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    Kat5 K(lysine) acetyltransferase 5 [ Mus musculus (house mouse) ]

    Gene ID: 81601, updated on 2-Nov-2024

    Summary

    Official Symbol
    Kat5provided by MGI
    Official Full Name
    K(lysine) acetyltransferase 5provided by MGI
    Primary source
    MGI:MGI:1932051
    See related
    Ensembl:ENSMUSG00000024926 AllianceGenome:MGI:1932051
    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
    PLIP; CPLA2; Tip55; Tip60; Htatip; Htatip1
    Summary
    Enables chromatin binding activity; histone acetyltransferase activity; and transcription coactivator activity. Involved in several processes, including nervous system development; positive regulation of regulatory T cell differentiation; and sperm DNA condensation. Acts upstream of or within positive regulation of transcription by RNA polymerase II. Part of NuA4 histone acetyltransferase complex and transcription regulator complex. Is expressed in several structures, including central nervous system; eye; genitourinary system; hemolymphoid system gland; and limb. Orthologous to human KAT5 (lysine acetyltransferase 5). [provided by Alliance of Genome Resources, Nov 2024]
    Expression
    Ubiquitous expression in CNS E14 (RPKM 21.5), testis adult (RPKM 19.9) and 28 other tissues See more
    Orthologs
    NEW
    Try the new Gene table
    Try the new Transcript table

    Genomic context

    See Kat5 in Genome Data Viewer
    Location:
    19 A; 19 4.34 cM
    Exon count:
    15
    Annotation release Status Assembly Chr Location
    RS_2024_02 current GRCm39 (GCF_000001635.27) 19 NC_000085.7 (5653042..5661584, complement)
    108.20200622 previous assembly GRCm38.p6 (GCF_000001635.26) 19 NC_000085.6 (5603014..5611556, complement)

    Chromosome 19 - NC_000085.7Genomic Context describing neighboring genes Neighboring gene ovo like zinc finger 1 Neighboring gene adaptor-related protein complex 5, beta 1 subunit Neighboring gene STARR-seq mESC enhancer starr_45393 Neighboring gene CapStarr-seq enhancer MGSCv37_chr19:5601776-5601959 Neighboring gene ribonuclease H2, subunit C Neighboring gene STARR-positive B cell enhancer ABC_E11011 Neighboring gene v-rel reticuloendotheliosis viral oncogene homolog A (avian) Neighboring gene signal-induced proliferation associated gene 1

    Genomic regions, transcripts, and products

    Expression

    • Project title: Mouse ENCODE transcriptome data 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)
    • Endonuclease-mediated (4) 
    • Gene trapped (2) 
    • Targeted (11)  1 citation

    Pathways from PubChem

    Interactions

    Products Interactant Other Gene Complex Source Pubs Description

    General gene information

    Markers

    Clone Names

    • MGC117972

    Gene Ontology Provided by MGI

    Function Evidence Code Pubs
    enables DNA-binding transcription factor binding IEA
    Inferred from Electronic Annotation
    more info
     
    enables DNA-binding transcription factor binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables chromatin binding IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    enables chromatin binding IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables chromatin binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables histone H2A acetyltransferase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables histone H2A acetyltransferase activity ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    enables histone H2AK5 acetyltransferase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H2AK5 acetyltransferase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables histone H4K16 acetyltransferase activity IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    enables histone H4K16 acetyltransferase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables histone H4K16 acetyltransferase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables histone acetyltransferase activity IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    enables histone acetyltransferase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables metal ion binding IEA
    Inferred from Electronic Annotation
    more info
     
    enables peptide 2-hydroxyisobutyryltransferase activity IEA
    Inferred from Electronic Annotation
    more info
     
    enables peptide butyryltransferase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables peptide butyryltransferase activity ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    enables peptide crotonyltransferase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables peptide crotonyltransferase activity ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    enables peptide-lysine-N-acetyltransferase activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables peptide-lysine-N-acetyltransferase activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables phospholipase binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables protein binding IPI
    Inferred from Physical Interaction
    more info
    PubMed 
    enables protein-containing complex binding ISO
    Inferred from Sequence Orthology
    more info
     
    enables transcription coactivator activity IDA
    Inferred from Direct Assay
    more info
    PubMed 
    enables transcription coactivator activity ISO
    Inferred from Sequence Orthology
    more info
     
    enables transcription coactivator activity ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    enables transcription coregulator activity IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    Process Evidence Code Pubs
    involved_in DNA damage response IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in DNA damage response ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in DNA damage response, signal transduction by p53 class mediator resulting in transcription of p21 class mediator ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in DNA damage response, signal transduction by p53 class mediator resulting in transcription of p21 class mediator ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in DNA repair-dependent chromatin remodeling IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in DNA repair-dependent chromatin remodeling ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in apoptotic process IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in cellular response to estradiol stimulus IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in cellular response to estradiol stimulus ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cellular response to glucose starvation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in cellular response to glucose starvation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in cellular response to glucose stimulus IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in double-strand break repair ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in double-strand break repair via homologous recombination IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    involved_in double-strand break repair via homologous recombination IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in double-strand break repair via homologous recombination ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in establishment of mitotic spindle orientation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in establishment of mitotic spindle orientation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in innate immune response IEA
    Inferred from Electronic Annotation
    more info
     
    acts_upstream_of_positive_effect lipid droplet disassembly ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_positive_effect lipid droplet disassembly ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in negative regulation of DNA-templated transcription ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of double-strand break repair via homologous recombination IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in negative regulation of double-strand break repair via homologous recombination ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of interleukin-2 production IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in negative regulation of interleukin-2 production ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in negative regulation of transcription by RNA polymerase II IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in negative regulation of transcription by RNA polymerase II ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in neural tube development IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in neurogenesis IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in nucleotide-excision repair ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in nucleotide-excision repair ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in peptidyl-lysine acetylation IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in peptidyl-lysine acetylation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of DNA-templated transcription ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of DNA-templated transcription ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of DNA-templated transcription NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in positive regulation of aggrephagy IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in positive regulation of aggrephagy ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of attachment of mitotic spindle microtubules to kinetochore ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of attachment of mitotic spindle microtubules to kinetochore ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of autophagy IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of autophagy ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of circadian rhythm IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in positive regulation of double-strand break repair via homologous recombination ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of double-strand break repair via homologous recombination ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of innate immune response ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of mitotic sister chromatid segregation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of mitotic sister chromatid segregation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of myoblast differentiation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of myoblast differentiation ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of protein acetylation IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in positive regulation of protein acetylation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of regulatory T cell differentiation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in positive regulation of regulatory T cell differentiation ISO
    Inferred from Sequence Orthology
    more info
     
    acts_upstream_of_or_within positive regulation of transcription by RNA polymerase II IGI
    Inferred from Genetic Interaction
    more info
    PubMed 
    involved_in positive regulation of transcription by RNA polymerase II ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of transcription by RNA polymerase II ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in positive regulation of triglyceride biosynthetic process ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in positive regulation of triglyceride biosynthetic process ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    involved_in proteasome-mediated ubiquitin-dependent protein catabolic process IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in proteasome-mediated ubiquitin-dependent protein catabolic process ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in protein acetylation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in protein modification process ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in regulation of apoptotic process NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in regulation of cell cycle ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in regulation of chromatin organization NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in regulation of double-strand break repair NAS
    Non-traceable Author Statement
    more info
    PubMed 
    involved_in regulation of hematopoietic stem cell differentiation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in regulation of transcription by RNA polymerase II IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    acts_upstream_of_or_within regulation of transcription by RNA polymerase II IDA
    Inferred from Direct Assay
    more info
    PubMed 
    involved_in response to ionizing radiation IEA
    Inferred from Electronic Annotation
    more info
     
    involved_in response to ionizing radiation ISO
    Inferred from Sequence Orthology
    more info
     
    involved_in sperm DNA condensation IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    involved_in spermatid development IMP
    Inferred from Mutant Phenotype
    more info
    PubMed 
    Component Evidence Code Pubs
    part_of NuA4 histone acetyltransferase complex IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    part_of NuA4 histone acetyltransferase complex IDA
    Inferred from Direct Assay
    more info
    PubMed 
    part_of NuA4 histone acetyltransferase complex ISO
    Inferred from Sequence Orthology
    more info
     
    part_of NuA4 histone acetyltransferase complex ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    part_of Swr1 complex ISO
    Inferred from Sequence Orthology
    more info
     
    part_of Swr1 complex ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in chromatin IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    located_in chromatin ISO
    Inferred from Sequence Orthology
    more info
     
    located_in cytoplasm ISO
    Inferred from Sequence Orthology
    more info
     
    located_in cytoplasm ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in cytosol IEA
    Inferred from Electronic Annotation
    more info
     
    located_in cytosol ISO
    Inferred from Sequence Orthology
    more info
     
    located_in intracellular membrane-bounded organelle ISO
    Inferred from Sequence Orthology
    more info
     
    located_in kinetochore ISO
    Inferred from Sequence Orthology
    more info
     
    located_in kinetochore ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in mitotic spindle pole IEA
    Inferred from Electronic Annotation
    more info
     
    located_in mitotic spindle pole ISO
    Inferred from Sequence Orthology
    more info
     
    located_in nucleolus ISO
    Inferred from Sequence Orthology
    more info
     
    located_in nucleolus ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in nucleoplasm ISO
    Inferred from Sequence Orthology
    more info
     
    located_in nucleoplasm TAS
    Traceable Author Statement
    more info
     
    part_of nucleosome ISO
    Inferred from Sequence Orthology
    more info
     
    is_active_in nucleus IBA
    Inferred from Biological aspect of Ancestor
    more info
     
    located_in nucleus ISO
    Inferred from Sequence Orthology
    more info
     
    located_in nucleus ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    located_in nucleus NAS
    Non-traceable Author Statement
    more info
    PubMed 
    located_in perinuclear region of cytoplasm IEA
    Inferred from Electronic Annotation
    more info
     
    part_of piccolo histone acetyltransferase complex ISO
    Inferred from Sequence Orthology
    more info
     
    part_of piccolo histone acetyltransferase complex ISS
    Inferred from Sequence or Structural Similarity
    more info
     
    part_of protein-containing complex ISO
    Inferred from Sequence Orthology
    more info
     
    located_in site of double-strand break IEA
    Inferred from Electronic Annotation
    more info
     
    is_active_in site of double-strand break ISO
    Inferred from Sequence Orthology
    more info
     
    part_of transcription regulator complex IDA
    Inferred from Direct Assay
    more info
    PubMed 

    General protein information

    Preferred Names
    histone acetyltransferase KAT5
    Names
    60 kDa Tat-interactive protein
    HIV-1 tat interactive protein 1, 60 kDa homolog
    HIV-1 tat interactive protein 1, homolog
    Tat-interactive protein-60
    histone acetyltransferase HTATIP
    lysine acetyltransferase 5
    protein 2-hydroxyisobutyryltransferase KAT5
    protein acetyltransferase KAT5
    protein crotonyltransferase KAT5
    NP_001186176.1
    NP_001186177.1
    NP_001186178.1
    NP_001349299.1
    NP_001349300.1
    NP_001349301.1
    NP_848752.1
    XP_006531923.1

    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_001199247.1NP_001186176.1  histone acetyltransferase KAT5 isoform 2

      See identical proteins and their annotated locations for NP_001186176.1

      Status: VALIDATED

      Description
      Transcript Variant: This variant (2) lacks an in-frame exon in the 5' coding region, compared to variant 1, and encodes a shorter isoform (2, also known as beta), compared to isoform 1.
      Source sequence(s)
      AF528196, CJ080877, DV648619
      Consensus CDS
      CCDS89314.1
      UniProtKB/TrEMBL
      A2VDH4
      Related
      ENSMUSP00000158391.2, ENSMUST00000236229.2
      Conserved Domains (2) summary
      PLN00104
      Location:43452
      PLN00104; MYST -like histone acetyltransferase; Provisional
      pfam11717
      Location:765
      Tudor-knot; RNA binding activity-knot of a chromodomain
    2. NM_001199248.1NP_001186177.1  histone acetyltransferase KAT5 isoform 3

      See identical proteins and their annotated locations for NP_001186177.1

      Status: VALIDATED

      Description
      Transcript Variant: This variant (3) has multiple differences in the 5' coding region, compared to variant 1. It encodes a shorter isoform (3, also known as gamma), compared to isoform 1.
      Source sequence(s)
      AF528195, AK160077, CJ080877, DV648619
      Consensus CDS
      CCDS89313.1
      UniProtKB/TrEMBL
      A0A494BAP7, A2VDH4
      Related
      ENSMUSP00000158181.2, ENSMUST00000235701.2
      Conserved Domains (2) summary
      PLN00104
      Location:76485
      PLN00104; MYST -like histone acetyltransferase; Provisional
      pfam11717
      Location:4098
      Tudor-knot; RNA binding activity-knot of a chromodomain
    3. NM_001199249.1NP_001186178.1  histone acetyltransferase KAT5 isoform 4

      Status: VALIDATED

      Description
      Transcript Variant: This variant (4) has multiple differences in the 5' and 3' coding regions, compared to variant 1. It encodes a shorter isoform (4, also known as delta) with a unique C-terminus, compared to isoform 1.
      Source sequence(s)
      AK143674, CA317697, CJ080877
      Related
      ENSMUST00000235788.2
      Conserved Domains (2) summary
      pfam11717
      Location:4098
      Tudor-knot; RNA binding activity-knot of a chromodomain
      cl17182
      Location:200378
      NAT_SF; N-Acyltransferase superfamily: Various enyzmes that characteristicly catalyze the transfer of an acyl group to a substrate
    4. NM_001362370.1NP_001349299.1  histone acetyltransferase KAT5 isoform 5

      Status: VALIDATED

      Source sequence(s)
      AC126029, AF528196, BY269987, DV648619
      Related
      ENSMUST00000236964.2
      Conserved Domains (1) summary
      PLN00104
      Location:7293
      PLN00104; MYST -like histone acetyltransferase; Provisional
    5. NM_001362371.1NP_001349300.1  histone acetyltransferase KAT5 isoform 5

      Status: VALIDATED

      Source sequence(s)
      AC126029, DV648619
      Conserved Domains (1) summary
      PLN00104
      Location:7293
      PLN00104; MYST -like histone acetyltransferase; Provisional
    6. NM_001362372.1NP_001349301.1  histone acetyltransferase KAT5 isoform 6

      Status: VALIDATED

      Source sequence(s)
      AC126029, DV648619
      UniProtKB/TrEMBL
      A2VDH4
      Conserved Domains (2) summary
      PLN00104
      Location:76537
      PLN00104; MYST -like histone acetyltransferase; Provisional
      pfam11717
      Location:4098
      Tudor-knot; RNA binding activity-knot of a chromodomain
    7. NM_178637.2NP_848752.1  histone acetyltransferase KAT5 isoform 1

      See identical proteins and their annotated locations for NP_848752.1

      Status: VALIDATED

      Description
      Transcript Variant: This variant (1) represents the longest transcript and encodes the longest isoform (1, also known as alpha).
      Source sequence(s)
      AF528195, CJ080877, DV648619
      Consensus CDS
      CCDS29472.1
      UniProtKB/Swiss-Prot
      A0A494B9U8, A1L394, Q3YFI9, Q8CGZ3, Q8CGZ4, Q8CHK4, Q8VIH0
      UniProtKB/TrEMBL
      A2VDH4, Q3UJQ1
      Related
      ENSMUSP00000109271.3, ENSMUST00000113641.4
      Conserved Domains (2) summary
      PLN00104
      Location:43504
      PLN00104; MYST -like histone acetyltransferase; Provisional
      pfam11717
      Location:765
      Tudor-knot; RNA binding activity-knot of a chromodomain

    RNA

    1. NR_155544.1 RNA Sequence

      Status: VALIDATED

      Source sequence(s)
      AC126029, DV648619
      Related
      ENSMUST00000236883.2

    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_000085.7 Reference GRCm39 C57BL/6J

      Range
      5653042..5661584 complement
      Download
      GenBank, FASTA, Sequence Viewer (Graphics)

    mRNA and Protein(s)

    1. XM_006531860.3XP_006531923.1  histone acetyltransferase KAT5 isoform X1

      See identical proteins and their annotated locations for XP_006531923.1

      UniProtKB/TrEMBL
      A2VDH4
      Conserved Domains (2) summary
      PLN00104
      Location:227513
      PLN00104; MYST -like histone acetyltransferase; Provisional
      cd18985
      Location:1477
      CBD_TIP60_like; chromo barrel domain of human tat-interactive protein 60, and similar proteins