Effects of breed proportion and components of heterosis for semen traits in a composite cattle breed

The aim of this study was to estimate the non‐additive genetic effects of the dominance component of heterosis as well as epistatic loss on semen traits in admixed Swiss Fleckvieh, a composite of Simmental (SI) and Red Holstein Friesian (RHF) cattle. Heterosis is the additional gain in productivity or fitness of cross‐bred progeny over the mid‐purebred parental populations. Intralocus gene interaction usually has a positive effect, while epistatic loss generally reduces productivity or fitness due to lack of evolutionarily established interactions of genes from different breeds. Genotypic data on 38,205 SNP of 818 admixed, as well as 148 RHF and 213 SI bulls as the parental breeds were used to predict breed origin of alleles. The genomewide locus‐specific breed ancestries of individuals were used to calculate effects of breed difference as well as the dominance component of heterosis, while proxies for two definitions of epistatic loss were derived from 100,000 random pairs of loci. The average Holstein Friesian ancestry in admixed bulls was estimated 0.82. Results of fitting different linear mixed models showed including the dominance component of heterosis considerably improved the model adequacy for three of the four traits. Inclusion of epistatic loss increased the accuracy of the models only for our new definition of the epistatic effect for two traits, while the other definition was so highly correlated with the dominance component that statistical separation was impossible.     

Optimal genetic contribution selection in Danish Holstein depends on pedigree quality

The aim of this study was to assess the importance of pedigree depth when performing optimal contribution selection as implemented in the software program EVA. This was done by applying optimal genetic contribution in the breeding program of the major Danish Dairy breed Danish Holstein. In the analyses earlier breeding decisions were considered by including all AI waiting- and young bulls and contract matings. Twenty potential sires, 2169 potential dams, 1421 AI-bulls and 754 contract matings plus pedigree animals were included. Results showed that the outcome was very dependent on quality of pedigree, also for information going more than 25 years (5–7 generations) back. The analyses showed that EVA works satisfactorily as a management tool for planning of breeding schemes with respect to contributions of sires of sons at population level in maximising the genetic gain, while controlling the increase in future inbreeding. The more weight put on the average additive genetic relationship in next generation relative to genetic merit, the lower the average merit of the matings, and the lower average additive genetic relationship among the chosen matings and the present breeding animals. Furthermore more weight on average additive genetic relationship gives a more diverse use of sires of sons. Given the potential sires and dams the average additive genetic relationship among the selected matings and the present breeding animals can be reduced from 0.1621 to 0.1495 at the cost of 0.7 genetic S.D. units at the total merit index. This reduction was obtained when selection was only on reduction of average relationship compared to selection only on genetic merit. Optimal genetic contribution selection is a promising tool for managing breeding schemes for populations facing inbreeding problems, such as Danish Holstein and other dairy breeds. However sufficient pedigree information is a necessity.     

Genomic estimation of additive and dominance effects and impact of accounting for dominance on accuracy of genomic evaluation in sheep populations

The objectives of this study were to estimate the additive and dominance variance component of several weight and ultrasound scanned body composition traits in purebred and combined cross‐bred sheep populations based on single nucleotide polymorphism (SNP) marker genotypes and then to investigate the effect of fitting additive and dominance effects on accuracy of genomic evaluation. Additive and dominance variance components were estimated in a mixed model equation based on “average information restricted maximum likelihood” using additive and dominance (co)variances between animals calculated from 48,599 SNP marker genotypes. Genomic prediction was based on genomic best linear unbiased prediction (GBLUP), and the accuracy of prediction was assessed based on a random 10‐fold cross‐validation. Across different weight and scanned body composition traits, dominance variance ranged from 0.0% to 7.3% of the phenotypic variance in the purebred population and from 7.1% to 19.2% in the combined cross‐bred population. In the combined cross‐bred population, the range of dominance variance decreased to 3.1% and 9.9% after accounting for heterosis effects. Accounting for dominance effects significantly improved the likelihood of the fitting model in the combined cross‐bred population. This study showed a substantial dominance genetic variance for weight and ultrasound scanned body composition traits particularly in cross‐bred population; however, improvement in the accuracy of genomic breeding values was small and statistically not significant. Dominance variance estimates in combined cross‐bred population could be overestimated if heterosis is not fitted in the model.     

There is room for selection in a small local pig breed when using optimum contribution selection: a simulation study

Selection progress must be carefully balanced against the conservation of genetic variation in small populations of local breeds. Well-defined breeding programs?with specified selection traits are rare in local pig breeds. Given the small population size,?the focus is often on the management of genetic diversity. However, in local breeds, optimum contribution selection can be applied to control the rate of inbreeding and to avoid reduced performance in traits with high market value. The aim of this study was to assess the extent to which a breeding program aiming for improved product quality in a small local breed would be feasible. We used stochastic simulations to compare 25 scenarios. The scenarios differed in?size of population, selection intensity of boars, type of selection (random selection, truncation selection based on BLUP breeding values, or optimum contribution selection based on BLUP breeding values), and heritability of?the selection trait. It was assumed that the local breed is used in an extensive system for a high-meat-quality market.?The?simulations showed that in the smallest population (300 female reproducers), inbreeding increased by 0.8% when selection was performed at random. With optimum contribution selection, genetic progress can be achieved that is almost as great as that with truncation selection based on BLUP breeding values (0.2 to 0.5 vs. 0.3 to 0.5 genetic SD, P < 0.05), but at a considerably decreased rate of inbreeding (0.7 to 1.2 vs. 2.3 to 5.7%, P < 0.01). This confirmation of the potential utilization of OCS even in small populations is important in the context of sustainable management and the use of animal genetic resources.     

Periodic rotational crosses. I. Breed and heterosis utilization

Periodic rotational crosses differ from conventional rotations by using sire breeds an unequal number of generations but in a regular sequence. Formulas for breed composition and heterosis in offspring from periodic rotations at equilibrium averaged over all generations of a cycle and for each generation of the cycle are presented. Coefficients of squares and products of breed differences and heterosis used to calculate inter-generational variance are also derived. Several specific periodic rotations utilizing a range of breed proportions were found to use from 70 to 95% as much heterosis as conventional rotations using the same number of breeds equally. Estimates of swine and cattle additive breed and heterosis effects taken from the literature were applied to the formulas. In both the swine and cattle examples, periodic rotations were found that equaled or exceeded the conventional rotations using the same number of breeds and also had lower inter-generational variance.     

Breed effects and heterosis in advanced generations of composite populations for preweaning traits of beef cattle

The effects of heterosis for gestation length, dystocia, calf survival, birth weight, 200-d weight, and ADG from birth to weaning were evaluated in F1, F2, and combined F3 and F4 generations in three composite populations. Breed effects were evaluated for the nine parental breeds (Red Poll, Hereford, Angus, Limousin, Braunvieh, Pinzgauer, Gelbvieh, Simmental, and Charolais) that contributed to the three composite populations. Breed effects were significant for all traits evaluated except survival at birth. The large differences among breeds in additive direct and additive maternal genetic effects offer a great opportunity to use the genetic differences among breeds to achieve and maintain optimum additive genetic (breed) composition to match genetic resources to a wide range of production-marketing ecosystems. There was no heterosis for gestation length. Mean heterosis for dystocia was significant estimated in F1 but not in F2 or in the combined F3 and F4 generations. Mean heterosis was not significant in any generation for survival at birth, to 72 h, and to weaning for the F1 generation; mean heterosis was significant for survival to weaning for the F2 generation and approached significance (P = .06) for the combined F3 and F4 generations. Mean heterosis over all composite populations and heterosis for each composite population were significant in all generations for weight at birth and at 200 d and for ADG from birth to weaning. Retained heterosis was not less than expected from retained heterozygosity in composite populations for the traits evaluated. These results suggest that heterosis for these traits likely is due to dominance effects and, thus, can be attributed to the recovery of accumulated inbreeding depression in the parental breeds.     

Genetic variation in composite and parental populations: expectations for levels of dominance and gene frequency.

Usefulness of multibreed composites depends on: 1) adaptation of the average breed effects combined, 2) amount of initial heterosis retained, and 3) the rate of further improvement from selection. Potential improvement could be increased, relative to parental stocks, by retained heterosis in reproductive rate and by any increase in additive genetic variation in the composite. A single-locus, two-allele model with additive plus dominant gene effects was used to clarify expected changes from varying: 1) degree of dominance (d) from none to over-dominance, 2) variance among parental lines in frequency of a dominant allele (Vq), and 3) mean gene frequency in the composite (qc). Simulation for a three-breed composite (1/2, 1/4, 1/4) showed that expected heterosis retained in the composite is proportional to d and Vq and is highest for any d and Vq when qc is intermediate. Percentage of change of the composite from parental mean in additive genetic variance (delta Vac,%) increases most with parental diversity (Vq) when d = 0 but changes with higher d from positive when q is less than .5 to negative when q is greater than .5. Therefore, the expected association of delta Vac with level of heterosis retained (delta H,%) is always smaller for higher d and changes from positive when qc less than .5 to negative for the more likely higher equilibrium values of qc. Thus, greater selection intensity (from a higher reproductive rate) in composites than in parent stocks may not be accompanied by increased genetic variability and change in response per unit of selection applied may be limited.     

Optimizing female allocation to reproductive technologies considering merit,inbreeding and cost in nucleus breeding programmes with genomic selection

Female reproductive technologies such as multiple ovulation and embryo transfer (MOET) and juvenile in vitro fertilization and embryo transfer (JIVET) have been shown to accelerate genetic gain by increasing selection intensity and decreasing generation interval. Genomic selection (GS) increases the accuracy of selection of young candidates which can further accelerate genetic gain. Optimal contribution selection (OCS) is an effective method of keeping the rate of inbreeding at a sustainable level while increasing genetic merit. OCS could also be used to selectively and optimally allocate reproductive technologies in mate selection while accounting for their cost. This study uses stochastic simulation to simulate breeding programmes that use a combination of artificial insemination (AI) or natural mating (N), MOET and JIVET with GS. OCS was used to restrict inbreeding to 1.0% increase per generation and also to optimize use of reproductive technologies, considering their effect on genetic gain as well as their cost. Two Australian sheep breeding objectives were used as an example to illustrate the methodology—a terminal sire breeding objective (A) and a dual‐purpose self‐replacing breeding objective (B). The objective function used for optimization considered genetic merit, constrained inbreeding and cost of technologies where costs were offset by a premium paid to the seedstock breeder investing in female reproductive technologies. The premium was based on the cumulative discounted expression of genetic merit in the progeny of a commercial tier in the breeding programme multiplied by the proportion of that benefit received by the breeder. With breeding objective B, the highest premium of 64% paid to the breeder resulted in the highest allocation of reproductive technologies (4%–10% for MOET and 19%–54% for JIVET) and hence the highest annual genetic gain. Conversely, breeding objective A, which had a lower dollar value of the breeding objective and a maximum of 5% mating types for JIVET and zero for MOET were optimal, even when highest premiums were paid. This study highlights that the level of investment in breeding technologies to accelerate genetic gain depends on the investment of genetic improvement returned to the breeder per index point gain achieved. It also demonstrates that breeding programmes can be optimized including allocation of reproductive technologies at the individual animal level. Accounting for revenue to the breeder and cost of the technologies can facilitate more practical decision support for beef and sheep breeders.     

Genomic prediction of growth in pigs based on a model including additive and dominance effects

Independent of whether prediction is based on pedigree or genomic information, the focus of animal breeders has been on additive genetic effects or ‘breeding values’. However, when predicting phenotypes rather than breeding values of an animal, models that account for both additive and dominance effects might be more accurate. Our aim with this study was to compare the accuracy of predicting phenotypes using a model that accounts for only additive effects (MA) and a model that accounts for both additive and dominance effects simultaneously (MAD). Lifetime daily gain (DG) was evaluated in three pig populations (1424 Pietrain, 2023 Landrace, and 2157 Large White). Animals were genotyped using the Illumina SNP60K Beadchip and assigned to either a training data set to estimate the genetic parameters and SNP effects, or to a validation data set to assess the prediction accuracy. Models MA and MAD applied random regression on SNP genotypes and were implemented in the program Bayz. The additive heritability of DG across the three populations and the two models was very similar at approximately 0.26. The proportion of phenotypic variance explained by dominance effects ranged from 0.04 (Large White) to 0.11 (Pietrain), indicating that importance of dominance might be breed‐specific. Prediction accuracies were higher when predicting phenotypes using total genetic values (sum of breeding values and dominance deviations) from the MAD model compared to using breeding values from both MA and MAD models. The highest increase in accuracy (from 0.195 to 0.222) was observed in the Pietrain, and the lowest in Large White (from 0.354 to 0.359). Predicting phenotypes using total genetic values instead of breeding values in purebred data improved prediction accuracy and reduced the bias of genomic predictions. Additional benefit of the method is expected when applied to predict crossbred phenotypes, where dominance levels are expected to be higher.     

Gene action and heterosis in lifetime traits of Friesian × Sahiwal crosses

Gene action and heterosis in seven lifetime traits of Holstein‐Friesian × Sahiwal crosses was studied by fitting genetic additive—dominance and Dickerson, Sheridan models. The additive—dominance model is found adequate for all the traits except ‘average milk yield per day of total productive life’. The heterosis measured from mid‐parent varied from 15% in ‘total life’ to 108% in ‘milk yield from all available lactations’. Heterosis measured from superior parent ranged from 6% in ‘total milk yield of first three lactations’ to 75% in ‘milk yield from all available lactations’. The larger heterosis observed in some of the lifetime traits is attributed as the direct consequence of longer productive life in F₁s compared to parental breeds. On the basis of the predicted performance of some of the nonavailable grades like 2/3 exotic, creation of synthetics is mooted.     

Genotyping strategies of selection candidates in livestock breeding programmes

Benefits of genomic selection (GS) in livestock breeding operations are well known particularly where traits are sex‐limited, hard to measure, have a low heritability and/or measured later in life. Sheep and beef breeders have a higher cost:benefit ratio for GS compared to dairy. Therefore, strategies for genotyping selection candidates should be explored to maximize the economic benefit of GS. The aim of the paper was to investigate, via simulation, the additional genetic gain achieved by selecting proportions of male selection candidates to be genotyped via truncation selection. A two‐trait selection index was used that contained an easy and early‐in‐life measurement (such as post‐weaning weight) as well as a hard‐to‐measure trait (such as intra‐muscular fat). We also evaluated the optimal proportion of female selection candidates to be genotyped in breeding programmes using natural mating and/or artificial insemination (NatAI), multiple ovulation and embryo transfer (MOET) or juvenile in vitro fertilization and embryo transfer (JIVET). The final aim of the project was to investigate the total dollars spent to increase the genetic merit by one genetic standard deviation (SD) using GS and/or reproductive technologies. For NatAI and MOET breeding programmes, females were selected to have progeny by 2 years of age, while 1‐month‐old females were required for JIVET. Genomic testing the top 20% of male selection candidates achieved 80% of the maximum benefit from GS when selection of male candidates prior to genomic testing had an accuracy of 0.36, while 54% needed to be tested to get the same benefit when the prior selection accuracy was 0.11. To achieve 80% of the maximum benefit in female, selection required 66%, 47% and 56% of female selection candidates to be genotyped in NatAI, MOET and JIVET breeding programmes, respectively. While JIVET and MOET breeding programmes achieved the highest annual genetic gain, genotyping male selection candidates provides the most economical way to increase rates of genetic gain facilitated by genomic testing.     

Periodic rotational crosses. II. Optimizing breed and heterosis use

Choice of the optimal number and sequence of breeds in a periodic cross was determined by comparing the trade-off between increased utilization of breed differences and decreased utilization of heterosis. It was shown that the change in mean efficiency resulting from adding the next best breed to the best conventional n-breed rotation is always less than the change in efficiency predicted from the increase in heterosis. Periodic rotations were generally optimized by decreasing the proportion of poorer performing breeds in the rotation. However, efficiency of periodic rotations can exceed that of the better breed even when the difference in additive breed effects for efficiency is almost twice the effect of heterosis on efficiency. The periodic rotation that was optimal also tended to have the lowest inter-generational variance. It was suggested that inter-generational variances of component traits, which are not necessarily minimized when crosses are selected on a combined efficiency trait, can be considered by including inter-generational variance in an index or by introducing maximum thresholds.     

Synthesis of direct and maternal genetic components of economically important traits from beef breed-cross evaluations

Published information on relative performance of beef breed crosses was used to derive combined estimates of purebred breed values for predominant temperate beef breeds. The sources of information were largely from the United States, Canada, and New Zealand, although some European estimates were also included. Emphasis was on maternal traits of potential economic importance to the suckler beef production system, but some postweaning traits were also considered. The estimates were taken from comparison studies undertaken in the 1970s, 1980s and 1990s, each with representative samples of beef breeds used in temperate agriculture. Weighting factors for breed-cross estimates were derived using the number of sires and offspring that contributed to that estimate. These weights were then used in a weighted multiple regression analysis to obtain single purebred breed effects. Both direct additive and maternal additive genetic effects were estimated for preweaning traits. Important genetic differences between the breeds were shown for many of the traits. Significant regression coefficients were estimated for the effect of mature weight on calving ease, both maternal and direct additive genetic, survival to weaning direct, and birth weight direct. The breeds with greater mature weight were found to have greater maternal genetic effects for calving ease but negative direct genetic effects on calving ease. A negative effect of mature weight on the direct genetic effect of survival to weaning was observed. A cluster analysis was done using 17 breeds for which information existed on nine maternal traits. Regression was used to predict breed-cross-specific heterosis using genetic distance. Only five traits, birth weight, survival to weaning, cow fertility, and preweaning and postweaning growth rate had enough breed-cross-specific heterosis estimates to develop a prediction model. The breed biological values estimated provide a basis to predict the biological value of crossbred suckler cows and their offspring.     

Effect of mate selection on fuzzy selective mating criteria in closed dairy multiple ovulation and embryo transfer nucleus programs

In order to control rates of response and inbreeding, mate selection using fuzzy selective mating criteria (FMC) was investigated in adult multiple ovulation and embryo transfer nucleus schemes for dairy cattle. Stochastic simulation was used to model the closed nucleus scheme. This mate selection was examined in four alternative mating and male selection schemes: (i) a hierarchical scheme; (ii) a hierarchical sibship scheme (two males per sibship); (iii) a factorial scheme (two sires per dam); and (iv) a factorial sibship scheme (two males per sibship and two sires per dam). Genetic response and inbreeding rate tended to be reduced by increasing the trade-off parameter of FMC between the expected breeding value and inbreeding of progeny. Inbreeding rates in all schemes were reduced by reducing the variance of family size through selection and the average coancestry of mating pairs through mate allocation.     

The crossbred sire: experimental results for sheep

Crossbreeding of sheep is practiced to exploit simultaneously the use of additive and nonadditive genetic effects. The goal is to achieve optimal levels of performance appropriate for defined systems of sheep production and marketing. Although the beneficial effects of individual and maternal heterosis on sheep production have been well documented and widely implemented, considerably less is known about the effects of paternal heterosis. Limited evidence suggests that crossbred rams are more sexually aggressive and exhibit greater testicular growth than do purebred rams. Average estimates of paternal heterosis effects were 1.4, -.7 and 2.3% for seasonal fertility, prolificacy and preweaning survival, respectively. The average effect of paternal heterosis on fertility during spring breeding was 29.5%. Progeny of crossbred and purebred sires were similar in birth weight, weaning weight and postweaning growth rate and in phenotypic variation for these growth traits. However, favorable paternal heterosis effects need not exist to warrant the use of crossbred sires. Composite or F1 sires can be used as an effective method to manage the composition of additive breed effects. For example, varying proportions of germ plasm from highly prolific breeds such as the Finnsheep and Romanov can be realized through the use of crossbred sires to set reproductive rates at desired levels. Crossbred sires may be used to a greater extent to optimize additive breed effects than to exploit effects of paternal heterosis. The role of composite breeds in managing both additive and nonadditive effects is discussed.     

Additive, dominance, and epistatic loss effects on preweaning weight gain of crossbred beef cattle from different Bos taurus breeds

(Co)variance components, direct and maternal breed additive, dominance, and epistatic loss effects on preweaning weight gain of beef cattle were estimated. Data were from 478,466 animals in Ontario, Canada, from 1986 to 1999, including records of both purebred and crossbred animals from Angus, Blonde d'Aquitaine, Charolais, Gelbvieh, Hereford, Limousin, Maine-Anjou, Salers, Shorthorn, and Simmental breeds. The genetic model included fixed direct and maternal breed additive, dominance, and epistatic loss effects, fixed environmental effects of age of the calf, contemporary group, and age of the dam x sex of the calf, random additive direct and maternal genetic effects, and random maternal permanent environment effects. Estimates of direct and maternal additive genetic, maternal permanent environmental and residual variances, expressed as proportions of the phenotypic variance, were 0.32, 0.20, 0.12, and 0.52, respectively. Correlation between direct and maternal additive genetic effects was -0.63. Breed ranking was similar to previous studies, but estimates showed large SE. The favorable effects of direct and maternal dominance (P < 0.05) on preweaning gain were equivalent to 1.3 and 2.3% of the phenotypic mean of purebred calves, respectively. The same features for direct and maternal epistatic loss effects were -2.2% (P < 0.05) and -0.1% (P > 0.05). The large SE of breed effects were likely due to multicollinearity among predictor variables and deficiencies in the dataset to separate direct and maternal effects and may result in a less reliable ranking of the animals for across breed comparisons. Further research to identify the causes of the instability of estimates of breed additive, dominance, and epistatic loss genetic effects, and application of alternative statistical methods is recommended.     

The impact of using old germplasm on genetic merit and diversity—A cattle breed case study

Artificial selection and high genetic gains in livestock breeds led to a loss of genetic diversity. Current genetic diversity conservation actions focus on long‐term maintenance of breeds under selection. Gene banks play a role in such actions by storing genetic materials for future use and the recent development of genomic information is facilitating characterization of gene bank material for better use. Using the Meuse‐Rhine‐Issel Dutch cattle breed as a case study, we inferred the potential role of germplasm of old individuals for genetic diversity conservation of the current population. First, we described the evolution of genetic merit and diversity over time and then we applied the optimal contribution (OC) strategy to select individuals for maximizing genetic diversity, or maximizing genetic merit while constraining loss of genetic diversity. In the past decades, genetic merit increased while genetic diversity decreased. Genetic merit and diversity were both higher in an OC scenario restricting the rate of inbreeding when old individuals were considered for selection, compared to considering only animals from the current population. Thus, our study shows that gene bank material, in the form of old individuals, has the potential to support long‐term maintenance and selection of breeds.     

Comparison of production systems for efficient use of indigenous pig breeds in developing countries

Conserving pig genetic resources and improving their productivity is important to increase returns over investment in developing countries. The purebred, first‐cross, rotational cross and backcross matings representing production systems based on pig breeds indigenous to the country and exotic pig breeds were investigated. The number of pigs in the nucleus and commercial herds necessary to produce a defined quantity of pork was considered. The amount of heterosis between the indigenous and exotic breeds, superiority in meat production, and degree of inferiority in reproductive performance of the exotic breed compared with that of the indigenous breed were investigated. The number of breeding pigs in the whole system was in the following order: pure breeding (PB) > first‐cross (F1) > rotational cross (RC) > backcross (BC) systems. The number of breeding pigs in the nucleus herds of the RC and BC systems was smaller than that in the nucleus herds of the PB and F1 systems. The degree of inferiority in reproductive performance of the exotic breed compared with that of the indigenous breed affected the efficiency of the production system.     

Genetic correlations between production with disease resistance and fertility in dairy cattle and consequences for total merit selection

Twelve years of data from progeny‐test results of 1486 Swedish Red and White (SRB) and 756 Swedish Black and White (SLB) AI bulls were analysed to provide estimates of genetic correlations between yield of protein and three health‐ and fertility traits. For both breeds, the correlations were unfavourable (r_G= —0.13 to —0.37). The effects of negative genetic correlations (compared to a situation with genetic correlations with zero values) on the b‐values in the total merit index of bulls were rather small but the effects on the estimated genetic gain were large. The effect of not including health and fertility traits in the index, although included in the breeding goal, resulted in a 9–10% reduced accuracy of estimated breeding values for total merit and thus a corresponding loss in total gain.     

Maternal heterosis and grandmaternal effects in beef cattle: postweaning growth and carcass traits

Crossbred steer and heifer progeny from 5-, 6- and 7-yr-old dams produced in a four-breed diallel crossing experiment involving the Brown Swiss, Red Poll, Hereford and Angus maternal grandsires and maternal granddams were evaluated for postweaning growth and carcass traits to estimate breed mean maternal heterosis, maternal heterosis for specific breed cross females, average maternal heterosis for all crosses, breed grandmaternal effects and net breed effects in crosses. All progeny evaluated were born in 1979 and 1980 and were sired by 7/8 or 15/16 Simmental bulls. Average maternal heterosis was significant for 200-d weight in heifers but not in steers and was not significant for final weight (444-d) in either heifers or steers. The effects of maternal heterosis on postweaning growth were not important. Differences among breeds in mean maternal heterosis values were small for growth-related traits. Breeds did not differ (P greater than .05) in grandmaternal effects for growth-related traits; Brown Swiss tended to be highest, Red Poll lowest, with Hereford and Angus intermediate. Differences in net breed effects in crosses favored Brown Swiss over the three other breeds and were generally significant for growth traits. Average maternal heterosis, though generally positive, was not significant for carcass traits on either an age-constant or weight-constant basis. Differences among breeds were small in grandmaternal effects, specific heterosis and net effects in crosses for carcass traits associated with both weight or composition; generally the Brown Swiss breed was favored on carcass traits associated with weight in the age-constant analysis and generally had a higher lean-to-fat ratio than the three other breeds in both the age-constant and weight-constant analyses.