GENETIC AND MATERNAL INFLUENCES ON BRAIN AND BODY SIZE IN RANDOMBRED HOUSE MICE

Genetic and environmental (chiefly maternal) variance and covariance components were estimated for brain and body size in randombred house mice of three different ages (one, three, and five months). Heritabilities estimated from regressions of offspring on their five-month-old male parents were fairly low over all three ages, averaging about 0.2 for brain size and about 0.3 for body size. Heritability estimates from female parents were higher, however, presumably because of the influence of maternal-environmental components of variance. The total maternal impact was estimated from full-sib analyses and, for the more reliable three- and five-month ages, averaged 23% for brain size and 20% for body size. Phenotypic, genetic, and environmental correlations and regressions of brain and body size also were calculated by parent-offspring and sib-correlation techniques, the phenotypic correlations generally decreasing from about 0.4 at one month to 0.2 at three and five months. Genetic correlations of brain and body size estimated from covariances of offspring on male parents were negative whereas those from female parents were positive in sign, and this as well as positive maternal correlations was taken as evidence of the influence of maternal sources of covariance for these traits. It was concluded that, in addition to direct genetic effects, indirect genetic sources of variance and covariance mediated through the prenatal maternal environment are quite important in the determination of brain size and its association with body size.     

Comparison of year-of-exam- and age-matched estimates of heritability in the Framingham Heart Study data

Several different approaches can be used to examine generational and temporal trends in family studies. The measurement of offspring and parents can be made over a short period of time with parents and offspring having quite different ages, or measurements can be made at the same ages but with decades between parent and offspring measures. A third approach, used in the Framingham Heart Study, has repeated examinations across a broad range of age and time, and provides a unique opportunity to compare these approaches. Parents and offspring were matched both on (year of exam) and on age. Heritability estimates for systolic blood pressure, body mass index, height, weight, cholesterol, and glucose were obtained by regressing offspring on midparent values with and without adjustment for age. Higher estimates of heritability were obtained for age-matched than for year-of-exam-matched data for all traits considered. For most traits, estimates of the heritability of the change over time (slope) of the trait were near zero. These results suggest that the optimal design to identify genetic effects in traits with large age-related effects may be to measure parents and offspring at similar ages and not to rely on age-adjustment or longitudinal measures to account for these temporal effects.     

GENETIC AND SOCIAL INHERITANCE OF BODY AND EGG SIZE IN THE BARNACLE GOOSE (BRANTA LEUCOPSIS)

We present heritability estimates for final size of body traits and egg size as well as phenotypic and genetic correlations between body and egg traits in a recently established population of the barnacle goose (Branta leucopsis) in the Baltic area. Body traits as well as egg size were heritable and, hence, could respond evolutionarily to phenotypic selection. Genetic correlations between body size traits were significantly positive and of similar magnitude or higher than the corresponding phenotypic correlations. Heritability estimates for tarsus length obtained from full-sib analyses were higher than those obtained from midoffspring-midparent regressions, and this indicates common environment effects on siblings. Heritabilities for tarsus length obtained from midoffspring-mother regressions were significantly higher than estimates from midoffspring-father regressions. The results suggest that this discrepancy is not caused by maternal effects through egg size, nor by extra-pair fertilizations, but by a socially inherited foraging site fidelity in females.     

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.     

Genetic and environmental effects on semen traits in Lacaune and Manech tête rousse AI rams (Open Access publication)

Data from 51 107 and 11 839 ejaculates collected on rams of the "Lacaune" and "Manech tête rousse" breeds, respectively, were analysed to determine environmental and genetic factors affecting semen production traits (ejaculate volume, semen concentration, number of spermatozoa and motility) in young (≤1 year) and adult (≥2 years) rams. Fixed effects and variance components were estimated using multiple trait animal models within each breed. For all traits, the main environmental effects identified were year, season, number of ejaculations, daily variation, interval from previous to current collection and age. Heritability estimates were moderate for volume, concentration and number of spermatozoa (0.12 to 0.33) and lower for motility (0.02 to 0.14). Genetic correlations between ages differed from 1 for all traits (0.14 to 0.90), indicating that semen characteristics corresponded to different traits in young and adult rams. Genetic and phenotypic correlations among traits within age category were globally similar for the different breeds and categories of animals.     

Estimation of genetic parameters for post-weaning traits of Kermani sheep

The objective of the present study was to estimate genetic parameters for post-weaning traits in Kermani sheep. Traits were included 6-month weight (6MW), 9-month weight (9MW), yearling weight (YW), greasy fleece weight at first shearing (GFW) and greasy fleece weights at various shearings (RFW). Data and pedigree information used in this research were collected at Breeding Station of Kermani sheep during 1993–2004. Genetic parameters were estimated with single- and multi-traits analysis using restricted maximum likelihood (REML) procedures, under animal models. Log likelihood ratio test indicated the most appropriate model for 6MW and 9MW should included direct additive genetic effects as well as maternal permanent environmental effects. However the most appropriate model for YW and GFW had only the direct additive genetic effects. The effects of sex, age of dam and year of birth were significant on body weight traits (P < 0.01). GFW was influenced significantly by sex and year of birth (P < 0.01) but was not affected by age of dam (P > 0.05). Type of birth was no significant effect on studied traits (P > 0.05). Also, the age of lamb at weighing time was a significant influence on 6MW, 9MW and YW. Direct heritability estimates for 6MW, 9MW, YW and GFW were 0.32, 0.03, 0.15 and 0.15, respectively. Maternal permanent environmental estimates of 0.09 were obtained for 6MW and 9MW. Genetic correlation estimates between mentioned traits ranged from 0.51 to 0.99. Phenotypic correlations were generally lower than those of genetic correlation and varied from 0.05 to 0.79 for various traits. The environmental correlations estimates between GFW with growth traits were low, but between other traits were positive and high, ranged from 0.54 to 0.72. The value of repeatability estimated for greasy fleece weight was 0.22.     

Development of a selection index to improve market value of cultured Atlantic salmon (Salmo salar).

Genetic parameters were estimated on growth and development traits using analyses of variance and covariance of 42 full-sib families from a select line, 39 full-sib families from a control line of the same strain, and 37 full-sib families from another unselected strain. The traits included percent 1+ smolts, percent sexually immature fish after 1 year in seawater (nongrilse), and fork length measured at intervals throughout the production cycle. The number of fish sampled per family was 30 of 300-1500 for all freshwater traits and 15-60, i.e., all individuals present, for all seawater traits. Heritability estimates of freshwater traits had a broad range (0.15-0.61), but estimates were lower for seawater traits (0.06-0.29). There was a general reduction in heritability estimates for traits measured at increasing ages both in freshwater and seawater. These reductions may in part represent declines of maternal and common environmental effects that inflate heritability estimated from full-sib families. The genetic correlations indicated positive associations between freshwater and seawater growth parameters. However, the correlations decreased with increasing intervals between measurements. A selection index was developed using the estimates of the genetic and phenotypic parameters for three economically important traits: percent 1 + smolts, percent nongrilse, and harvest length. The seawater traits, percent nongrilse, and fork length at 17 months (harvest length) were the most important in the index.     

Direct and social genetic parameters for growth and fin damage traits in Atlantic cod (Gadus morhua)

Background

The aim of the study was to estimate genetic parameters for direct and social genetic effects (SGE) for growth and welfare traits in farmed Atlantic cod (Gadus morhua). A SGE refers to the effect of an individual’s genes on trait performance of its social partners. In total, 2100 individually tagged juveniles from 100 families at an average age of 222 days post-hatching were used. Each family was separated into three groups of seven fish, and were randomly assigned to 100 experimental tanks, together with fish from two other families. Body weight and length of the first, second and third dorsal fin and the caudal fin measured by digital image analysis were measured at the start of the experiment, after two weeks, and after six weeks. Fin erosion was scored subjectively after six weeks. Variance components estimated using a conventional animal model were compared to those of an animal model including a SGE.

Results

Heritabilities from the conventional animal model ranged from 0.24 to 0.34 for body weight and 0.05 to 0.80 for fin length. Heritabilities for fin erosion were highest for the first dorsal fin (0.83 ± 0.08, mean ± standard error) and lowest for the third dorsal fin (0.01 ± 0.04). No significant SGE were found for body weight, whereas SGE for fin lengths were significant after two and six weeks. Contributions to the total heritable variance were equal to 21.5% (6.1 ± 2.1) for the direct effect, 33.1% (9.4 ± 3.2) for the direct-social covariance, and 45.4% (12.9 ± 4.1) for the social variance for length of the first dorsal fin. For fin erosion, SGE were only significant for the second and third dorsal fin.

Conclusions

Including SGE for fin length and fin erosion in the animal model increased the estimated heritable variation. However, estimates of total heritable variances were inaccurate and a larger experiment is needed to accurately quantify total heritable variance. Despite this, our results demonstrate that considering social breeding values for fin length or fin erosion when selecting fish will enable us to improve response to selection for welfare traits in Atlantic cod juveniles.     

Genetic parameters for direct and maternal effects on body weights of Draa goats

Genetic parameters and (co)variance components were estimated for weights at birth and at 30, 90 and 180 days of age for Draa goat maintained at Ouarzazate station over a period of 18 years (1988–2005). Records of 1498 kids, the progeny of 46 sires and 404 dams were used in the study. Analyses were carried out by restricted maximum likelihood. Six different animal models including or ignoring maternal genetic or permanent environmental effects were fitted for all traits. The Model 2 with only permanent environmental maternal effects seemed most suitable. Estimates of direct heritability from this model were 0.16 for birth weight and 0.07, 0.11 and 0.11 for weights at 30, 90 and 180 days, respectively. Maternal heritability estimates varied from 0.00 to 0.24 for all traits according to the model used (Models 4–6). Bivariate analysis by Model 2 was also used to estimate genetic correlations between traits. The estimates of genetic and phenotypic correlations among weights were positive and intermediate to high in value. Despite the low estimated heritabilities of body weight traits of Draa goat, there is a small genetic variability that may be exploited to improve growth performance.     

Estimating quantitative genetic parameters in wild populations: a comparison of pedigree and genomic approaches

The estimation of quantitative genetic parameters in wild populations is generally limited by the accuracy and completeness of the available pedigree information. Using relatedness at genomewide markers can potentially remove this limitation and lead to less biased and more precise estimates. We estimated heritability, maternal genetic effects and genetic correlations for body size traits in an unmanaged long‐term study population of Soay sheep on St Kilda using three increasingly complete and accurate estimates of relatedness: (i) Pedigree 1, using observation‐derived maternal links and microsatellite‐derived paternal links; (ii) Pedigree 2, using SNP‐derived assignment of both maternity and paternity; and (iii) whole‐genome relatedness at 37 037 autosomal SNPs. In initial analyses, heritability estimates were strikingly similar for all three methods, while standard errors were systematically lower in analyses based on Pedigree 2 and genomic relatedness. Genetic correlations were generally strong, differed little between the three estimates of relatedness and the standard errors declined only very slightly with improved relatedness information. When partitioning maternal effects into separate genetic and environmental components, maternal genetic effects found in juvenile traits increased substantially across the three relatedness estimates. Heritability declined compared to parallel models where only a maternal environment effect was fitted, suggesting that maternal genetic effects are confounded with direct genetic effects and that more accurate estimates of relatedness were better able to separate maternal genetic effects from direct genetic effects. We found that the heritability captured by SNP markers asymptoted at about half the SNPs available, suggesting that denser marker panels are not necessarily required for precise and unbiased heritability estimates. Finally, we present guidelines for the use of genomic relatedness in future quantitative genetics studies in natural populations.     

Genetic Effects of Acute Spermatogonial X-Irradiation of the Laboratory Rat

The genetic effects of one generation of spermatogonial X-irradiation in rats, by a single dose of 600r in one experiment and by a fractionated dose of 450r in another, were measured in three generations of their descendants. Estimates of dominant lethal mutation rates—(2 to 3) x 10 ^-4/gamete/r—from litter size differences between irradiated and nonirradiated stock were consistent with previous estimates from rats and mice. Similar consistency was found for estimates of sex-linked recessive mutation rates—(1 to 2) x 10^-4 chromosome/r—from male proportions within strains; however, when measured in crossbreds the proportion of males was higher in the irradiated than in the nonirradiated lines. This inconsistency in results is in keeping with the contradictory results reported for recessive sex-linked lethal mutation rates in mice. The effects used to estimate recessive lethal mutation rates which were unusually high—(2 to 14) x 10^-4/gamete/r—were not significant. Other factors that could have contributed to the observed effects are postulated.     

QTL mapping of <Emphasis Type="Italic">Sclerotinia</Emphasis> midstalk-rot resistance in sunflower

In many sunflower-growing regions of the world, Sclerotinia sclerotiorum (Lib.) de Bary is the major disease of sunflower (Helianthus annuus L.). In this study, we mapped and characterized quantitative trait loci (QTL) involved in resistance to S. sclerotiorum midstalk rot and two morphological traits. A total of 351 F₃ families developed from a cross between a resistant inbred line from the germplasm pool NDBLOS and the susceptible line CM625 were assayed for their parental F₂ genotype at 117 codominant simple sequence repeat markers. Disease resistance of the F₃ families was screened under artificial infection in field experiments across two sowing times in 1999. For the three resistance traits (leaf lesion, stem lesion, and speed of fungal growth) and the two morphological traits, genotypic variances were highly significant. Heritabilities were moderate to high (h²=0.55–0.89). Genotypic correlations between resistance traits were highly significant (P<0.01) but moderate. QTL were detected for all three resistance traits, but estimated effects at most QTL were small. Simultaneously, they explained between 24.4% and 33.7% of the genotypic variance for resistance against S. sclerotiorum. Five of the 15 genomic regions carrying a QTL for either of the three resistance traits also carried a QTL for one of the two morphological traits. The prospects of marker-assisted selection (MAS) for resistance to S. sclerotiorum are limited due to the complex genetic architecture of the trait. MAS can be superior to classical phenotypic selection only with low marker costs and fast selection cycles.     

Genetic parameter estimates for pre-weaning weight traits in Dorper sheep

Genetic parameters were estimated for birth-, 42-day, and 100-day (weaning) weight in the Dorper flock of the Glen Agricultural Institute in South Africa. Direct heritability estimates of 0.11, 0.28 and 0.20 and maternal heritability estimates of 0.10, 0.10 and 0.10 were obtained for body weights at birth, 42 and 100 days, respectively. The corresponding genetic correlation estimates between direct and maternal effects were 0.35, −0.63 and −0.58, respectively. Both direct and maternal genetic correlation estimates among the traits were of moderate to high magnitude and positive. It is concluded that the traits can be improved by selection with no serious antagonisms among traits studied.     

Intraspecific variation in seed dispersal of a Neotropical tree and its relationship to fruit and tree traits

The distribution of wind‐dispersed seeds around a parent tree depends on diaspore and tree traits, as well as wind conditions and surrounding vegetation. This study of a neotropical canopy tree, Platypodium elegans, explored the extent to which parental variation in diaspore and tree traits explained (1) rate of diaspore descent in still air, (2) distributions of diaspores dispersed from a 40‐m tower in the forest, and (3) natural diaspore distributions around the parent tree. The geometric mean rate of descent in still air among 20 parents was highly correlated with geometric mean wing loading^1/2 (r = 0.84). However, diaspore traits and rate of descent predicted less variation in dispersal distance from the tower, although descent rate⁻¹ consistently correlated with dispersal distance. Measured seed shadows, particularly their distribution edges, differed significantly among six parents (DBH range 62–181 cm) and were best fit by six separate anisotropic dispersal kernels and surveyed fecundities. Measured rate of descent and tree traits, combined in a mechanistic seed dispersal model, did not significantly explain variation among parents in natural seed dispersal distances, perhaps due to the limited power to detect effects with only six trees. Seedling and sapling distributions were at a greater mean distance from the parents than seed distributions; saplings were heavily concentrated at far distances. Variation among parents in the distribution tails so critical for recruitment could not be explained by measured diaspore or tree traits with this sample size, and may be determined more by wind patterns and the timing of abscission in relation to wind conditions. Studies of wind dispersal need to devote greater field efforts at recording the “rare” dispersal events that contribute to far dispersal distances, following their consequences, and in understanding the mechanisms that generate them.     

Mixed model analysis of a selection experiment for food intake in mice

Data from 23 generations of mice selected for increased and reduced appetite were analysed by Restricted Maximum Likelihood fitting an animal model with litters as additional random effects. Traits considered were food intake between 4 and 6 weeks of age adjusted for 4-week body weight (AFI), the selection criterion, and body weight at 6 weeks (6WW). Selection was carried out within families. A high and a low selection line and a control were maintained in each of three replicates. Analyses were performed for each replicate separately taking subsets of the data spanning different numbers of generations. Overall estimates of heritabilities were 0.15 for AFI, which agreed well with realized heritability estimates, and 0.42 for 6WW. The litter variance, expressed as a proportion of the phenotypic variance, was 0.21 for both traits, yielding intraclass correlations of full-sibs of 0.29 and 0.42, respectively. Similar results were obtained for variances of each trait using univariate and multivariate analyses. From the latter, estimates of correlations between the two traits were 0.46 for additive genetic, -0.19 for litter and 0.31 for residual effects, resulting in a phenotypic correlation of 0.23. Analyses of data from generations 2-7, 8-13 and 14-23 separately showed a marked decrease in genetic variance and heritability in later generations for both traits. Heritabilities of AFI, for instance, were 0.24, 0.10 and 0.07, respectively. These changes could not be attributed to the effects of inbreeding or of selection in an infinitesimal model and suggested that some change in variance due to change in gene frequency had occurred during the course of the experiment.     

Primate population studies at Polonnaruwa. II. Heritability of body measurements in a natural population of toque macaques (Macaca sinica)

The heritability of quantitative traits, or the proportion of phenotypic variation due to additive genetic or heritable effects, plays an important role in determining the evolutionary response to natural selection. Most quantitative genetic studies are performed in the laboratory, due to difficulty in obtaining genealogical data in natural populations. Genealogies are known, however, from a unique 20-year study of toque macaques (Macaca sinica) at Polonnaruwa, Sri Lanka. Heritability in this natural population was, therefore, estimated. Twenty-seven body measurements representing the lengths and widths of the head, trunk, extremities, and tail were collected from 270 individuals. The sample included 172 offspring-mother pairs from 39 different matrilineal families. Heritabilities were estimated using traditional mother-offspring regression and maximum likelihood methods which utilize all genealogical relationships in the sample. On the common assumption that environmental (including social) factors affecting morphology were randomly distributed across families, all but two of the traits (25 of 27) were significantly heritable, with an average heritability of 0.51 for the mother-offspring analysis and 0.56 for the maximum likelihood analysis. Heritability estimates obtained from the two analyses were very similar. We conclude that the Polonnaruwa macaques exhibit a comparatively moderate to high level of heritability for body form. © 1992 Wiley-Liss, Inc.     

籼粳杂交稻穗部性状的遗传效应及其与环境互作

采用包括基因型与环境互作效应的加性-显性-加性×加性上位性遗传模型,分析了不同环境下籼粳杂交稻穗部性状的遗传特点.结果表明,除了主穗粒数的加性与环境互作和二次枝梗数的显性与环境互作不显著外,其他性状均存在显著和极显著的加性、显性、加性×加性上位性遗传效应及其与环境的互作效应,其中均以显性效应为主,显性与环境互作效应对枝梗性状的影响较为明显.遗传率分析表明,各性状的普通广义遗传率最大,互作遗传率也有一定作用.杂种优势预测表明,除了一次、二次枝梗数外,其他性状均表现正向的杂种优势,基因型与环境互作只影响杂种优势表达的程度,而不改变其方向.遗传效应预测值结果表明,IR66158-37、IR65600-85、明恢63和R6694个亲本可以明显改良杂交后代多数穗部性状,且环境影响程度较小,可作为优良亲本列于育种计划中.     

Inheritance of body size in the Barnacle Goose under different environmental conditions

Heritabilities, genetic variances and covariances for body size traits, i.e. tarsus length, head length and body mass, were estimated under different environmental conditions in a Barnacle Goose (Branta leucopsis) population. Under poor growth conditions, that is, when average body size of fully grown offspring in a given cohort was small, the offspring-parent regressions and full-sib analyses yielded heritability estimates not significantly different from zero. By contrast, when growth conditions were normal or good the heritability estimates were generally significantly positive. Comparisons of genetic covariance estimates indicated that they also differed across the analysed environmental conditions. This result, together with similar results obtained in studies of passerine birds, suggests that genotype-environment interactions might be frequent within the range of environments normally encountered by birds in natural populations. If general, such results might question the validity of assuming approximate constancy of additive genetic variances and covariances over time and environments in evolutionary models.     

Effectiveness of and Financial Returns to Voluntary Medical Male Circumcision for HIV Prevention in South Africa: An Incremental Cost-Effectiveness Analysis

BackgroundEmpirical studies and population-level policy simulations show the importance of voluntary medical male circumcision (VMMC) in generalized epidemics. This paper complements available scenario-based studies (projecting costs and outcomes over some policy period, typically spanning decades) by adopting an incremental approach—analyzing the expected consequences of circumcising one male individual with specific characteristics in a specific year. This approach yields more precise estimates of VMMC’s cost-effectiveness and identifies the outcomes of current investments in VMMC (e.g., within a fiscal budget period) rather than of investments spread over the entire policy period.Methods/FindingsThe model has three components. We adapted the ASSA2008 model, a demographic and epidemiological model of the HIV epidemic in South Africa, to analyze the impact of one VMMC on HIV incidence over time and across the population. A costing module tracked the costs of VMMC and the resulting financial savings owing to reduced HIV incidence over time. Then, we used several financial indicators to assess the cost-effectiveness of and financial return on investments in VMMC. One circumcision of a young man up to age 20 prevents on average over 0.2 HIV infections, but this effect declines steeply with age, e.g., to 0.08 by age 30. Net financial savings from one VMMC at age 20 are estimated at US$617 at a discount rate of 5% and are lower for circumcisions both at younger ages (because the savings occur later and are discounted more) and at older ages (because male circumcision becomes less effective). Investments in male circumcision carry a financial rate of return of up to 14.5% (for circumcisions at age 20). The cost of a male circumcision is refinanced fastest, after 13 y, for circumcisions at ages 20 to 25. Principal limitations of the analysis arise from the long time (decades) over which the effects of VMMC unfold—the results are therefore sensitive to the discount rate applied, and more generally to the future course of the epidemic and of HIV/AIDS-related policies pursued by the government.ConclusionsVMMC in South Africa is highly effective in reducing both HIV incidence and the financial costs of the HIV response. The return on investment is highest if males are circumcised between ages 20 and 25, but this return on investment declines steeply with age.     

Estimation of genetic parameters of semen characteristics and reproductive traits in AI boars

(Co)variance components and further genetic parameters of boar semen characteristics and reproductive traits were estimated using the REML procedure applied to multi-trait animal models. The calculations were based on data from 210,733 ejaculates stemming from 2862 AI boars and collected from 1990 to 1997 in insemination stations for boars in the Czech Republic. Equal model equations for all traits included the AI station and the breed or breed combination as fixed effects, the interval between two collections for the boar as covariable and the animal and residual effects as random effects. The following heritabilities were estimated: semen volume 0.58, sperm concentration 0.49, progressive motion of spermatozoa 0.38, abnormal spermatozoa 0.34, number of total spermatozoa 0.42, number of insemination doses 0.40, number of piglets born alive 0.08, total number of piglets born 0.05 and conception rate 0.29. Heritabilities and genetic correlations were estimated on average values for each boar.