Nae1 (NEDD8 activating enzyme E1 subunit 1) - Rat Genome Database

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Gene: Nae1 (NEDD8 activating enzyme E1 subunit 1) Rattus norvegicus
Analyze
Symbol: Nae1
Name: NEDD8 activating enzyme E1 subunit 1
RGD ID: 619945
Description: Predicted to enable protein heterodimerization activity; ubiquitin activating enzyme activity; and ubiquitin protein ligase binding activity. Predicted to contribute to NEDD8 activating enzyme activity. Predicted to be involved in several processes, including mitotic DNA replication checkpoint signaling; protein modification by small protein conjugation; and ubiquitin-dependent protein catabolic process. Located in early endosome. Biomarker of middle cerebral artery infarction. Orthologous to human NAE1 (NEDD8 activating enzyme E1 subunit 1); PARTICIPATES IN neddylation pathway; proteasome degradation pathway involving cullin-dependent ubiquitin ligases; Alzheimer's disease pathway; INTERACTS WITH 1,3,5-trinitro-1,3,5-triazinane; 17alpha-ethynylestradiol; 2,4-dinitrotoluene.
Type: protein-coding
RefSeq Status: VALIDATED
Previously known as: amyloid beta precursor protein binding protein 1; amyloid beta precursor protein-binding protein 1, 59 kDa; amyloid protein-binding protein 1; APP-binding protein 1; APP-BP1; Appbp1; LOC291815; NEDD8-activating enzyme E1 regulatory subunit; RGD1563180; similar to amyloid beta precursor protein binding protein 1
RGD Orthologs
Human
Mouse
Chinchilla
Bonobo
Dog
Squirrel
Pig
Green Monkey
Naked Mole-Rat
Alliance Genes
More Info more info ...
Latest Assembly: mRatBN7.2 - mRatBN7.2 Assembly
Position:
Rat AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
GRCr819452,462 - 478,591 (+)NCBIGRCr8
mRatBN7.219446,015 - 472,145 (+)NCBImRatBN7.2mRatBN7.2
mRatBN7.2 Ensembl19446,000 - 472,371 (+)EnsemblmRatBN7.2 Ensembl
UTH_Rnor_SHR_Utx19441,827 - 468,061 (+)NCBIRnor_SHRUTH_Rnor_SHR_Utx
UTH_Rnor_SHRSP_BbbUtx_1.0191,208,624 - 1,234,856 (+)NCBIRnor_SHRSPUTH_Rnor_SHRSP_BbbUtx_1.0
UTH_Rnor_WKY_Bbb_1.019483,838 - 510,076 (+)NCBIRnor_WKYUTH_Rnor_WKY_Bbb_1.0
Rnor_6.019653,510 - 680,378 (+)NCBIRnor6.0Rnor_6.0rn6Rnor6.0
Rnor_6.0 Ensembl19653,510 - 680,378 (+)EnsemblRnor6.0rn6Rnor6.0
Rnor_5.019646,826 - 673,676 (+)NCBIRnor5.0Rnor_5.0rn5Rnor5.0
RGSC_v3.419382,556 - 408,667 (+)NCBIRGSC3.4RGSC_v3.4rn4RGSC3.4
RGSC_v3.119382,555 - 408,667 (+)NCBI
Celera19446,357 - 467,621 (+)NCBICelera
Cytogenetic Map19p14NCBI
JBrowse: View Region in Genome Browser (JBrowse)
Model


Gene-Chemical Interaction Annotations     Click to see Annotation Detail View
Gene Ontology Annotations     Click to see Annotation Detail View

Cellular Component

Molecular Function

Molecular Pathway Annotations     Click to see Annotation Detail View
References

References - curated
# Reference Title Reference Citation
1. APP-BP1 mediates APP-induced apoptosis and DNA synthesis and is increased in Alzheimer's disease brain. Chen Y, etal., J Cell Biol. 2003 Oct 13;163(1):27-33.
2. ASPP2 inhibits APP-BP1-mediated NEDD8 conjugation to cullin-1 and decreases APP-BP1-induced cell proliferation and neuronal apoptosis. Chen Y, etal., J Neurochem 2003 May;85(3):801-9.
3. APP-BP1, a novel protein that binds to the carboxyl-terminal region of the amyloid precursor protein. Chow N, etal., J Biol Chem 1996 May 10;271(19):11339-46.
4. Phylogenetic-based propagation of functional annotations within the Gene Ontology consortium. Gaudet P, etal., Brief Bioinform. 2011 Sep;12(5):449-62. doi: 10.1093/bib/bbr042. Epub 2011 Aug 27.
5. Rat ISS GO annotations from GOA human gene data--August 2006 GOA data from the GO Consortium
6. Focal transient ischemia increases APP-BP1 expression in neural progenitor cells. Joo Y, etal., Neuroreport. 2011 Mar 30;22(5):200-5. doi: 10.1097/WNR.0b013e32834449cf.
7. KEGG: Kyoto Encyclopedia of Genes and Genomes KEGG
8. Rab5 mediates an amyloid precursor protein signaling pathway that leads to apoptosis. Laifenfeld D, etal., J Neurosci. 2007 Jul 4;27(27):7141-53. doi: 10.1523/JNEUROSCI.4599-06.2007.
9. Rat ISS GO annotations from MGI mouse gene data--August 2006 MGD data from the GO Consortium
10. Electronic Transfer of LocusLink and RefSeq Data NCBI rat LocusLink and RefSeq merged data July 26, 2002
11. A new NEDD8-ligating system for cullin-4A. Osaka F, etal., Genes Dev. 1998 Aug 1;12(15):2263-8. doi: 10.1101/gad.12.15.2263.
12. KEGG Annotation Import Pipeline Pipeline to import KEGG annotations from KEGG into RGD
13. GOA pipeline RGD automated data pipeline
14. ClinVar Automated Import and Annotation Pipeline RGD automated import pipeline for ClinVar variants, variant-to-disease annotations and gene-to-disease annotations
15. Data Import for Chemical-Gene Interactions RGD automated import pipeline for gene-chemical interactions
16. Comprehensive gene review and curation RGD comprehensive gene curation
17. Insights into the ubiquitin transfer cascade from the structure of the activating enzyme for NEDD8. Walden H, etal., Nature 2003 Mar 20;422(6929):330-4.
Additional References at PubMed
PMID:10207026   PMID:10722740   PMID:12477932   PMID:12740388   PMID:18627766   PMID:21151996  


Genomics

Comparative Map Data
Nae1
(Rattus norvegicus - Norway rat)
Rat AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
GRCr819452,462 - 478,591 (+)NCBIGRCr8
mRatBN7.219446,015 - 472,145 (+)NCBImRatBN7.2mRatBN7.2
mRatBN7.2 Ensembl19446,000 - 472,371 (+)EnsemblmRatBN7.2 Ensembl
UTH_Rnor_SHR_Utx19441,827 - 468,061 (+)NCBIRnor_SHRUTH_Rnor_SHR_Utx
UTH_Rnor_SHRSP_BbbUtx_1.0191,208,624 - 1,234,856 (+)NCBIRnor_SHRSPUTH_Rnor_SHRSP_BbbUtx_1.0
UTH_Rnor_WKY_Bbb_1.019483,838 - 510,076 (+)NCBIRnor_WKYUTH_Rnor_WKY_Bbb_1.0
Rnor_6.019653,510 - 680,378 (+)NCBIRnor6.0Rnor_6.0rn6Rnor6.0
Rnor_6.0 Ensembl19653,510 - 680,378 (+)EnsemblRnor6.0rn6Rnor6.0
Rnor_5.019646,826 - 673,676 (+)NCBIRnor5.0Rnor_5.0rn5Rnor5.0
RGSC_v3.419382,556 - 408,667 (+)NCBIRGSC3.4RGSC_v3.4rn4RGSC3.4
RGSC_v3.119382,555 - 408,667 (+)NCBI
Celera19446,357 - 467,621 (+)NCBICelera
Cytogenetic Map19p14NCBI
NAE1
(Homo sapiens - human)
Human AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
GRCh381666,802,878 - 66,830,976 (-)NCBIGRCh38GRCh38hg38GRCh38
GRCh38.p14 Ensembl1666,802,875 - 66,873,256 (-)EnsemblGRCh38hg38GRCh38
GRCh371666,836,781 - 66,864,879 (-)NCBIGRCh37GRCh37hg19GRCh37
Build 361665,394,282 - 65,422,380 (-)NCBINCBI36Build 36hg18NCBI36
Build 341665,394,282 - 65,422,370NCBI
Celera1651,345,091 - 51,373,189 (-)NCBICelera
Cytogenetic Map16q22.1NCBI
HuRef1652,710,930 - 52,739,089 (-)NCBIHuRef
CHM1_11668,243,864 - 68,271,962 (-)NCBICHM1_1
T2T-CHM13v2.01672,597,172 - 72,625,270 (-)NCBIT2T-CHM13v2.0
Nae1
(Mus musculus - house mouse)
Mouse AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
GRCm398105,237,660 - 105,261,269 (-)NCBIGRCm39GRCm39mm39
GRCm39 Ensembl8105,237,660 - 105,261,269 (-)EnsemblGRCm39 Ensembl
GRCm388104,511,028 - 104,534,656 (-)NCBIGRCm38GRCm38mm10GRCm38
GRCm38.p6 Ensembl8104,511,028 - 104,534,637 (-)EnsemblGRCm38mm10GRCm38
MGSCv378107,034,928 - 107,058,537 (-)NCBIGRCm37MGSCv37mm9NCBIm37
MGSCv368107,400,153 - 107,423,744 (-)NCBIMGSCv36mm8
Celera8108,742,436 - 108,767,075 (-)NCBICelera
Cytogenetic Map8D3NCBI
cM Map853.04NCBI
Nae1
(Chinchilla lanigera - long-tailed chinchilla)
Chinchilla AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
ChiLan1.0 EnsemblNW_00495543323,333,742 - 23,358,579 (-)EnsemblChiLan1.0
ChiLan1.0NW_00495543323,333,797 - 23,358,255 (-)NCBIChiLan1.0ChiLan1.0
NAE1
(Pan paniscus - bonobo/pygmy chimpanzee)
Bonobo AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
NHGRI_mPanPan1-v21876,304,088 - 76,332,248 (-)NCBINHGRI_mPanPan1-v2
NHGRI_mPanPan11682,216,589 - 82,244,743 (-)NCBINHGRI_mPanPan1
Mhudiblu_PPA_v01647,119,548 - 47,147,679 (-)NCBIMhudiblu_PPA_v0Mhudiblu_PPA_v0panPan3
PanPan1.11666,221,829 - 66,246,648 (-)NCBIpanpan1.1PanPan1.1panPan2
PanPan1.1 Ensembl1666,221,829 - 66,246,865 (-)Ensemblpanpan1.1panPan2
NAE1
(Canis lupus familiaris - dog)
Dog AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
CanFam3.1582,462,690 - 82,486,097 (+)NCBICanFam3.1CanFam3.1canFam3CanFam3.1
CanFam3.1 Ensembl582,462,744 - 82,486,100 (+)EnsemblCanFam3.1canFam3CanFam3.1
Dog10K_Boxer_Tasha582,451,391 - 82,474,791 (+)NCBIDog10K_Boxer_Tasha
ROS_Cfam_1.0582,899,329 - 82,922,733 (+)NCBIROS_Cfam_1.0
ROS_Cfam_1.0 Ensembl582,899,354 - 82,922,729 (+)EnsemblROS_Cfam_1.0 Ensembl
UMICH_Zoey_3.1582,722,479 - 82,745,880 (+)NCBIUMICH_Zoey_3.1
UNSW_CanFamBas_1.0582,409,395 - 82,433,102 (+)NCBIUNSW_CanFamBas_1.0
UU_Cfam_GSD_1.0583,048,382 - 83,071,880 (+)NCBIUU_Cfam_GSD_1.0
Nae1
(Ictidomys tridecemlineatus - thirteen-lined ground squirrel)
Squirrel AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
HiC_Itri_2NW_02440934942,483,986 - 42,505,936 (+)NCBIHiC_Itri_2
SpeTri2.0 EnsemblNW_00493647517,083,077 - 17,105,133 (-)EnsemblSpeTri2.0SpeTri2.0 Ensembl
SpeTri2.0NW_00493647517,083,175 - 17,105,098 (-)NCBISpeTri2.0SpeTri2.0SpeTri2.0
NAE1
(Sus scrofa - pig)
Pig AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
Sscrofa11.1 Ensembl627,450,632 - 27,540,118 (-)EnsemblSscrofa11.1susScr11Sscrofa11.1
Sscrofa11.1627,511,541 - 27,540,142 (-)NCBISscrofa11.1Sscrofa11.1susScr11Sscrofa11.1
NAE1
(Chlorocebus sabaeus - green monkey)
Green Monkey AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
ChlSab1.1552,502,729 - 52,531,447 (-)NCBIChlSab1.1ChlSab1.1chlSab2
ChlSab1.1 Ensembl552,502,546 - 52,525,262 (-)EnsemblChlSab1.1ChlSab1.1 EnsemblchlSab2
Vero_WHO_p1.0NW_02366604723,380,340 - 23,409,070 (+)NCBIVero_WHO_p1.0Vero_WHO_p1.0
Nae1
(Heterocephalus glaber - naked mole-rat)
Naked Mole-Rat AssemblyChrPosition (strand)SourceGenome Browsers
JBrowseNCBIUCSCEnsembl
HetGla_female_1.0 EnsemblNW_00462474619,658,632 - 19,687,884 (+)EnsemblHetGla_female_1.0HetGla_female_1.0 EnsemblhetGla2
HetGla 1.0NW_00462474619,648,365 - 19,687,884 (+)NCBIHetGla_female_1.0HetGla 1.0hetGla2

Variants

.
Variants in Nae1
312 total Variants
miRNA Target Status

Predicted Target Of
Summary Value
Count of predictions:150
Count of miRNA genes:119
Interacting mature miRNAs:128
Transcripts:ENSRNOT00000048221
Prediction methods:Microtar, Miranda, Rnahybrid, Targetscan
Result types:miRGate_prediction

The detailed report is available here: Full Report CSV TAB Printer

miRNA Target Status data imported from miRGate (http://mirgate.bioinfo.cnio.es/).
For more information about miRGate, see PMID:25858286 or access the full paper here.


QTLs in Region (mRatBN7.2)
The following QTLs overlap with this region.    Full Report CSV TAB Printer Gviewer
RGD IDSymbolNameLODP ValueTraitSub TraitChrStartStopSpecies
7411590Foco7Food consumption QTL 76.80.001eating behavior trait (VT:0001431)feed conversion ratio (CMO:0001312)19124688055Rat
10054132Srcrt9Stress Responsive Cort QTL 92.870.0017blood corticosterone amount (VT:0005345)plasma corticosterone level (CMO:0001173)19127355345Rat
631678Cm9Cardiac mass QTL 94.270.0001aorta mass (VT:0002845)aorta weight (CMO:0000076)19128982497Rat
631681Cm12Cardiac mass QTL 123.330.00053heart left ventricle mass (VT:0007031)heart left ventricle weight to body weight ratio (CMO:0000530)19128982497Rat
1549847Bss8Bone structure and strength QTL 84lumbar vertebra strength trait (VT:0010574)vertebra ultimate force (CMO:0001678)19131963836Rat
9590298Uminl5Urine mineral level QTL 53.590.001urine mineral amount (VT:0015086)urine electrolyte level (CMO:0000593)19136824771Rat
8552935Pigfal10Plasma insulin-like growth factor 1 level QTL 105.7blood insulin-like growth factor amount (VT:0010479)plasma insulin-like growth factor 1 level (CMO:0001299)19136824771Rat
9590250Scort11Serum corticosterone level QTL 1123.450.001blood corticosterone amount (VT:0005345)plasma corticosterone level (CMO:0001173)19136824771Rat
9590090Insglur8Insulin/glucose ratio QTL 810.810.001blood insulin amount (VT:0001560)calculated plasma insulin level (CMO:0002170)19136824771Rat


Expression


RNA-SEQ Expression
High: > 1000 TPM value   Medium: Between 11 and 1000 TPM
Low: Between 0.5 and 10 TPM   Below Cutoff: < 0.5 TPM

alimentary part of gastrointestinal system circulatory system endocrine system exocrine system hemolymphoid system hepatobiliary system integumental system musculoskeletal system nervous system renal system reproductive system respiratory system appendage
High
Medium 3 42 49 33 19 33 8 9 74 35 39 11 8
Low 1 8 8 8 2 2
Below cutoff

Sequence


RefSeq Acc Id: ENSRNOT00000048221   ⟹   ENSRNOP00000045593
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2 Ensembl19446,000 - 472,307 (+)Ensembl
Rnor_6.0 Ensembl19653,510 - 680,376 (+)Ensembl
RefSeq Acc Id: ENSRNOT00000076948
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2 Ensembl19446,000 - 472,307 (+)Ensembl
Rnor_6.0 Ensembl19653,510 - 680,378 (+)Ensembl
RefSeq Acc Id: ENSRNOT00000112150   ⟹   ENSRNOP00000077419
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
mRatBN7.2 Ensembl19450,135 - 472,371 (+)Ensembl
RefSeq Acc Id: NM_032072   ⟹   NP_114461
RefSeq Status: VALIDATED
Type: CODING
Position:
Rat AssemblyChrPosition (strand)Source
GRCr819452,462 - 478,591 (+)NCBI
mRatBN7.219446,015 - 472,145 (+)NCBI
Rnor_6.019653,510 - 680,378 (+)NCBI
Rnor_5.019646,826 - 673,676 (+)NCBI
RGSC_v3.419382,556 - 408,667 (+)RGD
Celera19446,357 - 467,621 (+)NCBI
Sequence:
Protein Sequences
Protein RefSeqs NP_114461 (Get FASTA)   NCBI Sequence Viewer  
GenBank Protein AAD09247 (Get FASTA)   NCBI Sequence Viewer  
  AAI27482 (Get FASTA)   NCBI Sequence Viewer  
Ensembl Protein ENSRNOP00000045593
  ENSRNOP00000045593.4
  ENSRNOP00000077419.1
  ENSRNOP00000083806.1
GenBank Protein Q9Z1A5 (Get FASTA)   NCBI Sequence Viewer  
RefSeq Acc Id: NP_114461   ⟸   NM_032072
- UniProtKB: Q9Z1A5 (UniProtKB/Swiss-Prot),   A1L122 (UniProtKB/TrEMBL),   F1M7W7 (UniProtKB/TrEMBL)
- Sequence:
RefSeq Acc Id: ENSRNOP00000045593   ⟸   ENSRNOT00000048221
RefSeq Acc Id: ENSRNOP00000077419   ⟸   ENSRNOT00000112150
Protein Structures
Name Modeler Protein Id AA Range Protein Structure
AF-Q9Z1A5-F1-model_v2 AlphaFold Q9Z1A5 1-534 view protein structure

Transcriptome

eQTL   View at Phenogen
WGCNA   View at Phenogen
Tissue/Strain Expression   View at Phenogen

Promoters
RGD ID:13700910
Promoter ID:EPDNEW_R11433
Type:initiation region
Name:Nae1_1
Description:NEDD8 activating enzyme E1 subunit 1
SO ACC ID:SO:0000170
Source:EPDNEW (Eukaryotic Promoter Database, http://epd.vital-it.ch/)
Experiment Methods:Single-end sequencing.
Position:
Rat AssemblyChrPosition (strand)Source
Rnor_6.019653,507 - 653,567EPDNEW

Additional Information

Database Acc Id Source(s)
AGR Gene RGD:619945 AgrOrtholog
BioCyc Gene G2FUF-6647 BioCyc
BioCyc Pathway PWY-7899 [protein NEDDylation] BioCyc
BioCyc Pathway Image PWY-7899 BioCyc
Ensembl Genes ENSRNOG00000033133 Ensembl, ENTREZGENE, UniProtKB/TrEMBL
Ensembl Transcript ENSRNOT00000048221 ENTREZGENE
  ENSRNOT00000048221.4 UniProtKB/TrEMBL
  ENSRNOT00000076948.2 UniProtKB/TrEMBL
  ENSRNOT00000112150.1 UniProtKB/TrEMBL
Gene3D-CATH NAD(P)-binding Rossmann-like Domain UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
IMAGE_CLONE IMAGE:8370668 IMAGE-MGC_LOAD
InterPro APP-BP1 UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
  ThiF/MoeB/HesA UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
  ThiF_NAD_FAD-bd UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
  Ubiquitin-activating_enz UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
KEGG Report rno:84019 UniProtKB/TrEMBL
MGC_CLONE MGC:156584 IMAGE-MGC_LOAD
NCBI Gene 84019 ENTREZGENE
PANTHER NEDD8-ACTIVATING ENZYME E1 REGULATORY SUBUNIT UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
  PTHR10953 UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
Pfam ThiF UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
PhenoGen Nae1 PhenoGen
PIRSF APP-BP1 UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
RatGTEx ENSRNOG00000033133 RatGTEx
Superfamily-SCOP SSF69572 UniProtKB/Swiss-Prot, UniProtKB/TrEMBL
UniProt A0A8I5ZKA7_RAT UniProtKB/TrEMBL
  A0A8I6A0P5_RAT UniProtKB/TrEMBL
  A1L122 ENTREZGENE, UniProtKB/TrEMBL
  F1M7W7 ENTREZGENE
  Q9Z1A5 ENTREZGENE, UniProtKB/Swiss-Prot
UniProt Secondary F1M7W7 UniProtKB/TrEMBL


Nomenclature History
Date Current Symbol Current Name Previous Symbol Previous Name Description Reference Status
2015-07-29 Nae1  NEDD8 activating enzyme E1 subunit 1  RGD1563180  similar to amyloid beta precursor protein binding protein 1  Data merged from RGD:1563180 737654 PROVISIONAL
2008-04-30 RGD1563180  similar to amyloid beta precursor protein binding protein 1   RGD1563180_predicted  similar to amyloid beta precursor protein binding protein 1 (predicted)  'predicted' is removed 2292626 APPROVED
2008-04-25 Nae1  NEDD8 activating enzyme E1 subunit 1  Appbp1  amyloid beta precursor protein binding protein 1  Nomenclature updated to reflect human and mouse nomenclature 1299863 APPROVED
2006-03-07 RGD1563180_predicted  similar to amyloid beta precursor protein binding protein 1 (predicted)  LOC291815  similar to amyloid beta precursor protein binding protein 1  Symbol and Name status set to approved 1299863 APPROVED
2006-02-09 LOC291815  similar to amyloid beta precursor protein binding protein 1      Symbol and Name status set to provisional 70820 PROVISIONAL
2004-12-14 Appbp1  amyloid beta precursor protein binding protein 1    APP-binding protein 1  Name updated 1299863 APPROVED
2002-08-07 Appbp1  APP-binding protein 1      Symbol and Name status set to provisional 70820 PROVISIONAL

RGD Curation Notes
Note Type Note Reference
gene_physical_interaction human homolog binds the carboxyl (C) terminus of the amyloid precursor protein (APP) and is a component of the bipartite activating enzyme for the ubiquitin-like protein NEDD8 634621