我国近海常见磷虾遗传多样性与分化的分子生态学研究
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摘要
磷虾是一类大型的海洋浮游动物，数量大，分布广，是海洋浮游动物的重要类群之一，在海洋生态系统中具有重要地位，尤其在南大洋，是维系海洋初级生产者与更高营养阶的主要环节之一；同时，磷虾又是许多经济鱼类的主要饵料，近年来，磷虾作为新兴的渔业资源正引起人们的高度重视。全世界海洋中磷虾类近90种，中国近海磷虾类迄今已记录2科7属47种，其中以太平洋磷虾、长额磷虾、宽额假磷虾和中华假磷虾为主要优势种。它们的生态特性和分布中心各有不同，可作为研究海流和水团的指示生物。
本文采用聚丙烯酰胺凝胶电泳，通过等位酶技术分析了我国近海太平洋磷虾、宽额假磷虾和中华假磷虾种群的遗传多样性与遗传分化情况，为进一步了解磷虾的起源和系统进化提供实验证据，也为海洋浮游生物多样性保护提供基础资料。
(一)太平洋磷虾遗传多样性与分化研究：
1．本文分析了黄、东海2个航次，8个站位的太平洋磷虾群体，在7个酶系统中，共检测到9个酶位点，即：α淀粉酶(1个位点，3个等位基因)，R淀粉酶(1个位点，2个等位基因)，酯酶(3个位点，Est5、Est6和Est7，除Est7为单态外，Est5和Est6各有2个等位基因)，苹果酸脱氢酶(1个位点，2个等位基因)，苹果酸酶(1个位点，2个等位基因)，乳酸脱氢酶(1个位点，4个等位基因)，磷酸葡萄糖转氨酶(1个位点，5个等位基因)。
2．除E1站位因样品数太少未进行遗传分析外，其余7个站位太平洋磷虾群体具有较高的遗传多样性：各站位平均每个位点等位基因有效数A_e为1.2845～1.4165，预期杂合度H_e为0.1457～0.2271，多态位点百分数P为42.86～87.50％；所有站位太平洋磷虾总的多态位点百分数P=88.89％，预期杂合度H_e=0.3140±0.2552，A_e=1.6628±0.6585。
3．7个站位太平洋磷虾群体间遗传分化程度较高，G_(st)=0.4837，说明群体中总的遗传多样性48.37％来自各站位间的遗传变异，51.63％属于站位内的遗传变异，与F统计量F_(st)=0.5123基本一致。站位间的基因流N_m=0.2380，表明各站位间基因交流有限，反映出各站位间存在着隔离。
我国近海常见磷虾遗传多样性与分化的分子生态学研究
各站位间遗传距离为D二0.3383(0.0032一0.7641)，遗传一致度I=
0.7482(0.4658一0.9968)，聚类分析表明:黄、东海太平洋磷虾可分为
2个种群，一个由处于黄海冷水团中央的群体构成，另一个由黄海冷水
团边缘和东海群体组成。海流、水团以及生境选择差异可能是导致黄、
东海太平洋磷虾群体间生化遗传差异的主要原因。
(二)宽额假磷虾遗传多样性和遗传分化研究
1.本文对东海外海和南海2个站位宽额假磷虾群体进行了分析，在检测
的9个酶系统中，共检测到11个酶位点:天冬氨酸转氨酶(l个位
点，2个等位基因)，碱性磷酸酶(2个位点，A加了和A加2各有2个等
位基因)，R淀粉酶(l个位点，2个等位基因)，醋酶(2个位点，
Es巧和Est7各有2个等位基因)，苹果酸脱氢酶(l个位点，3个等
位基因)，苹果酸酶(l个位点，2个等位基因)，乳酸脱氢酶(l个
位点，4个等位基因)，磷酸葡萄糖转氨酶(l个位点，3个等位基因);
a淀粉酶为单态。其中东海E7站位未对ALP和AAT进行分析，Est7
仅在E7站位有表达。
2.2个站位宽额假磷虾群体遗传多样性水平最高:各站位平均每个位点
等位基因有效数A。为1 .3791一1 .6534，预期杂合度He为0.224)
0.3269，多态位点百分数P为70.00一75.00%;所有站位宽额假磷虾
总的多态位点百分数P二90.91%，预期杂合度He=0.3336士0.1 961，
A。=1 .6310士0.1509。
3.2个站位宽额假磷虾群体间遗传分化程度较高，Gs，=0.2804，说明群
体中总的遗传多样性28.04%来自各站位间的遗传变异，71.%%属于
站位内的遗传变异，接近F统计量Fs，二0.3引O。站位间的基因流呱
二O，4832，表明站位间存在着隔离、基因交流有限。站位间遗传距离
为D二0.2749，遗传一致度I=0.75%，结果表明:E7站位和06A站
位的宽额假磷虾分别代表着不同的地理种群，空间隔离是东、南海宽
额假磷虾群体间生化遗传差异的主要原因。
(三)中华假磷虾遗传多样性
1.厦门港中华假磷虾在检测的9个酶系统中，共检测到10个酶位点:
我国近海常见磷虾遗传多样性与分化的分子生态学研究
天冬氨酸转氨酶、a淀粉酶、酣酶和苹果酸酶均为l个位点，具2个
等位基因;乳酸脱氢酶和苹果酸脱氢酶(l个位点，3个等位基因)，
磷酸葡萄糖转氨酶(l个位点，4个等位基因);碱性磷酸酶(2个位点，
月加了有2个等位基因);AIPZ和R淀粉酶为单态。
中华假磷虾遗传多样性最小，平均每个位点等位基因有效数A。二
1.2624士0.3363，预期杂合度He为0.1“1士0.1766，多态位点百分数
P为80.00%
中华假磷虾和宽额假磷虾的遗传距离为0.7870，遗传一致度了二
0.4696，种f司界限明显。
The Order Euphausiacea consists of 86 or 87 species distributed over the ocean of the world, plays an important role in marine ecosystem, particularly in southern ocean, and is the key link of marine primarily productivity and higher trophic levels. Krill fisheries are relatively recent developments.
47 species belonging to seven genera of two families in krill were recorded in China seas. Among them, Euphausia pacifica, Pseudeuphausia latifrons and P. sinica are dominant species that play a key role in the China sea. By means of the vertical slab polyacrylamide gel electrophoresis, genetic heterogeneity of three krill was investigated, in order to elucidate the genetic diversity and differentiation among them. 1. Genetic diversity and differentiation of E. pacifica
E. pacifica were collected at eight stations in the Yellow Sea and East China Sea between 2000-2001 as part of 973 Chinese Globec. Seven enzyme systems were assayed as follow, a-l,4-glucan-4-glucaflhydrolase (E.C. 3.2.1.1, a-AMY), a-l,6-glucano-hydrolase (E.C. 3.2.1.9, R-AMY), esterase (E.C. 3.1.1.1, EST), malate dehydrogenase (E.C. 1.1.1.37, MDH), malic enzyme (E.C. 1.1.1.40, ME), lactate dehydrogenase (E.C. 1.1.1.27, LDH), and glucose phosphate isomerase (E.C. 5.3.1.9, PGI).
The proportion of polymorphic loci per station ranged from 42.86 to 87.50%, the mean value over all stations was 88.89%. Ivfean expected heterozygosity per locus (He) calculated over the 9 loci analyzed ranged from 0.1457 to 0.2271, the average over all stations was 0.3140+0.2552. The effective number of alleles per locus varied between 1.2845 and 1.4165, the mean value over all stations was 1.6628?.6585.
Genetic differentiation was high among the stations, the mean FST for all loci was 0.5123. The gene flow Nm was 0.2380<1, indicated that gene drift would promote the
genetic differentiation. The Nei's genetic distance(D) between the stations was 0.3383 (0.0032-0.7641), and the genetic identity was 0.7482 (0.4658-0.9968). The arrangement of the various stations on an UPGMA dendrogram on the basis of D produced two main clusters, one formed by the stations E2 and 1 -4, and the other by the remaining stations. The genetic subdivision was due to the inche variation and circulation vicariance. The stations E2 and 1-4 were located at the cold water mass area of the central Yellow Sea, which characterized by low temperature, high salinity and stable theromocline would generate a retention mechanism that promoted the formation of separate, self-supporting stocks of krill.
2 Genetic diversity and differentiation of P. latifrons
Specimens of P. latifrons were collected from the East China Sea and the South China Sea. The zymogram phenotypes of aspartate aminotransferase (E. C. 2.6.1.1, AAT), alkaline phosphatase (E. C. 3.1.3.1, ALP), a-amylase (a-AMY), R-amylase (R-AMY), esterase (EST), lactate dehydrogenase (LDH), raalate dehydrogenase (MDH), malic enzyme (ME), and phosphoglweoisomerase (PGI) were scored.
The proportion of polymorphic loci per station ranged from 70.00 to 75.00%, the mean value over two stations was 90.91%. Mean expected heterozygosity per locus (He) calculated over the 11 loci analyzed ranged from 0.2240 to 0.3269, the average over two stations was 0.3336?.1961. The effective number of alleles per locus varied between 1.3791 and 1.6534, the mean value over two stations was 1.6310+0.1509.
The standardized variation in allele frequencies of P. latifrons populations between two stations was 0.3410, and the gene flow was 0.4832, suggesting that the magnitude of genetic differentiation between two populations was relatively high. The Nei's genetic distance between two stations was 0.2749, and the genetic identity was 0.7596. The genetic subdivision was due to the geographic barrier.
3 Genetic diversity of P. sinica
The genetic structure of P. sinica in Xiamen harbour was examined by analyzing the zymogram phenotypes of aspartate aminotransferase (AAT), alkaline phosphatase
(ALP), a-amylase (o-AMY), R-amylase (R-AMY), esterase (EST), lactate deh
本文采用聚丙烯酰胺凝胶电泳，通过等位酶技术分析了我国近海太平洋磷虾、宽额假磷虾和中华假磷虾种群的遗传多样性与遗传分化情况，为进一步了解磷虾的起源和系统进化提供实验证据，也为海洋浮游生物多样性保护提供基础资料。
(一)太平洋磷虾遗传多样性与分化研究：
1．本文分析了黄、东海2个航次，8个站位的太平洋磷虾群体，在7个酶系统中，共检测到9个酶位点，即：α淀粉酶(1个位点，3个等位基因)，R淀粉酶(1个位点，2个等位基因)，酯酶(3个位点，Est5、Est6和Est7，除Est7为单态外，Est5和Est6各有2个等位基因)，苹果酸脱氢酶(1个位点，2个等位基因)，苹果酸酶(1个位点，2个等位基因)，乳酸脱氢酶(1个位点，4个等位基因)，磷酸葡萄糖转氨酶(1个位点，5个等位基因)。
2．除E1站位因样品数太少未进行遗传分析外，其余7个站位太平洋磷虾群体具有较高的遗传多样性：各站位平均每个位点等位基因有效数A_e为1.2845～1.4165，预期杂合度H_e为0.1457～0.2271，多态位点百分数P为42.86～87.50％；所有站位太平洋磷虾总的多态位点百分数P=88.89％，预期杂合度H_e=0.3140±0.2552，A_e=1.6628±0.6585。
3．7个站位太平洋磷虾群体间遗传分化程度较高，G_(st)=0.4837，说明群体中总的遗传多样性48.37％来自各站位间的遗传变异，51.63％属于站位内的遗传变异，与F统计量F_(st)=0.5123基本一致。站位间的基因流N_m=0.2380，表明各站位间基因交流有限，反映出各站位间存在着隔离。
我国近海常见磷虾遗传多样性与分化的分子生态学研究
各站位间遗传距离为D二0.3383(0.0032一0.7641)，遗传一致度I=
0.7482(0.4658一0.9968)，聚类分析表明:黄、东海太平洋磷虾可分为
2个种群，一个由处于黄海冷水团中央的群体构成，另一个由黄海冷水
团边缘和东海群体组成。海流、水团以及生境选择差异可能是导致黄、
东海太平洋磷虾群体间生化遗传差异的主要原因。
(二)宽额假磷虾遗传多样性和遗传分化研究
1.本文对东海外海和南海2个站位宽额假磷虾群体进行了分析，在检测
的9个酶系统中，共检测到11个酶位点:天冬氨酸转氨酶(l个位
点，2个等位基因)，碱性磷酸酶(2个位点，A加了和A加2各有2个等
位基因)，R淀粉酶(l个位点，2个等位基因)，醋酶(2个位点，
Es巧和Est7各有2个等位基因)，苹果酸脱氢酶(l个位点，3个等
位基因)，苹果酸酶(l个位点，2个等位基因)，乳酸脱氢酶(l个
位点，4个等位基因)，磷酸葡萄糖转氨酶(l个位点，3个等位基因);
a淀粉酶为单态。其中东海E7站位未对ALP和AAT进行分析，Est7
仅在E7站位有表达。
2.2个站位宽额假磷虾群体遗传多样性水平最高:各站位平均每个位点
等位基因有效数A。为1 .3791一1 .6534，预期杂合度He为0.224)
0.3269，多态位点百分数P为70.00一75.00%;所有站位宽额假磷虾
总的多态位点百分数P二90.91%，预期杂合度He=0.3336士0.1 961，
A。=1 .6310士0.1509。
3.2个站位宽额假磷虾群体间遗传分化程度较高，Gs，=0.2804，说明群
体中总的遗传多样性28.04%来自各站位间的遗传变异，71.%%属于
站位内的遗传变异，接近F统计量Fs，二0.3引O。站位间的基因流呱
二O，4832，表明站位间存在着隔离、基因交流有限。站位间遗传距离
为D二0.2749，遗传一致度I=0.75%，结果表明:E7站位和06A站
位的宽额假磷虾分别代表着不同的地理种群，空间隔离是东、南海宽
额假磷虾群体间生化遗传差异的主要原因。
(三)中华假磷虾遗传多样性
1.厦门港中华假磷虾在检测的9个酶系统中，共检测到10个酶位点:
我国近海常见磷虾遗传多样性与分化的分子生态学研究
天冬氨酸转氨酶、a淀粉酶、酣酶和苹果酸酶均为l个位点，具2个
等位基因;乳酸脱氢酶和苹果酸脱氢酶(l个位点，3个等位基因)，
磷酸葡萄糖转氨酶(l个位点，4个等位基因);碱性磷酸酶(2个位点，
月加了有2个等位基因);AIPZ和R淀粉酶为单态。
中华假磷虾遗传多样性最小，平均每个位点等位基因有效数A。二
1.2624士0.3363，预期杂合度He为0.1“1士0.1766，多态位点百分数
P为80.00%
中华假磷虾和宽额假磷虾的遗传距离为0.7870，遗传一致度了二
0.4696，种f司界限明显。
The Order Euphausiacea consists of 86 or 87 species distributed over the ocean of the world, plays an important role in marine ecosystem, particularly in southern ocean, and is the key link of marine primarily productivity and higher trophic levels. Krill fisheries are relatively recent developments.
47 species belonging to seven genera of two families in krill were recorded in China seas. Among them, Euphausia pacifica, Pseudeuphausia latifrons and P. sinica are dominant species that play a key role in the China sea. By means of the vertical slab polyacrylamide gel electrophoresis, genetic heterogeneity of three krill was investigated, in order to elucidate the genetic diversity and differentiation among them. 1. Genetic diversity and differentiation of E. pacifica
E. pacifica were collected at eight stations in the Yellow Sea and East China Sea between 2000-2001 as part of 973 Chinese Globec. Seven enzyme systems were assayed as follow, a-l,4-glucan-4-glucaflhydrolase (E.C. 3.2.1.1, a-AMY), a-l,6-glucano-hydrolase (E.C. 3.2.1.9, R-AMY), esterase (E.C. 3.1.1.1, EST), malate dehydrogenase (E.C. 1.1.1.37, MDH), malic enzyme (E.C. 1.1.1.40, ME), lactate dehydrogenase (E.C. 1.1.1.27, LDH), and glucose phosphate isomerase (E.C. 5.3.1.9, PGI).
The proportion of polymorphic loci per station ranged from 42.86 to 87.50%, the mean value over all stations was 88.89%. Ivfean expected heterozygosity per locus (He) calculated over the 9 loci analyzed ranged from 0.1457 to 0.2271, the average over all stations was 0.3140+0.2552. The effective number of alleles per locus varied between 1.2845 and 1.4165, the mean value over all stations was 1.6628?.6585.
Genetic differentiation was high among the stations, the mean FST for all loci was 0.5123. The gene flow Nm was 0.2380<1, indicated that gene drift would promote the
genetic differentiation. The Nei's genetic distance(D) between the stations was 0.3383 (0.0032-0.7641), and the genetic identity was 0.7482 (0.4658-0.9968). The arrangement of the various stations on an UPGMA dendrogram on the basis of D produced two main clusters, one formed by the stations E2 and 1 -4, and the other by the remaining stations. The genetic subdivision was due to the inche variation and circulation vicariance. The stations E2 and 1-4 were located at the cold water mass area of the central Yellow Sea, which characterized by low temperature, high salinity and stable theromocline would generate a retention mechanism that promoted the formation of separate, self-supporting stocks of krill.
2 Genetic diversity and differentiation of P. latifrons
Specimens of P. latifrons were collected from the East China Sea and the South China Sea. The zymogram phenotypes of aspartate aminotransferase (E. C. 2.6.1.1, AAT), alkaline phosphatase (E. C. 3.1.3.1, ALP), a-amylase (a-AMY), R-amylase (R-AMY), esterase (EST), lactate dehydrogenase (LDH), raalate dehydrogenase (MDH), malic enzyme (ME), and phosphoglweoisomerase (PGI) were scored.
The proportion of polymorphic loci per station ranged from 70.00 to 75.00%, the mean value over two stations was 90.91%. Mean expected heterozygosity per locus (He) calculated over the 11 loci analyzed ranged from 0.2240 to 0.3269, the average over two stations was 0.3336?.1961. The effective number of alleles per locus varied between 1.3791 and 1.6534, the mean value over two stations was 1.6310+0.1509.
The standardized variation in allele frequencies of P. latifrons populations between two stations was 0.3410, and the gene flow was 0.4832, suggesting that the magnitude of genetic differentiation between two populations was relatively high. The Nei's genetic distance between two stations was 0.2749, and the genetic identity was 0.7596. The genetic subdivision was due to the geographic barrier.
3 Genetic diversity of P. sinica
The genetic structure of P. sinica in Xiamen harbour was examined by analyzing the zymogram phenotypes of aspartate aminotransferase (AAT), alkaline phosphatase
(ALP), a-amylase (o-AMY), R-amylase (R-AMY), esterase (EST), lactate deh
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