Reviews in Aquaculture Reproduction of greater amberjack (Seriola dumerili) and other members of the family Carangidae Journal: Reviews in Aquaculture Manuscript ID RAQ-11-20-0300.R3 Manuscript Type: Review Date Submitted byFor the Review Only n/a Author: Complete List of Authors: Corriero, Aldo; University of Bari Aldo Moro, Emergency and Organ Transplantation, Section of Veterinary Clinics and Animal Production Wylie, Matthew; New Zealand Institute for Plant and Food Research Ltd, Seafood Production Unit Nyuji, Mitsuo ; Japan Fisheries Research and Education Agency, Fisheries Resources Institute Zupa, Rosa; University of Bari Aldo Moro, Emergency and Organ Transplantation, Section of Veterinary Clinics and Animal Production Mylonas, Constantinos ; Hellenic Center for Marine Research, Institute of Marine Biology, Biotechnology and Aquaculture fish reproduction, fish rearing in captivity, fish reproductive dysfunction, Keywords: fish gametogenesis, fish reproduction control, Carangidae Page 1 of 84 Reviews in Aquaculture Reproduction of greater amberjack (Seriola dumerili) and other members of the family 2 Carangidae 4 Corriero Aldo1, Wylie Matthew J.2, Nyuji Mitsuo3, Zupa Rosa1 and Mylonas Constantinos C.4 6 1 Department of Emergency and Organ Transplantation, Section of Veterinary Clinics and Animal Production, University of Bari Aldo Moro, Valenzano (Bari), Italy For Review Only 8 2 The New Zealand Institute for Plant and Food Research Limited, Seafood Production Unit, 293–297 Port Nelson, Nelson 7010, New Zealand 10 3 Fisheries Resources Institute, Japan Fisheries Research and Education Agency, Yokohama, Japan 12 4 Institute of Marine Biology, Biotechnology and Aquaculture, Hellenic Center for Marine Research, Heraklion, Crete, Greece 14 Correspondence: Constantinos C. Mylonas, Institute of Marine Biology, Biotechnology and 16 Aquaculture, Hellenic Center for Marine Research, P.O. box 2214, Heraklion, 71003, Crete, Greece. Email: [email protected] 1 Reviews in Aquaculture Page 2 of 84 18 Table of Contents 20 ABSTRACT .......................................................................................................................................................3 1. INTRODUCTION...........................................................................................................................................4 22 2. REPRODUCTIVE BIOLOGY OF THE FAMILY CARANGIDAE..............................................................................7 2.1. GREATER AMBERJACK .........................................................................................................................................8 24 2.1.1. Ovary structure and oogenesis..............................................................................................................8 2.1.2. Testis structure and spermatogenesis.................................................................................................10 26 2.1.3. Sex differentiation, sexual maturity and reproductive cycles..............................................................10 2.1.4. Endocrine control of reproduction ......................................................................................................14 28 2.2. OTHER SERIOLA SPP. ........................................................................................................................................15 2.2.1. Oogenesis and spermatogenesis.........................................................................................................15 30 2.2.2. Sexual maturity and reproductive cycles.............................................................................................16 2.2.3. Endrocrine controlFor of reproduction Review .....................................................................................................20 Only 32 2.3. OTHER CARANGIDS ..........................................................................................................................................23 2.3.1. Oogenesis and spermatogenesis.........................................................................................................23 34 2.3.2. Sexual maturity and reproductive cycles.............................................................................................24 3. REPRODUCTIVE FUNCTION IN CAPTIVITY AND SPONTANEOUS SPAWNING ...............................................26 36 3.1. GREATER AMBERJACK .......................................................................................................................................26 3.2. OTHER SERIOLA SPP. ........................................................................................................................................32 38 3.3. OTHER CARANGIDS ..........................................................................................................................................34 4. HORMONAL MANIPULATIONS OF REPRODUCTIVE FUNCTION AND INDUCED SPAWNING .........................37 40 4.1. GREATER AMBERJACK .......................................................................................................................................37 4.2. OTHER SERIOLA SPP. ........................................................................................................................................44 42 4.3. OTHER CARANGIDS ..........................................................................................................................................47 5. CONCLUDING REMARKS ............................................................................................................................50 44 6. REFERENCES ..............................................................................................................................................54 2 Page 3 of 84 Reviews in Aquaculture 46 Abstract The family Carangidae contains several species of aquaculture interest, including the 48 amberjacks, yellowtails and trevallies. Among them, the greater amberjack (Seriola dumerili), the Japanese amberjack or yellowtail (Seriola quinqueradiata) and the yellowtail kingfish 50 (Seriola lalandi) are considered the species with the highest potential for commercial aquaculture. Understanding the reproductive physiology-biology, spawning kinetics and 52 production characteristics in captivity is of utmost importance for the domestication of any animal, and developing broodstock management methods and therapies to optimize egg 54 production and overcomeFor potential Review reproductive dysfunctions Only are essential. The present article reviews the available literature on the reproductive biology of the Carangidae species of 56 interest for the aquaculture industry, both in the wild and under farming conditions. The reproductive traits of wild and farmed fish, whenever available, were compared in order to 58 improve the understanding of the reproductive dysfunctions occurring in captivity. Finally, the hormonal maturation and spawning induction protocols examined so far to ameliorate the 60 reproductive dysfunctions and obtain fertile gametes are summarized, and their effectiveness in the different rearing conditions are discussed. 62 3 Reviews in Aquaculture Page 4 of 84 64 1. Introduction The genus Seriola (family Carangidae) includes 12 species that are distributed in all 66 tropical and temperate waters (Table 1). Some of them have been notable species for aquaculture worldwide, while other carangid species have also been considered as potential 68 cultured species. Among them, the greater amberjack (Seriola dumerili) has attracted significant interest in Europe and Japan since the 1990s, because of its fast growth and 70 cosmopolitan distribution and appreciation. However, failure to control reproduction in captivity has prevented its commercial production (Ottolenghi et al. 2004). With the need to 72 diversify aquaculture worldwide,For aReview renewed interest inOnly studying the reproductive biology of greater amberjack and developing methods to control egg production in captivity has emerged 74 (Nyuji et al. 2016; Zupa et al. 2017b), and a significant body of information has been produced in recent years for this species. For other members of the Seriola family, such as the 76 Japanese amberjack or yellowtail (S. quinqueradiata), which is a very important fishery resource in Japan, aquaculture research had already begun in the 1970s (Kagawa 1992; 78 Nakada 2002; Yamazaki et al. 2002). Following the work on yellowtail, more recently the reproductive biology and physiology of yellowtail kingfish (S. lalandi) have also been studied 80 (Nakada 2002) and this species is currently reared commercially worldwide. The greater amberjack is a cosmopolitan species found throughout the temperate zone, 82 where it spawns naturally from February to April in the Gulf of Mexico (Wells & Rooker 2004), from April to June in Japan (Kawabe et al. 1996, 1998; Nyuji et al. 2016), from May 84 to July in the Mediterranean Sea (Marino et al. 1995a) and from April to October in the Canary Islands (Jerez et al. 2006). Yellowtail kingfish has been considered to exist as 86 geographically separate populations and its aquaculture has spread from Japan to Australia, Chile, Mexico, and California (Sicuro & Luzzana 2016). However, Martinez-Takeshita et al. 88 (2015) recently proposed that these different populations are actually genetically distinct 4 Page 5 of 84 Reviews in Aquaculture species and named them using the following names: yellowtail kingfish has been reserved for 90 fish in the Southern Hemisphere,