{"entryType":"UniProtKB reviewed (Swiss-Prot)","primaryAccession":"P36223","uniProtkbId":"GLND_AZOVI","entryAudit":{"firstPublicDate":"1994-06-01","lastAnnotationUpdateDate":"2026-09-02","lastSequenceUpdateDate":"1994-06-01","entryVersion":108,"sequenceVersion":1},"annotationScore":4.0,"organism":{"scientificName":"Azotobacter vinelandii","taxonId":354,"lineage":["Bacteria","Pseudomonadati","Pseudomonadota","Gammaproteobacteria","Pseudomonadales","Pseudomonadaceae","Azotobacter"]},"proteinExistence":"1: Evidence at protein level","proteinDescription":{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"Bifunctional uridylyltransferase/uridylyl-removing enzyme"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"UTase/UR"}]},"alternativeNames":[{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"Bifunctional [protein-PII] modification enzyme"}},{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"Bifunctional nitrogen sensor protein"}}],"includes":[{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"[Protein-PII] uridylyltransferase"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"PII uridylyltransferase"},{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"UTase"}],"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"2.7.7.59"}]}},{"recommendedName":{"fullName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"[Protein-PII]-UMP uridylyl-removing enzyme"},"shortNames":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"UR"}],"ecNumbers":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"3.1.4.-"}]}}]},"genes":[{"geneName":{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"glnD"},"synonyms":[{"value":"nfrX"}]}],"comments":[{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11320130"},{"evidenceCode":"ECO:0000269","source":"PubMed","id":"1683868"}],"value":"Modifies, by uridylylation and deuridylylation, the PII regulatory protein GlnK, in response to the nitrogen status of the cell that GlnD senses through the glutamine level. Under low glutamine levels, catalyzes the conversion of the PII protein and UTP to PII-UMP and PPi, while under higher glutamine levels, GlnD hydrolyzes PII-UMP to PII and UMP (deuridylylation). Thus, controls uridylylation state and activity of the PII protein, and plays an important role in the regulation of nitrogen fixation and metabolism"}],"commentType":"FUNCTION"},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"[protein-PII]-L-tyrosine + UTP = [protein-PII]-uridylyl-L-tyrosine + diphosphate","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:13673"},{"database":"Rhea","id":"RHEA-COMP:12147"},{"database":"Rhea","id":"RHEA-COMP:12148"},{"database":"ChEBI","id":"CHEBI:33019"},{"database":"ChEBI","id":"CHEBI:46398"},{"database":"ChEBI","id":"CHEBI:46858"},{"database":"ChEBI","id":"CHEBI:90602"}],"ecNumber":"2.7.7.59","evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}]}},{"commentType":"CATALYTIC ACTIVITY","reaction":{"name":"[protein-PII]-uridylyl-L-tyrosine + H2O = [protein-PII]-L-tyrosine + UMP + H(+)","reactionCrossReferences":[{"database":"Rhea","id":"RHEA:48600"},{"database":"Rhea","id":"RHEA-COMP:12147"},{"database":"Rhea","id":"RHEA-COMP:12148"},{"database":"ChEBI","id":"CHEBI:15377"},{"database":"ChEBI","id":"CHEBI:15378"},{"database":"ChEBI","id":"CHEBI:46858"},{"database":"ChEBI","id":"CHEBI:57865"},{"database":"ChEBI","id":"CHEBI:90602"}],"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}]}},{"commentType":"COFACTOR","cofactors":[{"name":"Mg(2+)","evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"cofactorCrossReference":{"database":"ChEBI","id":"CHEBI:18420"}}]},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"Uridylyltransferase (UTase) activity is inhibited by glutamine, while glutamine activates uridylyl-removing (UR) activity"}],"commentType":"ACTIVITY REGULATION"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"Has four distinct domains: an N-terminal nucleotidyltransferase (NT) domain responsible for UTase activity, a central HD domain that encodes UR activity, and two C-terminal ACT domains that seem to have a role in glutamine sensing"}],"commentType":"DOMAIN"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000269","source":"PubMed","id":"11320130"}],"value":"Cells lacking this gene are not viable unless a second mutation occurs, either spontaneously or by design, resulting in the inability of glutamine synthetase (GS) to be adenylylated"}],"commentType":"DISRUPTION PHENOTYPE"},{"texts":[{"evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}],"value":"Belongs to the GlnD family"}],"commentType":"SIMILARITY"}],"features":[{"type":"Chain","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":899,"modifier":"EXACT"}},"description":"Bifunctional uridylyltransferase/uridylyl-removing enzyme","featureId":"PRO_0000192717"},{"type":"Domain","location":{"start":{"value":461,"modifier":"EXACT"},"end":{"value":583,"modifier":"EXACT"}},"description":"HD","evidences":[{"evidenceCode":"ECO:0000255","source":"PROSITE-ProRule","id":"PRU01175"}]},{"type":"Domain","location":{"start":{"value":706,"modifier":"EXACT"},"end":{"value":789,"modifier":"EXACT"}},"description":"ACT 1","evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}]},{"type":"Domain","location":{"start":{"value":816,"modifier":"EXACT"},"end":{"value":899,"modifier":"EXACT"}},"description":"ACT 2","evidences":[{"evidenceCode":"ECO:0000255","source":"HAMAP-Rule","id":"MF_00277"}]},{"type":"Region","location":{"start":{"value":1,"modifier":"EXACT"},"end":{"value":342,"modifier":"EXACT"}},"description":"Uridylyltransferase"},{"type":"Region","location":{"start":{"value":343,"modifier":"EXACT"},"end":{"value":705,"modifier":"EXACT"}},"description":"Uridylyl-removing"}],"keywords":[{"id":"KW-0378","category":"Molecular function","name":"Hydrolase"},{"id":"KW-0460","category":"Ligand","name":"Magnesium"},{"id":"KW-0511","category":"Molecular function","name":"Multifunctional enzyme"},{"id":"KW-0535","category":"Biological process","name":"Nitrogen fixation"},{"id":"KW-0548","category":"Molecular function","name":"Nucleotidyltransferase"},{"id":"KW-0677","category":"Domain","name":"Repeat"},{"id":"KW-0808","category":"Molecular function","name":"Transferase"}],"references":[{"referenceNumber":1,"citation":{"id":"1683868","citationType":"journal article","authors":["Contreras A.","Drummond M.","Bali A.","Blanco G.","Garcia E.","Bush G.","Kennedy C.","Merrick M."],"citationCrossReferences":[{"database":"PubMed","id":"1683868"},{"database":"DOI","id":"10.1128/jb.173.24.7741-7749.1991"}],"title":"The product of the nitrogen fixation regulatory gene nfrX of Azotobacter vinelandii is functionally and structurally homologous to the uridylyltransferase encoded by glnD in enteric bacteria.","publicationDate":"1991","journal":"J. Bacteriol.","firstPage":"7741","lastPage":"7749","volume":"173"},"referencePositions":["NUCLEOTIDE SEQUENCE [GENOMIC DNA]","FUNCTION","COMPLEMENTATION OF E.COLI GLND MUTANT","ROLE IN NITROGEN FIXATION"],"referenceComments":[{"value":"OP / UW136","type":"STRAIN"}]},{"referenceNumber":2,"citation":{"id":"11320130","citationType":"journal article","authors":["Colnaghi R.","Rudnick P.","He L.","Green A.","Yan D.","Larson E.","Kennedy C."],"citationCrossReferences":[{"database":"PubMed","id":"11320130"},{"database":"DOI","id":"10.1099/00221287-147-5-1267"}],"title":"Lethality of glnD null mutations in Azotobacter vinelandii is suppressible by prevention of glutamine synthetase adenylylation.","publicationDate":"2001","journal":"Microbiology","firstPage":"1267","lastPage":"1276","volume":"147"},"referencePositions":["FUNCTION AS PII URIDYLYLTRANSFERASE","DISRUPTION PHENOTYPE"],"referenceComments":[{"value":"OP / UW136","type":"STRAIN"}]}],"uniProtKBCrossReferences":[{"database":"EMBL","id":"X59610","properties":[{"key":"ProteinId","value":"CAA42173.1"},{"key":"Status","value":"-"},{"key":"MoleculeType","value":"Genomic_DNA"}]},{"database":"PIR","id":"S24223","properties":[{"key":"EntryName","value":"S24223"}]},{"database":"RefSeq","id":"WP_012702416.1","properties":[{"key":"NucleotideSequenceId","value":"NZ_AP043912.1"}]},{"database":"AlphaFoldDB","id":"P36223","properties":[{"key":"Description","value":"-"}]},{"database":"SMR","id":"P36223","properties":[{"key":"Description","value":"-"}]},{"database":"GeneID","id":"309096015","properties":[{"key":"Description","value":"-"}]},{"database":"OMA","id":"GLMQFDL","properties":[{"key":"Fingerprint","value":"-"}]},{"database":"GO","id":"GO:0008773","properties":[{"key":"GoTerm","value":"F:[protein-PII] uridylyltransferase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-UniRule"}]},{"database":"GO","id":"GO:0008081","properties":[{"key":"GoTerm","value":"F:phosphoric diester hydrolase activity"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-UniRule"}]},{"database":"GO","id":"GO:0006808","properties":[{"key":"GoTerm","value":"P:regulation of nitrogen utilization"},{"key":"GoEvidenceType","value":"IEA:UniProtKB-UniRule"}]},{"database":"CDD","id":"cd04899","properties":[{"key":"EntryName","value":"ACT_ACR-UUR-like_2"},{"key":"MatchStatus","value":"1"}]},{"database":"CDD","id":"cd04900","properties":[{"key":"EntryName","value":"ACT_UUR-like_1"},{"key":"MatchStatus","value":"1"}]},{"database":"CDD","id":"cd00077","properties":[{"key":"EntryName","value":"HDc"},{"key":"MatchStatus","value":"1"}]},{"database":"CDD","id":"cd05401","properties":[{"key":"EntryName","value":"NT_GlnE_GlnD_like"},{"key":"MatchStatus","value":"1"}]},{"database":"FunFam","id":"1.10.3090.10:FF:000005","properties":[{"key":"EntryName","value":"Bifunctional uridylyltransferase/uridylyl-removing enzyme"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"3.30.460.10","properties":[{"key":"EntryName","value":"Beta Polymerase, domain 2"},{"key":"MatchStatus","value":"1"}]},{"database":"Gene3D","id":"1.10.3090.10","properties":[{"key":"EntryName","value":"cca-adding enzyme, domain 2"},{"key":"MatchStatus","value":"1"}]},{"database":"HAMAP","id":"MF_00277","properties":[{"key":"EntryName","value":"PII_uridylyl_transf"},{"key":"MatchStatus","value":"1"}]},{"database":"InterPro","id":"IPR045865","properties":[{"key":"EntryName","value":"ACT-like_dom_sf"}]},{"database":"InterPro","id":"IPR002912","properties":[{"key":"EntryName","value":"ACT_dom"}]},{"database":"InterPro","id":"IPR010043","properties":[{"key":"EntryName","value":"GlnD"}]},{"database":"InterPro","id":"IPR060736","properties":[{"key":"EntryName","value":"GlnD_3rd"}]},{"database":"InterPro","id":"IPR060737","properties":[{"key":"EntryName","value":"GlnD_helical"}]},{"database":"InterPro","id":"IPR003607","properties":[{"key":"EntryName","value":"HD/PDEase_dom"}]},{"database":"InterPro","id":"IPR006674","properties":[{"key":"EntryName","value":"HD_domain"}]},{"database":"InterPro","id":"IPR043519","properties":[{"key":"EntryName","value":"NT_sf"}]},{"database":"InterPro","id":"IPR013546","properties":[{"key":"EntryName","value":"PII_UdlTrfase/GS_AdlTrfase"}]},{"database":"InterPro","id":"IPR002934","properties":[{"key":"EntryName","value":"Polymerase_NTP_transf_dom"}]},{"database":"NCBIfam","id":"NF001366","properties":[{"key":"EntryName","value":"PRK00275.1"},{"key":"MatchStatus","value":"1"}]},{"database":"NCBIfam","id":"TIGR01693","properties":[{"key":"EntryName","value":"UTase_glnD"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47320","properties":[{"key":"EntryName","value":"BIFUNCTIONAL URIDYLYLTRANSFERASE/URIDYLYL-REMOVING ENZYME"},{"key":"MatchStatus","value":"1"}]},{"database":"PANTHER","id":"PTHR47320:SF1","properties":[{"key":"EntryName","value":"BIFUNCTIONAL URIDYLYLTRANSFERASE_URIDYLYL-REMOVING ENZYME"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF01842","properties":[{"key":"EntryName","value":"ACT"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF27444","properties":[{"key":"EntryName","value":"GlnD_3rd"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF27446","properties":[{"key":"EntryName","value":"GlnD_5th"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF08335","properties":[{"key":"EntryName","value":"GlnD_UR_UTase"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF01966","properties":[{"key":"EntryName","value":"HD"},{"key":"MatchStatus","value":"1"}]},{"database":"Pfam","id":"PF01909","properties":[{"key":"EntryName","value":"NTP_transf_2"},{"key":"MatchStatus","value":"1"}]},{"database":"PIRSF","id":"PIRSF006288","properties":[{"key":"EntryName","value":"PII_uridyltransf"},{"key":"MatchStatus","value":"1"}]},{"database":"SMART","id":"SM00471","properties":[{"key":"EntryName","value":"HDc"},{"key":"MatchStatus","value":"1"}]},{"database":"SUPFAM","id":"SSF55021","properties":[{"key":"EntryName","value":"ACT-like"},{"key":"MatchStatus","value":"1"}]},{"database":"SUPFAM","id":"SSF109604","properties":[{"key":"EntryName","value":"HD-domain/PDEase-like"},{"key":"MatchStatus","value":"1"}]},{"database":"SUPFAM","id":"SSF81301","properties":[{"key":"EntryName","value":"Nucleotidyltransferase"},{"key":"MatchStatus","value":"1"}]},{"database":"SUPFAM","id":"SSF81593","properties":[{"key":"EntryName","value":"Nucleotidyltransferase substrate binding subunit/domain"},{"key":"MatchStatus","value":"1"}]},{"database":"PROSITE","id":"PS51671","properties":[{"key":"EntryName","value":"ACT"},{"key":"MatchStatus","value":"2"}]},{"database":"PROSITE","id":"PS51831","properties":[{"key":"EntryName","value":"HD"},{"key":"MatchStatus","value":"1"}]}],"sequence":{"value":"MPQVDPDLFDPGQFQAELALKSSPIPAYKKALRCAREVLDARFQEGRDIRRLIEDRAWFVDQILALAWNRFDWSEDADIALIAVGGYGRGELHPYSDIDLLILMDGADHEVFREPIEGFLTLLWDIGLEVGQSVRSLAECAEEAQADLTVITNLMESRTIAGPEHLRQRMQEVTSAQRMWPSRAFFLAKRDEQKTRHARYNDTEYNLEPNVKGSPGGLRDIQTLLWIARRQFGTINLHAMVGQGFLLESEYTLLASSQEFLWKVRYALHMLAGRAEDRLLFDLQRQIAGLLGYEDSDAKLAVERFMQKYYRVVLGIAELTELVFQHFEEVILPGDAAGRVEPLNERFQVRDGYLEVTHAGVFQETPSALLEIFVLLARRPEIRGVRADTIRLLRDHRYLIDDAFRRDPHNTGLFIELFKSRQGIHRNLRRMNRYGILGRYLPEFGHIVGQMQHDLFHIYTVDAHTLNLIKNLRKLFWPELAEKYPLASKLIEKLPKPELIYLAGLYHDIGKGRGGDHSELGAADALAFCQRHDLPAMDTQLIVWLVRNHLLMSTTAQRKDLSDPQVIFDFAQKVRDQTYLDYLYVLTVADINATNPTLWNSWRASLLRQLYTETKHALRRGLEQPVGREEQIRQTQKAALDILVRSGTDPDDAEHLWTQLGDDYFLRHTSSDIAWHTEAILQHPSSGGPLVLIKETTQREFEGATQIFIYAPDQHDFFAVTVAAMDQLNLSIHDARVITSTSQFTLDTYIVLDADGGSIGNNPARIQEIRQGLVEALRNPADYPTIIQRRVPRQLKHFAFAPQVTIQNDALRPVTILEIIAPDRPGLLARIGKIFLDFDLSLQNAKIATLGERVEDVFFVTDAHNQPLSDPELCARLQLAIAEQLADGDSYIQPSRISI","length":899,"molWeight":102580,"crc64":"96AF63BC5EDE3C9D","md5":"502C767773ACAA0A624C0FE5175B4D66"},"extraAttributes":{"countByCommentType":{"FUNCTION":1,"CATALYTIC ACTIVITY":2,"COFACTOR":1,"ACTIVITY REGULATION":1,"DOMAIN":1,"DISRUPTION PHENOTYPE":1,"SIMILARITY":1},"countByFeatureType":{"Chain":1,"Domain":3,"Region":2},"uniParcId":"UPI000012B724"}}