The complementary effect of melatonin in accelerating ovulation and spawning of the Javaen barb Systomus rubripinnis (Valenciennes 1842)Abstract
The Javaen barb Systomus rubripinnis is an endemic fish not domesticated in Indonesia. This study aims to evaluate the effectiveness of gonadotropin and melatonin hormones in inducing ovulation of female Javaen barb. A total of 12 female fish (BW: 142.12 ± 18.08 g; egg diameter 1.0-1.2 mm) were selected to be injected with a combination of different hormones. The treatments were ovaprimTM at a dose of 0.6 mL/kg without melatonin (L0.6M0), ovaprimTM at 0.3 mL/kg with 0.25 mg/kg melatonin (L0.3M0.25), 500 IU/kg of human chorionic gonadotropin with 0.25 mg/kg melatonin (H500M0.25), and 0.25 mg/kg melatonin (M0.25). The L0.6M0 and L0.3M0.25 treatments were injected twice. The first injection was 40% of the total dose, while the rest (60%) was injected 6 hours after the first injection. Melatonin injection was carried out at the same time as the first injection. In the H500M0.25 treatment, melatonin injection was carried out 24 hours after HCG injection. Fish injected with ovaprimTM with and without melatonin had the fastest latency period, and ovulation occurred in all fish. The H500M0.25 treatment had an ovulation rate of 66.7%, while those injected with only melatonin (M0.25) did not ovulate. The number of ovulated eggs, fertilization and hatching rate from ovaprimTM injected broodstock were higher than those of HCG. In contrast, the larvae’s survival rate, body weight, and length were similar. In conclusion, ovaprimTM is practical in inducing ovulation of Javaen barb, and melatonin has a complementary effect on Javaen barb ovulation.
Keywords:
gonadotropin; hormone; induction; reproduction; survival rate
Resumo
O barbo de Java Systomus rubripinnis é um peixe endêmico não domesticado na Indonésia. Este estudo teve como objetivo avaliar a eficácia dos hormônios gonadotrofina e melatonina na indução da ovulação de fêmeas de barbo de Java. Um total de 12 fêmeas (PC: 142,12 ± 18,08 g; diâmetro do ovo 1,0-1,2 mm) foi selecionado para receber uma combinação de diferentes hormônios. Os tratamentos foram ovaprimTM na dose de 0,6 mL/kg sem melatonina (L0,6M0), ovaprimTM a 0,3 mL/kg com 0,25 mg/kg de melatonina (L0,3M0,25), 500 UI/kg de gonadotrofina coriônica humana com 0,25 mg/kg de melatonina (H500M0,25) e 0,25 mg/kg de melatonina (M0,25). Os tratamentos L0,6M0 e L0,3M0,25 foram injetados duas vezes. A primeira injeção foi de 40% da dose total, enquanto o restante (60%) foi injetado 6 horas após a primeira injeção. A injeção de melatonina foi realizada ao mesmo tempo que a primeira injeção. No tratamento H500M0,25, a injeção de melatonina foi realizada 24 horas após a injeção de HCG. Peixes injetados com ovaprimTM com e sem melatonina apresentaram o período de latência mais rápido, e a ovulação ocorreu em todos os peixes. O tratamento H500M0,25 apresentou uma taxa de ovulação de 66,7%, enquanto aqueles injetados apenas com melatonina (M0,25) não ovularam. O número de ovos ovulados, a fertilização e a taxa de eclosão dos reprodutores injetados com ovaprimTM foram maiores do que os do HCG. Em contraste, a taxa de sobrevivência, o peso corporal e o comprimento das larvas foram semelhantes. Em conclusão, o ovaprimTM é prático na indução da ovulação do barbo de Java e a melatonina tem um efeito complementar na ovulação do barbo de Java.
Palavras-chave:
gonadotrofina; hormônio; indução; reprodução; taxa de sobrevivência
1. Introduction
The Javaen barb Systomus rubripinnis (Valenciennes, 1842) (Cyprinidae) is fish species which that originally occur in Southeastern Asia, however, it is widely introduced in several locations (Fricke et al., 2025). It could be developed as a cultured fish due to its economic value (2-4 USD/Kg). Apart from being an edible fish species, Javaen barb fish have the potential to be developed as ornamental fish because of their unique color (Dewi et al., 2021). Meeting the consumption needs of Javaen barb fish still relies on the natural catchment. Catching activities carried out continuously will decrease the number of wild populations. Nowadays, it is challenging to find this fish spesies. This condition could impact the loss of one of the biological resources inland. In conserving natural resources and increasing Javaen barb populations in nature, hatchery activities are needed to support the availability of seeds throughout the year and sustainable aquaculture activities (Passini et al., 2019; Teletchea and Fontaine, 2014; Lorenzen et al., 2012).
Ovulation is one of the reproductive processes that occur in fish breeding activities. Ovulation is the last step in the fish reproductive cycle, where cycle regulation starts from the brain–the hypothalamus–pituitary–gonads (Zamri et al., 2022; Muñoz-Cueto et al., 2020; Blanco, 2020; Sipos et al., 2019). The addition of exogenous hormones is expected to increase the availability of reproductive hormones in the fish's body so that during the process of gonadal maturation and environmental influences, it can cause the brain to release GnRH, which causes the pituitary to produce gonadotropin hormones (LH=luteinizing hormone) (Zamri et al., 2022; Saleh et al., 2020; Ali et al., 2015). The LH hormone can increase egg development in the final maturation process of fish gonads and encourage fish to ovulate (Nguyen et al., 2019; Mosha, 2018). Some of the gonadotropin hormone products that are often used in fish breeding in the world to accelerate the process of final gonadal maturation and ovulation are LHRHa+AD (luteinizing hormone-releasing hormone analog) and HCG (human chorionic gonadotropin), and their combinations (Zamri et al., 2022; Zohar, 2021; Radona et al., 2020; Nosrati et al., 2019; Wahbi et al., 2017; Zadmajid, 2016).
LHRHa is a hormone from the protein group produced by the hypothalamus, consisting of 10 amino acids (decapeptides). It has a relatively short half-life in blood circulation and is easily decomposed in the body. In commercial products, ovaprimTM consists of 20 µg salmon gonadotropin hormone-releasing hormone (SGnRH) and 10 mg of domperidone (Sipos et al., 2019; Acharjee et al., 2017; Zadmajid, 2016; DiMaggio et al., 2014). HCG is a chorionic gonadotropin that contains lots of LH, which can potentially increase egg development in the final gonad maturation process of fish. HCG was first discovered in 1927 in the blood and urine of pregnant women (Elakkanai et al., 2015). Another hormone product that can be used to increase reproductive performance is melatonin. Melatonin is a hormone produced by the pineal gland. It functions as an MPF (maturation-promoting factor) inducer. It also increases MIH (maturation-inducing hormone) functionality which can stimulate the final process of oocyte maturation (Mondal et al., 2019). Several studies of ovulation induction in native fish using the LHRHa+AD and HCG hormones have been carried out, including mahseer the Neolissochilus soro (Valenciennes 1842) (Cyprinidae) (Farastuti et al., 2014), the tinfoil barb Barbonymus schwanenfeldii (Bleeker 1854) (Cyprinidae) (Dewantoro et al., 2017; Novitasari et al., 2014), the Asian striped catfish Hemibagrus nemurus (Valenciennes 1840) (Bagridae) (Subagja and Prakoso, 2018), the clown knife fish Chitala chitala (Hamilton 1822) (Notopteridae) (Setijaningsih et al., 2018), and the striped snakehead Channa striata (Bloch 1793) (Channidae) (Radona et al., 2020). However, not much has been done for induction applications using melatonin to improve reproductive performance in fish. So far, studies have solely used melatonin as an enrichment ingredient in fish feed. It was reported to have a direct action on follicles in the zebrafish Danio rerio (Hamilton 1822) (Danionidae) (increased germinal vesicle breakdown; GVBD) (Carnevali et al., 2011), and in the common carp Cyprinus carpio Linnaeus 1758 (Cyprinidae). It was also observed to stimulate growth and gonad maturation (Maitra et al., 2013), and accelerate gonadal maturation in tilapia (Wardhani, 2020).
This research evaluated the reproductive characteristic and success of Javaen barb spawning induced with ovaprimTM, HCG, and melatonin hormones. This hormone combination was used to analyze the ovulatory response of Javaen barb broodstock and is expected to accelerate the ovulation process and become an alternative solution to the problem of limited availability of the Javaen barb fish seeds.
2. Materials and Methods
This study was conducted at the Germplasm Research Station, Cijeruk, an installation unit of the Research Institute for Freshwater Aquaculture and Fisheries Extension (BRPBATPP), Ministry of Marine Affairs and Fisheries Republic of Indonesia.
2.1. Fish sample
The test fish used were caught in nature and maintained in a controlled environment. The mature female Javaen barb (BW: 142.12 ± 18.08 g and BL: 17.75 ± 0.38 cm) was selected as many as 12 fish with 1-1.2 mm oocytes diameter through ovarian cannulation. Meanwhile, 15 mature males (BW:120.28 ± 6.47 g and BL: 16.23 ± 0.51 cm) were selected based on sperm production through stripping. The broodstock was kept separately in a rectangular concrete pond measuring 2 m × 5 m with a water level of 1 m (10 tons) and water temperature ranging between 24 and 28 °C. The Javaen barb fish were first marked to facilitate observation of each treatment using a tagging number.
2.2. Induce ovulation
This study was conducted experimentally using a completely randomized design with four treatments and three replications each. Each replication consisted of 3 female. Fish were injected with LHRHa+AD (ovaprimTM) 0.6 mL/kg without the addition of melatonin (abbreviated as L0.6M0 treatment), ovaprimTM 0.3 mL/kg with melatonin 0.25 mg/kg (L0.3M0.25), HCG 500 IU/kg plus melatonin 0.25 mg/kg (H500M0.25), and melatonin 0.25 mg/kg (M0.25). L0.6 and L0.3 injections were carried out two times. The first injection used 40% of the treatment dose, while the second injected used 60% (6 hours after the first injection). The M0.25 injection was performed once, along with the first injection. In the H500M0.25 treatment, melatonin injection was carried out 24 hours after HCG injection. Before the injection, fish were anesthetized with Ocean Free 5 mL/L for five minutes.
2.3. Parameters observation
The ovulation latency period (hours), ovulation rate (%), gonadosomatic index (GSI) (%), number of ovulation eggs, fertilization rate (%), hatching rate (%), survival of 3-day-old larvae (%), and body weight and body length of fully yolk sack-absorbed larvae were observed. The latency time and ovulation rate were observed and calculated based on a study by Radona et al. (2020) in striped snakehead Channa striata. The number of ovulated eggs was calculated based on the gravimetric method referring to Ameer et al. (2021) in African catfish. 100 ovulated eggs were fertilized and stocked into baskets with a diameter of 11.5 cm, a water level of about 2.5 cm, and water temperature ranging from 26 to 28oC. A total of 12 baskets were prepared, representing four treatments and three replications. The FR was observed 2 hours after fertilization, while the HR was observed after 24 hours. The SR was observed three days after hatching. Larvae body length was measured using a microscope equipped with an ocular micrometer (Olympus stereo SZ 61), and weight measurements were carried out using a digital balance (KERN ABJ 220-4NM) with a measurement accuracy of 0.1 mg.
2.4. Statistical analysis
Data were tabulated and analyzed based on the analysis of variance (ANOVA) at p<0.05. Differences among treatments were analyzed further with Duncan's test using SPSS version 18.
3. Results
3.1. Latency period, ovulation, fertilization, and hatching rates
The latency time and ovulation rate of Javaen barb after hormone induction are presented in Table 1. Eight individuals were ovulated based on the observations on 12 spawned Javaen barbs. Javaen barb broodstock induced by ovaprimTM without melatonin (L0.6M0) and with melatonin (L0.3M0.25) showed the fastest latency time (680 ± 7 minutes and 685 ± 7 minutes, respectively), and all fish were ovulated. The H500M0.25 treatment had an ovulation rate of 66.7%, while those injected with only melatonin (M0.25) did not ovulate (Table 1). The GSI and fecundity parameters showed the same results as the latent time and ovulation rate parameters. Hormone induction with L0.6M0 and a combination of gonadotropin and melatonin hormones (L0.3M0.25) had the highest GSI and fecundity values. The GSI values were 12.58 ± 1.34% and 12.37 ± 1.23%, respectively. The fecundity values were 55,675 ± 5,103 eggs per female and 49,172 ± 7,353 eggs per female, respectively, where these values indicated significantly different (P<0.05) compared to the treatments using a combination of human chorionic gonadotropin and melatonin hormones induction (H500M0.25) and control (M0.25) (Table 2).
Latency time of ovulation, ovulation, and spawning rate of Javaen barb broodstock induced with a combination of ovaprimTM, HCG, and melatonin hormones.
Gonadosomatic index (GSI) and numbers of ovulated eggs of Javean barb broodstock induced with a combination of ovaprimTM, HCG, and melatonin hormones.
Reproductive performance (FR, HR, and SR) of Javaen barb induced by ovaprimTM, HCG, and melatonin hormones are presented in Table 3. The induction treatment, L0.3M0.25 had the highest FR value (P<0.05) with 95.00 ± 1.41% compared to the other induction treatments. In terms of HR, the induction treatment with L0.6 and hormone induction with L0.3M0.25 showed no different (P>0.05) with values of 51.08 ± 1.16% and 50.17 ± 0.66%, respectively, where the values produced by the two treatments showed a significant difference (P<0.05) compared to the induction treatment with H500M0.25. Meanwhile, the survival rate for each hormone induction treatment showed no difference (P>0.05).
Fertilization, hatching and larval survival rate of Javaen barb broodstock induced with ovaprimTM, HCG, and melatonin hormones.
3.2. Larval performance
The body length and weight of post-hatching and post-yolk sack absorption of Javaen barb larvae are presented in Table 4. All parameters of larval characteristics showed the same value for all treatments (P>0.05).
Larval performance of Javaen barb broodstock induced with a combination of ovaprimTM, HCG, and melatonin hormones.
4. Discussion
Induction using gonadotropin hormone can increase gonadal maturity and reproductive performance outside the fish spawning season (Zamri et al., 2022; Ameer et al., 2021; Acharjee et al., 2017; Zadmajid, 2016; Selvaraj et al., 2012). Tropical cyprinid fish will generally experience spawning periods during the rainy season (Mujtahidah et al., 2019). Differences in latency ovulation time, ovulation rate, GSI values, and fecundity that occurred among hormone induction (ovaprimTM 0.6 mL/kg), (ovaprimTM 0.3 mL/kg combined with melatonin 0.25 mg/kg) and the induction treatment using hormones (HCG 500 IU/kg + melatonin 0.25 mg/kg) is thought to be due to the influence of differences in the induced hormone content and the dose. Different amounts and hormone content can increase the efficiency and effectiveness of the artificial spawning process (Zamri et al., 2022; Dhas et al., 2017). Induction using ovaprimTM 0.6 mL/kg and ovaprimTM 0.3 mL/kg combined with melatonin 0.25 mg/kg showed the optimal results with significantly different GSI and fecundity values compared to the treatment induction using HCG 500 IU/kg + melatonin 0.25 mg/kg (P>0.05). According to DiMaggio et al. (2013, 2014), the effect of dose and hormone content induced during the spawning process on reproductive performance in fish is something that must be considered because it relates to the spawning frequency of fish species and egg quality.
The use of LHRHa+AD in accelerating the final gonad maturation and ovulation process and increasing reproductive performance has been proven successful in many fish species worldwide. Ameer et al. (2021) reported that using ovaprim at a dose of 0.5 mL/kg in African catfish Clarias gariepinus (Burchell 1822) (Clariidae) accelerated ovulation latency time, successful spawning, and increased reproductive performance (fertilization rate, hatching rate, fecundity, and larval survival) compared to using hMG (human menopausal gonadotropin). DiMaggio et al. (2014) stated that using ovaprim at a dose of 0.5 mL/kg in pigfish Orthopristis chrysoptera (Linnaeus 1766) (Haemulidae) resulted in better spawning performance with better quality eggs and larvae. Subagja and Prakoso (2018) also stated that using the LHRHa+AD hormone at 0.5 mL/kg in Asian redtail catfish Hemibagrus nemurus (Valenciennes 1840) (Bagridae) showed more optimal fertilization and hatching rates. Furthermore, Nosrati et al. (2019) reported that using LHRHa+AD (at doses of 10 and 20 μg/kg) increased reproductive performance in Caspian shemaya fish Alburnus chalcoides (Güldenstädt 1772) (Leuciscidae) with a 100% ovulation rate. In male fish, LHRHa+AD induction can also significantly increase GSI and sperm volume (Cejko and Krejszeff, 2016; Zadmajid, 2016). In the individual development phase, GSI will increase until spawning (Aberkane et al., 2018; Dopeikar et al., 2015). On the other hand, the use of HCG at a dose of 500 IU/kg combined with melatonin at a dose of 0.25 mg/kg in this study did not perform best in reproducing Javaen barb. This result could have happened because the dose used was not optimal, or the function of melatonin as a hormone to stimulate the ovulation process in fish combined with HCG has not been practical. This result is in line with the research of DiMaggio et al. (2014), which stated that HCG induction in pigfish Orthopristis chrysoptera at doses of 500, 1000, 2000, and 4000 IU/kg did not give optimal results. Furthermore, Żarski et al. (2017) reported that induction using ovaprimTM in Eurasian perch Perca fluviatilis (Linnaeus 1758) (Percidae) provided good sperm quality and volume compared to induction using HCG. The same results were found with the long spine scraper Paracapoetra trutta (Heckel 1843) (Cyprinidae) (Zadmajid, 2016) and the levantine scraper Paracapoeta damascina (Valenciennes 1842) (Cyprinidae) (Zadmajid et al., 2018). Several studies reported that using HCG would be effective when combined with LHRHa+AD (Sahadan et al., 2022; Zamri et al., 2022; Radona et al., 2020; Dhas et al., 2017; Hafeez-ur-Rehman et al., 2015).
In this study, the induction of ovaprimTM at a dose of 0.3 mL/kg combined with melatonin at a dose of 0.25 mg/kg showed the same reproductive performance as the induction of ovaprimTM at a dose of 0.6 mL/kg. This result indicates that melatonin (dose 0.25 mg/kg) could substitute 0.3 ml/kg ovaprimTM. These results support the statement of Maitra and Hasan (2016) that melatonin can stimulate the release of GnRH from the hypothalamus and increase the concentration of gonadotropin hormones in the pituitary, affecting the LH hormone. In addition, melatonin can stimulate antioxidants and reduce stress levels to improve reproductive performance. Furthermore, Falcon and Zohar (2018) and Hardeland et al. (2011) stated that melatonin and gonadotropins have complementary mechanisms of action, where melatonin controls the reproduction and spawning of fish. Fertilization and hatching are essential parameters for assessing the accuracy of a hormone and the dose used to induce ovulation and spermatozoa production (Ameer et al., 2021). In this study, induction using the ovaprimTM hormone had better reproductive performance results than HCG induction. Installation with ovaprimTM of 0.3 mL/kg combined with melatonin 0.25 mg/kg had the best results in FR parameters compared to hormone induction treatment with ovaprimTM of 0.6 mL/kg. This result showed that increasing the dose of induced gonadotropin hormone can harm reducing FR (Dimaggio et al., 2013; Subagja and Prakoso, 2018). So far, induction of gonadotropin hormones can improve the quality of eggs and larvae (Dewantoro et al., 2017; DiMaggio et al., 2013, 2014) but has not affected the length and weight characters of Javaen barb larvae in the larval stage. In this study, induction using the hormone ovaprimTM 0.6 mL/kg and a combination of hormones ovaprimTM 0.3 mL/kg + melatonin 0.25 mg/kg only showed the best ovulation performance and reproductive biology characteristics.
5. Conclusion
Combining melatonin hormone (0.25 mg/kg) and ovaprimTM (0.3 mL/kg) can accelerate the final gonad maturation process in Javaen barb fish. Melatonin can substitute 50% of the commercial spawning induction hormone premix (GnRH+AD) in ovulation induction and spawning of Javaen barb fish.
Acknowledgements
This study was supported by the Research Institute for Freshwater Aquaculture and Fisheries Extension (SP DIPA-32.12.2.403829/2021), Ministry of Marine Affairs and Fisheries, Republic of Indonesia. We would like to thank Prof. Anang Hari Kristanto for his advice and Hassane Nadio from Hi English Course for proofreading the present article. We would also thank Mrs. Wahyulia Cahyanti, Mr. Sudarmaji, Mr. Sirodiana, Miss. Fera Permata Putri, Mr. Heppy Aprilistianto, Mr. Ujang Heri, and Mr. Yudi Mulyadi for their assistance during this research.
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