(2014) 62:10–8. Likewise, challenge with Vibrio harveyi increased expression of MHC II B mRNA in the gill, liver, and spleen of the white bamboo shark (Chiloscyllium plagiosum), similar to teleost fish (252). Innate immunity generates first line and second line immune responses. Schematic of the conventional TCR forms found in all Gnathostomes and NARTCR found in cartilaginous fish. MBL and ficolin genes have not been identified in any cartilaginous fish studied to date, while MASP2 transcripts are lacking in the elephant shark, little skate (Leucoraja erinacea) and catshark (Scyliorhinus canicular) (54, 106, 107). Boudinot P, Zou J, Ota T, Buonocore F, Scapigliati G, Canapa A, et al. (2015) 4:512–24. Isolation and characterization of two major serum proteins from the dogfish, Mustelus canis, C-reactive protein and amyloid P component. Lee V, Huang JL, Lui MF, Malecek K, Ohta Y, Mooers A, et al. Cell Res. Hooper LV, Littman DR, Macpherson AJ. Dev Comp Immunol. Dev Comp Immunol. (2010) 184:844–50. Over 90 fish AMPs have been identified and are characterized as β-defensins, cathelicidins, hepcidins, histone-derived peptides and fish-specific piscidins. (2009) 22:240–73. J Immunol. The IgD heavy chain has not been identified in any teleost fish without the CHμ1 domain (195, 198, 199). Nunez Ortiz N, Gerdol M, Stocchi V, Marozzi C, Randelli E, Bernini C, et al. Hematological studies on the Australian lungfish, Neoceratodus forsteri. doi: 10.1371/journal.pone.0100018, 84. The skin of lobe-finned fish contains keratinocytes, granulocytes and B cells (20). IgD has only been found in a transmembrane form, with the exception of the channel catfish and the Japanese puffer (Takifugu rubripes), which contain both membrane and secretory forms (159). Haynes L, Fuller L, McKinney EC. doi: 10.1016/j.fsi.2018.04.016, 113. Blood and inflammatory cells of the lungfish Lepidosiren paradoxa. The second line of defence is through phagocytes; this is again produced by innate immunity. doi: 10.1002/immu.200310022, 146. Fish Shellfish Immunol. Loong AM, Hiong KC, Wong WP, Chew SF, Ip YK. However, due to their unique position in the evolution of adaptive immunity, more functional studies are now being applied to cartilaginous fish. Innate Immunity: Innate immunity refers to a naturally occurring immunity by the genetic constituents and the physiology of a person. doi: 10.1073/pnas.1000072107, 38. Hart S, Wrathmell AB, Harris JE. In lungfish, IgM, IgW, and IgN have been identified, while in coelacanths two forms of IgW has been discovered (Figure 3B) [reviewed in (158)]. Immunobiology of the Shark. (2011) 30:609–17. (2018) 9:1523. doi: 10.3389/fimmu.2018.01523, 78. (2015) 53:112–22. Not all species of cartilaginous fish exhibit all three types of granulocytes; for example, only G1 and G3 granulocytes have been found in Thornback rays (Raya clavate) and small eyed rays (Raja microcellata) (68). doi: 10.1080/01926230701320337. Kobayashi K, Tomonaga S, Kajii T. A second class of immunoglobulin other than IgM present in the serum of a cartilaginous fish, the skate, Raja kenojei: isolation and characterization. Whereas, innate immunity is present in all multicellular organisms, an adaptive immune system, based on VLRs, B-like, and T-like cells is found in jawless vertebrate, while an adaptive immune system, based on an Ig/TCR/MHC system, evolved with the appearance of jawed vertebrates. (2004) 173:7317–23. Magnadottir B. Innate immunity of fish (overview). (1988) 12:561–71. doi: 10.1016/0145-305X(84)90063-6, 46. Krishnan J, Selvarajoo K, Tsuchiya M, Lee G, Choi S. Toll-like receptor signal transduction. (2008) 8:533–44. Neumann NF, Stafford JL, Barreda D, Ainsworth AJ, Belosevic M. Antimicrobial mechanisms of fish phagocytes and their role in host defense. Innate immunity refers to a naturally occurring immunity by the genetic constituents and the physiology of a person. 254. (2009) 146:295–306. (2002) 56:130–48. IgM in the serum of cartilaginous fish is found in two different states, a monomeric 7S and pentameric 19S, which are present in approximately equal amounts (183). Adv Exp Med Biol. An immunochemical and cellular analysis of the B cell response. IV. Front Immunol. The lack of this domain does not interfere with the ability to interact with Igα/Igβ signaling molecules (179). The TCR is found in two forms: a heterodimer of α and β chains (αβ-TCR) or a heterodimer of γ and δ chains (γδ-TCR), linked by disulphide bonds. (2008) 181:3310–22. Mol Immunol. PLoS ONE. doi: 10.1016/j.fsi.2008.09.019, 263. doi: 10.4049/jimmunol.1600164, 203. Dev Comp Immunol. doi: 10.1016/S0145-305X(01)00033-7, 144. Organisms must constantly protect themselves from harm caused by pathogens like viruses and bacteria. (2016) 37:724–37. Finally, sharks use AID and SHM to diversify the shark TCR-γ and -α chains (27, 259). Rumfelt LL, Avila D, Diaz M, Bartl S, McKinney EC, Flajnik MF. (2014) 47:309–18. The development of high-quality, well-assembled, and annotated genomes in fish species will allow the identification of more immune-relevant transcripts, such as NITRs. The secretory pathway has been identified in many different fish species. doi: 10.1016/j.molimm.2006.09.010, 285. Sites of immune cell production in elasmobranch fishes. Saurabh S, Sahoo PK. doi: 10.1016/j.fsi.2017.03.037, 152. doi: 10.1016/0305-0491(85)90448-1, 178. doi: 10.1007/s00239-003-2566-7, 297. doi: 10.1042/bj3260877, 110. Solbakken MH, Torresen OK, Nederbragt AJ, Seppola M, Gregers TF, Jakobsen KS, et al. The mammalian complement system is composed of ~30 proteins that make up three activation pathways: the classical pathway, activated by antibody-antigen complexes and thus a bridge between innate and adaptive immunity; the alternative pathway, which is independent of antibodies and activated directly by pathogens; and the lectin pathway which is activated by the binding of the mannose-binding lectin (MBL), or ficolin, to mannose (or other sugar) residues present on the pathogen surface (102). McKinney EC, Flajnik MF. The J chain, which is required for IgM polymerization and secretion into the mucosa, has not been found in teleost fish, and therefore, tetrameric IgM is polymerized by interchain disulfide bonds (180). NS was supported by an NSERC PGS D fellowship, MR was supported by NSERC Discovery Grant 341304-2012, and SC was supported by NSERC Discovery Grant 2017-04630 and a Memorial University of Newfoundland Seed grant. The development of antibodies that specifically detect fish proteins, such as CD4 and CD8, is necessary to fully understand the homing and recirculation of B and T cells in fish. doi: 10.1016/j.dci.2008.05.008, 180. Antigens that are to be presented by MHC I are processed via the immunoproteasome and transferred to the endoplasmic reticulum (ER) by transporter associated with antigen processing (TAP) where they associate with MHC I and are eventually transported to the cell membrane. Merrifield DL, Harper GM, Dimitroglou A, Ringà E, Davies SJ. (2006) 43:1418–23. Adaptive Immunity: Adaptive immunity develops allergic reactions; immediate and delayed hypersensitivity. Mol Cell Biochem. In teleost fish, the interbranchial septum is reduced and contains a single caudal opening of the operculum, rather than multiple openings while in cartilaginous fish, the gills are supported for almost their entire length by an interbranchial septum with multiple branchial slits or gill openings (23). (2006) 50:185–95. (2010) 11:486. doi: 10.1186/1471-2164-11-486, 169. Adaptive Immunity: Lymph nodes, spleen, and lymphoid tissues are the barriers of the adaptive immunity. Amemiya CT, Alfoldi J, Lee AP, Fan S, Philippe H, Maccallum I, et al. Similarly, components of all three pathways have been characterized in several cartilaginous fish species, where they have been found to have hemolytic properties (113–116). AMPs, also known as host defense peptides, that are generally oligopeptides with a varying number of amino acids that are generally positively charged and play a major role in the innate immune system. Adaptive Immunity: Adaptive immunity possesses a higher diversity. In this regard, there are many factors or agents that … Adaptive Immunity: Adaptive immunity exhibits a higher potency. After engulfment, the phagosome, containing the pathogen, binds to a lysosome, forming a phagolysosome, where the pathogen is killed by various means including the production of ROS and NO (57). Several of these AMPs have been cloned and subsequent functional studies have demonstrated antiviral and antibacterial activities against a variety of pathogens, demonstrating that AMPs from teleost fish exhibit many if not all of the characteristics of other vertebrate AMPs (131–134). Fish Shellfish Immunol. doi: 10.1016/j.fsi.2010.06.004, 198. Ma TY, Wu JY, Gao XK, Wang JY, Zhan XL, Li WS. Putative CD28, CLA-4, and B7 genes have been annotated in the elephant shark genome, while CD28 has been identified in the coelacanth genome, however the function of these co-receptors in many fish species remains to be fully investigated (54, 173). Innate vs Adaptive or Acquired Immunity, 10 minute simplified video 1: Why the name? Dev Comp Immunol. Characterization and expression studies of Gaduscidin-1 and Gaduscidin-2; paralogous antimicrobial peptide-like transcripts from Atlantic cod (Gadus morhua). Mol Biol Evol. (1983) 15:437–55. Pilstrom L, Warr GW, Stromberg S. Why is the antibody response of Atlantic cod so poor? (2013) 33:479–92. (2011) 35:898–905. (1993) 39:447–59. Serum CRP-like protein profile in common carp Cyprinus carpio challenged with Aeromonas hydrophila and Escherichia coli lipopolysaccharide. doi: 10.1016/S0145-305X(98)00002-0, 114. Kuroda N, Figueroa F, O'hUigin C, Klein J. Rectangles represent Ig super-family domains; V represent variable domains (white), C represents constant domains (purple) and NAR V represents extra variable domain in NARTCR (green). Paralogous TLR4 and TLR8 genes have been identified in zebrafish (D. rerio) (81, 93), TLR8 in rainbow trout (76) and TLR3 and TLR7 in common carp (C. carpio) (84), while multiple copies of TLR7, TLR8, TLR9, TLR22, and TLR25 have been identified in the Atlantic cod (87). INTRODUCTION The immune system is comprised of two arms that work together to protect the body – the innate and adaptive immune systems. For example, β-defensin has been characterized in gilthead seabream, where it demonstrated antimicrobial activity against V. anguillarum, while in Nile tilapia (O. niloticus) β-defensin has shown an inhibitory effect on the growth of Escherichia coli DH5α and Streptococcus agalactiae (135). This is known as innate immunity. Dev Comp Immunol. doi: 10.1016/j.molimm.2014.05.009, 127. (2008) 32:1385–97. Lungfish macrophages are described to have typical vertebrate macrophage morphology (59, 60). Fish Shellfish Immunol. Immunogenetics. Interestingly, while MHC I and II are conserved in most jawed vertebrates, Gadiformes, such as the Atlantic cod, have lost the genes for MHC II and CD4, a co-receptor on T cells that interacts with MHC II (87, 88, 220, 240). doi: 10.1016/j.vaccine.2011.04.003, 292. (2007) 23:1127–51. (2016) 64:167–77. Two C3 variants have been described in the nurse shark and the small-spotted catshark, while two C4 gene haven been identified in the elephant shark and hammerhead shark (54, 106, 107). (2005) 174:6608–16. Dis Aquat Organ. (2010) 27:2576–86. doi: 10.4049/jimmunol.173.1.349, 175. CRP and SAP are considered major APPs (e.g., their concentrations may increase up to 1,000-fold) and are the most extensively studied APPs in fish. Criscitiello MF. For example, CRP and SAP expression in Atlantic salmon head kidney leukocytes are upregulated in response to r-IL-Iβ and r-IFNγ, two cytokines that stimulate APP production in mammals, suggesting that the acute phase response is evolutionarily conserved (151). Bernard D, Riteau B, Hansen JD, Phillips RB, Michel F, Boudinot P, et al. Panopoulou G, Poustka AJ. 42. doi: 10.1016/j.dci.2008.04.004. Groot Kormelink T, Mol S, de Jong EC, Wauben MHM. New York, NY: Garland Science (2002). Transcription of signal-3 cytokines, IL-12 and IFNαβ, coincides with the timing of CD8αβ up-regulation during viral infection of common carp (Cyprinus carpio L). Chen H, Kshirsagar S, Jensen I, Lau K, Simonson C, Schluter SF. In addition, no AID was found in the African lungfish using RT-qPCR (228). Dev Comp Immunol. Clem LW. Sahoo M, Edholm ES, Stafford JL, Bengten E, Miller NW, Wilson M. B cell receptor accessory molecules in the channel catfish, Ictalurus punctatus. Dev Comp Immunol. (2005) 26:581–7. (2014) 511:E7–9. In teleost fish, IgM is multimerized into a tetrameric form, although there have been reports of a monomeric IgM form in some teleost species (176, 177). Ray-finned fish comprise 99% of the Osteichthyes, of which 96% are from the infraclass Teleostei (9, 12). Identification and characterization of a novel intraepithelial lymphoid tissue in the gills of Atlantic salmon. doi: 10.1093/gbe/evw250, 241. Overgard AC, Hordvik I, Nerland AH, Eikeland G, Patel S. Cloning and expression analysis of Atlantic halibut (Hippoglossus hippoglossus) CD3 genes. The existence of M1 and M2 cell populations, as well as EVs, have yet to be examined in cartilaginous and lobe-finned fish. J Biol Chem. (1997) 326(Pt 3):877–81. doi: 10.1046/j.1365-3083.2002.01116.x, 214. Lymphoid aggregates, as well as macrophages and granular cells, have been found in the spiral valve of various shark and ray species (42, 43). IgD is found in many vertebrate classes, including teleost fish and acipenseriformes (a group of fish that phylogenetically links elasmobranches, teleosts, and sturgeons). Nonaka M, Nonaka M, Irie M, Tanabe K, Kaidoh T, Natsuume-Sakai S, et al. Nikolich-Zugich J, Slifka MK, Messaoudi I. doi: 10.1016/j.fsi.2009.04.003, 167. Fish granulocytes exhibit a wide variation in morphology, numbers and types of cells between species causing much confusion regarding their nomenclature. (1984) 8:803–12. For example, double positive T cells were observed in the thymic cortex of sea bass, while single CD4+ or CD8 α+ cells were found in the thymic medulla, similar to that of mammals (166). (2011) 6:e23417. Wx, Chan SM Hoyheim B, Helland R, Kaplan G, et al use and! Not taken https: //www.ncbi.nlm.nih.gov/books/NBK26827/, 261 about these two varieties is shared here 66 ) identified only type. Phagocytes serve as internal barriers, it is also involved in immune responses Verma AD, Xia B. CD4 +. 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