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Effects of marker-assisted selection under different initial frequency of QTL favorable allele.

A ten-generation continuous selection experiment on a single trait in a closed population was carried out by stochastic simulation. It assumed that the trait was controlled by polygenes and a single autosomal bi-allelic marked quantitative trait locus (QTL). Individual breeding values were estimated through animal model marker-assisted best linear unbiased prediction (MB-LUP), and breeding animals were selected according to their breeding values. The effects of three levels initial frequencies of QTL favorable allele on marker-assisted selection (MAS) were studied. The results showed that it would gain higher genetic response when implementing MAS on those traits with lower initial frequency of QTL favorable allele. When the initial frequency of QTL favorable allele was low, although the generation number required for fixing QTL favorable allele in the population would be lengthened, its frequency would be increased faster. However, the initial frequency of QTL favorable allele had little effects on the inbreeding rates.

Algorithms↗

Clinical biochemical parameters of the endangered Catalonian donkey breed: normal values and the influence of sex, age, and management practices effect.

Twelve clinical biochemical parameters were determined in 97 animals of all age groups and both sexes of the endangered Catalonian donkey breed. Significant sex differences were observed for phospholipid concentration (P<0.01). Evaluating the effect of management practices on the various parameters showed significant differences for total bilirubin (P<0.001) and creatinine (P<0.05) concentrations and gamma-glutamyltransferase (P<0.05) activity. Moreover, it was observed that inorganic phosphorus concentration decreased with age (P<0.001), whereas albumin and triglyceride concentrations increased with age (P<0.01 and P<0.001, respectively). Comparison of biochemical ranges obtained for the Catalonian donkey breed with reference ranges for other breeds and populations (Mammoth, USA donkeys, UK donkeys, Indian donkeys and Poitou donkeys), indicated that most values were similar, with the exceptions of enzymatic activities mainly. The results reported in the present study could serve as reference ranges for donkey populations.

Age Factors↗

Natural selection and inheritance of breeding time and clutch size in the collared flycatcher.

Many characteristics of organisms in free-living populations appear to be under directional selection, possess additive genetic variance, and yet show no evolutionary response to selection. Avian breeding time and clutch size are often-cited examples of such characters. We report analyses of inheritance of, and selection on, these traits in a long-term study of a wild population of the collared flycatcher Ficedula albicollis. We used mixed model analysis with REML estimation ("animal models") to make full use of the information in complex multigenerational pedigrees. Heritability of laying date, but not clutch size, was lower than that estimated previously using parent-offspring regressions, although for both traits there was evidence of substantial additive genetic variance (h2 = 0.19 and 0.29, respectively). Laying date and clutch size were negatively genetically correlated (rA = -0.41 +/- 0.09), implying that selection on one of the traits would cause a correlated response in the other, but there was little evidence to suggest that evolution of either trait would be constrained by correlations with other phenotypic characters. Analysis of selection on these traits in females revealed consistent strong directional fecundity selection for earlier breeding at the level of the phenotype (beta = -0.28 +/- 0.03), but little evidence for stabilising selection on breeding time. We found no evidence that clutch size was independently under selection. Analysis of fecundity selection on breeding values for laying date, estimated from an animal model, indicated that selection acts directly on additive genetic variance underlying breeding time (beta = -0.20 +/- 0.04), but not on clutch size (beta = 0.03 +/- 0.05). In contrast, selection on laying date via adult female survival fluctuated in sign between years, and was opposite in sign for selection on phenotypes (negative) and breeding values (positive). Our data thus suggest that any evolutionary response to selection on laying date is partially constrained by underlying life-history trade-offs, and illustrate the difficulties in using purely phenotypic measures and incomplete fitness estimates to assess evolution of life-history trade-offs. We discuss some of the difficulties associated with understanding the evolution of laying date and clutch size in natural populations.

Animals↗

Validation of an approximate approach to compute genetic correlations between longevity and linear traits.

The estimation of genetic correlations between a nonlinear trait such as longevity and linear traits is computationally difficult on large datasets. A two-step approach was proposed and was checked via simulation. First, univariate analyses were performed to get genetic variance estimates and to compute pseudo-records and their associated weights. These pseudo-records were virtual performances free of all environmental effects that can be used in a BLUP animal model, leading to the same breeding values as in the (possibly nonlinear) initial analyses. By combining these pseudo-records in a multiple trait model and fixing the genetic and residual variances to their values computed during the first step, we obtained correlation estimates by AI-REML and approximate MT-BLUP predicted breeding values that blend direct and indirect information on longevity. Mean genetic correlations and reliabilities obtained on simulated data confirmed the suitability of this approach in a wide range of situations. When nonzero residual correlations exist between traits, a sire model gave nearly unbiased estimates of genetic correlations, while the animal model estimates were biased upwards. Finally, when an incorrect genetic trend was simulated to lead to biased pseudo-records, a joint analysis including a time effect could adequately correct for this bias.

Longevity↗

Comparison of index selection and best linear unbiased prediction for simulated layer poultry data.

The advantage of using best linear unbiased prediction (BLUP) of breeding value over different selection indexes was examined for a sex-limited trait in a simulated layer poultry population. The base breeding population consisted of 30 males and 300 females that were unrelated to each other. Heritability for different analyses was assumed to be either .1, .2, or .5. Each generation was reproduced from two hatches each year, with a hatch variance of 3.165% of the phenotypic variance, except for one simulation, in which it was assumed to be 40% to test the effect of a large fixed effect. Parents were selected on 1) BLUP of breeding value, 2) optimum selection index (individual, full-, and half-sibs), 3) classical selection index (as for optimum, but index weight constant across generations), 4) reduced selection index (individual and full-sibs only), or 5) combination of classical and BLUP. The relative selection response with the selection indices compared to the BLUP estimates (except for the reduced selection index) were from 94.5 to 99.4% of BLUP. Inbreeding was higher in the BLUP selected populations, which could offset any advantage of BLUP if the populations were structured so that inbreeding could rise too rapidly.

Animals↗

Prevalence of spondylosis deformans and estimates of genetic parameters for the degree of osteophytes development in Italian Boxer dogs.

The aim of this study was to assess the prevalence of spondylosis deformans and to investigate genetic aspects of the degree of osteophytes development (DOD) in the Italian Boxer dog population. A total of 849 Boxer dogs was radiographed on the thoracic, lumbar, and sacral regions of the spine and scored for DOD. Grading of DOD was performed for all 20 intervertebral sites comprised within the first thoracic site (site T1-T2) and the site between the seventh lumbar and the first sacral vertebra (site L7-S1). Scores for DOD ranged from 0 (no osteophytes development) to 3 (presence of a bony spur formed by osteophytes on adjoining vertebrae). The first five thoracic sites exhibited no variation for DOD and were not considered in the analysis. The prevalence of spondylosis deformans was 84%, and frequency of dogs showing at least one intervertebral site that scored 3 for DOD was 50%. Scores for DOD at different sites were analyzed as different traits. Nongenetic effects influencing DOD scores were sex, age at screening, and the kennel. Posterior densities of heritability (h2) were estimated using a univariate Bayesian analysis. Eight sites exhibited a posterior probability greater than 0.8 for h2 > 10% and were considered in a multivariate restricted maximum likelihood analysis. Estimated h2 from multivariate analysis ranged from 25 to 48% (SE from 5 to 7%). Three sites exhibited h2 estimates greater than 40%. Genetic correlations for DOD scored at different sites ranged from 0.07 to 0.96. All thoracic sites had estimated correlations larger than 0.85 with other thoracic sites. Genetic correlation between the first and the second lumbar site was 0.91. Correlations between thoracic sites and the first two lumbar sites ranged from 0.5 to 0.9. Sites L6-L7 and L7-S1 also exhibited weak relationships with all remaining sites. Breeding values of dogs for DOD at the eight sites were predicted using estimated covariance matrices. A selection index for DOD was computed from predicted breeding values and a set of relative weighting factors produced by a panel of veterinarians. The index was the most important effect influencing phenotypic differences between dogs for average DOD score, number of affected sites, and number of sites with a DOD score > 1 (P < 0.001). The degree of osteophytes development is a trait showing exploitable additive genetic variance, and breeding programs for decreasing prevalence and severity of spondylosis deformans might focus on this trait.

Age Factors↗

Genetic evaluation and best prediction of lactation persistency.

Cows with high persistency tend to produce less milk than expected at the beginning of lactation and more than expected at the end. Best prediction of persistency was calculated as a function of a trait-specific standard lactation curve and a linear regression of test-day deviations on days in milk. Regression coefficients were deviations from a balance point to make yield and persistency phenotypically uncorrelated. The objectives of this study were to calculate (co)variance components and breeding values for best predictions of persistency of milk (PM), fat (PF), protein (PP), and SCS (PSCS) in Holstein cows. Data included 8,682,138 lactations from 4,375,938 cows calving since 1997, and 39,354 sires were evaluated. Sire estimated breeding values (EBV) for PM, PF, and PP were similar and ranged from -0.70 to 0.75 for PM; EBV for PSCS ranged from -0.37 to 0.28. Regressions of sire EBV on birth year were near zero (<0.003) but positive for PM, PF, and PP, and negative for PSCS. Genetic correlations of PM, PF, and PP with PSCS were moderate and favorable, indicating that increasing SCS decreases yield traits, as expected. Genetic correlations among yield and persistency were low to moderate and ranged from -0.09 (PSCS) to 0.18 (PF). This definition of persistency may be more useful than those used in test-day models, which are often correlated with yield. Routine genetic evaluations for persistency are feasible and may allow for improved predictions of yield traits. As calving intervals increase, persistency may have greater value.

Animals↗

Evaluating longevity of composite beef females using survival analysis techniques.

Objectives were to 1) identify risk factors affecting the longevity of beef females, 2) evaluate the utility of measures collected early in life in predicting longevity, and 3) estimate the heritability of longevity when females were culled primarily for not being pregnant following a 45-d breeding season. Data were from 1,379 Composite Gene Combination (CGC; 1/2 Red Angus, 1/4 Charolais, 1/4 Tarentaise) cows born from 1982 through 1999 at the USDA-ARS, Fort Keogh Livestock and Range Research Laboratory, Miles City, MT, and first calving at approximately 2 yr of age. The length of productive life was modeled using Cox regression to identify factors affecting the longevity of beef females. Age at first calving and calf birth weight did not influence longevity. Cows that experienced dystocia were at greater risk of being culled than those that calved without assistance (P < 0.01). On average, as breeding value for cow weight increased, the risk of being culled decreased (P < 0.01), whereas the risk of being culled increased with increasing maternal breeding values for preweaning gain (P < 0.05). Traits measured before 1 yr of age were not useful in predicting the subsequent longevity of cows. The heritability of functional longevity was estimated to be 0.14. Relatively low heritability and the lack of indicators of longevity expressed early in life suggest that genetic improvement of longevity will be difficult. Matching the genetic potential of cows for size and milk production to the production environment such that rebreeding performance is not compromised by concurrent lactation seems to be a consideration in retaining beef females when open cows are culled.

Animals↗

Cryptic evolution in a wild bird population.

Microevolution is expected to be commonplace, yet there are few thoroughly documented cases of microevolution in wild populations. In contrast, it is often observed that apparently heritable traits under strong and consistent directional selection fail to show the expected evolutionary response. One explanation proposed for this paradox is that a genetic response to selection may be masked by opposing changes in the environment. We used data from a 20-year study of collared flycatchers (Ficedula albicollis) to explore selection on, and evolution of, a heritable trait: relative body weight at fledging ('condition'). Despite consistent positive directional selection, on both the phenotypic and the additive genetic component (breeding values, estimated from an animal model) of condition, the mean phenotypic value of this trait in the population has declined, rather than increased, over time. Here we show that, despite this decline, the mean breeding value for condition has increased over time. The mismatch between response to selection at the levels of genotype and phenotype can be explained by environmental deterioration, concealing underlying evolution. This form of cryptic evolution may be common in natural environments.

Animals↗

Challenges and opportunities for integrating genetically modified animals into traditional animal breeding plans.

Techniques have been developed to introduce specific genes from one species into the germplasm of another, including livestock. This paper reviews reports on evaluation, selection, and breeding procedures for introduction and multiplication of transgenes in breeding populations of livestock. Before transgenes are introduced and multiplied in commercial breeding populations, it is necessary to test transgenics extensively for both favorable and unfavorable transgene effects. Parent stock used to produce transgenic founder animals should be selected to excel in polygenic breeding value for economically important traits, especially if founder animals or their sons are to be used directly by AI in commercial populations. However, polygenic breeding value of founder transgenic animals and inbreeding depression are of negligible importance if a transgene is introgressed into a selection nucleus population from three or four generations of backcrossing. Transgenic development should be economically viable for traits with major effects on net merit, especially in the dairy industry, where transgenic sires can be used extensively by AI. A minimum effect of about 10% of the mean (or one phenotypic standard deviation) is a reasonable approximation of the minimum transgene effect that would be necessary to justify introgression into a nucleus swine population. A transgene effect of 10 to 20% of the mean would be required for most economic traits in beef cattle.

Animals↗

Genetic and statistical analyses of strong selection on polygenic traits: what, me normal?

We develop a general population genetic framework for analyzing selection on many loci, and apply it to strong truncation and disruptive selection on an additive polygenic trait. We first present statistical methods for analyzing the infinitesimal model, in which offspring breeding values are normally distributed around the mean of the parents, with fixed variance. These show that the usual assumption of a Gaussian distribution of breeding values in the population gives remarkably accurate predictions for the mean and the variance, even when disruptive selection generates substantial deviations from normality. We then set out a general genetic analysis of selection and recombination. The population is represented by multilocus cumulants describing the distribution of haploid genotypes, and selection is described by the relation between mean fitness and these cumulants. We provide exact recursions in terms of generating functions for the effects of selection on non-central moments. The effects of recombination are simply calculated as a weighted sum over all the permutations produced by meiosis. Finally, the new cumulants that describe the next generation are computed from the non-central moments. Although this scheme is applied here in detail only to selection on an additive trait, it is quite general. For arbitrary epistasis and linkage, we describe a consistent infinitesimal limit in which the short-term selection response is dominated by infinitesimal allele frequency changes and linkage disequilibria. Numerical multilocus results show that the standard Gaussian approximation gives accurate predictions for the dynamics of the mean and genetic variance in this limit. Even with intense truncation selection, linkage disequilibria of order three and higher never cause much deviation from normality. Thus, the empirical deviations frequently found between predicted and observed responses to artificial selection are not caused by linkage-disequilibrium-induced departures from normality. Disruptive selection can generate substantial four-way disequilibria, and hence kurtosis; but even then, the Gaussian assumption predicts the variance accurately. In contrast to the apparent simplicity of the infinitesimal limit, data suggest that changes in genetic variance after 10 or more generations of selection are likely to be dominated by allele frequency dynamics that depend on genetic details.

Fourier Analysis↗

Effect of total allelic relationship on accuracy of evaluation and response to selection.

Total allelic relationship (TA) as a possible alternative to the pedigree-derived additive genetic relationship (RA) is defined. The TA measures the actual allelic identity between individuals for loci segregating for the trait concerned. Its value was studied by simulation in populations of different family structure, different numbers of loci, different numbers of alleles per locus, and different heritability levels. The alternative types of relationship matrices were used in mixed model equations to derive best linear unbiased prediction estimates (EBV) of breeding values (BV). Accuracies of evaluations were calculated as correlations of EBV with true breeding values. In populations with random selection and mating, EBVTA derived using TA had higher accuracies than EBVRA derived using RA. In populations with selection, EBVTA was more accurate and resulted in higher responses than selection on EBVRA. We conclude that not accounting for variation in average measures of relationship and identity in state can be important sources of variance of prediction error, and taking account of them increases the accuracy of selection.

Alleles↗

Effect of including relationships in the estimation of genetic parameters of beef calves.

Variances and covariances for birth weight, gain from birth to weaning (ADG), and 205-d weight were obtained from a sire-dam model and a sire-maternal grandsire model for a herd of Angus and a herd of Hereford cattle. Estimates of direct additive genetic variance (sigma 2A), maternal additive genetic variance (sigma 2M), covariance between direct and maternal additive genetic effects (sigma AM), permanent environmental variance (sigma 2PE), and residual variance (sigma 2e) were obtained both with and without the inverse of the numerator relationship matrix (A-1) included. Estimates of heritability for direct genetic effects (h2A), maternal genetic effects (h2M), and the correlation between direct and maternal effects (rAM) for birth weight were .37, .18, and -.01 in Angus and .53, .23, and -.19 in Herefords, respectively, for the analyses without A-1. For the analyses with A-1, estimates of h2A, h2M, and rAM were .42, .22, and -.12 for Angus and .58, .22, and -.13 for Herefords, respectively. Estimates of h2A, h2M, and rAM for ADG were .43, .15, and -.44 in Angus and .52, .38, and -.03 in Herefords, respectively, without A-1. With A-1, estimates of h2A, h2M, and rAM were .57, .15, and -.32 for Angus and .58, .39, and -.05 for Herefords, respectively. Estimates of h2A, h2M, and rAM for 205-d weight were .49, .15, and -.46 for Angus and .58, .43, and -.06 for Herefords, respectively, without A-1. With A-1, estimates of h2A, h2M, and rAM were .63, .16, and -.36 for Angus and .66, .43, and -.08 for Herefords, respectively. Estimates of h2A were higher with A-1 than without A-1, but estimates of h2M were similar. Using variances and covariances obtained from analyses including A-1 generally gave higher estimates of direct breeding values than using variances and covariances obtained from analyses not including A-1. Both Pearson product-moment and Spearman rank correlations were high (.99) between estimates of breeding values from the two analyses, although some changes in rank did occur.

Analysis of Variance↗

What is genetic quality?

Mate choice is favored by indirect selection if choosy females mate with males of high genetic quality. We believe, however, that testing hypotheses about indirect selection has been constrained by how we conceptualize and therefore empirically measure male genetic quality. Here, we argue that genetic quality is the breeding value of an individual for total fitness. We can therefore learn little about genetic quality from measures of only a few fitness components. We explain breeding value for total fitness, drawing on concepts from life-history theory and quantitative genetics, and suggest how approaches incorporating these insights might result in empirical progress.

Journal Article↗

Genetic trend for milk yield in Guzerat herds participating in progeny testing and MOET nucleus schemes.

Genetic trends for 305-day milk yield (P305) in Brazilian Guzerat herds under selection were compared. Data from 4898 lactations of 3179 purebred and crossbred cows from various regions of Brazil were used. Milk yield was adjusted for mature age and the contemporary groups were defined as herd and calving year. Genetic parameters were estimated using the MTDFREML program. The model included the random effects of animals and permanent environment, and herd-calving year, calving season and genetic composition as fixed effects. Genetic trends were estimated by linear regression of weighted average estimated breeding values as a function of calving year. The average P305 was 2065 +/- 922 kg and the heritability was 0.23 +/- 0.03. The annual genetic trend in estimated breeding values of cows for P305 was 7.09 +/- 0.71 kg between 1987 and 2004, and 6.47 +/- 2.35 kg between 1997 and 2004. For cows born and raised in the multiple ovulation and embryo transfer (MOET) nucleus, this trend was 36.46 +/- 24.54 kg/year between 1997 and 2004, 183.14 +/- 47.94 kg/year between 1997 and 2000, and 9.13 +/- 19.19 kg/year between 2001 and 2004. An average inbreeding coefficient of 0.04 was found for inbred MOET cows in 2004. Increasing the size of the family and introducing new progenies changed reliabilities and predicted transmitting ability estimates of MOET sires. In conclusion, there was a positive genetic trend for milk yield in the MOET nucleus at low inbreeding coefficients due to the increased accuracy and estimated genetic merit, but no changes in the average milk yield were observed.

Animals↗

Reduction of selection differentials in finite populations with a nested full-half sib family structure.

The effect of family structure is of increasing importance in modern breeding schemes, because increased intraclass correlations between relatives due to improved breeding value estimation methods use all family information, and increased family sizes are possible with improved reproduction rates. In addition, reduction of the generation intervals in modern breeding schemes leads to increased intraclass (family) correlations, because young animals have little information on individual or on progeny performance. This paper derives an approximation for the selection differential in a population divided into families. The result is then extended to an approximation for the selection differentials in populations that are divided into full sib families within paternal half sib families. The approximation is compared with Monte Carlo results, from which it is concluded that the approximation is satisfactory (i.e., rarely more than 5% in error). In some practical situations the approximation is shown to be not more than 2% in error. With high intraclass correlations and few animals selected, the reduction of the selection differentials is maximal. When breeding values are based on family information and the family structure is not accounted for, overestimation of the selection differentials can be up to 61%.

Animal Husbandry↗

Evaluation of cytoplasmic genetic effects in beef cattle using an animal model.

Mixed model techniques were used to evaluate the importance of cytoplasmic genetic effects on beef cattle performance. Birth weight (BWT), preweaning average daily gain (ADG), weaning weight (WT205), postweaning gain (PG), ultrasonic backfat thickness (FAT) and predicted milk yield (MILK) data were collected in two herds of Hereford cattle located at Plymouth and Raleigh, North Carolina. Cytoplasmic lines were determined based on the foundation female in the maternal lineage of each animal. An animal model was used to account for all nuclear genetic variation among animals within herds. Direct breeding values were estimated for all animals with records and their parents for all traits. For MILK, permanent environmental effects were estimated for animals with multiple records. For preweaning traits, maternal breeding values and permanent maternal environmental effects also were estimated. In all analyses, F-tests for cytoplasmic effects were not significant. Probability values approached significance (P = .15 to P = .10) only for PG and FAT at Plymouth. Assumptions regarding the ratios of genetic and environmental variances and covariances had no effect on F-tests. Results contrast with earlier analyses of the same data in which nuclear genetic effects were accounted for by including sire and maternal grandsire in the statistical model. This study failed to show that cytoplasmic genetic effects were important sources of variation in performance; residual additive genetic effects were confounded with cytoplasmic lines for these herds. Because cytoplasmic sources may be regarded as founder effects, further research is needed in other populations.

Animals↗

Analysis of litter size and days to lambing in the Ripollesa ewe. I. Comparison of models with linear and threshold approaches.

The analysis focused on model fitting of 2 ewe reproductive traits, litter size, and days to lambing (interval between the introduction of the ram into the flock and the subsequent parturition of the ewes). The experimental data set of the Universitat Autònoma of Barcelona flock was used, including 1,598 records of litter size and 1,699 records of days to lambing from 376 Ripollesa ewes between 1986 and 2005. Univariate and bivariate models were considered as beginning points with linear or threshold approximation for litter size. Model fitting was evaluated in terms of goodness-of-fit and predictive ability, using the mean square error and the correlation between phenotypic and predicted records (rho(y,ŷ)) as reference parameters. The bivariate model was preferable for both variables, minimizing mean square error and maximizing rho(y,ŷ). A threshold approximation for litter size was preferable over a linear approximation. Models were also compared with a simulation study, comparing the correlation coefficient between simulated and predicted breeding values (rho(a,â)). The bivariate threshold model was favored, with a rho(y,ŷ) of 0.677 and 0.834 for litter size and days to lambing, respectively. Correlation coefficients between simulated and predicted breeding values in the bivariate linear model were reduced slightly to 0.651 and 0.831, respectively, and they were lowest with linear univariate models (0.642 and 0.802). Although the bivariate models for ewe litter size and days to lambing were more accurate than the univariate models, the threshold approaches showed a greater advantage under the bivariate model. For the purpose of genetic evaluation of litter size in sheep, use of the threshold-linear model seems justified. In the Ripollesa breed, the evaluation of litter size can benefit from recording birth weight.

Animals↗