偏顶蛤的性腺发育和生殖周期
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摘要
为了探讨偏顶蛤的繁殖规律,自2013年10月至2014年9月,每月采集大连海区野生偏顶蛤(Modiolus modiolus)样品1次,测定采样点水温,采用组织学方法分析其性腺的季节变化。结果表明,偏顶蛤性腺发育以1年为1个周期,可分为增殖期、生长期、成熟期、排放期、休止期5个时期。偏顶蛤卵子的发生可分为卵原细胞期、生长初期、生长中期、生长后期和成熟期。雄性生殖细胞的发生经历了精原细胞、初级精母细胞、次级精母细胞、精细胞和精子等时期,精原细胞分为A型和B型2种类型。该海区偏顶蛤性腺指数和肥满度最低值均出现在1月,分别为17.54%和18.23%;二者最高值均出现在8月,分别为32.25%和38.43%。偏顶蛤繁殖期为6—11月,繁殖期水温在16.8~22.1℃;雌雄性腺发育基本同步,雄性发育略快,雌雄异体,二者性比接近1∶1。本研究为偏顶蛤的天然资源保护及大规模人工增养殖的开展提供了基础资料。
The wild Modiolus modiolus resource has experienced a dramatic decline in abundance over the past few decades because of overfishing, declining seawater quality, and a lack of proper protection. The reproductive cycle and gonadal development of M. modiolus from the coast of Heishijiao, Dalian city were investigated from October 2013 to September 2014 to develop management strategies and optimize hatchery-based seed production. Specimens were collected monthly, and seawater column temperatures were recorded simultaneously. Shell size, total weight, tissue weight, and gonad weight of M. modiolus were measured to calculate fatness and the gonadosomatic index, and histological techniques were used to analyze the seasonal gonadal cycle. The histological analysis showed a unimodal gametogenic cycle with one spawning season(August–October) annually. Although shellfish gonadal development stages have no unified standard, M. modiolus gonadal development was divided into five stages according to previous studies: the proliferating stage, growing stage, maturing stage, spawning stage, and resting stage. Oogenesis was divided into the oogonial stage, early growth stage, middle growth stage, late growth stage, and mature stage, according to the oocyte developmental characteristics and the proportions of different oocyte developmental stages in the follicle. Spermatogenesis of M. modiolus included five stages of spermatogonium, primary spermatocyte, secondary spermatocyte, spermatid, and spermatozoon. Spermatogonia were morphologically divided into types A and B. Type A spermatogonia distributed in all developmental stages, and their nuclear chromatin was arranged loosely. Type B spermatogonia occurred during the proliferating stage, and the nuclear chromatin was concentrated. Thus, types A and B spermatogonia were considered differentiated cells and stem cells, respectively. The peak gonadosomatic index of 32.25% and fatness of 38.43% were observed in January, and the maximum values for both occurred in August. The breeding season was from June to November, when the average seawater temperature was 16.8–22.1℃. M. modiolus gonads matured by the end of May, but they did not spawn until August of the rainy season, indicating that the mature gonads of the parental shellfish only provided a material basis for reproduction and that the reproductive process requires particular conditions for induction, such as temperature and salinity. The developmental rate of the male gonad was slightly faster than that of the female. Gonochorism existed, and the female to male ratio was 1︰1. These results provide basic information about M. modiolus reproduction to help with natural resources protection, and will aid in developing a large-scale artificial breeding program.
The wild Modiolus modiolus resource has experienced a dramatic decline in abundance over the past few decades because of overfishing, declining seawater quality, and a lack of proper protection. The reproductive cycle and gonadal development of M. modiolus from the coast of Heishijiao, Dalian city were investigated from October 2013 to September 2014 to develop management strategies and optimize hatchery-based seed production. Specimens were collected monthly, and seawater column temperatures were recorded simultaneously. Shell size, total weight, tissue weight, and gonad weight of M. modiolus were measured to calculate fatness and the gonadosomatic index, and histological techniques were used to analyze the seasonal gonadal cycle. The histological analysis showed a unimodal gametogenic cycle with one spawning season(August–October) annually. Although shellfish gonadal development stages have no unified standard, M. modiolus gonadal development was divided into five stages according to previous studies: the proliferating stage, growing stage, maturing stage, spawning stage, and resting stage. Oogenesis was divided into the oogonial stage, early growth stage, middle growth stage, late growth stage, and mature stage, according to the oocyte developmental characteristics and the proportions of different oocyte developmental stages in the follicle. Spermatogenesis of M. modiolus included five stages of spermatogonium, primary spermatocyte, secondary spermatocyte, spermatid, and spermatozoon. Spermatogonia were morphologically divided into types A and B. Type A spermatogonia distributed in all developmental stages, and their nuclear chromatin was arranged loosely. Type B spermatogonia occurred during the proliferating stage, and the nuclear chromatin was concentrated. Thus, types A and B spermatogonia were considered differentiated cells and stem cells, respectively. The peak gonadosomatic index of 32.25% and fatness of 38.43% were observed in January, and the maximum values for both occurred in August. The breeding season was from June to November, when the average seawater temperature was 16.8–22.1℃. M. modiolus gonads matured by the end of May, but they did not spawn until August of the rainy season, indicating that the mature gonads of the parental shellfish only provided a material basis for reproduction and that the reproductive process requires particular conditions for induction, such as temperature and salinity. The developmental rate of the male gonad was slightly faster than that of the female. Gonochorism existed, and the female to male ratio was 1︰1. These results provide basic information about M. modiolus reproduction to help with natural resources protection, and will aid in developing a large-scale artificial breeding program.
引文
[1]Song J,Cheng L,Chang Y Q,et al.Analysis and evaluation of nutrient composition in different tissues of Modiolus modiolus[J].Journal of Dalian Ocean University,2014,29(2):167–170.[宋坚,程龙,常亚青,等.偏顶蛤不同组织营养成分的分析及评价[J].大连海洋大学学报,2014,29(2):167–170.]
[2]Qi Z Y,Ma X T,Wang Z R,et al.Mollusca of Huanghai and Bohai[M].Beijing:Agriculture Press,1987:164–166.[齐钟彦,马绣同,王祯瑞,等.黄渤海的软体动物[M].北京:农业出版社,1987:164–166.]
[3]Chang K M,Wu J F.Study on artificial propagation of mussel Mytilus coruscus[J].South China Fisheries Science,2007,3(3):26–30.[常抗美,吴剑锋.厚壳贻贝人工繁殖技术的研究[J].南方水产,2007,3(3):26–30.]
[4]Firth L B,Knights A M,Bell S S.Air temperature and winter mortality:Implications for the persistence of the invasive mussel,Perna viridis in the intertidal zone of the south-eastern United States[J].J Exp Mar Biol Ecol,2011,400(1):250–256.
[5]Li T W,Li C M.Histology and ultrastructure of male reproductive system of Mytilus edulis Linnaeus(Mollusca:Lamellibranchia)[J].Zoological Research,1999,20(3):168–171.[李太武,李春茂.贻贝雄性生殖系统的组织学和超微结构[J].动物学研究,1999,20(3):168–171.]
[6]Lin Z H,Shan L Z,Chai X L,et al.Study on the gonad development and reproductive cycle of Meretrix meretrix Linnaeus[J].Journal of Fisheries of China,2004,28(5):510–514.[林志华,单乐州,柴雪良,等.文蛤的性腺发育和生殖周期[J].水产学报,2004,28(5):510–514.]
[7]Lin Z H,Shan L Z,Chai X L,et al.The reproductive biology of hard clam Mercenaria mercenaria(Linnaeus,1758)[J].Oceanologia et Limnologia Sinica,2005,36(5):430–436.[林志华,单乐洲,柴雪良,等.硬壳蛤(Mercenaria mercenaria)繁殖生物学研究[J].海洋与湖沼,2005,36(5):430–436.]
[8]Liao C Y,Xu Y F,Wang Y L.Reproductive cycle of the scallop Chlamys Farreri(Jones et Etpreston)at Qingdao[J].Journal of Fisheries of China,2005,7(1):1–13.[廖承义,徐应馥,王远隆.栉孔扇贝的生殖周期[J].水产学报,2005,7(1):1–13.]
[9]Cao F J,Liu Y,Zhang C F,et al.Studies on the sex gonad development and reproductive cycle of Lutraria sieboldii[J].Oceanologia et Limnologia Sinica,2012,43(5):976–982.[曹伏君,刘永,张春芳,等.施氏獭蛤(Lutraria sieboldii)性腺发育和生殖周期的研究[J].海洋与湖沼,2012,43(5):976–982.]
[10]Ruan F T,Gao S,Li L,et al.Gonad development and biochemical composition in the ark shell Scapharca broughtoniifrom coast of Shandong Peninsula[J].Journal of Fisheries of China,2014,38(1):47–55.[阮飞腾,高森,李莉,等.山东沿海魁蚶繁殖周期与生化成分的周年变化[J].水产学报,2014,38(1):47–55.]
[11]Gao Y M,Tian B,Yu Y G,et al.The gonadal development and reproductive cycle of Japanese scallop Patinopecten yessoensis in Tahe Bay in Dalian[J].Journal of Dalian Ocean University,2007,22(5):335–339.[高悦勉,田斌,于永刚,等.大连塔河湾海区虾夷扇贝的性腺发育与繁殖规律[J].大连水产学院学报,2007,22(5):335–339.]
[12]Zhao Z J,Li F X,Ke C H.Studies on the sex gonad development and reproductive cycle of clam Paphia Undulata[J].Journal of Fisheries of China,1991,15(1):1–8.[赵志江,李复雪,柯才焕.波纹巴非蛤的性腺发育和生殖周期[J].水产学报,1991,15(1):1–8.]
[13]Lv H,Yu D Z.Conadial development of Pacific oyster(Crassostrea gigas)and the relationship between condition index and accumulative temperature[J].Journal of Fishery Sciences of China,1998,5(1):18–24.[吕豪,于德州.太平洋牡蛎控温条件下的性腺发育,条件指数与积温的关系[J].中国水产科学,1998,5(1):18–24.]
[14]Chang Y Q,Song J,Wang G D,et al.Seeds breeding of hard clam Mercenaria mercenaria[J].Journal of Fishery Sciences of China,2002,9(1):43–47.[常亚青,宋坚,王国栋,等.硬壳蛤的人工育苗技术[J].中国水产科学,2002,9(1):43–47.]
[15]Li X,Liang J,Yan X W,et al.The relationship between water temperature and periodic variation of histology with gonad development of Manila clam Ruditapes philippinarum in different redions[J].Journal of Fisheries of China,2007,31(6):853–859.[李霞,梁峻,闫喜武,等.人工控温促熟条件下不同海域菲律宾蛤仔的性腺发育[J].水产学报,2007,31(6):853–859.]
[16]Wang R C,Wang Z P,Zhang J Z,et al.Science of Marine Shellfish Culture[M].Qingdao:Ocean University of China Press,2008.[王如才,王昭萍,张建中,等.海水贝类养殖学[M].青岛:中国海洋大学出版社,2008.]
[17]Wu H L,Wang H Y.Developmental stage of the gonad of the clam Paphia(Paratapes)undulata[J].Natural Science Journal of Hainan University,2002,20(1):41–47.[吴洪流,王红勇.波纹巴非蛤性腺发育分期的研究[J].海南大学学报:自然科学版,2002,20(1):41–47.]
[18]Yan B L,Xu X H,Zheng J S,et al.Study on the development of sex gonad and reproductive cycle of Scapharca subcrenata(Lischke)[J].Transactions of Oceanology and Limnology,2006(4):92–98.[阎斌伦,许星鸿,郑家声,等.毛蚶的性腺发育和生殖周期[J].海洋湖沼通报,2006(4):92–98.]
[19]Wourms J P.Reproduction of marine invertebrates[M].Pacific Grove:The Boxwood Press,1987:50–179.
[20]Sastry A N.Temperature effects in the reproduction of the bay scallop Argopecten irradians[J].Biol Bull,1966,130:118–143.
[21]Loosanoff V L,Davis H C.Temperature requirements for maturation of gonads of Northern oyster[J].Biol Bull,1952,103:80–96.
[22]Wu H L,Wu C Y,Chen X F,et al.Histology studies on the female gonad of Paphia(Paratapes)undulata(Born)[J].Natural Science Journal of Hainan University,2000,18(2):170–174.[吴洪流,伍春雨,陈雪芬,等.波纹巴非蛤雄性生殖腺的组织学观察[J].海南大学学报:自然科学版,2000,18(2):170–174.]
[2]Qi Z Y,Ma X T,Wang Z R,et al.Mollusca of Huanghai and Bohai[M].Beijing:Agriculture Press,1987:164–166.[齐钟彦,马绣同,王祯瑞,等.黄渤海的软体动物[M].北京:农业出版社,1987:164–166.]
[3]Chang K M,Wu J F.Study on artificial propagation of mussel Mytilus coruscus[J].South China Fisheries Science,2007,3(3):26–30.[常抗美,吴剑锋.厚壳贻贝人工繁殖技术的研究[J].南方水产,2007,3(3):26–30.]
[4]Firth L B,Knights A M,Bell S S.Air temperature and winter mortality:Implications for the persistence of the invasive mussel,Perna viridis in the intertidal zone of the south-eastern United States[J].J Exp Mar Biol Ecol,2011,400(1):250–256.
[5]Li T W,Li C M.Histology and ultrastructure of male reproductive system of Mytilus edulis Linnaeus(Mollusca:Lamellibranchia)[J].Zoological Research,1999,20(3):168–171.[李太武,李春茂.贻贝雄性生殖系统的组织学和超微结构[J].动物学研究,1999,20(3):168–171.]
[6]Lin Z H,Shan L Z,Chai X L,et al.Study on the gonad development and reproductive cycle of Meretrix meretrix Linnaeus[J].Journal of Fisheries of China,2004,28(5):510–514.[林志华,单乐州,柴雪良,等.文蛤的性腺发育和生殖周期[J].水产学报,2004,28(5):510–514.]
[7]Lin Z H,Shan L Z,Chai X L,et al.The reproductive biology of hard clam Mercenaria mercenaria(Linnaeus,1758)[J].Oceanologia et Limnologia Sinica,2005,36(5):430–436.[林志华,单乐洲,柴雪良,等.硬壳蛤(Mercenaria mercenaria)繁殖生物学研究[J].海洋与湖沼,2005,36(5):430–436.]
[8]Liao C Y,Xu Y F,Wang Y L.Reproductive cycle of the scallop Chlamys Farreri(Jones et Etpreston)at Qingdao[J].Journal of Fisheries of China,2005,7(1):1–13.[廖承义,徐应馥,王远隆.栉孔扇贝的生殖周期[J].水产学报,2005,7(1):1–13.]
[9]Cao F J,Liu Y,Zhang C F,et al.Studies on the sex gonad development and reproductive cycle of Lutraria sieboldii[J].Oceanologia et Limnologia Sinica,2012,43(5):976–982.[曹伏君,刘永,张春芳,等.施氏獭蛤(Lutraria sieboldii)性腺发育和生殖周期的研究[J].海洋与湖沼,2012,43(5):976–982.]
[10]Ruan F T,Gao S,Li L,et al.Gonad development and biochemical composition in the ark shell Scapharca broughtoniifrom coast of Shandong Peninsula[J].Journal of Fisheries of China,2014,38(1):47–55.[阮飞腾,高森,李莉,等.山东沿海魁蚶繁殖周期与生化成分的周年变化[J].水产学报,2014,38(1):47–55.]
[11]Gao Y M,Tian B,Yu Y G,et al.The gonadal development and reproductive cycle of Japanese scallop Patinopecten yessoensis in Tahe Bay in Dalian[J].Journal of Dalian Ocean University,2007,22(5):335–339.[高悦勉,田斌,于永刚,等.大连塔河湾海区虾夷扇贝的性腺发育与繁殖规律[J].大连水产学院学报,2007,22(5):335–339.]
[12]Zhao Z J,Li F X,Ke C H.Studies on the sex gonad development and reproductive cycle of clam Paphia Undulata[J].Journal of Fisheries of China,1991,15(1):1–8.[赵志江,李复雪,柯才焕.波纹巴非蛤的性腺发育和生殖周期[J].水产学报,1991,15(1):1–8.]
[13]Lv H,Yu D Z.Conadial development of Pacific oyster(Crassostrea gigas)and the relationship between condition index and accumulative temperature[J].Journal of Fishery Sciences of China,1998,5(1):18–24.[吕豪,于德州.太平洋牡蛎控温条件下的性腺发育,条件指数与积温的关系[J].中国水产科学,1998,5(1):18–24.]
[14]Chang Y Q,Song J,Wang G D,et al.Seeds breeding of hard clam Mercenaria mercenaria[J].Journal of Fishery Sciences of China,2002,9(1):43–47.[常亚青,宋坚,王国栋,等.硬壳蛤的人工育苗技术[J].中国水产科学,2002,9(1):43–47.]
[15]Li X,Liang J,Yan X W,et al.The relationship between water temperature and periodic variation of histology with gonad development of Manila clam Ruditapes philippinarum in different redions[J].Journal of Fisheries of China,2007,31(6):853–859.[李霞,梁峻,闫喜武,等.人工控温促熟条件下不同海域菲律宾蛤仔的性腺发育[J].水产学报,2007,31(6):853–859.]
[16]Wang R C,Wang Z P,Zhang J Z,et al.Science of Marine Shellfish Culture[M].Qingdao:Ocean University of China Press,2008.[王如才,王昭萍,张建中,等.海水贝类养殖学[M].青岛:中国海洋大学出版社,2008.]
[17]Wu H L,Wang H Y.Developmental stage of the gonad of the clam Paphia(Paratapes)undulata[J].Natural Science Journal of Hainan University,2002,20(1):41–47.[吴洪流,王红勇.波纹巴非蛤性腺发育分期的研究[J].海南大学学报:自然科学版,2002,20(1):41–47.]
[18]Yan B L,Xu X H,Zheng J S,et al.Study on the development of sex gonad and reproductive cycle of Scapharca subcrenata(Lischke)[J].Transactions of Oceanology and Limnology,2006(4):92–98.[阎斌伦,许星鸿,郑家声,等.毛蚶的性腺发育和生殖周期[J].海洋湖沼通报,2006(4):92–98.]
[19]Wourms J P.Reproduction of marine invertebrates[M].Pacific Grove:The Boxwood Press,1987:50–179.
[20]Sastry A N.Temperature effects in the reproduction of the bay scallop Argopecten irradians[J].Biol Bull,1966,130:118–143.
[21]Loosanoff V L,Davis H C.Temperature requirements for maturation of gonads of Northern oyster[J].Biol Bull,1952,103:80–96.
[22]Wu H L,Wu C Y,Chen X F,et al.Histology studies on the female gonad of Paphia(Paratapes)undulata(Born)[J].Natural Science Journal of Hainan University,2000,18(2):170–174.[吴洪流,伍春雨,陈雪芬,等.波纹巴非蛤雄性生殖腺的组织学观察[J].海南大学学报:自然科学版,2000,18(2):170–174.]