M. L. Hebart

Opportunities for genetic improvement of lamb survival

F. D. Brien, M. L. Hebart, D. H. Smith, J. E. Hocking Edwards, J. C. Greeff, K. W. Hart, G. Refshauge, T. L. Bird-Gardiner, G. Gaunt, R. Behrendt, M. W. Robertson, G. N. Hinch, K. G. Geenty and J. H. J. van der Werf

A review of the genetic and epigenetic factors affecting lamb survival

Poor lamb survival pre-weaning is a major source of reproductive inefficiency in Australian sheep flocks. While nutrition and management options have been extensively researched and promoted to improve lamb survival, the present review focuses on the prospects for obtaining genetic gain and helps identify selection strategies for boosting such gains to improve overall reproductive efficiency in the Australian sheep industry. Estimated heritability for lamb survival using linear model analysis is low, although use of threshold models suggests that heritability could be higher, which, if true, could help explain the substantial genetic gains obtained in long-term selection experiments. Epigenetic mechanisms may hinder selection and quantitative trait-loci identification through confounding and/or masking genetic variances and co-variances. With sufficient information, these effects could be considered in genetic evaluations by identifying those components that are amenable to selection. Regarding indirect selection, finding effective criteria for improving lamb survival has proved elusive. Most measures of maternal behaviour, temperament and lambing difficulty researched are poorly correlated genetically with lamb survival. Of lamb behaviours and thermo-genic indicators studied, latency to bleat following handling by humans is moderately genetically correlated with lamb survival, as is neonatal rectal temperature. Industry application remains to be adequately explored for the more promising of these measures. Finally, in lieu of direct selection for lamb survival, there is merit in selecting for multiple-rearing ability or its equivalent, possibly with additional selection criteria for lamb survival and reproductive efficiency.

Genesis, design and methods of the Beef CRC Maternal Productivity Project

Australian seedstock cattle breeders have expressed concerns that while there has been genetic improvement in feedlot and abattoir performance of cows, it could have led to a decline in maternal productivity, especially under variable nutritional conditions. This paper describes a substantial project with two components designed to address these issues. The first sub-project was to monitor bodyweight and composition of 7760 young Angus and Hereford cows as they experience variable physiological states (pregnancy and lactation) and seasons. This was conducted on large numbers in seedstock herds. The second sub-project was to monitor more regularly bodyweight, body composition, and calf rearing performance of 500 Angus cows that are genetically divergent for either fat or residual feed intake at two research centres. This also included two levels of nutrition and recording of weekly feed intake of small groups of cows for at least three parities to allow reporting of genotype × nutrition effects on maternal productivity and efficiency. Results from the project are reported in a series of papers with each one having a defined focus.

Divergent breeding values for fatness or residual feed intake in Angus cattle. 5. Cow genotype affects feed efficiency and maternal productivity

Cow bodyweight gain, calf weaning weight, feed intake and maternal productivity of 500 Angus cows, in 64 replicate groups, were measured over three parities at two locations (Struan and Vasse) as part of the Beef CRC Maternal Productivity Project. The cows were sourced as heifers from the top and bottom 10% of BREEDPLAN Rib Fat EBV (High-Fat and Low-Fat), and from High and Low residual feed intake (RFI) selection lines (High-RFI and Low-RFI). Each of the four genotypes were run under High- and Low-Nutrition (measured as feed on offer) at both sites. The High-Fat cows were 7% more efficient at producing weaner calves under Low-Nutrition than were the Low-Fat cows. This was driven primarily by the 4% difference between the lines in weaning rate. When weaning rate differences were accounted for (as covariate), there was no difference between the Fat lines in the efficiency of weaner weight production. When the weight gain of the cow was included as an output in addition to calf weaning weight, there was also no difference between the Fat lines in efficiency. Low-RFI cows were always more efficient at producing weaner calves than were the High-RFI cows. This was primarily driven through a 7% reduction in annual feed intake (across both nutrition treatments). However, the Low-RFI cows were leaner, had 6.3% lower weaning rate and calved on average 5.4 days later than did the High-RFI cows. Furthermore, the largest differences in feed intake were in spring when feed availability is greatest. In the context of the results herein, a balanced breeding program should include selection for improved reproduction and low RFI.

Divergent breeding values for fatness or residual feed intake in Angus cattle. 1. Pregnancy rates of heifers differed between fat lines and were affected by weight and fat

The pregnancy rate of heifers affects the efficiency and profitability of beef herds. Heifers extreme in rib fatness (Fat) or post-weaning residual feed intake (RFI) estimated breeding values (EBVs) were evaluated for their pregnancy rates at two locations in the southern agricultural regions of Australia (Struan and Vasse) as part of the Beef Cooperative Research Centre Maternal Productivity Project. Heifers divergent in Fat (High-Fat and Low-Fat) had differences in fat depth pre-joining at the 12/13th rib (4.4 mm vs 3.5 mm) and P8 rump site (6.1 mm vs 4.8 mm). This was associated with significant differences in pregnancy rates over a 9-week joining period (91.5% vs 83.0%) and an even larger difference when calculated over a 6-week joining period (77.3% vs 65.0%). Heifers divergent in RFI (Vasse only) also differed in rib fat (7.6 mm vs 6.4 mm) and P8 fat (11.0 vs 9.2 mm), but not significantly in pregnancy rates between the two RFI (High-RFI and Low-RFI) genotypes following a 9-week (92.4% vs 88.5%) or 6-week (81.2% vs 73.7%) joining period. The phenotypic analysis of the Fat and RFI heifers together indicated that weight and fat depth were the largest contributing factors to variation in pregnancy rates, and age and pre-joining weight gain were not significant. These phenotypic characteristics indicated that producers can manage heifers to particular weight and fat combinations to improve heifer conception rates. Associations of BREEDPLAN EBVs with heifer fertility showed that a shorter days-to-calving EBV had the biggest impact (P < 0.001) on heifer pregnancy rates and rib fat and scrotal size EBVs were close to significant (P < 0.10).

Divergent genotypes for fatness or residual feed intake in Angus cattle. 3. Performance of mature cows

This experiment evaluated the productivity of 500 Angus cows that differed in genetic merit for either subcutaneous rib fat depth (Fat) or residual feed intake (RFI) based on estimated breeding values (EBVs) and managed under two levels of nutrition. Reproductive rate over four calving opportunities in mature cows and growth performance of progeny to weaning was assessed. Level of nutrition significantly affected all body composition traits for both Fat and RFI line cows. Cows on High-Nutrition were 14–16% heavier (P < 0.001) than those on Low-Nutrition. Differences in EBVs for fatness were reflected in phenotypic fatness at maturity. High-RFI line cows were fatter for both scanned rump (P8) and rib (RIB) fat depth relative to their Low-RFI contemporaries. Of those cows that were lactating, there was no significant effect of line or nutrition on pregnancy rate or days to calving (DC). There was, however, a trend (P < 0.1) in the Low-Fat line cows towards longer DC compared with the High-Fat line cows. There was no significant effect of either line or nutrition on calf birthweight. Calves with mothers on High-Nutrition were 8% heavier at weaning (P < 0.001) than those on Low-Nutrition. Lower EBVs for RFI was associated with higher 200-day growth EBV and heavier calves at weaning. Current carcass BREEDPLAN EBVs can be used to select for changes in cow body composition if desired. In this experiment, Angus cows selected for lower RFI or with below-average fatness EBV and had raised a calf at every previous opportunity were not compromised in pregnancy rate or DC at maturity under varying nutrition such as can be experienced during normal seasonal conditions in southern Australia. However, selection for lower RFI was associated with lower weaning rate (P < 0.05), which warrants further investigation to confidently predict the implications for commercial cattle production.

Phenotypic and genotypic analysis of a barebreech trait in Merino sheep as a potential replacement for surgical mulesing.

The potential for adopting a genetic solution to protect sheep from blowfly strike on the breech was investigated in a flock of sheep that contained several animals expressing a trait characterised by low wool coverage over the breech and through a wide channel from the anus to the udder or scrotum. A scoring system (1, bare to 5, woolly) was developed and used to determine the heritability of the trait and its phenotypic and genetic correlations with other traits of importance in a sheep enterprise. In comparison to animals with woolly breeches, the skin in the breech of animals with a low bareness score was characterised by a low density of follicles producing short, medullated fibres, with histological evidence of immune rejection and follicular atrophy. The bareness score of progeny was influenced by the score of their respective sires suggesting a strong genetic component. The heritability of bareness score was moderate to high (h2 = 0.45 ± 0.02, 0.53 ± 0.01 and 0.38 ± 0.02 at lamb, hogget and adult ages, respectively). The lactation status and age of ewes influenced their bareness score, resulting in a low repeatability (0.42) of the trait between ages in females. Genetic correlations between bareness score and most other economically important traits were low. The weight of belly wool and the weight of skirtings was genetically related to bareness score (rg = +0.52 and +0.48 respectively), indicating that animals with barer breeches tend genetically towards lighter belly wool weights and lower weight of skirtings at wool classing. Selection and breeding for bareness score should achieve relatively rapid progress towards fixing the trait in a flock and without adverse effects on other important traits. Caution should be exercised in extrapolating these results to other bloodlines and environments where genetic mechanisms or environmental influences may be different.

Divergent genotypes for fatness or residual feed intake in Angus cattle. 2. Body composition but not reproduction was affected in first-parity cows on both low and high levels of nutrition

This paper reports a subset of results from the Beef Cooperative Research Centre-funded Maternal Productivity Project. This research aimed to describe the response of Angus cows of different and divergent genotypes to variable nutritional environments over five breeding seasons. Cows selected for a divergence in either fat depth (HFat vs LFat) or residual feed intake (RFI: HRFI vs LRFI) based on mid-parent estimated breeding values (EBV) for those traits were allocated in replicate groups to either high or low nutritional treatments at two different sites, namely the Vasse Research Centre in Western Australia and the Struan Research Centre in South Australia. The traits reported in this paper include output traits (birth and weaning weight of calves, liveweight change of cows), change traits (change in Rib Fat, P8 fat, eye muscle area and liveweight between specified time points) and reproductive traits [pregnancy rates, percentage calves born alive and days to calving at the days to calving at the second calving opportunity (DC2)]. Having had their first calf, the vulnerability of these young cows to nutritional restriction and how it may adversely affect rebreeding was examined. HFat and HRFI cows were fatter, heavier and had greater eye muscle area than LFat and LRFI, respectively, at all times during the breeding cycle on both levels of nutrition. There was no difference in either days-to-calving or pregnancy rates after the second mating between genotypes. Equally, nutritional treatment had no effect on these traits in this cohort of cows. There was evidence for an implied genetic correlation between Rib Fat EBV, DC2 and pregnancy rates of –0.38 that suggests that selection for leanness may result in reduced fertility of the herd but the effect was not significant herein. As long as producers record the phenotype for both traits and select cows with favourable DC2 as well as low fatness, these problems can be avoided, owing to only 22% of variation in pregnancy rates being explained by DC2 and Rib Fat EBV. Producers can largely be confident that selection for leanness, or increased feed efficiency, has little impact on productivity as long as cows are in adequate body condition to remain healthy and productive.

Wool fibre crimp is determined by mitotic asymmetry and position of final keratinisation and not ortho- and para-cortical cell segmentation.

Crimp, a distinguishing feature of sheep fibres, significantly affects wool value, processing and final fabric attributes. Several explanations for fibre bending have been proposed. Most concentrate on relative differences in the physicochemical properties of the cortical cells, which comprise the bulk of the fibre. However, the associations between cortical properties and fibre crimp are not consistent and may not reflect the underlying causation of fibre curvature (FC). We have formulated a mechanistic model in which fibre shape is dictated primarily by the degree of asymmetry in cell supply from the follicle bulb, and the point at which keratinisation is completed within the follicle. If this hypothesis is correct, one would anticipate that most variations in fibre crimp would be accounted for by quantitative differences in both the degree of mitotic asymmetry in follicle bulbs and the distance from the bulb to the point at which keratinisation is completed. To test this hypothesis, we took skin biopsies from Merino sheep from sites producing wool differing widely in fibre crimp frequency and FC. Mitotic asymmetry in follicle bulbs was measured using a DNA-labelling technique and the site of final keratinisation was defined by picric acid staining of the fibre. The proportion of para- to ortho-cortical cell area was determined in the cross-sections of fibres within biopsy samples. Mitotic asymmetry in the follicle bulb accounted for 0.64 (P < 0.0001) of the total variance in objectively measured FC, while the point of final keratinisation of the fibre accounted for an additional 0.05 (P < 0.05) of the variance. There was no association between ortho- to para-cortical cell ratio and FC. FC was positively associated with a subjective follicle curvature score (P < 0.01). We conclude that fibre crimp is caused predominantly by asymmetric cell division in follicles that are highly curved. Differential pressures exerted by the subsequent asymmetric cell supply and cell hardening in the lower follicle cause fibre bending. The extent of bending is then modulated by the point at which keratinisation is completed; later hardening means the fibre remains pliable for longer, thereby reducing the pressure differential and reducing fibre bending. This means that even highly asymmetric follicles may produce a straight fibre if keratinisation is sufficiently delayed, as is the case in deficiencies of zinc and copper, or when keratinisation is perturbed by transgenesis. The model presented here can account for the many variations in fibre shape found in mammals.

Perinatal wool follicle attrition coincides with elevated perinatal circulating cortisol concentration in Merino sheep

The development of the follicle population in Merino sheep skin was studied from 30 days pre-partum to 112 days post-partum in single- and twin-born Merino lambs. The total number of primary follicles estimated from primary follicle density and skin surface area did not change over this period, while secondary follicle number increased to Day 28 post-partum in singles and Day 56 post-partum in twins. Twins had 14% fewer total follicles than singletons (P   0.05). Immediately post-partum there was a 36% decrease (P   0.05) occurred in the single lambs. This attrition coincided with a surge in plasma cortisol concentration that commenced ~12 days before birth and persisted for 6–12 days post-partum. The surge in plasma cortisol concentration occurred in both single and twin lambs, commencing on Day 138 of gestation; however, the peak cortisol concentration and the period over which cortisol remained elevated was greater in twin lambs than in singletons (P < 0.001). This study confirms a previous finding of a perinatal reduction in S/P ratio in Merino sheep. The reduction was greater in twin lambs than in singletons but the follicles of twins recovered rapidly so that there was little difference in final S/P ratio between birth types after Day 30 post-partum. The postnatal follicle reduction followed the perinatal surge in plasma cortisol concentration and appeared to reflect the magnitude of the cortisol surge in that twins, which displayed a higher peak cortisol concentration, had a greater reduction in active follicles than singletons.