Dev Comp Immunol. doi: 10.1007/s00251-002-0473-5, 238. The former include T lymphocytes and their subsets identified by CD markers, B lymphocytes and natural killer (NK) cells. (2006) 50:185–95. doi: 10.1016/j.virol.2006.01.019, 267. 1. Isolation and characterization of two major serum proteins from the dogfish, Mustelus canis, C-reactive protein and amyloid P component. J Mol Evol. (2010) 29:594–9. In brief, two forms of IFNγ, produced by Th1 cells, IFNγ, and IFNγ rel, have been identified in teleost fish including Atlantic salmon, rainbow trout, and ginbuna crucian carp (Carassius carassius), while one form has been identified in fugu (278–280). Adaptive Immunity: Adaptive immunity refers to an acquired immunity, which is mediated by T cells and B cells and is characterized by an immunological memory. Due to the large number of teleost species, as well their economic importance, there have been many genomic and functional immunological studies completed on teleost fish. doi: 10.1016/0145-305X(88)90072-9. 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. Chang CI, Zhang YA, Zou J, Nie P, Secombes CJ. Shark immunity bites back: affinity maturation and memory response in the nurse shark, Ginglymostoma cirratum. doi: 10.3390/biology4030525, 25. A comparative study of pentraxin-like proteins in different fish species. Srisapoome P, Ohira T, Hirono I, Aoki T. Genes of the constant regions of functional immunoglobulin heavy chain of Japanese flounder, Paralichthys olivaceus. Castro CD, Flajnik MF. Two homologs of the IL-17 family, IL-17B and IL-17D have been identified in teleost fish, as well as several isoforms of molecules termed IL-17A/F1-3, IL-17C, and IL-17E (290). Liu Z, Liu S, Yao J, Bao L, Zhang J, Li Y, et al. This structure is conserved in almost all vertebrates (255). They diverged from a jawless common ancestor with the lineage leading to other bony vertebrates. Innate Immunity: Innate immunity refers to a naturally occurring immunity by the genetic constituents and the physiology of a person. (2011) 35:1309–16. doi: 10.1016/0165-2427(86)90115-7, 45. 254. Fish Shellfish Immunol. The immune response is non-specific in innate immunity, and it is specific in adaptive immunity. These differences indicate that not only is pathogen recognition quite diverse in fish, it can also be lineage-specific. (2014) 322:415–37. (1996) 240:143–9. doi: 10.4049/jimmunol.172.10.5851, 215. The spleen of the African lungfish Protopterus annectens: freshwater and aestivation. Evolutionary variation of immunoglobulin mu heavy chain RNA processing pathways: origins, effects, and implications. doi: 10.1016/j.dci.2008.05.008, 180. The T cell receptor (TCR) is always membrane bound and once stimulated via interaction with antigen presented by the pAPC, in the presence of co-stimulation, the T cell can be activated to function as a helper (CD4+) T cell, a regulatory (CD4+) T cell or a cytotoxic (CD8+) T cell. (2013) 496:311–6. Mol Immunol. doi: 10.1002/bies.201200005, 221. J Immunol. doi: 10.1016/j.molimm.2014.05.009, 127. In addition, a novel TLR with sequence similarity to TLR4 and TLR13 in mammals, and TLR21 in teleost fish, has been identified in the whale shark (Rhincodon typus) (80, 83). Similar to mammals, functionally distinct subpopulations of macrophages exist in bony fish. 10:2292. doi: 10.3389/fimmu.2019.02292. The immune system consists of a collection of molecules, cells, and tissues, which protect the body from various pathogens and toxins. Sugamata R, Suetake H, Kikuchi K, Suzuki Y. Teleost B7 expressed on monocytes regulates T cell responses. Fish Shellfish Immunol. Cell-mediated immunity is mediated by cytotoxic T cells, which induce the cell death of infected cells. Adaptive Immunity: Adaptive immunity is known as acquired immunity. The Atlantic salmon genome provides insights into rediploidization. TLR2, TLR3, TLR6, and TLR9 have been identified in the gray bamboo shark (Chiloscyllium griseum) genome whereas no TLR6 or TLR10 homolog has been identified in teleost fish. Available Online at: https://www.ncbi.nlm.nih.gov/books/NBK26827/, 261. Meyer A, Van de Peer Y. Sahoo M, Edholm ES, Stafford JL, Bengten E, Miller NW, Wilson M. B cell receptor accessory molecules in the channel catfish, Ictalurus punctatus. Distributed as sentinels throughout the body, DC are poised to capture Ag, migrate to draining … Definition. Recombinant IFNγ (r-IFNγ) was found to increase the expression of anti-viral and inflammation-relevant genes, as well as increase ROS and NO production in zebrafish, rainbow trout and goldfish macrophages, indicating a similar function as mammalian IFNγ (281, 282). doi: 10.3390/biology4040881, 56. Quiniou SM, Sahoo M, Edholm ES, Bengten E, Wilson M. Channel catfish CD8α and CD8β co-receptors: characterization, expression and polymorphism. These include macrophages, monocytes, neutrophils and dendritic cells and are found in both bony and cartilaginous fish (57, 95–98). doi: 10.1016/j.dci.2016.03.009. (162), Roux et al. (2003) 55:57–61. Two sub-isotypes of IgM have been identified in Atlantic salmon and brown trout (Salmo trutta), reflecting the pseudotetraploid state of salmonid genomes (181, 182). (2009) 27:768–72. Proc Natl Acad Sci USA. (275, 276) and Secombes and Wang (277). 4: The antimicrobial system of the innate immunity is shown by complement, leukins, plakins, lactic acid, lactoperoxidase and interferon Near TJ, Eytan RI, Dornburg A, Kuhn KL, Moore JA, Davis MP, et al. doi: 10.1016/j.fsi.2009.04.003, 167. Rumfelt LL, McKinney EC, Taylor E, Flajnik MF. Expression, purification, crystallization and preliminary X-ray diffraction analysis of nurse shark β2-microglobulin. doi: 10.1016/j.molimm.2003.10.001, 82. doi: 10.1016/j.fsi.2005.04.004, 103. Cambridge, MA: Academic Press (2016). Elements of the adaptive response also support the function of the innate immune system. doi: 10.1016/j.vetimm.2013.10.012, 235. (2014) 46:448–60. The activation of the adaptive immunity occurs in response to an overcoming of the second line barriers by a particular pathogen. Kawai T, Akira S. The roles of TLRs, RLRs and NLRs in pathogen recognition. Naturally concurrent infections of bacterial and viral pathogens in disease outbreaks in cultured Nile tilapia (Oreochromis niloticus) farms. Sato A, Sultmann H, Mayer WE, Klein J. Mhc class I gene of African lungfish. (2016) 5:e23. The African coelacanth genome provides insights into tetrapod evolution. 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). Alt FW, Rathbun G, Oltz E, Taccioli G, Shinkai Y. Immunobiology of the Shark. (2008) 180:7461–70. The secretion of EVs was not induced by CpG in a splenocyte culture (containing mostly B cells) suggesting that the EVs were likely produced by macrophages or dendritic cells in the head kidney leukocyte culture (64). Fish Shellfish Immunol. Multiple splice forms of IgW exist in cartilaginous fish other than the original six C domains (IgW-long), including a two C domain (IgW-short) form and a four C domain form (188, 200, 201). Edholm ES, Bengten E, Wilson M. Insights into the function of IgD. Dev Comp Immunol. doi: 10.1007/s004290050113, 34. However, in order to understand the functioning of each, some basic knowledge about these two varieties is shared here. Fish Shellfish Immunol. It can take up to 28 months before the antigen-specific titer levels return to pre-immunization levels once the Ig response has reached a plateau (183). (2013) 40:11–9. A recent study by Heimroth et al. doi: 10.1016/S0145-305X(02)00041-1, 301. (2004) 40:759–71. They are subdivided into two classes, the Actinopterygii (ray-finned fish) and the Sarcopterygii (lobe-finned fish) (Figure 1). Dev Comp Immunol. J Exp Zool. Evolution of duplicated IgH loci in Atlantic salmon, Salmo salar. Fish Shellfish Immunol. doi: 10.1002/jlb.61.2.141, 59. McKinney EC, Flajnik MF. doi: 10.1093/molbev/msq147, 8. It is proposed that mature B cells are released into the blood where they encounter antigen and mature into plasma blasts or plasma cells. Solbakken MH, Rise ML, Jakobsen KS, Jentoft S. Successive losses of central immune genes characterize the gadiformes' alternate immunity. *IgM for lungfish only; no IgM in the coelacanth. Adaptive Immunity: Lymph nodes, spleen, and lymphoid tissues are the barriers of the adaptive immunity. Serum levels of IgNAR are much lower than IgM and it is unknown if the J chain is required for IgNAR multimer formation (8). Dev Comp Immunol. J Biol Chem. doi: 10.1016/S0145-305X(82)80011-6, 96. Biology. If a pathogen is ingested, it will encounter the GI tract, which, like the skin and gills, contains both innate and adaptive immune cellular components. Identification of a transcript encoding a soluble form of toll-like receptor 5 (TLR5) in Atlantic salmon during Aeromonas salmonicida infection. doi: 10.1016/j.molimm.2007.10.010, 266. (2009) 27:397–406. Dooley H, Flajnik MF. (2006) 43:1080–7. doi: 10.4049/jimmunol.167.3.1325, 298. 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. (2002) 169:2488–97. Most of the mammalian complement components have homologs in various teleost species, including rainbow trout (105), zebrafish (110), and channel catfish (111), among many others, and their functions have been well-characterized [reviewed in (56, 112)]. Identification and characterization of a variant of the third component of complement (C3) in rainbow trout (Salmo gairdneri) serum. Hohman VS, Stewart SE, Rumfelt LL, Greenberg AS, Avila DW, Flajnik MF, et al. Immunogenetics. doi: 10.1007/s00251-014-0769-2, 161. (2015) 518:209–23. A cold-blooded view of adaptive immunity. Active and passive immunity are two types of adaptive immunity. Ramirez-Gomez F, Greene W, Rego K, Hansen JD, Costa G, Kataria P, et al. New York, NY: Garland Science (2001). doi: 10.1016/j.molimm.2006.09.010, 285. CpG-induced secretion of MHCIIbeta and exosomes from salmon (Salmo salar) APCs. Clin Lab Med. Adaptive Immunity: Adaptive immunity possesses a higher diversity. Dis Aquat Organ. Cho SH, Lee BD, An H, Eun JB, Kenojeinin I. Antimicrobial peptide isolated from the skin of the fermented skate, Raja kenojei. doi: 10.4049/jimmunol.173.12.7317, 211. Identification and characterization of a novel intraepithelial lymphoid tissue in the gills of Atlantic salmon. All TCR have a short cytoplasmic tail and therefore need to partner with CD3, a complex of transmembrane proteins with intracellular domains containing the conserved motif known as immunoreceptor tyrosine-based activation motif (ITAM). Fillatreau S, Six A, Magadan S, Castro R, Sunyer JO, Boudinot P. The astonishing diversity of Ig classes and B cell repertoires in teleost fish. Both CD28 and CLTA4, as well as orthologs of B7.1 and B7.2, have been identified in several teleost species (271–273). Criscitiello MF. doi: 10.1002/1521-4141(200003)30:3<858::AID-IMMU858>3.0.CO;2-M, 120. (2005) 57:458–63. doi: 10.1038/ni1166, 168. Multiple C3 forms have been identified in teleost fish and cartilaginous fish. doi: 10.1002/9781118993705, 69. (2004) 172:3635–43. Knowing the site of hematopoiesis in lobe-finned fish would allow for isolation of these cells and experiments that would lead to a better understanding of immune cells in these species. doi: 10.1111/j.1469-7580.2008.00943.x, 26. A major function of B cells, as well as other pAPCs such as macrophages and dendritic cells, is to process and present antigen to activate T cells. The vertebrate immune system is divided into 2 subsystems—the innate immune system and the adaptive immune system. (2016) 61:13–24. Innate immunity is orchestrated through phagocytes (Macrophages, and Neutrophils) and Natural Killer cells. The channel catfish genome sequence provides insights into the evolution of scale formation in teleosts. 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). J Biochem. Components of the innate immune system include physical barriers such as the skin, cellular processes such as phagocytosis and humoral components such as soluble proteins (1). Molecular cloning and expression analysis of major histocompatibility complex class IIB gene of the Whitespotted bambooshark (Chiloscyllium plagiosum). Nakao M, Mutsuro J, Obo R, Fujiki K, Nonaka M, Yano T. Molecular cloning and protein analysis of divergent forms of the complement component C3 from a bony fish, the common carp (Cyprinus carpio): presence of variants lacking the catalytic histidine. (2011) 48:1432–8. The key difference between these two segments is that, innate immunity is present from the point of birth while acquired immunity … Evidence for B cell development in the anterior kidney is supported by expression of RAG-1/2 and terminal deoxynucleotidyl transferase (TdT), and the resulting development of immature B cells with membrane Ig on their surface. doi: 10.1016/j.ydbio.2006.05.006, 39. Cell Res. doi: 10.1016/j.fsi.2009.05.020, 116. Characterization studies of CD3 in teleost fish have identified a conserved structure of CD3 between teleost fish and mammals (268, 269). ISRN Immunol. The immune response is broken down into innate immunity, which an organism is born with, and adaptive immunity, which an organism … While all three RIG-I-like receptors have been characterized in teleost fish, only RIG-I and MDA5 have been identified in the elephant shark and coelacanth genomes (54, 78). Ward JW. (2013) 4:28. doi: 10.3389/fimmu.2013.00028, 199. (2007) 44:2078–86. Tissue Cell. (1999) 49:367–75. Kurobe T, Hirono I, Kondo H, Saito-Taki T, Aoki T. Molecular cloning, characterization, expression and functional analysis of Japanese flounder Paralichthys olivaceus Fas ligand. Harun NO, Wang T, Secombes CJ. doi: 10.1111/j.1574-6968.1989.tb03441.x, 122. Origin of immunoglobulin isotype switching. Several other studies in fish, including rainbow trout and turbot (Psetta maxima), have since shown that neutralizing Ab can be induced against a variety of viral, bacterial and parasitic pathogens and vaccines (217, 218). Striking antigen recognition diversity in the Atlantic salmon T-cell receptor alpha/delta locus. 255. Identification of sturgeon IgD bridges the evolutionary gap between elasmobranchs and teleosts. (2017) 63:384–93. (2015) 4:525–39. Cheesman SE, Neal JT, Mittge E, Seredick BM, Guillemin K. Epithelial cell proliferation in the developing zebrafish intestine is regulated by the Wnt pathway and microbial signaling via Myd88. doi: 10.1016/j.fsi.2010.08.016, 256. What is the difference between innate and adaptive immunity? (2005) 6:295–302. Fish Shellfish Immunol. Zhang J, Kong X, Zhou C, Li L, Nie G, Li X. Toll-like receptor recognition of bacteria in fish: ligand specificity and signal pathways. For example, the West African lungfish has three IgM isotypes, while the spotted lungfish (P. dolloi) has two. Dong HT, Nguyen VV, Le HD, Sangsuriya P, Jitrakorn S, Saksmerprome V, et al. Boshra H, Li J, Sunyer JO. doi: 10.1002/jmor.1051640102, 68. Adaptive Immunity: Adaptive immunity only occurs in response to exposure to an external factor. Aust J Zool. An equally important function of the skin is the ability to secrete mucus, which acts as both a physical barrier, by trapping pathogens, and a chemical barrier (16). Lysozyme is one of the most studied innate immune components in fish. The secretory pathway releases granular toxins such as perforin and serine proteases called granzymes which work together to induce apoptosis (293). Dev Comp Immunol. Bridle A, Nosworthy E, Polinski M, Nowak B. Immunogenetics. doi: 10.1007/s00251-003-0542-4, 242. Comparative studies in fish such as those discussed here are essential for developing a comprehensive understanding of the evolution of the immune system. In: Immunobiology: The Immune System in Health and Disease. doi: 10.1016/j.dci.2016.08.016, 293. (2011) 150:635–48. Buonocore F, Randelli E, Trisolino P, Facchiano A, de Pascale D, Scapigliati G. Molecular characterization, gene structure and antibacterial activity of a g-type lysozyme from the European sea bass (Dicentrarchus labrax L.). Pettinello R, Dooley H. The immunoglobulins of cold-blooded vertebrates. It is unknown how the GALT in these species respond to infection and if it is in a similar manner as teleost fish and mammals. Dev Comp Immunol. Star B, Jentoft S. Why does the immune system of Atlantic cod lack MHC II? (2003) 23:441–71. Inoue JG, Miya M, Lam K, Tay BH, Danks JA, Bell J, et al. (2012) 36:665–79. doi: 10.1016/j.dci.2018.05.009, 86. Oncotarget. doi: 10.1371/journal.pone.0023417, 138. Similar to teleost fish, the immune response time of IgM in cartilaginous fish is much longer than in mammals. Innate type of immunity is generally inherited from parents and passed to … Innate Immunity: Temperature, pH, skin, and mucous membranes are the barriers of the innate immunity. J Immunol. (2012) 68(Pt 4):460–3. IgM is the only teleost isotype for which sub-isotypes have been identified. Immunobiology of the Shark. doi: 10.1038/nri2356, 51. In addition to inflammation, CRP and SAP have been shown to activate the complement pathways and play a role in the clearance of apoptotic cells (143, 145). doi: 10.1016/j.dci.2014.02.018, 237. Proc Natl Acad Sci USA. Dev Comp Immunol. S.N. Salmonids have an extraordinary complex type I IFN system: characterization of the IFN locus in rainbow trout Oncorhynchus mykiss reveals two novel IFN subgroups. Roca FJ, Sepulcre MA, Lopez-Castejon G, Meseguer J, Mulero V. The colony-stimulating factor-1 receptor is a specific marker of macrophages from the bony fish gilthead seabream. Chondrou MP, Mastellos D, Zarkadis IK. IgT, a primitive immunoglobulin class specialized in mucosal immunity. Figure 2 illustrates these three pathways, along with some of the associated proteins. J Comp Physiol B. IgM is the most prevalent Ab in both bony and cartilaginous fish plasma and can be found in both secreted and transmembrane forms. Humoral immunity and cell mediated immunity are two types of adaptive immunity. However, the response time of teleost IgM is much slower than in mammals, taking 3–4 weeks after immunization before specific titers are detected. While VDJ recombination has been characterized in fish [reviewed in (158, 165)], the process of negative and positive selection of developing B and T cells has not been fully elucidated, although these processes possibly occur in a similar manner as mammals. Fish Shellfish Immunol. doi: 10.2174/0929866524666161121142221, 142. 49. Innate Immunity: Innate immunity does not produce allergic reactions. doi: 10.1371/journal.pone.0001463, 111. doi: 10.1038/nature17164, 13. Negative selection occurs when a double positive T cells binds to MHC I or II with a high enough affinity to receive an apoptotic signal. (2005) 35:936–45. Ann N Y Acad Sci. Xiao FS, Wang YP, Yan W, Chang MX, Yao WJ, Xu QQ, et al. Parra D, Korytar T, Takizawa F, Sunyer JO. “2211 Cooperation Between Innate and Immune Responses” By OpenStax College – Anatomy & Physiology, Connexions Web site., Jun 19, 2013. In order to gain a better understanding of the lobe-finned fish immune system, the site(s) of hematopoiesis must be determined. doi: 10.1016/j.fsi.2006.10.005, 61. Fänge R. Lymphomyeloid tissues and blood cell morphology in elasmobranchs. As well, transcriptome analysis of the Indonesian coelacanth, Latimeria menadoensis, genome identified SAP encoding transcripts (157), however, to our knowledge, no other studies examining CRP or SAP in coelacanths or lungfish have been reported. The IgH locus of the channel catfish, Ictalurus punctatus, contains multiple constant region gene sequences: different genes encode heavy chains of membrane and secreted IgD. There are over 1,000 species of cartilaginous fish, which are divided into two subclasses: Elasmobranchii (sharks, rays, skates, and sawfish) and Holocephali (chimeras) (7). Mol Immunol. doi: 10.1016/j.molimm.2009.10.010, 286. Zwollo P, Cole S, Bromage E, Kaattari S. B cell heterogeneity in the teleost kidney: evidence for a maturation gradient from anterior to posterior kidney. Dev Comp Immunol. Innate Immunity: Innate immunity possesses a less diversity. Jenkins JA, Ourth DD. The three complement pathways with associated proteins. However, AID from teleost fish, specifically zebrafish, Japanese puffer and catfish can catalyze CSR in vitro in mammalian AID−/− lymphocytes suggesting that teleost AID has the full catalytic functions capable for CSR (225, 232). doi: 10.1038/nri3622, 63. The corporation between first line, second line, and third line defenses are shown in figure 2. doi: 10.1016/j.dci.2015.06.009, 149. Heimroth RD, Casadei E, Salinas I. They are members of the pentraxin family of APPs, are present in the body fluids of vertebrates and invertebrates, and are commonly associated with the acute phase response of inflammation (143). The immune system is composed of two subsystems—the innate immune system and the adaptive immune system. Serum CRP-like protein profile in common carp Cyprinus carpio challenged with Aeromonas hydrophila and Escherichia coli lipopolysaccharide. Immunogenetics. Implications of the new cells involved. The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. What are the Similarities Between Innate and Adaptive Immunity      – Outline of Common Features 4. BMC Genomics. Saliva, tears, skin, mucus linings, stomach acid, and some of the bacteria in the gut serve as barriers, preventing the entering of pathogens to the tissues of the body. Yazawa R, Cooper GA, Hunt P, Beetz-Sargent M, Robb A, Conrad M, et al. Both bony and cartilaginous fish have been shown to develop immunological memory (i.e., the ability to respond more rapidly and effectively to a pathogen that has been previously encountered). The adaptive immune system is highly specific to a particular antigen and can provide long-lasting immunity (2). PLoS ONE. (2013) 35:1824–8. (2002) 54:418–30. Structural analysis of the nurse shark (new) antigen receptor (NAR): molecular convergence of NAR and unusual mammalian immunoglobulins. Scand J Immunol. Grant KR. Ultimately, these pathways induce activation of the C3 convertase, which cleaves inactive C3 into C3a, an anaphylatoxin that acts as a chemotactic factor and aids in inflammation, and C3b, which acts as an opsonin, as well as an activator of downstream complement proteins leading to the formation of the membrane attack complex (103, 104). Chen H, Kshirsagar S, Jensen I, Lau K, Covarrubias R, Schluter SF, et al. J Immunol. doi: 10.1038/ni1389, 101. Key Difference – Innate Immunity vs Acquired Immunity Innate immunity and acquired immunity are two important and different segments of the immune system that act together to defend the body against infection and disease. The BCR includes the membrane-bound antibody (Ab) and the Ig-α/Ig-β (CD79a/b) heterodimer, which is involved in signal transduction. TLR27 was first identified and thought to only be found in the coelacanth genome but has since been identified in the spotted gar (Lepisosteus oculatus) and elephant shark (78, 89). doi: 10.1038/nature14438, 5. In both fish and mammals, tissue injury, infection and inflammation induce immune cells, such as macrophages, to secrete various cytokines into the bloodstream, which stimulate hepatocytes to produce and release acute phase proteins (APPs) (141, 142). Finally, sharks use AID and SHM to diversify the shark TCR-γ and -α chains (27, 259). 1. Flajnik MF. (2008) 32:204–12. doi: 10.1006/fsim.2000.0320, 32. A V-less form of IgW has been identified in both the spiny dogfish (S. acanthias) and the nurse shark but represents only 8% of the IgW transcripts analyzed (200). Innate vs. Adaptive Immunity The immune system is classically subdivided into two major categories: the adaptive and innate – even though the distinctions are in no way mutually exclusive. Dark blue circles represent heavy chain domains, light blue circles represent light chain domains (158–162). (2004) 16:589–97. The scales of cartilaginous fish are called placoid scales, also known as denticles (16). Palti Y. Toll-like receptors in bony fish: from genomics to function. Ig heavy chain genes and their locus in grass carp Ctenopharyngodon idella. doi: 10.1016/j.dci.2009.11.011, 289. Dev Comp Immunol. Be design, the innate immune … (1987) 67:639–45. Flajnik MF, Kasahara M. Origin and evolution of the adaptive immune system: genetic events and selective pressures. G1 granulocytes, referred to as heterophils or fine eosinophilic granulocytes, are often the most common granulocyte in cartilaginous fish. In: Smith S, Sim R, Flajnik M, editors, Immunobiology of the Shark. doi: 10.1111/j.1365-2109.2007.01883.x, 121. (2006) 177:61–9. Differentiate between innate immunity and acquired immunity. Sepahi A, Cordero H, Goldfine H, Esteban MA, Salinas I. Symbiont-derived sphingolipids modulate mucosal homeostasis and B cells in teleost fish. Riera Romo M, Perez-Martinez D, Castillo Ferrer C. Innate immunity in vertebrates: an overview. Nunez Ortiz N, Gerdol M, Stocchi V, Marozzi C, Randelli E, Bernini C, et al. Observations on the blood of the Australian lungfish, Neoceratodus-Forsteri Klefft. (2001) 25:807–25. (2011) 35:1336–45. Depending on the species, differences may occur such as duplication of individual V, D, or J segments, or tandem duplication of C domain exons such as that found in Atlantic salmon and zebrafish (167, 168). M1 (classically activated macrophages) are characterized by production of pro-inflammatory cytokines such as TNFα and IL-1β and production of ROS and NO, whereas M2 (alternatively activated macrophages) are linked to immunosuppression, wound repair and increased levels of arginase and anti-inflammatory cytokines such as interleukin (IL)-10 (55, 57, 61). (2008) 24:412–25. Janeway CA Jr, Travers P, Walport M, Shlomchik MJ. (2007) 35:495–516. p. 3–68. In the adaptive immune system, macrophages are one type of professional antigen presenting cell (pAPC) that can present phagocytosed materials to the T lymphocytes of the adaptive immune system through a process termed antigen presentation. Dev Comp Immunol. The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. doi: 10.1073/pnas.1304319110, 18. Innate immunity generates first line and second line immune responses. 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. When an innate immune cell encounters a pathogen, it will recognize a pathogen-associated molecular pattern (PAMP) found on the pathogen. Characterization of the immunoglobulin repertoire of the spiny dogfish (Squalus acanthias). Science. doi: 10.1002/jez.b.22558, 174. Characterization of γδ T cells from zebrafish provides insights into their important role in adaptive humoral immunity. Meijer AH, Gabby Krens SF, Medina Rodriguez IA, He S, Bitter W, Ewa Snaar-Jagalska B, et al. (2008) 67:160–8. Mucus from teleost fish contains a combination of lectins, lysozymes, complement proteins, and antimicrobial peptides (AMPs); all of which play a critical role in neutralizing pathogens (16, 21). Contrast host innate resistance with adaptive immunity 2. Many homologs of mammalian PRRs have been identified in fish. Upon activation of CD4+ cells, naïve cells can differentiate into specific subsets including Th1, Th2, Th17, and inducible T-regulatory (Treg) cells; each subset defined by their cytokine production (274). Betz UA, Mayer WE, Klein J. Dodds AW, Smith SL, Levine RP, Willis AC. doi: 10.1016/0161-5890(84)90037-3, 189. PLoS ONE. The secretory pathway has been identified in many different fish species. (2004) 56:292–300. Genome Biol Evol. (2011) 30:894–901. (2011) 6:e18832. 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. Aybar L, Shin DH, Smith SL. doi: 10.1016/S0145-305X(01)00033-7, 144. Adaptive Immunity: Adaptive immunity can last for the lifetime or for a short period of time. J Immunol. (2006) 109:183–7. A comprehensive understanding of the evolution of the immune system will continue to develop as more comparative research on cartilaginous fish, lobed-finned fish, and ray-fined fish is completed. A foreign substance the anterior ( or head ) kidney Wang YP, Yan W, Wang JY Zhan. Jmm, Prado EJR, Moraes AC, Eto SF, Medina Rodriguez IA He. 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