J.R. Briegel

Ewe lambs with higher breeding values for growth achieve higher reproductive performance when mated at age 8 months

We studied the relationships among growth, body composition and reproductive performance in ewe lambs with known phenotypic values for depth of eye muscle (EMD) and fat (FAT) and Australian Sheep Breeding Values for post-weaning live weight (PWT) and depth of eye muscle (PEMD) and fat (PFAT). To detect estrus, vasectomized rams were placed with 190 Merino ewe lambs when on average they were 157 days old. The vasectomized rams were replaced with entire rams when the ewe lambs were, on average, 226 days old. Lambs were weighed every week and blood was sampled on four occasions for assay of ghrelin, leptin and ß-hydroxybutyrate. Almost 90% of the lambs attained puberty during the experiment, at an average live weight of 41.4 kg and average age of 197 days. Ewe lambs with higher values for EMD (P < 0.001), FAT (P < 0.01), PWT (P < 0.001), PEMD (P < 0.05) and PFAT (P < 0.05) were more likely to achieve puberty by 251 days of age. Thirty-six percent of the lambs conceived and, at the estimated date of conception, the average live weight was 46.9 ± 0.6 kg and average age was 273 days. Fertility, fecundity and reproductive rate were positively related to PWT (P < 0.05) and thus live weight at the start of mating (P < 0.001). Reproductive performance was not correlated with blood concentrations of ghrelin, leptin or ß-hydroxybutyrate. Many ewe lambs attained puberty, as detected by vasectomized rams, but then failed to become pregnant after mating with entire rams. Nevertheless, we can conclude that in ewe lambs mated at 8 months of age, higher breeding values for growth, muscle and fat are positively correlated with reproductive performance, although the effects of breeding values and responses to live weight are highly variable.

Feed intake, body composition, and plasma metabolic hormones in Merino sheep that differ genetically in fleece weight or fibre diameter

Negative genetic correlations result in sheep selected for fleece weight having reduced fatness and reproductive turnoff. Both of these characteristics depend mainly on energy metabolism, but the links between wool growth and energy metabolism are poorly defined. The present study examined aspects of energy metabolism (body composition, feed intake, and metabolic hormones) in Merino sheep with high or low estimated breeding values for clean fleece weight (CFW) or fibre diameter (FD). The groups were selected to have similar mean liveweights. High CFW sheep had a lower proportion of fat (P < 0.01) and more lean tissue (P < 0.05) in their body, and ate more of a medium-quality diet (P < 0.05). Intake of a straw diet with low digestibility that imposed a physical constraint on feed intake was not associated with CFW. When fed at the same level (1.2 × calculated maintenance), high CFW sheep had lower plasma concentrations of glucose (P < 0.01), insulin (P < 0.05), and insulin-like growth factor-1 (P < 0.05), and higher concentrations of growth hormone (P < 0.001). Responses of plasma insulin or glucose to treatment with insulin, adrenaline, or propranolol were independent of CFW. No differences were found between the high and low FD groups, except that the plasma cortisol response to adrenocorticotrophic hormone (ACTH) was higher in sheep with low FD. Calculations indicate that the lower fat reserves and plasma insulin and glucose concentrations in the high CFW sheep are sufficient to account for previous reports of reduced number of lambs weaned in high CFW sheep. Furthermore, the results indicate that high CFW sheep have smaller metabolic energy reserves to withstand unfavourable feed conditions, and so may be less able to thrive and reproduce successfully in harsh feed environments.

Protein metabolism in skin and muscle of sheep selected for or against staple strength

Two experiments were carried out to determine the mechanisms underlying the reduced effect of nutritional status on wool growth rate in Merino sheep that have been selected for high staple strength (SS). In Expt 1, each group of 6 young sheep of SS+ and SS– genotypes were fed at 0.4 or 1.1 times maintenance, and in Expt 2, groups of 8 sheep of each genotype were fed at 1.1 and 1.8 times maintenance. In both experiments, rates of protein synthesis in skin, muscle, gut, rumen, and liver were determined using a flooding dose of labelled phenylalanine. Feed intake and the digestibility of feed were not affected by genotype. Neither dissection of the carcasses at slaughter, nor deuterated water analysis in Expt 1, detected any differences between the genotypes in body composition. The feeding level affected the total daily amount of protein synthesised in all the organs examined, and the fractional rate of protein synthesis was affected by feeding level in all organs except the liver. The fractional synthesis rate of protein was less responsive to feeding level in the SS+ sheep in both skin and muscle (P < 0.05), but not in the liver, jejunum, or rumen. Total protein synthesis in muscle, and the estimated rate of protein degradation, were also less responsive to feeding level in the SS+ sheep (P < 0.05). We conclude that sheep genetically selected for high or low SS have altered local mechanisms in both skin and muscle that control the way they respond to nutrition. These findings provide a mechanism by which selection for wool growth rate also affects body metabolism.

Differences in survival and neonatal metabolism in lambs from flocks selected for or against staple strength

In the field, we found that lamb mortality was lower (P < 0.01) in a flock selected for wool staple strength (SS+) than in a flock selected against staple strength (SS-), or a random-bred control flock (R); mortality rates calculated from 4200 records were 17.0, 22.8, and 24.1%, respectively. The proportion of twins was similar in all flocks, but birth weights differed (P < 0.01), being 4.40, 4.25, and 4.14 kg, respectively. Biochemical profiles were measured in lambs from 20 ewes from each flock around the time of birth in an animal-house study, to define potential causes of the genetic difference in survival. In the intensive study, birth weight was similar for all groups; however, gestation length was slightly shorter in the SS+ lambs (P < 0.05 for SS+ v. R). All lambs had low plasma concentrations of the essential amino acids and glucose at birth, compared with after suckling. Plasma concentrations of glucose, non-esterified fatty acids (NEFA), and blood urea nitrogen (BUN) at birth differed between lambs from the flocks with the greatest difference in mortality (i.e. SS+ v. R); that is, SS+ lambs had higher concentrations of glucose (1.84 v. 1.23 mM; P < 0.01) and NEFA (1.52 v. 1.08 mM; P < 0.05), and lower concentrations of BUN (7.7 v. 9.7 mM; P < 0.01) at birth, and for the next 9 h. The semimembranosus and semitendinosus muscle of the SS+ lambs also had lower concentrations of ATP (7.4 v. 5.0 μmol/g, respectively) compared with lambs from the other flocks, immediately post-partum. Muscle phosphocreatinine was also slightly lower in the SS+ lambs. No differences between the genotypes were observed in reserves of glycogen, carcass fat, brown adipose tissue, or in neonatal thyroid hormones. In summary, sheep selected for high wool staple strength had lower lamb mortality than the other 2 flocks. This was accompanied by metabolic differences at birth that can be associated with greater metabolic maturity, rather than greater energy reserves.

Responses of wool growth rate and body reserves to nutrition in Merino ewes: a potential biological link between production and fitness

This study examined whether the low body-fat reserves in sheep with high estimated breeding values (EBVs) for clean fleece weight (CFW) reported previously are affected by nutritional history, and second whether the effect may be related to differences in the variation in fibre diameter (CVfd). Groups of 11 20-month-old Merino ewes with high and low EBVs for CFW and for CVfd were compared in a 2 × 2 design at low bodyweight, then fed ad libitum for 100 days and re-measured. The response of wool growth rate to feed supply (WRF) was estimated as the slope of the regression against time of clean wool mass collected from mid-side patches at intervals of 30–40 days throughout the experiment. High CFW sheep had greater feed intake relative to liveweight and liveweight gain (P = 0.02), but did not differ significantly in other characteristics from low CFW sheep. High CVfd sheep had lower plasma concentrations of insulin (P = 0.02), IGF-1 (P = 0.03), and albumin (P = 0.02) throughout the study, and had less fat when in poor body condition (P = 0.02). The WRF was greater in both the high CFW (P = 0.003) and the high CVfd (P = 0.004) genotypes. When studied in poor body condition, sheep with a high WRF had lower liveweight (P < 0.001), lower body condition score (P < 0.001), lower plasma albumin (P < 0.001), and higher plasma growth hormone (P = 0.02), but these relationships weakened or disappeared after ad libitum feeding. Sheep with high WRF also had lower plasma concentrations of insulin (P = 0.002) and IGF-1 (P = 0.008) throughout the study, which may have brought about the increased responsiveness of protein and energy metabolism to nutrition. The results indicate that genetic selection for wool characteristics can affect the responsiveness of wool growth rate to nutrient supply. Sheep that are highly responsive grow more wool when offered abundant feed, but may have lower body nutrient reserves when on limited feed.

Relationships among body composition, circulating concentrations of leptin and follistatin, and the onset of puberty and fertility in young female sheep

The onset of puberty depends on the attainment of critical body mass, so should also be affected by increases in the rate of accumulation of muscle and adipose tissue. Adipose tissue and reproduction are linked by leptin. For muscle, a link has not yet been identified, although one possibility is follistatin. We assessed the relationships among circulating concentrations of follistatin and leptin and the rates of growth and accumulation of muscle and fat during pubertal development in female sheep. We used 326 animals with known phenotypic values for live weight (LW), depths of eye muscle (EMD) and fat (FAT), and known breeding values at post-weaning age for body mass (PWT) and depths of eye muscle (PEMD) and fat (PFAT). Leptin concentration was positively correlated with values for EMD, PEMD, FAT, PFAT, LW and PWT (P<0.001), whereas follistatin concentration was negatively correlated with values for EMD and PWT (P<0.001), and PEMD (P<0.01) and FAT (P<0.05). Leptin concentration was negatively related to age and positively related to live weight at first oestrus and the proportion of females that attained puberty (P≤0.05), and to fertility and reproductive rate (P<0.01). Follistatin concentration was negatively related to live weight at first oestrus and to fertility (P<0.01) and reproductive rate (P<0.05). There were positive correlations (P<0.001) between muscle accumulation and leptin concentration, and between muscle accumulation and reproductive performance. We conclude that leptin and follistatin are probably both involved in effects of accelerated accumulation of muscle and adipose tissues on the onset of puberty.

Plasma insulin concentrations and amino acid turnover in Merino sheep with high or low fleece weight

Although sheep with a relatively high fleece weight have reduced fat deposition and a lower reproductive rate, research has failed to identify any specific metabolic demand that growing a fleece places on the physiology of the sheep. This paper reports two experiments in which the effect of fleece weight was examined in ewes infused intravenously with amino acids. The first experiment was carried out when the ewes were 107 days pregnant, and the second 2 years later in non-pregnant ewes from the same flock. The ewes were derived initially from groups that differed in staple strength, but there was no effect of staple strength group on the characteristics measured in either experiment. In Expt 2, ewes were also infused with a bolus of l-[ring-d5] phenylalanine, and the enrichment in plasma determined by GC/mass spectrometry over the next 24 h. In both experiments, fasting plasma insulin concentrations were lower (P < 0.05) in ewes with a high fleece weight, and this difference continued during infusion in Expt 2 (P < 0.05). In Expt 1, infusion of ewes with amino acids resulted in higher (P < 0.05) plasma concentrations of α-amino nitrogen (indicating amino acids) in the ewes with a higher fleece weight, and in Expt 2, ewes with a high fleece weight had a 19% higher rate of appearance of endogenous phenylalanine (P < 0.05). We conclude that sheep with high wool growth rates have higher whole-body protein turnover rate, which may be achieved in part by lower insulin concentrations. Lower insulin in turn provides a mechanism through which wool growth rate may influence energy availability to other tissues.

Carcass and meat characteristics of sheep with an additional growth hormone gene

Effects of high growth hormone (GH) activity on body composition and some aspects of meat quality were examined in sheep transgenic for an additional copy of the ovine GH gene, as a tool to explore the biological importance of the GH axis in sheep selected commercially for meat production. Carcasses of 16 GH and 25 control mixed-sex sheep aged 45 months, and 6 GH and 6 control ewes aged 20 months, were measured. The dressing percentage was lower in the GH sheep (P < 0.001). The GH sheep had similar muscle mass to controls, but the weight of their fat depots was reduced (P < 0.001) to approximately 40% of controls, whereas limb-bone mass was 43% greater (P < 0.001) than controls. Fore and hind limbs were equally affected. Skin and most internal organs were heavier, particularly the pancreas, kidney, alimentary canal, and the liver. The concentration of intramuscular fat in the GH sheep was only 27% that of controls (P < 0.001), whereas the average pH of muscle 24 h after slaughter and the melting point of subcutaneous fat were both increased (P < 0.05). Similar changes in organ weights and body composition have been observed in sheep selectively bred to enhance lamb growth rate or to decrease fatness, suggesting that relative GH activity contributed to the outcomes of those experiments. This study indicates the importance of a multi-trait breeding objective to ensure that mechanisms associated with GH do not impair meat quality.

Effect of glucose or amino acid infusion during pregnancy in ewes with different wool growth responsiveness to nutrition

The reason why wool growth is suppressed in pregnancy is unclear. The present study examined the response to the nutrient drains imposed by pregnancy in 3 genotypes that had been selected for low (SS-) or high (SS+) staple strength, and a random control line. These flocks differ in the responsiveness of their protein metabolism to nutrient supply. Three groups of 20 pregnant ewes were fed to maintain conceptus-free liveweight, and were challenged with an infusion of glucose or amino acids in mid (Day 104) or late (Day 138) pregnancy. Infusion of glucose in late pregnancy caused a lower increment in plasma glucose concentration (P < 0.001) compared with mid-pregnancy, whereas plasma insulin responses to glucose infusion were not significantly different across time. In contrast, infusion of amino acids caused a greater increase in plasma concentrations of α-amino nitrogen in all genotypes in late pregnancy (P < 0.01), whereas the insulin response to amino acids was reduced (P < 0.001). Despite slower clearance of amino acids and decreased wool protein synthesis, the concentrations of blood urea nitrogen (BUN) were elevated in late pregnancy. The genotypes did not differ in their response to infusion with amino acids, but the SS- genotype had higher basal glucose and insulin concentrations. We conclude that the suppression in wool growth as pregnancy progresses was accompanied by greater oxidation but slower clearance of amino acids from plasma, associated with a lower insulin response. This defect in insulin response was selective for amino acids, because it was not observed in response to glucose infusion.