IndraLab

Statements


USP28 affects MYC
2 6 1 | 1 2 66 33
USP28 binds MYC.
6 | 1 19 33
6 | 1 18 27

sparser
"Moreover, addition of unphosphorylated degron peptide strongly reduced the interaction between USP28 and c-MYC, presumably by titration of Usp28 protein, while the p-Thr58/p-Ser62 peptide did not affect the interaction ( xref )."

sparser
"Recent studies showed that USP28 was involved in cancer-related pathways, and regulated physiological homeostasis of ubiquitination process, DNA-damage response, and cell cycle during genotoxic stress, which suggested that USP28 could be a promising target for cancer therapy. xref USP28 required for Myc function was screened. xref USP28 bound to Myc through an interaction with Fbw7α, and catalyzed the deubiquitination of Myc, thereby promoting its stabilization and contributing to tumor cell growth in colon and breast cancers. xref , xref USP28 can also bind to and deubiquitinate some proteins involved in DNA-damage pathways."

No evidence text available

sparser
"Currently, the mechanism that USP28 interacts with c-Myc and antagonizes c-Myc degradation is controversial."

reach
"XREF_FIG shows that ADI-PEG20 treatment enhances the interaction between USP-28 and c-Myc."

sparser
"These data suggest that Usp28 binding to c-Myc can occur via the degron motif in an Fbw7-independent fashion."

sparser
"Collectively, these data suggest that lactate enhances the USP28-MYC interaction via generation of a local acidic microenvironment ( xref )."

reach
"The deubiquitinating enzyme USP28 reportedly binds to the N-terminus of FBXW7a for deubiquitination degradation and then induces c-Myc to promote GBM [118–120]."
| PMC

No evidence text available

reach
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7alpha and c-Myc, which increased the levels of Ub-c-Myc."
USP28 binds MYC and Fbw7α. 2 / 2
| 2

sparser
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7α and c-Myc, which increased the levels of Ub-c-Myc."

sparser
"Although further studies did not show that hypoxia signaling impaired the association of USP28 with Fbw7α and c-Myc, cellular USP28 protein levels were decreased by Ni ions and hypoxia in A549 cells."
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."
USP28 binds MYC and C171. 1 / 1
| 1

reach
"Its crystal structure showed that USP28 interacts with MYC through C171 and the MBI domain."
USP28 binds MYC and USP. 1 / 1
| 1

sparser
"Importantly, our results establish that the MYC MBI domain (1–95 amino acids) interacts directly with the USP domain (160–652 amino acids) of USP28."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
USP28 activates MYC.
1 | 1 1 23
USP28 activates MYC. 10 / 26
1 | 1 1 23

reach
"USP28 has also been reported to antagonize ubiquitin-dependent degradation of the oncogene product MYC as well as JUN and Notch ."

reach
"FBXW7-185aa reduced the half-life of c-Myc by antagonizing USP28 induced c-Myc stabilization."

reach
"This peptide supposes to repress glioma malignancy through USP28 induced c-Myc stabilization."

reach
"Circ-FBXW7, highly expressed in normal human brain, has been reported to encode a novel 21-kDa protein, named FBXW7-185aa, that reduces the half-life of c-Myc by antagonizing USP28-induced c-Myc stabilization, thus acting as tumor suppressor in glioblastoma cells (34)."

reach
"Since FBW7gamma lacks the USP28 interaction motif of FBW7alpha, Usp28 selectively antagonize nucleoplasmic MYC degradation while not affecting nucleolar degradation."

reach
"In a previous study, circFBXW7 was found to be downregulated in glioma, where it is translated into a 185-amino acid (aa) protein (FBXW7-185aa) that competitively interacts with the deubiquitinating enzyme USP28, preventing USP28 from binding to FBXW7 and antagonizing USP28 induced c-Myc stabilization."

trips
"FBXW7-185aa reduced the half-life of c-Myc by antagonizing USP28-induced c-Myc stabilization."

reach
"A study by Zhang et al demonstrated that circ-FBXW7 encodes FBXW7-185aa, a protein consisting of 185 amino acids that can significantly inhibit tumour cell processes in vivo and reduce the half-life of c-Myc by antagonizing the stabilization effect of c-Myc induced by USP28."

reach
"Here, using murine genetic models, we determined that USP28 antagonizes the ubiquitin dependent degradation of c-MYC, a known USP28 substrate, as well as 2 additional oncogenic factors, c-JUN and NOTCH1, in the intestine."

reach
"Upregulation of FBXW7-185aa inhibited proliferation and cell cycle progression and reduced the half-life of c-Myc by antagonizing USP28 induced c-Myc stabilization in glioblastoma, while knockdown of FBXW7-185aa promoted malignant phenotypes in vitro and in vivo [XREF_BIBR]."
| PMC
USP28 inhibits MYC.
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USP28 inhibits MYC. 8 / 8
| 8

reach
"Thereafter, qRT‐PCR assays were performed on FBXW7, SKP2, USP28, and USP36, which confirmed that Dem could interact with FBXW7 and downregulate the expression of c‐Myc."

reach
"In addition, nickel and hypoxia exposure decreased USP28, a c-Myc de-ubiquitinating enzyme, contributing to a higher steady state level of c-Myc ubiquitination and promoting c-Myc degradation."

reach
"In colon carcinoma, USP28 antagonizes the activity of the SCFFBW7 ubiquitin ligase complex to regulate the Myc stability and further promotes cell differentiation [XREF_BIBR]."

reach
"In addition, nickel and hypoxia exposure decreased USP28, a c-myc de-ubiquitinating enzyme, contributing to a higher steady state level of c-myc ubiquitination and promoting c-myc degradation."

reach
"Furthermore, overexpressing USP28 inhibited FBXW7-185aa-induced c-Myc turnover (XREF_FIG)."

reach
"Moreover, FBXW7-185aa expression positively correlates with glioblastoma patient survival and negatively correlates with malignant phenotypes by inhibiting USP28-induced c-Myc stabilization (Yang et al., 2018)."

reach
"Moreover, circFBXW7 effectively inhibits glioma proliferation and cell cycle progression by antagonizing USP28 induced c-Myc stabilization [XREF_BIBR]."

reach
"FBXW7-185aa competitively interacts with USP28 and acts as a tumor-suppressive " decoy " that prevents USP28 binding to FBXW7alpha, thus antagonizing USP28 induced c-Myc stabilization."
USP28 bound to FBXW7-185aa inhibits MYC. 1 / 1
| 1

reach
"The above evidence collectively elucidates that FBXW7-185aa competitively interacted with USP28 and " freed " FBXW7alpha to degrade c-Myc."
USP28 deubiquitinates MYC.
1 1 | 6
USP28 deubiquitinates MYC. 8 / 8
1 1 | 6

"Usp28, an ubiquitin-specific protease, binds to myc through an interaction with fbw7alpha, an f-box protein that is part of an scf-type ubiquitin ligase. Therefore, it stabilizes myc."

reach
"Recent evidence has revealed that c-Myc can be deubiquitylated and regulated by USP28, USP36 and USP37."

reach
"The resultant weakly acidic microenvironment enhances the deubiquitination of MYC by USP28, improves the stability of MYC, and activates the promoter of Slug, ultimately promoting the stem cell-like properties of breast cancer."

reach
"The product of this is lactic acid that changes the pH of the local environment which then promotes the formation of a weakly acidic microenvironment, thus consequently enhancing the deubiquitination of MYC by the deubiquitination enzyme USP28 and improving the stability of MYC [38]."

reach
"USP28 directly deubiquitinates and stabilizes MYC."

reach
"An early study showed that USP28 deubiquitinates c-Myc via interacting with Fbw7alpha whereas a recent study reveals that USP37 deubiquitinates c-Myc independently of Fbw7 and c-Myc phosphorylation."

reach
"Identified in a retroviral shRNA library screen, USP28 has been shown to decrease MYC polyubiquitination and increase MYC stability by antagonizing the activity of the SCF FBW7 ubiquitin ligase complex [XREF_BIBR]."
USP28 increases the amount of MYC.
| 6
Modified USP28 increases the amount of MYC. 4 / 4
| 4

reach
"Overexpression of USP28 in BR cells enhances c-Myc expression and hence increases ASS1 transcription upon arginine deprivation, and consequently leads to cell survival."

reach
"As shown in XREF_FIG, overexpression of USP28 in A549 prevented the loss of c-Myc, suggesting that USP28 was involved in c-Myc degradation."

reach
"Intriguingly, both ectopic expression of Usp28 in wild-type MEFs (+/+ +) and complete knockout of Usp28 (-/-) strongly and equivalently increased levels of Myc, Jun, and Notch."

reach
"As expected, depletion of USP28 remarkably decreased MYC protein and reversed epinephrine induced increase in MYC protein expression (XREF_SUPPLEMENTARY and XREF_FIG), whereas overexpression of USP28 enhanced MYC expression (XREF_SUPPLEMENTARY)."
USP28 increases the amount of MYC. 2 / 2
| 2

reach
"Moreover, USP28 positively regulated the protein level of c-Myc, and c-Myc negatively regulated the radiosensitivity of ECA109 and ECA109R cells."

reach
"Importantly, USP28 WT remarkably increased protein expression of MYC, whereas USP28 Mut reversed lactate induced MYC and SLUG (XREF_FIG)."
USP28 decreases the amount of MYC.
| 2
Modified USP28 decreases the amount of MYC. 1 / 1
| 1

reach
"These outcomes indicated that a decreased USP28 protein level caused less USP28 binding to the c, Myc, and Fbw7alpha complex, and sequentially increased the steady state ubiquitination levels of c-Myc."
USP28 decreases the amount of MYC. 1 / 1
| 1

reach
"In human tumor cell lines, shRNA mediated depletion of Usp28 downregulates Myc levels and attenuates proliferation."
USP28 ubiquitinates MYC.
| 1
USP28 leads to the ubiquitination of MYC. 1 / 1
| 1

reach
"In vivo and in vitro ubiquitination assays showed that FBXW7-185aa overexpression antagonized the de-ubiquitination of c-Myc induced by USP28 and increased c-Myc ubiquitination."
USP28 is modified
3 | 2 61
USP28 is phosphorylated.
3 | 30
USP28 is phosphorylated. 10 / 19
| 19

sparser
"The findings that support this conclusion are the following: (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2-mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5'-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2-UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5'-AZA-induced apoptosis; and (vii) combinational therapies with 5'-AZA and ectopic USP28 expression enhances the effects of 5'-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."

sparser
"The increased basal levels of DNA damage presented in cancer cells could induce phosphorylation of USP28, thereby allowing the stabilization of USP28 substrates independently of E3-ligases in cancer cells."
| PMC

sparser
"USP28 is also phosphorylated by ATR/ATM and is involved in the response to DNA damage by virtue of its ability to deubiquitinate and stabilize FBW7 (see below), the ubiquitin ligase involved in the degradation of the myc transcription factor ( xref )."

sparser
"The downstream ATM kinase CHK2 could be a potential candidate to phosphorylates USP28 following DNA damage."
| PMC

sparser
"ATM-mediated phosphorylation of USP28 and UCK1 has an opposite effect on their association with KLHL2."

sparser
"Inhibition of ATR via VE-821, but not ATM using KU55933, decreased the phosphorylation of USP28 and strongly reduced the DUB activity of USP28, resulting in an overall reduction of USP28 abundance (Fig.  xref )."

sparser
"We then examined the biological significance of ATM-mediated phosphorylation of USP28."

sparser
"The USP25 activity may not be affected upon exposure to an inhibitor blocking the phosphorylation of USP28 on serine 67 and serine It could be another therapeutic option to target the binding between ATM and USP28 and reduce the phosphorylation of USP28, thus decreasing its enzymatic activity in cancer cells with increased DNA damage levels."
| PMC

sparser
"In response to ionizing radiation, USP28 is phosphorylated in an ATM-dependent manner, potentially providing a mechanism for control of USP28 activity [3] ."

sparser
"The subsequent phosphorylation of USP28 resulted in an increased USP28 enzymatic activity and stabilization of USP28 substrates, such as MYC and ∆Np63."
USP28 is phosphorylated on S67. 6 / 6
1 | 5

sparser
"The USP25 activity may not be affected upon exposure to an inhibitor blocking the phosphorylation of USP28 on serine 67 and serine It could be another therapeutic option to target the binding between ATM and USP28 and reduce the phosphorylation of USP28, thus decreasing its enzymatic activity in cancer cells with increased DNA damage levels."
| PMC

sparser
"In response to IR, USP28 becomes phosphorylated on Ser 67 and Ser 714 in an ATM-dependent manner [ xref ]."

sparser
"Considering the phosphorylation of USP28 on serine 67 and serine 714 regulates its enzymatic activity, it could be also possible to develop drugs targeting the phospho-sites."
| PMC

sparser
"As previously indicated, it has been proved that phosphorylation of USP28 on serine 67 and serine 714 increases its enzymatic activity upon DNA damage [ xref ]."
| PMC

No evidence text available

sparser
"Upon DNA damage, the phosphorylation of USP28 on serine 67 and serine 714 by the kinase ATM increases its enzymatic activity [ xref ]."
| PMC
USP28 is phosphorylated on S714. 5 / 5
1 | 4

sparser
"Upon DNA damage, the phosphorylation of USP28 on serine 67 and serine 714 by the kinase ATM increases its enzymatic activity [ xref ]."
| PMC

sparser
"Considering the phosphorylation of USP28 on serine 67 and serine 714 regulates its enzymatic activity, it could be also possible to develop drugs targeting the phospho-sites."
| PMC

No evidence text available

sparser
"As previously indicated, it has been proved that phosphorylation of USP28 on serine 67 and serine 714 increases its enzymatic activity upon DNA damage [ xref ]."
| PMC

sparser
"In response to IR, USP28 becomes phosphorylated on Ser 67 and Ser 714 in an ATM-dependent manner [ xref ]."
USP28 is phosphorylated on S131. 1 / 1
1 |

No evidence text available
USP28 is phosphorylated on serine. 1 / 1
| 1

sparser
"This interaction results in the phosphorylation of USP28 on serine 67 and 714 upon exposure to ionizing radiation in an ATM-dependent manner [ xref , xref ]."
| PMC
USP28 is phosphorylated on tyrosine. 1 / 1
| 1

sparser
"In addition, no phosphorylation of USP28 on tyrosine residue is referenced in PhosphoSitePlus™."
USP28 is sumoylated.
| 18
USP28 is sumoylated. 10 / 13
| 13

sparser
"SUMO2-conjugated exogenous USP28 was pulled down by Ni + beads under denaturing conditions, and SUMOylated USP28 protein was detected by anti-Flag antibody."

sparser
"SUMOylation of USP28 impaired its deubiquitinating activity on HIF-1α."

sparser
"We found that USP28 is SUMOylated in normoxia but rapidly deSUMOylated under hypoxia condition."

sparser
"293T cells were co-transfected with HIF-1α, HRE-driven luciferase with renilla as an internal control, and Flag-tagged vector control, Flag-tagged USP28 WT or Flag-tagged USP28 K99R. Our co-transfection data found that the highest HIF-1α luciferase activity was achieved in USP28 K99R transfection group (Fig.  xref b), suggesting that USP28 SUMOylation regulates the transcriptional activity of HIF-1α."

sparser
"The explicit interactions between SENP1 and USP28 prompted us to further examine whether SENP1 could be a deSUMOylation enzyme for USP28 SUMOylation."

sparser
"USP28 SUMOylation regulates its deubiquitinating activity against HIF-1α."

sparser
"N-terminal sumoylation of USP28 decreases its enzymatic activity [ xref , xref ]."
| PMC

sparser
"We found that without HIF-1α, the SUMOylation of USP28 was increased (Fig.  xref d)."

sparser
"In the present study, we proved that USP28 was SUMOylated under normoxic condition but rapidly deSUMOylated during hypoxia."

sparser
"Meanwhile, SENP1 KD HCT116 cells showed increased SUMOylated USP28 when compared with SENP1 WT cells (Fig.  xref b), and overexpression of SENP1 WT, but not SENP1 CA, further increased HIF-1α abundance in the presence of USP28 (Fig.  xref a)."
USP28 is sumoylated on K99. 5 / 5
| 5

sparser
"The present study found that the deubiquitinase USP28 was SUMOylated at K99 under normoxic condition but rapidly deSUMOylated by SENP1 during hypoxia."

sparser
"The K99R mutant USP28 showed much stronger efficiency against HIF-1α ubiquitination in vivo compared with USP28 WT (Fig.  xref ), suggesting that SUMOylation of USP28 at K99 inhibits its deubiquitinating activity on HIF-1α."

sparser
"A study showed that SUMOylation at Lys99 residues of USP28 could inhibit the activity of USP28 ( xref ), indicating potential therapeutic role in colon cancer cells and NSCLC cells ( xref ; xref ; xref )."

sparser
"SUMOylation of USP28 at K99 or direct fusion of SUMO2 to USP28 inhibits its DUB activity [128] ."

sparser
"Therein, the USP28-inactivating sumoylation of USP28 at K99 occurring under normoxia is reversed by direct interaction with SENP1 [ xref ]."
USP28 is desumoylated.
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USP28 is desumoylated. 10 / 12
| 12

sparser
"Taken together, these data suggested that hypoxia and HIF-1α promoted USP28 deSUMOylation."

sparser
"It is well known that HIF-1α is the master regulator in hypoxia response, therefore HIF-1α could potentially regulate USP28 deSUMOylation."

sparser
"Our data further elucidated that SENP1-mediated USP28 deSUMOylation was indispensable for its deubiquitinating activity towards HIF-1α, indicating SENP1 associated with USP28 to fully activate the HIF-1α pathway."

sparser
"Here, we report that USP28 is a SUMOylated protein in normoxia with moderate deubiquitinating activity towards HIF-1α in vitro, while hypoxia and HIF-1α activate USP28 through SENP1-mediated USP28 deSUMOylation to further accumulate HIF-1α protein in cells."

sparser
"SENP1 deSUMOylated and enhanced USP28 deubiquitinating activity towards HIF-1α."

sparser
"Our results indicate a novel mechanism by which SENP1-mediated deSUMOylation of USP28 stabilized HIF-1α during hypoxia."

sparser
"To investigate the significance of USP28 deSUMOylation, we analyzed the protein levels of HIF-1α in cells expressing either Flag-tagged vector control, Flag-tagged USP28 WT or Flag-tagged USP28 K99R."

sparser
"Hence, induced expression of SENP1 under hypoxia promotes desumoylation and activation of USP28, which then can contribute to further stabilisation of HIFs by their deubiquitynation activity ( xref )."

sparser
"This raised the question whether SENP1 was the enzyme responsible for USP28 deSUMOylation."

sparser
"Hypoxia and HIF-1α promoted USP28 deSUMOylation."
USP28 is ubiquitinated.
| 1
USP28 is ubiquitinated. 1 / 1
| 1

sparser
"LDHA ablation also rescued USP28 ubiquitination that was inhibited by epinephrine ( xref )."
USP28 is methylated.
| 1
USP28 is methylated. 1 / 1
| 1

reach
"Our results clearly demonstrate that ATG7 knockdown inhibited usp28 transcription through mediates hypermethylation of usp28 promoter."
USP28 is produced.
| 1
USP28 is produced. 1 / 1
| 1

reach
"Moreover, upregulated TET1 directly demethylates USP28 promoter, thereby enhancing USP28 transcription and expression, which binds to CD44s protein, and remove the ubiquitin group from the ubiquitinated CD44s protein, resulting in stabilization of CD44s protein to mediate stem like property of human BC cells."
USP28 affects TP53
1 | 3 31 26
USP28 binds TP53.
| 2 24
| 2 12

sparser
"Cuella-Martin et al. show that 53BP1 bridges interactions between p53 and the deubiquitinating enzyme USP28, promoting p53-DNA interactions to globally enhance p53-dependent transcriptional programs."

sparser
"The 53BP1 BRCT Domain Mediates Bivalent Interactions with p53 and USP28."

reach
"53BP1 controls p53-dependent responses through direct binding of p53 and USP28 (Cuella-Martin et al., 2016) and promotes NHEJ-mediated DSB repair by inhibiting DNA end resection (Panier and Boulton, 2013; Setiaputra and Durocher, 2019)."

sparser
"Interestingly, the tandem BRCT domain is known to interact with p53 and USP28 ( xref ; xref ; xref ; xref )."

sparser
"Because 53BP1 is known to bind USP28 as well as p53, a possible scenario is that it bridges the two proteins so that USP28 can deubiquitinate p53, preventing the tumor suppressor from being targeted to the proteasome for degradation."
| PMC

sparser
"Thus, our data identify an important role for 53BP1-bridged p53-USP28 interaction in regulating p53 function in human cells."

sparser
"Here, 53BP1’s C-terminal BRCT domains bridge p53’s interaction with the deubiquitinating enzyme USP28, catalysing de-ubiquitination events that stimulate p53 binding to, and activation of, target gene promoters xref ."

reach
"This would be consistent with a putative cooperation between p53, 53BP1, and USP28 in tumor suppression."

sparser
"Furthermore, USP28 and p53 bind the BRCT repeats at different interfaces, and it has been shown that 53BP1 and USP28 share a coregulatory role in supporting p53 functions [20] ( Figure 1 B,C)."

sparser
"53BP1-dependent p53 modulation requires both auto-oligomerization and tandem-BRCT domain-mediated bivalent interactions with p53 and the ubiquitin-specific protease USP28."
| 9

sparser
"To determine whether V1544, G1560 and/or G1593 regulate the interaction of 53BP1 with p53 and/or USP28, we immunoprecipitated HA-tagged 53BP1 WT and mutant protein complexes from HEK293T cells ( xref )."

sparser
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats ( xref ; xref )."

sparser
"In line with our transcriptomic analyses ( xref ), our data collectively reveal a function for 53BP1-dependent bivalent interactions with USP28 and p53 in enhancing p53-promoter element interactions, thereby amplifying p53-dependent transcriptional programs."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [14,19] ."

sparser
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28-dependent deubiquitination and activation of p53, leading to cell cycle arrest xref , xref ( xref )."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."

sparser
"We therefore hypothesize that 53BP1-USP28 complexes interact with nucleoplasmic p53 pools, where they function to prime p53 DNA-binding activity."

sparser
"Strikingly, unlike unstressedp21 -/- cells that mostly lacked nuclear p53, in the stressed condition, p53 not only was stabilized in the nucleus, but also formed bright nuclear foci of various sizes co-localizing with 53BP1 and USP28 in ~30% of the cell population ( xref ), suggesting that 53BP1, USP28 and p53 interact with each other after a stressed mitosis, consistent with the known interaction between 53BP1 and p53 or USP28 ( xref ; xref ; xref )."

sparser
"These data show that 53BP1 binds independently to p53 and USP28 via distinct BRCT domain surfaces and pointed toward a potential cooperative role for USP28-53BP1 complexes in p53 regulation."
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
USP28 activates TP53.
| 3 19 2
USP28 activates TP53. 10 / 24
| 3 19 2

eidos
"Following p53 localization under such conditions , 53BP1 and USP28 can mediate p53 activation and p21-dependent cell cycle arrest [ 119 ] ."

reach
"53BP1 and USP28 mediate p53 dependent cell cycle arrest in response to centrosome loss and prolonged mitosis."

reach
"53BP1 and USP28 somehow measure mitotic duration and activate p53 upon mitotic exit."

reach
"Loss of 53BP1, USP28, and TRIM37 all prevented p53 stabilization and G1 arrest following centrosome loss."

sparser
"Previous reports showed that 53BP1 and USP28 activate p53, preventing the proliferation of cells that have an increased chance of mitotic errors [ xref ]."
| PMC

reach
"Knockout of USP28 did not alter basal levels of p53 or prevent p53 stabilization in response to doxorubicin induced DNA damage (XREF_FIG and Fig."

eidos
"USP28Delta cells were used as the control because inactivation of USP28 prevents p53 activation and G1 arrest that is observed as a consequence of delayed mitosis following centrosome loss in RPE1 cells13 ."

reach
"Following p53 localization under such conditions, 53BP1 and USP28 can mediate p53 activation and p21-dependent cell cycle arrest [119]."

reach
"Previous reports showed that 53BP1 and USP28 activate p53, preventing the proliferation of cells that have an increased chance of mitotic errors [104]."
| PMC

reach
"Deletion of TP53BP1, USP28, or TRIM37 prevented p53 elevation in response to centrosome loss but did not affect cytokinesis failure induced arrest or p53 elevation after doxorubicin induced DNA damage."
USP28 deubiquitinates TP53.
1 | 7
USP28 deubiquitinates TP53. 8 / 8
1 | 7

reach
"While 53BP1, USP28, and p53 have not yet been demonstrated to form a ternary complex, USP28 was able to deubiquitinate p53 in vitro [9]."

reach
"The signalling pathway involves 53BP1 and the deubiquitylase USP28 acting in a complex to deubiquitylate and stabilise p53, which in turn controls cell fate."

reach
"Because 53BP1 is known to bind USP28 as well as p53, a possible scenario is that it bridges the two proteins so that USP28 can deubiquitinate p53, preventing the tumor suppressor from being targeted to the proteasome for degradation."
| PMC

reach
"However, it was not clear whether USP28 deubiquitinates p53 to stabilize it or regulates its DNA binding function, or whether its impact on p53 is indirect, via other factors."

reach
"While 53BP1, USP28, and p53 have not yet been demonstrated to form a ternary complex, USP28 was able to deubiquitinate p53 in vitro [XREF_BIBR]."

reach
"The nuclear p53 accumulation caused by overexpression of wild type USP28 was not due to a specific increase in p53 mRNA levels (XREF_FIG), further supporting our observation that USP28 deubiquitinates p53 for protein stabilization."

"its interacting protein USP28 that can directly deubiquitinate p53 in vitro and ectopically stabilize p53 in vivo"

reach
"Intriguingly, 53BP1 mediates p53 activation independently of its DNA repair activity, but requiring its interacting protein USP28 that can directly deubiquitinate p53 in vitro and ectopically stabilize p53 in vivo."
USP28 decreases the amount of TP53.
| 2
USP28 decreases the amount of TP53. 2 / 2
| 2

reach
"Conversely, depletion of USP28 resulted in significantly increased K48 ubiquitination and simultaneously diminished p53 level, which could be fully reversed by addition of recombinant USP28 (XREF_FIG)."

reach
"Notably, USP28 knockdown cells had decreased expression of p53, p21 and p16 INK4a, suggesting that the effect of USP28 on cell proliferation was mediated by regulating the expression of p53, p21 and p16 INK4a."
USP28 inhibits TP53.
| 1
USP28 inhibits TP53. 1 / 1
| 1

reach
"Supporting this, tumor cells silenced for USP28 or 53BP1 have previously been shown to prevent p53 elevation and growth arrest in response to prolonged prometaphase in cancer cells with increased propensity of mitotic errors."
USP28 affects FBXW7
1 1 | 42 17
USP28 binds FBXW7.
1 | 7 17
1 | 7 16

sparser
"USP28 also interacted with FBW7 under physiological conditions, as seen by endogenous FBW7 coimmunoprecipitating with endogenous USP28 (Fig. S2 H)."

reach
"The protein FBXW7-185aa functions as a tumor suppressor by competitively binding with USP28, and preventing USP28 binding to FBXW7α, subsequently inhibiting USP28-induced c-myc stabilization."

sparser
"Alternatively, USP21 could be acting directly to deubiquitylate the substrates of the cullin RING ligase, in an analogous fashion to USP28, which associates with the cullin-1-adapter FBW7 to deubiquitylate a range of substrates, including MYC ( xref )."

sparser
"In unstressed cells, USP28 forms a complex with FBXW7 and antagonizes substrate ubiquitination, however, after DNA damage this complex dissociates promoting FBXW7-dependent substrate degradation []."

No evidence text available

sparser
"Studies have identified the molecular mechanisms responsible for the intrinsic resistance include loss of PTEN , loss of NF1, CCND1 amplification, COT overexpression, mutations in RAC1 , and loss of the USP28-FBW7 complex [ xref , xref – xref ]."

reach
"Discovery of an analog of the Fbw7 and Usp28 complex in Drosophila demonstrates that such complexes represent an evolutionary conserved regulatory paradigm (Li et al., 2013)."

sparser
"Usp28 binds to Fbw7, and Usp28 was shown to be recruited to c-Myc protein indirectly through Fbw7 (the “piggyback” model) ( xref )."

sparser
"USP28 can bind to FBXW7 and inhibit ubiquitination modification of c-Myc by the latter, thus increasing the protein stability of c-Myc ( xref )."

sparser
"Several mechanisms of intrinsic resistance have been described among whom loss of PTEN, cyclin D1 amplification, loss of NF1, RAC1 P29S mutation, HGF secretion, MAP3K8 overexpression, loss of the USP28FBW7 complex, low levels of CD8 tumor-infiltrating T cells and increased expression of the immune inhibitory molecule PD-L1 ( xref – xref )."
| 1

sparser
"In contrast to VDU2 which directly interacts with HIF-1α, USP28 forms a ternary complex with HIF-1α via its association with HIF-1α bound Fbw7 (Flügel et al., xref )."
USP28 inhibits FBXW7.
| 15
| 14

reach
"Other proteins regulating C-MYC stability include : CIP2A, a C-MYC-interacting protein that specifically inhibits PP2A activity against C-MYC; USP28, a de-ubiquitinating enzymes that antagonizes FBW7 and promotes C-MYC stability and TRUSS, a receptor for DDB1 (damage specific DNA binding protein 1)-CUL4 (Cullin 4) E3 ligase complex."

reach
"Steady-state levels of Fbw7 in Usp28 -/- MEFs were rescued by MG132 treatment, demonstrating that Fbw7 was degraded by the proteasome in the absence of Usp28."

reach
"For Myc, we also examined the effects of knock-down of USP28, a Ub specific protease that antagonizes Fbw7 dependent Myc destruction [XREF_BIBR]."

reach
"On the other hand, downregulation of Fbw7, induced by Usp28 loss is either not sufficient to promote embryonic lethality, associated with Fbxw7 knockout, or does not occur in lineages, in which Fbw7 i[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Unexpectedly, we found that genetic deletion of Usp28 rescued the lethality of Fbw7 deficient primary fibroblasts."

reach
"A study has proposed a novel mechanism by which USP28 upregulates angiogenesis by antagonizing GSK-3beta (glycogen synthase kinase-3beta) and FBW7 dependent degradation of HIF-1alpha (hypoxia inducible factor-1alpha), a major regulator of angiogenesis, carcinogenesis, and various processes by which cells adapt to hypoxic conditions."

reach
"USP28 antagonizes FBW7, an F-box protein and an important component of the SKP1-CUL1-F-BOX (SCF)-type E3 ubiquitin ligase that targets key transcriptional factors to ubiquitin-directed proteasome degradation."

reach
"The Usp28 deubiquitinase antagonizes Fbw7 mediated turnover of multiple oncoproteins, including Myc, Jun, and Notch, and promotes tumorigenesis in the intestine."

reach
"In tumor cells, USP28 antagonizes the activity of the F-box and WD repeat domain containing 7 (FBXW7) to promoting the stability of c-Myc."

reach
"Fbw7 dependent substrate ubiquitination is antagonized by the Usp28 deubiquitinase."
USP28 bound to MYC inhibits FBXW7. 1 / 1
| 1

reach
"As a ubiquitin specific protease, Usp28 binds to Myc and antagonizes the activity of Fbw7, thereby stabilizing Myc."
USP28 deubiquitinates FBXW7.
1 | 11
USP28 deubiquitinates FBXW7. 10 / 12
1 | 11

reach
"In addition, the autocatalytic ubiquitylation of Fbw7 can be antagonized by the deubiquitinase Usp28 [XREF_BIBR]."

reach
"To test whether Usp28 directly deubiquitinates Fbw7, we used an in vitro assay with immunopurified SCF (Fbw7) and recombinant E1 and E2 enzymes, which leads to the accumulation of autoubiquitinated Fb[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Indeed, experiments in MEFs and human tumor cells demonstrate that Usp28 directly deubiquitinates and stabilizes Fbw7."

"We discuss mechanisms that regulate ubiquitination by Fbw7, including ubiquitin-specific proteases such as USP28 that counteract Fbw7 activity and thereby stabilise oncoproteins."

reach
"In addition, USP28 antagonizes FBW7 mediated ubiquitination and stabilizes HIF-1alpha [XREF_BIBR, XREF_BIBR]."

reach
"Usp28 preferentially deubiquitinates Fbw7 and at intermediate levels maintains stable Fbw7 but does not allow excessive Fbw7 substrate stabilization."

reach
"USP28 deubiquitinates and stabilizes FBW7 resulting in enhanced degradation of FBW7 substrates."

reach
"Ubiquitin specific peptidase 28 (USP28), a deubiquitinating enzyme can also repress autocatalytic ubiquitination and degradation of FBXW7 (Diefenbacher et al., 2014; Schulein-Volk et al., 2014)."

reach
"It was recently shown that the deubiquitinase Usp28 deubiquitinates and stabilizes Fbxw7 [XREF_BIBR]."

reach
"Indeed, wild-type Usp28, but not the catalytically inactive C171A mutant, promoted Fbw7 deubiquitination in vivo."
USP28 activates FBXW7.
| 5
USP28 activates FBXW7. 5 / 5
| 5

reach
"In agreement with this hypothesis, treatment of Usp28 -/- MEFs with PiB, a specific Pin1 inhibitor, attenuated degradation of ectopic Fbw7."

reach
"In contrast, complete absence of Usp28 triggers autocatalytic turnover of Fbw7 and leads to Fbw7 substrate stabilization.The most likely explanation for such nonlinear pattern of regulation is that Fb[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"This result suggests that Usp28 preferentially stabilizes Fbw7 due to more efficient binding and is consistent with the idea that Fbw7 targets Usp28 to its substrates.We concluded that stabilization o[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Preferential targeting of Fbw7 by Usp28 shifts the default activity of this system toward Fbw7 substrate ubiquitination and provides a mechanism to maintain physiological levels of multiple proto-onco[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, FBXW7 induced degradation of MYC is fine tuned by the de-ubiquitylation enzyme USP28 [XREF_BIBR], which slows down poly-ubiquitylation, as well as by the E3 ligase betaTRCP, which counteracts FBXW7 by conjugating mixed Ub chains to MYC that disfavors degradation."
USP28 decreases the amount of FBXW7.
| 2
USP28 decreases the amount of FBXW7. 2 / 2
| 2

reach
"Knockdown of Usp28 in human tumor cells also decreased endogenous Fbw7 levels, suggesting that regulation by Usp28 is a conserved mechanism that maintains Fbw7 stability."

reach
"Overexpressing USP28 reduced the expression of FBXW7 and suppressed FBXW7-185aa-induced c-Myc destabilization (XREF_FIG H)."
USP28 ubiquitinates FBXW7.
| 1
USP28 leads to the ubiquitination of FBXW7. 1 / 1
| 1

reach
"In contrast, complete knockout of Usp28 triggers autocatalytic ubiquitination and degradation of Fbw7, sparing Fbw7 substrates."
USP28 increases the amount of FBXW7.
| 1
Modified USP28 increases the amount of FBXW7. 1 / 1
| 1

reach
"In contrast, in the intestine and cerebellum, complete loss of Usp28 did not reduce Fbw7 levels but downregulated Fbw7 substrates.Mechanistic analysis revealed that this biphasic response to Usp28 los[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
MYC affects USP28
6 | 1 20 33
MYC binds USP28.
6 | 1 19 33
6 | 1 18 27

sparser
"Moreover, addition of unphosphorylated degron peptide strongly reduced the interaction between USP28 and c-MYC, presumably by titration of Usp28 protein, while the p-Thr58/p-Ser62 peptide did not affect the interaction ( xref )."

sparser
"Recent studies showed that USP28 was involved in cancer-related pathways, and regulated physiological homeostasis of ubiquitination process, DNA-damage response, and cell cycle during genotoxic stress, which suggested that USP28 could be a promising target for cancer therapy. xref USP28 required for Myc function was screened. xref USP28 bound to Myc through an interaction with Fbw7α, and catalyzed the deubiquitination of Myc, thereby promoting its stabilization and contributing to tumor cell growth in colon and breast cancers. xref , xref USP28 can also bind to and deubiquitinate some proteins involved in DNA-damage pathways."

No evidence text available

sparser
"Currently, the mechanism that USP28 interacts with c-Myc and antagonizes c-Myc degradation is controversial."

reach
"XREF_FIG shows that ADI-PEG20 treatment enhances the interaction between USP-28 and c-Myc."

sparser
"These data suggest that Usp28 binding to c-Myc can occur via the degron motif in an Fbw7-independent fashion."

sparser
"Collectively, these data suggest that lactate enhances the USP28-MYC interaction via generation of a local acidic microenvironment ( xref )."

reach
"The deubiquitinating enzyme USP28 reportedly binds to the N-terminus of FBXW7a for deubiquitination degradation and then induces c-Myc to promote GBM [118–120]."
| PMC

No evidence text available

reach
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7alpha and c-Myc, which increased the levels of Ub-c-Myc."
USP28 binds MYC and Fbw7α. 2 / 2
| 2

sparser
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7α and c-Myc, which increased the levels of Ub-c-Myc."

sparser
"Although further studies did not show that hypoxia signaling impaired the association of USP28 with Fbw7α and c-Myc, cellular USP28 protein levels were decreased by Ni ions and hypoxia in A549 cells."
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."
USP28 binds MYC and C171. 1 / 1
| 1

reach
"Its crystal structure showed that USP28 interacts with MYC through C171 and the MBI domain."
USP28 binds MYC and USP. 1 / 1
| 1

sparser
"Importantly, our results establish that the MYC MBI domain (1–95 amino acids) interacts directly with the USP domain (160–652 amino acids) of USP28."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
MYC inhibits USP28.
| 1
MYC inhibits USP28. 1 / 1
| 1

reach
"Indeed, HIF increases MYC phosphorylation at Thr58, triggering MYC ubiquitination, and decreases the de-ubiquitinating enzyme USP28, promoting MYC proteasome dependent degradation [XREF_BIBR]."
USP28 affects TP53BP1
5 1 1 | 14 22
USP28 binds TP53BP1.
5 1 | 12 22
5 1 | 12 10

sparser
"Furthermore, a 53BP1-USP28 complex was identified as an activator of p53-mediated transactivation ( xref , xref , xref , xref )."

sparser
"In particular, the TTD residues V1544 and G1560, located opposite to the methyl-K/R binding pocket ( xref ) ( xref ), define a non-canonical TTD interaction surface that promotes the association of 53BP1 with USP28."

sparser
"Different from the interaction of USP28 with LSD1, USP28-mediated p53BP1 stabilization only occurs after DNA damage and no effect is found in the absence of DNA damage, suggesting that the interaction between USP28 with p53BP1 is regulated by DNA damaging."

reach
"Furthermore, a 53BP1-USP28 complex was identified as an activator of p53-mediated transactivation (Cuella-Martin et al., 2016a, Fong et al., 2016, Lambrus et al., 2016, Meitinger et al., 2016)."

sparser
"Here, we describe the TTD as the second 53BP1 domain, in addition to the tandem BRCT, that regulates the USP2853BP1 interaction ( xref ) ( xref ; xref )."

reach
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28 dependent deubiquitination and activation of p53, leading to cell cycle arrest XREF_BIBR, XREF_BIBR (XREF_FIG)."

reach
"Taken together, a body of evidence supports a model in which a 53BP1 and USP28 complex modulates p53 activity in response to centrosome loss."

No evidence text available

reach
"For example, the DNA damage response pathway is central to the maintenance of genomic stability and both members of a key DDR complex, the TP53BP1 and USP28 complex, which are substrates of the ATM kinase, were identified within the top 110 TSGs (q-value < 0.15, XREF_FIG, XREF_FIG)."

sparser
"USP28 has been reported to regulate TP53 abundance in a USP28-TP53BP1 cell cycle-dependent fashion [ xref – xref ]."
| 9

sparser
"To determine whether V1544, G1560 and/or G1593 regulate the interaction of 53BP1 with p53 and/or USP28, we immunoprecipitated HA-tagged 53BP1 WT and mutant protein complexes from HEK293T cells ( xref )."

sparser
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats ( xref ; xref )."

sparser
"In line with our transcriptomic analyses ( xref ), our data collectively reveal a function for 53BP1-dependent bivalent interactions with USP28 and p53 in enhancing p53-promoter element interactions, thereby amplifying p53-dependent transcriptional programs."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [14,19] ."

sparser
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28-dependent deubiquitination and activation of p53, leading to cell cycle arrest xref , xref ( xref )."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."

sparser
"We therefore hypothesize that 53BP1-USP28 complexes interact with nucleoplasmic p53 pools, where they function to prime p53 DNA-binding activity."

sparser
"Strikingly, unlike unstressedp21 -/- cells that mostly lacked nuclear p53, in the stressed condition, p53 not only was stabilized in the nucleus, but also formed bright nuclear foci of various sizes co-localizing with 53BP1 and USP28 in ~30% of the cell population ( xref ), suggesting that 53BP1, USP28 and p53 interact with each other after a stressed mitosis, consistent with the known interaction between 53BP1 and p53 or USP28 ( xref ; xref ; xref )."

sparser
"These data show that 53BP1 binds independently to p53 and USP28 via distinct BRCT domain surfaces and pointed toward a potential cooperative role for USP28-53BP1 complexes in p53 regulation."
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
USP28 activates TP53BP1.
| 2
USP28 activates TP53BP1. 2 / 2
| 2

reach
"Different from the interaction of USP28 with LSD1, USP28 mediated p53BP1 stabilization only occurs after DNA damage and no effect is found in the absence of DNA damage, suggesting that the interaction between USP28 with p53BP1 is regulated by DNA damaging."

reach
"USP28 mediates centrosome loss induced G1 arrest through its deubiquitinase activity, and acts downstream of 53BP1 to stabilize p53."
USP28 deubiquitinates TP53BP1.
1 |
USP28 deubiquitinates TP53BP1. 1 / 1
1 |
TP53BP1 affects USP28
5 1 | 14 22
TP53BP1 binds USP28.
5 1 | 12 22
5 1 | 12 10

sparser
"Furthermore, a 53BP1-USP28 complex was identified as an activator of p53-mediated transactivation ( xref , xref , xref , xref )."

sparser
"In particular, the TTD residues V1544 and G1560, located opposite to the methyl-K/R binding pocket ( xref ) ( xref ), define a non-canonical TTD interaction surface that promotes the association of 53BP1 with USP28."

sparser
"Different from the interaction of USP28 with LSD1, USP28-mediated p53BP1 stabilization only occurs after DNA damage and no effect is found in the absence of DNA damage, suggesting that the interaction between USP28 with p53BP1 is regulated by DNA damaging."

reach
"Furthermore, a 53BP1-USP28 complex was identified as an activator of p53-mediated transactivation (Cuella-Martin et al., 2016a, Fong et al., 2016, Lambrus et al., 2016, Meitinger et al., 2016)."

sparser
"Here, we describe the TTD as the second 53BP1 domain, in addition to the tandem BRCT, that regulates the USP2853BP1 interaction ( xref ) ( xref ; xref )."

reach
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28 dependent deubiquitination and activation of p53, leading to cell cycle arrest XREF_BIBR, XREF_BIBR (XREF_FIG)."

reach
"Taken together, a body of evidence supports a model in which a 53BP1 and USP28 complex modulates p53 activity in response to centrosome loss."

No evidence text available

reach
"For example, the DNA damage response pathway is central to the maintenance of genomic stability and both members of a key DDR complex, the TP53BP1 and USP28 complex, which are substrates of the ATM kinase, were identified within the top 110 TSGs (q-value < 0.15, XREF_FIG, XREF_FIG)."

sparser
"USP28 has been reported to regulate TP53 abundance in a USP28-TP53BP1 cell cycle-dependent fashion [ xref – xref ]."
| 9

sparser
"To determine whether V1544, G1560 and/or G1593 regulate the interaction of 53BP1 with p53 and/or USP28, we immunoprecipitated HA-tagged 53BP1 WT and mutant protein complexes from HEK293T cells ( xref )."

sparser
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats ( xref ; xref )."

sparser
"In line with our transcriptomic analyses ( xref ), our data collectively reveal a function for 53BP1-dependent bivalent interactions with USP28 and p53 in enhancing p53-promoter element interactions, thereby amplifying p53-dependent transcriptional programs."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [14,19] ."

sparser
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28-dependent deubiquitination and activation of p53, leading to cell cycle arrest xref , xref ( xref )."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."

sparser
"We therefore hypothesize that 53BP1-USP28 complexes interact with nucleoplasmic p53 pools, where they function to prime p53 DNA-binding activity."

sparser
"Strikingly, unlike unstressedp21 -/- cells that mostly lacked nuclear p53, in the stressed condition, p53 not only was stabilized in the nucleus, but also formed bright nuclear foci of various sizes co-localizing with 53BP1 and USP28 in ~30% of the cell population ( xref ), suggesting that 53BP1, USP28 and p53 interact with each other after a stressed mitosis, consistent with the known interaction between 53BP1 and p53 or USP28 ( xref ; xref ; xref )."

sparser
"These data show that 53BP1 binds independently to p53 and USP28 via distinct BRCT domain surfaces and pointed toward a potential cooperative role for USP28-53BP1 complexes in p53 regulation."
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
TP53BP1 deubiquitinates USP28.
| 1
TP53BP1 leads to the deubiquitination of USP28. 1 / 1
| 1

reach
"Mechanistically, as well as its role in DNA repair, 53BP1 promotes p53 signalling, by coordinating ubiquitin specific peptidase 28 (USP28)-mediated p53 deubiquitination."
TP53BP1 activates USP28.
| 1
TP53BP1 activates USP28. 1 / 1
| 1

reach
"Given that p53 activity is quenched via MDM2 dependent p53 ubiquitination, it is tempting to speculate that a 53BP1 dependent targeting of USP28 into p53 protein complexes might counteract such events."
USP28 affects KDM1A
4 1 | 26 9
USP28 binds KDM1A.
4 | 10 9
4 | 10 9

No evidence text available

sparser
"Our results are in accordance with an older study that showed that the USP28LSD1 axis controlled CSC-like properties in human breast cancer [ xref ]."

No evidence text available

reach
"In this study, we found that USP28 interacted with and stabilized LSD1 from ubiquitination."

sparser
"Different from the interaction of USP28 with LSD1, USP28-mediated p53BP1 stabilization only occurs after DNA damage and no effect is found in the absence of DNA damage, suggesting that the interaction between USP28 with p53BP1 is regulated by DNA damaging."

sparser
"The interaction between LSD1 and USP28 is specific, as LSD1 could not interact with USP7 and USP25 under similar conditions ( xref )."

reach
"Together, these data indicate that USP28 interacts with LSD1 and this interaction is mediated through the AO domain of LSD1 and N-terminal region of USP28."

sparser
"To investigate whether USP28 interacted with LSD1, we co-expressed Flag-LSD1 and HA-USP28 in HEK293 cells."

reach
"To examine whether interaction of HDAC5 with LSD1 and USP28 complex deacetylates LSD1 or USP28, in vitro protein acetylation assays was first carried out by incubating GST tagged recombinant HDAC5 protein with cellular pull-down of LSD1-FLAG or USP28-FLAG by IP, and immunoprecipitates of IgG was incubated with recombinant HDAC5 protein as negative control of assays (XREF_FIG)."

reach
"USP28 interacted with and stabilized LSD1 via deubiquitination."
USP28 activates KDM1A.
| 6
USP28 activates KDM1A. 6 / 6
| 6

reach
"In agreement with this notion, knockdown of USP28 reduced the protein level and functional activity of LSD1, but these could be rescued by exogenous LSD1 expression (XREF_FIG)."

reach
"Overexpression of USP28 largely reversed HDAC5-KD-induced LSD1 protein degradation, suggesting a role of HDAC5 as a positive regulator of LSD1 through upregulation of USP28 protein."

reach
"To test whether USP28 directly regulates protein stability of LSD1, we co-expressed LSD1 with USP28 or vector control in HEK293 cells and examined the degradation of LSD1."

reach
"First, our data indicates that USP28 is a specific DUB for LSD1, as knockdown of USP28 reduced LSD1 stability, whereas overexpression of USP28 stabilized LSD1."

reach
"Overexpression of USP28 largely reversed HDAC5-knockdown-induced LSD1 protein degradation, suggesting HDAC5 positively regulates LSD1 by stabilizing USP28."

reach
"Interestingly, the downregulation of LSD1 by two independent USP28 shRNAs in HCT116 cells could be restored by MG132 treatment (XREF_FIG), indicating that USP28 enhances LSD1 stabilization through a deubiquitination event."
USP28 increases the amount of KDM1A.
| 4
USP28 increases the amount of KDM1A. 3 / 3
| 3

reach
"In this study, we found that USP28 was the major factor responsible for LSD1 stabilization in cancer cell lines and tumor samples, and that knockdown of USP28 decreased the level of LSD1 and suppressed CSC like properties in vitro and tumorigenicity in vivo."

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)."

reach
"When these DUBs were co-expressed with LSD1 in HEK293 cells, we noticed that USP28 significantly increased LSD1 level, similar to that treated with MG132 (XREF_SUPPLEMENTARY)."
Modified USP28 increases the amount of KDM1A. 1 / 1
| 1

reach
"The steady-state level of LSD1 was enhanced by ectopic USP28 expression in a dose dependent manner (XREF_FIG)."
USP28 deubiquitinates KDM1A.
1 | 3
USP28 deubiquitinates KDM1A. 3 / 3
1 | 2

reach
"We found that WT-USP28, but not CI-USP28, specifically removed LSD1 ubiquitination (XREF_FIG), indicating that USP28 directly deubiquitinated LSD1."

reach
"USP7 and USP28 inhibited LSD1 ubiquitination and stabilized LSD1 protein level [XREF_BIBR, XREF_BIBR]."
Modified USP28 leads to the deubiquitination of KDM1A. 1 / 1
| 1

reach
"However, co-expression of WT-USP28, but not CI-USP28, almost completely abolished LSD1 ubiquitination (lane 3 vs. lane 4, XREF_FIG)."
USP28 inhibits KDM1A.
| 2
USP28 inhibits KDM1A. 2 / 2
| 2

reach
"Overexpression of USP28 largely reversed HDAC5-knockdown-induced LSD1 protein degradation, suggesting HDAC5 positively regulates LSD1 by stabilizing USP28."

reach
"To understand whether HDAC5 may stabilize LSD1 protein through upregulation of USP28 protein stability, a rescue study was carried out in MDA-MB-231 and MCF10A-CA1a cells using concurrent transfection of HDAC5 siRNA and USP28 expression plasmids, and showed that overexpression of USP28 completely blocked the destabilization of LSD1 by HDAC5 depletion (XREF_FIG, XREF_SUPPLEMENTARY)."
USP28 decreases the amount of KDM1A.
| 1
USP28 decreases the amount of KDM1A. 1 / 1
| 1

reach
"Our study showing that increase of LSD1 protein expression by Jade-2 siRNA and decrease of LSD1 protein expression by USP28 siRNA in MDA-MB-231 cells confirmed the roles of Jade-2 and USP28 as LSD1 ubiquitin ligase and deubiquitinase in breast cancer cells (XREF_FIG; XREF_SUPPLEMENTARY)."
USP28 affects STAT3
| 27 8
USP28 binds STAT3.
| 7 8
| 7 8

reach
"USP28 interacts with STAT3 and decreases its polyubiquitination level."

sparser
"USP28 interacts with STAT3 and decreases its polyubiquitination level."

sparser
"We also found that USP28 interacted with STAT3, and stabilized STAT3 by decreasing its polyubiquitination, which indicated that STAT3 was a potential substrate of USP28 ( xref and xref )."

reach
"Immunoprecipitation was performed to assess whether USP28 interacted with STAT3 or deubiquitinated STAT3."

sparser
"USP28 interacted with STAT3 and decreased the polyubiquitination of STAT3, thereby increasing the stability of STAT3."

reach
"USP28 interacted with STAT3 and decreased the polyubiquitination of STAT3, thereby increasing the stability of STAT3."

reach
"We also found that USP28 interacted with STAT3, and stabilized STAT3 by decreasing its polyubiquitination, which indicated that STAT3 was a potential substrate of USP28 (XREF_FIG and XREF_FIG)."

sparser
"To evaluate whether USP28 interacted with STAT3, the reciprocal Co-IP was performed."

sparser
"Additionally, the Co-IP in xref showed that endogenous STAT3 also interacted with endogenous USP28."

sparser
"As shown in xref , the Co-IP revealed that exogenous USP28 interacted with exogenous STAT3 protein."
USP28 activates STAT3.
| 13
USP28 activates STAT3. 10 / 13
| 13

reach
"Moreover, the CHX chase assay revealed that overexpression of USP28 prolonged the half-life of STAT3 protein in NSCLC cells, which further suggested that USP28 stabilized STAT3 protein (XREF_FIG)."

reach
"USP28 increases the stability of STAT3."

reach
"These results demonstrated that USP28 was functional in NSCLC cells, and promoted NSCLC cell growth by inducing STAT3 signaling."

reach
"As shown in XREF_FIG, overexpression of USP28 upregulated STAT3 protein in a dose dependent manner, but the mRNA level of STAT3 was not changed obviously (XREF_FIG)."

reach
"In addition, the immunoblotting assay also revealed that overexpression of USP28 enhanced the STAT3 signaling in NSCLC cells (XREF_FIG)."

reach
"USP28 is highly expressed and mediates STAT3 signaling by stabilizing STAT3 in NSCLC cells, which suggests that USP28 can be a potential target for NSCLC therapy in the future."

reach
"Herein, we identified that USP28, a deubiquitinase aberrantly upregulated in patients with ADPKD, selectively removed K48-linked polyubiquitination and reversed protein degradation of signal transducer and activator of transcription 3 (STAT3)."

reach
"As shown in XREF_FIG, overexpression of USP28 significantly upregulated STAT3 derived luciferase activity, but another USP isoform, USP25, had no effect on STAT3 luciferase activity."

reach
"USP28 mediates STAT3 signaling in NSCLC cells."

reach
"As stated before, USP28 mediated STAT3 signaling by stabilizing STAT3 protein, which suggested that USP28 was functional in NSCLC cells."
USP28 increases the amount of STAT3.
| 5
USP28 increases the amount of STAT3. 5 / 5
| 5

reach
"In contrast, knockdown of USP28 suppressed STAT3 derived luciferase activity (XREF_FIG), and inhibited the expression of STAT3 in NSCLC cells (XREF_FIG)."

reach
"Whole cell lysates were also prepared for immunoblotting, which showed that USP28 upregulated the expression level of STAT3 (XREF_FIG)."

reach
"In addition, the immunoblotting also showed that knockdown of USP28 downregulated STAT3 expression in vivo (XREF_FIG)."

reach
"The whole cell lysates also revealed that USP28 increased the protein level of STAT3 (XREF_FIG)."

reach
"In contrast, knockdown of USP28 downregulated the protein level of STAT3 in NSCLC cells (XREF_FIG), but the mRNA level did not show significant change (XREF_FIG)."
USP28 inhibits STAT3.
| 1
USP28 inhibits STAT3. 1 / 1
| 1

reach
"Herein, we identified that USP28, a deubiquitinase aberrantly upregulated in patients with ADPKD, selectively removed K48-linked polyubiquitination and reversed protein degradation of signal transducer and activator of transcription 3 (STAT3)."
USP28 deubiquitinates STAT3.
| 1
USP28 deubiquitinates STAT3. 1 / 1
| 1

reach
"As shown in XREF_FIG, the IP indicated that STAT3 itself was polyubiquitinated and USP28 deubiquitinated the polyubiquitination of STAT3."

reach
"Functional assays demonstrated that overexpression of USP28 promoted cell proliferation and aerobic glycolysis of colorectal cancer, while USP28 inhibition could reverse these effects."

reach
"USP28 increases aerobic glycolysis and promotes cell proliferation upon stabilization of FOXC1 [50]."
| PMC

reach
"Further, lack of USP28 promotes a more malignant state of breast cancer cells, indicated by an epithelial-to-mesenchymal (EMT) transition, elevated proliferation, migration, and angiogenesis as well as a decreased adhesion."

reach
"Down-regulation of USP28 promoted NSCLC cells proliferation and induced cells apoptosis."

reach
"Previous reports showed that 53BP1 and USP28 activate p53, preventing the proliferation of cells that have an increased chance of mitotic errors [104]."
| PMC

reach
"Moreover, the upregulation of CCND1 and USP28 partially reverses the impaired proliferation and promoted apoptosis rate induced by miR-3940-5p in A549 cells (XREF_FIG, E-H)."

reach
"XREF_BIBR reported that overexpression of USP28 promotes NSCLC cell proliferation and apoptosis inhibition and is associated with poor prognosis in NSCLC patients."

reach
"We found that up-regulation of USP28 in NSCLC cell lines promoted cells proliferation."

reach
"USP28 contributes to the proliferation and metastasis of gastric cancer."

eidos
"USP28 increases aerobic glycolysis and promotes cell proliferation upon stabilization of FOXC1 [ 50 ] ."
| PMC

reach
"Downregulation of CCND1 and USP28 Inhibits Proliferation and Induces Apoptosis in A549 Cells."

reach
"Downregulating USP28 expression can inhibit the proliferation and growth of lung cancer cells [XREF_BIBR, XREF_BIBR]."

reach
"Moreover, 53BP1 or USP28 deletion restored NPC proliferation and brain size without correcting the upstream centrosome defects or extended mitosis."

reach
"MTT assays showed that USP28 overexpression reduced cell proliferation in MDA-MB-231 and MCF7 cells, and these effects were partially reversed by transfection with miR-500a-5p mimics."

reach
"CCK-8 assay shows that si-CCND1 and si-USP28 significantly inhibit A549 cell proliferation compared with si-controls (XREF_FIG, E and F)."

reach
"Additionally, according to the study by Popov, et al., depletion of USP28 in HeLa and LS174T cells can inhibit cell growth and proliferation due to its inability to enhance c-Myc stability [XREF_BIBR]."

reach
"Importantly, silencing KLHL2 or USP28 overexpression not only inhibited AML cell proliferation but also sensitized AML cells to 5 '-AZA-induced apoptosis in vitro and in vivo."
FBXW7 affects USP28
1 | 11 17
FBXW7 binds USP28.
1 | 7 17
1 | 7 16

sparser
"USP28 also interacted with FBW7 under physiological conditions, as seen by endogenous FBW7 coimmunoprecipitating with endogenous USP28 (Fig. S2 H)."

reach
"The protein FBXW7-185aa functions as a tumor suppressor by competitively binding with USP28, and preventing USP28 binding to FBXW7α, subsequently inhibiting USP28-induced c-myc stabilization."

sparser
"Alternatively, USP21 could be acting directly to deubiquitylate the substrates of the cullin RING ligase, in an analogous fashion to USP28, which associates with the cullin-1-adapter FBW7 to deubiquitylate a range of substrates, including MYC ( xref )."

sparser
"In unstressed cells, USP28 forms a complex with FBXW7 and antagonizes substrate ubiquitination, however, after DNA damage this complex dissociates promoting FBXW7-dependent substrate degradation []."

No evidence text available

sparser
"Studies have identified the molecular mechanisms responsible for the intrinsic resistance include loss of PTEN , loss of NF1, CCND1 amplification, COT overexpression, mutations in RAC1 , and loss of the USP28-FBW7 complex [ xref , xref – xref ]."

reach
"Discovery of an analog of the Fbw7 and Usp28 complex in Drosophila demonstrates that such complexes represent an evolutionary conserved regulatory paradigm (Li et al., 2013)."

sparser
"Usp28 binds to Fbw7, and Usp28 was shown to be recruited to c-Myc protein indirectly through Fbw7 (the “piggyback” model) ( xref )."

sparser
"USP28 can bind to FBXW7 and inhibit ubiquitination modification of c-Myc by the latter, thus increasing the protein stability of c-Myc ( xref )."

sparser
"Several mechanisms of intrinsic resistance have been described among whom loss of PTEN, cyclin D1 amplification, loss of NF1, RAC1 P29S mutation, HGF secretion, MAP3K8 overexpression, loss of the USP28FBW7 complex, low levels of CD8 tumor-infiltrating T cells and increased expression of the immune inhibitory molecule PD-L1 ( xref – xref )."
| 1

sparser
"In contrast to VDU2 which directly interacts with HIF-1α, USP28 forms a ternary complex with HIF-1α via its association with HIF-1α bound Fbw7 (Flügel et al., xref )."
FBXW7 inhibits USP28.
| 2
FBXW7 inhibits USP28. 1 / 1
| 1

reach
"Furthermore, expression of dominant negative Fbw7 or Fbw7 targeting shRNAs enhanced proliferation of Usp28 +/- MEFs, suggesting that their proliferative defect is, at least in part, mediated by the ac[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
FBXW7-S205A inhibits USP28. 1 / 1
| 1

reach
"Stable expression of the S205A mutant Fbw7, but not of the wild-type protein, strongly downregulated Fbw7 substrates and attenuated proliferation of the Usp28 -/- MEFs."
FBXW7 activates USP28.
| 2
FBXW7 activates USP28. 2 / 2
| 2

reach
"Furthermore, expression of dominant negative Fbw7 or Fbw7 targeting shRNAs enhanced proliferation of Usp28 +/- MEFs, suggesting that their proliferative defect is, at least in part, mediated by the ac[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"This result suggests that Usp28 preferentially stabilizes Fbw7 due to more efficient binding and is consistent with the idea that Fbw7 targets Usp28 to its substrates.We concluded that stabilization o[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
TP53 affects USP28
| 3 24
TP53 binds USP28.
| 2 24
| 2 12

sparser
"Cuella-Martin et al. show that 53BP1 bridges interactions between p53 and the deubiquitinating enzyme USP28, promoting p53-DNA interactions to globally enhance p53-dependent transcriptional programs."

sparser
"The 53BP1 BRCT Domain Mediates Bivalent Interactions with p53 and USP28."

reach
"53BP1 controls p53-dependent responses through direct binding of p53 and USP28 (Cuella-Martin et al., 2016) and promotes NHEJ-mediated DSB repair by inhibiting DNA end resection (Panier and Boulton, 2013; Setiaputra and Durocher, 2019)."

sparser
"Interestingly, the tandem BRCT domain is known to interact with p53 and USP28 ( xref ; xref ; xref ; xref )."

sparser
"Because 53BP1 is known to bind USP28 as well as p53, a possible scenario is that it bridges the two proteins so that USP28 can deubiquitinate p53, preventing the tumor suppressor from being targeted to the proteasome for degradation."
| PMC

sparser
"Thus, our data identify an important role for 53BP1-bridged p53-USP28 interaction in regulating p53 function in human cells."

sparser
"Here, 53BP1’s C-terminal BRCT domains bridge p53’s interaction with the deubiquitinating enzyme USP28, catalysing de-ubiquitination events that stimulate p53 binding to, and activation of, target gene promoters xref ."

reach
"This would be consistent with a putative cooperation between p53, 53BP1, and USP28 in tumor suppression."

sparser
"Furthermore, USP28 and p53 bind the BRCT repeats at different interfaces, and it has been shown that 53BP1 and USP28 share a coregulatory role in supporting p53 functions [20] ( Figure 1 B,C)."

sparser
"53BP1-dependent p53 modulation requires both auto-oligomerization and tandem-BRCT domain-mediated bivalent interactions with p53 and the ubiquitin-specific protease USP28."
| 9

sparser
"To determine whether V1544, G1560 and/or G1593 regulate the interaction of 53BP1 with p53 and/or USP28, we immunoprecipitated HA-tagged 53BP1 WT and mutant protein complexes from HEK293T cells ( xref )."

sparser
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats ( xref ; xref )."

sparser
"In line with our transcriptomic analyses ( xref ), our data collectively reveal a function for 53BP1-dependent bivalent interactions with USP28 and p53 in enhancing p53-promoter element interactions, thereby amplifying p53-dependent transcriptional programs."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [14,19] ."

sparser
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28-dependent deubiquitination and activation of p53, leading to cell cycle arrest xref , xref ( xref )."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."

sparser
"We therefore hypothesize that 53BP1-USP28 complexes interact with nucleoplasmic p53 pools, where they function to prime p53 DNA-binding activity."

sparser
"Strikingly, unlike unstressedp21 -/- cells that mostly lacked nuclear p53, in the stressed condition, p53 not only was stabilized in the nucleus, but also formed bright nuclear foci of various sizes co-localizing with 53BP1 and USP28 in ~30% of the cell population ( xref ), suggesting that 53BP1, USP28 and p53 interact with each other after a stressed mitosis, consistent with the known interaction between 53BP1 and p53 or USP28 ( xref ; xref ; xref )."

sparser
"These data show that 53BP1 binds independently to p53 and USP28 via distinct BRCT domain surfaces and pointed toward a potential cooperative role for USP28-53BP1 complexes in p53 regulation."
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
TP53 activates USP28.
| 1
TP53 activates USP28. 1 / 1
| 1

reach
"Moreover, purified USP28 was found to directly deubiquitinate p53 in vitro (XREF_FIG), raising potentially a direct role of USP28 in stabilizing p53 in vivo."
KDM1A affects USP28
4 | 12 9
KDM1A binds USP28.
4 | 10 9
4 | 10 9

No evidence text available

sparser
"Our results are in accordance with an older study that showed that the USP28LSD1 axis controlled CSC-like properties in human breast cancer [ xref ]."

No evidence text available

reach
"In this study, we found that USP28 interacted with and stabilized LSD1 from ubiquitination."

sparser
"Different from the interaction of USP28 with LSD1, USP28-mediated p53BP1 stabilization only occurs after DNA damage and no effect is found in the absence of DNA damage, suggesting that the interaction between USP28 with p53BP1 is regulated by DNA damaging."

sparser
"The interaction between LSD1 and USP28 is specific, as LSD1 could not interact with USP7 and USP25 under similar conditions ( xref )."

reach
"Together, these data indicate that USP28 interacts with LSD1 and this interaction is mediated through the AO domain of LSD1 and N-terminal region of USP28."

sparser
"To investigate whether USP28 interacted with LSD1, we co-expressed Flag-LSD1 and HA-USP28 in HEK293 cells."

reach
"To examine whether interaction of HDAC5 with LSD1 and USP28 complex deacetylates LSD1 or USP28, in vitro protein acetylation assays was first carried out by incubating GST tagged recombinant HDAC5 protein with cellular pull-down of LSD1-FLAG or USP28-FLAG by IP, and immunoprecipitates of IgG was incubated with recombinant HDAC5 protein as negative control of assays (XREF_FIG)."

reach
"USP28 interacted with and stabilized LSD1 via deubiquitination."
KDM1A inhibits USP28.
| 1
KDM1A inhibits USP28. 1 / 1
| 1

reach
"Disrupting USP28 destabilizes LSD1 protein, which decreases breast cancer stem cell like characteristics in vitro and suppresses tumorigenicity in vivo [XREF_BIBR]."
KDM1A activates USP28.
| 1
KDM1A activates USP28. 1 / 1
| 1

reach
"Although our results indicate that LSD1 is a major target of USP28 in breast cancer, it does not rule out the possibility that USP28 promotes CSC like traits through both LSD1 stabilization and inhibition of Myc degradation."
USP28 affects UCK1
| 12 9
USP28 binds UCK1.
| 6 8
| 6 4

sparser
"Although KLHL2 and UCK1 formed binary complexes, USP28 did not directly bind to UCK1."

reach
"Given that several DUBs are recruited to their substrates indirectly through binding to the E3 ligase XREF_BIBR, we analyzed whether KLHL2 mediated the interaction between UCK1 and USP28 in vivo."

reach
"USP28 binds to UCK1 via KLHL2."

reach
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28."

reach
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 (XREF_SUPPLEMENTARY B)."

sparser
"Given that several DUBs are recruited to their substrates indirectly through binding to the E3 ligase xref , we analyzed whether KLHL2 mediated the interaction between UCK1 and USP28 in vivo ."

sparser
"Our data strongly suggest that KLHL2 mediates the interaction between UCK1 and USP28."

reach
"Our data strongly suggest that KLHL2 mediates the interaction between UCK1 and USP28."

sparser
"USP28 does not bind directly with UCK1, but binds via KLHL2 [ xref ]."

reach
"The findings that support this conclusion are the following : (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2 mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5 '-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2 and UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5 '-AZA-induced apoptosis; and (vii) combinational therapies with 5 '-AZA and ectopic USP28 expression enhances the effects of 5 '-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."
USP28 binds UCK1 and KLHL2. 3 / 3
| 3

sparser
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28 (Figure xref G)."

sparser
"USP28 binds to UCK1 via KLHL2."

sparser
"The findings that support this conclusion are the following: (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2-mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5'-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2-UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5'-AZA-induced apoptosis; and (vii) combinational therapies with 5'-AZA and ectopic USP28 expression enhances the effects of 5'-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
USP28 deubiquitinates UCK1.
| 3
USP28 deubiquitinates UCK1. 3 / 3
| 3

reach
"We then evaluated whether USP28 could deubiquitinate UCK1."

reach
"In vitro deubiquitination assay further confirmed that WT-USP28, but not USP28-CA, efficiently deubiquitinated UCK1, indicating that USP28 served as direct DUB for UCK1."

reach
"Conversely, USP28 silencing in HEK293T markedly increased the ubiquitination of endogenous UCK1, implying that endogenous UCK1 was also a target of USP28."
USP28 ubiquitinates UCK1.
| 1 1
USP28 leads to the ubiquitination of UCK1. 2 / 2
| 1 1

reach
"We next examined the effect of USP28 on KLHL2 mediated ubiquitination of UCK1."

sparser
"Conversely, USP28 silencing in HEK293T markedly increased the ubiquitination of endogenous UCK1 (Figure xref F), implying that endogenous UCK1 was also a target of USP28."
USP28 inhibits UCK1.
| 1
USP28 inhibits UCK1. 1 / 1
| 1

reach
"Conversely, depletion of endogenous USP28 by shRNA in MV4-11 cells reduced the half-life of UCK1."
USP28 activates UCK1.
| 1
USP28 activates UCK1. 1 / 1
| 1

reach
"These data were further supported by the observation that USP28 significantly rescued elevated instability of UCK1 protein in the presence of KLHL2."
ATM affects USP28
4 | 9 6
ATM phosphorylates USP28.
4 | 8 6
ATM phosphorylates USP28. 10 / 11
| 5 6

sparser
"These novel findings establish a direct connection between the ubiquitination of UCK1 by KLHL2 and phosphorylation of USP28 and UCK1 by ATM, which are involved in mediating drug resistance against 5'-AZA in AML patients."

sparser
"In response to DNA damage, phosphorylation of the deubiquitinase USP28 by DDR kinases, ATM, and ATR leads to deubiquitination of either GATA4 or its associated factor [ xref ]."

reach
"In response to 5 '-AZA, UCK1 and USP28 are phosphorylated by ATM, instigating entirely different effects on the protein protein interaction."

reach
"The downstream ATM kinase CHK2 could be a potential candidate to phosphorylates USP28 following DNA damage."
| PMC

sparser
"Once all the three conservative SQ sites (S67, S495, and S714), in USP28, were mutated to alanine, the pS/TQ antibody completely could not recognize the immunoprecipitated Flag-USP28-3S/A (Figure xref D), suggesting that in response to 5'-AZA-induced stress, ATM phosphorylates USP28 at all these three sites."

sparser
"We next asked whether ATM or ATR phosphorylates USP28 upon CPPD treatment."

sparser
"In response to 5'-AZA, UCK1 and USP28 are phosphorylated by ATM, instigating entirely different effects on the protein-protein interaction."

reach
"USP28 is also phosphorylated by ATR and ATM and is involved in the response to DNA damage by virtue of its ability to deubiquitinate and stabilize FBW7 (see below), the ubiquitin ligase involved in the degradation of the myc transcription factor."

sparser
"It cannot be excluded the possibility that kinases other than ATM could phosphorylate USP28 under DNA damage stress."
| PMC

reach
"In response to DNA damage, phosphorylation of the deubiquitinase USP28 by DDR kinases, ATM, and ATR leads to deubiquitination of either GATA4 or its associated factor [XREF_BIBR]."
ATM phosphorylates USP28 on S714. 2 / 2
1 | 1

No evidence text available

reach
"Upon DNA damage, the phosphorylation of USP28 on serine 67 and serine 714 by the kinase ATM increases its enzymatic activity [10]."
| PMC
ATM phosphorylates USP28 on S495. 1 / 1
1 |

No evidence text available
ATM phosphorylates USP28 at position 67. 1 / 1
| 1

reach
"Upon DNA damage, the phosphorylation of USP28 on serine 67 and serine 714 by the kinase ATM increases its enzymatic activity [10]."
| PMC
ATM phosphorylates USP28 on S67. 1 / 1
1 |

No evidence text available
ATM phosphorylates USP28 on serine. 1 / 1
| 1

reach
"Upon DNA damage, the phosphorylation of USP28 on serine 67 and serine 714 by the kinase ATM increases its enzymatic activity [10]."
| PMC
ATM phosphorylates USP28 on S720. 1 / 1
1 |

No evidence text available
ATM activates USP28.
| 1
ATM activates USP28. 1 / 1
| 1

reach
"Since 5 '-AZA treatment can activate ATM in AML cells and the ATM pathway can prevent the apoptosis induction in 5 '-AZA resistant cells XREF_BIBR, we sought to determine whether ATM modulated the effect of USP28 on the UCK1 deubiquitination in response to 5 '-AZA."
UCK1 affects USP28
| 7 8
UCK1 binds USP28.
| 6 8
| 6 4

sparser
"Although KLHL2 and UCK1 formed binary complexes, USP28 did not directly bind to UCK1."

reach
"Given that several DUBs are recruited to their substrates indirectly through binding to the E3 ligase XREF_BIBR, we analyzed whether KLHL2 mediated the interaction between UCK1 and USP28 in vivo."

reach
"USP28 binds to UCK1 via KLHL2."

reach
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28."

reach
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 (XREF_SUPPLEMENTARY B)."

sparser
"Given that several DUBs are recruited to their substrates indirectly through binding to the E3 ligase xref , we analyzed whether KLHL2 mediated the interaction between UCK1 and USP28 in vivo ."

sparser
"Our data strongly suggest that KLHL2 mediates the interaction between UCK1 and USP28."

reach
"Our data strongly suggest that KLHL2 mediates the interaction between UCK1 and USP28."

sparser
"USP28 does not bind directly with UCK1, but binds via KLHL2 [ xref ]."

reach
"The findings that support this conclusion are the following : (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2 mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5 '-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2 and UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5 '-AZA-induced apoptosis; and (vii) combinational therapies with 5 '-AZA and ectopic USP28 expression enhances the effects of 5 '-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."
USP28 binds UCK1 and KLHL2. 3 / 3
| 3

sparser
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28 (Figure xref G)."

sparser
"USP28 binds to UCK1 via KLHL2."

sparser
"The findings that support this conclusion are the following: (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2-mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5'-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2-UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5'-AZA-induced apoptosis; and (vii) combinational therapies with 5'-AZA and ectopic USP28 expression enhances the effects of 5'-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
UCK1 activates USP28.
| 1
UCK1 activates USP28. 1 / 1
| 1

reach
"Conversely, USP28 silencing in HEK293T markedly increased the ubiquitination of endogenous UCK1, implying that endogenous UCK1 was also a target of USP28."
STAT3 affects USP28
| 7 8
| 7 8

reach
"USP28 interacts with STAT3 and decreases its polyubiquitination level."

sparser
"USP28 interacts with STAT3 and decreases its polyubiquitination level."

sparser
"We also found that USP28 interacted with STAT3, and stabilized STAT3 by decreasing its polyubiquitination, which indicated that STAT3 was a potential substrate of USP28 ( xref and xref )."

reach
"Immunoprecipitation was performed to assess whether USP28 interacted with STAT3 or deubiquitinated STAT3."

sparser
"USP28 interacted with STAT3 and decreased the polyubiquitination of STAT3, thereby increasing the stability of STAT3."

reach
"USP28 interacted with STAT3 and decreased the polyubiquitination of STAT3, thereby increasing the stability of STAT3."

reach
"We also found that USP28 interacted with STAT3, and stabilized STAT3 by decreasing its polyubiquitination, which indicated that STAT3 was a potential substrate of USP28 (XREF_FIG and XREF_FIG)."

sparser
"To evaluate whether USP28 interacted with STAT3, the reciprocal Co-IP was performed."

sparser
"Additionally, the Co-IP in xref showed that endogenous STAT3 also interacted with endogenous USP28."

sparser
"As shown in xref , the Co-IP revealed that exogenous USP28 interacted with exogenous STAT3 protein."
CHEK2 affects USP28
| 6 8
CHEK2 binds USP28.
| 4 7
| 4 1

sparser
"It was previously shown that USP28 interacts with CHK2 [ xref ]."
| PMC

reach
"USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2 mediated polyubiquitylation and proteasomal degradation of CHK2 [XREF_BIBR]."

reach
"Similar to FBXW7, USP28 forms a complex with PIRH2 and CHK2, thus antagonizing the ubiquitination of CHK2 by PIRH2.The protein PIRH2 (also called RCHY1) is a promising candidate for maintaining the previously demonstrated USP28 “piggyback” model."
| PMC

reach
"We show that the deubiquitylating enzyme USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2 mediated polyubiquitylation and proteasomal degradation of CHK2."

reach
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 21."
| 5

sparser
"Similar to FBXW7, USP28 forms a complex with PIRH2 and CHK2, thus antagonizing the ubiquitination of CHK2 by PIRH2."
| PMC

sparser
"We show that the deubiquitylating enzyme USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK221."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2 [ xref ]."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 xref ."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
CHEK2 phosphorylates USP28.
| 2 1
CHEK2 phosphorylates USP28. 3 / 3
| 2 1

reach
"However, analyses of the USP28 sequence did not reveal any CHK2 phosphorylation motif, suggesting that USP28 cannot be phosphorylated by CHK2."
| PMC

sparser
"However, analyses of the USP28 sequence did not reveal any CHK2 phosphorylation motif, suggesting that USP28 cannot be phosphorylated by CHK2."
| PMC

reach
"The downstream ATM kinase CHK2 could be a potential candidate to phosphorylates USP28 following DNA damage."
| PMC
USP28 affects cell growth
| 12 1
USP28 activates cell growth.
| 8
| 8

reach
"As shown in XREF_FIG, knockdown of USP28 significantly inhibited NSCLC cell growth in both A549 and H1299 cells."

reach
"These results demonstrated that USP28 was functional in NSCLC cells, and promoted NSCLC cell growth by inducing STAT3 signaling."

reach
"Overexpression of USP28 accelerated PC cell growth, whereas USP28 knockdown impaired PC cell growth both in vitro and in vivo."

reach
"XREF_BIBR Previous study has reported that USP28 was overexpressed in NSCLC and overexpression of USP28 promoted NSCLC cell growth, but its mechanism was still unknown in NSCLC cells."

reach
"Further, we found that USP28 promoted PC cell growth by facilitating cell cycle progression and inhibiting apoptosis."

reach
"Furthermore, USP28 regulates the expression of MYC protein, which is essential in USP28 induced cell growth in glioma cells."

reach
"Importantly, USP28 promoted MYC expression, which was required for USP28 induced cell growth in human glioma."

reach
"Moreover, knockdown of USP28 inhibited cell growth of NSCLC cells in vitro and delayed NSCLC tumor growth in vivo."
USP28 inhibits cell growth.
| 4 1
| 4 1

reach
"Knockdown of USP28 suppressed the cell growth of NSCLC both in vitro and in vivo."

sparser
"Moreover, knockdown of USP28 inhibited cell growth of NSCLC cells in vitro and delayed NSCLC tumor growth in vivo."

reach
"Previous study demonstrated that depletion of USP28 inhibited both cell-cycle and cell growth through regulation of MYC abundance XREF_BIBR."

reach
"Knockdown of USP28 inhibits NSCLC cell growth."

reach
"Additionally, according to the study by Popov, et al., depletion of USP28 in HeLa and LS174T cells can inhibit cell growth and proliferation due to its inability to enhance c-Myc stability [XREF_BIBR]."
USP28 affects CLSPN
1 1 1 | 9 1
USP28 deubiquitinates CLSPN.
1 | 3
USP28 deubiquitinates CLSPN. 4 / 4
1 | 3

reach
"Claspin is a bona fide USP28 substrate, as Claspin deubiquitination by USP28 could also be reconstituted in vitro."

reach
"Thus, USP28 deubiquitinates Claspin to enable the activation of a CHK1-dependent G -checkpoint , and protects checkpoint-mediators to facilitate apoptosis ."

reach
"De-ubiquitinase USP28 inhibits ubiquitination of Claspin protein and its proteasomal degradation [XREF_BIBR]."
USP28 binds CLSPN.
1 1 | 1 1
1 1 | 1

No evidence text available

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."

No evidence text available
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
USP28 activates CLSPN.
| 4
USP28 activates CLSPN. 4 / 4
| 4

reach
"These results suggest that USP28 specifically enhances Claspin protein stabilization through a de-ubiquitination event."

reach
"The specificity of DUBs, however, is poorly understood, and it is currently unknown how Usp28 targets Claspin but not Plk1."

reach
"Additionally, in this system, USP28 only targets Claspin but not PLK1 (both of which are subject to ubiquitination by the same E3 ligase)."

reach
"In conclusion, when faced with DNA damage, HNF-1beta-overexpressing cells conferred them with cell survival activity that is mediated through the USP28 mediated Claspin stabilization and then persistent Chk1 activation."
USP28 decreases the amount of CLSPN.
| 1
USP28 decreases the amount of CLSPN. 1 / 1
| 1

reach
"Knockdown of endogenous USP28 suppressed the Claspin expression and p-Chk1 activation and cell viability."
MiR-3940-5p affects USP28
| 12
MiR-3940-5p inhibits USP28.
| 5
MiR-3940-5p inhibits USP28. 5 / 5
| 5

reach
"As illustrated in XREF_FIG, A-D, compared with control cells, CCND1 and USP28 are downregulated at both mRNA and protein levels by ectopic expression of miR-3940-5p; however, pcDNA-CCND1 and pcDNA-USP28 robustly overturn this decreased expression of CCND1 and USP28 induced by miR-3940-5p."

reach
"Furthermore, animal study also confirmed that overexpression of miR-3940-5p suppresses the growth of NSCLC tumors and negatively regulates CCND1 and USP28 in vivo."

reach
"These results confirmed our speculation that miR-3940-5p inhibits NSCLC cell proliferation and promotes apoptosis by targeting CCND1 and USP28."

reach
"In line with binding data, the results show that miR-3940-5p markedly suppresses the luciferase activity of the WT CCND1 or USP28 3 ' UTR without effect on the MUT 3 ' UTR, which indicates that CCND1 and USP28 are direct targets of miR-3940-5p (XREF_FIG, C and D)."

reach
"Results show that CCND1 and USP28 protein expression levels are decreased compared with the control group (XREF_FIG C), suggesting that miR-3940-5p negatively regulates CCND1 and USP28 in vivo."
MiR-3940-5p activates USP28.
| 5
MiR-3940-5p activates USP28. 5 / 5
| 5

reach
"These results confirmed our speculation that miR-3940-5p inhibits proliferation and induces apoptosis in NSCLC cells by targeting CCND1 and USP28."

reach
"In a different study, however, miR-3940-5p was identified to target USP28 and CCND1 (Cyclin D1) in NSCLC."

reach
"Cell proliferation and apoptosis assays showed that miR-3940-5p suppressed proliferation and promoted apoptosis in NSCLC cells, and silencing CCND1 and USP28 both recapitulated the effects of miR-3940-5p on NSCLC cells."

reach
"Therefore, miR-3940-5p may target CCND1 and USP28 to inhibit NSCLC growth."

reach
"Based on these findings, we speculated that miR-3940-5p might target CCND1 and USP28 to inhibit NSCLC growth."
MiR-3940-5p decreases the amount of USP28.
| 2
MiR-3940-5p decreases the amount of USP28. 2 / 2
| 2

reach
"As illustrated in XREF_FIG, A-D, compared with control cells, CCND1 and USP28 are downregulated at both mRNA and protein levels by ectopic expression of miR-3940-5p; however, pcDNA-CCND1 and pcDNA-USP28 robustly overturn this decreased expression of CCND1 and USP28 induced by miR-3940-5p."

reach
"In addition, a significant negative correlation was found between expressions of miR-3940-5p versus CCND1 mRNA (r = -0.794; XREF_FIG D) and USP28 mRNA (r = -0.709; XREF_FIG E), indicating that miR-3940-5p either directly or indirectly suppressed CCND1 and USP28 transcript levels."
| 12
| 9

reach
"Moreover, the upregulation of CCND1 and USP28 partially reverses the impaired proliferation and promoted apoptosis rate induced by miR-3940-5p in A549 cells (XREF_FIG, E-H)."

reach
"Down-regulation of USP28 promoted NSCLC cells proliferation and induced cells apoptosis."

reach
"Down-regulation of USP28 induced cell apoptosis."

reach
"Resistance to vincristine in DLBCL by disruption of p53-induced cell cycle arrest and apoptosis mediated by KIF18B and USP28."

reach
"Hence, silencing of USP28 antagonized apoptosis induced by caspase-8 depletion and also reverted accumulation and activation of p53 (XREF_FIG)."

reach
"USP28 was identified as an interaction partner of 53BP1 [125], and loss of USP28 has been reported to lead to IR-induced apoptosis in H460 cells, in a similar manner to what has been seen in Chk2, p53 and PUMA null mice."

reach
"Moreover, FACS apoptosis analysis indicated that down-regulation of USP28 expression by siRNA increased the NSCLC cells apoptosis rate."

reach
"Downregulation of CCND1 and USP28 Inhibits Proliferation and Induces Apoptosis in A549 Cells."

reach
"Commentary on: Rovsing et al. Resistance to vincristine in DLBCL by disruption of p53-induced cell cycle arrest and apoptosis mediated by KIF18B and USP28."
| 3

reach
"These results indicate that CCND1 and USP28 are upregulated in A549 cells, and knockdown of CCND1 or USP28 obviously restrains cell proliferation and promotes apoptosis."

reach
"Further, we found that USP28 promoted PC cell growth by facilitating cell cycle progression and inhibiting apoptosis."

reach
"XREF_BIBR reported that overexpression of USP28 promotes NSCLC cell proliferation and apoptosis inhibition and is associated with poor prognosis in NSCLC patients."
USP28 affects CHEK2
| 5 7
USP28 binds CHEK2.
| 4 7
| 4 1

sparser
"It was previously shown that USP28 interacts with CHK2 [ xref ]."
| PMC

reach
"USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2 mediated polyubiquitylation and proteasomal degradation of CHK2 [XREF_BIBR]."

reach
"Similar to FBXW7, USP28 forms a complex with PIRH2 and CHK2, thus antagonizing the ubiquitination of CHK2 by PIRH2.The protein PIRH2 (also called RCHY1) is a promising candidate for maintaining the previously demonstrated USP28 “piggyback” model."
| PMC

reach
"We show that the deubiquitylating enzyme USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2 mediated polyubiquitylation and proteasomal degradation of CHK2."

reach
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 21."
| 5

sparser
"Similar to FBXW7, USP28 forms a complex with PIRH2 and CHK2, thus antagonizing the ubiquitination of CHK2 by PIRH2."
| PMC

sparser
"We show that the deubiquitylating enzyme USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK221."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2 [ xref ]."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 xref ."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
USP28 activates CHEK2.
| 1
USP28 activates CHEK2. 1 / 1
| 1

reach
"Interestingly, in another study, CHK2 is shown to be directly regulated by USP28."
USP28 affects BRAF
1 | 10
USP28 inhibits BRAF.
| 7
USP28 inhibits BRAF. 7 / 7
| 7

reach
"The loss of USP28 promoted MAPK activation and resistance to RAF inhibitor therapy by stabilizing BRAF in cell culture and in vivo models."

reach
"Both knockdown of USP28 and FBW7 significantly enhanced endogenous BRAF stability (XREF_FIG)."

reach
"Loss of USP28 enhances BRAF stabilization and confers resistance to vemurafenib in melanoma."

reach
"Loss of USP28 mediated BRAF degradation drives resistance to RAF cancer therapies."

reach
"Having established that USP28 and FBW7 reduces BRAF stability, we tested the effect of USP28 and FBW7 depletion on BRAF expression."

reach
"Importantly, of the 59 patients harboring BRAF V600E mutations 27% (16/59) displayed a> 50% decrease in USP28 mRNA expression levels, suggesting that in tumors harboring BRAF alterations, loss of USP28 may further increase the tumorigenic potential of these tumors by stabilizing BRAF and enhancing downstream MAPK activation."

reach
"However, we noted that loss of USP28 or FBW7 did not fully prevent BRAF degradation suggesting that BRAF degradation may occur through mechanisms independent of the USP28 and FBW7 axis."
USP28 activates BRAF.
| 2
USP28 activates BRAF. 2 / 2
| 2

reach
"As USP28 deubiquitinates and stabilizes FBW7, allowing the FBW7 and SCF ligase complex to bind and degrade substrates containing a Cdc4 phosphodegron motif, we hypothesized that forced expression of USP28 would target BRAF for degradation."

reach
"As expected USP28 mediated BRAF stability led to enhanced pERK levels after vemurfenib treatment compared with wild-type USP28 cells."
USP28 deubiquitinates BRAF.
1 |
USP28 deubiquitinates BRAF. 1 / 1
1 |

"Here, we demonstrate that the deubiquitinating enzyme USP28 functions through a feedback loop to destabilize RAF family members."
USP28 decreases the amount of BRAF.
| 1
Modified USP28 decreases the amount of BRAF. 1 / 1
| 1

reach
"Indeed, overexpression of USP28 decreased the concentrations of ectopically expressed and endogenous BRAF levels (XREF_FIG)."
TP53BP1 affects TP53
| 11
| 9

sparser
"To determine whether V1544, G1560 and/or G1593 regulate the interaction of 53BP1 with p53 and/or USP28, we immunoprecipitated HA-tagged 53BP1 WT and mutant protein complexes from HEK293T cells ( xref )."

sparser
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats ( xref ; xref )."

sparser
"In line with our transcriptomic analyses ( xref ), our data collectively reveal a function for 53BP1-dependent bivalent interactions with USP28 and p53 in enhancing p53-promoter element interactions, thereby amplifying p53-dependent transcriptional programs."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [14,19] ."

sparser
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28-dependent deubiquitination and activation of p53, leading to cell cycle arrest xref , xref ( xref )."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."

sparser
"We therefore hypothesize that 53BP1-USP28 complexes interact with nucleoplasmic p53 pools, where they function to prime p53 DNA-binding activity."

sparser
"Strikingly, unlike unstressedp21 -/- cells that mostly lacked nuclear p53, in the stressed condition, p53 not only was stabilized in the nucleus, but also formed bright nuclear foci of various sizes co-localizing with 53BP1 and USP28 in ~30% of the cell population ( xref ), suggesting that 53BP1, USP28 and p53 interact with each other after a stressed mitosis, consistent with the known interaction between 53BP1 and p53 or USP28 ( xref ; xref ; xref )."

sparser
"These data show that 53BP1 binds independently to p53 and USP28 via distinct BRCT domain surfaces and pointed toward a potential cooperative role for USP28-53BP1 complexes in p53 regulation."
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."
SENP1 affects USP28
| 3 8
SENP1 binds USP28.
| 1 6
| 1 6

reach
"The co-immunoprecipitation and GST Pull-down assays were used to determine the interaction between USP28 and SENP1."

sparser
"Firstly, we tested whether SENP1 could interact with USP28."

sparser
"These data indicate that the deSUMOylation enzyme SENP1 directly interacted with USP28."

sparser
"SENP1 directly interacts with USP28."

sparser
"In addition, GST pull-down assay was performed to further study if the interaction between USP28 and SENP1 was direct."

sparser
"The co-immunoprecipitation and GST Pull-down assays were used to determine the interaction between USP28 and SENP1."

sparser
"The explicit interactions between SENP1 and USP28 prompted us to further examine whether SENP1 could be a deSUMOylation enzyme for USP28 SUMOylation."
SENP1 desumoylates USP28.
| 2 2
SENP1 desumoylates USP28. 3 / 3
| 1 2

sparser
"SENP1 strongly desumoylates USP28 upon hypoxia [ xref ]."
| PMC

sparser
"SENP1 deSUMOylated and enhanced USP28 deubiquitinating activity towards HIF-1α."

reach
"SENP1 strongly desumoylates USP28 upon hypoxia [14]."
| PMC
Modified SENP1 leads to the desumoylation of USP28. 1 / 1
| 1

reach
"Hence, induced expression of SENP1 under hypoxia promotes desumoylation and activation of USP28, which then can contribute to further stabilisation of HIFs by their deubiquitynation activity (XREF_FIG)."
USP28 affects RCHY1
1 | 4 5
| 5

sparser
"Similar to FBXW7, USP28 forms a complex with PIRH2 and CHK2, thus antagonizing the ubiquitination of CHK2 by PIRH2."
| PMC

sparser
"We show that the deubiquitylating enzyme USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK221."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2 [ xref ]."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 xref ."
1 | 4

reach
"USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2 mediated polyubiquitylation and proteasomal degradation of CHK2 [XREF_BIBR]."

reach
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 21."

No evidence text available

reach
"We show that the deubiquitylating enzyme USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2 mediated polyubiquitylation and proteasomal degradation of CHK2."

reach
"Similar to FBXW7, USP28 forms a complex with PIRH2 and CHK2, thus antagonizing the ubiquitination of CHK2 by PIRH2.The protein PIRH2 (also called RCHY1) is a promising candidate for maintaining the previously demonstrated USP28 “piggyback” model."
| PMC
USP28 affects cell cycle
| 9
USP28 activates cell cycle.
| 7
| 7

reach
"Our results furthermore explain how USP28 may modulate the p53 dependent cell cycle checkpoint, thereby controlling tumor cell fate decisions as previously described."

reach
"In addition to arresting centrosome deficient cells, p53, 53BP1, and USP28 are all involved in the DNA damage response (DDR), raising the possibility that centrosome loss causes a cell cycle arrest because it somehow induces DNA damage."
| PMC

reach
"Further, we found that USP28 promoted PC cell growth by facilitating cell cycle progression and inhibiting apoptosis."

reach
"We showed that 53BP1 and USP28 are required to trigger p53 and p21- dependent cell cycle arrest, evoking an irreversible stress response that selects against unfit cells with disturbed mitosis (XREF_FIG)."

reach
"Consistently, overexpression of the wild type USP28 but not USP28 CI in normal, unstressed cells caused ectopic nuclear p53 accumulation and cell cycle arrest uniformly across the entire population (100%, XREF_FIG; not shown)."

reach
"53BP1 and USP28 mediate p53 dependent cell cycle arrest in response to centrosome loss and prolonged mitosis."

reach
"Similarly, western blot analyses revealed that the total p53 levels in 53BP1 -/- or USP28 -/- cells were kept low during acentrosomal cell division in the presence of mitotic delay (XREF_FIG), indicating that 53BP1 and USP28 function upstream of p53 to initiate cell cycle arrest in response to centrosome loss."
USP28 inhibits cell cycle.
| 2
| 2

reach
"As expected, USP28 overexpression further decreased the relative expression of cell cycle related genes (CDK4, CDK6, Cyclin D1, pRB, and E2F1) induced by 5 '-AZA, whereas p15 and p27 expression was significantly enhanced."

reach
"Conversely, USP28 silencing accelerated the cell cycle progression to G2/M phase."
USP28 affects HIF1A
1 | 4 4
USP28 inhibits HIF1A.
| 4
USP28 inhibits HIF1A. 4 / 4
| 4

reach
"GSK-3beta regulates cell growth, migration, and angiogenesis via Fbw7 and USP28 dependent degradation of HIF-1alpha."

reach
"In conclusion, knockdown of USP28 enhanced the radiosensitivity of EC cells by destabilizing c-Myc and enhancing the accumulation of HIF-1alpha."

reach
"In their following study, they found that GSK3beta regulated cell growth, migration and angiogenesis via Fbw7 and USP28 dependent degradation of HIF1alpha [9]."

reach
"A study has proposed a novel mechanism by which USP28 upregulates angiogenesis by antagonizing GSK-3beta (glycogen synthase kinase-3beta) and FBW7 dependent degradation of HIF-1alpha (hypoxia inducible factor-1alpha), a major regulator of angiogenesis, carcinogenesis, and various processes by which cells adapt to hypoxic conditions."
USP28 sumoylates HIF1A.
| 2
USP28 sumoylates HIF1A. 2 / 2
| 2

sparser
"To investigate the impact of USP28 SUMOylation on HIF-1α, we generated Flag-USP28 K99R mutant plasmid."

sparser
"Due to lack of the deSUMOylation activity by HIF-1α, we hypothesized that hypoxia and HIF-1α decrease USP28 SUMOylation through a deSUMOylation enzyme."
USP28 binds HIF1A.
| 2
| 1

sparser
"In contrast to VDU2 which directly interacts with HIF-1α, USP28 forms a ternary complex with HIF-1α via its association with HIF-1α bound Fbw7 (Flügel et al., xref )."
| 1

sparser
"To investigate whether HIF-1α interacts with the USP28 promoter, since this factor was required for the loss of USP28 , another ChIP assay using anti-HIF-1α antibody was conducted."
USP28 deubiquitinates HIF1A.
1 |
USP28 deubiquitinates HIF1A. 1 / 1
1 |
TP53 affects TP53BP1, and USP28
| 9
| 9

sparser
"To determine whether V1544, G1560 and/or G1593 regulate the interaction of 53BP1 with p53 and/or USP28, we immunoprecipitated HA-tagged 53BP1 WT and mutant protein complexes from HEK293T cells ( xref )."

sparser
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats ( xref ; xref )."

sparser
"In line with our transcriptomic analyses ( xref ), our data collectively reveal a function for 53BP1-dependent bivalent interactions with USP28 and p53 in enhancing p53-promoter element interactions, thereby amplifying p53-dependent transcriptional programs."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [14,19] ."

sparser
"While much remains to be learned about how the mitotic surveillance pathway functions to survey centrosomes, a plausible model is that in response to centrosome loss, 53BP1 binds to USP28 and p53 to facilitate USP28-dependent deubiquitination and activation of p53, leading to cell cycle arrest xref , xref ( xref )."

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."

sparser
"We therefore hypothesize that 53BP1-USP28 complexes interact with nucleoplasmic p53 pools, where they function to prime p53 DNA-binding activity."

sparser
"Strikingly, unlike unstressedp21 -/- cells that mostly lacked nuclear p53, in the stressed condition, p53 not only was stabilized in the nucleus, but also formed bright nuclear foci of various sizes co-localizing with 53BP1 and USP28 in ~30% of the cell population ( xref ), suggesting that 53BP1, USP28 and p53 interact with each other after a stressed mitosis, consistent with the known interaction between 53BP1 and p53 or USP28 ( xref ; xref ; xref )."

sparser
"These data show that 53BP1 binds independently to p53 and USP28 via distinct BRCT domain surfaces and pointed toward a potential cooperative role for USP28-53BP1 complexes in p53 regulation."
LDHA affects USP28
| 7 2
LDHA activates USP28.
| 5
LDHA activates USP28. 5 / 5
| 5

reach
"Chronic stress induced epinephrine enhances LDHA dependent metabolic activity, which increases lactate and augments USP28 that serves to stabilize the MYC protein."

reach
"Also, high glucose consistently triggered HK2 and LDHA expression and stimulated the USP28 and MYC axis in breast cancer cells (XREF_SUPPLEMENTARY)."

reach
"Following treatment with the LDHA inhibitor sodium oxamate, we found a reduction in USP28 stabilization induced by LDHA in a dose dependent manner (XREF_FIG and XREF_SUPPLEMENTARY)."

reach
"LDHA generating lactate enhances the USP28 signaling."

reach
"Silencing of LDHA significantly reversed induction of USP28 and MYC by epinephrine (XREF_FIG), while silencing of HK2 displayed no change on the effect of epinephrine (XREF_SUPPLEMENTARY)."
LDHA binds USP28.
| 2
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."
LDHA ubiquitinates USP28.
| 1
LDHA leads to the ubiquitination of USP28. 1 / 1
| 1

reach
"LDHA ablation also rescued USP28 ubiquitination that was inhibited by epinephrine (XREF_FIG)."
LDHA decreases the amount of USP28.
| 1
LDHA decreases the amount of USP28. 1 / 1
| 1

reach
"Moreover, the reduction in expression of USP28 caused by LDHA knockdown was reversed by MG132 (XREF_FIG)."
USP28 affects Ubiquitin
| 8
| 8

reach
"Additionally, a deubiquitinase, USP28, has been reported to antagonize ubiquitin dependent proteasomal degradation of c-Myc."

reach
"Interestingly, USP28 can also antagonize the ubiquitin dependent degradation of two additional oncogenic proteins, c-Jun and Notch1, expanding the substrate list that is shared by both USP28 and FBW7."

reach
"Here, using murine genetic models, we determined that USP28 antagonizes the ubiquitin dependent degradation of c-MYC, a known USP28 substrate, as well as 2 additional oncogenic factors, c-JUN and NOTCH1, in the intestine."

reach
"These results showed that the UIM of USP28 most likely enhances the cleavage of Ub chains by binding to the proximal Ub of K11-linked di-Ub and positioning the substrate correctly in the active site."

reach
"USP28 has also been reported to antagonize ubiquitin-dependent degradation of the oncogene product MYC as well as JUN and Notch 105 ."

reach
"We transfected HeLa cells with FLAG tagged Myc or Fbw7 together with HA tagged ubiquitin and increasing amounts of Usp28 and recovered ubiquitinated proteins by immunoprecipitation."

reach
"Interestingly, USP28 can also antagonize the ubiquitin-dependent degradation of two additional oncogenic proteins, c-Jun and Notch1, expanding the substrate list that is shared by both USP28 and FBW7."

reach
"USP28 has also been reported to antagonize ubiquitin-dependent degradation of the oncogene product MYC as well as JUN and Notch ."
USP28 affects USP28
| 8
USP28 decreases the amount of USP28.
| 2
USP28 decreases the amount of USP28. 2 / 2
| 2

reach
"In addition, Ni ions and hypoxia increased the levels of the gene silencing mark dimethylated H3 lysine 9 at USP28 promoter region, which suppressed USP28 gene expression, further depleting the cell of this c-Myc salvaging enzyme."

reach
"Ni (II) and hypoxia increased the levels of the gene silencing mark H3K9me2 (substrate of iron dependent dioxygenase) at USP28 promoter region, which suppressed USP28 gene expression."
USP28 activates USP28.
| 2
USP28 activates USP28. 2 / 2
| 2

reach
"These findings suggested that ATM mediated phosphorylation of USP28 promotes dissociation of USP28 from KLHL2, destabilizing UCK1."

reach
"Nickel ions or hypoxia could impact the de-ubiquitination process by directly inhibiting USP28 activity or indirectly decreasing USP28 activity through either the dissociation of USP28 from the c, Myc, and Fbw7alpha complex or by a reduction of cellular USP28 levels."
USP28 sumoylates USP28.
| 1
USP28 sumoylates USP28 on K99. 1 / 1
| 1

reach
"Therein, the USP28 inactivating sumoylation of USP28 at K99 occurring under normoxia is reversed by direct interaction with SENP1 [XREF_BIBR]."
USP28 methylates USP28.
| 1
USP28 leads to the methylation of USP28. 1 / 1
| 1

reach
"Our results clearly demonstrate that ATG7 knockdown inhibited usp28 transcription through mediates hypermethylation of usp28 promoter."
USP28 inhibits USP28.
| 1
USP28 inhibits USP28. 1 / 1
| 1

reach
"Thus, complete loss of Usp28 promotes Pin1 dependent degradation of Fbw7 by the proteasome, most likely via enhanced autocatalytic ubiquitination.Destabilization of Fbw7 in Usp28 -/- cells suggested t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 increases the amount of USP28.
| 1
Unmethylated USP28 increases the amount of USP28. 1 / 1
| 1

reach
"Upregulated TET1 then directly demethylates the ubiquitin specific peptidase 28 (USP28) promoter, thereby enhancing USP28 transcription and expression."
∆NP63 affects USP28
| 7
USP28 binds ∆NP63. 7 / 7
| 7

sparser
"The ∆Np63USP28 axis is required to maintain the identity of SCC cells."

sparser
"To assess whether the observed genetic alterations are attributed to the USP28∆Np63 axis in SCC, we next analysed “Hallmark” gene sets of reported USP28 substrates NOTCH1, MYC and AP1‐c‐JUN, in USP28 and ∆Np63 knock‐down A‐431 cells (Fig  xref J and K)."

sparser
"Therefore, we were able to show that the USP28∆Np63 axis is implicated in the regulation and control of the expression of SCC marker genes as well as genes controlling cell fate and SCC identity."

sparser
"To investigate a therapeutic potential of targeting the USP28∆Np63 axis in SCC cells of different origins, we used a set of human cancer cell lines, comprising the pancreas lines PANC‐1 (ADC) and BXPC‐3 (SCC); cervical cancer cell lines HeLa (ADC), SiHa and Ca Ski (SCC); the head‐and‐neck cell line Detroit 562 (SCC); and the lung cell lines H1299 (ADC) and LUDLU‐1 adh (SCC)."

sparser
"While the interaction between USP28 and ∆Np63 is independent of the catalytic site within the DUB, our results show that the active cysteine at position 171 is essential to facilitate the stabilizing effect on the ∆Np63 protein."

sparser
"Alternatively, SCC tumors express high levels of USP28 and the oncogenic transcription factors NOTCH1, c-JUN, c-MYC, and ∆Np63 ( xref B). squamous tumors depend on the USP28-∆Np63 axis to maintain SCC cell identity and induce the formation of tumors in vivo."
| PMC

sparser
"Finally, we targeted the USP28∆Np63 axis via AZ1 in SCC cell lines and compared proliferative responses to a respective ADC line (Figs  xref G and H, and xref D)."
7 |
Valproic acid increases the amount of USP28.
6 |
Valproic acid increases the amount of USP28. 6 / 6
6 |

No evidence text available

No evidence text available

No evidence text available

No evidence text available

No evidence text available

No evidence text available
Valproic acid methylates USP28.
1 |
1 |

No evidence text available
USP28 affects ∆NP63
| 7
USP28 binds ∆NP63. 7 / 7
| 7

sparser
"The ∆Np63USP28 axis is required to maintain the identity of SCC cells."

sparser
"To assess whether the observed genetic alterations are attributed to the USP28∆Np63 axis in SCC, we next analysed “Hallmark” gene sets of reported USP28 substrates NOTCH1, MYC and AP1‐c‐JUN, in USP28 and ∆Np63 knock‐down A‐431 cells (Fig  xref J and K)."

sparser
"Therefore, we were able to show that the USP28∆Np63 axis is implicated in the regulation and control of the expression of SCC marker genes as well as genes controlling cell fate and SCC identity."

sparser
"To investigate a therapeutic potential of targeting the USP28∆Np63 axis in SCC cells of different origins, we used a set of human cancer cell lines, comprising the pancreas lines PANC‐1 (ADC) and BXPC‐3 (SCC); cervical cancer cell lines HeLa (ADC), SiHa and Ca Ski (SCC); the head‐and‐neck cell line Detroit 562 (SCC); and the lung cell lines H1299 (ADC) and LUDLU‐1 adh (SCC)."

sparser
"While the interaction between USP28 and ∆Np63 is independent of the catalytic site within the DUB, our results show that the active cysteine at position 171 is essential to facilitate the stabilizing effect on the ∆Np63 protein."

sparser
"Alternatively, SCC tumors express high levels of USP28 and the oncogenic transcription factors NOTCH1, c-JUN, c-MYC, and ∆Np63 ( xref B). squamous tumors depend on the USP28-∆Np63 axis to maintain SCC cell identity and induce the formation of tumors in vivo."
| PMC

sparser
"Finally, we targeted the USP28∆Np63 axis via AZ1 in SCC cell lines and compared proliferative responses to a respective ADC line (Figs  xref G and H, and xref D)."
USP28 affects miR-3940-5p
| 7
USP28 activates miR-3940-5p. 7 / 7
| 7

reach
"CCND1 and USP28 Are Targets of miR-3940-5p."

reach
"We searched the TargetScan and miRanda databases and found that CCND1 and USP28 are potential targets of miR-3940-5p."

reach
"It was verified that CCND1 and USP28 were direct targets of miR-3940-5p, and the phenotypes of NSCLC cell proliferation and apoptosis imposed by miR-3940-5p could be attenuated by overexpression of CCND1 or USP28."

reach
"Furthermore, we verified that CCND1 and USP28 were direct targets of miR-3940-5p and also found that the effects of NSCLC cell proliferation and apoptosis by miR-3940-5p were attenuated by overexpression of CCND1 or USP28."

reach
"In combination with a significant negative correlation between expressions of miR-3940-5p versus CCND1 mRNA and USP28 mRNA in NSCLC tissues, these results suggest that CCND1 and USP28 are functional targets of miR-3940-5p and miR-3940-5p affects proliferation and apoptosis in A549 cells by targeting CCND1 and USP28."

reach
"We searched for miR-3940-5p targets by using TargetScan and miRanda online tools and found that CCND1 and USP28 were potential targets of miR-3940-5p."

reach
"However, overexpression of CCND1 and USP28 fails to fully restore the effect of miR-3940-5p on proliferation and apoptosis probably due to a sufficiently high endogenous level of CCND1 and USP28 already existing in A549 cells."
| 7
USP28 activates angiogenesis.
| 6
| 6

reach
"A study has proposed a novel mechanism by which USP28 upregulates angiogenesis by antagonizing GSK-3β (glycogen synthase kinase-3β) and FBW7-dependent degradation of HIF-1α (hypoxia-inducible factor-1α), a major regulator of angiogenesis, carcinogenesis, and various processes by which cells adapt to hypoxic conditions."

reach
"USP28 also upregulates angiogenesis by antagonizing GSK-3β (glycogen synthase kinase-3β) and FBW7-dependent degradation of HIF-1α (hypoxia-inducible factor-1α), a major regulator of angiogenesis, carcinogenesis, and various processes by which the cell adapts to hypoxic conditions."

reach
"USP28 Deficiency Promotes Breast and Liver Carcinogenesis as well as Tumor Angiogenesis in a HIF independent Manner."

reach
"Further, lack of USP28 promotes a more malignant state of breast cancer cells, indicated by an epithelial-to-mesenchymal (EMT) transition, elevated proliferation, migration, and angiogenesis as well as a decreased adhesion."

reach
"Correction : USP28 Deficiency Promotes Breast and Liver Carcinogenesis as well as Tumor Angiogenesis in a HIF independent Manner."

reach
"A study has proposed a novel mechanism by which USP28 upregulates angiogenesis by antagonizing GSK-3beta (glycogen synthase kinase-3beta) and FBW7 dependent degradation of HIF-1alpha (hypoxia inducible factor-1alpha), a major regulator of angiogenesis, carcinogenesis, and various processes by which cells adapt to hypoxic conditions."
USP28 inhibits angiogenesis.
| 1
| 1

reach
"Moreover, GSK3beta and FBW7 dependent HIF-1alpha degradation can be antagonized by the USP28 (ubiquitin specific peptidase-28), suggesting that FBW7 and USP28 could reciprocally regulate cell migration and angiogenesis in a HIF-1alpha-dependent manner [XREF_BIBR]."
USP28 affects SENP1
| 1 6
| 1 6

reach
"The co-immunoprecipitation and GST Pull-down assays were used to determine the interaction between USP28 and SENP1."

sparser
"Firstly, we tested whether SENP1 could interact with USP28."

sparser
"These data indicate that the deSUMOylation enzyme SENP1 directly interacted with USP28."

sparser
"SENP1 directly interacts with USP28."

sparser
"In addition, GST pull-down assay was performed to further study if the interaction between USP28 and SENP1 was direct."

sparser
"The co-immunoprecipitation and GST Pull-down assays were used to determine the interaction between USP28 and SENP1."

sparser
"The explicit interactions between SENP1 and USP28 prompted us to further examine whether SENP1 could be a deSUMOylation enzyme for USP28 SUMOylation."
USP28 affects MAPK
| 7
USP28 inhibits MAPK.
| 5
USP28 inhibits MAPK. 5 / 5
| 5

reach
"XREF_BIBR In a proportion of melanoma patients, USP28 was deleted, and loss of USP28 enhanced MAPK activity through the stabilization of RAF family members, which suggested that USP28 was a key factor in BRAF inhibitor resistance."

reach
"The loss of USP28 promoted MAPK activation and resistance to RAF inhibitor therapy by stabilizing BRAF in cell culture and in vivo models."

reach
"Importantly, of the 59 patients harboring BRAF V600E mutations 27% (16/59) displayed a> 50% decrease in USP28 mRNA expression levels, suggesting that in tumors harboring BRAF alterations, loss of USP28 may further increase the tumorigenic potential of these tumors by stabilizing BRAF and enhancing downstream MAPK activation."

reach
"Our results show that loss of USP28 enhances MAPK activity through the stabilization of RAF family members and is a key factor in BRAF inhibitor resistance."

reach
"Collectively our results show that loss of USP28 enhances downstream MAPK activation through the stabilization of BRAF, leading to decreased sensitivity to combination therapies involving BRAF inhibitors dabrafenib or vemurafenib (XREF_FIG)."
USP28 activates MAPK.
| 2
USP28 activates MAPK. 2 / 2
| 2

reach
"Because USP28 is frequently deleted in melanoma and USP28 depletion leads to BRAF stability and enhanced MAPK kinase in HEK293T cells, we asked if interfering with USP28 expression conferred a similar response in BRAF (V600E) melanoma cell lines."

reach
"In line with previous results, depletion of USP28 in all the melanoma cell lines tested resulted in increased stabilization of BRAF and enhanced downstream MAPK activation (XREF_FIG)."
USP28 affects KLHL2
1 | 2 4
1 | 2 1

No evidence text available

sparser
"Consistent with our data, computational modeling of structures predicted by the ZDOCK and PyMol software showed that Ser145 is the critical amino acid mediating the direct binding of UCK1 to KLHL2 (Figure xref J) while Ser67 phosphorylation disrupts the interaction between USP28 and KLHL2 (Figure xref K)."

reach
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28."

reach
"Consistent with our data, computational modeling of structures predicted by the ZDOCK and PyMol software showed that Ser145 is the critical amino acid mediating the direct binding of UCK1 to KLHL2 while Ser67 phosphorylation disrupts the interaction between USP28 and KLHL2."
USP28 binds UCK1 and KLHL2. 3 / 3
| 3

sparser
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28 (Figure xref G)."

sparser
"USP28 binds to UCK1 via KLHL2."

sparser
"The findings that support this conclusion are the following: (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2-mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5'-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2-UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5'-AZA-induced apoptosis; and (vii) combinational therapies with 5'-AZA and ectopic USP28 expression enhances the effects of 5'-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."
USP28 affects Flag
| 7
| 7

sparser
"293T cells were co-transfected with Flag-USP28, His-SUMO2, GFP-SENP1 WT or SENP1 catalytic mutant (SENP1 CA)."

sparser
"To test this, a Flag-USP28 expression construct was transfected into T24T cells; as expected, USP28 over-expression led to increased CD44 protein expression ( xref )."

sparser
"We found that bacterial purified GST-SENP1 protein was able to capture the Flag-USP28 protein purified from 293T cells."

sparser
"Flag-USP28 WT or Flag-USP28 K99R was co-transfected with His-ubiquitin, FBXW7, and HA-HIF-1α into 293T cells for 36 h, and MG132 was added 4 h before cells were harvested."

sparser
"Flag-USP28 and GFP-SENP1 plasmids were co-transfected into 293T cells for 36 h."

sparser
"To investigate the impact of USP28 SUMOylation on HIF-1α, we generated Flag-USP28 K99R mutant plasmid."

sparser
"Cells were harvested, and the immunoprecipitation of Flag-USP28 revealed the presence of GFP-SENP1 protein (Fig.  xref a), and SENP1 was also present in immunoprecipitates of endogenous USP28 in HCT116 cell lysate (Fig.  xref b)."
PRKD1 affects USP28
3 | 2 2
PRKD1 binds USP28.
3 | 1
3 | 1

sparser
"Consistent with this possibility, we found that USP28 and PRKD1 were stably associated in a co-immunoprecipitation assay ( xref )."

No evidence text available

No evidence text available

No evidence text available
PRKD1 phosphorylates USP28.
| 2 1
PRKD1 phosphorylates USP28. 3 / 3
| 2 1

reach
"We further showed that PRKD1 phosphorylates USP28, which interacts with and stabilizes ZNF304 from proteolytic degradation."

sparser
"We further showed that PRKD1 phosphorylates USP28, which interacts with and stabilizes ZNF304 from proteolytic degradation."

reach
"XREF_FIG shows that PRKD1 could phosphorylate a USP28 peptide containing the second, more conserved predicted phosphorylation site (XREF_FIG)."
KLHL2 affects USP28
1 | 2 4
1 | 2 1

No evidence text available

sparser
"Consistent with our data, computational modeling of structures predicted by the ZDOCK and PyMol software showed that Ser145 is the critical amino acid mediating the direct binding of UCK1 to KLHL2 (Figure xref J) while Ser67 phosphorylation disrupts the interaction between USP28 and KLHL2 (Figure xref K)."

reach
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28."

reach
"Consistent with our data, computational modeling of structures predicted by the ZDOCK and PyMol software showed that Ser145 is the critical amino acid mediating the direct binding of UCK1 to KLHL2 while Ser67 phosphorylation disrupts the interaction between USP28 and KLHL2."
USP28 binds UCK1 and KLHL2. 3 / 3
| 3

sparser
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28 (Figure xref G)."

sparser
"USP28 binds to UCK1 via KLHL2."

sparser
"The findings that support this conclusion are the following: (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2-mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5'-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2-UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5'-AZA-induced apoptosis; and (vii) combinational therapies with 5'-AZA and ectopic USP28 expression enhances the effects of 5'-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."
HIF1A affects USP28
| 2 5
HIF1A desumoylates USP28.
| 2
HIF1A desumoylates USP28. 2 / 2
| 2

sparser
"Hypoxia and HIF-1α promoted USP28 deSUMOylation."

sparser
"Taken together, these data suggested that hypoxia and HIF-1α promoted USP28 deSUMOylation."
HIF1A binds USP28.
| 2
| 1

sparser
"In contrast to VDU2 which directly interacts with HIF-1α, USP28 forms a ternary complex with HIF-1α via its association with HIF-1α bound Fbw7 (Flügel et al., xref )."
| 1

sparser
"To investigate whether HIF-1α interacts with the USP28 promoter, since this factor was required for the loss of USP28 , another ChIP assay using anti-HIF-1α antibody was conducted."
HIF1A activates USP28.
| 1 1
HIF1A activates USP28. 2 / 2
| 1 1

reach
"Here, we report that USP28 is a SUMOylated protein in normoxia with moderate deubiquitinating activity towards HIF-1alpha in vitro, while hypoxia and HIF-1alpha activate USP28 through SENP1 mediated USP28 deSUMOylation to further accumulate HIF-1alpha protein in cells."

sparser
"Here, we report that USP28 is a SUMOylated protein in normoxia with moderate deubiquitinating activity towards HIF-1α in vitro, while hypoxia and HIF-1α activate USP28 through SENP1-mediated USP28 deSUMOylation to further accumulate HIF-1α protein in cells."
HIF1A decreases the amount of USP28.
| 1
HIF1A decreases the amount of USP28. 1 / 1
| 1

reach
"As shown in XREF_FIG, knockdown of both HIF-1alpha and HIF-2alpha attenuated the decrease of USP28 protein level induced by Ni ions or hypoxia."
Flag affects USP28
| 7
| 7

sparser
"293T cells were co-transfected with Flag-USP28, His-SUMO2, GFP-SENP1 WT or SENP1 catalytic mutant (SENP1 CA)."

sparser
"To test this, a Flag-USP28 expression construct was transfected into T24T cells; as expected, USP28 over-expression led to increased CD44 protein expression ( xref )."

sparser
"We found that bacterial purified GST-SENP1 protein was able to capture the Flag-USP28 protein purified from 293T cells."

sparser
"Flag-USP28 WT or Flag-USP28 K99R was co-transfected with His-ubiquitin, FBXW7, and HA-HIF-1α into 293T cells for 36 h, and MG132 was added 4 h before cells were harvested."

sparser
"Flag-USP28 and GFP-SENP1 plasmids were co-transfected into 293T cells for 36 h."

sparser
"To investigate the impact of USP28 SUMOylation on HIF-1α, we generated Flag-USP28 K99R mutant plasmid."

sparser
"Cells were harvested, and the immunoprecipitation of Flag-USP28 revealed the presence of GFP-SENP1 protein (Fig.  xref a), and SENP1 was also present in immunoprecipitates of endogenous USP28 in HCT116 cell lysate (Fig.  xref b)."
ATR affects USP28
| 3 4
ATR phosphorylates USP28.
| 3 3
ATR phosphorylates USP28. 6 / 6
| 3 3

reach
"In response to DNA damage, phosphorylation of the deubiquitinase USP28 by DDR kinases, ATM, and ATR leads to deubiquitination of either GATA4 or its associated factor [XREF_BIBR]."

reach
"ATR phosphorylates USP28 and increases its enzymatic activity."

sparser
"We next asked whether ATM or ATR phosphorylates USP28 upon CPPD treatment."

sparser
"ATR phosphorylates USP28 and increases its enzymatic activity."

sparser
"In response to DNA damage, phosphorylation of the deubiquitinase USP28 by DDR kinases, ATM, and ATR leads to deubiquitination of either GATA4 or its associated factor [ xref ]."

reach
"USP28 is also phosphorylated by ATR and ATM and is involved in the response to DNA damage by virtue of its ability to deubiquitinate and stabilize FBW7 (see below), the ubiquitin ligase involved in the degradation of the myc transcription factor."
ATR binds USP28.
| 1
| 1

sparser
"Notably, binding of p-ATR and ∆Np63 to USP28 was increased after CPPD treatment (Fig.  xref )."
ATG7 affects USP28
| 7
ATG7 increases the amount of USP28.
| 3
ATG7 increases the amount of USP28. 2 / 2
| 2

reach
"ATG7 promoted USP28 protein expression, which directly bound to and deubiquinated CD44s protein and inhibited CD44s protein degradation."

reach
"The results showed that knockdown of ATG7 decreased human usp28 promoter driven reporter transcription activity (XREF_FIG), revealing that ATG7 promotes usp28 promoter transcription."
Modified ATG7 increases the amount of USP28. 1 / 1
| 1

reach
"In this study, we found that ATG7 overexpression increased USP28 protein expression."
ATG7 activates USP28.
| 2
ATG7 activates USP28. 2 / 2
| 2

reach
"The results showed that knockdown of ATG7 decreased human usp28 promoter driven reporter transcription activity (XREF_FIG), revealing that ATG7 promotes usp28 promoter transcription."

reach
"To investigate the mechanisms of ATG7 upregulating USP28 protein, we first examined usp28 mRNA levels in ATG7 knockdown transfectants."
ATG7 demethylates USP28.
| 1
ATG7 leads to the demethylation of USP28. 1 / 1
| 1

reach
"Consistently, TET1 overexpression also abolished ATG7 knockdown caused usp28 promoter hypermethylation (XREF_FIG)."
ATG7 decreases the amount of USP28.
| 1
ATG7 decreases the amount of USP28. 1 / 1
| 1

reach
"Herein, we found that specific knockdown ATG7 resulted in usp28 promoter hypermethylation as comparison to scramble nonsense transfectants in human BC cells, and increased USP28 expression with 5-aza-2-deoxycytidine treatment, strongly indicating that the promoter hypermethylation was responsible for USP28 transcription downregulation due to ATG7 knockdown."
USP28 affects proteolysis
| 6
USP28 activates proteolysis.
| 5
| 4

reach
"Overexpression of USP28 largely reversed HDAC5-KD-induced LSD1 protein degradation, suggesting a role of HDAC5 as a positive regulator of LSD1 through upregulation of USP28 protein."

reach
"Herein, we identified that USP28, a deubiquitinase aberrantly upregulated in patients with ADPKD, selectively removed K48-linked polyubiquitination and reversed protein degradation of signal transducer and activator of transcription 3 (STAT3)."

reach
"Overexpression of USP28 largely reversed HDAC5-knockdown-induced LSD1 protein degradation, suggesting HDAC5 positively regulates LSD1 by stabilizing USP28."

reach
"ATG7 promoted USP28 protein expression, which directly bound to and deubiquinated CD44s protein and inhibited CD44s protein degradation."
Modified USP28 activates proteolysis. 1 / 1
| 1

reach
"Overexpression of USP28 largely reversed HDAC5-KD induced LSD1 protein degradation, suggesting a role of HDAC5 as a positive regulator of LSD1 through upregulation of USP28 protein."
USP28 inhibits proteolysis.
| 1
| 1

reach
"Overexpression of USP28 largely reversed HDAC5-knockdown-induced LSD1 protein degradation, suggesting HDAC5 positively regulates LSD1 by stabilizing USP28."
USP28 affects UBC
6 |
6 |

No evidence text available

No evidence text available

No evidence text available

No evidence text available

No evidence text available

No evidence text available
USP28 affects Neoplasms
| 1 5
USP28 activates Neoplasms.
| 5
| 5

reach
"In SCC tumors, the inhibition of USP28 reduces tumor growth and increases cell death [17]."
| PMC

reach
"The identification of several DUB inhibitors, including inhibitors of USP28, which regulate various oncogenes (e.g., c-Myc) and USP25, which regulates TRAF (tumor necrosis factor receptor-associated factor) signalling, provide alternative approaches to induce degradation of potential “un-druggable” molecules in cancer.Gerry Melino (University of Cambridge, UK) presented data demonstrating the p53 family protein TAp73, facilitates ubiquitin-dependent degradation of HIF1α, thereby suppressing tumor progression."

reach
"Importantly, in mice carrying the tumors, USP28 deletion reduced tumor size and prolonged lifespan18."

reach
"In a mouse model of colorectal cancer, in contrast to the FBW7-deletion phenotype, intestine-specific deletion of USP28 reduced intestinal tumors."

reach
"Moreover, we concentrate on the impacts of USP28 on diverse hallmarks of cancer and discuss whether USP28 can accelerate or inhibit tumor progression."
USP28 inhibits Neoplasms.
| 1
| 1

eidos
"Moreover , we concentrate on the impacts of USP28 on diverse hallmarks of cancer and discuss whether USP28 can accelerate or inhibit tumor progression ."
UBC affects USP28
6 |
6 |

No evidence text available

No evidence text available

No evidence text available

No evidence text available

No evidence text available

No evidence text available
TET1 affects USP28
| 6
TET1 increases the amount of USP28.
| 3
TET1 increases the amount of USP28. 2 / 2
| 2

reach
"Moreover, upregulated TET1 directly demethylates USP28 promoter, thereby enhancing USP28 transcription and expression, which binds to CD44s protein, and remove the ubiquitin group from the ubiquitinated CD44s protein, resulting in stabilization of CD44s protein to mediate stem like property of human BC cells."

reach
"Collectively, our results strongly demonstrate that ATG7 overexpression mediated TET1 upregulation contributes to usp28 promoter demethylation, thereby increasing USP28 expression, consequently resulting in CD44s protein accumulation, and finally enhancing BC stem sphere formation, invasion, and lung metastatic activity."
Modified TET1 increases the amount of USP28. 1 / 1
| 1

reach
"TET1 overexpression markedly promoted USP28 and CD44a protein expression, and rescued cancer stem like properties of T24T (shATG7) cells (XREF_FIG and XREF_FIG)."
TET1 demethylates USP28.
| 2
TET1 demethylates USP28. 2 / 2
| 2

reach
"Collectively, our results strongly demonstrate that ATG7 overexpression mediated TET1 upregulation contributes to usp28 promoter demethylation, thereby increasing USP28 expression, consequently resulting in CD44s protein accumulation, and finally enhancing BC stem sphere formation, invasion, and lung metastatic activity."

reach
"Collectively, our results demonstrate that ATG7 autophagic removal of AUF1 promotes tet1 mRNA stabilization, which demethylates usp28 promoter, and in turn resulting in accumulation of CD44s protein, finally promoting cancer stem like properties, invasion and lung metastatic activity of human high invasive BC cells as shown in XREF_FIG."
TET1 decreases the amount of USP28.
| 1
TET1 decreases the amount of USP28. 1 / 1
| 1

reach
"Moreover, ATG7 inhibition stabilized AUF1 protein and thereby reduced tet1 mRNA stability and expression, which was able to demethylate usp28 promoter, reduced USP28 expression, finally promoting CD44s degradation."
HDAC5 affects USP28
| 5 1
HDAC5 activates USP28.
| 3
HDAC5 activates USP28. 3 / 3
| 3

reach
"We further demonstrated that HDAC5 promoted the protein stability of USP28, a bona fide deubiquitinase of LSD1."

reach
"Alternatively, histone deacetylase 5 (HDAC5) promotes USP28 stability and positively regulates the protein abundance of the Lysine-specific histone demethylase 1A (LSD1) [19]."
| PMC

reach
"Both LSD1 and histone deacetylases (HDACs) facilitate breast cancer proliferation, and interestingly, HDAC5 could promote the stability of USP28."
HDAC5 binds USP28.
| 1 1
| 1 1

reach
"In vitro pull-down assay using His tag recombinant LSD1 protein incubated with USP28-FLAG protein indicated a direct interaction of HDAC5 and USP28 (XREF_SUPPLEMENTARY), and HDAC5 overexpression significantly attenuated USP28 polyubiquitination (XREF_SUPPLEMENTARY)."

sparser
"In vitro pull-down assay using His-tag recombinant LSD1 protein incubated with USP28-FLAG protein indicated a direct interaction of HDAC5 and USP28 ( xref ), and HDAC5 overexpression significantly attenuated USP28 polyubiquitination ( xref )."
HDAC5 deubiquitinates USP28.
| 1
Modified HDAC5 leads to the deubiquitination of USP28. 1 / 1
| 1

reach
"In vitro pull-down assay using His tag recombinant LSD1 protein incubated with USP28-FLAG protein indicated a direct interaction of HDAC5 and USP28 (XREF_SUPPLEMENTARY), and HDAC5 overexpression significantly attenuated USP28 polyubiquitination (XREF_SUPPLEMENTARY)."
CASP8 affects USP28
| 4 2
CASP8 inhibits USP28. 6 / 6
| 4 2

reach
"Furthermore, Caspase-8 cleaves and inactivates USP28 to overcome the p53-dependent G2/M checkpoint in cancer cells [18]."
| PMC

reach
"In the nucleus, caspase-8 cleaves and inactivates the ubiquitin specific peptidase 28 (USP28), preventing USP28 from de-ubiquitinating and stabilizing wildtype p53."

reach
"In the nucleus, caspase-8 cleaves and inactivates the ubiquitin specific peptidase 28 (USP28), preventing USP28 from de-ubiquitinating and stabilizing wildtype p53."
| PMC

sparser
"We further provide evidence that tumor cells expressing high and nuclear levels of caspase-8 have defective p53-dependent apoptosis because caspase-8 cleaves and inactivates USP28 in cells with delayed or compromised mitosis ( xref )."

sparser
"Furthermore, Caspase-8 cleaves and inactivates USP28 to overcome the p53-dependent G2/M checkpoint in cancer cells [ xref ]."
| PMC

reach
"Thus, nuclear caspase-8 cleaves and thereby inactivates ubiquitin-specific peptidase 28 (USP28), the protein which stabilizes the wild-type p53 protein."
Nickel(2+) affects USP28
| 5
Nickel(2+) decreases the amount of USP28.
| 4
Nickel(2+) decreases the amount of USP28. 4 / 4
| 4

reach
"Although further studies did not show that hypoxia signaling impaired the association of USP28 with Fbw7alpha and c-Myc, cellular USP28 protein levels were decreased by Ni ions and hypoxia in A549 cells."

reach
"However, USP28 protein expression was decreased by Ni ions and hypoxia in the A549 cells (XREF_FIG)."

reach
"Our study demonstrated that Nickel and hypoxia exposure increased c-myc T58 phosphorylation and decreased USP28 protein levels in cancer cells, which both lead to enhanced c-myc ubiquitination and proteasomal degradation."

reach
"Here, it has been demonstrated that Ni ions and hypoxia increased H3K9m2 levels at the USP28 promoter and repressed USP28 gene expression."
Nickel(2+) increases the amount of USP28.
| 1
Nickel(2+) increases the amount of USP28. 1 / 1
| 1

reach
"As shown in XREF_FIG, knockdown of both HIF-1alpha and HIF-2alpha attenuated the decrease of USP28 protein level induced by Ni ions or hypoxia."
MiR-4295 affects USP28
| 5
MiR-4295 activates USP28.
| 4
MiR-4295 activates USP28. 4 / 4
| 4

reach
"Accumulating evidence indicated that miR-4295 may target USP28 in non small lung cancer and high expression of USP28 promoted tumour cells proliferation 22."

reach
"Thus, our data indicated that USP28 was targeted by miR-4295."

reach
"Our data indicated that miR-4295 may have targeted USP28 in NSCLC."

reach
"USP28 was targeted by miR-4295."
MiR-4295 decreases the amount of USP28.
| 1
Modified miR-4295 decreases the amount of USP28. 1 / 1
| 1

reach
"Nevertheless, the overexpression of miR-4295 inhibits the expression of USP28, which is an oncogene in lung cancer, indicating a tumor suppressive role of miR-4295 [35]."
USP28 affects ZNF304
2 1 | 1 1
USP28 binds ZNF304.
2 | 1
2 | 1

No evidence text available

sparser
"The co-immunoprecipitation experiment of xref shows that USP28 and ZNF304 were physically associated."

No evidence text available
USP28 deubiquitinates ZNF304.
1 | 1
USP28 leads to the deubiquitination of ZNF304. 2 / 2
1 | 1

No evidence text available

reach
"Notably, unlike wild-type USP28, a USP28 derivative containing a mutation in the PRKD1 phosphorylation site, USP28 (S899A), was unable to reduce ZNF304 ubiquitination (XREF_FIG)."
USP25 affects USP28
2 | 2 1
USP25 binds USP28.
2 | 1
2 | 1

No evidence text available

sparser
"Furthermore, cellular assays were conducted to confirm that Vismodegib could interact with the evolutionarily related deubiquitinases USP28 and USP25 and down-regulate the levels of the two enzymes' substrate proteins c-Myc, Notch1 and Tankyrase-1/2."

No evidence text available
USP25 dephosphorylates USP28.
| 2
USP25 leads to the dephosphorylation of USP28 on serine. 1 / 1
| 1

reach
"The USP25 activity may not be affected upon exposure to an inhibitor blocking the phosphorylation of USP28 on serine 67 and serine It could be another therapeutic option to target the binding between ATM and USP28 and reduce the phosphorylation of USP28, thus decreasing its enzymatic activity in cancer cells with increased DNA damage levels."
| PMC
USP25 leads to the dephosphorylation of USP28 at position 67. 1 / 1
| 1

reach
"The USP25 activity may not be affected upon exposure to an inhibitor blocking the phosphorylation of USP28 on serine 67 and serine It could be another therapeutic option to target the binding between ATM and USP28 and reduce the phosphorylation of USP28, thus decreasing its enzymatic activity in cancer cells with increased DNA damage levels."
| PMC
SGH affects USP28
| 1 4
SGH inhibits USP28.
| 1 2
SGH inhibits USP28. 3 / 3
| 1 2

reach
"All data together indicated that the downregulation of c-Myc, HK2, PKM2, LDHA, PHGDH, PSAT1, cyclin D1, and CDK6 in lung cancer cells was related to the ubiquitin protein degradation pathway, which was the result that the USP28 was inhibited by SGH."

eidos
"All data together indicated that the downregulation of c-Myc , HK2 , PKM2 , LDHA , PHGDH , PSAT1 , cyclin D1 , and CDK6 in lung cancer cells was related to the ubiquitin-protein degradation pathway , which was the result that the USP28 was inhibited by SGH ."

reach
"In terms of mechanism of action, SGH downregulated cell cycle- and nucleotide synthesis related proteins to block cell cycle at G0/G1 phase, reduced the expression levels of glycolytic metabolic enzymes to inhibit glycolysis, and downregulated the important cancer transcription factor c-Myc and the therapeutic target deubiquitinase USP28."
SGH decreases the amount of USP28.
| 2
SGH decreases the amount of USP28. 2 / 2
| 2

reach
"A 24 h treatment of SGH at 2 microM for PC9 cells and 5 microM for H157 cells downregulated USP28 and c-Myc expressions (** p < 0.01 or *** p < 0.001, XREF_FIG A, B and Figure S7A, B)."

reach
"Interestingly, SGH also downregulated the levels of USP28 and c-Myc."
RCHY1 affects CHEK2
| 5
| 5

sparser
"Similar to FBXW7, USP28 forms a complex with PIRH2 and CHK2, thus antagonizing the ubiquitination of CHK2 by PIRH2."
| PMC

sparser
"We show that the deubiquitylating enzyme USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK221."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and antagonizes PIRH2-mediated polyubiquitylation and proteasomal degradation of CHK2 [ xref ]."

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 xref ."
Vismodegib affects USP28
| 4
Vismodegib binds USP28.
| 3

reach
"Multiple biophysical and biochemical techniques including NMR, ITC, thermal shift assay, HDX-MS and site directed mutagenesis analysis were then used to characterize the interaction between Vismodegib and USP28."

reach
"Furthermore, cellular assays were conducted to confirm that Vismodegib could interact with the evolutionarily related deubiquitinases USP28 and USP25 and down-regulate the levels of the two enzymes ' substrate proteins c-Myc, Notch1 and Tankyrase-1/2."

reach
"Herein, we report our efforts to solve the cocrystal structure of Vismodegib bound to USP28 for the first time and subsequent structure-based optimization leading to a series of Vismodegib derivatives as potent USP28 inhibitors."
Vismodegib inhibits USP28.
| 1
| 1

reach
"USP28 and USP25 are down-regulated by Vismodegib in vitro and in colorectal cancer cell lines."

reach
"Functional assays demonstrated that overexpression of USP28 promoted cell proliferation and aerobic glycolysis of colorectal cancer, while USP28 inhibition could reverse these effects."

reach
"USP28 increases aerobic glycolysis and promotes cell proliferation upon stabilization of FOXC1 [50]."
| PMC

reach
"USP28 promotes aerobic glycolysis of colorectal cancer by increasing stability of FOXC1."

reach
"In conclusion, USP28 enhanced cell viability and aerobic glycolysis of colorectal cancer by stabilizing FOXC1, suggesting that USP28-FOXC1 might be a novel therapeutic avenue for colorectal cancer."

reach
"Here, we report that USP28 promotes the activity of the Wnt signaling pathway through maintaining the stability of TCF7L2."

eidos
"Here , we report that USP28 promotes the activity of the Wnt signaling pathway through maintaining the stability of TCF7L2 ."

eidos
"we reported that USP28 promotes the activity of the Wnt signaling pathway through maintaining the stability of TCF7L2 ."

eidos
"Our results show that USP28 contributes to the activity of the Wnt signaling pathway through stabilizing TCF7L2 ( Figure 7 ) , at least partially , as it also regulates other members of the TCF / LEF family transcription factors ( Figure 2A ) as well as Forkhead box protein M1 ( FOXM1 ) , another transcription factor involved in Wnt signaling ."
USP28 affects TCF7L2
| 1 3
USP28 activates TCF7L2.
| 1 1
USP28 activates TCF7L2. 2 / 2
| 1 1

reach
"Here, we report that USP28 promotes the activity of the Wnt signaling pathway through maintaining the stability of TCF7L2."

eidos
"As shown in Figure S3D , the absence of Usp28 resulted in the decreases of TCF7L2 levels as well as cyclin D1 levels ."
USP28 increases the amount of TCF7L2.
| 1
USP28 increases the amount of TCF7L2. 1 / 1
| 1

reach
"By regulating the levels of TCF7L2, USP28 modulates the Wnt/β-catenin signaling in liver cancer and USP28 depletion or inhibition by a small molecule inhibitor leads to a halt of growth in liver cancer cells."
USP28 deubiquitinates TCF7L2.
| 1
USP28 deubiquitinates TCF7L2. 1 / 1
| 1

reach
"USP28 Deubiquitinates TCF7L2 to Govern the Action of Wnt Signaling Pathway in Hepatic Carcinoma."
USP28 affects PRKD1
3 | 1
3 | 1

sparser
"Consistent with this possibility, we found that USP28 and PRKD1 were stably associated in a co-immunoprecipitation assay ( xref )."

No evidence text available

No evidence text available

No evidence text available
USP28 affects MDC1
1 1 | 1 1
1 1 | 1

No evidence text available

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."

No evidence text available
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
USP28 affects FOXC1
| 3 1
USP28 activates FOXC1.
| 3
USP28 activates FOXC1. 3 / 3
| 3

reach
"In conclusion, USP28 enhanced cell viability and aerobic glycolysis of colorectal cancer by stabilizing FOXC1, suggesting that USP28-FOXC1 might be a novel therapeutic avenue for colorectal cancer."

reach
"Moreover, proteasome inhibitor, MG132, could rescue USP28 silence induced degradation of FOXC1."

reach
"Protein expression of Forkhead Box C1 (FOXC1) was increased by USP28 over-expression, whereas knockdown of USP28 aggravated cycloheximide (CHX; protein synthesis inhibitor) stimulated decrease of FOXC1."
USP28 binds FOXC1.
| 1
USP28 binds FOXC1. 1 / 1
| 1

sparser
"In conclusion, USP28 enhanced cell viability and aerobic glycolysis of colorectal cancer by stabilizing FOXC1, suggesting that USP28-FOXC1 might be a novel therapeutic avenue for colorectal cancer."
USP28 affects ERK
| 3 1
USP28 inhibits ERK.
| 1
USP28 inhibits ERK. 1 / 1
| 1

reach
"Collectively these results suggest that USP28 expression is regulated by MAPK activity and may function through a feedback loop to negatively regulate ERK signaling."
USP28 increases the amount of ERK.
| 1
USP28 increases the amount of phosphorylated ERK. 1 / 1
| 1

reach
"As expected USP28 mediated BRAF stability led to enhanced pERK levels after vemurfenib treatment compared with wild-type USP28 cells."
USP28 decreases the amount of ERK.
| 1
Modified USP28 decreases the amount of phosphorylated ERK. 1 / 1
| 1

reach
"In contrast, ectopic expression of USP28, but not a catalytically inactive mutant, repressed phospho-ERK levels (XREF_FIG)."
USP28 binds ERK.
| 1
| 1

sparser
"Consistent with this finding, vemurafenib resistance in USP28 knockdown cell lines was associated with sustained ERK phosphorylation ( xref ; and Fig. S5, C and D)."
| 2 2
USP28 inhibits E3_Ub_ligase.
| 2
| 2

reach
"USP28 was shown to bind c-Myc through an interaction with FBW7alpha and antagonize its E3 ligase activities in the nucleus, leading to Myc stabilization and tumor cell proliferation."

reach
"USP28 antagonizes FBW7, an F-box protein and an important component of the SKP1-CUL1-F-BOX (SCF)-type E3 ubiquitin ligase that targets key transcriptional factors to ubiquitin-directed proteasome degradation."
| 2

sparser
"A deubiquitinase USP28 binds to the SCF Fbw7 ubiquitin E3 ligase, stabilizing Myc, and therefore promoting cell proliferation [ xref ]."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
USP28 affects CDKN1A
| 2 2
USP28 binds CDKN1A.
| 2
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."
USP28 activates CDKN1A.
| 2
Mutated USP28 activates CDKN1A. 1 / 1
| 1

reach
"Consistent with this, the expression of WT or ubiquitin binding mutant USP28 rescued p21 and MDM2 induction defects in USP28Delta cells, whereas expression of USP28 C171A did not (XREF_SUPPLEMENTARY B)."
USP28 activates CDKN1A. 1 / 1
| 1

reach
"We showed that 53BP1 and USP28 are required to trigger p53 and p21- dependent cell cycle arrest, evoking an irreversible stress response that selects against unfit cells with disturbed mitosis (XREF_FIG)."
MDC1 affects USP28
1 1 | 1 1
1 1 | 1

No evidence text available

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."

No evidence text available
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
CLSPN affects USP28
1 1 | 1 1
1 1 | 1

No evidence text available

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."

No evidence text available
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
3 |
Trichostatin A increases the amount of USP28. 3 / 3
3 |

No evidence text available

No evidence text available

No evidence text available
Lipopolysaccharide, E coli O55-B5 increases the amount of USP28.
2 |
2 |

No evidence text available

No evidence text available
Lipopolysaccharide, E coli O55-B5 decreases the amount of USP28.
1 |
1 |

No evidence text available
ZNF304 affects USP28
2 | 1
2 | 1

No evidence text available

sparser
"The co-immunoprecipitation experiment of xref shows that USP28 and ZNF304 were physically associated."

No evidence text available
USP28 affects vismodegib
| 3

reach
"Multiple biophysical and biochemical techniques including NMR, ITC, thermal shift assay, HDX-MS and site directed mutagenesis analysis were then used to characterize the interaction between Vismodegib and USP28."

reach
"Furthermore, cellular assays were conducted to confirm that Vismodegib could interact with the evolutionarily related deubiquitinases USP28 and USP25 and down-regulate the levels of the two enzymes ' substrate proteins c-Myc, Notch1 and Tankyrase-1/2."

reach
"Herein, we report our efforts to solve the cocrystal structure of Vismodegib bound to USP28 for the first time and subsequent structure-based optimization leading to a series of Vismodegib derivatives as potent USP28 inhibitors."

reach
"Usp28 deficiency promoted tumor cell differentiation accompanied by decreased proliferation, which suggests that USP28 acts similarly in intestinal homeostasis and colorectal cancer models."

reach
"Knockdown of USP28 directly increases the expression of differentiation genes but indirectly suppresses the expression of pluripotent molecules."

reach
"Knockdown of USP28 induces differentiation and suppresses self-renewal in CSCs derived from MMTV-Wnt1 mice."
USP28 affects USP25
2 | 1
2 | 1

No evidence text available

sparser
"Furthermore, cellular assays were conducted to confirm that Vismodegib could interact with the evolutionarily related deubiquitinases USP28 and USP25 and down-regulate the levels of the two enzymes' substrate proteins c-Myc, Notch1 and Tankyrase-1/2."

No evidence text available
USP28 affects SUMO2
3 |
3 |

No evidence text available

No evidence text available

No evidence text available
USP28 affects Notch
| 3
USP28 activates Notch.
| 2
USP28 activates Notch. 2 / 2
| 2

reach
"USP28 has also been reported to antagonize ubiquitin-dependent degradation of the oncogene product MYC as well as JUN and Notch ."

reach
"USP28 has also been reported to antagonize ubiquitin-dependent degradation of the oncogene product MYC as well as JUN and Notch 105 ."
USP28 increases the amount of Notch.
| 1
Modified USP28 increases the amount of Notch. 1 / 1
| 1

reach
"Intriguingly, both ectopic expression of Usp28 in wild-type MEFs (+/+ +) and complete knockout of Usp28 (-/-) strongly and equivalently increased levels of Myc, Jun, and Notch."
| 2

reach
"Knockdown of USP28 resulted in LSD1 destabilization, leading to the suppression of cancer stem cell (CSC)-like characteristics invitro and inhibition of tumorigenicity invivo, which can be rescued by ectopic LSD1 expression."

reach
"Although our results indicate that LSD1 is a major target of USP28 in breast cancer, it does not rule out the possibility that USP28 promotes CSC like traits through both LSD1 stabilization and inhibition of Myc degradation."

reach
"Knockdown of USP28 inhibits CSC like traits in vitro and suppresses tumorigenicity in vivo."
USP28 affects JUN
| 2 1
USP28 increases the amount of JUN.
| 1
Modified USP28 increases the amount of JUN. 1 / 1
| 1

reach
"Intriguingly, both ectopic expression of Usp28 in wild-type MEFs (+/+ +) and complete knockout of Usp28 (-/-) strongly and equivalently increased levels of Myc, Jun, and Notch."
USP28 binds JUN.
| 1
| 1

sparser
"The ChIP results of xref show, consistent with the bioinformatic prediction, that in DLD-1 cells cJUN was bound to the USP28 promoter."
USP28 activates JUN.
| 1
USP28 activates JUN. 1 / 1
| 1

reach
"Interestingly, USP28 can also antagonize the ubiquitin-dependent degradation of two additional oncogenic proteins, c-Jun and Notch1, expanding the substrate list that is shared by both USP28 and FBW7."
USP28 affects GSK3B
| 1 2
USP28 inhibits GSK3B.
| 1 1
USP28 inhibits GSK3B. 2 / 2
| 1 1

reach
"Moreover, GSK3beta and FBW7 dependent HIF-1alpha degradation can be antagonized by the USP28 (ubiquitin specific peptidase-28), suggesting that FBW7 and USP28 could reciprocally regulate cell migration and angiogenesis in a HIF-1alpha-dependent manner [XREF_BIBR]."

eidos
"APC , adenomatous polyposis coli ; DVL , Disheveled ; Fz , frizzled ; GSK3 , glycogen synthase kinase-3 ; LRP , lipoprotein receptor-related ; USP28 , ubiquitin-specific protease 28 As Usp28-deficient mice develop normally , the role of Usp28 in the normal Wnt signaling during embryonic development must be minimal or its absence could be mitigated by unspecified redundant mechanisms , for example , suppressing GSK3beta activity so that TCF7L2 is not ubiquitinated in the first place ."
USP28 activates GSK3B.
| 1
USP28 activates GSK3B. 1 / 1
| 1

reach
"A study has proposed a novel mechanism by which USP28 upregulates angiogenesis by antagonizing GSK-3beta (glycogen synthase kinase-3beta) and FBW7 dependent degradation of HIF-1alpha (hypoxia inducible factor-1alpha), a major regulator of angiogenesis, carcinogenesis, and various processes by which cells adapt to hypoxic conditions."
USP28 affects FBXW7-185aa
| 3
USP28 binds FBXW7-185aa. 3 / 3
| 3

reach
"Instead, FBXW7-185aa binds to USP28, which is a de-ubiquitinating enzyme that stabilizes c-Myc by inhibiting FBXW7alpha activity."

reach
"The above evidence collectively elucidates that FBXW7-185aa competitively interacted with USP28 and " freed " FBXW7alpha to degrade c-Myc."

reach
"XREF_BIBR Mechanistic studies demonstrated that FBXW7-185aa competitively interacted with the deubiquitinating enzyme USP28, preventing USP28 from binding to FBXW7, thereby upregulating FBXW7 expression."
USP28 affects F-box
| 3
USP28 activates F-box.
| 2
USP28 activates F-box. 2 / 2
| 2

reach
"Furthermore, USP28 is mutated in human cancer cells and is reported to antagonize the tumour suppressor F-box and WD40 domain-containing protein 7 (FBW7) 31 , highlighting the potential for USP28 inhibitors in various tumours, especially colorectal cancer 104 ."

reach
"Furthermore, USP28 is mutated in human cancer cells and is reported to antagonize the tumour suppressor F-box and WD40 domain-containing protein 7 (FBW7) , highlighting the potential for USP28 inhibitors in various tumours, especially colorectal cancer ."
USP28 inhibits F-box.
| 1
USP28 inhibits F-box. 1 / 1
| 1

reach
"USP28 antagonizes FBW7, an F-box protein and an important component of the SKP1-CUL1-F-BOX (SCF)-type E3 ubiquitin ligase that targets key transcriptional factors to ubiquitin-directed proteasome degradation."
USP28 affects CD44
| 1 2
| 1 2

sparser
"Binding of USP28 to CD44 standard (CD44s) protein leads to removal of the ubiquitin group from the ubiquitinated CD44s protein, resulting in the stabilization of CD44s protein to mediate the stem-like property of human bladder cancer cells."

reach
"Binding of USP28 to CD44 standard (CD44s) protein leads to removal of the ubiquitin group from the ubiquitinated CD44s protein, resulting in the stabilization of CD44s protein to mediate the stem-like property of human bladder cancer cells."

sparser
"Consistently, our previous study found that ATG7 regulates CD44 protein expression though USP28 binding to CD44 and promotes the debiquitination of CD44, consequently, increasing its protein stabilization [ xref ]."
USP28 affects ABCC9
| 3
| 2

sparser
"The full length compared with the truncated forms of the mef2c, lrrfip1, usp28 and abcc9 genes was decreased in the hnRNP A1 heterozygous mice compared with the hnRNP A1 +/+ mice ( xref b )."

sparser
"Previous studies demonstrated that there were different splicing forms of the mef2c, lrrfip1, usp28 and abcc9 genes during muscle development [ xref – xref ]."
| 1

sparser
"The alternative splicing forms of usp28 and abcc9 changed in rbm24 defect mice, causing dilated atria [ xref ]."
UCK1 affects KLHL2
| 3
USP28 binds UCK1 and KLHL2. 3 / 3
| 3

sparser
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28 (Figure xref G)."

sparser
"USP28 binds to UCK1 via KLHL2."

sparser
"The findings that support this conclusion are the following: (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2-mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5'-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2-UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5'-AZA-induced apoptosis; and (vii) combinational therapies with 5'-AZA and ectopic USP28 expression enhances the effects of 5'-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."
SUMO2 affects USP28
3 |
3 |

No evidence text available

No evidence text available

No evidence text available
KLHL2 affects UCK1, and USP28
| 3
USP28 binds UCK1 and KLHL2. 3 / 3
| 3

sparser
"IP and IB analyses suggested interactions between KLHL2, UCK1, and USP28 (Figure xref G)."

sparser
"USP28 binds to UCK1 via KLHL2."

sparser
"The findings that support this conclusion are the following: (i) the binding of UCK1 to the E3 ligase KLHL2 causes its polyubiquitination at K81 and degradation; (ii) the DUB USP28 binds to UCK1 through interaction with KLHL2 and antagonizes KLHL2-mediated effect on UCK1; (iii) UCK1 deubiquitination by USP28 is dependent on the phosphorylation status of USP28; (iv) USP28 is phosphorylated by 5'-AZA-activated ATM, resulting in disassociation of KLHL2 from USP28 and UCK1 destabilization; (v) ATM also phosphorylates UCK1 at S145, significantly enhancing the KLHL2-UCK1 complex formation; (vi) the knockdown of KLHL2 not only significantly inhibits AML cell proliferation, but also sensitizes AML cells to 5'-AZA-induced apoptosis; and (vii) combinational therapies with 5'-AZA and ectopic USP28 expression enhances the effects of 5'-AZA and induces synergistic lethality at cellular level and in the AML mouse model, which is, however, observed in cells deficient in USP28."
Hypoxia affects USP28
| 3
Hypoxia activates USP28.
| 2
Hypoxia activates USP28. 2 / 2
| 2

eidos
"Results Here , we report that USP28 is a SUMOylated protein in normoxia with moderate deubiquitinating activity towards HIF-1alpha in vitro , while hypoxia and HIF-1alpha activate USP28 through SENP1-mediated USP28 deSUMOylation to further accumulate HIF-1alpha protein in cells ."

eidos
"Our data display that hypoxia modulates the activity of USP28 , which leads to the question whether hypoxia also regulates USP28 SUMOylation ."
Hypoxia inhibits USP28.
| 1
| 1

eidos
"Results Hypoxia enhanced the deubiquitinating activity of USP28 on HIF-1alpha It has been reported that USP28 was down-regulated by hypoxia at both mRNA and protein levels [ 4 ] ."
HDAC affects USP28
| 3
HDAC increases the amount of USP28.
| 2
HDAC increases the amount of USP28. 2 / 2
| 2

reach
"Additional mechanism suggests that histone deacetylases 5 (HDAC5) promotes stability of USP28, thus enhancing stability of LSD1.Overexpression of USP28, HDAC5, and LSD1 is positively correlated in breast tumors."

reach
"16 Similarly, decreased HDAC activity promotes the acetylation of Htt protein and also causes an increase in Ube2e3, SUMo2, and USp28 expression."
HDAC decreases the amount of USP28.
| 1
HDAC decreases the amount of USP28. 1 / 1
| 1

reach
"16 Similarly, decreased HDAC activity promotes the acetylation of Htt protein and also causes an increase in Ube2e3, SUMo2, and USp28 expression."
FBXW7-185aa affects USP28
| 3
USP28 binds FBXW7-185aa. 3 / 3
| 3

reach
"Instead, FBXW7-185aa binds to USP28, which is a de-ubiquitinating enzyme that stabilizes c-Myc by inhibiting FBXW7alpha activity."

reach
"The above evidence collectively elucidates that FBXW7-185aa competitively interacted with USP28 and " freed " FBXW7alpha to degrade c-Myc."

reach
"XREF_BIBR Mechanistic studies demonstrated that FBXW7-185aa competitively interacted with the deubiquitinating enzyme USP28, preventing USP28 from binding to FBXW7, thereby upregulating FBXW7 expression."
CD44 affects USP28
| 1 2
| 1 2

sparser
"Binding of USP28 to CD44 standard (CD44s) protein leads to removal of the ubiquitin group from the ubiquitinated CD44s protein, resulting in the stabilization of CD44s protein to mediate the stem-like property of human bladder cancer cells."

reach
"Binding of USP28 to CD44 standard (CD44s) protein leads to removal of the ubiquitin group from the ubiquitinated CD44s protein, resulting in the stabilization of CD44s protein to mediate the stem-like property of human bladder cancer cells."

sparser
"Consistently, our previous study found that ATG7 regulates CD44 protein expression though USP28 binding to CD44 and promotes the debiquitination of CD44, consequently, increasing its protein stabilization [ xref ]."
ABCC9 affects USP28
| 3
| 2

sparser
"The full length compared with the truncated forms of the mef2c, lrrfip1, usp28 and abcc9 genes was decreased in the hnRNP A1 heterozygous mice compared with the hnRNP A1 +/+ mice ( xref b )."

sparser
"Previous studies demonstrated that there were different splicing forms of the mef2c, lrrfip1, usp28 and abcc9 genes during muscle development [ xref – xref ]."
| 1

sparser
"The alternative splicing forms of usp28 and abcc9 changed in rbm24 defect mice, causing dilated atria [ xref ]."
Vemurafenib affects USP28
| 2
| 2

reach
"It is important to note that in both cell lines tested vemurafenib treatment enhanced the expression USP28, indicative of a USP28 feedback loop in both these BRAF mutant cell lines (XREF_FIG)."

reach
"Pharmacodynamics studies demonstrated that depletion of USP28 led to a robust retention of ERK phosphorylation in tumors treated with vemurafenib (XREF_FIG)."
Phenylmercury acetate increases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
2 |
Panobinostat increases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
P-chloromercuribenzoic acid increases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
MiR-500a-5p affects USP28
| 1 1
MiR-500a-5p inhibits USP28.
| 1
MiR-500a-5p inhibits USP28. 1 / 1
| 1

eidos
"Collectively , our data suggested that high levels of miR-500a-5p in CAFs are transferred into tumor cells and downregulates USP28 expression , which promotes cell proliferation , metastasis and EMT in breast cancer ( Figure 8) ."
MiR-500a-5p activates USP28.
| 1
MiR-500a-5p activates USP28. 1 / 1
| 1

reach
"Therefore, these data indicated that miR-500a-5p promotes breast cancer progression and metastasis by sponging USP28."
MiR-216b affects USP28
| 2
MiR-216b inhibits USP28.
| 1
MiR-216b inhibits USP28. 1 / 1
| 1

reach
"These results indicated that miR-216b downregulated USP28 and c-Myc signaling in HCC cells."
MiR-216b activates USP28.
| 1
MiR-216b activates USP28. 1 / 1
| 1

reach
"Further studies showed that overexpression of miR-216b inhibited HCC cell growth, and molecular investigations revealed that miR-216b targeted USP28 and inhibited its expression in HCC cells."
2 |
Mercury dibromide increases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
Lactate affects USP28
| 2
Lactate activates USP28. 2 / 2
| 2

reach
"LDHA generating lactate enhances the USP28 signaling."

reach
"Interestingly, the lactate induced weak acidic environment promoted USP28 signaling as well as hydrochloric acid and acetic acid (XREF_FIG)."
2 |
Hsa-miR-363-3p decreases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
Entinostat affects USP28
2 |
Entinostat increases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
Dioxygen affects USP28
1 | 1
Dioxygen inhibits USP28.
| 1
| 1

reach
"Reactive oxygen species dependent down-regulation of tumor suppressor genes PTEN, USP28, DRAM, TIGAR, and CYLD under oxidative stress."
Dioxygen decreases the amount of USP28.
1 |
Dioxygen decreases the amount of USP28. 1 / 1
1 |

No evidence text available
Bisphenol A affects USP28
2 |
Bisphenol A decreases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
Belinostat affects USP28
2 |
Belinostat increases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
USP28 affects vemurafenib
| 2
| 2

reach
"As USP28 knockdown inhibited the ability of vemurafenib to attenuate ERK phosphorylation, we reasoned that in these cell lines BIM (Bcl-2-interacting mediator of cell death) accumulation would be down-regulated."

reach
"USP28 mediates vemurafenib sensitivity in vivo."
USP28 affects ubiquitination
| 2
USP28 inhibits ubiquitination. 2 / 2
| 2

sparser
"Overexpression of USP28 inhibited FBW7-depedent ubiquitination of c-MYC and cyclin E1, both of which are oncoproteins."

sparser
"USP7 and USP28 inhibited LSD1 ubiquitination and stabilized LSD1 protein level [ xref , xref ]."
USP28 affects tumor growth
| 2
USP28 activates tumor growth. 2 / 2
| 2

eidos
"Moreover , the DUB USP7 , USP13 , USP22 , USP28 , USP36 and USP37 stabilize c-Myc , thereby stimulating tumor growth [ 155 , 157 , 171-174 ] ."
| PMC

eidos
"In SCC tumors , the inhibition of USP28 reduces tumor growth and increases cell death [ 17 ] ."
| PMC
USP28 affects signaling
| 2
USP28 activates signaling. 2 / 2
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53–p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."
USP28 affects miR-500a-5p
| 2
USP28 inhibits miR-500a-5p.
| 1
USP28 inhibits miR-500a-5p. 1 / 1
| 1

reach
"In accordance with these findings, our study revealed that USP28, as the target gene of miR-500a-5p, reversed the ability of miR-500a-5p to promoted breast cancer cell proliferation and metastasis, indicating that USP28 functions as a breast cancer suppressor."
USP28 activates miR-500a-5p.
| 1
USP28 activates miR-500a-5p. 1 / 1
| 1

reach
"These findings suggested that MUT1 contained the affirmative mutant binding sites in the 3 '-UTR of USP28, thus indicating that USP28 is a direct target of miR-500a-5p."
| 2

reach
"Moreover, GSK3beta and FBW7 dependent HIF-1alpha degradation can be antagonized by the USP28 (ubiquitin specific peptidase-28), suggesting that FBW7 and USP28 could reciprocally regulate cell migration and angiogenesis in a HIF-1alpha-dependent manner [XREF_BIBR]."

reach
"Transwell assays showed that USP28 overexpression reduced both cell migration and invasion in MDA-MB-231 and MCF7 cells, and these effects were partially reversed by transfection with miR-500a-5p mimics."
USP28 affects cell death
| 1 1
| 1 1

reach
"In SCC tumors, the inhibition of USP28 reduces tumor growth and increases cell death [17]."
| PMC

eidos
"In SCC tumors , the inhibition of USP28 reduces tumor growth and increases cell death [ 17 ] ."
| PMC
| 2

reach
"In contrast, in a detailed study on the function of USP28 in breast cancer XREF_BIBR, Richter et al. found that USP28 deficiency in breast cancer cells enhances conversion toward a more aggressive phenotype by promoting EMT, proliferation, migration, angiogenesis, and decreased adhesion."

reach
"Further, lack of USP28 promotes a more malignant state of breast cancer cells, indicated by an epithelial-to-mesenchymal (EMT) transition, elevated proliferation, migration, and angiogenesis as well as a decreased adhesion."
USP28 affects Wnt signaling
| 2
USP28 activates Wnt signaling. 2 / 2
| 2

eidos
"To explore how USP28 might be involved in HCC development , we undertook a Gene Set Enrichment Analysis and found a significant positive correlation between USP28 expression and the expression of Wnt signaling pathway genes ( Figure 1D ) , suggesting that USP28 might contribute to Wnt signaling ."

eidos
"Taken together , these data strongly suggest that USP28 contributes to Wnt signaling through maintaining the stability of TCF / LEF transcription factors ."
USP28 affects USP21
2 |
2 |

No evidence text available

No evidence text available
USP28 affects TET1
| 2
USP28 activates TET1. 1 / 1
| 1

reach
"Upregulated TET1 then directly demethylates the ubiquitin specific peptidase 28 (USP28) promoter, thereby enhancing USP28 transcription and expression."
Unmethylated USP28 activates TET1. 1 / 1
| 1

reach
"Upregulated TET1 then directly demethylates the ubiquitin specific peptidase 28 (USP28) promoter, thereby enhancing USP28 transcription and expression."
USP28 affects SNAI2
| 2
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."

reach
"Further, we found that USP28 promoted PC cell growth by facilitating cell cycle progression and inhibiting apoptosis."

reach
"Overexpression of USP28 accelerated PC cell growth, whereas USP28 knockdown impaired PC cell growth both in vitro and in vivo."
USP28 affects PLK1
| 2
USP28 activates PLK1. 2 / 2
| 2

reach
"Additionally, in this system, USP28 only targets Claspin but not PLK1 (both of which are subject to ubiquitination by the same E3 ligase)."

reach
"The specificity of DUBs, however, is poorly understood, and it is currently unknown how Usp28 targets Claspin but not Plk1."

reach
"USP28 contributes to the proliferation and metastasis of gastric cancer."

reach
"MiR-500a-5p was transferred from CAFs to the cancer cells, and subsequently promoted proliferation and metastasis by binding to ubiquitin specific peptidase 28 (USP28)."
USP28 affects MEF2C
| 2
| 2

sparser
"The full length compared with the truncated forms of the mef2c, lrrfip1, usp28 and abcc9 genes was decreased in the hnRNP A1 heterozygous mice compared with the hnRNP A1 +/+ mice ( xref b )."

sparser
"Previous studies demonstrated that there were different splicing forms of the mef2c, lrrfip1, usp28 and abcc9 genes during muscle development [ xref – xref ]."
USP28 affects LDHA
| 2
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."
USP28 affects KITLG
| 2
USP28 inhibits KITLG.
| 1
USP28 inhibits KITLG. 1 / 1
| 1

reach
"USP28 antagonizes FBW7, an F-box protein and an important component of the SKP1-CUL1-F-BOX (SCF)-type E3 ubiquitin ligase that targets key transcriptional factors to ubiquitin-directed proteasome degradation."
USP28 deubiquitinates KITLG.
| 1
USP28 deubiquitinates KITLG. 1 / 1
| 1

reach
"In this setting, USP28 deubiquitinates the SCF component FBW7, allowing FBW7 to act as a substrate recognition factor targeting substrates for proteosomal mediated degradation."
USP28 affects JUN_family
| 2
| 2

reach
"USP28 has also been reported to antagonize ubiquitin-dependent degradation of the oncogene product MYC as well as JUN and Notch 105 ."

reach
"USP28 has also been reported to antagonize ubiquitin-dependent degradation of the oncogene product MYC as well as JUN and Notch ."
| 2

sparser
"Consistent with this notion, no complex formation was observed upon co-transfection of HA-USP28 and UCK1 in HEK293T cells, but coexpression of KLHL2 strongly stimulated the interaction between HA-USP28 and UCK1 ( xref A)."

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
USP28 affects HDAC5
| 1 1
| 1 1

reach
"In vitro pull-down assay using His tag recombinant LSD1 protein incubated with USP28-FLAG protein indicated a direct interaction of HDAC5 and USP28 (XREF_SUPPLEMENTARY), and HDAC5 overexpression significantly attenuated USP28 polyubiquitination (XREF_SUPPLEMENTARY)."

sparser
"In vitro pull-down assay using His-tag recombinant LSD1 protein incubated with USP28-FLAG protein indicated a direct interaction of HDAC5 and USP28 ( xref ), and HDAC5 overexpression significantly attenuated USP28 polyubiquitination ( xref )."
USP28 affects Fbw7α
| 2
USP28 binds MYC and Fbw7α. 2 / 2
| 2

sparser
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7α and c-Myc, which increased the levels of Ub-c-Myc."

sparser
"Although further studies did not show that hypoxia signaling impaired the association of USP28 with Fbw7α and c-Myc, cellular USP28 protein levels were decreased by Ni ions and hypoxia in A549 cells."
USP28 affects Fbw7alpha
| 2
USP28 binds Fbw7alpha.
| 1
USP28 binds Fbw7alpha. 1 / 1
| 1

reach
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7alpha and c-Myc, which increased the levels of Ub-c-Myc."
USP28 activates Fbw7alpha.
| 1
USP28 activates Fbw7alpha. 1 / 1
| 1

reach
"Nickel ions or hypoxia could impact the de-ubiquitination process by directly inhibiting USP28 activity or indirectly decreasing USP28 activity through either the dissociation of USP28 from the c, Myc, and Fbw7alpha complex or by a reduction of cellular USP28 levels."
USP28 affects Cyclin
| 2
Modified USP28 leads to the deubiquitination of Cyclin. 1 / 1
| 1

reach
"Overexpression of USP28 inhibited FBW7-depedent ubiquitination of c-MYC and cyclin E1, both of which are oncoproteins."
Modified USP28 leads to the deubiquitination of Cyclin on E1. 1 / 1
| 1

reach
"Overexpression of USP28 inhibited FBW7-depedent ubiquitination of c-MYC and cyclin E1, both of which are oncoproteins."
| 2
Modified USP28 activates Cell Survival. 1 / 1
| 1

reach
"Overexpression of USP28 in BR cells enhances c-Myc expression and hence increases ASS1 transcription upon arginine deprivation, and consequently leads to cell survival."
| 1

reach
"In conclusion, USP28 enhanced cell viability and aerobic glycolysis of colorectal cancer by stabilizing FOXC1, suggesting that USP28-FOXC1 might be a novel therapeutic avenue for colorectal cancer."
| 1 1
| 1 1

reach
"USP28 also enhances intestinal tumorigenesis."

eidos
"USP28 promotes tumorigenesis and has carcinogenic potential through deubiquitination ."
USP28 affects COMTD1
2 |
2 |

No evidence text available

No evidence text available
USP28 affects CD44s
| 2
USP28 binds CD44s. 2 / 2
| 2

reach
"Further studies illustrated that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing its protein stabilization."

reach
"Here, we discovered that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing this protein stability."
USP28 affects CD44s protein
| 2
Modified USP28 increases the amount of CD44s protein. 2 / 2
| 2

reach
"Intriguingly, ectopic expression of Flag-USP28 in T24T (shATG7) cells promoted CD44s protein expression and human BC sphere formation, revealing that the increased USP28 expression positively correlated with CD44s upregulation."

reach
"As expected, USP28 overexpression in T24T (shATG7 # 1) cells led to increased CD44s protein expression, as well as restoration of sphere formation (XREF_FIG - XREF_FIG)."
USP28 affects CD44 protein stability
| 2
USP28 activates CD44 protein stability. 2 / 2
| 2

eidos
"On the other hand , ISO also downregulated USP28 expression , which in turn reduced CD44 protein stability ."

eidos
"ISO attenuated CD44 protein stability by down-regulating USP28 in human BC cells Because ISO treatment not only inhibited endogenous CD44 expression but also reduced ectopic CD44 protein expression in T24T cell , the study next examined if down-regulation of CD44 expression due to ISO was a result of changes in degradation of CD44 protein ."
USP28 affects C-Cdh1
| 2
USP28 activates C-Cdh1. 2 / 2
| 2

reach
"The APC and C-Cdh 1 target Claspin, for instance, is protected from APC/C mediated down-regulation by USP28 [XREF_BIBR], and also 53BP1 and Chk2 were reported to be stabilized by USP28 upon DNA damage [XREF_BIBR]."

reach
"The APC and C-Cdh 1 target Claspin, for instance, is protected from APC/C mediated down-regulation by USP28 [XREF_BIBR], and also 53BP1 and Chk2 were reported to be stabilized by USP28 upon DNA damage [XREF_BIBR]."
USP28 affects C-Cdh1 targets
| 2
USP28 inhibits C-Cdh1 targets. 2 / 2
| 2

reach
"This model is supported by the observation that USP28 depletion does not lead to general degradation of APC and C-Cdh 1 targets, and the observation that Emi1 association with the APC/C does not appear to be controlled by DNA damage [XREF_BIBR]."

reach
"This model is supported by the observation that USP28 depletion does not lead to general degradation of APC and C-Cdh 1 targets, and the observation that Emi1 association with the APC/C does not appear to be controlled by DNA damage [XREF_BIBR]."
USP28 affects C
| 2
USP28 bound to MYC decreases the amount of C. 1 / 1
| 1

reach
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7alpha and c-Myc, which increased the levels of Ub-c-Myc."
USP28 bound to Fbw7alpha decreases the amount of C. 1 / 1
| 1

reach
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7alpha and c-Myc, which increased the levels of Ub-c-Myc."
USP28 affects BRCT
| 2
| 2

reach
"Furthermore, USP28 and p53 bind the BRCT repeats at different interfaces, and it has been shown that 53BP1 and USP28 share a co-regulatory role in supporting p53 functions [XREF_BIBR] (XREF_FIG)."

reach
"Furthermore, USP28 and p53 bind the BRCT repeats at different interfaces, and it has been shown that 53BP1 and USP28 share a coregulatory role in supporting p53 functions [20]."
USP28 affects ATR
| 1 1
USP28 deubiquitinates ATR.
| 1
USP28 deubiquitinates ATR. 1 / 1
| 1

reach
"Mechanistically, USP28 deubiquitinates and stabilizes the specificity factors and mediators of ATM and ATR signaling by protecting them from ubiquitination-mediated proteasome degradation."
USP28 binds ATR.
| 1
| 1

sparser
"Notably, binding of p-ATR and ∆Np63 to USP28 was increased after CPPD treatment (Fig.  xref )."
USP28 affects ATM
| 2
USP28 deubiquitinates ATM.
| 1
USP28 deubiquitinates ATM. 1 / 1
| 1

reach
"Mechanistically, USP28 deubiquitinates and stabilizes the specificity factors and mediators of ATM and ATR signaling by protecting them from ubiquitination-mediated proteasome degradation."
USP28 activates ATM.
| 1
USP28 activates ATM. 1 / 1
| 1

reach
"This early disassembly of the MRN complex could be prevented by altering the ubiquitination site of Nbs1 or by expressing a deubiquitinase, Usp28, which sufficiently restored homologous recombination repair and ATM, a major checkpoint kinase against DNA DSBs, activation abilities in RTS and RecQL4-depleted cells."
USP28 affects APC
| 2
USP28 activates APC. 2 / 2
| 2

reach
"The APC and C-Cdh 1 target Claspin, for instance, is protected from APC/C mediated down-regulation by USP28 [XREF_BIBR], and also 53BP1 and Chk2 were reported to be stabilized by USP28 upon DNA damage [XREF_BIBR]."

reach
"The APC and C-Cdh 1 target Claspin, for instance, is protected from APC/C mediated down-regulation by USP28 [XREF_BIBR], and also 53BP1 and Chk2 were reported to be stabilized by USP28 upon DNA damage [XREF_BIBR]."
USP21 affects USP28
2 |
2 |

No evidence text available

No evidence text available
TP53BP1 affects CDKN1A
| 2
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."
SNAI2 affects USP28
| 2
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."
POU2F1 affects USP28
2 |
POU2F1 decreases the amount of USP28. 2 / 2
2 |

No evidence text available

No evidence text available
MYC affects SNAI2
| 2
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."
MYC affects Fbw7α
| 2
USP28 binds MYC and Fbw7α. 2 / 2
| 2

sparser
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7α and c-Myc, which increased the levels of Ub-c-Myc."

sparser
"Although further studies did not show that hypoxia signaling impaired the association of USP28 with Fbw7α and c-Myc, cellular USP28 protein levels were decreased by Ni ions and hypoxia in A549 cells."
MEF2C affects USP28
| 2
| 2

sparser
"The full length compared with the truncated forms of the mef2c, lrrfip1, usp28 and abcc9 genes was decreased in the hnRNP A1 heterozygous mice compared with the hnRNP A1 +/+ mice ( xref b )."

sparser
"Previous studies demonstrated that there were different splicing forms of the mef2c, lrrfip1, usp28 and abcc9 genes during muscle development [ xref – xref ]."
LDHA affects SNAI2
| 2
SNAI2 binds USP28, LDHA, and MYC. 2 / 2
| 2

sparser
"Immunohistochemical staining was performed on these tissues, and the results revealed that high serum epinephrine was positively associated with high LDHA, USP28, MYC, and SLUG protein expression ( xref and xref )."

sparser
"Our data showed that high epinephrine in patient serum was positively associated with high LDHA, USP28, MYC, and SLUG expression and conferred lower overall survival and disease-free survival rates compared with those of patients with low epinephrine levels."
Hydrogen-Ion Concentration leads to the deubiquitination of USP28. 2 / 2
| 2

reach
"Chronic stress induced epinephrine activated LDHA to generate lactate, and the adjusted pH directed USP28 mediated deubiquitination and stabilization of MYC."

reach
"This leads to a switch to lactate production, and the adjusted pH then directs USP28 mediated deubiquitination and stabilization of MYC, thereby promoting stem like traits in breast cancer."
| 2

sparser
"Consistent with this notion, no complex formation was observed upon co-transfection of HA-USP28 and UCK1 in HEK293T cells, but coexpression of KLHL2 strongly stimulated the interaction between HA-USP28 and UCK1 ( xref A)."

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
Fbw7α affects USP28
| 2
USP28 binds MYC and Fbw7α. 2 / 2
| 2

sparser
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7α and c-Myc, which increased the levels of Ub-c-Myc."

sparser
"Although further studies did not show that hypoxia signaling impaired the association of USP28 with Fbw7α and c-Myc, cellular USP28 protein levels were decreased by Ni ions and hypoxia in A549 cells."
| 2

sparser
"A deubiquitinase USP28 binds to the SCF Fbw7 ubiquitin E3 ligase, stabilizing Myc, and therefore promoting cell proliferation [ xref ]."

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
COMTD1 affects USP28
2 |
2 |

No evidence text available

No evidence text available
CEP131 affects USP28
| 1 1
CEP131 inhibits USP28. 2 / 2
| 1 1

sparser
"We observed biochemical inhibition of USP25 and USP28 by AZ1 with IC 50 s of 3.47 ± 0.73 μM (n = 4) and 0.98 ± 0.02 μM (n = 2) respectively, in line with reported values for the compound."

eidos
"26 HepG2 and HuH6 cells were treated with different concentrations of AZ1 for 48 h , and AZ1 prominently decreased the levels of USP28 itself and that of TCF7L2 , as well as the target genes of TCF7L2 , AXIN2 , and Cyclin D1 ( Figure 6A ) ."
CDKN1A affects USP28
| 2
| 2

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53p21 signaling as well as screening for the upstream components that transduce the s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Important future directions include dissecting the interactions between 53BP1 and USP28 that activate downstream p53-p21 signaling, as well as screening for the upstream components that transduce the signal (see xref )."
CD44s affects USP28
| 2
USP28 binds CD44s. 2 / 2
| 2

reach
"Further studies illustrated that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing its protein stabilization."

reach
"Here, we discovered that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing this protein stability."
BRCT affects USP28
| 2
| 2

reach
"Furthermore, USP28 and p53 bind the BRCT repeats at different interfaces, and it has been shown that 53BP1 and USP28 share a co-regulatory role in supporting p53 functions [XREF_BIBR] (XREF_FIG)."

reach
"Furthermore, USP28 and p53 bind the BRCT repeats at different interfaces, and it has been shown that 53BP1 and USP28 share a coregulatory role in supporting p53 functions [20]."
ABCC9 affects MEF2C
| 2
| 2

sparser
"The full length compared with the truncated forms of the mef2c, lrrfip1, usp28 and abcc9 genes was decreased in the hnRNP A1 heterozygous mice compared with the hnRNP A1 +/+ mice ( xref b )."

sparser
"Previous studies demonstrated that there were different splicing forms of the mef2c, lrrfip1, usp28 and abcc9 genes during muscle development [ xref – xref ]."
Vorinostat affects USP28
1 |
Vorinostat increases the amount of USP28. 1 / 1
1 |

No evidence text available
Vitamin C affects USP28
| 1
| 1

eidos
"And that vitamin C reversed the induction of stem-like phenotype by inhibition of stress-induced LDH-A , and suppressing the lactic acid generation and the USP28 / MYC / SLUG pathway in BCSCs ."
Unspecified redundant mechanisms affects USP28
| 1
Unspecified redundant mechanisms activates USP28. 1 / 1
| 1

eidos
"APC , adenomatous polyposis coli ; DVL , Disheveled ; Fz , frizzled ; GSK3 , glycogen synthase kinase-3 ; LRP , lipoprotein receptor-related ; USP28 , ubiquitin-specific protease 28 As Usp28-deficient mice develop normally , the role of Usp28 in the normal Wnt signaling during embryonic development must be minimal or its absence could be mitigated by unspecified redundant mechanisms , for example , suppressing GSK3beta activity so that TCF7L2 is not ubiquitinated in the first place ."
Triclosan affects USP28
1 |
Triclosan decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Tributylstannane decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Sodium arsenite increases the amount of USP28. 1 / 1
1 |

No evidence text available
ShRNA-mediated affects USP28
| 1
ShRNA-mediated activates USP28. 1 / 1
| 1

eidos
"( A ) Western blot analysis of TCF7L2 in HEK293T cells treated with MG132 ( 20 muM ) for 3 h followed USP28 depletion by shRNA-mediated knockdown ."
Set benzylaminoethanol affects USP28
| 1
Set benzylaminoethanol inhibits USP28. 1 / 1
| 1

eidos
"Recently , a high throughput screening ( HTS ) identified the first set of benzylaminoethanol compounds AZ1 , AZ2 , and AZ4 that specifically inhibit USP28 ( 99 ) ."
| 1
Rigosertib sodium activates mutated USP28. 1 / 1
| 1

reach
"The combination of these results stresses the need for further investigation to determine the precise mechanism of rigosertib specifically targeting USP28 mutant cells."
1 |
Nickel sulfate decreases the amount of USP28. 1 / 1
1 |

No evidence text available
Nickel atom affects USP28
1 |
Nickel atom decreases the amount of USP28. 1 / 1
1 |

No evidence text available
New affects USP28
| 1
New inhibits USP28. 1 / 1
| 1

eidos
"Liu et al. ( 100 ) reported a new compound 19 , a [ 1,2,3 ] triazolo [ 4,5 - d ] pyrimidine derivative , that specifically inhibits USP28 and reduces gastric cancer cell proliferation and EMT with a better IC50 than that of AZ1 ."
Mono(2-ethylhexyl) phthalate decreases the amount of USP28. 1 / 1
1 |

No evidence text available
MiR-92b-3p affects USP28
| 1
MiR-92b-3p activates USP28. 1 / 1
| 1

reach
"we found that there was a significant negative relation between the level of miR-92b-3p and USP28 at protein level and reversed the down regulation of miR-92b-3p by hypoxia can suppress the proliferation of pulmonary artery smooth muscle cells by targeting USP28."
MiR-4295 maturation affects USP28
| 1
MiR-4295 maturation inhibits USP28. 1 / 1
| 1

eidos
"The present study found that ISO treatment led to time-dependent increases in Dicer protein abundance , which , in turn , subsequently promoted miR-4295 maturation that resulted in down-regulation of USP28 , acceleration of CD44 protein degradation , and ultimately brought about a significant reduction in stem-like properties of the human BC cells ."
MiR-363-3p affects USP28
| 1
MiR-363-3p inhibits USP28. 1 / 1
| 1

reach
"Furthermore, miR-363-3p suppresses the proliferation of human hepatocellular carcinoma cells by targeting S1PR1 or USP28 [48]."
MiR-143-3p affects USP28
| 1
MiR-143-3p increases the amount of USP28. 1 / 1
| 1

reach
"Mechanistically, LINC01426 acted as a competing endogenous RNA (ceRNA) for miR-143-3p to upregulate USP28 expression."
Methylmercury chloride increases the amount of USP28. 1 / 1
1 |

No evidence text available
Leflunomide affects USP28
1 |
Leflunomide decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-93-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-92a-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-615-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-5589-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-519d-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-493-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-4729 decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-4668-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-3940-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-367-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-340-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-32-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-25-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-221-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-20b-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-20a-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-20a-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-181a-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-17-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-153-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-1301-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-129-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-122-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-miR-106b-5p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-let-7g-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-let-7c-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Hsa-let-7a-2-3p decreases the amount of USP28. 1 / 1
1 |

No evidence text available
Hif2 affects USP28
| 1
Hif2 increases the amount of USP28. 1 / 1
| 1

reach
"There were two scenarios that resulted in an increase in ubiquitinated c-Myc levels in A549 cells : the first involved an increased phosphorylation of c-Myc at T58; and the second involved a decrease in USP28 protein levels partially mediated by HIF-1 and HIF-2."
H treatment SGH muM PC9 cells muM H157 cells affects USP28
| 1
H treatment SGH muM PC9 cells muM H157 cells inhibits USP28. 1 / 1
| 1

eidos
"A 24 h treatment of SGH at 2 muM for PC9 cells and 5 muM for H157 cells downregulated USP28 and c-Myc expressions ( * * p < 0.01 or * * * p < 0.001 , Figure 4A , B and Figure S7A , B ) ."
1 |
Formaldehyde decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Dorsomorphin increases the amount of USP28. 1 / 1
1 |

No evidence text available
Diclofenac affects USP28
1 |
Diclofenac increases the amount of USP28. 1 / 1
1 |

No evidence text available
Decabromodiphenyl ether increases the amount of USP28. 1 / 1
1 |

No evidence text available
D pyrimidine derivative affects USP28
| 1
D pyrimidine derivative inhibits USP28. 1 / 1
| 1

eidos
"Liu et al. ( 100 ) reported a new compound 19 , a [ 1,2,3 ] triazolo [ 4,5 - d ] pyrimidine derivative , that specifically inhibits USP28 and reduces gastric cancer cell proliferation and EMT with a better IC50 than that of AZ1 ."
CircRNA FBXW7 affects USP28
| 1
CircRNA FBXW7 inhibits USP28. 1 / 1
| 1

eidos
"The microRnas miR-92b-3p , miR-216b , miR-500a-5p , and miR-3940-5p negatively regulate USP28 expression blocking its translation [ 22,23,24,25 ] while the circRNA FBXW7 reduce USP28 activity , decreasing substrate recognition [ 26 ] ."
| PMC
CircMED27 affects USP28
| 1
CircMED27 increases the amount of USP28. 1 / 1
| 1

reach
"Our mechanistic investigations revealed that circMED27 functions as a competing endogenous RNA (ceRNA) for miR-655-3p to upregulate ubiquitin-specific peptidase 28 (USP28) expression."
Chloroquine affects USP28
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Chloroquine increases the amount of USP28. 1 / 1
1 |

No evidence text available
Cadmium dichloride increases the amount of USP28. 1 / 1
1 |

No evidence text available
Brought stem-like properties human BC cells affects USP28
| 1
Brought stem-like properties human BC cells activates USP28. 1 / 1
| 1

eidos
"The present study found that ISO treatment led to time-dependent increases in Dicer protein abundance , which , in turn , subsequently promoted miR-4295 maturation that resulted in down-regulation of USP28 , acceleration of CD44 protein degradation , and ultimately brought about a significant reduction in stem-like properties of the human BC cells ."

No evidence text available
1 |
Benzo[a]pyrene decreases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |
Aluminium atom decreases the amount of USP28. 1 / 1
1 |

No evidence text available
ZCCHC8 affects USP28
1 |
1 |

No evidence text available
ZC3H4 affects USP28
1 |
1 |

No evidence text available
UbV.USP28 affects USP28
| 1
UbV.USP28 inhibits USP28. 1 / 1
| 1

reach
"Next, we explored whether UbV.USP28 could inhibit the function of USP28 in live cells."
UbV.28 affects USP28
| 1
USP28 binds UbV.28. 1 / 1
| 1

reach
"Among them, UbV.28 was able to bind USP28 and inhibit the hydrolysis of the poly-Ub chains but not Ub-AMC, a mono-Ub fluorogenic substrate lacking a proximal Ub [117] ."
USP8 affects USP28
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No evidence text available
USP7 affects USP28
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1 |

No evidence text available
USP51 affects USP28
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1 |

No evidence text available
USP5 affects USP28
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1 |

No evidence text available
USP4 affects USP28
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1 |

No evidence text available
USP33 affects USP28
1 |
1 |

No evidence text available
USP28Delta 53BP1Delta affects USP28
| 1
USP28Delta 53BP1Delta activates USP28. 1 / 1
| 1

reach
"Moreover, the N3 resistance of a USP28Delta 53BP1Delta double-knockout line was not enhanced over single mutants (XREF_FIG C), confirming USP28 and 53BP1 function epistatically in this context."
1 |
1 |

No evidence text available
USP28 affects vincristine
| 1
| 1

reach
"We also identify the genes KIF18B and USP28 of which CRISPR/Cas9-directed knockout induces vincristine resistance across two DLBCL cell lines."
USP28 affects viability gastric cancer cells
| 1
USP28 activates viability gastric cancer cells. 1 / 1
| 1

eidos
"To validate our hypothesis that inhibition USP28 could decrease the viability of gastric cancer cells , we established an USP28 knockdown ( KD ) HGC-27 cell line using USP28 siRNA transfection ( Fig. 3C ) ."
USP28 affects ubiquitination transcription 7-like TCF7L2
| 1
USP28 activates ubiquitination transcription 7-like TCF7L2. 1 / 1
| 1

eidos
"Ubiquitin-specific protease 28 ( USP28 ) interacts with and modulates ubiquitination of transcription factor 7-like 2 ( TCF7L2 ) ."
USP28 affects ubiquitination modification c-Myc
| 1
USP28 inhibits ubiquitination modification c-Myc. 1 / 1
| 1

eidos
"USP28 can bind to FBXW7 and inhibit ubiquitination modification of c-Myc by the latter , thus increasing the protein stability of c-Myc ( 89 ) ."
USP28 affects ubiquitination LEF1
| 1
USP28 inhibits ubiquitination LEF1. 1 / 1
| 1

eidos
"The knockdown of USP28 did result in an increase in the ubiquitination of LEF1 ( Figure S3F ) but did not increase that of TCF7L1 ( Figure S3G ) ."
USP28 affects turnover
| 1
USP28 inhibits turnover. 1 / 1
| 1

sparser
"Furthermore, overexpressing USP28 inhibited FBXW7-185aa-induced c-Myc turnover ( xref )."
USP28 affects tumor-suppressor genes
| 1
USP28 activates tumor-suppressor genes. 1 / 1
| 1

reach
"Lastly, USP28 also acts as a tumor suppressor, inducing H2A-mediated transcriptional activation of tumor-suppressor genes [43]."
| PMC
USP28 affects trisulfur
| 1
| 1

reach
"Using an ATP based cell viability assay (Cell Titer-Glo), we screened a small library of 316 FDA approved chemical compounds and identified the PLK1 inhibitor rigosertib as a compound that selectively impairs the viability of USP28 depleted cells (XREF_FIG and Table S3)."
USP28 affects treatment tumors
| 1
USP28 inhibits treatment tumors. 1 / 1
| 1

eidos
"The inhibition of USP28 was particularly effective in the treatment of SCC tumors using murine models [ 17 ] ."
| PMC
USP28 affects translation
| 1
| 1

reach
"The microRnas miR-92b-3p, miR-216b, miR-500a-5p, and miR-3940-5p negatively regulate USP28 expression blocking its translation [22,23,24,25] while the circRNA FBXW7 reduce USP28 activity, decreasing substrate recognition [26]."
| PMC
USP28 affects target c-MYC
| 1
USP28 inhibits target c-MYC. 1 / 1
| 1

eidos
"c-MYC exemplifies a classically `` undruggable '' protein [ 57 ] , however its protein level is regulated by USP28 activity [ 58 ] raising the possibility of USP28 inhibition to target c-MYC levels [ 59,60 ] ."
USP28 affects stability PIM1 hypoxia
| 1
USP28 activates stability PIM1 hypoxia. 1 / 1
| 1

eidos
"In conclusion , USP28 blocked the ubiquitination and increased the stability of PIM1 / 2 , particularly in hypoxia ."
USP28 affects stability FBW7-substrates cancer cells
| 1
USP28 activates stability FBW7-substrates cancer cells. 1 / 1
| 1

eidos
"USP28 can counteract the activity of FBW7 and promote the stability of FBW7-substrates in cancer cells 40 ."
USP28 affects stability CHK2
| 1
USP28 activates stability CHK2. 1 / 1
| 1

eidos
"Considering that CHK2 phosphorylates and stabilizes p53 , inducing its apoptotic activity upon DNA damage , it can be deduced that USP28 acts as a tumor suppressor when it increases the stability of CHK2 ."
| PMC
USP28 affects radiosensitivity esophageal cancer cells
| 1
USP28 inhibits radiosensitivity esophageal cancer cells. 1 / 1
| 1

eidos
"The loss of USP28 enhances the radiosensitivity of esophageal cancer cells via the c-Myc pathway [ 38 ] ."
USP28 affects protein stability c-Myc
| 1
USP28 activates protein stability c-Myc. 1 / 1
| 1

eidos
"USP28 can bind to FBXW7 and inhibit ubiquitination modification of c-Myc by the latter , thus increasing the protein stability of c-Myc ( 89 ) ."

eidos
"The resultant weakly acidic microenvironment enhances the deubiquitination of MYC by USP28 , improves the stability of MYC , and activates the promoter of Slug , ultimately promoting the stem cell-like properties of breast cancer ."
USP28 affects p53 protein stability
| 1
USP28 activates p53 protein stability. 1 / 1
| 1

eidos
"The mechanism proposed was that 53BP1 and USP28 promote p53 protein stability specifically in the context of mitotic stress , but not by DNA damage or by Nutlin-3 ."
USP28 affects p21-dependent cell cycle
| 1
USP28 inhibits p21-dependent cell cycle. 1 / 1
| 1

eidos
"Following p53 localization under such conditions , 53BP1 and USP28 can mediate p53 activation and p21-dependent cell cycle arrest [ 119 ] ."
USP28 affects p-Chk1 protein
| 1
USP28 activates p-Chk1 protein. 1 / 1
| 1

reach
"These results suggest that USP28 specifically enhances p-Chk1 protein stabilization through a de-ubiquitination event of Claspin protein."
USP28 affects oncogenic transformation respiratory cells
| 1
USP28 activates oncogenic transformation respiratory cells. 1 / 1
| 1

eidos
"USP28 enables oncogenic transformation of respiratory cells and its inhibition potentiates molecular therapy targeting mutant EGFR , BRAF and PI3K ."
USP28 affects oncogenic reprogramming
| 1
USP28 activates oncogenic reprogramming. 1 / 1
| 1

eidos
"Here , we report that the ubiquitin carboxyl-terminal hydrolase 28 ( USP28 ) enables oncogenic reprogramming by regulating the protein abundance of proto-oncogenes such as c-JUN , c-MYC , NOTCH and incrementNP63 at early stages of malignant transformation ."
USP28 affects mitosis
| 1
USP28 activates mitosis. 1 / 1
| 1

reach
"53BP1 and USP28 mediate p53 dependent cell cycle arrest in response to centrosome loss and prolonged mitosis."
USP28 affects miR-500a-5p breast cancer cell metastasis
| 1
USP28 inhibits miR-500a-5p breast cancer cell metastasis. 1 / 1
| 1

eidos
"In accordance with these findings , our study revealed that USP28 , as the target gene of miR-500a-5p , reversed the ability of miR-500a-5p to promoted breast cancer cell proliferation and metastasis , indicating that USP28 functions as a breast cancer suppressor ."
USP28 affects melanoma cell resistance
| 1
USP28 inhibits melanoma cell resistance. 1 / 1
| 1

eidos
"Therefore , USP28 depletion increased BRAF protein levels and melanoma cell resistance to BRAF inhibitors [ 57 ] ."
| PMC
| 1
| 1

reach
"Following p53 localization under such conditions, 53BP1 and USP28 can mediate p53 activation and p21-dependent cell cycle arrest [119]."
USP28 affects lifespans
| 1
USP28 inhibits lifespans. 1 / 1
| 1

eidos
"USP28 gene expression correlated with poor prognosis and shortened lifespans in SCC patients [ 17 ] ."
| PMC
USP28 affects inhibition potentiates molecular therapy targeting mutant BRAF PI3K
| 1
USP28 activates inhibition potentiates molecular therapy targeting mutant BRAF PI3K. 1 / 1
| 1

eidos
"USP28 enables oncogenic transformation of respiratory cells and its inhibition potentiates molecular therapy targeting mutant EGFR , BRAF and PI3K ."

reach
"This early disassembly of the MRN complex could be prevented by altering the ubiquitination site of Nbs1 or by expressing a deubiquitinase, Usp28, which sufficiently restored homologous recombination repair and ATM, a major checkpoint kinase against DNA DSBs, activation abilities in RTS and RecQL4-depleted cells."
USP28 affects growth lung cancer cells
| 1
USP28 activates growth lung cancer cells. 1 / 1
| 1

eidos
"Downregulating USP28 expression can inhibit the proliferation and growth of lung cancer cells [ 8,23 ] ."
USP28 affects growth HepG2 Figures 1F S2A B HuH6 Figures 1G
| 1
USP28 activates growth HepG2 Figures 1F S2A B HuH6 Figures 1G. 1 / 1
| 1

eidos
"Indeed , the depletion of USP28 led to much slowed growth of HepG2 ( Figures 1F and S2A , B ) and HuH6 ( Figures 1G and S2C , D ) cells ."
USP28 affects different isoforms
| 1
USP28 activates different isoforms. 1 / 1
| 1

eidos
"USP28 can generate different isoforms by alternative splicing ."
| PMC
USP28 affects cellular MYC
| 1
USP28 activates cellular MYC. 1 / 1
| 1

eidos
"By inhibiting USP28 , these compounds indeed reduce cellular MYC levels , induce apoptosis , and inhibit cell proliferation in a dose-dependent manner ( 99 ) ."
USP28 affects cell
| 1
USP28 inhibits cell. 1 / 1
| 1

eidos
"Previous reports showed that 53BP1 and USP28 activate p53 , preventing the proliferation of cells that have an increased chance of mitotic errors [ 104 ] ."
| PMC
USP28 affects cell viability HGC-27 cells
| 1
USP28 activates cell viability HGC-27 cells. 1 / 1
| 1

eidos
"Expectedly , USP28 knockdown also resulted in decrease of cell viability of HGC-27 cells ( Fig. 3D ) , indicating that the level of USP28 plays a crucial role on the cell viability ."
USP28 affects c-MYC ubiquitylation
| 1
USP28 inhibits c-MYC ubiquitylation. 1 / 1
| 1

eidos
"In addition , a decrease in USP28 expression contributed to a higher level of c-MYC ubiquitylation and thus promoted degradation of this oncoprotein [ 138 ] ."
USP28 affects breast cancer cell
| 1
USP28 activates breast cancer cell. 1 / 1
| 1

eidos
"In accordance with these findings , our study revealed that USP28 , as the target gene of miR-500a-5p , reversed the ability of miR-500a-5p to promoted breast cancer cell proliferation and metastasis , indicating that USP28 functions as a breast cancer suppressor ."
USP28 affects aerobic glycolysis
| 1
USP28 activates aerobic glycolysis. 1 / 1
| 1

eidos
"USP28 increases aerobic glycolysis and promotes cell proliferation upon stabilization of FOXC1 [ 50 ] ."
| PMC
USP28 affects aerobic glycolysis colorectal cancer
| 1
USP28 activates aerobic glycolysis colorectal cancer. 1 / 1
| 1

eidos
"USP28 promotes aerobic glycolysis of colorectal cancer by increasing stability of FOXC1 ."
USP28 affects accumulate HIF-1alpha protein cells
| 1
USP28 activates accumulate HIF-1alpha protein cells. 1 / 1
| 1

eidos
"Results Here , we report that USP28 is a SUMOylated protein in normoxia with moderate deubiquitinating activity towards HIF-1alpha in vitro , while hypoxia and HIF-1alpha activate USP28 through SENP1-mediated USP28 deSUMOylation to further accumulate HIF-1alpha protein in cells ."
USP28 affects ZCCHC8
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1 |

No evidence text available
USP28 affects ZC3H4
1 |
1 |

No evidence text available
USP28 affects Wnt/β-catenin
| 1
USP28 activates Wnt/β-catenin. 1 / 1
| 1

reach
"By regulating the levels of TCF7L2, USP28 modulates the Wnt/β-catenin signaling in liver cancer and USP28 depletion or inhibition by a small molecule inhibitor leads to a halt of growth in liver cancer cells."
USP28 affects VHL
| 1
USP28 leads to the ubiquitination of VHL. 1 / 1
| 1

reach
"This recruits Fbw7 and USP28, which then mediates HIF-1alpha ubiquitination and subsequent VHL independent proteasomal degradation [XREF_BIBR]."
| 1

eidos
"In conclusion , USP28 blocked the ubiquitination and increased the stability of PIM1 / 2 , particularly in hypoxia ."
USP28 affects UbV.28
| 1
USP28 binds UbV.28. 1 / 1
| 1

reach
"Among them, UbV.28 was able to bind USP28 and inhibit the hydrolysis of the poly-Ub chains but not Ub-AMC, a mono-Ub fluorogenic substrate lacking a proximal Ub [117] ."
USP28 affects USP8
1 |
1 |

No evidence text available
USP28 affects USP7
1 |
1 |

No evidence text available
USP28 affects USP51
1 |
1 |

No evidence text available
USP28 affects USP5
1 |
1 |

No evidence text available
USP28 affects USP4
1 |
1 |

No evidence text available
USP28 affects USP33
1 |
1 |

No evidence text available
USP28 affects USP28Delta cells
| 1
USP28 inhibits USP28Delta cells. 1 / 1
| 1

eidos
"USP28Delta cells were used as the control because inactivation of USP28 prevents p53 activation and G1 arrest that is observed as a consequence of delayed mitosis following centrosome loss in RPE1 cells13 ."
USP28 affects USP20
1 |
1 |

No evidence text available
USP28 affects USP15
1 |
1 |

No evidence text available
USP28 affects USP14
1 |
1 |

No evidence text available
USP28 affects USP
| 1
USP28 binds MYC and USP. 1 / 1
| 1

sparser
"Importantly, our results establish that the MYC MBI domain (1–95 amino acids) interacts directly with the USP domain (160–652 amino acids) of USP28."
USP28 affects USO1
1 |
1 |

No evidence text available
USP28 affects UCHL5
1 |
1 |

No evidence text available
USP28 affects UCHL1
1 |
1 |

No evidence text available
USP28 affects TWF1
1 |
1 |

No evidence text available
USP28 affects TTC39A
| 1
| 1

sparser
"To further examine the USP28-LSD1 axis in regulating CSC-like properties in human breast cancer, we measured tumorsphere formation in BT549 and MCF7 cells with knockdown of USP28."
USP28 affects TRUSS
| 1
USP28 activates TRUSS. 1 / 1
| 1

reach
"Other proteins regulating C-MYC stability include : CIP2A, a C-MYC-interacting protein that specifically inhibits PP2A activity against C-MYC; USP28, a de-ubiquitinating enzymes that antagonizes FBW7 and promotes C-MYC stability and TRUSS, a receptor for DDB1 (damage specific DNA binding protein 1)-CUL4 (Cullin 4) E3 ligase complex."
USP28 affects TRIM63
1 |
1 |

No evidence text available
USP28 affects TRIM55
1 |
1 |

No evidence text available
USP28 affects TCF7L1
| 1
USP28 inhibits TCF7L1. 1 / 1
| 1

eidos
"The knockdown of USP28 did result in an increase in the ubiquitination of LEF1 ( Figure S3F ) but did not increase that of TCF7L1 ( Figure S3G ) ."
USP28 affects Snail1
| 1
USP28 increases the amount of Snail1. 1 / 1
| 1

reach
"When these DUBs were co-expressed with Snail1 in HEK293 cells, we found that USP12, Dub3 and USP28 significantly increased Snail1 levels, similar to results obtained when cells were treated with the proteasome inhibitor MG132 (XREF_SUPPLEMENTARY)."
USP28 affects SYK
| 1
| 1

sparser
"In addition, SYK does not bind to USP28 in the same assay ( Fig. 1 B, left panel)."
USP28 affects STAMBP
1 |
1 |

No evidence text available
USP28 affects STAM
1 |
1 |

No evidence text available
USP28 affects ST6GAL1
1 |
1 |

No evidence text available
USP28 affects SENP8
1 |
1 |

No evidence text available
USP28 affects SENP2
1 |
1 |

No evidence text available
USP28 affects RP23-446O17.1
1 |
USP28 binds RP23-446O17.1. 1 / 1
1 |

No evidence text available
USP28 affects RP23-437L13.2
1 |
USP28 binds RP23-437L13.2. 1 / 1
1 |

No evidence text available
USP28 affects RFFL
1 |
1 |

No evidence text available
USP28 affects RBM24
| 1
| 1

sparser
"The alternative splicing forms of usp28 and abcc9 changed in rbm24 defect mice, causing dilated atria [ xref ]."
USP28 affects RB1
| 1
USP28 inhibits RB1. 1 / 1
| 1

reach
"As expected, USP28 overexpression further decreased the relative expression of cell cycle related genes (CDK4, CDK6, Cyclin D1, pRB, and E2F1) induced by 5 '-AZA, whereas p15 and p27 expression was significantly enhanced."
| 1

eidos
"Independently , USP28 has been reported to modulate the activity of the Myc proto-oncogene [ 126 ] ."
USP28 affects Proteasome
| 1
| 1

reach
"miR-363 directly inhibits a ubiquitin specific protease, USP28 to promote proteasome mediated degradation of Myc in human hepatocellular carcinoma [XREF_BIBR]."

reach
"Inhibition of deubiquitinase USP28 attenuates cyst growth in autosomal dominant polycystic kidney disease."
USP28 affects PLA2G2A
1 |
1 |

No evidence text available
USP28 affects PJA1
1 |
1 |

No evidence text available
USP28 affects PIN1
| 1
USP28 activates PIN1. 1 / 1
| 1

reach
"Thus, complete loss of Usp28 promotes Pin1 dependent degradation of Fbw7 by the proteasome, most likely via enhanced autocatalytic ubiquitination.Destabilization of Fbw7 in Usp28 -/- cells suggested t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 affects PIM1/2
| 1
USP28 binds PIM1/2. 1 / 1
| 1

reach
"Interestingly, coimmunoprecipitation assays showed an increased interaction between PIM1/2 and USP28 in response to hypoxia, which correlated with reduced ubiquitination and increased protein stability."
USP28 affects PIM
| 1
USP28 activates PIM. 1 / 1
| 1

reach
"The overexpression of USP28 increased PIM protein stability and total levels in both normoxia and hypoxia, and USP28-knockdown significantly increased the ubiquitination of PIM1 and PIM2."
USP28 affects PFAS
1 |
1 |

No evidence text available
USP28 affects OTUB1
1 |
1 |

No evidence text available

reach
"Transwell assays showed that USP28 overexpression reduced both cell migration and invasion in MDA-MB-231 and MCF7 cells, and these effects were partially reversed by transfection with miR-500a-5p mimics."
USP28 affects NUMA1
| 1
| 1

sparser
"Using mass spectrometry and immunoprecipitation, we found that USP28 interacts with NuMA1 and affects centrosome clustering."
USP28 affects NSMCE2
1 |
1 |

No evidence text available
USP28 affects NSCLC
| 1
USP28 activates NSCLC. 1 / 1
| 1

reach
"The exact mechanism by which USP28 promotes NSCLC cell proliferation is undetermined yet."
USP28 affects MVP
| 1
USP28 increases the amount of MVP. 1 / 1
| 1

reach
"Our results show that silencing of USP28 reduces expression of MVP enzymes and lowers metabolic flux into this pathway."
USP28 affects MTNR1A
1 |
1 |

No evidence text available
USP28 affects MRPS7
1 |
1 |

No evidence text available
USP28 affects MAP3K1
1 |
1 |

No evidence text available
USP28 affects LIN28A
| 1
USP28 inhibits LIN28A. 1 / 1
| 1

reach
"USP28, through its deubiquitinating activity, antagonizes LIN28A protein turnover by reversing its proteasomal degradation."
USP28 affects KRAS
| 1
USP28 increases the amount of KRAS. 1 / 1
| 1

reach
"However, in contrast to the results in KRAS positive CRC cell lines, in WI-38 cells KRAS failed to increase ZNF304 protein levels (XREF_FIG) or significantly stimulate USP28 and PRKD1 transcription (XREF_FIG), explaining at least in part why KRAS expression does not result in INK4-ARF silencing."
USP28 affects JOSD1
1 |
1 |

No evidence text available
USP28 affects ISO-induced BC cell sphere
| 1
USP28 inhibits ISO-induced BC cell sphere. 1 / 1
| 1

eidos
"Moreover , USP28 expression attenuated the ISO-induced inhibition of BC cell sphere formation ( Fig. 2F & S2B ) ."
USP28 affects Hypoxia
| 1
USP28 activates Hypoxia. 1 / 1
| 1

reach
"The overexpression of USP28 increased PIM protein stability and total levels in both normoxia and hypoxia, and USP28-knockdown significantly increased the ubiquitination of PIM1 and PIM2."
USP28 affects Glioma
| 1
USP28 activates Glioma. 1 / 1
| 1

reach
"Ectopic USP28 expression promoted proliferation of SW1783 glioma cells both in vitro and in vivo, and conferred enhanced tumorigenicity in a nude mouse model."
USP28 affects GYS
| 1
USP28 activates GYS. 1 / 1
| 1

reach
"A study has proposed a novel mechanism by which USP28 upregulates angiogenesis by antagonizing GSK-3beta (glycogen synthase kinase-3beta) and FBW7 dependent degradation of HIF-1alpha (hypoxia inducible factor-1alpha), a major regulator of angiogenesis, carcinogenesis, and various processes by which cells adapt to hypoxic conditions."
USP28 affects G1
| 1
USP28 activates G1. 1 / 1
| 1

eidos
"USP28Delta cells were used as the control because inactivation of USP28 prevents p53 activation and G1 arrest that is observed as a consequence of delayed mitosis following centrosome loss in RPE1 cells13 ."
USP28 affects Forkhead
| 1
USP28 increases the amount of Forkhead. 1 / 1
| 1

reach
"Protein expression of Forkhead Box C1 (FOXC1) was increased by USP28 over-expression, whereas knockdown of USP28 aggravated cycloheximide (CHX; protein synthesis inhibitor) stimulated decrease of FOXC1."
USP28 affects FOXK1
1 |
1 |

No evidence text available
USP28 affects FBW7alpha
| 1
USP28 binds FBW7alpha. 1 / 1
| 1

reach
"The effect is limited to FBW7alpha because Usp28 specifically interacts with FBW7alpha and this interaction mediates complex formation between MYC and Usp28."
USP28 affects E3-ligases
| 1
USP28 binds E3-ligases. 1 / 1
| 1

reach
"It is not possible to exclude the possibility that E3-ligases other than FBXW7 or PIRH2 could form a complex with USP28, allowing for USP28 substrate recognition."
| PMC
USP28 affects Dnmt1
| 1
| 1

sparser
"USP28 binds to the UIM of USP28 and inhibits hydrolysis of Ub chains both in vitro and in cells, and these findings suggest that targeting of UIMs may be an alternative means of inhibiting DUBs including USP28, USP37 and potentially others."
| 1

reach
"As an example, the cleavage of USP28 by Caspase-8 inactivates the DUB and is required to overcome the p53-dependent G2/M DNA damage checkpoint [18]."
| PMC
USP28 affects Cysts
| 1
USP28 activates Cysts. 1 / 1
| 1

reach
"Inhibition of deubiquitinase USP28 attenuates cyst growth in autosomal dominant polycystic kidney disease."
USP28 affects Cyclin_E
| 1
USP28 decreases the amount of Cyclin_E. 1 / 1
| 1

reach
"In addition, overexpression of miR-216b suppressed the substrates ' expression of USP28, for example, c-Myc, and miR-216b overexpression also inhibited Cyclin E expression as well as upregulating p27 expression, both of which were the downstream signals of c-Myc."

eidos
"While reports have indicated that a lack of USP28 could reduce colon cancer in mice [ 114 ] and that USP28 could also be a prognostic marker in bladder [ 115 ] and gastric cancers [ 116 ] , a lack of USP28 promoted liver cancer and correlated with a worse survival of patients with invasive ductal breast carcinoma : Mouse xenograft experiments with USP28-lacking breast cancer cells supported these findings [ 117 ] ."
USP28 affects CRC
| 1
| 1

sparser
"As for JUN, it is known as transcription factor AP-1, is involved in activated KRAS-mediated transcriptional activation of USP28 by binding to the USP28 promoter in CRC cells."
USP28 affects COPS5
| 1
| 1

sparser
"All of these except USP25 form a cluster, within which USP28 and COPS5 form a heart-specific subcluster."
USP28 affects CIT
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1 |

No evidence text available
USP28 affects CHEK1
| 1
USP28 inhibits CHEK1. 1 / 1
| 1

reach
"We next examined whether USP28 knock-down (TOV21G cells transfected with si-USP28) reduces the levels of Chk1 protein."
USP28 affects CHEK
| 1
USP28 activates CHEK. 1 / 1
| 1

reach
"This early disassembly of the MRN complex could be prevented by altering the ubiquitination site of Nbs1 or by expressing a deubiquitinase, Usp28, which sufficiently restored homologous recombination repair and ATM, a major checkpoint kinase against DNA DSBs, activation abilities in RTS and RecQL4-depleted cells."
USP28 affects CDKN2A
| 1
USP28 increases the amount of CDKN2A. 1 / 1
| 1

reach
"Notably, USP28 knockdown cells had decreased expression of p53, p21 and p16 INK4a, suggesting that the effect of USP28 on cell proliferation was mediated by regulating the expression of p53, p21 and p16 INK4a."
USP28 affects CDK6
| 1
USP28 inhibits CDK6. 1 / 1
| 1

reach
"As expected, USP28 overexpression further decreased the relative expression of cell cycle related genes (CDK4, CDK6, Cyclin D1, pRB, and E2F1) induced by 5 '-AZA, whereas p15 and p27 expression was significantly enhanced."
USP28 affects CDK4
| 1
USP28 inhibits CDK4. 1 / 1
| 1

reach
"As expected, USP28 overexpression further decreased the relative expression of cell cycle related genes (CDK4, CDK6, Cyclin D1, pRB, and E2F1) induced by 5 '-AZA, whereas p15 and p27 expression was significantly enhanced."
USP28 affects CCNE1
1 |
USP28 deubiquitinates CCNE1. 1 / 1
1 |

". This study pointed towards USP28 as an oncogene, potentially acting via stabilisation of c-Myc and cyclin E1."
USP28 affects CCND1
| 1
USP28 inhibits CCND1. 1 / 1
| 1

eidos
"As shown in Figure S3D , the absence of Usp28 resulted in the decreases of TCF7L2 levels as well as cyclin D1 levels ."
USP28 affects C171
| 1
USP28 binds MYC and C171. 1 / 1
| 1

reach
"Its crystal structure showed that USP28 interacts with MYC through C171 and the MBI domain."
| 1

eidos
"However , the underlying mechanisms of breast cancer suppression by USP28 are not clear ."
USP28 affects BRAF protein
| 1
USP28 inhibits BRAF protein. 1 / 1
| 1

eidos
"Therefore , USP28 depletion increased BRAF protein levels and melanoma cell resistance to BRAF inhibitors [ 57 ] ."
| PMC
USP28 affects BRAF inhibitor
| 1
USP28 inhibits BRAF inhibitor. 1 / 1
| 1

reach
"In an attempt to find a way to overcome USP28 mediated BRAF inhibitor resistance, we performed a synthetic lethal screen using a library of FDA approved small molecules."
USP28 affects BCHE
| 1
Modified USP28 leads to the deubiquitination of BCHE. 1 / 1
| 1

reach
"Overexpression of USP28 inhibited FBW7-depedent ubiquitination of c-MYC and cyclin E1, both of which are oncoproteins."
USP28 affects BAP1
1 |
1 |

No evidence text available
USP28 affects ATXN3L
1 |
1 |

No evidence text available
USP28 affects ATG7
| 1
USP28 increases the amount of ATG7. 1 / 1
| 1

reach
"The results showed that knockdown of ATG7 decreased human usp28 promoter driven reporter transcription activity (XREF_FIG), revealing that ATG7 promotes usp28 promoter transcription."
USP28 affects ASS1
| 1
Modified USP28 increases the amount of ASS1. 1 / 1
| 1

reach
"Overexpression of USP28 in BR cells enhances c-Myc expression and hence increases ASS1 transcription upon arginine deprivation, and consequently leads to cell survival."
USP28 affects AQR
1 |
1 |

No evidence text available
USP20 affects USP28
1 |
1 |

No evidence text available
USP15 affects USP28
1 |
1 |

No evidence text available
USP14 affects USP28
1 |
1 |

No evidence text available
USP affects USP28
| 1
USP28 binds MYC and USP. 1 / 1
| 1

sparser
"Importantly, our results establish that the MYC MBI domain (1–95 amino acids) interacts directly with the USP domain (160–652 amino acids) of USP28."
USO1 affects USP28
1 |
1 |

No evidence text available
| 1

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
UCHL5 affects USP28
1 |
1 |

No evidence text available
UCHL1 affects USP28
1 |
1 |

No evidence text available
TWF1 affects USP28
1 |
1 |

No evidence text available
TTC39A affects USP28
| 1
| 1

sparser
"To further examine the USP28-LSD1 axis in regulating CSC-like properties in human breast cancer, we measured tumorsphere formation in BT549 and MCF7 cells with knockdown of USP28."
TRIM63 affects USP28
1 |
1 |

No evidence text available
TRIM55 affects USP28
1 |
1 |

No evidence text available
TP53BP1 affects MDC1
| 1
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
TP53 affects MYC
| 1

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
Streptoglutarimide H affects USP28
| 1
Streptoglutarimide H inhibits USP28. 1 / 1
| 1

reach
"Streptoglutarimide H (SGH) Downregulated Deubiquitinase USP28 and Cancer Transcription Factor c-Myc."
SYK affects USP28
| 1
| 1

sparser
"In addition, SYK does not bind to USP28 in the same assay ( Fig. 1 B, left panel)."
STAMBP affects USP28
1 |
1 |

No evidence text available
STAM affects USP28
1 |
1 |

No evidence text available
ST6GAL1 affects USP28
1 |
1 |

No evidence text available
SENP8 affects USP28
1 |
1 |

No evidence text available
SENP2 affects USP28
1 |
1 |

No evidence text available
SASS6 affects USP28
| 1
SASS6 inhibits USP28. 1 / 1
| 1

reach
"Importantly, knockout of the essential centriole component SAS6 also triggered a 53BP1, USP28, and p53 dependent G1 arrest, demonstrating that the cell cycle arrest was likely to be caused by centrosome loss and not by the loss of Plk4 activity per se [XREF_BIBR]."
RP23-446O17.1 affects USP28
1 |
USP28 binds RP23-446O17.1. 1 / 1
1 |

No evidence text available
RP23-437L13.2 affects USP28
1 |
USP28 binds RP23-437L13.2. 1 / 1
1 |

No evidence text available
RNAi affects USP28
| 1
RNAi inhibits USP28. 1 / 1
| 1

reach
"Although RNAi mediated knockdown of USP28, USP32 and USP36 showed that their depletion resulted in the increase of ub-K119-H2A, only USP28 depleted cells showed increased cell proliferation."
RFFL affects USP28
1 |
1 |

No evidence text available
RBM24 affects USP28
| 1
| 1

sparser
"The alternative splicing forms of usp28 and abcc9 changed in rbm24 defect mice, causing dilated atria [ xref ]."
RAS affects USP28
| 1
RAS activates USP28. 1 / 1
| 1

reach
"Thus, cJUN is responsible, at least in part, for RAS mediated transcriptional upregulation of USP28."
PR-619 affects USP28
| 1
PR-619 inhibits USP28. 1 / 1
| 1

eidos
"To date , only few small-molecule USP28 inhibitors have been reported , of which PR-619 inhibits USP28 with an EC50 value of about 8 mumol / L13 ."
PLA2G2A affects USP28
1 |
1 |

No evidence text available
PJA1 affects USP28
1 |
1 |

No evidence text available
PIM1/2 affects USP28
| 1
USP28 binds PIM1/2. 1 / 1
| 1

reach
"Interestingly, coimmunoprecipitation assays showed an increased interaction between PIM1/2 and USP28 in response to hypoxia, which correlated with reduced ubiquitination and increased protein stability."
PFAS affects USP28
1 |
1 |

No evidence text available
OTUB1 affects USP28
1 |
1 |

No evidence text available
NUMA1 affects USP28
| 1
| 1

sparser
"Using mass spectrometry and immunoprecipitation, we found that USP28 interacts with NuMA1 and affects centrosome clustering."
NSMCE2 affects USP28
1 |
1 |

No evidence text available
NOTCH1 affects USP28
| 1
NOTCH1 activates USP28. 1 / 1
| 1

reach
"Levels of NICD were only partially rescued by proteasome inhibition, suggesting that additional mechanisms contribute to the downregulation of NICD in Usp28 +/- cells."
MYC affects USP, and USP28
| 1
USP28 binds MYC and USP. 1 / 1
| 1

sparser
"Importantly, our results establish that the MYC MBI domain (1–95 amino acids) interacts directly with the USP domain (160–652 amino acids) of USP28."
MYC affects TP53
| 1

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
MYC affects C171
| 1
USP28 binds MYC and C171. 1 / 1
| 1

reach
"Its crystal structure showed that USP28 interacts with MYC through C171 and the MBI domain."
MTNR1A affects USP28
1 |
1 |

No evidence text available
MRPS7 affects USP28
1 |
1 |

No evidence text available
MIR4295 affects USP28
| 1
| 1

eidos
"In addition , this inhibition of miR-4295 expression not only increased USP28 3 ' - UTR activity but also promoted USP28 and CD44 protein expression ( Fig. 5B & 5C ) ."
MEF2A affects USP28
1 |
MEF2A decreases the amount of USP28. 1 / 1
1 |

No evidence text available
MDC1 affects TP53BP1
| 1
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
MAP3K1 affects USP28
1 |
1 |

No evidence text available
KRAS affects USP28
| 1
KRAS increases the amount of USP28. 1 / 1
| 1

reach
"However, in contrast to the results in KRAS positive CRC cell lines, in WI-38 cells KRAS failed to increase ZNF304 protein levels (XREF_FIG) or significantly stimulate USP28 and PRKD1 transcription (XREF_FIG), explaining at least in part why KRAS expression does not result in INK4-ARF silencing."
JUN affects USP28
| 1
| 1

sparser
"The ChIP results of xref show, consistent with the bioinformatic prediction, that in DLD-1 cells cJUN was bound to the USP28 promoter."
JOSD1 affects USP28
1 |
1 |

No evidence text available
ISO affects USP28
| 1
ISO inhibits USP28. 1 / 1
| 1

eidos
"The current study found that ISO inhibited USP28 expression , thereby promoting CD44 protein degradation and suppressing human BC sphere formation ."
Hemagglutinins affects UCK1, and USP28
| 1

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
HNRNPD affects USP28
| 1
HNRNPD decreases the amount of USP28. 1 / 1
| 1

reach
"As expected, AUF1 knockdown led to increased expression of TET1, USP28, and CD44s protein (XREF_FIG and XREF_FIG), as well as promotion of cancer stem like cell sphere formation (XREF_FIG and XREF_FIG)."
HNF1B affects USP28
| 1
HNF1B activates USP28. 1 / 1
| 1

reach
"Endogenous expression of HNF-1beta in TOV21G cells promotes the upregulation of USP28 at protein levels as determined by Western blot."
HK2 affects USP28
| 1
HK2 activates USP28. 1 / 1
| 1

reach
"Also, high glucose consistently triggered HK2 and LDHA expression and stimulated the USP28 and MYC axis in breast cancer cells (XREF_SUPPLEMENTARY)."
HIF1A affects FBXW7
| 1
| 1

sparser
"In contrast to VDU2 which directly interacts with HIF-1α, USP28 forms a ternary complex with HIF-1α via its association with HIF-1α bound Fbw7 (Flügel et al., xref )."
HIF1 affects USP28
| 1
HIF1 increases the amount of USP28. 1 / 1
| 1

reach
"There were two scenarios that resulted in an increase in ubiquitinated c-Myc levels in A549 cells : the first involved an increased phosphorylation of c-Myc at T58; and the second involved a decrease in USP28 protein levels partially mediated by HIF-1 and HIF-2."
Graphical abstract new affects USP28
| 1
Graphical abstract new inhibits USP28. 1 / 1
| 1

eidos
"Graphical abstract A new series of [ 1,2,3 ] triazolo [ 4,5 - d ] pyrimidine derivatives were identified to inhibit USP28 ."
Fbw7alpha affects USP28
| 1
USP28 binds Fbw7alpha. 1 / 1
| 1

reach
"This decline in total protein resulted in decreased endogenous USP28 binding to Fbw7alpha and c-Myc, which increased the levels of Ub-c-Myc."
FOXK1 affects USP28
1 |
1 |

No evidence text available
FOXC1 affects USP28
| 1
USP28 binds FOXC1. 1 / 1
| 1

sparser
"In conclusion, USP28 enhanced cell viability and aerobic glycolysis of colorectal cancer by stabilizing FOXC1, suggesting that USP28-FOXC1 might be a novel therapeutic avenue for colorectal cancer."
FBXW7 affects HIF1A, and USP28
| 1
| 1

sparser
"In contrast to VDU2 which directly interacts with HIF-1α, USP28 forms a ternary complex with HIF-1α via its association with HIF-1α bound Fbw7 (Flügel et al., xref )."
FBW7alpha affects USP28
| 1
USP28 binds FBW7alpha. 1 / 1
| 1

reach
"The effect is limited to FBW7alpha because Usp28 specifically interacts with FBW7alpha and this interaction mediates complex formation between MYC and Usp28."
ERK affects USP28
| 1
| 1

sparser
"Consistent with this finding, vemurafenib resistance in USP28 knockdown cell lines was associated with sustained ERK phosphorylation ( xref ; and Fig. S5, C and D)."
EPAS1 affects USP28
| 1
EPAS1 decreases the amount of USP28. 1 / 1
| 1

reach
"As shown in XREF_FIG, knockdown of both HIF-1alpha and HIF-2alpha attenuated the decrease of USP28 protein level induced by Ni ions or hypoxia."
E3_Ub_ligase affects TP53
| 1

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
E3-ligases affects USP28
| 1
USP28 binds E3-ligases. 1 / 1
| 1

reach
"It is not possible to exclude the possibility that E3-ligases other than FBXW7 or PIRH2 could form a complex with USP28, allowing for USP28 substrate recognition."
| PMC
Dnmt1 affects USP28
| 1
| 1

sparser
"USP28 binds to the UIM of USP28 and inhibits hydrolysis of Ub chains both in vitro and in cells, and these findings suggest that targeting of UIMs may be an alternative means of inhibiting DUBs including USP28, USP37 and potentially others."

reach
"In response to DNA damage, phosphorylation of the deubiquitinase USP28 by DDR kinases, ATM, and ATR leads to deubiquitination of either GATA4 or its associated factor [XREF_BIBR]."
Cyclin affects USP28
| 1
Cyclin activates USP28. 1 / 1
| 1

reach
"The non-amplified and cyclin E1 hi group had significantly increased USP28, while the amplified and cyclin E1 hi cancers had significantly lower FBXW7 expression consistent with a role for both in stabilizing cyclin E1."
CRC affects USP28
| 1
| 1

sparser
"As for JUN, it is known as transcription factor AP-1, is involved in activated KRAS-mediated transcriptional activation of USP28 by binding to the USP28 promoter in CRC cells."
COPS5 affects USP28
| 1
| 1

sparser
"All of these except USP25 form a cluster, within which USP28 and COPS5 form a heart-specific subcluster."
CLSPN affects TP53BP1
| 1
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
CIT affects USP28
1 |
1 |

No evidence text available
CHEK2 affects TP53
| 1

sparser
"Another potential candidate is UBR5, an E3 ligase that interacts with several reported USP28 substrates such as CHK2, p53, and c-MYC [ xref , xref , xref , xref ]."
| PMC
C171 affects USP28
| 1
USP28 binds MYC and C171. 1 / 1
| 1

reach
"Its crystal structure showed that USP28 interacts with MYC through C171 and the MBI domain."
BRAF affects USP28
| 1
BRAF activates USP28. 1 / 1
| 1

reach
"Collectively, these data demonstrate that BRAF ubiquitination and stability is directly regulated through an interplay between the FBW7 and SCF ubiquitin ligase complex and the deubiquitinating enzyme USP28 leading to enhanced MAPK activity."
BCHE affects USP28
| 1
BCHE activates USP28. 1 / 1
| 1

reach
"The non-amplified and cyclin E1 hi group had significantly increased USP28, while the amplified and cyclin E1 hi cancers had significantly lower FBXW7 expression consistent with a role for both in stabilizing cyclin E1."
BAP1 affects USP28
1 |
1 |

No evidence text available
Air Pollutants, Occupational decreases the amount of USP28. 1 / 1
1 |

No evidence text available
ATXN3L affects USP28
1 |
1 |

No evidence text available
ATM/IR affects USP28
| 1
ATM/IR phosphorylates USP28. 1 / 1
| 1

reach
"ATM/IR Dependent Phosphorylation of USP28."
ASS1 affects USP28
| 1
ASS1 increases the amount of modified USP28. 1 / 1
| 1

reach
"Overexpression of USP28 in BR cells enhances c-Myc expression and hence increases ASS1 transcription upon arginine deprivation, and consequently leads to cell survival."
AQR affects USP28
1 |
1 |

No evidence text available
ABCC9 affects RBM24, and USP28
| 1
| 1

sparser
"The alternative splicing forms of usp28 and abcc9 changed in rbm24 defect mice, causing dilated atria [ xref ]."
6-propyl-2-thiouracil decreases the amount of USP28. 1 / 1
1 |

No evidence text available
3-hydroxyisovaleric acid increases the amount of USP28. 1 / 1
1 |

No evidence text available
1 |

No evidence text available
1,3-dinitrobenzene increases the amount of USP28. 1 / 1
1 |

No evidence text available
| 1
(R)-adrenaline increases the amount of USP28. 1 / 1
| 1

reach
"Together, these findings illustrate that epinephrine enhances USP28 expression via induction of LDHA and metabolic rewiring (XREF_FIG)."
1 |

No evidence text available

No evidence text available