Contemporary divergence in early life history in grayling (Thymallus thymallus)

<p>Abstract</p> <p>Background</p> <p>Following colonization of new habitats and subsequent selection, adaptation to environmental conditions might be expected to be rapid. In a mountain lake in Norway, Lesjaskogsvatnet, more than 20 distinct spawning demes of grayling have been established since the lake was colonized, some 20-25 generations ago. The demes spawn in tributaries consistently exhibiting either colder or warmer temperature conditions during spawning in spring and subsequent early development during early summer. In order to explore the degree of temperature-related divergence in early development, a multi-temperature common-garden experiment was performed on embryos from four different demes experiencing different spring temperatures.</p> <p>Results</p> <p>Early developmental characters were measured to test if individuals from the four demes respond differently to the treatment temperatures. There was clear evidence of among-deme differences (genotype - environment interactions) in larval growth and yolk-to-body-size conversion efficiency. Under the cold treatment regime, larval growth rates were highest for individuals belonging to cold streams. Individuals from warm streams had the highest yolk-consumption rate under cold conditions. As a consequence, yolk-to-body-mass conversion efficiency was highest for cold-deme individuals under cold conditions. As we observed response parallelism between individuals from demes belonging to similar thermal groups for these traits, some of the differentiation seems likely to result from local adaptation</p> <p>Conclusion</p> <p>The observed differences in length at age during early larval development most likely have a genetic component, even though both directional and random processes are likely to have influenced evolutionary change in the demes under study.</p

The Chromosome-Level Genome Assembly of European Grayling Reveals Aspects of a Unique Genome Evolution Process Within Salmonids

Salmonids represent an intriguing taxonomical group for investigating genome evolution in vertebrates due to their relatively recent last common whole genome duplication event, which occurred between 80 and 100 million years ago. Here, we report on the chromosome-level genome assembly of European grayling (Thymallus thymallus), which represents one of the earliest diverged salmonid subfamilies. To achieve this, we first generated relatively long genomic scaffolds by using a previously published draft genome assembly along with long-read sequencing data and a linkage map. We then merged those scaffolds by applying synteny evidence from the Atlantic salmon (Salmo salar) genome. Comparisons of the European grayling genome assembly to the genomes of Atlantic salmon and Northern pike (Esox lucius), the latter used as a nonduplicated outgroup, detailed aspects of the characteristic chromosome evolution process that has taken place in European grayling. While Atlantic salmon and other salmonid genomes are portrayed by the typical occurrence of numerous chromosomal fusions, European grayling chromosomes were confirmed to be fusion-free and were characterized by a relatively large proportion of paracentric and pericentric inversions. We further reported on transposable elements specific to either the European grayling or Atlantic salmon genome, on the male-specific sdY gene in the European grayling chromosome 11A, and on regions under residual tetrasomy in the homeologous European grayling chromosome pairs 9A-9B and 25A-25B. The same chromosome pairs have been observed under residual tetrasomy in Atlantic salmon and in other salmonids, suggesting that this feature has been conserved since the subfamily split.Peer reviewe

Contemporary temperature-driven divergence in a Nordic freshwater fish under conditions commonly thought to hinder adaptation

BACKGROUND: Evaluating the limits of adaptation to temperature is important given the IPCC-predicted rise in global temperatures. The rate and scope of evolutionary adaptation can be limited by low genetic diversity, gene flow, and costs associated with adaptive change. Freshwater organisms are physically confined to lakes and rivers, and must therefore deal directly with climate variation and change. In this study, we take advantage of a system characterised by low genetic variation, small population size, gene flow and between-trait trade-offs to study how such conditions affect the ability of a freshwater fish to adapt to climate change. We test for genetically-based differences in developmental traits indicating local adaptation, by conducting a common-garden experiment using embryos and larvae from replicate pairs of sympatric grayling demes that spawn and develop in natural cold and warm water, respectively. These demes have common ancestors from a colonization event 22 generations ago. Consequently, we explore if diversification may occur under severely constraining conditions. RESULTS: We found evidence for divergence in ontogenetic rates. The divergence pattern followed adaptation predictions as cold-deme individuals displayed higher growth rates and yolk conversion efficiency than warm-deme individuals at the same temperature. The cold-deme embryos had a higher rate of muscle mass development. Most of the growth- and development differences occurred prior to hatch. The divergence was probably not caused by genetic drift as there was a strong degree of parallelism in the divergence pattern and because phenotypic differentiation (Q(ST)) was larger than estimated genetic drift levels (microsatellite F(ST)) between demes from different temperature groups. We also document that these particular grayling populations cannot develop successfully at temperatures above 12°C, whereas other European populations can, and that increasing the muscle mass development rate comes at the cost of some skeletal trait development rates. CONCLUSIONS: This study shows that genetically based phenotypic divergence can prevail even under conditions of low genetic variation and ongoing gene flow. Furthermore, population-specific maximum development temperatures along with musculoskeletal developmental trade-offs may constrain adaptation

Life history and demographic determinants of effective/census size ratios as exemplified by brown trout (Salmo trutta)

A number of demographic factors, many of which related to human-driven encroachments, are predicted to decrease the effective population size (Ne) relative to the census population size (N), but these have been little investigated. Yet, it is necessary to know which factors most strongly impact Ne, and how to mitigate these effects through sound management actions. In this study, we use parentage analysis of a stream-living brown trout (Salmo trutta) population to quantify the effect of between-individual variance in reproductive success on the effective number of breeders (Nb) relative to the census number of breeders (Ni). Comprehensive estimates of the Nb/N ratio were reduced to 0.16–0.28, almost entirely due to larger than binomial variance in family size. We used computer simulations, based on empirical estimates of age-specific survival and fecundity rates, to assess the effect of repeat spawning (iteroparity) on Ne and found that the variance in lifetime reproductive success was substantially higher for repeat spawners. Random family-specific survival, on the other hand, acts to buffer these effects. We discuss the implications of these findings for the management of small populations, where maintaining high and stable levels of Ne is crucial to extenuate inbreeding and protect genetic variability.publishedVersio

When phenology matters: age–size truncation alters population response to trophic mismatch

Climate-induced shifts in the timing of life-history events are a worldwide phenomenon, and these shifts can de-synchronize species interactions such as predator–prey relationships. In order to understand the ecological implications of altered seasonality, we need to consider how shifts in phenology interact with other agents of environmental change such as exploitation and disease spread, which commonly act to erode the demographic structure of wild populations. Using long-term observational data on the phenology and dynamics of a model predator–prey system (fish and zooplankton in Windermere, UK), we show that age–size truncation of the predator population alters the consequences of phenological mismatch for offspring survival and population abundance. Specifically, age–size truncation reduces intraspecific density regulation due to competition and cannibalism, and thereby amplifies the population sensitivity to climate-induced predator–prey asynchrony, which increases variability in predator abundance. High population variability poses major ecological and economic challenges as it can diminish sustainable harvest rates and increase the risk of population collapse. Our results stress the importance of maintaining within-population age–size diversity in order to buffer populations against phenological asynchrony, and highlight the need to consider interactive effects of environmental impacts if we are to understand and project complex ecological outcomes

Spatio-temporal patterns in pelvic reduction in threespine stickleback (Gasterosteus aculeatus L.) in Lake Storvatnet

Questions: The pelvic girdle with associated spines is an integrated anti-predator defence apparatus, and is assumed to protect against piscivores in the threespine stickleback. On the other hand, it might be costly to produce the pelvic apparatus in ion-poor and mineralchallenging freshwater. Hypothesis: Stickleback with a reduced pelvic apparatus should use more shelter and be more nocturnal, avoiding predation risk. In contrast, stickleback with a well-developed pelvic apparatus should have reduced mortality during ontogeny in encounters with piscivores and thus have a longer expected lifespan. Given these two life-history strategies, we expect assortative mating as a result of divergent selection. Organism: Marine and freshwater threespine stickleback (Gasterosteus aculeatus L.). Places and times: Two representative ancestral marine populations and 36 freshwater populations in northwestern Norway (Lake Storvatnet, the main focus of the study, and three lakes downstream of it). Material was collected from 2006 to 2009. Analytical methods: We categorized nominal pelvic apparatus development (CPS morphs) in all fish, and measured metrics associated with these categories in a subsample. We also studied temporal, spatial, and habitat variation in the distribution of pelvic morphs in Lake Storvatnet. In this population, and downstream populations, we contrasted the detailed pelvic morphology with the measured genetic diversity (microsatellites), also estimating gene flow. In Lake Storvatnet, we tested for genetic divergence and signs of potential build-up of reproductive isolation via assortative mating among the observed nominal categories of pelvic reduction (CPS). Results: Pelvic reduction was seen only in Lake Storvatnet, where more than 50% of fish had a reduced pelvis. The distribution of pelvic morphs was stable over time and did not differ between habitats. The proportion of fish with pelvic reduction decreased with age. Freshwater stickleback tended to have a smaller pelvis than marine fish. The Lake Storvatnet stickleback were genetically differentiated from the downstream Lake Gjerhaugsvatn population, and both of these were different from the marine populations, with little gene flow among populations. No apparent genetic structure was found between CPS morphs within Lake Storvatnet. However, genetic factorial correspondence axes were significantly correlated with pelvic principal component axes in Lake Storvatnet, suggesting some phenotype × genetic association. Conclusion: The weak association between phenotypes and genetic structure observed in this study may reflect the build-up of early steps of reproductive isolation. Given time, such mechanisms may lead to the evolution of assortative mating, which may drive adaptive pelvic morphs (niche peaks), further resulting in genetically divergent populations and pelvic morphs

Thermo-fluid dynamics modelling of steam electrolysis in fully-assembled tubular high-temperature proton-conducting cells

Electrolysis based on renewable energies offers a promising carbon-free solution for hydrogen generation and storage. The recent developments of proton ceramic electrolysis cells operating at intermediate temperatures bear promise of superior energy efficiency compared to oxide ion conducting electrolytes. Here, a proton ceramic Single Engineering Unit (SEU) design is optimized for steam electrolysis using a computational fluid dynamics (CFD) model implemented in a COMSOL Multiphysics software. The SEU is an all-in-one tubular cell arrangement that constitutes the smallest electrolysis unit and enables efficient, adaptable pressurized hydrogen generation. The parametrical modelling study is conducted for two adiabatic operation scenarios with distinct steam conversion rates and tested for multiple key parameters, namely internal and external chamber pressures and inlet stream temperature. The modelling results show that low steam conversions enable operation at higher current densities and that the thermoneutral voltage for a fixed steam conversion is highly sensitive to the process conditions and operation modes. The increment of the pressure of the generated hydrogen implies a reduced production rate at thermoneutral voltage, although it can be compensated with an enhanced steam pressure or a reduced inlet temperature. Additionally, the introduction of a porous medium as the SEU current collector in the steam chamber enhances heat transport within this chamber. The area specific resistance of the system determines the current density, enforcing an adaption of the area of the electrolyser to satisfy the target hydrogen production and energy efficiency. The resulting proposed SEU design and adapted operational parameters allow effective delivery of pressurized dry hydrogen for a wide range of conditions and applications.publishedVersio

Size- and stage-dependence in cause-specific mortality of migratory brown trout

Evidence‐based management of natural populations under strong human influence frequently requires not only estimates of survival but also knowledge about how much mortality is due to anthropogenic vs. natural causes. This is the case particularly when individuals vary in their vulnerability to different causes of mortality due to traits, life history stages, or locations. Here, we estimated harvest and background (other cause) mortality of landlocked migratory salmonids over half a century. In doing so, we quantified among‐individual variation in vulnerability to cause‐specific mortality resulting from differences in body size and spawning location relative to a hydropower dam. We constructed a multistate mark–recapture model to estimate harvest and background mortality hazard rates as functions of a discrete state (spawning location) and an individual time‐varying covariate (body size). We further accounted for among‐year variation in mortality and migratory behaviour and fit the model to a unique 50‐year time series of mark–recapture–recovery data on brown trout (Salmo trutta ) in Norway. Harvest mortality was highest for intermediate‐sized trout, and outweighed background mortality for most of the observed size range. Background mortality decreased with body size for trout spawning above the dam and increased for those spawning below. All vital rates varied substantially over time, but a trend was evident only in estimates of fishers' reporting rate, which decreased from over 50% to less than 10% throughout the study period. We highlight the importance of body size for cause‐specific mortality and demonstrate how this can be estimated using a novel hazard rate parameterization for mark–recapture models. Our approach allows estimating effects of individual traits and environment on cause‐specific mortality without confounding, and provides an intuitive way to estimate temporal patterns within and correlation among different mortality sources

Model prediction vs. reality--testing the predictions of a European eel (Anguilla anguilla) stock dynamics model against the in situ observation of silver eel escapement in compliance with the European eel regulation

A direct monitoring of European silver eel (Anguilla anguilla, L) escapement from rivers and estuaries has been proven to be challenging, and a Europe-wide documentation of escaping silver eel numbers therefore hardly seems realistic. To reinforce management decisions, policy-makers are thus widely reliant on the accuracy of escapement models. A 3-year programme of silver eel escapement monitoring was undertaken to compile model input data and revise an eel population model (German Eel Model II; GEM II) already used in the decision-making process of management authorities. By compiling necessary input data and analysing vital system-specific population characteristics, it was possible to compare the documented silver eel escapement with the modelled potential silver eel escapement. Resulting model predictions were close to actually monitored escapement numbers, which were distinctly lower than reference escapement values for the same freshwater system given in the implementation report of the German Eel Management Plans. Applying different commercial and recreational catch scenarios revealed the sensitivity of the model. The results show the potential of the GEM II and highlight the importance of high-quality input data to use model predictions as the basis for management measures

A systematic review of randomised controlled trials on the effectiveness of exercise programs on lumbo pelvic pain among postnatal women

Background: A substantial number of women tend to be affected by Lumbo Pelvic Pain (LPP) following child birth. Physical exercise is indicated as a beneficial method to relieve LPP, but individual studies appear to suggest mixed findings about its effectiveness. This systematic review aimed to synthesise evidence from randomised controlled trials on the effectiveness of exercise on LPP among postnatal women to inform policy, practice and future research. Methods: A systematic review was conducted of all randomised controlled trials published between January 1990 and July 2014, identified through a comprehensive search of following databases: PubMed, PEDro, Embase, Cinahl, Medline, SPORTDiscus, Cochrane Pregnancy and Childbirth Group’s Trials Register, and electronic libraries of authors’institutions. Randomised controlled trials were eligible for inclusion if the intervention comprised of postnatal exercise for women with LPP onset during pregnancy or within 3 months after delivery and the outcome measures included changes in LPP. Selected articles were assessed using the PEDro Scale for methodological quality and findings were synthesised narratively as meta-analysis was found to be inappropriate due to heterogeneity among included studies. Results: Four randomised controlled trials were included, involving 251 postnatal women. Three trials were rated as of ‘good’ methodological quality. All trials, except one, were at low risk of bias. The trials included physical exercise programs with varying components, differing modes of delivery, follow up times and outcome measures. Intervention in one trial, involving physical therapy with specific stabilising exercises, proved to be effective in reducing LPP intensity. An improvement in gluteal pain on the right side was reported in another trial and a significant difference in pain frequency in another. Conclusion: Our review indicates that only few randomised controlled trials have evaluated the effectiveness of exercise on LPP among postnatal women. There is also a great amount of variability across existing trials in the components of exercise programs, modes of delivery, follow up times and outcome measures. While there is some evidence to indicate the effectiveness of exercise for relieving LPP, further good quality trials are needed to ascertain the most effective elements of postnatal exercise programs suited for LPP treatment