. 2009;9(7):465–479. doi: 10.1038/nri2569

Table 1.

C-type lectin receptors, pathogen recognition and signalling

CLR^*	Expression	Glycan PAMPs	Pathogenic (exogenous) ligands	Signalling motif or adaptor	Signalling proteins involved	Immunological outcome	Refs
*Group II CLRs (asialoglycoprotein receptor family)*
DC-SIGN (CD209)	Myeloid DCs	High mannose and fucose (Le^X, Le^Y, Le^A, Le^B)	˙ Mycobacterium tuberculosis, Mycobacterium leprae, BCG, Lactobacilli spp. and Helicobacter pylori ˙ HIV-1, measles virus, dengue virus, SARS coronavirus and filoviruses ˙ Candida albicans ˙ Leishmania spp. ˙ Tick Ixodes scapularis saliva protein Salp15, peanut allergen Ara h1 and Schistosoma mansoni soluble egg antigens	None	LSP1, LARG, RhoA, Ras proteins, RAF1, Src kinases and PAKs	˙ Upregulation of TLR-induced IL-10 production ˙ Induction of T_H1, T_H2, T_H17 and regulatory T cell differentiation^‡ ˙ Inhibition of T_H1 cell differentiation^‡ ˙ Induction of regulatory T cells^‡ ˙ Antigen presentation	3,20,22,26,29,68,85–93
Langerin (CLEC4K, CD207)	Langerhans cells and dermal DC subset	High mannose, fucose (Le^Y, Le^B) and GlcNAc	˙ M. leprae ˙ HIV-1	Putative proline-rich domain	ND	˙ M. leprae endocytosis and antigen presentation ˙ HIV-1 internalization and degradation	84,94,95
MGL (CLEC10A, CD301)	Myeloid DCs and macrophages	Terminal GalNAc (Tn antigen, LDN)	˙ Filoviruses ˙ S. mansoni	None	ND	ND	96
CLEC5A (MDL1)	Monocytes and macrophages	ND	Dengue virus	DAP10 and DAP12	ND	Induction of TNF production	13,69,97
*Group II CLRs (asialoglycoprotein receptor family; dectin 1 subfamily)*
Dectin 1 (CLEC7A)	Myeloid DCs, monocytes, macrophages and B cells	β-1,3-glucan	˙ M. tuberculosis and Mycobacterium abscessus ˙ C. albicans, Aspergillus fumigatus, Pneumocystis carinii, Penicillium marneffei, Coccidioides posadasii and Histoplasma capsulatum	YxxL^§	SYK, PLCγ2, CARD9, BCL-10, MALT1, NIK and RAF1	˙ Induction of T_H1 and T_H17 cell differentiation through induction of IL-1β, IL-6, IL-12 and IL-23 production ˙ Induction of TNF and CXCL2 production ˙ Phagocytosis ˙ LTB₄ synthesis	4,5,17,42,43,59,65,98
MICL (CLEC12A, DCAL2)	Myeloid DCs, monocytes, macrophages and neutrophils	ND	ND	ITIM	SHP1, SHP2 and ERK	Inhibition of TLR4-induced IL-12 production	14,16
CLEC2 (CLEC1B)	Platelets	ND	˙ HIV-1 ˙ Snake venom protein rhodocytin	YxxL^§	SYK, LAT, RAC1 and PLCγ2	ND	99–101
DNGR1 (CLEC9A)	BDCA3⁺ DCs, monocytes and B cells	ND	ND	YxxL^§	SYK	˙ Induction of TNF production ˙ Antigen cross-presentation	78,82,102
CLEC12B	Macrophages	ND	ND	ITIM	SHP1 and SHP2	ND	103
*Group II CLRs (asialoglycoprotein receptor family; DCIR subfamily)*
Dectin 2 (CLEC6A)	Myeloid DCs, pDCs, monocytes, macrophages, B cells and neutrophils	High mannose	˙ M. tuberculosis ˙ C. albicans, Trichophyton rubrum, A. fumigatus, Microsporum audouinii and Paracoccoides brasiliensis ˙ House dust mite Dermatophagoides pteronyssinus allergens	FcRγ	Src kinases and SYK	˙ Induction of TNF and IL-6 production ˙ Cysteinyl leukotriene synthesis	10,58
BDCA2 (CLEC4C, CD303)	pDCs, monocytes, macrophages and neutrophils	ND	ND	FcRγ	SYK, BTK, BLNK and PLCγ2	˙ Inhibition of TLR9-induced type I IFN, TNF and IL-6 production ˙ Upregulation of TLR9-induced IL-10 production	11,31,33
Mincle (CLEC4E)	Myeloid DCs, monocytes and macrophages	α-mannose	Malassezia spp.	FcRγ	SYK and CARD9	˙ Induction of TNF, IL-10 and CXCL2 production	12,18
DCIR (CLEC4A)	Myeloid DCs, pDCs, monocytes, macrophages, B cells and neutrophils	ND	HIV-1	ITIM	SHP1 and SHP2	˙ Inhibition of TLR8-induced TNF and IL-12 production ˙ Inhibition of TLR9-induced TNF and IFNα production	6,15,38
*Group I CLRs (mannose receptor family)*
Mannose receptor (CD206)	Myeloid DCs and macrophages	High mannose, fucose and sulphated sugars (sLe^x)	˙ M. tuberculosis, Mycobacterium kansasii, Francisella tularensis, Klebsiella pneumoniae and Streptococcus pneumoniae ˙ HIV-1 and dengue virus ˙ C. albicans, Cryptococcus neoformans and P. carinii ˙ Leishmania spp.	ND	CDC42, RhoB, PAKs and ROCK1	Phagocytosis and antigen presentation	104–106
DEC205 (LY75, CD205)	Myeloid DCs	ND	ND	ND	ND	Endocytosis and antigen presentation	76
^*Alternative names are provided in brackets.
^‡Effect on T cell differentiation is ligand dependent.
^§In this motif, x denotes any amino acid. BCG, Mycobacterium bovis bacillus Calmette–Guérin; BCL-10, B cell lymphoma 10; BDCA, blood dendritic cell antigen protein; BLNK, B cell linker; BTK, Bruton's tyrosine kinase; CARD9; caspase-recruitment domain family, member 9; CDC42, cyclin-dependent kinase 42; CLEC, C-type lectin domain family member; CLR, C-type lectin receptor; CXCL2, CXC-chemokine ligand 2; DC, dendritic cell; DCIR, DC immunoreceptor; DC-SIGN, DC-specific ICAM3-grabbing non-integrin; dectin, DC-associated C-type lectin; ERK, extracellular signal-regulated kinase; FcRγ, Fc receptor γ-chain; GalNAc, N-acetylgalactosamine; GlcNAc, β1,6-N-acetylglucosamine; IL, interleukin; ITIM, immunoreceptor tyrosine-based inhibitory motif; LARG, leukaemia-associated Rho guanine nucleotide exchange factor; LAT, linker for activation of T cells; LSP1, lymphocyte-specific 1; LTB₄, leukotriene B₄; MALT1, mucosa-associated lymphoid tissue lymphoma translocation gene; MICL, C-type lectin-like receptor; mincle, macrophage-inducible C-type lectin; ND, not determined; NIK, nuclear factor-κB-inducing kinase; PAK, p21-activated kinase; PAMP, pathogen-associated molecular pattern; pDC, plasmacytoid DC; PLCγ2; phospholipase Cγ2; RHO, Ras homologue; ROCK1, Rho-associated coiled-coil-containing protein kinase 1; SARS, severe acute respiratory syndrome; SHP, SH2-domain-containing protein tyrosine phosphatase; sLe, sialyl Lewis; SYK, spleen tyrosine kinase; TLR, Toll-like receptor; T_H, T helper; TNF, tumour necrosis factor.