Reviewed, UniProtKB/Swiss-Prot O35987 (NSF1C_RAT)

Names and origin

Protein names

Recommended name:
    NSFL1 cofactor p47
Alternative name(s):
    p97 cofactor p47
    XY body-associated protein XY40

Gene names

Name:

Nsfl1c

Organism

Rattus norvegicus (Rat)

Taxonomic identifier

10116 [NCBI]

Taxonomic lineage

Eukaryota › Metazoa › Chordata › Craniata › Vertebrata › Euteleostomi › Mammalia › Eutheria › Euarchontoglires › Glires › Rodentia › Sciurognathi › Muroidea › Muridae › Murinae › Rattus

Protein attributes

Sequence length	370 AA.
Sequence status	Complete.
Sequence processing	The displayed sequence is not processed.
Protein existence	Evidence at protein level.

General annotation (Comments)

Function	Reduces the ATPase activity of VCP. Necessary for the fragmentation of Golgi stacks during mitosis and for VCP-mediated reassembly of Golgi stacks after mitosis. May play a role in VCP-mediated formation of transitional endoplasmic reticulum (tER). Ref.2 Ref.7 Ref.9 Ref.11
Subunit structure	Part of a ternary complex containing STX5A, NSFL1C and VCP. NSFL1C forms a homotrimer that binds to one end of a VCP homohexamer. The complex binds to membranes enriched in phosphatidylethanolamine-containing lipids and promotes Golgi membrane fusion. Interaction with VCIP135 leads to dissociation of the complex via ATP hydrolysis by VCP. Binds ubiquitin and mono-ubiquitinated proteins via its N-terminal UBA-like domain when bound to VCP.
Subcellular location	Nucleus. Golgi apparatus › Golgi stack. Note: Predominantly nuclear in interphase cells. Bound to the axial elements of sex chromosomes in pachytene spermatocytes. A small proportion of the protein is cytoplasmic, associated with Golgi stacks. Ref.2 Ref.5 Ref.13
Tissue specificity	Highly expressed in heart, brain, spleen, lung, liver, muscle, kidney and testis. Ref.2 Ref.5 Ref.1
Developmental stage	Highly expressed in pachytene spermatocytes during spermatogenesis.
Post-translational modification	Phosphorylated during mitosis. Phosphorylation inhibits interaction with Golgi membranes and is required for the fragmentation of the Golgi stacks during mitosis. Ref.13 Ref.14 Ref.15
Sequence similarities	Belongs to the NSFL1C family. Contains 1 SEP domain. Contains 1 UBX domain.

Ontologies

Keywords
Cellular component	Golgi apparatus Nucleus
Ligand	Lipid-binding
PTM	Phosphoprotein
Technical term	3D-structure Direct protein sequencing
Gene Ontology (GO)
Biological process	Golgi organization Ref.10 Inferred from physical interaction. Source: RGD membrane fusion Ref.10 Inferred from direct assay. Source: RGD
Cellular component	Golgi stack Inferred from electronic annotation. Source: UniProtKB-SubCell nucleus Inferred from electronic annotation. Source: UniProtKB-SubCell
Molecular function	lipid binding Inferred from electronic annotation. Source: UniProtKB-KW protein binding Ref.2 Inferred from physical interaction. Source: IntAct
Complete GO annotation...

Binary interactions

With	Entry	#Exp.	IntAct	Notes
Vcp	P46462	2	EBI-1993760,EBI-399011

Sequence annotation (Features)

Feature key

Position(s)

Length

Description

Graphical view

Feature identifier

Molecule processing

Chain

1 – 370

370

NSFL1 cofactor p47

PRO_0000210990

Regions

Domain

179 – 244

66

SEP

Domain

291 – 368

78

UBX

Motif

109 – 115

7

Nuclear localization signal

Motif

172 – 175

4

Nuclear localization signal

Amino acid modifications

Modified residue

74

1

Phosphoserine By similarity

Modified residue

114

1

Phosphoserine Ref.14 Ref.15

Modified residue

140

1

Phosphoserine; by CDC2

Modified residue

165

1

Phosphoserine By similarity

Modified residue

167

1

Phosphotyrosine By similarity

Modified residue

176

1

Phosphoserine Ref.15

Modified residue

272

1

Phosphoserine By similarity

Experimental info

Mutagenesis

41

1

F → A: Reduces ubiquitin binding and Golgi reassembly. Ref.11

Mutagenesis

57

1

T → A: No effect on phosphorylation. Ref.13

Mutagenesis

112

1

K → T: Strongly reduces nuclear location. Ref.13

Mutagenesis

114

1

S → A: No effect on phosphorylation. Ref.13

Mutagenesis

140

1

S → A: Abolishes phosphorylation by CDC2. Ref.13

Mutagenesis

140

1

S → D: Strongly reduces binding to Golgi membranes. Ref.13

Mutagenesis

173

1

R → T: Strongly reduces nuclear location. Ref.13

Mutagenesis

272

1

S → A: No effect on phosphorylation. Ref.13

Mutagenesis

301

1

R → A: Reduced interaction with VCP. Ref.17

Mutagenesis

342 – 345

4

TFPN → AG: Strongly reduced interaction with VCP. Ref.17

Mutagenesis

343

1

F → S: Reduced interaction with VCP. Ref.17

Mutagenesis

345

1

N → A: Reduced interaction with VCP. Ref.17

Secondary structure

1

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370

Helix Strand Turn

Details...

Helix

4 – 15

12

Helix

20 – 29

10

Helix

35 – 43

9

Beta strand

181 – 187

7

Beta strand

190 – 192

3

Beta strand

198 – 200

3

Helix

206 – 215

10

Helix

221 – 224

4

Beta strand

227 – 229

3

Beta strand

232 – 237

6

Turn

239 – 241

3

Helix

275 – 286

12

Beta strand

298 – 302

5

Turn

303 – 306

4

Beta strand

307 – 311

5

Helix

318 – 328

11

Helix

331 – 333

3

Beta strand

337 – 341

5

Turn

342 – 345

4

Helix

355 – 358

4

Beta strand

364 – 369

6

Sequences

	Sequence		Length	Mass (Da)
	O35987-1 [UniParc]. Last modified January 1, 1998. Version 1. Checksum: 60C81402E5C58134	FASTA	370	40,680
	10 20 30 40 50 60 MAEERQDALR EFVAVTGAEE DRARFFLESA GWDLQIALAS FYEDGGDEDI VTISQATPSS 70 80 90 100 110 120 VSRGTAPSDN RVTSFRDLIH DQDEEEEEEE GQRFYAGGSE RSGQQIVGPP RKKSPNELVD 130 140 150 160 170 180 DLFKGAKEHG AVAVERVTKS PGETSKPRPF AGGGYRLGAA PEEESAYVAG ERRRHSGQDV 190 200 210 220 230 240 HVVLKLWKTG FSLDNGDLRS YQDPSNAQFL ESIRRGEVPA ELRRLAHGGQ VNLDMEDHRD 250 260 270 280 290 300 EDFVKPKGAF KAFTGEGQKL GSTAPQVLNT SSPAQQAENE AKASSSILIN EAEPTTNIQI 310 320 330 340 350 360 RLADGGRLVQ KFNHSHRISD IRLFIVDARP AMAATSFVLM TTFPNKELAD ENQTLKEANL 370 LNAVIVQRLT

References

	« Hide 'large scale' referencesShow 'large scale' references »
[1]	"Molecular characterization and expression pattern of XY body-associated protein XY40 of the rat." Alsheimer M., Imamichi Y., Heid H., Benavente R. Chromosoma 106:308-314(1997) [PubMed: 9297509] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA], PROTEIN SEQUENCE OF 200-209; 283-292 AND 323-328, TISSUE SPECIFICITY. Strain: Wistar. Tissue: Spermatocyte.
[2]	"p47 is a cofactor for p97-mediated membrane fusion." Kondo H., Rabouille C., Newman R., Levine T.P., Pappin D., Freemont P., Warren G. Nature 388:75-78(1997) [PubMed: 9214505] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [MRNA], PROTEIN SEQUENCE OF 11-22; 94-101; 157-172; 189-214; 260-282; 323-346 AND 357-368, MASS SPECTROMETRY, FUNCTION, INTERACTION WITH VCP, SUBCELLULAR LOCATION, TISSUE SPECIFICITY. Tissue: Liver.
[3]	"The status, quality, and expansion of the NIH full-length cDNA project: the Mammalian Gene Collection (MGC)." The MGC Project Team Genome Res. 14:2121-2127(2004) [PubMed: 15489334] [Abstract] Cited for: NUCLEOTIDE SEQUENCE [LARGE SCALE MRNA]. Tissue: Lung.
[4]	Lubec G., Chen W.-Q. Submitted (APR-2007) to UniProtKB Cited for: PROTEIN SEQUENCE OF 200-214 AND 283-301, MASS SPECTROMETRY. Strain: Sprague-Dawley. Tissue: Hippocampus.
[5]	"Meiosis-specific protein selectively associated with sex chromosomes of rat pachytene spermatocytes." Smith A., Benavente R. Proc. Natl. Acad. Sci. U.S.A. 89:6938-6942(1992) [PubMed: 1495983] [Abstract] Cited for: TISSUE SPECIFICITY, SUBCELLULAR LOCATION.
[6]	"Syntaxin 5 is a common component of the NSF- and p97-mediated reassembly pathways of Golgi cisternae from mitotic Golgi fragments in vitro." Rabouille C., Kondo H., Newman R., Hui N., Freemont P., Warren G. Cell 92:603-610(1998) [PubMed: 9506515] [Abstract] Cited for: INTERACTION WITH STX5A.
[7]	"The p47 co-factor regulates the ATPase activity of the membrane fusion protein, p97." Meyer H.H., Kondo H., Warren G. FEBS Lett. 437:255-257(1998) [PubMed: 9824302] [Abstract] Cited for: FUNCTION.
[8]	"A complex of mammalian ufd1 and npl4 links the AAA-ATPase, p97, to ubiquitin and nuclear transport pathways." Meyer H.H., Shorter J.G., Seemann J., Pappin D., Warren G. EMBO J. 19:2181-2192(2000) [PubMed: 10811609] [Abstract] Cited for: INTERACTION WITH VCP.
[9]	"Role of p97 and syntaxin 5 in the assembly of transitional endoplasmic reticulum." Roy L., Bergeron J.J.M., Lavoie C., Hendriks R., Gushue J., Fazel A., Pelletier A., Morre D.J., Subramaniam V.N., Hong W., Paiement J. Mol. Biol. Cell 11:2529-2542(2000) [PubMed: 10930451] [Abstract] Cited for: FUNCTION.
[10]	"Phospholipid species act as modulators in p97/p47-mediated fusion of Golgi membranes." Pecheur E.-I., Martin I., Maier O., Bakowsky U., Ruysschaert J.-M., Hoekstra D. Biochemistry 41:9813-9823(2002) [PubMed: 12146947] [Abstract] Cited for: INTERACTION WITH MEMBRANES.
[11]	"Direct binding of ubiquitin conjugates by the mammalian p97 adaptor complexes, p47 and Ufd1-Npl4." Meyer H.H., Wang Y., Warren G. EMBO J. 21:5645-5652(2002) [PubMed: 12411482] [Abstract] Cited for: FUNCTION, MUTAGENESIS OF PHE-41.
[12]	"VCIP135, a novel essential factor for p97/p47-mediated membrane fusion, is required for Golgi and ER assembly in vivo." Uchiyama K., Jokitalo E., Kano F., Murata M., Zhang X., Canas B., Newman R., Rabouille C., Pappin D., Freemont P., Kondo H. J. Cell Biol. 159:855-866(2002) [PubMed: 12473691] [Abstract] Cited for: INTERACTION WITH VCIP135.
[13]	"The localization and phosphorylation of p47 are important for Golgi disassembly-assembly during the cell cycle." Uchiyama K., Jokitalo E., Lindman M., Jackman M., Kano F., Murata M., Zhang X., Kondo H. J. Cell Biol. 161:1067-1079(2003) [PubMed: 12810701] [Abstract] Cited for: PHOSPHORYLATION, MUTAGENESIS OF THR-57; LYS-112; SER-114; SER-140; ARG-173 AND SER-272, SUBCELLULAR LOCATION.
[14]	"Phosphoproteomic analysis of rat liver by high capacity IMAC and LC-MS/MS." Moser K., White F.M. J. Proteome Res. 5:98-104(2006) [PubMed: 16396499] [Abstract] Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-114, MASS SPECTROMETRY. Tissue: Liver.
[15]	"Quantitative phosphoproteomics of vasopressin-sensitive renal cells: regulation of aquaporin-2 phosphorylation at two sites." Hoffert J.D., Pisitkun T., Wang G., Shen R.-F., Knepper M.A. Proc. Natl. Acad. Sci. U.S.A. 103:7159-7164(2006) [PubMed: 16641100] [Abstract] Cited for: PHOSPHORYLATION [LARGE SCALE ANALYSIS] AT SER-114 AND SER-176, MASS SPECTROMETRY. Tissue: Kidney.
[16]	"Solution structure and interaction surface of the C-terminal domain from p47: a major p97-cofactor involved in SNARE disassembly." Yuan X., Shaw A., Zhang X., Kondo H., Lally J., Freemont P.S., Matthews S. J. Mol. Biol. 311:255-263(2001) [PubMed: 11478859] [Abstract] Cited for: STRUCTURE BY NMR OF 282-370, INTERACTION WITH VCP.
[17]	"Structural basis of the interaction between the AAA ATPase p97/VCP and its adaptor protein p47." Dreveny I., Kondo H., Uchiyama K., Shaw A., Zhang X., Freemont P.S. EMBO J. 23:1030-1039(2004) [PubMed: 14988733] [Abstract] Cited for: X-RAY CRYSTALLOGRAPHY (2.9 ANGSTROMS) OF 244-370, INTERACTION WITH VCP, MUTAGENESIS OF ARG-301; 342-THR--ASN-345; PHE-343 AND ASN-345.
[18]	"Structure, dynamics and interactions of p47, a major adaptor of the AAA ATPase, p97." Yuan X., Simpson P., McKeown C., Kondo H., Uchiyama K., Wallis R., Dreveny I., Keetch C., Zhang X., Robinson C., Freemont P., Matthews S. EMBO J. 23:1463-1473(2004) [PubMed: 15029246] [Abstract] Cited for: STRUCTURE BY NMR OF 1-46 IN COMPLEX WITH UBQUITIN, STRUCTURE BY NMR OF 171-246, INTERACTION WITH VCP.

Cross-references

Sequence databases

EMBLGenBankDDBJ

Y10769 mRNA. Translation: CAA71742.1.
AB002086 mRNA. Translation: BAA21659.1.
BC072464 mRNA. Translation: AAH72464.1.

IPI

IPI00214262.

RefSeq

NP_114187.1.

UniGene

Rn.2771

3D structure databases

PDBPDBe

Entry	Method	Resolution (Å)	Chain	Positions	PDBsum
1I42	NMR	-	A	282-370	[»]
1JRU	NMR	-	A	282-370	[»]
1S3S	X-ray	2.90	G/H/I	244-370	[»]
1V92	NMR	-	A	1-46	[»]
1VAZ	NMR	-	A	171-246	[»]

SMR

O35987. Positions 171-270.

ModBase

Search...

Protein-protein interaction databases

IntAct

O35987. 15 interactions.

PTM databases

PhosphoSite

O35987.

Genome annotation databases

Ensembl

ENSRNOG00000008604. Rattus norvegicus. [Contig view]

GeneID

83809.

KEGG

rno:83809.

Organism-specific databases

RGD

619952. Nsfl1c.

Phylogenomic databases

HOVERGEN

O35987.

Gene expression databases

ArrayExpress

O35987.

GermOnline

ENSRNOG00000008604. Rattus norvegicus.

Family and domain databases

InterPro

IPR012989. SEP.
IPR001012. UBX.
[Graphical view]

Pfam

PF08059. SEP. 1 hit.
PF00789. UBX. 1 hit.
[Graphical view]

SMART

SM00553. SEP. 1 hit.
SM00166. UBX. 1 hit.
[Graphical view]

PROSITE

PS51399. SEP. 1 hit.
PS50033. UBX. 1 hit.
[Graphical view]

ProtoNet

Search...

Other Resources

NextBio

616417.

Entry information

Entry name

NSF1C_RAT

Accession

Primary (citable) accession number: O35987

Entry history

Integrated into UniProtKB/Swiss-Prot:	January 16, 2004
Last sequence update:	January 1, 1998
Last modified:	June 16, 2009
This is version 81 of the entry and version 1 of the sequence. [Complete history]

Entry status

Reviewed (UniProtKB/Swiss-Prot)

Annotation project

HPI (Human Proteome Initiative)