Spawning induction and hormonal levels during final oocyte maturation in the silver perch (Bidyanus bidyanus)
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1 Aquaculture 229 (2004) Spawning induction and hormonal levels during final oocyte maturation in the silver perch (Bidyanus bidyanus) Berta Levavi-Sivan a, *, Rima Vaiman a, Ofer Sachs b, Itai Tzchori b a Department of Animal Sciences, Faculty of Agricultural, Food and Environmental Quality Sciences, The Hebrew University, P.O. Box 12, Rehovot 76100, Israel b Aquaculture Research Station-DOR, M.P Hof Hacarmel 30820, Israel Received 18 November 2002; received in revised form 11 April 2003; accepted 15 April 2003 Abstract The silver perch (Bidyanus bidyanus Mitchell) (Teraponidae), is a native Australian freshwater fish that, due to its high potential for aquaculture, was introduced to Israeli fish farming. The objective of this study was to find an optimal method for inducing spawning of silver perch. The agents tested for this purpose were: human chorionic gonadotropin (hcg; 150 or 200 IU/kg BW); salmon gonadotropin releasing hormone analogue (sgnrha at 10, 20, 30, or 40 Ag/kg BW); mammalian GnRH analogue (mgnrha at 30 Ag/kg) and the combination of sgnrha at 20 Ag/kg and domperidone at 5 mg/kg BW. Based on spawning success and relative fecundity, sgnrha at the dose of 30 Ag/kg was found to be more efficient than hcg, mgnrha or sgnrha with domperidone. Since domperidone does not improve the GnRHa effect on final oocyte maturation (FOM) and spawning, it is suggested that the dopaminergic inhibition during the stages of FOM in the silver perch is weak. Therefore, the use of GnRHa alone is sufficient to induce spawning in this fish. Immunoreactive gonadotropin (IR-GtH) and estradiol levels increased after a single injection of sgnrha, and peaked after 24 h. Plasma levels of 17a,20h-dihydroxy-4-pregnen-3-one (17,20-P) also increased significantly 24 h after the injection of mgnrha, 12 h before spawning, suggesting that 17,20-P is the maturation-inducing steroid in silver perch. In order to reveal whether the heterologous gonadotropin may elicit an immunological reaction, silver perch was subjected to prolonged treatment with hcg. This treatment resulted in no detectable titer of antibodies against the mammalian gonadotropin. In conclusion, although hcg has no deleterious effects in this fish, and is the more commonly used for spawning induction, sgnrha at 30 Ag/kg is the recommended * Corresponding author. Tel.: ; fax: address: sivan@agri.huji.ac.il (B. Levavi-Sivan) /$ - see front matter D 2004 Elsevier Science B.V. All rights reserved. doi: /s (03)
2 420 B. Levavi-Sivan et al. / Aquaculture 229 (2004) treatment for spawning induction of female silver perch under the conditions prevailing in Israeli aquaculture. D 2004 Elsevier Science B.V. All rights reserved. Keywords: GnRH; Silver perch; Final oocyte maturation; Domperidone; Estradiol; GTH; 17a,20h-dihydroxy-4- pregnen-3-one; Human chorionic gonadotropin; Spawning induction 1. Introduction The silver perch (Bidyanus bidyanus Mitchell) (Teraponidae), is a freshwater fish native to Australia with high potential for aquaculture due to its rapid growth under diverse conditions (Rowland et al., 1995). Due to its high production rates, the silver perch was chosen to enrich the variety of commercial fish species in Israel. Introduction of silver perch in this country was based on broodfish collected from wild populations in the Murray Darling River in Australia. In their native habitat, silver perch spawn during the summer floods when water temperatures are above 20 jc. During drought years, there is little or no spawning. The spawning and hatchery techniques for the silver perch are well developed in Australia (Rowland, 1984; Rowland et al., 1995). The silver perch does not spawn naturally under the conditions prevailing in Israeli aquaculture. Gonadotropin releasing hormone (GnRH) is a principal stimulator of follicle stimulating hormone (FSH = GtH I) and luteinizing hormone (LH = GtH II) release in teleost fish (reviewed by Van Der Kraak et al., 1998). In addition to the stimulatory effect of hypothalamic factors, LH secretion in teleost is also believed to be under inhibitory control by the hypothalamus. The gonadotropin-inhibiting factor (GRIF) was identified as dopamine (reviewed by Peter et al., 1986). Dopamine directly inhibits the GnRH-induced LH response in a wide array of teleosts including goldfish (Carassius auratus; Peter et al., 1991), common carp (Cyprinus carpio; Billard et al., 1983), European eel (Anguilla anguilla; Dufour et al., 1988), catfish (Clarias gariepinus; De Leeuw et al., 1986), and tilapia (Oreochromis niloticus Oreochromis aureus; Levavi-Sivan et al., 1995). Final oocyte maturation and spawning is usually achieved in these species only after combining GnRHa treatment with a dopamine receptor antagonist. The importance of dopamine inhibitory regulation varies among different teleosts; it is relatively weak in salmonids and some perciforms, thus blockade of dopamine inhibition only slightly potentiates GnRHinduced LH secretion (Peter et al., 1991). Other perciform fish can spawn after induction with GnRHa alone as reported in sea bream (Sparus aurata; Zohar, 1988), Atlantic croaker (Micropogonias undulates; Copeland and Thomas, 1989), and sea bass (Dicentrarchus labrax; Prat et al., 2001). GnRH stimulates LH release by the pituitary, which in turn promotes the maturational competence of oocytes and the production of the maturationinducing steroid (MIS) from the ovary. In many fish, it is the progestogen 17a, 20h dihydroxy-4-pregnen-3-one. The MIS secreted by the induced follicle promotes the synthesis in the oocyte of the maturation promoting factor responsible for the resumption of the meiotic division and consequently ovulation (Nagahama, 1994). In Australian aquaculture, silver perch are left to spawn naturally or are induced to spawn with human
3 B. Levavi-Sivan et al. / Aquaculture 229 (2004) chorionic gonadotropin (hcg) injections (Rowland, 1984). We suspected that repeated treatment with the heterologous gonadotropin may generate antibodies against the gonadotropin jeopardizing the future fertility of the broodstock. Therefore, objectives of the current study were: (1) to discover if silver perch treated with hcg develop anti-hcg antibodies. (2) to establish a protocol for spawning induction employing GnRHa which will be based on hormonal profile, spawning success and relative fecundity, and (3) to investigate the importance of dopaminergic inhibition on LH release, FOM and ovulation in silver perch. 2. Materials and methods 2.1. Fish Adult silver perch, B. bidyanus, of both genders, originally captured in the wild, were purchased from a commercial hatchery in Australia and brought to the Aquaculture Research Station-DOR in Israel. The fish were kept in tanks under natural photoperiod and temperature (31j30VN, 34j45VE). Fish were fed twice a day (at 9 AM and 4 PM) with commercial pellets (Hepalim), composed of 47% protein, 11% fat, 9% ash and 2.2% cellulose at 0.5% of the biomass and the food excess was regularly removed. The experiments were carried out in the summer of 2000, when water temperature was jc. During the experiment, each group of fish were kept in separate well-aerated 2500-l tanks, with well running water Antibodies against hcg Control male silver perch fish were 1 2 years old that had never been subjected to hcg treatment. hcg-sensitized male fish were 3 years old and had been injected more than four times with hcg (200 IU/kg; Sigma), over the last 2 years. They were boosted with hcg at the same dose three additional times, at 2-week intervals, before bleeding. Human Chorionoc Gonadotropin (5 Ag) was iodinated by the Iodogen method (Salachinski et al., 1981), with 0.5 ACi 125 I (Amersham, UK). The labeled standard was separated from the free iodine by chromatography on Sephadex G-25 minicolumn. The binding assays were performed generally according to Van der Kraak et al. (1989). Serum samples and labeled hcg were diluted in phosphate buffer, ph 8.0, containing 0.1% BSA. For binding assays, 100 Al serum diluted 1:10 and 1:50 were incubated overnight at room temperature with 125 I-hCG (30,000 cpm). Bound hormone was precipitated by the addition of 400 Al cold 40% polyethylene glycol-4000 and centrifugation for 30 min at 2000 g, at4jc. The unbound fraction was aspirated and the radioactivity of the bound fraction was counted. Positive controls, in which 125 I-hCG was incubated with anti-hcg serum (donated by Dr. Venktaesh, India), was added to ensure the reliability of the assay. In each assay, 125 I-hCG was incubated with serial dilutions of anti-hcg serum (1:1000 1:10,000) and yielded a standard curve with high correlation coefficient (0.965 F 0.003).
4 422 B. Levavi-Sivan et al. / Aquaculture 229 (2004) LH determination Plasma LH in silver perch was determined using a heterologous radioimmunoassay developed for another perciform fish (tilapia; Bogomolnaya et al., 1989) following the modified procedure of Levavi-Sivan and Yaron (1992). The second antibody employed was donkey anti-rabbit IgG bound to magnetic beads (Amerlex-M, Amersham). In order to verify the accuracy of the heterologous assay, serial dilutions of silver perch plasma and pituitary extract were determined in the same assay together with the standard and all were found to be parallel (Fig. 1). Sensitivity and the intra- and inter-assay coefficients of variance were as described in Levavi-Sivan et al. (1995). Since the standard tilapia GtH (tagth) used in this study was isolated and purified from pituitaries of adult fish collected during the spawning season (Bogomolnaya et al., 1989), it is assumed that tagth corresponds to GtH II or LH as defined in the salmon (Kawauchi et al., 1989) Steroid determinations Estradiol (E2) and 17a,20h-dihydroxy-4-pregnen-3-one (17,20-P) were measured by enzyme linked immunosorbent assay (ELISA), according to Cuisset et al., (1994), and Nash et al. (2000), using acetylcholinesterase as a label. Samples consisting of 250 Al of plasma were extracted with diethyl ether (for E2) or with ethyl acetate (for 17,20-P). The anti-e2 and anti 17,20-P used were as previously described (Levavi-Zermonsky and Yaron, 1986; Yaron and Levavi-Zermonsky, 1986). All samples were analyzed in duplicate, and for each ELISA plate, a separate standard curve was run. The lower limit Fig. 1. Regression lines obtained from serial dilutions of silver perch pituitary extract (diamond), plasma (circle) and standard tilapia gonadotropins (square) by radioimmunoassay. Log-Logit transformation.
5 of detection was 1.8 and 1.7 pg/ml for E2 and 17,20-P respectively. The intra- and interassay coefficients of variance for both assays were less than 5% and 11%, respectively. Steroid levels, determined by ELISA, were validated by verifying that serial dilutions were parallel to the standard curves Experimental design and spawning induction In order to determine the readiness of fish for spawning induction manipulation, fish were anaesthetized using clove oil (0.2 ml/l water), and oocyte samples were obtained via the genital pore using a 1-mm-diameter tygon tubing. The diameter of 100 oocytes from each female was recorded; only fish with oocytes larger than 700 Am were chosen for spawning induction. Fish were injected intraperitonealy with the following agents: hcg (150 or 200 IU/kg BW; ([D-Arg 6,Pro 9 -NEt]-salmon GnRH; Bachem; sgnrha (10, 20, 30, or 40 Ag/kg); ([D-Ala 6,Pro 9 -NEt]-mammalian GnRH, Sigma; mgnrha; 30 Ag/kg; or sgnrha at 20 Ag/kg combined with domperidone (5 mg/kg; Sigma). The experiment was conducted from May to July. Two injected males and six injected females were placed into 1000-l tanks and water temperature was maintained at jc. The tanks were examined for the presence of eggs starting 35 h after injection. Thirty minutes after spawning, the water-hardened eggs were removed from the tank, their volume was measured and they were incubated in 60-l Zuger funnels. The next experiment was conducted in order to find out whether dopaminergic inhibition of LH release exists in the silver perch during final oocyte maturation. Sixty mature female silver perch were divided into four groups and injected as follows: sgnrha dissolved in saline was injected at 20 Ag/kg; domperidone dissolved in DMSO was injected at 5 mg/kg BW; sgnrha + domperidone at the same doses or saline + DMSO. Domperidone and sgnrha were injected in separate injections at the same time. A 0.7-ml blood sample was taken, at the times indicated, from the caudal vessels into heparinised syringes; the plasma was separated by centrifugation and stored at 20 jc until further analysis. In another experiment, 15 females received one i.p. injection of mgnrha dissolved in saline at the dose 30 Ag/kg. Blood was sampled before injection, (time 0), and 12, 18, 24 and 36 h thereafter. The control group was sampled before injection and, 18 and 36 thereafter Statistical analysis B. Levavi-Sivan et al. / Aquaculture 229 (2004) Data are presented as mean F standard error of mean (S.E.M.). The significance of the differences between group means of hormone levels was determined by one-way analysis of variance (ANOVA) followed by Newman Keuls test. For testing parallelism between different regression lines, we used Analysis of Covariance at: edu/~harsham/business-stat/otherapplets/anocov.htm. 3. Results The binding of 125 I-hCG to the fish serum was similar in control (18.65 F 0.66%; n = 11) and hcg-injected silver perch (19.74 F 0.07%; n = 15). This indicates that silver
6 424 Table 1 Effect of various agents on spawning parameters Treatment Dose No. of injected females No. of spawned females Female body weight (g) Spawning ratio 1 (%) Egg volume (ml/kg) Latency period (h) Relative fecundity 2 ( 10 3 ) hcg 150 IU/kg F F F 0.50 a F a hcg 200 IU/kg F F F 0.34 a F 7.60 a mgnrha 30 Ag/kg F F F 0.92 a F a sgnrha 10 Ag/kg F F F 1.62 a F b sgnrha 20 Ag/kg F F F 2.00 a F a sgnrha 30 Ag/kg F F F 1.17 a F b sgnrha 40 Ag/kg F F F 1.25 a F a sgnrha + Domperidone 20 Ag/kg + 5 mg/kg F F F 1.80 a F a Female body weight, egg volume, latency period and relative fecundity are expressed as mean F S.E.M. Means designated by the same letter do not differ significantly from each other ( p>0.05). 1 Spawning ratio: the number of females which ovulated after injection divided by the total number of injected females. 2 Relative fecundity: total number of eggs/kg body weight of treated females. B. Levavi-Sivan et al. / Aquaculture 229 (2004)
7 B. Levavi-Sivan et al. / Aquaculture 229 (2004) perch treated repeatedly with hcg do not produce antibodies against the heterologous gonadotropin. Linearized tagth (LH) standard curve and serial dilutions of plasma or pituitary extract from silver perch are shown in Fig. 1. Each point is the mean of triplicate determinations of serial dilutions of a single sample. These dilution curves were found to be parallel to that of tagth standard, indicating that silver perch LH is immunologically similar to the tilapia LH and can be measured by the same RIA. When the data of standard curves were transformed (LOGIT) to a linear plot, the correlation coefficients of the lines were F and F for silver perch pituitary extract and plasma, respectively (n = 3 for each curve). Analysis of variance of the slopes for the different regression lines showed that the three slopes do not differ from each other ( p>0.05). This study validates the heterologous RIA for determining the levels of silver perch LH in plasma and pituitary. The next experiment was design in order to test the ability of various agents to induce spawning induction in the silver perch. The highest dose of hcg used (200 IU/kg BW) was more efficient in spawning rate than the lower dose (150 IU/kg BW) (97.4% vs. 67.7%, respectively), with no significant difference in the fecundity (Table 1). In order to find the most efficient dose of sgnrha for spawning induction of female silver perch, we tested the range of doses found to be potent in other species (10 40 Ag/kg). The most efficient doses in terms of spawning percentage were 30 or 40 Ag/kg. However, females treated with sgnrha at the dose 30 Ag/kg had significantly higher fecundity ( F vs F for 30 or 40 Ag/kg, respectively). Treatment with the combination of sgnrha and the dopamine antagonist domperidone yielded only 80% spawning and lower fecundity (Table 1). Injection of females with mgnrha (30 Ag/kg) resulted in high spawning percentage but significantly lower fecundity. No significant Fig. 2. Concentration of IR-GtH (mean F S.E.M., n = 8 11) in the plasma of female silver perch following injection of saline (control); sgnrha (20 Ag/kg; GnRH); domperidone (5 mg/kg; DOM; or both agents at the same doses (GnRH + DOM). Spawning was recorded after 39.2 F 1.80 h after injection. Statistically significant differences between groups are shown by different letters.
8 426 B. Levavi-Sivan et al. / Aquaculture 229 (2004) Fig. 3. Estradiol levels in (mean F S.E.M., n = 8 11) the plasma of female silver perch following injection of saline (control); sgnrha (20 Ag/kg; GnRH); domperidone (5 mg/kg; DOM; or both agents at the same doses (GnRH + DOM). Spawning was recorded after 39.2 F 1.80 h after injection. Statistically significant differences between groups are shown by different letters. difference was noted in the spawning latency (the time interval between the injection and the spawning) between fish treated with either of the hormones or doses. The next experiment was conducted in order to determine whether dopaminergic inhibition of LH is predominant in the silver perch and whether GnRH treatment for spawning induction should be combined with a dopamine antagonist. Females were treated with sgnrha (20 Ag/kg) alone or in combination with the dopamine receptor antagonist domperidone. The fluctuations in plasma concentration of IR-LH observed in the control group were not significant (Fig. 2). However, significantly higher levels of IR- Fig. 4. Concentration of IR-GtH (mean F S.E.M., n = 7 10) in the plasma of female silver perch following injection of saline (control) or mgnrha (30 Ag/kg; GnRH). Spawning was recorded after 38.0 F 0.92 h after injection. Statistically significant differences between groups are shown by different letters.
9 B. Levavi-Sivan et al. / Aquaculture 229 (2004) Fig. 5. Concentration of 17,20-P (mean F S.E.M., n = 7 10) in the plasma of female silver perch following injection of saline (control) or mgnrha (30 Ag/kg; GnRH). Spawning was recorded after 38.0 F 0.92 h after injection. Statistically significant differences between groups are shown by different letters. LH were recorded in the group injected with GnRHa alone compared to the group injected with domperidone only (DOM) or in combination with GnRH reaching a peak after 24 h and later declining (Fig. 2). Spawning occurred 39.2 F 1.80 h after the injection and IR- LH levels remained low after spawning. Similar results were obtained in plasma E2 levels. The highest levels of E2 were recorded 24 h after injection with GnRH alone and remained low after spawning (Fig. 3). No change in plasma concentration of IR-LH was observed in the saline-injected controls (Fig. 4). However, significantly higher concentrations of the hormone compared to the control were recorded already 12 h after injection of mgnrha (30 Ag/kg; Fig. 4). The levels were still high 38 h after injection when the females were in the process of spawning. The levels of 17a,20h-dihydroxy-4-pregnen-3-one (17,20-P) in the same fish gradually increased after injection and peaked 12 h before the fish spawned (Fig. 5) but the levels were low during spawning. 4. Discussion Introduction of a new species into Israeli aquaculture required the establishment of a protocol for spawning induction in order to be able to complete the whole life cycle of the fish under the conditions prevailing locally. The silver perch does not spawn naturally in Israel. Although spawning induction has been successfully achieved in Australia using hcg, it was suspected that such procedure may lead to the generation of antibodies against the heterologous hormone and when the same treatment will be applied in subsequent years, the fish will either need higher doses or will not respond (Donaldson and Hunter, 1983; Mylonas and Zohar, 2001). Nevertheless, no significant binding of 125 I-hCG was detected in serum samples of silver perch previously subjected four times to hcg for spawning induction and in addition, three times before blood sampling. Moreover, females
10 428 B. Levavi-Sivan et al. / Aquaculture 229 (2004) treated with the same dose of hcg (200 IU/kg) for more than 3 years showed no reduction in their response (Table 1). Similarly, antibodies against hcg were not detected in silver carp broodstock or in goldfish immunized with hcg administered singly or in combination with adjuvant (Van der Kraak et al., 1989). In contrast, significant titers of hcg antibodies were observed in the circulation of striped bass treated with hcg (Mylonas and Zohar, 2001). This discrepancy can be explained by the different methods used for determining the antibody titer, the different protocols used for immunization or, most likely to difference between fish species. Hormonal manipulations for the induction of FOM, ovulation, spermiation and spawning have made possible the control of reproductive processes of cultured fish, and have contributed significantly to the sophistication and expansion of the aquaculture industry. Various GnRH analogs have been used successfully to induce ovulation in a number of fish species. Mylonas and Zohar (2001) listed 21 species where ovulation was induced using GnRHa at doses ranging from 1 to 600 Ag/kg body weight. However, the responsiveness of individual fish, in terms of number of ovulations and number of eggs ovulated, was highly variable. Although the present work showed that the silver perch do not form antibodies against hcg, and the spawning ratio of females induced by hcg (200 IU/kg) did not differ significantly from females induced by sgnrha (30 40 Ag/kg), the relative fecundity was lower in the former. Based on the spawning ratio and relative fecundity, sgnrha at the dose 30 Ag/kg seems to be the most effective and is recommended for spawning induction of female silver perch under the conditions prevailing in the Israeli aquaculture. The silver perch ovary is defined as having a group synchronous development that leads to multiple spawnings (Rowland, 1984). Nevertheless, in the present study only a single spawning occurred after injecting hcg or a GnRH analog in a single injection. In Australia, the mean latency period after injection of 200 IU/kg hcg was 35.7 F 2.4 and all (n = 7) the fish spawned (Rowland, 1984), while in our study the same dose of hcginduced spawning with a mean latency of 41 F 0.33 and 97% (n = 38) of the fish spawned. This study was also designed to answer the question of whether dopaminergic inhibition of LH release is predominant similar to the situation in tilapia, carp, loach (Paramisgurnus dabryanus), catfish or goldfish (Levavi-Sivan et al., 1995; Lin et al., 1989; De Leeuw et al., 1986; Chang et al., 1984, respectively) or whether the dopaminergic inhibition is weak or absent, similar to the situation in the sea bass, Atlantic croacker, or striped bass (reviewed by Mylonas and Zohar, 2001). The results clearly show that the use of a dopamine antagonist does not potentiate the effect of GnRHa treatment in inducing FOM and spawning in the silver perch and that GnRHa alone is sufficient to increase LH, E2 and 17,20-P levels and to induce spawning. The injection of mgnrha resulted in elevation of LH in the plasma similar to the response to sgnrha, but with a longer duration of peak levels. However, sgnrha was more effective than mgnrha in spawning success and induced higher fecundity. The reason for the longer effect of mgnrh is not certain but it is likely that its metabolic clearance is slower than that of sgnrha, as was shown for the sea bream (Goren et al., 1990). Postvitellogenic silver perch ovaries, containing oocytes with a diameter larger than 700 Am, responded to the GnRH-stimulated LH surge by an enhanced output of estradiol.
11 B. Levavi-Sivan et al. / Aquaculture 229 (2004) Plasma E2 levels increased significantly after 12 h, in response to GnRHa injection, and peaked after 24 h. A similar increase in E2 levels from postvitellogenic oocytes before final oocyte maturation was also reported in the carp (Levavi-Zermonsky and Yaron, 1986), longfinned eels (Anguilla dieffenbachia; Lokman et al., 2001); and gilthead seabream (Gothilf et al., 1997). Since the silver perch is a group synchronous spawner, the ovaries of spawning fish contain at this stage, not only maturing follicles but also follicles at earlier stages of growth and development. Therefore, the increase in E2 levels may be associated with the ongoing induction of vitellogenin and oocyte growth. Plasma 17,20-P levels showed a significant increase 24 h after injection of mgnrha which occurred 12 h before actual spawning. This is consistent with findings in other species that during oocyte maturation the follicle shifts from the secretion of estradiol (C18) to the secretion of 17a,20h, dihydroxy-4-pregnene-3-one (17,20-P; C21), probably through the activation of 20h-hydroxy steroid dehydrogenase (Nagahama, 1994; Nagahama et al., 1994). Our results indicates that 17,20-P is also involved in the final maturation of the oocytes in the silver perch. A similar rise in 17,20-P level around final maturation and ovulation was reported in the Atlantic and coho salmons and rainbow trout (Salmo salar, Oncorhynchus kisutch, Salmo gairdneri; Wright and Hunt, 1982); ayu (Plecoglossus altivelis; Hirose et al., 1983); goldfish (Kagawa et al., 1983), African catfish (Lambert and van den Hurk, 1982); carp (Levavi-Zermonsky and Yaron, 1986); gilthead seabream (Kadmon et al., 1985; Gothilf et al., 1997), striped trumpeter (Latris lineate; Morehead et al., 1998), and Japanese yellowtail (Seriola quinqueradiata; Rahman et al., 2001). 5. Conclusion Our results show that the injection of hcg to silver perch fish does not results in the formation of antibodies against the mammalian gonadotropin. We also showed that domperidone does not improve the GnRHa effect on FOM and that GnRHa alone is sufficient for spawning induction in the silver perch. Based on the spawning ratio and relative fecundity, sgnrha at the dose of 30 Ag/kg was found to be more efficient than hcg, mgnrha or sgnrh + domperidone, and hence is the recommended treatment for spawning induction of female silver perch under the conditions prevailing in Israel. Acknowledgements This work was supported by a grant from the Israeli Organization of Fisheries. References Billard, R., Alagarswami, K., Peter, R.E., Breton, B., Potentialisation par le pimozide des effets du LHRH- A sur la sécrétion gonadotrope hypophysaire, l ovulation et la spermiation chez la carpe commune (Cyprinus carpio). C. R. Acad. Sci. (Paris) 296, Bogomolnaya, B., Yaron, Z., Hilge, V., Graesslin, D., Lichtenberg, V., Abraham, M., Isolation and radioimmunoassay of a steroidogenic gonadotropin of tilapia. Isr. J. Aquac.-Bamidgeh 41, Chang, J.P., MacKenzie, D.S., Gould, D.R., Peter, R.E., Effects of dopamine and norepinephrine on in vitro
12 430 B. Levavi-Sivan et al. / Aquaculture 229 (2004) spontaneous and gonadotropin-releasing hormone-induced gonadotropin release by dispersed cells or fragments of the goldfish pituitary. Life Sci. 35, Copeland, P.A., Thomas, P., Control of gonadotropin release in the Atlantic croaker (Micropogonias undulatus): evidence for lack of dopaminergic inhibition. Gen. Comp. Endocrinol. 74, Cuisset, B., Pradelles, P., Kime, D.E., Kuhn, E.R., Babin, P., Le Menn, F., Enzyme immunoassay for 11- ketotestosterone using acetylcholineesterase as label. Application to the measurement of 11-ketotestosterone in plasma of Siberian sturgeon. Comp. Biochem. Physiol. 108, De Leeuw, R., Goos, H.J.Th., Van Oordt, P.G.W.J., The dopaminergic inhibition of the gonadotropinreleasing hormone-induced gonadotropin release: an in vitro study with fragments and cell suspensions from pituitaries of African catfish, Clarias gariepinus (Burchell). Gen. Comp. Endocrinol. 63, Donaldson, E.M., Hunter, G.A., Induced final maturation, ovulation and spermiation in cultured fishes. In: Hoar, W.S., Randall, D.J., Donaldson, E.M. (Eds.), Fish Physiology. Reproduction, vol. 9B. Academic Press, Orlando, FL, pp Dufour, S., Lopez, E., Le Menn, F., Le Belle, N., Baloche, S., Fontaine, Y.A., Stimulation of gonadotropin release and ovarian development by administration of a gonadoliberin agonist and of dopamine antagonists in female silver eel pretreated with estradiol. Gen. Comp. Endocrinol. 70, Goren, A., Zohar, Y., Fridkin, M., Elhanati, E., Koch, Y., Degradation of gonadotropin releasing hormone in the gilthead seabream, Sparus aurata: I. Cleavage of native salmon GnRH, mammalian LHRH and their analogs in the pituitary. Gen. Comp. Endocrinol. 79, Gothilf, Y., Meiri, I., Elizur, A., Zohar, Y., Preovulatory changes in the levels of three gonadotropinreleasing hormone-encoding messenger ribonucleic acids mrnas gonadotropin h-subunit mrnas, plasma gonadotropin, and steroids in the female gilthead seabream, Sparus aurata. Biol. Reprod. 57, Hirose, K., Nagahama, Y., Adachi, S., Wakabayashi, K., Changes in serum concentrations of gonadotropins, 17a-hydroxyprogesteroneand 17a,20h-dihydroxy-4-pregnen-3-one during synthetic LH-RH-induced final oocyte maturation and ovulation in the ayu Plecoglossus altivelis. Bull. Jpn. Soc. Sci. Fish. 49, Kadmon, G., Yaron, Z., Gordin, H., Sequence of gonadal events and estradiol levels in Sparus aurata (L.) under two photoperiod regimes. J. Fish Biol. 26, Kagawa, H., Young, G., Nagahama, Y., Changes in plasma steroid hormone levels during gonadal maturation in female goldfish Carassius auratus. Bull. Jpn. Soc. Sci. Fish. 49, Kawauchi, H., Suzuki, K., Itoh, H., Swanson, P., Naito, N., Nagahama, Y., Nozaki, M., Nakai, Y., Itoh, S., The duality of fish gonadotropins. Fish Physiol. Biochem. 7, Lambert, J.G.D., van den Hurk, R., Steroidogenesis in the ovaries of the African catfish, Clarias lazera, before and after an HCG induced ovulation. In: Richter, C.J.J., Goos, H.J.Th. (Eds.), Reproductive Physiology of Fish. Pudoc, Wageningen, pp Levavi-Sivan, B., Yaron, Z., Involvement of cyclic adenosine monophosphate in the stimulation of gonadotropin secretion from the pituitary of the teleost fish, tilapia. Mol. Cell. Endocrinol. 85, Levavi-Sivan, B., Ophir, M., Yaron, Z., Possible sites of dopaminergic inhibition of gonadotropin release from the pituitary of a teleost fish, tilapia. Mol. Cell. Endocrinol. 109, Levavi-Zermonsky, B., Yaron, Z., Changes in gonadotropin and ovarian steroids associated with oocyte maturation during spawning induction in the carp. Gen. Comp. Endocrinol. 62, Lin, H.R., Van der Kraak, G., Zhou, X.J., Liang, J.Y., Peter, R.E., Rivier, J.E., Vale, W.W., Effects of [D-Arg 6,Trp 7, Leu 8, Pro 9 Net]-luteinizing hormone-releasing hormone (sgnrh-a) and [D-Ala 6, Pro 9 Net]- luteinizing hormone releasing hormone (LHRH-A), in combination with pimozide or domperidone, on gonadotropins release and ovulation in the Chinese loach and common carp. Gen. Comp. Endocrinol. 69, Lin, H.R., Peng, C., Van der Kraak, G., Peter, R.E., Dopamine inhibits gonadotropins secretion in the Chinese loach. Fish Physiol. Biochem. 6, Lokman, P.M., Wass, R.T., Suter, H.C., Scott, S.G., Judge, K.F., Young, G., Changes in steroid hormone profiles and ovarian histology during salmon pituitary-induced vitellogenesis and ovulation in female New Zealand longfinned eels, Anguilla dieffenbachii Gray. J. Exp. Zool. 289, Morehead, D.T., Pankhurst, N.W., Ritar, A.J., Effect of treatment with LHRH analogue on oocyte maturation, plasma sex steroid levels and egg production in female striped trumpeter Latris lineata (Latrididae). Aquaculture 169,
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