Human leukocyte antigen - Knowledge (XXG)

1125:(SFVT) approach for HLA genetic analysis that categorizes HLA proteins into biologically relevant smaller sequence features (SFs), and their variant types (VTs). Sequence features are combinations of amino acid sites defined based on structural information (e.g., beta-sheet 1), functional information (e.g., peptide antigen-binding), and polymorphism. These sequence features can be overlapping and continuous or discontinuous in the linear sequence. Variant types for each sequence feature are defined based upon all known polymorphisms in the HLA locus being described. SFVT categorization of HLA is applied in genetic association analysis so that the effects and roles of the epitopes shared by several HLA alleles can be identified. Sequence features and their variant types have been described for all classical HLA proteins; the international repository of HLA SFVTs will be maintained at IMGT/HLA database. A tool to convert HLA alleles into their component SFVTs can be found on the Immunology Database and Analysis Portal (ImmPort) website. 1650:

receptors and receptor isoforms. Over the years, serotyping antibodies became more refined as techniques for increasing sensitivity improved and new serotyping antibodies continue to appear. One of the goals of serotype analysis is to fill gaps in the analysis. It is possible to predict based on 'square root','maximum-likelihood' method, or analysis of familial haplotypes to account for adequately typed alleles. These studies using serotyping techniques frequently revealed, in particular for non-European or north East Asian populations many null or blank serotypes. This was particularly problematic for the Cw locus until recently, and almost half of the Cw serotypes went untyped in the 1991 survey of the human population.

395:. T-cells have receptors that are similar to B-cell receptors, and each T-cell recognizes only a few MHC class II-peptide combinations. Once a T-cell recognizes a peptide within an MHC class II molecule, it can stimulate B-cells that also recognize the same molecule in their B-cell receptors. Thus, T-cells help B-cells make antibodies to the same foreign antigens. Each HLA can bind many peptides, and each person has 3 HLA types and can have 4 isoforms of DP, 4 isoforms of DQ and 4 Isoforms of DR (2 of DRB1, and 2 of DRB3, DRB4, or DRB5) for a total of 12 isoforms. In such heterozygotes, it is difficult for disease-related proteins to escape detection. 849:

could not completely distinguish alleles and so stopped at this level. The third through fourth digits specify a nonsynonymous allele. Digits five through six denote any synonymous mutations within the coding frame of the gene. The seventh and eighth digits distinguish mutations outside the coding region. Letters such as L, N, Q, or S may follow an allele's designation to specify an expression level or other non-genomic data known about it. Thus, a completely described allele may be up to 9 digits long, not including the HLA-prefix and locus notation.

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designated as Dw types. Serotyped DR1 has cellularly defined as either of Dw1 or of Dw20 and so on for other serotyped DRs. Table shows associated cellular specificities for DR alleles. However, cellular typing has inconsistency in the reaction between cellular-type individuals, sometimes resulting differently from predicted. Together with difficulty of cellular assay in generating and maintaining cellular typing reagents, cellular assay is being replaced by DNA-based typing method.

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receptors are capable of binding an almost endless variation of peptides of 9 amino acids or longer in length, protecting interbreeding subpopulations from nascent or epidemic diseases. Individuals in a population frequently have different haplotypes, and this results in many combinations, even in small groups. This diversity enhances the survival of such groups, and thwarts evolution of epitopes in pathogens, which would otherwise be able to be shielded from the immune system.

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have fewer alleles, combinations of A1:B1 can produce a theoretical potential of 7,755 DQ and 5,270 DP αβ heterodimers, respectively. While nowhere near this number of isoforms exist in the human population, each individual can carry 4 variable DQ and DP isoforms, increasing the potential number of antigens that these receptors can present to the immune system.

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been detected at least five times in unrelated individuals via the application of a sequence-based typing (SBT) method, or at least three times via a SBT method and in a specific haplotype in unrelated individuals. Rare alleles are defined as those that have been reported one to four times, and very rare alleles as those reported only once.

194:), all of which are the HLA Class1 group, present peptides from inside the cell. For example, if the cell is infected by a virus, the HLA system brings fragments of the virus to the surface of the cell so that the cell can be destroyed by the immune system. These peptides are produced from digested proteins that are broken down in the 1833:

the world where Western Europeans have migrated. The "A3-B7-DR2-DQ1" is more widely spread, from Eastern Asia to Iberia. The Super-B8 haplotype is associated with a number of diet-associated autoimmune diseases. There are 100,000s of extended haplotypes, but only a few show a visible and nodal character in the human population.

408:. Because of the importance of HLA in transplantation, the HLA loci are some of the most frequently typed by serology and PCR. It has been shown that high resolution HLA typing (HLA-A, HLA-B, HLA-C, HLA-DRB1, HLA-DQB1 and HLA-DPB1) may be relevant in transplantation to identify a full match, even when the donor is related. 91: 287:. They may protect against cancers or fail to protect (if down-regulated by an infection). HLA may also be related to people's perception of the odor of other people, and may be involved in mate selection, as at least one study found a lower-than-expected rate of HLA similarity between spouses in an isolated community. 679:

enteropathy-associated T-cell lymphoma cases. More often, however, HLA molecules play a protective role, recognizing increases in antigens that are not tolerated because of low levels in the normal state. Abnormal cells might be targeted for apoptosis, which is thought to mediate many cancers before diagnosis.

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variant of the nucleotide (DNA) sequence at a locus, such that each allele differs from all other alleles in at least one (single nucleotide polymorphism, SNP) position. Most of these changes result in a change in the amino acid sequences that result in slight to major functional differences in the protein.

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Studies of the variable positions of DP, DR, and DQ reveal that peptide antigen contact residues on class II molecules are most frequently the site of variation in the protein primary structure. Therefore, through a combination of intense allelic variation and/or subunit pairing, the class II peptide

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Studies of humans and animals imply a heterozygous selection mechanism operating on these loci as an explanation for this variability. One proposed mechanism is sexual selection in which females are able to detect males with different HLA relative to their own type. While the DQ and DP encoding loci

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For many populations, such as the Japanese or European populations, so many patients have been typed that new alleles are relatively rare, and thus SSP-PCR is more than adequate for allele resolution. Haplotypes can be obtained by typing family members in areas of the world where SSP-PCR is unable to

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There are several types of serotypes. A broad antigen serotype is a crude measure of identity of cells. For example, HLA A9 serotype recognizes cells of A23- and A24-bearing individuals. It may also recognize cells that A23 and A24 miss because of small variations. A23 and A24 are split antigens, but

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However, the gene frequencies of the most common alleles (>5%) of HLA-A, -B, -C and HLA-DPA1, -DPB1, -DQA1, -DQB1, and -DRB1 from South America have been reported from the typing and sequencing carried out in genetic diversity studies and cases and controls. In addition, information on the allele

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MHC loci are some of the most genetically variable coding loci in mammals, and the human HLA loci are no exceptions. Despite the fact that the human population went through a constriction several times during its history that was capable of fixing many loci, the HLA loci appear to have survived such

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Gene typing is different from gene sequencing and serotyping. With this strategy, PCR primers specific to a variant region of DNA are used (called sequence-specific primers). If a product of the right size is found, the assumption is that the HLA allele has been identified. New gene sequences often

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In the end, a workshop, based on sequence, decides which new allele goes into which serogroup either by sequence or by reactivity. Once the sequence is verified, it is assigned a number. For example, a new allele of B44 may get a serotype (i.e. B44) and allele ID i.e. B*44:65, as it is the 65th B44

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A representative cellular assay is the mixed lymphocyte culture (MLC) and used to determine the HLA class II types. The cellular assay is more sensitive in detecting HLA differences than serotyping. This is because minor differences unrecognized by alloantisera can stimulate T cells. This typing is

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Modern HLA alleles are typically noted with a variety of levels of detail. Most designations begin with HLA- and the locus name, then * and some (even) number of digits specifying the allele. The first two digits specify a group of alleles, also known as supertypes. Older typing methodologies often

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These haplotypes can be used to trace migrations in the human population because they are often much like a fingerprint of an event that has occurred in evolution. The Super-B8 haplotype is enriched in the Western Irish, declines along gradients away from that region, and is found only in areas of

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variation in the rate at which of individual HLA alleles are detected, attempts have been made to categorize alleles at each expressed HLA locus in terms of their prevalence. The result is a catalog of common and well-documented (CWD) HLA alleles, and a catalogue of rare and very rare HLA alleles.

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Although the number of individual HLA alleles that have been identified is large, approximately 40% of these alleles appear to be unique, having only been identified in single individuals. Roughly a third of alleles have been reported more than three times in unrelated individuals. Because of this

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In order to create a typing reagent, blood from animals or humans would be taken, the blood cells allowed to separate from the serum, and the serum diluted to its optimal sensitivity and used to type cells from other individuals or animals. Thus, serotyping became a way of crudely identifying HLA

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Common HLA alleles are defined as having been observed with a frequency of at least 0.001 in reference populations of at least 1500 individuals. Well-documented HLA alleles were originally defined as having been reported at least three times in unrelated individuals, and are now defined as having

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There are issues that limit this variation. Certain alleles like DQA1*05:01 and DQA1*05:05 encode proteins with identically processed products. Other alleles like DQB1*0201 and DQB1*0202 produce proteins that are functionally similar. For class II (DR, DP and DQ), amino acid variants within the

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Six loci have over 100 alleles that have been detected in the human population. Of these, the most variable are HLA B and HLA DRB1. As of 2012, the number of alleles that have been determined are listed in the table below. To interpret this table, it is necessary to consider that an allele is a

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Aside from the genes encoding the six major antigen-presenting proteins, many other genes, many involved in immune function, are located on the HLA complex. Diversity of HLAs in the human population is one aspect of disease defense, and, as a result, the chance of two unrelated individuals with

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Broad antigen types are still useful, such as typing very diverse populations with many unidentified HLA alleles (Africa, Arabia, Southeastern Iran and Pakistan, India). Africa, Southern Iran, and Arabia show the difficulty in typing areas that were settled earlier. Allelic diversity makes it

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Minor reactions to subregions that show similarity to other types can be observed to the gene products of alleles of a serotype group. The sequence of the antigens determines the antibody reactivities, and so having a good sequencing capability (or sequence-based typing) obviates the need for

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The large extent of variability in HLA genes poses significant challenges in investigating the role of HLA genetic variations in diseases. Disease association studies typically treat each HLA allele as a single complete unit, which does not illuminate the parts of the molecule associated with

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Donor-specific HLA antibodies have been found to be associated with graft failure in renal, heart, lung, and liver transplantation. These donor-specific HLA antibodies can exist pretransplant as consequence of sensitization to prior transplants or through pregnancies, but also occur de novo

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The image off to the side shows a piece of a poisonous bacterial protein (SEI peptide) bound within the binding cleft portion of the HLA-DR1 molecule. In the illustration far below, a different view, one can see an entire DQ with a bound peptide in a similar cleft, as viewed from the side.

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Some HLA-mediated diseases are directly involved in the promotion of cancer. Gluten-sensitive enteropathy is associated with increased prevalence of enteropathy-associated T-cell lymphoma, and DR3-DQ2 homozygotes are within the highest risk group, with close to 80% of gluten-sensitive

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HLA antibodies are typically not naturally occurring, and with few exceptions are formed as a result of an immunologic challenge to a foreign material containing non-self HLAs via blood transfusion, pregnancy (paternally inherited antigens), or organ or tissue transplant.

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or, shortened, B27). A parallel system that allowed more refined definition of alleles was developed. In this system, an "HLA" is used in conjunction with a letter, *, and a four-or-more-digit number (e.g., HLA-B*08:01, A*68:01, A*24:02:01N N=Null) to designate a specific

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Any cell displaying some other HLA type is "non-self" and is seen as an invader by the body's immune system, resulting in the rejection of the tissue bearing those cells. This is particularly important in the case of transplanted tissue, because it could lead to

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coefficient for these loci. In addition, some HLA loci are among the fastest-evolving coding regions in the human genome. One mechanism of diversification has been noted in the study of Amazonian tribes of South America that appear to have undergone intense

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a constriction with a great deal of variation. Of the 9 loci mentioned above, most retained a dozen or more allele-groups for each locus, far more preserved variation than the vast majority of human loci. This is consistent with a heterozygous or

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HLA typing has led to some improvement and acceleration in the diagnosis of celiac disease and type 1 diabetes; however, for DQ2 typing to be useful, it requires either high-resolution B1*typing (resolving *02:01 from *02:02), DQA1*typing, or DR

670:, it is the only effective means of discriminating between first-degree relatives that are at risk from those that are not at risk, prior to the appearance of sometimes-irreversible symptoms such as allergies and secondary autoimmune disease. 214:-positive or cytotoxic T-cells) that destroy cells. Some new work has proposed that antigens longer than 10 amino acids, 11-14 amino acids, can be presented on MHC I, eliciting a cytotoxic T-cell response. MHC class I proteins associate with 3190:

Gonzalez-Galarza FF, Mack SJ, Hollenbach J, Fernandez-Vina M, Setterholm M, Kempenich J, et al. (February 2013). "16(th) IHIW: extending the number of resources and bioinformatics analysis for the investigation of HLA rare alleles".

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recognize alleles and typing requires the sequencing of new alleles. Areas of the world where SSP-PCR or serotyping may be inadequate include Central Africa, Eastern Africa, parts of southern Africa, Arabia, S. Iran, Pakistan, and India.

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There are two parallel systems of nomenclature that are applied to HLA. The first, and oldest, system is based on serological (antibody based) recognition. In this system, antigens were eventually assigned letters and numbers (e.g.,

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Margaritte-Jeannin P, Babron MC, Bourgey M, Louka AS, Clot F, Percopo S, et al. (June 2004). "HLA-DQ relative risks for coeliac disease in European populations: a study of the European Genetics Cluster on Coeliac Disease".

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Cano P, Klitz W, Mack SJ, Maiers M, Marsh SG, Noreen H, et al. (May 2007). "Common and well-documented HLA alleles: report of the Ad-Hoc committee of the american society for histocompatiblity and immunogenetics".

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Senev A, Coemans M, Lerut E, Van Sandt V, Kerkhofs J, Daniëls L, Driessche MV, Compernolle V, Sprangers B, Van Loon E, Callemeyn J, Claas F, Tambur AR, Verbeke G, Kuypers D, Emonds MP, Naesens M (September 2020).

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frequencies of HLA-I and HLA-II genes for the European population has been compiled. In both cases the distribution of allele frequencies reveals a regional variation related with the history of the populations.

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Farjadian S, Naruse T, Kawata H, Ghaderi A, Bahram S, Inoko H (November 2004). "Molecular analysis of HLA allele frequencies and haplotypes in Baloch of Iran compared with related populations of Pakistan".

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There are three major and two minor MHC class II proteins encoded by the HLA. The genes of the class II combine to form heterodimeric (αβ) protein receptors that are typically expressed on the surface of

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Nunes JM, Buhler S, Roessli D, Sanchez-Mazas A (May 2014). "The HLA-net GENE pipeline for effective HLA data analysis and its application to 145 population samples from Europe and neighbouring areas".

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Valluri V, Valluei V, Mustafa M, Santhosh A, Middleton D, Alvares M, et al. (August 2005). "Frequencies of HLA-A, HLA-B, HLA-DR, and HLA-DQ phenotypes in the United Arab Emirates population".

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DR protein (DRA:DRB1*0101 gene products) with bound Staphylococcal enterotoxin ligand (subunit I-C), view is top down showing all DR amino acid residues within 5 Angstroms of the SEI peptide.

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Oshima M, Deitiker P, Ashizawa T, Atassi MZ (May 2002). "Vaccination with a MHC class II peptide attenuates cellular and humoral responses against tAChR and suppresses clinical EAMG".

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result in an increasing appearance of ambiguity. Because gene typing is based on SSP-PCR, it is possible that new variants, in particular in the class I and DRB1 loci, may be missed.

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between variable alleles and loci within each HLA gene class. Less frequently, longer-range productive recombinations through HLA genes have been noted producing chimeric genes.

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The other MHC class II proteins, DM and DO, are used in the internal processing of antigens, loading the antigenic peptides generated from pathogens onto the HLA molecules of

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Middleton D, Gonzalez F, Fernandez-Vina M, Tiercy JM, Marsh SG, Aubrey M, et al. (December 2009). "A bioinformatics approach to ascertaining the rarity of HLA alleles".

3647:"Eplet Mismatch Load and De Novo Occurrence of Donor-Specific Anti-HLA Antibodies, Rejection, and Graft Failure after Kidney Transplantation: An Observational Cohort Study" 1999:"The Role of Major Histocompatibility Complex in Organ Transplantation- Donor Specific Anti-Major Histocompatibility Complex Antibodies Analysis Goes to the Next Stage" 853: 168: 376:

Through a similar process, proteins (both native and foreign, such as the proteins of viruses) produced inside most cells are displayed on HLAs (to be specific,

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uses the HLAs to differentiate self cells and non-self cells. Any cell displaying that person's HLA type belongs to that person and is therefore not an invader.

269:. Predicting which (fragments of) antigens will be presented to the immune system by a certain HLA type is difficult, but the technology involved is improving. 1696:

For example, SSP-PCR within the clinical situation is often used for identifying HLA phenotypes. An example of an extended phenotype for a person might be:

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is extremely low. HLA genes have historically been identified as a result of the ability to successfully transplant organs between HLA-similar individuals.

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allele discovered. Marsh et al. (2005) can be considered a code book for HLA serotypes and genotypes, and a new book biannually with monthly updates in

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Matsumura M, Fremont DH, Peterson PA, Wilson IA (August 1992). "Emerging principles for the recognition of peptide antigens by MHC class I molecules".

2917:"A new HLA map of Europe: Regional genetic variation and its implication for peopling history, disease-association studies and tissue transplantation" 391:

When bound, peptides are presented to T-cells. T-cells require presentation via MHC molecules to recognize foreign antigens—a requirement known as

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serological reactions. Therefore, different serotype reactions may indicate the need to sequence a person's HLA to determine a new gene sequence.

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necessary to use broad antigen typing followed by gene sequencing because there is an increased risk of misidentifying by serotyping techniques.

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post-transplantation. There is a clear link between the risk of HLA antibody sensitisation and the donor-recipient HLA (molecular) mismatch.

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Shankarkumar U, Prasanavar D, Ghosh K, Mohanty D (May 2003). "HLA A*02 allele frequencies and B haplotype associations in Western Indians".

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Bodmer JG, Marsh SG, Albert ED, Bodmer WF, Dupont B, Erlich HA, et al. (May 1992). "Nomenclature for factors of the HLA system, 1991".

3115: 4129: 2128:"Empirical Evaluation of the Use of Computational HLA Binding as an Early Filter to the Mass Spectrometry-Based Epitope Discovery Workflow" 2177:

Galbraith W, Wagner MC, Chao J, Abaza M, Ernst LA, Nederlof MA, et al. (1991). "Imaging cytometry by multiparameter fluorescence".

753: 666:. Current serotyping can resolve, in one step, DQ8. HLA typing in autoimmunity is being increasingly used as a tool in diagnosis. In 3782: 3765: 3360: 1881: 4063: 3872: 123: 1146:

While the current CWD and rare or very rare designations were developed using different datasets and different versions of the

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Bouzid R, de Beijer MT, Luijten RJ, Bezstarosti K, Kessler AL, Bruno MJ, Peppelenbosch MP, Demmers JA, Buschow SI (May 2021).

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An HLA haplotype is a series of HLA "genes" (loci-alleles) by chromosome, one passed from the mother and one from the father.

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Illustration of an HLA-DQ molecule (magenta and blue) with a bound ligand (yellow) floating on the plasma membrane of the cell

4139: 4124: 2831:"Identification of Novel Candidate Epitopes on SARS-CoV-2 Proteins for South America: A Review of HLA Frequencies by Country" 2730: 1635:(or rarely, D locus). Every two years, a nomenclature is put forth to aid researchers in interpreting serotypes to alleles. 4134: 1122: 3514:

Apanius V, Penn D, Slev PR, Ruff LR, Potts WK (1997). "The nature of selection on the major histocompatibility complex".

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Antibodies against disease-associated HLA haplotypes have been proposed as a treatment for severe autoimmune diseases.

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and other diseases. People with certain HLA antigens are more likely to develop certain autoimmune diseases, such as

249:) present antigens from outside of the cell to T-lymphocytes. These particular antigens stimulate multiplication of 4093: 3856: 2364:"The Case for High Resolution Extended 6-Loci HLA Typing for Identifying Related Donors in the Indian Subcontinent" 4083: 3792: 2091:

Burrows SR, Rossjohn J, McCluskey J (January 2006). "Have we cut ourselves too short in mapping CTL epitopes?".

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The phenotype exampled above is one of the more common in Ireland and is the result of two common genetic

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encoded by HLAs are those on the outer part of body cells that are (in effect) unique to that person. The

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Number of variant alleles at class I loci according to the IMGT-HLA database, last updated October 2018:

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Parham P, Ohta T (April 1996). "Population biology of antigen presentation by MHC class I molecules".

2618:"Post-copulatory genetic matchmaking: HLA-dependent effects of cervical mucus on human sperm function" 3559: 3550:

Wedekind C, Seebeck T, Bettens F, Paepke AJ (June 1995). "MHC-dependent mate preferences in humans".

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may be used to give rise to a sibling with matching HLA, although there are ethical considerations.

3050:"Novel sequence feature variant type analysis of the HLA genetic association in systemic sclerosis" 873: 692: 651: 619: 523: 373:, which then produce a variety of effects and cell-to-cell interactions to eliminate the pathogen. 77: 2616:

Jokiniemi A, Magris M, Ritari J, Kuusipalo L, Lundgren T, Partanen J, Kekäläinen J (August 2020).

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antibodies specific to either are typically used more often than antibodies to broad antigens.

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receptor's peptide-binding cleft tend to produce molecules with different binding capability.

643: 320: 277: 266: 2452: 3707: 3666: 3658: 3610: 3567: 3523: 3488: 3453: 3417: 3382: 3313: 3305: 3251: 3200: 3147: 3069: 3061: 2982: 2974: 2928: 2889: 2852: 2842: 2794: 2686: 2678: 2637: 2629: 2588: 2580: 2539: 2531: 2492: 2375: 2334: 2326: 2274: 2263:"Serotyping for homotransplantation. XVII. Preliminary studies of HL-A subunits and alleles" 2233: 2225: 2186: 2149: 2139: 2100: 2065: 2020: 2010: 1943: 1935: 1749:

In general, this is identical to the extended serotype: A1,A3,B7,B8,DR3,DR15(2), DQ2,DQ6(1)

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Karp DR, Marthandan N, Marsh SG, Ahn C, Arnett FC, Deluca DS, et al. (February 2010).

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Marsh SG, Albert ED, Bodmer WF, Bontrop RE, Dupont B, Erlich HA, et al. (April 2010).

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HLAs have other roles. They are important in disease defense. They are the major cause of

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Jokiniemi A, Kuusipalo L, Ritari J, Koskela S, Partanen J, Kekäläinen J (November 2020).

1971: 1150:, the approximate fraction of alleles at each HLA locus in each category is shown below. 3563: 3292:

Mack SJ, Cano P, Hollenbach JA, He J, Hurley CK, Middleton D, et al. (April 2013).

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Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences

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Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences

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MHC class I proteins form a functional receptor on most nucleated cells of the body.

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Disease-related peptides fit into these "slots" much like a hand fits into a glove.

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4 β-chains (only 3 possible per person), encoded by HLA-DRB1, DRB3, DRB4, DRB5 loci

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with respect to certain genetic characteristics. This has led to a field known as

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in length. Foreign antigens presented by MHC class I attract T-lymphocytes called

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Agarwal RK, Kumari A, Sedai A, Parmar L, Dhanya R, Faulkner L (September 2017).

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to produce antibodies to that specific antigen. Self-antigens are suppressed by

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Kurkó J, Besenyei T, Laki J, Glant TT, Mikecz K, Szekanecz Z (October 2013).

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Mignot E, Lin L, Rogers W, Honda Y, Qiu X, Lin X, et al. (March 2001).

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Nakamura T, Shirouzu T, Nakata K, Yoshimura N, Ushigome H (September 2019).

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List of human leukocyte antigen alleles associated with cutaneous conditions

863: 142: 3721: 3680: 3662: 3622: 3571: 3500: 3465: 3429: 3327: 3263: 3212: 3159: 3083: 3018: 2996: 2942: 2901: 2866: 2700: 2651: 2633: 2602: 2553: 2504: 2389: 2348: 2330: 2247: 2229: 2190: 2163: 2112: 2034: 1957: 3579: 3535: 3394: 3294:"Common and well-documented HLA alleles: 2012 update to the CWD catalogue" 2806: 2288: 2198: 2077: 983: 903: 4073: 4040: 4035: 4025: 4008: 4003: 3998: 3993: 3988: 3976: 3971: 3696:"Preimplantation diagnosis for Fanconi anemia combined with HLA matching" 3065: 2015: 1829:(By serotyping A3-Cw7-B7-DR15-DQ6 or the older version "A3-B7-DR2-DQ1") 1644: 1536: 1502: 1468: 1434: 1404: 1374: 1344: 1310: 814: 807: 793: 380:) on the cell surface. Infected cells can be recognized and destroyed by 354: 338: 258: 65: 3694:

Verlinsky Y, Rechitsky S, Schoolcraft W, Strom C, Kuliev A (June 2001).

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Salmon J, Wallace DJ, Dubois EL, Kirou KA, Hahn B, Lehman TA (2007).

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Number of variant alleles at class II loci (DM, DO, DP, DQ, and DR):

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HistoCheck HLA matching tool for organ and stem cell transplantation

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dbMHC Home, NCBI's database of the Major Histocompatibility Complex

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Requena D, Médico A, Chacón RD, Ramírez M, Marín-Sánchez O (2020).

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binds with major and minor gene subunits to produce a heterodimer.

3914: 3909: 3904: 3899: 3894: 3889: 1276: 1242: 1208: 862: 751: 738: 734: 730: 723: 718: 713: 313: 191: 187: 183: 89: 3837:

Rare Alleles Project at the AlleleFrequencies Net Database (AFND)

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Hurley CK (1997). "DNA-based typing of HLA for transplantation."

3097: 2214:"Mammalian social odours: attraction and individual recognition" 1893: 1802:

which is called ' 'super B8' ' or ' 'ancestral haplotype' ' and

706:

There are three major and three minor MHC class I genes in HLA.

107: 3845: 2261:

Singal DP, Mickey MR, Mittal KK, Terasaki PI (November 1968).

254: 211: 90: 618:

HLA types are inherited, and some of them are connected with

3811: 2520:"Genetics of rheumatoid arthritis - a comprehensive review" 122:. The HLA system is also known as the human version of the 3807:

American Society for Histocompatibility and Immunogenetics

3787: 218:, which, unlike the HLA proteins, is encoded by a gene on 161:. The proteins encoded by certain genes are also known as 3802:

British Society for Histocompatibility and Immunogenetics

988: 908: 3801: 3806: 349:(APCs) engulf the pathogen through a process called 4051: 3923: 3880: 3116:"Immunology Database and Analysis Portal (ImmPort)" 2461:. Philadelphia: Lippincott Williams & Wilkins. 76: 64: 56: 51: 34: 3817:Allele Frequencies at Variable Immune related loci 2963:"Nomenclature for factors of the HLA system, 2010" 1109:DRB3, DRB4, DRB5 have variable presence in humans 357:from the pathogen are digested into small pieces ( 198:. In general, these particular peptides are small 3019:"Statistics < IMGT/HLA < IPD < EMBL-EBI" 2423:Mitchell RS, Kumar V, Abbas AK, Fausto N (2007). 361:) and loaded on to HLA antigens (to be specific, 3347:Leffell MS, Donnenberg AD, Rose NR, eds. (1997) 2313:Taylor CJ, Bolton EM, Bradley JA (August 2011). 1142:Table of HLA alleles in each prevalence category 2308: 2306: 2451:Values are given for Caucasians, according to 1917: 1915: 1913: 854:History and naming of human leukocyte antigens 3857: 1121:disease. Karp D. R. et al. describes a novel 8: 3756:Zsolt Harsanyi and Richard Hutton, Richard, 3133: 3131: 3129: 2915:Sanchez-Mazas A, Buhler S, Nunes JM (2013). 2524:Clinical Reviews in Allergy & Immunology 257:-positive T cells), which in turn stimulate 153:, which means that they have many different 3185: 3183: 3181: 3179: 3177: 3175: 3173: 3171: 3169: 2417: 2415: 2413: 2411: 2409: 2407: 2405: 2403: 2401: 2399: 2368:Biology of Blood and Marrow Transplantation 2003:International Journal of Molecular Sciences 3864: 3850: 3842: 1627:. HLA loci can be further classified into 768:Major MHC class II proteins only occur on 3824:at the U.S. National Library of Medicine 3758:Genetic Prophecy: Beyond the Double Helix 3711: 3670: 3339: 3337: 3317: 3073: 2986: 2932: 2856: 2846: 2690: 2641: 2592: 2543: 2447: 2445: 2379: 2338: 2278: 2237: 2212:Brennan PA, Kendrick KM (December 2006). 2153: 2143: 2024: 2014: 1947: 2956: 2954: 2952: 1152: 1129:Common, well-documented and rare alleles 410: 129:Mutations in HLA genes may be linked to 3287: 3285: 3283: 3281: 3279: 3277: 3275: 3273: 3236: 3234: 3232: 3230: 3193:International Journal of Immunogenetics 3104:from the original on 24 September 2006. 3012: 3010: 3008: 3006: 1909: 1878:hematopoietic stem cell transplantation 46:Schematic representation of MHC class I 3029:from the original on 20 September 2012 2754:Evolution and the Spiral of Technology 141:. The HLA gene complex resides on a 3 31: 7: 2723:"HLA Nomenclature @ hla.alleles.org" 1116:Sequence feature variant type (SFVT) 2573:American Journal of Human Genetics 867:Codominant expression of HLA genes 365:). They are then displayed by the 118:responsible for regulation of the 25: 3783:European Bioinformatics Institute 1882:preimplantation genetic diagnosis 789:α-chain encoded by HLA-DPA1 locus 687:There is evidence for non-random 157:, allowing them to fine-tune the 4064:Minor histocompatibility antigen 3873:Major histocompatibility complex 3552:Proceedings. Biological Sciences 3528:10.1615/critrevimmunol.v17.i2.40 3458:10.1111/j.1399-0039.2004.00302.x 3422:10.1111/j.1399-0039.2005.00441.x 3152:10.1111/j.1399-0039.2009.01361.x 2979:10.1111/j.1399-0039.2010.01466.x 2622:Proceedings. Biological Sciences 2497:10.1111/j.0001-2815.2004.00237.x 2280:10.1097/00007890-196811000-00005 827:α-chain encoded by HLA-DRA locus 642:(systemic lupus erythematosus), 124:major histocompatibility complex 40: 2733:from the original on 2 May 2018 1972:"Human leukocyte antigen (HLA)" 984: 904: 291:identical HLA molecules on all 3516:Critical Reviews in Immunology 1866:HLA matching for sick siblings 149:at 21.3. HLA genes are highly 1: 3493:10.1016/S0198-8859(03)00032-6 3349:Handbook of Human Immunology. 1123:sequence feature variant type 126:(MHC) found in many animals. 3387:10.1016/0198-8859(92)90079-3 3256:10.1016/j.humimm.2007.01.014 544:Systemic lupus erythematosus 498:Systemic lupus erythematosus 420:Diseases with increased risk 412:HLA and autoimmune diseases 4130:Genes on human chromosome 6 2799:10.1126/science.272.5258.67 2459:Dubois' lupus erythematosus 1876:In some diseases requiring 27:Genes on human chromosome 6 4156: 4094:Cluster of differentiation 3779:IMGT/HLA Sequence Database 2427:. Philadelphia: Saunders. 2381:10.1016/j.bbmt.2017.05.030 1869: 1642: 851: 341:enters the body, specific 285:organ transplant rejection 94:HLA region of Chromosome 6 4084:Human blood group systems 3760:, London: Granada, 1982 ( 3615:10.1080/08916930290022270 2683:10.1038/s41437-020-0350-8 2536:10.1007/s12016-012-8346-7 1922:Choo SY (February 2007). 1837:Role of allelic variation 1609:Serotype and allele names 1108: 913: 897:Tables of variant alleles 552: 506: 471:21-hydroxylase deficiency 429: 276:encode components of the 39: 3826:Medical Subject Headings 3822:Human+leukocyte+antigens 3713:10.1001/jama.285.24.3130 3351:pp. 521–55, Boca Raton: 3054:Human Molecular Genetics 2848:10.3389/fimmu.2020.02008 2105:10.1016/j.it.2005.11.001 1940:10.3349/ymj.2007.48.1.11 1022: 1005: 1002: 999: 996: 993: 916: 770:antigen-presenting cells 763:antigen-presenting cells 584:Diabetes mellitus type 1 569:Diabetes mellitus type 1 534:Diabetes mellitus type 1 367:antigen-presenting cells 347:antigen-presenting cells 18:Human Leukocyte Antigens 4089:Cell adhesion molecules 4059:Human leukocyte antigen 3797:The Anthony Nolan Trust 2835:Frontiers in Immunology 2756:. Trafford Publishing. 2425:Robbins Basic Pathology 2145:10.3390/cancers13102307 2070:10.1126/science.1323878 838:antigen-presenting cell 648:inclusion body myositis 167:, as a result of their 114:in humans which encode 106:) system or complex of 100:human leukocyte antigen 35:Human leukocyte antigen 3739:The Compatibility Gene 3663:10.1681/ASN.2020010019 3572:10.1098/rspb.1995.0087 2634:10.1098/rspb.2020.1682 2331:10.1098/rstb.2011.0030 2230:10.1098/rstb.2006.1931 2191:10.1002/cyto.990120702 1928:Yonsei Medical Journal 868: 757: 628:ankylosing spondylitis 456:Acute anterior uveitis 436:Ankylosing spondylitis 333:In infectious diseases 329: 272:HLAs corresponding to 225:HLAs corresponding to 178:HLAs corresponding to 159:adaptive immune system 95: 4140:Blood antigen systems 4125:Human MHC haplogroups 3793:HLA Informatics Group 1643:Further information: 866: 852:Further information: 755: 317: 210:(also referred to as 116:cell-surface proteins 93: 4135:Transfusion medicine 2093:Trends in Immunology 2016:10.3390/ijms20184544 632:rheumatoid arthritis 620:autoimmune disorders 559:Rheumatoid arthritis 513:Autoimmune hepatitis 406:transplant rejection 3564:1995RSPSB.260..245W 3098:"IMGT/HLA Database" 2791:1996Sci...272...67P 2752:Shennan DH (2006). 2455:(right column) in: 2062:1992Sci...257..927M 1604:Examining HLA types 1171:No. well-documented 985: 905: 874:balancing selection 813:β-chain encoded by 806:α-chain encoded by 792:β-chain encoded by 693:genetic matchmaking 413: 371:CD4+ helper T cells 147:chromosome 6, p-arm 131:autoimmune diseases 3066:10.1093/hmg/ddp521 3017:EBI Web Services. 2628:(1933): 20201682. 869: 758: 709:Major MHC class I 446:Reactive arthritis 411: 399:In graft rejection 330: 267:regulatory T cells 169:historic discovery 96: 4102: 4101: 4069:Blood transfusion 3751:978-0-241-95675-5 3737:Daniel M. Davis, 3310:10.1111/tan.12093 3205:10.1111/iji.12030 2934:10.1159/000360855 2894:10.1111/tan.12356 2763:978-1-55212-518-2 2468:978-0-7817-9394-0 2434:978-1-4160-2973-1 2325:(1575): 2312–22. 2224:(1476): 2061–78. 1797:A1-Cw7-B8-DR3-DQ2 1601: 1600: 1176:% well-documented 1148:IMGT/HLA Database 1113: 1112: 979: 978: 683:In mate selection 644:myasthenia gravis 611: 610: 278:complement system 173:organ transplants 88: 87: 16:(Redirected from 4147: 3866: 3859: 3852: 3843: 3726: 3725: 3715: 3691: 3685: 3684: 3674: 3651:J Am Soc Nephrol 3641: 3635: 3634: 3598: 3592: 3591: 3547: 3541: 3539: 3511: 3505: 3504: 3481:Human Immunology 3476: 3470: 3469: 3440: 3434: 3433: 3405: 3399: 3398: 3375:Human Immunology 3370: 3364: 3341: 3332: 3331: 3321: 3289: 3268: 3267: 3244:Human Immunology 3238: 3225: 3224: 3187: 3164: 3163: 3135: 3124: 3123: 3122:on 26 July 2011. 3118:. Archived from 3112: 3106: 3105: 3094: 3088: 3087: 3077: 3045: 3039: 3038: 3036: 3034: 3014: 3001: 3000: 2990: 2958: 2947: 2946: 2936: 2912: 2906: 2905: 2877: 2871: 2870: 2860: 2850: 2826: 2820: 2818: 2774: 2768: 2767: 2749: 2743: 2742: 2740: 2738: 2719: 2713: 2712: 2694: 2662: 2656: 2655: 2645: 2613: 2607: 2606: 2596: 2564: 2558: 2557: 2547: 2515: 2509: 2508: 2479: 2473: 2472: 2449: 2440: 2438: 2419: 2394: 2393: 2383: 2374:(9): 1592–1596. 2359: 2353: 2352: 2342: 2310: 2301: 2300: 2282: 2258: 2252: 2251: 2241: 2209: 2203: 2202: 2174: 2168: 2167: 2157: 2147: 2123: 2117: 2116: 2088: 2082: 2081: 2056:(5072): 927–34. 2045: 2039: 2038: 2028: 2018: 1994: 1988: 1987: 1985: 1983: 1968: 1962: 1961: 1951: 1919: 1828: 1824: 1820: 1816: 1812: 1808: 1794: 1790: 1786: 1782: 1778: 1774: 1746: 1742: 1738: 1734: 1730: 1726: 1722: 1718: 1714: 1710: 1706: 1702: 1153: 1006:Theor. possible 986: 906: 729:Minor genes are 652:Sjögren syndrome 524:Sjögren syndrome 486:Behçet's Disease 414: 327: 216:β2-microglobulin 202:, of about 8-10 44: 32: 21: 4155: 4154: 4150: 4149: 4148: 4146: 4145: 4144: 4105: 4104: 4103: 4098: 4047: 3919: 3876: 3870: 3788:hla.alleles.org 3775: 3734: 3729: 3693: 3692: 3688: 3657:(9): 2193–204. 3643: 3642: 3638: 3600: 3599: 3595: 3558:(1359): 245–9. 3549: 3548: 3544: 3513: 3512: 3508: 3478: 3477: 3473: 3446:Tissue Antigens 3442: 3441: 3437: 3410:Tissue Antigens 3407: 3406: 3402: 3372: 3371: 3367: 3342: 3335: 3298:Tissue Antigens 3291: 3290: 3271: 3240: 3239: 3228: 3189: 3188: 3167: 3140:Tissue Antigens 3137: 3136: 3127: 3114: 3113: 3109: 3096: 3095: 3091: 3047: 3046: 3042: 3032: 3030: 3016: 3015: 3004: 2967:Tissue Antigens 2960: 2959: 2950: 2927:(3–4): 162–77. 2914: 2913: 2909: 2882:Tissue Antigens 2879: 2878: 2874: 2828: 2827: 2823: 2785:(5258): 67–74. 2776: 2775: 2771: 2764: 2751: 2750: 2746: 2736: 2734: 2727:hla.alleles.org 2721: 2720: 2716: 2664: 2663: 2659: 2615: 2614: 2610: 2566: 2565: 2561: 2517: 2516: 2512: 2485:Tissue Antigens 2481: 2480: 2476: 2469: 2456: 2450: 2443: 2435: 2422: 2420: 2397: 2361: 2360: 2356: 2312: 2311: 2304: 2267:Transplantation 2260: 2259: 2255: 2211: 2210: 2206: 2176: 2175: 2171: 2125: 2124: 2120: 2090: 2089: 2085: 2047: 2046: 2042: 1996: 1995: 1991: 1981: 1979: 1978:. December 2020 1970: 1969: 1965: 1921: 1920: 1911: 1907: 1890: 1874: 1868: 1852: 1839: 1826: 1822: 1818: 1814: 1810: 1806: 1795:(By serotyping 1792: 1788: 1784: 1780: 1776: 1772: 1759: 1744: 1740: 1736: 1732: 1728: 1724: 1720: 1716: 1712: 1708: 1704: 1700: 1690: 1682:Tissue Antigens 1669: 1667:Gene sequencing 1660: 1658:Cellular typing 1647: 1641: 1623:at a given HLA 1611: 1606: 1202: 1197: 1192: 1187: 1182: 1177: 1172: 1167: 1162: 1157: 1144: 1131: 1118: 951:Minor Antigens 922:Major Antigens 899: 879:gene conversion 861: 856: 846: 746: 701: 685: 676: 624:type I diabetes 616: 614:In autoimmunity 603:Coeliac disease 579: 401: 393:MHC restriction 337:When a foreign 335: 319: 301: 145:stretch within 135:type I diabetes 47: 28: 23: 22: 15: 12: 11: 5: 4153: 4151: 4143: 4142: 4137: 4132: 4127: 4122: 4117: 4115:Gene complexes 4107: 4106: 4100: 4099: 4097: 4096: 4091: 4086: 4081: 4076: 4071: 4066: 4061: 4055: 4053: 4049: 4048: 4046: 4045: 4044: 4043: 4038: 4033: 4028: 4023: 4013: 4012: 4011: 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Retrieved 1975: 1966: 1934:(1): 11–23. 1931: 1927: 1875: 1860: 1857: 1853: 1844: 1840: 1831: 1825: ; DQB1 1821: ; DQA1 1817: ; DRB1 1804: 1801: 1791: ; DQB1 1787: ; DQA1 1783: ; DRB1 1770: 1765: 1763: 1760: 1751: 1748: 1698: 1695: 1691: 1681: 1678: 1674: 1670: 1661: 1652: 1648: 1633:MHC class II 1612: 1203:categorized 1145: 1136: 1132: 1119: 980: 909:MHC class I 900: 891: 887: 883: 870: 847: 844:Nomenclature 835: 767: 759: 728: 708: 705: 702: 686: 677: 660: 617: 402: 390: 386: 382:CD8+ T cells 375: 363:MHC class II 351:phagocytosis 336: 302: 289: 282: 271: 227:MHC class II 224: 177: 162: 128: 112:chromosome 6 103: 99: 97: 29: 4079:Calgranulin 3882:MHC class I 3381:(1): 4–18. 3199:(1): 60–5. 2099:(1): 11–6. 1688:Phenotyping 1629:MHC class I 1196:% very rare 1003:-B3 to -B5 859:Variability 689:mate choice 578:HLA-DR3 and 378:MHC class I 261:-producing 204:amino acids 196:proteasomes 180:MHC class I 151:polymorphic 52:Identifiers 4109:Categories 3741:, London, 1905:References 1850:Antibodies 1766:haplotypes 1757:Haplotypes 1639:Serotyping 1161:No. common 664:serotyping 656:narcolepsy 417:HLA allele 78:Membranome 3353:CRC Press 3221:205192491 2709:220947699 2179:Cytometry 1813: ; B 1809: ; C 1779: ; B 1775: ; C 1201:% alleles 674:In cancer 299:Functions 71:IPR037055 4074:Arrestin 3745:, 2014 ( 3722:11427142 3681:32764139 3623:12389643 3588:34971350 3501:12691707 3466:15496201 3430:16029430 3328:23510415 3264:17462507 3213:23198982 3160:19793314 3102:Archived 3084:19933168 3027:Archived 2997:20356336 2943:24861861 2902:24738646 2867:33013857 2841:: 2008. 2815:22209086 2731:Archived 2701:32747723 2671:Heredity 2652:32811307 2603:11179016 2554:23288628 2505:15140032 2390:28603069 2349:21727137 2297:32428268 2248:17118924 2164:34065814 2113:16297661 2035:31540289 1976:MS Trust 1958:17326240 1888:See also 1645:Serotype 1570:All Loci 1181:No. rare 1166:% common 815:HLA-DQB1 808:HLA-DQA1 794:HLA-DPB1 522:Primary 359:peptides 355:Proteins 339:pathogen 305:proteins 259:antibody 200:polymers 164:antigens 133:such as 66:InterPro 3875:classes 3672:7461684 3631:5690960 3580:7630893 3560:Bibcode 3536:9094452 3395:1399721 3319:3634360 3075:2807365 2988:2848993 2858:7494848 2807:8600539 2787:Bibcode 2779:Science 2692:7553942 2643:7482290 2594:1274481 2545:3655138 2453:Page 61 2340:3130422 2289:5696819 2239:1764843 2199:1782829 2155:8150281 2132:Cancers 2078:1323878 2058:Bibcode 2050:Science 2026:6769817 1949:2628004 1616:HLA-B27 1198:alleles 1193:alleles 1188:alleles 1183:alleles 1178:alleles 1173:alleles 1168:alleles 1163:alleles 1104:11,431 1087:34,528 1071:16,036 778:T cells 774:B cells 598:HLA-DQ8 594:HLA-DQ2 554:HLA-DR4 548:2 to 3 508:HLA-DR3 502:2 to 3 493:HLA-DR2 481:HLA-B51 466:HLA-B47 431:HLA-B27 345:called 328: 263:B-cells 155:alleles 4016:HLA-DR 3984:HLA-DQ 3967:HLA-DP 3950:HLA-DO 3933:HLA-DM 3828:(MeSH) 3764: 3749: 3720: 3679: 3669: 3629: 3621: 3586: 3578: 3534: 3499: 3464: 3428: 3393: 3359: 3326: 3316: 3262: 3219: 3211: 3158: 3082: 3072: 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