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HMGCR Gene

protein-coding   GIFtS: 71
GCID: GC05P074635

3-Hydroxy-3-Methylglutaryl-CoA Reductase

(Previous name: 3-hydroxy-3-methylglutaryl-Coenzyme A reductase)
  See HMGCR-related diseases
at  

(According to 1HGNC, 2Entrez Gene,
3UniProtKB/Swiss-Prot, 4UniProtKB/TrEMBL, 5OMIM, 6GeneLoc, 7Ensembl, 8DME, 9miRBase, 10fRNAdb, 12H-InvDB, 13NCBI, 14NONCODE, and/or 15RNAdb)
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Aliases
3-Hydroxy-3-Methylglutaryl-CoA Reductase1 2     HMG-CoA Reductase2 3
3-Hydroxy-3-Methylglutaryl-Coenzyme A Reductase1 2     EC 1.1.1.343 8
3-Hydroxy-3-Methylglutaryl CoA Reductase (NADPH)1 2     LDLCQ32 5
Hydroxymethylglutaryl-CoA Reductase1 2     EC 1.1.18

External Ids:    HGNC: 50061   Entrez Gene: 31562   Ensembl: ENSG000001131617   OMIM: 1429105   UniProtKB: P040353   

Export aliases for HMGCR gene to outside databases

Previous GC identifers: GC05P073440 GC05P074869 GC05P074671 GC05P074717 GC05P074668 GC05P069838


(According to Entrez Gene, GeneCards, Tocris Bioscience, Wikipedia's Gene Wiki, PharmGKB,
UniProtKB/Swiss-Prot, and/or UniProtKB/TrEMBL)
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Entrez Gene summary for HMGCR Gene:
HMG-CoA reductase is the rate-limiting enzyme for cholesterol synthesis and is regulated via a negative feedback
mechanism mediated by sterols and non-sterol metabolites derived from mevalonate, the product of the reaction
catalyzed by reductase. Normally in mammalian cells this enzyme is suppressed by cholesterol derived from the
internalization and degradation of low density lipoprotein (LDL) via the LDL receptor. Competitive inhibitors of
the reductase induce the expression of LDL receptors in the liver, which in turn increases the catabolism of
plasma LDL and lowers the plasma concentration of cholesterol, an important determinant of atherosclerosis.
Alternatively spliced transcript variants encoding different isoforms have been found for this gene. (provided by
RefSeq, Aug 2008)

GeneCards Summary for HMGCR Gene:
HMGCR (3-hydroxy-3-methylglutaryl-CoA reductase) is a protein-coding gene. Diseases associated with HMGCR include smith-lemli-opitz syndrome, and xanthomatosis. GO annotations related to this gene include protein homodimerization activity and NADPH binding.

UniProtKB/Swiss-Prot: HMDH_HUMAN, P04035
Function: Transmembrane glycoprotein that is the rate-limiting enzyme in cholesterol biosynthesis as well as in
the biosynthesis of nonsterol isoprenoids that are essential for normal cell function including ubiquinone and
geranylgeranyl proteins

summary for HMGCR Gene:
3-hydroxy-3-methyl-glutaryl-CoA reductase (HMG-CoA reductase) catalyzes the rate limiting step in
cholesterol biosynthesis - the conversion of 3-hydroxy-3-methyl-glutaryl-CoA into mevalonic acid. This
transmembrane enzyme is bound to the endoplasmic reticulum and is ubiquitously expressed. When cholesterol
levels falls, HMG-CoA reductase is transcriptionally upregulated by sterol regulatory element binding
protein (SREBP), which binds to the sterol regulatory element (SRE) in the 5' region of the HMG-CoA gene.
When cholesterol levels are elevated, short-term regulation of HMG-CoA is mediated by Ser872 phosphorylation
by AMP-activated protein kinase and long-term regulation is mediated by the increased sensitivity of this
enzyme to proteolytic breakdown. HMG-CoA reductase is the pharmalogical target for statins and the human
gene encoding this enzyme is localized to 5q13.3-q14.

Gene Wiki entry for HMGCR (HMG-CoA reductase) Gene

PharmGKB "VIP" summary for HMGCR


(According to GeneLoc and/or HGNC, and/or
Entrez Gene (NCBI build 37),
and/or miRBase,
Genomic Views according to UCSC (hg19) and Ensembl (release 75), Regulatory elements and Epigenetics data according to QIAGEN, and/or SwitchGear Genomics)
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RefSeq DNA sequence at NCBI GenBank:
NC_000005.9  NC_018916.2  NT_034772.7  
Regulatory elements:
   Regulatory transcription factor binding sites in the HMGCR gene promoter:
         SREBP-1c   SREBP-1b   PPAR-alpha   PPAR-gamma1   CBF-A   SREBP-1a   c-Myb   PPAR-gamma2   CP1C   CBF(2)   
         Other transcription factors

SwitchGear Promoter luciferase reporter plasmidHMGCR promoter sequence
   Search Chromatin IP Primers for HMGCR

Epigenetics:
DNA Methylation CpG Assay Predesigned for Pyrosequencing in human, mouse, rat HMGCR


Genomic Location:
Genomic View: UCSC Golden Path with GeneCards custom track

Entrez Gene cytogenetic band: 5q13.3-q14   Ensembl cytogenetic band:  5q13.3   HGNC cytogenetic band: 5q13.3-q14

HMGCR Gene in genomic location: bands according to Ensembl, locations according to (and/or Entrez Gene and/or Ensembl if different)
HMGCR gene location

GeneLoc information about chromosome 5         GeneLoc Exon Structure

GeneLoc location for GC05P074635:  view genomic region     (about GC identifiers)

Start:
74,632,154 bp from pter      End:
74,657,929 bp from pter
Size:
25,776 bases      Orientation:
plus strand

(According to 1UniProtKB, HORDE, 2neXtProt, Ensembl, and/or Reactome, Modification sites according to PhosphoSitePlus, Specific Peptides from DME, RefSeq according to NCBI, PDB rendering according to OCA and/or Proteopedia, Recombinant Proteins from EMD Millipore, R&D Systems, GenScript, Enzo Life Sciences, OriGene, Novus Biologicals, Sino Biological, ProSpec, Cloud-Clone Corp., eBioscience, and/or antibodies-online,
Biochemical Assays by EMD Millipore, R&D Systems, OriGene, GenScript, Cell Signaling Technology, Enzo Life Sciences, Cloud-Clone Corp., eBioscience, and/or antibodies-online, Antibodies by EMD Millipore, R&D Systems, Cell Signaling Technology, OriGene, Novus Biologicals, Thermo Fisher Scientific, Abcam, Cloud-Clone Corp, antibodies-online, and/or others.)
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UniProtKB/Swiss-Prot: HMDH_HUMAN, P04035 (See protein sequence)
Recommended Name: 3-hydroxy-3-methylglutaryl-coenzyme A reductase  
Size: 888 amino acids; 97476 Da
Subunit: Homodimer. Interacts (via its SSD) with INSIG1; the interaction, accelerated by sterols, leads to the
recruitment of HMGCR to AMFR/gp78 for its ubiquitination by the sterol-mediated ERAD pathway. Interacts with
UBIAD1
Sequence caution: Sequence=BAH12375.1; Type=Frameshift; Positions=122;
Selected PDB 3D structures from and Proteopedia for HMGCR (see all 22):
1DQ8 (3D)        1DQ9 (3D)        1DQA (3D)        1HW8 (3D)        1HW9 (3D)        1HWI (3D)    
Secondary accessions: B7Z3Y9 Q8N190
Alternative splicing: 3 isoforms:  P04035-1   P04035-2   P04035-3   (Most highly expressed transcript in skin, esophagus, and uterine cervix)

Explore the universe of human proteins at neXtProt for HMGCR: NX_P04035

Explore proteomics data for HMGCR at MOPED

Post-translational modifications: 

  • N-glycosylated. Deglycosylated by NGLY1 on release from the endoplasmic reticulum (ER) in a sterol-mediated manner1
  • Undergoes sterol-mediated ubiquitination and ER-association degradation (ERAD). Accumulation of sterols in the
    endoplasmic reticulum (ER) membrane, triggers binding of the reductase to the ER membrane protein INSIG1. This
    INSIG1 binding leads to the recruitment of the ubiquitin ligase, AMFR/gp78, initiating ubiquitination of the
    reductase. The ubiquitinated reductase is then extracted from the ER membrane and delivered to cytosolic 26S
    proteosomes by a mechanism probably mediated by the ATPase Valosin-containing protein VCP/p97. Lys-248 is the
    main site of ubiquitination. Ubiquitination is enhanced by the presence of a geranylgeranylated protein1
  • Ubiquitination2 at Lys89, Lys248, Lys288, Lys299, Lys381, Lys396, Lys401, Lys460, Lys474, Lys480,
                                 Lys502, Lys619, Lys633, Lys662, Lys704, Lys715, Lys864, Lys873
  • Glycosylation2 at Asn281, Asn296, Asn419, Asn518, Asn870
  • Modification sites at PhosphoSitePlus
  • Selected DME Specific Peptides for HMGCR (P04035) (see all 40)
     PMATTEG  GRGKSVV  GSAMAGS  LWQFYLS 


    See HMGCR Protein Expression from SPIRE MOPED, PaxDB, and MaxQB

    REFSEQ proteins (2 alternative transcripts): 
    NP_000850.1  NP_001124468.1  

    ENSEMBL proteins: 
     ENSP00000426745   ENSP00000287936   ENSP00000340816   ENSP00000427340   ENSP00000409100  
     ENSP00000421378   ENSP00000420871  

    HMGCR Human Recombinant Protein Products:

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    Novus Biologicals HMGCR Lysate
    Browse Sino Biological Recombinant Proteins
    Browse Sino Biological Cell Lysates
    Browse ProSpec Recombinant Proteins
    Cloud-Clone Corp. Proteins for HMGCR

     
    Search eBioscience for Proteins for HMGCR 

     
    antibodies-online proteins for HMGCR (2 products) 

     
    antibodies-online peptides for HMGCR

    HMGCR Antibody Products:

    EMD Millipore Mono- and Polyclonal Antibodies for the study of HMGCR
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    antibodies-online antibodies for HMGCR (43 products) 

    HMGCR Assay Products:

    Browse Kits and Assays available from EMD Millipore
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    GenScript Custom Assay Services for HMGCR
    Browse Enzo Life Sciences for kits & assays
    Cloud-Clone Corp. ELISAs for HMGCR
    Cloud-Clone Corp. CLIAs for HMGCR
    Search eBioscience for ELISAs for HMGCR 
    antibodies-online kits for HMGCR (24 products) 


    (According to HGNC, IUPHAR, InterPro, ProtoNet, UniProtKB, and/or BLOCKS, Sets of similar genes according to GeneDecks)
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    IUPHAR Guide to PHARMACOLOGY protein family classification: hydroxymethylglutaryl-CoA reductase
    Lanosterol biosynthesis pathway

    Selected InterPro protein domains (see all 9):
     IPR009023 HMG_CoA_Rdtase_NAD(P)-bd
     IPR000731 SSD
     IPR004554 HMG_CoA_Rdtase_eu_arc
     IPR004816 HMG_CoA_Rdtase_metazoan
     IPR023282 HMG_CoA_Rdtase_N

    Graphical View of Domain Structure for InterPro Entry P04035

    ProtoNet protein and cluster: P04035

    1 Blocks protein domain: IPB002202 Hydroxymethylglutaryl-coenzyme A reductase

    UniProtKB/Swiss-Prot: HMDH_HUMAN, P04035
    Similarity: Belongs to the HMG-CoA reductase family
    Similarity: Contains 1 SSD (sterol-sensing) domain


    Find genes that share domains with HMGCR           About GenesLikeMe


    (According to 1UniProtKB, Genatlas, LifeMap Discovery™, IUBMB, and/or 2DME, Human phenotypes from GenomeRNAi, Animal models from MGI Mar 06 2013, genOway,
    transcription factor targeting from QIAGEN and/or HOMER, miRNA Gene Targets from miRTarBase, shRNA from OriGene, siRNAs from OriGene, QIAGEN, microRNA from QIAGEN, SwitchGear Genomics, Gene Editing from DNA2.0, Clones from OriGene, GenScript, Sino Biological, DNA2.0, Vector BioLabs and/or Addgene, Cell Lines from GenScript, ESI BIO, In Situ Hybridization Assays from Advanced Cell Diagnostics, Flow cytometry from eBioscience, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene.)
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    Molecular Function:

         UniProtKB/Swiss-Prot Summary: HMDH_HUMAN, P04035
    Function: Transmembrane glycoprotein that is the rate-limiting enzyme in cholesterol biosynthesis as well as in
    the biosynthesis of nonsterol isoprenoids that are essential for normal cell function including ubiquinone and
    geranylgeranyl proteins
    Catalytic activity: (R)-mevalonate + CoA + 2 NADP(+) = (S)-3-hydroxy-3-methylglutaryl-CoA + 2 NADPH
    Enzyme regulation: Regulated by a negative feedback mechanism through sterols and non-sterol metabolites derived
    from mevalonate. Inhibited by statins, a class of hypolipidemic agents used as pharmaceuticals to lower
    cholesterol levels in individuals at risk from cardiovascular disease due to hypercholesterolemia. Inhibition of
    HMGCR in the liver stimulates the LDL-receptors, which results in an increased clearance of LDL from the
    bloodstream and a decrease in blood cholesterol levels. The first results can be seen after one week of statin
    use and the effect is maximal after four to six weeks

         Genatlas biochemistry entry for HMGCR:
    3-hydroxy-3-methylglutaryl-CoA reductase,catalyzing a rate-limiting step in cholesterol biosynthesis

         Enzyme Numbers (IUBMB): EC 1.1.1.341 2 EC 1.1.12

         Gene Ontology (GO): Selected molecular function terms (see all 9):    About this table

    GO IDQualified GO termEvidencePubMed IDs
    GO:0004420hydroxymethylglutaryl-CoA reductase (NADPH) activity IEA--
    GO:0005515protein binding IPI--
    GO:0008158hedgehog receptor activity ----
    GO:0016616oxidoreductase activity, acting on the CH-OH group of donors, NAD or NADP as acceptor ----
    GO:0042282hydroxymethylglutaryl-CoA reductase activity IDA2991281
         
    Find genes that share ontologies with HMGCR           About GenesLikeMe


    Phenotypes:
         2 MGI mutant phenotypes (inferred from 1 allele(MGI details for Hmgcr):
     mortality/aging  normal 

    Find genes that share phenotypes with HMGCR           About GenesLikeMe

    Animal Models:
         MGI mouse knock-out Hmgcrtm1Ishi for HMGCR

       genOway: Develop your customized and physiologically relevant rodent model for HMGCR

    miRNA
    Products:
        
    miRTarBase miRNAs that target HMGCR:
    hsa-mir-92a-3p (MIRT048980), hsa-mir-193b-3p (MIRT016598), hsa-mir-29c-3p (MIRT020381), hsa-mir-615-3p (MIRT040026), hsa-mir-335-5p (MIRT018888), hsa-mir-32-5p (MIRT028289)

    Block miRNA regulation of human, mouse, rat HMGCR using miScript Target Protectors
    Selected qRT-PCR Assays for microRNAs that regulate HMGCR (see all 39):
    hsa-miR-300 hsa-miR-513a-5p hsa-miR-29a hsa-miR-374a* hsa-miR-374c hsa-miR-149 hsa-miR-3613-3p hsa-miR-556-3p
    SwitchGear 3'UTR luciferase reporter plasmidHMGCR 3' UTR sequence
    Inhib. RNA
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    OriGene RNAi products in human, mouse, rat for HMGCR
    Predesigned siRNA for gene silencing in human, mouse, rat HMGCR

    Gene Editing
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    DNA2.0 Custom Protein Engineering Service for HMGCR

    Clone
    Products:
         
    OriGene clones in human, mouse for HMGCR (see all 11)
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    GenScript: all cDNA clones in your preferred vector (see all 2): HMGCR (NM_000859)
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    DNA2.0 Custom Codon Optimized Gene Synthesis Service for HMGCR
    Vector BioLabs ready-to-use adenovirus/AAV for human, mouse, rat HMGCR
    Addgene plasmids for HMGCR 

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    Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for HMGCR

    Flow Cytometry
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    eBioscience FlowRNA Probe Sets ( VA1-12329) for HMGCR 


    (According to UniProtKB, COMPARTMENTS Subcellular localization database, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene.)
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    Subcellular locations from UniProtKB/Swiss-Prot
    HMDH_HUMAN, P04035: Endoplasmic reticulum membrane; Multi-pass membrane protein
    Subcellular locations from COMPARTMENTS: 

    CompartmentConfidence
    endoplasmic reticulum5
    peroxisome5
    extracellular3
    plasma membrane3
    cytosol2
    mitochondrion2
    nucleus2
    cytoskeleton1

    Gene Ontology (GO): Selected cellular component terms (see all 6):    About this table

    GO IDQualified GO termEvidencePubMed IDs
    GO:0005778peroxisomal membrane IDA17180682
    GO:0005783endoplasmic reticulum IDA17180682
    GO:0005789endoplasmic reticulum membrane TAS2991281
    GO:0016020membrane ----
    GO:0016021integral component of membrane IEA--

    Find genes that share ontologies with HMGCR           About GenesLikeMe


    (SuperPaths according to PathCards, Pathways according to R&D Systems, Cell Signaling Technology, KEGG, PharmGKB, BioSystems, Sino Biological, Reactome, Tocris Bioscience, GeneGo (Thomson Reuters), QIAGEN, and/or UniProtKB, Sets of similar genes according to GeneDecks, Interaction Networks according to QIAGEN, and/or STRING, Interactions according to 1UniProtKB, 2MINT, 3I2D, and/or 4STRING, with links to IntAct and Ensembl, Ontologies according to Gene Ontology Consortium 01 Apr 2014 via Entrez Gene).
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    SuperPaths for HMGCR About   (see all 14)  
    See pathways by source

    SuperPathContained pathways About
    1superpathway of cholesterol biosynthesis
    superpathway of cholesterol biosynthesis0.85
    Terpenoid backbone biosynthesis0.34
    Cholesterol biosynthesis0.85
    superpathway of geranylgeranyldiphosphate biosynthesis I (via mevalonate)0.00
    Cholesterol biosynthesis0.62
    mevalonate pathway I0.00
    2Regulation of Cholesterol Biosynthesis by SREBP (SREBF)
    Regulation of Cholesterol Biosynthesis by SREBP (SREBF)0.75
    SREBP signalling0.31
    Activation of Gene Expression by SREBP (SREBF)0.75
    3Metabolism
    Metabolism0.38
    Metabolism of lipids and lipoproteins0.37
    Metabolic pathways0.38
    4Statin Pathway
    Statin Pathway0.73
    Statin Pathway, Pharmacodynamics0.73
    5Fatty acid, triacylglycerol, and ketone body metabolism
    Fatty acid, triacylglycerol, and ketone body metabolism0.65
    PPARA Activates Gene Expression0.63
    Regulation of Lipid Metabolism by Peroxisome proliferator-activated receptor alpha (PPARalpha)0.65


    Find genes that share SuperPaths with HMGCR           About GenesLikeMe

    Pathways by source                                                                                                                                                                 See SuperPaths
    Show all pathways

    1 Downloadable PowerPoint Slide of GeneGlobe Pathway Central Maps for HMGCR
        AMPK Enzyme Complex Pathway

    Selected BioSystems Pathways for HMGCR (see all 9)
        Cholesterol biosynthesis
    Statin Pathway
    AMPK signaling
    Integrated Breast Cancer Pathway
    SREBP signalling

    3 PharmGKB Pathways for HMGCR
        Bisphosphonate Pathway, Pharmacodynamics
    Metformin Pathway, Pharmacodynamic
    Statin Pathway, Pharmacodynamics

    3 Kegg Pathways  (Kegg details for HMGCR):
        Terpenoid backbone biosynthesis
    Metabolic pathways
    Bile secretion

    UniProtKB/Swiss-Prot: HMDH_HUMAN, P04035
    Pathway: Metabolic intermediate biosynthesis; (R)-mevalonate biosynthesis; (R)-mevalonate from acetyl-CoA: step
    3/3

        Pathway & Disease-focused RT2 Profiler PCR Arrays including HMGCR: 

              Lipoprotein Signaling & Cholesterol Metabolism in human mouse rat
              Prostate Cancer in human mouse rat
              Fatty Liver in human mouse rat

    Interactions:

        Search GeneGlobe Interaction Network for HMGCR

    STRING Interaction Network Preview (showing 5 interactants - click image to see 25)

    Selected Interacting proteins for HMGCR (P040351, 3 ENSP000002879364) via UniProtKB, MINT, STRING, and/or I2D (see all 149)
    InteractantInteraction Details
    GeneCardExternal ID(s)
    ENSG00000206286Q8N1B43I2D: score=1 
    ENSG00000224455Q8N1B43I2D: score=1 
    ENSG00000225590Q8N1B43I2D: score=1 
    ENSG00000228425Q8N1B43I2D: score=1 
    ENSG00000236014Q8N1B43I2D: score=1 
    About this table

    Gene Ontology (GO): Selected biological process terms (see all 26):    About this table

    GO IDQualified GO termEvidencePubMed IDs
    GO:0006695cholesterol biosynthetic process TAS--
    GO:0006743ubiquinone metabolic process IEA--
    GO:0007568aging IEA--
    GO:0007584response to nutrient IEA--
    GO:0008284positive regulation of cell proliferation ----

    Find genes that share ontologies with HMGCR           About GenesLikeMe



    (Chemical Compounds according to UniProtKB, Enzo Life Sciences, EMD Millipore, Tocris Bioscience, ApexBio, HMDB, BitterDB, and/or Novoseek, Ligands according to IUPHAR, and Drugs according to DrugBank, Enzo Life Sciences, and/or PharmGKB)
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    Browse Small Molecules at EMD Millipore
       Enzo Life Sciences drugs & compounds for HMGCR
      Browse compounds at ApexBio 

    Compounds for HMGCR available from Tocris Bioscience    About this table
    CompoundAction CAS #
    Fluvastatin sodiumHMG-CoA reductase inhibitor[93957-55-2]
    SimvastatinHMG-CoA reductase inhibitor[79902-63-9]
    Pitavastatin calciumHMG-CoA reductase inhibitor[147526-32-7]
    Atorvastatin hemicalcium saltHMG-CoA reductase inhibitor[134523-03-8]
    LovastatinHMG-CoA reductase inhibitor[75330-75-5]

    8 HMDB Compounds for HMGCR    About this table
    CompoundSynonyms CAS #PubMed Ids
    3-Hydroxy-3-methylglutaryl-CoA(S)-3-hydroxy-3-methylglutaryl-CoA (see all 15)1553-55-5--
    AtorvastatinAtorvastatin acid (see all 4)134523-00-5--
    Coenzyme AAcetoacetyl coenzyme A sodium salt (see all 21)85-61-0--
    Mevalonic acid(3RS)-Mevalonate (see all 24)150-97-0--
    NADPAdenine-nicotinamide dinucleotide phosphate (see all 18)53-59-8--
    NADPH2'-(dihydrogen phosphate) 5'-(trihydrogen pyrophosphate) Adenosine 5'-ester with 1,4-dihydro-1-b-D-ribofuranosylnicotinamide (see all 23)53-57-6--
    Pravastatin3beta-Hydroxycompactin (see all 7)81093-37-0--
    Simvastatin(+)-Simvastatin (see all 28)79902-63-9--

    Selected DrugBank Compounds for HMGCR (see all 17)    About this table
    CompoundSynonyms CAS #TypeActionsPubMed Ids
    Lovastatin6 alpha-Methylcompactin (see all 4)75330-75-5targetinhibitor12405293 12172803 12082550 12369143 12204108 11752352 12214063 19249983
    Rosuvastatin-- 287714-41-4targetinhibitor11501230 11256847 16128575 11752352 12481202 12539816 12773150 11772327
    Mevastatin-- 73573-88-3targetinhibitor12671581 12433810 12852453 11752352 12677170 12518039 10592235
    Atorvastatin-- 134523-00-5targetinhibitor15748157 15109613 12808485 11752352 12769127 15474503
    CerivastatinCerivastatin sodium (see all 2)145599-86-6targetinhibitor10740137 11057870 11470741 11752352 10193776 11448925
    SimvastatinSimvastatin [Usan:Ban:Inn] (see all 4)79902-63-9targetinhibitor12671581 12433810 12852453 12677170 11752352 12518039
    NADHbeta-DPNH (see all 18)606-68-8target--14511985 15755483 17139284 15152097 17016423
    (S)-Hmg-Coa-- --target--17139284 17016423
    1,4-Dithiothreitol-- --target--17139284 17016423
    2'-Monophosphoadenosine 5'-Diphosphoribose-- --target--17139284 17016423

    Selected IUPHAR Ligands for HMGCR (hydroxymethylglutaryl-CoA reductase) (see all 38)    About this table
    LigandTypeActionAffinityPubmed IDs
    (3R,5S,6E)-7-[4-(4-fluorophenyl)-1-phenyl-2-(propan-2-yl)-1H-pyrrol-3-yl]-3,5-dihydroxyhept-6-enoate
    InhibitorInhibition112153213
    (3R,5R)-7-[1-cyclohexyl-4-(4-fluorophenyl)-2-(propan-2-yl)-1H-pyrrol-3-yl]-3,5-dihydroxyheptanoate
    InhibitorInhibition112153213
    (3R,5R)-7-[4-(4-fluorophenyl)-1-phenyl-2-(propan-2-yl)-1H-pyrrol-3-yl]-3,5-dihydroxyheptanoate
    InhibitorInhibition112153213
    (3R,5S,6E)-7-[4-(4-fluorophenyl)-1,2-bis(propan-2-yl)-1H-pyrrol-3-yl]-3,5-dihydroxyhept-6-enoate
    InhibitorInhibition10.32153213
    (3R,5S,6E)-7-[1-cyclohexyl-4-(4-fluorophenyl)-2-(propan-2-yl)-1H-pyrrol-3-yl]-3,5-dihydroxyhept-6-enoate
    InhibitorInhibition102153213
    (3R,5R)-7-(5-{[(4-cyanophenyl)methyl]carbamoyl}-3-(4-fluorophenyl)-4-phenyl-1-(propan-2-yl)-1H-pyrrol-2-yl)-3,5-dihydroxyheptanoate
    InhibitorInhibition9.717574412
    (3R,5R)-7-[4-(4-fluorophenyl)-2-methyl-1-(propan-2-yl)-1H-pyrrol-3-yl]-3,5-dihydroxyheptanoate
    InhibitorInhibition9.572153213
    (3R,5R)-7-[3-(4-fluorophenyl)-8-oxo-7-phenyl-1-(propan-2-yl)-1H,4H,5H,6H,7H,8H-pyrrolo[2,3-c]azepin-2-yl]-3,5-dihydroxyheptanoate
    InhibitorInhibition9.5217574411
    (3R)-6-[2-(4-fluorophenyl)-4-[(4-fluorophenyl)sulfanyl]-6-(propan-2-yl)phenoxy]-3,5-dihydroxyhexanoate
    InhibitorInhibition9.31656041
    (3R,5R)-7-[4-(4-fluorophenyl)-2,5-dimethyl-1-phenyl-1H-pyrrol-3-yl]-3,5-dihydroxyheptanoate
    InhibitorInhibition9.242153213

    Selected Novoseek inferred chemical compound relationships for HMGCR gene (see all 134)    About this table
    Compound   -log (P-Val)   Hits   PubMed IDs for Articles with Shared Sentences (# sentences)
    simvastatin 95.9 368 1627634 (5), 2083515 (4), 2077527 (3), 8521762 (3) (see all 99)
    pravastatin 95.6 259 8318553 (2), 8366183 (2), 9542477 (2), 1904355 (2) (see all 99)
    mevalonate 95.3 226 17412884 (3), 1908464 (3), 1524429 (3), 9237866 (2) (see all 99)
    lovastatin 95.3 267 11205904 (4), 12843138 (3), 10146977 (2), 8082513 (2) (see all 99)
    fluvastatin 93.4 145 8882381 (8), 1854361 (3), 7604801 (3), 8198018 (2) (see all 99)
    atorvastatin 93.4 174 9793596 (4), 17651990 (3), 19930424 (3), 9485553 (3) (see all 99)
    cerivastatin 92.4 84 11129127 (5), 14720086 (4), 17151817 (2), 11470741 (2) (see all 59)
    cholesterol 92.3 1117 10170452 (4), 10713930 (4), 10791271 (4), 11716837 (4) (see all 99)
    3-hydroxy-3-methylglutaryl-coa 92.1 294 2307197 (6), 9441587 (6), 12514264 (6), 7797599 (5) (see all 99)
    geranylgeranyl pyrophosphate 89 27 19464372 (2), 10996358 (1), 11884371 (1), 15850779 (1) (see all 20)

    6 PharmGKB related drug/compound annotations for HMGCR gene    About this table
    Drug/compound PharmGKB Annotation
    atorvastatin  CA  
    fluvastatinCA  
    hmg coa reductase inhibitorsCA  
    lovastatinCA  
    pravastatinCA  
    simvastatinCA  



    Find genes that share compounds with HMGCR           About GenesLikeMe



    (Secondary structures according to fRNAdb,
    GenBank/EMBL/DDBJ Accessions according to
    Unigene (Build 236 Homo sapiens; Apr 25 2013) or GenBank,
    RefSeq according to Entrez Gene,
    DOTS (version 10), and/or AceView, transcript ids from Ensembl with links to UCSC,
    exon structure from GeneLoc, alternative splicing isoforms according to ASD and/or ECgene,
    siRNAs from OriGene, QIAGEN, shRNA from OriGene, microRNA from QIAGEN, SwitchGear Genomics,
    Tagged/untagged cDNA clones from OriGene, GenScript, DNA2.0, Vector BioLabs, and/or Addgene, Primers from OriGene, and/or QIAGEN, Flow cytometry from eBioscience )
    About This Section

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    REFSEQ mRNAs for HMGCR gene (2 alternative transcripts): 
    NM_000859.2  NM_001130996.1  

    Unigene Cluster for HMGCR:

    3-hydroxy-3-methylglutaryl-CoA reductase
    Hs.628096  [show with all ESTs]
    Unigene Representative Sequence: NM_000859
    13 Ensembl transcripts including schematic representations, and UCSC links where relevant:
    ENST00000511206(uc010izn.1) ENST00000287936(uc011cst.2 uc003kdp.3 uc003kdq.3 uc010izo.3 uc010izp.3)
    ENST00000343975 ENST00000509431 ENST00000507942 ENST00000442032 ENST00000504466
    ENST00000515776 ENST00000508070 ENST00000509085 ENST00000511986 ENST00000514315
    ENST00000512053
    miRNA
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    Selected qRT-PCR Assays for microRNAs that regulate HMGCR (see all 39):
    hsa-miR-300 hsa-miR-513a-5p hsa-miR-29a hsa-miR-374a* hsa-miR-374c hsa-miR-149 hsa-miR-3613-3p hsa-miR-556-3p
    SwitchGear 3'UTR luciferase reporter plasmidHMGCR 3' UTR sequence
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    GenScript: all cDNA clones in your preferred vector (see all 2): HMGCR (NM_000859)
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    Vector BioLabs ready-to-use adenovirus/AAV for human, mouse, rat HMGCR
    Addgene plasmids for HMGCR 
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    Flow Cytometry
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    eBioscience FlowRNA Probe Sets ( VA1-12329) for HMGCR 

    Additional mRNA sequence: 

    AK292892.1 AK296499.1 AK299655.1 AK312437.1 AY429541.1 AY429542.1 AY429543.1 AY429544.2 
    BC024180.1 BC033692.1 M11058.1 M62627.1 M62633.1 

    15 DOTS entries:

    DT.447815  DT.99928400  DT.100794644  DT.120859642  DT.100794651  DT.97844838  DT.100794648  DT.120859653 
    DT.40257342  DT.95071605  DT.95071606  DT.95071631  DT.95071640  DT.100794646  DT.97835796 

    Selected AceView cDNA sequences (see all 342):

    AU132001 AI968551 BQ045225 BQ774835 AL518628 AA503060 AI381010 BX954962 
    BG152850 T68833 BM984122 AI056875 CF264725 BQ688371 F13542 AI961446 
    AA577170 AW974542 F04965 F04434 AI962842 CA432613 AI056859 D29148 

    GeneLoc Exon Structure

    5 Alternative Splicing Database (ASD) splice patterns (SP) for HMGCR    About this scheme

    ExUns: 1 ^ 2 ^ 3 ^ 4 ^ 5 ^ 6 ^ 7a · 7b ^ 8 ^ 9 ^ 10a · 10b ^ 11 ^ 12 ^ 13 ^ 14 ^ 15 ^ 16a · 16b ^ 17 ^ 18a · 18b ^ 19a · 19b ^ 20a · 20b
    SP1:                                      -                                               -                                         -     -                     
    SP2:                                      -                                                                                         -     -                     
    SP3:                                                                                                                                                            
    SP4:                                                                                                                                                            
    SP5:                                                                                                                                                            


    ECgene alternative splicing isoforms for HMGCR

    (RNA expression data according to H-InvDB, NONCODE, miRBase, and RNAdb, Expression images according to data from BioGPS, Illumina Human BodyMap, and CGAP SAGE, Sets of similar genes according to GeneDecks, in vivo and in vitro expression data from LifeMap Discovery™, Protein expression images according to data from SPIRE 1MOPED, 2PaxDb, and 3MaxQB, plus additional links to SOURCE, and/or BioGPS, and/or UniProtKB,
    PCR Arrays from QIAGEN, Primers from OriGene, and/or QIAGEN, In Situ Hybridization Assays from Advanced Cell Diagnostics)
    About This Section

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    HMGCR expression in normal human tissues (normalized intensities)
    See probesets specificity/sensitivity at GeneAnnot
    About this imageBioGPS <intensity>2/3
    CGAP TAG: GAAGTGGCAG
    HMGCR Expression
    About this image


    HMGCR expression in embryonic tissues and stem cells    About this table
    Data from LifeMap, the Embryonic Development and Stem Cells Database
     selected tissues (see all 8) fully expand
     
     Brain (Nervous System)    fully expand to see all 4 entries
             Cerebral Cortex
     
     Bone (Muscoskeletal System)
             Bone Marrow
     
     Trophoblast (Extraembryonic Tissues)
             Trophoblast Cells Trophoblast
     
     Testis (Reproductive System)
             Leydig Cells Testis Interstitium
     
     Limb (Muscoskeletal System)
             Limb Bud
    HMGCR Protein expression data from MOPED1, PaxDb2 and MaxQB3    About this image

    HMGCR Protein Expression

    SOURCE GeneReport for Unigene cluster: Hs.628096
        Pathway & Disease-focused RT2 Profiler PCR Arrays including HMGCR: 
              Lipoprotein Signaling & Cholesterol Metabolism in human mouse rat
              Prostate Cancer in human mouse rat
              Fatty Liver in human mouse rat

    Primer
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    Advanced Cell Diagnostics RNAscope RNA in situ hybridization assays for HMGCR

    (Orthologs according to 1,2HomoloGene (2older version, for species not in 1newer version), 3euGenes, 4SGD , 5MGI Mar 06 2013, with possible further links to Flybase and/or WormBase, and/or 6Ensembl pan taxonomic compara , Gene Trees according to Ensembl and TreeFam)
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    This gene was present in the common ancestor of eukaryotes.

    Orthologs for HMGCR gene from Selected species (see all 29)    About this table
    Organism Taxonomic
    classification
    Gene Description Human
    Similarity
    Orthology
    Type
    Details
    mouse
    (Mus musculus)
    Mammalia Hmgcr1 , 5 3-hydroxy-3-methylglutaryl-Coenzyme A reductase1, 5 86.55(n)1
    93.57(a)1
      13 (50.65 cM)5
    153571  NM_008255.21  NP_032281.21 
     966489675 
    chicken
    (Gallus gallus)
    Aves HMGCR1 3-hydroxy-3-methylglutaryl-CoA reductase 79.09(n)
    86.29(a)
      395145  NM_204485.1  NP_989816.1 
    lizard
    (Anolis carolinensis)
    Reptilia HMGCR6
    3-hydroxy-3-methylglutaryl-CoA reductase
    84(a)
    1 ↔ 1
    2(14885587-14911381)
    African clawed frog
    (Xenopus laevis)
    Amphibia LOC3977502 X.laevis 3-hydroxy-3-methylglutaryl-coenzyme A reductase 77.06(n)    M29258.1 
    zebrafish
    (Danio rerio)
    Actinopterygii hmgcra1 3-hydroxy-3-methylglutaryl-Coenzyme A reductase a 70.29(n)
    78.34(a)
      559054  NM_001079977.1  NP_001073446.1 
    fruit fly
    (Drosophila melanogaster)
    Insecta Hmgcr1 , 3 germ-cell migration
    hydroxymethylglutaryl-CoA more3
    HMG Coenzyme A reductase1
    44(a)3
    53.44(n)1
    51.14(a)1
      3 95B13
    428031  NM_170089.21  NP_732900.11 
    worm
    (Caenorhabditis elegans)
    Secernentea F08F8.23
    hmgr-11
    Hydroxymethylglutaryl-coA reductase3
    hmgr-11
    46(a)3
    54(n)1
    48.16(a)1
      III(7430911-7433596)3
    1760471  NM_066225.51  NP_498626.21 
    baker's yeast
    (Saccharomyces cerevisiae)
    Saccharomycetes HMG2(YLR450W)4
    HMG21
    One of two isozymes of HMG-CoA reductase that convert more4
    HMG21
    57.36(n)1
    59.15(a)1
      12(1032627-1035764)4
    8511711, 4  NP_013555.11, 4 
    thale cress
    (Arabidopsis thaliana)
    eudicotyledons HMG11 HMG1 59.33(n)
    57.18(a)
      843982  NM_106299.3  NP_177775.2 
    rice
    (Oryza sativa)
    Liliopsida Os.1642 Oryza sativa (japonica cultivar-group) cDNA cloneJ more 73.22(n)    AK069555.1 


    ENSEMBL Gene Tree for HMGCR (if available)
    TreeFam Gene Tree for HMGCR (if available)

    (Paralogs according to 1HomoloGene,
    2Ensembl, and 3SIMAP, Pseudogenes according to 4Pseudogene.org Build 68)
    About This Section

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    (SNPs/Variants according to the 1NCBI SNP Database, 2Ensembl, 3PupaSUITE, 4UniProtKB, and DNA2.0, Linkage Disequilibrium by HapMap, Structural Variations(CNVs/InDels/Inversions) from the Database of Genomic Variants, Mutations from the Human Gene Mutation Database (HGMD), the Human Cytochrome P450 Allele Nomenclature Database, and the Locus Specific Mutation Databases (LSDB), Blood group antigen gene mutations by BGMUT, Resequencing Primers, Cancer Mutation PCR Arrays and Assays, and Copy Number PCR Arrays from QIAGEN)
    About This Section

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    Selected SNPs for HMGCR (see all 586)    About this table                                 

    Genomic DataTranscription Related DataAllele Frequencies
    SNP IDValidClinical
    significance
    Chr 5 posSequence#AA
    Chg
    TypeMore#Allele
    freq
    PopTotal
    sample
    More
    ----------
    rs725387121,2
    C--74639574(+) ATGAC-/AA/AC 
            
    TATAT
    2 -- int10--------
    rs1996996091,2
    C--74640967(+) TTTTT-/TAAGAG 2 -- int10--------
    rs114438961,2
    C--74644785(+) GAATT-/AAAAAA 2 -- int11Minor allele frequency- A:0.50CSA 2
    rs560213021,2
    C--74651576(+) AAAAA-/AGTGTG 2 -- int10--------
    rs172385681,2
    C,F--74924070(+) ATATAG/TAATTT 2 -- us2k15Minor allele frequency- T:0.03NS NA 346
    rs172449391,2
    C--74924095(+) AAACTA/CGACAA 2 -- us2k15Minor allele frequency- C:0.02NS NA 346
    rs1498455371,2
    C--74924133(+) AACTCA/CTATTC 2 -- us2k10--------
    rs37617381,2
    C,F,H--74924138(+) ATATTC/TCTATT 2 -- us2k127Minor allele frequency- T:0.06NS EA NA CSA WA 3168
    rs172383301,2
    C,F,H--74924261(+) CCTGTA/GACTTC 2 -- us2k125Minor allele frequency- G:0.06NS EA NA CSA WA 2810
    rs172383371,2
    C,F--74924298(+) AGCGGC/GATTAA 2 -- us2k1 tfbs35Minor allele frequency- G:0.02NS NA 264

    HapMap Linkage Disequilibrium report for HMGCR (74632154 - 74657929 bp)

    Structural Variations
         Database of Genomic Variants (DGV) 3 variations for HMGCR:    About this table    
    Variant IDTypeSubtypePubMed ID
    esv268588CNV Insertion20981092
    nsv882192CNV Loss21882294
    dgv6183n71CNV Loss21882294

    Human Gene Mutation Database (HGMD): HMGCR
    Site Specific Mutation Identification with PCR Assays
    SeqTarget long-range PCR primers for resequencing HMGCR
    DNA2.0 Custom Variant and Variant Library Synthesis for HMGCR

    (in which this Gene is Involved, According to MalaCards, OMIM, UniProtKB, the University of Copenhagen DISEASES database, Novoseek, Genatlas, GeneTests, GAD, HuGE Navigator, and/or TGDB.)
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    OMIM gene information: 142910    OMIM disorders: --

    19 diseases for HMGCR:    
    About MalaCards
    smith-lemli-opitz syndrome    xanthomatosis    cholesteryl ester storage disease    creatine phosphokinase, elevated serum
    sitosterolemia    defective apolipoprotein b-100    lichen planus pemphigoides    cerebrotendinous xanthomatosis
    hyperlipoproteinemia type iii    atherosclerosis    spondylosis    seizure disorder
    aortic valve disease    hypercholesterolemia    familial hypercholesterolemia    familial hyperlipidemia
    hypertriglyceridemia    coronary artery disease    myocardial infarction

    6 diseases from the University of Copenhagen DISEASES database for HMGCR:
    Heart disease     Atherosclerosis     Cerebrovascular accident     Familial hyperlipidemia
    Diabetes mellitus     Myopathy

    Find genes that share disorders with HMGCR           About GenesLikeMe

    Selected Novoseek inferred disease relationships for HMGCR gene (see all 95)    About this table

    Disease   -log (P-Val)   Hits   PubMed IDs for Articles with Shared Sentences (# sentences)
    hypercholesterolemia 90.7 260 11794823 (3), 10791271 (3), 8520093 (3), 8882381 (3) (see all 99)
    rhabdomyolysis 81.8 61 12841814 (2), 11773716 (2), 11573861 (2), 7873090 (2) (see all 54)
    hyperlipidemia 79.9 74 9110123 (4), 10356659 (4), 9801932 (3), 10999499 (2) (see all 59)
    dyslipidemia 79.2 67 15285698 (2), 2149540 (2), 8595642 (2), 15725039 (2) (see all 58)
    coronary heart disease 77.5 74 1899896 (2), 8983865 (2), 9202359 (2), 9674175 (2) (see all 64)
    atherosclerosis 76.4 64 8913414 (2), 9683927 (2), 7604791 (1), 9537338 (1) (see all 56)
    hypercholesterolemia familial 74.1 20 1814804 (2), 2312032 (1), 1987991 (1), 1412289 (1) (see all 19)
    myopathy 74.1 49 19078619 (4), 8725265 (2), 12241128 (2), 9300786 (1) (see all 39)
    cardiovascular diseases 72.2 72 15962651 (2), 10912119 (1), 11228171 (1), 15922259 (1) (see all 68)
    coronary artery disease 68.9 81 9248989 (6), 10713930 (2), 10559508 (2), 14727930 (2) (see all 69)

    Genetic Association Database (GAD): HMGCR
    Human Genome Epidemiology (HuGE) Navigator: HMGCR (58 documents)

    Export disorders for HMGCR gene to outside databases

    (in PubMed. Associations of this gene to articles via 1Entrez Gene, 2UniProtKB/Swiss-Prot, 3HGNC, 4GAD, 5PharmGKB, 6HMDB, 7DrugBank, 8UniProtKB/TrEMBL, 9 Novoseek, and/or 10fRNAdb)
    About This Section

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    PubMed articles for HMGCR gene, integrated from 10 sources (see all 2082):
    (articles sorted by number of sources associating them with HMGCR)
        Utopia: connect your pdf to the dynamic
    world of online information

    1. Common SNPs in HMGCR in micronesians and whites associated with LDL-cholesterol levels affect alternative splicing of exon13. (PubMed id 18802019)1, 4, 9 Burkhardt R....Breslow J.L. (Arterioscler. Thromb. Vasc. Biol. 2008)
    2. Variants in the HMG-CoA reductase (HMGCR) gene influence component phenotypes in polycystic ovary syndrome. (PubMed id 19327767)1, 4, 9 Xu N....Goodarzi M.O. (Fertil. Steril. 2010)
    3. Effect of ScrF I polymorphism in the 2nd intron of the HMGCR gene on lipid-lowering response to simvastatin in Chinese diabetic patients. (PubMed id 17870053)1, 4, 9 Ying S....Gao Y.Y. (Biochem. Biophys. Res. Commun. 2007)
    4. Genetic variation in 3-hydroxy-3-methylglutaryl CoA reductase modifies the chemopreventive activity of statins for colorectal cancer. (PubMed id 20403997)1, 4, 9 Lipkin S.M....Gruber S.B. (Cancer Prev Res (Phila) 2010)
    5. (TTA)n polymorphism in 3-hydroxy-3-methylglutaryl-coenzyme A and response to atorvastatin in coronary artery disease patients. (PubMed id 19067673)1, 4, 9 Noriega V....Prieto J.C. (amp 2009)
    6. Interaction between HMGCR and ABCA1 cholesterol-related genes modulates Alzheimer's disease risk. (PubMed id 19446537)1, 4, 9 RodrA-guez-RodrA-guez E....Combarros O. (Brain Res. 2009)
    7. A paucimorphic variant in the HMG-CoA reductase gene is associated with lipid-lowering response to statin treatment in diabetes: a GoDARTS study. (PubMed id 18815589)1, 4, 9 Donnelly L.A....Palmer C.N. (Pharmacogenet. Genomics 2008)
    8. Accelerated degradation of HMG CoA reductase mediated by binding of insig-1 to its sterol-sensing domain. (PubMed id 12535518)1, 2, 9 Sever N.... DeBose-Boyd R.A. (Mol. Cell 2003)
    9. Gene-gene and gene - clinical factors interaction in acute myocardial infarction: a new detailed risk chart. (PubMed id 20388088)1, 4, 9 Licastro F....Ferrari R. (Curr. Pharm. Des. 2010)
    10. A combination of proatherogenic single-nucleotide polymorphisms is associated with increased risk of coronary artery disease and myocardial infarction in Asian Indians. (PubMed id 19558216)1, 4, 9 Poduri A....Talwar K.K. (DNA Cell Biol. 2009)

    (in PubMed, OMIM, and NCBI Bookshelf)
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     ANDOR
    Aliases
    Disorders
    Free Text  

      Query String
    PubMed
    OMIM
    NCBI Bookshelf
      (Note: In FireFox, select the above section and copy using Ctrl-C)

    (According to Entrez Gene, HGNC, AceView, euGenes, Ensembl, miRBase, ECgene, Kegg, and/or H-InvDB)
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    Entrez Gene: 3156 HGNC: 5006 AceView: HMGCR Ensembl:ENSG00000113161 euGenes: HUgn3156
    ECgene: HMGCR Kegg: 3156 H-InvDB: HMGCR

    (According to HUGE)
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    (According to PharmGKB, ATLAS, HORDE, IMGT, LEIDEN, UniProtKB/Swiss-Prot, UniProtKB/TrEMBL, and/or others, e.g. Wikipedia and GeneReviews, via UniProtKB/Swiss-Prot)
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    NameDescription
    PharmGKB entry for HMGCR Pharmacogenomics, SNPs, Pathways

    (Patent information from GeneIP,
    Licensable technologies from WIS Yeda, Salk, Tufts,
    IP news from LifeMap Sciences, Inc.)
    About This Section

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    Patent Information for HMGCR gene:
    Search GeneIP for patents involving HMGCR

    GeneCards and IP:
    Japan Patent Office Licenses GeneCards     European Patent Office Licenses GeneCards     Improving the IP Search



    (Antibodies, recombinant proteins, and assays from EMD Millipore, R&D Systems, OriGene, QIAGEN, GenScript, Cell Signaling Technology, Novus Biologicals, Sino Biological, Enzo Life Sciences, Abcam, ProSpec, Cloud-Clone Corp., Thermo Fisher Scientific, eBioscience, antibodies-online, and/or others, Gene Editing from DNA2.0. Clones from OriGene, GenScript, Sino Biological, DNA2.0, SwitchGear Genomics, Vector BioLabs, Addgene, Cell lines from GenScript, and ESI BIO, Flow cytometery from eBioscience, PCR Arrays from QIAGEN, Drugs and/or compounds from EMD Millipore, Tocris Bioscience, Enzo Life Sciences, and/or ApexBio, In Situ Hybridization Assays from
    Advanced Cell Diagnostics, Animal models from genOway)
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    GeneCards Homepage - Last full update: 7 May 2014 - Incrementals: 9 May 2014 , 2 Jun 2014 , 26 Jun 2014 , 30 Jun 2014 , 21 Aug 2014 , 8 Sep 2014 , 7 Oct 2014

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