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    Eif2ak4 eukaryotic translation initiation factor 2 alpha kinase 4 [ Mus musculus (house mouse) ]

    Gene ID: 27103, updated on 27-Nov-2024

    Summary

    Official Symbol
    Eif2ak4provided by MGI
    Official Full Name
    eukaryotic translation initiation factor 2 alpha kinase 4provided by MGI
    Primary source
    MGI:MGI:1353427
    See related
    Ensembl:ENSMUSG00000005102 AllianceGenome:MGI:1353427
    Gene type
    protein coding
    RefSeq status
    VALIDATED
    Organism
    Mus musculus
    Lineage
    Eukaryota; Metazoa; Chordata; Craniata; Vertebrata; Euteleostomi; Mammalia; Eutheria; Euarchontoglires; Glires; Rodentia; Myomorpha; Muroidea; Muridae; Murinae; Mus; Mus
    Also known as
    GCN2; MGCN2; 2610011M03
    Summary
    Enables eukaryotic translation initiation factor 2alpha kinase activity and tRNA binding activity. Involved in several processes, including GCN2-mediated signaling; learning or memory; and regulation of translational initiation. Acts upstream of or within cellular response to starvation; endoplasmic reticulum unfolded protein response; and regulation of translation. Located in cytoplasm. Is expressed in several structures, including gut; hemolymphoid system gland; liver; lung; and ovary. Human ortholog(s) of this gene implicated in pulmonary venoocclusive disease 2. Orthologous to human EIF2AK4 (eukaryotic translation initiation factor 2 alpha kinase 4). [provided by Alliance of Genome Resources, Nov 2024]
    Expression
    Ubiquitous expression in ovary adult (RPKM 8.2), limb E14.5 (RPKM 7.9) and 28 other tissues See more
    Orthologs
    NEW
    Try the new Gene table
    Try the new Transcript table

    Genomic context

    See Eif2ak4 in Genome Data Viewer
    Location:
    2 E5; 2 59.4 cM
    Exon count:
    41
    Annotation release Status Assembly Chr Location
    RS_2024_02 current GRCm39 (GCF_000001635.27) 2 NC_000068.8 (118219065..118305715)
    108.20200622 previous assembly GRCm38.p6 (GCF_000001635.26) 2 NC_000068.7 (118388583..118475234)

    Chromosome 2 - NC_000068.8Genomic Context describing neighboring genes Neighboring gene fibrous sheath-interacting protein 1 Neighboring gene STARR-seq mESC enhancer starr_05456 Neighboring gene STARR-positive B cell enhancer ABC_E7849 Neighboring gene STARR-positive B cell enhancer mm9_chr2:118089004-118089305 Neighboring gene STARR-seq mESC enhancer starr_05457 Neighboring gene RIKEN cDNA 1700054M17 gene Neighboring gene G protein-coupled receptor 176 Neighboring gene STARR-seq mESC enhancer starr_05459 Neighboring gene STARR-positive B cell enhancer ABC_E1594 Neighboring gene STARR-positive B cell enhancer ABC_E9525 Neighboring gene aldo-keto reductase family 1, member B3 pseudogene Neighboring gene signal recognition particle 14

    Genomic regions, transcripts, and products

    Expression

    • 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

    Bibliography

    GeneRIFs: Gene References Into Functions

    What's a GeneRIF?

    Variation

    Alleles

    Alleles of this type are documented at Mouse Genome Informatics  (MGI)
    • Chemically induced (ENU) (1) 
    • Endonuclease-mediated (4) 
    • Targeted (3)  1 citation

    Interactions

    Products Interactant Other Gene Complex Source Pubs Description

    General gene information

    Markers

    Gene Ontology Provided by MGI

    Function Evidence Code Pubs
    enables 3-phosphoinositide-dependent protein kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables AMP-activated protein kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables ATP binding IEA
    Inferred from Electronic Annotation
    more info
     
    enables DNA-dependent protein kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables Rho-dependent protein serine/threonine kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables eukaryotic initiation factor eIF2 binding IC
    Inferred by Curator
    more info
    PubMed 
    enables eukaryotic translation initiation factor 2alpha kinase activity IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    enables eukaryotic translation initiation factor 2alpha kinase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables eukaryotic translation initiation factor 2alpha kinase activity IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    enables eukaryotic translation initiation factor 2alpha kinase activity IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    enables eukaryotic translation initiation factor 2alpha kinase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables histone H2AS1 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H2AS121 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H2AT120 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H2AXS139 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H2BS14 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H2BS36 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3S10 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3S28 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3S57 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3T11 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3T3 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3T45 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H3T6 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H4S1 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables protein binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables protein serine kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables protein serine/threonine kinase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables ribosomal protein S6 kinase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables tRNA binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    Process Evidence Code Pubs
    involved_in DNA damage checkpoint signaling IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in GCN2-mediated signaling IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in GCN2-mediated signaling IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in GCN2-mediated signaling ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in T cell activation involved in immune response IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in adaptive immune response IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in cellular response to UV IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in cellular response to amino acid starvation IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in cellular response to amino acid starvation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in cellular response to amino acid starvation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in cellular response to amino acid starvation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cellular response to cold ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cellular response to cold ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    acts_upstream_of_or_within cellular response to starvation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in chromatin remodeling IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in defense response to virus IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in eiF2alpha phosphorylation in response to endoplasmic reticulum stress ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in eiF2alpha phosphorylation in response to endoplasmic reticulum stress ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    acts_upstream_of_or_within endoplasmic reticulum unfolded protein response IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in learning IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in long-term memory IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in negative regulation by host of viral genome replication IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of CREB transcription factor activity IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in negative regulation of neuron differentiation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    acts_upstream_of_or_within negative regulation of translation IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    involved_in negative regulation of translational initiation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in negative regulation of translational initiation in response to stress IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in negative regulation of translational initiation in response to stress IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in neuron projection extension IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of adaptive immune response IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of defense response to virus by host IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of gene expression IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of long-term synaptic potentiation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of protein phosphorylation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of translational initiation in response to starvation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of translational initiation in response to starvation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in protein autophosphorylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in protein phosphorylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in protein phosphorylation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in regulation of cytoplasmic translational initiation in response to stress IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in regulation of feeding behavior IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in regulation of translational initiation IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    involved_in regulation of translational initiation ISS
    Inferred from Sequence or Structural Similarity
    more info
    PubMed 
    involved_in regulation of translational initiation by eIF2 alpha phosphorylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in regulation of translational initiation by eIF2 alpha phosphorylation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in regulation of translational initiation by eIF2 alpha phosphorylation ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within regulation of translational initiation in response to stress IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in response to endoplasmic reticulum stress IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    involved_in viral translation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    Component Evidence Code Pubs
    is_active_in cytoplasm IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    located_in cytoplasm IDA
    Inferred from Direct Assay
    more info
    PubMed 
    is_active_in cytosol IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    located_in cytosolic ribosome ISS
    Inferred from Sequence or Structural Similarity
    more info
    PubMed 
    is_active_in nucleus IBA
    Inferred from Biological aspect of Ancestor
    more info
     

    General protein information

    Preferred Names
    eIF-2-alpha kinase GCN2
    Names
    GCN2 eIF2alpha kinase
    GCN2-like protein
    NP_001171277.1
    NP_001342312.1
    NP_001408350.1
    NP_001408351.1
    NP_038747.2
    XP_006499689.1
    XP_006499690.1
    XP_017174398.1
    XP_036018145.1

    NCBI Reference Sequences (RefSeq)

    NEW Try the new Transcript table

    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_001177806.2NP_001171277.1  eIF-2-alpha kinase GCN2 isoform 2

      See identical proteins and their annotated locations for NP_001171277.1

      Status: VALIDATED

      Description
      Transcript Variant: This variant (2) differs in the 5' UTR and coding sequence and lacks an alternate internal exon in the 5' end compared to variant 1, that causes a frameshift. The resulting isoform (2) has a shorter and distinct N-terminus compared to isoform 1.
      Source sequence(s)
      AL929163
      Consensus CDS
      CCDS50669.1
      UniProtKB/TrEMBL
      A2AUM2
      Related
      ENSMUSP00000099586.4, ENSMUST00000102527.10
      Conserved Domains (4) summary
      PLN02972
      Location:9581378
      PLN02972; Histidyl-tRNA synthetase
      cd14012
      Location:187425
      PK_eIF2AK_GCN2_rpt1; Pseudokinase domain, repeat 1, of eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cd14046
      Location:470886
      STKc_EIF2AK4_GCN2_rpt2; Catalytic domain, repeat 2, of the Serine/Threonine kinase, eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cl00266
      Location:12841528
      HGTP_anticodon; HGTP anticodon binding domain, as found at the C-terminus of histidyl, glycyl, threonyl and prolyl tRNA synthetases, which are classified as a group of class II aminoacyl-tRNA synthetases (aaRS). In aaRSs, the anticodon binding domain is responsible for ...
    2. NM_001355383.2NP_001342312.1  eIF-2-alpha kinase GCN2 isoform 3

      Status: VALIDATED

      Description
      Transcript Variant: This variant (3) lacks three alternate internal exons in the 5' end compared to variant 1, that causes a frameshift. The resulting isoform (3) has a shorter and distinct N-terminus compared to isoform 1.
      Source sequence(s)
      AL929163
      UniProtKB/TrEMBL
      A2AUM2
      Conserved Domains (4) summary
      cd14012
      Location:109347
      PK_eIF2AK_GCN2_rpt1; Pseudokinase domain, repeat 1, of eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cd14046
      Location:392808
      STKc_EIF2AK4_GCN2_rpt2; Catalytic domain, repeat 2, of the Serine/Threonine kinase, eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cl00266
      Location:12061450
      HGTP_anticodon; HGTP anticodon binding domain, as found at the C-terminus of histidyl, glycyl, threonyl and prolyl tRNA synthetases, which are classified as a group of class II aminoacyl-tRNA synthetases (aaRS). In aaRSs, the anticodon binding domain is responsible for ...
      cl28413
      Location:8801300
      HisS; Histidyl-tRNA synthetase [Translation, ribosomal structure and biogenesis]
    3. NM_001421421.1NP_001408350.1  eIF-2-alpha kinase GCN2 isoform 2

      Status: VALIDATED

      Source sequence(s)
      AL929163
    4. NM_001421422.1NP_001408351.1  eIF-2-alpha kinase GCN2 isoform 4

      Status: VALIDATED

      Source sequence(s)
      AL929163
    5. NM_013719.4NP_038747.2  eIF-2-alpha kinase GCN2 isoform 1

      Status: VALIDATED

      Description
      Transcript Variant: This variant (1) represents the longest transcript and encodes the longest isoform (1).
      Source sequence(s)
      AL929163
      Consensus CDS
      CCDS16576.1
      UniProtKB/Swiss-Prot
      A2AUM3, A2AUM4, Q6GQT4, Q6ZPT5, Q8C5S0, Q8CIF5, Q9CT30, Q9CUV9, Q9ESB6, Q9ESB7, Q9ESB8, Q9QZ05
      UniProtKB/TrEMBL
      A2AUM0
      Related
      ENSMUSP00000005233.6, ENSMUST00000005233.12
      Conserved Domains (7) summary
      smart00220
      Location:5891000
      S_TKc; Serine/Threonine protein kinases, catalytic domain
      smart00591
      Location:26137
      RWD; domain in RING finger and WD repeat containing proteins and DEXDc-like helicases subfamily related to the UBCc domain
      COG0124
      Location:10491485
      HisS; Histidyl-tRNA synthetase [Translation, ribosomal structure and biogenesis]
      cd14012
      Location:299537
      PK_eIF2AK_GCN2_rpt1; Pseudokinase domain, repeat 1, of eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cd14046
      Location:582998
      STKc_EIF2AK4_GCN2_rpt2; Catalytic domain, repeat 2, of the Serine/Threonine kinase, eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cl00266
      Location:13941640
      HGTP_anticodon; HGTP anticodon binding domain, as found at the C-terminus of histidyl, glycyl, threonyl and prolyl tRNA synthetases, which are classified as a group of class II aminoacyl-tRNA synthetases (aaRS). In aaRSs, the anticodon binding domain is responsible for ...
      cl00268
      Location:10601384
      class_II_aaRS-like_core; Class II tRNA amino-acyl synthetase-like catalytic core domain. Class II amino acyl-tRNA synthetases (aaRS) share a common fold and generally attach an amino acid to the 3' OH of ribose of the appropriate tRNA. PheRS is an exception in that it attaches ...

    RefSeqs of Annotated Genomes: GCF_000001635.27-RS_2024_02

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

    Reference GRCm39 C57BL/6J

    Genomic

    1. NC_000068.8 Reference GRCm39 C57BL/6J

      Range
      118219065..118305715
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_036162252.1XP_036018145.1  eIF-2-alpha kinase GCN2 isoform X2

      UniProtKB/TrEMBL
      A2AUM1, A2AUM2
      Related
      ENSMUSP00000106498.2, ENSMUST00000110874.8
      Conserved Domains (5) summary
      smart00591
      Location:959
      RWD; domain in RING finger and WD repeat containing proteins and DEXDc-like helicases subfamily related to the UBCc domain
      PLN02972
      Location:9921412
      PLN02972; Histidyl-tRNA synthetase
      cd14012
      Location:221459
      PK_eIF2AK_GCN2_rpt1; Pseudokinase domain, repeat 1, of eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cd14046
      Location:504920
      STKc_EIF2AK4_GCN2_rpt2; Catalytic domain, repeat 2, of the Serine/Threonine kinase, eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cl00266
      Location:13181562
      HGTP_anticodon; HGTP anticodon binding domain, as found at the C-terminus of histidyl, glycyl, threonyl and prolyl tRNA synthetases, which are classified as a group of class II aminoacyl-tRNA synthetases (aaRS). In aaRSs, the anticodon binding domain is responsible for ...
    2. XM_017318909.3XP_017174398.1  eIF-2-alpha kinase GCN2 isoform X1

      UniProtKB/Swiss-Prot
      A2AUM3, A2AUM4, Q6GQT4, Q6ZPT5, Q8C5S0, Q8CIF5, Q9CT30, Q9CUV9, Q9ESB6, Q9ESB7, Q9ESB8, Q9QZ05
      Conserved Domains (5) summary
      smart00591
      Location:26137
      RWD; domain in RING finger and WD repeat containing proteins and DEXDc-like helicases subfamily related to the UBCc domain
      PLN02972
      Location:10431463
      PLN02972; Histidyl-tRNA synthetase
      cd14012
      Location:272510
      PK_eIF2AK_GCN2_rpt1; Pseudokinase domain, repeat 1, of eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cd14046
      Location:555971
      STKc_EIF2AK4_GCN2_rpt2; Catalytic domain, repeat 2, of the Serine/Threonine kinase, eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cl00266
      Location:13691613
      HGTP_anticodon; HGTP anticodon binding domain, as found at the C-terminus of histidyl, glycyl, threonyl and prolyl tRNA synthetases, which are classified as a group of class II aminoacyl-tRNA synthetases (aaRS). In aaRSs, the anticodon binding domain is responsible for ...
    3. XM_006499626.5XP_006499689.1  eIF-2-alpha kinase GCN2 isoform X4

      UniProtKB/TrEMBL
      A2AUM2
      Conserved Domains (4) summary
      PLN02972
      Location:9471367
      PLN02972; Histidyl-tRNA synthetase
      cd14012
      Location:176414
      PK_eIF2AK_GCN2_rpt1; Pseudokinase domain, repeat 1, of eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cd14046
      Location:459875
      STKc_EIF2AK4_GCN2_rpt2; Catalytic domain, repeat 2, of the Serine/Threonine kinase, eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cl00266
      Location:12731517
      HGTP_anticodon; HGTP anticodon binding domain, as found at the C-terminus of histidyl, glycyl, threonyl and prolyl tRNA synthetases, which are classified as a group of class II aminoacyl-tRNA synthetases (aaRS). In aaRSs, the anticodon binding domain is responsible for ...
    4. XM_006499627.5XP_006499690.1  eIF-2-alpha kinase GCN2 isoform X5

      UniProtKB/TrEMBL
      A2AUM2
      Related
      ENSMUSP00000106494.2, ENSMUST00000110870.8
      Conserved Domains (4) summary
      PLN02972
      Location:9201340
      PLN02972; Histidyl-tRNA synthetase
      cd14012
      Location:149387
      PK_eIF2AK_GCN2_rpt1; Pseudokinase domain, repeat 1, of eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cd14046
      Location:432848
      STKc_EIF2AK4_GCN2_rpt2; Catalytic domain, repeat 2, of the Serine/Threonine kinase, eukaryotic translation Initiation Factor 2-Alpha Kinase 4 or General Control Non-derepressible-2
      cl00266
      Location:12461490
      HGTP_anticodon; HGTP anticodon binding domain, as found at the C-terminus of histidyl, glycyl, threonyl and prolyl tRNA synthetases, which are classified as a group of class II aminoacyl-tRNA synthetases (aaRS). In aaRSs, the anticodon binding domain is responsible for ...