青岛近岸金乌贼繁殖群体形态特征及遗传分化
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摘要
为探究青岛近岸金乌贼(Sepia esculenta)繁殖群体结构及其结群历时长、产卵期内亲体规格递减的结群现象,运用形态学度量方法结合微卫星分子标记技术,对青岛薛家岛近岸前、中、后3个不同洄游时期金乌贼繁殖群体的形态特征和遗传分化水平进行统计分析,探究其亲缘关系并量化差异水平。主要结果如下:形态学主成分分析(PCA)结果显示,前3个主成分的累积贡献率为60.067%,低于85%;判别分析(D_A)结果显示, 3个时期繁殖群体的判别准确率介于66.7%～82.1%,两者的散点图结果一致,均显示3个群体未明显占据不相重叠的区域。微卫星标记结果表明, 3个时期繁殖群体的遗传距离D_A介于0.12～0.16,群体遗传分化指数为0.0014～0.0064,利用Structure判断最佳理论组群为1,表明其亲缘关系较近。综合分析认为,虽青岛近海不同洄游时期金乌贼繁殖群体的规格差异较大,但形态学参数差异较小,遗传距离较近且遗传分化水平较低(F_(st)<0.05),未呈现明显的群体分化。
Population differentiation and structure are important topics of evolutionary biology, and morphology and genetics are the main methods of researching these topics. Sepia esculenta migrates annually from the overwintering field to the spawning ground, and for a long time, it has been thought that the breeding period of its inshore populations was as long as two months. In addition, during the grouping period, the specification of S. esculenta arriving in different stages showed a decreasing trend with arrival time. Thus, the objectives of this study were to characterize the population structure and clustering phenomenon in S. esculenta over time, and discover why the parental specification diminished during spawning of the mature animals in Qingdao. The morphological characteristics and genetic differentiation during three different migratory periods(early, middle, and late) of S.esculenta inhabiting the coastal waters of Xuejia Island were analyzed using morphology and DNA markers. The results showed that the cumulative contribution rate of the first three principal components was 60.067% in the principal component analysis(PCA), which was lower than 85%. The discrimination accuracy ranged from 66.7%to 82.1% in the discriminant analysis(D_A). The scatter plots of PCA and D_A showed a similar result, which is the species was not clearly dispersed among these three migratory periods. Analysis based on microsatellite markers indicated that genetic distances(D_A) ranged from 0.12 to 0.16 and the genetic differentiation index(Fst) ranged from 0.0014 to 0.0064 in S. esculenta from the three different migratory periods. The optimal number of theoretical groups was 1 according to the genetic structural analysis. These findings suggested that there was no obvious population differentiation among early, middle, and late migratory periods of S. esculenta in Qingdao, owing to the low values of morphological differences, genetic distances, and genetic differentiation, although the population showed a large difference in mantle length and body weight in specifications. The differences in the swimming ability of small and large individuals may be the main reason behind the long period of S. esculenta's clustering and the decrease of its specification over time. It is undeniable that the differences in morphology and genetics among populations of marine cephalopods populations of fish are unpredictable yet significant. Macroscopic identification methods, such as morphological studies, are vulnerable to the influence of living environment, and at the same time, marine fish with good dispersal ability usually exhibit very low genetic differentiation within a wide geographic range. Therefore, the strong athletic ability of Sepia esculenta may also lead to a similar genetic structure of its populations.
Population differentiation and structure are important topics of evolutionary biology, and morphology and genetics are the main methods of researching these topics. Sepia esculenta migrates annually from the overwintering field to the spawning ground, and for a long time, it has been thought that the breeding period of its inshore populations was as long as two months. In addition, during the grouping period, the specification of S. esculenta arriving in different stages showed a decreasing trend with arrival time. Thus, the objectives of this study were to characterize the population structure and clustering phenomenon in S. esculenta over time, and discover why the parental specification diminished during spawning of the mature animals in Qingdao. The morphological characteristics and genetic differentiation during three different migratory periods(early, middle, and late) of S.esculenta inhabiting the coastal waters of Xuejia Island were analyzed using morphology and DNA markers. The results showed that the cumulative contribution rate of the first three principal components was 60.067% in the principal component analysis(PCA), which was lower than 85%. The discrimination accuracy ranged from 66.7%to 82.1% in the discriminant analysis(D_A). The scatter plots of PCA and D_A showed a similar result, which is the species was not clearly dispersed among these three migratory periods. Analysis based on microsatellite markers indicated that genetic distances(D_A) ranged from 0.12 to 0.16 and the genetic differentiation index(Fst) ranged from 0.0014 to 0.0064 in S. esculenta from the three different migratory periods. The optimal number of theoretical groups was 1 according to the genetic structural analysis. These findings suggested that there was no obvious population differentiation among early, middle, and late migratory periods of S. esculenta in Qingdao, owing to the low values of morphological differences, genetic distances, and genetic differentiation, although the population showed a large difference in mantle length and body weight in specifications. The differences in the swimming ability of small and large individuals may be the main reason behind the long period of S. esculenta's clustering and the decrease of its specification over time. It is undeniable that the differences in morphology and genetics among populations of marine cephalopods populations of fish are unpredictable yet significant. Macroscopic identification methods, such as morphological studies, are vulnerable to the influence of living environment, and at the same time, marine fish with good dispersal ability usually exhibit very low genetic differentiation within a wide geographic range. Therefore, the strong athletic ability of Sepia esculenta may also lead to a similar genetic structure of its populations.
引文
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[2]Li J Y.On the breeding and migration of the golden cuttlefish,Sepia esculenta Hoyle,living in Yellow Sea[J].Journal of Shandong College of Oceanology,1963(2):69-108.[李嘉泳.金乌贼Sepia esculenta Hoyle在黄渤海的结群生殖和洄游[J].山东海洋学院学报,1963(2):69-108.]
[3]Zhen Y J,Chen X Z,Cheng J H,et al.Biological Resources and Environment in the East China Sea Continental Shelf[M].Shanghai:Shanghai Scientific&Technical Publishers,2003:722-727.[郑元甲,陈雪忠,程家桦,等.东海大陆架生物资源与环境[M].上海:上海科学技术出版社,2003:722-727.]
[4]Yamada U,Tagawa M,Kishita S,et al.Fishes of the East China Sea and the Yellow Sea[M].Nagasaki:Seikai National Fisheries Research Laboratory,1986:208-461.
[5]Frédérich B,Liu S Y V,Dai C F.Morphological and genetic divergences in a coral reef damselfish,Pomacentrus coelestis[J].Evolutionary Biology,2012,39(3):359-370.
[6]Xu S Y,Zhang H,Liu B Z,et al.Morphological study of Sebastiscus marmoratus among populations in China and Japan[J].Acta Hydrobiologica Sinica,2013,37(5):960-966.[徐胜勇,张辉,柳本卓,等.中日褐菖鲉群体形态学比较研究[J].水生生物学报,2013,37(5):960-966.]
[7]Han Z,Xiao Y S,Gao T X.Comparison of morphological characteristics of 9 Larimichthys polyactis populations in China[J].South China Fisheries Science,2012,8(3):25-33.[韩真,肖永双,高天翔.中国近海9个小黄鱼群体的形态学比较研究[J].南方水产科学,2012,8(3):25-33.]
[8]Mwanja M T,Muwanika V,Nyakaana S,et al.Population morphological variation of the Nile perch(Lates niloticus,L.1758),of East African Lakes and their associated waters[J].African Journal of Environmental Science and Technology,2011,5(11):941-949.
[9]Cavalcanti M J,Monteiro L R,Lopes P R D.Landmark-based morphometric analysis in selected species of serranid fishes(Perciformes:Teleostei)[J].Zoological Studies,1999,38(3):287-294.
[10]Sun C F,Ye X,Dong J J,et al.Genetic diversity analysis of six cultured populations of Macrobrachium rosenbergii using microsatellite markers[J].South China Fisheries Science,2015,11(2):20-26.[孙成飞,叶星,董浚键,等.罗氏沼虾6个养殖群体遗传多样性的微卫星分析[J].南方水产科学,2015,11(2):20-26.]
[11]Liu L W,Chen X J,Xu Q H,et al.Isolation and genetic diversity of microsatellite DNA of Ommastrephes bartramii in the North Pcific Ocean[J].Acta Ecology Sinica,2014,34(23):6847-6854.[刘连为,陈新军,许强华,等.北太平洋柔鱼微卫星标记的筛选及遗传多样性[J].生态学报,2014,34(23):6847-6854.]
[12]Wolfus G M,Garcia D K,Alcivar-Warren A.Application of the microsatellite technique for analyzing genetic diversity in shrimp breeding programs[J].Aquaculture,1997,152(1-4):35-47.
[13]Chareontawee K,Poompuang S,Na-Nakorn U,et al.Genetic diversity of hatchery stocks of giant freshwater prawn(Macrobrachium rosenbergii)in Thailand[J].Aquaculture,2007,271(1-4):121-129.
[14]Zhang Q Q,Chen J,Jiang X Y,et al.Establishment of DNAfingerprinting and analysis on genetic structure of different Parabramis and Megalobrama populations with microsatellite[J].Journal of Fisheries of China,2014,38(1):15-22.[张倩倩,陈杰,蒋霞云,等.不同鳊鲂鱼类群体微卫星DNA指纹图谱的构建和遗传结构分析[J].水产学报,2014,38(1):15-22.]
[15]Dong Z Z.On the geographical distribution of the cephalopods in the chinese waters[J].Oceanologia et Limnologia Sinica,1978,9(1):108-118.[董正之.中国近海头足类的地理分布[J].海洋与湖沼,1978,9(1):108-118.]
[16]Natsukari Y,Tashiro M.Neritic squid resources and cuttlefish resources in Japan[J].Marine and Freshwater Behaviour and Physiology,1991,18(3):149-226.
[17]Cheng J S,Zhu J S.Study on feeding characteristics and trophic levels of main economic invertebrates in the Yellow Sea[J].Acta Oceanologia Sinica,1997,19(6):102-108.[程济生,朱金声.黄海主要经济无脊椎动物摄食特征及其营养层次的研究[J].海洋学报,1997,19(6):102-108.]
[18]Wei L Z.Biology of Sepia esculenta in the coastal waters of Rizhao[D].Qingdao:Ocean University of China,2005:2-3.[韦柳枝.山东日照近海金乌贼生物学研究[D].青岛:中国海洋大学,2005:2-3.]
[19]Yagi T.On the growth of the shell in Sepia esculenta Hoyle caught in Tokyo Bay[J].Bulletin of the Japanese Society of Scientific Fisheries,1960,26(7):646-652.
[20]Wei Z B.Preliminary observation of S.esculenta habits[J].Chinese Journal of Zoology,1964,12(3):132-134.[魏臻邦.金乌贼生活习性的初步观察[J].动物学杂志,1964,12(3):132-134.]
[21]Hao Z L.Studies on the behavior and marking technology of Sepia esculenta[D].Qingdao:Ocean University of China,2007:61-67.[郝振林.金乌贼的行为习性及标志技术的研究[D].青岛:中国海洋大学,2007:61-67.]
[22]Wang L,Zhang X M,Ding P W,et al.Reproductive behavior and mating strategy of Sepia esculenta[J].Acta Oceanologia Sinica,2017,37(6):1871-1880.[王亮,张秀梅,丁鹏伟,等.金乌贼繁殖行为与交配策略[J].生态学报,2017,37(6):1871-1880.]
[23]Zhao H J,Wei B F,Hu M,et al.Preliminary study on Sepia esculenta oosperm hatching and effects of different adhesion substrates[J].Transactions of Oceanology and Limnology,2004,64(3):64-68.[赵厚钧,魏邦福,胡明,等.金乌贼受精卵孵化及不同材料附着基附卵效果的初步研究[J].海洋湖沼通报,2004,64(3):64-68.]
[24]Fujita T,Hirayama I,Matsuoka T,et al.Spawning behavior and selection of spawning substrate by cuttlefish Sepia esculenta[J].Bulletin of the Japanese Society of Scientific Fisheries,1997,63(2):145-151.
[25]Sambrook J,Russell D W.Molecular Cloning:A Laboratory Manual,3rd edition[M].New York:Cold Spring Harbor Laboratory Press,2001:479-482.
[26]Yuan Y J,Liu S F,Bai C C,et al.Isolation and characterization of new 24 microsatellite DNA markers for golden cuttlefish(Sepia esculenta)[J].International Journal of Molecular Sciences,2012,13(1):1154-1160.
[27]Zheng X D,Ikeda M,Barinova A.Isolation and characterization of microsatellite DNA loci from the golden cuttlefish,Sepia esculenta Hoyle(Cephalopoda)[J].Molecular Ecology Notes,2007,7(1):40-42.
[28]Yasuda J.Some ecological note on the cuttlefish,Sepia esculenta Hoyle[J].Nihon-Suisan-Gakkai-Shi,1951,16(8):350-356.
[29]Mcgrath B,Jackson G.Egg production in the arrow squid Nototodarus gouldi(Cephalopoda:Ommastrephidae),fast and furious or slow and steady?[J].Marine Biology,2002,141(4):699-706.
[30]Lou A R,Sun R Y,Li Q F,et al.Basic Ecology[M].Beijing:Higher Education Press,2002:24-25.[娄安如,孙濡泳,李庆芬,等.基础生态学[M].北京:高等教育出版社,2002:24-25.]
[31]Zheng X D,Ikeda M,Kong L F,et al.Genetic diversity and population structure of the golden cuttlefish,Sepia esculenta(Cephalopoda:Sepiidae)indicated by microsatellite DNAvariations[J].Marine Ecology,2009,30(4):448-454.
[32]Moghadam H Y,Zolgharnein H,Aliabadi M A,et al.Genetic analysis of cuttlefish Sepia Pharaonis(Ehrenberg,1831)populations in persian gulf with microsatellite makers[J].Journal of Shellfish Research,2015,34(2):1-5.
[33]Cabranes C,Fernandez-Rueda P,Martinez J L.Genetic structure of Octopus vulgaris around the Iberian Peninsula and Canary Islands as indicated by microsatellite DNA variation[J].ICES Journal of Marine Science,2008,65(1):12-16.
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