雌性阿根廷滑柔鱼性腺发育阶段的脂肪酸组成及其变化
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摘要
阿根廷滑柔鱼是典型的头足类,资源生物量与其生长发育过程中营养物质积累及生殖投入密切相关。为了获知雌性阿根廷滑柔鱼性腺发育各阶段的脂肪酸组成及其变化特点,研究采用氯仿甲醇法和气相色谱法检测分析胴体、消化腺和卵巢的脂肪酸种类及其含量。结果表明,雌性阿根廷滑柔鱼胴体、消化腺和卵巢3个组织的脂肪酸均以多不饱和脂肪酸(PUFA)为主,饱和脂肪酸(SFA)次之,单不饱和脂肪酸(MUFA)的总含量最低。胴体组织的脂肪酸种类最全面,共检测出28种脂肪酸,消化腺和卵巢分别检测出21种和17种脂肪酸。检测发现3个组织中SFA均以C16∶0脂肪酸最为丰富,MUFA以C18∶1n9c和C20∶1脂肪酸为主,PUFA的DHA(C22∶6n3)最丰富,EPA(C20∶5n3)次之。胴体的SFA和MUFA在成熟期达到最大值,在产卵期显著降低;消化腺的PUFA保持相对稳定水平,但是其含量显著低于胴体和卵巢;卵巢持续积累脂肪酸,在产卵期时MUFA和PUFA显著降低。研究表明,雌性阿根廷滑柔鱼胴体、消化腺和性腺3个组织的脂肪酸种类具有组织特殊性,而主要脂肪酸具有一致性。性腺发育过程中,胴体的SFA和MUFA可能转化为能量供给卵巢发育,消化腺的PUFA则向胴体和卵巢转移,卵巢则持续积累脂肪酸供给卵子发生。
Argentinean short-fin squid, Illex argentinus, is an important Cephalopod species, characterized by fast growth, short lifespan and semelparous reproduction. It has been assumed that the stock biomass is closely related to the nutrients acquisition, accumulation and subsequent allocation to reproduction during the sexual maturation. In order to understand the fatty acid composition and its changes during sexual maturation in female Illex argentinus, the fatty acid profiles and their relative content in multiple tissues, namely mantle tissue, digestive gland and ovary, were investigated by the methods of chloroform-methanol and gas chromatography. The results indicated that the polyunsaturated fatty acids(PUFA) were the most abundant fatty acid class for the tissues investigated, followed by saturated fatty acids(SFA) and monounsaturated fatty acids(MUFA). A total of 28 fatty acid profiles were detected in mantle tissue, while digestive gland and ovary were had 21 fatty acid profiless and 17 fatty acid profiles, respectively. It was found that most of SFA content was presented as C16∶0, MUFA content as C18∶1 n9 c and C20∶1, and PUFA content as docosahexaenoic acid(DHA; 22∶6 n3) and eicosapentaenoic acid(EPA; 20∶5 n3). During the sexual maturation, both SFA content and MUFA content in mantle tissue attained the highest level at mature stage, and then decreased significantly at spawning stage, probably indicating these fatty acids were mobilized to fuel reproduction. PUFA content in digestive gland was significantly lower than that in mantle tissue or ovary, even though it maintained a relatively stable status throughout sexual maturation. The ovary continuously accumulated fatty acid content, but showed a significant decrease in MUFA content and PUFA content during the spawning period, possibly suggesting that these fatty acids were increasingly allocated to oogenesis. These evidence indicated that the fatty acid profiles in Illex argentinus showed tissue-specific characteristics, while the major fatty acid profiles were consistent in mantle tissue, digestive gland and ovary. In the mantle tissue, both SFA and MUFA were possibly mobilized to fuel reproductive growth during sexual maturation. In the digestive gland, the PUFA content might be continuously transferred to mantle and ovary with maturation. Regarding fatty acids in the ovary, the content might be continuously accumulated and subsequently used to support oogenesis during the sexual maturation.
Argentinean short-fin squid, Illex argentinus, is an important Cephalopod species, characterized by fast growth, short lifespan and semelparous reproduction. It has been assumed that the stock biomass is closely related to the nutrients acquisition, accumulation and subsequent allocation to reproduction during the sexual maturation. In order to understand the fatty acid composition and its changes during sexual maturation in female Illex argentinus, the fatty acid profiles and their relative content in multiple tissues, namely mantle tissue, digestive gland and ovary, were investigated by the methods of chloroform-methanol and gas chromatography. The results indicated that the polyunsaturated fatty acids(PUFA) were the most abundant fatty acid class for the tissues investigated, followed by saturated fatty acids(SFA) and monounsaturated fatty acids(MUFA). A total of 28 fatty acid profiles were detected in mantle tissue, while digestive gland and ovary were had 21 fatty acid profiless and 17 fatty acid profiles, respectively. It was found that most of SFA content was presented as C16∶0, MUFA content as C18∶1 n9 c and C20∶1, and PUFA content as docosahexaenoic acid(DHA; 22∶6 n3) and eicosapentaenoic acid(EPA; 20∶5 n3). During the sexual maturation, both SFA content and MUFA content in mantle tissue attained the highest level at mature stage, and then decreased significantly at spawning stage, probably indicating these fatty acids were mobilized to fuel reproduction. PUFA content in digestive gland was significantly lower than that in mantle tissue or ovary, even though it maintained a relatively stable status throughout sexual maturation. The ovary continuously accumulated fatty acid content, but showed a significant decrease in MUFA content and PUFA content during the spawning period, possibly suggesting that these fatty acids were increasingly allocated to oogenesis. These evidence indicated that the fatty acid profiles in Illex argentinus showed tissue-specific characteristics, while the major fatty acid profiles were consistent in mantle tissue, digestive gland and ovary. In the mantle tissue, both SFA and MUFA were possibly mobilized to fuel reproductive growth during sexual maturation. In the digestive gland, the PUFA content might be continuously transferred to mantle and ovary with maturation. Regarding fatty acids in the ovary, the content might be continuously accumulated and subsequently used to support oogenesis during the sexual maturation.
引文
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[33] BOYLE P,RODHOUSE P.Cephalopods:ecology and fisheries[M].Oxford,UK:Blackwell Science,2005:464.
[34] 陈斌,冯健,吴彬,等.饥饿对太平洋鲑(Oncorhynchus spp.)鱼体脂肪与脂肪酸的影响[J].海洋与湖沼,2012,43(6):1247-1253.CHEN B,FENG J,WU B,et al.The effects of starvation on fat and fatty acids composition in Pacific salmon (Oncorhynchus spp.)[J].Oceanologia et Limnologia Sinica,2012,43(6):1247-1253.
[35] LIN D M,CHEN X J,CHEN Y,et al.Sex-specific reproductive investment of summer spawners of Illex argentinus in the southwest Atlantic[J].Invertebrate Biology,2015,134(3):203-213.
[36] LAPTIKHOVSKY V V,NIGMATULLIN C M.Egg size,fecundity,and spawning in females of the genus Illex (Cephalopoda:Ommastrephidae)[J].ICES Journal of Marine Science,1993,50(4):393-403.
[37] LIN D M,CHEN X J,CHEN Y,et al.Ovarian development in Argentinean shortfin squid Illex argentinus:group-synchrony for corroboration of intermittent spawning strategy[J].Hydrobiologia,2017,795(1):327-339.
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[39] REIS D B,GARCíA-HERRERO I,RIERA R,et al.An insight on Octopus vulgaris paralarvae lipid requirements under rearing conditions[J].Aquaculture Nutrition,2015,21(6):797-806.
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[2] KJ?RSVIK E,MANGOR-JENSEN A,HOLMEFJORD I.Egg quality in fishes[J].Advances in Marine Biology,1990,26:71-113.
[3] SARGENT J R,BELL J G,BELL M V,et al.Requirement criteria for essential fatty acids[J].Journal of Applied Ichthyology,1995,11(3/4):183-198.
[4] 高淳仁,雷霁霖.不同脂肪源对真鲷幼鱼生长、存活及体内脂肪酸组成的影响[J].中国水产科学,1999,6(3):55-60.GAO C R,LEI J L.Effects of different lipids on growth,survival and tissue fatty acid composition of juvenile Pagrosomus major[J].Journal of Fishery Sciences of China,1999,6(3):55-60.
[5] IVERSON S J.Tracing aquatic food webs using fatty acids:from qualitative indicators to quantitative determination[M]//KAINZ M,BRETT M T,ARTS M T.Lipids in Aquatic Ecosystems.New York:Springer,2009:281-308.
[6] PHILLIPS K L,JACKSON G D,NICHOLS P D.Predation on myctophids by the squid Moroteuthis ingens around Macquarie and Heard Islands:stomach contents and fatty acid analyses[J].Marine Ecology Progress Series,2001,215:179-189.
[7] PHILLIPS K L,NICHOLS P D,JACKSON G D.Lipid and fatty acid composition of the mantle and digestive gland of four Southern Ocean squid species:implications for food-web studies[J].Antarctic Science,2002,14(3):212-220.
[8] 黄亮,吴莹,张经.脂肪酸标志水生生态系统营养关系的研究[J].海洋科学,2009,33(3):93-96.HUANG L,WU Y,ZHANG J.Studies on nutrient relation of aquatic ecosystem using fatty acids[J].Marine Sciences,2009,33(3):93-96.
[9] RODHOUSE P G K,ARKHIPKIN A I,LAPTIKHOVSKY V,et al.Illex argentinus,argentine shortfin squid[J].Advances in Squid Biology,Ecology and Fisheries.Part II,2013,23(4):109-148.
[10] ARKHIPKIN A I.Squid as nutrient vectors linking Southwest Atlantic marine ecosystems[J].Deep Sea Research Part II:Topical Studies in Oceanography,2013,95:7-20.
[11] FAO.Review of the state of world marine fishery resources[M].Rome:Food and Agriculture Organization of the United Nations,2011.
[12] 陈新军,陆化杰,刘必林,等.大洋性柔鱼类资源开发现状及可持续利用的科学问题[J].上海海洋大学学报,2012,21(5):831-840.CHEN X J,LU H J,LIU B L,et al.Current exploitation and some scientific issues in the sustainable utilization of Ommastrephidae[J].Journal of Shanghai Ocean University,2012,21(5):831-840.
[13] ARKHIPKIN A.Age,growth,stock structure and migratory rate of pre-spawning short-finned squid Illex argentinus based on statolith ageing investigations[J].Fisheries Research,1993,16(4):313-338.
[14] CHIU T Y,CHIU T S,CHEN C S.Movement patterns determine the availability of Argentine shortfin squid Illex argentinus to fisheries[J].Fisheries Research,2017,193:71-80.
[15] 林东明,陈新军,魏嫣然,等.阿根廷滑柔鱼雌性个体肌肉和性腺组织能量积累及其生殖投入[J].水产学报,2017,41(1):70-80.LIN D M,CHEN X J,WEI Y R,et al.Energy accumulation of both somatic and reproductive tissues and its allocation to reproduction in Argentinean short-fin squid (Illex argentinus)[J].Journal of Fisheries of China,2017,41(1):70-80.
[16] LIN D M,CHEN X J,WEI Y R,et al.The energy accumulation of somatic tissue and reproductive organs in post-recruit female Illex argentinus and the relationship with sea surface oceanography[J].Fisheries Research,2017,185:102-114.
[17] GARCíA-GARRIDO S,HACHERO-CRUZADO I,GARRIDO D,et al.Lipid composition of the mantle and digestive gland of Octopus vulgaris juveniles (Cuvier,1797) exposed to prolonged starvation[J].Aquaculture International,2010,18(6):1223-1241.
[18] GARCíA S,DOMINGUES P,NAVARRO J C,et al.Growth,partial energy balance,mantle and digestive gland lipid composition of Octopus vulgaris (Cuvier,1797) fed with two artificial diets[J].Aquaculture Nutrition,2011,17(2):e174-e187.
[19] MONROIG ó,GUINOT D,HONTORIA F,et al.Biosynthesis of essential fatty acids in Octopus vulgaris (Cuvier,1797):Molecular cloning,functional characterisation and tissue distribution of a fatty acyl elongase[J].Aquaculture,2012,360-361:45-53.
[20] REIS D B,ACOSTA N G,ALMANSA E,et al.In vivo metabolism of unsaturated fatty acids in Octopus vulgaris hatchlings determined by incubation with 14C-labelled fatty acids added directly to seawater as protein complexes[J].Aquaculture,2014,431:28-33.
[21] WAKO Y,ISHIKAWA S,NAKAYA H.Comparison of fatty acids in liver lipids from various sizes of squid (Illex argentinus)[J].Bioscience,Biotechnology,and Biochemistry,1993,57(12):2181-2183.
[22] ARKHIPKIN A I.Reproductive system structure,development and function in cephalopods with a new general scale for maturity stages[J].Journal of northwest Atlantic Fishery Science,1992,12:63-74.
[23] ICES.Report of the workshop on sexual maturity staging of cephalopods (WKMSCEPH)[R].Livorno,Italy:ICES,2010:14-17.
[24] 林东明,陈新军,方舟.西南大西洋阿根廷滑柔鱼夏季产卵种群繁殖生物学的初步研究[J].水产学报,2014,38(6):843-852.LIN D M,CHEN X J,FANG Z.Preliminary study on reproductive biology of summer spawning stock of Illex argentinus in the southwestern Atlantic Ocean[J].Journal of Fisheries of China,2014,38(6):843-852.
[25] 中华人民共和国国家质量监督检验检疫总局,中国国家标准化管理委员会.GB/T 22223—2008 食品中总脂肪、饱和脂肪(酸)、不饱和脂肪(酸)的测定水解提取-气相色谱法[S].北京:中国标准出版社,2008:1-16.General Administration of Quality Supervision,Inspection and Quarantine of the People's Republic of China,Standardization Administration of China.GB/T 22223-2008 Determination of total fat saturated fat and unsaturated fat in foods-Hydrolytic extraction-Gas chromatography[S].Beijing:Standards Press of China,2008:1-16.
[26] ZAR J H.Biostatistical analysis[M].4th ed.Upper Saddle River,New Jersey:Prentice Hall,1999,
[27] NAVARRO J C,VILLANUEVA R.Lipid and fatty acid composition of early stages of cephalopods:an approach to their lipid requirements[J].Aquaculture,2000,183(1/2):161-177.
[28] NAVARRO J C,VILLANUEVA R.The fatty acid composition of Octopus vulgaris paralarvae reared with live and inert food:deviation from their natural fatty acid profile[J].Aquaculture,2003,219(1/4):613-631.
[29] ROSA R,COSTA P R,NUNES M L.Effect of sexual maturation on the tissue biochemical composition of Octopus vulgaris and O.defilippi (Mollusca:Cephalopoda)[J].Marine Biology,2004,145(3):563-574.
[30] FLUCKIGER M,JACKSON G D,NICHOLS P,et al.An experimental study of the effect of diet on the fatty acid profiles of the European Cuttlefish (Sepia officinalis)[J].Marine Biology,2008,154(2):363-372.
[31] SARGENT J,BELL G,MCEVOY L,et al.Recent developments in the essential fatty acid nutrition of fish[J].Aquaculture,1999,177(1/4):191-199.
[32] JAYA-RAM A,KUAH M K,LIM P S,et al.Influence of dietary HUFA levels on reproductive performance,tissue fatty acid profile and desaturase and elongase mRNAs expression in female zebrafish Danio rerio[J].Aquaculture,2008,277(3/4):275-281.
[33] BOYLE P,RODHOUSE P.Cephalopods:ecology and fisheries[M].Oxford,UK:Blackwell Science,2005:464.
[34] 陈斌,冯健,吴彬,等.饥饿对太平洋鲑(Oncorhynchus spp.)鱼体脂肪与脂肪酸的影响[J].海洋与湖沼,2012,43(6):1247-1253.CHEN B,FENG J,WU B,et al.The effects of starvation on fat and fatty acids composition in Pacific salmon (Oncorhynchus spp.)[J].Oceanologia et Limnologia Sinica,2012,43(6):1247-1253.
[35] LIN D M,CHEN X J,CHEN Y,et al.Sex-specific reproductive investment of summer spawners of Illex argentinus in the southwest Atlantic[J].Invertebrate Biology,2015,134(3):203-213.
[36] LAPTIKHOVSKY V V,NIGMATULLIN C M.Egg size,fecundity,and spawning in females of the genus Illex (Cephalopoda:Ommastrephidae)[J].ICES Journal of Marine Science,1993,50(4):393-403.
[37] LIN D M,CHEN X J,CHEN Y,et al.Ovarian development in Argentinean shortfin squid Illex argentinus:group-synchrony for corroboration of intermittent spawning strategy[J].Hydrobiologia,2017,795(1):327-339.
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