Table 1.
C-type superfamily, their carbohydrate ligands and protein expression in human organs.
|
Common Name (HUGO Name if Different) |
Gene Symbol | Carbohydrate Preferential Affinity | Protein Expression in the Organs |
|---|---|---|---|
| Proteoglycans or lecticans | |||
| Aggrecan | ACAN | Hyaluronic acid [25] | Cartilage, soft tissue |
| Brevican | BCAN | Hyaluronic acid [26,27] | Brain |
| Neurocan | NCAN | Hyaluronic acid [28] | Brain |
| Versican | VCAN | Hyaluronic acid [29] | Brain |
| FRAS1 related extracellular matrix 1 | FREM1 | b) | Adrenal gland, appendix, colon, duodenum, epididymis, kidney, lung, pancreas, placenta, rectum, salivary gland, small intestine, stomach, testis, tonsil, thyroid gland |
| Type II transmembrane receptors | |||
| Blood Dendritic Cell Antigen 2 (C-type lectin domain family 4 member C) | CLEC4C | Gal-β-(1-3 or 1-4)-GlcNAc-β-(1-2)-Man trisaccharides [30,31] | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, muscle tissues, pancreas, proximal digestive tract, skin |
| DC-SIGN (CD209 molecule) | CD209 | High N-linked d-Mannose- oligosaccharides, and branched l-fucose, both with free OH-3 and OH-4. (N-linked glycans, N-acetyl-d-glucosamine, Lewis a, b, x and y) [32] | Bone marrow, lung |
| DC-SIGN2 | CLEC4M | High N-linked d-Mannose- oligosaccharides, branched l-fucose, N-linked glycans, N-acetyl-d-glucosamine, Lewis a, b and y | Brain, gastrointestinal tract, lung |
| Dectin-2 (C-type lectin domain containing 6A) | CLEC6A | α-(1-2) or α-(1-4) mannans [33] and other high-α-d-mannose carbohydrates [34] | Blood |
| Dendritic cell immunoreceptor (DCIR) (C-type lectin domain family 4 member A) | CLEC4A | Mannose, fucose and weakly interacts with N-acetylglucosamine [35] | Bone marrow, spleen, lung |
| Fc fragment of IgE receptor II | FCER2 | Mannose [36], immunoglobulin E, CD21, galactose [37] | Lymph node, bone marrow, spleen, appendix, tonsil, skin |
| Hepatic Asialoglycoprotein Receptor 1 | ASGR1 | Terminal β-d-galactose and N-acetylgalactosamine units [38] | Stomach, liver, gallbladder |
| Hepatic Asialoglycoprotein Receptor 2 | ASGR2 | Terminal β-d-galactose and N-acetylgalactosamine units [38] | Liver |
| Kupffer Cell receptor (C-type lectin domain family 4 member F) | CLEC4F | Galactose, fucose, and N-acetylgalactosamine [39] | Liver |
| Langerin (CD207 molecule) | CD207 | High-mannose oligosaccharides, mannose, N-acetylglucosamine, fucose. Note that OH-3 and OH-4 should be free for recognition, and preferentially equatorial. N-acetylmannosamine showed less affinity; thereby axial derivatives should be avoided. Sulfated mannosylated glycans, keratan sulfate and β-glucans [40] | Lymph node, tonsil, skin, spleen |
| Liver sinusoidal epithelial cell lectin (LSECtin) (C-type lectin domain family 4 member G) | CLEC4G | Mannose, N-acetylglucosamine and fucose [41] | Lymph node, brain, colon, kidney, liver, testis |
| Macrophage Asialoglycoprotein Receptor | CLEC10A | Terminal galactose and N-acetylgalactosamine residues [42] | Bone marrow, brain, lymph node, oral mucosa, skin, spleen, tonsil |
| Macrophage C-type Lectin (MCL) | CLEC4D | Trehalose 6,6′-dimycolate, α-d-mannans18 (however it was suggested that MCL is not a carbohydrate-binding lectin) [43] | Bone marrow, lung, lymph node, spleen, tonsil |
| MINCLE (C-type lectin domain family 4 member E) | CLEC4E | α-mannose, trehalose-6′6-dimycolate, glucose [19] | a) |
| Collectins | |||
| Collectin-K1 (collectin subfamily member 11) | COLEC11 | High mannose oligosaccharides with at least a mannose-α-(1-2)-mannose residue [44] | a) |
| Collectin-L1 (collectin subfamily member 10) | COLEC10 | Galactose, mannose, fucose, N-acetylglucosamine, N-acetylgalactosamine [45] | a) |
| Mannose-binding lectin 2 | MBL2 | Mannose, fucose, N-acetylglucosamine [46] | Liver |
| Pulmonary surfactant protein 1 (surfactant protein A1) | SFTPA1 | N-acetylmannosamine, l-fucose, mannose, glucose, poorly to galactose. Preferentially oligosaccharides [47] | Lung |
| Pulmonary surfactant protein 2 (surfactant protein A2) | SFTPA2 | N-acetylmannosamine, l-fucose, mannose, glucose, poorly to galactose. Preferentially oligosaccharides [47] | Lung |
| Pulmonary surfactant protein B (surfactant protein B) | SFTPB | b) | Lung |
| Pulmonary surfactant protein C (surfactant protein C) | SFTPC | Lipopolysaccharides [47] | Lung |
| Pulmonary surfactant protein D (surfactant protein D) | SFTPD | Maltose, glucose, mannose, poorly to galactose. Preferentially oligosaccharides [47] | Lung |
| Scavenger receptor with CTLD (SRCL) (collectin subfamily member 12) | COLEC12 | d-galactose, l- and d-fucose, N-acetylgalactosamine (internalizes specifically in nurse-like cells), sialyl Lewis X, or a trisaccharide and asialo-orosomucoid (ASOR). May also play a role in the clearance of amyloid-beta in Alzheimer disease [48] | Brain, lung, placenta |
| Selectins | |||
| Selectin E | SELE | Sialyl Lewis x, a [49] | Bone marrow, colon, nasopharynx |
| Selectin L | SELL | Sialyl Lewis x [50] | Appendix, bone marrow, lymph node, spleen, tonsil |
| Selectin P | SELP | Sialyl Lewis x [49] | Bone marrow, colon |
| Natural Killer (NK) | |||
| C-type lectin domain family 2 member L | CLEC2L | b) | Brain, skeletal muscle |
| C-type lectin domain containing 5A | CLEC5A | Fucose, mannose, N-acetylglucosamine, N-acetylmuramic acid-β(1-4)-N-acetylglucosamine [51] | Blood |
| CD72 molecule | CD72 | b) | Appendix, bone marrow, lymph node, spleen, tonsil |
| Killer cell lectin-like receptor G1 | KLRG1 | Mannose [52] | Appendix, cervix (uterine), colon, duodenum, small intestine, stomach, tonsil |
| Killer cell lectin-like receptor G2 | KLRG2 | b) | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, muscle tissues, pancreas, proximal digestive tract, skin |
| CD69 molecule | CD69 | Fucoidan (weak). N-acetylamine was reported but not supported by a second report. Does not bind glucose, galactose, mannose, fucose or N-acetylglucosamine [53] | Appendix, bone marrow, lymph node, spleen, tonsil |
| Killer cell lectin-like receptor F1 | KLRF1 | Predicted to not bind carbohydrates [54] | Blood |
| C-type lectin domain family 2 member B | CLEC2B |
b) Known to bind to KLRF1 |
Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, muscle tissues, proximal digestive tract, skin |
| Oxidized low-density lipoprotein receptor 1 | OLR1 | Predicted to not bind to carbohydrates [55] | a) |
| Killer cell lectin-like receptor D1 | KLRD1 | α-(2-3)-linked NeuAc on multi-antennary N-glycan, heparin, sulfate-containing polysaccharides [56] | a) |
| C-type lectin domain family 1 member A | CLEC1A | b) [57] | a) |
| C-type lectin domain family 1 member B | CLEC1B | Predicted to not bind to carbohydrates [58] | a) |
| C-type lectin domain family 12 member B | CLEC12B | b) | a) |
| C-type lectin-like 1 | CLECL1 | Predicted to not bind to carbohydrates [21] | a) |
| C-type lectin domain family 12 member A | CLEC12A | b) | Bone marrow, lung, spleen |
| DNGR (C-type lectin domain containing 9A) | CLEC9A | Specific interactions were not discovered yet, although it is known that this lectin binds to α-actin filaments and β-spectrin [59] | a) |
| C-type lectin domain family 2 member A | CLEC2A | b) | Skin |
| Dectin-1 (C-type lectin domain containing 7A) | CLEC7A | β-(1-3)- and β-(1-6)-d-Glycans (neither mono- or short oligosaccharides/polymers are recognized) [60] | Blood, bone marrow |
| C-type lectin domain family 2 member D | CLEC2D | High molecular weight sulfated glycosaminoglycans [61] | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, muscle tissues, pancreas, proximal digestive tract, skin |
| Killer cell lectin-like receptor B1 | KLRB1 | Terminal Gal-α-(1-3)-Gal, N-acetyllactosamine. [62] Sucrose octasulphate [63] | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, muscle tissues, pancreas, proximal digestive tract, skin |
| Killer cell lectin-like receptor C1 | KLRC1 | b) | a) |
| Killer cell lectin-like receptor C2 | KLRC2 | b) | a) |
| Killer cell lectin-like receptor C3 | KLRC3 | b) | Colon, duodenum, small intestine, stomach, tonsil |
| Killer cell lectin-like receptor C4 | KLRC4 | b) | a) |
| Killer cell lectin-like receptor K1 | KLRK1 | α-(2-3)-NeuAc-containing N-glycans [64], heparin, heparan sulfate [56] | Appendix, lymph node, spleen, tonsil |
| Macrophage Mannose Receptor (MMR) | |||
| Endo180 (Mannose receptor C type 2) | MRC2 | Mannose, fucose, N-acetylglucosamine [65] | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, muscle tissues, pancreas, proximal digestive tract, skin |
| Lymphocyte antigen 75 | LY75 | Predicted to not bind carbohydrates [65] | Appendix, breast, bronchus, cervix (uterine), duodenum, endometrium, fallopian tube, gallbladder, liver, lung, lymph node, nasopharynx, pancreas, placenta, rectum, spleen, stomach, thyroid gland, tonsil, urinary bladder, |
| Mannose receptor C-type 1 c) | MRC1 | Mannose, fucose, glucose, N-acetylglucosamine [66] (C-type) 4-O-sulphated GalNAc (R-type) | Colon, endometrium, kidney, lung, rectum, skin, soft tissue, testis |
| Phospholipase A2 receptor | PLA2R1 | Predicted to not bind carbohydrates [65] but known to bind collagen | Kidney |
| Free C-type Lectin Domains (CTLDs) | |||
| C-type lectin domain containing 19A | CLEC19A | b) | a) |
| Lithostathine-alpha (Regenerating family member 1 alpha) | REG1A | b) | Duodenum, pancreas, small intestine, stomach |
| Lithostathine-beta (Regenerating family member 1 beta) | REG1B | b) | Duodenum, pancreas, small intestine, stomach |
| Regenerating family member 3 alpha | REG3A | Peptidoglycan (binding affinity increases with the length of the carbohydrate moiety) [67] | Appendix, duodenum, skin, small intestine, stomach |
| Regenerating family member 3 gamma | REG3G | Peptidoglycan [67] | a) |
| Regenerating family member 4 | REG4 | Mannans, heparin [67] | Appendix, colon, duodenum, rectum, small intestine |
| Type I receptors | |||
| Chondrolectin | CHODL | b) [68] | Appendix, colon, duodenum, rectum, small intestine, testis |
| Layilin | LAYN | Hyaluronan [69] | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, muscle tissues, pancreas, proximal digestive tract, skin |
| Tetranectin | |||
| Cartilage-derived C-type lectin (C-type lectin domain family 3 member A) | CLEC3A | Expected to bind sulfated polysaccharides such as heparin [70] | a) |
| Stem cell growth factor (SCGF) (C-type lectin domain containing 11A) | CLEC11A | b) | Bone marrow, soft tissue |
| Tetranectin (C-type lectin domain family 3 member B) | CLEC3B | Sulfated polysaccharides such as heparin [70] | a) |
| Polycystin | |||
| Polycystin 1 like 3, transient receptor potential channel interacting | PKD1L3 | Predicted to not bind carbohydrates | a) |
| Polycystin 1, transient receptor potential channel interacting | PKD1 | Predicted to bind galactosyl and glucosyl residues. Might bind oligosaccharides with mannosyl moieties [71] | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, pancreas, proximal digestive tract, skin |
| Attractin | |||
| Attractin | ATRN | b) | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, endocrine tissues, female tissues, gastrointestinal tract, kidney and urinary bladder, lung, male tissues, pancreas, proximal digestive tract, skin |
| Attractin-like 1 | ATRNL1 | b) | a) |
| CTLD/acidic neck | |||
| CD302 molecule | CD302 | b) [72] | a) |
| Proteoglycan 2, pro eosinophil major basic protein | PRG2 | Heparin [73] | Bone marrow, placenta |
| Proteoglycan 3, pro eosinophil major basic protein 2 | PRG3 | b) | Bone marrow |
| Endosialin | |||
| CD93 molecule | CD93 | b) | Bone marrow, brain, colon, kidney, lung, spleen |
| C-type lectin domain containing 14A | CLEC14A | b) | Appendix, brain, cervix (uterine), colon, duodenum, esophagus, gallbladder, heart muscle, kidney, lung, pancreas, prostate, rectum, skin, small intestine, stomach, testis |
| Endosialin (CD248 molecule) | CD248 | b) | Adipose and soft tissue, bone marrow and lymphoid tissues, brain, female tissues, gastrointestinal tract, kidney and urinary bladder, muscle tissues, pancreas, skin |
| Thrombomodulin | THBD | b) | Cervix (uterine), colon, esophagus, lymph node, oral mucosa, placenta, skin, tonsil, urinary bladder, vagina |
| Others | |||
| C-type lectin domain family 18 member A | CLEC18A | Fucoidan, β-glucans, β-galactans [74] | a) |
| Prolectin (C-type lectin domain containing 17A) | CLEC17A | Terminal α-d-mannose and fucose residues [75] | Appendix, lymph node, spleen, stomach, tonsil |
| DiGeorge syndrome critical region gene 2 | DGCR2 | b) | Pancreas |
| FRAS1 related extracellular matrix 1 | FREM1 | b) | Adrenal gland, appendix, colon, duodenum, epididymis, kidney, lung, pancreas, placenta, rectum, salivary gland, small intestine, stomach, testis, tonsil, thyroid gland |