Table 1.

A selection of helminth glycan structures involved in innate and adaptive immunity.

Glycan	Structure	Species	Antigenic?	Receptors	Effects/functions	Reference
LN Lewis x (Lex)		S. mansoni* S. mansoni*	No Yes	DC-SIGN, MR (weak)	Granuloma induction B cell proliferation, IL-10 production, TH2 bias, immunosuppressive, antibodies can mediate complement lysis of leukocytes	van de Vijver et al. (227) Srivatsan et al. (59), Velupillai and Harn (102), Nyame et al. (63), van Die et al. (117), Meevissen et al. (120)
Poly-Lex		S. mansoni*	Yes			Srivatsan et al. (59), van Roon et al. (228), Mandalasi et al. (60)
LDN LDNF		S. mansoni* Schistosoma spp.*, H. contortus, T. spiralis	Yes Yes	MGL; galectin-3 DC-SIGN; MGL; MR; CD62E (E-selectin)	Granuloma induction; possible molecular mimicry with snail host; antibodies to LDN lyse schistosomula in vitro DC maturation; antibodies correlate with protection to H. contortus	Srivatsan et al. (27), Neeleman et al. (28), Nyame et al. (122, 224), van den Berg et al. (100), van Vliet et al. (119), van de Vijver et al. (227), Meevissen et al. (120), Yoshino et al. (284) Srivatsan et al. (27), Nyame et al. (223), van Die et al. (117), van Vliet et al. (119), van Liempt et al. (118), Meevissen et al. (120), van Stijn et al. (128)
Poly-LDNF		S. mansoni	Yes	DC-SIGN		Kawar et al. (289), van Liempt et al. (118), Wuhrer et al. (48, 229)
FLDN LDN-DF		S. mansoni S. mansoni	Yes Yes	DC-SIGN	Stimulates IL-10, IL-6, and TNF-α production by PBMC	Naus et al. (234), van Remoortere et al. (233), de Boer et al. (236), Meevissen et al. (120), Frank et al. (51) van der Kleij et al. (116), Naus et al. (234), Frank et al. (51), van Remoortere et al. (233)
FLDNF		S. mansoni	Yes		Antibodies to FLDNF are protective in rats	Geyer et al. (50), Grzych et al. (171, 221, 222), Wuhrer et al. (46), Kantelhardt et al. (49)
DF-LDN-DF		S. mansoni	Yes		Diagnostic in urine	Robijn et al. (243, 250)
Core β2 Xyl, core α3 Fuc		S. mansoni	Yes		Th2 biasing of DCs	Faveeuw et al. (114), Meevissen et al. (120)
Circulating cathodic antigen (CCA)		S. mansoni	Yes		Diagnostic (urine, sera)	Deelder et al. (65), van Dam et al. (245)
Circulating anodic antigen (CAA)		S. mansoni	Yes		Diagnostic (urine, sera), forms antibody-antigen complexes	Deelder et al. (65), Vermeer et al. (244)
Tyvelose		T. spiralis	Yes		Antibodies to tyvelose are protective and diagnostic	Ellis et al. (30), Reason et al. (31), McVay et al. (215), Bolás-Fernandez and Corral Bezara (72)
Galα1-3GalNAc		H. contortus	Yes		Antibodies to Galα1-3GalNAc are protective	van Stijn et al. (257)
Gal-Fuc		C. elegans, A. suum	Yes	Endogenous and fungal galectins; human Gal-1	Fungal CGL2 kills C. elegans	Yan et al. (54), Butschi et al. (80), Takeuchi et al. (57, 58)
Galα1-4Galβ1-3GalNAc		Echinococcus spp.	Yes		Diagnostic	Koizumi et al. (254), Díaz et al. (255)
PC-glycan		Filarial and GI nematodes, E. granulosus	Yes		Anti-inflammatory (both Th1 and Th2); nematode development	Fletcher et al. (41), Peters et al. (217), Paschinger et al. (37), Rzepecka et al. (142), Grabitzki et al. (39)
Methylated Fuc/Gal		Toxocara spp.	Yes			Khoo et al. (43), Koizumi et al. (44)
PC-glycolipids		Ascaris spp.	Yes		Inhibits LPS-induced B cell proliferation and macrophage, IL-12 release; induces PBMC to produce Th1 cytokines	Lochnit et al. (132), Deehan et al. (133), van Riet et al. (134)

Defined determinants (common names are indicated) recognized by either antibodies or glycan-binding proteins and lectins are indicated in the blue background boxes, along with the known helminths expressing the determinant. Note that many structures are composite examples, and that some complex and branched glycans may possess one or more of these determinants, such as outer branch LDN-DF and inner core Xyl and Fuc residues. The asterisk indicates glycan motifs that can also be made in mammalian hosts, but are not necessarily cross reactive, due to differences in surrounding structures. Their antigenicity is also noted if they have been confirmed as antibody targets, as well as any known receptors and demonstrated functions, in vitro or in vivo, with selected corresponding references.