Skip Header

You are using a version of browser that may not display all the features of this website. Please consider upgrading your browser.
Entry version 223 (02 Jun 2021)
Sequence version 2 (11 Oct 2005)
Previous versions | rss
Add a publicationFeedback
Protein

Progranulin

Gene

GRN

Organism
Homo sapiens (Human)
Status
Reviewed-Annotation score:

Annotation score:5 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
-Experimental evidence at protein leveli <p>This indicates the type of evidence that supports the existence of the protein. Note that the 'protein existence' evidence does not give information on the accuracy or correctness of the sequence(s) displayed.<p><a href='/help/protein_existence' target='_top'>More...</a></p>

<p>This section provides any useful information about the protein, mostly biological knowledge.<p><a href='/help/function_section' target='_top'>More...</a></p>Functioni

Secreted protein that acts as a key regulator of lysosomal function and as a growth factor involved in inflammation, wound healing and cell proliferation (PubMed:28541286, PubMed:28073925, PubMed:18378771, PubMed:28453791, PubMed:12526812).

Regulates protein trafficking to lysosomes and, also the activity of lysosomal enzymes (PubMed:28453791, PubMed:28541286).

Facilitates also the acidification of lysosomes, causing degradation of mature CTSD by CTSB (PubMed:28073925).

In addition, functions as wound-related growth factor that acts directly on dermal fibroblasts and endothelial cells to promote division, migration and the formation of capillary-like tubule structures (By similarity).

Also promotes epithelial cell proliferation by blocking TNF-mediated neutrophil activation preventing release of oxidants and proteases (PubMed:12526812).

Moreover, modulates inflammation in neurons by preserving neurons survival, axonal outgrowth and neuronal integrity (PubMed:18378771).

By similarity5 Publications

Promotes proliferation of the epithelial cell line A431 in culture.

Inhibits epithelial cell proliferation and induces epithelial cells to secrete IL-8.

1 Publication

Stabilizes CTSD through interaction with CTSD leading to maintain its aspartic-type peptidase activity.

1 Publication

<p>The <a href="http://www.geneontology.org/">Gene Ontology (GO)</a> project provides a set of hierarchical controlled vocabulary split into 3 categories:<p><a href='/help/gene_ontology' target='_top'>More...</a></p>GO - Molecular functioni

GO - Biological processi

<p>UniProtKB Keywords constitute a <a href="http://www.uniprot.org/keywords">controlled vocabulary</a> with a hierarchical structure. Keywords summarise the content of a UniProtKB entry and facilitate the search for proteins of interest.<p><a href='/help/keywords' target='_top'>More...</a></p>Keywordsi

Molecular functionCytokine

Enzyme and pathway databases

Pathway Commons web resource for biological pathway data

More...
PathwayCommonsi
P28799

Reactome - a knowledgebase of biological pathways and processes

More...
Reactomei
R-HSA-6798695, Neutrophil degranulation

SIGNOR Signaling Network Open Resource

More...
SIGNORi
P28799

<p>This section provides information about the protein and gene name(s) and synonym(s) and about the organism that is the source of the protein sequence.<p><a href='/help/names_and_taxonomy_section' target='_top'>More...</a></p>Names & Taxonomyi

<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section provides an exhaustive list of all names of the protein, from commonly used to obsolete, to allow unambiguous identification of a protein.<p><a href='/help/protein_names' target='_top'>More...</a></p>Protein namesi
Recommended name:
Progranulin1 Publication
Short name:
PGRN1 Publication
Alternative name(s):
AcrograninBy similarity
Epithelin precursor1 Publication
Glycoprotein of 88 KdaBy similarity
Short name:
GP88
Short name:
Glycoprotein 88
Granulin precursor1 Publication
PC cell-derived growth factorBy similarity
Short name:
PCDGF1 Publication
Proepithelin2 Publications
Short name:
PEPI1 Publication
Cleaved into the following 8 chains:
Alternative name(s):
Granulin G
Alternative name(s):
Granulin F
Alternative name(s):
Epithelin-2By similarity
Granulin B
Alternative name(s):
Epithelin-1By similarity
Granulin A
Alternative name(s):
Granulin C
Alternative name(s):
Granulin D
Alternative name(s):
Granulin E
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section indicates the name(s) of the gene(s) that code for the protein sequence(s) described in the entry. Four distinct tokens exist: 'Name', 'Synonyms', 'Ordered locus names' and 'ORF names'.<p><a href='/help/gene_name' target='_top'>More...</a></p>Gene namesi
Name:GRNImported
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section provides information on the name(s) of the organism that is the source of the protein sequence.<p><a href='/help/organism-name' target='_top'>More...</a></p>OrganismiHomo sapiens (Human)
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section shows the unique identifier assigned by the NCBI to the source organism of the protein. This is known as the 'taxonomic identifier' or 'taxid'.<p><a href='/help/taxonomic_identifier' target='_top'>More...</a></p>Taxonomic identifieri9606 [NCBI]
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section contains the taxonomic hierarchical classification lineage of the source organism. It lists the nodes as they appear top-down in the taxonomic tree, with the more general grouping listed first.<p><a href='/help/taxonomic_lineage' target='_top'>More...</a></p>Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresPrimatesHaplorrhiniCatarrhiniHominidaeHomo
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section is present for entries that are part of a <a href="http://www.uniprot.org/proteomes">proteome</a>, i.e. of a set of proteins thought to be expressed by organisms whose genomes have been completely sequenced.<p><a href='/help/proteomes_manual' target='_top'>More...</a></p>Proteomesi
  • UP000005640 <p>A UniProt <a href="http://www.uniprot.org/manual/proteomes%5Fmanual">proteome</a> can consist of several components.<br></br>The component name refers to the genomic component encoding a set of proteins.<p><a href='/help/proteome_component' target='_top'>More...</a></p> Componenti: Chromosome 17

Organism-specific databases

Human Gene Nomenclature Database

More...
HGNCi
HGNC:4601, GRN

Online Mendelian Inheritance in Man (OMIM)

More...
MIMi
138945, gene

neXtProt; the human protein knowledge platform

More...
neXtProti
NX_P28799

Eukaryotic Pathogen, Vector and Host Database Resources

More...
VEuPathDBi
HostDB:ENSG00000030582.16

<p>This section provides information on the location and the topology of the mature protein in the cell.<p><a href='/help/subcellular_location_section' target='_top'>More...</a></p>Subcellular locationi

Keywords - Cellular componenti

Lysosome, Secreted

<p>This section provides information on the disease(s) and phenotype(s) associated with a protein.<p><a href='/help/pathology_and_biotech_section' target='_top'>More...</a></p>Pathology & Biotechi

<p>This subsection of the 'Pathology and Biotech' section provides information on the disease(s) associated with genetic variations in a given protein. The information is extracted from the scientific literature and diseases that are also described in the <a href="http://www.ncbi.nlm.nih.gov/sites/entrez?db=omim">OMIM</a> database are represented with a <a href="http://www.uniprot.org/diseases">controlled vocabulary</a> in the following way:<p><a href='/help/involvement_in_disease' target='_top'>More...</a></p>Involvement in diseasei

Ubiquitin-positive frontotemporal dementia (UP-FTD)3 Publications
The disease is caused by variants affecting the gene represented in this entry.
Disease descriptionFrontotemporal dementia (FTD) is the second most common cause of dementia in people under the age of 65 years. It is an autosomal dominant neurodegenerative disease.
Related information in OMIM
Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'Sequence' section describes natural variant(s) of the protein sequence.<p><a href='/help/variant' target='_top'>More...</a></p>Natural variantiVAR_0444519A → D in UP-FTD; no significant difference in the total mRNA between cases and controls; although the mutant protein is expressed it is not secreted and appears to be trapped within an intracellular compartment. 2 PublicationsCorresponds to variant dbSNP:rs63751243EnsemblClinVar.1
Ceroid lipofuscinosis, neuronal, 11 (CLN11)1 Publication
The disease is caused by variants affecting the gene represented in this entry.
Disease descriptionA form of neuronal ceroid lipofuscinosis characterized by rapidly progressive visual loss due to retinal dystrophy, seizures, cerebellar ataxia, and cerebellar atrophy. Cognitive decline may also occur. Neuronal ceroid lipofuscinoses are progressive neurodegenerative, lysosomal storage diseases characterized by intracellular accumulation of autofluorescent liposomal material.
Related information in OMIM

Keywords - Diseasei

Neurodegeneration, Neuronal ceroid lipofuscinosis

Organism-specific databases

DisGeNET

More...
DisGeNETi
2896

GeneReviews a resource of expert-authored, peer-reviewed disease descriptions.

More...
GeneReviewsi
GRN

MalaCards human disease database

More...
MalaCardsi
GRN
MIMi607485, phenotype
614706, phenotype

Open Targets

More...
OpenTargetsi
ENSG00000030582

Orphanet; a database dedicated to information on rare diseases and orphan drugs

More...
Orphaneti
275864, Behavioral variant of frontotemporal dementia
314629, CLN11 disease
100070, Progressive non-fluent aphasia
100069, Semantic dementia

The Pharmacogenetics and Pharmacogenomics Knowledge Base

More...
PharmGKBi
PA28998

Miscellaneous databases

Pharos NIH Druggable Genome Knowledgebase

More...
Pharosi
P28799, Tbio

Genetic variation databases

BioMuta curated single-nucleotide variation and disease association database

More...
BioMutai
GRN

Domain mapping of disease mutations (DMDM)

More...
DMDMi
77416865

<p>This section describes post-translational modifications (PTMs) and/or processing events.<p><a href='/help/ptm_processing_section' target='_top'>More...</a></p>PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'PTM / Processing' section denotes the presence of an N-terminal signal peptide.<p><a href='/help/signal' target='_top'>More...</a></p>Signal peptidei1 – 17Sequence analysisAdd BLAST17
<p>This subsection of the 'PTM / Processing' section describes the extent of a polypeptide chain in the mature protein following processing or proteolytic cleavage.<p><a href='/help/chain' target='_top'>More...</a></p>ChainiPRO_000001269318 – 593ProgranulinAdd BLAST576
<p>This subsection of the 'PTM / Processing' section describes the position and length of an active peptide in the mature protein.<p><a href='/help/peptide' target='_top'>More...</a></p>PeptideiPRO_000001269418 – ?47ParagranulinAdd BLAST30
PeptideiPRO_0000012695?58 – ?113Granulin-1Add BLAST56
PeptideiPRO_0000012696123 – 179Granulin-2Add BLAST57
PeptideiPRO_0000012697206 – 261Granulin-3Add BLAST56
PeptideiPRO_0000012698281 – 336Granulin-4Add BLAST56
PeptideiPRO_0000012699364 – 417Granulin-5Add BLAST54
PeptideiPRO_0000012700442 – ?496Granulin-6Add BLAST55
PeptideiPRO_0000012701?518 – ?573Granulin-7Add BLAST56

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/help/ptm%5Fprocessing%5Fsection">PTM / Processing</a> section specifies the position and type of each covalently attached glycan group (mono-, di-, or polysaccharide).<p><a href='/help/carbohyd' target='_top'>More...</a></p>Glycosylationi118N-linked (GlcNAc...) asparagine1 Publication1
<p>This subsection of the PTM / Processing":/help/ptm_processing_section section describes the positions of cysteine residues participating in disulfide bonds.<p><a href='/help/disulfid' target='_top'>More...</a></p>Disulfide bondi126 ↔ 1391 Publication
Disulfide bondi133 ↔ 1491 Publication
Glycosylationi236N-linked (GlcNAc...) asparagineSequence analysis1
Glycosylationi265N-linked (GlcNAc...) asparagine2 Publications1
Disulfide bondi284 ↔ 2961 Publication
Disulfide bondi290 ↔ 3061 Publication
Disulfide bondi297 ↔ 3141 Publication
Disulfide bondi307 ↔ 3211 Publication
Disulfide bondi315 ↔ 3281 Publication
Disulfide bondi322 ↔ 3351 Publication
Disulfide bondi366 ↔ 3781 Publication
Glycosylationi368N-linked (GlcNAc...) asparagine1 Publication1
Disulfide bondi372 ↔ 3881 Publication
Disulfide bondi397 ↔ 4101 Publication
Disulfide bondi404 ↔ 4161 Publication
Glycosylationi530N-linked (GlcNAc...) asparagine1 Publication1

<p>This subsection of the <a href="http://www.uniprot.org/help/ptm%5Fprocessing%5Fsection">PTM/processing</a> section describes post-translational modifications (PTMs). This subsection <strong>complements</strong> the information provided at the sequence level or describes modifications for which <strong>position-specific data is not yet available</strong>.<p><a href='/help/post-translational_modification' target='_top'>More...</a></p>Post-translational modificationi

Cleaved by ELANE; proteolysis is blocked by SLPI and is concentration- and time-dependent and induces CXCL8/IL-8 production; granulin-3 and granulin-4 are resistant to ELANE (PubMed:12526812, PubMed:28743268). Cleaved by CTSL in lysosome thus regulating the maturation and turnover of progranulin within the lysosome (PubMed:28743268).2 Publications

Keywords - PTMi

Disulfide bond, Glycoprotein

Proteomic databases

Encyclopedia of Proteome Dynamics

More...
EPDi
P28799

jPOST - Japan Proteome Standard Repository/Database

More...
jPOSTi
P28799

MassIVE - Mass Spectrometry Interactive Virtual Environment

More...
MassIVEi
P28799

MaxQB - The MaxQuant DataBase

More...
MaxQBi
P28799

PaxDb, a database of protein abundance averages across all three domains of life

More...
PaxDbi
P28799

PeptideAtlas

More...
PeptideAtlasi
P28799

PRoteomics IDEntifications database

More...
PRIDEi
P28799

ProteomicsDB: a multi-organism proteome resource

More...
ProteomicsDBi
54499 [P28799-1]
54500 [P28799-2]
81234

Consortium for Top Down Proteomics

More...
TopDownProteomicsi
P28799-2 [P28799-2]
P28799-3 [P28799-3]

PTM databases

GlyConnect protein glycosylation platform

More...
GlyConnecti
1290, 5 N-Linked glycans (1 site), 3 O-Linked glycans (1 site)

GlyGen: Computational and Informatics Resources for Glycoscience

More...
GlyGeni
P28799, 7 sites

iPTMnet integrated resource for PTMs in systems biology context

More...
iPTMneti
P28799

MetOSite database of methionine sulfoxide sites

More...
MetOSitei
P28799

Comprehensive resource for the study of protein post-translational modifications (PTMs) in human, mouse and rat.

More...
PhosphoSitePlusi
P28799

<p>This section provides information on the expression of a gene at the mRNA or protein level in cells or in tissues of multicellular organisms.<p><a href='/help/expression_section' target='_top'>More...</a></p>Expressioni

<p>This subsection of the 'Expression' section provides information on the expression of a gene at the mRNA or protein level in cells or in tissues of multicellular organisms. By default, the information is derived from experiments at the mRNA level, unless specified 'at protein level'.<br></br>Examples: <a href="http://www.uniprot.org/uniprot/P92958#expression">P92958</a>, <a href="http://www.uniprot.org/uniprot/Q8TDN4#expression">Q8TDN4</a>, <a href="http://www.uniprot.org/uniprot/O14734#expression">O14734</a><p><a href='/help/tissue_specificity' target='_top'>More...</a></p>Tissue specificityi

In myelogenous leukemic cell lines of promonocytic, promyelocytic, and proerythroid lineage, in fibroblasts, and very strongly in epithelial cell lines. Present in inflammatory cells and bone marrow. Highest levels in kidney.

<p>This subsection of the 'Expression' section reports the experimentally proven effects of inducers and repressors (usually chemical compounds or environmental factors) on the level of protein (or mRNA) expression (up-regulation, down-regulation, constitutive expression).<p><a href='/help/induction' target='_top'>More...</a></p>Inductioni

Increased in response to lysosome alkalization.1 Publication

Gene expression databases

Bgee dataBase for Gene Expression Evolution

More...
Bgeei
ENSG00000030582, Expressed in esophagus mucosa and 246 other tissues

ExpressionAtlas, Differential and Baseline Expression

More...
ExpressionAtlasi
P28799, baseline and differential

Genevisible search portal to normalized and curated expression data from Genevestigator

More...
Genevisiblei
P28799, HS

Organism-specific databases

Human Protein Atlas

More...
HPAi
ENSG00000030582, Low tissue specificity

<p>This section provides information on the quaternary structure of a protein and on interaction(s) with other proteins or protein complexes.<p><a href='/help/interaction_section' target='_top'>More...</a></p>Interactioni

<p>This subsection of the <a href="http://www.uniprot.org/help/interaction%5Fsection">'Interaction'</a> section provides information about the protein quaternary structure and interaction(s) with other proteins or protein complexes (with the exception of physiological receptor-ligand interactions which are annotated in the <a href="http://www.uniprot.org/help/function%5Fsection">'Function'</a> section).<p><a href='/help/subunit_structure' target='_top'>More...</a></p>Subunit structurei

Progranulin is secreted as a homodimer (PubMed:23364791).

Interacts with SLPI; interaction protects progranulin from proteolysis (PubMed:12526812).

Interacts (via region corresponding to granulin-7 peptide) with CTSD; stabilizes CTSD and increases its proteolytic activity (PubMed:28453791).

Interacts (via region corresponding to granulin-7 peptide) with SORT1; this interaction mediates endocytosis and lysosome delivery of progranulin; interaction occurs at the neuronal cell surface in a stressed nervous system (PubMed:21092856).

Interacts with PSAP; facilitates lysosomal delivery of progranulin from the extracellular space and the biosynthetic pathway (PubMed:26370502).

Forms a complex with PSAP and M6PR; PSAP bridges the binding between progranulin and M6PR (PubMed:26370502).

Forms a complex with PSAP and SORT1; progranulin bridges the interaction between PSAP and SORT1; facilitates lysosomal targeting of PSAP via SORT1; interaction enhances PSAP uptake in primary cortical neurons (PubMed:28541286).

Interacts (via regions corresponding to granulin-2 and granulin-7 peptides) with GBA; this interaction prevents aggregation of GBA-SCARB2 complex via interaction with HSPA1A upon stress (PubMed:27789271).

Interacts (via region corresponding to granulin-7 peptide) with HSPA1A; mediates recruitment of HSPA1A to GBA and prevents GBA aggregation in response to stress (PubMed:27789271).

7 Publications

<p>This subsection of the '<a href="http://www.uniprot.org/help/interaction%5Fsection">Interaction</a>' section provides information about binary protein-protein interactions. The data presented in this section are a quality-filtered subset of binary interactions automatically derived from the <a href="https://www.ebi.ac.uk/intact/">IntAct database</a>. It is updated at every <a href="http://www.uniprot.org/help/synchronization">UniProt release</a>.<p><a href='/help/binary_interactions' target='_top'>More...</a></p>Binary interactionsi

P28799
With#Exp.IntAct
ADAMTSL4 - isoform 3 [Q6UY14-3]3EBI-747754,EBI-10173507
AK3 [Q9UIJ7]3EBI-747754,EBI-3916527
ANAPC11 [Q9NYG5]3EBI-747754,EBI-2130187
APLN [D3DTF8]3EBI-747754,EBI-22002556
ARFGAP1 [Q8N6T3]3EBI-747754,EBI-716933
ARFGAP1 - isoform 3 [Q8N6T3-3]3EBI-747754,EBI-10694449
ASB3 - isoform 3 [Q9Y575-3]3EBI-747754,EBI-14199987
ASB9 - isoform 3 [Q96DX5-3]3EBI-747754,EBI-25843552
ASCL4 [Q6XD76]3EBI-747754,EBI-10254793
ASIC4 - isoform 3 [Q96FT7-4]3EBI-747754,EBI-9089489
BAG6 - isoform 2 [P46379-2]3EBI-747754,EBI-10988864
BAK1 [Q16611]3EBI-747754,EBI-519866
BAP18 [Q8IXM2]3EBI-747754,EBI-4280811
BECN1 [Q14457]3EBI-747754,EBI-949378
BMF [Q96LC9]3EBI-747754,EBI-3919268
BPESC1 [Q9GZL8]3EBI-747754,EBI-25861458
BRK1 [Q8WUW1]3EBI-747754,EBI-2837444
BTRC [Q9Y297]3EBI-747754,EBI-307461
C12orf74 [Q32Q52]3EBI-747754,EBI-12891828
C1orf216 [Q8TAB5]3EBI-747754,EBI-747505
C22orf39 [Q6P5X5]3EBI-747754,EBI-7317823
C22orf39 - isoform 2 [Q6P5X5-2]3EBI-747754,EBI-10692329
C4orf17 [Q53FE4]3EBI-747754,EBI-715110
C7orf31 [Q8N865]3EBI-747754,EBI-10174456
C7orf50 [Q9BRJ6]3EBI-747754,EBI-751612
CACNA1A [O00555]2EBI-747754,EBI-766279
CAMK1G [Q96NX5]3EBI-747754,EBI-3920838
CAPN15 [O75808]3EBI-747754,EBI-6149008
CCDC88C [A0A0A0MR69]3EBI-747754,EBI-12954453
CDK9 - isoform 2 [P50750-2]3EBI-747754,EBI-12029902
CHD1 - isoform 2 [O14646-2]3EBI-747754,EBI-10961487
CIAO2B [Q9Y3D0]3EBI-747754,EBI-744045
CITED2 [Q99967]3EBI-747754,EBI-937732
CLEC11A [Q9Y240]3EBI-747754,EBI-3957044
CLIP3 [Q96DZ5]3EBI-747754,EBI-12823145
CLPP [Q16740]3EBI-747754,EBI-1056029
CNPY3 [Q9BT09]3EBI-747754,EBI-2835965
CNST - isoform 3 [Q6PJW8-3]3EBI-747754,EBI-25836090
COPS3 [Q9UNS2]3EBI-747754,EBI-350590
CRCT1 [Q9UGL9]3EBI-747754,EBI-713677
CREB5 - isoform 3 [Q02930-3]3EBI-747754,EBI-10192698
CRY2 [Q49AN0]3EBI-747754,EBI-2212355
CSTA [P01040]3EBI-747754,EBI-724303
CTSD [P07339]4EBI-747754,EBI-2115097
CXCL5 [P42830]3EBI-747754,EBI-12175919
CXorf38 [Q8TB03]3EBI-747754,EBI-12024320
CYB5A [P00167]3EBI-747754,EBI-1047284
CYSRT1 [A8MQ03]3EBI-747754,EBI-3867333
DBN1 [Q16643]5EBI-747754,EBI-351394
DCAF8 - isoform 2 [Q5TAQ9-2]3EBI-747754,EBI-25842815
DLGAP2 - isoform 3 [Q9P1A6-3]3EBI-747754,EBI-12019838
DLX2 [Q07687]3EBI-747754,EBI-3908234
DMRT3 [Q9NQL9]3EBI-747754,EBI-9679045
DNASE1L1 [P49184]3EBI-747754,EBI-20894690
ECM1 [Q16610]3EBI-747754,EBI-947964
EED - isoform 2 [O75530-2]3EBI-747754,EBI-11132357
EIF5B [O60841]3EBI-747754,EBI-928530
ELAPOR1 - isoform 3 [Q6UXG2-3]3EBI-747754,EBI-12920100
EPS8L1 - isoform 3 [Q8TE68-3]3EBI-747754,EBI-21574901
EPS8L2 [Q9H6S3]3EBI-747754,EBI-3940939
FABP7 [O15540]3EBI-747754,EBI-10697159
FAF1 [Q9UNN5]3EBI-747754,EBI-718246
FAM111B [Q6SJ93]3EBI-747754,EBI-6309082
FAM131C [Q96AQ9]4EBI-747754,EBI-741921
FAM74A6 [Q5TZK3]3EBI-747754,EBI-10247271
FAM76B - isoform 3 [Q5HYJ3-3]6EBI-747754,EBI-11956087
FAM98C [Q17RN3]3EBI-747754,EBI-5461838
FAM9A [Q8IZU1]3EBI-747754,EBI-8468186
FANCL [Q9NW38]3EBI-747754,EBI-2339898
FASTKD1 [Q53R41]3EBI-747754,EBI-3957005
FBH1 [Q8NFZ0]3EBI-747754,EBI-724767
FBLN5 [Q9UBX5]3EBI-747754,EBI-947897
GATA1 - isoform 2 [P15976-2]3EBI-747754,EBI-9090198
GATA2 - isoform 2 [P23769-2]3EBI-747754,EBI-21856389
GFOD1 [Q9NXC2]3EBI-747754,EBI-8799578
GLI4 [P10075]3EBI-747754,EBI-14061927
GLRX3 [O76003]9EBI-747754,EBI-374781
GNE - isoform 2 [Q9Y223-2]6EBI-747754,EBI-11975289
GOLGA2P5 [Q9HBQ8]3EBI-747754,EBI-22000587
GPR141 [Q7Z602]3EBI-747754,EBI-21649723
GPSM3 [Q9Y4H4]3EBI-747754,EBI-347538
H2AP [O75409]3EBI-747754,EBI-6447217
H3-5 [Q6NXT2]3EBI-747754,EBI-2868501
H3C12 [P68431]3EBI-747754,EBI-79722
hCG_2001421 [A8K0U2]3EBI-747754,EBI-25843825
HHEX [Q03014]3EBI-747754,EBI-747421
HOXA1 [P49639]18EBI-747754,EBI-740785
HOXC4 [P09017]3EBI-747754,EBI-3923226
IGFBP4 [P22692]3EBI-747754,EBI-2831948
IL16 - isoform 2 [Q14005-2]3EBI-747754,EBI-17178971
ILF3 [Q9NXX0]3EBI-747754,EBI-743980
ING4 [Q9UNL4]3EBI-747754,EBI-2866661
IQCF2 [Q8IXL9]3EBI-747754,EBI-10238842
IRAG1 - isoform 3 [Q9Y6F6-3]3EBI-747754,EBI-25840037
ISCA2 [Q86U28]3EBI-747754,EBI-10258659
KEAP1 [Q14145]3EBI-747754,EBI-751001
KIF1A [Q12756]3EBI-747754,EBI-2679809
KLF15 [Q9UIH9]3EBI-747754,EBI-2796400
KLF3 [P57682]3EBI-747754,EBI-8472267
KLHL20 [Q9Y2M5]3EBI-747754,EBI-714379
KRT34 [O76011]3EBI-747754,EBI-1047093
KRTAP1-1 [Q07627]3EBI-747754,EBI-11959885
KRTAP1-5 [Q9BYS1]3EBI-747754,EBI-11741292
KRTAP10-7 [P60409]3EBI-747754,EBI-10172290
KRTAP10-8 [P60410]3EBI-747754,EBI-10171774
KRTAP11-1 [Q8IUC1]3EBI-747754,EBI-1052037
KRTAP12-1 [P59990]3EBI-747754,EBI-10210845
KRTAP13-2 [Q52LG2]3EBI-747754,EBI-11953846
KRTAP13-3 [Q3SY46]3EBI-747754,EBI-10241252
KRTAP15-1 [Q3LI76]3EBI-747754,EBI-11992140
KRTAP19-7 [Q3SYF9]3EBI-747754,EBI-10241353
KRTAP26-1 [Q6PEX3]8EBI-747754,EBI-3957672
KRTAP5-9 [P26371]3EBI-747754,EBI-3958099
KRTAP6-1 [Q3LI64]3EBI-747754,EBI-12111050
KRTAP6-2 [Q3LI66]3EBI-747754,EBI-11962084
KRTAP8-1 [Q8IUC2]3EBI-747754,EBI-10261141
LASP1 - isoform 2 [Q14847-2]3EBI-747754,EBI-9088686
LCA5L [O95447]3EBI-747754,EBI-8473670
LCE1A [Q5T7P2]3EBI-747754,EBI-11962058
LCE1B [Q5T7P3]3EBI-747754,EBI-10245913
LCE1D [Q5T752]3EBI-747754,EBI-11741311
LCE1E [Q5T753]3EBI-747754,EBI-11955335
LCE2A [Q5TA79]3EBI-747754,EBI-10246607
LCE2B [O14633]3EBI-747754,EBI-11478468
LCE2D [Q5TA82]3EBI-747754,EBI-10246750
LCE3C [Q5T5A8]3EBI-747754,EBI-10245291
LCE3E [Q5T5B0]3EBI-747754,EBI-10245456
LCE4A [Q5TA78]4EBI-747754,EBI-10246358
LHX6 [Q9UPM6]3EBI-747754,EBI-10258746
LHX8 [Q68G74]3EBI-747754,EBI-8474075
LOC401296 [A2RU56]3EBI-747754,EBI-9088215
LOXL4 [Q96JB6]3EBI-747754,EBI-749562
LPIN1 [Q14693]3EBI-747754,EBI-5278370
LVRN - isoform 4 [Q6Q4G3-4]3EBI-747754,EBI-25862057
MALT1 - isoform 2 [Q9UDY8-2]3EBI-747754,EBI-12056869
MAP1LC3B [Q9GZQ8]3EBI-747754,EBI-373144
MAP3K5 [Q99683]3EBI-747754,EBI-476263
MAX - isoform 4 [P61244-4]3EBI-747754,EBI-25848049
MBD4 - isoform 2 [O95243-2]3EBI-747754,EBI-6448717
MEOX2 [Q6FHY5]3EBI-747754,EBI-16439278
MNDA [P41218]3EBI-747754,EBI-2829677
MOGAT3 - isoform 3 [Q86VF5-3]3EBI-747754,EBI-25840143
MRPS17 [Q9Y2R5]3EBI-747754,EBI-1046443
MSH5 - isoform 4 [O43196-4]3EBI-747754,EBI-25860238
MSRB3 - isoform 2 [Q8IXL7-2]3EBI-747754,EBI-10699187
MYLPF [Q96A32]3EBI-747754,EBI-1390771
MYNN [Q9NPC7]3EBI-747754,EBI-3446748
NACAD [O15069]3EBI-747754,EBI-7108375
NDN [Q99608]3EBI-747754,EBI-718177
NDUFAF4 [Q9P032]3EBI-747754,EBI-2606839
NFX1 [Q12986]3EBI-747754,EBI-2130062
NGEF [Q8N5V2]3EBI-747754,EBI-718372
NLK [Q9UBE8]7EBI-747754,EBI-366978
NLRP1 [Q96AM0]3EBI-747754,EBI-25860999
NOTCH1 [Q6IAD4]3EBI-747754,EBI-25860267
NR1D2 [Q14995]3EBI-747754,EBI-6144053
NUFIP2 [Q7Z417]10EBI-747754,EBI-1210753
OIP5 [O43482]3EBI-747754,EBI-536879
OTUB1 [Q96FW1]3EBI-747754,EBI-1058491
OTX1 [P32242]10EBI-747754,EBI-740446
P2RY6 [Q15077]3EBI-747754,EBI-10235794
P4HB [P07237]4EBI-747754,EBI-395883
PALM - isoform 2 [O75781-2]3EBI-747754,EBI-16399860
PARP11 - isoform 4 [Q9NR21-5]3EBI-747754,EBI-17159452
PCGF5 - isoform 2 [Q86SE9-2]3EBI-747754,EBI-25861637
PER1 [O15534]3EBI-747754,EBI-2557276
PERP [Q96FX8]3EBI-747754,EBI-17183069
PGLYRP3 [Q96LB9]3EBI-747754,EBI-12339509
PHF7 [Q9BWX1]3EBI-747754,EBI-4307517
PHLDA1 [A2BDE7]3EBI-747754,EBI-14084211
PIAS1 [O75925]3EBI-747754,EBI-629434
PLA2G12A [Q9BZM1]3EBI-747754,EBI-3916751
PLEKHG4 - isoform 2 [Q58EX7-2]3EBI-747754,EBI-21503705
PLLP [Q9Y342]3EBI-747754,EBI-3919291
PLPPR1 [Q8TBJ4]3EBI-747754,EBI-18063495
POLR3F [Q9H1D9]3EBI-747754,EBI-710067
POU4F2 [Q12837]6EBI-747754,EBI-17236143
PP9974 [P09565]3EBI-747754,EBI-10196507
PRKAA2 [P54646]3EBI-747754,EBI-1383852
PRKAB2 [O43741]4EBI-747754,EBI-1053424
PRPS2 [P11908]3EBI-747754,EBI-4290895
PSAP [P07602]5EBI-747754,EBI-716699
PSMB10 [P40306]3EBI-747754,EBI-603329
PSMB8 - isoform 2 [P28062-2]3EBI-747754,EBI-372312
PTMA [Q8TBK9]3EBI-747754,EBI-1056327
PTPMT1 [Q8WUK0]3EBI-747754,EBI-7199479
PUM1 [Q14671]3EBI-747754,EBI-948453
PXN [Q7Z7K5]3EBI-747754,EBI-25841978
QARS1 [P47897]3EBI-747754,EBI-347462
RBM14 [Q96PK6]3EBI-747754,EBI-954272
RCHY1 - isoform 4 [Q96PM5-4]3EBI-747754,EBI-21252376
RHPN1 [Q8TCX5]3EBI-747754,EBI-746325
RNF112 [Q9ULX5]3EBI-747754,EBI-25829984
RNF138 [Q8WVD3]3EBI-747754,EBI-749039
RNF14 [Q9UBS8]3EBI-747754,EBI-2130308
RNF208 [Q9H0X6]3EBI-747754,EBI-751555
RPS15A [P62244]3EBI-747754,EBI-347895
RUSC1-AS1 [Q66K80]3EBI-747754,EBI-10248967
RYBP [Q8N488]3EBI-747754,EBI-752324
SCAMP4 [Q969E2]3EBI-747754,EBI-4403649
SDC2 [P34741]3EBI-747754,EBI-1172957
SEM1 [P60896]3EBI-747754,EBI-79819
SEMA4G - isoform 3 [Q9NTN9-3]3EBI-747754,EBI-9089805
SEPTIN6 [Q14141]3EBI-747754,EBI-745901
SERINC3 [Q13530]3EBI-747754,EBI-1045571
SGTA [O43765]7EBI-747754,EBI-347996
SHFL - isoform 3 [Q9NUL5-3]3EBI-747754,EBI-22000547
SHOX2 - isoform 3 [O60902-3]3EBI-747754,EBI-9092164
SMAD9 - isoform B [O15198-2]3EBI-747754,EBI-12273450
SMCP [P49901]3EBI-747754,EBI-750494
SMURF1 - isoform Short [Q9HCE7-2]3EBI-747754,EBI-9845742
SNRNP40 [Q96DI7]3EBI-747754,EBI-538492
SORT1 [Q99523]3EBI-747754,EBI-1057058
SPATA3 [A0A024R4B0]3EBI-747754,EBI-14123856
SPATA8 [Q6RVD6]3EBI-747754,EBI-8635958
SPINK2 [P20155]3EBI-747754,EBI-10200479
SPRED2 [Q7Z698]3EBI-747754,EBI-7082156
SPRY2 [O43597]3EBI-747754,EBI-742487
SPRY4 [Q9C004]3EBI-747754,EBI-354861
SPSB3 [Q6PJ21]3EBI-747754,EBI-3937206
STUB1 [Q9UNE7]3EBI-747754,EBI-357085
SUMF2 [Q8NBJ7]3EBI-747754,EBI-723091
SYT16 - isoform 3 [Q17RD7-3]3EBI-747754,EBI-25861603
TASOR2 [Q5VWN6]3EBI-747754,EBI-745958
TAT [P17735]3EBI-747754,EBI-12046643
TAX1BP1 [Q86VP1]3EBI-747754,EBI-529518
TBPL1 [P62380]3EBI-747754,EBI-716225
TCEAL8 [Q8IYN2]3EBI-747754,EBI-2116184
TDG [Q13569]3EBI-747754,EBI-348333
TEAD1 - isoform 2 [P28347-2]3EBI-747754,EBI-12151837
TEX19 [Q8NA77]3EBI-747754,EBI-13323487
TIMM17B [O60830]3EBI-747754,EBI-2372529
TLE1 [Q04724]3EBI-747754,EBI-711424
TLE5 - isoform 2 [Q08117-2]3EBI-747754,EBI-11741437
TMEM61 [Q8N0U2]3EBI-747754,EBI-25830583
tmp_locus_54 [Q53NU3]3EBI-747754,EBI-10242677
TMUB2 - isoform 4 [Q71RG4-4]3EBI-747754,EBI-25831574
TNFRSF1A [P19438]4EBI-747754,EBI-299451
TNFRSF1B [P20333]5EBI-747754,EBI-358983
TRIM35 - isoform 2 [Q9UPQ4-2]3EBI-747754,EBI-17716262
TRMT61B [Q9BVS5]3EBI-747754,EBI-3197877
TRNT1 - isoform 3 [Q96Q11-3]3EBI-747754,EBI-25861172
TSC22D4 [Q9Y3Q8]3EBI-747754,EBI-739485
TSPAN4 [O14817]3EBI-747754,EBI-8652667
TSSC4 [A0A024RCB9]3EBI-747754,EBI-25860845
TTC1 [Q99614]3EBI-747754,EBI-742074
TTC23 - isoform 3 [Q5W5X9-3]3EBI-747754,EBI-9090990
TUT7 - isoform 5 [Q5VYS8-5]3EBI-747754,EBI-9088812
UTP23 [Q9BRU9]3EBI-747754,EBI-5457544
VASN [Q6EMK4]3EBI-747754,EBI-10249550
VDAC2 [P45880]3EBI-747754,EBI-354022
VPS37A [Q8NEZ2]3EBI-747754,EBI-2850578
VPS37A - isoform 2 [Q8NEZ2-2]3EBI-747754,EBI-10270911
VSX2 [P58304]3EBI-747754,EBI-6427899
WDR61 [Q9GZS3]3EBI-747754,EBI-358545
WWOX - isoform 5 [Q9NZC7-5]3EBI-747754,EBI-12040603
YAF2 - isoform 2 [Q8IY57-5]3EBI-747754,EBI-12111538
YIF1A [O95070]3EBI-747754,EBI-2799703
YY1 [P25490]4EBI-747754,EBI-765538
ZBTB24 - isoform 2 [O43167-2]3EBI-747754,EBI-25842419
ZFP64 [Q9NTW7]3EBI-747754,EBI-711679
ZKSCAN8 [Q15776]3EBI-747754,EBI-2602314
ZNF124 [Q15973]3EBI-747754,EBI-2555767
ZNF138 [P52744]3EBI-747754,EBI-10746567
ZNF180 - isoform 4 [Q9UJW8-4]3EBI-747754,EBI-12055755
ZNF239 [Q16600]3EBI-747754,EBI-8787052
ZNF296 [Q8WUU4]3EBI-747754,EBI-8834821
ZNF366 [Q8N895]3EBI-747754,EBI-2813661
ZNF444 - isoform 2 [Q8N0Y2-2]3EBI-747754,EBI-12010736
ZNF488 - isoform 2 [Q96MN9-2]3EBI-747754,EBI-25831733
ZNF497 [Q6ZNH5]3EBI-747754,EBI-10486136
ZNF524 [Q96C55]3EBI-747754,EBI-10283126
ZNF57 [Q68EA5]3EBI-747754,EBI-8490788
ZNF572 [Q7Z3I7]3EBI-747754,EBI-10172590
ZNF625 - isoform 2 [Q96I27-2]3EBI-747754,EBI-12038525
ZNF641 - isoform 2 [Q96N77-2]3EBI-747754,EBI-12939666
ZNF670 [Q9BS34]3EBI-747754,EBI-745276
ZNF696 [Q9H7X3]3EBI-747754,EBI-11090299
ZNF697 [Q5TEC3]3EBI-747754,EBI-25845217
ZNF774 [Q6NX45]3EBI-747754,EBI-10251462
ZNF791 [Q3KP31]3EBI-747754,EBI-2849119
ZSCAN26 [Q16670]3EBI-747754,EBI-3920053
ZSCAN9 [O15535]3EBI-747754,EBI-751531
A0A384ME253EBI-747754,EBI-10211777
Q7L8T73EBI-747754,EBI-25831943
Q7Z7833EBI-747754,EBI-9088990
Hoxa1 [P09022] from Mus musculus.2EBI-747754,EBI-3957603
Isoform 2 [P28799-2]
With#Exp.IntAct
APLN [D3DTF8]3EBI-25860013,EBI-22002556
ATP5IF1 [Q9UII2]3EBI-25860013,EBI-718459
CCDC26 [Q8TAB7]3EBI-25860013,EBI-10271580
CDK9 - isoform 2 [P50750-2]3EBI-25860013,EBI-12029902
CREB5 - isoform 3 [Q02930-3]3EBI-25860013,EBI-10192698
DLK1 [P80370]3EBI-25860013,EBI-21555397
DPYSL4 [O14531]3EBI-25860013,EBI-719542
DVL3 [Q92997]3EBI-25860013,EBI-739789
FABP7 [O15540]3EBI-25860013,EBI-10697159
FAM131C [Q96AQ9]3EBI-25860013,EBI-741921
FAM76B - isoform 3 [Q5HYJ3-3]3EBI-25860013,EBI-11956087
hCG_1820408 [Q8N7T0]3EBI-25860013,EBI-25858908
HOXA1 [P49639]3EBI-25860013,EBI-740785
LCE2A [Q5TA79]3EBI-25860013,EBI-10246607
MSRB3 - isoform 2 [Q8IXL7-2]3EBI-25860013,EBI-10699187
NR1D2 [Q14995]3EBI-25860013,EBI-6144053
PP9974 [P09565]3EBI-25860013,EBI-10196507
PUM1 [Q14671]3EBI-25860013,EBI-948453
PXN [Q7Z7K5]3EBI-25860013,EBI-25841978
SCAMP4 [Q969E2]3EBI-25860013,EBI-4403649
SDC2 [P34741]3EBI-25860013,EBI-1172957
SIRT3 [Q9NTG7]3EBI-25860013,EBI-724621
SOX14 [O95416]3EBI-25860013,EBI-9087806
SPATA3 [A0A024R4B0]3EBI-25860013,EBI-14123856
SPRED1 [Q7Z699]3EBI-25860013,EBI-5235340
TAT [P17735]3EBI-25860013,EBI-12046643
TCP10L [Q8TDR4]3EBI-25860013,EBI-3923210
TMUB2 - isoform 4 [Q71RG4-4]3EBI-25860013,EBI-25831574
TP53TG5 [Q9Y2B4]3EBI-25860013,EBI-21870909
YY1 [P25490]3EBI-25860013,EBI-765538
ZFHX2 [Q9C0A1]3EBI-25860013,EBI-25850811
ZNF180 - isoform 4 [Q9UJW8-4]3EBI-25860013,EBI-12055755
ZNF366 [Q8N895]3EBI-25860013,EBI-2813661
A0A087WZY13EBI-25860013,EBI-13387614
Q7L8T73EBI-25860013,EBI-25831943
Granulin-1 (PRO_0000012695)
With#Exp.IntAct
CTSD [P07339]2EBI-21335602,EBI-2115097
Granulin-2 (PRO_0000012696)
With#Exp.IntAct
CTSD [P07339]2EBI-21335615,EBI-2115097
Granulin-3 (PRO_0000012697)
With#Exp.IntAct
CTSD [P07339]2EBI-21335629,EBI-2115097
Granulin-4 (PRO_0000012698)
With#Exp.IntAct
CTSD [P07339]2EBI-21335642,EBI-2115097
Granulin-5 (PRO_0000012699)
With#Exp.IntAct
CTSD [P07339]2EBI-21335656,EBI-2115097
Granulin-6 (PRO_0000012700)
With#Exp.IntAct
CTSD [P07339]2EBI-21335669,EBI-2115097
Granulin-7 (PRO_0000012701)
With#Exp.IntAct
CTSD [P07339]2EBI-21335682,EBI-2115097

GO - Molecular functioni

Protein-protein interaction databases

The Biological General Repository for Interaction Datasets (BioGRID)

More...
BioGRIDi
109153, 170 interactors

CORUM comprehensive resource of mammalian protein complexes

More...
CORUMi
P28799

Database of interacting proteins

More...
DIPi
DIP-41742N

Protein interaction database and analysis system

More...
IntActi
P28799, 421 interactors

Molecular INTeraction database

More...
MINTi
P28799

STRING: functional protein association networks

More...
STRINGi
9606.ENSP00000053867

Miscellaneous databases

RNAct, Protein-RNA interaction predictions for model organisms.

More...
RNActi
P28799, protein

<p>This section provides information on the tertiary and secondary structure of a protein.<p><a href='/help/structure_section' target='_top'>More...</a></p>Structurei

Secondary structure

1593
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details

3D structure databases

SWISS-MODEL Repository - a database of annotated 3D protein structure models

More...
SMRi
P28799

Database of comparative protein structure models

More...
ModBasei
Search...

Protein Data Bank in Europe - Knowledge Base

More...
PDBe-KBi
Search...

Miscellaneous databases

Relative evolutionary importance of amino acids within a protein sequence

More...
EvolutionaryTracei
P28799

<p>This section provides information on sequence similarities with other proteins and the domain(s) present in a protein.<p><a href='/help/family_and_domains_section' target='_top'>More...</a></p>Family & Domainsi

<p>This subsection of the 'Family and domains' section provides information about the sequence similarity with other proteins.<p><a href='/help/sequence_similarities' target='_top'>More...</a></p>Sequence similaritiesi

Belongs to the granulin family.Curated

Keywords - Domaini

Repeat, Signal

Phylogenomic databases

evolutionary genealogy of genes: Non-supervised Orthologous Groups

More...
eggNOGi
KOG4296, Eukaryota

Ensembl GeneTree

More...
GeneTreei
ENSGT00470000042293

The HOGENOM Database of Homologous Genes from Fully Sequenced Organisms

More...
HOGENOMi
CLU_026274_0_0_1

InParanoid: Eukaryotic Ortholog Groups

More...
InParanoidi
P28799

Identification of Orthologs from Complete Genome Data

More...
OMAi
PQNTVCN

Database of Orthologous Groups

More...
OrthoDBi
162721at2759

Database for complete collections of gene phylogenies

More...
PhylomeDBi
P28799

TreeFam database of animal gene trees

More...
TreeFami
TF319678

Family and domain databases

Gene3D Structural and Functional Annotation of Protein Families

More...
Gene3Di
3.10.20.500, 7 hits

Integrated resource of protein families, domains and functional sites

More...
InterProi
View protein in InterPro
IPR000118, Granulin
IPR039036, Granulin_fam
IPR037277, Granulin_sf

The PANTHER Classification System

More...
PANTHERi
PTHR12274, PTHR12274, 1 hit

Pfam protein domain database

More...
Pfami
View protein in Pfam
PF00396, Granulin, 7 hits

Simple Modular Architecture Research Tool; a protein domain database

More...
SMARTi
View protein in SMART
SM00277, GRAN, 7 hits

PROSITE; a protein domain and family database

More...
PROSITEi
View protein in PROSITE
PS00799, GRANULINS, 7 hits

<p>This section displays by default the canonical protein sequence and upon request all isoforms described in the entry. It also includes information pertinent to the sequence(s), including <a href="http://www.uniprot.org/help/sequence%5Flength">length</a> and <a href="http://www.uniprot.org/help/sequences">molecular weight</a>. The information is filed in different subsections. The current subsections and their content are listed below:<p><a href='/help/sequences_section' target='_top'>More...</a></p>Sequences (3+)i

<p>This subsection of the <a href="http://www.uniprot.org/help/sequences%5Fsection">Sequence</a> section indicates if the <a href="http://www.uniprot.org/help/canonical%5Fand%5Fisoforms">canonical sequence</a> displayed by default in the entry is complete or not.<p><a href='/help/sequence_status' target='_top'>More...</a></p>Sequence statusi: Complete.

<p>This subsection of the <a href="http://www.uniprot.org/help/sequences%5Fsection">Sequence</a> section indicates if the <a href="http://www.uniprot.org/help/canonical%5Fand%5Fisoforms">canonical sequence</a> displayed by default in the entry is in its mature form or if it represents the precursor.<p><a href='/help/sequence_processing' target='_top'>More...</a></p>Sequence processingi: The displayed sequence is further processed into a mature form.

This entry describes 3 <p>This subsection of the 'Sequence' section lists the alternative protein sequences (isoforms) that can be generated from the same gene by a single or by the combination of up to four biological events (alternative promoter usage, alternative splicing, alternative initiation and ribosomal frameshifting). Additionally, this section gives relevant information on each alternative protein isoform.<p><a href='/help/alternative_products' target='_top'>More...</a></p> isoformsi produced by alternative splicing. AlignAdd to basket

This entry has 3 described isoforms and 14 potential isoforms that are computationally mapped.Show allAlign All

Isoform 1 (identifier: P28799-1) [UniParc]FASTAAdd to basket

This isoform has been chosen as the <div> <p><b>What is the canonical sequence?</b><p><a href='/help/canonical_and_isoforms' target='_top'>More...</a></p>canonicali sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

« Hide
        10         20         30         40         50
MWTLVSWVAL TAGLVAGTRC PDGQFCPVAC CLDPGGASYS CCRPLLDKWP
60 70 80 90 100
TTLSRHLGGP CQVDAHCSAG HSCIFTVSGT SSCCPFPEAV ACGDGHHCCP
110 120 130 140 150
RGFHCSADGR SCFQRSGNNS VGAIQCPDSQ FECPDFSTCC VMVDGSWGCC
160 170 180 190 200
PMPQASCCED RVHCCPHGAF CDLVHTRCIT PTGTHPLAKK LPAQRTNRAV
210 220 230 240 250
ALSSSVMCPD ARSRCPDGST CCELPSGKYG CCPMPNATCC SDHLHCCPQD
260 270 280 290 300
TVCDLIQSKC LSKENATTDL LTKLPAHTVG DVKCDMEVSC PDGYTCCRLQ
310 320 330 340 350
SGAWGCCPFT QAVCCEDHIH CCPAGFTCDT QKGTCEQGPH QVPWMEKAPA
360 370 380 390 400
HLSLPDPQAL KRDVPCDNVS SCPSSDTCCQ LTSGEWGCCP IPEAVCCSDH
410 420 430 440 450
QHCCPQGYTC VAEGQCQRGS EIVAGLEKMP ARRASLSHPR DIGCDQHTSC
460 470 480 490 500
PVGQTCCPSL GGSWACCQLP HAVCCEDRQH CCPAGYTCNV KARSCEKEVV
510 520 530 540 550
SAQPATFLAR SPHVGVKDVE CGEGHFCHDN QTCCRDNRQG WACCPYRQGV
560 570 580 590
CCADRRHCCP AGFRCAARGT KCLRREAPRW DAPLRDPALR QLL
Length:593
Mass (Da):63,544
Last modified:October 11, 2005 - v2
<p>The checksum is a form of redundancy check that is calculated from the sequence. It is useful for tracking sequence updates.</p> <p>It should be noted that while, in theory, two different sequences could have the same checksum value, the likelihood that this would happen is extremely low.</p> <p>However UniProtKB may contain entries with identical sequences in case of multiple genes (paralogs).</p> <p>The checksum is computed as the sequence 64-bit Cyclic Redundancy Check value (CRC64) using the generator polynomial: x<sup>64</sup> + x<sup>4</sup> + x<sup>3</sup> + x + 1. The algorithm is described in the ISO 3309 standard. </p> <p class="publication">Press W.H., Flannery B.P., Teukolsky S.A. and Vetterling W.T.<br /> <strong>Cyclic redundancy and other checksums</strong><br /> <a href="http://www.nrbook.com/b/bookcpdf.php">Numerical recipes in C 2nd ed., pp896-902, Cambridge University Press (1993)</a>)</p> Checksum:i4E5947F1B4EDE619
GO
Isoform 2 (identifier: P28799-2) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     377-531: Missing.

Show »
Length:438
Mass (Da):46,991
Checksum:i558B72CBAF3B991F
GO
Isoform 3 (identifier: P28799-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-71: MWTLVSWVAL...QVDAHCSAGH → MAITAAHGAS...LPAVKTGCTA
     72-251: Missing.

Show »
Length:413
Mass (Da):44,132
Checksum:i0E3767A44BE314EC
GO

<p>In eukaryotic reference proteomes, unreviewed entries that are likely to belong to the same gene are computationally mapped, based on gene identifiers from Ensembl, EnsemblGenomes and model organism databases.<p><a href='/help/gene_centric_isoform_mapping' target='_top'>More...</a></p>Computationally mapped potential isoform sequencesi

There are 14 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
EntryEntry nameProtein names
Gene namesLengthAnnotation
K7EQ05K7EQ05_HUMAN
Progranulin
GRN
436Annotation score:

Annotation score:2 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7EKL3K7EKL3_HUMAN
Progranulin
GRN
388Annotation score:

Annotation score:2 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7EPL0K7EPL0_HUMAN
Progranulin
GRN
191Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7EMR1K7EMR1_HUMAN
Progranulin
GRN
172Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7EQK6K7EQK6_HUMAN
Progranulin
GRN
142Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7ERM2K7ERM2_HUMAN
Progranulin
GRN
189Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7EJY4K7EJY4_HUMAN
Progranulin
GRN
163Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7EK92K7EK92_HUMAN
Progranulin
GRN
136Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7EM89K7EM89_HUMAN
Progranulin
GRN
149Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
K7ENI2K7ENI2_HUMAN
Progranulin
GRN
111Annotation score:

Annotation score:1 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
There are more potential isoformsShow all

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'Sequence' section reports difference(s) between the canonical sequence (displayed by default in the entry) and the different sequence submissions merged in the entry. These various submissions may originate from different sequencing projects, different types of experiments, or different biological samples. Sequence conflicts are usually of unknown origin.<p><a href='/help/conflict' target='_top'>More...</a></p>Sequence conflicti219S → H AA sequence (PubMed:2268320).Curated1
Sequence conflicti290C → S AA sequence (PubMed:8471426).Curated1
Sequence conflicti386W → H AA sequence (PubMed:2268320).Curated1
Sequence conflicti407G → R in AAA58617 (PubMed:1542665).Curated1
Sequence conflicti428K → E in BAD96242 (Ref. 8) Curated1
Sequence conflicti434A → G in AAA58617 (PubMed:1542665).Curated1
Sequence conflicti454Q → G in AAA58617 (PubMed:1542665).Curated1
Sequence conflicti460L → Q in AAA58617 (PubMed:1542665).Curated1
Sequence conflicti547R → A in AAA58617 (PubMed:1542665).Curated1
Sequence conflicti567A → R in AAA58617 (PubMed:1542665).Curated1

Natural variant

Feature keyPosition(s)DescriptionActionsGraphical viewLength
Natural variantiVAR_0444519A → D in UP-FTD; no significant difference in the total mRNA between cases and controls; although the mutant protein is expressed it is not secreted and appears to be trapped within an intracellular compartment. 2 PublicationsCorresponds to variant dbSNP:rs63751243EnsemblClinVar.1
Natural variantiVAR_06462519R → W1 PublicationCorresponds to variant dbSNP:rs63750723EnsemblClinVar.1
Natural variantiVAR_06462655R → W1 PublicationCorresponds to variant dbSNP:rs1555610922Ensembl.1
Natural variantiVAR_06462769A → T1 PublicationCorresponds to variant dbSNP:rs199944486Ensembl.1
Natural variantiVAR_064628119Missing 1 PublicationCorresponds to variant dbSNP:rs758168578Ensembl.1
Natural variantiVAR_064629120S → Y1 PublicationCorresponds to variant dbSNP:rs63750043EnsemblClinVar.1
Natural variantiVAR_064630182T → M1 PublicationCorresponds to variant dbSNP:rs63750479EnsemblClinVar.1
Natural variantiVAR_064631221C → S1 PublicationCorresponds to variant dbSNP:rs758322775Ensembl.1
Natural variantiVAR_064632275P → L1 PublicationCorresponds to variant dbSNP:rs529849967Ensembl.1
Natural variantiVAR_064633376D → N1 PublicationCorresponds to variant dbSNP:rs143030899EnsemblClinVar.1
Natural variantiVAR_064634398S → L1 PublicationCorresponds to variant dbSNP:rs148213321Ensembl.1
Natural variantiVAR_064635433R → Q1 PublicationCorresponds to variant dbSNP:rs114248177EnsemblClinVar.1
Natural variantiVAR_014830515G → A1 PublicationCorresponds to variant dbSNP:rs25647EnsemblClinVar.1
Natural variantiVAR_064636564R → H1 PublicationCorresponds to variant dbSNP:rs971443926Ensembl.1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'Sequence' section describes the sequence of naturally occurring alternative protein isoform(s). The changes in the amino acid sequence may be due to alternative splicing, alternative promoter usage, alternative initiation, or ribosomal frameshifting.<p><a href='/help/var_seq' target='_top'>More...</a></p>Alternative sequenceiVSP_0534721 – 71MWTLV…CSAGH → MAITAAHGASTAVQTGDPAS KDQVTTPWVPSSALIVSSNA RTSPRAVLWSMAPGGAAPCP RLPAVKTGCTA in isoform 3. 1 PublicationAdd BLAST71
Alternative sequenceiVSP_05347372 – 251Missing in isoform 3. 1 PublicationAdd BLAST180
Alternative sequenceiVSP_001837377 – 531Missing in isoform 2. 2 PublicationsAdd BLAST155

Sequence databases

Select the link destinations:

EMBL nucleotide sequence database

More...
EMBLi

GenBank nucleotide sequence database

More...
GenBanki

DNA Data Bank of Japan; a nucleotide sequence database

More...
DDBJi
Links Updated
X62320 mRNA Translation: CAA44196.1
AF055008 mRNA Translation: AAC09359.1
M75161 mRNA Translation: AAA58617.1
AY124489 mRNA Translation: AAM94026.1
BT006844 mRNA Translation: AAP35490.1
AK023348 mRNA Translation: BAB14535.1
AK222522 mRNA Translation: BAD96242.1
CH471178 Genomic DNA Translation: EAW51599.1
CH471178 Genomic DNA Translation: EAW51600.1
BC000324 mRNA Translation: AAH00324.1
BC010577 mRNA Translation: AAH10577.1

The Consensus CDS (CCDS) project

More...
CCDSi
CCDS11483.1 [P28799-1]

Protein sequence database of the Protein Information Resource

More...
PIRi
JC1284, GYHU

NCBI Reference Sequences

More...
RefSeqi
NP_002078.1, NM_002087.3 [P28799-1]
XP_005257310.1, XM_005257253.1 [P28799-1]

Genome annotation databases

Ensembl eukaryotic genome annotation project

More...
Ensembli
ENST00000053867; ENSP00000053867; ENSG00000030582 [P28799-1]
ENST00000639447; ENSP00000492014; ENSG00000030582 [P28799-2]

Database of genes from NCBI RefSeq genomes

More...
GeneIDi
2896

KEGG: Kyoto Encyclopedia of Genes and Genomes

More...
KEGGi
hsa:2896

UCSC genome browser

More...
UCSCi
uc002igp.2, human [P28799-1]

Keywords - Coding sequence diversityi

Alternative splicing

<p>This section provides links to proteins that are similar to the protein sequence(s) described in this entry at different levels of sequence identity thresholds (100%, 90% and 50%) based on their membership in UniProt Reference Clusters (<a href="http://www.uniprot.org/help/uniref">UniRef</a>).<p><a href='/help/similar_proteins_section' target='_top'>More...</a></p>Similar proteinsi

<p>This section is used to point to information related to entries and found in data collections other than UniProtKB.<p><a href='/help/cross_references_section' target='_top'>More...</a></p>Cross-referencesi

<p>This subsection of the <a href="http://www.uniprot.org/manual/cross%5Freferences%5Fsection">Cross-references</a> section provides links to various web resources that are relevant for a specific protein.<p><a href='/help/web_resource' target='_top'>More...</a></p>Web resourcesi

Atlas of Genetics and Cytogenetics in Oncology and Haematology

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
X62320 mRNA Translation: CAA44196.1
AF055008 mRNA Translation: AAC09359.1
M75161 mRNA Translation: AAA58617.1
AY124489 mRNA Translation: AAM94026.1
BT006844 mRNA Translation: AAP35490.1
AK023348 mRNA Translation: BAB14535.1
AK222522 mRNA Translation: BAD96242.1
CH471178 Genomic DNA Translation: EAW51599.1
CH471178 Genomic DNA Translation: EAW51600.1
BC000324 mRNA Translation: AAH00324.1
BC010577 mRNA Translation: AAH10577.1
CCDSiCCDS11483.1 [P28799-1]
PIRiJC1284, GYHU
RefSeqiNP_002078.1, NM_002087.3 [P28799-1]
XP_005257310.1, XM_005257253.1 [P28799-1]

3D structure databases

Select the link destinations:

Protein Data Bank Europe

More...
PDBei

Protein Data Bank RCSB

More...
RCSB PDBi

Protein Data Bank Japan

More...
PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
1G26NMR-A281-311[»]
2JYENMR-A281-337[»]
2JYTNMR-A364-417[»]
2JYUNMR-A364-417[»]
2JYVNMR-A123-179[»]
6NUGNMR-A284-307[»]
SMRiP28799
ModBaseiSearch...
PDBe-KBiSearch...

Protein-protein interaction databases

BioGRIDi109153, 170 interactors
CORUMiP28799
DIPiDIP-41742N
IntActiP28799, 421 interactors
MINTiP28799
STRINGi9606.ENSP00000053867

PTM databases

GlyConnecti1290, 5 N-Linked glycans (1 site), 3 O-Linked glycans (1 site)
GlyGeniP28799, 7 sites
iPTMnetiP28799
MetOSiteiP28799
PhosphoSitePlusiP28799

Genetic variation databases

BioMutaiGRN
DMDMi77416865

Proteomic databases

EPDiP28799
jPOSTiP28799
MassIVEiP28799
MaxQBiP28799
PaxDbiP28799
PeptideAtlasiP28799
PRIDEiP28799
ProteomicsDBi54499 [P28799-1]
54500 [P28799-2]
81234
TopDownProteomicsiP28799-2 [P28799-2]
P28799-3 [P28799-3]

Protocols and materials databases

Antibodypedia a portal for validated antibodies

More...
Antibodypediai
1406, 660 antibodies

The DNASU plasmid repository

More...
DNASUi
2896

Genome annotation databases

EnsembliENST00000053867; ENSP00000053867; ENSG00000030582 [P28799-1]
ENST00000639447; ENSP00000492014; ENSG00000030582 [P28799-2]
GeneIDi2896
KEGGihsa:2896
UCSCiuc002igp.2, human [P28799-1]

Organism-specific databases

Comparative Toxicogenomics Database

More...
CTDi
2896
DisGeNETi2896

GeneCards: human genes, protein and diseases

More...
GeneCardsi
GRN
GeneReviewsiGRN
HGNCiHGNC:4601, GRN
HPAiENSG00000030582, Low tissue specificity
MalaCardsiGRN
MIMi138945, gene
607485, phenotype
614706, phenotype
neXtProtiNX_P28799
OpenTargetsiENSG00000030582
Orphaneti275864, Behavioral variant of frontotemporal dementia
314629, CLN11 disease
100070, Progressive non-fluent aphasia
100069, Semantic dementia
PharmGKBiPA28998
VEuPathDBiHostDB:ENSG00000030582.16

GenAtlas: human gene database

More...
GenAtlasi
Search...

Phylogenomic databases

eggNOGiKOG4296, Eukaryota
GeneTreeiENSGT00470000042293
HOGENOMiCLU_026274_0_0_1
InParanoidiP28799
OMAiPQNTVCN
OrthoDBi162721at2759
PhylomeDBiP28799
TreeFamiTF319678

Enzyme and pathway databases

PathwayCommonsiP28799
ReactomeiR-HSA-6798695, Neutrophil degranulation
SIGNORiP28799

Miscellaneous databases

BioGRID ORCS database of CRISPR phenotype screens

More...
BioGRID-ORCSi
2896, 11 hits in 995 CRISPR screens

ChiTaRS: a database of human, mouse and fruit fly chimeric transcripts and RNA-sequencing data

More...
ChiTaRSi
GRN, human
EvolutionaryTraceiP28799

The Gene Wiki collection of pages on human genes and proteins

More...
GeneWikii
Granulin

Database of phenotypes from RNA interference screens in Drosophila and Homo sapiens

More...
GenomeRNAii
2896
PharosiP28799, Tbio

Protein Ontology

More...
PROi
PR:P28799
RNActiP28799, protein

The Stanford Online Universal Resource for Clones and ESTs

More...
SOURCEi
Search...

Gene expression databases

BgeeiENSG00000030582, Expressed in esophagus mucosa and 246 other tissues
ExpressionAtlasiP28799, baseline and differential
GenevisibleiP28799, HS

Family and domain databases

Gene3Di3.10.20.500, 7 hits
InterProiView protein in InterPro
IPR000118, Granulin
IPR039036, Granulin_fam
IPR037277, Granulin_sf
PANTHERiPTHR12274, PTHR12274, 1 hit
PfamiView protein in Pfam
PF00396, Granulin, 7 hits
SMARTiView protein in SMART
SM00277, GRAN, 7 hits
PROSITEiView protein in PROSITE
PS00799, GRANULINS, 7 hits

MobiDB: a database of protein disorder and mobility annotations

More...
MobiDBi
Search...

<p>This section provides general information on the entry.<p><a href='/help/entry_information_section' target='_top'>More...</a></p>Entry informationi

<p>This subsection of the 'Entry information' section provides a mnemonic identifier for a UniProtKB entry, but it is not a stable identifier. Each reviewed entry is assigned a unique entry name upon integration into UniProtKB/Swiss-Prot.<p><a href='/help/entry_name' target='_top'>More...</a></p>Entry nameiGRN_HUMAN
<p>This subsection of the 'Entry information' section provides one or more accession number(s). These are stable identifiers and should be used to cite UniProtKB entries. Upon integration into UniProtKB, each entry is assigned a unique accession number, which is called 'Primary (citable) accession number'.<p><a href='/help/accession_numbers' target='_top'>More...</a></p>AccessioniPrimary (citable) accession number: P28799
Secondary accession number(s): D3DX55
, P23781, P23782, P23783, P23784, Q53HQ8, Q53Y88, Q540U8, Q9BWE7, Q9H8S1, Q9UCH0
<p>This subsection of the 'Entry information' section shows the date of integration of the entry into UniProtKB, the date of the last sequence update and the date of the last annotation modification ('Last modified'). The version number for both the entry and the <a href="http://www.uniprot.org/help/canonical%5Fand%5Fisoforms">canonical sequence</a> are also displayed.<p><a href='/help/entry_history' target='_top'>More...</a></p>Entry historyiIntegrated into UniProtKB/Swiss-Prot: December 1, 1992
Last sequence update: October 11, 2005
Last modified: June 2, 2021
This is version 223 of the entry and version 2 of the sequence. See complete history.
<p>This subsection of the 'Entry information' section indicates whether the entry has been manually annotated and reviewed by UniProtKB curators or not, in other words, if the entry belongs to the Swiss-Prot section of UniProtKB (<strong>reviewed</strong>) or to the computer-annotated TrEMBL section (<strong>unreviewed</strong>).<p><a href='/help/entry_status' target='_top'>More...</a></p>Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program
DisclaimerAny medical or genetic information present in this entry is provided for research, educational and informational purposes only. It is not in any way intended to be used as a substitute for professional medical advice, diagnosis, treatment or care.

<p>This section contains any relevant information that doesn't fit in any other defined sections<p><a href='/help/miscellaneous_section' target='_top'>More...</a></p>Miscellaneousi

Keywords - Technical termi

3D-structure, Direct protein sequencing, Reference proteome

Documents

  1. Human chromosome 17
    Human chromosome 17: entries, gene names and cross-references to MIM
  2. Human entries with genetic variants
    List of human entries with genetic variants
  3. Human variants curated from literature reports
    Index of human variants curated from literature reports
  4. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
  5. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
  6. SIMILARITY comments
    Index of protein domains and families
UniProt is an ELIXIR core data resource
Main funding by: National Institutes of Health

We'd like to inform you that we have updated our Privacy Notice to comply with Europe’s new General Data Protection Regulation (GDPR) that applies since 25 May 2018.

Do not show this banner again