypothesized that COL17A1 might mediate the functional effect of USP22 on LUAD progression."
CDK11B affects USP22
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sparser
"USP22 interacts with CDK11B."

sparser
"Taken together, these results indicated that USP22 interacts with CDK11B in hepatocellular carcinoma cells."
CDC20 affects USP22
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CDC20 inhibits USP22. 2 / 2
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"Therefore, APC and CDC20 E3 ligase complex promotes USP22 protein degradation, presumably allowing CCNB1 degradation for cells to exit M phase."

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"In contrast, the APC and CDC20 E3 ligase complex negatively regulates USP22 activity by targeting it for degradation, presumably allowing CCNB1 downregulation so that cells can exit mitosis and enter anaphase."
CCND1 affects USP22
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CCND1 activates USP22. 2 / 2
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"Strikingly, inhibiting downstream CCND1/CDK activity with PD-0332991 in USP22-overexpressing cells was sufficient to reverse the effect of USP22 on cell-cycle progression."

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"Levels of CCND1 in the cytoplasm decreased following depletion of either USP22 or USP2; however, both the chromatin-bound and chromatin-free pools of CCND1 were sensitive to USP22 depletion, while USP2 depletion had little effect on either pool."
Atx7 affects USP22
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Atx7 inhibits USP22. 2 / 2
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"Moreover, polyQ expansion of Atx7 decreases the deubiquitinating activity of USP22 and, consequently, increases the level of monoubiquitinated H2B."

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"The mutation of the zinc finger domain in Atx7 that disrupts its interaction with USP22 dramatically abolishes sequestration of USP22."
ATXN7L3 affects AR
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AR binds USP22 and ATXN7L3. 2 / 2
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sparser
"This result together with the observed interaction between the AR and both ATXN7L3 and USP22 suggested that these proteins are recruited together to the promoters of AR-dependent genes upon ligand ind[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"To further investigate this hypothesis, we analyzed putative interactions of USP22 or ATXN7L3 with AR in mammalian cells."

reach
"The association between USP22 and AP4 and liver, but not lymph node, metastasis may be due to these proteins driving blood stream, but not lymphatic, metastasis of CRC cells."

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"Consistent with the above results, ChIP analysis revealed that USP22 binds the promoter region of AP4."