Comparisons of reproductive function and fatty acid fillet quality between triploid and diploid farm Atlantic salmon (Salmo salar).

Triploidy could prevent escaped farm salmon breeding in the wild, while also improving nutrient quality within farmed fillets. Despite these potential advantages, triploid Atlantic salmon have not been widely used in aquaculture, and their reproductive function has yet to be fully evaluated. Here, we compare reproductive function and fillet composition between triploid and diploid farm salmon under standard aquaculture rearing conditions. We show that female triploids are sterile and do not develop gonads. By contrast, males produce large numbers of motile spermatozoa capable of fertilizing wild salmon eggs. However, compared with diploids, reproductive development and survival rates of eggs fertilized by triploid males were significantly reduced, with less than 1% of eggs sired by triploid males reaching late-eyed stages of development. Analyses of fillets showed that total lipid and fatty acid quantities were significantly lower in triploid than in diploid Atlantic salmon fillets. However, when fatty acids were normalized to total lipid content, triploid fillets had significantly higher relative levels of important omega-3 long-chain polyunsaturated fatty acids. Our results show that: (i) escaped triploid farm salmon are very unlikely to reproduce in the wild and (ii) if able to match diploid fillet lipid content, triploid farm salmon could achieve better fillet quality in terms of essential fatty acids.

Farmed Atlantic salmon Salmo salar L. parr may reduce early survival of wild fish.

The study examined the density-mediated effects on growth, survival and dispersal of wild and farmed Atlantic salmon Salmo salar offspring in the period immediately following emergence, using a substitutive design. In small confined stream channels, wild parr coexisting with farmed parr had a significantly poorer survival, than wild parr alone. Density did not affect this relationship. In larger unconfined stream channels, wild parr coexisting with farmed parr entered a downstream trap in higher numbers than wild parr in allopatry. The results suggests that during the earliest life stages, farmed S. salar can outcompete wild S. salar, resulting in a reduced survival of wild S. salar.

Microsatellite standardization and evaluation of genotyping error in a large multi-partner research programme for conservation of Atlantic salmon (Salmo salar L.).

Microsatellite genotyping is a common DNA characterization technique in population, ecological and evolutionary genetics research. Since different alleles are sized relative to internal size-standards, different laboratories must calibrate and standardize allelic designations when exchanging data. This interchange of microsatellite data can often prove problematic. Here, 16 microsatellite loci were calibrated and standardized for the Atlantic salmon, Salmo salar, across 12 laboratories. Although inconsistencies were observed, particularly due to differences between migration of DNA fragments and actual allelic size ('size shifts'), inter-laboratory calibration was successful. Standardization also allowed an assessment of the degree and partitioning of genotyping error. Notably, the global allelic error rate was reduced from 0.05 ± 0.01 prior to calibration to 0.01 ± 0.002 post-calibration. Most errors were found to occur during analysis (i.e. when size-calling alleles; the mean proportion of all errors that were analytical errors across loci was 0.58 after calibration). No evidence was found of an association between the degree of error and allelic size range of a locus, number of alleles, nor repeat type, nor was there evidence that genotyping errors were more prevalent when a laboratory analyzed samples outside of the usual geographic area they encounter. The microsatellite calibration between laboratories presented here will be especially important for genetic assignment of marine-caught Atlantic salmon, enabling analysis of marine mortality, a major factor in the observed declines of this highly valued species.

Seawater tolerance in Atlantic salmon, Salmo salar L., brown trout, Salmo trutta L., and S. salar × S. trutta hybrids smolt.

High levels of hybridization between Atlantic salmon (Salmo salar) and brown trout (Salmo trutta) have been reported in the Gyrodactylus salaris infected Rivers Vefsna and Driva in Norway. The survival and behaviour during the sea phase of such hybrids is unknown. The reported work documents ionoregulatory status after 24 h seawater challenge tests (24hSW) and gill Na+/K+-ATPase (NKA) activity of migrating wild smolts of Atlantic salmon, brown trout and hybrids at two sampling dates during the 2006 smolt run in River Driva. Salmon, trout and hybrids contributed to 27, 52 and 21% of the catches, respectively. The large contribution of hybrids suggests both a high hybridization rate and a high survival rate from fry to smolt. Both salmon and hybrids had a well-developed seawater tolerance at the time of downstream migration, revealed by small ionoregulatory effects and no or low mortality rates during the 24hSW tests. The trout were not fully adapted to seawater, and high mortality rates were observed (71 and 92%) during the 24hSW tests. The NKA activity was not significantly different between salmon and hybrids. Most of the hybrids were physiologically capable of direct entry to full strength seawater. The incomplete seawater tolerance in trout compared to salmon corresponds well with differences in life-history patterns between these two species. The life history strategy of the hybrids during the sea phase is not known, and further investigations on the marine behaviour and survival is needed to evaluate the role of hybrids in the risk of spreading G. salaris to nearby river systems.

Genetically enhanced growth causes increased mortality in hypoxic environments.

Rapid growth and development are associated with several fitness-related benefits. Yet, organisms usually grow more slowly than their physiological maximum, suggesting that rapid growth may carry costs. Here we use coho salmon (Oncorhynchus kisutch) eggs of wild and transgenic genotypes to test whether rapid growth causes reduced tolerance to low levels of oxygen (hypoxia). Eggs were exposed to four different durations of hypoxia, and survival and growth were recorded until the end of the larval stage. Survival rates decreased with increasing duration of hypoxia, but this decrease was most pronounced for the transgenic group. Larval mass was also negatively affected by hypoxia; however, transgenic genotypes were significantly larger than wild genotypes at the end of the larval stage. Oxygen can be a limiting factor for survival and development in a wide range of organisms, particularly during the egg stage. Thus, the reduced ability of fast-growing genotypes to cope with low oxygen levels identified in the present study may represent a general constraint on evolution of rapid growth across taxa.

Parallel evolution of ecomorphological traits in the European whitefish Coregonus lavaretus (L.) species complex during postglacial times.

The extensive phenotypic polymorphism in the European whitefish has triggered evolutionary research in order to disentangle mechanisms underlying diversification. To illuminate the ecological distinctiveness in polymorphic whitefish, and evaluate taxonomic designations, we studied nine Norwegian lakes in three watercourses, which each harboured pairs of divergent whitefish morphs. We compared the morphology and life history of these morphs, documented the extent of genetic differentiation between them, and contrasted the niche use of sympatric morphs along both the habitat and resource axes. In all cases, sympatric morphs differed in the number of gill rakers, a highly heritable trait related to trophic utilization. Individual growth rate, age and size at maturity, diet and habitat use also differed between morphs within lakes, but were remarkably similar across lakes within the same morph. Microsatellite analyses confirmed for all but one pair that sympatric morphs were significantly genetically different, and that similar morphs from different lakes likely have a polyphyletic origin. These results are most compatible with the process of parallel evolution through recurrent postglacial divergence into pelagic and benthic niches in each of these lakes. We propose that sparsely and densely rakered whitefish sympatric pairs may be a likely case of ecological speciation, mediated in oligotrophic lakes with few trophic competitors.

Evolutionary history of the European whitefish Coregonus lavaretus (L.) species complex as inferred from mtDNA phylogeography and gill-raker numbers.

We compared mitochondrial DNA and gill-raker number variation in populations of the European whitefish Coregonus lavaretus (L.) species complex to illuminate their evolutionary history, and discuss mechanisms behind diversification. Using single-strand conformation polymorphism (SSCP) and sequencing 528 bp of combined parts of the cytochrome oxidase b (cyt b) and NADH dehydrogenase subunit 3 (ND3) mithochondrial DNA (mtDNA) regions, we documented phylogeographic relationships among populations and phylogeny of mtDNA haplotypes. Demographic events behind geographical distribution of haplotypes were inferred using nested clade analysis (NCA) and mismatch distribution. Concordance between operational taxonomical groups, based on gill-raker numbers, and mtDNA patterns was tested. Three major mtDNA clades were resolved in Europe: a North European clade from northwest Russia to Denmark, a Siberian clade from the Arctic Sea to southwest Norway, and a South European clade from Denmark to the European Alps, reflecting occupation in different glacial refugia. Demographic events inferred from NCA were isolation by distance, range expansion, and fragmentation. Mismatch analysis suggested that clades which colonized Fennoscandia and the Alps expanded in population size 24 500-5800 years before present, with minute female effective population sizes, implying small founder populations during colonization. Gill-raker counts did not commensurate with hierarchical mtDNA clades, and poorly with haplotypes, suggesting recent origin of gill-raker variation. Whitefish designations based on gill-raker numbers were not associated with ancient clades. Lack of congruence in morphology and evolutionary lineages implies that the taxonomy of this species complex should be reconsidered.

Morphological divergence and origin of sympatric populations of European whitefish (Coregonus lavaretus L.) in Lake Femund, Norway.

Combining morphological and genetic analysis, we compared patterns of diversification within and between morphs among sympatric European whitefish (Coregonus lavaretus L.) populations in Lake Femund, Norway. Seven external populations, from potential colonization routes into Lake Femund were included. We found that deep-, shallow-, river- and bay spawning populations are distinct morphs in Lake Femund. Within morphs, populations range from being similar genetically (Fst=0-0.005) among deep-spawning populations to being highly differentiated (Fst=0.153) between bay-spawning populations. Between morphs, genetic differences ranged from a low (Fst=0.008-0.022) between deep- and shallow-spawning populations to high difference (Fst=0.125-0.143) between shallow- and bay-spawning populations. A higher proportion of molecular variance was seen among (3.9%) than within morphs (2.8%). The adaptive gene combinations behind the four morphs seem to have originated within the lake, although the lake could have been colonized from more than one source population.

Lifetime success and interactions of farm salmon invading a native population.

Farm Atlantic salmon escape and invade rivers throughout the North Atlantic annually, which has generated growing concern about their impacts on native salmon populations. A large-scale experiment was therefore undertaken in order to quantify the lifetime success and interactions of farm salmon invading a Norwegian river. Sexually mature farm and native salmon were genetically screened, radio tagged and released into the River Imsa where no other salmon had been allowed to ascend. The farm fishes were competitively and reproductively inferior, achieving less than one-third the breeding success of the native fishes. Moreover, this inferiority was sex biased, being more pronounced in farm males than females, resulting in the principal route of gene flow involving native males mating with farm females. There were also indications of selection against farm genotypes during early survival but not thereafter. However, evidence of resource competition and competitive displacement existed as the productivity of the native population was depressed by more than 30%. Ultimately, the lifetime reproductive success (adult to adult) of the farm fishes was 16% that of the native salmon. Our results indicate that such annual invasions have the potential for impacting on population productivity, disrupting local adaptations and reducing the genetic diversity of wild salmon populations.

A meta-analysis of fluctuating asymmetry in relation to heterozygosity.

Fluctuating asymmetry, the random departure from perfect bilateral symmetry, is a common measure of developmental instability that has been hypothesized to be inversely correlated with heterozygosity. Although this claim has been widely repeated, several studies have reported no such association. Therefore, we test the generality of this association, using meta-analysis, by converting test statistics for the relationship between heterozygosity (H) and fluctuating asymmetry (FA) into a common effect size, the Pearson's product-moment correlation coefficient. We have analysed a database containing 41 studies with a total of 118 individual samples. Overall we found an unweighted mean negative effect size; r=-0.09 (i. e. a negative correlation between H and FA). Significant heterogeneity in effect size was mainly caused by a difference between ectothermic and endothermic animals, and to a lesser extent by the use of different study designs (i.e. within-population vs. among-populations). Mean effect size for endothermic animals was positive and significantly different from the mean effect size for ectothermic animals. Only for within-population studies of ectothermic animals did we find a significantly negative effect size (r=-0.23 +/- 0.09). The distribution of effect sizes in relation to sample size provided little evidence for patterns typical of those produced by publication bias. Our analysis suggests, at best, only a weak association between H and FA, and heterozygosity seems to explain only a very small amount of the variation in developmental instability among individuals and populations (r2=0.01 for the total material).

Analysis of genetic structure and dispersal patterns in a population of sea beet.

A model of the migration pattern in a metapopulation of sea beet (Beta vulgaris L. ssp. maritima), based on the continuous distributions of seed and pollen movements, is fitted to gene frequency data at 12 isozyme and RFLP loci by maximum likelihood by using an approximation of the simultaneous equilibrium distribution of the gene frequencies generated by the underlying multivariate stochastic process of genetic drift in the population. Several alternative restrictions of the general model are fitted to the data, including the island model, a model of complete isolation, and a model in which the seed and pollen dispersal variances are equal. Several likelihood ratio tests between these alternatives are performed, and median bias in the estimated parameters is corrected by using parametric bootstrapping. To assess the fit of the selected model, the predicted covariances are compared with covariances computed from the data directly. The dependency of estimated parameters on the ratio between effective and absolute subpopulation sizes, which is treated as a known parameter in the analysis, is also examined. Finally, we note that the data also appear to contain some information about this ratio.

Stochastic Dispersal Processes in Plant Populations

A dispersal model for airborne pollen based on assumptions about wind directionality, gravity, and a wind threshold at which pollen is taken by the wind is developed, using a three dimensional diffusion approximation. The bivariate probability distribution of pollen receipt by flowers at the same height as the pollen source is derived. Gravity, vertical random movements, and vegetation density turn out to have similar effects on this distribution. Maximum likelihood methods for estimating the combined parameters from data with multiple point or continuous pollen sources, and one or more plant varieties, are developed. Using an example data set from the literature, it is shown that our model gives a better fit than more traditional descriptive dispersal models of the form e-ar b. We also show that estimates of important properties of the dispersal distribution, such as the variances, become considerably smaller using our model than for the more traditional models. Finally, we discuss potential extensions and evolutionary implications of these types of models. Copyright 1997 Academic Press

Developmental stability and environmental stress in Salmo salar (Atlantic salmon).

We have studied the developmental stability (measured as fluctuating asymmetry of five meristic characters) of three populations of Atlantic salmon Salmo salar (rivers Imsa, Lone and Ogna, western Norway). All three populations were both sampled in the wild, and hatched and reared in a common environment in a hatchery (with water from the river Imsa) from fertilization until smoltification. Both the Imsa and Lone hatchery populations have been sea-ranched in the Imsa for 10 years, whereas the Ogna populations is novel to the hatchery environment. Individual biochemical heterozygosity was scored at 50 loci, of which 11 were polymorphic. There was no correlation between biochemical heterozygosity and fluctuating asymmetry at the individual level, neither when tested within groups nor when tested between groups. There were no differences in fluctuating asymmetry between wild and hatchery Imsa and Lone fish, indicating that the hatchery environment did not disrupt early developmental homeostatic processes. However, the Ogna hatchery fish had significantly elevated levels of fluctuating asymmetry compared to the wild Ogna fish, indicating that the hatchery environment was hostile. The Ogna hatchery fish also had significantly higher fluctuating asymmetry than the Imsa hatchery and the Lone hatchery fish. Maladaptation to the hatchery environment is the most likely explanation for the increased asymmetry in river Ogna fish.

Inferring patterns of migration from gene frequencies under equilibrium conditions.

A new maximum likelihood method to simultaneously estimate the parameters of any migration pattern from gene frequencies in stochastic equilibrium is developed, based on a model of multivariate genetic drift in a subdivided population. Motivated by simulations of this process in the simplified case of two subpopulations, problems related to the nuisance parameter q, the equilibrium gene frequency, are eliminated by conditioning on the observed mean gene frequency. The covariance matrix of this conditional distribution is calculated by constructing an abstract process that mimics the behavior of the original process in the subspace of interest. The approximation holds as long as there is limited differentiation between subpopulations. The bias and variance of estimates of long-range and short-range migration in a finite stepping stone model are evaluated by fitting the model to simulated data with known values of the parameters. Possible ecological extensions of the model are discussed.

Optimal age at sexual maturity of sympatric and experimentally allopatric cutthroat trout and Dolly Varden charr.

Reproductive potentials of transplanted curthroat trout (Salmo clarki) and Dolly Varden charr (Salvelinus malma) and of their donor stocks were estimated from life history data. We found good agreement between observed and predicted age at maturity in all populations, and cannot reject the hypothesis that the fish matured at the age maximizing the overall lifetime reproductive potential ([Formula: see text]). Our estimates were insensitive to probable variations in female fecundity, adult mortalityrate and maximum body length. Small changes in either juvenile mortality-rate or individual growth-rate had marked effects on the estimations, as did changes in the Malthusian parameter (r). Three alternative mechanistic explanations of how age at maturity is determined could be rejected. We suggest that fish are able to adjust the maturity age non-genetically to changes in growth-rate, and that temporal variations in juvenile survival-rate allow coexistence of genotypes coding for different ages at maturity at the same growth-rate.