Affinage

MRAP

Melanocortin-2 receptor accessory protein · UniProt Q8TCY5

Length
172 aa
Mass
19.1 kDa
Annotated
2026-06-10
100 papers in source corpus 12 papers cited in narrative 12 extracted findings
Cross-family judge vs UniProt: Affinage preferred faithfulness: 6/6 claims corpus-supported (100%)

Mechanistic narrative

Synthesis pass · prose summary of the discoveries below

MRAP (melanocortin 2 receptor accessory protein) is a single-pass membrane protein that serves as an essential chaperone and signaling co-factor for the ACTH receptor MC2R, and loss-of-function mutations in MRAP cause familial glucocorticoid deficiency type 2 (PMID:15654338, PMID:16868047). It adopts an unusual antiparallel homodimeric topology in which both N- and C-termini face the extracellular side, and this dual-topology dimer forms a stable complex with MC2R that escorts the receptor out of the endoplasmic reticulum to the plasma membrane, where MC2R becomes glycosylated and competent to signal (PMID:18077336). MRAP performs two genetically separable functions—trafficking of MC2R and enabling ACTH binding/cAMP signaling at the cell surface—mapped to distinct N-terminal regions (PMID:19028547). Structurally, MRAP1 makes a sharp extracellular kink that acts as a 'seat-belt' stabilizing ACTH binding and MC2R activation within the MC2R–Gs–MRAP1 complex (PMID:36588120). Beyond MC2R, MRAP is a bidirectional regulator of the melanocortin receptor family, promoting MC2R surface expression while reducing responsiveness of MC1R, MC3R, MC4R, and MC5R, in part by trapping MC5R intracellularly and blocking its homodimerization (PMID:19329486, PMID:19535343). MRAP also binds Gαs to potentiate PKA-dependent, ACTH-induced lipolysis in adipocytes (PMID:29217655), and is required in vivo for adrenal progenitor differentiation and cortex zonation through WNT4/β-catenin and sonic hedgehog signaling (PMID:29879378).

Mechanistic history

Synthesis pass · year-by-year structured walk · 10 steps
  1. 2005 High

    Established MRAP's existence and core role by linking a human disease gene to MC2R biology, answering why some glucocorticoid-deficiency patients lack MC2R mutations.

    Evidence SNP mapping and mutation identification in FGD patients with co-immunoprecipitation showing MRAP–MC2R interaction and ER-to-surface trafficking

    PMID:15654338

    Open questions at the time
    • Molecular topology of MRAP unresolved
    • Mechanism distinguishing trafficking from signaling not defined
  2. 2006 Medium

    Confirmed that loss-of-function MRAP mutations are causative for FGD by identifying a homozygous truncating mutation in an affected family.

    Evidence Gene sequencing of patient DNA/fibroblasts identifying an exon 3 frameshift (L31X)

    PMID:16868047

    Open questions at the time
    • Single family
    • Mechanistic consequence inferred from prior functional work, not tested here
  3. 2007 High

    Resolved MRAP's membrane topology, revealing an unprecedented antiparallel homodimer that forms the functional unit complexing with MC2R.

    Evidence Epitope topology mapping, glycosylation mutagenesis, co-IP and surface immunoprecipitation in cells

    PMID:18077336

    Open questions at the time
    • Stoichiometry of MRAP:MC2R complex not defined
    • Atomic details of dimer interface unresolved
  4. 2008 High

    Separated MRAP's two activities by mutagenesis, showing trafficking of MC2R and enabling surface receptor binding/signaling are genetically distinct.

    Evidence Deletion/alanine-substitution mutagenesis with surface expression, ACTH binding, and cAMP assays

    PMID:19028547

    Open questions at the time
    • Structural basis of the signaling-enabling function not yet shown
    • How specific N-terminal residues contribute to ligand binding unclear
  5. 2009 High

    Broadened MRAP from an MC2R-specific chaperone to a bidirectional regulator of the entire melanocortin receptor family, including opposing effects on MC2R versus MC5R.

    Evidence Co-IP, surface expression, BiFC, and cAMP assays across MC1R–MC5R with MRAP and MRAP2

    PMID:19329486 PMID:19535343

    Open questions at the time
    • Mechanism by which MRAP traps MC5R intracellularly not fully defined
    • Physiological relevance of non-MC2R modulation untested in vivo
  6. 2012 Medium

    Placed MRAP dependence in an evolutionary context, showing MC2R MRAP-independence in an early-diverging vertebrate and a feed-forward transcriptional co-induction with MC2R in human adrenal tissue.

    Evidence Heterologous CHO expression of elephant shark MC2R; ACTH/AngII stimulation of primary adrenal cultures with mRNA quantification

    PMID:22419722 PMID:22919056

    Open questions at the time
    • Single-lab functional reconstitutions
    • Direct mechanism of co-regulation not established
  7. 2017 High

    Extended MRAP function beyond receptor trafficking by identifying Gαs as a binding partner mediating ACTH-driven lipolysis in adipocytes.

    Evidence Proteomics screen, co-IP, Gαs-binding mutant, 3T3-L1 knockdown/overexpression, and fat-specific transgenic mice

    PMID:29217655

    Open questions at the time
    • Structural basis of MRAP–Gαs interaction unknown
    • Whether Gαs binding contributes to MC2R signaling not tested
  8. 2018 High

    Defined MRAP's in vivo requirement for adrenal development, linking it to progenitor differentiation and cortex zonation pathways.

    Evidence Mrap-knockout mouse with histology, hormone measurement, and WNT4/β-catenin and sonic hedgehog pathway analysis

    PMID:29879378

    Open questions at the time
    • Whether developmental phenotype is MC2R-dependent or independent unresolved
    • Direct molecular link between MRAP and these pathways not established
  9. 2022 Medium

    Proposed MRAP1 as a broad-spectrum modulator of metabolic GPCRs beyond melanocortin receptors.

    Evidence Single-cell transcriptomics, co-IP across many GPCRs, surface translocation and cAMP assays, in vivo hypothalamic AAV knockdown

    PMID:36314066

    Open questions at the time
    • Most interactions not deeply validated
    • Single-lab screen; functional relevance of most GPCR pairs unestablished
  10. 2023 High

    Provided the structural mechanism of MRAP-assisted ACTH recognition through a cryo-EM 'seat-belt' model of the MC2R–Gs–MRAP1 complex.

    Evidence Cryo-EM structure determination with mutagenesis and signaling validation

    PMID:36588120

    Open questions at the time
    • Structure of MRAP with non-MC2R melanocortin receptors absent
    • Conformational dynamics of the antiparallel dimer during activation not resolved

Open questions

Synthesis pass · forward-looking unresolved questions
  • How MRAP's bidirectional and broad GPCR modulation, Gαs coupling, and developmental roles are mechanistically integrated remains open.
  • No unified model connecting receptor chaperone, Gαs scaffold, and developmental functions
  • Structural basis of inhibitory effects on MC1R/MC3R/MC4R/MC5R unknown

Mechanism profile

Synthesis pass · controlled-vocabulary classification · explore literature graph →
Molecular activity
GO:0098772 molecular function regulator activity 4 GO:0060090 molecular adaptor activity 2
Localization
GO:0005783 endoplasmic reticulum 2 GO:0005886 plasma membrane 2
Pathway
R-HSA-162582 Signal Transduction 3 R-HSA-9609507 Protein localization 2 R-HSA-1266738 Developmental Biology 1
Partners
GNASMC1RMC2RMC3RMC4RMC5R
Complex memberships
MC2R–Gs–MRAP1 complexMRAP antiparallel homodimer

Evidence

Reading pass · 12 per-paper findings extracted from the source corpus
Year Finding Method Journal Conf PMIDs
2005 MRAP (melanocortin 2 receptor accessory protein) interacts with the ACTH receptor (MC2R) and is required for trafficking of MC2R from the endoplasmic reticulum to the cell surface; loss-of-function mutations in MRAP cause familial glucocorticoid deficiency type 2. SNP array mapping, mutation identification, co-immunoprecipitation/interaction assay in cell-based system Nature genetics High 15654338
2007 MRAP forms antiparallel homodimers at the plasma membrane — a previously uncharacterized membrane protein topology. Both N- and C-terminal ends of MRAP face the extracellular side, MRAP is glycosylated on both ends, co-immunoprecipitation of differentially tagged MRAPs confirmed homodimerization, and the antiparallel homodimer forms a stable complex with MC2 receptor. Without MRAP, MC2R is retained in the ER; with MRAP, MC2R is glycosylated and reaches the plasma membrane where it signals in response to ACTH. Epitope topology mapping with antibodies, glycosylation mutagenesis, co-immunoprecipitation, selective surface immunoprecipitation, cell-surface signaling assay Proceedings of the National Academy of Sciences of the United States of America High 18077336
2008 MRAP facilitates MC2R trafficking to the plasma membrane and is absolutely required for ACTH binding and cAMP stimulation. A short sequence just N-terminal to the transmembrane domain is essential for dual (antiparallel) topology; the transmembrane region is not essential for topology but both regions are necessary for function. Deletion or alanine-substitution of other N-terminal regions yields MRAP mutants that support surface expression of MC2R but not receptor signaling, identifying two distinct actions of MRAP: trafficking of MC2R and enabling surface receptor binding/signaling. Deletion and alanine-substitution mutagenesis, cell-surface expression assay, cAMP signaling assay, ACTH binding assay Molecular and cellular endocrinology High 19028547
2009 MRAP and its homologue MRAP2 interact with all five melanocortin receptors (MC1R–MC5R). Interaction with MC2R results in surface expression and signaling. In contrast, MRAP and MRAP2 reduce responsiveness of MC1R, MC3R, MC4R, and MC5R to NDP-MSH, making MRAP and MRAP2 bidirectional regulators of the MCR family. Co-immunoprecipitation, cell-surface expression assay, cAMP signaling assay (ligand stimulation) Proceedings of the National Academy of Sciences of the United States of America High 19329486
2009 MRAP has opposite effects on MC2 and MC5 receptors: it promotes MC2R surface expression while trapping MC5R intracellularly. MRAP forms stable complexes with both receptors (co-precipitation), but MC2R–MRAP dimers are at the plasma membrane while MC5R–MRAP dimers are intracellular. MRAP additionally blocks MC5R homodimerization. The regions of MRAP required for these opposing effects on MC2R vs. MC5R differ. ELISA cell-surface assay, bimolecular fluorescence complementation (BiFC/YFP), co-immunoprecipitation, microscopy The Journal of biological chemistry High 19535343
2018 MRAP deficiency in mice causes impaired adrenal progenitor cell differentiation and disrupted cortex zonation. Mrap-null mice die at birth (rescued by corticosterone administration to dams) and surviving adults have isolated glucocorticoid deficiency with small adrenal glands, impaired capsular morphology, and dysregulation of WNT4/β-catenin and sonic hedgehog pathways in progenitor cells. Mrap-knockout mouse model, histology, pathway analysis (WNT4/β-catenin, sonic hedgehog), hormone measurement FASEB journal High 29879378
2017 MRAP interacts with Gαs as a novel binding partner (identified by unbiased proteomics screen and confirmed by co-immunoprecipitation). An MRAP mutant that cannot associate with Gαs fails to augment PKA activation and lipolytic response to ACTH. MRAP knockdown in 3T3-L1 adipocytes reduces ACTH-induced lipolysis; overexpression increases it. Transgenic mice overexpressing MRAP specifically in fat show increased lipolytic response to ACTH and resistance to high-fat diet-induced obesity. Proteomics screen, co-immunoprecipitation, MRAP knockdown/overexpression in 3T3-L1 adipocytes, Gαs-binding mutant, transgenic mouse model, lipolysis and metabolic assays Diabetes High 29217655
2023 Cryo-EM structure of the ACTH-bound MC2R–Gs–MRAP1 complex revealed that MRAP1 has a sharp kink at its extracellular region and exerts a 'seat-belt' effect that stabilizes ACTH binding and MC2R activation. Mutagenesis confirmed the structural basis of ACTH recognition by MC2R and the mechanism of receptor activation assisted by MRAP1. Cryo-electron microscopy (cryo-EM) structure determination, mutagenesis analysis, functional signaling assays Cell research High 36588120
2012 Expression of a cartilaginous fish (elephant shark) MC2R ortholog in CHO cells reaches the plasma membrane and signals in response to ACTH without requiring co-expression of exogenous MRAP, demonstrating that MRAP-independence is a feature of this early-diverging vertebrate MC2R and that the requirement for MRAP1 in MC2R trafficking and ACTH-selective signaling emerged after the divergence of cartilaginous and bony fishes. Heterologous expression in CHO cells, cell-surface trafficking assay, cAMP ligand-selectivity assay Endocrinology Medium 22919056
2022 MRAP1 (and MRAP2) were identified as broad-spectrum GPCR modulators interacting with 36 and 46 metabolic-related GPCRs, respectively, beyond the melanocortin receptor family. Co-expression of MRAP1/2 with these GPCRs altered surface translocation, constitutive activities, and ligand-stimulated downstream signaling. Hypothalamic MRAP2 knockdown (via AAV-shRNA) increased body weight in mice, and MRAP2 was required for CRF/CRHR1-mediated suppression of feeding. Single-cell transcriptome analysis, co-expression co-immunoprecipitation (protein-protein interaction), cell-surface translocation assays, cAMP signaling assays, in vivo AAV knockdown in mouse hypothalamus Clinical and translational medicine Medium 36314066
2012 In human adrenocortical tissues, MRAP and MC2R expression are co-induced by ACTH (~11-fold and ~20-fold, respectively) and by angiotensin II, consistent with a feed-forward mechanism facilitating MC2R cell-surface availability; MRAP2 expression was suppressed by forskolin and PMA. MRAP and MRAP2 levels were significantly lower in adrenocortical carcinomas than other adrenal tissues. Primary adrenal culture stimulation experiments, mRNA quantification, correlation analysis with hormone levels The Journal of clinical endocrinology and metabolism Medium 22419722
2006 A novel MRAP frameshift mutation (7-base deletion in exon 3, causing L31X stop codon) was identified in homozygous form in affected individuals from a family with familial glucocorticoid deficiency, confirming that loss-of-function MRAP mutations cause FGD. Gene sequencing of DAX-1, MC2R, and MRAP in patient-derived DNA/fibroblasts The Journal of clinical endocrinology and metabolism Medium 16868047

Source papers

Stage 0 corpus · 100 papers · ranked by NIH iCite citations
Year Title Journal Citations PMID
1993 Optimized survival of hippocampal neurons in B27-supplemented Neurobasal, a new serum-free medium combination. Journal of neuroscience research 2038 8377226
2005 Mutations in MRAP, encoding a new interacting partner of the ACTH receptor, cause familial glucocorticoid deficiency type 2. Nature genetics 325 15654338
2011 Th17 cells expressing KIR3DL2+ and responsive to HLA-B27 homodimers are increased in ankylosing spondylitis. Journal of immunology (Baltimore, Md. : 1950) 229 21248258
2015 HLA-B27. Annual review of immunology 218 25861975
2014 HLA-B27 and human β2-microglobulin affect the gut microbiota of transgenic rats. PloS one 204 25140823
2009 MRAP and MRAP2 are bidirectional regulators of the melanocortin receptor family. Proceedings of the National Academy of Sciences of the United States of America 197 19329486
1993 Susceptibility to inflammatory disease in HLA-B27 transgenic rat lines correlates with the level of B27 expression. Journal of immunology (Baltimore, Md. : 1950) 185 8473755
2002 Cell-surface expression and immune receptor recognition of HLA-B27 homodimers. Arthritis and rheumatism 183 12428240
1997 HLA-B27 polymorphism and worldwide susceptibility to ankylosing spondylitis. Tissue antigens 178 9062966
1999 Inflammatory disease in HLA-B27 transgenic rats. Immunological reviews 161 10450519
2013 HLA-B27 misfolding and ankylosing spondylitis. Molecular immunology 151 23993278
2007 Melanocortin-2 receptor accessory protein MRAP forms antiparallel homodimers. Proceedings of the National Academy of Sciences of the United States of America 141 18077336
2003 Lymphoblastoid cells express HLA-B27 homodimers both intracellularly and at the cell surface following endosomal recycling. European journal of immunology 137 12616495
2001 Leukocyte receptor complex-encoded immunomodulatory receptors show differing specificity for alternative HLA-B27 structures. Journal of immunology (Baltimore, Md. : 1950) 130 11698424
1989 HLA-B27 and HLA-A2 subtypes: structure, evolution and function. Immunology today 129 2679637
2010 From HLA-B27 to spondyloarthritis: a journey through the ER. Immunological reviews 126 20193000
2018 The microbiome and HLA-B27-associated acute anterior uveitis. Nature reviews. Rheumatology 104 30301938
2004 HLA-B27 heavy chain homodimers are expressed in HLA-B27 transgenic rodent models of spondyloarthritis and are ligands for paired Ig-like receptors. Journal of immunology (Baltimore, Md. : 1950) 102 15265899
1983 The cross-tolerance hypothesis, HLA-B27 and ankylosing spondylitis. British journal of rheumatology 102 6418259
2006 The pathogenetic role of HLA-B27 and its subtypes. Autoimmunity reviews 98 17289555
2001 HLA-B27 and genetic predisposing factors in spondyloarthropathies. Current opinion in rheumatology 97 11555726
1985 Organization, sequence and expression of the HLA-B27 gene: a molecular approach to analyze HLA and disease associations. Immunobiology 96 3912316
1984 HLA-B27 and the immune response to enterobacterial antigens in ankylosing spondylitis. Clinical and experimental immunology 93 6607143
2018 Effects of HLA-B27 on Gut Microbiota in Experimental Spondyloarthritis Implicate an Ecological Model of Dysbiosis. Arthritis & rheumatology (Hoboken, N.J.) 85 29287307
2008 Structure and function of the melanocortin2 receptor accessory protein (MRAP). Molecular and cellular endocrinology 85 19028547
2003 Formation of HLA-B27 homodimers and their relationship to assembly kinetics. The Journal of biological chemistry 82 14684742
2009 Opposite effects of the melanocortin-2 (MC2) receptor accessory protein MRAP on MC2 and MC5 receptor dimerization and trafficking. The Journal of biological chemistry 75 19535343
1992 HLA-B27 expression modulates gram-negative bacterial invasion into transfected L cells. Journal of immunology (Baltimore, Md. : 1950) 74 1588045
2000 Survival of hippocampal and cortical neurons in a mixture of MEM+ and B27-supplemented neurobasal medium. Free radical biology & medicine 73 10754261
2002 Psoriatic spondyloarthropathy: a comparative study between HLA-B27 positive and HLA-B27 negative disease. Seminars in arthritis and rheumatism 71 12077714
2013 Polymorphism of HLA-B27: 105 subtypes currently known. Current rheumatology reports 69 23990399
2021 Recent Developments in HLA B27 Anterior Uveitis. Frontiers in immunology 62 33469457
2015 HLA-B27 and psoriatic disease: a modern view of an old relationship. Rheumatology (Oxford, England) 60 26289052
1999 The role of HLA-B27 in spondyloarthritis. Immunogenetics 59 10602882
2002 HLA B27 in health and disease: a double-edged sword? Rheumatology (Oxford, England) 58 12154202
1979 HLA B27 and defects in the T-cell system in Whipple's disease. European journal of clinical investigation 57 93049
2011 An update on the genetics of HLA B27-associated acute anterior uveitis. Ocular immunology and inflammation 55 21428748
2022 Both Disease Activity and HLA-B27 Status Are Associated With Gut Microbiome Dysbiosis in Spondyloarthritis Patients. Arthritis & rheumatology (Hoboken, N.J.) 54 35818337
2006 HLA-B27 and the pathogenesis of spondyloarthropathies. Immunology letters 52 17129613
1993 Location of gp330/alpha 2-m receptor-associated protein (alpha 2-MRAP) and its binding sites in kidney: distribution of endogenous alpha 2-MRAP is modified by tissue processing. European journal of cell biology 52 8223699
1989 Molecular mimicry in HLA-B27-related arthritis. Annals of internal medicine 52 2672931
2017 The interplay between HLA-B27 and ERAP1/ERAP2 aminopeptidases: from anti-viral protection to spondyloarthritis. Clinical and experimental immunology 51 28759104
2011 What is the role of HLA-B27 in spondyloarthropathies? Autoimmunity reviews 49 21296192
2000 HLA-B27 alone rather than B27-related class I haplotypes contributes to ankylosing spondylitis susceptibility. Human immunology 49 10717805
2004 The immunobiology of HLA-B27: variations on a theme. Current molecular medicine 48 15011956
2010 The role of HLA-B27 in spondyloarthritis. The Journal of rheumatology 47 21123333
2012 Expression of HLA-B27 causes loss of migratory dendritic cells in a rat model of spondylarthritis. Arthritis and rheumatism 46 22674414
2008 Peptides: the cornerstone of HLA-B27 biology and pathogenetic role in spondyloarthritis. Tissue antigens 46 18489433
2010 HLA-B27: what's new? Rheumatology (Oxford, England) 43 20083539
1997 HLA-B27-derived peptides as autoantigens for T lymphocytes in ankylosing spondylitis. Arthritis and rheumatism 43 9365095
2018 Salmonella exploits HLA-B27 and host unfolded protein responses to promote intracellular replication. Annals of the rheumatic diseases 41 30355574
1978 HLA B27 and the genetics of ankylosing spondylitis. Annals of the rheumatic diseases 41 107868
2020 Emerging roles of melanocortin receptor accessory proteins (MRAP and MRAP2) in physiology and pathophysiology. Gene 39 32679290
2009 Subtypes of HLA-B27: history and implications in the pathogenesis of ankylosing spondylitis. Advances in experimental medicine and biology 38 19731628
2012 Identification of an MRAP-independent melanocortin-2 receptor: functional expression of the cartilaginous fish, Callorhinchus milii, melanocortin-2 receptor in CHO cells. Endocrinology 37 22919056
2002 HLA-B27: natural function and pathogenic role in spondyloarthritis. Arthritis research 37 12110134
1976 HLA-B27 and modified bone formation. Lancet (London, England) 37 55536
2022 T cell Repertoire Profiling and the Mechanism by which HLA-B27 Causes Ankylosing Spondylitis. Current rheumatology reports 36 36197645
2004 Animal models of HLA-B27-associated diseases. Current molecular medicine 36 15011957
2004 Recognition of classical and heavy chain forms of HLA-B27 by leukocyte receptors. Current molecular medicine 35 15011960
2013 HLA-B27 and antigen presentation: at the crossroads between immune defense and autoimmunity. Molecular immunology 34 23916069
2009 HLA-B27 misfolding and spondyloarthropathies. Prion 34 19363299
1984 Polymorphonuclear cell motility, ankylosing spondylitis, and HLA B27. Annals of the rheumatic diseases 34 6608924
2016 HLA-B27 Anterior Uveitis: Immunology and Immunopathology. Ocular immunology and inflammation 33 27245590
2009 Structural basis for T cell alloreactivity among three HLA-B14 and HLA-B27 antigens. The Journal of biological chemistry 33 19617632
1999 HLA-B27 associated spondyloarthropathy, an autoimmune disease based on crossreactivity between bacteria and HLA-B27? Annals of the rheumatic diseases 32 10491358
1996 Seronegative spondyloarthropathies and HLA-B27 subtyes: a study in Asian Indians. Clinical rheumatology 32 8835495
2021 Intraocular dendritic cells characterize HLA-B27-associated acute anterior uveitis. eLife 30 34783307
2018 The Role of Autophagy in the Degradation of Misfolded HLA-B27 Heavy Chains. Arthritis & rheumatology (Hoboken, N.J.) 29 29342507
2006 Unusual presentation of familial glucocorticoid deficiency with a novel MRAP mutation. The Journal of clinical endocrinology and metabolism 29 16868047
2009 The role of B27 heavy chain dimer immune receptor interactions in spondyloarthritis. Advances in experimental medicine and biology 28 19731637
2008 Induction of HLA-B27 heavy chain homodimer formation after activation in dendritic cells. Arthritis research & therapy 28 18759962
1998 MICA gene polymorphisms and HLA-B27 subtypes in Japanese patients with HLA-B27-associated acute anterior uveitis. Investigative ophthalmology & visual science 27 9501876
1994 Molecular analysis of HLA-B27 haplotypes in Caucasoids. Frequencies of B27-Cw in Jewish and Spanish populations. Human immunology 27 7860357
1990 B27+ disease versus B27- disease. Scandinavian journal of rheumatology. Supplement 27 2259879
2023 Structural basis of signaling regulation of the human melanocortin-2 receptor by MRAP1. Cell research 26 36588120
2018 Elephant shark melanocortin receptors: Novel interactions with MRAP1 and implication for the HPI axis. General and comparative endocrinology 26 30468718
2018 MRAP deficiency impairs adrenal progenitor cell differentiation and gland zonation. FASEB journal : official publication of the Federation of American Societies for Experimental Biology 25 29879378
2014 HLA-B27 subtype oligomerization and intracellular accumulation patterns correlate with predisposition to spondyloarthritis. Arthritis & rheumatology (Hoboken, N.J.) 25 24692163
2009 HLA-B27 misfolding and spondyloarthropathies. Advances in experimental medicine and biology 25 19731632
1996 HLA-B27 structure, function, and disease association. Current opinion in rheumatology 25 8864580
2023 HLA-B27, axial spondyloarthritis and survival. Annals of the rheumatic diseases 24 37679034
2016 Does the Microbiome Cause B27-related Acute Anterior Uveitis? Ocular immunology and inflammation 24 27002532
2009 Implications of structural and thermodynamic studies of HLA-B27 subtypes exhibiting differential association with ankylosing spondylitis. Advances in experimental medicine and biology 24 19731629
2019 HLA-B27 alters BMP/TGFβ signalling in Drosophila, revealing putative pathogenic mechanism for spondyloarthritis. Annals of the rheumatic diseases 23 31563893
2016 Non-conventional forms of HLA-B27 are expressed in spondyloarthritis joints and gut tissue. Journal of autoimmunity 23 27036372
2014 HLA-B27, but not HLA-B7, immunodominance to influenza is ERAP dependent. Journal of immunology (Baltimore, Md. : 1950) 23 24835397
2013 Expression of aberrant HLA-B27 molecules is dependent on B27 dosage and peptide supply. Annals of the rheumatic diseases 23 23625978
2012 Melanocortin 2 receptor-associated protein (MRAP) and MRAP2 in human adrenocortical tissues: regulation of expression and association with ACTH responsiveness. The Journal of clinical endocrinology and metabolism 22 22419722
2007 Profiling of microRNA expression by mRAP. Nature protocols 22 18079713
2013 The biochemistry and immunology of non-canonical forms of HLA-B27. Molecular immunology 21 23910730
1990 Expression of HLA-B27 in transgenic mice is dependent on the mouse H-2D genes. The Journal of experimental medicine 21 2212952
2022 Identification of MRAP protein family as broad-spectrum GPCR modulators. Clinical and translational medicine 20 36314066
2018 Influence of TNF and IL17 Gene Polymorphisms on the Spondyloarthritis Immunopathogenesis, Regardless of HLA-B27, in a Brazilian Population. Mediators of inflammation 20 29849482
2012 Functional expression of Squalus acanthias melanocortin-5 receptor in CHO cells: ligand selectivity and interaction with MRAP. European journal of pharmacology 20 22314217
2016 ERAP1 reduces accumulation of aberrant and disulfide-linked forms of HLA-B27 on the cell surface. Molecular immunology 19 27107845
2000 Colitis in HLA-B27/beta 2 microglobulin transgenic rats. International reviews of immunology 19 10723676
2000 HLA-B27 and immunogenetics of spondyloarthropathies. Current opinion in rheumatology 19 10910175
1988 Molecular biology of the HLA-B27 locus. British journal of rheumatology 19 3042071
2017 Regulation of Lipolytic Response and Energy Balance by Melanocortin 2 Receptor Accessory Protein (MRAP) in Adipocytes. Diabetes 18 29217655

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