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ALOX5 (Arachidonate 5-Lipoxygenase).

Written2006-07Sreeparna Banerjee, Seda Tuncay
Department of Biology, Office: Z-16/Lab: B-59, Middle East Technical University, 06531 Ankara, Turkey

(Note : for Links provided by Atlas : click)

Identity

Alias_symbol (synonym)5-LOX
Other alias5-LO
EC 1.13.11.34
leukotriene A4 synthase
5LPG
LOG5
HGNC (Hugo) ALOX5
LocusID (NCBI) 240
Atlas_Id 42985
Location 10q11.21  [Link to chromosome band 10q11]
Location_base_pair Starts at 45374166 and ends at 45446119 bp from pter ( according to hg19-Feb_2009)  [Mapping ALOX5.png]
Local_order Genes flanking ALOX5, in centromere to telomere direction on 10q11, are:
  • OR6D1P 10q11.21 olfactory receptor, family 6, subfamily D, member 1 pseudogene.
  • LOC643413 10q11.21 hypothetical protein LOC643413.
  • OR13A1 10q11.21 olfactory receptor, family 13, subfamily A, member 1.
  • ALOX5 10q11.2 arachidonate 5-lipoxygenase.
  • MARCH8 10q11.21 membrane-associated ring finger (C3HC4) 8.
  • LOC653306 10q11.21 similar to membrane-associated ring finger (C3HC4) 8.
  • ANUBL1 10q11.21 AN1, ubiquitin-like, homolog (Xenopus laevis).
    		•
    Fusion genes
    (updated 2016)    		ALOX5 (10q11.21) / ABCA13 (7p12.3)

    DNA/RNA

     
      Diagram of the ALOX5 gene. Exons are represented by purple boxes (in scale). Exons 1 to 14 are from the 5' to 3' direction.
    Description ALOX5 gene spans a region of 71,88 kb and has 14 exons, the sizes being 192, 199, 82, 123, 107, 173, 147, 204, 87, 179, 122, 101, 171 and 606 bps. ALOX5 gene has 5 CpG islands and 3' end of the gene for cellular modulator of immune recognition (c-MIR).
    Transcription ALOX5 gene promoter (H. sapiens) lacks the TATA box and has eight GC-boxes within 180 bp from the major transcription initiation site (at-65 in relation to ATG), five of which are in tandem (-176 to - 147). Consensus-binding sites for the transcription factor serum protein 1 (SP1), and early growth-response protein 1(EGR-1) exists in this region. A Vitamin D receptor binding site has been located in a positive regulatory region (-779 to -229) of the ALOX5 promoter. Several other consensus-binding sites for transcription factors such as GATA, glucocorticoid receptors and NFKB also exist. DNA methylation and histone deacetylase are also strongly involved in ALOX5 expression.
    Pseudogene No pseudogenes have been reported for ALOX5.

    Protein

    Note The ALOX5 gene encodes a member of the lipoxygenase gene family, 5-LOX, which catalyzes the synthesis of leukotrienes (LT) from arachidonic acid. Leukotrienes are responsible for a series of inflammatory and allergic conditions. 5-LOX is also unique in requiring the 5-LOX activating protein (FLAP), a nuclear trans-membrane protein that plays an essential role in the transfer of arachidonic acid to 5-LOX. FLAP can also bind to MK-886, a compound that blocks LT biosynthesis.
    Description 5-LOX is a 77.9 kDa protein consisting of 673 amino acids. The enzyme requires calcium, iron and ATP as cofactors. The enzyme activity is also stimulated by the presence of microsomal membranes and trace amounts of lipid hydroperoxides. The protein has a catalytic domain and a regulatory domain. The regulatory domain, which controls leukotriene synthesis and binds calcium, nucleotides and phospholipids also has a PLAT (Polycystin-1, Lipoxygenase, alpha-Toxin) domain.
    Expression 5-LOX protein is expressed in bone marrow derived cells such as monocytes/macrophages, mast cells, B-lymphocytes, polymorphonuclear leukocytes, dendritic cells and foam cells of human atherosclerotic tissues, as well as spleen, thymus brain, spinal cord, skeletal muscle, pancreas, prostate, kidney and lung in humans.
    Localisation Subcellular location of 5-LOX protein is the cytoplasm or nucleoplasm. 5-LOX is largely cytosolic in resting peritoneal macrophages, monocytes, neutrophils, monocytes and eosinophils. By contrast, alveolar macrophages and mast cells contain cytosolic and intranuclear fractions of the enzyme. Leukotriene synthesis capacity is determined by a calcium independent nuclear import of 5-lipoxygenase. Three nuclear localization sequence (NLS) exist, Leu-111 to Asp-121; Asp-156 to Asp-166 and Val-514 to Leu-535.
    Function 5-LOX, a monomeric enzyme, catalyzes the conversion of arachidonic acid to 5(S)-hydroperoxy-6-trans-8, 11, 14-cis-eicosatetraenoic acid (5(S)-HETE), and further dehydration to the allylic epoxide 5(S)-trans-7,9-trans-11,14-cis-eicosatetrenoic acid (leukotriene A4). The LTA4 intermediate is then converted to LTB4 by LTA4 hydrolase. LTB4 attracts leukocytes and are important for the inflammatory response.

    5-LOX migrates to the nuclear membrane upon cellular activation leading to LTB4 biosynthesis. This function depends on calcium dependent binding of the N-terminal C2 domain of 5-LOX to phospholipids resulting in the release of fatty acid substrates for enzyme action.

    Phosphorylation of 5-LOX on Ser-271 by MAPK-activating protein (MAPKAP) kinase 2, Ser-663 by extracellular signal-regulated kinases (ERK-2) and Ser-523 by protein kinase A (PKA) catalytic subunit has been shown to stimulate 5-LOX activity.

    In addition, overexpression of 5-LOX was shown to promote senescence-like growth arrest in human and mouse embryo fibroblasts via a p53/p21-dependent pathway, by regulating reactive oxygen species production, independent of telomerase activity. Thus, a senescence-like growth arrest may be of significance in the pathogenesis of 5-LOX-associated disorders.

    Homology C. familiaris: LOC477753, similar to Arachidonate 5-lipoxygenase
    R. norvegicus: ALOX5, arachidonate 5-lipoxygenase
    M. musculus: ALOX5, arachidonate 5-lipoxygenase
    A. thaliana: AT3G22400 iron ion binding / lipoxygenase
    O. sativa: OSJNBb0017F17.2, putative lipoxygenase

    Mutations

    Note A family of mutations in the G+C-rich transcription factor binding region of ALOX5 has been identified in which several Sp1 and Egr-1 binding motifs are altered in the region of 176 to 147 bp upstream from the ATG translation start site. These mutations alter transcription factor binding and may play a role in 5-LOX gene expression in vivo. A haplotype containing polymorphisms in a negative regulatory region of the ALOX5 promoter (G-1752A and G-1699A) may influence colon cancer risk in Caucasians. In addition, the genetic variant of tandem repeat (GGGCGG; Sp1-binding motif) in ALOX5 promoter in group of Korean aspirin intolerant asthma patients has been associated with the severity of airway hyper-responsiveness.

    Implicated in

    Note
      
    Entity Esophageal cancer
    Disease Immunohistochemistry analyses of 5-LOX expression in 161 esophageal tissue indicated that the enzyme was expressed in 79% (127/161) of cancer tissues but in only 13% (4/32) of normal esophageal mucosa. 5-LOX was also expressed in 8 esophageal cancer cell lines examined. In addition, 5-LOX inhibitors AA861 and REV5901 increased cell viability and apoptosis in the esophageal cancer cell lines.
      
      
    Entity Pancreatic cancer
    Disease 5-LOX expression is upregulated human pancreatic cancer cells. The 5-LOX metabolite 5(S)-HETE was shown to stimulate proliferation, as well as the proliferation of the mitogenic intracellular tyrosine kinases, MEK/ERK and PI3 kinase/AKT.
      
      
    Entity Colorectal cancer
    Disease Exposure to cigarette smoke extract (CSE) was shown to enhance 5-LOX protein expression in the inflammation-associated colonic adenomas. The effects of CSE on colon cancer cells were mediated by 5-LOX DNA demethylation. In addition, an up-regulation of matrix metalloproteinase-2 (MMP-2) and vascular endothelial growth factor (VEGF), key angiogenic factors for tumorigenesis, were also observed. These effects were reversed by treating the colon cancer cells with dual 5-LOX and COX-2 inhibitors.
      
      
    Entity Atherosclerosis
    Disease 5-LOX, known to generate proinflammatory LTs, is highly expressed in the arterial walls of atherosclerotic patients, with the number of enzyme expressing lesion leukocytes increasing during disease progression. All constituents of the 5-LOX pathway are significantly expressed in human diseased arteries, thereby supporting a model of atherogenesis, whereby 5-LOX pathway dependent inflammatory circuits composed of leukocytes, smooth muscle cells and endothelial cells evolve within blood vessels during late stages of lesion development.
      
      
    Entity Asthma
    Disease LTs and their receptors play an important role in the pathogenesis of asthma. Th2 cytokines, interleukins-4 and -13 can upregulate cysteinyl leukotriene 1 receptor expression. In addition, cysteinyl LTs favour an allergic phenotype by upregulating type 2 cytokine expression and decreasing type 1 cytokine expression. Polymorphisms of the 5-LOX promoter have also been associated with the development of asthma.
      
      
    Entity Immune response and tissue homeostasis
    Note The products of the ALOX5 pathway, particularly LTs, are lipid messengers that act on the immune response system and tissue homeostasis. Their abnormal production can induce several diseases such as asthma, inflammation, atherosclerosis, basophilic leukaemia, oedema, exercise-induced asthma, anaphylaxis, psoriasis, bronchial spasms and allergic rhinitis.
    Oncogenesis Alterations in the 5-LOX pathway can result in the aberrant formation of its products, hydroxyeicosatetraenoic acids or leukotrienes. This can, in turn, increase cellular proliferation and survival and suppress apoptosis of human cells and thereby play a significant role in human carcinogenesis.
      

    Bibliography

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    Multiple signal pathways are involved in the mitogenic effect of 5(S)-HETE in human pancreatic cancer.
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    GC-rich sequences in the 5-lipoxygenase gene promoter are required for expression in Mono Mac 6 cells, characterization of a novel Sp1 binding site.
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    PMID 15308583
     
    Stress-induced nuclear export of 5-lipoxygenase.
    Hanaka H, Shimizu T, Izumi T
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    PMID 16165096
     
    Increased 5-lipoxygenase expression and induction of apoptosis by its inhibitors in esophageal cancer: a potential target for prevention.
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    Naturally occurring mutations in the human 5-lipoxygenase gene promoter that modify transcription factor binding and reporter gene transcription.
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    Mutations in the human 5-lipoxygenase gene.
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    Clinical reviews in allergy & immunology. 1999 ; 17 (1-2) : 59-69.
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    Polymorphism of tandem repeat in promoter of 5-lipoxygenase in ASA-intolerant asthma: a positive association with airway hyperresponsiveness.
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    Allergy. 2005 ; 60 (6) : 760-765.
    PMID 15876305
     
    Trichostatin A and structurally related histone deacetylase inhibitors induce 5-lipoxygenase promoter activity.
    Klan N, Seuter S, Schnur N, Jung M, Steinhilber D
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    Biochimica et biophysica acta. 2005 ; 1736 (1) : 30-37.
    PMID 16081317
     
    Protein kinase A inhibits leukotriene synthesis by phosphorylation of 5-lipoxygenase on serine 523.
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    PMID 15280375
     
    Multiple nuclear localization sequences allow modulation of 5-lipoxygenase nuclear import.
    Luo M, Pang CW, Gerken AE, Brock TG
    Traffic (Copenhagen, Denmark). 2004 ; 5 (11) : 847-854.
    PMID 15479450
     
    5-lipoxygenase and FLAP.
    Peters-Golden M, Brock TG
    Prostaglandins, leukotrienes, and essential fatty acids. 2003 ; 69 (2-3) : 99-109.
    PMID 12895592
     
    5-Lipoxygenase-activating protein homodimer in human neutrophils: evidence for a role in leukotriene biosynthesis.
    Plante H, Picard S, Mancini J, Borgeat P
    The Biochemical journal. 2006 ; 393 (Pt 1) : 211-218.
    PMID 16144515
     
    Expression of 5-lipoxygenase and leukotriene A4 hydrolase in human atherosclerotic lesions correlates with symptoms of plaque instability.
    Qiu H, Gabrielsen A, Agardh HE, Wan M, Wetterholm A, Wong CH, Hedin U, Swedenborg J, Hansson GK, Samuelsson B, Paulsson-Berne G, Haeggström JZ
    Proceedings of the National Academy of Sciences of the United States of America. 2006 ; 103 (21) : 8161-8166.
    PMID 16698924
     
    Regulation of 5-lipoxygenase enzyme activity.
    Rådmark O, Samuelsson B
    Biochemical and biophysical research communications. 2005 ; 338 (1) : 102-110.
    PMID 16122704
     
    The discovery of the leukotrienes.
    Samuelsson B
    American journal of respiratory and critical care medicine. 2000 ; 161 (2 Pt 2) : S2-S6.
    PMID 10673217
     
    Egr-1 and Sp1 interact functionally with the 5-lipoxygenase promoter and its naturally occurring mutants.
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    American journal of respiratory cell and molecular biology. 1998 ; 19 (2) : 316-323.
    PMID 9698605
     
    Analysis of the 5-lipoxygenase promoter and characterization of a vitamin D receptor binding site.
    Sorg BL, Klan N, Seuter S, Dishart D, Rådmark O, Habenicht A, Carlberg C, Werz O, Steinhilber D
    Biochimica et biophysica acta. 2006 ; 1761 (7) : 686-697.
    PMID 16750418
     
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    DNA methylation regulates 5-lipoxygenase promoter activity.
    Uhl J, Klan N, Rose M, Entian KD, Werz O, Steinhilber D
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    PMID 12751760
     
    Extracellular signal-regulated kinases phosphorylate 5-lipoxygenase and stimulate 5-lipoxygenase product formation in leukocytes.
    Werz O, Bürkert E, Fischer L, Szellas D, Dishart D, Samuelsson B, Rådmark O, Steinhilber D
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    Werz O, Szellas D, Steinhilber D, Rådmark O
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    Contributory role of 5-lipoxygenase and its association with angiogenesis in the promotion of inflammation-associated colonic tumorigenesis by cigarette smoking.
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    Citation

    This paper should be referenced as such :
    Banerjee, S ; Tuncay, S
    ALOX5 (arachidonate 5-lipoxygenase)
    Atlas Genet Cytogenet Oncol Haematol. 2006;10(4):243-246.
    Free journal version : [ pdf ]   [ DOI ]
    On line version : http://AtlasGeneticsOncology.org/Genes/ALOX5ID42985ch10q11.html

    External links

    Nomenclature
    HGNC (Hugo)ALOX5   435
    Cards
    AtlasALOX5ID42985ch10q11
    Entrez_Gene (NCBI)ALOX5  240  arachidonate 5-lipoxygenase
    Aliases5-LO; 5-LOX; 5LPG; LOG5
    GeneCards (Weizmann)ALOX5
    Ensembl hg19 (Hinxton)ENSG00000012779 [Gene_View]
    Ensembl hg38 (Hinxton)ENSG00000012779 [Gene_View]  chr10:45374166-45446119 [Contig_View]  ALOX5 [Vega]
    ICGC DataPortalENSG00000012779
    TCGA cBioPortalALOX5
    AceView (NCBI)ALOX5
    Genatlas (Paris)ALOX5
    WikiGenes240
    SOURCE (Princeton)ALOX5
    Genetics Home Reference (NIH)ALOX5
    Genomic and cartography
    GoldenPath hg38 (UCSC)ALOX5  -     chr10:45374166-45446119 +  10q11.21   [Description]    (hg38-Dec_2013)
    GoldenPath hg19 (UCSC)ALOX5  -     10q11.21   [Description]    (hg19-Feb_2009)
    EnsemblALOX5 - 10q11.21 [CytoView hg19]  ALOX5 - 10q11.21 [CytoView hg38]
    Mapping of homologs : NCBIALOX5 [Mapview hg19]  ALOX5 [Mapview hg38]
    OMIM152390   600807   
    Gene and transcription
    Genbank (Entrez)AB208946 AK225603 AK315223 AW403395 BC034464
    RefSeq transcript (Entrez)NM_000698 NM_001256153 NM_001256154 NM_001320861 NM_001320862
    RefSeq genomic (Entrez)
    Consensus coding sequences : CCDS (NCBI)ALOX5
    Cluster EST : UnigeneHs.89499 [ NCBI ]
    CGAP (NCI)Hs.89499
    Alternative Splicing GalleryENSG00000012779
    Gene ExpressionALOX5 [ NCBI-GEO ]   ALOX5 [ EBI - ARRAY_EXPRESS ]   ALOX5 [ SEEK ]   ALOX5 [ MEM ]
    Gene Expression Viewer (FireBrowse)ALOX5 [ Firebrowse - Broad ]
    SOURCE (Princeton)Expression in : [Datasets]   [Normal Tissue Atlas]  [carcinoma Classsification]  [NCI60]
    GenevisibleExpression in : [tissues]  [cell-lines]  [cancer]  [perturbations]  
    BioGPS (Tissue expression)240
    GTEX Portal (Tissue expression)ALOX5
    Protein : pattern, domain, 3D structure
    UniProt/SwissProtP09917   [function]  [subcellular_location]  [family_and_domains]  [pathology_and_biotech]  [ptm_processing]  [expression]  [interaction]
    NextProtP09917  [Sequence]  [Exons]  [Medical]  [Publications]
    With graphics : InterProP09917
    Splice isoforms : SwissVarP09917
    PhosPhoSitePlusP09917
    Domaine pattern : Prosite (Expaxy)LIPOXYGENASE_1 (PS00711)    LIPOXYGENASE_2 (PS00081)    LIPOXYGENASE_3 (PS51393)    PLAT (PS50095)   
    Domains : Interpro (EBI)LipOase    LipOase_C    LipOase_CS    LipOase_Fe_BS    LipOase_mml    PLAT/LH2_dom   
    Domain families : Pfam (Sanger)Lipoxygenase (PF00305)    PLAT (PF01477)   
    Domain families : Pfam (NCBI)pfam00305    pfam01477   
    Domain families : Smart (EMBL)LH2 (SM00308)  
    Conserved Domain (NCBI)ALOX5
    DMDM Disease mutations240
    Blocks (Seattle)ALOX5
    PDB (SRS)2ABV    3O8Y    3V92    3V98    3V99   
    PDB (PDBSum)2ABV    3O8Y    3V92    3V98    3V99   
    PDB (IMB)2ABV    3O8Y    3V92    3V98    3V99   
    PDB (RSDB)2ABV    3O8Y    3V92    3V98    3V99   
    Structural Biology KnowledgeBase2ABV    3O8Y    3V92    3V98    3V99   
    SCOP (Structural Classification of Proteins)2ABV    3O8Y    3V92    3V98    3V99   
    CATH (Classification of proteins structures)2ABV    3O8Y    3V92    3V98    3V99   
    SuperfamilyP09917
    Human Protein AtlasENSG00000012779
    Peptide AtlasP09917
    HPRD01065
    IPIIPI00218916   IPI01012189   IPI00418599   
    Protein Interaction databases
    DIP (DOE-UCLA)P09917
    IntAct (EBI)P09917
    FunCoupENSG00000012779
    BioGRIDALOX5
    STRING (EMBL)ALOX5
    ZODIACALOX5
    Ontologies - Pathways
    QuickGOP09917
    Ontology : AmiGOleukotriene production involved in inflammatory response  arachidonate 5-lipoxygenase activity  iron ion binding  protein binding  extracellular region  extracellular space  nuclear envelope  nuclear envelope  nuclear envelope lumen  nucleoplasm  cytosol  cytosol  leukotriene metabolic process  nuclear matrix  leukotriene biosynthetic process  lipoxygenase pathway  nuclear membrane  secretory granule lumen  neutrophil degranulation  oxidation-reduction process  ficolin-1-rich granule lumen  lipoxin metabolic process  
    Ontology : EGO-EBIleukotriene production involved in inflammatory response  arachidonate 5-lipoxygenase activity  iron ion binding  protein binding  extracellular region  extracellular space  nuclear envelope  nuclear envelope  nuclear envelope lumen  nucleoplasm  cytosol  cytosol  leukotriene metabolic process  nuclear matrix  leukotriene biosynthetic process  lipoxygenase pathway  nuclear membrane  secretory granule lumen  neutrophil degranulation  oxidation-reduction process  ficolin-1-rich granule lumen  lipoxin metabolic process  
    Pathways : BIOCARTAEicosanoid Metabolism [Genes]   
    Pathways : KEGGArachidonic acid metabolism    Serotonergic synapse    Ovarian steroidogenesis    Toxoplasmosis   
    REACTOMEP09917 [protein]
    REACTOME PathwaysR-HSA-6798695 [pathway]   
    NDEx NetworkALOX5
    Atlas of Cancer Signalling NetworkALOX5
    Wikipedia pathwaysALOX5
    Orthology - Evolution
    OrthoDB240
    GeneTree (enSembl)ENSG00000012779
    Phylogenetic Trees/Animal Genes : TreeFamALOX5
    HOVERGENP09917
    HOGENOMP09917
    Homologs : HomoloGeneALOX5
    Homology/Alignments : Family Browser (UCSC)ALOX5
    Gene fusions - Rearrangements
    Polymorphisms : SNP and Copy number variants
    NCBI Variation ViewerALOX5 [hg38]
    dbSNP Single Nucleotide Polymorphism (NCBI)ALOX5
    dbVarALOX5
    ClinVarALOX5
    1000_GenomesALOX5 
    Exome Variant ServerALOX5
    ExAC (Exome Aggregation Consortium)ALOX5 (select the gene name)
    Genetic variants : HAPMAP240
    Genomic Variants (DGV)ALOX5 [DGVbeta]
    DECIPHERALOX5 [patients]   [syndromes]   [variants]   [genes]  
    CONAN: Copy Number AnalysisALOX5 
    Mutations
    ICGC Data PortalALOX5 
    TCGA Data PortalALOX5 
    Broad Tumor PortalALOX5
    OASIS PortalALOX5 [ Somatic mutations - Copy number]
    Somatic Mutations in Cancer : COSMICALOX5  [overview]  [genome browser]  [tissue]  [distribution]  
    Mutations and Diseases : HGMDALOX5
    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
    LOVD (Leiden Open Variation Database)TBsLVD Tuberculosis susceptibility Locus Variation Database
    BioMutasearch ALOX5
    DgiDB (Drug Gene Interaction Database)ALOX5
    DoCM (Curated mutations)ALOX5 (select the gene name)
    CIViC (Clinical Interpretations of Variants in Cancer)ALOX5 (select a term)
    intoGenALOX5
    NCG5 (London)ALOX5
    Cancer3DALOX5(select the gene name)
    Impact of mutations[PolyPhen2] [SIFT Human Coding SNP] [Buck Institute : MutDB] [Mutation Assessor] [Mutanalyser]
    Diseases
    OMIM152390    600807   
    Orphanet
    MedgenALOX5
    Genetic Testing Registry ALOX5
    NextProtP09917 [Medical]
    TSGene240
    GENETestsALOX5
    Target ValidationALOX5
    Huge Navigator ALOX5 [HugePedia]
    snp3D : Map Gene to Disease240
    BioCentury BCIQALOX5
    ClinGenALOX5
    Clinical trials, drugs, therapy
    Chemical/Protein Interactions : CTD240
    Chemical/Pharm GKB GenePA46
    Clinical trialALOX5
    Miscellaneous
    canSAR (ICR)ALOX5 (select the gene name)
    Probes
    Litterature
    PubMed260 Pubmed reference(s) in Entrez
    GeneRIFsGene References Into Functions (Entrez)
    CoreMineALOX5
    EVEXALOX5
    GoPubMedALOX5
    iHOPALOX5
    REVIEW articlesautomatic search in PubMed
    Last year publicationsautomatic search in PubMed

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