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UniProtKB - P04233 (HG2A_HUMAN)

Entry version 225 (10 Feb 2021)
Sequence version 3 (16 Apr 2002)
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Protein

HLA class II histocompatibility antigen gamma chain

Gene

CD74

Organism
Homo sapiens (Human)
Status
Reviewed-Annotation score:

Annotation score:5 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 protein 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

Plays a critical role in MHC class II antigen processing by stabilizing peptide-free class II alpha/beta heterodimers in a complex soon after their synthesis and directing transport of the complex from the endoplasmic reticulum to the endosomal/lysosomal system where the antigen processing and binding of antigenic peptides to MHC class II takes place. Serves as cell surface receptor for the cytokine MIF.
Binds to the peptide-binding site of MHC class II alpha/beta heterodimers forming an alpha-beta-CLIP complex, thereby preventing the loading of antigenic peptides to the MHC class II complex until its release by HLA-DM in the endosome.1 Publication
Stabilizes the conformation of mature CTSL by binding to its active site and serving as a chaperone to help maintain a pool of mature enzyme in endocytic compartments and extracellular space of antigen-presenting cells (APCs). Has antiviral activity by stymieing the endosomal entry of Ebola virus and coronaviruses, including SARS-CoV-2 (PubMed:32855215). Disrupts cathepsin-mediated Ebola virus glycoprotein processing, which prevents viral fusion and entry. This antiviral activity is specific to p41 isoform (PubMed:32855215).By similarity1 Publication

<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

Complete GO annotation on QuickGO ...

GO - Biological processi

Complete GO annotation on QuickGO ...

<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

Molecular functionChaperone
Biological processAdaptive immunity, Immunity

Enzyme and pathway databases

Pathway Commons web resource for biological pathway data

More...PathwayCommonsi		P04233

Reactome - a knowledgebase of biological pathways and processes

More...Reactomei		R-HSA-202733, Cell surface interactions at the vascular wall
R-HSA-2132295, MHC class II antigen presentation

SIGNOR Signaling Network Open Resource

More...SIGNORi		P04233

Protein family/group databases

MEROPS protease database

More...MEROPSi		I31.002

Transport Classification Database

More...TCDBi		9.A.75.1.1, the mhc ii receptor (mhc2r) family

<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:
HLA class II histocompatibility antigen gamma chainCurated
Alternative name(s):
HLA-DR antigens-associated invariant chain
Ia antigen-associated invariant chain
Short name:
Ii
CD_antigen: CD74
Cleaved into the following chain:
Class-II-associated invariant chain peptide1 Publication
Short name:
CLIP1 Publication
<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:CD74Imported
Synonyms:DHLAG
<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>OrganismiHomo sapiens (Human)
<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 identifieri9606 [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 lineageiEukaryota › Metazoa › Chordata › Craniata › Vertebrata › Euteleostomi › Mammalia › Eutheria › Euarchontoglires › Primates › Haplorrhini › Catarrhini › Hominidae › Homo
<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
  • UP000005640 <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 5

Organism-specific databases

Human Gene Nomenclature Database

More...HGNCi		HGNC:1697, CD74

Online Mendelian Inheritance in Man (OMIM)

More...MIMi		142790, gene

neXtProt; the human protein knowledge platform

More...neXtProti		NX_P04233

Eukaryotic Pathogen, Vector and Host Database Resources

More...VEuPathDBi		HostDB:ENSG00000019582.14

<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

Extracellular region or secreted Cytosol Plasma membrane Cytoskeleton Lysosome Endosome Peroxisome ER Golgi apparatus Nucleus Mitochondrion Manual annotation Automatic computational assertionGraphics by Christian Stolte & Seán O’Donoghue; Source: COMPARTMENTS

Topology

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/help/subcellular%5Flocation%5Fsection">'Subcellular location'</a> section describes the subcellular compartment where each non-membrane region of a membrane-spanning protein is found.<p><a href='/help/topo_dom' target='_top'>More...</a></p>Topological domaini1 – 46CytoplasmicSequence analysisAdd BLAST46
<p>This subsection of the <a href="http://www.uniprot.org/help/subcellular%5Flocation%5Fsection">'Subcellular location'</a> section describes the extent of a membrane-spanning region of the protein. It denotes the presence of both alpha-helical transmembrane regions and the membrane spanning regions of beta-barrel transmembrane proteins.<p><a href='/help/transmem' target='_top'>More...</a></p>Transmembranei47 – 72Helical; Signal-anchor for type II membrane proteinSequence analysisAdd BLAST26
Topological domaini73 – 296ExtracellularSequence analysisAdd BLAST224

Keywords - Cellular componenti

Cell membrane, Endoplasmic reticulum, Endosome, Golgi apparatus, Lysosome, Membrane

<p>This section provides information on the disease(s) and phenotype(s) associated with a protein.<p><a href='/help/pathology_and_biotech_section' target='_top'>More...</a></p>Pathology & Biotechi

<p>This subsection of the 'Pathology and Biotech' section provides information on the disease(s) associated with genetic variations in a given protein. The information is extracted from the scientific literature and diseases that are also described in the <a href="http://www.ncbi.nlm.nih.gov/sites/entrez?db=omim">OMIM</a> database are represented with a <a href="http://www.uniprot.org/diseases">controlled vocabulary</a> in the following way:<p><a href='/help/involvement_in_disease' target='_top'>More...</a></p>Involvement in diseasei

A chromosomal aberration involving CD74 is found in a non-small cell lung tumor. Results in the formation of a CD74-ROS1 chimeric protein.1 Publication

Mutagenesis

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the <a href="http://www.uniprot.org/manual/pathology%5Fand%5Fbiotech%5Fsection">'Pathology and Biotech'</a> section describes the effect of the experimental mutation of one or more amino acid(s) on the biological properties of the protein.<p><a href='/help/mutagen' target='_top'>More...</a></p>Mutagenesisi225 – 226PG → RR or DD: Decreases inhibition of Ebola virus infection. 1 Publication2
Mutagenesisi225 – 226PG → TT: No effect on inhibition of Ebola virus infection. 1 Publication2

Sites

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection describes interesting single amino acid sites on the sequence that are not defined in any other subsection. This subsection can be displayed in different sections ('Function', 'PTM / Processing', 'Pathology and Biotech') according to its content.<p><a href='/help/site' target='_top'>More...</a></p>Sitei208 – 209Breakpoint for translocation to form a CD74-ROS1 fusion protein2

Organism-specific databases

DisGeNET

More...DisGeNETi		972

Open Targets

More...OpenTargetsi		ENSG00000019582

The Pharmacogenetics and Pharmacogenomics Knowledge Base

More...PharmGKBi		PA26236

Miscellaneous databases

Pharos NIH Druggable Genome Knowledgebase

More...Pharosi		P04233, Tchem

Chemistry databases

ChEMBL database of bioactive drug-like small molecules

More...ChEMBLi		CHEMBL4692

IUPHAR/BPS Guide to PHARMACOLOGY

More...GuidetoPHARMACOLOGYi		2840

Genetic variation databases

BioMuta curated single-nucleotide variation and disease association database

More...BioMutai		CD74

Domain mapping of disease mutations (DMDM)

More...DMDMi		20178292

<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 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_00000679541 – 296HLA class II histocompatibility antigen gamma chainAdd BLAST296
<p>This subsection of the 'PTM / Processing' section describes the position and length of an active peptide in the mature protein.<p><a href='/help/peptide' target='_top'>More...</a></p>PeptideiPRO_000044888697 – 120Class-II-associated invariant chain peptide1 PublicationAdd BLAST24

Amino acid modifications

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'PTM / Processing' section specifies the position and type of each modified residue excluding <a href="http://www.uniprot.org/manual/lipid">lipids</a>, <a href="http://www.uniprot.org/manual/carbohyd">glycans</a> and <a href="http://www.uniprot.org/manual/crosslnk">protein cross-links</a>.<p><a href='/help/mod_res' target='_top'>More...</a></p>Modified residuei25PhosphoserineBy similarity1
<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>Glycosylationi130N-linked (GlcNAc...) asparagine1
Glycosylationi136N-linked (GlcNAc...) asparagine1 Publication1
Glycosylationi203O-linked (GalNAc...) threonine2 Publications1
<p>This subsection of the PTM / Processing":/help/ptm_processing_section section describes the positions of cysteine residues participating in disulfide bonds.<p><a href='/help/disulfid' target='_top'>More...</a></p>Disulfide bondi213 ↔ 232PROSITE-ProRule annotation1 Publication
Disulfide bondi243 ↔ 250PROSITE-ProRule annotation1 Publication
Disulfide bondi252 ↔ 271PROSITE-ProRule annotation1 Publication
Glycosylationi256N-linked (GlcNAc...) asparagine1 Publication1
Glycosylationi281O-linked (GalNAc...) serine1 Publication1

<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

N- and O-glycosylated. O-glycosylated with core 1 or possibly core 8 glycans.4 Publications

Keywords - PTMi

Disulfide bond, Glycoprotein, Phosphoprotein, Proteoglycan

Proteomic databases

jPOST - Japan Proteome Standard Repository/Database

More...jPOSTi		P04233

MassIVE - Mass Spectrometry Interactive Virtual Environment

More...MassIVEi		P04233

MaxQB - The MaxQuant DataBase

More...MaxQBi		P04233

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

More...PaxDbi		P04233

PeptideAtlas

More...PeptideAtlasi		P04233

PRoteomics IDEntifications database

More...PRIDEi		P04233

ProteomicsDB: a multi-organism proteome resource

More...ProteomicsDBi		51689 [P04233-1]
51690 [P04233-2]
51691 [P04233-3]

PTM databases

GlyConnect protein glycosylation platform

More...GlyConnecti		659, 9 N-Linked glycans (2 sites), 1 O-Linked glycan (1 site)

GlyGen: Computational and Informatics Resources for Glycoscience

More...GlyGeni		P04233, 9 sites, 1 O-linked glycan (4 sites)

iPTMnet integrated resource for PTMs in systems biology context

More...iPTMneti		P04233

Comprehensive resource for the study of protein post-translational modifications (PTMs) in human, mouse and rat.

More...PhosphoSitePlusi		P04233

SwissPalm database of S-palmitoylation events

More...SwissPalmi		P04233

<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

<p>This subsection of the 'Expression' section provides information on the expression of a gene at the mRNA or protein level in cells or in tissues of multicellular organisms. By default, the information is derived from experiments at the mRNA level, unless specified 'at protein level'.<br></br>Examples: <a href="http://www.uniprot.org/uniprot/P92958#expression">P92958</a>, <a href="http://www.uniprot.org/uniprot/Q8TDN4#expression">Q8TDN4</a>, <a href="http://www.uniprot.org/uniprot/O14734#expression">O14734</a><p><a href='/help/tissue_specificity' target='_top'>More...</a></p>Tissue specificityi

In B cells, represents 10% of total CD74 expression.1 Publication
In B cells, represents 70% of total CD74 expression.1 Publication

Gene expression databases

Bgee dataBase for Gene Expression Evolution

More...Bgeei		ENSG00000019582, Expressed in spleen and 246 other tissues

ExpressionAtlas, Differential and Baseline Expression

More...ExpressionAtlasi		P04233, baseline and differential

Genevisible search portal to normalized and curated expression data from Genevestigator

More...Genevisiblei		P04233, HS

Organism-specific databases

Human Protein Atlas

More...HPAi		ENSG00000019582, Tissue enhanced (blood)

<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

Homotrimer. In the endoplasmic reticulum (ER) it forms a heterononameric MHC II-Ii complex: 3 MHC class II molecules (heterodimers of an alpha and a beta subunit) bind to the CD74 homotrimer (also known as invariant chain or HLA class II histocompatibility antigen gamma chain). In the endosomal/lysosomal system, the CD74 component undergoes sequential degradation by various proteases, including CTSS and CTSL, leaving a small fragment termed CLIP (class-II-associated invariant chain peptide) attached to the MHC class II molecule (alpha-beta-CLIP complex). This processed complex interacts with HLA_DM and HLA_DO heterodimers in order to release CLIP and facilitate the binding of antigenic peptides to the MHC class II molecules.

Interacts with CD44; this complex is essential for the MIF-induced signaling cascade that results in B cell survival.

By similarity2 Publications

Interacts with the mature form of CTSL; the complex survive in neutral pH environment.

1 Publication

<p>This subsection of the '<a href="http://www.uniprot.org/help/interaction%5Fsection">Interaction</a>' section provides information about binary protein-protein interactions. The data presented in this section are a quality-filtered subset of binary interactions automatically derived from the <a href="https://www.ebi.ac.uk/intact/">IntAct database</a>. It is updated at every <a href="http://www.uniprot.org/help/synchronization">UniProt release</a>.<p><a href='/help/binary_interactions' target='_top'>More...</a></p>Binary interactionsi

Hide details
P04233
With#Exp.IntAct
CD44 [P16070]9EBI-2622890,EBI-490245
CXCR2 [P25025]2EBI-2622890,EBI-2835281
CXCR4 [P61073]4EBI-2622890,EBI-489411
EIF3E [P60228]2EBI-2622890,EBI-347740
HSPA1A [P0DMV8]4EBI-2622890,EBI-11052499
KPNB1 [Q14974]2EBI-2622890,EBI-286758
dnaK [P0A6Y8] from Escherichia coli (strain K12).8EBI-2622890,EBI-542092
Isoform p35 [P04233-2]
With#Exp.IntAct
BET1 [O15155]3EBI-12222807,EBI-749204
CLDN19 - isoform 2 [Q8N6F1-2]3EBI-12222807,EBI-12256978
CLEC7A - isoform 6 [Q9BXN2-6]3EBI-12222807,EBI-11989440
CMTM7 [Q96FZ5]3EBI-12222807,EBI-2807956
EDDM3B [P56851]3EBI-12222807,EBI-10215665
ERG28 [Q9UKR5]3EBI-12222807,EBI-711490
FDFT1 [P37268]3EBI-12222807,EBI-714550
FXYD6 [Q9H0Q3]3EBI-12222807,EBI-713304
LPAR3 [Q9UBY5]4EBI-12222807,EBI-12033434
NRAC [Q8N912]4EBI-12222807,EBI-12051377
PGA4 [P0DJD7]3EBI-12222807,EBI-12957629
PLP1 - isoform DM-20 [P60201-2]3EBI-12222807,EBI-12188331
PLPP4 [Q5VZY2]3EBI-12222807,EBI-10485931
POMGNT1 [Q8WZA1]3EBI-12222807,EBI-3912424
PTCH1 - isoform M [Q13635-3]3EBI-12222807,EBI-14199621
RTP2 [Q5QGT7]3EBI-12222807,EBI-10244780
SEC22A [Q96IW7]3EBI-12222807,EBI-8652744
SERP1 [Q9Y6X1]3EBI-12222807,EBI-10329948
SLC35B1 [P78383]3EBI-12222807,EBI-12147661
SMIM1 [B2RUZ4]3EBI-12222807,EBI-12188413
TEX11 - isoform 3 [Q8IYF3-3]3EBI-12222807,EBI-11523345
TMEM120B [A0PK00]3EBI-12222807,EBI-10171534
TMEM243 [Q9BU79]3EBI-12222807,EBI-12887458
TMEM254 [Q8TBM7]3EBI-12222807,EBI-11956809
TMEM60 [Q9H2L4]3EBI-12222807,EBI-2852148
TMEM97 [Q5BJF2]3EBI-12222807,EBI-12111910
UPK2 [O00526]4EBI-12222807,EBI-10179682

GO - Molecular functioni

Complete GO annotation on QuickGO ...

Protein-protein interaction databases

The Biological General Repository for Interaction Datasets (BioGRID)

More...BioGRIDi		107410, 138 interactors

The Eukaryotic Linear Motif resource for Functional Sites in Proteins

More...ELMi		P04233

Protein interaction database and analysis system

More...IntActi		P04233, 200 interactors

Molecular INTeraction database

More...MINTi		P04233

STRING: functional protein association networks

More...STRINGi		9606.ENSP00000009530

Chemistry databases

BindingDB database of measured binding affinities

More...BindingDBi		P04233

Miscellaneous databases

RNAct, Protein-RNA interaction predictions for model organisms.

More...RNActi		P04233, protein

<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

Secondary structure

1296
Legend: HelixTurnBeta strandPDB Structure known for this area
Show more details

3D structure databases

Biological Magnetic Resonance Data Bank

More...BMRBi		P04233

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

More...SMRi		P04233

Database of comparative protein structure models

More...ModBasei		Search...

Protein Data Bank in Europe - Knowledge Base

More...PDBe-KBi		Search...

Miscellaneous databases

Relative evolutionary importance of amino acids within a protein sequence

More...EvolutionaryTracei		P04233

<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>Domaini210 – 271Thyroglobulin type-1PROSITE-ProRule annotationAdd BLAST62

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>Regioni225 – 226Required for interaction with CTSL2 Publications2

<p>This subsection of the 'Family and domains' section provides general information on the biological role of a domain. The term 'domain' is intended here in its wide acceptation, it may be a structural domain, a transmembrane region or a functional domain. Several domains are described in this subsection.<p><a href='/help/domain_cc' target='_top'>More...</a></p>Domaini

Antiviral activity requires delivery of the thyroglobulin domain to the endosomal membrane.1 Publication

Keywords - Domaini

Signal-anchor, Transmembrane, Transmembrane helix

Phylogenomic databases

evolutionary genealogy of genes: Non-supervised Orthologous Groups

More...eggNOGi		KOG1214, Eukaryota

Ensembl GeneTree

More...GeneTreei		ENSGT00390000008961

InParanoid: Eukaryotic Ortholog Groups

More...InParanoidi		P04233

Database of Orthologous Groups

More...OrthoDBi		1190626at2759

Database for complete collections of gene phylogenies

More...PhylomeDBi		P04233

TreeFam database of animal gene trees

More...TreeFami		TF317779

Family and domain databases

Conserved Domains Database

More...CDDi		cd00191, TY, 1 hit

Gene3D Structural and Functional Annotation of Protein Families

More...Gene3Di		1.10.870.10, 1 hit
4.10.800.10, 1 hit

Integrated resource of protein families, domains and functional sites

More...InterProi		View protein in InterPro
IPR043530, CD74_antigen
IPR015386, MHC_II-assoc_invar/CLIP_MHC-bd
IPR022339, MHC_II-assoc_invar_chain
IPR011988, MHC_II-assoc_invariant_trimer
IPR036613, MHCII_invariant_trimer_sf
IPR000716, Thyroglobulin_1
IPR036857, Thyroglobulin_1_sf

Pfam protein domain database

More...Pfami		View protein in Pfam
PF09307, MHC2-interact, 1 hit
PF08831, MHCassoc_trimer, 1 hit
PF00086, Thyroglobulin_1, 1 hit

PIRSF; a whole-protein classification database

More...PIRSFi		PIRSF001992, CD74_antigen, 1 hit

Protein Motif fingerprint database; a protein domain database

More...PRINTSi		PR01990, CD74ANTIGEN

Simple Modular Architecture Research Tool; a protein domain database

More...SMARTi		View protein in SMART
SM00211, TY, 1 hit

Superfamily database of structural and functional annotation

More...SUPFAMi		SSF48305, SSF48305, 1 hit
SSF57610, SSF57610, 1 hit

PROSITE; a protein domain and family database

More...PROSITEi		View protein in PROSITE
PS00484, THYROGLOBULIN_1_1, 1 hit
PS51162, THYROGLOBULIN_1_2, 1 hit

<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>Sequences (5+)i

<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.

This entry describes 5 <p>This subsection of the 'Sequence' section lists the alternative protein sequences (isoforms) that can be generated from the same gene by a single or by the combination of up to four biological events (alternative promoter usage, alternative splicing, alternative initiation and ribosomal frameshifting). Additionally, this section gives relevant information on each alternative protein isoform. This section is only present in reviewed entries, i.e. in UniProtKB/Swiss-Prot.<p><a href='/help/alternative_products' target='_top'>More...</a></p> isoformsi produced by alternative splicing and alternative initiation. AlignAdd to basket

This entry has 5 described isoforms and 5 potential isoforms that are computationally mapped.Show allAlign All

Isoform p451 Publication (identifier: P04233-1) [UniParc]FASTAAdd to basket
Also known as: Long

This isoform has been chosen as the <div> <p><b>What is the canonical sequence?</b><p><a href='/help/canonical_and_isoforms' target='_top'>More...</a></p>canonicali sequence. All positional information in this entry refers to it. This is also the sequence that appears in the downloadable versions of the entry.

« Hide
        10         20         30         40         50
MHRRRSRSCR EDQKPVMDDQ RDLISNNEQL PMLGRRPGAP ESKCSRGALY 
        60         70         80         90        100
TGFSILVTLL LAGQATTAYF LYQQQGRLDK LTVTSQNLQL ENLRMKLPKP 
       110        120        130        140        150
PKPVSKMRMA TPLLMQALPM GALPQGPMQN ATKYGNMTED HVMHLLQNAD 
       160        170        180        190        200
PLKVYPPLKG SFPENLRHLK NTMETIDWKV FESWMHHWLL FEMSRHSLEQ 
       210        220        230        240        250
KPTDAPPKVL TKCQEEVSHI PAVHPGSFRP KCDENGNYLP LQCYGSIGYC 
       260        270        280        290 
WCVFPNGTEV PNTRSRGHHN CSESLELEDP SSGLGVTKQD LGPVPM
Length:296
Mass (Da):33,516
Last modified:April 16, 2002 - v3
<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:i27A13F252D5FB91D
GO
Isoform p351 Publication (identifier: P04233-2) [UniParc]FASTAAdd to basket
Also known as: Short

The sequence of this isoform differs from the canonical sequence as follows:
     209-272: Missing.

Show »
Length:232
Mass (Da):26,399
Checksum:i5144439D0FD27C99
GO
Isoform 3 (identifier: P04233-3) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     148-160: NADPLKVYPPLKG → SHWNWRTRLLGWV
     161-296: Missing.

Show »
Length:160
Mass (Da):18,328
Checksum:i95CB3D7226497DE8
GO
Isoform p411 Publication (identifier: P04233-4) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-16: Missing.

Show »
Length:280
Mass (Da):31,492
Checksum:i5BAC4D0AB16A8EDD
GO
Isoform p331 Publication (identifier: P04233-5) [UniParc]FASTAAdd to basket

The sequence of this isoform differs from the canonical sequence as follows:
     1-16: Missing.
     209-272: Missing.

Show »
Length:216
Mass (Da):24,375
Checksum:i7EE2DB296EB80DFD
GO

<p>In eukaryotic reference proteomes, unreviewed entries that are likely to belong to the same gene are computationally mapped, based on gene identifiers from Ensembl, EnsemblGenomes and model organism databases.<p><a href='/help/gene_centric_isoform_mapping' target='_top'>More...</a></p>Computationally mapped potential isoform sequencesi

There are 5 potential isoforms mapped to this entry.BLASTAlignShow allAdd to basket
EntryEntry nameProtein names
Gene namesLengthAnnotation
H0YBZ2H0YBZ2_HUMAN
HLA class II histocompatibility ant...
CD74
316Annotation score:

Annotation score:5 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>
H0YBZ8H0YBZ8_HUMAN
HLA class II histocompatibility ant...
CD74
145Annotation score:

Annotation score:2 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>
E7ESL3E7ESL3_HUMAN
HLA class II histocompatibility ant...
CD74
152Annotation score:

Annotation score:2 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>
E7EQJ3E7EQJ3_HUMAN
HLA class II histocompatibility ant...
CD74
122Annotation score:

Annotation score:1 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>
E7ER71E7ER71_HUMAN
HLA class II histocompatibility ant...
CD74
52Annotation score:

Annotation score:1 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>

<p>This subsection of the 'Sequence' section reports difference(s) between the protein sequence shown in the UniProtKB entry and other available protein sequences derived from the same gene.<p><a href='/help/sequence_caution' target='_top'>More...</a></p>Sequence cautioni

The sequence AAA36304 differs from that shown. Reason: Erroneous initiation. Truncated N-terminus.Curated

Experimental Info

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'Sequence' section reports difference(s) between the canonical sequence (displayed by default in the entry) and the different sequence submissions merged in the entry. These various submissions may originate from different sequencing projects, different types of experiments, or different biological samples. Sequence conflicts are usually of unknown origin.<p><a href='/help/conflict' target='_top'>More...</a></p>Sequence conflicti167R → T in CAA27047 (PubMed:3001652).Curated1

Alternative sequence

Feature keyPosition(s)DescriptionActionsGraphical viewLength
<p>This subsection of the 'Sequence' section describes the sequence of naturally occurring alternative protein isoform(s). The changes in the amino acid sequence may be due to alternative splicing, alternative promoter usage, alternative initiation, or ribosomal frameshifting.<p><a href='/help/var_seq' target='_top'>More...</a></p>Alternative sequenceiVSP_0609041 – 16Missing in isoform p41 and isoform p33. 1 PublicationAdd BLAST16
Alternative sequenceiVSP_037869148 – 160NADPL…PPLKG → SHWNWRTRLLGWV in isoform 3. 1 PublicationAdd BLAST13
Alternative sequenceiVSP_037870161 – 296Missing in isoform 3. 1 PublicationAdd BLAST136
Alternative sequenceiVSP_005331209 – 272Missing in isoform p35 and isoform p33. 5 PublicationsAdd BLAST64

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
K01144 mRNA Translation: AAA36304.1 Different initiation.
X00497 mRNA Translation: CAA25192.1
X00497 mRNA Translation: CAA25193.1
X03339 Genomic DNA Translation: CAA27046.1
X03340 Genomic DNA Translation: CAA27047.1
M13560 M13559 Genomic DNA Translation: AAA36033.1
BT019505 mRNA Translation: AAV38312.1
AK292076 mRNA Translation: BAF84765.1
AK297889 mRNA Translation: BAG60210.1
AC011372 Genomic DNA No translation available.
AC011388 Genomic DNA No translation available.
CH471062 Genomic DNA Translation: EAW61727.1
CH471062 Genomic DNA Translation: EAW61728.1
CH471062 Genomic DNA Translation: EAW61729.1
CH471062 Genomic DNA Translation: EAW61730.1
CH471062 Genomic DNA Translation: EAW61731.1
BC018726 mRNA Translation: AAH18726.1
BC024272 mRNA Translation: AAH24272.1

The Consensus CDS (CCDS) project

More...CCDSi		CCDS34276.1 [P04233-3]
CCDS47308.1 [P04233-2]
CCDS47309.1 [P04233-1]

Protein sequence database of the Protein Information Resource

More...PIRi		A93981, HLHUG

NCBI Reference Sequences

More...RefSeqi		NP_001020329.1, NM_001025158.2 [P04233-3]
NP_001020330.1, NM_001025159.2 [P04233-1]
NP_004346.1, NM_004355.3 [P04233-2]

Genome annotation databases

Ensembl eukaryotic genome annotation project

More...Ensembli		ENST00000009530; ENSP00000009530; ENSG00000019582 [P04233-1]
ENST00000353334; ENSP00000230685; ENSG00000019582 [P04233-2]
ENST00000377795; ENSP00000367026; ENSG00000019582 [P04233-3]

Database of genes from NCBI RefSeq genomes

More...GeneIDi		972

KEGG: Kyoto Encyclopedia of Genes and Genomes

More...KEGGi		hsa:972

UCSC genome browser

More...UCSCi		uc003lsc.4, human [P04233-1]

Keywords - Coding sequence diversityi

Alternative initiation, Alternative splicing

<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
K01144 mRNA Translation: AAA36304.1 Different initiation.
X00497 mRNA Translation: CAA25192.1
X00497 mRNA Translation: CAA25193.1
X03339 Genomic DNA Translation: CAA27046.1
X03340 Genomic DNA Translation: CAA27047.1
M13560 M13559 Genomic DNA Translation: AAA36033.1
BT019505 mRNA Translation: AAV38312.1
AK292076 mRNA Translation: BAF84765.1
AK297889 mRNA Translation: BAG60210.1
AC011372 Genomic DNA No translation available.
AC011388 Genomic DNA No translation available.
CH471062 Genomic DNA Translation: EAW61727.1
CH471062 Genomic DNA Translation: EAW61728.1
CH471062 Genomic DNA Translation: EAW61729.1
CH471062 Genomic DNA Translation: EAW61730.1
CH471062 Genomic DNA Translation: EAW61731.1
BC018726 mRNA Translation: AAH18726.1
BC024272 mRNA Translation: AAH24272.1
CCDSiCCDS34276.1 [P04233-3]
CCDS47308.1 [P04233-2]
CCDS47309.1 [P04233-1]
PIRiA93981, HLHUG
RefSeqiNP_001020329.1, NM_001025158.2 [P04233-3]
NP_001020330.1, NM_001025159.2 [P04233-1]
NP_004346.1, NM_004355.3 [P04233-2]

3D structure databases

Select the link destinations:

Protein Data Bank Europe

More...PDBei

Protein Data Bank RCSB

More...RCSB PDBi

Protein Data Bank Japan

More...PDBji
Links Updated
PDB entryMethodResolution (Å)ChainPositionsPDBsum
1A6AX-ray2.75C103-117[»]
1ICFX-ray2.00I/J210-274[»]
1IIENMR-A/B/C134-208[»]
1L3HNMR-A210-274[»]
1MUJX-ray2.15C97-121[»]
3PDOX-ray1.95C102-120[»]
3PGCX-ray2.66C/F106-120[»]
3PGDX-ray2.72C/F106-120[»]
3QXAX-ray2.71C/F103-117[»]
3QXDX-ray2.30C/F103-117[»]
4AENX-ray2.20C106-120[»]
4AH2X-ray2.36B106-120[»]
4X5WX-ray1.34C102-120[»]
5KSUX-ray2.73C/F103-117[»]
5KSVX-ray2.19C109-123[»]
BMRBiP04233
SMRiP04233
ModBaseiSearch...
PDBe-KBiSearch...

Protein-protein interaction databases

BioGRIDi107410, 138 interactors
ELMiP04233
IntActiP04233, 200 interactors
MINTiP04233
STRINGi9606.ENSP00000009530

Chemistry databases

BindingDBiP04233
ChEMBLiCHEMBL4692
GuidetoPHARMACOLOGYi2840

Protein family/group databases

MEROPSiI31.002
TCDBi9.A.75.1.1, the mhc ii receptor (mhc2r) family

PTM databases

GlyConnecti659, 9 N-Linked glycans (2 sites), 1 O-Linked glycan (1 site)
GlyGeniP04233, 9 sites, 1 O-linked glycan (4 sites)
iPTMnetiP04233
PhosphoSitePlusiP04233
SwissPalmiP04233

Genetic variation databases

BioMutaiCD74
DMDMi20178292

Proteomic databases

jPOSTiP04233
MassIVEiP04233
MaxQBiP04233
PaxDbiP04233
PeptideAtlasiP04233
PRIDEiP04233
ProteomicsDBi51689 [P04233-1]
51690 [P04233-2]
51691 [P04233-3]

Protocols and materials databases

ABCD curated depository of sequenced antibodies

More...ABCDi		P04233, 19 sequenced antibodies

Antibodypedia a portal for validated antibodies

More...Antibodypediai		2245, 2129 antibodies

The CPTC Antibody Portal

More...CPTCi		P04233, 1 antibody

The DNASU plasmid repository

More...DNASUi		972

Genome annotation databases

EnsembliENST00000009530; ENSP00000009530; ENSG00000019582 [P04233-1]
ENST00000353334; ENSP00000230685; ENSG00000019582 [P04233-2]
ENST00000377795; ENSP00000367026; ENSG00000019582 [P04233-3]
GeneIDi972
KEGGihsa:972
UCSCiuc003lsc.4, human [P04233-1]

Organism-specific databases

Comparative Toxicogenomics Database

More...CTDi		972
DisGeNETi972

GeneCards: human genes, protein and diseases

More...GeneCardsi		CD74
HGNCiHGNC:1697, CD74
HPAiENSG00000019582, Tissue enhanced (blood)
MIMi142790, gene
neXtProtiNX_P04233
OpenTargetsiENSG00000019582
PharmGKBiPA26236
VEuPathDBiHostDB:ENSG00000019582.14

GenAtlas: human gene database

More...GenAtlasi		Search...

Phylogenomic databases

eggNOGiKOG1214, Eukaryota
GeneTreeiENSGT00390000008961
InParanoidiP04233
OrthoDBi1190626at2759
PhylomeDBiP04233
TreeFamiTF317779

Enzyme and pathway databases

PathwayCommonsiP04233
ReactomeiR-HSA-202733, Cell surface interactions at the vascular wall
R-HSA-2132295, MHC class II antigen presentation
SIGNORiP04233

Miscellaneous databases

BioGRID ORCS database of CRISPR phenotype screens

More...BioGRID-ORCSi		972, 3 hits in 875 CRISPR screens

ChiTaRS: a database of human, mouse and fruit fly chimeric transcripts and RNA-sequencing data

More...ChiTaRSi		CD74, human
EvolutionaryTraceiP04233

The Gene Wiki collection of pages on human genes and proteins

More...GeneWikii		CD74

Database of phenotypes from RNA interference screens in Drosophila and Homo sapiens

More...GenomeRNAii		972
PharosiP04233, Tchem

Protein Ontology

More...PROi		PR:P04233
RNActiP04233, protein

The Stanford Online Universal Resource for Clones and ESTs

More...SOURCEi		Search...

Gene expression databases

BgeeiENSG00000019582, Expressed in spleen and 246 other tissues
ExpressionAtlasiP04233, baseline and differential
GenevisibleiP04233, HS

Family and domain databases

CDDicd00191, TY, 1 hit
Gene3Di1.10.870.10, 1 hit
4.10.800.10, 1 hit
InterProiView protein in InterPro
IPR043530, CD74_antigen
IPR015386, MHC_II-assoc_invar/CLIP_MHC-bd
IPR022339, MHC_II-assoc_invar_chain
IPR011988, MHC_II-assoc_invariant_trimer
IPR036613, MHCII_invariant_trimer_sf
IPR000716, Thyroglobulin_1
IPR036857, Thyroglobulin_1_sf
PfamiView protein in Pfam
PF09307, MHC2-interact, 1 hit
PF08831, MHCassoc_trimer, 1 hit
PF00086, Thyroglobulin_1, 1 hit
PIRSFiPIRSF001992, CD74_antigen, 1 hit
PRINTSiPR01990, CD74ANTIGEN
SMARTiView protein in SMART
SM00211, TY, 1 hit
SUPFAMiSSF48305, SSF48305, 1 hit
SSF57610, SSF57610, 1 hit
PROSITEiView protein in PROSITE
PS00484, THYROGLOBULIN_1_1, 1 hit
PS51162, THYROGLOBULIN_1_2, 1 hit

ProtoNet; Automatic hierarchical classification of proteins

More...ProtoNeti		Search...

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 nameiHG2A_HUMAN
<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: P04233
Secondary accession number(s): A8K7R1
, B4DNE8, D3DQG3, D3DQG4, Q14597, Q29832, Q5U0J8, Q8SNA0, Q8WLP6
<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: March 20, 1987
Last sequence update: April 16, 2002
Last modified: February 10, 2021
This is version 225 of the entry and version 3 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
DisclaimerAny medical or genetic information present in this entry is provided for research, educational and informational purposes only. It is not in any way intended to be used as a substitute for professional medical advice, diagnosis, treatment or care.

<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

3D-structure, Direct protein sequencing, Reference proteome

Documents

  1. Human cell differentiation molecules
    CD nomenclature of surface proteins of human leucocytes and list of entries
  2. Human chromosome 5
    Human chromosome 5: entries, gene names and cross-references to MIM
  3. MIM cross-references
    Online Mendelian Inheritance in Man (MIM) cross-references in UniProtKB/Swiss-Prot
  4. PDB cross-references
    Index of Protein Data Bank (PDB) cross-references
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