{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,26]],"date-time":"2026-09-26T03:04:26Z","timestamp":1790391866208,"version":"4.1.0"},"reference-count":131,"publisher":"MDPI AG","issue":"3","license":[{"start":{"date-parts":[[2018,3,3]],"date-time":"2018-03-03T00:00:00Z","timestamp":1520035200000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001809","name":"National Science Foundation of China","doi-asserted-by":"publisher","award":["31530069"],"award-info":[{"award-number":["31530069"]}],"id":[{"id":"10.13039\/501100001809","id-type":"DOI","asserted-by":"publisher"}]},{"name":"AoShan Talents Cultivation Program Supported by Qingdao National Laboratory for Marine Science and Technology","award":["2017ASTCP-OS13"],"award-info":[{"award-number":["2017ASTCP-OS13"]}]},{"name":"Modern Agro-industry Technology Research System","award":["CARS-49"],"award-info":[{"award-number":["CARS-49"]}]},{"name":"Dalian High Level Talent Innovation Support Program","award":["2015R020"],"award-info":[{"award-number":["2015R020"]}]},{"name":"Talented Scholars in Dalian Ocean University"},{"name":"Natural Science Foundation of Liaoning","award":["20170520056"],"award-info":[{"award-number":["20170520056"]}]}],"content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["IJMS"],"abstract":"<jats:p>Self-nonself discrimination is a common theme for all of the organisms in different evolutionary branches, which is also the most fundamental step for host immune protection. Plenty of pattern recognition receptors (PRRs) with great diversity have been identified from different organisms to recognize various pathogen-associated molecular patterns (PAMPs) in the last two decades, depicting a complicated scene of host-pathogen interaction. However, the detailed mechanism of the complicate PAMPs\u2013PRRs interactions at the contacting interface between pathogens and hosts is still not well understood. All of the cells are coated by glycosylation complex and thick carbohydrates layer. The different polysaccharides in extracellular matrix of pathogen-host are important for nonself recognition of most organisms. Coincidentally, massive expansion of PRRs, majority of which contain recognition domains of Ig, leucine-rich repeat (LRR), C-type lectin (CTL), C1q and scavenger receptor (SR), have been annotated and identified in invertebrates by screening the available genomic sequence. The phylum Mollusca is one of the largest groups in the animal kingdom with abundant biodiversity providing plenty of solutions about pathogen recognition and immune protection, which might offer a suitable model to figure out the common rules of immune recognition mechanism. The present review summarizes the diverse PRRs and common elements of various PAMPs, especially focusing on the structural and functional characteristics of canonical carbohydrate recognition proteins and some novel proteins functioning in molluscan immune defense system, with the objective to provide new ideas about the immune recognition mechanisms.<\/jats:p>","DOI":"10.3390\/ijms19030721","type":"journal-article","created":{"date-parts":[[2018,3,6]],"date-time":"2018-03-06T07:37:25Z","timestamp":1520321845000},"page":"721","update-policy":"https:\/\/doi.org\/10.3390\/mdpi_crossmark_policy","source":"Crossref","is-referenced-by-count":80,"title":["Pathogen-Derived Carbohydrate Recognition in Molluscs Immune Defense"],"prefix":"10.3390","volume":"19","author":[{"given":"Weilin","family":"Wang","sequence":"first","affiliation":[{"name":"Liaoning Key Laboratory of Marine Animal Immunology, Dalian Ocean University, Dalian 116023, China"},{"name":"Functional Laboratory of Marine Fisheries Science and Food Production Processes, Qingdao National Laboratory for Marine Science and Technology, Qingdao 266235, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Xiaorui","family":"Song","sequence":"additional","affiliation":[{"name":"Liaoning Key Laboratory of Marine Animal Immunology, Dalian Ocean University, Dalian 116023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lingling","family":"Wang","sequence":"additional","affiliation":[{"name":"Liaoning Key Laboratory of Marine Animal Immunology, Dalian Ocean University, Dalian 116023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Linsheng","family":"Song","sequence":"additional","affiliation":[{"name":"Liaoning Key Laboratory of Marine Animal Immunology, Dalian Ocean University, Dalian 116023, China"},{"name":"Functional Laboratory of Marine Fisheries Science and Food Production Processes, Qingdao National Laboratory for Marine Science and Technology, Qingdao 266235, China"},{"name":"Liaoning Key Laboratory of Marine Animal Immunology and Disease Control, Dalian Ocean University, Dalian 116023, China"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"1968","published-online":{"date-parts":[[2018,3,3]]},"reference":[{"key":"ref_1","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1034\/j.1600-065X.2000.917309.x","article-title":"Innate immune recognition: Mechanisms and pathways","volume":"173","author":"Medzhitov","year":"2000","journal-title":"Immunol. Rev."},{"key":"ref_2","doi-asserted-by":"crossref","first-page":"197","DOI":"10.1146\/annurev.immunol.20.083001.084359","article-title":"Innate immune recognition","volume":"20","author":"Janeway","year":"2002","journal-title":"Annu. Rev. Immunol."},{"key":"ref_3","doi-asserted-by":"crossref","first-page":"52","DOI":"10.1111\/imr.12576","article-title":"Building conventions for unconventional lymphocytes","volume":"279","author":"Pasman","year":"2017","journal-title":"Immunol. Rev."},{"key":"ref_4","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1038\/nri3141","article-title":"How do plants achieve immunity? Defence without specialized immune cells","volume":"12","author":"Spoel","year":"2012","journal-title":"Nat. Rev. Immunol."},{"key":"ref_5","unstructured":"Dantzler, W.H. (2011). Invertebrate Circulatory Systems. Handbook of Physiology, Comparative Physiology, Oxford University Press."},{"key":"ref_6","doi-asserted-by":"crossref","first-page":"371","DOI":"10.1146\/annurev-ecolsys-120213-091627","article-title":"Animal Phylogeny and Its Evolutionary Implications","volume":"45","author":"Dunn","year":"2014","journal-title":"Annu. Rev. Ecol. Evol. Syst."},{"key":"ref_7","doi-asserted-by":"crossref","first-page":"65","DOI":"10.1016\/j.fsi.2015.03.013","article-title":"The immune system and its modulation mechanism in scallop","volume":"46","author":"Song","year":"2015","journal-title":"Fish Shellfish Immun."},{"key":"ref_8","doi-asserted-by":"crossref","first-page":"112","DOI":"10.1016\/j.biosystems.2017.10.003","article-title":"The evolution of the Glycomic Codes of extracellular matrices","volume":"164","author":"Buckeridge","year":"2018","journal-title":"BioSystems"},{"key":"ref_9","doi-asserted-by":"crossref","first-page":"819","DOI":"10.1038\/nature06246","article-title":"Recognition of microorganisms and activation of the immune response","volume":"449","author":"Medzhitov","year":"2007","journal-title":"Nature"},{"key":"ref_10","doi-asserted-by":"crossref","first-page":"102","DOI":"10.1016\/j.biosystems.2017.07.003","article-title":"The sugar code: Why glycans are so important","volume":"164","author":"Gabius","year":"2018","journal-title":"BioSystems"},{"key":"ref_11","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1093\/glycob\/cww086","article-title":"Biological Roles of Glycans","volume":"27","author":"Varki","year":"2017","journal-title":"Glycobiology"},{"key":"ref_12","doi-asserted-by":"crossref","first-page":"524","DOI":"10.1016\/j.tibtech.2004.07.012","article-title":"Sugar-mediated ligand-receptor interactions in the immune system","volume":"22","author":"Rudd","year":"2004","journal-title":"Trends Biotechnol."},{"key":"ref_13","doi-asserted-by":"crossref","first-page":"5723","DOI":"10.1002\/ejoc.201100296","article-title":"Carbohydrates and Immunology: Synthetic Oligosaccharide Antigens for Vaccine Formulation","volume":"2011","author":"Morelli","year":"2011","journal-title":"Eur. J. Org. Chem."},{"key":"ref_14","doi-asserted-by":"crossref","first-page":"567","DOI":"10.1016\/j.sbi.2008.08.001","article-title":"Microbial recognition of human cell surface glycoconjugates","volume":"18","author":"Imberty","year":"2008","journal-title":"Curr. Opin. Struct. Biol."},{"key":"ref_15","doi-asserted-by":"crossref","first-page":"1437","DOI":"10.1042\/BST0351437","article-title":"Building an immune system from nine domains","volume":"35","author":"Palssonmcdermott","year":"2007","journal-title":"Biochem. Soc. Trans."},{"key":"ref_16","doi-asserted-by":"crossref","first-page":"191","DOI":"10.1111\/j.1462-5822.2008.01260.x","article-title":"The leucine-rich repeat domain in plant innate immunity: A wealth of possibilities","volume":"11","author":"Padmanabhan","year":"2009","journal-title":"Cell. Microbiol."},{"key":"ref_17","doi-asserted-by":"crossref","first-page":"8693","DOI":"10.1038\/srep08693","article-title":"Massive expansion and functional divergence of innate immune genes in a protostome","volume":"5","author":"Zhang","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_18","doi-asserted-by":"crossref","first-page":"24","DOI":"10.1016\/j.dci.2014.02.004","article-title":"Multigene families of immunoglobulin domain-containing innate immune receptors in zebrafish: Deciphering the differences","volume":"46","author":"Rodrigueznunez","year":"2014","journal-title":"Dev. Comp. Immunol."},{"key":"ref_19","doi-asserted-by":"crossref","first-page":"3312","DOI":"10.1007\/s11434-012-5155-6","article-title":"Sequence variability of fibrinogen-related proteins (FREPs) in Crassostrea gigas","volume":"57","author":"Zhang","year":"2012","journal-title":"Chin. Sci. Bull."},{"key":"ref_20","doi-asserted-by":"crossref","first-page":"111","DOI":"10.1007\/978-3-319-20819-0_5","article-title":"Fibrinogen-Related Proteins (FREPs) in Mollusks","volume":"57","author":"Adema","year":"2015","journal-title":"Results Probl. Cell Differ."},{"key":"ref_21","doi-asserted-by":"crossref","first-page":"4313","DOI":"10.1242\/jeb.085191","article-title":"Variable immune molecules in invertebrates","volume":"216","author":"Cerenius","year":"2013","journal-title":"J. Exp. Biol."},{"key":"ref_22","doi-asserted-by":"crossref","first-page":"155","DOI":"10.1016\/S0378-1119(01)00444-9","article-title":"Structure of two FREP genes that combine IgSF and fibrinogen domains, with comments on diversity of the FREP gene family in the snail Biomphalaria glabrata","volume":"269","author":"Leonard","year":"2001","journal-title":"Gene"},{"key":"ref_23","doi-asserted-by":"crossref","first-page":"485","DOI":"10.1016\/j.fsi.2015.01.021","article-title":"Highly diverse fibrinogen-related proteins in the Pacific oyster Crassostrea gigas","volume":"43","author":"Huang","year":"2015","journal-title":"Fish Shellfish Immun."},{"key":"ref_24","doi-asserted-by":"crossref","first-page":"1119","DOI":"10.1016\/j.dci.2008.03.001","article-title":"Fibrinogen-bearing protein genes in the snail Biomphalaria glabrata: Characterization of two novel genes and expression studies during ontogenesis and trematode infection","volume":"32","author":"Zhang","year":"2008","journal-title":"Dev. Comp. Immunol."},{"key":"ref_25","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1016\/j.gene.2004.07.003","article-title":"Representation of an immune responsive gene family encoding fibrinogen-related proteins in the freshwater mollusc Biomphalaria glabrata, an intermediate host for Schistosoma mansoni","volume":"341","author":"Zhang","year":"2004","journal-title":"Gene"},{"key":"ref_26","doi-asserted-by":"crossref","first-page":"465","DOI":"10.1016\/j.dci.2009.12.007","article-title":"In search of the origin of FREPs: Characterization of Aplysia californica fibrinogen-related proteins","volume":"34","author":"Gorbushin","year":"2010","journal-title":"Dev. Comp. Immunol."},{"key":"ref_27","doi-asserted-by":"crossref","first-page":"526","DOI":"10.1038\/nature11696","article-title":"Insights into bilaterian evolution from three spiralian genomes","volume":"493","author":"Simakov","year":"2012","journal-title":"Nature"},{"key":"ref_28","doi-asserted-by":"crossref","first-page":"251","DOI":"10.1126\/science.1088069","article-title":"Diversification of Ig Superfamily Genes in an Invertebrate","volume":"305","author":"Zhang","year":"2004","journal-title":"Science"},{"key":"ref_29","doi-asserted-by":"crossref","first-page":"175","DOI":"10.1177\/1753425908093800","article-title":"Expression profiling and binding properties of fibrinogen-related proteins (FREPs), plasma proteins from the schistosome snail host Biomphalaria glabrata","volume":"14","author":"Zhang","year":"2008","journal-title":"Innate Immun."},{"key":"ref_30","doi-asserted-by":"crossref","first-page":"17","DOI":"10.1159\/000321882","article-title":"The Primary Role of Fibrinogen-Related Proteins in Invertebrates Is Defense, Not Coagulation","volume":"3","author":"Hanington","year":"2011","journal-title":"J. Innate Immun."},{"key":"ref_31","doi-asserted-by":"crossref","first-page":"334","DOI":"10.1016\/j.dci.2010.10.007","article-title":"Individual sequence variability and functional activities of fibrinogen-related proteins (FREPs) in the Mediterranean mussel (Mytilus galloprovincialis) suggest ancient and complex immune recognition models in invertebrates","volume":"35","author":"Romero","year":"2011","journal-title":"Dev. Comp. Immunol."},{"key":"ref_32","doi-asserted-by":"crossref","first-page":"56","DOI":"10.1016\/j.fsi.2008.07.019","article-title":"A fibrinogen-related protein from bay scallop Argopecten irradians involved in innate immunity as pattern recognition receptor","volume":"26","author":"Zhang","year":"2009","journal-title":"Fish Shellfish Immun."},{"key":"ref_33","doi-asserted-by":"crossref","first-page":"221","DOI":"10.1016\/j.fsi.2014.03.025","article-title":"A new fibrinogen-related protein from Argopecten irradians (AiFREP-2) with broad recognition spectrum and bacteria agglutination activity","volume":"38","author":"Yang","year":"2014","journal-title":"Fish Shellfish Immun."},{"key":"ref_34","doi-asserted-by":"crossref","first-page":"810","DOI":"10.1093\/glycob\/cwp063","article-title":"Natural ligands for CD33-related Siglecs?","volume":"19","author":"Varki","year":"2009","journal-title":"Glycobiology"},{"key":"ref_35","doi-asserted-by":"crossref","first-page":"922","DOI":"10.1093\/glycob\/cwm065","article-title":"Human-specific expression of Siglec-6 in the placenta","volume":"17","author":"Hayakawa","year":"2007","journal-title":"Glycobiology"},{"key":"ref_36","doi-asserted-by":"crossref","first-page":"136","DOI":"10.1016\/j.dci.2016.03.026","article-title":"A novel siglec (CgSiglec-1) from the Pacific oyster (Crassostrea gigas) with broad recognition spectrum and inhibitory activity to apoptosis, phagocytosis and cytokine release","volume":"61","author":"Liu","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"ref_37","doi-asserted-by":"crossref","first-page":"196","DOI":"10.1111\/j.1365-2567.2009.03100.x","article-title":"Junctional adhesion molecule (JAM)-B supports lymphocyte rolling and adhesion through interaction with \u03b14\u03b21 integrin","volume":"128","author":"Ludwig","year":"2009","journal-title":"Immunology"},{"key":"ref_38","doi-asserted-by":"crossref","first-page":"211","DOI":"10.1016\/j.dci.2015.09.011","article-title":"A novel junctional adhesion molecule A (CgJAM-A-L) from oyster (Crassostrea gigas) functions as pattern recognition receptor and opsonin","volume":"55","author":"Liu","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"ref_39","doi-asserted-by":"crossref","first-page":"4631","DOI":"10.1073\/pnas.1000093107","article-title":"Human leucine-rich repeat proteins: A genome-wide bioinformatic categorization and functional analysis in innate immunity","volume":"108","author":"Ng","year":"2011","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_40","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1016\/j.dci.2015.09.006","article-title":"An LRR-only protein representing a new type of pattern recognition receptor in Chlamys farreri","volume":"54","author":"Wang","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"ref_41","doi-asserted-by":"crossref","first-page":"952","DOI":"10.1126\/science.1134301","article-title":"Genomic Insights into the Immune System of the Sea Urchin","volume":"314","author":"Rast","year":"2006","journal-title":"Science"},{"key":"ref_42","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1093\/bfgp\/els007","article-title":"The amphioxus genome provides unique insight into the evolution of immunity","volume":"11","author":"Dishaw","year":"2012","journal-title":"Brief Funct. Genom."},{"key":"ref_43","doi-asserted-by":"crossref","first-page":"1082","DOI":"10.4161\/auto.7.9.16464","article-title":"Leucine-rich repeat (LRR) proteins: Integrators of pattern recognition and signaling in immunity","volume":"7","author":"Ng","year":"2011","journal-title":"Autophagy"},{"key":"ref_44","doi-asserted-by":"crossref","first-page":"1367","DOI":"10.4049\/jimmunol.0903128","article-title":"Alternative Adaptive Immunity in Jawless Vertebrates","volume":"185","author":"Herrin","year":"2010","journal-title":"J. Immunol."},{"key":"ref_45","doi-asserted-by":"crossref","first-page":"1112","DOI":"10.1101\/gr.069674.107","article-title":"Genomic analysis of the immune gene repertoire of amphioxus reveals extraordinary innate complexity and diversity","volume":"18","author":"Huang","year":"2008","journal-title":"Genome Res."},{"key":"ref_46","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1093\/intimm\/dxh186","article-title":"Toll-like receptors in innate immunity","volume":"17","author":"Takeda","year":"2004","journal-title":"Int. Immunol."},{"key":"ref_47","doi-asserted-by":"crossref","first-page":"588","DOI":"10.1073\/pnas.95.2.588","article-title":"A family of human receptors structurally related to Drosophila Toll","volume":"95","author":"Rock","year":"1998","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_48","doi-asserted-by":"crossref","first-page":"182","DOI":"10.1016\/j.dci.2016.07.010","article-title":"A new non-phagocytic TLR6 with broad recognition ligands from Pacific oyster Crassostrea gigas","volume":"65","author":"Wang","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"ref_49","doi-asserted-by":"crossref","unstructured":"Zhang, Y., He, X., Yu, F., Xiang, Z., Li, J., Thorpe, K.L., and Yu, Z. (2013). Characteristic and Functional Analysis of Toll-like Receptors (TLRs) in the lophotrocozoan, Crassostrea gigas, Reveals Ancient Origin of TLR-Mediated Innate Immunity. PLoS ONE, 8.","DOI":"10.1371\/journal.pone.0076464"},{"key":"ref_50","doi-asserted-by":"crossref","first-page":"1287","DOI":"10.1016\/j.fsi.2013.02.014","article-title":"The first Toll receptor from the triangle-shell pearl mussel Hyriopsis cumingii","volume":"34","author":"Ren","year":"2013","journal-title":"Fish Shellfish Immun."},{"key":"ref_51","doi-asserted-by":"crossref","first-page":"198","DOI":"10.1016\/j.dci.2014.02.020","article-title":"A novel Toll like receptor with two TIR domains (HcToll-2) is involved in regulation of antimicrobial peptide gene expression of Hyriopsis cumingii","volume":"45","author":"Ren","year":"2014","journal-title":"Dev. Comp. Immunol."},{"key":"ref_52","doi-asserted-by":"crossref","first-page":"189","DOI":"10.1016\/j.fsi.2017.02.015","article-title":"HcToll3 was involved in anti-Vibrio defense in freshwater pearl mussel, Hyriopsis cumingii","volume":"63","author":"Zhang","year":"2017","journal-title":"Fish Shellfish Immun."},{"key":"ref_53","doi-asserted-by":"crossref","first-page":"511","DOI":"10.1016\/j.dci.2010.12.005","article-title":"A primitive Toll-like receptor signaling pathway in mollusk Zhikong scallop Chlamys farreri","volume":"35","author":"Wang","year":"2011","journal-title":"Dev. Comp. Immunol."},{"key":"ref_54","doi-asserted-by":"crossref","first-page":"205","DOI":"10.1016\/j.fsi.2017.12.064","article-title":"The various components implied the diversified Toll-like receptor (TLR) signaling pathway in mollusk Chlamys farreri","volume":"74","author":"Wang","year":"2018","journal-title":"Fish Shellfish Immun."},{"key":"ref_55","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.dci.2016.01.013","article-title":"Two novel LRR-only proteins in Chlamys farreri: Similar in structure, yet different in expression profile and pattern recognition","volume":"59","author":"Wang","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"ref_56","doi-asserted-by":"crossref","first-page":"188","DOI":"10.1016\/j.dci.2017.08.005","article-title":"The versatile functions of LRR-only proteins in mollusk Chlamys farreri","volume":"77","author":"Wang","year":"2017","journal-title":"Dev. Comp. Immunol."},{"key":"ref_57","doi-asserted-by":"crossref","first-page":"2753","DOI":"10.1074\/JBC.X600004200","article-title":"Lectins: Carbohydrate-specific Reagents and Biological Recognition Molecules","volume":"282","author":"Sharon","year":"2007","journal-title":"J. Biol. Chem."},{"key":"ref_58","doi-asserted-by":"crossref","first-page":"6179","DOI":"10.1111\/j.1742-4658.2005.05031.x","article-title":"The C-type lectin-like domain superfamily","volume":"272","author":"Zelensky","year":"2005","journal-title":"FEBS J."},{"key":"ref_59","first-page":"241","article-title":"The immune role of C-type lectins in molluscs","volume":"8","author":"Wang","year":"2011","journal-title":"ISJ-Invert. Surviv. J."},{"key":"ref_60","doi-asserted-by":"crossref","first-page":"19","DOI":"10.1111\/j.1600-065X.1998.tb01185.x","article-title":"The C-type lectin superfamily in the immune system","volume":"163","author":"Weis","year":"1998","journal-title":"Immunol. Rev."},{"key":"ref_61","doi-asserted-by":"crossref","first-page":"139","DOI":"10.1007\/s11033-013-2846-2","article-title":"Molecular cloning and expression of a C-type lectin gene from Venerupis philippinarum","volume":"41","author":"Mu","year":"2014","journal-title":"Mol. Biol. Rep."},{"key":"ref_62","doi-asserted-by":"crossref","first-page":"1034","DOI":"10.1016\/j.dci.2008.02.001","article-title":"A novel C-type lectin from abalone, Haliotis discus discus, agglutinates Vibrio alginolyticus","volume":"32","author":"Wang","year":"2008","journal-title":"Dev. Comp. Immunol."},{"key":"ref_63","doi-asserted-by":"crossref","first-page":"466","DOI":"10.1016\/j.fsi.2014.09.032","article-title":"HdhCTL1 is a novel C-type lectin of abalone Haliotis discus hannai that agglutinates Gram-negative bacterial pathogens","volume":"41","author":"Zhang","year":"2014","journal-title":"Fish Shellfish Immun."},{"key":"ref_64","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/j.fsi.2016.06.007","article-title":"A novel C-type lectin with four CRDs is involved in the regulation of antimicrobial peptide gene expression in Hyriopsis cumingii","volume":"55","author":"Zhao","year":"2016","journal-title":"Fish Shellfish Immun."},{"key":"ref_65","doi-asserted-by":"crossref","first-page":"210","DOI":"10.1016\/j.gene.2013.05.029","article-title":"Molecular cloning and characterization of perlucin from the freshwater pearl mussel, Hyriopsis cumingii","volume":"526","author":"Lin","year":"2013","journal-title":"Gene"},{"key":"ref_66","doi-asserted-by":"crossref","first-page":"598","DOI":"10.1016\/j.fsi.2008.07.015","article-title":"Characterization, tissue expression, and immunohistochemical localization of MCL3, a C-type lectin produced by Perkinsus olseni-infected Manila clams (Ruditapes philippinarum)","volume":"25","author":"Kim","year":"2008","journal-title":"Fish Shellfish Immun."},{"key":"ref_67","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1016\/j.cbpb.2008.01.006","article-title":"Purification and antibacterial characterization of a novel isoform of the Manila clam lectin (MCL-4) from the plasma of the Manila clam, Ruditapes philippinarum","volume":"150","author":"Takahashi","year":"2008","journal-title":"Comp. Biochem. Phys. B"},{"key":"ref_68","doi-asserted-by":"crossref","first-page":"1119","DOI":"10.1016\/j.dci.2006.03.005","article-title":"Analysis of EST and lectin expressions in hemocytes of Manila clams (Ruditapes philippinarum) (Bivalvia: Mollusca) infected with Perkinsus olseni","volume":"30","author":"Kang","year":"2006","journal-title":"Dev. Comp. Immunol."},{"key":"ref_69","doi-asserted-by":"crossref","first-page":"99","DOI":"10.1016\/j.dci.2017.05.025","article-title":"The oyster immunity","volume":"80","author":"Wang","year":"2018","journal-title":"Dev. Comp. Immunol."},{"key":"ref_70","doi-asserted-by":"crossref","first-page":"780","DOI":"10.1016\/j.dci.2009.01.003","article-title":"Cflec-4, a multidomain C-type lectin involved in immune defense of Zhikong scallop Chlamys farreri","volume":"33","author":"Zhang","year":"2009","journal-title":"Dev. Comp. Immunol."},{"key":"ref_71","doi-asserted-by":"crossref","first-page":"10068","DOI":"10.1038\/srep10068","article-title":"CfLec-3 from scallop: An entrance to non-self recognition mechanism of invertebrate C-type lectin","volume":"5","author":"Yang","year":"2015","journal-title":"Sci. Rep."},{"key":"ref_72","doi-asserted-by":"crossref","first-page":"591","DOI":"10.1016\/j.dci.2011.10.002","article-title":"A multi-CRD C-type lectin with broad recognition spectrum and cellular adhesion from Argopecten irradians","volume":"36","author":"Wang","year":"2012","journal-title":"Dev. Comp. Immunol."},{"key":"ref_73","doi-asserted-by":"crossref","first-page":"183","DOI":"10.1038\/360183a0","article-title":"Engineering galactose-binding activity into a C-type mannose-binding protein","volume":"360","author":"Drickamer","year":"1992","journal-title":"Nature"},{"key":"ref_74","doi-asserted-by":"crossref","first-page":"495","DOI":"10.1016\/j.cbpa.2010.04.004","article-title":"Identification and molecular characterization of a mucosal lectin (MeML) from the blue mussel Mytilus edulis and its potential role in particle capture","volume":"156","author":"Espinosa","year":"2010","journal-title":"Comp. Biochem. Phys. A"},{"key":"ref_75","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1007\/s10719-011-9326-5","article-title":"Lectins influence chondrogenesis and osteogenesis in limb bud mesenchymal cells","volume":"28","author":"Monsefi","year":"2011","journal-title":"Glycoconj. J."},{"key":"ref_76","doi-asserted-by":"crossref","first-page":"286","DOI":"10.1016\/j.fsi.2007.11.014","article-title":"A lectin (CfLec-2) aggregating Staphylococcus haemolyticus from scallop Chlamys farreri","volume":"24","author":"Zheng","year":"2008","journal-title":"Fish Shellfish Immun."},{"key":"ref_77","doi-asserted-by":"crossref","first-page":"707","DOI":"10.1016\/j.fsi.2009.02.017","article-title":"A novel C-type lectin (Cflec-3) from Chlamys farreri with three carbohydrate-recognition domains","volume":"26","author":"Zhang","year":"2009","journal-title":"Fish Shellfish Immun."},{"key":"ref_78","doi-asserted-by":"crossref","first-page":"333","DOI":"10.1016\/j.dci.2016.08.019","article-title":"The sequence variation and functional differentiation of CRDs in a scallop multiple CRDs containing lectin","volume":"67","author":"Huang","year":"2017","journal-title":"Dev. Comp. Immunol."},{"key":"ref_79","doi-asserted-by":"crossref","unstructured":"Yang, J., Wang, L., Zhang, H., Qiu, L., Wang, H., and Song, L. (2011). C-Type Lectin in Chlamys farreri (CfLec-1) Mediating Immune Recognition and Opsonization. PLoS ONE, 6.","DOI":"10.1371\/journal.pone.0017089"},{"key":"ref_80","doi-asserted-by":"crossref","first-page":"1274","DOI":"10.1016\/j.dci.2010.07.004","article-title":"An ancient C-type lectin in Chlamys farreri (CfLec-2) that mediate pathogen recognition and cellular adhesion","volume":"34","author":"Yang","year":"2010","journal-title":"Dev. Comp. Immunol."},{"key":"ref_81","doi-asserted-by":"crossref","first-page":"26854","DOI":"10.1074\/jbc.M601251200","article-title":"Lectin from the Manila Clam Ruditapes philippinarum Is Induced upon Infection with the Protozoan Parasite Perkinsus olseni","volume":"281","author":"Kim","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_82","doi-asserted-by":"crossref","first-page":"220","DOI":"10.1016\/j.fsi.2016.10.030","article-title":"Comparative study of two single CRD C-type lectins, CgCLec-4 and CgCLec-5, from pacific oyster Crassostrea gigas","volume":"59","author":"Jia","year":"2016","journal-title":"Fish Shellfish Immun."},{"key":"ref_83","doi-asserted-by":"crossref","first-page":"3086","DOI":"10.4049\/jimmunol.179.5.3086","article-title":"A Galectin of Unique Domain Organization from Hemocytes of the Eastern Oyster (Crassostrea virginica) Is a Receptor for the Protistan Parasite Perkinsus marinus","volume":"179","author":"Tasumi","year":"2007","journal-title":"J. Immunol."},{"key":"ref_84","doi-asserted-by":"crossref","first-page":"229","DOI":"10.1016\/j.margen.2011.06.004","article-title":"A tandem-repeat galectin involved in innate immune response of the pearl oyster Pinctada fucata","volume":"4","author":"Wang","year":"2011","journal-title":"Mar. Genom."},{"key":"ref_85","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.dci.2010.08.007","article-title":"A multidomain galectin involved in innate immune response of pearl oyster Pinctada fucata","volume":"35","author":"Zhang","year":"2011","journal-title":"Dev. Comp. Immunol."},{"key":"ref_86","doi-asserted-by":"crossref","first-page":"326","DOI":"10.1016\/j.fsi.2009.11.016","article-title":"An immune responsive multidomain galectin from bay scallop Argopectens irradians","volume":"28","author":"Song","year":"2010","journal-title":"Fish Shellfish Immun."},{"key":"ref_87","doi-asserted-by":"crossref","first-page":"592","DOI":"10.1016\/j.dci.2011.01.006","article-title":"A galectin with quadruple-domain from bay scallop Argopecten irradians is involved in innate immune response","volume":"35","author":"Song","year":"2011","journal-title":"Dev. Comp. Immunol."},{"key":"ref_88","doi-asserted-by":"crossref","first-page":"1131","DOI":"10.1016\/j.dci.2008.03.002","article-title":"Noble tandem-repeat galectin of Manila clam Ruditapes philippinarum is induced upon infection with the protozoan parasite Perkinsus olseni","volume":"32","author":"Kim","year":"2008","journal-title":"Dev. Comp. Immunol."},{"key":"ref_89","doi-asserted-by":"crossref","first-page":"750","DOI":"10.1016\/j.fsi.2010.12.025","article-title":"F-type lectin involved in defense against bacterial infection in the pearl oyster (Pinctada martensii)","volume":"30","author":"Chen","year":"2011","journal-title":"Fish Shellfish Immun."},{"key":"ref_90","doi-asserted-by":"crossref","first-page":"29583","DOI":"10.1074\/jbc.M605687200","article-title":"The Phylogenetically Conserved Molluscan Chitinase-like Protein 1 (Cg-Clp1), Homologue of Human HC-gp39, Stimulates Proliferation and Regulates Synthesis of Extracellular Matrix Components of Mammalian Chondrocytes","volume":"281","author":"Badariotti","year":"2006","journal-title":"J. Biol. Chem."},{"key":"ref_91","doi-asserted-by":"crossref","first-page":"514","DOI":"10.1016\/j.fsi.2014.08.006","article-title":"Characteristic and functional analysis of a ficolin-like protein from the oyster Crassostrea hongkongensis","volume":"40","author":"Xiang","year":"2014","journal-title":"Fish Shellfish Immun."},{"key":"ref_92","doi-asserted-by":"crossref","first-page":"564","DOI":"10.1016\/j.fsi.2017.05.057","article-title":"Molecular and functional characterization of a glycosylated Galactose-Binding lectin from Mytilus californianus","volume":"66","year":"2017","journal-title":"Fish Shellfish Immun."},{"key":"ref_93","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1007\/s10126-017-9728-x","article-title":"Purification, Biochemical Characterization, and Amino Acid Sequence of a Novel Type of Lectin from Aplysia dactylomela Eggs with Antibacterial\/Antibiofilm Potential","volume":"19","author":"Carneiro","year":"2017","journal-title":"Mar. Biotechnol."},{"key":"ref_94","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.imlet.2004.06.015","article-title":"Structural and functional anatomy of the globular domain of complement protein C1q","volume":"95","author":"Kishore","year":"2004","journal-title":"Immunol. Lett."},{"key":"ref_95","doi-asserted-by":"crossref","first-page":"635","DOI":"10.1016\/j.dci.2011.01.018","article-title":"The C1q domain containing proteins of the Mediterranean mussel Mytilus galloprovincialis: A widespread and diverse family of immune-related molecules","volume":"35","author":"Gerdol","year":"2011","journal-title":"Dev. Comp. Immunol."},{"key":"ref_96","doi-asserted-by":"crossref","first-page":"612","DOI":"10.1016\/j.fsi.2016.10.010","article-title":"Identification and function of a novel C1q domain-containing (C1qDC) protein in triangle-shell pearl mussel (Hyriopsis cumingii)","volume":"58","author":"Huang","year":"2016","journal-title":"Fish Shellfish Immun."},{"key":"ref_97","doi-asserted-by":"crossref","first-page":"281","DOI":"10.1016\/j.fsi.2008.06.003","article-title":"A novel C1q-domain-containing protein from Zhikong scallop Chlamys farreri with lipopolysaccharide binding activity","volume":"25","author":"Zhang","year":"2008","journal-title":"Fish Shellfish Immun."},{"key":"ref_98","doi-asserted-by":"crossref","unstructured":"Wang, L., Wang, L., Zhang, H., Zhou, Z., Siva, V., and Song, L. (2012). A C1q Domain Containing Protein from Scallop Chlamys farreri Serving as Pattern Recognition Receptor with Heat-Aggregated IgG Binding Activity. PLoS ONE, 7.","DOI":"10.1371\/journal.pone.0043289"},{"key":"ref_99","doi-asserted-by":"crossref","first-page":"427","DOI":"10.1016\/j.fsi.2012.05.032","article-title":"A novel C1qDC protein acting as pattern recognition receptor in scallop Argopecten irradians","volume":"33","author":"Wang","year":"2012","journal-title":"Fish Shellfish Immun."},{"key":"ref_100","doi-asserted-by":"crossref","first-page":"42","DOI":"10.1016\/j.dci.2017.09.014","article-title":"Comparative study of three C1q domain containing proteins from pacific oyster Crassostrea gigas","volume":"78","author":"Lv","year":"2018","journal-title":"Dev. Comp. Immunol."},{"key":"ref_101","doi-asserted-by":"crossref","first-page":"577","DOI":"10.1189\/jlb.0104014","article-title":"Activation of murine macrophages by Neisseria meningitidis and IFN-\u03b3 in vitro: Distinct roles of class A scavenger and toll-like pattern recognition receptors in selective modulation of surface phenotype","volume":"76","author":"Mukhopadhyay","year":"2004","journal-title":"J. Leukoc. Biol."},{"key":"ref_102","doi-asserted-by":"crossref","first-page":"1160","DOI":"10.1111\/j.1462-5822.2009.01326.x","article-title":"Scavenger receptors: Role in innate immunity and microbial pathogenesis","volume":"11","author":"Areschoug","year":"2009","journal-title":"Cell. Microbiol."},{"key":"ref_103","doi-asserted-by":"crossref","first-page":"227","DOI":"10.1016\/j.dci.2010.09.010","article-title":"A novel scavenger receptor-cysteine-rich (SRCR) domain containing scavenger receptor identified from mollusk mediated PAMP recognition and binding","volume":"35","author":"Liu","year":"2011","journal-title":"Dev. Comp. Immunol."},{"key":"ref_104","doi-asserted-by":"crossref","first-page":"837","DOI":"10.1038\/nri3089","article-title":"Peptidoglycan recognition proteins: Modulators of the microbiome and inflammation","volume":"11","author":"Royet","year":"2011","journal-title":"Nat. Rev. Immunol."},{"key":"ref_105","doi-asserted-by":"crossref","first-page":"684","DOI":"10.1073\/pnas.0507656103","article-title":"Dual strategies for peptidoglycan discrimination by peptidoglycan recognition proteins (PGRPs)","volume":"103","author":"Swaminathan","year":"2006","journal-title":"Proc. Natl. Acad. Sci. USA"},{"key":"ref_106","doi-asserted-by":"crossref","first-page":"646","DOI":"10.1016\/j.fsi.2007.01.023","article-title":"Molecular cloning and characterization of a short type peptidoglycan recognition protein (CfPGRP-S1) cDNA from Zhikong scallop Chlamys farreri","volume":"23","author":"Su","year":"2007","journal-title":"Fish Shellfish Immun."},{"key":"ref_107","doi-asserted-by":"crossref","first-page":"1300","DOI":"10.1016\/j.dci.2010.08.006","article-title":"Peptidoglycan recognition protein of Chlamys farreri (CfPGRP-S1) mediates immune defenses against bacterial infection","volume":"34","author":"Yang","year":"2010","journal-title":"Dev. Comp. Immunol."},{"key":"ref_108","doi-asserted-by":"crossref","first-page":"1768","DOI":"10.1016\/j.molimm.2009.01.022","article-title":"A novel peptidoglycan recognition protein containing a goose-type lysozyme domain from the Pacific oyster, Crassostrea gigas","volume":"46","author":"Itoh","year":"2009","journal-title":"Mol. Immunol."},{"key":"ref_109","doi-asserted-by":"crossref","first-page":"30","DOI":"10.1016\/j.dci.2013.10.011","article-title":"Involvement of Pacific oyster CgPGRP-S1S in bacterial recognition, agglutination and granulocyte degranulation","volume":"43","author":"Iizuka","year":"2014","journal-title":"Dev. Comp. Immunol."},{"key":"ref_110","doi-asserted-by":"crossref","first-page":"3335","DOI":"10.1007\/s11033-009-9920-9","article-title":"Molecular characterization and expression analysis of lipopolysaccharide and beta-1,3-glucan-binding protein (LGBP) from pearl oyster Pinctada fucata","volume":"37","author":"Zhang","year":"2010","journal-title":"Mol. Biol. Rep."},{"key":"ref_111","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1016\/j.aquaculture.2004.03.012","article-title":"cDNA cloning and mRNA expression of the lipopolysaccharide- and beta-1,3-glucan-binding protein gene from scallop Chlamys farreri","volume":"239","author":"Su","year":"2004","journal-title":"Aquaculture"},{"key":"ref_112","doi-asserted-by":"crossref","first-page":"825","DOI":"10.1016\/j.fsi.2010.07.025","article-title":"CfLGBP, a pattern recognition receptor in Chlamys farreri involved in the immune response against various bacteria","volume":"29","author":"Yang","year":"2010","journal-title":"Fish Shellfish Immun."},{"key":"ref_113","doi-asserted-by":"crossref","first-page":"715","DOI":"10.1016\/j.fsi.2007.11.012","article-title":"Isolation, purification and characterization of \u03b2-1,3-glucan binding protein from the plasma of marine mussel Perna viridis","volume":"24","author":"Jayaraj","year":"2008","journal-title":"Fish Shellfish Immun."},{"key":"ref_114","doi-asserted-by":"crossref","first-page":"1607","DOI":"10.3389\/fimmu.2017.01607","article-title":"DM9 Domain Containing Protein Functions As a Pattern Recognition Receptor with Broad Microbial Recognition Spectrum","volume":"8","author":"Jiang","year":"2017","journal-title":"Front. Immunol."},{"key":"ref_115","doi-asserted-by":"crossref","first-page":"29135","DOI":"10.1038\/srep29135","article-title":"Identification, Characterization, and X-ray Crystallographic Analysis of a Novel Type of Mannose-Specific Lectin CGL1 from the Pacific Oyster Crassostrea gigas","volume":"6","author":"Unno","year":"2016","journal-title":"Sci. Rep."},{"key":"ref_116","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1016\/j.coi.2015.01.007","article-title":"Non-canonical activation of inflammatory caspases by cytosolic LPS in innate immunity","volume":"32","author":"Yang","year":"2015","journal-title":"Curr. Opin. Immunol."},{"key":"ref_117","doi-asserted-by":"crossref","first-page":"187","DOI":"10.1038\/nature13683","article-title":"Inflammatory caspases are innate immune receptors for intracellular LPS","volume":"514","author":"Shi","year":"2014","journal-title":"Nature"},{"key":"ref_118","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1016\/j.dci.2016.03.015","article-title":"Caspase-3 serves as an intracellular immune receptor specific for lipopolysaccharide in oyster Crassostrea gigas","volume":"61","author":"Xu","year":"2016","journal-title":"Dev. Comp. Immunol."},{"key":"ref_119","doi-asserted-by":"crossref","first-page":"330","DOI":"10.1016\/j.dci.2017.09.002","article-title":"The self-activation and LPS binding activity of executioner caspase-1 in oyster Crassostrea gigas","volume":"77","author":"Lu","year":"2017","journal-title":"Dev. Comp. Immunol."},{"key":"ref_120","first-page":"422","article-title":"Lectin activities of cytokines: Functions and putative carbohydrate-recognition domains","volume":"1572","author":"Cebo","year":"2002","journal-title":"BBA-Biomembr."},{"key":"ref_121","doi-asserted-by":"crossref","first-page":"107","DOI":"10.1007\/978-1-4615-0065-0_8","article-title":"Lectin domains on cytokines","volume":"535","author":"Zanetta","year":"2003","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_122","doi-asserted-by":"crossref","first-page":"49","DOI":"10.1042\/bj3180049","article-title":"Interleukin 2 is a lectin that associates its receptor with the T-cell receptor complex","volume":"318","author":"Zanetta","year":"1996","journal-title":"Biochem. J."},{"key":"ref_123","doi-asserted-by":"crossref","first-page":"5685","DOI":"10.1074\/jbc.M008662200","article-title":"Recombinant Human Interleukins IL-1\u03b1, IL-1\u03b2, IL-4, IL-6, and IL-7 Show Different and Specific Calcium-independent Carbohydrate-binding Properties","volume":"276","author":"Cebo","year":"2001","journal-title":"J. Biol. Chem."},{"key":"ref_124","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1016\/j.dci.2015.08.002","article-title":"CgIL17-5, an ancient inflammatory cytokine in Crassostrea gigas exhibiting the heterogeneity functions compared with vertebrate interleukin17 molecules","volume":"53","author":"Xin","year":"2015","journal-title":"Dev. Comp. Immunol."},{"key":"ref_125","doi-asserted-by":"crossref","first-page":"66","DOI":"10.1007\/978-1-4419-8059-5_4","article-title":"Earthworm Immunity","volume":"708","author":"Bilej","year":"2010","journal-title":"Adv. Exp. Med. Biol."},{"key":"ref_126","doi-asserted-by":"crossref","first-page":"145","DOI":"10.1007\/978-3-642-18670-7_6","article-title":"Functional convergence of invertebrate and vertebrate cytokine-like molecules based on a similar lectin-like activity","volume":"34","author":"Beschin","year":"2004","journal-title":"Prog. Mol. Subcell. Biol."},{"key":"ref_127","doi-asserted-by":"crossref","first-page":"624","DOI":"10.1016\/j.cell.2016.05.076","article-title":"Hexokinase Is an Innate Immune Receptor for the Detection of Bacterial Peptidoglycan","volume":"166","author":"Wolf","year":"2016","journal-title":"Cell"},{"key":"ref_128","doi-asserted-by":"crossref","first-page":"1549","DOI":"10.3389\/fimmu.2017.01549","article-title":"Exploring Non-Metabolic Functions of Glycolytic Enzymes in Immunity","volume":"8","author":"Seki","year":"2017","journal-title":"Front. Immunol."},{"key":"ref_129","doi-asserted-by":"crossref","first-page":"318","DOI":"10.1016\/j.dci.2017.09.001","article-title":"A GTP-dependent Phosphoenolpyruvate Carboxykinase from Crassostrea gigas involved in immune recognition","volume":"77","author":"Lv","year":"2017","journal-title":"Dev. Comp. Immunol."},{"key":"ref_130","doi-asserted-by":"crossref","first-page":"282","DOI":"10.1016\/j.fsi.2016.03.153","article-title":"The modulation of haemolymph arginine kinase on the extracellular ATP induced bactericidal immune responses in the Pacific oyster Crassostrea gigas","volume":"54","author":"Jiang","year":"2016","journal-title":"Fish Shellfish Immun."},{"key":"ref_131","doi-asserted-by":"crossref","first-page":"23R","DOI":"10.1093\/glycob\/cwm005","article-title":"Evolution of carbohydrate antigens\u2014Microbial forces shaping host glycomes?","volume":"17","author":"Bishop","year":"2007","journal-title":"Glycobiology"}],"container-title":["International Journal of Molecular Sciences"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.mdpi.com\/1422-0067\/19\/3\/721\/pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2025,10,11]],"date-time":"2025-10-11T14:57:23Z","timestamp":1760194643000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.mdpi.com\/1422-0067\/19\/3\/721"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2018,3,3]]},"references-count":131,"journal-issue":{"issue":"3","published-online":{"date-parts":[[2018,3]]}},"alternative-id":["ijms19030721"],"URL":"https:\/\/doi.org\/10.3390\/ijms19030721","relation":{},"ISSN":["1422-0067"],"issn-type":[{"value":"1422-0067","type":"electronic"}],"subject":[],"published":{"date-parts":[[2018,3,3]]}}}