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.

Chemical composition and particle size influence the toxicity of nanoscale plastic debris and their co-occurring benzo(α)pyrene in the model aquatic organisms Daphnia magna and Danio rerio.

Little is known about how particle chemical composition and size might influence the toxicity of nanoscale plastic debris (NPD) and their co-occurring chemicals. Herein, we investigate the toxicity of 3 × 1010 particles/L polyethylene (PE, 50 nm), polypropylene (PP, 50 nm), polystyrene (PS, 200 and 600 nm), and polyvinyl chloride (PVC, 200 nm) NPD and their co-occurring benzo(a)pyrene (BaP) to Daphnia magna and Danio rerio. During the 21 days of exposure to PE 50 nm and PS 200 nm, the number of broods produced by D. magna decreased compared to other treatments. Exposure to BaP alone did not produce any effects on the reproduction of the daphnids, however, the mixture of BaP with PS (200 or 600 nm) or with PE (50 nm) reduced the number of broods. Exposure of D. rerio embryos to PE 50 nm, PS 200 nm, and PS 600 nm led to a delay in the hatching. The presence of PS 200 nm and PVC 200 nm eliminated the effects of BaP on the hatching rate of zebrafish. Our findings suggest that data generated for the toxicity of one type of NPD, e.g. PVC or PS may not be extrapolated to other types of NPD.

Pre-fertilization exposure of sperm to nano-sized plastic particles decreases offspring size and swimming performance in the European whitefish (Coregonus lavaretus).

Exposure of aquatic organisms to micro- and nano-sized plastic debris in their environment has become an alarming concern. Besides having a number of potentially harmful impacts for individual organisms, plastic particles can also influence the phenotype and performance of their offspring. We tested whether the sperm pre-fertilization exposure to nanoplastic particles could affect offspring survival, size, and swimming performance in the European whitefish Coregonus lavaretus. We exposed sperm of ten whitefish males to three concentrations (0, 100 and 10 000 pcs spermatozoa-1) of 50 nm carboxyl-coated polystyrene spheres, recorded sperm motility parameters using computer assisted sperm analysis (CASA) and then fertilized the eggs of five females in all possible male-female combinations. Finally, we studied embryonic mortality, hatching time, size, and post-hatching swimming performance of the offspring. We found that highest concentration of plastic particles decreased sperm motility and offspring hatching time. Furthermore, sperm exposure to highest concentration of plastics reduced offspring body mass and impaired their swimming ability. This suggests that sperm pre-fertilization exposure to plastic pollution may decrease male fertilization potential and have important transgenerational impacts for offspring phenotype and performance. Our findings indicate that nanoplastics pollution may have significant ecological and evolutionary consequences in aquatic ecosystems.

Global data set of long-term summertime vertical temperature profiles in 153 lakes.

Climate change and other anthropogenic stressors have led to long-term changes in the thermal structure, including surface temperatures, deepwater temperatures, and vertical thermal gradients, in many lakes around the world. Though many studies highlight warming of surface water temperatures in lakes worldwide, less is known about long-term trends in full vertical thermal structure and deepwater temperatures, which have been changing less consistently in both direction and magnitude. Here, we present a globally-expansive data set of summertime in-situ vertical temperature profiles from 153 lakes, with one time series beginning as early as 1894. We also compiled lake geographic, morphometric, and water quality variables that can influence vertical thermal structure through a variety of potential mechanisms in these lakes. These long-term time series of vertical temperature profiles and corresponding lake characteristics serve as valuable data to help understand changes and drivers of lake thermal structure in a time of rapid global and ecological change.

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.

Deeper waters are changing less consistently than surface waters in a global analysis of 102 lakes.

Globally, lake surface water temperatures have warmed rapidly relative to air temperatures, but changes in deepwater temperatures and vertical thermal structure are still largely unknown. We have compiled the most comprehensive data set to date of long-term (1970-2009) summertime vertical temperature profiles in lakes across the world to examine trends and drivers of whole-lake vertical thermal structure. We found significant increases in surface water temperatures across lakes at an average rate of + 0.37 °C decade-1, comparable to changes reported previously for other lakes, and similarly consistent trends of increasing water column stability (+ 0.08 kg m-3 decade-1). In contrast, however, deepwater temperature trends showed little change on average (+ 0.06 °C decade-1), but had high variability across lakes, with trends in individual lakes ranging from - 0.68 °C decade-1 to + 0.65 °C decade-1. The variability in deepwater temperature trends was not explained by trends in either surface water temperatures or thermal stability within lakes, and only 8.4% was explained by lake thermal region or local lake characteristics in a random forest analysis. These findings suggest that external drivers beyond our tested lake characteristics are important in explaining long-term trends in thermal structure, such as local to regional climate patterns or additional external anthropogenic influences.

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.

Do Metabolic Traits, Vulnerability to Angling, or Capture Method Explain Boldness Variation in Eurasian Perch?

The pace-of-life syndrome (POLS) concept predicts that individuals with high baseline metabolic rates demonstrate high boldness, aggressiveness, and activity, especially in food acquisition, with associated relatively greater energy requirements. In fishes, these behaviors may increase individual vulnerability to angling. To test the predictions of the POLS concept, we quantified individual standard metabolic rate (SMR) and boldness in both wild-caught and hatchery-reared Eurasian perch (Perca fluviatilis). We found both SMR and boldness to be repeatable traits but detected no correlation between them. Individual vulnerability to angling was assessed in the hatchery-reared perch, but we found no difference in boldness or SMR between vulnerable and nonvulnerable perch. Wild-caught perch were ice fished using either natural or artificial bait, and we observed no differences in boldness or SMR with respect to bait type or capture order. Our findings do not support the predictions of the POLS concept and, consistent with earlier studies in perch, suggest that angling may not drive selection against boldness in this species.

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.

Latitudinal variation in sexual dimorphism in life-history traits of a freshwater fish.

Sexual dimorphism is common across the animal kingdom, but the contribution of environmental factors shaping differences between the sexes remains controversial. In ectotherms, life-history traits are known to correlate with latitude, but sex-specific responses are not well understood. We analyzed life-history trait variation between the sexes of European perch (Perca fluviatilis L.), a common freshwater fish displaying larger female size, by employing a wide latitudinal gradient. We expected to find sex-dependent latitudinal variation in life-history variables: length at age, length increment, and size at maturity, with females showing consistently higher values than males at all latitudes. We further anticipated that this gender difference would progressively decrease with the increasingly harsh environmental conditions toward higher latitude. We hypothesized that growth and length increment would decrease and size/age at maturity would increase at higher latitudes. Our results confirmed female-biased sexual size dimorphism at all latitudes and the magnitude of sexual dimorphism diminished with increase in latitude. Growth of both sexes decreased with increase in latitude, and the female latitudinal clines were steeper than those of males. Hence, we challenge two predominant ecological rules (Rensch's and Bergmann's rules) that describe common large-scale patterns of body size variation. Our data demonstrate that these two rules are not universally applicable in ectotherms or female-biased species. Our study highlights the importance of sex-specific differences in life-history traits along a latitudinal gradient, with evident implications for a wide range of studies from individual to ecosystems level.

Phenotypic plasticity in growth and fecundity induced by strong population fluctuations affects reproductive traits of female fish.

Fish are known for their high phenotypic plasticity in life-history traits in relation to environmental variability, and this is particularly pronounced among salmonids in the Northern Hemisphere. Resource limitation leads to trade-offs in phenotypic plasticity between life-history traits related to the reproduction, growth, and survival of individual fish, which have consequences for the age and size distributions of populations, as well as their dynamics and productivity. We studied the effect of plasticity in growth and fecundity of vendace females on their reproductive traits using a series of long-term incubation experiments. The wild parental fish originated from four separate populations with markedly different densities, and hence naturally induced differences in their growth and fecundity. The energy allocation to somatic tissues and eggs prior to spawning served as a proxy for total resource availability to individual females, and its effects on offspring survival and growth were analyzed. Vendace females allocated a rather constant proportion of available energy to eggs (per body mass) despite different growth patterns depending on the total resources in the different lakes; investment into eggs thus dictated the share remaining for growth. The energy allocation to eggs per mass was higher in young than in old spawners and the egg size and the relative fecundity differed between them: Young females produced more and smaller eggs and larvae than old spawners. In contrast to earlier observations of salmonids, a shortage of maternal food resources did not increase offspring size and survival. Vendace females in sparse populations with ample resources and high growth produced larger eggs and larvae. Vendace accommodate strong population fluctuations by their high plasticity in growth and fecundity, which affect their offspring size and consequently their recruitment and productivity, and account for their persistence and resilience in the face of high fishing mortality.

Male Investments in High Quality Sperm Improve Fertilization Success, but May Have Negative Impact on Offspring Fitness in Whitefish.

Many ejaculate traits show remarkable variation in relation to male social status. Males in disfavoured (subordinate) mating positions often invest heavily on sperm motility but may have less available resources on traits (e.g., secondary sexual ornaments) that improve the probability of gaining matings. Although higher investments in sperm motility can increase the relative fertilization success of subordinate males, it is unclear whether status-dependent differences in sperm traits could have any consequences for offspring fitness. We tested this possibility in whitefish (Coregonus lavaretus L.) by experimentally fertilizing the eggs of 24 females with the sperm of either highly-ornamented (large breeding tubercles, dominant) or less-ornamented (small tubercles, subordinate) males (split-clutch breeding design). In comparison to highly-ornamented individuals, less-ornamented males had higher sperm motility, which fertilized the eggs more efficiently, but produced embryos with impaired hatching success. Also offspring size and body condition were lower among less-ornamented males. Furthermore, sperm motility was positively associated with the fertilization success and offspring size, but only in highly-ornamented males. Together our results indicate that male investments on highly motile (fertile) sperm is not necessarily advantageous during later offspring ontogeny and that male status-dependent differences in sperm phenotype may have important effects on offspring fitness in different life-history stages.

Both male and female sexual ornaments reflect offspring performance in a fish.

Secondary sexual characters are often expressed in both sexes (mutual ornamentation), but are less often studied simultaneously. We studied the adaptive signaling function of male and female ornamentation in a mutually ornamented fish, the whitefish Coregonus lavaretus. In an experimental design in which nongenetic environmental effects were minimized, we found that highly ornamented females, males, and their parental combinations had offspring with better swimming performance and predator-avoidance ability than less ornamented individuals or combinations. Furthermore, highly ornamented females had larger offspring that also had higher yolk volume than less ornamented individuals. Offspring swimming performance was not dependent on offspring size and was only weakly affected by yolk volume, which suggest that swimming performance and measured morphological traits are independent fitness measures. In conclusion, mutual ornamentation of whitefish may signal the quality of individuals in both sexes, which may indicate ongoing directional selection for these ornamental traits. However, offspring fitness traits were also dependent on parental combination, which suggests that genetic compatibility effects may weaken the directional selection and the indicator value of the ornamentation.

Consistency of standard metabolic rate in relation to life history strategy of juvenile Atlantic salmon Salmo salar.

Temporal consistency of relative standard metabolic rate (rSMR) of individual Atlantic salmon Salmo salar juveniles in three stocks was investigated. The stocks differed in their geographical origin and natal habitat (fresh water, brackish water, seawater). Oxygen consumption measurements of hatchery-reared fish hatched in February 2005 were conducted three times: in early spring 2006, in autumn 2006, and in late spring 2007. The results partly disagreed and partly agreed with earlier studies, in which temporal consistency of SMR in juvenile salmonids have been addressed. In the first period from early spring to autumn, no correlation between rSMR statuses of individuals was found whereas in the second period from autumn to late spring, fish were mainly observed to maintain their relative SMR level indicating a significant repeatability of individual rSMR status over the latter period. Furthermore, a relationship between rSMR status and life history strategy was found: post-smolts and smolts had higher SMR than non-smolts.