Genetic changes in pink salmon Oncorhynchus gorbuscha Walbaum during acclimatization in the White sea basin

Genetic parameters of pink salmon introduced into the White Sea basin in 1985 and 1998 were compared to the corresponding parameters of the donor population from the Ola River (Magadan oblast). The detected genetic differences indicate that colonization of a new area is accompanied by impoverishment of the gene pool of the native population. This effect was particularly marked in the odd-year line of pink salmon introduced in 1985. The probable causes of these genetic changes are discussed.     

Genetic divergence in pink salmon introduced into the European north of Russia: microsatellite and allozyme variation analysis

The 1985 introduction into the European North of Russia resulted in the formation of a large stock of pink salmon of the odd-year breeding line. To assess the divergence of the new population and the role of various microevolutionary factors, variation of four microsatellite loci and fifteen genes encoding proteins (allozymes) in samples of fish, running for spawning in rivers of the new area, and in samples from the donor population of the Ola River (Magadan oblast). In the generations 8 and 9 of the introduced pink salmon of the odd-year line, the genetic diversity (the number of alleles and the mean heterozygosity) both at allozyme and at microsatellite loci was significantly lower, than that in the donor population. The explanations of the decline in diversity are discussed. The first evidence for spatial genetic divergence in transplanted fish within the new area has been obtained; the divergence level may be comparable with that characteristic of native populations.     

Genetic divergence in pink salmon introduced into the European North of Russia: Microsatellite and allozyme variation analysis

The 1985 introduction into the European North of Russia resulted in the formation of a large stock of pink salmon of the odd-year broodline. To assess the divergence of the new population and the role of various microevolutionary factors, variation of four microsatellite loci and fifteen genes encoding proteins (allozymes) in samples of fish, running for spawning in rivers of the new area, and in samples from the donor population of the Ola River (Magadan region). In the generations 8 and 9 of the introduced pink salmon of the odd-year line, the genetic diversity (the number of alleles and the mean heterozygosity) both at allozyme and at microsatellite loci was significantly lower, than that in the donor population. The explanations of the decline in diversity are discussed. The first evidence for spatial genetic divergence in transplanted fish within the new area has been obtained; the divergence level may be comparable with that characteristic of native populations.     

Experimental microevolution: transplantation of pink salmon into the European North

Human-mediated translocations of species beyond their native ranges can enhance evolutionary processes in populations introduced to novel environments. We studied such processes in several generations of pink salmon Oncorhynchus gorbuscha introduced to the European North of Russia using a set of morphological and life-history traits as well as molecular genetic markers with different selective values: protein-coding loci, mtDNA, microsatellites, and MHC. The introduction of reproductively isolated pink salmon broodlines of odd and even years yielded different results. The odd-year broodline established self-reproducing local populations in many rivers of new range, but sustainable changes in external morphology, reproduction, and life-history, as well as the impoverishment of the gene pool occurred. Their successful colonisation of the new range resulted in specialisation manifested in the rapid directional shifts in some highly heritable phenotypic traits accompanied by increased homozygosity at molecular markers as a consequence of genetic drift and selective processes. The returns of transplanted pink salmon of even-year broodline decreased sharply already in the second generation, but there was no marked reduction of genetic diversity. Our data, as well as the analysis of the history of all pink salmon transplantations beyond the species range, demonstrate comparatively greater success of introduced odd-year broodline and permit to assume different adaptive plasticity of the even- and odd-year broodlines in pink salmon, what is most likely determined by differences in their evolutionary histories. Population genetic data suggest that the even-year broodline probably diverged from the odd-year broodline relatively recently and, due to the founder effect, may have lost a part of its genetic variation with which adaptive plasticity potential is associated.     

Straying of hatchery salmon in Prince William Sound,Alaska

The straying of hatchery salmon may harm wild salmon populations through a variety of ecological and genetic mechanisms. Surveys of pink (Oncorhynchus gorbuscha), chum (O. keta) and sockeye (O. nerka) salmon in wild salmon spawning locations in Prince William Sound (PWS), Alaska since 1997 show a wide range of hatchery straying. The analysis of thermally marked otoliths collected from carcasses indicate that 0–98% of pink salmon, 0–63% of chum salmon and 0–93% of sockeye salmon in spawning areas are hatchery fish, producing an unknown number of hatchery-wild hybrids. Most spawning locations sampled (77%) had hatchery pink salmon from three or more hatcheries, and 51% had annual escapements consisting of more than 10% hatchery pink salmon during at least one of the years surveyed. An exponential decay model of the percentage of hatchery pink salmon strays with distance from hatcheries indicated that streams throughout PWS contain more than 10% hatchery pink salmon. The prevalence of hatchery pink salmon strays in streams increased throughout the spawning season, while the prevalence of hatchery chum salmon decreased. The level of hatchery salmon strays in many areas of PWS are beyond all proposed thresholds (2–10%), which confounds wild salmon escapement goals and may harm the productivity, genetic diversity and fitness of wild salmon in this region     

Gonadal Development of Diploid and Triploid Pink Salmon (<Emphasis Type="Italic">Oncorhynchus gorbuscha</Emphasis>) from the White Sea

Only males (fully mature and failing to mature) and intersexes have been found among triploids of pink salmon. Possible mechanisms of genetic sex determination in pink salmon are discussed. The possibility of using triploids to regulate sizes of artificial pink salmon populations is also discussed. Triploids of pink salmon are deemed inefficient to use in aquaculture. Delayed maturation is observed in some diploid females reared in farming cages.     

Estimates of population differentiation in pink salmon Oncorhynchus gorbuscha with using of the microsatellite markers may be underestimated due to high population sizes

Data on the variation at eight microsatellite loci in the Far East salmon Oncorhynchus gorbuscha samples caught in 1984–1985 and 2001–2006 are analyzed. F-statistics indices at all levels of the hierarchical spatial structures are very small. At the same time, the differentiation between populations (according to F _ST estimates) in the odd-year broodline of pink salmon does not exceed the temporal variation within populations. In the even-year broodline, the F-statistics indices at the interregional and intraregional levels are significantly greater than those in the odd-year broodline. F _ST estimates (averaged over the same set of loci) vary widely within the range: the highest values are observed in populations of the coast of North America (except Alaska) and in the new range in the European North of Russia, whereas in the populations of the Asian part of the range and Alaska they are one order of magnitude smaller. The causes of the heterogeneity of the estimates of genetic differentiation within the range and between the broodlines of odd and even years are discussed. Since the mean population size estimates were correlated with the F _ST values, it was assumed that the effect of random genetic drift, the main factor of population divergence in selectively neutral loci, weakens with an increase in the population size. Because of the greater population sizes in pink salmon compared to other salmon species, as well due to the uneven distribution of populations of different size, the usage of microsatellite markers may lead to an underestimation of the true divergence of populations and their regional groups and, consequently, to an overestimation of genetic migration.     

Parallel signatures of selection in temporally isolated lineages of pink salmon

Studying the effect of similar environments on diverse genetic backgrounds has long been a goal of evolutionary biologists with studies typically relying on experimental approaches. Pink salmon, a highly abundant and widely ranging salmonid, provide a naturally occurring opportunity to study the effects of similar environments on divergent genetic backgrounds due to a strict two‐year semelparous life history. The species is composed of two reproductively isolated lineages with overlapping ranges that share the same spawning and rearing environments in alternate years. We used restriction‐site‐associated DNA (RAD) sequencing to discover and genotype approximately 8000 SNP loci in three population pairs of even‐ and odd‐year pink salmon along a latitudinal gradient in North America. We found greater differentiation within the odd‐year than within the even‐year lineage and greater differentiation in the southern pair from Puget Sound than in the northern Alaskan population pairs. We identified 15 SNPs reflecting signatures of parallel selection using both a differentiation‐based method (BAYESCAN) and an environmental correlation method (BAYENV). These SNPs represent genomic regions that may be particularly informative in understanding adaptive evolution in pink salmon and exploring how differing genetic backgrounds within a species respond to selection from the same natural environment.     

Genetic stock structure of odd-year pink salmon, Oncorhynchus gorbuscha (Walbaum), from Washington and British Columbia and potential mixed-stock fisheries applications

We used electrophoresis to determine the number and characteristics of genetically distinct stocks of odd-year pink salmon in Washington and southern British Columbia. We analysed 5128 fish from 52 collections (taken in 1985, 1987 and 1989). We observed genetic variation at 53 enzyme-coding loci, 19 of which were polymorphic at the P_o-95 level in at least one stock. Genotypic proportions conformed to Hardy-Weinberg expectations in nearly all cases. The genetic profiles of individual populations were generally stable over the three cycle years studied. Significant differences in allele frequencies at sAAT-3* , PEP-LT* and PGDH* for several stocks were, however, noted between this study and previously reported data for pink salmon. We used G-tests and cluster analysis of genetic distances to evaluate genetic interrelationships among collections and to define genetically distinct stocks. Differentiation among stocks exhibited a clear geographic pattern with three major clusters of stocks recognizable: (1) Hood Canal and Washington Strait of Juan de Fuca stocks, (2) Puget Sound, Fraser River, and southern Canada South Coast stocks, and (3) northern Canada South Coast stocks and Canada North Coast stocks. Computer simulations using 14 and 28 loci, and sample sizes of 15C600, demonstrated that accurate estimates of stock-group composition could be obtained for pink salmon fisheries having a considerable range of stock compositions. The simulations revealed that approximately 50% fewer fish were required to obtain a given level of precision of stock group composition estimates with 28 loci as with the set of 14 loci used in previous investigations.     

Annual changes in the population size of the salmon louse Lepeophtheirus salmonis (Copepoda: Caligidae) on high-seas Pacific Salmon (Oncorhynchus spp.), and relationship to host abundance

The population size of the salmon louse, Lepeophtheirus salmonis, was monitored annually in the summers of 1991–1997 by examining six species of Pacific salmon (Oncorhynchus spp.) caught by surface long-lines in oceanic offshore waters of the North Pacific Ocean and Bering Sea. The annual copepod population size on all salmonids caught was estimated by combining the calculated number of copepods carrying on each salmonid species. The copepod population fluctuated markedly from year to year, which resulted largely from marked annual changes in abundance of pink salmon (O. gorbuscha). Since pink salmon were most frequently and heavily infected and since their abundance changed every year, the copepod population was high in the years when this salmonid species was abundant, but low when it was rare. On the contrary, chum salmon (O. keta) did not show high prevalence and intensity of infection, but the annual abundance of this host species was consistently high, i.e. chum salmon carried many copepods every year. Copepods on other salmonid species (sockeye salmon O. nerka, coho salmon O. kisutch, chinook salmon O. tshawytscha, and steelhead trout O. mykiss) constantly formed a small percentage of the total copepod population. Both chum and pink salmon are the most important hosts in terms of their substantial contribution to support the copepod population, but the importance as hosts of each species is definitely different between the species. Chum salmon is a stable important host supporting the copepod population at a relatively high level every year, while the number of copepods on pink salmon annually exhibits marked fluctuations, and this salmonid species is regarded as an unstable important host.     

Size-selective mortality of Sea of Okhotsk pink salmon in the ocean in the winter and spring

The mortality of Sea of Okhotsk pink salmon in the winter and spring varies significantly from year to year, which complicates forecasts of its arrival in the following year based on data on the downstream migration of fry and surveys in the fall. The size-selective mortality of pink salmon was studied and the possibility of using the size and weight parameters of juveniles for predicting their return was evaluated through measurements of scale increments in juvenile pink salmon that were caught in the southern Sea of Okhotsk in the fall of 2007 and 2008 and in fish of these year classes that came back to spawn. In the 2007 year class, which had a low overwinter survival level in the ocean, the average scale increments for the first year of life were considerably smaller than those in adult fish that returned to the spawning grounds. In the pink salmon of 2008, which had a very high level of overwinter survival, the values of scale increments in juveniles and adults were similar. This confirms the hypothesis of a critical size and a critical period, according to which slowly growing juveniles that do not accumulate enough energy reserves for summer are eliminated in the winter to a greater extent as compared to fast-growing fish. Correlation analysis revealed a significant negative relationship between the size and weight of juvenile pink salmon and their mortality in the ocean. After conducting further and more extensive studies this will allow using the size parameters of juvenile pink salmon as one of predictors of its return for the year following the fall surveys. These results emphasize how important it is to take the size and growth rate of juvenile salmon into account when forecasting their return.     

Genetic markers of adaptive processes in the Far Eastern pink salmon <Emphasis Type="Italic">Oncorhynchus gorbuscha</Emphasis>: Allelic diversity at the locus of major histocompatibility complex MHC I-<Emphasis Type="Italic">A1</Emphasis>

To clarify allelic diversity at the locus of major histocompatibility complex MHC class I-A1 in the Far Eastern pink salmon Oncorhynchus gorbuscha, sequencing of the electrophoretic alleles isolated from the gel (DGGE alleles) was performed. In 47 individuals, the genotypes of which consisted of ten DGGE alleles, 18 MHC I-A1 nucleotide sequences were revealed, and thus, eight cryptic alleles not detected by electrophoresis were identified. Eleven of these alleles were identified earlier in pink salmon from Hokkaido, Alaska, and British Columbia, and seven, possibly, were unique to the populations from some Far Eastern regions. Six of the previously determined DGGE alleles corresponded to more than one nucleotide sequence. However, the sequences attributed to the same DGGE allele differed on average by less than 1 nucleotide. These findings point to sufficient sensitivity of the DGGE method, although the genetic diversity and differentiation estimates obtained with it will obviously be somewhat underestimated. Considerable predominance of nonsynonymous substitutions over the synonymous ones in the codons of the MHC I-A1 antigen-binding site confirms the presence of positive selection aimed at providing the population resistance to local spectrum of pathogens. Refinement of the allelic composition of the adaptively important MHC genetic marker will contribute to more complete understanding of the adaptive genetic structure of pink salmon as an important element of the overall population structure of the species.     

Climate warming causes phenological shift in Pink Salmon, Oncorhynchus gorbuscha, behavior at Auke Creek, Alaska

Thirty‐four years (1972–2005) of water temperature data and extensive biological observations at Auke Creek, Alaska indicate a general warming trend that affected the native pink salmon (Oncorhynchus gorbuscha) population. Serial environmental records at nearby Auke Bay, Alaska over 46 years show trends of increasing air and sea surface temperatures. Trends of increased total precipitation and earlier date of ice out on nearby Auke Lake also occurred, but not at significant rates. Average water temperatures during the incubation of pink salmon in Auke Creek increased at a rate of 0.03 °C yr^?1 over the 34‐year period. For the 1972–2005 broods, midpoints of fry migrations from Auke Creek ranged between April 2 and May 7, and there was a trend of earlier migration of pink salmon fry at a rate of ? 0.5 days yr^?1. The migration timing of adult salmon into Auke Creek also showed a trend toward earlier timing. The earlier adult migration combined with warmer incubation temperatures are related to earlier migration of pink salmon fry. If the observed warming trend continues, Auke Creek may become unsuitable habitat for pink salmon. Given the trend for salmon fry to migrate earlier, a larger portion of the population may become mismatched with optimum environmental conditions during their early marine life history. If salmon adults continue to migrate into the creek earlier when water temperatures are commonly high, it will result in increased prespawning mortality.     

Sex Determination Model in Pink Salmon <Emphasis Type="Italic">Oncorhynchus gorbuscha</Emphasis> (Walbaum, 1792) (Salmonidae,Osteichthyes) Controlled by Multi-Copy Genes Located in Sex Chromosomes

This article is devoted to presenting the hypothesis explaining the fact of a considerable prevalence of phenotypic males among the triploid pink salmon as well as the regular occurrence of intersexes, which were revealed by us. This hypothesis also explains the large proportion (in some cases) in pink salmon populations of the individuals whose genetic sex does not match the phenotypic sex. We assume that the genes encoding the factors that contribute to the transformation of individuals into males (but not the marker sequences of the Y chromosome) are present not only in the Y chromosome of pink salmon but also in the X chromosome, although in smaller quantities.     

On possible re-distribution of pink salmon <Emphasis Type="Italic">Onchorhynchus gorbuscha</Emphasis> between the reproduction areas of different stocks in the Sakhalin-Kuril region

The scale structure of pink salmon Oncorhynchus gorbuscha inhabiting southern Sakhalin and Iturup Island were studied in 2014. These locations are characterized by the unusual dynamics of catches and biological parameters of the fish that come to spawn in different areas in the Sakhalin-Kuril region. The pink salmon that has originated from Iturup Island appeared en masse in the waters of southern Sakhalin, according to the analysis of the scleritogram fragments that reflect the fish growth during the first months of life. This is the first confirmation of the hypothesis of the fluctuating stocks of pink salmon approved by the ichthyological methods. It is suggested that wide-scale straying of pink salmon takes place in the years of dominance shift between the generations of even and odd spawning years.     

Hypotheses concerning the decline and poor recovery of Pacific herring in Prince William Sound, Alaska

This paper updates previous reviews of the 1993 stock decline of Pacific herring (Clupea pallasi) in Prince William Sound, Alaska, and focuses on hypotheses about subsequent poor recovery. Recent age structured assessment modeling with covariate analysis indicates that the population dynamics of the sound’s herring are influenced by oceanic factors, nutrition, and, most substantially, hatchery releases of juvenile pink salmon. For the 1993 decline, poor nutrition remains the most probable cause with disease a secondary response. Concerning poor recovery, we examined 16 potential factors and found three to be causal: oceanic factors, poor nutrition, and hatchery releases of juvenile pink salmon. Absences of strong year classes at both Sitka and Prince William Sound after 1993 indicate the action of large-scale ocean processes. Beyond regional-scale environmental factors, two factors specific to the sound influence the population dynamics of herring and are likely impeding recovery. First, pink salmon fry releases have increased to about 600 million annually and may disrupt feeding in young herring, which require adequate nutrition for growth and overwintering survival. Juvenile pink salmon and age-1 herring co-occur in nearshore areas of bays in late spring and summer, and available data on dietary overlap indicates potential competition between the age-1 juvenile herring and juvenile pink salmon. Field studies demonstrate that juvenile herring reduce food intake substantially in the presence of juvenile pink salmon. Second, overwintering humpback whales may consume potentially large amounts of adult herring, but further studies must confirm to what extent whale predation reduces herring biomass.     

Comparative phylogeography of the two pink salmon broodlines: an analysis based on a mitochondrial DNA genealogy

Over most of their natural northern Pacific Ocean range, pink salmon (Oncorhynchus gorbuscha) spawn in a habitat that was repeatedly and profoundly affected by Pleistocene glacial advances. A strictly two-year life cycle of pink salmon has resulted in two reproductively isolated broodlines, which spawn in alternating years and evolved as temporal replicates of the same species. To study the influence of historical events on phylogeographical and population genetic structure of the two broodlines, we first reconstructed a fine-scale mtDNA haplotype genealogy from a sample of 80 individuals and then determined the geographical distribution of the major genealogical assemblages for 718 individuals sampled from nine Alaskan and eastern Asian even- and nine odd-year pink salmon populations. Analysis of restriction site states in seven polymerase chain reaction (PCR)-amplified mtDNA regions (comprising 97% of the mitochondrial genome) using 13 endonucleases resolved 38 haplotypes, which clustered into five genealogical lineages that differed from 0.065 to 0.225% in net sequence divergence. The lineage sorting between broodlines was incomplete, which suggests a recent common ancestry. Within each lineage, haplotypes exhibited star-like genealogies indicating recent population growth. The depth of the haplotype genealogy is shallow ( approximately 0.5% of nucleotide sequence divergence) and probably reflects repeated decreases in population size due to Pleistocene glacial advances. Nested clade analysis (NCA) of geographical distances showed that the geographical distribution observed for mitochondrial DNA (mtDNA) haplotypes resulted from alternating influences of historical range expansions and episodes of restricted dispersal. Analyses of molecular variance showed weak geographical structuring of mtDNA variation, except for the strong subdivision between Asian and Alaskan populations within the even-year broodline. The genetic similarities observed among and within geographical regions probably originated from postglacial recolonizations from common sources rather than extensive gene flow. The phylogeographical and population genetic structures differ substantally between broodlines. This can be explained by stochastic lineage sorting in glacial refugia and perhaps different recolonization routes in even- and odd-year broodlines.     

Na+/K+-ATPase Activity in Smolts of Pink Salmon Oncorhynchus gorbuscha (Walbaum, 1792) from the White Sea Exposed to Fresh,Estuarine, and Sea Water

Doklady Biological Sciences - The Na+/K+-ATPase (NKA) activity in smolts of pink salmon Oncorhynchus gorbuscha (a salmon species introduced in 1959 into the rivers of the Kola Peninsula) was...     

Variability of cytochrome b gene of mitochondrial DNA of pink salmon Oncorhynchus gorbuscha (Walbaum) from the rivers of the continental coast of the Sea of Okhotsk and Zavyalov Island

Data on the structure and variability of the nucleotide sequence of the cytochrome b (cytb) gene fragment of mitochondrial DNA (mtDNA) in pink salmon of adjacent generations from the rivers of the continental coast of the Sea of Okhotsk and Zavyalov Island were obtained. The differences between adjacent generations were discovered. By the marker studied, pink salmon of the even-year line are characterized by a high level of genetic variability. It is shown that reproductive isolation has led to accumulation of specific mutations of the mtDNA cytb gene in each line and, consequently, to their divergence. Analysis of the data showed that the share of intrapopulation genetic variability of northern Sea of Okhotsk pink salmon accounts for about 91%. The intergroup component that was calculated for adjacent generations of the species is relatively small and is 9%, which seems to be indicative of a relatively recent divergence of adjacent generations of salmon.     

Sex ratio control in pink salmon (Oncorhynchus gorbuscha) and chum salmon (O. keta) populations: the possible causes and mechanisms of changes in the sex ratio

Long-term changes in the sex ratio have been studied in pink salmon (Oncorhynchus gorbuscha) and chum salmon (O. keta) populations of Kamchatka and Sakhalin. It has been demonstrated that these changes are an adaptation to population dynamics: an increase in the population size is accompanied by a shift towards a higher proportion of males; a decrease in population size, by a shift towards a higher proportion of females. The correspondence between morphological and molecular characters in populations of the two species has been analyzed in order to determine the mechanism of sex ratio control. In some pink salmon and chum salmon populations, there is a discrepancy between sex identifications based on morphological characters and molecular markers. This discrepancy is assumed to be accounted for by sex inversion mechanisms, which may be population- or region-specific. In two cases, it has been found that the sex ratio discrepancy in populations is related to the numbers of fish in subsequent generations. These findings suggest that sex inversion may be related to population size control.