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NRAM4_ARATH
ID   NRAM4_ARATH             Reviewed;         512 AA.
AC   Q9FN18; Q0WWH0; Q9SNV8;
DT   11-APR-2003, integrated into UniProtKB/Swiss-Prot.
DT   01-MAR-2001, sequence version 1.
DT   03-AUG-2022, entry version 138.
DE   RecName: Full=Metal transporter Nramp4;
DE            Short=AtNramp4;
GN   Name=NRAMP4; OrderedLocusNames=At5g67330; ORFNames=K8K14.5;
OS   Arabidopsis thaliana (Mouse-ear cress).
OC   Eukaryota; Viridiplantae; Streptophyta; Embryophyta; Tracheophyta;
OC   Spermatophyta; Magnoliopsida; eudicotyledons; Gunneridae; Pentapetalae;
OC   rosids; malvids; Brassicales; Brassicaceae; Camelineae; Arabidopsis.
OX   NCBI_TaxID=3702;
RN   [1]
RP   NUCLEOTIDE SEQUENCE [MRNA], FUNCTION, AND INDUCTION.
RC   STRAIN=cv. Columbia;
RX   PubMed=10781110; DOI=10.1073/pnas.97.9.4991;
RA   Thomine S., Wang R., Ward J.M., Crawford N.M., Schroeder J.I.;
RT   "Cadmium and iron transport by members of a plant metal transporter family
RT   in Arabidopsis with homology to Nramp genes.";
RL   Proc. Natl. Acad. Sci. U.S.A. 97:4991-4996(2000).
RN   [2]
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=cv. Columbia;
RX   PubMed=9501997; DOI=10.1093/dnares/4.6.401;
RA   Nakamura Y., Sato S., Kaneko T., Kotani H., Asamizu E., Miyajima N.,
RA   Tabata S.;
RT   "Structural analysis of Arabidopsis thaliana chromosome 5. III. Sequence
RT   features of the regions of 1,191,918 bp covered by seventeen physically
RT   assigned P1 clones.";
RL   DNA Res. 4:401-414(1997).
RN   [3]
RP   GENOME REANNOTATION.
RC   STRAIN=cv. Columbia;
RX   PubMed=27862469; DOI=10.1111/tpj.13415;
RA   Cheng C.Y., Krishnakumar V., Chan A.P., Thibaud-Nissen F., Schobel S.,
RA   Town C.D.;
RT   "Araport11: a complete reannotation of the Arabidopsis thaliana reference
RT   genome.";
RL   Plant J. 89:789-804(2017).
RN   [4]
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA].
RC   STRAIN=cv. Columbia;
RX   PubMed=14593172; DOI=10.1126/science.1088305;
RA   Yamada K., Lim J., Dale J.M., Chen H., Shinn P., Palm C.J., Southwick A.M.,
RA   Wu H.C., Kim C.J., Nguyen M., Pham P.K., Cheuk R.F., Karlin-Newmann G.,
RA   Liu S.X., Lam B., Sakano H., Wu T., Yu G., Miranda M., Quach H.L.,
RA   Tripp M., Chang C.H., Lee J.M., Toriumi M.J., Chan M.M., Tang C.C.,
RA   Onodera C.S., Deng J.M., Akiyama K., Ansari Y., Arakawa T., Banh J.,
RA   Banno F., Bowser L., Brooks S.Y., Carninci P., Chao Q., Choy N., Enju A.,
RA   Goldsmith A.D., Gurjal M., Hansen N.F., Hayashizaki Y., Johnson-Hopson C.,
RA   Hsuan V.W., Iida K., Karnes M., Khan S., Koesema E., Ishida J., Jiang P.X.,
RA   Jones T., Kawai J., Kamiya A., Meyers C., Nakajima M., Narusaka M.,
RA   Seki M., Sakurai T., Satou M., Tamse R., Vaysberg M., Wallender E.K.,
RA   Wong C., Yamamura Y., Yuan S., Shinozaki K., Davis R.W., Theologis A.,
RA   Ecker J.R.;
RT   "Empirical analysis of transcriptional activity in the Arabidopsis
RT   genome.";
RL   Science 302:842-846(2003).
RN   [5]
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA] OF 4-512.
RC   STRAIN=cv. Columbia;
RA   Totoki Y., Seki M., Ishida J., Nakajima M., Enju A., Kamiya A.,
RA   Narusaka M., Shin-i T., Nakagawa M., Sakamoto N., Oishi K., Kohara Y.,
RA   Kobayashi M., Toyoda A., Sakaki Y., Sakurai T., Iida K., Akiyama K.,
RA   Satou M., Toyoda T., Konagaya A., Carninci P., Kawai J., Hayashizaki Y.,
RA   Shinozaki K.;
RT   "Large-scale analysis of RIKEN Arabidopsis full-length (RAFL) cDNAs.";
RL   Submitted (JUL-2006) to the EMBL/GenBank/DDBJ databases.
RN   [6]
RP   FUNCTION, SUBCELLULAR LOCATION, TISSUE SPECIFICITY, INDUCTION, AND
RP   DISRUPTION PHENOTYPE.
RC   STRAIN=cv. Wassilewskija;
RX   PubMed=16270029; DOI=10.1038/sj.emboj.7600864;
RA   Lanquar V., Lelievre F., Bolte S., Hames C., Alcon C., Neumann D.,
RA   Vansuyt G., Curie C., Schroeder A., Kraemer U., Barbier-Brygoo H.,
RA   Thomine S.;
RT   "Mobilization of vacuolar iron by AtNRAMP3 and AtNRAMP4 is essential for
RT   seed germination on low iron.";
RL   EMBO J. 24:4041-4051(2005).
RN   [7]
RP   FUNCTION.
RX   PubMed=20181755; DOI=10.1104/pp.109.150946;
RA   Lanquar V., Ramos M.S., Lelievre F., Barbier-Brygoo H., Krieger-Liszkay A.,
RA   Kraemer U., Thomine S.;
RT   "Export of vacuolar manganese by AtNRAMP3 and AtNRAMP4 is required for
RT   optimal photosynthesis and growth under manganese deficiency.";
RL   Plant Physiol. 152:1986-1999(2010).
CC   -!- FUNCTION: Vacuolar metal transporter involved in intracellular metal
CC       homeostasis. Can transport iron (Fe), manganese (Mn) and cadmium (Cd).
CC       Regulates metal accumulation under Fe starvation. Acts redundantly with
CC       NRAMP3 to mobilize vacuolar Fe and provide sufficient Fe during seed
CC       germination. In association with NRAMP3, required for optimal growth
CC       and photosynthesis under Mn deficiency. Exports Mn from vacuoles in
CC       leaf mesophyll cells, making Mn available for functional photosystem II
CC       in chloroplasts. {ECO:0000269|PubMed:10781110,
CC       ECO:0000269|PubMed:16270029, ECO:0000269|PubMed:20181755}.
CC   -!- SUBCELLULAR LOCATION: Vacuole membrane {ECO:0000269|PubMed:16270029};
CC       Multi-pass membrane protein {ECO:0000269|PubMed:16270029}.
CC   -!- TISSUE SPECIFICITY: Expressed in vascular tissues.
CC       {ECO:0000269|PubMed:16270029}.
CC   -!- INDUCTION: By iron starvation. {ECO:0000269|PubMed:10781110,
CC       ECO:0000269|PubMed:16270029}.
CC   -!- DISRUPTION PHENOTYPE: No visible phenotype under normal growth
CC       condition, but slightly enhanced resistance of root growth in presence
CC       of Cd and increased accumulation of Mn and Zn under Fe starvation.
CC       {ECO:0000269|PubMed:16270029}.
CC   -!- SIMILARITY: Belongs to the NRAMP (TC 2.A.55) family. {ECO:0000305}.
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DR   EMBL; AF202540; AAF13279.1; -; mRNA.
DR   EMBL; AB007645; BAB09018.1; -; Genomic_DNA.
DR   EMBL; CP002688; AED98329.1; -; Genomic_DNA.
DR   EMBL; AY136467; AAM97132.1; -; mRNA.
DR   EMBL; BT003406; AAO30069.1; -; mRNA.
DR   EMBL; AK226381; BAE98528.1; -; mRNA.
DR   RefSeq; NP_201534.1; NM_126133.4.
DR   AlphaFoldDB; Q9FN18; -.
DR   SMR; Q9FN18; -.
DR   BioGRID; 22110; 7.
DR   IntAct; Q9FN18; 5.
DR   STRING; 3702.AT5G67330.1; -.
DR   TCDB; 2.A.55.2.18; the metal ion (mn(2+)-iron) transporter (nramp) family.
DR   PaxDb; Q9FN18; -.
DR   PRIDE; Q9FN18; -.
DR   ProteomicsDB; 239057; -.
DR   EnsemblPlants; AT5G67330.1; AT5G67330.1; AT5G67330.
DR   GeneID; 836868; -.
DR   Gramene; AT5G67330.1; AT5G67330.1; AT5G67330.
DR   KEGG; ath:AT5G67330; -.
DR   Araport; AT5G67330; -.
DR   TAIR; locus:2158242; AT5G67330.
DR   eggNOG; KOG1291; Eukaryota.
DR   HOGENOM; CLU_020088_5_1_1; -.
DR   InParanoid; Q9FN18; -.
DR   OMA; SHKEPQR; -.
DR   OrthoDB; 666470at2759; -.
DR   PhylomeDB; Q9FN18; -.
DR   PRO; PR:Q9FN18; -.
DR   Proteomes; UP000006548; Chromosome 5.
DR   ExpressionAtlas; Q9FN18; baseline and differential.
DR   Genevisible; Q9FN18; AT.
DR   GO; GO:0016021; C:integral component of membrane; IEA:UniProtKB-KW.
DR   GO; GO:0000325; C:plant-type vacuole; HDA:TAIR.
DR   GO; GO:0009536; C:plastid; HDA:TAIR.
DR   GO; GO:0005774; C:vacuolar membrane; IDA:TAIR.
DR   GO; GO:0005773; C:vacuole; HDA:TAIR.
DR   GO; GO:0015086; F:cadmium ion transmembrane transporter activity; IBA:GO_Central.
DR   GO; GO:0005384; F:manganese ion transmembrane transporter activity; IBA:GO_Central.
DR   GO; GO:0042742; P:defense response to bacterium; IMP:TAIR.
DR   GO; GO:0055072; P:iron ion homeostasis; IMP:TAIR.
DR   GO; GO:0006828; P:manganese ion transport; IMP:TAIR.
DR   GO; GO:2000379; P:positive regulation of reactive oxygen species metabolic process; IGI:TAIR.
DR   GO; GO:0010039; P:response to iron ion; IEP:TAIR.
DR   GO; GO:0009624; P:response to nematode; HEP:TAIR.
DR   HAMAP; MF_00221; NRAMP; 1.
DR   InterPro; IPR001046; NRAMP_fam.
DR   PANTHER; PTHR11706; PTHR11706; 1.
DR   Pfam; PF01566; Nramp; 1.
DR   PRINTS; PR00447; NATRESASSCMP.
DR   TIGRFAMs; TIGR01197; nramp; 1.
PE   2: Evidence at transcript level;
KW   Ion transport; Iron; Iron transport; Membrane; Reference proteome;
KW   Transmembrane; Transmembrane helix; Transport; Vacuole.
FT   CHAIN           1..512
FT                   /note="Metal transporter Nramp4"
FT                   /id="PRO_0000212601"
FT   TRANSMEM        52..72
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        80..100
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        129..149
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        161..181
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        189..209
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        235..255
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        277..297
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        323..343
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        371..391
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        402..422
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        440..460
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   TRANSMEM        468..488
FT                   /note="Helical"
FT                   /evidence="ECO:0000255"
FT   CONFLICT        246
FT                   /note="H -> N (in Ref. 5; BAE98528)"
FT                   /evidence="ECO:0000305"
FT   CONFLICT        467
FT                   /note="L -> V (in Ref. 1; AAF13279)"
FT                   /evidence="ECO:0000305"
SQ   SEQUENCE   512 AA;  56385 MW;  89F2CE2359DF3324 CRC64;
     MSETDRERPL LASEERAYEE TEKVLIVGID EEEDADYDDD PGNSPKFSWK KLWLFTGPGF
     LMSIAFLDPG NLESDLQAGA IAGYSLIWLL MWATAIGLLI QLLSARLGVA TGRHLAELCR
     EEYPTWARMV LWIMAEIALI GADIQEVIGS AIAIKILSNG LVPLWAGVVI TALDCFIFLF
     LENYGIRKLE AVFAILIATM ALAFAWMFGQ TKPSGTELLV GALVPKLSSR TIKQAVGIVG
     CIIMPHNVFL HSALVQSREV DPKKRFRVKE ALKYYSIEST GALAVSFIIN VFVTTVFAKS
     FYGTEIADTI GLANAGQYLQ DKYGGGFFPI LYIWAIGVLA AGQSSTITGT YAGQFIMGGF
     LNLKMKKWVR ALITRSCAII PTMIVALVFD SSDSMLDELN EWLNVLQSVQ IPFAVIPLLC
     LVSNEQIMGS FKIQPLVQTI SWIVAALVIA INGYLMVDFF SGAATNLILL VPVIIFAIAY
     VVFVLYLISR GLTYTPWQLV ASSHKEPQRD DE
 
 
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