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Inulin affects nutrition and metabolism in many animals. Although inulin is widely used in the diet of teleosts, its mechanism of action is unknown. Here, we investigated the effect of inulin (2 %) on the intestinal microbiome and metabolism in rainbow trout (Oncorhynchus mykiss) selected for growth and survival when fed a 100 % plant-based diet (suave) and a control line (temoin). Metabolic responses to the two factors (line and inulin) in liver, intestine, muscle and adipose were tissue-specific, with line and interaction between the two factors influencing overall expression in liver. In the intestine, inulin and line and in muscle, line influenced the expression of metabolic genes. Microbiota between the mucus and digestive contents was significantly different, with genera from Proteobacteria being more abundant in the mucus, whereas genera from the Firmicutes and Planctomycetes being more abundant in contents. Effect of inulin and interaction between factors on the microbiome was evident in contents. The significant taxa of control and inulin-fed groups differed greatly with Streptococcus and Weissella being significantly abundant in the inulin-fed group. There was a general trend showing higher levels of all SCFA in temoin group with propionic acid levels being significantly higher. An operational taxonomic unit (OTU) belonging to the Ruminococcaceae was significantly abundant in suave. The tissue-specific correlations between OTU and gene expression may indicate the link between microbiome and metabolism. Together, these results suggest that line and inulin impact the gene expression in a tissue-specific manner, possibly driven by specific OTUs enriched in inulin-fed groups and suave.

International audience; The importance of poly-unsaturated fatty acids (PUFAs) in food is crucial for the animal and human development and health. As a complementary strategy to nutrition approaches, genetic selection has been suggested to improve fatty acids (FAs) composition in farmed fish. Gas chromatography (GC) is used as a reference method for the quantification of FAs; nevertheless, the high cost prevents large scale phenotyping as needed in breeding programs. Therefore, a calibration by means of Raman scattering spectrometry has been established in order to predict FA composition in rainbow trout Onchorhynchus mykiss adipose tissue. FA composition of visceral adipose tissue was analysed by both GC and Raman micro-spectrometry techniques on 268 individuals fed with three different feeds, which have different FA compositions. Among the possible regression methods, the ridge regression method, was found to be efficient to establish calibration models from the GC and spectral data. The best cross-validated R2 values were obtained for total PUFAs, omega-6 (Ω-6) and omega-3 (Ω-3) PUFA (0.79, 0.83 and 0.66, respectively). For individual Ω-3 PUFAs, α-linolenic acid (ALA, C18:3), eicosapentaenoic acid (EPA, C20:5) and docosahexenoic acid (DHA, C22:6) were found to have the best R2 values (0.82, 0.76 and 0.81, respectively). This study demonstrates that Raman spectroscopy could be used to obtain good correlation coefficients on adipocytes allowing to predict PUFAs, and calibration models can be used to predict PUFAs contents for large scale and high throughput phenotyping in rainbow trout

Background In response to major challenges regarding the supply and sustainability of marine ingredients in aquafeeds, the aquaculture industry has made a large-scale shift toward plant-based substitutions for fish oil and fish meal. But, this also led to lower levels of healthful n−3 long-chain polyunsaturated fatty acids (PUFAs)—especially eicosapentaenoic (EPA) and docosahexaenoic (DHA) acids—in flesh. One potential solution is to select fish with better abilities to retain or synthesise PUFAs, to increase the efficiency of aquaculture and promote the production of healthier fish products. To this end, we aimed i) to estimate the genetic variability in fatty acid (FA) composition in visceral fat quantified by Raman spectroscopy, with respect to both individual FAs and groups under a feeding regime with limited n-3 PUFAs; ii) to study the genetic and phenotypic correlations between FAs and processing yields- and fat-related traits; iii) to detect QTLs associated with FA composition and identify candidate genes; and iv) to assess the efficiency of genomic selection compared to pedigree-based BLUP selection. Results Proportions of the various FAs in fish were indirectly estimated using Raman scattering spectroscopy. Fish were genotyped using the 57 K SNP Axiom™ Trout Genotyping Array. Following quality control, the final analysis contained 29,652 SNPs from 1382 fish. Heritability estimates for traits ranged from 0.03 ± 0.03 (n-3 PUFAs) to 0.24 ± 0.05 (n-6 PUFAs), confirming the potential for genomic selection. n-3 PUFAs are positively correlated to a decrease in fat deposition in the fillet and in the viscera but negatively correlated to body weight. This highlights the potential interest to combine selection on FA and against fat deposition to improve nutritional merit of aquaculture products. Several QTLs were identified for FA composition, containing multiple candidate genes with indirect links to FA metabolism. In particular, one region on Omy1 was associated with n-6 PUFAs, monounsaturated FAs, linoleic acid, and EPA, while a region on Omy7 had effects on n-6 PUFAs, EPA, and linoleic acid. When we compared the effectiveness of breeding programmes based on genomic selection (using a reference population of 1000 individuals related to selection candidates) or on pedigree-based selection, we found that the former yielded increases in selection accuracy of 12 to 120% depending on the FA trait. Conclusion This study reveals the polygenic genetic architecture for FA composition in rainbow trout and confirms that genomic selection has potential to improve EPA and DHA proportions in aquaculture species.

For the sustainable development of aquaculture, vegetable ingredients have been used to replace the traditional ingredients fishmeal and fish oil in aquafeeds. On the other hand, selective breeding has also been implemented at research level to obtain strains that are adapted to feeding on the plant-based diet. To better understand the underlying mechanisms prompting the adaptation to plant-based diets in fish, we investigated the hepatic expression of several microRNAs (miRNAs) that are involved in the post-transcriptional regulation of cholesterol and lipid metabolism at 8 h and 16 h after the last meal in two lines of rainbow trout: one selected for better adaptation to plant-based diets and the corresponding control line. Both groups were fed either a fishmeal and fish oil based diet or a 100% plant-based diet. Results showed that the expression of miR-33a in liver was greatly elevated in trout fed the plant-based diet, while the expression of miR-122 and miR-128 was much higher in the selected line at 8 h after the last meal regardless of the diet. Furthermore, our results indicated that some genes involved in immune processes (caspase 6 apoptosis-related cysteine peptidase like 2, casp6l2) and cAMP signal transduction (phosphodiesterase 4B cAMP-specific a, pde4ba) were also potentially regulated by miRNAs. They were newly identified as putative direct targets of miRNAs and affected in trout fed the plant-based diet. Though further investigations are still needed to establish a valid relationship between miRNAs and their target genes, our study found miR-33a, miR-122 and miR-128 as potential candidates for further study and provided new perspectives to understand the role of miRNAs in the selective breeding for adaptation to the plant-based diets.

Within the context of a growing aquaculture production coupled with a plateau of the production in the main components of aquafeeds (fish oil and fishmeal), recent studies have typically focused on replacing these feedstuffs with terrestrial plant ingredients for cultured carnivorous aquatic species, such as rainbow trout (Oncorhynchus mykiss). Substitution rates without adverse effects have, however, reached their limit. One potential way forward would be to take advantage of the genetic variability that exists in the salmonid population. However, to date, little is known about the underlying molecular mechanisms responsible for this genetic variability. The aim of the present research was to understand why some genotypes are better able to utilize plant-based diets devoid of marine resources. In this regard, three isogenic lines of rainbow trout (R23h, AB1h, and A22h), with similar growth when fed marine resources-based diets and which differ greatly in their responses to a plant-based diet, were fed with either a complete plant-based diet (V diet) or a marine resources-based diet (M diet) since first-feeding. Fish traits and the hepatic transcriptome of these three genotypes were compared after 5 months of feeding. First, differences in the ability to grow with the V diet observed amongst genotypes was not due to higher feed intake, but instead due to differences in feed efficiency. The comparison of transcriptome profiles revealed 575 (R23h vs. AB1h), 1,770 (R23h vs. A22h), and 2,973 (AB1h vs. A22h) probes differentially expressed amongst the three genotypes when fed the V diet. Interestingly, R23h and AB1h fish, which were the least affected by the V diet, exhibited the highest growth. These results demonstrate that these fish were able to maintain a high level of energy production and protein synthesis. Moreover, these genotypes were also able to activate pathways linked to lipid and cholesterol metabolisms, such as the biosynthesis of long-chain polyunsaturated fatty acids. Finally, as previously, immunity seems to also play an important role in the ability of fish to use the V diet, and further studies are needed to understand the mechanisms by which immunity interacts with growth.

One of the top priorities of the aquaculture industry is the genetic improvement of economically important traits in fish, such as those related to processing and quality. However, the accuracy of genetic evaluations has been hindered by a lack of data on such traits from a sufficiently large population of animals. The objectives of this study were thus threefold: (i) to estimate genetic parameters of growth-, yield-, and quality-related traits in rainbow trout (Oncorhynchus mykiss) using three different phenotyping technologies [invasive and non-invasive: microwave-based, digital image analysis, and magnetic resonance imaging (MRI)], (ii) to detect quantitative trait loci (QTLs) associated with these traits, and (iii) to identify candidate genes present within these QTL regions. Our study collected data from 1,379 fish on growth, yield-related traits (body weight, condition coefficient, head yield, carcass yield, headless gutted carcass yield), and quality-related traits (total fat, percentage of fat in subcutaneous adipose tissue, percentage of fat in flesh, flesh colour); genotypic data were then obtained for all fish using the 57K SNP Axiom® Trout Genotyping array. Heritability estimates for most of the 14 traits examined were moderate to strong, varying from 0.12 to 0.67. Most traits were clearly polygenic, but our genome-wide association studies (GWASs) identified two genomic regions on chromosome 8 that explained up to 10% of the genetic variance (cumulative effects of two QTLs) for several traits (weight, condition coefficient, subcutaneous and total fat content, carcass and headless gutted carcass yields). For flesh colour traits, six QTLs explained 1–4% of the genetic variance. Within these regions, we identified several genes (htr1, gnpat, ephx1, bcmo1, and cyp2x) that have been implicated in adipogenesis or carotenoid metabolism, and thus represent good candidates for further functional validation. Finally, of the three techniques used for phenotyping, MRI demonstrated particular promise for measurements of fat content and distribution, while the digital image analysis-based approach was very useful in quantifying colour-related traits. This work provides new insights that may aid the development of commercial breeding programmes in rainbow trout, specifically with regard to the genetic improvement of yield and flesh-quality traits as well as the use of invasive and/or non-invasive technologies to predict such traits.

Background Rainbow trout is a significant fish farming industry under temperate climates. Female reproduction traits play an important role in the economy of breeding companies with the sale of fertilized eggs. The objectives of this study are threefold: to estimate the genetic parameters of female reproduction traits, to determine the genetic architecture of these traits by the identification of quantitative trait loci (QTL), and to assess the expected efficiency of a pedigree-based selection (BLUP) or genomic selection for these traits. Results A pedigreed population of 1,343 trout were genotyped for 57,000 SNP markers and phenotyped for seven traits at 2 years of age: spawning date, female body weight before and after spawning, the spawn weight and the egg number of the spawn, the egg average weight and average diameter. Genetic parameters were estimated in multi-trait linear animal models. Heritability estimates were moderate, varying from 0.27 to 0.44. The female body weight was not genetically correlated to any of the reproduction traits. Spawn weight showed strong and favourable genetic correlation with the number of eggs in the spawn and individual egg size traits, but the egg number was uncorrelated to the egg size traits. The genome-wide association studies showed that all traits were very polygenic since less than 10% of the genetic variance was explained by the cumulative effects of the QTLs: for any trait, only 2 to 4 QTLs were detected that explained in-between 1 and 3% of the genetic variance. Genomic selection based on a reference population of only one thousand individuals related to candidates would improve the efficiency of BLUP selection from 16 to 37% depending on traits. Conclusions Our genetic parameter estimates made unlikely the hypothesis that selection for growth could induce any indirect improvement for female reproduction traits. It is thus important to consider direct selection for spawn weight for improving egg production traits in rainbow trout breeding programs. Due to the low proportion of genetic variance explained by the few QTLs detected for each reproduction traits, marker assisted selection cannot be effective. However genomic selection would allow significant gains of accuracy compared to pedigree-based selection.

International audience; Fillet yield (i.e. the proportion of edible muscle in a fish) is a key economic trait for species sold as fillets. Its genetic improvement is complicated by several of its characteristics 1) it is a ratio trait, 2) its numerator (fillet weight) and denominator (body weight) are strongly correlated (correlations in the range 0.89-0.99), 3) it offers little phenotypic variation and 4) it cannot be measured on alive breeding candidates. In a former study, we showed that it could be improved by selection, especially with three selection indices, fillet yield, residual fillet weight and a ratio-specific linear index. However, it is well known that the heritability of ratio traits does not permit a reliable prediction of genetic gains. As predictability of genetic gains is a key requirement to define breeding programs, we investigated how genetic gains in fillet yield could be predicted by the genetic parameters of fillet yield, of residual fillet weight and of the component traits of the linear index. To this end, we compared simulated genetic gains with those estimated by classical prediction methods. This was done using real sets of genetic parameters obtained in nine populations of rainbow trout, European sea bass, gilthead sea bream and common carp. We show that the genetic parameters of fillet yield cannot be used to reliably predict genetic gains in fillet yield. Conversely, selection index theory using a linear index, combining either fillet weight and body weight or fillet weight and waste weight, provides almost perfect prediction of gains. Still, it is highly sensitive to the precision of the genetic and phenotypic correlations estimates, which should not be rounded to less than three decimals for fillet weight and body weight, while two decimals are appropriate for fillet weight and waste weight. A simple, reasonably precise alternative to the linear index is the use of residual fillet weight (the residual of the regression of fillet weight on body weight) as a surrogate for fillet yield.

BackgroundRainbow trout is a significant fish farming species under temperate climates. Female reproduction traits play an important role in the economy of breeding companies with the sale of fertilized eggs. The objectives of this study are threefold: to estimate the genetic parameters of female reproduction traits, to determine the genetic architecture of these traits by the identification of quantitative trait loci (QTL), and to assess the expected efficiency of a pedigree-based selection (BLUP) or genomic selection for these traits.ResultsA pedigreed population of 1343 trout were genotyped for 57,000 SNP markers and phenotyped for seven traits at 2 years of age: spawning date, female body weight before and after spawning, the spawn weight and the egg number of the spawn, the egg average weight and average diameter. Genetic parameters were estimated in multi-trait linear animal models. Heritability estimates were moderate, varying from 0.27 to 0.44. The female body weight was not genetically correlated to any of the reproduction traits. Spawn weight showed strong and favourable genetic correlation with the number of eggs in the spawn and individual egg size traits, but the egg number was uncorrelated to the egg size traits. The genome-wide association studies showed that all traits were very polygenic since less than 10% of the genetic variance was explained by the cumulative effects of the QTLs: for any trait, only 2 to 4 QTLs were detected that explained in-between 1 and 3% of the genetic variance. Genomic selection based on a reference population of only one thousand individuals related to candidates would improve the efficiency of BLUP selection from 16 to 37% depending on traits.ConclusionsOur genetic parameter estimates made unlikely the hypothesis that selection for growth could induce any indirect improvement for female reproduction traits. It is thus important to consider direct selection for spawn weight for improving egg production traits in rainbow trout breeding programs. Due to the low proportion of genetic variance explained by the few QTLs detected for each reproduction traits, marker assisted selection cannot be effective. However genomic selection would allow significant gains of accuracy compared to pedigree-based selection.

International audience; Phenotypic plasticity is a key component of the ability of organisms to respond to changing environmental conditions. In this study, we aimed to study the establishment of DNA methylation marks in response to an environmental stress in rainbow trout and to assess whether these marks depend on the genetic background. The environmental stress chosen here was temperature, a known induction factor of epigenetic marks in fish. To disentangle the role of epigenetic mechanisms such as DNA methylation in generating phenotypic variations, nine rainbow trout isogenic lines with no genetic variability within a line were used. For each line, half of the eggs were incubated at standard temperature (11°C) and the other half at high temperature (16°C), from eyed-stage to hatching. In order to gain a first insight into the establishment of DNA methylation marks in response to an early temperature regime (control 11°C vs. heated 16°C), we have studied the expression of 8 dnmt3 (DNA methyltransferase) genes, potentially involved in de novo methylation, and analysed global DNA methylation in the different rainbow trout isogenic lines using LUMA (LUminometric Methylation Assay). Finally, finer investigation of genome-wide methylation patterns was performed using EpiRADseq, a reduced-representation library approach based on the ddRADseq (Double Digest Restriction Associated DNA) protocol, for six rainbow trout isogenic lines. We have demonstrated that thermal history during embryonic development alters patterns of DNA methylation, but to a greater or lesser extent depending on the genetic background.

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International audience; Fillet yield (i.e. the proportion of edible flesh from a given amount of fish) is a trait of high economic interest for species sold as fillets. Improving it by selective breeding is not an easy task, as it is a ratio trait, which causes mathematical difficulties. It is moreover a specific ratio where numerator ( fillet weight) and denominator (body weight) are very strongly correlated (genetic and phenotypic correlations in the range 0.89 – 0.99). This led some authors to conclude that they have the same genetic and phenotypic basis, precluding selection for improved fillet yield. In this study, we propose to study fillet yield as a component of two traits, fillet weight and waste weight (waste weight = body weight- fillet weight, so the sum of head, bones, fins and viscera weight), which we expect to be less correlated. Using data from 5 batches of fish from 3 species (sea bass, sea bream, rainbow trout), we show that as expected, fillet weight and waste weight are less correlated than fillet weight and body weight (on average, r A = 0.91 and r P = 0.88 vs. r A = 0.987, r P = 0.981). We used stochastic simulation to generate genotypes and phenotypes of fish using those genetic parameters for fillet weight and waste weight. We simulated 10 generations of selection for increased fillet yield using nine selection indices. Five indices had rather similar performances, residual fillet weight (the residual of the regression of fillet weight to body weight), fillet yield, fillet to waste ratio, restricted selection index (a linear index aimed at improving fillet weight while keeping waste weight constant) and linear index (optimized to improve fillet/waste ratio). With these indices, the average selection gain was +0.66% of fillet yield per generation (range 0.30 to 0.95% using real genetic parameters from 5 fish batches). Selection for the difference between fillet weight and waste weight was 30% less efficient, while selection for increased fillet weight or increased body weight was 55 – 65% less efficient. Selection against waste weight had a null or even negative impact on fillet yield. Factors favorable to higher selection gains are low initial fillet yield, different heritabilities and CVs of fillet weight and waste weight, low genetic correlation and high phenotypic correlation of fillet weight and waste weight. These results suggest that although fillet weight and body weight are strongly correlated and proportional to each other, moderate selection gains on fillet yield are possible. We consider it would be useful to add waste weight in the parameters recorded in future genetic studies on fillet yield.

When compared with fish meal and fish oil, plant ingredients differ not only in their protein content and amino acid and fatty acid profiles but are also devoid of cholesterol, the major component of cell membrane and precursor of several bioactive compounds. Based on these nutritional characteristics, plant-based diets can affect fish physiology and cholesterol metabolism. To investigate the mechanisms underlying cholesterol homeostasis, rainbow trout were fed from 1 g body wt for 6 mo with a totally plant-based diet (V), a marine diet (M), and a marine-restricted diet (MR), with feed intake adjusted to that of the V group. The expression of genes involved in cholesterol synthesis, esterification, excretion, bile acid synthesis, and cholesterol efflux was measured in liver. Results showed that genes involved in cholesterol synthesis were upregulated in trout fed the V diet, whereas expression of genes related to bile acid synthesis ( cyp7a1) and cholesterol elimination ( abcg8) were reduced. Feeding trout the V diet also enhanced the expression of srebp-2 while reducing that of lxrα and miR-223. Overall, these data suggested that rainbow trout coped with the altered nutritional characteristics and absence of dietary cholesterol supply by increasing cholesterol synthesis and limiting cholesterol efflux through molecular mechanisms involving at least srebp-2, lxrα, and miR-223. However, plasma and body cholesterol levels in trout fed the V diet were lower than in fish fed the M diet, raising the question of the role of cholesterol in the negative effect of plant-based diet on growth.

Rainbow trout (Oncorhynchus mykiss) was recognized as a typical ‘glucose-intolerant’ fish and poor dietary carbohydrate user. Our first objective was to test the effect of dietary carbohydrates themselves (without modification of dietary protein intake) on hepatic glucose gene expression (taking into account the paralogs). The second aim was to research if two isogenic trout lines had different responses to carbohydrate intake, showing one with a better use dietary carbohydrates. Thus, we used two isogenic lines of rainbow trout (named A32h and AB1h) fed with either a high carbohydrate diet or a low carbohydrate diet for 12 weeks. We analysed the zootechnical parameters, the plasma metabolites, the hepatic glucose metabolism at the molecular level and the hormonal-nutrient sensing pathway. Globally, dietary carbohydrate intake was associated with hyperglycaemia and down regulation of the energy sensor Ampk, but also with atypical regulation of glycolysis and gluconeogenesis in the liver. Indeed, the first steps of glycolysis and gluconeogenesis catalysed by the glucokinase and the phospenolpyruvate carboxykinase are regulated at the molecular level by dietary carbohydrates as expected (i.e. induction of the glycolytic gck and repression of the gluconeogenic pck); by contrast, and surprisingly, for two other key glycolytic enzymes (phosphofructokinase enzyme – pfkl and pyruvate kinase – pk) some of the paralogs (pfklb and pklr) are inhibited by carbohydrates whereas some of the genes coding gluconeogenic enzymes (the glucose-6-phosphatase enzyme g6pcb1b and g6pcb2a gene and the fructose1-6 biphosphatase paralog fbp1a) are induced. On the other hand, some differences for the zootechnical parameters and metabolic genes were also found between the two isogenic lines, confirming the existence of genetic polymorphisms for nutritional regulation of intermediary metabolism in rainbow trout. In conclusion, our study determines some new and unexpected molecular regulations of the glucose metabolism in rainbow trout which may partly lead to the poor utilization of dietary carbohydrates and it underlines the existence of differences in molecular regulation of glucose metabolism between two isogenic lines which provides arguments for future selection of rainbow trout., Summary: Using isogenic lines, this study determines some new, unexpected molecular regulation of the glucose metabolism in rainbow trout, which may partly lead to the poor utilization of dietary carbohydrates.

International audience; Marker-based parentage assignment provides the opportunity to investigate factors of efficiency for mixed-family designs and factorial mating. In such designs, family size is both uncontrolled and small, which may be thought to limit the accuracy of estimated breeding values (EBVs). The objective of this work was to estimate the accuracy of EBVs of growth and quality traits in a large factorial mating design and in commercial breeding conditions. An expected six hundred full-sib families of rainbow trout Oncorhynchus mykiss (2042 fish in total) were produced by ten factorial matings of six dams with ten sires. Fish were phenotyped for body weight, carcass yield, fillet yield, fillet fat content and fi llet colour, and family information was recovered using microsatellite markers. The accuracy of EBVs was estimated using or removing individual performance to mimic combined family selection (with individual phenotype) or sib selection (without individual phenotype). The traits investigated had medium to high heritability (0.17–0.58). High to very high accuracy (0.630–0.817) was estimated for combined family selection. The accuracy of sib selection (not using individual phenotype) was 18–22% lower (0.542–0.638), but remained in the upper range reported for such traits. This level of accuracy was higher than those reported in conventional breeding programs using separate family rearing. This was true even for families with a very low number of full-sibs. Individual EBV accuracy was more closely linked to the total number of full- and half-sibs of each fish than to its number of full-sibs. We hypothesize that this was due to the factorial mating, which led to a high number of the genetic ties between sibs. These results highlight the possibility of introducing precise estimated breeding values for quality traits into combined or sib selection in breeding programs when using mixed families from factorial designs and marker- based parentage assignment in aquaculture species

Ajouter clé UT WOS; Describing and explaining the geographic within-species variation in phenotypes ("phenogeography") in the sea over a species distribution range is central to our understanding of a variety of eco-evolutionary topics. However, phenogeographic studies that have a large potential to investigate adaptive variation are overcome by phylogeographic studies, still mainly focusing on neutral markers. How genotypic and phenotypic data could covary over large geographic scales remains poorly understood in marine species. We crossed 75 noninbred sires (five origins) and 26 dams (two origins; each side of a hybrid zone) in a factorial diallel cross in order to investigate geographic variation for early survival and sex ratio in the metapopulation of the European sea bass (Dicentrarchus labrax), a highly prized marine fish species. Full-sib families (N = 1,950) were produced and reared in a common environment. Parentage assignment of 7,200 individuals was performed with seven microsatellite markers. Generalized linear models showed significant additive effects for both traits and pleiotropy between traits. A significant nonadditive genetic effect was detected. Different expression of traits and distinct relative performances were found for reciprocal crosses involving populations located on each side of the main hybrid zone located at the Almeria-Oran front, illustrating asymmetric reproductive isolation. The poor fitness performance observed for the Western Mediterranean population of sea bass is discussed as it represents the main source of seed hatchery production, but also because it potentially illustrates nonadaptive introgression and maladaptation.

Selection to improve processing yields relies on sib selection, in which live candidates are ranked according to their family breeding value. This approach limits genetic progress, as it only exploits genetic variability between families and not within them. Indirect criteria measured on live candidates could overcome this limitation. The present study (1) proposes a procedure to identify indirect criteria to predict processing yields in rainbow trout (head, carcass and fillet yields), (2) estimates genetic parameters of these indirect criteria, and (3) predicts relative genetic gains in processing yields using full-sib selection or indirect individual selection on those indirect criteria. DNA-pedigreed all-female rainbow trout Oncorhynchus mykiss (n = 2029, 1631.0 +/- 355.6 g) from 600 families produced from 100 sires and 60 dams were characterized by external and internal non-lethal morphological measures using digital pictures and real time ultrasound tomography. Nineteen landmarks were recorded on the digital pictures to define the outline of the body, head and lateral line. Their coordinates were used to calculate different lengths, heights and areas. Five different internal thicknesses were measured by ultrasound tomography. In the first phase of this study, processing yields were predicted using multiple linear regressions including both external and internal morphometric variables. In a second phase, the heritability of the predicted values and their genetic correlations with real processing yields were estimated using animal models. Predicted yields exhibited intermediate heritabilities (0.25-0.28) that were half the value of heritabilities for real processing yields (0.47-0.55), but had high genetic correlations with these real yields (0.87-0.90). The relative efficiency of indirect selection (IS) on these indirect criteria was compared to theoretical mass selection (MS) or sib selection (FS) with different family sizes (10 or 100) and two different selection pressures (10% or 40%). At the same selection pressure (10%, with 100 sibs per family %), full-sib selection created genetic progress 49.6% to 60.5% higher than indirect selection according to the processing yield targeted. However, when sib-selection pressure was limited to a more realistic between family selection pressure (40% and 10 sibs per family), indirect selection with 10% selection pressure was 21.9% to 32.7% more efficient than sib selection.

Aquaculture of carnivorous species has strongly relied on fish meal and fish oil for feed formulation; however, greater replacement by terrestrial plant-based products is occurring now. This rapid change in dietary environment has been a major revolution and has to be taken into consideration in breeding programs. The present study analyzes potential consequences of this nutritional tendency for selective breeding by estimating genetic parameters of BW and growth rates estimated by the thermal growth coefficient (TGC) over different periods with extremely different diets. European sea bass (Dicentrarchus labrax L.) from a factorial cross (1,526 fish) between 25 sires and 9 dams were used to estimate heritabilities and genotype by diet interaction. Starting 87 d after fertilization (2.5 g), one-half of the sea bass were fed a diet containing marine products (M), and the other one-half were fed a totally plant-based (PB) diet (without any fish meal or fish oil). The fish were individually tagged, reared in a recirculated system, and genotyped at 13 microsatellites to rebuild parentage of individuals. Body weight and TGC were measured for 335 d until fish fed the M diet reached 108.3 g of BW. These traits were significantly less in fish fed the PB diet (P 0.1). Survival was significantly less in fish fed the PB diet (PB=64.7%, M=93.7% after 418 d, P

Sea bass is a major species in Mediterranean aquaculture, and is now being subject to selective breeding programmes for faster growth. In terrestrial species, it was demonstrated that fast growth may be linked to a correlated degradation of fitness traits. In this experiment, we evaluated 600 young sea bass from a factorial mating of 76 sires and 13 dams. The sires were from four genetic groups, wild (W), domesticated (D), and selected for growth (2 groups, M and P). The 600 offspring were submitted to two acute confinement stress challenges at 6 weeks intervals, and plasma cortisol at one hour post stress was measured. The same fish were also submitted to two swimming challenges at a 5 days interval, where the maximum sustained swimming speed ( U max ) of each fish was evaluated. Parentage was assessed by genotyping of 12 microsatellites. 554 fish had both valid parentage and phenotypes. Cortisol had a low repeatability (r = 0.30 between the two successive measurements) while repeatability was moderate for U max (r = 0.62). However, genetic correlations between successive measurements were very high (> 0.96) for both traits, indicating that successive measurements were related to the same trait. Heritability was moderate for mean post-stress cortisol (h 2 = 0.34 ± 0.09) and U max (h 2 = 0.48 ± 0.08). When U max was expressed in m.s − 1 , it was negatively correlated to cortisol (r A = − 0.48 ± 0.08) and weakly correlated to body weight (r A = 0.12 ± 0.16), but figures changed when it was expressed in Body Lengths.s − 1 (h 2 = 0.55 ± 0.08, r A = − 0.10 ± 0.19 with cortisol and r A = − 0.64 ± 0.07 with body weight, respectively). Cortisol was moderately negatively correlated with body weight (r A = − 0.36 ± 0.18). The four lines did not differ for cortisol or U max , but when U max was expressed in BL.s − 1 it tended to be lower in the two selected lines — which were also significantly larger. However, this is likely due to a phenotypic decrease of relative U max with increasing body size. We conclude that selection for growth and/or domestication should not impact maximum sustained swimming speed in the European sea bass, but may tend to favour animals with low cortisol responsiveness. These traits could be used to orientate functional capabilities other than productivity in sea bass. Statement of relevance We estimate heritability of cortisol stress response and (for the first time) in European sea bass, as well as their correlations with growth. We show moderate correlations, and no correlated response to selection for growth. We also provide methods to evaluate these traits on large number of fishes. This can be useful to monitor and design breeding programmes.

Background The achievement of sustainable feeding practices in aquaculture by reducing the reliance on wild-captured fish, via replacement of fish-based feed with plant-based feed, is impeded by the poor growth response seen in fish fed high levels of plant ingredients. Our recent strategy to nutritionally program rainbow trout by early short-term exposure to a plant-based (V) diet versus a control fish-based (M) diet at the first-feeding fry stage when the trout fry start to consume exogenous feed, resulted in remarkable improvements in feed intake, growth and feed utilization when the same fish were challenged with the diet V (V-challenge) at the juvenile stage, several months following initial exposure. We employed microarray expression analysis at the first-feeding and juvenile stages to deduce the mechanisms associated with the nutritional programming of plant-based feed acceptance in trout. Results Transcriptomic analysis was performed on rainbow trout whole fry after 3 weeks exposure to either diet V or diet M at the first feeding stage (3-week), and in the whole brain and liver of juvenile trout after a 25 day V-challenge, using a rainbow trout custom oligonucleotide microarray. Overall, 1787 (3-week + Brain) and 924 (3-week + Liver) mRNA probes were affected by the early-feeding exposure. Gene ontology and pathway analysis of the corresponding genes revealed that nutritional programming affects pathways of sensory perception, synaptic transmission, cognitive processes and neuroendocrine peptides in the brain; whereas in the liver, pathways mediating intermediary metabolism, xenobiotic metabolism, proteolysis, and cytoskeletal regulation of cell cycle are affected. These results suggest that the nutritionally programmed enhanced acceptance of a plant-based feed in rainbow trout is driven by probable acquisition of flavour and feed preferences, and reduced sensitivity to changes in hepatic metabolic and stress pathways. Conclusions This study outlines the molecular mechanisms in trout brain and liver that accompany the nutritional programming of plant-based diet acceptance in trout, reinforces the notion of the first-feeding stage in oviparous fish as a critical window for nutritional programming, and provides support for utilizing this strategy to achieve improvements in sustainability of feeding practices in aquaculture. Electronic supplementary material The online version of this article (doi:10.1186/s12864-016-2804-1) contains supplementary material, which is available to authorized users.

The composition of feed for farmed salmonids has strongly evolved during the last decades due to the substitution of fishery-derived fish oil and fishmeal by ingredients of plant origin. Little information is available regarding the effects of this transition on adaptive capacities in fish. Two rainbow trout isogenic lines, known for their divergent ability to grow on a plant-based diet (PBD), were fed for seven months from first feeding either a fully PBD or a control marine-resources diet and were compared for their growing and survival capacities over time and their behavioral and stress responses at similar sizes but different ages. Although fish displayed similar appetitive behaviour, the two lines were highly affected by the PBD translated in decreased growth and apathetic behaviour, but also stronger stress responses displayed by stronger cortisol increases and more stress-related behaviour when isolated. The two lines were found to be similarly sensitive to a PBD for the assessed stress-related parameters, but one line displayed a lower survival during the early rearing period. Overall, these results suggest that a PBD supplied to fish from the alevin stage has strong effects on physiological and behavioural parameters, with possible impairment of fish welfare, but also genome-dependent survival.

This preliminary study assessed genotype _ diet interaction in late growth of European sea bass (Dicentrarchus labrax) fed with either a ￠sh meal (FM)- or a ￠sh oil (FO)-based diet (M) or an all-plant-based (PB) diet. Atotal of 550 ￠sh from 224 families were reared together and tagged. DNAwas sampled and microsatellites were used to assign parentage. When ￠sh weight was 192 _ 54 g, two tanks were fed with M (FM: 100%; FO: 100%) and two others with PB (FM: 0% and FO: 0%). Body weight (BW), fork length (FL) and ￠llet lipid content (CorrFat) were analysed with a linear model and with REML methodology.We observed no signi￠cant di!erences between groups, but a slightly lower (P50.03) daily growth coe⁄cient in sea bass fed PB than in those fed M. Heritability estimates of BWdi!ered signi￠cantly from zero (PB: 0.37 _ 0.18; M: 0.47 _ 0.24). Sire _ diet interactions were signi￠cant and genetic correlations ranged between 0.51 and 0.87, showing genotype _ diet interaction for BWand CorrFat. For the ￠rst time, genetic parameters in the context of total replacement of marine ￠shery by-products were estimated in European sea bass, showing re-ranking of family performances with extremely contrasted diets.

Malformations in farmed fish are a prime research topic as it has a major impact on fish welfare and on the economical value of the product. In this study, the genetic component of spine deformities was estimated in European sea bass, a major species of Mediterranean aquaculture, with the use of a posteriori reconstruction of pedigree with microsatellites. Our population exhibited a very high malformation rate (81%),probably due to intense swimming of fish during pre-growing The heritability of global spine deformities was evaluated to be 0.21 +/- 0.04 on the underlying scale (033 +/- 0.06 for lordosis, 0.13 +/- 0.04 for scoliosis). Deformities scored by an internal or external examination turned out to be genetically the same trait (genetic correlations>0.9), although phenotypic correlations were moderate to high (0.45 +/- 0.04 to 0.92 +/- 0.01) due to difficulties in scoring the less severe deformities. Along time (from 100 to 800 g mean weight), lordosis had a constant heritability, with very high genetic correlations between stages (>0.9). its incidence increased from 39 to 53% indicating that the malformation was probably present from the beginning of the rearing period but that its expression increased overtime. A positive linkage between spine deformities and growth rate was demonstrated both by the higher early weight of affected fish and by the positive genetic correlation between weight and malformation (from 0.21 +/- 0.15 to 0.40 +/- 0.14), although deformities have a negative phenotypic impact on growth. However, heritabilities of malformation traits calculated here are only estimates that may indirectly describe the sensitivity of fish to the particular pre-growing environment as a function of different body weight genotypes. A quantification of realised heritability would therefore be necessary before concluding on the existence of a genetic basis for spinal malformations such as lordosis and scoliosis. Nevertheless and by precaution, selection for fast growth rate needs careful monitoring of deformities, and selection against malformations may counteract impacts of selection for fast growth. (c) 2009 Elsevier B.V. All rights reserved.

Genetic parameters for carcass quality traits were estimated in 27 families of sea bass (3 dams x 9 sires factorial mating design), raised mixed in the same tanks starting before hatching. O. spring parentage was determined a posteriori using 6 microsatellite loci. Carcass quality traits were recorded at 818 days post fertilization (mean standard length: 32.6 +/- 3.1 cm). Genetic parameters were estimated from the sire half sib variance and covariance components. Heritability of body weight (BW) and carcass processing traits (standardized to body weight) percent head weight (Head%), percent viscera weight (Viscera%) and percent visceral fat weight (VisceFat%) were relatively high ranging from 0.48 +/- 0.15 (Viscera%) to 0.87 +/- 0.23 (Head%); the estimate of heritability for fillet yield (Fillet%) was lower (0.25 +/- 0.10) but was significantly greater than zero. Body weight was positively correlated to Fillet%, Viscera%, and VisceFat%, and negatively correlated to Head%. These results indicate that these carcass processing traits can be modified by directional selection and that selection for greater BW would lead to an increase of Fillet%, Viscera% and VisceFat% and a decrease of Head%. Muscle lipid content (MuscleLipid%) was determined using two indirect methods: measurements with a Torry Fish Fatmeter (R) (TorryLipid) and determination of the percentage of dry matter content (MuscleDry%) via desiccation. Both measures were highly correlated to chemical measurements of MuscleLipid%. Regression analysis indicated a superior predictive value of TorryLipid suggesting that MuscleLipid% may be evaluated via rapid, non lethal measurements with a Torry Fish Fatmeter. Heritability estimates of TorryLipid and MuscleDry% differed significantly from zero (TorryLipid: 0.28 +/- 0.12, MuscleDry%: 0.36 +/- 0.14) indicating that MuscleLipid% could be lowered by directional selection. TorryLipid and MuscleDry% were weakly correlated to body weight and carcass processing traits suggesting that simultaneous improvement of MuscleLipid% and other carcass quality traits may require definition of multi-trait selection indices.

Terrestrial plant resources are increasingly used as substitutes for fish meal and fish oil in fish feed in order to reduce the reliance of aquaculture on marine fishery resources. Although many studies have been conducted to assess the effects of such nutritional transition, no whole breeding cycles of fish fed diets free from marine resources has been reported to date. We therefore studied the reproductive performance of trout after a complete cycle of breeding while consuming a diet totally devoid of marine ingredients and thus of n-3 long chain polyunsaturated fatty acids (n-3 LC-PUFAs) that play a major role in the formation of ova. Two groups of female rainbow trout were fed from first feeding either a commercial diet (C, marine and plant ingredients), or a 100% plant-based diet (V, blend of plant proteins and vegetable oils). Livers, viscera, carcasses and ova were sampled at spawning and analyzed for lipids and fatty acids. Although the V-diet was devoid of n-3 LC-PUFAs, significant amounts of EPA and DHA were found in livers and ova, demonstrating efficient bioconversion of linolenic acid and selective orientation towards the ova. Some ova were fertilized to assess the reproductive performance and offspring survival. We observed for the first time that trout fed a 100% plant-based diet over a 3-year breeding cycle were able to produce ova and viable alevins, although the ova were smaller. The survival of offspring from V-fed females was lower (-22%) at first spawning, but not at the second. Our study showed that, in addition to being able to grow on a plant-based diet, rainbow trout reared entirely on such a diet can successfully produce ova in which neo-synthesized n-3 LC-PUFAs are accumulated, leading to viable offspring. However, further adjustment of the feed formula is still needed to optimize reproductive performance.

Dans un contexte de demande croissante en protéines animales, l’élevage de précision et la sélection animale doivent viser à accroître la compétitivité des systèmes de production et à réduire leur impact environnemental. Les animaux doivent être plus efficaces pour transformer leur ration (réduire l’ingéré tout en maintenant une production stable), en acceptant des ressources alimentaires variées (protéines ou fibres) et en minimisant leurs rejets (totaux ou leurs composantes N et P, GES). Toutefois, l’efficacité alimentaire est un phénotype complexe qui doit être phénotypé finement pour comprendre les processus biologiques sous-jacents avant d’identifier les leviers d’action pour l’améliorer (efficacité digestive vs métabolique ; efficacité énergétique vs protéique). Ce phénotypage doit être réalisé en conditions variées tant sur le plan du génotype, du niveau alimentaire que de la composition de la ration (teneur en fibres, en acides aminés). Il s’agit aussi d’envisager une approche prédictive de l’ingéré et de l’efficacité alimentaire basée sur des paramètres peu coûteux et faciles à mesurer (croissance, bio-marqueurs…) dans le cadre d’un élevage de précision ou de programmes de sélection génétique, car le phénotypage à large échelle de l’ingéré reste coûteux et parfois difficile à mettre en oeuvre. Pour évaluer l’efficience globale des systèmes de productions animales, il s’agit de considérer les réponses des animaux pour des génotypes et des régimes alimentaires contrastés en termes d’efficacité alimentaire et de rejets en quantité et en qualité (GES, N, P), mais aussi de robustesse des animaux (comportement alimentaire, activité générale, santé)., In a context of an increasing demand for animal proteins, precision farming and animal breeding must be goal-oriented to improve production competitiveness and to reduce environmental impact. Animals have to be more efficient in transforming their diet (reducing their feed intake while maintaining production performance), accepting various feeds (protein and fibers) and reducing their wastes (total, N, P, GHG…). However, feed efficiency is a complex phenotype that needs to be deeply phenotyped. This will allow unravelling the biological processes to identify ways to improve feed efficiency (digestive versus metabolic efficiency; energetic versus protein efficiency) according to the genotype, the feed level and the diet composition (fibre content, AA content). Predictive biology of feed intake and feed efficiency is also needed using cost-effective and easy to record traits (growth, biomarkers…) for precision farming and for selective breeding, because large-scale phenotyping for feed intake remains costly. To assess the efficiency of livestock production systems, there is a need to reconsider animal responses according to contrasted genotypes and feeds not only for feed efficiency and wastes in quantity and quality (GHG, N, P), but also for robustness of animals (feeding behavior, general activity, health).

Chantier qualité GA; International audience; Sea bass is a major species in Mediterranean aquaculture, but has a distribution area ranging from North Atlantic to South Mediterranean, with a population structure previously revealed by population genetics. To test the farming performances of wild sea bass populations, we produced a partial diallel cross mating scheme, using sires originating from North Atlantic (NAT), South Atlantic (SAT), West Mediterranean (WEM), North-East Mediterranean (NEM) and South-East Mediterranean (SEM). Fifteen sires per origin were mated in a full-factorial design using artificial fertilization with 9 NAT dams and 17 WEM dams, producing 10 population crosses and 1950 potential full-sib families. All fish were reared together, then tagged at an average weight of 20 g and distributed to four different sites (1800 fish per site). They were grown to an objective of 200 g mean weight, where 737 to 775 fish were slaughtered in each site, and their parentage was recovered using 6 to 7 microsatellite loci, resulting in 98.9% unique assignments. All populations had similar growth rates until tagging size (20 g), but differences appeared later on. No heterosis appeared for growth rate, and genotype by environment interaction (G × E) at the population level was limited, with a significant re-ranking only in one rearing site, while strong G × E for growth rate was observed within populations. Populations were different in shape, muscular fat content, and carcass yield, but not in fillet yield. In general, heterosis was absent and G × E was very limited between populations. No "ideal" population combining all favorable traits was identified. Differences between extreme populations ranged between 3 and 49% of the mean, depending on the traits. Interestingly, in almost all cases, these differences were within the reach of one generation of intense (5%) phenotypic selection.

The study of inter-individual variability of personality in fish is a growing field of interest but the genetic basis of this complex trait is still poorly investigated due to the difficulty in controlling fish genetic origin and life history. When available, isogenic lines that allow performing independent tests on different individuals having identical genotype constitute a very relevant experimental material to disentangle the genetic and environmental components of behavioural individuality. We took advantage of heterozygous isogenic lines to investigate the personality in rainbow trout through the analysis of their reactions to different experimental situations. To this end, seven to ten rainbow trout isogenic lines were screened for their spatial exploratory behaviour, their flight response toward a stressor and their risk taking behaviour. Results showed that some lines seemed less sensitive to new events or environmental changes and could be defined as low responsive, while others were very sensitive and defined as high responsive. The use of isogenic lines highlighted the importance of genetic factors, in combination with life history, in the expression of personality in domesticated fish.

The evolution of genetic variability is studied in six successive generations of a population originating from wild Helix aspersa. During the first three generations (G1 to G3), no artificial selection was applied. During the next three generations (G4 to G6), two lines were reared: a control line (C) and a line (S) selected for increased adult weight. Genetic variability is described by genealogical parameters (inbreeding, number of founders, effective number of founders and ancestors, effective number of remaining genomes) and by the additive genetic variance in adult weight. A large decrease in all parameters was observed between G1 and G2, suggesting strong natural selection: additive genetic variance in adult weight (transformed data) decreased from 0.0119 +/- 3.8 x 10(-3) to 0.0070 +/- 1.7 x 10(-3) (P0.05) and effective number of ancestors from 97.4 to 67.0. Selection also caused a large decrease during the first generation: additive genetic variance was 0.0079 +/- 2.1 x 10(-3) in G3 and 0.0040 +/- 1.1 x 10(-3) after the first selection cycle (P0.02). At the same time, the effective number of ancestors decreased from 59.2 to 29.5 and 24.2. This decrease is consistent with the theory of selection and the Bulmer effect.

In ¢sh, sexual maturation is often considered to be a problem because it perturbs growth and product quality. Therefore, it is common to select against early-maturing males in commercial breeding programmes (Gjedrem 2000). Genetic determinism of age at sexual maturation has been extensively studied in many species, and even some QTLs have been found (e.g. Gjerde 1986; Hankin, Nicholas & Downey 1993; Longalong, Eknath & Bentsen 1999; Kause, Ritola, Paananen, Mantysaari & Eskelinen 2003; Haidle, Janssen, Gharbi, Moghadam, Ferguson & Danzmann 2008). However, the eiect of sexual maturation on heritability estimates for other traits is not documented. In this paper, we present data collected during the beginningof sexualmaturationof a group of rainbow trout and showhow it biases heritability estimations. The ¢sh studied were issued froma full-factorial mating between two dams and 45 sires. Fish were all reared in the same tank since the eyed stage under a natural photoperiod and pedigrees were redrawn using10 microsatellites. Fishwere harvested in April at 17 months of age. They were killed on ice and several growth and quality traits were determined. Body weight was the main trait studied in this paper. The sex of the ¢sh was recorded by visual inspection of the gonads. For females, none of the individuals showed signs of maturation. For males, observers attempted to diierentiate non-maturing and maturing males but it turned out to be di⁄cult because we were at the very beginning of the maturation (this rainbow trout strain usually spawns in November^ December). However, we recorded gonads weight to enable calculation of the gonado-somatic index (GSI5100 gonads weight/body weight), which was used to determine which males were maturing or not.We studied several GSI thresholds: 0.1, 0.2, 0.3, 0.4 and 0.5. For each threshold, males with GSI above the threshold were considered to be maturing males. We studied sex eiect and heritability estimates for each threshold. Heritability was estimated using VCE5 (Groeneveld & Kovac 1990) with a sire model and sex and dam as ¢xed eiects. Dam was set as a ¢xed eiect because there were only two dams in our mating design, and therefore between-dams variance is of no interest. Two dams were used instead of one in order to avoid confusion of dominance eiects with additive genetic eiects in the between-sires variance. Moreover, this design was demonstrated to be e⁄cient for estimating heritabilities when the total number of oispring analysed is ¢xed (Dupont-Nivet, Vandeputte & Chevassus 2002). First, we analysed all the datasets with a sex eiect comprising three levels (female, male and maturing male). Second, maturing males were removed from the dataset and the sex effect was set to two levels (male and female). Using ‘FAP’ software (Taggart 2007), 87% of ¢sh could be attributed to their parents. Oispring from two sires with less than three oispring were removed from the data set. Four hundred and thirty-three animals were ¢nally analysed. The sex ratio was well equilibrated with 217 females, i.e., 50.1%. In Table 1, Aquaculture Research, 2010, 41, e878^e880 doi:10.1111/j.1365-2109.2009.02448.x