Microsatellite identification of individual sockeye salmon in Barkley Sound, British Columbia

Population structure of sockeye salmon Oncorhynchus nerka in Barkley Sound, British Columbia, Canada was examined by analysis of microsatellite variation at 14 loci in three populations sampled in each of 3 years. The mean F _ST over all 14 loci was 0·063. Differences among populations accounted for 12 times the variation observed among years within populations. The number of alleles present at a locus was related to the power of the locus to provide accurate identification of individuals to population. The more alleles that were present at a locus, the greater was the power of the locus for individual identification. Individuals were correctly classified to one of three lakes of origin at a rate of 89%, and to one of two river drainages at a rate of 96%.     

DNA Microarray for Rapid Detection of Mitochondrial DNA Haplotypes of Chum Salmon

For use in genetic stock identification, we developed an oligonucleotide (DNA) microarray hybridization method for rapid and accurate detection of nucleotide sequence variations in 20 previously identified variable nucleotide sites in about 500 bp within the 5 half of the control region of mitochondrial DNA of chum salmon (Oncorhynchus keta). The method includes immobilization of synthesized oligonucleotides containing respective polymorphic sites on a glass slide precoated with polycarbodiimide resin, a 2-hour hybridization with DNA microarray of biotinylated polymerase chain reaction fragments spanning the 5 variable portion followed by short washing, and visualization of hybridization signals by conventional ABC method and scanner-assisted computation of signal intensity on a computer. The entire process of hybridization and detection was completed within 4 hours. The resulting DNA microarray could detect all of the single nucleotide mutations and therefore could be used to identity the sequence variations defining 30 mtDNA haplotypes of chum salmon as revealed previously by nucleotide sequence analysis.     

Tendencies of Density Dynamics in Reophilic and Lacustrine Sockeye Salmon Oncorhynchus nerkain the Basin of Azabach'e Lake

Temporal series of density in the spawners of two sockeye salmon populations from Azabach'e Lake (Kamchatka River basin) were analyzed using periodic functions. It has been found that the quality of describing the density dynamics for sockeye salmon by such models is very high (R ²= 0.943 and R ²= 0.905 for early and late populations, respectively). The density (for early sockeye salmon) and total number (for late sockeye salmon) distributions of the reproductive fraction of the populations have been developed and analyzed.     

Parallel signatures of selection at genomic islands of divergence and the major histocompatibility complex in ecotypes of sockeye salmon across Alaska

Understanding the genetic mechanisms that facilitate adaptive radiation is an important component of evolutionary biology. Here, we genotyped 82 neutral SNPs, seven SNPs in islands of divergence identified in a previous study (island SNPs), and a region of the major histocompatibility complex (MHC) in 32 populations of sockeye salmon to investigate whether conserved genes and genomic regions are involved in adaptive radiation. Populations representing three ecotypes were sampled from seven drainages with differing habitats and colonization histories spanning a range of 2,000 km. We found strong signatures of parallel selection across drainages at the island SNPs and MHC, suggesting that the same loci undergo divergent selection during adaptive radiation. However, patterns of differentiation at most island SNPs and the MHC were not associated with ecotypes, suggesting that these loci are responding differently to a mosaic of selective pressures. Our study provides some of the first evidence that conserved genomic islands may be involved in adaptive divergence of salmon populations. Additionally, our data provide further support for the hypothesis that sockeye salmon inhabiting rivers unconnected to lakes harbour similar genetic diversity across large distances, are likely the ancestral form of the species, and have repeatedly recolonized lake systems as they have become available after glacial recession. Finally, our results highlight the value and importance of validating outlier loci by screening additional populations and regions, a practice that will hopefully become more common in the future.     

Major histocompatibility complex diversity is positively associated with stream water temperatures in proximate populations of sockeye salmon

Local adaptation to heterogeneous environments generates population diversity within species, significantly increasing ecosystem stability and flows of ecosystem services. However, few studies have isolated the specific mechanisms that create and maintain this diversity. Here, we examined the relationship between water temperature in streams used for spawning and genetic diversity at a gene involved in immune function [the major histocompatibility complex (MHC)] in 14 populations of sockeye salmon (Oncorhynchus nerka) sampled across the Wood River basin in south‐western Alaska. The largest influence on MHC diversity was lake basin, but we also found a significant positive correlation between average water temperature and MHC diversity. This positive relationship between temperature and MHC diversity appears to have been produced by natural selection at very local scales rather than neutral processes, as no correlation was observed between temperature and genetic diversity at 90 neutral markers. Additionally, no significant relationship was observed between temperature variability and MHC diversity. Although lake basin was the largest driver of differences in MHC diversity, our results also demonstrate that fine‐scale differences in water temperature may generate variable selection regimes in populations that spawn in habitats separated by as little as 1 km. Additionally, our results indicated that some populations may be reaching a maximum level of MHC diversity. We postulate that salmon from populations in warm streams may delay spawning until late summer to avoid thermal stress as well as the elevated levels of pathogen prevalence and virulence associated with warm temperatures earlier in the summer.     

Interannual Variations of the Body Length in Sockeye Salmon Oncorhynchus nerka Smolts of Lake Kuril'skoe in Relation to External Factors

A nonlinear regression model of body length dynamics was constructed for age smolts greater than two years old of the sockeye salmon Oncorhynchus nerka of Lake Kuril'skoe (Kamchatka). The analysis of variance showed that about 93% of the variation of body length is accounted for by the fish population density and water temperature. The reasons for the increase in the body length of smolts after volcanic ashfall are discussed.     

Signals of heterogeneous selection at an MHC locus in geographically proximate ecotypes of sockeye salmon

The genes of the major histocompatibility complex (MHC) are an important component of the vertebrate immune system and can provide insights into the role of pathogen‐mediated selection in wild populations. Here, we examined variation at the MHC class II peptide‐binding region in 27 populations of sockeye salmon (Oncorhynchus nerka), distributed among three distinct spawning ecotypes, from a complex of interconnected rivers and lakes in south‐western Alaska. We also obtained genotypes from 90 putatively neutral single nucleotide polymorphisms for each population to compare the relative roles of demography and selection in shaping the observed MHC variation. We found that MHC divergence was generally partitioned by spawning ecotype (lake beaches, rivers and streams) and was 30 times greater than variation at neutral markers. Additionally, we observed substantial differences in modes of selection and diversity among ecotypes, with beach populations displaying higher levels of directional selection and lower MHC diversity than the other two ecotypes. Finally, the level of MHC differentiation in our study system was comparable to that observed over much larger geographic ranges, suggesting that MHC variation does not necessarily increase with increasing spatial scale and may instead be driven by fine‐scale differences in pathogen communities or pathogen virulence. The low levels of neutral structure and spatial proximity of populations in our study system indicate that MHC differentiation can be maintained through strong selective pressure even when ample opportunities for gene flow exist.     

Genetic Stock Identification of Chum Salmon in the Bering Sea and North Pacific Ocean Using Mitochondrial DNA Microarray

A newly developed DNA microarray was applied to identify mitochondrial (mt) DNA haplotypes of more than 2200 chum salmon in the Bering Sea and North Pacific Ocean in September 2002 and also 2003, when the majority of maturing fish were migrating toward their natal river. The distribution of haplotypes occurring in Asian and North American fish in the surveyed area was similar in the 2 years. A conditional maximum likelihood method for estimation of stock compositions indicated that the Japanese stocks were distributed mainly in the north central Bering Sea, whereas the Russian stocks were mainly in the western Bering Sea. The North American stocks were abundant in the North Pacific Ocean around the Aleutian Islands. These results indicate that the Asian and North American stocks of chum salmon are nonrandomly distributed in the Bering Sea and the North Pacific Ocean, and further the oligonuleotide DNA microarray developed by us has a high potential for identification of stocks among mixed ocean aggregates of high-seas chum salmon.     

Gene diversity analysis of mitochondrial DNA, microsatellites and allozymes in landlocked Atlantic salmon

This study investigates the patterns of genetic diversity detected in allozymes, mtDNA, and microsatellites, in order to assess their relative efficacy to differentiate sympatric landlocked salmon populations and to estimate changes in genetic diversity between wild and first-generation hatchery fish. Overall, the three genetic markers indicated a genetic differentiation between two sympatric populations of Lake Saint-Jean, Québec. MtDNA and microsatellites also showed significant differences between wild and first-generation hatchery fish originating from the same river. Allozyme analysis was the most limited approach due to the low genetic diversity detected and the necessity to kill specimens. Although low polymorphism was found in mtDNA, it was the most discriminant marker between wild populations. Microsatellite analysis appears to be a promising approach due to its high sensitivity in differentiating wild populations, in detecting changes in allele composition between wild and first-generation hatchery fish and its potential for increased resolution by augmenting the number of polymorphic loci. Given the benefits and disadvantages of the three methods, the combination of mtDNA and microsatellite analyses will best address our research objectives.     

Genetic characterization of specific pathogen‐free rhesus macaque (Macaca mulatta) populations at the California National Primate Research Center (CNPRC)

A study based on 14 STRs was conducted to understand intergenerational genetic changes that have occurred within the California National Primate Research Center's (CNPRC) regular specific pathogen‐free (SPF) and super‐SPF captive rhesus macaque populations relative to their conventional founders. Intergenerational genetic drift has caused age cohorts of each study population, especially within the conventional population, to become increasingly differentiated from each other and from their founders. Although there is still only minimal stratification between the conventional population and either of the two SPF populations, separate derivation of the regular and super‐SPF animals from their conventional founders has caused the two SPF populations to remain marginally different from each other. The regular SPF and, especially, the super‐SPF populations have been influenced by the effects of differential ancestry, sampling, and lost rare alleles, causing a substantial degree of genetic divergence between these subpopulations. The country of origin of founders is the principal determinant of the MHC haplotype composition of the SPF stocks at the CNPRC. Selection of SPF colony breeders bearing desired genotypes of Mamu‐A^*01 or ‐B^*01 has not affected the overall genetic heterogeneity of the conventional and the SPF research stocks. Because misclassifying the ancestry of research stocks can undermine experimental outcomes by excluding animals with regional‐specific genotypes or phenotypes of importance, understanding founder/descendent genetic relationships is crucial for investigating candidate genes with distinct geographic origins. Together with demographic management, population genetic assessments of SPF colonies can curtail excessive phenotypic variation among the study stocks and facilitate successful production goals. Am. J. Primatol. 72:587–599, 2010. © 2010 Wiley‐Liss, Inc.     

Differences in Responses of Chinook Salmon to Climate Shifts: Implications for Conservation

Understanding how organisms respond to climate is critical for focusing the debate about ways to recover imperiled or manage exploited species. However, efforts to understand climate effects on biota are complicated by differences among species in life history and physiology. Even within a species it is not clear if different populations will react similarly to large-scale climate trends. Climate regimes exhibit basin-wide effects similar to the El Ni no Southern Oscillation but persist for decades. In the North Pacific Ocean, two regime shifts (abrupt changes from one regime to another) occurred in 1976–1977 and 1989–1990 and had wide ranging effects on many species. We examined the response of chinook salmon from 9 evolutionary significant units (ESUs) to the regime shifts. While there was an average decline in spawner numbers associated with the regime shifts, ESUs did not respond in a uniform manner: some ESUs declined, some did not respond and one may have increased. Four ESUs currently listed under the Endangered Species Act may have declined more across regime boundaries than did the five non-listed ones. Interpretation of this result depends on two ESUs: the Snake River spring/summer run and the Central Valley fall run. The Snake River ESU had the largest decline and most sampling effort. If this ESU was excluded from the analysis, there was no evidence that listed and non-listed stocks responded differently to the regimes. The Central Valley ESU is currently a candidate for listing. If this ESU is considered to be a threatened or endangered, then listed ESUs declined more on average than did non-listed ESUs across the regime boundaries regardless of the Snake River ESU. As a whole, these results suggest that long-term climate trends are important to the dynamics of chinook populations and that sub-units of a species (here ESUs) can respond differently to these regimes.     

Revisiting the Freedom of Fishing and Legal Obligations on States Not Party to Regional Fisheries Management Organizations

For decades, states have cooperated through regional fisheries management organizations (RFMOs) on the conservation and management of living marine resources on the high seas. Nonmembers, or third states not bound by their decisions, have been an Achilles’ heel. In this article, the legal status of RFMO nonmembers and the freedom of fishing are examined. It is concluded that RFMO nonmembers have concrete obligations with respect to established fisheries subject to RFMO regulation.     

MHC class IIB alleles contribute to both additive and nonadditive genetic effects on survival in Chinook salmon

The genes of the major histocompatibility complex (MHC) are found in all vertebrates and are an important component of individual fitness through their role in disease and pathogen resistance. These genes are among the most polymorphic in genomes and the mechanism that maintains the diversity has been actively debated with arguments for natural selection centering on either additive or nonadditive genetic effects. Here, we use a quantitative genetics breeding design to examine the genetic effects of MHC class IIB alleles on offspring survivorship in Chinook salmon (Oncorhynchus tshawytscha). We develop a novel genetic algorithm that can be used to assign values to specific alleles or genotypes. We use this genetic algorithm to show simultaneous additive and nonadditive effects of specific MHC class IIB alleles and genotypes on offspring survivorship. The additive effect supports the rare-allele hypothesis as a potential mechanism for maintaining genetic diversity at the MHC. However, contrary to the overdominance hypothesis, the nonadditive effect led to underdominance at one heterozygous genotype, which could instead reduce variability at the MHC. Our algorithm is an advancement over traditional animal models that only partition variance in fitness to additive and nonadditive genetic effects, but do not allocate these effects to specific alleles and genotypes. Additionally, we found evidence of nonrandom segregation during meiosis in females that promotes an MHC allele that is associated with higher survivorship. Such nonrandom segregation could further reduce variability at the MHC and may explain why Chinook salmon has one of the lowest levels of MHC diversity of all vertebrates.     

Geographic variation of the major histocompatibility complex in Eastern Atlantic grey seals (Halichoerus grypus)

Pathogen-driven balancing selection maintains high genetic diversity in many vertebrates, particularly in the major histocompatibility complex (MHC) immune system gene family, which is often associated with disease susceptibility. In large natural populations where subpopulations face different pathogen pressures, the MHC should show greater genetic differentiation within a species than neutral markers. We examined genetic diversity at the MHC-DQB locus and nine putatively neutral microsatellite markers in grey seals (Halichoerus grypus) from eight United Kingdom (UK) colonies, the Faeroe Islands and Sable Island, Canada. Five DQB alleles were identified in grey seals, which varied in prevalence across the grey seal range. Among the seal colonies, significant differences in DQB allele and haplotype frequencies and in average DQB heterozygosity were observed. Additionally, the DQB gene exhibited greater differentiation among colonies compared with neutral markers, yet a weaker pattern of isolation by distance (IBD). After correcting for the underlying IBD pattern, subpopulations breeding in similar habitats were more similar to one another in DQB allele frequencies than populations breeding in different habitats, but the same did not hold true for microsatellites, suggesting that habitat-specific pathogen pressure influences MHC evolution. Overall, the data are consistent with selection at MHC-DQB loci in grey seals with both varying selective pressures and geographic population structure appearing to influence the DQB genetic composition of breeding colonies.     

A comparison of variability and population structure for major histocompatibility complex and microsatellite loci in California coastal steelhead (Oncorhynchus mykiss Walbaum)

The major histocompatibility complex (MHC) contains genes integral to immune response in vertebrates. MHC genes have been shown to be under selection in a number of vertebrate taxa, making them intriguing for population genetic studies. We have conducted a survey of genetic variation in an MHC class II gene for steelhead trout from 24 sites in coastal California and compared this variation to that observed at 16 presumably neutral microsatellite loci. A high amount of allelic variation was observed at the MHC when compared to previously published studies on other Pacific salmonids. Elevated nonsynonymous substitutions, relative to synonymous substitutions, were detected at the MHC gene, indicating the signature of historical balancing selection. The MHC data were tested for correlations to and deviations from the patterns found with the microsatellite data. Estimates of allelic richness for the MHC gene and for the microsatellites were positively correlated, as were estimates of population differentiation (F(ST)). An analysis for F(ST) outliers indicates that the MHC locus has an elevated F(ST) relative to the neutral expectation, although a significant result was found for only one particular geographical subgroup. Relatively uniform allele frequency distributions were detected in four populations, although this finding may be partially due to recent population bottlenecks. These results indicate that, at the scale studied here, drift and migration play a major role in the observed geographical variability of MHC genes in steelhead, and that contemporary selection is relatively weak and difficult to detect.     

Dynamics of Mhc evolution in birds and crocodilians: amplification of class II genes with degenerate primers

Genes of the major histocompatibility complex (Mhc) are the most polymorphic functional loci in mammalian populations, but little is known of Mhc variability in natural populations of nonmammalian vertebrates. To help extend such studies to birds and relatives, we present a pair of degenerate primers that amplify polymorphic segments of one chain (the β chain) of the class II genes from the major histocompatibility complex (Mhc) of archosaurs (birds + crocodilians). The primers target two conserved regions lying within portions of the antigen-binding site (ABS) encoded by the second exon and amplify multiple genes from both genomic DNA and cDNA. The pattern of nucleotide substitution in ABS codons of 51 sequences amplified and cloned from five species of passerine birds and an alligator (Alligator mississippiensis) indicates that archosaurian class II β genes are subject to selective forces similar to those operating in mammalian populations. Hybridization of a genomic clone generated by the primers revealed highly polymorphic bands in a sample of Florida scrub jays (Aphelocoma coerulescens coerulescens). Because the primers amplify only part of the ABS from multiple class II genes, they will be useful primarily for generating species specific clones, thereby providing a critical inroad to more detailed structural and evolutionary studies.     

Major histocompatibility complex heterozygote advantage and widespread bacterial infections in populations of Chinook salmon (Oncorhynchus tshawytscha)

Despite growing evidence for parasite-mediated selection on the vertebrate major histocompatibility complex (MHC), little is known about variation in the bacterial parasite community within and among host populations or its influence on MHC evolution. In this study, we characterize variation in the parasitic bacterial community associated with Chinook salmon ( Oncorhynchus tshawytscha ) fry in five populations in British Columbia (BC), Canada across 2 years, and examine whether bacterial infections are a potential source of selection on the MHC. We found an unprecedented diversity of bacteria infecting fry with a total of 55 unique bacteria identified. Bacterial infection rates varied from 9% to 29% among populations and there was a significant isolation by distance relationship in bacterial community phylogenetic similarity across the populations. Spatial variation in the frequency of infections and in the phylogenetic similarity of bacterial communities may result in differential parasite-mediated selection at the MHC across populations. Across all populations, we found evidence of a heterozygote advantage at the MHC class II, which may be a source of balancing selection on this locus. Interestingly, a co-inertia analysis indicated only susceptibility associations between a few of the MHC class I and II alleles and specific bacterial parasites; there was no evidence that any of the alleles provided resistance to the bacteria. Our results reveal a complex bacterial community infecting populations of a fish and underscore the importance of considering the role of multiple pathogens in the evolution of host adaptations.     

Variation in the timing of river entry of Atlantic salmon （Salmo salar L. ） in the Baltic

The timing of river entry in the Atlantic salmon is known to depend on genetic, demographic and environmental factors, but little is known about the relative magnitude of among population and among year variation and covariation in this respect in natural state Atlantic salmon rives. To investigate this, variability in the timing of river entry in three historical Finnish Atlantic salmon populations were analyzed using salmon trap data collected during 1870- 1902. The analyses reveled that 1 ） the timing of river entry differed substantially and consistently among the rivers, and that 2） variation among the rivers was much larger than variation among years. Annual variations were not explained by regional environmental conditions, whereas in one river the timing of the local flood peak was a significant predictor of the timing of river entry. Differences in the timing of salmon entry to geographically closely situated rivers suggests that a regionally fixed opening date for coastal fisheries might not be the best management strategy as it may lead to uneven exploitation of salmon populations from different rivers [ Current Zoology 55 （5） ： 342 - 349, 2009] .     

Variation in the timing of river entry of Atlantic salmon (Salmo salar L.) in the Baltic

The timing of river entry in the Atlantic salmon is known to depend on genetic,demographic and environmental factors,but little is known about the relative magnitude of among population and among year variation and covariation in this respect in natural state Atlantic salmon rives.To investigate this,variability in the timing of river entry in three historical Finnish Atlantic salmon populations were analyzed using salmon trap data collected during 1870-1902.The analyses reveled that 1)the timing of river e...     

Applications of DNA barcoding to fish landings: authentication and diversity assessment

DNA barcoding methodologies are being increasingly applied not only for scientific purposes but also for diverse real-life uses. Fisheries assessment is a potential niche for DNA barcoding, which serves for species authentication and may also be used for estimating within-population genetic diversity of exploited fish. Analysis of single-sequence barcodes has been proposed as a shortcut for measuring diversity in addition to the original purpose of species identification. Here we explore the relative utility of different mitochondrial sequences (12S rDNA, COI, cyt b, and D-Loop) for application as barcodes in fisheries sciences, using as case studies two marine and two freshwater catches of contrasting diversity levels. Ambiguous catch identification from COI and cyt b was observed. In some cases this could be attributed to duplicated names in databases, but in others it could be due to mitochondrial introgression between closely related species that may obscure species assignation from mtDNA. This last problem could be solved using a combination of mitochondrial and nuclear genes. We suggest to simultaneously analyze one conserved and one more polymorphic gene to identify species and assess diversity in fish catches.