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    Hif1a hypoxia inducible factor 1, alpha subunit [ Mus musculus (house mouse) ]

    Gene ID: 15251, updated on 14-Nov-2017
    Official Symbol
    Hif1aprovided by MGI
    Official Full Name
    hypoxia inducible factor 1, alpha subunitprovided by MGI
    Primary source
    MGI:MGI:106918
    See related
    Ensembl:ENSMUSG00000021109 Vega:OTTMUSG00000019740
    Gene type
    protein coding
    RefSeq status
    REVIEWED
    Organism
    Mus musculus
    Lineage
    Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha; Muroidea; Muridae; Murinae; Mus; Mus
    Also known as
    MOP1; bHLHe78; AA959795; HIF1alpha; HIF1-alpha; HIF-1-alpha
    Summary
    This gene encodes the alpha subunit which, along with the beta subunit, forms a heterodimeric transcription factor that regulates the cellular and developmental response to reduced oxygen tension. The transcription factor has been shown to regulate genes involved in several biological processes, including erythropoiesis and angiogenesis which aid in increased delivery of oxygen to hypoxic regions. The transcription factor also plays a role in the induction of genes involved in cell proliferation and survival, energy metabolism, apoptosis, and glucose and iron metabolism. Alternative splicing results in multiple transcript variants encoding different isoforms. [provided by RefSeq, Sep 2015]
    Orthologs
    Location:
    12 C3; 12 31.99 cM
    Exon count:
    16
    Annotation release Status Assembly Chr Location
    106 current GRCm38.p4 (GCF_000001635.24) 12 NC_000078.6 (73901324..73947554)
    Build 37.2 previous assembly MGSCv37 (GCF_000001635.18) 12 NC_000078.5 (75008854..75048517)

    Chromosome 12 - NC_000078.6Genomic Context describing neighboring genes Neighboring gene protein kinase C, eta Neighboring gene predicted gene, 34189 Neighboring gene predicted gene 8075 Neighboring gene predicted gene 15283 Neighboring gene S100 calcium binding protein A11, pseudogene Neighboring gene small nuclear RNA activating complex, polypeptide 1

    • Project title: Mouse ENCODE transcriptome data
    • Description: RNA profiling data sets generated by the Mouse ENCODE project.
    • BioProject: PRJNA66167
    • Publication: PMID 25409824
    • Analysis date: n/a

    GeneRIFs: Gene References Into FunctionsWhat's a GeneRIF?

    Products Interactant Other Gene Complex Source Pubs Description

    Markers

    Homology

    Gene Ontology Provided by MGI

    Function Evidence Code Pubs
    DNA binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    DNA binding IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    DNA binding transcription factor activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    DNA binding transcription factor activity ISO
    Inferred from Sequence Orthology
    more info
    PubMed 
    E-box binding IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    Hsp90 protein binding ISO
    Inferred from Sequence Orthology
    more info
     
    RNA polymerase II transcription factor activity, sequence-specific DNA binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    RNA polymerase II transcription factor activity, sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    enzyme binding ISO
    Inferred from Sequence Orthology
    more info
     
    histone acetyltransferase binding ISO
    Inferred from Sequence Orthology
    more info
     
    histone deacetylase binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    nuclear hormone receptor binding ISO
    Inferred from Sequence Orthology
    more info
     
    p53 binding ISO
    Inferred from Sequence Orthology
    more info
     
    protein binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    protein complex binding ISO
    Inferred from Sequence Orthology
    more info
     
    protein dimerization activity IEA
    Inferred from Electronic Annotation
    more info
     
    protein domain specific binding ISO
    Inferred from Sequence Orthology
    more info
     
    protein heterodimerization activity ISO
    Inferred from Sequence Orthology
    more info
     
    protein kinase binding ISO
    Inferred from Sequence Orthology
    more info
     
    sequence-specific DNA binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    contributes_to sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    transcription factor activity, RNA polymerase II distal enhancer sequence-specific binding ISO
    Inferred from Sequence Orthology
    more info
     
    contributes_to transcription factor activity, RNA polymerase II distal enhancer sequence-specific binding ISO
    Inferred from Sequence Orthology
    more info
     
    transcription factor activity, RNA polymerase II transcription factor binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    transcription factor activity, transcription factor binding ISO
    Inferred from Sequence Orthology
    more info
     
    transcription factor binding ISO
    Inferred from Sequence Orthology
    more info
     
    transcriptional activator activity, RNA polymerase II proximal promoter sequence-specific DNA binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    transcriptional activator activity, RNA polymerase II transcription regulatory region sequence-specific DNA binding ISO
    Inferred from Sequence Orthology
    more info
     
    ubiquitin protein ligase binding ISO
    Inferred from Sequence Orthology
    more info
     
    Process Evidence Code Pubs
    B-1 B cell homeostasis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    angiogenesis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    axonal transport of mitochondrion ISO
    Inferred from Sequence Orthology
    more info
     
    blood vessel development IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    blood vessel morphogenesis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    camera-type eye morphogenesis IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    cardiac ventricle morphogenesis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    cartilage development IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    cell differentiation IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    cell differentiation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    cellular iron ion homeostasis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    cellular response to hypoxia IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    cellular response to hypoxia ISO
    Inferred from Sequence Orthology
    more info
     
    cerebral cortex development IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    collagen metabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    connective tissue replacement involved in inflammatory response wound healing IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    digestive tract morphogenesis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    dopaminergic neuron differentiation IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    dopaminergic neuron differentiation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    elastin metabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    embryonic hemopoiesis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    embryonic placenta development IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    embryonic placenta development IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    epithelial cell differentiation involved in mammary gland alveolus development IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    epithelial to mesenchymal transition IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    glucose homeostasis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    heart looping IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    hemoglobin biosynthetic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    hypoxia-inducible factor-1alpha signaling pathway IDA
    Inferred from Direct Assay
    more info
    PubMed 
    intestinal epithelial cell maturation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    iris morphogenesis IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    lactate metabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    lactation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    muscle cell cellular homeostasis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of TOR signaling IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of apoptotic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of apoptotic process ISO
    Inferred from Sequence Orthology
    more info
     
    negative regulation of bone mineralization IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of gene expression ISO
    Inferred from Sequence Orthology
    more info
     
    negative regulation of growth IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of mesenchymal cell apoptotic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of neuron apoptotic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of ossification IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of oxidative stress-induced neuron intrinsic apoptotic signaling pathway ISO
    Inferred from Sequence Orthology
    more info
     
    negative regulation of reactive oxygen species metabolic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of thymocyte apoptotic process IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    negative regulation of transcription from RNA polymerase II promoter ISO
    Inferred from Sequence Orthology
    more info
     
    negative regulation of vasoconstriction ISO
    Inferred from Sequence Orthology
    more info
     
    neural crest cell migration IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    neural fold elevation formation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    outflow tract morphogenesis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    oxygen homeostasis IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    oxygen homeostasis ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of angiogenesis ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of apoptotic process ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of autophagy IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    positive regulation of autophagy of mitochondrion IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    positive regulation of blood vessel endothelial cell migration ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of cell proliferation ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of epithelial cell migration IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    positive regulation of erythrocyte differentiation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    positive regulation of gene expression ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of gluconeogenesis ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of hormone biosynthetic process ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of insulin secretion involved in cellular response to glucose stimulus IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    positive regulation of macroautophagy IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    positive regulation of neuroblast proliferation IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    positive regulation of pri-miRNA transcription from RNA polymerase II promoter ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of receptor biosynthetic process ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of smooth muscle cell proliferation ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of transcription from RNA polymerase II promoter IDA
    Inferred from Direct Assay
    more info
    PubMed 
    positive regulation of transcription from RNA polymerase II promoter IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    positive regulation of transcription from RNA polymerase II promoter IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    positive regulation of transcription from RNA polymerase II promoter ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of transcription from RNA polymerase II promoter in response to hypoxia ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of transcription, DNA-templated ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of vascular endothelial growth factor production ISO
    Inferred from Sequence Orthology
    more info
     
    positive regulation of vascular endothelial growth factor receptor signaling pathway IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    regulation of aerobic respiration IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    regulation of catalytic activity IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    regulation of cell proliferation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    regulation of gene expression IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    regulation of gene expression ISO
    Inferred from Sequence Orthology
    more info
     
    regulation of glycolytic process IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    regulation of thymocyte apoptotic process IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    regulation of transcription from RNA polymerase II promoter in response to oxidative stress IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    regulation of transcription from RNA polymerase II promoter in response to oxidative stress ISO
    Inferred from Sequence Orthology
    more info
     
    regulation of transcription, DNA-templated IDA
    Inferred from Direct Assay
    more info
    PubMed 
    regulation of transcription, DNA-templated ISO
    Inferred from Sequence Orthology
    more info
    PubMed 
    regulation of transforming growth factor beta2 production ISO
    Inferred from Sequence Orthology
    more info
     
    response to fungicide ISO
    Inferred from Sequence Orthology
    more info
     
    response to hypoxia IDA
    Inferred from Direct Assay
    more info
    PubMed 
    response to hypoxia IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    response to hypoxia IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    response to hypoxia ISO
    Inferred from Sequence Orthology
    more info
    PubMed 
    response to muscle activity IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    retina vasculature development in camera-type eye IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    signal transduction ISO
    Inferred from Sequence Orthology
    more info
     
    transcription from RNA polymerase II promoter ISO
    Inferred from Sequence Orthology
    more info
     
    transcription, DNA-templated IEA
    Inferred from Electronic Annotation
    more info
     
    vascular endothelial growth factor production ISO
    Inferred from Sequence Orthology
    more info
     
    vasculature development IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    visual learning IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    Component Evidence Code Pubs
    RNA polymerase II transcription factor complex ISO
    Inferred from Sequence Orthology
    more info
     
    cytoplasm ISO
    Inferred from Sequence Orthology
    more info
     
    cytosol ISO
    Inferred from Sequence Orthology
    more info
     
    motile cilium IDA
    Inferred from Direct Assay
    more info
    PubMed 
    nuclear body ISO
    Inferred from Sequence Orthology
    more info
     
    nuclear speck IDA
    Inferred from Direct Assay
    more info
    PubMed 
    nucleoplasm ISO
    Inferred from Sequence Orthology
    more info
     
    nucleus IDA
    Inferred from Direct Assay
    more info
    PubMed 
    nucleus ISO
    Inferred from Sequence Orthology
    more info
     
    protein complex ISO
    Inferred from Sequence Orthology
    more info
     
    transcription factor complex ISO
    Inferred from Sequence Orthology
    more info
     
    Preferred Names
    hypoxia-inducible factor 1-alpha
    Names
    ARNT-interacting protein

    RefSeqs maintained independently of Annotated Genomes

    These reference sequences exist independently of genome builds. Explain

    These reference sequences are curated independently of the genome annotation cycle, so their versions may not match the RefSeq versions in the current genome build. Identify version mismatches by comparing the version of the RefSeq in this section to the one reported in Genomic regions, transcripts, and products above.

    mRNA and Protein(s)

    1. NM_001313919.1NP_001300848.1  hypoxia-inducible factor 1-alpha isoform 1

      Status: REVIEWED

      Description
      Transcript Variant: This variant (1) represents the longest transcript and encodes the longest isoform (1).
      Source sequence(s)
      AC124712, AK034087
      UniProtKB/Swiss-Prot
      Q61221
      Conserved Domains (6) summary
      smart00091
      Location:94149
      PAS; PAS domain
      cd00083
      Location:1669
      HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...
      cd00130
      Location:240339
      PAS; PAS domain; PAS motifs appear in archaea, eubacteria and eukarya. Probably the most surprising identification of a PAS domain was that in EAG-like K+-channels. PAS domains have been found to bind ligands, and to act as sensors for light and oxygen in ...
      pfam08447
      Location:255340
      PAS_3; PAS fold
      pfam08778
      Location:800836
      HIF-1a_CTAD; HIF-1 alpha C terminal transactivation domain
      pfam11413
      Location:565594
      HIF-1; Hypoxia-inducible factor-1
    2. NM_001313920.1NP_001300849.1  hypoxia-inducible factor 1-alpha isoform 3

      Status: REVIEWED

      Description
      Transcript Variant: This variant (3) has an alternate 5' structure, which results in the use of a downstream start codon, and uses an alternate in-frame splice site in the 3' coding region compared to variant 1. The encoded isoform (3) has a shorter N-terminus compared to isoform 1.
      Source sequence(s)
      AC124712
      Consensus CDS
      CCDS83968.1
      UniProtKB/Swiss-Prot
      Q61221
      UniProtKB/TrEMBL
      A0A0R4J1E9
      Related
      ENSMUSP00000106088.1, OTTMUSP00000021323, ENSMUST00000110461.7, OTTMUST00000047241
      Conserved Domains (6) summary
      smart00091
      Location:81136
      PAS; PAS domain
      cd00083
      Location:356
      HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...
      cd00130
      Location:227326
      PAS; PAS domain; PAS motifs appear in archaea, eubacteria and eukarya. Probably the most surprising identification of a PAS domain was that in EAG-like K+-channels. PAS domains have been found to bind ligands, and to act as sensors for light and oxygen in ...
      pfam08447
      Location:242327
      PAS_3; PAS fold
      pfam08778
      Location:773809
      HIF-1a_CTAD; HIF-1 alpha C terminal transactivation domain
      pfam11413
      Location:538567
      HIF-1; Hypoxia-inducible factor-1
    3. NM_010431.2NP_034561.2  hypoxia-inducible factor 1-alpha isoform 2

      See identical proteins and their annotated locations for NP_034561.2

      Status: REVIEWED

      Description
      Transcript Variant: This variant (2) uses an alternate in-frame splice site in the 5' coding region, compared to variant 1. The encoded isoform (2) is shorter than isoform 1.
      Source sequence(s)
      AC124712, AK034087, AK150367, CX241289
      Consensus CDS
      CCDS25977.1
      UniProtKB/Swiss-Prot
      Q61221
      UniProtKB/TrEMBL
      Q3UCW2
      Related
      ENSMUSP00000021530.7, OTTMUSP00000021321, ENSMUST00000021530.7, OTTMUST00000047239
      Conserved Domains (6) summary
      smart00091
      Location:93148
      PAS; PAS domain
      cd00083
      Location:1568
      HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...
      cd00130
      Location:239338
      PAS; PAS domain; PAS motifs appear in archaea, eubacteria and eukarya. Probably the most surprising identification of a PAS domain was that in EAG-like K+-channels. PAS domains have been found to bind ligands, and to act as sensors for light and oxygen in ...
      pfam08447
      Location:254339
      PAS_3; PAS fold
      pfam08778
      Location:799835
      HIF-1a_CTAD; HIF-1 alpha C terminal transactivation domain
      pfam11413
      Location:564593
      HIF-1; Hypoxia-inducible factor-1

    RefSeqs of Annotated Genomes: Mus musculus Annotation Release 106 details...Open this link in a new tab

    The following sections contain reference sequences that belong to a specific genome build. Explain

    Reference GRCm38.p4 C57BL/6J

    Genomic

    1. NC_000078.6 Reference GRCm38.p4 C57BL/6J

      Range
      73901324..73947554
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_006515478.3XP_006515541.1  hypoxia-inducible factor 1-alpha isoform X1

      Conserved Domains (6) summary
      smart00091
      Location:94149
      PAS; PAS domain
      cd00083
      Location:1669
      HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...
      cd00130
      Location:240339
      PAS; PAS domain; PAS motifs appear in archaea, eubacteria and eukarya. Probably the most surprising identification of a PAS domain was that in EAG-like K+-channels. PAS domains have been found to bind ligands, and to act as sensors for light and oxygen in ...
      pfam08447
      Location:255340
      PAS_3; PAS fold
      pfam08778
      Location:786822
      HIF-1a_CTAD; HIF-1 alpha C terminal transactivation domain
      pfam11413
      Location:551580
      HIF-1; Hypoxia-inducible factor-1
    2. XM_017314961.1XP_017170450.1  hypoxia-inducible factor 1-alpha isoform X2

      UniProtKB/Swiss-Prot
      Q61221
      Conserved Domains (6) summary
      smart00091
      Location:93148
      PAS; PAS domain
      cd00083
      Location:1568
      HLH; Helix-loop-helix domain, found in specific DNA- binding proteins that act as transcription factors; 60-100 amino acids long. A DNA-binding basic region is followed by two alpha-helices separated by a variable loop region; HLH forms homo- and heterodimers, ...
      cd00130
      Location:239338
      PAS; PAS domain; PAS motifs appear in archaea, eubacteria and eukarya. Probably the most surprising identification of a PAS domain was that in EAG-like K+-channels. PAS domains have been found to bind ligands, and to act as sensors for light and oxygen in ...
      pfam08447
      Location:254339
      PAS_3; PAS fold
      pfam08778
      Location:785821
      HIF-1a_CTAD; HIF-1 alpha C terminal transactivation domain
      pfam11413
      Location:550579
      HIF-1; Hypoxia-inducible factor-1
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