ID   Q1K9H2_I33A0            Unreviewed;       498 AA.
AC   Q1K9H2; B4URF1; Q1I2B5;
DT   30-MAY-2006, integrated into UniProtKB/TrEMBL.
DT   30-MAY-2006, sequence version 1.
DT   27-MAR-2024, entry version 114.
DE   RecName: Full=Nucleoprotein {ECO:0000256|HAMAP-Rule:MF_04070};
DE   AltName: Full=Nucleocapsid protein {ECO:0000256|HAMAP-Rule:MF_04070};
DE            Short=Protein N {ECO:0000256|HAMAP-Rule:MF_04070};
GN   Name=NP {ECO:0000256|HAMAP-Rule:MF_04070,
GN   ECO:0000256|RuleBase:RU361251, ECO:0000313|EMBL:ABF21292.1};
OS   Influenza A virus (strain A/Wilson-Smith/1933 H1N1) (Influenza A virus
OS   (strain A/WS/1933 H1N1)).
OC   Viruses; Riboviria; Orthornavirae; Negarnaviricota; Polyploviricotina;
OC   Insthoviricetes; Articulavirales; Orthomyxoviridae; Alphainfluenzavirus;
OC   Alphainfluenzavirus influenzae; Influenza A virus.
OX   NCBI_TaxID=381518 {ECO:0000313|EMBL:ABF21292.1, ECO:0000313|Proteomes:UP000131702};
OH   NCBI_TaxID=8782; Aves (birds).
OH   NCBI_TaxID=9606; Homo sapiens (Human).
OH   NCBI_TaxID=9823; Sus scrofa (Pig).
RN   [1] {ECO:0007829|PDB:2IQH}
RP   X-RAY CRYSTALLOGRAPHY (3.20 ANGSTROMS) OF 8-498.
RX   PubMed=17151603; DOI=10.1038/nature05379;
RA   Ye Q., Krug R.M., Tao Y.J.;
RT   "The mechanism by which influenza A virus nucleoprotein forms oligomers and
RT   binds RNA.";
RL   Nature 444:1078-1082(2006).
RN   [2] {ECO:0000313|Proteomes:UP000131702}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RA   Ghedin E., Sengamalay N., Zaborsky J., Feldblyum T., Subbu V., Shumway M.,
RA   Sitz J., Katzel D., Koo H., Salzberg S.L., Griesemer S., Ginocchio C.,
RA   StGeorge K., Bennett R., Taylor J., Bao Y., Bolotov P., Dernovoy D.,
RA   Kiryutin B., Lipman D.J., Tatusova T.;
RT   "The NIAID Influenza Genome Sequencing Project.";
RL   Submitted (MAR-2006) to the EMBL/GenBank/DDBJ databases.
RN   [3] {ECO:0000313|EMBL:ABF21292.1, ECO:0000313|Proteomes:UP000131702}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=A/Wilson-Smith/1933 {ECO:0000313|EMBL:ABF21292.1};
RA   Mbawuike I.N., Zhang Y., Yamada R.E., Nino D., Bui H.-H., Sette A.,
RA   Couch R.B.;
RT   "Complete Genome Sequencing And Analysis Of Selected Influenza Virus
RT   Vaccine Strains Spanning Six Decades (1933-1999).";
RL   Submitted (APR-2006) to the EMBL/GenBank/DDBJ databases.
RN   [4] {ECO:0000313|Proteomes:UP000131702}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=A/WSN/1933 TS61 {ECO:0000313|EMBL:ABF47959.1};
RG   The NIAID Influenza Genome Sequencing Consortium;
RL   Submitted (APR-2006) to the EMBL/GenBank/DDBJ databases.
RN   [5] {ECO:0000313|EMBL:ABF47959.1, ECO:0000313|Proteomes:UP000148454}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=A/WSN/1933 TS61 {ECO:0000313|EMBL:ABF47959.1};
RA   Spiro D., Ghedin E., Sengamalay N., Halpin R., Boyne A., Zaborsky J.,
RA   Feldblyum T., Subbu V., Sparenborg J., Shumway M., Sitz J., Katzel D.,
RA   Koo H., Salzberg S.L., Griesemer S., St George K., Bennett R., Taylor J.,
RA   Bennink J.R., Yewdell J.W., Bao Y., Bolotov P., Dernovoy D., Kiryutin B.,
RA   Lipman D.J., Tatusova T.;
RT   "The NIAID Influenza Genome Sequencing Project.";
RL   Submitted (MAY-2006) to the EMBL/GenBank/DDBJ databases.
RN   [6] {ECO:0000313|EMBL:ACF54602.1}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=A/WSN/1933 {ECO:0000313|EMBL:ACF54602.1};
RA   Spiro D., Halpin R., Boyne A., Bera J., Ghedin E., Hostetler J.,
RA   Fedorova N., Kim M., Zaborsky J., Overton L., Djuric K., Sarmiento M.,
RA   Sitz J., Katzel D., Webster R.G., Hoffmann E., Krauss S., Naeve C.,
RA   Bolotov P., Bao Y., Sanders R., Dernovoy D., Kiryutin B., Lipman D.J.,
RA   Tatusova T.;
RT   "The NIAID Influenza Genome Sequencing Project.";
RL   Submitted (JUL-2008) to the EMBL/GenBank/DDBJ databases.
RN   [7] {ECO:0000313|Proteomes:UP000113251}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RA   Spiro D., Halpin R., Boyne A., Bera J., Ghedin E., Hostetler J.,
RA   Fedorova N., Hine E., Overton L., Djuric K., Sarmiento M., Sitz J.,
RA   Katzel D., Manojkumar R., Devis R., Fulvini A., Silverman J., Le J.,
RA   Kilbourne E.D., Pokorny B., Bucher D., Orff E., Minieri J., Onodera S.,
RA   Huang L., Bao Y., Sanders R., Dernovoy D., Kiryutin B., Lipman D.J.,
RA   Tatusova T.;
RT   "The NIAID Influenza Genome Sequencing Project.";
RL   Submitted (JUL-2008) to the EMBL/GenBank/DDBJ databases.
RN   [8] {ECO:0000313|EMBL:ACF54602.1}
RP   NUCLEOTIDE SEQUENCE [LARGE SCALE GENOMIC DNA].
RC   STRAIN=A/WSN/1933 {ECO:0000313|EMBL:ACF54602.1};
RA   Ming Y.M., Sheng G.Y., Jing W., Wei Z.W.;
RL   Submitted (JUL-2008) to the EMBL/GenBank/DDBJ databases.
RN   [9] {ECO:0007829|PDB:3RO5}
RP   X-RAY CRYSTALLOGRAPHY (2.66 ANGSTROMS) OF 8-498.
RX   PubMed=21896751; DOI=10.1073/pnas.1107906108;
RA   Gerritz S.W., Cianci C., Kim S., Pearce B.C., Deminie C., Discotto L.,
RA   McAuliffe B., Minassian B.F., Shi S., Zhu S., Zhai W., Pendri A., Li G.,
RA   Poss M.A., Edavettal S., McDonnell P.A., Lewis H.A., Maskos K., Mortl M.,
RA   Kiefersauer R., Steinbacher S., Baldwin E.T., Metzler W., Bryson J.,
RA   Healy M.D., Philip T., Zoeckler M., Schartman R., Sinz M.,
RA   Leyva-Grado V.H., Hoffmann H.H., Langley D.R., Meanwell N.A., Krystal M.;
RT   "Inhibition of influenza virus replication via small molecules that induce
RT   the formation of higher-order nucleoprotein oligomers.";
RL   Proc. Natl. Acad. Sci. U.S.A. 108:15366-15371(2011).
RN   [10] {ECO:0007829|PDB:3TG6}
RP   X-RAY CRYSTALLOGRAPHY (3.00 ANGSTROMS) OF 8-498.
RA   Edavettal S.C., Pearce B.C., Lewis H.A., Schneeweis L., Deminie C.,
RA   Discotto L., Carpenter B., Yanchunas J., Langley D.R., Metzler W., Gao M.,
RA   Poss M.A., Maskos K., Mortl M., Keifersauer R., Steinbacher S., Healy M.,
RA   Phillip T., Zoeckler M., Schartman R., McAuliffe B., Minassian B.F.,
RA   Shi S., Zhu S., Pendri A., Li G., Sinz M., Meanwell N.A., Krystal M.,
RA   Gerritz S.W., Cianci C., Baldwin B.T., Bryson J.W., McDonnell P.A.;
RT   "Biophysical and Structural Characterization of a Novel Class of Influenza
RT   Virus Inhibitors.";
RL   Submitted (AUG-2011) to the PDB data bank.
RN   [11] {ECO:0007829|PDB:4IRY}
RP   X-RAY CRYSTALLOGRAPHY (2.80 ANGSTROMS) OF 1-221 AND 227-498.
RX   PubMed=23269829; DOI=10.1128/mBio.00467-12;
RA   Ye Q., Guu T.S., Mata D.A., Kuo R.L., Smith B., Krug R.M., Tao Y.J.;
RT   "Biochemical and structural evidence in support of a coherent model for the
RT   formation of the double-helical influenza A virus ribonucleoprotein.";
RL   MBio 4:e00467-e00412(2012).
RN   [12] {ECO:0007829|PDB:2YMN}
RP   STRUCTURE BY ELECTRON MICROSCOPY (20.00 ANGSTROMS).
RX   PubMed=23180774; DOI=10.1126/SCIENCE.1227270;
RA   Moeller A., Kirchdoerfer R.N., Potter C.S., Carragher B., Wilson I.A.;
RT   "Organization of the influenza virus replication machinery.";
RL   Science 338:1631-1634(2012).
RN   [13] {ECO:0007829|PDB:4BBL}
RP   STRUCTURE BY ELECTRON MICROSCOPY (18.00 ANGSTROMS) OF 8-498.
RX   PubMed=23180776; DOI=10.1126/SCIENCE.1228172;
RA   Arranz R., Coloma R., Chichon F.J., Conesa J.J., Carrascosa J.L.,
RA   Valpuesta J.M., Ortin J., Martin-Benito J.;
RT   "The structure of native influenza virion ribonucleoproteins.";
RL   Science 338:1634-1637(2012).
RN   [14] {ECO:0007829|PDB:4DYA, ECO:0007829|PDB:4DYB}
RP   X-RAY CRYSTALLOGRAPHY (2.40 ANGSTROMS) OF 8-498.
RA   Lewis H.A., Baldwin E.T., Steinbacher S., Maskos K., Mortl M.,
RA   Kiefersauer R., Edavettal S., McDonnell P.A., Pearce B.C., Langley D.R.;
RT   "To be determined.";
RL   Submitted (FEB-2012) to the PDB data bank.
RN   [15] {ECO:0007829|PDB:5TJW}
RP   X-RAY CRYSTALLOGRAPHY (3.23 ANGSTROMS).
RX   PubMed=27965447; DOI=10.1128/mBio.01569-16;
RA   Hanke L., Knockenhauer K.E., Brewer R.C., van Diest E., Schmidt F.I.,
RA   Schwartz T.U., Ploegh H.L.;
RT   "The Antiviral Mechanism of an Influenza A Virus Nucleoprotein-Specific
RT   Single-Domain Antibody Fragment. .";
RL   MBio 7:e01569-e01516(2016).
CC   -!- FUNCTION: Encapsidates the negative strand viral RNA, protecting it
CC       from nucleases. The encapsidated genomic RNA is termed the
CC       ribonucleoprotein (RNP) and serves as template for transcription and
CC       replication. The RNP needs to be localized in the host nucleus to start
CC       an infectious cycle, but is too large to diffuse through the nuclear
CC       pore complex. NP comprises at least 2 nuclear localization signals that
CC       are responsible for the active RNP import into the nucleus through
CC       cellular importin alpha/beta pathway. Later in the infection, nclear
CC       export of RNPs are mediated through viral proteins NEP interacting with
CC       M1 which binds nucleoproteins. It is possible that nucleoprotein binds
CC       directly host exportin-1/XPO1 and plays an active role in RNPs nuclear
CC       export. M1 interaction with RNP seems to hide nucleoprotein's nuclear
CC       localization signals. Soon after a virion infects a new cell, M1
CC       dissociates from the RNP under acidification of the virion driven by M2
CC       protein. Dissociation of M1 from RNP unmasks nucleoprotein's nuclear
CC       localization signals, targeting the RNP to the nucleus.
CC       {ECO:0000256|HAMAP-Rule:MF_04070}.
CC   -!- SUBUNIT: Homomultimerizes to form the nucleocapsid. May bind host
CC       exportin-1/XPO1. Binds to viral genomic RNA. Protein-RNA contacts are
CC       mediated by a combination of electrostatic interactions between
CC       positively charged residues and the phosphate backbone and planar
CC       interactions between aromatic side chains and bases.
CC       {ECO:0000256|HAMAP-Rule:MF_04070}.
CC   -!- INTERACTION:
CC       Q1K9H2; Q1K9H2: NP; NbExp=4; IntAct=EBI-6050688, EBI-6050688;
CC       Q1K9H2; Q8WV44: TRIM41; Xeno; NbExp=3; IntAct=EBI-6050688, EBI-725997;
CC   -!- SUBCELLULAR LOCATION: Virion {ECO:0000256|HAMAP-Rule:MF_04070,
CC       ECO:0000256|RuleBase:RU361251}. Host nucleus {ECO:0000256|HAMAP-
CC       Rule:MF_04070, ECO:0000256|RuleBase:RU361251}.
CC   -!- PTM: Late in virus-infected cells, may be cleaved from a 56-kDa protein
CC       to a 53-kDa protein by a cellular caspase. This cleavage might be a
CC       marker for the onset of apoptosis in infected cells or have a specific
CC       function in virus host interaction. {ECO:0000256|HAMAP-Rule:MF_04070}.
CC   -!- SIMILARITY: Belongs to the influenza viruses nucleoprotein family.
CC       {ECO:0000256|HAMAP-Rule:MF_04070, ECO:0000256|RuleBase:RU361251}.
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DR   EMBL; DQ508906; ABF21292.1; -; Genomic_RNA.
DR   EMBL; CY010791; ABF47959.1; -; Genomic_RNA.
DR   EMBL; CY034135; ACF54602.1; -; Viral_cRNA.
DR   PDB; 2IQH; X-ray; 3.20 A; A/B/C=8-498.
DR   PDB; 2YMN; EM; 20.00 A; A/B/C/D/E/F=1-498.
DR   PDB; 3RO5; X-ray; 2.66 A; A/B=8-498.
DR   PDB; 3TG6; X-ray; 3.00 A; A/B=8-498.
DR   PDB; 4BBL; EM; 18.00 A; A/B/C/D/E/F/G/H/I/J/K/L/M/N/O/P/Q/R/S/T/U/V/W/X=8-498.
DR   PDB; 4DYA; X-ray; 2.75 A; A/B=8-498.
DR   PDB; 4DYB; X-ray; 2.80 A; A/B=8-498.
DR   PDB; 4DYN; X-ray; 2.40 A; A/B=8-498.
DR   PDB; 4DYP; X-ray; 2.82 A; A/B=8-498.
DR   PDB; 4DYT; X-ray; 3.00 A; A/B/C=8-498.
DR   PDB; 4IRY; X-ray; 2.80 A; A/B=1-401, A/B=430-498.
DR   PDB; 5TJW; X-ray; 3.23 A; A=1-498.
DR   PDBsum; 2IQH; -.
DR   PDBsum; 2YMN; -.
DR   PDBsum; 3RO5; -.
DR   PDBsum; 3TG6; -.
DR   PDBsum; 4BBL; -.
DR   PDBsum; 4DYA; -.
DR   PDBsum; 4DYB; -.
DR   PDBsum; 4DYN; -.
DR   PDBsum; 4DYP; -.
DR   PDBsum; 4DYT; -.
DR   PDBsum; 4IRY; -.
DR   PDBsum; 5TJW; -.
DR   PDBsum; 6H9G; -.
DR   PDBsum; 6I7B; -.
DR   PDBsum; 6I85; -.
DR   PDBsum; 6J1U; -.
DR   SMR; Q1K9H2; -.
DR   DIP; DIP-60419N; -.
DR   DIP; DIP-61893N; -.
DR   IntAct; Q1K9H2; 113.
DR   ABCD; Q1K9H2; 6 sequenced antibodies.
DR   Proteomes; UP000113251; Genome.
DR   Proteomes; UP000131702; Genome.
DR   Proteomes; UP000148454; Genome.
DR   GO; GO:0019029; C:helical viral capsid; IEA:UniProtKB-UniRule.
DR   GO; GO:0042025; C:host cell nucleus; IEA:UniProtKB-SubCell.
DR   GO; GO:1990904; C:ribonucleoprotein complex; IEA:UniProtKB-KW.
DR   GO; GO:0019013; C:viral nucleocapsid; IEA:UniProtKB-UniRule.
DR   GO; GO:0042802; F:identical protein binding; IPI:IntAct.
DR   GO; GO:0003723; F:RNA binding; IEA:UniProtKB-UniRule.
DR   GO; GO:0005198; F:structural molecule activity; IEA:UniProtKB-UniRule.
DR   GO; GO:0046718; P:viral entry into host cell; IEA:UniProtKB-KW.
DR   GO; GO:0075732; P:viral penetration into host nucleus; IEA:UniProtKB-UniRule.
DR   HAMAP; MF_04070; INFV_NCAP; 1.
DR   InterPro; IPR002141; Flu_NP.
DR   Pfam; PF00506; Flu_NP; 1.
DR   SUPFAM; SSF161003; flu NP-like; 1.
PE   1: Evidence at protein level;
KW   3D-structure {ECO:0007829|PDB:2IQH, ECO:0007829|PDB:2YMN};
KW   Capsid protein {ECO:0000256|ARBA:ARBA00022561, ECO:0000256|HAMAP-
KW   Rule:MF_04070};
KW   Helical capsid protein {ECO:0000256|ARBA:ARBA00022497, ECO:0000256|HAMAP-
KW   Rule:MF_04070};
KW   Host nucleus {ECO:0000256|ARBA:ARBA00022562, ECO:0000256|HAMAP-
KW   Rule:MF_04070}; Host-virus interaction {ECO:0000256|HAMAP-Rule:MF_04070};
KW   Ribonucleoprotein {ECO:0000256|ARBA:ARBA00023274, ECO:0000256|HAMAP-
KW   Rule:MF_04070};
KW   RNA-binding {ECO:0000256|ARBA:ARBA00022884, ECO:0000256|HAMAP-
KW   Rule:MF_04070};
KW   Viral nucleoprotein {ECO:0000256|ARBA:ARBA00023086, ECO:0000256|HAMAP-
KW   Rule:MF_04070};
KW   Viral penetration into host nucleus {ECO:0000256|ARBA:ARBA00022524,
KW   ECO:0000256|HAMAP-Rule:MF_04070};
KW   Virion {ECO:0000256|HAMAP-Rule:MF_04070, ECO:0000256|RuleBase:RU361251};
KW   Virus entry into host cell {ECO:0000256|HAMAP-Rule:MF_04070}.
FT   REGION          1..21
FT                   /note="Disordered"
FT                   /evidence="ECO:0000256|SAM:MobiDB-lite"
FT   MOTIF           1..18
FT                   /note="Unconventional nuclear localization signal"
FT                   /evidence="ECO:0000256|HAMAP-Rule:MF_04070"
FT   MOTIF           198..216
FT                   /note="Bipartite nuclear localization signal"
FT                   /evidence="ECO:0000256|HAMAP-Rule:MF_04070"
SQ   SEQUENCE   498 AA;  56377 MW;  91390B2963EAB5A3 CRC64;
     MATKGTKRSY EQMETDGERQ NATEIRASVG KMIDGIGRFY IQMCTELKLS DYEGRLIQNS
     LTIERMVLSA FDERRNKYLE EHPSAGKDPK KTGGPIYRRV DGKWRRELIL YDKEEIRRIW
     RQANNGDDAT AGLTHMMIWH SNLNDATYQR TRALVRTGMD PRMCSLMQGS TLPRRSGAAG
     AAVKGVGTMV MELIRMIKRG INDRNFWRGE NGRRTRIAYE RMCNILKGKF QTAAQRTMVD
     QVRESRNPGN AEFEDLIFLA RSALILRGSV AHKSCLPACV YGSAVASGYD FEREGYSLVG
     IDPFRLLQNS QVYSLIRPNE NPAHKSQLVW MACHSAAFED LRVSSFIRGT KVVPRGKLST
     RGVQIASNEN METMESSTLE LRSRYWAIRT RSGGNTNQQR ASSGQISIQP TFSVQRNLPF
     DRPTIMAAFT GNTEGRTSDM RTEIIRLMES ARPEDVSFQG RGVFELSDEK ATSPIVPSFD
     MSNEGSYFFG DNAEEYDN
//