微卫星标记对黑龙江流域大麻哈鱼遗传多样性的研究

采用 12个微卫星标记 ,对中国 3个大麻哈鱼洄游群体 (乌苏里江、黑龙江和绥芬河 )的遗传多样性进行了检测。计算出各个种群的基因杂合度、遗传多样性和各个座位的多态信息含量。结果表明 ,3个大麻哈鱼洄游种群的平均基因杂合度分别为 :0 .6 732、0 5 995、0 .6 917,种群遗传多样性分别为 0 .70 82、0 .6 5 11、0 .76 16。这些结果表明大麻哈鱼遗传多样性还比较丰富 ,其资源的恢复具有良好的前景 ,说明当前中国大麻哈鱼资源数量下降并非由遗传因素引起 ,主要原因可能是由于过度捕捞和水域环境污染等人为因素造成。人工增殖放流为恢复中国大麻哈鱼资源起到了重要作用 ,但目前大麻哈鱼的小种群极易产生遗传瓶颈的现状也应引起人们高度重视     

Genetic analysis of four wild chum salmon<Emphasis Type="Italic">Oncorhynchus keta</Emphasis> populations in China based on microsatellite markers

Synopsis To assess the genetic variation and population structure of wild chum salmon in China, we analyzed microsatellite loci for populations in the Amur, Wusuli, Suifen Current and the Tumen rivers. We evaluated expected heterozygosity with two estimators of genetic differentiation (F_ST and G_ST) and Nei’s standard genetic distance. The average expected heterozygosity across the 10 loci was 0.65 in the Wusuli River and the Suifen Current River, 0.64 in the Amur River and 0.66 in the Tumen River, The results of this study show that the recent declines in chum salmon have not led to low levels of genetic variability in China. The proportion of inter-population subdivision among chum salmon was between 5.7 and 6.8%. According to the estimator used, the NJ tree based on Nei’s standard genetic distance indicated that there were two different branches (the Sea of Okhotsk branch and the Sea of Japan branch), the Amur River and the Wusuli River populations were closer, while the Suifen Current River and the Tumen River clustered together. The genetic test for population bottlenecks provided no evidence for a significant genetic signature of population decline, which is consistent with the record of the four populations we have in the last few years.     

Population structure of chum salmon Oncorhynchus keta in the Russian Far East, as revealed by microsatellite markers

Chum salmon populations in the Russian Far East have a complex multi-level genetic structure. A total of 53 samples (2446 fish) were grouped into five major regional clusters: the southern Kurils, eastern Sakhalin, southwestern Sakhalin, the Amur River, and a northern cluster. The northern cluster consists of chum salmon populations from a vast geographical region, including Chukotka, Kamchatka, and the continental coast of the Sea of Okhotsk. However, the degree of its genetic differentiation is low, 1.9%. In contrast, the southern population cluster exhibits much higher variation; for example, differentiation between chum salmon groups within Sakhalin Island reaches 4.6%, and the differentiation between Iturup Island and Sakhalin Island chum salmon is 7.7%. This suggests that southern populations of Asian chum salmon have a more ancient evolutionary history than northern populations. In contrast to the available data, our study indicates a great deviation of southwestern Sakhalin populations from other Sakhalin chum salmon. The Russian Far East chum salmon are genetically diverse and show statistically significant differentiation even within small geographic localities. This can be used to assign samples of unknown origins to definite local populations.     

Genetic diversity of Asian chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> (Salmonidae,Salmoniformes) and the dynamics of gene pools in Sakhalin populations under artificial reproduction

The estimates of genetic diversity in populations of chum salmon Oncorhynchus keta from different regions of Sakhalin Island, Iturup Island, and the Anadyr’ River were obtained on the basis of analysis of allozyme variability. These estimates together with our published earlier data on chum salmon from the Amur River basin and the rivers of the northern coast of the Sea of Okhotsk demonstrate pronounced regional genetic differentiation in the Asian part of the fish range. The intraregional level of interpopulation genetic diversity was maximum on Sakhalin Island (G _ST = 6.6%) and was small on Iturup Island (G _ST = 0.9%) and the northern coast of the Sea of Okhotsk (G _ST = 0.6%). Interpopulation genetic diversity of Sakhalin chum salmon was almost commensurable to the diversity of the whole pool of studied Asian populations (G _ST = 7.6%) and would be presented more completely in baselines assigned for genetic identification of mixed stocks. It was demonstrated that the character and degree of genetic differentiation between populations of chum salmon from the main hatcheries situated in different regions of the Sakhalin oblast and connected to one another by numerous transplantations of fertilized eggs did not change significantly during an approximately 20-year period of our observation, and this fact suggests low efficiency of such transplantations.     

Genetic differentiation among chum salmon, Oncorhynchus keta (Walbaum), from Primorye and Sakhalin

Mitochondrial DNA (mtDNA) was examined in six wild populations of chum salmon ( Oncorhynchus keta Walbaum) distributed over the Primorye Region which extends approximately 1000 km along the Sea of Japan coast, and six populations from Sakhalin, using restriction enzyme analysis. By means of two restriction enzymes ( Bam HI and Eco 81I) eight mtDNA clonal lines were revealed in the 346 chum salmon studied. Mitochondrial DNA variants grouped the fish into two major clusters representing the Primorye and Sakhalin regions. Analysis of different chum salmon generations showed stability in the temporal population genetic structure in the three Primorye populations studied, but instability in the Sakhalin population of the Naiba River. We succeeded in detecting four major mtDNA clonal variants in Primorye chum salmon. The geographic distribution of clonal frequencies in Primorye populations has a clinal U-shape associated with the north-south axis of the Primorye region. On the whole, Sakhalin populations are less heterogeneous than Primorye ones. Two hatchery-seeded stocks from south west Sakhalin showed no mtDNA clonal variation.     

Microsatellite variation reveals weak genetic structure and retention of genetic variability in threatened Chinook salmon (<Emphasis Type="Italic">Oncorhynchus tshawytscha</Emphasis>) within a Snake River watershed

Pacific salmon (Oncorhynchus spp.) have been central to the development of management concepts associated with evolutionarily significant units (ESUs), yet there are still relatively few studies of genetic diversity within threatened and endangered ESUs for salmon or other species. We analyzed genetic variation at 10 microsatellite loci to evaluate spatial population structure and genetic variability in indigenous Chinook salmon (Oncorhynchus tshawytscha) across a large wilderness basin within a Snake River ESU. Despite dramatic 20th century declines in abundance, these populations retained robust levels of genetic variability. No significant genetic bottlenecks were found, although the bottleneck metric (M ratio) was significantly correlated with average population size and variability. Weak but significant genetic structure existed among tributaries despite evidence of high levels of gene flow, with the strongest genetic differentiation mirroring the physical segregation of fish from two sub-basins. Despite the more recent colonization of one sub-basin and differences between sub-basins in the natural level of fragmentation, gene diversity and genetic differentiation were similar between sub-basins. Various factors, such as the (unknown) genetic contribution of precocial males, genetic compensation, lack of hatchery influence, and high levels of current gene flow may have contributed to the persistence of genetic variability in this system in spite of historical declines. This unique study of indigenous Chinook salmon underscores the importance of maintaining natural populations in interconnected and complex habitats to minimize losses of genetic diversity within ESUs.     

Mitochondrial DNA variation in populations of the chum salmon Oncorhynchus keta (Walbaum) from rivers of the Prymorye and Sakhalin

Mitochondrial DNA (mtDNA) variation was studied using restriction fragment length polymorphism (RFLP) in chum salmon populations from three rivers in southern Primorye and one river in Sakhalin Island. Significant differences were detected between the samples from Primorye and Sakhalin Island. No differences were found between the samples from the rivers of Primorye, which can be explained by a high rate of gene flow due to transplantation of spawn from one river to another. The effect of fish breeding on the chum salmon populations correlated with the indices of haplotype and nucleotide diversity (h and pi, respectively). The lowest diversity was found in the completely artificial population from the Ryazanovka River; the highest, in natural populations from the Narva and Naiba rivers. Frequencies of haplotypes in consecutive generations were significantly different, which confirms the effects of genetic drift on the small-size chum salmon populations of Primorye.     

Self-sustaining populations, population sinks or aggregates of strays: chum (Oncorhynchus keta) and Chinook salmon (Oncorhynchus tshawytscha) in the Wood River system, Alaska

Small populations can provide insights into ecological and evolutionary aspects of species distributions over space and time. In the Wood River system in Alaska, USA, small aggregates of Chinook (Oncorhynchus tshawytscha) and chum salmon (O. keta) spawn in an area dominated by sockeye salmon (O. nerka). Our objective was to determine whether these Chinook and chum salmon are reproductively isolated, self-sustaining populations, population sinks that produce returning adults but receive immigration, or strays from other systems that do not produce returning adults. DNA samples collected from adult chum salmon from 16 streams and Chinook salmon from four streams in the Wood River system over 3 years were compared to samples from large populations in the nearby Nushagak River system, a likely source of strays. For both species, microsatellite markers indicated no significant genetic differentiation between the two systems. Simulations of microsatellite data in a large source and a smaller sink population suggested that considerable immigration would be required to counteract the diverging effects of genetic drift and produce genetic distances as small as those observed, considering the small census sizes of the two species in the Wood River system. Thus, the Wood River system likely receives substantial immigration from neighbouring watersheds, such as the Nushagak River system, which supports highly productive runs. Although no data on population productivity in the Wood River system exist, our results suggest source-sink dynamics for the two species, a finding relevant to other systems where salmonid population sizes are limited by habitat factors.     

The genetic structure of chum salmon (Oncorhynchus keta Walbaum) populations inferred from the nucleotide variation of the mitochondrial DNA cytochrome b gene

The nucleotide sequences of a fragment of the mitochondrial DNA cytochrome b gene were determined in 12 chum salmon populations from the Russian Far East. The level of genetic diversity in the chum salmon populations from the Iturup Island, northern coast of the Sea of Okhotsk, and Anadyr' River was found to be higher than in the populations from Kamchatka and Sakhalin, which may be related to the history of their origin and dispersal. The proportions of intrapopulation genetic variability (F(ct)) and interpopulation genetic variability within the groups (F(sc)) account for 90.87 and 0.9%, respectively, and the intergroup component (F(st)) comprises 8.23%. The predominance of one haplotype, B1, which is common for all populations studied, and a low share of intergroup variability suggest the beginning of colonization by the species of the given region from a common source (group of founders) and a relatively recent time of divergence of the chum salmon populations from the region examined.     

Microsatellite variability in chum salmon of Primorye

In this study, 12 samples of chum salmon from the southern and central parts of Primorye were studied with ten microsatellite loci. All studied localities of chum salmon of Primorye formed three main genetically different groups: (1) the Narva–Barabashevka–Ryazanovka cluster of southern Primorye, (2) Kievka River, and (3) Avvakumovka River. The revealed genetic heterogeneity of chum salmon showed clear population structure in accordance with the geographical location of the samples. The study suggests that, for the purposes of artificial reproduction of chum salmon, it is desirable to perform egg planting with regard to the described population structure of chum salmon of Primorye.     

Rapid expansion of an enhanced stock of chum salmon and its impacts on wild population components

A harvested stock of chum salmon homing to Kurilskiy Bay, Iturup Island, consists of two genetically distinct river populations that reproduce in two rivers that drain into the bay and are characterized by limited gene flow. One of these is small and can be regarded as wild, whereas the other is much larger and, until recently, was composed of naturally reproducing components spawning in the river??s mainstem and tributaries, with almost no hatchery reproduction during the past two decades. The only human impact on reproduction of the chum salmon stock was regulation of the escapement, with officially accepted limits to avoid ??over-escapement??. Recently the hatchery began to release a large amount of chum salmon juveniles. As confirmed by data on variation in both age composition and microsatellite DNA, first-generation hatchery-origin fish that returned from the first large releases occupied spawning grounds and presumably competed directly with, and potentially displaced wild fish. The most dramatic example is a genetically distinct beach-spawning form of chum salmon that was swamped by much more numerous hatchery-origin fish of the river-spawning form. In order to restore and support naturally reproduced population components, careful estimation of the carrying capacity of natural spawning grounds is necessary with efforts to increase escapement to these habitats. We also recommend concerted efforts to restore and conserve a unique beach-spawning population of chum salmon. We further recommend development of a marking program for direct estimation of straying and evaluation of ecological and genetic impacts of hatchery fish on neighboring wild and natural populations.     

The variability in chum salmon Oncorhynchus keta (Walbaum) mitochondrial DNA and its connection with the paleogeological events in the northwest Pacific

The results of examining mtDNA variability in populations of chum salmon Oncorhynchus keta from the rivers of the basins of the seas of Japan and Okhotsk and in the chum salmon seasonal races of the Amur River are presented. A significant level of polymorphism between the majority of the populations studied was detected. The groups of chum salmon from the seas of Japan and Okhotsk displayed the most pronounced differences. Analysis of genetic variability demonstrated that periodic paleontologic and climatic changes in the past of this region were the most probable factor that caused the divergence of these populations. The advances and retreats of glaciers and the accompanying regressions and transgressions of the ocean level caused isolation of chum salmon in the refugia belonging hypothetically to the paleo-Shuifen and paleo-Amur regions. These population groups diverged presumably 350–450 thousand years ago. Differences between the seasonal races of the Amur chum salmon are insignificant, and their emergence dates back to the period of the last Wisconsinian glaciation. Probably, the main isolation factor now is the genetically determined time of spawning.     

The variation in chum salmon Oncorhynchus keta (Walbaum) mitochondrial DNA and its connection with the paleogeographic events in the Northwest Pacific

The results of examining mtDNA variation in populations of chum salmon Oncorhynchus keta from the rivers of the basins of the seas of Japan and Okhotsk and in the chum salmon seasonal races of the Amur River are presented. A significant level of polymorphism between the majority of the populations studied was detected. The groups of chum salmon from the Japan and Okhotsk Seas displayed the most pronounced differences. Analysis of genetic variation demonstrated that periodic paleontologic and climatic changes in the past of this region were the most probable factor that caused the divergence of these populations. The advances and retreats of glaciers and the accompanying regressions and transgressions of the ocean level caused isolation of chum salmon in the refugia belonging hypothetically to the paleo-Suifun and paleo-Amur regions. These population groups diverged presumably 350-450 thousand years ago. Differences between the seasonal races of the Amur chum salmon are insignificant, and their emergence dates back to the period of the last Wisconsin glaciation. Probably, the main isolation factor now is the genetically determined time of spawning.     

Genetic variability and differentiation of populations of chum salmon Oncorhynchus keta (Walbaum) from the southern Russian Far East

Allozyme variation of populations of chum salmon Oncorhynchus keta from southern Russian Far East was examined. Of 55 loci screened, 31 were polymorphic. Within-population variation accounted for most of the allele diversity; FST averaged over loci was 0.052. Linkage disequilibrium was found in less than 5% of locus pairs in the chum population examined. Analysis of within- and among-population variance components of linkage disequilibrium using D-statistics (Ohta, 1982) showed that most genetic variation was distributed among populations.     

Patterns of genetic diversity in population complexes of pacific chum salmon <Emphasis Type="Italic">Oncorhynchus keta</Emphasis> Walbaum,from Asia and America,inferred from allozyme polymorphism data

Based on the data of Russian and foreign researchers, a database, consisting of 100 allozyme-coding loci examined in 288 chum salmon populations from Asia and Northern America, was constructed. Using G-test, genetic heterogeneity of Asian population samples of chum salmon was evaluated. Correlations between the frequencies of major alleles and geographic latitude of the mouths of native rivers were estimated. Using the methods of Nei and Cavalli-Sforza and Edwards, for different local chum salmon stock groups the genetic distances at the number of polymorphic enzyme loci were determined. Analysis of these distances made it possible to evaluate the patterns of genetic diversity in regional population groups from the Russian Far East, Japan, and North America. The proportions of genetic variation at each hierarchical level, identified in accordance with the geographical positions of the populations, were estimated through partitioning of variation in Asian populations into within and between-population components. It was demonstrated that intraspecific genetic structure of chum salmon corresponded geographic subdivision into regional population groups.     

Interregional differentiation of chum salmon from Sakhalin and South Kurils infered from microsatellite markers

Variability at ten microsatellite loci was examined in wild and hatchery populations of chum salmon from the Sakhalin Island and Southern Kuril Islands, Iturup and Kunashir. Substantial genetic differences between Sakhalin and South Kurils chum salmon (the differentiation theta reached 6.0%) were revealed. Statistically significant differences between chum salmon from Iturup and that from Kunashir were demonstrated, as well as between the chum salmon populations from different rivers within the islands. It was shown that in different types of population comparisons, required different marker sets most informative were.     

Genetic Variation and Differentiation among Populations of Chum Salmon Oncorhynchus ketaWalbaum from Southern Russian Far East

Allozyme variation of populations of chum salmon Oncorhynchus ketafrom southern Russian Far East was examined. Of 55 loci screened, 31 were polymorphic. Within-population variation accounted for most of the allele diversity; F _STaveraged over loci was 0.052. Linkage disequilibrium was found in less than 5% of locus pairs in the chum population examined. Analysis of within- and among-population variance components of linkage disequilibrium using D-statistics (Ohta, 1982) showed that most genetic variation was distributed among populations.     

Interregional differentiation of chum salmon from Sakhalin and South Kurils inferred from microsatellite markers

Variability at ten microsatellite loci was examined in wild and hatchery populations of chum salmon (Oncorhynchus keta Walbaum) from the Sakhalin Island and Southern Kuril Islands, Iturup and Kunashir. Substantial genetic differences between Sakhalin and South Kurils chum salmon (the differentiation reached 6.0%) were revealed. Statistically significant differences between chum salmon from Iturup and those from Kunashir were demonstrated, as well as between the chum salmon populations from different rivers within the islands. It was shown that in different types of population comparisons, different marker sets were most informative.     

Genetic Structure of Chum Salmon (Oncorhynchus keta) Populations in the Lower Columbia River: Are Chum Salmon in Cascade Tributaries Remnant Populations?

The lower Columbia River drainage once supported a run of over a million chum salmon. By the late 1950s, the run had decreased to often a few hundred fish. With the exception of Grays River near the coast and an aggregation of chum salmon spawning in creeks and the main stem near Bonneville Dam in the Columbia Gorge, most populations were thought to be extinct. However, chum salmon consistently return in low numbers to tributaries originating in the Cascade Range: the Cowlitz, Lewis, and Washougal rivers. To assess whether Cascade spawners were strays or remnants of former populations, chum salmon from the Coastal, Cascade and Gorge ecoregional zones were characterized at 17 microsatellite loci. Significant heterogeneity in genotype distributions was detected between zones and collections formed regional groups in a neighbor-joining tree. Cascade collections had higher allelic richness and private alleles, and the Cowlitz River supported genetically divergent fall and summer runs, the only summer chum salmon run extant in the Columbia River drainage. We propose that chum salmon in the Cascade zone are remnants of original populations. We attribute the divergence between zonal groups to diverse ecological conditions in each zone, which promoted regional genetic adaptation, and to genetic drift experienced in small populations.     

Genetic structure of freshwater Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis> L.) populations from the lakes Onega and Ladoga of northwest Russia and implications for conservation

Freshwater Atlantic salmon (Salmo salar L.) populations are a living example of adaptation to the changing conditions caused by glacial cycles. The uniqueness of these populations is emphasized by almost complete resistance to the dangerous parasite Gyrodactylus salaris. In Europe, freshwater salmon populations occur primarily in north-western Russia in the republic of Karelia. These systems include Lakes Ladoga and Onega, the two largest lakes in Europe, each of which harbours a number of freshwater salmon spawning rivers. We used microsatellite markers to study the genetic structure and temporal stability of 11 freshwater salmon populations in Russian Karelia. Populations clustered according their region of origin. Although temporal variation in allele frequencies was observed in the majority of temporal comparisons, various lines of evidence demonstrated that this influence was relatively minor compared to spatial variation that explained eight times more of the variability than temporal variation. Temporal stability tended to occur in populations from rivers with a higher linear lake coefficient. The high level of genetic structuring observed in both lake systems and the apparent low level of migration between populations suggests that treating each river as a separate management unit is recommended. In addition, as the number of populations is large, the best strategy for such fine scale management would be to ensure that the level of natural reproduction in each river is sufficient to sustain the population. A prioritization strategy for population conservation based on estimating the relative roles of different evolutionary forces shaping the gene pools highlighted a number of populations where further monitoring is warranted and also identified populations which could be prioritized for conservation as living gene banks in the event that conservation resources are limited. This prioritization agreed well with the occurrence of temporal (in)stability.