Analysis of the Genetic Structure of Northwestern Bering Sea Walleye Pollock, Theragra chalcogramma

The intraspecific structure of major populations of the walleye pollock Theragra chalcogramma from the northwestern part of the Bering Sea was studied. Specimens from Navarin, Olyutor and Shirshov shoals were sampled in the 2001 spawning season. Preliminary cluster analysis (TREECON) of PCR-RAPD data revealed the existence of cluster with low level of bootstrap support, which generally corresponds to geographic localization of the shoals. Processing of microsatellite sequence data (loci Tch12, Tch14, Tch15 and Tch18) with GENEPOP demonstrated the deviation from HWE and differentiation between the samples. The value of the inter-population variance (F_st = 0.02) corresponded to published data on marine stocks, which were subject to high levels of gene flow. The Shirshov group was found to be equidistant from the Navarin and Olyutor groups, with the genetic distance between the latter two being significantly less. Analysis of the microsatellite loci inferred disparities in their selective capacities. Loci Tch12 and Tch15 are found to be highly homozygous with low levels of polymorphism. Tch12 clearly follows the ‘3 band’ pattern. Tch15 shows sign of linkage disequilibrium. Comparison of F_st values for the loci with the mean standardized variance f₀ suggests that Tch12 is under pressure of disruptive selection, while Tch15 is rather neutral and Tch14 and Tch18 are under the influence of balance selection. Also a discussion is conducted in the context of undeterminate group migration.     

Microsatellite Variability of Pacific Herring <Emphasis Type="Italic">Clupea pallasii</Emphasis> Valenciennes, 1847 from the Sea of Okhotsk and Bering Sea

The genetic variability of ten microsatellite loci was examined in samples of the herring from the Sea of Okhotsk and the Bering Sea. All loci were polymorphic; the expected heterozygosity estimates varied in the range of 0.3–94.3% (mean 66.7%). The degree of genetic differentiation of the herring was statistically significant (θ = 1.38%). The level of pairwise genetic differentiation F_ST was–0.002–0.046; R_ST was–0.003–0.166. Genetic differentiation of the herring from the Sea of Okhotsk and the Bering Sea correlated with the spatial-geographic structure of the species in the studied range on the basis of F_ST (P = 0.001).     

Genetic differentiation of yellowfin sole (<Emphasis Type="Italic">Limanda aspera</Emphasis>) from Taui Bay in the Okhotsk Sea

The population genetic structure of yellowfin sole inhabiting the northern part of the Okhotsk Sea has been investigated. The genetic heterogeneity of samples collected in geographically remote areas of Taui Bay was determined. The value of genetic differentiation (G _ST = 2.39%) in the studied samples was slightly lower than in those from the Bering Sea (G _ST = 4.25%) (Grant et al., 1983).     

Genetic variations in Clupea pallasii herring from Sea of Okhotsk based on microsatellite markers

The genetic variations among spawning groups of herrings from different spawning grounds of the northwestern part of the Sea of Okhotsk was assessed using ten microsatellite loci. All loci were polymorphic with the expected heterozygosity estimates varying at different loci in the range of 0.7–95.0% (with a mean of 68.5%). The degree of genetic differentiation displayed by the herrings from the Sea of Okhotsk was not statistically significant (θ = 0.74%). The level of pairwise genetic differentiation F _ST varied in the range of 0.002–0.014, nor was it statistically significant in all comparison pairs between the herring samples.     

The genetic variability of the red king crab, <Emphasis Type="Italic">Paralithodes camtschatica</Emphasis> (Tilesius, 1815) (Anomura,Lithodidae) introduced into the Barents Sea compared with samples from the Bering Sea and Kamchatka region using eleven microsatellite loci

The intentional introduction of red king crab, Paralithodes camtschatica (Tilesius, 1815) in the Barents Sea represent one of a few successful cases and one that now supports a commercial fishery. Introductions of alien species into new environments are often associated with genetic bottlenecks, which cause a reduction in the genetic variation, and this could be important for the spreading potential of the species in the Atlantic Ocean. Red king crab samples collected in the Varangerfjord located on the Barents Sea (northern Norway) were compared with reference crab samples collected from the Bering Sea and Kamchatka regions in the Pacific Ocean. All samples were screened for eleven microsatellite loci, based on the development of species-specific primers. The observed number of alleles per locus was similar, and no reduction in genetic variation, including gene diversity and allelic richness, was detected between the Varangerfjord sample and the reference sample from Okhotsk Sea near Kamchatka, indicating no genetic bottlenecking at least for the microsatellite loci investigated. The same results were found in comparison with the sample from Bering Sea. The level of genetic differentiation among the samples, measured as overall F _ST across all loci, was relatively low (0.0238) with a range of 0.0035–0.1000 for the various loci investigated. The largest pairwise F _ST values were found between the Bering Sea and Varangerfjord/Barents Sea samples, with a value of 0.0194 across all loci tested. The lowest value (0.0101) was found between the Varangerfjord and Kamchatka samples. Genetic differentiation based on exact tests on allele frequencies revealed highly significant differences between all pairwise comparisons. The high level of genetic variation found in the Varangerfjord/Barents Sea sample could be of significance with respect to further spreading of the species to other regions in the North Atlantic Ocean.     

Mitochondrial DNA Variation in Northwestern Bering Sea Walleye Pollock, Theragra chalcogramma (Pallas)

Genetic variability of the highly valuable gadid species, walleye pollock, Theragra chalcogramma (Pallas 1811), from five spatially separated northwestern Bering Sea areas (Ozernoi Bay, Olutorskyi Bay, Koryak shelf, Navarin region, and Anadyr Gulf) was investigated. Haplotype diversity within the samples ranged from 0.8788±0.0393 to 0.9436±0.0162. Nucleotide diversity within the samples ranged from 0.0108 to 0.0127. Nucleotide diversity among the regional collections ranged from 0% to 0.18%. Walleye pollock from the Anadyr Gulf appeared genetically separate from the other four samples in a clustering of genetic distances. Total heterogeneity among all five samples was significant, while there was no heterogeneity among the four samples excluding that from the Anadyr Gulf. Pair-wise comparisons using tests for heterogeneity and F_st supported the dendrogram of genetic distances in that only the collection from the Anadyr Gulf was significantly different from the others. The observed pattern of genetic differentiation among walleye pollock from the northwestern Bering Sea presumably emerged as a result of a population of the species subdividing in the northernmost part of its geographic range, though further analysis is needed to verify this supposition.     

Reproductive conditions of walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> (Gadidae) off the northeastern coast of Sakhalin Island,Sea of Okhotsk

The spatial distribution of eggs and larvae of the walleye pollock Theragra chalcogramma is considered in respect to dynamics of oceanologic processes, nutrients, chlorophyll а and zooplankton off the northeastern coast of Sakhalin Island in spring 2012. It is shown that the effect of severe temperature regime in the near-bottom horizons in the western Sea of Okhotsk during the spawning period of walleye pollock becomes milder due to specific features of water dynamics. The egg distribution is determined by mesoscale eddies in the region. The species survival depends on the effect of such environmental factors as freshwater discharge from the Amur River, eddy structure in waters of the Sea of Okhotsk, and dynamics of phytoplankton and zooplankton development.     

Study of the filiform gonads of walleye pollock <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> inhabiting the Sea of Okhotsk and the Bering Sea

The histological study of the adult specimens of walleye pollock Theragra chalcogramma carrying filamentous gonads evidenced that this group comprised males, females, and sterile fish. The specimens carrying the filamentous gonads have been found mostly in the elder age groups; no maturing of the oocytes or sperm cells has been registered. The frequency of occurrence of such specimens was relatively low: 0.07–0.13% in the Sea of Okhotsk and up to 0.25% in the Bering Sea. We conclude that these specimens have nearly no effect on the reproductive success of this species.     

Spawning of walleye pollock (<Emphasis Type="Italic">Theragra chalcogramma</Emphasis>) off the Southwestern coast of Kamchatka

The results of ichthyoplankton surveys conducted in 2007–2008 showed that waters off Southwest Kamchatka and North Kuril Islands were areas of mass spawning of walleye pollock. The peak spawning occurred during the last 10 days of April and in early May, which was much later than the peak at the main spawning site off West Kamchatka. The spawning activity of walleye pollock near the southwestern shores of Kamchatka is a regular event, as the analysis of archive materials shows. This gave us grounds to suspect the existence of a southern site that coincided well with the mass spawning in the Shelikhov Gulf in its timing and scale, but was missed during standard ichthyoplankton surveys conducted in early April. After analyzing the growth rates of spawners, the assumption was made that the southern spawning was performed by the East Kamchatkan walleye pollock population, whose mass spawning usually occurs in late April-early May. According to the data of 2008, the estimated biomass of walleye pollock spawning in the area of the Ozernovskaya basin in late April was nearly 600000 tons. Regular monitoring of the southern spawning is proposed by means of additional ichthyoplankton surveys south of 53° N, including the Okhotsk Sea waters of the North Kuril Islands, in late April—early May.     

Variability of the Characteristics of the First Scale Annulus and Length of Year-Old Fish in the Bering Sea Walleye Pollock Theragra chalcogramma

Geographic and interannual variability of the number of annuli and the radius of the first scale annulus, as well as the retrospective length of year-old walleye pollock from the western, northern, and eastern parts of the Bering Sea, was examined using data for the years 1995, 1996, 1998, and 1999. The characteristics of the first scale annulus were varying. By these parameters, walleye pollock from the western Bering Sea significantly differs statistically from walleye pollock of the eastern Bering Sea. With respect to number of annuli, the radius of the first ring, and the retrospective length of year-old specimens, the walleye pollock from the Navarinskii region occupies an intermediate position between fish from the western and eastern parts of the Bering Sea. Interannual variability of the three parameters was found for walleye pollock from the Navarinskii region.     

A mitogenomic approach to the taxonomy of pollocks: <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> and <Emphasis Type="Italic">T. finnmarchica</Emphasis> represent one single species

Background  

The walleye pollock (Theragra chalcogramma) and Norwegian pollock (T. finnmarchica) are confined to the North Pacific and North Atlantic Oceans, respectively, and considered as distinct species within the family Gadidae. We have determined the complete mtDNA nucleotide sequence of two specimens of Norwegian pollock and compared the sequences to that of 10 specimens of walleye pollock representing stocks from the Sea of Japan and the Bering Sea, 2 specimens of Atlantic cod (Gadus morhua), and 2 specimens of haddock (Melanogrammus aeglefinus).     

Some biological features and productive characteristics of <Emphasis Type="Italic">Oithona similis</Emphasis> (Copepoda) in the Sea of Okhotsk and western Bering Sea

The biological peculiarities of Oithona similis were described based on the materials from 29 complex surveys that were conducted by the TINRO Center in the Sea of Okhotsk and the western Bering Sea from 1986 to 2005 and the production of the species was estimated. In this area, O. similis produces four generations a year, viz., one in the spring, one in the fall, and two in the summer. The peak of spawning in May and June coincides with the maximum development of phytoplankton. Somatic production of O. similis in both seas becomes higher during the summer season. In this species, the largest increment in body weight begins with the copepodite stage CIV, when basic nutrients are accumulated. The average specific daily production of O. similis in the epipelagic zone of the Sea of Okhotsk and Bering Sea was 0.0092 and 0.01 in the spring, 0.043 and 0.031 in the summer, and 0.023 and 0.014 in the fall, respectively. The total somatic production of this species reached 28.318 million tons in the Sea of Okhotsk and 4.811 million tons in the Bering Sea. The Sea of Okhotsk is more favorable for the development of O. similis.     

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.     

Springtime microprotozoan abundance and biomass in the southeastern Bering Sea and Shelik of Strait, Alaska

We surveyed springtime biomass and abundance of the >20 µmmicroprotozoa in surface waters of the SE Bering Sea and Shelikof Strait, Alaska. This study was part of the Fisheries OceanographyCoordinated Investigations (FOCI) program examining processeswhich affect the recruitment variability of walleye pollock(Theragra chalcogramma). Microprotozoa are a potential preyresource for larval pollock which has not been previously examined.In both areas, the >20 µm microprotozoa were predominantlydinoflagellates and ciliates. At the time of sampling (May 1990in Shelikof Strait and April 1992 in the SE Bering Sea), thespring diatom bloom was under way in Shelik of Strait, but notin the SE Bering Sea. Heterotrophic dinoflagellates dominatedthe microprotozoan assemblage in Shelik of Strait, but not inthe SE Bering Sea. In the SE Bering Sea. total microprotozoanabundances ranged from 300 to 6233 organisms 1^–1 and biomassfrom 0.58 to 9.73 µg C 1^–1. In Shelik of Strait,abundance and biomass were higher, ranging from 850 to 14 960organisms 1^–1 and from 1.29 to 70.73 µg C 1^–1,respectively. These biomass levels are comparable to those reportedfrom other coastal and oceanic regions. Microprotozoan biomasslevels were sufficient to support the estimated metabolic needsof first-feeding larval walleye pollock. It remains to be shownwhether larval pollock use this resource.     

Genetic differentiation of pink salmon Oncorhynchus gorbuscha Walbaum in the Asian part of the range

Genetic variation at 19 enzyme (including 11 polymorphic) and 10 microsatellite loci was examined in the population samples of odd-and even-broodline pink salmon from the southern part of Sakhalin Island, Southern Kuril Islands, and the northern coast of the Sea of Okhotsk. The estimates of relative interpopulation component of genetic variation for the allozyme loci, per broodline, were on average 0.43% (G _ST), while over the microsatellite loci it was 0.26% (the ?_ST coefficient, F-statistics based on the allele frequency variance), and 0.90% (the ρ_ST coefficient, R-statistics based on the allele size variance). The values of interlinear component constituted 2.34, 0.31, and 1.05% of the total variation, respectively. Using the allozyme loci, statistically significant intralinear heterogeneity was demonstrated among the regions, as well as among the populations of southern Sakhalin. Multidimensional scaling based on the allozyme data demonstrated regional clustering of the sample groups, representing certain populations during the spawning run or in different years. Most of the microsatellite loci examined were found to be highly polymorphic (mean heterozygosity > 0.880). The estimates of interlinear, interregional, and interpopulation variation over these loci in terms of ?_ST values were substantially lower than in terms of ρ_ST values. Regional genetic differentiation, mostly expressed at the allozyme loci between the populations from the northern Sea of Okhotsk and the Sakhalin and Kuril group of populations, was less expressed at the microsatellite loci. The differentiation between these regions observed can be considered as the evidence in favor of a large-scale isolation by distance characterizing Asian pink salmon. It is suggested that in pink salmon, low genetic differentiation at neutral microsatellite loci can be explained by extremely high heterozygosity of the loci themselves, as well as by the migration gene exchange among the populations (the estimate of the gene migration coefficient inferred from the “private” allele data constituted 2.6 to 3.4%), specifically, by the ancient migration exchange, which occurred during postglacial colonization of the range     

Social and reproductive behavior of a captive group of walleye pollock,Theragra chalcogramma

Synopsis The social and reproductive behavior of a group of four male and seven female walleye pollock,Theragra chalcogramma, were observed in a large tank. Pollock spent most of their time swimming in a loose aggregation near the surface. Males descended from the aggregation more often than females to follow and make physical contact with other males as well as with females. The difference between males and females in the frequency of diving in our tank is consistent with the reported pattern of depth segregation of the sexes in natural pollock spawning aggregations. The frequency of social interactions increased when pollock became reproductively active and was higher at night and during twilight when most of the spawning occurred. Male interactions with females most frequently involved physical contact, while male interactions with other males were more often limited to following. There was no indication that male-male interactions result in the formation of stable social dominance relationships that determine priority of access to mates, as has been suggested previously for walleye pollock. Rather, following and contact interactions appear to promote male identification of potential mates and encounters with ripe females. The possible functional significance of male social interactions is discussed in relation to reports on natural walleye pollock spawning aggregations.     

Assessment of the consumption of walleye pollock eggs by nekton and jellyfish in the northern Sea of Okhotsk in the spring

In the northern Sea of Okhotsk, nekton and jellyfish consumed as many as 831 × 10⁹ walleye pollock eggs per day in 2011. The nekton exerted the highest pressure, viz., 98.3% of the overall predation on pollock egg by aquatic animals. Of the entire quantity of consumed eggs, 55.9% were eaten by herring, 35.9% by walleye pollock, 6.5% by Sakhalin sole, and 1.7% by jellyfish. Among jellyfish, scyphomedusae Cyanea capillata and Chrysaora melonaster, as well as the hydromedusa Tima sachalinensis consumed the largest quantities of eggs. The total consumption of pollock egg by aquatic animals in 2011 was estimated at 42.4 × 10¹², or 11.4% of the entire quantity of eggs that were spawned by walleye pollock in the waters of the northern part of the sea. The total amount of pollock eggs that were eaten by herring and pollock together for 51 days in 2011 amounted to 38.9 × 10¹², which was 5.7 times as much as that in 2002. Thus, a significant growth of predation on pollock eggs by their main consumers, viz., herring and walleye pollock, was observed in 2011. This was caused by an increase in the populations of both species during the recent years and also by a higher concentration of pollock eggs.     

Evaluation of the productivity of walleye pollock generations in the northern Sea of Okhotsk

Based on data on the abundances of various age groups in the northern Sea of Okhotsk in 1984–2008, the productivity of 33 generations of walleye pollock born between 1975 and 2007 was evaluated in two ways. The first approach allowed a retrospective estimation by abundance of the first nine one-year age groups, and the second one was a perspective estimation using the abundance index of the nearest recruitment—3- and 4-year-olds—that enabled us to evaluate the productivity of a generation before it enters into spawning and commercial stocks. These productivity estimates were compared to the data of annual catches and the dynamics of spawning stock for many years. A high value of walleye pollock stock was shown to be supported by several productive and highly productive generations, a medium value—by one productive and one moderately productive or by two moderately productive generations, and a low value—by one productive, or moderately productive, and low productive generations. In addition, overall mortality rates were calculated for generations with various productivity levels. The greatest variability of mortality rates was observed in immature walleye pollock; the maximum mortality rate was typical for the recruitment of highly abundant generations, and the minimum, for the least abundant ones. The lowest mortality was recorded at the age of mass maturation. Mature individuals of generations with various productivity levels showed insignificant differences in overall mortality rates.     

The size composition of the pelagic eggs of walleye pollack <Emphasis Type="Italic">Theragra chalcogramma</Emphasis> in spawning areas of the northern Sea of Okhotsk

The data on the size composition of the pollack developing eggs of walleye pollack collected in the northern Sea of Okhotsk in 2001 during mass spawning are given. The eggs caught in western Kamchatka and in Shelikhov Bay had the greatest diameter; they were somewhat less in the northern Okhotsk Sea area; and the smallest eggs were found in eastern Sakhalin. The clinal variability of the size composition of walleye pollack eggs was observed in the northern Sea of Okhotsk, and in the western and eastern parts it had a local pattern. Parameters such as spawning female size composition, temperature, and salinity, as well their influence on the size composition of eggs, were considered.Original Russian Text Copyright © 2004 by Biologiya Morya, Ovsyannikov.     

Population genetics of Atlantic bluefin tuna,Thunnus thynnus (Linnaeus, 1758), in the Mediterranean: implications for its conservation management

The movement of Atlantic bluefin tuna (ABFT) across international boundaries necessitates traceability strategies that would provide more accurate information needed for stock assessment. The Mediterranean Sea is one of the main contributors to ABFT reproduction and global population genetic diversity. In the present study this genetic variability was investigated using 193 samples of adult bluefin tuna from Spain, Turkey and Malta – a longitudinal distance of 3400 km. Analysed were 13 microsatellite loci (eight of which were newly‐tested) as genetic markers for the population study. Allele richness measured per locus and sampling location varied from 1.89 to 8.88, taking into account rarefaction. ABFT private alleles were detected in each of the three sampling sites. No significant spatial genetic divergence was found between pairs at the studied locations (F_ST values <0.0001; P‐values >0.05). Bayesian clustering analysis corroborated a single and panmictic ABFT population in the Mediterranean Sea. Statistical power analyses indicated a high probability of detecting genetic differentiation and population structure with the sample size and microsatellites used, even at an F_ST value of 0.005. From the results it may be postulated that migrating ABFT during the spawning season are allowing gene flow within the Mediterranean Sea. The complex interplay of movements, including plasticity in the selection of spawning sites with increasing age and environmental conditions, require multiple and new fisheries monitoring and management techniques in order to target the ABFT long‐term conservation effectively.