Selection among critically endangered landlocked salmon (Salmo salar m. sebago) families in survival and growth traits across early life stages and in different environments.

Endangered wild fish populations are commonly supported by hatchery propagation. However, hatchery-reared fish experience very different selective pressures compared to their wild counterparts, potentially causing genotype-by-environment interactions (G × E) in essential fitness traits. We experimentally studied early selection in a critically endangered landlocked Atlantic salmon population, first from fertilization to the swim-up stage in a common hatchery setting, and thereafter until the age of 5 months in two contrasting rearing environments. Swim-up progeny were moved either to standard indoor hatchery tanks involving conventional husbandry or to seminatural outdoor channels providing only natural food. After the first summer, sampled survivors were assigned to their families by genotyping. Early survival until the swim-up stage was mostly determined by maternal effects, but also involved significant variation due to sires and full-sib families (potential genetic effects). High on-growing survival in hatchery tanks (88.7%) maintained a more even distribution among families (relative share 1.5%-4.2%) than the seminatural environment (0.0%-5.4%). This heterogeneity was mostly maternal, whereas no independent paternal effect occurred. Heritability estimates were high for body size traits in both environments (0.62-0.69). Genetic correlations between the environments were significantly positive for body size traits (0.67-0.69), and high body condition in hatchery was also genetically linked to rapid growth in the seminatural environment (0.54). Additive and phenotypic growth variation increased in the seminatural environment, but scaling effects probably played a less significant role for G × E, compared to re-ranking of genotypes. Our results suggest that not only maternal effects, but also genetic effects, direct selection according to the environmental conditions experienced. Consistently high genetic variation in growth implies that, despite its low overall genetic diversity and long history in captive rearing (>50 years), this landlocked Atlantic salmon population still possesses adaptive potential for response to change from hatchery rearing back to more natural conditions.

Exposure of gametes to aged nano-sized plastic particles during fertilization can influence early larval development in the European whitefish (Coregonus lavaretus).

Plastic pollution has been a growing environmental concern for decades, increasingly affecting both marine and freshwater ecosystems worldwide. Nano-sized plastic particles (NPs) potentially have various toxicological impacts on aquatic organisms and the ecosystem; however, less is known about their possible adverse effects on the reproductive biology and offspring traits of fishes. The present study investigated whether an acute exposure of gametes to aged NPs during fertilization affects offspring early mortality, hatching time, body size at hatching or swimming performance of larvae in a common freshwater fish, the European whitefish (Coregonus lavaretus). Using a replicated full-factorial breeding design, we fertilized the eggs of seven females with the milt of seven males both under exposure medium containing aged 270 nm polystyrene NPs and under control medium. In comparison with the control group, exposure of gametes to NPs increased larval body length slightly but significantly, whereas the embryo mortality, hatching time, and larval swimming performance were not affected. Maternal identity affected significantly all the studied offspring traits while paternal identity only affected the offspring length. Our results suggest that the studied acute exposure of gametes to aged NPs might have interfered normal embryonic development by affecting larval size, but this did not seemingly compromise offspring performance.

The effects of environmental enrichment on hatchery-performance, smolt migration and capture rates in landlocked Atlantic salmon.

Enrichment of rearing environment with natural elements has been suggested to improve the welfare and post-release survival of cultured fish. We studied the combined effects of shelter structures, periodical water flow and water level changes on pre- and post-release performance of critically endangered landlocked Atlantic salmon (Salmo salar m. sebago). Relative to standard (plain) rearing tanks, provision of enrichment improved fish condition factor and survival during the first year of rearing when most mortality was attributable to parasitic and bacterial infections. The consequent higher density in enriched tanks probably induced greater growth variation and more dorsal fin damages than found in fish of standard tanks. Possibly this was partly due to the applied changes in water level. Experimentally determined smolt migration tendency at age 3 did not differ, on average, between the rearing groups, but enriched-reared fish showed clearly less variation in total movement activity than standard-reared fish. Experimental angling in earthen ponds did not suggest divergent vulnerability between the differentially reared fish at age 3, but decreased condition during the preceding growth season increased vulnerability to fishing. Based on long-term post-stocking tag returns in large-lake fisheries, fish length at release but not rearing method affected the capture rates of fish released at age 2. When released at age 3 the fish grown in enriched environment had a higher risk to be captured with stationary gears and earlier by hook and line gears compared to standard-reared conspecifics. Earlier time of maximal smolt migration activity was associated with an increased risk of being captured. We suggest that environmental enrichment may modulate growth- and behavior-related qualities that indirectly increased the vulnerability to fishing in natural conditions but not in experimental setting. The favorable effects of enrichment on early survival encourages adopting enriched rearing practices in supportive breeding of landlocked salmon.

Ultra-acute exposure to cadmium does not impair whitefish sperm motility.

Cadmium (Cd) exposure can impair the traits of aquatic animals associated with reproduction. In natural lakes Cd is typically detected at concentrations below 0.001 mg l-1 . The authors investigated the impact of ultra-acute Cd exposure on sperm motility in European whitefish (Coregonus lavaretus). They activated sperm with water containing various nominal concentrations of Cd and recorded sperm motility parameters. Only the highest Cd concentration (500 mg l-1 ) was associated with decreased sperm swimming velocity and increases in both the percentage of static cells and curvature of the sperm swimming trajectory. The results indicate that environmentally realistic concentrations of Cd during the sperm motility activation are not critically harmful to male C. lavaretus fertilization potential.

Do whitefish (Coregonus lavaretus) larvae show adaptive variation in the avoidance of microplastic ingestion?

The presence of microplastics in aquatic ecosystems has recently received increased attention. Small plastic particles may resemble natural food items of larval fish and other aquatic organisms, and create strong selective pressures on the feeding traits in exposed populations. Here, we examined if larval ingestion of 90 µm polystyrene microspheres, in the presence of zooplankton (Artemia nauplii, mean length = 433 µm), shows adaptive variation in the European whitefish (Coregonus lavaretus). A full-factorial experimental breeding design allowed us to estimate the relative contributions of male (sire) and female (dam) parents and full-sib family variance in early feeding traits, and also genetic (co)variation between these traits. We also monitored the magnitude of intake and elimination of microplastics from the alimentary tracts of the larvae. In general, larval whitefish ingested small numbers of microplastics (mean = 1.8, range = 0-26 particles per larva), but ingestion was marginally affected by the dam, and more strongly by the full-sib family variation. Microsphere ingestion showed no statistically significant additive genetic variation, and thus, no heritability. Moreover, microsphere ingestion rate covaried positively with the ingestion of Artemia, further suggesting that larvae cannot adaptively avoid microsphere ingestion. Together with the detected strong genetic correlation between food intake and microplastic intake, the results suggest that larval fish do not readily possess additive genetic variation that would help them to adapt to the increasing pollution by microplastics. The conflict between feeding on natural food and avoiding microplastics deserves further attention.

Offspring phenotype is shaped by the nonsperm fraction of semen.

In a large majority of animal species, the only contribution of males to the next generation has been assumed to be their genes (sperm). However, along with sperm, seminal plasma contains a wide array of extracellular factors that have many important functions in reproduction. Yet, the potential intergenerational effects of these factors are virtually unknown. We investigated these effects in European whitefish (Coregonus lavaretus) by experimentally manipulating the presence and identity of seminal plasma and by fertilizing the eggs of multiple females with the manipulated and unmanipulated semen of several males in a full-factorial breeding design. The presence of both own seminal plasma and foreign seminal plasma inhibited sperm motility, and the removal of own seminal plasma decreased embryo survival. Embryos hatched significantly earlier after both semen manipulations than in control fertilizations; foreign seminal plasma also increased offspring aerobic swimming performance. Given that our experimental design allowed us to control potentially confounding sperm-mediated (sire) effects and maternal effects, our results indicate that seminal plasma may have direct intergenerational consequences for offspring phenotype and performance. This novel source of offspring phenotypic variance may provide new insights into the evolution of polyandry and mechanisms that maintain heritable variation in fitness and associated female mating preferences.

Experimental crossbreeding reveals strain-specific variation in mortality, growth and personality in the brown trout (Salmo trutta).

Although hybridization between populations with low genetic diversity may induce heterosis, it can also lead to reduced fitness of hybrid offspring through outbreeding depression and loss of local adaptations. Using a half-sib mating design, we studied on brown trout (Salmo trutta) how hybridization of migratory hatchery-strain females with males from various strains would affect early mortality, growth and personality in F1 offspring. No differences in mortality or alevin body length were found between the crossing groups by the end of the yolk-sac stage. At later developmental stages, higher mortality and slower growth in one of the geographically distant hybrid groups indicated potential outbreeding depression. The personality component indicating boldness and exploration tendency showed fairly low genetic variation and no phenotypic differences among the crossing groups while the personality component related to freezing behavior indicated stronger freezing responses in the purebred and local cross strain when compared to the two other strains. However, the purebred hatchery strain possessed stronger additive genetic tendency for boldness and explorative behavior, and weaker genetic tendency for freezing behavior, when compared to the wild × hatchery hybrid group. Our results add to the cumulating evidence of risks related to the stocking of fish strains from non-native origins.

Sperm pre-fertilization thermal environment shapes offspring phenotype and performance.

The sperm pre-fertilization environment has recently been suggested to mediate remarkable transgenerational consequences for offspring phenotype (transgenerational plasticity, TGB), but the adaptive significance of the process has remained unclear. Here, we studied the transgenerational effects of sperm pre-fertilization thermal environment in a cold-adapted salmonid, the European whitefish (Coregonus lavaretus). We used a full-factorial breeding design where the eggs of five females were fertilized with the milt of 10 males that had been pre-incubated at two different temperatures (3.5°C and 6.5°C) for 15 h prior to fertilization. Thermal manipulation did not affect sperm motility, cell size, fertilization success or embryo mortality. However, offspring that were fertilized with 6.5°C-exposed milt were smaller and had poorer swimming performance than their full-siblings that had been fertilized with the 3.5°C-exposed milt. Furthermore, the effect of milt treatment on embryo mortality varied among different females (treatment×female interaction) and male-female combinations (treatment×female×male interaction). Together, these results indicate that sperm pre-fertilization thermal environment shapes offspring phenotype and post-hatching performance and modifies both the magnitude of female (dam) effects and the compatibility of the gametes. Generally, our results suggest that short-term changes in sperm thermal conditions may have negative impact for offspring fitness. Thus, sperm thermal environment may have an important role in determining the adaptation potential of organisms to climate change. Detailed mechanism(s) behind our findings require further attention.

Thermal sensitivity of growth indicates heritable variation in 1-year-old rainbow trout (Oncorhynchus mykiss).

BACKGROUND: Rainbow trout is an important aquaculture species, which has a worldwide distribution across various production environments. The diverse locations of trout farms involve remarkable variation in environmental factors such as water temperature, which is of major importance for the performance of fish. Thus, robust fish that could thrive under different and suboptimal thermal conditions is a desirable goal for trout breeding. Using a split-family experimental design (40 full-/half-sib groups) for a rainbow trout population derived from the Finnish national breeding program, we studied how two different rearing temperatures (14 and 20 °C) affect feed intake, growth rate and feed conversion ratio in 1-year-old fish. Furthermore, we quantified the additive genetic (co-)variation for daily growth coefficient (DGC) and its thermal sensitivity (TS), defined as the slope of the growth reaction norm between the two temperatures. RESULTS: The fish showed consistently lower feed intake, faster growth and better feed conversion ratio at the lower temperature. Heritability of TS of DGC was moderate ([Formula: see text]). The co-heritability parameter derived from selection index theory, which describes the heritable variance of TS, was negative when the intercept was placed at the lower temperature (-0.28). This resulted in moderate accuracy of selection. At the higher temperature, co-heritability of TS was positive (0.20). The genetic correlation between DGC and its TS was strongly negative (-0.64) when the intercept was at the lower temperature and positive (0.38) but not significantly different from zero at the higher temperature. CONCLUSIONS: The considerable amount of genetic variation in TS of growth indicates a potential for selection response and thus for targeted genetic improvement in TS. The negative genetic correlation between DGC and its TS suggests that selection for high growth rate at the lower temperature will result in more temperature-sensitive fish. Instead, the correlated response of TS is less pronounced if the selection for a higher DGC occurred at the higher temperature. It seems possible to control the correlated genetic change of TS while selecting for fast growth across environments, especially if measurements from both environments are available and breeding values for reaction norm slope are directly included in the selection index.

Genetic (co)variance of rainbow trout (Oncorhynchus mykiss) body weight and its uniformity across production environments.

BACKGROUND: When rainbow trout from a single breeding program are introduced into various production environments, genotype-by-environment (GxE) interaction may occur. Although growth and its uniformity are two of the most important traits for trout producers worldwide, GxE interaction on uniformity of growth has not been studied. Our objectives were to quantify the genetic variance in body weight (BW) and its uniformity and the genetic correlation (rg) between these traits, and to investigate the degree of GxE interaction on uniformity of BW in breeding (BE) and production (PE) environments using double hierarchical generalized linear models. Log-transformed data were also used to investigate whether the genetic variance in uniformity of BW, GxE interaction on uniformity of BW, and rg between BW and its uniformity were influenced by a scale effect. RESULTS: Although heritability estimates for uniformity of BW were low and of similar magnitude in BE (0.014) and PE (0.012), the corresponding coefficients of genetic variation reached 19 and 21%, which indicated a high potential for response to selection. The genetic re-ranking for uniformity of BW (rg = 0.56) between BE and PE was moderate but greater after log-transformation, as expressed by the low rg (-0.08) between uniformity in BE and PE, which indicated independent genetic rankings for uniformity in the two environments when the scale effect was accounted for. The rg between BW and its uniformity were 0.30 for BE and 0.79 for PE but with log-transformed BW, these values switched to -0.83 and -0.62, respectively. CONCLUSIONS: Genetic variance exists for uniformity of BW in both environments but its low heritability implies that a large number of relatives are needed to reach even moderate accuracy of selection. GxE interaction on uniformity is present for both environments and sib-testing in PE is recommended when the aim is to improve uniformity across environments. Positive and negative rg between BW and its uniformity estimated with original and log-transformed BW data, respectively, indicate that increased BW is genetically associated with increased variance in BW but with a decrease in the coefficient of variation. Thus, the scale effect substantially influences the genetic parameters of uniformity, especially the sign and magnitude of its rg.

Female ornaments revisited - are they correlated with offspring quality?

The evolution and signalling content of female ornamentation has remained an enduring challenge to evolutionary biologists, despite the fact that secondary sexual characters are widespread in females. While females usually invest significant amounts of their resources, including carotenoids, in offspring, all the resources allocated to elaborate ornamentation reduce resources available for other purposes. This may in turn constrain female fitness leading to dishonest female signalling. We review the literature for empirical studies on mutually ornamented species with conventional sex roles, by focusing on the association between female ornaments and quality of their offspring. We found 43 papers where 33 (77%) are bird-studies, nine (21%) are on fishes, and one (2%) is a lizard-study. Nine of these report negative, 14 non-existing, and 20 positive associations between female ornament and offspring quality. Eighteen of the bird studies (55%) show a positive association between the two traits investigated, whereas five (15%) of the studies report a negative association. The number of fish studies, although few, is skewed in the opposite direction with two (22%) and four (44%) studies supporting positive and negative association, respectively. A minority of studies on carotenoids-based ornaments reports a positive association (4 of 18 studies, or 22%) between the traits, which is low compared to studies on non-carotenoids-based ornaments (16 of 25 studies, or 64%). The above-mentioned relative large number of studies with negative association, especially common in studies on fishes and in carotenoids-based-ornaments, challenges the generality of the direct selection hypothesis to account for female fineries. This is important because this hypothesis seems to have strong support in recent literature on the topic. In the present paper, we also propose possible explanations for the observed differences between taxa and suggest directions and ideas for future research on the evolution of female ornamentation.

Genetics of microenvironmental sensitivity of body weight in rainbow trout (Oncorhynchus mykiss) selected for improved growth.

Microenvironmental sensitivity of a genotype refers to the ability to buffer against non-specific environmental factors, and it can be quantified by the amount of residual variation in a trait expressed by the genotype's offspring within a (macro)environment. Due to the high degree of polymorphism in behavioral, growth and life-history traits, both farmed and wild salmonids are highly susceptible to microenvironmental variation, yet the heritable basis of this characteristic remains unknown. We estimated the genetic (co)variance of body weight and its residual variation in 2-year-old rainbow trout (Oncorhynchus mykiss) using a multigenerational data of 45,900 individuals from the Finnish national breeding programme. We also tested whether or not microenvironmental sensitivity has been changed as a correlated genetic response when genetic improvement for growth has been practiced over five generations. The animal model analysis revealed the presence of genetic heterogeneity both in body weight and its residual variation. Heritability of residual variation was remarkably lower (0.02) than that for body weight (0.35). However, genetic coefficient of variation was notable in both body weight (14%) and its residual variation (37%), suggesting a substantial potential for selection responses in both traits. Furthermore, a significant negative genetic correlation (-0.16) was found between body weight and its residual variation, i.e., rapidly growing genotypes are also more tolerant to perturbations in microenvironment. The genetic trends showed that fish growth was successfully increased by selective breeding (an average of 6% per generation), whereas no genetic change occurred in residual variation during the same period. The results imply that genetic improvement for body weight does not cause a concomitant increase in microenvironmental sensitivity. For commercial production, however, there may be high potential to simultaneously improve weight gain and increase its uniformity if both criteria are included in a selection index.

Genetic and potential non-genetic benefits increase offspring fitness of polyandrous females in non-resource based mating system.

BACKGROUND: The adaptive significance of female polyandry is currently under considerable debate. In non-resource based mating systems, indirect, i.e. genetic benefits have been proposed to be responsible for the fitness gain from polyandry. We studied the benefits of polyandry in the Arctic charr (Salvelinus alpinus) using an experimental design in which the material investments by the sires and maternal environmental effects were controlled. RESULTS: Embryonic mortality showed a strong paternal genetic component, and it was lower in polyandrously fertilized offspring (sperm competition of two males) than in monandrous fertilizations. We also found that high sperm velocity was associated with low offspring mortality, but not with the size of the offspring or their yolk volume. Although no male effect was found on the size of the offspring yolk reserves, yolk volume was higher in offspring from polyandrous matings than offspring of the either of the two males when mated monandrously. CONCLUSIONS: In support of the "good sperm hypothesis, we found that sperm velocity was positively associated with offspring fitness. In addition, our results suggest that polyandrous females gain genetic advantage (higher offspring survival) from this behavior, but that some benefits of polyandry (larger yolk volume) may not be explained solely by the additive genetic effects. This suggests that sperm competition environment may intensify the selection on genetically superior sperm which in turn may produce offspring that have superior yolk reserves. However, as high sperm velocity was not associated with larger yolk volume, it is possible that also some other non-genetic effects may contribute to offspring fitness. The potential role of polyandrous mating in inbreeding avoidance is discussed.