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Aliases for LRP4 Gene

Aliases for LRP4 Gene

  • LDL Receptor Related Protein 4 2 3 5
  • Multiple Epidermal Growth Factor-Like Domains 7 3 4
  • MEGF7 3 4
  • LRP-4 3 4
  • LRP10 3 4
  • Low Density Lipoprotein Receptor-Related Protein 4 2
  • Low-Density Lipoprotein Receptor-Related Protein 4 3
  • KIAA0816 4
  • CMS17 3
  • SOST2 3
  • CLSS 3

External Ids for LRP4 Gene

Previous GeneCards Identifiers for LRP4 Gene

  • GC11M048816
  • GC11M047756
  • GC11M046910
  • GC11M046842
  • GC11M046834
  • GC11M046584
  • GC11M046919
  • GC11M048070
  • GC11M048796
  • GC11M049899
  • GC11M047088
  • GC11M047932
  • GC11M050786
  • GC11M051581
  • GC11M052243
  • GC11M052903
  • GC11M053571

Summaries for LRP4 Gene

Entrez Gene Summary for LRP4 Gene

  • This gene encodes a member of the low-density lipoprotein receptor-related protein family. The encoded protein may be a regulator of Wnt signaling. Mutations in this gene are associated with Cenani-Lenz syndrome. [provided by RefSeq, May 2010]

GeneCards Summary for LRP4 Gene

LRP4 (LDL Receptor Related Protein 4) is a Protein Coding gene. Diseases associated with LRP4 include Cenani-Lenz Syndactyly Syndrome and Myasthenic Syndrome, Congenital, 17. Among its related pathways are ECM proteoglycans and Degradation of the extracellular matrix. Gene Ontology (GO) annotations related to this gene include calcium ion binding and scaffold protein binding. An important paralog of this gene is LRP2.

UniProtKB/Swiss-Prot for LRP4 Gene

  • Mediates SOST-dependent inhibition of bone formation. Functions as a specific facilitator of SOST-mediated inhibition of Wnt signaling. Plays a key role in the formation and the maintenance of the neuromuscular junction (NMJ), the synapse between motor neuron and skeletal muscle. Directly binds AGRIN and recruits it to the MUSK signaling complex. Mediates the AGRIN-induced phosphorylation of MUSK, the kinase of the complex. The activation of MUSK in myotubes induces the formation of NMJ by regulating different processes including the transcription of specific genes and the clustering of AChR in the postsynaptic membrane. Alternatively, may be involved in the negative regulation of the canonical Wnt signaling pathway, being able to antagonize the LRP6-mediated activation of this pathway. More generally, has been proposed to function as a cell surface endocytic receptor binding and internalizing extracellular ligands for degradation by lysosomes. May play an essential role in the process of digit differentiation (By similarity).

Gene Wiki entry for LRP4 Gene

Additional gene information for LRP4 Gene

No data available for CIViC summary , Tocris Summary , PharmGKB "VIP" Summary , fRNAdb sequence ontologies and piRNA Summary for LRP4 Gene

Genomics for LRP4 Gene

GeneHancer (GH) Regulatory Elements for LRP4 Gene

Promoters and enhancers for LRP4 Gene
GeneHancer (GH) Identifier GH Type GH
Score
GH Sources Gene Association Score Total Score TSS distance (kb) Number of Genes Away Size (kb) Transcription Factor
Binding Sites
Gene Targets
GH11I046916 Promoter/Enhancer 2.3 EPDnew FANTOM5 Ensembl ENCODE dbSUPER 562.4 -0.2 -232 5.5 HDGF PKNOX1 FOXA2 ARID4B SIN3A ZNF48 ZNF121 GLIS2 ZNF143 FOS LRP4 CHRM4 C11orf49
GH11I047213 Promoter/Enhancer 2.3 EPDnew Ensembl ENCODE dbSUPER 16.8 -296.7 -296698 3.7 HDGF PKNOX1 CLOCK FOXA2 SMAD1 MLX ARID4B SIN3A DMAP1 IRF4 DDB2 LRP4 PACSIN3 NUP160 FNBP4 ENSG00000270060 ZNF408 C11orf49 ATG13 ENSG00000256897
GH11I047182 Promoter/Enhancer 2.7 EPDnew FANTOM5 Ensembl ENCODE dbSUPER 13.5 -267.7 -267726 8.2 PKNOX1 CLOCK FOXA2 MLX ZFP64 ARID4B SIN3A DMAP1 ZNF2 ZBTB7B PACSIN3 ENSG00000271350 LRP4 C11orf49 MADD NR1H3 MIR6745
GH11I047175 Promoter/Enhancer 2.3 EPDnew Ensembl ENCODE dbSUPER 15.4 -259.1 -259098 4.7 HDGF HNRNPUL1 PKNOX1 CLOCK FOXA2 ARNT ARID4B SIN3A DMAP1 ZNF2 ARFGAP2 MIR6745 NUP160 FNBP4 ENSG00000270060 ENSG00000200090 DDB2 RPS10P19 ZNF408 LRP4
GH11I047142 Enhancer 1.1 Ensembl ENCODE dbSUPER 29.9 -224.6 -224625 1.6 CLOCK NFIB MAX SIN3A ZBTB40 ZNF664 ZNF335 CTBP1 GATA3 POLR2A LRP4 C11orf49 F2 ENSG00000255520 GC11P047160
- Elite GeneHancer and/or Elite GeneHancer-gene association Download GeneHancer data dump

GeneHancers around LRP4 on UCSC Golden Path with GeneCards custom track

Top Transcription factor binding sites by QIAGEN in the LRP4 gene promoter:

Genomic Locations for LRP4 Gene

Genomic Locations for LRP4 Gene
chr11:46,856,717-46,918,642
(GRCh38/hg38)
Size:
61,926 bases
Orientation:
Minus strand
chr11:46,878,268-46,940,193
(GRCh37/hg19)

Genomic View for LRP4 Gene

Genes around LRP4 on UCSC Golden Path with GeneCards custom track

Cytogenetic band:
LRP4 Gene in genomic location: bands according to Ensembl, locations according to GeneLoc (and/or Entrez Gene and/or Ensembl if different)
Genomic Location for LRP4 Gene
GeneLoc Logo Genomic Neighborhood Exon StructureGene Density

RefSeq DNA sequence for LRP4 Gene

Proteins for LRP4 Gene

  • Protein details for LRP4 Gene (UniProtKB/Swiss-Prot)

    Protein Symbol:
    O75096-LRP4_HUMAN
    Recommended name:
    Low-density lipoprotein receptor-related protein 4
    Protein Accession:
    O75096
    Secondary Accessions:
    • B2RN39
    • Q4AC85
    • Q5KTZ5

    Protein attributes for LRP4 Gene

    Size:
    1905 amino acids
    Molecular mass:
    212045 Da
    Quaternary structure:
    • Homooligomer. Interacts with MUSK; the heterodimer forms an AGRIN receptor complex that binds AGRIN resulting in activation of MUSK (By similarity). Interacts (via the extracellular domain) with SOST; the interaction facilitates the inhibition of Wnt signaling (PubMed:21471202). Interacts with MESD; the interaction promotes glycosylation of LRP4 and its cell-surface expression (By similarity).
    SequenceCaution:
    • Sequence=BAE19679.1; Type=Erroneous initiation; Note=Translation N-terminally shortened.; Evidence={ECO:0000305};

neXtProt entry for LRP4 Gene

Post-translational modifications for LRP4 Gene

  • Ubiquitination at posLast=17531753
  • Glycosylation at posLast=264264, posLast=498498, posLast=719719, isoforms=901, posLast=10771077, isoforms=1415, isoforms=1467, and posLast=16881688

Other Protein References for LRP4 Gene

No data available for DME Specific Peptides for LRP4 Gene

Domains & Families for LRP4 Gene

Gene Families for LRP4 Gene

HGNC:
Human Protein Atlas (HPA):
  • Disease related genes
  • Predicted membrane proteins
  • Predicted secreted proteins

Suggested Antigen Peptide Sequences for LRP4 Gene

Graphical View of Domain Structure for InterPro Entry

O75096

UniProtKB/Swiss-Prot:

LRP4_HUMAN :
  • Belongs to the LDLR family.
Family:
  • Belongs to the LDLR family.
genes like me logo Genes that share domains with LRP4: view

Function for LRP4 Gene

Molecular function for LRP4 Gene

GENATLAS Biochemistry:
low density lipoprotein receptor-related protein 4,abundant in heart
UniProtKB/Swiss-Prot Function:
Mediates SOST-dependent inhibition of bone formation. Functions as a specific facilitator of SOST-mediated inhibition of Wnt signaling. Plays a key role in the formation and the maintenance of the neuromuscular junction (NMJ), the synapse between motor neuron and skeletal muscle. Directly binds AGRIN and recruits it to the MUSK signaling complex. Mediates the AGRIN-induced phosphorylation of MUSK, the kinase of the complex. The activation of MUSK in myotubes induces the formation of NMJ by regulating different processes including the transcription of specific genes and the clustering of AChR in the postsynaptic membrane. Alternatively, may be involved in the negative regulation of the canonical Wnt signaling pathway, being able to antagonize the LRP6-mediated activation of this pathway. More generally, has been proposed to function as a cell surface endocytic receptor binding and internalizing extracellular ligands for degradation by lysosomes. May play an essential role in the process of digit differentiation (By similarity).

Phenotypes From GWAS Catalog for LRP4 Gene

Gene Ontology (GO) - Molecular Function for LRP4 Gene

GO ID Qualified GO term Evidence PubMed IDs
GO:0005509 calcium ion binding IEA --
GO:0005515 protein binding IPI 21471202
GO:0017147 Wnt-protein binding IBA --
GO:0030971 receptor tyrosine kinase binding IEA,ISS --
GO:0034185 apolipoprotein binding IEA --
genes like me logo Genes that share ontologies with LRP4: view
genes like me logo Genes that share phenotypes with LRP4: view

Human Phenotype Ontology for LRP4 Gene

HPO Id HPO Name Alternative Ids Definition Synonyms

Animal Models for LRP4 Gene

MGI Knock Outs for LRP4:

miRNA for LRP4 Gene

No data available for Enzyme Numbers (IUBMB) , Transcription Factor Targets and HOMER Transcription for LRP4 Gene

Localization for LRP4 Gene

Subcellular locations from UniProtKB/Swiss-Prot for LRP4 Gene

Cell membrane; Single-pass type I membrane protein.

Subcellular locations from

COMPARTMENTS
Extracellular space Cytosol Plasma membrane Cytoskeleton Lysosome Endosome Peroxisome ER Golgi Apparatus Nucleus Mitochondrion 0 1 2 3 4 5 Confidence
COMPARTMENTS Subcellular localization image for LRP4 gene
Compartment Confidence
plasma membrane 5
extracellular 2
mitochondrion 2
nucleus 2
cytosol 2
peroxisome 1

Subcellular locations from the

Human Protein Atlas (HPA)
  • Mitochondria (1)
  • Nucleoli (1)
  • Nucleus (1)
See all subcellular structures

Gene Ontology (GO) - Cellular Components for LRP4 Gene

GO ID Qualified GO term Evidence PubMed IDs
GO:0005886 plasma membrane IBA,ISS --
GO:0009986 cell surface IEA,ISS --
GO:0014069 postsynaptic density ISS,IEA --
GO:0016020 membrane IEA --
GO:0016021 integral component of membrane IEA --
genes like me logo Genes that share ontologies with LRP4: view

Pathways & Interactions for LRP4 Gene

genes like me logo Genes that share pathways with LRP4: view

Pathways by source for LRP4 Gene

2 Reactome pathways for LRP4 Gene

Gene Ontology (GO) - Biological Process for LRP4 Gene

GO ID Qualified GO term Evidence PubMed IDs
GO:0001822 kidney development IDA 20381006
GO:0001932 regulation of protein phosphorylation IEA --
GO:0001942 hair follicle development IEA --
GO:0006897 endocytosis IEA --
GO:0007275 multicellular organism development IEA --
genes like me logo Genes that share ontologies with LRP4: view

No data available for SIGNOR curated interactions for LRP4 Gene

Drugs & Compounds for LRP4 Gene

No Compound Related Data Available

Transcripts for LRP4 Gene

Unigene Clusters for LRP4 Gene

Low density lipoprotein receptor-related protein 4:
Representative Sequences:

Alternative Splicing Database (ASD) splice patterns (SP) for LRP4 Gene

No ASD Table

Relevant External Links for LRP4 Gene

GeneLoc Exon Structure for
LRP4
ECgene alternative splicing isoforms for
LRP4

Expression for LRP4 Gene

mRNA expression in normal human tissues from GTEx, Illumina, BioGPS, and CGAP SAGE for LRP4 Gene

mRNA differential expression in normal tissues according to GTEx for LRP4 Gene

This gene is overexpressed in Brain - Caudate (basal ganglia) (x4.8).

Protein differential expression in normal tissues from HIPED for LRP4 Gene

This gene is overexpressed in Plasma (57.4) and Heart (9.2).

Integrated Proteomics: protein expression in normal tissues and cell lines from ProteomicsDB, PaxDb, and MOPED for LRP4 Gene



Protein tissue co-expression partners for LRP4 Gene

NURSA nuclear receptor signaling pathways regulating expression of LRP4 Gene:

LRP4

SOURCE GeneReport for Unigene cluster for LRP4 Gene:

Hs.4930

mRNA Expression by UniProt/SwissProt for LRP4 Gene:

O75096-LRP4_HUMAN
Tissue specificity: Expressed in bone; present in osteoblasts and osteocytes. No expression is observed in osteoclast. Expressed in several regions of the brain.

Evidence on tissue expression from TISSUES for LRP4 Gene

  • Nervous system(4.8)
  • Muscle(4.4)
  • Intestine(2.8)

Phenotype-based relationships between genes and organs from Gene ORGANizer for LRP4 Gene

Germ Layers:
  • ectoderm
  • endoderm
  • mesoderm
Systems:
  • cardiovascular
  • digestive
  • immune
  • integumentary
  • nervous
  • respiratory
  • skeletal muscle
  • skeleton
  • urinary
Organs:
Head and neck:
  • brain
  • cerebellum
  • cerebrospinal fluid
  • cheek
  • chin
  • cranial nerve
  • ear
  • eye
  • eyelid
  • face
  • forehead
  • head
  • jaw
  • larynx
  • mandible
  • maxilla
  • meninges
  • mouth
  • neck
  • nose
  • outer ear
  • pharynx
  • skull
  • tongue
  • tooth
  • vocal cord
Thorax:
  • chest wall
  • clavicle
  • diaphragm
  • esophagus
  • lung
  • rib
  • rib cage
  • scapula
  • sternum
Abdomen:
  • kidney
  • stomach
Pelvis:
  • pelvis
  • ureter
Limb:
  • ankle
  • arm
  • digit
  • elbow
  • femur
  • fibula
  • finger
  • foot
  • forearm
  • hand
  • hip
  • humerus
  • knee
  • lower limb
  • nail
  • radius
  • shin
  • shoulder
  • thigh
  • tibia
  • toe
  • ulna
  • upper limb
  • wrist
General:
  • blood
  • blood vessel
  • peripheral nerve
  • peripheral nervous system
  • skin
  • spinal column
  • spinal cord
  • vertebrae
  • white blood cell
genes like me logo Genes that share expression patterns with LRP4: view

No data available for mRNA expression in embryonic tissues and stem cells from LifeMap Discovery for LRP4 Gene

Orthologs for LRP4 Gene

This gene was present in the common ancestor of animals.

Orthologs for LRP4 Gene

Organism Taxonomy Gene Similarity Type Details
chimpanzee
(Pan troglodytes)
Mammalia LRP4 33 34
  • 99.67 (n)
platypus
(Ornithorhynchus anatinus)
Mammalia LRP4 34
  • 94 (a)
OneToOne
dog
(Canis familiaris)
Mammalia LRP4 33 34
  • 92.62 (n)
cow
(Bos Taurus)
Mammalia LRP4 33 34
  • 91.53 (n)
oppossum
(Monodelphis domestica)
Mammalia LRP4 34
  • 91 (a)
OneToOne
rat
(Rattus norvegicus)
Mammalia Lrp4 33
  • 90.87 (n)
mouse
(Mus musculus)
Mammalia Lrp4 33 16 34
  • 90.43 (n)
chicken
(Gallus gallus)
Aves LRP4 33 34
  • 80.13 (n)
lizard
(Anolis carolinensis)
Reptilia LRP4 34
  • 88 (a)
OneToOne
tropical clawed frog
(Silurana tropicalis)
Amphibia lrp4 33
  • 74.32 (n)
zebrafish
(Danio rerio)
Actinopterygii CABZ01083440.1 34
  • 81 (a)
OneToOne
lrp4 33
  • 69.39 (n)
African malaria mosquito
(Anopheles gambiae)
Insecta AgaP_AGAP000037 33
  • 55.74 (n)
fruit fly
(Drosophila melanogaster)
Insecta CG8909 35 33 34
  • 52.24 (n)
sea squirt
(Ciona savignyi)
Ascidiacea -- 34
  • 55 (a)
OneToOne
Species where no ortholog for LRP4 was found in the sources mined by GeneCards:
  • A. gosspyii yeast (Ashbya gossypii)
  • Actinobacteria (Mycobacterium tuberculosis)
  • African clawed frog (Xenopus laevis)
  • Alicante grape (Vitis vinifera)
  • alpha proteobacteria (Wolbachia pipientis)
  • amoeba (Dictyostelium discoideum)
  • Archea (Pyrococcus horikoshii)
  • baker's yeast (Saccharomyces cerevisiae)
  • barley (Hordeum vulgare)
  • beta proteobacteria (Neisseria meningitidis)
  • bread mold (Neurospora crassa)
  • Chromalveolata (Phytophthora infestans)
  • common water flea (Daphnia pulex)
  • corn (Zea mays)
  • E. coli (Escherichia coli)
  • filamentous fungi (Aspergillus nidulans)
  • Firmicute bacteria (Streptococcus pneumoniae)
  • fission yeast (Schizosaccharomyces pombe)
  • green algae (Chlamydomonas reinhardtii)
  • honey bee (Apis mellifera)
  • K. lactis yeast (Kluyveromyces lactis)
  • loblloly pine (Pinus taeda)
  • malaria parasite (Plasmodium falciparum)
  • medicago trunc (Medicago Truncatula)
  • moss (Physcomitrella patens)
  • orangutan (Pongo pygmaeus)
  • pig (Sus scrofa)
  • rainbow trout (Oncorhynchus mykiss)
  • rice (Oryza sativa)
  • rice blast fungus (Magnaporthe grisea)
  • schistosome parasite (Schistosoma mansoni)
  • sea anemone (Nematostella vectensis)
  • sea urchin (Strongylocentrotus purpuratus)
  • sorghum (Sorghum bicolor)
  • soybean (Glycine max)
  • stem rust fungus (Puccinia graminis)
  • sugarcane (Saccharum officinarum)
  • thale cress (Arabidopsis thaliana)
  • tomato (Lycopersicon esculentum)
  • toxoplasmosis (Toxoplasma gondii)
  • Trichoplax (Trichoplax adhaerens)
  • wheat (Triticum aestivum)
  • worm (Caenorhabditis elegans)

Evolution for LRP4 Gene

ENSEMBL:
Gene Tree for LRP4 (if available)
TreeFam:
Gene Tree for LRP4 (if available)

Paralogs for LRP4 Gene

Paralogs for LRP4 Gene

(13) SIMAP similar genes for LRP4 Gene using alignment to 3 proteins:

genes like me logo Genes that share paralogs with LRP4: view

Variants for LRP4 Gene

Sequence variations from dbSNP and Humsavar for LRP4 Gene

SNP ID Clin Chr 11 pos Variation AA Info Type
rs1020244365 uncertain-significance, Syndactyly Cenani Lenz type 46,890,114(-) G/A coding_sequence_variant, genic_upstream_transcript_variant, missense_variant
rs10769215 benign, Syndactyly Cenani Lenz type 46,893,047(-) A/G coding_sequence_variant, genic_upstream_transcript_variant, synonymous_variant
rs10838623 benign, Syndactyly Cenani Lenz type 46,857,966(-) G/A 3_prime_UTR_variant, genic_downstream_transcript_variant
rs10838631 benign, Syndactyly Cenani Lenz type 46,893,032(-) T/C coding_sequence_variant, genic_upstream_transcript_variant, synonymous_variant
rs111383894 benign, Syndactyly Cenani Lenz type 46,918,567(-) G/A 5_prime_UTR_variant, genic_upstream_transcript_variant, upstream_transcript_variant

Structural Variations from Database of Genomic Variants (DGV) for LRP4 Gene

Variant ID Type Subtype PubMed ID
esv3418464 CNV insertion 20981092
nsv310 CNV insertion 18451855

Variation tolerance for LRP4 Gene

Residual Variation Intolerance Score: 0.982% of all genes are more intolerant (likely to be disease-causing)
Gene Damage Index Score: 21.64; 99.31% of all genes are more intolerant (likely to be disease-causing)

Additional Variant Information for LRP4 Gene

Human Gene Mutation Database (HGMD)
LRP4
SNPedia medical, phenotypic, and genealogical associations of SNPs for
LRP4

No data available for Polymorphic Variants from UniProtKB/Swiss-Prot for LRP4 Gene

Disorders for LRP4 Gene

MalaCards: The human disease database

(17) MalaCards diseases for LRP4 Gene - From: HGMD, OMIM, ClinVar, GTR, Orphanet, Swiss-Prot, DISEASES, and GeneCards

- elite association - COSMIC cancer census association via MalaCards
Search LRP4 in MalaCards View complete list of genes associated with diseases

UniProtKB/Swiss-Prot

LRP4_HUMAN
  • Cenani-Lenz syndactyly syndrome (CLSS) [MIM:212780]: A congenital malformation syndrome defined as complete and complex syndactyly of the hands combined with malformations of the forearm bones and similar manifestations in the lower limbs. It is characterized by fusion and disorganization of metacarpal and phalangeal bones, radius and ulnar shortening, radioulnar synostosis, and severe syndactyly of hands and feet. {ECO:0000269 PubMed:20381006}. Note=The disease is caused by mutations affecting the gene represented in this entry.
  • Myasthenic syndrome, congenital, 17 (CMS17) [MIM:616304]: A form of congenital myasthenic syndrome, a group of disorders characterized by failure of neuromuscular transmission, including pre-synaptic, synaptic, and post-synaptic disorders that are not of autoimmune origin. Clinical features are easy fatigability and muscle weakness affecting the axial and limb muscles (with hypotonia in early-onset forms), the ocular muscles (leading to ptosis and ophthalmoplegia), and the facial and bulbar musculature (affecting sucking and swallowing, and leading to dysphonia). The symptoms fluctuate and worsen with physical effort. {ECO:0000269 PubMed:24234652}. Note=The disease is caused by mutations affecting the gene represented in this entry.
  • Sclerosteosis 2 (SOST2) [MIM:614305]: A sclerosing bone dysplasia characterized by a generalized hyperostosis and sclerosis leading to a markedly thickened skull, with mandible, ribs, clavicles and all long bones also being affected. Due to narrowing of the foramina of the cranial nerves, facial nerve palsy, hearing loss and atrophy of the optic nerves can occur. Sclerosteosis is clinically and radiologically very similar to van Buchem disease, mainly differentiated by hand malformations and a large stature in sclerosteosis patients. {ECO:0000269 PubMed:21471202, ECO:0000269 PubMed:24234652}. Note=The disease is caused by mutations affecting the gene represented in this entry.

Additional Disease Information for LRP4

Genetic Association Database
(GAD)
Human Genome Epidemiology Navigator
(HuGE)
ATLAS of Genetics and Cytogenetics in Oncology and Haematology
genes like me logo Genes that share disorders with LRP4: view

No data available for Genatlas for LRP4 Gene

Publications for LRP4 Gene

  1. Identification of high-molecular-weight proteins with multiple EGF-like motifs by motif-trap screening. (PMID: 9693030) Nakayama M … Ohara O (Genomics 1998) 2 3 4 58
  2. LRP4 third β-propeller domain mutations cause novel congenital myasthenia by compromising agrin-mediated MuSK signaling in a position-specific manner. (PMID: 24234652) Ohkawara B … Ohno K (Human molecular genetics 2014) 3 4 58
  3. Bone overgrowth-associated mutations in the LRP4 gene impair sclerostin facilitator function. (PMID: 21471202) Leupin O … Kneissel M (The Journal of biological chemistry 2011) 3 4 58
  4. Genetic predictors of fibrin D-dimer levels in healthy adults. (PMID: 21502573) Smith NL … Hayward C (Circulation 2011) 3 44 58
  5. Analysis of recently identified osteoporosis susceptibility genes in Han Chinese women. (PMID: 20554715) Liu JM … Ning G (The Journal of clinical endocrinology and metabolism 2010) 3 44 58

Products for LRP4 Gene

Sources for LRP4 Gene

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