LOCUS       NM_004476               4110 bp    mRNA    linear   PRI 09-OCT-2024
DEFINITION  Homo sapiens folate hydrolase 1 (FOLH1), transcript variant 1,
            mRNA.
ACCESSION   NM_004476
VERSION     NM_004476.3
KEYWORDS    RefSeq; MANE Select.
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 4110)
  AUTHORS   Li,R.X., Hao,Y. and Ettel,M.
  TITLE     Expression of PSMA in Tumor-Associated Vasculature Predicts Poorer
            Survival in Patients With Hepatocellular Carcinoma and Is Likely
            Associated With PD-L1
  JOURNAL   Int J Surg Pathol 32 (7), 1248-1255 (2024)
   PUBMED   38321782
  REMARK    GeneRIF: Expression of PSMA in Tumor-Associated Vasculature
            Predicts Poorer Survival in Patients With Hepatocellular Carcinoma
            and Is Likely Associated With PD-L1.
REFERENCE   2  (bases 1 to 4110)
  AUTHORS   Zhou,W., Halder,S., Herwald,S., Ghijsen,M., Shafi,G., Uttarwar,M.,
            Rosen,E., Franc,B. and Kishore,S.
  TITLE     Frequent Amplification and Overexpression of PSMA in Basallike
            Breast Cancer from Analysis of The Cancer Genome Atlas
  JOURNAL   J Nucl Med 65 (7), 1004-1006 (2024)
   PUBMED   38664014
  REMARK    GeneRIF: Frequent Amplification and Overexpression of PSMA in
            Basallike Breast Cancer from Analysis of The Cancer Genome Atlas.
            Publication Status: Online-Only
REFERENCE   3  (bases 1 to 4110)
  AUTHORS   Mlcochova,P., Barinka,C., Tykvart,J., Sacha,P. and Konvalinka,J.
  TITLE     Prostate-specific membrane antigen and its truncated form PSM'
  JOURNAL   Prostate 69 (5), 471-479 (2009)
   PUBMED   19107881
  REMARK    GeneRIF: PSM' protein is likely not generated by alternative
            splicing of the PSMA gene but by different mechanism, probably via
            an endoproteolytic cleavage of the full-length PSMA.
REFERENCE   4  (bases 1 to 4110)
  AUTHORS   Lambert,L.A. and Mitchell,S.L.
  TITLE     Molecular evolution of the transferrin receptor/glutamate
            carboxypeptidase II family
  JOURNAL   J Mol Evol 64 (1), 113-128 (2007)
   PUBMED   17160644
REFERENCE   5  (bases 1 to 4110)
  AUTHORS   Davis,M.I., Bennett,M.J., Thomas,L.M. and Bjorkman,P.J.
  TITLE     Crystal structure of prostate-specific membrane antigen, a tumor
            marker and peptidase
  JOURNAL   Proc Natl Acad Sci U S A 102 (17), 5981-5986 (2005)
   PUBMED   15837926
  REMARK    GeneRIF: Structure and relevance to the development of
            chemotherapeutics and cancer-imaging agents.
REFERENCE   6  (bases 1 to 4110)
  AUTHORS   Pangalos,M.N., Neefs,J.M., Somers,M., Verhasselt,P., Bekkers,M.,
            van der Helm,L., Fraiponts,E., Ashton,D. and Gordon,R.D.
  TITLE     Isolation and expression of novel human glutamate carboxypeptidases
            with N-acetylated alpha-linked acidic dipeptidase and dipeptidyl
            peptidase IV activity
  JOURNAL   J Biol Chem 274 (13), 8470-8483 (1999)
   PUBMED   10085079
REFERENCE   7  (bases 1 to 4110)
  AUTHORS   Troyer,J.K., Beckett,M.L. and Wright,G.L. Jr.
  TITLE     Detection and characterization of the prostate-specific membrane
            antigen (PSMA) in tissue extracts and body fluids
  JOURNAL   Int J Cancer 62 (5), 552-558 (1995)
   PUBMED   7665226
REFERENCE   8  (bases 1 to 4110)
  AUTHORS   Su,S.L., Huang,I.P., Fair,W.R., Powell,C.T. and Heston,W.D.
  TITLE     Alternatively spliced variants of prostate-specific membrane
            antigen RNA: ratio of expression as a potential measurement of
            progression
  JOURNAL   Cancer Res 55 (7), 1441-1443 (1995)
   PUBMED   7882349
REFERENCE   9  (bases 1 to 4110)
  AUTHORS   Israeli,R.S., Powell,C.T., Corr,J.G., Fair,W.R. and Heston,W.D.
  TITLE     Expression of the prostate-specific membrane antigen
  JOURNAL   Cancer Res 54 (7), 1807-1811 (1994)
   PUBMED   7511053
REFERENCE   10 (bases 1 to 4110)
  AUTHORS   Israeli,R.S., Powell,C.T., Fair,W.R. and Heston,W.D.
  TITLE     Molecular cloning of a complementary DNA encoding a
            prostate-specific membrane antigen
  JOURNAL   Cancer Res 53 (2), 227-230 (1993)
   PUBMED   8417812
COMMENT     REVIEWED REFSEQ: This record has been curated by NCBI staff. The
            reference sequence was derived from M99487.1, AC118273.4 and
            CA432757.1.
            This sequence is a reference standard in the RefSeqGene project.
            
            On Nov 22, 2018 this sequence version replaced NM_004476.2.
            
            Summary: This gene encodes a type II transmembrane glycoprotein
            belonging to the M28 peptidase family. The protein acts as a
            glutamate carboxypeptidase on different alternative substrates,
            including the nutrient folate and the neuropeptide
            N-acetyl-l-aspartyl-l-glutamate and is expressed in a number of
            tissues such as prostate, central and peripheral nervous system and
            kidney. A mutation in this gene may be associated with impaired
            intestinal absorption of dietary folates, resulting in low blood
            folate levels and consequent hyperhomocysteinemia. Expression of
            this protein in the brain may be involved in a number of
            pathological conditions associated with glutamate excitotoxicity.
            In the prostate the protein is up-regulated in cancerous cells and
            is used as an effective diagnostic and prognostic indicator of
            prostate cancer. This gene likely arose from a duplication event of
            a nearby chromosomal region. Alternative splicing gives rise to
            multiple transcript variants encoding several different isoforms.
            [provided by RefSeq, Jul 2010].
            
            Transcript Variant: This variant (1) encodes the longest isoform
            (1).
            
            Sequence Note: This RefSeq record was created from transcript and
            genomic sequence data 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##
            Transcript exon combination :: M99487.1, AF176574.1 [ECO:0000332]
            RNAseq introns              :: mixed sample support SAMEA1965299,
                                           SAMEA1966682 [ECO:0006172]
            ##Evidence-Data-END##
            
            ##RefSeq-Attributes-START##
            MANE Ensembl match     :: ENST00000256999.7/ ENSP00000256999.2
            RefSeq Select criteria :: based on conservation, expression
            ##RefSeq-Attributes-END##
            COMPLETENESS: full length.
PRIMARY     REFSEQ_SPAN         PRIMARY_IDENTIFIER PRIMARY_SPAN        COMP
            1-2567              M99487.1           69-2635
            2568-3895           AC118273.4         119133-120460       c
            3896-4023           CA432757.1         104-231             c
            4024-4024           AC118273.4         119004-119004       c
            4025-4108           CA432757.1         19-102              c
            4109-4110           AC118273.4         118918-118919       c
FEATURES             Location/Qualifiers
     source          1..4110
                     /organism="Homo sapiens"
                     /mol_type="mRNA"
                     /db_xref="taxon:9606"
                     /chromosome="11"
                     /map="11p11.12"
     gene            1..4110
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /note="folate hydrolase 1"
                     /db_xref="GeneID:2346"
                     /db_xref="HGNC:HGNC:3788"
                     /db_xref="MIM:600934"
     exon            1..311
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     CDS             194..2446
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /EC_number="3.4.17.21"
                     /note="isoform 1 is encoded by transcript variant 1;
                     glutamate carboxylase II; N-acetylated alpha-linked acidic
                     dipeptidase 1; pteroylpoly-gamma-glutamate
                     carboxypeptidase; folylpoly-gamma-glutamate
                     carboxypeptidase; membrane glutamate carboxypeptidase;
                     glutamate carboxypeptidase 2; glutamate carboxypeptidase
                     II; NAALADase I; cell growth-inhibiting gene 27 protein;
                     N-acetylated-alpha-linked acidic dipeptidase I"
                     /codon_start=1
                     /product="glutamate carboxypeptidase 2 isoform 1"
                     /protein_id="NP_004467.1"
                     /db_xref="CCDS:CCDS7946.1"
                     /db_xref="GeneID:2346"
                     /db_xref="HGNC:HGNC:3788"
                     /db_xref="MIM:600934"
                     /translation="MWNLLHETDSAVATARRPRWLCAGALVLAGGFFLLGFLFGWFIK
                     SSNEATNITPKHNMKAFLDELKAENIKKFLYNFTQIPHLAGTEQNFQLAKQIQSQWKE
                     FGLDSVELAHYDVLLSYPNKTHPNYISIINEDGNEIFNTSLFEPPPPGYENVSDIVPP
                     FSAFSPQGMPEGDLVYVNYARTEDFFKLERDMKINCSGKIVIARYGKVFRGNKVKNAQ
                     LAGAKGVILYSDPADYFAPGVKSYPDGWNLPGGGVQRGNILNLNGAGDPLTPGYPANE
                     YAYRRGIAEAVGLPSIPVHPIGYYDAQKLLEKMGGSAPPDSSWRGSLKVPYNVGPGFT
                     GNFSTQKVKMHIHSTNEVTRIYNVIGTLRGAVEPDRYVILGGHRDSWVFGGIDPQSGA
                     AVVHEIVRSFGTLKKEGWRPRRTILFASWDAEEFGLLGSTEWAEENSRLLQERGVAYI
                     NADSSIEGNYTLRVDCTPLMYSLVHNLTKELKSPDEGFEGKSLYESWTKKSPSPEFSG
                     MPRISKLGSGNDFEVFFQRLGIASGRARYTKNWETNKFSGYPLYHSVYETYELVEKFY
                     DPMFKYHLTVAQVRGGMVFELANSIVLPFDCRDYAVVLRKYADKIYSISMKHPQEMKT
                     YSVSFDSLFSAVKNFTEIASKFSERLQDFDKSNPIVLRMMNDQLMFLERAFIDPLGLP
                     DRPFYRHVIYAPSSHNKYAGESFPGIYDALFDIESKVDPSKAWGEVKRQIYVAAFTVQ
                     AAAETLSEVA"
     exon            312..417
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            418..604
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            605..706
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            707..832
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            833..1019
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1020..1113
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1114..1212
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1213..1298
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1299..1418
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1419..1501
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1502..1565
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1566..1633
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1634..1725
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1726..1816
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            1817..2081
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            2082..2163
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            2164..2256
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     exon            2257..4110
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /inference="alignment:Splign:2.1.0"
     regulatory      2545..2550
                     /regulatory_class="polyA_signal_sequence"
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /note="hexamer: AATAAA"
     polyA_site      2567
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
     regulatory      4082..4087
                     /regulatory_class="polyA_signal_sequence"
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /note="hexamer: AATAAA"
     polyA_site      4110
                     /gene="FOLH1"
                     /gene_synonym="FGCP; FOLH; GCP2; GCPII; mGCP; NAALAD1;
                     PSM; PSMA"
                     /note="major polyA site"
ORIGIN      
        1 agtgcactct agaaacactg ctgtggtgga gaaactggac cccaggtctg gagcgaattc
       61 cagcctgcag ggctgataag cgaggcatta gtgagattga gagagacttt accccgccgt
      121 ggtggttgga gggcgcgcag tagagcagca gcacaggcgc gggtcccggg aggccggctc
      181 tgctcgcgcc gagatgtgga atctccttca cgaaaccgac tcggctgtgg ccaccgcgcg
      241 ccgcccgcgc tggctgtgcg ctggggcgct ggtgctggcg ggtggcttct ttctcctcgg
      301 cttcctcttc gggtggttta taaaatcctc caatgaagct actaacatta ctccaaagca
      361 taatatgaaa gcatttttgg atgaattgaa agctgagaac atcaagaagt tcttatataa
      421 ttttacacag ataccacatt tagcaggaac agaacaaaac tttcagcttg caaagcaaat
      481 tcaatcccag tggaaagaat ttggcctgga ttctgttgag ctagcacatt atgatgtcct
      541 gttgtcctac ccaaataaga ctcatcccaa ctacatctca ataattaatg aagatggaaa
      601 tgagattttc aacacatcat tatttgaacc acctcctcca ggatatgaaa atgtttcgga
      661 tattgtacca cctttcagtg ctttctctcc tcaaggaatg ccagagggcg atctagtgta
      721 tgttaactat gcacgaactg aagacttctt taaattggaa cgggacatga aaatcaattg
      781 ctctgggaaa attgtaattg ccagatatgg gaaagttttc agaggaaata aggttaaaaa
      841 tgcccagctg gcaggggcca aaggagtcat tctctactcc gaccctgctg actactttgc
      901 tcctggggtg aagtcctatc cagatggttg gaatcttcct ggaggtggtg tccagcgtgg
      961 aaatatccta aatctgaatg gtgcaggaga ccctctcaca ccaggttacc cagcaaatga
     1021 atatgcttat aggcgtggaa ttgcagaggc tgttggtctt ccaagtattc ctgttcatcc
     1081 aattggatac tatgatgcac agaagctcct agaaaaaatg ggtggctcag caccaccaga
     1141 tagcagctgg agaggaagtc tcaaagtgcc ctacaatgtt ggacctggct ttactggaaa
     1201 cttttctaca caaaaagtca agatgcacat ccactctacc aatgaagtga caagaattta
     1261 caatgtgata ggtactctca gaggagcagt ggaaccagac agatatgtca ttctgggagg
     1321 tcaccgggac tcatgggtgt ttggtggtat tgaccctcag agtggagcag ctgttgttca
     1381 tgaaattgtg aggagctttg gaacactgaa aaaggaaggg tggagaccta gaagaacaat
     1441 tttgtttgca agctgggatg cagaagaatt tggtcttctt ggttctactg agtgggcaga
     1501 ggagaattca agactccttc aagagcgtgg cgtggcttat attaatgctg actcatctat
     1561 agaaggaaac tacactctga gagttgattg tacaccgctg atgtacagct tggtacacaa
     1621 cctaacaaaa gagctgaaaa gccctgatga aggctttgaa ggcaaatctc tttatgaaag
     1681 ttggactaaa aaaagtcctt ccccagagtt cagtggcatg cccaggataa gcaaattggg
     1741 atctggaaat gattttgagg tgttcttcca acgacttgga attgcttcag gcagagcacg
     1801 gtatactaaa aattgggaaa caaacaaatt cagcggctat ccactgtatc acagtgtcta
     1861 tgaaacatat gagttggtgg aaaagtttta tgatccaatg tttaaatatc acctcactgt
     1921 ggcccaggtt cgaggaggga tggtgtttga gctagccaat tccatagtgc tcccttttga
     1981 ttgtcgagat tatgctgtag ttttaagaaa gtatgctgac aaaatctaca gtatttctat
     2041 gaaacatcca caggaaatga agacatacag tgtatcattt gattcacttt tttctgcagt
     2101 aaagaatttt acagaaattg cttccaagtt cagtgagaga ctccaggact ttgacaaaag
     2161 caacccaata gtattaagaa tgatgaatga tcaactcatg tttctggaaa gagcatttat
     2221 tgatccatta gggttaccag acaggccttt ttataggcat gtcatctatg ctccaagcag
     2281 ccacaacaag tatgcagggg agtcattccc aggaatttat gatgctctgt ttgatattga
     2341 aagcaaagtg gacccttcca aggcctgggg agaagtgaag agacagattt atgttgcagc
     2401 cttcacagtg caggcagctg cagagacttt gagtgaagta gcctaagagg attctttaga
     2461 gaatccgtat tgaatttgtg tggtatgtca ctcagaaaga atcgtaatgg gtatattgat
     2521 aaattttaaa attggtatat ttgaaataaa gttgaatatt atatatagtt atgtgagtgt
     2581 ttatatatgt gtgtgtttat attgtttatc ttctccctat ggattaaaac tgaatttcat
     2641 aattataaga ggttattctg aagtggaaaa atttaactca gtattaaatc taaggagaat
     2701 ggcctaatat agtaaaactc tcatctggca ttatcaggga atcaagtcta atctattcat
     2761 gtcacttcac acagaagaaa acatcagtat gtcagagagc acactgggga atatgcacaa
     2821 gattatccca agccagaggc ctcacggcct acctggccag cctgggctga gaggatcact
     2881 atctcagcac actatttggg aaatggatca aatcacactt ttagtaaatg ttatcactct
     2941 atagcataag aaataattat tttttattta tataaaaggc tatagtataa aatatatgta
     3001 tagtaattaa atgaacactt gtgaacctaa tagccatatg aagaaaataa catttctaat
     3061 atctttggat gccccatgta ctaatgacag ttatgctttt gcattttctt gaattttatg
     3121 tttatttatc tttcctctgt cattatttat aattttatca cacatggctg tatcctttac
     3181 atgttttggc attatgtatt tttgaacttt ttgtaaagac aatcatacca tgtgtaattt
     3241 tcagggactt gatttttttc attgactttt aagggttcaa atatattatc actgtggctg
     3301 tagtttgcca tattttgctg atatagagca ttcattcaca tgagggtagg attcagggtc
     3361 catcaagaca gagaaaacat acagtaatgt gaatagggaa agttaatatg aagaattatt
     3421 aattgttaca gcattggaac aatgaaatat tgtctagtaa tatgtaaaga gaagtctcaa
     3481 gaatatgtga tgagcagatg taaggaattg ctcttgtctc catggtgaat ttggagcagc
     3541 caatgaagag tcccctcaca ttgtggcctc gctcaaagtt aagaagtcgc tgtagtgttg
     3601 cccttgaaga atctgcttca aattgacact tcagaactcc ccagaaactt gtcttctggg
     3661 ccaatgtgta aagctgttta tgaagaaatg tcaagccaga ggggctctac tacaaatttg
     3721 gcaaaggaca atttcaggag aagctcttgg ccgctgggtt ctcctggcca ccatgaactt
     3781 caggaagtgg gtgccatagc agcagcctga actacagaat ctgggcactg gtgtagctct
     3841 gtatgccctc cgtgtcagat gctggagatg tcatttgcat tgccagagtt tgccaagggt
     3901 gcacacagaa agcagattga aaagcaccct cttggaacat ctctccaatg ccttctactc
     3961 acaaagttta acatcattaa cacgtgacaa agaagaacta tttaatgggc ccagatctat
     4021 ttatgaagac aatcaagtgg gagtttggag tggataaccc aaatttggat aactggtgaa
     4081 taataaaatg tatttatttc tgctggtgta
//
