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Mark P Lokman - One of the best experts on this subject based on the ideXlab platform.

  • spatiotemporal expression of activin receptor like kinase 5 and bone morphogenetic protein receptor type ii in the ovary of shortfinned eel Anguilla australis
    Comparative Biochemistry and Physiology B, 2021
    Co-Authors: Ali Falahati, Erin L. Damsteegt, Yuichi Ozaki, Vahid Zadmajid, Kaitlyn J Freeman, Mark P Lokman
    Abstract:

    In the eel ovary, the expression of growth differentiation factor-9 (Gdf9) appears to be largely confined to the germ cell in early stages of oogenesis. However, both the target tissue and the function of Gdf9 in fish remain unknown. This study aimed to describe the abundance and localization of activin receptor-like kinase-5 (Alk5) and bone morphogenetic protein receptor type II (Bmpr2), which together mediate the Gdf9 signal, in the ovary of a basal teleost, the shortfinned eel, Anguilla australis, during early folliculogenesis. The cDNA encoding eel alk5 and bmpr2 genes were cloned, characterized and the transcript abundances of these receptors quantified by quantitative real-time PCR. Ovarian transcript abundance for both receptors, along with that of gdf9 and of its paralogue bmp15, increased from the previtellogenic to early vitellogenic stage. Localization of receptor mRNAs by in situ hybridization revealed that these receptors are located in the somatic cells surrounding the oocyte. Furthermore, tissue distribution analysis showed that the expression of alk5 and bmpr2 were highest in ovary and thyroid, respectively. Unexpectedly, however, bmpr2 mRNA levels were lower in the ovary than in any of the other 17 tissues examined, and indeed, lower than ovarian gdf9 transcript abundance. These findings, together with the ovarian expression pattern of Gdf9, suggest that Gdf9, and conceivably, Bmp15, from the oocyte can signal through receptors that are located on the somatic cells surrounding the oocyte; this, in turn, facilitates elucidation of the function of these growth factors during oogenesis in teleost fish.

  • induction of oocyte development in previtellogenic eel Anguilla australis
    General and Comparative Endocrinology, 2020
    Co-Authors: Matthew J. Wylie, Yukinori Kazeto, Anh Tuan Nguyen, Jolyn H Z Chia, Mark P Lokman
    Abstract:

    Abstract The role of gonadotropins during early ovarian development in fish remains little understood. Concentrations of gonadotropins were therefore experimentally elevated in vivo by administration of recombinant follicle-stimulating hormone (rec-Fsh) or human chorionic gonadotropin (hCG) and the effects on ovarian morphology, sex steroid levels and mRNA levels of genes expressed in pituitary and ovary examined. Hormones were injected thrice at weekly intervals in different doses (20, 100 or 500 µg/kg BW for rec-Fsh and 20, 100 or 500 IU/kg BW for hCG). All treatments, especially at the highest doses of either rec-Fsh or hCG, induced ovarian development, reflected in increased oocyte size and lipid uptake. Both gonadotropins up-regulated follicle-stimulating hormone receptor (fshr) mRNA levels and plasma levels of estradiol-17β (E2). Exogenous gonadotropins largely decreased the expression of follicle-stimulating hormone β-subunit (fshb) and had little effect on those of luteinizing hormone β-subunit (lhb) in the pituitary. It is proposed that the effects of hCG on ovarian development in previtellogenic eels could be indirect as a significant increase in plasma levels of 11-ketotestosterone (11-KT) was found in eels treated with hCG. Using rec-Fsh and hCG has potential for inducing puberty in eels in captivity, and indeed, in teleost fish at large.

  • expression of gonadotropin subunit and gonadotropin receptor genes in wild female new zealand shortfinned eel Anguilla australis during yellow and silver stages
    General and Comparative Endocrinology, 2019
    Co-Authors: Anh Tuan Nguyen, Yukinori Kazeto, Jolyn H Z Chia, Mark P Lokman
    Abstract:

    Abstract Despite tremendous importance of follicle-stimulating hormone (Fsh) and luteinizing hormone (Lh) as primary controllers of reproductive development, information on the expression profiles of the genes encoding gonadotropin subunits and gonadotropin receptors (Fshr and Lhr) in wild eels are essentially non-existent. This study investigated pituitary fshb and lhb mRNA levels and ovarian fshr and lhr mRNA levels of wild shortfinned eels, Anguilla australis at different stages of oogenesis. Protein expression of Fsh in the pituitary was also quantified and visualized using slot blot and immunohistochemistry. Pituitary fshb and lhb mRNA levels showed a differential expression pattern, fshb mRNA levels increasing significantly from the perinucleolus (PN) to the oil droplet stage (OD) before slightly decreasing (not significantly) in the early vitellogenic stage (EV). A similar trend was observed in relative Fsh protein levels analyzed by slot blot and immunohistochemistry, but this trend was not reflected in the plasma levels of sex steroids. In contrast, pituitary lhb mRNA levels increased significantly from the PN to EV stage. A higher expression of Fsh at both mRNA and protein levels in the pituitary of eels at the OD stage compared to other investigated stages suggests that synthesis of Fsh production in the pituitary may reach a peak at the OD stage. In the ovary, transcript abundances of fshr and lhr gradually increased during previtellogenic follicle growth, but markedly and significantly increased thereafter. Taken together, our data suggest i) that Fsh release may be very limited, or absent, prior to onset of puberty in shortfinned eels and ii) that Lh is not functionally important in this fish during the EV stage.

  • triacylglyceride physiology in the short finned eel Anguilla australis the effects of androgen
    American Journal of Physiology-regulatory Integrative and Comparative Physiology, 2016
    Co-Authors: Erin L. Damsteegt, Yuichi Ozaki, Sally P A Mccormick, Mark P Lokman
    Abstract:

    The importance of androgens (especially 11-ketotestosterone) during previtellogenesis in eels is well established. In wild pubertal migrants, circulating 11-ketotestosterone levels correlate with a...

  • triacylglyceride physiology in the short finned eel Anguilla australis changes throughout early oogenesis
    American Journal of Physiology-regulatory Integrative and Comparative Physiology, 2015
    Co-Authors: Erin L. Damsteegt, Ali Falahatimarvast, Sally P A Mccormick, Mark P Lokman
    Abstract:

    During certain stages in an animal's life cycle, energy requirements may exceed energy intake from the diet. The spawning migration of temperate eels is a textbook example of negative energy balance, forcing these fish to rely on stored fats (triacylglycerides) to provide their muscles with energy for swimming and their growing oocytes with the nutrients needed to develop and support healthy offspring. We predicted broad implications of this great need for endogenous triacylglycerides in terms of their packaging, transport, and ovarian uptake. To test this, serum lipid concentrations and transcript abundances of intestinal and hepatic triacylglyceride packagers and ovarian triacylglyceride modifiers and receivers were investigated throughout previtellogenesis (feeding phase) and into early vitellogenesis (fasting phase) in short-finned eels. A switch from exogenous to endogenous triacylglyceride packaging was seen as the liver upregulated transcript levels of apolipoprotein B and microsomal triacylglyceride transport protein and downregulated those of apolipoprotein E and lipoprotein lipase. In the intestine, the reverse response was observed. Furthermore, ovarian transcript abundances of triacylglyceride modifiers and receivers increased (apolipoprotein E, lipoprotein lipase, and vitellogenin receptor), indicative of increased triacylglyceride uptake during previtellogenesis. We propose that increased hepatic apolipoprotein B production is a conserved vertebrate response to prolonged periods of negative energy balance.

John A. Donald - One of the best experts on this subject based on the ideXlab platform.

  • nitric oxide control of the dorsal aorta and the intestinal vein of the australian short finned eel Anguilla australis
    The Journal of Experimental Biology, 2004
    Co-Authors: Brett L Jennings, Bradley Randal Scott Broughton, John A. Donald
    Abstract:

    SUMMARY This study investigated the mechanisms by which nitric oxide (NO) regulates the dorsal aorta and the intestinal vein of the Australian short-finned eel Anguilla australis . NADPH diaphorase histochemistry and immunohistochemistry using a mammalian endothelial nitric oxide synthase (NOS) antibody could not demonstrate NOS in the endothelium of either blood vessel; however, NOS could be readily demonstrated in the endothelium of the rat aorta that was used as a control. Both blood vessels contained NADPH diaphorase positive nerve fibres and nerve bundles, and immunohistochemistry using a neural NOS antibody showed a similar distribution of neural NOS immunoreactivity in the perivascular nerves. In vitro organ bath physiology showed that a NO/soluble guanylyl cyclase (GC) system is present in the dorsal aorta and the intestinal vein, since the soluble GC inhibitor oxadiazole quinoxalin-1 (ODQ; 10 –5 mol l –1 ) completely abolished the vasodilatory effect of the NO donor, sodium nitroprusside (SNP; 10 –4 mol l –1 ). In addition, nicotine (3×10 –4 mol l –1 ) mediated a vasodilation that was not affected by removal of the endothelium. The nicotine-mediated dilation was blocked by the NOS inhibitor, N ω -nitro-l-arginine (l-NNA; 10 –4 mol l –1 ), and ODQ (10 –5 mol l –1 ). More specifically, the neural NOS inhibitor, N ω -propyl-l-arginine (10 –5 mol l –1 ), significantly decreased the dilation induced by nicotine (3×10 –4 mol l –1 ). Furthermore, indomethacin (10 –5 mol l –1 ) did not affect the nicotine-mediated dilation, suggesting that prostaglandins are not involved in the response. Finally, the calcium ionophore A23187 (3×10 –6 mol l –1 ) caused an endothelium-dependent dilation that was abolished in the presence of indomethacin. We propose the absence of an endothelial NO system in eel vasculature and suggest that neurally derived NO contributes to the maintenance of vascular tone in this species. In addition, we suggest that prostaglandins may act as endothelially derived relaxing factors in A. australis .

  • nitric oxide control of the dorsal aorta and the intestinal vein of the australian short finned eel Anguilla australis
    The Journal of Experimental Biology, 2004
    Co-Authors: Brett L Jennings, Bradley Randal Scott Broughton, John A. Donald
    Abstract:

    This study investigated the mechanisms by which nitric oxide (NO) regulates the dorsal aorta and the intestinal vein of the Australian short-finned eel Anguilla australis. NADPH diaphorase histochemistry and immunohistochemistry using a mammalian endothelial nitric oxide synthase (NOS) antibody could not demonstrate NOS in the endothelium of either blood vessel; however, NOS could be readily demonstrated in the endothelium of the rat aorta that was used as a control. Both blood vessels contained NADPH diaphorase positive nerve fibres and nerve bundles, and immunohistochemistry using a neural NOS antibody showed a similar distribution of neural NOS immunoreactivity in the perivascular nerves. In vitro organ bath physiology showed that a NO/soluble guanylyl cyclase (GC) system is present in the dorsal aorta and the intestinal vein, since the soluble GC inhibitor oxadiazole quinoxalin-1 (ODQ; 10(-5) mol l(-1)) completely abolished the vasodilatory effect of the NO donor, sodium nitroprusside (SNP; 10(-4) mol l(-1)). In addition, nicotine (3 x 10(-4) mol l(-1)) mediated a vasodilation that was not affected by removal of the endothelium. The nicotine-mediated dilation was blocked by the NOS inhibitor, N(omega)-nitro-L-arginine (L-NNA; 10(-4) mol l(-1)), and ODQ (10(-5) mol l(-1)). More specifically, the neural NOS inhibitor, N(omega)-propyl-L-arginine (10(-5) mol l(-1)), significantly decreased the dilation induced by nicotine (3 x 10(-4) mol l(-1)). Furthermore, indomethacin (10(-5) mol l(-1)) did not affect the nicotine-mediated dilation, suggesting that prostaglandins are not involved in the response. Finally, the calcium ionophore A23187 (3 x 10(-6) mol l(-1)) caused an endothelium-dependent dilation that was abolished in the presence of indomethacin. We propose the absence of an endothelial NO system in eel vasculature and suggest that neurally derived NO contributes to the maintenance of vascular tone in this species. In addition, we suggest that prostaglandins may act as endothelially derived relaxing factors in A. australis.

L. F. Gibson - One of the best experts on this subject based on the ideXlab platform.

  • control of saprolegniosis in the eel Anguilla australis richardson by aeromonas media strain a199
    Aquaculture, 2004
    Co-Authors: M. J. Lategan, Fraser R. Torpy, L. F. Gibson
    Abstract:

    The probiotic activity of Aeromonas media strain A199 for the control of saprolegniosis at low water temperatures was trialled in a laboratory-based challenge using the eel, Anguilla australis Richardson. Eels were challenged in the presence of a physiological and physical stress akin to that preceding the winter outbreaks of saprolegniosis in farms. The concentration of Saprolegnia parasitica cysts was 9±1.2 cysts/ml for the initial challenge. Morbidity due to saprolegniosis was low, at 27% in A199-treated tanks, in comparison to the 44% recorded for the non-treated control tanks. The subsequent addition of live A199 cultures to the treatment tanks resulted in the recovery of fish suffering from saprolegniosis (p=0.038), with temperature appearing to be a factor regulating the effect of A199. It was concluded that A199 is a potential agent for the management of winter outbreaks of saprolegniosis in eels.

  • Control of saprolegniosis in the eel Anguilla australis Richardson, by Aeromonas media strain A199
    'Elsevier BV', 2004
    Co-Authors: M. J. Lategan, Torpy Fr, L. F. Gibson
    Abstract:

    The probiotic activity of Aeromonas media strain A199 for the control of saprolegniosis at low water temperatures was trialled in a laboratory-based challenge using the eel, Anguilla australis Richardson. Eels were challenged in the presence of a physiological and physical stress akin to that preceding the winter outbreaks of saprolegniosis in farms. The concentration of Saprolegnia parasitica cysts was 9±1.2 cysts/ml for the initial challenge. Morbidity due to saprolegniosis was low, at 27% in A199-treated tanks, in comparison to the 44% recorded for the non-treated control tanks. The subsequent addition of live A199 cultures to the treatment tanks resulted in the recovery of fish suffering from saprolegniosis (p=0.038), with temperature appearing to be a factor regulating the effect of A199. It was concluded that A199 is a potential agent for the management of winter outbreaks of saprolegniosis in eels. © 2004 Elsevier B.V. All rights reserved

  • antagonistic activity of aeromonas media strain a199 against saprolegnia sp an opportunistic pathogen of the eel Anguilla australis richardson
    Journal of Fish Diseases, 2003
    Co-Authors: M. J. Lategan, L. F. Gibson
    Abstract:

    A bacteriocin-like inhibitory substance (BLIS) produced by Aeromonas media strain A199 inhibited the growth of Saprolegnia sp. in vitro, an opportunistic pathogen isolated from affected eels, Anguilla australis (Richardson). The presence of BLIS in solid media inhibited the growth of the vegetative state of the aquatic mould as well as the germination of cysts. Uninhibited growth was, however, observed in the presence of inactive BLIS, suggesting that the in vitro antagonism derived from the BLIS of A199. In four independent in vivo tank observations of fish affected with saprolegniosis, the daily addition of A199 to tank water contributed to the subsequent swift recovery of affected hosts from invasion by this opportunistic pathogen.

Brett L Jennings - One of the best experts on this subject based on the ideXlab platform.

  • nitric oxide control of the dorsal aorta and the intestinal vein of the australian short finned eel Anguilla australis
    The Journal of Experimental Biology, 2004
    Co-Authors: Brett L Jennings, Bradley Randal Scott Broughton, John A. Donald
    Abstract:

    SUMMARY This study investigated the mechanisms by which nitric oxide (NO) regulates the dorsal aorta and the intestinal vein of the Australian short-finned eel Anguilla australis . NADPH diaphorase histochemistry and immunohistochemistry using a mammalian endothelial nitric oxide synthase (NOS) antibody could not demonstrate NOS in the endothelium of either blood vessel; however, NOS could be readily demonstrated in the endothelium of the rat aorta that was used as a control. Both blood vessels contained NADPH diaphorase positive nerve fibres and nerve bundles, and immunohistochemistry using a neural NOS antibody showed a similar distribution of neural NOS immunoreactivity in the perivascular nerves. In vitro organ bath physiology showed that a NO/soluble guanylyl cyclase (GC) system is present in the dorsal aorta and the intestinal vein, since the soluble GC inhibitor oxadiazole quinoxalin-1 (ODQ; 10 –5 mol l –1 ) completely abolished the vasodilatory effect of the NO donor, sodium nitroprusside (SNP; 10 –4 mol l –1 ). In addition, nicotine (3×10 –4 mol l –1 ) mediated a vasodilation that was not affected by removal of the endothelium. The nicotine-mediated dilation was blocked by the NOS inhibitor, N ω -nitro-l-arginine (l-NNA; 10 –4 mol l –1 ), and ODQ (10 –5 mol l –1 ). More specifically, the neural NOS inhibitor, N ω -propyl-l-arginine (10 –5 mol l –1 ), significantly decreased the dilation induced by nicotine (3×10 –4 mol l –1 ). Furthermore, indomethacin (10 –5 mol l –1 ) did not affect the nicotine-mediated dilation, suggesting that prostaglandins are not involved in the response. Finally, the calcium ionophore A23187 (3×10 –6 mol l –1 ) caused an endothelium-dependent dilation that was abolished in the presence of indomethacin. We propose the absence of an endothelial NO system in eel vasculature and suggest that neurally derived NO contributes to the maintenance of vascular tone in this species. In addition, we suggest that prostaglandins may act as endothelially derived relaxing factors in A. australis .

  • nitric oxide control of the dorsal aorta and the intestinal vein of the australian short finned eel Anguilla australis
    The Journal of Experimental Biology, 2004
    Co-Authors: Brett L Jennings, Bradley Randal Scott Broughton, John A. Donald
    Abstract:

    This study investigated the mechanisms by which nitric oxide (NO) regulates the dorsal aorta and the intestinal vein of the Australian short-finned eel Anguilla australis. NADPH diaphorase histochemistry and immunohistochemistry using a mammalian endothelial nitric oxide synthase (NOS) antibody could not demonstrate NOS in the endothelium of either blood vessel; however, NOS could be readily demonstrated in the endothelium of the rat aorta that was used as a control. Both blood vessels contained NADPH diaphorase positive nerve fibres and nerve bundles, and immunohistochemistry using a neural NOS antibody showed a similar distribution of neural NOS immunoreactivity in the perivascular nerves. In vitro organ bath physiology showed that a NO/soluble guanylyl cyclase (GC) system is present in the dorsal aorta and the intestinal vein, since the soluble GC inhibitor oxadiazole quinoxalin-1 (ODQ; 10(-5) mol l(-1)) completely abolished the vasodilatory effect of the NO donor, sodium nitroprusside (SNP; 10(-4) mol l(-1)). In addition, nicotine (3 x 10(-4) mol l(-1)) mediated a vasodilation that was not affected by removal of the endothelium. The nicotine-mediated dilation was blocked by the NOS inhibitor, N(omega)-nitro-L-arginine (L-NNA; 10(-4) mol l(-1)), and ODQ (10(-5) mol l(-1)). More specifically, the neural NOS inhibitor, N(omega)-propyl-L-arginine (10(-5) mol l(-1)), significantly decreased the dilation induced by nicotine (3 x 10(-4) mol l(-1)). Furthermore, indomethacin (10(-5) mol l(-1)) did not affect the nicotine-mediated dilation, suggesting that prostaglandins are not involved in the response. Finally, the calcium ionophore A23187 (3 x 10(-6) mol l(-1)) caused an endothelium-dependent dilation that was abolished in the presence of indomethacin. We propose the absence of an endothelial NO system in eel vasculature and suggest that neurally derived NO contributes to the maintenance of vascular tone in this species. In addition, we suggest that prostaglandins may act as endothelially derived relaxing factors in A. australis.

Graham Young - One of the best experts on this subject based on the ideXlab platform.

  • 11 ketotestosterone and igf i increase the size of previtellogenic oocytes from shortfinned eel Anguilla australis in vitro
    Reproduction, 2007
    Co-Authors: Mark P Lokman, Kataraina A N George, Sean L Divers, Michael Algie, Graham Young
    Abstract:

    Previtellogenic ovarian fragments from eel, Anguilla australis, were cultured in vitro in a chemically defined medium containing steroids and/or peptide hormones for 18 days in order to investigate their involvement in control of early oocyte growth. 11-Ketotestosterone (11-KT), but not estradiol-17beta, induced a significant 10-20% increase in diameters of previtellogenic oocytes and oocyte nuclei in a dose-dependent manner. Effects were greatest for 100 nM 11-KT, a dose that is within the physiological range seen in very early vitellogenic eels in the wild. The effect was not accompanied by obvious ultrastructural changes in the oocytes other than an apparent increase in nuclear size. Similarly, treatment with recombinant human IGF-I resulted in increased oocyte diameters, whereas no such effect was seen after treatment with heterologous insulin, GH, leptin, or human chorionic gonadotropin. Interestingly, lipid supplementation also resulted in an increase in oocyte diameter, and greater radioactivity in ovarian explants following incubation with (14)C-triglycerides and 11-KT, but not FSH, suggesting that the androgen may play a role in lipid accumulation into the oocyte. Our results implicate hormones from both the reproductive and the metabolic axes in control of previtellogenic oocyte growth in a teleost fish.

  • effects of steroid and peptide hormones on in vitro growth of previtellogenic oocytes from eel Anguilla australis
    Fish Physiology and Biochemistry, 2003
    Co-Authors: Mark P Lokman, Kataraina A N George, Graham Young
    Abstract:

    To investigate the hormonal control of previtellogenic oocyte growth in eel, Anguilla australis, ovarian explants were incubated with or without different doses of steroid (11-ketotestosterone, estradiol-17β) or peptide hormones (ovine growth hormone, human insulin-like growth factor-I, human chorionic gonadotropin, porcine insulin) for 18 days. 11-Ketotestosterone, but not estradiol-17β, significantly increased oocyte diameters at physiological doses (30–100 ng/mL). Similarly, incubation with insulin-like growth factor-I, but not any of the other peptide hormones tested, resulted in increased oocyte diameters at high physiological doses (≥30 ng/mL). These data implicate androgens and hormones from the metabolic axis in mediating growth of previtellogenic oocytes in eel.

  • 11-Ketotestosterone induces silvering-related changes in immature female short-finned eels, Anguilla australis.
    Comparative Biochemistry and Physiology A-molecular & Integrative Physiology, 2001
    Co-Authors: Detlef H. Rohr, Peter S. Davie, P. Mark Lokman, Graham Young
    Abstract:

    Abstract The developmental transition from a residential, immature ‘yellow’ eel to a migratory, maturing adult ‘silver’ eel is accompanied by many morphological changes that appear to be under endocrine control. High circulating levels of the teleost, and usually male-specific, androgen 11-ketotestosterone (11-KT) are found in migrating female short-finned eels, Anguilla australis . We examined the role of this steroid in silvering by implanting immature, female short-finned eels either with blank vehicles or with vehicles containing 11-KT. Six weeks after they had received the implants, eels treated with 11-KT had developed ‘chisel-shaped’ snouts and black pectoral fins with tapered ends, and the size of their eyes had increased significantly. 11-KT treated eels had a thicker dermis than control eels and an epidermis with fewer or no mucous cells. Ventricular mass at the end of the experiment was two-fold larger than in control eels. 11-KT treated eels also had larger livers and gonads. Ovaries contained predominantly cortical alveolus stage III oocytes, as opposed to the smaller gonads of control eels containing previtellogenic stage II oocytes. All of these changes correspond to changes during the developmental transition from yellow to silver eels in the wild. This demonstrates that silvering in eels is under endocrine control and that the presumed male-specific steroid 11-KT is capable of inducing silvering-related changes in a female teleost. We discuss how species-specific responses to 11-KT may differ depending on tissue-specific androgen receptor abundance and how a dual demand on liver function can explain the apparently positive effects of 11-KT on liver growth.

  • induced spawning and early ontogeny of new zealand freshwater eels Anguilla dieffenbachii and a australis
    New Zealand Journal of Marine and Freshwater Research, 2000
    Co-Authors: Mark P Lokman, Graham Young
    Abstract:

    Abstract Knowledge on the reproductive biology of New Zealand freshwater eels (Anguilla australis Richardson and A. dieffenbachii Gray) is limited to changes associated with gametogenesis in the adults; details pertaining to embryonic and larval development are not available, but are of obvious benefit for a better understanding of the eel life history and for artificial propagation programmes, whether for aquaculture or conservation purposes. Therefore, eels were artificially matured and eggs fertilised with sperm from either species to investigate early ontogeny. Only eggs from A. australis hatched successfully after c. 45 h of incubation at room temperature, yielding larvae of c. 2.5 mm in length. Larvae survived for up to 5 days, by which time they had attained a length of 5.3 mm. Scanning electron microscopy of 1‐ and 2‐day‐old larvae revealed the presence of prominent free neuromasts on the head and along the flanks. Although hatching of New Zealand eels was achieved for the first time, it is strong...