LOCUS       NM_001400784            4781 bp    mRNA    linear   PRI 13-DEC-2024
DEFINITION  Homo sapiens ubiquitin specific peptidase 28 (USP28), transcript
            variant 23, mRNA.
ACCESSION   NM_001400784
VERSION     NM_001400784.1
KEYWORDS    RefSeq.
SOURCE      Homo sapiens (human)
  ORGANISM  Homo sapiens
            Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi;
            Mammalia; Eutheria; Euarchontoglires; Primates; Haplorrhini;
            Catarrhini; Hominidae; Homo.
REFERENCE   1  (bases 1 to 4781)
  AUTHORS   Jin,C., Einig,E., Xu,W., Kollampally,R.B., Schlosser,A., Flentje,M.
            and Popov,N.
  TITLE     The dimeric deubiquitinase USP28 integrates 53BP1 and MYC functions
            to limit DNA damage
  JOURNAL   Nucleic Acids Res 52 (6), 3011-3030 (2024)
   PUBMED   38227944
  REMARK    GeneRIF: The dimeric deubiquitinase USP28 integrates 53BP1 and MYC
            functions to limit DNA damage.
REFERENCE   2  (bases 1 to 4781)
  AUTHORS   Meitinger,F., Belal,H., Davis,R.L., Martinez,M.B., Shiau,A.K.,
            Oegema,K. and Desai,A.
  TITLE     Control of cell proliferation by memories of mitosis
  JOURNAL   Science 383 (6690), 1441-1448 (2024)
   PUBMED   38547292
  REMARK    GeneRIF: Control of cell proliferation by memories of mitosis.
REFERENCE   3  (bases 1 to 4781)
  AUTHORS   Karapurkar,J.K., Colaco,J.C., Suresh,B., Tyagi,A., Woo,S.H.,
            Jo,W.J., Ko,N., Singh,V., Hong,S.H., Oh,S.J., Kim,K.S. and
            Ramakrishna,S.
  TITLE     USP28 promotes tumorigenesis and cisplatin resistance by
            deubiquitinating MAST1 protein in cancer cells
  JOURNAL   Cell Mol Life Sci 81 (1), 145 (2024)
   PUBMED   38498222
  REMARK    GeneRIF: USP28 promotes tumorigenesis and cisplatin resistance by
            deubiquitinating MAST1 protein in cancer cells.
            Publication Status: Online-Only
REFERENCE   4  (bases 1 to 4781)
  AUTHORS   Zhou,W., Chen,J. and Wang,J.
  TITLE     Comprehensive prognostic and immunological analysis of Ubiquitin
            Specific Peptidase 28 in pan-cancers and identification of its role
            in hepatocellular carcinoma cell lines
  JOURNAL   Aging (Albany NY) 15 (13), 6545-6576 (2023)
   PUBMED   37450415
  REMARK    GeneRIF: Comprehensive prognostic and immunological analysis of
            Ubiquitin Specific Peptidase 28 in pan-cancers and identification
            of its role in hepatocellular carcinoma cell lines.
REFERENCE   5  (bases 1 to 4781)
  AUTHORS   Li,J., Peng,J., Wu,L., Shen,X., Zhen,X., Zhang,Y., Ma,H., Xu,Y.,
            Xiong,Q., Zhu,Q. and Zhang,P.
  TITLE     The deubiquitinase USP28 maintains the expression of the
            transcription factor MYCN and is essential in neuroblastoma cells
  JOURNAL   J Biol Chem 299 (7), 104856 (2023)
   PUBMED   37230388
  REMARK    GeneRIF: The deubiquitinase USP28 maintains the expression of the
            transcription factor MYCN and is essential in neuroblastoma cells.
REFERENCE   6  (bases 1 to 4781)
  AUTHORS   Popov,N., Herold,S., Llamazares,M., Schulein,C. and Eilers,M.
  TITLE     Fbw7 and Usp28 regulate myc protein stability in response to DNA
            damage
  JOURNAL   Cell Cycle 6 (19), 2327-2331 (2007)
   PUBMED   17873522
  REMARK    GeneRIF: Usp28 dissociates from Fbw7alpha in response to UV
            irradiation, providing a mechanism how Fbw7-mediated degradation of
            Myc is enhanced upon DNA damage.
REFERENCE   7  (bases 1 to 4781)
  AUTHORS   Popov,N., Wanzel,M., Madiredjo,M., Zhang,D., Beijersbergen,R.,
            Bernards,R., Moll,R., Elledge,S.J. and Eilers,M.
  TITLE     The ubiquitin-specific protease USP28 is required for MYC stability
  JOURNAL   Nat Cell Biol 9 (7), 765-774 (2007)
   PUBMED   17558397
  REMARK    GeneRIF: High expression levels of USP28 are found in colon and
            breast carcinomas, and stabilization of MYC by USP28 is essential
            for tumour-cell proliferation.
REFERENCE   8  (bases 1 to 4781)
  AUTHORS   Zhang,D., Zaugg,K., Mak,T.W. and Elledge,S.J.
  TITLE     A role for the deubiquitinating enzyme USP28 in control of the
            DNA-damage response
  JOURNAL   Cell 126 (3), 529-542 (2006)
   PUBMED   16901786
  REMARK    GeneRIF: Using a human cell line that faithfully recapitulated the
            Chk2-p53-PUMA pathway, we show that USP28 is required to stabilize
            Chk2 and 53BP1 in response to DNA damage.
REFERENCE   9  (bases 1 to 4781)
  AUTHORS   Puente,X.S., Sanchez,L.M., Overall,C.M. and Lopez-Otin,C.
  TITLE     Human and mouse proteases: a comparative genomic approach
  JOURNAL   Nat Rev Genet 4 (7), 544-558 (2003)
   PUBMED   12838346
  REMARK    Review article
REFERENCE   10 (bases 1 to 4781)
  AUTHORS   Valero,R., Bayes,M., Francisca Sanchez-Font,M., Gonzalez-Angulo,O.,
            Gonzalez-Duarte,R. and Marfany,G.
  TITLE     Characterization of alternatively spliced products and
            tissue-specific isoforms of USP28 and USP25
  JOURNAL   Genome Biol 2 (10), RESEARCH0043 (2001)
   PUBMED   11597335
  REMARK    GeneRIF: molecular cloning of USP28 & characterization of
            alternatively spliced products and tissue-specific isoforms
COMMENT     REVIEWED REFSEQ: This record has been curated by NCBI staff. The
            reference sequence was derived from AP001874.4, AP003170.4 and
            KF459544.1.
            
            Summary: The protein encoded by this gene is a deubiquitinase
            involved in the DNA damage pathway and DNA damage-induced
            apoptosis. Overexpression of this gene is seen in several cancers.
            [provided by RefSeq, Oct 2016].
            
            Sequence Note: The RefSeq transcript and protein were derived from
            genomic sequence to make the sequence consistent with the reference
            genome assembly. The genomic coordinates used for the transcript
            record were based on transcript alignments.
            
            Publication Note:  This RefSeq record includes a subset of the
            publications that are available for this gene. Please see the Gene
            record to access additional publications.
            
            ##Evidence-Data-START##
            RNAseq introns :: mixed sample support SAMEA1965299, SAMEA1966682
                              [ECO:0006172]
            ##Evidence-Data-END##
PRIMARY     REFSEQ_SPAN         PRIMARY_IDENTIFIER PRIMARY_SPAN        COMP
            1-128               AP001874.4         7291-7418           c
            129-261             AP003170.4         108788-108920       c
            262-367             AP003170.4         97950-98055         c
            368-527             AP003170.4         96885-97044         c
            528-614             AP003170.4         90536-90622         c
            615-752             AP003170.4         89707-89844         c
            753-826             AP003170.4         88207-88280         c
            827-906             AP003170.4         87154-87233         c
            907-1055            AP003170.4         85484-85632         c
            1056-1183           AP003170.4         83520-83647         c
            1184-1279           AP003170.4         79892-79987         c
            1280-1459           AP003170.4         73945-74124         c
            1460-1668           AP003170.4         71461-71669         c
            1669-1739           AP003170.4         70172-70242         c
            1740-1968           AP003170.4         68563-68791         c
            1969-2160           AP003170.4         65350-65541         c
            2161-2300           AP003170.4         64585-64724         c
            2301-2486           AP003170.4         64238-64423         c
            2487-2582           AP003170.4         62776-62871         c
            2583-2761           AP003170.4         61155-61333         c
            2762-2840           AP003170.4         60960-61038         c
            2841-2920           AP003170.4         60085-60164         c
            2921-3044           AP003170.4         59445-59568         c
            3045-3240           AP003170.4         57770-57965         c
            3241-3524           KF459544.1         112-395
            3525-4781           AP003170.4         54162-55418         c
FEATURES             Location/Qualifiers
     source          1..4781
                     /organism="Homo sapiens"
                     /mol_type="mRNA"
                     /db_xref="taxon:9606"
                     /chromosome="11"
                     /map="11q23.2"
     gene            1..4781
                     /gene="USP28"
                     /note="ubiquitin specific peptidase 28"
                     /db_xref="GeneID:57646"
                     /db_xref="HGNC:HGNC:12625"
                     /db_xref="MIM:610748"
     exon            1..128
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     CDS             72..3416
                     /gene="USP28"
                     /EC_number="3.4.19.12"
                     /note="isoform 20 is encoded by transcript variant 23;
                     ubiquitin thioesterase 28; deubiquitinating enzyme 28;
                     ubiquitin-specific-processing protease 28; ubiquitin
                     thiolesterase 28"
                     /codon_start=1
                     /product="ubiquitin carboxyl-terminal hydrolase 28 isoform
                     20"
                     /protein_id="NP_001387713.1"
                     /db_xref="GeneID:57646"
                     /db_xref="HGNC:HGNC:12625"
                     /db_xref="MIM:610748"
                     /translation="MTAELQQDDAAGAADGHGSASNGDITQAVSLLTDERVKEPSQDT
                     VATEPSEVEGSAANKEVLAKVIDLTHDNKDDLQAAIALSLLESPKIQADGRDLNRMHE
                     ATSAETKRSKRKRCEVWGENPNPNDWRRVDGWPVGLKNVGNTCWFSAVIQSLFQLPEF
                     RRLVLSYSLPQNVLENCRSHTEKRNIMFMQELQYLFALMMGSNRKFVDPSAALDLLKG
                     AFRSSEEQQQDVSEFTHKLLDWLEDAFQLAVNVNSSPRNKSENPMVQLFYGTFLTEGV
                     REGKPFCNNETFGQYPLQVNGYRNLDECLEGAMVEGDVELLPSDHSVKYGQERWFTKL
                     PPVLTFELSRFEFNQSLGQPEKIHNKLEFPQIIYMDRYMYRSKELIRNKRECIRKLKE
                     EIKILQQKLERYVKYGSGPARFPLPDMLKYVIEFASTKPASESCPPESDTHMTLPLSS
                     VHCSVSDQTSKESTSTESSSQDVESTFSSPEDSLPKSKPLTSSRSSMEMPSQPAPRTV
                     TDEEINFVKTCLQRWRSEIEQDIQDLKTCIASTTQTIEQMYCDPLLRQVPYRLHAVLV
                     HEGQANAGHYWAYIYNQPRQSWLKYNDISVTESSWEEVERDSYGGLRNVSAYCLMYIN
                     DKLPYFNAEAAPTESDQMSEVEALSVELKHYIQEDNWRFEQEVEEWEEEQSCKIPQME
                     SSTNSSSQDYSTSQEPSVASSHGVRCLSSEHAVIVKEQTAQAIANTARAYEKSGVEAA
                     LSELKEAEPKKPMPQETNLAEQSEQPPKANDAESTAQPNSEVSEVEIPSVGRILVRSD
                     ADGYDEEVMLSPAMQGVILAIAKARQTFDRDGSEAGLIKAFHEEYSRLYQLAKETPTS
                     HSDPRLQHVLVYFFQNEAPKRVVERTLLEQFADKNLSYDERSISIMKVAQAKLKEIGP
                     DDMNMEEYKKWHEDYSLFRKVSVYLLTGLELYQKGKYQEALSYLVYAYQSNAALLMKG
                     PRRGVKESVIALYRRKCLLELNAKAASLFETNDDHSVTEGINVMNELIIPCIHLIINN
                     DISKDDLDAIEVMRNHWCSYLGQDIAENLQLCLGEFLPRLLDPSAEIIVLKEPPTIRP
                     NSPYDLCSRFAAVMESIQGVSTVTVK"
     exon            129..261
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            262..367
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            368..527
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            528..614
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            615..752
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            753..826
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            827..906
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            907..1055
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            1056..1183
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            1184..1279
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            1280..1459
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            1460..1668
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            1669..1739
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            1740..1968
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            1969..2160
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            2161..2300
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            2301..2486
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            2487..2582
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            2583..2761
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            2762..2840
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            2841..2920
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            2921..3044
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            3045..3240
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     exon            3241..4781
                     /gene="USP28"
                     /inference="alignment:Splign:2.1.0"
     regulatory      4763..4768
                     /regulatory_class="polyA_signal_sequence"
                     /gene="USP28"
                     /note="hexamer: AATAAA"
     polyA_site      4781
                     /gene="USP28"
                     /note="major polyA site"
ORIGIN      
        1 ctagtcctga gaggctgggc cggcggcggc tgcggcggga gaccggtgac ccgcggctgg
       61 gcgcctcggc catgactgcg gagctgcagc aggacgacgc ggccggcgcg gcagacggcc
      121 acggctcggc cagtaatggt gacattactc aggcagtcag ccttctcact gatgagagag
      181 ttaaggagcc cagtcaagac actgttgcta cagaaccatc tgaagtagag gggagtgctg
      241 ccaacaagga agtattagca aaagttatag accttactca tgataacaaa gatgatcttc
      301 aggctgccat tgctttgagt ctactggagt ctcccaaaat tcaagctgat ggaagagatc
      361 ttaacaggat gcatgaagca acctctgcag aaactaaacg ctcaaagaga aaacgctgtg
      421 aagtctgggg agaaaacccc aatcccaatg actggaggag agttgatggt tggccagttg
      481 ggctgaaaaa tgttggcaat acatgttggt ttagtgctgt tattcagtct ctctttcaat
      541 tgcctgaatt tcgaagactt gttctcagtt atagtctgcc acaaaatgta cttgaaaatt
      601 gtcgaagtca tacagaaaag agaaatatca tgtttatgca agagcttcag tatttgtttg
      661 ctctaatgat gggatcaaat agaaaatttg tagacccgtc tgcagccctg gatctattaa
      721 agggagcatt ccgatcatct gaggaacagc agcaagatgt gagtgaattc acacacaagc
      781 tcctggattg gctagaggac gcattccagc tagctgttaa tgttaacagc agtcccagga
      841 acaaatctga aaatccaatg gtgcagctgt tctatggtac tttcctgact gaaggggttc
      901 gtgaaggaaa acccttttgt aacaatgaga ccttcggcca gtatcctctt caggtaaacg
      961 gttatcgcaa cttagacgag tgtttggaag gggccatggt ggagggtgat gttgagcttc
     1021 ttccctccga tcactcggtg aagtatggac aagagcgttg gtttacaaag ctacctccag
     1081 tgttgacctt tgaactctca agatttgagt ttaatcagtc ccttgggcag ccagagaaaa
     1141 ttcacaataa gctggaattt cctcagatta tttatatgga caggtacatg tacaggagca
     1201 aggagcttat tcgaaataag agagagtgta ttcgaaagtt gaaggaggaa ataaaaattc
     1261 tgcagcaaaa attggaaagg tatgtgaaat atggctcagg cccagctcgg ttcccgctcc
     1321 cggacatgct gaaatatgtt attgaatttg ctagtacaaa acctgcctca gaaagctgtc
     1381 cacctgaaag tgacacacat atgacattac cactttcttc agtgcactgc tcggtttctg
     1441 accagacatc caaggaaagt acaagtacag aaagctcttc tcaggatgtt gaaagtacct
     1501 tttcttctcc tgaagattct ttacccaagt ctaaaccact gacatcttct cggtcttcca
     1561 tggaaatgcc ttcacagcca gctccacgaa cagtcacaga tgaggagata aattttgtta
     1621 agacctgtct tcagagatgg aggagtgaga ttgaacaaga tatacaagat ttaaagactt
     1681 gtattgcaag tactactcag actattgaac agatgtactg cgatcctctc cttcgtcagg
     1741 tgccttatcg cttgcatgca gttcttgttc atgaaggaca agcaaatgct ggacactatt
     1801 gggcctatat ctataatcaa ccccgacaga gctggctcaa gtacaatgac atctctgtta
     1861 ctgaatcttc ctgggaagaa gttgaaagag attcctatgg aggcctgaga aatgttagtg
     1921 cttactgtct gatgtacatt aatgacaaac taccctactt caatgcagag gcagccccaa
     1981 ctgaatcaga tcaaatgtca gaagtggaag ccctatctgt ggaactcaag cattacattc
     2041 aggaggataa ctggcggttt gagcaggaag tagaggagtg ggaagaagag cagtcttgca
     2101 aaatccctca aatggagtcc tccaccaact cctcatcaca ggactactct acatcacaag
     2161 agccttcagt agcctcttct catggggttc gctgcttgtc gtctgagcat gctgtgattg
     2221 taaaggagca aactgcccag gctattgcaa acacagcccg tgcctatgag aagagcggtg
     2281 tagaagcggc actgagtgag cttaaagaag ctgaacccaa gaagcccatg ccccaggaaa
     2341 caaaccttgc agagcagtca gaacagcccc caaaggctaa tgatgcagag tctactgccc
     2401 agcctaattc tgaggtctct gaagtcgaga ttcccagtgt gggaaggatt ctggttagat
     2461 ctgatgcaga tggatatgat gaggaggtga tgctgagccc tgccatgcaa ggggtcatcc
     2521 tggccatagc taaagcccgt cagacctttg accgagatgg gtctgaagca gggctgatta
     2581 aggcattcca tgaagaatac tccaggctct atcagcttgc caaagagacc cccacctctc
     2641 acagtgatcc tcgacttcag catgtccttg tctacttttt ccaaaatgaa gcacccaaaa
     2701 gggtagtaga acgaaccctt ctggaacagt ttgcagataa aaatcttagc tatgatgaaa
     2761 gatcaatcag cattatgaag gtggctcaag cgaaactgaa ggaaattggt ccagatgaca
     2821 tgaatatgga agagtacaag aagtggcatg aagattatag tttgttccga aaagtgtctg
     2881 tgtatctcct aacaggccta gaactctatc aaaaaggaaa gtaccaagag gcactttcct
     2941 acctggtata tgcctaccag agcaatgctg ccctgctgat gaaggggccc cgccgggggg
     3001 tcaaagaatc cgtgattgct ttataccgaa gaaaatgcct tctggagctg aatgccaaag
     3061 cagcttctct ttttgaaaca aatgatgatc actccgtaac tgagggcatt aatgtgatga
     3121 atgaactgat catcccctgc attcacctta tcattaataa tgacatttcc aaggatgatc
     3181 tggatgccat tgaggtcatg agaaaccatt ggtgctctta ccttgggcaa gatattgcag
     3241 aaaatctgca gctgtgccta ggggagtttc tacccagact tctagatcct tctgcagaaa
     3301 tcatcgtctt gaaagagcct ccaactattc gacccaattc tccctatgac ctatgtagcc
     3361 gatttgcagc tgtcatggag tcaattcagg gagtttcaac tgtgacagtg aaataagctc
     3421 ccacatgttc aaggcccatt ctggttcctg gctgcctgcc tcttgcacag aagttcgttg
     3481 tcatagtgct caccttggga aaaggattag gtgggcacat aagattccga tcagacccca
     3541 accatgctgc atgtgtaaag aaggattgaa aataaaattg cactttttag gtacaaaatc
     3601 ataaaagctg tttcactaga aaaggcagaa agcagtgtat taaggtgttg aattacgcca
     3661 gaagacctga aatgccttgt acctacaaca atgcttaggc ttttctaagc ctcttgccac
     3721 ttttaaaatt atccttcagg cataaatatt tttgacagca gaatagaaga atgattcatg
     3781 agaacctgaa ccagatgaac agctactagt tattttatca aatacagatg acatttaaaa
     3841 attcttaact acaagagatt agaaatataa accttgcctg gctcttgcca ggagataaca
     3901 aaatgggttg ctgatgaact gcaccctttt acatgtgggt agaatataag ctcacatggc
     3961 agtgagatgt tgaaaagtca aaagagacct gtctctctcc tttcttttct atctttaaac
     4021 cagaaaacct catactcagt cctcagtgaa agaaagtaaa gtattaagga ctttagacag
     4081 aagagcattg tgtaacttga ctgaagatca tccattaata gttattaggc atttaggtaa
     4141 aattttctaa tacctaaaaa ttgtcaaaaa cagtcaatag ggctactgct ggcccaaaga
     4201 ccatttaggt ccacctcctc ttttttgctc tttttttttt tctgtgacag tttcactgtg
     4261 tcgcccaggc tggcgttcag tggtgcaatc tcagctcact gcaaactctg tctcctgggc
     4321 tcaagtgatt ctcgtgcctc agcctcccga atagctggaa ttacgggcat gcaccaccac
     4381 acctggctaa tttttgtatt tttaatagag atggggtttc accatattgg ccaggctgat
     4441 ctctaactcc tggcctcaag tgatctatct gcctccctca gcctcccaaa gtctgggatt
     4501 gcagacaagt catcgtaccc ggccttcttt tttgccctta aaagtaaggg atgtgggttt
     4561 gtacaaaaaa aaaaaaaaaa aaaaaaaaac cagcatacat atgcaaaact atatatatat
     4621 gtatatgtag agaaaaatac ttcccattga tcatttttaa aaggcttctg attggatatt
     4681 gtgttttaac caaattttaa agattaatgg aatcatgaaa gggaaaaaaa ttgatacaac
     4741 tatgcagatt ttataaatgt gcaataaaag tatttgtttt a
//
