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. Author manuscript; available in PMC: 2009 Nov 3.
Published in final edited form as: Curr Protoc Immunol. 2008 May;CHAPTER:Appendix–1X. doi: 10.1002/0471142735.ima01xs81

Paired Receptor Systems of the Innate Immune System

Eriko Yamada 1, Daniel W McVicar 1
PMCID: PMC2771913  NIHMSID: NIHMS117187  PMID: 18491293

Introduction

Upon infection, the host immune system must integrate a constellation of extracellular signals to orchestrate the migration and activation of the multiple cell types required for the development an appropriate response. The host’s response must be strong enough to eliminate the pathogen, but at the same time regulated so as not to inflict unwarranted damage to bystander tissues. Cells of the innate immune system are the first to be activated following infection, and therefore, typically play both a vital early effector role and a key regulatory role in both the development and attenuation of the inflammatory response. In addition, innate immune cells are critical in shaping the subsequent adaptive immune response.

As an adaptation to this critical role, innate immune cells express many regulatory receptors. Unlike T cells and B cells, the response of innate immune cells is not generally regulated through antigen specific receptors. Instead, they detect infection through more generalized methods such as the recognition of pathogen-associated molecular patterns or screening for indicators of stress such as modification or absence of MHC class I molecules. Since innate immune responses are not antigen specific, the system requires tight regulatory controls that interpret signals from several receptor-ligand interactions. One mechanism for this regulation is the expression of Paired Receptor Systems.

Paired receptors tune the inflammatory response. Many of the inhibitory receptors within paired systems participate in self tolerance by recognizing ubiquitously expressed endogenous molecules such as MHC class I molecules, cell surface sialic acid modified proteins, or CD200. During infection these markers of self are either downregulated or modified. At the same time, stress related molecules are expressed. The activating siblings of Paired Receptor Systems may recognize these “alert” molecules up-regulated as a consequence of infection or may bind constitutively expressed ligands that gain advantage as inhibitory ligands are downregulated. Alternatively, activating siblings may directly recognize pathogen associated or encoded ligands. Perhaps resolution of this issue will follow the positive identification of more activating ligands. Regardless, it is clear that these paired systems play critical roles in overall regulation of innate immunity.

In this table we review paired receptors involved in the innate immune system. For purposes of this review we define “Paired Receptors” as families of related membrane proteins that show the following characteristics: 1) encoded by different genes, but located as a gene cluster on a given chromosomes, 2) have significant homology within their extracellular domains, 3) expressed on overlapping immune populations and, 4) confirmed to have both activating and inhibitory members. For simplicity in nomenclature, whenever possible we have used the official gene nomenclature provided by NCBI and listed all other names in the alias field. In addition, we have provided a listing of both mouse and human paired systems as some families exist in both species whereas others are important in the study of either human or murine models of disease.

Table 1.

Mouse

Family Receptor Cellular Expression Chromosome location Other names Receptor characteristics Cytoplasmic tyrosine motifs Ligands(s) Action References
CD200r Cd200r1 Granulocytes, Monocytes, DCs, Macrophages, Mst cells, T cells, NKT cells, B cells 16 CD200R, Mox2r, OX2R Type I transmembrane protein, 2 Ig-like domains None CD200 Inhibition Wright GJ. et al. 2000. Immunity. 13: 233
Wright GJ. et al. 2003. J Immunol. 171:3034
Vieites JM. et al. 2003. Gene. 311: 99
Hatherley D. et al. 2005. J Immunol.175: 2469
CD200r Cd200r3 Basophils, Mast cells, Monocytes, DCs, B cells 16 mCD200RLb Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None Unknown Activation Voehringer D. et al. 2004. J Biol Chem. 279: 54117
Wright GJ. et al. 2003. J Immunol. 171: 3034
Hatherley D. et al. 2005. J Immunol.175: 2469
CD200r Cd200r4 Macrophages, Monocytes, Granulocytes, Mast cells, DCs, NK, NKT, B cells 16 mCD200RLa, Cd200r2 Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None Unknown Activation Wright GJ. et al. 2003. J Immunol. 171: 3034
Hatherley D. et al. 2005. J Immunol.175: 2469
CD300 4732429D16Rik Granulocytes, Macrophages, Mast cells 11 CLM5, LMIR4, clm-5, RP23-331L12.3 Type I transmembrane protein, 1 Ig-like domain, Associates with FcR γ chain None Unknown Activation Izawa K. et al. 2007. J Biol Chem. Apr 16.
CD300 CD300a Macrophages, Mast cells, DCs, Granulocytes, B cells, Eosinophils 11 LMIR1, Clm8, MMAC8, Pigr4, mcpir1, MAIR-Ia Type I transmembrane, 1 Ig-like domain ITIMs Unknown Inhibition Munitz A. et al. 2006. J Allergy Clin Immunol. 118: 1082
Kumagai H. et al. 2003. Biochem Biophys Commun. 307: 719
Yotsumoto K. et al. 2003. J Exp Med. 198: 223
Bachelet I. et al. 2005. J Immunol. 175: 7989
CD300 Cd300b (AAY56360) Myeloid cells 11 mIREM-3 Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 and other unidentified molecules None Unknown Activation Martinez-Barriocanal A.and Sayos J. 2006. J Immunol. 177: 2819
CD300 Cd300c Monocytes, Neutrophils, T cells, B cells 11 Clm6 Type I transmembrane protein, Charged lysine residues in TM None Unknown Activation? Chung DH. et al. 2003. J Immunol. 171: 6541
CD300 CD300d Macrophages, a subset of B cells, Mast cells 11 LMIR2, CLM4, DIgR1, Igsf7, Cd300c, MAIR-II Type I transmembrane, Ig-like receptor, Associates with either DAP12, DAP10 or FcR γ chain None Unknown Activation Kumagai H. et al. 2003. Biochem Biophys Commun. 307: 719
Nakahashi C et al. 2007. J Immunol. 178: 765
Yotsumoto K. et al. 2003. J Exp Med. 198: 223
Turnbull IR. et al. 2007. Nature Rev. 7: 155
CD300 Cd300e Myeloid cells 11 CLM2; TREM5; Cd300le Type I transmembrane protein, Charged lysine residue in TM None Unknown Activation? Chung DH. et al. 2003. J Immunol. 171: 6541
CD300 Cd300lf DCs 11 CLM1, CLIM1, Digr2, IREM1, Pigr3, IgSF13, F730004D16Rik, LMIR3 Type I transmembrane protein, 1 Ig-like domain 2 ITIMs Unknown Inhibition Shi L. et al. 2006. Blood. 108: 2678
Álvarez-Errico D. et. al. 2007. J Immunol. 178: 808
Izawa K. et al. 2007. J Biol Chem. Apr 16.
Clecs Clec4a2 DCs, Neutrophils, B cells, Monocytes, Macrophages 6 DCIR, Dcir1, Clecsf6 Type II trasmembrane protein, C-type lectin ITIMs Unknown Inhibition Bates EE. et al. 1999. J Immunol. 163: 1973
Kanazawa N. et al. 2004. Immunobiology. 209: 179
Flornes LM. et al. 2004. Immunogenetics. 56: 506
Kanazawa N. et al. 2002. J Invest. Dermatol. 118: 261
Clecs Clec4b1 DCs, Monocytes, Macrophages, B cells 6 DCAR, Aplra2, Clec4b, mDcar2, DCARβ Type II transmembrane protein, C-type lectin, Associates with FcR γ chain None Unknown Activation Kanazawa N. et al. 2004. Immunobiology. 209: 179
Kanazawa N. et al. 2003. J Biol Chem. 278: 32645
Robinson MJ. et al. 2006. Nature Immunol. 7: 1258
Clecs Clec4n Monocytes, DCs, Polymorphonuclear cells 6 Clecsf10, dectin2 Type II transmemrane protein, C-type lectin, FcR γ chain? None weak affinity to mannose Activation Aragane Y.et al. 2003. J Immunol. 171: 3801
Taylor PR. et al. 2005. Eur J Immunol. 35: 2163
Robinson MJ. et al. 2006. Nature Immunol. 7: 1258
Fernandes MJ. et al. 1999. Cancer Res. 59: 2709
Clecs Clec5a Macrophages, Monocytes 6 Mdl1, Ly100, Clecsf5 Type II transmembrane protein, C-type lectin, Associates with DAP12 None Unknown Activation Bakker A.B. et al. 1999. Proc Natl Acad Sci USA. 96: 9692
Clecs Clec7a Macrophages, DCs, Monocytes, T cells, Polymorphonuclear cells 6 BGR, Clecsf12, β-GR Type II trasmembrane protein, C-type lectin ITAM-like motif (YxxL) β-1,3-linked and β-1,6-linked glucans Avtivation Ariizumi K. et al. 2000. J Bio Chem. 275: 20157
Klrb Klrb1a NK cells, NKT cells 6 Ly55a, NKR-P1A, Nkrp1-a, NKR-P1 2 C-type lectin, Type II transmembrane protein, Believed to associate with FcR γ chain YxxL motif Clr Activation or Inhibition Ryan JC. et al. 2001. Immunol Rev. 181:126
Giorda et al. 1990. Science. 249: 1298
Plougastel et al. 2001. Immunogenetics 53: 592
Klrb Klrb1b NK cells, NKT cells 6 Ly55b, NKR-P1B, Nkrp1-b, NKR-P1 34 C type lectin, Type II transmembrane protein ITIM Unknown Inhibition Carlyle JR. et al. 2004. Proc Natl Acad Sci USA. 101: 3527
Trowsdale et al. 2001. Immunol Rev 181: 20
Ryan JC. et al. 2001. Immunol Rev. 181: 126
Giorda et al. 1990. Science. 249: 1298
Plougastel et al. 2001. Immunogenetics 53: 592
Klrb Klrb1c NK cells, NKT cells 6 117187 C type lectin, Type II transmembrane protein, Associates with FcR γ chain none Unknown Activation Ryan JC. et al. 2001. Immunol Rev. 181: 126
Giorda et al. 1990. Science. 249: 1298
Plougastel et al. 2001. Immunogenetics 53: 592
Klrb Klrb1d NK cells, NKT cells 6 Ly55d, Nkrp1d, NKR-P1D C type lectin, Type II transmembrane protein ITIM Clr-b Inhibition Ryan JC. et al. 2001. Immunol Rev. 181: 126
Giorda et al. 1990. Science. 249: 1298
Plougastel et al. 2001. Immunogenetics 53: 592
Iizuka K. et al. 2003. Nat Immunol. 4: 801
Klrb Klrb1f NK cells, CD8+ T cells, DCs 6 Nkrp1f, NKR-P1F, Nkrp1-f C type lectin, Type II transmembrane protein, Charged arginine residue in TM None Clr-g Activation? Iizuka K. et al. 2003. Nat Immunol. 4: 801
Tian W. et al. 2005. Cell Immunol. 234: 39
Plougastel B. et al. 2001. Immunogenetics. 53: 209
Ly49 Ly49A NK cells, NKT cells, some T cells 6 - C-type lectins homodimers, Ly49A-like subfamily of Inhibitors ITIM Dd,k Inhibition Karlhofer et al. 1992. Nature 358: 66
Smith et al. 1994. J. Immunol. 153: 1068
Chan et al. 1989. J. Immunol. 142: 1727
Hanke et al. 1999. Immunity 11: 67
Ly49 Ly49B Myeloid cells 6 - C-type lectin; homodimers, Distantly related to inhibitory Ly49s ITIM Dd, Db, Dk, Kb, Kd Inhibition Wong et al. 1991. J. Immunol. 147: 1417
Scarpellino et al. 2007. J. Immunol. 178: 1277
Gays et al. 2006. J. Immunol. 177: 5840
Ly49 Ly49C NK cells, NKT cells, some T cells 6 - C-type lectins, Homodimers, Ly49C/I-like subfamily of inhibitors ITIM Db,d,k, Kd Inhibition Hanke et al. 1999. Immunity 11: 67
Brennan et al. 1996. J. Exp. Med. 184: 2085
Stoneman et al. 1995. J. Exp. Med. 182: 305
Ly49 Ly49D NK cells 6 - C-type lectins, Homodimers, Associates with DAP12, Ly49A-like subfamily of activators None Dd,r,sp Activation Ortaldo et al. 1999. J. Immunol. 163: 5269
Smith et al. 1994. J. Immunol. 153: 1068
Mason et al. 1996. J. Exp. Med. 184: 2119.
Ly49 Ly49E NK cells, NKT cells, some T cells 6 - C-type lectins, Homodimers, Ly49C/I-like subfamily of inhibitors ITIM ? Inhibition Makrigiannis et al. 2001. J. Immunol. 166: 5034
Smith et al. 1994. J. Immunol. 153: 1068
Gays et al. 2005. J. Immunol. 175: 2938
Fraser et al. 2002. Eur. J. Immunol. 32: 868
Ly49 Ly49F NK cells, NKT cells, some T cells 6 - C-type lectins, Homodimers, Ly49C/I-like subfamily of inhibitors ITIM ? Inhibition Smith et al. 1994. J. Immunol. 153: 1068
Gays et al. 2005. J. Immunol. 175: 2938
Ly49 Ly49G NK cells, NKT cells, some T cells 6 - C-type lectins; homodimers, Ly49G-like subfamily of inhibitors ITIM Dd, Ld Inhibition Hanke et al. 1999. Immunity 11: 67
Mason et al. 1995. J. Exp. Med. 182: 293
Smith et al. 1994. J. Immunol. 153: 1068
Ly49 Ly49H NK cells 6 - C-type lectin, Homodimer, Associates with DAP12, Ly49C/I-like subfamily of activators None Db, and m157 in B6 mice Activation Brennan et al. 1996. J. Exp. Med. 184: 2085
Gosselin et al. 1999. J. Leuk. Biol. 66: 165
Ly49 Ly49I NK cells, NKT cells, some T cells 6 - C-type lectins, Homodimers, Ly49C/I-like subfamily of inhibitors ITIM Kd, Dk, Dd (in B6 mice), m157 (in 129 mice) Inhibition Hanke et al. 1999. Immunity 11: 67
Makrigiannis et al. 2001. J. Immunol. 166: 5034
Ly49 Ly49J NK cells, NKT cells, some T cells 6 - C-type lectins, Homodimers, Ly49C/I-like subfamily of inhibitors ITIM ? Inhibition McQueen et al. 1999. Immunogenetics 48: 174
Ly49 Ly49L NK cells 6 - C-type lectin, Homodimer, Associates with DAP12, Ly49C/I-like subfamily of activators None Unknown Activation McQueen et al. 1999. Immunogenetics 48: 174
Makrigiannis et al. 2000. J. Leuk. Biol. 68: 765
Ly49 Ly49O NK cells, NKT cells, some T cells 6 - C-type lectins homodimers, Ly49A-like subfamily of Inhibitors ITIM Db,d,k,Ld Inhibition Makrigiannis et al. 1999. J. Immunol. 163: 4931
Ly49 Ly49P NK cells 6 - C-type lectins, Homodimers, Associates with DAP12, Ly49A-like subfamily of activators None Dd,k(in NOD mouse strain) Activation Makrigiannis et al. 1999. J. Immunol. 163: 4931
Silver et al. 2000. J. Immunol. 165: 1771
Ly49 Ly49Q pDC 6 - C-type lectin; homodimers, Distantly related to inhibitory Ly49s ITIM Kb Inhibition Makrigiannis et al. 2002. Genomics 79: 437
Tai et al. 2007. Mol. Immunol. 44: 2638
Toyama-Sorimachi et al. 2005. J. Immunol. 174: 4621
Scarpellino et al. 2007. J. Immunol. 178: 1277
Ly49 Ly49R NK cells 6 - C-type lectins, Homodimers, Associates with DAP12, Ly49A-like subfamily of activators None Db,d,k,Ld Activation Makrigiannis et al. 2001. J. Immunol. 166: 5034
Ly49 Ly49S NK cells, NKT cells, some T cells 6 - C-type lectins, Homodimers, Ly49C/I-like subfamily of inhibitors ITIM ? Inhibition Makrigiannis et al. 2001. J. Immunol. 166: 5034
Ly49 Ly49T NKT cells, some T cells 6 - C-type lectins, Homodimers, Ly49G-like subfamily of inhibitors ITIM ? Inhibition Makrigiannis et al. 2001. J. Immunol. 166: 5034
Ly49 Ly49U NK cells 6 - C-type lectin, Homodimer, Associates with DAP12, Ly49C/I-like subfamily of activators None Db Activation Makrigiannis et al. 2001. J. Immunol. 166: 5034
Ly49 Ly49V NK cells, NKT cells, some T cells 6 - C-type lectins homodimers, Ly49A-like subfamily of Inhibitors ITIM Dd, b,k,Kb,d,kLd Inhibition Makrigiannis et al. 2001. J. Immunol. 166: 5034
Ly49 Ly49W 6 - C-type lectin, Homodimer, Associates with DAP12, Ly49G-like subfamily of activators Dd, k (in NOD mouse strain) Unknown Activation Silver et al. 2001. J. Immunol. 166: 2333
NKG2 NKG2A/B NK cells, Activated CD8+, NKT cells 6 CD159A; Klrc1 C-type lectin, Heterodimerizes with CD94 2 ITIMs Qa-1(b) Inhibition Salcedo et al. 1998. Eur J Immunol. 28: 4356
Vance et al. 1998. J Exp Med. 188: 1841
Smith et al. 2001. Immunol Rev. 181: 115
Barten et al. 2001. Trends Immunol. 22: 52
Lohwasser et al. 1999. Eur J Immunol. 29: 755
NKG2 NKG2C NK cells, Activated CD8+ cells, NKT cells 6 Klrc2 C-type lectin, Heterodimerizes with CD94, Associates with DAP12 None Qa-1(b) Activation Salcedo et al. 1998. Eur J Immunol. 28: 4356
Vance et al. 1999. J Exp Med. 190: 1801
Smith et al. 2001. Immunol Rev. 181: 115
Barten et al. 2001. Trends Immunol. 22: 52
Lohwasser et al. 1999. Eur J Immunol. 29: 755
NKG2 NKG2D NK cells, Activated CD8+, LPS-activated macrophages 6 None C-type lectin, Associates with DAP10 None H60, RAE-1 Activation Ho et al. 1998. Proc Natl Acad Sci USA. 95: 6320
Smith et al. 2001. Immunol Rev. 181: 115
Cerwenka et al. 2000. Immunity. 12: 721
Lanier LL. 2003. Curr. Opin. Immunol. 15: 308
NKG2 NKG2D-S (short form) NK cells, T 6 None C-type lectin, Associates with DAP12 and DAP10 None H60, RAE-1 Activation Gilfillan S et al. 2002. Nature Immunol. 3: 1150
Diefenbach A et al. 2002. Nature Immunol. 3: 1142
Lanier LL. 2003. Curr Opin Immunol. 15: 308
NKG2 NKG2E Activated CD8+ cells 6 Klrc3 C-type lectin, Heterodimerizes with CD94, Associates with DAP12 None Qa-1(b) Activation Vance RE. et al. 1999. J Exp Med. 190: 1801
O'Callaghan CA. et al. 2000. Microbes Infect. 2: 371
PILR Pilra Granulocytes, Macrophages 5 FDF03 1 Ig-like domain ITIM CD99 like molecules Inhibition Fournier NJ. et al. 2000. Immunol. 165: 1197
Shiratori I. et al. 2004. J Exp Med. 199: 525
PILR Pilrb1 NK cells, Macrophages, DCs, Granulocytes 5 Fdact, Pilrb, FDFACT 1 Ig-like domain, Associates with DAP12 None CD99 like molecules Activation Fournier NJ. et al. 2000. Immunol. 165: 1197
Shiratori I. et al. 2004. J Exp Med. 199: 525
PIR Lilrb3 Mast cells, Macrophages, B cells, Granulocytes, DCs, Neurons 7 Gp91, Lilrb3, Pirb 6 Ig-like domains 1 ITIM and 2 ITSMs H-2Kb, 2Kk, 2Kd, 2Dd, 2Ld, HLA-G Inhibition Takai T. 2005. Immunology. 115: 433
Chen CC. et al. 2002. Immunol Res. 26: 191
Liang S. et al. 2002. Eur J Immunol. 32: 2418
Ujike A et al. 2002. Nat Immunol. 3: 542
Syken J. et al. 2006. Science. 313: 1795
PIR Pira1 Mast cells, Macrophages, DCs, Granulocytes 7 Pir, 6M21, Ly89, PIR-A1 6 Ig-like domains, Associates with FcR γ chain None H-2 molecules, HLA-G Activation Kubagawa H. et al. 1997. Proc Natl Acad Sci USA. 94: 5261
Takai T. 2005. Immunology. 115: 433
Colonna M. 2003. Nature Rev Immunol. 3: 1
Siglec Siglec-3 Monocytes, Macrophages 7 Cd33, gp67 Type I transmembrane protein, 2 Ig-like domains ITIM and ITSM Sialic acids Inhibition Brinkman-Van der Linden EC. et al. 2003. Mol Cell Biol. 23: 4199
Crocker PR. and Varki A. 2001. Immunology. 103: 137
Crocker PR. et al. 2002. Curr Op Structure Biol. 12: 609
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
Siglec Siglece Neutrophils, Monocytes, Macrophages, Eosinophils, NK cells, DCs, Some B cells 7 Cd170, Siglec5, Siglec9, Siglecl1, mSiglec-E Type I transmembrane protein, 3 Ig-like domains ITIM and ITSM Sialic acids Inhibition Aizawa H. et al. 2003. Genomics. 82: 521
Zhang JQ. 2004. Eur. J. Immunol. 34: 1175
Yu Z. et al. 2001. Biochem J. 353: 483
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
Siglec Siglecf Eosinophils 7 Siglec5; mSiglec-F Type I transmembrane protein, 4 Ig-like domains ITIM and ITSM Sialic acids (α2,3-linkages > α2,6-linkages) Inhibition Zhang M. et al. 2007. Blood. 109: 4280
Aizawa H. et al. 2003. Genomics. 82: 521
Angata T. et al. 2001. J Biol Chem. 276: 45128
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
Siglec Siglecg Eosinophils, B cells 7 Siglec-G; Siglec10; mSiglec-G Type I transmembrane protein, 5 Ig-like domains ITIM and ITSM Sialic acids Inhibition Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
Aizawa H. et al. 2003. Genomics. 82: 521
Siglec Siglech Plasmacytoid DCs 7 Siglec-H Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None Sialic acids Inhibition of IFNα pathway Blasius AL. et al. 2006. Blood. 107: 2474
Blasius AL. et al. 2006. Trends Immunol. 27: 255
Zhang J. et al. 2006. Blood. 107: 3600
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
SIRP Sirpα Myeloid cells, Neurons 2 Sirp-α, P84, SIRP, SHP-1, CD172a, Ptpns1, SHPS-1, Bit 3 Ig-like domains ITIMs CD47, Surfactant proteins A and D Inhibition Sano S. et al. 1997. FEBS Lett. 411: 327
Barclay AN. and Brown M.H. 2006. Nature Rev Immunol. 6: 457
SIRP Sirpβ1 Macrophages, Neutrophils 3 Sirpβ, SIRP-β 3 Ig-like domains, Associates with DAP12 None Unknown Activation Lahoud MH. et al. 2006. J Immunol. 177: 372
Hayashi A. et al. 2004. J Biol Chem. 279: 29450
Barclay AN. and Brown M.H. 2006. Nature Rev Immunol. 6: 457
TREM Trem1 A subset of monocytes and macrophages, Neutrophils, Granulocytes 17 - Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 None Unknown Activation Kelker MS. Et al. 2004. J Mol Biol. 344: 1175
Murakami Y. et al. 2007. J Immunol. 178: 1144
Chung DH. et al. 2002. Eur J Immunol. 32: 59
Bouchon A. et al. 2001. Nature. 410: 1103
TREM Trem2 Activated macrophages, DCs, Osteoclasts, Microglia 17 Trem2a, Trem2b, Trem2c Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 None Unknown Activation/Inhibition Daws MR. et al. 2003. J. Immunol. 171: 594
Turnbull IR et al. 2006. J Immunol. 177: 3520
Hamerman JA. et al. 2006. J Immunol. 177: 2051
Humphrey MB. et al. 2006. J Bone Miner Res. 21: 237
TREM Trem3 Myeloid cells 17 - Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 None Unknown Activation Chung DH. et al. 2002. Eur J Immunol. 32: 59

Table 2.

Human

Family Receptor Cellular Expression Chromosome location Other names Receptor characteristics Cytoplasmic tyrosine motifs Ligands(s) Action References
CD200R CD200R1 Macrophages, Neutrophils, Monocytes, DCs, Mast cells, T cells, NKT cells, B cells 3 OX2R, MOX2R, CD200R, HCRTR2 Type I transmembrane protein, 2 Ig-like domains None CD200 Inhibition Minas K. et al. 2006. Crit Rev Immunol. 26: 213
Wright GJ et al. 2003. J Immunol. 171: 3034
CD200R CD200R2 Basophils, Mast cells 3 CD200RLa Type I transmembrane protein, 2 Ig-like domain, Associates with DAP12 None Unknown Activation Wright GJ. et al. 2003. J Immunol. 171: 3034
Voehringer D. 2004. J Biol Chem. 279: 54117
CD300 CD300A Eosinophils, Mast cells 17 IRC1, IRC2, IRp60, IGSF12, CMRF35H, CMRF35H9 Type I transmembrane protein, 1 Ig-like domain, 3 ITIMs Unknown Inhibition Munitz A. et al. 2006. Blood. 107: 1996
Bachelet I. et al. 2005. J Immunol. 175: 7989
CD300 CD300C Monocytes, Neutrophils, T cells, B cells 17 CMRF35A1, CMRF35A, CMRF35, IGSF16 Type I trasnmembrane protein, 1 Ig-like domain, Adaptor molecule unknown None Unknown Activation Clark GL. et al. 2001. Tissue Antigens. 57: 415
CD300 CD300E Monocytes, Myeloid DCs, Macrophages 17 CLM2, IREM2, CD300LE Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 None Unknown Activation Aguilar H. et al. 2004. J Immunol. 173: 6703
Clark GJ. et al. 2007. J Immunother. 30: 303
CD300 CD300LB Myeloid cells 17 CLM7, IREM3, TREM5 Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 and unidentified molecules Tyrosine-based motif Unknown Activation Martinez-Barriocanal A.and Sayos J. 2006. J Immunol. 177: 2819
CD300 CD300LF Myeloid cells 17 CLM1, NKIR, IREM1, CD300f, IgSF13 Type I transmembrane protein, 1 Ig-like domain 2 ITIMs Unknown Inhibition Marquez J.A. et al. 2007. J Mol Biol. 367: 310
Alvarez-Errico D. et al. 2007. J Immunol. 178: 808
Alvarez-Errico D. et al. 2004. Eur J Immunol. 34: 3690
CLEC CLEC12A Monocytes, Macrophages, DCs, Granulocytes 12 CLL1, MICL, CLL-1, DCAL-2 Type II transmembrane protein, C-type lectin 1 ITIM Unknown Inhibition Marshall AS. et al. 2004. J Biol Chem. 279: 14792
Bakker AB. et al. 2004 Cancer Res. 64: 8443
Robinson MJ. et al. 2006. Nat Immunol. 7: 1258
CLEC CLEC2B Macrophages, Granulocytes, Monocytes 12 AICL, IFNRG1, CLECSF2 Type II trasmembrane protein, C-type lectin ? NKp80 Activation Welte S. et al. 2006. Nature Immunol. 7: 1334
CLEC CLEC2D NK cells 12 LLT1, OCIL, CLAX Type II transmembrane protein, C-type lectin ? KLRB1 (NKR-P1A) Activation Boles KS. et al. 1999. Immunogenetics. 50: 1
Lebbink RJ. and Meyaard L. 2007. Mol Immunol. 44: 2153
Aldemir H. et al. 2005. J Immunol. 176: 7791
Rosen DB. Et al. 2005. J immunol. 176: 7796
Mathew PA. et al. 2004. Mol Immunol. 40: 1157
CLEC CLEC4A DCs, Neutrophils, B cells, Monocytes, Macrophages 12 DCIR, LLI, DDB27, CLECSF6, HDCGC13P Type II trasmembrane protein, C-type lectin 1 ITIM Unknown Inhibition Kanazawa N. et al. 2004. Immunobiology. 209: 179
Bates EE. et al. 1999. J Immunol. 163: 1973
CLEC CLEC4C Plasmacytoid DCs, Monocytes, Macrophages, Neutrophils 12 DLEC, HECL, BDCA2, CD303, CLECSF7, CLECSF11 Type II trasmembrane protein, C-type lectin, Adaptor molecules unknown None Unknown Activation or Inhibition Dzionek A. et al. 2001. J Exp Med. 194: 1823
Dzionek A. et. al. 2002. Hum Immunol. 63: 1133
Kanazawa N. et al. 2004. Immunobiology. 209: 179
CLEC CLEC6A Monocytes, Macrophages, Neutrophils, DCs, B cells, Activated CD4+ T cells 12 CLECSF10, dectin-2 Type II trasmembrane protein, C-type lectin, Associates with Fc γ receptor None Hyphal components of some fungi Activation Sato et al. 2006. J Biol Chem. 281: 38854
Gavino AC. et al. 2005. Exp Dermatol. 14: 281
Robinson MJ. et al. 2006. Nat Immunol. 7: 1258
Kanazawa N. et al. 2004. Immunobiology. 209: 179
CLEC CLEC7A Macrophages, DCs, Monocytes, T cells, B cells, Eosinophils, Polymorphonuclear cells 12 DECTIN1, CLECSF12, BGR Type II trasmembrane protein, C-type lectin ITAM-like motif (YxxL) β-1,3-linked and β-1,6-linked glucans Activation Gersuk GM. et al. 2006. J Immunol. 176: 3717
Underhill DM. et al. 2005. Blood. 106: 2543
Robinson MJ. et al. 2006. Nat Immunol. 7: 1258
Lee MS. et al. 2007. Mol Cells. 23: 1
Willnent JA. et al. 2005. Eur J Immunol. 35: 1539
CLEC KLRB1 NK cells, NKT cells, T cells 12 NKR-P1A, NKR-P1, NKR, CD161, CLEC5B Type II trasmembrane protein, C-type lectin ? CLEC2D (LLT1) Activation or Inhibition Lanier LL. et al. 1994. J Immunol. 153: 2417
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CLEC KLRG1 Some NK cells and T cells 12 2F1, MAFA, MAFA-L, CLEC15A, MAFA-2F1, MGC13600 Type II transmembrane protein, C-type lectin 1 ITIM Cadherins Inhibition Lebbink RJ. and Meyaard L. 2007. Mol Immunol. 44: 2153
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KIR KIR2DL1 NK, NKT cells, T cells 19 NKAT, 47.11, CL-42, NKAT1, p58.1, CD158a, KIR-K3, KIR-K9, KIR221, KIR-K64, KIR-K65 Type I transmembrane protein, 2 Ig-like domains 2 ITIMs HLA-C N77/K80 Inhibition Stewart C.A. et al. 2005. Proc Natl Acad Sci USA. 102: 13224
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KIR KIR2DL2 NK cells, NKT cells, T cells 19 NKAT6, p58.2, CD158b, CD158B1 Type I transmembrane protein, 2 Ig-like domains 2 ITIMs HLA-C S77/N80 Inhibition Boyington JC. et al. 2001. Mol Immunol 38: 1007
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KIR KIR2DL3 NK, NKT cells, T cells 19 p58, CL-6, NKAT, GL183, NKAT2, CD158b, NKAT2A, NKAT2B, NKR-K7, CD158B2, KIR-K15, KIR-K7c, p58 KIR, KIR-023GB Type I transmembrane protein, 2 Ig-like domains 2 ITIMs HLA-C S77/N80 Inhibition Colonna M. and Samaridis J. Science. 1995. 268: 405
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KIR KIR2DL4 CD56high NK cells and IL-2 activated NK cells 19 G9P, CD158D, KIR103, KIR103AS Type I transmembrane protein, 3 Ig-like domains, Associates with FcR γ chain, Recruits SHP-2 ITIM HLA-G Activation/Inhibition Faure M. and Long EO. 2002. J Immuno. 168: 6208
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KIR KIR2DL5A NK cells, T cells 19 CD158F, KIR2DL5, KIR2DL5.1, KIR2DL5.3 Type I transmembrane protein, 3 Ig-like domains 2 ITIMs Unknown Inhibition Gomez-Lozano N. et al. 2002. Immunogenetics. 54: 314
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KIR KIR2DL5B NK cells, T cells 19 KIR2DL5, KIR2DL5.2, KIR2DLX Type I transmembrane protein, 3 Ig-like domains 2 ITIMs Unknown Inhibition Gomez-Lozano N. et al. 2002. Immunogenetics. 54: 314
Estefania E. et al. 2007. J Immunol. 178: 4402
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KIR KIR2DS1 NK cells 19 p50.1; CD158H; CD158a Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None HLA-C weakly, Peptide specificity is similar to KIR2DL1 Activation Lanier LL. et al. Annu Rev Immunol. 2005. 23: 225
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KIR KIR2DS2 NK cells 19 NKAT5, CD158J, CD158b, p58 KIR Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None Very low affinity to HLA-C S77/N80 with uncertainty Activation Snyder MR. et al. 2004. J Immunol 173: 3725
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KIR KIR2DS3 NK cells 19 NKAT7 Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None Unknown Activation Becker S. et al. 2003. Hum Immunol 64: 183
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KIR KIR2DS4 NK cells, NKT cells 19 KKA3, KIR1D, NKAT8, CD158I, KIR412 Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None Non-MHC class I protein Activation Lanier L.L. et al. Annu Rev Immunol. 2005. 23: 225
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KIR KIR2DS5 NK cells 19 NKAT9, CD158G Type I transmembrane protein, 2 Ig-like domains, Associates with DAP12 None Unknown Activation Becker S. et al. 2003. Hum Immunol. 64: 183
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KIR KIR3DL1 NK cells, NKT cells, T cells 19 KIR, CL-2, NKB1, AMB11, CL-11, NKAT3, NKB1B, KIR-G1, NKAT10, CD158E1, CD158E2, KIR3DS1, CD158E1/2 Type I transmembrane protein, 3 Ig-like domains 2 ITIMs HLA-A-Bw4, HLA-B-Bw4 Inhibition Lanier LL. et al. Annu Rev Immunol. 2005. 23: 225
Stewart CA. et al. 2003. J Immunol. 170: 6073
Yawata M. et al. 2006. J Exp Med. 203: 633
Carr WH. Et al. 2005. J Immunol. 175: 5222
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KIR KIR3DL2 NK cells, T cells 19 p140, NKAT4, CD158K, NKAT4B Type I transmembrane protein, 3 Ig-like domains 2 ITIMs HLA-A Inhibition Chan HW. et al. 2003. J Exp Med. 197: 245
Meenagh A. et al. 2004. Tissue Antigens. 64: 226
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KIR KIR3DL3 NK cells, T cells 19 KIR44, KIRC1, CD158Z, KIR3DL7, KIR3DL3var Type I transmembrane protein, 3 Ig-like domains 2 ITIMs Unknown Inhibition Jones DC. et al. 2006. Immunogenetics. 58: 614
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KIR KIR3DS1 NK cells, NKT cells, T cells 19 NKAT10, CD158E2 Type I transmembrane protein, 3 Ig-like domains, Associates with DAP12 None Unknown Activation Pascal V. et al. 2007. J Immunol. in press
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LILR LILRA1 Monocytes 19 LIR6, CD85I 4 Ig-like domains, Associates with FcR γ chain None Recognize classical and b2m free heavy chain form of HLA-B27 Activation Borges L. et al. 1997. J. Immunol. 159: 5192
Colonna M. 2003. Nature Rev Immunol. 3:1
Allen RL. et al. 2001. J Immunol. 167: 5543
Tedla N. et al. 2003. Proc Natl Acad Sci USA. 100: 1174
Brown D. et al. 2004. Tissue Antigens. 64: 215
LILR LILRA2 Eosinophils, Basophils, Monocytes, Neutrophils 19 LIR7, ILT1, CD85h 4 Ig-like domains, Associates with FcR γ chain None MHC class I? Activation Colonna M. 2003. Nature Rev Immunol. 3: 1
Tedla N. et al. 2003. Proc Natl Acad Sci USA. 100: 1174
Sloane DE. et al. 2004. Blood. 104: 2832
Brown D. et al. 2004. Tissue Antigens. 64: 215
LILR LILRA3 Soluble molecules 19 e3, HM31, HM43, ILT6, LIR4, CD85E, LIR-4 4 Ig-like domains None MHC class I? Activation Colonna M. 2003. Nature Rev Immunol. 3: 1
LILR LILRA4 Plasmacytoid DCs 19 ILT7, CD85g 4 Ig-like domains, Associates with FcR γ chain None Unknown Activation or Inhibition Brown D. et al. 2004. Tissue Antigens. 64: 215
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LILR LILRA5 Neutrophils, Monocytes, Myelomonocytic cells 19 CD85, LIR9, CD85F, ILT11, LILRB7 2 Ig-like domains, Associates with FcR γ chain, Extracellular domain can be a soluble molecule None Unknown Do not bind to classical and nonclassical MHC class I Activation Borges L. et al. 2003. Blood. 101: 1484
Brown D. et al. 2004. Tissue Antigens. 64: 215
Shiroishi M. et al. 2006. J Bio Chem. 281: 19536
LILR LILRB1 Monocytes, DCs, NK cells, T cells, B cells, Neutrophils, Eosinophils 19 LIR1, ILT2, CD85, CD85j 4 Ig-like domains 4 ITIMs MHC class I molecules (Preferentially HLA-G), HCMV UL18 Inhibition Tedla N. et al. 2003. Proc Natl Acad Sci USA. 100: 1174
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Shiroishi M. et al. 2006. Prod Natl Acd Sci USA. 103: 16412
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Cosman D. et al. 1997. J Immunol. 7: 273
LILR LILRB2 Myelomonocytic lineage, Eosinophils, Monocytes 19 CD85D, ILT4, LIR-2, LIR2, MIR-10, MIR10 4 Ig-like domains 3 ITIMs Recognize many MHC class I molecules, Preferentially HLA-G, free heavy chains of HLA-B27, HCMV UL18 Inhibition Tedla N. et al. 2003. Proc Natl Acad Sci USA. 100: 1174
Colonna M. 2003. Nature Rev Immunol. 3: 1
Shiroishi M. et al. 2006. Prod Natl Acd Sci USA. 103: 16412
Allen RL. et al. 2001. J Immunol. 167: 5543
Brown D. et al. 2004. Tissue Antigens. 64: 215
LILR LILRB3 Eosinophils, Basophils, Monocytes, Neutrophils 19 HL9, ILT5, LIR3, CD85A 4 Ig-like domains 4 ITIMs Unknown, Does not bind to B27 Inhibition Tedla N. et al. 2003. Proc Natl Acad Sci USA. 100: 1174
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Sloane DE. et al. 2004. Blood. 104: 2832
Brown D. et al. 2004. Tissue Antigens. 64: 215
LILR LILRB4 Monocytes 19 LILR5, LIR5, ILT3, CD85K 2 Ig-like domains 3 ITIMs Unknown, Does not bind to B27 Inhibition Colonna M. 2003. Nature Rev Immunol. 3:1
Tedla N. et al. 2003. Proc Natl Acad Sci USA. 100: 1174
Brown D. et al. 2004. Tissue Antigens. 64: 215
Vlad G. et al. 2006. Int Immunopharmacol. 6: 1889
LILR LILRB5 Monocytes 19 LILR8, CD85C 4 Ig-like domains 2 ITIMs Unknown Inhibition Colonna M. 2003. Nature Rev Immunol. 3: 1
Tedla N. et al. 2003. Proc Natl Acad Sci USA. 100: 1174
Brown D. et al. 2004. Tissue Antigens. 64: 215
Borges L. et al. 1997. J Immuno. 159: 5192
NKG2 KLRC1 NK, Activated CD8+, NKT cells 12 NKG2, NKG2A, CD159A Type II trasmemrbane protein, C type lectin, Heterodimerizes with CD94 2 ITIMs HLA-E Inhibition Lanier LL. et al. Annu Rev Immunol 2005. 23: 225
Kabat A. et al. 2002. J. Immunol. 169: 1948
Michaelsson J. et al. 2002. J Exp Med. 196: 1403
Brett KK. 2005. J Immunol. 174: 2878
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NKG2 KLRC2 NK, Activated CD8+ cells, NKT cells 12 NKG2C, CD159c Type II transmembrane protein, C type lectin, Heterodimerizes with CD94, associates with DAP12 None HLA-E Activation Lanier LL. et al. Annu Rev Immunol. 2005. 23: 225
Hikami K. et al. 2003. Genes Immun. 4: 160
Houchins JP. et al. 1991. J Exp Med. 173: 1017
Brett KK. 2005. J Immunol. 174: 2878
NKG2 KLRC3 NK, Activated CD8+ cells 12 NKG2E Type II transmembrane protein, C type lectin, Heterodimerizes with CD94, Associates with DAP12 None HLA-E Activation Lanier LL et al. 1998. Immunity. 8: 693
Brett KK. 2005. J Immunol. 174: 2878
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NKG2 KLRK1 NK, CD8+ T, γδT, LPS-activated macrophages 12 CD314, NKG2D, KLR Type II trasnemembrane protein, C type lectin, Homodimer, Associates with DAP10 None MICA/MICB, ULBP Activation Raulet DH. 2003. Nat Rev Immunol 3: 781
Lanier LL. 2003. Curr Opin Immunol. 15: 308
Groh V. et al. 2002. Nature. 419: 734
Sutherland CL. et al. 2002. J Immunol. 168: 671
PILR PILRA Monocytes, Macrophages, Granulocytes 7 PILRα, FDF03 1 Ig-like domain ITIMs CD99 related molecule Inhibition Mousseau DD. et al. 2000. J Biol Chem. 275: 4467
Koga T. et al. 2004. Nature. 428: 758
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PILR PILRB Leukocytes 7 PILRβ, FDFACT1, FDFACT2 1 Ig-like domain, Charged lysine residue in TM None Unknown Activation Mousseau DD. et al. 2000. J Biol Chem. 275: 4467
Fournier N. et al. 2000. J Immunol. 165: 1197
SIGLEC SIGLEC10 Eosinophils, Monocytes, NK cells, Neutrophils, B cells 19 SLG2, PRO940 Type I transmembrane protein, 5 Ig like-domains ITIM and ITSM Sialic acids (α2,3 linkages = α2,6-linkages) Inhibition Li N. et al. 2001. J Biol Chem. 276: 28106
Clark HF. et al. 2003. Genome Res. 13: 2265
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
SIGLEC SIGLEC11 Macrophages, Microglia 19 - Type I transmembrane protein, 5 Ig-like domains ITIM and ITSM Sialic acids (α2,8-linkages) Inhibition Hayakawa T. et al. 2005. Science. 309: 1693
Angata T et al. 2002. J Biol Chem. 277: 24466
Clark HF. et al. 2003. Genome Res. 13: 2265
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
SIGLEC SIGLEC14 Hematopoietic cells 19 - 3 Ig-like domains, Associates with DAP12 None Sialic acids Activation Angata T. et al. 2006. FASEB J. 20: 1964
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SIGLEC SIGLEC3 Myeloid progenitor cells, Monocytes, Activated T cells, Activated NK cells 19 CD33, p67, FLJ00391 Type I transmembrane protein, 2 Ig-like domains ITIM and ITSM Sialic acids (α2,6-linkages > α2,3-linkages) Inhibition Hernandez-Caselles T. et al. 2006. J Leukoc Biol 79: 46
Crocker PR. et al. 2002. Curr Op Structure Biol. 12: 609
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SIGLEC SIGLEC5 Neutrophils, Macrophages, Monocytes, Basophils, B cells 19 CD170, OBBP2, CD33L2, OB-BP2 Type I transmembrane protein, 4 Ig-like domains ITIM and ITSM Sialic acid (α2,3-linkages = α2,6-linkages > α2,8-linkages) Inhibition Angata T. et al. 2006. FASEB J. 20: 1964
Avril T. et al. 2005. J Biol Chem. 280: 19843
Erickson-Miller CL. et al. 2003. Exp Hematol. 31: 382
Ghannadan M. et al. 2002. Int Arch Allergy Immunol. 127: 299
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
SIGLEC SIGLEC6 Neutrophils, B cells, Placental trophoblasts 19 CD33L, OBBP1, CD33L1, CDw327 Type I transmembrane protein, 3 Ig-like domains ITIM and ITSM Sialyl Tn antigen Inhibition Patel N. et al. 1999. J Biol Chem. 274: 22729
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
SIGLEC SIGLEC7 Granulocytes, Monocytes, NK cells, CD8+ T cells 19 p75, QA79, AIRM1, CDw328, D-siglec, p75/AIRM1 Type I transmembrane protein, 3 Ig-like-domains ITIM and ITSM Sialic acids (α2,8-linkages > α2,6-linkages > α2,3-linkages) Inhibition Nicoll G. et al. 1999. J Biol Chem. 274: 34089
Angata T and Varki A. 2000. Glycobiology. 10: 431
Attrill H. et al. 2006. Biochem. J. 397: 271
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SIGLEC SIGLEC8 Eosinophils, Basophils, Mast cells 19 SAF2, MGC59785, SIGLEC8L Type I transmembrane protein, 3 Ig-like domains ITIM and ITSM Sialic acids (α2,3-linkages > α2,6-linkages) Inhibition Nutku E. et al. 2000. Blood. 101: 5014
Floyd H. et al. 2000. J. Biol. Chem. 275: 861
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
SIGLEC SIGLEC9 NK cells, T cells, B cells, Monocytes, Neutrophils 19 CDw329, OBBP-LIKE Type I transmembrane protein, 3 Ig-like domains ITIM and ITSM Sialic acids (α2,3 linkages = α2,6-linkages) Inhibition Angata T. and Varki A. 2000. J Biol Chem. 275: 22127
Ikehara Y. et al. 2004. J Biol Chem. 279: 43117
Rapoport EM. et al. 2006. Biochemistry Mosc. 71: 496
Crocker PR. et al. 2007. Nature Rev Immunol. 7: 255
SIRP SIRP-α Macrophages, Monocytes, DCs, Granulocytes, Neurons, Astrocytes 20 CD172a, SHPS1, P84, MYD-1, BIT, PTPNS1 3 Ig-like domains ITIMs CD47, Surfactant proteins A and D Inhibition Gardai SJ. et al. 2003. Cell. 115: 13
Seiffert M. et al. 1999. Blood. 94: 3633
Barclay A.N. and Brown M.H. 2006. Nature Rev Immunol. 6: 457
SIRP SIRP-β1 Macrophages, Neutrophils 20 CD172b, DKFZp686A05192, RP4-576H24.1 3 Ig-like domains, Forms disulphide-linked dimer, Associates with DAP12 None Unknown Activation Dietrich J. et al. 2000. J Immunol. 164: 9
Tomasello E. et al. 2000. Eur J Immunol. 30: 2147
Barclay AN. and Brown MH. 2006. Nature Rev Immunol. 6: 457
SIRP SIRP-β2 NK cells, T cells 20 CD172g, SIRPγ, SIRP-B2, bA77C3.1 3 Ig-like domains, No lysine to associate with adaptor molecules None CD47 No signals Piccio L. et al. 2005. Blood. 105: 2421
Barclay AN. and Brown MH. 2006. Nature Rev Immunol. 6: 457
TREM TREM-1 A subset of monocytes and macrophages, Neutrophils, Granulocytes 6 - Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 None Unknown Activation Bouchon A. et al. 2000. J Immunol. 164: 4991
Chung DH. et al. 2002. Eur J Immunol. 32: 59
Colonna M. et al. 2003. Nature Rev Immunol. 3: 1
TREM TREM-2 Macrophages, DCs, Osteoclasts, Microglia 6 Trem2a, Trem2b, Trem2c Type I transmembrane protein, 1 Ig-like domain, Associates with DAP12 None Unknown Activation or Inhibition Bouchon A. et al. 2000. J Immunol. 164: 4991
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