Tormod Ådnøy

Effects on Production Traits of Haplotypes Among Casein Genes in Norwegian Goats and Evidence for a Site of Preferential Recombination

In goat milk the most abundant proteins are the casein genes, CSN1S1, CSN2, CSN1S2, and CSN3. Mutations have been identified within these genes affecting the level of gene expression, and effects on milk production traits have been reported. The aim of this study was to detect polymorphisms (SNPs) in the casein genes of Norwegian goats, resolve haplotype structures within the loci, and assess the effect of these haplotypes on milk production traits. Four hundred thirty-six Norwegian bucks were genotyped for 39 polymorphic sites across the four loci. The numbers of unique haplotypes present in each locus were 10, 6, 4, and 8 for CSN1S1, CSN2, CSN1S2, and CSN3, respectively. The effects of the CSN1S1 haplotypes on protein percentage and fat kilograms were significant, as were the effects of CSN3 haplotypes on fat percentage and protein percentage. A deletion in exon 12 of CSN1S1, unique to the Norwegian goat population, explained the effects of CSN1S1 haplotypes on fat kilograms, but not protein percentage. Investigation of linkage disequilibrium between all possible pairs of SNPs revealed higher levels of linkage disequilbrium for SNP pairs within casein loci than for SNP pairs between casein loci, likely reflecting low levels of intragenic recombination. Further, there was evidence for a site of preferential recombination between CSN2 and CSN1S2. The value of the haplotypes for haplotype-assisted selection (HAS) is discussed.

Effect of milk- and TRIS-based extenders on the fertility of sheep inseminated vaginally once or twice with liquid semen

We studied the influence of two different extenders, a milk-based versus a TRIS-based extender, using a split-sample technique, on fertility after single and double vaginal inseminations in natural estrous in Norwegian Crossbred ewes. Semen from 21 Norwegian Crossbred rams, all aged approximately 0.5 years, was used for AI of totally 561 Norwegian Crossbred ewes housed at 37 different farms. The farmers performed the inseminations themselves. The ewes were allocated to four parallel groups based on the two extenders and single or double inseminations (2 x 2). The farmers were recommended to inseminate the ewes between 12 and 24 h after detection of natural standing estrous. Vaginal insemination with cooled liquid semen diluted in the milk-based extender resulted in a statistically significant (P<0.01) better fertility of about 10% units both as 25-day NR (non return rate)-and lambing rates, compared with semen diluted in the TRIS-based extender. Double inseminations gave significantly higher (P=0.03) fertility results for both extenders expressed as 25-day NR results, but was not quite statistically significant when expressed as lambing rates (P=0.06) compared with single insemination. The overall 25-day NR results for the milk-based extender (66.4%) after single inseminations is in accordance with both the national results (67.1%) based on vaginal inseminations of 11,377 ewes, as well as with the results from a previous study in the same region achieving a 25-day NR results of 63.3%. In conclusion, liquid ram semen diluted in a milk-based extender and vaginally inseminated once in natural heat, with a semen dose of 150 x 10(6) spermatozoa, gave acceptable fertility results and is to be recommended as the method of choice in Norway.

Effect of vaginal and cervical deposition of semen on the fertility of sheep inseminated with frozen-thawed semen

The effect of vaginal and cervical deposition of frozen-thawed semen on the fertility of sheep was tested in a field trial in which 543 Norwegian crossbred ewes aged between six months and five-and-a-half years from 10 farms were inseminated after natural oestrus. Cervical insemination with 200 x 106 spermatozoa resulted in 25-day non-return and lambing rates of 75·4 and 72·7 per cent, respectively, and vaginal insemination gave rates of 71·3 and 67·4 per cent; the cervical inseminations produced significantly higher lambing rates (P=0·04). There were significant differences between the lambing rates for different rams (P=0·006) and different farmers (P=0·003), and there was a significant interaction between farmer and deposition site (P=0·03). After vaginal insemination fertility was encouragingly high, but the results varied with the farmer, and different flock and management conditions.

Effect of deposition site and sperm number on the fertility of sheep inseminated with liquid semen.

The effect of the deposition site and the numbers of sperm on the fertility of sheep was tested in a field trial in which 1292 Norwegian crossbred ewes aged between six months and five-and-a-half years from 52 farms were inseminated with liquid semen after natural oestrus. Cervical insemination with 150 x 106 and 75 x 106 spermatozoa resulted in 25-day non-return rates of 63.7 and 56.1 per cent, and vaginal insemination gave non-return rates of 63.3 and 56.6 per cent, respectively. There was no significant difference between the cervical and vaginal inseminations, but the inseminations with 150 x 106 spermatozoa gave significantly higher non-return rates (P=0.004). There were significant differences between the non-return rates for different rams (P<0.0001) and farmers (P=0.0002) but the age of the ewe had no significant effect.

Casein SNP in Norwegian goats: additive and dominance effects on milk composition and quality

BackgroundThe four casein proteins in goat milk are encoded by four closely linked casein loci (CSN1S1, CSN2, CSN1S2 and CSN3) within 250 kb on caprine chromosome 6. A deletion in exon 12 of CSN1S1, so far reported only in Norwegian goats, has been found at high frequency (0.73). Such a high frequency is difficult to explain because the national breeding goal selects against the variants effect.MethodsIn this study, 575 goats were genotyped for 38 Single Nucleotide Polymorphisms (SNP) located within the four casein genes. Milk production records of these goats were obtained from the Norwegian Dairy Goat Control. Test-day mixed models with additive and dominance fixed effects of single SNP were fitted in a model including polygenic effects.ResultsSignificant additive effects of single SNP within CSN1S1 and CSN3 were found for fat % and protein %, milk yield and milk taste. The allele with the deletion showed additive and dominance effects on protein % and fat %, and overdominance effects on milk quantity (kg) and lactose %. At its current frequency, the observed dominance (overdominance) effects of the deletion allele reduced its substitution effect (and additive genetic variance available for selection) in the population substantially.ConclusionsThe selection pressure of conventional breeding on the allele with the deletion is limited due to the observed dominance (overdominance) effects. Inclusion of molecular information in the national breeding scheme will reduce the frequency of this deletion in the population.

The effect of initial weight of the ewe on later reproductive effort in domestic sheep (Ovis aries)

In ungulates, females typically need to reach a critical threshold body weight in order to reproduce. Females born in ‘poor’ years may lose 1 year of reproduction, as they reach the critical body mass 1 year later. Some studies report also a lasting effect of low initial body weight into prime age, while several other studies have shown that nearly all prime-age females ovulate or are pregnant. However, the quality of the offspring (as measured by offspring body weight) is often not considered. We tested whether female ungulates with an initially low body weight (at around weaning) reach the same reproductive output as females with higher initial weights, and thus whether or not they are able to compensate for their initial low weight. Data on body weight of 123 533 lambs of domestic sheep Ovis aries from 73 299 litters that in turn derived from 32 359 different ewes were used. All ewes and lambs had been free-ranging on outlying pastures in Norway during the entire summer season. Ewes with an initially low body weight produced smaller offspring throughout their lifespan. Ewes with an initially low body weight also produced fewer lambs at first and second parturition, but lifetime number of offspring was not related to ewes’ initial weight. We concluded that bigger was better: the smaller offspring of ewes with a low initial weight demonstrate a lasting cost on reproductive effort, with consequences for animal production systems and management of wild ungulates.

The Norwegian sheep breeding scheme: description, genetic and phenotypic change

The developments in Norwegian sheep breeding since the early 1990s are reviewed. For the largest breeding population, the Norwegian White Sheep, results are presented for both genetic and phenotypic changes. Of the nine traits that make up the aggregate genotype, the largest gain per year, in per cent of the corresponding phenotypic average, was found for carcass grade (1.66%) and carcass weight (0.99%), number of lambs born at 1, 2 and 3 years of age (0.32% to 0.60%) and the maternal effect on weaning weight (0.26%). For fat grade, a genetic deterioration was estimated. This may be due to the too small weighting of this trait in the aggregate genotype and the true genetic parameters being somewhat different from the estimates in the prediction of breeding values. For lamb as well as ewe fleece weight, genetic change was close to zero - interpreted as mainly a correlated response to other traits in the aggregate genotype. Data for the two traits of fleece weight were, respectively, selected and few. Thus, phenotypic change was calculated for all traits except for fleece weight, and in addition for number of lambs at weaning, being indirectly selected for through number of lambs born. For all traits, with the exception of fat grade, advantageous phenotypic change was estimated. For weaning and carcass weight, the phenotypic change was less than the genetic change, while the opposite was observed for carcass and fat grade and number of lambs born. The latter traits can be more easily controlled by environmental actions, and the results thus exemplify the interdependency between environmental and genetic change.

Differences in choice of diet between sheep breeds grazing mountain pastures in Norway

In the present study, the foraging behaviour of ewes from three common Norwegian sheep breeds; the long-tailed Dala sheep, and the two short-tailed breeds Spæl sheep and Norwegian Fur sheep was studied. Based on difference in body size and digestive anatomy, we expected the lighter short-tailed ewes to browse more on woody plants than the Dala ewes. The experiments were conducted during two consecutive summers within the flocks’ usual grazing areas, and we sampled the proportion of grazing time that ewes spent feeding on woody plant species. We tested the hypothesis by applying generalized linear mixed models to the feeding behaviour data. As predicted, the short-tailed ewes spent substantially more of their grazing time browsing on trees, bushes and heather compared to the Dala ewes. We discuss the results in the context of maintaining the, now threatened, pastoral landscapes of mountainous regions of Norway; short-tailed breeds should be more effective than the Dala breed in reducing tree and bush encroachment, but more detailed studies are needed to quantify the breed differences in this respect.

Allele Interaction – Single Locus Genetics Meets Regulatory Biology

Background Since the dawn of genetics, additive and dominant gene action in diploids have been defined by comparison of heterozygote and homozygote phenotypes. However, these definitions provide little insight into the underlying intralocus allelic functional dependency and thus cannot serve directly as a mediator between genetics theory and regulatory biology, a link that is sorely needed. Methodology/Principal Findings We provide such a link by distinguishing between positive, negative and zero allele interaction at the genotype level. First, these distinctions disclose that a biallelic locus can display 18 qualitatively different allele interaction sign motifs (triplets of +, – and 0). Second, we show that for a single locus, Mendelian dominance is not related to heterozygote allele interaction alone, but is actually a function of the degrees of allele interaction in all the three genotypes. Third, we demonstrate how the allele interaction in each genotype is directly quantifiable in gene regulatory models, and that there is a unique, one-to-one correspondence between the sign of autoregulatory feedback loops and the sign of the allele interactions. Conclusion/Significance The concept of allele interaction refines single locus genetics substantially, and it provides a direct link between classical models of gene action and gene regulatory biology. Together with available empirical data, our results indicate that allele interaction can be exploited experimentally to identify and explain intricate intra- and inter-locus feedback relationships in eukaryotes.

Comparison of fertility results after vaginal insemination using different thawing procedures and packages for frozen ram semen

BackgroundThe effect of different thawing procedures for ram semen frozen in minitubes and mini straws on the fertility of sheep was tested in a field trial.MethodsAltogether, 719 Norwegian Crossbred ewes, aged between six months and six-and-a-half years from 8 farms, were inseminated vaginally in natural oestrus with frozen-thawed semen. Minitubes were thawed at 70°C for 8 sec (T70) and mini straws either at 50°C for 9 sec (S50) or at 35°C for 12 sec (S35).ResultsVaginal insemination with 200 × 106 spermatozoa resulted in 25-days non-return rates of 63.2, 59.6, and 62.5% (overall 61.8%), respectively, and lambing rates of 56.8, 55.0, and 59.2% (overall 57.0%), respectively. No significant effect on fertility (as 25-days non-return- or lambing rate) was seen for straw type/thawing temperature (P = 0.5/0.5), but semen filled in mini straws and thawed at 35°C resulted numerically in the highest lambing rate (59.2%). A significant effect was, however, seen for farmer (P = >0.0001/>0.0001) and ram (P = 0.009/0.002). Moreover, age of the ewes had a significant effect on the NR rate (0.007), but not on lambing rate (P = 0.2).ConclusionA vaginal deposition of frozen ram semen containing approximately 200 × 106 spermatozoa, filled in mini straws and thawed at 35°C is a simplified technique that under field conditions and used on a do-it-yourself regime gives acceptable lambing rates in Norway.