BREEDING AND GENETICS SYMPOSIUM: Climate change and selective breeding in aquaculture.

Aquaculture is the fastest growing food production sector and it contributes significantly to global food security. Based on Food and Agriculture Organization (FAO) of the United Nations, aquaculture production must increase significantly to meet the future global demand for aquatic foods in 2050. According to Intergovernmental Panel on Climate Change (IPCC) and FAO, climate change may result in global warming, sea level rise, changes of ocean productivity, freshwater shortage, and more frequent extreme climate events. Consequently, climate change may affect aquaculture to various extents depending on climatic zones, geographical areas, rearing systems, and species farmed. There are 2 major challenges for aquaculture caused by climate change. First, the current fish, adapted to the prevailing environmental conditions, may be suboptimal under future conditions. Fish species are often poikilothermic and, therefore, may be particularly vulnerable to temperature changes. This will make low sensitivity to temperature more important for fish than for livestock and other terrestrial species. Second, climate change may facilitate outbreaks of existing and new pathogens or parasites. To cope with the challenges above, 3 major adaptive strategies are identified. First, general 'robustness' will become a key trait in aquaculture, whereby fish will be less vulnerable to current and new diseases while at the same time thriving in a wider range of temperatures. Second, aquaculture activities, such as input power, transport, and feed production contribute to greenhouse gas emissions. Selection for feed efficiency as well as defining a breeding goal that minimizes greenhouse gas emissions will reduce impacts of aquaculture on climate change. Finally, the limited adoption of breeding programs in aquaculture is a major concern. This implies inefficient use of resources for feed, water, and land. Consequently, the carbon footprint per kg fish produced is greater than when fish from breeding programs would be more heavily used. Aquaculture should use genetically improved and robust organisms not suffering from inbreeding depression. This will require using fish from well-managed selective breeding programs with proper inbreeding control and breeding goals. Policymakers and breeding organizations should provide incentives to boost selective breeding programs in aquaculture for more robust fish tolerating climatic change.

Genetic parameters for carcass weight, conformation and fat in five beef cattle breeds.

Profitability of beef production can be increased by genetically improving carcass traits. To construct breeding value evaluations for carcass traits, breed-specific genetic parameters were estimated for carcass weight, carcass conformation and carcass fat in five beef cattle breeds in Finland (Hereford, Aberdeen Angus, Simmental, Charolais and Limousin). Conformation and fat were visually scored using the EUROP carcass classification. Each breed was separately analyzed using a multitrait animal model. A total of 6879-19 539 animals per breed had phenotypes. For the five breeds, heritabilities were moderate for carcass weight (h 2=0.39 to 0.48, s.e.=0.02 to 0.04) and slightly lower for conformation (h 2=0.30 to 0.44, s.e.=0.02 to 0.04) and carcass fat (h 2=0.29 to 0.44, s.e.=0.02 to 0.04). The genetic correlation between carcass weight and conformation was favorable in all breeds (r G=0.37 to 0.53, s.e.=0.04 to 0.05), heavy carcasses being genetically more conformed. The phenotypic correlation between carcass weight and carcass fat was moderately positive in all breeds (r P=0.21 to 0.32), implying that increasing carcass weight was related to increasing fat levels. The respective genetic correlation was the strongest in Hereford (r G=0.28, s.e.=0.05) and Angus (r G=0.15, s.e.=0.05), the two small body-sized British breeds with the lowest conformation and the highest fat level. The correlation was weaker in the other breeds (r G=0.08 to 0.14). For Hereford, Angus and Simmental, more conformed carcasses were phenotypically fatter (r P=0.11 to 0.15), but the respective genetic correlations were close to zero (r G=-0.05 to 0.04). In contrast, in the two large body-sized and muscular French breeds, the genetic correlation between conformation and fat was negative and the phenotypic correlation was close to zero or negative (Charolais: r G=-0.18, s.e.=0.06, r P=0.02; Limousin: r G=-0.56, s.e.=0.04, r P=-0.13). The results indicate genetic variation for the genetic improvement of the carcass traits, favorable correlations for the simultaneous improvement of carcass weight and conformation in all breeds, and breed differences in the correlations of carcass fat.

Genotype-by-environment interaction of growth traits in rainbow trout (Oncorhynchus mykiss): a continental scale study.

Rainbow trout is a globally important fish species for aquaculture. However, fish for most farms worldwide are produced by only a few breeding companies. Selection based solely on fish performance recorded at a nucleus may lead to lower-than-expected genetic gains in other production environments when genotype-by-environment (G × E) interaction exists. The aim was to quantify the magnitude of G × E interaction of growth traits (tagging weight; BWT, harvest weight; BWH, and growth rate; TGC) measured across 4 environments, located in 3 different continents, by estimating genetic correlations between environments. A total of 100 families, of at least 25 in size, were produced from the mating 58 sires and 100 dams. In total, 13,806 offspring were reared at the nucleus (selection environment) in Washington State (NUC) and in 3 other environments: a recirculating aquaculture system in Freshwater Institute (FI), West Virginia; a high-altitude farm in Peru (PE), and a cold-water farm in Germany (GER). To account for selection bias due to selective mortality, a multitrait multienvironment animal mixed model was applied to analyze the performance data in different environments as different traits. Genetic correlation (rg) of a trait measured in different environments and rg of different traits measured in different environments were estimated. The results show that heterogeneity of additive genetic variances was mainly found for BWH measured in FI and PE. Additive genetic coefficient of variation for BWH in NUC, FI, PE, and GER were 7.63, 8.36, 8.64, and 9.75, respectively. Genetic correlations between the same trait in different environments were low, indicating strong reranking (BWT: rg = 0.15 to 0.37, BWH: rg = 0.19 to 0.48, TGC: rg = 0.31 to 0.36) across environments. The rg between BWT in NUC and BWH in both FI (0.31) and GER (0.36) were positive, which was also found between BWT in NUC and TGC in both FI (0.10) and GER (0.20). However, rg were negative between BWT in NUC and both BWH (-0.06) and TGC (-0.20) in PE. Correction for selection bias resulted in higher additive genetic variances. In conclusion, strong G × E interaction was found for BWT, BWH, and TGC. Accounting for G × E interaction in the breeding program, either by using sib information from testing stations or environment-specific breeding programs, would increase genetic gains for environments that differ significantly from NUC.

Accounting for early rearing density effects on growth in the genetic evaluation of rainbow trout, Oncorhynchus mykiss.

In fish breeding, full-sib families are often kept in separate tanks until individuals are large enough to be tagged and pooled. This practice induces substantial environmental variation common to full sibs (VFS) in BW. We used multigeneration data on rainbow trout to investigate how variation among families in early rearing density affects BW at different ages and environments (fresh water and sea), and whether variance parameters and ranking of breeding candidates change when density is either excluded or included as a regression term in a multitrait animal model. Increasing density displayed a consistent negative relationship with full-sib tank-mean BW at the end of fry-stage when family sizes were equalized (r2 of linear regressions 11 to 31%). In 4 out of 6 year classes, the significant negative association between density and BW also remained through the family-tank period until tagging at 6 mo of age (r2=3 to 19%). In some year classes, early density had a carry-over effect on later BW means, reaching up to the age of >2 yr (after the second and third growing season). Yet, the association was generally weaker at later ages and varied from nonexistent to both significantly negative and positive (r2=0 to 6%). For each BW, the inclusion of early density in genetic model primarily captured the variance that was otherwise attributable to VFS. The reduction of VFS was most pronounced in tagging BW (21% difference between the models), where common environmental effects were moderately high (c2=0.18 in the model without density). For later BW traits, the difference in VFS was 6 to 8% (c2=0.04 to 0.05). The changes in genetic, residual, and phenotypic variances were generally small in the model, including density. Similarly, only a slight change in the heritability estimate of any BW was found (differences of h2 0.2 to 1.3% between models). Correlations between EBV obtained by the 2 models were highly positive in each BW trait (r range 0.94 to 1.00), indicating that ranking of the breeding candidates remained consistent, regardless of whether density was accounted for or not. Our findings highlight the importance of equalizing family sizes to decrease VFS in rainbow trout growth. If selection is practiced for EBV of BW recorded at an age of >2 yr, and equalization is done early enough, the effects induced by different initial density can be sufficiently corrected for by the common full-sib effect in models used for genetic evaluation.

Merino ewes can be bred for body weight change to be more tolerant to uncertain feed supply.

Sheep in Australia experience periods with different feed supply causing them to gain and lose BW during the year. It is more efficient if ewes lose less BW during periods of poor nutrition and gain more BW during periods of good nutrition. We investigated whether BW loss during periods of poor nutrition and BW gain during periods of good nutrition are genetically different traits. We used BW measurements from 2,336 adult Merino ewes managed over 5 yr in a Mediterranean climate in Katanning, Australia. Body weight loss is the difference between 2 BW measured 42 d apart during mating, a period of poor nutrition. Body weight gain is the difference between 2 BW measured 131 d apart during a period of good nutrition between prelambing and weaning. We estimated variance compnents of BW change using 3 methods: 1) as a trait calculated by subtracting the first BW from the second, 2) multivariate analysis of BW traits, and 3) random regression analysis of BW. The h(2) and genetic correlations (rg) estimated using the multivariate analysis of BW and the BW change trait were very similar whereas the random regression analysis estimated lower heritabilities and more extreme negative genetic correlations between BW loss and gain. The multivariate model fitted the data better than random regression based on Akaike and Bayesian information criterion so we considered the results of the multivariate model to be more reliable. The heritability of BW loss (h(2) = 0.05-0.16) was smaller than that of BW gain (h(2) = 0.14-0.37). Body weight loss and gain can be bred for independently at 2 and 4 yr of age (rg = 0.03 and -0.04) whereas at 3 yr of age ewes that genetically lost more BW gained more BW (rg = -0.41). Body weight loss is genetically not the same trait at different ages (rg range 0.13-0.39). Body weight gain at age 3 yr is genetically the same trait at age 4 yr (rg = 0.99) but is different between age 2 yr and the older ages (rg = 0.53 and 0.51). These results suggest that as the ewes reach their mature BW, BW gain at different ages becomes the same trait. This does not apply to BW loss. We conclude that BW change could be included in breeding programs to breed adult Merino ewes that are more tolerant to variation in feed supply.

Heat stress effects on farrowing rate in sows: genetic parameter estimation using within-line and crossbred models.

The pork supply chain values steady and undisturbed piglet production. Fertilization and maintaining gestation in warm and hot climates is a challenge that can be potentially improved by selection. The objective of this study was to estimate 1) genetic variation for farrowing rate of sows in 2 dam lines and their reciprocal cross; 2) genetic variation for farrowing rate heat tolerance, which can be defined as the random regression slope of farrowing rate against increasing temperature at day of insemination, and the genetic correlation between farrowing rate and heat tolerance; 3) genetic correlation between farrowing rate in purebreds and crossbreds; and 4) genetic correlation between heat tolerance in purebreds and crossbreds. The estimates were based on 93,969 first insemination records per cycle from 24,456 sows inseminated between January 2003 and July 2008. These sows originated from a Dutch purebred Yorkshire dam line (D), an International purebred Large White dam line (ILW), and from their reciprocal crosses (RC) raised in Spain and Portugal. Within-line and crossbred models were used for variance component estimation. Heritability estimates for farrowing rate were 0.06, 0.07, and 0.02 using within-line models for D, ILW, and RC, respectively, and 0.07, 0.07, and 0.10 using the crossbred model, respectively. For farrowing rate, purebred-crossbred genetic correlations were 0.57 between D and RC and 0.50 between ILW and RC. When including heat tolerance in the within-line model, heritability estimates for farrowing rate were 0.05, 0.08, and 0.03 for D, ILW, and RC, respectively. Heritability for heat tolerance at 29.3°C was 0.04, 0.02, and 0.05 for D, ILW, and RC, respectively. Genetic correlations between farrowing rate and heat tolerance tended to be negative in crossbreds and ILW-line sows, implying selection for increased levels of production traits, such as growth and reproductive output, is likely to increase environmental sensitivity. This study shows that genetic selection for farrowing rate and heat tolerance is possible. However, when this selection is based solely on purebred information, the expected genetic progress on farrowing rate and heat tolerance in crossbreds (commercial animals) would be inconsequential.

Defining desired genetic gains for rainbow trout breeding objective using analytic hierarchy process.

Distributing animals from a single breeding program to a global market may not satisfy all producers, as they may differ in market objectives and farming environments. Analytic hierarchy process (AHP) is used to estimate preferences, which can be aggregated to consensus preference values using weighted goal programming (WGP). The aim of this study was to use an AHP-WGP based approach to derive desired genetic gains for rainbow trout breeding and to study whether breeding trait preferences vary depending on commercial products and farming environments. Two questionnaires were sent out. Questionnaire-A (Q-A) was distributed to 178 farmers from 5 continents and used to collect information on commercial products and farming environments. In this questionnaire, farmers were asked to rank the 6 most important traits for genetic improvement from a list of 13 traits. Questionnaire B (Q-B) was sent to all farmers who responded to Q-A (53 in total). For Q-B, preferences of the 6 traits were obtained using pairwise comparison. Preference intensity was given to quantify (in % of a trait mean; G%) the degree to which 1 trait is preferred over the other. Individual preferences, social preferences, and consensus preferences (Con-P) were estimated using AHP and WGP. Desired gains were constructed by multiplying Con-P by G%. The analysis revealed that the 6 most important traits were thermal growth coefficient (TGC), survival (Surv), feed conversion ratio (FCR), condition factor (CF), fillet percentage (FIL%), and late maturation (LMat). Ranking of traits based on average Con-P values were Surv (0.271), FCR (0.246), TGC (0.246), LMat (0.090), FIL% (0.081), and CF (0.067). Corresponding desired genetic gains (in % of trait mean) were 1.63, 1.87, 1.67, 1.29, 0.06, and 0.33%, respectively. The results from Con-P values show that trait preferences may vary for different types of commercial production or farming environments. This study demonstrated that combination of AHP and WGP can be used to derive desired gains for a breeding program and to quantify differences due to variations market demand or production environment.

Genetic parameters for feed utilization traits in Japanese quail.

Feed costs substantially affect the efficiency of poultry operations, justifying genetic improvement of feed utilization by selection. The current research was conducted to estimate genetic variance for the 4-wk feed conversion ratio (FCR) and its genetic correlations with BW, BW gain (WG), feed intake (FI), and residual feed intake (RFI) in Japanese quail. The data analyzed originated from a line selected for low FCR for 3 generations. In each generation, 35 sires and 70 dams were used as parents for the next generation. Body weight and WG were recorded on a total of 1,226 individuals, whereas FCR, RFI, and FI were recorded on 505 family groups. The results showed that heritability estimates (±SE) of BW at 28 d of age and WG between 7 and 28 d of age were 0.22 ± 0.05 and 0.28 ± 0.06, respectively. For FI, FCR, and RFI, significant genetic variances were estimated. Genetic correlations of FCR between 7 and 28 d of age with WG and FI between 7 and 28 d of age were -0.45 ± 0.09 and 0.24 ± 0.08, respectively. This implies that a low FCR is genetically related to a high WG and low FI. The genetic correlation between FCR from 7 to 28 d of age and RFI from 7 to 28 d of age was 0.26 ± 0.08, indicating that the 2 alternative feed efficiency traits are genetically different traits, and that the correlated genetic response in one of them in response to selection on the other is likely to be only moderate. Genetic correlations of RFI from 7 to 28 d of age with WG and FI between 7 and 28 d of age were 0.08 ± 0.04 and 0.74 ± 0.11, respectively. This reflects the fact that RFI is phenotypically independent of WG, which tends to make the genetic correlation between RFI and WG low as well. In conclusion, all the traits analyzed displayed significant genetic variance, allowing their genetic improvement by selection, yet the alternative feed utilization traits, FCR and RFI, displayed different genetic characteristics.

Quality and production trait genetics of farmed European whitefish, Coregonus lavaretus.

We present here phenotypic and genetic parameters for the major quality and production traits of farmed European whitefish. A total of 70 families were produced by mating each of 45 sires to an average of 1.6 dams and each of the 52 dams to an average of 1.3 sires. A total of 2,100 individuals were recorded for survival, and 507 individuals for growth and quality-related traits. The 4 major results were as follows: first, all traits exhibited nonzero heritabilities except for fillet gaping and fillet protein%. The heritabilities for the production traits were harvest weight (0.42 ± 0.10), gutted weight (0.40 ± 0.10), fillet weight (0.36 ± 0.09), maturity score (0.27 ± 0.11, on liability scale), survival (0.19 ± 0.05, on liability scale), carcass% (0.14 ± 0.07), and fillet% (0.11 ± 0.06). The heritabilities for the quality traits were condition factor (0.49 ± 0.10), fillet lipid% (0.37 ± 0.10), muscle texture (0.30 ± 0.09), Distell lipid reading (0.26 ± 0.09), fillet lightness (0.16 ± 0.07), fillet gaping (0.04 ± 0.06), and fillet protein% (0.04 ± 0.06). Second, the quality traits that were significantly genetically correlated with each other were all related to lipid deposition. Increasing fillet lipid% (an undesired change in whitefish) was genetically related to desired lighter fillet color [genetic correlation (r(G)) = 0.70 ± 0.22] and to undesired greater condition factor (0.39 ± 0.17). None of the other genetic correlations between condition factor, fillet lipid%, muscle texture, fillet lightness, fillet gaping, and fillet protein% were significant. Third, BW and gutted weight were genetically related to the quality traits that were genetically related to lipid deposition. Increasing harvest weight was genetically related to high fillet lipid% (r(G) = 0.59 ± 0.14), lighter fillet color (0.61 ± 0.25), and to greater condition factor (0.60 ± 0.12). All other genetic correlations of harvest weights with the quality traits were nonsignificant, indicating that rapid growth was not genetically related to gaping and softer flesh. Fourth, none of the genetic correlations of carcass%, fillet%, maturity, and survival with the quality traits were significant, implying weak genetic integration between the traits. Yet, marginally significant genetic correlations were found for fillet lipid% with maturity score (r(G) = -0.46 ± 0.24) and survival (0.36 ± 0.19). These results provide the genetic basis for assessing the potential to improve product quality via selective breeding.

Response to selection for feed conversion ratio in Japanese quail.

We investigated the effect of selection for 4-wk feed conversion ratio (FCR) on genetic improvement of FCR, BW, weight gain (WG), feed intake (FI), and residual FI (RFI) in Japanese quail. The F line was selected for reduced FCR and the C line was maintained as a randombred control. In each generation, 35 sires and 70 dams were used as parents for the next generation. Three generations of selection were performed. Realized heritability for FCR was calculated as the ratio of cumulative selection response to the cumulative selection differential, and additionally, genetic response was quantified as the difference between the means of selection and control lines. The results showed that realized heritability for FCR after 3 generations of selection was 0.67. The mean FCR in F line and C line in the last generation was 2.13 and 2.61, respectively. This is 18.4% cumulative genetic improvement, or 6.1% improvement per generation. In the last generation, the means of F and C lines were 193 and 166 g for BW at age 28 d (16.4% total increase, or 5.5% per generation), 184 and 158 g for WG (17.2% total higher gain and 5.7% per generation), 393 and 413 g for FI (4.9% total higher consumption and 1.6% per generation), and -24.5 and 10.2 for RFI (-34.7 g of cumulative gain; -11.6 g per generation), respectively. These results show that selection to decrease FCR increases BW and WG and decreases FI and RFI as a correlated response.

Quantitative genetic architecture of parasite-induced cataract in rainbow trout, Oncorhynchus mykiss.

Parasites impose costs on their hosts. The capability to fight against them is of great advantage, but may also be traded off with other traits. Although often observed at the phenotypic level, our knowledge of the extent to which such trade-offs are genetically determined is relatively poor. We tested this possibility with a farmed rainbow trout population suffering from natural Diplostomum spp. infections that cause cataracts in fish. We estimated the heritability of cataract severity and examined phenotypic and genetic correlations between cataract and a set of performance traits measured three times during a 3-year rearing period. A cataract score was used as an indicator of the host's capability to resist and/or tolerate the parasite. Our results showed moderate heritability for the cataract. Nevertheless, we found no evidence for a genetic or phenotypic trade-off between parasite resistance/tolerance and the measured performance traits. Initial body weight was not correlated with the cataract score. Phenotypic and genetic correlations of cataract severity with body mass and condition measured in the second and third year were strongly negative, indicating reduced growth and condition in fish with a high cataract score. The reduced body size and condition in cataract-bearing fish were probably reflected in the phenotypic association between a high cataract score and delayed maturity age in females. Put together, our study did not provide evidence of genetic or phenotypic trade-offs between Diplostomum resistance/tolerance and a number of performance traits. Therefore, selection for lessened Diplostomum-caused cataracts is unlikely to have a negative impact on the studied performance traits.

Direct and indirect selection of visceral lipid weight, fillet weight, and fillet percentage in a rainbow trout breeding program.

We assessed whether visceral lipid weight, fillet weight, and percentage fillet from BW, 3 traits laborious to record, could be genetically improved by indirect selection on more easily measured traits in farmed rainbow trout. Visceral lipid is discarded as waste during slaughter, influencing production efficiency and production costs. Fillet weight and fillet percentage directly influence economic returns in trout production. The study comprised 3 steps. First, we assessed the degree to which selection on percentage of visceral weight from BW indirectly changes visceral lipid weight and the size of intestines and internal organs. The phenotypic analysis of weights of viscera, intestines, visceral lipid, liver, and gonads measured from 40 fish revealed that phenotypic selection against visceral weight was most strongly directed to visceral lipid, and to a lesser degree to intestines and gonads. Because genetic relationships among these traits were not established, it is not known whether indirect selection leads to genetic responses. Second, we examined whether direct selection for the fillet traits could be replaced by indirect selection on BW, eviscerated BW, visceral weight, visceral percentage, head volume, and relative head volume (head volume relative to BW). The selection index calculations based on the quantitative genetic parameters obtained from multigenerational pedigree data showed that genetic improvement of fillet percentage through direct selection (selection accuracy, r(TI) = 0.54) was equally efficient compared with indirect selection on visceral percentage ( r(TI) = 0.54). Genetic improvement of fillet weight through direct selection (r(TI) = 0.56) was always more efficient than indirect selection, yet indirect selection for eviscerated BW ( r(TI) = 0.50) was almost as efficient as direct selection. Third, the expected genetic responses to alternative selection indices showed that improved fillet percentage was mainly a result of a moderate decrease in visceral weight rather than of a major increase in absolute fillet weight. Moreover, fillet percentage is challenging to improve, even if it exhibits moderate heritability (h(2) = 0.29). This is because fillet percentage displays low phenotypic variation. In conclusion, fillet weight and fillet percentage can be increased by indirect selection against visceral percentage and for high eviscerated BW.

Genetic relationships of body composition and feed utilization traits in European whitefish (Coregonus lavaretus L.) and implications for selective breeding in fishmeal- and soybean meal-based diet environments.

Body composition traits have potential use in fish breeding programs as indicator traits for selective improvement of feed efficiency. Moreover, feed companies are increasingly replacing traditional fish meal (FM) based ingredients in feeds for carnivorous farmed fish with plant protein ingredients. Therefore, genetic relationships of composition and feed utilization traits need to be quantified for both current FM-based and future plant-based aquaculture feeds. Individual whole-body lipid% and protein%, daily gain (DG), ADFI, and G:F (daily gain/daily feed intake) were measured on 1,505 European whitefish (Coregonus lavaretus) from 70 half/full-sib families reared in a split-family design with either a typical FM or a novel soybean meal (SBM) based diet. Diet-specific genetic parameters were estimated with multiple-trait animal models. Lipid% was significantly greater in the FM diet group than in the SBM group, even independent of final BW or total feed intake. In both diets, lipid% showed moderate heritability (0.12 to 0.22) and had positive phenotypic and genetic correlations with DG (0.37 to 0.82) and ADFI (0.36 to 0.88). Therefore, selection against lipid% can be used to indirectly select for lower feed intake. Protein% showed low heritability (0.05 to 0.07), and generally very weak or zero correlations with DG and ADFI. In contrast to many previous studies on terrestrial livestock, lipid% showed zero or very weak phenotypic and genetic correlations with G:F. However, selection index calculations demonstrated that simultaneous selection for high DG and reduced lipid% could be used to indirectly increase G:F; this strategy increased absolute genetic response in G:F by a factor of 1.5 to 1.6 compared with selection on DG alone. Lipid% and protein% were not greatly affected by genotype-diet environment interactions, and therefore, selection strategies for improving body composition within current FM diets should also improve populations for future SBM diets.

Feed efficiency of rainbow trout can be improved through selection: different genetic potential on alternative diets.

To assess the genetic potential for selection of increased feed efficiency in rainbow trout (Oncorhynchus mykiss), we estimated the heritabilities and correlations for BW, daily weight gain (DG), and daily feed intake (DFI). Body weight was recorded 5 times, and DG and DFI 3 times during a feeding trial lasting 22 mo. To test the hypothesis that phenotypic and genetic parameters were influenced by a nutritional environment, fish were fed either a modern normal protein diet (NP, 40 to 45% protein and 30 to 33% lipid) or an alternative high protein diet (HP, 50 to 56% protein, 20 to 24% lipid) in a split-family design. Results showed that there were no large differences in heritabilities between the diets. Average heritability for DFI over both diets and different fish ages was low (average h2 = 0.10), indicating that modest genetic changes in response to selection can be obtained. Average heritabilities for BW and DG over both diets and different fish ages were 0.28 and 0.33, respectively. The NP diet enabled fish to express a wide range of BW, as shown by the increased coefficients of phenotypic variation for BW. Fish fed the HP diet showed increased phenotypic variation for DFI in > 750-g fish. On the NP diet, genetic correlations of DFI with DG and BW were very strong for 750- to 2,000-g fish. In contrast, on the HP diet, the respective correlations were moderate to low, revealing more genetic potential to change growth and feed intake simultaneously in opposite directions. An analysis of the predicted selection responses showed that selection solely for high DG improved feed efficiency as a correlated genetic response. Simultaneous selection for high DG and reduced DFI, in turn, may increase genetic gain in feed efficiency by a factor of 1.2 compared with selection solely for DG. However, variation for growth and feed intake and the relationships between these traits were different in different nutritional environments, leading to divergent genetic responses on the alternative diets.

Genetic associations of prolificacy with performance, carcass, meat quality, and leg conformation traits in the Finnish Landrace and Large White pig populations.

The objective of this study was to estimate genetic associations of prolificacy traits with other traits under selection in the Finnish Landrace and Large White populations. The prolificacy traits evaluated were total number of piglets born, number of stillborn piglets, piglet mortality during suckling, age at first farrowing, and first farrowing interval. Genetic correlations were estimated with two performance traits (ADG and feed:gain ratio), with two carcass traits (lean percent and fat percent), with four meat quality traits (pH and L* values in longissimus dorsi and semimembranosus muscles), and with two leg conformation traits (overall leg action and buck-kneed forelegs). The data contained prolificacy information on 12,525 and 10,511 sows in the Finnish litter recording scheme and station testing records on 10,372 and 9,838 pigs in Landrace and Large White breeds, respectively. The genetic correlations were estimated by the restricted maximum likelihood method. The most substantial correlations were found between age at first farrowing and lean percent (0.19 in Landrace and 0.27 in Large White), and fat percent (-0.26 in Landrace and -0.18 in Large White), and between number of stillborn piglets and ADG (-0.38 in Landrace and -0.25 in Large White) and feed:gain (0.27 in Landrace and 0.12 in Large White). The correlations are indicative of the benefits of superior growth for piglets already at birth. Similarly, the correlations indicate that age at first farrowing is increasing owing to selection for carcass lean content. There was also clear favorable correlation between performance traits and piglet mortality from birth to weaning in Large White (r(g) was -0.43 between piglet mortality and ADG, and 0.42 between piglet mortality and feed:gain), but not in Landrace (corresponding correlations were 0.26 and -0.22). There was a general tendency that prolificacy traits were favorably correlated with performance traits, and unfavorably with carcass lean and fat percents, whereas there were no clear associations between prolificacy and meat quality or leg conformation. In conclusion, accuracy of estimated breeding values may be improved by accounting for genetic associations between prolificacy, carcass, and performance traits in a multitrait analysis.

Seasonality and genetic architecture of development time and body size of the birch feeding sawfly Priophorus pallipes.

We tested, using the sawfly Priophorus pallipes feeding on leaves of mountain birch, whether the expression of genetic (co)variation of larval development time and body size can be altered by exposing larvae to diets with differential seasonal changes in quality. In nature, larvae feed mainly on mature leaves, but occasionally they are forced to consume senescing leaves. Sixty families were assayed on three experimentally simulated diets: mature leaves of high quality, senescing leaves of rapidly declining quality, and senesced leaves of low quality. The intuitively obvious positive phenotypic and genetic correlations between development time and final mass were observed when the larvae consumed leaves of stable high quality, but low and declining food quality prevented long-growing individuals and families from achieving high final mass, switching the correlations to close to zero or negative in these treatments. The amount of genetic variation for body size showed a non-linear change across the diet quality gradient, whereas genetic variation for development time increased with decreasing diet quality. The among-trait difference in the degree reaction norms crossed along the diet gradient caused the changes in the expression of genetic (co)variation within the environments. Our results show that seasonally varying diet quality induces dramatic changes in the genetic (co)variation of development time and body size, and that simultaneous analysis of reaction norms and environment-specific expression of genetic (co)variation is necessary for the understanding of the genetic characteristics underlying the construction of phenotypes in heterogeneous environments.

Seasonally varying diet quality and the quantitative genetics of development time and body size in birch feeding insects.

Genetic variance-covariance structures (G), describing genetic constraints on microevolutionary changes of populations, have a central role in the current theories of life-history evolution. However, the evolution of Gs in natural environments has been poorly documented. Resource quality and quantity for many animals and plants vary seasonally, which may shape genetic architectures of their life histories. In the mountain birch-insect herbivore community, leaf quality of birch for insect herbivores declines profoundly during both leaf growth and senescence, but remains stable during midsummer. Using six sawfly species specialized on the mountain birch foliage, we tested the ways in which the seasonal variation in foliage quality of birch is related to the genetic architectures of larval development time and body size. In the species consuming mature birch leaves of stable quality, that is, without diet-imposed time constraints for development time, long development led to high body mass. This was revealed by the strongly positive phenotypic and genetic correlations between the traits. In the species consuming growing or senescing leaves, on the other hand, the rapidly deteriorating leaf quality prevented the larvae from gaining high body mass after long development. In these species, the phenotypic and genetic correlations between development time and final mass were negative or zero. In the early-summer species with strong selection for rapid development, genetic variation in development time was low. These results show that the intuitively obvious positive genetic relationship between development time and final body mass is a probable outcome only when the constraints for long development are relaxed. Our study provides the first example of a modification in guild-wide patterns in the genetic architectures brought about by seasonal variation in resource quality.

Adaptive sex ratio variation in pre-industrial human (Homo sapiens) populations?

Sex allocation theory predicts that in a population with a biased operational sex ratio (OSR), parents will increase their fitness by adjusting the sex ratio of their progeny towards the rarer sex, until OSR has reached a level where the overproduction of either sex no longer increases a parent's probability of having grandchildren. Furthermore, in a monogamous mating system, a biased OSR is expected to lead to lowered mean fecundity among individuals of the more abundant sex. We studied the influence of OSR on the sex ratio of newborns and on the population birth rate using an extensive data set (n = 14,420 births) from pre-industrial (1775-1850) Finland. The overall effect of current OSR on sex ratio at birth was significant, and in the majority of the 21 parishes included in this study, more sons were produced when males were rarer than females. This suggests that humans adjusted the sex ratio of their offspring in response to the local OSR to maximize the reproductive success of their progeny. Birth rate and, presumably, also population growth rate increased when the sex ratio (males:females) among reproductive age classes approached equality. However, the strength of these patterns varied across the parishes, suggesting that factors other than OSR (e.g. socioeconomic or environmental factors may also have influenced the sex ratio at birth and the birth rate.