金乌贼微卫星标记开发及两个野生群体遗传多样性比较
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摘要
为进一步了解金乌贼Sepia esculenta野生群体遗传多样性水平,采用磁珠富集法从金乌贼基因组中开发了微卫星标记,并应用青岛(QD)金乌贼野生群体对其多态性进行评价,进而比较了青岛和长江口(CJK)两个野生群体的遗传差异。结果表明:开发标记的种群遗传学评价显示,在269个克隆子中,有192个含有微卫星序列(71.38%),基于该序列设计85对引物,其中20对通过筛选,完美型占60%,非完美型占10%,复合型占30%;经群体验证,表观等位基因数(A)为2～17不等,平均为8.15,观测杂合度(H_O)分布范围为0.146～0.936,平均为0.630,期望杂合度(H_E)分布范围为0.172～0.930,平均为0.702;其中5个位点不符合哈迪—温伯格(Hardy-Weinberg)平衡预期,存在零等位基因可能是其偏离平衡的原因;应用本研究开发的引物对2个近源种扩增,开发的20个标记位点中有6个在针乌贼Sepia andreana中表现为多态,4个位点在曼氏无针乌贼Sepiella maindroni中表现为多态;应用开发的11个位点对青岛和长江口两个野生群体进行遗传差异分析显示,所有位点中,青岛群体共检测到163个等位基因,长江口群体共检测到152个等位基因,每个位点的等位基因数为5～28和6～26不等,平均等位基因数分别为14.818和13.818;青岛群体独有等位基因19个,观测杂合度分布范围为0.250～0.936,平均为0.731,期望杂合度分布范围为0.265～0.930,平均为0.771,多态信息含量为0.223～0.946,平均为0.746;长江口群体独有等位基因8个,观测杂合度分布范围为0.500～0.896,平均为0.731,期望杂合度分布范围为0.623～0.960,平均为0.857,多态信息含量为0.614～0.948,平均为0.845;群体间遗传分化较弱(F_(st)值为0.032 5),群体分配分析结果表明,两个种群中所有个体正确分配到各自种群的概率分别为86.4%和84.0%。研究表明,本试验中开发的微卫星标记位点多样性水平略低于前人的研究,但有一定的跨种扩增通用性,长江口群体多样性水平略高于青岛群体,尽管两个群体间遗传分化程度不高,但存在明显差异。
Microsatellite loci were isolated and characterized through the magnetic beads enrichment method, and polymorphism evaluation and genetic differences were investigated in two wild populations(Qingdao, QD and Yangtze River Delta, CJK) of golden cuttlefish Sepia esculenta along coastal waters in China to explore the genetic diversity of the species. The population genetic evaluation of microsatellite markers indicated that in 269 clones 192 clones contained microsatellite sequences(71.38%), and 85 pairs of primers were designed based on these sequences, among which 20 pairs were clearly amplified and shown to be polymorphic, perfect SSR(accounting for 60%), imperfect SSR(10%), and compound SSR(30%). The numbers of alleles per locus were ranged from 2 to 17(average 8.15), the observed from 0.146 to 0.936(average 0.630)and expected heterozygosities from 0.172 to 0.930(mean 0.702). Fifteen of twenty loci was conformed to Hardy-Weinberg equilibrium and no significant linkage disequilibrium was found between any pair of loci after Bonferroni correction. Six and four loci displayed successful cross-species amplification in two other closely related species, Sepia andreana and Sepiella maindroni. A total of 163 alleles were detected in QD population, and 152 alleles in CJK population in 11 polymorphic loci analysis of population genetic diversity. The number of alleles was ranged from 5 to 28, with average allele number of 14.818, and 6 to 26, with average allele number of 13.818. There were 19 unique alleles in QD population,with observed heterozygosity from 0.250 to 0.936, with an average of 0.731, expected heterozygosity from 0.265 to 0.930 with an average of 0.771, polymorphic information content(PIC) from 0.223 to 0.946 with an average 0.746. There were 8 unique alleles in CJK population, with observation heterozygosity from 0.500 to 0.896(average 0.731), expected heterozygosity from 0.623 to 0.960(average 0.857), and PIC from 0.614 to 0.948 with an average 0.845. The genetic differentiation value(F_(st)) of 0.032 5 was observed between the populations. The population distribution analysis revealed that the probability that all individuals in the two populations were correctly assigned to their respective populations was 86.4% in QD population and 84.0% in CJK population. The findings indicated that the level of diversity of microsatellite loci developed in this study was slightly lower than that in previous studies, with a certain cross-species amplification rate. Population diversity was higher in CJK than that in QD, though low degree of genetic differentiation between the two populations, with significant differences.
Microsatellite loci were isolated and characterized through the magnetic beads enrichment method, and polymorphism evaluation and genetic differences were investigated in two wild populations(Qingdao, QD and Yangtze River Delta, CJK) of golden cuttlefish Sepia esculenta along coastal waters in China to explore the genetic diversity of the species. The population genetic evaluation of microsatellite markers indicated that in 269 clones 192 clones contained microsatellite sequences(71.38%), and 85 pairs of primers were designed based on these sequences, among which 20 pairs were clearly amplified and shown to be polymorphic, perfect SSR(accounting for 60%), imperfect SSR(10%), and compound SSR(30%). The numbers of alleles per locus were ranged from 2 to 17(average 8.15), the observed from 0.146 to 0.936(average 0.630)and expected heterozygosities from 0.172 to 0.930(mean 0.702). Fifteen of twenty loci was conformed to Hardy-Weinberg equilibrium and no significant linkage disequilibrium was found between any pair of loci after Bonferroni correction. Six and four loci displayed successful cross-species amplification in two other closely related species, Sepia andreana and Sepiella maindroni. A total of 163 alleles were detected in QD population, and 152 alleles in CJK population in 11 polymorphic loci analysis of population genetic diversity. The number of alleles was ranged from 5 to 28, with average allele number of 14.818, and 6 to 26, with average allele number of 13.818. There were 19 unique alleles in QD population,with observed heterozygosity from 0.250 to 0.936, with an average of 0.731, expected heterozygosity from 0.265 to 0.930 with an average of 0.771, polymorphic information content(PIC) from 0.223 to 0.946 with an average 0.746. There were 8 unique alleles in CJK population, with observation heterozygosity from 0.500 to 0.896(average 0.731), expected heterozygosity from 0.623 to 0.960(average 0.857), and PIC from 0.614 to 0.948 with an average 0.845. The genetic differentiation value(F_(st)) of 0.032 5 was observed between the populations. The population distribution analysis revealed that the probability that all individuals in the two populations were correctly assigned to their respective populations was 86.4% in QD population and 84.0% in CJK population. The findings indicated that the level of diversity of microsatellite loci developed in this study was slightly lower than that in previous studies, with a certain cross-species amplification rate. Population diversity was higher in CJK than that in QD, though low degree of genetic differentiation between the two populations, with significant differences.
引文
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[7] 汪金海,韩松,郑小东,等.金乌贼(Sepia esculenta)繁殖模式的分子学鉴定[J].海洋与湖沼,2017,48(1):184-189.
[8] 韦柳枝,高天翔,韩志强,等.日照近海金乌贼生物学的初步研究[J].中国海洋大学学报:自然科学版,2005,35(6):923-928.
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[10] 金洋,薛张芝,张洪超,等.金乌贼肌肉中三甲胺脱甲基酶的分离纯化及酶学性质[J].水产学报,2017,41(6):845-853.
[11] 刘长琳,赵法箴,陈四清,等.金乌贼胚胎发育阶段主要生化成分的变化[J].中国海洋大学学报:自然科学版,2016,46(11):62-72.
[12] 刘长琳,阮飞腾,秦搏,等.野生金乌贼成体肌肉的营养成分分析及评价[J].海洋科学,2016,40(8):42-48.
[13] 刘长琳,刘思玮,赵法箴,等.金乌贼(Sepia esculenta)早期发育阶段相关酶活性的变化[J].渔业科学进展,2016,37(6):105-109.
[14] 周维武.金乌贼(Sepia esculenta Hoyle)人工孵化与培育技术[J].现代渔业信息,2007,22(11):27-29.
[15] 陈四清,刘长琳,庄志猛,等.饵料、盐度对金乌贼幼体生长的影响[J].渔业现代化,2008,35(6):23-25,32.
[16] 刘长琳,庄志猛,陈四清,等.金乌贼亲体驯养与繁殖特性研究[J].渔业现代化,2009,36(2):34-42.
[17] Zheng Xiaodong,Zhao Jiaming,Xiao Shu,et al.Isozymes analysis of the golden cuttlefish Sepia esculenta (Cephalopoda:Sepiidea)[J].Journal of Ocean University of China,2004,3(1):48-52.
[18] Wei Liuzhi,Gao Tianxiang,Zhang Xiumei.Isozymes analysis of Sepia esculenta (Cephalopoda:Sepiidea)[J].Journal of Fishery Sciences of China,2005,12(5):449-555.
[19] Zheng X D,Ikeda M,Kong L F,et al.Genetic diversity and population structure of the golden cuttlefish,Sepia esculenta (Cephalopoda:Sepiidea) indicated by microsatellite DNA variations[J].Marine Ecology,2009,30,448-454.
[20] Zheng Xiaodong,Ikeda M,Barinova A,et al.Isolation and characterization of microsatellite DNA loci from the golden cuttlefish,Sepia esculenta Hoyle (Cephalopoda)[J].Molecular Ecology Notes,2007,7(1):40-42.
[21] 郑小东,许然,池田実,等.基于COI基因的西北太平洋金乌贼种群遗传学研究[J].中国海洋大学学报:自然科学版,2017,47(9):55-61.
[22] 孙孝文,张晓锋,赵莹莹,等.水产生物微卫星标记技术研究进展及其应用[J].中国水产科学,2008,15(4):689-703.
[23] 徐浩,鲁翠云,孙孝文.利用164个微卫星标记分析镜鲤家系的遗传多样性和经济性状[J].大连海洋大学学报,2013,28(3):247-253.
[24] 董颖,杨瑞,姜志强,等.微卫星标记在人工养殖小体鲟种群的数据分析方法比较和遗传多样性分析[J].大连海洋大学学报,2016,31(5):516-521.
[25] 李云霞,李娇,丁君.等.基于微卫星标记的刺参群体遗传结构分析及与经济性状的相关性研究[J].大连海洋大学学报,2013,28(5):438-444.
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