{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,7,1]],"date-time":"2026-07-01T18:40:34Z","timestamp":1782931234735,"version":"3.54.5"},"reference-count":66,"publisher":"Springer Science and Business Media LLC","issue":"1","license":[{"start":{"date-parts":[[2026,2,2]],"date-time":"2026-02-02T00:00:00Z","timestamp":1769990400000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"},{"start":{"date-parts":[[2026,2,2]],"date-time":"2026-02-02T00:00:00Z","timestamp":1769990400000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/creativecommons.org\/licenses\/by\/4.0"}],"funder":[{"DOI":"10.13039\/501100009827","name":"Alexandria University","doi-asserted-by":"crossref","id":[{"id":"10.13039\/501100009827","id-type":"DOI","asserted-by":"crossref"}]}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Sci Rep"],"abstract":"<jats:title>Abstract<\/jats:title>\n                  <jats:p>\n                    This study evaluated and compared the effectiveness of\n                    <jats:italic>Bacillus<\/jats:italic>\n                    species probiotics and formalin-killed bacterin to control vibriosis caused by\n                    <jats:italic>Vibrio anguillarium<\/jats:italic>\n                    in European eel (\n                    <jats:italic>Anguilla anguilla<\/jats:italic>\n                    ) elvers. Fish were allocated into four triplicate groups defined as follows: a control non-vaccinated (CON) group that fed on a basal diet without additives, two groups fed on diets supplied with\n                    <jats:italic>Bacillus<\/jats:italic>\n                    species probiotics (PRO; Sanolife PRO-F) in a dose of 0.5\u00a0g \/kg diet (PRO-F1) and 1.0\u00a0g \/kg diet (PRO-F2), and the fourth group was vaccinated with immersion formalin-killed bacterin and got a booster vaccination at 14th day (VACC). The experiment lasted for 28 days. In the end, the immune responses, including antibody titers, lysozyme activities (in serum and skin mucous), and bactericidal activity, were evaluated. The relative percent survival (RPS) after intraperitoneal challenge with\n                    <jats:italic>V. anguillarium<\/jats:italic>\n                    was calculated after an additional 10-day observation period. A one-way ANOVA followed by Dunnett\u2019s multiple comparison test was used to compare the experimental groups (VACC, PRO-F1, and PRO-F2) with the CON group. The results revealed significantly higher antibody titers, lysozyme activities (in serum and skin mucous), and serum bactericidal activity in the VACC and PRO-treated groups over the CON group. Of interest, the lysozyme activities (in serum and skin mucous) and antibody titers were significantly higher in the VACC group over both PRO groups. After the experimental infection, it was noticed that both VACC and PRO significantly protected eel elvers against\n                    <jats:italic>V. anguillarium<\/jats:italic>\n                    compared to the CON group. In addition, the RPS were 41.67%, 29.17%, and 37.50% in the VACC, PRO-F1, and PRO-F2 groups, respectively. This study unveils the vital roles of PRO and VACC and their effective roles in enhancing innate and mucosal immunities and protecting European eel elvers against vibriosis caused by\n                    <jats:italic>V. anguillarium<\/jats:italic>\n                    . Based on our findings, immersion vaccination emerges as the superior strategy for managing vibriosis in eels. This approach offers greater protection than feed-based probiotics, underscoring its potential as a viable vaccination method in eel aquaculture.\n                  <\/jats:p>","DOI":"10.1038\/s41598-026-35298-8","type":"journal-article","created":{"date-parts":[[2026,2,2]],"date-time":"2026-02-02T07:14:16Z","timestamp":1770016456000},"update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":1,"title":["Comparative efficacy of Bacillus probiotics and formalin-killed bacterin against Vibrio anguillarum in European eel elvers"],"prefix":"10.1038","volume":"16","author":[{"given":"Hany M.R.","family":"Abdel-Latif","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nashwa","family":"Abdel-Razek","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Riad H.","family":"Khalil","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,2,2]]},"reference":[{"key":"35298_CR1","unstructured":"Nielsen, T. & Prouzet, P. Capture-based aquaculture of the wild European eel (Anguilla anguilla). Capture-based aquaculture. Global overview. FAO Fisheries Technical Paper, (508), 141\u2013168.\u200f (2008)."},{"key":"35298_CR2","doi-asserted-by":"crossref","unstructured":"Chart, H. & Munn, C. B. Experimental Vibriosis in the Eel (Anguilla anguilla). Fish Diseases, 39\u201344 (1980).","DOI":"10.1007\/978-3-642-67854-7_7"},{"key":"35298_CR3","doi-asserted-by":"publisher","first-page":"317","DOI":"10.1111\/j.1095-8649.1976.tb03955.x","volume":"8","author":"DH McCarthy","year":"1976","unstructured":"McCarthy, D. H. Vibrio disease in eels. J. Fish Biol. 8, 317\u2013320. https:\/\/doi.org\/10.1111\/j.1095-8649.1976.tb03955.x (1976).","journal-title":"J. Fish Biol."},{"key":"35298_CR4","unstructured":"Austin, B. & Austin, D. A. Vibrionaceae representatives. In: Bacterial Fish Pathogens (ed. by B. Austin & D.A. Austin), pp. 270\u2013276. Ellis Horwood Ltd, Chichester. (1999)."},{"key":"35298_CR5","doi-asserted-by":"publisher","first-page":"117","DOI":"10.3147\/jsfp.6.117","volume":"6","author":"Y Jo","year":"1972","unstructured":"Jo, Y. & Muroga, K. Vibrio anguillarum isolated from eels cultured in freshwater ponds. Fish Pathology 6, 117\u2013119 (1972).","journal-title":"Fish Pathology"},{"key":"35298_CR6","doi-asserted-by":"publisher","first-page":"187","DOI":"10.3354\/dao035187","volume":"35","author":"I Dalsgaard","year":"1999","unstructured":"Dalsgaard, I., H\u00f8i, L., Siebeling, R. J. & Dalsgaard, A. Indole-positive Vibrio vulnificus isolated from disease outbreaks on a Danish eel farm. Dis. Aquat. Org. 35, 187\u2013194 (1999).","journal-title":"Dis. Aquat. Org."},{"key":"35298_CR7","doi-asserted-by":"publisher","first-page":"285","DOI":"10.1111\/j.1365-2761.2006.00719.x","volume":"29","author":"B Fouz","year":"2006","unstructured":"Fouz, B., Larsen, J. L. & Amaro, C. Vibrio vulnificus serovar A: an emerging pathogen in European anguilliculture. J. Fish Dis. 29, 285\u2013291. https:\/\/doi.org\/10.1111\/j.1365-2761.2006.00719.x (2006).","journal-title":"J. Fish Dis."},{"key":"35298_CR8","unstructured":"B, A. & DA, A. Bacterial fish pathogens, disease of farmed and wild fish, 6th edn. Springer, Dordrecht. (2006)."},{"key":"35298_CR9","unstructured":"Bergman, A. M. Die roten Beulenkrankheit des Aales. Ber Bayerischen Biol Versuchs\u2013Station2 (in German), 10\u201354 (1909)."},{"key":"35298_CR10","doi-asserted-by":"publisher","first-page":"643","DOI":"10.1111\/j.1365-2761.2011.01279.x","volume":"34","author":"I Frans","year":"2011","unstructured":"Frans, I. et al. Vibrio anguillarum as a fish pathogen: virulence factors, diagnosis and prevention. J. Fish Dis. 34, 643\u2013661. https:\/\/doi.org\/10.1111\/j.1365-2761.2011.01279.x (2011).","journal-title":"J. Fish Dis."},{"key":"35298_CR11","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquaculture.2022.737951","volume":"551","author":"HMR Abdel-Latif","year":"2022","unstructured":"Abdel-Latif, H. M. R. et al. Shrimp vibriosis and possible control measures using probiotics, postbiotics, prebiotics, and synbiotics: A review. Aquaculture 551, 737951. https:\/\/doi.org\/10.1016\/j.aquaculture.2022.737951 (2022).","journal-title":"Aquaculture"},{"key":"35298_CR12","doi-asserted-by":"publisher","first-page":"585","DOI":"10.1111\/raq.12188","volume":"10","author":"ME Hickey","year":"2018","unstructured":"Hickey, M. E. & Lee, J.-L. A comprehensive review of Vibrio (Listonella) anguillarum: ecology, pathology and prevention. Rev. Aquac. 10, 585\u2013610. https:\/\/doi.org\/10.1111\/raq.12188 (2018).","journal-title":"Rev. Aquac."},{"key":"35298_CR13","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquaculture.2021.737514","volume":"547","author":"S Yilmaz","year":"2022","unstructured":"Yilmaz, S. et al. Probiotics, prebiotics, and synbiotics used to control vibriosis in fish: A review. Aquaculture 547, 737514. https:\/\/doi.org\/10.1016\/j.aquaculture.2021.737514 (2022).","journal-title":"Aquaculture"},{"key":"35298_CR14","doi-asserted-by":"publisher","first-page":"1137","DOI":"10.1111\/j.1462-2920.2006.01054.x","volume":"8","author":"FC Cabello","year":"2006","unstructured":"Cabello, F. C. Heavy use of prophylactic antibiotics in aquaculture: a growing problem for human and animal health and for the environment. Environ. Microbiol. 8, 1137\u20131144. https:\/\/doi.org\/10.1111\/j.1462-2920.2006.01054.x (2006).","journal-title":"Environ. Microbiol."},{"key":"35298_CR15","doi-asserted-by":"publisher","first-page":"139","DOI":"10.1046\/j.1365-2621.1998.3320139.x","volume":"33","author":"DJ Alderman","year":"1998","unstructured":"Alderman, D. J. & Hastings, T. S. Antibiotic use in aquaculture: development of antibiotic resistance \u2013 potential for consumer health risks*. Int. J. Food Sci. Technol. 33, 139\u2013155. https:\/\/doi.org\/10.1046\/j.1365-2621.1998.3320139.x (1998).","journal-title":"Int. J. Food Sci. Technol."},{"key":"35298_CR16","doi-asserted-by":"publisher","first-page":"255","DOI":"10.1046\/j.1365-2621.2003.00671.x","volume":"38","author":"K Holmstr\u00f6m","year":"2003","unstructured":"Holmstr\u00f6m, K. et al. Antibiotic use in shrimp farming and implications for environmental impacts and human health. Int. J. Food Sci. Technol. 38, 255\u2013266. https:\/\/doi.org\/10.1046\/j.1365-2621.2003.00671.x (2003).","journal-title":"Int. J. Food Sci. Technol."},{"key":"35298_CR17","doi-asserted-by":"publisher","first-page":"640","DOI":"10.1111\/raq.12344","volume":"12","author":"R Lulijwa","year":"2020","unstructured":"Lulijwa, R., Rupia, E. J. & Alfaro, A. C. Antibiotic use in aquaculture, policies and regulation, health and environmental risks: a review of the top 15 major producers. Rev. Aquac. 12, 640\u2013663. https:\/\/doi.org\/10.1111\/raq.12344 (2020).","journal-title":"Rev. Aquac."},{"key":"35298_CR18","doi-asserted-by":"publisher","first-page":"89","DOI":"10.1586\/14760584.4.1.89","volume":"4","author":"I Sommerset","year":"2005","unstructured":"Sommerset, I., Kross\u00f8y, B., Biering, E. & Frost, P. Vaccines for fish in aquaculture. Expert Rev. Vaccines 4, 89\u2013101. https:\/\/doi.org\/10.1586\/14760584.4.1.89 (2005).","journal-title":"Expert Rev. Vaccines"},{"key":"35298_CR19","doi-asserted-by":"publisher","first-page":"1317","DOI":"10.1007\/s10499-023-01218-0","volume":"32","author":"MM Abotaleb","year":"2024","unstructured":"Abotaleb, M. M. et al. Efficacy of combined inactivated vaccines against Vibrio alginolyticus and Streptococcus agalactiae infections in Nile tilapia in Egypt. Aquacult. Int. 32, 1317\u20131334. https:\/\/doi.org\/10.1007\/s10499-023-01218-0 (2024).","journal-title":"Aquacult. Int."},{"key":"35298_CR20","doi-asserted-by":"publisher","first-page":"210","DOI":"10.1016\/j.fsi.2019.04.066","volume":"90","author":"A Adams","year":"2019","unstructured":"Adams, A. Progress, challenges and opportunities in fish vaccine development. Fish Shellfish Immunol. 90, 210\u2013214. https:\/\/doi.org\/10.1016\/j.fsi.2019.04.066 (2019).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR21","doi-asserted-by":"publisher","first-page":"183","DOI":"10.3354\/dao045183","volume":"45","author":"B Fouz","year":"2001","unstructured":"Fouz, B. et al. Field testing of a vaccine against eel diseases caused by Vibrio vulnificus. Dis. Aquat. Org. 45, 183\u2013189 (2001).","journal-title":"Dis. Aquat. Org."},{"key":"35298_CR22","doi-asserted-by":"publisher","first-page":"93","DOI":"10.1016\/S1050-4648(03)00036-6","volume":"16","author":"MD Esteve-Gassent","year":"2004","unstructured":"Esteve-Gassent, M. D., Fouz, B. & Amaro, C. Efficacy of a bivalent vaccine against eel diseases caused by Vibrio vulnificus after its administration by four different routes. Fish Shellfish Immunol. 16, 93\u2013105. https:\/\/doi.org\/10.1016\/S1050-4648(03)00036-6 (2004).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR23","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.aquaculture.2004.07.006","volume":"241","author":"MD Esteve-Gassent","year":"2004","unstructured":"Esteve-Gassent, M. D., Barrera, R. & Amaro, C. Vaccination of market-size eels against vibriosis due to Vibrio vulnificus serovar E. Aquaculture 241, 9\u201319. https:\/\/doi.org\/10.1016\/j.aquaculture.2004.07.006 (2004).","journal-title":"Aquaculture"},{"key":"35298_CR24","doi-asserted-by":"publisher","first-page":"9","DOI":"10.1016\/j.aquaculture.2003.10.006","volume":"231","author":"MD Esteve-Gassent","year":"2004","unstructured":"Esteve-Gassent, M. D., Fouz, B., Barrera, R. & Amaro, C. Efficacy of oral reimmunisation after immersion vaccination against Vibrio vulnificus in farmed European eels. Aquaculture 231, 9\u201322. https:\/\/doi.org\/10.1016\/j.aquaculture.2003.10.006 (2004).","journal-title":"Aquaculture"},{"key":"35298_CR25","doi-asserted-by":"publisher","unstructured":"Abdel-Latif, H. M. R. et al. Control of yersiniosis in rainbow trout, Oncorhynchus mykiss: innovative non-antibiotic feed-based strategies. Annals of Animal Science 0, https:\/\/doi.org\/10.2478\/aoas-2024-0087 (2023).","DOI":"10.2478\/aoas-2024-0087"},{"key":"35298_CR26","doi-asserted-by":"publisher","first-page":"36","DOI":"10.1016\/j.fsi.2021.07.007","volume":"117","author":"MT El-Saadony","year":"2021","unstructured":"El-Saadony, M. T. et al. The functionality of probiotics in aquaculture: An overview. Fish Shellfish Immunol. 117, 36\u201352. https:\/\/doi.org\/10.1016\/j.fsi.2021.07.007 (2021).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR27","doi-asserted-by":"publisher","first-page":"147","DOI":"10.1016\/S0044-8486(99)00187-8","volume":"180","author":"FJ Gatesoupe","year":"1999","unstructured":"Gatesoupe, F. J. The use of probiotics in aquaculture. Aquaculture 180, 147\u2013165. https:\/\/doi.org\/10.1016\/S0044-8486(99)00187-8 (1999).","journal-title":"Aquaculture"},{"key":"35298_CR28","doi-asserted-by":"crossref","unstructured":"Mondal, S., Mondal, D., Mondal, T. & Malik, J. in Bacterial Fish Diseases (eds Gowhar Hamid Dar et al.) 351\u2013378 (Academic Press, 2022).","DOI":"10.1016\/B978-0-323-85624-9.00024-5"},{"key":"35298_CR29","doi-asserted-by":"publisher","first-page":"466","DOI":"10.1016\/j.fsi.2016.04.023","volume":"54","author":"BT Birhanu","year":"2016","unstructured":"Birhanu, B. T. et al. Immunomodulation of Lactobacillus pentosus PL11 against Edwardsiella tarda infection in the head kidney cells of the Japanese eel (Anguilla japonica). Fish Shellfish Immunol. 54, 466\u2013472. https:\/\/doi.org\/10.1016\/j.fsi.2016.04.023 (2016).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR30","doi-asserted-by":"publisher","first-page":"756","DOI":"10.1016\/j.fsi.2012.11.028","volume":"34","author":"J-S Lee","year":"2013","unstructured":"Lee, J.-S. et al. Effects of dietary supplementation of Lactobacillus pentosus PL11 on the growth performance, immune and antioxidant systems of Japanese eel Anguilla japonica challenged with Edwardsiella tarda. Fish Shellfish Immunol. 34, 756\u2013761. https:\/\/doi.org\/10.1016\/j.fsi.2012.11.028 (2013).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR31","doi-asserted-by":"publisher","first-page":"201","DOI":"10.1016\/j.fsi.2016.12.035","volume":"61","author":"S Lee","year":"2017","unstructured":"Lee, S. et al. Comparative evaluation of dietary probiotics Bacillus subtilis WB60 and Lactobacillus plantarum KCTC3928 on the growth performance, immunological parameters, gut morphology and disease resistance in Japanese eel. Anguilla japonica. Fish & Shellfish Immunology 61, 201\u2013210. https:\/\/doi.org\/10.1016\/j.fsi.2016.12.035 (2017).","journal-title":"Anguilla japonica. Fish & Shellfish Immunology"},{"key":"35298_CR32","doi-asserted-by":"publisher","first-page":"283","DOI":"10.1016\/j.fsi.2018.09.031","volume":"83","author":"S Lee","year":"2018","unstructured":"Lee, S. et al. Synergistic effects of dietary supplementation of Bacillus subtilis WB60 and mannanoligosaccharide (MOS) on growth performance, immunity and disease resistance in Japanese eel. Anguilla japonica. Fish & Shellfish Immunology 83, 283\u2013291. https:\/\/doi.org\/10.1016\/j.fsi.2018.09.031 (2018).","journal-title":"Anguilla japonica. Fish & Shellfish Immunology"},{"key":"35298_CR33","doi-asserted-by":"crossref","unstructured":"Jang, W. J. et al. Effect of Bacillus sp. supplementation diet on survival rate and microbiota composition in artificially produced eel larvae (Anguilla japonica). Frontiers in Microbiology, 2036 (2022).","DOI":"10.3389\/fmicb.2022.891070"},{"key":"35298_CR34","volume-title":"Studies on Bacterial Diseases Affecting Some Cultured Marine Fishes in Alexandria governorate","author":"H Abdel-Latif","year":"2013","unstructured":"Abdel-Latif, H. Studies on Bacterial Diseases Affecting Some Cultured Marine Fishes in Alexandria governorate (Alexandria University, Egypt, Faculty of Veterinary Medicine, 2013)."},{"key":"35298_CR35","doi-asserted-by":"publisher","DOI":"10.1016\/j.aqrep.2022.101385","volume":"27","author":"MAM El-Son","year":"2022","unstructured":"El-Son, M. A. M., Elshopakey, G. E., Rezk, S., Eldessouki, E. A. A. & Elbahnaswy, S. Dietary mixed Bacillus strains promoted the growth indices, enzymatic profile, intestinal immunity, and liver and intestinal histomorphology of Nile tilapia. Oreochromis niloticus. Aquaculture Reports 27, 101385. https:\/\/doi.org\/10.1016\/j.aqrep.2022.101385 (2022).","journal-title":"Oreochromis niloticus. Aquaculture Reports"},{"key":"35298_CR36","doi-asserted-by":"publisher","first-page":"91","DOI":"10.3354\/dao043091","volume":"43","author":"R Collado","year":"2000","unstructured":"Collado, R., Fouz, B., Sanju\u00e1n, E. & Amaro, C. Effectiveness of different vaccine formulations against vibriosis caused by Vibrio vulnificus serovar E (biotype 2) in European eels Anguilla anguilla. Dis. Aquat. Org. 43, 91\u2013101 (2000).","journal-title":"Dis. Aquat. Org."},{"key":"35298_CR37","doi-asserted-by":"publisher","unstructured":"Esteve-Gassent, M. D., Nielsen, M. E. & Amaro, C. The kinetics of antibody production in mucus and serum of European eel (Anguilla anguilla L.) after vaccination against Vibrio vulnificus: development of a new method for antibody quantification in skin mucus. Fish & Shellfish Immunology 15, 51\u201361, https:\/\/doi.org\/10.1016\/S1050-4648(02)00138-9 (2003).","DOI":"10.1016\/S1050-4648(02)00138-9"},{"key":"35298_CR38","doi-asserted-by":"publisher","first-page":"2475","DOI":"10.1128\/iai.65.6.2475-2479.1997","volume":"65","author":"C Amaro","year":"1997","unstructured":"Amaro, C., Fouz, B., Biosca, E. G., Marco-Noales, E. & Collado, R. The lipopolysaccharide O side chain of Vibrio vulnificus serogroup E is a virulence determinant for eels. Infect. Immun. 65, 2475\u20132479 (1997).","journal-title":"Infect. Immun."},{"key":"35298_CR39","doi-asserted-by":"publisher","first-page":"237","DOI":"10.1016\/S0044-8486(00)00345-8","volume":"188","author":"PH Klesius","year":"2000","unstructured":"Klesius, P. H., Shoemaker, C. A. & Evans, J. J. Efficacy of single and combined Streptococcus iniae isolate vaccine administered by intraperitoneal and intramuscular routes in tilapia (Oreochromis niloticus). Aquaculture 188, 237\u2013246. https:\/\/doi.org\/10.1016\/S0044-8486(00)00345-8 (2000).","journal-title":"Aquaculture"},{"key":"35298_CR40","doi-asserted-by":"publisher","unstructured":"Shoemaker, C. A., LaFrentz, B. R. & Klesius, P. H. Vaccination of sex reversed hybrid tilapia (Oreochromis niloticus\u00a0\u00d7\u00a0O.\u00a0aureus) with an inactivated Vibrio vulnificus vaccine. Biologicals 39, 424\u2013429, https:\/\/doi.org\/10.1016\/j.biologicals.2011.08.004 (2011).","DOI":"10.1016\/j.biologicals.2011.08.004"},{"key":"35298_CR41","doi-asserted-by":"publisher","first-page":"45","DOI":"10.1016\/j.aquaculture.2012.04.033","volume":"354\u2013355","author":"CA Shoemaker","year":"2012","unstructured":"Shoemaker, C. A., LaFrentz, B. R. & Klesius, P. H. Bivalent vaccination of sex reversed hybrid tilapia against Streptococcus iniae and Vibrio vulnificus. Aquaculture 354\u2013355, 45\u201349. https:\/\/doi.org\/10.1016\/j.aquaculture.2012.04.033 (2012).","journal-title":"Aquaculture"},{"key":"35298_CR42","unstructured":"Harrigan, W. F. & McCance, M. E. Laboratory methods in microbiology. (Academic press, 2014)."},{"key":"35298_CR43","first-page":"447","volume":"49","author":"DF Amend","year":"1981","unstructured":"Amend, D. F. Potency testing of fish vaccines. Dev. Biol. Stand. 49, 447\u2013454 (1981).","journal-title":"Dev. Biol. Stand."},{"key":"35298_CR44","doi-asserted-by":"publisher","first-page":"65","DOI":"10.1111\/j.1365-2761.2006.00695.x","volume":"29","author":"ME Nielsen","year":"2006","unstructured":"Nielsen, M. E. & Esteve-Gassent, M. D. The eel immune system: present knowledge and the need for research. J. Fish Dis. 29, 65\u201378. https:\/\/doi.org\/10.1111\/j.1365-2761.2006.00695.x (2006).","journal-title":"J. Fish Dis."},{"key":"35298_CR45","doi-asserted-by":"crossref","unstructured":"Krogdahl, \u00c5., Kortner, T. M. & L\u00f8kka, G. in Feed and Feeding for Fish and Shellfish (ed Vikas Kumar) 405\u2013459 (Academic Press, 2025).","DOI":"10.1016\/B978-0-443-21556-8.00005-3"},{"key":"35298_CR46","doi-asserted-by":"publisher","first-page":"525","DOI":"10.3390\/biology4030525","volume":"4","author":"I Salinas","year":"2015","unstructured":"Salinas, I. The Mucosal Immune System of Teleost Fish. Biology 4, 525\u2013539 (2015).","journal-title":"Biology"},{"key":"35298_CR47","doi-asserted-by":"publisher","unstructured":"Santos, R. A. et al. Isolation and Characterization of Fish-Gut Bacillus spp. as Source of Natural Antimicrobial Compounds to Fight Aquaculture Bacterial Diseases. Marine Biotechnology 23, 276\u2013293, https:\/\/doi.org\/10.1007\/s10126-021-10022-x (2021).","DOI":"10.1007\/s10126-021-10022-x"},{"key":"35298_CR48","doi-asserted-by":"publisher","first-page":"820","DOI":"10.1016\/j.fsi.2019.02.010","volume":"87","author":"FKA Kuebutornye","year":"2019","unstructured":"Kuebutornye, F. K. A., Abarike, E. D. & Lu, Y. A review on the application of Bacillus as probiotics in aquaculture. Fish Shellfish Immunol. 87, 820\u2013828. https:\/\/doi.org\/10.1016\/j.fsi.2019.02.010 (2019).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR49","doi-asserted-by":"publisher","first-page":"2","DOI":"10.1016\/j.fsi.2010.02.017","volume":"29","author":"SK Nayak","year":"2010","unstructured":"Nayak, S. K. Probiotics and immunity: A fish perspective. Fish Shellfish Immunol. 29, 2\u201314. https:\/\/doi.org\/10.1016\/j.fsi.2010.02.017 (2010).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR50","doi-asserted-by":"publisher","first-page":"223","DOI":"10.1111\/j.1365-2109.2007.01883.x","volume":"39","author":"S Saurabh","year":"2008","unstructured":"Saurabh, S. & Sahoo, P. K. Lysozyme: an important defence molecule of fish innate immune system. Aquac. Res. 39, 223\u2013239. https:\/\/doi.org\/10.1111\/j.1365-2109.2007.01883.x (2008).","journal-title":"Aquac. Res."},{"key":"35298_CR51","doi-asserted-by":"publisher","first-page":"5029","DOI":"10.1111\/are.15989","volume":"53","author":"SM Aboleila","year":"2022","unstructured":"Aboleila, S. M., Salah, A., Mohamed, R. A. & Diab, A. M. Combined effect of a mixture of Bacillus species and vaccination on haemato-biochemical parameters, immune response and transcriptomic changes of Nile tilapia challenged with Streptococcus iniae. Aquac. Res. 53, 5029\u20135044. https:\/\/doi.org\/10.1111\/are.15989 (2022).","journal-title":"Aquac. Res."},{"key":"35298_CR52","doi-asserted-by":"publisher","unstructured":"Kord, M. I. et al. The Immunostimulatory Effects of Commercial Feed Additives on Growth Performance, Non-specific Immune Response, Antioxidants Assay, and Intestinal Morphometry of Nile tilapia, Oreochromis niloticus. Frontiers in Physiology Volume 12 - 2021, https:\/\/doi.org\/10.3389\/fphys.2021.627499 (2021).","DOI":"10.3389\/fphys.2021.627499"},{"key":"35298_CR53","doi-asserted-by":"publisher","first-page":"496","DOI":"10.1016\/j.fsi.2015.03.004","volume":"44","author":"KM Selim","year":"2015","unstructured":"Selim, K. M. & Reda, R. M. Improvement of immunity and disease resistance in the Nile tilapia, Oreochromis niloticus, by dietary supplementation with Bacillus amyloliquefaciens. Fish Shellfish Immunol. 44, 496\u2013503. https:\/\/doi.org\/10.1016\/j.fsi.2015.03.004 (2015).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR54","doi-asserted-by":"publisher","first-page":"477","DOI":"10.1016\/j.fsi.2015.01.017","volume":"43","author":"G SongLin","year":"2015","unstructured":"SongLin, G. et al. A novel recombinant bivalent outer membrane protein of Vibrio vulnificus and Aeromonas hydrophila as a vaccine antigen of American eel (Anguilla rostrata). Fish Shellfish Immunol. 43, 477\u2013484. https:\/\/doi.org\/10.1016\/j.fsi.2015.01.017 (2015).","journal-title":"Fish Shellfish Immunol."},{"key":"35298_CR55","doi-asserted-by":"publisher","DOI":"10.1002\/mbo3.766","volume":"8","author":"S Guo","year":"2019","unstructured":"Guo, S. et al. Immunogenicity of a bivalent protein as a vaccine against Edwardsiella anguillarum and Vibrio vulnificus in Japanese eel (Anguilla japonica). MicrobiologyOpen 8, e00766. https:\/\/doi.org\/10.1002\/mbo3.766 (2019).","journal-title":"MicrobiologyOpen"},{"key":"35298_CR56","doi-asserted-by":"crossref","unstructured":"Austin, B. & McIntosh, D. Natural antibacterial compounds on the surface of rainbow trout, Salmo gairdneri Richardson. Journal of Fish Diseases 11 (1988).","DOI":"10.1111\/j.1365-2761.1988.tb00550.x"},{"key":"35298_CR57","doi-asserted-by":"publisher","first-page":"153","DOI":"10.1016\/0145-305X(94)00063-L","volume":"19","author":"MR Wilson","year":"1995","unstructured":"Wilson, M. R. et al. cDNA sequences and organization of IgM heavy chain genes in two holostean fish. Dev. Comp. Immunol. 19, 153\u2013164. https:\/\/doi.org\/10.1016\/0145-305X(94)00063-L (1995).","journal-title":"Dev. Comp. Immunol."},{"key":"35298_CR58","doi-asserted-by":"publisher","unstructured":"Cuesta, A., Meseguer, J. & Esteban, M. A. Total serum immunoglobulin M levels are affected by immunomodulators in seabream (Sparus aurata L.). Veterinary Immunology and Immunopathology 101, 203\u2013210, https:\/\/doi.org\/10.1016\/j.vetimm.2004.04.021 (2004).","DOI":"10.1016\/j.vetimm.2004.04.021"},{"key":"35298_CR59","doi-asserted-by":"publisher","first-page":"444","DOI":"10.1111\/anu.12176","volume":"21","author":"JS Lee","year":"2015","unstructured":"Lee, J. S. et al. Evaluation and characterization of a novel probiotic Lactobacillus pentosus PL11 isolated from Japanese eel (Anguilla japonica) for its use in aquaculture. Aquac. Nutr. 21, 444\u2013456. https:\/\/doi.org\/10.1111\/anu.12176 (2015).","journal-title":"Aquac. Nutr."},{"key":"35298_CR60","doi-asserted-by":"publisher","unstructured":"Kord, M. I. et al. The Immunostimulatory Effects of Commercial Feed Additives on Growth Performance, Non-specific Immune Response, Antioxidants Assay, and Intestinal Morphometry of Nile tilapia, Oreochromis niloticus. Frontiers in Physiology 12, https:\/\/doi.org\/10.3389\/fphys.2021.627499 (2021).","DOI":"10.3389\/fphys.2021.627499"},{"key":"35298_CR61","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquaculture.2022.738948","volume":"563","author":"HMR Abdel-Latif","year":"2023","unstructured":"Abdel-Latif, H. M. R., Chaklader, M. R., Shukry, M., Ahmed, H. A. & Khallaf, M. A. A multispecies probiotic modulates growth, digestive enzymes, immunity, hepatic antioxidant activity, and disease resistance of Pangasianodon hypophthalmus fingerlings. Aquaculture 563, 738948. https:\/\/doi.org\/10.1016\/j.aquaculture.2022.738948 (2023).","journal-title":"Aquaculture"},{"key":"35298_CR62","doi-asserted-by":"publisher","unstructured":"Amenyogbe, E. et al. The exploitation of probiotics, prebiotics and synbiotics in aquaculture: present study, limitations and future directions. : a review. Aquaculture International 28, 1017\u20131041, https:\/\/doi.org\/10.1007\/s10499-020-00509-0 (2020).","DOI":"10.1007\/s10499-020-00509-0"},{"key":"35298_CR63","doi-asserted-by":"publisher","first-page":"66","DOI":"10.1016\/j.rvsc.2017.01.016","volume":"115","author":"G Banerjee","year":"2017","unstructured":"Banerjee, G. & Ray, A. K. The advancement of probiotics research and its application in fish farming industries. Res. Vet. Sci. 115, 66\u201377. https:\/\/doi.org\/10.1016\/j.rvsc.2017.01.016 (2017).","journal-title":"Res. Vet. Sci."},{"key":"35298_CR64","doi-asserted-by":"publisher","first-page":"2205","DOI":"10.1111\/are.15073","volume":"52","author":"EM Moustafa","year":"2021","unstructured":"Moustafa, E. M. et al. Efficacy of Bacillus probiotic mixture on the immunological responses and histopathological changes of Nile tilapia (Oreochromis niloticus, L) challenged with Streptococcus iniae. Aquac. Res. 52, 2205\u20132219. https:\/\/doi.org\/10.1111\/are.15073 (2021).","journal-title":"Aquac. Res."},{"key":"35298_CR65","doi-asserted-by":"crossref","unstructured":"Wang, X., Zhang, P. & Zhang, X. Probiotics Regulate Gut Microbiota: An Effective Method to Improve Immunity. Molecules 26 (2021).","DOI":"10.3390\/molecules26196076"},{"key":"35298_CR66","doi-asserted-by":"publisher","unstructured":"Zhang, W., Liao, Z., Hu, F., Chen, X. & Huang, X. Protective immune responses of recombinant outer membrane proteins OmpF and OmpK of Aeromonas hydrophila in European eel (Anguilla anguilla). Aquac. Res. 50, 3559\u20133566. https:\/\/doi.org\/10.1111\/are.14311 (2019).","DOI":"10.1111\/are.14311"}],"container-title":["Scientific Reports"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35298-8","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35298-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35298-8.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,2,2]],"date-time":"2026-02-02T10:00:34Z","timestamp":1770026434000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41598-026-35298-8"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,2,2]]},"references-count":66,"journal-issue":{"issue":"1","published-online":{"date-parts":[[2026,12]]}},"alternative-id":["35298"],"URL":"https:\/\/doi.org\/10.1038\/s41598-026-35298-8","relation":{},"ISSN":["2045-2322"],"issn-type":[{"value":"2045-2322","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,2,2]]},"assertion":[{"value":"9 August 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"5 January 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 February 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"order":1,"name":"Ethics","group":{"name":"EthicsHeading","label":"Declarations"}},{"value":"The authors declare no competing interests.","order":2,"name":"Ethics","group":{"name":"EthicsHeading","label":"Competing interests"}},{"value":"The study is reported in accordance with ARRIVE guidelines. It was verified by the Local Experimental Animal Care Committee, Faculty of Veterinary Medicine, Alexandria University, Egypt. Ethical Consent was obtained from the Institutional Animal Care and Use Committee, Alexandria University, on 12th December 2022 with an Ethical Code (ALEXU-IACUC-013\/2022\/12\/-3R\/4P\/186).","order":3,"name":"Ethics","group":{"name":"EthicsHeading","label":"Institutional review board statement"}}],"article-number":"4367"}}