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Home , Galectin, Mannose receptor, SIGLEC

Tables of Representative Carbohydrate-Binding To date numerous CBPs have been identified and demonstrated or implicated in mediating various cellular events through -carbohydrate interactions. The Consortium will target representatives of three major families of CBPs 1) the C-type family (including the , Table 1), the Siglec family (including CD22, Table 2) and the family (Table 3). In addition, within the last five years receptors (TCR) have been documented to recognize carbohydrate when presented by non-classical (CD1) and classical major histocompatibility complex (MHC) antigen presenting molecules. For the purposes of this project, the sub-family of TCR and the corresponding presenting molecules represent a fourth family (Table 4) that will be investigated. In addition to these four families, several other CBPs have been identified which are not homologous members of these families, but that still fall within the scope of the program. Please see the Consortium CBP DB for a comprehensive listing of targets within the scope of the Consortium. Table 1. C-Type CBP Group Species Cell types Comments CD23a 2 Human B cells ¥ Binds CD11/CD18 via carbohydrate Murine ¥ Binds IgE via protein determinant subset ¥ Binds T cells to promote primary DC-SIGNb 2 Human ¥ binding motif ¥ Binds ICAM3 as ligand, also HIV gp120 DC-SIGNRc 2 Human Dendritic cells ¥ May be a co-receptor with DC-SIGN Dendritic cells in ¥ Dendritic cells in spleen, and skin Dectin-1d 2 Murine Spleen, thymus and (Langerhans) Skin (langerhan) Dectin-2e 2 Murine Same as Dectin-1 ¥ Similar expression pattern to dectin-1 HECLf 2 Human ¥ Putative cytoplasmic localization Erythroid precursors Langering 2 Human Langerhan cells ¥ Inducer of Birbeck granules (epidermal DC) Layilinh 2 Murine Numerous ¥ Located in membrane ruffles ¥ Binds talin MCLi 2 Murine Macrophage ¥ Macrophage restricted Minclej 2 Murine Macrophage ¥ Inducible C-type lectin (TNF, IL-6, INF-γ ) ¥ Binds Gal/GalNAc MMGLk 2 Murine ¥ Removes asialo- from serum ¥ Implicated in DTH Endothelial cells ¥ Binds SLeX containing carbohydrates l Murine E- 4 ¥ Mediates leukocyte trafficking to sites of Human Platelets ¥ Binds SLeX containing O-linked sugars on PSGL-1 P-selectinm 4 Murine Endothelial ¥ Mediates leukocyte trafficking to sites of Human inflammation ¥ Mediates platelet binding to Neutrophils ¥ Binds sulfated-SLex containing carbohydrates on O- Lymphocytes linked n Murine L-selectin 4 Human ¥ Mediates trafficking to sites of inflammation ¥ Mediates lymphocyte recirculation DEC-205o 6 Murine Dendritic ¥ Endocytic receptor involved in antigen processing Human Thymic endothelial ¥ Existing knock out mouse Macrophages ¥ Endocytic receptor Fibroblasts ¥ Binds GlcNAc Endo180p 6 Human Murine Endothelial Chondrocytes Macrophages ¥ Role in innate immunity, possible uptake of antigen Mannose (GalNAc- Hepatic endothelial for presentation by CD1 and MHC class II q 6 Murine SO4) receptor ¥ Mannose specific as monomer in macrophages ¥ GalNAc-SO4 specific as dimer in endothelial cells Phospholipase A(2) 6 Human Various ¥ Binds to phosholipase A2 through peptide receptorr Murine determinant

References: a 1-3; b 4-7; c 5; d 8; e 9; f 10; g 11; h 12; i 13; j 14; k 15-19; l 20; m 21-23, 20; n 24-26, 20; o 27-29; p 30; q 31-36; r 37-40 Table 2. Siglec Species Cell types Comments (# Ig Domais) ¥ Subsets of resident and inflammatory macrophages /Siglec-1a Human ¥ Does not require unmasking to mediate binding Macrophages (17) Murine ¥ Does not have -based motifs in cyt. Tail ¥ Prefers 2,3 linked over 2,6 linkage ¥ Regulator of activation ¥ Recruits SHP-1 to cytoplasmic ITIM motifs Human CD22/Siglec-2b (7) B cells ¥ Potential role in B cell homing to bone marrow Murine ¥ Binding site masked on resting B cells ¥ Strong preference for 2,6-linked sialic acids ¥ Commonly used as a marker of early myeloid cells Myeloid Human ¥ Co-cross-linking with Fc receptor inhibits activation CD33/Siglec-3c (2) progenitors Murine ¥ Two cytoplasmic ITIM-like motifs ¥ Recruits SHP-1 and SHP-2 to ITIM motifs ¥ Role in maintenance of , inhibitor of axonal growth Human MAG/Siglec-4d (4) ¥ Two forms with and without ITIM motif Murine Schwann cells ¥ Single cytoplasmic tyrosine-based motif ¥ Prefers 2,3 linked sialic acid over 2,6 linkage Monocytes ¥ Presence of two ITIM-like motifs Siglec-5e (4) Human Neutrophils ¥ Binds 2,3- 2,6- and 2,8-linked sialic acids ¥ Identified as low affinity Trophoblast Siglec-6f (3) Human ¥ Presence of two ITIM-like motifs B cells ¥ Restricted specificity for Sialyl Tn structure ¥ Identified as NK cell inhibitory receptor Natural killer cells ¥ Alternatively spliced form lacking D2 identified Siglec-7g (3) Human Monocytes ¥ Presence of two ITIM-like motifs ¥ Binds equally to 2,3 and 2,6-linked sialic acids ¥ Alt. Spliced forms have two ITIM-like motifs Siglec-8h (3) Human ¥ Prefers 2,3 linked sialic acid over 2,6 linkage Monocytes ¥ Presence of two ITIM-like motifs Siglec-9I (3) Human Neutrophils ¥ Binds equally to 2,3 and 2,6-linked sialic acids NK-like cells ¥ Presence of two ITIM-like motifs Siglec-10j (5) Human B cells ¥ Binds equally to 2,3 and 2,6-linked sialic acids Siglec-11(5) Human Under study ¥ Under investigation References a 41, 42; b 43-46; c 47, 48; d 47, 48, 9, 49, 50; e 51, 52; f 52; g 53-55; h 56; i 57, 58; j59 Table 3. Galectin / Species Structure Organs/Cell types Comments Galectin-1a • Probably most widely distributed galectin Abundant in most • Human • Binds terminal _-galactosides organs (muscle, heart, • Mouse • Occurs as a 14.9 kDa non-covalent dimer lung, liver, lymph • Rat • Expressed in epithelial endothelial, fibroblasts, and node, thymus, colon) smooth muscle cells b Galectin-2 • Thought to be expressed at a minor level • Human Stomach epithelial • Occurs as a 14.5 kDa non-covalent dimer • Rat cells

c Galectin-3 • N-terminus has a -like domain • Human Colon, macrophages, • Also known as the Mac-2 suface antigen • Mouse many other epithelial • Occurs as a 27.5 kDa monomer • Rat and fibroblastic cells

d Galectin-4 • Occurs as a divalent monomeric protein with • Human tandem CRDs that may have different sugar binding • Mouse Gastrointestinal specificity • Rat

e Galectin-5 • The single CRD has 85% homology to the CRD of • Rat Erythrocytes galectin-9

f Galectin-6 • 85% identical to galectin-6 • Human • Galectin-4 and galectin-6 genes are closely linked Gastrointestinal • Mouse

g Galectin-7 • Used as a marker of stratified • Human Skin • Mouse

h Galectin-8 • Occurs as a divalent monomeric protein with • Human Liver, kidney, cardiac tandem CRDs (joined by a link peptide) that may • Mouse muscle, lung, and have different sugar binding specificity • Rat brain • Its link peptide is totally unique from Galectin4/6

I Galectin-9 • Galectin-9 and its (most probably) allelic variant • Human ecalectin, are potent chemoattractants Thymus, kidney, • Mouse • Occurs as a divalent monomeric protein with Hodgkin’s lymphoma • Rat tandem CRDs (joined by a link peptide) that may have different sugar binding specificity j Galectin-10 • Most abundant protein in eosinophils, formerly (Charcot-Leyden called Charcot-Leyden Crystal Protein Crystal Protein) • Previously thought to have lysophospholipase Eosinophils, A. Human activity, but recently shown to be distinct • Mouse • Unlike galectin-1, galectin-10 binds mannose

k Galectin-11 (Grifin) • May represent a new lens crystallin, (galectin- related inter-fiber protein) Lens • Rat • Exhibits lack of affinity for simple β-galactosides • Like galectin-1, galectin-11 can dimerize References: a 60-62; b 63; c 64; d 65; e 66; f 67; g 68; h 69; i 70; j 71; k 72 Table 4. T cells Reactive with Antigens Containing Carbohydrates Antigen T cell Antigen presenting molec. Peptides Classical class I or • Glycopeptides αβ T cells Classical class II • Glycopeptides γδ T cells None—direct recognition • Synthetic glycosphingolipids Microbial Undefined microbial αβ T cells CD1a • Glucose monomycolate • αβ T cells CD1b • Phosphatidyl inositols • Hexosyl-1-phosphoisoprenoids αβ T cells CD1c • Mannosyl-phosphodolichols • Undefined microbial γδ T cells CD1c Autologous or synthetic glycolipids • Undefined autologous αβ T cells CD1a • Brain glycosphingolipids including αβ gangliosides and sulfatide T cells CD1b • Undefined autologous αβ T cells CD1c • α-galactosylceramide NK T cells CD1d • Undefined autologous References:

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