Preliminary data on variation of four microsatellite loci in Pacific herring <Emphasis Type="Italic">Clupea pallasii</Emphasis>

Variation of microsatellite loci Cpa110, Cpa113, Cpa4, and Cpa7 was for the first time examined in Pacific-type herring Clupea pallasii from the White Sea (Cl. pallasii marisalbi), the Kara Sea (Cl. pallasii suworowi), the Sea of Okhotsk, and Lake Nerpich’e, Kamchatka Bay, northwestern Pacific (Cl. pallasii pallasii). All loci exhibitedhigh genetic diversity. The estimates of expected heterozygosity varied from 41.5 to 95.6% (mean, 82%). The level of pairwise genetic differentiation F _st at all microsatellite loci varied from 0.005 to 0.076 (0.019, on average) and t was statistically significant (p < 0.05) in most of the pairs of herring samples. Estimates of genetic differentiation among the herring of one subspecies were lower than between the groups belonging to different subspecies.     

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 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.     

Analysis of microsatellite loci variations in herring (Clupea pallasii marisalbi) from the White Sea

The genetic diversity among spawning groups of herring from different parts of the White Sea was assessed using ten microsatellite loci. All loci were polymorphic with the expected heterozygosity estimates varying in the range of 12.7–94.1% (mean was 59.5%). The degree of genetic differentiation displayed by White Sea herring was statistically significant (θ = 2.03%). The level of pairwise genetic differentiation F _ST was 0–0.085, and it was statistically significant in most of the comparison pairs between the herring samples. A hierarchical analysis of molecular variance (AMOVA) revealed the statistically significant differentiation of White Sea herring. 96.59% genetic variation was found within the samples and 3.41% variation was found among the populations. The main component of interpopulation diversity (1.85%) falls at the differences between two ecological forms of herring, spring- and summer-spawning. Within the spring-spawning form, the presence of local stocks in Kandalaksha Bay, Onega Bay, and Dvina Bay was demonstrated.     

Temporal stability of the population genetic structure of the White Sea herring <Emphasis Type="Italic">Сlupea pallasii marisalbi</Emphasis>

The length–age distributions and genetic variation of ten microsatellite loci were examined in samples of herring from the Chupa Inlet of the Kandalaksha Gulf of the White Sea. The samples were collected on the same spawning grounds over a period of 13 years and during the same spawning season. The obtained data demonstrate stability of the length structure and genetic characteristics (allele and genotype frequencies, allelic richness, and heterozygosity). The degree of temporal genetic differentiation was not statistically significant, θ = 0.0038 (CI,–0.0006 to 0.006). Spatial differentiation of the White Sea herring far outweighs the level of temporal variability, which makes it possible to suggest the important role of reproductive isolation in the formation of the population genetic structure of herring.     

Population structure and variability of Pacific herring (<Emphasis Type="Italic">Clupea pallasii</Emphasis>) in the White Sea,Barents and Kara Seas revealed by microsatellite DNA analyses

Pacific herring, Clupea pallasii, have recently colonised the northeast Atlantic and Arctic Oceans in the early Holocene. In a relatively short evolutionary time, the herring formed a community with a complex population structure. Previous genetic studies based on morphological, allozyme and mitochondrial DNA data have supported the existence of two herring subspecies from the White Sea and eastern Barents and Kara Seas (C. p. marisalbi and C. p. suworowi, respectively). However, the population structure of the White Sea herring has long been debated and remains controversial. The analyses of morphological and allozyme data have previously identified local spawning groups of herring in the White Sea, whereas mtDNA markers have not revealed any differentiation. We conducted one of the first studies of microsatellite variation for the purpose of investigating the genetic structure and relationship of Pacific herring among ten localities in the White Sea, the Barents Sea and the Kara Sea. Using classical genetic variance-based methods (hierarchical AMOVA, overall and pairwise F _ST comparisons), as well as the Bayesian clustering, we infer considerable genetic diversity and population structure in herring at ten microsatellite loci. Genetic differentiation was the most pronounced between the White Sea (C. p. marisalbi) versus the Barents and Kara seas (Chesha–Pechora herring, C. p. suworowi). While microsatellite variation in all C. pallasii was considerable, genetic diversity was significantly lower in C. p. suworowi, than in C. p. marisalbi. Also, tests of genetic differentiation were indicating significant differentiation within the White Sea herring between sympatric summer- and spring-spawning groups, in comparison with genetic homogeneity of the Chesha–Pechora herring.     

Evidence for Two Highly Differentiated Herring Groups at Goose Bank in the Barents Sea and the Genetic Relationship to Pacific Herring, Clupea pallasi

Genetic studies on Atlantic herring, Clupea harengus, have generally revealed a low level of genetic variation over large geographic areas. Genetically distinct herring populations in some of the Norwegian fjords are exceptions, and juvenile herring from the large oceanic herring, Norwegian Spring Spawners (NSS), are often found in mixture with local fjord populations as well as widely distributed in the Barents Sea. Research surveys in the eastern Barents Sea (Goose Bank) in 1993, 1994 and 2001 included collection of herring samples for allozyme analyses. As expected the results identified juveniles from NSS stock, but an additional unique group of herring (low vertebrae number), being almost fixed for alternative alleles at several allozyme loci, was detected. In some cases, the two groups of herring were taken in the same trawl catches as documented by highly significant departure from Hardy—Weinberg expectation with large excess of homozygotes providing evidence for population mixing. Large genetic differences (Nei's genetic distance = 1.53; F_ST = 0.754) were detected in pairwise comparisons based on five allozyme loci. The two herring groups were also compared with reference samples of Pacific herring, Clupea pallasi, including one sample from Japan Sea and three Alaskan samples. UPGMA dendrogram based on five allozyme loci revealed a close genetic relationship between the low vertebrae herring in the Barents Sea and the group of samples of Pacific herring. Although significant different in allele frequencies, one of the herring samples clustered together with the reference sample from Bering Sea with genetic distance of 0.008 and F_ST value of 0.032. The close genetic relationship found in this paper, suggest a re-evaluation of the taxonomic status of the Barents Sea herring populations investigated.     

Detecting population structure in a high gene-flow species,Atlantic herring (Clupea harengus): direct,simultaneous evaluation of neutral vs putatively selected loci

In many marine fish species, genetic population structure is typically weak because populations are large, evolutionarily young and have a high potential for gene flow. We tested whether genetic markers influenced by natural selection are more efficient than the presumed neutral genetic markers to detect population structure in Atlantic herring (Clupea harengus), a migratory pelagic species with large effective population sizes. We compared the spatial and temporal patterns of divergence and statistical power of three traditional genetic marker types, microsatellites, allozymes and mitochondrial DNA, with one microsatellite locus, Cpa112, previously shown to be influenced by divergent selection associated with salinity, and one locus located in the major histocompatibility complex class IIA (MHC-IIA) gene, using the same individuals across analyses. Samples were collected in 2002 and 2003 at two locations in the North Sea, one location in the Skagerrak and one location in the low-saline Baltic Sea. Levels of divergence for putatively neutral markers were generally low, with the exception of single outlier locus/sample combinations; microsatellites were the most statistically powerful markers under neutral expectations. We found no evidence of selection acting on the MHC locus. Cpa112, however, was highly divergent in the Baltic samples. Simulations addressing the statistical power for detecting population divergence showed that when using Cpa112 alone, compared with using eight presumed neutral microsatellite loci, sample sizes could be reduced by up to a tenth while still retaining high statistical power. Our results show that the loci influenced by selection can serve as powerful markers for detecting population structure in high gene-flow marine fish species.     

Intraspecies structure of the pacific herring <Emphasis Type="Italic">Clupea pallasii</Emphasis> valenciennes, 1847 (Clupeidae: Clupeiformes) in the Sea of Japan and the southern Sea of Okhotsk inferred from the variability of mitochondrial DNA control

The differentiation of local herring populations from the Sea of Japan and the southern Sea of Okhotsk was assessed on the basis of mtDNA (mitochondrial DNA) control region variability. Three samplings of sea herring and one sampling of lake herring were studied. The genetic analysis showed a significant (P < 0.01) difference between lake and sea herring samplings and no differentiation among sea samplings (P > 0.05). AMOVA analysis revealed that only 4.11% of the variability was attributed to the interpopulation component, while variability within populations was 95.89%. It is suggested that the reason for the weak differentiation of local sea herring populations may be an intensive gene flow.     

New host and locality records for two protozoan (Myxosporea, Coccidia) parasites of marine fishes

This is the first report of the myxosporean Ortholinea orientalis from Atlantic herring Clupea harrengus. It infects the kidney tubules and previously was known from Pacific herring Clupea pallasii and navaga Eleginus navaga in the White Sea and North Pacific Ocean. This is also the first report of the coccidian Eimeria raibauti from Norway pout Trisopterus esmarkii. It infects the epithelium of the pyloric ceca and previously was known only from poorcod Trisopterus minutus in the Mediterranean Sea. The new records are both from the northern North Sea.     

Population genomic evidence for adaptive differentiation in the Baltic Sea herring

Detecting and estimating the degree of genetic differentiation among populations of highly mobile marine fish having pelagic larval stages is challenging because their effective population sizes can be large, and thus, little genetic drift and differentiation is expected in neutral genomic sites. However, genomic sites subject to directional selection stemming from variation in local environmental conditions can still show substantial genetic differentiation, yet these signatures can be hard to detect with low‐throughput approaches. Using a pooled RAD‐seq approach, we investigated genomewide patterns of genetic variability and differentiation within and among 20 populations of Atlantic herring in the Baltic Sea (and adjacent Atlantic sites), where previous low‐throughput studies and/or studies based on few populations have found limited evidence for genetic differentiation. Stringent quality control was applied in the filtering of 1 791 254 SNPs, resulting in a final data set of 68 182 polymorphic loci. Clear differentiation was identified between Atlantic and Baltic populations in many genomic sites, while differentiation within the Baltic Sea area was weaker and geographically less structured. However, outlier analyses – whether including all populations or only those within the Baltic Sea – uncovered hundreds of directionally selected loci in which variability was associated with either salinity, temperature or both. Hence, our results support the view that although the degree of genetic differentiation among Baltic Sea herring populations is low, there are many genomic regions showing elevated divergence, apparently as a response to temperature‐ and salinity‐related natural selection. As such, the results add to the increasing evidence of local adaptation in highly mobile marine organisms, and those in the young Baltic Sea in particular.     

A microsatellite-based estimation of clonal diversity and population subdivision in Zostera marina, a marine flowering plant

We examined the genetic population structure in eelgrass (Zostera marina L.), the dominant seagrass species of the northern hemisphere, over spatial scales from 12 km to 10 000 km using the polymorphism of DNA microsatellites. Twelve populations were genotyped for six loci representing a total of 67 alleles. Populations sampled included the North Sea (four), the Baltic Sea (three), the western Atlantic (two), the eastern Atlantic (one), the Mediterranean Sea (one) and the eastern Pacific (one). Microsatellites revealed substantial genetic variation in a plant group with low allozyme diversity. Average expected heterozygosities per population (monoclonal populations excluded) ranged from 0.32 to 0.61 (mean = 0. 48) and allele numbers varied between 3.3 and 6.7 (mean = 4.7). Using the expected frequency of multilocus genotypes within populations, we distinguished ramets from genetic individuals (i.e. equivalent to clones). Differences in clonal diversity among populations varied widely and ranged from maximal diversity (i.e. all ramets with different genotype) to near or total monoclonality (two populations). All multiple sampled ramets were excluded from further analysis of genetic differentiation within and between populations. All but one population were in Hardy-Weinberg equilibrium, indicating that Zostera marina is predominantly outcrossing. From a regression of the pairwise population differentiation with distance, we obtained an effective population size Ne of 2440-5000. The overall genetic differentiation among eelgrass populations, assessed as rho (a standardized estimate of Slatkin's RST) was 0.384 (95% CI 0.34-0.44, P < 0.001). Genetic differentiation was weak among three North Sea populations situated 12-42 km distant from one another, suggesting that tidal currents result in an efficient exchange of propagules. In the Baltic and in Nova Scotia, a small but statistically significant fraction of the genetic variance was distributed between populations (rho = 0.029-0. 053) at scales of 15-35 km. Pairwise genetic differentiation between European populations were correlated with distance between populations up to a distance of 4500 km (linear differentiation-by-distance model, R2 = 0.67). In contrast, both Nova Scotian populations were genetically much closer to North Sea and Baltic populations than expected from their geographical distance (pairwise rho = 0.03-0.08, P < 0.01). A biogeographical cluster of Canadian with Baltic/North Sea populations was also supported using a neighbour-joining tree based on Cavalli-Sforza's chord distance. Relatedness between populations may be very different from predictions based on geographical vicinity.     

Marine landscapes and population genetic structure of herring (Clupea harengus L.) in the Baltic Sea

Numerically small but statistically significant genetic differentiation has been found in many marine fish species despite very large census population sizes and absence of obvious barriers to migrating individuals. Analyses of morphological traits have previously identified local spawning groups of herring (Clupea harengus L.) in the environmentally heterogeneous Baltic Sea, whereas allozyme markers have not revealed differentiation. We analysed variation at nine microsatellite loci in 24 samples of spring-spawning herring collected at 11 spawning locations throughout the Baltic Sea. Significant temporal differentiation was observed at two locations, which we ascribe to sympatrically spawning but genetically divergent 'spawning waves'. Significant differentiation was also present on a geographical scale, though pairwise F(ST) values were generally low, not exceeding 0.027. Partial Mantel tests showed no isolation by geographical distance, but significant associations were observed between genetic differentiation and environmental parameters (salinity and surface temperature) (0.001 < P < or = 0.099), though these outcomes were driven mainly by populations in the southwestern Baltic Sea, which also exhibits the steepest environmental gradients. Application of a novel method for detecting barriers to gene flow by combining geographical coordinates and genetic differentiation allowed us to identify two zones of lowered gene flow. These zones were concordant with the separation of the Baltic Sea into major basins, with environmental gradients and with differences in migration behaviour. We suggest that similar use of landscape genetics approaches may increase the understanding of the biological significance of genetic differentiation in other marine fishes.     

Characterization of a major histocompatibility class II A gene (Clha-DAA) with an embedded microsatellite marker in Atlantic herring (Clupea harengus L.)

An Atlantic herring major histocompatibility class II A ( Clha-DAA ) cDNA sequence has been characterized and was shown to encode a leader peptide, alpha-1 domain, alpha-2 domain, connecting peptide, transmembrane and cytoplasmic region. The Clha-DAA protein sequence has all the characteristics of a teleost class II A protein with conserved cysteines in both the alpha-1 and the alpha-2 domains and two potential N-linked glycosylation sites. Exon 2 sequences encoding the polymorphic alpha-1 domain from different individuals were analysed and revealed the presence of at least two loci. The Clha-DAA gene consists of four exons and three short introns. Four unique intron 3 sequences from multiple individuals were obtained and were shown to contain a (TG)_n microsatellite sequence. Primers were optimized such that only a single microsatellite locus designated Clha-DAA-INTR3 was amplified. Four herring populations from the North Sea and the Baltic Sea were genotyped for Clha-DAA-INTR3 . In total, 16 Clha-DAA-INTR3 alleles were detected; the distribution of the alleles showed no deviation from Hardy–Weinberg expectation. Levels of genetic differentiation among samples were of similar magnitude as have been reported earlier for neutral microsatellite loci between northern North Sea and Baltic Sea herring populations.     

Variability of Morphobiological Characteristics of the Pacific Herring Clupea pallasii from Arctic and Pacific Populations

Journal of Ichthyology - A comparative analysis of the biological and morphometric characteristics of the Pacific herring Clupea pallasii from different parts of the range, from the White Sea to...     

An analysis of allozyme variation in herring <Emphasis Type="Italic">Clupea pallasii</Emphasis> from the White and Barents Seas

An analysis of genetic variation is made according to four allozyme loci of reproductive (spawning) groups of oligovertebrate herring Clupea pallasii collected in various bays of the White Sea and the south-eastern part of the Barents Sea in 1995–2002. The temporal stability of genetic characteristics during several years is shown. The analysis of genetic variation revealed a significant difference between herring from the south-eastern part of the Barents Sea and spring-spawning and summer-spawning herring from inner bays of the White Sea. The analysis of geographic variation of genetic characteristics of spawning aggregations revealed the change in frequencies of alleles of loci LDH-2* and MDH-4* from the north-east to the south-west along the coast.     

Analysis of genetic diversity and phylogenetic relationships among red jungle fowls and Chinese domestic fowls

Genetic diversity and phylogenetic relationships among 568 individuals of two red jungle fowl subspe- cies (Gallus gallus spadiceus in China and Gallus gallus gallus in Thailand) and 14 Chinese domestic chicken breeds were evaluated with 29 microstaellite loci, the genetic variability within population and genetic differentiation among population were estimated, and then genetic diversity and phylogenetic relationships were analyzed among red jungle fowls and Chinese domestic fowls. A total of 286 alleles were detected in 16 population with 29 microsatellite markers and the average number of the alleles observed in 29 microsatellite loci was 9.86±6.36. The overall expected heterozygosity of all population was 0.6708±0.0251, and the number of population deviated from Hardy-Weinberg equilibrium per locus ranged from 0 to 7. In the whole population, the average of genetic differentiation among population, measured as FST value, was 16.7% (P<0.001), and all loci contributed significantly (P<0.001) to this differentiation. It can also be seen that the deficit of heterozygotes was very high (0.015) (P<0.01). Reynolds' distance values varied between 0.036 (Xiaoshan chicken-Luyuan chicken pair) and 0.330 (G. gallus gallus-Gushi chicken pair). The Nm value ranged from 0.533 (between G. gallus gallus and Gushi chicken) to 5.833 (between Xiaoshan chicken and Luyuan chicken). An unrooted consensus tree was constructed using the neighbour-joining method and the Reynolds' genetic distance. The heavy-body sized chicken breeds, Luyuan chicken, Xiaoshan chicken, Beijing Fatty chicken, Henan Game chicken, Huainan Partridge and Langshan chicken formed one branch, and it had a close genetic relationship between Xiaoshan chicken-Luyuan chicken pair and Chahua chicken-Tibetan chicken pair. Chahua chicken and Tibetan chicken had closer genetic relationship with these two subspecies of red jungle fowl than other domestic chicken breeds. G. gallus spadiceus showed closer phylogenetic relationship with Chinese domestic chicken breeds than G. gallus gallus. All 29 microstaellite loci in this study showed high levels of polymorphism and significant genetic differentiation was observed among two subspecies of red jungle fowl and 14 Chinese domestic chicken breeds. The evolutional dendrogram is as follows: evolutional breeds→primitive breeds (Chahua chicken and Tibetan)→red jungle fowl in China (G. gallus spadiceus)→red jungle fowl in Thailand (G. gallus gallus). The results supported the theory that the domestic fowls might originate from different subspecies of red jungle fowl and Chinese domestic fowls had independent origin.     

Genetic diversity of silver pomfret (Pampus argenteus) in the Southern Yellow and East China Seas

Silver pomfret (Pampus argenteus) is an important fisheries resource in China, but harvest management lacks identification of management units (genetic stocks or populations). Using amplified fragment length polymorphism (AFLP), we investigated the population genetic structure of silver pomfret from the Yellow and East China Seas. In 143 individuals examined from six locations (three locations in the Yellow Sea, three in the East China Sea), a total of 194 loci were detected by six primer combinations. In all loci, 149 loci (76.8%) were polymorphic, and the percentage of polymorphic loci varied within samples from 62.9% to 67.0%. Among the six samples, Nei’s genetic diversity was 0.23 ± 0.20–0.25 ± 0.21, and Shannon’s index was 0.34 ± 0.28–0.36 ± 0.29. Analyses revealed significant genetic differentiation among the six samples, indicating at least six separate populations of silver pomfret in the Southern Yellow and East China Seas.     

Genetic Relationship between Lake and Marine Forms of Pacific Herring Clupea pallasii

Journal of Ichthyology - Polymorphism in the variable fragment of the mtDNA control region (D-loop, 373 bp) and four microsatellite loci in 683 specimens of the Pacific herring Clupea pallasii...