F. D. Brien

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.

Neonatal lamb mortality: factors associated with the death of Australian lambs

The objective of the present study was to examine the factors associated with the death of neonatal lambs. Postmortem autopsy data were collected from 3198 newborn lambs in the Sheep CRCs Information Nucleus Flock situated in various environments throughout southern Australia. The proportion dying by category from highest to lowest was starvation–mismothering (25%), stillbirth (21%), birth injury (18%), dystocia (9%), death in utero–prematurity (10%), predation (7%), cold exposure (5%), undiagnosed (4%), infection (1%) or misadventure (1%). Factors best explaining the probability of lambs falling into a death category included both birth type and birthweight for dystocia, stillbirth, starvation–mismothering and death in utero–prematurity. The probability of a lamb falling into any category was predicted at the mean birthweight, within birth type. Single-born lambs were more likely to die from dystocia and stillbirth, while twin lambs were more likely to die from birth injury, starvation–mismothering or from undiagnosed causes. Triplet lambs were more likely to die from starvation–mismothering or death in utero–prematurity. Sire type (Merino, maternal or terminal) did not affect the proportions of lambs within any category. The proportions lost to each cause of death were largely consistent among locations, despite the rate of death varying. Dystocia, stillbirth and birth injury, as evidenced by the presence of oedema around the head and neck or by lesions of the central nervous system, accounted for 48% of autopsied lambs. We conclude that for improvements to occur in the rates of lamb survival, the Australian sheep industry must focus on minimising losses due to dystocia, stillbirth, birth injury and starvation.

Effect of periconceptional nutrition on the growth, behaviour and survival of the neonatal lamb

Periconceptional nutrition (PCN) can influence foetal hypothalamo-pituitary adrenal (HPA) axis function and alter cortisol secretion with possible consequences for maturation and growth of major organs, gestation length and behaviour. We examined effects of PCN on phenotype and survival of the neonatal lamb in 466 Merino ewes allocated to treatments providing 70%, 100% and 150% respectively, of maintenance requirements for 17 days prior and 6 days after insemination. Gestation length and birth weight for lambs in PCN treatment groups was similar (P > 0.05) but low PCN decreased the size of the neonate (crown-rump-length and metacarpal length P < 0.05). A subset of lambs euthanased at 5 days of age further showed that low PCN decreased the amount of peri-renal fat (P < 0.05) and increased liver mass (P < 0.05) while high PCN increased neck thymus and ovary mass (P < 0.05). Neonatal lambs from low PCN ewes returned faster to their mothers after release (P < 0.05) and contacted the udder in the shortest time (P < 0.05). Significant interactions between PCN treatment and sex (P < 0.05) and between PCN treatment and ewe age (P < 0.05) were also observed for time lambs took to follow the ewe. Survival of lambs was similar but potential differences may have been masked by favourable weather conditions. In conclusion, this study provides evidence of significant changes in lamb growth and development dependent on PCN and, for the first time, links these changes with significant changes in behaviour of the neonate. The impact of these effects on lamb survival and potential reproductive capacity of female offspring remains to be determined.

Thermogenesis and physiological maturity in neonatal lambs: a unifying concept in lamb survival

Lamb mortality represents reproductive wastage and an animal welfare concern. While lambs are thought to be at a thermogenic advantage following birth in comparison to other species, death from exposure can still be a major contributor to lamb mortality, largely because of the inclement conditions often prevailing at lambing. For this reason, thermogenesis has been studied extensively in neonatal lambs. Heat is produced in the neonatal lamb by shivering and non-shivering thermogenesis. The latter is heat generated by metabolism of brown adipose tissue (BAT) found largely in the thorax and peri-renal areas of the newborn lamb. Brown adipose tissue differs from normal adipose tissue in that it contains densely packed mitochondria, a high cytochrome c content and a vast vascular network. Heat is generated in BAT by uncoupling of the proton conductance mechanism from ATP production, resulting in heat production instead of stored energy. The ability of lambs to resist cooling differs among individuals and this is likely to be due to both genetic and phenotypic factors. The heritability of cold resistance is moderate-to-high and polymorphic gene markers associated with energy homeostasis and cold-related mortality have been identified. Additionally, several aspects of the phenotype of the lamb have been associated with cold resistance. Most relate to properties of the coat, skin and bodyweight, the latter being particularly important, presumably through effects on surface area to volume ratios and subsequent heat loss. The ability of the neonate to achieve the transition from intra- to extra-uterine life has been termed physiological maturity and is associated with the ability to activate appropriate neuro-endocrinological and behavioural changes that are consistent with homeostasis of energy metabolism. Ways to alter physiological maturity of the lamb, such as nutrition, pharmacology and genetic selection, have been identified, and while these show promising results with regards to thermoregulation, a key limitation of their application has been the lack of a repeatable, representative model of neonatal cold stress. An estimation of the non-shivering component potential of a lamb’s ability to thermoregulate can be derived from norepinephrine challenges, but more useful models of real-world cold stress are climate chambers or controlled water bath tests. Further use of repeatable test models such as these with appropriate neuroendocrine and metabolic metrics will identify key components and markers of physiological maturity associated with lamb thermogenesis and survival.

Effect of previous reproductive performance on current reproductive rate in South Australian Merino ewes

Reproductive performance of ewes in the current year may be influenced by their performance in the previous year and by the ability of seasonal and management conditions to facilitate full recovery before the next breeding cycle. This possibility was tested by analysing reproductive data from 2100 South Australian Merino ewes mated to Merino rams annually from 1991 to 1997. The model fitted to the data included the fixed effects of previous reproductive performance (PRP) (no lambs born; single or multiple lambs born, none reared; single lambs born, singles reared; multiples born, singles reared; multiples born, multiples reared), age of ewe (2.5, 3.5, 4.5 years), year (1992–1997), bloodline (A, B, L, X) and all two-way interactions. Current-year parameters were net reproduction rate (NRR, lambs weaned of ewes joined) and its components of fertility (ewes lambing of ewes joined), fecundity (lambs born of ewes lambing) and lamb survival (lambs weaned of lambs born). Other parameters were lamb liveweight at weaning per ewe joined and pre-mating liveweight. In addition, repeatability values were calculated across ages for these parameters. NRR was higher in the current year for those groups of ewes that reared lambs in the previous year; this was associated with the highly productive multiple-bearing and -rearing groups having a liveweight at the next mating similar compared to that of groups that did not rear lambs. Repeatability values of 0.27 for NRR and 0.40 for fertility suggest there are genetic and permanent environmental components contributing to the differences observed among the PRP groups for NRR, particularly so for fertility. Selecting ewes after their first lambing opportunity on fertility alone at a rate of 90% would give a non-cumulative increase in fertility of 3.4% at subsequent lambing opportunities. Hence, improvements in the current ewe flock for NRR by graziers managing flocks of the South Australian Merino strain are likely to come from (1) identifying and selecting those ewes of high fertility from early records and (2) attending to management factors impacting on fecundity and lamb survival.

The impact of selecting for increased ewe fat level on reproduction and its potential to reduce supplementary feeding in a commercial composite flock

In southern Australian grazing systems, energy availability typically exceeds energy requirements. Grazing systems are likely to have higher profitability if ewes can utilise this relatively cheap feed by gaining more condition (muscle and fat) and then mobilising it when feed is expensive. The present paper focusses on the importance of genetic merit for fat on the lifetime productivity of the ewe within a commercial operation. The analysis was carried out on a maternal composite stud flock in Holbrook, New South Wales. Ultrasound fat and muscle depth were measured on 2796 ewes as lambs at post-weaning and as adult ewes, to determine the genetic relationship between young and adult body composition. The hypothesis of the paper is that selection for increased fat at young age will improve body condition of adult ewes, which will lead to improved reproduction and potentially reduced requirements for supplementary feeding. Given that the difference in feed cost between times of abundance and shortage is likely to be larger, the system may become more profitable despite being less efficient (owing to increased feed intake). Our results indicated that selection for scanned post-weaning fat and muscle depth in lambs should lead to increased fat muscle and body condition in the breeding ewe because of strong genetic correlation estimates between the post-weaning and adult traits of 0.68–0.99. The influence of body composition traits on reproduction (number of lambs weaned) within the stud flock was weak. However, it is hypothesised that by genetically increasing post-weaning fat depth in ewes, producers could improve profitability by reducing the demands for supplementary feeding.

Variation in physiological profiles may explain breed differences in neonatal lamb thermoregulation

Ability to adapt rapidly from the uterine environment to self-thermoregulation following birth is a vital requirement for neonatal lamb survival. This investigation reports factors that could explain differences in thermoregulation among breeds that differ in lamb survival. Breeds such as the Merino and Border Leicester have previously been shown to be divergent for birthweight, cold resistance and lamb survival. Cross-bred (Poll Dorset Border Leicester (PDBL, n = 9) and Poll Dorset Merino (PDM, n = 25)) and pure-bred (Border Leicester (BL, n = 35) and Merino (M, n = 46)) lambs were recorded for the thermogenic measures rectal temperature at birth, cold resistance (time for rectal temperature to fall to 35°C while in a cooled water bath) and cold recovery (time to restore rectal temperature after cold exposure) at 1 day of age. In pure-bred lambs, 1 kg increase in weight resulted in a 0.25°C increase in rectal temperature at birth (P < 0.001) and 4.2 min increase in cold resistance (P < 0.001). In contrast, cross-bred lambs did not exhibit any relationship between birthweight and rectal temperature at birth, although they displayed a 3.2 min greater cold resistance for every 1 kg increase in birthweight (P < 0.001). BL-derived lambs were more cold resistant than M lambs (cross-bred: PDBL, 67.1 ± 2.5 min; PDM, 56.4 ± 1.6 min; P < 0.01; and pure-bred: BL, 58.1 ± 1.5 min; M, 53.2 ± 1.3 min; P < 0.01). The quadratic relationship of glucose concentration over time during cold exposure differed with lamb breed. PDBL exhibited higher peak glucose concentrations than did PDM (11.0 mmol/L and 8.9 mmol/L, respectively; P < 0.01). BL took longer to reach peak glucose concentration (50 min) than did M (40 min) and this peak value was higher (BL, 9.4 mmol/L; M, 7.7 mmol/L; P < 0.001). In conclusion, variations in birthweight and glucose metabolism are associated with breed differences in thermogenesis of neonatal lambs.

Ewe culling and retention strategies to increase reproductive rates in Merino sheep

The present empirical study of data from three Merino resource flocks of varying genetic backgrounds run in different environments investigated the impact of various selection strategies aimed at culling poor performers (‘passengers’) or retaining good doers (‘performers’) longer, or both, on the flock reproductive performance. Four strategies were investigated and applied either individually or in combination, to quantify their impact on the litter size, rearing type and breeding efficiency (number of lambs weaned per ewe per year) of the whole flock. The main benefit from implementing any of the strategies was a decrease in the proportion of dry ewes in each flock (1–7%). There was little difference between the single strategies of culling twice-dry ewes or culling ewes that twice lambed and lost their lambs. When retaining a cohort of older ewes, it was more efficient to select these on the basis of their lifetime reproductive performance. There was only a marginal benefit gained by combining strategies and, in some cases, a single strategy was still better in terms of breeding efficiency. Commercial producers can improve the reproductive rate of these flocks by culling twice-dry ewes on the basis of their pregnancy scan data.

Litter size at lambing influences genetic evaluation of maternal rearing ability

The genetic parameters for maternal rearing-ability of ewes were investigated by ignoring or defining the rearing ability trait separately by litter-size class (singles versus multiples) using multi-generational data from three, fully pedigreed Merino flocks differing in prolificacy, production level and environment. Genetic correlations (ra) between litter size (LSIZE) and the percentage of lambs surviving (PLSURV) were more negative with an increasing mean flock litter size (ra: –0.21 ± 0.17 to –0.73 ± 0.08), while the corresponding range in phenotypic correlations (rp) was substantially lower (rp: –0.15 ± 0.01 to –0.33 ± 0.01). Rearing-ability traits defined separately by litter-size class were highly correlated (ra: 0.49 ± 0.19 to 0.64 ± 0.38), but not genetically identical traits in the most prolific flock. Defining rearing-ability traits separately by litter-size class reduced the antagonistic genetic correlations between LSIZE and PLSURV to between –0.39 ± 0.14 and 0.14 ± 0.28, through accommodating the change in mean and variance of PLSURV with LSIZE. Similarly, linear transformation (TSURV) of PLSURV within each litter size to a common mean and variance reduced the antagonistic trend in genetic correlations between LSIZE and TSURV to range between –0.05 ± 0.17 and –0.43 ± 0.13. Since genetic correlations are low to moderate between TSURV and LSIZE, it is possible to select for improvements in both the number of lambs born and lamb survival simultaneously. Defining rearing-ability traits separately by litter-size class and the role of transformation will be investigated further for improving the accuracy of genetic evaluation for rearing ability across a range of flocks, breeds and environmental conditions using more extensive industry data.

Season and reproductive status rather than genetic factors influence change in ewe weight and fat over time. 4. Genetic relationships of ewe weight and fat score with fleece, reproduction and milk traits

The profitability of a sheep enterprise is greatly influenced by the ability of the ewe to produce a lamb annually. This paper examines the between- and within-breed association between the liveweight and fat score of the adult ewe and key fleece, reproduction and milk production traits. The study analysed the performance of 2846 first-cross Merino and Corriedale ewes from the maternal central progeny test (MCPT) over their first three production cycles. Breed (9 sire breeds) and within-breed (91 sires) correlations were calculated from bivariate correlations of adult liveweight and fat score with fleece, reproduction and milk production traits. The mean liveweight and fatness of the breeds were low to moderately negatively correlated with greasy fleece weight (–0.55 and –0.25, respectively) and moderate to strongly positively correlated with fibre diameter (0.88 and 0.41, respectively). However, the within-breed correlations were generally low to negligible. The genetic relationships between ewe fat score and reproductive traits at the breed and within-breed level was generally low to negligible and, in some cases, negative under production conditions where the ewes were well managed and averaged a fat score of 3.5 pre-joining and became too fat across the production cycle. Strong breed correlations between ewe fat score and milk fat concentration (0.88) suggest that selection of fatter sire breeds will lead to increased fat in their progenies’ milk, which was associated with larger lambs. The potential advantages of selecting for more fat to improve the maternal performance of the ewe seem to be limited in enterprises where the production system can easily be adjusted to restrict the influence of temporary and long-term feed deficiencies, as was achieved within the MCPT research flocks.