Modelling and estimation of genotype by environment interactions for production traits in French dairy cattle

Background

Genotype by environment interactions are currently ignored in national genetic evaluations of dairy cattle. However, this is often questioned, especially when environment or herd management is wide-ranging. The aim of this study was to assess genotype by environment interactions for production traits (milk, protein, fat yields and fat and protein contents) in French dairy cattle using an original approach to characterize the environments.

Methods

Genetic parameters of production traits were estimated for three breeds (Holstein, Normande and Montbéliarde) using multiple-trait and reaction norm models. Variables derived from Herd Test Day profiles obtained after a test day model evaluation were used to define herd environment.

Results

Multiple-trait and reaction norm models gave similar results. Genetic correlations were very close to unity for all traits, except between some extreme environments. However, a relatively wide range of heritabilities by trait and breed was found across environments. This was more the case for milk, protein and fat yields than for protein and fat contents.

Conclusions

No real reranking of animals was observed across environments. However, a significant scale effect exists: the more intensive the herd management for milk yield, the larger the heritability.     

Relationships between genotype x environment interactions and rank orders for a set of genotypes tested in different environments

Multilocation trials in plant breeding lead to cross-classified data sets with rows=genotypes and columns=environments, where the breeder is particularly interested in the rank orders of the genotypes in the different environments. Non-identical rank orders are the result of genotype x environment interactions. Not every interaction, however, causes rank changes among the genotypes (rank-interaction). From a breeder's point of view, interaction is tolerable only as long as it does not affect the rank orders. Therefore, the question arises of under which circumstances does interaction become rank-interaction. This paper contributes to our understanding of this topic. In our study we emphasized the detection of relationships between the similarity of the rank orders (measured by Kendall's coefficient of concordance W) and the functions of the diverse variance components (genotypes, environments, interaction, error). On the basis of extensive data sets on different agricultural crops (faba bean, fodder beet, sugar beet, oats, winter rape) obtained from registration trials (1985–1989) carried out in the Federal Republic of Germany, we obtained the following as main result: W ₂ ^g /( ₂ ^g + ₂ ^v ) where ₂ ^g =genotypic variance and ₂ ^v = ₂ ^ge + ₂ ^o /L with ₂ ^ge =interaction variance, ₂ ^o =error variance and L=number of replications.     

Assessing the impact of natural service bulls and genotype by environment interactions on genetic gain and inbreeding in organic dairy cattle genomic breeding programs

The objective of the present study was to compare genetic gain and inbreeding coefficients of dairy cattle in organic breeding program designs by applying stochastic simulations. Evaluated breeding strategies were: (i) selecting bulls from conventional breeding programs, and taking into account genotype by environment (G×E) interactions, (ii) selecting genotyped bulls within the organic environment for artificial insemination (AI) programs and (iii) selecting genotyped natural service bulls within organic herds. The simulated conventional population comprised 148 800 cows from 2976 herds with an average herd size of 50 cows per herd, and 1200 cows were assigned to 60 organic herds. In a young bull program, selection criteria of young bulls in both production systems (conventional and organic) were either ‘conventional’ estimated breeding values (EBV) or genomic estimated breeding values (GEBV) for two traits with low (h²=0.05) and moderate heritability (h²=0.30). GEBV were calculated for different accuracies (r_mg), and G×E interactions were considered by modifying originally simulated true breeding values in the range from r_g=0.5 to 1.0. For both traits (h²=0.05 and 0.30) and r_mg⩾0.8, genomic selection of bulls directly in the organic population and using selected bulls via AI revealed higher genetic gain than selecting young bulls in the larger conventional population based on EBV; also without the existence of G×E interactions. Only for pronounced G×E interactions (r_g=0.5), and for highly accurate GEBV for natural service bulls (r_mg>0.9), results suggests the use of genotyped organic natural service bulls instead of implementing an AI program. Inbreeding coefficients of selected bulls and their offspring were generally lower when basing selection decisions for young bulls on GEBV compared with selection strategies based on pedigree indices.     

Evaluation of dry matter intake and daily weight gain predictions of the Cornell Net Carbohydrate and Protein System with local breeds of beef cattle in China

Three feeding trials with different dietary treatments were conducted to evaluate predictions of dry matter intake and average daily gain of Chinese local beef cattle by Cornell Net Carbohydrate and Protein System (CNCPS) version 5.0. In Trial 1, 60 growing Jin Nan yellow steers (local yellow cattle breed originated from Shanxi Province) were assigned to two treatments with five pens in each treatment and six animals in each pen. Trial 2 was conducted with 72 Lu Xi yellow growing steers (local yellow cattle breed originated from Shandong Province) in 12 conventional pens. These animals were assigned to four treatments with six animals in each pen. In Trial 3, 60 Lu Xi yellow steers were allocated to four treatments with five animals in each pen. Average daily dry matter intake (DMI) for each steer was measured as a mean of each pen. All the data about animals, environment, management and feeds required by the CNCPS model were collected, and model predictions were generated for animals of each pen. Model predictions were evaluated by mean bias, mean square prediction error (MSPE) and regression between the observed and predicted values. Animals in 9 of 10 treatments showed a very close model-predicted DMI values to the observed. In another treatment, animals had a mean bias of −0.72 kg DM/day (TRT 10 in Trial 3). With overall data included, the regression equation between observed and predicted DMI was: Y_OBS = 1.10X_CNCPS − 0.78 (R² = 0.82; P<0.0001), with an intercept not different (P=0.32) from zero, but a slope still different (P<0.0001) from 1. Although the model-predicted ADG values were very close to the observed values in all trials, most of the predicted values were always lower than the observed values with absolute mean bias less than 0.2. When all data were pooled, the regression equation between observed and predicted ADG was: Y_OBS = 1.07X_CNCPS + 0.02 (R² = 0.73; P<0.0001), with an intercept not different from zero (P=0.88), but a slope that differed from unity (P<0.0001). Results suggest that the CNCPS model is an acceptable model to predict dietary DM intake and average daily gain of Chinese local beef cattle breeds. Further studies are warranted to introduce model adjustment for Chinese conditions.     

Genotype by environment interactions for fitness in hybrid genotypes of Avena barbata

We examined genotype (G) by environment (E) interactions for fitness in mesic and xeric ecotypes of the self-fertilizing annual grass, Avena barbata and their recombinant inbred hybrid progeny. Fitness was assayed (1) in experimental water and nutrient treatments in the greenhouse and (2) in common gardens in each ecotype's native habitat. G x E interactions were significant in the greenhouse. Nevertheless, the same recombinant genotypes tended to have high fitness across all water and nutrient treatments. G x E interactions were less pronounced in the field, and were driven by the contrast between the uniformly low survivorship at the mesic site in 2004 and genetic variation in fitness at the other years/site combinations. Moreover, the mesic ecotype consistently outperformed the xeric in both field and greenhouse. Several of the recombinant genotypes outperformed the parents in the novel greenhouse treatments, but these genotypes did not outperform the mesic parent in field trials. Indeed, it is only in the comparison between field and greenhouse environments that there was a noticeable change in the identity of the most-fit genotype. The results provide evidence that hybridization can create genotypes that are better adapted to newer environments such as those imposed in our greenhouse experiments.     

Characterisation of terminal sire sheep farm systems,based on a range of environmental factors: a case study in the context of genotype by environment interactions using Charollais lambs

The objective of this study was to define different terminal sire flock environments, based on a range of environmental factors, and then investigate the presence of genotype by environment interactions (G×E) between the environments identified. Data from 79 different terminal sire flocks (40 Texel, 21 Charollais and 18 Suffolk), were analysed using principal coordinate and non-hierarchical cluster analyses, the results of which identified three distinct environmental cluster groups. The type of grazing, climatic conditions and the use of vitamins and mineral supplements were found to be the most important factors in the clustering of flocks. The presence of G×E was then investigated using data from the Charollais flocks only. Performance data were collected for 12 181 lambs, between 1990 and 2010, sired by 515 different sires. Fifty six of the sires had offspring in at least two of the three different cluster groups and pedigree information was available for a total of 161 431 animals. Traits studied were the 21-week old weight (21WT), ultrasound muscle depth (UMD) and log transformed backfat depth (LogUFD). Heritabilities estimated for each cluster, for each trait, ranged from 0.32 to 0.45. Genetic correlations estimated between Cluster 1 and Cluster 2 were all found to be significantly lower than unity, indicating the presence of G×E. They were 0.31 (±0.17), 0.68 (±0.14) and 0.18 (±0.21) for 21WT, UMD and LogUFD, respectively. Evidence of sires re-ranking across clusters was also observed. Providing a suitable strategy can be identified, there is potential for the optimisation of future breeding programmes, by taking into account the G×E observed. This would enable farmers to identify and select animals with an increased knowledge as to how they will perform in their specific farm environment thus reducing any unexpected differences in performance.     

Genotype by production environment interaction for birth and weaning weights in a population of composite beef cattle

The objectives of the present study were: (1) to evaluate the importance of genotype×production environment interaction for the genetic evaluation of birth weight (BW) and weaning weight (WW) in a population of composite beef cattle in Brazil, and (2) to investigate the importance of sire×contemporary group interaction (S×CG) to model G×E and improve the accuracy of prediction in routine genetic evaluations of this population. Analyses were performed with one, two (favorable and unfavorable) or three (favorable, intermediate, unfavorable) different definitions of production environments. Thus, BW and WW records of animals in a favorable environment were assigned to either trait 1, in an intermediate environment to trait 2 or in an unfavorable environment to trait 3. The (co)variance components were estimated using Gibbs sampling in single-, bi- or three-trait animal models according to the definition of number of production environments. In general, the estimates of genetic parameters for BW and WW were similar between environments. The additive genetic correlations between production environments were close to unity for BW; however, when examining the highest posterior density intervals, the correlation between favorable and unfavorable environments reached a value of only 0.70, a fact that may lead to changes in the ranking of sires across environments. The posterior mean genetic correlation between direct effects was 0.63 in favorable and unfavorable environments for WW. When S×CG was included in two- or three-trait analyses, all direct genetic correlations were close to unity, suggesting that there was no evidence of a genotype×production environment interaction. Furthermore, the model including S×CG contributed to prevent overestimation of the accuracy of breeding values of sires, provided a lower error of prediction for both direct and maternal breeding values, lower squared bias, residual variance and deviance information criterion than the model omitting S×CG. Thus, the model that included S×CG can therefore be considered the best model on the basis of these criteria. The genotype×production environment interaction should not be neglected in the genetic evaluation of BW and WW in the present population of beef cattle. The inclusion of S×CG in the model is a feasible and plausible alternative to model the effects of G×E in the genetic evaluations.     

Potential for adaptive evolution at species range margins: contrasting interactions between red coral populations and their environment in a changing ocean

Studying population‐by‐environment interactions (PEIs) at species range margins offers the opportunity to characterize the responses of populations facing an extreme regime of selection, as expected due to global change. Nevertheless, the importance of these marginal populations as putative reservoirs of adaptive genetic variation has scarcely been considered in conservation biology. This is particularly true in marine ecosystems for which the deep refugia hypothesis proposes that disturbed shallow and marginal populations of a given species can be replenished by mesophotic ones. This hypothesis therefore assumes that identical PEIs exist between populations, neglecting the potential for adaptation at species range margins. Here, we combine reciprocal transplant and common garden experiments with population genetics analyses to decipher the PEIs in the red coral, Corallium rubrum. Our analyses reveal partially contrasting PEIs between shallow and mesophotic populations separated by approximately one hundred meters, suggesting that red coral populations may potentially be locally adapted to their environment. Based on the effective population size and connectivity analyses, we posit that genetic drift may be more important than gene flow in the adaptation of the red coral. We further investigate how adaptive divergence could impact population viability in the context of warming and demonstrate differential phenotypic buffering capacities against thermal stress. Our study questions the relevance of the deep refugia hypothesis and highlights the conservation value of marginal populations as a putative reservoir of adaptive genetic polymorphism.     

Contrasting levels of genotype by environment interaction for life history and morphological traits in invasive populations of Zaprionus indianus (Diptera: Drosophilidae)

It has been demonstrated that phenotypic plasticity and genotype by environment interaction are important for coping with new and heterogeneous environments during invasions. Zaprionus indianus Gupta (Diptera: Drosophilidae) is an Afrotropical invasive fly species introduced to the South American continent in 1999. This species is generalist and polyphagous, since it develops and feeds in several different fruit species. These characteristics of Z. indianus suggest that phenotypic plasticity and genotype by environment interaction may be important in this species invasion process. In this sense, our aim was to investigate the role of genetic variation for phenotypic plasticity (genotype by environment interaction) in Z. indianus invasion of the South American continent. Specifically, we quantified quantitative genetic variation and genotype by environment interactions of morphological and life history traits in different developmental environments, that is, host fruits. This was done in different populations in the invasive range of Z. indianus in Argentina. Results showed that Z. indianus populations have considerable amounts of quantitative genetic variation. Also, genotype by environment interactions was detected for the different traits analyzed in response to the different developmental environments. Interestingly, the amounts and patterns of these parameters differed between populations. We interpreted these results as the existence of differences in evolutionary potential between populations that have an important role in the short‐ and long‐term success of the Z. indianus invasion process.     

Genetic parameters and genotype–environment interactions for skeleton deformities and growth traits at different ages on gilthead seabream (Sparus aurata L.) in four Spanish regions

One of the most important problems of fish aquaculture is the high incidence of fish deformities, which are mainly skeletal. In this study, genetic parameters on gilthead seabream (Sparus aurata L.) for skeleton deformities at different ages (179, 269, 389, 539 and 689 days) and their correlations with growth traits were estimated, as were as their genotype × environment interactions (G × E) at harvesting age. A total of 4093 offspring from the mass spawning of three industrial broodstocks belonging to the PROGENSA^® breeding programme were mixed and on‐grown by different production systems in four Spanish regions: Canary Islands (tanks and cage), Andalusia (estuary), Catalonia (cage) and Murcia (cage). Parental assignment was inferred using the standardized SMsa1 microsatellite multiplex PCR. From three broodstocks, 139 breeders contributed to the spawn and a total of 297 full‐sibling families (52 paternal and 53 maternal half‐sibling families) were represented. Heritabilities at different ages were medium for growth traits (0.16–0.48) and vertebral deformities (0.16–0.41), and low for any type of deformity (0.07–0.26), head deformities (0.00–0.05) and lack of operculum (0.06–0.11). The genetic correlations between growth and deformity traits were medium and positive, suggesting that to avoid increasing deformities they should be taken into account in breeding programmes when growth is selected. The G × E interactions among the different facilities were weak for length and deformity and strong for growth rate during this period. These results highlight the potential for the gilthead seabream industry to reduce the prevalence of deformities by genetic improvement tools.     

Comparing the consequences of natural selection,adaptive phenotypic plasticity,and matching habitat choice for phenotype–environment matching,population genetic structure,and reproductive isolation in meta‐populations

Organisms commonly experience significant spatiotemporal variation in their environments. In response to such heterogeneity, different mechanisms may act that enhance ecological performance locally. However, depending on the nature of the mechanism involved, the consequences for populations may differ greatly. Building on a previous model that investigated the conditions under which different adaptive mechanisms (co)evolve, this study compares the ecological and evolutionary population consequences of three very different responses to environmental heterogeneity: matching habitat choice (directed gene flow), adaptive plasticity (associated with random gene flow), and divergent natural selection. Using individual‐based simulations, we show that matching habitat choice can have a greater adaptive potential than plasticity or natural selection: it allows for local adaptation while protecting genetic polymorphism despite global mating or strong environmental changes. Our simulations further reveal that increasing environmental fluctuations and unpredictability generally favor the emergence of specialist genotypes but that matching habitat choice is better at preventing local maladaptation by individuals. This confirms that matching habitat choice can speed up the genetic divergence among populations, cause indirect assortative mating via spatial clustering, and hence even facilitate sympatric speciation. This study highlights the potential importance of directed dispersal in local adaptation and speciation, stresses the difficulty of deriving its operation from nonexperimental observational data alone, and helps define a set of ecological conditions which should favor its emergence and subsequent detection in nature.