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NPY1R (neuropeptide Y receptor Y1)

Written2010-06Massimiliano Ruscica, Elena Dozio, Luca Passafaro, Paolo Magni
Dipartimento di Endocrinologia, Fisiopatologia e Biologia Applicata, Universita degli Studi di Milano, Italy (MR, LP, PM); Dipartimento di Morfologia Umana e Scienze Biomediche Citta Studi, Universita degli Studi di Milano, Italy (ED)

(Note : for Links provided by Atlas : click)

Identity

Alias_namesNPYR
Other alias
HGNC (Hugo) NPY1R
LocusID (NCBI) 4886
Atlas_Id 44260
Location 4q32.2  [Link to chromosome band 4q32]
Location_base_pair Starts at 163323965 and ends at 163332795 bp from pter ( according to hg19-Feb_2009)  [Mapping NPY1R.png]

DNA/RNA

Note History: The human NPY1R cDNA was cloned from a human brain cDNA library. The NPY1R was the first to be characterized, when the expression pattern of an orphan receptor was recognized to overlap with the distribution of NPY in brain. NPY receptors belong to the large superfamily of G-protein-coupled receptors. Many of these receptor genes lack introns, supporting the proposition that they were created via RNA-mediated transpositional events. Differently from the other NPY receptor isoforms NPY1R is the only one containing a single 97-base pairs (bp) intron in the coding region following the fifth transmembrane domain.
 
  Exon/intron structure and splice sites in the 5'UTR of the human NPY1R gene.
Description A 14-kilobase pair (kb) region of genomic DNA encoding the human neuropeptide Y Y1-receptor gene including 3'- and 5'- flanking sequences is localized to chromosome 4. It encompasses 8632 bp of DNA (4q32.2) between 164245117 and 164253748 bp. The overall sequence of the gene consists of approximately 10 kb. The genomic structure presents a 6-kb intron situated approximately 150 bp upstream of the start codon within the 5'-untranslated region (5'UTR), as well as a small intron within the coding region.
The human NPY1R gene is divided into three exons: exon 1 (115 bp), exon 2 (850 bp), and exon 3 (1749 bp). In particular, the NPY1R gene contains three alternative exon 1 sequences (80, 110, and 106 bp) located 6.4, 18.4, and 23.9 kb upstream of exon 2. Exon 1A is located 6.4 kb upstream of exon 2; exon 1B was found a further 12 kb upstream exon 1A, and exon 1C another 5.5 kb upstream of exon 1B. These alternative 5' exons allow the regulation of tissue-specific expression of the receptor. The first 57 nucleotides of the 5'UTR of the human NPY1R mRNA are separated by a 6-kb intron from the second exon. The second intron 97 bp, containing an in-frame stop codon, is located at nucleotide 908 in the protein coding region after the fifth transmembrane domain between exon 2 and 3. Moreover, as shown by Nakamura, mouse NPY1R gene contains an alternate exon 4 located over 15 kb downstream of exon 3.
Transcription The human cDNA encodes a protein of 384 amino-acids (aa) in lenght that is preceded by approximately 200 bp of 5'UTR sequence.

Protein

Note The NPY Y1 receptor subtype (Y1R) was the first to be cloned in the rat, and subsequently in human and mouse. This receptor is as conserved as its ligand (NPY) throughout evolution and mammalian and non-mammalian species. The complete NPY1-36 molecule is necessary for NPY to bind to Y1R. Any proteolytic process leading to alterations in the NH2-terminal domain essentially abolishes the ability of NPY to bind to Y1R. Therefore, NH2-terminally truncated NPY fragments such as NPY2-36, or NPY3-36 have little or no affinity for the Y1R. Modification of COOH-terminal residues does not affect agonist binding. Thus, it has been established that the NH2 terminus is essential for NPY to activate Y1R. The pharmacological profile of the Y1R is characterized by high affinity for NPY, PYY and the corresponding analogs containing Pro34 and low affinity for the N-terminally truncated analogs and for PP.
 
  Figure A. Y1R affinity for various PP-family hormones and their C-terminal sequences.
Figure B. Homo sapiens Neuropeptide Y receptor type 1 (384 aa).
Description Y1R has seven putative transmembrane domains associated with G-protein (GPCR). In the N-terminal portion Y1R presents potential sites of glycosylation and in the second extracellular loop, four extracellular cysteines in position 33, 113, 198 and 296 which may form two disulfide bridges (Cys 33 and 296; Cys 133 and 198). Phosphorylation sites are present in the intracellular domain (cysteine in the C-terminal portion at position 338). These cysteines may also explain the capability of palmitate residues to bind to the receptor. As observed for many GPCR, Y1R is internalized together with its ligand into endosomes and recycled to the cell surface within 60 minutes upon agonist stimulation. Moreover, Y1R is able to form homodimers.
Expression The expression of the human NPY Y1 receptor has been studied extensively by using immunohistochemical methods, in situ hybridization experiments and reverse-transcription polymerase chain reaction (RT-PCR; mRNA detection). The human NPY1R is expressed in both central nervous system (i.e., cerebral cortex, thalamus and amigdala) and periphery (i.e., heart, kidneys, gastrointestinal tract, as well as blood vessels).

Nervous system
The NPY Y1R is widely distributed in the central nervous system. A study conducted on four normal human brains revealed that high levels of Y1R receptor mRNA were expressed in cortical areas and in the claustrum, while moderate levels were present in the nucleus accumbens, caudate nucleus, putamen, amygdaloid nuclei and arcuate and paraventricular nuclei of the hypothalamus. Moreover, a study conducted on prefrontal cortex of subjects affected by bipolar disorder, major depression, or schizophrenia revealed a progressive age-related decline in the expression of Y1R mRNA associated with a lack of coexpression with NPY neurons. Interestingly, there was no significant effect of suicide as a cause of death on Y1R mRNA expression levels. In fact, subjects with suicide as a cause of death tended to have higher Y1R mRNA expression levels, but these individuals were among the youngest ones (45 years old) in the population studied.

Periphery
Peripherally, Y1Rs are expressed mainly in arteries and veins, where they are associated with vasoconstriction and potentiation of other vasoconstrictors of neurogenic origin. Although limited, there is evidence of prejunctional Y1R inhibition of neurotransmitter release. Nonetheless, NPY Y1R is primarily located postjunctionally on vascular smooth muscle cells.
1) Colon
In vitro receptor autoradiography ([125I]PYY) performed on normal human colonic tissue obtained from nine patients showed that Y1R is distributed only in vessels. No measurable levels of subtype Y1 was identified in smooth muscle, mucosa, muscularis mucosae, as well as in lymphoid follicles, myoenteric and submucosal plexus.
2) Heart
A study conducted on 20-week old fetal human hearts showed that Y1R is present on right ventricular endocardial endothelial cells. In particular, it is highly expressed at the level of the nucleus specifically at the perinucleoplasm and nuclear membrane levels, while lower levels were detected in the cytoplasm and the plasma membrane.
3) Dental pulp
NPY Y1R proteins were present in solubilized membrane preparations of both healthy and inflamed human gingival tissue by Western blotting. Major immunoreactive bands were detected at approximately 55 kDa due to a glycosylated form of the native receptor protein. By using the SwissProt glycosylation prediction packages NetNGlyc and NetOGly, authors confirmed that the human Y1R has potential N- and O-glycosylation sites. The expression of Y1R protein in both healthy and inflamed gingival tissue suggests that NPY could act via the Y1R to exert its tonic effects. Moreover, Y1R was expressed in human dental pulp with evidence of increased expression in carious compared with noncarious teeth. Y1R were localized to nerve fibres and inflammatory cells in the dental pulp of carious teeth.
4) Achilles tendons
Y1R is expressed in the tenocytes in the Achilles tendon. Specifically, Y1R is present within the smooth muscle of the blood vessel walls, but not in the endothelial layer of calcaneal tendons.
5) Skin
In human tissues, RT-PCR and immunocytochemistry studies suggested that Y1R is the primary receptor in human cutaneous circulation, supporting the findings that local non-noradrenergic mechanisms are entirely Y1R-based. Skin blood flow in humans is controlled through two branches of the sympathetic nervous system: a vasoconstrictor system and an active vasodilator system of uncertain neurotransmitter. In this context, NPY showed a vasoconstrictor effect in human subcutaneous arteries that had been dissected out of the abdominal regions from patients who underwent nonvascular disease surgeries (e.g., hernia). NPY decreased cutaneous blood flow via Y1R, with evidence for the additional involvement of postjunctional Y2R. This ability of NPY and Y1R to affect skin vascular conductance varies in accordance with relative innervations at specific sites.

Localisation NPY Y1R is a seven transmembrane receptor which has all the characteristics of the GPCR family including potential glycosylation sites in the N-terminal portion and in the second extra-cellular loop.
Function NPY has been demonstrated to be involved in mitogenic pathways and stimulate cell proliferation via the Y1R. The activation of Y1R is generally associated with reduction of cAMP accumulation, increase of intracellular free calcium concentration ([Ca2+]i), and modulation of the MAPK pathway via several signaling molecules, including the protein kinase C (PKC).
Y1R has been involved in several NPY-induced responses, such as activation of neuroendocrine axes, vasoconstriction, anxiolysis, as well as the stimulation of food intake. Moreover, Y1R mediates emotional behavior, stress response, and ethanol consumption.
The prototype of NPY Y1R-mediated responses is vasoconstriction. Specifically, the physiological role of the Y1R subtype was demonstrated in mice lacking Y1R expression, which show no blood pressure response to NPY, but a normal response to norepinephrine. Y1R knockout mice have normal blood pressure, suggesting that the Y1R does not play a crucial role in maintaining blood pressure homeostasis in unstimulated conditions. However, Y1R has been also involved in other NPY-induced responses, such as stimulation of food intake and activation of neuroendocrine axes. In particular, Y1R and Y5R, both expressed in hypothalamic regions involved in the control of feeding, represent the most likely candidates for mediating the appetite stimulatory capacity of NPY. Mice lacking Y1R showed an increased body weight due to a low-energy expenditure rather than high-energy intake. In fact, these mice had a decreased metabolic rate secondary to decreased locomotor activity and movement associated thermogenesis.
Homology The human Y1R subtype shares closest aa identity with the Y4R subtype (42%) and the non-active, human form of the y6 subtype (51%).

Mutations

Note In 2004 Ramanathan described a case of autism in which a 19 megabase on chromosome 4q, spanning 4q32 to 4q34, was detected. Being involved in the deletion, those genes which are abundantly expressed in the brain, Y1R and Y5R resulted implicated. In this context, being the neuroproliferative effect of NPY in the hippocampus mediated through the neuropeptide Y Y1R, the authors postulate that the effect of NPY on learning and memory may be mediated through NPY neurogenesis.
Okahisa et al. described that genetic variants of rs7687423 of the NPY1R gene may alter the subjective effects of methamphetamine and result in susceptibility to dependence. Because NPY1R mRNA changes were observed in peripheral tissues and the brain in schizophrenia patients, these findings may also indicate that the NPY1R gene is involved in vulnerability to methamphetamine-induced psychosis because almost all of the analyzed subjects with methamphetamine dependence had comorbid methamphetamine psychosis.

Implicated in

Note
  
Entity Various tumors
Note NPY receptors are mainly expressed in specific endocrine tumors and epithelial malignancies as well as in embryonal tumors. In endocrine tumors, NPY receptors are present in steroid hormone producing tumors, namely adrenal cortical adenomas, carcinomas, ovarian granulosa cell tumors, Sertoli-Leydig cell tumors, and in catecholamine producing tumors, i.e. pheochromocytomas and paragangliomas. Based on pharmacological displacement experiments, in addition to tumor cells, intra- and peritumoral blood vessels express Y1Rs. The Y1R-expressing blood vessels are mainly small and medium-sized arteries.
  
  
Entity Prostate cancer
Note Prostate cancer represents one of the most common malignant diseases among men in the Western world. It is initially androgen dependent and it may later progress to the androgen-independent stage, which is associated with a lack of efficacy of the available hormonal therapy. This tumoral progression appears to be promoted at least in part by several growth factors and neurohormones. Within this context, we showed that Y1R protein is expressed in three human prostate cancer cell lines (LNCaP -androgen dependent-, DU145 and PC3 -androgen independent-) and that NPY treatment reduced the proliferation of LNCaP and DU145 cells and increased that of PC3 cells. Interestingly, the Y1R antagonist BIBP3226 abolished such effects, suggesting a mandatory role of Y1-R in this process. Moreover, these effects are associated with a clone-specific pattern of intracellular signaling activation, including a peculiar time-course of MAPK/ERK1/ERK2 phosphorylation (long-lasting in DU145 and transient in PC3 cells).
  
  
Entity Breast cancer
Note Breast cancer accounts for almost 1/3 of all incident cases of cancer in women. Interestingly, the expression of NPY-Rs has been found in 85% of primary breast cancer in a series of 95 cases, and in 100% of lymph node metastases of receptor-positive primaries, where Y1R expression predominated and was often present in high density and great homogeneity. In normal breast tissue, however, Y1R was only found in a minority of the cases and concomitantly with Y2R, which seemed to be predominant in non-neoplastic breast. The neoplastic condition of breast tissue may thus induce a switch of expression from Y2R to Y1R. Moreover, a functional interplay between estrogen and Y1R has been shown in a human breast cancer cell line responsive to this steroid, where estrogen was found to increase Y1R expression, which in turn negatively regulated estrogen-stimulated cell proliferation.
  
  
Entity Pheochromocytoma and paraganglioma
Note The frequency of NPY receptors (NPYRs) expression in pheochromocytomas and paragangliomas was found to be 35% and 61%, respectively. Both Y1R and Y2R are expressed, with a higher density of Y2R in paragangliomas than in pheochromocytomas, whereas the density of Y1R is comparably low in both tumor categories. NPYRs, mainly Y1R and Y5R, are also expressed in the Ewing's sarcoma family of tumors, other related neural crest-derived tumors, where activation of these receptors has been reported to regulate cell proliferation, as shown in the SK-N-MC cell line, an Ewing's sarcoma family of tumors expressing Y1R, where NPY has been shown to inhibit cell growth.
  

Bibliography

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Citation

This paper should be referenced as such :
Ruscica, M ; Dozio, E ; Passafaro, L ; Magni, P
NPY1R (neuropeptide Y receptor Y1)
Atlas Genet Cytogenet Oncol Haematol. 2011;15(3):283-287.
Free journal version : [ pdf ]   [ DOI ]
On line version : http://AtlasGeneticsOncology.org/Genes/NPY1RID44260ch4q32.html


External links

Nomenclature
HGNC (Hugo)NPY1R   7956
Cards
AtlasNPY1RID44260ch4q32
Entrez_Gene (NCBI)NPY1R  4886  neuropeptide Y receptor Y1
AliasesNPY1-R; NPYR
GeneCards (Weizmann)NPY1R
Ensembl hg19 (Hinxton)ENSG00000164128 [Gene_View]
Ensembl hg38 (Hinxton)ENSG00000164128 [Gene_View]  chr4:163323965-163332795 [Contig_View]  NPY1R [Vega]
ICGC DataPortalENSG00000164128
TCGA cBioPortalNPY1R
AceView (NCBI)NPY1R
Genatlas (Paris)NPY1R
WikiGenes4886
SOURCE (Princeton)NPY1R
Genetics Home Reference (NIH)NPY1R
Genomic and cartography
GoldenPath hg38 (UCSC)NPY1R  -     chr4:163323965-163332795 -  4q32.2   [Description]    (hg38-Dec_2013)
GoldenPath hg19 (UCSC)NPY1R  -     4q32.2   [Description]    (hg19-Feb_2009)
EnsemblNPY1R - 4q32.2 [CytoView hg19]  NPY1R - 4q32.2 [CytoView hg38]
Mapping of homologs : NCBINPY1R [Mapview hg19]  NPY1R [Mapview hg38]
OMIM162641   
Gene and transcription
Genbank (Entrez)A26481 AB209237 AK296624 AK298282 AK312578
RefSeq transcript (Entrez)NM_000909
RefSeq genomic (Entrez)
Consensus coding sequences : CCDS (NCBI)NPY1R
Cluster EST : UnigeneHs.733329 [ NCBI ]
CGAP (NCI)Hs.733329
Alternative Splicing GalleryENSG00000164128
Gene ExpressionNPY1R [ NCBI-GEO ]   NPY1R [ EBI - ARRAY_EXPRESS ]   NPY1R [ SEEK ]   NPY1R [ MEM ]
Gene Expression Viewer (FireBrowse)NPY1R [ Firebrowse - Broad ]
SOURCE (Princeton)Expression in : [Datasets]   [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
GenevisibleExpression in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
BioGPS (Tissue expression)4886
GTEX Portal (Tissue expression)NPY1R
Protein : pattern, domain, 3D structure
UniProt/SwissProtP25929   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
NextProtP25929  [Sequence]  [Exons]  [Medical]  [Publications]
With graphics : InterProP25929
Splice isoforms : SwissVarP25929
PhosPhoSitePlusP25929
Domaine pattern : Prosite (Expaxy)G_PROTEIN_RECEP_F1_1 (PS00237)    G_PROTEIN_RECEP_F1_2 (PS50262)   
Domains : Interpro (EBI)GPCR_Rhodpsn    GPCR_Rhodpsn_7TM    NPY1_rcpt    NPY_rcpt   
Domain families : Pfam (Sanger)7tm_1 (PF00001)   
Domain families : Pfam (NCBI)pfam00001   
Conserved Domain (NCBI)NPY1R
DMDM Disease mutations4886
Blocks (Seattle)NPY1R
PDB (SRS)2F1U    2IIL   
PDB (PDBSum)2F1U    2IIL   
PDB (IMB)2F1U    2IIL   
PDB (RSDB)2F1U    2IIL   
Structural Biology KnowledgeBase2F1U    2IIL   
SCOP (Structural Classification of Proteins)2F1U    2IIL   
CATH (Classification of proteins structures)2F1U    2IIL   
SuperfamilyP25929
Human Protein AtlasENSG00000164128
Peptide AtlasP25929
HPRD01215
IPIIPI00018252   IPI00963848   IPI00556057   IPI00965803   IPI00964874   IPI00965299   IPI00965510   
Protein Interaction databases
DIP (DOE-UCLA)P25929
IntAct (EBI)P25929
FunCoupENSG00000164128
BioGRIDNPY1R
STRING (EMBL)NPY1R
ZODIACNPY1R
Ontologies - Pathways
QuickGOP25929
Ontology : AmiGOpeptide YY receptor activity  pancreatic polypeptide receptor activity  outflow tract morphogenesis  neuropeptide Y receptor activity  protein binding  plasma membrane  integral component of plasma membrane  glucose metabolic process  G-protein coupled receptor signaling pathway, coupled to cyclic nucleotide second messenger  adenylate cyclase-inhibiting G-protein coupled receptor signaling pathway  neuropeptide signaling pathway  chemical synaptic transmission  locomotory behavior  feeding behavior  blood circulation  regulation of blood pressure  sensory perception of pain  regulation of multicellular organism growth  
Ontology : EGO-EBIpeptide YY receptor activity  pancreatic polypeptide receptor activity  outflow tract morphogenesis  neuropeptide Y receptor activity  protein binding  plasma membrane  integral component of plasma membrane  glucose metabolic process  G-protein coupled receptor signaling pathway, coupled to cyclic nucleotide second messenger  adenylate cyclase-inhibiting G-protein coupled receptor signaling pathway  neuropeptide signaling pathway  chemical synaptic transmission  locomotory behavior  feeding behavior  blood circulation  regulation of blood pressure  sensory perception of pain  regulation of multicellular organism growth  
Pathways : KEGGNeuroactive ligand-receptor interaction   
REACTOMEP25929 [protein]
REACTOME PathwaysR-HSA-418594 [pathway]   
NDEx NetworkNPY1R
Atlas of Cancer Signalling NetworkNPY1R
Wikipedia pathwaysNPY1R
Orthology - Evolution
OrthoDB4886
GeneTree (enSembl)ENSG00000164128
Phylogenetic Trees/Animal Genes : TreeFamNPY1R
HOVERGENP25929
HOGENOMP25929
Homologs : HomoloGeneNPY1R
Homology/Alignments : Family Browser (UCSC)NPY1R
Gene fusions - Rearrangements
Polymorphisms : SNP and Copy number variants
NCBI Variation ViewerNPY1R [hg38]
dbSNP Single Nucleotide Polymorphism (NCBI)NPY1R
dbVarNPY1R
ClinVarNPY1R
1000_GenomesNPY1R 
Exome Variant ServerNPY1R
ExAC (Exome Aggregation Consortium)NPY1R (select the gene name)
Genetic variants : HAPMAP4886
Genomic Variants (DGV)NPY1R [DGVbeta]
DECIPHERNPY1R [patients]   [syndromes]   [variants]   [genes]  
CONAN: Copy Number AnalysisNPY1R 
Mutations
ICGC Data PortalNPY1R 
TCGA Data PortalNPY1R 
Broad Tumor PortalNPY1R
OASIS PortalNPY1R [ Somatic mutations - Copy number]
Somatic Mutations in Cancer : COSMICNPY1R  [overview]  [genome browser]  [tissue]  [distribution]  
Mutations and Diseases : HGMDNPY1R
LOVD (Leiden Open Variation Database)Whole genome datasets
LOVD (Leiden Open Variation Database)LOVD - Leiden Open Variation Database
LOVD (Leiden Open Variation Database)LOVD 3.0 shared installation
BioMutasearch NPY1R
DgiDB (Drug Gene Interaction Database)NPY1R
DoCM (Curated mutations)NPY1R (select the gene name)
CIViC (Clinical Interpretations of Variants in Cancer)NPY1R (select a term)
intoGenNPY1R
NCG5 (London)NPY1R
Cancer3DNPY1R(select the gene name)
Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
Diseases
OMIM162641   
Orphanet
MedgenNPY1R
Genetic Testing Registry NPY1R
NextProtP25929 [Medical]
TSGene4886
GENETestsNPY1R
Target ValidationNPY1R
Huge Navigator NPY1R [HugePedia]
snp3D : Map Gene to Disease4886
BioCentury BCIQNPY1R
ClinGenNPY1R
Clinical trials, drugs, therapy
Chemical/Protein Interactions : CTD4886
Chemical/Pharm GKB GenePA258
Clinical trialNPY1R
Miscellaneous
canSAR (ICR)NPY1R (select the gene name)
Probes
Litterature
PubMed73 Pubmed reference(s) in Entrez
GeneRIFsGene References Into Functions (Entrez)
CoreMineNPY1R
EVEXNPY1R
GoPubMedNPY1R
iHOPNPY1R
REVIEW articlesautomatic search in PubMed
Last year publicationsautomatic search in PubMed

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