Effects of Carica papaya seed powder as a plant-based aromatase inhibitor on growth performance, feed utilisation, reproductive health, biochemical indices, and intestinal histomorphology in hybrid red tilapia (Oreochromis spp.) https://doi.org/10.12982/VIS.2026.014
Article Sidebar
Main Article Content
Abstract
Plant-derived aromatase inhibitors, such as Carica papaya, are increasingly employed in aquaculture for sex reversal, immunostimulant and growth enhancement. However, their effects on fish physiology, particularly nutrient utilisation, body composition, and serum biochemistry, remain poorly documented, especially in hybrid red tilapia. This study evaluated the effects of dietary inclusions of Carica papaya seed powder (CPSP) at dosage levels 0, 30, 60, and 90 g/kg feed on fish physiological parameters. A completely randomised design with three replicates per treatment was used, comprising a 60-day treatment phase and a 90-day recovery phase. During the treatment, higher CPSP (60–90 g/kg) significantly increased weight gain and protein efficiency ratio (PER) (p < 0.05) but impaired feed conversion ratio (FCR) (p < 0.05), with no significant differences in specific growth rate (SGR) (p > 0.05). At the recovery phase, fish previously fed 60–90 g/kg CPSP exhibited significantly improved SGR, FCR, and PER (p < 0.05). Lipid, ash, and moisture content were significantly elevated in fish from the 90 g/kg group (p< 0.05), while carbohydrate markedly declined (p > 0.05). Reproductive metrics showed that 90 g/kg CPSP significantly increased 11-ketotestosterone (p< 0.05) with suppression in oestradiol and GSI (p < 0.05). Serum biochemistry indicated lower aspartate aminotransferase (AST) and alanine aminotransferase (ALT) but higher alkaline phosphatase (ALP), albumin, and globulin in fish from 90 g/kg CPSP groups (p < 0.05). Cholesterol, glucose, and total protein levels markedly improved (p < 0.05). For improved fish health, 60 g/kg CPSP is recommended. Future research should prioritise scalable methods to commercialise papaya seeds as a cost-effective feed additive and sex-reversal agent in aquaculture.
Article Details
This work is licensed under a Creative Commons Attribution 4.0 International License.
Publishing an article with open access in Veterinary Integrative Sciences leaves the copyright with the author. The article is published under the Creative Commons Attribution License 4.0 (CC-BY 4.0), which allows users to read, copy, distribute and make derivative works from the material, as long as the author of the original work is cited.
References
Abd El-Naby, A.S., El Asely, A.M., Hussein, M.N., Khattaby, A E.R., Sabry, E.A., Abdelsalam, M., Samir, F., 2024. Effects of dietary fermented Saccharomyces cerevisiae extract (Hilyses) supplementation on growth, hematology, immunity, antioxidants, and intestinal health in Nile tilapia. Sci. Rep. 14(1), 1–14.
Abo-Taleb, H.A., Mabrouk, M.M., El-Tabakh, M.A., Abdelzaher, O.F., Nabeeh, A., Elokaby, M.A., Ashour, M., Sallam, A.E., Mansour, A.T., El-feky, M.M., 2024. The potential use of daphnia meal as substitute for fishmeal in diets of hybrid red tilapia affects growth performance, activities of digestive enzymes, antioxidant, immune status and intestinal histological parameters. Anim. Physiol. Anim. Nutr. 108(3), 752-763
Abu-Elala, N.M., Khattab, M.S., AbuBakr, H.O., Helmy,S., Hesham, A., Younis, N.A., Dawood, M.A.O., El Basuini, M.F., 2023. Neem leaf powder (Azadirachta indica) mitigates oxidative stress and pathological alterations triggered by lead toxicity in Nile tilapia (Oreochromis niloticus). Sci Rep. 13(1), 1–16.
Adham, K.G., Ibrahim, H.M., Hamed, S.S., Saleh, R.A., 2002. Blood chemistry of the Nile tilapia, Oreochromis niloticus (Linnaeus, 1757) under the impact of water pollution. Aquat. Ecol. 36(4), 549–557.
Adi, G., Uedeme-Naa,B., Okon, M.A., 2020. Comparative study of testis histology and haematology of Clarias gariepinus exposed to phytochemicals of Moringa oleifera and Carica papaya leaf powder. Int. J. Fish. Aquat. Stud. 8(2), 301-306.
Afonso, L.O.B., Wassermann, G.J., De Oliveira, R.T. 2001. Sex reversal in Nile tilapia (Oreochromis niloticus) using a nonsteroidal aromatase inhibitor. J. Exp. Zool. 290(2), 177–181.
AOAC, 2005. Official methods of analysis of Association of official agriculture chemists, 17th edition (2nd Version). AOAC, Maryland.
Asghar, M., Khan, N., Fatima, M., Arslan, M., Davies, S.J., Ul Haque, N., 2024. Feasibility of replacing fish oil with sunflower oil on the growth, body composition, fatty acid profile, antioxidant activity, stress response, and blood biomarkers of Labeo rohita. PLOS ONE. 19(3), e0299195.
Ashraf, A., Sabu, S., Sasidharan, A., Sunooj, K.V., 2024. Natural feed supplements from crustacean processing side streams for improved growth of finfishes and crustaceans. J. Anim. Physiol. Anim. Nutr. 2024, 1-26.
Ayodipupo Babalola, B., Ifeolu Akinwande, A., Otunba, A.A., Ebenezer Adebami, G., Babalola, O., Nwuofo, C., 2024. Therapeutic benefits of Carica papaya: A review on its pharmacological activities and characterization of papain. Arab. J. Chem. 17(1), 105-369.
Ayotunde, E.O., Offem, B.O., Okey, I.B., Ikpi, G.U., Ochang, S.N., Agbam, N.E., Omini, D.E., 2010. Toxicity of pawpaw (Carica papaya) seed powder to sharptooth catfish Clarias gariepinus fingerlings and effects on haematological parameters. Int. J. Fish. Aquat. 2(3), 71–78.
Azarm, H.M., Lee, S.M., 2014. Effects of partial substitution of dietary fish meal by fermented soybean meal on growth performance, amino acid and biochemical parameters of juvenile black sea bream Acanthopagrus schlegeli. Aquac. Res. 45(6), 994–1003.
Azizah, L.S., Kismiyati, Fasya, A.H., 2019. Effectiveness of pepaya leaf extract (Carica Papaya L.) to control ectoparasite argulus on common carp (Cyprinus Carpio). IOP Conf. Ser. Earth Environ. Sci. 236(1), 012-106.
Badran, A.A., Magouz, F.I., Zaineldin, A.I., Abdo, S.E., Amer, A.A., Gewaily, M.S., Dawood, M.A.O., 2024. Using a blend of oilseed meals in the diets of Nile tilapia (Oreochromis niloticus): effects on the growth performance, feed utilization, intestinal health, growth, and metabolic-related genes. BMC Vet. Res. 20(1), 1–14.
Basavaraja, N., Raghavendra, C.H., 2017. Hormonal sex reversal in red tilapia (Oreochromis niloticus and Oreochromis mossambicus) and inheritance of body colour in O. mossambicus and red tilapia: implications for commercial farming. Aquac. Int. 25(3), 1317–1331.
Benli, A.C.K., Yildiz, H.Y., 2004. Blood parameters in Nile tilapia (Oreochromis niloticus L.) spontaneously infected with Edwardsiella tarda. Aquac. Res. 35(14), 1388–1390.
Chen, L., Wang, L., Cheng, Q., Tu, Y.X., Yang, Z., Li, R.Z., Luo, Z.H., Chen, Z.X., 2020. Anti-masculinization induced by aromatase inhibitors in adult female zebrafish. BMC Genomics. 21(1), 1–10.
Chen, Q., Zhao, H., Huang, Y., Cao, J., Wang, G., Sun, Y., Li, Y., 2016. Effects of dietary arginine levels on growth performance, body composition, serum biochemical indices and resistance ability against ammonia-nitrogen stress in juvenile yellow catfish (Pelteobagrus fulvidraco). Anim. Nutr. 2(3), 204.
Chuhila, Y.J., Mwita, C.J., Chibwana, F.D., 2024. Aquaculture spillage: a gateway to establishment and colonization of non-indigenous tilapias (Pisces, Cichlidae) in the Pangani Catchment, northern Tanzania. Hydrobiologia. 851(1), 67–86.
Correia, D., Escarcega-Miranda, B., Barreto-Curiel, F., Mata-Sotres, J., Del-Rio-Zaragoza, O., Viana, M.T., Rombenso, A.N., Correia, D., Escarcega-Miranda, B., Barreto-Curiel, F., Mata-Sotres, J., Del-Rio-Zaragoza, O., Viana, M.T., Rombenso, A.N., 2019. Growth performance and body composition of hybrid red tilapia (Oreochromis mossambicus × O. aureus) fed with different protein levels raised in saltwater. Lat. Am. J. Aquat. Res. 47(5), 853–859.
Dawood, M.A.O., Koshio, S., 2020. Application of fermentation strategy in aquafeed for sustainable aquaculture. Rev. Aquac. 12(2), 987–1002.
Dinh, Q.M., Nguyen, T.H.D., 2022. Procedure for performing a fixed microscopic specimen of the gonads of fish. Vet. Integr. Sci. 20(3), 645–657.
Doering, J.A., Villeneuve, D.L., Tilton, C.B., Kittelson, A.R., Blackwell, B.R., Kahl, M.D., Jensen, K.M., Poole, S.T., Cavallin, J.E., Cole, A.R., Dean, K.N., LaLone, C.A., Ankley, G.T., 2021. Assessing effects of aromatase inhibition on fishes with group-synchronous oocyte development using western mosquitofish (Gambusia affinis) as a model. Aquat. Toxicol. 232, 105741.
Eissa, E.S.H., Awlya, O.F., Abusudah, W.F., Qadhi, A.H., Abd El-Aziz, Y.M., Rinthong, P.O., Abdelnour, S.A., Munir, M.B., Eissa, M.E.H., Sakr, S.E.S., 2024. Curcumin’s effects on growth indices, histological scores, blood metabolites, redox state, immunity, and antioxidant-related genes of red tilapia (Oreochromis sp.). Aquac. Int. 1–16.
Eissa, E.S.H., El-Sayed, A.F.M., Ghanem, S.F., Dighiesh, H.S., Abd Elnabi, H.E., Hendam, B.M., Elleithy, A.A., Eissa, M.E.H.,Abd El-Aziz, Y.M., 2024. Dietary mannan-oligosaccharides enhance hematological and biochemical parameters, reproductive physiology, and gene expression of hybrid red tilapia (Oreochromis niloticus x O. mossambicus). Aquac. 581, 740453.
Ekinadose, O., Wokeh, O.K., Orose, E., 2021. Histological changes in the gills, livers and muscles of Oreochromis niloticus fed with steriod, natural hormone and pawpaw seed based diets. Ege J. Fish. Aquat. Sci. 38(4), 487–497.
El-Sayed Ali, T., El-Sayed, A.M., Eissa, MA.R.,Hanafi, H.M., 2018. Effects of dietary Biogen and sodium butyrate on hematological parameters, immune response, and histological characteristics of Nile tilapia (Oreochromis niloticus) fingerlings. Aquac. Int. 26(1), 139–150.
Enyidi, U., Nduh-Nduh, A., 2016. Application of phytogenics as first feed of larval African catfish Clarias gariepinus. J. Adv. Biol. Biotechnol. 5(3), 1–10.
Fan, S., Shi, H.J., Peng, Y.X., Huang, Y.Q., Jiang, M.Y., Li, G.L., Wang, D.S., Jiang, D.N., 2023. Dietary aromatase inhibitor treatment converts XY gsdf homozygous mutants to sub-fertile male in Nile tilapia (Oreochromis niloticus). Aquac. 569, 739381.
FAO, 2024. The State of World Fisheries and Aquaculture 2024 – Blue Transformation in action. Available online: https://doi.org/10.4060/cd0683en.
Farrag, F., Khalil, F., Mehrim, A., Refaey, M., 2013. Pawpaw (Carica papaya) seeds powder in Nile Tilapia (Oreochromis niloticus) diet 1-growth performance, survival, feed utilization, carcass composition of fry and fingerlings. J. Anim. Poul. Produc. 4(6), 363–379.
Fazio, F., 2019. Fish hematology analysis as an important tool of aquaculture: a review. Aquac. 500, 237–242.
Fazio, F., Costa, A., Capparucci, F., Costa, G., Parrino, V., Arfuso, F., 2024. Automated hematological approach and protein electrophoretic pattern in tilapia (Oreochromis niloticus): an innovative and experimental model for aquaculture. Animals. 14(3), 392.
Fazio, F., Filiciotto, F., Marafioti, S., Di Stefano, V., Assenza, A., Placenti, F., Buscaino, G., Piccione, G., Mazzola, S., 2012. Automatic analysis to assess haematological parameters in farmed gilthead sea bream (Sparus aurata Linnaeus, 1758). Mar. Freshw. Behav. Physiol. 45(1), 63–73.
Gao, Z.X., Wang, H.P., Wallat, G., Yao, H., Rapp, D., O’Bryant, P., MacDonald, R., Wang, W.M., 2010. Effects of a nonsteroidal aromatase inhibitor on gonadal differentiation of bluegill sunfish Lepomis macrochirus. Aquac. Res. 41(9), 1282–1289.
Ghosal, I., Chakraborty, S.B., 2020. Production of monosex all-male Nile tilapia using ethanol extract of Tribulus Terrestris seeds. Proc. Zool. Soc. 73(2), 188–191.
Ghosal, I., Mukherjee, D., Chakraborty, S.B., 2021. The effects of four plant extracts on growth, sex reversal, immunological and haemato-biochemical parameters in Nile tilapia, Oreochmomis niloticus (Linnaeus, 1758). Aquac. Res. 52(2), 559–576.
Gomes, E.F., Rema, P., Kaushik, S.J., 1995. Replacement of fish meal by plant proteins in the diet of rainbow trout (Oncorhynchus mykiss): digestibility and growth performance. Aquac. 130(2–3), 177–186.
Gule, T.T., Geremew, A., 2022. Dietary strategies for better utilization of aquafeeds in tilapia farming. Aquac. Nutr. 2022, 463307.
Haihambo, W., Gabriel, N.N., 2022. Sodium bicarbonate and acetic acid: an effective anaesthetic for three spotted tilapia, Oreochromis andersonii fingerlings. J. Appl. Anim. Ethics. Res. 4(2), 159–169.
Hamid, N.K.A., Somdare, P.O., Md Harashid, K.A., Othman, N.A., Kari, Z.A., Wei, L.S., Dawood, M.A.O., 2022. Effect of papaya (Carica papaya) leaf extract as dietary growth promoter supplement in red hybrid tilapia (Oreochromis mossambicus × Oreochromis niloticus) diet. Saudi J. Biol. Sci. 29(5), 3911–3917.
Hanan, M.Y., Amatul-Samahah, M.A., Jaapar, M.Z., Mohamad, S.N., 2022. The effects of field cricket (Gryllus bimaculatus) meal substitution on growth performance and feed utilization of hybrid red tilapia (Oreochromis spp.). Appl. Food. Res. 2(1), 100070.
Harikrishnan, M., Syanya, F.J., Khanna, A.R.N., Mumina, P., Mathia, W.M., 2024. Ecological implications of unintentional aquaculture escapees: an overview of risks, remediation strategies and knowledge gaps in the aquaculture sector of India and riparian East African countries. Mar. Fish. Sci. 37(4), 633-666.
Hastuti, S., Subandiyono, S., Hastuti, S., 2020. Aminotransferase, hematological indices and growth of tilapia (Oreochromis niloticus) reared in various stocking densities in aquaponic systems. AACL Bioflux. 13(2), 813-824.
Heluy, G.M., Ramos, L.R.V., Pedrosa, V.F., Sarturi, C., Figueiredo, P.G.P., Vidal, L.G.P., França, I.de.F., Pereira, M.M., 2020. Oregano (Origanum vulgare) essential oil as an additive in diets for Nile tilapia (Oreochromis niloticus) fingerlings reared in salinized water. Aquac. Res. 51(8), 3237–3243.
Hoseinifar, S.H., Mirvaghefi, A., Merrifield, D.L., Amiri, B.M., Yelghi, S., Bastami, K.D. 2011. The study of some haematological and serum biochemical parameters of juvenile beluga (Huso huso) fed oligofructose. Fish. Physiol. Biochem. 37(1), 91–96.
Hosseinpour Zelaty, A., Sayed Hassani, M.H., Sohrabi, T., Bagherzadeh Lakani, F., Mohseni, M., Pajand, Z., 2024. Effect of a commercial probiotic (Supersets® and Varna®) on growth performance, hematological indices, and immunological parameters in sturgeon hybrid bester (Huso huso ♂× Acipenser ruthenus ♀) fingerling. Iran. J. Fish. Sci. 23(2), 293–306.
Iipinge, L.N., 2019. Effect of pawpaw (Carica Papaya) seed meal on growth performance, feed utilization, survival and masculinization of sexualy undifferentiated three spotted Tilapia (Oreochromis Andersonii) fry. Available online: https://repository.unam.edu.na/handle.
Inokuchi, M., Hiroi, J., Kaneko, T., 2022. Why can mozambique tilapia acclimate to both freshwater and seawater? insights from the plasticity of ionocyte functions in the euryhaline teleost. Front. Physiol. 13, 914277.
Islam, R., Hossain, M.B., Islam, M.N., Islam, M.M., Islam, M.T., 2020. Nutrient composition of small indigenous fish species (SIS) from homestead ponds of Noakhali coast, Bangladesh. Egypt. J. Aquat. Biol. Fish. 24(7-Special issue), 943–954.
Jamal, R., Asad, F., Naz, S., Hussain, S.M., 2024. Comparative assessment of natural and synthetic reproductive inhibitors in Oreochromis niloticus. Int. J. Agric. Biol. Eng. 17(5), 284–292.
Jimoh, W.A., Ayeloja, A.A., Ojo, O.A., Ayodele, C.T., Alabi, A.E., Obinnakwelu, G.A., 2023. Effect of dietary cinnamon supplementation on the growth in length, haematology, serum biochemistry, intestinal microbiota and enzymes and histomorphological changes of the intestine, liver and the kidney of Heteroclarias (Clarias gariepinus ♀ × Heterobranchus bidorsalis♂). Vet. Integr. Sci. 21(3), 973–997.
Kamil, M.A., Abu-Bakar, L., Reduan, M.F.H., Kamaruzaman, I.N.A., Mahamud, S.N.A., Azmi, A.F.M., Rahman, M.S.A., Nordin, M.L., Shaari, R., Shean, S.C.S., 2023. The Use of medicinal plants in avian colibacillosis management: a review. Vet. Integr. Sci. 21(2), 507–522.
Khalil, F.F., Farrag, F.H., Mehrim, A.I., Refaey, M.M.A., 2014. Pawpaw (Carica papaya) seeds powder in Nile tilapia (Oreochromis niloticus) diets: 2 Liver status, sexual hormones and histological structure of the gonads. Egypt. J. Aquat. Biol. Fish. 18(1), 97–113.
Kumar, N., Antony Jesu Prabhu, P., Pal, A.K., Remya, S., Aklakur, M., Rana, R.S., Gupta, S., Raman, R.P., Jadhao, S.B., 2011. Anti-oxidative and immuno-hematological status of Tilapia (Oreochromis mossambicus) during acute toxicity test of endosulfan. Pestic. Biochem. Physiol. 99(1), 45–52.
Lal, J., Vaishnav, A., Singh, S.K., Biswas, P., Mehta, N.K., Meena, D.K., Waikhom, G., 2024. The potential of aromatase inhibitors in fish masculinization: A comprehensive review of applications, mechanisms and future perspectives. Int. Aquat. Res. 16(2), 113-142.
Lal, J., Biswas, P., Singh, S.K., Debbarma, R., Deb, S., Yadav, N.K.,Patel, A.B., 2023. Effects of dietary aromatase inhibitors on masculinization of rosy barb (Pethia conchonius): Evidence from growth, coloration and gonado-physiological changes. PLOS ONE. 18(11), e0287934.
Lohiya, N.K., Goyal, R.B., Jayaprakash, D., Ansari, A.S., Sharma, S., 1994 Antifertility effects of aqueous extract of Carica papaya seeds in male rats. Planta. Medica. 60(5), 400–404.
Loureiro Paschoalini, A., Boaventura, T.P., Costa Dos Santos, F.A., Bazzoli, N., Luz, R.K., Favero, G.C., 2024. Morphological and histochemical changes in the intestine of adult catfish (Lophiosilurus alexandri) subjected to chronic fasting and refeeding. J. Appl. Aquac. 2024, 1–15.
Magblenou, L.D., Ly, M.A., Kantoussan, J., Faye, R., Kane, Y., 2019. Effects of varying dietary levels of Carica papaya seed meal powder (PSM) on growth and histology of gonads in Oreochromis niloticus larvae. Int. J. Biol. Chem. Sci. 13(2), 624–633.
Majumdar, R., Benzie, J., Delamare-Deboutteville, J., Yossa, R., Khor, L., Verner-Jeffreys, D., Chadag, V., Mills, D.J., Smallhorn-West, P., Eriksson, H., Tigchelaar, M., Allison, E., Hossain, P.R., Tilley, A., Rossignoli, C., Adam, R., Gichohi, W., Siamudaala, V., Komugisha, B., Fleming, A., 2024. WorldFish Annual Report 2023. WorldFish, Malaysia, pp. 1-25.
Makkar, H.P.S., Francis, G., Becker, K., 2007. Bioactivity of phytochemicals in some lesser-known plants and their effects and potential applications in livestock and aquaculture production systems. Animal. 1(9), 1371–1391.
Mansour, A.T., Hamed, H.S., El-Beltagi, H.S., Mohamed, W.F., 2022. modulatory effect of papaya extract against chlorpyrifos-induced oxidative stress, immune suppression, endocrine disruption, and dna damage in female Clarias gariepinus. Int. J. Envi. Res. Pub. Heal. 19(8), 4640.
Marshall, J.A., Bielic, P.E., 1996. Periodicity of reproductive behaviour by the blue tilapia, Oreochromis aureus. Env. Biol. Fish. 47, 411–414.
Ng, W.K., Hanim, R., 2007. Performance of genetically improved Nile tilapia compared with red hybrid tilapia fed diets containing two protein levels. Aquac. Res. 38(9), 965–972.
Ngozi, G., 2016. Sex reversal of Nile Tilapia (Oreochromis Niloticus) using 17α-methyl-testosterone, wet testes from selected animals and plant extract. Int. J. Life Sci. Res. 4(3), 23–29.
Niyibizi, L., Vidakovic, A., Norman Haldén, A., Rukera Tabaro, S., Lundh, T., 2022. Aquaculture and aquafeed in Rwanda: current status and perspectives. J. Appl. Aquac. 35(98), 1-22.
Nwachi, O.F., Esa, Y.B., Christianus, A., Rahim, A.A., Kamarudin, M.S., 2020. Patterns of colour inheritance from crossbreeding between Red hybrid tilapia (Oreochromis sp.) and GIFT tilapia (Oreochromis niloticus). J. Environ. Biol. 41, 1289–1294.
Okomoda, V.T., 2017. Haematology and gonad histology of Oreochromis niloticus (Linnaeus, 1758) fed Carica papaya seed meal. Braz. J. Aquat. Sci. Tech. 21(1), 8–15.
Omeje, V.O., 2016. Effect of Pawpaw (Carica papaya) seed meal on the reproductive, endocrine and immune system of Mozambique tilapia (Oreochromis mossambicus) (PhD thesis). Stellenbosch university, pp. 1-25.
Omeje, V.O., Lambrechts, H., Brink, D., 2018. Effect of pawpaw (Carica Papaya) seed meal on sex determination, growth, and survival, of oreochromis mossambicus fry. Isr. J. Aquac. - Bamidgeh. 59(4), 1-12.
Omeje, V.O., Lambrechts, H., Brink, D., 2019. Hormonal profile and reproductive parameters of pre-vitellogenic mozambique tilapia (Oreochromis mossambicus) on pawpaw (carica papaya) seed meal. J. Agric. Sci. 11(2), 524.
Omeje, V.O., Lambrechts, H., Brink, D., 2020. Use of pawpaw (Carica papaya) seed in tilapia sex reversal. Revi. Agricul. Sci. 8, 230–242.
Owolabi, O.D., Abdulkareem, S.I., 2021. Carica papaya and Mangifera indica modulate haematological, biochemical and histological alterations in atrazine-intoxicated fish, Clarias gariepinus (Burchell 1822). J. Basic. Appl. Zool. 82(1), 1–18.
Palladino, A., De Felice, E., Attanasio, C., Barone, C.M.A., Crasto, A., D’Angelo, L., Giaquinto, D., Lambiase, C., Scocco, P., Serrapica, F., Maruccio, L., 2023. A morphological and ultrastructural study of the anterior digestive tract of adult Nile Tilapia Oreochromis niloticus. Animals. 13(3), 420.
Patel, H.V, Rani, K.S., Patani, P., 2024. A review on role of herbs: new perspectives and potential health benefits of citrullus lanatus & carica papaya seeds. J. Popul. Ther. Clin. Pharmacol. 31(1), 2189–2122.
Pongthana, N., Nguyen, N.H., Ponzoni, R.W., 2010. Comparative performance of four red tilapia strains and their crosses in fresh- and saline water environments. Aquac. 308, 109–114.
Ponzoni, R.W., Nguyen, N.H., Khaw, H.L., Hamzah, A., Bakar, K.R.A., Yee, H.Y., 2011. Genetic improvement of Nile tilapia (Oreochromis niloticus) with special reference to the work conducted by the World Fish Center with the GIFT strain. Rev. Aquac. 3(1), 27–41.
Radwan, M., Darweesh, K.F., Ghanem, S.F., Abdelhadi, Y., Kareem, Z.H., Christianus, A., Karim, M., Waheed, R.M., El-Sharkawy, M.A., 2023. Regulatory roles of Pawpaw (Carica papaya) seed extract on growth performance, sexual maturity, and health status with resistance against bacteria and parasites in Nile tilapia (Oreochromis niloticus). Aquac. Int. 31(5), 2475–2493.
Sadiq Bukhari, A., Syed Mohamed, H.E., Broos, K.V., Stalin, A., Singhal, R.K., Venubabu, P. 2012. Histological variations in liver of freshwater fish Oreochromis mossambicus exposed to 60Co gamma irradiation. J. Environ. Radioact. 113, 57–62.
Sayed, A.E.D, Moneeb, R.H., 2015. Hematological and biochemical characters of monosex tilapia (Oreochromis niloticus, Linnaeus, 1758) cultivated using methyltestosterone. J. Basic. Appl. Zool. 72, 36–42.
Séralini, G.E., Moslemi, S., 2001. Aromatase inhibitors: past, present and future. Mol. Cell. Endocrinol. 178(1–2), 117–131.
Singh, R., Singh, A., Tripathi, M., 2012. Effect of a non steroidal tamoxifen on the gonad and sex differentiation in Nile tilapia, Oreochromis niloticus. J. Environ. Biol. 33, 799–803.
Somdare, P.O., Hamid, N.K.A., Sul’Ain, M.D., 2023. Effect of different forms of Carica papaya leaf processing techniques on growth, body indices and survival rate of red hybrid tilapia, Oreochromis mossambicus × Oreochromis niloticus. IOP Conf. Ser. Earth. Environ. Sci. 1221(1), 012031.
Suman, U., Waikhom, G., Singh, S.K., Mangang, Y.A., Debbarma, R., Biswas, P., Das, P., Singh, S.B., 2024. Evidences of gonadal impairment and growth acceleration in barbonymus gonionotus fed papaw seed: sustainable green alternatives for energy portioning. J. Appl. Ichthyol. 2024(1), 5663091.
Syanya, F.J., Litabas, J.A., Mathia, W.M., Ntakirutimana, R., 2023. Nutritional fish diseases in aquaculture: a human health hazard or mythical theory: an overview. European J. Nutr. Food. Saf. 15(8), 41–58.
Syanya, F.J., Mahadevan, H., Khanna, A.R.N., 2025. The effects of a non-steroid aromatase inhibitor on hybrid red tilapia masculinization, growth, reproductive hormone profile, and economic efficiency in aquaculture. Aquac. Int. 33, 160.
Syanya, F.J., Mathia, W.M., Harikrishnan, M., 2023. Current status and trend on the adoption of fish feed additives for sustainable tilapia aquaculture production: a review. Asian. J. Fish. Aquat. Res. 22(3), 10–25.
Syanya, F.J., Khanna, A.N., Mahadevan, H., Mathia, W.M., Mumina, P., Mwangi, P.M., 2025. Dietary effects of saccharomyces cerevisiae and bifidobacterium longum probiotics cocktail on the growth and health of mozambique tilapia (oreochromis mossambicus) in aquaculture. Vet. Integr. Sci. 24(1), 1-21.
Syanya, F.J., Mahadevan, H., Khanna, A.R.N., Mathia, W.M., Mumina, P., 2025. Role of a non-steroidal aromatase inhibitor on growth performance, body composition, reproductive and serum biochemical indices, and intestinal histomorphology in hybrid red tilapia (oreochromis spp.). Fish. Physiol. Biochem. 51(1), 45.
Syed, R., Masood, Z., Ul Hassan, H., Khan, W., Mushtaq, S., Ali, A., Gul, Y., Jafari, H., Habib, A., Ishaq Ali Shah, M., Gabol, K., Gul, H., Ullah, A., 2022. Growth performance, haematological assessment and chemical composition of Nile tilapia, Oreochromis niloticus (Linnaeus, 1758) fed different levels of Aloe vera extract as feed additives in a closed aquaculture system. Saudi J. Biol. Sci. 29(1), 296–303.
Uchida, D., Yamashita, M., Kitano, T., Iguchi, T., 2004. An aromatase inhibitor or high water temperature induce oocyte apoptosis and depletion of P450 aromatase activity in the gonads of genetic female zebrafish during sex-reversal. Comp. Biochem. Physiol. A Mol. Integr. Physiol. 137(1), 11–20.
Vijayaram, S., Ringø, E., Ghafarifarsani, H., Hoseinifar, S.H., Ahani, S., Chou, C.C., 2024. Use of algae in aquaculture: a review. Fishes. 9(2), 63.
Watanabe, W.O., Ernst, D.H., Olla, B.L., Wicklund, R.I., 1989. Aquaculture of red tilapia Oreochromis sp. in marine environments: State of the art. Available online: https://archimer.ifremer.fr/doc/00000/1493/1127.pdf.
Watanabe, W.O., Losordo, T.M., Fitzsimmons, K., Hanley, F., 2002. Tilapia production systems in the americas: technological advances, trends, and challenges. Rev. Fish. Sci. 10(3–4), 465–498.
Wokeh, O.K., Orose, E., Wokeh, O.K., Orose, E., 2021. Use of dietary phytochemicals as control for excessive breeding in Nile Tilapia (Oreochromis niloticus): a review. GSC Biol. Pharm. Sci. 17(2), 152–159.
Yehia, Y.A., Greish, S.S., Kamel, M.S., Abd EL-Rahman, A.S. M., Ahmed, Y.M., Gaber, M., 2024. Potential role of Azolla pinnata on abamectin induced hepatorenal toxicity and oxidative stress in Nile tilapia. J. Appl. Plant. Prot. 13(1), 29–38.
Yostawonkul, J., Kitiyodom, S., Supchukun, K., Thumrongsiri, N., Saengkrit, N., Pinpimai, K., Hajitou, A., Thompson, K.D., Rattanapinyopituk, K., Maita, M., Kamble, M.T., Yata, T., Pirarat, N., 2023. Masculinization of Red Tilapia (Oreochromis spp.) using 17α-Methyltestosterone-Loaded Alkyl Polyglucosides Integrated into Nanostructured Lipid Carriers. Animals. 13(8) 1-25.
Zaki, F.M., Said, M.M., Tahoun, A.A., Amer, M., 2021. Evaluation of different sex reversal treatments in red tilapia hybrid. Egypt. J. Aquat. Biol. Fish. 25(1), 279-292.
Zuluaga-Hernández, C.D., Hincapié, C.A., Osorio, M., 2023. Non-conventional ingredients for Tilapia (Oreochromis spp.) feed: a systematic review. Fishes. 8(11), 556.
