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Entry version 86 (02 Jun 2021)
Sequence version 1 (30 Aug 2005)
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Protein

ADAMTS-like protein 4

Gene

Adamtsl4

Organism
Rattus norvegicus (Rat)
Status
Reviewed-Annotation score:

Annotation score:3 out of 5

<p>The annotation score provides a heuristic measure of the annotation content of a UniProtKB entry or proteome. This score <strong>cannot</strong> be used as a measure of the accuracy of the annotation as we cannot define the 'correct annotation' for any given protein.<p><a href='/help/annotation_score' target='_top'>More...</a></p>
-Experimental evidence at transcript leveli <p>This indicates the type of evidence that supports the existence of the protein. Note that the 'protein existence' evidence does not give information on the accuracy or correctness of the sequence(s) displayed.<p><a href='/help/protein_existence' target='_top'>More...</a></p>

<p>This section provides any useful information about the protein, mostly biological knowledge.<p><a href='/help/function_section' target='_top'>More...</a></p>Functioni

Positive regulation of apoptosis. May facilitate FBN1 microfibril biogenesis (By similarity).

By similarity

Caution

Although similar to members of the ADAMTS family, it lacks the metalloprotease and disintegrin-like domains which are typical of that family.Curated

<p>The <a href="http://www.geneontology.org/">Gene Ontology (GO)</a> project provides a set of hierarchical controlled vocabulary split into 3 categories:<p><a href='/help/gene_ontology' target='_top'>More...</a></p>GO - Molecular functioni

GO - Biological processi

<p>UniProtKB Keywords constitute a <a href="http://www.uniprot.org/keywords">controlled vocabulary</a> with a hierarchical structure. Keywords summarise the content of a UniProtKB entry and facilitate the search for proteins of interest.<p><a href='/help/keywords' target='_top'>More...</a></p>Keywordsi

Biological processApoptosis

Enzyme and pathway databases

Reactome - a knowledgebase of biological pathways and processes

More...
Reactomei
R-RNO-5173214, O-glycosylation of TSR domain-containing proteins

<p>This section provides information about the protein and gene name(s) and synonym(s) and about the organism that is the source of the protein sequence.<p><a href='/help/names_and_taxonomy_section' target='_top'>More...</a></p>Names & Taxonomyi

<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section provides an exhaustive list of all names of the protein, from commonly used to obsolete, to allow unambiguous identification of a protein.<p><a href='/help/protein_names' target='_top'>More...</a></p>Protein namesi
Recommended name:
ADAMTS-like protein 4
Short name:
ADAMTSL-4
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section indicates the name(s) of the gene(s) that code for the protein sequence(s) described in the entry. Four distinct tokens exist: 'Name', 'Synonyms', 'Ordered locus names' and 'ORF names'.<p><a href='/help/gene_name' target='_top'>More...</a></p>Gene namesi
Name:Adamtsl4Imported
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section provides information on the name(s) of the organism that is the source of the protein sequence.<p><a href='/help/organism-name' target='_top'>More...</a></p>OrganismiRattus norvegicus (Rat)
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section shows the unique identifier assigned by the NCBI to the source organism of the protein. This is known as the 'taxonomic identifier' or 'taxid'.<p><a href='/help/taxonomic_identifier' target='_top'>More...</a></p>Taxonomic identifieri10116 [NCBI]
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section contains the taxonomic hierarchical classification lineage of the source organism. It lists the nodes as they appear top-down in the taxonomic tree, with the more general grouping listed first.<p><a href='/help/taxonomic_lineage' target='_top'>More...</a></p>Taxonomic lineageiEukaryotaMetazoaChordataCraniataVertebrataEuteleostomiMammaliaEutheriaEuarchontogliresGliresRodentiaMyomorphaMuroideaMuridaeMurinaeRattus
<p>This subsection of the <a href="http://www.uniprot.org/help/names%5Fand%5Ftaxonomy%5Fsection">Names and taxonomy</a> section is present for entries that are part of a <a href="http://www.uniprot.org/proteomes">proteome</a>, i.e. of a set of proteins thought to be expressed by organisms whose genomes have been completely sequenced.<p><a href='/help/proteomes_manual' target='_top'>More...</a></p>Proteomesi
  • UP000002494 <p>A UniProt <a href="http://www.uniprot.org/manual/proteomes%5Fmanual">proteome</a> can consist of several components.<br></br>The component name refers to the genomic component encoding a set of proteins.<p><a href='/help/proteome_component' target='_top'>More...</a></p> Componenti: Chromosome 2

Organism-specific databases

Rat genome database

More...
RGDi
1561012, Adamtsl4

<p>This section provides information on the location and the topology of the mature protein in the cell.<p><a href='/help/subcellular_location_section' target='_top'>More...</a></p>Subcellular locationi

Keywords - Cellular componenti

Extracellular matrix, Secreted

<p>This section describes post-translational modifications (PTMs) and/or processing events.<p><a href='/help/ptm_processing_section' target='_top'>More...</a></p>PTM / Processingi

Molecule processing

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'PTM / Processing' section denotes the presence of an N-terminal signal peptide.<p><a href='/help/signal' target='_top'>More...</a></p>Signal peptidei1 – 24Sequence analysisAdd BLAST24
<p>This subsection of the 'PTM / Processing' section describes the extent of a polypeptide chain in the mature protein following processing or proteolytic cleavage.<p><a href='/help/chain' target='_top'>More...</a></p>ChainiPRO_000025796825 – 1030ADAMTS-like protein 4Sequence analysisAdd BLAST1006

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/help/ptm%5Fprocessing%5Fsection">PTM / Processing</a> section specifies the position and type of each covalently attached glycan group (mono-, di-, or polysaccharide).<p><a href='/help/carbohyd' target='_top'>More...</a></p>Glycosylationi451N-linked (GlcNAc...) asparagineSequence analysis1
Glycosylationi731N-linked (GlcNAc...) asparagineSequence analysis1

<p>This subsection of the <a href="http://www.uniprot.org/help/ptm%5Fprocessing%5Fsection">PTM/processing</a> section describes post-translational modifications (PTMs). This subsection <strong>complements</strong> the information provided at the sequence level or describes modifications for which <strong>position-specific data is not yet available</strong>.<p><a href='/help/post-translational_modification' target='_top'>More...</a></p>Post-translational modificationi

Glycosylated (By similarity). Can be O-fucosylated by POFUT2 on a serine or a threonine residue found within the consensus sequence C1-X2-(S/T)-C2-G of the TSP type-1 repeat domains where C1 and C2 are the first and second cysteine residue of the repeat, respectively. Fucosylated repeats can then be further glycosylated by the addition of a beta-1,3-glucose residue by the glucosyltransferase, B3GALTL. Fucosylation mediates the efficient secretion of ADAMTS family members. Also can be C-glycosylated with one or two mannose molecules on tryptophan residues within the consensus sequence W-X-X-W of the TPRs, and N-glycosylated. These other glycosylations can also facilitate secretion (By similarity).By similarity

Keywords - PTMi

Glycoprotein

Proteomic databases

PaxDb, a database of protein abundance averages across all three domains of life

More...
PaxDbi
Q4FZU4

PRoteomics IDEntifications database

More...
PRIDEi
Q4FZU4

PTM databases

GlyGen: Computational and Informatics Resources for Glycoscience

More...
GlyGeni
Q4FZU4, 2 sites

<p>This section provides information on the expression of a gene at the mRNA or protein level in cells or in tissues of multicellular organisms.<p><a href='/help/expression_section' target='_top'>More...</a></p>Expressioni

Gene expression databases

Bgee dataBase for Gene Expression Evolution

More...
Bgeei
ENSRNOG00000049385, Expressed in esophagus and 21 other tissues

Genevisible search portal to normalized and curated expression data from Genevestigator

More...
Genevisiblei
Q4FZU4, RN

<p>This section provides information on the quaternary structure of a protein and on interaction(s) with other proteins or protein complexes.<p><a href='/help/interaction_section' target='_top'>More...</a></p>Interactioni

<p>This subsection of the <a href="http://www.uniprot.org/help/interaction%5Fsection">'Interaction'</a> section provides information about the protein quaternary structure and interaction(s) with other proteins or protein complexes (with the exception of physiological receptor-ligand interactions which are annotated in the <a href="http://www.uniprot.org/help/function%5Fsection">'Function'</a> section).<p><a href='/help/subunit_structure' target='_top'>More...</a></p>Subunit structurei

Interacts with CTSB.

Interacts with FBN1 (By similarity).

By similarity

GO - Molecular functioni

Protein-protein interaction databases

STRING: functional protein association networks

More...
STRINGi
10116.ENSRNOP00000063998

<p>This section provides information on the tertiary and secondary structure of a protein.<p><a href='/help/structure_section' target='_top'>More...</a></p>Structurei

3D structure databases

SWISS-MODEL Repository - a database of annotated 3D protein structure models

More...
SMRi
Q4FZU4

Database of comparative protein structure models

More...
ModBasei
Search...

<p>This section provides information on sequence similarities with other proteins and the domain(s) present in a protein.<p><a href='/help/family_and_domains_section' target='_top'>More...</a></p>Family & Domainsi

Domains and Repeats

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/help/family%5Fand%5Fdomains%5Fsection">Family and Domains</a> section describes the position and type of a domain, which is defined as a specific combination of secondary structures organized into a characteristic three-dimensional structure or fold.<p><a href='/help/domain' target='_top'>More...</a></p>Domaini47 – 91TSP type-1 1PROSITE-ProRule annotationAdd BLAST45
Domaini681 – 740TSP type-1 2PROSITE-ProRule annotationAdd BLAST60
Domaini741 – 800TSP type-1 3PROSITE-ProRule annotationAdd BLAST60
Domaini803 – 865TSP type-1 4PROSITE-ProRule annotationAdd BLAST63
Domaini866 – 925TSP type-1 5PROSITE-ProRule annotationAdd BLAST60
Domaini926 – 982TSP type-1 6PROSITE-ProRule annotationAdd BLAST57
Domaini985 – 1022PLACPROSITE-ProRule annotationAdd BLAST38

Region

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'Family and Domains' section describes a region of interest that cannot be described in other subsections.<p><a href='/help/region' target='_top'>More...</a></p>Regioni73 – 150DisorderedSequence analysisAdd BLAST78
Regioni168 – 306DisorderedSequence analysisAdd BLAST139

Compositional bias

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'Family and Domains' section describes the position of regions of compositional bias within the protein and the particular type of amino acids that are over-represented within those regions.<p><a href='/help/compbias' target='_top'>More...</a></p>Compositional biasi178 – 198Polar residuesSequence analysisAdd BLAST21
Compositional biasi207 – 284Polar residuesSequence analysisAdd BLAST78

Keywords - Domaini

Repeat, Signal

Phylogenomic databases

evolutionary genealogy of genes: Non-supervised Orthologous Groups

More...
eggNOGi
KOG3538, Eukaryota
KOG4597, Eukaryota

Ensembl GeneTree

More...
GeneTreei
ENSGT00940000161136

The HOGENOM Database of Homologous Genes from Fully Sequenced Organisms

More...
HOGENOMi
CLU_000660_6_0_1

InParanoid: Eukaryotic Ortholog Groups

More...
InParanoidi
Q4FZU4

Identification of Orthologs from Complete Genome Data

More...
OMAi
QICNTIP

Database of Orthologous Groups

More...
OrthoDBi
414258at2759

Database for complete collections of gene phylogenies

More...
PhylomeDBi
Q4FZU4

Family and domain databases

Gene3D Structural and Functional Annotation of Protein Families

More...
Gene3Di
2.20.100.10, 6 hits

Integrated resource of protein families, domains and functional sites

More...
InterProi
View protein in InterPro
IPR010294, ADAM_spacer1
IPR010909, PLAC
IPR000884, TSP1_rpt
IPR036383, TSP1_rpt_sf

Pfam protein domain database

More...
Pfami
View protein in Pfam
PF05986, ADAM_spacer1, 1 hit
PF08686, PLAC, 1 hit
PF00090, TSP_1, 1 hit

Simple Modular Architecture Research Tool; a protein domain database

More...
SMARTi
View protein in SMART
SM00209, TSP1, 7 hits

Superfamily database of structural and functional annotation

More...
SUPFAMi
SSF82895, SSF82895, 6 hits

PROSITE; a protein domain and family database

More...
PROSITEi
View protein in PROSITE
PS50900, PLAC, 1 hit
PS50092, TSP1, 6 hits

<p>This section displays by default the canonical protein sequence and upon request all isoforms described in the entry. It also includes information pertinent to the sequence(s), including <a href="http://www.uniprot.org/help/sequence%5Flength">length</a> and <a href="http://www.uniprot.org/help/sequences">molecular weight</a>. The information is filed in different subsections. The current subsections and their content are listed below:<p><a href='/help/sequences_section' target='_top'>More...</a></p>Sequencei

<p>This subsection of the <a href="http://www.uniprot.org/help/sequences%5Fsection">Sequence</a> section indicates if the <a href="http://www.uniprot.org/help/canonical%5Fand%5Fisoforms">canonical sequence</a> displayed by default in the entry is complete or not.<p><a href='/help/sequence_status' target='_top'>More...</a></p>Sequence statusi: Complete.

<p>This subsection of the <a href="http://www.uniprot.org/help/sequences%5Fsection">Sequence</a> section indicates if the <a href="http://www.uniprot.org/help/canonical%5Fand%5Fisoforms">canonical sequence</a> displayed by default in the entry is in its mature form or if it represents the precursor.<p><a href='/help/sequence_processing' target='_top'>More...</a></p>Sequence processingi: The displayed sequence is further processed into a mature form.

Q4FZU4-1 [UniParc]FASTAAdd to basket
« Hide
        10         20         30         40         50
MELWLGRLWL YVMLLLLLLQ LCQDQELLGP SLQTPSEEDQ VPEGLWGPWG
60 70 80 90 100
RWASCSQPCG VGVQRRSRTC ELHPALSLPP RPPRHPEAPQ PRGQGSRPQT
110 120 130 140 150
PRDPQSLYRP QPRGRGGPLR GPASQVGREE TQEPRGAQRF RVRDPIKPGM
160 170 180 190 200
FGYGRVPFAL PLHRSRRLAH KPGQPKDSST AEETLPSQPP STEPASEKHS
210 220 230 240 250
PHMQPPELRA QSRSPSAETP RSGTAQTEVP SRTSSAPSDM GIPAPTSSFR
260 270 280 290 300
DSRSFQGSPE PRMPTSQGAE RQPHPFSPVT RSQLSRRHWR PPGSPHRSPD
310 320 330 340 350
GWLPLTRDSS PHWSLFAPSS PTPECSGESE QMRACSQEPC PPEQPDPRAL
360 370 380 390 400
QCAAFDSQEF MGQLYQWEPF TEVQGSQRCE LNCRPRGFRF YVRHTEKVQD
410 420 430 440 450
GTLCQPGSLD ICVAGHCLSP GCDGILGSGR RPDGCGVCGG DGSTCRLVSG
460 470 480 490 500
NLTDRGGPLG YQKILWIPAG ASHLRISQFR PSSNYLALRG PGGRSIINGN
510 520 530 540 550
WAVDPPGSYA AVGTVFQYNR PPREEGKGET LSAEGPTTQP VDVYMIFQED
560 570 580 590 600
NPGVFYQYVT SAAPESPSTM PPALQLQPEM LRGEPLLPSA PRPVRAPGTL
610 620 630 640 650
QRQARIPQVP PPTHVRTAMG SSAGYWKQVG HSECSASCGK GVWRPIFLCV
660 670 680 690 700
SRESGEELDE QSCAVGARPP ASPESCHRPP CPPYWEAGEW TSCSRSCGPG
710 720 730 740 750
TQHRQLLCRQ EFGGGGSSVP PERCGHLPRP NITQSCQLRL CGHWEISSPW
760 770 780 790 800
SQCSVRCGRG QRSRQVRCVG SNGHEVGKQE CASGPPPPPS REACDMGPCT
810 820 830 840 850
TAWFYSDWSS KCSAECGTGI QRRAVVCLRS GETLQGDPEA GSTEQGCPLR
860 870 880 890 900
SRPPDMRACS LGPCEKTWRW YTGPWSECSS ECGSGTQHRD IICVSKLGTK
910 920 930 940 950
FNVTSPSNCS HLPRPPALQP CQGQACEDQW FSTLWSPCSQ SCQGGVQTRE
960 970 980 990 1000
VQCLSSNHTL SSRCPPHLRP SRKRPCNSQP CNQRPDDQCK DSSPHCPLVV
1010 1020 1030
QARLCVYPYY TATCCRSCAH VLEQSQLEPA
Length:1,030
Mass (Da):112,785
Last modified:August 30, 2005 - v1
<p>The checksum is a form of redundancy check that is calculated from the sequence. It is useful for tracking sequence updates.</p> <p>It should be noted that while, in theory, two different sequences could have the same checksum value, the likelihood that this would happen is extremely low.</p> <p>However UniProtKB may contain entries with identical sequences in case of multiple genes (paralogs).</p> <p>The checksum is computed as the sequence 64-bit Cyclic Redundancy Check value (CRC64) using the generator polynomial: x<sup>64</sup> + x<sup>4</sup> + x<sup>3</sup> + x + 1. The algorithm is described in the ISO 3309 standard. </p> <p class="publication">Press W.H., Flannery B.P., Teukolsky S.A. and Vetterling W.T.<br /> <strong>Cyclic redundancy and other checksums</strong><br /> <a href="http://www.nrbook.com/b/bookcpdf.php">Numerical recipes in C 2nd ed., pp896-902, Cambridge University Press (1993)</a>)</p> Checksum:iE38CC6486E08546B
GO

Sequence databases

Select the link destinations:

EMBL nucleotide sequence database

More...
EMBLi

GenBank nucleotide sequence database

More...
GenBanki

DNA Data Bank of Japan; a nucleotide sequence database

More...
DDBJi
Links Updated
BC099119 mRNA Translation: AAH99119.1

NCBI Reference Sequences

More...
RefSeqi
NP_001029184.1, NM_001034012.1
XP_008759529.2, XM_008761307.2
XP_017446392.1, XM_017590903.1
XP_017446393.1, XM_017590904.1
XP_017446394.1, XM_017590905.1
XP_017446395.1, XM_017590906.1
XP_017446396.1, XM_017590907.1

Genome annotation databases

Ensembl eukaryotic genome annotation project

More...
Ensembli
ENSRNOT00000074156; ENSRNOP00000063998; ENSRNOG00000049385

Database of genes from NCBI RefSeq genomes

More...
GeneIDi
310670

KEGG: Kyoto Encyclopedia of Genes and Genomes

More...
KEGGi
rno:310670

<p>This section provides links to proteins that are similar to the protein sequence(s) described in this entry at different levels of sequence identity thresholds (100%, 90% and 50%) based on their membership in UniProt Reference Clusters (<a href="http://www.uniprot.org/help/uniref">UniRef</a>).<p><a href='/help/similar_proteins_section' target='_top'>More...</a></p>Similar proteinsi

<p>This section is used to point to information related to entries and found in data collections other than UniProtKB.<p><a href='/help/cross_references_section' target='_top'>More...</a></p>Cross-referencesi

Sequence databases

Select the link destinations:
EMBLi
GenBanki
DDBJi
Links Updated
BC099119 mRNA Translation: AAH99119.1
RefSeqiNP_001029184.1, NM_001034012.1
XP_008759529.2, XM_008761307.2
XP_017446392.1, XM_017590903.1
XP_017446393.1, XM_017590904.1
XP_017446394.1, XM_017590905.1
XP_017446395.1, XM_017590906.1
XP_017446396.1, XM_017590907.1

3D structure databases

SMRiQ4FZU4
ModBaseiSearch...

Protein-protein interaction databases

STRINGi10116.ENSRNOP00000063998

PTM databases

GlyGeniQ4FZU4, 2 sites

Proteomic databases

PaxDbiQ4FZU4
PRIDEiQ4FZU4

Genome annotation databases

EnsembliENSRNOT00000074156; ENSRNOP00000063998; ENSRNOG00000049385
GeneIDi310670
KEGGirno:310670

Organism-specific databases

Comparative Toxicogenomics Database

More...
CTDi
54507
RGDi1561012, Adamtsl4

Phylogenomic databases

eggNOGiKOG3538, Eukaryota
KOG4597, Eukaryota
GeneTreeiENSGT00940000161136
HOGENOMiCLU_000660_6_0_1
InParanoidiQ4FZU4
OMAiQICNTIP
OrthoDBi414258at2759
PhylomeDBiQ4FZU4

Enzyme and pathway databases

ReactomeiR-RNO-5173214, O-glycosylation of TSR domain-containing proteins

Miscellaneous databases

Protein Ontology

More...
PROi
PR:Q4FZU4

Gene expression databases

BgeeiENSRNOG00000049385, Expressed in esophagus and 21 other tissues
GenevisibleiQ4FZU4, RN

Family and domain databases

Gene3Di2.20.100.10, 6 hits
InterProiView protein in InterPro
IPR010294, ADAM_spacer1
IPR010909, PLAC
IPR000884, TSP1_rpt
IPR036383, TSP1_rpt_sf
PfamiView protein in Pfam
PF05986, ADAM_spacer1, 1 hit
PF08686, PLAC, 1 hit
PF00090, TSP_1, 1 hit
SMARTiView protein in SMART
SM00209, TSP1, 7 hits
SUPFAMiSSF82895, SSF82895, 6 hits
PROSITEiView protein in PROSITE
PS50900, PLAC, 1 hit
PS50092, TSP1, 6 hits

MobiDB: a database of protein disorder and mobility annotations

More...
MobiDBi
Search...

<p>This section provides general information on the entry.<p><a href='/help/entry_information_section' target='_top'>More...</a></p>Entry informationi

<p>This subsection of the 'Entry information' section provides a mnemonic identifier for a UniProtKB entry, but it is not a stable identifier. Each reviewed entry is assigned a unique entry name upon integration into UniProtKB/Swiss-Prot.<p><a href='/help/entry_name' target='_top'>More...</a></p>Entry nameiATL4_RAT
<p>This subsection of the 'Entry information' section provides one or more accession number(s). These are stable identifiers and should be used to cite UniProtKB entries. Upon integration into UniProtKB, each entry is assigned a unique accession number, which is called 'Primary (citable) accession number'.<p><a href='/help/accession_numbers' target='_top'>More...</a></p>AccessioniPrimary (citable) accession number: Q4FZU4
<p>This subsection of the 'Entry information' section shows the date of integration of the entry into UniProtKB, the date of the last sequence update and the date of the last annotation modification ('Last modified'). The version number for both the entry and the <a href="http://www.uniprot.org/help/canonical%5Fand%5Fisoforms">canonical sequence</a> are also displayed.<p><a href='/help/entry_history' target='_top'>More...</a></p>Entry historyiIntegrated into UniProtKB/Swiss-Prot: October 31, 2006
Last sequence update: August 30, 2005
Last modified: June 2, 2021
This is version 86 of the entry and version 1 of the sequence. See complete history.
<p>This subsection of the 'Entry information' section indicates whether the entry has been manually annotated and reviewed by UniProtKB curators or not, in other words, if the entry belongs to the Swiss-Prot section of UniProtKB (<strong>reviewed</strong>) or to the computer-annotated TrEMBL section (<strong>unreviewed</strong>).<p><a href='/help/entry_status' target='_top'>More...</a></p>Entry statusiReviewed (UniProtKB/Swiss-Prot)
Annotation programChordata Protein Annotation Program

<p>This section contains any relevant information that doesn't fit in any other defined sections<p><a href='/help/miscellaneous_section' target='_top'>More...</a></p>Miscellaneousi

Keywords - Technical termi

Reference proteome
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