IndraLab

Statements


USP28 affects MYC
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USP28 binds MYC.
10 | 1 29 41
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"XREF_BIBR USP28 was also required for Myc stability, and bound to Myc through an interaction with Fbw7alpha, an F-box protein that was a part of an Skp1-Cullins-F-box (SCF)-type ubiquitin ligase."

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

sparser
"Furthermore, we found that lactate enhanced the interaction of MYC only with USP28 WT ."

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"Past investigations revealed that although many FBW7 domains, including WD40, promoted FBW7’s binding to USP28, the amino terminus of FBW7 contributed to the interaction of FBW7, c-Myc and USP28 [5, 29]."

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"However, in the absence of FBW7, the direct interaction of USP28 and c-Myc was also observed, during which USP28 was more likely to bind to unphosphorylated peptide [31]."

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"Although the ubiquitination of c-Myc induced by lanatoside C can be turned over by USP28, what was further demonstrated is that lanatoside C can attenuate the interaction between USP28 and c-Myc."

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"For example, USP28 binds to Myc and causes the deubiquitination of Myc, resulting in protecting Myc from degradation (XREF_TABLE)."

sparser
"Previous studies have shown that USP28 does not bind to MYC directly but rather binds MYC through interaction with FBW7 ( xref )."

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"In fact, it was subsequently found that USP28 binds to MYC at the same motif recognized by Fbw7, but only when it is unphosphorylated, and promotes deubiquitination of MYC in the absence of Fbw7.104 Moreover, it has been reported that USP28 dissociates from Fbw7α in response to DNA damage, allowing MYC to be degraded by Fbw7α.105 Adding another layer of complexity in regulating MYC, USP28 was reported to control the stability of Fbw7 by antagonizing its auto-ubiquitination and degradation in specific tissues such as lung, pancreas and liver, leading to destabilization of Fbw7 targets and to dose-dependent effects in Usp28 knockout mice."

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"Usp28 recruited Fbw7 at all levels but bound Myc only at the highest level of expression."
Fbw7α binds MYC and USP28. 2 / 2
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"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."

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."
LDHA binds USP28, MYC, and SNAI2. 2 / 2
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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."

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 )."
MYC binds USP28 and USP. 1 / 1
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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.
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USP28 activates MYC. 10 / 41
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"This protein inhibits the development of malignant gliomas by antagonizing USP28-induced c-Myc stability and reducing the half-life of c-Myc."

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"USP28 can counteract the activity of FBW7 and promote c-MYC stability in cancer cells."

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"Yang et al (80) found that circFBXW7 encodes a novel 21 kDa protein, FBXW7-185aa, and that FBXW7-185aa inhibits the formation of malignant glioblastoma through antagonizing USP28-induced c-Myc stability by reducing the half-life of c-Myc."

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"USP28 antagonizes the Ub-dependent degradation of c-myc, as well as c-Jun and Notch, by targeting the Ub ligase Fbw7, thereby stabilizing the HIF-1α transcription factor [66] ."

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"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]."
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"Interestingly, Yang et al. [75] identified a circular RNA encoding circ-FBXW7, whose expression in GBM samples was lower compared to the surrounding normal tissue, being capable of antagonizing the stabilization of the oncoprotein c-MYC initiated by the deubiquitinase USP28; circ-FBXW7 expression was linked to greater survival in patients."

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"Therefore, FBXW-185aa prevents FBXW7alpha induced degradation by antagonizing USP28 induced c-Myc stabilization."

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

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

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"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."
USP28 inhibits MYC.
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USP28 inhibits MYC. 10 / 16
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"Degradation of c-MYC is inhibited by de-ubiquitination, and the ubiquitin-specific proteases USP28 and USP36 increase c-MYC stability by deubiquitination, protecting c-MYC from degradation ."

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"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]."

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"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."

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"Loss of USP28 promotes MYC ubiquitination (45), predicting a diminished MYC-PAF1 interaction and enhanced expression of MYC target genes in USP28-deficient cells."

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"Thus, the peptide reduces the half-life of c-Myc by antagonizing USP28-induced c-Myc stabilization.In triple-negative breast cancer (TNBC), circ-FBXW7 produces the same above-mentioned protein (FBXW7-[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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

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"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)."

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"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."

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"Such as the spanning junction open reading frame in circ-FBXW7 driven by internal ribosome entry site encodes a novel 21-kDa protein, which reduces the half-life of c-Myc by antagonizing USP28-induced c-Myc stabilization in mammals [12]."

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"Simultaneously, targeting USP28 promotes the degradation of c-MYC, resulting in cell cycle arrest and inhibition of DNA repair."
USP28 bound to FBXW7-185aa inhibits MYC. 1 / 1
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"The above evidence collectively elucidates that FBXW7-185aa competitively interacted with USP28 and " freed " FBXW7alpha to degrade c-Myc."
USP28 deubiquitinates MYC.
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USP28 deubiquitinates MYC. 10 / 14
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"USP28 directly deubiquitinates and stabilizes MYC."

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"USP28 monomers may antagonize ubiquitination of MYC by these enzymes, analogously to the mechanism described for Apc/c-mediated turnover of Claspin (79)."

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"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]."

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"In addition, the ubiquitination of c-Myc is inhibited by the deubiquitinase USP28, which might be carried on FBW7 and operate as a c-Myc protein stabiliser [60]."

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"[6693] USP28 also deubiquitinates Myc salvaging it from proteosomal degradation."

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"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."

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"Indeed, USP28 de-ubiquitinates and stabilizes MYC, the USP28/MYC interaction being mediated by the F-box protein, FBW7, which targets MYC for degradation by the proteasome."

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"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]."

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

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"Up to now, it has been reported that USP28 suppresses the ubiquitination and degradation of c-Myc through interaction with FBW7α ( Popov et al., 2007 )."
USP28 increases the amount of MYC.
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Modified USP28 increases the amount of MYC. 4 / 4
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"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."

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"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."

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

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"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. 4 / 4
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"Moreover, USP28 positively regulated the protein level of c-Myc, and c-Myc negatively regulated the radiosensitivity of ECA109 and ECA109R cells."

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"Similarly, USP28 is a DUB that stabilizes the c-Myc transcription factor, a potent oncogene in a wide variety of human cancers, and inhibiting USP28 is expected to reduce the levels of c-Myc, and downregulate its activities (Weisberg et al., 2017; Wrigley et al., 2017; Zhou et al., 2018)."

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"Importantly, USP28 WT remarkably increased protein expression of MYC, whereas USP28 Mut reversed lactate induced MYC and SLUG (XREF_FIG)."

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"USP28 WT but not CA mutant also enhanced levels of c-Myc and mature SREBP1a (Fig. S3c, d), indicating a similar mode of regulation."
USP28 decreases the amount of MYC.
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USP28 decreases the amount of MYC. 3 / 3
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"As shown in Figure 5A, depleting USP28 did cause decreases in c-MYC levels in TNBC cells."

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"In human tumor cell lines, shRNA mediated depletion of Usp28 downregulates Myc levels and attenuates proliferation."

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"Moreover, inactivation of USP28 in established APC min/+ tumours reduced c-Myc protein levels, decreased tumour cell proliferation, and increased tumour cell differentiation and death, resulting in in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 ubiquitinates MYC.
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USP28 leads to the ubiquitination of MYC. 2 / 2
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"Here, we provide evidence that dimerization of USP28 attenuates deubiquitination of MYC and limits recruitment of the elongation factor PAF1c."

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"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 affects FBXW7
1 1 | 70 25
USP28 binds FBXW7.
1 | 18 25
1 | 18 25

sparser
"Loss of the USP28-FBW7 complex."

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"Past investigations revealed that although many FBW7 domains, including WD40, promoted FBW7’s binding to USP28, the amino terminus of FBW7 contributed to the interaction of FBW7, c-Myc and USP28 [5, 29]."

sparser
"Therefore, the loss of the USP28-FBW7 complex could stabilize BRAF to potentiate activating downstream MAPK pathway, promoting the therapeutic resistance to BRAF-targeted therapy."

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 )."

sparser
"Collectively, the loss of USP28-FBW7 complex-mediated BRAF degradation stimulates the intrinsic resistance to MAPK inhibition targeted therapy in melanoma [ xref , xref ]."

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"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)."

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"In case of the Fbw7 and Usp28 complex, we suggest that wiring control of the ligase and its substrates to a single DUB ensures that stabilization of proto-oncoproteins during homeostatic processes occ[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Due to mutations in RAC1, loss of PTEN, NF1, CCND1, USP28-FBW7 complex, COT overexpression, and CCND1 amplification, the BRAF kinase enzyme developed resistance over the commercially available BRAF inhibitors."

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
"Past investigations revealed that although many FBW7 domains, including WD40, promoted FBW7’s binding to USP28, the amino terminus of FBW7 contributed to the interaction of FBW7, c-Myc and USP28 [ xref , xref ]."
USP28 deubiquitinates FBXW7.
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USP28 deubiquitinates FBXW7. 10 / 21
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"USP28 can antagonize the process of FBW7 autoubiquitination, which can promote the degradation of BRAF and then lead to the attenuation of MAPK signaling [16, 115], suggesting that USP28 can suppress cell proliferation."

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

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

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"USP28 deubiquitinates and stabilizes FBW7, allowing the WD40 repeat sequence in FBW7 to bind and degrade substrates containing Cdc4 phosphorylation motifs, and it was found that the Cdc4 phosphorylation structural domains are widely present in human RAF subtypes."

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"However, it has recently emerged that USP28 also directly de-ubiquitinates Fbw7, thereby controlling its protein stability [95] ."

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"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)."

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"For instance, USP7 can promote cancer development by downregulating p53 levels through deubiquitination of mouse double minute 2 (MDM2); lysine 63 deubiquitinase (CYLD), which is frequently mutated in nasopharyngeal carcinoma, inhibits cell invasion and metastasis by interfering with the NF-κB signaling pathway; USP28 is an oncogenic factor that deubiquitinates F-box and WD repeat domain containing 7 (FBXW7), thereby enhancing the stability of RAF family members and inhibiting activation of the MEEK pathway."

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"Indeed, wild-type Usp28, but not the catalytically inactive C171A mutant, promoted Fbw7 deubiquitination in vivo."

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"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]"

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"Usp28 preferentially deubiquitinates Fbw7 and at intermediate levels maintains stable Fbw7 but does not allow excessive Fbw7 substrate stabilization."
USP28 inhibits FBXW7.
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"Loss of the deubiquitinase Usp28 monoallele stabilizes and facilitates Fbw7 mediated substrate degradation, whereas, complete knockout of Usp28 promotes the degradation of Fbw7, leading to the accumulation of Fbw7 substrates."

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

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"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."

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"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."

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"Consequently, half of the normal dose of Usp28 (in the Usp28 +/- cells) is adequate to maintain stable Fbw7 but is not sufficient to antagonize Fbw7 mediated substrate degradation."

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

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"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]."

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"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]"

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"Finally, Fbw7 activity is antagonized by the ubiquitin-specific proteases Usp28 and Usp36, which may be readily amenable to inhibition with small molecules."

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"The Usp28 deubiquitinase antagonizes Fbw7 mediated turnover of multiple oncoproteins, including Myc, Jun, and Notch, and promotes tumorigenesis in the intestine."
USP28 activates FBXW7.
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USP28 activates FBXW7. 10 / 10
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"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]"

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"In several tissues, including lung, liver and pancreas, deletion of Usp28 downregulates Fbw7 and promotes accumulation of its substrates."

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"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]"

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"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]"

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"Knockout of Usp28 triggered Fbw7 degradation and accumulation of Fbw7 substrates, resulting in oncogenic transformation."

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"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."

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"USP28, a deubiquitinating enzyme, causes BRAF degradation by stabilizing FBXW7 [29]."

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

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"Similarly, USP28 prevents FBXW7 self-degradation in chronic lymphocytic leukemia (CLL), leading to elevated levels of Notch1 (Close et al., 2018)."

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"Intriguingly, deletion of Usp28 also suppresses intestinal tumor development in the absence of Fbw7, possibly via Fbw7-independent regulation of substrates like Notch, Jun and Myc [102] ."
USP28 ubiquitinates FBXW7.
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USP28 leads to the ubiquitination of FBXW7. 3 / 3
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"In contrast, complete knockout of Usp28 triggers autocatalytic ubiquitination and degradation of Fbw7, sparing Fbw7 substrates."

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"USP28 deletion is associated with 53.2%-9.7% of breast cancer cases, which reveals that USP28 gene deletion-mediated disruption of FBXW7 auto-ubiquitination might correlate with breast cancer development in certain way, but this remains to be further investigated [51]."

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"Complete knockdown of USP28 in mice triggers the auto-ubiquitination of FBXW7 and leads to substrate accumulation, while maintaining the single allele deletion of USP28 partially restores FBXW7 expression, and substrate protein levels are reduced in all mouse tissues [50]."
USP28 decreases the amount of FBXW7.
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USP28 decreases the amount of FBXW7. 3 / 3
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"Overexpressing USP28 reduced the expression of FBXW7 and suppressed FBXW7-185aa-induced c-Myc destabilization (XREF_FIG H)."

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"Complete knockdown of USP28 in mice triggers the auto-ubiquitination of FBXW7 and leads to substrate accumulation, while maintaining the single allele deletion of USP28 partially restores FBXW7 expression, and substrate protein levels are reduced in all mouse tissues [50]."

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"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."
MYC affects USP28
10 | 1 36 41
MYC binds USP28.
10 | 1 29 41
10 | 1 29 35

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"XREF_BIBR USP28 was also required for Myc stability, and bound to Myc through an interaction with Fbw7alpha, an F-box protein that was a part of an Skp1-Cullins-F-box (SCF)-type ubiquitin ligase."

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

sparser
"Furthermore, we found that lactate enhanced the interaction of MYC only with USP28 WT ."

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"Past investigations revealed that although many FBW7 domains, including WD40, promoted FBW7’s binding to USP28, the amino terminus of FBW7 contributed to the interaction of FBW7, c-Myc and USP28 [5, 29]."

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"However, in the absence of FBW7, the direct interaction of USP28 and c-Myc was also observed, during which USP28 was more likely to bind to unphosphorylated peptide [31]."

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"Although the ubiquitination of c-Myc induced by lanatoside C can be turned over by USP28, what was further demonstrated is that lanatoside C can attenuate the interaction between USP28 and c-Myc."

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"For example, USP28 binds to Myc and causes the deubiquitination of Myc, resulting in protecting Myc from degradation (XREF_TABLE)."

sparser
"Previous studies have shown that USP28 does not bind to MYC directly but rather binds MYC through interaction with FBW7 ( xref )."

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"In fact, it was subsequently found that USP28 binds to MYC at the same motif recognized by Fbw7, but only when it is unphosphorylated, and promotes deubiquitination of MYC in the absence of Fbw7.104 Moreover, it has been reported that USP28 dissociates from Fbw7α in response to DNA damage, allowing MYC to be degraded by Fbw7α.105 Adding another layer of complexity in regulating MYC, USP28 was reported to control the stability of Fbw7 by antagonizing its auto-ubiquitination and degradation in specific tissues such as lung, pancreas and liver, leading to destabilization of Fbw7 targets and to dose-dependent effects in Usp28 knockout mice."

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"Usp28 recruited Fbw7 at all levels but bound Myc only at the highest level of expression."
Fbw7α binds MYC and USP28. 2 / 2
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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."

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."
LDHA binds USP28, MYC, and SNAI2. 2 / 2
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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."

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 )."
MYC binds USP28 and USP. 1 / 1
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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.
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MYC inhibits USP28. 3 / 3
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"To demonstrate that, we next sought to determine if reexpressing c-MYC could rescue USP28 depletion."

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"Indeed, HIF increases MYC phosphorylation at Thr58, triggering MYC ubiquitination, and decreases the de-ubiquitinating enzyme USP28, promoting MYC proteasome dependent degradation [XREF_BIBR]."

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"Here we show that dimerization of USP28 limits deubiquitination of MYC, thereby controlling interactions with PAF1c."
MYC increases the amount of USP28.
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MYC increases the amount of USP28. 2 / 2
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"Potential candidates affected by RSV include oncogenic regulators c-JUN and c-MYC, which activate USP28 transcription (38–40)."

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"Expression of Usp28, but not of its catalytic inactive mutant, increased levels of Myc and its half-life."
MYC activates USP28.
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MYC activates USP28. 2 / 2
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"Furthermore, c-Myc reversed the inhibitory effect of USP28 on the radiosensitivity of EC cells."

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"In colorectal cancer, MYC has been shown to activate USP28, a deubiquitinase that in turn stabilizes MYC, JUN, and NOTCH (Diefenbacher et al. 2014)."
USP28 is modified
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USP28 is phosphorylated.
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USP28 is phosphorylated. 10 / 30
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"We also provided evidence that ATM-mediated phosphorylation of USP28 resulted in its disassociation from KLHL2 and UCK1 destabilization."

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"ATM/IR-Dependent Phosphorylation of USP28."

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"We raised antibodies against the phosphoserine 67 and phosphoserine 714 and used them to confirm that endogenous USP28 is phosphorylated in response to IR ( Figure 7 D)."

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"Furthermore, there appears to be a basal level of USP28 phosphorylation even in the absence of IR treatment for Ser67, suggesting the presence of some constitutive ATM signaling, which could contribut[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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

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

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

sparser
"Analysis of USP28 phosphorylation levels."

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"In brief, unphosphorylated USP28 requires complex formation with another E3-ligase, such as FBXW7, for substrate recognition, whereas phosphorylated USP28 can interact and deubiquitinate substrates independently of E3-ligases ( xref C)."
| PMC

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"The downstream ATM kinase CHK2 could be a potential candidate to phosphorylates USP28 following DNA damage."
| PMC
USP28 is phosphorylated on S67. 5 / 6
1 | 5

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

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"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

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"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 ]."

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"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
USP28 is phosphorylated on S714. 4 / 5
1 | 4

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

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
"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
"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
USP28 is phosphorylated on serine. 2 / 2
| 2

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

sparser
"We found that three serine residues of USP28 were specifically phosphorylated in response to IR, and two of these, serine 67 and serine 714, are SQ sites, which are potential ATM/ATR phosphorylation s[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
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.
| 22
USP28 is sumoylated. 10 / 17
| 17

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

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

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

sparser
"Do other post-translational modifications, such as SUMOylation of USP28 or methylation of 53BP1, or multimerization of USP28 or 53BP1, regulate the MSP ( xref ; xref )?"

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
"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)."

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

sparser
"Our data display that hypoxia modulates the activity of USP28, which leads to the question whether hypoxia also regulates USP28 SUMOylation."

sparser
"N-terminal SUMOylation of USP28 decreases its enzymatic activity."

sparser
"A study determined that SUMOylation of USP28 usually occurs at lysine 99, which is in the UIM domain of the USP28 N-terminus."
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
"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 ]."

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
"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."
USP28 is desumoylated.
| 13
USP28 is desumoylated. 10 / 13
| 13

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

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

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
"Taken together, these data suggested that hypoxia and HIF-1α promoted 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
"SENP1 deSUMOylated and enhanced USP28 deubiquitinating activity towards HIF-1α."

sparser
"In turn, the stabilisation of HIF-1α in hypoxia leads to activation of USP28 through SENP1-mediated USP28 deSUMOylation to further increase transcriptional activity of HIF-1 in human cancer cell lines [ xref , xref ]."

sparser
"SENP1-mediated deSUMOylation of USP28 enhanced its deubiquitinating activity towards HIF-1α, and thereby prevented FBXW7-dependent degradation of HIF-1α during hypoxia."
USP28 is methylated.
| 1 4
USP28 is methylated. 5 / 5
| 1 4

sparser
"Next, the USP28 methylation landscape was also analyzed."

sparser
"These findings indicated that USP28 methylation is significantly associated with mRNA levels in various cancers."

sparser
"Therefore, the identification of aberrations in USP28 methylation patterns may provide a promising avenue for the development of molecular biomarkers for tumors."

sparser
"Subsequently, we further evaluated the influence of USP28 methylation status on prognosis in multiple cancers."

reach
"Our results clearly demonstrate that ATG7 knockdown inhibited usp28 transcription through mediates hypermethylation of usp28 promoter."
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 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
2 1 | 3 47 27
USP28 activates TP53.
| 3 31 4
USP28 activates TP53. 10 / 38
| 3 31 4

sparser
"Multiple studies indicate that P53 is a downstream target protein of USP28, as USP28 can stabilize and activate P53 [ xref , xref , xref , xref – xref ]."

reach
"Understanding how 53BP1 and USP28 elevate p53 in response to centrosome loss and extended mitotic duration, and determining whether centrosome loss is an independent input into the p53 circuit or triggers p53 elevation because it leads to sequential prolonged mitoses, are important future goals arising from the results described here."

reach
"Knockout of 53BP1 or USP28 prevents the stabilization of P53 in cells that experience prolonged mitosis, indicating that 53BP1 and USP28 lie upstream of P53 in this signaling axis (Fong et al. 2016; Lambrus et al. 2016; Meitinger et al. 2016)."

reach
"A genome-wide CRISPR/Cas9 screen of centrinone-treated cells revealed that depletion of TP53BP1, USP28, or TRIM37 inhibits p53 elevation and alleviates proliferation arrest upon centrosome loss [ 41 ][MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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

reach
"USP28 depletion decreased p53 protein, but not p53 transcript, levels in BJ cells undergoing replicative senescence (Fig. 2E; Supplemental Fig. S3E,F)."

reach
"53BP1 responds to mitotic stress, which prolongs mitosis, or to DNA damage and triggers the stabilization of p53 by the deubiquitinase USP28 to stop the proliferation of potentially damaged cells."

reach
"USP28 overexpression increased the levels of multiple senescence markers, including p53, p21 , and GATA4 proteins but not p16, again suggesting that USP28 may be inducing senescence in part through the p53 and GATA4 branches (Fig. 3C–E; Supplemental Fig. S3K,L)."

reach
"XREF_BIBR Intriguingly, 53BP1 and USP28 mediated p53 dependent cell cycle arrest in response to centrosome loss and prolonged mitosis."

reach
"Furthermore, knockout of USP28 prevented P53 up-regulation, consistent with the notion that P53 activation in these microcephaly models occurs through MSP signaling."
USP28 binds TP53.
2 | 3 23
2 | 3 14

sparser
"53BP1 controls p53-dependent responses through direct binding of p53 and USP28 ( xref ) and promotes NHEJ-mediated DSB repair by inhibiting DNA end resection ( xref ; xref )."

sparser
"53BP1-dependent p53 modulation requires both auto-oligomerization and tandem-BRCT domain-mediated bivalent interactions with p53 and the ubiquitin-specific protease 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
"This hypothesis is further supported by biochemical work showing that both p53 and USP28 interact with the tandem-BRCT domains of 53BP1 [ xref – xref ], which are dispensable for the DNA damage response role of 53BP1 [ xref ], but are required for the centrosome surveillance pathway [ xref ]."

sparser
"Even after DNA damage, where USP28 recruitment to 53BP1 foci is visible, a stable MSP complex was not detected and mutations in the BRCT or TTD domains of 53BP1 selectively disrupted the p53 and USP28 interactions, respectively ( xref ; xref )."

sparser
"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 ] ( xref )."

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
"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 ."

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

reach
"The MSP, as well as the transcriptional response to N3 or DSBs (IR or TOP2 inhibition) requires interactions between the 53BP1-TTD and USP28, as well as the 53BP1-BRCT and p53 (Figure 1A)."
| 8

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

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , 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 deubiquitinates TP53.
1 | 10
USP28 deubiquitinates TP53. 10 / 11
1 | 10

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
"However, USP28 also promotes tumour suppressive pathways, for example by deubiquitinating TP53, a function that is counteracted by nuclear caspase 8 during tumour relapse [5]."

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."

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."

reach
"The tumor suppressor p53, which participates in cell cycle arrest, cell apoptosis and DNA repair, has been demonstrated to be deubiquitinated and stabilized by USP28 [102, 103]."

reach
"USP28 can directly deubiquitinate P53 and stabilize mutated P53, enhancing the ability of cancer cells to invade."

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
"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
"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
"They also observed that the deubiquitylase activity of USP28 was necessary to promote p53-dependent responses.Whether USP28 deubiquitylates p53 directly or modifies one of its coterie of binding prote[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 increases the amount of TP53.
| 2
USP28 increases the amount of TP53. 1 / 1
| 1

reach
"Consistent with the lower expression of p53, p21 and p16, USP28-depleted 2BS cells had less blue-colored cells ( Fig. 6 C), and the numbers of senescence-positive cells in USP28 knockdown cells were a[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 increases the amount of mutated TP53. 1 / 1
| 1

reach
"Moreover, we found that USP28 overexpression rescued mutant p53 levels in ΔFBXO42 or ΔCCDC6 RPE1 cells, indicating that USP28 is downstream of FBXO42/CCDC6 (Appendix Fig. 5D)."
USP28 decreases the amount of TP53.
| 1
USP28 decreases the amount of TP53. 1 / 1
| 1

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)."
USP28 affects TP53BP1
7 1 1 | 28 41
USP28 binds TP53BP1.
7 1 | 23 40
7 1 | 23 31

sparser
"Meitinger et al. performed a genome-wide CRISPR/Cas9 screen in RPE-1 cells and identified a 53BP1-USP28 module which induces G1-phase cell cycle arrest after centrosome loss ( xref )."

sparser
"Using this antibody we examined the association of endogenous USP28 and 53BP1 ( Figure 1 C)."

sparser
"As shown in Figures 1 D and 1E, 53BP1 specifically interacts with USP28 but not USP25 or USP7."

sparser
"DNA damage did not dramatically change the interaction between USP28 and 53BP1 ( Figure 1 F)."

reach
"USP28 dimerization is stimulated by 53BP1, which selectively binds USP28 dimers."

reach
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats."

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
"Indeed, work from the Holland, Oegema, and Tsou laboratories has implicated the 53BP1-USP28 module in the p53-dependent response to prolonged mitotic arrest and to numerical centrosomal abnormalities [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"We verified the binding and specificity of the interactions between USP28 and 53BP1 by transiently expressing HA-USP28, HA-USP25, or HA-USP7 in 293T cells."

sparser
"Activation of p53-transcription by TIRR depletion relied on 53BP1-TTD interactions with p53-K382me2 and USP28, however it remains unclear if USP28 interacts with 53BP1 in this context ( xref )."
| 8

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

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
USP28 activates TP53BP1.
| 3
USP28 activates TP53BP1. 3 / 3
| 3

reach
"USP28 mediates centrosome loss induced G1 arrest through its deubiquitinase activity, and acts downstream of 53BP1 to stabilize p53."

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
"However, USP28 loss did not strongly impair DDR signaling or DNA repair, consistent with previous work that found the 53BP1-BRCT domains were not required for the DNA repair functions of 53BP1(Figure 1A) (Knobel et al., 2014; Ward et al., 2006)."
USP28 inhibits TP53BP1.
| 1 1
| 1 1

sparser
"USP28 inhibits 53BP1 and maintains its assembly with PIKKS and its substrate complex, which promotes checkpoint activation and leads to cell cycle arrest [88] ."

reach
"USP28 inhibits 53BP1 and maintains its assembly with PIKKS and its substrate complex, which promotes checkpoint activation and leads to cell cycle arrest [88] ."
USP28 deubiquitinates TP53BP1.
1 | 1
USP28 deubiquitinates TP53BP1. 1 / 2
1 | 1

reach
"USP28 also modulates the DNA damage response (DDR) in cancer cells by deubiquitinating and stabilising the checkpoint kinase 2 (CHK2), the TP53-binding protein 1 (TP53BP1) and Claspin (CLSPN) [6, 7], thereby preventing apoptosis but establishing cell cycle arrest to facilitate DNA repair [8]."
TP53BP1 affects USP28
7 1 | 29 40
TP53BP1 binds USP28.
7 1 | 23 40
7 1 | 23 31

sparser
"Meitinger et al. performed a genome-wide CRISPR/Cas9 screen in RPE-1 cells and identified a 53BP1-USP28 module which induces G1-phase cell cycle arrest after centrosome loss ( xref )."

sparser
"Using this antibody we examined the association of endogenous USP28 and 53BP1 ( Figure 1 C)."

sparser
"As shown in Figures 1 D and 1E, 53BP1 specifically interacts with USP28 but not USP25 or USP7."

sparser
"DNA damage did not dramatically change the interaction between USP28 and 53BP1 ( Figure 1 F)."

reach
"USP28 dimerization is stimulated by 53BP1, which selectively binds USP28 dimers."

reach
"USP28 and p53 both bind to 53BP1 through the tandem C-terminal BRCT repeats."

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
"Indeed, work from the Holland, Oegema, and Tsou laboratories has implicated the 53BP1-USP28 module in the p53-dependent response to prolonged mitotic arrest and to numerical centrosomal abnormalities [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"We verified the binding and specificity of the interactions between USP28 and 53BP1 by transiently expressing HA-USP28, HA-USP25, or HA-USP7 in 293T cells."

sparser
"Activation of p53-transcription by TIRR depletion relied on 53BP1-TTD interactions with p53-K382me2 and USP28, however it remains unclear if USP28 interacts with 53BP1 in this context ( xref )."
| 8

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

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
TP53BP1 activates USP28.
| 3
TP53BP1 activates USP28. 3 / 3
| 3

reach
"We show that USP28 dimerization is stimulated by 53BP1—the major binding partner of USP28 and a key mediator protein in cellular response to DNA damage and replicative stress (41,86–89)."

reach
"Depletion of 53BP1 in p19 Nras cells diminished USP28 dimerization (Figure 5C; Supplementary Figure S5E), mimicking the effect of etoposide."

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."
TP53BP1 inhibits USP28.
| 2
| 2

reach
"Incubation of total cell lysates with the Ub-VME/Ub-VS probes showed that depletion of 53BP1 stimulated USP28 deubiquitinase activity (Figure 5D, Supplementary Figure S5G), whereas the abundance of USP28 was not changed (Supplementary Figure S5H and I)."

reach
"We concluded that depletion of 53BP1 promotes formation of USP28 monomers and stabilizes MYC."
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."

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
"Recently, upregulation of USP28 has been reported to be associated with poor prognosis in NSCLC patients and promote NSCLC cell proliferation XREF_BIBR."

reach
"Although the functions of ΔNp63 are independent of p53, these results indicate that USP28 promotes the cell cycle through its impacts on ΔNp63 and then accelerates cell proliferation."

reach
"Therefore, mediated by LSD1, USP28 can accelerate the cell cycle and promote cell proliferation."

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

reach
"Overexpression of USP28 promoted NSCLC cells proliferation, and was associated with poor prognosis in NSCLC patients."

reach
"Up-regulation of USP28 promoted NSCLC cells proliferation."

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

reach
"This suggests that Usp28 in different cellular contexts is a mediator of proliferation."

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 bound to FBXW7 activates cell population proliferation. 1 / 1
| 1

reach
"USP28 also binds the E3 ubiquitin ligase FBW7 to stabilize MYC and promote cell proliferation [33] ( Fig. 3 a)."

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

reach
"Deubiquitinases have potential deep clinical impacts as is the case of USP28, that when overexpressed, may inhibit AML proliferation and sensitizes AML to the hypomethylating agent azacytidine [ 23 ],[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP28 depletion significantly increased the proliferation rate and strikingly delayed senescence (Fig. 2A–C; Supplemental Fig. S3A)."

reach
"LncRNA-SNHG14/miRNA-3940-5p/NAP12 mRNA may be potential for CRC diagnosis and prognosis.8 miR-3940-5p negatively regulates CUL7 and activates NF-κB to promote gliomagenesis in human gliomas.38 miR-3940-5p targets CCND1 and USP28 to inhibit proliferation and induce apoptosis in NSCLC cells."

reach
"CCK8 and EdU assay showed that si-USP28 reduced PASMCs proliferation ( Fig. 4 A and B)."

reach
"Conversely, USP28 downregulation may significantly inhibit the proliferation and migration of GC cells; however, the effects of USP28 expression on the proliferation and migration of gastric epithelial mucosal cell lines were not significant (46)."

reach
"Functional experiments showed that down-regulation of USP28 expression effectively inhibited the proliferation, migration and invasion of HCC cells, while overexpression of USP28 produced the opposite effect."

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
"Recently, it was shown that loss of USP28 or 53BP1, two upstream components of MSP, can rescue NPC proliferation in microcephaly models with centrosome defects (Phan et al. 2021)."

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)."
FBXW7 affects USP28
1 | 22 25
FBXW7 binds USP28.
1 | 18 25
1 | 18 25

sparser
"Loss of the USP28-FBW7 complex."

reach
"Past investigations revealed that although many FBW7 domains, including WD40, promoted FBW7’s binding to USP28, the amino terminus of FBW7 contributed to the interaction of FBW7, c-Myc and USP28 [5, 29]."

sparser
"Therefore, the loss of the USP28-FBW7 complex could stabilize BRAF to potentiate activating downstream MAPK pathway, promoting the therapeutic resistance to BRAF-targeted therapy."

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 )."

sparser
"Collectively, the loss of USP28-FBW7 complex-mediated BRAF degradation stimulates the intrinsic resistance to MAPK inhibition targeted therapy in melanoma [ 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)."

reach
"In case of the Fbw7 and Usp28 complex, we suggest that wiring control of the ligase and its substrates to a single DUB ensures that stabilization of proto-oncoproteins during homeostatic processes occ[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Due to mutations in RAC1, loss of PTEN, NF1, CCND1, USP28-FBW7 complex, COT overexpression, and CCND1 amplification, the BRAF kinase enzyme developed resistance over the commercially available BRAF inhibitors."

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
"Past investigations revealed that although many FBW7 domains, including WD40, promoted FBW7’s binding to USP28, the amino terminus of FBW7 contributed to the interaction of FBW7, c-Myc and USP28 [ xref , xref ]."
FBXW7 inhibits USP28.
| 2
FBXW7 inhibits USP28. 2 / 2
| 2

reach
"USP28 antagonises FBXW7, resulting in MYC stabilisation [11–13]."

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 activates USP28.
| 2
FBXW7 activates USP28. 2 / 2
| 2

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
"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]"
USP28 affects STAT3
4 1 | 30 12
USP28 binds STAT3.
4 | 7 12
4 | 7 12

sparser
"USP28 interacted with STAT3, and increased the stability of STAT3 by inducing its deubiquitination."

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
"This interaction between STAT3 and USP28 was further confirmed by IP with anti-USP28 antibody and then blotting with anti-STAT3 antibody, wherein STAT3 protein was also found in the USP28 immunoprecip[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"The fact that USP28 interacted with STAT3, and both proteins were upregulated in ADPKD kidneys prompted us to speculate that USP28 may regulate STAT3 as a deubiquitinase."

sparser
"The deubiquitinating enzyme USP28 mediates STAT3 signaling in non-small-cell lung cancer cells, and USP28 interacts with STAT3 and enhances its stability by inducing the deubiquitination of STAT3 [ xref ]."

sparser
"A previous study demonstrated that the deubiquitinating enzyme USP28 directly interacts with STAT3 and increases its stability. xref Thus, regulation of STAT3 acetylation and recruitment of E3 ligase or deubiquitinating enzymes might be associated with the regulation of STAT3 protein stability by SA14."

sparser
"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."

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

sparser
"To evaluate whether USP28 interacted with STAT3, the reciprocal Co-IP was performed."
USP28 activates STAT3.
| 15
USP28 activates STAT3. 10 / 15
| 15

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

reach
"USP28 increases the stability of STAT3."

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
"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
"The deubiquitinating enzyme USP28 mediates STAT3 signaling in non-small-cell lung cancer cells, and USP28 interacts with STAT3 and enhances its stability by inducing the deubiquitination of STAT3 [22]."

reach
"As stated before, USP28 mediated STAT3 signaling by stabilizing STAT3 protein, which suggested that USP28 was functional in NSCLC cells."

reach
"In this study, the deubiquitinating enzyme USP28 was found to mediate STAT3 signaling in NSCLC cells."

reach
"In non-small-cell lung cancer, USP28 appears to mediate STAT3 signaling through deubiquitination and stabilization (21)."

reach
"These results demonstrated that USP28 was functional in NSCLC cells, and promoted NSCLC cell growth by inducing STAT3 signaling."
USP28 increases the amount of STAT3.
| 5
USP28 increases the amount of STAT3. 5 / 5
| 5

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)."

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

reach
"Whole cell lysates were also prepared for immunoblotting, which showed that USP28 upregulated the expression level of STAT3 (XREF_FIG)."
USP28 deubiquitinates STAT3.
1 | 2
USP28 deubiquitinates STAT3. 2 / 3
1 | 2

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

reach
"The deubiquitinating enzyme USP28 mediates STAT3 signaling in non-small-cell lung cancer cells, and USP28 interacts with STAT3 and enhances its stability by inducing the deubiquitination of STAT3 [22]."
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 affects KDM1A
5 1 | 30 11
USP28 binds KDM1A.
5 | 10 11
5 | 10 11

sparser
"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."

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)."

sparser
"Additionally, the level and activity of the histone demethylase lysine specific demethylase 1 (LSD1) can be controlled by USP28 through a deubiquitination event, during which an amine oxidase domain in LSD1 and the N-terminal region of USP28 are responsible for the LSD1 interaction with USP28 [ xref ]."

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

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

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
"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."

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

reach
"In our study, we found that knockdown of Myc did not affect the binding between USP28 and LSD1 (XREF_SUPPLEMENTARY)."

sparser
"USP28 interacts with LSD1."
USP28 deubiquitinates KDM1A.
1 | 6
USP28 deubiquitinates KDM1A. 5 / 6
1 | 5

reach
"USP28 functions as a bona fide deubiquitinating enzyme for LSD1, as it interacts with and stabilizes LSD1 through the process of deubiquitination."

reach
"The results above suggest that USP28 may stimulate EMT via its deubiquitination of LSD1 and Snail and then suppress EMT-related molecules.In addition to the molecules involved in EMT, other molecules, such as Lin28A, which is a crucial regulator of invasiveness, can also accelerate metastasis [84]."

reach
"Therefore, USP28 can maintain Notch signaling, through which cell differentiation is inhibited.Moreover, LSD1, which can be deubiquitinated by USP28, has the capacity to regulate pluripotency markers such as octamer-binding transcription factor 4 (Oct4) and SRY-related HMG box-containing 2 (SOX2) and the differentiation gene p21 [9, 83, 91]."

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 activates KDM1A.
| 6
USP28 activates KDM1A. 6 / 6
| 6

reach
"These findings highlight the critical role of USP28-mediated LSD1 stabilization in maintaining CSC-like properties and driving tumorigenicity.In glioblastoma, the deubiquitinase USP22 facilitates LSD1 stabilization in a glycogen synthase kinase 3β (GSK3β)-dependent manner ."

reach
"Overexpression of HDAC5 significantly increases USP28 expression in cell lines and reduces USP28 polyubiquitination, supporting the idea that HDAC5 stabilizes LSD1 by enhancing its deubiquitinase prot[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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
"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."

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
"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."
USP28 inhibits KDM1A.
| 4
USP28 inhibits KDM1A. 4 / 4
| 4

reach
"In the human gastric cancer cell line HGC-27, the level of USP28 decreases in a dose-dependent manner of compound 19, leading to the degradation of the substrate LSD1 and total c-MYC through the prote[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Overexpression of HDAC5 significantly increases USP28 expression in cell lines and reduces USP28 polyubiquitination, supporting the idea that HDAC5 stabilizes LSD1 by enhancing its deubiquitinase prot[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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)."

reach
"Overexpression of USP28 largely reversed HDAC5-knockdown-induced LSD1 protein degradation, suggesting HDAC5 positively regulates LSD1 by stabilizing USP28."
USP28 increases the amount of KDM1A.
| 3
USP28 increases the amount of KDM1A. 3 / 3
| 3

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)."

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."
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)."
TP53 affects USP28
2 | 5 24
TP53 binds USP28.
2 | 3 23
2 | 3 14

sparser
"53BP1 controls p53-dependent responses through direct binding of p53 and USP28 ( xref ) and promotes NHEJ-mediated DSB repair by inhibiting DNA end resection ( xref ; xref )."

sparser
"53BP1-dependent p53 modulation requires both auto-oligomerization and tandem-BRCT domain-mediated bivalent interactions with p53 and the ubiquitin-specific protease 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
"This hypothesis is further supported by biochemical work showing that both p53 and USP28 interact with the tandem-BRCT domains of 53BP1 [ xref – xref ], which are dispensable for the DNA damage response role of 53BP1 [ xref ], but are required for the centrosome surveillance pathway [ xref ]."

sparser
"Even after DNA damage, where USP28 recruitment to 53BP1 foci is visible, a stable MSP complex was not detected and mutations in the BRCT or TTD domains of 53BP1 selectively disrupted the p53 and USP28 interactions, respectively ( xref ; xref )."

sparser
"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 ] ( xref )."

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
"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 ."

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

reach
"The MSP, as well as the transcriptional response to N3 or DSBs (IR or TOP2 inhibition) requires interactions between the 53BP1-TTD and USP28, as well as the 53BP1-BRCT and p53 (Figure 1A)."
| 8

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

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , 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.
| 2 1
TP53 activates USP28. 3 / 3
| 2 1

reach
"In 2 out of 3 cell line screens, the fitness attenuation effect of TP53-/- background cells was stronger in USP28 than in the neighboring ATM gene, supporting the causal role of USP28 in that segment (Additional file 3: Data S5)."

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."

sparser
"Homozygosity of either Usp28 or 53bp1 loss reduced p53 activation and rescued brain size in both Sas4 and Cep63 microcephaly models ( xref )."
USP28 affects SOX9
| 21 7
USP28 binds SOX9.
| 9 7
| 9 7

reach
"PARPi promotes the interaction between SOX9 and USP28."

sparser
"PARPi promotes the interaction between SOX9 and USP28."

sparser
"The GST pull-down assay was implemented to validate the direct interaction between SOX9 and USP28 (Fig. xref )."

sparser
"The endogenous interaction between USP28 and SOX9 was confirmed in SKOV3 and UWB1.289 cells (Fig. xref )."

sparser
"To explore the impact of PARPi on the interaction between USP28 and SOX9, cells were treated with olaparib for 48 h."

reach
"The GST pull-down assay was implemented to validate the direct interaction between SOX9 and USP28 (Fig. 2A)."

sparser
"In conclusion, these data indicated a direct interaction between USP28 and SOX9, suggesting a potential role of USP28 in the regulation of SOX9 ubiquitination."

reach
"The endogenous interaction between USP28 and SOX9 was confirmed in SKOV3 and UWB1.289 cells (Fig. 2F, G)."

reach
"To explore the impact of PARPi on the interaction between USP28 and SOX9, cells were treated with olaparib for 48 h."

reach
"The Co-IP results showed that olaparib treatment enhances the binding between USP28 and SOX9 (Fig. 2H)."
USP28 increases the amount of SOX9.
| 4
USP28 increases the amount of SOX9. 4 / 4
| 4

reach
"In contrast, the overexpression of USP28 increased both endogenous and exogenous protein levels of SOX9 in a dose-dependent manner (Fig. 3B and Fig. S3B, C)."

reach
"However, the catalytically inactive form (USP28 ) or the ΔUSP truncation mutant of USP28 (USP28 ) failed to enhance the expression of SOX9 (Fig. 3C)."

reach
"The pharmacological inhibitor of USP28, AZ1, decreased the protein levels of USP28 itself and that of SOX9 in a dose-dependent manner (Fig. 3J)."

reach
"Firstly, we investigated whether USP28 can modulate the expression level of SOX9."
USP28 activates SOX9.
| 4
USP28 activates SOX9. 4 / 4
| 4

reach
"Collectively, our results revealed that USP28 prevents SOX9 proteasomal degradation by eliminating K48-linked polyubiquitin chains."

reach
"Knockdown of USP28 reduced the binding of SOX9 to the promoter regions of SMARCA4, UIMC1, and SLX4 (Fig. S9A)."

reach
"Additionally, USP28 promoted olaparib resistance by stabilizing SOX9 protein and enhancing DNA damage repair."

reach
"Furthermore, the USP28 specific inhibitor AZ1 reduced SOX9 protein stability and increased the sensitivity of ovarian cancer cells to olaparib."
USP28 ubiquitinates SOX9.
| 2
USP28 leads to the ubiquitination of SOX9. 2 / 2
| 2

reach
"In addition, USP28 specific inhibitor AZ1 dose-dependently increased the ubiquitination of SOX9 and mitigated the decrease in SOX9 ubiquitination induced by olaparib treatment (Fig. 4F, G)."

reach
"USP28 mediates the ubiquitination of SOX9 as a deubiquitinase."
USP28 deubiquitinates SOX9.
| 2
USP28 deubiquitinates SOX9. 2 / 2
| 2

reach
"Overexpression of WT USP28, but not its catalytically inactive form (USP28 ) or the ΔUSP truncation mutant (USP28 ), induced SOX9 deubiquitination and inhibited its degradation."

reach
"In contrast, overexpression of USP28, but not its inactive form or ΔUSP truncation, led to decreased ubiquitination of SOX9 (Fig. 4B), suggesting that the catalytic activity of USP28 is essential for SOX9 deubiquitination."
USP28 affects UCK1
1 4 1 | 12 9
USP28 binds UCK1.
4 | 6 8
4 | 6 4

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

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

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

reach
"USP28 binds to UCK1 via KLHL2."

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

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

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."

sparser
"Although KLHL2 and UCK1 formed binary complexes, USP28 did not directly bind to UCK1."
KLHL2 binds UCK1 and USP28. 3 / 3
| 3

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

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
"USP28 binds to UCK1 via KLHL2."

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
USP28 deubiquitinates UCK1.
1 1 | 4
USP28 deubiquitinates UCK1. 3 / 4
1 | 3

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."

reach
"We then evaluated whether USP28 could deubiquitinate UCK1."
USP28 leads to the deubiquitination of UCK1 on K81. 1 / 2
1 | 1

reach
"Additionally, USP28 can reverse the polyubiquitination of uridine cytidine kinase 1 (UCK1) at K81 catalyzed by KLHL2, which is required for the link between USP28 and KLHL2 [27]."
USP28 ubiquitinates UCK1.
| 1 1
USP28 leads to the ubiquitination of UCK1. 2 / 2
| 1 1

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."

reach
"We next examined the effect of USP28 on KLHL2 mediated ubiquitination 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."
KDM1A affects USP28
5 | 10 11
5 | 10 11

sparser
"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."

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)."

sparser
"Additionally, the level and activity of the histone demethylase lysine specific demethylase 1 (LSD1) can be controlled by USP28 through a deubiquitination event, during which an amine oxidase domain in LSD1 and the N-terminal region of USP28 are responsible for the LSD1 interaction with USP28 [ xref ]."

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

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

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
"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."

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

reach
"In our study, we found that knockdown of Myc did not affect the binding between USP28 and LSD1 (XREF_SUPPLEMENTARY)."

sparser
"USP28 interacts with LSD1."
ATM affects USP28
1 3 1 | 11 9 1
ATM phosphorylates USP28.
1 2 | 8 8 1
ATM phosphorylates USP28. 10 / 15
1 | 5 8 1

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
"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]."

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

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

sparser
"USP28 is required for p53 and p21 accumulation following p53 activation ( xref E–G) and is phosphorylated on at least three sites by ATM in response to IR ( xref )."

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

reach
"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, suggesting that in response to 5 '-AZA-induced stress, ATM phosphorylates USP28 at all these three sites."

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."
ATM phosphorylates USP28 on S714. 1 / 3
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 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 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 activates USP28.
1 | 2 1
ATM activates USP28. 3 / 4
1 | 2 1

reach
"In line with this model, our data indicate that ATM-dependent activation of USP28 stimulates origin firing early after genotoxic stress."

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."

sparser
"In line with this model, our data indicate that ATM-dependent activation of USP28 stimulates origin firing early after genotoxic stress."
ATM binds USP28.
1 | 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
USP28 affects CLASP1
| 17 7
USP28 binds CLASP1.
| 4 7
| 4 7

reach
"The interaction between USP28 and MAST1 was observed using a Duolink in situ proximity ligation assay (PLA) kit (Cat."

sparser
"The in situ USP28-MAST1 interaction was observed as red signals (PLA dots) when USP28 and MAST1 were immunostained together, but not when they were stained with individual antibodies (Fig.  xref C)."

sparser
"On the basis of the above observations, we hypothesized that the interaction between USP28 and MAST1 might influence on MAST1 protein turnover."

reach
"We further used Duolink PLA assays to validate the interaction between USP28 and MAST1 under physiological conditions."

reach
"To delineate the molecular mechanism by which USP28 regulates MAST1 protein, we first examined the interaction between USP28 and MAST1 under physiological conditions."

reach
"On the basis of the above observations, we hypothesized that the interaction between USP28 and MAST1 might influence on MAST1 protein turnover."

sparser
"The high mRNA expression score of USP28 was proportional with MAST1 expression (Fig.  xref C; Supplementary Table S5) and showed a positive correlation between USP28-MAST1 exhibiting r value of 0.36 (Fig.  xref D)."

sparser
"We envision that targeting the USP28-MAST1 axis along with cisplatin treatment might be an alternative therapeutic strategy to overcome cisplatin resistance in cancer patients."

sparser
"To delineate the molecular mechanism by which USP28 regulates MAST1 protein, we first examined the interaction between USP28 and MAST1 under physiological conditions."

sparser
"These findings confirm that USP28 interacts with MAST1 protein at both endogenous and exogenous levels."
USP28 inhibits CLASP1.
| 5
USP28 inhibits CLASP1. 5 / 5
| 5

reach
"Likewise, Annexin-V-positive cells were increased in USP28-KO clones compared with USP28-depleted cells reconstituted with USP28 or MAST1 (Fig. 5G), indicating that loss of USP28 sensitizes cells to cisplatin-mediated DNA damage and induced cell death.Finally, we wished to investigate the molecular mechanism of USP28 regulation on MAST1-driven cisplatin-resistance."

reach
"An anchorage-independent colony formation assay demonstrated that USP28-KO clones showed reduced colony numbers compared with the mock control, while the colony-forming ability was increased in USP28-depleted cells reconstituted with USP28 or MAST1 (Fig. 6A and Supplementary Fig. 15A)."

reach
"Similarly, cellular invasion and migration rate were significantly hindered in USP28-KO clones, whereas USP28-depleted cells reconstituted with USP28 or MAST1 showed reversed results (Fig. 6B, C and Supplementary Fig. 15B, C)."

reach
"Next, we measured the ability of USP28-depleted cells to impair tumor growth in NSG mice by subcutaneously injecting USP28-KO cells and USP28-depleted cells reconstituted with either USP28 or MAST1 obtained from Fig. 5A."

reach
"However, mice bearing USP28-depleted cells reconstituted with USP28 or MAST1 displayed increased tumor size, volume and weight (Fig. 6D, E, Supplementary Table S6)."
USP28 increases the amount of CLASP1.
| 4
USP28 increases the amount of CLASP1. 4 / 4
| 4

reach
"USP28 positively regulates MAST1 protein levels."

reach
"Together, these results suggest that USP28 positively regulates MAST1 protein levels."

reach
"Together, these results indicate that loss of USP28 decreased MAST1 protein level and suppressed tumor growth upon cisplatin treatment."

reach
"Furthermore, we showed that the loss of USP28 downregulates MAST1 protein level and sensitizes cells for cisplatin toxicity during tumor growth, leading to reduced tumor size, volume and weight.In conclusion, our study demonstrates that USP28 could enhance MAST1-driven cisplatin resistance by stabilizing MAST1 protein level in cancer."
USP28 deubiquitinates CLASP1.
| 3
USP28 deubiquitinates CLASP1. 3 / 3
| 3

reach
"USP28 promotes tumorigenesis and cisplatin resistance by deubiquitinating MAST1 protein in cancer cells."

reach
"USP28 deubiquitinates MAST1 protein and extends its half-life."

reach
"USP28 deubiquitinates MAST1."
USP28 activates CLASP1.
| 1
USP28 activates CLASP1. 1 / 1
| 1

reach
"Additionally, CRISPR/Cas9-mediated gene knockout of USP28 in A549 and NCI-H1299 cells blocked MAST1-driven cisplatin resistance, resulting in suppressed cell proliferation, colony formation ability, migration and invasion in vitro."
STAT3 affects USP28
4 | 7 12
4 | 7 12

sparser
"USP28 interacted with STAT3, and increased the stability of STAT3 by inducing its deubiquitination."

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
"This interaction between STAT3 and USP28 was further confirmed by IP with anti-USP28 antibody and then blotting with anti-STAT3 antibody, wherein STAT3 protein was also found in the USP28 immunoprecip[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"The fact that USP28 interacted with STAT3, and both proteins were upregulated in ADPKD kidneys prompted us to speculate that USP28 may regulate STAT3 as a deubiquitinase."

sparser
"The deubiquitinating enzyme USP28 mediates STAT3 signaling in non-small-cell lung cancer cells, and USP28 interacts with STAT3 and enhances its stability by inducing the deubiquitination of STAT3 [ xref ]."

sparser
"A previous study demonstrated that the deubiquitinating enzyme USP28 directly interacts with STAT3 and increases its stability. xref Thus, regulation of STAT3 acetylation and recruitment of E3 ligase or deubiquitinating enzymes might be associated with the regulation of STAT3 protein stability by SA14."

sparser
"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."

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

sparser
"To evaluate whether USP28 interacted with STAT3, the reciprocal Co-IP was performed."
USP28 affects Neoplasms
| 1 20
USP28 activates Neoplasms.
| 19
| 19

reach
"Together, these results indicate that loss of USP28 decreased MAST1 protein level and suppressed tumor growth upon cisplatin treatment."

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

reach
"Many studies have demonstrated that USP28 can promote the deubiquitination and stabilization of many oncoproteins, implying that USP28 can stimulate the progression of many tumors."

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

reach
"The enzyme has been found to be overexpressed in bladder cancer, glioma, gastric cancer, etc. [12] Besides, multiple studies have shown that the knockdown of USP28 or its inhibition by chemical inhibi[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP28 can induce tumor angiogenesis directly and indirectly through the deubiquitination of HIF-1α and c-Myc."

reach
"The tumor-promoting role of USP28 and relevant investigations are currently dominant; nevertheless, several studies have demonstrated its oncostatic effects."

reach
"USP28 plays a crucial role in tumor progression, and inhibiting USP28 can suppress the growth of various tumors [1]."

reach
"Intriguingly, deletion of Usp28 also suppresses intestinal tumor development in the absence of Fbw7, possibly via Fbw7-independent regulation of substrates like Notch, Jun and Myc [102] ."
USP28 inhibits Neoplasms.
| 1 1
| 1 1

reach
"Depletion of USP28 reduced tumor size and increased the lifespan of tumor-bearing mice.The chromatin modulator LSD1 controls cellular pluripotency through histone demethylation and is overexpressed in[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

eidos
"Moreover , we concentrate on the impacts of USP28 on diverse hallmarks of cancer and discuss whether USP28 can accelerate or inhibit tumor progression ."
| 19
| 11

reach
"Remarkably, both apoptosis and microcephaly were rescued by the removal of 53bp1, Usp28 or p53 in Sas‐4 conditional mutants [69, 141]."

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

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

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."

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

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

reach
"LncRNA-SNHG14/miRNA-3940-5p/NAP12 mRNA may be potential for CRC diagnosis and prognosis.8 miR-3940-5p negatively regulates CUL7 and activates NF-κB to promote gliomagenesis in human gliomas.38 miR-3940-5p targets CCND1 and USP28 to inhibit proliferation and induce apoptosis in NSCLC cells."

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)."
| 8

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
"Moreover, cell apoptosis is triggered by a reduction in USP28 [192]."

reach
"However, some studies have indicated that USP28 deficiency leads to apoptosis [42]."

reach
"miR-3940-5p can target CCND1 and USP28 to inhibit the growth of NSCLC cells, and overexpression of CCND1 and USP28 can attenuate miR-3940-5P-induced proliferation and apoptosis of NSCLC cells (88)."

reach
"The specific mechanism by which USP28 deficiency induces apoptosis is still unclear."

reach
"Loss of USP28, renders cells resistant to IR-induced apoptosis, similar to what has been seen with Chk2, p53 and PUMA null mice."

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 SREBF2
2 | 10 7
USP28 binds SREBF2.
2 | 4 7
2 | 4 7

sparser
"However, it is possible that the interaction between SREBP2 and USP28 is indirect and involves additional, unknown proteins."

sparser
"It has also been demonstrated that USP28 binds to mature SREBP2, leading to its deubiquitination and stabilization."

sparser
"While both proteins localise to and interact in the nucleus, we did observe some level of interaction of SREBP2 and USP28 also in the cytoplasm."

sparser
"Moreover, mature SREBP2 binds to USP28, leading to deubiquitination and stabilization, as well as regulation of the mevalonate pathway (MVP), which drives the synthesis of cholesterol, thus driving tu[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Notably, it is unclear whether the interaction between SREBP2 and USP28 is direct or whether it involves additional proteins."

reach
"However, it is possible that the interaction between SREBP2 and USP28 is indirect and involves additional, unknown proteins."

reach
"While both proteins localise to and interact in the nucleus, we did observe some level of interaction of SREBP2 and USP28 also in the cytoplasm."

reach
"Notably, it is unclear whether the interaction between SREBP2 and USP28 is direct or whether it involves additional proteins.USP28 is highly upregulated in LSCC, where it stabilises the transcription factor ΔNP63 that is essential for the maintenance of the squamous lineage [16]."

reach
"Moreover, mature SREBP2 binds to USP28, leading to deubiquitination and stabilization, as well as regulation of the mevalonate pathway (MVP), which drives the synthesis of cholesterol, thus driving tu[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"We also show that USP28 binds to mature SREBP2, leading to its deubiquitination and stabilisation."
USP28 increases the amount of SREBF2.
| 2
USP28 increases the amount of SREBF2. 2 / 2
| 2

reach
"Deletion of USP28 accelerated the loss of both full-length and mature SREBP2 compared to wild-type cells (Fig. 3a, b)."

reach
"This showed that wild-type USP28 increases the levels of mature SREBP2, which was not observed with the CA mutant (Fig. 3d)."
USP28 deubiquitinates SREBF2.
| 2
USP28 deubiquitinates SREBF2. 2 / 2
| 2

reach
"USP28 deubiquitinates and stabilises SREBP2."

reach
"Furthermore, USP28 WT but not CA mutant decreased levels of mature SREBP2 in a pull-down assay using 6His-tagged K48-only ubiquitin (Fig. 3e), while silencing of USP28 increased the amount of SREBP2 ubiquitination detected by immunoprecipitation in A431 cells (Fig. 3f)."
USP28 decreases the amount of SREBF2.
| 1
USP28 decreases the amount of SREBF2. 1 / 1
| 1

reach
"Furthermore, USP28 WT but not CA mutant decreased levels of mature SREBP2 in a pull-down assay using 6His-tagged K48-only ubiquitin (Fig. 3e), while silencing of USP28 increased the amount of SREBP2 ubiquitination detected by immunoprecipitation in A431 cells (Fig. 3f)."
USP28 activates SREBF2.
| 1
USP28 activates SREBF2. 1 / 1
| 1

reach
"Furthermore, silencing of USP28 in A431 prevented the stabilisation of mature SREBP2 following proteasome inhibition by MG-132 (Fig. 3c)."
USP28 affects CLSPN
1 1 1 | 15 1
USP28 deubiquitinates CLSPN.
1 | 7
USP28 deubiquitinates CLSPN. 7 / 8
1 | 7

reach
"USP28 also modulates the DNA damage response (DDR) in cancer cells by deubiquitinating and stabilising the checkpoint kinase 2 (CHK2), the TP53-binding protein 1 (TP53BP1) and Claspin (CLSPN) [6, 7], thereby preventing apoptosis but establishing cell cycle arrest to facilitate DNA repair [8]."

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

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

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

reach
"However, it must be noted that USP28 can also deubiquitinate and stabilise other substrates not known to be associated with Fbw7, including Claspin [92] , Chk2 [93] , and LSD1 [94] , which could also [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, Claspin, which was discovered as a protein required for the activation of checkpoint kinase 1 (Chk1), can be deubiquitinated by USP28 through reversing the effect of ubiquitin ligase APC/C [67, 68]."

reach
"Thus, deubiquitination of Claspin caused by USP28 contributes to cell cycle arrest, which maintains ovarian clear cell carcinoma cell viability in response to a genotoxic stress [68]."
USP28 binds CLSPN.
1 1 | 2 1
1 1 | 2

reach
"USP28 phosphorylation regulates the complex formation of DNA checkpoint proteins and binds to checkpoint proteins 53BP1, Claspin, and MDC1, resulting in repairing DNA damage."

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
USP28 activates CLSPN.
| 5
USP28 activates CLSPN. 5 / 5
| 5

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."

reach
"USP28 has been reported to promote the proliferation of both tumor and normal cells by stabilizing Claspin [41]."

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

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

reach
"Additionally, in this system, USP28 only targets Claspin but not PLK1 (both of which are subject to ubiquitination by the same E3 ligase)."
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."
USP28 affects Flag
| 18
| 18

sparser
"We used USP28-KO clones exhibiting reduced MAST1 protein level and the cells were reconstituted either with Flag-USP28 or Myc-MAST1 for further functional experiments (Fig.  xref A; Supplementary Fig. 8)."

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

sparser
"To this end, we transfected both A549 and HEK293 cells with increasing concentrations of Flag-USP28 and analyzed MAST1 protein level."

sparser
"To examine the deubiquitinated modification of TRIM21 by USP28, we evaluated the ubiquitination level of TRIM21 in HL-1 cells expressing Flag-USP28 (USP28 OE ) and found that USP28 OE promoted the deubiquitination of TRIM21 (Fig. xref D)."

sparser
"However, the half-life of MAST1 protein was rescued when USP28-KO cells were reconstituted with Flag-USP28 (Fig.  xref J, lane 9–12; Supplementary Fig. 4A, lane 9–12), while USP28CA failed to increase the half-life of MAST1 protein (Fig.  xref K, lane 9–12; Supplementary Fig. 4B, lane 9–12)."

sparser
"Here, we performed a transcriptome in Ang II-challenged HL-1 cells expressing Flag-USP28 (USP28 OE ) and showed that USP28 OE negatively regulates antioxidant activity and biological oxidations pathways (Fig. xref A and xref A-B)."

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

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
"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)."
UCK1 affects USP28
4 | 6 8
4 | 6 4

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

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

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

reach
"USP28 binds to UCK1 via KLHL2."

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

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

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."

sparser
"Although KLHL2 and UCK1 formed binary complexes, USP28 did not directly bind to UCK1."
KLHL2 binds UCK1 and USP28. 3 / 3
| 3

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

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
"USP28 binds to UCK1 via KLHL2."

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
Flag affects USP28
| 18
| 18

sparser
"We used USP28-KO clones exhibiting reduced MAST1 protein level and the cells were reconstituted either with Flag-USP28 or Myc-MAST1 for further functional experiments (Fig.  xref A; Supplementary Fig. 8)."

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

sparser
"To this end, we transfected both A549 and HEK293 cells with increasing concentrations of Flag-USP28 and analyzed MAST1 protein level."

sparser
"To examine the deubiquitinated modification of TRIM21 by USP28, we evaluated the ubiquitination level of TRIM21 in HL-1 cells expressing Flag-USP28 (USP28 OE ) and found that USP28 OE promoted the deubiquitination of TRIM21 (Fig. xref D)."

sparser
"However, the half-life of MAST1 protein was rescued when USP28-KO cells were reconstituted with Flag-USP28 (Fig.  xref J, lane 9–12; Supplementary Fig. 4A, lane 9–12), while USP28CA failed to increase the half-life of MAST1 protein (Fig.  xref K, lane 9–12; Supplementary Fig. 4B, lane 9–12)."

sparser
"Here, we performed a transcriptome in Ang II-challenged HL-1 cells expressing Flag-USP28 (USP28 OE ) and showed that USP28 OE negatively regulates antioxidant activity and biological oxidations pathways (Fig. xref A and xref A-B)."

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

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
"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)."
USP28 affects SNAI1
| 12 5
USP28 binds SNAI1.
| 3 5
| 3 5

sparser
"Such blockade was found to inhibit the interaction of USP28 with Snail1."

sparser
"This revealed that USP28 binds to and stabilizes Snail1 via deubiquitination, and these effects are dependent on GSK-3β-mediated phosphorylation."

sparser
"Subsequently, reciprocal co-immunoprecipitation assays confirmed that USP28 interacted with Snail1 in BC cells ( xref H)."

sparser
"LiCl (40 μM) treatment greatly weakened the mutual interaction between USP28 and Snail1 ( xref A)."

sparser
"These results demonstrated that the phosphorylation of the SRD domain was essential for the interaction of USP28 with Snail1."

reach
"LiCl (40 μM) treatment greatly weakened the mutual interaction between USP28 and Snail1 (Fig. 6A)."

reach
"This revealed that USP28 binds to and stabilizes Snail1 via deubiquitination, and these effects are dependent on GSK-3β-mediated phosphorylation."

reach
"Subsequently, USP28 interacted with and deubiquitinated Snail1; these effects were dependent on GSK-3beta-mediated phosphorylation."
USP28 deubiquitinates SNAI1.
| 5
USP28 deubiquitinates SNAI1. 5 / 5
| 5

reach
"Subsequently, USP28 interacted with and deubiquitinated Snail1; these effects were dependent on GSK-3beta-mediated phosphorylation."

reach
"Moreover, USP28 mediated the Pard3 silencing-induced deubiquitination and stabilization of Snail1, thereby promoting BC migration and invasion."

reach
"As expected, ectopic expression of wild-type USP28 reduced the polyubiquitination of Snail1 in a dose dependent manner (Fig. 5E)."

reach
"As Fig. 5F shown, wild-type USP28 instead of the DUB dead mutant USP28C 1A reduced the polyubiquitination of Snail1, whereas USP28 silencing increased Snail1 polyubiquitination."

reach
"The results above suggest that USP28 may stimulate EMT via its deubiquitination of LSD1 and Snail and then suppress EMT-related molecules.In addition to the molecules involved in EMT, other molecules, such as Lin28A, which is a crucial regulator of invasiveness, can also accelerate metastasis [84]."
USP28 ubiquitinates SNAI1.
| 2
USP28 leads to the ubiquitination of SNAI1. 2 / 2
| 2

reach
"As Fig. 5F shown, wild-type USP28 instead of the DUB dead mutant USP28C 1A reduced the polyubiquitination of Snail1, whereas USP28 silencing increased Snail1 polyubiquitination."

reach
"Moreover, USP28 mediated the Pard3 silencing-induced deubiquitination and stabilization of Snail1, thereby promoting BC migration and invasion."
USP28 decreases the amount of SNAI1.
| 2
USP28 decreases the amount of SNAI1. 2 / 2
| 2

reach
"USP28 knockdown dramatically decreased the protein levels of Snail1, whereas USP28 overexpression had the opposite effect (Fig. 5A and B)."

reach
"The expression of EMT-related markers increased, while the expression of N-cad, FN1, ZEB1, snail, and slug was reduced by binding to ubiquitin-specific peptidase 28."
SOX9 affects USP28
| 9 7
| 9 7

reach
"PARPi promotes the interaction between SOX9 and USP28."

sparser
"PARPi promotes the interaction between SOX9 and USP28."

sparser
"The GST pull-down assay was implemented to validate the direct interaction between SOX9 and USP28 (Fig. xref )."

sparser
"The endogenous interaction between USP28 and SOX9 was confirmed in SKOV3 and UWB1.289 cells (Fig. xref )."

sparser
"To explore the impact of PARPi on the interaction between USP28 and SOX9, cells were treated with olaparib for 48 h."

reach
"The GST pull-down assay was implemented to validate the direct interaction between SOX9 and USP28 (Fig. 2A)."

sparser
"In conclusion, these data indicated a direct interaction between USP28 and SOX9, suggesting a potential role of USP28 in the regulation of SOX9 ubiquitination."

reach
"The endogenous interaction between USP28 and SOX9 was confirmed in SKOV3 and UWB1.289 cells (Fig. 2F, G)."

reach
"To explore the impact of PARPi on the interaction between USP28 and SOX9, cells were treated with olaparib for 48 h."

reach
"The Co-IP results showed that olaparib treatment enhances the binding between USP28 and SOX9 (Fig. 2H)."
USP28 affects Ubiquitin
| 11 4
USP28 activates Ubiquitin.
| 8
| 8

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

reach
"USP28 could modulate the lifespan of c-Myc, c-JUN, Notch1, and ΔNp63 through erasing ubiquitin modifications from them [ 8 , 9 ]."

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

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
"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
"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
"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 antagonizes the Ub-dependent degradation of c-myc, as well as c-Jun and Notch, by targeting the Ub ligase Fbw7, thereby stabilizing the HIF-1α transcription factor [66] ."
USP28 binds Ubiquitin.
| 3 4
| 3 4

reach
"For the structure of USP34-Ub-PA complex, the structures of unbound USP34 and ubiquitin (derived from the structure of USP28 covalently bound to Ub, PDB ID: 6H4H) 35 were used as search models in mole[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"In the nucleus, after caspase-8-mediated cleavage at specific sites on USP28, the 30 kDa fragment encoding the UBA and UIM domains is lost, thus affecting the binding of USP28 to ubiquitin and leading[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"A structure of human USP28 bound to Ubiquitin-PA was downloaded from PDB (6HEK) ( Gersch, Wagstaff et al. 2019 ) and visualized in Discovery Studio to identify the Ub bound interface residues on USP28[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"We have also highlighted the region that USP28 binds with Ub."

sparser
"Ub binding to USP28 is similar to other USPs such as USP7/HAUSP and USP21 ( Ernst et al., 2013; Hu et al., 2002 )."

sparser
"For the structure of USP34-Ub-PA complex, the structures of unbound USP34 and ubiquitin (derived from the structure of USP28 covalently bound to Ub, PDB ID: 6H4H) 35 were used as search models in mole[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"We have also highlighted the region that USP28 binds with Ub."
SENP1 affects USP28
2 | 4 9
SENP1 binds USP28.
2 | 1 6
2 | 1 6

sparser
"SENP1 directly interacts with USP28."

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

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

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

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.
| 3 3
SENP1 desumoylates USP28. 5 / 5
| 2 3

sparser
"SENP1 strongly deSUMOylates USP28 and stabilizes the hypoxia-inducible factor 1α (HIF-1α) during hypoxia [41] ( Fig. 3 b)."

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

reach
"SENP1 strongly desumoylates USP28 upon hypoxia [14]."
| PMC

reach
"SENP1 strongly deSUMOylates USP28 and stabilizes the hypoxia-inducible factor 1α (HIF-1α) during hypoxia [41] ( Fig. 3 b)."

sparser
"SENP1 deSUMOylated and enhanced USP28 deubiquitinating activity towards HIF-1α."
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 cell growth
| 13 1
USP28 activates cell growth.
| 8
| 8

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

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

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

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

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
"Overexpression of USP28 accelerated PC cell growth, whereas USP28 knockdown impaired PC cell growth both in vitro and in vivo."
USP28 inhibits cell growth.
| 5 1
| 5 1

reach
"Knockdown of USP28 inhibits NSCLC cell growth."

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

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

reach
"Furthermore, USP28 is shown to regulate cell proliferation in U2OS cells, because depletion of USP28 promotes cell growth and DNA replication."

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
"Knockdown of USP28 suppressed the cell growth of NSCLC both in vitro and in vivo."
USP28 affects cell cycle
| 14
USP28 activates cell cycle.
| 11
| 11

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

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
"Therefore, mediated by LSD1, USP28 can accelerate the cell cycle and promote cell proliferation."

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

reach
"USP28 and SPINT2 mediate cell cycle arrest after whole genomedoubling."
| DOI

reach
"In addition to promoting the cell cycle and enhancing aerobic glycolysis, USP28 can also stimulate cell proliferation."

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."

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
"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
"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)."
USP28 inhibits cell cycle.
| 3
| 3

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
"These extended mitosis trigger the activation of 53BP1 and USP28 which eventually blocks the cell cycle in G1 through a p53‐ and p21‐dependent pathway (Wong et al, 2015; Fong et al, 2016; Lambrus et al, 2016; Meitinger et al, 2016; preprint: Meitinger et al, 2022)."

reach
"Conversely, USP28 silencing accelerated the cell cycle progression to G2/M phase."
MiR-3940-5p affects USP28
| 13
MiR-3940-5p inhibits USP28.
| 6
MiR-3940-5p inhibits USP28. 6 / 6
| 6

reach
"MiR-3940-5p suppresses the proliferation of non-small cell lung cancer cells by targeting cyclin D1 and ubiquitin specific peptidase-28 (Ren et al., 2017)."

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
"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
"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
"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
"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."

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

reach
"These results confirmed our speculation that miR-3940-5p inhibits proliferation and induces apoptosis in NSCLC cells by targeting CCND1 and USP28."
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."
USP28 affects USP28
| 13
USP28 activates USP28.
| 4
USP28 activates USP28. 4 / 4
| 4

reach
"Loss of USP28 promotes MYC ubiquitination (45), predicting a diminished MYC-PAF1 interaction and enhanced expression of MYC target genes in USP28-deficient cells."

reach
"However, a trend towards increased USP28 protein expression was detected in CD3/CD28 activated helper T cells at 72h compared to resting naïve T cells ( Figure 1C ), suggesting that USP28 protein levels are induced by T cell activation.To better understand the role of USP28 in T cells, USP28 knockout (USP28 ) and their littermate control mice (USP28 ) were generated as previously described (24)."

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."

reach
"These findings suggested that ATM mediated phosphorylation of USP28 promotes dissociation of USP28 from KLHL2, destabilizing UCK1."
USP28 inhibits USP28.
| 3
USP28 inhibits USP28. 3 / 3
| 3

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]"

reach
"Similarly to the case of NRF2, the deubiquitinases can prevent the degradation of HIF-1α so their activity must be taken into consideration when developing a possible therapy based on the increased HIF-1α ubiquitination.For example, the ubiquitination of HIF-1α mediated by FBXW7, analyzed earlier in this review, may be inhibited by the action of the Ubiquitin Specific Peptidase 28 (USP28), which acts as a FBXW7 regulator [102]."

reach
"However, a trend towards increased USP28 protein expression was detected in CD3/CD28 activated helper T cells at 72h compared to resting naïve T cells ( Figure 1C ), suggesting that USP28 protein levels are induced by T cell activation.To better understand the role of USP28 in T cells, USP28 knockout (USP28 ) and their littermate control mice (USP28 ) were generated as previously described (24)."
USP28 deubiquitinates USP28.
| 2
USP28 deubiquitinates USP28. 2 / 2
| 2

reach
"Indeed, USP28 de-ubiquitinates and stabilizes MYC, the USP28/MYC interaction being mediated by the F-box protein, FBW7, which targets MYC for degradation by the proteasome."

reach
"USP28 is the first DUB discovered to deubiquitinate and stabilize MYC.103 USP28 interacts with MYC via Fbw7α and reverses Fbw7α-mediated ubiquitination of MYC in the nucleoplasm."
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 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 affects CHEK2
1 | 6 6
USP28 binds CHEK2.
| 4 6
| 4 1

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

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

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

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
| 4

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

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

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 deubiquitinates CHEK2.
1 | 2
USP28 deubiquitinates CHEK2. 2 / 3
1 | 2

reach
"However, it must be noted that USP28 can also deubiquitinate and stabilise other substrates not known to be associated with Fbw7, including Claspin [92] , Chk2 [93] , and LSD1 [94] , which could also [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"USP28 also modulates the DNA damage response (DDR) in cancer cells by deubiquitinating and stabilising the checkpoint kinase 2 (CHK2), the TP53-binding protein 1 (TP53BP1) and Claspin (CLSPN) [6, 7], thereby preventing apoptosis but establishing cell cycle arrest to facilitate DNA repair [8]."
SREBF2 affects USP28
2 | 4 7
2 | 4 7

sparser
"However, it is possible that the interaction between SREBP2 and USP28 is indirect and involves additional, unknown proteins."

sparser
"It has also been demonstrated that USP28 binds to mature SREBP2, leading to its deubiquitination and stabilization."

sparser
"While both proteins localise to and interact in the nucleus, we did observe some level of interaction of SREBP2 and USP28 also in the cytoplasm."

sparser
"Moreover, mature SREBP2 binds to USP28, leading to deubiquitination and stabilization, as well as regulation of the mevalonate pathway (MVP), which drives the synthesis of cholesterol, thus driving tu[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"Notably, it is unclear whether the interaction between SREBP2 and USP28 is direct or whether it involves additional proteins."

reach
"However, it is possible that the interaction between SREBP2 and USP28 is indirect and involves additional, unknown proteins."

reach
"While both proteins localise to and interact in the nucleus, we did observe some level of interaction of SREBP2 and USP28 also in the cytoplasm."

reach
"Notably, it is unclear whether the interaction between SREBP2 and USP28 is direct or whether it involves additional proteins.USP28 is highly upregulated in LSCC, where it stabilises the transcription factor ΔNP63 that is essential for the maintenance of the squamous lineage [16]."

reach
"Moreover, mature SREBP2 binds to USP28, leading to deubiquitination and stabilization, as well as regulation of the mevalonate pathway (MVP), which drives the synthesis of cholesterol, thus driving tu[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"We also show that USP28 binds to mature SREBP2, leading to its deubiquitination and stabilisation."
CHEK2 affects USP28
| 6 7
CHEK2 binds USP28.
| 4 6
| 4 1

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

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

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

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
| 4

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

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

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

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

sparser
"However, analyses of the USP28 sequence did not reveal any CHK2 phosphorylation motif, suggesting that USP28 cannot be phosphorylated by CHK2."
| PMC
ATR affects USP28
2 2 | 5 4
ATR phosphorylates USP28. 7 / 7
| 3 4

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 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]."

sparser
"The latest research has found that ATR phosphorylates USP28 (S67 and S714) and increases its enzymatic activity, further confirming that targeting the USP28-Np63 axis in sensitizing squamous cell tones down this DNA damage response pathways [ xref ]."

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

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
"ATR phosphorylates USP28 and increases its enzymatic activity."

sparser
"We next asked whether ATM or ATR phosphorylates USP28 upon CPPD treatment."
ATR phosphorylates USP28 on S714. 1 / 3
1 1 | 1

reach
"The latest research has found that ATR phosphorylates USP28 (S67 and S714) and increases its enzymatic activity, further confirming that targeting the USP28-Np63 axis in sensitizing squamous cell tones down this DNA damage response pathways [46]."
ATR phosphorylates USP28 on S67. 1 / 3
1 1 | 1

reach
"The latest research has found that ATR phosphorylates USP28 (S67 and S714) and increases its enzymatic activity, further confirming that targeting the USP28-Np63 axis in sensitizing squamous cell tones down this DNA damage response pathways [46]."
USP28 affects BRAF
| 12
USP28 inhibits BRAF.
| 8
USP28 inhibits BRAF. 8 / 8
| 8

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

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

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."

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

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
"USP28, a deubiquitinating enzyme, causes BRAF degradation by stabilizing FBXW7 [29]."

reach
"Loss of USP28 mediated BRAF degradation drives resistance to RAF cancer therapies."
USP28 activates BRAF.
| 3
USP28 activates BRAF. 3 / 3
| 3

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."

reach
"Therefore, the attenuation of BRAF and MAPK signaling caused by USP28 can inhibit EMT as well [16]."
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)."
Vismodegib affects USP28
| 11
Vismodegib binds USP28.
| 8

reach
"Finally, compound 14 was obtained by the deprotection of the Boc group.Based on the crystal structure of USP28 bound with Vismodegib, we initially explored the optimal fragment located in the solvent-[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Through the extensive optimization, we finally obtained several potent USP28 inhibitors, and further demonstrated the anticancer utility of the obtained compounds via the cell proliferation test and t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Therefore, the anti-cancer effects of three compounds could mainly be attributed to their inhibition on USP28 and might not involve the Hedgehog-Smoothened pathway.In summary, we obtained and solved t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"X-ray crystallography study demonstrated that Vismodegib was bound to USP28 protein via a novel allosteric pocket."

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
"We discovered that the marketed anticancer drug Vismodegib ( 1 ) could directly bind to USP28, with an IC 50 value of 4.4 μM in the enzymatic inhibition assay [ 18 ]."

reach
"In current study, we firstly solved the X-ray structure of Vismodegib bound to USP28."
Vismodegib inhibits USP28.
| 3
| 3

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

reach
"Indeed, vismodegib decreases the activity of USP25 and USP28 as well as their substrates, such as c-MYC, in CRC cells[30,89]."

reach
"Compared to Vismodegib, which is a moderate inhibitor of USP28 (IC 50 = 4.81 ± 1.65 μM), the inhibition activities of 9l , 9o and 9p against the enzyme are much stronger."
CLASP1 affects USP28
| 4 7
| 4 7

reach
"The interaction between USP28 and MAST1 was observed using a Duolink in situ proximity ligation assay (PLA) kit (Cat."

sparser
"The in situ USP28-MAST1 interaction was observed as red signals (PLA dots) when USP28 and MAST1 were immunostained together, but not when they were stained with individual antibodies (Fig.  xref C)."

sparser
"On the basis of the above observations, we hypothesized that the interaction between USP28 and MAST1 might influence on MAST1 protein turnover."

reach
"We further used Duolink PLA assays to validate the interaction between USP28 and MAST1 under physiological conditions."

reach
"To delineate the molecular mechanism by which USP28 regulates MAST1 protein, we first examined the interaction between USP28 and MAST1 under physiological conditions."

reach
"On the basis of the above observations, we hypothesized that the interaction between USP28 and MAST1 might influence on MAST1 protein turnover."

sparser
"The high mRNA expression score of USP28 was proportional with MAST1 expression (Fig.  xref C; Supplementary Table S5) and showed a positive correlation between USP28-MAST1 exhibiting r value of 0.36 (Fig.  xref D)."

sparser
"We envision that targeting the USP28-MAST1 axis along with cisplatin treatment might be an alternative therapeutic strategy to overcome cisplatin resistance in cancer patients."

sparser
"To delineate the molecular mechanism by which USP28 regulates MAST1 protein, we first examined the interaction between USP28 and MAST1 under physiological conditions."

sparser
"These findings confirm that USP28 interacts with MAST1 protein at both endogenous and exogenous levels."
USP28 affects TRIM21
| 10
| 10

sparser
"Our study illustrates a cardiomyocyte-specific USP28-TRIM21 axis in regulating hypertrophic cardiomyopathy and indicates USP28 as a pharmacological target for cardiac hypertrophy."

sparser
"USP28 binds TRIM21 to regulate deubiquitinated modification and stability of TRIM21."

sparser
"We then confirmed the endogenous interaction between USP28 and TRIM21 in Ang II-induced HL-1 and heart tissues using co-IP assay (Fig. xref C)."

sparser
"Subsequently, the ability of USP28-C171A to maintaining the protein stability of TRIM21 was also decreased, compared with wildtype USP28 (Fig. xref J), although USP28-C171A still bound to TRIM21 ( xref F)."

sparser
"Taken together, USP28 binds to TRIM21 and regulates K48-linked deubiquitination and stability of TRIM21 via its active site C171 (Fig. xref K)."

sparser
"Therefore, the role of USP28-TRIM21 axis in Ang II-induced oxidative stress was examined."

sparser
"We then have investigated the binding of USP28 and TRIM21 and the maintenance of TRIM21 stabilization by USP28 via K48-linked deubiquitination."

sparser
"Apart from the USP28-TRIM21 interaction, USP28 has been reported to deubiquitinate some other substrate proteins xref , such as Cyclin E1 xref and NICD1 xref ."

sparser
"Given TRIM21 is not feasible as a therapeutic target for cardiac hypertrophy, due to its widespread expression in multi-organs, the cardiomyocyte-specific USP28-TRIM21 axis may avoid the potential and systemic side-effects induced by targeting TRIM21 alone, suggesting that targeting USP28 is a more specific strategy for cardiac hypertrophy therapy."

sparser
"Conclusion: Our study illustrates a cardiomyocyte-specific USP28-TRIM21 axis in regulating hypertrophic cardiomyopathy and presents USP28 as a potential target for the treatment of cardiac hypertrophy."
USP28 affects LIN28A
2 | 5 3
USP28 binds LIN28A.
2 | 1 3
2 | 1 3

sparser
"Specifically, the present study assessed whether RSV would disrupt the interactions between Lin28A and USP28 or prevents ERK1/2-mediated phosphorylation at S200."

sparser
"These findings support the hypothesis that disruption of the Lin28A-USP28 interaction, which stabilizes Lin28A, could underlie the mechanism of RSV of reducing Lin28A protein expression levels at higher doses."

reach
"Specifically, the present study assessed whether RSV would disrupt the interactions between Lin28A and USP28 or prevents ERK1/2-mediated phosphorylation at S200.The MTT assay results in the present study agreed with previous reports (31,32), indicating that RSV inhibited NCCIT cell viability in a dose-dependent manner with an IC of 148 µM. All experiments were performed using the same concentrations (25–150 µM)."

sparser
"For example, Haq et al ( xref ) reported that Lin28A interacts with USP28 and deubiquitinates it, maintaining the protein stability and prolonging its half-life."
USP28 increases the amount of LIN28A.
| 2
USP28 increases the amount of LIN28A. 2 / 2
| 2

reach
"Notably, USP28 knockout alone in the NCCIT cells was insufficient to completely reduce Lin28A protein expression, indicating additional regulatory mechanisms."

reach
"Given that the degradation of Lin28A was by the proteasomal pathway, this could explain the stability of Lin28A protein as USP28 deubiquitinase was also downregulated at high RSV concentrations, and at 72 and 96 h. Inhibition of USP28 using siUSP28 decreases the levels of Lin28A, but the reduction is less pronounced compared to the decrease observed after RSV treatment."
USP28 deubiquitinates LIN28A.
| 2
USP28 deubiquitinates LIN28A. 2 / 2
| 2

reach
"In addition, Lin28A, which can be deubiquitinated by USP28, is associated with cell proliferation."

reach
"However, the deubiquitinase ubiquitin-specific protease 28 (USP28) was reported to reverse Lin28A ubiquitination, preventing its degradation and increasing stability."
USP28 affects KLHL2
4 | 2 4
4 | 2 1

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."
KLHL2 binds UCK1 and USP28. 3 / 3
| 3

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

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
"USP28 binds to UCK1 via KLHL2."
USP28 affects HIF1A
1 | 6 3
USP28 inhibits HIF1A.
| 6
USP28 inhibits HIF1A. 6 / 6
| 6

reach
"PMID :11746980 [36] Flugel D, Gorlach A, Kietzmann T. 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
"GSK-3beta regulates cell growth, migration, and angiogenesis via Fbw7 and USP28 dependent degradation 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
"Flugel D, Gorlach A, Kietzmann T. Gsk-3beta regulates cell growth, migration, and angiogenesis via Fbw7 and USP28 dependent degradation of HIF-1alpha."

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

sparser
"To investigate the impact of USP28 SUMOylation on HIF-1α, we generated Flag-USP28 K99R mutant plasmid."
USP28 binds HIF1A.
1 | 1
1 | 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 affects DNA Damage
| 10
USP28 activates DNA Damage.
| 7
| 7

reach
"Additionally, USP28 promoted olaparib resistance by stabilizing SOX9 protein and enhancing DNA damage repair."

reach
"Monomeric USP28 induces replication-dependent DNA damage."

reach
"Western blot analysis of γH2AX came to the same conclusions, showing that inhibition of USP28 induces DNA damage (Fig. 7F, I), while overexpression of USP28 led to an alleviated DNA damage upon olaparib treatment (Fig. 7G)."

reach
"Arrest of cell cycle by serum deprivation also decreased and equalized γH2AX levels in both cell lines (Supplementary Figure S3C), suggesting that DNA damage induced by monomeric USP28 requires DNA replication.Assessment of DNA breakage using neutral comet assay (60) showed an increased level of DSBs in cells expressing USP28-M compared to USP28-WT (Figure 3D)."

reach
"In previous experiments, we found that targeting USP28 with AZ1 induces DNA damage in non-small cell lung cancer while also inhibiting DNA repair."

reach
"Abrogation of USP28 induces DNA damage and cell death."

reach
"Summarizing the research findings above, we can link USP28 inhibition and AZ1-induced events such as cell cycle alterations, apoptosis, immune activation, and DNA damage: Targeting USP28 induces DNA damage, activating Noxa-mediated mitochondrial apoptosis; DNA damage and mitochondrial apoptosis release dsDNA and mtDNA which activate tumor cell immunogenicity via the cGAS-STING signaling pathway; simultaneously targeting USP28 leads to c-MYC degradation, causing cell cycle arrest and inhibition of DNA repair, further promoting DNA damage-induced cell apoptosis mediated by Noxa protein, thereby enhancing tumor cell immunogenicity mediated by dsDNA and mtDNA.Platinum-based drugs are commonly used DNA-damaging agents widely employed in the treatment of malignant tumors."
USP28 inhibits DNA Damage.
| 3
| 3

reach
"Likewise, Annexin-V-positive cells were increased in USP28-KO clones compared with USP28-depleted cells reconstituted with USP28 or MAST1 (Fig. 5G), indicating that loss of USP28 sensitizes cells to cisplatin-mediated DNA damage and induced cell death.Finally, we wished to investigate the molecular mechanism of USP28 regulation on MAST1-driven cisplatin-resistance."

reach
"Western blot analysis of γH2AX came to the same conclusions, showing that inhibition of USP28 induces DNA damage (Fig. 7F, I), while overexpression of USP28 led to an alleviated DNA damage upon olaparib treatment (Fig. 7G)."

reach
"The dimeric deubiquitinase USP28 integrates 53BP1 and MYC functions to limit DNA damage."
TRIM21 affects USP28
| 10
| 10

sparser
"Our study illustrates a cardiomyocyte-specific USP28-TRIM21 axis in regulating hypertrophic cardiomyopathy and indicates USP28 as a pharmacological target for cardiac hypertrophy."

sparser
"USP28 binds TRIM21 to regulate deubiquitinated modification and stability of TRIM21."

sparser
"We then confirmed the endogenous interaction between USP28 and TRIM21 in Ang II-induced HL-1 and heart tissues using co-IP assay (Fig. xref C)."

sparser
"Subsequently, the ability of USP28-C171A to maintaining the protein stability of TRIM21 was also decreased, compared with wildtype USP28 (Fig. xref J), although USP28-C171A still bound to TRIM21 ( xref F)."

sparser
"Taken together, USP28 binds to TRIM21 and regulates K48-linked deubiquitination and stability of TRIM21 via its active site C171 (Fig. xref K)."

sparser
"Therefore, the role of USP28-TRIM21 axis in Ang II-induced oxidative stress was examined."

sparser
"We then have investigated the binding of USP28 and TRIM21 and the maintenance of TRIM21 stabilization by USP28 via K48-linked deubiquitination."

sparser
"Apart from the USP28-TRIM21 interaction, USP28 has been reported to deubiquitinate some other substrate proteins xref , such as Cyclin E1 xref and NICD1 xref ."

sparser
"Given TRIM21 is not feasible as a therapeutic target for cardiac hypertrophy, due to its widespread expression in multi-organs, the cardiomyocyte-specific USP28-TRIM21 axis may avoid the potential and systemic side-effects induced by targeting TRIM21 alone, suggesting that targeting USP28 is a more specific strategy for cardiac hypertrophy therapy."

sparser
"Conclusion: Our study illustrates a cardiomyocyte-specific USP28-TRIM21 axis in regulating hypertrophic cardiomyopathy and presents USP28 as a potential target for the treatment of cardiac hypertrophy."
OAZ1 affects USP28
| 10
OAZ1 inhibits USP28.
| 7
OAZ1 inhibits USP28. 7 / 7
| 7

reach
"Prieto-Garcia et al demonstrated that AZ1 blocks USP28-dependent stabilization of ΔNp63, leading to a decrease in SCC transplanted tumor size and number (Prieto-Garcia et al, 2020)."

reach
"Inspired by the strong effect on viral replication, we explored USP25 as an antiviral drug target using the small molecule AZ1, which effectively inhibits USP25 and USP28 enzymatic activity ."

reach
"Differently substituted molecules (AZ1: -F/-CF ; AZ2: -O-CF ; AZ4: 3x-F) inhibited USP28/USP25 in a high nanomolar to low micromolar range, while potency was significantly diminished for the non-fluoro-substituted variant AZ3 (Wrigley et al, 2017)."

reach
"In addition to SGH, AZ1 is another inhibitor of USP28 that is quite significant for the treatment of lung cancer."

reach
"Kinetic analysis of full-length USP28 inhibition by AZ1 revealed a non-competitive mode of inhibition (Wrigley et al, 2017)."

reach
"The results showed that AZ1 could effectively inhibit USP28 in non-small cell lung cancer (Fig.S1A)."

reach
"AZ1 is the first reported inhibitor of USP28, demonstrating an ability to bind to and inhibit USP28, while also exhibiting dual activity against USP25 [60]."
OAZ1 binds USP28.
| 2
| 2

reach
"AZ1 binds to USP28 at the Ub-tail binding channel—distal end of the cleft, at the intersection between the thumb, palm, and finger subdomains."

reach
"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs."
OAZ1 increases the amount of USP28.
| 1
OAZ1 increases the amount of USP28. 1 / 1
| 1

reach
"The pharmacological inhibitor of USP28, AZ1, decreased the protein levels of USP28 itself and that of SOX9 in a dose-dependent manner (Fig. 3J)."
KLHL2 affects USP28
4 | 2 4
4 | 2 1

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."
KLHL2 binds UCK1 and USP28. 3 / 3
| 3

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

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
"USP28 binds to UCK1 via KLHL2."
HDAC5 affects USP28
| 10
HDAC5 activates USP28.
| 5
HDAC5 activates USP28. 5 / 5
| 5

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

reach
"Chunyu Cao et al. reported that HDAC5 directly interacted with and stabilized the LSD1 protein, and HDAC5 promoted the stability of USP28 thus indirectly preventing the ubiquitination-dependent degradation of LSD1 in breast cancer cell lines ."

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
"As reported by Chunyu Cao et al., HDAC5 directly interacted with and stabilized the LSD1 protein, and HDAC5 promoted the stability of USP28 thus indirectly preventing the ubiquitination-dependent degradation of LSD1 in breast cancer cell lines ."
HDAC5 deubiquitinates USP28.
| 4
HDAC5 leads to the deubiquitination of USP28. 3 / 3
| 3

reach
"HDAC5 blocks USP28 polyubiquitination."

reach
"Additionally, belonging to class IIa HDACs, histone deacetylase 5 (HDAC5) attenuates the polyubiquitination of USP28 through its direct interaction with USP28, suggesting that HDAC5 has the ability to stabilize USP28 [54–56]."

reach
"Overexpression of HDAC5 significantly increases USP28 expression in cell lines and reduces USP28 polyubiquitination, supporting the idea that HDAC5 stabilizes LSD1 by enhancing its deubiquitinase prot[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
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)."
HDAC5 increases the amount of USP28.
| 1
HDAC5 increases the amount of USP28. 1 / 1
| 1

reach
"Overexpression of HDAC5 significantly increases USP28 expression in cell lines and reduces USP28 polyubiquitination, supporting the idea that HDAC5 stabilizes LSD1 by enhancing its deubiquitinase prot[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 9

reach
"Then, a series of biochemical and biophysical methods were applied to characterize the interaction between USP28 and Otilonium Bromide at both the molecular and the cellular level."

reach
"The K48-linked di-Ub hydrolysis experiment was repeated for three times, and the representative data was shown.Carr-Purcell-Meiboom-Gill (CPMG) and saturation transfer difference (STD) NMR experiments[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, two biophysical methods including one-dimensional NMR and thermal shift assay were used to verify the interaction between Otilonium Bromide and USP28."

reach
"Here in this research, after the confirmation of the interaction between Otilonium Bromide and USP28, the immunoprecipitation assay was applied to validate the involvement of USP28 in the ubiquitinati[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"After the characterization of the interaction between USP28 and Otilonium Bromide, cellular experiments were conducted to explore the novel therapeutic potential of the drug and also the related under[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"According to the results derived from the cellular thermal shift assay and the cellular Ub-PA assay, Otilonium Bromide could bind to USP28 and inhibit its covalent linking with ubiquitin in cellular c[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 3

reach
"Since the possible binding surface revealed by the HDX-MS data for Otilonium Bromide in USP28 is spatially far from the catalytic core of the enzyme, we then conclude that Otilonium Bromide inhibits t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Otilonium Bromide (OB), which is known as an antimuscarinic and calcium channel blocker and used for the treatment of uncontrollable abdominal pain, was identified to inhibit USP28′s enzymatic activit[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Besides, the Tm of USP28 negatively shifted with a value of 9.30 ℃ after the incubation with Otilonium Bromide, which indicates that the inhibitor could downregulate the thermal stability of USP28 ( F[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 affects SENP1
2 | 1 6
2 | 1 6

sparser
"SENP1 directly interacts with USP28."

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

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

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

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 RCHY1
1 | 4 4
1 | 4

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 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
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 21."
| 4

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

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

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 xref ."
CASP8 affects USP28
| 7 2
CASP8 inhibits USP28.
| 5 2
CASP8 inhibits USP28. 7 / 7
| 5 2

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

reach
"Similarly, USP28 can be cleaved and inactivated by caspase-8, preventing USP28 from deubiquitinating and stabilizing p53, thereby overriding the p53-dependent G2/M cell cycle checkpoint and eventually leading to treatment resistance [113]."

reach
"Thus, nuclear caspase-8 cleaves and thereby inactivates ubiquitin-specific peptidase 28 (USP28), the protein which stabilizes the wild-type p53 protein."

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
"In the nucleus, caspase-8 cleaves and inactivates the ubiquitin specific peptidase 28 (USP28), preventing USP28 from de-ubiquitinating and stabilizing wildtype p53."
| 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."
CASP8 activates USP28.
| 2
CASP8 activates USP28. 2 / 2
| 2

reach
"Accordingly, recombinant caspase-8 produced a robust and concentration dependent cleavage of recombinant USP28 in vitro and in whole cell lysates (XREF_SUPPLEMENTARY and XREF_SUPPLEMENTARY)."

reach
"Elevated nuclear expression of caspase-8 was recently reported to promote cleavage and inactivation of the ubiquitin-specific peptidase 28 (USP28), resulting in de facto p53 protein loss and cell cycle progression in a p53-proficient melanoma tumor model [34]."
USP28 affects vismodegib
| 8

reach
"Finally, compound 14 was obtained by the deprotection of the Boc group.Based on the crystal structure of USP28 bound with Vismodegib, we initially explored the optimal fragment located in the solvent-[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Through the extensive optimization, we finally obtained several potent USP28 inhibitors, and further demonstrated the anticancer utility of the obtained compounds via the cell proliferation test and t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"Therefore, the anti-cancer effects of three compounds could mainly be attributed to their inhibition on USP28 and might not involve the Hedgehog-Smoothened pathway.In summary, we obtained and solved t[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"X-ray crystallography study demonstrated that Vismodegib was bound to USP28 protein via a novel allosteric pocket."

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
"We discovered that the marketed anticancer drug Vismodegib ( 1 ) could directly bind to USP28, with an IC 50 value of 4.4 μM in the enzymatic inhibition assay [ 18 ]."

reach
"In current study, we firstly solved the X-ray structure of Vismodegib bound to USP28."

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
"USP28 promotes tumor progression and glycolysis by stabilizing PKM2/Hif1-α in cholangiocarcinoma."

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

reach
"In addition to promoting the cell cycle and enhancing aerobic glycolysis, USP28 can also stimulate cell proliferation."

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

reach
"After stabilizing FOXC1, USP28 enhances aerobic glycolysis and promotes cell proliferation."

reach
"Ubiquitin-specific protease 28 (USP28) can promote glycolysis through mediating PKM2, which in turn promotes the progression of Cholangiocarcinoma [37]."

reach
"Functional assays demonstrated that overexpression of USP28 promoted cell proliferation and aerobic glycolysis of colorectal cancer, while USP28 inhibition could reverse these effects."
USP28 affects USP25
3 | 1 4
USP28 binds USP25.
3 | 4
3 | 4

sparser
"Recent findings show that it also binds and inhibits USP28 and USP25."

sparser
"In a recent study by H. Wang and colleagues (Wang et al, xref ), VSM was shown to bind and inhibit USP28 and USP25."

sparser
"However, through molecular docking prediction, it was found, currently, that known inhibitors can simultaneously bind to USP25 and USP28."

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."
USP28 deubiquitinates USP25.
| 1
USP28 deubiquitinates USP25. 1 / 1
| 1

reach
"Whether USP28 can deubiquitinate the substrates of USP25, such as the BCR-ABL protein, needs further exploration [133]."
USP28 affects MAPK
| 8
USP28 inhibits MAPK.
| 6
USP28 inhibits MAPK. 6 / 6
| 6

reach
"It has been reported that loss of USP28 stabilizes BRAF and enhances MAPK pathway activity, and eventually promotes resistance to BRAF inhibitors in melanoma cells."

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)."

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
"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
"This confirms that loss-of-function mutations in USP28 can induce MAPK pathway activation by stimulating the fibrosarcoma-ERK pathway."
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 FOXM1
| 8
USP28 deubiquitinates FOXM1.
| 6
USP28 deubiquitinates FOXM1. 4 / 4
| 4

reach
"USP28 deubiquitinates and stabilizes the forkhead box M1 (FOXM1) transcription factor, which, in turn, activates the Wnt/β-catenin pathway and subsequently the expression of many EMT-related genes [146,195]."

reach
"Additionally, two other studies demonstrated that deubiquitylation and stabilization of FoxM1 by USP5 and USP28 contribute to its nuclear accumulation, which facilitates the recruitment of β-catenin to the Wnt target gene promoter and activates the Wnt signaling pathway (Chen et al., 2016; Chen L. et al., 2021)."

reach
"USP28 deubiquitinates and stabilizes FOXM1, which subsequently leads to the activation of the Wnt/β-linked protein pathway and promotes the progression of pancreatic cancer [110]."

reach
"Interestingly, another study found that USP28 can deubiquitinate and stabilize FOXM1, thus promoting cell proliferation through the Wnt/β-catenin pathway (Chen et al., 2021a)."
USP28 deubiquitinates FOXM1 on M1. 2 / 2
| 2

reach
"In another study, a reduction in G0/G1-phase cells and an increase in S-phase cells were observed in pancreatic cancer cells with ectopic expression of USP28, and this advancement in cell proliferation was mediated by the deubiquitination of Forkhead box M1 (FOXM1), which is induced by USP28 [4]."

reach
"USP28 deubiquitinates and stabilizes the forkhead box M1 (FOXM1) transcription factor, which, in turn, activates the Wnt/β-catenin pathway and subsequently the expression of many EMT-related genes [146,195]."
USP28 activates FOXM1.
| 2
USP28 activates FOXM1. 2 / 2
| 2

reach
"The function of USP28 is to elevate FOXM1 stability and after that, stimulation of Wnt can occur to promote PC progression [ 155 ]."

reach
"USP28-mediated stabilization of FOXM1 significantly promoted nucleus β-catenin trans -activation, which in turn resulted in the activation of the Wnt/β-catenin pathway and pancreatic cancer progressio[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
TP53BP1 affects TP53
| 8
| 8

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

sparser
"Previous work has shown that both p53 and USP28 directly interact with 53BP1 through its BRCT domains [ xref , xref ]."
SNAI1 affects USP28
| 3 5
| 3 5

sparser
"Such blockade was found to inhibit the interaction of USP28 with Snail1."

sparser
"This revealed that USP28 binds to and stabilizes Snail1 via deubiquitination, and these effects are dependent on GSK-3β-mediated phosphorylation."

sparser
"Subsequently, reciprocal co-immunoprecipitation assays confirmed that USP28 interacted with Snail1 in BC cells ( xref H)."

sparser
"LiCl (40 μM) treatment greatly weakened the mutual interaction between USP28 and Snail1 ( xref A)."

sparser
"These results demonstrated that the phosphorylation of the SRD domain was essential for the interaction of USP28 with Snail1."

reach
"LiCl (40 μM) treatment greatly weakened the mutual interaction between USP28 and Snail1 (Fig. 6A)."

reach
"This revealed that USP28 binds to and stabilizes Snail1 via deubiquitination, and these effects are dependent on GSK-3β-mediated phosphorylation."

reach
"Subsequently, USP28 interacted with and deubiquitinated Snail1; these effects were dependent on GSK-3beta-mediated phosphorylation."
∆NP63 affects USP28
| 7
USP28 binds ∆NP63. 7 / 7
| 7

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

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
"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)."

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
"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)."
Ubiquitin affects USP28
| 3 4
| 3 4

reach
"For the structure of USP34-Ub-PA complex, the structures of unbound USP34 and ubiquitin (derived from the structure of USP28 covalently bound to Ub, PDB ID: 6H4H) 35 were used as search models in mole[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"In the nucleus, after caspase-8-mediated cleavage at specific sites on USP28, the 30 kDa fragment encoding the UBA and UIM domains is lost, thus affecting the binding of USP28 to ubiquitin and leading[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"A structure of human USP28 bound to Ubiquitin-PA was downloaded from PDB (6HEK) ( Gersch, Wagstaff et al. 2019 ) and visualized in Discovery Studio to identify the Ub bound interface residues on USP28[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"We have also highlighted the region that USP28 binds with Ub."

sparser
"Ub binding to USP28 is similar to other USPs such as USP7/HAUSP and USP21 ( Ernst et al., 2013; Hu et al., 2002 )."

sparser
"For the structure of USP34-Ub-PA complex, the structures of unbound USP34 and ubiquitin (derived from the structure of USP28 covalently bound to Ub, PDB ID: 6H4H) 35 were used as search models in mole[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"We have also highlighted the region that USP28 binds with Ub."
USP28 affects ∆NP63
| 7
USP28 binds ∆NP63. 7 / 7
| 7

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

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
"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)."

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
"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)."
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
"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."

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
"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."
| 4

reach
"Overall, USP28 appears to alter the early phase of T cell activation and proliferation.3.5 USP28 contributes to Th17 cell differentiation and iTreg cell function."

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."

reach
"Usp28 deficiency promoted tumor cell differentiation accompanied by decreased proliferation, which suggests that USP28 acts similarly in intestinal homeostasis and colorectal cancer models."
| 3

reach
"Knockdown of USP28 could induce the differentiation of CSCs while attenuate their self-renewal, which can be reversed by LSD1 expression."

reach
"In support of these findings, USP28 knockdown impaired the self-renewal capacity of cancer stem cells (CSCs) derived from MMTV-Wnt1 tumors and promoted their differentiation."

reach
"Let-7 is a tumor suppressor that also plays an essential role in the acceleration of cell differentiation [92, 93]; hence, we infer that USP28 may inhibit cell differentiation through its indirect function on let-7."
| 6

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

reach
"Through the miR‐500a‐5p/ubiquitin specific peptidase 28 (USP28) axis, exosomal miR‐500a‐5p in CAFs could promote cancer cell proliferation, EMT, and metastasis [57]."

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)."

reach
"In NCCIT cells, wound healing assays and Matrigel cell invasion assays revealed that the USP28 activates the process of metastasis through its deubiquitination of Lin28A [74]."

reach
"Pard3 deletion led to the up-regulation of USP28, which promoted BC migration and migration via Snail1 stabilization.Given its essential role in tumor metastasis and EMT, Snail1 is considered a key regulator of aggressive phenotypes in tumors."

reach
"USP28 is a member of deubiquitinase family, and the aberrant expression and functional dysregulation of USP28 might lead to the development and metastasis of malignant tumors [12,13] ."
| 1

reach
"In addition, USP28 knockdown combined with sorafenib inhibited tumor growth and metastasis in tumor xenograft mice model."
USP25 affects USP28
3 | 4
3 | 4

sparser
"Recent findings show that it also binds and inhibits USP28 and USP25."

sparser
"In a recent study by H. Wang and colleagues (Wang et al, xref ), VSM was shown to bind and inhibit USP28 and USP25."

sparser
"However, through molecular docking prediction, it was found, currently, that known inhibitors can simultaneously bind to USP25 and USP28."

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."
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 )."
PRKD1 phosphorylates USP28.
| 2 1
PRKD1 phosphorylates USP28. 3 / 3
| 2 1

reach
"XREF_FIG shows that PRKD1 could phosphorylate a USP28 peptide containing the second, more conserved predicted phosphorylation site (XREF_FIG)."

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

reach
"We further showed that PRKD1 phosphorylates USP28, which interacts with and stabilizes ZNF304 from proteolytic degradation."
LDHA affects USP28
| 5 2
LDHA activates USP28.
| 5
LDHA activates USP28. 5 / 5
| 5

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

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)."

reach
"LDHA generating lactate enhances the USP28 signaling."

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
"Chronic stress induced epinephrine enhances LDHA dependent metabolic activity, which increases lactate and augments USP28 that serves to stabilize the MYC protein."
LDHA binds USP28.
| 2
LDHA binds USP28, MYC, and SNAI2. 2 / 2
| 2

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."

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 )."
| 6

reach
"Then, a series of biochemical and biophysical methods were applied to characterize the interaction between USP28 and Otilonium Bromide at both the molecular and the cellular level."

reach
"The K48-linked di-Ub hydrolysis experiment was repeated for three times, and the representative data was shown.Carr-Purcell-Meiboom-Gill (CPMG) and saturation transfer difference (STD) NMR experiments[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"In addition, two biophysical methods including one-dimensional NMR and thermal shift assay were used to verify the interaction between Otilonium Bromide and USP28."

reach
"Here in this research, after the confirmation of the interaction between Otilonium Bromide and USP28, the immunoprecipitation assay was applied to validate the involvement of USP28 in the ubiquitinati[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"After the characterization of the interaction between USP28 and Otilonium Bromide, cellular experiments were conducted to explore the novel therapeutic potential of the drug and also the related under[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"According to the results derived from the cellular thermal shift assay and the cellular Ub-PA assay, Otilonium Bromide could bind to USP28 and inhibit its covalent linking with ubiquitin in cellular c[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

reach
"To test whether USP28 activity is sufficient for senescence induction, USP28 was overexpressed and found to induce senescence (as indicated by growth arrest and positive SA-β-Gal staining) (Fig. 3A,B; Supplemental Fig. S3I,J) comparable with p21 overexpression."

reach
"USP28 overexpression increased the levels of multiple senescence markers, including p53, p21 , and GATA4 proteins but not p16, again suggesting that USP28 may be inducing senescence in part through the p53 and GATA4 branches (Fig. 3C–E; Supplemental Fig. S3K,L)."

reach
"To determine the dependency of p53 and GATA4 in USP28-induced senescence, we combined doxycycline-inducible (Tet-on) USP28 overexpression with either p53 inhibition (via E6 expression) or GATA4 depletion in BJ fibroblasts."

reach
"E6 or GATA4 depletion blunted USP28-induced senescence (Supplemental Fig. S4A)."

reach
"While either p53 or GATA4 depletion via shRNA reduced USP28-induced senescence as measured by SA-β-Gal activity, codepletion of p53 and GATA4 resulted in a synergistic effect (Fig. 3D)."

reach
"Together, these results indicate that p53 and GATA4 each independently contributes to USP28-induced senescence."
USP28 affects ZNF304
2 1 | 2 1
USP28 binds ZNF304.
2 | 1 1
2 | 1 1

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

reach
"The findings revealed a robust physical association between USP28 and ZNF304, a key player in cancer metastasis [29]."
USP28 deubiquitinates ZNF304.
1 | 1
USP28 leads to the deubiquitination of ZNF304. 1 / 2
1 | 1

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)."
USP28 affects HM26
| 6
HM26 binds USP28. 6 / 6
| 6

sparser
"Finally, the growth of strain USP28-HM26 did not change in the presence of cobalt with respect to the control since nickel, arsenic, and lithium showed a different growth than the control."

sparser
"Finally, the results of the phenotypic characterization of overexpression strains (USP5-HM26, USP21-HM26, and USP28-HM26), for all stress conditions tested, were in line with what was expected based on the previous classification of these proteins by RT-PCR."

sparser
"Phenotypic characterization of overexpression strains H. mediterranei HM26 USP5-HM26, USP21-HM26 and USP28-HM26."

sparser
"The phenotypic characterization of the three overexpression strains (USP5-HM26, USP21-HM26, and USP28-HM26) was performed by analyzing its growth compared with the control strain H. mediterranei HM26 in the same culture conditions as in the RT-PCR assay."

sparser
"However, analysis of H. mediterranei transcriptome (Esclapez et al. xref ) reveals that USP28-HM26 shows decreased expression in nitrogen starvation (48 h) in comparison to the control conditions."

sparser
"A significant difference was detected in the growth of USP5-HM26 and USP28-HM26 strains; they grew much faster, showing changes in specific growth rates and doubling times, and reached higher values in the stationary phase."
TET1 affects USP28
| 6
TET1 increases the amount of USP28.
| 3
TET1 increases the amount of USP28. 3 / 3
| 3

reach
"Ten-eleven translocation1 (Tet1), a DNA demethylase, can promote the expression of USP28 through facilitating demethylation of USP28 promoter."

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."
TET1 demethylates USP28.
| 3
TET1 demethylates USP28. 3 / 3
| 3

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."

reach
"Ten-eleven translocation1 (Tet1), a DNA demethylase, can promote the expression of USP28 through facilitating demethylation of USP28 promoter."

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."
SGH affects USP28
| 1 5
SGH inhibits USP28.
| 1 3
SGH inhibits USP28. 4 / 4
| 1 3

reach
"In terms of the mechanism, SGH inhibits USP28 and downregulates the important cancer transcription factor c-MYC by downregulating the cell cycle and nucleotide synthesis-related proteins to block the [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

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."

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."

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 ."
SGH decreases the amount of USP28.
| 2
SGH decreases the amount of USP28. 2 / 2
| 2

reach
"Interestingly, SGH also downregulated the levels of USP28 and c-Myc."

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)."
LIN28A affects USP28
2 | 1 3
2 | 1 3

sparser
"Specifically, the present study assessed whether RSV would disrupt the interactions between Lin28A and USP28 or prevents ERK1/2-mediated phosphorylation at S200."

sparser
"These findings support the hypothesis that disruption of the Lin28A-USP28 interaction, which stabilizes Lin28A, could underlie the mechanism of RSV of reducing Lin28A protein expression levels at higher doses."

reach
"Specifically, the present study assessed whether RSV would disrupt the interactions between Lin28A and USP28 or prevents ERK1/2-mediated phosphorylation at S200.The MTT assay results in the present study agreed with previous reports (31,32), indicating that RSV inhibited NCCIT cell viability in a dose-dependent manner with an IC of 148 µM. All experiments were performed using the same concentrations (25–150 µM)."

sparser
"For example, Haq et al ( xref ) reported that Lin28A interacts with USP28 and deubiquitinates it, maintaining the protein stability and prolonging its half-life."
KRAS affects USP28
| 5 1
KRAS activates USP28. 6 / 6
| 5 1

sparser
"JUN is involved in the transcriptional activation of USP28 in colorectal cancer (CRC) cells by activated KRAS."

reach
"The results of XREF_FIG showed that KRAS promoted the transcriptional upregulation of USP28."

reach
"Another molecule, KRAS, which plays a vital role in the signal transduction of most growth factor receptors, has been validated to increase the activity of c-Jun and upregulate USP28 [44–47]."

reach
"Furthermore, the reduction in USP28 in cells with c-Jun loss and the CHIP results indicate that c-Jun can make a connection with the USP28 promoter, revealing that c-Jun is responsible for the upregulation of USP28 caused by KRAS [47] (Fig. 3)."

reach
"JUN transcript is also responsible for KRAS-mediated transcriptional activation of USP28 in CRC."

reach
"JUN is involved in the transcriptional activation of USP28 in colorectal cancer (CRC) cells by activated KRAS."
HM26 affects USP28
| 6
HM26 binds USP28. 6 / 6
| 6

sparser
"Finally, the growth of strain USP28-HM26 did not change in the presence of cobalt with respect to the control since nickel, arsenic, and lithium showed a different growth than the control."

sparser
"Finally, the results of the phenotypic characterization of overexpression strains (USP5-HM26, USP21-HM26, and USP28-HM26), for all stress conditions tested, were in line with what was expected based on the previous classification of these proteins by RT-PCR."

sparser
"Phenotypic characterization of overexpression strains H. mediterranei HM26 USP5-HM26, USP21-HM26 and USP28-HM26."

sparser
"The phenotypic characterization of the three overexpression strains (USP5-HM26, USP21-HM26, and USP28-HM26) was performed by analyzing its growth compared with the control strain H. mediterranei HM26 in the same culture conditions as in the RT-PCR assay."

sparser
"However, analysis of H. mediterranei transcriptome (Esclapez et al. xref ) reveals that USP28-HM26 shows decreased expression in nitrogen starvation (48 h) in comparison to the control conditions."

sparser
"A significant difference was detected in the growth of USP5-HM26 and USP28-HM26 strains; they grew much faster, showing changes in specific growth rates and doubling times, and reached higher values in the stationary phase."
HIF1A affects USP28
1 | 1 4
HIF1A desumoylates USP28.
| 2
HIF1A desumoylates USP28. 2 / 2
| 2

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

sparser
"Hypoxia and HIF-1α promoted USP28 deSUMOylation."
HIF1A binds USP28.
1 | 1
1 | 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

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."

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."
ATG7 affects USP28
| 6
ATG7 activates USP28.
| 3
ATG7 activates USP28. 3 / 3
| 3

reach
"To investigate the mechanisms of ATG7 upregulating USP28 protein, we first examined usp28 mRNA levels in ATG7 knockdown transfectants."

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
"Mechanistically, ATG7 upregulates the deubiquitinase USP28, which removes the ubiquitin group from CD44 protein leading to stabilization of CD44 protein to control stem-like proprieties in bladder can[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
ATG7 increases the amount of USP28.
| 2
ATG7 increases the amount of 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
"ATG7 promoted USP28 protein expression, which directly bound to and deubiquinated CD44s protein and inhibited CD44s protein degradation."
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."
MiR-4295 affects USP28
| 5
MiR-4295 activates USP28.
| 4
MiR-4295 activates USP28. 4 / 4
| 4

reach
"USP28 was targeted by miR-4295."

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
"Accumulating evidence indicated that miR-4295 may target USP28 in non small lung cancer and high expression of USP28 promoted tumour cells proliferation 22."
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]."
| 5
| 5

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."

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
"USP28 can induce tumor angiogenesis directly and indirectly through the deubiquitination of HIF-1α and c-Myc."

reach
"USP28 Deficiency Promotes Breast and Liver Carcinogenesis as well as Tumor Angiogenesis in a HIF independent Manner."
| 3

reach
"The experiments in vitro confirmed that USP28 could promote cell proliferation, migration, and invasion in the HCC cell lines."

reach
"Finally, the functional experiments confirmed that USP28 significantly promoted proliferation, invasion, and migration, which agrees with previous tumor studies [11, 67, 68]."

reach
"The results revealed that the knockdown of USP28 also dramatically decreased the migration and invasion of HCCLM3 and Huh-7."

reach
"Functional experiments showed that down-regulation of USP28 expression effectively inhibited the proliferation, migration and invasion of HCC cells, while overexpression of USP28 produced the opposite effect."

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 MDC1
1 1 | 2 1
1 1 | 2

reach
"USP28 phosphorylation regulates the complex formation of DNA checkpoint proteins and binds to checkpoint proteins 53BP1, Claspin, and MDC1, resulting in repairing DNA damage."

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
USP28 affects H2AC20
| 5
| 5

sparser
"To test if USP28 binds Histone H2A directly, the interaction between USP28 and histone H2A was further tested by GST pull-down assay."

sparser
"As shown in Fig. 4 C, both H2A and ub-K119-H2A were detected in the GST-USP28 pulldown complex, but not in the GST control complex, demonstrating that the interactions between USP28 and H2A is direct."

sparser
"Furthermore, interactions between USP28 and H2A was tested by the Ni-NTA pull down assay using his-tagged H2A and HA-tagged USP28."

sparser
"Similar to removing the K-48 linked ubiquitin chains of p53, studies have illustrated that USP28 can interact with H2A directly and reduce the level of ubiquitination at K119 of histone H2A (ub-K119-H2A) [ xref ]."

sparser
"To determine if USP28 directly interacts with H2A, immunoprecipitation assays were performed to detect interaction between the endogenous USP28 and H2A in U2OS cells."
| 5
USP28 activates DNA replication.
| 3

reach
"Expression of monomeric USP28 promotes PAF1c recruitment, diminishes TRCs and stimulates DNA replication."

reach
"Nascent chromatin capture assays (48) showed an increased enrichment of RNAPII and CTR9 on nascent DNA in USP28-M cells (Figure 2F), suggesting that monomeric USP28 stimulates DNA replication in the vicinity of RNAPII/PAF1c-bound sites."

reach
"USP28 monomers stimulate DNA replication."
| 2

reach
"We propose that dimerization of USP28 prevents ectopic DNA replication at transcriptionally active chromatin to maintain genome stability."

reach
"Furthermore, USP28 is shown to regulate cell proliferation in U2OS cells, because depletion of USP28 promotes cell growth and DNA replication."
| 2 3
| 2 3

eidos
"USP28 contributes to tumorigenesis through modulating the lifespan of oncogenic factors such as c-Myc and DeltaNp63 , and it has been identified as a potential target for anti-cancer drug development ."

reach
"Taken together, these results suggest that USP28 could potentially modulate tumorigenesis and cancer progression by exerting control over the epigenetic state of cancer cells."

eidos
"USP28 promotes tumorigenesis and has carcinogenic potential through deubiquitination ."

reach
"In addition, deubiquitinating enzymes USP22 [200], USP28 [201], and OTUD6A [202] stabilize c-Myc and promote tumorigenesis in breast cancer, squamous cell lung carcinoma, and prostate cancer, respectively."

reach
"Additionally, USP28 and USP10 contribute to tumorigenesis by regulating the DNA damage response and stabilizing p53 under stress conditions [19–21]."
USP28 affects CDKN1A
| 5
USP28 activates CDKN1A.
| 3
USP28 activates CDKN1A. 3 / 3
| 3

reach
"Consistent with the notion that deubiquitination of ub-K119-H2A leads to transcriptional activation, we showed that depletion of USP28 inhibits the transcriptional activity of p53, p21 and p16 INK4a ."

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
"USP28 overexpression increased the levels of multiple senescence markers, including p53, p21 , and GATA4 proteins but not p16, again suggesting that USP28 may be inducing senescence in part through the p53 and GATA4 branches (Fig. 3C–E; Supplemental Fig. S3K,L)."
USP28 increases the amount of CDKN1A.
| 2
USP28 increases the amount of CDKN1A. 2 / 2
| 2

reach
"Furthermore, USP28 depletion reduced levels of basal and nutlin-3a-induced p21 expression (Fig. 2F), suggesting that USP28 may also affect p21 levels during replicative senescence, likely through p53."

reach
"Consistent with the lower expression of p53, p21 and p16, USP28-depleted 2BS cells had less blue-colored cells ( Fig. 6 C), and the numbers of senescence-positive cells in USP28 knockdown cells were a[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 affects CD44
1 | 2 2
USP28 binds CD44.
1 | 1 2
1 | 1 2

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 ]."

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."
USP28 deubiquitinates CD44.
| 1
USP28 deubiquitinates CD44. 1 / 1
| 1

reach
"Following the upregulation of USP28 induced by ATG7, CD44 can be deubiquitinated and stabilized by USP28, which mediates the enhancement of the cancer stem cell-like properties of bladder cancer."
MDC1 affects USP28
1 1 | 2 1
1 1 | 2

reach
"USP28 phosphorylation regulates the complex formation of DNA checkpoint proteins and binds to checkpoint proteins 53BP1, Claspin, and MDC1, resulting in repairing DNA damage."

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
H2AC20 affects USP28
| 5
| 5

sparser
"To test if USP28 binds Histone H2A directly, the interaction between USP28 and histone H2A was further tested by GST pull-down assay."

sparser
"As shown in Fig. 4 C, both H2A and ub-K119-H2A were detected in the GST-USP28 pulldown complex, but not in the GST control complex, demonstrating that the interactions between USP28 and H2A is direct."

sparser
"Furthermore, interactions between USP28 and H2A was tested by the Ni-NTA pull down assay using his-tagged H2A and HA-tagged USP28."

sparser
"Similar to removing the K-48 linked ubiquitin chains of p53, studies have illustrated that USP28 can interact with H2A directly and reduce the level of ubiquitination at K119 of histone H2A (ub-K119-H2A) [ xref ]."

sparser
"To determine if USP28 directly interacts with H2A, immunoprecipitation assays were performed to detect interaction between the endogenous USP28 and H2A in U2OS cells."
Cep131 affects USP28
| 5
Cep131 binds USP28.
| 3
| 3

sparser
"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs.  xref and  xref )."

sparser
"AZ1 binds to USP28 at the Ub-tail binding channel—distal end of the cleft, at the intersection between the thumb, palm, and finger subdomains."

sparser
"However, the observation that the three structurally and chemically different compounds AZ1, VSM, and FT206 interact with USP28 in the same cleft, yet exert their inhibitory activity for USP28 and its close homolog USP25 without significantly affecting other DUBs of the USP family, prompted us to further investigate the origin of this selectivity."
Cep131 inhibits USP28.
| 2
Cep131 inhibits USP28. 2 / 2
| 2

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."

sparser
"The results showed that AZ1 could effectively inhibit USP28 in non-small cell lung cancer (Fig. xref A)."
CLSPN affects USP28
1 1 | 2 1
1 1 | 2

reach
"USP28 phosphorylation regulates the complex formation of DNA checkpoint proteins and binds to checkpoint proteins 53BP1, Claspin, and MDC1, resulting in repairing DNA damage."

reach
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [XREF_BIBR]."
| 1

sparser
"Moreover, USP28 binds checkpoint proteins 53BP1, Claspin, and Mdc1 [ xref ]."
Nickel(2+) affects 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
"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."

reach
"However, USP28 protein expression was decreased by Ni ions and hypoxia in the A549 cells (XREF_FIG)."
MiR-500a-5p affects USP28
| 1 3
MiR-500a-5p activates USP28.
| 3
MiR-500a-5p activates USP28. 3 / 3
| 3

reach
"The author also demonstrated that miR-500a-5p promoted proliferation and metastasis through binding to ubiquitin-specific peptidase 28 (USP28) [98]."

reach
"Therefore, these data indicated that miR-500a-5p promotes breast cancer progression and metastasis by sponging USP28."

reach
"MiR-500a-5p in CAF-derived EVs in breast cancer promotes proliferation and metastasis ability by targeting and reducing ubiquitin-specific peptidase 28 (USP28) [41]."
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) ."
ZNF304 affects USP28
2 | 1 1
2 | 1 1

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

reach
"The findings revealed a robust physical association between USP28 and ZNF304, a key player in cancer metastasis [29]."
USP28 affects cell death
| 1 3
USP28 inhibits cell death.
| 1 1
| 1 1

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

reach
"In SCC tumors, the inhibition of USP28 reduces tumor growth and increases cell death [17]."
| PMC
USP28 activates cell death.
| 2
| 2

reach
"Abrogation of USP28 induces DNA damage and cell death."

reach
"Remarkably, the ablation of p53, 53bp1, or Usp28 in addition to Sas‐4 or Sass6 rescues the cell death and allows the double mutant embryos to develop 1 day further, when they resemble cilia mutants (Fig. 4) [51, 70, 100]."

reach
"Here, we report 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 ."

eidos
"we reported 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 ."
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 )."
USP28 affects PPARA
| 4
| 4

sparser
"Mechanistically, USP28 directly interacted with PPARα, deubiquitinated and promoted Mfn2 transcription by stabilizing PPARα (Lys152), finally blocking the morphofunctional defects of mitochondria ( xref )."

sparser
"Moreover, a recent study has demonstrated that USP28 can directly interact with PPARα (Lys152) and exert a protective effect in DCM by regulating mitochondrial homeostasis through the PPARα-Mfn2 axis."

sparser
"Also, we did not find the potential PPARα-USP28 interaction in TAC-induced mouse heart from our LC-MS/MS dataset."

sparser
"RNA sequencing, immunoprecipitation and mass spectrometry analysis, chromatin immunoprecipitation assays, chromatin immunoprecipitation sequencing, and protein pull-down assay mechanistically revealed that USP28 directly interacted with PPARα (peroxisome proliferator-activated receptor α), deubiquitinating and stabilizing PPARα (Lys152) to promote Mfn2 (mitofusin 2) transcription, thereby impeding mitochondrial morphofunctional defects."
USP28 affects MVP
| 4
USP28 increases the amount of MVP.
| 3
USP28 increases the amount of MVP. 3 / 3
| 3

reach
"Silencing of USP28 can limit the expression of MVP enzymes with a lower metabolic flux, and a dual USP28/25 inhibitor reduces viability of LSCC cells [155]."

reach
"We found that depletion of USP28 reduces the expression of multiple MVP proteins, most prominently enzymes that catalyse the sequential condensation of acetoacetyl-CoA to form isoprenoid chains that are used for protein prenylation and for the synthesis important cellular metabolites, including CoQ10, dolichol and cholesterol."

reach
"Our results show that silencing of USP28 reduces expression of MVP enzymes and lowers metabolic flux into this pathway."
USP28 activates MVP.
| 1
USP28 activates MVP. 1 / 1
| 1

reach
"Depletion of USP28 significantly reduced the proportion of labelled CoQ10, indicating downregulation of the MVP, while cholesterol labelling was only mildly affected (Fig. 1i)."
USP28 affects GST
| 4
| 4

sparser
"Briefly, purified GST-USP28 was incubated with histone H2A. The complex was then pulled down by GSH agarose beads, and the bound proteins were eluted with glutathione and analyzed by SDS-PAGE."

sparser
"GST-USP28 was cloned into pGEX vector for expression in bacteria."

sparser
"As shown in Fig. 4 C, both H2A and ub-K119-H2A were detected in the GST-USP28 pulldown complex, but not in the GST control complex, demonstrating that the interactions between USP28 and H2A is direct."

sparser
"USP28 antibodies were generated by immunizing rabbits with a GST-USP28 fusion protein consisting of GST fused on its C terminus to the N-terminal 1–332 amino acids of USP28 and affinity purified (Beth[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 affects FOXC1
| 4
USP28 activates FOXC1. 4 / 4
| 4

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."

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

reach
"After stabilizing FOXC1, USP28 enhances aerobic glycolysis and promotes cell proliferation."

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."
USP28 affects FBXW7-185aa
| 4
FBXW7-185aa binds USP28. 4 / 4
| 4

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."

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

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

reach
"Moreover, FBXW7-185aa can competitively bind USP28 to antagonizing USP28-induced c-Myc stabilization, thus suppressing glioma cell proliferation and cell cycle [74] ."
RCHY1 affects CHEK2
| 4
| 4

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

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

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

sparser
"Mechanistically, USP28 directly interacted with PPARα, deubiquitinated and promoted Mfn2 transcription by stabilizing PPARα (Lys152), finally blocking the morphofunctional defects of mitochondria ( xref )."

sparser
"Moreover, a recent study has demonstrated that USP28 can directly interact with PPARα (Lys152) and exert a protective effect in DCM by regulating mitochondrial homeostasis through the PPARα-Mfn2 axis."

sparser
"Also, we did not find the potential PPARα-USP28 interaction in TAC-induced mouse heart from our LC-MS/MS dataset."

sparser
"RNA sequencing, immunoprecipitation and mass spectrometry analysis, chromatin immunoprecipitation assays, chromatin immunoprecipitation sequencing, and protein pull-down assay mechanistically revealed that USP28 directly interacted with PPARα (peroxisome proliferator-activated receptor α), deubiquitinating and stabilizing PPARα (Lys152) to promote Mfn2 (mitofusin 2) transcription, thereby impeding mitochondrial morphofunctional defects."
GST affects USP28
| 4
| 4

sparser
"Briefly, purified GST-USP28 was incubated with histone H2A. The complex was then pulled down by GSH agarose beads, and the bound proteins were eluted with glutathione and analyzed by SDS-PAGE."

sparser
"GST-USP28 was cloned into pGEX vector for expression in bacteria."

sparser
"As shown in Fig. 4 C, both H2A and ub-K119-H2A were detected in the GST-USP28 pulldown complex, but not in the GST control complex, demonstrating that the interactions between USP28 and H2A is direct."

sparser
"USP28 antibodies were generated by immunizing rabbits with a GST-USP28 fusion protein consisting of GST fused on its C terminus to the N-terminal 1–332 amino acids of USP28 and affinity purified (Beth[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
FBXW7-185aa affects USP28
| 4
FBXW7-185aa binds USP28. 4 / 4
| 4

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."

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

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

reach
"Moreover, FBXW7-185aa can competitively bind USP28 to antagonizing USP28-induced c-Myc stabilization, thus suppressing glioma cell proliferation and cell cycle [74] ."
CHEK2 affects RCHY1, and USP28
| 4
| 4

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

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

sparser
"USP28 forms a complex with PIRH2 and CHK2 and counteracts the negative regulation of PIRH2 on CHK2 xref ."
CD44 affects USP28
1 | 1 2
1 | 1 2

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 ]."

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."
MiR-216b affects USP28
| 3
MiR-216b inhibits USP28.
| 2
MiR-216b inhibits USP28. 2 / 2
| 2

reach
"These results indicated that miR-216b downregulated USP28 and c-Myc signaling in HCC cells."

reach
"RSV could also boost miR-216b expression, which inhibits USP28 mRNA in hepatocellular carcinoma cells (44)."
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."
CircMED27 affects USP28
| 3
CircMED27 increases the amount of USP28. 3 / 3
| 3

reach
"22 In this study, we concluded that circMED27 induces the lenvatinib resistance of HCC by upregulating the expression of USP28 and sponging miR-655-3p.In summary, we demonstrate that, for the first time, a circMED27 was upregulated in HCC serum and tissues."

reach
"More importantly, circMED27 upregulated ubiquitin-specific peptidase 28 expression to revert LVN’s resistance via sponging miR-655-3p."

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."
USP28 affects proteolysis
| 3
USP28 activates proteolysis.
| 2
| 2

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
"ATG7 promoted USP28 protein expression, which directly bound to and deubiquinated CD44s protein and inhibited CD44s protein degradation."
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 TCF7L2
| 1 2
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 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 KRT1
| 1 2
| 1 2

sparser
"The interaction between USP28 and KRT1 was confirmed by immunoprecipitation and immunofluorescence staining."

sparser
"Mechanistic investigations demonstrated that USP28 interacted with KRT1 and exerted deubiquitination on KRT1, thereby maintaining the stability of KRT1."

reach
"The interaction between USP28 and KRT1 was confirmed by immunoprecipitation and immunofluorescence staining."
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
"USP28 could modulate the lifespan of c-Myc, c-JUN, Notch1, and ΔNp63 through erasing ubiquitin modifications from them [ 8 , 9 ]."
USP28 affects HYCC1
| 3
USP28 activates HYCC1. 3 / 3
| 3

reach
"The experiments in vitro confirmed that USP28 could promote cell proliferation, migration, and invasion in the HCC cell lines."

reach
"Furthermore, two deubiquitinating enzymes, USP9X and USP28, have significant expression in HCC and could similarly promote HCC cell proliferation by regulating the activity of this pathway (128, 129)."

reach
"Additionally, our molecular biology experiments validated that the knockdown of USP28 substantially reduced the proliferative and invasive abilities of the HCC cell lines.Conclusions: Our study suggests that USP28 could potentially serve as a biomarker for cancer immunologic infiltration and poor prognosis, with potential applications in developing novel cancer treatment strategies."
USP28 affects GSK3B
| 1 2
USP28 inhibits GSK3B.
| 1 1
USP28 inhibits GSK3B. 2 / 2
| 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 ."

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 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 Fbw7α
| 3
Fbw7α binds MYC and USP28. 2 / 2
| 2

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."

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."
USP28 binds Fbw7α. 1 / 1
| 1

sparser
"Supporting this notion is that USP36 and USP28 cooperatively further increase MYC levels. xref Thus, it is likely that the Fbw7α-USP28 and Fbw7γ-USP36 axis dynamically control the life cycle of MYC in different cell compartments ( xref )."
USP28 affects FOXK1
1 | 2
USP28 binds FOXK1.
1 | 1
1 | 1

reach
"Furthermore, we demonstrated that ubiquitin specific peptidase 28 (USP28) interacts with and targets FOXK1 for deubiquitination and stabilization."
USP28 deubiquitinates FOXK1.
| 1
USP28 deubiquitinates FOXK1. 1 / 1
| 1

reach
"Our research showed that FOXK1 is deubiquitinated by USP28 and facilitates cell proloferation and radioresistance by activating the Hippo pathway, suggesting that FOXK1 may act as a potential radiosensitizing target for lung cancer."
| 1 2

reach
"USP28 also binds the E3 ubiquitin ligase FBW7 to stabilize MYC and promote cell proliferation [33] ( Fig. 3 a)."

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 Cep131
| 3
| 3

sparser
"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs.  xref and  xref )."

sparser
"AZ1 binds to USP28 at the Ub-tail binding channel—distal end of the cleft, at the intersection between the thumb, palm, and finger subdomains."

sparser
"However, the observation that the three structurally and chemically different compounds AZ1, VSM, and FT206 interact with USP28 in the same cleft, yet exert their inhibitory activity for USP28 and its close homolog USP25 without significantly affecting other DUBs of the USP family, prompted us to further investigate the origin of this selectivity."

reach
"In previous experiments, we found that targeting USP28 with AZ1 induces DNA damage in non-small cell lung cancer while also inhibiting DNA repair."

reach
"Further, the high expression of the deubiquitinating enzyme USP28 was targeted by miR-4295, promoting non-small cell lung cancer cell proliferation [45]."

reach
"As a target gene of miR-4295, the high expression of USP28 will promote the occurrence of non-small cell lung cancer,14 promote the proliferation, metastasis, and angiogenesis of cancer cells, and inhibit the apoptosis of cancer cells.34 To sum up, the low expression of miR-4295 will weaken the targeted inhibitory effect on USP28, thus showing a cancer-promoting effect."

reach
"The USPs family is the largest and most important DUB family, and many of these elements are involved in the occurrence and development of multiple cancers, such as USP1, USP4, USP7, USP22, and USP28, which promote or inhibit the development and progression of colorectal, breast, and hepatocellular carcinomas via deubiquitining the hub genes of cancer-related signaling pathways [10, 11]."

reach
"Deubiquitinase USP28 promotes the malignant progression and radio-resistance of hepatocellular carcinoma by stabilizing WDHD1."

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 ] ."
UCK1 affects KLHL2
| 3
KLHL2 binds UCK1 and USP28. 3 / 3
| 3

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

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
"USP28 binds to UCK1 via KLHL2."
PARD3 affects USP28
| 3
PARD3 decreases the amount of USP28. 3 / 3
| 3

reach
"Previous studies have reported that Pard3 can regulate the stability of Snail1, however there is no clear report on whether Pard3 can regulate the expression of MMP9 by regulating the stability of Snail1.In this study we found that Pard3 silencing activated the NFκB pathway and promoted the transcription of USP28."

reach
"In addition, Pard3 silencing was found to increase the transcription of USP28 by activating the NFκB pathway.DUBs encompass approximately 90 cysteine proteases and metalloproteases, all of which play an important role in deubiquitinating and stabilizing target proteins [36]."

reach
"Moreover, Pard3 silencing led to activation of the NFkappaB pathway, promoting the expression of USP28."
KRT1 affects USP28
| 1 2
| 1 2

sparser
"The interaction between USP28 and KRT1 was confirmed by immunoprecipitation and immunofluorescence staining."

sparser
"Mechanistic investigations demonstrated that USP28 interacted with KRT1 and exerted deubiquitination on KRT1, thereby maintaining the stability of KRT1."

reach
"The interaction between USP28 and KRT1 was confirmed by immunoprecipitation and immunofluorescence staining."
KLHL2 affects UCK1, and USP28
| 3
KLHL2 binds UCK1 and USP28. 3 / 3
| 3

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

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
"USP28 binds to UCK1 via KLHL2."
JUN affects USP28
| 2 1
JUN increases the amount of USP28.
| 2
JUN increases the amount of USP28. 2 / 2
| 2

reach
"Potential candidates affected by RSV include oncogenic regulators c-JUN and c-MYC, which activate USP28 transcription (38–40)."

reach
"In contrast, c-JUN [50] and c-MYC [19] upregulate USP28 expression by binding to the USP28 promoter to initiate its transcription ( Fig. 3 d)."
JUN binds 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."
Hypoxia affects USP28
| 3
Hypoxia activates USP28.
| 2
Hypoxia activates USP28. 2 / 2
| 2

eidos
"Our data display that hypoxia modulates the activity of USP28 , which leads to the question whether hypoxia also regulates USP28 SUMOylation ."

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 ."
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 ] ."
Fbw7α affects USP28
| 3
Fbw7α binds MYC and USP28. 2 / 2
| 2

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."

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."
USP28 binds Fbw7α. 1 / 1
| 1

sparser
"Supporting this notion is that USP36 and USP28 cooperatively further increase MYC levels. xref Thus, it is likely that the Fbw7α-USP28 and Fbw7γ-USP36 axis dynamically control the life cycle of MYC in different cell compartments ( xref )."
| 1 2

reach
"USP28 also binds the E3 ubiquitin ligase FBW7 to stabilize MYC and promote cell proliferation [33] ( Fig. 3 a)."

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
DDR affects USP28
| 3
DDR inhibits USP28. 3 / 3
| 3

reach
"Overall, our research provides new insights into the DDR blocking effect, cellular toxicity mechanism, and proliferation inhibition mechanism of USP28 inhibition."

reach
"We propose that during an unperturbed cell cycle, DDR signaling due to endogenous DNA lesions or fork stalling can disrupt USP28 dimers to transiently activate USP28 for localized origin firing."

reach
"DDR signaling diminishes USP28 dimerization."
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)."
MiR-363-3p affects USP28
| 2
MiR-363-3p inhibits USP28. 2 / 2
| 2

reach
"However, ectopic expression of miR-363-3p in human hepatocellular carcinoma cell lines causes a decrease in USP28, indicating that USP28, c-Myc and miR-363-3p constitute a negative feedback loop [8]."

reach
"Furthermore, miR-363-3p suppresses the proliferation of human hepatocellular carcinoma cells by targeting S1PR1 or USP28 [48]."
Lactate affects USP28
| 2
Lactate activates USP28. 2 / 2
| 2

reach
"Interestingly, the lactate induced weak acidic environment promoted USP28 signaling as well as hydrochloric acid and acetic acid (XREF_FIG)."

reach
"LDHA generating lactate enhances the USP28 signaling."
Dioxygen affects USP28
| 2
| 2

reach
"Reactive oxygen speciesdependent down-regulation of tumor suppressor genes PTEN, USP28, DRAM, TIGAR, and CYLD under oxidative stress."

reach
"Reactive oxygen species dependent down-regulation of tumor suppressor genes PTEN, USP28, DRAM, TIGAR, and CYLD under oxidative stress."
WDHD1 affects USP28
| 1 1
| 1 1

reach
"After UbiBrowser database analysis, the interaction between USP28 and WDHD1 was verified using GST pull-down and Co-immunoprecipitation (CoIP) assay."

sparser
"After UbiBrowser database analysis, the interaction between USP28 and WDHD1 was verified using GST pull-down and Co-immunoprecipitation (CoIP) assay."
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

reach
"As a DUB, USP28, which was discovered due to its homology with USP25, can contribute to protein deubiquitination as well [3]."

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 ."

reach
"Through the miR‐500a‐5p/ubiquitin specific peptidase 28 (USP28) axis, exosomal miR‐500a‐5p in CAFs could promote cancer cell proliferation, EMT, and metastasis [57]."

reach
"Pard3 deletion led to the up-regulation of USP28, which promoted BC migration and migration via Snail1 stabilization.Given its essential role in tumor metastasis and EMT, Snail1 is considered a key regulator of aggressive phenotypes in tumors."
| 2

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."

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]."
| 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 WDHD1
| 1 1
| 1 1

reach
"After UbiBrowser database analysis, the interaction between USP28 and WDHD1 was verified using GST pull-down and Co-immunoprecipitation (CoIP) assay."

sparser
"After UbiBrowser database analysis, the interaction between USP28 and WDHD1 was verified using GST pull-down and Co-immunoprecipitation (CoIP) assay."
USP28 affects STAT5
| 1 1
| 1 1

sparser
"Although that study did not report changes in STAT5 levels, a similar interaction between USP28 and STAT5 would be consistent with our results in which ubiquitination of STAT5 in the absence of USP28 would lead to its proteasomal degradation ( xref )."

reach
"Although that study did not report changes in STAT5 levels, a similar interaction between USP28 and STAT5 would be consistent with our results in which ubiquitination of STAT5 in the absence of USP28 would lead to its proteasomal degradation (46)."
USP28 affects SNAI2
| 2
LDHA binds USP28, MYC, and SNAI2. 2 / 2
| 2

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."

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 )."
USP28 affects RECQL5
| 2
USP28 deubiquitinates RECQL5.
| 1
USP28 deubiquitinates RECQL5. 1 / 1
| 1

reach
"USP28 interacts with and deubiquitinates RECQL5."
USP28 binds RECQL5.
| 1
| 1

reach
"USP28 interacts with and deubiquitinates RECQL5."

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."
USP28 affects PIM1
1 | 1
1 | 1

sparser
"In contrast, a recent study has proposed that ubiquitination of both Pim-1 and Pim-2 is important in regulating the stability of the protein, when under conditions of hypoxia, during hypoxia, deubiquitinase USP28 preferentially binds to Pim-1 and -2 and prevents their degradation [ xref ], suggesting that alternative regulatory mechanisms of Pim kinase degradation may take place under hypoxic conditions."
USP28 affects OAZ1
| 2
| 2

reach
"AZ1 binds to USP28 at the Ub-tail binding channel—distal end of the cleft, at the intersection between the thumb, palm, and finger subdomains."

reach
"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs."
USP28 affects 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 antagonizes the Ub-dependent degradation of c-myc, as well as c-Jun and Notch, by targeting the Ub ligase Fbw7, thereby stabilizing the HIF-1α transcription factor [66] ."
USP28 affects NPPA
| 2
| 2

sparser
"In the area of excipient monographs, the committee has worked on the following substances: sodium chloride [USP27-NF22], Ammonium sulfate [USP27-NF22], Cetrimonium bromide [USP27-NF22], Asparagine [PF[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"A recent example is 〈 1043 〉 Ancillary Materials for Cell, Gene, and Tissue-Engineered Products official USP28-NF23, 1st supplement authored by the Gene and Cell Therapy Committee (GCT) which will pro[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 2
USP28 inhibits Microcephaly.
| 1
| 1

reach
"Recently, it was shown that loss of USP28 or 53BP1, two upstream components of MSP, can rescue NPC proliferation in microcephaly models with centrosome defects (Phan et al. 2021)."
USP28 activates Microcephaly.
| 1
| 1

reach
"Remarkably, both apoptosis and microcephaly were rescued by the removal of 53bp1, Usp28 or p53 in Sas‐4 conditional mutants [69, 141]."
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
LDHA binds USP28, MYC, and SNAI2. 2 / 2
| 2

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."

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 )."
USP28 affects JUN_family
| 2

sparser
"In CRC cells, the JUN gene binds to the USP28 promoter and is involved in KRAS-mediated transcriptional activation to promote CRC formation."

sparser
"In CRC cells, JUN interacts with the promoter of USP28 [ xref ]."
USP28 affects HIF1
| 2
USP28 deubiquitinates HIF1.
| 1
USP28 leads to the deubiquitination of HIF1. 1 / 1
| 1

reach
"Similar to c-Myc, hypoxia-inducible factor-1α (HIF-1α) is also one of the substrates of USP28 and FBW7, and the deubiquitination of HIF-1α catalyzed by USP28 requires the mediation of FBW7 as well."
USP28 activates HIF1.
| 1
USP28 activates HIF1. 1 / 1
| 1

reach
"For instance, USP9X influences apoptosis by targeting MCL-1 [244, 245] or regulates TGFβ signaling via KDM4C [246], while USP7 and USP24 target p53 [247, 248], USP13 targets PTEN [249], USP53 interacts with DNA damage binding protein 2 (DDB2) [250], and USP28 modulates HIF-1α [251]."
USP28 affects GATA4
| 1 1
USP28 binds GATA4.
| 1
| 1

sparser
"The USP28GATA4 axis described here may provide a mechanistic insight into GATA4 regulation."
USP28 activates GATA4.
| 1
USP28 activates GATA4. 1 / 1
| 1

reach
"USP28 depletion caused marked reduction of GATA4 accumulation (Fig. 2G,H)."
USP28 affects FT206
| 2
FT206 binds USP28. 2 / 2
| 2

reach
"Computational protonation state analysis with PropKa (Søndergaard et al, 2011) suggests that the proximity of both side chains leads to a doubly protonated H261 thereby rendering engagement in multiple bonds with E366 and with the AZ1 trifluoro-methyl group possible.VSM and FT206 bind to USP28 at similar sites located further towards the wider and open Ub-tail proximal position of the USP28-inhibitor-binding cleft (Fig. 1A,B)."

reach
"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs."
USP28 affects FN1
| 2
USP28 decreases the amount of FN1. 2 / 2
| 2

reach
"The expression of EMT-related markers increased, while the expression of N-cad, FN1, ZEB1, snail, and slug was reduced by binding to ubiquitin-specific peptidase 28."

reach
"Additionally, USP28 has been shown to reduce FN-1 and increase E-cadherin levels, which can inhibit tumor invasion and migration [3]."
| 2

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

reach
"Thus, the decreases in c-MYC levels resulted from the depletion of USP28 might underlie the requirement of the DUB in these cells, and the regulation on RecQ helicases might not be as important in terms of TNBC cell proliferation."
USP28 affects DNA repair
| 2
| 2

reach
"Additionally, USP38 and USP28 both interact with DNA damage response proteins (GO response to ionizing radiation, FDR =7.0 × 10 and 3.0 × 10 for USP38 and USP28 respectively), supporting the hypothesis that these DUBs are involved in DNA repair (Figure 3b)."

reach
"The alterations in DNA damage (Fig. 5C) and inhibition of DNA repair (Fig. 5D) induced by USP28 silencing were also consistent with AZ1 treatment."

reach
"For instance, USP28 promoted the growth of TNBC cells by regulating a DNA damage checkpoint , while MYSM1 sensitized TNBC cells to cisplatin treatment ."

reach
"For example, USP28 activates the DNA damage response upon centrosomal stress."
USP28 affects Colitis
| 2
USP28 activates Colitis. 2 / 2
| 2

reach
"These results led us to hypothesize that USP28 may play a role in T cell homeostasis or function.3.2 USP28 protects mice against the early development of DSS-induced colitis."

reach
"In this study we have shown that USP28 plays a protective role in inflammation associated with DSS-induced colitis."
USP28 affects CDKN2A
| 2
USP28 increases the amount of CDKN2A.
| 1
USP28 increases the amount of CDKN2A. 1 / 1
| 1

reach
"Consistent with the lower expression of p53, p21 and p16, USP28-depleted 2BS cells had less blue-colored cells ( Fig. 6 C), and the numbers of senescence-positive cells in USP28 knockdown cells were a[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 activates CDKN2A.
| 1
USP28 activates CDKN2A. 1 / 1
| 1

reach
"Consistent with the notion that deubiquitination of ub-K119-H2A leads to transcriptional activation, we showed that depletion of USP28 inhibits the transcriptional activity of p53, p21 and p16 INK4a ."
USP28 affects CD44s
| 2
USP28 binds CD44s. 2 / 2
| 2

reach
"Here, we discovered that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing this protein stability."

reach
"Further studies illustrated that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing its protein stabilization."
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
"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 ."

eidos
"On the other hand , ISO also downregulated USP28 expression , which in turn reduced CD44 protein stability ."
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 ATM
1 | 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
USP28 affects APC_C
| 2
USP28 activates APC_C. 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 ABCC9
| 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 ]."
STAT5 affects USP28
| 1 1
| 1 1

sparser
"Although that study did not report changes in STAT5 levels, a similar interaction between USP28 and STAT5 would be consistent with our results in which ubiquitination of STAT5 in the absence of USP28 would lead to its proteasomal degradation ( xref )."

reach
"Although that study did not report changes in STAT5 levels, a similar interaction between USP28 and STAT5 would be consistent with our results in which ubiquitination of STAT5 in the absence of USP28 would lead to its proteasomal degradation (46)."
SNAI2 affects USP28
| 2
LDHA binds USP28, MYC, and SNAI2. 2 / 2
| 2

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."

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 )."
| 1 1
Radiation, Ionizing phosphorylates USP28 on S67. 1 / 1
| 1

sparser
"Moreover, these phosphorylation events are dramatically reduced in ATM null cells ( Figure 7 E), indicating that the IR-induced phosphorylation on serine 67 and serine 714 of USP28 are ATM dependent a[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
Radiation, Ionizing leads to the phosphorylation of USP28. 1 / 1
| 1

reach
"We raised antibodies against the phosphoserine 67 and phosphoserine 714 and used them to confirm that endogenous USP28 is phosphorylated in response to IR ( Figure 7 D)."
PIM1 affects USP28
1 | 1
1 | 1

sparser
"In contrast, a recent study has proposed that ubiquitination of both Pim-1 and Pim-2 is important in regulating the stability of the protein, when under conditions of hypoxia, during hypoxia, deubiquitinase USP28 preferentially binds to Pim-1 and -2 and prevents their degradation [ xref ], suggesting that alternative regulatory mechanisms of Pim kinase degradation may take place under hypoxic conditions."
NPPA affects USP28
| 2
| 2

sparser
"In the area of excipient monographs, the committee has worked on the following substances: sodium chloride [USP27-NF22], Ammonium sulfate [USP27-NF22], Cetrimonium bromide [USP27-NF22], Asparagine [PF[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"

sparser
"A recent example is 〈 1043 〉 Ancillary Materials for Cell, Gene, and Tissue-Engineered Products official USP28-NF23, 1st supplement authored by the Gene and Cell Therapy Committee (GCT) which will pro[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
MYC affects SNAI2
| 2
LDHA binds USP28, MYC, and SNAI2. 2 / 2
| 2

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."

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 )."
MYC affects Fbw7α
| 2
Fbw7α binds MYC and USP28. 2 / 2
| 2

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."

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."
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 ]."
LEP affects USP28
| 1 1
LEP inhibits USP28. 2 / 2
| 1 1

sparser
"Here, we applied a recently reported USP28-specific inhibitor OB and showed that selective inhibition of USP28 by OB abrogates TAC-induced established hypertrophic heart failure and oxidative stress, indicating OB as a potential candidate for the treatment of heart failure."

reach
"Otilonium Bromide (OB), which is known as an antimuscarinic and calcium channel blocker and used for the treatment of uncontrollable abdominal pain, was identified to inhibit USP28′s enzymatic activit[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
LDHA affects SNAI2
| 2
LDHA binds USP28, MYC, and SNAI2. 2 / 2
| 2

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."

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 )."
JUN_family affects USP28
| 2

sparser
"In CRC cells, the JUN gene binds to the USP28 promoter and is involved in KRAS-mediated transcriptional activation to promote CRC formation."

sparser
"In CRC cells, JUN interacts with the promoter of USP28 [ xref ]."
Hydrogen-Ion Concentration leads to the deubiquitination of USP28. 2 / 2
| 2

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."

reach
"Chronic stress induced epinephrine activated LDHA to generate lactate, and the adjusted pH directed USP28 mediated deubiquitination and stabilization of MYC."
Fbw7α affects MYC, and USP28
| 2
Fbw7α binds MYC and USP28. 2 / 2
| 2

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."

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."
FT206 affects USP28
| 2
FT206 binds USP28. 2 / 2
| 2

reach
"Computational protonation state analysis with PropKa (Søndergaard et al, 2011) suggests that the proximity of both side chains leads to a doubly protonated H261 thereby rendering engagement in multiple bonds with E366 and with the AZ1 trifluoro-methyl group possible.VSM and FT206 bind to USP28 at similar sites located further towards the wider and open Ub-tail proximal position of the USP28-inhibitor-binding cleft (Fig. 1A,B)."

reach
"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs."
FOXK1 affects USP28
1 | 1
1 | 1

reach
"Furthermore, we demonstrated that ubiquitin specific peptidase 28 (USP28) interacts with and targets FOXK1 for deubiquitination and stabilization."
Cyclin affects USP28
| 2
Cyclin activates USP28. 2 / 2
| 2

reach
"It had an important function in inhibition of NSCLC (non small cell lung cancer) development that was mediated by cyclin D1 and ubiquitin specific peptidase28 (USP28) which target miR-3940-5p (Sun et al. 2013, Ren et al. 2017)."

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."
CD44s affects USP28
| 2
USP28 binds CD44s. 2 / 2
| 2

reach
"Here, we discovered that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing this protein stability."

reach
"Further studies illustrated that USP28 could directly bind to CD44s and promoted the deubiquitination of CD44s, in turn increasing its protein stabilization."
CADPE affects USP28
| 2
CADPE inhibits USP28.
| 1
CADPE inhibits USP28. 1 / 1
| 1

reach
"Similar to lanatoside C, caffeic acid 3,4-dihydroxyphenethyl ester (CADPE), which is derived from Teucrium pilosum, can downregulate USP28 and then contribute to the decrease in c-Myc."
CADPE decreases the amount of USP28.
| 1
CADPE decreases the amount of USP28. 1 / 1
| 1

reach
"It has been reported that CADPE can downregulate USP28 protein levels and Notch1, Cyclin E, and c-MYC protein levels in leukemia cells, suggesting that CADPE can be used as a c-MYC inhibitor in the tr[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
ABCC9 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 ]."
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 ."
Smo-1 affects USP28
| 1
Smo-1 inhibits USP28. 1 / 1
| 1

reach
"SUMO can suppress the activation of USP28 through direct interaction with the catalytic domain of USP28 [17, 50]."

reach
"Furthermore, we developed USP28-recruiting cGAS DUBTACs that effectively stabilized cGAS, elevated the cGAS-STING signaling pathway, and elicited an antiproliferative effect."
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 ."
| 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."
RecQ affects USP28
| 1
| 1

sparser
"However, one must bear in mind that USP28 regulates a large number of proteins (including c-MYC) implicated in the proliferation and survival of tumor cells, and the USP28-RecQ axis is only part of the action."
MiR-655-3p affects USP28
| 1
MiR-655-3p inhibits USP28. 1 / 1
| 1

reach
"Consistent with our expectations, our results indicated that miR-655-3p overexpression significantly decreased the USP28 mRNA and protein levels in HCC cells (Figures 5C–5E)."
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."
MCherry affects USP28
| 1
| 1

sparser
"The sequence encoding the vhhGFP4 nanobody (GFPNb) was synthesized as a gBlock from IDT based on available sequence (Addgene #35575) and cloned upstream of mCherry, mCherry-USP7 and mCherry-USP28 cons[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
Estrone affects USP28
| 1
Estrone 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."
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
| 1
| 1

reach
"Similar to lanatoside C, caffeic acid 3,4-dihydroxyphenethyl ester (CADPE), which is derived from Teucrium pilosum, can downregulate USP28 and then contribute to the decrease in c-Myc."
ArgR affects USP28
| 1
ArgR inhibits USP28. 1 / 1
| 1

reach
"CT1113 also inhibits USP25’s close homologue USP28 with a similar if not better efficacy ."
VSM affects USP28
| 1
VSM binds USP28. 1 / 1
| 1

reach
"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs."
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
UbV.28 binds USP28. 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 ΔFBXO42
| 1
USP28 activates ΔFBXO42. 1 / 1
| 1

reach
"Moreover, we found that USP28 overexpression rescued mutant p53 levels in ΔFBXO42 or ΔCCDC6 RPE1 cells, indicating that USP28 is downstream of FBXO42/CCDC6 (Appendix Fig. 5D)."
USP28 affects Δ
| 1
USP28 activates Δ. 1 / 1
| 1

reach
"USP28 could modulate the lifespan of c-Myc, c-JUN, Notch1, and ΔNp63 through erasing ubiquitin modifications from them [ 8 , 9 ]."
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 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 ub-K119-H2A
| 1
USP28 deubiquitinates ub-K119-H2A. 1 / 1
| 1

reach
"Elevated ub-K119-H2A has the capacity to reduce p53, p21 and p16 , indicating that USP28, which can deubiquitinate and lead to a decrease in ub-K119-H2A, has the ability to enhance the suppression of carcinogenesis by mediating p53, p21 and p16 [32]."
USP28 affects turnover
| 1
USP28 inhibits turnover. 1 / 1
| 1

sparser
"Furthermore, overexpressing USP28 inhibited FBXW7-185aa-induced c-Myc turnover ( xref )."
USP28 affects tetraubiquitin
| 1
USP28 deubiquitinates tetraubiquitin. 1 / 1
| 1

reach
"As shown in Figure 7 F, wild-type but not the catalytic-inactive USP28 robustly deubiquitinated tetraubiquitin in vitro, indicating that USP28 is indeed a bona fide DUB.Next, we examined if indeed the[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
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 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 recQ
| 1
| 1

sparser
"However, one must bear in mind that USP28 regulates a large number of proteins (including c-MYC) implicated in the proliferation and survival of tumor cells, and the USP28-RecQ axis is only part of the action."
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 p53_family
| 1
USP28 deubiquitinates p53_family. 1 / 1
| 1

reach
"Collectively, the simplest model accounting for the observed results is that 53BP1 acts as a scaffold that promotes the deubiquitylation of the p53 DNA-binding domain by USP28 in the nucleoplasm ( Fig[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
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 olaparib
| 1
| 1

reach
"Furthermore, the USP28 specific inhibitor AZ1 reduced SOX9 protein stability and increased the sensitivity of ovarian cancer cells to olaparib."

reach
"53BP1 and USP28 mediate p53 dependent cell cycle arrest in response to centrosome loss and prolonged mitosis."
USP28 affects 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."
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
USP28 affects mCherry
| 1
| 1

sparser
"The sequence encoding the vhhGFP4 nanobody (GFPNb) was synthesized as a gBlock from IDT based on available sequence (Addgene #35575) and cloned upstream of mCherry, mCherry-USP7 and mCherry-USP28 cons[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
| 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 lenvatinib
| 1
| 1

reach
"As shown in Figure 5J, USP28 knockdown significantly enhanced the sensitivity of PLC/PRF/5 and HCCLM3 to lenvatinib."
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 fibrosarcoma-ERK
| 1
USP28 inhibits fibrosarcoma-ERK. 1 / 1
| 1

reach
"This confirms that loss-of-function mutations in USP28 can induce MAPK pathway activation by stimulating the fibrosarcoma-ERK pathway."

reach
"Co-deletion of USP28, however, allows embryogenesis to proceed significantly further [97]."
USP28 affects different isoforms
| 1
USP28 activates different isoforms. 1 / 1
| 1

eidos
"USP28 can generate different isoforms by alternative splicing ."
| PMC

reach
"Therefore, the inhibition of the enzymatic activity of USP28 might promote the cellular degradation of the substrate proteins via ubiquitin-proteasome system."
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 bleomycin
| 1
| 1

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"Sensitivity to the DSB-inducing reagent, bleomycin, also increased mostly by the inactivation of both USP2 and USP28 deubiquitinases (Fig. 7C)."
USP28 affects argR
| 1
USP28 inhibits argR. 1 / 1
| 1

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"Here, we report that genetic depletion of USP28 or using CT1113, a potent small molecule targeting USP28, can strongly destabilize NOTCH1 and inhibit the growth of T-ALL cells."
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 Wnt/β-catenin
| 1
USP28 activates Wnt/β-catenin. 1 / 1
| 1

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"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 VSM
| 1
VSM binds USP28. 1 / 1
| 1

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"AZ1, VSM, and FT206 bind to USP28 at different overlapping positions inside the same hydrophobic cleft, located at the intersection of the USP-thumb and palm subdomains in the center of the concave part of the USP S1-site, which acts as a binding surface for the globular domain of the distal Ub moiety of the cleaved substrate (Figs."
USP28 affects UbV.28
| 1
UbV.28 binds USP28. 1 / 1
| 1

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"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 USP
| 1
MYC binds USP28 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 TWSG1
| 1
USP28 activates TWSG1. 1 / 1
| 1

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"Analysis of mutational patterns in USP28 also supported a TSG role."
USP28 affects Snail1
| 1
USP28 increases the amount of Snail1. 1 / 1
| 1

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"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 SLC4A1
| 1
| 1

reach
"Though ZDOCK and ClusPro gave different binding conformation of FBXW7 to the same UCID region of USP28, Pepsite2 validated that WD repeat region of FBXW7 binds to USP28 which was predicted by ZDOCK ( [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 affects RB1
| 1
USP28 decreases the amount of RB1. 1 / 1
| 1

reach
"Additionally, we found that USP28 knockdown increased the level of phospho-Rb, which further confirmed the hypothesis that USP28 depletion decreased the transcription of the tumour suppressors ( Fig. [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 affects PPARG
| 1
| 1

sparser
"Mechanistically, the hepatic deubiquitinase USP28 directly binds to PPARγ, preventing its ubiquitination and subsequent degradation, thereby maintaining the integrity of the PPARγ signaling pathway."
USP28 affects PPAR
| 1
USP28 deubiquitinates PPAR on K152. 1 / 1
| 1

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"RNA sequencing, immunoprecipitation and mass spectrometry analysis, chromatin immunoprecipitation assays, chromatin immunoprecipitation sequencing, and protein pull-down assay mechanistically revealed that USP28 directly interacted with PPARα (peroxisome proliferator-activated receptor α), deubiquitinating and stabilizing PPARα (Lys152) to promote Mfn2 (mitofusin 2) transcription, thereby impeding mitochondrial morphofunctional defects."
USP28 affects PLK3
| 1
USP28 leads to the ubiquitination of PLK3. 1 / 1
| 1

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"More interestingly, Plk3 also destabilizes SIAH2 in a kinase activity dependent manner.55 Thus, a mutual regulatory mechanism exists between Plk3 and SIAH2, which may functions to fine-tune the HIF-1 signaling.USP28, a deubiquitinase that suppresses the stability of MYC and HIF-1a,36,60,61 also appears to indirectly contribute to suppression of Plk3 by hypoxia and nickel.55 It has been shown that USP28 can be suppressed by nickel via HIF-, the ubiquitin-proteasome system-, and DNA methylation dependent mechanisms.62 USP28 prevents the suppression of Plk3 by nickel, suggests that suppression of Plk3 deubiquitination by USP28 in response to nickel could contribute to the elevated degradation of Plk3 by the ubiquitin-proteasome system."
USP28 affects PIM1/2
| 1
PIM1/2 binds USP28. 1 / 1
| 1

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"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

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"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 PARD3
| 1
USP28 decreases the amount of PARD3. 1 / 1
| 1

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"The cell motility assays showed that USP28 over-expression rescued the motility of BC cells impaired due to Pard3 overexpression, but Snail1 silence blocked this function (Fig. S2A)."

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 NFE2L2
| 1
USP28 decreases the amount of NFE2L2. 1 / 1
| 1

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"There was no alleviation of the liver injury at all (Supplementary Fig. 3a–c) and the loss of Usp28 did not cause an upregulation of NRF2 target gene expression (Supplementary Fig. 3d)."
USP28 affects NFASC
| 1
USP28 inhibits NFASC. 1 / 1
| 1

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"In addition, Pard3 silencing was found to increase the transcription of USP28 by activating the NFκB pathway.DUBs encompass approximately 90 cysteine proteases and metalloproteases, all of which play an important role in deubiquitinating and stabilizing target proteins [36]."
USP28 affects NBN
| 1
USP28 leads to the deubiquitination of NBN. 1 / 1
| 1

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"As K48-linked ubiquitination of NBS1 by SKP2 E3 ligase also occurs in cells with intact RECQL4 helicase and overexpression of USP28 reduces the ubiquitination of NBS1, these results suggest that there may be a novel control mechanism that plays an important role in the DSB response by ubiquitination and deubiquitination of NBS1."
USP28 affects Mice
| 1
USP28 inhibits Mice. 1 / 1
| 1

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"As a result, USP28 overexpression attenuated cardiac remodeling and dysfunction, lipid accumulation, and mitochondrial impairment in high-fat diet/streptozotocin-induced type 2 diabetes mice."
USP28 affects MiR-500a-5p
| 1
MiR-500a-5p binds USP28. 1 / 1
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"MiR-500a-5p is transferred from CAF to cancer cells and subsequently promotes proliferation and metastasis by binding to ubiquitin specific peptidase 28 (USP28)."
| 1
| 1

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"Therefore, the attenuation of BRAF and MAPK signaling caused by USP28 can inhibit EMT as well [16]."
USP28 affects KRT5
| 1
USP28 increases the amount of KRT5. 1 / 1
| 1

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"Prior work has also suggested that USP28 may contribute to the basal phenotype, as the downregulation of USP28 led to decrease of the basal marker CK14, in association with a shift from basal to luminal cellular morphology [54], and in lung cancer the deletion of USP28 reduces the number of lesions with expression of CK5 [55]."
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 Hypertrophy
| 1
| 1

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"Xie et al. [24] reported that USP28 regulates mitochondrial homeostasis via the PPARα-Mfn2 axis and modulates cardiac hypertrophy in diabetic cardiomyopathy."
| 1

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
USP28 affects H2AX
| 1
USP28 deubiquitinates H2AX. 1 / 1
| 1

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"Inhibition of Dub activity by N-ethylmaleimide (NEM) prevented deubiquitination of H2AX by Dub3 and to support the specificity of the assay, we demonstrate that USP28, an aspecific Dub, previously rep[MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
USP28 affects GYS
| 1
USP28 activates GYS. 1 / 1
| 1

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"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 Forkhead box
| 1
USP28 leads to the deubiquitination of Forkhead box on M1. 1 / 1
| 1

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"In another study, a reduction in G0/G1-phase cells and an increase in S-phase cells were observed in pancreatic cancer cells with ectopic expression of USP28, and this advancement in cell proliferation was mediated by the deubiquitination of Forkhead box M1 (FOXM1), which is induced by USP28 [4]."
USP28 affects FBW7-E3 ligase
| 1
USP28 inhibits FBW7-E3 ligase. 1 / 1
| 1

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"In the case of HIF1A, the deubiquitylase USP28 can antagonize the FBW7-E3 ligase to prevent HIF1A from degradation [14,23,25]."
USP28 affects ERK
| 1
USP28 inhibits ERK. 1 / 1
| 1

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"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 affects E3-ligases
| 1
E3-ligases binds USP28. 1 / 1
| 1

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"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."
USP28 affects Cysts
| 1
USP28 activates Cysts. 1 / 1
| 1

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"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

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"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."
USP28 affects Cyclin
| 1
USP28 activates Cyclin. 1 / 1
| 1

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"USP28 increases the stabilization of c-MYC and cyclin E1, thereby promoting the proliferation of cancer cells [39]."

reach
"FT206 (57) inhibited the cell growth of lung squamous cell carcinoma (LSCC), which was mediated by USP28."
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 CCDC6
| 1
USP28 activates CCDC6. 1 / 1
| 1

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"Moreover, we found that USP28 overexpression rescued mutant p53 levels in ΔFBXO42 or ΔCCDC6 RPE1 cells, indicating that USP28 is downstream of FBXO42/CCDC6 (Appendix Fig. 5D)."
| 1

eidos
"However , the underlying mechanisms of breast cancer suppression by USP28 are not clear ."
USP28 affects BRAF protein
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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 ATG7
| 1
USP28 increases the amount of ATG7. 1 / 1
| 1

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"The results showed that knockdown of ATG7 decreased human usp28 promoter driven reporter transcription activity (XREF_FIG), revealing that ATG7 promotes usp28 promoter transcription."
USP affects USP28
| 1
MYC binds USP28 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."
| 1

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
Ten-eleven translocation1 affects USP28
| 1
Ten-eleven translocation1 increases the amount of USP28. 1 / 1
| 1

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"Ten-eleven translocation1 (Tet1), a DNA demethylase, can promote the expression of USP28 through facilitating demethylation of USP28 promoter."
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

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"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)."
SLC4A1 affects USP28
| 1
| 1

reach
"Though ZDOCK and ClusPro gave different binding conformation of FBXW7 to the same UCID region of USP28, Pepsite2 validated that WD repeat region of FBXW7 binds to USP28 which was predicted by ZDOCK ( [MISSING/INVALID CREDENTIALS: limited to 200 char for Elsevier]"
RECQL5 affects USP28
| 1
| 1

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"USP28 interacts with and deubiquitinates RECQL5."
RAS affects USP28
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RAS activates USP28. 1 / 1
| 1

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"Thus, cJUN is responsible, at least in part, for RAS mediated transcriptional upregulation of USP28."
PPARG affects USP28
| 1
| 1

sparser
"Mechanistically, the hepatic deubiquitinase USP28 directly binds to PPARγ, preventing its ubiquitination and subsequent degradation, thereby maintaining the integrity of the PPARγ signaling pathway."
PIM1/2 affects USP28
| 1
PIM1/2 binds USP28. 1 / 1
| 1

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"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."
MiR-500a-5p affects USP28
| 1
MiR-500a-5p binds USP28. 1 / 1
| 1

reach
"MiR-500a-5p is transferred from CAF to cancer cells and subsequently promotes proliferation and metastasis by binding to ubiquitin specific peptidase 28 (USP28)."
MYC affects USP, and USP28
| 1
MYC binds USP28 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
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 ) ."
MDC1 affects TP53BP1
| 1
| 1

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

sparser
"Conversely, silencing KLHL2 significantly decreased the amount of endogenous UCK1 that was bound to HA-USP28 ( xref B)."
HIF1 affects USP28
| 1
HIF1 increases the amount of USP28. 1 / 1
| 1

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"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."
HDAC affects USP28
| 1
HDAC increases the amount of USP28. 1 / 1
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"16 Similarly, decreased HDAC activity promotes the acetylation of Htt protein and also causes an increase in Ube2e3, SUMo2, and USp28 expression."
GATA4 affects USP28
| 1
| 1

sparser
"The USP28GATA4 axis described here may provide a mechanistic insight into GATA4 regulation."
FBXO42 affects USP28
| 1
FBXO42 inhibits USP28. 1 / 1
| 1

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"Genetic ablation of FBXO42 or CCDC6 in PANC-1 cells also reduced USP28 protein abundance, without affecting USP28 transcript levels (Fig. 3I; Appendix Fig. 5C), indicating that the post-translational regulation of USP28 by FBXO42/CCDC6 is not unique to RPE1 cells."
ERK affects USP28
| 1
ERK phosphorylates USP28. 1 / 1
| 1

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"This indicates that both stabilization pathways (USP28 and Lin28A phosphorylation by MAPK/ERK) can be blocked by RSV to reduce Lin28A protein expression."
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
E3-ligases binds USP28. 1 / 1
| 1

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"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."
CircMED27 affects USP28
| 1
CircMED27 increases the amount of USP28. 1 / 1
| 1

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"CircMED27 promoted lenvatinib resistance of HCC through sponging miR-655-3p and increasing the expression of USP28 (68)."
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 ]."
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
CENPJ affects USP28
| 1
CENPJ activates USP28. 1 / 1
| 1

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"The acentriolar cells in the Cenpj null embryos activate a cell death pathway that is dependent on 53BP1 (gene name Trp53bp1), USP28 (gene name Usp28), and p53 (gene name Trp53), which is known as the mitotic surveillance pathway (Lambrus and Holland, 2017; Xiao et al., 2021)."
CCDC6 affects USP28
| 1
CCDC6 inhibits USP28. 1 / 1
| 1

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"Genetic ablation of FBXO42 or CCDC6 in PANC-1 cells also reduced USP28 protein abundance, without affecting USP28 transcript levels (Fig. 3I; Appendix Fig. 5C), indicating that the post-translational regulation of USP28 by FBXO42/CCDC6 is not unique to RPE1 cells."
BRAF affects USP28
| 1
BRAF activates USP28. 1 / 1
| 1

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"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."