Humoral factors of innate immunity in fish
DOI:
https://doi.org/10.33910/2687-1270-2025-6-3-279-294Keywords:
fish immunity, humoral factors, aquaculture, innate immunity, complement system, lysins, lectins, pentraxins, interferons, chemokinesAbstract
The study of humoral factors of innate immunity in bony fish (Teleostei) is of critical importance for advancing national aquaculture. Although fish possess both innate and adaptive immune systems, as do more phylogenetically advanced vertebrates, data from model organisms of other taxonomic classes cannot be directly extrapolated. Poikilothermy in combination with an aquatic environment fundamentally shapes the development and function of all aspects of the teleost immune system. Aquaculture enterprises are invested not only in enhancing productivity but also in sustainable, prophylactic strategies to mitigate the risk of pathological outbreaks. This article details the principal components of nonspecific humoral defense in fish: lysins (including antimicrobial peptides, the complement system, and lysozyme), lectins (such as mannose-binding lectins, calcium-dependent lectins, and pentraxins), transferrin, metallothioneins, interferons, and chemokines. We outline their roles in pathogen defense and the regulation of inflammatory responses. The nonspecific arm constitutes the foundational defense mechanism in teleosts. A comprehensive understanding of these humoral factors is essential for identifying key targets for effective disease prevention in aquaculture.
References
ЛИТЕРАТУРА
Алексеева, И. Г., Артамонов, Н. А. (2023) Сапролегниоз рыб в промышленном рыбоводстве. В кн.: Ветеринарная медицина: связь поколений как фактор устойчивого развития России. Материалы Международной конференции. Омск: Изд-во Омского государственного аграрного университет имени П. А. Столыпина, с. 153–158.
Апаликова, О. В., Киселева, М. Н., Митрюшкина, Д. К. и др. (2025) Молекулярно-генетические подходы к изучению устойчивости рыб к болезням. Труды ВНИРО, т. 199, с. 142–156. https://doi.org/10.36038/2307-3497-2025-199-142-156
Ахмедова, С. Д. (2014) Показатели гуморального иммунитета у пациентов с микотическим поражением кожи и ее придатков. Медицинские новости, № 3, с. 77–79.
Вавиленкова, Ю. А. (2012) Современные представления о системе интерферона. Вестник Смоленской государственной медицинской академии, т. 11, № 2, с. 74–82.
Вылка, М. М., Лютиков, А. А., Дьякова, С. А. и др. (2025) Микробиота кишечника производителей сиговых на примере Coregonus nasus в аквакультуре в нерестовый период. Экологическая генетика, т. 23, № 3, с. 225–234. https://doi.org/10.17816/ecogen679052
Делягин, В. М., Скворцова, Ю. В., Раймулла, Е. А. (2025) Остеоиммунология. Лечение и профилактика, т. 15, № 2, с. 5–19.
Дрошнев, А. Е., Завьялова, Е. А., Гулюкин, М. И., Хлунов, О. В. (2012) Современная вакцинопрофилактика радужной форели против вибриоза. Российский ветеринарный журнал. Сельскохозяйственные животные, № 1, с. 31–33.
Дун, С., Шилин, М. Б., Леонтьева, Е. О. (2024) Развитие новых технологий защиты рыб от инфекций в условиях аквакультуры. Арктика и инновации, т. 2, № 3, с. 64–82. https://doi.org/10.21443/3034-1434-2024-2-3-64-82
Егорова, Е. В., Кренев, И. А., Оборин, Н. Н., Берлов, М. Н. (2023) Антимикробная активность системы комплемента. Медицинский академический журнал, т. 23, № 2, с. 31–45. https://doi.org/10.17816/MAJ322841
Жандалгарова, А. Д., Микряков, Д. В., Грозеску, Ю. Н. и др. (2025) Иммуномодулирующее действие комплекса бифидо- и лактобактерий на неспецифический гуморальный иммунитет гибрида осетровых рыб. Известия Российской академии наук. Серия биологическая, № 2, с. 249–254.
Завьялова, Е. А., Дрошнев, А. Е., Булина, К. Ю. (2019) Вакцинация в аквакультуре: история, закономерности и особенности проведения работ. Российский журнал «Проблемы ветеринарной санитарии, гигиены и экологии», № 1 (29), с. 95–100.
Зубарева, А. А., Лютиков, А. А., Скорик, Ю. А. (2024) Создание прототипов ДНК-вакцин для аквакультуры на основе природных полисахаридов. В кн.: К. В. Колончин, М. М. Мельник (ред.). Рыбохозяйственная наука. История, современность, перспективы. Материалы Международной научно-практической конференции, посвященной 110-летию создания «ГосНИОРХ» им. Л. С. Берга. М.: Всероссийский научно- исследовательский институт рыбного хозяйства и океанографии, с. 173–175.
Кочнева, А. А., Курицын, А. Е., Мурзина, С. А. (2025) Выявление in silico антимикробных последовательностей белков мозга радужной форели в аспекте комплексной переработки отходов аквакультуры. Доклады Российской академии наук. Науки о жизни, т. 520, с. 50–56. https://doi.org/10.7868/S3034505725010099
Кузьмина, В. В. (2016) Влияние цинка и меди на активность протеаз пищеварительного тракта, обеспечивающих неспецифическую защиту рыб. Труды ВНИРО, т. 162, с. 64–72.
Меркулова, Н. Л., Грехнева, Е. В., Чуйкова, С. В. (2021) Изучение возможности использования эпидермальной слизи Clarias gariepinus как нового источника фармакологически ориентированных препаратов. В кн.: А. Г. Габибов, М. А. Островский (ред.). III объединенный научный форум физиологов, биохимиков и молекулярных биологов. VII съезд биохимиков России. X российский симпозиум «Белки и пептиды». VII съезд физиологов СНГ. Т. 2. М.: Перо, с. 138.
Микряков, В. Р., Микряков, Д. В. (2015) Иммунологическая индикация здоровья рыб. Вопросы ихтиологии, т. 55, № 1, с. 119–123. https://doi.org/10.7868/S0042875215010129
Пронина, Г. И. (2014) Возможность повышения иммунной устойчивости гидробионтов в аквакультуре. Известия Оренбургского государственного аграрного университета, № 3 (47), с. 180–182.
Пронина, Г. И., Иванов, А. А., Маннапов, А. Г., Саная, О. В. (2021) Иммунитет пойкилотермных гидробионтов. Известия Тимирязевской сельскохозяйственной академии, вып. 2, с. 71–91. https://doi.org/10.26897/0021-342X-2021-2-71-91
Субботкин, М. Ф., Субботкина, Т. А. (2020) Влияние заражения и инъекций субстанций различной природы на лизоцим карповых рыб (Cyprinidae) (обзор). Биология внутренних вод, № 2, с. 180–191. https://doi.org/10.31857/S0320965220020151
Субботкина, Т. А., Субботкин, М. Ф. (2003) Особенности активности лизоцима некоторых видов костистых рыб р. Волги. Биология внутренних вод, № 1, с. 102–107.
Суворова, Т. А., Микряков, Д. В., Пронина, Г. И. и др. (2025) Некоторые показатели неспецифического иммунитета леща Abramis brama водохранилищ Средней Волги. Биология внутренних вод, т. 18, № 1, с. 226–231.
Яковенко, П. П., Авдеев, А. С. (2024) Роль цитокинов в иммунитете костистых рыб. В кн.: Н. Е. Горковенко (ред.). Актуальные проблемы ветеринарной медицины: состояние и решения: сборник статей по материалам Международной научно-практической конференции, посвященной 50-летию со дня основания факультета ветеринарной медицины. Краснодар: Изд-во Кубанского государственного аграрного университета им. И. Т. Трубилина, с. 193–197.
Agbede, S. A., Adedeji, O. B., Adeyemo, O. K. (2012) Tissues and orgars involved in the non-specific defence mechanism in fish: A review. Journal of Applied Sciences Research, vol. 8, no. 5, pp. 2493–2496.
Alejo, A., Tafalla, C. (2011) Chemokines in teleost fish species. Developmental and Comparative Immunology, vol. 35, no. 12, pp. 1215–1222. https://doi.org/10.1016/j.dci.2011.03.011
Bacon, K., Baggiolini, M., Broxmeyer, H. et al. (2002) Chemokine/chemokine receptor nomenclature. Journal of Interferon and Cytokine Research, vol. 22, no. 10, pp. 1067–1068. https://doi.org/10.1089/107999002760624305
Bayne, C. J., Gerwick, L. (1996) The acute phase response and innate immunity of fish. Developmental and Comparative Immunology, vol. 25, no. 8–9, pp. 725–743. https://doi.org/10.1016/s0145-305x(01)00033-7
Beck, G., Habicht, G. S. (2007) Immunity and the invertebrates. Scientific American, vol. 275, no. 5, pp. 60–66. https://doi.org/10.1038/scientificamerican1196-60
Bhatt, P., Kumaresan, V., Palanisamy, R. et al. (2018) A mini review on immune role of chemokines and its receptors in snakehead murrel Channa striatus. Fish and Shellfish Immunology, vol. 72, pp. 670–678. https://doi.org/10.1016/j.fsi.2017.11.036
Birkemo, G. A., Lüders, T., Andersen, Ø. et al. (2003) Hipposin, a histone-derived antimicrobial peptide in Atlantic halibut (Hippoglossus hippoglossus L.). Biochimica et Biophysica Acta (BBA) — Proteins and Proteomics, vol. 1646, no. 1-2, pp. 207–215. https://doi.org/10.1016/s1570-9639(03)00018-9
Boshra, H., Li, J., Sunyer, J. O. (2006) Recent advances on the complement system of teleost fish. Fish and Shellfish Immunology, vol. 20, no. 2, pp. 239–262. https://doi.org/10.1016/j.fsi.2005.04.004
Cocito, C. (1983) Properties of virginiamycin-like antibiotics (synergimycins), inhibitors containing synergistic components. In: F. E. Hahn (eds.). Modes and mechanisms of microbial growth inhibitors. Berlin; Heidelberg: Springer Publ., pp. 296–332. https://doi.org/10.1007/978-3-642-68946-8_19
Dong, X., Li, J., He, J. et al. (2016) Anti-infective mannose receptor immune mechanism in large yellow croaker (Larimichthys crocea). Fish and Shellfish Immunology, vol. 54, pp. 257–265. https://doi.org/10.1016/j.fsi.2016.04.006
Dong, X., Shilin, M. B., Apalikova, O. V. et al. (2021) The anti-infective immune mechanism of the CCL2 and CCL3 chemokines in the large yellow croaker (Larimichthys crocea). Journal of Applied Ichthyology, vol. 37, no. 6, pp. 807–996. https://doi.org/10.1111/jai.14214
Dunkelberger, J. R., Song, W.-C. (2010) Complement and its role in innate and adaptive immune responses. Cell Research, vol. 20, no. 1, pp. 34–50. https://doi.org/10.1038/cr.2009.139
East, L., Isacke, C. M. (2002) The mannose receptor family. Biochimica et Biophysica Acta (BBA) — General Subjects, vol. 1572, no. 2-3, pp. 364–386. https://doi.org/10.1016/s0304-4165(02)00319-7
Ellis, A. E. (1987) Inhibition of the Aeromonas salmonicida extracellular protease by α2-macroglobulin in the serum of rainbow trout. Microbial Pathogenesis, vol. 3, no. 3, pp. 167–177. https://doi.org/10.1016/0882-4010(87)90093-3
Ellis, A. E. (2001) Innate host defense mechanism of fish against viruses and bacteria. Developmental and Comparative Immunology, vol. 25, no. 8-9, pp. 827–239. https://doi.org/10.1016/s0145-305x(01)00038-6
Freedman, S. J. (1991) The role of alpha 2-macroglobulin in furunculosis: A comparison of rainbow trout and brook trout. Comparative Biochemistry and Physiology Part B: Comparative Biochemistry, vol. 98, no. 4, pp. 549–553. https://doi.org/10.1016/0305-0491(91)90252-9
Furnes, C., Seppola, M., Robertsen, B. (2009) Molecular characterisation and expression analysis of interferon gamma in Atlantic cod (Gadus morhua). Fish and Shellfish Immunology, vol. 26, no. 2, pp. 285–292. https://doi.org/10.1016/j.fsi.2008.12.002
Gazi, U., Martinez-Pomares, L. (2009) Influence of the mannose receptor in host immune responses. Immunobiology, vol. 214, no. 7, pp. 554–561. https://doi.org/10.1016/j.imbio.2008.11.004
Hellio, C., Pons, A. M., Beaupoil, C. et al. (2002) Antibacterial, antifungal and cytotoxic activities of extracts from fish epidermis and epidermal mucus. International Journal of Antimicrobial Agents, vol. 20, no. 3, pp. 214–219. https://doi.org/10.1016/S0924-8579(02)00172-3
Holland, M. C. H., Lambris, J. D. (2002) The complement system of teleosts. Fish and Shellfish Immunology, vol. 12, no. 5, pp. 399–420. https://doi.org/10.1006/fsim.2001.0408
Huang, Y., Yu, X.-Y., Luo, P. et al. (2023) Three novel L-type lectins from obscure puffer Takifugu obscurus promote antimicrobial immune response. Developmental and Comparative Immunology, vol. 149, article 105046. https://doi.org/10.1016/j.dci.2023.105046
Klockars, M., Roberts, P. (1976) Stimulations of phagocytosis by human lysozyme. Acta Haematologica, vol. 55, no. 5-2, pp. 289–295. https://doi.org/10.1159/000208029
Kugapreethan, R., Wan, Q., Nilojan, J., Lee, J. (2018) Identification and characterization of a calcium-dependent lily-type lectin from black rockfish (Sebastes schlegelii): Molecular antennas are involved in host defense via pathogen recognition. Developmental and Comparative Immunology, vol. 81, pp. 54–62. https://doi.org/10.1016/j.dci.2017.11.006
Kurtz, J. (2005) Specific memory within innate immune systems. Trends in Immunology, vol. 26, no. 4, pp. 186–192. https://doi.org/10.1016/j.it.2005.02.001
Laing, K. J., Secombes, C. J. (2004) Chemokines. Developmental and Comparative Immunology, vol. 28, no. 5, pp. 443–460. https://doi.org/10.1016/j.dci.2003.09.006
Liu, J., Liu, X., Wang, Z., Zhang, Q. (2022) Immunological characterization and function analysis of L-type lectin from spotted knifejaw, Oplegnathus punctatus. Frontiers in Immunology, vol. 13, article 993777. https://doi.org/10.3389/fimmu.2022.993777
Liu, X., Tang, X., Wang, L. et al. (2014) Molecular cloning and expression analysis of mannose receptor in blunt snout bream (Megalobrama amblycephala). Molecular Biology Reports, vol. 41, no. 7, pp. 4601–4611. https://doi.org/10.1007/s11033-014-3331-2
Maier, V. H., Dorn, K. V., Gudmundsdottir, B. K., Gudmundsson, G. H. (2008) Characterisation of cathelicidin gene family members in divergent fish species. Molecular Immunology, vol. 45, no. 14, pp. 3723–3730. https://doi.org/10.1016/j.molimm.2008.06.002
Mu, Y., Zhou, S., Ding, N. et al. (2019) Molecular characterization of a new fish specific chemokine CXCL_F6 in large yellow croaker (Larimichthys crocea) and its role in inflammatory response. Fish and Shellfish Immunology, vol. 84, pp. 787–794. https://doi.org/10.1016/j.fsi.2018.10.068
Natnan, M. E., Low, C.-F., Chong, C.-M. et al. (2021) Integration of omics tools for understanding the fish immune response due to microbial challenge. Frontiers in Marine Science, vol. 8, article 668771. https://doi.org/10.3389/fmars.2021.668771
Neves, J. V., Wilson, J. M., Rodrigues, P. N. S. (2009) Transferrin and ferritin response to bacterial infection: The role of the liver and brain in fish. Developmental and Comparative Immunology, vol. 33, no. 7, pp. 848–857. https://doi.org/10.1016/j.dci.2009.02.001
Nomiyama, H., Hieshima, K., Osada, N. et al. (2008) Extensive expansion and diversification of the chemokine gene family in zebrafish: Identification of a novel chemokine subfamily CX. BMC Genomics, vol. 9, article 222. https://doi.org/10.1186/1471-2164-9-222
Pajuelo, D., Lee, C.-T., Roig, F. J. et al. (2015) Novel host-specific iron acquisition system in the zoonotic pathogen Vibrio vulnificus. Environmental Microbiology, vol. 17, no. 6, pp. 2076–2089. https://doi.org/10.1111/1462-2920.12782
Redmond, A. K., Zou, J., Secombes, C. J. et al. (2019) Discovery of all three types in cartilaginous fishes enables phylogenetic resolution of the origins and evolution of interferons. Frontiers in Immunology, vol. 10, article 1558. https://doi.org/10.3389/fimmu.2019.01558
Robertsen, B. (2006) The interferon system of teleost fish. Fish and Shellfish Immunology, vol. 20, no. 2, pp. 172–191. https://doi.org/10.1016/j.fsi.2005.01.010
Robertsen, B., Bergan, V., Røkenes, T. et al. (2003) Atlantic salmon interferon genes: Cloning, sequence analysis, expression, and biological activity. Journal of Interferon and Cytokine Research, vol. 23, no. 10, pp. 601–612. https://doi.org/10.1089/107999003322485107
Salton, M. R. J., Ghuysen, J. M. (1959) The structure of di- and tetra-saccharides released from cell walls by lysozyme and streptomyces F1 enzyme and the β (1→4) N-acetylhexosaminidase activity of these enzymes. Biochimica et Biophysica Acta, vol. 36, no. 2, pp. 552–554. https://doi.org/10.1016/0006-3002(59)90205-7
Saurabh, S., Sahoo, P. K. (2008) Lysozyme: An important defence molecule of fish innate immune system. Aquaculture Research, vol. 39, no. 3, pp. 233–239. https://doi.org/10.1111/j.1365-2109.2007.01883.x
Smith, N. C., Rise, M. L., Christian, S. L. (2019) A comparison of the innate and adaptive immune systems in cartilaginous fish, ray-finned fish, and lobe-finned fish. Frontiers in Immunology, vol. 10, article 2292. https://doi.org/10.3389/fimmu.2019.02292
Stafford, J. L., Belosevic, M. (2003) Transferrin and the innate immune response of fish: identification of a novel mechanism of macrophage activation. Developmental and Comparative Immunology, vol. 27, no. 6–7, pp. 539–554. https://doi.org/10.1016/s0145-305x(02)00138-6
Sun, B., Lei, Y., Cao, Z. et al. (2019) TroCCL4, a CC chemokine of Trachinotus ovatus, is involved in the antimicrobial immune response. Fish and Shellfish Immunology, vol. 86, pp. 525–535. https://doi.org/10.1016/j.fsi.2018.11.080
Tasumi, S., Ohira, T., Kawazoe, I. et al. (2002) Primary structure and characteristics of a lectin from skin mucus of the Japanese eel Anguilla japonica. Journal of Biological Chemistry, vol. 277, no. 30, pp. 27305–27311. https://doi.org/10.1074/jbc.M202648200
Tkachenko, H., Grudniewska, J. (2017) Antioxidant defense in the brain tissue of rainbow trout (Oncorhynchus mykiss Walbaum) immunized by anti-Aeromonas vaccine. Aktual’nye problemy intensivnogo razvitiya zhivotnovodstva, no. 20-2, pp. 326–331.
Wang, J., Meng, Z., Wang, G. et al. (2020) A CCL25 chemokine functions as a chemoattractant and an immunomodulator in black rockfish, Sebastes schlegelii. Fish and Shellfish Immunology, vol. 100, pp. 161–170. https://doi.org/10.1016/j.fsi.2020.02.063
Wang, T., Liang, J., Xiang, X. et al. (2019) Functional identification and expressional responses of large yellow croaker (Larimichthys crocea) interleukin-8 and its receptor. Fish and Shellfish Immunology, vol. 87, pp. 470–477. https://doi.org/10.1016/j.fsi.2019.01.035
Wang, X.-A., Ma, A.-J., Sun, Z.-B. (2021) Genetic parameters of seven immune factors in turbot (Scophthalmus maximus) infected with Vibrio anguillarum. Journal of Fish Diseases, vol. 44, no. 3, pp. 263–271. https://doi.org/10.1111/jfd.13320
Wangkahart, E., Secombes, C. J., Wang, T. (2019) Studies on the use of flagellin as an immunostimulant and vaccine adjuvant in fish aquaculture. Frontiers in Immunology, vol. 9, article 3054. https://doi.org/10.3389/fimmu.2018.03054
Wu, C., Zhao, X., Babu, V. S. et al. (2018) Distribution of mannose receptor in blunt snout bream (Megalobrama amblycephala) during the embryonic development and its immune response to the challenge of Aeromonas hydrophila. Fish and Shellfish Immunology, vol. 78, pp. 52–59. https://doi.org/10.1016/j.fsi.2018.03.049
Yu, L., Li, C.-H., Chen, J. (2019) A novel CC chemokine ligand 2 like gene from ayu Plecoglossus altivelis is involved in the innate immune response against Vibrio anguillarum. Fish and Shellfish Immunology, vol. 87, pp. 886–896. https://doi.org/10.1016/j.fsi.2019.02.019
Zhao, X., Liu, L., Hegazy, A. M. et al. (2015) Mannose receptor mediated phagocytosis of bacteria in macrophages of blunt snout bream (Megalobrama amblycephala) in a Ca2+-dependent manner. Fish and Shellfish Immunology, vol. 43, no. 2, pp. 357–363. https://doi.org/10.1016/j.fsi.2015.01.002
Zheng, F., Asim, M., Lan, J. et al. (2015) Molecular cloning and functional characterization of mannose receptor in zebra fish (Danio rerio) during infection with Aeromonas sobria. International Journal of Molecular Sciences, vol. 16, no. 5, pp. 10997–11012. https://doi.org/10.3390/ijms160510997
Zou, J., Secombes, C. J. (2011) Teleost fish interferons and their role in immunity. Developmental and Comparative Immunology, vol. 35, no. 12, pp. 1376–1387. https://doi.org/10.1016/j.dci.2011.07.001
Zou, J., Carrington, A., Collet, B. et al. (2005) Identification and bioactivities of IFN-γ in rainbow trout Oncorhynchus mykiss: The first Th1-type cytokine characterized functionally in fish. The Journal of Immunology, vol. 175, no. 4, pp. 2484–2494. https://doi.org/10.4049/jimmunol.175.4.2484
Zou, J., Gorgoglione, B., Taylor, N. G. H. et al. (2014) Salmonids have an extraordinary complex type I IFN system: Characterization of the IFN locus in rainbow trout Oncorhynchus mykiss reveals two novel IFN subgroups. The Journal of Immunology, vol. 193, no. 5, pp. 2273–2286. https://doi.org/10.4049/jimmunol.1301796
REFERENCES
Agbede, S. A., Adedeji, O. B., Adeyemo, O. K. (2012) Tissues and orgars involved in the non-specific defence mechanism in fish: A review. Journal of Applied Sciences Research, vol. 8, no. 5, pp. 2493–2496. (In English)
Ahmedova, S. D. (2014) Pokazateli gumoral’nogo immuniteta u patsientov s mikoticheskim porazheniem kozhi i ee pridatkov [Indices of humoral immunity in patients with mycotic lesions of skin and its appendages]. Meditsinskie novosti, no. 3, pp. 77–79. (In Russian)
Alejo, A., Tafalla, C. (2011) Chemokines in teleost fish species. Developmental and Comparative Immunology, vol. 35, no. 12, pp. 1215–1222. https://doi.org/10.1016/j.dci.2011.03.011 (In English)
Alekseeva, I. G., Artamonov, N. A. (2023) Saprolegnioz ryb v promyshlennom rybovodstve [Saprolegniosis of fish in industrial fish culture]. In: Veterinarnaya meditsina: svyaz’ pokolenij kak faktor ustojchivogo razvitiya Rossii. Materialy Mezhdunarodnoj konferentsii [Veterinary medicine: Generational connection as a factor of sustainable development in Russia. Proceedings of the International conference]. Omsk: Omsk State Agrarian University named after P. Stolypin Publ., pp. 153–158. (In Russian)
Apalikova, O. V., Kiseleva, M. N., Mitryushkina, D. K. et al. (2025) Molekulyarno-geneticheskie podkhody k izucheniyu ustojchivosti ryb k boleznyam [Molecular genetic approaches to the study of fish resistance to diseases]. Trudy VNIRO, vol. 199, pp. 142–156. https://doi.org/10.36038/2307-3497-2025-199-142-156 (In Russian)
Bacon, K., Baggiolini, M., Broxmeyer, H. et al. (2002) Chemokine/chemokine receptor nomenclature. Journal of Interferon and Cytokine Research, vol. 22, no. 10, pp. 1067–1068. https://doi.org/10.1089/107999002760624305 (In English)
Bayne, C. J., Gerwick, L. (1996) The acute phase response and innate immunity of fish. Developmental and Comparative Immunology, vol. 25, no. 8–9, pp. 725–743. https://doi.org/10.1016/s0145-305x(01)00033-7 (In English)
Beck, G., Habicht, G. S. (2007) Immunity and the invertebrates. Scientific American, vol. 275, no. 5, pp. 60–66. https://doi.org/10.1038/scientificamerican1196-60 (In English)
Bhatt, P., Kumaresan, V., Palanisamy, R. et al. (2018) A mini review on immune role of chemokines and its receptors in snakehead murrel Channa striatus. Fish and Shellfish Immunology, vol. 72, pp. 670–678. https://doi.org/10.1016/j.fsi.2017.11.036 (In English)
Birkemo, G. A., Lüders, T., Andersen, Ø. et al. (2003) Hipposin, a histone-derived antimicrobial peptide in Atlantic halibut (Hippoglossus hippoglossus L.). Biochimica et Biophysica Acta (BBA) — Proteins and Proteomics, vol. 1646, no. 1-2, pp. 207–215. https://doi.org/10.1016/s1570-9639(03)00018-9 (In English)
Boshra, H., Li, J., Sunyer, J. O. (2006) Recent advances on the complement system of teleost fish. Fish and Shellfish Immunology, vol. 20, no. 2, pp. 239–262. https://doi.org/10.1016/j.fsi.2005.04.004 (In English)
Cocito, C. (1983) Properties of virginiamycin-like antibiotics (synergimycins), inhibitors containing synergistic components. In: F. E. Hahn (eds.). Modes and mechanisms of microbial growth inhibitors. Berlin; Heidelberg: Springer Publ., pp. 296–332. https://doi.org/10.1007/978-3-642-68946-8_19 (In English)
Delyagin, W. M., Scvortsova, Yu. V., Raimulla, Ye. A. (2025) Osteoimmunologiya [Osteoimmunology]. Lechenie i profilaktika, vol. 15, no. 2, pp. 5–19. (In Russian)
Dong, X., Li, J., He, J. et al. (2016) Anti-infective mannose receptor immune mechanism in large yellow croaker (Larimichthys crocea). Fish and Shellfish Immunology, vol. 54, pp. 257–265. https://doi.org/10.1016/j.fsi.2016.04.006 (In English)
Dong, X., Shilin, M. B., Apalikova, O. V. et al. (2021) The anti-infective immune mechanism of the CCL2 and CCL3 chemokines in the large yellow croaker (Larimichthys crocea). Journal of Applied Ichthyology, vol. 37, no. 6, pp. 807–996. https://doi.org/10.1111/jai.14214 (In English)
Dong, X., Shilin, M. B., Leonteva, E. O. (2024) Razvitie novykh tekhnologij zashchity ryb ot infektsij v usloviyakh akvakul’tury [Development of improved technologies for protecting fish from infections in aquaculture]. Arktika i innovatsii — Arctic and Innovations, vol. 2, no. 3, pp. 64–82. https://doi.org/10.21443/3034-1434-2024-2-3-64-82 (In Russian)
Droshnev, A. E., Zavyalova, E. A., Gulukin, M. I., Hlunov, O. B. (2012) Sovremennaya vaktsinoprofilaktika raduzhnoj foreli protiv vibrioza [Modern vaccinal prevention of rainbow trout against vibriosis]. Rossijskij veterinarnyj zhurnal. Sel’skokhozyajstvennye zhivotnye, no. 1, pp. 31–33. (In Russian)
Dunkelberger, J. R., Song, W.-C. (2010) Complement and its role in innate and adaptive immune responses. Cell Research, vol. 20, no. 1, pp. 34–50. https://doi.org/10.1038/cr.2009.139 (In English)
East, L., Isacke, C. M. (2002) The mannose receptor family. Biochimica et Biophysica Acta (BBA) — General Subjects, vol. 1572, no. 2–3, pp. 364–386. https://doi.org/10.1016/s0304-4165(02)00319-7 (In English)
Ellis, A. E. (1987) Inhibition of the Aeromonas salmonicida extracellular protease by α2-macroglobulin in the serum of rainbow trout. Microbial Pathogenesis, vol. 3, no. 3, pp. 167–177. https://doi.org/10.1016/0882-4010(87)90093-3 (In English)
Ellis, A. E. (2001) Innate host defense mechanism of fish against viruses and bacteria. Developmental and Comparative Immunology, vol. 25, no. 8-9, pp. 827–239. https://doi.org/10.1016/s0145-305x(01)00038-6 (In English)
Egorova, E. V., Krenev, I. A., Oborin, N. N., Berlov, M. N. (2023) Antimikrobnaya aktivnost’ sistemy komplementa [Antimicrobial activity of the complement system]. Meditsinskij akademicheskij zhurnal — Medical Academic Journal, vol. 23, no. 2, pp. 31–45. https://doi.org/10.17816/MAJ322841 (In Russian)
Freedman, S. J. (1991) The role of alpha 2-macroglobulin in furunculosis: A comparison of rainbow trout and brook trout. Comparative Biochemistry and Physiology Part B: Comparative Biochemistry, vol. 98, no. 4, pp. 549–553. https://doi.org/10.1016/0305-0491(91)90252-9 (In English)
Furnes, C., Seppola, M., Robertsen, B. (2009) Molecular characterisation and expression analysis of interferon gamma in Atlantic cod (Gadus morhua). Fish and Shellfish Immunology, vol. 26, no. 2, pp. 285–292. https://doi.org/10.1016/j.fsi.2008.12.002 (In English)
Gazi, U., Martinez-Pomares, L. (2009) Influence of the mannose receptor in host immune responses. Immunobiology, vol. 214, no. 7, pp. 554–561. https://doi.org/10.1016/j.imbio.2008.11.004 (In English)
Hellio, C., Pons, A. M., Beaupoil, C. et al. (2002) Antibacterial, antifungal and cytotoxic activities of extracts from fish epidermis and epidermal mucus. International Journal of Antimicrobial Agents, vol. 20, no. 3, pp. 214–219. https://doi.org/10.1016/S0924-8579(02)00172-3 (In English)
Holland, M. C. H., Lambris, J. D. (2002) The complement system of teleosts. Fish and Shellfish Immunology, vol. 12, no. 5, pp. 399–420. https://doi.org/10.1006/fsim.2001.0408 (In English)
Huang, Y., Yu, X.-Y., Luo, P. et al. (2023) Three novel L-type lectins from obscure puffer Takifugu obscurus promote antimicrobial immune response. Developmental and Comparative Immunology, vol. 149, article 105046. https://doi.org/10.1016/j.dci.2023.105046 (In English)
Klockars, M., Roberts, P. (1976) Stimulations of phagocytosis by human lysozyme. Acta Haematologica, vol. 55, no. 5-2, pp. 289–295. https://doi.org/10.1159/000208029 (In English)
Kochneva, A. A., Kuritsyn, A. E., Murzina, S. A. (2025) Vyyavlenie in silico antimikrobnykh posledovatel’nostej belkov mozga raduzhnoj foreli v aspekte kompleksnoj pererabotki otkhodov akvakul’tury [Identification in silico of antimicrobial sequences of rainbow trout brain proteins in the context of integrated in the context of complex recycling of aquaculture wastes]. Doklady Rossijskoj akademii nauk. Nauki o zhizni, vol. 520, pp. 50–56. https://doi.org/10.7868/S3034505725010099 (In Russian)
Kugapreethan, R., Wan, Q., Nilojan, J., Lee, J. (2018) Identification and characterization of a calcium-dependent lily-type lectin from black rockfish (Sebastes schlegelii): Molecular antennas are involved in host defense via pathogen recognition. Developmental and Comparative Immunology, vol. 81, pp. 54–62. https://doi.org/10.1016/j.dci.2017.11.006 (In English)
Kurtz, J. (2005) Specific memory within innate immune systems. Trends in Immunology, vol. 26, no. 4, pp. 186–192. https://doi.org/10.1016/j.it.2005.02.001 (In English)
Kuz’mina, V. V. (2016) Vliyanie tsinka i medi na aktivnost’ proteaz pishchevaritel’nogo trakta, obespechvayushchikh nespetsficheskuyu zashchitu ryb [Effect of zinc and copper on the gut protease activity providing non-specific protection of fish]. Trudy VNIRO, vol. 162, pp. 64–72. (In Russian)
Laing, K. J., Secombes, C. J. (2004) Chemokines. Developmental and Comparative Immunology, vol. 28, no. 5, pp. 443–460. https://doi.org/10.1016/j.dci.2003.09.006 (In English)
Liu, J., Liu, X., Wang, Z., Zhang, Q. (2022) Immunological characterization and function analysis of L-type lectin from spotted knifejaw, Oplegnathus punctatus. Frontiers in Immunology, vol. 13, article 993777. https://doi.org/10.3389/fimmu.2022.993777 (In English)
Liu, X., Tang, X., Wang, L. et al. (2014) Molecular cloning and expression analysis of mannose receptor in blunt snout bream (Megalobrama amblycephala). Molecular Biology Reports, vol. 41, no. 7, pp. 4601–4611. https://doi.org/10.1007/s11033-014-3331-2 (In English)
Maier, V. H., Dorn, K. V., Gudmundsdottir, B. K., Gudmundsson, G. H. (2008) Characterisation of cathelicidin gene family members in divergent fish species. Molecular Immunology, vol. 45, no. 14, pp. 3723–3730. https://doi.org/10.1016/j.molimm.2008.06.002 (In English)
Merkulova, N. L., Grekhneva, E. V., Chujkova, S. V. (2021) Izuchenie vozmozhnosti ispol’zovaniya epidermal’noj slizi Clarias gariepinus kak novogo istochnika farmakologicheski orientirovannykh preparatov [Studying the possibility of using the epidermal mucus of Clarias gariepinus as a new source of pharmacologically oriented drugs]. In: A. G. Gabibov, M. A. Ostrovskij (eds.). III ob’edinennyj nauchnyj forum fiziologov, biokhimikov i molekulyarnykh biologov. VII s’ezd biokhimikov Rossii. X rossijskij simpozium “Belki i peptidy”. VII s’ezd fiziologov SNG [III Joint scientific forum of physiologists, biochemists, and molecular biologists. VII Congress of Russian biochemists. X Russian symposium “Proteins and peptides”. VII Congress of CIS physiologists]. Vol. 2. Moscow: Pero Publ., p. 138. (In Russian)
Mikryakov, V. R., Mikryakov, D. V. (2015) Immunologicheskaya indikatsiya zdorov’ya ryb [Immunological indication of fish health]. Voprosy ikhtiologii — Journal of Ichthyology, vol. 55, no. 1, pp. 119–123. https://doi.org/10.7868/S0042875215010129 (In Russian)
Mu, Y., Zhou, S., Ding, N. et al. (2019) Molecular characterization of a new fish specific chemokine CXCL_F6 in large yellow croaker (Larimichthys crocea) and its role in inflammatory response. Fish and Shellfish Immunology, vol. 84, pp. 787–794. https://doi.org/10.1016/j.fsi.2018.10.068 (In English)
Natnan, M. E., Low, C.-F., Chong, C.-M. et al. (2021) Integration of omics tools for understanding the fish immune response due to microbial challenge. Frontiers in Marine Science, vol. 8, article 668771. https://doi.org/10.3389/fmars.2021.668771 (In English)
Neves, J. V., Wilson, J. M., Rodrigues, P. N. S. (2009) Transferrin and ferritin response to bacterial infection: The role of the liver and brain in fish. Developmental and Comparative Immunology, vol. 33, no. 7, pp. 848–857. https://doi.org/10.1016/j.dci.2009.02.001 (In English)
Nomiyama, H., Hieshima, K., Osada, N. et al. (2008) Extensive expansion and diversification of the chemokine gene family in zebrafish: Identification of a novel chemokine subfamily CX. BMC Genomics, vol. 9, article 222. https://doi.org/10.1186/1471-2164-9-222 (In English)
Pajuelo, D., Lee, C.-T., Roig, F. J. et al. (2015) Novel host-specific iron acquisition system in the zoonotic pathogen Vibrio vulnificus. Environmental Microbiology, vol. 17, no. 6, pp. 2076–2089. https://doi.org/10.1111/1462-2920.12782 (In English)
Pronina, G. I. (2014) Vozmozhnost’ povysheniya immunnoj ustojchivosti gidrobiontov v akvakul’ture [The possibility to increase the immune resistance of hydrobionts in aquaculture]. Izvestiya Orenburgskogo gosudarstvennogo agrarnogo universiteta — Izvestia Orenburg State Agrarian University, no. 3 (47), pp. 180–182. (In Russian)
Pronina, G. I., Ivanov, A. A., Mannapov, A. G., Sanaya, O. V. (2021) Immunitet pojkilotermnykh gidrobiontov [Immune system of poikilothermic aquatic organisms]. Izvestiya Timiryazevskoj sel’skokhozyajstvennoj akademii — Izvestiya of Timiryazev Agricultural Academy, no. 2, pp. 71–91. https://doi.org/10.26897/0021-342X-2021-2-71-91 (In Russian)
Redmond, A. K., Zou, J., Secombes, C. J. et al. (2019) Discovery of all three types in cartilaginous fishes enables phylogenetic resolution of the origins and evolution of interferons. Frontiers in Immunology, vol. 10, article 1558. https://doi.org/10.3389/fimmu.2019.01558 (In English)
Robertsen, B. (2006) The interferon system of teleost fish. Fish and Shellfish Immunology, vol. 20, no. 2, pp. 172–191. https://doi.org/10.1016/j.fsi.2005.01.010 (In English)
Robertsen, B., Bergan, V., Røkenes, T. et al. (2003) Atlantic salmon interferon genes: Cloning, sequence analysis, expression, and biological activity. Journal of Interferon and Cytokine Research, vol. 23, no. 10, pp. 601–612. https://doi.org/10.1089/107999003322485107 (In English)
Salton, M. R. J., Ghuysen, J. M. (1959) The structure of di- and tetra-saccharides released from cell walls by lysozyme and streptomyces F1 enzyme and the β (1→4) N-acetylhexosaminidase activity of these enzymes. Biochimica et Biophysica Acta, vol. 36, no. 2, pp. 552–554. https://doi.org/10.1016/0006-3002(59)90205-7 (In English)
Saurabh, S., Sahoo, P. K. (2008) Lysozyme: An important defence molecule of fish innate immune system. Aquaculture Research, vol. 39, no. 3, pp. 233–239. https://doi.org/10.1111/j.1365-2109.2007.01883.x (In English)
Smith, N. C., Rise, M. L., Christian, S. L. (2019) A comparison of the innate and adaptive immune systems in cartilaginous fish, ray-finned fish, and lobe-finned fish. Frontiers in Immunology, vol. 10, article 2292. https://doi.org/10.3389/fimmu.2019.02292 (In English)
Stafford, J. L., Belosevic, M. (2003) Transferrin and the innate immune response of fish: identification of a novel mechanism of macrophage activation. Developmental and Comparative Immunology, vol. 27, no. 6-7, pp. 539–554. https://doi.org/10.1016/s0145-305x(02)00138-6 (In English)
Subbotkin, M. F., Subbotkina, T. A. (2020) Vliyanie zarazheniya i in’ektsij substantsij razlichnoj prirody na lizotsim karpovykh ryb (Cyprinidae) (obzor) [Effect of infection and injections of substances of different origin on lysozyme in cyprinids (Cyprinidae) (review)]. Biologiya vnutrennikh vod, no. 2, pp. 180–191 https://doi.org/10.31857/S0320965220020151 (In Russian)
Subbotkina, T. A., Subbotkin, M. F. (2003) Osobennosti aktivnosti lizotsima nekotorykh vidov kostistykh ryb r. Volgi [Patterns of the lysozyme activity in some species of teleost fishes from the Volga river]. Biologiya vnutrennikh vod, no. 1, pp. 102–107. (In Russian)
Sun, B., Lei, Y., Cao, Z. et al. (2019) TroCCL4, a CC chemokine of Trachinotus ovatus, is involved in the antimicrobial immune response. Fish and Shellfish Immunology, vol. 86, pp. 525–535. https://doi.org/10.1016/j.fsi.2018.11.080 (In English)
Suvorova, T. A., Mikryakov, D. V., Pronina, G. I. et al. (2025) Nekotorye pokazateli nespetsificheskogo immuniteta leshcha Abramis brama vodokhranilishch Srednej Volgi [Some indicators of nonspecific immunity of bream Abramis brama in reservoirs of the middle Volga]. Biologiya vnutrennikh vod, vol. 18, no. 1, pp. 226–231. (In Russian)
Tasumi, S., Ohira, T., Kawazoe, I. et al. (2002) Primary structure and characteristics of a lectin from skin mucus of the Japanese eel Anguilla japonica. Journal of Biological Chemistry, vol. 277, no. 30, pp. 27305–27311. https://doi.org/10.1074/jbc.M202648200 (In English)
Tkachenko, H., Grudniewska, J. (2017) Antioxidant defense in the brain tissue of rainbow trout (Oncorhynchus mykiss Walbaum) immunized by anti-Aeromonas vaccine. Aktual’nye problemy intensivnogo razvitiya zhivotnovodstva, no. 20-2, pp. 326–331. (In English)
Vavilenkova, J. A. (2012) Sovremennye predstavleniya o sisteme interferona [Modern conception of interferon system]. Vestnik Smolenskoj gosudarstvennoj meditsinskoj akademii — Vestnik of the Smolensk State Medical Academy, vol. 11, no. 2, pp. 74–82. (In Russian)
Vylka, M. M., Lyutikov, A. A., Dyakova, S. A. et al. (2025) Mikrobiota kishechnika proizvoditelej sigovykh na primere Coregonus nasus v akvakul’ture v nerestovyj period [Gut microbiota of healthy and saprolegniosis-affected whitefish (Coregonus nasus) producers in aquaculture]. Ecologicheskaya genetika — Ecological Genetics, vol. 23, no. 3, pp. 225–234. https://doi.org/10.17816/ecogen679052 (In Russian)
Wang, J., Meng, Z., Wang, G. et al. (2020) A CCL25 chemokine functions as a chemoattractant and an immunomodulator in black rockfish, Sebastes schlegelii. Fish and Shellfish Immunology, vol. 100, pp. 161–170. https://doi.org/10.1016/j.fsi.2020.02.063 (In English)
Wang, T., Liang, J., Xiang, X. et al. (2019) Functional identification and expressional responses of large yellow croaker (Larimichthys crocea) interleukin-8 and its receptor. Fish and Shellfish Immunology, vol. 87, pp. 470–477. https://doi.org/10.1016/j.fsi.2019.01.035 (In English)
Wang, X.-A., Ma, A.-J., Sun, Z.-B. (2021) Genetic parameters of seven immune factors in turbot (Scophthalmus maximus) infected with Vibrio anguillarum. Journal of Fish Diseases, vol. 44, no. 3, pp. 263–271. https://doi.org/10.1111/jfd.13320 (In English)
Wangkahart, E., Secombes, C. J., Wang, T. (2019) Studies on the use of flagellin as an immunostimulant and vaccine adjuvant in fish aquaculture. Frontiers in Immunology, vol. 9, article 3054. https://doi.org/10.3389/fimmu.2018.03054 (In English)
Wu, C., Zhao, X., Babu, V. S. et al. (2018) Distribution of mannose receptor in blunt snout bream (Megalobrama amblycephala) during the embryonic development and its immune response to the challenge of Aeromonas hydrophila. Fish and Shellfish Immunology, vol. 78, pp. 52–59. https://doi.org/10.1016/j.fsi.2018.03.049 (In English)
Yakovenko, P. P., Avdeev, A. S. (2024) Rol’ tsitokinov v immunitete kostistykh ryb [Role of cytokines in immunity of bony fishes]. In: N. E. Gorkovenko (ed.). Aktual’nye problemy veterinarnoj meditsiny: sostoyanie i resheniya. Sbornik statej po materialam Mezhdunarodnoj nauchno-prakticheskoj konferentsii, posvyashchennoj 50-letiyu so dnya osnovaniya fakul’teta veterinarnoj meditsiny [Current veterinary medicine issues: Status and solutions. Collection of articles based on the materials of the International scientific and practical conference dedicated to the 50th anniversary of the Faculty of Veterinary Medicine]. Krasnodar: Kuban State Agrarian University named after I. T. Trubilin Publ., pp. 193–197. (In Russian)
Yu, L., Li, C.-H., Chen, J. (2019) A novel CC chemokine ligand 2 like gene from ayu Plecoglossus altivelis is involved in the innate immune response against Vibrio anguillarum. Fish and Shellfish Immunology, vol. 87, pp. 886–896. https://doi.org/10.1016/j.fsi.2019.02.019 (In English)
Zavyalova, E. A., Droshnev, A. E., Bulina, K. Yu. (2019) Vaktsinatsiya v akvakul’ture: istoriya, zakonomernosti i osobennosti provedeniya rabot [Vaccination in aquaculture: History, patterns and features of the work]. Rossijskij zhurnal Problemy veterinarnoj sanitarii, gigieny i ekologii — Russian Journal Problems of Veterinary Sanitation, Hygiene and Ecology, no. 1 (29), pp. 95–100. (In Russian)
Zhandalgarova, A. D., Mikryakov, D. V., Grozesku, Yu. N. et al. (2025) Immunomoduliruyushchee dejstvie kompleksa bifido- i laktobakterij na nespetsificheskij gumoral’nyj immunitet gibrida osetrovykh ryb [Immunomodulatory effect of bifidoand lactobacilli complex on non-specific humoral immunity of sturgeon fish hybrids]. Izvestiya Rossijskoj akademii nauk. Seriya biologicheskaya — Proceedings of the Russian Academy of Sciences. Biological Series, no. 2, pp. 249–254. (In Russian)
Zhao, X., Liu, L., Hegazy, A. M. et al. (2015) Mannose receptor mediated phagocytosis of bacteria in macrophages of blunt snout bream (Megalobrama amblycephala) in a Ca2+-dependent manner. Fish and Shellfish Immunology, vol. 43, no. 2, pp. 357–363. https://doi.org/10.1016/j.fsi.2015.01.002 (In English)
Zheng, F., Asim, M., Lan, J. et al. (2015) Molecular cloning and functional characterization of mannose receptor in zebra fish (Danio rerio) during infection with Aeromonas sobria. International Journal of Molecular Sciences, vol. 16, no. 5, pp. 10997–11012. https://doi.org/10.3390/ijms160510997 (In English)
Zou, J., Secombes, C. J. (2011) Teleost fish interferons and their role in immunity. Developmental and Comparative Immunology, vol. 35, no. 12, pp. 1376–1387. https://doi.org/10.1016/j.dci.2011.07.001 (In English)
Zou, J., Carrington, A., Collet, B. et al. (2005) Identification and bioactivities of IFN-γ in rainbow trout Oncorhynchus mykiss: The first Th1-type cytokine characterized functionally in fish. The Journal of Immunology, vol. 175, no. 4, pp. 2484–2494. https://doi.org/10.4049/jimmunol.175.4.2484 (In English)
Zou, J., Gorgoglione, B., Taylor, N. G. H. et al. (2014) Salmonids have an extraordinary complex type I IFN system: Characterization of the IFN locus in rainbow trout Oncorhynchus mykiss reveals two novel IFN subgroups. The Journal of Immunology, vol. 193, no. 5, pp. 2273–2286. https://doi.org/10.4049/jimmunol.1301796 (In English)
Zubareva, A. A., Lyutikov, A. A., Skorik, Yu. A. (2024) Sozdanie prototipov DNK-vaktsin dlya akvakultury na osnove prirodnykh polisakharidov [Creation of DNA vaccines for aquaculture based on natural polysaccharides]. In: K. V. Kolonchin, M. M. Mel’nik (eds.) Rybokhozyajstvennaya nauka. Istoriya, sovremennost’, perspektivy. Materialy Mezhdunarodnoj nauchno-prakticheskoj konferentsii, posvyashchennoj 110-letiyu sozdaniya “GosNIORKH im. L. S. Berga” [Fisheries science. History, modernity, prospects. Materials of the International scientific and practical conference dedicated to the 110th anniversary of the establishment of the L. S. Berg State Research Institute]. Moscow: Russian Federal Research Institute of Fisheries and Oceanography Publ., pp. 173–175. (In Russian)
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